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VOL. 49 

Founded by William Henry Leggett 1870 




Jean Broadhurst 
James Arthur Harris 
George T. Hastings 
Marshall Avery Howe 

Michael Levine 
George Edward Nichols 
Arlow Burdette Stout 

Norman Taylor 





Published fob the Ciub 
The Intelligencer Printing Company 

Lancaster. Pennsylvania 








Pessin, Louis J. Epiphyllous plants of certain regions In Jamaica 

(plate i) I 

DuRAND, Elias J. The genus Catinella 15 

MuNZ, Philip A., and Johnston, Ivan M. Miscellaneous notes on 

plants of Southern California — I 31 

Farr, Clifford H. Quadripartition by furrowing in Sisyrinchium 

(plate 2) 51 

Round, Eda M. A modern plant fossil 63 

Hazen, Tracy E. The phylogeny of the genus Brachiomonas (plates 



Stevens, O. A. New records and other notes on North Dakota plants. 93 

YuNCKER, T. G. Three new species of Cuscuta from Mexico 107 

Hazen, Tracy E, New British and American species of Lobomonas: a 

study in morphogenesis of motile algae (plates 5, 6) 123 

Anderson, Flora. The development of the flower and embryogeny of 

Martynia louisiana (plates "^^ ^) 141 

Overholts,-L. O. Mycological notes for 1920 (plate 9) 163 

Graff, Paul W- Unreported plants from Glacier National Park 175 

OsTERHOUT, Geo. E, Two new plants from Colorado 183 

Arthur, Joseph Charles, New species of Uredineae — XIV 189 

Smith, Charles Piper. Studies in the genus Lupinus — -VII. L. suc- 

culentus and L. niveus 197 

Howe, Marshall A., and Hollick, Arthur. A new American fossil 

hepatic , 207 

Cockerell, T. D. a. a new genus of fossil Liliaceae 211 

Graff, Paul W. Philippine Basidiomycetes — V 

DosDALL, Louise. Occurrence of the pyncial stage of Puccinia Taraxaci 235 

Bartram, Edwin B. Midwinter botanizing in southern Arizona 237 

Britton, N. L., and Rose, J. N. T\vo new genera of Cactaceae 251 

RusBY, H. H. New species of trees of medical interest from Bolivia. . . 259 
Ashe, W. W, Notes on trees and shrubs of the southeastern United 

States 265 

Tharp, B. C, Commelinantia, a new genus of Commelinaceae (plates 

10, 11) , . 269 

Rowxee, W. W. The genus Costus in Central America (plates 12-15). . 283 

Shimek, B. Quercus lyrata in Iowa (plates 16, 17) 293 

PoRTERPiELD, W. M., Jr. References to the algae in the Chinese classics . 297 

Dodge, B. O. A Lachnea with a botryose conidial stage 301 

• • ■ 



Dates of Publication 

DoRAN, W. L, Effect of external and internal factors on the germination 

of fungous spores 3^3 

MuNZ» Philip A., and Johnston, Ivan M. Miscellaneous notes on 

plants of Southern California — II 349 

Mackenzie, Kenneth Kent. Notes on Carex — XII 361 

Oldenbush, Carrie. Stimulation of plants by carbon disulphide, . . 375 

Index to American botanical literature, 23, 45, 65, lii. 159, 185. 215, 

253^ 277, 307»34i» 391 

Index to volume 49 399 

Dates of Publications 

No. I, for January. 
No. 2, for February. 
No. 3, for March. 
No. 4, for April. 
No. 5, for May. 
No. 6, for June. 
No. 7, for July. 
No. 8, for August. 
No. 9, for September 
No. 10, for October. 
No. II, for November, 
No. 12, for December. 

Pages I- 30. 

31- 50. 

51- 74- 





Issued March 21, 1922. 

April 4, 1922. 

April 12, 1922. 

May 18, 1922. 

June 15, 1922. 

July 8, 1922. 

August 10, 1922. 

August 31, 1922. 

September 22, 1922. 

October 11, 1922. 

November 9, 1922. 

January 27, 1923. 


Page 2, line 8, for *'arboreum*' read "arbornm/' 
Page 6, footnote, for *'Carag7{ia'* read '' Caraguata. 
Page 14, line 7, for ' 


14, line 7, for ''Plate 5'' read "Plate i." 

Page 14, line 2 from bottom, for '' Caragtita'' read *' Caragtmta/'' 

Page 32, line 11 from bottom, for ''mutictis*' read "mutica,*' 

Page 38, line 13 from bottom, for *'racemosa** read **racemosum.'' 

Page 76, line i, for. ''gareement'* read ''agreement." 

Page 77, line 12 from bottom, for "genera" read '^general, " 

Page 92, line 4, for "ertracted" read "retracted." 

Page 100, line 4, for "Amsinkia" read "Amsinckia." 

Page 1 01, line 13, for "siberica" read "sibirica." 

Page 109, line 6, for "wartius" read "Martins." 

Page 109, line 10 from bottom, for "falsi" read "false," 

Page 125, line 7 from bottom, insert the word "are" after "sure," 

Page 127, line 16, insert a semicolon after "posteriorlbus. " 

Errata ( Continued) 

Page 127 

Page 143 
Page 156 
Page 166 
Page 167 
Page 180 
Page 189 
Page 195 
Page 197 
Page 208 
Page 224 
Page 226 
Page 228 
Page 241 
Page 242 
Page 242 
Page 244 
Page 245 
Page 247 
Page 248 
Page 271 
Page 271 
Page 275 
Page 291 

line 23, for "zsooporis" read "zoosporis/' 

Fig. 7, the lower right hand "S'' should be "S*." 








ne 12, for "1902" read "1901." 

nes I and 6 from bottom, for **hyaliua*' read **liyaliiins,** 

ne 14, for "hyalina" read "iiyalinus." 

nc 12, for "Pentstemon*' read "Penstemon.*' 

ne 9, for *'Pentslemon'* read *' Penstetnon." 

ne 5, for *' Bermudiauum*^ read ^' Bermiidiana, 
ne 8 from bottom, omit "with." 
after explanation of Fig, i add **X 3." 




Page 292, 1 
Page 320, 1 
Page 323. 1 

ne 16, for **corrugatus'* read *'corniga[a/* 
ne 9, for '' Polys tit ictus'' read '' Polysticfus." 

ne 18, for "mulitplex" read " 

ne 16, for "Notholeana" read "NoTHOLAENA. " 

ne I, for '*machrostachya" read "macrostachya/' 

ne 16, for "ringens" read "rigens. " 

ne 8 from bottom, for " Vauquelina" read "Vauquelinia. " 

nes 24 and 26, for "Parosella" read 





ne 7, for "cholorotica" read "chlorotica. 

ne 12 from bottom, for "Pentstemon" read ''Penstemon. 

ne 22, for "spathe" read "sheath of leaf." 

ne 6 from bottom, for "leaves" read "branches. 

nc 4, for **Tinant{a'' read " Tradescanlia." 

ne 15 from bottom, for "herbarium of Cornell University" read 

S. National Herbarium." 

ne 19, for "Aented" read "Oersted." 

nes 5 and 13 from bottom, for ** Enlothia" read '* Endothia. 

ne 7, for '^Flasmopora*' read *' Plasmopara." 



Vol. 49 

No. 1 




JANUARY, 1922 

Epiphyllous plants of certain regions in Jamaica* 

Louis J. Pessin 

(with plate I AND ONE TEXT FIGURE) 


The great variety and luxuriant growth of the epiphyllous 
plants seen in Jamaica in the summer of 1919 led the writer to 
study them from two points of view: first, their distribution in 
relation to climatic conditions; second, their histological relation 
to the leaves bearing them, in order to determine whether these 
epiphytes are to any degree parasitic. The study of the distri- 


bution and environment of the epiphyllous plants was made in 
the field. The relation of the'epiphyll to its supporting leaf was 




of the fact that they have long interested botanists. As early as 
1875 Berkeley (i) reported the occurrence of epiphyllous lichens 
growing on the leaves of the tea plant; some of these apparently 
epiphyllous forms showed marked parasitic adaptation. Such 
forms were found in abundance in warm moist regions. Two of 
the forms observed by Berkeley were regarded as species belonging 


to the genus Strigula. 

A few years after Berkeley's paper appeared, Cunningham (2) 
reported on an epiphyllous lichen and an epiphyllous alga, My- 
coidea parasitica^ parasitic on leaves of Mangifera indica^ Croton 
and Rhododendron. 

[The Bulletin for December (48 : 315-363) was issued February 28. 1922.I 
* Botanical Contribution No. 69 from the Johns Hopkins University. 


2 Pessin: Epiphyllous plants of Jamaica 

Marshall Ward (6) worked on an epiphyllous lichen which he 
believed to be the one worked on by Cunningham* He observed 
no penetration of the tissues of the supporting leaf by the lichen 
and therefore decided that it was not parasitic- He did find, 


however, one or more cross walls in the palisade cells of the sup- 
porting leaf beneath the epiphylls, which formed three or four 
layers of these cells instead of the normally single layer. 

Schmidle (5) found species of Trentepohlia arhoreum and T. 
aurea as well as Phycopeltis microcystis quite common as epi- 
phyllous forms in Samoa. Massart (3) found that some orchids 
pass their entire existence on a single leaf. He found also that 
the epiphyllous floras differ in different regions. Algae and 

liverworts are the most common epiphyllous forms, while mosses 
acquire the epiphyllous habit only in very humid ravines. 

Observations and methods 

Specimens of epiphyllous plants were collected in various 
regions in the Blue Mountains of Jamaica. The field observations 
were made during June and July, 1919, in the following places: 
Cinchona Plantation (5 ,000 feet) , Morce's Gap (4,934 feet) , 
New Haven Gap (5,600 feet), Green River Valley (3,000 feet), 
Markham Hill (5,800 feet), Vinegar Hill Road (4,500 feet), Clyde 
River Valley (4,500 feet) and Blue Mountain Peak (7,428 feet). 

Material was either pressed or preserved in glycerine in 4 per 
cent formaldehyde, or in 60 per cent alcohol. For careful histo- 
logical study, material was fixed in chromoacetic acid and pre- 
served in 70 per cent alcohol until finally dehydrated, embedded, 
cut and stained with the Flemming's triple stain. In some cases 
material was left unstained for comparison. 

The Cinchona Plantation- is situated on a spur running south 
from the Blue Mountains at five thousand feet above sea level. 
The temperature varies between the extremes of 48*^ and 82^ F. 
The vegetation of the garden about the residence of the Cinchona 
Plantation consists largely of introduced plants, many of which 
are Himalayan, Australian and Cape of Good Hope species. 
There are also native tree ferns, orchids, bromeliads, and many 
other transplanted native species. The banks along the trails 
outside the garden are covered mostly by native ferns interspersed 

Pessin: Epiphyllous plants of Jamaica 3 

with, or on the north side of Cinchona Hill largely replaced by, 
Hedychittm coronaritim. 

During the wet season Cinchona is enveloped in clouds for days 
at a stretch and the rains are very heavy and frequent. In spite 
of this abundance of moisture at certain times of the year, epi- 
phyllous plants are rather scarce on the south side of Cinchona 
Hill. Only one hepatic, two species of algae and a few kinds of 
lichens were found growing there. Of these Trentepohlia grows 
in damp or shaded places on the ground, on the bark of trees and 
on the leaves of certain plants. Lichens, the most common of 
which belong to the genus Sticta, grow abundantly on certain 
plants. Some of the leaves of Callistemon lanceolatunty for instance, 
are almost completely covered by this lichen. In a few cases the 
leaves of Hedychium were found sprinkled with a reddish disk- 
like alga belonging to the genus Phycopeltis. The epiphyllous 
flora of this region is evidently not a rich one (see Table I, A). 

At Morce's Gap, tw^o miles north of Cinchona and at about 
the same level, is found a characteristic tropical montane rain 
forest. Here moisture is abundant at all seasons of the year. On 
the windward side of the mountains everything reeks with mois- 
ture; clouds drift constantly through these dense forests and rain 
falls frequently during eight months of the year. The dense 
vegetation, except on the ridges, is usually dripping wet. Very 
rarely do the sun's rays penetrate to the floor of these forests. 
Ferns carpet the floor and cover the walls of the steep-sided 
ravines. Tree ferns, such as Alsophila and Cyathea, are very 
common, many of them having trunks four or five inches thick and 
thirty feet high or more. Species of the genera Marattia, Danaea, 
Polypodium and Elaphoglossum are very abundant. Such forms 
as Blechnum attenuatum and many species of Trichomanes, Hymeno- 
phyllum, Polypodium and Elaphoglossum have been compelled to 
adopt either the climbing or the epiphytic habit in the intense 
competition for sunlight, Podocarpus, Hedyosmum, species of 
Piper and many other dicotyledonous trees and shrubs, together 
with numerous herbaceous plants, are scattered through these 
woods, increasing the struggle for existence and making the 

competition for light more keen. 

In such an environment epiphyllous plants are very abundant. 


Epipijyllous plants of Jamaica 


Supporting plant 





A. Cinchona Plan- 
tation, 5000 ft. alt.; 
■ South exposure. 


Callistemon lanceolatum 


Parmelia sp. 
Sticia sp. 
Usnea sp. 

A form of Pilocar- 

Cololejeunea diaphana 



Hedychium coronariurn 

Phycopeltis sp. 
Trentepohlia aurea 

Lopadium sp. 




Pittosporum iohira 

Phycopeltis sp. 



B. Morce's Gap, 
4934 ft. alt.; north 

Elaphoglossunt latifo- 

Desmidium sp. 
Nostoc sp. 
Phycopeltis sp. 
Pinnularia sp. 
Scytonema sp. 
Staurastrum sp. 


Cololejeunea sp. 
Cololejeunea diaphana 
Crossotolej eunea sp. 


Metzgeria furcata 
Microlejeunea sp. (?) 


Hedychium coronarium 


Phycopeliis sp. 

Crossotolej eunea sp. 
Aphanolejeunea sp. 
Metzgeria furcata 
Cololejeunea diaphana 
pellucida Schiffn. 



Hedyosmum arhor- 
escens Sw, 


Crossotolej eunea sp. 
Lejeunea Jlava 

■* * 









Supporting plant 





B. Morce's Gap — 

Mangifera indica 

Phycopeltis sp. 

L^n determined si>ecies* 

Podocarpus coriaceus 

Phycopeltis sp. 


Crossotolejeunea sp. 
Lejeunea Jlava 

. Polypodium sp. 

Undetermined species* 



Clastohryum ameri- 


C. New Haven Gap, 
5600 ft. alt.; north 

Stelis ophioglossoides 

Phycopeltis sp. 
Trentepohlia aurea 


Trichomanes sp. 



Crossotolejeunea sp. 


Blechnum attenuatum 

Phycopeltis sp. 
Pinnularia sp. 
Nosioc sp. 
Scytonema sp. 



Cololejeunea diaphana 
Crossotolejeunea sp. 



Lophocolea sp. 
Metzgeria furcata \ 
Taxilejeunea sp. 

re plans 


Clusia haielioides 


Pinnularia sp. 

Nostoc sp. 
Scytonema sp. 



Parmelia sp, 


Aphanolejeunea sp. 




Cololejeunea diaphana 
Lejeu7iea Jlava 
Metzgeria furcata 
Microlejeunea sp. 

M iltenothamnium 
re plans 

* Up to the present the writer has been unable to have some of the epiphyllous lichens definitely determined. 


New Haven Gap 


D. Vinegar Hill 
Road, 4500 ft. alt.: 
north and \vest 

Supporting plant 

Monoclea CoUschei 

Plagiockila sp. 

Rhododendron sp 

Asplenium sp 

Clusia havelioides 

lati folium^ 

TABLE I— Continued 


Nostoc sp. 
Phycopeltis sp, 
Pinnularia sp. 

Chroococcus sp 
Gleocapsa sp. 
Nostoc sp. 
Phycopeltis sp. 
Scytonema sp. 


Phylloporina epiphylla 
Undetermined species 




Riccardia sp 



Cololejeunea sp. 
Cololejeunea diaphana 
Crossotolejeunea sp. 


Melzgeria furcata 

Cololejeunea sp. 


Lejeunea flava 
Lophocolea sp- 
Metzgeria furcata 
Taxilejeunea sp. 

Cololejeunea diaphana 
Crossotolejeunea sp. 
Lejeunea flava 
Melzgeria furcata 


* On a leaf of this species the seed plant Caraguta SinienesH was likewise found living as an epiphyll. 


D. Vinegar Hill 
Road — Continued. 

E- Blue Mountain 
Peak. 7428 ft. alt.; 
northeast expo- 

F. Clyde River Val- 
ley, 4000 ft. alt.; 
north and west 

TABLE I — Continued. 


Supporting plant 
Epidendrtim sp. 


Polypodium Phyllztidis 

Podocarpus coriaceus 

Phycopeltis sp. 

Citrus Limonium 

Phycopeltis sp. 
Scytonema sp. 
Trentepohlia aurea 


Undetermined species. 

Coenogonium sp. 
Collema sp. 

Lecanora sp. 
Pannaria rubiginosa 

Pyrenula sp. 

Undetermined species. 




Cololejeunea sp. 
Cololejeunea diaphana 
Crossotolejeunea sp. 




Lophocolea sp. 
Melzgeria furcata 
Taxilejeunea sp. 

Fissidens sp 

Aphanolejeunea sp 
Cololejeunea sp. 


Lejeunea flava 
Melzgeria furcata 

8 Pessin: Epiphyllous plants of Jamaica 

Here one finds plants of many types growing on the soil; on these 
grow various plants as epiphytes and on these in turn grow other 
epiphytes. A remarkable diversity of epiphytic forms can be 
found on one plant, and some of the forms are wonderfully adapted 
to their epiphytic mode of life. These epiphytes occur on every 
patch of dead or living plant surface that receives even a moderate 
supply of light. One finds on a single leaf here such forms as 
algae, lichens, hepatics, mosses and, at times, even seed plants, 
not to mention occasional fungi and minute epiphyllous animals. 
It is rather strange that not a single fern prothallus has been 
found on any of the leaves of this region, though numerous spo- 
rangia were present and probably spores also on many different 
kinds of leaves. In Table I, B, are listed the different kinds of 
epiphyllous forms found near Morce's Gap. ^ 

The climatic conditions of New Haven Gap do not differ 
markedly from those of Morce*s Gap. The temperature is nearly 
the same, ranging annually from 40.5*^ to 83"^ F., and the floor of 
the forest is constantly moist and shaded. Here again one finds 
the characteristic rain forest vegetation. The epiphyllous plants 
here are quite as abundant as at Moreens Gap and consist mainly 
of algae, hepatics, and mosses. The leaves of the climbing plants 
of this locality are well inhabited by epiphyllous forms. The 
leaves of Clusia havetioides and of such ferns as Blechmim atten- 
uatum, Elaphoglossum latijolium and Poly podium Phyllitidis serve 
most frequently as the substratum for epiphylls (see Plate i). 
A glance at Table T, C, gives one a fair notion of the groups of 
epiphyllous plants found in the New Haven Gap region. 



it presents a splendid view of the ravines. The crowns of the tree 
ferns carpet many side ravines of the valley. The huge lianes, 
the numerous orchids, the wild pines and the purplish filmy ferns 

m and Polypodium 
plaschnichianum) , i 


spread over the banks. One does not 
have to walk off the trail to find numerous epiphyllous plants. 
The upper leaves of many of the low trees on the Vinegar Hill 
Road are completely covered by them. Table I, D, shows the 
great variety of epiphyllous forms found in this region. 

Pessin: Epiphyllous plants of Jamaica 9 

On Table I, E and F, are indicated the epiphyllous plants 
observed on the Blue Mountain Peak and in the Clyde River 
Valley. In the former, though there is always an abundance of 
moisture, the epiphyllous flora is not a rich one, owing probably 
to the frequent heavy rains, the rather low temperature and the 
occasional strong winds. Those of the Clyde River Valley occur 
chiefly on trees growing along streams, where there is sufficient 
shade and a humid atmosphere. 

Fig. I. Part of transverse section of a leaf of Elaphoglossum laiifolium (E) 
with the rhizoids (Rh) of an epiphyllous liverwort (L^) pressed into its cuticle. This 
liverwort in turn bears another epiphyllous liverwort (L^), which is likewise attached 
by its rhizoids (Rh). 

In general, observations in these Blue Mountains of Jamaica 
show that the epiphyllous members of each group of plants are 
limited to certain regions where climatic conditions are suitable 
for such growth. Thus, Algae, Hepaticae, and Musci usually 
thrive best on the windward side of the mountains and are abun- 
dant only in very damp and very shady woods. Epiphyllous 
lichens, on the other hand, do not require very warm, humid, and 
shady places but are also found In relatively dry regions. It was 
rather difficult to ascertain with any degree of definiteness the 
exact age of the leaves which bore epiphyllous plants, but it is 
safe to state that none are present on very young leaves. As a 
rule they do not appear on leaves until the latter are one or two 


seasons old. 

In order to determine the relation between the epiphyllous 
forms and the plants on which they grow, cross sections were 
made of the leaves of Elaphoglossum laiifolium, Clusia havetioides, 

10 Pessin: Epiphyllous plants of Jaiviaica 

Polypodium Phyllitidis and Mangifera indica. The leaves of 

Elaphoglossum latifolium were those chiefly used because of the 

abundance on them of epiphyllous liverworts in various stages of 

development. Sections of leaves of Mangifera indica were also 

studied to determine the relation of an epiphyllous lichen to the 

mango leaf. 

Sections of the leaves bearing epiphyllous hepatics showed 

characteristic thickened, club-like rhizoids arising in groups from 

the stem of the creeping liverwort. These rhizoids are sunken 
into the cuticle of the supporting leaf as if pressed in by some 

external force. The rhizoids broaden at their lower ends and 
flatten out as is shown in Text-fig. i. This is true even of the 
rhizoids of one epiphyllous liverwort when growing upon another, 
for one sometimes finds two or more tiers of epiphylls growing 
upon a fern or a dicotyledon. The rhizoids of the upper hepatic 
are in such cases decidedly flattened against the delicate leaves 
of the lower liverworts. This too is shown in the figure. The 
important point to be noted is that the rhizoids of the epiphyllous 
form were not observed to penetrate the tissues of the supporting 
leaf. Nor was there evident any abnormality in the internal 
structure of the supporting leaf, such as might conceivably be 
produced by the shade or by the weight of the epiphylls on the 
leaf or, possibly, by some substance such as an organic acid 
excreted by the rhizoids on the liverwort. 

The leaves of Mangifera indica bore numbers of orange-red 
disk-like plants of a species of Phycopeltis. Among these orange- 
red specks there were also minute grey patch-like lichens resem- 
bling those described by Cunningham (3) and Ward (9). The 
relation between these lichens and the mango leaf on which they 
grew has not been definitely determined. The sections of the 
mango leaf thus far studied showed no modifications of its tissues 
and no penetration by the lichens. 


The problem of the evolutionary origin and individual develop- 
ment of the epiphyllous plants is a complex one which still requires 
much careful investigation. Schimper (4) observed that epiphytic 
seed plants commonly originate on their support through seeds. 

Pessin: Epiphyllous plants of Jamaica 11 

The scattered seeds germinate on the trunk, branches, or leaves 
of a tree or shrub and, with adequate moisture and other nutritive 
material, develop into epiphytes. He divides the fruits and seeds 


of epiphytes into three categories: first, those transported by 
birds, monkeys and other tree-inhabiting animals; second, those 
which are very light and may be transported by the wind and 
which may readily catch in the crevices of the bark and germinate 
there; third, those seeds which are especially adapted to be carried 
by the wind. Such seeds usually possess either hairs or wings or 
inflated coats which aid their distribution. 

It is conceivable that a great many of the epiphyllous forms 
here being considered arise from spores, for spores have been found 
on many leaves. The epiphyllous liverworts evidently arise 
often from gemmae, for many such gemmae in the process of 
development have been found on leaves. The spores of liverworts, 
on the contrary, have rarely been found germinating on leaves. 
The observations thus far made indicate that many an epiphyllous 
hepatic or moss makes its start on the host by creeping up the 
stem of the support from the ground or up the petiole from the 
stem. As it climbs upward it dies off from below and when its 
younger part reaches the leaves it spreads over the surface and 
becomes strictly an epiphyllous form. Secondly, epiphyllous 
forms may be lodged upon the host by the agency of water. 
When during a rain the water flows down over the leaves and 
branches of a tree in the rain forest, it breaks off the hepatics and 
mosses and other plants from the surface of the stem or leaf and 
washes them down on the leaves below, where they continue to 
grow, covering the leaf in the course of time with a dense mat. 
Such is probably the explanation of the fact that in most cases 
the lower leaves possess the greatest number of epiphyllous plants, 
though, of course, lower leaves are older, better shaded, and are 
surrounded by moist air. It is probable also that the epiphyllous 
plants found on the dorsal sides of certain overturned leaves arise 
in the manner just described- The fact that no epiphyllous 
plants were observed on the lower sides of leaves still in normal 
position Is evidence that these plants are propagated largely by 
these fragments. If it were commonly by spores we should 
expect these, at least occasionally, to stick to the underside of a 

12 Pessix: Epiphyllous plants of Jamaica 

leaf. Such cases, however, are extremely rare. In a third possible 
method of dispersal, spores, gemmae, or seeds may be washed down 
by the dripping water from above, or may be blown from adjacent 
plants by the wind. This method is particularly common among 
algae and lichens. The first two types of vegetative propagation 
seem to be the commoner ones, for in many instances the epiphyl- 
lous forms growing on the stem have actually been observed to 
spread over the surface of the leaf. Moreover, many of the forms 
found on the leaves occur likewise on the stem of the supporting 
plant. This is particularly true of hepatics and mosses. In 
lichens, however, the case may be different. There the alga, as 
well as the component fungus, may often be present separately on 
the leaf; each may reach the leaf in the form of loose spores or 
fragments, and there combine to give rise to the lichen itself. 

Massart (3) failed to find in Java any epiphyllous pteridophytes 
and phanerogams. In the tabulated lists of epiphyllous plants of 


Jamaica in this paper it will be noted that no pteridophytes have 
been found. Although many fern sporangia were observed among 
the epiphylls by the writer, no germinating fern spores or fern 
prothallia were seen. The writer, however, found a few epiphyl- 
lous phanerogams. These were very young plants of the genus 
Caraguata attached to the surface of the leaf, and apparently doing 
well. How much further their development w^ould continue is 
hard to tell, for the substratum is not firm enough to support a 
mature wild pine; before the epiphyllous wild pine could have time 
to mature, the supporting leaf would probably be shed. The fact 
is, at all events, that no mature wild pines were found growing on 
leaves. According to Massart, the epiphyllous plants do not 
constitute a similar flora everywhere; in some localities this consists 
solely of thallophytes, in others of bryophytes. This is also true 
for Jamaica, where in certain regions only one type occurs, while 
in other regions numerous diverse forms are present. 

Massart's suggestion that the leaf-inhabiting fungus has been 


derived from the epiphyllous plant is at least favored by the rela- 
tion to its host of the alga Cephaleuros, which may have either the 
parasitic or the epiphytic mode of life. On the other hand, there 
are still some fungi which possess the epiphyllous habit, such as 
Metiola astrina and a few others. There seems little doubt that 

Pessin: Epiphyllous plants of Jamaica 13 

a careful study of epiphyllous plants would throw some light on 
the origin of parasitism, such as that of leaf-infesting fungi. 


■a ■ 

!• Epiphyllous plants of diverse selected regions in Jamaica 
were 'studied in the field and collected for laboratory study in 

2. The results of the field observations were tabulated so as 
to give at a glance a general idea of the distribution of these 
epiphyllous plants in relation to the altitude^ average temperature, 
and average humidity of their various habitats. 

3. The possible modes of attaining the epiphyllous position 


(a) Continual growing forward and dying ofif below, of hepatics 

and mosses, until they reach and cover the leaf. 

(b) By distribution of spores, gemmae or seeds through the 
agency of rain and wind. This is especially true for algae and 

(c) Propagation by displaced fragments of mature lichens, 
hepatics, and mosses. 

4. An abundance of humidity, moderate temperature, and a 
relatively calm atmosphere encourage an abundant epiphyllous 

5. Histological studies of epiphyll-bearing leaves of Elapho- 
glossnm latifolium, Clusia havetioides and Mangifera indica revealed 
no apparent penetration of the leaf by any part or organ of the 

6. The rhizoids of epiphyllous hepatics are especially adapted 
for the epiphyllous habit by becoming thickened, by flattening 
out and by being pressed into the cuticle of the leaf, thus firmly 
attaching the epiphyll to the leaf and preventing its being blown 
or washed off easily. 

7- It is conceivable that the shade produced by the epiphyll, 
the accumulation of the organic material, the possible excretion 
of acids and other substances by its rhizoids may, have some 
physiological effect on the supporting leaf. 

8. A careful study of the epiphyllous mode of life will, it is 
believed, bring out some close relations between parasitism and 

14 Pessin: Epiphyllous plants of Jamaica 

the epiphyllous habit. Parasitism is possibly an advanced, 
specialized phase of epiphytism. 

The writer is indebted to the Botanical Department of the 
Johns Hopkins University for the opportunity of going to Jamaica. 
The writer wishes to thank particularly Professor Duncan S. 
Johnson for suggesting this problem, for taking the photographs 


reproduced on Plate 5, and for helpful advice in carrying on the 
work. Acknowledgments are due Professor Alexander W. Evans 
for the determination of most of the epiphyllous hepatics; to 
Mrs. N. L. Britton of the New York Botanical Garden for the 
determination of some of the epiphyllous mosses; and to Pro- 
fessors C. C. Plitt and Lincoln W. Riddle for the determination 
of certain epiphyllous lichens. 

Literature cited 

1. Berkeley, M, J. On the thread blight of tea. Quart. Jour. Microsc. 

Sci. 15: 130-133- /• i-4' 1875. 

2. Cunningham, D. D. On Mycoidea parasitica. Trans. Linn. Soc. 

London IL i: 301-316, pL 42, 43. iSjj. 

3. Massart, J. Les veg^taux epiphylles. Ann. Jard. Bet. Buitenzorg. 

Suppl. 2: 103-108. 1898. 

4. Schimper, A, F. W, Botanische Mittheilungen aus den Tropen 2. 

Die epiphytische Vegetation Americas. Jena. 1888. 

5. Schmidle, W. Einige Baumalgen aus Samoa. Hedwigia 36: 277- 

287. pL 6~g. 1897. 

6. Ward, H. M. On the structure, development and life history of a 

tropical epiphyllous lichen. Trans. Linn. Soc. London IL 2: 87- 
119. pi. 18-21. 1883. 

E^xiplanatlon of plate X 

Fig. I. Leaf of PoJ>/?oJimw PAyh'/2jj5, showing epiphyllous hepatics and mosses. 

Fig. 2. Leaf of Elaphoglossuyn latifolium, showing a luxuriant growth of epiphyl- 
lous plants; a small wild pine, Caragula Sintenesii, can be seen on the right hand 

The genus Catinella 


The genus Catinella was established by Boudier* for a single 

species, of rather indefinite relationship, which has commonly 

been identified as Peziza oUvacea Batsch. While Batsch's plant 

* is indeterminable when tested according to present day standards 


of specific limitations, the fungus which has gone by that name is 
well known, being clearly described by Massee,t and nicely figured 
by Boudier.J The synonymy listed below emphasizes not only 
the truth of Boudier's statement that the species has been in- 
cluded sometimes in one genus, sometimes in another, but that 
it has been referred by various writers to several different families 
as well. The inoperculate asci and intense reaction to caustic 
potash remove it from the Pezizaceae, where the bright color and 
fleshy consistency of the fresh young plants at first seem to place 
it, and to which It has been referred by Saccardo, Rehm and others. 


The somewhat gelatinous nature of the excipulum has suggested 
the Bulgariaceae to Fries, Karsten and others, but that character 


is certainly not conspicuous enough to associate the plant with 
Bulgaria ingttinans or Sarcosoma ruftim. Moreover, a truly 
gelatinous tissue does not become friable on drying. The present 
writer is inclined to the opinion of Berkeley, Phillips, Massee and 
Boudier that its affinities are more properly with the Patellari- 
aceae, such, for example, as Karschia. The genus may be charac- 
terized as follows: 

A genus of the Patellariaceae. Ascomata fleshy and somewhat 
gelatinous when fresh, becoming friable when dry, sessile, attached 
to the substratum by radiating, dark fibers; excipulum entirely 
parenchymatous, becoming deep violet with KOH; asci opening 
by a pore, not blue with iodine; spores eight, continuous, pale 
brown; asci and paraphyses agglutinated at the tips to form an epi- 

* Hist. Class. Disc. Eur. 150. 1907, 
t Brit. Fungus-Fl. 4: 94. 1895. 
t Icon. Myc. 3. pi. 452. 


16 Durand: The genus Catinella 

Species two, as follows 

Spores 7-1 1 X 4-5 M- 
Spores 13--15 X 6-7 ju. 

1. C nigrO'Olivacea 

2. C. elastica. 


. Catinella nigro-olivacea (L. v. S.) Durand, comb, nov. 


pi. 12 J. 51, 1783. 

Peziza nigro-olivacea L, v. S, Syn, Car. 121. 1822. 

Patellaria pulla ^ nigro-olivacea Fr. Syst. 2: 160. 

Bulgaria nigrita Fr. Elench. 2: 16. 1830. 

Lemalis rufo-olivacea L. v, S. Syn. N, Am. n. 1089. 1834. 

Rhizina nigro-olivacea Curr. Trans. Linn. Soc. 24: 494. pi. 51. 

/. 10-12 {fid. 






(fide Massee 

14: 108. 

pplanata B. & Br. J 


Peziza fuscocar pa Ell. & Hoi. Jour. Myc. i: 5. 1885. 
Patellaria olivacea Phill. Brit. Disc. 361. 1887 (Batsch ?). 


1889 (Batsch ?). 


Pezicula viridi-atra Sacc. Syll. 8: 315. 1889. 

Phaeopezia Jiiscocarpa Sacc. Syll. 8: 474. 1889. 

Bulgariella pulla jS nigro-olivacea Sacc. Syll. 8: 638. 

Bulgariella nigrita Sacc. Syll. 8: 638. 1889. 

Patinella violacea Sacc. Syll. 8: 770. 1889 {fide Massee). 

Patinella olivacea Sacc. Syll. 8: 770. 1889 (Batsch ?). 

Patinella hirneola Sacc. Syll. 8: 771. 1889 {fide Massee). 

Patinella applanata Sacc. Syll. 8: 771. 1889. 

Ilumaria marchica Rehm, Rabenh. Kypt.-Fl. i^: 952. 1894. 



Aleurina niarchica Sacc. & Syd. Syll. 16: 739. 1902. 

fuscocarpa Morgan, Jour. Myc 
fuscocarpa Sacc. & Syd. SvII. 11 


Catinella olivacea Boud. Hist. Class. Disc. Eur. 150. 1907 
(Batsch ?). 

Ascomata solitary or two or three together, sessile, attached to 
the substratum by numerous, radiating, dark brown fibers most 

Durand: The genus CatInella 17 

conspicuous ifi young plants; at first spherically closed, then open- 
ing out so as to become successively cup-shaped, saucer-shaped, 
finally applanate, with a permanently upturned margin; very 
young plants greenish yellow, becoming deeper and darker green, 
finally blackish olive when old, fleshy and somewhat gelatinous 
when fresh, 3-10 mm. in diameter, the majority 3-5 mm. On 
drying the disk becomes olive-black, the exterior brownish, some- 
what furfuraceous or vertically striate, and the texture friable. 
Exciptdum about 400 // thick at the base, entirely parenchymatous, 
the ectal cells 18-20 ju in diameter, polygonal, with dark brown 
walls, becoming gradually smaller and yellow toward the hy- 
menium, the superficial cells of the margin and sides projecting to 
form short, stout, obtuse, septate, flexuous, brown pili causing the 
surface to appear furfuraceous, those at the base much longer and 
radiating so as to form a small subiculum covering the substratum 
2-3 mm. beyond the margin. On treatment with i per cent 
caustic potash the hymenium or whole flesh becomes deep violet 
and frequently yields a violet solution. Hymenium about I20~ 
140 /i thick. Asci narrowly cylindric-clavate, apex rounded, not 
blue with iodine, 65-90x3-6/1 (mostly 70-80 /x). Spores eight, 
uniseriate, elliptic or elliptic-clavate, usually slightly narrowed 
near the middle so as to appear slipper-shaped, continuous, two- 
guttulate, pale olive-brown to deep brown, 7-1 1 x 4-5 m (majority 
8-10 At) ; Paraphyses cylindric, septate, rarely branched, somewhat 
longer than the asci, the tips agglutinated with amorphous matter 
to form an epithecium. 

On rotten wood, especially of old, moist, carious, decorticated 
logs, not abundant but widely distributed in eastern North America 
and the West Indies; also in Europe and Ceylon. 

This is one of our most peculiar and attractive discomycetes. 
"In its young state it is truly PezizaA\k^^ and very beautiful." 
While easily recognizable at sight when once understood, it has 
been described as new over and over again until very few fungi 
have a greater multiplicity of names. While this may be the 
species to which Batsch applied the name Peziza olivacea, his brief 
description and crude figures are much too inadequate to justify 
more than surmise, and may apply equally well to numerous other 
species. The apparent rarity of our plant in central Europe makes 
it still more improbable that Batsch had seen it. P. olivacea is 
simply one of the numerous names of fungi handed down from the 
pre-Persoonian period, the mycological stone age, w^hich are ab- 
solutely indeterminable according to present taxonomic standards, 

18 Durand: The genus Catinella 

and which should therefore be dropped completely and no longer 
allowed to encumber the mycological literature. 

The first certainly identifiable name applied to the plant*under 
consideration is the one adopted in the present paper* While 
Schweinitz's type, from North Carolina, is missing from his her- 
barium, there is a specimen marked ''rotten wood, Beth,*' under 
the n2ime'' Lemalis pulla fi nigro-olivascens Schw. No. 1089." This 
label involves a curious switching of names and an error in spelling 
as follows: Peziza nigro-olivacea L. v. S. was included by Fries in 
his Systema as ^^Patellaria pulla ^ nigro-olivacea,'' In 1834, 
Schweinitz transferred the species to the genus Lemalis^ and 
changed the specific name so that it appeared as ''ioSq. 4. L. 
rtifo-oUvacea, L. v. S., Syn. Car. 1220, Peziza nigro-olivacea; Salem 
'et Bethlehem differt specifice a priori [L. ptilla Fr,]", At a later 
date, Dr. Michener, in mounting and arranging Schweinitz's fungi, 
restored the original specific name (nigro-olivacea) to the label, 
but misspelled it *^ nigro-olivascens.^^ The presence of numbers and 
references In each case leaves no doubt that the specimen now in 
the Schweinltzian Herbarium is the one referred to in 1834 as from 

Although Fries stated that he had seen a specimen of Peziza 
nigro-olivacea, there seems to be no Schweinltzian material so 
labeled in the Friesian Herbarium, at Upsala. However, there is a 
specimen called ''Lemalis rufo-olivacea Schw.," from Curtis, as well 
as one marked ''Peziza applanata ex herb. Schweinitz.*' While 
both are typical Catinella nigro-olivacea, the latter is quite dif- 
ferent from the specimen in Schweinitz*s own herbarium called 
P. applanata. 

In 1830, Fries described as Bulgaria nigrita a plant collected 
in Russia by Weinmann. So far as the writer can discover, no 
more recent collection has been referred to this species, which has 
remained practically unknown for nearly a century. In the 
Friesian Herbarium is a specimen marked ''Bulgaria nigrita, 
Petersburg," in Ellas Fries's own hand, which is in all f)robability 
the original type. When the writer saw it, in 1904, he was at once 
impressed with its gross resemblance to the American plant. Sub- 
sequent microscopical examination removed all doubts as to their 
specific identity. 

Durand: The gexus Catinella 19 

During the Berkeleyan period this species was collected occa- 
sionally in various parts of the worid, and almost as often described 
as new. Material from Cuba (Wright, No. 369) was given the 

name Peziza viridi-atra B. & C. Examination of the type, at 

Kew, shows it to be identical with the Schweinitzian plant, 

Saccardo placed it in Pezicula. Specimens from Ceylon were 

called Patellaria applanata B. & Br. The writer has not seen the 
type, but Massee, on examination, declared it to be Patinella 


olivacea (C nigro-olivacea). On the basis of the same evidence 
the same writer included P. violacea B. & Br. and P. hirneola 
B. iSc Br., also from Ceylon, as synonyms. In the original descrip- 
tion of P. applanata the authors mentioned a "United States 
specimen." Material so labeled in the herbaria of Fries and the 
Philadelphia Academy of Science, collected in South Carolina, 
by Curtis, has been examined, and certainly belongs here. Addi- 
tional collections reported from Connecticut (Wright), Pennsyl- 

vania (M 

(Curtis) have not been 


No specimen of Rhizina nigro-olivacea Curr. has been available 

to the w^riter. 


after examination of the type) place it as a synonym of P. olivacea, 
there seems to be no doubt of its identity with C. nigro-olivacea. 

Peziza fuse ocar pa Elb & Hoi. w^as described from material col- 
lected in Iowa by Holway. It has been referred to Phaeopezig^ by 
Saccardo, to Humaria by Morgan, and to Aleurina by Saccardo & 


Sydow, and by Rehm. This specific name is the one under which 
the plant here considered has commonly been reported or dis- 
tributed in America. The type is identical with Peziza nigro- 
olivacea L. v. S. The writer has received it from many localities 
and has had abundant opportunity to study it in the fresh con- 
dition in all stages of development. 

Of all the references of the present species made by Saccardo, 
the most curious is that where Peziza nigro-olivacea L. v. S. is 
made a synonym of Mollisia umbonata (Pers.) Sacc. To Per- 
soon's specific description are appended verbatim Fries's observa- 
tions comparing Schweinitz's plant with Patellaria ptilla Fr., a ^ 
totally different plant from Persoon's. This confusion may be 
due to a mixing of notes. 

20 Durand: The genus Catinella 

Finally, the plant here considered was described as Humaria 
marchica by Rehm, from German material distributed by Sydow 
and examined by the writer. 

While it has been demonstrated that Catinella nigro-oUvacea 

has passed under about thirty different names, it may well be that 
others have escaped the writer's attention. 

In the citation of specimens the following abbreviations have 
been used: CU, Herbarium of Cornell University; D, Durand 
Herbarium; F, Fries Herbarium; K, Kew Herbarium; Mo, Her- 
barium of the Missouri Botanical Garden; NY, Herbarium of the 
New York Botanical Garden; Phil, Philadelphia Academy; S, 
Schweinitz Herbarium. 

Material examined 

Russia: Petersburg (F, type of Bulgaria nigrila Fr.). 

Germany: Brandenburg, Sydow (NY, cotype of Humaria 
marchica Rehm, in Sydow, Myc. March. 295^); Moravia, Petrak 
(D, as Aleurina olivacea [Batsch] v. Hohn.), 

Ontario: Carleton Place, Macoun (NY); London, Dearness 
(CU, D, E, & E. N. Am. Fungi 2325). 

New Hampshire: Warren, L. W, Riddle (D). 

New York: Ithaca, Atkinson (D); Varna, Reddick (D); 
Honeoye, Durand (D); Adirondack Mountains, Catskill Moun- 
tains, Old Forge, Kasoag, Big Indian, South Pond, Elisabethtown, 
Peck (A, as Patellaria olivacea), 

Pennsylvania: Bethlehem, Schweinitz (S); Springtown, Witte 
(NY); Mauch Chunk (Phil) ; Chester County, Michener (Phil). 

North Carolina: Blowing Rock, Durand (D), 

South Carolina: Curtis (F, Phil). 

Louisiana: Cloutierville & Abita Springs, Langlois (NY). 

Arkansas: Camden, C. /. Humphrey (D). 

Ohio: Preston, Morgan (NY). 

Minnesota: Lakeville (D) ; St. Louis River, Holway (XY, Mo). 

North Dakota: Fargo, Seaver (NY, Mo, N. Dak. Fung. ^5, 
as Phaeopezia Juscocarpa) . 

Iowa: Mt. Pleasant, Seaver (D); Decorah, Holway (NY, type 
of Peziza Juscocarpa). 

Cuba: Wright 360 (K, type of Peziza viridi-atra) 

Durand: The genus Catinella 21 

2. Catinella elastica (Pat. & Gail.) Durand, comb. nov. 


Phaeopezia elastica Pat. & Gail. Bull. Soc. Myc. Fr. 4: 99. 


Aleurina elastica (Pat. & Gail.) Sacc. & Syd. Syll. 16: 739. 
1902; Rehm, Ann. Myc. i: 515. 1903. 
Ascomata sessile, solitary or in groups, 1-3 mm. in diameter, 

rich dark brown externally, black within, attached at the base by 
radiating brown fibers consisting of septate hyphae; margin 
thickened. Excipulum parenchymatous, of polygonal cells, the 
ectal ones with brown walls, those at the sides of the cup giving 
rise to short, septate, flexuous, brown pili, those at the base to 
stout, brown, radiating hyphae, Asci cylindric-clavate, apices 
rounded, not blue with iodine. Spores eight, uniseriate, elliptic, 
or elliptic-clavate, or slipper-shaped, continuous, brown, two- 
guttulate, 13-15 X 6-7 ju. Paraphyses cylindric, hyaline, septate, 
tips agglutinated with amorphous matter. 

Material examined 

Venezuela: on dead wood, Mapire, May, 1887, A, Gaillard 6 


This species seems to differ from the preceding only in the 
larger spores. In the herbarium of the New York Botanical 
Garden is a specimen of what appears to be the original and only 
collection. The above description is that of the authors supple- 
mented by notes drawn from the above mentioned specimen. 
In the original description the tissue is spoken of as elastic, some- 
what like that of Bulgaria. The dried material is friable. It 
breaks up so that no complete ascus has been seen. The material 
becomes deep violet on treatment with KOH. 

University of Minnesota 



The aim of this Index is to include all current botanical literature written b> 
Americans, published iq America, or based upon American material ; the word Amer- 
ica being used in the broadest sense. 

Reviews, and papers that relate exclusively to forestry, agriculture, horticulture, 
manufactured products of vegetable origin, or laboratory methods are not included, and 
no attempt is made to index the literature of bacteriology. An occasional exception is 
made in favor of some paper appearing in an American periodical which is devoted 
wholly to botany. Reprints are not mentioned unless they differ from the original in 
e important particular. If users of the Index will call the attention of the editor 
to errors or omissions, their kindness will be appreciated. 

This Index is reprinted monthly on cards, and furnished in this form to sub- 
scribers at the rate of three cents for each card. Selection of cards is not permitted: 
each subscriber must take all cards published during the term of his subscription. 
Correspondence relating to the card issue should be addressed to the Treasurer of 
the Torrey Botanical Club. 

Andres, H. Piroleen-Studien. Verh. Bot. Ver. Brandenburg 56: 1-70. 
6 Je 1914. 

Arber, A. Leaves of the Helobieae. Bot. Gaz. 72: 31-38. pL 7- 
16 Jl 1921. 

Arthur, J. C. Specialization and fundamentals in botany. Am. Jour, 
Bot. 8: 275-285. 30 Je 1921, 

Bakhuizen Van den Brink, R. C. Revisio generis Avicenniae. Bull. 
Jard. Bot. Buitenzorg III. 3: 199-226. pL 14-22, My 192 1. 

Bamhart, J. H. Biographical notices of persons mentioned in the 
Schweinitz-Torrey correspondence. Mem. Torrey Club 16: 290- 
300. 16 Jl 192T. 

Blake, S. F. A new Aspilia from Trinidad- Proc, Biol. Soc. Washing- 
ton 34: 119,120. 30jei92i. 

Blake, S. F. New Meliaceae from Mexico. Proc. Biol. Soc. Washing- 
ton 34: 115-118. 3oJei92i. 

New species in Cedrela (i) and Guarea (2). 

Bradshaw, R. V. Oregon willows. Am. Hot- 27; 56-59. My 1921. 


24 Index to American botanical literature 

Britton, N. L. The cacti of Trinidad. Bull. Dept. Agr. Trinidad & 

Tobago 19: 81-87. 192 1. 

Carano, E. Nuove ricerche sulla embriologia delle Asteraceae. Annali 
di Botanica 15: 97-196. pi. 4-12 -f /. i, 2, 28 F 1921. 

Campbell, D. H. Professor H. Bruchmann. Science 11. 54: 67, 68. 
22 JI 1921. 

Campbell, D. H. The eusporangiate ferns and the stelar theory. Am. 

Jour. Bot. 8: 303-314-/' ^'7- 3^ Je 1921. 
Campbell, D. H. Die Verbreitung gewisser Lebermoose der malaiischen 

Region. Jahrb. Wissensch. Bot. 56: 365-373. 1915. 

Chamberlain, C. J, Helmut Bruchmann. Bot.. Gaz. 72: 45-47. 16 
Jl 1921. [With portrait.] 

Child, C. M* Certain aspects of the problem of physiological correla- 
tion. Am. Jour. Bot. 8; 286-295. 30 Je 1921. 

Clark, S. P- Sweet clover in Arizona. Univ. Arizona Coll. Agr. 
Circ. 34: 1-7. Ap 1921. 

[Clute, W. N.j A new form of Denlaria. Am. Bot. 27; 69. My 1921. 

Deniaria laciniata var. dichronum from Iowa. 

Collins, J. L. Inbreeding and crossbreeding. Jour. Hered. 12: 89-93, 
/. JI-J4- II Jl 1921. 

The effect of inbreeding and crossbreeding in a wild plant [Crepis capillaris] 

i i 

of the sunflower family.** 

Conard, H. S. An outline for vascular plants. Science II. 54: 15, 16, 
I Jl 1921. 

Cook, M. T., & Martin, W. H. Potato diseases in New Jersey. New 
Jersey Agr. Exp. Sta. Circ. 122: 1-39./. 1-21. F 1921* 

Cook, M. T., & Poole, R. F. Diseases of sweet potatoes. New Jersey 
Agr. Exp. Sta. Circ. 123: 1-24./. 7-77. Ap 1921. 

Douin, C. La famille de Cephaloziellac^es. Soc. Bot. France Mem. 
29: 1-90./. i~io2. 28 D 1920. 

Includes as American novelties, the genus Evansia, and species in Prionolobus (i), 
Evansia (i), and Cephaloziella (9). 

East, E. M., & Jones, D. F. Round tip tobacco— a plant "made to 
order." Jour. Hered. 12: 51-56. pL J +/. i~S, 11 Jl 1921, 

Felt, E. P. Key to American insect galls. New York State Mus. Bull. 
200: 1-310. pL 1-16 +/. 1-250. I Au 1917. 

Fernald, M. L. The geographic distribution of hybrids. Science IL 
54: 7Z. 74. 22 Jl 1921. 

Folsom, D. Potato leafroll. Maine Agr. Exp. Sta. Bull. 297: 37-52. 
/. 26-33^ Ap 1921. 

Index to American botanical literature 25 

Fromme, F. D., & Wingard, S. A. Varietal susceptibility of beans to 
rust. Jour. Agr. Research 21: 385-404. pL 69-73. 15 Je 1921, 

Glover, W. O. Blister canker of apple and its control. New York Agr. 
Exp. Sta. Bull, 485: 1-71. pL i-i^ +/. i~8. Ja 1921. 

Hansen, A. A. Protect our vanishing wild plants! Conservationist 4: 
67-70. My 1921. [Illust.] 

HarmS| H. Uber die Fluoresceriz des Aufgusses der Rinde von Pninus 
serotina. Verh. Bot. Ver. Brandenburg 59: 146-150. 15 Ja 1918. 


Harms, H. Uber Fluorescenz-Erscheinungen bei dem Holze der 
Leguminose Eysetihardtia amorphoides H. B. K. Verh. Bot. Ver. 
Brandenburg 56: 184-197.' 18 Ja 1915. 

Harms, H, Nachtrage und Verbesserungen 2u meinem Aufsatze iiber 
Fluorescenzerscheinungen. Verh. Bot. Ver. Brandenburg 57: 191- 
202. I Mr 1916, 

Harms, H. Ernst Ule. Verh. Bot. Ver. Brandenburg 57: 150-184. 
I Mr 1916. [With portrait.] 

Harris, J. A. Tissue weight and water content in a tetracotyledonous 
mutant of Phaseohis vulgaris, Proc. Soc. Exper. Biol, and Med. 18: 
207-209, 192 1. 

Harvey, E. M., & Mumeek, A. E. The relation of carbohydrates and 
nitrogen to the behavior of apple spurs. Oregon Agr. Coll. Exp. 
Sta. Bull. 176: 1-47./. 1-12. Ap 1921. 

Hayes, H. K., & Gardner, R. J. Breeding crop plants. Pp. i-xvii 
+ 1-328./. 1-66. New York. 1921. 

Hume, A. M., Loomis, H., & Hutton, J- G. Water as a limiting factor 
in the growth of sweet clover {M[elilottis] alba). South Dakota 
Agr. Exp. Sta. Bull. 191 : 257-298. pL J, 2. ]\ 1920. 

Hutchinson, C. B. Heritable characters of maize — VII. Shrunken 
endosperm. Jour. Hered. 12: 76-83./. 20-24. n J' 192I' 

Hutchinson, J. The genera of Fumariaceae and their distribution. 
Kew Bull. Misc. Inform. 1921: 97-115. 1921. 

Illick, J. S. The white pine. Am. For. 27: 422-426. Jl 1921. 



Jones, L. R., Walker, J. C, & Tisdale, W. B. Fusarium resistant 
cabbage, Univ. Wisconsin Agr. Exp. Sta. Research Bull. 48: i-34- 
/. i~io. N 1920. 

Judd, C. S. The Alahee tree. Hawaiian For. & Agric. 18: I33-I37- 
Je 1921. [Illust.] 

26 Index to American botanical literature 

Kraemer, H. Plant colors. Am. Jour. Pharm. 93: 414-416. Je 1921. 

Kraemer, H. Some experiments on the modification of color in plants. 
Am. Jour. Pharm. 93: 416-418. Je 1921. 

Kunkel, L. O. A possible causative agent for the mosaic disease of 
corn. Bull. Exp. Sta. Hawaiian Sugar PL Assoc. 3: 1-15. ph 4-15 
+ f. J, 2. 9 Jl 1921. 

Lee, H. A. The relation of stocks to mottled leaf of Citrus trees. 
Philipp. Jour. Sci. 18: 85-93. pL 1-3, Ja 1921. 

Loesener, T. Plantae Selerianae. — IX. Verh, Bot. Ver. Brandenburg 
58: 129-157. I My 1917. 

Includes new American species in Nemaslylis (i). Iresine (i), Dapknopsts (i), 
and Aniseia (i). 


Lonay, H. Sur qu61ques genres rares ou critiques de Renonculac&s. 
Bull. Soc. Bot. Belgique45: 191-204. 4 D 1908. 

Long, W. H- Notes on new or rare species of rusts. Bot. Gaz. 72: 
39-44. 16 Jl 1 92 1. 

Includes new species in Gymno sporangium (i), and Ravenelia (3). 

MacDougal, D, T. Water deficit and the action of vitaniines, amino- 
compounds, and salts on hydration. Am. Jour. Bot. 8; 296-302. 
30 Je 192 1. 


Martin, W. H. Studies on tomato leaf-spot control. New Jersey 
Agr. Exp. Sta, Bull. 345: 1-42. pL i + f. i. N 1920. 

Maxon, W. R. A neglected fern paper. Proc. Biol. Soc. Washington 
34: 111-114. 30 Je 1921. 

Refers to Watt's Catalog of Ferns, published in Canadian Naturalist II. 13: 
157-160. 1867. 

Merrill, E. D. New Philippine Moraceae. Philipp. Jour. Sci. i8: 
49-69. Ja 192 1. 

Merrill, E. D. New Philippine Myrtaceae. Philipp. Jour, Sci. 18: 
289-308. ]\Ir 1921. 

Mulford, F- L. Oaks for ornamental planting. Am. For. 27: 461-467- 
Jl 1921. [Illust.] 

Nash, G» V, Acacia armata. Addisonia 6: 13. pL igg, 29 Je 1921- 

Nash, G. V. Acacia leprosa. Addisonia 6: 7. pL iq6. 29 Je 1921. 

Nash, G. V. Acacia lineata. Addisonia 6: 15. pL 200. 29 Je 1921. 

Nash, G. V. Acacia longifolia floribunda. Addisonia 6: ii- pL igS^ 
29 Je 1921. 

Nash, G. V. Acacia Nabonnandi. Addisonia 6: 9. pL ig/. 29 Je 

Index to American botanical literature 27 

Nash, G. V. Acacia pubescens, Addisonia 6: 3, 4. pL IQ4. 29 Je 

Nash, G. V. Acacia pitlchella. Addisonia 6: 5. pL ig$. 29 Je 1921. 

Nash, G. V. Acacia scaptiliformis, Addisonia 6: i, 2. pL iqj. 29 Je 

Nelson, E. W. Lower California and its natural resources. Mem. 
Nat. Acad. Sci. 16^; 1-171. pi. 1-35. 1921. 


Neville, H. O. Pineapples in Cuba. Cuba Rev. 19: 13-28. Jl 1921. 

Osterhout, W. J. V., Thaxter, R,, & Feraald, M. L. Lincoln Ware 
Riddle. Science IL 54: 9. i Jl 1921, 

Parish, S. B. A reminiscence of roses. Am. Bot. 27: 60, 61. My 1921. 

Patouillard, N. Clathrotrichum, nouveau genre d'hyphomyc^tes. Bull. 
Soc. MycoL France 37: 33-35. 15 Ap 1921, 

Parasitic on Selaria in Ecuador. 

Pittier, H. Acerca del genero Gyranthera Pittier. Bol. Com. e Indust. 
[Venezuela] 13: 428-433. Ja 1921. 

Includes G. caribensis sp. nov. 


Pittier, H. Arboles y arbustos nuevos de Venezuela — L Bol. Com. e 
Indust. [Venezuela] 13: 417-428. Ja 1921. 

Includes new species in Inga (i). Raputia (i). Esenheckia (i). Trichilia (i), 

Stillingia (i), Gustavia (i). Mouriria (i), Oxythece (i), Pouteria (i), and Sider- 
oxylum (i). 

Rapp, C. W. Bacterial blight of beans. Oklahoma Agr. Exp. Sta. 

Bull. 131: 1-39-/- -r-//. Jl 1920. 
Reinking, O- A. Citrus diseases of the Philippines, southern China, 

Indo-China, and Siam. Philipp. Agric. 9: 121-179, pL 1-14. F 

Reinking, O. A. Diseases of economic plants in Indo-China and Siam. 
Philipp, Agric. 9: 181-183. F 1921. 

Rolfe, R. A. Wercklea Intea [Malvaceae]. Kew Bull. Misc. Inform. 
1921: 118, 119. 1921. 

New species from Costa Rica. 

Rydberg, P. A. Tw^o new species from Arizona. Am. Bot. 27: 61-63. 
My 1 92 1. 

Eriogonum CliUei Rydb. and Foresiiera arizonica (A. Gray) Rydb. 

Sands, W. N. Plants common to the West Indies and Malaya. Agric. 
News 20: 163; 182, 183. 28 My 1921. 

_ k 

28 Index to American botanical literature 

Schmitz, H., & Daniels, A. S. Studies in wood decay — I. Laboratory 
tests on the relative durability of some western coniferous woods 
with particular reference to those growing in Idaho. School For- 
estry Univ, Idaho Bull, i: i-ii. Jl 1921. 

Schmitz, H. Studies in wood decay — IL Enzyme action in Polyporus 
volvatus Peck and Fomes ignarius (L.) Gillet. Jour. Gen. Physiol. 3: 
795-800. 20 Jl 1 92 1. 

Shear, C. L., & Stevens, N". E. (Editors). The correspondence of 
Schweinitz and Torrey. Mem. Torrey Club 16: 119-300, ph 6, 7. 
16 Jl 1921. 

Sherman, H. Respiration of dormant seeds. Bot. Gaz. 72: 1-30. 
/. 1-4. 16 Jl 1921. 

Soth, M. E. Goat-weed in Idaho. Am. Bot. 27: 70. My 1921. 

Sousa Brito, E. C. de. Algumas plantas uteis do Brasil, o hybridismo 
de uma dellas. Arch. Esc. Sup. Agric. e Med. Vet. .4: 25-54. pL 
10-16. D 1920. 

Stadler, L, J., & Helm, C. A. Corn in Missouri — -L Corn varieties 

and their improvement. Missouri Agr. Exp. Sta. Bull. 181: 1-51. 
/. J-7. Mr 1921. 

Stahel, G. De Sclerotium-ziekte van de Liberiakoffie in Suriname 
veroorzaakt door Sclerotium coffetcolum nov. spec. Dept. Landb. 
Suriname Bull. 42: 1-34. pL i-ii, Ja 1921. 

Stevens, F. L. The relation of plant pathology to human welfare- 



Stout, A. B, Types of flowers and intersexes in grapes with reference 
to fruit development. New York Agr. Exp. Sta. Techn. Bull. 82: 
1-16. pL 7-7. Ja 1921. 

Sydow, H. & P. Novae fungorum species — XVL Ann. Mycol. 18: 
154-160. Ap 1921. 

Includes new American species in Septohasidium (i). Uromyces (i). Aecidium (i), 
Catacauma (i), Taphrina (i), 

Taubenhaus, J. J, A study of the black and the yellow molds of ear 

Bull. Texas. Agr. Exp. Sta. 270: 3-38./. i-io. O 1920. 

aus, J. J., & Mally, F. W. Pink root disease of onions and its 


control in Texas. Texas Agr. Exp. Sta. Bull. 273; 1-42. / 


W. R. Additions to the flora of Mount Desert, Maine. Rhodora 

23: 65-68. 20 Ap 1921. 

Index to American botanical piterature ' 29 

Thatcher, R. W. The chemistry of plant life. Pp. i-xvi + 1-268, 
New York. 1921. 

Thomson, G. E. Hegari in Arizona. Univ. Arizona Coll. Agr. Circ. 
33: 1-4. Ap 1921. [Illust.] 

Thomas, R. C. Botrytis rot and wilt of tomato. Bull. Ohio Agr. Exp. 
Sta. 6: 59-62. Ap 1921. [Illust.] 

Thurston, H. W. A note on the corrosive sublimate treatment for 

the control of Rhizoctonia. Phytopathology 11: 150, 151. Mr 

Tiffany, L. H. Algal food of the young gizzard shad. Ohio Jour, Sci, 
21: 113-122. F 1921. 

Tiffany, L. H. New forms of Oedogonium. Ohio Jour. Sci. 21: 272, 
273. pL J. Je 1921. 

2 new si)ecies from Ohio. 

Tisdale, W. H,, & Jenkins, J. M. Straighthead of rice and its control. 
U. S. Dept. Agr. Farm. Bull. 1212: I~i6. /. 1-6. Je 1921. 

Tourney, J. W. Damage to forests and other vegetation by smoke, 
ash and fumes from manufacturing plants in Naugatuck Valley, 
Connecticut. Jour. For. 19: 367-373./. j~6. Ap 1921. 

Trelease, W. The Jack Oak {Quercus ellipsoidalis). Trans. Illinois 
Acad. Sci. 12: 108-118. pL 1JQ-14J, 1919- 

Trelease, W« North American pipers of the section Ottonia, Am. 
Jour. Bot. 8: 212-217. P^- 5~^' 3^ i\p 1921. 

Tucker, E. M. Bibliographical notes. Jour. Arnold Arbor. 2: 181- 
184. Ja 1921. 

Discussion of issuance of Nouveau Duhamel and of Torrey and Gray's Flora of 
North America. 

Tucker, E. M. Catalogue of the library of the Arnold x'\rboretum of 


Harvard University. Publ. Arnold Arbor. 6. l: 1-782. 1914; 
2: 1-542- 1917- 

Compiled under the direction of Charles Sprague Sargent. 


Ule, E. Unter den Indianern des nordlichen Amazonas- Verh. 
Naturw. Vereins Hamburg HI. 21: LXXVI, LXXVII. 1914. 

A botanical and anthropological journey. 

Uphof, J. C. T. Die Waldflora im Staate Washington. Vegations- 

bilder XHI. 7: // pp. + pL 37-42* 1920. 
Urban, I. Novitates haitienses. Notizbl. Bot. Gart. und Mus. Berlin- 

Dahlem 7: 495-499- I Mr 1921. 

Includes new species in Pilea (i), Phthirusa (i). Cassia (i), Sehastiana (i)- 
Ahutilon (i) and Eugenia (i). 

30 Index to American botanical literature 

Urban, L tfber die Malvaceen-Gattung Montezuma Moc. et Sess6. 
Notizbl. Bot. Gart, und Mus. Berlin-Dahlem 7: 543, 544. I Mr 

Wakefield, E. M. Mosaic diseases of plants. West Indian Bull. 18: 
197-206. [192 1,] 

Walker, E.R. The gametophytes of Eqatsettim laevigatiim, Bot. Gaz. 
71: 378-391- Pl' 2Ji 24 +/. 1-3. 16 My 1921. 

f 1- 

Walker, J. C. Onion smudge. Jour, Agr. Research 20: 685-721.^/. 

SO'Ss +/. i~io. I F 1921. 

Walker, J. C. Rust of onion followed by a secondary parasite- Phyto- 
pathology 11: 87-90./. J, 2. F 1921. 

Weatherby, C. A. A form of Ilex opaca. Rhodora 23 : 1 18, 1 19. 19 My 

Weir, J. Rv Thelephom terrestris, T. fimbriata, and T. caryophyllea 

■ L 

on forest tree seedlings. Phytopathology 11 : 141-144. pL 5, Mr 

Wells, B, W. Evolution of zoocecidia. Bot, Gaz. 71: 358-377. pL 21^ 
22. 16 My 1921. 

Wells, B.W. A phenomenal shoot. Science II. 54: 13, 14. I Jl 192 1. 

Patilownia tomentosa. 


Weston, W. H. Another conidial Sclerospora of Philippine maize. 
Jour, Agr. Research 20: 669-684. pi. 76-7g. 1 F 1921. 

Whaley, W- Properties of wood. A chapter from Lucien Chancerel's , 
Botanique Forestiere. Jour. For. 19: 414-420. Ap 192 1. 

White, O. E. The pollination of flowers. Brooklyn Bot. Gard. Leafl. 
9 n. 3, 4: 1-15. 4 My 1921. 

Wiegand, K. M, Amelanchier amabilis, a new name. Rhodora 23: 
48. 5 Ap 1921. 

Wiegand, K. M. Echinochloa in North America. Rhodora 23; 49- 
65. 20 Ap 1921. 

Includes E. faltidigena sp. nov. and several new varieties. 

Wieland, G- R. Monocarpy and pseudomonocarpy in the cycadeoids. 
Am. Jour. Bot. 8; 218-230. pL g-12. 30 Ap 1921. 

Williams, C. B. Report on the Froghopper Blight of sugar-cane in 
Trinidad. Mem. Dept- Agn Trinidad and Tobago i: 1-170. pi- 
I'll +/. 1-32. Ja 1921. 

Considers also **The Root Disease of Sugar Cane" caused by fungi. 

Yunker, T. G. A Curious abnormality in Ciiscula cvspidata. Am. Bot. 
27: 48, 49. My 1921. Illlust.] 

Bull. Torrey Club 

Volume 49, plate i 





•"■"^^^J T M lM - ^ ^^^^ 


4 ^ 

L /Jir^i^^'cfi^jf "i-i^-yWi-^ 



Vol. 49 

No. 2 





^ Miscellaneous notes on plants of Southern California — I 

Philip A, Munz and Ivan M. Johnston 

Recent botanical collecting has added to our knowledge of the 
flora of Southern California and has made possible the presenta- 
tion of the notes given in this paper. Collections cited as having 
been made by '* M & H" were by Munz and Harwood, those by 
" ilf , J &f H^' were by Munz^ Johnston and Harwood ; other coUec- 
tions are not abbreviated- Specimens of all the plants mentioned 
are in the C. F. Baker Herbarium of Pomona College. 


Notkolaena sinuata var. integerrima Hook. Sp. Fil. 5: 108. 1864. 
Locally abundant on a rocky hillside in a gulch back of the 
Bonanza King Mine, Providence Mountains, Mohave Desert, 
M &" Hjsso and M,J&H 4224. Our material is a fine match for 
the figure given by Eaton (Ferns of North America pL jp, f. i. 
1879). New to California. 

WoODSiA scopuuNA D. C. Eaton 

Woodsia scop 


This fern, infrequent in the higher mountains of the middle 
and northern parts of the state, must be accredited a place in the 
fern-Hst of Southern California, as a few plants were collected in 
the Providence Mountains several miles south of the Bonanza 
King Mines. The single colony discovered, M, J & H 4212, was 
growing in the shelter of boulders in the bed of a dry gulch in the 
lower portions of the pinyon belt. The soil in which the plant 

iThe Bulletin for January (49: 1-30. pi. . 


32 MuNZ AND Johnston: Plants of California 

grew was damp, not saturated, gravel; a water hole was situated 
a few hundred feet down the canyon and there was apparently a 
meager supply of water from underground sources. On the 
adjacent canyon walls were such typical xerophilous ferns as 
Notholaena tenera, M, J &" H 42J2; N. Parryi, Mj J & H 42j6; 
Cheilanlhes Feei, M, J & H 421$; C Covillei, M, J & H 4015; 
Gymnogramme triangularis ^ M^ J & H 4021; and Pellaea compacta, 
M, J & H 4018. 

Notholaena californica D. C. Eaton 

Notholaena californica t). C. Eaton, Bull. Torrey Club 10: 27. 


The junior author collected a few plants of this fern high upon 
a precipitous south-facing cliff at Deadman Point, about ten miles 
southeast of Victorville. The collected plants were associated 
with an abundance of Notholaena Parryi and with fewer Cheilanthes 
Covillei and Gymnogramme triangularis. This collection now 
marks the most northern station for this species, at least in Cali- 
fornia, and at the same time makes the species first known as an 
elernent of the Mohave Desert flora. Maxon in his recent dis- 
cussion ( Contr. U. S, Nat. Herb. 17: 603. 1916) has cited many 
specimens that show the distribution of this neat little plant. 
Johnston (Bull. So. Cal. Acad. 17: 64. 1918) has recorded a 
second coastal and what is at the same time the most westerly 
known station on the mainland. 

Triodia mutica (Torr.) Benth. 
Tricuspis muticus Torr. U. S. Rep. Expl. Miss. Pacif. 4 : 156. 1857. 
Triodia mutica Benth.; Wats. Proc. Am. Acad. 18 : 180. 1883. 

Abundant on a rocky pinyon-clad hillside near the Bonanza 
King Mine, in the Providence Mountains, M, J &" H 4132. Pre- 
viously known in California only from a collection made in the 
central Sierra Nevada (cf. Jepson, Fl. California 141. 1912). 

Determination bv Mrs. Chase. 

Agave utahensis Engelm. 

Agave utahense Engelm. Bot. King. Exp. 497. 1871. 

Common in rocky places in the pinyon belt of the Providence 

Mountains. M. 


MuNZ AND Johnston: Plants of California 33 

Brandegee (Zee 5: 153. 1903). Mulford 
made by Parish at Ivanpah (Rep. Miss( 

1 896) . 

Yucca baccata Terr. 


Mex. Bound. Surv. 221, 1859. 
necies in the Providence Mounta 


1918) and at 

r ■ 

a much eariier date by Dr. Engelmann (Trans. St. Louis Acad, 3 : 
44- 1 873)? so that all that remains for us to do is to add a few 
notes on its habits* This species and F. mohavensis are very 
common in the vicinity of the Bonanza King Mine but, although 
growing in the same region, differ widely in habitats and 
habits, as well as in technical characters. F. baccata grows on 
rocky, sunny hillsides in the pinyon belt and is a very late-flowering 


plant; for but few individuals were in flower during our visit to 


the mountains from May 21 to 24, 1920. On the other hand, 
F. mohavensis grows along the foot of the mountains, below the 
pinyon belt, and flowers very early; for on the May visit fruit 

r ■ ■ 

was set, flowering having been observed on an earlier visit during 
the last week in March. Our observations fully substantiated 
Mr, Parish's statement that F. baccata was recognizable in the 
field by the lighter color of the leaves and the universal acaulescent 

Structurally the two species are very distinct. Y. baccata has 
large campanulate flowers, whose perianth segments do not spread 
frorn the base but remain ^rect and appressed about the ovary 
for a short distance before spreading, thus suggesting the tubular 
condition characteristic of the genus Samtiela Trel. In F. 


mohavensis the shorter segments spread from the very base and 
are more curved, thus making the flowers globose in shape. 

The meager material that is available of F. baccata suggests 
that there may be a major and a minor flower form. Mr. Brande- 
gee*s collection, reported as F. mohavensis (Zoe 5: 153. 1903), 

+ _ 

Lemmon's 1884 collection at Ft. Mohave, and our If, / ^ H 4114 
all represent plants with conspicuously large flowers, the perianth- 
segments of which vary between 8 and 9.5 cm. in length. On the 
other hand, Parish 102S1 from the north end of the Providence 
Mountains and our M, J & H 41 ij from the south end of these 

34 MuNz AND Johnston: Plants of California 

mountains have strikingly smaller corollas, the perianth-segments 
measuring 6-7 cm. in length. Under the original description 
Dr. Torrey quotes Bigelow to the effect that this species has 
perianth lobes 2^^ or 3 inches (6.25-7.5 cm.) long, while Wooton 
and Standley (Contr. U. S. Nat. Herb. 19: 135. 1915) give them as 



1902) says that they are "about 75 mm." long; these facts would 
indicate that our large-flowered plants have flowers large for the 
species. It is to be hoped that future collections will be made 
with especial attention to this species, In order to ascertain the 
exact nature and extent of the floral variation here indicated. 

In passing, it should be noted that the style-characters given by 
Trelease (Rep. Missouri Bot. Gard. 13 : 46. 1902 ; and Contr. U. S. 
Nat. Herb. 23: 91. 1920) and the flower- and foliage-characters 

given by Sargent (Gard. & For. 9: 104. 1896) seem to be il- 
lusionary, and that the distinctness between our western F. mo- 
havensis and the Texan plant originally called Y. haccata var. 
macrocarpa (Torrey, Lc.) is open to strong doubt. We would use 


the name F. macrocarpa (Torr.) Coville (Contr. U. S. Nat. Herb. 
4: 202. 1893) in preference to Y. mohavensis Sarg., were it not 
for Engelmann's F. macrocarpa (Bot. Gaz. 6: 224. 1881); the de- 
notation of F. macrocarpa Engelm. is uncertain from the literature 
(Trelease, op. ciL 98. 1902; Sargent, lx.\ and Engelmann, Bot. 
Gaz. 7: 17. 1882); and it Is thought best to make no change in 
our usage until it has been definitely decided whether Engel- 
mann's species be a synonym of F, Schottii or actually identical 
with Torrey 's var, macrocarpa, as Wooton and Standley have 


treated it* In other words we use F. mohavensis in a wide sense 
and Include therein F. haccata macrocarpa Torr., though appreci- 
ating the fact that F. mohavensis may not be the correct name for 


such a concept 

Dr. Merriam (N. Am. Fauna 7: pi. 12, 14. 1893) has given 
two plates that show the very different forms which characterize 
F. haccata and F. mohavensis. Although his plate of the latter 
represents the dominant and ordinary phase of that species, yet 
we observed a phase with unbranched trunks and usually longer 
leaves, whose habit is strongly suggested by the plants of a very 
different species, figured In pi. j of the third report of the Missouri 
Botanical Garden. 

MuNZ AND Johnston: Plants of California 35 

Naias flexilis (Willd.) Rostk. & Schmidt 

Catilinia flexilis Willd. Abh. Akad. Berlin 95. 1803. 

Naias flexilis Rostk. & Schmidt, Fl. Sedin. 382. 1824, 

Previously reported by Davidson from Soldiers' Home' near 

Los Angeles; collected in 1919, Street & Williams 2688, at Laguna 

Lakes near Laguna Beach, about fifty miles southeast of earlier 

records. The shallow pools known as the Laguna Lakes have a 

rather interesting lot of aquatic plants, such as Zannichellia 

palustrisy Echinodorus cordifolitis, Marsilea vestita, and Elatine 

Eriogonum apiculatum Wats. 

Eriogonum apiculatum Wats. Proc. Am. Acad. 17: 378. 1882. 

This species, said by Jepson (Fl. CaHfornia 407. 1914) to 
occur on ''Mt. San Jacinto, 7800 to 8200 feet" and on '*Cuya- 
maca Mt.," has been collected at 5,500 feet, August 5, 1918, Mrs. 
Mary F. Spencer p8g, in open woods on Palomar Mountain in San 
Diego County. This station, in addition to being of altitudinal 
interest, is considerably west of the others mentioned. 

MOLLUGO verticillata L, 

Mollugo verticillata L. Sp. PI. 89. 1753. 

Said by Jepson (Fl. CaHfornia 460. 1914) to be ''sparingly 
naturalized" and given as collected by Davidson in Los Angeles. 
Abrams (Fl. Los Angeles 127. 1917) names also Laguna in Orange 
County. We can report this plant from near Beaumont, on the 
line between Riverside and San Bernardino Counties, Mtinz 2614^ 
March 1919, growing on a dry hillside in the chaparral at about 
2000 feet. 

Herniaria cinerea DC. 

Herniaria cinerea DC. Fl. Fr. Suppl. 375. 1815. 

This species, naturalized from southern Europe, can now be re- 
ported from Southern California, Munz & Johnston 4406, Pomona 
College Campus, Claremont, where it is fairly well established, but 
threatened by impending grading and building. Our plants are 
well matured and some of them had ripened their seeds by the 

end of March. 

AxEMONE tuberosa Rydb. 

Anemone tuberosa Rydb. Bull. Torrey Club 29: 151. 1902. 

Previously recorded from California as collected in the Pana- 

mint Mountains, 1891, Coville & Funston; and in 1849 by Fremont, 

36 MuNz AND Johnston: Plants of California 

locality not given. It was found abundant at the eastern base of 
the Providence Mountains on rocky slopes just below the pinyon 
belt, M. & H 3555 and M, J. & H 4207. 

Myosurus cupulatus Wats, 

b ■ 

Myosurus cupulatus Wats. Proc. Amer. Acad. 17: 362, 1882. 

Canyon side in the Upper Sonoran Zone at 4,000 feet, Provi- 
dence Mountains, M, J & H 4152, May 22, 1920; at this time 
the plants were quite dead, having matured their seeds. Pre- 
viously known only from a few collections in Arizona, 

Jepsonia Parryi (Torr..) Small 
Saxijraga Parryi Torr. Bot. Mex. Bound. Surv- 69, pL 25. 1859. 
Jepsonia Parryi Small, Bull. Torrey Club 23: 18. 1896. 

In January, 192 1, Mr. Gordon Nicholson detected a very 
populous colony in a small side canyon running up into the Santa 
Ana Mountains from the Santa Ana Canyon, and very near the 
Orange-Riverside County line. The plants grew at about 1,000 
ft. altitude in clay soil, on a rather open grassy slope facing west. 
We have only specimens in leaf, but these check closely with plate 
25 in the Mexican Boundary Survey Report- The type came 
from San Diego County where It has been collected several times 
since, but outside of that county it has been known from but two 
collections, both in Riverside County. Dr. Hall found it at 
Double Butte near Winchester, and Mr. Parish has reported it 
from Palm Springs (Muhlenbergia 3: 123. 1907). 


Linum spergulinum Gray 




Mrs. Mary Sp 

is as described by Gray. 

June 25, 1920- The appendage on the petal 

HoLACANTHA Emoryi Gray 



M. J a? H 


M, J & 

Amboy on flat stony desert. Both these specimens are more 

"MuNZ AND Johnston: Plants of California 37 

compact than Emory's figures in the Mexican Boundary Survey, 
but a specimen sent us by Mr. S. B. Parish, collected by a Mr: 
Childs at the Hayfields in the Chuckawalla Valley, is of more 
open habit than the one in Emory's plate. Both of our collec- 
tions were very near the road and from large conspicuous shrubs, 
three to four feet high, forming wide gray masses of interlacing 
thorns; it seems strange that such conspicuous plants so near the 
road have been collected so seldom. Our Ludlow station is no 
doubt the one reported by Mrs. Ferris (Bull. So. California Acad. 
i8: 13. 191 9), but the one near Amboy is some distance from 
those reported by Davidson (ibid. 19: 55. 1920). 

Tragia ramosa Torr. 
Tragia ramosa Torr. Ann. Lye. N. Y. 2: 245. 1826. 

Locally common in a gravelly and stony canyon bottom in the 
Lower Sonoran Zone at the eastern base of the Providence Moun- 
tains, M, J & H. 421Q. The first collection that has been re- 
ported from California. ' ' 

Abutilon parvulum Gray 
Ahutilon parvulum Gray. PI. Wright. i:^i. 1852. 


In rocky ground at the base of the Providence Mountains near 
a deserted mining camp several miles south of the Bonanza King 
Mine; rather common locally; M, J & H 4206, First record 

from California. 

Frasera Parryi Torr. 

Frasera Parryi Torr. Bot. Mex. Bound. Surv. 156. 1859. 

Collected June 7, 1919, in the foothills of the San Gabriel 
Mountains between Live Oak and San Dimas Canyons, by Mrs. 
Fitch, C. F. Baker Herbarium No. 6594^ Long ago reported from 
"east of . . . Los Angeles" by Brewer (Bot. California i: 484. 
1876), but not again detected on the south slope of the San 
Gabriel range until the present. Frequent in the San Bernardino 
and San Jacinto ranges, where it inhabits pine-clad slopes of the 
Transition Zone. The present collection was a single plant and 
grew in the lower chaparral belt at about 1700 feet altitude. 

Salvia mohavensis Greene 
Audiberiia capitata Gray, Proc. Am. Acad. 7: 387. 1868. 
Salvia mohavensis Greene, Pittonia 2: 235. 1892. 

38 MuNz AND Johnston: Plants of California 

Collected on the crest of a rocky outlying spur of the Turtle 
Mts. southwest of Needles, M & Hj^og, It was also found rather 
commonly on rocky places in the Providence Mountains, which 
constitute the type locality. The Parish brothers long ago col- 
lected it in the mountains near Camp Cady, The Turtle Mountain 
station was at only about 1,200 feet altitude; plants were beginning 
to flower there on the first of April, In the Providence Moun- 


tains the species was observed from the base to near the crest, 
from 3,000 to 7,000 feet altitude, where only a few plants were in 


bud in the last week in May, Field observation would indicate 
that the species is restricted to rocky places, paying little attention 
to the lines between the Sonoran Zones, 

Cryptantha maritima Greene 

Krynitzkia ramosissima 


Au 1885. 
20: 277. 


Au 1885. 

Krynitzkia maritima Greene, Bull. Cal. Acad, i: 204. 
Cryptantha maritima Greene, Pittonia i: 116. 1887. 

Attention should be called to the fact that the plants currently 
going under the name of C. ramosissima Greene should properly 
be designated as C. maritima. As Coville indicated twenty-nine 
years ago (Contr. U. S. Nat. Herb. 4: 165. 1893) K. ramosissima is 
an untenable name for the plant in question, Gray having originally 
proposed it as a substitute, on the grounds of applicability, for 
the specific name racemosa. Gray clearly indicated that Eri- 
trichiiim racemosa Wats, and K. ramosissima Gray were synony- 
mous, but Greene took the latter name and gave it a wholly new 
connotation, making it cover certain misdetermined specimens 
cited by Gray under the name K, ramosissima. K. ramosissima 
Greene is a redefinition of K, ramosissima Gray, and as this latter 
unquestionably a synonym of C. racemosa, the former, technically 
having the same type as the latter, since it is merely a redefinition 
of it, cannot be considered more than an improper interpretation 

ramosissima Gray. Neither the American nor the Vienna 



currently used in today! 

C. maritima was described from Guadalupe Island specimens, 
but despite its remote insular habitat, like many other species, 

MuNZ AND Johnston:. Plants of California 39 

it IS identical with the mainland form. Gray (in the supplement 
of the Synoptical Flora, p. 428), attempted to distinguish between 
the insular plant and C ramosissima Greene, but when applied to 
a series of any size, his characters utterly fail to divide the two 
forms and certainly do not segregate the material into anything 
suggesting geographical lines. Vasey and Rose (Proc. U. S. 
Nat. Mus. 11: 532, 1888) and Brandegee (Bot. Gaz. 27: 453. 
1899) have expressed doubts as to the distinctness of C ramosis- 
sima and C maritima, but now with a fine series for. study, we 
feel that their doubts can be increased to the point of certainty. 
Every character presented by the Guadalupe specimens, or for 
that matter by any of the coastal plants, can be exactly duplicated 
in specimens from the desert interior. Though it is unfortunate 
that a name such as ''maritima'* should be applied to a plant so 

characteristic of the driest portions of the desert area, yet there 
is no technical reason for coining a new name or for accepting any 


other old one. 

Among the California representatives of the genus, C maritima 
is unique in its possession of dark reddish stems. This species, 
along with C recurvata, possesses but two ovules, a condition dif- 


degee, Z.c). 



Hillmanii Nels. & Ken. Proc. Biol. Soc. Wash 

19: 157. 1906. 

This plant must now be accredited a place in the California 
flora. It would seem that it enters the state only in the region of 



fine collection, My J & H 4222, near the Bonanza King Mine. 

C Hillmanii is very close to C. gracilis Osterhout (Bull. Torrey 
Club 30: 236.. 1903) and at most is but a poor variety of the 
latter species. Abundant collections of the species may cause 
C. Hillmanii to be reduced outright, but the single isotype of 
C. gracilis in the University of California collection seems to differ 
from all the specimens of C. Hillmanii in its lower and more slender 
habit and less congested inflorescence; therefore, for the time 

40 MuNz AND Johnston: Plants of California 

I I 

being, C Hillmanii can be retained in varietal rank. The type 
of C. gracilis came from Glenwood Springs in the mountains of 
Colorado and it is entirely possible that the differences detected 
are environmental in origin. Among California cryptanthas, 
C. Hillmanii can be recognized by its single smooth nutlet and 
usually fulvous, densely pilose and inconspicuously bristly calyx. 

^ Penstemon Munzii Johnston, sp. nov. 

Plants with several coarse, erect, loosely tufted glabrate stems 
that become at least five dm. high; leaves all opposite, entire and 
glabrate; basal leaves ovate- or lanceolate-spatulate with winged 
petioles that about equal the blade, becoming 7 cm. long and 
2.5 cm. wide; the lower cauline leaves oblanceolate, the upper 
ones broadly sessile and lanceolate^ while the leaves of the in- 
jRorescence are minute and linear-subulate; inflorescence narrow, 
the flowers in strict one- to three-flowered cymules; corolla bright 
red, 2 cm. long, narrowly funnelform-tubular, evenly though but 
slightly ampHated upward, strongly and conspicuously bilabiate, 
glabrous within; upper two lobes of the corolla erect, about 7 mm. 
long, united for~ about two-thirds of their length, lower three 
lobes of the corolla strongly reflexed, about 6 mm. long, united 
for nearly half their length; anther-sacs glabrous, obscurely rugu- 
lose or papillose, adnate if at all only near the base, ovate-oblong, 
2-2.5 mm. long, their inner sides paralleling each other or forming 
a small angle; dehiscent by a slit extending between two-thirds 
and three-fourths the way to the base, sharply dentate along the 
line of dehiscence; sterile filament glabrous, somewhat flattened, 
emarginate; sepals broadly ovate, acute, 3-5 mm. long, scarious 
margined; pedicels about as long as the sepals; fruit unknown. 

Known only from the type specimen, collected in May, 1920, 
Munz, Johnston, df Harwood 42^1, on a high exposed ridge in the 
pinyon belt near the Bonanza King Mine on the east slope of the 
Providence Mountains, Southeastern California. The type is 
No. 7534 in the Baker Herbarium of Pomona College. 

Certainly of the Barhati and probably nearest P. harbatus 
Torreyi (Benth.) Gray, but very different from it in its color, 
which is a brighter, lighter, and more yellowish red, in the meas- 

J I 

urements of the corolla, and in the size as well as non-divergence 
of the anther-sacs. P. Munzii is a lower, stouter, and less grace- 
ful plant than most of the forms of P. harbatus (Cav.) Roth. 

From P. Eatoni Gray and from P. subulatus Jones, the other 
two red penstemons found in this part of the desert area, both of 

MuNZ AND Johnston; Plants of California 41 

which are in the Centranthifolii, this species can be quickly sep- 
arated by its unique tone of red and by its smaller, stouter, de- 
cidedly bilabiate corollas. It is .a well marked species and a 


fitting addition to the interesting penstemon flora of the Providence 



V Penstemon Clevelandii var. Stephensi (Brandegee) comb. nov. 

Penstemon Stephensi Brandegee, Zoe $: 151. 1903. 

During one of our trips to the Providence Mountains we made 
a small collection of this little known plant, which is apparently 

rare in the type region; for Mr. Brandegee informs us that he 
collected only enough for the type sheet, a full one, while our 
collection, M, J & 1142/4^ was enough for only a few duplicates. 
A critical study of our collection and of the type seems to in- 
dicate that P. Stephensi finds a very close relative in P. Clevelandi 
Gray, a species restricted to the canyons about the Colorado Des- 

r ^^ 

ert and in the mountains to the west of it. Our collection of P. 
Stephensi, as well as the type, is so similar to certain collections 
of P. Clevelandi {e.g,, Hall 1160 and 2i4g and Parish Bros. 1216) 
that as far as gross aspect, size and vegetative characters are con- 
cerned, they might be of one and the same collection. There is, 
however, one character which separates the two collections of P. 
Stephensi from all of the many collections of P. Clevelandi; this is the 
lack of bearding on the sterile filament. Though the corollas of 
the two forms are alike in size, color, and shape, the sterile filamen! 
in P. Stephensi is absolutely bald, while it is densely bearded in 
P. Clevelandi. Appreciating, therefore, that we are concerned 
here only with a unit-difference, we feel that P. Stephensi Is noth- 
ing but a geographic race of P. Clevelandi and, because of it, worthy 

only of a subordinate rank. 

Penstemon Palmeri Gray 

Penstemon Palmeri Gray, Proc. Am. Acad. 7: 379. 1868. 

We found this species to be the most comnion and most 
spectacular penstemon in the Providence Mountains, M, J of H. 
4276. About the Bonanza King Mine it was common about the 
foot of the mountains and especially so in small draws, where it 
grew in large colonies. This being bur first acquaintance in the 

42 MuNZ AND Johnston: Plants of California 

field with the typical plant, and knowing quite intimately the 
low, much-branched, few-flowered montane plant, currently going 
under the name of P. Palmeri, we were much surprised to discover 
that this tall, virgate, many-flowered, desert plant was the typical 
form of P. Palmeri. While unquestionably, the montane plant 
is a close ally of the taller interior form, and as herbarium material 
shows, intergrades with it, yet we feel that so pronounced an ex- 


treme ought to be given nomenclatorial recognition, and so, for 
us, the mountain form becomes: 

*^ Penstemon Palmeri var. Grinnellii (Eastw.) comb. nov. 

Penstemon Grinnellii Eastw. Bull. Torrey Club 32: 207. 1905. 

The type of this form came from Mt. Wilson and is the form 
rather common in rocky ground in the pine belt, and to a less ex- 
tent in the upper chaparral belt, in the mountains of Southern 
California from the San Jacinto Range northward. With this 
restriction the species itself becomes strictly deserticolous. In 
its extreme the variety is characterized by its looser, shorter and 
fewer flowered inflorescence, lower and more slender stature and 
non-glaucous leaves. 

Penstemon calcareus Brandegee 

Penstemon calcareus Brandegee, Zoe 5: 152. 1903. Not Jones, 

1 908. 
Penstemon desertorum Jones, Contr, W. Bot. 12: 59. 1908. 

We succeeded in making a fair-sized collection of this peculiar 
and interesting little plant, M, J &f H. 4154^ The species is known 
from but two other collections, all of them from the Providence 

Mountains, where it erows in crevices of rorks on vprl-ir;il rliff?^. 



species; for isotypes of the former plant are exact matches for 


Mr. J 

uniform series in aspect and size, as well as in structural characters. 
P. calcareus Jones (Contr. W. Bot. 12: 60. 1908) is a homo- 

(Bot. Gaz. 27: 455. 1899). 


MuNz AND Johnston: Plants of California 43 

^Penstemon antirrhinoides var. microphyllus (Gray) comb, nov- 
Penstemon microphyllus Gray, Pac. R. R, Rep. 4: 119. 1856. 
Penstemon Plummerae Abrams, Bull. Torrey Club 33: 445. 1906. 

In California this interior form of P. antirrhinoides Benth. is 
known only from the Providence Mountains. Mr. Brandegee 
collected and reported it (Zoe 5: 151. 1903) as P. antirrhinoides, 
while we found it to be rather common on the rocky sides of a 
small canyon back of the Bonanza King Mine. Although quite 
familiar with P. antirrhinoides, as represented on the coastal slopes, 
we could not detect in this desert plant any peculiarities of habit 
or size, though while observing the plant in the field we agreed 
that there was some intangible dififerencej a difference which was 
later discovered to reside in the grayish hue resulting from a 
cinereous pubescence. 

This shrub was described by Gray from flowerless specimens 
collected by Bigelow on the ''Williams Fork of the Colorado." 
Based on insufficient material, the plant has remained an obscure 
one. It is not surprising, therefore, that Dr. Abrams, upon 
meeting so obscure a plant, treated it as though it were unde- 
scribed- It can hardly be doubted that P. Plummerae and P. 
microphyllus are synonyms, for Dr. Gray's meager description 
applies completely to P. Plummerae, while the types of both are 
from the same phytogeographical area. Significant evidence is 
found also in the fact that this is the only shrubby Penstemon re- 
vealed by recent collecting in Arizona. 

Dr. Abrams has indicated the characters separating this form 
from the typical P. antirrhinoides, which ranges on the hillsides 
near the coast from San Pedro Martir Mountains northward to 
near San Bernardino, and which is characterized by its very 
broadly ovate sepals and glabrate or puberulent twigs. The 
variety microphyllus, on the other hand, is strictly an interior 
form, ranging from the Providence Mountains eastward into 
western Arizona, and is marked by its distinctly cinereous twigs 
and oblong-ovate or ovate-lanceolate sepals. While the three- 
fold difference residing in the sepals, pubescence and range, argues 
much for specific distinctness of these plants, the relationship is 
so close and so obvious that we feel it best to treat them only 
as well marked geographic forms. 


MuNZ AND Johnston: Plants of California 

Penstemon albomarginatus Jones 

Jones, Contn W 



distinct species. We 

the plant near Lavic, where it 

grew in the deep sand of a wash that ran up between the tongues 


The wash was a stone's 

throw from the Sante Fe Trail and near the grade crossing west of 
Lavic The species is very aptly named, for the white margining 
to the leaves and sepals is very striking. It is apparently rare 
and we can get trace of but three other collections: Good Springs, 
Nevada, and Yucca, Arizona, reported by Professor Jones; and 
Jean Station, Nevada, collected by Mrs. Brandegee (specimen 
in the herbarium of the University of California). Professor 
Jones collected the type in sand and, as our plant grew in similar 
material, the species is probably arenicolous and is to be looked for 
in sandy places. The type and our collection, M, J &f H 4204, 
were both collected in May and were at the time setting seed. 

^ Aplopappus Gooddingi (Nels.) comb, nov, 


Sideranthus Gooddingi Nels. Bot. Gaz. 37: 266. 1904. 

A single plant, M & H j6oj, collected on a gravelly hillside 
about eleven miles west of Needles. The present collection is the 
first one from California, but is remarkably like the typical plants 
from Nevada, It is characterized by its perennial root and tall 
strict stems and large and multiradiate heads, which are up to 
16 mm. wide. A. gracilis the nearest relative of the desert regions, 
is a lower annual with ascending stems, smaller (i cm. or less wide) 
heads and fewer rays, eight to sixteen; its leaves are more setose 
than in A. Gooddingi. 

Pomona College^ 

Claremont, California 



The aim of this Index fs to include all current botanical literature written t>j 
Americans, published in America, or based upon American material ; the word Amer- 
ica being used in the broadest sense. 

Reviews, and papers that relate exclusively to forestry, agriculture, horticulture, 

manufactured products oi* vegetable origin, or laboratory methods are not included, and 

no attempt is made to index the literature of bacteriology. An occasional exception is 

made in favor of some paper appearing in an American periodical which is devoted 

wholly to botany. Reprints are not mentioned unless they diifer from the original in 

some important particular. If users of the Index will call the attention of the editor 

to errors or omissions, their kindness will be appreciated. 

This Index is reprinted monthly on cards, and furni-^hed in this form to subscribers 
at the rate of three cents for each card. Selections of cards are not permitted; each 

subscriber must take all cards published during the terra of his subscription, Corre 
spondence relating to the card issue should be addressed to the Treasurer of the Torrcy 
Botanical Club. 


Knuth, R, Zwei rieue Dioscorea aus Brasilien, Notizbl. Bot. Gart. 
und Mus. Berlin-Dahlem 7: 538, 539. i Mr 1921. 

Korstian, C. F., Hartley, C*, Watts, L. F., & Hahn, G. G. A chlorosis 

of conifers corrected by spraying with ferrous sulphate. Jour. 
Agr. Research 21: 153-171./. 1-4. 2 My 1921. 

Kranzlin, F» Orchidaceae Dusenianae novae. Arkiv Bot. 16^: 1-30 
6 Ap 1920. 

Includes new species from Brazil (23), and Mexico (4), the latter collected by 


Lee, H. A. The increase in resistance to citrus canker with the ad- 
vance in maturity of Citrus trees. Phytopathology 11: 70-73. 
F 1921. 


» " L- 

Lehman, S. G. Soft rot of pepper fruits. Phytopathology 11: 85- 
87. F 1921. 


Lindstrom, E. W- Concerning the inheritance of green and yellow 
pigments in maize seedlings. Genetics 6: 91-110. Ja 1921. 

Livingston, B. E., & Hawkins, L. A. The water-relation between plant 
and soil. Carnegie Instit. Washington Publ. 204: 1-48. /. J-jl 

10 Mr 1915. 


46 Index to American botanical literature 

Lloyd, F. E. Guayule {Parthenium argentatam Gray). A rubber- 
plant of the Chihuahuan desert. Carnegie Instit. Washington 
Publ. 139: 1-213. pL 1-46 +/. i~20, 27 JI igii- 

MacDougal, D. T. A new high temperature record for growth. Sci- 
ence II. 53: 370-372. 15 Ap 1921. 

MacDougal, D. T., & Spalding, E. S. The water-balance of succulent 
plants. Carnegie Instit. Washington Publ. i|i: 1-77. pi- iS +/- 
i-id. 29 D 1910. 

Macpherson, G* E. Comparison of development in dodder and morn- 
ing glory. Bot. Gaz. 71: 392-398. pL 25-27. 16 My 1921, 


Malme, G. O. A.-N. Asclepiadaceae riograndenses adjectis notulis de 
ceteris Asclepiadaceis in Brasilia extratropica, Uruguay et Misiones 
coUectis. Arkiv Bot. 16^^: 1-34. 23 D 1920. 

Includes new species in Roulinia (i), Calostigma (i). 

Mameli de Calvino, E, Estudios anatomicos y fisiologicos sobre la 
cana de azucar en Cuba. Estac. Exper. Agron. Santiago, Cuba, 
Bol. 46: 1-49./. 7-2/. 

Marsh, C. D., & Clawson, A, B. Poisonous properties of the whorled 
milkweeds Asdepias ptimila and A,.verticillata var. Geyen\ U. S. 
Dept. Agr. Bull. 942; 1-14. pi. 1-3 +/. 7, 2. 4 My 1921. 

Mason, T. G. A note on some recent researches on the cotton plant in 
the West Indies, with special reference to St. Vincent. West Indian 
Bull. 18: 184-197.^/. J, 2. [1921.] 


Mason, T. G. The water-balance of the plant and its significance in 
crop production. West Indian Bull. 18: 157-184. [1921.] 

Massey, L. M. Experimental data on losses due to crown-canker ol 
rose. Phytopathology 11: 125-134. Mr 1921. 

Matz, J. Infection and nature of the yellow stripe disease of cane 
(mosaic, mottling, etc.). Jour. Dept. Agr. Porto Rico 3*: 65-82. 


/. 2-/7. O I919. 

Matz, J. Obeservaciones en la gomosis de la cana en Puerto Rico. 
Revista Agr. Puerto Rico 6*: 33-39- 30 Ap 1921. [Illust.] 

[Maxon, W. R., & Eallip, E. P.] Botanical exploration in Jamaica 
[1920]. Smithson. Misc. Coll. 72^; 49-54,/, 61-66, 1921. 

Mayes, J. The practical control of weeds in Missouri meadows and 
pastures. Bull. Missouri Board Agr. 18: 1-36. Je 1920. [Illust,] 

McAtee, W. L. Notes on Viburnum and the assemblage Caprifoli- 
aceae. Bull. Torrey Club 48: 149-154./. j. 25 My 1921. 

McDougall, W. B. Thick-walled root hairs of Gleditsia and related 
genera. Am. Jour. Bot. 8: 171-175./. j-j. 3 Ap 1921. 

Index to American botanical literature 47 

McLeaii) F. T. A study of the structure of the stomata of two species 
of Citrus in relation to citrus canker. Bull. Torrey Club 48: loi- 
106./. I. 4 Ap 1921. 

McLean, F. T., & Lee, H- A. The resistance to citrus canker of Citrus 
nobilis and a suggestion as to the production of resistant varieties in 

other Citrus species. 

/. 7. Mr 1921. 

McNair, J. B- A study of Rhus diversiloba with special reference to its 
toxicity. Am. Jour. Bot. 8: 127-146. pL 2 + f, /, 2. 3 Ap 1921. 

McNair, J. B. The morphology and anatomy of Rhus diversiloba. 
Am. Jour. Bot. 8: 179-191. pL j, 4. 30 Ap 1921. 

McNair, J. B. The transmission of Rhus poison from plant to person. 
Am. Jour. Bot. 8: 238-250. 24 My 1921. 

