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Full text of "Journal of the Arnold Arboretum."

JOURNAL 

OF THE 

ARNOLD ARBORETUM 

HARVARD UNIVERSITY 

ALFRED REHDER 
JOSEPH H. FAULL and CLARENCE E. KOBUSKI 

VOLUME XVI 




Reprinted with the permission of the 
Arnold Arboretum of Harvard University 
KRAUS REPRINT CORPORATION 

New York 



0E( 



1968 



DATES OF ISSUE 

No. 1, (pp. 1-143, pi. 119-128) issued January 25, 1935. 

No. 2, (pp. 145-271, pi. 129-139) issued April 24, 1935. 

No. 3, (pp. 273-365, pi. 140-154) issued July 10, 1935. 

No. 4, (pp. 367-483, pi. 155-165) issued October 25, 1935. 



Printed in U.S.A. 



TABLE OF CONTENTS 

S.TUD1ES IN BORAGINACEAE, X. ThE BoRAGINACEAE OF NORTHEASTERN 

South America. By Ivan M. Johnston 1 

and one text figure. By Alfred Rehder 65 

Notes on some of the Ebenaceae and Verbenaceae of the Solo- 
mon Islands Collected on the Arnold Arboretum Expedition, 
1930-1932. With plates 120-122. By R. C. Bakhuisen van den Brink 68 
An Endemic Sophora from Rumania. With plates 123 and 124 and 
one text figure. By Edgar Anderson 76 

tum. By Ernest J. Palmer 81 

The Hosts of Gymnosporangium globosum Farl. and their Rela- 
tive Susceptibility. With plates 125-128 and four text figures. 
By /. L. Ma^Lachlan 98 

A Preliminary Note on Life History Studies of European 
Species of Milesia. By LiUian M. Hunter 143 

Studies in the Boraginaceae, XL By Ivan M. Johnston 145 

Loranthaceae Collected in the Solomon Islands by L. J. Brass 
and S. F. Kajewski on the Arnold Arboretum Expedition, 
1930-1932. With plate 129. By B. H. Danser 206 

Chromosome Numbers in the Hamamelidaceae and their Phylo- 
genetic Significance. With three text figures. By Edgar Ander- 
son and Karl Sax 210 

Chromosome Structure in the Meiotic Chromosomes of Rhoeo 
discolor Hance. With plates 130 and 131. By Karl Sax 216 

Elaioplasts in Iris: A Morphological Study. With plates 132- 
137. By Anna F. FauU 225 

Notes on Yucca. With plates 138 and 139. By Susan Delano 
McKelvey 268 

The Visible Structure of the Secondary Wall and its Signifi- 
cance IN Physical and Chemical Investigations of Tracheary 
Cells and Fibers. With plates 140-149. By /. IV. Bailey and 
Thomas Kerr 273 

The Effect of Temperature on Nuclear Differentiation in 
Microspore Development. With one text figure and plate 150. By 
Karl Sax 301 

Notes on the Ligneous Plants Described by Leveille from East- 
ern Asia. By Alfred Rehder 311 

Huodendron, a New Genus of Styracaceae. With one text figure 
and plates 151 and 152. By Alfred Rehder 341 

Studies in Theaceae, I. Eurya Subgen. Ternstroemiopsis. With 
plate 153. By Clarence E. Kobuski 347 

Two New Species of Crataegus from Missouri. With two text 
figures. By Ernest J. Palmer 353 



iv TABLE OF CONTENTS 

New Hybrids from the Arnold Arboretum. By Edgar Anderson 
and Alfred Rehdcr 358 

Hypodermella Hiratsukae, a New Species of Hypodermataceae 
FROM Japan. With plate 154. By Grant D. Darker 364 

The Hosts, Life History and Control of Gymnosporangium 
CLAViPEs C. and p. With plates 155-160. By Ivan H. Crowell ... 367 

The Dispersal of Viable Basidiospores of the Gymnosporan- 
gium Rusts. With two text figures. By /. D. MacLachlan 411 

Chromosome Structure and Behavior in Mitosis and Meiosis. 
With plates 161-164. By Hally JoUivette Sax and Karl Sax 423 

The Flora of San Felix Island. With plate 165. By Ivan M. 
Johnston 440 

Some New Trees and Shrubs from Mexico. By Alfred Rehder. . 448 

New Facts Concerning Cephalosporium Wilt of Elms. By D. B. 
Creager 453 

The Arnold Arboretum during the Fiscal Year ended June 30, 
1935; The Arboretum; The Pathological Laboratory; The Cyto- 
genetic Laboratory; The Herbarium; The Library; Bibliography of 
the published writings of the staff and students, July 1, 1934— June 
30, 1935 455 

Staff of the Arnold Arboretum 1934-1935 469 



JOURNAL 

OF THE 

ARNOLD ARBORETUM 

^I JANUARY, 1935 



^UDIES IN BORAGINACEAE, X 

THE BORAGINACEAE OF 

NORTHEASTERN SOUTH AMERICA 

Ivan M. Johnston 

The present paper is a critical account of the Boraginaceae known 
from British, Dutch and French Guiana and the adjoining portions of 
Brazil, north and east of, the Amazon and the Rio Negro. A general 
account, it is preliminary to a treatment of the Dutch Guianan species 
of the family which Prof. A. A. Pulle has invited me to prepare for his 
"Flora of Surinam." 

The borages of the Guianas have been long neglected, ^uch frag- 
mentary work as has been done on them has been restricted to the nar- 
row political boundaries. Though various species of the group have 
been described from the Guianas, some of them among the first based 
upon South American material, the identity of the types has remained 
obscure, and material in herbaria has continued to be named largely by 
guess or has been left to accumulate unidentified. The great reference 
works, such as DeCandolle's Prodromus, or Martius's Flora Brasiliensis, 
resolve little of the confusion that seems always to have enveloped our 
knowledge of the Guianan Boraginaceae. They added little to the 
observations all ready long available in the writings of Lamarck and 
Poiret. Indeed, so little known and confused were the Guianan species 
of Cordia and Tomnejortia, that a few years ago, during my studies 
of the Brazilian species of these genera, Contr. Gray Herb. 92: 1-89 
(1930), I was forced to pass over, undiscussed, the very evident rela- 
tions existing between the species of the Guianas and those of northern 
Brazil, and forced to admit that certain of the obscure species (several 
of them not even listed in the Index Kewensis) might be identical and 
older than the ones I was forced to accept. A careful study of the 
Guianan Boraginaceae has been long needed. 



2 JOURNAL OF THE ARNOLD ARBORETUM Lvol. xvi 

The conspicuous relationship evident among the Guianan Boragi- 
naceae is that with Brazil, most of the species extending into and about 
the Amazon Basin or having their immediate relatives there. The 
affinities westward in Venezuela are not so numerous nor so pronounced. 
Except for Trinidad (which after all is floristically close to that of east- 
ern Venezuela) direct relations to the northward are negligible. Among 
the Guianan Boraginaceae only the group Cordia § Pilkordia has devel- 
oped any number of local species. The relations of these local endemics 
are in the Amazon Basin where the species of this group are not local 
but widely distributed. The Guianas are a marked endemic center for 
Pilkordia comparable with the centers of that group found in south- 
eastern Brazil, northern V^enezuela and adjacent Colombia, and the 
West Indies. 

I have treated in this paper all the borages known north and east of 
the Amazon, the Rio Negro and the eastern boundary of Venezuela. 
The monotypic Lepidocordia is endemic to this area. Of the 38 species 
definitely known from this large area only two, Cordia multispicata and 
Cordia naidophila are at present unknown from British, Dutch or 
French Guiana. Several other species approach our area, reaching the 
Orinoco Valley from the westward. Among these species those which 
may eventually be found in northwestern British Guiana are Cordia 
globosa (Jacq.) HBK., Cordia alba Jacq. and Bourreria cumanensis 
(Loefl.) O. E. Schulz. The writings of Schomburgk, Aublet and others 
have listed various West Indian species from the Guianas. Some of 
these records are evidently based upon misidentifications, others, how- 
ever, I am convinced, are simply unfortunate guesses as to what the 
authors believed might be found there. Most of these questionable 
records relate to species common and widespread at low altitudes in the 
Antilles. This group of plants is poorly represented on the Guianan 
coastal area, probably because of adverse winds and currents and the 
unfavorably humid conditions. 

In the preparation of this report I have examined practically all the 
types concerned and have studied most of the large or important Gui- 
anan collections in Europe and United States. Studies have been made 
at Kew, London, Leiden, Utrecht, Copenhagen, Berlin, Munich, Geneva 
and Paris. Large loans of critical material have been obligingly sent 
for further, more leisurely study at the Arnold Arboretum and the Gray 
Herbarium from Kew, London, Leiden, Utrecht, Berlin, Paris and New 
York. Particular mention, however, is to be made of the large loan 
from the Botanical Museum at Utrecht. This material, assembled 
through the inspiration of Prof. Pulle and kindly made available to me 



1935] JOHNSTON, STUDIES IN BORAGINACEAE 3 

by him, consists of numerous series of copious specimens collected 
over a number of years, at different seasons, from various numbered 
individual trees or shrubs, in the Forest Reserves of Dutch Guiana. 
Through the examination of this remarkable record of seasonal varia- 
tion I have been able to establish unquestionably the specific identity of 
certain seasonal forms heretofore troublesome to identify. Of great 
help in the preparation of this report the collections have been generally 
instructive to me personally. It has been a privilege to have such a 
convincing demonstration of the nature and extent of seasonal variation 
in individual trees and shrubs of the Tropics. 

The following abbreviations have been used in designating the source 
of the material cited. B. W. — collections by the Forest Service 
(Boschwezen) of Dutch Guiana; ,AA — Arnold Arboretum; BD — 
Botanical Museum at Berlin; BM — British Museum of Natural His- 
tory; DC — Prodromus Herbarium of DeCandolle at Geneva; Del — 
Delessert Herbarium at Geneva; G — Gray Herbarium; K — herba- 
rium at Kew; Leid — herbarium at Leiden; NY — New York Botani- 
cal Garden; US — U. S. National Herbarium; Utr — Utrecht 
Herbarium. 

Stigmas 2 or 4, simple; inflorescence cymose-paniculate or 
spicate or globose, the branches not distinctly scorpioid; 
erect broad-leaved trees and shrubs. 
Stigmas 4, capitate or clavate, borne on a conspicuous slender, 

twice lobed or cleft style 1. Cordia, p. 3. 

Stigmas 2, conic, sessile on the apex of the ovary, short and 

inconspicuous 2. Lepidocordia, p. 45. 

Stigma 1, consisting of an anulate fertile base and a more or less 
developed sterile frequently bifid apical portion; inflo- 
rescence with distinctly scorpioid branches or the flowers 
cauline and solitary in the internodes. 

Fruit baccate; clambering shrubs 3. Tourncfortia, p. 46. 

Fruit dry; herbs or small sufifrutescent plants . .4. Hdiotropimn.p. 57. 
1. Cordia [Plumier] Linnaeus, Gen. 87 (1754). 
Trees or shrubs, usually with broad leaves. Inflorescence a loosely 
paniculate or glomerate or capitate or spicate cyme. Calyx usually 
5-toothed or 5-10-lobed, usually persistent. Corolla white, yellow or 
reddish, small to conspicuous, usually 5-merous, rarely 6-lS-merous, 
salvefform or subrotate to funnelform or subtubular. Stamens as many 
as the corolla-lobes, exserted or included, filaments short or long. Ovary 
4-ceIled, ovules 1-4. Style terminal, well developed, 2-lobed or 2-parted, 
the branches each 2-lobed. Stigmas 4, capitate or clavate, small. Fruit 
unlobed, a drupe with a bony pit and mucilaginous or dry exocarp, or 



4 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

the walls dry and papery, 1-4-celled. Endosperm none. Cotyledons 
plicate. 

A very large genus of diverse habit and structures; widely distributed 
throughout the Tropics. Centering in America. Type Species: C. 
sebestena L. 

On the grounds that the original "Cordia" of Plumier, Nov. PI. Amer. 
Gen. 13, tab. 14 (1703), which was accepted and validated by Lin- 
naeus, is not a member of the Boraginaceae, the generic name Cordia has 
been recently discarded for the present concept by Dr. von Friesen, Bull. 
Soc.Bot. Geneve, ser. 2,24: 131-4(1933). With this I can not agree. It 
is pointed out by von Friesen that Plumier's illustration shows a 2- 
celled ovary and a simply bifid style and that the generic description of 
Cordia (based upon Plumier's plate and description) given by Linnaeus 
in the Genera Plantarum of 1754, pg. 87, also calls for these structures. 
Dr. von Friesen believes they are structures of some genus outside of the 
Boraginaceae. I believe they are structures of Cordia sebestena faultily 
described from inaccurate drawings. 

I have had the privilege of studying, in the library of the Natural 
History Museum at Paris, the amazing series of volumes of plates and 
manuscripts accumulated by Plumier during his visits (1689-97) to 
the West Indies. In one of these volumes of manuscript, 6 : tab. 64-66, 
are found fine drawings labeled "Cordia nucis iuglandis folio, flore 
purpureo." The best of these original drawings, made in the West Indies 
by Plumier, fills a folio page and shows a characteristic branch of Cordia 
sebestena bearing leaves, flowers and fruit. In the corner of the page 
are the details of flower and fruit, differing only in arrangement from 
those printed in Plumier's Genera. The structures of style and ovary 
are quite alike in both. This may be verified by a comparison of the 
small plate in Plumier's Genera and the good copy of the original folio 
plate published in Burmann's edition of Plumier's Plantarum Ameri- 
canarum, fasc. 5, tab. 105 (1757). The later plate is identified as C. 
sebestena by Urban, Rep. Spec. Nov. Beiheft 5: 60 (1920). Von Frie- 
sen, 1. c. 135, however, believes that only the leafy, fiower-bearing branch 
is C. sebestena and that the disputed unattached analytic details belong 
to some other genus. Since, however, the disputed details are an inte- 
gral part of the original drawing of Plumier, which consists mainly of a 
flowering and fruiting branch unquestionably of C. sebestena, and since 
the details, as far as one can compare them, are quite like homologous 
structures shown growing attached to the flowering and fruiting branch 
of C. sebestena, I feel there is every reason for believing that, however 
inaccurate, they were intended to show the structures of that species. 



1935] JOHNSTON, STUDIES IN BORAGINACEAE 5 

It is to be recalled that Plumier's drawings were made long before the 
work of Linnaeus on the Sexual System directed particular attention and 
gave special importance to the number and structure of the internal parts 
of the flower and fruit. Plumier, and Linnaeus who copied from him, 
may have given erroneous descriptions of the fruit and style of Cordia 
sebestena but since they were trying to describe that species I believe we 
should retain their name for the genus containing it. 

Plumier's drawing was made on the island of St. Thomas. The fol- 
lowing quotation from his manuscript gives the type-locality of C. sebes- 
tena in some detail. "Martio plantarum florentem frutusque matures 
ferentem adinveni apud insulam Sancti Thomae, juxta Littus quoddam 
La Baye du nord vocitatum, sinum scilicet ad septenttrionalem plagam 
ipsi Arci oppositum." 



Corolla marcescent; fruit cylindrical, dry, with a fibrous charta- 
ceous coat, not bony, closely invested by the tube of the per- 
sistent corolla and the strongly ribbed cylindrical calyx, at 
maturity flower disarticulating from the inflorescence and 
the calyx and corolla and the enclosed fruit falling away 
together, with the spreading corolla-lobes acting as a para- 
chute ; pubescence stellate ; axis of inflorescence usually tun- 
nelled or inflated and inhabited by ants 1. C. alliodora. 

Corolla withering after anthesis and soon deciduous ; fruit usu- 
ally baccate, with a bony ovoid or globose stone ; pubescence 

Corolla red or orange, large ; calyx becoming fleshy and com- 
pletely enveloping the dry fruit and even adnate to it 

Corolla white or yellow; fruit juicy, not adnate to the dry 

Corolla-lobes longer than broad; calyx explanate at ma- 
turity ; inflorescence usually large and loosely branched ; 
trees or shrubs with usually horizontal branches and 

Petioles of well developed leaves 15-40 (usually 20-30) 

mm. long ; ovary and fruit glabrous ; inflorescences 

terminating leafy branchlets. 

Leaves glabrous and lustrous above, 15-40 cm. long; 

calyx ca. 5 mm. long; stone obliquely ovoid, ca. 

18 mm. long; explanate calyx 10-13 mm. broad 

3. C. fallax. 

Leaves strigose and rather dull above, 10-27 cm. long; 
calyx 2.5-3.5 mm. long; stone transversely com- 
pressed-ovoid, ca. 10 mm. long; explanate calyx 

5-7 mm. broad 4. C. tctrandra. 

Petioles of well developed leaves 3-15 (usually 5-10) 



JOURNAL OF THE ARNOLD ARBORETUM Ivol. x 

mm. long; inflorescence usually borne at the forks of 
the dichotomous stems. 
Ovary and style hairy; fruit mostly pubescent. 

Leaves glabrous above or practically so; veins less 
conspicuously rebranched than in next; calyx 
usually apiculate, opening somewhat irregularly. 
Stems with conspicuous subnodal swellings that 
serve as ant-domatia ; calyx tending to disinte- 
grate at maturity and showing a fibrous struc- 
ture; plant usually conspicuously bristly 

••••; 5. C. nodes. 

Stems without subnodal swellings, not myrmeco- 

philous ; calyx not with fibrous structure; plant 

not bristly. 

Fruit glabrous ; calyx with a fine minute strigose- 

puberulence; lower leaf-surfaces glabrous 

or practically so 6. C. lacvifron 

Fruit strigose; calyx strigose ; lower leaf-sur- 
faces pubescent. 
Lower surfaces of leaves evidently bearing 
numerous erect slender hairs; inflo- 
rescence stiffish but loose and open. . .7. C. Spruce 
Lower surfaces of leaves apparently glabrous, 
but really bearing scattered minute in- 
conspicuous very short ascending hairs; 
inflorescence dense with short rigid 

crowded branches 8. C. ncrvosc 

Leaves abundantly hairy above, veins repeatedly re- 
branched; calyx opening by 5 triangular lobes. 
Calyx prominently and regularly 10-ribbed, 4-5 

ovoid 9. C. fnlrc 

Calyx not ribbed, 2-4 mm. long. 
Upper surface of leaves velvety, with very abund- 
ant slender erect or ascending hairs ; leaves 
usually strongly dimorphic about the stem- 
forks (the normal elongate leaves usually 
opposed by much smaller suborbicular 
ones) ; fruit clothed with abundant slender 
appressed usually tawny hairs ; stone ascend- 
ing, ovoid 10. C. ioqucn 

Upper surface of leaves simply strigose or mi- 
nutely scabrous ; leaves homomorphic. 
Lower surface of leaves green (drying 
brown), .scabrid with very short sparse 
hairs; leaves lanceolate; fruit minutely 
strigose ; stone ascending ovoid 

•■■•■ 11. C. scahnjolic 

Lower surface of leaves pallid, with a feltv 



JOHNSTON, STUDIES IN BORAGINACEAE 7 

hairs; leaves broadly lanceolate to ovate; 
fruit glabrescent ; stone transversely ovoid 

12. C. bicolor. 

)vary, style, and fruit glabrous. 
Lower surface of leaves pallid with a felty covering 

of abundant slender appressed hairs 12. C. bicolor. 

Lower surface of leaves not felty with a pallid 

indument. 

Stone globose or depressed globose, quite rugose ; 

calyx with abundant long slender hairs on the 

inner surface which project beyond the edge of 

the calyx-lobes and appear as a dense pale cili- 

ation on their margins. 

Calyx outside covered with abundant slender 

silky hairs ; leaves rather thin, more or less 

dimorphic at the stem-forks, lower surface 

much paler than the upper 13. C. sericicalyx. 

Calyx sparsely strigose outside ; leaves rigid, 
homomorphic, lower surface scarcely paler 

than upper 14. C. panicularis. 

Stone ovoid or ellipsoid, smooth, erect, elongate; 
calyx strigose on the inner surface, the hairs 
projecting beyond the calyx-lobes sparse and 

Hairs on lower leaf-surface erect. 

Lower leaf-surface somewhat scabrid with 

minute stout hairs; leaves large, 15-28 

cm. long, with evidently falcate midrib ; 

branchlets with short erect hairs . . .15. C. Sagotii. 
Lower leaf-surface velvety with long slender 

hairs; leaves moderate-sized, 8-18 cm. 

long, midrib weakly falcate; branchlets 

Hairs on lower leaf-surface appressed. 

Flower-buds elongate, obovoid, 4-5 mm. long ; 

leaves 8-20 cm. long; inflorescence large 

and stiffish ; Lower Amazon and the 

Guianas. 

Leaf-blades broadest at or above middle, 

". . . 17. C. exaltata. 

Leaf-blades broadest at or below the middle 
drying a bright warm brown 

17a. var. melanonenra. 

Flower buds subglobose, 2-3 mm. long ; leaves 

6-11 cm. long; inflorescence slender and 

usually small; Upper Amazon . . .18. C. naidophila. 

la-lobes distinctly broader than long ; calyx cupulate or 

ylindrical at maturity ; inflorescence dense, globose or 

picate, or exceptionally a small loose cyme ; shrubs 



JOURNAL OF THE ARNOLD ARBORETUM 



witli erect or ascending I)ranche>^ 


;, frequently 


sub- 




scandent. 








Corolla very large, 35-50 mm. long, 


, the slender 


tube 




abruptly and much expanded into i 


I coarse cylindrical 




throat; inflorescence capitate; tip i 


of calyx-lobe. 


; con- 




spicuously long-attenuate 


19. 


C. granc 


iiflora 


Corolla small, about 5 mm. long or le; 


is, tube and t 






weakly differentiated. 








Inflorescence glomerate or cymose 


20. 


C. polycephala 


Inflorescence distinctly spicate. 








Leaves not hairy above, merely more or less v 


erru- 




cose or muriculate, elongate 


; spikes tern 


linal ; 




petioles not decurrent on the 


peduncles 








21. C, 


, macrost 


achya 



decurrent on the subtended peduncle. 
Calyx-lobes in bud lacking free tips ; inflorescence 
becoming quite loosely flowered and elongate ; 
upper surface of leaves strigose, somewhat 

lustrous 22. C. Schomburgkii. 

Calyx-lobes in the bud with projecting free tips : 
inflorescence dense and stout ; upper surface 
of leaves with stiff erect or ascending hairs 
which arise from bulbous bases, surface not 
lustrous. 
Calyx hairy all over, lobes narrowly triangular. 

long acuminate 2i. C. tomentosa. 

Calyx with tube nearly glabrous, lobes broadly 

triangular, short acuminate 24. C. muUispicaia. 

1. Cordia alliodora (R. & P.) Chamisso ex DeCandolIe, Prodr. 9: 
472 (1845); Johnston, Contr. Gray Herb. 92: 13 (1930). Cerdana 
alliodora Ruiz & Pavon, FI. Peruv. 2: 47, tab. 184 (1799). Cardm 
trichotoma sensu Sandwith, Kew Bull. 1933: 335 (1933). 

Tree up to 20 m. tall; branchlets sparingly to densely stellate- 
pubescent; leaves oblong or lanceolate to elliptic, usually broadest at or 
above the middle, i-2, cm. broad, 1-2 dm. long, base acute or obtuse, 
apex acuminate, margin entire, upper surface stellate-pubescent or 
glabrate, lower surface paler, stellate-tomentose or glabrescent, 5-7 
pairs of veins, petiole 1-3 cm. long; inflorescence terminal, loosely and 
widely branched, 1-3 dm. thick, the flowers crowded on the branches, 
the axis commonly inflated, gall-like, irregular, usually serving as an 
ant-domatium; calyx cylindrical, with ten prominent ribs, densely 
stellate-tomentose, 4-6 mm. long, 2-2.5 mm. thick, lobes 5, inconspicu- 
ous; corolla white, drying brown, marcescent, lobes oblong, 5-7 mm. 
long, 1.5-3.5 mm. broad, spreading; fruit sausage-shaped with fibrous 



1935] JOHNSTON, STUDIES IN BORAGINACEAE 9 

chartaceous wall, ca. 5 mm. long, completely envelop)ed by the tube of 
the persistent corolla and by the ensheathing calyx-tube and falling 
away enclosed by them. 

Headwaters of the Rio Branco in northern Brazil and adjacent south- 
ern British Guiana; northern Venezuela and Colombia and southward 
along the Andes and northward in Central America and the West Indies. 

British Guiana: north side of Kanuku Mts., ca. 10 miles east of the 
Takutu River, ca. 135 m. alt., small tree, 4.5 m. tall, trunk ca. 8 cm. thick, 
in secondary forest near edge of savanna, fl. pure white, Oct. 10, 1931, 
Forest Dept. Brit. Guiana D230/2221 (K) ; Pirara (Marakanata), Rupu- 
nuni Savannas, ca. 120 m. alt., tree ca. 20 m. tall, trunk 11 m. to fork, 
4 dm. thick, in sandy soil on patch of savanna-forest on top of ridge, Oct. 
21, 1931, Forest Dept. Brit. Guiana D195/2186 (K). 

Brazil: Mniam, tributary of Suruma River, Nov. 1909, Ule 8290 (K, 
BD, Del) ; Limao, lower Cotinga River, Sept. 1927, Tate 140 (NY). 

Although previously I have cited one of the above collections as C. 
trichotoma, Contr. Gray Herb. 92: 15 (1930), I am now of the opinion 
that all the material from the upper Rio Branco watershed is more 
closely related to C. alliodora. The corolla-lobes in our plants are 3-3.5 
mm. broad. The stems are simply tunneled by ants. There are no dis- 
torted, gall-like thickenings in the axis of the inflorescence. Compared 
with large series of C. trichotoma and C. alliodora our plants seem most 
like the latter species in gross aspect. The colony on the Rio Branco 
was probably derived from northern Venezuela where only C. alliodora 
is known. Strangely C. alliodora seems to be rare or absent in the Ori- 
noco Valley and in the other parts of the wet tropical forests of north- 
eastern South America. 

Previously I have attempted to maintain the Argentine, Paraguayan 
and Brazilian plant, ranging to the east and south of the Amazon Basin, 
as a species distinct from C. alliodora. I am now of the belief that this 
plant, called C. trichotoma in my treatment of the Brazilian species, is 
distinguished from C. alliodora only by its larger flowers, and that it had 
best be classified as a variety of that latter species. The correct tri- 
nomial for the large-fiowered Brazilian form is Cordia alliodora var. 
tomentosa A. DC. 

2. Cordia sebestena Linnaeus, Sp. PI. 190 (1753). Cordia spe- 
ciosa Salisbury, Prodr. Ill (1796); DeCandolle, Prodr. 9: 476 (1845) ; 
Pulle, Enum. PI. Surinam 397 (1906). 

Tree or shrub 1^7 m. tall; branchlets with a fine soft curly pu- 
bescence and scattered much coarser appressed hairs; leaves ovate to 
elliptic or subcordate, 9-16 cm. long, 5-14 cm. broad, broadest below 
the middle, base obtuse or rounded or subcordate, apex obtuse to coarsely 



10 JOURNAL OF THE ARNOLD ARBORETLM [vol. xvi 

short-acuminate, margin entire, upper surface with scattered short stiff 
appressed hairs, the hairs usually arising from minute pustulate disks, 
lower surface glabrescent or sparsely strigose, with 5-6 pairs of veins; 
petiole slender, 1-4 cm. long; inflorescence corymbose, usually ter- 
minal, ascendingly branched; calyx firm, strigose and densely brown 
puberulent, elongate in the bud, 12-15 mm. long, 3-5 mm. thick, open- 
ing by several unequal teeth ca. 2-3 mm. long, at maturity becoming 
much expanded by the enlarging fruit which it encloses, 3-4 cm. long; 
corolla orange or scarlet, funnelform, tube twice length of the eylindrical 
calyx; fruit bony, dry, ovoid, pointed, 1-2 cm. long, completely and 
tightly invested by the juicy white accrescent calyx. 

Native on the islands of the Caribbean and probably also along the 
coasts of Venezuela, Colombia and Central America; frequently culti- 
vated in the Tropics. 

British Guiana: Botanic Gardens, Georgetown, cultivated, Aug. 1905, 
collector not given 7976/6915 (BD). 

Dutch Guiana : Surinam, cultivated, tree (y-9 m., fl. red, Dec. 1837, 
Splitgerber 312 (Leid) ; Cottica district near Plant. Alliance, Aug. 1901, 
Went 280 (Utr) ; Paramaribo. Fockc 1371 (Utr). 

French Guiana: indefinite. 1802, Gabriel (Del). 

3. Cordia fallax, sp. nov. Cordia guianensis Klotzsch ex Schom- 
burgk. Fauna u. Fl. Brit. Guian. 960 (1848), nomen; not C. gujanensis 
(Desv.) R. & S. (1819), nor C. guianensis R. & S. ex DC. (1845). 

Arbor 5-10 m. alta; ramulis brunnescentibus cum pilis abundantibus 
brevibus erectis velutinis; foliis homomorphis ellipticis vel obovato- 
oblongis 15-40 cm. longis 6-18 cm. latis ad medium vel paullo supra 
medium latioribus minute glanduloso-punctulatis, basi rotundis vel sub- 
cordatis ad obtusis vel late acutis, apice breviter acuminatis, margine 
integerrimis vel rariter leviter sinuatis, supra lucentibus in costa et 
nervis primariis pilos inconspicuos gerentibus ceteris glabris vel sub- 
glabris, subtus pilis plus minusve abundantibus gracillimis ascendentibus 
molliter vestitis, nervis 7-8-jugatis, costa falcato-curvatis, petiolis 15-30 
mm. longis; cymis ramulos foliatos terminantibus laxissime ramosis ad 
3 dm. diametro; calyce in alabastro obovoideo extus indumento brun- 
nescente velutino molli vestito, intus glaberrimo, ad anthesin ca. 5 mm. 
longo (lobis deltoideis 5), fructifero explanato 10 14 mm. lato; corolla 
1 cm. longa, lobis obovatis extus glabris. filamentis basim versus pilosis; 
ovario et stylo glaberrimo; fructu glabro; nuce valde rugoso oblique 
ovoideo acuminato ca. 18 mm. longo. 