Menaul, P. Note on the formation of hydrocyanic acid in plants. 

Jour. Biol. Chem. 46: 297. Ap 1921. 

Merriman, M 

Millefolium L. Torreya 

/. 7-5. Ap 1921. 

Morse, W. J. The transference of potato late blight by insects. 
Phytopathology 11: 94-96. F 1921. 

Mousley, H. Further notes on the orchids of Hatley, Stanstead Co., 
Quebec, 1920. Canad. Field Nat. 34: 169-173. 12 Myi92i- 

Muttkowski, R. A. Copper in animals and plants. Science 11. 53: 

453. 454- 13 My 1921, 
Needham, J, G. A biological examination of Lake George, N. Y. 

Sci. Mo. 12: 434-438. My 1921. 

Nelson, J- C. Additions to the flora of western Oregon during 1920, 
Torreya 21: 24-28. Ap 1921. 

Oakley, R. A*, & Evans, M. W. Rooting stems in timothy. Jour. 
Agr. Research 21: 173-178. pL jp, 40. 2 My 1921. 

Orton, C. R- Lightning injury to potato and cabbage. Phytopathol- 
ogy II: 96-98./. 7, F 1 92 1. 

Palmer, E, J. Botanical reconnaissance of southern Illinois. Jour. 
Arnold Arbor. 2: 129-153. Ja 1921. 

Peattie, D. C. An interesting habitat. Rhodora 23; 69-71, 20 Ap 

Pennell, F. W. "Veronica" in North and South America- Rhodora 
23; 1-22, 29-41. 1921. 

Includes Veronica caienata sp. nov.» and several new combinations. 

Pieters, A. J., & Kephart, L. W. Annual white sweet clover and strains 
of the biennial form. U. S. Dept. Agr. Dept. Circ. 167: 1-21. /. 
1-8. Ap 1921. 

48 Index to American botanical literature 

Piper, C. V. Two new legumes from Mexico and Costa Rica. Proc- 
- Biol. Soc, Washington 34: 41, 42. 31 Mr 192 1. 

Pkaseolus chiapasanus and Calopogonium ferrugineum sp. nov. 

Pool, R. J., Weaver, J- E., & Jean, F. C. Further studies in the ecotone 

between prairie and woodland. Univ. Nebraska Stud. 18: 1-47. 
. /. 1-17. 191S. 
Pritchard, F. J., & Porte, W. S. Collar-rot of tomato. Jour. Agr. 

Research 21: 179-184. pL 41-43. 2 My 1921. 

Includes VerHcillium Lycopersici sp. nov. 

Puttemans, A. Gloeosporiiim Bombacis, n. sp. Bull. Soc. Path. Veg. 

7; 74, 75. I S 1920. 

Puttemans, A. Sur VOidhim du Chene au Bresil. Bull. Soc. Path. 
Veg. 7: 37-40. I Je 1920. 

Rathbun, A. E. Methods of direct inoculation with damplng-off fungi, 


Phytopathology 11: 80-84./. 1-3. F 1921. 

Record, S* J. Further notes on intercellular canals in dicotyledonous 
woods. Jour, For. 19: 255-266. Mr 1921. 

Reed, H. S. Growth and sap concentration. Jour. Agr. Research 

21; 81-98./. i-y. I Ap 1921. 

Rehdefj A. Azalea or Loiseleuria. Jour. Arnold Arbor. 2: 156-159. 
Ja 1921. 

Rehder, A. Philadelphus verrucosus Schrader spontaneous in Illinois. 
Jour. Arnold Arbor. 2: 153-156. Ja 1921. 

Ricalton, J. Famous and interesting trees. Am. For. 27: 216-224. 
Ap 1921, [Illust-l 

Richards, H. M. Acidity and gas interchange in Cacti. Carnegie 
Instit. Washington Publ. 209: 1-107./. i~6, 11 Au 1915. 

Riker, A. J. Chondriomes in Chara. Bull. Torrey Club 48; 141-148. 
^ pi. 3. 25 My 1921. 


Rosen, H. R. Furthern observations on a bacterial root and stalk rot 
of field corn. Phytopathology 11: 74-79./- 1-4- F 1921. 

Roth, G. Die aussereuropaischen Laubmoose. Nachtr.i [zu Band i]: 
1-18. pL J, 2. 25 Ap 1912; Xachtr. 2: 1-8. pL i. i N 1913: 
Nachtr. 3: 1-6. pi. i. 18 1915. 

Rowlee, W. W. Tropical trees with light-weight wood- Jour. New' 
York Bot. Card. 22: 75-78. Ap 1921. . 

Rusby, H, H. Guide to the Economic Museum of the New York 
Botanical Garden. Bull. New York Bot. Gard. 11 : 1-318. 30 Ap 
192 1. 

Includes a few new combinations, such as Acyntha Kirkii (Baker) Rusby, page 23- 

Index to American botanical literature 49 

iSaccardo, P. A. Mycetes Boreali-Americani a d. Doct. J, R. Weir 

_ J 

(Spokane, Washington) an. MCMXIX communicati. Nuov. 
Giorn. Bot. Ital. II. 27: 75-88. D 1920. 

Includes 30 new species. 

Sanders, E. M. The natural regions of Mexico. Geog. Rev. ii: 212- 

226. f, 1-5. Ap 1921. 

Includes a vegetation map of Mexico. 

Savelli, R. Anomalie delle plantule e anomalie di germinazione in 
Nicotiana, Nuov. Giorn. Bot. Ital. II. 27: 129-153. D 1920. 

Schaffner, J. H. Additions to the catalog of Ohio vascular plants for 
1920, Ohio Jour. Sci. 21: 128-135. F 1921. 

r ^ h 

Schaffner, J. H. Reversal of the sexual state in certain types of 

monecious inflorescences. Ohio Jour. Sci. 21: 185-198. pL i, 2, 
Ap 192 1. 

Schill, J. F. Crassulacea-Statistiek en geografischeverspreiding (1737-- 
1910). Succulenta 3: 12-15. 9 Mr 1921. 

Schlechter, R. Die Gattung Promenaea Ldl. Notizbl. Bot. Gart, 
und Mus. Berlin-Dahlem 7: 467-481. i Mr 1921, 

Includes 5 new species from Brazil. 


Schlechter, R. Uber einige neue Orchidaceen aus Colombia. Notizbl. 

■ h 

Bot. Gart. und Mus. Berlin-Dahlem 7: 527-532. I Mr 1921. 

New species la Stelis (i), Pleurotkallis (4), Epidendrum (i). 

Schxiltz, E. S,, & Folsom, D. Leafroll, net-necrosis and spindling- 
sprout of the Irish potato. Jour. Agr. Research 21 : 47-80. pL 1-12. 

I Ap 1921. 

Scofield, C. S. Cotton rootrot in the San Antonio rotations. Jour. 
Agr. Research 21: 1 17-125. 2 My 1921. 

Sears, P. B. Vegetation mapping. Science II. 53: 325, 326. 8 Ap 

[Shantz, H- L]. Botanical collecting in Africa [i9i9f 1920]. Smithson. 

Misc. Coll. 72®: 31-38./- 44-^49' 1921- 
Shaw, G. R. The genus Pinus. Publ. Arnold Arbor. 5: 1-96. pL 1-39. 


Shreve, E. B, The daily march of transpiration in a desert perennial. 

Carnegie Instit. Washington Publ. 194: 1-64. pL I +/. 1-22. 13 

Ap 1914. 
Shreve, F.. The vegetation of a desert mountain range as conditioned 


by climatic factors. Carnegie Instit. Washington Publ. 217: 1-112. 

pL i-j6 -f/. 1-18. 28 O 1915. 


50 Index to American botanical literature 

ShuU, G. H. Bursa Biirsa-pastoris and Bjirsa Heegeri: biotypes and 
hybrids. Carnegie Instit. Washington Publ. 112: 1-57. pL 1-4 +/. 
1-23. 1909. 

Skottsberg, C, Botanische Ergebnisse der Schwedischen Expedition 
nach Patagonien und deni Feuerlande 1907-1909. VIIL Marine 
Algae I- Phaeophyceae. Kungl. Svenska Vetenskapakad. Handh 
61^^: 1-56./. 1-20. 1921. 

Includes Gononema and Cladochroa, gen. nov., and new species in Strehlonema (i), 
Myrionema (2), Lithoderma (i), Ralfsia {1), Elachista (2), Mesogloea (i), Scyto- 
ihamnus (i). 

Small, J. K- Old trails and new discoveries. Jour. New York Bot 
Gard. 22: 25-40. pi. 253, 2^4. F 1921; 22: 49-64. pL 255, 256, 
Mr 192 1. 

Collecting in southern Florida- Includes Diospyros Mosieri, sp. nov. 

Smith, E, F., & McKenney, R. E. -B. A dangerous tobacco disease 
appears in the United States. U. S. Dept. Agr. Dept. Circ. 174: 
1-6. Ap 1921. 

Snell, W. H. The relation of the moisture content of wood to its decay, 
with special reference to the spraying of log piles. Pulp and Paper 
Mag. 19: 53I-533-/- ^» 2. 19 My 1921. 

Standley, P. C. Flora of Glacier National Park, Montana. Contrib. 
U. S. Nat. Herb. 22: 235-438 + I- XVII. pi. 33-52, 192 1. 

Standley, P. C- Rubiaceae {pars). N. Am. Fl. 32: 87-158. 10 My 

Includes new species in Pinarophyllon (i), Deppea (i), Bouvardia (12), Ravnia (i), 
Hillia (i), Exostema (5), and Tontanea (i). 

Steil, W. N- The distribution of the archegonia and the antheridia 
on the prothallia of some homosporous leptosporangiate ferns. 
Trans. Am. Micros. Soc. 38: 271-273. O 1919. 

Steil, W. N. Secondary prothallia of Nephrodium hirtipes Hk. Trans. 
Am. Micros. Soc. 38: 229-234, pL 25, 26. Jl 1919. 

Stevenson, J. A. Enfermedades del citro en Puerto Rico. Revista 

Agric, Puerto Rico 4®: 9-19. Je 1920. 

[Corrected reprint.] 

Stockberger, W. W. Ginseng culture. U. S. Dept. Agr. Farmers' Bull. 
1184: 1-16. f. 1-3. Ap 1921. 

Strausbaughi P. D. Dormancy and hardiness in the plum. Bot. 
Gaz. 71: 337-357-/' ^-4- 16 My 1921. 

Strayer, O. B. Rejuvenating pecan trees. Am. For. 27: 238. Ap 
1921. , 

Vol. 49 

No. 3 




MARCH, ^922 

Quadripartition by furrowing In Sisyrlnchium 

Clifford H. Farr 
(with plate 2) 


Since the publication of the author's paper (5) on Magnolia 
there have appeared a few contributions to the subject of the 
quadripartition of pollen- mother-cells by furrowing. In 191 8 
Tackholm and Soderberg (24) presented an interesting di:>covery 
of the occurrence of quadripartition and bipartition respectively 
in two species of Aristolochia, a genus of Dicotyledons; Aristo- 
lochia fimbriata was found to have successive bipartition, just 
as Samuelson (19) had reported in this genus, but Aristolochia 
SipJio was found to have quadripartition of the pollen -mother- 
cells after the homoeotypic mitosis. It is stated that there is 
a slight thickening of the spindle fibers after the heterotypic 
division in the latter species, but no figures of quadripartition 
are shown in this paper. In Vinca rosea the authors conclude 
that quadripartition must occur because of the tetrahedral 
arrangement of the microspores within the mother wall. It 
will be remembered that Frye and Blodgett in 1905 (8) found 
successive bipartition in the sister genus Apocynum^ and the 
microspores here are sometimes arranged tetrahedrally. The 
authors (24) state that after the homoeotypic division in Aris- 
tolochia Sipho walls are laid down as fine plates on the equator 
of the spindle, and that in Albizzia lophanta, a Dicotyledon, 
and Dioscorea qiiinquefolia, a Monocotyledon, quadripartition 

[The Bulletin for February (49: 31-50) was issued April 4, 1922.] 


— ■ 

52 Farr: Quadripartition in Sisyrinchium 

is accomplished by equatorial plates on the spindles. Quadri- 
partition by cell-plates has never been satisfactorially described 
and figured in any of the higher plants in connection with a 
study employing modern methods of microtechnique; it is there- 
fore much to be desired that the authors present these instances 
in detail as soon as possible. 

Florin (7) in 19 1 8 found that quadripartition occurs in the 
spore-formation of one of the acrogj^nous Jungermanniales but 
does not give the details of the process as to whether it is by 
furrowing or cell-plates. In 1919, Miss Digby (3) published 
an extensive chromosomal study of Osmunda in which she refers 
to the formation of cell-plates after the first reduction division. 
These apparently persist through the second mitosis without 
forming completed partitions, and then other cell-plates are 
formed at right angles to these. The figures of these stages 
are drawn from material of 0. pahistris var- aurea although the 
description seems to apply to all four of the types studied, 
including 0. regalis. Smith (20) in 1900 described and figured 
these stages of the latter species- He concluded that the cell- 
plate of the first division was ephemeral and that the division 
was finally accomplished by the simultaneous formation of six 

I * 

cell-plates on the six spindles respectively, thus accomplishing 
quadripartition by cell-plates. During 1920 Yamaha (27) pub- 
lished a study of cell-plate formation in the archesporial and 
spore-mother-cells of Psilotum triqiietrum. He found that the 
nearly complete cell-plate which is formed after the first meiotic 
division breaks up into a granular mass, which persists until 
the close of the second nuclear division. At this time it becomes 
transformed into connecting fibers upon which a cell-plate is laid 
down between the non-sister nuclei, while other cell-plates are 
being formed on the connecting fibers betw'een the sister nuclei 
of the second division. In this way a quadripartition of the 
cell is accomplished by cell-plates. Yamaha concludes that the 
cell-plate has a duplex nature from the first, and that it does not 
split after formation as Timberlake believed. It is to be noted 
that in quadripartition by furrowing the partition is duplex from 
the first. Yamaha suggests that the invagination of the plasma- 
membrane may play some part in the later stages in these cells 
of Psilotum, It therefore seems to me that on account of this 
duplex nature of the partition and in the light of the recent work 
on quadriparition by furrowing that further work on cell-div- 
ision of Psilotum would be warranted. 

Farr: Quadripartitiox in Sisyrinchium 53 

Gu6rin (10) has made a study of the development of the 
anthers and pollen of the labiates, but passes over the matter 
of division of the mother-cell into the four microspores as ex- 
hibiting nothing of particular interest. Valleau (26) in connec- 
tion with his study of sterility in the strawberry has published 
a careful account of pollen-formation. He describes in detail 
the formation of the material about the protoplast which in 
other forms has been considered as the thickening of the cell- 
wall. However, in Fragaria, it seems to be a secretion from the 
protoplast rather than the thickening of the wall. With regard 
to the formation of the partitions the author does not com:nit 
himself. He states: 

The daughter nuclei are soon formed, and walls are laid down between 
them, dividing the cytoplasnl evenly. The cells gradually split apart, 
separating the four microspores and allowing the entrance between them 
of the viscous material. 

It might be concluded from this description that cell-plates 
accomplish the partitioning and that the invagination of the 
peripheral material is simply incidental to the rounding up 
process. No figures of these stages are show^n. Quite recently 
R. R. Gates (9) has published a preliminary account of reduction 
divisions in the pollen-mother -cells of Lacttica sativa. In this 
paper there is described a quadripartition by furrowing rather 
than by cell-plates, just as the writer (4) had previously reported 
for Helianthtis and Ambrosia^ as well as in other groups of 
Dicotyledons. No drawings are given by Gates, but a para- 
graph is devoted to a description of the process. He mentions 
that furrows may be formed either in the presence or the absence 
of spindle fibers but does not describe the latter instance any 
farther. It is stated that an ephemeral cell-plate occasionally 
is found after the heterotypic division, but it never functions 


and no cell-plate is present after the homoeotypic mitosis. 

The most careful recent study of this question is that of 
Mrs. Wanda K. Farr (6), in which she gives figures and des- 
criptions of quadripartition in the poUen-mother-cells of Cobaea 
scandens. These support the writer's contention (4) that 
cell-division by furrowing is common in the pollen-mother-cells 
of Dicotyledons. It is interesting to note that the first descrip- 
tion of cell-division in any plant is probably that by Brongniart 
In 1827 on the pollen-mother-cells of Cobaea scandens (2), in 
which it is indicated that the process is furrowing, though, of 

54 Farr; Quadripartitiox in Sisyrixchium 

course, he had no realization of the internal mechanism of the 
cell. This work was followed by that of Mirbel (17) who 
presented his paper in 1832, though it was not published until 
later. He described and figured quadripartition by furrowing 
in the pollen-mother-cells of Cucurbita Pepo. A little later 
Von Mohl (18) published figures and descriptions of the same 
process in other plants. It now seems that the interpretation 
given by these first observers was more nearly correct than that 
which has been given by most students of cell-division In higher 
plants in more recent years. 

Up to the present no detailed study has been made of quadri- 
partition in Monocotyledons, and it was with the idea of com- 
paring the situation in this group with that found In Dicotyledons 
that the present study was undertaken. Several cases of 
quadripartition In Monocotyledons are known. In 1915, L. 
Guignard published two papers (12, 13) on the occurrence of 
quadripartition in Monocotyledons. In the first paper he 
includes a comprehensive review of the literature on that subject, 
as well as on blpartition in the reduction-divisions of Dicotyledons. 
Quadripartition in Monocotyledons and blpartition in Dicoty- 
ledons -at the time of reduction-divisions are unquestionably 
the exception in the method of pollen formation in these two 
groups respectively. So rare is their occurrence in fact that 
Van Tieghem (25) used them as characters of prime Importance 
in distinguishing the two classes of Angiosperms, thereby 
including the Nympheaceae with the Monocotyledons. Guig- 
nard had previously (11) reported quadripartition in six genera 
of the Orchldaceae, and in the first (12) of his recent publications 
adds four genera (six species) of Liliaceae and six genera (sixteen 
species) of the Irldaceae. To this list the second paper (13) 
contributes three more genera (three species) to the number 
of the Irldaceae having this method. To the Liliaceae may be 
added Strasburger 's (21, p. 151) findings in A sphodelus, and it will 
be remembered that Hofmelster found it In Naias (14, p. 636). 
In 1917, Tackholm and Soderberg published a paper (23) in 
which they discuss tetrad formation in Monocotyledons. In 
addition to its occurrence in the above-mentioned groups they 
refer to quadripartition being reported by Hofmelster in Trades- 
cantia, by Rosenberg in Anther icum and by Von Mohl and others 
In the Juncaceae. So that they find it reported in six families 
of the Monocotyledons, namely: Liliaceae, Juncaceae, Iridacaea, 

Farr; Quadripartition in Sisyrinchium 55 

Commelinaceae, Orchidaceae, and Cyperaceae. The last-named 
is included because of the work of Juel on Carex, of which it 
it may be questioned whether the term quadripartition is 

applicable. In 19 18 the same authors (24) published again, 
adding another species of Monocotyledons to this list, namely, 

Dioscorea quinquefolia. This brings the total number of genera 
up to nineteen or twenty. In Canna they report a tetrahadral 
arrangement of the microspores but do not take this as evidence 
of quadripartition. 

Guignard (12) states that in four species of Iris there is 
occasionally a suggestion of a partition after the heterotypic 
nuclear division but this is always ephemeral. After the 
homoeotypic mitosis in Sisyrinchium, Antholyza, Freesia, Ixia, 
and Monbretia, when the fibers of the central spindle are formed 
between the four nuclei, it is stated that rather frequently there 
is noticeable a faint thickening on the internal face of the mother 
wall at the places which are to become the points of insertion 
of the partitions. Further than this no details of quadripartition 
are given, nor are drawings or photomicrographs shown to give 
evidence as to whether the process is accomplished by cell-plates 
or by constriction furrows. It is not unlikely that the ephemeral 
equatorial differentiations which Guignard found in these 
Iridaceae, and which Tackholm and Soderbeg reported for 
Aristolochia Sipho are orange zones, such as the writer (5) has 
shown to occur in Magnolia, Tackholm and Soderberg (24) 
do not describe the process of quadripartition in Dioscorea 
except to state that it is accomplished by equatorial plates 
being formed on the spindles. 

It thus appears desirable that a study be made of the details 
of the process of quadripartition in a Monocotyledon. The 
writer has discussed at some length the literature on this subject 
in his first paper in 1916 (4). At that time he presented a 
description and drawings of quadripartition in Nicotiana and 
other Dicotyledons which led to the conclusion that no cell-plates 
are formed, but that division is accomplished by furrows very 
much as in animal cells. Mrs. Wanda K. Farr (6) has more 
recently shown that the same type of cytokinesis occurs in 
Cobaea scandens as was found in Nicotiana. In 1918, the writer 
(5) supplemented the work of Guignard on Magnolia, in which 
the latter showed incipient furrows during interkinesis, but did 
not present the stages following the origin of the tetranuclear 


56 Farr: Quadripartition in Sisyrinchium 

condition. It was found that after the disappearance of the 
ephemeral orange zone a furrow developed on the margin of the 
equator of the heterotypic spindle. However, the development 
of this furrow is arrested when it is only about one-fourth 
completed and the homoeotypic nuclear division ensues. The 
division of the cell is finally accomplished by the completion 
of this furrow and the formation of two other furrows at approxi- 
mately right angles to it. Although Nymphaea has not as yet 
been thoroughly investigated the work of Lubimenko and Maige 
(16) indicates that it is similar in this respect to Magnolia. 

The following study was made upon material of Sisyrinchium 
Biishii Bicknell, collected near College Station, Texas, and 
identified by Dr. A. S, Hitchcock. This species proved especi- 
ally fine for study, both because of the excellent fixation and 
because of the distribution of the mitotic figures in the anthers. 
In Nicotiana the mother-cells of a given anther are all in very 
nearly the same stage of division, and hence to secure an un- 
broken series of stages it is necessary to take material from several 
different anthers. In Magnolia the mother-cells within the 
anther were at different stages but there w^as no special arrange- 
ment of them. In either of these cases the objection might 
be raised that a cell-plate was overlooked due to the study of 
an incomplete series of stages. It was therefore highly desirable 
that a form be found in which the stages were arranged In order 
within the anther from one end to the other, so that by studying 
a number of anthers it could be determined with certainty that no 
steps have been omitted. Such a situation is that which Sisy- 
rinchiicm Bushii presents. The mother-cells at one end of the 
anther may be in metaphase of the heterotypic division and 
those at the other end in metaphase of the homoeotypic. This 
is the condition shown in Fig. i.. In Fig. 2, the cells at the Inner 
end are in metaphase of the homoeotypic division and those at 
the outer end are in the tetranuclear stage. It is evident that 
the study of a number of such anthers is sure to reveal every 
stage, however transitory. It will be remembered that a similar 
condition exists in the testes of Batracoceps, and among plants 
it was found by Strasburger (22) in the anthers of Fritillaria and 
by D'Angremond (1) in the banana. 

The pollen-mother-cells of Sisyrinchium are smaller than 
those characteristic of many Monocotyledons. Miss Kliene- 
berger (15) has recently published a study of the size of nuclei 

Farr: Quardtpartition in Sisvrinchium 57 

of Monocotyledons, in which she finds that contrary to the 
general opinion the nuclei of most Monocotyledons are small. 
It is only in the Iridaceae and parts of the Liliaceae, Amaryl- 
lidaceae and Convallariaceae that large nuclei are found. How- 
ever, her study is not concerned with pollen-mother-cells 
especially. It would be interesting to find how great the 
differences in size of pollen-mother-cells may be among Mono- 
cotyledons, and their relation to the size of the nuclei. The 
tetranucleate pollen-mother-cells of Sisyrinchium just before 
division are 36-40 microns in diameter, which is just about the 
size of those of Nicotiana. 

The mother-cell wall is likewise thickened to about the same 
extent as is that of Nicotiana, that is, on the average about one- 
tenth of the diameter of the cell. The process of thickening 
of the cell-wall begins about the time of synapsis, but the greatest 
development is during interkinesis. The cells are more compact 
within the anther than they are in either Nicotiana or Magnolia^ 
but nevertheless are quite loosely disposed during diakinesis 
and later stages of reduction. They are somewhat closer 
together toward the end of the process than in the earlier stages 
doubtless due to the enlargement of the cells and the thickening 
of the walls. There seems to be no evidence that the gelatinous 
substance enveloping the protoplast during the later stages of 
reduction-divisions is a secretion from the protoplast, as Valleau 
(26) suggests, rather than a product of the cell-wall. The 
middle lamellae remain quite distinct throughout these stages 
and careful observation of the material both of Sisyrinchium 
and of the other forms previously studied leads the writer to 
the conclusion that it is the secondary lamellae of these pollen- 
mother-cells which during reduction-divisions take on the 
colloidal property of imbibition and swell to many times their 
original thickness. 

The heterotypic division results in two discoid nuclei being 
organized at opposite ends of a rather long central spindle. No 
indication of a cell-plate or other equatorial differentiation is 
to be discerned at any time during interkinesis. Nor is there 
any indication of furrowing. The nuclei gradually become more 
nearly spherical (Fig, i) and finally, before the second nuclear 
division begins, they are almost perfect spheres. They appar- 
ently do not come into contact with the plasma membrane. The 
fibers of the central spindle disappear almost entirely during in- 

58 Farr: Quadripartition in Sisyrixchium 

terkinesis, and the integrity of the spindle as such becomes en- 
tirely lost. The fact that all stages of interkinesis may be found 
within the extent of a eingle anther (Fig. i) makes it possible 
to arrive at an index of the relative time required for that stage. 
Judging from the extent of the various stages within the anther 
it would seem that the time involved in interkinesib is about 
equal to that involved in either the first or the second reduction- 

In the metaphase of the homoeotypic nuclear division the 
spindles are in some cases parallel and in some at right angles 
to each other, and in still others at positions intermediate 
between these two extremes. When the nuclei are reconstituted 
they may occasionally be found in a single plane as is bhown in 
the outer end of the anther in Fig. 2. But more frequently the 
nuclei are tetrahedrally arranged (Fig. 3). Quite a number of 
cellii show intermediate orientations. It is interesting to note 
that the type of division of the cell is the same, regardless of 


the orientation of the nuclei, just as was previously reported 
for Magnolia (5). 

The nuclei become reorganized in much the same manner 
as after the heterotypic division. They, however, do not become 
as large as in the previous division and they move apart so that 
they become closely appressed to the plasma membrane. Al- 
most as soon as the nuclear membranes appear spindles are 
organized connecting the nuclei which are not already connected 
by the two homoeotypic spindles. There are thus six spindles 
in all within the cell, but the integrity of some of these seems 

almost lost especially in cases where the four nuclei are in one 

The time elasping between the conclusion of nuclear division 
and the beginning of cytokinesis is evidently relatively long. 
This is indicated by the fact that all of the cells of a single anther 
may be in these stages, those at one end just coming out of 
karyokinesis and those at the other end not yet showing signs 
of the formation of partitions. It thus appears that this period 
is at least as long as that involved in either the heterotypic or 
homoeotypic mitosis; which is quite in harmony with the writer's 
previous suggestion (5) that cell-plate formation and furrowing 
do not occur at corresponding periods in the cell-cycle, but that 
cell-plate formation follows very soon after nuclear division, if 
indeed the two processes may not frequently overlap. Furrow- 

Farr: Quadripartitiox in Sisyrinchium 59 

ing on the other hand takes place only after a considerable time 
has elasped following the re-formation of the nuclei. If cell- 
plates appeared in thib division we would most certainly expect 
to find them in such an anther as that shown in Fig. 2, but none 
such are present. 

The process of cytokinesis in the quadripartition of the 
pollen-mother-cells of Sisyrinchium is by furrowing and is 
practically identical with that found in Nicotiana. It does not 
seem necessary to repeat the details of the description of the 
process here, inasmuch as there is no difference from that which 
has been published before (4). In cells where the nuclei are 
tetrahedrally arranged the plasma membrane invaginates most 
rapidly at the points equidistant from the three nuclei and 
therefore a central triangular area is formed (Fig. 3). In cells 
in which the nuclei are in one plane no such central triangular 
area appears, but the spindles have the Same relation to each 
other, as was shown in those cells of Magnolia (5) in which the 


arrangement was of this type. 

Literature cited 

1. D'Angremond, A. Parthenokarpie und Samenbildung be 

Bananen, Flora 107: 57-110, pL 4-11. 1915. 

2. Brongniart, Adolph. M^moire sur la generation et le de- 

veloppement de Fembryon de la v^getaux phan^rogames. 
Ann. Sci. Nat. 12: 14-53,145-17^,225-296. pL j4-^4. 

3. Digby, L. On the archesporial and meiotic mitoses of 

Osmunda. Ann. Bot. 33: 135-172. pi. 8-12. 1919. 

4. Farr, C. H. Cytokinesis of the pollen-mother-cells of certain 

• Dicotyledons. Mem. New York Bot. Garden 6: 253- 
317. pi. 27-2g. 19 16, 

5. Cell division by furrowing in Magnolia. Am* Jour* 

Bot. 5: 379-395- M 30-32- 1918- 

6. Farr, Wanda K. Cell division of the pollen-mother-cells of 

Cohaea scandens alba. Bull. Torrey Club 47: 325-338. 

pL 14. 1920. 

7. Florin, R. Cytologische Bryophytenstudien I. UberSpor- 

enbildung bei Chiloscyphtis polyanthus (L.) Corda. 
Ark. for Bot, 15i«: i-io. pL i + f. J, 2. 1918. 

8. Frye, T. C, & Blodgett, E. B. Contribution to the life- 

history of Apocynum androsaemifolium. Bot. Gaz. 40 r 

49-53- pl' 2. 1905. 

6o Farr: Quadripartition in* Sisyrixchium 

9. Gates, R. R- A preliminary account of the meiotic phe- 

nomena in pollen mother-cells and tapetum of lettuce 
{Lactuca sativa). Proc. Roy. Soc. London, B. 91: 216- 
. /. I, 2. 1920. 

10. Guerin, Paul, Developpement de I'anthcre et du pollen 


des Labiees. Compt. Rend. Acad. Sci. Paris 168: 
182-185, 1919. 

1 1 . Guignard, L- Recherches sur le developpement de 1 'an there 

et du pollen des Orchidees. Ann. Sci. Nat. Bot. VL 
14: 26-45. pL 2. 1882. 

12. Sur la formation du pollen. Compt. Rend. Acad. 

Sci. Paris. 160: 428-433. 19 15. 

13. Nouvelles observations sur la formation du pollen 

chez certaines Monocotylddones. Compt. Rend- Acad. 
Sci. Paris. 161: 623-625. 1915. 

14. Hofmeister, W. Neue Beitrage zur Kenntniss der Embrj'o- 

bllding der Phanerogamen. 11. Monocotyledonen. Ab- 
handl. Konigl. Sachs. Ges. Wiss. 7: 629-760. pL 1-2$* 

1 861. 

Emmy. Ueber die Grosse und Beschaffenheit 

der Zellkerne mit besonderer Berucksichtigung der 
Systematik. Beih. Bot. Centralb. 35^: 219-278, pi i. 

16. Lubimenko, W., & Maige, A. Recherches cytologiques sur 

le developpement des cellules-meres du pollen chez les 
Nympheacees. Rev. G6n. Bot. 19: 401-425, 433-458, 

474-505. 1907. 

17. Mirbel, M. Complement des observations sur le Mar- 

chantia polymor pJm , suivi de recherches sur les meta- 
morphoses des utricules, et sur rorigine, les develqppe- 
ments et la structure de I'anthere et du pollen des 
veg^taux phan^rogames. M^m. Acad. Roy. Sci. 13: 
375-436. pi. i-io. 1835. 

18. Mohl, Hugo von. Ueber die Vermehrung der Pflanzen- 

zellen durch Theilung. Tubingen. 1835. 

19. Samuelson, G. Ueber die PoUenentwickelung von Anona 

und Aristolochia und ihre systematische Bedeutung. 
Svensk. Bot. Tidsk. 8: 181-189. 1914. 

20. Smith, W. R. The achromatic spindle in the spore mother 

cells of Osmunda regalis. Bot. Gaz. 30: 361-377. pi. 22. 

21. Strasburger, E. Zellbildung und Zelltheilung. Ed. 3. Jena. 


Farr: Quadripartition in Sisyrinchium 6i 

22. Das botanische Praktikum. Jena. 1884. 

23. Tackholm G., & Soderberg, E. Ueber die PoIIenent- 

wickelung bei Cinnomomum nebst Erorterungen iiber 
die phylogenetische Bedeutung des Pollentypus. Ark. 
for Bot, 15^: 1-12. 1917. 

24. Neue Beispiele der simultanen und sukzesslven 

Wandbildung in den PoIIen-mutterzellen- Svensk. Bot- 
Tidskr. 12: 189-201. /, i-p. 1918. 

25. Tieghem, Ph. Van. L 'oeuf du plantes considere comme base 

de la classification. Ann. Sci. Nat, Bot, VI. 14: 286- 
464. 1 90 1. 

26. Valleau, W. D. Sterility in the strawberry. Jour. Agr. 

Research 12: 613-6.70 pL B~E + 35, 36. 1918. 

27. Yamaha, Gihei. Einige Beobachtungen iiber die Zellteilung 

in der Archesporen und Sporenmutterzellen von Psilotum 
triquetrum Sw., mit besonderer Riicksicht auf die Zell- 
plattenbildung. Bot. Mag. Tokyo 34: 1 17-129. /. 1-20. 


Description of plate 2 

Fig. I. Photomicrograph of a longitudinal section of an anther of 
Sisyrinchium Bushii, The cells in the outer end are in the metaphase of 

the heterotypic and those at the inner end are in metaphase of the homoeo- 
typic division. 

Fig. 2. Photomicrograph of an anther at a slightly later stage of develop- 
ment. The cells at the inner end are in metaphase of the homoeotypic 
mitosis and those at the outer end are in the tetranucleate stage just prior 
to furrow formation. 

FiG, 3. A drawing of a cell in cytokinesis. The furrows are partly 
formed and the central triangular area is appearing. 

Bull. Torrey Club 

Volume 49, plate 2 

F19 2 

F.g \ 



A modern plant fossil 

Ed A M. Round 
(with oxe text figure) 

While collecting botanical materials in East Killingly^ 
Connecticut, the writer came upon a so-called "cedar swamp '* 
in which ferns like Woodwardia angustifolia Smith and Aspidium 
simulatum Davenp. grew, together with sphagnum and several 
species of iiverw^ort. Woodsmen had visited the locality recently 
and cut the large hemlock trees, from the stumps of which 
broad shelf fungi belonging to the Polyporaceae had developed. 
One of these fungi had been removed from its host by a previous 
visitor and thrown upon a rock, over which flowed the waters 
of the peat bog. Examination of this water-soaked 
showed it to be tough and pliable like India rubber, due perhaps 
to the resinous nature of the hemlock tree on which the fungus 
grew and the preservative qualities of the water from the peat 
bog. The specimen was in such good condition that the writer 
contemplated drying it for further study, when a cluster of the 
same fungi was found in situ upon a large hemlock stump, 
somewhat overgrown by small black birch trees. Examination 
of one of these specimens revealed not only the ordinary lines 
upon its surface but also several very clear tracings of black 
birch leaves, the details of which were as exact as obtain on 
many rocks with fossil leaves impressed thereon (Fig. i). 

In searching for an explanation of this phenomenon, it 
appeared that the black birch leaves had touched the surface 
of the shelf fungus when it was damp and growing rapidly, so 
that the leaves had been completely overgrown by the fungoid 
strands while still attached to the parent plant. Over the whole 
surface could be found examples of leaves in process of being ab- 
sorbed or incorporated into the fungus, parts of them being 
encrusted while other parts of the same leaves were free, dry 
and easily broken. Still other cases showed leaves scarcely 
touching the fungus but evidently somewhat invaded by its 
hyphae, as they were beginning to brown and seemed to be dying. 

Many ideas are advanced to account for the formation 



. Round: A Modern Plant Fossil 

of fossils.* Given, however, materials of as tough a nature 
as the shelf fungus from the hemlock stump, on which a leaf 
had been encrusted or replaced by hyphae of the same tough- 

FiG. I, Black birch leaves "fossilized'' on a shelf fungus, X H- Original 
specimen now in the Paleontological Laboratory, Brown University. 

ness, one has but to postulate the submergence of the fungus 



It appears, therefore, that the fungus above described with 
its encrusted black birch leaves illustrates a method whereby 
fossils may have formed in the past or may develop under mod- 
conditions. It may therefore be popularly designated as 
exemplifying a "modern plant fossil." 


Pirsson & Schuchert, Text book of geology, p. 435, 1915. 
t See Shinier, An introduction to the study of fossils, p. 3, 1918 



The aim of this Index is to include all current botanical literature written by 
Americans, published in America, or based upon American material; the word 
America being used in the broadest sense. 

Reviews, and papers that relate exclusively to foresty, agriculture, horti- 
culture, manufactured products of vegetable origin, or laboratory methods 
are not included, and no attempt is made to index the literature of bacteriology. 
An occasional exception is made in favor of some paper appearing in an Ameri- 
can periodical which is devoted wholly to botany. Reprints are not mentioned 
unless they differ from the original in some Important particular. If users 
of the Index will call the attention of the editor to errors or omissions, their 
kindness will be appreciated. 


This Index Is reprinted monthly on cards, and furnished in this form to 
subscribers at the rate of three cents for each card. Selections of cards are 
not permitted; each subscriber must take all cards published during the term 
of his subscription. Correspondence relating to the card issue should be 
addressed to the Treasurer of the Torrey Botanical Club. 

Adams, J. F. Rusts on conifers in Pennsylvania. Pennsylvania 

Agn Exp. Sta. Bui. i6o: 3-30. f, i-io. D 1919. 

Adams, J. F. Sexual fusions and development of the sexual 

organs in the peridermiums. Pennsylvania Agr. Exp. Sta. 
Bull. 160: 31-76. pL 1-5+ f. 1-8. D 1919. 

Albertson, A- O. Nantucket wild flowers. Pp. i-xlv+ 1-442. 

1921. [Illust.] 

Arny, A. C. Further experiments in field technic in plot tests. 



Ashe, W. W- A new shrubby buckeye. Jour. Washington 

Acad. Sci. 3: 424. 19 S 1913. 

Aesctihis microcarpay sp. nov., from Georgia. 

Barnes, C. R. Physiology, In Coulter, J. M., Barnes, C. R., & 

Cowles, H. C, A textbook of botany 295-484./. 6ig-6gg. 

Bamhart, J. H. Otto Kuntze. Bull. Charleston Mus. 9: 65- 

68. D 1913. 

Berry, E. W. An Eocene flora from Trans-Pecos, Texas. U. S. 

Geol. Surv. Professional Paper 125-A: 1-9. pL 1-3 -i- f. 

ly 2. 12 S I9I9. 

Includes new species in Geonomites (i) and Ilex (i). 


66 Index to American Botanical Literature 

Berry, E. W. The teaching of paleobotany. Bull. Geol. See, 

Amer. 31: 389-394. 30 S 1920. 

Blake, S. F. The American species of Maximilianea (Cochlo- 

spermum). Jour. Washington Acad. Sci, 11: 125-132,/. i, 
2, 19 Mr. 1921. * 

Includes I new species from Venezuela. 

Blake, S. F. The anay, a new edible-fruited relative of the 

avocado. Jour. Washington Acad. Sci. 9: 457-462. 4 O 

Includes 2 new species of Hufelandia from Central America. 

Blake, S. F. A preliminary revision of the North American 


Jour. Washingt 

/■ I, 2, 4 J 

Includes 2 new species. 

Blake, S, F. Revision of Ichthyomethta, a genus of plants used 

for poisoning fish. Jour. Washington Acad. Sci. 9: 241- 

. 4 My 1919. 

Includes 2 new species from tropical America. 

Blake, S. F. Revision of the true mahoganies (Swietenia). 

Jour. Washington Acad. Sci. 10: 286-297. /. j, 2. 19 My 

Includes i new species from Mexico. 

Briggs, L. J. The living plant as a physical system. Jour. 

Washington Acad, Sci. 7: 89-1 11. /. 1-6. 19 F 1917. 

Briggs, L. J., & Shantz, H. L. The wilting coefficient for 

different plants and its indirect determination. Jour. 
Washington Acad. Sci. 1: 228-232, 19 N 1911. 

Britton, E. G. Bahama mosses. Bryologist 24: 17-19- pL /• 

25 Jl 1921. 

Britton, E. G., & Holzinger, J. M. The rediscovery of Physcomi- 

trium pygmaeum James. Bryologist 24: 26, 27. 25 Jl 

Britton, N. L. Further botanical studies in Trinidad. Jour. 

New York Bot. Gard. 22: 93-102. My 192 1. 

Brown, B. Cretaceous Eocene correlation in New Mexico, 

Wyoming, Montana, Alberta. Bull. Geol. Soc. Amer. 25:- 
355-380. 15 S 1914. 

Buchholz J. T. Studies concerning the evolutionary status of 

polycotyledony. Am. Jour. Bot. 6: 106-119. /. 1-23. Mr 

Index to American Botanical Literature 67 

Carey, C. L, On the gross structure of an agar gel. Bull. 

Torrey Club 48: 173-182. /. 1-4. i Au 1921. 

Clements, F, E- Scope and significance of paleo-ecology. Bull. 

Geol. Soc. Amer. 29: 369-374. 30 Je 1918. 

Cobb, F,, & Bartlett, H. H. On Mendelian inheritance in 

crosses between mass-mutating and non-mass-mutating 
strains of Oenothera pratincola. Jour. Washington Acad. 
Sci. 9: 462-483. 4 O 1919. 

Collins, G. N. Intolerance of maize to self-fertilization. Jour. 

Washington Acad. Sci. 9: 309-312. 4 Je 1919. 

Collins, G. N. New place effect in maize. Jour. Agr. Research 

12: 231-243. 4 F 1918. 

Collins, G. N. The origin of maize. Jour. Washington Acad. 

Sci. 2: 520-530. 19 D 1912. 

Collins, G. N., &Kenipton J. K. A hybrid between Tripsacum 

and Etichlaena. Jour. Washington Acad. Sci. 4: 114-117. 
4 Mr 1914. 

Collins, G. N., & Kempton, J. H. A teosinte-maize hybrid. 

Jour. Agr. Research 19: 1-37. pi. 1-7 + J^ 1-33^ i Ap 1920. 

Cook, O. F. Agriculture and native vegetation in Peru. Jour. 

Washington Acad. Sci. 6: 284-293. f. /, 2, 19 My 1916. 

Cook, O. F. Evolution through normal diversity. Jour. Wash- 
ington Acad. Sci. 9: 192-197. 4 Ap 1919. 

Cook, O. F. Glaticothea, a new genus of palms from Lower 

California. Jour. Washington Acad. Sci. 5: 236-241. 4 
Ap 1915. 

Cook, O. F. Ivory palms in Panama. Jour. Washington Acad. 

Sci. 3: 138-143. 4 Mr 1913. 

Includes 4 new species of Phytelephas. 

Cook, O. F, Jointed leaves of Amygdalaceae. Jour. Washing- 
ton Acad. Sci. 2: 218-220/ 4 My 1912. 

Cook, O. F. The Mascarene cabbage palm as a new genus. 

Jour. Washington Ac 

Linoma Cook, gen. nov. 

127- 4 Mr 

Cook, O. F. Morphology and evolution of leaves. J 

Washington Acad. Sci. 6: 537-547- ^9 S 1916. 

Cook, O. F. A new generic name for the sapote. J 

Washington Acad. Sci. 3: 158-160. 19 Mr 1913. 

Achradelphia Cook., nom. nov. 

68 Index to American Botanical Liter aure 

Cook, O. F. A new genus of palms allied to Archonto phoenix. 
Jour. Washington Acad. Sci. 5: 116-122. 19 F 1915. 

Loroma Cook, from Australia. 

Cook, O. F. Pomegranate flowers dimorphic. Jour. Washing- 
. ton Acad. Sci, 2: 434-437. /. /, 2, 4 N 19 12. 

Cook, O. F. Quichua names of sweet potatoes. Jour. Wash- 
ington Acad. Sci. 6: 86-90, 19 F 1916. 

Cook, O. F, Seedling morphology in palms and grasses. Jour. 
Washington Acad. Sci. 7: 420-425. 19 Jl 191 7- 

Cook, O. F. Tribroma, Sinew genus of tropical trees related to 

Theobroma. Jour. Washington Acad. Sci. 5: 28 
Ap 1915. 

Based on Theobroma hicolor H, & B, of tropical America. 


Cook, O. F., & Cook, R. C. The maho, or mahogua, as a trans- 

Pacific plant, J 
IQ Mr IQ18. 

8: 153-170. 

Cook, O. F,, & Cook, R. C. Polynesian names of sweet potatoes. 

Jour. Washington Acad. Sci. 6: 339-347. 
ter, J. M. Morphology, in Coulter, J. M 




J. M 

A textbook of 

botany (for colleges and universities). Pp. 1-964 +A-L 
+ a-q. /. 1-1234. 1910-1911. 

Coville, F. V. The formation of leaf mold. Jour. Washington 

Acad. Sci. 3: 77-89. 4 F 1913. 

les, H. C. Ecology, in Coulter 

Cowles, H. C, A textbook of 
1234. 191 1. 

J. M 

/. 700- 

Cunningham, C. C. Study of the relation of the length of 
kernel to the yield of corn {Zea Mays indentata). Jour. 
Agr. Research 21: 427-438. pi. 80-87. i Jl 1921. 

Dufrenoy, J. The biological significance of false witches '- 

brooms in ericaceous plants. Jour. Washington Acad. Sci. 
8-' 527-532. 19 S 1918. 

Fairchild, D. A hunter of plants. Nat. Geog. Mag. 36: 57-77- 

Jl 1919. [lUust.] 

Account of the work of Frank N. Meyer. 

Index to American Botanical Literature 6^ 

Femald, M. L. The American representatives of Arenaria 

sajanensis. Rhodora 21: 12-17. 10 F 19 19. 

Includes Arenaria mar cescensy sp. nov., from Newfoundland and Quebec. 

Femald, M. L. American variations of Arenaria verna. 


Rhodora 21: 21, 22. 10 F 1919. 

Fernald, M. L. The earlier names for Alsinopsis. Rhodora 

21: 9-12. 10 F 1919. 

Femald, M. L. The specific identity of Arenaria groenlandica 

and A, glabra. Rhodora 21: 17-21. 10 F 1919. 

Femald, M, L. The type of the genus Alsine. Rhodora 21: 

7-9. 10 F 1919. 

Femald, M. L. The unity of the genus Arenaria. Rhodora 

21: 1-7. 10 F 1919. 

Includes many new combinations. 

Goodspeed, T. H., McGee, J. M., & Hodgson, R. W. Note on 

the effects of illuminating gas and its constituents in causing 
abscission of flowers in Nicotiana and Citrus. Univ. 
CaHfornia Publ. Bot. 5: 439-450. 28 D 1918, [Corrected 

Griffiths, D., & Juenemann, H, E, Commercial Dutch-bulb 

culture in the United States. U. S. Dept. Agr. Bull. 797: 
1-50. /. 1-32. 22 N 1919. 

Harper, R. M. The supposed southern limit of the eastern 

hemlock. Torreya 19: 198, 199. O 1919. 

Harper, R. M. Water and mineral content of 

fern. Am. Fern Jour. 9: 99-103. 24 Ja 1920, 

Polypodium polypodioides (L.) Hitchcock. 

Herre, A. W. C. T. Supplement to the lichen flora of the 

Santa Cruz Peninsula, California. Jour. Washington Acad. 
Sci, 2: 380-386. 19 S 1912. 

Includes Thelocarpon albomarginatum, sp. nov. 

Hitchcock, A. S. History of the Mexican grass, Ixophorus 

an epiphytic 

uniselus. Jour. Wa 

4 N 

Hodges, R. S. Ringworm of the nails. Arch. Dermat. & 

Syphil. 4: 1-26. /. 1-12. Jl 192 1. 

Caused by species of Trichophyton. 

Hollick, A. Qtiercus heterophylla in the Clove Valley [New 

York]. Proc. Staten Id. Assoc. Arts & Sci. 7: 32-34- P^- 
4. 1918. 

yo Index to American Botanical Literature 

Hopkins, A. D. The bioclimatic law. Jour. Washington Acad. 

Sci. 10: 34-40 19 Ja 1920. 

Howe, M. A. A little-known mangrove of Panama. Jour. 

New York Bot. Gard. 12: 61-72. /. i6-2j. Ap 1911; 

Jamieson, C. O., & WoUenweber, H, W. An external dry rot 

of potato tubers caused by Fusariiim trichothecioides , 
Wollenb. Jour. Washington Acad. Sci. 2: 146-152. /. J. 
19 Mr 1912. ' 

Jennings, O. E. New or noteworthy plants from northwestern 

Ontario. I. Jour. Washington Acad. Sci. 10: 453-460. 4 


Includes new species and varieties in Lysias (i), Kneiffia (i), Pyrola 
(3), Scutellaria (i), Stachys (3). 

Kearney, T. H. Plant life on saline soils. Jour. Washington 
Acad. Sci. 8: 109-125. 4 Mr 1918. 


Kellerman, K. F. The effect of salts of boron upon the distri- 
bution of desert vegetation. Jour. Washington Acad. Sci- 
10: 481-486. 19 O 1920. 

Kellerman, M, Phototypes, a means for wide distribution of 

type material. Jour. Washington Acad. Sci. 2: 346-348. 
19 Au 1912. 

Kempton, J. H. The ancestry of maize. Jour. Washington 

Acad. Sci. 9: 3-1 1. 4 Ja 1919. 

Kempton, J. H. A correlation between endosperm color and 

albinism in maize. Jour. Washington Acad. Sci. 7: 146- 
149. 19 Mr 1917. 

Kempton, J. H. Linkage between brachytic culms and peri- 
carp and cob color in maize. Jour. Washington Acad. Sci. 
11: 13-20. 4 Ja 1921. 

Knowlton, F. H. A catalogue of the Mesozolc and Cenozoic 

plants of North America. U. S. Geol. Surv. Bull. 696: i- 
815. 1919. 

Knowlton, F. H. Cretaceous-Tertiary boundary in the Rocky 

Mountain region. Bull. Geol. Soc. Amer. 25: 325-340. 
15 S 1914. 

Knowlton, F. H. Description of a supposed new fossil species 

of maize from Peru. Jour. Washington Acad. Sci. 9: 134- 
136. 4 Mr 191Q. 

Index to American Botanical Literature 71 


Knowlton, F. H, Relations between the Mesozoic floras of 

North and South America. Bull. Geol. Soc. Amer. 29: 
607-614. 30 D 1918. 

Lamb, W- H. Moreh oak, a new name for Quercus morehus 

Kellogg. Jour. Washington Acad. Sci. 6: 657, 658. 19 N 

Lewton, F. L. The Australian fugosias. Jour. Washington 
■ Acad. Sci. 5: 303-309. 4 My 1915. 

Lewton, F. L. The history of kidney cotton. Jour. Wash- 
ington Acad. Sci. 10: 591-597. /. /, 2. 19 D 1920. 

[Lumsden, D.] Hand-list of trees and shrubs native and exotic, 
including the Coniferae, grown in the arboretum [of Walter 
Reed General Hospital, Washington, D, C.]. pp. 1-66. 
I Mr 1921. 

Maxon, W. R. A new Anemia from Mexico. Jour. Washington 

Acad. Sci. 8: 199, 200. 4 Ap 1918. 

Anemia Makrinii Maxon. 


McCulloch, H. L. A bacterial disease of Gladiolus, Science 

H. 54: 115, 116. 5 Au 1921. 

Bacterium m^arginatum, sp. nov. 

McDougall, W. B. The classification of symbiotic phenomena. 

Plant World 21: 250-256. O 1918. 

McLaren, D. California's flora. Parks & Recreation 4: 275- 

279. Jl 1921. [Ulust.] 

McMurran, S. M- Walnut blight in the eastern United States. 

U. S. Dept. Agr. Bull. 611: 1-7. pL J, 2, 10 D 1917. 

Metcalf, F. P. Notes on North Dakota plants. Jour. Wash- 
ington Acad. Sci. 10: 188-198. 4 Ap 1920. 

MoxJey, G. L. Some vacation lichens, Bryologist 24: 24, 25. 

25 Jl 1921. 

Nichols, G. E. The vegetation of Connecticut. VI. The 

plant associations of eroding areas along the seacoast. 
Bull. Torrey Club 47: 89-117. /. /-^. 31 Mr 1920. 


Norton, J. B, The eastern and the western migrations of 

Smilax into North America. Jour. Washington Acad, 
Sci. 6: 281-283. /• ^' ^9 My 1916. 

Norton, J, B. Spring flowers in the fall. Jour. Washington 

Acad Sci. 5: 285-287. /. i. 19 Ap 1915. 

72 IxDEX TO American Botanical Literature 


Parish, S. B, A supplementary bibliography of the Southern 

California flora. Bull. S. Cahfornia Acad. Sci. 19: 24-29, 
Ja 1920. 

Pearson, W. H, Notes on a small collection of hepatics from 

Oregon. Bryologist 24: 21. 25 Jl 192 1. 

Piper, C. V. A new genus of Leguminosae. Jour. Washington 

Acad. Sci. 10: 432,433. 19 S 1920. 

Monoplegma, gen, nov., from Costa Rica. 

Pittier, H. Inophloeum, a new genus of the mulberry family. 
Jour. Washington Acad. Sci. 6: 112-114. 4 Mr 1916. 

From Panama and adjacent Colombia. 

Pittier, H. Notes on the genus Swartzia in Panama and 

Guatemala. Jour. Washington Acad. Sci. 11: 155-160- 
4 Ap 1921, 

Includes 2 new species. 

Pittier, H. On the origin of chicle with descriptions of two 

new species of Achras. Jour, Washington Acad. Sci. 9: 
431-438. 19 S 1919. 

New species from Guatemala and Panama. 

Pittier, H. Two new species of Bursera. Jour. Washington 

Acad. Sci. 11: 229, 230. 19 My 192 1. 

From Panama and Guatemala. 

Pittier, H. The Venezuelan mahogany, a hitherto undescribed 

species of the genus Swietenia. Jour. Washington Acad. 
Sci. 10: 32-34. 19 Ja 1920. 

Swtbtenia Candotlei, sp. nov. 

Pritchard, F. J. Relation of horse nettle {Solanum caroUnense) 

to leafspot of tomato (Septoria Lycopersici). Jour. Agr. 
Research 21: 501-506. pL 95~99- i Jl 1921. 

Ramaley, F. The role of sedges in some Colorado plant 

communities. Am. Jour. Bot. 6: 120- 130. /. 7, 2. Mr 1919. 

Richards, B. L. Pathogenicity of Corticium vasum on the 

potato as affected by soil temperature. J 

21: 459-482. pi. 88-93. i Jl 1921. 

Ricker, P. L. A sketch of botanical activity in the District 

of Columbia and vicinity. Jour. Washington Acad. Sci. 
8: 487-498. 19 Au 1918; 516-521. 19 S 1918. 

Ricker, P. L. A synopsis of the Chinese and Formosan species 

of Albizzia. J 
Ap 1918. 


Index to American Botanical Literature ^i^ 

Rose, J. N. A new Echeveria from Mexico. Jour. Washington 

Acad. Sci. i: 267-269. f, 7, 4 D 191 1. 

Echeveria lutea Rose. 

Rydberg, P. A. Notes on Rosaceae — XIIL Bull, Torrey Club 

48: 159-172. I Au 1921. 

Saflford, W. E, Acacia cornigera and its allies. Jour. Washing- 
ton Acad. Sci. 4: 356-368. 19 Jl 1914. 

Includes 10 new species from Mexico and Central America. 

Saflford, W, E. Chenopodium Nuttalliae,' a food plant of the 

Aztecs. Jour. Washington Acad. Sci. 8: 521-527. /. i-j. 
19 S 1918. 

A new species from Mexico. 

Saflford, W. E. Identity of cohoba, the narcotic snuff of ancient 

Haiti. Jour. Washington Acad, Sci. 6: 547-562. /. i-j. 
19 S 1916. 

Saflford, W. E. New or imperfectly known species of bull-horn 

acacias. Jour, Washington Acad. Sci. t 

Includes 3 new species of Acacia horn Mexico. 


Saflford, W. E. Notes on the genus Dahlia with descriptions 

of two new species from Guatemala. Jour. Washington 

Acad. Sci. 9: 364-373- /• ^-4^ I9 Jl I9i9- 
Saflford, W, E. Papiialthia mariannae, a new species of 

Annonaceae from the island of Guam. Jour. Washington 
Acad. Sci. 2: 459-463. /. i, 2. 19 N 1912. 

Saflford, W. E. Proposed classification of the genus Rollinia 

with descriptions of several new species. Jour. Washington 
Acad Sci. 6: 370-384. /. j-j. 19 Je 1916. 

Includes 3 new species from tropical America. 

Saflford, W. E. Rolliniopsis, a new genus of Annonaceae from 

Brazil. Jour. Washington Acad. Sci. 6: 197-204. /. J, 2. 

19 Ap 1916. 

Includes 2 new species. 

Saflford, W. E, Synopsis of the genus Datura, Jour. Washing- 
ton Acad. Sci. 11: 173-189. /. 1-3. 19 Ap 1921. 

Includes i new species from Colombia and 4 from Ecuador. 

Shreve, F. A map of the vegetation of the United States. 

Geog. Rev, 3: 119-125. F 1917 [with map.] 

Solis, O. Importancia de los jardines y algunas especies vegetale 

exoticas. Rev. Agricola 6: 78-84. Je. 192 1 [Illust.] 

74 Index to American Botanical Literature 

Standley, P. C. Ammocodon, a new genus of AUioniaceae, from 


the southwestern United States. Jour. Washington Acad. 
Sci. 6: 629-631. 4 N 1916. 

Standley, P. C. Blepharidium, a new genus of Rubiaceae from 
Guatemala. Jour. Washington Acad. Sci. 8: 58-60. 4 F 

Standley> P. C. Comparative notes on the floras of New Mexico 
and Argentina. Jour. Washington Acad. Sci. 6: 236^244. 
4 My 1916. 

Standley, P. C. A new species of Rondeletia from Mexico. 
Jour. Washington Acad. Sci. 8: 126, 127. 4 Mr 1918. 

Rondeletia Rekoi Standley. 

Standley, P. C. The North American species of Agonandra, 
Jour. Washington Acad. Sci. 10: 505-508. 4 N 1920. 

Includes 2 new species from Mexico. 

Standley, P. C, The North American species of Genipa. Jour. 


Washington Acad. Sci. 8: 639-643. 4 D 1918. 

Includes 2 new species from Panama. 


Standley, P. C. Notes on Orlhopterygitim htmucid. Jour. 
Washington Acad. Sci. 5: 628-631 - / J. 19 N 1915. 

Standley, P. C. Omiltemia, a new genus of Rubiaceae from 

Mexico. Jour. Wash 


Standley, P. C. Tidestromia, a nev^ generic name. Jour. 
Washington Acad. Sci. 6: 69, 70. 4 F 191 6. 

Xew name for Cladolhrix S. Wats., 1880, preoccupied by Cladothrix 
Cohn, 1875. 

Stevens, F. L. Bacteriology in plant pathology. Trans. Am. 

Microscop, Soc. 36: 5-12. Ja 1917. 

Swanton, J. R. Note on the aboriginal name *'aje". Jour. 



W. T, Chaetosp 


citrous fruits. Jour. Washington Acad. Sci. 3: 99-102. 

19 F 1913. 

From the Philippine Islands, 


Thaxter, R. Preliminary descriptions of new species of Rickia 

and Trenomyces. Proc. Am. Acad. Arts. & Sci. 48: 365- 
386. S 1912. 

Includes new American species in Rickia (i) and Trenomyces (4). 

Vol. 49 

No. 4 




APRIL, 1922 

The phylogeny of the genus Brachiomonas 

Tracy E. Hazen 

(with plates 3 AND 4 AND FIVE TEXT FIGURES) 

BrachiomonaSy one of the oddest and most Interesting genera 
of the Chlamydomonas group, has not until now been reported 
outside of western Europe. The genus was established by 
Bohlin (1) In 1898 with two species, differing in details of form, 
found in Sweden near Stockholm. The more slender species, 
B. gracilis, has not been reported elsewhere. The type species, 
B. suhmarina, had been collected about three years earlier by 
Lagerheim (6, p. 7) near Tromsj^, Norway, and recorded as a 
nomen nudum which was taken up by Bohlin. A somewhat 



It appears 

that Dangeard (3, p. 74) had observed the same form about 
ten years earlier than Bohlin, at Luc-sur-Mer, France, but had 
postponed publishing it until gametes could be found. Chodat 
(2, p. 143, /. 66) has recorded the species from Ajaccio, Corsica, 
showing that It Is not exclusively of northern habjtat. 

West (12) in 1908 reported a form collected in brackish 
marshes at Sheerness, England, near the mouth of the Thames, 
which he rather hesitatingly referred to B. stibmarina, since It 
seemed to be somewhat Intermediate between Bohlin 's two 
species. In July, 1920, at CuUercoats, near Newcastle-upon- 
Tyne, and In August at Plymouth, England, I found abundant 
material In small rock pools, showing forms similar to those des- 
cribed by West, together with specimens agreeing more with 
Bohlin 's type. Earlier In the same season (June 25) I had 
found on the coast of Norway at Valdersund, near Trondhjem, 

[The Bulletin for March (49: 51-74. pi 2) was issued April 12, 1922.] 


76 The phylogeny of the genus Brachiomonas 

a few specimens which showed precise gareement with Bohlin's 
description of B. submarina. Again a few days later, at Aale- 
sund, south of Trondhjem, I collected typical material, and here 
in great abundance. At this station some pools were almost 
exclusively filled with slender forms corresponding to Bohlin's 
B. gracilis; in other pools a mixture of the two species, with 
perplexing intermediate forms, appeared. Observing pools con- 
taining a natural pure culture, one would scarcely question 
the distinctness of the two species: when both were found 
together in the same pool, the variet}^ of forms seen presented a 
very strong suggestion of hybridization between the two species. 
New material collected in New York in March and April, 
1 92 1, has furnished forms (Figs. 5-7) almost precisely like the 
slender cells illustrated by West, together with others showing 
every gradation to the broadest type of B, suhmarina. These 
slender forms closely resemble Bohlin's /. ib of B. gracilis, 
but I am now convinced that West was entirely justified in 
including them in the very variable broader species. We seem 
not to have in America, and I did not find in England the ex- 
treme forms represented by Bohlln 's /. la and ic of B. gracilis. 
Whether they are sufficiently distinct to warrant the main- 
tenance of the second species is a question which would be more 
convincingly settled through studies by pure culture methods. 
I first found Brachiomonas suhmarina in March, 1907, in 
a small rock pool on the shore of Long Island Sound at Twin 
Island, Pelham Bay, New York. It was found In almost pure 
culture, coloring the water green from its abundance, and per- 
sisted in the motile condition for some weeks when brought to 
the laboratory. Although the same station was visited many 
times during the following twelve years, no Brachiomonas was 
found until -the middle of November, 1919, when it was again 
abundant in the same pool, and occurred sparingly in neighbor- 
ing rock hollows of similar character. I found it repeatedly 
in the same pools until the first of December, in spite of hard 
frosts and Ice formation during these two weeks. The habitat 
appears to be very like that of Bohlin's type station, small 
hollows so high above ordinary tide limits as to seem to be filled 
only with rain water, which are nevertheless sometimes dashed 
by waves so as to become brackish, as evidenced by taste and 
by the salt incrustation left on the rock margin upon evaporation. 
About the time of the collection in 1919, an unusually high tide 
was reported, and It is possible that this may have washed the 

The phylogeny of the gen"us Brachiomoxas 77 

Brachiomonas into the pool from the open sea. Such an occur- 
rence might account for the sporadic appearance of the species 
at this station, though it is much more probable that it is usually 
present in a resting state and that conditions were not favorable 
for the production of zoospores at the time of other visits to the 
station. A strange point is that, although careful examination 
of the specimens collected in the spring of 1907 shows them all 
to be like Bohlin's type, as were also the few specimens found 
there in September, 1920, and abundant material from March to 
June and in September, 192 1, only a few such individuals were 
found in the fall of 19 19 (Figs. 1,2), while most of the material 
showed a distinct difference in form (Figs. 8-16). This can 
hardly be interpreted as a mere seasonal variation. New mater- 
ial collected the last of February of the present season, from 
pools still coated with ice, consisted very largely of the new 
obtuse form, but after a week or more in the laboratory this 
form had mostly been replaced by individuals corresponding to 
Bohlin's type, present only in small numbers at the time of 
collection. Whether this is a case of the new type actually 
giving rise to the standard type through reproduction, or merely 
an example of the regular succession of growth of different organ- 
isms familiar in collections of protozoa, is a matter calling for 
more extended observation- For the determination of the per- 
manent status of the new form, pure culture methods are highly 
desirable, but thus far I have been unable to grow Brachiomonas 
on seawater agar, which succeeds admirably with other chlamy- 
domonads of similar habitat. For the present it will be con- 
venient to discuss this divergent plant under the name 

Brachiomonas submarina Bohlin, forma obtusa f. nov. 

This new form is entirely like Bohlin's type in its genera 
shape; that Is, it possesses a long posterior extension or horn 
and four shorter median arms directed backwards, often with 
depressions between them; the anterior portion might be roughly 
likened to the shape of an umbrella with four ribs. There is 
often a wedge-shaped beak through which the cilia emerge, 
though sometimes the apex appears merely rounded. The 
protoplast may fill the arms and posterior horn entirely (Fig. 8) 
or partially (Fig. 12) or not at all. The massive chromatophore 
may be equally variable; that is the protoplasm in the horn 
and in the arms may be all green, as also in the t>'plcal form 
(Fig, 2), or it may be colorless, sometimes appearing sharply 

78 The phylogeny of the genus Brachiomonas 


delimited from the central green mass (Fig. 8), while In other 
cases the green and colorless regions merge into one another 
almost imperceptibly (Fig. i). The chromatophore is here 
certainly less definitely or constantly organized than in most 
species of Chlamydomonas ] in fact it would perhaps be more 
accurate not to speak of a definite chromatophore but merely of 
cytoplasm with chlorophyll more or less diffused through it. 
This variability in the extent of the distribution of the chlorophyll 
in the protoplasm is very similar to the condition in Hydro- 
dictyon, to which Harper (5, p. 179) has called attention. West 
(12) states that in his specimens of B, stibmarina the chroma- 
tophore never extended into the arms. This was undoubtedly 
due to the fact that he never saw his material when freshly 
collected, but only after It had been sent from near the mouth 
of the Thames to Birmingham. I find that very generally in 
freshly collected specimens the chromatophore fills the arms 
and posterior horn, but often retracts from them soon after 
being subjected to the less favorable light and aeration conditions 
of captivity. 

Imbedded in the chromatophore in a subparletal position, 
regularly on the side next to the stigma, is a single relatively 
large pyrenoid; in vigorous young specimens this is generally 
situated in front of the middle of the cell; often in older Individ- 
uals it becomes posterior in position. The conspicuous red 
eye-spot is linear in form ; or, sometimes at least, it may be very 
narrowly wedge-shaped, pointed in front and broader behind; 
it lies in the depression midway between two arms. Most 
frequently when the cell is at rest the pyrenoid and stigma lie 
in a median ventral position (Fig. 6). On the other side of the 
cell from the pyrenoid the nucleus may sometimes be detected 
without staining, lying in a colorless mass of cytoplasm, best 
seen in polar view (Fig. 13). When the pyrenoid lies in its 
characteristic anterior position the nucleus is likely to be some- 
what posterior (Figs. 15, 17); if the pyrenoid becomes posterior 
the nucleus is then more often median in position. In older 
stages vacuoles often develop in the chromatophore (FiG. 4) 
as described by West (12). 

In hanging drop cultures, or in jars kept for some time in 
the laboratory, many individuals may be found which show little 
or no trace of the four lateral arms (Figs. 14, 15). Bohlin has 
shown such a form in one dividing individual. The cell wall is 
of so firm a character that it is difficult to imagine how the arms 

The phylogeny of the genus Brachiomonas 79 

could be modified or suppressed after they are once formed; 
nevertheless in practically all examples of division seen, I have 
found the arms well developed before the escape of the daughter 
cells (Figs. 20, 21) so that such cells can hardly be looked upon 
as a reversion to the simpler ancestral form. 

. The main difference between this new form and Bohlin's 
type lies in the fact that the arms are here always rounded and 
obtuse, while in the type form they are sharp pointed and often 
more slender. The characters of the form may be summarized 
as follows: 

Brachiomonas submarina Bohlin, forma obtusa f. nov, 
B. celluHs vegetativis figura structuraque formae typicae 
similibus, sed brachiis cornuque posteriore obtusis; cytoplasmate 
et chromatophoro alias in extrema ipsa extendentibus, alias ex 
eis plus minusve retractis. 

Longit. cellularum 15-32/i (saepius 20-25/1); lat. cell, 15- 
22 fx (saepius ca. 18^). 

Hab. in aquis subsalsis lacunarum saxearum. Twin Island, 
Pelham Bay, New York; Nov., 1919, Feb.-March, 1922. 

Chodat's figures (2, /. 66) Indicate that the form collected 
in Corsica may be identical with the one here described, though 
it is possible that the difference between his sketches and those 
of Bohlin may be due to the personal equation in drawing. 

As indicated above, I have found this obtuse form only at 
one station. Bohlin's type, as represented by my Fig. 1-3, I 
have also collected early in September during each of the past 
two vears at Bass Point, Nahant. Massachusetts- It should 



In older cultures numerous cells were found which showed 
the entire protoplast contracted into a globose mass (Fig. 16). 
These showed no further development in this material; but in 
fresh hanging drop cultures of B, submarina followed for a much 
shorter time in Norway, such cells presently became quiescent 
and developed a new spherical wall closely investing the pro- 
toplast. Soon these aplanospores showed their chromatophore 
turning from green to a tawny tint, and finally the orig-nal 
wall of the zoospores gradually disappeared. 

Practically all genera allied to Chlamydomonas possess two 
contractile vacuoles pulsating more or less in alternation, 
situated near the base of the cilia. In Brachiomonas these 

appear to be entirely absent. 
/. II; 14, /, loi c), it is tru< 


8o The phylogexy of the genus Brachiomonas 

beautiful original sketches in color, preserved with his collection 
of drawings in the South Kensington Museum, show that these 
spots are brown colored granules suspended in the fluid of a 
large permanent vacuole. The absence of contractile vacuoles 
I have been inclined to regard as another mark of the highly 
specialized character of the genus, since they are regularly to 
be expected in primitive members of this group and its ancestors. 
It is possible, however, that this peculiarity is merely an incident 
due to the special habitat; for Btitschli has remarked the general 
absence of contractile vacuoles in marine Ciliates and Rhizopods, 
and D'Arcy Thompson (11, pp. 165 and 264) regards this 
absence in such cases as ''no more than a physical consequence 
of the different conditions of existence In fresh water and in 

^ w 

salt. " The marine species, Chlamydomonas hrachyura West, 
besides exhibiting other points of resemblance to Brachiomonas^ 
appears also to have no contractile vacuoles; this species was 
discovered In association with a new Carteriaceous genus 
Platymonas, in which West (13) found two small contractile 
vacuoles, which, however, ''could only be observ^ed with dif- 
ficulty." In two brackish water species of Platymonas which 
I have investigated with considerable care, I am unable to 
satisfy myself that I ever see contractile vacuoles. Lewis has 
just published figures of one of these species, P. suhcordiformis 
(Wille) Hazen,* which likewise show no contractile vacuoles- 
Furthermore, two new brackish water forms of the primitive 
family Polyblepharidaceae, one a minute species of Pyramimonas 
and the other a new genus most closely related to Polyhlepharides, 
which have turned up in my latest collections of Brachiomonas, 
made while this paper was in the hands of the printer, very 
certainly possess no contractile vacuoles. Over against these 
several records of their absence, however, attention may be 
called to their presence (though here very small) in Chlamy- 
domonas caudata Wille, discussed below, which must be con- 
sidered as regularly a marine organism. 

Asexual reproduction is accomplished by internal division 
into four or eight (very rarely perhaps two) daughter cells. 
The division very generally begins while the cell is swimming 
actively, and often the typical cell shape is acquired by the 
daughter cells or zoospores and they move with their own cilia 
inside the mother cell-wall while it continues in active movement 


♦ Notes from the Woods Hole Laborator>', — 1921* Platymonas suhcordi- 
formis (Wille) Hazen, Rhodora 23: 249-251, pL ijj, /. /-/p. Mr 1922. . 

The phylogexy of the genus Brachiomonas 8i 

by its own cilia. Bohlln states that the first division is longi- 
tudinal, and certainly this is regularly the case. Chodat 
presents a figure (2, /. 66), and I have seen two or three such 
cases, where the first division appears to be transverse, but it 
is quite possible that such an appearance is due to an unusual 
shifting of the position of the cell contents after a proper longi- 
tudinal division. The subsequent divisions have not been 
described or figured by previous observers, and are ordinarily 
difficult to follow because of the continual active movement 
of the mother cell. I was so fortunate as to find one very 
favorable case in B. suhmarina f. obiusa, and later many similar 
ones in the type form, which I believe to be entirely representa- 
tive. This individual became quiescent after the first division 
was completed (Figs. 22, 23) ;it presented its anterior pole upward, 
so that it was possible to see easily that the second plane of 
cleavage is also longitudinal and perpendicular to the first. 
Immediately after the second cleavage a slight stretching and 
shifting of the daughter portions took place, so that one of them 
appeared somewhat underneath two others (Fig. 24). The 
third cleavage appears to cut each of the four daughter portions 
across its longer diameter, that is the plane seems to be essentially 
transverse to the previous divisions and to the axis of the mother 
cell. The pyrenoid disappears before the first cleavage occurs, 
and the stigma fades out during this process. A new and 
rather small pyrenoid appears in each daughter portion after 
the second cleavage, and probably another pyrenoid arises de 
novo in four of the eight daughter cells. About an hour after 
the completion of the third cleavage the eight daughter cells 
had acquired the typical form of the arms, and also the eye- 
spot and cilia (Fig. 27); after about another hour they escaped 
from the mother cell-wall. In certain cases it is clear that the 
cilia of the mother cell remain connected by a strand of proto- 
plasm to one of the daughter portions during division (Fig. 19) 
or even until the eight zoospores acquire their final form. 
Whether this is regularly the case appears doubtful. In mate- 
rial identified as Brachiomonas gracilis Bohlin, at Aalesund, 
Norway, I observed numerous cases of division which followed 

the course outlined above- 
In the case of the production of only four daughter cells 
it is clear that the cytoplasmic polarity of the mother cell 
would be directly transferred to each of the new zoospores. 
But when division proceeds further it is less clear how the 

82 The phylogeny of the genx^s Brachiomoxas 

polarity may be transmitted: for, as stated above, the third 
plane appears to cut the four daughter portions transversely 
in such a manner that half of the resulting eight would retain 
the original anterior cytoplasm, while the four posterior zoospores 
would seem to be under the necessity of regenerating (so to 
speak) their anterior pole. It is possible that in the slight 
shifting of the daughter portions following so quickly upon the 
first tw^o divisions there is actually more rotation than is apparent 
so that the third division may effect a properly qualitative 
bipartition, but that appears doubtful from the cases observed. 
The whole problem of polarity in chlamydomonads is one of 
great interest demanding a degree of care in observation which 
hardly any species up to the present has received. 

Brachiomonas sim 



later at CuUercoats, near Newcastle-upon-Tyne, and In August 
at Plymouth, England. It was found in small rock pools of 
brackish water, sometimes in association with one of the other 


species of Brachiomonas^ then again In practically a natural 
pure culture. 

At first sight the organism looked like a Chlamydomonas, 
many Individuals (Figs. 31, 39) showing somewhat the general 
oval, posteriorly pointed form characteristic of C. caudata 
Wille (Figs. 46-50), which occurred near by, though never in the 
same pools. But presently it was noticed that even the cells 
most nearly approaching C. caudata in form were nevertheless 
more like Brachiomonas submarina in internal organization, 
showing a similar linear stigma and large lateral and anterior 
pyrenoid; furthermore many individuals show small bumps or 
protuberances slightly back of the middle region, immediately 
suggesting the arms of Brachiomonas, though never really at- 
taining that character (Figs. 28, 29). The posterior horn, often 
rather elongated, shows a characteristic curv-e not seen in B. 
submarina. Generally the protoplast was conspicuously fur- 
nished with vacuoles, but whether this is a constant character 
is somewhat doubtful, inasmuch as C. caudata in similar pools 
was often similarly vacuolate at this time. The English ma- 
terial, moreover, was much less vacuolate. There is great 
variability in the development of the lateral protuberances; if 
formed at all they appear before the escape of the zoospores 

The phylogeny of the genus Brachiomonas 83 

from the mother cell, and the form then attained persists 
through the life of the individual In newly formed zoospores, 
usually, but not always, the protoplast and chromatophore ex- 
tend into the posterior horn so as to fill it completely. In 
older individuals the protoplast gradually retracts so that a 
considerable portion of the horn, or finally all of it is empty. 
The pyrenoid and red eye-spot regularly lie on the same side of 
the cell, and the nucleus on the opposite side; as in B. sub- 
marina, this is most clearly shown in cells momentarily resting 
with one of the poles directed upward (Fig. 32). 

As in the other species, asexual reproduction is accomplished 
by division of the protoplast into four or eight daughter cells 
which develop the characteristic form while the mother cell 
still retains its cilia in active movement (Figs. 35, 36). 

In sexual reproduction, gametes may be formed to the 
number of thirty-two in one mother cell, which retains its 


motile condition until they are completely developed (Fig. 40) 
just as in the case of asexual reproduction.. The gametes are 
similar in form to the asexual cells, but much smaller, lacking 
a cell wall, and furnished with relatively longer cilia: they 
also seemed to be more apt to show angularity in the middle 
region than the asexual cells. Usually there was a slight differ- 
entiation between the pair, a smaller and more fusiform gamete 
conjugating with one slightly larger and more angular (Figs. 
41, 42). Meeting by the ciliated anterior ends, the fusion is 
lateral, resulting in a quadriciliate zygote of ovoid -pyrif or m 
shape which swims actively, and may for some time show the 
two nuclei, two pyrenoids, and two linear eye-spots (FiG. 43); 
when momentarily at rest the four long cilia trail backwards. 
Finally the nuclei fuse, the cilia disappear (Fig. 44) the form 
becomes spherical, and the thick wall of the resting zygospore 
is developed (Fig. 45). After some days the cell contents 
assume a reddish orange color similar to that of the aplanospore 

of this species and of B. suhmarina. 

Brachiomonas simplex sp. nov. B. cellulis vegetativis 
majoribus, plerumque oviformibus quidem sed ad posteriorem 
partem in cornu curvatum productis, brachiis quae ceterae 
hujus generis species habent aut omnino suppressis aut evolutis 
tantummodo in 2-4 gibbos exiguuos subangulares vel rotundatos; 
ad polum autem anteriorem globosis vel papilla conica par\'ula 
exornatis: chromatophoro cellulam prope complente aut non 
longe in cornu posterius se extendente, pyrenoidem lateralem 
modo in medio modo a fronte habente, saepe vacuola majuscula 

84 The phylogeny of the genus Brachiomonas 

plura praebente: puncto rubro (stlgmate) lineari prope pyre- 
noidem sito: nucleo ad latus oppositum cellulae intra partem 
chromatophori excavatam complexo: ciliis binis ca. % breviori- 

bus quam cellula instructis. 

Propagatio fit per cellulae vegetatlvae impigre natantis 
divisionem in 4 vel 8 zoosporas, quae tegumenti scissura erum- 
punt. Praeterea aplanosporae, primo virides, postea paulatim 
ad colorem fulvum transmutatae, per contractionem protoplasti 
in massam globosam ad cellulae partem anteriorem, ut in B, 
submarina, efficiuntur. 

Generatio fit per copulationem gametarum inter se subae- 
qualium et nudarum, quae cellulis vegetativis forma quidem 
similes sunt, sed perparv^ae et ciliis quam corpus ipsum multo 
longioribus praeditae, quaeque per divisionem cellulae vegeta- 
tivae protoplasti etiam in 32 individua parturiuntur. Zygo- 
sporae globose, tegumento levi vestltae, diametro ca. 12^. 

Longit. cellularum veg. 30-48^, lat. iS~2^fi, Longit. game- 
tarum 13-15/X, lat. 6-8yn; longit. ciliorum ca. 18^. 

Hab. in aquis lacunarum saxearum subsalsis. Aalesund, 
Norway, 28 June, 1920; CuUercoats, Northumberland, England, 
19 July, 1920; Plymouth, England, 11 August, 1920. 

Because of the fact that unarmed individuals appear in 
older cultures of B, suhmarina, one might be tempted to inquire 
whether -S, simplex is merely a form of that species. Never- 
theless in the natural pure cultures seen both at Aalesund and 
at Plymouth, B. simplex maintained its characteristic simplicity 
of form from the earliest development of the daughter cells 
to maturity, and even when it was found intermingled with one 
of the other species it showed a distinctive character, parti- 
cularly in the curving of the posterior horn, which rendered 
It always recognizable. Again, the variability of form exhibited 
by this new species might readily suggest for it a hybrid origin. 
We know little of hybridization among algae, but a very inter- 
esting preliminary report of its occurrence in Chlamydomonas 
has recently been published by Pascher (7). It may be remarked 
that Pascher 's heterozygous cells showed great diversity in 
internal organization, while here in B. simplex only variability 
of external form is found. From a careful consideration of all 
the conditions of its habitat and its behavior, I am convinced 
that B, simplex is to be regarded as a distinct species which 
represents the transition from Chlamydomonas to the more 
characteristic species of Brackiomonas. 

There is some resemblance between this species and Chlamydo- 
monas brachyura West (13) a form which developed in a tank of 
seawater sent from Plymouth to Birmingham in I9i5, and for 

■ ■ 

The phylogeny of the genus Brachiomonas 85 

which I searched vainly at Plymouth in 1920, Nevertheless 
since C brachyura lacks a red eye-spot and Is pointed instead 
of rounded at the anterior end, we may perhaps find more 
suggestion of the origin of Brachiomonas in the species mentioned 


at the beginning of this section, namely, 

Chlamydomoxas caudata Wille 


This very Interesting and apparently rare species was found 
June 28, 1920, at Aalesund, Non\^ay, in small rock pools of the 
same character as indicated in the original account in 1903 
(15, pp. 115-118, 135, pL J, f. 4~ii) and my determination of the 
species was later confirmed by Professor Wille, from living mater- 
ial which I carried to him in Christianla. In one pool was a practi- 
cally pure culture, giving an intense green color to the brackish 
water; in another pool close by was a considerable admixture 
of C siibcaudata Wille, which was certainly the most abundant 
species of the region, now as in 1902. Why C. caudata should 
occur in only three or four out of many similar pools is an inter- 
esting question. It was also surprising that it could not be 
found on the English coast, where the conditions appeared to 
be much the same, and where I did find several of the other 


brackish -water species which were associated with C. caudata 

at Aalesund. This species appears not to have been found 

elsewhere than at this original station. 

The elongated oval posteriorly pointed form of the zoospores 

is very notable in comparison with the rounded form of the 
commonly known species of CJilamydomonas , The strong 
resemblance to the more reduced forms of Brachiomonas sim- 
plex is equally striking; so much so. In fact, as intimated above, 
that upon superficial observ^ation the two species might be 
mistaken for closely related members of the same genus. Ordi- 
narily in active individuals, especially In newly formed zoo- 
spores, the protoplast conforms rather closely to the shape of 
the cell-wall, being merely rounded at the point of insertion of 


* Plavfair, to be sure, has reported [Proc. Linn. Soc. New South Wales 43: 
515. 1918] a form found apparently in fresh water of which he writes: 
'*The only tailed species of Chlamydomonas; It is impossible to make any 
mistake in the identification," Since, however, Playfair states that his 
specimens lacked pyrenoid and stigma, and his figure shows a distinctly 
different morphology of the anterior portion of the cell, including a different 
position of the nucleus, there can be no justification for trying to identify his 
form with a species so well defined as C caudata, at least until cultural evidence 
is brought forward to support the attempt. 


The fhylogeny of the genus Brachiomonas 


the cilia, and tapering to an acute point at the posterior end 
Fig. 46), Older individuals, kept for 


hanging drops, show the cell-wall much thickened, or the pro- 
toplast contracted and rounded posteriorly and produced into 
more or less of a beak at the anterior end (Figs. 47, 48). The 
broad truncate anterior papilla of the cell-wall (Hautwarze) 
impresses me as being rather more prominent than shown in 



Figs. 46-50. Chlamydomonas caudata Wille 

46. Young cell, "dorsal view/ ' showing typica Iposition of nucleus, stigma, 
and pyrenold. 47. Older cell with thickened wall, 48. Similar cell in 
"lateral view." 49. Typical arrangement of daughter cells. 50. Unusual 
arrangement of daughter cells. All, X 720, approximately. 

Wille 's (15) figures, and the red eye-spot, first described as 



my mature specimens always a strongly oval disc, sometimes 
with an anterior pointed end, showing its thinness in the fact 
that the edge view was narrowly linear. These differences 
may be readily accounted for by differences of magnification 
during observ'ation. The longitudinal streaking of the chro- 
matophore noted in the original description, and even Intro- 
duced In Wille 's key to the species, was never distinguishable 
in vigorous material of either this species or C. subcaudata m 
1920; only certain individuals kept for ten days or more in 

The phylogeny of the genus Brachiomonas ^7 

hanging drop cultures showed almost a lattice-like appearance 
of the chromatophore. Although the chromatophore is hollowed 
out about as far as the middle of the cell, or sometimes even 
more deeply, nevertheless the position of the nucleus is notably- 
less central than might be expected, for it lies approximately 
in the anterior third of the cell. 

A point of considerable interest is the position of the stigma 
and contractile vacuoles. Regularly the eye-spot is so placed 
that when the cilia are stretched out in a plane parallel to the 
stage of the microscope the stigma lies on the apparent right 
side of the cell and in a slightly upper focus, or with the cell 
revolved 180° the stigma lies on the apparent left side and at a 
slightly lower focus; in such a position only one of the small 
contractile vacuoles is visible. If the cell is revolved only 90' 
from the first position, so that one of the cilia lies nearer the 
observer and the other at a somewhat lower focus, the stigma 
shows its disc-like shape in an upper central focus (Fig. 48) 
and the two contractile vacuoles may be seen side by side at 
the same time, though one is always smaller than the other, 
on account of the alternation in their pulsations. This position 
of the stigma and cilia in one plane and the contractile vacuoles 
In another plane perpendicular to it Indicates a dorsi-ventral 
character of the cell which appears to me to be the rule In many 
species of Chlamydomonas, though it seems to have received 


little or no attention from students of the group. 

The dorsiventral differentiation in Brachiomonas, as men- 
tioned above and indicated in Figs. 6, 7, 13, 15, and 32, Is 
slightly different; there the cilia normally stretch out in a plane 
perpendicular to that which passes through the stigma, pyrenold, 


and nucleus, and most commonly the zoospore comes to rest 
with the stigma lying exactly in the middle on the ventral side, 
or (less often) on the dorsal side, toward the observer. In 
many of the figures which seem to indicate a different relative 
position of the eye-spot and cilia, the latter were actually lying 
In a plane oblique to that of the microscope stage, which could 
not easily be shown In the drawings. 

* There are, it Is true, many published figures of Chtamydomonas which 
show the two contractile vacuoles side by side apparently in the same plane 
with the cilia; but in cells exhibiting so much movement it is very easy to 
transfer what is seen in one view to a sketch representing in general a different 
view, unless one is keenly on the lookout for such a point. Probably also 
many of our somewhat classic figures are not made from camera lucida draw- 
ings of quiescent individuals, as they should be to determine such features. 

88 The phylogenyof the genus Brachiomonas 

In the asexual reproduction of C caudata, Wille describes 
the division as longitudinal, after the disappearance of the cilia 
of the mother cell, resulting in the formation of four zoospores. 
On several occasions when I examined material with my travel 
microscope in the field in late afternoon, or freshly collected 
material in early evening, I found numerous individuals which 
contained the four fully formed zoospores while the mother, 
cell was still swimming actively- It was only in the more rarely 
seen divisions of material kept for a day or two after collection 
that I found non-motille cells in process of reproduction; such 
cells may have lost the power of movement earlier because of 
poorer aeration than in the normal habitat. This division of 
actively motile cells I have found to be normal in several other 
species of Chlamydomonas, I was unable to find examples 
which would actually show more of the details of the process 
of division than indicated by Wille, who merely furnishes one 
figure of completed division, I did, however, observe that a 
large majority of cases showed two of the daughter cells with ' 
their anterior end directed forwards, and two with the anterior 
end directed backwards (Fig. 49). In other species of Chlamydo- 


monas such as arrangement of the daughter cells is character- 
stically found when the first plane of division appears transverse 
to the longitudinal axis of the mother cell, or when, though 
fundamentally longitudinal, the plane becomes placed trans- 
versely by a rotation of the entire protoplast. During the 
past season I have had the opportunity of observing this 
latter method many times in a new species to be described in a 
future paper: this species furnished one exceptional case in 
which both divisions, carefully followed, were strictly longi- 
tudinal, resulting in four zoospores all headed toward the 
anterior end of the mother cell. This exceptional arrangement 
I also found in rare instances in C caudata (Fig. 50). Wille 's 
figures (15, pi. j, /. -8) which shows the same arrangement, 
appears, therefore to be untypical, and I am obliged to believe 
that the first division of this species, though properly longitudi- 
nal, would regularly show as a transverse cleavage, in consequence 
of a rotation of the protoplast to such an extent that its anterior 
pole comes to lie in a position nearly in the middle of the length 
of the cell. 


Reichenow (9, 10, pp, 35-39) it is true, has expressed the view 
that the division in all species of Chlamydomonas may be funda- 
mentally longitudinal, an apparent transverse division being al- 

The phylogeny of the genus Brachiomonas 89 

ways preceeded, in his opinion, by a rotation of the protoplast, so 
that he beHeves the manner of cell division to have no phyloge- 
netic significance. Attractive as this view is in its possibility of 
simplifying the idea of direct transmission of polarity to the 
daughter cells (and my own recent studies on Chlamydomonas 
somewhat tend to support such a view) nevertheless we still 
lack sufficiently definite and accurate observations to permit 
the denial that an actual transverse division may occur In some 

If in C. caudata the first cleavage plane is actually or by 
rotation of the protoplast transverse (as appears to me most 
probable), and if we accept the current view derived from Dill 
(4) that the most primitive species are those in which both 
divisions are longitudinal, then C, caudata could hardly be re- 
garded as an actual transitional form between the ordinary 
ovoid Chlamydomonas type and the Brachiomonas group, since 
all species of the latter genus appear to have retained the primi- 
tive mode of division by two strictly longitudinal cleavages, 
in spite of their advancement in other respects. Nevertheless, 
in its general morphology, and in the similarity of its method of 
formation of aplanospores (rarely found in other species of 
Chlamydomonas), C, caudata must be very close to the ancestral 
line which has given rise to the four forms of Brachiomonas 
recognized above; and these present a rather close ascending 
series, with perhaps a definite orthogenetic tendency beginning 
with B. simplex^ which is barely emerging from the Chlamydo- 
monas stage, through J5, suhmarina /. ohtusa and then B. sub- 
marina (type) to jB, gracilis, an extremely slender form, so far 
removed from any Chlamydomonas type as to justify thoroughly 
the retention of Brachiomonas as a genus. 

In certain graphic schemes of the phylogeny of the chlamydo- 
monads — e. g. those of Wille (15) and West (14, p. 163)— the genus 
Lohomonas Dangeard is indicated as representing an inter- 
mediate stage between Chlamydomonas and Brachiomonas] but 
the two known species of Lohomonas, as well as tw^o new species 
to be described by the writer in the next number of this journal, 
are very minute forms, and in their general morphology exhibit 
by no means so good a connection as is shown by C caudata 
and B. simplex. Lohomonas must be regarded as a special 
offshoot from the Chlamydomonas line not leading to anything 
higher, so far as we know at present. 

Barxard College, 
Columbia University 

90 The phylogeny of the genus Brachiomonas 

Literature cited 

1. Bohlin, K. Zur Morphologie und Biologie einzelliger 

Algen. Ofvers. K, Vet. Akad. Forhand. 54: 507-529. 


2. Chodat, R. Algues vertes de la Suisse, Berne. 1902. 

3. Dangeard, P. A. M^moire sur les Chlamydomonadin^es 

ou rhistoire d'une cellule. Le Botaniste 6: 65-292. /. i- 
20. 1899. 

4. Dill, O. Die Oattung Chlamydomonas und ihre nachsten 

Verwandten, Jahrb. Wiss. Bot. 28: 323-358. 1895, 

5. Harper, R. A. The structure of protoplasm. Am. Jour. 

Bot. 6: 273-300. 1919. 

6. Lagerheim, G. Studien iiber artkische Cryptogamea. 
Troms^ Mus. Aarhefter 17: 1-24. 1895. 

7. Pascher, A. Uber die Kreuzung einzelliger, haploider 

Organismen : Chlamydomonas, . Ber. Deutsch. Bot. Ges. 
• 34: 228-246. /. 7-5. 1916. 

8. Printz, H. Kristianatraktens Protococcoideer. Vid.-Selsk. 
Skrift. I. Mat. Naturv. Klasse 1913«: 1-123. pi. 1-7 + /. 
/, 2. 1914. 

9. Reichenow, E. Untersuchuugen an Haematococcus pluvi- 

alls und eingen anderen Flagellaten. Sitz.-ber. Ges. Naturf. 
Freunde Berlin 1909: 85-91. 1909. 

10. . Untersuchungen an Haematococcus pluvialis 

nebst Bemerkungen iiber andere Flagellaten. Arb. Kais. 
Gesundheitsamte 33: 1-45. pi. i, 2. 1909. 

11. Thompson, D'Arcy W. On growth and form. Cambridge. 


12. West, G. S. Some critical green algae. 2 Brachiomonas 
submarina. Jour. Linn. Soc. Bot. 38: 281-283. pi. 20. 
f. 7-19. 1908. 

13. . Algological notes XX, XXI. Jour. Bot. 54: 

2-5. /. J, 2. 1916. 

14. . Cambridge Botanical Handbooks. Algae 1. 

Cambridge. 1916. 

15. Wille, N. Algologische Notizen XI. Uber die Gattung 
Chlamydomonas. Nyt Mag. Naturvid. 41: 109-162. pi 
3, f. 4~ii. 1903. 

Description of plates 3 and 4 

All drawings made with the aid of camera lucida from living mateaal 
in hanging drop cultures (or in some cases freshly fixed with osmic acid vapor) : 
Plate 3 done with Leitz 1/12 in. oil immers. obi. and compens. oc. 6, magnifica- 

The phylogeny of the genus Brachiomonas 91 

tion about 1440 diameters; plate 4 done with Spencer 2 mm. oil immers. obj. 
and Leitz compens. oc, 6, magnification about 1300 diameters; both reduced 
in reproduction to about 720 diameters. 

Plate 3 
Figs, 1-7. Brachiomonas submarina Bohh'n, type form 

Fig. I. Vegetative cell; posterior horn filled with colorless cytoplasm. 

Fig. 2. Chromatophore filling the extremities: Dec, 19x9- 

Fig. 3. Colorless cytoplasm in extremities; stigma and pyrenoid lying 
on the under side of the cell: March, 192 1. 

Fig. 4. Anterior polar view of similar cell; vacuoles in chromatophore. 

Fig. 5. Slender form similar to Bohlin's/. ib of 5. gracilis (fixed wii.h 
osmic acid vapor). 

Fig. 6. Moderately slender form: showing typical position taken by 
living zoospore temporarily at rest, the pyrenoid and stigma lying in the 
center of the under side (osmic vapor): April 1921. 

Fig. 7. Anterior polar view of similar cell, showing cilia stretching out 
in plane perpendicular to that in which the nutleLs, pyrenoid, and stigma 
topically lie. 

Figs. 8-27. Brachiomonas submarina forma obtusa Hazen 

Fig. 8, Chromatophore contracted, arms and horn filled with colorless 

cytoplasm (osmic vapor): November, 1919. 

Fig?. 9, 10. Arms empty, colorless cytoplasm extending into posterior 

horn (osmic vapor). 

Fig. II, Anterior polar view of similar cell, living. 

Fig. 12. Arms and horn nearly empty (osmic vapor). 

Fig. 13. Posterior polar view of similar cell, living, showing typical re- 
lative positions of organs as in Fig. 7; a large colorless cytoplasmic region 
around the nucleus. 

Fig. 14. Older cell showing no trace of arms. 

Fig, 15. Cell from old culture with only a suggestion of lateral arms; 
stigma and pyrenoid lying underneath the nucleus. 

Fig. 16. Senescent cell with contracted protoplast, preparing to form an 

aplanospore: December, 19 19. 

Fig, 17, Young cell showing nucleus posterior to pyrenoid. . 

Fig. 18. Posterior polar view of similar cell. 

Fig. 19. First division of motile cell; 10:30 p. m. (o^mic vapor). 

Fig. 20, Eight zoospores fully formed in motile mother cell. 

Fig. 21, Four zoospores: cilia of mother cell no longe/ present. 

Figs. 22-27. Successive stages in division resulting in eight zoospores 
(November, 1919): Fig. 22 at 2:40 p. m.; Fig. 23 at 3:40; Fig. 24 at 4:50; 
Fig. 25 at 5:30; Fig. 26 at 6:20; Fig. 27 at 7:40. The zoospores escaped 
before 8:40. 

Plate 4 
Brachiomonas pimpi-ex Hazen 

Drawn 29 June — 4 July, 1920, Aalesund, Norway. 
Figs. 28, 29. Typical mature vegetative cells: in Fig. 29 the stigma lies 
slightly underneath. 

Fig. 30. Anterior polar view of similar cell. 


92 The phylogeny of the genus Brachiomonas 

Fig, 31. Cell on which the "bumps" representing lateral arms are un- 

Fig. 32. Anterior polar view of similar cell. 

Fig. 33. Protoplast ertracted from one of the "bumps" (osmic vapor). 

Fig. 34. Large older cell with more posterior pyrenoid. 

Fig. 35 Eight zoospores in motile cell; 12:45 A. M. 

Fig. 36. Four zoospores in motile cell. 

Fig. 37. Young zoospore free; chromatophore filling posterior horn. 

Fig. 38. Three of the four zoospores have escaped through a triangular 
rent in the wall of the mother cell. 

Fig. 39 Similar to Fig. 31, but with more typical position of pyrenoid 

(osmic vapor) 

Fig. 40. Thirty-two gametes in motile mother cell; 10 A M. 

Figs. 41, 42. Gametes at beginning of conjugation (osmic vapor). 

Fig. 43. Motile zygote, momentarily resting; nuclei not fused. 

Fig. 44. Zygote, cilia having disappeared, nuclei fused; two stigmata still 

Fig. 45. Zygospore twenty-four hours after conjugation. 



In the living cells the pyrenoid actually appears upon casual 
observation to be green because imbedded in the green chromatophore. It 
is believed, however, that the printing of the pyrenoids in the same sepia 

as the cell-wall and protoplasmic outlines represents more truthfully their 
real nature. 

Note. As final proof of this paper is received, the writer has 
just made a mid-April excursion to the Massachusetts coast, 
where he has collected typical Brachiomonas suhmarina Bohlin 
from Cape Ann to Buzzards Bay, at Rockport, Gloucester, 
Marblehead, Nahant, and Fair Haven. There has been no in- 
dication of other species of Brachiomonas or of Chlamydomonas 
caudata at these stations. 

Bull. Torr 



IMli 49, PLATE ^ . 





















BlM.. ToKK» V C'l I'H 

\'oi.UMH 49, I'LATF 4 












New records and other notes on North Dakota plants* 

O. A. Stevens 


The present paper is supplementary to Bergman's Flora of 
North Dakota (in Sixth Bien. Rept. Agr. Coll. Survey [191 1- 
1912]. 1918) and, as such, will follow the same arrangement and 
nomenclature. The purpose is to record additional species, 
designated by a dagger (f), and other notes of special interest. 
During the years 1910 to 1920 I have had opportunity to visit 
nearly every part of the state and, incidental to economic work, 
to obtain many specimens of interest. Some of these were 
noted In the Flora and others have been reported by Lunell 

Nat. 4-6. 1917-1919). I have also included in the 
paper a few corrections which have come to notice and revised 
certain groups in which I have become especially Interested. 
Where not otherw^ise stated, the specimens listed have been 
collected by myself. 

Lepidium ramosissimUxM a. Nels. I do not know why 
Bergman reduced this to a synonym of L. densiflorum Schrad. 
The two species are very different and may be distinguished as 

follows: 1 J -u 

L. densiflorum. Annual, stem usually simple below and wtth 

several long racemes above; basal and lower stem leaves 

serrate; pods distinctly margined and broadest toward the 


L. ramosissimum. Biennial, bushy branched from the base; 
basal and lower stem leaves pinnatlfid, the lobes Imear or 
oblanceolate; pods not margined above, elliptic m outhne. 
The following specimens belong to L. ramosissimum : Tower 
City, Bergman 870; Valley City, Aug. 13, 1912, (first year 
rosettes); Courtenay, June 20, 1911; Kensal, July 26, 1915, 
(rosettes, also flowering stems); Manfred, July 12, 1913; Sykes- 
ton, July 15, 1891, Bolley; Sheyenne, July 4- 1908, Lunell; 
Leeds, Aug, i, 1912; Bathgate, July 22, 1892, Lee; Ganbetta; 
Bell 264 ; Rugby, Bergman 261 1. This species is quite common 
throughout the central part of the state, growing about streets 
and dooryards rather than in fields. At Langdon on July 19, 
1 9 18, I noticed the rose ttes of L. ramoisssim um fairly covering 

* Contribution from Department of Botany, North Dakota AgricuUnral 
College and Experiment Station. 