Endemic to British Guiana. 



1935] JOHNSTON, STUDIES IN BORAGINACEAE 11 

1920, Hitchcock 17563 (type, Gray Herb.; isotype, NY) ; upper Rupununi 
River near Dadanawa, ca. lat. 2° 45' N., tree 5 m. tall, June 10, 1922, La 
Cruz 1484 (NY); indefinite, 1844, Schomburgk 875/1510b (K) ; indefi- 
nite, 1841, Schomburgk 875 (BD, Del, P) ; indefinite, Schomburgk 1510 
(BD, TYPE of C. guianensis; G). 

Although bearing various numbers and different data, the material 
from Schomburgk cited above agrees so completely in details of ma- 
turity, pressing, etc., that one may recognize it as consisting of parts of 
a single collection. The material at Berlin bears Klotzsch's binomial. 
This, however, has never been associated with a description and is fur- 
ther invaHd by reason of being a homonym. Schomburgk, 1. c, reported 
C. guianensis KI. only from the banks of the Barama River and there 
is every reason for believing that this is indeed the source of the Schom- 
burgk material mentioned. 

The species has been confused with C. tetrandra, although it is readily 
distinguished from that species by having glabrous upper leaf-surfaces 
and in being noticeably larger in all its parts. Its relations are with that 
group of species of Venezuela and Colombia which is exemplified by C. 
bogotensis Benth. Its very large leaves, hairy beneath, quickly dis- 
tinguish it from these much more westerly species. 

4. Cordia tetrandra Aublet, Hist. PI. Guian. Fr. 1:222, tab. 87 
(1775); Poiret, Encyc. 7: 42 (1806); PuUe, Enum. PI. Surinam 397 
(1906); Johnston, Contr. Gray Herb. 92: 55 (1930). Lithocardium 
tetrandrum (Aubl.) Kuntze, Rev. Gen. 2: 976 (1891). Cordia cordi- 
jolia Humboldt, Bonpland & Kunth, Nov. Gen. et Sp. 3: 70 (1818); 
DeCandolle, Prodr. 9 : 483 ( 1845) . Lithocardium cordijolium (HBK.) 
Kuntze, Rev. Gen. 2: 976 (1891). Cordia muneco Humboldt, Bonpland 
& Kunth, Nov. Gen. et Sp. 7: 207 (1825); DeCandolle, Prodr. 9: 486 
(1845). Lithocardium muneco (HBK.) Kuntze, Rev. Gen. 2:977 
(1891). Borellia asper Rafinesque, Sylva Tellur. 41 (1838). Cordia 
umbraculijera DeCandolle, Prodr. 9: 484 (1845); Schomburgk, Fauna 
u. Fl. Brit. Guian. 960 (1848); Fresenius in Martius, Fl. Bras. 8^ : 16 
(1857). Lithocardium umbracuUjerum (DC.) Kuntze, Rev. Gen. 
2: 977 (1891). 

Tree, 3-12 m. tall; branchlets pallid, tomentose with abundant curved 
spreading short hairs; leaves homomorphic, ovate to elliptic or oblong 
or lance-ovate, broadest either below or just above the middle (usually 
the latter), 1-1.7 dm. long, 5-14 cm. broad, base more or less oblique, 
obtuse or rounded or subcordate, apex obtusish to acute, the very tip 
blunted (not acuminate), under surface green, sparsely strigose, sec- 
ondary venation obscure, lower surface much paler, more or less brown- 
ish with rather abundant short slender curved hairs which spring from 



12 JOURNAL OF THE ARNOLD ARBORETUM Tvol. xvi 

the much rebranched veins, usually velvety, with 7 10 pairs of veins; 
petioles well developed, 2-5 cm. long; cymes usually terminating leafy 
branchlets, loosely branched, 1-3 dm. broad; calyx obovoid in bud, 
densely covered with fine appressed hairs, inside sparsely strigose or 
hispidulous; calyx at anthesis 2.5-3.5 mm. long, with 4-5 more or less 
equal deltoid lobes, in fruit explanate and 5-7 mm. broad; corolla white, 
prevailingly 5-merous, 4-5 mm. long, glabrous, lobes elongate, filaments 
exserted, hairy at base; ovary and style glabrous; fruit glabrous; stone 
very rugose, transversely compressed-ovoid, ca. 1 cm. long, pulp white 

Northeastern coast of Brazil (Maranhao and Para), northern South 
America and southward along the Andes to Bolivia; frequently cul- 

British Guiana: Rockstone, banks of the Essequibo, 1921, Glcason 
865 (K) ; Demerara River, May 1889, Jenman 4878 (K) ; Demerara, 
Parker (K, DC) ; Platburg Creek, Canje River, fruit glutinous, yellowish 
green, 1914, Hohcnkcrk 631 (K) ; indefinite, large tree, flowers yellowish 
white, 1837, Schomburgk 408 (DC, type of C. umbracuUfera; isotypes, 
G, K, BD, P). 

Dutch Guiana: upper Nickerie River, Feb. 1915, B. IV. 1074 (Utr) ; 
near Paramaribo, 1910, native collector (Utr) ; near Paramaribo, tree, 
fl. white, 1844, Kappler, ed. Hohenacker 1619 (Utr, P) ; Plant. Jagtlust, 5 
m. tall, 1913, Soeprato 6E (Utr); Plant. Osembo-Onverwacht, 1913, 

B. W. 6229 (Utr) ; Plant. Slootwijk, tree 5 m. tall, Soeprato lOH (Utr) ; 
Watramiri, tree no. 1568, fruit edible, mucilaginous, June 4, 1916, B. W . 
1836 (Utr) ; Watramiri, tree no. 1568, Feb. 7, 1917, B. W. 2659 (Utr) ; 
Watramiri, tree no. 1568, Feb. 18, 1920, B. W. 4551 (Utr) ; Watramiri, 
tree no. 1568, Dec. 7, 1920, B. IV. 4974 (Utr) ; Surinam, tree 9-12 m. tall 
with broad horizontal branches, usually cultivated, fl. white, Nov. 1837, 
Splitgerher 123 (Leid) ; Surinam, 1841, Berthoud-Conlon 553 (BM) ; 
Surinam, Hostmann 355 (K, BM, BD, Del, P) ; Surinam, Hostmann 
(Utr). 

French Guiana: Mana, March 1854, Melinon 215 (P) ; Mana. 1857, 
Sagot (P) ; lies du Salut, fruit white, glutinous, 1854, Sagot 445 (K, BM, 
P); He de Cayenne, 1851, Sagot (P) ; Cayenne, Aublet (BM. type of 

C. tetrandra) ; Mahoury near Cayenne, Sagot (P) ; indefinite, Martin ex 
herb. Rudge (BM), LePrieur 252 (Del, P), Perrottet (P), Gabriel (Del) 
and Poitcau (K). 

Aublet reports C. tetrandra from the He de. Cayenne and from the 
mainland of French Guiana. I have examined specimens from his per- 
sonal herbarium, now at the British Museum, as well as a duplicate from 
it now in the Swartz herbarium at Stockholm. His description, his 
illustration, and the two specimens, leave no doubt as to the exact iden- 
tity of C. tetrandra. Aublet's name is inapt, the species is practically 
always pentandrous. 



19351 JOHNSTON, STUDIES IN BORAGINACEAE 13 

The species appears 'to be indigeneous only in a broad band of wet 
tropical forest about the northern margin of South America, where it 
seems to be most common at low altitudes on the coastal plain. On the 
east base of the Andes it is known from Peru and Bolivia. I have seen 
no material from the dryer portions of Brazil to the south of the Amazon 
Basin that is indubitably from wild plants. Brade, Bol. Mus. Nac. Rio 
Janeiro 8: 35 (1932), however, has recently reported it from Manaos. 
The collections by Spruce from the mouth of the Rio Negro, reported 
(sub C. umbraculijera) in the Flora Brasiliensis, 8^ : 16 (1857), is not 
C. tetrandra, but the material subsequently made the type of C. Sprucei 
Mez. I have reported, Contr. Gray Herb. 92: 55 (1930), C. tetrandra 
from Ceara and Pernambuco. This was incorrect and is the result of a 
bad clerical error. The specimens actually represent C. toqueve. I am 
indebted to Mr. Killip, in lit., and to Mr. Brade, 1. c. 34, for the correc- 
tion of this unfortunate error. 

The wood of C. tetrandra has been described by Pfeiffer, De Hout- 
soort. V. Surinam. 1 : 444 ( 1926). In the herbarium at Utrecht the col- 
lection, cited above, from the Plantation of Osembo-Onverwacht bears 
the annotation, "Pfeiffers Woods of Surinam no. 59." 

Aublet states that the species is called "Bois Margarite" and "Arbre 
a parasol." The following vernacular names are associated with the 
specimens cited above, Kakuru or Clammy C\xexry-^Hohenkerk 631 ; 
Kakhoro' (Arow.), Tafrabom (Nig. Eng.) and Alatoeloeka (Kar.) — 
B. W. 107 A; Tafelboom— 5. W. 6229 and Split gerber 123; Tafelboom 
(Sur. Dutch), Tafraboom (Nig. Eng.), Boggi lobbi (Saram.), Toen- 
balobbi (Saram.), Kakhoro (Arow.), Araatroekoe {K^v.)—Watramiri 
tree no. 1568; Bois parasol— 5a^of 445; Roquei— Sagt)^ at Cayenne 
and Mahoury. 

5. Cordia nodosa Lamarck, Tab. Encyc. 1:422 (1791); Poiret, 
Encyc. 7:43 (1806); Schomburgk, Fauna u. Fl. Brit. Guian. 960 
1848); Fresenius in Martins, Fl. Bras. 8M 16, tab. 5 (1857); Bailey, 
Bot. Gaz. 77: 32-49, tab. 6-7 (1924); Johnston, Contr. Gray Herb. 
92:46 (1930). Lithocardium nodosum (Lam.) Kuntze, Rev. Gen. 
2: 977 (1891). Cordia hirsuta Willdenow, Sp. PL 1: 1076 (1798); 
Meyer, Prim. Fl. Esseq. 114 (1818). Firensia hirsuta (Willd.) 
Rafinesque, Sylva Tellur. 40 ( 1838) . Cordia jormicarum Hoffmannsegg 
ex Roemer & Schultes, Syst. 4: 800 (1819). Cordia miranda DeCan- 
dolle, Prodr. 9: 475 (1845). Lithocardium mirandum (DC.) Kuntze, 
Rev.' Gen. 2:977 (1891). Cordia hispidissima DeCandolle, Prodr. 
9: 475 (1845). Lithocardium hispidissimum (DC.) Kuntze, Rev. Gen. 
2: 977 (1891). Cordia nodosa var. hispidissima (DC.) Fresenius in 



14 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

Martius, Fl. Bras. 8' : 17 (1857). Cordia nodosa var. angustijoUa Fre- 
senius in Martius, Fl. Bras. 8' : 17 (1857). Cordia umbrosa Spruce ex 
Rusby, Bull. Torr. Bot. CI. 26: 147 (1899). Cordia volubilis Pittier, 
(Explor. Bot. Cuenca de Maracaibo p. 41) Bol. Comer, e Indust. 4: ? 
(1923); Jour. Wash. Acad. Sci. 19: 184 (1929). Cordia collococa 
sensu Aublet, Hist. PI. Guian. Fr. 1: 219, tab. 86 (1775). 

Shrub or tree, 2-11 m. tall; stems bearing stiff spreading brownish 
bristles which are usually abundant but may be sparse or nearly absent; 
the stems below each fork abruptly and asymmetrically enlarged and con- 
taining a cavity usually serving as an ant-domatium; leaves usually 
subopposite or whorled, more or less heteromorphic, somewhat lustrous 
on both surfaces, lanceolate to nearly elliptic, broadest near the middle, 
10-35 cm. long, 3-28 cm. broad, base obtuse, apex acuminate, margin 
entire, upper surface with impressed veins, more or less bullate, with a 
few hairs along the midrib, lower surface paler, with very scattered 
bristles on the veins, with 6-10 pairs of veins, these repeatedly re- 
branched and anastomosing, petiole 2-5 mm. long, bristly; inflorescence 
cymose-paniculate, loose or dense, 2-10 cm. in diameter, bristly and 
usually also with minute curly brownish pubescence, borne at the forks 
of the stem; calyx usually somewhat puberulent and strigose, more or 
less bristly especially about the apiculate apex, papery in texture and 
very obscurely ribbed, opening irregularly to form several very irregular 
lobes, frequently persisting and eventually breaking up into fibers ; calyx 
in the bud ovoid or elHpsoid, ca. 5 mm. long; corolla white, tube 4-6 mm. 
long, lobes 2-3 mm. long, filaments hairy at base, 3-4 mm. long; style 
and ovary hairy; fruit usually more or less bristly; stone transversely 
ovoid, 13-17 mm. long. 

In British, Dutch and French Guiana and widely distributed in the 
Amazon Basin; also in the headwaters of the Orinoco (in southern Vene- 
zuela and eastern Colombia) and in northwestern Venezuela. 

British Guiana : Amakura River, 5 m. tall, March 1923, La Cruz 3430 
(G); Barima River, March 1896, Jcnnian 7055 (K) ; Kamakusa, upper 
Mazaruni, ca. long. 59" 50', 1922-23, La Cruz 2887 and 4231 (G) ; Macouria 
River, Nov. 1886, Jenman 2391 and 2392 (K) ; Tumatumari, dense upland 
forest, shrub 2.5-6 m. tall, 1921, Glcason 311 (K) ; Kaieteur Falls, Potaro 
River, 1923, La Cruz 4407 (G) ; island in Cuyuni River below Kamaria 
Falls, 18 dm. tall, 1920, Bailey 40 (G) ; Kartabo region, second growth 
forests, 1920, Bailey 29, 41 and 42 (G) ; Bonasika Creek, at sea-level, 
Anderson 66 (K) ; Moraballi Creek, small tree up to 6 m. tall, in low 
brush and clearings in mixed forest, fl. white, fruit bristly, becoming pale 
red, Aug. 15, 1929, Sandwith 72 (K. BD) : Rockstone, dense upland forest, 
2.5-3 m. tall. 1921. Glcason 583 (K) ; Blue Mts., Demerara, fruit red. 



1935] JOHNSTON, STUDIES IN BORAGINACEAE 15 

River, May 1889, Jenman 4924 (K) ; Malaroo Creek, Corantyne River, 
small tree, 3-6 m. tall, Oct. 1879, im Thurn (K, P) ; indefinite, Schom- 
burgk 904 (K) and 984 (K, BM, BD, P). 

Dutch Guiana : Kaboeri Reserve, Corantyne River, tree no. 684, Nov. 
1920 and Aug. 1922, B. W. 4835 and 5986 (Utr) ; mouth of Lucie River, 
Corantyne River, 1910, Hulk 315 (Utr) ; way to Kwatta, Paramaribo, 
June 1916, Samuels 237 (G, Leid, BD, P) ; Station at Groningen, forest. 
May 1916, Sanmels 123 (G, K, Leid, BD, P) ; Watramiri reserve, Sara- 
macca River, June 1918, B. W . 3864 (Utr) ; Watramiri reserve, tree no. 
1652, mature fruit yellow, soft and sweet, Dec. 1916, B. IV. 2488 (Utr) ; 
Watramiri, tree no. 1652, used for tea, May, 1916, B. W. 1911 (Utr) ; 
Watramiri reserve, tree no. 1652, April 1917, Oct. 1917, July 1918, and 
Feb. 1920, B. W. nos. 2756, 3309, 3872 and 4541 (Utr) ; Watramiri reserve, 
tree no. 1652, fl. 'white, B. W. 4012 (Utr); Watramiri reserve, tree no. 
1652, March 1919, ripe fruit sordid white, B. W. 4301 (Utr) ; Sectie O. 
reserve, upper Para River, tree no. 800, fl. light green, leaves used for tea, 
Aug. 1916, B. IV. 2306 (Utr) ; woods near Poelebantji, tree 4-6 m. tall, 
Feb. 1845, Kegel 691 (Utr) ; Brownsberg. Surinam River, tree 10 m. tall, 
trunk 1 dm. thick, fl. sordid white, Sept. 1915, B. W. 727 (Utr) ; Browns- 
berg reserve, tree no. 1174, fl. white, dried leaves used as a substitute for 
tea, fruit yellow, globose, soft and juicy, Nov. 1916, B. W. 2498 (Utr) ; 
Brownsberg reserve, tree no. 1174, fl. white, Sept. 1918, B. IV. 4002 
(Utr); Brownsberg reserve, tree no. 1174, March 1917, Feb. 1919 and 
March 1921, B. IV. nos. 2721, 4265 and 5075 (Utr) ; Brownsberg reserve, 
tree no. 1174. fl. sordid white, with strong odor, Sept. 1923, B. W. 6227 
(Utr) ; Brownsberg reserve, tree no. 1174, fl. sordid white, odor strong, 
Nov. 1924, B. W. 6684 (Utr) ; Brownsberg summit, July 1924, fl. white, 
B. W. nos. 6634 and 6722 (Utr) ; woods near Raleigh Falls, Coppename 
River, hispid tree, fl. yellowish white, fruit white, hispid, Sept. 11, 1933, 
Lanjouw 788 (Utr). 

French Guiana: Maroni, 1864, Melinc 
Maroni River, Sagot (P) ; He Portal, fruit 
June 1857, Sagot 446 (P) ; Acarouani, tree, corolla pale yellowish, sepals 
4, stamens 4, Oct. 1854, Sagot 446 (P) ; Acarouani, 1854, Sagot (P) ; 
Acarouani, fruit red, April 1858, Sagot 446 (P) ; Acarouani, 1859, Sagot 
(P); Acarouani, fruit pale yellow, 1854, Sagot 446 (P) ; Godebert, 
Wachenheim 410 (F) ; in loco Macaya ad praedicem Patuis, Richard (P) 
vicinity of Cavenne, hill above Grant's Road, Montabo, shrub, 1921, Broad- 
way 543 (G, k) ; Cayenne, Martin (BM, BD, P) ; Cayenne, March 1859, 
Sagot (P) ; Cavenne, Patris (BM, Del) ; indefinite, 1850, Leprieur (BM, 
P) ; indefinite, Auhlet (BM, type of C. nodosa) ; indefinite, Perrottet 214 
(Del, DC) and Poiteau (K, BD, P). 

Brazil: Carmo, Rio Branco, Sept. 1, 1924, Bequaert (G) : Surumu, 
Serra do Mairary. Rio Branco, tree or shrub 2-8 m. tall, fl. white, Nov. 
1909, Ule 8456 (K, BD) ; Rio Negro below mouth of Xibaru, betw. Bar- 
cellos and Sao Gabriel. Dec. 1854 Spruce 3790 (NY, K, Del) ; Rio Cumina, 
Sampaio 5136 and 5148 (BD) ; near Montalegre, Nov. 24, 1873, Traill 
561 (K) ; Prainha, Dec. 17, 1873, Traill 562 (K). 



16 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

tall, Jan. 1853, Spruce 3281 (G). 

This is a classic ant-plant. The results of an anatomical study and a 
review of the more important literature on this plant have been published 
recently by I. W. Bailey, Bot. Gaz. 77: 32-49, tab. 6-7 (1924). Accord- 
ing to this author the peculiar subnodal structures serving as ant-domatia 
are "formed by an invagination of epidermal, cortical, and fibrovascular 
tissues which originate in the axil of one of the leaves of the false verti- 
cil, and which develops into the interior of a more or less symmetrical 
or unilateral, subnodal enlargement of the cauline axis." 

The attention drawn by the complex ant-domatia has, I believe, 
blinded students to the evident relationships of this remarkable plant. 
The flowers and fruit are very similar to those found in C. Sprucei and 
its relatives. The apiculate, papery, irregularly disrupted calyx, the 
hairy ovary and style, and the transversely ovoid stone, not to mention 
the glabrous upper leaf-surfaces, comparatively stiff and contracted 
inflorescence, etc., all indicate close relations with that group of upper 
Amazon and Guianan species. Cordia nodosa has only three notable 
peculiarities, its subnodal swellings, its bristly indument, and its fibrous 
calyx. In the past the species has been placed in a special section, 
Physoclada, of the genus Cordia. I have become so impressed with its 
obvious relations with C. Sprucei and allies, however, that I now am 
quite content to associate it with these species in the section Pilicordia. 

The species is very variable both in the size and shape of its leaves, 
and in the abundance of the bristles on its herbage. This variation 
seems to be ecological in origin. In any case I can find no evidence that 
it is in any way geographically correlated. It should be noted that 
collectors have given the fruit in British Guiana as red. In Dutch and 
French Guiana the fruit is given as white or whitish in numerous cases, 
and once as yellow. 

Aublet gives the Carib name for the plant as "Achira-mourou." The 
following vernacular names are associated with specimens cited: Coura- 
belli ants T^Xsini— Anderson 66; Yluri-hee-levT-kou— Par/^er 272; 
Hurneyreyroko — Sandwith 12; Awelemoeloe (Kar.) — B. W. 727; 
Marribonsoehoedoe (Neg. Eng.), Horowejoreroko (Arow.), Arreuonoe 
(Kar.)— Tree no. 1652 at Watramiri; Mattoe toenbalobbi (Sar.), 
Horowe, joee lokko, Hoereuereroko (Arow.), and Awali emoeloe, Aloeko 
uonore (Kar.)— Tree 800 at Sectie 0. 

6. Cordia laevifrons, sp. no v. 

Arbor minor vel frutex, dichotome ramosus; ramulis fuscis, apicem 
versus dense puberulis mox glabrescentibus; foliis vix crassis ellipticis 



1935] JOHNSTON, STUDIES IN BORAGINACEAE 17 

ad lanceolato-oblongis vel oblongo-obovatis 12-25 cm. longis 6-14 cm. 
latis saepe ad medium vel supra medium latioribus, basi obtusis vel plus 
minusve rotundis vel late acutis, apice saepe abrupte acuminatis, supra 
lucentibus saepe in costa pilos paucos adpressos gerentibus ceteris 
glaberrimis, subtus pallidioribus glabris vel sparsissime minutissimeque 
ascendenter adpresseque pubescentibus, nervis primariis 6-8-jugatis, 
nervis tertiariis obscuris, petiolis 5-18 mm. longis; cymis saepissime in 
furcis ramulorum ortis, laxe graciliterque ramosis 3-15 cm. diametro, 
pedunculo gracili; corolla alba glabra, tubo 6 mm. longo calycem 
superante, lobis 2 mm. longis rotundis latis, filamentis 4 mm. longis 
longe exsertis basim versus pilosis; calyce in alabastro anguste obovato 
4-5 mm. longo extus dense puberulento (intus subglabro) obscure 
lateque 10-costato, apice plus minusve apiculato ad anthesin in lobos 
irregulares lacerulatos disrupto, fructifero explanato; stylo et apice 
ovarii sparse minuteque hispidulo; fructu glaberrimo; nuce transverse 
ovoideo 10-14 mm. longo. 

Endemic to French and Dutch Guiana. 

Dutch Guiana: Lucie River, a small tree 6 m. tall, fl. white, April 12, 
1926, B. W. 6999 (Utr) ; forest near Abontjeman, May 1910, natwe col- 
lector 236 (type, Utrecht). 

French Guiana : Maroni River, 130 km. upstream, fruit edible, 1877, 
Crevanx (P) ; along the Maroni, 1861, Melinon 16, 59, 254 and 271 (P) ; 
along the Maroni, 1863, Melinon 283 (P) ; Maroni, along road to St. Lau- 
rent, clearings, 15 dm. tall, fl. white, Oct. 1876, Melinon 225 (P) ; St. Jean, 
2 m. tall, fl. white, May 16, 1914, Benoist 1230 (P). 

A relative of C. Sprucei notable chiefly for its rather thin, nearly 
glabrous leaves, puberulent obscurely ribbed calyces, and quite glabrous 
fruits. As with other relatives of C. Sprucei the veins of the leaves are 
not so finely rebranched as is common in this section of the genus. 
Crevaux gives the bush-negro name of the plant as "Tiki Topichi." 

7. Cordia Sprucei Mez, Bot. Jahrb. 12:549 (1890); Johnston, 
Contr. Gray Herb. 92: 53 (1930). Lithocardium Sprucei (Mez) 
Kuntze, Rev. Gen. 2: 977 (1891). 

Tree 4-5 m. tall, branching dichotomous; branchlets dark, sparsely 
short-hirsute or with short incurved hairs; leaves drying brown, sub- 
homomorphic, elliptic or oblong-obovate, 12-23 cm. long, 6-11.5 cm. 
broad, broadest at or above the middle, apex acuminate, base acute to 
truncate, upper surface somewhat lustrous, hairy along the midrib and 
with scattered hairs along the principal veins but otherwise glabrous, 
lower surface with rather abundant short soft erect hairs, petiole 5-10 
mm. long, veins in 6-9 pairs, tertiary veins tending to be obscure; in- 



18 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

florescence rather loosely though rigidly branched, ca. 1 dm. thick, 
usually borne at the forks of the stem, rarely terminal; calyx well cov- 
ered with short incurving tawny hairs, obscurely ribbed, obovate in the 
bud and apiculate, 4-5 mm. long, ca. 2.5 mm. thick, bursting rather 
irregularly at the apex into 2-5 broad rather thin teeth ; corolla white, 
tube 4-5 mm. long, lobes broad ca. 1.5 mm. long, filaments hairy at 
base, 4 mm. long; ovary densely hairy above the middle; fruit yellow, 
strigose; stone transversely ellipsoid, 1-1.5 cm. long. 

Known only from the Rio Negro of Brazil and from French Guiana. 

French Guiana : "in Sylvis doeciduis Fluvii Kourou, ad casam indi 
Felix," Nov., Richard (P). 

Brazil: Barra do Rio Negro, 1850-51, Spruce 1019 (Munich, type; 
BD, frag; G, photo.) ; vicinity of Barra, 1850-51, Spruce (G, K, BM) ; 
Barra to Matiriho, Jan. 1851, Spruce 1234 (K, BM) ; Barra, fruit yellow, 
transversely oblong, April 1851, Spruce 1234 (K, BM, Del) ; Sao Gabriel, 
Rio Negro, ca. 90 m. alt., 1930-31, Holt & Blake 608 (G). 

Richard's label gives the following field data concerning his collection 
from French Guiana, — "frutex 3-4 ped., ramis diffuse patentibus, dicho- 
tomis; fl. albidi; ramillis cymae recurvis et, inexpansis floribus, revo- 
lutis." The collections are remarkably similar to Spruce's material from 
the lower Rio Negro, except that in one of Richard's two sheets the 
branches of the inflorescence are somewhat tawny tomentulose. 

Since discussing the type of C. Sprucei, 1. c, I have examined the 
actual type-specimen at Munich. The specimen has the following 
familiar printed label reading, "In vicinibus Barra, Prov. Rio Negro, 
coll. R. Spruce, Dec-March, 1850-51." The collector's number, in 
script, is "1019." The specimen is that cited under C. umbraculifera in 
the Flora Brasiliensis. It is evidently part of the same collection as the 
unnumbered specimens I have seen at the Gray Herbarium, at Kew, and 
at the British Museum. 

This species not only has relatives in C. nervosa and C. laevijrons of 
the Guianas but also in undescribed trees of the Putumayo and the 
Huallaga of eastern Peru. The glabrous upper leaf-surfaces and the 
somewhat papery irregularly disrupted apiculate calyces are characters 
of this group of species. 

8. Cordia nervosa Lamarck, Tab. Encyc. 1:422 (1791); Poiret, 
Encyc. 7: 47 (1806); DeCandolle, Prodr. 9: 484 (1845). Lithocardium 
nervosum (Lam.) Kuntze, Rev. Gen. 2: 977 (1891). Cordia calophylla 
Vahl, Ecolog. 3: 5 (1807); DeCandolle, Prodr. 9: 486 (1845). Litho- 
cardium calopkyllum (Vahl) Kuntze, Rev. Gen. 2: 976 (1891). 

Shrub or small tree, up to 5 m. tall; branchlets closely and antrorsely 
strigose; leaves homomorphic, stiff and coriaceous, with an arcuate mid- 



1935] JOHNSTON, STUDIES IN BORAGINACEAE 19 

rib, broadly lanceolate to elliptic or lance-oblong, 10-25 cm. long, 4-10 
cm. broad, margin weakly recurved, apex acuminate, base rounded to 
acute and usually more or less oblique and asymmetrical, upper surface 
glossy, smooth and quite glabrous, lower surface drying brown, dull, 
somewhat scabrous with abundant short inconspicuous hairs, with 8-10 
pairs of primary veins, these connected by simple branches, the secondary 
branches of the veins absent or very obscure ; petiole canaliculate, stiff, 
5-10 mm. long; inflorescence small and compact, 1-4 cm. long, peduncles 
very short or none, branches slender, strictly forked, bearing flowers on 
only one side and in age studded with the elevated pedicellar flower- 
attachments, becoming rigid and woody in age and more or less spread- 
ing or deflexed, persistent long after the falling of the fruit; calyx obo- 
void in bud, 4-5 mm. long, minutely short-strigose, more or less apicu- 
late, not at all ribbed, sparsely strigose inside, bursting apically and the 
lobes torn and irregular, in fruit explanate; corolla white, tube ca. 5 mm. 
long, lobes broad, ca. 3 mm. long, filaments very hairy; ovary glabrous 
or sparsely hairy towards the apex; style usually sparsely hairy; fruit 
minutely and abundantly strigose, pulp bright red, glutinous, insipid; 
stone transversely ovoid, 10-13 mm. long. 

French Guiana and adjacent Brazil ; British Guiana. 

British Guiana: Kaieteur Savanna, spreading shrub 18 dm. tall, 1881, 
Jcmnan 1062 (K). 

French Guiana: Cayenne, 1857, Melinon (P) ; Gourdonville, Kourou 
River, shrub, fl. white, Sept. 25, 1914, Benoist 1618 (P) ; in umbrosis sylvis 
praedii Dm. [ ?] Patuis, Richard (P) ; indefinite, herb. Lamarck (Paris, 
TYPE of C. nervosa) ; indefinite, von Rohr 152 (herb. Vahl, type of C. 
calophylla; BM, isotype) ; indefinite, 1859, Leprieur (Del); indefinite, 
1819-21, Poiteau (K, Del). 

Brazil: Counany, Oct. 13, 1895, "Chapeo del Sol," Hubcr 1032 (Boiss). 

The type of C. nervosa in the Lamarck herbarium is so very similar 
to the material collected by Richard (in the General Herbarium at Paris) 
that I believe they are parts of a single collection or, in other words, 
that the type of C. nervosa was collected by Richard. Unfortunately, I 
have been unable to identify Richard's locality with any degree of con- 
fidence. At Paris I found on the label of a very different species the 
following more explicit mention of the probable locality, i. e. "in loco 
Dm. [spelling?] Macaya ad praedicem Dm. [?] Patuis." There was 
formerly a sugar plantation called Macaya on the He de Cayenne sev- 
eral kilometers east of Matoury. This may have been that referred to 
by Richard, for he is known to have collected extensively about the 
Island of Cayenne as well as over most of the French Guianan coastal 



20 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

It is interesting to note that Richard has appended to his specimen 
a manuscript name, under Collococcus, in which the same specific epithet 
is used as was subsequently published by Vahl. Richard and von Rohr 
were both in the Guianas about 1785. These facts naturally make one 
wonder if there may not have been some meeting or some exchange of 
material between these two botanists and possibly if Vahl's type may 
not have had the same source as that of Lamarck. 

The species is an unusually distinct one, being notable because of its 
suppressed tertiary leaf-veins and small dense subsessile inflorescence. 
Its closest relations are with C. Sprucei and C. laevijrons which have 
similar somewhat papery, irregularly dehiscent apiculate calyces. In 
C Sprucei the veins are more repeatedly branched than in C. nervosa, 
though generally less so than in other species of the section Pilicordia. 

9. Cordia fulva, sp. nov. 

Arbor vel frutex, dichotome ramosus; ramulis brunneis cum pilis 
brevibus divergentibus abundantissimis velutinis; foliis subhomomor- 
phis crassiusculis ellipticis vel ovatis 10-22 cm. longis 7-11 cm. latis 
saepe ad medium vel supra medium latioribus, basi obtusis vel rotundis 
vel rariter acutis, apice breviter acuminatis, supra scabris pilis brevibus 
rigidusculis ascendentibus vestitis, subtus saepe brunnescentibus velu- 
tinis in nervis et nervulis elevatis et numerosis pilos graciles erectos 
abundantes gerentibus, nervis 5-7-jugatis, petiolis brevibus; cymis in 
furcis ramulorum ortis vel rariter terminalibus, rigidis brunneo-velu- 
tinis laxe ramosis, floribus in ramulis plus minusve congestis; calyce in 
alabastro obovato 4-5 mm. longo 2-i mm. crasso evidenter 10-costato 
extus brunneo-velutino intus supra medium strigoso; lobis 5 deltoideis; 
corolla alba, tubo 5 mm. longo, lobis ca. 2.5 mm. longis et latis, filamentis 
4-5 mm. longis basim versus pilosis; ovario apicem versus dense 
longeque pilosis; fructu evidenter velutino; nuce transverse ovoideo 
1-1.5 cm. longo. 

Known only from northern Dutch Guiana and adjacent French 
Guiana. 

Dutch Guiana 
lector 227 (Utr) ; , 
(Utr). 

French Guiana: Maroni, shrub 3 m. tall, in clearings, fl. white, 
branches horizontal, 1877, Melinon 137 (type, Gray Herb.; isotype, 
Paris) ; Maroni, Melinon ^55 (G, P) ; Maroni, Wachenheim 75 (G, P) ; 
indefinite, 1862, Melinon 82 (P) ; indefinite, 1821, Perrottet (P). 

Related to C. trichoclada DC. and C. Ckamissoniana Don, of eastern 
Brazil, this species differs from the former in its velvety rather than 



1935] JOHNSTON, STUDIES IN BORAGINACEAE 21 

bristly stems, more softly hairy scarcely bullate leaves, looser less stiffly 
branched larger inflorescences and slightly smaller calyces, and from 
C. Chamissoniana in its more hairy leaves and conspicuously ribbed, 

10. Cordia toqueve Aublet, Hist. PI. Guian. Fr. 1:228, tab. 90 
(1775); Poiret, Encyc. 7:44 (1806); DeCandolle, Prodr. 9:488 
(1845); Johnston, Contr. Gray Herb. 92: 52 (1930). Litkocardium 
toqueve (Aubl.) Kuntze, Rev. Gen. 2:977 (1891). Cordia hetero- 
phylla Poiret, Diet. Sci. Nat. 10: 409 (1818) ; Willdenow ex Roemer & 
Schultes, Syst. 4:800 (1819); Chamisso, Linnaea 4:480 (1829); 
DeCandolle, Prodr. 9:487 (1845). Litkocardium heterophyllum 
(Poir.) Kuntze, Rev. Gen. 2: 977 (1891). Cordia pubescens Will- 
denow ex Roemer & Schultes, Syst. 4:800 (1819). Litkocardium 
pubescens (Willd.) Kuntze, Rev. Gen. 2:977 (1891). Toquera to- 
mentosa Rafinesque, Sylva Tellur. 40 (1838). Cordia hebecarpa 
DeCandolle, Prodr. 9: 488 (1845). Litkocardium kebecarpum (DC.) 
Kuntze, Rev. Gen. 2:977 (1891). 

Tree; branchlets velvety with abundant soft more or less curled 
brown hairs ; leaves strongly dimorphic, upper surface with rather abund- 
ant short straight ascending hairs, lower surface velvety with curved 
soft slender spreading hairs from the prominent and numerous veins 
and veinlets; larger principal leaves very broadly lanceolate to lance- 
ovate, broadest towards the base, 1-3 dm. long, 6-15 cm. broad, above 
the middle contracted to an acute or acuminate apex, base obtuse to 
rounded, somewhat oblique; smaller sort of leaves more or less orbicular, 
5-12 cm. long, 5-11 cm. broad, broadly obtuse or even subcordate at 
base, rounded or acuminate at apex; inflorescence loosely and slenderly 
branched, 1-1.5 dm. broad; calyx obovoid in bud, 2.5-3 mm. long, open- 
ing by 5 triangular lobes, unribbed, strigose inside, outside covered with 
a dense indument of appressed slender curved hairs; calyx becoming 
somewhat cupulate at maturity, ca. 1.5 mm. deep; corolla white, tube 
2.5 mm. long, lobes 1.5 mm. long, filaments 2 mm. long, hairy near base; 
ovary densely hairy at apex; fruit abundantly tawny-strigose, style- 
base forming a short eccentric beak; stone ovoid, strictly ascending, 
ca. 1 cm. long. 

Confined to French Guiana and eastern Brazil. 

French Guiana : vicinity of Cayenne, small tree, fruit yellowish, May 
16, 1921, Broadway 201 (G) ; near Cayenne, fl. yellow, 1897, Soubiron 
(P); near Cayenne, July 1841, Melinon 243 (Leid, P) ; Cayenne, Feb. 
1859, Saqot (P) ; Cayenne, Martin (K) ; Cayenne, Leblond, ex Mus. 
Paris 348 (BD, P) ; Cayenne, ex Mits. Paris sine no. (G. BD) ; Cayenne. 



22 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

Herb. Willd. sub no. 4574 (BD, type of C. heterophylla WiUd.) ; Cayenne, 
Herb. Poiret (P, type of C. heterophylla Poir.) ; indefinite, Aublet (BM, 
TYPE of C. toqueve); indefinite, Poiteau (K) ; indefinite, 1859, Leprieur 
(Del). 

The original material of this species was collected by Aublet in clear- 
ings made by the Caribs about 15 leagues up the Sinnamary River. 
The tree was called "Toqueve" by these inhabitants of French Guiana. 
I have seen Aublet's specimen at the British Museum. This material, 
taken in conjunction with Aublet's illustration and lengthy description, 
leaves no doubt as to the correctness of the present application of the 

At Paris among Poiret's specimens (in the Cosson collections) there 
is a fragmentary specimen of the present species accompanied by a label 
in Poiret's script reading, Cordia heterophylla, folia altera majora et 
minora, rami asperi, hirti. Caienne. Added to the label in another, and 
unrecognized hand is "diet, des Sc. nat. herb. Poiret." Poiret stated 
that Cordia heterophylla was seen in the Desfontaines herbarium. The 
specimen in the Poiret collections at Paris is, I believe, a fragment of 
the type of C. heterophylla Poir., now probably conserved at Florence. 

The binomial "C. heterophylla" is found on a specimen of C. toqueve 
in Willdenow's herbarium at Berlin and was published by Roemer & 
Schultes a year after Poiret's published use of the name. The specimen 
is also given as from "Cayenne" and may be a duplicate of the specimen 
described by Poiret. 

Cordia toqueve is readily distinguished among the South American 
species by its tawny velvety indument, strikingly dimorphic leaves, and 
conspicuously hairy fruit. It is known only from Bahia, Ceara and 
Pernambuco in Brazil, and from near Cayenne in French Guiana. The 
Brazilian plant, which is quite like that from the Guianas, has been de- 
scribed as C. hebccarpa DC. 

11. Cordia scabrifolia A. DeCandolIe, Prodr. 9: 485 (1845) ; John- 
ston, Contr. Gray Herb. 92: 53 (1930); Brade, Bol. Mus. Nac. Rio 
Janeiro 8; 34 (1932). 

Tree or shrub up to 15 m. tall, branching dichotomous; branchlets 
dark, abundantly and minutely antrorse-strigose ; leaves homomorphic, 
ovate- to oblong-lanceolate, 11-18 cm. long, 4-7 cm. broad, broadest 
near the middle, apex acuminate, base acute to somewhat rounded; upper 
surface drying dark, abundantly and very minutely antrorse-strigose, 
lower surface drying light, bearing numerous very minute very short 
appressed hairs on the abundant veins and veinlets, the hairs tending 
to be directed centripetally towards the middle of the veinlet-areoles, 



1935] JOHNSTON, STUDIES IN BORAGINACEAE 23 

petiole 4-9 mm. long, veins in 5-8 pairs, repeatedly rebranched; in- 
florescence usually borne at the forks of the stems, pedunculate, slender, 
loosely branched, 4-10 cm. broad; calyx obovoid in the bud, ca. 3 mm. 
long, densely strigose, opening by 5 subequal triangular lobes, strigose 
inside, base substipitate ; corolla white, tube ca. 3 mm. long, lobes ovate, 
ca. 2 mm. long, filaments exserted, hairy at base; style and at least the 
apex of the ovary hairy; fruit densely and minutely strigose; stone 
ovoid, ascending, ca. 1 cm. long. 

Probably restricted to the Amazon Basin; doubtfully reported from 
British Guiana. 

British Guiana: indefinite, Schomburgk 911 (Boiss, type; isotypes, 
K, BM, BD, P). 

Brazil: Prainha, Nov. 1873, Traill 563 (K) ; Rio Cumina, Oct.-Nov. 
1928, Sampaio 5367, 5505 and 5510 (BD) ; Rio Negro-gapo above Cabu- 
quena, Dec. 1851, Spruce 1942 (K, BM). 

Except for the type-collection, Schomburgk 911, which is given as 
from British Guiana, all known collections of C. scabrijolia come only 
from within the Amazon Basin. I suspect that the type also came from 
the Amazon watershed and from what is now Brazil. Schomburgk, 
Fauna u. Fl. Brit. Guiana 960 (1848), reports the species from the 
upper Essequibo. It is significant, however, that no other collectors 
have found it in British Guiana and that, in the notes of Robert Schom- 
burgk, plant no. 911, which certainly seems to apply to this species, 
refers to a collection almost certainly from the Rio Negro watershed. 
In the list no. 911 has no locality indicated but the adjacent numbers, 
where the locality is indicated in several instances, all do come from 
Barcellos on the Rio Negro. 

12. Cordia bicolor A. DeCandolle, Prodr. 9:485 (1845); Pulle, 
Enum. PI. Surinam 397 (1906). Lithocardium bicolor (A. DC.) 
Kuntze, Rev. Gen. 2: 976 (1891). Cordia dichotoma Klotzsch ex 
Schomburgk, Fauna u. Fl. Brit. Guiana 1084 (1848), nomen; not Forst. 
(1797). Lithocardium Lockartii Kuntze, Rev. Gen. 2:438 (1891). 
Cordia Lockartii Kuntze, Rev. Gen. 2:438 (1891), in synonymy. 
Cordia trichostyla Pittier, Contr. U. S. Nat. Herb. 18: 252, fig. 102 
(1920). Cordia carnosa Rusby, Three Hundred N. Sp. So. Amer. PI. 
104 (1920). Cordia coriacea Killip, Jour. Wash. Acad. Sci. 17: 329 
(1927). Cordia sericicalyx sensu Johnston, Contr. Gray Herb. 92: 54 
(1930). 

Shrub or small tree up to 6 m. tall, branching dichotomous; branch- 
lets angulate, velvety with very abundant spreading usually tawny short 
hairs; leaves homomorphic, ovate to more or less broadly lanceolate, 



24 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

broadest at or below the middle, 8-16 cm. long, 2.5-7 cm. broad, apex 
acuminate, base acute to rounded, upper surface dark, finely strigose, 
lower surface very pale, covered by appressed slender hairs that are 
borne on the veins and veinlets and which converge over and cover the 
veinlet-areoles; veins in 5-7 pairs, repeatedly rebranched; petiole 5-10 
mm. long; inflorescence usually at the forks of the stem, loosely 
branched; calyx in bud ca. 4 mm. long, obovoid, clothed with abundant 
short appressed more or less tawny hairs, inner surface with numerous 
appressed longer white hairs, opening by 5 subequal triangular lobes; 
corolla white, tube ca. 3 mm. long, lobes ca. 2.5 mm. long, filaments 
hairy near base; style and upper part of ovary hairy or these exception- 
ally glabrous; fruit glabrescent; stone transversely ovoid, over 1 cm. 
long. 

Occurring in the Amazon headwaters of Bolivia; east and south of 
the Amazon Basin in Brazil; and across northern South America from 
Dutch Guiana to eastern Colombia; apparently very sporadic in occur- 
rence, widely distributed but not common. Also in Central America and 
in the southern-most West Indies. 

British Guiana: Oreala, Corantyne River, Oct. 1879, Jcmnan 7 (K) ; 
indefinite, Schombiirgk "109" (K) ; indefinite, Schomburgk 601 (BM). 

Dutch Guiana: indefinite, Hostmann 406 (DC, type of C. bicolor; 
isotypes, K, BD, Del, P) ; indefinite, Hostmann & Kappler 406 (Munich) ; 
indefinite, Kappler 406 (P). 

Brazil: Roraima, 1200 m. alt., corolla white, Feb. 1910, Ule 8748 (K, 
BD); Roraima, 1842-43, Schomburgk 678 (Del, P) ; indefinite, Schom- 
burgk 678/1032 (K) ; indefinite, Nov. 1842, Schomburgk 1032 (BD, type 
of C. dichotoma). 

Venezuela: Lower Orinoco, 1896, Rusbv & Squires 418 (NY, type 
oiC.carnosa). 

In my paper on the Brazilian cordias I quite incorrectly applied the 
name C. sericicalyx to this present species. The type of C. sericicalyx 
is Schomburgk 109. There is material of C. bicolor at Kew and the 
British Museum, however, which bears Schomburgk's no. 109. Mr. 
Killip has suggested to me that this may have resulted from inverting 
no. 601. Reexamination of the specimens makes me believe this sug- 
gestion is correct, especially since one finds in Robert Schomburgk's 
notes (for nos. 589-700 incl.), which were forwarded from Curasscuraka 
(on the Rupununi at the base of the Annai Hills) on Feb. 1838, that 
no. 601 bears the following appropriate data, "a shrub with brown bark 
growing like the table tree, leaves green, below silvery, calyx light 
green, petals white, filaments and anthers cream-colored." No locality 
for no. 601 is given, but it is certain that the gamut of numbers to which 



1935] JOHNSTON, STUDIES IN BORAGINACEAE 25 

it belongs was collected either during the exploration south to the 
Akarai Mts., on the Brazilian border, or about the Annai savannas. 

Schomburgk's nos. 678 and 1032 appear on printed labels as from 
"Roraima." Schomburgk approached the base of that mountain from 
the south (Brazilian territory) along the Kukenam River. It will be 
noted that Schomburgk, Fauna u. Fl. Brit. Guiana 1084 (1848), reports 
C. dkhotoma only from the banks of that Brazilian stream. 

13. Cordia sericicalyx A. DeCandolle, Prodr. 9: 485 (1845) ; Pulle, 
Enum. PI. Surinam 397 (1906); not Johnston, Contr. Gray Herb. 92: 
54 (1930). Litkocardium sericicalyx (A. DC.) Kuntze, Rev. Gen. 
2:977 ( 1891 ) . Cordia sericicalyx var. latijolia Miquel, Stirp. Surinam. 
140 (1850). CorrfwierewwBritton, Bull.Torr. Bot. C1.50: 54 (1923). 

Tree 3-6 m. tall, branching dichotomous; branchlets slender, finely 
strigose; leaves conspicuously heteromorphic, firm but rather thin and 
smooth, much paler beneath, finely strigose on both surfaces, veins re- 
peatedly branched and anastomosing: larger leaves oblong to broadly 
lanceolate, 11-25 cm. long, 5-12 cm. broad, usually broadest below the 
middle, base obtuse or broadly acute, midrib somewhat arcuate, veins 
in 7-11 pairs, petiole densely strigose, 5-20 mm. long; smaller leaves 
elliptic to orbicular-ovate, 6-8 cm. long, 3-7.5 cm. broad; cymes loose, 
5-20 cm. broad, branches slender, usually borne at the forks of the 
stems; calyx sessile, 5-toothed, outside completely covered with minute 
appressed silky hairs, inside above the middle densely villous-strigose 
with the hairs projecting a little beyond the edge of the calyx and so 
appearing as ciliate margins of the lobes; calyx at anthesis 3-4 mm. 
long, 1.5-2.5 mm. broad, unribbed, in fruit becoming explanate; corolla 
ca. 5 mm. long, glabrous, lobes obovate, filaments exserted, pilose near 
base; ovary and style glabrous; fruit glabrous; stone very rough, de- 
pressed, weakly asymmetric 8-10 mm. broad. 

Ranging from Dutch Guiana westward to the Orinoco and apparently 
to Trinidad. 

British Guiana: indefinite, Schomburgk 109 (DC, type of C. serici- 
calyx; isotypes BD, P). 

Dutch Guiana: Wayombo River near Cornells Kondre, tall tree, fl. 
greenish white, Jan. 23, 1915, B. W. 824 (Utr) ; Para District, in forest, 
Feb.-April, 1844, Kappler 1510 (Utr, type of var. latijolia; isotypes, Leid, 
BD, Del, P). 

Venezuela: lower Orinoco. 1896, Rusbv & Squires 282 (NY, K) and 
259 (K). 

In my paper on the Brazilian species of Cordia I incorrectly applied 
the name C. sericicalyx to the concept properly called C. bicolor. Some 



26 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

of the details of this confusion will be found discussed under C. bicolor 
in the present paper. 

The type of C. serkkalyx is Schomburgk 109, given merely as from 
British Guiana. The species is not listed in Schomburgk's "Fauna und 
Flora von Britisch Guiana," although possibly it may be the basis for 
the otherwise unintelligible report of C. heterophylla, 1. c. 960 (1848), 
which is listed as on the Demerara and Essequibo. Of all the cordias 
collected by the Schomburgks, Cordia serkkalyx has the most pro- 
nounced dimorphic leaves and hence is the one most apt to be associated 
with the appropriate but incorrect name, C. heterophylla. 

The type of C. serkkalyx var. latijolia Miquel has been examined. It 
is not separable from the typical form of the species. Miquel gives the 
type of his variety as "Kappler 1500." The type-specimen, however, is 
clearly numbered "1510"/ 

In defining C. serkkalyx I have excluded a very closely related form 
which seems to replace our Guianan plant across .the northern parts of 
Venezuela and Colombia. This form, called C. opaca Rusby, is a more 
slender plant with more elongate, frequently more or less ribbed, fulvous 
calyces, very sparsely hairy, short filaments, and firmer, usually more 
lanceolate leaf-blades. 

The specimen of C. serkkalyx from Cornells Kondre is associated 
with two vernacular names. These are Omose (Kar.) and Kakoro 

14. Cordia panicularis Rudge, PI. Rar. Guian. 30, tab. 46 (1805). 
Lithocardium pankulare (Rudge) Kuntze, Rev. Gen. 2: 976 (1891). 

Shrub or large tree, 3-20 m. tall, dichotomous or trichotomous; 
branchlets drying dark, sparsely strigose, loosely branched; leaves sub- 
homomorphic, lucent, elliptic to lance-elliptic or broadly lanceolate, 1-2 
dm. long, 4-7.5 cm. broad, broadest at or just below the middle, mature 
leaves usually drying quite brown, apex long-acuminate, base obtuse, 
both surfaces sparsely strigose or beset with minute very short ascending 
hairs arising from inconspicuous pustulate bases, lower surface slightly 
the paler; veins in 5-6 pairs, evident, repeatedly rebranched; petiole 
5-10 mm.; cymes loose, 5-30 cm. broad, borne at the forks of the stem; 
calyx sessile, 5-toothed, 4-5 mm. long, subcylindric or obconic-cylindric, 
terete or very obscurely angulate, outside rather sparsely strigose, inside 
densely villous-strigose, becoming explanate in fruit; corolla glabrous, 
ca. 8 mm. long, lobes obovate to oblong, 2-3 mm. long, filaments hairy 
near base; ovary and style glabrous; fruit glabrous; stone very rugose, 
depressed, weakly asymmetric, 8-10 mm. broad. 

Known only from the Guianas. 



1935] JOHNSTON, STUDIES IN BORAGINACEAE 27 

British Guiana: Potaro Landing, clearings and roadsides, shrub 3^.5 
m. tall, 1921, Gleason 259 (NY, K) ; Tumatumari, dense upland forest, 
tall shrub, almost vine-like, 1921, Gleason 156 (NY, K) ; Bootooba, sand- 
hill in forest, Oct. 1924, Per sand i^-^ (BD, NY). 

Dutch Guiana: near Patricksavanna, in forest. May 1910, native col- 
lector 182 (Utr) ; Brownsberg Summit, June 1924, tree 20 m. tall, B. W 
6513 Knd 6519 (Utr.). 

French Guiana: Acarouani, fl. white, 1857, Sagot 448 (K, P) ; Aca- 
rouani, 1854, Sagot 448 (P) ; Acarouani, medium-sized tree or shrub, Oct. 
1856, Sagot (K, P) ; Cayenne, Martin ex herb. Rudge (BM, type) ; 
Cayenne, Martin (K, P) ; Cayenne, Martin 151 (P). 

The type of C. panictdaris is a young flowering branch bearing leaves 
not yet stiff and somewhat coriaceous as they become in the mature 
state. Sagot has distributed under an unpublished name, sub no. 448, 
collections of this species made over several years which show both the 
young and the mature foliage. Although quite different in appearance 
I am confident that C. panictdaris and C. sericicalyx are close relatives. 
The rough depressed glabrous fruits in these species are notably similar. 
The specimen from Brownsberg Summit is given as called "Berg 
Tafraboom." 

15. Cordia Sagotii, sp. nov. Cardia coriacea Sagot ex Benpist, 
Archives Bot. 5, Mem. 1: 257 (1933), not Killip (1927). 

Frutex vel arbor 5-10 m. alta, dichotome ramosa; ramulis scabridis 
pilos minutos numerosos erectos vel ascendentes e basi incrassatos geren- 
tibus; foliis homomorphis ellipticis vel late lanceolatis vel oblongis rigide 
coriaceis 15-30 cm. longis 7-14 cm. latis medium versus latioribus, basi 
rotundis vel obtusis vel acutis aliquantum obliquis, apice breviter acumi- 
natis, supra sparse inconspicueque brevi-strigosis, subtus pilis numerosis 
brevissimis erectis asperatis, nervis 6-8-jugatis abundanter ramosis; 
petiolis 5-10 mm. longis; cymis saepe in furcis ramulorum ortis sed 
rariter ut videtur lateralibus, laxissime ramosis 1-3 dm. crassis; calyce 
in alabastro obovoideo ca. 3 mm. longo sparse striguloso, lobis triangu- 
laribus 5; corolla alba fragrante, tubo 4-5 mm. longo, lobis 1.5-2 mm. 
longis, filamentis 5-6 mm. longis basim versus pilosis; ovario et stylo 
glaberrimis; fructu glabro flavo; nuce anguste ovoideo erecto ca. 15 mm. 
longo laevi. 

Known only from northeastern Dutch Guiana and northwestern 
French Guiana. 

Dutch Guiana: Sectie O. Reserve, fl. white, Nov. 14, 1917, B. W. 
3414 (Utr) ; Sectie O. Reserve, tree no. 505, Oct. 23, 1916, B. W. 1194 

(Utr) ; Sectie O. Reserve, tre ^" " " '" 

345 (Utr) ; Zanderii I. Rese 



28 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

1915, Oct. 1917, Nov. 1919, April 1920, Jan. 1921 and Nov. 1921, B. W. 
1137, 3357, 4433, 4631, 5037 and 5565 (Utr) ; Zander! j I. Reserve, tree no. 
230, fl. white, fragrant, Feb. 1, 1917, Feb. 14, 1917, July 1917, Oct. 1917, 
and Nov. 1918, B. W. 2679, 2673, 3026, 3372 and 4067 (Utr.) 

French Guiana: banks of the Maroni, 1861, Melinon 243 (P) ; Gode- 
bert, 1920-21, Wachenheim 81 and 207 (P) ; Charvein, fl. white fragrant, 
shrub 10 m. tall, Dec. 9, 1913, Benoist 318 (P) ; Acarouani, medium-sized 
tree, mature leaves very coriaceous fl.. white, fragrant, Dec. 1856, Sagot 
447 (type of C. coriacea Sagot and C. Sagotii, Paris) ; Acarouani, 1856, 
Sagot 447 (K, P) ; Acarouani, tall shrub, fl. white, fragrant, mature leaves 
very coriaceous, 1857, Sagot 447 (K, BD, P) ; Acarouani, 1855, Sagot 447 
(K) ; indefinite, 1863, Melinon 63/96; indefinite, 1862, Mclmon 412 (P) ; 
indefinite, 1802, Gabriel (Del). 

This species is represented in many herbaria of Europe by various 
collections of Sagot (all numbered 447). These are all determined as 
C. coriacea Sagot, a name unpublished until Benoist recently used it in 
his work on the timbers of French Guiana. Benoist has given an ade- 
quate botanical description (in French) of Sagot's species. The lengthy 
description of the wood-structures and the vernacular name, given by 
him, however, are based upon Melinon 88 which represents Cordia hirta. 
Since Benoist's formal botanical description is evidently based upon 
Sagot's collections and since his binomial was also evidently derived 
from them, I am content to accept Sagot's species as formally established 
by Benoist's description, especially since the description was not drawn 
to include Melinon's herbarium specimen. Among the several collections 
of Sagot, obtained in various states of maturity over several years (but 
all distributed under the same collection-number) I have selected the 
specimen at Paris collected in Dec. 1856 as the type of C. coriacea Sagot. 
Unfortunately the name C. coriacea Sagot is a homonym of an earlier 
published name. Furthermore its publication by Benoist, because of 
the lack of a Latin diagnosis, is invalid. For these reasons the plant is 
described as a new species, C. Sagotii. The type is the same as that 
selected for C. coriacea Sagot. 

The species is readily recognized by its very large rigid coriaceous 
leaves which have the midrib markedly bowed and the lower surface 
scabrous with minute short erect hairs. The glabrous fruit is erectly 
ovoid and yellow when mature. 

The following vernacular names have been found on the labels accom- 
panying specimens from Dutch Guiana— Tafraboom (Surinam Dutch), 
Kakoro (Arow.), Anoemalatti (Nig. Eng.), Danlieba (Sar.), Dokoa or 
Dokka (Arow.) and Anaakara (Kar.) — B. W. 345 ; Tafrabom, Arow- 
troeka (Arow.) and Kokoro konokodikoro (Kar.) — B. W. 1194; Tafel- 
boom (Surinam. Dut.), Tafraboom (Nig. Eng.), Boggi lobbi and 



1935] JOHNSTON, STUDIES IN BORAGINACEAE 29 

Toenbalobbi (Sar.), Kakoro (Arow.) and Aratroeka (Kar.)— Tree 176 
at Zanderij I. 

16. Cordia hirta, sp. nov. 

Arbor summum ad 20 m. alta, dicho- vel trichotome ramosa; ramulis 
brunneis saepe fulvo-hirsutis; foliis subhomomorphis ellipticis vel 
oblongo-oblanceolatis ad medium vel supra medium latioribus 8-18 cm. 
longis 3.5-7 cm. latis, basi acutis vel obtusis vel plus minusve rotundis, 
apice abrupte breviterque acuminatis, supra plus minusve scabris pilis 
abundantibus brevibus rigidis ascendentibus vestitis, subtus plus 
minusve velutinis pilis gracilioribus erectioribus ornatis, nervis 7-9- 
jugatis, petiolis hirsutis 4-9 mm. longis; cymis in furcis ramulorum ortis 
laxe ramosis 1-3 dm. crassis pilis rigidis erectis brunneis vestitis; calyce 
sessili in alabastro obovoideo 3-4 mm. longo vix costato extus brunneo- 
hispido intus sparse strigoso, lobis 5 triangularibus erectis ; calyce fruc- 
tifero explanato; corolla 5-6 mm. longa, lobis ca. 1.5 mm. longis rotundis 
recurvatis, filamentis exsertis basim versus pilosis; ovario et stylo glaber- 
rimo; fructu glaberrimo; nuce anguste ellipsoidea valde ascendente 
15-18 mm. longa vix rugosa. 