94 Stevens: North Dakota plants 

the ground. In a dooryard at Courtenay the species grew 
mixed with i. densiflonim. The latter had racemes about 3 cm. 
long with numerous weJl-developed pods, while the plants of L. 
ramosissimum had only a few half -developed pods. 

t Lepidium Draba L. Several years ago (Ann. Rept. 
North Dakota Exp. Sta, 22: 80. 1912) I called attention to the 
possibility of this being introduced in Turkestan alfalfa seed. 
On June 6, 1918, a flowering plant was sent me by Mr. H. A. 
Nelson of Ray. [A well estabhshed colony has since been found 
along the railroad track at Moorhead, Minnesota, May 22, 1921.I 

t Sisymbrium Loeselii L. Determination verified by Paul 
C. Standley and specimens deposited in herbaria of the U. S. 
National Museum and the New York Botanical Garden. 

Annual, 6-9 dm. high, sparingly hirsute with simple flattened 
hairs, these less abundant above but present on upper stems 
and pedicels; leaves runcinate, the terminal segment hastate 
and irregularly denticulate, lower stem leaves sometimes I dm. 
long; racemes becoming 3-4 dm. long, pedicels widely spreading 
one third to one half the length of the slender pods, which are 
ascending and about 3 cm. long with three-ner\ed valves. 

Devil's Lake, July 15, 1920 (common about the streets); 
Tappen, July 12, 1919 (a single plant in the street; several in 
an old field of Bromus inermis Leyss. several years before). 
The description was drawn from the growing plants and speci- 
mens collected at Devil's Lake. The plant has somewhat the 
aspect of Sophia intermedia Rydb., but the leaves are quite 
diff'erent and the racemes are fewer but longer and coarser. 
The relative height of flowers and young pods mentioned by 
Koch and others seems scarcely distinctive. In this plant the 
pods do not surpass the flowers in normally developed racemes, 
but examples are frequent where they do so, apparently as a 
result of arrested development. 

Erucastrum Pollichii Schimp. & Spenn. Park. River, 
Aug. 8, 1913; Williston, Aug. 15, 1915; Oakes, July 18, 1919J 
Cooperstown, Aug. 25. 1919. This has now been found at 
many places, chiefly along the railroad tracks. The seeds have 
been identified in several samples of timothy and millet coming 
from near Grand Forks or a short distance north or south of 
this locality. The pods are not flattened, as stated in Bergman's 
key, but rounded, the valves with a rather prominent mid- 
nerve; racemes leafy bracted. The plants grow vigorously m 
late fall as shown by a flowering specimen collected at Fargo, 
Oct. 26, 1914, 

Stevens: North Dakota plants 95 

Malva verticillata L. I have not seen the specimen 
which Bergman referred to this species but am inclined to think 
that it was M. horealis Wallm., which is the common plant at 
Fargo and other places in the state. A specimen of what seems 

to be M, crispa L. f was sent by A. G. Sorlie, Grand Forks, 
Oct. 10, 1916. 

Euphorbia Esula L. Wahpeton, June 3, 1920, Geo, P. 
Wolf; Hazelton, July 15, 1919, in field of Bromus inermis; New 
Rockford, July, 191 1, /. K Campbell; Mona, June 5, 1914, 
Albert Graves; Ray, July 2, 1917, H. A. Nelson, In one field 
near Fargo patches are scattered over 20 acres. Apparently 
a bad weed, spreading by the roots. Time and manner of in- 
troduction unknown. 

t SiLENE DiCHOTOMA Ehrh. Edgeley, July, 25, 191 7, 
Brenckle; Langdon, July 19, 191 8; Regent, July 20, 1916, A. 
C Goldtrap, In timothy fields. Plants grown from seed 

proved to be coarse 



t SiLEXE Fabaria (L.) Sibth. & Sm. Specimens determined 

by Paul C. Standley. 

Biennial; glaucous, much branched, 6-9 dm. high. Basal 
and lowest stem leaves narrowed at the base, larger stem leaves 
ovate, sessile; flowering stems dichotomous with a flower 
in the fork, each branch with several remote clusters of three 
to seven flowers each. Flowers about i cm. wide, white, opening 
at night; petals cleft to the throat, not appendaged, the lobes 
oblanceolate, rounded at the apex; calyx indistinctly fifteen- 
ribbed, closely investing the ovate horny capsule, but not 
especially enlarged in fruit. 

Venturia, July 16, 1919; Oakes, July 18, 1919. Lower stems 
and leaves with very much the aspect of Vaccaria Vaccaria, 
A specimen in the herbarium of Dr. J. F. Brenckle was collected 
at Kulm by him in 1916, and he states that he had found the 
plant there previous to that date. Apparently well established 
at Oakes and Venturia along roadsides. Dr. Standley writes 
that this is probably the first time it has been collected in this 


t Lychnis alba Mill. Devil's Lake, July 13, 1920. 


For several years I have been interested in C. album and 
related forms, especially in regard to seed characters. The 
plants are rather difficult to collect as the seeds are slow In 
maturing and the lower leaves are likely to be lost by the time 
the fruiting branches are well matured. With respect to the 
adherence of the pericarp it is to be noted that it rubs off more 
f^asily in fresh material. 

96 Stevens: North Dakota plants 

. In the fall of 1916 I collected twenty-six specimens in the 
vicinity of Fargo and submitted them to Dr. Standley, who 
determined them as follows: 

C album (4); 

t C paganutn Reichenb. (11); 

C paganuntj approaching C. album (8); 

t C. ferulatum Lunell (3). 
Apparently C- pagamim is the commonest form here. I 
have not as yet been able to make much further progress in 
clearing up the relations of these forms but wish to offer what 
few notes I have- 



open, exposing the fruit. While this does not seem t 
constantly true for C. ferulatum, I find it also in a rather variable 
degree in the C. album specimens, being most prominent in 
well-matured plants. No. it, determined as C paganum, has 


calyces closed in specimen collected Aug. 25, but fruit from same 
plant collected in September shows them quite open. As to C 
paganum, most of the plants were either so badly infested by an 
insect or so late in maturing that very little mature seed was 
obtained. From material at hand I would describe the seeds 
as follows: 

C album — 1.2-1.4 mm. wide, upper surface flattened and with 

shallow, oblong pits (which show through the pericarp). 
C. paganum — 1.2-14 mm. wide, upper surface dull, neither 

flattened nor pitted. 
ferulatum — 1.2 


■ ^^ 

C. lepiophlyllum — 0.9 mm. wide, not flattened, smooth and shining. 

The size of the seed of C. paganum is given by Standley 
(N. Am. Fl. 21: 21. 1916) as 1.3-2 mm. In one of the twenty- 
six specimens, a plant collected near the woods along the Red 
River, the seeds are variable in size, measuring 1.4-2 mm., but 
in all the others they are scarcely larger than those of C. album. 

The three specimens of C. ferulatum differ among themselves. 
One, collected in an old garden (No. 3), which I take 
to be typical, is pale grayish green, simple below and with short 
branches above, the leaves oblong-rhombic with four or five 
short teeth on each side; another (No. 9, labelled by Standley 
as apparently a form of C. ferulatum) Is bushy branched with 
narrow entire leaves. The third (No. 2) is in a rather advanced 
condition with lower leaves mostly fallen; the flowering branches 

Stevens: North Dakota plants 97 


are unusually thick and dense and the seeds are pitted on the 
upper side. I am inclined to think it is a plant of C. album with 
an unusual amount of well-matured fruits and therefore with 
the spreading calyces unusually prominent. 

The seeds of these species are described by Standley as 
"nearly smooth, black and shining" except in C femlatum, 
where they are said to be ''puncticulate/' but I find in our 
material that C. album has pitted seeds and C, femlatum 
smooth (excepting as in No, 2 just noted; type material from 
Lunell also examined). The seeds of C, leptophyllum seem to be 
quite easily distinguished from those of the other common field 
species by their smaller size. 

t Chenopodium pratericola Rydb. Narrows, Aug. 12, 
I9i3» Lunell. 


Wahpeton, Aug. 6, 1919. Along a 

street by buildings. Dr. Standley, who examined a specimen, 
writes that this species is perhaps only a form of A.hastata L., 
but we seem to have no other specimens which would be referred 
to this form. 

t Suckleya Suckleyana (Torr.) Rydb. Belfield July, 4, 



Following Standley 's treatment of this genus (N- Am. FK 
21: 79-80. 19 16) I would place our material as follows; 

C hyssopifolium L. Mandan, Wright 610. 

t C nitidum Kit. Sandhills near Anselm, Aug. 21, 19 18. 

t C. villosum Rydb. Sandhills near Anselm, Aug. 21, 
1918; also all other specimens referred by Bergman to C. hyssopi- 
folium, except Wright 610. The Fargo specimen of C villosum 
was collected along the railroad (doubtless introduced in gravel) 
and no plants have been observed since. It was a well matured 
plant with very little pubescence except on the bracts. The 
Anselm plants are young, upright, rather slender, and densely 

Polygonum Hydropiper L. Hankinson, July 30, 1919. 
Lunell (Am. Mid. Nat. 5: 184. 1918) has referred the Ft. Ransom 
specimen to P. punctatum leptostachyum (Meisn.) Small, but 
it has dull akenes and flowers in the lower axils, as in P. 

AscLEPiAs SYRiACA L. Westficld, July 16, 1919. This does 
not seem to be common so far west. The specimen reported 

gS Stevens: North Dakota plants 


speciosa Torr.; it is stenle but has the characteristic pointea 
leaves of that species. The specimen reported by Bergman as 
possibly a hybrid of the two species seems scarcely different 
from A. syriaca. 


Mr. W. W 

has examined our material of A. verticillata L. and refers the 
following to A. pumila: Esther, Bell 542; Fleak, Bell 1370. 



of Bergman 's catalogue should be referred to this species instead 
of to A. viridiflora Ell. 

t Veronica maritima L. Kongsberg, Sept. 191 8, Fred 


Schmidt, Jr. A few plants in a -grove of planted trees. Deter- 
mined by F. W. PennelU 

CuscuTA ARVENSis Beyr. The Logan County record is by 
Brenckle, Aug. 18, 1912. We have also the following records: 
McLeod,J?e//j5*5: Valley City, July 26, 1913; Enderlin, Aug. 20, 
1918. Dr. T. G. Yuncker, who has examined portions of the 
Logan County and Valley City specimens, writes that he would 
call them C. pentagona Engelm, The other specimens seem to 
be the form which he calls C. calycina Engelm. The former 
were growing upon plants of the dry prairie, while the latter 
preferred those of riverbanks and similar localities. 

CuscuTA Gronovii Willd. Fargo, Aug, 14, 1890, Waldron, 
Is the only specimen which I consider typical. When preparing 
my paper on Ctisctita (Am. Jour. Bot. 3: 185-188. 1916) I was 
unable to find fresh material, but all that I have found at Fargo 
since that time agrees better with what I had called C plattensis 
A. Nels. The habitat of C. plattensis, as stated by Nelson, 
hardly indicates this plant and I become doubtful of its identity. 
Dr. Yuncker writes me that an examination of the type of C. 
plattensis shows my plant to be quite different, although the 
description of the species as given by Nelson agrees. He refers 
my specimens to C. Gronovii var. ctirta Engelm. In a flax field at 
Ft. Ransom, where considerable damage was caused, this form 
and C. Coryli Engelm. were both present, sometimes separate 
and at others tangled together on the same host plant. The 
field was a small one next to the woods of the river. 

Stevens: North Dakota plants 99 


Following the treatment of the western forms by E Nelson 
(Ninth Rept. Wyoming Agr. College, 1899), our plants would 
seem to be separated more satisfactorily as follows: 

Leaves 5-10 mm. long; flowering stems one-flowered. P. Hoodii. 

Leaves 1^2 cm. long; flowering stems two-to several- flowered: more up- 
right, the bark peeling off" in shreds. P. andicola. 

Phlox andicola (Britton) E. Nelson. P. Douglasii of 
Bergman's Flora, in part. This seems to be better separated 
from P. Hoodii Rich, by the above characters than by the size 
of the flowers. The Medora and Washburn specimens referred 
by Bergman to P. Douglasii Hook, are evidently P, Hoodii, 

t Lappula cenchrusoides a. Nels. Marmarth, July 4, 
1918, in sandy soil of river valley. Habit of growth similar to 
Z. occidentalis (Wats.) Greene, nutlets similar to those of L. 


Lapptda (L,) Karst. but larger, the spines longer, and with a row 
of elongated tubercles on the middle of the back of the nutlets. 
The following, previously referred to L. Lappula, also belong 
here: Medora, Bergman 12^6; same locality, June 19, 1910; 
Williston, Bell 25. 


(A. Nels.) Nels. & Macb 

Marmarth, same as preceding. Often three of the nutlets of 
a flower have the spines confluent, forming a spreading border, 
the fourth nutlet with simple spines. In the specimens col- 
lected, however, there seems a decided tendency for the spines 
to be simple on all nutlets of the lower flowers. There are also 
nutlets with spines of intermediate form, broadened but not 
united- Counts on three plants show nutlets as follows: 

Plant No, I — 80 united, 71 simple, 40 intermediate; 

Plant No. 2 — 61 united, 106 simple, 34 intermediate; 

Plant No. 3 — 47 united, 47 simple, 14 intermediate. 
The color of the corolla is an uncertain character, white 
flowers often appearing bluish in the dried specimens. 

t Lappula texana homosperma (A. Nels.) Nels. & Macbn 

Marmarth, same as preceding, occasional plants in patches of 
L, occidentalis, all plants dried up and only fruits collected. 
From a planting of fruits the following season about twenty-five 
plants of L. occidentalis were obtained from fruits of that species. 
No plants were secured from the few fruits of the other two 


Lake Ibsen (Leeds) 

July 4 and i7, 1914, Lunell; Pleasant Lake, J 

loo Stevens: North Dakota plants 



191 8, in loose burned clay around large boulders of the same 
jnaterial. ' 

Amsinkia Menziesii (Lehm.) Nels. & Macbr. A. inter- 
media F. &M. Rugby July 7, I9i7,and July 27, 1918, a number 
of plants along the railroad track near the station; the first col- 
lection In flower, the second in fruit. Bergman 's specimen from 
Pembina is evidently this species also, instead of A, lycopsoides 

t Lamium amplexicaule L. Langdon, July 18, 1918, a 
quantity in a dooryard* 



several plants in a field of Bromtis inermis. Several other 
introduced plants were also in the field: Potentilla argentea L., 
Campamda sibirica L., Chrysanthemum Leucanthemum L. and 
Bromus tectorum L. To be consistent with Bergman's nomen- 
clature this plant should perhaps be referred to Moldavica but 
I have not thought it desirable to make such a change at this 


time. The plant is different in appearance from D, 
Nutt., being slender and having very small flowers. 

t Lycopus communis Bicknell, Pleasant Lake, July 26, 
1912, Lunell', Anselm, Aug. 29, 1920 (common in aJder swamps). 
The Anselm plant has been verified by P. A. Rydberg. 

t Salvia lanceolata Willd. In fields. Carrington, Aug. 
27, 1919; Mandan, Sept. 13, 1920. 

Chamaerhodos erecta (L). Bunge. This was included in 
the seventh edition of Gray's Manual, the record being appar- 
ently based on a specimen collected near Crookston, Minn. 



1901). I had thought that 


gravel pit. It is probably one of the instances of a plains species 
occuring on the eastern edge of the Red River Valley. I have 
not seen the plant east of the hills along the Sheyenne River at 
Sheyenne and Valley City. 


May 26, 1917. One 

tree is in a thicket east of the fair grounds, and quite a number 
occur in a bend of the river two miles farther north. 

t Medicago falcata L. Oakes, July 18, 1919, a plant 
along the railroad tracks. 

Stevens: North Dakota plants ioi 

t Astragalus parviflorus (Pursh) MacM. Marmarth 

July 3, 1918. Q 

t Hedysaruw 


single pod collected. Reported by Arthur (N. Am. Fl. 7: 450. 


t EvoNYMUS atropurpureus Jacq. Owego, Sept. 1916, 

R. A. Shunk. 

t LoMATiUM macrocarpum (Nutt.) Cov. & Rose. 
May 18, 1918, Lunell, and fruits collected at Minot in J 
^eem properly referred to this species. 



i have seen a specimen collected 

by A. H. Shunk along the Sheyenne River near Anselm. 



plant found in a field of Bromus inermis. 


Wallr, A specimen in the 

Gray Herbarium from Leeds, Aug. 21, 1902, Lunell, is referred 
by Millspaugh and Sherff (Field Mus. Nat. Hist. Bot Ser. 4: 


33 • 1 9 1 9) to this species. Some specimens of Xanthium 
collected along the river at Fargo in 1919, to show variations in 
burs, were determined by Sherff as X. italicum Mor. {X. 
canadense of the Flora) and X. acerostim Greene, f with the com- 
ment that the latter was perhaps not distinct. 

t Chrysothamnus nauseosus (Pall.) Britton. Williston, 

Aug. II, 1915, on the hills along the Missouri River about fifteen 
miles southeast of the city. A plant of quite different appear- 
ance from C graveolens (Nutt.) Greene, w^hich is common there 
and on the buttes in the bad lands — the crown low, and with 
gray branches only 1-2 dm. long. This is the plant referred by 
LunelJ (Am. Mid. Nat. 5: 41. 1918) at my suggestion to C. 
formosus Greene, but it evidently is not that species. 

Helianthus giganteus L, The form referred to in Berg- 
man 's Flora under this name is quite common in the central 
(and western?) part of the state, but I am as yet in doubt as 
to its status. It is evidently Lunell's H. nitidus (Am. Mid. 
Nat. 1: 235. 1914). Specimens from Valley City and New 
Rockford were determined by Standley as if. tuberostis. The 
Fargo specimens are -BT. Maximiliani Schrad. 

Helianthus grosse-serratus Martens, The Fargo speci- 

men of the Flora is certainly H. Maximiliani. The stem 
is only slightly hispid above and the leaves are broader and more 
nearly flat than usual. The Kenmare specimen {Bergman 2/44) 
belongs to the preceding form. 

I02 Stevens: North Dakota plants 

t Madia glomerata Hook. Spring Brook, Aug. 17, 1915^ 

a quantity in a prairie slough near the town. 

t Artemisia pabularis (A. Nels.) Rydb. Mandan, Sept. 11, 

1020. Determined by Rydberg. There seems to be no other 
specimen among our material which resembles this. Mr. 
Thysell of the Northern Plains Station showed me the plants, 
only two places where they were growing being known. 

t Senecio manitobensis Greenman. Bottineau, July 8, 
1917. Determined by J. Lunell. In the meadow of a dried up 
pond in the Turtle Mts.; Towner, July 13, 1911, Lunell (speci- 
men In the Gray Herbarium). 

t Centaurea picris Pall. Spring Brook, Aug. 11, 1918. 
Determined by Standley. . Received from Jacob Widman who 
referred to it as abundant in a field. Especially a few years 
ago the akenes were often found in Turkestan alfalfa seed, but 
this is apparently the first record of its having become established. 

Sonchus arvensis L, Ellendale, July 24, 1919. Several 
spots around buildings in the town. 

fSoNCHUS ULIGINOSUS Bieb. S. arvensis oi Bergman's Flora, 
Fargo, Aug. i, 1916; Oakes, July 18, 1919; Crosby, June 7, 1919, 

/. II, Phelps; WiUiston, Aug. 1915. Specimens of this and of 
the preceding were examined by Standley, who stated that no 
North American specimens were in the U. S. National Museum 
collections. A specimen sent to the New York Botanical 
Garden was reported by Rydberg as apparently new to this 
country*. This seems rather strange in view of the fact that it 
is the common form through the Red River Valley, and westward 
in the northern part of North Dakota to somewhat beyond Devil's 
Lake, southern Manitoba, and western Minnesota. It is said 
to have first appeared at-Portage Plains, Manitoba, about 1900. 

This form diflfers from 5. arvensis in the absence of glandular 
hairs on the upper stems and involucres. While I have not 
been able to compare carefully fresh material, I believe the heads 
are smaller, paler, and the rays more inclined to become recurved. 
Vegetative reproduction is by long horizontal, roots and not 
rootstocks as commonly stated. The development of the 
terminal head is often stopped before flowering and several flower- 
ing branches arise just below it. 

* Small has recently listed the species as an addition to the American 

flora, from Pennsylv^ania, where It was collected in 192 1 (see Torreya 21: loO. 

Stevens: North Dakota plants 103 

Lactuca virosa L. Many European authors describe this 
as having horizontal leaves and black, broadly margined akenes. 
This does not apply to our plants. The pinnatifid-leaved form 
is only occasional in the state (Kulm, Brenckle 782 \ Bottineau, 
July 23, 1918; Fargo, Aug. 17, 1920). Pammel's notes (Rhodora 


20: 180-181, 191 8) on the replacement of the entire-leaved form 
by the pinnatifid in Iowa suggest that It will be interesting to 
watch for a similar case here. 

t Crepis occidentalis Nutt. Bowman, June 23, 1918. 
Quite common on Twin Buttes. 

t Crepis capillaris Roth. Fargo, Aug, 26, 1920; Willow 
City, F. M. Rich, 

t Spirodela polyrhiza (L). Schleid. Fargo, Aug. 30, 1919, 

a few plants among masses of Lemna. 

t Cyperus diandrus Torr. Anselm, Aug. 1916, R, A. Shunk. 

t Cyperus esculentus L. Fargo, Oct. 2, 1920. 

SciRPUS VALiDus Vahl. I cannot support Bergman's separa- 

occidentalis (W. 

There do seem to be tw^ 

Manual (ed. 7, /. 

rigid panicles and longer spikelets. The specimens, however, do 
not match Bergman's separation. Both of these forms were 
found in a small patch at Oakes on July 18, 1919, the first 
occupying a definite part of the patch. Specimens of each were 
identified by Agnes Chase as S, validus. 

Carex obtusata Liljebe. Verified by Standley. Fargo, 
June 24, 1920. Quite abundant, at least in one place in low 
prairie. Kensal, Bergman, ^744} is alsothis in stead of C. 
stenophylla Wahlenb. 

The following additional species of Carex were recorded by 
Lunell (Am. Mid. Nat. 3: 234-237. 1916) from specimens deter- 
mined by Mackenzie: C Hookeriana Dewey, C. athrostachya 
Olney, C. tenera Dewey, C. praticola Rydb., C Emoryi Dewey, 
C, scirpiformis Mackenzie, C Parryana Dew^ey, C laeviconica 
Dewey and C atherodes Spreng. 

t Syntherisma sanguinale (L.) Dulac. Fargo. Oct, 2, 

t Panicum perlongum Nash. Verified by Hitchcock. 
Fargo, June 24, 1920, Very abundant In one place in low 

104 Stevens: North Dakota plaxts 



1919: Fargo, Aug. 16, 1920; Steele, Sept. 10, 1920. - 

Festuca viridula Vasey. Lunell (Am. Mid. Nat. 4: 224 

191 7) has reported this species from Dunseith. I have examined 

carefully a specimen received from him ("det. by U. S. Dept. 

Agr.") but can see no reason why it should not be referred to 

F. Hallii (Vasey) Piper. 

Bromus japoxicus Thunb. Fargo, Oct. 1918. Determined 

by Agnes Chase. All of the specimens referred by Bergman to 

B. commutatus Schrad., and several others not reported, seem 

to belong here. 

t LoLiUM RiGiDUM DuTHiEi Hook- Determined by Agnes 

Annual: culms 3 to 4 dm. high, rather stiff, somewhat 
scabrous above; leaves upright, 5 mm. wide, I dm. or more long, 
glabrous or nearly so; glume about three-fourths as long as the 
well developed spikelets, equalling the younger or shorter ones; 
spikelets five- to seven-flowered, the larger 2 cm. long; lemmas 
8 to 10 mm. long bearing an awn 7 to 12 mm. long. 

Described from a specimen collected July 7, 19^9^ grown 
from seeds found in a sample of wheat from the northeastern 
part of the state. An early maturing annual, not so large as 
Zr. temtdentum L. and quite different in the longer, lanceolate, 
awned florets. The culms commonly bear a branch from the 
first node, some of these In the specimen cited being only I dm. 
long with spikes barely protruding. The Milton specimen cited 
by Bergman under L. temulentum belongs here, both it and plants 
from the 1 919 culture having been examined by Mrs- Chase. 
The plant seems well established in that vicinity as evidenced 
by these cases, by material received at one or two other times 
and by seeds found in several samples of wheat. 

t LoLiUM RExMOTUM Schrank. In flax plots, Mandan, 1918. 


Three species of this genus are of great economic importance 
in North Dakota. Having had occasion to pay particular 
attention to these, I find that as a rule, descriptions and figures 
seem to have been made from immature material, causing certain 
characters to be overlooked. The spikelets are well figured by 
Hillman (Bur. PI. Ind., U. S. Dept. Agr. Circ. 73- I9ii)- ^ 
offer the following descriptions: 

Stevens: North Dakota plants 105 

A. repens (L.) Beauv. Splkelets distinctly articulated to 
the rachis (so that they break away readily, with a rounded 
base), and with a strong transverse impression about I mm. 
above the base. Glumes from half as long to nearly equalling 
the spikelet, strongly nerved, with about a dozen short teeth 
on the keel* near the apex which is acute or obtuse, often bearing 
an awn 2-10 mm. long; margins of the glumes thin and translu- 
cent except toward the base where they are indurated, rounded, 
usually slightly separated exteriorly and quite widely interiorly. 

A. Smithii Rydb. Spikelets not articulated (breaking away 
with a rough base) and only slightly impressed. Glumes 
about one-half as long as the spikelets, indistinctly nerved and 
without distinct teeth on the back; margins thin on lower third 
(meeting exteriorly), abruptly narrowed to a long stiff, acumin- 
ate point. 

A, tenerum Vasey. Spikelets not articulated and only 
slightly impressed. Glumes about equalling the spikelets, acute, 
strongly nerved, the nerves all rough with small teeth, margins 
nearly straight. 

t Agropyron dasystachum (Hook.) Scribn. A specimen 
from Bottineau, July 7, 1920, is referred here (''form which has 
been called subvillostim*') by Hitchcock. Bowman, June, 30 
1918, and Kenmare, July 15, 1913, are the same, and I think 
all the specimens previously referred to A. molle (excepting 
perhaps Bell 383) belong here. 

t Dryopteris Thelypteris (L.) a. Gray* Anselm, R, A. 

t Dryopteris spinulosa (Retz.) Kuntze. Another speci- 
men of the same collection as the preceding seems to belong here, 
as does also Walhalla, Bergman 2000. 

Since the above was written I have received from Mr. F. P. 
Metcalf a reprint of an article (Jour. Washington Acad. Sci. 10: 
188-198. 1920) in which he reports eighteen additional species for 
the state. Several of the records suggest the probability of an 
error of indentification or of a difference in interpretation. In 
reply to my inquiry regarding a few of them Dr. Standley 
wrote that he has been able to find but one of the specimens, 
Rumex Britannica L. (McLean County), and that that seemed 
to be correctly determined. 

* This is not actually the central nerve, the broader part of the glume being 
exterior and only one or two nerves on the side next the rachis. 


Three new species of Cuscuta from Mexico 

t. g. yuncker 
(with three text figures) 

In studying collections of Cuscuta from Mexico three species 
have been discovered which, it is beheved, have not been 
hitherto described. Descriptions of these new species, together 
with camera lucida sketches, are given below. With the addi- 
tion of these the number of species of Cuscuta now known to 
occur in Mexico is thirty-six. 

Cuscuta dentatasquamata sp. no v. 

Stems medium to slender. Flowers reddish, 3-4 mm, long, 
on pedicels shorter than or mostly about equal to the flowers, 
in rather compact, cymose clusters; calyx deep, lobes exceeding 
the corolla tube, triangular, acute, slightly overlapping at the 
base, thickened in the median region, forming a small carina 
on most of the lobes; corolla campanulate, lobes triangular, 
acute, upright or spreading, shorter than the tube and with 
slightly uneven margins; stamens shorter than the corolla lobes, 
filaments stoutish and about equal to the oval or orbicular 
anthers; scales about reaching the stamens, dentate, with 

Fig. 1. Cuscuta dentatasquamata Yuncker 
A. Flower, X 5. B. Opened corolla, X 5- C. Individual scale, X 10. 
D. Opened calyx, X 5. E. Capsule, X 5. 

few processes, bridged at about the middle, not firmly adherent 
to the corolla; styles slender, about equal to, or longer than, the 
depressed-globose ovary. Capsule large, depressed-globose, 
thin, somewhat irregularly clrcumscissile, carrying the withered 
corolla about it; seeds about 1.5 mm. long, globose, two to four 
in each capsule, hilum oblong, transverse. 

This species falls in section Eugrammica, subsection Lepti- 
LOBAE, near Cuscuta Choisiana Yuncker. It differs from that 



Yunker: New species of.Cuscuta 

species, however, in the color of the flowers, and in the shape of 
the scales and of the calyx and corolla lobes. It superficially 
resembles Cuscufa partita Choisy, but differs from that species 
in possessing shorter corolla lobes, dentate scales, shorter fila- 
ments and larger flowers- 

The only specimen examined was from Los Pinitos, Sonora, 



Stems rather coarse.. Flo^wers 4-5 mm. long, on pedicels 
about as long as, or shorter than, the flowers, in cymose clusters; 
calyx shorter than the corolla tube, lobes ovate-orbicular, over- 
lapping, somewhat fleshy toward the base, margins very uneven; 
corolla campanulate, lobes much shorter than the tube, erect 
or spreading, overlapping, obtuse or rarely slightly acutish, 
margins uneven ; stamens shorter than the lobes, filaments 
stoutish, about equal to, or slightly longer than, the oval anthers; 
scales not reaching the stamens, oblong, moderately fringed 

■ r - 

Fig. 2. CuscuTA cozumeliensis Yuncker 

A. Flower, X 5. B. Opened corolla, X 5. C. Opened calyx, X 5- D- 
Individual scale, X 10. E. Ovary, X 5. 

with medium length processes, bridged at about a quarter of 
their height; styles slender, much longer than the depressed- 
globose ovary. The only specimen seen of this species is 
fragmentary and not ripe enough to show capsules. The ovary, 
however, gives indications that the capsule would be definitely 
circumscissile and probably carry the withered corolla about it. 


This species falls in section Eugrammica, subsection Ob- 
TUsiLOBAE, near Cuscuta macrocephala Schaffner, from which it 
differs in the stamens not being sessile as they are in that species, 
and in having smaller stigmas and longer processes on the scales. 

Yuncker: New species of Cuscuta 




which it differs in having shorter corolla lobes and shorter scales; 

with Cuscuta corymbosa grandifl 

Engelm., from which it 


ments and with broader scales; and with Ctiscuta iinctoria 
wartius, from which it differs in its larger flowers and pro- 
portionately shorter corolla lobes and shorter scales. 

The specimen examined was from Cozumel Island, Yucatan, 
Mexico (Gaunter go), the type, in the Gray Herbarium. 

Cuscuta durangana sp. nov. 

Stems slender. Flowers membranous, about 2 mm. long, on 
pedicels mostly as long as, or longer than, the flowers, in cymose 
panicles; calyx about as long as the corolla or scarcely reaching 
the sinus; lobes ovate, obtuse or, rarely, acutish, not over- 
lapping; corolla campanulate; lobes ovate, obtuse, spreading, 
later becoming reflexed, about as long as the tube; stamens short- 
er than the lobes, the large, 
oval, versatile anthers about 
equal to the stoutish filaments; 
scales about reaching the sta- 
mens, obovate or spatulate, 
fringed about the upper half, 
bridged low; styles slender and 
longer than the globose ovary. 
Capsule depressed-globose, 





withered corolla about it; ap- 
parently mostly two-seeded; Fig. 3. Cuscuta durangana Vuncker 

A. Flower, X 5 B. Opened corolla, 
X 5 C. Opened calyx, X 5- D- Cap- 
sule, X 5. E. Individual scale^ X 10. 

seeds ovate, slightly rostrate, 
about 1.5 mm. long, hilum 

oblong, oblique. 

This species falsi in section Eugrammica, subsection Obtusi- 
LOBAE, near Cuscuta applanata Engelm. It differs from this 
species, however, in having longer pedicels and a looser inflores- 
cence and in not exhibiting keels on the calyx. It superficially 
resembles Cuscuta umhellata H. B. K. The obtuse corolla and 
calyx lobes easily distinguish this species, however. 

The only specimen examined was from Durango, Mexico 
[Endlich 268), the type, in the herbaruim of the Botanical 
Institute at Dahlem (a small bit in the writer's herbarium). . 

DePauw University 



The aim of this Index is to incline all current botanical literature written by 
Americans, published in "' ' ""'ca, or based upon American material; the word 
America being used in the ►_. t sense. 

Reviews, and papers that relate exclusively to foresty, agriculture, horti- 
culture, manufactured products of vegetable origin, or laboratory methods 
are not included, and no attempt Is made to index the literature of bacteriology. 
An occasional exception is made in favor of some paper appearing in an Ameri- 
can periodical which is devoted wholly to botany. Reprints are not mentioned 
unless they differ from the original in some important particular. If users 
of the Index will call the attention of the editor to errors or omissions, their 
kindness will be appreciated. 

This Index is reprinted monthly on cards, and furnished in this form to 
subscribers at the rate of three cents for each card. Selections of cards are 
not permitted; each subscriber must take all cards published during the term 
of his subscription. Correspondence relating to the card issue should be 
addressed to the Treasurer of the Torrey Botanical Club, 

Acosta> C. Coleccion de maderas cubanas. Revista Agr, Com. 

y Trab. 3: 55, F 1920. 

Acosta, C. El tararaco [Hippeastrum], Revista Agr. Com. y 

Trab. 3: 56. F 1920. [lUust.] 

Acosta, C. Sobre el cayeput [Melaleuca leucadendron L.] 


/. J, 2. N. 1919 

J. M. Estudio de los platanos y guineos cubanos. Re- 


/. 1-28. D 1920. 

Andrews, A. L. Notes on North American Sphagnum. VI- 

Bryologist 18: 1-6. 16 Ja 1915. 

Andrews, A. L. Notes on North American Sphagnum. VII. 

Bryologist 20: 84-89. 13 O 1917. 

Andrews, A. L. Tortula caroliniana, new species. Bryologist 

23: 72-76. pL 5. 2 D 1920. 

From North Carolina. 

Arber, A. Leaves of certain amaryllids. Bot. Gaz. 72: 102- 

105. /. 1-8. 15 Au 1921. 

Arber, A. The leaf structure of the Iridaceae, considered in 

theory. Ann. Bot. 35: 301-336. 

/. 1-66. Jl 


112 Index to American botanical literature 

Ball, C. R. The relation of crop-plant botany to human welfare 

Am. Jour. Bot. 8: 323-338- Jl 1921. 
Ballou, C. H, Los mejores mangos para Cuba. Revista Agr 
. Com. yTrab. 1: 187-191. Ap 1918. [Illust.] 

Bennett, A. Sparganium angustifolium Michx. Jour. Bot. 59 

235, 236. Au 1921. 


Papaya . Bot 

Gaz. 72: 97-101./. 1-6. 15 Au 1921. 

Berry, E. W. A fossil sea bean from Venezuela. Am. Jour. 

Sci. 50: 310-313./. I. O 1920. 

Berry, E. W. Age of the Dakota flora. Am. Jour. Sci. 50: 

387-390. N 1920. 

Berry, E, W. A palm nut from the Miocene of the Canal Zone. 

Proc. U. S. Nat. Mus. 59: 21, 22. /. 1-3. 1921. 
Berry. E.W. Paleobotany: a sketch of the origin and evolution 

of floras. Smithsonian Report 1918: 289-407. pi. 1-6 

+ /. 1-42. 192 1. 

Berry, E. W. Tertiary fossil plants from Costa Rica. Proc. 

U. S. Nat. Mus. 59: 169-185. pi. 22-27. 1921. 

Includes 8 new species. 

Blakeslee, A. F., Welch, D. S., & Cartledge, J. L. Technique 

in contrasting mucors. Bot. Gaz. 72: 162-172. /. 1-2. 
15 S 1921. 

Blasquez, A. La vegetacion las lluvias. Revista Agricola 6: 

333-345- Je 1920. 
Bois, D., & Gerome, J. La chilacayote du Mexique (courge de 

Siam): Cucurbita ficifolia Bouche (C melanosperma Al. 

Braun). Bull. Mus. Hist. Natur. 26: 675-678. 1920. 

Borden, A. D. A biological study of the red jdate-palm scale. 
Phoenicococctis Marlatii. Jour. Agr. Research 2 1 : 659- 
667. pL 127-130. I Au 192 1. 

Botto, A. La sweet tussac, Phalaris hiilbosa Cav, Revista 

Facult. Agron. y Veter. Univ. La Plata IL 12: 226-242. /. 

1-4. 1916; 13: 1-76./. j-p. 1918. 

Britton, N. L- Stewardson Brown. Jour. N. Y. Bot. Gard. 22: 

110-112, Je 1921. 

Bruner, S. C. Enfermedades de la vid [Vitis] en Cuba. Re- 
vista Agr. Com. y Trab. 1: 406-409. Au 1918. [Illust.] 

Index to American botanical literature 113 

Bruner, S. C. La enferraedad del "mosaico" o de '^Rayas 

amarillas" de la cana de azucar en Cuba. Revista Agr. 
Com, y Trab. 2: 437-441- /- i. 2. S 1919. 

Bruner, S. C. La '' Phomopsis'' de, la berenjena. Revista 

Agr, Com. y Trab. 1: 468, 469. S 1918. 

Bruner, S. C. La pudricion negra del cacao, Revista Agr. 

Com. y Trab. 2: 630-636. D 1919, [Illust.] 

Calvino, M. El zacate prodigio {Tripsacum latijolium, Hitch- 
cock). Revista Agr. Com. y Trab. 3: 62-67. F 1920. 


Calvino, M. Estudio sobre el cultivo de la soya [Soja max] 

en Cuba. Revista Agr. Com> y Trab. 3; 124-131, My 


1920. [Illust.] . 

Calvino, M. La lechuga [Lachica sativa]j su cultivo y apro- 
vechamiento. Revista Agr, Com, y Trab. 1: 411-418. 
Au 1918. [Illust.] 

Calvino, M. La luz y la produccion vegetal, Revista Agr. 

Com. y Trab. 3: 532-536. Ap 1921, [Illust.] 

Calvino, M. La jicama de agua (PachyrJdztts tuherosiis) 
Revista Agr, Com. y Trab. 2: 84-87. Mr 1919. [Illust.] 

Calvino, M. Nuevos variedades de cana de azucar. Rev. 
Agr. Com, y Trab. 3: 436-440. Ja 1921. [Illust.] 

Carpenter, C. W. Morphological studies of the Pythium-like 
fungi associated with root rot in Hawaii. Bull. Exp. 
Sta. Hawaiian Sugar PI, Assoc, Bot. 3: 59-65. pL 16-23. 
Au 1921, 

Child, C. M. Le probleme de Tintegration physiologique. 
Scientia 30: 1 15-126. i Au 1921, 

Clute, W. N. Note and comment. Am, Bot. 27: 116-113. 

Au 1921, 

Clute, W. N. The Victoria water lily. Am. Bot. 27: 81-86. 

Au 1921. 
Collins, J. L. Rever^on in composites. Jour, Hered. 12: 129- 

133- /. 16-IQ 10 Au 1921, 
Conard, H. S. Fossil plants and classification. Am. Bot. 27: 

95-101. Au 1921. 
Coulter, J. M., & Land, W. J. G. A homosporous American 

Lepidostrobus. Bot. Gaz. 72: 106-108. 15 Au 1921. 

114 Index to American botanical literature 

Cremata, M. Una excursi6n botanica a Isla de Pinos. Revista 

Agr. com. y Trab. 3: 47-49. F 1920. [Illust.] 

Cruz, F. B. El cultivo del tabaco en Cuba. Revista Agr. Com. 
yTrab. 1: 172-183. Ap 1918. [Illust.] 

Cummings, C. E. Mushrooms. Hobbles 2: 3-17. S 1921. 

Dachnowski, A. P. Peat deposits and their evidence of climatic 

changes. Bot. Gaz. 72: 57-89. /. 1-12. 15 Au 1921. 

Davidson, A. New species from southern California. Bull. S, 

California Acad. Sci. 20: 49-53. Au 192 1 [Illust,] 

• • 

Dusen, P., & Neger, F. W. Uber Xylopodien. Beih. Bot. 

Centralbl. 38: 258-317. /. i-io. 25 My 1921. 

Studies of Brazilian plants. 

Duursma, G. D. Pilocereussen. Floralia 42: 376-378. 17 

Je 192 1. [Illust.] 

Duursma, G. D. Rhipsalideae. Floralia 42: 378, 379. 17 

Je 1921. 

Eames, E. A. An unusual form of Hahenaria clavellata. 
Rhodora 23: 126-127. pL iji. 26 Au 1921. 

Edgerton, C. W., & Tiebout, G. L. The mosaic disease of the 

Irish potato and the use of certified potato seed. Louisiana 
Bull. 181: 1-15. /. i-j. Au 1921. 

Elorduy, S. T. Selecci6n del tabaco. Rev. Agricola 5: 8oi- 

8io. /. i-p. Ap 1921; 6: 14-18. /. 10-12, My 1921. 

Emerson, R. A The genetic relations of plant colors in maize. 

Cornell Univ. Agr. Exp. Sta. Mem. 39: I-156. pi. l-ii. 
Mr 1921. 

Felippone, F. Contribution a la flore bryologique de I'Uruguay. 

Fasc. 3: 40-45. 191 7. [Illust.] 

Includes new species in Didymodon (i), Leptodonlium (l), Fdbronia (1). 

Feustel, H. Anatomic und Biologic der Gymnospermenbl atter. 

Beih. Bot. Centralbl. 38: i'J'7-2^7. 25 My 1921. 

Fortun, G. M. Notas sobre una excursion a " El Retire " [Cuba]. 

Revista Agr. Com. y Trab. 3: 410-413. D 1920. [Illust.] 

Residence of Jose Blain. 

Fortun, G. M., & Bniner, S. C. Investigacioncs sobre la en- 

fermedad del " mosaic© " o "rayas amarillas" dela cana 
de azucar. Revista Agr. Com. y Trab. 3: 441-445. Ja 
192 1. [Illust.] 


Eraser, G. The dwarf trees of Vancouver Island. Gard. Chron. 

70: 102. /. 40. 20 Au 192 1. 

Index to American botanical literature 115 

Fred, E. B. The fixation of atmospheric nitrogen by inoculated 

soybeans. Soil, Sci. 11: 469-472. pL i~j. Je 1921. 

Fred, E. B., Wright, W. H., & Frazier, W. C. Field tests on the 

inoculation of canning peas. Soil Sci. 11: 479-491. pL 
1-3. Je 192 1, 

Frye, T. C. Notes on useful and harmful mosses. Bryologist 

23: 71. 2 D 1920. 

Gagnepain, F. Classification des Eugenia. Bull. Soc. Bot. 

France 64: 94-103. 191 7. 

Gile, P. L., & Carrero, J. O. Assimilation of nitrogen, phospho- 
rus and potassium by com when nutrient salts are confined 
to diflferent roots. Jour. Agr. Research 21: 545-573. /. I. 
15 Jl 1921. 

Girola, C. D. Contribucion al curso de cultivas industriales. 
Revista Facult. Agron. y Veter. Univ. La Plata II. 10: 

83-113- 1913- 
Girola, C. D. El cultivo de la yerba-mate. {Ilex paraguari- 
ensis St. Hil.) Revista Facult. Agron. y Veter. Univ. La 

Plata II. 11: 163-184. /. 1-4. 1915; 12: 196-225. /, 1-6. 

Girola, C. D. Monografia sobre el canamo {Cannabis sativa L.). 

Revista Facult. Agron. y Veter. Univ. La Plata IL 10: 9- 
68. /. 1-13. 1913. 

Girola, C. D* Observaciones sobre el cultivo del henequen. 

Revista Facult. Agron. y Veter. Univ. La Plata 1 1. 11: 
101-151. /. j-jp. 1915. 

Grant, M. Saving the redwoods. Nat. Geog, Mag. 37: 519- 

536. Je 1920. 

Griffiths, D. The production of the Easter lily In northern 

climates. U. S. Dept. Agr, Bull. 962: 1-31. /. 1-14. 12 Au 

Grosovich, J» M. Las principales malezas de los sembrados. 
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Grove, W. B. Species placed by Saccardo in the genus Phoma. 
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Includes 2 new American combinations in Phovtopsis. 

Hardy, M. E. Earth stars. Am. Bot. 27: 86, 87 Au 192 1. 

Harris, J. A. Leaf-tissue production and water content in a 

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ii6 Index to American botanical literature 

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Heim, A. Charakterpflanzen der Halbinsel Niederkalifomien. 
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Hohenkerk, L. S. Botanical identifications of British Guiana 
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HoUick, A. Loco weeds, Nat. Hist. 21: 85-91. F 1921. 

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Index to American botaxIcal literature 117 

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120 Index to American botanical literature 

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New species from Guatemala. 


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122 Index to American botanical literature 

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Vol. 49 

No. 5 




MAY, 1922 

New British and American species of Lobomonas: 

a study in morphogenesis of motile algae 


Tracy E. Hazen 

(with plates 5 AND 6) 

The genus Lobomonas was established in 1899 by Dangeard 
(3) on a single species, L. Francei, found somewhat frequently 
in the vicinity of Poitiers, France, This species had been figured . 
twenty-one years previously by Stein (7, pL 13, /. 77, 18) as a 
form of Chlamydomonas ptilvisculus Ehrenb. Golenkin also 
appears to have had this species In Russia, and to have confused 
it with another genus, for one of his figures (5, /. 19) y described 
as a reduced form of Pteromonas alata (Cohn) Seligo, can hardly 
be anything else than a young cell of L, Francei, Apparently 
the species has not been studied or scarcely even reported others- 
wise, except by Dangeard- In 1902 Chodat (2) transferred to 
the newly founded genus his species Chlamydomonas stellataj 
briefly described six years .before (1); he even appears to have 
been doubtful of the distinctness of his form from Dangeard's 
type, though both species have been accepted by West (8, p. 172) 

and Wille (9, p. 19), who have reproduced the original illustra- 

The genus presents a cell organization almost precisely like 
that of Chlamydomonas, probably its nearest relative, with the 
exception that its outer wall is furnished with variable irregulari- 
ties or protuberances, which in the type species are frequently 
more developed on the posterior part of the cell, while in L. 
stellata the more uniformly triangular lobes are figured as cover- 
ing the wall nearly to the region of insertion of the cilia. It 



124 Hazen: New species of Lobomonas 


is probable that Lobomonas has escaped the notice of collectors 
to some extent because of the minute size of the cells, but doubt- 
less it is actually one of the rarest genera of the Chlamydomonas 
group. The two new species now presented serve to emphasize 
the unity and distinctness of the genus, and also provide certain 
features of division and conjugation not known hitherto, which 
indicate a fundamental parallelism with Chlamydomonas. 

Lobomonas pentagonia sp. no v. 

This species, the first of the genus to be reported in England, 
I believe, was discovered in considerable abundance, together 
w^ith two species of Pteromonas, at Ham Common, Surrey, near 
Kew, in 1920. The habitat was the border of a small shallow 
pond at the west end of the common, a place frequented by cows 
and horses, and therefore supplied with water containing a 
considerable amount of nitrogenous organic matter in solution: 
in fact the general conditions were very like those of the much 
smaller pool where Lobomonas rostrat-a, to be described presently, 
was found with another species of Pteromonas in New Jersey. 

Like all other species of the genus, L. pentagonia is very 
minute, but it is more constant in form than L. Francei or L. 
rostrata. When seen in side view the cell appears rather pent- 
agonal in outline (Figs. 1-4), but careful focussing shows that 
the angles do not all lie in the same plane. Polar views often 
show five to eight protuberances (Fig. 14), likewise not all at 
the same level. One of the most symmetrical specimens, when 
resting with its ciliated end turned upward, shows four anterior 
protuberances and, alternating with them, four others at the 
posterior end of the zoospore (Fig. 5) ; the number, however. Is 
variable. The protoplast, including the chromatophore, in 
young individuals usually extends into and fills the protuber- 
ances, and ends anteriorly in a somewhat obtuse beak, to which 
the two ciha are attached (Fig. i). In older specimens the pro- 
toplast usually retracts from one or more of the wall protuberances 
and leaves them empty; they do not appear like dense gelatinous 
structures. The chromatophore is more or less hollowed out 
in the common Chlamydomonas fashion but thickened in the 
region w^here the pyrenoid lies in a somewhat lateral position. 
The two alternately pulsating vacuoles lie in a plane nearly 
perpendicular to that passing through the two extended cilia 
(Fig. 5), as in most species of Chlamydomonas, so that in the 
ordinary face view of the cell only one is usually seen. The 

Hazen: New species of Lobomonas 125 

narrow rod-shaped red eye-spot lies in or just beneath the plasma 
membrane slightly in front of the middle of the cell. Its position 
with relation to the cilia is more like that which it occupies in 
Brachiomonas than that in Chlamydomonas , as brought out 
recently by the writer (6), though the dorsiventral differentiation 
here is perhaps less constant and definite than in the two related 

Asexual reproduction. Dangeard (3) reports that his 
attempts to cultivate L. Frdncei in a moist chamber did 
not succeed, and that it was difficult to obtain the multipli- 
cation of the organism. He shows that the cells come to rest 
and generally become rounded in shape, then divide into four 
or eight daughter cells which escape as zoospores. He gives 
no figure to support Wille's (8) surmise that division is longitu- 
dinal. In our two new species I found that very generally after 
motile cells were mounted in a hanging drop they would for the 
most part come to rest in a few hours and proceed to divide, 
though in many cases the daughter cells failed to become motile 
or escape. In both species pyrenoids were not seen in any 
dividing specimens until the daughter cells had begun to take 
on the typical form, so that it would appear necessary to con- 
clude that in this genus, as in Brachiomonas and in some species 
of Chlamydomonas ^ the pyrenoid disappears before the first cleav- 
age, and that one is formed de novo in each daughter cell. 

In L, pentagonia the first division plane usually appears at 
first sight to be transverse to the longitudinal axis of the cell, 
but several cases were observed which lead to the conviction 
that there is regularly a rotation of the protoplast during or 
before the beginning of cleavage. Figs. 6 and 7 show two stages 
of division where the protoplast has revolved to an oblique 
position when the cleavage begins; probably the rotation was 
delayed more than usual in this case; here the original contrac- 
tile vacuoles persist after the division of the nucleus. Fig. 8 
shows a case where what I am sure the original vacuoles persisted 
until the end of cleavage into two daughter cells. In case of the 
formation of four zoospores the second plane of cleavage is per- 
pendicular to the first, i. e. it lies in a longitudinal axis of the 
mother cell: it may lie in a single plane through the two halves 
(Fig. ii), or the second division plane in one half cell may be 
perpendicular to that in the other (Fig, 12). 

126 Hazex: New species of Lobomonas 

Sexual reproduction apparently has not been observed hith- 
erto in this genus. I considered it good fortune, therefore, to dis- 
cover three or four cases of conjugation taking place almost 


simultaneously in one hanging drop. The gametes have the 
general form and character of the vegetative cells, but are much 
smaller, provided with an excessively delicate cell wall, and the 
cilia are longer than the cell body. In the first case seen, one 
gamete was rather broad and pillow-shaped, with a distinctly 
visible cell wall, while the other was narrower and probably more 
nearly cylindrical, and furnished with a wall so delicate that it 
was detected only in its subsequent behavior. The two gametes 
became engaged by. their cilia, but not otherwise in contact, 
and remained in this position for at least a half hour, with slight 
dancing movement (Fig. 15); the ciUa, most if not all of this 
time, were trailed backward along the sides of the narrower 
gamete. The first movement toward joining was in a sudden 
break of the anterior papilla of the broad gamete, whereby a 
broader papilla of colorless cytoplasm surged forward and pres- 
ently plastered itself on the beak of the unchanged gamete (Fig. 
16), the remainder of the protoplast of the broader gamete then 
distinctly withdrawing from the posterior part of its wall (Fig. 
17). Now for a period of about a half hour the remainder of 
the protoplast of the broader gamete was gradually oozing out 
of its wall and into the narrower gamete: even yet the narrower 
gamete hardly showed a wall, but it seemed evident that one 
must be present, since the posterior part of this gamete rigidly 
retained its original form (Figs. 19, 20). After this point more 
active ciliary movement carried the zygote beyond possibility 
of observation. 

A second case showed a similar figure of the narrow gamete 
remaining rigid for as long a time as it could be followed. In 
all probability this wall of the narrow gamete is finally thrown 
off separately, permitting the rounding up of the plasmatic mass 
to form a spherical zygospore. A third case showed the gametes 
more nearly equal, and It was clear that the walls of both were 
practically alike in character and not easily abandoned (Fig. 21), 
so that fusion was long delayed. 

There is in this species apparently little differentiation be- 
tween the gametes ; indeed one might be inclined to regard the 
difference in size as merely accidental. Nevertheless from the 
behavior of the cilia, that is, both pairs for the most of the time 
stretching back alongside the narrower gamete, one may assume 

Hazen: New species of Lobomoxas 127 

a certain degree of differentiation. The conjugation of this species 

presents a close parallel to that described by Goroschankin for 

Chlamydomonas reticulata (5, p. 126, pL 3,j, 6-8). He states 

that the gametes show Httle difference in size, and that they 

sometimes simultaneously slip out of their walls; but more 

frequently, after the beginning of conjugation, one of the gametes 

first throws off its wall and takes on a globose form, then the 

second does likewise, and then the two rounded masses go on to 

complete fusion. 

The subjoined Latin diagnosis presents the chief character- 

istics of this species: 

Lobomonas pentagonia sp. nov. X. cellulis vegetativis minu- 
tis, membrana a latere aspectata forma aliquanto pentagonia 
sed anguHs rotundatis (verrucis) haud omnibus in eodem piano 
a vertice aspectata rotundata cum 5-8 Vermel's, aliisanterioribus, 
aliis posterioribus protoplasto vel membranae conformali vel 
plus minus contracto et ellipsoideo, cum rostello conico ad quod 
cilia bina cellulae longitudinem fere adequantia affiguntur; 

chromatophoro excavato, pyrenoidem unum sublateralem port- 
ante, et in parte excavata nucleum lateralem includente; 
stigmate bacilliformi paululum ante mediam cellulam sito; 
vacuolis contractilibus binis in rostello cytoplasmatlco positis. 

Fropagatio fit 2 aut 4 zsooporis intra cellulae matricalis 
membranam ortis, divisione priore visa quasi transversaria 
propter protoplast! rotationem sed vero longitudinali. 

Generatiofitgametis parvulis, vel subaequalibus vel aliquan- 
to disparibus, membrana tenuissima vestitis, inter se binatim 

Longit. cell. veg. 10-13 V-'y l^t. 9-10 ^. Longit. gametarum 
ca. 8 ^\ lat. 4-5 yi; longit. ciliorum ca. 13 y-. 

Hab. in stagni margine. Ham Common, Surrey, England, 
1-7 Aug. 1920. 

Lobomonas rostrata sp. nov. 

This form, the first representative of the genus to be reported 
in America, at first sight appeared very similar to L. Francei 
Dangeard (3, p. 115), but careful study disclosed differences as 
important as those which distinguish most species of Chlamy- 
domonaSj so that I feel obliged to regard it as a new species. I 
first found a few individuals in examining a collection of Gonium 
pectorale and an undetermined species of Chlamydomonas y ob- 
tained the last of September, 1919, from a rain-water pool of 
a highway in the southern part of Englewood, New Jersey. Lat- 
er it was interesting to discover that the species had been collect- 

128 Hazen: New species of Lobomonas 

ed about a week earlier in a much deeper pool, about half a mile 
distant from that just mentioned; this discovery was due to the 
fact that a fine colony of the Lobomonas developed on an agar 
plate containing a sample of my first collection of Pkromonas 
from this pool. The new species continued to appear sporadically 
in later gatherings from the same pool, the last being made 
November 12; it was also collected in October, together with 
Chlamydomonas metastigma Stein, from another rain-water pool 
in a wheel-rut, not far from the one first mentioned, though 
separated from it by railroad tracks bordered by a deep ditch 
on either side. The first mentioned wheel-rut yielded a few 
individuals in the following season. A few specimens of this 
species were also discovered during the past summer in a col- 
lection from a similar wheel-rut in northern Vermont: here it 
was accompanied by Gonium pectorale, Pandorina, Chlorogonium, 
Chlamydomonas, and a very interesting new form of Polyhle- 
pharideSj to be described in a forthcoming paper. The Lobomo- 
nas never appeared to be abundant like its associates in the 
same pools; usually not more than a dozen or two specimens 
turned up in one hanging drop mount. 

The vegetative cells or zoospores of this species most common- 
ly have a somewhat obpyriform shape (Fig. 22~2'j) though they 
are sometimes almost ellipsoid. In younger individuals the 
cell wall is so delicate and close-lying as to be indistinguishable 
for the most part, but in older cells it is well developed (Figs. 
28, 29) ; it is generally produced into a variable number of lobes, of 
which from five to seven or sometimes as many as ten appear 
in a face view; that these lobes are developed on all sides of the 
cell is clearly shown in a polar view (Fig. 30). At the anterior, 
usually broader end, the wall is extended into a truncate, wedge- 
shaped beak, or possibly more typically this takes the form of 
a more or less double papilla (Fig. 33) ; on account of the minute 
size of the organism it is often most difficult to see clearly the 
exact structure of this protuberance, which is one of the most 
characteristic features distinguishing this species from the two 
hitherto described in Europe. Sometimes the wall appears to 
be uniformly thin, sometimes thickened at the end of the lobes, 
and sometimes considerably thickened throughout. The proto- 
plast, indistinguishable outwardly from the bright green chro- 
matophore, fills the lobes in young individuals; in older cells it 
retracts more or less, so as to leave some or all of the lobes color- 
less. In such cases it is not easy to determine whether the 

Hazen: New species of Lobomonas 129 

lobe IS a dense, gelatinous structure, or simply membranous and 
separated by a space from the protoplast. In certain cases the 
latter interpretation is clearly indicated, for the. tip of the lobe 
is manifestly of thickened grayish wall substance, with a clear 
space inside (FiG. 37). The cilia are attached at a single point 
to the anterior end of the protoplast, which is usually obtuse, 
though it may have a slight beak; they immediately diverge 
at a wide angle to pass separately through the papilla of the wall, 
and are often seen in quiescent individuals stretching out stiffly 
in the form of a v; their length is as variable as the cell outline, 
often being less than the cell length, but perhaps more character- 
istically distinctly greater than the cell length. At the base of 
the cilia are the alternately pulsating vacuoles, lying regularly 
in such a position that a line passing through the two is perpen- 
dicular to the plane in which the quiescent cilia lie, so that only 
one of the vacuoles is seen when both cilia are equally clear, but 
both may appear at the same level when one of the cilia is behind 
the other (Fig. 29), The single pyrenoid occupies a lateral position 
in the deeply hollowed out chroma tophore (Figs, 24, 26), contrast- 
ing sharply with the axial pyrenoid in a massive ch roma tophore 
described and figured by Dangeard (3) in Lobomonas Francei. 
The small rod-shaped red eye-spot lies in front of the middle 
of the cell, but apparently not in a constant position with ref- 
erence to planes passing through the cilia and contractile vacu- 
oles, as is the rule in Pteromonas and in many species of Chlamy- 



Reproduction. The earliest stages of division found pre- 
sented the appearance of a cleavage transverse to the longi- 
tudinal axis of the cell. More careful consideration, however, 



vacuoles, lying in what appears to be the original colorless an- 
terior cytoplasm, now appear on the side of the cell. Even In 
such a case as that shown in Fig. 37, there is a colorless central 
region which can only be explained on the supposition that the 
anterior end of the protoplast had revolved ninety degrees from 
its original position, here not clockwise as in FiG. 36, but in a 
vertical plane with reference to the observer. The mother cell 
retains very much of its original form throughout the process of 
division, instead of rounding up, which Dangeard describes as 
being the general rule for L. Francei. Here also there may be 
four or eight daughter cells formed, and they regularly show the 

130 Hazen: New species of Lobomonas 

typical obpyriform and lobed shape, and sometimes even show 
the protoplast somewhat separated from the new cell wall, before 
escaping from the mother cell. The escape appears to be accom- 
plished by a gradual softening and disintegration of the wall of the 
mother cell, rather than by rupture at a single point (Fig, 40). 
The eight daughter cells of this figure are so small as to occasion 
the surmise that they might be gametes, but conjugation was 

not seen in this species. 

The chief characteristics distinguishing this species from 

L. Francei Dang, are: the general obpyriform shape, the well 

developed anterior beak or papilla, the lateral pyrenoid, and the 

persistence of the form of the mother cell during division. The 

description may be summarized as follows: 

Lobomonas rostrata sp. nov, L. cellulis vegetatlvis plus 
mlnusve obpiriformibus, rarius ellipsoideis; membrana cellulae 
in verrucas plures quarum 5-7 vel etiam 10 in facie una apparent 
producta, atque in polo anteriore rostello cuneiformi sen papilla 
subduplici instructa, per quod rostellum procurrunt cilia bina 
longitudine corpus cellulae adaequantia vel longiora, vel bre- 
viora : protoplasto primum membranae verrucas complente, 
deinde plus minus contracto et ellipsoideo: chroma tophoro valde 
excavato, pyrenoidem unum lateralem portante: stigma te bacil- 
liformi, paullulum ante mediam cellulam site: nucleo majore 
nunc paene centralis nunc laterali, rarius posteriori; vacuolis 
contractilibus binis juxta papillam anteriorem suppositis. 

Propagatio fit protoplasto cellulae vegetatlvae jam immobilis 
diviso in 4 aut 8 zoosporas, quae formam typicam priusquam 
evaderunt ex cellulae matricalis membrana adipiscuntur. 
Copula tio haud observata. 

Longit, cell. veg. 5-12 ^; lat. 4-8 [l; long, ciliorum ca. 5~I4 ^• 

Hab. in aqua pluviali quae coUigitur In viis terrenis, et in 
lacuna quadem lutulenta In pascuo pecuario. Englewood, New 
Jersey, Sept.-Nov. 1919, Sept. 1920: Shelburne, Vermont, 3 
Aug. i92i. 

It has been already pointed out elsewhere (6) that Lobomonas 
is to be regarded as a special offshoot from Chlamydomonas, 
not leading to any higher group so far as we know at present. 
It might be thought simple to derive the genus directly from 
the Polyblepharidaceae, even from the genus Dtmaliella, which 
clearly appears to be the immediate ancestor of Chlamydomonas, 
since it has all the features of cell organization of the latter 
genus except for the lack of a firm cell wall. When, however, 
it is recalled that the gametes of Lobomonas, reported above for 
the first time, possess cell walls, ft will be more natural to look for 

Hazen: New species of Lobomoxas 131 

its ancestry among those species of Chlamydomonas which posses 
walled gametes, and are therefore ranked as the primitive mem- 
bers of the genus, since their vegetative cells and gametes 
differ (visibly) only in point of size. Our two new species of 
Lobomonas, moreover, also resemble a number of the relatively 
primitive members of Chlamydomonas in their method of cell 
division, namely through a cleavage which is fundamentally 
longitudinal but early shifts to a transverse position. The 
question then arises, what influences led to the divergence of 
cell form, which is practically the sole basis of separation between 
the species forming the small genus Lobomonas and the much 
larger number comprised in Chlamydomonas. 

I ^ 

A consideration of morphogenesis in primitive algae 

This whole problem of the origin and inheritance of cell 
form in primitive organisms is one of very great interest which 
has received comparatively little attention. Perhaps the most 
extended discussion of the question has been furnished by D'Arcy 
Thompson (22), who regards surface tension as the paramount 
factor in the determination of cell form. Though at one point' 
he admits that 'the physical cause of the localized inequalities 
of surface tension remains unknown', and at other times hints 
that an internal chemical heterogeneity may have some influence 
in connection with such difi^erences in surface tension, never- 
theless again and again he reiterates his main thesis in regard 
to unicellular organisms, 'that not only their general config- 
uration but also their departures from symmetry may be correlated 
with the molecular forces manifested in their fluid or semi-fluid 
surfaces'. This explanation seems to us entirely inadequate 
and not in harmony with the general weight of evidence. For 
the particular organisms considered here and in the previous 
paper on Brachiomonas (6) we can offer little direct evidence; 
but there are certain facts which suggest that the conception of 
the non-homogeneity of the protoplasmic structure of the cell, 
as developed by Rhumbler (19) and Harper (12) supplies a 
much more workable hypothesis than the idea of mere surface 

tension forces. 

Passing over Rhumbler 's work on protozoa, the most 

thorough treatment of morphogenesis within a small group 

of primitive plants is found in the studies of Harper (11, 12) 

on Pediastriim. He believes that the general four-lobed form 

132 Hazen: New species of Lobomonas 

of the cells familiar in most species of Pediaslrum may well 
have arisen in evolution as a consequence of the pressure and 
contact relations of the young cells in the sixteen-celled colony, 
regarding them merely as surface tension globules: nevertheless 
he has shown repeatedly that this four-lobed form does not de- 
pend in ontogeny upon the forces which may have been respon- 
sible for it originally, but that it is inherited and may reach 
full expression when there is the least possible contact with other 
cells of the colony. Repeatedly Harper calls attention to the 


view that though surface tension is commonly acting as a 
morphogenetic factor, nevertheless 'it is the inherited anomo- 
genous consistency of the cells which is of most significance in 
determining their form.' In strictly unicellular organisms like 
Lobomonas and Brachiomonas, there is an absence of the inter- 
action of contact and pressure stimuli which are important 
influences in the variation of the Pediastrum colony; never- 
theless in the fundamental organization of the cell the factors 
must be parallel in the main. In both cases the lobed form may 
be regarded as adaptive for the general metabolism of the cell. 
In Pediastrum the development of spines is usually looked upon 
as a case of primitive differentiation for protection, and in 
Lobomonas and Brachiomonas the lobes might easily be conceived 
of as subserving a similar function: in point of fact, I have ob- 
served that when these forms are found in the same pools with 
smooth-walled ovoid Chlamydomonas cells, it is the latter that 
are devoured by protozoa, while they rarely or never prey upon 
the lobed forms, even though the latter are smaller. However, 
when two species of Chlamydomonas are present together, some- 
times one is rapidly consumed by protozoa while the other is 
ignored. The anti-selectionist, moreover, might fairly inquire 
why it is that these genera of bizarre form have produced very 
few species, in comparison with the extraordinarily successful 
genus Chlamydomonas, which has attained well nigh three score 



characteristic of Meutliche Metabolle.' I cannot find that 
Dangeard uses this term in connection with this genus; but as 
defined by him elsewhere* as amoeboid movement, or used 

*0n designe sous le nom de metabolie une sorte de mouvement par con- 
traction du corps partlculier k quelques Euglenes, Amibes, Monades, etc. 
C*est ce que nous appelons mouvement amiboide. [Dangeard: Recherches 
sur les algues inferieures. . Ann. Sci. Nat, Bot. VIL 7: 144, 1888.] 

Hazen; New species of Lobomonas 133 

more exactly by other recent writers (especially in connection 
with the Polyblepharidaceae) in the sense of euglenoid or amoe- 
boid change of form — aiid this was the usage of Perty*, by whom 
the word appears to have been coined in 1852 — the term Is mis- 
leading when applied to Lobomonas: or at least it can be used only 
in a very restricted sense in connection with this genus. The 
young cells of Lobomonas of course do undergo a certain change 
of outline during the formative process within the wall of the 
mother cell : ordinarily essentially the mature form is acquired 
and fixed before their escape, and I have found no evidence that 
it is appreciably altered afterward during activity, though, in 
appearance only, the mature cell may be distinctly reminiscent 
of Amoeba. Nevertheless, this idea of amoeboid change of 
form is most suggestive in a discussion of the formative period 
of such genera as we have under consideration, and for this re- 
stricted period I believe we are entirely justified in drawing a 
parallel with the results of certain recent researches on the 
production of pseudopodia. 

A brief survey of this work may be useful in this connection. 
McClendon (16) has attempted to explain amoeboid movement 
as due to local increase in permeability, the Amoeba simply re- 
ceding from the side on which the permeability has been in- 
creased. Its author himself admits the difficulty of explaining 
positive reactions by this theory, and it certainly does not offer 
any sufficient explanation for the assumption of the character- 
istic cell-form shown by our developing chlamydomonads. The 
closely related hypothesis that production of pseudopodia may 
be accounted for by local variation in surface tension has been 
widely invoked. The re-statement of this theory lately made 
by Thompson (22) may here be passed over, even as this stim- 
ulating writer has all but ignored the evidence which has been 
accumulating against the surface tension explanation. Both 
the permeability idea and the surface tension explanation as 
ordinarily employed are objectionable, in that they depend too 

*MetaboHe. Hirunter verstehe ich die durch innere Vorgdnge bedingte 
wechselnde Gestaltanderung. Seit langem bei den Amiben bekannt ist sie 
bei den Infusorien so viel als nicht beachtet worden. [Perty: Zur Kenntniss 
kleinster Lebensformen, p. 127. Berne. 1852.] 

Since this term MetaboUe seems to be unfamiliar to American botanists 
because of its very restricted usage, and since it has even been rendered as 
equivalent to metabolism in a recent German-English dictionary for chemists, 
it appears to be worth while to call attention to this original definition. 

134 Hazen: New species of Lobomonas 

much upon external environmental influences to account for 
the requisite local variability, though this element is not neces- 
sarily involved in either theory. 

The beginning of the more recent undermining of the surface 
tension explanation of form change is found in the work of 
Jennings (14), who from painstaking direct observation reported 
that the currents in a moving Amoeba as a whole 'are not similar 
to those of a drop of inorganic fluid that is moving or elongating 
as a result of a local increase or decrease in surface tension', and 
in particular that *the movements of material in a forming 
pseudopodium are not like those in a projection which is pro- 
duced in a drop of inorganic fluid as a result of a local decrease in 

surface tension'. 


to the conclusion 'that changes in the surface tension 'of the 
body are not the primary factors in the movements and reac- 
tions of Amoeba'. Similar conclusions from direct observation 
Were announced tw^o years later by Dellinger (10), whose clever 
photographs of Amoeba viewed from the side reveal the locomo- 
tion of the organism as a sort of 'walking,' rather than a flowing of 
a fluid substance upon the substratum. These negatory con- 
clusions have been reinforced and extended by a series of recent 
researches in the field of microdissection (15, 21, 13). From 
these there is general agreement that the structure of Amoeba 
(and probably of many other rhizopods and cells of primitive 
organization) is not of the nature of a simple fluid mass governed 
chiefly by surface tension forces, but rather that it is a high^ 
non-homogeneous system, consisting of comparatively fluid 
endoplasm surrounded by an ectoplasm w^hich has often the 
character of a semi-rigid gel, possessing a considerable elasticity, 
though the two regions may very probably grade into one an- 
other imperceptibly. 

Finally, the experiments of Hyman (13) reveal in each 
pseudopodium of Amoeba a gradient in susceptibility to potas- 
sium cyanide, the susceptibility being greatest at the distal end 
and decreasing proximally. This susceptibility gradient is re- 
garded as being a metabolic gradient which arises before the 
pseudopodium appears, 'and hence the metabolic change which 
produces increased susceptibility is the primary cause of pseud- 
opodium formation.' Liquefaction or solation is regarded as 
the cause of the extension of a pseudopodium, and coagulation 
or gelation as the cause of its retraction: the liquefaction is be- 
lieved to be brought about by the metabolic change just re- 

Hazen: New species of Lobomoxas 135 

ferred to. This theory that amoeboid movemnt is due to alter- 
ations of the colloidal state is only in the nature of confirm- 
ation and extension of the view advanced more than forty 
years ago by Montgomery (17, 18) that protoplasmic movement 
of amoeboid organisms consists in *an alternate expansion and 
contraction of organic substance': Montgomery even anticipated 
Hyman in expressing the idea that the liquefaction which occas- 
ions pseudopodium advancement is itself due to metabolic changes, 
while he anticipated Rhumbler (19) in the idea of a non-homo- 
geneity of the primitive protoplasmic mass which permits various 

functions to be carried on in different regions at the same 

This somewhat lengthy excursus (which is yet only the 
briefest possible summary of a voluminous literature) has been 
introduced here only to bring before botanists, to whom the field 
may be unfamiliar, facts which it is believed may be directly 
applied in the case of the chlamydomonads we are considering. 
The form development of Lohomonas, Brachiomonas, and Ptero- 
monas must be essentially amoeboid for a brief period during 
the organization of the daughter cells, and we are justified 
in assuming that their lobes and excrescences are the expression 
of the same non-homogeneous organization of the protoplast 

as is characteristic of Amoeba. 

For this view, furthermore, we may draw an additional 
parallel from the results of microdissection. In the developing 
oogonium of FucuSy Seifriz reports (21) that the viscosity of the 
protoplasm changes from liquid consistency in the young uni- 
nucleate stage to slightly viscous consistency when the division 
into eight eggs is just complete, and to decidedly viscous consis- 
tency (just under the viscosity of glycerine) in the mature nor- 
mally discharged egg: that is, in Seifriz 's scale of ten grades of 
viscosity — the first attempt on the part of microdissectionists 
at standardization in this matter — the variation is from grade 
3 to grade 6. Yet further, from the behavior of disintegrating 
eggs of Fucus, Seifriz (20) finds that the process may be localized 
in such a manner as to indicate 'a gross structure of the egg 
plasm, f e., the protoplasm is composed of many centers of activity 
in which different chemical reactions take place/ I liave recently 
found a condition almost precisely similar in the case of a newly 
discovered polyblepharid genus which it is hoped may soon be 
published. The cells of this species, though surrounded only 
by an exceedingly delicate protoplasmic membrane, are never- 

136 Hazen: New species of Lobomonas 

theless able to' maintain for extended periods an elaborately 
eight-ridged or winged prismatic form, and in disintegration 
often break only at one or two points, thus permitting the greater 
body of the protoplasm to remain practically intact, while only 
small streams ooze out. From these observations we are justified 
in concluding that such cells as those of Brachiomonas and Lo- 
homonaSy could we apply methods of microdissection, would be 
found to have protoplasm of a comparatively fluid consistency 
during division, but that local increase in viscosity gradually 
permits the fixation of the characteristic lobed form of the cell. 
When it is recalled that in these two genera the pyrenoids regular- 
ly disappear in preparation for cell division, and are reorganized 
with the maturing of the daughter cells, it will readily be seen 
that in this reorganization, combined with the ordinary processes 
involved in division of cells inheriting differentiation of polarity 
in at least three axes or planes, there is abundant room for the 
play of such metabolic changes as might well account for con- 
siderable differences in viscosity in different parts of the develop- 
ing daughter protoplasts. It is this non-homogeneity of struct- 
ure, involving very likely chemical as well as physical differences, 
which may be regarded as the dominant factor in the determina- 
tion of form in such organisms. From this standpoint, the prob- 
lem as to how this characteristic form may be transmitted in 
heredity does not seem so insoluble as it does on the assumption 
of form determining unit factors in a germ plasm. 

One further point may be emphasized. It is stated by 
Thompson (22) that when 'owing to some heterogeneity of the 
substance' the operation of uniform surface tension forces is 
modified so as to result In the production of the ellipsoid cell 
characteristic of yeast, for example, 'this or any other asym- 
metrical form, once acquired, may be retained by virtue of the 
solidification and consequent rigidity of the membranous wall 
of the cell.* In the case of the organisms with which we are 
here concerned at least, I am confident, the development of the 
cell wall is not a necessary condition of the maintenance of 
specific form; for the new polyblepharid mentioned above, and 
others of the same group, as well as the gametes of Brachiomonas, 
are able to preserv^e essentially the same form for long periods, 
in spite of the fact that they are clothed only with a protoplasmic 
membrane of such excessive thinness that it is practically unde- 
monstrable. It is then, owing to relative viscosity in their 

Hazen: New species of Lobomonas 


protoplasmic substance that these motile organisms are able 
to attain and maintain their specific form. 

In both Brachiomonas and Lobomonas we have noted that 
not infrequently mature cells exhibit a more or less rounded form 
of the protoplast inside of the cell wall, while the latter main- 
tains the typical lobed form as fixed in the formative period. 
Such conditions are not improbably produced by shrinkage due 
to loss of a certan amount of water from the protoplast, accom- 
panied by a tendency to decrease in viscosity of the outer layer, 
in which case, surface tension would tend to bring about a more 
rounded form. In other words, the maturing protoplast might 
be said to show a tendency to revert to what might be considered 
the ancestral form in preparation for reproduction. 

It is a pleasure to acknowledge my debt to Professor R. A. 
Harper for numerous stimulating discussions of problems con- 
nected with morphogensis, and for reading this paper in manu- 

Barnard College, 
CoLUMBL\ University 



138 Hazen: New species of Lobomonas 

Literature cited 

A. In connection with Lobomonas 
l.Chodat, R. Mat^riaux pour servir a Thistolre des Protococ- 

coidees V. Bull. Herb. Boiss. 4: 273-280. 1896. 

2. ^Algues vertes de la Suisse. Berne. 1902. 

3. Dangeard, P. A. Memoire sur les Chlamydomonac 

rhistoire d'une cellule. 


4. Golenkin, M. Pteromonas alata Cohn. Bull. Soc, Imp. 

pL II. 1891. 


5. Goroschankin, I. N. 


logic und Systematik der Chlamydomonaden, Bull. 
Soc. Imp. Nat. Moscou 5: 101-142. pi, 1-3. 1891, 

6. Hazen, T. E. The phylogeny of the genus Brachiomonas . 

Bull. Torrey Club 49: 75-92. pi. 3,4^ 1922. 

7. Stein, F. Der Organismus der Infusionsthiere 3^ Leipzig. 


8. West, G. S. Cambridge Botanical Handbooks. Algae L 

Cambridge. 19 16. 

9. Wille, N. Chlorophyceae. Engler & PrantI, Nat. Pflanz- 

enfam 1^, Nachtr. 1911. 

B. On the problem of morphogenesis 

10, Dellinger, O. P- Locomotion of Amoebae and allied forms. 

Jour. Exp. Zool. 3: 337-358. pi i, 2 + f, 1-27. 1906. 

11. Harper, R. A. The evolution of cell types and contact and 

pressure responses in Pediastrum. Mem. Torrey Club 
17: 21Q-240. /. i'2y. 1918. 

12. Organization, reproduction, and inheritance in Pedta- 

strum. Proc. Am. Phil. Soc. 57: 375-439. pl- 3* ^ + /• 
I-3S. 1918. 

13. Hyman, L* H. Metabolic gradients in Amoeba and their 

relation to the mechanism of amoeboid movement. 
Jour. Exp. Zool. 24: 55-99. /. 1-14^ ^9^7- 

14. Jennings, H. S: Contributions to the study of the behavior 

of lower organisms. The movements and reactions of 
Amoeba. Carnegie Inst. Washington PubL 16: 13 ^^ 
234. /. 30''7S^ 1904- 

15. Kite, G. L. Studies on the physical properties of proto- 

plasm. I. The physical properties of protoplasm of 

Hazen: New species of Lobomonas 139 

certain animal and plant cells. Am. Jour. Physiol. 32: 
T46-164. 1913. 

16. McClendon, J, F. Ameboid motion and tropisms. Physi- 

cal chemistry of vital phenomena, pp. 148-156. Prince- 
ton. 191 7. 

17. Montogomery, E. The elementary functions and the 

primitive organization of protoplasm. St. Thorn. Hosp. 
Rep. 9: 75-100. 1879. 

18. Zur Lehre von der IMuskelcontraktion. 11, Dieamo- 

boide Bewegung. Arch. Gesam. Physiol. 25; 499-509. 
pig. 1881. 

19. Rhumbler, L. Der Aggregatzustand und die physikalischen 

Besonderheiten des lebenden Zellinhaltes. Zeits. Allg. 
Physiol. 1: 279-388. 1902, 

20. Seifriz, W. Observations on the structure of protoplasm 

by aid of microdissection. 


21. Viscosity values of protoplasm as determined by micro- 
dissection. Bot. Gaz. 70: 360-386. 1920. 

22, Thompson, D'Arcy W. On growth and form, Cambridge. 


Explanation of plates 5 and 6 

Figures drawn with camera lucida from living materia! kept in hanging 
drops (VanTieghem cells of standard height of 5 mm.):Leitz compens. oc. 
12 was used in combination with oil immers. obj. 1-12 inch, or Spencer 2 mm. 
The drawings have been reduced one half in reproduction, making the present 
magnification approximately 1150 diameters for FiGS. 1-21 and 41, and 1375 

diameters for Figs. 22-40. 

Plate 5 

Lobomonas pentagonia Hazen 

Figs. 1-5, Typical vegetative cells or zoospores: in Figs. 1-3 the 
stigma lies on the under side of the cell. 

F1G.5. Anterior polar view: four anterior lobes emtpy, chromatophore 

filling four posterior lobes. 

Fig. 6. Protoplast rotating clockwise in preparation for division; 7 p. m. 

Fig. 7. The same cell at 7:05 p. m; beginning of cleavage. , 

Fig. 8. Division into two zoospores completed at 6:30 p. M. 

Fig. 9, Another individual: pyrenoid and cilia beginning to appear In 
daughter cells; nucleus in most advanced lobe: 10:30 P. M. 

Fig, 10. Pentagonal form attained by daughter cells. 

Fig. II. Four daughter cells in one plane. 

Fig. 12. Two daughter cells with axes perpendicular to those of the 

other pair; 

140 Hazen: New species of Lobomonas 

Fig. 13- An unusually simple form: cilia contracted into a ball. 

Fig. 14. Cell in posterior polar view. 

Figs. 15-20. Stages in conjugation of a pair of slightly unequal gametes: 
Fig. 15 at 1:45 p. m.; Fig. 16 at 2:15; Fig. 17 at 2;i6; Fig. 18 at 2:25; Fig. 
19 at 2:40; Fig. 20 at 2:45. 

Fig. 21, Zygote formed by conjugation of equal walled gametes. 


Plate 6 


Lobomonas rostrata Hazen 

Figs. 22-27. Typical young vegetative cells. 

Figs. 28, 29, Mature cells of less common form, but with well developed 
wall, Fig. 29 showing edge view of beak. 

Fig. 30. Anterior polar view. 

Figs. 31, 32. Cells showing persistence of form some time after loss of 

Fig. 33. Relatively large mature cell, pyrenoid and stigma underneath. 

Fig. 34. The same cell 24 hours later: four daughter cells completely 
organized except for cilia, 11 p. M. 

Fig. 35, One of the four daughter cells, one day later, not very well 
formed, probably because of unfavorable conditions. 

Fig. 36. Beginning of cleavage, after clockwise rotation of protoplast; 

9:30 p. M. 


Fig. 37. Similar division, after rotation of protoplast in vertical plane; 

10:30 p. M. 

Fig. 38. Second cleavage Just completed; 10:30 P. M, 


Fig. 39. Eight daughter cells (perhaps gametes) 6 p. M. 

Fig. 40. The same at 9:30 p. m.: daughter cells held by gelatinized wall 
of mother cell: cell a moved out sluggishly at 8:30; at 10 p. M. cell b backed 
out and presently rested on end, cilia downward as at bK 

Fig. 41. Mature cell from Shelburne, Vermont, August 3, 1921. All other 
figures from New Jersey material, in 1919. 

Bl'Ll. Tukrev i-'tre 

\'0LI ME 49, PLATE 5 
















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The development of the flower and embryogeny of 

Martynia louisiana 


Flora Anderson. 



The most common and widely distributed representative 
of the Martyniaceae in temperate North America is Martynia 
lotiisiana Mill. Although the Hterature on the family is chiefly 
taxonomic, the descriptions are meagre and confusing. AU the 
species have, at some time, been placed in the genus Martynia. 
The ten or more known species comprise three genera, native 
to the warmer parts of the western hemisphere. 

Efforts were made to secure seeds and to grow plants of the 
different species for comparison, in order, if possible, to work 
out the taxonomy of the family. Seeds were obtained from 
seed-houses both in the United States and in England and also 
from the Bureau of Plant Industry, the University of Texas 
and the Mississippi Agricultural College. 

From seeds secured through the Bureau of Plant Industry, 
quite a number of plants of Martynia louisiana were grown — 
all the plants producing viable seeds — and one specimen of 
Martynia lutea Lindl. which, although maturing seemingly 
normal fruit from selfed flowers, produced no viable seeds. 
Reciprocal crosses of Martynia lutea and Martynia louisiana, 
and of Martynia lutea and Martynia fragrans Lindl. failed. 

From seeds furnished by seed-houses, the University of 
Texas and the Mississippi Agricultural College, quite a number 
of plants of Martynia louisiana and Martynia fragrans were 
grown in Montgomery County, Indiana, during the summers 
1915-1919- The two species mentioned above cross very readily, 
producing in all cases viable seeds. These plants, grown in the 
open in rich soil and under practically the same conditions, 
showed very few differences except in the color of the corolla, 
which is more or less a variable character. The similarity ol 
the plants seems to indicate that they belong to the same species, 
but more cultures must be studied before this point canbedef- 

initely determined. 


142 Anderson: Embryogeny of Martynia Louisiana 

In as much' as no morphologicarwork seemed to have been 
done on this family, a study of Martynia lonisiana and Martynia 
fragrans was undertaken, the results of which are given in the 
following pages. While the study was based mainly on Martynia 
lottisiayia no structural differences between the two species were 

Material and methods 

The material used for this study was collected from plants 

of Martynia lonisiana and Martynia fragrans grown in the field, 

in Montgomery County, Indiana. Collections were usually rnade 

during the month of August, w^hen the plants were most vigorous. 

The apices of racemes, small flowers and stamens, pistils and 

young fruits in various stages of development w^ere killed and 

fixed in a I per cent chromo-acetic acid or in chrom-osmic-acetic 

acid one to two hours and then In chromo-acetic acid twenty -four 

to thirty-six hours, washed, dehydrated, and embedded through 

chloroform. To determine the time necessary for the pollen 

tube to reach the egg, pollination was controlled, and, beginning 

at one hour after the application of the pollen, collections were 

made at inter\'als of one hour for fourteen consecutive hours. 

These collections were of whole pistils w^hich were killed and 

fixed in chrom-osmic-acetic acid one to two hours and then in 

chromo-acetic acid thirty-six hours. Beginning thirty hours 

after pollination, collections of pistils and young fruits were 

made at intervals of six hours for two hundred and four hours. 

Sections were cut 4-10 (x, most of them 5 yL thick; those of whole 

pistils were longitudinal and 4-5 ^ in thickness. Both modified 

triple and Heidenhain's iron-alum-haematoxylin were used for 

staining, the former being more satisfactory. 

The development of the flower 

As in most inflorescences, the flower of Martynia louisiana 
arises as a lateral branch (Fig. i, X) in the axil of a 
bract (Fig. i, B). The first bract arises as a lateral papilla 

near the apex of the main axis of the raceme (Fig. 2, B'). 
This bract grows rapidly and about the time its tip reaches 
the apex of the raceme, a rather broad protuberance appears 
in the axil (Fig. i, X). With further growth of the axis of the 
raceme other bracts appear, and in the axils of these the primordia 
of other flowers develop (Fig. 4, X). As the rudiment of 

Anderson: Embryogeny of Martynia Louisiana 


the" flower grows it becomes broader and somewhat oblique 
at the apex (Figs. 2 and 3, X). On the adaxial side of 
the primordium of the flower there appears a protuberance 
or ridge, the upper lobe of the calyx, extending about one third 
the distance around it (Fig. 3, S). Very soon the other Icbes 
of the calyx appear as small protuberances similar to the ad- 
axial lobe. The individual lobes appear as separate protuber- 
ances, but later grow up as a tube except on the abaxial side of 



Figs. 1-7, Development of the flower. Figs. 1-3, X 45; Figs. 3-7, 
X 30. B. bract; X, flower primordium; S, S*, calyx lobe; P," corolla lobe; A, 
stamen; O, pistil; b, bracteole. 

the flower. At this point, the two lower lobes of the calyx are 
united only at the base and in the mature flower overlap. The 
adaxia) lobe of the calyx is the largest, and in the bud covers the 
other lobes (Figs. 4 and 7, S). The calyx lobe, shown atS' in 
Fig. 4, is the edge of one of the lateral lobes and appears smaller 
than it really is. Just within the calyx lobes are seen small 

144 Anderson: Embryogeny of Martynia Louisiana 


protuberances (Fig. 4, P), the primordia of the corolla lobes, 
which are alternate with those of the calyx. Arising simul- 
taneously with the corolla lobes (Fig. 5, P) and alternate with 
them are the epipetalous stamens (Fig. 5, A). Figs. 6 and 7 
are other sections of the flower shown in Fig. 5. Fig. 6 is a 
median section through the primordia of two lobes (P) of the 
corolla, while Fig. 7 is a median section through the primordia 
of two stamens (A). A comparsion of these two figures will give 
the relative size of the parts at this stage. There are primordia 

Figs. 8-13. Development of the flower (coii.), X 30- B- bract; b, 
bracteole; S, calyx lobe; P, F\ corolla lobe; A, A\ stamen; O, pistil; L, pla- 
centa; C, ovarian cavity. 


of five stamens, but very early the one between the posterior 
pair becomes aborted (Figs, id and 14, A'). The pistil is the last 
cycle to appear, and in Fig. 5 is seen as abroad protuberance (O;. 
In Figs. 5-9, at B, are seen cross sections of the bract which 
curves over the flower. These figures are of the flower as viewed 
from the adaxial side. And in the same figures at b, are seen 
sections of the two bracteoles. The two bracteoles arise as 

Anderson: Embryogeny of Martynia Louisiana 


protuberances similar to those of the lobes of the calyx, but 
earlier than the calyx lobes. The bracteoles develop inferior 
to the calyx, opposite each other and at right angles to the 
bract (Fig. 10, b, B). Since Fig. 9 is a median section through 
the flower, only the edge of each bracteole is shown. At this 
stage, they are large enough to cover the young flower. 

Figs. 8 and 9 show the further development of the parts 
of the flower. The calyx (S) is large enough to cover the other 
floral parts. An almost median section of two lateral lobes (P) 
of the corolla is shown in Fig. 9, while a cross section of the tip 
of the lower lobe is shown at P^ in Fig. 8. The lower lobe of 
of the corolla is the largest and in 
the bud is covered by the others. 
The manner in which the corolla lobes 
overlap is shown in Fig. ii. A 
median section through two of the 
stamens (Fig. 8, A) of the same flower 
shows the further development of 
the stamens seen at A, Fig. .7, As in 
most cases, the development of the 
anther precedes that of the filament 
(Fig. 8, A). However, at this time 
the tissue of the anther is undiff'eren- 

FiG. 14. Immature stamen. 

tiated. At the time 

the pollen A, aborted stamen. 

Fig. 15. Mature 

mother-cells are formed the filament 

is still shorter than the anther. The \^^J^ enarge . 


H, stigma 

filaments remain short (Fig. 14) until 

Fig. 16. Mature pistil and 

almost the time of an thesis, when they stamens in position. . H, stigma 
elongate rapidly, bringing the anthers lobes. 

into the throat of the corolla, just below the lips of the stigma 
(F.iG 16). The stamens are didynamous, the posterior pair 
being the longer. Just before an thesis, the lobes of the anther 
diverge, forming a straight. line transverse to the filament. At 
the same time, the filaments of the corresponding pairs of 
stamens converge in such a way as to bring the diverging 
anthers into contact (Fig. 16) along the line of their linear 
dehiscence. At time of shedding, the rather large pollen grains 
are characterized by an exine that is thickened in the form 
of hexagonal areas. 

As was stated previously, the pistil appears as a rather broad 
protuberance (Fig, 5, O), but soon a depression is formed in 
the center (Fig. 9, O). The primordium of the pistil continues 

146 Anderson: Embryogexy of Martyxia Louisiana 


to grow up as a ring enclosing a central ovarian cavity^ as is 
seen in cross section (Fig. id, C). At a very early stage in the 
growth of the pistil, two parietal placentae appear. These 
are seen as outgrowths from the walls of the cavity (Figs, io 
and II, L). These outgrowths extend toward the center of the 
cavity and soon each forms two lobes (Fig. 12, L) which develop 
into the two lamellae of each placenta, resembling in cross 
section the letter T (Fig. 13, L). The placentae grow together 
at the base, thus forming a short columella. A cross section 
through the base of the ovary would appear as two-celled, but 
no ovules are developed from this region of the placentae. The 
upper part of the pistil is tubular and more narrow, forming 
a long, slender, hollow style which terminates in an unequally 


Development of megaspore and embryo-sac 


The young ovule first appears as a small, nipple-like protub^ 
erance upon the placenta. Very early in its development, a 
hypodermal cell near the tip becomes differentiated and may be 
distinguished from the surrounding cells by its large nucleus and 
dense cytoplasm (Plate 7, fig. 17). This cell, the megaspore 
mother-cell, grows rapidly, keeping pace with the lengthenmg 
nucellus. At this stage, the nucellus consists of merely the en- 
larged megaspore mother-cell, covered by the epidermis (Fig. 18). 
At the base of the nucellus the single integument appears as a fold, 
at first more pronounced on the side next to the cavity of the 
, (Fig. 18, I). By the more rapid growth of this side the 
ovule soon becomes sharply curved, so that the long axis of the 
nucellus is now parallel with that of the funiculus. At this 
stage, the integument extends around the nucellus, which has 
become more elongated (Fig. 19). With further growth the 
integument soon equals and finally exceeds the nucellus 1^1^^- 
20. and 21). The megaspore mother-cell has also grown rapidly 
and when fully developed is still enclosed by only a single layer 
of nucellus (Fig. 21). The nucleus, which has increased in 
size preparatory to the first division and which has had a posi- 
tion near the center of the cell (Fig. 20) , comes to lie nearer the- 
chalazal end, where it undergoes the first or heterotypic division 
(Fig. 21). All the spindles of this stage found were rather small, 
as compared with the size of the cell. Whether cell formation, 
follows this first nuclear division was not ascertained, but the: 

AxDE.Rso:: Embryogeny of Martyxia Louisiana 147 

typical axial row of four megaspores Is finally formed (Fig. 22), 
and, as is usually the case, the innermost megaspore, the one 
next the chalazal end, persists to form the embryo-sac (Figs. 
22 and 23). The upper three potential megaspores are soon 
destroyed by the rapid growth of the functional one (Fig. 23.) 
By the time the tetrad is formed, the nucellus has reached 
its maximum growth, and the cells of the integument adjacent 
to the nucellus have become dense in cytoplasm and now func- 
tion as a nutritive jacket (Fig. 22). In the meantime the 
integument has grown and exceeds considerably the length of 
the nucellus (Fig. 2;^^). 

The functional megaspore does not increase much in size 
before the first nuclear division but, after this first division, it 
elongates rapidly and occupies a space almost the length of 
the nucellus (Figs. 23 and 24). As the megaspore elongates 
the two nuclei keep their relative positions — one in each end 
of the cell. Most of the cytoplasm of the cell is in the ends, 
thus leaving a large vacuole in the center. A few strands of 
cytoplasm penetrate the vacuole and form a connection between 
the masses at each end of the cell. The rapid growth of this 
cell has consumed the three upper potential megaspores and the 
nucellus around the center of the spore (Fig. 24). Before the 
second nuclear division of the megaspore takes place, the upper 
three megaspores have practically disappeared and the functional 
one now extends nearly the length of the nucellus, which is rapid- 
ly being used as food (Fig. 25). At this time the central vacuole 
is not so pronounced — ^at least in some cases. The spindles of 
the second nuclear division may be seen — one in each end of the 
cell. With the exception of a few cells at the chalazal end, the 
nucellus is consumed by the growing megaspore, which comes 
to occupy the space left by the disorganizing tissue. There 
is no evidence of a nucellar cap being formed as Merrell reported 
for Silphuim (3) and other plants that have a similar develop- 
ment. The nuclei now undergo another division which results 
in eight nuclei — four in each end of the cell. Next follows the 
formation of the three cells of the egg-apparatus and the three 
antipodals. One nucleus from each end of the embryo-sac, the 
polar nuclei, remains free in the cytoplasm. At this stage, the 
embryo-sac is considerably elongated, being about seven times as 
long as wide. The micropylar end, containing the egg-apparatus, 
is enlarged to about one and a half times the diameter of the 
other part of the sac. This enlarged portion comprises nearly one 

148 Anderson: Embryogeny of Martynia Louisiana 

third the length of the embryo-sac. The egg-apparatus consists 
of the large pear-shaped egg-cell, which extends far down into 
the sac cavity, and the two large synergids (Plate 8, fig. 26, 
G and Sn). The lower part of the egg in which the nucleus lies is 
well filled with cytoplasm, but the upper part is vacuolate. The 
synergids, one on either side of the egg, extend down about two 
thirds the length of the egg-cell. Each synergid has a diameter 
nearly equaling that of the egg and a centrally located nucleus, 
which is a little smaller than the egg nucleus. As is frequently 
found in other angiosperms, a fairly large vacuole lies below the 
nucleus In the synergid (Fig. 26, Sn). 

In the chalazal end of the embryo-sac are the three, rather 
long, slender, well-developed antipodals (Fig. 26, N). In most 
instances, the nuclei of the antipodals are fairly large and the 
cytoplasm has the character of that of the body of the embryo- 


After the formation of the cells at either end of the embryo-sac 
the polar nuclei move to the center and lie side by side for some 
time (Fig. 26, Pn), but before the pollen tube reaches the embryo- 
sac, the two polar nuclei fuse (Fig. 27). Sections were made of 
ovaries from flowers in which pollination had not occurred and 
it was found that the polar nuclei had fused (Fig. 28). So the 
fusion of the polar nuclei is in no way due to a stimulus from 
the pollen tube. The fusion nucleus does not approach the egg- 
apparataus, as Kanda reported for Verbena (2), but remains in 
the center of the embryo-sac. . Each polar nucleus has a very 
prominent nucleolus and after fusion the two nucleoli are very 
noticeable for some time (Fig. 27), but later only one large 
nucleolus is usually seen (Fig. 28). 


The polar nuclei seem to unite about the time of anthesis, 
for conditions like those represented in Fig. 26 are observed in 
ovaries fixed just before the expansion of the corolla lobes, while 
the fusion nucleus is seen in flowers that have been open one of 
two hours. Most of the flowers open early in the morning, and 
shortly after daylight the bees, which are the chief pollinizing 
agents, are busy gathering nectar from the lower portion of the 
corolla tube. 

Anderson: Embryogeny of Martynia Louisiana 149 

The tw^o lobes of the sensitive stigma stand open just above 
the open anthers (Fig. 16), in the upper portion of the throat 
of the corolla, and the bee, entering the corolla tube, brushes 
the stigma, thus depositing the pollen that may be present on 
the upper part of its body and securing a new supply from the 
open anthers just below as it walks down the tube. The stigma 
is very sensitive when the flower first expands and the slightest 
touch will cause the lobes to close. If pollen be present 
they usually remain closed, but, in the absence of pollen, they re- 
open in from five to ten minutes. If pollen is witheld the stigma 
lobes may be made to close and open many times, but after one 
to three days the stigmas tend to lose their sensitiveness. 
How^ever, this sensitiveness is not an index to the receptiveness 
of the stigmatic surface, for flowers hand pollinated after the 
stigma ceased to be sensitive and even after the corolla dropped, 
produced viable seeds. Viable seeds were also secured from 
flowers hand pollinated in the bud before the stigma lobes nor- 
mally opened. This was accomplished by gently forcing pollen 
between the closed lobes by means of a needle. In most flowers 
thus treated the stigma lobes never opened. At this time the 
anthers are still closed, but pollen from these as well as from 
open anthers was shown to be effective. 


Usually the pollen grains germinate immediately on the stig- 
matic surface, as was readily seen in material fixed one hour 
after the pollen was applied. At this time, many of the tubes 
were well into the tissue of the style. The growth of the pollen 
tube through the tissue, usually two or three cells from the 
stylar canal, is very rapid, for fertilization frequently occurs 
within six hours after pollination. The pollen tube is filled 
with plastids and granules that take such a dense stain that it 
is impossible to demonstrate the presence of the sperm nuclei, 
but, since the long, slender crescentic generative cell In the mature 
pollen grain shows only one nucleus, it is thought that the for- 
mation of the two sperms takes place in the tube. The pollen 
tube, as It grows nearly straight down through the style to the 
placenta, is comparatively broad and seems to retain its densely 
staining quality throughout the greater part of its length. 

I50 Axderson: Embryogeny of Martynia Louisiana 

^ h' 




i i 







On reaching the ovule, the pollen tube 
grows through the micropyle and enters 
the embr^^o-sac, sometimes a little to one 
side, crushing one or both the synergtds. 
The tip of the pollen tube usually covers 
the egg-cell, thus obscuring the nucleus of 
the latter (Fig. 32). ^^n some instances it 
seems that the tube enters the egg-cell at 
the upper end and transverses the entire 
length of it (FiG- 31). In all cases ob- 
served, there was evidence that the tube 
had discharged the sperm nuclei in such 
a way that one of them came in direct 
contact with the egg nucleus (FiG. 30). 
Only the sperms and a little of the con- 
tents from the tip leave the pollen tube 

(Figs. 30 and 31). The fate of .the 

second sperm nucleus was not ascertained, 
but it seems evident that it plays no part 
in the formation of the endosperm. The 
second sperm nucleus may not leave the 
tube, but this point can not be demon- 
strated until some method of fixing and 
staining can be found that will reveal the 
presence of the nuclei among the granules 
in the cytoplasm. The pollen tube in the 
micropyle and in the upper part of the 
embryo-sac retains its form and densely 
staining quality long after the embryo-sac 
is filled with endosperm (Fig. 35). 

The actual union of sperm and egg nu- 
cleus is similar to that in other angio- 
sperms. The instances of fertilization 
observed were similar to that of Lilitini 
Martagon, figured by Mottler (4). For 
Verbena, Kanda (2) figures the sperm 
nucleus in contact with the egg-cell and 
speaks of it as fertilization. In all the 
fertilization staees observed in Martynia, 

Fig. 35. Embryo-sac filled with endosperm, 

X 140 

pollen tube 

U, suspensor 

Anderson: Embryogeny of MartyxNia Louisiana 151 

the sperm has been in direct contact withthe egg nucleus. 
This would seem to indicate that the pollen tube carries the 
sperm into the egg-cell. 