French Guiana to eastern Venezuela, apparently in the mountains 
back from the coast. 

Dutch Guiana: forest nea 
m. tall, Aug. 26, 1920, Pulle 

B. IV. 6149 (Utr). 

French Guiana: indefinite, 1863, Melinon 88 (P) ; indefinite, 1863, 
Melinon 113 (P). 

Venezuela : Arabopo, slopes of Roraima, ^ mile above Arabopo 
Swamp, 1260 m. alt., Jan. 1, 1928, Tate 259 (type. New York); (??) 
Arabopo, slopes of Roraima, Jan. 1, 1928, Tate 255 (NY). 

This plant evidently frequents the hills. No locality is given for 
Melinon's collections but his vernacular names, "Bois Calalon de mon- 
tagne" and "Cedre Calalon de serre basse" suggest that the specimens 
may have come from high ground up the Maroni where he is known to 
have collected. It is to be noted that the wood sample accompanying 
Melinon 88 is that described by Benoist, Arch. Bot. 5, Mem. 1: 267 
(1933), under the incorrect name of C. coriacea Sagot {= C. Sagotii). 
The species is a well marked one with its closest relation probably in 

C. Sagotii, from which it is quickly separable by its smaller leaves, and 
more copious longer brown pubescence. 

17. Cordia exaltata Lamarck, Tab. Encyc. 1: 422 (1791); Poiret, 
Encyc. 7:47 (1806); DeCandolle, Prodr. 9:484 (1845). Lithocar- 
dium exaltatum (Lam.) Kuntze, Rev. Gen. 2:977 (1841). Cordia 



30 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

mucronata Poiret, Diet. Sci. Nat. 10: 410 (1818). Cordia scabrida 
Martius ex Fresenius in Martius, Fl. Bras. 8' : 11, tab. 9, fig. 12 (1857); 
Johnston, Contr. Gray Herb. 92: 62 (1930). Lithocardium scabridum 
(Mart.) Kuntze, Rev. Gen. 2: 439 (1891). 

Shrub or tree, 2-24 m. tall, branching dichotomous; branchlets dark, 
sparsely strigose; leaves somewhat scabrid, sparsely strigose on both 
surfaces, commonly more or less heteromorphic, usually drying oliva- 
ceous or muddy brown, tertiary veinlets evident; smaller leaves ovate 
to orbicular, 4-9 cm. long; larger leaves elliptic to broadly oblanceolate, 
broadest at or above the middle, 8-20 cm. long, 4-10 cm. broad, apex 
abruptly short-acuminate, base acute to somewhat rounded, veins in 
6-8 pairs, petiole 5-10 mm. long, inflorescence borne at forks of stem, 
loosely branched, 1-2 dm. broad, somewhat scabrous, sparsely strigose or 
finely hirsute ; calyx sessile, sparsely strigose outside, strigose inside the 
tube, pubescent on the inner face of the 5 regular triangular teeth, in 
the bud elongate, 4-5 mm. long, 1.5-2 mm. thick, apex rounded, not 
ribbed, in fruit explanate; corolla white, tube 5-7 mm. long, lobes ca. 
2 mm. long, ovate; filaments long exserted, hairy at base; ovary and 
style glabrous; fruit glabrous, orange-yellow or red; stone ellipsoid, 
erect or nearly so, 1-1.5 cm. long, not irregularly roughened. 

Lower Amazon Valley and along the coast into French Guiana. 

French Guiana: Cayenne, Martin (P); "Cayenne, Martin" (herb. 
Poiret, TYPE of C. mucronata) ; Cayenne, Perrottet (Del, P)j 
Leprieiir sine no. (K, Del, DC) and 164 (K, P) 
Del); indefinite, Richard (K, Del) ' • " ' 
marck, type of C. cxaltata). 



960 (1848),nomen. 

A forma typica differt foliis ellipticis vel oblongo-ovatis medio vel 
infra medium latissimis in sicco plus minusve spadiceis apice longe acu- 
minatis; inflorescentia saepe lata et laxa. 

Known only from middle and western British Guiana. 

British Guiana: Gravee Creek, Kaituma River, N.W. Dist., fairly 
high tree, Oct. 28, 1908, Anderson 103 (K) ; Assakatta, lat. 7° 45' N., long. 
59° 5' W., Sept. 1923, La Crnz 4322 (G) ; Waini River, lat. 8° 20' N., 
long. 59° 40' W., tree 6 m. tall, April 1923, La Cms 3855 (G) ; Santa Rosa, 
Moruka River, Pomeroon Dist., tree 9 m. tall, fl. white, Aug. 1921, La Cruz 
1015 (G) ; Moruka River, July 1927, La Cruz 4522 (G) ; Waramuri 
Mission, Moruka River, 9 m. tall, fl. white, Oct. 1922, La Cruz 2399 (G) ; 
Tabla, Pomeroon Dist., fl. white, Sept. 1921, La Cruz 1224 (G) ; banks of 
the Pomeroon, Aug. 1843, Schomburgk 1398 (BD, type of C. melano- 
ncura) ; Mazaruni River, Aug. 1889, Jcnnvan 5475 (NY, K, BM) ; oppo- 



1935] JOHNSTON, STUDIES IN BORAGINACEAE 31 

site Bartica, April 1887, Jemnan 3625 (K, NY) ; Moraballi Creek near 
Bartica, tree 24 m. tall, trunk 25 cm. thick, fl. white, Aug. 26, 1929, Sand- 
rvith 118 (K); Essequibo River, Aug. 1889, Jenntan 5817 (K) ; indefinite, 
Schomburgk 842/1398b (K), 842 (Del) and 840 (P). 

The type of Cordia exaltata is said to have come from Richard. In 
the General Herbarium at Paris there are two good specimens of this 
species collected by Richard, one bearing mature, and the other very 
young fruit. Neither collection is provided with exact geographical data 
though they are provided with the collector's detailed field-notes. The 
specimen with young fruit, of which Lamarck's type may be a duplicate, 
is given as ''arbor 12-25 pedalis, trunco recto cortice griseo laevissimo, 
ramis expansis." The other specimen has the following (abbreviated) 
field-notes. Arbor 20-40 pedalis, ramis patentibus ramosis saepius ter- 
natis, divaricatis; folia sparsa pulchre viridia; bacca elliptica, laevis- 
sima, glabra, nitens, flavescens; pulpa ^lutinosa viscosa; in sylvis variis; 
dec. fructus maturant. 

Poiret's material of his C. mucronata consists of two small fragmen- 
tary specimens which are so much alike that I believe they are parts of 
a single collection. Leaves, flowers, and fruit are represented. The 
material is undoubtedly conspecific with the type of C. exaltata. Martin 
probably collected it about La Gabrielle to the southeast of Cayenne. 
The species enters French Guiana from the coastal area of Brazil and 
probably does not extend much northwest of Cayenne. 

Cordia scabrida, which has been repeatedly collected in the area about 
the city of Para and also about Santarem further up the Amazon, is 
evidently conspecific with the plant of French Guiana. Exploration will 
no doubt show that it is present along the north bank of the lower 
Amazon and in the coastal forests of Brazil towards the Guianan border. 

The characters I am able to give for the plant of British Guiana, 
which I call Cordia exaltata var. melanoneura, do not separate it sharply 
from typical C. exaltata, but I am of the opinion that the British 
Guianan plant merits at least varietal and possibly even specific rank. 
Geographically, it is separated from C. exaltata by half of French, half 
of British, and all of Dutch Guiana. The leaves usually differ in shape 
and in the characteristic warm brown they assume in drying. Most of 
the material of the species and variety can be sorted rapidly and accu- 
rately merely on the basis of differences in gross aspect. The species 
C. melanoneura first appeared in Schomburgk's book, 1. c, where it is 
said to grow on the banks of the Pomeroon River. This is in the very 
region in which the var. melanoneura has been most collected. The 
following vernacular names appear on herbarium specimens of the vari- 



JOURNAL OF THE ARNOLD ARBORETUM 

Tree—Sandwith 118; 



18. Cordia naidophila, sp. nov. 

Arbor minor dichotome ramosa; ramulis fuscatis dense adpresseque 
cinereo-hirsutulis; foliis homomorphis ovatis vel elliptico-ovatis 6-11 
cm. longis 4-5 cm. latis medium versus vel infra medium latioribus, basi 
acutis vel rotundis, apice graciliter acuminatis, utrinque scabris pilis 
abundantibus brevibus rigidis antrorse adpressis asperatis, nervis 5-8- 
jugatis, nervulis abundanter ramosis, petiolis ca. 5 mm. longis; cymis in 
furcis ramulorum ortis laxe graciliterque ramosis; calyce strigo in ala- 

strigoso, lobis 5 triangularibus; corolla alba tubo 2-3 mm. longa, lobis 
ca. 2 mm. longis, filamentis ca. 2.5 mm. longis basim versus pilosis; 
ovario et stylo glaberrimo; fructu glabro; nuce obovoideo erecto ca. 12 

In the upper Amazon Basin, particularly in the drainage of the Rio 
Negro, Brazil. 

Venezuela : near San Carlos, headwaters of the Rio Negro, 1853-54, 
Spruce 2960 (G). 

Brazil: Manaos, Agricultura 
corolla creamy white, Oct. 13, 
Herb.) ; Barra, Oct. 1819, Ma, 
Ule 5191 (BD). 

This plant of the upper Ama 
vestris Fresen., of the coastal j 
in having the upper leaf-surfaces, dull rather than more or less glossy 
and abundantly rather than very sparsely hairy. The lower surface of 
the leaves in C. naidophila is regularly strigose in a manner quite like 
the upper surfaces. In true C. silvestris the lower face of the leaves is 
very finely strigose and usually coarsely hairy as well ; the upper face is 
sparsely hairy or nearly glabrous. Specimens of C. naidophila have 
been referred to C. silvestris by Fresenius in Martins, Fl. Bras. 8' : 13 
(1857) and by Johnston, Contr. Gray Herb. 92: 61 (1930). 

19. Cordia grandiflora (Desv.) Humboldt, Bonpland & Kunth, 
Nov. Gen. et Sp. 3: 77 (1818) ; Fresenius in Martins, Fl. Bras. 8': 21 
(1857) ; Johnston, Contr. Gray Herb. 92: 21 (1930). Varronia grandi- 
flora Desvaux, Jour, de Bot. 1: 273 (1809); Poiret, Encyc. Suppl. 3: 
730 (1814) ; von Friesen, Bull. Soc. Bot. Geneve, ser. 2, 24: 170, fig. 8 
(1933). Lithocardium grandiflorum (Desv.) Kuntze, Rev. Gen. 2: 977 
(1891). Varronia lantanoides Willdenow ex Chamisso, Linnaea 4 : 492 
(1829), in synonymy. Cordia ruja Klotzsch in Schomburgk, Fauna u. 



1935] JOHNSTON, STUDIES IN BORAGINACEAE 33 

Fl. Brit. Guiana 960 (1848), nomen. Varronia grandiflora var. glabrata 
von Friesen, Bull. Soc. Bot. Geneve ser. 2, 24: 171, fig. 8e (1933). 
Varronia grandiflora var. Sprucei von Friesen, Bull. Soc. Bot. Geneve 
ser. 2,24: 148 (1933), nomen. 

Shrub, 1-3.5 m. tall; stems clothed with slender appressed hairs; 
leaves triangular-ovate to lanceolate, with rather conspicuous veins, 
4-10 cm. long, 1.5-5 cm. broad, base quite obtuse, margin conspicuously 
crenate-dentate, upper surface with slender appressed or ascending hairs, 
usually strigose, lower surface with shorter and more slender hairs, 
petioles 1-2 cm. long; inflorescence capitate, terminal, 1.5-2 cm. thick, 
peduncle 5-15 cm. long; calyx coarsely strigose, 7-10 mm. long, lobes 
long attenuate, the tips linear and free (ca. 2 mm. long) in the bud; 
corolla white, very large, 3.5-5 cm. long, tube 5-9 mm. long and 2 mm. 
thick, abruptly expanded into the coarse cylindrical throat (9-14 mm. 
thick) ; fruit ca. 9 mm. long, elongate, invested by calyx nearly to its 
apex. 

British Guiana to central Venezuela and southward into the Amazon 
Valley. 

British Guiana: Essequibo. Jan. 1842, Schomburgk 358 (BD, type 
of C. rufa) ; Essequibo, Appun 2514 (K) ; Mamette, Rupununi River, Oct. 
\%d,9,Jenman5533 (K). 

Venezuela: Ciudad Bolivar, ca. 35 m. alt., June 1931, Holt & Blake 
740 (G) ; Paloma, lower Orinoco, March 1896, Rusby & Squires 14 (G, 
BM, BD) ; Angostura, Morits (BD). 

Brazil: Rio Trombetas, vicinity of Obidos, shrub ca. 25 dm. tall, grow- 
ing into the water, corolla white, Dec. 1849, Spruce 515 (G, K, BM) ; Rio 
Branco, herbaceous, growing by side of river, corolla white, Sept. 1858, 
Schomburgk 817 (K, BM, Del). 

This remarkable species was first collected by Humboldt & Bonpland 
near San Fernando de Apure, Venezuela, just west of the great bend of 
the Orinoco. The material collected was described as Varronia grandi- 
flora by Desvaux. Cordia rufa Klotzsch is a name based upon a collec- 
tion made by Schomburgk (no. 358) on the Essequibo in British Guiana. 
It has never been described. Schomburgk in his catalogue mentions it 
only from the upper Essequibo. The Schomburgk brothers, however, 
made two collections of this species. A study of Robert Schomburgk's 
field-notes at Kew shows that his collection no. 817, labeled and dis- 
tributed as from British Guiana, is in fact from Brazil, from the Rio 
Branco almost certainly between Sao Joaquin and Pirara. 

20. Cordia polycephala (Lam.), comb. nov. Varronia polycephala 
Lamarck, Tab. Encyc. 1: 418 (1791). Lantana corymbosa Linnaeus, 
Sp. PI. 628 (1753), not C. corymbosa Willd. ex R. & S. (1819). Var- 



34 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

ronia monosperma Jacquin, PI. Rar. Hort. Schoenbr. 1: 18, tab. 39 
(1797). Cordia monosperma (Jacq.) Roemer & Schultes, Syst. 4: 463 
(1819). Varronia dichotoma Ruiz & Pavon, FI. Peruv. 2: 2i, tab. 146 
(1799), not C. dichotoma Forst. (1786). Varronia ulmijolia Jussieu 
ex Dumont-Courset, Le Bot. Cult. ed. 1, 2: 148 (1802), nomen. Cordia 
ulmijolia (Juss.) DeCandolle, Prodr. 9:494 (1845), not C. ulmijolia 
Spreng. (1825). Varronia corymbosa Desvaux, Jour, de Bot. 1: 275 
(1809), not "V. corymbosa L." ex Desf. (1804), nomen. Cordia 
corymbosa (Desv.) Don, Gen. Syst. 4: 383 (1838), not C. corymbosa 
Willd. ex R. & S. (1819). Cordia bijurcata Roemer & Schultes, Syst. 
4:466 (1819). Cordia corymbosa of Urban, Symb. Ant. 4:519 
(1910), and most subsequent authors; Johnston, Contr. Gray Herb. 
92: 30 (1930). Cordia patens sensu Pulle, Enum. PI. Surinam 398 
(1906). — Periclymenum rectum, salviae joliis majoribus, etc., Sloane, 
Nat. Hist. Jamaica 2: 83, tab. 194, fig. 3 (1725). Ulmi angustijoliae 
jacie Baccijera Jamaicensis etc., Plukenet, Phytogr. tab. 328, fig. 5 
(1691) and Almag. Bot. 393 (1694). 

Slender shrub, 1-5 m. tall, frequently subscandent ; stems with 
appressed or somewhat spreading indument of intermixed short and long 
hairs; leaves ovate to lanceolate, 2-12 cm. long, 1-5 cm. broad, base 
acute to obtuse or nearly rounded, apex acute, margin toothed or sub- 
entire, upper surface bearing numerous minute limy tuberculations, very 
sparsely strigose, under surface usually brown, commonly finely and 
densely tomentulose, on the veins bearing slender appressed or ascend- 
ing coarser hairs; petiole slender, 3-10 mm. long, decurrent 1-3 mm. on 
the subtended branchlet and peduncle; cymes usually densely glom- 
erate, subglobose, 5-15 mm. thick, rarely expanding and becoming 
loosely flowered and the branches more or less evidently dichasial and 
scorpioid; peduncles axillary and terminal; calyx strigose. 2-3 mm. 
long, lobes broad, the tips not free in the bud, at maturity calyx tightly 
investing the fruit nearly to the apex; corolla white, 4-5 mm. long; 
fruit red, stone 4-5 mm. long, ovoid or subglobose. 

Very widely distributed in the warm parts of America. 

British Guiana : Pirara, Tan. 1842, Schomburgk 601 (BD) ; savannas, 
shrub 3-4 m. tall, fl. white, Schomburgk 382 (K, BM, Leid. BD, DC, P) : 
Nigate, Nealu, Corantyne River, a few feet tall, Oct. 1879, Jcumaii 367 
(K) ; indefinite, Schomburgk 384 (BM). 

Dutch Guiana: Matappi, Corantyne River, June 18, 1916. B. W. 
2153 and 2174 (Utr) ; common on backlands of Paramaribo Gardens, 
2-2.5 m. tall, June 1910. Stockdalc 8823 (K) ; near Paramaribo, frequent, 
shrub 2.5-3 m. tall, fl. white, Nov. 1837, Splitgcrhcr 72 (Leid) ; near Para- 
maribo. Fockc 755 and 1132 (Utr) ; near Paramari!)o, shrub 2 ni. tall. Kuy- 



1935] JOHNSTON, STUDIES IN BORAGINACEAE 35 

per 20 (Utr); near Paramaribo, 1910, native collector no. 117 (Utr) ; 
near Paramaribo, Kegel 109 (P) ; edge of forest near Paramaribo, fl. vel- 
low, 1844, Kapplered.Hohenacker 1570 (P) ; Plant. Liberte, edge of forest, 
fl. white, shrub 2.5 m. tall, 1933, Lanjouw 223 (Utr) ; Plant. Rust en Werk, 
Aug. 5, 1913, shrub 4 m. tall, Soeprato 60 (Utr) ; Slootwijk, Commewyne 
River, roadside shrub 1 m. tall, July 1913, Soeprato 29G (Utr) ; Bloemendal 
Boite, June 4, 1913, 2 m. tall, Soeprato 25 A (Utr) ; Leonsberg, Aug. 8, 1913, 
Soeprato 127 (Utr); near Plant. Jagtlust, Aug. 1901, Went 64 (Utr); 
Para District, 1919, Kuyper 20 (Utr) ; lower Commewyne River near 
Plant. Maasstroom, Focke 1358 (Utr) ; indefinite, Hostmann 292 (K, BD, 
P), Hostmann sine no. (Leid), Focke 1355 (G, Utr), and Leschenaidt 
(P) ; indefinite, 1841, Berthoud-Coulon 551 (BM). 

Brazil: Rio Negro, Jan. 11, 1887, Moura 568 (BD) ; between mouth of 
Rio Negro and the Capoenas, shrub 2.5 m. tall, July 1851, Spruce 1695 
(K, BM); Igarape Burete, Pracua, Rio Surumu, Feb. 1909, Ule 7962 
(BD). 

Venezuela: lower Orinoco, 1896, Rusby & Squires 309 (G). 

This is the most widely distributed species of Cordia in America and, 
in many regions, one of the commonest. Its nomenclatorial history is 
involved. Its list of synonyms is a long one. Above I have given merely 
the oldest names, those published down to and including the Systema 
by Roemer & Schultes, vol. 4, in 1819. Since 1845 the plant has been 
called either C. ulmijolia or C. corymbosa. A study of these names 
proves them to be illegitimate on several grounds. 

The name C. ulmijolia was sponsored by DeCandolle, Prodr. 9: 494 
( 1845) . Those who followed him cited the name "Cordia ulmijolia Juss. 
in Bum. Cour., Bot. Cult. 2: 148 (1802)." A study of the work cited, 
however, shows that the name was actually published under Varronia 
and that no description was provided. It is a mere garden name! 
Varronia ulmijolia Juss. was accepted by no one until DeCandolle took 
it up and transferred it to Cordia. Previously, however, Sprengel, Syst. 
1: 653 (1825), had applied the binomial Cordia ulmijolia to another 
concept. Cordia ulmijolia (Juss.) DC. is hence a late homonym, besides 
being based upon a mere garden name. 

The name C. corymbosa seems to have been introduced by Urban, 
Symb. Ant. 4: 519 (1910). It has been universally accepted in recent 
years. The basic synonymy given by Urban and repeated by subsequent 
authors is as follows, — Cordia corymbosa (L.) Don, Gen. Syst. 4: 383 
(IS3&) and Lantana corymbosa L.Sp.Fl. 628 (1753). The name-bring- 
ing synonym is based upon several pre-Linnaean references all of which 
refer to our concept. Cordia corymbosa Don, however, is not based 
upon Lantana corymbosa but upon Varronia corymbosa Desvaux, Jour, 
de Bot. 1: 275 (1809). This latter name is expressly a renaming of C. 
monospermy Jacq. Both Desvaux I. c. 277 and Don, 1. c. 385, cite Lan- 



36 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

tana corymbosa L. among the synonyms of Varronia (or Cordia) 
lineatal ! The name Cordia corymbosa Don, can not be used for our 
plant in any case. It is antedated by Cordia corymbosa Willd. ex 
R. & S., Syst. 4: 801 (1819). Hence, since the name Cordia corymbosa 
Don is not based upon Lantana corymbosa L., and since it is invalid 
through being a later homonym, it must be rejected. 

Although it is the oldest name applied to our plant, Lantana corym- 
bosa L. can not be transferred to Cordia because of two earlier published 
homonyms. The next oldest name used for our plant is Varronia poly- 
cepkala Lam., which is based upon a phrase-name and illustration given 
by Plukenet. Transferred to Cordia Lamarck's name becomes the cor- 
rect one for the present concept. 

Various names cited by me in my formal list of synonyms have, at 
one time or another, been involved with binomials which I would refer 
to other species. The most important of these are Varronia lineata L. 
and V. humilis Jacq. Since these two names seem primarily based upon 
the notes and description given by Browne, Nat. Hist. Jamaica 172, tab. 
13, fig. 1 (1759), 1 identify them with the plant current as C. globosa 
(Jacq.) HBK. Browne's plant has the short pedunculate, terminal, 
globose flower-heads and the subulate calyx-lobes of that common West 
Indian species. 

The Guianan plants represent the typical West Indian form of Cordia 
polycephala, which is characterized by having the lower surfaces of the 
leaves densely and finely tomentulose. In South America, the typical 
form is found in our area, and in the country south along the Andes. 
One collection from Dutch Guiana, according to the label, is called, 
"Man blala oema" {native collector 117). 

21. Cordia macrostachya (Jacq.) Roemer & Schultes, Syst. 4: 461 
(1819). Varronia macrostachya Jacquin, Enum. PI. Insul. Carib. 14 
(1760) and (C. macrostachia) Select. Stirp. Amer. 41 (1763); Desvaux, 
Jour, de Bot. 1: 272 (1809). Varronia guianensis Desvaux, Jour, de 
Bot. 1: 270 (1809). Cordia gujanensis (Desv.) Roemer & Schultes, 
Syst. 4: 460 (1819); (C. guianensis) Meyer, Nov. Acta Acad. Caes.- 
Leop. Car. 12-': 778 (1825). Montjolya guianensis (Desv.) von 
Friesen, Bull. Soc. Bot. Geneve ser. 2, 24: 181 (1933). Cordia inter- 
rupta DeCandolle, Prodr. 9: 491 (1845). Cordia oxyphylla DeCan- 
dolle, Prodr. 9: 492 (1845). Lithocardium oxypkyllum Kuntze, Rev. 
Gen. 2:977 (1891). Cordia graveolens Humboldt, Bonpland & 
Kunth, Nov. Gen. et Sp. 3: 74 (1818); Miquel, Stirp. Surinam. 141 
(1850) ; Pulle, Enum. PI. Surinam 398 (1906). Varonia martinicensis 
sensu Aublet, Hist. PI. Guian. Fr. 1: 232 (1775); of Poiret, Encyc. 



1935] JOHNSTON, STUDIES IN BORAGINACEAE 37 

4: 264 (1797), as to Leblond 370. ? Cordia cylindrostachya sensu 
Schomburgk, Fauna u. Fl. Brit. Guian. 960 (1848). ? C. salicina sensu 
Garcke, Linnaea 22: 68 (1849). Varronia cylindrostachya sensu 
Graham, Ann. Carnegie Mus. 22: 240 (1934). 

Shrub 1 2.5 m. tall, younger parts bearing minute resinous granules; 
stems more or less strigose or with stiff incurved hairs; leaves lanceolate 
to oblong-ovate, elongate, 5-10 (-20) cm. long, 1-3 (-10) cm. broad, 
base obtuse or acute, contracted into a petiole 5-20 (-30) mm., long, 
apex obtuse to acute, margin denticulate to evidently dentate or sinuate- 
dentate, upper surface smooth to scabrous and characteristically gla- 
brous, bearing small limy tuberculations or murications which are low 
or somewhat prominent (rarely each bears a very short erect hair), lower 
surface pale, with soft curved hairs on the veins and veinlets; peduncles 
terminal, distinct from the petioles, 2-10 cm. long, slender; spikes 5-10 
cm. long, becoming rather loosely flowered at maturity; calyx granu- 
late and usually somewhat strigose, ca. 3 mm. long at anthesis, the tips 
of the triangular lobes not free in the bud; corolla white, ca. 5 mm. long; 
fruit red, invested to beyond the middle by the cupulate calyx; stone 
broadly ovoid, 4-5 mm. long. 

Northern South America and northward into Central America and 
the West Indies. 

British Guiana : Frechal, dry savanna, shrub, fl. white, fruit red, Sept. 
6, 1927, Tate 37 (NY) ; Pomeroon River, 25 dm. tall, 1922-23, La Cruz 
3041-3145 (G) ; Demerara, coastal region, 12 dm. tall, 1881, Jcnman 
1501 (K, P) ; vicinity of Demerara, 1824, Parker (DC, type of C. inter- 
rupta) ; Georgetown, wild land in the Botanic Garden, fl. white, Oct. 1919. 
Hitchcock 16535 (G, NY) ; coast lands, June 1889, Jenman 5206 (NY). 

Dutch Guiana: Corantyne River, 1911, Hulk 99 (Utr); near Para- 
maribo, 1910, native collector 41 (Utr) ; near Paramaribo, 2-3 m. tall, 
Essed 120 (Utr) ; near Paramaribo, 1904, Essed 120 (Utr) ; Paramaribo, 
Aug. 1901, Went 305 (Utr) ; Paramaribo, June 1903, Versteeg 464 (Utr) ; 
Paramaribo, Jan. 1901, Went 570 (Utr) ; Paramaribo, shrub 1-2 m. tall, 
Kuyper 34 (Utr) ; near Paramaribo, shrub 2-2.5 m. tall, fl. white, fruit 
red, Nov. 1837, Splitgerber 44 (Utr) ; Paramaribo near Agricultural Ex- 
periment Station, marshy land, fl. white, shrub ca. 1 m. tall, 1933, Lanjouw 
65 (Utr) ; near Paramaribo on road to Plant. Leonsberg, Aug. 1920, Pidle 
H48 (Utr) ; Leonsberg, 1913, Soeprato 135 (Utr) ; between Kwatta and 
Paramaribo, Feb. 28, 1900, Tulleken 67 (Leid) ; Div. Q, forest of Agric. 
Experim. Station, Paramaribo, 1916, Samuels 60 (G, K, Leid, BD, P) ; 
La Liberte, 1 m. tall, 1913, Soeprato 235 (Utr) ; Post Sommelsdijk, lower 
Commewyne River, July 1913, Soeprato 37 (Utr) ; Plant. Domburg near 
Surinam River, Aug. 23, 1900, TuUeken 265 (Leid) ; Para District, in 
woods, shrub, fl. white, April 1838, Splitgerber 1160 (Leid) ; Matappica, 
Dec, Fockc 279 (Utr) ; Lawa River, Oct. 1903, Versteeg 298 (Utr) ; upper 



38 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

Commewyne River, in forest, Focke 213 (Utr) ; indefinite, Focke 282 and 
446 (Utr), Tulleken 17 (Leid), Hostmann sine no. (Leid) and Hostmann 
323 (K, BM, BD, Del, P) ; indefinite, 1823-24, Leschenault (Leid, P). 

French Guiana: Acarouani, cultivated, "van inodora," 1856, Sagot 
(P); Mana, shrub, leaves fragrant, 1857, Sagot 444 (K, P) ; Mana, 
shrubby, leaves fragrant, fl. white, Sagot 444 (P) ; lies du Salut, shrub 
5-15 dm. tall, fragrant, fruit red, March 1856, Sagot (P) ; lies du Salut, 
shrub with fragrant leaves, 1854, Sagot 444 (K, BM, P) ; Cayenne, von 
Rohr (BM) ; Cayenne, 1853, Rothery 202 (K, BM) and sine no. (BM) ; 
near Cayenne in savannas, shrub, fl. white, fruit rose-colored, June 1921, 
Broadway 573 (G, NY, K) ; Cayenne, aromatic shrub, fl. white, fruit red, 
April 1897, Soubiron (P) ; Cayenne, very common about town, Richard 
(Paris, TYPE of V. guianensis) ; near Cayenne, shrub near sea, fl. white, 
fruit red, April 1921, Broadway 49 (G) ; indefinite, 1820, Perrottet 212 
(DC, TVi'E of C. interrupta); indefinite, 1792, Leblond 370 (Del, 
Lamarck). 

The type of Varronia macrostachya Jacq. comes from Cartagena. It 
is briefly described, and then largely through comparison with V. curas- 
savica Jacq., but with little doubt is evidently conspecific with our com- 
mon Guianan shrub. Our plants are remarkably similar to some I have 
seen from Cartagena. Among the spicate varronias of the coastal 
area of northern South America C. macrostachya is characterized by its 
glabrous upper leaf-surfaces, usually large leaves, and elongate slender 
terminal spikes. In dry localities the leaves tend to become smaller and 
the spikes short. In these phases the plant is distinguished from C. 
curassavica Jacq. only by the absence of hairs on the upper side of the 

Desvaux's Varronia guianensis evidently applies to this plant. The 
vernacular name, "Montjoly," and Aublet's discussion which are men- 
tioned by Desvaux, both apply to our plant. There is a sheet from the 
Desvaux collections at Paris which is determined as V. guianensis by 
Desvaux. The specimen is labeled merely as from South America, 
except that it is given as a shrub, no other data concerning it is given 
on the accompanying label. The specimen, however, is almost cer- 
tainly a duplicate of one, also at Paris, collected by Richard and labeled 
as "frequentissima in suburbanis, Cayenne." This information as to 
locality is that given by Desvaux as the source of his species. I con- 
sider the specimens as types of the species. 

The type of C. interrupta was collected by Perrottet. It is given 
merely as from French Guiana and it consists of a leaf of C. tomentosa 
and a branch of C. macrostachya. A comparison of this type-material 
in the Prodromus Herbarium with other Perrottet material in the De- 
lessert collections, at Geneva, shows such a close agreement in details of 



1935] JOHNSTON, STUDIES IN BORAGINACEAE 39 

discoloration, etc., etc., that we may well believe them parts of one 
collection. The material in the Delessert Herbarium is labeled as from 
"Mana, 1820, Perrottet." 

Humboldt and Bonpland collected material near Angostura on the 
Orinoco which seems to be good C. macrostachya. The material, how- 
ever, was described as a new species, Cordia graveolens HBK. Further 
up the Orinoco, perhaps in very shaded humid locations, there have been 
collected plants evidently related to C. graveolens, though differing in 
having the upper leaf-surfaces quite smooth and bearing only minute 
very scattered microscopic limy disks and resinous granulations. This 
form was described as Cordia polystachys HBK. and C. canescens 
Willd., from the Mapure. Spruce (no. 3012) has collected a quite 
similar plant near the Brazil- Venezuela border. The type of another 
species, C. spicata Willd., given as from Angostura, seems to be an essen- 
tially similar form. 

The material given as collected by Richard in Cayenne by Poiret, 
Encyc. 4:264 (1797), under the name Varronia curassavica, differs 
from all Guianan plants and is, I believe, West Indian. Desvaux, Jour, 
de Bot. 1:271 (1808), and later Poiret, Encyc. Suppl. 3:728^29 
(1814), associate this specimen with V. angustijolia West, of the island 
of St. Croix. This is probably correct. 

In the Lamarck herbarium there are only two sheets labelled V. mar- 
tinkensis. This material belongs to Leblond no. 370 and is the basis of 
the description of V. martinicensis by Poiret, Encyc. 4: 264 (1797). 
Desvaux referred the plant to V. curassavica, but I consider it quite 
representative of C, macrostachya. 

It should be noted that plants of eastern Brazil (from Ceara south- 
ward to Rio Grande do Sul), which I have treated as Cordia verhenacea 
DC, cf. Johnston, Contr. Gray Herb. 92: 25 (1930), is verly closely 
related to C. macrostachya and perhaps should be accepted as a form 
of it. 

The present species seems to be well kno 
the name "Montjoly." In several collectic 
plant is given as called "Blaka oema." 

22. Cordia Schomburgkii DeCandolle, Prodr. 9:490 (1845); 
Schomburgk, Fauna u. Fl. Brit. Guiana 960 (1848); Garcke, Linnaea 
22: 68 (1849); Pulle, Enum. PI. Surinam 397 (1906). Lithocardium 
Schomburgkii (DC.) Kuntze, Rev. Gen. 2: 977 (1891). Cordia lucida 
Splitgerber ex Pulle, Enum. PI. Surinam 397 (1906), nomen sub C. 
Aubletii. Cordia tobagensis Urban in Fedde, Rep. Spec. Nov. 16: 39 
(1919). Cordia tobagensis var. Broadwayi Urban in Fedde, Rep. Spec. 



40 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

Nov. 16: 40 (1919). (?) Cordia Aubletii sensu Schomburgk, Fauna u. 
Fl. Brit. Guiana 960 (1848). Cordia polystachya sensu Scliomburgk, 
Fauna u. Fl. Brit. Guiana 1151 (1848). Cordia patens var. polycephala 
sensu Miquel, Stirp. Surinam. 140 ( 1850). Cordia Aubletii sensu Pulle, 
Enum. PI. Surinam 397 (1906). Varronia guianensis sensu Graham, 
Ann. Carnegie Mus. 22: 239 (1934). 

Shrub 1-4 m. tall; stems with a thin indument of fine curved pu- 
bescence and intermixed short coarse curved bristles; leaves ovate to 
elliptic or lance-elliptic, 5-10 cm. long, 2-7 cm. broad, base obtuse or 
nearly rounded, apex acute, margin entire to sharply dentate, upper sur- 
face drying brown, usually somewhat lustrous, distinctly and simply 
strigose, lower surface pale, covered with a fine minute curved puberu- 
lence, petiole slender 6-9 mm. long, decurrent 5-8 mm. on the subtended 
branchlet or peduncle; peduncles axillary, slender, ascending, up to 
9 cm. long; spikes elongate and loosely flowered, 5-15 cm. long; calyx 
nearly glabrous, bearing numerous resinous granules, with a few bristles 
about the apex, the tips of the deltoid lobes not free in the bud, calyx 
at anthesis or when sterile vase-shaped or funnel-form and 3-4 mm. 
long, at maturity expanded by the enlarging fruit; corolla white, ca. 4 
mm. long; fruit red, tightly ensheathed by the calyx; stone ovoid 
4-5 mm. long. 

Known only from British, Dutch and French Guiana and from Trini- 
dad and Tobago. 

British Guiana : banks of the Barama, Oct. 1843, Schomburgk 1510 
(BD); Tumatumari, Potaro River, along trail in forest, shrub 2.5-3 m. 
tall, fl. white, Jan. 1920, Hitchcock 17382 (G, NY) ; Penal Settlement, 
1911, Hitchc\ock 17150 (K) ; upper Mazaruni River, long. 60° 10' W., fl. 
yellow, 8-12 dm. tall, Sept. 1922, La Cms 2343 and 2260 (G, NY) ; Maza- 
runi, Appitn 293 (K) ; Kamakusa, upper Mazaruni, long. 59° 50' W., Nov. 
1922, La Cruz 2882 (G, NY) ; Kyk-over-all, near Kartabo, bushy shrub 
3 m. tall, July 1924, Graham 215 (NY) ; Kartabo, large shrub in clearing. 
Aug. 11, 1920, Bailey 170 (G) ; Kalacoon, shrub in clearing, fruit red, 
Aug. 23, 1920, Bailey 170 (G) ; Kalacoon, shrub 15 dm. tall, along creek, 
June 1924, Graham 133 (NY); banks of Rupununi, May 1843, Schom- 
burgk 1304 (BD, as C. polystadiva, det. Kl.) ; Demerara, Parker (K) ; 
Kamuni Creek, Demerara River, March 1889, Jenman 4919 (K) ; Malali, 
Demerara River, lat. 5° 35' N., 1922, La Cruz 2678 (G. NY) ; Christian- 
burg, Demerara River, 1910, Anderson 553 (K) ; Vryheid, Demerara 
River, fl. white. Feb. 15, 1924, Linder 61 (G, NY) ; between Demerara and 
Berbice rivers, ca. lat. 5° 50' N., 18 dm. tall, fl. white. La Cruz 1594 (G, 
NY); Berbice, 1837, Schomburgk 406 (Del); indefinite, 1838, Schom- 
burgk 406 (DC, TYPE of C. Schomburqkii: isotypes, K, BM, Leid, BD, 
P) : Canje Creek. Aug. 1908. Bartlctt (NY). 

Dutch Guiana: upper Nickerie River, Feb. 24, 1915, B. W. 1035 



1935] JOHNSTON, STUDIES IN BORAGINACEAE 41 

(Utr) ; Saramacca River, Dec. 1902 and Jan. 1903, Pulle 125 and 417 
(Utr) ; Heidoti, Saramacca River, 1920, B. W. 4621 (Utr) ; road near 
Paramaribo, 1842, Focke 816 and 760 (Utr); near Paramaribo, shrub 
2.5-3.5 m. tall, fl. white, Dec. 1837, Splitgcrber 206 (Leiden, type of C. 
Uicida) ; Plant. Liberte, lower Surinam, edge of forest, shrub 2-3 m. tall, 
fl. white, 1933, Lanjouw 224 (Utr) ; Para District, forest, fl. white, April 
1838, Splitgerher 1161 (Leid) ; Para District, June 1904, Versteeg 507 
(Utr) ; Plant. Guineesche Vriendschap, 1915, Soeprato 313 (Utr) ; Caro- 
lina Creek, Para River, May 1921, B. IV. 5123 (Utr) ; near Guyana placer 
mines, Oct. 1909, Boldingh 3918H (Utr) ; Kadjoe, Surinam River, May 
1910, natwe collector (Utr) ; Koemba Rapids, upper Surinam River, July 
1908, Tresling 232 (Utr, BD) ; indefinite, Hoshnann (Leid), Hostmann 
295 (Utr, P), Hostmann 877 (G, K, BM, Utr, BD, Munich, Del, P), 
Kap pier 877 (P) and Focke (K). 

French Guiana: St. Laurent du Maroni, shrub 2-3 m. tall, fl. white, 
Jan. 15, 1914, Benoist 604 (P) ; St. Jean, shrub 2 m. tall, fl. white. May 
16, 1914, Benoist 1223 (P) ; Mana, fl. white, 1858, Sagot 1169 (K, BM, 
P) ; Godebert, Jan. 1920, Wachenheim 79 (K, BM, P), 101 (G, K, BM, 
P) andJ52 (P). 

The species is most nearly related to C. Poeppigii DC, of eastern 
Peru and to C. jerruginea (Lam.) R. & S. of the northern Andes and 
Central America. It is readily recognized by its peculiar calyx. This, 
when not distorted by the enlarging fruit, is funnel-form or vase-shaped 
and is abruptly expanded from a short tube (ca. 1-1.5 mm. thick and 
1-2 mm. long) or even from the narrow base. The calyx-tube is cov- 
ered with numerous minute resinous granules. The ascending, more 
or less deltoid, lobes are somewhat strigose outside especially towards 
their apices. 

The following vernacular names have been found on the labels of the 
specimens indicated. Black sage — Anderson 553; Waijanaka erepa- 
\oe—B. W. 4621; Baka Otxtvai— Tresling 232; Blakka hoema— 5. W . 
5123 and Makoeja pipa (Kar.), Kaboejakoro diamaroe (Arow.) and 
Blakka wintje (Nig. Eng.)— 5. W. 1035. 

23. Oordia tomentosa Lamarck ex Roemer & Schultes, Syst. 4: 459 
(1819). Varronia tomentosa Lamarck, Tab. Encyc. 1:419 (1791); 
Poiret, Encyc. 4: 264 (1797); Desvaux, Jour, de Bot. 1: 268 (1808). 
Lithocardium tomentosum (Lam.) Kuntze, Rev. Gen. 2:977 (1891). 
Montjolya tomentosa (Lam.) von Friesen, Bull. Soc. Bot. Geneve ser. 
2, 24: 183 (1933). Cordia Aubletii DeCandoUe, Prodr. 9: 490 (1845). 
Lithocardium Aubletii (DC.) Kuntze, Rev. Gen. 2: 976 (1891). 

Shrub, 1-4 m. tall ; stems clothed with a mixture of curved ascending 
coarse and slender hairs; leaves lanceolate to ovate-lanceolate or ovate, 
7^15 cm. long, 2.5-7 cm. broad, base obtuse or somewhat rounded, apex 
acute or somewhat acuminate, margin irregularly dentate, upper sur- 



42 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

face dull, scabrous, with numerous stiff ascending bristles which arise 
from more or less bulbous bases, lower surfaces pallid, more or less 
tomentose with abundant fine short interlaced hairs; petioles 5-10 cm. 
long, decurrent 2-6 mm. on the subtended branchlet or peduncle; pe- 
duncles axillary, ascending, 2-10 cm. long; spike broadly clavate, very 
dense, 1.5-4 cm. long, at anthesis ca. 7 mm. thick, increasing to about 
twice that thickness in fruit; calyx densely hairy or tomentose with 
intermixed resinous granules, ca. 5 mm. long at anthesis, much accrescent 
in maturity, the tips of the lobes evidently free in the bud, calyx-lobes 
narrowly triangular, very elongate, acuminate; corolla white; fruit 
tightly invested by the calyx at maturity; stone ca. 4 mm. long. 

Known only from French and Dutch Guiana. 

Dutch Guiana: Voltz Mts., open formation, small shrub 2 m. tali, fl. 
white, Aug. 22, 1920, Pulle 252 (Utr). 

French Guiana: Cayenne, open ground near Baduel, July 10, 1921, 
Broadway 720 (G) ; Cayenne, shrub 4.5 m. tall, fl. white, Feb. 1859, Sagot 
(P) ; Cayenne, S. Marca savanna near Mt. Baduel, 1867, Jelski (BD) ; 
Cayenne, 1859, Sagot 1315 (K, BM, P) ; Cayenne, Martin (K), Leprieur 
(K, Del, P) and Aublct (BM) ; indefinite, Martin ex herb. Rudge (BM) ; 
indefinite. Poiteau (K, BD, Del) ; indefinite, 1820, Perrottet 211 (DC). 

The present species is most closely related to C. multispicata Cham, 
of eastern Brazil, from which it differs in having the calyx coarsely stri- 
gose or tomentose all over, rather than nearly or quite glabrous. The 
type of Varronia tomentosa in the Lamarck Herbarium is labeled as 
coming from Jussieu and is devoid of any geographic data. It consists 
of a leaf and a fragment of inflorescence. These, however, evidently 
represent the much collected plant of the vicinity of Cayenne which is 
treated here. 

There must remain some question as to the proper disposition of the 
name Cordia Aubletii DC. The plant actually described by DeCandolle 
is Perrottet (no. 211), which represents C. tomentosa. The reference 
by DeCandolle to "Varronia Martinicensis Aubl. guian. 232 non Jacq.", 
which might stand as the basis for the name chosen by him, is quite 
ambiguous. Aublet quoted a name and a descriptive phrase from 
Jacquin which apply to a West Indian species. The several lines of 
discussion by Aublet, concerning the fragrance of the herbage, the color 
of the fruit, the vernacular name and the local uses of the plant, in fact 
all the original data, all apply to C. mcacrostachya Jacq. Consequently 
if the name C. Aubletii is taken as founded upon the reference to Aublet's 
work, the species justly should become a synonym of C. macrostachya. 
I have preferred, however, to associate the name C. Aubletii with De- 
Candolle's specimen from Perrottet and the description of that specimen 



19351 JOHNSTON, STUDIES IN BORAGINACEAE 43 

published in the Prodromus. It may be further noted that the speci- 
men of Hostmann 877, which the younger DeCandoUe, in a foot-note, 
cited as representing C. Aubletii, is in fact representative of C. 
Schomburgkii, 

In the British Museum there is a collection of C. tomcntosa made by 
Aublet. Among his manuscripts I have seen a good description of the 
species mentioned. It is possible that Aublet identified the plant as 
Varronia globosa and that the report of V. globosa in his book, 1. c. 
1: 232 (1775), may be based upon his collection of C. 



24. Cordia multispicata Chamisso, Linnaea 4: 490 (1829); Frese- 
nius in Martius, Fl. Bras. 8^: 17, tab. 6 (1857) ; Johnston, Contr. Gray 
Herb. 92: 29 (1930). Lithocardium multispkatum (Cham.) Kuntze, 
Rev. Gen. 2: 977 (1891). Cordia bahiensis DeCandolle, Prodr. 9: 489 
(1845). Varronia spicata Salzmann ex DeCandolle, Prodr. 9:489 
(1845), as synonym. 

Shrub becoming subscandent, 1-3 m. tall; stems clothed with a mix- 
ture of curved ascending coarse and slender hairs; leaves ovate to ovate- 
lanceolate, 4-10 cm. long, 2.5-6 cm. broad, base obtuse to rounded, apex 
acute to somewhat acuminate, margin crenate or dentate, upper surface 
not lustrous, scabrous, bearing numerous coarse short ascending hairs 
from bulbous bases, lower surface pale, bearing slender soft curved hairs 
on the nervation; petioles slender, 4-10 mm. long, decurrent 2-5 mm. 
on the subtended peduncle or branchlet; peduncle axillary, 1-6 cm. 
long, ascending; spike broadly clavate to nearly subcylindrical, 1-7 cm. 
long, short and dense to rather loose and elongate, 7-10 mm. thick; 
calyx hairy on the lobes, the tube practically glabrous, bearing numerous 
resinous granules, tip of lobes short but free in the bud, lobes broadly 
triangular and short acuminate; calyx ca. 3 mm. long at anthesis, 
accrescent; corolla white, 4-5 mm. long; fruit ensheathed by the calyx 
at maturity; stone ca. 4 mm. long. 

Brazilian, from the mouth of the Amazon southward to Rio Janeiro. 

Brazil: beach at Prainha, Nov. 26, 1873, Traill 561 (K). 

This species which has been much collected about the mouth of the 
Amazon, particularly in the vicinity of Para, is clearly related to C. to- 
mentosa of French and Dutch Guiana. Collecting on the Brazilian 
coast north of the Amazon will no doubt reveal the presence of the species 
in that region and possibly may produce forms transitional to C. tomen- 
tosa as well. Cordia multispicata seems to differ from C. tomentosa 
chiefly in its calyx which tends to be entirely glabrous and evidently 
resinous-granulate below the shorter nearly deltoid lobes. In C. tomen- 



44 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

tosa the calyx-lobes are distinctly more elongate, being narrowly tri- 
angular, and have more elongate tips. The calyx-tube in the Guianan 
plant is hairy all over. In fact, except for the free lobe-tips of the calyx, 
C. multispkata is more suggestive of C. Schomburgkii than of C. 
tomentosa. 

Doubtful and excluded Species 

Cordia flavescens Aublet, Hist. PI. Guian. Fr. 1:226, tab. 89 
(1775); Poiret, Encyc. 7: 43 (1806); Schomburgk, Fauna u. Fl. Brit. 
Guiana 960 (1848); Johnston, Contr. Gray Herb. 92:63 (1930). 
Lithocardium flavescens (Aubl.) Kuntze, Rev. Gen. 2:977 (1891). 
Cordia sarmentosa Lamarck, Tab. Encyc. 1 : 422 (1791). Cordia erhi- 
tioides Lamarck ex Dietrich, Synop. 1:612 (1839), in synonymy. 
Firensia Scopoli, Intr. 157 (1777). 

According to Aublet, 1. c. 227, the original material of this species was 
collected on the He de Cayenne and on the adjacent mainland of French 
Guiana near Tonnegrande. The species is based upon mixed material 
consisting of (1) flowers of some species of Cordia, and (2) fruiting 
branches of Ocotea commutata Nees, of the Lauraceae. Since the spe- 
cies is based upon two entirely discordant elements it must be rejected 
under article 51 of the International Rules of Nomenclature. At the 
British Museum and at the museum in Stockholm there are specimens of 
this species collected by Aublet. These specimens represent only the 
Ocotea. 

Cordia lutea Lamarck, Tab. Encyc. 1: 421 (1791). 

A specimen of this species in the Delessert Herbarium is labeled as 
collected in Cayenne by Perrottet. The species is known only from 
the semi-arid regions of western Peru and Ecuador and is not even to be 
expected from the wet Guianan coast. The data on the specimen at 
Geneva are obviously incorrect. I have had previous occasions to ques- 
tion the accuracy of the geographic data associated with the Perrottet 
collections in the Delessert Herbarium. 

Cordia scandens Poiret, Diet. Sci. Nat. 10: 410 (1818). 

This species is given as collected by Martin in Cayenne. I have seen 
the fragmentary type in Poiret's herbarium and ample specimens of 
evidently the same collection in the General Herbarium, also at Paris. 
A search in the herbarium proved that this authentic material is refer- 
rable to Dichapetalum vestitum Baill., var. scandens [Benth.] Baillon 
in Martius, Fl. Bras. 12' : 372 (1886). The correct name for that 
plant, consequently, is Dichapetalum scandens (Poir.), comb. nov. 

Cordia tetraphylla Aublet, Hist. PI. Guian. Fr. 1:224, tab. 88 
(1775); Poiret. Encyc. 7: 42 (1806); Schomburgk, Fauna u. Fl. Brit. 



1935] JOHNSTON, STUDIES IN BORAGINACEAE 45 

Guian. 1151 (1848); Fresenius in Martius, Fl. Bras. 8': 13 (1857); 
Pulle, Enum. PI. Surinam 397 (1906); Johnston, Contr. Gray Herb. 
92: 64 (1930). Lithocardium tetraphyllum (Aubl.) Kuntze, Rev. Gen. 
2: 977 (1891). Firensia lutea Rafinesque, Sylva Tellur. 40 (1838). 

Aublet, 1. c. 226, states that this shrub is very common on the sand 
near Kourou and westward along the French Guiana coast to Sinnamary. 
The description given by Aublet is a mixture, based partly upon the flow- 
ers of some species of Cordia and partly upon a leafy fruiting branch of 
Buchenavia capitata (Vahl) Eichl. of the Combretaceae. The name 
based upon entirely discordant elements must be discarded. 

Cordia Myxa sensu Schomburgk, Fauna u. Fl. Brit. Guiana 830 
(1848). 

Cordia curassavica sensu Schomburgk, Fauna u. Fl. Brit. Guiana 
830 (1848). 

Cordia martinicensis sensu Schomburgk, Fauna u. Fl. Brit. Guiana 
830 (1848). 

The three names above cited are listed by Schomburgk in his cata- 
logue of the plants of British Guiana. The first two are given as culti- 
vated. The last is given as growing wild on plantations and in ditches 
in the coastal region. I suspect that all three names represent mis- 
determinations. 

2. Lepidocordia Ducke, Archiv. Jard. Bot. Rio Janeiro 4: 170 
(1925). 

Large tree with broad leaves. Inflorescence a dichotomously branched 
corymbose panicle. Calyx 5-lobed, persistent. Corolla white, small, 
5-merous, with short tube and spreading lobes. Stamens 5, short-ex- 
serted. Ovary 4-celled. Stigmas 2, conic, sessile on the apex of the 
ovary. Fruit unlobed, a drupe, breaking up into 2 flattened bony 2- 
seeded nutlets. Endosperm present. Cotyledons flat. 

A monotype endemic to our area. Its immediate relationships are 

1. Lepidocordia punctata Ducke, Archiv. Jard. Bot. Rio Janeiro 
4: 171, tab. 22 (1925); Sandwith, Kew Bull. 1933: 335 (1933). 

Tree 15-30 m. tall, trunk irregularly and very deeply fluted; leaves 
oblong-lanceolate, broadest at or above the middle, 1-2 dm. long, 2-7 
cm. broad, with 6-9 pairs of veins, base obtuse to acute, apex acuminate, 
upper surface with inconspicuous short erect hairs on the veins and 
veinlets, minutely and abundantly white-pustulate, lower surface darker 
with ascending or appressed hairs on the principal veins, petiole 1-2 cm. 
long- inflorescence stiff, 5-10 cm. broad, flowers crowded at the ends of 



46 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

the branches, not scorpioid; calyx l-l mm. long at anthesis, in fruit 
twice as large and persistent, lobes lance-ovate, acuminate; corolla 
white, 2-2.5 mm. long, lobes ovate and about as long as the tube; fila- 
ments flattened, glabrous, ca. 1 mm. long, inserted in the middle of the 
corolla tube; anthers small; ovary glabrous; fruit erect, narrowly obo- 
void or ellipsoid, glabrous, lustrous, red, 5-7 mm. long, ca. 3 mm. thick. 

British Guiana: right bank of the Rewa River, ca. 14 miles SSE. of 
tall, trunk deeply 
low hill with red 
, D91, record 
no. 2082 (K). 

Brazil : Rio Branco de Obidos, northeast of Obidos in the forest called 
"Repartimento," State of Para, non-flooded forest, medium-sized tree 
with fluted trunk, fl. white, Dec. 15, 1913, Diickc, Jard. Bot. Rio Jan. no. 
17864 & Herb. Anias. Mus. Para no. 15152 (K, BD, isotypes) ; hills near 
the Rio Brariquinho, a tributary of the Rio Branco de Obidos, large tree 
with trunk excavated and sulcate, fl. white, fruit red, Jan. 27, 1918, Diicke, 
Jard. Bot. ,Rio Jan. no. 17863 & Herb. Amaz. Mus. Para no. 16958 (K, BD) ; 
forest on Rio Branco de Obidos, elevated place near mouth of Rio Bran- 
quinho, large tree with white flowers, Nov. 1, 1919, Duckc, Jard. Bot Rio 
Jan. HO. 11406 (BD). 

This remarkable tree is known only from the collections cited. Ducke 
found it on a minor tributary of the Amazon, just to the northeast of 
Obidos, State of Para, Brazil. He has made several collections at this 
type-locality. Members of the Forest Service of British Guiana recently 
discovered a second locality on the Rewa (or Illiwa) River, between the 
Rupununi River and the upper Essequibo between 3° and 4° N. lat. 
This new station is about 650 km. east-northeast of the type-station near 
the Amazon, and separated from it by the basin of the Rio Trombetas 
and the headwaters of the Essequibo River. There is every reason to 
believe that Lepidocordia will be found in the intervening region when 
it is reasonably well explored. 

3. Tournefortia Linnaeus, Gen. 68 (1754). 

Shrubs or woody vines with broad leaves. Inflorescence consisting of 
scorpioid racemes or spikes borne in dichotomous panicles. Calyx per- 
sistent, usually 5-lobed. Corolla white or yellowish, small, usually 5- 
merous, with cylindrical tube and spreading limb. Stamens usually 5, 
borne on the corolla-tube, included; filaments short. Ovary 4-celled, 
style terminal and solitary. Stigma sessile or borne on a distinct style, 
peltate or conic, fertile on the sides, apex usually bifid. Fruit a drupe, 
lobed or unlobed at maturity breaking up into 2-4 bony nutlets. Nut- 
lets 1-2-seeded, frequently with 1-2 empty cavities. Endosperm thin. 
Cotyledons flat. 



1935] JOHNSTON, STUDIES IN BORAGINACEAE ^ 

A genus of about 100 variable and ill-defined species; widespread i 
the Tropics but evidently centering in America. Type Species, T. hh 



Fruit deeply 4-lobed; embryos curved; corolla-lobes linear or 
long acuminate. § Cyphocycma. 
Corolla-tubes short, 1.5-2.3 mm. long, constricted at the 
throat, lobes linear, nearly as long as the tube ; fruit white ; 

plant very slender 1- T. volnbilis. 

Corolla-tube elongate, 3-8 mm. long, not constricted at throat ; 
lobes broadened below middle, half length of tube or less ; 
fruit yellowish. 
Herbage sptirsely and inconspicuously short-strigose or gla- 
brous, even when young 2. T. syringaefolia. 

Herbage evidently and usually abundantly hairy, especially 

when young 3. 7. paniculata. 

Fruit obscurely if at all lobed ; embryos straight; corolla-lobes 
broad and rounded. § Eutournefortia. 
Style well developed, 2-3 mm. long, evident even on the mature 
fruit; throat of corolla inflated; fruiting calyces fre- 
quently pedicellate ■ -4. T. Ulei. 

Style short, the stigma apparently sessile on the mature fruit ; 

throat of corolla constricted ; calyx sessile even in fruit. 

Stems with pale short curving appressed hairs or glabrous ; 

corolla-tube 4-5 mm. long 5. T. hicolor. 

Stems with spreading brown or tawny bristles ; corolla-tube 
5-8 mm. long. 
Leaves dull, with abundant long slender hairs ; stems with 

abundant bristles 2-4 mm. long 6. T. cuspidate. 

Leaves glossy, with only scattered hairs along the veins, 
practically glabrous ; stems with only scattered short 

bristles 1-2 mm. long 7. T. melanochaeta. 

1. Tournefortia volubilis Linnaeus, Sp. PI. 140 (1753). T. flori- 
bunda sensu Schomburgk, Fauna u. Fl. Brit. Guian. 1084 (1848). 

Slender vine, densely clothed with slender curved hairs; branchlets 
slender; leaves lanceolate to ovate-lanceolate, 2-10 cm. long, 1-5 cm. 
broad, base rounded or obtuse, apex acuminate, surface densely strigose 
or velvety-tomentose, pale or tawny; inflorescence very slender and 
loosely branched, the spikes becoming 2-10 cm. long; calyx 1-2 mm. 
long, weakly accrescent, lobes subulate, reaching to beyond middle of 
corolla-tube; corolla white, tube 1.5-2.3 mm. long, strigose, throat con- 
stricted, lobes linear, 1-2 mm. long, spreading; fruit white with black 
dots. 4-lobed, the lobes subglobose, breaking up into single-seeded nut- 
lets; style developed. 

Eastern British Guiana, Venezuela, Colombia and Ecuador, and 
northward in the West Indies and Central America. 



JOURNAL OF THE ARNOLD ARBORETUM 



I believe that the three above cited specimens are parts of a single 
collection and the basis upon which Schomburgk, 1. c. 1084, reported 
T. floribunda from the southern slopes of Roraima. Part of the south 
slope of Roraima belongs to Venezuela, and it is quite possible that 
Schomburgk's specimens may have come from within Venezuelan rather 
than Brazilian territory. 

Schomburgk, op. cit. pg. 830, reports T. volubilis as occurring about 
abandoned plantations near the coast of British Guiana, and Aublet, 
Hist. PI. Guian. Fr. 1 : 117 (1775), lists it as occurring in French Guiana. 
I have seen no specimens to substantiate either of these two records. 
I consider the accuracy of both records extremely questionable. 