The endosperm 

The primary endosperm nucleus, resulting from the union 
of the two polar nuclei, occupies a position near the center of 
the embryo-sac. Usually before the pollen tube enters the sac 
this endosperm nucleus divides and a transverse wall is formed 
between the two resulting nuclei, thus dividing the embry-sac 
into two chambers (Fig. 29), At this time, the cytoplasm in 
the embryo-sac has a frothy appearance due to the many small 
vacuoles present. This frothy character seems to appear after 
the fusion of the polar nuclei (FiG. 28). Very" conspicuous, 
densely staining plastid primordia and granules are present in 
the cytoplasm. The synergids seem to have increased in length 
and to have become more vacuolate (Fig. 29). The egg-cell 
remains apparently unchanged.. The antipodals are more 
elongated and sometimes contain a granular, densely staining 
cytoplasm .(Fig., 29). After the wall dividing the embryo-sac 
into two chambers is formed, the nuclei move to a nearly central 
position in their respective cells. And, as one would infer, the 
endosperm cell at the micropylar end of the sac grows more 
rapidly than the one at the chalazal end (Fig. 31). The pollen 
tube usually reaches the embryo-sac when the endosperm is in 
a tw^o-celled stage. In Fig. 31, it may be seen that fertilization 
has taken place. The outline of the tip of the pollen tube is shown 
by a dotted line in the lower part of the egg-cell, and the shading 
represents the relative density of the contents in different parts 
of the tube. Only the tip seems to have discharged its contents. 
A remnant of one synergid still shows. The synergids, if 
not crushed out by the entrance of the pollen tube, are soon ab- 
sorbed. The endosperm cell in the micropylar end of the sac 
is very large and contains a large nucleus with Irregular oudine 
and a prominent nucleolus. Many plastlds are still seen in the 
frothy cytoplasm of both endosperm cells, and some are present 
in the antipodals, which continue to grow, especially in length 
(Fig. 31). The increase in size of the antipodals is better illus- 
trated in Fig. 32. Their cytoplasm is somewhat vacuolate at 
this stage and the large nuclei contain 'very prominent nucleoli. 
Usually the first division of the endosperm cell is transverse 

IS2 Anderson: Embryogeny of Martynia Louisiana 

and the second longitudinal, the one in the micropylar end divid- 
ing before the one in the chalazal end; but, in Fig. 32, it appears 
as if the first division had been longitudinal and the second 
transverse. The cytoplasm in the large cells around the fertilized 
egg contains many large as well as small vacuoles, while 
that of the lower cells contains only small ones. Very few 
plastids are present at this stage. They are seldom noticeable 
In later stages of endosperm development. Fig. 32 shows also 
the old densely staining pollen tube in the micropyle and upper 
end, of the embryo-sac, its tip covering the fertilized egg. By 

focusing down on the tip of the tube, the 
nucleus of the fertilized egg is seen and is 
represented in the drawing by a dotted line 
(Fig. 32). In the meantime, the whole em- 
bryo-sac has been enlarging at the expense 


of the integument which also has grown. 
The enlargement is more pronounced at first 
in the micropylar end where the fertilized 
egg is elongating. The layer of cells of the 
integument adjacent to the embryo-sac is 
always dense and rich in cytoplasm and func- 
tions as a nutritive jacket to the growing 
endosperm. Fig. 33 shows the further de- 
velopment of the endosperm, and the 
elongation of the fertilized egg. The old 
pollen tube is still very prominent in the 

Fig. 36 Outline ^^^^^PY^^ 2ind upper end of the sac, obscur- 
of ovule, X 44. 1, integ- ^^g the Upper portion of the fertilized egg-cell. 

ument;M, embryo-sac The cytoplasm in the large endosperm cells 
E, embryo; U^suspen- of the micropylar end is very vacuolate, due 
sor; T, old pollen tube, p^bably to the rapid growth of the cells and 
to the loss of food to the developing embryo. Only a few 
endosperm cells develop in this end of the sac, for very soon 
a long suspensor is formed which brings the embryo into the 
chalazal end. The endosperm cells in the middle of the sac 
are smaller and richer in cytoplasm than the upper ones. The 
antipodals have nearly reached their maximum growth. They 
seem to function as elaborators of food and persist long after 
the embryo-sac is well filled with endosperm (Fig. 35). In 
several cases, four antipodals, all well developed, were found 
(Fig. 33). The extra one had probably been formed by the 

Anderson: Embryogeny of Martynia Louisiana 153 

division of one of the three normal ones. Fig. 34, a cross section 
of the chalazal end of an embryo-sac in the same stage as the 
above, shows the arrangement of the four antipodals. At this 
stage, the nuclei of the antipodals appear as densely staining 
homogeneous masses. 

As the embryo with its very long suspensor grows through 
the center of the elongating embryo-sac, large endosperm cells 
rich in cytoplasm are formed, but when it reaches the lower 
part of the sac where the first division of the embryo takes 
place, the endosperm growth practically ceases in the middle 
and upper part of the sac. The cells of this part of the sac are 
rather large and similar in shape to those first formed, while 
those in the lower part are smaller and not so rich in cytoplasm 
(Fig. 35). It is also seen from Fig. 35 that the embryo-sac has 
enlarged considerably, especially the lower part which contains 
the developing embryo. The nutritive jacket is still very active 
in the elaboration of food for the growing endosperm. Fig. 36 
shows in outline the relative size of the embryo, endosperm and 
integument at the time the embryo is in the two-cell stage as 
seen in Fig. 35. 

The embryo 



After fertihzation, the egg with its nucleus occupying a 
nearly central position in the lower part of the cell grows rapidly 
in length (Fig. 33). Its manner of growth down through the 
endosperm is similar to that of a pollen tube, for it becomes a 
long narrow cell with the nucleus retaining a position near the 
growing tip. No division seems to take place until the distal 
end has transversed nearly two thirds the length of the embryo- 
sac. The lower portion of the cell is rather dense in cytoplasm 
and stains deeply, but the upper portion has very little and 
stains so faintly that the course of its growth is hard to trace. 

After the distal end has reached the lower portion of the 
embryo-sac, a transverse division occurs, which forms a very long, 
slender suspensor cell and a short, rounded distal one, the em- 
bryonal cell (Fig. 37). In most instances the cytoplasm of the 
embyonal cell contains many plastids and other densely staining 
granules, while that of the suspensor is ver>^ poor in plastids. 
After the first division two others occur almost simultaneously, 
The suspensor cell divides, forming a short cell next the embryo- 
nal cell, w^hich has enlarged preparatory to division, and. at the 

154 Anderson: Embryogeny of Martyxia Louisiana 

same time, the enlarged embryonal cell divides longitudinally 
forming a two-celled embryo (Figs. 38 and 35). The second 
division of the embryo is usually transverse and the third longi- 
tudinal, thus forming the octant stage The further growth of 
the embryo is illustrated in Figs. 39-43 and is so like the well 
known development of Capsella that details are unnecessary. 
The very long suspensor, shown in Fig. 35 in dotted 
lines, is almost indentical with that of Trapella sinensis Oliv. as 

Pl- 7, / 


Figs. 37-43. Development of the embryo. Figs. 37-38, X 500; Figs. 
39-42, X 175; Fig. 43, X 6. E, embry^o; U, suspensor; C, cotyledon; R, 
radicle; K, chalazal end of embr>'o-sac; Q, micropylar end of embryo-sac; 
N, remains of antipodals; V, soft, inner portion of seed coat; W, hard, warty 
portion of seed coat. 

serratum as figured by Billings (1,/. S^) is also similar in struct- 
ure but does not push the embryo as far down into the embryo-sac 
as that of Martynia, although the shape of the embryo-sac and 
the development of the endosperm are much the same in both 
species. Usually a second and sometimes a third division 
occurs in the suspensor, resulting in two or three short cells 
next the embryo, the innermost of which seems to contribute to 
its formation. 

Fig. 43 shows a section of a young seed containing a nearly 
mature embryo. The radicle (R) is short and thick while the 

Anderson: Embryogeny of Martynia Louisiana 155 

cotyledons (C) are very large and fleshy." This figure shows 
also a little endosperm which, in the mature seed, comprises 
a thin, transparent covering for the embryo. The embryo has 
developed in such a way that it practically fills the lower part 
of the sac, while the upper and middle portions (Q) form in the 
mature seed a little brown tube on the micropylar end of the 
remaining endosperm. There is also a small brown knob at 
the chalazal end (K) marking that part of the embryo-sac for- 
merly occupied by the large antipodals, the remains of which 
appear at N. The broken line in this figure represents the 
demarcation in the tissue forming the seed coat. That region 
of tissue next the endosperm (V) matures into the soft inner 
portion of the seed coat, while the walls of the cells forming the 
outer portion (W) become thick and lignlfied, thus maturing 


black in color. 


The flowers develop in the axils of protective bracts. The 
cycles of the flower appear in the following order: calyx, corolla 
and epipetalous stamens, and pistil. 

Of the four potential megaspores formed, only one is function- 

The embryo-sac is long and narrow, with the micropylar end 

considerably enlarged. The large egg-cell extends far down 
into the sac cavity, with the two synergids on either side. The 
three antipodals are long, slender cells that increase in size and 

persist as elaborators of food. 

The polar nuclei move to a position near the center of the 
sac where they unite before the pollen tube reaches the ovule. 

Fertilization usually occurs within six to eight hours after 


The rather large pollen tube stains densely throughout the 

greater part of its length and persists in the micropyle and upper 

end of the embryo-sac long after the endosperm is formed. 

The first division of the endosperm cell, which is usually 
transverse, occurs before fertilization. At first, a few cells 
are formed in the upper and middle portions of the embryo-sac, 
but the bulk of the endosperm is in the lower part of the sac, 
where the embryo develops. 

The suspensor is very long and narrow and pushes the embryo 

156 Anderson: Embryogeny of Martynia Louisiana 

into the lower portion of the embryo-sac, where the first division 
takes place. The development of the embryo proper is like 
that of CapseJla. 

I take opportunity here to express my obligations to Pro- 
fessor H. W. Anderson for the use of the Wabash College Bot- 
anical Laboratory during the summers of 191 5 and 1916 and to 
Professor D. M. Mottier for valuable assistance and criticism 
in connection with this study. 

Indiana University 

Literature cited 



1. Billings, Frederick H. Beitrage zur Kenntniss der Samenent- 

wickelung- Flora 88; 253-318. 1902. 


2. KandE) M. Field and laboratory studies of Verbena, Bot. 

Gaz. 69: 54-71- P^^ ^^9 + /• ^0-17, 27, 54-^3^ ^9^75^ 1920. 

3. Merrell, W. D. A contribution to the life history of 

Silphium. Bot. Gaz. 29: 99-133. pL 3-10. 1900. 

4. Mottier, D. M. Fecundation in plants. Carnegie Inst, Wash- 

ington PubL 15. 1904. 

5. Oliver, F. W. On the structure, development, and affinities 

of Trapella, Oliv., a new genus of Pedalineae. Ann. Bot. 

2: 75-115- pl' 5~9^ 1888. 

Explanation of plates 7 and 8 

Plate 7 

All figures, x 400 

Fig. 17. Vertical section through the nuceUus; megaspore mother-cell 

Fig. 18. Elongated nuceUus with megaspore mother-cell; beginning of 
integument, i. 

Fig. 19. Vertical section of young ovule turned on its axis; the integu- 
ment reaching half the length of the nucellus. 

Fig. 20, Nearly mature megaspore mother-cell enclosed in a single 
layer of nucellus. 

Fig. 21. Heterotypic spindle in megaspore mother-cell. 
Fig. 22. Tetrad, the upper three potential megaspores disorganizing. 
Nutritive jacket formed from the layer of integument adjacent the nucellus. 
Fig. 23. Section of young ovule showing the single integument and 

first division of the megaspore; the upper three potential megaspores dis- 

Figs. 24 and 25. Further development of the megaspore; the central 
portion of nucellus disorganizing. 

Bull. Torrey Club 

Volume 49, plate 7 

W- i 


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Bull. Torkey Ciub 

Volume 49. plate 8 














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Anderson: Embryogeny of Martynia Louisiana 


Plate 8 


Figs. 26-33. longitudinal sections of embryo-sacs. Fig. 34, cross section 
of antipodals. 

Fig. 26. Mature embryo-sac; polar nuclei side by side near the center 
of the sac; G, egg-cell; Sn, synergid; N, antipodals; Pn, polar nuclei, X 300. 

Fig. 27. Mature embryo-sac; polar nuclei fusing, X 300. 

Fig, 28. Mature embryo-sac; polar nuclei fused, X 350. 

Fig. 29. Endosperm two-celled; egg not yet fertilized, X 350. 

Fig. 30. Fertilization. Sperm and egg nuclei in contact; old pollen 
tube quite prominent; G, egg-cell; T, pollen tube; Sn, synergid, X 740. 

Figs. 31 and 32. Development of endosperm and elongation of anti- 
podals, X 350. 

Fig. 33. Development of endosperm and embryo; four antipodals, N; 
E, elongating embryo; T, old pollen tube, X 350. 

Fig. 34. Cross section of chalazal end of embryo-sac showing four anti- 
podals, N, X 350- 




The aim of this Index is to include all current botanical literature written by 
Americans, published in America, or based upon American material; the word 
America being used in the broadest sense. 

Reviews, and papers that relate exclusively to foresty, agriculture, horti- 
culture, manufactured products of vegetable origin, or laboratory methods 
are not included, and no attempt is made to index the literature of bacteriology. 
An occasional exception is made in favor of some paper appearing in an Ameri* 
can periodical which is devoted wholly to botany. Reprints are not mentioned 
unless they differ from the original in some important particular. If users 
of the Index will call the attention of the editor to errors or omissions, their 
kindness will be appreciated. 

This Index is reprinted monthly on cards, and furnished in this form to 
subscribers at the rate of three cents for each card. Selections of cards are 
not permitted; each subscriber must take all cards published during the term 
of his subscription. Correspondence relating to the card issue should be 
addressed to the Treasurer of the Torrey Botanical Club. 

Adams, J. F. Observations on the spores of Sichzophyllum 
commune. Torreya 21: 98-100. D 1921. [Illust.] 

Allen, W. E. Some work on marine phytoplankton. Trans. 
Am. Micr. Soc. 40: 177-181. O 1921. 

Allen, W. E. The investigation of ocean pasturage. Ecology 

2: 215-219. 3 D 1921. 

Ames, O. Additions to the orchid flora of Panama. Proc. Biol. 

Soc, Washington 34: 149-154- 21 D 1921, 

Includes 7 new species. 


Ames, O. Eria gigantea. Gigantic eria. Addisonia 6: 41, 

42, pL 213. 24 O 1921. 


Anderson, E. G* The inheritance of salmon silk color in maize. 

Cornell Univ. Agr. Exp. Sta, Mem. 48: 539-554. N 1921. 

Anderson, P. J., & Ickis, M. G. Massachusetts species of 

Helvella. Mycologia 13: 201-229. pL 11, 12, 8 O 1921. 

Arthur, J. C. Memoranda and index of cultures of Uredineae, 

1899-1917. Mycologia 13: 230-262. 8 O 1921. 

Ashe, W. W. Notes on Rhododendron. Rhodora 23: 177-179. 

24 O 1921. 


i6o Index to American botanical literature 

Bailey, L* H. Magnolia stellata. Addisonia 6: 37, 38. pL 211. 

24 O 1921. 

Ball, C. R. Notes on willows of sections Pentandreae and Nigrae. 

Bot. Gaz. 72: 220-236. /. 1-4. 15 O 192 1. 

Includes 2 new varieties of Salix from California. 


Barker, E. E. Bud variation in the sugar cane. Jour. Hered- 
12: 271-274,/. 2/. 23 S 1921. 

Bechtel, A. R. The floral anatomy of the Urticales. Am. 


14 N 1921. 


Bird, H. Soil acidity in relation to insects and plants. Ecology 

2: I93-I97- 3^ 1921. 

Birge, E. A., & Juday, C. Further limnological observations 

on the Finger Lakes of New York. Bull. Bur. Fisheries 
37; 211-252. /, 7-5, 8 O 1921. 

Includes notes on plankton botany. 

Blakeslee, A. F., Cartledge, J. L., & Welch, D, S. Sexual 

dimorphism in Ctmninghamella. Bot. Gaz. 72: 185-219. 
/. J. 15 O 1921. 

Boynton, K, R. Bergenia crassijolia. Thick-leaved saxifrage. 

Addisonia 6: 43. pL 214. 24 O 192 1. 

Boynton, K, R. Helichrysum bracleatum. Strawflower. 

Addisonia 6: 46. pL 213. 24 O 192 1. 

Bradshaw, R. V. Cynostinis echinatus in Oregon. Torreya 

21: 81-83. O 1921. 

Braun, E. L. Composition and source of the flora of the Cin- 
cinnati (Ohio) region. Ecology 2: 161-180./. j. 3D 1921. 

Briggs, G, Para and Paspahim: two introduced grasses of 

Guam. Guam Agr. Exp. Sta. Bull, i: 1-44. pL 1-6. 2 
D 1921, 

Britten, E. G. Clerodendron Thompsonae, Mrs. Thompson's 

clerodendron, Addisonia 6: 39, 40, pi, 212. 24 O 1921. 

Britton, N. L. Lathyncs latifolins. Everlasting pea. Addiso- 
nia 6: 35. pL 210, 24 O 192 1. 

Britten, N. L. Plant nomenclature: more suggestions. Jour. 

Bot. 59: 296, 297. O 1921. 

Brotherston, R. P. The pentstemon. Gard- Chron. 70: 208. 

22 O. 1921. 

Index to American botanical literature i6i 

Burlingame, L. L. Variation and heredity in Ltipinus. Am. 

Nat. 55: 427-448. /. j-j. O 1921. 

Cayla, V. A propos de la qualite du cauotchouc d'Hevea 
brasiliensis, ses variations, leurs causes. Importance 
economlque pour la production de I'Amazone. Arch. 
Escol. Super. Agr. e Med. Veter. 5; 81-96. S 1921. 


Chace, E. M., Wilson, C. P., & Church, C. G. The composition 

of California lemons. U. S. Dept. Agr. Bull. 993: 1-18. 
/. 1-4. 15 O 192 1. 

Chamberlain, C, J. Growth rings in a monocotyl. Bot, Gaz. 
72: 293-304./. 1-16, 15 N 1921. 

Clark, A. H. A new classification of animals. Bull. Instit. 
Oceanogr. 400: 1-24. 20 S 1921. 

Includes a summary of plant classification. 

Clements, F. E. Drouth periods and climatic cycles. Ecology 
2: 181-188. 3 D 1921. 

Coker, W. C, & Beardslee, H, C. The collybias of North 
Carolina. Jour. Mitchell Sci. Soc. 37: 83-107. pL i +/. 4-2^, 
D 1921. 

Includes Collyhia lilacina, sp. nov. 

Cook, F. C. Absorption of copper from the soil by potato 
plants. Jour, Agr. Research 22: 281-287. 29 O 192 1. 

Cook, M. T. The blossom blight of the peach. Phytopathology 

11: 290-294. pL 12. 15 N 192 1. 

Cook, M- T. Falling foliage. Phytopathology 11: 337-339. 

9 D 1921. 

Cook. M. T. Peach yellows and little peach. Bot. Gaz, 72; 

25^255. pi. 6, 7. 15 O 1921. 

Cook, M. T. Wilting caused by walnut trees. Phytopathology 

11: 346. 9 D 1921. 

Cook, O. F. Causes of shedding in cotton. Jour. Hered. 12: 

199-204./. s~6. 23 S 1 92 1. 

Coville, F. V. Directions for blueberry culture, 192 1. U. S. 

Dept. Agr, Bull. 974: 1-24. pL i-2g. 15 O 1921. 

Creswell, C. F., & Bid well, G- L. Composition of cotton seeds, 

U. S. Dept. Agr. Bull. 948: 1-221. pL i-\-maps i-ii. lo S 


i62 Index to American botanical literature 

Dana, B. F. Two new Sclerotinia diseases. Phytopathology 

11: 225-228. pi. 8. 6 O 1921. 

Doolittle, S. P. Overwintering of the bacterial wilt of cucur- 
bits. Phytopathology 11: 299, 300. 15 N 1921. 

Dupler, A. W. The air chambers of Reboulia hemisphaerica. 
Bull. Torrey Club 48: 241-252./. 1-22. 13 D 1921. 

Ehlers, J. H. Paniciim virgahim var. cubense in Michigan. 

Rhodora 23: 200. 15 D 1921. 

Eldridge, A. E. Woodland flowers of the prairie region. Am. 

For. 27: 714-716. N 1921. [Illust.] 

Emerson, F. W. Subterranean organs of bog plants. Bot. 
Gaz. 72: 359-374- /• ^-^^- I5 D 1921. 

Emerson, R. A. Genetic evidence of aberrant chromosome 
behavior in maize endosperum. Am. Jour. Bot. 8: 411-424. 
/. /• 14 N 1921. 

Ensign, M. R. Area of vein-islets in leaves of certain plants 

as an age determinant. Am. Jour. Bot. 8: 433-441. pL 
23. 19 D 1921. 

Evans, A. W. The genus Riccardia in Chile. Trans, Con- 
necticut Acad. Sci. 25: 93-209. /. 2-IJ. D 192 1. 


Includes 3 new species and 17 new combinations, 

Fairman, C- E. The fungi of our common nuts and pits. Proc. 

Rochester Acad. Sci. 6; 73-115, pL 15-20. S 192 1. 

Includes Schizocapnodiunij gen. nov., and 27 new species in various genera. 

Fehlinger, H. Brasilianische Faserpflanzen, Tropenpflanzer 
24: 173-176. D 1921. 

Fujioka, M., & Takahaski, K. On the cause of the darkening 
of the heartwood of Cryplomeria japomca Don. Jour. For. 
19: 844-866. D 1921. 

Gardner, G. A. Observations sur les articles du Reverend Pere 
Fontanel, S. J. Nat. Canad. 48: 49-56. S 1921. 

Gardner, M. W., & Gilbert, W. W. Field tests with cucumber 

angular leafspot and anthracnose. Phytopathology 11: 
^98, 299. 15 N 192 1. 

Gardner, M. W., & Kendrick, J. B. Soybean mosaic. Jour. 
Agr. Research 22: 111-113. pi. 18 ig. 8 O 1921. 

Vol. 49 

No. 6 




JUNE, 1922 

Mycological notes for 1920* 


I. Zythia resinae (Ehrenb.) Karst. 

For several years there has been noted in the mo.untains 
near State College a small inconspicuous fungus always confined 
to resin exudate on species of two- and three-needle pines. It 
has been found especially abundant in the vicinity of Charter 
Oak and Greenwood Furnace, Pennsylvania. Its habitat is 
the resin mass inhabited by the larvae of the "pitch midge," 
Cecidomyia resinicola Osten-Sacken. The origin of the resin 
exudate has not been studied by the writer, but E. P. Feltf 
states (on the authority of Miss L. S* Eckel) that the resin flow 
from the tree is not the result of an injury caused by the 
midges, but that they '*take advantage of normal exudations 
of resin on w^hich to oviposit," although '*the flow of this 
substance may be increased to some extent by the larvae 
rasping the living tissues." At any rate the final resin masses 
are subglobose or depressed globose and 1.5-2,5 cm. in diameter 
(Plate 9, fig. 2), They may occur on one side of the limb only, 
as stated and illustrated by Felt, or the exudate may entirely 
encircle the limb, though I have seen no instances of the saccate 
type of resin mass illustrated by Felt. These masses are in 

*Contribution from the Department of Botany, The Pennsylvania 
State College, No. 38. **Some mycological notes for 1919" was published in 
Mycologia 12; 135-142. 1920. 

fN. y. State Mus. Memoir 8: 410-412. 1906. 

[The Bulletin for May (49: 123-162. pL 5-8) was issued June 15, 1922.] 


i64 Mycological notes for 1920 


all cases found near the ends of the branches and always at the 

forks of the twigs. 

Eventually the fungus produces its pycnidia on this hardened 
and blackened resin mass. These are minute elongate- 
ovoid or almost conical bodies (Plate, 9, fig. r, at left), yellow 
or amber-colored, and large enough to be easily visible, on close 
inspection, to the unaided eye. The ostiole is usually quite 
prominent, and may often be seen with a hand lens. In texture 
the pycnidia are waxy and easily crushed when mounted In 
water. The spores are globose, smooth, hyaline, and very small, 
measuring only about 2 ^J^ in diameter. 

The nature of the pycnidium places the fungus In the Nee- 
trioidaceae of the Fungi Imperfect!. After several unsuccessful 
attempts at identification, the species was determined for me 
as Zythia resinae by Mrs. F. W. Patterson, who wrote that my 
specimens represented the only American material she had seen. 
Later Dr. Seaver identified my plants as being the conidial 
stage of Nectriella resinae (Fr.) Sacc, as listed by Saccardo. 
Nedriella is a genus of the true Nectriaceae, order Hypocreales, 
and consequently with an ascus in the life history. Examination 
of material collected at various times over a period of several 
years has failed to show on the same substratum any ascospore- 
producing structure comparable to the perithecium of a Nectriella. 
There is produced, however, on the same resin masses that bear 
the pycnidia an apothecial fructification that has been referred 
to Biatorella resinae (Fr.) Mudd., as listed below, 

2. Biatorella resinae (Fr.) Mudd. 

In company with Zythia resinae, as described above, there 
frequently grows an apothecial fruiting body that was determined 
as Biatorella resinae by Dr. L. W. Riddle, to whom my specimens 
were sent by Dr. Seaver. The very small apothecia are about 
I mm. in diameter and yellowish or amber-colored (Plate 9, fig. 
I, at right). They have been collected on resin exudate in- 
habited by the larvae of the "pitch midge*' on Pintis virginiana 
at Charter Oak and on P. ponderosa at Greenwood Furnace. 
The curious habitat and the frequent association of these apo- 
thecia wuth the pycnidia of the Zythia suggest a connection 
between the two, but if the Zythia is the pycnidial stage of the 
Nectriella such could hardlv be the rase. 

Mycological notes for 1920 


3. PiLACRE Petersii B. & Br. 

This rare species was found for the first time in the writer's 
experience on a dead snag of Acer rubrum, July i, 1920, on Stone 
Creek, Huntingdon County, Pennsylvania. Hundreds of the 
small brown stipitate fruiting heads were present on a small 
area (Plate 9, fig, 4), and their abundance suggested that 
this species is probably not so rare as has been supposed but, 
because of its small size and inconspicuous coloration, is usually 


Figs. 1-4. Pilacre Petersii B. & Br. 

I. Hyphae with clamp connections. 2 and 3. Basidia with spores. 
4. Spores. 

overlooked. Cross-septate basidia characteristic of the Auricu- 

lariales are always difficult to demonstrate, but the writer has 

located them w^ith more ease in this species than in any other, 

though not readily demonstrable at best (Text figs. 2 and 3). 

The fungus departs widely in consistency and habit from other 

members of the order. The following descriptiv^e notes are 

appended : 

Sporophores gregarious on the substratum, sometimes grow- 
ing from a slight superficial gray mycelium, 2-5 mm. high, 
consisting of a slender stalk bearing a single subglobose or 
hemisphaeric head; head 0.5-2.5 mm. in diameter, at first gray- 
pruinose, then deep brown, covered by a thin membrane that 
disappears at maturity, filled with a brown spore mass mixed 
with hyphae and powdery at maturity; spores sessile on the 
basidia, subglobose, smooth, brown, 5-7.5 \i diameter; basidia 
transversely three-septate (four-celled), hyaline, straight or 
curved, produced laterally on somewhat larger hyphae, 15-24^1. 
long, 2-3 yi in diameter, the terminal cell bearing its spore 
apicaJly, the other cells each with a lateral spore produced on 
either side of the basidium if straight, but on the convex surface 
if curved; hyphae hyaline, then brown, somewhat branched, 
with cross walls and clamp connections, 2-3 [i. in diameter; 

i66 Mycological notes for 1920 

stem central, grayish white, 0.2-0.35 mm. in diameter. [Text 
FIGS. 1-4.] 

On dead wood of deciduous trees. 

4. TuLAsxELLA ViOLAE (Quel.) Boud, & Gal. 

The genus TtdasneUa, recently monographed for this 
country by Burt, is an anomalous genus, CorticiumA^ke in 
appearance, but hardly so in structure. It is characterized by 
the presence of large sterigmate-like processes, four in number 
(usually), that are produced on a basidium-like organ, and that 
in turn bear each a hyaline spore. Only three species are list- 


ed by Burt, all of narrow geographical range and none hitherto 
reported from Pennsylvania. 

A small collection of T. Violae was made in the mountains of 
Center County, Pennsylvania, on January 29, 1921, on the inside 
of a gum (Nyssa sylvatica) log by the writer's mycology class while 
on a. collecting trip. The discovery of the specimens is credited 
to Mr. W. A. Kuntz. 

The color of these specimens was flesh pink or livid pink 
(RIdgway) when collected, but it is not probable that they will 
long retain these colors in the herbarium. In aspect the plants 
resemble very much a thin species of Corticium^ and casual ob- 
serv^ations of free hand sections show nothing very different 
from sections of species of that genus. Spores were abundant 
in the collection, however, and it was only when the usual 


attempt was made to substantiate their connections to basidia 
that the real nature of the hymenium became apparent. With 
some difficulty the large clavate or elongate sterigmata were 
made out, sometimes four in number and sometimes apparently 
but three. After the development of the sterigmata the basidia 
appear to collapse, or at least in a spore-bearing condition the 
group of sterigmata is so large as to give this impression. The 
spores are broadly ellipsoid to globose,- and in these specimens 
have two or three large conspicuous guttulae The spores 
measure 5.5-7.5 x 5 yi. No cystidia are present. 

5. Dacryomyces hyalina Quel. 

The species of the genus Dacryomyces are not well known in 
this country, partly because, in spite of their usually bright 
colors, they dry down and are rather inconspicuous in dry 
weather, when collectors are most likely to be in the field. D. 
hyalina, as determined for me by C, G. Lloyd in 1919, was 

Mycological notes for 1920 


found for the first time in Pennsylvania on January 29, 192 1, 
on a fallen log of Pinus Strohus. The dried plants appear as 
small scaly growths, dark in color, but when wet they swell 
up and take on a more waxy appearance and gelatinous texture. 
Fresh specimens measure 1-2 mm. in diameter, or by confluence 


as much as 3 mm. The color is a wax>^ yellow in young speci- 
mens, verging to olivaceous or brownish when dried. The 
fructification is almost disk-shaped, being considerably flattened, 
and the upper surface may be smooth or thrown into serpentine 
folds. The hymenium is composed of the usual bifurcate 
basidia (Text fig. 7), each with two spores. The spores (Text 
FIG. 5) are short-cylindric though often narrowed at one end, 
hyahne, one-celled, and measure 10-12 x 3-4 \i. The hyphae in 


Figs. 5-8. Dacryomyces hyalina Quel. 

5. Mature spores. 6. Hypha with clamp connection, 7. Mature 
basidia, 8. Hypha with scabrous wall and clamp connection* 


the trama vary from 4 [^ to 6 [i. in diameter and the walls of the 
larger ones appear scabrous. Branching and anastomoses are 
common, but cross walls are not abundant. Contrary to the 
usual condition in the • Tremellales and the Dacr>'omyce tales, 
clamp connections are found on the hyphae, though not abundant. 
On the older scabrous hyphae they are rather inconspicuous 
(Text fig. 8) but on the younger hyphae they are not difficult 

to locate (Text fig. 6). 

Good figures of this plant were recently published by Lloyd 
(Myc. Notes, No. 58, p. 828. 1919), who reports the species 
from North Carolina. The name employed to designate the 
species is open to criticism. The plants seem to be specifically 
distinct from D. deliquescens, which they resemble microscopical- 
ly as to spores and basidia, since the latter seems to lack the large 

i68 Mycological notes for 1920 

scabrous-walled hyphae and the clamp connections of D. hyalinus. 
I have not made, for D. deliquescens, an exhaustive search for 
clamp connections, but have failed to find them in the mounts 
I have made. Neither have I ever seen D. deliquescens assume 
the dark colors on drying, characteristic of the present species. 


6. Stereum radiatum Peck 

The fourth Pennsylvania collection of this species was made 
in October 2J, I920, on coniferous boards in the greenhouse at 
State College, A fifth collection, made in the Ottawa Valley, 
Canada, by R. J. Blair in 1917, is in my herbarium. Three of 
these collections were taken from structural coniferous timbers, 
four of them being from Tstiga canadensis. The specific iden- 
tity of the substratum of the fifth collection was not determined, 
except that it was recognized as a coniferous host. The first 
collection, made here in 1915, was determined by Dr. Burt. 

The species is one of the easiest of the stereums to recognize. 
The plants are almost entirely resupinate and the hymenium 
is in all collections a uniform cinnamon brown or Sudan brown, 
except that where the fresh specimens are wounded on the 
hymenium they immediately turn black and remain so on dry- 
ing. The hymenial surface is radiately lined with obtuse ribs 
in all cases. The pileus where developed is black on top in 
dried specimens and usually decidedly rugose. 

Spores have not been certainly seen; in one specimen, how- 
ever, small rounded hyaline spore-like bodies, 4-5 [x in diameter, 
were present on the hymenium but were not seen attached to 
basidia. No cystidia are present. Sections of the hymenium 
sometimes become pea green in KOH solution. 

7. Merulius fugax Fn 

This rare and interesting species was collected on a conif- 
erous board in the greenhouse at the Pennsylvania State College, 
on October 26, 1920, It is a bright colored species, my field 
notes recording a salmon buff color, varying to pinkish cinnamon 
for the hymenial surface, and with the margin considerably 
lighter. The specimen was rather young and the gyrose- 
plicate hymenium was developed only in the center of the fruc- 
tification. The plant is separable from the substratum when 
fresh but as soon as it becomes dry it adheres tightly, although 
the fructification is quite membranous in that condition. Micro- 
scopic examination of sections through it shows the character- 


Mycological notes for 1920 169 

istic features of the species, which are the oblong-ellipsoidal to 
subglobose spores and the large conspicuously incrusted hyphae 
of the subhymenium next the substratum. These hyphae are 
4-6 [X in diameter and possess both cross walls and clamps. The 
spores under the microscope are all but hyaline, although the 
species is included among the Leptospori by Dn Burt, who 
states that they sometimes appear slightly yellowish. No 
cystidia are present. 

Meruliiis siibaurantiactis Peck is recorded as a synonym by 
Dr- Burt, and my sections of the type of that species show struc- 
tures identical with those of M.fugaXj except that no incrusted 
hyphae are present next the substratum. Apparently no other 
species described from America has similar spores. 

8. SoLEXiA FAscicuLATA (Pers.) Fr. 

Plants rather crowded on decorticated wood, consisting 
of isolated cylinders not more than i mm. high and 0.2 mm. 
thick, whitish or slightly brownish, externally compactly tomen- 
tose; hymenium lining the hollow cylinders, of clavate basidia 
without cystidia; spores not seen attached to basidia but free- 
floating, hyaline, thin-walled, and 5-7 [jl in diameter. 

On coniferous w^ood. Reitz Gap, Center County, Pennsyl- 
vania, September 20, 1920, /. F. Adams. 

According to C. G. Lloyd, who made the determination, 
S. villosa Fr. is probably the same species. These specimens 
depart from the available descriptions of S, fasciculata, w^hich 
call for a plant 2-7 mm. high. The plants under the hand lens 
(Plate 9, fig. 3), resemble a simple Mucronella in which genus, 
however, the basidia are external. 



Collected on the banks of Stone Creek, Huntingdon County^ 
Pennsylvania, July 9, 1920. 

A rare species, listed by Murrill as occurring in New England 
and New York, The uniform pale indigo-blue color of cap, 
tubes, and stem is unique among the pore fungi. A single 
specimen was collected, from w^hich the following notes were made: 

Plant stipitate, more or less uniformly pale indigo-blue; 
pileus nearly orbicular, 4.5 x 3.5 x 0.3-0.7 cm., smoky indigo, 
glabrous, dry, slightly rugose; margin lobed, deflexed; context 
white, 1-5 mm. thick, taste mild; tubes about 2 mm. long, 
pale blue within, the mouths pale indigo-blue, angular, thin- 
walled, entire, averaging two per mm., decurrent on the stem, 

170 Mycological notes for 1920 

not separable from the context; stem excentric, nearly lateral, 
deep mdigo-blue, pallid within, glabrous, 2.5 cm. long, 0.5 cm. 
thick; spores subglobose, hyaline, 4-5 [x in diameter; cystidia 

The dried plant is dark bluish black on the pileus and stem, 

and the hymenium has become brown. 

10, Polyporus compactus sp. nov. 

Plants resupinate to narrowly reflexed or cushion-shaped, 
the pileus not more than i cm. broad, and 1.5 cm. thick, in re- 
supinate condition 3-8 mm, thick, white or gray in color, corky- 
watery when fresh, firm and compact when dry, glabrous or 
practically so, the margin rounded and obtuse; context light 
wood color, friable but compact w^hen dry, taste mild; tubes 
oblique, less than 2 mm. long, often lacking or poorly developed, 
the mouths subcircular, averaging three or four per mm., white; 
dissepiments rather thick and rigid; basidiospores ellipsoid or 
ovoid, often with a slightly truncate apex, hyaline, 7.5-9 x 4.5-6 
[x; chlamydospores (?) abundant on the hymenium and the 
hyphae of the context, subglobose to broadly ellipsoid, colorless 
or somewhat greenish or yellowish under the microscope, 7.5-9 
X 6-7.5 l^ or 6-7.5 ^ in diameter; basidia large, pyriform, 9-10 
jjL in diameter; cystidia mixed with the basidia as pointed, narrow, 
flask-shaped organs 3-4 [i in diameter; hyphae of two kinds: 
[a) large hyaline, unstaining hyphae, often thick-walled, abruptly 
breaking up at one end into several to many smaller branches 
that gradually taper out to the extremity, 3-5 [x in diameter in 
the larger portions, lacking cross w^alls and clamps; (&) small 
deeply staining and much branched hyphae bearing at least 
some of the chlamydospores, with a few cross walls and clamps, 

diameter 2-3.5 {^^ [Plate 9, figs. 5 and 6; Text figs. 9-14-] 

On dead standing trunks or on stumps of Quercus, especially 
Q. alba. 

Specimens are at hand as follows, the first mentioned being 
designated as the type: 

New Brunswick, New Jersey, on dead standing Quercus alba, 
August 25, 1920, Overholts& West, No. 7873; Lamar, Clinton 
County, Pennsylvania, on dead standing Quercus alba, July 26, 
1920, Overholts Herb. No. 7525; State College, Pennsylvania, on 
oak stump, October, 1920, No. 7173; New Brunswick, New 
Jersey, on dead Quercus alba, August 21, 1918, E. West J4J, 
No. 7584; New Brunswick, New Jersey, on dead standing oak 
trunks, June 16, 1919, E. West 414, No. 7526. 

After this manuscript was submitted for publication, the 
specimens that had been previously designated as the types and 
from which the photographs for this article were made, were 

Mycological notes for 1920 



destroyed by a fire that burned the photographic studio where 
the collection was kept at the time. This has necessitated 
designating my herbarium No. 7873, as the type of the species 
rather than No. 7525, from which the photographs were made, 
and which is therefore no longer in existence. This will explain 
any discrepancy that might appear in the failure, in later years, 
to find a collection in my herbarium with the exact aspect 
presented in the photographs. 

Figs. 9-14. Polyporus compactus Overholts 

9. Small portion of the hymenium showing basidla, cystidia, and attached 
spores. 10. Mature basldiospores. 11, Branched hypha. 12. Chlamydo- 
spores. 13, Production of chlamydospores. 14. Hypha with clamp con- 

Of the collections here cited, only the first and the last named 
show basidia and basidiospores, while all bear the chlamydospore 

The real status of the so-called chlamydospore condition has 
not been determined, as no cultures have been attempted. That 
the knob-like grow^ths as shown in Plate 9 involve this species 
can scarcely be questioned- It has' been suggested that these 
growths may represent a diseased condition and that the bodies 
here designated as chlamydospores may be in reality the spores 

172 Mycological notes for 1920 

of the parasite. They are not produced in chains but formed 
singly as the termini of smallhyphaedeeply staining with eosin. 
This fungus has' been under careful observ^ation for sev- 
eral years, but only the chlamydospore condition was known 
until 1920. A well-developed pileus is never present. In some 
respects the species show^s resemblance to P. EUisiantis Murrill, 
which however is found on coniferous wood and has entirely 
different spores. The cushion-shaped or nodular type of sporo- 
phore, bearing only the chlamydospore condition, has the context 
more or less zoned and with a disagreeable odor as of strong lard. 

II. FoMES Bakeri Murrlll 


I found this species to be quite common, by far the most 


observed it in comoanv with Mr. E. West 

Its only host in that locality is the river birch, Betula nigra, 
which is its host also in the Mississippi Valley. The species 
was described and illustrated in my notes for 1919, where 
reference was made to a collection said to have been taken from 
Betula lutea near Philadelphia in 1915 by Rhoads. In all prob- 
ability the host for this collection was B. nigra, as B. lutea is 
not found in that locality, according to Dr. Rhoads. 

In connection with this species it was of interest to find 
recently in the Schweinitz Herbarium at Philadelphia a specimen 
of Fomes under the name of JPolyporus annosus Fr. (Syn. N. Am. 
Fungi No, 401) that is undoubtedly F. Bakeri, although it seems 
not to have been recognized by those who have worked over 
the Schwelnitzlan material in recent years. Lloyd once referred 
this specimen to F. rimosus, noting, how^ever, its similarity to 
F. robustus Karst., which he has subsequently regarded as identical 
with F. Bakeri. This Schweinitzian collection bears no date but 
was made certainly prior to 183 1 and was listed by Schweinitz 
as occurring on Betula. 

12. Phallogaster saccatus Morgan 

A rare plant, placed in the Phallaceae by some largely on the 
foetid nature of the internal deliquescent gleba, but in appearance 
more allied to the Lycoperdaceae, Two specimens were collect- 
ed at Westport, Pennsylvania, August 11, 1920. Excellent 


been published by Lloyd. 

Mycological notes for 1920 173 

Explanation of plate 9 


Fig. I, Enlarged photograph of a ^mall area of the resin exudate on which 
Zyihia resinae (Ehrenb.) Karst. and Biatorella resinae (Fr.) Mudd, grow side 
by side. The elongate bodies in the left hand corner are the pycnidia of the 
Zythia, while the larger disk-shaped bodies to the right are the apothecia 
of the Biatorella. Photographed with Micro-Tessar 10 X lens, by E. T. Kirk. 

Fig. 2, Resin exudate inhabited by the pitch midge and bearing the 
fruiting bodies of Zythia resinae and Biatorella resinae, X i- Photo by the 

Fig. 3, Solenia fasciculata (Pers.) Fr,, showing the isolated white-tipped 
cylinders in which the hymenium Is produced. Photographed with Micro- 


Tessar 10 X lens, by C. B. Neblette. 

Fig. 4, Pilacre Peter sii B. & Br., natural size, Photo by the writer. 

Fig. 5, Rounded abnormal growth of Polyporus campadus Overholts, 
as usually found on vertical surfaces, and bearing only the chlamydospore 
condition, X I. Photo by the writer. 

Fig. 6, Poroid form of Polyporus compadus, showing the largely resupi- 
nate manner of growth, X l- Photo by C. B. Neblette. 

Bull. Torrey Club 

Volume 49, plate 9 






f. POT vpoRTJc; rOMPACTUS Overhol 

Unreported plants from Glacier National Park 


Paul W. Git\FF 

Our national parks are each season attracting a greater 
number of vacationists drawn to them by the beauties of nature, 
some only by the great attractions of mountain scenery but 
many by all the varying phases of nature to be enjoyed in these 
national playgrounds. To the person of botanical or zoological 
interests Glacier Park is one of the most attractive. Its high 
rugged peaks, deep valleys, numerous glaciers and lakes, streams 
and cascades offer a great amount of variation. The Continen- 
tal Divide passes through the Park with the result that the 


drainage of the w^estern portion is into the Columbia River, 
while the northeastern part slopes toward the Hudson Bay, and 
the eastern into the Missouri River drainage, A greater amount 
of variation will be found here and with less sophistication than 
in any other of our park areas. 

In the herbarium of the University of Montana Biological 
Station at Yellow Bay are a number of plants which have been 
collected at the Park. Among these are a number which have 
not as yet been reported from that region even in Standley's 
recent publication.* The present list is offered as an addition 
and slight contribution to the knowledge of the flora of this region. 
Of the thirty- two species here listed twelve have not before been 
reported from the state of Montana. The names of these 
twelve are preceded by a dagger (f). Two of these, Car ex Hele- 
onastes Ehrh., and Salix reticulata L., have not, so far as I am 
aware, been reported from the United States. They are Canadian 
species which have entered this country in the high mountains 
of this region, both being extreme northern forms. 

t I. Danthonia piketorum Piper, Fl. Northw. Coast 46. 1915. 
D, spicata var pinetorum Piper, Erythea 7: 103. 1899. D, 
thermalis Scribn, U, S. Dept. Agr. Div. Agrost. Circ. 30: 
5. 1901. 
Mineral Park, August 10, 1 910, M, E. Jones, at 1800 meters 


This is a northwest coast plant which has been reported as 

♦Flora of Glacier National Park. Contr. U. S. Nat. Herb. 22: 235-438 

pi' 33-^52^ 1 92 1 


176 Unreported plants from Glacier National Park 

being found to the eastward In Wyoming and Idaho, but not as 
yet in the state of Montana. 

2. Poa Cusickii Vasey, Contr. U. S. Nat, Herb. 1: 271. 1893. 

Mineral Park, August 8, 1910, M. E, Jones, at 2400 meters 


Reported from British Columbia, south to Oregon and east- 
ward through Idaho and western Montana. 



3. PoA nevadensis Vasey, Bull. Torrey Club 10:66. 1883. 
Greenwood's Camp, August 15,- 1910, M, £. Jones, at an 

elevation of 1350 meters. 

Reported from British Columbia southward to Arizona, 
and eastward to North Dakota and Colorado. 

4. Bromus eximius (Shear) Piper, Contr. U. S. Nat. Herb. 11: 

143. 1906. B. vulgaris var. eximius Shear, U. S. Dept. 
Agr. Div. Agrost. Bull. 23: 44. 1900. 

Greenwood's Camp, August 15, 1910, M. E. Jones, at 1350 
meters elevation. 


This Is primarily a Pacific coast plant, being found from 
British Columbia southward to CaUfornia. To the eastward 
of this range it Is found In the states of Montana and Wyoming. 

5. Eriophorum polystachyon L. Sp. PL 52. 1753, 

Mineral Park, August 12, 1910, M. E. Jones, at 1800 meters 

This is primarily a Canadian species which has been found 
to the southward in the Rockies and In the North Atlantic States. 


165. 1903. 



meters elevation. 

A Pacific coast and Canadian species, also reported southward 
in the Atlantic States. 

7. CARExDEWEYANASchwein, Ann. Lye. NewYork 1:65. 1824. 
Waterton Lake, August 17, 1910, M. E, Jones, at 1200 
meters elevation. 

This is a northern species with a range extending southward 
in both the Appalachians and Rockies. It has been reported from 

Montana, but not from the Pprlr rpainn 

Unreported plants from Glacier National Park 177 

8. Carex festiva Dewey, Am. Jour. Sci. 29: 246. 1835. 

Sperry Glacier, August 27, 1909, M. E. Jones, at 2250 

meters elevation; Gnnnell Lake, August 10, 1911, M. J. Elrod, 
at 2100 meters elevation. 

Found from California to the Arctic regions on the coast, 
and in the Rocky Mountains 

t9. Carex foenea Willd. Enum. 957. 1809. 

Waterton Lake, August 17, 1910, M. E. Jones, at an eleva- 
tion of 1200 meters 

This is primarily a Canadian species which has crept down 
into our range. Its range is from New Brunswick to Penn- 
sylvania, thence northwestward to Minnesota, British Columbia, 
and the Yukon. 

10. Carex Goodenovii J. Gay, Ann. Sci. Nat. II, 11 : 191. 1839. 
C. caespitosa Gooden., Trans. Linn. Soc. 2: 195. pL 21 j 1794. 

Not Linn. 1753. 
Lake Mc Donald, August 2, 1910. M, E. Jones, at 900 meters 

elevation. Mineral Park, August 10, 1910, M. -E. Jones, at 
an altitude of 1800 meters. 

Found from Newfoundland and Pennsylvania westward to 
Alaska and Colorado. 

t II. Carex Heleonastes Ehrh.; L. f. Suppl. 414- 178 1. 

Sperry Glacier, August 2^, 1909, M, E, Jones^ at an altitude 
of 2250 meters, growing in a damp locality among moss. 


While reported from the Upper Temperate and Alpine 
Life Zones in Canada, from Ontario and the Hudson Bay region 
westward to British Columbia, this species does not seem to 
have been reported from this side the "Border. " It is essentially 
a northern plant which has found its way into the United States 
in the high altitudes of the Park. 

ti2. Carex hormathodes Fernald, Rhodora 8: 165. 1906. 
Blackfoot Glacier, August 30, 1909, M. E. Jones, at 2100 

- 4i 

meters elevation. 

Reported previously from British Columbia, but not from 

the northwestern United States. 

ti3. Carex monile Tuckerm. Enum. Meth. 20. 1843. 

Grinnell Lake, August 20, 1911, M, J. Elrod, at 1500 meters 


This differs from C. vesicaria L., with which it is sometimes 
confused, in the possession of three stigmas rather than two 
and in the presence of more than one staminate spike. The 
variety colorata Bailey has been reported from Yellowstone 

178 Unreported plants from Glacier National Park 

Park, but the typical form of the species has not been reported 

from Montana. 

Reported south of our range in Wyoming, and across the 

United States. It is also found along the Pacific coast from. 

California to Alaska. 

ti4. Carex polygama Schkuhr, Riedgr. 1: 84, 1801. 

Waterton Lake, August 17, 1910, M. £. Jones^ at an elevation 
of 1200 meters. 

Reported well to the northward on both coasts, and across 
the United States from California through Utah, Colorado, and 
Missouri to the Atlantic coast. It is a southern species which 
has found a congenial habitat to the northward on both sea- 

fiS- Carex saxatilis L. Sp. PI. 976. 1753. 

• Bowman Lake, August 19, 1910, M, E. Jones, at 1200 meters 


C. saxatilis is an arctic species found across North America 

from Labrador to Alaska. It has been reported as entering 

the United States in the mountainous portion of Maine, but 

has not been reported from the western states except in the 

high mountains of Colorado. It is essentially an arctic plant, 

and is found also in the extreme northern portion of Europe and 

b _ 

16. Salix orbicularis Anders.; De Candolle, Prodromus 16^; 

300. 1868. 

Mountain, August, ion, M. 


meters elevation. 

Reported previously from the Sperry Glacier region under 
the name S, reiictdata L. . 


This is an extreme northwestern species found from Alaska 
to Hudson Bay, and southward into the United States in the 
Rocky Mountains. 

ti7. Salix reticulata L. Sp. PL 1018. 1753. Not Porter & 

Blackfoot Glacier, August 31, 1909, M. E. Jones , at an ele- 
vation of 2100 meters. . - 

This species has been considered previously as purely Cana- 
dian. It has been reported from the Canadian Rockies and 
northward to Alaska and Hudson Bay. Salix saximontana 
Rydb. has been reported from this region w^ith S. reticulata L. 


Unreported plants from Glacier National Park 179 

as a synonym, an error for S. reticulata Porter & Coulter, a 
quite distinct species. 

18. Eriogonum ovalifolium Nutt. Jour. Philad. Acad. 7; 
50. pi. 8. 1834. 

Brown's Pass, August 13, 1911, M. 7. Elrod,a,t 2580 meters 

A western species found from New Mexico northward through 
California and Colorado to British Columbia, 


19. Polygonum bistortoides Pursh, var. linearifolium 

(Wats.) Small, Bull. Torrey Club 19: 252. 1892. 
Iceberg Lake, August 18, 191 1, M. J. Elrod, at an elevation 
of 1800 meters. 

Found from - New Mexico northward within the Rocky 
Mountains through Canada to the Arctic regions. 

I - F - 

20. SiLENE DouGLASii Hook. Fl. Bor. Am. 1: 88. 1840. 
Sperry Glacier, August 2^, 1909, M,.E. Jones, at 2250 meters 


Rather a common plant in the Rockies, and on the coast 
from Washington to California. 

21. Arenaria subcongesta (Wats.) Rydb., Bull. Torrey Club 
24: 244, 1897. 

Brown's Pass, August 13, 1911, M. J. Elrod, at 2580 meters 

This species is common in the Rocky Mountain region. It 
seems to be the same as A. lithophila Rydb. and A. Biirkei 
Howell and should possibly be considered only a variety of 
A. conges ta Nutt. 

22. Aragallus monticola (Gray) Greene, Pittonia 3: 212. 


Mineral Park, August 8, 19 10, M. E. Jones, at 2100 meters 

elevation; Brown's Pass, August 13, 1911, M. J. Elrod, at 2580 
meters elevation. 

Not uncommon in the mountains from Wyoming northward. 

23. PoLEMONiUM coNFERTUM Gray, Proc. Acad. Philadelphia 
1863:73. 1864. 

Brown's Pass, August 13, 1911, M. J. Elrod, at 2580 meters 


A high elevation plant found in the mountains of Utah, 

Colorado, Wyoming, and Montana; occasionally in northern 
New Mexico and California. 

i8o Unreported plants from Glacier National Park 


24. Phacelia Franklinii (R. Br.) Gray, Manual, Ed. 2, 329. 


Mineral Park, August 8, 1910, M, E, JoneSj at 2400 meters 
elevation; Brown's Pass, August 13, 191 1, M, /. Elrod, at 2580 

meters elevation. 

Wyoming, and Michig 

to Hudson Bay and Alaska. 

t 25. Lappula cill\ta (Doi 





26. Pentstemon Menziesii Hook. Fl. Bor. Amer. 2: 98. 1840- 
P.fruticosus (Pursh) Greene, Pittonia 2: 239. 1892. 
Sperry Glacier, August 27, 1909, M, E, Jones, at 2250 meters 

elevation; Brown's Pass, August 13, 1911, M. J. Elrod, at 2580 

meters elevation. 

Found from California and Colorado northward to British 
Columbia and Alberta. 

t27. Veronica Cusickii Gray, Syn. Flor. 2: 288. 1878- 

Mineral Park, August 9, 1910, M. E, Jones, at 1800 meters 
elevation . 

Previously reported only from the high mountain regions of 
the three states, Washington, Oregon, and Idaho. 

28. Valeriana occidentalis Heller, Bull. Torrey Club 25: 
269. 1898. V. sylvatica Banks; Richards, Frankl. Journ. 
Ed. 2, Appendix 2. 1823. Not Schmidt, 1795. V.wyomtng- 

ensis E. Nels. Erythea 7: 167. 1899. 

meters elevation. 

M. E. Jones, at 2100 



to Colorado and Utah. 


29. SoLiDAGO PuRSHii Porter, Bull. Torrey Club 21:311. 1894. 
S. hiimilis Pursh, FI. Am. Sept. 543. 1814. Not Mill. 


meters elevation. 


Rocky Moiun tains from New Mexico 

Columbia and northward. 


Unreported plants from Glacier National Park i8i 

30. Erigeron grandiflorus Hook. FI. Bor. Am. 2; 18. pL i2j. 

Sperry Glacier, August 27^ 1909, M. E. Jones, at an eleva- 
tion of 2250 meters. 

Reported from Colorado to the extreme north; more es- 
pecially in the high elevations of the Canadian Rockies. 

t 31. Antennaria alpixa (L.) Gaertn. Fr. etSem.2;4io. 1791. 

Gnaphalium alpinum L. Sp. PI. 856. 1753- 

This IS essentially a Canadian species found across the con- 
tinent from Labrador to Alaska, but extending southward into 
the United States in the Rockies and Sierras. 



32, Senecio aureus L. sp, PL 870. 1753. 

Brown's Pass, August 13, 1911, M. /. Elrod, at 2580 meters 
elevation; Iceberg Lake, August 18, 19x1,^. J. Elrod, at 1800 

meters elevation. 

This species is close to Rydberg's S. crocatus of Colorado, 
Wyoming, and the northern Rockies. Its range is from Alaska 
to California, and across the continent. 

University of Montana Biological Station, 
Yellow Bay, Montana 

Two new plants from western Colorado 


Geo. E. Osterhout 

^ Nuttallia marginata sp. nov. 

Seemingly a biennial, the stem 3 dm. or more high, white^ 
smooth below, roughish pubescent above, the branches alternate, 
the lower ones elongate; the lower leaves oblanceolate, sinuate 
dentate, 6-8 cm. long, 5-7 mm. wide, seesile by a narrow base, 
the upper leaves smaller, coarsely dentate, sessile, all of them 
roughish with a barbed pubescence ; the flowers yellow, numerous in 
a cyme of alternate branches, the whole flower — capsule and petals 
2 cm. long, sometimes subtended by a narrow entire bract, five 
outer petals 13 mm. long, the claws short and broad, the upper 
portion ovate, pointed, the outside pubescent, five inner petals 
smaller, spatu'ate; the capsule 8-1 1 mm. long, rounded at the 
base, harshly pubescent; the sepals lanceolate, 7 mm. long, 
pubescent like the capsule; the style 10 mm. long, stigmatic 
at the tip; seeds many, slightly over 2 mm. long, 1.5 mm. wide, 
the margin very narrow. 

The only locality where I have collected this Nuttallia is 
De Beque, in Mesa County, on the hills north of the town. 
The type specimens (No. 5842) were collected August 22, 1918, 
The species begins to blossom near the first of June. It is peculiar 
In Its narrowly margined seeds and pubescent petals. 

■^Acrolasia humilis sp. nov. 

Annual, I-I.5 dm. high, the stem white, smooth below, 
pubescent with slender barbed hairs above, the first branches 
from near the base, but the branches few and distant; the low- 
est leaves spatulate, on short petioles, 3-5 cm. long, 5 mm. or 
more wide, the succeeding leaves sessile, becoming wider and 
ovate above, about 2 cm. long, all entire and pubescent beneath, 
glabrous above or nearly so; the first flowers sessile in the axil 
of the branches, the later ones clustered and sessile at the ends 
of the branches, the petals five, yellow, broadly obovate, veined, 
2.5-3 nim. long, the calyx lobes linear, 2 mm. long; the stamens 
ten to fifteen, the filaments slender, the anthers globose; the 
capsule becoming about i cm. long, 3 mm. wide, rough with 
small barbed pubescence, and longer slender hairs not barbed; 
the seeds prismatic and minutely muricate. 



I also collected it in the vicinity of Eckert, in Delta County. 
On account of the entire leaves it rather resembles Acrolasia 


1 84 


latifolia Rydb. or A. dispersa (Wats.) Rydb., but it really be- 
longs with A. albicaulis (Dougl.) Rydb. The seeds are similar 
to the latter, and are in the capsule in three rows, not in a single 


Windsor, Colorado 



The aim of this Index Is to Include all current botanical literature written by 
Americans, published In America, or based upon American material; the word 
America being used In the broadest sense. 

Reviews, and papers that relate exclusively to foresty, agriculture, horti- 
culture, manufactured products of vegetable origin, or laboratory methods 
are not included, and no attempt Is made to index the literature of bacteriology. 
An occasional exception Is made in favor of some paper appearing In an Ameri- 
can periodical which is devoted wholly to botany. Reprints are not mentioned 
unless they differ from the original in some important particular. If users 
of the Index will call the attention of the editor to errors or omissions, their 
kindness will be appreciated. 

This Index is reprinted monthly on cards, and furnished in this form to 
subscribers at the rate of three cents for each card. Selections of cards are 
not permitted; each subscriber must take all cards published during the term 
of his subscription. Correspondence relating to the card issue should be 
addressed to the Treasurer of the Torrey Botanical Club. 

Gardner, M. W-, & Kendrick, J. B. Turnip mosaic. Jour. 
Agr, Research 22: 123, pi 20 15 O 192 1. 

Garrett, A. O. Some introduced plants of Utah. Torreya 21: 

76-79, O 1921. 

Gatin, C. L. (Premiere contribution a I'etude) De Tembryon 
et de la germination des Aracees. Ann. Sci. Nat. Bot- X. 3: 
145-180. pL i-io. 'N 1921. 

Gericke, W. F. Root development of wheat seedlings. Bot; 

Gaz. 32: 404-406./. /. 15 D 1921. 

Gleason, H. A. Botanizing in British Guiana. Jour. New 

York Bot. Gard. 22: 161-168. S 1921. 


Gleason, H. A. A rearrangement of the Bolivian species of 

Centropogon and Siphocampylus. Bull. Torrey Club 48: 
189-201. 26 S 1921. 

Includes 2 new species of Siphocampylus. 

Goldring, W. Annual rings of growth in Carboniferous wood. 

Bot. Gaz. 72: 326-330. pi 14, 15 N. 1921. 

Goss, R* W. Temperature and humidity studies of some 

Fusaria rots of the Irish potato (Solanum tuberosum), Joui , 
Agr. Research 22: 65-79 pL 10. 11. 8 O 1921. 


1 86 Index to American botanical literature 

GriflEee, F. Comparative vigor of F2 wheit cDs.ei an:! their 
parents. Jour. A;jr. R-search 22: 53-63- 8 O 192 1. 

Grimes, E. J. A new station for Pogonia affinis. Rhodora 

23: 195-197. 15 D I92I, 

Guppy, H. B. America's contribution to the story of the plant- 
world. Jour. Ecol. 9: 90-94. S 192 1. 

Harlan, H. V., & Pope, M. N. Ash content of the awn, rachis, 
palea, and kernel of barley during growth and maturation. 
Jour. Agr. Research 22: 433-449. f-iS- 19 N 192 1. 

Harms, H. Drei neue Legumlnosen aus Venezuela. Notizbl. 
Bot. Gart. u. Mus. Berlin-Dahlem 8: 51, 52. I S 1921. 

Species of Calliandra (i), Pilhecolobium (i), and Piptadenia (l). 

Harms, H. Kakteen als Wirtspflanzen. Monatschr. Kakteenk. 
31: 184-186. D 1921. 

Harms, H. Kakteen und Sigillarien, Monatschr, Kakteenk. 

31: 145-148. O 1921. 

Harper, R. M. Cape Cod vegetation. Xorreya 21: 91-98. 

D 1921. [Illust.] 

Harrington, G. T. Optimum temperatures for flower seed ger- 
mination. Bot. Gaz, 72: 337-358. /. i-io. 15 D 192 1. 

Harris, J. A., Sinnott, E. W., Pennypacker, J. Y., & Durham, 

G. B. The vascular anatomy of hemitrimerous seedlings of 
Pliaseohis vulgaris. Am. Jour. Bot. 8; 375-381. 14 N 192 1. 

Harris, J. A., Sinnott, E. W., Pennypacker, J. Y., & Durham, 

G. B. The interrelationship of the number of the two 
types of vascular bundles in the transition zone of the 
axis of Phaseolus vulgaris. Am. Jour. Bot, 8: 425-432. 
19 D 1921. 

Harshberger, J. W. The artistic anatomy of trees. Nat. Hist. 

21: 387-397- O 1921. [Illust.] 

■ , 

Harter, L. L., & Weimar, J. L. A comparison of the pectinase 

produced by different species of Rhkopus. Jour. Agr. Re- 
search 22: 371-377- /- ^. 2. 12 N 1921. 

Harter, L. L-, Weimar, J. L., & Lauritzen, J. I. The decay of 

sweet potatoes {Ipomoea batatas) produced by different 
species of Rhizop us. Phytopathology 11: 279-284. 15 N 
192 1. 

Index to American botanical literature 187 

Hastings, G. T. The forest flora of Grassy Sprain Ridge [New 

York], Torreya 21: 73-76. O 192 1. 


Hastings, G. T. The boy scouts and conservation of wild 

flowers. Torreya 21; 83, 84. O 1921. 

Hauman, L. Deux graminees geantes de la flore argentine. 

Physis 5: 52-56. 31 O 192 1. 

Includes Sporoholus maximus, sp. nov. 

Hauman, L,, & Parodi, L. R. Los parasitos vegetales de las 

plantas cultivadas en la Republica Argentina. Revista 
Facult. Agron. y Veter. 3: 227-274. /. 1-4, D 192 1. 


Hawkins, L. A. A physiological study of grapefruit ripening 

and storage. Jour. Agr. Research 22: 263-279./. J. 29 O 

Heald, F. D. The relation of spore load to the per cent of stink- 
ing smut appearing in the crop. Phytopathology 11: 269^ 
278. 15 N 1921. 

Heller, H. H. Phylogenetic position of the bacteria. Bot. 

Gaz. 32: 390-396. 15 D 1921. 

Heller, H. H. Suggestions concerning a rational basis for the 

classification of the anaerobic bacteria. Jour. Bact. 6: 
521-553. N 1921. 

Studies in pathogenic anaerobes IV. 

Hoerner, G* R. Miscellaneous studies on the crown rust of 

oats. Am. Jour. Bot. 8: 452-457. pL 24. 19 D 192 1. 

Holm, T. Morphological study of Carya alba and Jiiglans 

nigra. Bot. Gaz. 32: 375-389. p!. 75, 16 + /. J. 15 D 

Holm, T. Recent botanical publications from the United 

States National Museum. Am. Midi. Nat. 7: 165-180. 
N 1921. 

Considerable original matter contained in a review. 

Hopkins, E. F. Studies on the Cercospora leaf spot of bur 

clover. Phytopathology 11: 311-318. pL j, 14 + /. «^, 9- 
9 D 1921. 

Howard, G, E. Extraction and separation of the pigments of 

Nereocystis Luetkeana, Publ. Puget Sound Biol. Sta. 
3: 79-91. ph 14. I O 1921. 

i88 Index to American botanical literature 

Howe, C. G. Pectic material in root hairs. Bot. Gaz. 72: 
313-320. 15 N 1921. 


Hubert, E. E. Notes on sap stain fungi. Phytopathology 11: 
214-224. pi 7 + f. 1-4' 6 1 92 1. 

Huhnholz, P, Echinocactus Anisitsii K. Sch. Monatschr. 
Kakteenk. 31: 150, 151. O 1921. 

niick, J. S. The American walnuts. Am. For. 27: 699-704, 
N 1921. [Illust.] 

InmaDj O. L. Comparative studies on respiration XX, The 
cause of partial recovery. Jour, Gen. Phy§ioL 4: 1 71-175- 
/. r, 2, 20 N 1921. 

Jackson, H. S., & Mains, E. B. Aecial stage of the orange 

leafrust of wheat, Puccinia iriticina Eriks. Jour. Agr. 
Research 22: 151-172, ph 21 + /. J. 15 O 1921. 

* w 

Jeffrey, E. C, The geographical distribution of hybrids. 
Science II. 54; 517. 25 N 1921. 

Johnson, D. S. Invasion of virgin soil in the tropics. Bot- 
Gaz, 72: 305-312. /. j, 2. 15 N 1921. 

Johnson, D. S. Polypodium vulgare as an epiphyte. Bot, 
Gaz. 72: 237-244. /. j-j. 15 O 192 1, 

Jones, F. R., & Tisdale, W. B. Effect of soil temperature upon 

the development of nodules on the roots of certain legumes. 


/. 1-4. I O 1921- 

Jones, L. R., & Doolittle, S. P. Angular leaf-spot of cucumber. 

Phytopathology 11: 297, 298. 15 N 1921. 

Jones, L. R., & Keitt, G. W. Eugene Washburn Roark. Phyto- 
pathology 11: 327. 9 D 192 1. 

Kempton, j. H. Heritable characters of maize. HI. Bra- 

chytic culms. V. Adherence. Jour. Heredity 11: iH" 
115./. 12. 6 My 1920; 11: 317-322./. 16-19. 23 Mr 1921. 

Kempton, J. H. Heritable characters of maize. VHI. White 

sheathes. Jour. Heredity 12: 224-226. /. 18. 23 S 1921- 

Kendrick, J. B., & Gardner, M. W. Seed transmission of soy- 
bean bacterial blight. Phytopathology 11: 340-342- P^- 
17. 9 D 1921. 

Vol. 49 

No. 7 




JULY, 1922 

New species of Uredlneae — ^XIV 


Joseph Charles Arthur 

In the tenth article in this series of new species an error 
was made in the determination of the host for Uromyces fuscains 
(BuIL Torrey Club 45: 142. 1918), which should be Rumex 
paucifolius Nutt., and not ''Polygonum alpinum AIL" as 
printed. It was due to the persistent efforts ol 



Pentstemon (see Fungi Utahenses 244)^ that the facts in the case 
were finally established.- In July, 1920, Mr. Garrett visited 



he Rumex paucifolius . Upon my presenting the situation to 
Professor Aven Nelson, he looked up the field notes and the 
herbarium specimens collected in Idaho at the same place and 
time as the type of U. fuscatus and came to the conclusion that 
the host of this collection also was R. paucifolius. Only these two 
localities for the species are known up to the present time. 
The correction of these errors was complicated by a similar 
uncertainty and error in the host names for Puccinia Polygoni- 
alpini Cruch. & Mayor (Dicaeoma Polygoni-alpini Arth. N. Am. 
Flora 7:385. 1920), for which only tAvo collections were known 
at that time, a third one having been added from Greenville, 
California, since the printing of the Flora. Rumex paucifolius^ 
host of the UromyceSj can not be told from Polygonum alpinum^ 
host of the Pucciniaj when the fruiting parts are absent, hence 
the difficulties that have arisen. 


Contribution from the Botanical Department of Purdue Universitj 
Agricultural Experiment Station- 
[The Bulletin fo/ June (49: 163-188. pL g) was issued July 8, 1922 ] 


190 Arthur: New species of Uredineae — XIV 

In the twelfth article of this series, under Puccinia offuscala 
(Bull Torrey Club 47: 469. 1920), the name Uredo Zorniae 
Dietel (Hedwigia 38: 257. 1899) should have been given as a 
synonym. This name was applied to Ule's 22q6 from BraziL 
Although pycnia have not been seen yet there is circumstantial 
evidence to indicate that no aecia are produced, and therefore 
the species Is probably a Bullaria^ and should be called B. 
Zorniae (Dietel) comb. nov. It is a specie's of the New World, 
to which the name Puccinia Zorniae McAlpine does not apply* 

In the same article, under Pucciniastrum americanum {Bull. 
Torrey Club 47:468. 1920), Ruhus occidentalis was erroneously 
given as a host for this species. The error was called to my 
attention by Dr. J. J. Davis of the University of Wisconsin, 
and was confirmed by subsequent examination under the 
microscope. The type of P. arcticum americanum Farl , is on Ruhus 
neglectus Peck, collected at Bussey, near Cambridge, Massachu- 
setts. This collection is first mentioned in the original article 
by Farlow (Rhodora 10: 16. 1908) as on R, occidentalis near 
Cambridge, but later in the article is referred to as on R. neglectus, 
Bussey, October, 1875. The correction in the name of the host 
is credited to M. L. Fernald. I have been able to ascertain these 
facts through the assistance of Dr. R. Thaxter, who has carefully 
examined the original material in the Farlow Herbarium. 

Melampsoropsis roanensis sp. nov. 

IL Uredinia hypophyllous, in crowded groups of 2-8 on 
somewhat discolored spots, roundish or ovoid, 0.3-0.8 mm.long, 
prominent, early dehiscent by a central or lateral slit, leaving 
a border or a cap-like covering of epidermis, pulverulent; 
peridium delicate, uniform in thickness; urediniospores catenulate, 
ellipsoid, 20-26 by 32-40 yi; wall colorless, 1.5-2.5 ^ thick, closely 
and finely verrucose on one side grading to coarsely verrucose 
on opposite side with somewhat deciduous cylindric tubercles. 

2 p. long. 

gular and often confluent groups 


axy, orange-red, tardily naked; teliosp 

16-20 ^ broad, almost or quite as long, in a series 65-100 
long; wall colorless, thin, i tx, smooth; contents orange-re 


Michx. (Ericaceae) 


On Rhododendron punctatum Andr., LeConte Mountain 
'ier County, Tennessee, altitude 6600 feet, Tune 8, 1921 

Arthur: New species of Uredineae — XIV 19 1 

11, in, H. F, Bain, communicated by John A. Stevenson 5812 

The urediniospores of this interesting species are much 
larger and more prominently verrucose than those of the common 
form in Europe, AI, (Chrysomyxa) Rhododendri, This is the 
first rust on Rhododendron seen from the eastern United States, 
although the European species has been ascribed to this country 


by a number of authors. Duggar says (Fungous Diseases 432, 


1909), *'In the United States this fungus is particularly common 
in the mountains of the east, and southward as far as the southern 
limits of the Appalachians." The statement does not apply to 
any true rust on Rhododendron, Roan Mountain and LeConte 
Mountain are about 100 miles apart. The rust was reported 
abundant on the summit of LeConte Mountain.. Mycologists 
who visit the summits of these mountains and those nearby 
should be on the lookout for the aecia, which may be expected 
to occur on species of Picea, 

Cronartium stalactiforme Arthur & Kern, comb. nov. 

Peridermmm stalactiforme Arth. & Kern, Bull. Torrey Club 
33: 419. 1906. 

This Cronartinm belongs to a group of three species which 
have been much confused, both in nomenclature and in structural 
characters. All three species have uredinia and telia on Castilleja 
and related genera of scrophulariaceous hosts, which are difficult 
to distinguish except by means of cultures. The aecia of the 
three species, however, occuring on pine trunks and branches, 
can be identified with considerable certainty, even by their gross 
appearance. The aecia of C. stalactiforme are confined to the bark 
and do not produce woody galls. They often extend over con- 
siderable areas, and cause little or no swelling. The individual 
aecia arc low and flattened, with a circumscissile deliiscence. 

filamentosum (Peck) 



upon the host, but the individual aecia are more elongated, 
often becoming cylindric, and have noticeable filaments extend- 
ing from top to bottom through the individual aecia. There are 
only rudiments of such filaments in C. stalactiforme. 

Cronartinm coleosporioides (Diet. & Holw.) Arth. {Perider- 
miuni Harknessii Moore, 1876) differs from the two preceding 
species by producing a woody gall, often of considerable size and 

192 Arthur: New species of Uredixeae — XIV 

abruptly rising from the healthy tissue. The individual aecia are 
low and coalesce into a more or less continuous mass, and are 
entirely without internal filaments. 

There has been much confusion regarding the application of 
the name Peridermium Harknessii, owning in the first place to 
the obscure way in which it was first published*, and in the 
second place to the later inclusion of the Pacific Coast forms of 
the very similar P. Cerebrum, whose uredinia and telia occur 

on leaves of oak. 

The name was announced at a meeting of the San Francisco 
Microscopical Society in July, 1876, by J. P. Moore, who 
exhibited specimens and explained the nature of the fungus, 
and who read a letter from H. W. Harkness describing the 
species in an informal but accurate manner. The collection to 



* Our attention was directed to the early history of this name by Mr. E. 
Bethel in a letter dated January 22, 1922. Through the assistance of Mr. 
W, M. Hepburn, of the Purdue University Library, and Professor W. C. 
Blasdale, of the University of California, the following facts have been estab- 

In the minute book of the San Francisco Microscopical Society this re- 
cord occurs: "July 20, 1876. Mr. J. P. Moore read a paper written by Dr. 
H. W. Harkness on a new variety of fungus infesting Pinus ponderosa near 
Colfax, and he proposed "the name Peridermium Harknessii, which was 
adopted." There is also in the archives of the society a newspaper clipping 
giving a more extended account of the matter as presented to the society. 
This clipping, Professor Blasdale has ascertained, came from the issue of the 
Daily Alta California of San Francisco, of Saturday, July 22, 1876, volume 
28. It consists of the Harkness paper in full, and a paragraph stating how 
Moore proposed the name. 

The society did not issue any printed account of its proceedings during 
this period of its existence, but occasionally, as the subject matter warranted, 
the secretary transmitted a more or less formal minute to the London Monthly 
Microscopical Journal. In the issue of that journal for September I, 1876, 
an account of the meeting of July 20 is given in the same words used in the 
newspaper clipping, with three additional paragraphs. 

What is referred to as a paper by Dr. Harkness is ih the form of a letter 
to the secretary of the society. It begins with the sentence: "I have today 
forwarded for the Society's Cabinet a specimen of Peridermium," etc. This 
type material was destroyed in the great San Francisco fire, being bulky and 
in a box separate from other type collections, as Mr. Bethel writes, who has 
consulted the herbarium of the society both before and since the fire, and as 
stated by Meinecke (Phytopathology 10: 281. 1920), who also calls atten- 
tion to the duplicate specimen at the New York Botanical Garden, 

Arthur: New species of Uredineae — XIV 193 

about fifty miles west of Lake Tahoe. Harkness says the fungus 
"appears both on the limbs and trunks of young trees of tlie 
variety Pimis ponderosa, generally forming a complete circle 
around the trees, its sporidia appearing as a zone of bright 
orange yellow. The spores first germinate beneath the cuticle, 
which it destroys. Owing to the irritation of this presence an 
abnormal thickening of the cambium is produced, which in 
turn gives place to an excessive growth of woody fibre. This 
process being repeated from time to time a large bulbous ex- 
pansion is soon formed, so that as often occurs a stem of but an 
inch in diameter is enlarged to that of four or five. Above this 
bulb the further development of the stem is retarded, or arrested 
altogether, its place being supplied by a dense tuft of minute 


branches.*' Among other matters it is also stated that ^'in the 
\icinity of Colfax the fungus appears to be limited to an area 
of but a few acres in extent. Within that area, however, it is 
destroying the young growth." 

The quotation is taken from the report by the secretary of 
the San Francisco society sent to the Monthly Microscopical 
Journal of London, and printed on page 164 in volume 16, 
bearing the date of September i, 1876. This periodical was 
maintained by the Royal Microscopical Society. 

The name, host and locality are cited in Harkness & Moore's 
Pacific Coast Fungi in 1880, still without confusion with any 
other form. But when Harkness published technical descriptions 
of "New species of Californian fungi" in the Bulletin of the 
California Academy of Science in 1884, other forms and hosts 
were incorporated, and from this time on the application of the 
name has been various. 

The chief source of confusion has been in failing to rec- 
ognize the Pacific Coast form of Peridermiiim Cerebrum Peck, 
which occurs along the coast of California, and is especially 
abundant at Monterey on Pinus radiata {P. insignis). The 
present writers feel especially culpable for this confusion, for 
they placed P. Harknessii under P. Cerebrum as a synonym 
in their paper on North American species of Peridermiiim on 

pine (Mycologia 6: 133. I9i4)» and have been followed by 
Meinecke, In his paper on Peridermi'um Harknessii and Cro~ 
nartium Querctium (Phytopathology 6: 225-240. 1916), and 
by others. 

The present knowledge of the Castilleja rusts indicates that 
the correct application of the names should be as follows; 

194 Arthur: New species of Uredineae — XIV 

Cronarlhim filamentosuni (Peck) Hedge. {Peridermittm fila- 

menlosum Peck, 1882); 

Cronartium coleosporwides (Diet. & Holw,) Arth. (Uredo 

coleosporwides Diet. & Holw. 1893); 

Cronartium stalactiforme Arth. & Kern (Peridermmm stalacii- 

forme Arth. & Kern, 1906). 

Should it be deemed advisable to combine these three species 
as subspecie's, races, or forms, the name to be employed for the 
ag.s^rcgate species, using the oldest specific name, would be 
Cronarlhim Harknessii (Moore) Meinecke, Phytopathology 

10: 282. 1920. 

DIABOLE gen. nov. 

Cycle of development includes only subcuticular telia, with 
a possibility of pycnia. 

Telia somewhat indefinite in extent, without paraphyses. 
Teh'ospores free, usually paired on a common pedicel, one-celled, 
more or less adhering laterally; wall colored, usually verrucose, 
at least above; pore one, in upper part of cell. 

Diabole cubensis comb, nov- 

-L 1 

?) cubense Arth. Mem. Ton 


Mimosa pigra L. {M 

L. ) and was at first assigned with many misgivings to the genus 
Uromychdium. That connection has become more and more 
uncertain, and it is now believed to constitute a new and very 

distinctive genus. 


Puccinia Plucheae (Sydow) comb. nov. 

Uredo Plucheae Sydow, Ann. Myc. 1: 333. 1903. 

Uredo hiocellata Arth. Bull. Torrey Club 33: 517. 1906. 

Puccinia hiocellata Vestergr. Micr. Rar. Sel. 1267, 1908. 

The teliospores of this rust have been collected on two hosts, 
Pluchea fastigiata Griseb. and P. Quitoc DC, in Argentina, 
and issued In Vestergren 's exsiccati, Nos. 1267 and 1368. The 
rust occurs on a number of species of Pluchea in Florida, Guat- 
emala and the West Indes, but no teliospores have yet been 
found in North America. ' 

Uredo nominata sp. nov. 

11. Uredinia ampliig:nous, grouped upon discolored areas, 
5-10 mm. across, oblong or linear, 04-5 mm, long, somewhat 

Arthur: New species of Uredineae — XIV 195 

tardily naked, pulverulent, light-brown, ruptured epidermis 
evident; urediniospores ellipsoid or obovoid, 18-23 by 23-32 [x; 
wall light cinnamon-brown, 1.5-2 \x thick, moderately echinulate, 
the pores 3 or 4, approximately equatorial. 

On Sisyrinchium Bermtidianum L. {Iridaceae), St. Davids, 

Bermuda, February 26 — March 9, 1908, Stewardson Brotvn 611. 

A number of rusts have been described on Sisyrinchium, both 

from North and South America, but representatives of them are 

rare and scanty in herbaria. Judging from descriptions and 


sidered distinct. 

Uredo cumula sp. nov. 

II. Uredinia amphigenous, somewhat aggregate or scattered, 
round, applanate, small, 0.1-0.4 mm. across, soon naked, pulveru- 
lent, cinnamon-brown, conspicuous, ruptured epidermis notice- 
able; urediniospores broadly ellipsoid or obovate, small, 18-20 
by 22-25 ^\ v^^\\ pale cinnamon-brown, thin, i ^a or less, finely 
echinulate, the pores 2, equatorial, indistinct. 

On Buchnera elongata Sw. {Scrophidariaceae) , Herradura, 
Cuba, March 24, 192 1, John R, Johnston 2^jo. The species 
is notable for its small delicate spores, although when massed on 
the surface of the plant they are of ordinary conspicuousness. 
The generic name of the host is sometimes written Bnechnera. 

Uredo curvata sp. nov. 

II. Uredinia hypophyllous, loosely grouped on slightly 
discolored areas 3-5 mm. across, round, small, 0.1-0.3 nim. 
in diameter, soon naked, pulverulent, dark chestnut-brown, 
ruptured epidermis inconspicuous; paraphyscs numerous, per- 
ipheral, incurved, hyphoid or somewhat clavate, slender, about 
7 by 30 pi, the wall thin, less than i ji, slightly thicker above, 
1-2 [X, colorless below, light chestnut-brown above, sometimes 
one- or two-septate; urediniospores obovate, 14-16 by 26-32 [x, 
very strongly cur\^ed; wall chestnut-brown, uniformly thin, 
I fx, with one indistinct pore on the concave side below the 
equator, evenly echinulate-verrucose. 

On Inga vera Willd, {Mimosaceae), San Diego de los Banos, 
Cuba, March 26, 1 921, John R. Johnston 2540, The species 
is remarkable for its ver}- much curv^ed spores with one pore on 
the indented side, being in this respect similar to the uredinia of 
Piiccinia invaginata Arth., which occurs in Cuba on the rhamna- 
ceous genus Gouania. 

Aecidium Yuccae sp. nov. 

O. Pycnia amphigenous, in small and crowded groups, 
inconspicuous, honey-yellow, subepidermal, small, globoid. 


Arthur: New species of Uredinae — XIV 

80-115 yi in diameter; ostiolar filaments prominent, 45-65 [i 

long, agglutinated Into a column, 

I. Aecia amphigenous, irregularly arranged m loose groups, 
cylindric, 0.3-0.5 mm. in diameter and somewhat higher ;peridium 
firm, erect, the margin finely erose; peridial cells angularly 
globoid or ellipsoid in face view, rectangular or somewhat 
rhomboidal In section, 16-18 by 23-35 l^» abutted or slightly over- 
lapping, the outer wall 3-5 ^ thick, smooth, the inner wall thin- 
ner, about 2 jx, finely verrucose; aeciospores globoid or broadly 
ellipsoid, 16-20 by 19-24 ^; wall colorless, thin, I [i or less, very 
closely and finely verrucose. . 


On Yucca glmica Nutt. {Dracdenaceae), Crawford, Nebraska, 
June 28, 192 1, A. 0. Garrett. An interesting species, as it comes 
from the semi-arid, sandhill region, where a number of grass rusts 
of restricted range occur, to one of which it undoubtedly is 

■ L 

genetically connected. Only a few groups of aecia on the 
upper part of a young leaf were secured by Mr. Garrett, who 
kindly submitted all the material for study. 

Purdue University, 
Lafayette, Indiana 

Studies in the genus Luplnus — VII. L, succulentus and 



Charles Piper Smith 

(with four text figures) 



In accounting for the North American annuals and biennials 
of the subgenus Lupinus, I have found it advisable to recognize 
six groups, as indicated and contrasted in the key below. These 
groups, however, do not provide for the Mexican annuals, 
L. bilineahis Benth., L, Hart^vegi Lindb, and L. Barkeri LindL, 
nor for the Costa Rican L. Clarkei Oersted, as satisfiictory 
material of these species has not been available for my study. 

Keel petals ciliate on their lower edges near the claw, com- 

monly ciliate above, also, near the claw. 
Racemes shorter than their peduncles. 
Racemes longer than their peduncles. 

Flowers verticillate; leaflets glabrous above. 



Flowers scattered; leaflets more or less hairy above. Sf arsiflori. 

Keel petals non-ciliate on their lower edges. 
Keel petals ciliate above near the point. 
Keel petals non-ciliate on both upper and lower edges. 
Flowers verticillate; leaflets hairy above. 
Flowers with scattered 

Leaflets hairy above, 2-5 mm, wide; pods 10-15 

mm. long; seeds 1.5-3 rr^i^i- long. 
Leaflets glabrous above, 6-12 mm. wide; pods 30 50 
mm. long; seeds 4-6 mm. long. 





Fig. 75. I. Stiversiani; 2. Succulenti; 3. Sparsiflori; 4. Micranthi; 

5. CoNCiNNi; 6. Subcarnosi. 

Fig. 75 is here inserted to emphasize and contrast the above 


igS Smith: Studies in the genus Lupinus — VII 

indicated variations in the keel. Four of these groups, namely, 
the Sparsiflori, Stiversiani, Co?idn?ii, and Snbcarnosi, have been 
treated In the last two papers of this series (Bull. Torrey Club 
47:487-509. 1920; 48:219-234. 1921). This paper will con- 



The one species included here is so distinct from all the other 
annuals of our region that I do not feel justified in placing it in 
any of the other groups recognized. The ciliation of the keel 
is too constant to be ignored, though nowhere properly described. 
This is the group some years ago indicated by me under the 
name Affines (Muhlenbergia 6: 134. 191 1). 

la. Lupinus succulentus Dough; C. Koch, Wochenschrift 

Gaertn. Pflanzenkunde 4: 277. 1861. [Fig. 76.] 

Stout, succulent or fistulous, 2-6 dm. tall, branched, nearly 
glabrous or sparsely appressed-pubescent (rarely villous) : leaves 
several; petioles slender, 6-12 cm. long, one to three times as 
long as their leaflets, stipules linear-setaceous, 10-12 mm. 
long, the free part widely divergent; leaflets seven to nine, 
cuneate or cuneate-obovate, rounded, truncate, or emarginate 
at apex, usually apiculate, glabrous above, sparsely appressed- 
pubescent beneath, 20-70 mm. long, 6-16 mm. wide; peduncles 
2-8 cm. long, racemes 6-30 cm. long, flowers subverticillate in 
about four to eight whorls or groups, spreading in anthesis, 
becoming ascending upon withering, 12-17 mm. long; bracts 
early deciduous, linear, 6-10 mm, long; pedicels spreading- 
pubcsccnt with very short hairs, 46 mm. long; calyx bracteolate, 
subappressed-pubescent, the upper lip deeply two-toothed or 
bifid, about 5 mm. long, the lower lip lanceolate, entire and acute 
or three-toothed, 7-8 mm. long; banner suborbicular, about 
14 X 13 mm. including the claw, glabrous, blue with yellow 
center turning violet, or rarely bluish white, wings 12-14 mm. 
long, about 8 mm. wide, blue or rarely nearly white, more or 
less ciHate at the base above the claw, keel stout, 12-14 mm. 
long, the point upturned, distinctly ciliate near the claw both 
above and below, purple-tipped or orange-tipped in the albinos, 
otherwise whitish; pods about 50 mm. long, 9-10 mm. wide, 
loosely pubescent or villous with hairs 0.5 to 1.5 mm. long, 
ovules eight to ten; seeds oblong, 3.5-5 nim, long, much marbled 
with dark brown, w^ith a pair of contiguous w^hitish spots 
embracing the raphe, the hilum deeply sunken in a protruding 
ring: axial root normally vertical. 

This is the plant that has been known as L. affinis Agardh, 
since Watson reviewed the genus in 1873 (Proc. Am. Acad. 
8: 517), but the specimens so marked in the Lindley Herbarium 

bMrtH: Studies in the genus Lupinus — VII 



(now at Cambridge University, England) arc not this species. 


Indeed, Dr. Greene, in 1891 (Flora Franciscana 1:40), expressed 
doubt that our robust, succulent annual is the true L. affirn's, 
and I have long felt that Agardh s description does not satis- 
factorily apply to same. It remained, however, for Miss Alice 
Eastwood to unearth the needed evidence, and her photograph 
of Lindley 's specimen labelled L. succulenttis shows said specimen 
to be of this species. Likewise, her photographs of the specimciis 
labelled L. affinis Agardh show equally well that same should 
not be referred to this robust species, but rather to L, nanus^ 
a species which will be treated in a later paper. These are the 
conclusions arrived at by Miss Eastwood with the specimens 
before her. 

Koch knew^ the species as a garden plant under the name 
L. succtdentus and quotes Biedenfeld's Garten Jahrbuch as, his 



Fig. 76. Lupinus succulentus Dough i. C. P. Smith 1406 (CPS); 2. 

K. Brandegee (UC 81964); 3. Edna Hannibal (DS 87569); 4. L. R, 
Ahrams 4224 (DS); 5. K. Brandegee {\]C 187796), 

only authority for attributing the name to Douglas. He also 
states that the same species was distributed from Darmstadt 
as L. Liebmanni and from Geneva as L. ptirpitreus Del. Con- 
sidering the species to be undescribed, he drew up in 1861 a 
description in German, accrediting the species to Douglas and 
suggesting that its native home must be assumed to be California 
or Oregon. As his description is not readily available in this 
country, it is reproduced here: 

Die ganze, mehr neidrig bleibende Pflanze ist mit Ausnahme der 
Oberflache der Blatter sehr fein behaart; 9 langlichkeilformige Blattchen 
mit zuriickgebogener Spitze; Nebenblattchen wenig angewachsen, borsten- 

200 Smith: Studies in the genus Lupinus — VII 

formig, aufrecht; Aehre gestielt, kurz, aus wenigen (meist nur 3) Quirlen 
bestchend; Deckblatter langer als die Knospen; Deckblattchen klein und 
oft undeutlich; Oberlippe 2 theilig, kiirzer als die ganze Unterlippe;. 
Schiffchen an der Basis des Randes gewimi^ert. 

Dicse durch ihre dunkelen, schwarzvioletten Bluthen ausgezeichuete 
und sehr zu enipfehlende Art scheint noch gar nicht beschrieben zu sein. 
Obwohl Douglas als Autor angegeben 1st und man vermuthen muss, dass 
Kalifornien oder das Oregon-Gebiet Vaterland sind, so haben wir den Namen 
doch nur in Biedenf eld's Garten Jahrbucbe (im I. Hefte S. 292) gefunden. 
Aus welcher Zeitschrift der Verfasser die Pflanze entlehnte, wissen wir 
nicht. Neuerdings ist sic weider aus dem Darmstadter botanischcn Garten 


als L. L i e b ni a n n i , aus dem Genfer als L. p u r p u r e u s Del. ver- 
brietet worden. Wir vermuthen, dass die Namen in dem Verzeichnisse irgend 
eines botanischen Gartens aufgestellt wurden. 

Koch also here recorded his opinion that the species seems 
to be related to L. arvensis Benth. and to L. ramosissimtis Benth,, 
a Colombian species collected by Hartweg, and especially to 
the plant illustrated in plate eleven of the Botanical Register 
for 1847 and there named L. Ehrenbergii. Evidently some one 
claimed this species to be the same as L. densiflorus Benth.; 
for Koch, in 1867 (Ind. Sem. Hort. BeroL, App. 1: ii), published 
a Latin diagnosis and stated that it differed from L. densiflorus 
in its fragile stems and branches and violet flowers. Since this 
description is likewise difficult of access, I reproduce most of 
it here: 


Annuus, humiHs, ramosus, fragilis, puberulus; stipulae setiformes, 
erectae, paululum adnatae; folia 9-foliolata, 3-pollicaris; petiolus long- 
itudine folia aequans; . . . foliolis supra glaberrimus subtus pubcrulis 
oblongis, sed ad basin magis attenuatis, apice breviter cuspidate, recurvo; 

. . . spica verticillata, verticilHsdistantibus; . . . labium supcrius 
bifidum, inferius lanceolatum, integrum; .... vexillum et alae violaceae, 

. . , carina albida, ad partem superiorem violacea, apice flavescente; 
legurrien pilosum, inter scmina constrictum. 

Species per longum tani tcmpus in hortis culta e California nee non e 
terris mexicanis sine dubio allata, accredit ad L. densiflorus Benth., differt 
tamen fragilitate caulis ramosuniquc et floribus violaccis. Nusquam 
descripta esse videtur, sed nupperimc iterum in hortis botanicis nominibus: 
Lupini Liebmannii et purpurei occurrit* 

There is a sheet of this species, marked '^Lupinus sticculcntus 
Douglas, 1843", in a large old collection of European garden 
plants recently secured by Stanford University. Thus, I consider 
it advisable to accept this name for this species, which otherwise 
is without a proper botanical name. Douglas could hardly have 
overlooked the species during his sojourn in California, and 

certainly a more appropriate name could not be found. I cannot 

Smith: Studies in the genus Lupinus — VII 201 

follow Watson (Proc. Am, Acad. 8:538. 1873; and Bib. Index 
236. 1878) in assuming that this '*is probably but a garden 
form" of L. densiflorus, nor accept his application of Agardh's 
name to this species. Koch might have said more about the 
pods and seeds, but his 'Megumen . . , inter seniina 
constrictum" and his comparison of the species with L. arvensis, 
L. ramostssimtis, and L. Ehrenhergii do not point to a confusion 
with L. densiflortis. 

The species is so common in the San Francisco Bay region 
that I will omit citation of many specimens examined. An albino 
form (petals pale bluish white) was common in 1921, In and near 
the mouth of Alum Rock Canyon, Santa Clara County. This 
was mostly in pure patches, but sometimes associated with 
pale blue, or the normal, dark blue, form. Seeds were secured 
from marked colonies, but those from the palest-flowered plants 
were heavily pigmented, perhaps averaging even darker, rather 

than paler, than is normal for the species. 

California . Alameda County : Berkeley, Feb. , 1 899, 
H. P, Chandler 2JQ (UC); Livermore, April, 1904, A. A. Heller 
7320 (B, UC); Niles, June, 1918, H. A, Walker 5006 (UC); 
Sunol Valley, June, 1916, L. R. Abrams 5^99 (DS). Butte 
County: Chico, A. A. Heller 12983 (UCX), Colusa County: 
College City, 1905, Alice King (UC); Sycamore Slough, Sacra- 
mento River, April, 191 7, R. S. Ferris 621 (DS). Contra Costa 
County: Brentwood, May, 1893, ^- Eastwood (CA) ; Bryon 
Springs, March, 1914, A, Eastwood (CA) ; Concord, March, 1914, 
A, Eastwood (CA) ; Martinez, April, 1862, W. H, Brewer gg6 
(UC). Fresno County: Alcalde, March, 1892, T. S. Brandegee 
(CA); Alcalde, March, 1893, ^^ Eastwood (CA) ; Huron, March, 
1893, ^' Eastwood (CA). Kern County: Sunset Oil Wells, 
March, 1893, A. Eastwood (CA). Los Angeles County: Avalon, 
Santa Catalina Island, May, 1920, G, L. Moxley 732 (CPS); 
Claremont, April, 1904, C, F. Baker (CA) ; Elysian Hills, Feb., 
1903, E. Braunton (UC); Los Angeles, April, T901, E. D. Palmer 
(UC); Redondo, San Pedro Hills, March, 1903, L. R. Abrams 
3141 (DS); Santa Catalina Island, May, 1916, X, Brandegee 
(UC). Monterey County: Carmel-by-the-Sea, March, 1910, 
Alice D, Randall (DS) ; Castroville, T. S. Brandegee (CA); 
Jolon Grade from King City, April, 1920, C. D. Duncan 78 (DS) ; 
Pacific Grove, Pebble Beach, April, 1909, L, R. Abrams 4224 
(DS); Posts, June, 1893, A. Eastwood (CA). Riverside County: 


202 Smith: Studies in the genus Lupinus — VII 


Hcmet, May, 1904, C. F. Baker 4iSg {UC, DS). Sacramento 
Coimty; Sacramento, April and May, Edna Hannibal (DS). 
San Benito County: Idria, May, 1893, ^^ Eastuvod (CA) ; 
San Benito, May, 1918, A, Eashvood (CA). San Bernardino 
Comity: San Bernardino, April, 1888, S. B. & W, F, Parish 

1901, 5. B. Parish (DS). San Diego County: 
Campo, April, 1920, A. Eastwood (CA) ; La Jolla, April, 1903, 
T. 5. Brandegee (UC), and March, 1914, F. £. &" £. S, Clements 
(UC); Linda Vista, May, 1894, T. S, Brandegee (UC); Point 
Loma, April, 1913, A. Eastwood (CA) ; San Diego, June, 1885, 
Fanny £. Fish (UC), and March, 1889, C. R. Orctilt (CA), also 
April, 1905, K. Brandegee (DS, UC), San Francisco County: 
Twin Peaks trail, April, 192 1, Bertha Dold 105 (CPS). San 
Joaquin County: Tracy, April, 

1903, C. F. Baker 2908 (CA). 
,uis Obispo, May, 1882, M. E. 
r (CA), July, 191 1, X". Brandegee 


March, 1882, Mrs. R, W\ Summers {VC). San Mateo County: 
Crystal Springs Lake, April, 1896, A. Eastwood (UC) ; Portola, 
May, 1903, A. D. E. Elmer 4827 (CA, DS); La Honda summit, 
May, 1900, W. R. Dudley (DS); San Mateo ravine, April, 1894, 

W. R. Dudley (DS); San Pedro, June, 1903, A. D. E. Elmer 4681 
(CA, DS). Santa Barbara County: Ellwood, May, 1908, A. 
Eastwood 2 (CA); Santa Cruz Island, April, 1888, T. S. Brandegee 
(UC), July-Aug., 1886, E. L. Greene (CA), and June, 1918, 
A. Eashvood (CA); Santa Maria Rivei, June-July, 1906, A. East- 
ivood J2Q (CA). Santa Clara County: Alum Rock Park, April, 
1907, A. A. Heller 8471 (CPS, DS), Feb., 192 1, C. P. Smith 3212., 
albino, and 3213, normal colored (CPS); Loma Piieta, April, 
1894, J.B. Davy 631 (UC); Los Altos, April, 1894, W. R. 
Dudley (DS); Saratoga, June, 1915,!,. R. Abrams $260 (DS) ; 
Stanford Uni\ersity, April, 1903, A. D. E. Elmer ^.gio (CA, DS), 
and May, 1902, C. F. Baker 858 {Ch,\^C)-C3im^he\\, March, 
1921, Claribel Boesch loi (CPS); Evergreen, March, i()2\,Lofta 
Bland loi (CPS) ; Madrone Packwood School, April, 192 1, Mrs. 
A. F. Cochrane 102 (CPS). Santa Cruz' County: Chittenden, 
April, 1921, C. P. Smith 3259 (CPS); Ellicott, June, 1908, C. P. 
Smith 1455 (CPS), Shasta County: Anderson, April, 1914, L. 
E. Smith J3Q (CA). Solano County: Cordelia, April, 1902, Heller 
& Broun 5369 (B, DS) ; Vacaville, May, 1891, W. L. Jepson 
(DS), and May, 1903, C.F. Baker 5071 {T>'^/\JC). Sonoma 

Smith: Studies in the genus Lupinus — VII 203 

County: Petaluma Valley, April, 1908, C. P. Smith 1382 (CPS). 
Sutter County: Marysville Buttes, April, 1915,-4. A, Heller 
11,792 (CA, DS, UCX); West Butte, April, 1917, R. S. Ferris 
690 (DS); Yuba City, April, 1891, W, L. Jepson (UC). Te- 
hama County: Red Bluff, May, 1914, L. £. Smith 670 (CA). 
Ventura County: Ojai Valley, April, 1896, F, W. Hubby 41 
(UC). Yolo County: Davis, April, 1915, P. B. Kennedy 21 
(UCX); Yolo, June, 1914. C, P. Smith, seed only (CPS). 

Lower California, Carysito, April, 1885, C. R, Orcutt (CA). 

Arizona. Gila County: Roosevelt Dam, April 191 7, -4. East- 
wood 6286 (CA), and May, 19 19, A. Eastwood (CA). 