2. Tournefortia syringaefolia Vahl, Symb. 3: 2i (1794). Messer- 
schmidia syringifolia (Vahl) Roemer & Schules, Syst. 4: 543 (1819); 
Don, Gen. Syst. 4: 370 (1838). T. peruviana Poiret, Encyc. Suppl. 
4; 425 (1816); Urban, Symb. Ant. 4: 524 (1910); Johnston, Contr. 
Gray Herb. 92: 78 (1930). T. swinamensis A. DeCandolle, Prodr. 
9: 526 (1845); Schomburgk, Fauna u. Fl. Brit. Guian. 961 and 1151 
(1848) ; Miquel, Stirp. Surinam. 138 (1850) ; Pulle, Enum. PI. Surinam. 
398 (1906). T. Hostmanni Klotzsch ex Schomburgk, Fauna u. Fl. 
Brit. Guian. 1151 (1848), nomen. T. maculata sensu Lamarck. Tab. 
Encyc. 1: 416 (1791) and Poiret, Encyc. 5: 357 (1804), as to plants 
in Herb. Lam. T. laurijolia sensu DeCandolle, Prodr. 9: 522, adnot. 
(1845). T. joetidissima sensu De Vriese, Nederl. Kruidkund. Arch. 
1: 347 (1848). 

Shrubby vine; branchlets inconspicuously short-pubescent; leaves 
ovate to lance-elliptic or broadly lanceolate, 4-10(-15) cm. long. 
2-5 (-8) cm. broad, base acute to rounded, apex acuminate, lower face 
paler than upper, both faces very sparsely and very inconspicuously 
short-strigose (even when immature), usually abundantly and very 
minutely tuberculate, petioles 7-15 mm. long; inflorescence slender, 
loosely branched, terminal, 5-15 cm. broad, spikes usually less than 
5 cm. long even in fruit; calyx 1-1.5 mm. long, weakly accrescent, lobes 
subulate to ovate; pedicels 0-1 mm. long at anthesis, in fruit becoming 
1-5 mm. long and usually much thickened; corolla greenish white, tube 
3-4 or even 8 mm. long, limb 3-4 or even 5 mm. broad, lobes 1-1.3 or 
even 2.5 mm. long, spreading, broad below the middle and above coarsely 
long-acuminate; fruit yellow or yellowish, usually spotted with black, 
conspicuously 4-lobed, the lobes subglobose, breaking up into 4 single- 
seeded nutlets; style developed. 



1935] JOHNSTON, STUDIES IN BORAGINACEAE 49 

From northern Brazil and Peru northward into the West Indies and 
Central America. 

British Guiana: Berbice, Jan. 1896, Jcnman 6925 (K). 

Dutch Guiana: Plant. Jagtlust, 1913, Soeprato 38E (Utr) ; Para Dis- 
trict, scandent shrub, corolla somewhat greenish white, April 1838, Split- 
gerber 1159 (Leid, P) ; Brownsberg Summit, liana with greenish flowers, 
1924, B. W. 6650 (Utr) ; banks of lower Commewyne River, Dec. 1842, 
flowers greenish, Focke 750 (Utr) ; indefinite, Hostmann 289 (BD, type 
of T. Hostmanni Kl. ; K, BM, P), Hostmann 951 (Boiss, type of T. siiri- 
namensis; G, K, BM, Utr, BD, Deles, P), Hostmann ed. Hohenacker 1721 
(P), Kappler951 (P), Kappler 1721 (Utr) and Focke 180 (Leid). 

French Guiana: Cayenne, von Rohr (Copenhagen, type of T. syrin- 
gaefolia, BM, isotype) ; Cayenne, Martin (K) ; Cayenne, Rudge (BM) ; 
near Cayenne, Feb. 12, 1845, Rothcry 207 (K), sine no. (BM) ; Montabo, 
Cayenne, 1866, Jelski (BD). 

I have seen the types of all the species listed above. Vahl's species 
evidently belongs here and is notable chiefly for having the leaf-blade 
ovate (in accord with its name) rather than lance-oblong as is usually 
common in this plant. The common form of the species in the Guianas 
and in northwestern South America is well exemplified by the types of 
T. Hostmanni and T. surinam^ensis . All the material from French 
Guiana, including the type of T. syringaejolia, has corolla-tubes a few 
millimeters longer than in other South American plants. Similar 
elongate corolla-tubes, however, are found in the West Indian plants 
that have been classified, along with all South American forms, as T. 
peruviana. Possibly the difference in corolla-length may merit some 
nomenclatorial recognition, particularly as both the Cayenne and West 
Indian forms with elongate corollas also tend to have more ample leaf- 
blades than the short-tubed plants. 

3. Tournefortia paniculata Cham. var. spigeliaeflora (A. DC), 
comb. nov. Tournefortia spigeliae flora A. DeCandolle, Prodr. 9: 525 
(1845); Schomburgk, Fauna u. Fl. Brit. Guian. 1151 (1848); John- 
ston, Contr. Gray Herb. 92: 81 (1930). 

Similar to T. syringaejolia Vahl, differing only in the more abundant 
and usually more slender and tawny hairs on the herbage, particularly 
on the immature leaves. 

About the margins of the Amazon Basin in southwestern British 
Guiana, Colombia and Peru ; also in Costa Rica. 

British Guiana : Rupununi near Pirara, Feb. 1842, Schomburgk 669 
(BD) ; Pirara, etc., Schomburgk 427 (BM, Deles, P) ; near Pirara, 1838, 
Schomburgk 749 (K, BM, Leid, BD, Deles; DC, type); indefinite, 
Schomburgk 427 /669b (K). 

The elusiveness and scarcity of characters separating the many rec- 



50 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

ognizable species of Tourmfortia are well exemplified by the case of 
T. paniculata. This plant, centering in Brazil and having a distribution 
generally to the south and east of its close relative, T. syringaejolia, is 
separated from its relative only by quantity of pubescence. This differ- 
ence is usually very real and tangible. When not, it at least gives to 
the two forms a perceptible though almost intangible difference in 
aspect, which coupled with their distinctly natural and credible wide 
geographic range, leads one inevitably to the conviction that two large 
genetic entities are concerned. These two species would be united by a 
stern judge of species-behavior. In his treatment of Tournejortia, how- 
ever, he would be forced to create specific aggregations so large as to be 
all inclusive and indefinite, or to assemble under his aggregates such an 
array of subspecific categories as to be cumbersome and impractical. 
Difficulties in defining and describing species are encountered repeatedly 
in Tournejortia and force the student of the genus to abandon rigid and 
preconceived notions concerning species values. Testing the subjective 
matters of plant aspect by the facts of geographic distribution, he 
must gropingly work out natural concepts of the incipient and unfor- 
tunately not yet sharply definable species. The test of the resulting 
classification of the species of Tournejortia is not the imposing number 
and decisiveness of the key-characters, but the objectivity of the con- 
cept judged by one studying masses of material of it and of the group to 
which it belongs. A study of material of T. paniculata and T. syringae- 
jolia does justify these two concepts and, as in many other similar cases 
in Tournejortia (e. g. T. hirsutissima and T. bicolor), does lend support 
to the universal recognition of these weak but practicable concepts. 

The present variety is that race of T. paniculata, with tawny some- 
what shaggy pubescence, long corolla-tubes and long acuminate corolla- 
lobes, which occurs about the head-waters of the Amazon and is geo- 
graphically separated from the typical form of T. paniculata, of eastern 
Brazil and Paraguay, by the great tracts of the Amazon Basin. It is a 
weak variety but certainly worthy of some recognition. The extreme 
form is well exemplified by the type-collection of T. spigeliaejiora, col- 
lected by Schomburgk and labeled as from Pirara. In Robert Schom- 
burgk's notes at Kew the type-number (749) appears in a list of plants 
(nos. 701-769) sent out to the coast from Pirara on June 25, 1838. In 
this list only six numbers are provided with definite localities, these all 
being given as from Pirara. Richard Schomburgk in his published cata- 
logue, 1. c. 1151, lists T. spigeliaejiora only from the "vicinity of Pirara 
at the edge of the oasis." This locality is on the margin of the Rio 
Branco watershed. 



1935] JOHNSTON, STUDIES IN BORAGINACEAE 51 

4. Tournef ortia Ulei Vaupel, Notizbl. Bot. Gart. Berlin 6:186 
(1914); Johnston, Contr. Gray Herb. 92: 70 (1930). T. Miquelii 
Macbride, Proc. Amer . Acad. 51:541(1916). T. syringae folia of most 
authors, e. g. Miquel, Stirp. Surinam. 137, tab. 41 (1850) ; Pulle, Enum. 
PI. Surinam 398 (1906), excl. of Splitgerber 841. 

Shrub or liana; branchlets puberulent; leaves rather thin, ovate or 
ovate-elliptic or rarely broadly lanceolate, 6-17 cm. long, 3-8 cm. broad, 
base obtuse, apex acuminate, surfaces glabrous except for inconspicuous 
puberulence on the veins beneath, frequently with numerous scattered 
minute usually pale tuberculations; inflorescence loosely branched, the 
racemes loosely flowered and becoming 2-10 cm. long; calyx 1.5-2 mm. 
long, weakly accrescent, with triangular or subulate lobes, sessile or 
shortly and distinctly pedicellate; corolla 7-8 mm. long, greenish white, 
tube 3-4 mm. long, throat 1-1.5 mm. long, inflated, limb 2-3 mm. broad, 
lobes ovate ca. 1 mm. long; fruit glabrous, 4-5 mm. thick, not quite so 
long, broadest below middle; style well developed, becoming 2-3 mm. 
long, usually persistent; stigma clavate. 

Known from the Guianas and in the head-waters of the Amazon south 
to Bolivia, 

British Guiana: Arawak Matope, Cuyuni River, Oct. 1904, Bartlett 
8333 (K) ; upper Demerara River, Sept. 1887, Jenman 4117 (K, NY). 

Dutch Guiana : Surinam River near Bergendal, Focke 1308 (Utr, type 
of T. Miquelii) ; road near Brownsberg, 1910, native collector 170 (Utr) ; 
Commewyne River, Focke (K) ; Surinam, Focke 121 (Leid). 

French Guiana: Maroni River near Apatou, Oct. 1901, Went 458 
(Utr) ; He Portal, Maroni River, May 1857, Sagot 1011 (K, P) ; St. Jean, 
fl. greenish. May 18, 1914, Benoist 1247 (P) ; Charvein, fl. green, Jan. 
20, 1914, Benoist 648 (P) ; Roura, 1858, Sagot (P) ; Cayenne, 1859, Sagot 
(P) ; indefinite, Poiteau (K, BD). 

In the Guianas this plant has been generally misdetermined as T. 
syringae j alia, a name properly applicable to the very different plant that 
has been called T. peruviana. The first name for our very distinct spe- 
cies is T. Ulei, based upon material collected by Ule in extreme south- 
western Brazil. The name T. Miquelii Macbr., is based upon a plate 
published by Miquel, 1. c. Since this is evidently drawn from the speci- 
men at Utrecht collected at Bergendal by Focke (no. 1308), that speci- 
men may be considered as the type of T. Miquelii. It is quite like 
T. Ulei. 

5. Tournefortia bicolor Swartz, Prodr. 40 (1788) and Fl. Ind. Dec. 
1: 344 (1797); Johnston, Contr. Gray Herb. 92: 69 (1930). T. lae- 
vigata Lamarck, Tab. Encyc. 1: 416 (1791); Fresenius in Martius, Fl. 
Bras. 8': 49 (1857). T. laevigata var. latijolia DeCandolle, Prodr. 



52 JOURNAL OF THE ARNOLD ARBORETUM fvoL. xvi 

9: 519 (1845); Schomburgk, Fauna u. Fl. Brit. Guian. 1151 (1848). 
T. glabra Aublet, Hist. PI. Guian. Fr. 1: 118 (1775), not Linn. (1753). 
T. Aubletii Macbride, Proc. Amer. Acad. 51 : 541 (1916). 

Shrub 1-5 m. tall, becoming subscandent; branchlets with weak short 
usually sparse ascending appressed hairs, rarely glabrous or puberulent ; 
leaves ovate to elliptic or lance-ovate, 5-14 cm. long, 3-9 cm. broad, sub- 
coriaceous, base obtuse to rounded, apex acute, upper surface slightly 
lustrous, bearing a few scattered weak short appressed hairs, smooth or 
bearing very minute inconspicuous papillae, lower surface slightly more 
hairy than upper; petioles 5-15 mm. long; inflorescence dense, branched, 
5-20 cm. broad, racemes becoming 1-4 cm. long; calyx sparsely strigose 
at anthesis with lobes lanceolate or ovate, 1-2.5 mm. long, weakly 
accrescent, usually sessile; corolla white, tube 4-5 mm. long, strigose 
outside, about twice length of calyx, limb 6-7 mm. broad; fruit white, 
very fleshy, ca. 8 mm. long, glabrous; stigma subsessile. 

Widely distributed in the American Tropics. 

British Guiana: Barima River. 2.5-3.5 m. tall. fl. white, March 1923, 
La Cruz 2372 and 2372 (G) ; Issorora, Aruka River, wet forest, tree 10 
m., Jan. 1923, Hitchcock 17560 (G, NY) ; Arawak Matope, Cuyuni River, 
fl. white, Oct. 1904, Bartlctt 8222 (K) ; near Pirara, Feb. 1842, Schom- 
burgk (BD). 

Dutch Guiana: Coppename River, fl. white Sept. 1901, Boon 1112 
(Utr) ; banks of Surinam River below Kabel, liana, fl. white, 1933, Lan- 
jouw 1221 (Utr) ; Brownsberg, tree no. 66, B. IV. 2242 (Utr) ; Browns- 
berg Summit, 1924, B. W. 6512 and 6714 (Utr) ; Brownsberg, 1915, B. IV. 
711 (Utr) ; Tapanahoni River, scandent shrub, fl. white, Oct., Kappler ed. 
Hohenackcr 2094 (BD) ; Maroni River, shrub 4-5 m., July 1904, Vcrsteeg 
714 (Utr) ; indefinite, 1862, Kappler 127 (Leid). 

French Guiana: La Mana, 1823-24, Leschenault (P) ; Cayenne, 
1786-91, z'on Rohr (BM) ; indefinite, 1820, Pcrrottet (Deles). 

5a. Toumefortia bicolor var. calycosa Bonn. Smith, Bot. Gaz. 
14: 27 (1889); Johnston, Contr. Gray Herb. 92: 70 (1930). T.Schom- 
burgkii DeCandolle, Prodr. 9: 517 (1845); Schomburgk, Fauna u. Fl. 
Brit. Guian. 961 (1848). T. alba Splitgerber ex DeVriese, Nederl. 
Kruidkund. Arch. 1:347 (1848); Schomburgk, Fauna u. Fl. Brit. 
Guian. 1151 (1848). (?) T. coriacea Vaupel, Bot. Jahrb. 54, Beibl. 
119: 3 (1916). 

Calyx 3-4 mm. long, the lobes linear or lanceolate; plant tending to 
be slightly more pubescent than in the species. 

Dutch and British Guiana and southwestward across the Amazon 
Basin to Peru and Ecuador; also in Guatemala and Honduras. 

British Guiana: Rockstone, bank of Essequibo, July 31, 1921, Gleason 
897 (NY, K) ; bank of Corantyne River, shrub 2.5-3 m. tall, Sept. 1878. 



1935] JOHNSTON, STUDIES IN BORAGINACEAE 53 

im Thurn (K, P) ; Oreala, Corantyne River, trailing over bushes, Nov. 
1879, Jenman 120 (P) ; Epira, Corantvne River, trailing over bushes, Nov. 
1879, Jenman 65 (P) ; Berbice, 1837, Schombnrgk 70 (Deles) ; indefinite, 
Schomhurgk 70 (DC, type of T. Schomhurgkii; BM, Leid, BD, P) ; 
indefinite, ligneous twiner, fl. white, Schomhurgk 70 (K). 

Dutch Guiana: Apoera Island, Corantyne River, fl. white, June 22, 
1916. B. W. 2043 (Utr) ; Wilhelmina Range, Peak no. 1200, June 9, 1926, 
B. IV. 7060 (Utr) ; near Plant. Catharina Sophia, Saramacca River, shrub 
1-2.5 dm. high, in shade, fl. white, April 1838, Splitgerber 841 (Leid, type 
of T. alba; isotype, P). 

Brazil: Rio Negro near confluence with Rio Solimoes, May 1851, 
Spruce 1491 (G, K, BM). 

The widely distributed "T. bkolor Sw." is reported by Schomburgk, 
1. c. 830, as cultivated as a decorative shrub in the coastal regions of 
British Guiana. Tournejortia glabra Aubl. is based entirely upon an 
unpublished plate by Plumier, (manuscripts at Paris vol. 6, tab. 53). 
This plate represents T. bkolor and is drawn from specimens obtained 
at Leogane in Haiti. The name T. Aubletii Macbr., is a mere renaming 
of T. glabra Aubl., because of an earlier Linnaean homonym. In a strict 
sense, therefore, both T. glabra Aubl. and T. Aubletii Macbr. are really 
West Indian plants. 

The var. calycosa is based upon material from Guatemala. In Cen- 
tral America it appears to be rare and restricted to eastern Guatemala 
and adjacent Honduras where it is found in the same regions as T. bi- 
color and T. hirsutissima. In South America it occurs in the region to 
the east and south of the Orinoco Basin and hence far separated from 
T. hirsutissima L., which in South America is known only from north- 
ern Venezuela and Colombia. While the var. calycosa is not known to 
grow with T. bicolor in South America, it does occur in Ecuador, Peru 
and the Guianas where the species has been much collected. The variety 
seems to be a plant of more wet forests than those usually selected by 
T. bicolor. A study of T. bicolor from all parts of its very extensive 
range shows it to be remarkably uniform in the size of its calyx. The 
uniformly and evidently more elongate calyx-lobes of the var. calycosa 
merit some nomenclatorial recognition particularly since this variation 
seems to be geographically localized. 

The variety has been frequently confused with T. hirsutissima, but 
that species may be readily distinguished from both T. bicolor and the 
var. calycosa by its more abundant, more spreading hairs on the herb- 
age, and particularly by the stiff erect or ascending hairs (usually from 
a somewhat bulbose base) that according to abundance give a hirsute to 
velvety covering to the upper surfaces of the leaves. The fruit in T. 
hirsutissima, furthermore, is usually hairy. In T. bicolor and variety 



54 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

the somewhat glossy upper leaf-surfaces have weak scattered and com- 
monly inconspicuous appressed hairs. The fruit is glabrous. The stems 
are usually sparsely hairy or glabrous. The calyx of T. hirsutissima is 
very variable in length. It may be short to elongate in the various parts 
of the range of the species. The two species, T. bkolor and T. kirsutis- 
sima, are very closely related and the differences separating them are 
almost exclusively those of pubescence mentioned. The difference seems 
to be decisive, however, and the resulting concepts natural and 
practicable. 

The type of T. alba is the common form of the var. calycosa, it has 
the pubescence, particularly on the stems, sparse and scarcely, if at all, 
more abundant than is commonly found in T. bkolor. The type of T. 
Schomburgkii agrees with other collections from British Guiana in hav- 
ing conspicuously and rather densely hairy stems. In this regard it 
tends to suggest T. hirsutissima with which it has been confused. 
Schomburgk's original collection of T. Schomburgkii (no. 70), was dis- 
tributed labeled as from "British Guiana" or from Berbice. In his cata- 
logue, 1. c. 961, he states that it grows on the banks of the Essequibo and 
makes no mention of Berbice. In closing it may be added that the type 
of T. alba, Splitgerber 841, was incorrectly cited under T. syringaejolia 
(equals T. Ulei) by Pulle, Enum. PI. Surinam 398 (1906). 

6. Tournefortia cuspidata Humboldt, Bonpland & Kunth, Nov. 
Gen. et Sp. 3: 83 (1818). T. obscura A. DeCandolle, Prodr. 9: 517 
(1845); Schomburgk, Fauna u. Fl. Brit. Guian. 961 (1848); Fresenius 
in Martius, Fl. Bras. 8^:49, adnot. (1857); Johnston, Contr. Gray 
Herb. 92: 68 (1930). T. setifera Urban & Ekman, Ark. Bot. 22a, 
no. 17: pg. 94 (1929). T. hirsutissima sensu Pulle, Enum. PI. Surinam 
398 (1906). 

Shrub or liana; branchlets pubescent, also conspicuously shaggy with 
abundant slender spreading brown hairs 2-4 mm. long; leaves lance- 
ovate to lanceolate, 7-15 cm. long, 3-6 cm. broad, base rounded or 
obtuse, apex acuminate, both surfaces with abundant appressed slender 
elongate hairs; petioles 5-13 mm. long; inflorescence stiffly and loosely 
branched, the spikes becoming 1-3 cm. long and crowded at the ends 
of the elongate branches; calyx-lobes subulate or linear, 7-9 mm. long 
at anthesis, weakly accrescent, sparsely long-hairy and short-strigose; 
corolla white, tube 5-8 mm. long, densely strigose outside, limb 4-6 
mm. broad, lobes broad; fruit white, fleshy, compressed ovoid, prob- 
ably ca. 8 mm. long, glabrous, more or less verrucose; stigma sessile. 

Northern South America (Dutch Guiana to Colombia) and south- 
ward, in the upper reaches of the Amazon Basin, to Bolivia, doubtfully 
from eastern Brazil; Central America; West Indies. 



1935] JOHNSTON, STUDIES IN BORAGINACEAE 55 

British Guiana: Batima, March 1896, Jenman 7118 (K) ; banks of 
the Quitaro, 1837, Schomburgk 571 (DC, type of T. ohscura; G, K, BM, 
Leid, BD) ; Berbice, June 1889, VVaby ex Jenman 5157 (K, BM) ; Ber- 
bice, Bunnann (Deles) ; Demerara, Parker (K). 

Dutch Guiana : Matappi, Corantyne River, liana, fl. white, June 18, 
1916, B. IV. 2168 (Utr) ; Brownsberg Summit, liana, fl. white, July 3, 
1924, B. W. 6570 (Utr) ; Goddo, upper Surinam River, Jan. 29, 1926, 
Stahd 134 (Utr) ; Pikien River, fl. white, July 1908, Tresling 203 (Utr) ; 
Maroni River near Armina Falls, small shrub, fl. white, 1933, Lanjouw 
526 (Utr) ; indefinite, Hostmann 227 (K, BM, Utr, BD, Deles, P). 

I have examined the types of the species above cited. They are evi- 
dently conspecific! The type of T. obscura is Schomburgk 571, labeled 
as from the banks of the Quitaro. The specimen was collected by Rob- 
ert Schomburgk. A study of his manuscript list at Kew shows that no. 
571 falls within the gamut (no. 511-588) of numbered collections sent 
out from Curassawaka (on the Essequibo) in Dec. 1837. According to 
the list these numbers apply to specimens from "the Quitaro in November 
and to a few on the River Rewa, but the greater part of the high num- 
bers from the mountains of Attarypou [Kanuku Mts.]." Only one 
number is provided with a definite locality, no. 581 being given as from 
the "mountains of Attarypou." The list gives the following field-notes 
for no. 571, "A ligneous twiner growing by river side, leaves light-green, 
a shade lighter below, calyx light green, petals and organs of fructifica- 
tion pure white, fruit a white berry." In Richard Schomburgk's pub- 
lished catalogue, 1. c. 961, T. obscura is reported from the banks of the 
Rupununi, Rewa and Quitaro rivers. 

In my paper on the Brazilian species of Tournejortia I reported this 
species, sub T. obscura, doubtfully from eastern Brazil. At Paris I have 
since seen a collection by Glaziou (no. 9981) labeled as from "Sao Joao 
da Barra" on Feb. 8, 1876. This locality is at the mouth of the Para- 
hyba River in the northeastern section of the State of Rio Janeiro. The 
same number is cited by Glaziou, Bull. Soc. Bot. France 57, Mem. 3e : 
478 (1910), under the name "T. Salzmanni" and as from "Sao Joao, 
pres Campos." At Kew this collection by Glaziou, no. 9981, is labeled 
as cultivated at Rio Janeiro. The species has evidently been collected 
in eastern Brazil but whether or not from cultivated plants is still to be 
settled. 

7. Tournefortia melanochaeta DeCandolle, Prodr. 9: 520 (1845). 

Shrub or liana; branchlets with scattered spreading brown hairs 1-2 
mm. long; leaves lance-ovate, 9-11 cm. long, 5-6 cm. broad, base obtuse 
to rounded, apex acuminate, both surfaces lustrous and glabrous except 
for a very few slender appressed hairs along the midrib and veins, 



56 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

petioles 10-13 mm. long; inflorescence stiffly and loosely branched, 
spikes becoming 10-15 mm. long and crowded at the ends of the 
elongate branches; calyx-lobes glabrous, lanceolate to linear, 4-7 mm. 
long, weakly accrescent; corolla white, tube 5-7 mm. long, densely stri- 
gose outside, limb 4-5 mm. broad, lobes broad; stigma sessile; fruit 
unknown but probably as in T. cuspidata. 

Known only from French Guiana. 

French Guiana: "Cayenne ou Guyana fran(;aise," Museum de Paris, 
1821 (DC, TYPE) ; Cayenne (Martin) ex Museo Horti Paris, 1819 (BD) ; 
Cayenne, Martin (G, P). 

The material cited is evidently all part of one large collection by 
Martin. It seems to be scarcely more than a glabrescent phase of T. 
cuspidata, a species which extends, from the westward, to the borders of 
French Guiana, though it is not as yet known to have been collected 
within that colony. 

Doubtful and Excluded Species 

Tournef ortia gnaphalodes (Linn.) R. Brown ex Roemer & Schultes, 
Syst. 4: 538 (1819). 

Schomburgk, Fauna u. Fl. Brit. Guian. 830 (1848), reports this plant 
from the coasts of British Guiana, while Lamarck, Encyc. 3: 94 (1789), 
and Aublet, Hist. PI. Guian. Fr. 1: 117 (1775) under the name Helio- 
tropium gnaphalodes, indicate its occurrence in French Guiana. The 
plant is widely distributed in the West Indies but does not reach south 
to Trinidad. I have seen no material of it from the Guianan coast and, 
furthermore, do not believe that it is native in the region. 

Tournefortia foetidissima Linnaeus, Sp. PL 140 (1753). 

Tournefortia hirsutissima Linnaeus, Sp. PI. 140 (1753). 

Tournefortia cymosa Linnaeus, Sp. PI. ed. 2, 202 (1762). 

The above three species are listed by Aublet, Hist. PI. Guian. Fr. 
1: 117-118 (1775), as occurring in French Guiana. They are West 
Indian species not known from the Guianas and, furthermore, not to be 
expected there. 

Tournef ortia incana (Meyer) Don, Gen. Syst. 4: 368 (1838), not 
Lamarck (1791). 

Tournef ortia Meyeri DeCandolle, Prodr. 9: 530 (1845). 

The above two names are based upon Messerschmidia incana Meyer 
from the mouth of the Essequibo. The plant seems to be a species of 
Ilcliotropium. 



1935] JOHNSTON, STUDIES IN BORAGINACEAE 57 

4. Heliotroplum [Tournef.] Linnaeus, Gen. 63 (1754). 

Mostly low herbaceous or suffrutescent plants, rarely shrubs; leaves 
small to large. Inflorescence of solitary, geminate or ternate scorpioid 
spikes or racemes or the flowers solitary cauline and internodal. Calyx 
persistent or deciduous, with 5 teeth or lobes. Corolla yellow, white or 
blue, small, 5-merous, tube cylindrical, limb spreading. Stamens 5, 
borne in the corolla-tube, included, filaments short. Ovary 4-celled; 
style terminal, solitary; stigma sessile or on a distinct style, peltate or 
conic, fertile on the side, apex bearing a conic or cylindrical sterile 
appendage that is usually bifid or bidentate. Fruit dry, lobed or un- 
lobed, at maturity breaking up into 2-4 bony nutlets. Nutlets 1-2- 
seeded, frequently with 1-2 sterile cavities. Endosperm developed. 

A large genus widely distributed in the warmer regions of the world. 
Type Species, H. europaeum L. 

Key to the Species 
Plant distinctly succulent, entirely glabrous, frequently somewhat 



into a conic apex 2. H. lagoense. 

Flowers borne in well developed scorpioid spikes or racemes. 
Corolla blue or purple; fruit glabrous, ribbed, angulate, 
deeply 2-lobed with the lobes horizontally divergent; 
plant coarse, erect, more or less hirsute with large evi- 
dently veined leaves 3-10 cm. broad 3. H. indicmn. 

Corolla white ; fruit strigose, rounded, weakly 4-lobed ver- 
tically ; plant rather slender, erect or decumbent ; leaves 
with appressed pubescence, very obscurely veined, less 
than 2 cm. broad. 
Leaf-blades elliptic to broadly oblanceolate, 5-20 mm. 
broad, petiole 4-15 mm. long; racemes mostly gemi- 
nate or ternate, bractless 4. H. procumbens. 

Leaf-blades linear to oblanceolate, 1-8 mm. broad, petiole 

1-2 mm. long; racemes always single, bearing small 

lanceolate bracts 1-2 mm. long. 

Corolla 4-6 mm. long, anthers joined at their apices; 

stigma borne on a short but evident style; stems 

comparatively coarse and stiff; leaves drying 

rather light colored 5. H. ternaturn. 

Corolla 2-2.5 mm. long, anthers not joined apically ; 
stigma sessile on the fruit; stems wiry, very slen- 
der, leaves usually drying dark colored 6. H. filifonne. 

1. Heliotropium curassavicum Linnaeus, Sp. PI. 1: 130 (1753); 
Schomburgk, Fauna u. Fl. Brit. Guian. 961 (1848); Johnston, Contr. 
Gray Herb. 81: 14 (1928). 



58 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

Annual or short-lived perennial, succulent, glabrous; stems prostrate 
or decumbent, 1-3 dm. long, ascendingly branched; leaves narrowly to 
broadly oblanceolate, 2-4 cm. long, 3-10 mm. broad, fleshy, frequently 
somewhat glaucous; flowers borne in bractless single or geminate scor- 
pioid spikes 1-10 cm. long; calyx ca. 1.5 mm. long at anthesis, over 
2 mm. long at maturity, sessile or subsessile, lobes broadly cuneate to 
triangular-ovate; corolla white, 2-3 mm. long, lobes ca. 1 mm. long, 
tube shorter than calyx; anthers 0.6-1 mm. long, sagittate, apex with 
acuminate appendage, not joined together; fruit weakly 4-lobed, epi- 
carp slightly fleshy and wrinkled in drying; style sessile, disk of stigma 
broad; nutlets 4, equal, oblong, 2-2.5 mm. long, 1-celled, 1-seeded. 

From the coast of British Guiana, Venezuela and Colombia south- 
ward along the west coast of South America to central Chile and Pata- 
gonia, and northward through the West Indies and in Central America 
to southern-most United States. 

British Guiana: Georgetown, Hitchcock 16572 (G, NY); indefinite, 
seashore, June, 1889, Jenman 5471 (US, BM) ; indefinite, Jemnan 2165 
and 4466 (NY). 

The species is listed as occurring in French Guiana by Aublet, Hist. 
PI. Guian. Fr. 1 : 117 (1775). I consider this record very question- 
able. It may be noted that I have examined and made dissections of 
the type of H. Lehmannianum Bruns, Mitt. Inst. Allg. Bot. Hamburg, 
8:69, fig. 10 (1929), recently described from the coast of southern 
Peru. My study has shown the type to be quit^ ordinary H. curassavi- 
cum. Bruns described and illustrated remarkable developments in 
corolla-lobing and in anther-shape. In the various dissections made on 
the type I could find no suggestion of remarkable structures, the corolla 
and the anthers agreeing perfectly with the familiar typical West Indian 
H. curassavicum. 

2. Heliotropium lagoense (Warm.) Gurke in Engler & Prantl, Nat. 
Pflanzenf. IV. Abt. 3a: 97 (1893) ; Johnston, Contr. Gray Herb. 81 : 49 
(1928). Schleidenia lagoensis Warming, Kjoeb. Vidensk. Meddel. 
1867: 15 (1868). Heliotropium trinitense Urban, Symb. Ant. 7: 350 
(1912). 

Annual herb with scattered slender appressed hairs; stems slender, 
prostrate, 5-30 cm. long, ascendingly branched; leaves oblanceolate, 
0.5-1.5 cm. long; flowers borne singly along the leafy stems, extra- 
axillary; calyx of 5 unequal lanceolate or cuneate lobes, at anthesis 
1.5-2 mm. long, becoming about twice as long in fruit; pedicels 1-3 mm. 
long, ascending; corolla white, 3-4 mm. long, funnelform; lobes ovate, 
1-1.5 mm. long; sinus rounded, plaited, occasionally with a minute 



1935] JOHNSTON, STUDIES IN BORAGINACEAE 59 

lobule; anthers oblong, bearing an obese hairy apical appendage nearly 
as large as the anther proper, anthers joined together by their append- 
ages; fruit glabrous or nearly so, subterete, base rounded, from at or 
below middle contracted upwardly into a broad conic or short-rostrate 
apex; nutlets 4, ca. 1.5-2 mm. long, single seeded. 

Northern Dutch Guiana, Trinidad, northern Venezuela, eastern 
Bolivia, and eastern and western Brazil; not common and apparently 
local and erratic in distribution. 

Dutch Guiana: "Suriname," Jan. 1885, Suringar (Leid). 

This species is frequently confused with H. filiforme, which it re- 
sembles in its wiry stems and slender leaves, but it is readily distinguished 
from that plant by its conic fruit and its cauline internodal flowers. 
Although Suringar's collection has no definite locality it is to be sup- 
posed that it is from the vicinity of Paramaribo wheje Suringar is known 
to have done most of his collecting in Dutch Guiana. 

3. Heliotropimn indicum Linnaeus, Sp. PI. 130 (1753); Aublet, 
Hist. PI. Guian. Fr. 1: 117 (1775); Pulle, Enum. PI. Surinam 399 
(1906); Johnston, Contr. Gray Herb. 81: 19 (1928). Heliophytum 
indicum (Linn.) DeCandolle, Prodr. 9:556 (1845); Schomburgk, 
Fauna u. Fl. Brit. Guian. 831 and 961 (1848). 

An erect coarse weedy annual herb, usually more or less pale-hirsute, 
1-10 dm. tall, mostly branched above the middle; leaves ovate or elliptic 
to broadly lanceolate, herbaceous, veined, 5-15 cm. long, 3-10 cm. 
broad, margin repand or undulate, apex acute, base obliquely acute to 
subcordate, petioles 4-10 cm. long, winged just below the leaf-blade; 
flowers borne in bractless single scorpioid spikes becoming 5-30 cm. 
long; calyx with subulate or cuneate lobes 2-2.5 mm. long, somewhat 
accrescent in fruit; corolla blue or violet or exceptionally white, salver- 
form, tube 2.5-4.5 mm. long, evidently surpassing the calyx, limb 2-4 
mm. broad; anthers elongate, the apices not united; style short and 
slender; fruit glabrous, strongly ribbed, deeply 2-lobed (the lobes 
spreading) and breaking up into 4 angulate nutlets 2-2> mm. long. 

A Pan-Tropic weed, probably of American origin. 

British Guiana: Comaca, Moruka River, La Cruz 1058 (NY, US) ; 
Waranuri Mission, Moruka River, Oct. 1922. La Cms 2600 (G, NY, US) ; 
Kamakusa, upper Mazaruni River, Nov. 1922, La Cruz 2748 (G, NY) ; 
Tumatumari, Gleason 370 (NY) ; Hyde Park, east bank of Demerara 
River, Dahlgren (FM) ; Demerara, Parker (K) ; Georgetown, weed 
along canal, fl. white, 1919, Hitchcock 16684 (G, NY, US) ; Epira, banks 
of Corantyne River, Nov. 1879, Jenman 54 (K) ; Corantyne River, Oct. 
1879, im Thurn (P) ; indefinite, Schomburgk 206 (BD) and 600 (K). 

Dutch Guiana: bank of Corantyne River, 1911, Hulk 33 (Utr) ; sand 



60 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

near Maripaston, Saramacca River, Nov. 1902, Pidle 4 (Utr) ; Para- 
maribo, fl. bluish, Kuyper 25 (Utr) ; near Paramaribo, fl. blue, Dec. 1837, 
Splitgerber 338 (Leid, P) ; Combee, Paramaribo, Aug. 1901, Went 319 
and 332 (Utr); Plant. Rust en Werk, 1913, Soeprato 57 (Utr); upper 
Surinam River near Saida, 1908, Trcsling 346 (Utr) ; Plant. Slootwijk, 
Commewyne River, 1913, Soeprato 15J (Utr) ; beach facing Cottica Mt., 
Law^a River, fl. white with yellowish throat, light lilac when young, Oct. 
1903, Verstceg 289 (Utr); Cottica River near Moengo, marshy ground, 
fl. light blue, 1933, Lanjouw 403 (Utr) ; indefinite, Focke 1370 (Utr). 

French Guiana: Maroni, Wachenheim 287 and 290 (P), 291 (BM) ; 
St. Jean, fl. pale blue, April 26, 1914, Benoist (P) ; St. Jean, fl. pale blue 
with yellowish center, March 8, 1914, Benoist 1154 (P) ; St. Laurent du 
Maroni, Jan. 1908, attractive to butterflies, LeMoult (P) ; Acarouani 
1855, about dwellings, fl. bluish, Sagot 449 (P) ; vicinity of Cayenne, fl. 
purple, May 6, 1921, Broadway 118 (K); Cayenne, Feb. 18, 1845, Rothery 
165 (BD) ; lies du Salut, 1854, fl. pale bluish, Sagot 449 (BM, P). 

Brazil: Obidos, Spruce 476 (K) ; Monte Alegre, 1873, Traill (K). 

Venezuela : Ciudad Bolivar, 1931, Holt & Blake 838 (G) ; Las Batillas, 
Passarge & Selwyn 301 (BD) ; Puerto Ayacucho, 1931, Holt & Blake 834 
(G). 

In Dutch Guiana this species is called "Kaha Kankay" {Tresling no. 
346) and "Kokorrode" {Pulle no. 4) and is given as being used with 
salt as a cure for gas on the stomach {Versteeg no. 289). According to 
Aublet, 1. c, it is called "Cret-de-Coq" in French Guiana and an infusion 
of the flowers used "pour arreter les pertes de sang chez les femmes." 

4. Heliotropium procumbens Miller, Diet. ed. 8, no. 10 (1768); 
Johnston, Contr. Gray Herb. 81: 52 (1928). Heliotropium inundatum 
Swartz, Prodr. 40 (1788). 

Annual herb with more or less abundant slender appressed hairs, plant 
usually cinereous; stems erect or decumbent, 1-5 dm. long, ascendingly 
branched ; leaves with elliptic, obovate or broadly oblanceolate blades, 
1-4 cm. long, 5-20 mm. broad, petioles slender 4-15 mm. long; flowers 
borne in slender scorpioid racemes, racemes mostly geminate or ternate, 
bractless, elongating, becoming 2-10 cm. long, peduncles 1-4 cm. long; 
calyx with 5 unequal lanceolate or linear lobes, at anthesis 0.5-1 mm. 
long, becoming 1-2.5 mm. long in fruit, pedicels ca. 0.5 mm. long; corolla 
white 1-5 mm. long, lobes ovate, short, with rounded sinus; anthers 
ovate, contracted apically into short narrow appendages, anthers not 
joined apically; fruit depressed globose, 4-lobed, strigose; stigma sessile; 
nutlets strigose, 0.5-1 mm. long. 

Northern Argentina northward through Tropical America to southern 
United States; rare in the very wet regions. 

British Guiana: indefinite: Appun 1762 (K). Schomburcfk 1024 (K, 
BD) and 1026 (K). 



1935] JOHNSTON, STUDIES IN BORAGINACEAE 61 

Brazil: Prainha, Traill (K) ; Alemquer, Spruce (K). 

5. Heliotropimn ternatum Vahl, Symb. Bot. 3: 21 (1794); John- 
ston, Contr. Gray Herb. 81: 69 (1928). Heliophytum passerinoides 
Klotzsch ex Schomburgk, Fauna u. Fl. Brit. Guian. 1152 (1848), nomen. 
Schleidenia Fumana Fresenius in Martius, Fl. Bras. 8^:40 (1857). 
Heliotropium Fumana (Fresen.) Giirke in Engler & Prantl, Nat. Pflan- 
zenf. IV. Abt. 3a: 97 (1893); Johnston, Contr. Gray Herb. 81: 71 
(1928). Heliotropium sp., Oliver, Trans. Linn. Soc. London, Bot. 
2: 279 (1887). Heliotropium strictissimum sensu N. E. Brown, Trans. 
Linn. Soc. London, Bot. 6: 51 (1901). Heliotropium jruticosum of 

Suffrutescent, with abundant ascending or appressed hairs; stems 
erect or decumbent, 1-5 dm. long, ascending branched; leaves lanceo- 
late to linear, revolute, 1-3 cm. long, 1-8 mm. broad, with slender 
petiole 1-2 mm. long; flowers borne in stiff scorpioid racemes; racemes 
single, elongating and becoming 2 15 cm. long, bearing scattered lanceo- 
late bracts 2-3 mm. long; calyx at anthesis 2-i mm. long, consisting of 
5 more or less unequal lanceolate lobes, becoming twice as large in 
maturity; pedicels 0.5-1 mm. long; corolla white, 4-6 mm. long, lobes 
ovate, sinus rounded and plicate; anthers ovate with short obtuse hairy 
apical appendages which are apically joined to one another; fruit de- 
pressed globose, 4-lobed, strigose; style short but evident; nutlets 1-1.5 
mm. long. 

Central America and the West Indies, northern Colombia and Vene- 
zuela, southern British Guiana and eastern Brazil; chiefly in open 
places or in dry thickets. 

British Guiana: Kamakot, Ireng River, 1884-85, Jenman 3 (US); 
Konkarmo, Ireng Valley, Nov. 16, 1884, im Thurn 3 (K, BM) ; Ireng 
Valley, Quelch &- McConnell 220 (K, BM) and 302 (K) ; Rupununi, 
Appun 2203 (K) ; Rupununi, May 1842, Schomburgk 573 (BD, type of 
H. passerinoides) ; Pirara, etc., 1841-42, Schomburgk 282 (P) ; indefinite, 
gravelly savanna, leaves appear farinaceous, fl. white, 1836, Schomburgk 
(K). 

In and about the West Indies H. ternatum is generally recognized as 
a variable species in habit, pubescence and leaf-shape. The typical and 
most common form of it has loosely revolute leaves 3-5 mm. broad and 
a loosely appressed indument of slender hairs. This plant appears to 
be rare or absent in our area. In the Guianas the species seems to be 
represented only by a form found in the savannas near the Brazilian 
border. This has linear leaves 1-3 mm. broad and is silky strigose. It 
is quite like plants from eastern Brazil that have been described as H. 
Fumana. In my monograph, 1. c. 81: 69-71 (1928), //. Fumana was 



62 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

treated as doubtfully distinct from H. ternatum. Subsequent consid- 
eration of its relation with H. ternatum, in the light of new material, has 
led me to the belief that it is only a pronounced form, possibly a savanna- 
ecad of that species. Much of the material from the dry eastern corner 
of Brazil, which formerly I placed in H. Fumana (and even some that 
I placed in H. saUcoides Cham.) I now refer unhesitatingly to H. terna- 
tum. Only the material with narrow sublinear leaves and distinctly silky 
strigose indument should be placed under H. Fumana. This form comes 
from more interior, more moist localities than those in which H. ter- 
natum is usually found. Transitional forms are common. If the linear- 
leaved, silky-strigose form of the savannas needs to be recognized, it 
may be called Heliotropium ternatum var. Fumana (Fresen.), comb. 

Both H. ternatum and the variety may be distinguished from H. saU- 
coides by having white rather than bright yellow corollas. The leaves 
also dry much lighter in color in H. ternatum and the pubescence of the 
herbage is not so strongly spreading and so coarse as in H. saUcoides. 
Warming's Schleidenia subracemosa, which I placed under H. saUcoides, 
I now refer to H. ternatum. Schleidenia dasycarpa Fresen. is also based 
upon material referable to H. ternatum. I have recently examined the 
types of these Brazilian species. 

The only Guianan specimen of this species at Berlin that was col- 
lected by Schomburgk (no. 573) is labeled as from the Rupununi. It 
bears Klotzsch's name, HeUophytum passerinoides, and is evidently the 
type of that undescribed species. Schomburgk, in his published list, 
however, gives H. passerinoides Kl. only from the savannas near the 
Takutu River. That stream joins the Ireng not far west of Pirara which 
in its turn is even a shorter distance west of the Rupununi. All of 
Schomburgk's specimens, under their various labels, probably came from 
the general region to the west of Pirara. From this region H. ternatum 
var. Fumana extends up the Ireng where others have collected it. 

The typical form of the species is either very rare or absent in the 
coastal region of the Guianas. Possibly the region is too wet. The 
description of the puzzling Messerschmidia incana Meyer, from the 
mouth of the Essequibo, suggests H. ternatum in all except the fruit. 
The doubt surrounding Meyer's plant, however, forces me to leave it 
unplaced. 

6. Heliotropium filiforme Lehmann, Getting. Gel. Anzeigen 1817: 
1515 (1817) and Asperif. 1:37 (1818); UeCandolle, Prodr. 9:545 
(1845); Pulle, Enum. PI. Surinam 399 (1906); Johnston, Contr. Gray 
Herb. 81 : 61 ( 1928) . Schleidenia fiUformis (Lehm.) Fresenius in Mar- 



1935] JOHNSTON, STUDIES IN BORAGINACEAE 63 

tius, Fl. Bras. 8^:40 (1857). Heliotropium helophilum Martius, 
Flora Regensb. 21^ Beibl. 4: 85 (1838) and Herb. Fl. Bras. p. 165, 
no. 267 (1841); DeCandolle, Prodr. 9:544 (1845); Schomburgk, 
Fauna u. Fl. Brit. Guian. 961 (1848) ; Miquel, Stirp. Surinam. 136, tab. 
40 (1850). 

Annual herb, sparingly strigose; stems slender, erect or decumbent, 
1-4 dm. long, ascendingly branched; leaves 1-2.5 cm. long, 1.5-3.5 mm. 
broad, oblanceolate, petiole very slender, 1-2 mm. long; inflorescence 
consisting of very slender scorpioid racemes, these solitary, elongating, 
becoming 2-15 cm. long, bearing minute scattered lanceolate bracts, 
1-2 mm. long; flowers 1-3 mm. distant, strict, numerous; calyx of 5 
unequal lanceolate or lance-ovate lobes, at anthesis 1.5-2 mm. long, be- 
coming about twice as long in fruit, pedicels becoming 0.5-1 mm. long 
in fruit; corolla white, funnelform, 2-2.5 mm. long; lobes ovate with 
broad open sinus, ca. 1 mm. long; anthers each contracted into a short 
puberulent apical appendage, not coherent; fruit depressed globose, 
obscurely 4-lobed, strigose; stigma sessile or subsessile; nutlets almost 
1 mm. long. 

Eastern Bolivia and Paraguay northward through Brazil to Vene- 
zuela and the Guianas; also in Trinidad and British Honduras; grow- 
ing in sand, usually near water. 

British Guiana: Essequibo River, Demerara, 1881, Jcnman 1095 (K, 
P) ; Essequibo, Jan. 1842, Schomburgk 321 (BD) ; upper Rupununi, 
Appun 2394 (K) ; Pirara, 1845, Schomburgk 228 (BD, P) ; Berbice, sandy 
soil, 1837, Schomburgk 351 (K, BD, P) ; indefinite, Schomburgk 228'/ 
321b (K). 

Dutch Guiana: Corantyne River, sandy places near Wonotobo, 1916, 
B. IV. 2866 (Utr); Avanavero Rapids, Kabalebo River, in sand, 1920, 
B. IV. 4653 (Utr); island in Lucie River, sandv soil, 1910, Hulk 398 
(Utr); lower Saramacca River, in sand, Nov. 1902, Pulle 69 (Utr); 
lower Surinam River, April 1846, Kapplcr. ed. Hohenackcr 1810 (Utr, 
BD); Maroni River, rocks at Armina Falls, 1901, Went 467 (Utr); 
Maroni River near Bonnidoro, in sand, fl. white, Kappler, cd. Hohenackcr 
2095 (BD) ; Lawa River, sandy flat, Oct. 1903, Vcrsteeg 277 (Utr) ; in- 
definite, 1862. Kappler 158 (Leid). 

French Guiana: Maroni, 1862, ,Rcch (P) ; Cavenne, Martin (P) : 
Cayenne, 1853, Rothery 209 (Cambr.) ; Oyapock, Oct. 1844, collector not 

Brazil: Monte Ale^re, Traill 570 (K) ; middle Rio Cumina. Dec. 24, 
1928, Sampaio 5906 (BD) ; Rio Cumina, Cataract of Tronca. Sept. 18. 
1928, Sampaio 5011 (BD) ; Rio Trombetas, Lag. Cavpuru. Traill 568 
(K) ; Barra Rio Negro, Spruce 1115 (G, K). 

Venezuela: Angostura. 1864. Grosoudy (P) ; Puerto de Tablas. Can- 
ton de Upata, 1864. Grosoudy (P); Tigrito. Passargc & Schvxn 515 
(BD) ; Las Botillas, Passargc & Sekvvn 302 (BD). 



64 JOURNAL OF THE ARNOLD ARBORETUM Fvol. xvi 

Doubtful and Excluded Species 

Heliotropium fruticosum L. ex Aublet, Hist. PI. Guian. Fr. 1: 117 
(1775). 

Although listed by Aublet I believe that this species does not occur 
in French Guiana. The binomial was applied to H. tematum in many 
of the older books. But neither this latter species nor the one properly 
called H. fruticosum is known or even to be expected in French Guiana. 

Heliotropium latifolium Willd. ex Schomburgk, Fauna u. Fl. Brit. 
Guian. 961 (1848). 

Listed by Schomburgk as distributed through the forests in the 
northern parts of British Guiana. The species (and the cited synonym, 
H. scorpioides HBK.) is a synonym of H. angiospermum Murray 
(= H. parvijiorum L.). Although this plant is known from eastern 
Brazil and from northern Venezuela, no specimens have been seen from 
the Guianas. 

Messerschmidia incana Meyer, Prim. Fl. Esseq. 92 (1818). Tour- 
nejortia incana Don, Gen. Syst. 4: 368 (1838), not Lamarck (1791). 
Tournejortia Meyeri DeCandolle, Prodr. 9: 530 (1845); Schomburgk, 
Fauna u. Fl. Brit. Guian. 1151 (1848). 

The description given by Meyer makes it evident that this plant must 
be a Heliotropium, rather than a Tournejortia, if indeed it really is a 
member of the Boraginaceae at all. The original description strongly 
suggests H. tematum Vahl in all but the fruit, which is given as con- 
sisting of two biovulate sub-trilocular nutlets. The type was collected 
on the west bank of the Essequibo River near its mouth, where it is 
given as growing in dry places. Until the type (probably at Goettingen) 
is examined the species must remain very puzzling and unplaced. 



REHDER, HANDELIODENDRON 



HANDELIODENDRON, A NEW GENUS OF SAPINDACEAE 



With plate 119 and one text figure 

Handeliodendron, gen. nov. 

Flores ut videntur hermaphroditi, symmetrici, satis parvi, albescentes; 
sepala 5, libera, imbricata, ovato-oblonga vel oblonga, obtusiuscula, uni- 
nervia, ciliolata, extus intusque puberula, basin versus ut pedicellus 
squamulis patelliformibus obsita; petala 4 vel interdum 5, sepalis duplo 
longiora, imbricata, oblonga, obtusiuscula, basin versus sensim in 
unguem attenuata, supra basin lamellis 2 elevatis instructa, extus ad- 
presse pubescentia, intus glabra, ciliolata, medio reflexa; discus lateralis 
pulvinaris, irregulariter lobulatus, latere staminibus opposite concavus, 
fere aeque latus quam altus; stamina 7, raro 8, inaequalia, longiora 
petalis subaequilonga, sed ob petala recurvata manifeste exserta, fila- 
mentis leviter sursum curvatis, apice excepto villoso-pilosis, antheris 
late ovalibus mucronulatis, infra medium dorsum affixis; ovarium late 
fusiforme, longiuscule stipitatum, in stylum brevem apice stigmatibus 3 
brevissimis conicis coronatum attenuatum, triloculare; ovula in quoque 
loculo 2, alterum erectum, alterum pendulum. Capsula piriformis, 
leviter 3-loba vel abortu 2-loba vel simplex, stipitata, loculicida, peri- 
carpio coriaceo brunneo lenticellis albidis consperso; semina in quaque 
capsula 1-4, ovoidea, testa crustaceo-coriacea, atra, nitida, hilo brunneo 
parvo elliptico, arillo albido duplici circiter semen medium tegente, e 
trichomatibus cohaerentibus constituto, exteriore hilum cingente ab 
interiore annulo incrassato separate. Embryo vix curvatus, cotyledoni- 
bus plano-convexis fere rectis, basi tantum curvatus et in radiculam 
dorsalem gracilem in plica testae immersam et fere ad micropylem de- 
scendentem contractus. 

Arbor alta, cortice griseo, ramulis hornotinis brunneis glabris, anno- 
tinis lenticellatis spadiceis. Folia opposita, glabra, digitata, foliolis 5 
inaequalibus petiolulatis ellipticis vel elliptico-obovatis, abrupte in acu- 
men caudatum productis, bais late cuneatis in petiolulum decurrentibus, 
supra laete viridibus subtus pallidioribus et glandulis scutellatis initio 
fusco-rubris demum nigrescentibus sparse vel sparsissime conspersis, 
pinnatinerviis, nervis utrinsecus 9-12 patentibus arcuatis supra levissime 
subtus magis elevatis margine anastomosantibus, costa supra elevata 
sed in canaliculo plus minusve immersa subtus manifeste elevata; petio- 



66 JOURNAL OF THE ARNOLD ARBORETUM 1vol. xvi 

lis gracilibus teretibus basi tantum leviter sukatis estipulatis. Panicula 
terminalis, plus minusve longe pedunculata, pyramidalis, laxa, multi- 
flora, pedicellis squamulosis exceptis glabra, ramulis oppositis, in 
dichasia pluraque triflora exeuntibus, pedicellis gracilibus squamulosis 
bracteis bracteolisque deciduis instructis; alabastra ovoidea. 

Species unica Chinae provinciae Kweichou incola. 

Handeliodendron Bodinieri (Levi.), comb, no v. 

Character generis. 

Petiolus 4-11 cm. longus; foliola basalia terminali saepe duplo minora, 
3-12 cm. longa et 1.5-6.5 cm. lata, petiolulis 1-15 mm. longis; panicula 
pedunculo 3-5 cm. longo excluso circa 10 cm. longa et lata, pedicellis 
2-5 mm. longis; sepala 2-i mm. longa; petala 9 mm. longa et 2 mm. 
lata; stamina 5-9 mm. longa, antheris 0.75 mm. longis; ovarium stylo 
brevissimo incluso 1.25 mm. longum, stipite 1.5 mm. longo. Capsula 
stipite circiter 1 cm. longo incluso 3.2 cm. longa; semina circa 1 cm. 

China. Kweichou: district de Ly-po, /. Cavaterie in herb. 
Bodinier, no. 2626, Sept. 1898 (fruit). May 11, 1899 "grand arbre" 
(holotype of Sideroxylon Bodinieri in herb. Leveille, Bot. Card. 
Edinb.); Mapo, Pingchow, alt. 500 m., common in light woods, Y. 
Tsiang, no. 6813, Aug. 30, 1930, "tree, bark dark gray, branchlets lenti- 
cellate, fruit reddish, seeds black" (in Herb. Nat.-hist. Mus. Wien ex 
Herb. Metrop. Mus. Nat. Hist. Acad. Sin. Nanking). 

Though Handeliodendron resembles in its opposite digitately 5-folio- 
late leaves the Hippocastanaceae, it shows in its other characters a closer 
affinity with the Sapindaceae and is best placed with the tribe Harpul- 
lieae on account of the 2-ovuled locules, the symmetrical flower, the 
dehiscent fruit, the not spirally curved embryo and the presence of a 
terminal leaflet. The genus exhibits a number of characters unusual or 
rare in the family, as the opposite digitate leaves, flowers with 7 stamens, 
stipitate ovary and a unilateral disk, and seeds with a double arillus 
and straight embryo. Opposite leaves are very rare in the family, they 
occur in Valenzuela and some species of Matayba, digitately 5-foliolate 
leaves are still rarer and are only found in a few species of Allophylus 
as in /4. dimorphophyllus Radlk., though ternate leaves occur in a num- 
ber of genera as Delavaya, Hypelate, Llagunoa, Thouinia, and Allo- 
phylus; also the double arillus is very rare. The wood, but not the 
bark and other parts of the plant, contains saponin according to Dr. 
Handel-Mazzetti. 

The solitary flowering specimen I have seen has only the terminal 



Arnold Aku. Vol. XVI 




1935] 



REHDER, HANDELIODENDRON 



flowers of the dichasia at the end of the branches fully open, all the 
other flowers are in bud. The open flowers as well as the flowers still 
in bud seem to have normal ovaries and normal anthers; all the open 
flowers have 4 petals, while in at least some of the flowers in bud I 
counted 5 petals; in one flower I found 8 stamens. The description of 
the seeds is based on notes and on drawings kindly furnished me by Dr. 
Handel-Mazzetti, since I did not feel at liberty to dissect the solitary 
fruit of the type specimen. He had identified Tsiang's no. 6813 which 




is in fruit as belonging to the same species as Bodinier's flowering speci- 
men, after I had sent him a photograph of the type of Sideroxylon 
Bodinieri Levi, with a detached flower. A photograph of Tsiang no. 
6813 kindly sent me with a detached fruit by Dr. Handel-Mazzetti con- 
firmed the identity of the two specimens. 

I take pleasure in naming this interesting and distinct new genus in 
honor of Dr. H. Handel-Mazzetti, who has collected extensively for 
several years in China and whose critical account of the plants of his 
and other collections in his Symbolae Sinicae is one of the most im- 
portant contributions to our knowledge of the flora of China. 



JOURNAL OF THE ARNOLD ARBORETUM 



NOTES ON SOME OF THE EBENACEAE AND VERBENACEAE 
OF THE SOLOMON ISLANDS COLLECTED ON THE 
ARNOLD ARBORETUM EXPEDITION, 1930-1932^ 

R. C. Bakhuizen van den Brink 

IVith plates 120-122 

EBENACEAE 

Diospyros ellipticifolia (Stokes) Bakhuizen, Enum. Mai. Eben. in 

Card. Bull. Str. Settl. 7(2) : 162 (1933). 

Maba clliptica J. R. et G. Forster. Char. Gen. PI. 122, tah. 6 (1776). 
Y s a b e 1 Island: Tiratona, alt. 600 m., Brass 3318, $ , flor., 
Dec. 8, 1932. — Vernacular name "Gaitutunu." 

Diospyros ferrea (Willd.) Bakhuizen, Enum. Mai. Eben. in Gard. 
Bull. Str. Settl. 7(2): 162 (1933). 

Maba buxifolia (Rottb.) A. L. Jussieu in Ann. Mus. Hist. Nat. 5: 418 

Diospjrros ferrea var. salomonensis Bakhuizen, var. nova. 

Subsimilis D. ellipticifoliae, sed staminibus 9, foliis submajoribus 
differt. 