The distribution of this plant is thus seen to be from Shasta 
County to northern Lower California, the occurrence at Roosevelt 
Dam, Arizona, being certainly due to a casual introduction 
of seed from California, Variations are not particularly well- 
marked, except in two cases, as indicated below: 

lb, ^Lupinus succulentus Layneae var. nov, 

Humilrs ramis decumbentibus, villosus pilis pandentibus 
2 mm. longis, leguminibus aeque villosis. 

Depressed, i dm. tall, with decumbent branches and short 
racemes, densely villous with hairs 2 mm. long even on the pods. 

California, San Mateo County: Farallon City, Junei 
1918, K. Brandegee (Type, UC 187796). 

Mrs. Brandegee 's maiden name was Mary Katharine Layne, 

ic' Lupinus succulentus Brandegeei var. nov. [Fig. 77.] 

Humilis, erectus, foHoHs maximis prope 20 mm. longis, 
floribus 9-10 mm. longis, vexlllo 10 mm, longo, 7.5 mm. lato, 
alls apice non truncatis, carina ad basin super paulum ciliata. 

Much reduced in stature and size of flowers; about I dm. tall, 
largest leaflets about 20 mm. long, flowers 9-10 mm. long, 
banner about iq x 7.5 mm., wings not truncate at the apex, 
keel less ciliate abo\e. 

Lower California. Comondu, Feb., 1889, T. S. Brandegee 

i c 

m , 

Fig. 77. Lupinus succulentus Brandegeei C. P. Smith. T. S, Bran- 
degee (UC 8349S). 

264 Smith: Studies in the genus Lupixus — Vll 


(Type, UC 83498); San Julio, April, 1889, T. S. Brandegce (UC), 
and April, 1919, T, 5. Brandegee (CA). 


L. micranthus and its nearest relatives comprise the largest 

. and most difficult group of the annual species of the subgenus 

Ltipinus Watson. Variations in shape and size of flowers, 

leaflets, pods, and seeds produce conspicuous extremes which 

have suggested ''new species" not a few in number. That 

botanists have appreciated these variations is evidenced by the 

follow^ing list of published names: 

L. hicolor Lindh Bot. Reg. ii\ pL nog. 1827. 

L. micranthus Doug},; Lindl. Bot. Reg. 15: pL 1251, 1829. 

L. t^amfi Dough; Benth. Tran. Hort. Soc. IL 1:409. 1835. 

L. affinis Agardh, Syn. Gen. Lup. 20. 1835. 

L. Aschenhornii S. Schauer, Linnaea 20: 739. 1847. 

L. micranthus microphyllus^2its, Vroc, hnv, hcdiA. 8:535. ^873. 

L, niveus Wats, ibid, 11: 126. 1876. 

L. trifidus Torr.; Wats. ibid. 12: 250. 1877. 

L, ckihuahuensis Wa,ts\ ibid. 21: 423. 1886. 

L. carnosulus Greene, Bull. Cal. Acad. 2: 144. 1886. . 

L, umbellatus Greene, ibid. 2: 145. 1886. 

L. pachylobus Greene, Pittonia i: 65. 1887. 

L. polycarpus Greene, ibid. 2: 171. 1888, 

L. rostratus Eastwood, Proc. Cal. Acad. II. 6: 424. pi. 56. 1896. 

Z, persistens Heller, Muhlenbergia 2; 62. 1905. 

X. vaUicqla Heller, ibid. 4; 40. 1908. 

L. apricus Greene, Leaflets 2: 67. 1910. 

L. vallicola apricus C. P. Smith, Muhlenbergia 6: 135. 191 1. 

L. hirstitulns Greene, Leaflets 2: 152. 191 1. 

L. sabulosns Heller, Muhlenbergia 7: 9. 191 1. 

L. Piper smithii HeUer, ibii. 7: 93, 191 1. 

L. strigulosns Gandoger. Bull. Soc. Bot, France 60: 461. 1913- 

In addition to accounting for these published names, the 
scope of these papers will necessitate disposing of some seven 
herbarium names. Of the above, only one species will be treated 
at this time, the remainder being scheduled for attention in 
subsequent papers. 

The name Bicolores would have been a more appropriate 
appellation for this group; but the name here employed has been 
in use for some time. The flowers measure 6-16 mm. in length 
and are always bicolored but are certainly not always small. 
They are usually verticillate; but the racemes are sometimes 
reduced to one umbel-like whorl, and in one form the flowers 
are strictly scattered. The keel is normally ciliate on the upper 

Smith: Studies in the genus Lupinus- — VII 205 

margin near the point, a character not shared by any other group 


of North American annual lupines; however, two of the forms 
lack this ciliation,' The upper surface of the leaves is almost 
always with at least a few short hairs. 

2. LupiNUS niveus Wats. Proc. Am. Acad. II : 126. 1876. [Fig. 78.] 

Apparently annual or biennial with persistent pctioled 
cotyledons, erect, 3-4 dm. tall, branched well above the base, 
densely velvety tomentose above the cotyledons, which are 
fleshy, glabrous, the blades about 20 mm. long, 12 mm. wide: 
leaves velvety on both sides, petioles 6-8 cm. long, leaflets 
eight or nine, rounded or angled at the apex, 30-40 mm. long, 
9-12 mm. wide: peduncles 5-8 cm', long, racemes 8-12 cm. long, 
loosely few-flowered, bracts deciduous, linear, 5-8 mm. long; 
flowers 10 mm. long, spreading or subdrooping, pedicels 4-6 mm. 
long; calyx practically ebracteolate, the lips equal in length, 
about 5 mm. long, the upper cleft or bifid, over 4 mm. wide, 
the lower bluntly two-toothed, barely 3 mm. wide, petals broad, 
deep blue, banner suborbicular, about 10 x 10 mm., greenish 

I cm . 

Fig. 78. Lupinus niveus Wats. E, Palmer 861 (G, US). 

yellow in the center, wings about 10 x 7 mm., only slightly 
ciliate on the upper margin near the ver>^ sharp upturned point; 
pods yellow, 40-50 mm. long, 8-10 mm. wide, stout, ovules four 
or five; seeds pale, obscurely spotted, mostly 5-6 mm. long. 

This is a very distinct, isolated, insular species not closely 
related to any other known species; but as I see it, better lined 
tip with L. nanus, at least for the present. For a long time 
I was misled by Watson's clause, ''allied to L. lencophyUus'\ 
and thus had difficulty in classifying this plant. 

Lower California: Guadelupe Island, 1875, E. Palmer 25. 
(G); 1889, E. Palmer 861 (G, US); 1893, F. Franceschi (US). 

My thanks are especially due to Miss Alice Eastwood for 
the use of unpublished notes and to Dr, J. H. Barnhart for 
important bibliographical help. 

The abbreviations used herein in the citation of specimens 
may be identified by reference to the following list: 


Smith: Studies in the genus Lupinus — VII 

B, Brooklyn Botanic Garden; 
CA, California Academy of Science; 
CPS, private herbarium of the writer; 
DS, Dudley Herbarium, Stanford University; 
G, Gray Herbarium, Harvard University; 
UC, Department of Botany, University of California; 
UCX, Division of Agronomy, University of California Exper- 
iment Station; 

US, United States National Herbarium. 

San Jose, California 


A new American fossil hepatic 
Marshall A. Howe and Arthur Hollick 

(with one text figure) 



Fossil Hepaticae are among the rarest of known paleobot- 
anica! remains. The total number of described species is less 
than twenty. Of these more than half are identifiable as living 
species of Jungermanniaceae, all but one of which are represented 
merely by fragments enclosed in amber of recent geologic age, 
found In the Old World. The remainder probal)]y all belong in 
the Marchantiaceae. 

Three species only have been heretofore recorded from 
America,* all of which are referable, either definitely or provis- 
ionally, to the latter family. It should be remarked, however, 
that the identity, as well as the botanical relationship of the 
species last cited, in our footnote, is questioned by Fontaine, who 
says (/. c, p. 54): *'It is similar to that of a Brachyphyllum and 
the plant may be really a twig of that conifer. " Incidently it may 
also be remarked that this is the only species recorded for any 
geological horizon older than Tertiary, and that the type of the 
species, from the Jurassic of England, was originally described 
as an alga under the name Fitcoides erectus Bean.f 

Several years ago a collection of fossil plant remains from 
Florissant, Colorado, w^as sent by Mr. T. D. A. Cockerell to the 
New York Botanical Garden for examination. Among them 
was what appeared to be a fragment of a bryophyte; but it was 
so small and imperfectly defined that it was not identified, 
even as to its probable class relationship, until an enlarged pho- 
tograph (Fig .1) was made. The specimen evidently represents a 

(a) Preissites Wardii Knowlton, Bull. Torrey Club 21: 45S. pi. 2ig. 1S94. 
Eocene (Fort Union formation), Burn's Ranch, thirty miles from nicndive, 

(b) Marchantia Pealei Knowlton, Proc. U. S. Nat. Mus. 35: 157. />/. ^5, 
1908. Eocene (Lance formation), Custer County, Montana. 

(c) ^' Marchaniites erectus (Bean) Seward?", ^M^ Fontaine in Ward, 
Monog. U. S. Geo!. Survey 48: 53. pL 6^ /. I, 2. 1905. Jurassic, Douglas 
County, Oregon. 

jln Leckenby, Quart. Jour. Geol. Soc. London 20: 81. pi. ii, f. ja, jb 
(erroneously numbered 2a, 2b on the plate). 1864. 



Howe & Rollick: A new fossil hepatic 

member of the Bryophyta and probably belongs to the family 
Jungermanniaceac of the class Hepaticae. 

Jungermanniopsis gen. nov. 

Plants slender and minute, consisting of stem and leaves and 
showing a pronounced dorsiventrallty, with ventral (postical) 
rbothairs and dorsal (antical) apparently one-ranked but pro- 
bably two-ranked ovate-acumhiate, often subfalcate, assurgent 
or subvertical (antically secund) leaves and with occasional 
indistinct suggestions of much smaller lateral lanceolate or 
Hgulate leaves. 

Jungermanniopsis Cocker ellii sp. nov, 


Stem ascending at apex; leaves 1-1.5 mm. long, 04-0.7 mm. 
in maximum widths contiguous, narrowly spaced, or slightly 
overlapping at extreme base, the inter\^als between the leaves 
commonl}^ resembling, the outlines of inverted leaves, the often 
subfalcate acuminations directed forwards, the acuminations 
sometimes accentuated by reflexions of the subapical margins, 
the margins entire or with indistinct indications of small teeth. 

Collector: T. D. A, Cockerell. 
Formation and locality : M 

shale, Florissant, 


Colorado (Station 14). 



The one fragment, upon 
which the genus and species are 
based, is about 12 or 13 mm. long 
and evidently lies on its side. 
The specimen shows a possible 
forking of the stem near the apex, 
with one branch ascending and 
the other descending, but the 
descending part is probably 
only a collection of roothairs. 
Among the living Hepaticae,. the pointed, antically secund leaves 
of Jungermanniopsis suggest the widely distributed Ilerherta 
aihinca (Dicks.) S. F. Gray; but there is no indication that the 
leaves are bilobed as in the genus Ilerherta, In its antically 
secund leaves, it resembles also such leafy Hepaticae as James- 
oniella atihim^talis (DC.) Steph. and certain species of Nardia, 
but in the form and looser disposition of the leaves it is very 
different from any of these. The possible presence of smaller 


ERELLii Howe & Hollick 


ligulate or lanceolate lateral leaves might, if fully substantiated, 
result in placing the plant with the Musci rather than with the 
Hepaticae. But the small size and general habit of the organism 

Musci. The 
the collector. 



A new genus of fossil Liliaceae 

t. d. a. cockerell 
(with one text figure) 

In 1873* Lesquereux reported a singular fossil from the 

Miocene shales of Florissant, Colorado, as Ophioglossum Alleni. 

In 1878 he figured itf and assigned it to the genus Sahinia, 

on account of its close resemblance to Heer's European fossil 

5. reticulata. HolIick| gave a new figure, and placed the plant 

in the genus Tmesipteris, where it remains in Knowlton's 

This so-called Tmesipteris Alleni is common in the Florissant 

shales but has remained an unsolved problem. A number of 
years ago, at the British Museum, I showed specimens to Dr. 
A..C. Seward and Dr. A. G. Nathorst. We could all agree that 
there was no relationship with Tmesipteris, Init it was im- 
possible to even guess at the true position of the species. Seward,§ 
when publishing on the subject later, reject- 
ed the reference to Tmesipteris, without 
being able to suggest another alternative. 
In I9I3-1I to avoid perpetuating an obvious 
error, I referred the plant to Carpolithcs, 
supposing it to represent some sort of 
fruiting body. No seeds could ever be 
seen, although the appearance suggested 
a flattened pod; but the published figures 
failed to show a thickening on the midrib, ^■'^- ^- Brachyruscus 
a little beyond the middle. In a very good ^^^T'/Y ''!!!" ■ 

c t T-1 . . m. , A. Lladode bearing 

speamen found at Florissant m November, pj^tjn^t^ fl„,,,,. B. 

192 1, there is a definite body at this point, flower, enlarged. C. 
with a diameter of nearly 3 mm. It looks Fruit on a mature clad- 
like a small berr>% with the contents ex- ^^^- ^- Emarginate 

truded (Fig. I, C). apexof a mature dadode. 

Probably the riddle would never have been solved but for 
the discovery of a younger specimen in the Florissant shales, 

* For bibliographical references see Knowlton, Mesozoic and Cenozolc 
plants of North America. U. S. Geological Survey Bull. 696: 626. 1909. 
t Tertiary Flora, pi. 5, /. //, 1878. 
X Bull. Torrey Club. 21: pL 205, f. 12. 1894. 
§ Fossil plants 2: 25. 1910. 
.^ Am. Jour. Sci. VI. 36: 500. 


212 Cockerell: a new genus of fossil Liliaceae 

at Station 14. This (Fig. i, A, B) shows that we have to 
do neither with a leaf, frond nor pod, but with a cladode. At- 
tached to the midrib is an indistinct mass, presumably a thin bract, 


upon which can be seen a dark object (Fig. i, B), which seems to 
agree very closely with the flower of Riiscus, The pistil is 
erect and distinct. The plant is dioecious, and the staminate 
flower is as yet unknown.' At first I thought there might be 
two flowers, but the supposed second one is, J believe, part of 
the midrib. 

As I could not identify the plant with Rusciis, I consulted 
Dr. Britton, who very kindly sent me material of Phyllonoma 

ifolia Willd 

It was at once 

apparent that the fossil had nothing to do with Phyllonoma, 
but returning to Rtiscus as the only alternative, I was able 
to recognize close morphological similarity. The genus Ritsciis 
is today limited to a few Palaearctic species, with lanceolate 
to ovate sharply pointed cladodes. The apical points are stiff 
and sharp, as any one who has handled butcher's broom will 
testify. At first sight the venation seems quite different from 
that of the fossil, but if we imagine the Rtiscus cladode broadened 
and abbreviated until the principal veins are nearly or quite 
transverse instead of longitudinal, the correspondence is exact. 
The published figures of the fossil do not show that there are 
principal veins, just as in Rtiscus. 

The morphological correspondence Is so exact that it becomes 
a question whether the fossil can be referred to Rtiscus. However, 
considering the broad form of the cladodes, with truncate or 
emarginate, mucronate apex, the essentially transverse veins, 
and the remoteness in time and space, it appears justifiable to 
cstablisli a distinct genus, Brachyrusciis, the species becoming 
Brachyrusciis Alleni sp. nov.* WTiether Heer's Salvinia ret- 

* Dr, Hollick writes that he is not convinced that the flowering cladode, 
which I have figured" and described, belongs to the same plant as the *' Tmes- 
iplcris** AllcnL I had considered the possibility of there being two different 
things, and am personally entirely convinced that all the specimens belong 
to a single species. However, I will designate the flowering specimen as the 
type of the genus and of a species which I will name B. AllenL This name 
will remain valid if my view is correct, only B. Alleni Cockerell will be a 
synonym of B, Alleni (Lesq.) Cockerell, If it should be decided that there 
are two quite different plants, B. Alleni Cockerell will still stand, only to 
.fall in case some author considers that two distinct species of Brachyrusciis 

are present. 

Cockerell: A new genus of fossil Liliaceae 213 

iculata Is congeneric, it is impossible to say; the reticulation, 
as figured, looks different, 

Riisctis today has its western limit in Madeira. The insular 
form, which I saw in cultivation in Funchal, is regarded as a 
race of R. hypophyllum L., but is perhaps specifically distinct. 
The name R. latifolius Lowe is preoccupied by Rafinesquc. 


Cladodes pyriform to suboval in outline, with a strong midrib ; 
apex obtuse, emarginate, with a stiff mucro representing the 
end of the midrib or morphological apex; principal veins diverging 
from the midrib approximately at right angles, except toward 
the apex, where they form angles of about 45 degrees; principal 
veins connected by veinlets, forming a coarse reticulation of 

quadrilateral or kite-shaped areas; inflorescence borne on 
midrib as in RiiscuSy with apparently a single flower, which (as 
imperfectly preserved) presents no characters to differentiate 
it from Riisciis, Type the following: 

Brachyruscus AUeni sp. nov. 

Cladode in the type specimen broad-pyriform, with tapering 
base, length about 18 mm. and breadth 12 mm. [in my opinion 
the larger cladodes (length about 31 mm. and breadth 20 mm.), 
with dark and distinct venation, belong to the same species]: 
stems unknown but doubtless woody. 



The aim of this Index is to include all current botanical literature written by 
Americans, published in America, or based upon American material; the word 
America being used in the broadest sense* 

Reviews, and papers that relate exclusively to foresty, agriculture, horti- 
culture, manufactured products of vegetable origin, or laboratory methods 
are not included, and no attempt is made to index the literature of bacteriology. 
An occasional exception is made in favor of some paper appearing in an Ameri- 
can periodical which is devoted wholly to botany. Reprints are not mentioned 
unless they differ from the original in some important particular. If users 
of the Index will call the attention of the editor to errors or omissions, their 
kindness will be appreciated. 

This Index is reprinted monthly on cards, and furnished in this form to 
subscribers at the rate of three cents for each card. Selections of cards are 
not permitted; each subscriber must take all cards published during the term 
of his subscription. Correspondence relating to the card issue should be 
addressed to the Treasurer of the Torrey Botanical Club. 


Kern, F. D. Distribution of Berberis vulgaris in Pennsylvania. 

Bull. Torrey Club 48: 263-269. 31 D 1921. 

Kern, F. D. The J. Roberts Lowrie Herbarium. Torreya 20: 

79-81. O 1921, 

Kern, F. D. Observations of the dissemination of the barberry. 

Ecology 2: 211-214. 3 D 1921. 

Keuchenius, P. E. Die Rindenbraune der Ilevea brasUiensis. 

Centralbl. Bakt. Parasitenk, u. Infektionskr. 55: 14-74- 
/. 1-33. 18 N 1921, 

Keys, A. The Agricultural Department, Dominica. Kew Bull. 

Misc. Inform. 1921: 67-85. 1921. 

Khadilkar, T. R. A sectorial chimera in maize. Jour. Heretl. 

12: 284./. 34. 23 S 1921. ' 

Klugh, A. B. The marine algae of the Passamaquoddy region, 

New Brunswick. Contrib. Canad. Biol. 1917: 79-84. pi. 
8. 1917. 

Klugh, A. B. Springtime in the woods. Am. For. 27: 317- 

319, 324. My 1921/ [Illust.] 


2i6 Index to American botanical literature 

Knowlton, C. H. Herbarium of Rev. W. P. Alcott. Rhodora 
23: 47. 5 Ap 1921. 

Knowlton, F. H. Criteria for determination of climate by means 
of fossil plants. Bull. Geol. Soc. Amer. 32: 353-358. i 
S 1921. 

Knox, A. A. The induction, development, and heritability 
of fasciations. Carnegie Tnstit. Washington Publ. 98: 1-18. 

pL 1-5 + /. I. 1908. 

[This is the second paper of this publcatiion.l. 


Knudson, L., & Ginsburg, S. Suggestions with respect to the 
measurement of osmotic pressure. Am. Jour. Bot. 8: 
164-170. /. /. 3 Ap 1921. 

Knunker, A, Hans Freiherr von Tiirckheim. Gartenflora 
70: 19-22, 15 Ja 1921. 

Von Tiirckheim collected chiefly in Guatemala 

Koch, E*, & Rumbold, C- Phoma on sweet sorghum. Phyto- 
pathology 11: 253-268. pL g-ii + /. 7-j. 15 N 1921. 


Krausel, R. 1st Taxodium distichiim oder Sequoia semper- 
virens Charakterbaum der deutschen Braunkohle? Ber. 
Deutsch. Bot. Ges, 39; 258-263. /. 1-3. 10 S 192 1. 

Includes a discussion and illustrations of jthe parenchyma of Taxodium 

Krieger, L- C. C. Is Amanita pantherina edible or poisonous? 
Mycologia 13: 270, 271. 8 192 1. 

Lachmund, H. G. Some phases in the formation of fire scars. 


O 1921. 

Leighty, C. E., & Boshnakian, S. Genetic behavior of the spelt 
form in crosses between Triticum spelta and Tritictim 
sativum. Jour. Agr. Research 22: 335-364. pL 33 + /. 
1-3. 12 N 1921. 

Leonian, L. H. Studies on the Valsa apple canker in New 

Mexico. Phytopathology 11: 236-243. 21 O 1921. 

Lindau, G. Eine neue A phelandra- Art. Notizbl. Bot. Gart. 

u. Mus. Berlin-Dahlcm 8: 50. i S 1921. 

Aphelandra Dusenii from Brazil. 

Index to American botanical literature 217 

Lipman, C. B. A contribution to oar knowledge of soil relation- 
ships with Citrus chlorosis. Phytopatholhgy 11: 301-305. 
9 D 1921- 

Loeb, J. The quantitative basis of the polar character of re. 
generation in Bryophyllum. Science II. 54: 521, 522- 
25 N 1921. 

Long, C. A. E. Some rare plants from Knox County, Maine. 

Rhodora 23; 198, 199. 15 D 1921. 

Lyon, C. J. Comparative studies on respiration, XVIII. 

Respiration and antagonism in Elodca, Am. Jour. But. 
8: 458-463. /. J, 2. 19 D 1921. 

Macoun, J. M., & Holm, T. The vascular plants of the arctic 

coast of America west of the looth meridian, collected by 
the Canadian Arctic Expedition, 1913-18. Rep. Canad. 
Arctic Exped- 1913-18. 5A: 5-25. pL i~j, 14 O 1921. 

Magrou, J. Symbiose et tuberisation. Ann, Sci. Nat. Bot. X. 
3: 181-275. P^' ^~9 +f'-^'Q' N 1921. 

Mains, E. B. Unusual rusts on Nyssa and Urticaslmm. Am. 
Jour. Bot. 8; 442-451. /. 1-6. 19 D 1921. 

Inchides Aplospora, gen. nov. 



weed {Asclepias mexicana) as a poisonous plant. U. S. 
Dept. Agr. Bull. 969: 1-16. pi 1-3 + /- J. 6 1921, 


Marsh, C. D., Clawson, A. B., Couch, J. F., & Marsh, H, West- 
em sneezeweed {Helenium Hoopesii) as a poisonous plant. 
U. S. Dept. Agr. Bull. 947: 1,2 +/. J-5. 11 O 1921. 

McWhorter, F. P. Destruction of mosses by lichens. Bot. 

Gaz. 72: 321-325. pi. 13. 15 N 1921. 

Mead, C. W. Indian corn or maize. Nat. Hist. 21: 408-413. 

O 1921. [Illust.] 

Menzies, D. Sir Archibald Menzies, R. N. Card. Chron. 

70: 324. 24 D 1921. 

Merrill, E. D. Review of the new species of plants proposed 

by N. L. Burman in Flora Indica. Philipp. Jour. Sci. 19: 

329-388. S 1921. 

21 8 Index to AMRRiCiVN botamical literature 

Metcalf , W. Notes on the Bishop puie {Pinus muricata) . Jour. 

For. 19: 886-902. D 192 1. [Illust.] 

Miller, H. G. Further studies on relation of sulphates to plant 
growth and composition. Jour. Agr, Research 22: loi-iio. 
8 O 1921. 

Moffatt, W. S. Why Illinois prairie flowers have disappeared. 
Nat. Stud. Rev. 17: 307-309. N 1921. 

Molfino, J. F. Contribucion a la flora de la region de Bahia 
Blanca (Argentina). Physis S: 1-27. 31 O 1921. [Illust.] 

Mulford, F. L. Trees with bright autumn foliage. Am, For. 
27: 648-655. O 1921. [Illust.] • 

Munns, E. N. Coulter pine. Jour. For. 19: 903-906. D 1921. 

Munns, E. N. Effect of location of seed upon germination. 
Bot. Gaz. 72: 256-260. 15 O 192 1. 

Orton, C. R., & Weiss, F. The reaction of first generation 
hybrid potatoes to the wart disease. Phytopathology 
11: 306-310. 9 D 1921. 

Osborne, T. B., Wakeman, A. J., & Leavenworth, C. S. The 

proteins of the alfalfa plant. Jour. Biol. Chem. 49: 63-91. 
N 192 1. 

Pascher, A. Uber die Oberelnstimmungen zwischen den Diato- 

meen, Heterokonten und Chrysomonaden. Ber. Deutsch. 
Bot. Ges. 39: 236-248. /. 1-6. 10 S 1921. 

Pemberton, C. C. Overgrowth of stumps of conifers. Canad. 
Field Nat. 35: 81-87./. i~4^ 22 N 1921. 

Perry, W. J. Some observations on the relation of soil moisture 

to height growth of pine saplings. Jour. For. 19: 752, 
753. N 1921. 

Perry, W. J. Yellow pine reproduction. Jour. For. 19: 622- 

631. 1921. 

Plaisance, G. P., & Hammer, B. W. The mannitol-producing 

organisms in silage. Jour, Bact. 6: 431-443. S 192 1. 

Pomeroy, C. S. Bud variation in Eleagnus. Jour.. Heredity 

12: 227-230. /. ig, 20. 23 S 192 1. 

Index to American botanical literature 219 

Puttick, G. F. The reaction of the Fi generation of a cross 

between a common and a durum wheat to two biologic 
forms of Fuccinia graminis. Phytopathology 11: 205- 
213. 6 O 1921, 

Raber, O. L. The effect upon permeability of polyvalent cations 

in combination with polyvalent anions. Am. Jour. Bot. 
8:382-385./. J. 14 N 1921. 

Studies of Laminaria. 

Raber, O. L- The effect upon permeability of (I) the same 

substance as cation and anion, and (II) changing the 
valency of the same ion. Am. Jour. Bot. 8: 464-470./. j, 2. 
19 D 1921. 

Rangel, E. Nota sobre dois fungos. Arch. Escol. Super. Agr. 
e Med. Veter. 5: 35-40, pL j-5. S 1921. 

Ostvaldla, gen. nov., and Peniotrkhum saccardiniim, sp^ nov. 

Record, S. J- Figure in wood. Am. For. 27: 611-617. O 

192 1. [Illust.J 


Rehder, A. Plant nomenclature: more suggestions. Jour. Bot. 

59: 289-294. O 1921. 

Reinking, O. A. Fiji disease of sugar cane in the PhilHppine 

Islands. Phytopathology 11 : 334^337- P^- ^5»-f^- 9 ^ 1921. 

Reinking, O. A., & Groff, G. W. The Kao Pan seedless Siamese 

pummelo and its culture. Philipp. Jour. Sci. 19: 389-437. 
pi. 1-16 + f. I. O 1921. 

Rendle, A, B. Elisia, an overlooked genus-name. Jour. Bot. 

59: 261-264. S 192 1. 

IncUifles reprint of the original publication of this genus and its 3 species 
by "Milano" in New Orleans Med. & Surg. Jour. 1847: 614-616. 1847. 

Renner, O. Das Rotnervenmerkmal der Onotheren. Ber. 

Deutsch. Bot. Ges. 39: 264-270. 10 S 1921. 

Richards, B. L. A dryrot canker of sugar beets. Jour." Agr. 

Research 22: 47-52. pi. 4-g. i O 1921. 

Roark, E. W. The Septoria leaf spot of Rubiis. Pliytopath- 

ology 11:328-333- 9 D 1921. 

Due to Mycosphaerella Riibi, sp. nov. 

220 Index to American botanical literature 

J. W 


caused by Phylloslida conges fa Heald and Wolf. ' Jour. 
Agr. Research 22: 365-370- P^- 34 + /• -f. 2. 12 N 1921. • 

Robinson, T. R. The bud-sport origin of a pink-fleshed grape- 
fruit in Florida. Jour. Hered. 12: 195-198. 23 S 192 1. 

Rogers, E. C. Influence of the period of transplanting western 
white pine seedlings upon their behavior in nursery and 
plantation. Jour. Agr. Research 22 : 33-46./. 1-7, i O 1921. 

Round, E. M. Odontopteris genuina in Rhode Island. Bot. 

Gaz. 32: 397-403- /■ ^S^ ^5 D 1921. 

Rumbold, C, & Tisdale, E. K. Notes on Phoma insidiosa Tass. 
found on Sudan grass. Phytopathology 11 : 345 9 D 192 1 . 

Ruschmann, W. Die Cohune- oder Corozoniisse Mittelame- 
rikas. Tropenpflanzer 24: 147, 148. O 192 1. 

Satterthwait, A. F. Notes on the food plants and distribution 
of certain billbugs. Ecology 2: 198-210. /.J. 3D 192 1. 

Schlechter, R. Die Thismieae. Notizbl. Bot. Gart. u. Mus. 
Berlin-Dahlem 8: 31-45. i S 192 1. 

Includes Triurocodon, gen. nov., from Brazil, and new American species 
in Sarcosiphon (i), Myostoma (i). 

Schuepp, O. Zur Theorie der Blattstellung. Ber. Deutsch. 

Bot. Ges. 39: 249-257./. i, 2. 10 S 1921. 

Schultz, E. S. A transmissable mosaic disease of Chinese cab- 
bage, mustard, and turnip. Jour. Agr. Research 22: 173- 
178. pi, B, 15 O 1921. 

Seymour, E. K., & McFarland, F. T. Loss from rye ergot. 
Phytopathology 11: 285-289./, /. 2, 15 N 1921. 

Shapovalov, M., & Edson, H. A. Blackleg potato tuber-rot under 
irrigation. Jour. Agr. Research 22: 81-82. pi. 13-16 + 
pL A. 8 O 1921. 

Showalter, A. M. Chromosomes of Conocephalum conicum. 

Bot. Gaz. 72: 245-249. pL 4, 5. 15 O 1921. 

Sievers, A. F., & Mclntrye, J. D. Changes in the composition 
of paprikas {Capsicum annmim) during the growing period. 


S 1921. 

Index to Amilrican botanical literature 22 1 

Sinnott, E* W. The relation between body size and organ 

size in plants. 

/. J, 2, O 192 1. 

Skinner, J. J., & Reid, F. R. Nutrient requirements of clover 
and wheat in solution cultures. Soil. Sci. 12: 287-296. pL 
J. O 1921. 

Small, J. K. Along the Juniata River [Pennsylvania]. Jour. 

New York Bot. Gard. 22; 168-171. S 1921. 

Small, J. K. Another Sonchiis for America, Torreya 21; 

100, loi. D 1921. 

Records the finding of the European Sonchiis uUginosiis ui Pennsylvania. 

Small, J. K. Cercis chinensis. Chinese redbud. Addisonia 

6: 33, 34. pL 209. 24 1921. 

Small, J. K, Monarda didyma. Oswego-tea. Addisonia 6 : 
47, 48. pL 216 24 192 1. 

Smiley, F- J. A report upon the boreal flora of the Sierra 

Nevada of California, Univ. California Publ. Bot. 9; i- 

423. pL 1-7. S 1921. 

Includes 21 new names or combinations. 

Spencer, E, R. Decay of Brazil nuts. Bot. Gaz. 72: 265-292. 
pi 8-12 + /. /-J. 15 N 1921. 

Includes new species in Pellioniella (i), Cephalosporium (i), Phomopsis 


Spink, H. M. The geographical distribution of commercial 
timber in the Pacific States of North America and its 
significance to the lumber industry. Scottish Geog. Mag. 

37: 257-266. 15 O, 1921. [Illust.] 

Sprague, T. A. The generic name Schizonotus. Jour. Bot. 59: 

^49~25i; S 1921. 

Sprague, T. A. Plant dermatitis. Jour. Bot. 59; 308-310. 

N 192 1. 

Steil, W. N. Vegetative reproduction and aposporous growths 

from the young sporophyte of Poly podium irioides. Bull. 
Torrey Club 48: 203-205. 26 S 192 1. 


Steil, W. N. The development of prothallia and anlheridia 

from the sex organs of Polypodiicm irioides. Bull. Torrey 
Ckib 48; 271-277. pi. 4 + f. i-4- 31 D 1921. 

Index to American botanical literature 

Sure, B., & Read, J. W. Biological analysis of the seed of the 
Georgia velvet bean, Stizolohmm Deeringianum. Jour. 
Agn Research 22: 5-15./. i~iS- I O 1921. 

Swanson, C. O. Hydrocyanic acid in Sudan grass. Jour. 
Agr. Research 22: 125-138. 15 O 1921, 

Tavares, J. S. Cecidologia brazileira. Broteria, Zool. 19: 
97-112. I D 1921. 

Taylor, F, B. The literature of diatoms. Trans. Am, Micr. 
Soc. 40: 187-194. O 1921. 

Taylor, M. W. Internal aecia of Puccinia albiperidia Arthur. 

Phytopathology 11: 343, 344-/- /. 9 D 1921. 

Taylor, N. Endemism in the Bahama flora. Ann. Bot. 35: 
523-532. O 1921. 

Taylor, N. Plant Immigrants and natives: a comparison, 
Brooklyn Bot. Gard. Leafl. IX. 9: 1-7. 26 O 192 1. 

Thaxter, R. Lincoln Ware Riddle. Rhodora 23: 181-184. 

15 D 1921. 

Thorn, C, & Lefevre, E. Flora of corn meal. Jour. Agr. 

Research 22: 179-188. 22 1921. 

Thompson, W, P. A botanical trip through German South-west 
Africa. Canad. Field Nat. 35: 74, 75. 25 O 192 1. 

Townsend, C. O. The beet-sugar industry in the United States. 


14 O 1921. 

Urban, I, Plantae Caribaeae. Notizbl. Bot. Gart. u. Mus. 

Berlin-Dahlem 8: 25-30. i S 192 1. 

Includes new species in Capparis (i), Sloanea (i), Pachira (i), Basana- 
cantha (i). 

Vaupel, F. Echinopsis mamillosa Giirke. Monatschr. Kakteenk. 

31:152-155. O1921. [lUust.] 

VinaU, H. W., & Getty, R. E. Sudan grass and related plants. 

U. S. Dept. Agr. Bull. 981: 1-68./. 1-25. 21 D 1921. 

Wagner, E.. Einiges iiber Malacocarpus. Monatschr. Kak- 
teenk. 31: 143, 144. S 1921. 

Vol. 49 

No. 8 




AUGUST, 1922 

Philippine Basidiomycetes — ^V* 

Paul W. Graff 

. PoRiA Persoon 

PoRiA LEUCOPLACA (Berk.) Sacc. SylL Fung. 6: 322. 1888. 
Poly poms leucoplactts Berk.; Hooker, Flora Nov. Zeland, 2: 
180. 1855. 

Luzon: Province of Benguet, Pauai, June, 1909, R, C. 
McGregor^ Bur. Set. 8720^ at an elevation of 2,100 m,, on decaying 

Collected previously in New Zealand and Java. 

Trametes Fries 

Trameies aspera (Jungh.) Bres. Hedwigia 53: 69. 1912. 
Poly poms asper Jungh. Flor, Crypt. Javae 1; 61. 1838. 
Polystictus asper Sacc. Syll. Fung. 6: 224. 1888. 
Luzon: vicinity of Manila, August, 1912, Sanchez 26, 57, 





*The papers of this series already published have appeared as follows. 
Additions to the basidiomycetous flora of the Philippines, Philip. Jour. Sci: 
8: (Bot.) 299-309. pL S-xo* 1913; Philippine Basidiomycetes, 11. Philip 
Jour. Sci. 9: (Bot.) 235-255. pL 2. 1914; Philippine Basidiomycetes — III. 
Bull. Torrey Club 45: 451-469. pL 13. 1918; Philippine Basidiomycetes 
IV. Bull. Torrey Club. 48: 285-295. 1922. 

[The Bulletin for July (49: 189-222) was issued August 10, 1922.] 


224 Graff: Philippine Basidiomycetes— V 

Polyporus incanus Lev.; Gaudichaud, Bot.Voy. Bonite 1: 183. 
pL ijy, f. 2. 1846. 

Luz(-n: Province of Bataan, Mount Mariveles, November, 
1912, P. W. Graff, Bur. Sci. S157, S158. 

This fungus was first collected in the Philippines by Gaudi- 
chaud-Eeaupr^, in 1836-37, on the visit of the corv^ette "La 
Bonite " to the port of Manila. The fungus has not been reported 
since, and it is with considerable pleasure that one of these 
three-quarter century old species is again collected. 

Trametes paleacea Fr. Nov. Symb. Myc. 97. 1851. 
Polyporus paleacetis Fr. Epicr. Myc. 471. 1838. 

Luzon: Province of Bataan, Mount Mariveles, November 
3-19, 1912, P. W. Graff, Bur. Sci. igogS, on a decaying los: at 
an elevation of 400 m. 

Previously reported from New Guinea. 

Trametes corrugatus (Pers.) Bres. Hedwigia 51: 316. 1912, 
Polyporus corrugatus Pers.; Gaudichaud, Bot. Voy. Uranie 

172. 1826. 
Polyporus fusco-hadius Pers.; Gaudichaud, Bot. Voy. Uranie 

[72. 1826. 
Polyporus scahrosus Pers.; Gaudichaud, Bot. Voy, Uranie 

172. 1826. 
Daedalea sanguinea Klotz, LInnaea 8: 481. 1833. 
Polyporus indecorus Jungh. Flor. Cr>pt. Javae 1: 51. 1838. 
Polyporus tegnlaris Lev. Ann. Sci. Nat. Bot. III. St 131. 1846. 
Hexagonia cruenta Mont. Syll. Gen. Spec. Crypt. 169. 1856. 
Polystictus Persoonii Cooke, Grevillea 14: 85. 1885. 
Trametes nitida Pat. Jour, de Bot. 4: 17. 1890. 
Earliella cubensis Murr. Bull. Torrey Club 32: 479. I9*^5- 
Earliclla corrugata Murr. Bull. Torrey Club 34: 468. 1907. 

Luzon: Province of Laguna, Mount Maquiling, Februar}% 
19 1 2, P. W. Graff, Bur. Sci. 1602^. 

This IS a much described species of universal tropical distri- 
bution. It was first described from material collected in the 


India, Ceylon, Java, Borneo 
and the American tropics. 

Graff: Philippine Basidiomycetes — ^V 225 

Daedalea Persoon 

Daedalea lurida Lev. Ann, Sci. Nat. Bot. III. 2: 189. 1844. 

415. 1908. 


, January, 1906, F. W. Foxii 
subconfragosa Murr.). MiXD 

■ District of Davao, Lake Laneo, Camp Keithley, July, 1907, 
Mary S. Clemens '' V'\ 

Murrill's species, described from Philippine material, seems 
to be -without doubt identical with that described by Leveille 
from Javan specimens, 

Daedalea Palisoti Fr. Syst. Myc. 2: 335. 1821. 

Daedalea amanitoides Beauv. FL Owar. 1: 44, pi. 2$, 1804. 
Daedalea repanda Pers.; Gaudichaud, Bot. Voy. Uranie 
168. 1826. 


Daedalea applanata Klotz. Linnaea 8: 481. 1833. 

Lenzites Palisoti Fr. Epicr. Myc. 404, 1838. 

Lenzites repanda Fr, Epicr. Myc. 404. 1838. 

Lenzites applanata Fr, Epicr. Myc. 404. 1838. 

Lenzites pallida Berk. Jour. Bot. 1: 146. 1842. 

Lenzites platypoda Lev. Ann. Sci. Nat. Bot. IIL 2: 180. 1844. 

Bur, Sci. 

5, July, 1906, M. 

Mount Maquilli 

W, Graff, 

tree trunk. CuLiON: October, 1903, Merrill 3531. Negros: 
Gimagaan River, Januarj^ 1904, Copeland 21. 

Daedalea indica Jungh., described from Javan material, 
should in all probability be added to the list of synonyms of 
this species. Daedalea, in the Polyporaceae, and Lenzites, in the 
Agaricaceae, are so closely related, and the species of one so 
often show a strong tendency to a variation toward the other, 
that considerable confusion has arisen in their determination. 
This is especially true in the case of this species and D.flavida Lev. 
in the Philippine material. In the latter species the variation 
even goes further, producing a Trametes form. 

The types of both Leveille 's and Berkeley's species are 
from material collected in the Philippines. This species is of 
very general tropical distribution. 

Daedalea pruinosa L6v. Ann. Sci. Nat. Bot. IIL 2: 199- 1844. 

226 Gr.\ff: Philippine Basidiomycetes — V 

Hexagonia glabra Lev. Ann. Sci, Nat. Bot. III. 5: 143. 1846. 
Luzon: Province of Laguna, Mount Maquiling, February, 
1912, P, W. Graff, Bur. ScL 16015. 

Collected previously in India and the Hawaiian Islands. 

Daedalea versatilis (Berk.) comb. nov. 

Trametes versatilis Berk. Jour. Bot. 1: 150. 1842. 
HexQgoniaciliata Klotz. Nov. Act.Acad. Nat. Cur. 19:(Suppl.) 

235- pi' 5, /' ^- 1843- 

PolyUitictiis versatilis Sacc. Syll. Fung. 6: 244. 1888. 

Fmialia versatilis Murr. Bull. Torrey Club 34: 469. I9C>7* 
Luzon : Province of Bataan, Lamao, February, 1904, Williams. 

As represented in Philippine material this species is of a de- 
cidedly daedaleoiu type. Occasionally Trametes-like forms are 
collected, showing that it may merge into the type of plant 
described by Berkeley. These forms are rare, however, as also 
are such as might tend toward the genus P oly stictiis . 

Collected in the American tropics and the Philippines- 

Elmerina Bresadola 

Elmerina cladophora (Berk.) Bres. Hedwigia 51: 319. 1912- 
Hexagonia cladophora Berk. Jour. Linn. Soc. Bot. 16: 47. 
. 1878. 
Luzon; Province of Nueva Ecija,Cabanatuan, October, 1908, 

R. C. McGregor, Bur. Sci. 52'/2] Province of Rizal, Jalajala, 

October, 1910, C, B, Robinson, Bur. Sci. Iig25. 
Reported only from the Philippines. 

Elmerina vespacea (Pers,) Bres. Hedwigia 51: 319. 19^2. 
Hexagonia vespacea Pers.; Gaudichaud, Bot. Voy. Uranie 

170. 1826. 
Lenzites aspera Klotz. Linnaea 8: 480. 1833. 
Polyporus lacerus Jungh. Flor. Crypt. Javae 1: 65. 1838. 
Daedalea inconcinna Berk. Jour. Bot. 1: 151. 1842. Not 


Lenzites platyphylla L6v. Ann. Sci. Nat. Bot. III. 2: 179. 1844. 
Daedalea aulaxina L6v. Ann. Sci. Nat. Bot. III. 2: 197. 1844. 



Hexagonia macrotrema Jungh.; Fries, Symb. Mv 

Graff: Philippine Basidiomycetes — V 227 

Hexagonia albida Berk. J 


Hexagonia Cesati Berk.; Cesati, Myc. Born. 8. 1879. 
Daedalea pruinosa Ces. Myc, Born. 7. 1879. Not Lcveill6. 
Daedalea ijitennedia Berk. Jour. Linn. Soc. Bot. 18: 385. 

Lenzites nivea Cooke, Grevillea 15: 94. 1886. 

Negros: Province of Negros Oriental, Dumaguete, April, 

1908, Elmer g86o. 

, Mauritius, Surinam,- and J 

Hexagoma Fries 

Hexagonia apiaria (Pers.) Fr. Epicr. Myc. 497. 1836. 

PoJyporiis apiarius Pers. ; Gaudichaud, Bot. Voy. Uranie 
172. 1826. 

Hexagonia Koenigii Berk. Ann. Nat. Hist. H, 10: 379. 1853. 

Luzon: Province of Bataan, November, 1909, H. M. Curran, 
For. Bnr. ig226. 

Collected in Java, Ceylon, and the Rawak Islands. 

Hexagonia Clemensiae (Murr.) comb. nov. 

Inonotiis Clemensiae Murr. Bull. Torrey Club 35: 401. 1908. 
Polyporns Clemensiae Sacc. & Trott.; Saccardo, Syll. Fung. 

21: 272 1912. 

Mindanao: District of Davao, Lake Laneo, Camp Keithley, 
September-October, 1907, Mary S. Clemens^ on dead wood. 
Not reported as yet outside the Philippine Islands. 

Hexagonia cucullata (Mont.) Murr. Bull. Torrey Club 31: 

332. 1904. 

Favolus cticullatiis Mont.; Ramon de la Sagra, Hist. Phys. 

Polit. Nat. Cuba 9: 378. 1841. 
Favolus curtipes Berk. & Curt. Jour. Bot. Kew Card. Miscel. 

1: 234. 1849. 
Hexagonia Taxodii Murr. Bull. Torrey Club 31: 332. 1904- 
Luzon, Province of Isabella, Palanan Bay, June, 1913, 

Escritor, Bur. Sci. 21185. 

Collected in Ceylon, the West Indies, tropical America, and 

eastern United States. 

Hexagonia subrubida (Murr.) comb. nov. 

Hapalopiltis subrubidus Murr. Bull. Torrey Club 34: 47^- 


228 Graff: Philippine Basidiomycetes — ^V 

Polyporus sabrubidus Sacc. & Trott.; Saccardo, Syll. Fung. 

21: 276. 1912, 

Luzon: Province of Bataan, Mount Mariveles, November, 
1904, Elmer 6gii, on dead tree branches. 

This species should' have been transferred to the genus 
Hexagonia by Saccardo and Trotter instead of to the genus 
Polyporus, to which it does not belong. 

So far only found in the Philippine Islands. 

Favolus Fries 

(Mont.) Bres. H 
us Mont. Ann. S 



princeps Berk. & Curt. Jour. Linn. Soc. Bot. 10: 

321. 1869. 

Luzon: Province of Nueva Vizcaya, Mount Umuguen, 



Favolus mulitplex Lev. Ann. Sci. Nat, Bot, IIL 2: 203. 1844. 

Luzon: Province of Bataan, Balanja Mountains, January, 
19 10, H. M, Cur ran, For. Bur. ip^32. 

According to the collector's note this fungus is edible and 
used by the Negritos of Bataan Province as a food. 

This species has been previously reported from Brazil and 


2; 203, 1844. 

(J ungh 

Jungh. Flor. Crypt. Javae 1: 75. 1838. 

Ilymenogramme spathulata Sacc, & Cub. ; Saccardo, Syll. Fung 
S: 653. 1887. 

Mindanao: District of Davao, Mount Apo. September 

1909, Elmer 11572. 


Gloeoporus Montagne 

Gloeoporus reticulatus (Fr.) comb. nov. 

Auricularia reticulata Fr. Epicr, Myc. 555. 1836. 
Gloeoporus conchoides Mont,; Ramon de la Sagra, Hist 
Phys, Polit. Nat. Cuba 9: 385. pL is, j. i, 1841. 

Graff: Philippine Basidiomycetes — V 229 

Thelephora dolosa Lev. Ann. Sci. Nat. Bot. III. 2: 209. 1844. 
Luzon: Province of Laguna, Mount Maquiling, Februarys 
1912, P. W. Graff, Bur, Sci. 15952. 

Collected previously in Brazil, Cuba, and Java. 

Gloeoforus dichrous (Fr.) Bres. Hedwigia 52: 74. 1912. 

Polyporus dichrous Fr. Obs. Myc. 1: 125. 1815. 

BoleHis thelephoroides Hook.; Kunth, Syn. Plant. 10. 1822. 

Boletus dichrous Spreng. Syst. Veget. 4: 475. 1828. 

Polyporus nigro-purpurascens Schwein. Syn. Fung. Amer, 
Bor. 360. 183 1. 

Polyporus pelleponis Seer. Mycogr. Suisse Champ. 73. 1833. 

Polyporus thelephoroides Fr. Epicr. Myc. 473. 1836. 

Gloeoporus candidus Speg. Fung. Guar. Pug. 1: 30. 1883. 

Polystictus thelephoroides Sacc. Syll. Fung. 4: 284. 1886. 

Luzon: Province of Rizal, January, 1906, F. W. Foxworthy, 
Bur. Sci.' yg. Mindanao: District of Davao, Mount Apo, 
May, 1909, Elmer J0747. 

Collected in southern Europe, the American tropics, and 

Laschia Fries 
Laschia minima (Jungh.) Sacc. Syll. Fung. 6: 409. 1888. 

Polyporus minimus Jungh. Flor. Cr>^pt- Javae 1: 64. 1838. 

Mindanao: Subprovince of Butuan, March-July, 1911 
Weber I22g^ on decaying woods. 
Reported previously from Java. 



Pleurotus Fries 
Pleurotus flabellatus Berk. & Br. Jour. Linn. Soc. Bot. 

11: 529. 1871. 

Luzon: Province of Benquet, Baguio, December 5, 19 10, 

E. Fenix, Bur. Sci. 12Q48, on dead wood. 

Collected previously in tropical America, Ceylon, and South 


Lentixus Fries 
Lentinus fusco-purpureus Kalchbr. Grevillea 8: I53- iS^^- 

230 Graff: Philippine Basidiomycetes — V 

Luzon: Province of Rizal, Bosoboso, July, 1906, M, Ramos, 

Bur, Sci. iiq6. 

This species, while allied to Lentimts Zeylieri Berk., seems 

to be distinct. . 

Described from material collected on the Richmond River, 

Australia, and not previously reported from elsewhere. 

Lentinus javanicus Lev. Ann. Sci, Nat. Bot. IIL 5: 118. 


Lentinus Decaisneanus Lev. Ann. Sci. Nat. Bot. IIL 5: 120. 

Lentinus infundihidiformis Berk. & Br. Jour. Linn. Soc. 

Bot. 14: 42. 1875. 
Luzon: Province of Bataan, Lamao, February 29, 1904. Cop- 
eland 175, 

Previously reported from Java. 

Lentinus polychrous L6v. Ann.ScI. Nat. Bot. IIL 2: 175. 1844. 
Lentinus praerigidus Berk. Jour. Bot. Kew Gard. Miscel. 

6: 132. 1854, 
Lentinus Kurzianus Berk. & Curr. Trans. Linn. Soc. 11. 

1: 120. pL 20y /. 2. 1876; 
Luzon: Province of Cagayan, Aparri, March, 1909, H. M, 
Curr an, For. Bur. 168 ig. 

Collected in Java, Sumatra, and eastern India. 

Lentinus sajor-caju (Rumph.) Fr. Epicr. Myc. 393. 1838. 

Agaricus sajor-caju Rumph. Herb. Amb. 11: 125, pL $6, 

fig. I. 1741-1755- 

Luzon: Province of Bataan, Mount Mariveles, January, 

1904, Merrill 3/04. 

Collected in Amboinia and Java. 

Lentinus TANGHiNiAE Lev. Ann. Sci. Nat. Bot. IIL 5: 119. 1846, 
Mindanao: District of Davao, Todaya, March, 1904, 

Copeland 516, 

Described from material collected in Madagascar. 

Lentinus velutinus Fr. Linnaea 5: 510. 1830. 

Scleroma velutinum Fr. Epicr. Myc. 392. 1838. 

Lentinus setiger Lev. Ann. Sci. Nat. Bot. IIL 2: 176. 1844. 

Lentinus hrachatus L^v.; Zollinger, Syst. Verz. Ind. Arch. 
^7- 1854. 

Graff: Philippine Basidiomycetes — V 231 



Mindanao: District of Davao, Todaya, April, 1904, Copeland, 
Collected previously in Brazil, Cuba, and Guiana. 

Lenzites Fries 


srziTES ACUTA Berk. Jour. Bot. 1: 146. 1842. 

Luzon: Province of Cagayan, March, 1909, Becani, For. Bur. 

The gills m this species have a somewhat wavy margin as 
well as being notched and, while the majority of the longer 
ones are forked, most of the shorter are distinct. 

This is the only collection w^hich has been made of this 
fungus in the Philippines since Cuming's collection of the type, 
between 1836 and 1840. In the meanwhile the species has been 
collected in northern India. 

Lenzites platyphylla Lev. Ann. Sci. Nat. Bot, III. 2: 179. 

1844. Not Cooke. 

Luzon": Province of Nueva Vizcaya, vicinity of Dupax, 
January-April, 19 12, R. C. McGregor, Bur. Sci. 14353 • 20284; 
Province of Bataan, Lamao, November, 1909, H. M, Ctirran^ 
For. Bur. igi88; Mount Mariveles, Januar>^ 1910, H. M. 
Curran, For. Bur. 19244a, Mindanao: District of Davao, 
Lake Lanao, Camp Keithley, September, 1907, Mary S. Clemens. 

Collected previously in JaY'a and Guiana. 

Lenzites striata (Swartz) Fr. Epicr. Myc, 406. 1838. 

Agaricus striatus Swartz, Prodr. Fl. Ind. Occ. 148. 1788. 

Mertilius striattis Lam. Encyc. M^th. Bot. 4: 127. 1797- 

Daedalea striata Fr. Syst. Myc. 1: 334. 1821. 

Luzon: Province of Bataan, Mount Mariveles, November 
3-19, 1912, P. W. Graff, Bur. Sci. 19044, at an elevation of 350 
m., on decaying and burned logs. 

Previously collected in tropical America, (including the 
West Indies,) Australia, and Borneo, 

ScHizoPHYLLUM Fries 

ScHizoPHYLLUM COMMUNE Fr, Syst. Myc. 1: 330. 1821. 
Agaricus multifidus Batsch, Elench. Fung. 1: 124. /. 126. 

Scaphophorus agaricoides Ehrh.; Nees, Hor. Phys, Berol. 94. 


232 Gilvff: Philippine Basidiomycetes — ^V 

Schizonia vulgaris Pers. Myc. Eur. 3: 14. 1828. 
Schizophyllum alneum Schrot.; Cohn, Krypt. Flor. Schles. 

3: 383^ 1887. 

July, 1906, M. 

Bur, Sci, 1 19s; Panay, Iloilo, January 2, 1904, Copeland ig. 
A fungus of universal distribution living on woody hosts of 
all sorts, both as a saprophyte and as a facultative parasite. After 
gaining entrance beneath the bark of living trees, through wounds, 
this species is able to slowly advance through the tissues which 
are apparently killed in advance of the progress of the fungus. 

Crepidotus Fries 

Crepidotus pogonatus Kalchbr. Grevillea 9: 131. 1880. 

Luzon: Province of Nueva Vizcaya, vicinity of Dupax, 
Januar>^ 14, 1913, -R. C. McGregor, Bur. Sci, 20231, on decaying 

Previously collected in East Africa. 

Panaeolus Fries 

Panaeolus papilionaceus (Bull.) Fr. Epicr. Myc. 236. 1836. 

Agaricus eguinus Alb. & Schw. Consp. Fung. Lus. 3. 1805. 

Agaricus campanulatus Linn. Sp. Plant. 1175. 1753- 
Agaricus carbon Batsch, Elench. Fung. 6. 1783. 
Agaricus papilionaceus Bull. Hist. Champ. France 561. 

Agaricus varius Pers. Icon. Des. Fung. 40. 1800. 
Panaeolus campanulatus Berk. Outl. Brit. Fung. 175. i860. 



H. M. 


Merrill 5008. Mindanao: District of Davao, March 5, 1904, 
Copeland 344, 

The formation of the new genus Copelandia, in which the 
sole distinction from Panaeolus is the presence of cystidia, 
is inviting for the future a confusion similar to that existing 
in the genera Panus and Lenlinus, where the only difference 
exists in the presence or absence of notched gills. This character- 
istic, even though at present confined to this one species, is 
not a sufficient distinction for a generic character, or to insure 

Graff: Philippine Basidiomycetes — V 233 

the possible acceptance of this as the type of a new genus. 
It seems the wisest course, therefore, to continue as before with 
the use of the Friesian name, at least till some character more 
positive may be found to warrant the change. 

Previously reported from the United States, Europe, South 


as the Phillippines. 

University of Montana, 
Missoula, Montana 

Occurrence of the pycnial stage of Puccinla Taraxacl* 


Louise Dosdall 

On April 29, 1920, dandelions were found on University Farm, 
St. Paul, heavily infected with rust. In addition to the fact that 
this was the earliest date on record at the University of the 
occurrence of Puccinia Taraxaci (Rabent.) Plow., this collection 
was of particular interest because pycnia were found accompany- 
ing the uredinia. As far as could be discovered from the lit- 
erature, this stage had never before been collected in North 
America, although both Plowright (1889) and Grove (1913) 

had reported its occurrence in England, and Jacky (1909) in 

In the spring of 1921 a careful watch was kept for the first 
appearance of dandelion rust to determine whether or not the 
pycnial stage was common in this region. The first pycnia 
were found on University Farm, April 17, 1921. Subsequently 
they were found in various localities around St. Paul and Minn- 
eapolis. If a careful search was made, the pycnial stage could 


A difference was 

noted in the abundance of the rust during the two seasons in 
which observations were made. In 1920 the pycnia were found 
accidentally in two localities. In both cases the infection on 
the plants was so conspicuous that the rust was observ^ed while 
one was walking hurriedly through the field. In 192 1 no pycnia 
could be found without the most diligent search. 

Through correspondence with Dr. H. S. Jackson, it was 
learned that the pycnial stage of Puccinia Taraxaci had been 
collected at Brookville, Indiana, in April, 1918, by C. A. Ludwig, 
and the specimens deposited in the Arthur Herbarium. This 
specimen was compared with the Minnesota collection and the 
rusts found to be Identical. In spite of the fact that collections 
are rare, it would seem that the occurrence of the pycnial stage 
must be rather common. 

The pycnia of Puccinia Taraxaci are amphigenous on yellow, 
slightly sw^ollen spots, which occur frequently along the midrib 

^ . ^ i_^ 

*Published, with the approval of the Director, as Paper No. 303 of the 
Journal Series of the Minnesota Agricultural Experiment Station. 


236 Dosdall: Pycnial stage of Puccinta Taraxaci 


of the leaf but are found sometimes on the blade. They are 
reddish brown in color, subglobose in shape, about 90[x in diameter 
and 76'tX in height. Ostiolar filaments are present. Uredinia 
appear on the same spots soon after the pycnia. 

Midwinter botanizing in southern Arizona 

Edwin B. Bartram 

As a large part of the literature relating to the desert flora 
of Southern Arizona is not readily accessible to any but pro- 
fessional botanists the following brief survey of the recognizable 
winter plants in the region tributary to Tucson may prove 
interesting and helpful to those fortunate enough to make the 

acquaintance of this fascinating country in the early months 
of the year. 

From a botanical point of view the area is obviously more 
closely related to the Mexican state of Sonora than it is to any 
of our possessions so that many of its most characteristic plants 
are not represented at all in Wooton and Standley's invaluable 
Flora of New Mexico. 

The preparation of specimens in this dry sunny climate is a 
simple operation; the surrounding mountains, or at least their 
foothills, are easy of access; and, as the representation of Arizona 
plants in most of our larger eastern herbaria is quite meagre, 
the ambitious collector finds an almost unlimited field in which 
to work. A surprising variety of flowers is evident even at this 
season and many plants whose period of bloom is past are re- 
cognizable by some perlstent vestige of fruit or flowers. 

Apart from such unique and unfamiliar plants as the sahuaro, 
chollas, candle bush, prickly pears and palo-verde, which claim 
the undivided attention of the newcomer until the novelty and 
charm of the first impressions are to a certain extent dulled by 
familiarity, the most striking feature of the desert vegetation is 
the isolation of various clearly defined plant communities. It is 
all desert, to be sure, but the wide gravel plains over whichCovillea 
gltitinosa is dominant and the barren spurs and talus slopes, 
sun baked, arid and almost devoid of plant life, excepting the 
hardiest drought-resisting species that have become inured to 
this environment, seem to resolve themselves readily into a 
series of barriers that from necessity restrict the distribution of 
many species to quite definitely circumscribed limits. 

Distances in Arizona are proverbially deceptive, but we were 




238 Bartram: Botanizing in southern Arizona 

least fifteen miles away Instead of under five miles as we had 
fondly imagined. The horses are willing and capable of covering 
thirty miles or more In a day without discomfort, but the first 
days experience proved conclusively that the one indispensable 
article of equipment was a good canteen; for, excepting a few 
canyons in the Catalina Range reaching well back Into the forested 
and snow-covered slopes, water is never available. The higher 
peaks are heavily snow-capped during most of the winter months, 
although snow rarely descends to the level of the plains. After 
a rainy night in Januarj^ it Is not unusual to see the snow line 
across the slopes as straight as though It had been drawn with 
a ruler. In the sheltered canyons where snow rarely falls and 
never persists we found many flowering plants In early January 
which were materialy increased in number and variety with 
each succeeding week. Many plants are distributed generally 
over entire ranges but in each canyon, walled in by the steep 
arid intervening ridges, we found some species that were wanting 
in every other locality we visited. 

Sabino Canyon in the Santa Catalina Mountains, distant 
some sixteen miles from Tucson, is one that we happened to 
reach quite frequently and will serve nicely as an example of 
these interesting localities. Leaving early we drive across the 
plains through miles of pungent greasewood with its curiously 
folded leaves and "bright yellow flowers, which even now are 
giving way to the little felted spheres of fruit. Thickets of 
cholla, six or eight feet high, varying in color from pale green to 
ruddy brown and brightened with pendant clusters of orange 
colored fruit, glow in the early sunlight. Cirsium neomexicantiniy 
Lesquerella Fendleri, and Baileya miiUiradiata are In bloom along 
the roadsides, the stiff wands of old fruit bend over colonies of 
gray-green A triplex canescens, and here and there an occasional 
Yucca elata lifts its brown panicle of old fruit above the shrub 
level. In a broad belt along the base of the mountains the 
sahuaro gives a grass green tinge to the foothills ten miles away 
where the arroyos are marked by wavering lines of mesqulte and 
desert willow. The hot rocky slopes flanking the mouth of 
the canyon are spotted with Opttntia Bigelovii, glistening like 
spicules of glass In the brilliant sunshine and undoubtedly the 
most vicious of all the chollas. A carpet of detached joints 
covers the ground about every plant, and woe to the unwary 

Bartram: Botanizing in southern Arizona 239 

walker who steps unguardedly into these precincts. The spines 
penetrate shoe leather like a steel needle and. refuse to be with- 
drawn. Nature guards her wards well in these desert places and 
the animals, profiting by the example, guard their dens with a 
pile of these bristling joints, carried one by one at a cost easy 
to be reckoned by anyone unfortunate enough to have brushed 
against a stem in an unguarded moment. 

Along the creek issuing from the canyon are many waifs 
from the higher zones, such as Juniperiis scopidoriim, Yucca 
baccate, Qttercus Emoryij Q. ohlongifolia and Q.hypoleuca, hut it is 
among the rocks on sheltered slopes in the canyon that we find 
the earliest flowers. Here the fair>' dusters, Calliandra eriophylla, 
are just spreading their delicate balls of pink bloom, and clumps 
of LesqiiereUa purpurea are established In many congenial crev- 
ices. Hermannia pauciflora with the flowers well concealed 
among the matted woody stems is easily overlooked, and a form of 
Anislotus puhernhis with showy golden yellow flowers and nearly 
leafless stems helps to brighten the otherwise barren ledges. 
Notholaena Lemmoni grows abundantly among the rocks of a 
dry sunny slope, while the shaded crevices and fissures give 
harbour to such species as Pellaea rmicronata, Cheilanthes Lind- 
heimeri and various mosses, among which Tortida inerniis, T, 
ruralisy Barbida Manniae and B. chloronolus are the most in 
evidence. Where almost any other living thing would wither and 
perish Agave Parryi and Dasylirion Wheeleri spread over the 
ledges and boulders in flourishing clumps that lend a charac- 
teristic touch to this unique landscape. 

The region about Picture Rocks was another productive 

locality that proved interesting enough to warrant trips at 
intervals of about ten days. Some twelve miles west of Tucson, 
near the far end of the Tucson Mountains in a broad gully 
fronting the old Indian pictographs, from which the place takes 
its name, Anemone sphenophylla was just coming into bloom on 
January tw^elfth, white clusters of Dryopefalon runcinalum 
brightened many of the damp rock fissures, the trim little 
Thysanocarpus amplectens grew sparingly in nearly every 
depression where sufiicient soil had accumulated to support a 
short span of life during the season of winter rains, while various 
winter annuals, such as Lepidium lastocarpum, Draha ciineiJoUa 
and Amsinckia Afen^i mi, were scattered sparingly in the shelter 

240 Bartram: Botanizing in southern Arizona 

of shrubs or rocks wherever a bit of moisture and shade was 
obtainable. Along the dry arroyos leading off through the 
mountains Anislotus hrachycarpus and A. trispermus were just 
opening their first flowers, and on the dry flanking talus slopes 
the omnipresent Encelia farinosa in full bloom was temporarily 
forgotten in the pleasure incident to finding a patch of Polygala 
macradenia with inconspicuous little purple flowers barely 
visible even at close range. 

A similar association of species, including DryopetaloUy 
Anemone and Thysanocarpus^v^^soh^orved on the shaded side 
of a steep ridge at Robles Pass^ where scattered pockets of soil 
had been formed by the disintegrating mosses and lichens, and 
no doubt a more thorough survey of the region would disclose 
many similar localities; but the complete isolation of these 
stations, one from another, is the one factor that seems to be 
repeatedly emphasized. 

Theorizing is always interesting, often productive, but some- 
times misleading, yet in the light of these facts one can hardly 
reject the inference that at some comparatively recent time the 
forms of plant life, not fitted to thrive in the arid gravel and 
rock formations of the desert as we find it today, were more 
widely distributed than they are now, and by progressive changes 
have been isolated by the rising tide of desert vegetation. 
Such a condition is plain in outline but difficult to translate 
into terms that are applicable w^hen the available data are so 
meagre. Inequality in the distribution of rainfall, the processes 
of erosion and disintegration and variations in climatic conditions 
have been some of the contributing causes to the spread and 
contraction of the various plant communities as we now know 
them, but to w^hat extent these influences have operated and in 
what manner the distribution has been modified by complicated 
cross currents of action and reaction are queries that may be 
more satisfactorily answered at some future time, when our 
knowledge of the subject will be more intimate and compre- 
hensive than it is now. 

The following list of species, collected between December 
23d, I9I9) ^i^d January 26th, 1920, may be considered as 
fairly representative of the region within a radus of fifteen 
miles from Tucson but is necessarily incomplete. Many of the 
plants listed w^ere in flower and nearly all of them provided with 

Bartram: Botanizing in southern Arizona 241 


fruit m a sufficiently good state of preservation to make them 
easily recognizable. 

For assistance in the determination of many of the species 
I am much indebted to the kindness of Dr. Paul C. Standley. 

Cheilanthes Lindheimeri Hook. Frequent on arid ledges and rocky 

Cheilanthes myriophylla Desv, Less common than the preceding. 

Cheilanthes Pringlei Davenp. The finely dissected delicate frondj 
of this species distinguished ic at once from its associates in this region. In" 
frequent in shaded rock clefts on the lower slopes. 

Cheilanthes Wrightii Hook. Inf/equent but easily recognized by 
the smooth green pinnules and lustrous channelled stipes. 

NoTHOLAENA HoOKERi D. C. Eaton. Rather common among rocks 
on arid hillsides. The characteristic outline of the fronds and the bright 
yellow color of the lower surface are unmistakable marks.' 

Notholeana sinuata (Sw.) Kaulf. Frequent among dry rocks on the 
lower mountain slopes. 


Notholaena sinuata integerrima Hook. More widely distributed 
than the typical form of the species. 

Notholaena bonariensis (Willd.) C. Chr. Rare among rocks near 
the mouth of Sabino Canyon. 

Notholaena Lemmoni D. C. Eaton. Abundant on a dry rocky slope 
in the mouth of Sabino Canyon but not found elsewhere. 

Pellaea mucronata D. C. Eaton. Ledges and crevices in the foothills. 
This seems to be the commonest fern of the region. 

Selaginella arizonica Maxon. A recently described species forming 
flat green mats among rocks in the arroyos. 

Selaginella ruficola Underw. The grayish color and erect habit 
are good field marks for this species. Near the mouth of Sabino Canyon 
it forms dense turfy mats over the dry ledges. 

Juniperus scopulorum Sarg. Bank of stream near the mouth of 
Sabino Canyon. Evidently a stray from the higher ridges. 

Ephedra trifurcata Torr. Abundant along washes in the gravel 

Ephedra antisyphilitica Meyer. Rare in an arroyo near Picture 
Rocks. The slender flagellate stems give this plant a very different aspect 
from the more common, stiffly erect, E. trifurcata. 

Ephedra viridis Coville. Collected but once in the dry foothills at 
Robles Pass, 

Andropogon saccharoides Sw. Frequent on the plain and foothill. 

Heteropogon contortus (L.) Roem. & Schult. More confined to the 
foothill slopes. 

242 Bartram: Botanizing in southern Arizona 

Chaetcchloa machrostachya (H. B. K.) Scribn. & Merr. Rare on 
talus slopes at Robles Pass. 

Chaetcchloa viridis (L.) Scribn, Introduced in fields and waste places. 
Aristida ADSCENSiONis L. Frequent on hillsides in the Tucson Mount- 


Aristida Wrightji Nash. Uncommon on dry slopes in the Tucson 

Aristida bromoides H. B. K, Common in the foothills. Spikelets 
developing through the winter. 

Aristida divergens Vasey. Dry slopes in the mouth of Sabino Canyon. 

Muhlenbergia dumosa Scribn, A unique species confined to ledges 
along the trail in Sabino Canyon at about 3000 feet elevation. 

Muhlenbergia microsperma (D C.) Kunth. Rare in the dry foothills. 

Muhlenbergia Porteri Scribn. Common in the shelter of shrubs 
on the gravel plains. 

Epicampes ringens Benth. Slopes in the mouth of Sabino Canyon. 

Bouteloua filiformis (Fourn.) Griffiths. Flowering plants collected 
in Sabino Canyon on January 3d. 

Bouteloua Rothrockii Vasey. The dark brown spikelets persist 
well through the winter. 

Bouteloua Parryi (Fourn.) Griffiths. This and the preceding species 
are abundant near Robles Pass. 

Trichloris fasciculata Fourn. Roadsides south of Tucson, 

Dasyochloa pulchella (H. B. K.) Willd. Generally distributed over 
the plains. 

Koeleria cristata (L.) Pers. Along streams in the foothills of the 
Santa Catalina Mountains. 

Eragrostis megastachya (Koel.) Link. A weed of waste places. 

PoA Bigelovii Va^ey & Scribn. Confined to damp ledges near Picture 

Cyperus cyrtolepis Torn & Hook. Edge of stream east of Pima 


JUNCUs NODOsus L. Edge of stream in Pima Canyon. 

Yucca elata Engelm. Single plants scattered over the plains but 
nowhere abundant. 

Yucca baccata Torn Thickets in the mouth of Sabino Canyon. Ap- 
parently washed down from the higher levels. 

Nolina microcarpa S. Wats. With the preceding, 

Dasyliriox Wheeleri S, Wats. On dry ledges in the foothills. 

DiPTEROSTEMON PAUCIFLORUS (Torr.) Rydb. Sparingly in flower along 
an arroyo near Picture Rocks. 

Agave Parryi Engelm. On dry ledges in the mouth of Sabino canyon. 

Bartram; Botanizing in southern Arizona 243 

PoPULUS WiSLiZENi (S, Wats.) Sarg. Catkins fully developed and 
falling by late January. 

QuERCUS ARizoNiCA Sarg. Scattered along the creek near the mouth 
of Sabino Canyon. 

QuERcus Emoryi Torn With the preceding but more abundant and 

QuERcus HYPOLEUCA Engelm. Sterile but easily recognized by the 
leathery leaves densely white woolly beneath. 

QuERcus OBLONGIFOLIA Torr. This and the preceding species of oaks 
are all confined to the lower end of Sabino Canyon, where they seem to exist 
as waifs carried down by the stream from higher levels in the region about 
Mt. Lemmon. 

Celtis pallida Torr, Frequent along arroyos in the Tucson Mountains. 

Parietaria obtusa Rydb. Localized on a shaded ledge near Picture 

PhoradeM)ron macrophyllum (Engelm.) Cockerell. Abundant on 
Poptdus in the plains and foothills. 

Phoradendrox cahfornicum Nutt. The abundant small red berries 
and bare stems make a very attractive combination. Chiefly on Mesqulte in 
the plains. 

Eriogonum polycladon Benth. Recognizable but long past flowering. 

EriogoN'um pinetorum Greene. Long past flowering but leaves and 
fruit persistent enough for identification. 

Eriogonum Wrightii Torr. Widely distributed over arid rocky 
slopes in the foothills. 

RuMEX hy'menosepalus Torr. Common In sandy areas throughout 
the plains. 

Polygonum punctatum Ell. Edge of stream in the foothills of the 
Santa Catalina Mountains. 

Atriplex cakescens (Pursh) Nutt. Several forms of this species 
are abundantly over the plains. 

Atriplex rosea L. Young plants in the sandy bed of an arroyo near 


Picture Rocks. 

Chekopodium pagaxum Reichenb. Sparingly introduced in waste places* 

Amaranthus paniculatus L. a common weed of fields and waste 

Wedeliella incarnata (L.) Cockerell. Flowering sparingly in sunny 
talus slopes in the foothills. 

Hesperoxia retrorsa (Heller) Standi. Dry slopes near Picture Rocks. 

Commicarpus scandens (L.) Standi. Localized in rock clefts along 
an arro>o west of Tucson. 

Axemoke sphekophvlla Poepp. Frequent among rocks along arroyos 
in the foothills. In full flower by mid-January. 

244 Bartram: Botanizing in southern Arizona 

Clematis Drummondii Torr. & Gray. Frequent on the plains west of 


EscHSCHOLTZiA MEXICANA Greene. A few flowers on sheltered banks 
in January. 

Thysanocarpus amplectens Greene. Locally abundant in damp 
pockets near Picture Rocks and at Robles Pass." Sparingly in flower on 
January 12th but plentiful and in mature fruit ten days later. The curious 
disk-shaped pods with perforated wing margins mark this off at once from 
all other crucifers of the region. 

Strepanthus arizonicus S. Wats. Frequent in the shade of bushea 
near Picture Rocks, 

Lepidium hirsutum Rydb. Infrequent on lower slopes of Santa Cat- 
alina Mountains. 


Lepidium lasiocarpum Nutt. Common in sandy plains and dry washes. 

Lesquerella Fendleri (Gray) S. Wats. Flowering early along dry 
roadsides and on open slopes in the foothills. 

Lesquerella purpurea (Gray) S. Wato. An attractive early flower 

of rock crevices in the foothills and lower slopes of the canyons. In vigor- 
ous tufts the stems sometimes attain a length of nearly two feet. 

Draba cuneifolia Nutt. Widely distributed on shaded knolls and 
banks along the arroybs. 

Arabis perennans S. Wats. The purple flowers give quite a touch of 
color to the ledges along the arroyos west of Tucson. 

Brassica nigra (L.) Koch. Occasional in cultivated fields. 

Sophia ochroleuca Wooton. Frequent in sheltered shady spots. 

Dryopetalon runcinatum Gray. Rather local in damp shaded 
rock pockets. The delicate white flowers form a pleasing contrast with their 
rough arid setting. 

Sedum Griffiths!! Rose. Ledges in Sabino Canyon. 

Fendlera ruficola Gray. Ledges in Sabino Canyon. Leaves and 
old fruit persistent. 

Philadelphus micrpohyllus Gray. Frequent on open rocky slopes 
in the foothills. In sheltered places the flowers were opened in profusion by 
mid January. 


In the mouth of Sabino Canyon. 

Vauquelina californica (Torr.) Sarg. Ledges in Sabino Canyon at 
about 300 feet elevation. 

Calliandra eriophylla Benth. One of the most attractive of the 
early spring flowers. Abundant on open rocky slopes in the foothills. 

Acacia coxstricta Benth. Frequent on the plains. Pods more per- 
sistent than A. Greggii, 

Acacia Greggii Gray. This and A, constricla are characteristic 
thorny shrubs of the gravel plains. 

Bartram: Botanizing in southern Arizona 245 

Prosopsis glandulosa Torr. This is the main source of firewood for 

the region. 

AcuAN Jamesii (Torr. & Gray) Kuntze. Pods persistent through the 

Cassia Covesii Gray, Scattered sparingly over the open gravel plains. 
Pods and leaves persistent. 

Hoffmanseggia densiflora (Benth.) Gray. On sandy plains. Pods 

Parkensonia microfhylla Torr. A thorny shrub or small tree of 
the foothills and adjacent plains. On sheltered sunny slopes a few flowers 
were in evidence by late January. 

Cercidium Torreyanum Sarg. A low nearly leafless shrub occuring 
sparingly along washes in the plains. 

Krameria glandulosa Rose & Painter. A straggling flowerless shrub 
of the plains with soft silky pubescent young growth. 

Anislotus brachycarpus (Benth.) Rydb. The broad leaflets and 
villous pubescence separate this species sharply from the following where 
they grow together. 

Anislotus trispermus (Greene) Woot. & Standi. With the preceding 
in a sandy bed of an arroyo near Picture Rocks. Just coming into flower. 

Anislotus puberulus (Benth.) Woot. & Standi. What seems to be 
a form of this species was in full flower on the ledges of Sabino Canyon at 
about 3000 feet elevation. 

Parosella Parryi (Torr. & Gray.) Heller. This inconspicuous little 
plant is quite typical of arid rocky hillsides in the foothills. 

Parosella Greggii (Gray) Heller. Abundant on rocky spurs and slopes 
near Agua Calgiente but not observed elsewhere. 

NissoLlA ScHOTTii (Torr.) Gray. Pods and leaves persistent. Frequent. 

Astragalus Nuttallianus DC. Local on the plains. In flower 
in January. 

Cracca tenella Gray. Localized on dry sunny ledges near Robles Pass. 

Galactia Wrightii Gray. Twining over low bushes in the foothills. 
Leaves and pods persistent. 

Olneya tesota Gray. Foothill slopes and nearby plains. 

Erodium cicutarium (L.) L'Her. Abundant in waste places and 
on open sandy plains. 

Erodium texanum Gray. Gravelly Knolls near Picture Rocks. 

Covillea glutinosa (Engelni.) Rydb. The most abundant and char- 
acterisitc shrub of the plains. 

KoEBERLiNiA SPINOSA Zucc. Frequent on knolls and in thickets in 
the plains. 

JANUSIA GRACILIS Gray. Twining about bushes on open rocky hillsides. 

246 Bartram: Botaxizing in southern Arizona 

RuTOSMA PURPUREA Woot. & Standi, Frequent on banks of the 
arroyos west of Tucson. 

PoLYGALA MACRADENIA Gray. Confined to a rocky hillside near Picture 

Euphorbia cafitellata Engelm. Forming quite showy mats on ledges 
and in crevices along the arroyos west of Tucson. 

Euphorbia pediculifera Engelm. A form with densely glandular 
pubescent stems and leaves is not uncommon. 

Euphorbia polycarpa Benth. This and the two preceding species 
of the genus are freely distributed over the arid ridges and talus slopes. They 
all belong to the group of perennials with entire pubescent leaves and vary 
considerably in habit and appearance. 

DiTAXis neomexicana (Muell. Arg,) Heller. The lax straggling habit 
of this species is very different from the stiff erect form of D, sericophylla. 

DiTAxis SERicoPHYLLA (Gray) Heller. With the preceding on rocky 
slopes at Robles Pass. 

Callitriche palustris L. In shallow water at the mouth of Sabino 

SiMMONDSiA CALIFORNICA Vutt. A Striking evergreen bush of the dry 
plains and foothills with leathery leaves and yellow flowers in drooping 
axillary clusters. 

DoDONAEA viscosA L. Abundant along the stream bank in the mouth 
of Sabino Canyon. 

CoNDALiA SPATHULATA Gray. Banks of arroyos near the foothills. 
Leaves and fruits persistent. 

Rhamxus tomentella Benth. Ledges in Sabino Canyon at about 
3000 feet elevation. 

Hibiscus Coulteri Harv. Flowering sparingly through the winter. 

Hibiscus denudatus Benth. The pale almost leafless stems and 
delicately tinted pink flowers give this plant an attractive personality. Rare 
on open slopes at Robles Pass. 

Gayoides crispum (L.) Small. Frequent on rocky slopes. 

AeuiiLON INCANUM (Link) Sweet. Common on rocky hillsides. Leaves 
and carpels persistent. 

Sphaeralcea ambigua Gray. A polymorphous species widely scattered 
over the region. Flowers white or pink and leaves varying greatly in outline. 

Sphaeralcea grossulariaefolia York. On rocky slopes at Robles Pass. 

Hermanxia pauciflora S. Wats. Confined to rocky slopes near the 
mouth of Sabino Canyon. 

Ayenia pusilla L. Frequent on the dry gravel plains. 

Fouquieria splendens Engelm. Although the leaves were well developed 
in sunny sheltered places no flowers were in evidence by the end of January. 

QPUNTIA FULGiDA Engelm. Common on the plains. 

Bartram: Botanizing in southern Arizona 247 

Opuntta arborescens Engelm. Forming impenetrab!e thickets on 
the plains, 

Opuxtia leptocaulis DC. The common slender stemmed cholla 
of the region. 

Opuntia Bigelovii Engelm. Frequent on arid rocky slopes in the 

X ■ P 

Opuxtia cholorotica Engelm. & Bigel. Oa ledges near the mouth of 
Sabino Canyon, 

Opuntia Toumeyi Rose. Frequent on the gravel plains. 

Mamillaria Grahami Engelm. Frequent in small colonies among 
rocks in the foothills. The diminutive scarlet pipe-shaped fruits are v^ery 

Echinocactus Wislizeni Engelm. Scattered sparingly over the foot- 
hill slopes. Some plants are well over a meter in height. 

Echinocereus Fendleri (Engelm,) Riimp! Forming clumps in 
Rravel plains near the foothills. 

Cereus giganteus Engelm. The sahuaro well merits the reputation of 
being one of the most unique and striking elements of the desert vegetation. 

Daucus pusillus Mx. Scarce on shaded knolls in the washes. 

Plumbago scandens L, In the foothills of the Santa Catahna Moun- 
tains. Leaves and fruit persistent. 

FoRESTERiA PHILLYREOIDES Torr. This is a Mexican type and Dr. 
Standley informs me they have but one other specimen from the United 
States, collected at Tucson by Pringle, The staminate flowers were fully 
developed by December 30th and a few leafy shoots supplied the foliage 
characters. On rocky slopes along the Ajo road west of Tucson. 

Fraxixus attenuata Jones. In the mouth of Sabino Canyon. 

Menodora scabra Gray. On arid slopes of the Tucson Mountains 

Haplophyton cimicidum a. DC. Rare on rocky talus slopes at 
Robles Pass. 

AscLEPiAS linaria Cav. Frequent on ledges in the lower slopes of 
the Santa Catalina Mountains, 

Philibertella linearis Gray. Twining over bushes on dry rocky hill- 

EvoLVULus LINIF0LIU5 L. An inconspicuous little perenial with fugitive 
flowers. Frequent among rocks and dry ledges in the foothills. 

Jacquemontia Pringlei Gray. Not uncommon on rocky hillsides. 
Leaves and capsules persistent. 

Pectocarya linearis (R. & P.) DC. Locally abundant on a gravel 
shelf near Picture Rocks. 

Amsinckia Menziesii (Lehm.) Nels. & Macb. Flowering with Thys- 
anocarpus and Vryopetalon on shaded ledges at Picture Rocks. 

248 Bartram: Botanizing in southern Arizona 

Verbena Wrightii Gray. This is one of the showiest of the early 
flowers in the foothills, where it blooms profusely in January. 

Verbena xutha Lehm. Rare and local on dry slopes of the Tucson Mts. 

LiPPiA Wrightii Gray. Rocky slopes in the foothills. Leaves and 
nutlets persistent. 

Marrubium vulgare L. a very common weed of roadsides and waste 

Mesosphaerum Emoryi (Torr.) Kuntze. Flowering through the winter 
and very characterisric of arid rocky slopes in the foothills. 

Hedeoma oblongifolia (Gray) Heller. Ledges in the mouth of Sabino 

Hedeoma nana (Torr.) Greene. In rock crevices along arroyos west 
of Tucson. The leaves and fruit of this and the preceding are persistent 
through the winter. 

Salvia pinguifolia (Fernald) Woot. & Standi. On rocky hillside at 
Robles Pass, Buds well developed but not in flow^er. 

NICOTIANA trigonoph\-lla Dunal. A common plant of dry ledges in 
the foothills. In flower throughout the winter. 

NICOTIANA glauca Graham. An introduced shrub of waste places. 

Lycium Cooperi Gray. Frequent on rocky slopes. 

Lycium Fremontii Gray. Ledges at the mouth of Sabino Canyon. 
In full flower on January 6th. 

Lycium parviflorum Gray. A thorny straggling shrub of the gravel 

LvciuM Torreyi Gray. Confined to the slopes near Picture Rocks. 

Physalis ixocarpa Brot. Infrequent in the Tucson Mountains. 

Antirrhinum antirrhiniflorum (Willd.) Hitchc. On slopes of the 
Tucson Mountains. Leaves and capsules persistent. 

ScROPHULARiA PARVIFLORA Woot. & Standi. Flowering sparingly on 
January 20th In the mouth of Pima Canyon, Santa Catalina Mountains. 

Pentstemon superbus a. Nels. Sandy washes in the foothills. Flow- 
ering freely in January. 

MiMULUS Langsdorfii Don. Shallow water in a brook east of the 
mouth of Pima Canyon. 

Mecardonia peduncularis (Benth.) Small. Abundant In a brook 

east of the mouth of Pima Canyon. This species seems to flower and mature 
fruit throughout the winter. 

Stemodia durantifolia (L.) Sw. In shallow water with the two 


DiAPEDiuM Torreyi (Gray) Woot. & Standi. Arid slopes near Agua 
Caliente. The large cordate bracts, fading white and papery, give this plant 
a unique appearance in the field. 

Bartram: Botanizing in southern Arizona 249 

CARLO\yRiGHTiA ARizoNiCA Gray. Leaves and capsules persistent 
enough for identification, 

SiPHONOGLOSSA LOXGIFLORA (Torr.) Gray. Frequent on arid slopes 
west of Tucson. Leaves and capsules persistent. 

RuELLiA TUBEROSA occiDENTALis Gray. Infrequent near Picture Rocks- 
Old capsules persistent, 

Chilopsis linearis (Cav.) Sweet. Along arroyos in the foothills- 

Stenolobium iNCrsuM Rose & Standi. Rocky ledges and crevices 
among the foothills. The persistent leaves and showy yellow capsules make 
this a clean and attractive plant throughout the winter. 

Galium stellatum Kellogg. This plant makes considerable growth 
from a diffuse woody base during the winter, but there was no evidence of 
bloom by January 30th. Frequent on ledges and rocky hillsides. 

Sambucus mexicana Presl. Frequent along water courses and in 
cultivation. In bloom by mid- January, 

Ptiloria pauciflora (Torr.) Raf. A common plant of the arid plains 
and foothills blooming through the winter. 

Perezia Wrightii Gray. In full flower on January 21st. Infrequent 
on arid rocky slopes near Picture Rocks. 

Trixis californica Kellogg. In full flower during January, Abundant 
and quite showy among rocks in the foothills. 

Hymenoclea monogyra Torr. & Gray. A tall slender weed-like 

shrub with persistent filiform leaves and winged fruit. In the mouth of Pima 

Franseria deltoidea Torr. One of the characteristic plants of dry 
gravel plains near the foothills. Buds well developed by January 30th 
but no flower. 

EuPATORiUM solidagixifolium Gray. Ledges at the mouth of Sabino 

Coleosanthus baccharideus (Gray) Kuntze. Infrequent on ledges 
near the mouth of Pima Canyon. Leathery leaves and flower heads very 

Coleosanthus Coulteri (Gray) Kuntze. Common on rocky hillsides, 

Coleosanthus Wrightii (Gray) Britton, Ledges in the mouth of 
Sabino Canyon. 

Gymxosperma COrymbosum DC. Confined to the mouth of Pima 

Gutierrezia glomerella Greene. Leaves and old heads persistent. 
Frequent on the plains. 

Sideranthus australis (Greene) Rydb. Common on the gravel plains. 

IsocOMA Hart\vegi (Gray) Greene. Forming dense clumps on ridges 
and knolls in the gravel plains. 

Chrysoma laricifolia (Gray) Greene. Rock clefts and ledges in the 
arid foothills. Flowering casually through the winter months. 

250 Bartram: Botanizing in southern Arizona 

Baccharis glutinosa Pers. Common along the banks of Rillito Creek- 

Baccharis sarothroides Gray. Abundant on the sand bars and 
banks of Rillito Creek. 

Erigeron divergens Torn & Gray, This is apparently an abnormal 
winter form of the species but very constant throughout the region. It is 
scattered freely over the rocky foothill slopes and flowers during January. 

Gnaphalium Wrightii Gray. Frequent on arid rocky slopes. 

Parthenium incanum H. B. K. A common low shrub of the foothills. 
In good conditions through the winter, 

Crassina pumila (Gray) Kuntze. Occurring sparingly on dry gravelly 

Encelia farinosa Gray. The distinctive gray color of many of the 

desert slopes is derived from the lustrous silvery leaves of this species. The 

showy yellow flowers are locally abundant, even in January, and it may 

well be numbered among the most satisfying and attractive plants of the 

Leptosyne arizonica Gray. Bank of a stream in the mouth of Pima 

PoROPHYLLUM JUNCIFORME Greene. A rush-like plant of ledges and 
rocky slopes in the foothills. 

Dysodia porophylloides Gray. Rare on ledges near Picture Rocks. 

Psilostrophe Cooperi (Gray) Greene. Fresh flowers among the 

persistent papery old blooms are found sparingly in January. Frequent on 
the gravel plains. 

Baileya multiradiata Harv, & Gray. Frequent and showy in 
sandy plains, especially along roadsides. Flowering freely by late January. 

Perityle dissecta Gray. An attractive little plant with curiously 
cleft leaves and persistent involucres. In rock clefts near the mouth of 
Pima Canyon. 

Bahia dealbata Gray. Arroyos In the Tucson Mountains. 

Helenium montanum Nutt. Localized along the bank of a stream 
east of the mouth of Pima Canyon. 

Artemisia albula Wooton. Arid foothill slopes Abundant near 
Agua Caliente. 

Artemisia Brittonu Rydb. Ledges in Sabino Canyon. 

Bebbia juncea (Benth.) Greene. An attractive slender stemmed 
plant of the sandy plains. 

Senecio Lemmom Gray. Flowering rather sparingly on rocky slopes 
in the foothills. 

CiRSiUM neomexicanum Gray, Infrequent along roadsides in the 
gravel plains. 

Bushkill, Pennsylvania 

Two new genera of Cactaceae 

N. L. Britton and J. N. Rose 

In our recent study of the genus Echinocactus , we have 
segregated a number of new genera, and restored several previ- 
iously pubHshed, among which are Malacocarpiis, Gymnocalycium , 
and Discocacius, all composed of species native to South America. 
Among the North American segregates are the two described 
below, which do not belong even to the same subtribe as Echi- 



Echinocacttis subgenus Thelocactus Schumann, Gesamtb- 
Kakteen 429. 1898. 

Cacti of medium size, one-jointed, solitary or caespitose, 
globose or somewhat depressed, spiny, often densely so; ribs 
few, low or even indistinct, often spiraled, divided into large 
tubercles, the flowering ones more or less grooved above; flowers 
from near the center of the plant, borne on very young tubercles, 
rather large for the subtribe, campanulate, diurnal; scales on 
the ovar>' usually few, their axils naked; fruit, so far as known, 
dry, dehiscing by a basal pore; seeds numerous, black, finely 
tuberculate, with a large basal hilum. 

Type species: Echinocacttis hexaedropJiorus Lemaire. 
The generic name refers to the tubercled ribs, 

1. Thelocactus hexaedrophorus (Lemaire) comb. nov. 
Eckinocacius hexaedropJiorus Lemaire, Cact. Gen. Nov- Sp. 

27. 1839. 

2. Thelocactus bicolor (Galeotti) comb, nov. 
Echinocactiis bicolor Galeotti in Pfeiffer, Abbild. Beschr. Cact. 

2: pi ^5. 1848. 

3 Thelocactus lophothele (Salm-Dyck) comb, nov. 




/ NEOLLOYDIA gen. nov. 

Small, more or less caespitose plants, fibrous-rooted, cylindrlc, 


ribs, grooved above; radial spines numerous, widely spreadmg; 


Brixton & Rose: New genera of Cactaceae 

central spines one to several, much stouter and longer than 
the radials; flowers large, pink or purple, subcentral from the 
axils of nascent tubercles, their segments widely spreading; 
fruit compressed-globose, dull colored, thin-walled, becoming 
papery, dry, with few scales or none; seeds globose, black, dull, 
tuberculate-roughened, with a large light basal scar. 

Type species: Mammillaria conoidea De CandoUe. 

The genus is dedicated to Professor Francis E. Lloyd 


I. Neolloydia conoidea (De Candolle) comb. nov. 

Mammillaria conoidea De Candolle, Mem. Mus. Hist, Nat, 
Paris. 17: 112. 1828. 

Echinocactus conoidetis Poselger, AUg. Gartenz. 21: 107. 1853, 

2/ Neolloydia Beguinii (Weber) comb. nov. 

Echinocactus Beguinii Weber in Schumann, Gesamtb, 
Kakteen 442. 1898. 




The aim of this Index is to include all current botanical literature written by 
Americans, published in America, or based upon American material; the word 
America being used in the broadest sense. 

Reviews, and papers that relate exclusively to foresty, agriculture, horti- 
culture, manufactured products of vegetable origin, or laboratory methods 
are not included, and no attempt is made to index the literature of bacteriology. 
An occasional exception is made in favor of some paper appearing in an Ameri- 
can periodical which is devoted wholly to botany. Reprints are not mentioned 
unless they diflfer from the original in some important particular. If users 
of the Index will call the attention of the editor to errors or omissions, their 
kindness will be appreciated. 

This Index is reprinted monthly on cards, and furnished in this form to 
subscribers at the rate of three cents for each card. Selections of cards are 
not permitted; each subscriber must take all cards pubUshed during the term 
of his subscription. Correspondence relating to the card issue should be 
addressed to the Treasurer of the Torrey Botanical Club. 

Andrews, F. M. The effect of aeration on plants, Proc. Indiana 
Acad. Sci, 1920: 147, 148. 1921. 

Andrews, F, M. The effect of centrifugal force on plants. 
Proc. Indiana Acad. Sci. 1920: 143-145. 192 1. 

Andrews, F. M. Phyllotaxis of Spectdaria perfoliata. Proc, 
Indiana Acad. Sci. 1920: 149, 150. 192 1, 

Andrews, F. M. Studies on Pollen III. Proc. Indiana Acad. 
Sci. 1920: 155-156. 1921. 

Baerthlein, K. Uber bakterielle Variabilitat, insbesondere 
sogenannte Bakterienmutationen. Centralbl. Bakt. Par- 
asitenk. u. Infekt. 81^: 369-435. 31 Jl 1918. 

Baker, F. C. The life of the Pleistocene or Glacial Period, 
Univ. Illinois Bull. 17: 1-476. pL 1-56. 7 Je 1920. 

Bardell, E. M. Production of root hairs in water. Univ. Wash* 
ington Fubl. Bot. 1: 1-9. i F 1915. 

Barrus, M. F. Bean anthracnose. Cornell Univ. Agr. Exp. 
Sta. Mem. 42: 101-214. pLi-8+f. lo-ig. Jl 1921. 


254 Index to American botanical literature 

Barton, J. E. The amount of standing timber in Kentucky. 
Min. and For. Resources Kentucky V. 1: 251-284. 

Beccari, O., & Rock, J. F. A monographic study of the genus 
Pritchardia. Mem. B. P. Bishop Mus. 8: 1-77. pL' 1-24. 


Includes 13 new species from the Hawaiian Islands. 

Belling, J. The behavior of homologous chromosomes in a 
triploid canna. Proc. Nat. Acad. Sci. 7: 197-201. /. i. 2. 
Jl. 1921. 

B[enedict], H. M. Further consideration of the sizes of vein 
islets of leaves as an age determinant. Science II. 55: 399, 
400. 14 Ap 1922. 

Benedict, R. C. Is Botrychiiim dissectiim a sterile mutant of 
B. obliquum? Am. Fern. Jour. 11: 53"55- 1921. 

Benedict, R. C. Nephrolepis nutrition. Am. Fern Jour. 11: 
41-43. 1921. [lUust.] 

Berry, E. W. Tertiary fossil plants from Venezuela. Proc. 
U. S. Nat. Mus. 59: 553-579- pl^ 107-109, 1921. 

Bowman, H. H. M. Histological variations in Rhizophora 
Mangle, Michigan Acad. Sci. Rep. 22: 129-134. pL g-12. 

Au 1921. 

Britton, N. L. George Valentine Nash. Jour. New York Bot. 
Gard. 22: 145-147. pL 261. Au 1921. 

Brown, F. ^B. H. Origin of the Hawaiian flora. Proc. Pan- 
Pacific Sci. Confer. 1: 131-142. 1921, 

Brown, L. L. Canadian Sitka spruce, its mechanical and phy- 
sical properties. Canada Dept, Inter. For. Bull. 71: i-39- 
/. 1-20, 1921, 

Brunswik, H. Uber Hesperidinspharite im lebenden Hautgewebe 
von Antherium Binotii Lenden, Ber, Deutsch. Bot. Ges. 
39: 208-212. Je 1921. 

Buscalioni, L,, & Muscatello, G. Studio monografico sulle 

specie americane del gen. ^' Sauratiia^^ Willd. Malpighia 

29: 1-32; 97-112. 1921. 

Includes Saxiratda intermedia sp. nov. from Guatemala. 

IxDEx TO American botanical literature 255 

Buscalioni, L. II legno crittogamico del fasclo vascolare seminale 
di talune angiosperme considerate nei suoi rapporti colle 
moderne teorle filogenetiche. Malpighia 29:46-80; 113-207. 
f^i-30. 1921. 

Butters, F. K. A new western species of Pellaea, Am- Fern 
Jour. 11: 39, 40. 192 1. 

Pellaea Su'ksdorfiana from Washington. 

Butters, F. K* Salvinia in Minnesota. Am. Fern Jour. 11: 
48-50. 1 92 1. 

Chapman, G, H. Tobacco investigations. Massachusetts Agr. 
Coll. Bull. 195: 1-38. pl.i~2,B. [1920] 

Chiovenda, E. La culla del cocco. Webbia 5: 199-294. 1921. 

Concerning the origin of the coco-palm. 

Chrlstensen, C. An overlooked species of Dryopteris Am. 
Fern Jour. 11: 44-46. 1921 

Dryopteris crypta (Underw. & Maxon) from Cuba. 

Conzatti, C, Monografia del Arbol de Santa 


Pp. 1-65. Oaxaca, Mexico. Mr 192 1. [Illust.] 

Account of a giant tree of Taxodium mucronatum Ten. 

Coons, G. H., & Levin, E. The relation of light to pycnidium 
formation in the Sphaeropsidales. Michigan Acad. Sci. 
Rep, 22: 209-213. Au 1921. 

Corrington, L. J. The Peltigeraceae. [The Ascomycetes of 
Ohio V]. Ohio Biol. Surv. 2: 354-362. pi. 14, 13. 28 My 

Coville, F, V. The influence of cold in stimulating the growth 
of plants. Ann. Rep. Smithsonian Inst. 1919: 281-291. 
pL j-27. 1 92 1. 

Dahlgreen, XJ. Phosphorescent animals and plants. Nat. Hist. 
22: 1-26. Ap 1922. 

Dallimore, W. The yellow pines of North America. KewBull. 
Misc. Inform. 1921: 330-335- 1921- 

Dalton, L. V. The plants and animals of Venezuela. In Dal ton 
L, v., Venezuela, 47-60. London. 1912. [Illust.] 

256 Index to American botanical literature 

Darlington, H. T. Contributions to the flora of Gogebic County, 
Michigan. Part I. Michigan Acad. Sci. Rep. 22: 147-176. 

Au 1921. 

Deam, C. C. Plants new to Indiana—IX. Proc. Indiana 
Acad. Sci. 1920: 225-228. 1921. 

Dow, C. M. Anthology and bibliography of Niagara Falls. 
Pp, I-1423. Albany. 1921. [Illust.] 

Chapter VI (pages 439-491) gives a chronologic discussion of the "flora 
and fauna." 

Dufrenoy, J, Anaerobic bacteria in plant tissues. Phyto- 
pathology 11: 344. Au 192 1. 

Dupret, H. Notes about the Drepanodadi of the vicinity of 
Montreal, Quebec, Canada. Bryologist 24: 36-39. 21 S 

Earle, F. S. S, M, Tracy as a botanist. Torreya 21: 64, 65. 
Au 192 1. 

East, E. M. A study of partial sterility in certain hybrids. 
Genetics 6: 311-365. /, i-iy. Jl 192 1. 

Studied crosses of Nicotiana, 

Emerson, R. A. Heritable characters of maize— IX. Crinkly 
leaf. Jour. Hered. 12: 267-270. /. 24-26. 23 S 1921. 

Enock, C. R. Natural History [of Peru]. In Enock, C. R.^ 
Peru 192-205. London. 1908. 

Espinosa Busies, M. R. Una enfermedad grave del trigo Chileno. 
Revista Chilena Hist. Nat 24: 79-86. pLj+f. 8. 31 Au 1920. 

Essary, S. H. Lespedeza (Japan clover). Univ. Tennessee 
Agr, Exp, Sta. Bull. 123: 1-28. Ap 1921. [Illust.] 

Evermann, B, W., & Clark, H. W. The flora of Lake Maxinkuckee 
and vicinity. In Evermann & Clark's Lake Maxinkuckee 
[Indiana], a physicial and biological survey 2: 117-447. 1920. 

Farwell, O. A, Notes on the Michigan flora — III. Michigan 
Acad. Sci. Rep. 22: 177-185. Au 1921. 

Includes new varieties and forms in various genera. 

Fink, B. The Lecideaceae. [The Ascomycetes of Ohio IV.] 
Ohio Biol. Surv. 2: 334^353- pl^ ^3- 28 My 1921. 

Includes new species in Lecidea (i), Bacidia {i),Buellia (i) , Rhizocar pan (i). 

Index to American botanical literature 2^^ 

Fink, B. Lincoln Ware Riddle, lichenist. Bryologist 24 : 
33-36. 21 S 1921. 

Fisher, G. C. A station for the ramshead lady 's-slippen Torreya 
21: 63, 64. Au 1921. 


Forbes, C. N. New Hawaiian plants— VII. Occ. Pap. B. P, 
Bishop Mus. 7: 33-39. pL 3-1 1. 1920. 

Includes 7 new species. 

Forbes, C. N. Notes on Marsilea villosa Kaulf. Occ. Pap. B. P. 
Bishop Mus. 7: 47-49. pi. 13^ 14, 1920. 

Forbes, C. N. Salient features of Hawaiian botany. Proc* 
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N., & Munro, G. C. A new Cyanea from Lanai, 

Hawaii. Occ. 

Cyanea Baldwinii, sp. nov. 


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Gassner, G. Ueber einen eigenartigen Uromyces auf Passijiora 
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Mr 1922. 

Includes Uromyces Appelianiis sp. nov. from Uruguay. 


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Je 1921. 

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Contains much false botanical matter, 

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258 Index to American botanical literature 

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120-123; 195-196; 255-256; 326-329; 374-377- 1921- 

Heimlich, L. F. Native plants of 


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Hoffmann, R, Flora of Berkshire County, Massachusetts. 
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Hotson, J« W* Sphignum used as a surgical aressing in Germany 
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F 1922. 

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Includes Trianiliellj gen. nov. and new combinations in many genera. 

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Vol. 49 

No. 9 





New species of trees of medical interest from Bolivia 

H. H, RusBY 

The botaincal work of the Mulford Biological Exploration of 
1921-22 was directed specially toward the investigation of 
medicinal plants. The distinctly pharmacological features of 
these plants will be discussed in the medical and pharmaceutical 
literature, but such unknown species as are encountered should 
be cited in botanical publications. Two problems of exceptional 
interest were connected with the botanical origin of the two 
drugs, *'coto*' and '*cocillana" or "guapl,"and their spurious 

It has never been known what trees yield the coto and paracoto 
barks, although evident that they pertain to the Lauraceae, 
For many years, no genuine bark of either has reached our 
markets, and the use of their worthless substitutes has resulted 
in destroying the medical reputation of these valuable drugs. 
Through the careful work of my associate, Dr. O. E. White, of 
the Brooklyn Botanic Garden, it is now possible to describe the 
genuine coto tree, as well as its principal substitute, and to provide 
for the supply of the genuine drug in future. The bark of a 
second substitute, with leaf-bearing branches, was also collected* 
but the species cannot be determined from this material, though 
undoubtedly pertaining to the genus Nectandra, 

Of coclllana or guapi bark, I collected not only the genuine* 
species, but also the only substitute that has as yet been seen In 
commerce, besides two other species which, for special reasons, 
are liable to so appear in future. 

The Bulletin for August (49: 223-258) was published August 31, 1922 



26o New species of trees from Bolivia 

Those interested further in these subjects should consult 
future numbers of the Journal of the American Pharmaceutical 


Nectandra coto Rusby, sp. nov. 

Young parts, inflorescence and lower leaf-surfaces densely 
and finely yellowish-tomentellate, the twigs stout. Mature 
leaves, including the petiole, 10-20 cm. long, 3-7 cm. wide, the 
petiole 2-3 cm. long, stout, sharply channelled above when dry, 
reddish browTi; blade thick and coriaceous, regularly oblanceo- 
late or slightly inequilateral, abruptly very short-pointed, the 
point acutish, the margin entire and sharply revolute, the upper 
surface glabrous and lustrous, drying brownish, the midrib 
reddish and channelled; lower surface pale, the midrib sharply 
prominent, the secondaries twelve to fourteen on each side, 
thin and crooked, lightly prominent, lightly anastomosing near 
the margin, the interspaces faintly reticulate. Inflorescence 
(very young in my specimen) axillary, the panicles small and 
loosely flowered, mostly long-peduncled, but shorter than their 
leaves even when in fruit, the peduncles and their branches 
coarsely and strongly angled. Bracts broadly ovate, thick, 
reddish, obtuse. Buds too young for dissection, appearing 
while the preceding mature fruit is still attached, tomentose. 
Calyx-limb wholly deciduous in fruit, the base almost concealing 
the fruit when young. Fruiting pedicel of variable length, 
thickened upward and becoming continuous with the calyx. 
Calyx-cup of mature fruit hard and thick, dark brown, roughly 
tuberculate, especially downward, broadly campanulate or 
subhemispherical, 2 cm. or more broad, the margin truncate, 
roughish, single but showing two distinct layers, the light brown 
fruit slightly projecting when mature, its summit lightly convex, 
finely granular or lightly mottled, bearing the very short, stout, 
acute style. 

Mulford Biological Exploration Nos. 1048 and 1050, Collect- 
ed by Dr. O. E. White as "Coto No. i " (No. 1048) and "CoLo 
No. la" (No. 1050), and under the native names **cotopiquiante'' 
and **coto fino." Mountains south of Huachi, near the Cocha- 

bamba River, Boli\ia, at an altitude of 2500 to 3000 feet, 
September 9, 1921, 

Dr. \Miite's field notes are as follows: 

No. 1048.— A tree 50 to 60 feet high, with few branches except near the 
top and these short in comparison with the height of the tree. Bark of a 
cinnamon-brown color, peeling with difficulty, aromatic and very pungent, a 
few seconds after being chewed, the odor pungent when fresh. Wood of a 
sandal-wood yellow and a satiny luster, the trunk turning brown after being 
peeled. Leaves whitish beneath, dark- to Hght-green and glossy aboAC. 
Young growth cinnamon-brown. Flower buds whitish. Cup of fruit brown- 

New species of trees from Bolivia 261 

green with brown spots, the fruit green, evidently of two generations on the 
same tree. 

It Is not possible to say with absolute certainty, from speci- 
mens collected in this stage of growth, whether this is a species 
of Nectandra or of Ocofea. The slender midribs and veins of the 
leaves are more those of Ocotea than of Nectandra, but the tomen- 


tose flower-buds are characteristic of Nectandra, the flowers of 
Ocotea being usually glabrous. There is no species of either 
Nectandra or Ocotea with which this can he identifioH 

Ocotea pseudo-aoto Rusby, sp. nov. 

Young growth and inflorescence whitish-puberulent, the 
lower surfaces of the mature leaves minutely and sparsely 
puberulent, the branchlets elongated and stout, little spreading, 
deep red, more or less angled. Leaves, including the petioles, 
6-12 cm. long, 1.5-3 cm. broad, the petioles 1-1.5 cm* long, 
deep red, the upper surface nearly plane. Blades thick, oblong, 
acuminate at base and with a very abrupt short and stout 
obtuse point at the summit, the margin entire, not revolute, the 
upper surface brown when dry, glabrous, slightly lustrous, the 
midrib slightly prominent, sharply so underneath, the six to 
eight pairs of secondaries slender, ascending, obscurely connecting 
near the margin. Panicles axillary, less than half the length of 
their leaves, sessile or subsessile, broad, rather dense as seen in 

the young state. Bracts thickish, red, broadly ovate. Perigone 
thin, glabrous, undeveloped in the specimen. The other floral 
parts are too young for characterization. 

Collected by Dr. O. E. White in the mountains south of 
Huachi, near the Cochabamba River, Bolivia, at an altitude of 
2500 to 300 feet, September 9, 1921. Mulford Biological 
Exploration No. 1051, collected as **Coto No. 2" and under the 
name "coto ordinario.'* It is to be noted that the term "ordin- 
ario," as applied by these natives to any substance or article, 
has the significance that the article is of inferior quality. 

Dr. White contributes the following field notes: 

Very large forest tree, growing with Nos. 1048 and 1050 and of the same 
general habit. Young buds only, and no fruit encountered. Bark thick and 
easily peeled, not specially aromatic and not pungent. The peeled wood 
remains white. Wood reddish-white internally. 

The same difficulty of identification presents itself with this 
species as with No. 1050. The buds are a little more developed 
and the perigone characters are found to correspond with those 
of Ocotea. 

262 New species of trees from Bolivia 


occurred in commerce in this country under both names, coto 
and paracoto, but sold under either name it is absolutely 
spurious and medicinally worthless, so far as known. 


Aerodiclidium benense Rusby, sp. nov. 

Specimen in mature fruiting stage. Glabrous, the twigs 
numerous, rather slender, terete, densely leafy at the summit. 


Petioles 5-10 mm. long, slender, sharply channelled above. 
Blades thick and coriaceous, lustrous above, lanceolate, abruptly 
acuminate at base and summit, with acute apex, the midrib 
and slender venation scarcely prominent above, sharply so 
beneath. Principal secondaries five or six on each side, with 
short intermediate ones, strongly upcurved and connecting near 
the margin, the reticulation fine and sharp. Margin entire, very 
slightly revolute. Panicles axillary, less than half the length of 
their leaves, stoutly peduncled, having one to four shortly and 
stoutly pedicelled fruits, the pedicels strongly thickened upward. 
Calyx-cup subhemispherical, about 1.5 cm. broad, hard and thick, 
heavily wrinkled, its truncate summit with thin, sharp margin 
projecting slightly beyond the strongly recurved limb so as to 
form a double mouth, the limb thin, irregularly crenate-dentate. 
Fruit about 2 cm. long and two-thirds as broad, oval to ovoid, 
light brown, glabrous, strongly wrinkled at the base and very 
shortly stipitate, the rounded summit topped by the very short 
stout style. 

Collected by Dr. O. E. White at the second portage of the 
Bopi River, Bolivia, at an altitude of about 2000 feet, August 
8, 192 1. Mulford Biological Exploration No. 649. 

Dr. White says: '*A small evergreen tree 20 to 30 feet high, 
trunk 6 Inches in diameter, growing in damp, partly shady 
places near the river bank. The fruit is green with white specks, 
the cup brilliant scarlet-red. Not common." 

Guarea Bangii Rusby, sp. nov. 

Only fruiting specimens seen. Glabrous, with the exception 
of the tomentose fruit, pedicels and peduncles, and a very slight 
short, sparse pubescence on the leaf rachis, petlolules and lower 
surface of the midribs. Rachis and petiole subterete, slender, 
together 7-20 cm. long, the petiole var^ang from one-eighth (in 
the larger leaves) to one-half (in the smaller ones) of the total 
length. Leaflets three to five pairs, opposite, the lower success- 
ively much smaller, 7-20 cm. long, including the petiolule, 
which is one- tenth to one-fifth of the total, 3-7 cm. wide, thin, 
pale green, oblanccolate, acute at the base, the summit very 

New species of trees from Bolivia 263 

abruptly contracted into a short and broad, mostly obtuse 
acumination, the margin entire and not revolute. Midrib 
usually slightly furrowed on its upper surface toward the base, 
underneath prominent and slender, like the seven to twelve 
pairs of mostly opposite secondaries, which are strongly up- 
curved near the margin, where they are obscurely confluent. 
Fruit-panicles much shorter than their leaves, sessile or sub- 
sessile, rather densely fruited, the rachls and pedicels very stout, 
the latter little longer than broad, annular-roughened, articulated 
with the very short stipe of the fruit, which is not so thick as its 
pedicel. Mature fruit usually a little more than 2 cm, long and 
nearly or quite as broad, pear-shaped when young, becoming 
nearly spherical, except for the narrow base, which remains 
unchanged to form a short thecaphore. Body of fruit obscurely 
costate or striate, tubercular-roughened and softly and finely 

First collected by Mr. Miguel Bang, on the hills near Tipuani. 
Collected also by H. H. Rusby in October, 1921, on the Mulford 
Biological Exploration, at Rurrenabaque, Bolivia, Nos. 779 and 
1590; also No. 2178 (**Guapi B'*). All the spurious cocillana 
bark that has reached our market up to the present has been 
derived from this species. 

A good-sized forest tree of the Mapiri and Beni River region, 
growing mostly on the hillsides in relatively dry soil, the bark 
roughly striate and shreddy, and peeling in long, fibrous, thin 

Guarea alborosea Rusby, sp. nov. 

Glabrous, the petiolule and lower surface of midrib finely 
verrucose or papillose in the dried state. Petiole and rachis 
rather stout, deep purple, together 2-4 dm. long, the petiole 
about one-fifth of the total length, flat on the upper surface, the 
leaflets five to ten pairs, nearly opposite, the lower somewhat 
smaller. Petiolules 4-8 mm. long, stout, lightly channeled above, 
like the midrib. Blades 10-20 cm. long, by 4-8 cm. wide, thick- 
ish, pale underneath, oval or slightly narrower either below or 
above the middle, the base rounded or obtuse, the summit very 
abruptly contracted into a short, narrow, acute point, the 
margin entire and not revolute, the venation not at all prominent 
above, sharply so underneath, the secondaries ten to fifteen on a 
side, not opposite, straight and slender, moderately spreading, 
falcate toward the margin, connected by the coarsely reticulate 
tertiaries. Inflorescence axillary, racemose, simple, or occasion- 
ally slightly branched at the base, the racemes recurved-pendu- 
lous, often 2-3 dm. long, 1.5-2 cm. broad, densely flowered, the 
rachis slender, strongly nodose by the flower-scars, the bractlets 
minute, the flowers divaricate. Pedicels slender, articulated 

264 New species of trees from Bolivia 

Into the broad concave base, the pedicel proper scarcely one- 
fifth the length of the flower, obscurely angled. Unopened 
cal>^ not seen. Mature flowers about 6 nun. long; the erect 
unexpanded corolla 2.5-3 i^im. broad. Calyx crateriform, 3 
mm. broad, closely investing the base of the corolla, deeply 
four-lobed, the lobes thick, purple, broadly ovate and obtuse. 
Petals 6 mm. long, 2 mm. broad, oblong, obtuse, slightly thick- 
ened at the purple or rose colored summit, the basal portion white. 
Stamen-tube three-fourths the length of the petals, broadly 
sulcate, slightly contracted below^ the mouth, which Is not 
quite so broad as the lower half, the margin shallowly eight- 
crenate, the small anthers equalling its lobes. Ovary and style 
appressed-pilose, the ovoid ovary about equalling the stout 
style in length, the latter reaching the summit of the tube. 
Immature fruit fig-shaped, abruptly contracted at the base, then 
tapering into the stout short pedicel. Mature fruit not seen. 

A small tree in the forest at Rurrenabaque, Bolivia, at an 
altitude of about 1000 feet. Collected by H. H. Rusby in 
flower and fruit, on the Mulford Biological Exploration, October 
6, 1921 (No. 797). 

The species is peculiar In Its very long slender raceme and in 


the handsome rose and white coloration of Its flowers. 

Notes on trees and shrubs of the southeastern United States 

W. W. Ashe 
I. The Castanea pumila group 

Castanea pumila Margaretta var. nov. 

A tree, sometimes attaining (in Arkansas) a height of 15 m. 
and a diameter of i m., but usually smaller and often a large 
single-stemmed shrub. Leaves sinuate-serrate with ten to 
twenty pairs of lateral veins; sun leaves, especially those on 
flowering twigs, oblong or lance-oblong, 8-15 cm. long, 3-5 cm. 
broad, acute and somewhat taper-pointed at each end, thick 
and firm, dark green above, close white-pubescent beneath exxept 
on the midrib and veins; shade leaves 5-12 cm, long, 3-6 cm. 
broad, obov^ate or sometimes oblong-obovate, obtuse, or rounded 
and abruptly pointed at the apex, cuneate at the base, thin, 
pale green on both sides, but more or less pubescent beneath. 
Stammate aments continuous, 8-12 cm. long, at an thesis 6-8 
mm. thick; calyx lobes 3-4 mm. long, obtuse; involucre of fruit 
bur-like, 2-4 cm. thick, canescent, comparatively thinly beset 
with branched rigid canescent spines, 3-6 mm. long, which as a 
rule are not sufficiently dense to conceal completely the involucre. 
Nut short-ovate, 11-15 mm. long, 7-12 mm. thick, pointed 
at the tip, shining, blackish brown. 

Dry soils or, at the upper edge of the longleaf pine lands of 
Texas and Louisiana, along the edges of sandy hummocks. 
The following collections have been made by the writer: Ala- 
bama, Winston County ; Mississippi, Jackson ; LouisiAxN^a, 
Natchitoches and Sabine (Type) Parishes ; Texas, Sabine County ; 
Oklahoma, McCurtaIn County; and Arkansas, common on the 

mountains. Named for Marearet Henrv Newland. 

pical C pumila (L,) M 

west and southwest of Tennessee and is separated from it by hav- 
ing the much shorter and smaller shade leaves of an obovate 
outline, thinner and often green beneath ;by the closer white pube- 
scence on the narrow and acute sun leaves (never velvety and 
rarely dull grayish); by the spines of the involucre being 
canescent well toward their tips (those of the typical C pumila 
being nearly glabrous when mature) and seldom so dcase as to 
conceal the involucre. Nut apparently not quite so plump as 

that of the type. 


266 Notes on treks and shrubs 

Castanea alnifolia Nutt. 

The western recorded limit of this dwarf shrub with its sub- 
terraneous stems is Louisiana.* It was collected by the writer 
in Newton County, Texas, during October, 1 921, under the 
same conditions as it grows further east and not exceeding 18 
inches in height. C. alnifolia puhescens Nutt. has all of the leaves 
pubescent beneath. 

^ Castanea floridana (Sarg,) comb nov. 

Castanea alnifolia floridana Sarg. Bot. Gaz. 67: 242. 1919. 

I have not seen authentic specimens of the tree which Dr. 
Sargent has recently described and which he separates from the 
typical C. alnifolia by its arborescent form, but it is believed 
that material collected near Charleston, South Carolina, in 1915 
represents it; and, if so, the differences of smaller fruit (which 
is more densely hispid), somewhat larger leaves with about two 
more pairs of lateral veins, and entirely different habit of growth 
justify specific rank. 

X Castanea NEGLECTA Dode, Bull. Soc. Dendr. France 8: 155. 

May, 1908. C pumila x C, denlata 

According to the description this hybrid is in general like C. 
ptimila. It differs, however, in having larger and less densely 
pubescent leaves, a larger involucre with irregular spines, longer 
aments, and a single larger nut. The distribution is given as 
extending from Maryland to North Carolina. As a rule hybrids 
between C. pumila and C. dentata have the nuts in pairs, although 
this is not necessarily the case. 

M. Dode has been kind enough to furnish the writer with a 
list of the specimens at the Museum of Paris, which he refers to C 
neglecta. Of the seven specimens listed, the locations of five are 
well within the ranges of both C pumila and C. dentala. These 
are: Small & Heller 152, and their collection of June 22, 1891, 
both from Blowing Rock, North Carolina; Pearson, Fairfax 
County, Virginia, 1844; Green, Maryland, 1839; BonpJand, 
Washington, D. C. Since the states only are given for the other 
two {Gihhs, South Carolina, and Vasey, Alabama, 1878), it is 
not possible to determine this. 

* Small, Flora S. E. U. S. Ed. 2, 348. 

Notes on trees and shrubs 267 


County, Georgia, and Yancey County, North Carolina, largely 
agrees with Dode's description of C. neglecta. In its gross 
characters it shows, as Dode states, a superficial resemblance 
to C dentata, having large pointed leaves, not very densely 
pubescent, and a large involucre with long, irregular, nearly 
glabrous and closely crowded spines; but there are two nuts to 
the involucre. Other material collected in flower in Sullivan 
County, Tennessee, shows similar leaf character. 

-^Castanea Ashei Sudworth, comb. nov. 
Castanea ptimila Ashei Sudworth, Am. For. 28: 300. 1922, 

This is the common chinquepin of the coastal plain of the 
Southern States. Its distribution begins in northeastern North 
Carolina and extends to Louisiana (and probably to south- 
eastern Texas). It is separated from C pumila by its smaller 
foliage, shorter and more slender aments, canescent spines on 
the involucre and prevailingly thicker nuts. Walter (FL Car. 

233- 1788) notes two forms (without description), praecox and 
scrotina. C. Ashei is possibly his forma praecox. 

The type of C. pumila (L.) Mill. {Fagus pumila L.) came from 
eastern Virginia (Gronovius, Flora Virginica 118. 1739), within 
which region the only known form is that which is here regarded 
as typical C. pumila. 

Key to Castanea pumila group (one nut to the involucre) 

Al = Alleghanian; C = Carolinian; A^ Austral. 
Spines of mature fruit densely set, glabrous or 

Leaves soft-pubescent beneath C, pumila (AI & C). 

Leaves 10-20 cm. long, more or less glabrate. . X C neglecta (Al). 

Spines of mature fruit not densely set, canescent or 

tips merely glabrate. 
Leaves 10-15 cm. long; 12-20 pairs of veins. 

All leaves closely white-pubescent be- 

■ neath C. Ashei {h). 

Interior leaves green beneath or glab- 
rate C pumila Margarella (C). 

Leaves 6-10 cm. long; 8-16 pairs of veins. 

Only uppermost leaves on twigs white- 
pubescent beneath. 
10-16 pairs of veins, non-stoloniferous. C, floridana (A). 
8-12 pairs of veins, a dwarf stoloniferous 

shrub C. alnijolia (A). 

All leaves white pubescent beneath C alnijolia ptcbescens (A) , 

268 Notes on trees and shrubs 

2. Miscellaneous species 


X Q 

lercus coloradensis hybr. nov. Q. virginiana Mill. X Q. 

macrocarpa Mx. 
Leaves heterophyllous, entire and oblong or undulate, sinuate 
or lobed, especially near the apex, with short abruptly acute 
or obtuse lobes, 4.5-10 cm. long, 2-4 cm. wide, thick and sub- 
coriaceous, dark green above, strongly reticulated and densely 
close white-pubescent beneath, revolute on the margins; twigs 
3-4 mm. thick, permanently close pale gray pubescent. Fruit 
2.5-3 mm. long, excluding peduncle (which is often of half the 
length of the fruit) ; cup covering one-half or more of the oblong 
nut, obconic or slightly rounded, covered with slender acute, 
closely imbricate scales, either thin at the top or much thickened 
by several overlapping rows of scales and incurved at the edge. 

Along the Colorado River above Marble Falls, Texas. While 
the overcup oak is rare in this region, the cup-character points 

Q. acuminata Houba or Q 


Quercus nigra elongata comb. nov. 

Querctis aquatica elongata Ait. Hort. Kew. ed. i, 3: 357. 1789. 

In the third edition of Alton's work there is a reference to a 
plate (Abbott, Insects Ga. 2: pL 2g.)y which figures the leaf- 
form described by Sargent under his Q. nigra tridentifera (Bot. 
Gaz. 65: 429. 1918). 


^Malus elongata pubens var. nov. 

In general similar to the type but having somewhat broader 
leaves, perntanently soft-pubescent beneath 

Head of Chattoga River, Rabun County, Georgia, October, 

1913, W. W, A. 

Malus platycarpa parvula var. nov. 

Foliage in general similar to that of type, but the leaves more 
ovate in outline and smaller; fruit of the same shape but much 



Commelinantia, a new genus cf the Commelinaceae 

B. C. Tharf 

(with plates io and ii) 

Few if any families of plants have members in more urgent 
need of field study than are those of the family Commelinaceae. 
Their evanescent corollas, which it is difficult even to bring 
from field to laboratory without injury; the delicate shades of 
color which are lost in pressing; and their succulent stems, 
which shrink tremendously in drying, make field study imperative 
if one is to get any adequate idea of these delicate details as they 
appear In the growing plants. Since 1919 I have had abundant 
opportunity for field study of a very interesting member of the 
family, a species first described by Torrey* as Tradescanlia 
anomala and latter transferred by C. B. Clarkef to the genus 
Tinantiaj where it has remained to the present time. 

Torrey *s description, based upon specimens from Texas, (one 
of Wright's, and one of Thurber's), preserved in the herbarium 
of the New York Botanical Garden, is surprisingly good, con- 
sidering that it was drawn wholly from dried material. Such 
details of general habit, sepals, petals (except color), filaments 
(except that of the posterior stamen), anthers, style, stigma, 
and ovules, as are pointed out, are correctly described. The 
greatest error into which his material led him seems to have 
been in considering the ovary and capsule two-celled. He also 
described the seeds as being '*as in the rest of the genus,'* which 
is incorrect. The wonder is that, having examined his material 
so carefully as to be able to write such an accurate description, 
he could feel justified in assigning this species to the genus 
Tradescanlia. That he was none too confident of its having been 
correctly assigned is abundantly testified by the specific name, 
by legendsf on the sheets containing his specimens, and by his 

*U. S. and Mex. Bound. Bot. 225. 1859. 

fDeCandolle, Monog. Phaner. 3: 287, 1881, 

t Wright's specimen is labelled "Wright, Texas, Commelyna anomala, Torr 
Tradescanlia anomala Torn," the word ''Commelyna'' being stricken out 
Thurber's specimen is labelled "Thurber, San Antonio Texas, May 1853 
Commelyna rhodantha, Torr." 



observation that "the species Is intermediate between Trades- 
cantia and Commelyna.^^ 

Clarke's description of Tinantia anomala appears to be based 
neither upon the excellent original by Torrey, nor upon any 
critical examination of material of the species. He characterizes 
the plant as glabrous, the leaves as ovate-cordate, the three 
longer stamens simply as '*naked above/' the three shorter as 
having "golden bearded antJiers," the stems as "dichotomously 
branched," the inflorescence as being "almost included in the 
subcomplicate uppermost leaf, "the petals as being "obovate- 
elliptic, bluish" (by inference all alike in size and appearance). 


With the exception of the capsule, the measurements of all 
structures are under-stated. His specimens may not have shown 
the basal linear-lanceolate, ciliate, leaves; his scant measure- 
ments may have been due to his materials having shrunken in 



drying, and his representation of the "uppermost leaf as 
subcomplicate" may have been due to his specimens having 
been somewhat wilted before being pressed ; but there seems no 
valid excuse for so inaccurate a description of stamens, or for 
characterizing the stems as dichotomously branched, or for 
totally ignoring the description (excellent so far as it goes) of 
Torrey. His observation, concerning the relationship existing 
between the type species (T.'ftigax) and the other species, that 
caeterae species . . . cum T. fugace . . . militant; 
et ad Tinantiam ex necessitate referendae sunt, nisi genus Tinantia 
ad Tradescantiam reducatur," seems to be in the nature of an 
apologetic attempt at justification, though it is difficult to see 
the logic in his statement. 

My own experience with an abundance of fresh material of 
the species in question constrains me to call attention to the 
following characteristics, which keep it from being properly 
placed in the genus Tinantia as characterized by Clarke : branches 
which break through the leaf sheaths; flowers in a simple ter- 
minal scorpioid raceme subtended by a single^ broad ^ erect, flat, 
cordate, clasping spathe; colored lateral petals much larger than 
the very small anterior white petal; stamens of four distinct 
kinds as regards size and pubescence. 

The character of the stamens and of the spathe and the habit 
of the branches breaking through the leaf sheaths are sufficient 
to exclude it from either Commelina or Tradescantia. The 
former has difformed stamens, all of whose filaments are entirely 


glabrous and some of whose anthers are sterile; a spathe which is 
folded so as to protect alike the flowers bud and the ripening 
capsule, the flower being projected above the edges of the spathe 
only during the very brief period of bloom : the latter has stamens 
all alike in size and pubescence and all fertile; the inflorescence 
subtended by one or more (usually two) unlike bracts, never by 
a spathe. Neither Commelina nor Tradescantia has branches 
which break through the leaf sheaths. This habit, howcv^er, is 


held in common with certain tropical genera to which the species 
has never been heretofore compared. Dr. Pennell has kindly 
gone over the whole family as represented by specimens in the 
New York Botanical Garden Herbarium and reports the following 
genera as having this habit: 

1. Callisia (tropical American) lacks spathe, has uniform petals and 

2. Campelia (tropical American) has spathe, but uniform petals and 
stamens, white. 


3. Cyanoiis (tropical Asia) has series of overlapping spathes. 

4. Dichorisandra (tropical American) lacks spathe, inflorescence a panicle. 

5. Floscopa (tropical American and Asia) lacks spathe, inflorescence a 

6. Forrestia (tropical Asia) lacks spathe, inflorescence congested erupts 

7. Pollia (tropical Asia) lacks spathe, inflorescence a panicle. 

8. Sauvellea (Cuba) has spathe, but petals and stamens uniform; repent. 

9. Sireptolirion (China) has spathe below cymes or panicle, but petals 
and stamens uniform, the filaments bearded. Stem twining. 

10, Tradescantella (Florida) lacks spathe, but petals and stamens uniform. 

II- Zehrina Ctropical American) has two bracts below the congested in- 
florescence; petals uniform, at base approximating to form a corolla-tube. 

The species diflfers from any of these generic groups in its 
combination of erect habit, single, flat, erect, spathe; unequal 
petals; and stamens of four forms. 

In addition to the several genera listed above, Dr. Pennell 
finds that Tradescantia Pringlei S. Wats.* also has leaves 
which break through the sheathes. Moreover, it has the following 
additional characters which ally it ver>^ closely to T. automata: 
general appearance ver>^ similar; inflorescence a simple scorpoid 
raceme (or frequently reduced to only one flower); stamens 
probably like those of 2". ajiomala in that there are four kinds, 

*Trans. Am. Acad. 26: 157. 1891, 


implied by the following statement: "stamens 6, unequal, the 
longer with filiform filaments bearded in the middle (or one 
naked) and broadly oblong anthers, the shorter filaments densely 
bearded in the middle with green gland-tipped hairs, the anthers 
orbicular; anther cells contiguous." The stamen corresponding 
to the posterior stamen in 7". anomala might, by casual examina- 
tion, have been counted as being like the postero-lateral ones. 

But r. Pringlei has the following characters In conflict with 
T, anomala: plant green, not glaucous; cauHne leaves uni- 
formly ovate, abruptly acuminate, none modified to form a 
definite *'spathe'' (though the slender peduncles of T. Pringlei 
frequently have one or more much reduced sheathes below the 
inflorescence), infloresc'ence axillary, petals uniform, all blue 
("purple" ?). Though T. Pringlei thus shows such strong 
resemblance to T, anomala in certain characteristic^, yet 
because of these important points of dissimilarity and because 
of insufficient material for critical study I feel constrained to be 
content with pointing to it as perhaps the closest existing relative 
of T, anomala, leaving the question of its assignment to the 
proper genus to be settled by a more critical study of more 
abundant material. Certainly it can not properly belong in 
the genus Tradescantial 

Confining my attention, therefore, in the present paper lo 
the placing of T, anomala, it seems not only justifiable but 
necessary to make it the type of a new genus.* Following the 
suggestion of Torrey that the species has certain characteristics 
which ally it to Tradescantia and Commelina and recognizing 
the fact that it was assigned to Tinantia more than forty years 
ago, I propose a sort of hybridization of these names, giving the 
new genus the name Commelinantia, The generic description 
which follows Is made broad enough to include some of the dis- 
tinctive characters of T. Pringlei, acting on the probability that 
this species will ultimately be assigned to the new genus. 


Annual succulent plants, somewhat resembling species of 
Commelina: stems 2-8 dm. long, erect, ascending, or decumbent, 

•Since 1 becjan the study of this sjx^cies with the view of determining the 
propriety of retaining it in Tinantia, I have learned that Dr. J. K. Small had 
decided to segregate it as a distinct genus in the next edition of his Flora of 
the Southeastern United States. 


glabrous, much branched at maturity, the branches ascending, 
breaking through the close-fitting leaf -sheaths: basal leaves 0.5- 
3.5 dm. long, acute or acuminate, smooth or slightly pubescent 
with a few long hairs, appearing in early winter after the fall 
rain; lower cauline leaves resembling the basal leaves, 0.5-2.5 dm. 
long, the petiole arising from the closed sheath; upper cauline 
haves generally shorter and broader than the lower; inflorescence 
a simple scorpoid, peduncled raceme, terminating branches and 
stems (or sometimes axillary and reduced to a single flower); 
flowers one or more (usually), each subtended by a small ovate 
bract; calyx of three similar, green, ovate, imbricated, persistent 
sepals; corolla showy, the petals equal or very unequal; stamens 
six, all fertile but dissimilar in size and in degree of pubescence; 
ovary oblong, three-celled, with two superimposed ovules in 
each cell, the stigma capitate; capsule loculicidally three-valved, 
the seeds greyish brown with copious endosperm surrounding 
the transverse embryo. 

Commelinantia anomala (Torr.) comb. nov. 

Tradescantia anomala Torr. U. S. & Mex. Bound. Bot. 


Tinantia anomala C, B. Clarke; De Candolle, Monog, Phaner. 
3: 287. 1881. 

Pea green, usually glaucescent throughout: stems clustered, 
at first simple but later much branched, more than one branch 
frequently arising at a single node (especially in the upper part 
of a stem late in the season); basal leaves 2-3.5 dm. long, linear- 
spatulate, attenuate into long ciliate petioles below, rather pale 
green, glaucescent, slightly pubescent with a few long hairs 
above; lower cauline leaves 1.5-2.5 dm. long; upper cauline 
leaves sessile at the top of the sheaths or short-petioled, 0.5-2 
dm. long, broadly or narrowly lanceolate or somewhat cordate 
and clasping at the base, acute or acuminate; inflorescence 
strictly terminal on stems and branches, subtended by a spathe 
strongly resembling the upper cauline leaves except in being 
erect, shorter (4-8 cm. long), broader, and strongly cordate- 
clasping at the base and in having no sheath ; flowers three to 
sixteen or more, the bracts persistent, glabrous and green, each 
usually 3-5 mm. long; sepals keeled, slightly hooded, blunt, 
9-12 mm. long, glandular-ciliate above the middle in the bud, 
a few apical cilia being persistent; corolla of two postero-lateral 
very showy lavender blue petals and a very small and in con- 
spicuous anterior white petal, the showy petals almost rhomboid, 
1. 5-1. 8 cm. long and somewhat broader than long, the small 
petal ovate-rhomboid, 3-4 mm. long, and longer than broad; 
stamens with filaments, of four distinct forms — (a) the posterior 
stameu densely bearded (except at the extreme apex, at the 


extreme base and in the lower two-thirds of the inner face) with 
showy lavender-purple hairs, (b) the two postero-lateral densely 
bearded above the middle with a collar of short lavender-purple 
yellow-tipped hairs, (c) the two antero-lateral bearded below 
the middle on the posterior margin with a tuft of short lavender 
hairs, and {d) the anterior stamen glabrous throughout— anthers 
of the three anterior stamens oblong, these stamens being mani- 
festly larger throughout than the posterior stamens, the latter 
with orbicular anthers; ovary constricted in the middle, the 
stigma terminating a slender bent white style; capsule oblong, 
6-8 mm, long; seeds 3-3.5 x 2-2.25 mm., rounded at one end, 
truncate at the other, with a curved longitudinal grooved hilum 
on the inner face, near the center of which lies an apparent 
eccentric perforation containing the minute embryo; endosperm 
not continuous but separated by an obscure commissure on one 
side and bearing a group of low broad, tuberculate ridges radi- 
ating from the sides of the embryo cavity, where it comes through 
to the outer surface. 

In rich, moist, shaded soil of ravines and rocky hillsides of 

the Edwards Plateau, Texas; February to October, but princi- 
pally March to May. Original localities, according to Torrey: 
^' shady woods on the Blanco, Comale, and other rivers, 
Texas; Wright, (No. 699.) San Antonio, Texas; Thtirber,'" 

A comparison of Torrey 's description with that submitted 
herewith will rev^eal several places wherein certain differences 
appear. These differences are: (i) in the height of the plant; 
(2) length of basal leaves; (3) number of flowers per raceme; 
(4) I can find no evidence of wings on the margin of the posterior 
filament and am at a loss to account for Torrey 's mention of such 
structures unless he examined the filament in question only 
from the flat naked inner face of pressed material ; (5) the ovary 
is uniformly three-celled with two superimposed ovules in each 
cavity, though failure of fertilization will of course cause abortion 
of any one or more of the six, with a consequent great variation 
in the appearance of the capsule; (6) the seeds, though super- 
ficially resembling those of Tradescantia, are different from any 
Tradescantia seeds I have examined in that the cavity containing 
the embryo extends entirely through the seed, the endosperm 
instead of being continuous around the embryo is separated by 
a commissure that extends to the embryo-cavity on one side, 
and the seeds of C. anomala are of greater size. Other points 
which are brought out in the description submitted herewith 
are not mentioned in that of Torrey. 

Bull. Torrev Club 

Volume 49, plate 10 


Bull. Torrey Club 

Volume 49, plate ii 






I am indebted to Dr. F. W. Pennell, now of the Philadelphia 
Academy of Sciences, ■ for helpful criticism and advice, for 
assistance in looking up the literature, and for access to a speci- 
men of Tinantia Pringlei. 

University of Texas. 

Explanation of plates 10 and 11 

Plate 10 

Plant of Commelinanfia anomala (Torr.) Tharp, showing typical appearance 
of a flourishing specimen in bloom. The specimen is somewhat wilted causing 
the spathe to fold slightly. The linear-spatulate basal leaves, characteristic 
of plants during winter months, have disappeared by the tune the stems 
have reached this stage of maturity. Scorpoid racemes seen in profile. Scale 
in inches. 

Plate ti 

Fig. I. Inflorescence of C. anomala, showing the position assumed by each 
flower as it comes into bloom. The spathe is somewhat wilted, causing it 
to roll slightly at the edges. Scale metric. 

Fig. 2. Dissected flower to show relative size and general appearance of parts. 

Fig. 3. Stamens from a bud several hours before anthesis; magnified eight 
diameters. The filaments have not attained nearly the length of those in an 
open flower, but the anthers are beginning to dehisce. Relative size and 
shape of mature anthers, and also the nature of the pubescence of filaments 
well shown. 



The aim of this Index is to include all current botanical literature written by 
Americans, published in America, or based upon American material; the word 
America being used in the broadest sense. 

Reviews, and papers that relate exclusively to foresty, agriculture, horti- 
culture, manufactured products of vegetable origin, or laboratory methods 
are not included, and no attempt is made to index the literature of bacteriology. 
An occasional exception is made in favor of some paper appearing in an Ameri- 


can periodical which is devoted wholly to botany. Reprints are not mentioned 
unless they differ from the original in some important particular. If users 
of the Index w'ill call the attention of the e3itor to errors or omissions, their 
kindness will be appreciated. 

This Index is reprinted monthly on cards, and furnished in this form to 
subscribers at the rate of three cents for each card. Selections of cards are 
not permitted; each subscriber must take all cards published during the term 
of his subscription. Correspondence relating to the card issue should he 
addressed to the Treasurer of the Torrey Botanical Glub. 

Johnson, A. G. .Wheat scab and its control. U. S. Dept. Agr, 
Farmers* Bull. 1224: 1-16, /. 1-12. O 1921. 

Juday, C. The plankton. In Evermann, B. W., & Clark, H. W. 
Lake Maxinkuckee, a i:)hysical and biological survey 2: 
105-116. 1920. 

Kaiser, G. B. Little journeys into mossland, IV. Bryologist 
24: 41-43. 21 S 1921. 

Kasai, M. On the morphology and some cultural results of 
Fusaritim Solani Mast. Appel et Wollenweber, an organism 
which causes dry rot in the Irish potato tubers. Ber. Ohara 

Instit. Landw. Forsch. 1: 510-542. pL 8-1 1. 1920. 

Kaufifman, C. H. A black rot of squash. Michigan Acad. 
Sci. Rep. 22: 2or, 202* Au 192 1. 

Kauffman, C- H. Collyhia slricHpes, developed in the laboratory. 
Michigan Acad. Sci. Rep. 22: 203, 204. Au 1921, 

Kauffman, C. H. Mortierella Baineri. Michigan Acad. Sci. 
Rep. 22: 195-198. pL 16. Au 1921. 

Kurtz, F, Atlas de lasplantes fosiles de la Republica Argentina. 
Actas Acad. Nac. Ci. 7: I33"i53- i9-i- 


278 Index to American botanical literature 

Laibach, F. Untersuchungen uber einige Septoria Arten und 
ihre Fahigkeit zur Bildung hoherer Fruchtformen III und IV. 
Zeitschr. Plfanzenkr. 31: 161-194. 1921. 

Lee, H, A. Citrus-canker control; a progress report of experi- 
ments. Philippine Jour. Sci. 19: 129-175. pL 1-2. Au 1921. 

Lloyd, J. U. Plant pharmacy. Am. Jour. Pharm. 94: 238-244. 
Ap 1922. 

Lowe, E. N, Plants of Mississippi. Mississippi Geol. Surv. 
Bull. 17: 1-293. F 1921. 

Ludwig, C. A. A curious variation in the common milkweed. 
Proc. Indiana Acad. Sci. 1920: 243-245. /. 1-6, 1921. 

Lyon, T. L., & Wilson, J. K. Liberation of organic matter by 
roots of growing plants. Cornell Univ. Agr. Exp. Sta. Mem. 
40: 1-44^/- /-p. Jl 1921. 

McLean, F- T. The permeability of Citrus leaves to w^ater. 
Philipp. Jour. Sci. 19: 115-124. /. j. Jl 1921. 

Marchionatto, J. B. Acacia visco Lorentz, Syn. A, platensis 
Manganaro. Revisto Facult, Agron. Univ. Nac. La Plata 
142: 90-95, 1921. 

Marcou, P. Cacao, cacahult ou cacaouete. Jour. Soc. Ameri- 

canistes Pasia II. 12: 65-67. 1920, 

Maxon, W. R. Notes on American ferns— XVIL Am. Fern 
Jour. 11: 33-39. 1921. 

Includes Selaginella scopulosum, sp. nov., from Montana. 

Molisch, H. Der Naturmensch als Entdecker auf botanischem 
Gebiete. Schr.Ver. Verbr. Naturw. Kenntn.Wien 54: 97-123. 
/. J, 2. 1914. 

Muir, F, The origin of the Hawaiian flora and fauna. Proc. 
Pan-Pacific Sci. Confer. 1: 143-146. 1921. 

Murrill, W. A. John Macoun. Mycologia 13: 264. 80 1921. 

Murrill, W. A. Edward T. Harper. Mycologia 13: 264, 265, 
80 1921. 

Nestler, A* Einige Beobachtungen an der Paprikafrucht. Ber. 
Deutsch. Bot, Ges. 39: 230-234. Je 192 1. 

Index to American botanical literature 279 

Newcombe, F. C. Response of sensitive stigmas to unusual 
stimuli. Michigan Acad. Scl, Rep. 22: 145, 146. Au 1921. 

Patouillard, N. Une nouvelle lepiote du Bresil (Lepiota Putte' 
maneii). Bull. Trimest. Soc. Mycol. 37; 81-83. 20 Jl 1921. 

Pemberton, C. C. Some notes of the growth of A rbiitus Menziesii 
Pursh. Canad. Field-Nat. 36: 21-26. /. 1-4, F 1922. 

Petry, E. J. Germination and growth of Ceanothus americanus 
as aflFected by heated soils. Michigan Acad. Sci. Rep. 22: 
135-143- pl' ^3, 14' Au 1921. 


Pfeiffer, H. Conspectus Cyperacearum in America meridional! 
nascentium. I. Genus Heleocharis R. Br. Herbarium 1921: 
65-69. 1 92 1. 


Praeger, R, L. An account of the genus Sednm as found in 
cultivation. Jour. Roy. Hort. Soc. 46: 1-3 14. /. 1-8$. 6 Je 

Reinking, O. A. Higher Basidiomycetes from the Philippines 
and their hosts, V. Philipp. Jour. Sci. 69: 91--114. Jl 1921. 

Renner, O. Heterogamie im weiblichen Geschlecht und Em- 

bryosackentwicklung bei dem Onotheren. Zeitschr. Bot, 
13: 609-621, /. 7-5. 1 92 1. 

Rhoads, A. S. Some new or little known hosts for wood-des- 
troying fungi. HI. Phytopathology 11: 319-326. Au 1921. 



Ridlon, H. C. A new Polypodium from Vermont. Am, Fern 
Jour. 11: 46-48. pL /. 1921. 

Polypodium vulgare f. rotundaitim. 

Rodway, J. ' Fauna and flora [of Guiana]. In Rodway, J., 
Guiana 237-248. London. 1912, 

Rudolph, B. A., & Franklin, H. J. Studies of cranberries during 
storage. Fungi studies. Massachusetts Agr. Coll. Bull. 
198: 88-92. /. /, 2. 1920. 


Safford, W. C. Cultivated plants of Polynesia and their vern- 
acular names, an index to the origin and migration of the 
Polynesiana. Proc. Pan-Pacific Sci. Confer. 1: 183-187. 192 1. 

28o Indbx to American botanical literature 

Sampson, H. C. An ecological survey of the prairie vegetation 
of Illinois. Bull. Illinois Nat. Hist. Surv. 13: 523-577- pl- 

Saunders, £• R. Address to the Botanical Section [British 
Association for the Advancement of Science, 1920]. British 
Assoc. Adv, Sci, Rep. 88: 169-190. 1920, 

A genetical address. 

Sax, K. Sterility in wheat hybrids. L Sterility relationships 
and endosperm development. Genetics 6: 399-416. Jl 1921. 

Scala, A. C. Contribucion al estudio histologico de la flora 
chilena. RcvistaChilena Hist. Nat. 24: 73-78. /. 5-7. 31 Au 

Schiiroff, P. N. Die Teilung des vegetativen PoUenkerns bei 
EichJiornia crassipes. Ber. Deutsch. Bot. Ges. 40: 60-63. 
/. 7. 23 F 1922. 

Serre, P. La culture du cacaoyer au Costa-Rica. Bull. Mus. 
Hist. Natur. 1921: 260-264. 1921. 

Showalter, A. M. An orthotropous ovule in Hyacinlhus orientalis 
L. Torreya 21: 62, 63. /. i, 2. Au 192 1, 

Silveira, A. A. Especies novae civitatis Minas Geraes [Brazil]. 
Arch. Mus. Nac. [Brazil] 23: 159-171, pL 1-5. 1921, 

New species in Aspidosperma (i), Eriocaulon (4). 

Small, J. K. Seminole bread— the conti. A history of the genus 

Zamia in Florida. Jour, New 
pL 2j8-2^Q, 192 1. 

Includes Zamia umbrosa, sp. nov. 


Smith, C. P. Studies in the genus Lupinns—Vl. The Stiver^ 
siani, Concinni and SuhcarnosL Bull. Torrey Club. 48: 
219-234. /. 67-7^. 210. 1921, 

Smith, A, L. Lichens. Pp. 1-464. /. 1-135. Cambridge, 
England. 1921, In series of Cambridge Botanical Handbooks. 

Smith, L. Sugar cane in St. Croix. Virgin Is. Agr. Exp. Sta. 
Bull 2: 1-23. pi. I + /. /, 2. 192 1. 


Spegazzini, C. Mycetes chilenses. Bol. Acad. Nac. Ci, [Arg- 
entina] 25: 1-124. 192 1. [lUust.] 

Includes 5 genera and 113 new specie and forms of Fungi. 

Index to American botanical literature 281 

Spegazzini, C. Una niieva especie argon tina del genero '' Pro- 
sopanche/' Anal. Soc. Ci. Argentina 92: 251-256. /. 1-4. 
D 1921. 

Prosopanche Mazzuchii sp. no v. 

Spegazzini, C, Plantas nuevas o intcresantes. Anal. Soc, Ci 
Argentinisa 92: 77-123. S 1921. 

Includes new species in Atriplex (i), Porlulaca (2), Talinum (i), EcUn- 
ocactus (i). 

Sprague, T. A. Alphabetical list of nomina conservanda [Pliaii- 
erogamae]. Kew Bull. Misc. Infonii. 1921: 321-326. 1921. 

Sprague, T. A. A revision of the genus J^c/o/ia. Kew Bull. Misc. 
Inform. 1921: 270-278. 1921. 

Includes 5 new species from tropical America. 

Sprague [T. A.] & Riley [ . ]. P etasto ma Broadway i,^\n\i^\\e 
et Riley [Bignoniaceae]. Kew Bull. Misc. Inform. 1921: 
311, 312. 1921. 

Sprague [T. A.j. Rinorca Ilummelii, Sprague [Violaceae]. Kew 
Bull. Misc. Inform. 1921: 307, 308. 1921. 

Stevens, O. A. Pants of Fargo, North Dakota, with dates of 
flowering Am. Midi. Nat. 7: 135-156. S 1921. 

Stewart, F. C. The velvet-stemmed Collybia- a wild winter 
mushroom. Ann. Rep. (Geneva) New York Agr. Exp. Sta. 
37: 208-227. 1919- [Illust.] 


St. John, H. A critical revision of Hydrangea arhorescens. 
Rhodora 23: 203-208. 9 Ja 1922. 

Stout, A, B. Sterility and fertility in species of Ilcmcrocallis, 
Torreya 21: 57-62. f. i~8. Au 1921. 

Stout, A. B. Studies of grai)es in cooperation with tlie Slate 
Experiment Station at Gene\a, N. Y. Jour. New York 
Bot. Gard. 22: 148-156, pL 262, My 192 1. 

Tahara, M. Cytologische Studien an einigen Komi)ositen. 
Jour. Coll. Sci. Tokyo Imper. Univ. 43": 1-53. pL 1-4. 
30 Mr 1921. 

Tilden, J. E. The study of Pacific Ocean algae. Proc. Pan-Pacif. 
Sci. Confer, 1: 207-209. 1921. 

282 Index to American botanical literature 

Torrey, G, S- Les conldies de Ctinninghamella echinulata 
Thaxter. Bull. Trimest. Soc. MycoL 37: 93-99- P^^ ^o, f. 
Q-IJ' 20 Jl 1921, 

Torrey, G. S. Coronella nivea Crouan. Bull. Trimest. Soc. 
Mycol. 37: 88-93. pL 10, f. i-S. 20 jl 1921. 

Turrill, W. B. Plants introduced to horticulture from Chile and 
Argentina (including Patagonia and Fuegia). Jour. Royt 
Hort. Soc. 46: 346-350. 6 Je 192 1. 

Urban, I. Flora domingensis. Symb. An till. 8: 481-860. 
I Jl 1921, 


Includes new species in Lyonia (3), Bumelia (i), Matastelma (i), Cordia 
(i), Tournefortia (i), Ruellia (i). 

Van Hook, J. M. Indiana fungi— V. Proc. Indiana Acad. 
Sci. 1920: 209-214. 192 1. 

■- ~ > 

Van Hook, J, M. The pycnidium of Cicinnobolus. Proc- 
Indiana Acad. Sci. 1920: 215, 216. /. j. 1921. 

Van Hook, J. M. A tricotyledonous bean. Proc. Indiana Acad- 
Sci. 1920: 217. /. i. 1921. 

L + 

Vaupel, F. Melocadus depresstis Hooker. Monatsh. Kakteenk. 
32: 38. Mr 1922. [Illust.] 

From Brazil. 

Walker, J, C, & Jones, L. R. Relation of soil temperature and 
other factors to onion smut infection. Jour. Agr. Research 
22: 235-262, pL 25-^27 + /. 7, 2. 29 O 1921. 

Warnstorf, C. Die Unterfamilie der Scapaniodcen (Spruce 1885). 
Hedwigia 63: 58-1 16, 1921. 

Includes 7 new American species, 

Weatherby, C. A. Old time Connecticut botanists and their 
herbaria IL Rhodora 23: 122-125. Au 1921; 171-177- 

Weatherby, C. A. Other records of Salivina natans in the United 
States. Am. Fern Jour. 11: 50-53. 1921. 

Additional tothose from Minnesota;see Butters, ''Salvinia in Minnesota." 

Weatherwax, P. Anomalies in mai^e and its relatives— L Bull. 
Torrey Club 48: 253-255. /, 7-5. 1921. 

Vol 49. 

No. 10 




OCTOBER, 1922 

The genus Costus in Central America 

w- w. rowlee 
(with plates 12-15) 

As at present organized the genus Costus L. is composed of 
about one hundred species, divided almost equally between the 
Eastern and Western Hemispheres. Very few species occur in 
both hemispheres. The American species are most abundant 
in regions of heavy rainfall, maximum humidity and high 
temperature of both air and soil. The genus ranges from one 
Tropic to the other; it occurs wherever conditions are favorable 
in hot, warm and temperate regions, but is not found in high 
and cold districts. 

In any given locality the species do not usually appear abun- 
dant, but are scattered about in deep woodlands and undisturbed 
jungle. The plants are herbaceous perennials with spirally 
arranged leaves. The stems or culms are simple and represent 
branches arising from a system of underground rootstocks. 
They thus form gregarious clusters, some of the culms lemaining 
sterile and others producing spikes of flowers. In some cases the 
mats are more extensive than in others, this depending partly 
upon age and partly upon the habit of the plant. The leaves are 
ovate to lanceolate in outline and invariably entire. 

In Spanish American countries members of the genus are 
called "canagria" or ''cana agria," meaning "bitter cane," 

[The Bulletin for September (49: 259-282. pL 10, 11) was issued Sep- 

tember 22, 1922.1 


284 Rowlee: Genus Costus in Central America 

all jointed and unbranched stems being known as "cana" and 
the tissues in the present instance being very bitter. The plants 
are fleshy throughout their whole body structure, and their 
tissues are saturated with water, so that it is difficult to make 
satisfactory' herbarium specimens. Certain parts of the plant 
and especially the spikes arc so saturated as they grow in the 
field that it is easy to wring water from them. In drying they 
change their form markedly, and it is therefore most desirable 
to make observ^ations on specimens growing under natural 
conditions. The flowers in many cases are delicately colored 
and become fragile when pressed. 

The genus Costus^ together with two or three other genera, 
belongs in the subfamily Costoideac of the family Zingiberaceae, 
the group being ver>^ distinct in its general characteristics from 
the other genera of the family. Two types have been recognized 
in the genus: in the first the floral spike is borne upon what may 
be called a normal leafy culm; in the second the floral spike 
develops upon a modified culm. In the first type the culms are 
designated as "not diverse;" in the second, as "diverse." 

The species heretofore recorded from the Western Continent 
outside of South America have all belonged to the group in 
which the culms are not diverse. In the present paper tw^o 
Central American species with diverse culms are reported and 
proposed as new. They are not closely related to each other, 
but each is represented by allied species in South America, The 
paper reports in addition an undescribed species from Central 
America In w^hich the culms are not diverse and discusses the 
previously described Central American species. 

Key to the Central American species 

A. Culms diverse, the spikes borne on short bracteate 

B. Bracts of spikes with foh'aceous appendages 

(Costa Rica) i. C. hradcattis 

BB. Bracts of spikes not appendiculate (Guate- 

ruala) 2. C sepacmtensis. 

AA. Culms not diverse, the spikes borne on ordinary 

leafy ones. 

B. Bracts of spikes with foHaceous appendages. 
C. Spikes ovoid, green. 

D. Plants large, 2-3 m. tall; leaves 
about 30 cm. long, fuscous-hirsute, 
especially on the sheaths (Panama). 3. C. villosissimiis. 

Rowlee: Genus Costus in Central America 


DD. Plants small; leaves about 16 cm. 
long (Mexico, Costa Rica and Pana- 
ma) 4. C hirsuius, 

CC. Spikes cylindrical, red. 

D. Appendages of bracts lanceolate, 
acuminate, upper sterile erect; 
leaves glabrous (Guatemala and Costa 

Rica) 5. C. Bakeri. 

DD. Appendages of bracts broadly ovate; 

leaves scabrous (Costa Rica) 6. C. lima. 

BB. Bracts of spikes not appendiculate. 

C. Callose lacking on back of bracts (Costa 

Rica) 7. C Malortieaniis. 

CC. Callose present on back of bracts. 

D. Spike ovoid; leaves sessile (Mexico). 
DD. Spike fusiform (Costa Rica and 

8. C. pulvendenlus. 

Guatemala) 9. C, sanguineus. 

DDD. Spike cylindrical or spherical. 

E. Plants small, about 2 m. high. 
F. Leaves glabrous. 

G. Spike red, cylindrical 

(Costa Rica and Panama) 10. C spicatus. 
GG. Spike green, spherical 

(Guatemala) II. C. congcstus, 

FF. Leaves hairy beneath (Costa 

Rica and Panama) 12. C. nutans. 

EE. Plant large, much more than 2 m. 


F. Labellum orange-red with 
yellow markings (Costa Rica 

and Guatemala) 13. C. splendens. 

FF. Labellum yellow (Costa 

Rica) 14. C. laxus. 

I. Costus bracteatus sp. iiov. 

Plant clothed with long black hairs; leafy culms up to 2 m. 
high, the lower leaves reduced to sheathing scales; flowering 
culms 40 cm. tall, the leaves reduced to sheaths except the four 
or five uppermost, which bear small blades and closely surround 
the spike. Uppermost leaves on the sterile culms large and thin, 
2i2 cm. long, 7 cm. wide, oblanceolate, acuminate at the apex 
and tapering below evenly to an acute base, ferruginous, hair>^ 
on both sides, the hairs arising from a bulbous base; midrib 
slender; venation 3.5/23;* petioles i cm. long, densely ferru- 
ginous-toinentose ; ligules 5~io mm. long, tomentose; sheaths 

h ■ 

* This fraction indicates that there are 3.5 primary and 23 secondary veins 
to the centimeter. 

286 Rowlee: Genus Costus ix Central America 

overlapping, spreading-hirsute where exposed and with long 
appressed needle-like hairs where included in the next lower 
sheath: lower leaves reduced to sheaths with oblique orifices: 
sheaths of the flowering culms particularly loose, the upper- 
most lea\es forming a transition to the bracts of the spike. 
Spike ovate-cylindrical, 13 cm. long, 6 cm. wide, conspicuously 
bracteate, the bracts oblong, 2-5 cm. long, 1-8 cm. wide, puberu- 
lent on the back and along the margins, strigillose within; 
appendages of the bracts narrowed at the base, ovate, acuminate, 
rust>' pubcruletit: bracteole on the right side of the flower, 
conduplicate, 2.5 cm. long, with a linear callose 4 cm. long on 
the back near the apex, acuminate, rusty -pubcrulcnt. Flower 
3.5 cm. long; ovary 5 cm. long, three-ribbed, the ribs densely 
tomentose, the facies less so : calyx i .3 cm. long, rusty-puberulent, 
the lobes equal, triangular, 0,3 cm, long, acute: corolla three- 
lobed, very thin and delicate, nearly equalling the labellum, 
nearly as broad as long, reddish orange in the upper part: sta- 
minodia equalling the labellum, also reddish orange at the tip; 
stigma spoon-shaped with a notched appendage on the back. 

[Plate 12.I 

Costa Rica: in woods on the hillside about a mile south of 
Sicjuirres, H. E. Stork {Rowlee & Stork 6/^), Specimen pre- 
served in the herbarium of Cornell University. 

Only a single mat of the species, including one flowering culm 
and several leafy culms, was found, although careful search 
was made for additional material. The plant differs from any 
other species known to us and is remarkable in its peculiar 
vesture and delicate texture. It appears to be most closely 
related to K. Schumann's Costus Dinklagei from Cameroon, 
Africa. The locality where this novelty was found is interesting. 
The old Spanish trail from the mouth of the Matina River leads 
westerly along the foot of the mountains to near Siquirres. 
It then ascends the mountains along the south side of the gorge 
of the Reventazon River to the central plateau, where the cities 
of Cartago and San Jose are located. Our plant was found near 
this trail. There was also found near the same place a spur 
(for horse-back riding) quite unlike any known to the present 
generation and probably dating back to the days of the buccaneer. 

2. Costus sepacuitensis sp. nov. 

Vegetative and fruiting culms diverse: leafy culms with large 
elliptic leaves (resembling those of C maximtis), at least 30 cm. 
long and 9 cm. wide, softly pubescent on both sides, margin 
ciliate; petiole 5 cm. long; ligule 1.5 cm. long, ciliate on the 

Rowlee: Genus Costus in Central America 287 

margin, pubescent on upper part, glabrescent below; venation 
5/25: flowering culm at least 30 cm, long, bearing only sheaths, 
the. latter closely embracing the culm, sparsely puberulent. 
Spike not much exceeding the culm in thickness and appearing 
like it in texture and in color (dark chestnut in dried specimens) : 
bracts broadly ovate, 4 cm. long, and 3 cm. wide, glabrous, 
highly polished especially within. Flowers not seen. Fruit 
obovoid, pubescent, 1.5 cm. long and 0.8 cm. wide, crowned with 
a deeply parted calyx; calyx lobes unequal, 1.3 cm. long, separated 
to near the base, ovate, acute. [Plate 13. 1 

Guatemala: near the Finca Sepacuite, Alta Verapaz, April 
13, 1902, 0. F. Cook & R, F. Griggs 5g6. Known only from the 
type specimen, which is in the U. S. National Herbarium. 

This remarkable plant is related to C geothyrsus K. Schum. 
of Ecuador and C erythrocoryne K. Schum. of Peru but differs 
from both in the shape and size of the spike, in the calyx, and 
in the vesture of the plant. Other species with diverse culms 
occur in South America and in the Eastern Hemisphere. 

3. Costus villosissimus Jacq. Fragm. 51. pL So. 1 800-1 810 

Specimens referable to this species from Central America 
and the West Indies exhibit considerable variation, although 
agreeing in stature and in having appendiculate green bracts 
and ovate spikes. The hairiness varies, for example, from tawny 
shagginess to an almost smooth condition, while the color of 
the flower varies from sulphur yellow to white. In all cases, 
however, the margin of the labellum is tinted with pink. The 
most typical specimen collected by H. E. Stork and the writer 
is No. 401, found in June, 1918, on the west side of the Panama 
Canal, opposite Balboa. The flowers are large and showy, the 
labellum being tubular, about 7 . 5 cm. and ivory white 
except for the rose red border. The whole plant and particularly 
the upper leaves are clothed with long tawny hairs. This 
specimen was 3 m. tall. Another specimen, also in bloom but 
only half as tall, is No. 325a, collected in September, 191 8, at 


It agrees with No. 401 in having tawny 

hairs and in the color of its flowers. 

The species, although apparently rare, is wiclely distributed in 
tropical America. The following additional specimens from 
Central America may be cited: near Balboa, Canal Zone, 
Panama, H. Pittier 3776, 3779, 6690; Province of Alta Verapaz, 
Guatemala, 0. F. Cook & R. F. Griggs 30S. 

288 Rowlee: Genus Costus in Central America 

4, Costus hirsutus C. PresI, Reliq. Haenk. 1: 112. 1830 

This is a much smaller plant than the foregoing, according 
to the writer's interpretation, and is evidently not more than 
50 cm. high. The spike also is smaller. The following specimen 
seems to be characteristic; on the cattle trail between the 
Chiriqui Lagoon and the divide, western Panama, H, E. Stork 
{Rowlee & Stork 1045). Another specimen to be associated with 
the species is the following: upper stretch of the Saraquipi 
Ri\'er, II, Pittier 14166, Both represent a diminutive C. villo- 
sissimiis. The published descriptions of C hirsutus are meager, 
and no drawing or photograph of the species has been seen by 
the writer. 

5. Costus Bakeri K. Schum.; Engler, Pflanzenreich 4^^: 

387. 1904 

The following specimens of this species, all from the uplands 
of central and western Guatemala, have been studied: Barranca 
del Samald, Dept. Retalhulen, 1700 ft. alt., /. Donnell Smith 
280J; Barranca de Eminencia, Dept. Amatitlan, 1400 ft. alt., 
J. Donnell Smith 2802 (Type) ; Volcan Tecuamburro, Dept. 
Santa Rosa, J. Donnell Smith ^650, The following specimen 
from central Costa Rica has likewise been examined: A, Ton- 
dtiz 282, 

C. Bakeri is characterized by having a cylindrical and appendi- 
culate spike, more or less red in color; the appendages of the 
bracts being acute and longer than w^ide. In these respects it 
agrees with C. comosus (Jacq.) Roscoe, of northern Colombia 
and Venezuela, a species with which it was originally confused. 
In C comosnSj however, the upper surface of the leaves is softly 
hairy, while in C. Bakeri it is perfectly glabrous. It is probable 
that intermediates betw^een these tw^o species will be found. 

6. Costus lima K. Schum.; Engler, Pflanzenreich 4^^:388. 1904 

Dr. Schumann apparently knew C, lima from the type speci- 
men only, which was collected at the following -station on the 
Pacific side of Costa Rica: near Punta Arenas, January, 1854, 
Scherzer, The species is frequent however, throughout the 
lowlands in the northern part of Costa Rica, being found also 
on the Atlantic side. The following additional specimens may 
be cited: Buenos Aires, near the continental divide, A, Tondiiz 
6659; Livingston on the Reventazon River, Rowlee & Stork 648; 
Siouirres. Rowlee & Stork 1^18. 

Rowlee: Genus Costus in Central America 289 

It agrees with C. Baked in having an appendiculate, cylindri- 
cal spike, often much shortened, but the color is more brilliantly 
red, making the spike very conspicuous. The appendages, 
moreover, differ in being shorter and less striking. The species 
is further characterized by the peculiar vesture of the very 
opaque leaves, the upper surface being clothed with shorter 
bulbous hairs, all pointing toward the apex, while the lower 
surface is densely. covered by a tawny tomentum. The plants 
are large, often 5-7 m. high, and the spike is about the size of 
an ear of corn. 

According to the original account C. lima is related to C 
Friedrichsenii O. G. Peters., a species described from plants 
cultivated at Berlin. Dr. Schumann states that it was probably 
introduced from Central America, but the writer has seen no 
specimen agreeing with the description. 

7. Costus Malortieanus Wendl. Hamb. Garten- u. 

Blumenzeit, 19: 30. 1863 

So far as the writer's observ^ations go C. Malortieanus is 
confined to the coastal plains of northeastern Costa Rica. The 
type locality is near the Sarapiqui River at the head of navigation, 
and the original description was drawn from specimens cultivated 
In Germany. The species grows also along the Reventazon 
River, where it emerges on to the lowlands. It is the gem of 
the genus. The plants are small, being less than i nt. high, and 
grow in wide spreading mats, flowering in July and August. 
The flowers, w^hich are not freely produced, are very beautiful, 
suggesting in their tints some of the tropical orchids. The 
leaves are broadly obovate, densely hairy, and obscurely variegat- 
ed with darker and lighter threads of green. Specimens in the 
greenhouses of tlie New York Botanical Garden, although under 
cultivation for several years, have not as yet produced flowers. 
Two other species of tropical North America with unappen- 
daged bracts and no callose have been described under the names 
C piciiis D. Don, and C mexicanus Liebm. Specimens of the 
first two ha\'e not been found by the writer, but both are ap- 
parently of Mexican origin, C. pictus being based on plants 
grown in England. 

:29o Rowlee: Genus Costus in Central America 

8. Costus pulverulentus C. Presl, Reliq. Haenk. 1:41.1830 
The original specimen of C pulvemlenius was collected in 


Mexico by Haenke, no more definite locality being mentioned. 
The writer has identified with it, somewhat doubtfully, the 
following specimen from Guatemala, originally referred to C. 
Malorlieanus: Rio Sis, Dept. Suchitepequez, 1300 ft. alt., /. 
Donnell Smith 2,800. This and the following six species agree 
in having a callose on the back of the bracts. 

9. Costus sanguineus Donn. Sm. Bot, Gaz. 31: 122. 1901 

This species Is the most frequent representative of the genus 
in northern Costa Rica, and a photograph of it has been repro- 
duced in Calvert's "A year of Costa Rican Natural History/' 
opposite page 257, under the incorrect name C. Malortieanus. 
It is not found on the beach but inhabits higher ground, often 
at an elevation of as much as 2000 feet. The type specimen 
was collected in 1901, at Cubilquitz, Dept. Alta Verapaz, 
Guatemala, about 1000 ft. alt., by H. von Tuerckheim (No- 
7686), but the species has been found in other Central American 
countries bounding the Carrlbbean Sea. The following specimens 
may be cited; Peralta, i?ou'/e* & Stork 45; Livingston on the 
Reventazon River, Rowlee & Stork 653; and Siquirres, Rowlee & 
Stork 13 II, 1 31 2. The plant is about 1.5 m. high, and the 

leaves are softly pubescent. The red spikes are fusiform and 
acute, when perfectly developed, as shown in Calvert's figure, 
but they are frequently deformed and much shortened- 

10. Costus spicatus (Jacq.) Sw. Prodr. Fl. Ind. Occ, 11, 

The original material of C- spicatus came from the 
Indies, where the species is widely distributed. It is found, 



Rica, between LImon and the mouth of the Matlna River. The 
following specimens may be referred to it: Chargres, 185a. 
A, Fendler 446; Rio Dulce, March, 1889, /. Donnell Smith; 
vicinity of Frio, province of Colon, August, 191 1, H, Pittier 
4137; Limon, Rowlee & Stork 674. The field notes that follow 
were made from No. 674. 

Gulms 1-2 m. high, many from a rather dense mat, leafy to near the base, 
whofe plant glabrous. .Leaves 18 cm. long, 7 cm. wide, elliptic, short-acuminate 

Rowlee: Genus Costus in Central America 291 

at the apex, subauriculatc at the base, shining green above, glacuous be- 
neath, petioles 6 mm. long, sheath shorter than the internodes, ligulc 5 mm. 
lung, three or four leaves approximate to the base of the spike, venation 7735. 
Si)Ike oblong, 7 cm. long, 2.3 cm. thick, very compact. Bracts nearly orbi- 
cular, 2.5-3 cm. in diameter, Bracteole 1. 5 cm. long, relatively broad, boat- 
shaped, dextrad. Ovary 5 cm. long, glabrous, subtrigonous, seeds vcr>^ numer- 
ous, angular with elatcr-Hke aril. Calyx 5 cm. long, campanulate and with 
very short broad lobes and red margins. Corolla orange-red, 2 cm. long, 
dilated at the apex, lobed to the middle, lobes oblong, acute. Labcllum 3 cm. 
long, obovate, subtrilobcd, yellow. Stamen 2.5 cm. long, lanceolate, obtuse, 
anther adnate above the middle. 

II. Costus congestus sp. nov. 

Stem less than i m. high. Leaves crowded near the summit; 
petioles 5 cm. long; blade 24 cm, long, 6 cm. wide, acuminate at 
the apex, tapering from abo\'e the middle toward the base, 
slightly glaucous beneath, green above, glabrous throughout. 
Spike spherical, about 4 cm. in length and breadth; lower bracts 
with leafy appendages; bract proper orbicular, red, with a 
callose line. Ovary glabrous, 4 mm. long; calyx divided to the 
base, the lobes unequal, oblong, rounded, at the apex; petals 
large, hyaline, obtuse; labellum 4 cm. long, [Plate 14. J 

Guatemala : 



Sniiih 2036 (Type); Santa Lucia, Dept. Esculntla, 1045 ft. alt., 
March 3, 1905, W. A. Kellerman 3284' Santa Barbara, Dept. 
Solola, 1370 ft. alt., August, 1891, /. Donnell Smith 137. Type 
specimen in the herbarivim of Cornell University. 

Mexico: vicinity of Choapam, 
July 28, 1894, E. W. Nelson qii. 

The species is glabrous in all its parts. It differs from C. 
spicatus (Jacq.) Sw., to which Baker referred some of the speci- 
mens, in the size of the flower and in the shape and size of the 

12. Costus nutans K. Schum.; Engler, Pflanzenreich 4^®: 407. 


The type material of C. nutans came from the following local- 
ity: near Aguacate, Costa Rica, September, 1857 (in flower), C. 

Hoffman 727, 

The only specimen seen by the writer was collected on Mount 
Pirri, Panama, on June 12, 1912, by E. A. Goldman (No. 1963) 
at an altitude of 1350 m. [Plate i 

292 Rowlee: Genus Costus in Central America 

13. Costus splendens Donn. Sm. & Tuerckheim; Doniien 

Smith, Bot. Gaz. 33: 260. 1902 

Coshis maximus K. Schum. ; Engler, Pflanzenreich 4**^: 405. 1904, 
Costus gigantcus KunUe, Rev. Gen. 2: 687. 1891, Not Ridley, 


Tlie species was based on a specimen collected in Guatemala 
at an altitude of 350 m., by H. von Tuerckheim (No. 8015), no 
more definite locality being mentioned. The writer has not seen 
the type specimen, but other material collected in Guatemala bv 
Von Tuerckheim and referred to C. splendens has been available 
for study. In this material the bracts bear a callose below^ the 
apex, although no structure of this sort is mentioned in the 
original description. Since these specimens are indistinguishable 
from C. maximus K. Schum., the name C, splendens will have to 
supplant the later name C, maximus, and the known range of 
the species wull extend from Guatemala to Panama. 

14. Costus laxus 0. G. Peters.; Martins, Fl. Brasil. S"^: 56. 



The type specimen was collected in Costa Rica by Aented 
in 1847, the name of the locality being illegible. The species 
is known also from Venezuela. 

Cornell Univeksity 


Explanation of plates 12-15 

Plate 12. Costus bracteatus Rowlee. 

Plate 13, Costus sepacuitensis Rowlee 

Plate 14. Costus congestus Rowlee. 

Plate 15. Costus nutans K. Schum. 

Bull. Torrey Club 

Volume 49, plate 12 

J V 





Bull. Torrey Club 

Volume 49, plate i,^ 


Bull, Torrey Club 

Volume 49, plate 14 



"^ > 





? ff 








. / 







Bull. Torrey Club 

Volume 49, plate 15 


Quercus lyrata in Iowa 

B. Shimek 

(with plates 1 6 AND 1 7) 

The discovery of the overcup oak, Quercus lyrata Walter, 
in Iowa, by the writer, so far extends the range of this species 
that it seems worthy of more detailed record. Sargent says 
that the species "is distributed from the valley of the Patunxent 
River in southern Maryland southward near the coast to western 
Florida, through the Gulf States to the valley of the Trinity 
River in Texas, through Arkansas and southwestern Missouri, 
where in a swamp near Allentown, there is a single specimen, 
the most northern known representative of the species west of 
the Mississippi River, to central Tennessee, southern Indiana, 
and Jasper County, Illinois." 

Other authors give the distribution less fully, but in all the 
references consulted the range falls within that here given. The 
herbarium of the Missouri Botanical Garden at St. Louis 
contains several specimens from the southern counties of Illinois, 
and from Butler Cojinty, Missouri, but these localities also 
lie within the territory given by Sargent. 

The Iowa specimens are located in Iowa County, near the 
tow^n of Amana. This not only extends the northern range 
west of the Mississippi, but the locality is more than two degrees 
of latitude north of the Maryland and Illinois localities, making 
this the northernmost point from which the species is known. 

Thus far three trees have been found. They are located in 


the low bottom land timber along the low^a River, the locality 
being subject to overflow and always quite moist. They are 
associated with the large, bottomland form of the bur oak, 
Querctis macrocarpa, a few trees of Q. hicolor, and the ordinary 
bottomland species of low^a valleys, such as Ulmus americana, 
Behila nigra, Popidus deltoides, Acer sacchariniim, A. Negundo, 
Platanus occidentalism and Juglans nigra. Other trees of the 
species will probably be found among the ''bur-oaks." 

The finest of the three specimens (shown in Plate i6, fig. i) 
is about 70 feet high, and the trunk measures 22 inches in diam- 
eter at a height of 3 feet. Its rate of growth is probably approxi- 

*Silva of North America 8: 48. 1895 


29+ Shimek: Quercis lyrata in Iowa 

mately the same as that of the large bottomland form of Q. 
macrocarpa, and a nearby specnnen of the latter, recently cut, 
measured 27 inches in diameter and exhibited about 120 rings, 
those from about 55 to 85 showing the most rapid growth. Our 
overcup oak trees are^ probably not less than a hundred years 
old, thus antedating the coming of the white man to this section. 
The trees have been observed both in flower and fruit. Sar- 
gent gives tlie time of flowering as March or April, but in this 
jiorthem locality the flowers do not appear until about the 
middle of May- 
Two of the trees produce acorns which are entirely enclosed 
in the cup ; the third has a partly open cup, a form not infrequent 
in the South. The peduncles of the cups are 6-16 mm. in length. 
The wall of the cup is 3-6 mm. thick or even thicker at the 
base, but thins upward. The scales of the cup arc prominently 
tubercular, the tubercles becoming smaller upward. The small 
opening has a short fringe. 

The nut is short-ovate, with a broad basal scar and is 16-19 
mm. in length, with about the same diameter. Its tops and 
sides are covered w^ith a short, grayish pubescence. For cup 
and acorns see Plate 16, fig, 2. 

The general aspect of the tree is not unlike that of the large 
form of Q. macrocarpa which grows in somewhat better drained 
spots in the same timber. The latter is here not gnarled or 
stunted, as is usually the case in more exposed localities. It 
is probable that our species has been mistaken for the latter in 
other localities. 

The leaves are distinctly difi"erent from those of the nearby 
bur-oaks. As compared with the latter they are thinner, with 
more distinct veinlets; lighter green; more irregularly lobed, 
with the terminal lobe rarely coarsely crenate; usually smaller 
and narrow^er; and the upper surface is more likely to have scat- 
tered short hairs at maturity, while the pubescence of the lower 
surface is much less dense. Plate 16, fig. 3, shows young 
leaves, and Plate 17 shows mature leaves taken in different 
years. The leaves and acorns shown on the plates are from the 
same tree. 

On the whole the Iowa specimens of Q. lyrata are quite 
typical . 

Shimek: Quercus lyrata in Towa 29^ 

Explanation of plates 16 and 17 

p*LATE 16 

Fig. I. Quercus lyrata Walt., a leafless tree, 

Fig. 2. Cups and acorns, x .55. 

Fig. 3. Young leaves and flowering twig^ x .42 

Plate 17 

Fig. I. Mature leaves of Quercus lyrata, x .24. 
Fig. 2. Old leaves, taken a year earlier, x .30, 

Bull. Torrky Club 

Volume 49, plate x6 



Bull. Torrkv Club 

Volume 49, plate 17 



References to the algae in the Chinese classics 


(with one text figure) 

The following notes on the algae referred to m ancient 
Chinese literature are at best sketchy, but are offered neverthe- 
less for what they are worth. The references to the texts and the 


ictor of Biology at St, John's Uni 
Mr. C- F. Wu, formerly of St. John 

The writer presents this paper with a view to throwing some new 
light on the development of science in general, and to intro- 
ducing to the West, besides, some evidences of Far Eastern 
activity in this line, which parallels, if not predates, that of 

Agriculture in China dates back to Shen Nung,'^ an emperor 
of the legendary period, 3000 B.C. He was said to be the first 
farmer and taught the people to till their fields. Since this 
mythical age, the people of China have been farmers primarily. 
As the Old Testament was essentially the expression of an 
agricultural and pastoral people and is, in consequence, re- 
plete with similes and references to plants and animals, so in the 
Chinese classics w^e find the farming life of the people, a life 
of continuous touch with nature, coming to expression In the 
frequently recurring allusions to animals and plants, and in the 
use of terms connected therewith. Long before the Aristotelian 
age of hearsay and philosophical conjecture, the ordinary facts 
of farming and floriculture had entered the realms of Chinese 
literature in the form of terms serving as figurative expressions 
for desirable characters and virtues. In order to have given 
time for everyone to become so familiar with these original 
terms that they crept unconsciously Into speech and literature 
as specific classifiers, observation of the form and structure of 
plants must already have proceeded far. This Is offered as 
one evidence of age-old familiarity with the facts of nature. 
The second is coincident with the first and deals with the ideo- 
graphs representing these terms and ideas. The discovery and 
use of the facts of nature called into being special ideographs to 

Appendix, "Peking," by Juliet Bredon 


298 Porterfield: Algae in the Chinese classics 

represent them, so that owing to the character of the language 
and its slow evolution, we feel that the appearance of these 
specific terms in the ancient literature places the knowledge 
back of them at a very early date. 




e— : 



Chinese character, as we are accustomed to term the ideograph, 

is built up of many radicals or elementary symbols supposed 

formerly to have been pictures which taken together represent 

an idea. The character for algae is 

Tsao (Fig. i) and may be resolved 

into four radicals, the ones for grass, 

water, wood, and mouth, the latter 

being repeated three times. The 

Fig. I. Tsao mouth radical is a simple rectangle, 

a, grass radical; &, mouth which when repeated three times and 

radical;., wood radical; ..wa j^^^^ j^ ^-^^ ^^^^ ^f ^ pyramid, 

radical;/, small ideograph mean- , ^11 1 ^ 

ing rank or segment. ^^^ ^* ^^^ t)^^"^^ ^"^ ^"^ ^" ^op, 

make up an ideograph which means 
rank or character. This latter smaller ideograph seems to 
carry with its meaning the idea of segmented, or possibly cellular 
(!) structure because of the shape of the thrice repeated box- 
shaped radicals. Evidently the idea in their minds from the 
character used was that of a grass-like, fibrous or stringy, 
cellular plant that grows in the water. Such an analysis leads 
us to believe that they had in general a good idea of what an 
alga is. 

The character for Tsao first appeared in one of the five classics, 
the Canon of Boems (Fig. i). In the chapter Chao Nan there 
appears the following passage, the romanized version of which 
is here given: ** 'yii bih bien tsao" (with respect to the collecting 
of algae). This term is still used today. The term for water 
plants is a much simpler character and therefore is much more 
general in its meaning. It has the grass and water radicals but 
no specifically descriptive ones. There can be no confusion 
then in the use of these terms. 

In support of the cellular or segmented idea of algae brought 
out in the analysis of the character, K'ung An-kuo, in the Canon 
of History, says that **an alga is an aquatic plant that has 
systematically arranged branching parts, and is used, therefore, 
(figuratively) to denote literature." He may here be referring 

Porterfield: Algae in the Chinese Classics 299 

to the diagrammatic structure of Hydrodictyon, Nearly two 
thousand years ago in a pond beside the temple of Han Voo Tee, 
algae were said to have grown to a length of nine feet. Among 
other characteristics they were reported as having the appear- 


ance of a net from w^hich certain water birds were said to have 
had great difficulty in extricating themselves. The people called 
this the water-net alga. The structure of such types was 
plainly visible under careful scrutiny, hence a netted or reti- 
culate structure was specifically attributed to algae. 

About 600 B. C, in a book entitled Sze tsen, occurs the follow- 
ing reference to algae: ''Some algae are a delicacy fit for the 
most honorable guest, even for the king himself." At the 
present time country people gather Nostoc for food. This is 

called "Heaven vegetable." Red algae are dried and eaten 
by farmers who live lear the sea. It is to some of these doubt- 
less that the quotation refers. 

The real knowledge in Chinese medicine as it exists today is 
based on the Chinese ''Materia Medica,'' the edition of which 
was begun four thousand years ago. The present edition was 
written two hundred years ago In the Ming dynasty. Among 
other plants, Kw'un Boo, or Laminaria is mentioned as being 
useful for medicinal purposes, for which it is calcined after being 
washed and sun-dried. It is a common practice in China to 
pack open cuts with ashes in order to stop the bleeding. Whether 
there is any discrimination shown as to what kinds of ashes are 
used the writer is not prepared to say. If there is, it is interesting 
to note that in case preference was shown for ashes of kelps, 
they must have realized that some medicinal virtue was con- 
tained In them. Iodine as an element was probably not known 
to the Chinese but they may have realized its presence as a 
virtuous remedy in other things. 

As in western countries the sea weeds of the China coast are 
used by the farmers nearby for fertilizer and also, when dried 
for fuel. Agar-agar is made by the Chinese out of certain species 
of sea-weeds, a well known fact to most scientists and technicians. 

The morphological characters of these marine forms seem to 
have been as well know^n as the freshwater forms, if not better. 
Many were said to be attached to stones. The large leafed 
forms were known as ox or horse algae. Some float on the top 
of the w^ater, others live at the bottom. Of the smaller forms, 

300 Porterfield: Algae in the Chinese Classics 

many have long silk-like filaments, the longer ones more than 
thirty ** segments" each, and others have the appearance of 

"uncombed hair." 

Metaphorically the term for algae is used in a very compli- 
mentary sense. The elegance and beauty of essays was often 


designated by or compared to that of the algae. The term was 
used in praise of the thinking of a learned man, signifying that 
his thoughts were as systematically ordered as the parts of an 
^lg3.. Judgment was in like manner complimented. The 
algae contributed not only to the language and literature of 
the Chinevse but also to their art and superstitions. In an old 
book called Zong Shu, we find reference to conventionalized 
designs derived from algae being included in the embroidery of 
their garments. When the figures of algae appeared on the ends 
of the roof beams of their houses in brilliant colors, we find that 
the underlying idea was protection from fire. Because algae 
were known to be water plants, any evidence of them on houses 
was a protection against fire in that the former invoked the aid 
of their native element to drive away the fire spirit. 

From this discussion we begin to realize that from direct 
references in ancient Chinese literature and an analysis of the 
ideograph, there is a possibility, if not a probability, that the 
knowledge of the algae as a distinct morphologic unit in the 
plant kingdom dates back to very early times, as compared 
with the state of knowledge in western countries. Moreover 
this knowledge seemed to be more wide spread amongst the 
people from the use of Tsao in a metaphorical sense, in its 
practical use as medicine, as food and fuel, and as fertilizer; as 
the basis of commercial products; and in the realization of Its 
decorative value and its superstitious meaning. 

A Lachnea with a botryose conidial stage 


B. O, Dodge 
(with seven text figures) 

While engaged in culturmg vspecies of Discomycetes for the 
purpose of studying the nature of the origin of the ascocarp, an 
interesting Lachnea was collected several different times in 1912 
on a variety of substrata in the vicinity of New York City. The 
apothecium of this fungus, which was identified* at the time as 
Lachnea abundans Karst., originates In a long winding multicel- 
lular ascogonlum, so similar to that described by Fraserf for 
L, cretea that, when we consider also the characters of the fruit 
bodies and mycelium, there appears to be little doubt that the 
fungi with which both of us were working are at least very 
closely related species. Belie v^Ing that the characteristics of the 


primordla are of fundamental importance In determining re- 
lationships, further discussion, other than the mention made 
in a paper published at that time, J seemed unnecessar^\ 

Miss Fraser does not state that she grew L, cretea in pure 
cultures. Her cultures were originally derived presumably by 
transfers from a plate culture overrun with moulds and some 
hyphomycetous fungus. It would be necessary to know whether 
L. cretea has a conidial stage like that which we have connected 
with L. abundans before we can be certain that the species are 
identical. In view of the discussions that have arisen since the 
publication by Seaver and Home* of a paper on Sclerotinia 
Geranii and another paper by Godfrey! on Sclerotinia Ricini^ 
both of these forms having Botrytis conidial stages, it has been 
thought advisable to point out that there are other types of 
Discomycetes which also have asexual fructifications closely 
resembling Botrytis, 

* The identification has been confirmed by Dr. F. J. Seaver, who is con- 
sidering L, cretea as a possible synonym. 

tAnn, Bot. 27: 553-563. 19^3- 
JBull. Torrey Club 41: 165 1914. 

* Mem. Torrey Club 17: 202-206. 1917. 
t Phytopathology 9: S^SS^J- I9^9- 


Dodge: Lach 

The conidia of Lachnea abundans are smooth at maturity 
while those of Sclerotinia Geranii are rough or warted, otherwise 
their conidial stages are very similar. The connection between 


abundans has been 

established repeatedly by cultures from single conidia and from 
single ascospores, apothecia arising in each case within a week 
or two after showing the spores. 


Fig. I. Part of a dichotomously branched conidiophore showing the en- 

upon which spores 

Fig. 2. End branches of a similar stage more highly magnified, spore 
buds arising from two of the globular heads. 

Fig. 3. Portion of an old conidiophore showing botr>'Osc clusters of 



Figs, 4 and 5. Germinated ascospore becomes a coll in the hypha; Fig. 5 
also shows ungerminated ascospores. 

Ascospores germinate very' readily when the inoculated culture 

Dodge: Lachnea with a botryose conidial stage 303 

medium is heated to 70° C. for about fifteen minutes. This 
method has been found effectual in inducing germiiiation of 
ascospores of certain other species of Discomycetes.J Tiic out- 
lines of the spore are generally obscured as it germinates and 
becomes a cell in the hypha (Figs. 4, 5,). Theconidia as well 
as the ascospores remain viable for a long time if kept dry in the 
laboratory, the former, sometimes living three or four 
years. Should several conidia be sowed together in a culture 
there follows at once a great amount of anastomosing of the 
germ tubes. 

The fungus grows well on almost any of the ordinary culture 
media. Conidiophores arise the second day In cultures on milk, 


bouillon, potato agar, etc. An agar medium in which the nutrient 
is a decoction from heated soil is especially favorable for the 
development of ascocarps. A potato dextrose agar gives an 
abnormal amount of conidial development. The conidia are 
formed on the spherical or knob-shaped ends of regularly dichoto- 
mously branched conidiophores, seven or eight such divisions 
often occurring. One might consider an aerial hypha whose 
main axis ends in a pair of sporophores as a part of the coni- 
dlophore system even though branches ultimately bearing conidia 
arise at irregular Intervals from It. Fig. i shows a branch of the 
third order at the time spore formation is just beginning. A 
portion of such a branch is further enlarged in FiG. 2, showing 
conidial buds from two of the ultimate branchlets. As spores 
mature they hang together in botryose clusters covering about 
eight pairs of end branches (Fig. 3, A). At B in this figure the 
conidia have been dislodged, exposing the extremities of the 
conidiophore subdivisions, now collapsed. The length of the 
conidiophore system and the number of times dichotomous 
branching occurs depend of course upon the kind of nutrient 
in the medium. On soil decoction agar the sporophores are very 
short, and there is very little of the aerial type of hyphae. The 
mycelial hyphae on the other hand branch more or less dichoto- 
mously and In this respect also the fungus is like the L. cretea 
studied by Fraser. The color of the aerial hyphae, conidiophores 
and conidia in mass varies from pale ochraceoiis buff to vinaceous 
buff (Ridgway), depending on age and vigor of growth. 

X Mycologia 4: 218-222. 1912. 

304 Dodge: Lachnea with a botryose cinidial stage 

•The conidia are spherical, smooth, faintly colored, pale 
ochraceous buff in mass, 7-9 [i in diameter. The ascogonia 
begin to appear, as noted, in about ten days, and mature fruit 
bodies will be formed within two or three weeks. Fraser lays 
considerable stress on the branching of the trichogyne end of 
the ascogonium of L. cretea, thinking such vegetative growth 
the "progressive degeneration" of Atkinson, indicates that the 
species is becoming apogamous. Many such abnormal or 
aborted ascogonia are always found in cultures of these Dis- 
comycetes and they should be carefully distinguished from those 


Fig, 6. Hairs from the margin of the apothecium, ascus with spores and 

Fig. 7. Part of a occtlon of a small apothecium showing the character of 
the (fells of the wall and of the margin. 

normal primoridia which develop into fruit bodies. Ascocarps of 
L, ahundans (Fig. 7) are 1-3 mm. in diameter and uniformly 
ochraceous in color. The hairs (Fig. 6) are about 150 ^ long, 
the asci 130-150 yi, and the ascospores 7-8 x 12-14 ]l. The 
paraphyses, which are enlarged rather abruptly at the ends, are 
4-5 yi broad. 

It Is certainly dangerous to place too much weight on the 
asexual spore forms in determining relationships of their ascogen- 
ous stages in the face of these apparently anomalous cases and 
such others as we find in Ascoholus magnificas, which is connected 

Dodge: Lachnea with a botryose conidial stage 305 

with a Papulospora^ similar to those forms hitherto thought to 
belong to Melanospora, If one, following Saccardo, for example, 
seeks to find a description which might apply to the conidial 
stage of Sclerotinia Geranii or of Lachnea abundans he turns 
directly to the forms along with Botrytis cinerea on the basis 
of the spore clusters. It seems to the writer, therefore, an im- 
possible task to determine the real generic affinities of species 
of form genera from a study of the conidial or pycnidial stages 
alone. Brierly* has recently published a paper on Botrytis 
cinerea. His argument is based entirely on the assumption that 
this species has no perfect or ascogenous stage. It is certainly 
unusual to state that a fungus has no ascocarpic stage simply 
on the basis of large numbers of cultures that produce only 
conidial fructifications. The recent work of Bensaude, Kniep 
and others, on several of the well knowm Basidiomycetes em- 
phasizes the importance of growing in pairs strains or races 
derived from different spores, at least as a last resort, in attempts 
to obtain the *' perfect" stages. We are finding more and more 
forms in which a strain, which is sterile when grown alone, still, 
when grown together with some other strain, at once takes 
part in the development of a "perfect" stage. 

There is no question that Lachnea abundans Is homothallic, 
a culture from a single conidium or a single ascospore being 
capable under suitable conditions of giving rise to ascocarps. 
Whatever may be the standing of the species now commonly 
referred to the form genus Botrytis, after study has revealed 
their ascomycetous connections, it Is interesting to find an 
'* imperfect'' fungus w^Ith a botryose conidial stage connected 
with a little Lachnea which may be so easily cultured on ordinary 

Bureau of Plant Industry, 
U. S. Department of Agriculture, 
Washington, D. C. 

* Mycologia 12: 115-134. 1920. 

* Phil. Trans. Roy. Soc. B. 210; 83-114. 1920. 



The aim of this Index is to include all current botanical literature written by 
Americans, published in America, or based upon American material; the word 
America being used in the broadest sense. 

Reviews, and papers that relate exclusively to foresty, agriculture, horti- 
culture, manufactured products of vegetable origin, or laboratory methods 
are not included, and no attempt is made to index the literature of bacteriology. 
An occasional exception is made In favor of some paper appearing in an Ameri- 
can periodical which is devoted wholly to botany. Reprints are not mentioned 
unless they differ from the original in some important particular. If users 
of the Index will call the attention of the editor to errors or omissions, their 

kindness will be appreciated. 

This Index is reprinted monthly on cards, and furnished in this form to 
subscribers at the rate of three cents for each card. Selections of cards are 
not permitted; each subscriber must take all cards published during the term 
of his subscription. Correspondence relating to the card issue should be 
addressed to the Treasurer of the Torrey Botanical Club. 

Rhoads, A, S. The pathology of Lupifius arboreus^ with special 

reference to the decay caused by two wound-parasites, 
Collyhia velutipes and Pleiirotus ostreatus. Phytopathology 
11: 389-404. pL i8~20. 16 F 1922. 

Rigg, G. B* Identity of cetain yellow pigments in plants and 

animals. Science II, 55: loi, 102. 27 Ja 1922. 

Robbins, W. W. Mosaic diseases of sugar beets. Phytopathol- 
ogy. 11: 349-365-/- i~8. 27 Ja 1922. 

Robertson, C. Flowers and insects, XXI. Data of anthecology, 


Bot. Gaz. 73: 148-152. 15 F 1922. 

Robertson, C. Flower seasons. Sci. Mo. 14: 201-203. F 1922. 

Robertson, C, The sunflower and its insect visitors. 'Ecology 

3: 17-21. 14 Mr 1922. 

Rock, J, F. Hunting the chaulmoogra tree. Nat. Geog. Mag. 

41: 243-276. Mr 1922. [Illust.] 

Tarakiogejios Kurzii of Siam. 

Rothgeb, B. E. Cultural experiments with grain sorghums in 

the Texas panhandle. U. S. Dept. Agr. Bull. 976: 1-43. 

/. /-//. 20 Ja 1922. 


3o8 Index to American Botanical Literature 

Rydberg, P. A. Phy togeographical notes on the Rocky Mountain 
region, X. Grasslands and other open formations of the 
Montane Zone of the Southern Rockies. Bull. Torrey 
Club 48: 315-326, 28 F 1922. 


Saccardo, P. A. Micetes Boreali-Americani. Giorn. Bot. ItaL 

27; 75-88. 1920. 

Sage, H. The royal palm (Oreodoxa regia). Am- For. 28: 85- 

88. F 1922. [Ilhist.] 

Sax, K., & Gowen, J. W. Productive and unproductive types 

of apple trees. Jour. Hered. 12: 291-300. 20 F 1922. 



Schaffner, J. H. Additions to the catalog of Ohio vascular 

plants for 1921. Ohio Jour. Sci. 22: 91-94. Ja 1922. 

H, The classification of plants. — XII, Ohio Jour. 

Schaffner, J. H, The classification of plants.— 

Sci. 22: 129-139. Mr 1922- 

Schaffner, J. H. Control of the sexual state in Arisaema tri- 
phyllitm and Arisaema Dracontium. Am. Jour. Bot, 9: 
72-78. 20 Mr 1922. 

Schaffner, J. H. Progression of sexual evolution in the plant 
kingdom. Ohio Jour. Sci. 22: 101-113. F 1922. 

Schlechter, R*, & Hoehne, F. C. Contribuisoes ao conheci- 
mento das Orquiddceas do Brasil. Anex. Mem. Inst. 
Butantan 1: 9-68. jrontis,-\-pL 1-14. Mr 1922. 

Includes Fractiunguis and Pseudostelis, gen. nov., 38 new species and 
2 new varieties. 

Schneifer, C. Notes on American willows — XI. Jour. Arnold 
Arbor. 2: 185-204, Ja 1922. 

Setchell, W. A. Aboriginal tobaccos. Am. Anthropologist 23: 

397-414. O-D 1921. 

Includes map. 

Severin, H. H. P. Minimum incubation periods of causative 
agent of curly leaf in beet leafhopper and sugar beet. 
Phytopathology 11: 424-429. /. 1-4. 16 F 1922. 

Index to American Botanical Literature 309 

Shear, C. L. Frederik Georg Emil Rostrup. Phytopathology 

12: 1-3. With portrait. 10 Mr 1922. 

Shull, G. H. Three new mutations in Oenothera Lamarckiana. 

Jour. Hered, 12: 354-363./. 8-16, O 1921. 

Sinnott, £• W., & Blakeslee, A. F. Structural changes asso- 
ciated with factor mutations and with chromosome, muta- 
tions in Datura. Proc. Nat. Acad. Sci. 8: 17-19. 15 F 1922. 


Sismey, E* D. A contribution to the algal flora of the Ojanagan 

[British Columbia], Canad. Field Nat. 35: 112-114. 3 

F 1922. 

Small, J. K. Historic trails, by land and by water. Jour. New 
York Bot. Gard. 22: 193-222. pL 263-166. D 1921. 

Record of exploration In Florida in Dec. 1919. 

Small, J. K. Wild pumpkins. Jour. New York Bot. Gard. 
23: 19-23. F 1922. 

[ey, F. J., & others. Weeds of Californi 
control. Monthly Bull. Dept. Agr. of Cc 
11: 1-22, 1-360. /, 1-138. F-Mr 1922. 


Spaulding, P. Investigations of the white pine blister-rust. 
U. S. Dept. Agr. Bull. 957: i-ioo. /. /-/J. 6 F 1922. 

Stephenson, H. T. Trees in winter. Am. For. 28: 79-84. F 

1922. [Illust.I 

Sterrett, W. D. A new oak from the Gulf States. Jour. Elisha 
Mitchell Sci. Soc. 37: 178, 179, Mr 1922. 

Quercus ashei sp. nov. 

Stout, A. B. Cyclic manifestation of sterility in Brassica peki- 

nensis and B. chinensis. Bot. Gaz. 73: TIO-132./. 1-7. 15 

F 1922. 

Sutherland, J. American trees in Scotland. Jour. For. 20: 71- 

74. Ja 1922. 

Sydow, H. Novae fungorum species — XVII. Ann. Mycol. 19: 

304-309- 1 92 1. 

Includes new American species in Cryotosphaeria (i), Cihhcra (i), and 


Phaeocibosia (i). 

Jio Index to American Botanical Literature 

Taylor, W- R- The embryogeny of Cystanthtis parviflonis 

Baker. Am, Jour. Bot. 8: 502-506. pL 15, 2,6. 15 F 1922. 

Trelease, W. The peltate peperomias of North America. 

Bot Gaz. 73: 133--147. pL 1-4. 15 F 1922. 

Includes 10 new species from Mexico and Central America. 

True, R. H. The significance of calcium for higher green plants. 

Science II. 55: 1-6. 6 Ja 1922. 

Uphof, J. C. T. Ecological relations of plants in southeastern 

Missouri. Am. Jour. Bot. 9: 1-17. pi. i, 2+ /. /-J. 21 F 

Vaupel, F^ Echinocactus Williamsii Lem. Monatsschr. Kak- 

teenk. 31: 186. D 192 1. [lUust.] 

Vaupel, F. Fruch tender Cerciis Strausii (H.) Vpl. Monatsschr. 

Kakteenk. 32: 8, Ja 1922. [Illust.] 

Vinall, H. N., & Cron, A. B. Improvement of sorghums by 
hybridization. Jour, Hered. 12: 435-443. frontispiece -\- 
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Wann, F. B., & Muenscher, W. C. A preliminary list of the 

38-41. 6 Mr 


Warburg, O. Die Pflanzenwelt. 3: 1-552. pL 1-27, +/, 1-278. 

Leipzig. 1922. 


Jour. 11: 109-113. pL 2. 31 M 

Webb, R. W. Studies in the physiology of the fungi. XV. 
Germination of the spores of certain fungi in relation to 
hydrogen ion concentration. Ann. Missouri Bot. Gard. 
8: 283-341./. i~jQ. S 1921. 

Weidman, R. H. Forest succession as a basis of the silvi- 
culture of western yellow pine. Jour. For. 19:877-885. 
D 1921. 

Index to American Botanical Literature 311 

Weingart, W. Cereus geometrizans Mast, und Verwandte. 
Monatsschr. Kakteenk. 32: 17-20. F 1922; 35-38. Mr 

Weingart, W. Cereus princepsHovt . Wiesb. Monatsschr. Kak- 
teenk. 32: 20. F 1922. [Illiist.] 

From Guatemala. 

Weir, J. R, Cenanginm pint phil urn n. sp., an undescribed 
canker-forming fungus on Piniis ponderosa and P. conlorla. 
Phytopathology 11: 294-296. pi. ij -\-f-i, 2, 15 N 1921. 

Weston, W- H. A note relative to the reappearance of the 

sugar cane downy mildew in the Philippines. Phytopath- 
ology 11: 371-375- 27 Ja 1922. 

Wherry E. T. Soil reaction in relation to plant growth. Am. 

Jour. Pharm. 94: 110-114. F 1922. 

Reported by Dr. H. Leffmann. 

Wiggans, R. G. A classification of the cultivated varieties of 

barley. Cornell Univ. Agr. Exp. Sta. 46: 369-456. pi. 34- 
j8 +/. 7-72. S 1 92 1. 

Williams, F. N. Critical notes on some species of Cerastium. 
Jour. Bot. 59: 324-329. N 1921; 59: 349-3vS3- D 1921.' 

Includes C. glutinosum glabratum, var nov., from Colombia. 

Williams, R. S. Mosses from British Guiana and Dominica, 

Lesser Antilles, collected by Miss E. F. Noel in 1914. 
Bryologist 24: 65, 66. pL 4. 15 F 1922. 

Macromilrium trinilense sp. nov. 

Willis, J. C. Endemic genera of plants in their relation to others. 
Ann. Bot. 35: 493-512- O 1921.' 

Wilson, E. H. The cherries of Japan. Publ. Arnold Arb. 
7. 1916. 

Wilson, E. H. The conifers and taxads of Japan. Publ. Arnold 
Arb. 8. 1916. 

Wilson. E. H. Notes from Australasia. II. The New Zealand 

forests. J 


Wilson, T. R. C. The effect of spiral grain on the strength of 
wood. Jour. For. 19: 740-747./. 1-4. N 1921. 

312 Index to American Botanical Literature 

Winston, J. R, Commercial tontrol of citrus scab. U. S. Dept. 

Agr. 215: 3-8. 15 F 1922. 

Wolf, F. A., & Shunk, I<V. Tolerance to acids of certain bac- 
terical plant pathogenes. Phytopathology 11 ; 244-250. 
21 O 1921, 

Woodcock, E. F,, & Zeeuw, R. de. The anatomy of the haustor- 
ial roots of Comandra. Michigan Acad. Sci. Rep, 22: 189- 
192. pL IS. Au 1921. 

Wright, W. G. Research work of the Dominion Forest Service. 

Jour. For, 20: 62-66. J a 1922. 

Young, H. C, & Bennett, C. W. Studies in parasitism I. Toxic 
substances produced by fungi. Michigan Acad. Sci. Rep. 
22: 205-208. Au 1921, 

Yuncker, T. G. A list of Indiana mosses. Proc. Indiana Acad. 
Sci. 1920: 231-242. 192 1. 

Yuncker, T. G. A species of Cusctita not hitherto reported 
from Indiana. Proc. Indiana Acad. Sci. 1920: 229. 192 1. 

Zahlbruckner, A. NeueFlechten — IX. Myc. 19:224-242. 1921. 

Includes new species in Opegrapha (2), Graphis (i), Graphina (3), En- 
terodyction (i). 

Zahn, K, H, Compositae — Ilieracitim, Pflanzenreich IV. 
280. 1-864. 18 O 192 1. 

Zavitz, E. J. Reforestation in Ontario. Jour. For. 20: 18-24. 

Ja 1922. 



Zeller, S. M.,'& Owens, C. E. European canker on the Pacific 

slope. Phytopathology 11: 464-468. /. i-j. 25 F 1922. 

Zeman, V. Los hongos de la "Fhalaris btdbosa." Revista 

Facult. Agron. 14: 179-184. 1921. 

Zeman, V. Bacteriosis del bananero (una nueva enfermedad). 
Revista Facult. Agron. Univ. Nac. La Plata 14: 17-30. 1921. 


Zundel. G. L. The efifects of treatment for bunt on the ger- 
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21 Mr 1922. 

Vol. 49 

No. 11 





Effect of external and internal factors on the germination 

of fungous spores* 

Wm, L- Doran 


(with two text figures) 

k J 

The greater part of the work 'which has been clone on th^ 
germination of fungous spores has been incidental to studies of 
the life histories of fungi. References to spore germination are 
in, mosi cases isolated and few attempts have been made to 
draw any general conclusions from the results obtained. Since 
all fungous diseases must begin with infection and since the 
germination of the spore is the forerunner of infection, further 
knowledge of the conditions which favor or inhibit spore germina- 
tion is of great practical importance. In this paper are des- 
cribed the results of the writer's study of the effect of various 
factors on the germination of fungous spores. 

It is becoming increasingly common for field studies of the 
efficiency of fungicides to be preceded by laboratory tests of 
the toxicity of fungicides to fungi. If fungous spores are to be 
germinated both in the presence and absence of fungicides, it 
is important that all other conditions be identical and at or near 
the optimum. It is therefore necessary that optimum conditions 
for the germination of the spores be known before the laboratory 
tests of fungicides are made. 

Frequent references are made in the literature to the effect of 
prevailing climatic conditions on the outbreak or severity of 

AcknowledgemenL — ^The author gratefully acknowledges the assistance 
and encouragement received from Dr. O, R. Butler in this work. 

[The Bulletin for October (49: 283-312. pi. 12-17) %'as issued October 

n, 1922.1 

314 DoRAx: Ox the germixatiox of fuxgous spores 

epiphytotics of fungous originr Field experiments and observa- 
tions on the effect of weather on diseases of plants are affected 


by a very complex set of factors. Conclusions cannot safely 
be drawn until the various factors have been separately studied 
and such study is more easily effected in the laboratory than 
in the field. 

Factors influencing the germination of fungous spores may be 
divided into two groups, internal and external. The internal 
factors are maturity of the spore, longevity of the spore, anima- 
tion, and a poorly understood factor which may be called vitality 
of the spore. The external factors are oxygen, presence of 
moisture either in the form of water vapor or water of conden- 
sation (rain or dew^), temperature, light, nutrient substances, 
toxic substances, and duration of the time period during which 
one or more than one of the factors acting jointly exert their 
influence on the spore of ihe fungus. 


The germination of the following fungous spores was studied: 

-conidia of 


-tin 2 a f ruck' gen a (Pers.) Schrl., spores of Alternaria Solani (E. 
& M,) Jones & Grout., spores of Botrytis cinerea Pers., spores of 
Rhizoi us nigricans Ehr., aeciospores and urediniospores of 
Cronarlium rihicola F. de Wal., aeciosopres of Gymnosporangium 
clavipes C. & P., and teliospores of Piiccinia Malvaceartim Mont. 
The spores of the fungi were obtained fresh from the living host 
in all cases except Alternaria Solani. Spores of parasitic fungi 
to be used for germination tests were found to be more depend- 

r r 

%ble when obtained from this source than when obtained from 
fungi growling on artificial media. A spore obtained from the 
living host is presumably possessed of its full natural vigor. 
A spore obtained from a fungus growing on artificial media may 
have been subjected to a debilitating influence, or may, on the 
other hand, have become unnaturally resistant to adverse con- 
ditions. In this connection it should be noted that Young and 
Cooper (1) found that when ViomereUa rufomaculans was grown 
on agar eight to fifteen days, The spores were more resistant 
to the toxic action of fungicides than w^hen the fungus had been 
grown on agar only four to eight days. If fresh spores from the 
living host are not obtainable and spores from artificial culture 
must be used, it is the experience of the writer that all spores 

DoRAN: On the germination 


the germination of which is to be studied should be of the same 
age. The germination of spores from a second culture transferred 
from the first should not be compared with the germination of 
spores from the first culture. 

Distilled water was the medium in which the spores were 
placed to germinate. As is subsequently described more in 
detail, it was soon evident that the distilled water contained an 
insufficient amount of air for spore germination. Consequently, 
the distilled water was in all cases aerated before being used. 
The aeration was accomplished by drawing air through the 
water for about thirty minutes by means of an aspirator and an 
Allihn gas washing bottle- 
On culture plate benches in moist chambers were placed 
glass slides and on them drops of water either containing spores 
m suspension or the spores were shaken off into the drops. It 
is important that all drops of water be approximately equal in 
size especially when the toxic action of fungicides is to be 
tested. Clark (2) studied spore fermination in hanging cultures. 
Duggar (3) used that method also and a modification of it, 
placing the cells In small Petri dishes. Mains (4) germinated 
fungous spores in hanging drops on the cover of a Petri dish. 
When the cardinal temoeratures for the termination of the 




temperature. Constant temperatures were secured by the use 
of a DeKhotinsky electric oven and a Hearson incubator. 

The next point determined was the minimum length of time 
in which all the viable spores of each fungus will germiiate 
at the optimum temperature. At least six hours more than this 
minimum time period was allowed to elapse before removing 
the slides from the moist chambers a id determining the relative 
number of spores germinated. If slightly less than sufficient 
time is allowed a narrower range of optimum temperatures 
will be found than if more time is allowed, resulting, when the 
results are plotted, in a temperature-germination curve more 
acute than flat at its summit. If too long a time passes before 


the germinating spores are counted, the germ tubes will have 
grown and branched so as to make counting difficult and un- 
certain. All data are based on relative number of spores germi- 

3i6 Doran: On the geriviination of fungous spores 

The spores were counted by the use of a micrometer disc 
ruled in squares and the help of a tallying register. The micro- 
meter disc ruled in squares is placed in the ocular, the slide 
bearing the drops of water containing spores is placed on the stage 
of the microscope and moved by means of a mechanical stage 
until the outer line on the micrometer disc appears as a tangent 
to the circumference of the drop. 'The germinating spores 
appearing between the first two lines of the micrometer disc 
are counted, and then the ungerminated spores in this area are 
counted. When one field as limited by the lines on the micro- 
meter disc is counted, the slide is moved enough to bring a new 
field into view for counting. This is continued until a total of 
fifteen hundred to two thousand spores, both germinated and 
ungerminated, have been counted. This necessitates the ex- 
amination of several or many drops. From the count obtained, 
the relative number of spores germinating is figured, germina- 


tion elsewhere being raised proportionately. This accuracy of 
counting is most necessary when the relation between tempera- 
ture and spore germination is being determined. It is not as 
necessary in the case of toxicity tests with fungicides except 
when the differences in toxicity are small. 

Relation of the viability of the spore to its age 

Maturity. — Until the spore has attained a certain age or 
degree of maturity, it cannot germinate. Even before a spore is 
really mature, it can germinate if other conditions are near 
enough to the optimum. For example, the range between the 
minimum and maximum temperature is considerably greater 
for the germination of a mature urediniospore of Puccinia 
Antirrhini than for the germination of an immature uredinio- 
spore of this fungus, collected the first day it breaks through the 
host surface- The immature spore may be prevented from 
germinating by the action of fungicides not of sufficient con- 
centration to prevent the germination of the mature spore. 
A solution containing 0.257 per cent copper sulphate prevents 
the germination of mature spores of Uromyces caryophyllinus. 
A solution containing only half as much copper sulphate prevents 
the germination of immature spores of this fungus. It should 
be understood that mature and immature spores may not 
differ morphologically. Spores here referred to as immature are 
so called because when collected young they did not germinate, 

Doran: Ox the germination of fungous spores 317 


although older spores from the same fungus did germinate. AH 
conclusions as to germination should be based upon the behavior 
of mature spores. 

It is possible by exercising care in obtaining spores to separate 
the mature from the immature to some extent at least. When a 
fungus has matured its spores, they are usually free'd in such 
a way that they may be disseminated by the movement of air 
or water. So far as possible, the natural method should be 
duplicated in obtaining spores for experimental purposes. WTien 
a scab lesion on the fruit or leaf of the apple is washed, it is 
probable that only mature conidia and perhaps a few dead 
conidia of Venturia inaegttalis are detached. But when a similar 
lesion on the fruit or leaf of the apple is brushed or scraped, it 
is probable that many immature conidia are also detached. In 
the case of all except the fungi having very short-lived spores 
the immature spores on a fresh lesion greatly outnumbered the 
dead spores. When the writer obtained conidia of Venturia 
inaegtialis by brushing apple scab lesions the relative number of 
conidia which were capable of germinating was only twenty-five 
as compared with the relative number of one hundred germinat- 
ing when the conidia were obtained by washing lesions with a 
stream of water from a pipette. When fungous spores are to be 
obtained for germination studies they should be freed from the ' 
host as gently as possible and in as nearly as possible the same 
way in which they would be removed in nature. In this connec- 
tion it is interesting to note that Jones (5) found that the 
ascospores of Pseudopeziza Trifolii and P. Medicaginis germinate 
readily when they are discharged naturally but not when they 
are crushed out of the ascus. 

Some attention has been given to the later maturing of spores 
which were detached from the fungus while still immature. 
Schaffnit (6) is of the opinion that unless fungous spores are 
Internally mature before they are detached from the fungus, 
they never gain power to germinate. But the results of Melhus 
and Durrell (7) indicate that some urediniospores can mature 
after being detached from the fungus. 

With the exception of the spores of Bolrytis cinerea, all the 
fungous spores studied by the writer were apparently killed if 

detached when immature. 

It is possible in some cases to distingiush between mature and 
immature spores by their relative position in the fruiting parts 

3l8 DORAN 

of the fungus. The only spores of Sphaerotheca mors-uvae which 
Foreman (8) induced to germinate were the large spores found 
at the tips of the chains of spores. Weimer (9) observed that 
the teliospores of Gymnosporangium Juniperi-virginianae which 
are situated at the apex of the spore horn germinate best and he 
attributed this to their maturing earlier. 

As the season advanced in the autumn the writer found it 
increasingly difficult to secure good germination of the conidia 
of Ventiiria inaequalis even when conidia were obtained from 
fresh lesions. This is not attributed to the presence of a greater 
number of spores too old to germinate, for they would probably 
have been washed away earlier. It is believed that a larger 
number of immature conidia of this fungus were present in the 
autumn. It is not unHkely that the process of maturing of the 
spore is slower when the temperature is lower. 

Very little work has been done on the relation of the viability 

of fungous spores to the conditions under which the host plant 

grew and in which the fungus developed. If certain conditions 

may retard the maturing of the spores, it is quite possible that 

other conditions during their development may entirely prevent 

their ever germinating. On the other hand, certain conditions 

during development of the spore may result in an extremely 

vigorous spore. Spores may be mistakenly designated as mature 

and Immature, on the basis of their behavior when placed under 

optimum conditions for germination, when they should properly 

be designated as spores of the same age, some vigorous and others 
without vigor. 

The Inability to germinate any fungous spore under all possible 
combinations of conditions may be temporarily explained by the 
supposition that the spores are either dead, non-fu ictioning or 
In a resting condition. There are perhaps many resting spores 
not now recognized as such. Reed and Crablll (10) have ad- 

the theory that a rest period is necessary 
)f Gymnosporangium Juniperi-virginianae t 



Longevity. — For all spores there is a 
variable with the environmental conditions, beyond which the 
spore cannot germinate. The spore may be alive and too old to 
germinate or it may be dead. A live spore too old to germinate 
under the range of conditions within which it formerly could 
germinate Is, according to the observation of the writer, able 



to germinate when all conditions approach the optimum. There 
comes a time when even this is not stimulative and the spore is 
then dead so far as we can telL Neglect of the age factor may 
render any study of spore germination uridependable. 

In studying the relation of the age of spores to their viability, 
points to be considered are the date of collection, the conditions 
in which the spores are placed after collecting, and the length 
of time spores remain viable after their collection, Aeciospores 
of Cronartium ribicola were found to germinate with diminishing 
vigor as the season advanced. The aeciospores of this fungus 
were collected on May 25th, June 4th and June 12th from the 
same sorus. On the first date, the relative number of spores 
germinating was one hundred, on the second date forty-nine 
and on the third date twenty-nine. Observations with this 
and other fungi indicate that although there is at first a sharp 

falling-off in viability, it is retained for a comparatively long 
time and only gradually lost- In any spore germination studies 
which involve comparisons, spores should be used which are 
known to be of the same age. 

Among the factors bearing upon the retention of viability 
after the collection of spores are moisture, temperature, and 
freedom from or connection with the parent. Detachment 
from the parent was not found to have any effect on the length 
of life of the aeciospores of Cronartium ribicola, viability de- 
creasing the same in both cases. When collected, the relative 
number of these spores germinating was one hundred, after 
having been collected thirty days the germination fell to seventy- 
three, after fifty days to five, and after fifty-five days no spores 


Freshly collected aeciospores of Cronartium ribicola were 

stored indoors at temperatures of 7"" C, 15° C, and 23'' C. These 
various temperatures had no effect upon the longevity of the 
spores. But aeciospores stored in moist air retained their 
-viability longer than those stored in dryer air. At least in the 
case of this fungus moisture is of more importance than tempera- 
ture in its effect on the length of life of the spore. This effect of 

moisture is 


— ^^ -w^ ^ ^ -74. 

moist air, and if other conditions are equal they should live 
longer in moist air than in dry air. These results agree with 
those of Anderson (11) who found that in a humid atmosphere 

320 DORAN 

the spores of Cylindrodadium scoparium live for several weeks 
but that in a dry atmosphere they die in fifteen da\'S. 

Under no conditions did aectospores of Cronartiiim ribicola 
germinate when more than eight weeks old. A few cases are 
recorded in the literature of aeciospores of this fungus living 
longer, but it is probable that in such cases germination is reduced 
to a trace and that even that small germination occurs only 


for a time. 

The conidia of Venturia inaequalis produced on the leaves 
are somewhat shorter lived than those produced on the fruit. 
According to Aderhold (12) the conidia of this fungus retain 
their viability not more than eight weeks. The writer found 
none of the conidia of this fungus to live more than six weeks on 
the fruit and not more than four weeks on the leaf. With the 
aging of the leaves in the fall, the life of the conidia becomes even 
shorter. Difficulty was experienced in germinating conidia 
from fruit in storage. The scab lesions, however, were con- 
taminated with CephaJothecium rosennij the presence ^f the 
conidia of which in the drops of water may bave been partly 
responsible for the failure of the conidia of Venturia inaequalis 
to germinate. As will be described later, the presence in a drop 
of water of the spores of several fungi prevents or retards the 
germination of all the spores as compared with germination in 
a drop of water containing only the spores of one fungus. 

One of the factors bearing upon the retention of viability by 
spores after their collection is their freedom from or connection 
with the fungus or each other. Anderson and Rankin (13) 
found that ascospores of Entothia parasitica which remained in 
the perithecia in the bark germinated after being stored dry for 
one year. But when ascospores were removed and separated 
for only five months they lost the power to germinate. According 
to Burrill (14) the spores of Glomerella rufomaculans remain 
viable for a long time when their soluble. protective coating in 
which they are imbedded is present. But if it is washed away, 
the spores soon lose the power of germination. Anderson and 
Rankin (13) found that if the pycnospores of Entothia parasitica 
are stored in a dry place they live at least one year. But if the 
pycnospore horns are placed in water and the water allowed to 
evaporate so as to leave the spores separated, they do not retain 
their viability more than one month. 

Doran: On the germination of fungous spores ^21 


of the spore. For th 




gether in their protective coatings be used. 

Exposure to light shortens the life of some spores, as shown 

by the results of Hoerner (15) with the uredlniospores of Piiccinia 

Some attention has been given to the longevity of the spores 
of the Uredinales. The sporidia are the shortest-lived spores of 
this group of fungi. According to Reed and Craybill (10) the 
sporidia of Gymnosporangium Jimiperi-virginianae do not live 

than six days m dry air. Spaulding (16) reports that the 
sporidia of Cronanium ribicola live less than ten minutes at 
room temperature with the humidity at ninety. Duff (17) 
found that two weeks after the collection of the urediniospores of 
Cronartmm ribicola their germination was so reduced as to be- 
come practically negligible. The life of a teliospore measured 
from the time of its formation to its death may be long, but the 
life of a teliospore if measured from the close of its normal resting 
period until its death is not loTig; it is apparently as short in 
some cases as the life of many non-resting spores. Melhus, 
Durrell, and Kirby (18) report that the teliospores of Puccinia 
gramiiiis .v^hiclwusXure in April, cannot germinate after the end of 
June following. 

The experience of the writer is to the effect that in most cases 
aeciospores are longer lived than urediniospores. The average 
life of urediniospores is between thirty and sixty days. The 
average life of the aeciospore is about 50 per cent greater. The 
several spore forms of the Uredinales behave alike in this, that 
as they grow older their range between maximum and minimum 
conditions for germination becomes narrow^er. 

External factors 

Temperature relation. — One of the most important external 
factors bearing upon the germination of fungous spores is the 
temperature relation. In order to draw. from the scattered data 
in the literature some more general conclusions than we now have 
as to the cardinal temperatures for the germination of fungous 
spores, Table I has been prepared. In it are given the cardinal 

322 Doran; On the germination of fungous spores 

temperatures and the range of temperature for germination of 
thirty fungous spores. 

If the optimum temperature for the germination of the spores 
of Plasmodiophora Brassicae, which Chupp (19) found to be 
27°-3o'' C, is typical for the Myxomycetes, then the temperature 
requirements for the germination of the spores of the slime moulds 
must be higher than those of fungi. 

A study of Table I shows for the germination of the spores of 
several groups of fungi the following average cardinal temperatures 
and average range between minimum and maximum temperatures. 
For the germination of the Phycomycetes the average cardinal 
temperatures are 1.2'' C, iS'' C, and 26.1° C, and the range is 
25,0 degrees Centigrade. For the germination of the aeciospores 
of the Uredinales, the average cardinal temperatures are 6.5 
C, 13.0° C. and 22.0"^ C. and the average range is 15.5 degrees 
Centigrade. The average cardinal temperatures for the ger- 
mination of the urediniospores are 5° C, 16.2*^ C, and 28.9^ C, 
and the range through which these spores can germinate is 
23.9 degrees Centigrade. In the case of teliospores the cardinal 
temperatures for germination are 74° C; 19.9° C, and 28.1° C, 
and the range for germination is 20.7 degrees Centigrade. 

Temperature limits are narrower for the production of sporidia 
by germinating teliospores than for the production of germ tubes 
only. This is illustrated by the results obtained by the writer 
with the teliospores of Pticcinia Malvacearum, At temperatures 
as low as 5° C. promycelia are produced but no sporidia. Above 
23"^ C. promycelia are produced but sporidia are rare. 

These results agree with those of Dietel (29) who found that 
above 23*^ C. the teliospores of Pticcinia graminis produce no 
sporidia but only germ tubes. Reed and Craybill (10) noticed 
that the teliospores of Qymno sporangium Juniperi-virginianae 
do not germinate when the temperature rises to 24° C. But 
above that temperature they sometimes develop promycelia. 

When the Fungi Imperfecti and the one representative of the 
imperfect stage of the Ascomycetes named in TabIe I are con- 
sidered collectively, it is seen that for the germination of these 
six>res the cardinal temperatures are 7.3^ C, 23.1^ C, and 
39.6° C. Between the minimum and maximum temperature 
limits for germination there is an average range of 32.3 degrees 




Cardinal temperatures and range of temperature for the germination 

of the spores ol representative fungi. 














Cardinal Temperatures 

Plasmodiophora Brassicae 

Plasmopora viticola, conidia . . . . 
P. viticola, conidia 


Cystopiis candidus, conidia 

Phytophlhora infestans, conidia 
Peronospora parasitica, conidia 
Gymnosporangium clavipes, aeci- 


Cronartium rihicola, aeclospores 
Puccinia Phlei-pratensis, ure- 


2°-3° C. 


P. coronata, uredlniospores . . . 

P. coronata, uredinlospores 

P. rubtgo-vera, urediniospores . . 

P. graminis, urediniospores 

P. dispersa, urediniospores . . . . 

P. Antirrhini, urediniospores. . . 

P. Sorghi, urediniospores 

Cronartium rihicola, uredinio- 

Uromyces caryophyllinus, ure- 


27°-30° C. 

25^-30° C. 

25°-35" C. 

10° C. 25° C. 



14° C. 
12° C. 


25° c 
19° c 

18° c. 
18° c. 


10°-12'' C 


30° C. 

30° c. 

30° C. 


31° c. 

Uromyces Trijolii, uredinio- 

Gymnosporangium Juniperi-vir- 



ginianae, teliospores 

G. Juniperi-virginianae, telio- 

i8°-20° C. 
10° C. 

14° c. 

14° C. 

14° C. 

16° C. 

25°-27° C 
20° C. 

25° c. 

25° c 

29° C 

34° C. 

11° C. 




Puccinia graminis, teliospores.. . 

P, graminis^ teliospores 

P, Lariciy teliospores , . . . . 

P. Malvacearum, teliospores. . . . 

P. graminis^ basidiospores 

Colletotrichum lagenarium 

Melanconium sp * 

Phyllosticta Antirrhini 

P, Antirrhini 18° C- 

Alternaria Solani 

15° c. 

23°-24° C. 29° C. 

29" C. 

22° C. 
20° C. 


14° c. 

i5°-20° C. 

22°-27° C. 

23°-27° C. 

25° c. 

23° c. 
25° c. 

30° C 

Septoria sp 

47° C. 
-3°C. 26°-28°C.'37°-45°C. 

24''-28'' C. 34° C. 













Numbers refer to the literature cited. 


DoRAX: On the germination of fungous spores 

For these several groups of fungi the minimum temperatures 
for spore germination occur between i° and 74*^ C. The opti- 
mum temperatures are all between 13° and 23.1° C, The maxi- 
mum temperatures for the germination of the spores occur 
between 22° and 39.6° C. The Phycomycetes have the lowest 
minimum temperatures, followed in order by the urediniospores, 
the aeciospores and the teliospores of the Uredinales, Teliospores 
and the spores of the Fungi Imperfecti have the highest, and about 
the same minimum temperature for germination. 

Of these five groups of fungous spores, aeciospores have the 
lowest optimum temperature for germination followed in turn 


16.2 18 19.9 23.1 









1 2 3 4 5 6.5 7 A 



Fig. X. Curv^es showing the cardinal temperatures for the germination 
of the spores of representative fungi. 

by urediniospores, conidia of Phycomycetes, teliospores, and 
then by the spores of the Fungi Imperfecti, which seem to have 
the highest optimum temperature for germination. 

Aeciospores have the lowest maximum temperature for 
germination, and the spores of the imperfect fungi have the 
highest. About midway between these two come urediniospores. 
Their maximum temperature for germination is higher than that 
of the conidia of the Phycomycetes. 

The spores of the Fungi Imperfecti can, it would appear, 

the greatest 

As com- 

pared with this, the range of the Phycomycetes is 77 per cent 

DoRAN: On the germination of fungous spores 325 

as great, that of the urediniospores is 74 per cent as great, that 
of the teliospores is 64 per cent as great, and the range of the 
aeciospores is only 48 per cent. 

- Fig. I shows the ranges and the cardinal temperatures for 
these groups, and their relative positions. 

The results obtained by the writer and others indicate that any 
departure from optimal conditions tends to narrow the tempera- 
ture limits within which fungous spores will germinate. The 
literature contains a very few references to the relation which 









12. 3 4 5 6 7. 8 9 JO II U"T3 14 15 \6 17 18 19 20 21 22 23 24 25 26 27 28 29 30 il 32, 

Fig. 2. Curve showing temperature limits for the germination of the 
conidia of Venturia inaeqtialis, 

exists between the temperatures required for the process of spore 
germination and the other environmental conditions prevailing 
during that period. Gardner (30) found that the minimum 
temperature for the germination of the spores of Colletotrichum 
lagenarium is 7° C. when the spores are in exposed drops of water, 
but it is 14° C. when the spores are in hanging drops of water. 
When these spores are In exposed drops of prune decoction, the 

temperature for their germination Is 4° C. It would 
appear that the presence of nutrient substance and the availa- 
bility of sufficient ox>'gen both tend to lower the minimum 


The work of the writer included the determination of the 
cardinal temperatures for the germination of the conidia of 
Venturia inaequalis, the spores of Botrytis cinerea, and those of 



DoRAN: On the germination of fungous spores 

Rhizopus nigricans. Cardinal temperatures for the germination 
of the other spores used had been previously determined (Doran, 


The results obtained with the conidia of Venturia inaequalis are 

shown in Table II and by the curve in Fig. 2. For the germina- 
lion of the conidia of Venturia inaequalis^ the minimum tem- 
perature is 3° C, the optimum temperature is I4°-I5° C, and 
the maximum temperature is 31^ C. The conidia can germinate 
through a range of twenty-eight degrees. 

This is an unusually long range. Since the causal fungus of 
apple scab is not especially exacting as regards temperature for 


the germination of the conidia the indications are that the 
distribution or severity of this disease, following the primary 
infection by the ascospores is dependent more upon rainfall 
than upon temperature. The conidia of Venturia inaequalis are 
disseminated most freely during June, July and August. As 
shown in FiG. 2, warm nights do not prevent their germination. 
A study of the temperature-germination curve in FiG. 2 shows 
that the falling off In germination of the conidia Is more rapid 
from the optimum to the minimum temperature than from the 
optimum to the maximum temperature. Over 40 per cent of the 
number of spores germinating at the optimum (i4°-i5^ C.) 
germinate at 9° C. and 24° C, respectively. 


Effect of temperature on the germination of the conidia of 

Venturia inaequalis 


Relative germination at different temperatures measured in degrees Centi- 





















20 52 
































































II 1 

2 1 


perature, as to all other external conditions, the relative number 
of spores germinating was taken as an indicator of the reaction 




of temperature. Johnson (26) and Melhus and Durrell (7) 
preferred to use the growth or length of the germ tubes as an 
indicator of the temperature reaction. To test the relative 
merits of these indicators, both were used by the writer in the 
case of germinating conidia of Venturia inaequalis. When the 
optimum temperature for germination was based upon length 
of germ tube rather than upon number of spores germinating 
the optimum was less sharply defined. The writer believes that 
to consider length of germ tube rather than number of spores 
germinating involves growth rather than germination and so 
should be studied separately, for there is no reason to believe 
that the optimum temperature for the growth of a fungus is the 
same as the optimum temperatures for spore germination. 
Melhus (22) found the optimum temperature for the germination 
of the conidia of PhytopMiora infestans to be I2°-I3'^ C. Jones, 
Giddings and Lutman (30) found that the optimum temperature 
for the growth of this fungus is between 16° and 19° C. The 
cardinal temperatures for the germination of the conidia of 
Botrytis cinerea were found to be minimum 7° C. to 17"^ C, and 
maximum 26"^ C. For the germination of the spores of Rhizopus 
nigricans, the minimum temperature was found to be lo"" C. 
and the optimum 19° C. to 20° C- All later studies of tlie germ- 
ination of Botrytis cinerea were made at 15° C. and of the germ- 
mation of Rhizopus nigricans at 19° C. 

Time relation, — The length of time required for the process 
of spore germination Is In itself an important factor. If before 
the elapse of the necessary time period the environmental 
conditions depart from the optimum, the germination process is 
impeded or stopped and the spore may even be killed. In all 
laboratory studies of spore germination, if conditions in nature 
are to be duplicated, no more time need be allowed for germina- 
tion than that in which nearly optimal conditions are likely to 


continue in nature. Probably most fungous spores begin and 
complete germination between darkness and daybreak. One of 
the principal natural protections of plants from pathogenic 
fungi is the failure of environmental conditions to remain be- 
tween minimum and maximum limits long enough for spore 
germination. Infection does not occur if during the protrusion 
and growth of the germ tube the necessary precipitated moisture 
evaporates or the air falls below a certain degree of humidity 
or the temperature passes beyond minimum or maximum limits. 

328 Doran: On the gerjviination of fungous spores. 

According to the observation of the writer, extremes of tempera- 
ture at this time, if not too long continued, have no more serious 


This is resumed when the temperature again approaches the 
optimum. But when the humidity of the air falls, the results 
to the spore are more serious. In the case of all spores worked 
with, and especially with conidia of Venttiria inaequaUs, it was 
found that a germinating spore can live but a short time in a dry 
environment, which to an ungerminated spore would be in no 
way detrimental. A return to optimal conditions proved such 
spores to be not merely inhibited but dead. 


ermmation an im 


by Venturia inaeqiialis . He found that the ascospores of this 


trees must remain wet eight or ten hours to be abundantly in- 



temperature is from the optimum for germination, the longer must 
the surface of any plant remain w^et in order for the spores on 
it to germinate and Infect the host. 

Duggar (3) found that after drying the germinating spores 
of Botrytis would grow no more. But the germinating spores of 
Aspergillus showed new growth after being dried several days. 
Duggar 's results with the parasitic form mentioned are in agree- 
ment with the results secured by the writer, who worked only 
with parasitic forms. 

The relation of environmental factors to the time required 
for spore germination has occasionally been noted. The amount 
of moisture present may affect tlie length of time in which the 
spore germinates. Taubenhaus (38) reports that the spores of 
Glomerella nijomaciilans from the sweet pea germinate In from 
six to twenty-four hours, depending on whether there is much 
or little moisture present in the atmosphere. 

The writer has noticed that spores germinating in moist air 
require a relatively longer time for the process than do the 
same spores germinating in or on precipitated moistqre. 

Anderson (11) observed that at 25^ C, which Is within optimal 
limits for the germination of the spores of Cylindrocladium 
scoparium, the spores of this fungus begin to germinate in two 
to three hours, but at lower temperatures a longer time is neces- 

Doran; On the germination of fungous spores 329 

sary for their germination, five hours at 12'' C. and twenty-four 
hours at 8^ C. Tisdale (39) found that at 4° C, which is their 
minimum temperature for germination, the spores of Didymellina 
iridis germinate in twenty-one hours, but at their optimum 
temperature of 20° C. to 26° G. these spores germinate in two 
and one half hours, and at their maximum temperature which 
is 30^ C, their germination requires eleven hours. 

The presence of a nutrient solution may hasten the germina- 
tion of some spores. Gardner (30) found that the spores of 
CoUetotrichtim lagenarium germinate in five hours in agar but 
require ten to twenty-four hours for their germination in dis- 
tilled water at the same temperature. 

It appears that immature spores and aged spores may germin- 
ate more slowly than mature spores. Melhus, Durrell, and 
Kirby (18) found that both before and after the teliospores of 
Puccinia graminis have completed their rest period, the time 
required for their germination is longer than that required just 
as they end their rest period in the spring. 

The results mentioned above indicate that all fungous spores 
germinate most rapidly when all environmental conditions are 
nearest to the optimum. 

Table III shows the time necessar^^ for the germination of 
twenty-three representative fungous spores. After a study of 
this table it is possible to draw certain conclusions as follows: 
The average time required for the germination of fungous spores 
is about twelve hours ; for the germination (indirect) of the conidia 
of the Phycomycetes it is four hours; for the ascospores of the 
Ascomycetes it is thirteen hours; for the germination of the 
chlamydospores of the Ustilaginales it is twenty-one hours; for 
the germination of the aeciospores of the Uredinales it is ten 
hours; for the germinations of the urediniospores of the Uredin- 
ales it is five hours; for the germination of the teliospores of the 





that the lenerth of time necessarv 

germination of the fungous spores named in Table IV is: aecio- 
spores of Cronartium ribicola twelve hours; urediniospores of 
Cronartium ribicola five and one-half hours; for the production 
of basidia by the teliospores oi Puccinia Malvacearum two hours; 
one hour more, a total of three hours are necessary for the 


Doran: On the germination of fungous spores 


production of sporidia by the teliospores of this fungus; aecio- 
spores of Gymno sporangium clavipes require four hours; conidia 
of Venturia inaegiialis about twenty-four hours; and conidia of 
Sclerotinia frucHgena five hours. These tests were all made at 



Length of time required for the germination of the spores of 

representative fungi 







Phytophthora infestans, conidia 
Cystopiis candidns, conidia. . . . 

Plasmopara viticola, conidia. . . . 
Venturia inaequalis, ascospores. 
Gtdgnardia Bid-wellii, ascospores 
Endothia parasitica, ascospores. . 
Didymellina Iridis, pycnospores. 

Hours necessary 
for germination 


Pseudopeziza Trifolii, ascospores 
Diplocarpon Rosae, conidia 

Glomerella rufomaculans, conidia 
Tilletia foetens , chlamydospores , 

Ustilago Hordei, chlamydospores 

Ustilago Trittcij chlamydospores 

Cronartium ribicola, aeclospores 

Puccinia Anlirrhiniy urediniospores 

P. Phlei-pralensiSf urediniospores 

P. Malvacearum, teliospores 

Gymno sporangium Juniper i-virginianaCy telio- 


Colletolrichum lagenarium 
C, Schizanthi 

Phyllosticta A ntirrhini 
P. Antirrhini 

Cylindrodadium scoparium 
Septoria Gladioli , 




















* Numbers refer to the literature cited 



tests were made at temperatures approaching the minimum 
and maximum temperatures for the germination of the spores of 
these fungi. Under these conditions, in every case, a longer 
time -was required for germination than at optimum temperatures. 
Oxygen relation. — It was observed that spores in the interior 


DoRAN: On the gerimin 


the surface. The conldia of Venturia inaequalis germinated 
only when they were on or near the surface of the water. Tclio- 
spores of Pticcinia Malvaceamm did not produce sporidia when 
deeply submerged in the drop. But when distilled water was 
used, which had previously been aerated, equally good germina- 
tion was secured whether or not the spores were submerged 
in the drops. 

DeBary (47) described the behavior of spores in water be- 
tween a cover glass and a glass slide. In this case^ the spores 


Time period necessary for the germination of representative 

fungous spores under" 


optimum conditions 

Fungus and spores 

No. of hours 

Spores germinating (relative numbers) 

Cronartium ribicola^ 

10 . 

1 00 

Cronartium ribicola^ 



r ! 


GymnosporangkiM davipeSy 





Puccinia Malvacearum, 


Producing basidia 100. Producing 

Producing basidiospores loo 

Sclerotinia fructigena, 



Venturia inaequalisy 




near the periphery of the cover glass germinate better than those 
near the center due to the relative amounts of air available. 
Duggar (3) found that a reduced oxygen supply retards spore 
germination, Blackman (48) noticed that if the germ tube of 
Phragmidium violaceum does not grow through the water and 
so reach the air it develops abnormally. Weimer (9) observ^ed 

that when the teliospores of Gymnosp 
virsinianae are covered with water they proc 


instead of the normal promycelia and basidiospores. Melh 

332 Doran: On the germination 

and Durrcll (7) found that when the uredmiospores of Puccinia 
coronata are submerged only a small percentage of them germinate 
as compared with the number germinating when they float on 


a drop of water. 

When spore germination tests w^ere made in non-aerated 
distilled water, the writer secured best results by sowing the 
spores on the surface of the water rather than immersing them. 
Some spores, however, will not float. According to Duggar (3) 
the spores of many of the Phycomycetes and Hymenomycetes 
usually sink. The specific gravity of the spores of several 
Hymenomycetes determined by Buller (49) were between 1.02 
and 1. 21. Since some spores sink, all distilled water for germina- 

■ tion studies should be aerated. 

At this point reference should be made to the injurious effect 
of the presence of fungous spores on the germination of the spores, 
of other fungi. Although the injurious effect of this competition 
between spores might be attributed to the secretion of toxic 
substances, there is little likelihood that this is the case. There 
is an indication that it is due to there being insufficient oxygen 
for all. The effect is much more marked in non-aerated than in 
aerated distilled water and is more marked in the interior of a 
drop of non-aerated distilled water than on its' surface* All 
spores of Uredinales which the wTiter has observed germinate 





in a drop of w^ater with the teliospores of Pttccinia Malvacearum^ 
the spores of the first three fungi may germinate but the telio- 


spores of Pttccinia Mahacearum do not germinate, although 
check teliospores in uncontaminated drops germinate perfectly. 
When in place of Puccinia Malvacearwn the conidia of Venkirio- 
inaequalis were used a similar result was obtained; they germin* 
ated far better when no other spores were present with them in 
the drop of water. WTien there is lack of sufficient ox>^gen for 
all, only those spores which require the least will germinate. 
It w^as also observed that when relatively few spores of one 
species were present in a drop they germinated better than 
when the drop was crowded. This, too, may be attributed to 
insufficient oxj^gen. In all spore germination studies where 
optimum conditions are desired; an effort should be made to 
exclude from the culture drop the spores of all fungi except the 

Doran: On the germin 


one under consideration, and there should not be present an 
excessive number of spores of that one fungus. 

There are a few references which have come to the attention 
of the writer on the deleterious effect of competition or crowding 
on germinating spores. Edgerton (50) noticed that when more 
than twelve or fifteen spores of Colletotrichum Lindemuthiantim 
are present in one cubic millimeter of water, they germinate 
more poorly than when a smaller number of spores are present. 
Taubenhaus (51) found it advisable to wash hollyhock leaves 
in water to remove the spores of saprophytic fungi preparatory 
to studying the germination of the teliospores of Puccinia. 
Malvaceamm. He does not indicate in what manner the pres- 
ence of the spores of other fungi interfere with the germination 

Light relation, — Opinion in the literature is not unanimous 
as to the effect of light on spore germination. According to 
DeBary (52) and Farlow (53) light inhibits the germination 
of the spores of the Oomycetes. Cuboni (54) concluded that 
intense light interferes with the germination of the conidia of 
Plasmopara viticola. Ward (55) found that the spores of the 
brome rust germinate as readily in light as in darkness. Melhus 
(22) observed no difference in the percentage of conidia of 
Cysioptts canididus germinating, nor in the time required for 
their germination, whether germination took place in light or 
darkness. Melhus (23) found that light does not interfere 


with the germination of the conidia of Phytophthora infeslans 
if the optimum temperature for their germination is not exceeded. 
Duff (17) found that exposure to glass filtered sunlight is not 
injurious to the urediniospores of Cronartiitm ribicola, providing 
the temperature does not rise too high. He found that exposure 

to the ultra-violet rays from an electric arc completely inhibited 
spore germination. Lauritzen (56) concluded from his invest- 
igations that light is not a limited factor in the infection of 
plants by fungi. 

The writer tested the effect of sunlight on the germination of 
the spores of Alternaria Solani and the conidia of Sclerotinia 
fructigena. The spores of both of these fungi germinate quite as 
well in sunlight, whether it be direct, diffuse, glass filtered or 
not glass filtered, as they do in darkness, provided that the 
conditions of temperature and moisture meanwhile remain near 
the optimum. The indications are that the spores of fungi 

334 Doran: On the germination of fungous spores 

germinate in either light or darkness, when the other environ- 
mental conditions are between maximum and minimum limits. 


It is, however, probable that the germination of fungous spores 
in nature usually takes place in darkness, since it is in the night 
that conditions of temperatures and moisture nearer the opti- 
mum for germination usually prevail for a longer time. 

Water relation, — An examination of the literature indicates 
that the moisture requirement of germinating spores is not 
uniform. Tulasne (57) was able to germinate teliosp ores of 
the Uredinales as well in a saturated atmosphere as in a drop 
of water. Patrigeon (20) found that for the germination of the 
conidia of Plasmopara viticola precipitated moisture is necessary 
and merely damp air is insufficient. According to DeBary 
(47) the spores of the Uredinales germinate better when they are 
merely in a moist atmosphere than when they are in water. 
Lesage (58) found that the spores of Penicillium glaticum can 
germinate in damp air so long as the humidity does not fall 
below 82 or 84 per cent. Taubenhaus (51) readily germinated 
the teliospores of Pticcinia Malvacearum in a humid atmosphere. 
Levin (59) was able to infect tomatoes with Septoria Ly coper sici 
by merely applying dry spores. Weimer (9) found the telio- 
spores of Gymnosporanghint Juniper i-virginianae unable to 
germinate when they were in contact with no moisture except 
that in the atmosphere. These spores did not germinate until 
the air became super-saturated, so that small drops of water 



(11) states that the spores of Cylindrocladiiim scoparium never 
germinate except when they are directly in water, a moist 
atmosphere being insufficient. According to Melhus and Durrell 
(7) the urediniospores of Puccinia coronata germinate only when 
they are in direct contact with water. Smiley (33) found that 
in the absence of a film of water, the spores of PhyUosticta 
Antirrhini do not germinate. Melhus, Durrell and Kirby (18) 
found that the sporidia of Pticcinia graminis germinate pro- 
fusely in drops of water, and but very poorly when their only 
moisture is obtained from the atmosphere. 

The writer tested the effect of moist air as compared with 


Table V. 


minate perfectly in moist air, as they do in water. The spores of 
Alternaria Solani and the conidia of Venturiainaequalis germinate 


Dor.\n: On the germination of fungous spores 


only 4 percent in morst air as compared with loo percent 
in water. This means that in nature the relative number 
of spores of these two last named fungi germinating in moist 
air is very small, but undoubtedly sufficient to cause some in- 



in moist air but require precipitated moisture. The conidia of 
Peronospora pygmaea were also found to require precipitated 
moisture rather than moist air for their germination. 

Of the spores studied by the writer and mentioned as above in 
the literature, in only five cases can they germinate well when 
their only source of moisture is tlie atmosphere. Three germinate 

poorly in moist air. Six germinated only in precipitated 
moisture. The evidence is insufficient to warrant the conclusion 
that any group of fungi characteristically obtain their water for 
germination principally in the form of vapor. When optimuni 
conditions for laboratory experiments on spore germination are 



Effect of moist air as compared with precipitated moisture 

on the germination of fungous spores 

Spore germinating (relative numbers) 

Moisture conditions 

A . Solani 

V, inae- 

G. clavi- S. friicti- 
pes gena 

In a drop of distilled water 
On dry slide in moist chamber 








p. pyg- 


to be obtained, water in the form of precipitated moisture rather 
than water vapor should be supplied. The question of too much 
water is probably closely related with the question of too little 
osy^gen . 


I. The germination of the following spores was studied; 
conidia of Venturia inaegtialis (Cke.) Wint., conidia of Sclerotinia 



& Grout, spores of Botrytis cinerea Pers., spores of 



2. Spores of parasitic fungi germinate better when obtained 
from the living host than when obtained from artificial media. 

336 Doilvn: On the germin-atiox of fungous spores 

3. Mature spores can germinate through a broader range of 
environmental conditions than can immature spores. 

4. Freshly mature spores can germinate through a broader 
range of environmental conditions than can old spores. As the 
spores age, viability at first decreases sharply after which it is 
only gradually lost. Longevity of spores is dependent on condi- 
tions of storage after detachment from the host. Moisture is 
of more importance than temperature in its effect on the length 

„of life of the spore. 

5. Spores of the Phycomycetes can germinate at the lowest 
minimum temperatures, followed in order by the urediniospores, 
the aeciospores, and the tellospores of the Uredlnales. Aecio- 
spores have the lowest optimum temperatures for germination 
followed in order by urediniospores, conidia of Phycomycetes, 
teliospores, and the spores of the Imperfect Fungi. 

6. The nearer all conditions are to the optimum, the shorter 
the time required for spore germination. 

7. Competition or crowding inhibits spore germination. 
This IS attributed to Insufficient oxygen. 

8. Spores of the fungi studied germinated in either light or 



of some spores if water vapor is available - 

New Hampshire Agricxtltural Experiment S 
Durham, New Hampshire 

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51. Taubenhaus, Jacob J. A contribution to our knowledge of 

the morphology and life history of Puccinia Malvaceanim 
Mont. Phytopathology 1: 55-62. pL 12.-14. I9ii, 

52. DeBary, A, Recherches sur le developpement de quelques 

champignons parasites. Ann. Sci. Nat. Bot. IV. 20: 5- 
148. pL I~i3^ 1863. 

53. Farlow, W. G. The American grape vine mildew. Bull. 

Bussey Inst. Art. 1: 419. 1876. 

54. Cuboni, J. C. La Perojiospora dei grappoli. Atti Congr. 

Nazion. Bot. Critt. m Parma, 91-108. 1887. 


55. Ward, H. Marshall. On the relations between host and 

parasite in the bromes and their brown rust, Puccinia. 
dispersa Erikss. Ann. Bot. 16: 233-315. 1902. 

56. Lauritzen, J. L. Relations of temprature and humidity 

to infection by certain fungi. Phytopathology 9: 7-35. 


57. Tulasne, L. R. Second memoire sur les Uredin^es et les 

Ustilaginees. Ann, Sci. 77-196. pL^-12. 

58. Lesage, P. Recherches experimen tales sur la germination 

des spores du PenicilUum glancwn. Ann. Sci. Nat. Bot. 
8: 309-322. 1895. 

59. Levin, Ezra. The leaf spot disease of tomato. Michigan 

Agr. Exp. Sta. Bull. 25: 1-15, 1916. 




The aim of this Index is to include all current botanical literature written by 
Americans, published in America, or based upon American material; the word 
America being used in the broadest sense. 

Revie\^^, and papers that relate exclusively to foresty, agriculture, horti- 
culture, manufactured products of vegetable origin, or laboratory methods 
are not included, and no attempt is made to index the literature of bacteriology. 
An occasional exception is made in favor of some paper appearing in an Ameri- 
can periodical which is devoted wholly to botany. Reprints are not mentioned 
unless they differ from the original in some important particular. If users 
of the Index will call the attention of the editor to errors or omissions, their 
kindness will be appreciated. 

This Index is reprinted monthly on cards, and furnished in this form to 
subscribers at the rate of three cents for ea,ch card. Selections of cards are 
not permitted; each subscriber must take all cards published during the term 
of his subscription. Correspondence relating to the card issue should be 
addressed to the Treasurer of the Torrey Botanical Club. 

Anderson, R* J., & Kulp, W. L. Analysis and composition of 

corn pollen. J 

F 1922* 

Anderson, P. J. Development and pathogensis of the onion 

smut fungus. Massachusetts Agr, Exp. Sta. 4: 99-133./. 
i~3' N 1921. 

Andrews, A. L. Further bryophytes from North Carolina (and 

Tennessee). Bryologist 24: 49-58. 9 Ja 192 1. 

Andrews, A. L. Notes on North American Sphagnum, IX. 

Bryologist 24: 81-86. 27 F 1922. 

Armstrong, G. M. Studies in the physiology of the fungi. 

XIV. Sulphur nutrition: the use of thiosulphate as in- 
fluenced by hydrogen-ion concentration. Ann. Missouri 
Bot. Gard. 8: 237-281. f. 1-21. S 1921. 

Arndt, C. H. The growth of field corn as affected by iron and 

aluminum salts. Am. Jour. Bot 9: 47~7i- P^- 4+1- -^"<5- 
20 Mr. 1922. 


Arthur, J. C* Uredlnales collected by Fred J. Seaver in Trinidad. 

Mycologia 14: 12-24. 6 Mr 1922. 

Includes 5 new species. 


342 Index to American botanical literature 

A-fthur, J. C, Uredinales collected by R. Thaxter and J^ B. 

Rorer in Trinidad. Bot.Gaz. 73: 58-69, /. 1-4. i8Ja 1922. 

Ashe, W. W. The eastern shrubby species of Rohinia. Jour. 

Elisha Mitchell Sci. Soc. 37: 175-177. Mr 1922. 

JR. grandiflora sp. no v. 


Ashe, W. W. Forest types of the Appalachians and White Moun- 
tains. Jour. Elisha Mitchell Sci, Soc. 37: 183-198. Mr 1922. 

Avten, J. T, The organic phosphorus content of some Iowa 

soils. Soil Sci. 13: 1 19-124. F 1922. 

Awschalom, M. Datos sobre la influencia del selenio en la 

vegetaci6n cuando sustituye al ion sulfurico en el liquido 

nutritivo de Knop. Revista. Facult. Agron. 14: 122-162, 

Baas-Becking^ L. G. M. The origin of the vascular structure 

in the genus Botrichitim; with notes on the general anatomy. 

Rec. Trav. Bot, Neerlandais 18: 333-372. pL 5^6 -^ f, 1-49- 

Bailey, J. W. Adventures in mossland. — ^Ahunt foi Desmatodon 

latijolitis (Hedw.) Brid. Bryologist 24: 86-88. 27 F 1922. 


Bailey, L. W. Diatoms from the Quill Lakes, Saskatchewan, 

and from Airchie, Alberta. Contr. Canadian Biol, ii: 
157-165, pi. I, 2:. 1921. - 

Barker, E. E. The architecture of the coco palm. Nature- 
study Rev. 18: 44-50. /. 1-4. Ja-F 1922. 

Benoist, R. Contribution a I'etude de la flore des Guyanes. 

Bull. Soc. Bot. France 68: 311-323, 2 Au 192 1. 

Berry, E. W. Additional occurrences of Pleistocene plants* 

Torreya 22: 10, 11. F 1922. 

Berry, E. W. Carboniferous plants from Peri\. Am. Jour. 

Sci, 203: 189-194. Mr 1922. 

Berry, E. W. The flora of the Woodbine sand at Arthur's 

Bluff. Texas. U. S. Geol. Sui 

+ f. II. 23 Mr. 1922. 

Includes Trochodendr aides, gen, nov. 


Index to American botanical literature 343 



the Finger Lakes of New York. Bull U. S. Bur. Fisheries 
37; 209-252, 1922. 

Includes notes on phytoplankton. 

4 ■ 

Blake, S. F. New Plants from Guatemala and Honduras- 

Contrib. U. S. Nat. Herb. 24 : 1-32. pi i-io + /. 1-4. 1922. 

Includes 46 new species. 


Blake., S. F., & Sprague, T. A. On the generic name Wikstroemia. 

our. Bot. 60: 52-54. F 1922. 

Blakeslee, A, F. Variations in Datura due to changes in chro- 
mosome number. Am. Nat. 56: 16-31. /. j-7. F 1922. 


Blomquist, H. L. Vascular anatomy of Angiopteris evecta^ 

Bot. Gaz. 73: 181-199. pL 5-8 + /. 1-8. M 1922. 


Botto, A. Una nueva forrajera para las regiones del norte. — El 

pasto de Rhodes. Revista Faculty Agron. 14: 170-178. 
/. 7-j. 1921. 


{Chloris Guyana Kunth.) 

Bouyoucos, G. A study of the fertility of the soils of Greece. 

Soil Sci. 13: 63-79. F 1922. 

Brand, A. Borraginaceae-Borraginoideae Cynoglosseae. Pflan-^ 

zenreich IV. 252: 1-183. f. 1-22, 22 N 1921. 

Broadway, W. E. Botanical collecting in French Guiana. Jour. 

New York Bot. Gard. 22: 177-183. O 1921. 


Burns, G. P. Eccentric growth and the formation of redwood 

in the main stem of conifers. Vermont Agr. Exp. Sta. 
Bull- 219: 1-16. pL 1-4 + /. i-io. Je 1920. 

Butler, E. J. Phytopathology in the United States. Kew Bulk 

Misc. Inform. 1922: 85-91. 1922. 


Canela, P, F. Sobre la presencia de una plaga en los fru tales 

en San Luis, BoL Minis. Agric. de la Nacion 26: 253-256. 

Cannon, W. A. Plant habits and habitats in the arid portions of 

South Australia. Carnegie Inst. Washington Publ. 308: 

1-139. pi' i~3^ + /. J-J^- 1921. 

344 Index to American botanical literature 

Caverhill, P. C. Forestrj^ problems in British Columbia, Jour. 

For. 20: 44-53. Ja 1922. 


Chardon, C. E. Contribution to our knowledge of the Pyre- 
nomycetes of Porto Rico. Mycologia 13: 279-300. pL 13- 

15 + /' ^-4' 21 Ja 1922. 

Includes 9 spp. nov. and.i comb. nov. 

Churchill, J. R. Cimicifuga racemosa in Massachusetts. 

Rhodora 23: 201-203. 9 Ja 1922. 

Clements, F. E. Aeration and air-content. The role of oxygen 
in root activity. Carnegie Inst. Washington Publ. 315: 

I-I83. I92I. 

Clokey, I. W. Carex notes. Am. Jour. Sci. 3; 88-^91. pL 2. 
Ja 1922. 

Includes Carex apoda and C Paysonisj spp. nov. 

CoateSi L. The "peach-almond" hybrid. Jour. Hered. 12: 
328, 329. 20 F 1922. [Illust.] 

Cocks, R. S. A list of the trees of Louisiana. Jour. Arnold 
Arbor. 2: 204-216. Ja 1922. 

Cockerell, T. D. A. Flora of Porto Santo. Torreya 22: 4-10. 
/. J, 2 F 1922. 

Coker, W. C, & Grant, F. A. A new genus of water mold 
related to Blastocladia. Jour. Elisha Mitchell Sci. Soc. 
37: 180-182. plj2, Mr 1922. 

Septocladia gen. nov. and 5. dichotoma sp. nov. 


Collins, G. N. Teosinte in Mexico. Jour. Hered. 12: 339-350. 

/. J-/. O 192 1. 

Conti, M. El campo experimental de hidrologia agricola. 
Revista Facult. Agron. 14: 97-121. 1921. [Illust.] 

Cooper, W. S. The ecological life history of certain species of 
Ribes and its application to the control of the white pine 
blister rust. Ecology 3: 7-16. /. 1-3. 25 F 1921. 

Cundftll, F., & Fawcett, W. Dr. Anthony Robinson of Jamaica. 
Jour. Bot. 60: 49-52. F 1922. 

Davis, B. M. Species, pure and impure. Science 11. 55: 107- 

114. 3 F 1922. 

Index to American botanical literature 345 

Demerec, M* Heritable characters of maize. X. Zebra striped 

leaves. Jour. Hered. 20; 406, 407. /. 12^ N 192 1. 


Denison, I, A. The nature of ceitaiii aluminum salts In the 

soil and their influence on ammonification and nitrification. 

Soil Sci. 13: 81-106. F 1922.^ 


Denny, F. E. Formulas for calculating number of fruits re- 
quired for adequate sample for analysis, Bot. Gaz. 7^: 
44-57- 18 Ja 1922, 

Detwiler, S. B. Blister rust appears in the Puget Sound region. 

Am. For, 28: 97, 98. F 1922. 

Dietel, P. Zur Umgrenzung der Gattung Pileolaria Cast. Ann. 

Mycol. 19: 300-303. 1921. 

Dodge, C, K. Miscellaneous papers on the botany of Michigan. 


15-234. 1921 

Plant lists. of various sections of state. Includes biography, portrait 
and map. 

Dunbar, J. How to grow rhododendrons. Jour. New York 

Bot. Gard, 22; 184-190. O 1921. 


Durand,E. J. Thegenus Catinella. Bull. Torrey Club 49: 15-21- 

21 Mr 1922. 

Dwight, T. W. Timber administration of the Dominion Forest 

Service. Jour. Forestry 20: 4-9. Ja 1922. 

EUiott, C. Sterility of oats. U. S. Dept. Agr. Bull. 1058: 1-8. 

pL 1-4. 6 Mr 1922. 

Emerson, R. A, The nature of bud variations as indicated by 

their mode of inheritance. Am. Nat. 56: 64-79. F 1922. 

Emerson, R. A., & Hutchinson, C. B. The relative frequency 

of crossing over in microspore and in megaspore develop- 
ment in maize. Genetics 6: 417-432. S 1921. • 


Mont, within 

the host plant and on Its surface. Phytopathology 11 : 459- 
463. 25 F 1922. 

Ezekiel, W. N. Some factors affecting the production of 

apothecia of Sderotinia cinerea. Phytopathology^ 1 1 : 495- 
499. 21 Mr 1922. 

346 Index to American botanical literature 

Faris, J. A. Violet root tip (Rhizoctonia crocoriim DC.) in the 

United States. Phytopathology 11: 412-423, 16 F 1922. 

Farr, C. A. Quadripartltion by furrowing in Sisyrinchium. BulL 

Torrey Club 49: 51-61. pL 2. Mr 1922. 

Farwell, O. A. Corrections in botanical nomenclature. Am* 

Midi. Nat. 8: 33-35. Ja 1922. 

Includes new combinations under Dilepymm (i) and Gardenia (3). 


Faull, J. H. Some problems of forest pathology in Ontario. 

Needle blight of white pine. Jour. For. 20 : 67-70- J^ 1922 . 

Fernald, M. L. The Gray Herbarium expedition to Nova 
Scotia. Rhodora 23: 89-111. pi, 130, 19 My 1921; 23: 

130^152. 26 Au 1921; 23: 153-171. 24 O 1921; 23: 184- 
195. 15 D 1921; 23: 223-245- 26 Ja 1922; 23: 257-278. 
pL 133^ 27 F 1922; 23: 284-300. 22 Ap 1922. 

Includes new species in Lophiola (i) and Antennaria (i), 12 new varieties 
and 2 new forms. 

Fink, B. An addition to the distribution of a rare fungus. 


Mycoldgia 14; 49, 50. 6 Mr 1922. 

Tylostema verrucosum in Ohio. 

Fitzpatrick, H. M., Thomas, H. E., & Kirby, R. S. The Ophio- 
bolus causing take-all of wheat. Mycologia 14: 30-37. 
pL 10 + /. I. 6 Mr 1922. 

Fromme, F. D. Incidence of loose smut in wheat varieties. 

Phytopathology 11: 507-510. 21 Mr 1922. 

Gail, F. W. Factors controlling distribution of Douglas fir In 
semi-arid regions of the Northwest. Ecology 2: 281-291. /. 

1-3. 20 Ja 1922. 

Gaudron, J. El peso de las raices del Manihot utilissima en 
relacion con la fasciaclon de los tallos. Arch. Asoc. Peru 
1: 20-37. pl' ^^ 2. 1921. 

Gericke, W. F. Certain relations between the protein content 
of wheat and the length of the growing period of the head- 
bearing stalks. Soil Sci. 13: 135-138. F 1922. 

Gerould, J. H. The dawn of the cell theory. Sci. Monthly 14 : 
268-277. Mr 1922. 

Index to Americax ik)taxical literature 347 

Girola, C. D. Agallas de corona del duraziiero, Bol. Minis. 
Agric. 26: 257-259. pi. i, 2 (i colored), ' Jl-S 1921., 

k y - ■ 

Pseudornonas fumefaciens. 

Girola, C. D. Cultivo de la Yerba mate Torrefaceion y pre- 

I r 

r| paracion del producto. Bol. ]\IInis Agric. 26: 447-488. /. 

1-23. O-D 1 92 1. 


Girola, C. D. Sobre algunas enfermedades de la papa. BoL 
Minis. Agric. 26: 260-264. P^- ^y -• Jl^S 1921. 

Gleason, H* A.. The witch-hazels. Jour. New York Bot. 

Gard. 23: 17-19. F 1922. 


Goebel, K. Erdwurzeln mit Velamen. Flora 115: 1-26. /. i, 2, 



Graff, P. W. Philippine Basidiomycetes — IV. Bull. Torrey 

Club 48: 285-295. 19 Ja 1922. 

Gress, E. M. Annotated list of fossil plants of the Dakota 

formation (Cretaceous) in the collections at the Carnegie 

Museum. Ann. Carnegie Mus. 13: 274-332. pL 14, 75. 

192 I. . 

Includes 3 spp, nov. 

Greaves, J. E. Influence of salts on bacterial activities of soil, 

Bot. Gaz. 73: 161-180. Mr 1922 

Greenman, J. M. Two new senecios from the West Indies. 

Ann. Missouri Bot. Gard. 8: 97-102. pL i-z. 9 Ja 1922. 

Includes 5. snbsquarrosus and S. Freemanii, spp. nov. 

Grout, A. J. Mosses of a Staten Island house and lot* Bryolo- 
gist 24: 64. 9 Ja 1922. 

Guilliermond, M. A. Nouvelles obserxations sur Forigine des 

plastides dans les phanerogames. Rev. Gen. Bot. 33: 401- 
419, 449-466. /. 1--8, 1921, 

Haasis, F. W. ^ Relations between soil-type and root foim of 

western yellow pine seedlings. Ecology 2:292-303. /. 1-3. 
20 Ja 1922. 


Hall, H. M., & Long, F. I/. Rubber content of North American 
plants. Carnegie Inst. Washington Publ. 313" i~^5- P^- 

1-3. 1921. 

348' Index to American botanical literature 

Hallermeier, M. 1st das Hangen der Bliiten eine Schutzein- 
' richtung? Flora 115: 75 lo^- 1922. 


Harms, H, Uber einige CanVa-arten aus Sudamerika mit 

besondere Berucksichtigung der peruanischen Arten. No- 
tizbl. Bot. Gart. u. Mus. Berlin-Dahlem 8: 91-100. I Ap 

Includes 4 spp. nov. 

Harper, R. M. A botanical bonanza in Tuscaloosa County, 
Alabama. " Jour. Elisha Mitchell Sci. Soc. 37: 153-160. 

_ V 

pi. 28. Mr 1922. 

^ ■ 

Harper, R. M. Geography of central Florida. Florida Geol. 

Surv. Ann. Rep. 13: 71-307. /. 1-43. 1921. 

Harshberger, J. W. Geographic names and terms of signi- 
ficance in plant geology and ecology. Bull. Geog. Soc. 
Philadelphia 20: 32--46. Ja 1922. 

Harvey, L. H. Yellow-white pine formation at Little Manister, 

Michigan. Bot. Gaz. 73: 26-43. /• ^^^' 18 Ja 1922. 

Haskell, R- J. Fhytophthora infestans on egg plant in the United 
Stutes. Phytopathology 11: 504, 505. 21 Mr 1922. 

Hayes, H. K., & Stakman, E. C. Resistance of barley to HeU 

minthosporiiim sativum P. K. B. Phytopathology 11: 405- 
411. 16 F 1922. 

Heller, H. H. Certain genera of the Clostridiaceae, Jour. Bact. 

7: 1-38. Ja 1922. 

Includes 25 new genera. 

Hibbard, P, L. Some experiments on reclamation of infertile 

alkali soils by means of gypsum and other treatments. 
Soil Sci» 13: 125-134. F 1922. 

Higgins, B. B. Notes on the morphology and systematic re- 
lationship of Sclerotium Rolfsii Sacc. Jour. Elisha Mitchell 

Sci. Soc, 37: 167-172. pL 2g. Mr 1922. 


Hoemer, G* R. Infection capabilities of crown rust of oats. 

Phytopathology 12: 4-15. 10 Mr 1922. 

Hollick, A. Local flora notes — ^Staten Island. Torreya 22: 

1-3- Ja-F 1922. 

Vol. 49. 

No. 12 






Miscellaneous notes on plants of Southern California — II 


Philip A. Munz and Ivan M. Johnston 

Lemna trisulca L. 

Lemna trisulca L. Sp. PL 970. 1753. 

This plant, reported by Abrams (FL Los Angeles 78. 1904) 
from Bear Valley in the San Bernardino Mountains and by 
Torrey (Pacific R. R. Rep. 4: 142. 1856) from San Gabriel 
Creek, has been collected by Miss G. Corwin at the mouth of 
Santa Ana Canyon near Mentone, where it was found in a 
ditch at about 2000 feet altitude. It is, therefore, apparently 
to be looked for at various points along the Santa Ana River 


Ophrys monophyllos L. Sp. PL 947. 1753* 

Microstylis monophyllos LindL Gen. Sp. Orch. 19, 1830. 

A few typical specimens of this species, F. W. Peirson 22^1^ 
were collected in marshy ground at 8000 feet elevation on the 
South Fork of the Santa Ana River in the San Bernardino 
Mountains, where they grew with such plants as Gentiana 
simplex, G. humilis, Parnassia cirrala, and Luztila comosan This 
is the first collection west of the Rocky Mountains, 

Phoradendron calif ornicum var. distans forma leucocarpum 

Trelease forma nova 

Like the variety distans, but having white berries. 



[The Bulletin for November (49: 313-348) was issued November 9, 
*The first paper of this series was published in the Bulletin for Feb- 

ruary, 1922 (49: 31-44), 


350 MuNZ AND Johnston: Plants of California — II 

Type: Twenty-nine Palms/ southern edge of Mohave 
Desert, May i, 1921, Munz 455c, (Baker Herb. 9547)- 

Very conspicuous as white-fruited clumps growing in bushes 
of mesquite with the typicaLred-fruited form. Specimens of 


this collection w^ere sent to Dr. Wm. Trelease who considered 
it worth recognition and permitted us to publish it. A second 
collection, Munz 4703, likewise on mesquite, from Andreas 
Canyon near Palm Springs, can also be reported. 

.. Eriogonum nodosum var. Jaegeri var. nov. 

With habit, inflorescence, and floral structures of the typical 
form of the species but with stems tomentose only below on the 
leaf-bearing portion, the upper parts of the stem and the in- 
florescence being glabrous with, at most, a little^ wool in the 
axils. Involucres glabrous or slightly floccose, the subtending 
bracts woolly on their inner edges, 

J I 

Type: Dry Morongo Wash, near Hole-In -the- Wall Springs, 
Riverside County, November 15, i<)2i,E. C. Jaeger 251 (Baker 
Herb. 9424). 

This glabrate form of E, nodosum Small (Bulb Torrey Club 
25: 49, 1898) was collected at the eastern base of the San 
Bernardino Mountains, between 2500 and 3000 feet altitude, 
and well deserves recognition because of the totally different 
aspect as compared with the typical form. It is reported as 
very plentiful by its finder, Mr. Edmund C. Jaeger, one of the 
most active collectors of the desert flora. 

^ Eriogonum fasciculatum var. flavoviride var. nov. 

With capitate inflorescence and general habits of the var. 
polijolhim T. & G., but lower in stature, about one foot high; 
Hght green in color instead of ashy; leaves glabrous and green 
above, but somewhat tomentose and pallid below; calyx glabrous 
without and hairy within. 

Type: Pinyon Wells, southern edge of Mohave Desert, 
April 30, 1921, Mtmz 4505, (Baker Herb, 9425). 

This peculiar yellowish green glabrate plant was found to 
be abundant in a sandy desert canyon in high Lower Sonoran 
Zone above Pinyon Wells. It was associated all along the 
canyon w^ith var, polifolium, but was very distinct from this in 
size and color. We have seen also material from Cottonwood 



MuNz AND Johnston: Plants of California — II 351 

ium but was smaller and earlier in anthesis. The relation of this 
glabrous variety to var. polifoUum is analogous to that existing 
between typical E.fascicidatum and the puhescentv^r. foliolostim. 

Allionia pumila Standley 

AlUonia piimila Standley, Contr. U. S. Nat. Herb. 12: 345. 

Allionia Brandegei Standley, h c. 346. 

The only Allionia hitherto reported from California is A. 
Brandegei J the type of which came from the Providence Mountains 
in the eastern part of the Mohave Desert. Two collections 
recently made by F, W. Peirson in the San Bernardino Mountains 
are of this species, which has recently been included In A. 
pumila by Standley (No. Am. Flora 21: 226. 1918). These 
collections not only extend the range of this species to the western 
part of the desert but even into the coastal drainage. One 
{Peirson 2257) from the Santa Ana River, at 4500 feet elevation, 
is twice as tall as the type of A. Brandegei and has the leaves 
broadly ovate and cordate, rather than oblong and scarcely 
cordate. The other plant {Peirson 18^0)^ from Cushenberry 
Grade, is more like Brandegee's type collection, being about 
ten inches high and with leaves ovate to oblong. 

Allionia nyctagixea Michx. 

Allionia nyctaginea Michx. Fl. Bor. Am. 1: 100. 1803. 
Allionia ovata Pursh, Fl. Am. Sept. 97. 18 14. 






to have been destroyed by the oil spray used by the railroad 
for weed destruction. This is apparently 
the state. 





ScopulopUla nitrophiloides Jones, Contr, W. Bot. 12: 5. 1908. 
EremoUthia Rixfordii Jepson, Fl. California 499. /. 100. 1914. 
This odd desert plant seems to be generically distinct from 
A chyr onychia, but Jones's ScopulopUla has precedence over 
the much later generic name EremoUthia of Jepson, 

352 MuNz AND Johnston: Plants of California — II 

/ Claytonia lanceolata var. Peirsonii var. nov. 

As in the typical form of the species, but with peduncle and 
primary axis of inflorescence shortened so as to make the flowers 
appear umbellate. 

Type: near Kelly's Cabin on Ontario Ridge, San Gabriel 
Mountains, at 8200 feet altitude, May 29, 1920, F. W. &. 
Mabel Peirson (Baker Herb. 10524). 

Known only from the eastern portion of the San Gabriel 
Mountains where, beside the type, we have seen a collection, 
F. W. Peirson 21^1, from the Devirs Backbone at 7000 feet 
altitude. It is probable that this variety will necessitate 
a reconsideration of the status of C umbellata Wats. (Bot- 
King Exp. 43. pL 6,f.4-5. 1871) of the northeastern part 
of California, for the principal character of that species is the 
same umbellate inflorescence that characterizes our variety. 
The new variety, however, may be told from C* umbellata 
by its shorter-pet ioled, broader leaves, which are widest below 
the middle (instead of above), and by its shorter stem, larger 
flowers, and isolated southern range. 

Thlaspi arvense L. 

Thlaspi arvense L. Sp. PI. 646. 1753. 

This crucifer, common in the Rocky Mountain region, can 
now be reported from Southern California {F. W. Peirson 66) 
at Henniger's Flats near Pasadena. The only other collection 
we have seen from California is by Chase from Adin, in Modoc 

^ Draba corrugata var. saxosa (Davidson) comb, nov, 
Draba saxosa Davidson, Bull. So. Cal. Acad. 19: 11. 1920* 

Peak may well be deserving of a varieta 
of a specific one. As pointed out in anot 
PI. World 22: 88. 1919) the San Jacinto 
matters of degree and is closely simulate< 
from the San Bernardino Mountains, 



MuNz AND Johnston: Plants of California — II 353 

• Sphaeralcea rosacea sp. nov. 

A shrubby perennial forming loovse clumps 10-15 d^- high; 
stems ashy, stellate-tomentose, slender, ascending, virgate ; 
leaves remote, ovate- or oblong-cordate and somewhat three- 
lobed, doubly crenate, stellate-pubescent but greener above, 
blade usually 2-2.8 cm. long and 1.5-2.5 cm. wide, petioles 
12-20 mm. long; inflorescence a very loose open panicle 2-4 
dm. long, the ascending branches remote and one- to four- 
flow^ered; calyx stellate-tomentose, at an thesis 8-11 mm. high 
and 5~7 mm. wide, with lanceolate or oblong-lanceolate lobes, 
at fruiting 10-14 ™™' ^<^ng, bractlets setaceous, deciduous, 
4-7 mm. long ; pedicels 3-20 mm. long; petals pinkish when 
fresh but drying a rose-violet, 17-21 mm. long, claw densely 
villous-ciliate; fruit much depressed, stellate-pubescent; carpels 
one-seeded, about 3.5 mm. high, oblong in outline, sides smooth 
or at least not reticulate, seminiferous only below, the upper 
half thinner and scarious. 

Type: Palm Springs, April 2, 191 7, Alice B. Chittenden 
(Herb. Cal. Acad.)- 

A very ornamental species that is known to us only from 
the vicinify of Palm Springs, Riverside County, where it is a 
common plant in rocky ground in the canyons opening into the 
Colorado Desert. Besides the type we have seen a collection 
by Parish, No. 4109, and one by Johnston. This plant is the 



Robinson (Synop. FL 1 : 315. 1897) in his review of the genus. 


distinct and probably not even an Immediate relative of that 
species, differing from it in habit and stature; in size, color 
and disposition of flowers; and in shape and texture of leaves. 
When we first saw the two species growling together, as they 
commonly do about Palm Springs, we never once thought 
of associating the two plants under one name; in fact, basing 
determination on superficial resemblances, the new species was 
taken to be a species of Malvastrum. The proposed species 
bears a considerable resemblance to certain of the shrubby mal- 
vastrums, having more or less woody stems and similarly colored 
flowers, but the technical characters of the carpels definitely 
ally it with Sphaeralcea. 

Viola Sheltoni Torr. 
Viola Sheltoni Torr.Pac.R.R. Rep. 4:67. pi. 2. 1856. 

354 MuNZ AND Johnston: Plants of California — II 


• Locally common on moist, shaded > oak-covered canyon-sides 
in the upper chaparral belt on the south slope of the San Antonio 
Mountains, where specimens have been collected In Charcoal 
Fork of Cucamonga Canyon, at 4200-4500 feet altitude, Johns- 
ton 1303 f and near Brown's Flats, at 4200 feet, Jolmston 1759- 
The specimens appear the same, character for character, as the 
common plant of Mendocino and Siskiyou Counties. The plant 
was known to range as far south as Mariposa County, where it 
was collected by Congdon, but between that station and the 
one in Southern California, which we now report, there lies a 
gap of over 280 miles in which the plant is not known to occur. 
This station for F. Sheltoni was announced as V. lobaia by 
Johnston (Bull, So. Calif, Acad. 17: 65. 191 8). 

Oenothera speciosa Nutt. 

speciosa Nutt, Tour. Acad, Phila 


Well establ 

the confluence with Ihe Santa Ana River. First collected 
several years ago by Johnston. Recently the following specimens 
have been collected near Pomona, all in moist soil along the 
stream: Mrs. Leningion^ Mnnz 4680, 4560. These discoveries 
add a new species and subgenus of Oenothera to the flora of Cal- 
ifornia. Our material is atypical in its rounded instead of ob- 
cordate petals and in being taller, some of the specimens attaining 
a height of four feet. 

^Oenothera cardiophylla var. splendens var. nov. 

A large flowered variation with hypanthium two-thirds as 




A very remarkable variation of the species and, because of 
its large flowers, probably of ornamental value. We know of 
but two collections, the one from The Needles by Grinnell and 
the other from the Chocolate Mountains by Childs. The 
foliage and habit of the plant are those characteristic of the 
species. No doubt some will consider this distinct enough for 
a species, but we are deterred from publishing it as such because 
of the immense amount of variation in the floral structures of 

MtJNZ AND Johnston: Plants of California— H 355 

cardiophylla, of which this 

able extreme. 

Chimaphila umbellata (LO Nutt. 

Pyrola umbellata L. Sp, PL 396- 1753. 
Chimaphila umbellata Nutt. Gen. 1: 27 
Chimaphila occidentalis Rydb. No. Am. 


f>ccmentalts Rydb. No. Am. Flora 29: 30. 1914. 

This species can now be reported from Southern California, 
having been collected in the Dollar Lake region of the San 
Bernardino Mountains at 8750 feet altitude, F. W. Peirson 82g. 
This discovery and that of Pyrola picta Smith, Johnston 2860, 
add two species of Pyrolaceae to the list prepared by Parish 
(PI. World 20: 247. 1917) for the San Bernardino mountains. 


Asclepias eriocarpa var. microcarpa var. nov. 

Follicles 3-4 cm. long; seeds 5 mm. long; coma 15 mm. long; 
pedicels 2-3.5 ^^ • ^^^Sl ^^ flowers and vegetative parts like the 
typical form of the species. 

Type: North Fork of Deep Creek, San Bernardino Moun- 
tains, on a sunny hillside at 6000 feet altitude, July 17, 1921, 
/. M. Johnston 2833 (Baker Herb. 9563.) 

This variety is the form of the species frequent through the 
pine belt of the San Bernardino Mountains and differs from the 
typical form, which is not uncommon on the plains at the foot 
of the mountains, in its much smaller follicles. Many plants 
of the valley-inhabiting A, eriocarpa Benth. have been seen by 
the authors, but none have been noted which failed to have 

Inrcrf^r thaTi in the varietv here described. A. erio- 

fruit much larger t 
carpa has follicles 8 

4-6 cm 

measure about 8 mm. in length 

mm. long. 

The small follicles can not be attributed to the blighting 
action of frost, for all produce perfect seeds and in all colonies seen 



exploration may reveal the fact that the pine-belt form of A . 
eriocarpa found in the San Jacinto and San Gabriel Mountains 
is also referable to this variety; fruiting material from these 
ranges is lacking. 




>rry * - 

b emardina sprnriDvr 

• Phlox doUchantha Brand; Engler, Pflanzenreich 4^^^: 67. /. 
i8,A-D. 1907; Nels. W. Am. Phloxes 28. 1898; Milliken, 
Univ. Cal. Pub* Bot. 2: 65. 1904. Not P. doKchantha Gray, 

Proc, Am, Acad. 22: 310. 1887. 

Type: In open pine forest, 6000 feet altitude, Bear Valley, 
June 1886, Parish 1818, (Univ. Calif. Herb. 194015). 

The new species indicated here is the plant from the San 
Bernardino Mountains that has been known as P. doUcantha. 
The type of P. doUchantha Gray, howe\'er, came from the Pahran- 
gat Mountains in southern Nevada and is evidently a form of 
what has been called P. Stanshuryi brevifolia and P. superba. 
In the recent treatments of Phlox by E. Nelson, Milliken, and 
Brand, the plant from the San Bernardino Mountains has 
uslirped the name of P. doUchantha to the exclusion of the 
original Nevada plants. P. bernardina differs from P. doUchantha 
and Its immediate relatives m its very elongated corolla-tube, 
which is over four, instead of less than three, times the length 
of the calyx; in having its stems not shrubby below but her- 
baceous to the ground; and in its habitat, which is not at low 
altitudes in desert mountains but in the open pine forests of 
the San Bernardino Mountains, The new species is separated 
from its nearest relative by over 100 miles. It is known only 
from San Bernardino Mountains and appears to have been 
collected there in Bear Valley only. 

Nama humifusum Brand 

Nama humifusum Brand, Beitrage Hydrophyll. 9, 191 1. 

In Southern California this species has been known only 
from the three localities (Soldiers^ Home, Los Angeles County, 
and Diamond Lake and Sweetwater Valley in San Diego County) 
given by Brand in the Pflanzenreich (4^^^: 146. 1913); the 
plant, however, may be reported from Laguna Canyon In Orange 

County, where it was collected in May, 1919, MunZy Street, 
& Williams 2681. 

Lycium Spencerae Macbrlde 

Lycium Spencerae Macbrlde, Contr. Gray Herb. 53: 18. 1918. 

The type of this species proves to be a flowering specimen of 

Primus fasciculala (Torr.) Gray (Proc. Am. Acad. 10: 70. 

MUNZ AND Johnston: Plants of California— II 357 

1874). The identifying of this lycioid almond with L. Spencer ae 
adjusts for local botanists the problem of explaining how so 
distinct a species, as this Lycium seemed to be, could so long 
go undetected in a locality so frequently visited as Cajon Pass. 
We are under obligations to Mr. F. Macbride for making at 
the Gray Herbarium the comparisons necessary for the establish- 
ing of the identity of the two species. 



Penstemon Cleveland! var. connatus var. nov. 

With habit of growth and flowers of the species, but with 
leavfes jagged-serrate, usually glabrous and with at least the 
four upper pairs connate-perfoliate. 

Type: near Van Deventers, southeastern base of the San 
Jacinto Mountains, June 1901, H. M. Hall 214Q (Univ. Calif. 
Herb. 54999)- 

A very well marked variation that Is apparently restricted 
to the eastern (Colorado Desert) base of the San Jacinto Moun- 
tains. Referable to this variety are the following: Hall 1160 
from tlie type locality, Parish Bros. 1216 and Johnston, April, 
191 7, from near Palm Springs, The most evident relationship 
is with the variety Stephensi of the eastern Mohave Desert, 
the new variety agreeing in its foliar characters but differing 
In its somewhat smaller corolla and beardless sterile stamens. 
In our recent discussion of the var. Stephensi (Bull. Torrey 
Club 46: 41, 1922) the specimens of P. Clevelandi Gray mentioned 
as simulating that variety in habit are in fact referable to var. 

P. Clevelandi' in its typical form occurs along the desert 
borders to the south of the range of the var. connatus. The 
northernmost locality seen for it is Coyote Canyon, Hall 2766. 
In the typical form the leaves are entire or rarely remotely 
dentate, never glaucous and are entirely free or, at most, only 
partially joined and not completely fused, with the suture line 
obliterated as In the variety. 

/Galium angustifolium var. pinetorum var. nov. 

A low herbaceous perennial, stems 2-4 dm. high, simple, 
erect, tufted or arising from along short, prostrate, persistent, 

subterranean stems. 

Type: Sierra Madre Mountains, Los Angeles County, 1893, 
A. Davidson (Univ. Calif. Herb. 28169). 

358 MuNz AND Johnston: Plants of California — II 


form of the species only in habit of growth. Additional speci- 
mens referable to this variety are the following: Santa Lucia 
Mountains, K. Brandegee, on the north, and Cuyamaca Moun- 
tains, r. 5. Brandegee, July 7, 1894, on the south. Plants from 
the San Jacinto Mountains, Hall 2S09, have the growth-habit 
of the variety, but are strictly branched and become 5-7 dm. 
high. The variety has been mentioned without a formal name 
by Johnston (PI. World 22: 118. 1919). 

• Galium Hallii sp. nov. 

Dioecious perennial, shrubby at the base, with relatively 
few long (4-6 dm.) decumbent qu^adrangular stems, younger 
stems short-hispid, older stems glabrate and with a shiny ex- 
foliating papery bark; leaves in fours, hirsute like the branchlets, 
light green, ovate-elliptical to elliptical, 5-1 1 mm. long, sessile, 
margins strongly revolute, midvein strong, lateral veins weak 
or absent; flowers borne in loose leafy clusters on the con- 
spicuously and gracefully recurved tips of the branchlets, yellow- 
ish, about 2 mm. in diameter, bearing a few stiff hairs; fruit 
black, slightly juicy, body 3 mm. in diameter, covered with 
dense villous hairs of 2 mm. length. 

Type: Coldwater Fork of Lytle Creek, San Gabriel Moun- 
tains, in gravelly ground at 5200-5700 feet altitude, July 7, 
191 8, Jchnston 206^ (Univ. Calif, Herb.)- 

Specimens examined: Coldwater Fork of Lytle Creek, San 
Gabriel Mountains, 5500 ft. alt, Johnston 1667; Lytle Creek, 


5750 ft. alt., Hall 122^; Rock Creek Canyon, San Gabriel Moun- 
tains, Abrams & McGregor 5g8; Seymour Creek, Mt. Pinos 


region, 6700 ft. alt., Hall 6^0$; Tehachepi, 1889, Greene; Erskin 
Creek, 4-5000 ft. alt., Pur pus 5349, 

This is a remarkably clear-cut undescribed species of the 
G, multiflorum-allmnce that has been confused with G. multi- 
floruniy G. stellatum, and G. occidentale; from all of these it 
certainly differs and can be distinguished by its noddi'ng flower- 
clusters. The plant has been recognized as distinct for several 
years, and the increasing material at hand seems to justify its 
description as a new species. It has been treated as ^^ Galium 
sp-" by Johnston (PI. World 22: 118. 1919). 

The range is eminently a natural one, for the plant occurs 
at and just below the lower limits of the pine belt in the rocky 
canyons that open into the extreme western portion of the Mohave 

MuNZ AND Johnston; Plants of California— II 359 

Desert. The type locality, the southernmost known station, is 
almost on the Los Angeles— San Bernardino County line; 
from this point the range extends northward to Erskin Creek 
in the Sierra Nevada of Kern County. 

It might be mentioned here that G. occidentale McClatchie 
(Erythea 2: 124. 1894) is a trivial form of G. calif ornicum 
H. & A., and that G. grande McClatchie (/. c.) is indistinguishable 
from G. piihens Gray. These identities are established by a 
study of authentic specimens. 

Bahia dissecta (Gray) Britton 

Amatiria dissecta Gray, Mem. Am. Acad. II. 4: 104. 1849, 
Bahia dissecta Britton, Trans. N. Y. Acad. Sci. 8: 68. 1888. 
Villanova dissecta Rydb. Bull. Torrey Club 37: 333. 1910. 
A mauriopsis dissecta Rydb. No. Am. FIora34 12,7. 1914- 
The first collection of this species to be reported from the 
state was made in the San Bernardino Mountains, F. W. Peir- 
son 224s, along the Santa Ana River, at 6500 feet elevation, 
August 24, 1920. 

Pomona College 

Claremont, California 

Notes on Carex — XII 

Kenneth Kent Mackenzie 

I* An inland relative of a salt marsh sedge 

The only representative of the Ovales which is an inhabitant 
of salt marshes, as far as I am aware, is Carex hormathodes 
Fernald, which is an abundant and characteristic species along 
the North American coasts from western Newfoundland to 
southeastern Virginia.* In 1902 attention was first drawn by 
Professor Fernald to an allied pla it found in fresh water swamps 
and reported by him as growing from Massachusetts to the 
District of Columbia. This was first treated by him slS Carex 
tenera Dewey var. Richii Fernald, and later on as Carex lior- 
mathodes var. Richii Fernald. 

In northern New Jersey this variety Richii is an abundant 
plant in places in swampy meadows, and with its graceful 
nodding heads is one of the most pleasing and attractive of our 
sedges. It is in every way a much more handsome plant than 
is Carex hormathodes. But not only does it have an entirely 
different habitat and a different appearance, but It has technical 
characters which are absolutely good. The perigynia have 
suborbicular bodies abruptly contracted into the beaks; the 
perigynia are loosely ascending or spreading in the spikes; 
the terminal spike is abruptly contracted at base below the 
perigynia, and the staminate portion is conspicuous; and the 
spikes are scattered in a very flexuous head. In Carex hormatho- 
des on the other hand the perigynia have oblong-ovate bodies 
much less abruptly contracted into the beaks; the perigynia 
are appressed; the terminal spike tapers at base, and the staminate 
portion is short; and the upper spikes are approximate. The 

* The Inland reports for this species are all erroneous, as also are the re- 
ports of Its occurrence from further south than southeastern Virginia. It 
however is to be expected at least as far south as North Carolina. The van 
invisa (Boott) Fernald represents nothing but stunted specimens. 


362 IVIackenzie: Notes on Carex— XII 

dtflferences between the two species are well brought out In the 
figures (No. 356-8) in Gray's ManuaL 



Bayard Long, both of whom are well acquinted with it in the 



Carex Richii (Femald) Mackenzie sp. nov. 

Carex tenera Dewey var. major Olney, Exsicc. fasc. 2, No, 15 
(in part). 1870, 

Carex tenera var. Richii Fernald, Proc, Am. Acad. 37: 475. 

Carex hormathodes Fernald var. Richii Fernald, Rhodora 8: 166. 

Densely cespitose, the rootstocks very short-prolonged, 
black, fibrillose, the culms 6-10 dm. high, erect, but very slender 
to base, nodding, sharply triangular, very rough beneath head, 
exceeding the leaves, dark brownish at base and clothed with 
the dried-up leaves of the previous year, the lower bladeless; 
leaves of the year two to four to a culm, on lower fourth, but 
widely separate, the lower nodes often exposed, the blades flat, 
1.5-2.5 mm, wide, usually 1-3 dm. long, long-attenuate,^ the 
sheaths green striate ventrally nearly to mouth, the ligule 
surrounding the culm, short-prolonged beyond base of blade; 
spikes four to eight, strongly separate in a moniHform, flexuous 
head, gynaecandrous, the staminate flowers conspicuous, 
tapering clavate or in the terminal truncate clavate at base, 
the pistillate part oVbicular to ovoid-orbiculur, 6~I2 mm. long, 
6-9 mm. wide, rounded at apex, and with some twenty to thirty 
spreadinjg or spreading-ascending perigynia with conspicuous 
tips; lower bracts setaceous, prolonged, the upper scale-like; 
scales lanceolate, hyaline and yellowish-brown tinged, acuminate 
or aristate from the sharp midvein, shorter and much narrower 
than the perigj^nia; perigynia very flat and thin except where 
distended by achene, membranceous, greenish, or in age straw- 
colored, 4-5 mm. long, 2,5 mm. wide, the body suborbicular, 
widest near middle, strongly winged to base, serrulate above 
middle, finely about ro-nerved on either side, round-truncate 
and sessile at base, abruptly contracted into the flat, serrulate, 
red-tipped, obliquely cut dorsally at length bidentate beak, 
more than half length of body; achenes lenticular, stipitate, 
oblong-obovoid, 1.5 mm. long, 0.75 mm. wide, yellowish-brown, 
apiculate; style slender, obscurely jointed with achene; stigmas 
two, reddish, slender, short* 


XII 363 

Specimens examined* 

Massachusetts: Middleboro, Murdoch, June, 1900 (N.E.) 
Canton, Churchill, June 17, 1900 (N.E.); Dover, Winslow, 
June 21, 1912 (N.E.); T>e6ha.m, Forbes, June 10, 1912 (N.E.); 
Stony Brook Reservation, Kidder, June 14, 1920 (N.E.); Sud- 
hmy, Rich et at. qq (Type), June 17, 1902 (N.E., H., P., N. Y., 
K.M.); Bedford, Murdoch, June 7, 1902 (N.E.); Weston, 
Weatherby , ]nnQ 20, 1909 (N.E.) ; West Cambridge, Pease 3084, 
May 27, 1903 (N.E.) ; South Sudbury, Miss Carr, June 30, 1897 
(N.E.); Cambridge, 5m/ (H,), also Hubbard, June 2, 1896 (N. 
E.), also Deane, June 8, 1887 (H.), also Femald, June 10, 1891 
(Col.); Stoneham, i^icA, June 5, 1894 (N.E.); Reading, Perkins, 
June 14, 1883 (N.E.), also Manning, June 8, 1882 (N.E.); 

Arlington, Wm. Boott, June 5, 1870 (H.); Mystic Pond, Wm. 

Boon. June 5, 1870 (H.); Amesbury, Eaton, June i, 1903 (H.); 

Sherborn, Loomis lojj, June 19, 1912 (K.M.). 
Rhode Island: Providence, Olney (N.Y.). 
Connecticut: Waterbury, Blewitt, June 15, 1911 (N.E.), 

also 1287, June 10, 1912 (N.E.) ; South Windsor, Woodward, 

June 2S, 1909 (N.E.); Columbia, Weatherby 4000, June 26, 

Pomfret, J 

Weatherby 4114, July 

Wright, May 

(H.); Sprague, Graves, June 13, 1902 (H.); Griswold, Graves, 
June 16, 1899 (H.); Killingly, Bissell, June 22, 1902 (H.); 
Thompson, Bissell, June 22, 1902 (H.); Oxford, Ilarger, June 5, 

?, June II, 1905 (P.) ; East Hartford, Weather- 

June 24, 1904 

by. May 25, 1903 
(K. M., H.) ; 

New York: Junius, Sartwell (H.), 

New Jersey: CXoster, Austin (H.); Newark, Livingston js> 
May 27, 1894 (H,); Crosswick's Creek, Britton, May 29, 1904 
(N. Y.) ; Passaic River, Bergen County (N.Y.) ; Oradell, Bergen 
County, Mackenzie 75^, June 19, 1904 (K.M.); Stirlington, 
Morris County, Mackenzie 1267, May 28, 1905 (K.M.); Mon- 

* Specimens are cited from herbaria as follows: Columbia University 
(Col.); Gray Herbarium, Harvard University (H.);K. K. Mackenzie (K.M); 
New England Botanical Club (N.E); New. York Botanical Garden (N.Y.); 
New York State Museum (Albany); Philadelphia Academy of Natural 

Sciences (P.). 


364 Mackenzie: Notes on Carex — XII 

mouth Junction, Mackenzie 49^3, May 26, 1912 (K.M,) ; Succa- 
sunna, Mackenzie 4128, June 13, 1909 (K.M.); Delanco, Burling- 
ton County, long 3474, May 19, 1910 (P.); Centre Square, 
Brintm, May 20, 1892 (P.) ; eight miles southwest of Mickleton, 
Heritage, May 25, 1892 (P.); Budd's Lake, Morris County, 
Heritage, May 28, 1895 (P.). 

Pennsylvania: Easton, Garber, 1868 (P.), also Porter, 
June 5, 1890 (P.); Lehigh Gap, Lehigh County, Pretz 1831, 
May 20, 1909 (P.); Slatington, Lehigh County, Pretz 10182, 
June 6, 1920 (P.), also Bachman 2132, June i, 1911 (P.) ; Strass- 
burg, Lehigh County, Pretz 335s, May 20, 191 1 (P.) ; Cementdn, 
Lehigh County, Pretz 10131, May 30, 1920 (P.); Rockhill, 
MacElwee, May 31, 1903 (P.); Quakertown, Bucks County, 
Marlindale (P.); Sellersville, Pretz, May 25, 1884, June i, 1888, 
and May 26, 1896 (P.), also Crawford, June i, 1989 (P).; Penn 
Valley, Bucks County, Crawford, June i, 1899 (P.); Tully- 
town, Crawford & Brown, 1902 (P.); Byberry, Martindale, 
June, 1864 (P,); Wissahickon, A. H. Smith (P.); Sugartown, 
Chester County, Pennell & Long ^628, June 26, 1912 (P.); 
Nottingham, Chester County, Pennell & Long 7483, June 22, 
1912 (P.); Mercersburg, Franklin County, ex herb. Porter, May 
25, 1847 (P.); Wyoming Valley, Thurston, 1892 (P.); Elk 
County, McMinn, 1868 (P.). 

Delaware: Wilmington, Commons, June 30, 1896 (P.); 
Vandyke, Newcastle County, Long May 24, 1909 (P.); Town- 
send, Commons, May 17, 1883 (P.), 

Indiana: Palmyra, Harrison County, Beam 23417, June 4, 
1917 (K.M., H), 

2, A segregate from Carex albolutescens 

Among the varieties of Carex slraminea Willd. recognized 
by Professor Bailey was var. cumulata Bailey, based on speci- 
mens collected in Pennsylvania by Pojter, in New Brunswick 
by Fowler and on the Saskatchewan Plains by Macoun. This 
variety, proposed in 1889, was maintained by Bailey in 1890 
in the 6th Edition of Gray's Manual (p. 622), where its dis- 
tribution was given as from Pennsylvania to New England and 
northward. It was said to be rare and to 
Three years later in 1893 " Professor Bailey placed the plant 

grow in dry gro 


Mackenzie : Notes ON Carex — XII 365 

ing to his previous publications concerning its distinguishing 
characters or range; and this treatment is followed by Kiikenthal 
in the Pflanzenreich (4^0: 209). By other authors Professor 



As shown however by the large number of specimens of 
Bailey's variety which have been collected in recent years, it 
not only has an entirely different range from Carex albolutescens 
but it has several constant and well-marked points of difference, 
and is clearly entitled to specific rank. It is a northern species 
ranging from Saskatchewan to New Brunswick and south to 
Massachusetts, Connecticut and northern Pennsylvania- It 
occurs in a great variety of habitats varying from wet to dry, but 
seems able to thrive in much drier places than can Carex 
albolutescens. In the southern part of its range it seems only 
to be recorded from the higher elevations. It is not a coastal 
plain plant like Carex albolutescens^ but like that species Is not 
a species of limestone areas. 

As above stated Carex albolutescens is typically a coastal 
plant. It reaches its northern limit in eastern Massachusetts 
but has a very extensive range to the south, being found as 
far south as northern South America. In the more southern 
part of its range it gets into the mountains. 

The best technical character to separate the northern plant 
from Carex albolutescens is that in it the perlgynia are nerveless 
ventrally, while in Carex albolutescens they are strongly several- 
nerved. But in addition to this the spikes are numerous (five 
to thirty) and densely aggregated ; the green perlgynia are more 
conspicuous than in Carex albolutescens, thus giving the heads 
a characteristic heavy, green, or in age brownish, appearance; 
the sheaths are looser, and the leaf -blades of the sterile culms 
are usually wider, being 4-5 mm. In width; in developed plants 
the lateral spikes also are truncate-rounded at base. 

Carex albolutescens has fewer spikes (three to ten), which 
are less densely aggregated ; and the heads have a very character- 
istic silvery-green appearance; its sheaths are tighter than in 
the northern plant and the leaf-blades of the sterile culms are 

* The perlgyniuni and the heavily spiked head, figured in the seventh 
edition of Gray's Manual (/. 35^) as Carex albolutescens, represent Bailey's 
plant, and the head with five spikes represents genuine Carex albolutescens. 

366 Mackenzie: Notes on Carex — ^XII 

normally narrower and but 2.5-3 mm. in widtli (although 
occasionally more) ; and the lateral spikes are round-tapering 

at base. 

The northern plant is evidently abundantly distinct and it 

should be treated as 


M acken 

Carex alala Torn var. pulchra Olney (mostly), Exsicc. fasc. 2, 
No. 23 (also Bailey Proc. Am. Acad. 22: 150 as synonym). 

Carex stramtnea Willd, var. cumulata Bailey, Mem. Torrey 
Club 1: 23. 1889. 

Carex alholulescens Schw. var* cumulata Bailey, Bull. Torrey 
Club 20:422. 1893. 

Very densely cespitose, the rootstock not prolonged, the 
culms 3-9 dm. high, stiff, sharply triangular, roughened beneath 
head, brownish at base aiid clothed with the dried-up leaves of 
the previous year, the lowest bladeless; leaves of the year with 
well-developed blades two to four to a culm, on lower third but 
not bunched, the lower nodes sometimes exposed, the blades 
flat, light-green, thickish, erect or ascending, 3-5 mm. wide, 
7-25 cm. long, roughened towards apex, the sheaths loose, 
strongly green striate ventrally and conspicuously prolonged be- 
yond base of blade and continuous with the ligule; sterile slioots 
elongated, the blades usually 4-5 mm. wide, occasionally nar- 
rower, erect, the sheaths very loose; spikes five to thirty, gynae- 
candrous, very closely aggregated into an ovoid or oblong head 2-4 
cm. long, 1-2 cm. thick, the spikes ovoid, 6~io mm. long, 5-8 
mm. thick, short-pointed at apex, the lateral truncate or rounded 
at base, the terminal clavate, greenish appearing or in age 



conspicuous; scales ovate, obtuse or acute, much narrower and 
shorter than the perigynia, white-hyaline with one- to three- 
ner\*ed green center; perig^^nia thin, plano-convex, 3-4 mm. 
long, 2.5 mm. wide, membranaceous, greenish or in age brownish, 
the body orbicular-obovate, widest at top, round-tapering and 
sessile at base, wing-margined nearly to base, serrulate above 
the middle, nerveless ventrally, lightly several-nerved dorsally, 
very abruptly contracted into the short, serrulate, flat, obliquely 
cut dorsally beak about 0.75 mm. long, about one-third to 
one-fourth the length of the body; achenes lenticular, oval- 
obovoid, at maturity yellowish brown, very short substipitate, 

2 mm. long, 1,25 mm» wide, bent-apiculate; style slender, 


Mackenzie: Notes on Carex — XII 367 


jointed with achene, at length deciduous; stigmas two, slender, 
Hght-reddish, short. 

Specimens examined 

New Brunswick: Bass River, Fowler , August 9 , 1872 (H, 
N.Y.); Miramichi, Fowler, August 24, 1894 (H.), also Williams 
& Fernald, July 25, 1902 (H.); Kent County, Fowler (Type), 
July 1870 (H., N.Y., Col. Albany, P.). 

Nova Scotia : Bell 's Island, Laliave River, Macoun 82318, 
August 10, 1910 (H.); Bridgewater, Macoun 82319, July 18, 
1910 (H.); Sprlnghill Junction, Pease & Long 20309, July 18, 
1920 (K.M.) ; Port Mouton, Fernald & Graves 20312, Atigust 18, 
1920 (K.M.); Middleton, Fernald et aL 20323, July 20, 1920 (K. 
M.) ; Broad River, Fernald & Bissell 20311, August 16, 1920 

Maine: Somesville, Fernald, September 22, 1892 (N.E.,H.); 
Bangor, Knight, August 7, 1904 (H.) ; Pembroke, Fernald 1439, 
July 17, 1909, (N.E., H.); Southport, Fernald, Augast i and 2, 
1894, (N.E., H); Kennebunk, Parlin 867, August 26, 1897 
(H., N.E.); Mt. Desert Island, Greenleaf, August 3, 1883 (N.E.), 
also Rand, Redfield, Faxon, Fernald, Williams, numerous col- 
lections 1890-1898 (H., N.E., Col., N.Y.); Milford, Fernald & 
Long 12928, August 25, 1916 (N.E.); Lovell, Pease 17307, 
August 30, 1918 (N.E.); Matinicus, C. A. E. Long, August 28, 

1919 (N. E.) ; summit of Mt. Battie, 1325 feet, Furbish, July 14, 
1903 (N.E.), also Bissell, August 13, 1913 (N.E.); Durham, 
Dinsmore, August, 1898 (N.E.); Bear Mountain, Livermore, 
Furbish, July 24 and 25, 1896 (N.E.); Kennebunk, Deane, 
July 5, 1894 (N.E.) ; Berwick, Parlin 1062, July 24, 1898 (N.E.) ; 
North Berwick, Fernald, July, 1897 (N.E.). 

New Hampshire: Nottingham, Eaton, June 20, 1900 (H.); 
Mt. Monadnock, 3000 ft., Harper, July 3, 1899 (N.E.), also "A. 
W. Z>.," August 7, 1916 (N.E.). 

Massachusetts: Mt. Watatic, Ashby, Knowlton, August 

9, 1908 (H.); Medford, W. Boott, June 30, 1861, (N.E.); July 
25, 1863 (H.); Edgewith, W. Booll, July, 1853 (H.); Ashland, 
Morong, July 10, 1879 (H.) ; West Manchester, Hubbard 193, 
October 11, 191 1 (N.E.); Middlesex Falls, Kidder, June 28, 

1920 (N.E.); Wilmington, Collins, June 2$, 1887 (N.E.); Mt. 
Washington, Berkshire County, Hoffman, August 23, 1914 

368 Mackenzie: Notes on Carex — XII 

(N.E.); Lake Undine Dome, Berkshire county, Hoffman, August 

27, 1902 (N. E.). 

Connecticut: East Windsor Hill, "A. W. Z>.", August 26, 
191 5 (N.E.); South Windsor, Bissell, July 30, 19 16 (N.E.); also 
Blewitt 1S02, July 10, 1912 (N.E,), also 3470, August 8, 1917 


New York: Taghkanick Range, Hoysradt, July-August 

(H.) ; Karner, Albany County, House, 6410, July 7, 1919 (Albany) ; 

Whitestown, Oneida County, Haberer 3760, August 22, 1912 


(Albany); Ft. Ann, Washington County, Burnham 4Q, June 19, 
1892 (N.Y.); Sams Point, alt, 2600 ft., i?w55y, August 20, 1896 
(CoL); Mt. Riga, Hoysradt, July 17, 1878 (Col.)- 

Pennsylvania: Naomi Pines, Porter, July i, 1893 (P.)t 

also Britton, Jiily 2, 1893 (Col.); Pocono Plateau, T. Greene^ 
September 4, 1861 (P.), also Porter, September 11, 1869 (P.), 
July25,i88i (P.), August 6,1891 (Col.,HO,and July i, 1893 (P.). 

Michigan: Alger, Wheeler, July 2, 1895 (H,); Omer, Arenac 
County, Wheeler, July 24, 1902 (H.); Duck Lake, Arenac 
County, Wheeler, 1900 (H.);Townsliip 18, n. 4 E., Section 24, 
Wheeler, August 8, 1900 (H.) ; Township 19, n. 6 E,, Section 16, 
Wheeler, August 9, 1900 (H.); Emerson, Chippewa County, 
Dodge, August 28 and September 4, 1914, (K.M.); Manistlque, 
Schoolcraft County, Dodge, September 22, 1915 (K.M.) ; Alpena, 
Wheeler, July 8, 1895 (H., CoL). 

Ontario: Lake of the Woods, Macoim jj, July 28, 1872 


Saskatchewan: Fort Ellice, Macoun 2ig, 1872 (H.)- 



In the northern part of the United States from Maine to 
Michigan and in southern Canada from Ontario to Manitoba, 
there is found a large, robust and handsome member of the 
Ovales, It has five to ten spikes, which normally are well sep- 
arated, but plants with approximate or subapproximate spikes 
are not uncommon. It is the plant treated by Professor Fernald 
as Carex festucacea Scik. (Proc. Am. Acad. 37: 477. pL j, /. 
4^-48), and also so treated in Gray's Manual (7th ed., p- 221. /. 
363). To it is added in the treatments referred to a variety 
brevior (Dewey) Fernald, distinguished by having but three to 
six approximate or subapproximate spikes. 

Mackenzie: Notes on Carex — XI I 369 


of Britten & Brown's Illustrated Flora, I treated these two 
plants together, because the number of heads and their degree 
of separation was not a constant character. Later, in a paper on 
Carex straminea and its allies (Bull. Torrey Club 42: 603. 1916), 
it was pointed out that the genuine Carex festucacea was a differ- 
ent plant than either of the above, and the name Carex brevier 
(Dewey) Mackenzie was applied to what I had treated as Carex 
festucacea Schk. in the Illustrated Flora. 

In the meantime Kiikenthal (Pflanzenreich 4^*^; 207), notic- 
ing the close general resemblance of Fernald's Carex festucacea 
to Carex Bicknellii Britton (Carex straminea var, Crawei 



here under discussion has characters which are constant in a 
very considerable series of specimens. As compared with Carex 
brevier it has the perigynia appressed-erect instead of spreading- 
ascending. The mature perigynia are membranaceous and 
strongly pellucid -puncticulate. They entirely lack the firm 
coriaceous texture so characteristic of Carex brevier and are 
also more nerved than in that species. These differences are 
well brought out in Fernald's figures above referred to, even 
the difference in the texture of the perigynia coming out. 

The culms of the northern plant are much developed the 
first year of their growth, and the old dried leaves towards the 
base of the culm are therefore very marked in the flowering year. 
In Carex brevier on the other hand the development the first 
year is much less, and the dried leaves are short and rather 
inconspicuous in the flowering year. The sheaths too in the 
northern plant are white-hyaline ventrally for but a short 
distance, while in Carex brevior this feature is very strongly 
developed, and they also differ from those of the latter plant in 
being strongly papillate dorsally. As stated by Fernald the 
northern plant has five to ten spikes which normally are distinct 
and well separate, and Carex brevior usually has fewer spikes 
which are approximate or sub-approximate. However, tliis is 
but a well-marked tendency and specimens of the northern 
plant with a congested head or Carex brevier with spikes as 
widely separated as in the northern plant are not uncommon. 
Carex Bicknellii can be readily distinguished by its larger 

370 Mackenzie: Notes on Carex — XII 

(5-5-7-5 mm 

peng^^'nia, which are 

nerved ventrally, and very 


than IS Carex hrevior. 

members of the Ovales 
h America and the cha 

which separate them dates from Professor Fernald's revision 
of the group in 1902. The plant which I have been here dis- 
cussing IS one which he has collected on a number of occasions- 


It is therefore one to which it is highly fitting to give his name. 
Unfortunately there is a poorly understood Japanese species 
to which his name has been given. Under the circumstances I 
trust that I will be pardoned in proposing for the plant I have 



Carex Merritt-Fernaldii Mackenzie sp. nov, 

*^ Carex feshicacea Schkuhr" Fernald, Proc. Am. Acad. 37: 477. 

pL 3, f. 47-8. 1902. 

Cespitose from short-prolonged, lignescent, black-fibrillose 
rootstocks, the culms 3-10 dm. high, slender but stiff, sharply 
triangular above, terete towards base, strongly biennial, exceeding 
the leaves, smooth or very slightly roughened beneath head, 
conspicuously clothed at base with the short -bladed leaves of 
the previous year, the lower bladeless; leaves with well-developed 
blades three to six to a culm, on the lower third but not bunched, 
the blades erect-ascending, dull- or yellowish-green, firm, 1.5-3 
mm. wide, usually 1--2 dm. long, strongly roughened towards 
apex especially on the margins, the sheaths tight, white-hyaline 
ventrally for a short distance below mouth only, strongly 

IS with 1 

insertion of 

Spikes four to ten (usually six to eight), aggregated to separate, 
the head 1.5-8 cm. long, 7-15 mm. wide, the spikes gyiiaecan- 
drous, subgloboseto ovoid, 7-15 mm. long, 5-9 mm. wide, with 
fifteen to thirty appressed-ascending perigynia above (the beaks 
appressed-ascending), blunt at apex, abruptly contracted into 
the short clavate basal staminate portion; bracts scale-like, the 
lowest often prolonged, 1-4 cm. long, the upper merely acuminate 
or short-awned; scales ovate, obtuse to short-cuspidate, yellow- 
ish brown with three-nerved green center and narrow hyaline 
margins, shorter and much narrower than the perigynia above; 
perlgj-nia suborbicular, 4-5 mm. long, 2.5-3,5 mm. wide, con- 
cave-convex, thickish over the achene but the walls membran- 


Mackenzie: Notes ON Carex— XII 371 

aceous, strongly pellucid-puncticulate, yellowish-green or in age 
yellowish-white, strongly winged to base, serrulate to below 
middle, truncate rounded at base, strongly several to many 
ner\^ed dorsally, nerveless to three to five-nerved ventrally, 
often only towards base, the wings one- to two-nerved, abruptly 
narrowed into the flat, serrulate, reddish -brown tipped, obliquely 
cut dorsally, bidentate beak, about i mm. long and less than 
one-third length of body, the margins of the orifice reddish- 
brown; achenes lenticular, 1.75 mm. long, 1.5 mm. wide, sub- 
stipitate, apiculate, brown, shining; style slender, straight, 
jointed with achene, deciduous; stigmas two, reddish brown, 
slender, long. 

Type locality; Orono, Maine, Fernald, July 3, 1897 
(Type in the Gray Herbarium of Harvard Universitv") . 

Specimens examined 

Maine: Seal Harbor, Redfield, July 14, 1890 (H., N.E., P.), 
also Williams July 25, 1890 (N.E) ; Canton, Parlin 2021, August 
8, 1906 (H.); North Berwick, Parlin 798, July 15, 1897, also 
May 1892 (H.); Mt. Desert Islatid, Rand, July 9, 1893, also 
July 15, 1894 (H.); Basin Mills, Knight, July 15, 1904 (H.); 
Orono, Fernald, June i, 1899 (N.E.), July 14, 1890 (N.E.), 
July 12, 1897 (N.E.), July 2 and 3, 1897 (H.); Bumford Point, 
Parlin, July 4, 1890 (H.); Canton^ Parlin 1972, July 10, 1906 
(H.), also 2225, 1907 (H.); Tatnic, Parlin, July 11, 1891, also 
June 1892, (N. E., H.); Orono, Fernald & Long 12953, July 22, 
1916; (N.E., P.); Milford, Fernald & Long 12953, August 25, 
1916 (N.E., P.) ; Farmington, Knowlton, July 9, 1911 (N.E., P.) ; 
Woodstock, Pease 17126, July 11, 1918 (N.E.); Canton, Parlin 
2094, June 25, 1907 (N.E.) ; Dixfield, Parlin 2461, June 28, 1908 
(N.E.); Gilead, Pease 16013, July 5, 1914 (N.E.); Rumford 
Point, Parlin, July 4, 1890 (N.E.); Dennysville, Fernald 1444, 
July 19, 1909 (N.E.); Columbia, Knmvlton, August 4, 1916 
(N.E., P.); West Pembroke, Fernald 1452, July 6, 1909 (N.E.); 
Big Hill, Fernald 1438, July 17, 1909 (N.E.); Leeds, Knowlton, 
July 23, 1916, (N.E., P.); New Gloucester, Knowlton, July 25, 
1920 (N.E.); Baldwin, Fernald et al. 12956, August 30, 1916 
(N.E., P.); Kennebunkport, Deane, July 12, 1894 (N. E.); 
North Berwick, Parlin 798, July 15, 1897 (N.E.); Alfred, 
Fernald & Long, 12954, August 10, 1916 (N.E., P.); Wells, 
Fernald & Long 12952, August 8. 1916 (N.E.., P.); North Ber- 

372 Mackenzie: Notes on Carex — XII 

wick, Parlin & Fernald, June 13-14. 1896 (N.E.); Tatnic Hill, 
Parlin dt Fernald, June 13-14, 1896 (N.E.); Cape Porpoise, 

July, 1901 (N.E.) ; South Berwick, Parlin, Ju 
Hermon, Knight 5035, June 30, 1906 (K.M 

July 15, 1905 (K.M.). 

' HAMPSHfRE: Trov, Rand & Robinson 507 


July 8, 1914, also July 

July 30, 1915 (N.E.); 


Vermont: Mason Hill, Pownal, Churchill, June i8, 1901 


Massachusetts: Amherst, B. P. Clark, June 6, 1881 (P.). 

New York: Wellesley Island, Jefferson County, Robinson 
& Maxon j, June 28, 1902 (H.); Thousand Island Park, Ball, 
July 1904 (K.M.); Middle Grove, Saratoga County, Peck 
(Albany) ; East Galway , Saratoga County, E. A . Burt (Albany) ; 
West Ft, Ann, Washington County, Bnrnham, June 27, 1918 
(Albany) . 

Ontario: Lake Nipigon, John Macoun, July 11, 1884 (H.); 
Kingston, Khigh, June 30, 1906 (H.); ''Portage River, Lake 
Superior", ex herb, C. E. Smith, August 3, 1865 (P.)- 

Michigan: Washington, ex herb. Dewey; Herb. (H.); Kew- 
enaw County , Far well 762, August 1890 (H.); Muskegon, Mc- 
Clatchie48 (H,); Douglas Lake, Emmet County, Ehlers, 2g6, 
2gg, July 17, 1920, also L. E. Smith i, July 3, 191 7 and 159, 
Aug. 20, I9I7(K.M.); Clifton, Farwell, July 18, 1890 (Albany); 
Isle Royale Williamson, 22^6 August 15, 19 12 (P.). 

Manitoba: Lake Winnipeg Valley, Bourgeau 1857 (H.). 

* * * 


After the above was written, and when on a visit to the Phil- 
adelphia Academy of Sciences, Mr. Bayard Long asked me to 
look at the type specimen of Carex alboltUescens Schw. preserved 
there. It was apparent that it was a young and immature 
specimen of the genuine Carex siraminea Schk. as interpreted by 
me (Bull. Torrey Club 42: 605-6), a conclusion which Mr. Long 
had himself arrived at. The previous identification of th^ plant 

Mackenzie : Notes ON Carex — XII 373 


of Schweinitz with the plant treated in this article and generally 
as Carex albGlutescens can be readily understood. The two 
species really stand next to each other, and as long as genuine 
Carex straminea was not understood and kept separate, it was 
a natural thing to do to identify the plant of Schweinitz in the 
way that was done. The name, however, has always been 
inappropriate, and one is glad to be able to reduce it to synonymy. 
It now appears that the plant treated in this article and gen- 
erally as Carex albokitescens {''Carex albolutescens Schw."Kuk- 
enth. in Engler, Pflanzenreich 4^*^: 208; Mackenzie in Britton 
& Brown, 111. Fl. (2d., ed.) 1: 385./. Q41 [excluding var. cumtdata 
Bailey]; Robinson & Fernald in A. Gray Man. (7th. ed.) 219. 
P^' 35^ [left hand] excluding var. cumulata Bailey; ^' Carex foenea 
Willd. " Boott, 111. Carex 3 : 118, pL 375) is without a name. It 
therefore gives me much pleasure to be able to name it for that 
excellent field man and acute student, Mr. Bayard Long, as Carex 
Longii. A specimen collected July 24, 1907, by Mr. Long at Cold 
Spring, Cape May County, New Jersey, and preserved in the 
herbarium of the Philadelphia Academy of Natural Sciences is 
designated as the type. 

Stimulation of plants by carbon disulphide 


Carrie Oldenbusch 

The problem of stimulation of plants by minute doses of 
chemicals, which are poisonous in higher concentrations, has 
long been an interesting one. Raulin (1) in 1869 was probably 
the first to undertake work of this kind, using a nutrient solution 
made up of a large number of substances as a culture medium. 
As the stimulant, Raulin added to his nutrient solution small 
amounts of metallic salts, such as silver nitrate, platinum di- 
chloride, or copper sulphate, and found that cultures of Asper- 
gillus niger, grown In these solutions produced a more luxuriant 
mycelium than cultures to which nothing had been added. 
Pfeffer (2) in 1895, by means of comprehensive experiments, de- 
termined that such a complicated nutrient solution was unnec- 
essary and originated a number of solutions which have been 
and are still extensively used in work of this kind. Benecke 
(3), in the same year, published a paper in which he also gave a 
much simpler normal solution than Raulin's but slightly different 
from Pfeffer's. 

Richards (4), using the latter 's solution with the addition 
of traces of zinc, iron, sodium, lithium, and other salts, obtained 
a heavier growth of Aspergillus niger and Penicillium glaucum 
than in cultures to which none of the above were added, demon- 
strating that Raulin's solutions really resulted in stimulation. 
Ono in 1900 corroborated Richards's results and also worked 
out the effect of these salts on certain algae. He found that 
algae as a rule have their point of greatest stimulation at a much 
lower concentration of the salt but in other respects are similar 
to fungi. Richards (5) and Ono (6) determined that the econ- 
omic coefficient of sugar is considerably lower in the cultures 
to which salts had been added than in the controls, proving that 
stimulated fungi require a smaller amount of carbohydrate 
food in proportion to the quantity of waste acid produced. 

Since this time many other investigators have worked on 
this problem, obtaining similar results. Kahlenberg and True 
(7) formulated a law which they gave as an explanation of this 
stimulation phenomenon. They claim that since the chemical 
and physical properties of solutions are due to the properties of 


376 Oldenbusch: Stimulation of plants 

the ions dissociated in the solution and of the undissociated 
molecules, then the physiological properties of a solution are due 
to the same factors. The authors worked out this theory with 
lupine seeds, the germination of which w^as accelerated by dilute 
solutions of electrolytic salts. They claimed that this stimu- 
lation was due to the hydrogen and metallic ions in the solutions. 
Heald (8) working with seeds of other plants obtained similar 
results and also attributed them to the electrolytic dissociation. 
Stevens (9) working with fungus spores applies this same theory. 
Iwanoff (10) claims that metallic salts are poisonous to plants In 
proportion to the atomic weight of the metal, the heavier metals 
being more poisonous than those of smaller atomic weight. In 
his opinion, the same is true of the alcohols, the lower alcohols 
(methyl and ethyl) being less poisonous than those of higher 
molecular content, such as propyl and butyl. Steinberg (11), 
in a recent paper on the stimulation of Aspergillus ^ attributes the 
stimulation to the increase in H ion content when zinc salts are 
added to the nutrient solution. It is not obvious, however, 
that the action of such oligodynamic substances is due to the 
hydrogen ion concentration alone, especially since we find non- 
electrolytes, such as ether, chloroform, and other anaesthetics, 
exerting a stimulatory Influence, Carbon disulphide, being a 
very volatile liquid, which is non-dissociable in aqueous solution, 
also comes in this category. 

Various authors have worked with ether and chloroform , 



send (12) In his experiments with Avena seeds found that if 




than under normal conditions. This acceleration lasts for some 
time and gradually disappears. In a later paper (13) Townsend 
verified his results wath other seeds: Zea Mays^ Cuciirhita, 

Pliaseolus. etc. 


atmosphere of HCN gas (14), germinated more rapidly than 
those not exposed although this advantage did not last for any 
appreciable time. Miss Latham (15), working with Sterigmato- 
cystis nigra^ determined that cultures of the fungus grown on 
nutrient media and exposed to chloroform vapor, produced a 
more luxuriant growth with a greater dry w^eight than similar 

Oldenbusch: Stimulation of plants 377 

cultures unexposed. Furthermore the oxalic acid formation was 
lower per unit dry weight in the stimulated cultures. 

Johannsen (16) was the first to discover that exposnig dor- 
mant buds to an atmosphere hnpregnated with ether forced the 
buds to open earlier than normal. Chloroform has the same 
effect but is more powerful, less being needed to bring about the 


same result. 

No attempt has been made to cover the whole field of the 
very voluminous literature on this subject. A comprehensive 
account may be found in Czapek's ''Biochemie der Pflanzen/' 
second edition, Vol. i, page 147 el seq., 1913, 

Among the investigators who experimented with carbon 
disulphide, none used it in gaseous form but either in pure liquid 


state or in aqueous solution. Sirker (17) in Japan found that 
mulberry bushes, grown in soil previously treated with CS2, pro- 
duced more branches, higher plants and more leaves with a 
greater dry weight for each individual leaf than bushes grown in 
untreated soil. Koch (18) in 1912 found the same thing with 
other plants. Koch asserts that the carbon disulphide is not 
used as a carbon source, nor does it act as an insecticide in 
killing off the harmful lower organisms in the soil, since under 
such circumstances a larger amount of the CS2 would be more 
beneficial. This, however, is not the case, a larger amount of 
the CS2 being harmful, and the stimulation takes place only when 


the compound is added in minute quantities. Fred (19) in the 
same year stated that carbon disulphide added to the soil in 
dilute solutions stimulated the growth of soil bacteria, the nitro- 
gen-fixing forms in particular, so that the quantity of nitrogen 
built up into higher compounds was measurably increased over 
normal. In a later paper Fred (21) corroborated his earlier 
results and concluded that while after treatment with carbon 
disulphide the soil showed a reduction in the number of* micro- 
organisms, it was followed by a great increase and an increase 
also in the by-products of their action. With relatively strong 
applications of carbondisulphide to the soil, corn and oats 
seemed deleteriously affected, while mustard and buckwheat 
were benefitted, as shown by increase in dry w^eight. In acid 
soil, clover is also stimulated. 

The experiments described below, which further test the 
effect of carbon disulphide on plants, were performed in the 

378 Oldenbusch: Stimulation of plants 

Botanical Laboratories of Barnard College, under the super- 
vision of Professor H. M. Ricliards, for whose advice and assis- 
tance the writer is greatly indebted. 

■ In this work, while no attempt was made to cover every type 
or condition of plant tissues, a number of widely differing kinds 
of material were employed. The experiments fall into three 
groups, each of which could profitably be elaborated beyond 
what was attempted here. These three categories are as given 
below and will be considered as separate topics in the following 


1. Stimulation of seeds and seedlings. 

2. Stimulation of buds. 

3. Stimulation of certain fungi. 

Carbon disulphlde is a clear, practically colorless liquid when 
pure, very inflammable and volatile. It has a disagreeable odor, 
especially when impure, because of the decomposition into other 
sulphur compounds, hydrogen sulphide in particular (see Graham, 
20). It is soluble in all proportions in alcohol, ether, and chlo- 
roform, but only slightly soluble in water. The solubility in 
water, according to Graham, varies inversely as the temperature, 
as follows: 

At 12^ -13^ C. 2.0 gms, CSa dissolve in i liter of water. 

" 15° -16** " 1.91 *' '* '' '* I '' '' " 

^' 25° -27^ '' 1.68 " " '' . " I '' *' 

" 30** -ss"* " 145 " " '* " I " " 

From these data, it was estimated by interpolation that at 
20*^-22*^ C. (average room temperature) about 1.8 gms. CS2 
dissolves in i liter of water. 

In the following paper, the concentrations of the solutions 
used were determined in terms of molecular solutions. The 
molecular weight of carbon disulphlde is 76.125*, and if 1.8 grams 

dissolve In i liter of water at approximately room temperature, 
then a saturated aqueous solution is 0.0235 of a molecular solu- 
tion. More dilute solutions were prepared from this, in fractions 
varying by 1/2000M. No attempt was made to determine or 
maintain the exact concentration closer than this, because carbon 
disulphide is so easily vaporized and a certain amount of that in 
solution is continually volatilizing out. In cases where the ex- 
periments lasted over a few days, the solutions were renewed at 
intervals of two or three days. 

Oldenbusch: Stimulation of plants 379 

I. Stimulation of seeds and seedlings 



The first phase taken up was to determhie if there was any 
stimulation of young seedlings by dikite solutions of carbon 
disulphide. Townsend (12, 13, 14) tested the effect of ether 
vapor and hydrogen cyanide gas on different kinds of seeds, by 
soaking seeds in water for twenty-four hours and then putting in 
an airtight damp chamber containing ether vapor. The exposed 
seeds germinated more quickly than unexposed controls, but 
whether left in ether chamber or removed immediately after 
germination, the stimulated seedlings soon lost their advantage, 
and five days after germination the control seeds equaled the 
stimulated in growth. Likewise dry seeds exposed to HCN gas 
for a short time and then soaked in water germinated more 
rapidly than seeds not treated with the hydrogen cyanide. 
However in every case, growth was equal after a week. 

In the following experiments, wheat grains (Triticum sativum) 
were used. One hundred grains were soaked in the CS2 solution 
for twenty-four hours. Seed pans were filled with moist sphag- 
num, covered with damp cheese cloth, and grains planted out on 
top of this. They were then covered with a second layer of 
cheese cloth, with a pad of moist sphagnum on top of the whole. 
Pans were kept under a shelf in the greenhouse for three or four 
days, taking care not to allow the sphagnum to dry out. The 
seedlings were then uncovered, epicotyls measured and the 
average taken of those germinated. It was found that the 
saturated solution completely inhibited growth, while the seed- 
lings in the other solutions grew as shown in Table I. 

Average growth in millimeters of epicotyls of wheat seedlings after 

being soaked in following csg solutions for 24 hours 

and left to germinate for 3 or 4 days 

Series i. Germinated 4 days 
Exp. A. B. C. Average Avg, per day 


377 37.7 94 

M/4,000 39-7 35-2 38.5 37-8 9.4 

M/6,000 40.6 29.2 34-9 87 

M/8,000 34-8 311 4L3 39-1 9.8 

M/i2,ooo 38.3 34.1 36.2 91 

M/i6,ooo 42.7 33-2 37-6 37.8 9.5 

M/22,000 41.2 28.9 434 37-8 9-5 

M/32,000 40.0 36.6 394 387 9-7 

M/40,000 41-5 41-5 10.4 

Control 35-8 33-0 36.6 35,1 8-8 

38o Oldenbusch; Stimulation of plants 

TABLE I (con,) 

Series II. Germinated 3 days 


Exp. D. E. Average Avg. per day 

M/2,000 13.6 lo.o 1 1.8 3.9 

M/4,000 13.3 13-8 136 4-5 

M/8,000 16.8 14.5 157 5-2 

M/i6,ooo 14.7 13-7 14-2 4-7 

M/22,000 14.9 14-2 14-6 4-9 

M/32,000 18.7 14.4 16.6 5.5 

M/40,000 12.7 15.3 14.0 4.7 

Control 1 1,1 13.6 13.3 4.4 

Although these results show some irregularities due to the 
comparatively small number of cases, they also reveal a marked 
stimulation of growth especially in the solutions between 

M/22,000 and M/4' 



four day instances, showing that the stimulation tends to 
diminish, and growth tends to approach the normal after an 
interval, just as Townsend found. 

An experiment with Vicia Faba seedlings w^as carried out in 
the following manner. The seeds were planted and allowed to 
germinate until the hypocotyls reached a convenient length to 
measure. A number 400 cc. bottles were almost filled with 

^_ \ 

Knop's nutrient solution and enough CS2 solution was added to 
make of desired concentration. The bottles were then fitted with 
corks, each cork having three holes bored in it, which were just 
small enough to prevent the seeds from falling through. The 
seedlings grown as above, were then washed in distilled water, 
hypocotylsm easured, and arranged in culture bottles so that 
seed rested on top of cork and hypocotyl extended through the 
hole into the solution. The bottles were placed in a dark 
chamber in the green house (temoerature varvine from 18° to 

and the daily growth computed. 


Oldenbusch: Stimulation of plants 381 





M/2,000 M/4,000 M/6,ooo M/8,ooo M/20,000 M/40,000 Control 
Exp. A. 


istday 5.3 6.5 13.0 13.7 7.0 

2nd " 2.2 5.4 n.7 16.1 9.5 

. Total 7.5 1 1.9 24.7 29.8 16.5 

Avg, 3.8 6.0 12.4 14.9 8.3 

Exp. B. 

1st da}- 18,2 18.2 22.7 22.9 17.8 18.9 

2nd " 194 23.4 21.6 21.0 16.2 19.6 

3rd " 21.4 19.0 19.7 15.8 8.1 14.4 

4th " 19.2 19.2 23.5 17.0 17.7 16.3 

Total 78.2 79.8 87.5 767 59.8 69.2 

Avg. 19.6 20.0 21.9 19.2 15.0 17.3 
















16.6 19.3 

16.0 17.5 

15-5 197 

48.1 56.5 

16.0 t8,8 

Exp. C. 

1st day 16,6 14.6 

2nd " 10.5 16.0 

3rd " 14.1 18.4 

Total 41.2 49.0 

Avg. 13.7 16.3 

Exp. D, 


1st day 9.5 16.8 10,8 20.0 39.0 1.5 19.8 

2nd *' 7.0 II. 5 ii.o 18.0 12.5 II. o 16.5 

3rd " 8.0 14.0 12.9 ' 24.5 18.5 13.2 17,0 

Total 24.5 42.3 34.7 62.5 70.0 25.7 53.3 

Avg. 8.2 14.1 II. 6 20.8 23.3 8.6 17.8 

Total average of the four experiments (i.e. twelve examples). 

8.6 14.0 15.2 21.4 19,6 13.2 15.6 

Table II shows a distinct stimulation in the growth of 
hypocotyls of Vicia seedlings, in the M/6,000 to M/20,000 CSx 

solutions, the average greatest stimulation being in the M/8,000 
dilution, although the M/20,000 shows a greater acceleration in 
some instances. In the M/40,000 concentration, the CS2 content 
has become so dilute that it no longer has any effect on growth. 
In the higher concentrations, the M/6,ooo solution caused a 
marked stimulation in Exp. A and B, although in other cases 

382 Oldenbusch: Stimulation of plants 

growth is not as great as normal in these dilutions. Although 
the M/4,000 and M/2,000 solutions do not entirely inhibit 
growth, they do retard it below normal. Complete inhibition 
was found in the saturated CS2 solution, but no dilutions were 
tried between that and the above. 


Koch (18) tried similar experiments with Helianthus annuus 
and Ciicurhita Pepo seeds, letting the seeds germinate and then 
placing them in the dark, with their hypocotyls In solutions 
containing o.i and 0.05 cc. CS2 to looo cc. water for thirteen 
to fourteen hours. In one case only, in the solution containing 
0.05 cc. CS2 to tJie liter(about M/i 50 solution), did he obtain 
growth higher than normal. This being only a single instance 
was probably an accident due to the solution being weaker 
than intended and cannot in any sense be called a stimulation. 
Koch did not try any solutions weaker than thi^ and therefore 
did not reach the stimulatory region but really only determined 
that carbon disulphide acted as an inhibitory agent. 

2. Stimulation of dormant twigs 

+ I 


Johannsen (16), investigating the effect of ether on dormant 
twigs, found that they were stimulated so that the buds opened 
sooner and developed more rapidly when exposed to ether vapor 


for twenty-four to forty-eight hours, than when unexposed. 
When using chloroform a much smaller amount was neces- 
sary to produce the same result (9 grams of chloroform giv- 
ing the same acceleration as 40 grams of ether). 

With these results in view, experiments were carried out to 
see if vaporized carbon disulphide would stimulate buds to open. 
Willow twigs were cut from trees outdoors which had been 
subjected to winter conditions and were probably frozen in 
spite of the general mildness of the winter. The lower three or 
four inches of the stem were placed in a bottle of water and a 
large bell-jar put over them. Pieces of filter paper, moistened 
with 10 cc. of carbon disulphide were quickly inserted under the 
bell-jar. The whole w^as placed in a pan of water making a 
water seal so that the carbon disulphide, which vaporizes almost 
immediately, could not escape from the bell-jar. The twigs 
were left in this atmosphere (approximately saturated with CS, 
gas) for periods ranging from ten minutes to eighteen hours. 
They were then removed, the stems were put into fresh water 
and with a control placed in the greenhouse. 

Oldenbusch: Stimulation of plants 383 


as follows. After three days, the twigs exposed ten and twenty 
minutes began to develop, the buds showing patches of green 
at the tips. After seven days, the buds on the twigs exposed 
thirty minutes were open considerably. One of the hour experi- 
ments showed slight greening of the upper buds but other speci- 
mens gave no evidence of development. After nine days, the 
control (not exposed to CS2 at all) had two buds developing. 
Twigs exposed ten, twenty and thirty minutes continued to show 
better bud development for two to three weeks longer, after which 
the condition of all tended to become the same. Twigs exposed 
for one and a half hours or for two hours started to open but 
after a day or so became brown and withered, while those 
exposed for four hours or more showed no signs of development 
at any time, indicating that the protoplasm itself must have 
been killed. 

3- Stimulation of certain fungi 

As stated above, there has been a vast amount of work done 
on stimulation of fungi by metallic salts and other chemicals. 
Because of this, a number of experiments were set up to test the 

effect of carbon disulphide on fungi. 

To determine the effect of dilute solutions of carbon disulphide 
on the germination of spores of Rhizopus nigricans and Slerig- 
matocystis nigra, hanging drop slides were used. Small glass 
rings were attached to microscope slides by means of a solution 
of dextrin in water (vaseline is not practicable since carbon 
disulphide is very soluble in it). The medium used for the ger- 
mination of the spores was prepared according to the following 
formula (see Richards, 4): 

Potassium pho phate,,, 0.5 gm. 

Magnesium sulphate 0,25 gm. 

Ammonium nitrate -, \ I-O gm- 

Ferrous sulphate 

5.0 gms 

Distilled water 100 cc. 

To this was added 5 gms. of gelatine, and the mixture was 
then heated in water bath. When the gelatine was completely 
solved, the medium was filtered tubed (10 cc. to the tube) and 
sterilized. Before the gelatine hardened, the CS, solution was 
added to make of desired concentration. 


Oldenbusch; Stimulation of plants 

A small drop of this gelatine was put on a clean cover glass 
and inoculated with fungus spores. After inverting the cover 
glass, and sealing on to the glass ring the slides were left standing 
at room temperature under a bell-jar containing moist filter 
paper to present slides from drying up. After some hours, 
slides were examined microscopically for germination, and where 
possible the germination-tubes were measured by means of a 

standardized micrometer eye-piece. 

According to Table III, solutions of carbon disulphide in 
nutrient gelatine varying from M/i 6,000 to M/40,000 in con- 
centration, stimulate spores of Rhizopus nigricans and Sterig- 
maiocystis nigra to germinate sooner than in the same media 

but without the CS2. The point of greatest stimulation lies 
between the M/32,000 and the M/40,000 solutions. 

In another type of experiment with Sterigmatocystis spores, 
the stimulation of growth was measured by the increase in dry 


Average growth in microns of 10 germination tubes of fungous 

nutrient media to which cs2 had been added 

Rhizopus nigricans 

SrORES, tnoculated 

Exp. A. , 

No, of M/800 M/8,000 M/i6»ooo M/22,000 M/32,000 M/40,000 Control 




1 1.9 


Exp. B. 



Exp, C. 











57.4 over 300 

17 over 300 

over 300 
If II 

over 300 

« 4( 

Exp. D. 

Exp. E, 


281. 1 


over 300 over 300 



148. 1 

127.4 over 300 

over 300 over 300 over 300 250, 






over 300 


■ t 









over 300 

over 300 over 300 

over 300 over 300 
II 11 II II 












over 300 45.9 











Oldenbusch: Stimulation of plants 


TABLE III (con.) 

Sterigmatocystis nigra 
Exp. A. 

No. of 

hours M/800 M/8,000 M/i6,ooo M/22,000 M/32,000 M/40,000 Control 


Exp. B. 



Exp. C. 


. 25 


Exp. D. 




over 300 

over 300 




over 300 over 300 224,9 











over 300 over 300 over 300 


over 300 

41 (I 





over 300 


over 300 

1 17-0 


over 300 over 300 

<i i< <l <l 


















over 300 

41 <l 


over 300 

over 300 




over 300 


weight of the germinating hyphae. In their experiments 150CC. 
Erlenmeyer flasks were used as cuhiire bottles and^ with pipettes 
and other apparatus, were cleansed with cleaning solution, rinsed 
in tap water a number of times, then in distilled water and finally 
in double distilled water. The same solution as in the former 
experiment, but without the gelatine and with double distilled 
water was employed as a culture medium. The trace of ferrous 
sulphate was introduced by adding one drop of a saturated 
solution of ferrous sulphate in double distilled water. 

After the nutrient solution was made up, it was inoculated 
in bulk with the Sterigmatocystis spores, and 50 cc. of the inocu- 
lated solution pipetted into each flask. The aqueous CS2 
solution (also prepared with double distilled water) was added 
last. The flasks were then corked by means of a mercury air 
trap, which would allow air to come into the flask, but would not 
allow any gas to escape from the flask and thus change the CSa 

386 Oldenbusch: SxiMtrLAXioN of plants 

After being stoppered, the flasks were left in a dark chamber 
at room temperature and, as a preventive measure, the controls 
were put in a different compartment from those to which carbon 
disulphide had been added. After five days, the flasks were 
removed, pads filtered off on weighed filter paper and dried in 
hot air oven at 70*^ to dry weight. No attempt was made to 
determine the COa or acid production, although this would have 
been an interesting point to investigate. 


Although growth was not wholly normal in the controls, 
presumably due to lack of adequate circulation which may have 
caused carbon dioxide to accumulate, nevertheless the gross 
appearance of the stimulated was strikingly different from that of 
control specimens. The solutions containing CS2 in the most 
optimum concentrations bore tough pads, thicker and con- 
siderably heavier than normal. All the stimulated pads were 
wrinkled and curled, and slightly yellow on the under surface. 
Practically no spores were produced, although in some cases a 
few yellow sporangia were observed. The control felts on the 
other hand were thin and tender in texture, and more or less 
covered with . black spores. However, if further experiments 
were to be carried out, a more perfect system of maintaining the 
concentration of CSa in the flasks would have to be devised in 
order to obtain absolutely normal controls, 



Dry weight in milligrams of cultures of Sterigmatocystis nigra. 


Exp, Control. M/80,000 M/40,000 M/20,000 M/io,ooo M/5,000 

No CSt 

A 439 838 800 740 585 270 

B 442 1028 815 748 675 370 

C 530 765 1035 495 490 

D 578 788 740 810 540 

Avg. 497 855 848 698 573 320 

The results of Table IV are sufficient to Indicate that there 

Is a marked stimulation in cultures to which carbon disulphide 
has been added. The region of greatest stimulation Is in the 
flasks M/8o,ooo and M/40,000 CS^ solutions, the more dilute 
concentration being slightly more advantageous than the other. 

Oldenbusch: Stimulation of plants 387 

Complete inhibition was not found in any of the experiments 
tried, although the flasks containing M/5,000 solution only pro- 
duced felts of approximately three-fourths of the normal weight. 

In some instances the M/io,ooo acted slightly as an inhibitor so 
that growth below normal was produced. 


All things considered, the results of this investigation lead 
one to conclude that carbon disulphlde acts on plants as many 
other chemical stimulants do. When applied either in aqueous 
solution or in vaporized condition it inhibits growth, if given in 
too concentrated a solution or if vapor is used for too long a time. 

When used in dilute amounts, carbon disulphide has the 
ability to stimulate: 

« ■ 

(i) Dormant protoplasm, as in the case where twigs exposed 
to CSj vapor for a short period of time quickly resumed their 
growth ; 

(2) Active protoplasm, as in case where the hypocotyls of 

Vicia seedlings elongated more rapidly upon the addition of 

small amounts of CS 
(3) Conidial spon 



and produced germination tubes elongating more rapidly, the 
stimulation causing also causes an Increase in dry weight. 

Carbon disulphide acts also as other stimulants do, in that the 
acceleration of growth lasts only for a certain period of time, 
and then growth tends to reach normal again. This fact was 
shown in the twig experiments and those with Triticum grains. 
The same would probably be true of the other experiments if 
left running for a sufficient length of time. 

Botanical Department, 
Barnard College, 
Columbia University, 
New York City 

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1 920- 1 922 

The aim of this Index is to include all current botanical literature written by 
Americans, published in America, or based upon American material; the word 
America being used in the broadest sense. 

Reviews, and papers that relate exclusively to foresty, agriculture, horti- 
culture, manufactured products of vegetable origin, or laboratory methods 
are not included, and no attempt is made to index the literature of bacteriology. 
An occasional exception is made in favor of some paper appearing in an Ameri- 
can periodical which is devoted wholly to botany. Reprints are not mentioned 
unless they differ from the original in some Important particular. If users 
of the Index will call the attention of the editor to errors or omissions, their 
kindness will be appreciated. 

This Index is reprinted monthly on cards, and furnished in this form to 
subscribers at the rate of three cents for each card. Selections of cards are 
not permitted; each subscriber must take all cards published during the term 
of his subscription. Correspondence relating to the card issue should be 
addressed to the Treasurer of the Torrey Botanical Club. 

HoUick, A. Paulownia in winter. Jour. New York Bot. Gard. 

23: 1-3. pL 167 y 268. Ja 1922. 

Holm, T. Contributions to the morphology, synonymy, and 

geographical distribution of arctic plants. Rep. Canad. 
Arctic Exped. 1913-18. 5^: 1-139. 10 F 1922 [Illust.], 


Holm, T. Seasonal dimorphism in Arisaema triphylhim. Am. 

Midi. Nat- 8: 42-48. /. 1-5. Mr 1922. 

Holm. T. Studies in the Cyperaceae XXXI 1 1. Am. Jour. Sci. 
3: 138-144. /. J-72. F 1922. 

House, H. D. Nomenclatorial notes on certain American plants 

II. Am. Midi. Nat. 8: 61-64. Mr 1922. 

Howe, M 

Algae. J 

Bot. Gard. 23: 23-24. F 1922. 


Hughes, D. K. The " serrato-ciliata" group of Tropaeolum. 

Kew Bull. Misc. Inform. 1922: 63-85. /. 1-4. 1922. 

Includes 16 spp. nov. 

Hunter, M. R. The present status of Scolopendrium in New- 
York State. Am. Jour. Bot. 9: 28-36. /. i, i. 2i F 1922. 



392 Index to American botanical literature 

Illick, J. S. The maples. Am. For. 28: 12-19. Ja 1922. 


Illick, J. S. What our Christmas trees are. Am. For, 27: 739" 

749. D 192 1. [Illust.] 

Illick, J. S. The sycamores. Am. For. 28: 145-150- Mr 1922. 


Illick, J. S. Woody plants of Franklin County, Pennsylvania. 

Pennsylvania Dept. For. Bull. 21: 5-42. I Ja 1921. 

Jackson, B. D. Thomas Nuttall (1786-1859). Jour. Bot. 60: 

57. 1922. 

Note, supplied by Mr. F. R. Dixon-Nuttall, as to late residence and 
burial place of Thomas Nuttall. 

Jepson, W. L. The giant Sequoia. In Hall, A. F., Handbook 
of Yosemite National Park [California], 235-246. pL 22, 23. 
New York. 1921. 

Joffe, J. S. Studies of sulfur oxidation in sulfur-floats-soil 

mixtures. Soil Sci. 13: 107-118, F 1922. 

Johnson, J. The relation of air temperature to certain plant 

diseases- Phytopathology 11: 446-458. pi. 21-^3. 25 F 


Jones, D. F. Hybridization in plant and animal improvement. 

Sci. Mo. 14: 5-23. Ja 1922. [Illust.] 

Jones, D. F. The indeterminate growth factor in tobacco and 

its effect upon development. 
S 1921. 



differences. Science II. 55: 348, 349, 31 Mr 


Jones, F. R., & Vaughan, R. E. Anthracnose of the garden pea. 

Phytopathology 11: 500-503. pi. ^5+/. 1-2. 21 Mr 1922. 

Judd, C. S. Kilanea National Park trees. Hawaiian For. & 

Agr. 18: 255-260. D 1921. [Illust.] 

Kaiser, G. B. Little journeys into rnossland. V. The old genus 

llypnum. Br>ologist 24: 58-60. 9 Ta 1922. 

Index to American botanical literature 393 

Kauffman, C. H. Studies in the genus Inocybe I. The species 

of Inocybe in Peck's collections. New York State Mus. 
Bull, 1919: 43-60. My-Je 1920. 

Kelly, J. P. Tlie synthesis of full coloration in Phlox. Science 

IL 55: 245. 3 Mr 1922. 

Kindle, E. M. Notes on the forests of southeastern Labrador. 

Geog. Rev. 12: 57-71. /. /-//. J a 1922. 

Klughy A. B. A correction concerning the life zones of Canada- 

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Knowlton, C. H., Ripley, W. S., & Weatherby, C. A. Third 

report of the committee on floral areas. Rh'odora 23 : 209- 
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Preliminary lists of New England Plants, XXVIIL 

Knowlton, H, E. Studies in pollen, with special refeience to 

longevity. Cornell Univ. Agr. Exp. Sta. 52 : 75 1-793. 
Ja 1922. 

Knudson, L. Non-symbiotic germination of orchid seeds. Hot. 

Gaz. 73: 1-25. /. i-j. Ja 1922. 

Korstian, C. F., & Fetherolf, N. J. Control of steam girdle of 

spruce transplants caused by excessive heat. Phytopath- 

ology 11: 485-490. /. 7-j. 21 Mr 1922. 



Minnesota. Univ. Minnesota 

/. i-j8. S 1 92 1. 

Krause, K. Rubiaceae peruvianae novae. Notizbl. Bot. Cart. 


103. I Ap 1922. 

Includes spp. nov. in Palicourea (l), Psychotria (2), Cepltaelis (i), 
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Krause, K., & Hoehne, F. C. Contribugoes conheclniento das 

Rubidceas do BrasII meridional. Anex. Mem. Inst. Butan- 

9-33. pi. 1-6. 1922 

Includes 6 new species. 


peppermint. J 

Chem. 50: 31-34- Ja 19^2 

394 Index to American botanical literature 

Krieger, L. C. C. Schizophyllum commune with a stipe. Myco 
logia 14: 47, 48, 6 Mr 1922. 

La Rue, C. D., & Bartlett, H. H. A demonstration of numerous 
distinct strains within the nominal species Pestalozzia 
Guepini Desm. Am. Jour. Bot. 9: 79-92. 20 Mr 1922. 

Lesdain, M. B. de. Lichenes prope Habanam in insula Cuba 

anno 1914 a cl. fratre Ars^ne Brouard lecti. Bryologist 24: 
68, 69. 15 F 1922. 

Levine, M. Studies on plant cancers. III. The nature of the 
soil as a determining factor in the health of the beet, 
Beta vulgaris, and its relation to the size and weight of the 
crown gall produced by inoculation with Bacterium tume- 
jaciens. Am. Jour. Bot. 8: 507-525. 15 F 1922. 

Lewis, 1. F., & Taylor, W. R. Notes from the Woods Hole 

Laboratory, 1921. Rhodora 23: 249-256. pL ijj+f, /, 2. 
27 F 1922. 

Includes i comb. nov. 

Loeb, J. Quantitative laws In regeneration. IIL The quanti- 
tative basis of polarity in regeneration. jWr. Gen. Physiol. 
4: 447, 461. /. 1-8. 20 M 1922. 

Experiments with Bryophyllum caJycinum. 

Laftfield, J. V. G. The behavior of stomata. Carnegie Inst. 
Washington Publ. 314: 1-104. pL 1-16 + /. 1-54. 1921. 

Ludwig, C. A. The control of angular leaf spot of cotton. 
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Macbride, T. H. Some of the ways of the slime mould. Mycolo- 

gia 13: 329^334- 21 Ja 1922. 



and P. apocrypta. Mycologia 13: 315-322. /. 1-4. 21 J 


Martin, G. W. Rhizophidium polysiphoniae in the United 

States. Bot. Gaz. 73: 236-238. /. i-io. Mr 1922. 

Maxon, W. R. Notes on American ferns— XVI 11. Am. Fern 
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Index to American botanical literiture 395 

McAlpine, D. Bitter pit in apples and pears: the latest results 

in preventive measures. Phytopathology 11: 366-370. 
27 Ja 1922. 

McDougall, W. B. Mycorhizas of coniferous trees. Jour. For. 

20: 255-260./. i-j. Mr 1922. 

McDougal, W. B. Symbiosis in a deciduous forest. L Bot. 

Gaz. 73: 200, 212. /, j-j. M 1922. 



McKinney, H. H., & Johnson, A. G. Wojnouncia graminis (Mc- 

Alp.) Sacc. & D. Sacc. on wheat in the United States. 
Phytopathology 11: 505, 506. 21 Mr 1922. 

Meade, R, M. Positions and movements of cotton leaves. 

Jour. Hered. 12: 444-448./. 7-g. D 1921. 


Meister, G. K. Natural hybridization of wheat and rye in 

Russia, /our. Hered. 12: 467-470. D 1921. 

Merl, E. M. Biologische Studien uber die Utriculariablase. 

Flora 115: 59-74. /• -^"J- 1922. 

Millar, W 

Western Canada. J 

For. 20: 10-17. Ja 1922. 

Moore, B. Humus and root systems in certain northeastern 

forests in relation to reproduction and competition. Jour. 
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Moore, B. Influence of certain soil factors on the growth of 

tree seedlings and wheat. Ecology 3 : 65-83. II /. 1-6, Ja 

Mulford, F. L. Broad-leaved evergreen shrubs for the South. 

Am. For. 28: 99-104. F 1922. [Illust] 

Murphy, P. A. The sources of infection of potato tubers with 

the blight fungus, Phytophthora infestans. Sci. Proc* 

pL 11-14. Au 1 92 1. 

Experinientss conducted in Canada. 

Murrill, W. A. A fragrant polypore. Mycologia 14: 46, 47 

6 Mr 1922. 

Trametes suaveolens. 

396 Index to American botanical literature 

Murrill, W. A. Hygrophorus caprinus. Mycotogia 14: 48, 49. 
6 Mr 1922. 

Murrill, W. A. Illustrations of fungi— XXXIII. Mycologia 

14: 25-29. pi. 1-9. 6 Mr 1922. 

Oliveira, A. A. Metodos practlcos para el cultivo de la yerba 

mate en el terrltorio de mlsiones. Bol. Minis. Agric. 26: 
395-402. O-D 1 92 1. 

Overholts, L. O. Diagnoses of American porias. — T. Mycologia 

14: i-ii, pU i-i-f. i~6. 6 Mr 1922. 

Palm, B. T. The false mildew of tobacco introduced Into the 

United States from the Dutch East Indies. Phytopathology 
11: 430-432. 16 F 1922. 

Palmer, E. J. The forest flora of the Ozark region. Jour. 
■ Arnold Arbor. 2: 216-232, Ja 1922. 

Parks, H. E. California hypogaeous Fungi — Tuberaceae. 

Mycologia 13: 301-314. 21 J4 1922. 

Paton, J. B. Pollen and pollen enzymes. Am. Jour. Hot. 8: 

471-501. 15 F 1922. 

Payson, E. B. Monograph of the genus Lesguerella. Ann. 

Missouri Bot. Gard. 8: 103-236. /. 1-34. 9 Ja 1922. 

Pearson, W. H. Ricciocarptis natans (L.) Corda, from Africa. 

Bryologlst 24: 69, 70. S 1921. 15 F 1922. 

in, L. J. Epiphyllous plants of certain regions ir 
Bulb Torrey Club 49: 1-14. ph i -^f, i. 21 M 


Peters, R. Moisture requirements of germinating seeds. 

Kansas Univ. Sci, Bull. 13: 23-37. /. J, 1, My 1920. 


Petzke, E. La Fourcroya giganlea (una planta textil). Bol 



Pfeiffer, H. Conspectus Cyperacearum in America meridional! 
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Piche, G. C. The forests of Quebec. Jour. For. 20: 25-43. Ja 


Index to American botanical literature 397 

Piper, C. V. The identification of Berberis Aguifolium and 

Berberis repens. Contrib. U. S. Nat. Herb. 20: 437-451. 
pL 24. 1922. 

Piper, C- V. An unusual type of proliferation in Agropyron 

cristatum. Jour. Hered. 20: 423./. 20, N 1921. 

Pittier, H. New or noteworthy plants from Colombia and 

Central America— 8. Contrib. U. S. Nat. Herb. 20: 453- 
490. pL ^/-jo. 1922. 

Includes new species in Virola (i), DialyantJiera (i), PUhecollobium (10), 
Machaerium (9), Protium (2), ZaniJioxylum (3), Eurya (i), Lucuma (2), 
Sideroxylon (i), Vitex (4), Calycophysum (2). 

Pittier, H. On the species of Dalbergia of Mexico and Central 

America. Jour. Washington Acad. Sci. 12 : 54-64. 4 F 1922. 

Includes 8 new species. 

Plitt, C. C. A preliminary report, with notes, on the lichens 

found near the Cinchona Botanical Station, Jamaica, 
British West Indies. Bryologist 24: 60-64. 9 J^ 1922; 24: 

70-74- 15 F 1922. 

Poole, R. F, Some recent investigations on the control of 

Sclerotinia Libertiana on the green house on the muck 
forms of Bergen County, New Jersey. Phytopathology 
12: 16-20. /. 1-3. 10 Mr 1922. 

Popenoe, W. ' The Andes berry. Jour. Hered. 12: 387-393. 
fro7itispiece + /. /-J. N T921. 

^ Ruhus glaucus. 

Popenoe, W. The frutilla or Chilean strawberry. Jour. Hered. 

12: 457-466./. 13-17. D 1921. 

Popenoe, W. The native home of the cherimoya. Jour. Hered. 

12: 331-336. 20 F 1922. [Illust.I 

Prince, G, H. Forestry administration in New Brunswick. 

Jour. For. 20: 54-61. Ja 1922. 

Pritchard, F. J*, & Porte, W. S» Effect of fertilizers and lime on 

control of tomato leaf spot (Septoria Lycopersici). Phyto- 
pathology 11: 433-441. /. 1-16. 25 F 1922, 

398 Index to American botanical literature 

Pulling, H. E. Biophysics as a point of view in plant physiology. 

Am, Jour. Bot. 9: 37-46. 21 F 1922. 

Ramsey, F. T. The Swazey barberry. Jour. Hered. 20: 426, 

427. /. 24. N 1921. 

Berberis Swazeyi 

Rayner, M. C. Nitrogen fixation in Ericaceae. Bot. Gaz. 73: 

226-235./. 1-4. M 1922. 

Record, S. J. Boxwoods of commerce. Bull Torrey Club 47: 

297-306. /. J. 31 D 1922. 

Record, S. J. Notes on woods. Science IL 55: 267-269; 10 

Mr 1922. 

Rehder, A. New species, varieties, and combinations from the 

herbarium and the collections of the Arnold Arboretum. 

Jour. Arnold Arbor. 3: 11 

Includeo i sp. nov. in Prunus, 


Reiche, C, Rasgos biologicos del abrojo. Mem. y Rev. Soc. 

Cien. 40; 41-44. pL i. O 1921. 

Opiintia tunicata. 

Renacco, R. Sobre algunas enfermedades parasitarias de las 

plantas cultivadas. Anal. Soc. Cien. Argentina 81: 62-70. 
Ja-F 1916. 

Round, E. M. A modern plant fossil. Bull. Torrey Club 49: 

63, 64./ J. 12 Ap 1922. 

Sargent, C. S. Notes on North American trees — IX. Jour- 
Arnold Arbor. 3: i-ii, Jl 1921. 

Includes u spp, nov. in Crataegus, 

Sears, P. B. Variations in cytology and gross morphology of 

Taraxacum. Bot. Gaz. 73: 308-325. pL p, 10. Ap 1922. 

P. B. Variation in cytology and gross morphology of 

Taraxacum. IL Senescence, rejuvenescence, and leaf var-. 

iation in Taraxacum. Bot. Gaz. 73: 425-446. /. /-p. 
Je 1922. 

rer, F. J. Phyllostictales. Phyllostictaceae (pars). North 
American Flora 6; 1-84. 5 Ap 1922, 



New names and the final members of new combinations are fn bold face type. 

Abutjlon incanum, 246; parvulum, 37 

Acacia constricta, 244; Greggii, 244 

Acerates lanuginosa, 98; viridiflora, 98 
Acer "" • 

Negundo, 293 ; rubrum, 1 65 ; 

saccharinum, 293 
Achyronychia, 351; Rixfordii, 351 
Acrolasia albicauHs, 184; dispersa, 184; 

humilis, 183; latifolia, 184 
Acuan Jamesii, 245 
Aecidium Yuccae, 195 
Aerodiclidium benense, 262 
Agaricus campanulatus, 232; Carbon, 

232; equlnus, 232; multifidus, 231; 

papiHonaceus, 232; Sajor-Caju, 230; 
striatus, 231; varius, 232 
Agave, 32; Parryi, 239, 242; utahense, 

Agropyron, 104; dasystachum, 105; 

-' 4^,-^^»-*»w, «. V'^ I ^A««^ k.j^<, *^,JI 

tenerum, 105 

Albizzia lophanta, 51 

Aleurina, 19; elastica, 21; fuscocarpa, 
16; marchica, 16; oUvacea, 20 

Algae in the Chinese classics. Refer- 
ences to the, 297 

Algae, ^ New British and American 

species of Lobomonas; a study in 

morphogenesis of motile, 123 
Allionia, 351; Brandegei, 351; nycta- 

ginea. 351; ovata, 35^; pumila, 351 
Alsophila, 3 

Alternaria, 332; Solani, 314, 323, 333- 

Amaranthus paniculatus, 243 
Amauria dissecta, 359 
Amauriopsis dissecta, 359 
Ambrosia, 53 

American fossil hepatic. A new, 207 
American species of Lobomonas: 
study in morphogenesis of motile 

algae, New British and, 123 
Amoeba,^ 133 135 
Amsinckia intermedia, 100; lycopsoides, 

100; Menziesii, 100, 239, 247 

Anderson, Flora, The development 

of the flower and embryogeny ot 

Martynia louisiana, 141 
Andropogon saccharoides, 241 
Anemone, 240; tuberosa, 35; spheno- 

phylla, 239, 243 

Anislotus brach^xarpus, 240, 245 ; 
puberulus, 239, 245; trispemius, 24b, 

Antennana alplna, i8i 

Anthericum, 54 

Anthol>jza, 55 

Antirrhinum antirrhiniflorum, 248 

Aphanolejeunea, 4, 5, 7; sicaefolia, 5 

AplopappusGooddingi,44; gracilis, 44 

Apocynum, 51 

Arabis perennans, 244 

Aragallus monticola, 179 

Arenaria Burkei, 179; congesta, 179; 

lithophila, 179; subcongesta, 179 
Aristida adscensionis, 242; bromoides, 

242; divergens, 242; Wrightii, 242 
Aristolochia, 51; fimbriata, 51; Sipho, 

Arizona, Midwinter botanizing in 

southern, 237 
Artemisia albula,25o; Brittonii, 250; 

pabularis, 102 

Arthur, Joseph Charles, New species 

ol Uredineae — XIV, 189 
Asclepias eriocarpa , 355 ; eriocarpa 

microcarpa, 355; linaria. 247; pum- 

ila, 98; speciosa, 98; syriaca, 97, 

98; verticiliata, 98 
Ascobolus, magnificus, 304 
Ashe, W- W., Notes on trees and shrubs 

of the southeastern United States, 

Aspergillus, 328, 376; nigcr, 375. 
Asphodelus, 54 
Aspidium simu latum, 63 

Asplenium, 6, 8 

Astragalus Nuttallianus, 245; parvi- 

florus, 101 

Atnplex canescens, 238, 243; hastata, 

97; patula, 97; rosea, 243 
Audibertia capitata, 37 
Auricularia reticulata, 228 
Avena, 376 
Ayenia pusilla, 246 


Baccharis glutinosa, 250; sarothroides, 

Bahia dealbata, 250; dissecta, 359 

Baileya multiradiata, 238, 250 

Barbula chloronotus, 239 ; Manniae, 239 




Bartram, Edwin B., Midwinter bot- 
anizing in southern Arizona, 237 
Basidiomycetes— V, Philippine, 223 
Bebbia juncea, 250 
Betula, 172; lutea, 172; nigra, 172,293 

Biatorella, 173; resinae, 164, 173 
Blechnuni, 8; attenuatum, 3, 5, 8 
Boletus dichrous, 229; thelephoroides, 


Bolivia, New species of trees of medical 
interest from, 259 

Botanizing in southern Arizona, Mid- 
winter, 237 

Botrytis, 301, 305, 328; cinerea, 305, 
314, 317, 325, 327, 335 

Bouteloua filiorfmis, 242; Parryl, 242; 
Rothrockii, 242 

Brachiomonas, 75-77, 79, 80, 82, 84, 
85. 87, 89.92. 125, 131, 132, 135-137; 

gracilis, 75, 76, 81, 89, 91; simplex, 

82-85, 89, 91; submarina, 75, 76, 78, 
79, 82-84, 89, 91, 92; submarina 

obtusa, 77, 79, 81, 89, 91 

Brachiomonas, The phylogeny of the 

genus, 75 
Brachyphyllum, 207 
Brachyruscus, 212, 213; Alleni,2ii- 

Brassica nigra, 244 

British and American species of 
Lobomonas: a study in morpho- 
genesis of motile algae, New, 123 

Brixton, N. L., & Rose, J. N., Two 

new genera of Cactaceae, 251 
Bromuscommutatus, I04;eximius, 176; 

inermis, 94, 95, 100, loi; japonicus, 

104; tectorum, 100; vulgaris exi- 

mius, 176 
Buchnera elongata, 195 
Buechnera, 195 
Bulgaria, 21; inquinans, 15; nigrlta, 

i6, 18, 20 

Bulgariella nigrita, 16; pulla nigro- 

ollvacea, 16 
Bullaria, 190; Zorniae, 190 

Cactaceae, Two new genera of, 251 

California — i, Miscellaneous notes on 
plants of Southern, 31; — 11^ 349 

Calliandra eriophylla, 239, 244 
Callisia, 271 

Callistemon lanceolatum, 3, 4 
Callitriche palustris, 246 
Campanula sibirica, loo^ loi 
Campelia, 271 
Canna, 55 

Capsella, 154, 156 

Caraguata, 12; Sintenesii, 6, 14 

Carbon disu Iphide, Stimulation of 

plants by, 375 

Carex, 55, 103, 369; alata pulchra, 366; 
albolutescens, 364, 365, 372, 373; 
albolutescens cumulata, 366, 373; 
aquatilis,i76;atherodes, 103; athros- 

tachya, 103; Bicknellii, 369; brevior, 
368-370; caespitosa, 177 ; cumu- 
lata, 366; Deweyana, 176; Emoryi, 
103; festiva, 177 ;testucacea, 368-370; 
foenea, 177, 373; Goodenovii, 177; 
Heleonastes, 175, 177; Hookeriana, 
103; hormathodes, 177, 361; hor- 
mathodes invisa, 361; hormathodes 

Richii, 361, 362; laeviconica, 103; 
Longii, 373; Merritt-Femaldii, 370; 

monile, i77;obtusata, 103; Parryana, 
103; polygama, 178; praticola, 103; 
Richii, 362; saxatilis, 178; scirpi- 
formis, 103; stenophylla, 103; stra- 
minea, 364, 369, 372, 373; straminea 
Crawei, 369; straminea cumulata, 
364, 366; tenera, 103; tenera major, 
362; tenera Richii, 361, 362; vesi- 
caria, 177 

Carex — XII, Notes on, 361 
Carlowrightia arlzonica, 249 
Carpolithes, 211 
Cassia Covesii, 245 

Castanea alnifolia, 266, 267; alnifolia 
floridana, 266; alnifolia pubescens, 
266, 267; Ashei, 267; dentata, 266, 
267; floridana, 266, 267; neglecta, 
266, 267; pumila, 265-267; pumila 
Ashei, 267;pumila Margaretta, 265, 
267; pumila praecox, 267; pumila 
serotina, 267 

Castilleja, 191, 193 

Catinella, I5;elastica, 16, 21; nigro- 
olivacea, 16, 18-20; olivacea, 16 

Catinella, The genus, 15 
Caulinia flexilis, 35 
Cecidomyia resinlcola, 163 
Celtis pallida, 243 
Centaurea picris, 102 
Central America, The genus Costus in, 

Cephaleurps, 12 

Cephalothecium roseum, 320 
Cercidium Torreyanum, 245 
Cereus giganteus, 247 

Chaetochloa macrostachya, 242; viri- 

dis, 242 
Chamaerhodos erecta, 100 
Cheilanthes Covillei, 32; Feei, 32; 

Lindheimeri, 239, 241; myriophylla, 

241; Pringlei, 241; Wrightii, 241 

Chenopodium, 95; album, 95-97; feru- 
latum, 96, 97; leptophyllum, 96, 97; 
paganum, 96, 243; pratericola, 97 

Chilopsis linearis, 249 




Chimaphila occidentalis, 355; nmbel- 
lata, 355 ^ 

Chinese classics, References to the 
algae in the, 297 

Chlamydomonas, 75, 78, 79, 82, 84, 85, 
87-89, 123-125, 127-132; brachyura 
80, 84, 85; caudata, 80, 82, 85, 86, 88, 
89, 92; metastigma, 128; pulvis- 
culus, 123; reticulata, 127; stellata, 
123; subcaudata, 85, 86 

Chlorogonium, 128 

ChroococcuSj 6 

Chrysanthemum Leucanthemum, 100 

Chrysoma laricifolia, 249 

Chrysomyxa, 191 

Chrysothamnus formosus, loi ; graveo- 
lens, loi; nauseosus, loi 

Cirsium neomexicanum, 238, 

Citrus Limonium, 7 

Cladosporiuni, 332 

Clastobryum americanum, 5 

Claytonia lanceolata Peirsonii, 352; 
umbellata 352 

Clematis Drummondii, 244 

Clusia havetioides, 5, 6, 8, 9, 13 

Cobaea scandens, 53, 55 

CocKERELL, T. D. A., A new genus of 
fossil Liliaceae, 211 

Coenogonium, 7 

Coleosanthus baccharideus, 249 ; 

Coulteri, 249; Wrightii, 249 
Collema, 7 

CoUetotrichum, 332; lagenarium, 323, 
325» 329t 330; Lindemuthianum, 333; 
Schizanthi, 330 
Cololejeunea, 4, 6, 7; diaphana, 4-7 
Colorado, Two new plants from west- 
ern, 183 

Commelina, 270-272 
Commeiinaceae, Commelinantia, A new 
genus of the, 269 

Commelinantia, 272; anomala, 273- 

Commelinantia, A new genus of the 

Commeiinaceae, 269 
Commelyna, 269, 270; anomala, 269; 

rhodantha, 269 
Commicarpus scandens, 243 
Condalia spathulata, 246 
Copelandia, 232; papilionacea, 232 
Corispermum, 97; hyssopifolium, 97; 

nitidum, 97; villosum, 97 
Corticium, 166 
Costus, 283, 284; Bakeri, 285, 288, 289; 

bracteatus, 284, 285, 292; comosust 
288; congestus, 285, 291,292; Dink- 

lagei, 286; erythrocoryne, 287; Fried- 
richsenii, 289; geothyrsus, 287; 
giganteus, 292; hirsutus, 285, 288; 
laxus, 285, 292; lima, 285, 288, 289; 

Malortieanus, 285, 289, 290; maxi- 
mus, 286, 292; niexicanus, 289; 
nutans, 285, 291 , 292 ; pictus, 289 ; pul- 
verulentus, 285, 290; sanguineus, 

285, 290; sepacuitensis, 284, 286, 
292; spicatus, 285, 290, 291; splcn- 
dens, 285, 292; villosissimus, 284, 
287, 288 

Costus in Central America, The genus, 

Covillea glutinosa, 237, 245 
Cracca tenella, 245 
Crassina pumila, 250 
Crataegus mollis, 100 
Crepidotus, 232; pogonatus, 232 
Crepis capillaris, 103; occidentalis, 103 
Cronartium, 191; coleosporioides, 191; 
194; filamentosum, 191, 194; Hark- 
nessii, 194; Quercuum, 193; ribicola, 

3i4» 319-321, 323» 329-331, 333» 
stalactiforme, 191, 194 

Crossotolejunea, 4-7 
Croton, I 

Cryptantha calycosa, 100; gracilis, 39, 

40; gracilis Hillmanii,39;Hillmanii, 

39, 40; maritima, 38, 39; racemosa, 

38; ramosissima,38,39; recurvata,39 

Cucurbita, 376; Pepo, 54, 382 

Cuscuta, 98, 107; applanata, 109; 

arvensis, 98; calycina, 98; Choisiana, 

107; Coryli, 98; corymbosa grandi- 

flora, 109; cozumeliensis, 108; 

dentatasquamata, 107 ; duran- 

gana, i09;floribunda, i09;Gronovii, 
98, Gronoviicurta, 98; macrocephala, 
108; partita, 108; pentagona, 98; 
plattensis, 98; tinctoria, 109; um- 
bellata, 109 

Cuscuta from Mexico, Three new 
species of, 107 

Cyanotis, 271 

Cyathea, 3 

Cylindrocladium scoparium, 320, 328, 

330y 334 
Cyperus cyrtolepis, 242; diandrus, 103; 

esculentus, 103 
Cystopus candidus, 323, 330, 333 

Dacryomyces, 166; deliquescens, 167, 
168; hyalinus, 166-168 

Daedalea, 225; amanitoides, 225; ap- 
planata, 225; aulaxina, 226; flavida, 
225; inconcinna, 226; indica, 225; 
intermedia, 227; lurida, 225; Palisoti, 
225; pruinosa, 225, 227;^ repanda, 
225; sanguinea, 224; striata, 231; 
subconfragosa. 225; versa tilis, 226 

Danaea, 3 . 

Danthonia pinetorum, 175; spicata 

pinetorum, 175; thermalis, 175 



Dasylirion Wheelen, 239, 242 

Dasyochloa pulchella, 242 

Daucus pusillus, 247 

Desmidium, 4 

Development of the flower and embryo- 

geny of Martynia louislana, The, 


Diabole, 194; cubensis, 194 

Diapedium Torreyi, 248 
Dicaeoma Polygoni-alpini, 189 
Dichorisandra, 271 
Didymellina Iridis, 329, 330 
Dioscorea, 55; quinquefolia, 51, 55 
Diplasiolejeunea pellucida, 4-7 
Diplocarpon Rosae, 330 
Dipterostemon pauciflorus, 242 
Discocactus, 251 
Ditaxis neomexicana, 246; sericophylla, 

Dodge, B. O., A Lachnea with a 

botryose conidial stage, 301 
Dodonaea viscosa, 246 
DoRAN, Wm. L., Effect of external and 

internal factors on the germination 

of fungous spores, 313 
DosDALL, Louise, Occurrence of the 

pycnial stage ot Puccini a Taraxaci, 

Draba corrugata, 352 ; corrugata 

saxosa, 352; cuneifolla, 239, 244; 

saxosa, 352 
Dracocephalum Moldavica, lOO; parvi- 

florum, 100; thymiflorum, 100 
Drepanolejeunea campanulata, 

crucianella, 7 
Dryopetalon, 240, 247 ; runcinatum, 

239. 244 
Dryopteris spinulosa, 105; Thely- 

teris, 105 
Dunaliella, 130 
DuRAND, Elias J. The genus Cati- 

nella, 15 
Dysodia porophylloides, 250 
Earliella corrugata, 224; cubensis, 224 
Echinocactus, 251 ; Beguinii, 252; 

bicolor, 251; conoideus, 252; hexae- 

drophorus, 251 ; lophothele, 251 ; 

Wislizeni, 247 

Echinocereus Fendleri, 247 

Echinodorus cordifolius, 35 

Effect of external and internal factors 

on the germination of fungous 
spores, 313 

Elaphoglossum, 3; latifolium, 4, 6, 8- 
io» 13, 14 

Elatine californica, 35 

Elmerina, 226; cladophora, 226; ves- 
pacea, 226 

Encelia farinosa, 240, 250 

Endothia parasitica, 320, 330 


Ephedra antisyphiUtica, 241 ; trif ur- 

cata, 241; viridis, 241 
Epicampes ringens, 242 
Epidendrum, 7 
Epiphyllous plants of certain regions 

in Jamaica, i 
Eragrostis megastachya, 242 
Eremolithia, 351; Rixtordii, 351 
Erigeron divergens, 250; grandiflorus, 


Eriogonuni aplculatum, 35; fasclcu- 
latum, 351 ; fasciculatumflavoviride, 
350; fasciculatum foliolosum, 351 ; 
fasciculatum polifolium, 350, 351 ; 
nodosum, 350; nodosum Jaegeri,35o; 

ovalifolium, 179; pinetorum, 243; 

polycladon, 243; Wrightii, 243 
Eriophorum polystachyon, 176 
Eritrichium racemosum, 38 
Erodium cicutarium, 245; texanum, 245 
Erucastrum Pollichii, 94 
Eschscholtzia mexicana, 244 
Eupatorium solidaginifoliuni, 249 
Euphorbia capitellata, 246; Esula, 95; 

pediculifera, 246; polycarpa, 246 
Evolvulus linifolius, 247 
Evonymus atropurpureus, loi 

Fagus pumila, 267 

Farr, Clifford H., Quadripartition 

by furrowing in Sisyrinchium, 51 
Favolus, 228; cucullatus, 227; curtipes, 

227; megaloporus, 228; multiplex, 

228; princeps, 228; spathulatus, 228 

Fendlera rupicola, 244 

Festuca Hallii, 104; viridula, 104 

Fissidens, 7 

Floscopa, 271 

Fomes, 172; Bakeri, 172; rimosus, 172; 
robustus, 172 

Forrestia, 271 

Foresteria phillyreoides, 247 

Fossil, A modern plant, 63 

Fossil hepatic, A new American, 207 

Fossil Liliaceae, A new genus of, 211 

Fouquleria splendens, 246 - 

Fragaria, 53 

Franserla deltoidea, 249 

Frasera Parryi, 37 

Fraxxnus attenuata, 247 

Freesia. 55 
Fritillaria, 56 

Fucoides erectus, 207 

Fucus, 135 

Funalia versatilis, 226, 

Fungous spores, Effect of external and 

internal factors on the germination 

of, 313 

Galactia Wrightii, 245 



Galium, 358 ; angustifoliumpinetonim, } 

^^^A.^^'^^^^"^^""^' 359; grande, 359; 
Halhi, 358; multiflorum, 358; occi- 

dentale, 358, 359 ; pubens, 359 ; 
stellatum, 249, 358 

Gayoides crispum, 246 

Geiitiana humilis, 349; simplex, 349 

Germination of fungous spores, Effect 

of external and internal factors on 
the, 313 

Glacier National Park, Unreported 

plants from, 175 
Gleocapsa, 6 
Gloeoporus, 228; candidus, 229; con- 

choides, 228; dichrous, 229; retic- 

ulatus, 228 

rufomaculans, 3 14, 320, 
328, 330 

Gnaphalium alpinum, 181; Wrightii, 

Gonium pectorale, 127, 128 

Gouania, 195 

Graff, Paul W., Philippine Basidio- 
mycetes — -V, 223; Unreported plants 
from Glacier National Park, 163 

Guarea alborosea, 263; Bang!!, 262 

Gui^nardia Bidwellii, 330 

Gutierrezia glomerella, 249 

Gymnocalycium, 251 

Gymnogramme triangularis, 32 

Gyninosperma corymbosum, 249 

Gymnosporangium clavipes, 323, 330, 
3ii, 334» 335; Juniperi-virginianae, 
318, 32i-323» 33O1 331, 334 

Hapalopilus subrubidus, 227 
Haplopnyton cimicidum, 247 
Hartmannia speciosa, 354 
Haze.v, Tracy E., New British and 
American species of Lobomonas: a 
study in morphogenesis of motile 
algae, 123; The phylogeny of the 
genus Brachiomonas, 75 
Hedeorna nana, 248; oblongifolia, 
Hedychium, 3; coronarlum, 3, 4 
Hedyosmum, 3; arborescens, 4 
Hedysarum, loi; cinerascens, loi 
Helenium montanum, 250 
Helianthus, 53 ; annuus, 382 ; giganteus, 
lOi ; grosse-serratus, loi ; Maxi- 
miliani, loi; nitidus, loi; tuberosus, 


Hepatic, A new American fossil, 207 
Herberta, 208; adunca, 208 
Hermannia pauciflora, 239, 246 
Herniaria cinerea, 35 
Hesperonia retrorsa, 243 
Heteropogon contortus, 241 
Hexagonia, 227, 228; albida, 227; 
apiaria, 227; Cesati, 227; ciliata, 226; 


cladophora, 226; Glemensiae, 227; 

cruenta, 224; cucuUata, 227; glabra, 

226; Koenigii, 227; macrotrema, 226; 

Molkenboeri, 226; subrubida, 227; 

Taxodii, 227; vespacea, 226 
Hibiscus Coulteri, 246; denudatus, 246 
Hoffnianseggia densiflora, 245 
Holacantha Emoryi, 36 

HoLLiCK, Arthur; Howe, Marshall 
A., &, A new American fossil 
hepatic, 207 

Howe, Marshall A., & Hollick, 

Arthur, A new American fossil 
hepatic, 207 

Humaria, 19; fuscocarpa, 16; mai-chica, 

16, 20; olivacea, 16 
Hydrodictyon, '^8, 2gg 
Hymenoclea monogyra, 249 
Hymenogramme spathulata, 228 
Hymenophyllum, 3 

Inga vera, 195 

Inonotus Clemensiae, 227 

low^a, Quercus lyrata in, 293 

Iris, 55 

Isocoma Hartwegi, 249 

Ixia, 55 

Jacquemontia Pringlei, 247 

Jamaica, Epiphyllous plants of certain 

regions in, i 
Jamesoniella autumnalis, 208 
Janusla gracilis, 245 
Jepsonia Parryi, 36. 

Johnston, Ivan M., Munz, Philip A. 

&, Miscellaneous notes on plants of 
Southern California — I, 31; — II, 349 

Juglans nigra, 293 

Juncus nodosus, 242 

Jungermanniopsis, 208; Gockerellli, 

Juniperus scopulorum, 239, 241 

Karschia, 15 

Koeberlinia spinosa, 245 
I Koeleria cristata, 242 
j Krameria glandulosa, 245 

Krynitzkia maritlma, 38; ramosissima, 

Lachnea, 301, 305; abundans. 301, 

302, 304, 305; cretea, 301, 303, 304 
Lachnea with a botryose conidial 

stage, A, 301 
Lactuca sativa, 53; virosa, 103 
Lamlnaria, 299 
Lamium amplexicaule, 100 
Lappula cenchrusoides, 99; ciliata, 180; 

cupulata foliosa, 99; floribunda, 99; 

Lappula, 99; occidentalis, 99; texana 

homosperma, 99 



Laschia, 229; minima, 229; spathulata, 

Lecanora, 7 
Lejeunea flava, 4-7 

Lemalis, 18; pulla, 18; pulla nigro- 
olivascens, 18; rufo-olivacea, 16, 18 
Lemna, 103; trisuica, 349 

Lentinus, 229, 232; brachatus, 230; 
Decaisneanus, 230; fusco-purpureus, 
229; iufundibuliformis, 230; javani- 
cus, 230; Kurzianus, 230; polychrous, 
230; praerigidus, 230; Sajor-Caju, 
230; setiger, 230; tanghiniae, 230; 
velutinus, 230; Zeyheri, 230 

Lenzites, 225, 231; acuta, 231; ap- 
planata, 225; aspera, 226; nivea, 227; 
Palisoti, 225; pallida, 225; platy- 
phylla, 226, 231; platypoda, 
repanda, 225; striata, 231 

Lepiidium densiflorum, 93, 94; Draba, 
94; hlrsutum, 244; lasiocarpum, 239, 
244; ramosissi,mum, 93, 94 

Leptqsyne arizonica, 250 

Lesquerella Fendleri, 238, 244; pur- 
purea, 239, 244 
Liiiaceae, A new genus of fossil, 211 
Lilium Martagon, 150 
Linum spergulinum, 36 
Lippia Wrightii, 248 

Lobomonas, 89, 123, 124, 128, 130- 

I33i 135-137; Francei, 123-125, 127, 
129, 130; pentogonia, 124, 125, 127, 
129, i39;rostrata, 124, 127, 130, 139; 
stellata, 123 

Lobomonas : a study 1 n morpho- 
genesis of motile algae, New British 
and American species of, 123 

Lolium remotum, 104; rigidum, 104; 
temulentum, 104 

Lomatium macrocarpum, loi 

Lopadium, 4 

Lophocolea, 5-7 

Lupinus, 197, 204; affinis, 198, ' 199, 
204; apricus, 204; arvensis, 200, 201; 
Aschenbornii, 204; Barkeri, 197; 

bicolor, 204; bilineatus, 197; car- 
nosulus, 204; chihuahuensis, 204; 
Clarkei, 197; densiflorus, 200, 201; 
Ehrenbergii, 200, 201 ; Hartwegi, 
197; hirsutulus, 204; leucophyllus, 
205; Liebmanni, 199; micranthus, 
204; micranthus microphyllus, 204; 
nanus, 199, 204, 205; niveus, 197, 
204, 205; pachylobus, 204; persistans, 
204; Pipersmithii, 204; polycarpus, 
204; purpureas, 199; ramosissimus, 
200, 201; rostratus, 204; sabulosus, 
204; strigulosus, 204; succulentus, 
197-200; succulentus Brandegeei, 

203; succulentus Layneae, 203; tri- 
fidus, 204; umbellatus, 204; vallicola, 
204; vallicola apricus, 204 
Lupinus succulentus and L. niveus, 
Studies in the genus Lupinus — VII, 

Luzula comosa, 349 

Lychnis alba, 95 

Lycium, 357; Cooperi, 248; Fremontii, 

248; parviflorum, 248; Spencerae, 

356, 357; Torreyi, 248 
Lycopus communis, 100 

Mackenzie, Kenneth Kent, Notes on 

Carex— XII, 361 
Madia glomerata, 102 
Magnolia, 51, 55-59 
Malacocarpus, 25 1 
Malus elongata, 268; elongata pubens, 

268; platycarpa parvula, 268 

Malva boreaHs, 95; crispa, 95; verti- 

cillata, 95 
Malvastrum, 353 
Mammlllaria conoidea, 252; Grahami, 

Mangifera indica, i, 5, 10, 13 
Marattia, 3 

Marchantia Pealei, 207 
Marchantltes erectus, 207 
Marrubium vulgare, 248 
Marsilea vestita, 35 
Martynia, 141, 150, 154; fragrans, 14T, 

142; louisiana, 141, 142; lutea, 141 
Martynia louisiana, The development 

of the flower and embryogeny of, 141 
Mecardonia peduncularis, 248 
Medicago falcata, 100 
Melampsoropsis Rhododendri, 191 ; 

roanensis, 190 

Melanconium, 323 

Melanospora, 305 

Menodora scabra, 247 

Merulius fugax, 168, 169; striatus, 231; 

subaurantiacus, 169 
Mesosphaerum Emoryi, 248 
Metiola astrina, 12 
Metzgeria furcata, 4-7 

Mexico, Three new species of Cus- 

cuta from, 107 
Microlejeunea, 4, 5 
Microstylis monophyllos, 349 
Midwinter botanizing in southern 

Arizona, 237 
Mimosa asperata, 194; pigra, 194 
Mimulus Langsdorfii, 248 

Miscellaneous notes on plants of 
Southern California — I, 31; — 11, 

A 349 ^ 

Mittenothamnium reptans, 5, 6 

Modern plant fossil. A, 63 



Mollisia umbonata. 19 

Mollugo verticillata, 35 
Monbretia, 55 

Monoclea Gottschei, 6 

Mucronella, 169 

M uhlenbergia dumosa, 242 ; micro- 

sperma, 242; Porteri, 242 
MuNZ, Philip A., & Johxston, Ivan 

M., Miscellaneous notes on plants of 
Southern Caliiorniia — I, 31; — II, 

Mycoidea parasitica, I 
Mycologxcal notes for, 1920, 163 
Myoporuni serratuni, 154 
Myosurus cupulatus, 36 

Naias, 54; flexilis, 35 

Nama humifusum, 356 

Nardia, 208 

Nectandra, 259, 261; coto^ 260 

Nectriella, 164; resinae, 164 

Neolloydia , 25 1 ; Beguinii , 252 ; conoi- 

dea 252 

New American fossil hepatic, A, 207 

New British and American species of 

Lobonionas: a study in morpho- 

gensis of motile algae, 123 - 

New genus of fossil Liliaceae, A, 211 

New genus of the Commelinaceae, 

Commelinantia, A, 269 

New plants from western Colorado, 
Two, 183 

New records and other notes on North 

Dakota plants, 93 
New species of trees of medical interest 

from Bolivia, 259 
New species of Uredineae — XIV, 189 
Nicotiana, 55-57, 591 glauca, 248; 

trigonophylla, 248 
Nissolia Schottii, 245 
Nolina microcarpa, 242 
North Dakota plants, New records 

and other notes on, 93 
Nostoc, 4-6, 299 

Occurrence of the pycnial stage of 

Puccinia Taraxaci, 235 
Ocotea, 261; pseudo-coto, 261 

Oenothera, 354; cardiophylla, 355; 
cardiophylla splendens, 354; speci- 
osa, 354 

Oldenbusch, Carrie, Stimulation of 

plants by carbon disulphide, 375 
Olneya tesota, 245 

Ophioglossum Alleni, 211 
Ophrys monophyllos, 349 

Opuntia arborescens, 247 ; Bigelovii, 
238, 247; chlorotica, 247; fulgida, 
246; leptocaulis, 247; Toumeyi, 247 

Osmunda, 52; .palustris aurea, 52; 
regalis, 52 

OsTERHOUT, Geo. E., Two new plants 
from western Colorado, 183 

OvERHOLTS, L. O., Mycological notes 
for 1920, 163 

Notes for 1920, Mycological, 163 

Notes on Carex — XII, 361 

Notes on North Dakota plants, New 

records and other, 93 
Notes on plants of Southern California, 

Miscellaneous, — I, 31; — II, 349 
Notes on trees and shrubs of the south- 
eastern United States, 265 
Notholaena bonariensis, 241 ; cali- 
fornica, 32; Hookeri, 241; Lemmoni, 
239, 241; Parryi, 32; sinuata, 241; 
sinuata integerrima, 31, 241 j tenera, 

Nuttallia, 183; marginata, 183 

Nymphaea, 56 
Nyssa sylvatica, 166 

Panaeolus, 232 ; campanulatus, 232 ; 

papilionaceus, 232 

Pandorina, 128 

Panicum perlongum, 103 

Pannaria rubiginosa, 7 
Panus, 232 

Papulospora, 305 

Parietaria obtusa, 243 

Parkinsonia microphylla, 245 

Parmelia, 4-6 

Parnassia cirrata, 349 

Parosela Greggii, 245; Parryi, 245 

Parthenium incanum, 250 

Patellaria applanata, 16, 19; hirneola, 
16; olivacea, 16, 20; puUa, 19; pulla 
nigro-olivacea, 16, 18; violacea, 16 

Patinella applanata, 16, 19; hirneola, 
16, 19; olivacea, 16, 19; violacea, 16, 

Pectocarya linearis, 247 

Pediastrum, 131, 132 

Pellaea compacta, 32; mucronata, 239, 


Penicillium glaucum, 334, 375 

Penstemon, 43, 189; albomarginatus, 
44; antirrhinoides, 43; antirrhinoides 

microphyllus, 43; barbatus, 40; 
barbatus Torreyi, 40; calcareus, 42; 
Cleveland!, 41, 357; Cleveland! con- 
natus, 357; Cleveland! Stephens!, 
41 1 357; desertorum, 42; Eaton i, 40; 
truticosus, 180; Grinnellii, 42; Men- 
ziesii, 180; microphyllus, 43;Munzii» 
40; Palmeri, 41, 42; Palmeri Grin- 
nellii, 42; petiolatus, 42; Pluniiuerae, 




43; Stephensi, 

sup)erbus, 248 
Perezia Wrightii, 249 
Peridermium, 193; Cerebrum, 192, 193 




•V, 223 

filamentosum, 191, 194; Harknessii, 

191-193; stalactiforme, 191, 194 
Perityle dissecta, 250^ 
Peronospora parasitica, 323 ; pyg- 

maea, 335 
Pessin, Louis J., Epiphyllous plants 

of certain regions in Jamaica, i 
Pezicula, 19; viridi-atra, 16 
Peziza applanata, 18; fuscocarpa, 16, 

19, 20; nigro-olivacea, i6» 18, 19; 

olivacea, 15-1^; viridi-atra, 16, 19,20 
Phacelia Franklinii, 180 

Phaeopezia, 19; elastica, 21 ; fusco- 
carpa, 16, 20; marchica, 16 
Phallogaster saccatus, 172 * 
Phaseolus, 376 

Phiiadelphus microphyllus, 
Philibertella linearis, 247 
Philippine Basidiomycetes 
Phlox, 99; andicola, 99; bernardina, 

356; dolichantha, 356; Douglasii, 99; 

Hoodii, 99; Stansburyi brevifolia, 

356; superba, 356 
Phoradendron californicum, 243; cali- 

fornicum distans leucocarpum, 349; 

macrophyllum, 243 
Phragmidium violaceum, 331 
Phycopeltis, 3-7, 10; microcystis, 2 
Phyllonoma, 212; ruscifolia, 212 
Phylloporina epiphylla, 6 
Phyllosticta Antirrhini, 323, 330 
Phylogeny of the genus Brachiomonas, 

The, 75. 
Physalis ixocarpa, 248 

Phytophthora infestans, 323, 327, 330, 

Picea, 191 

Pilacre Petersii, 165, 173 

Pinnularia, 4-6 

Pinus insignis, 

193; radiata, 

virginiana, 164 
Piper, 3 

Pittosporum tobira, 4 
Plagiochila, 6 

Plasmodiophora Brassicae, 322, 323 
Plasmopara viticola, 323, 330, 333, 334 

Platanus occidentalis, 293; Wrightii, 

Platymonas, 80; subcordiformis, 80 

Pleurotus, 229; flabellatus, 229 

Pluchea, 194; fastigiata, 194; Quitoc, 


Plumbago scandens, 247 

Poa Bigelovii, 242; Cusickii, 176; 

nevadensis, 176 
Podocarpus, 3; coriaceus, 5, 7 
Polemonijum confertum, 179 
Pollia, 271 

Pol yblephar ides, 80, 128 

1 93 1 ponderosa, 
193 ; Strobus, 


Polygala macradenia, 240, 246 
Polygonum alpinum, 189; bistortoides 

lijiearifoUum, 179; Hydropiper, 97; 
punctatum, 243; punctatum leptos- 
tachyum, 97 
Polypodium, 3, 5, 8; Phyllitidis, 7, 8, 

10, 14 

Polyporus, 228; annosus, 172; apiarius, 

227; asper, 223; caeruloporus, 169; 
Clemensiae, 227; compactus, 170, 
I7i» 173J corrugatus, 224; dichrous, 
229; ElUsianus, 172; fusco-badius, 
224; incanus, 224; indecorus, 224; 
lacerus, 226; leucoplacus, 223; mega- 
loporus, 228; minimus, 229; nigro- 

purpurascens, 229; paleaceus, 224; 

pelleporus, 229; scabrosus, 224; 

subrubidus, 228;tegularis, 224; thele- 

phoroides, 229 
Polystichum plaschnichianum, 8 
Polystictus, 226; asper, 223; Persoonii. 

224; thelephoroides, 229; versatilis, 




Poria, 223; leucoplaca, 223 
Porophyllum juniciforme, 250 
PoRTERFiELD, Jr. W. M., References 

to thealgaein the Chinese classics, 297 
Potent! lla argentea, 100 
Preissites Wardii, 207 
Prosopsis glandulosa, 245 
Prunus fasciculata, 356 
Pseudppeziza Medicaginis, 317; Tri- 


Psilostrophe Cooperi, 250 
Psilotum, 52; triquetrum, 52 
Pteromonas, 124, 128, 129, 135; alata, 

Ptiloria pauciflora, 249 

Puccinia, 189; Antlrrhini, 316, 323, 330; 

biocellata, 194; coronata, 321, 323, 

- 332, 334; dispersa, 323; graminis, 
32I7323, 329, 334; invaginata, 195; 
Larici, 323; Malvacearum, 314, 322, 

323, 329-334; offuscata, 190; Phlei- 
pratensis, 323, 330; Plucheae, 194; 

Polygoni-alpini, 189; rubigo-vera, 

323; Sorghi, 323; Taraxaci, 235; Zor- 

niae, 190 
Puccinia Taraxaci, Occurrence of the 

pycnial stage of, 235 
Pucciniastrum americanum, 190; arcti- 

cum americanum, 190 
Pyramimonas, 80 
Pyrenula, 7 
Pyrola picta, 355; umbellata, 355 

Quadripartition by furrowing in Sisy- 
irnchium, 51 



Quercus, 170; acuminata, 268; alba, 
170; aquatica, elongata 268; ari- 
zonlca, 243; bicobr, 293; colora- 
densis, 268; Emor>'i, 293, 243; 
hypoleuca, 239, 243; lyrata, 293-295; 
macrocarpa, 268, 293, 294; nig a 
elongata, 268; nigra tridentifera, 
268; oblongifolia, 239, 243; stellata, 
268; virginiana. 268 

Quercus lyrata in Iowa, 293 

Refeences to the algae in the Chinese 
classics, 297 

Rhamnus tomentella, 246 
Rhizina nigro-olivacea, 16, 19 
Rhizopus nigricans, 314, 326, 327, 335, 

^ 383^ 384 

Rhododendron, i, 6, 191; catawbiense, 

^190; punctatuni, 190 
Riccardia, 6 

Rose, J. N., Brixton, N. L., &, Two 

ne\v genera of Cactaceae. 251 

Round, Eda M., A modern plant fossil, 

RowxEE, W. W.. The genus Costus in 

Central America, 28;^ 
Rubus neglectus, 190; occidentalis, 190 
Ruellia tuberosa occidentalis, 249 
Runiex Britannica, 105; hymenose- 

palus, 243; paucifolius, 189 
RusBY. H. H., New species of trees of 

medical interest from Bolivia, 259 
Ruscus, 212, 213; hypoglossuni, 212; 

hypophyllum, 213; latifolius, 213 

Rutosma purpurea, 246 



Salix orbicularis, 178; reticulata, 175, 
178, 179; saximontana, 178 

Salvia lanceolata, 100; mohavensis, 37; 
pinguifolia, 248 

Salvinia, 211; reticulata, 211, 212 

Sambucus mexicana, 249 

Samuela, 33 

Sarcosoma rufum, 15 

Sauvellea, 271 

Saxifraga Parryi, 36 

Scaphophorus agaricoides, 231 

Schizonia vulgaris, 232 

Schizophyllum, 231; alneum,. 


commune, 231 
Scirpus occidentalis, 103; validus, 103 
Scleroma velutinum, 230 
Sclerotinia fructie^ena. '^14.. '^.lo. 

Senecio aureus, 181 ; crocatus, 181; 

Lemmoni, 250; manitobensis, I02 
Septoria, 323; Gladioli, 330; Lycoper^ 

sfci, 334 

Shimek, B., Quercus lyrata in Iowa, 

Shrubs of the southeastern United 

States, Notes on trees and, 265 
Sicyos angulatus, loi 
Sideranthus australis, 249; Gooddingi, 

Silene dichotoma, 95; Douglasii, 

Fabaria, 95; noctiflora, 95 
Silphium, 147 

Simmondsia californica, 246 
Siphonoglossa longiflora, 249 
Sisymbrium Loeselii, 94 

Sisyrinchium, 55~57^ 59» I95; 
mudiana, 195; Bushii, 56, 6r 

Sisyrinchium, Quadripartition by fur- 
rowing In, 51 
Smith, Charles Piper.. Studies in the 

genus Lupinus — VII. L. succulentus 
and L. niveus, 197 

Solenia fasciculata/ 169, 173; villosa, 

Solidago humiHs, 180; Purshii, 180 
Sonchus arvensis, 102; uliginosus, 102 
Sophia intermedia, 94; ochroleuca, 244 
Southern California, Miscellaneous 

notes on plants of, — -I, 31; — II, 349 
Sphaeralcea, 353; ambigua, 246, 353; 

grossulariaefolia, 246; rosacea, 353 
Spnaerotheca mors-uvae, 318 
Spirodela polyrhiza, 103 
Sporobolus asper, 104 
Staurastrum, 4 
Stelis ophioglossoides, 5 
Stemodia durantifolia, 248 
Stenolobium incisum, 249 
Stereum radiatum, 168 
Sterigmatocystis, 384, 385; nigra, 376, 

Stevens, O. A., New records and other 
notes on North Dakota plants, 93 

Sticta, 3, 4 

of plants by carbon 

333^ 335; Ger; 
Ricini, 301 

Scopulophila, 351 
Riifordii, 351 

Scrophularia parv 
Scytonema, 4-7 
Sedum Griffithsii 
Selaginella arizoni 




disulphide, 375 
Strepanthus arizonicus, 244 
Streptolirion, 271 

Strigula, i 

Studies in the genus Lupinus 

L, succulentus and L. niveus, 197 
Study in morphogensis of motile algae, 

New British and American species of 

Lobomonas: A, 123 
Suckleya Suckleyana, 97 
Syntherisma sanguinale, 103 

Taxtlejeunea, 5-7 



Tharp, B. C, Commelinantia, a new 

genus of the Commellnaceae, 269 
Thelephora dolosa, 229 

Thelocactus, 25 1 ; bicolor, 251; hexae- 
drophorus, 251; lophothele, 251 

Thiaspi arvense, 352 

Three new species of Cuscuta from 

Mexico, 107 
Thysanocarpus, 240, 247; amplectens, 

239. 244 
Tilletia foetens, 330 

Tinantia, 269, 270, 272; anomala, 270, 

273; fugax, 270; Pringlei, 275 
Tmesipteris, 211; Alleni, 212 
Tortula incnnls, 239; ruralis, 239 
Tradescantella, 271 
Tradescantia, 54, 269-273; anomala, 

269, 271-273; Pringlei, 271, 272 
Tragia ramosa, 37 
Trametes, 223; aspera, 223; corrugata, 

224; incana, 223; nitida, 224; palea- 

cea, 224; versatilis, 226 
Trapella sinensis, 154 
Trees and shrubs of the southeastern 

United States, Notes on, 265 
Trees of medical interest from Bolivia, 

New species of, 259 
Trentepohlia, 3; arborum, 2; aurea, 2, 

^4. 5. 7 ^ 

Trichloris fasciculata, 242 

Trichomanes, 3, 5 

Tricuspis mutica, 32 

Triodia mutica, 32 

Triticum, 387; sativum, 379 

Trixis californica, 249 

Tsuga canadensis, 168 

Tulasnella, 166; Violae, 

Two new genera of Cactaceae, 251 

Two new plants from western Colorado, 




Ulmus americana, 293 

United States, Notes on trees 
shrubs of the southeastern, 265 

Unreported plants from Glacier Nat- 
ional Park, 163 

Uredineae — XIV, New species of, 189 

Uredo biocellata, 194; coleosporioides, 

194; cumula, 195; curvata, 195; 

nominata, 194; Plucheae, 194; Zor- 

niae, 190 
Uromyces, 1 89 ; caryophylli nus, 3 1 6, 

323; fuscatus, 189; Trifolii, 323 
Uromycladium, 194; cubense, 194 
Usnea, 4 
Ustilago Hordei, 330; Tritici, 330 

Vaccaria Vaccaria, 95 

Valeriana occidentalis, 180; sylvatica, 

180; wyomingensis, 180 
Vauquelinia californica, 244 
Venturia inaequalis, 314, 317, 318,320, 

325-328, 330-332, 334. 335 
Verbena, 148, 150; Wrightii, 248; 

xutha, 248 
Veronica Cusickii, 180; maritima, 98 
Vicia, 381, 387; Faba, 380' 381 
Villanova dissecta, 359 

Vinca rosea, 51 

Viola Sheltoni, 353, 354 

Wedeliella incarnata, 243 
Woodsia scopulina, 31 
Woodwardia angustifolia, 


Xanthium, loi; acerosum, loi; cana 
dense, loi; italicum, loi; pennsyl 
vanicum, loi 

Yucca baccata, 33, 34, 239, 242 ; 

baccata macrocarpa, 34; elata, 238, 

242; glauca, 196; macrocarpa, 34; 

mohavensis, 33, 34; Schottii, 34 
YuNCKER, T. G.. Three new species of 

Cuscuta from Mexico, 107 

Zannichellia palustris, 35 

Zea Mays, 376 

Zebrina, 271 

Zythia, 164, 173; resinae, 163, 164, 173