Ramuli teretes, rugosi, dense tuberculatim lenticellati. Folia ellip- 
tica vel oblongo-lanceolata, basi obtusa vel rotundata, apice obtusa vel 
breviter obtuse acuminata, 5-20 cm. longa, 3-7 cm. lata, chartacea vel 
tenuiter coriacea, supra atro-viridia, nitida vel statu sicco subopaca, 
subtus pallidora, primum subtus appresse pubescentia, denique costa 
excepta utrinque glabra, nervis lateralibus utrinsecus 7-10 vel pluribus, 
utrinque subinconspicuis, venis reticulatis laxis utrinque subprominulis 
invisibilibus; petiolus semiteres, appresse rufo-pubescens, glabrescens, 
0.3-0.5 cm. longus. Flores masculi sessiles, 3-5-ni, cymosi vel in 



iJn 




S F Kajew^ki in continuation of his botanical work 






>retum in the New Hebrides and North Queensland, 


left 


Brisbane for Bougainville 


Island and the British Solomon Islands. He resigned 


in J 


une 1931 and in July 1932 


his place was taken by Mr. L. J . Brass, who remained 


there until December 1932 w 


hen he returned to Brisbane, and after a few weeks 






the Archbold Expedition of the American Museum of 




iiral Historv. New York. 


Mr. C. T. White, Brisbane, to whom the collections 




ement, advises that they have now been roughly sorted 




) families. Several of thes< 




and 


we have already received c 




Dr. 




n Brink. The descriptions of the new species, new 












the general account of the Bougainville and Solomon 


Isla 


nds collection to be publis 


hed at a later date.-Ed. 



19351 BAKHUIZEN, EBENACEAE AND VERBENACEAE 69 

pseudo-racemis dispositi; calyx urceolatus, 3-dentatus, in anthesi saepius 
rumpens, intus versus apicem appresse rufo-pubescens; corolla albida, 
versus lobos purpurascens; stamina 9, sublibera, glabra. Fructus ellip- 
soideus vel subglobosus, primum appresse pubescens, maturitate gla- 
brescens, flavescens, 1.5-2 cm. longus, 1.5-1.75 cm. diam.; calyx fruc- 
tifer subpatelliformis, marginibus reflexus, extus sparse appresse pu- 
bescens, glabrescens, rugulosus, intus toto superficie sericeus, 1-1.2 cm. 
diam.; semina oblonga, triquetra, utrinque acuta, facie recta, dorso con- 
vexa, a latere compressa, rugulosa, nigra, 1.2-1.5 cm. longa, 0.5-0.7 
cm. lata, 0.5 cm. crassa; albumen aequabile. 

Ulawa Island: Brass 2958, 9, fruct., Oct. 5, 1932. 
Ysabel Island: Jaukau, Brass 3152, S , flor., Nov. 19, 1932. — 
Vernacular names "Aibul" (under no. 2958) and "Gno-gno-finete" 
(underno. 3152). 

Diospyros insularis Bakhuizen. Enum. Mai. Eben. in Gard. Bull. 
Str. Settl. 7(2): 173 (1933). Plate 120, 121 

.Arbuscula. Folia elliptico-oblonga, basi cordata. apice breviter ob- 
tuse acuminata, supra atro-viridia, statu sicco olivacea, subopaca, nervis 
lateralibus 7-12 distantibus, secus marginem non vel indistincte anas- 
tomosantibus, utrinque prominentibus; petiolus subteres vel apicem 
versus subdepressus. Fructus axillaris, sessilis, plerumque solitarius, 
ellipsoideus vel subglobosus, utrinque rotundatus, primum sericeus. 
statu maturo glabratus, ruber, statu sicco niger, rugulosus, opacus. 2-2.5 
cm. diam.; calyx fructifer viridis, valde accrescens, extus glaber, 3.5-4 
cm. diam., tubo crasso, plano-cupuliforme, subquadrato, intus rufo- 
sericeo, rimo elevato, 2-2.5 cm. diam.. lobis late ovatis obtusis vel sub- 
orbicularibus, coriaceis, patulo-recurvis vel reflexis, striato-nervosis, 
utrinque glabris, 1-1.5 cm. longis, basi 1.5-1.75 cm. latis; semina usque 
ad 8, oblongo-ellipsoidea, triquetra, dorso convexa, a latere compressa, 
ca. 1.5 cm. longa, 0.7 cm. lata, 0.4-0.5 cm. crassa; testa rugulosa. nigra: 
albumen aequabile. 

Ysabel Island: Maringe, Brass 3166, 9 , fruct.. Nov. 22, 
1932. 

Diospyros maritima Blume, Bijdr. Flor. Ned. Ind. 669 (1825). 

Ysabel Island: Sigana, alt. 100 m., Brass 3450, 9 , fruct., 
Jan. 11, 1933. — Vernacular name "Gegila." 

Diospyros samoensis A. Gray in Proc. Amer. Acad. 5: 326 (1861). 

San Christoval Island: Star Harbor, .Brow 3073, '^ 
fior., Oct. 18, 1932. Ngela Group: Xayotana Island, Brass 



JOURNAL OF THE ARNOLD ARBORETUM 



VERBENACEAE 




Avicennia marina (Forsk.) Vierhapper, Beitr. Kennt. Flor 


. Sud- 


rab. in Denkschr. Akad. Wiss. Wien, 71: 435 (1907). 




Avicennia marina var. resinifera (Forst.) Bakhuizen, Rev 


. Gen. 


vie. in Bull. Jard. Bot. Buitenz., ser. 3(2): 210, tab. 16 (1921) 




Malaita Island: Quoimonapu, sea level, Kajewski 


2344, 


lec. 11, 1930. — Vernacular name "Bu-bula." 





CaJlicarpa pedunculata R. Brown, Trod. Flor. Nov. HoU. 513 
(1810). 

Guadalcanal Island: Berande River, sea level, Kajew- 
ski 2420, Jan. 7, 1930. — Vernacular name "Bau." 

Callicarpa pentandra Roxb., Flor. Ind. 1: 409 (1820) var. typica 
(Schau.) Bakhuizen, forma genuina Bakhuizen in H. J. Lam. & Bak- 
huizen, Rev. Verb, in Bull. Jard. Bot. Buitenz., ser. 3, 3(1) : 12 (1921). 

Bougainville Island: Kieta, sea level, Kajewski 1560, 
March 21, 1930; Kupei Gold Field, alt. 950 m., Kajewski 1643, April 
7, 1930; Kugu-maru, Buin, alt. 150 m., Kajewski 1841, June 9, 1930. 
San Cristoval Island: Waimamura, alt. 200 m.. Brass 
2625, August 10, 1932. — Vernacular name "Sor-ku-ku" (under no. 
1841). 

Callicarpa pentandra var. paloensis (Elm.) Bakhuizen, forma 
furfuracea Bakhuizen in H. J. Lam & Bakhuizen, Rev. Verb, in Bull. 
Jard. Bot. Buitenz., ser. 3, 30: 15 (1921). 

Malaita Island: Quoimonapu, sea level, Kajewski 2340, 
Dec. 11, 1930. Guadalcanal Island: Ma-massa, Konga, 
alt. 400 m., Kajewski 2485, Feb. 12, 1931; Vulolo, Tutuve Mt., alt. 1200 
m., Kajewski 2540, April 20, 1931. — Vernacular names, "Quoi-esa" 
(under no. 2340), "Kim-berri" (under no. 2485) and "Kimberi" (under 
no. 2540). 

Clerodendron Buchanani (Roxb.) Walpers, Rep. Bot. Syst. 4: 108 
(1845). 

Bougainville Island: Kajewski 1606, March 29, 1930; 
Karngu, Buin, sea level, Kajewski 2222, Oct. 6, 1930. San C r i s - 
toval Island: Waimamura, 5rfl5^ 3140, Oct. 1932. Ysabel 
Island : Sigana, alt. 20 m., Brass 3465, Jan. 13, 1933. — Vernacu- 
lar name "Arka-koo" (under no. 2222). 

Meded. Rijks Herb. Leiden, 



1935] BAKHUIZEN, EBENACEAE AND VERBENACEAE 71 

Bougainville Island: Kupei Gold Field, alt. 900 m, 
Kajewski 1687, April 11, 1930; Kugu-maru, Buin, alt. 150 m., Kajew- 
ski 1925, August 4, 1930; same locality, Kajewski 1978, August 22>, 
1930. Malaita Island: Quoi-mon-apu, sea level, Kajewski 2341, 
Dec. 11, 1930. Guadalcanal Island : Vulolo, Tutuve Mt., 
alt. 1200 m., Kajewski 2502, April 14, 1931. San C r i s t o v a 1 
Island: Hinuahaoro, alt. 900 m., Brass 2919, Sept. 22, 1932. 
Ysabel Island: Tiratofia, alt. 600 m.. Brass 3403, Dec. 29, 
1932. — Vernacular names "Koru-kopu" (under no. 1925), "E-ya- 
papor" (under no. 1978), "Kaka-fair" (under no. 2341), "Ambus-gor- 
le-Ie" (under no. 2502) and "Fuho" (under no. 3403). 

This species is closely related to C buruanum Miq. which differs in 
the much longer corolla-tube and also to C. injortunatum L., which has 
a glabrous corolla and a longer corolla-tube. Nevertheless all these 
species may perhaps be considered as only extreme forms of C. injor- 
tunatum L. 

Clerodendron inerme (L.) Gaertner, Fruct. Sem. Plant. 1:271, tab. 
75 (1788). 

Bougainville Island: Karngu, Buin, sea level, Kajew- 
ski 2244, Oct. 12, 1930. Guadalcanal Island: Berande, 
sea level, Kajewski 2407, Jan. 5, 1931. — Vernacular names "Pumb- 
arg-aru" (under no. 2244) and "A-la-loi-alugi" (under no. 2407). 

Paradaya amicorum (Seem.) Seemann in Jour. Bot. 3: 258 (1865). 
Clerodendron amicorum Seemann in Bonplandia, 10:249 (1862). 

Faradaya amicorum var. salomonensis Bakhiuzen, var. nova. 

Frutex flexuosus plerumque scandens, primum appresse pubescens, 
denique glabrescens, in ramulis florigeris cinereo-subsericeus. Folia 
valde variabilia, lanceolata-oblonga vel obovata, utrinque 
basi acute vel obtuse cuneata, apice breviter subacute 
utrinque glabra, 7-20 cm. longa, 3.5-10 cm. lata, nervis lateralibus dis- 
tantibus, utrinsecus 5-7. Inflorescentiae axillares vel in paniculis speci 
osis terminalibus dispositae, multifiorae, cymis subinde trichotomis, basi 
conspicue bracteolatis; bracteolae oblongo-ellipticae vel sublanceolatae. 
utrinque sericeae, 1-2.5 cm. longae, 0.3-1 cm. latae. Flores subparvi. 
in alabastro globosi, pedicellati; pedicelli teretes, graciles, cinereo- 
sericei, basi bracteolati, 0.5-1.5 cm. longi; calyx primum subclausus vel 
apice poro dehiscens, denique truncatus vel margine undatus, vel den- 
tatus vel etiam distincte lobatus, 0.5-0.6 cm. longus, 0.7-1 cm. diam., 
fructifer accrescens, saepius irregulariter ruptus, extus sparse pubescens, 
basi excepta glabrescens; corolla alba, hypocrateriformis, utrinque 



72 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

glabra, tubo variabili, 1-1.5 cm. longo, lobis ovatis vel suborbicularibus, 
glabris, margine cillatis; stamina longe exserta, glabra; ovarium quad- 
rangulare vel 4-lobatum, glabrum; stylus filiformis, teres, glaber, usque 
ad 3 cm. longus. Fructus submagnus, abortu 1-pyrenus; pyrena elongata, 
nucleis reductis appendiculiformibus basi suffulta, monosperma, glabra, 
3-4 cm. longa, 1.5-2 cm. diam. 

San Cristoval Island: Waimamura, alt. 50 m., 5rfl55 
2635, August 11, 1932. Ysabel Island: Tiratona, alt. 600 m., 
Brass 3399, Dec. 29, 1932. — Vernacular name "Naosokono." 

Perhaps this is not really different from the typical form, but it has a 
glabrous corolla. 

Gmelina moluccana (Bl.) Backer in Heyne, Nutt. Plant. Ned. Ind. 
4: 118 (1917); Bakhuizen in H. J. Lam & Bakhuizen, Rev. Verb, in 
Bull. Jard. Bot. Buitenz.. ser. 3, 3( 1 ) : 67 ( 192 1 ) . 

San Cristoval Island: Waimamura, sea level, J5rc55 
2860, Sept. 12, 1932. 

Gmelina salomonensis Bakhuizen, spec. nova. Plate 122 

Arbor speciosa. Ramuli crassi, teretes, novelli rufo-tomentosi, vetu- 
stiores glabrescentes, sparse lenticellati. Folia opposita, longe petio- 
lata, coriacea, ovata vel oblongo-elliptica, basi cordata vel subtruncata, 
apice acuminata, obtusa, integerrima, 15-35 cm. longa, 10-24 cm. lata, 
supra viridia, lucida, primum sparse pubescentia, denique costa et 
nervis exceptis glabra, subtus grisea, submoUiter rufo-tomentosa, basi 
nonnuUis glandulis parvis obsessa, costa supra leviter subtus valde 
prominente. utrinque glabrescente, nervis lateralibus utrinsecus 12-18 
pallidis, supra prominulis glabris, subtus prominentibus rufo-tomentosis. 
venis reticulatis utrinque prominulis. Inflorescentiae terminales, 
elongatae, paniculiformes, dense ramosae, infra foliatae, rufo-tomento- 
sae, bracteolatae, dense multiflorae, 20-30 cm. longae, 10-15 cm. diam.; 
bracteae parvae, lineari-oblongae, utrinque acuminatae acutae, utrinque 
tomentosae, mox deciduae. Flores minores, pedicellati; calyx cupuli- 
formis, regulariter obtuseque 5-dentatus, extus rufo-tomentellus, glan- 
dulis 2-4 parvis vestitus, intus glaber, 0.3-0.4 cm. longus et diam., fruc- 
tifer subaccrescens; corolla minor, inaequaliter 5-lobata, subbilabiata, 
utrinque pubescens, statu sicco ca. 1.5 cm. longa, tubo inferne angustato 
in faucem ventricosam ampliato, intus ad insertionem staminum longe 
hirsute, superne glabrato, calyce multo longiore, 0.7-1 cm. longo; 
stamina 4, didynamia, vix exserta, statu sicco 0.5-1 cm. longa, fila- 
mentis glabris; ovarium subglobosum vel obovoideum, apice subtrunca- 
tum, glaberrimum; stylus filiformis, teres, sparse pilosus, vix exsertus. 



1935] BAKHUIZEN, EBENACEAE AND VERBENACEAE 73 

Statu sicco ca. 1 cm. longus. Drupa minora, ovoideo-globosa, nitida, 
maturitate nigra, statu sicco 1.2-1.5 cm. diam.; calyx fructifer leviter 
excrescens, subapplanatus vel marginibus ref!exis, quinatus, 0.5-0.7 

Ysabel Island: Tiratofia, alt. 600 m., Brass 3309, Dec. 8, 
1932. — Vernacular name "Koko." 

This plant is intermediate between G. moluccana (Bl.) Backer and 
G. macrophylla (R. Br.) Benth. and may be a hybrid of these species. 
From G. moluccana it differs in the tomentose under side of the leaves 
and the villous calyx; from G. macrophylla in the terete branches, the 
elevated nerves and veins on the upper side of leaves, somewhat in the 
form of panicles, but especially in the small and regular 5 -toothed calyx. 

Petraeovitex multiflora (Sm.) Merr. var. salomonensis Bakhuizen, 
var. nova. 

Frutex scandens, gracilis; ramuli quadrangulares, primum tomentelli, 
glabrescentes. Folia opposita, ternata vel inaequaliter biternata; foliola 
3-9, minora, sessilia, chartacea; foliola lateralia ovato-oblonga vel 
oblongo-elliptica, basi rotundata, apice obtuse acuminata, 1-4.5 cm. 
longa, 0.5-2.3 cm. lata, foliolum terminate oblongo-lanceolatum, 
utrinque attenuatum, basi decurrens, acute acuminatum, apice obtusi- 
uscule acuminatum, 3.5-6 cm. longa, 1.5-2.5 cm. lata, omnia margine 
Integra, supra glabra, subtus primo farinaceo-tomentosa, denique gla- 
brescentia vel costa nervisque excepta glabra, nervis lateralibus 6-8 cum 
venis reticulatis utrinque prominulis ; petiolus communis 2-i cm. longus, 
petiolus lateralis 0.5-1 cm. longus, omnes superne canaliculati, cinereo- 
tomentelli. Inflorescentiae terminates, laxe paniculiformes, infra foli- 
atae, multiflorae, 30-50 cm. longae, 20-40 cm. diam.; bracteae minutae 
lineares, tomentellae, 0.15-0.25 cm. longae; cymulae breviter peduncu- 
latae vel superne subsessiles, 7-15-f!orae, tomentellae, 0.5-1 cm. diam., 
pedunculis 0-1 cm. longis. Flores parvi, subsessiles, albidi; calyx 5- 
dentatus, cinereo-tomentellus, 0.1-0.15 cm. diam.; corolla alba, extus 
minute pubescens vel glabrescens, in fauce albido-puberula, 5-lobata, 
lobis inaequalibus, reflexis; stamina 4, exserta, fauci inserta, glabra; 
stylus gracilis, exsertus, glaber, 0.3 cm. longus, stigmatibus bifidis; 
ovarium ovoideum, basi glabrum, apice cinereo-tomentellum. 

Bougainville Island: Kupei Gold Field, alt. 850 m., 
Kajewski 1686, April 11, 1930. 

This variety much resembles P. sumatrana H. J. Lam, but it has 
sessile leaflets, a cinereous-tomentose calyx and a hairy ovary. 

Premna integrifolia Linnaeus, Mant. 2: 252 (1771) s. 1. 



74 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

Bougainville Island: Kieta, sea level, Kajewski 1566, 
March 30, 1930; Kugu-maru, Buin, alt. 150 m., Kajewski 1842. 
Malaita Island: Quoimonapu, alt. 50 m., Kajewski 2330, 
Dec. 10, 1930. Guadalcanal Island: Vulolo, Tutuve Mt., 
alt. 1200 m., Kajewski 2503, April 14, 1931. San Cristoval 
Island: Waimamura, alt. 200 m., Brass 2624, August 10, 1932 (f. 
taitensis Schau.); Kirakira, Brass 2768, August 30, 1932; Star Har- 
bour, Brass 3132, Oct. 1932. — Vernacular names "Garlu" (under no. 
1842), "Qua-eu" (under no. 2330) and "Arru-arru" (under no. 2503). 

Teysmanniodendron Ahernianum (Merr.) Bakhuizen, comb. nov. 
Viiex Ahcrniana Merrill in Bur. Gov. Lab. Bull. 6: 18 (1903). 

A large sized tree, up to 50 m. high; bark grey or brown; v^^ood hard, 
brown; branchlets round, greyish, rufous-pubescent, glabrescent. Leaves 
2-3-foliolate, short petioled; petiole terete, rufous-tomentose, especially 
at the base and in the insertion of the petiolules, 4-5 cm. long; petiolules 
in all leaves equal, furrowed above, thickened and rufous-tomentose at 
the base only, otherwise glabrous, 2-4 cm. long. Leaflets oblong, coria- 
ceous, rather rigid, shining above, glabrous on both surfaces, except 
pubescent on the midrib beneath when young, 10-32 by 4-13 cm., 
reticulations of leaves very dense beneath. Cymes axillary, many- 
flowered, 15-30 cm. long; peduncles 1-2 in the axils, flattened, rufous- 
tomentose, 5-12 cm. long. Flowers rather small; calyx funnel-shaped, 
obscurely 5-dentate, rufous-sericeous, 0.4-0.5 cm. long, and wide; corolla 
with very short tube, glabrous at the base, otherwise sericeous, throat 
and base of the lip densely villous ; ovary globose, glabrous, biloculate. 
Fruit oblong or pear-shaped, purple green when ripe, shining and striate 
when dry, 1.5-2 cm. long, 1-1.5 cm. diam., one-seeded; exocarp coria- 
ceous, thin; seed oblong, 1 cm. long, 0.5-0.7 cm. diam.; fruiting calyx 
enlarged, cup-shaped, truncate, 0.5 cm. long, 0.5-0.8 cm. in diam. 

Ysabel Island: Tataba, alt. 50 m. Brass 3441, Jan. 5, 1933. 
Guadalcanal Island: Sorvorhio Basin, alt. 200 m., Kajew- 
ski 2715, Feb. 3, 1932. — Vernacular name "Seupa" (under no. 2715). 

Vitex cofassus Reinwardt ex Blume, Bijdr. Flor. Ned. Ind. 813 
(1826). 

Bougainville Island: Kieta, sea level, Kajewski 1533, 
March 17, 1930; Kugu-maru, Buin, alt. 150 m., Kajewski 1843, May 
28, 1930. Malaita Island: Quoimonapu, alt. 300 m., Kajew- 
ski 2381, Dec. 16, 1930. Guadalcanal Island: Berande 
River, Kajewski 2387, Dec. 24, 1930; Mamassa, Konga, alt. 400 m., 
Kajewski 2489, Feb. 13, 1931; Vulolo, Tutuve Mt., alt. 1200 m., Ka- 
jewski 2605, May 1, 1931. San Cristoval Island: Balego- 




DiOSl-YKOS INSULAUIS B: 



(D-O 



Q, 



:y 



It 



■^ 







Jour. Arnold Arb. Vol. XVI. 




1935] BAKHUIZEN, EBENACEAE AND VERBENACEAE 75 

Nagonago, alt. 350 m., Brass 2821, Sept. 5, 1932. Ysabel Island: 
Maringe, Bras :i 3154, Nov. 19, 1932 ; Tasia, Brass 3272, Dec. 5, 1932. — 
Vernacular names "Moi-kewie" (under no. 1843), "Father" (under no. 
2381), '.'Vada" (under no. 2387), "Vatha" (under no. 2489), "Vasa" 
(under no. 2605), "Hada" (under no. 2821), "Wara" (under no. 3154) 
and "Varha" (under no. 3272). 

EXPLANATION OF PLATES 
Plate 120 
Diospyros insiilaris Bakh. (type G. Peekel 4 A from New Ireland, 
S). — A. Flowering branch (X 3/2). — B and C. Leaf base seen from 
above and from below (X 1>4). — D. Inflorescence, flowers dropped 
(X2). — E-L. Male flowers with details: E-F. flowers from the inside 
and the outside (X2y2) ; G. longitudinal section of flower, seen from the 
inside (X 2y:') ; H. calyx, seen from the inside (X 2>^) ; I. rudimentary 
ovary (X 10) ; K. androeceum (X ZYz) ; L. stamens ( X 5). 

Plate 121 
Diospyros insiilaris Bakh. (type Brass 3166 from the Solomon Islands, 
9). — A. Leafy branch (X >4). — B. Fruiting branch (X yi). — 

C. Axis of the fruiting branch (X 2^4). — D. Peduncle of fruit at the 
apex (X 13^). — E-F. Fruiting calyx (X 5^): E. from the inside; 
F. from the outside. — G-H. Seeds (X 1>^) : H. seed in transverse 

Plate 122 

Diospyros samoensis A. Gray — A-B. Flowering branches of male 

plant (X ^). — C-F. Male flower bud with details : C. corolla bud ( X 3) : 

D. corolla bud cut lengthwise (X 3); E. stamens (X 5); F. anthers 
(X 10). — G. Flowering branch of female plant (X 3^). — H-N. Fe- 
male flower with details : H. flower buds from above and from below ( X 
2) ; I. calyx ( X 2) and calyx-lobe, seen from the inside ( X 2^) ; J- corolla 
bud (X 3); K. corolla bud cut lengthwise (X 3) ; L. ovary (X 3): 
M. anthers (X 15) ; N. ovary in transverse section (X 5). — R. Fruiting 
branch (X y^). — S-T. Fruiting calyx from the outside and from the 
inside (X 1 ) ; T. lobe of fruiting calyx from the outside (X 1 3^). — U. 
Seeds ( X 1>^). — V. Seed in cross section ( X 2). 



JOURNAL OF THE ARNOLD ARBORETUM 



AN Ex\DEMIC SOPHORA FROM RUMANIA 



//'///) plates 12S and 124 and one text figure 

One of the most interesting endemics of the Balkan peninsula is the 
Sophora discovered at Babadag by J. Prodan.' Through the kindness 
to visit this locality on September 4, 1934 and collected abundant fruit- 
of Dr. C. Georgescu of the Scoala Politechnica at Bucarest I was able 
ing material. Subsequent comparison with Asiatic material of Sophora 
alopccuroides L. in the herbaria of the Royal Botanic Garden at Kew 
and of the Arnold Arboretum has convinced me that the Rumanian plant 
deserves to be described as a distinct species and I take pleasure in nam- 
ing it after its discoverer. 

Sophora Prodanii, sp. nov. 

Herba suffruticosa, 5-7 dm. alta. Folia 5-10 cm. longa, imparipin- 
nata; foliola 19-25, oblongo-elliptica, 12 mm. longa, 7 mm. lata, mem- 
branacea, supra glabra, subtus pilos sparsos appressos gerentia. Race- 
mus densus. Flores ignoti, non visi. Legumen 5-7 cm. longum, gla- 
brescens; semina 3-7, luteo-fusca, 5 mm. longa. 

Known only from a single hilltop near Babadag, Rumania. 

Seremet, Babadag, Rumania, Edgar Anderson no. 85 (type), Sept. 
4, 1934 (specimens deposited in the herbaria of the Arnold Arboretum, 
Royal Botanic Garden Kew and British Museum of Natural History). 

An erect suffrutescent herb from an underground rootstalk; stems, 
erect, 5-7 dm. tall, slender, with ascending simple branches, subterete, 
dark green with fine, rather scattered, short, appressed hairs. Leaves 
alternate, imparipinnate, 5-10 cm. long; stipules wanting; leaflets 19- 
25, elliptic oblong to oblanceolate, up to 12 mm. long and 7 mm. wide 
when well developed, dark green, rather thin, becoming brittle when 
dry; apex rounded with a mucronate tip. Leaflets glabrous above, pu- 
bescent below with very scattered fine short appressed hairs; margin 
entire and somewhat revolute; midrib evident but veins weak and 
evident only beneath; petiolule about 1 mm. long. Inflorescence ter- 
minal, racemose, dense, sub-erect. Flowers not seen. Fruit terete, 
torose, wingless, indistinctly ribbed, indehiscent or tardily dehiscent, 

'Map. Bot. Lapok. 11: 231, 235 (1012). 



1935] ANDERSON, A SOPHORA FROM RUMANIA 77 

with sparse appressed hairs; pedicels in fruit 2-4 mm. long, strictly 
ascending. Seeds yellowish brown, 5 mm. long. 

The species is of very restricted distribution. It is at present known 
only from this one locality, the summit of a small hill near the ancient 
town of Babadag. It occurs over a space of several acres in the edge of 
the forest and persists as a weed in an adjoining field. Prodan (loc. cit.) 
in his account of the plant from Babadag identified it with S. alope- 
curoides L. but pointed out that it was much more nearly glabrous. 



S, 






ibiu_. ^, ,-:_„,:ZSse,c^i? 



® = Sophora Prodanii 

* — S. alopecuroides 

O = S. alopecuroides var. tomentosa 

♦ = Intermediate form collected by Gilliat Smith 

Sophora alopecuroides, sensu latiore, is a wide-spread species (see Fig. 
1), extending from central Asia to northern Asia Minor and the vicinity 
of Constantinople (Istambul). From central Asia to Asia Minor there 
is a progressive transition in pubescence, leaf size, and leaf texture. If 
only the two ends of the series existed they could easily be maintained 
as two separate species, a small-leaved species with appressed silky hairs 
from northern Asia Minor and a coarser species with spreading tomen- 



78 JOLRNAL OF THE ARNOLD ARBORETUM [vol. xvi 

tose pubescence from Central Asia. As early as 1850 Spach' had pro- 
posed the name 5. Jauberti for the Sophora from Asia Minor and in 1894 
Freyn and Sintenis- described Goebelia reticulata from northern Asia 
Minor, a name which was later transferred to Sophora by Hayek.^ Azna- 
vour* went so far as to advance the plants collected by him in the sub- 
urbs of Constantinople to the status of a variety, Buxbaumii of Goebelia 
reticulata. 

It is certainly true that there is marked geographical differentiation in 
5. alopecuroides, sensu latiore, but when a large series of specimens is 
examined, these local and regional differences are found to intergrade. 
Particularly interesting are two collections made by Mr. B. Gilliat Smith 
(nos. 1904 and 1714) in the neighborhood of Tabriz, Persia, which can 
be assigned with certainty neither to the form from Central Asia nor to 
that from Asia Minor. Since Tabriz is in the region where these two 
forms come together, it seems best to follow Boissier'' and Bornmiiller" 
and treat the Sophora from Asia Minor as S. alopecuroides L. and that 
from Central Asia as S. alopecuroides L. var. tomcntosa (Boiss.) Bornm. 
Further study will undoubtedly permit the separation of other geo- 
graphical varieties. The specimens I have seen from N. W. China 
which have been referred to S. alopecuroides are certainly different from 
those collected in Afghanistan and Persia. 

Taken as a whole, 5. alopecuroides and 5. Prodanii present a graded 
series in size and texture of leaflet, pubescence and color (Table I). 

.S". Prodanii S. alopecuroides S. alopecuroides 



upper side of appressed sil 

leaflets glabrous pubescence pubescei 

under side of scattered appressed appressed silky spreadir 

leaflets hairs pubescence pubescei 



*Magyar Bot. Lapok, 12: 
5F1. Or. 2: 628-620 (1872^ 
6Bot. Cent. Beih. 27: 347 



1935] ANDERSON, A SOPHORA FROM 

There is no more difference, if as much, between 5. Prodanii and S. alope- 
curoides from the neighborhood of the Bosphorus as between 5. alope- 
curoides from Asia Minor and S. alopecuroides var. tomentosa from 
Afghanistan. But in the latter case, there is a full set of intermediate 
forms from the intervening territory while in the former the inter- 
mediates which once undoubtedly existed have long since disappeared. 
In the 250 miles between Babadag and the Bosphorus no sophoras of 
this group have been collected. The differences between 5. alope- 
curoides and 5. Prodanii, though slight, include leaf texture and color 
as well as pubescence and general size. For this reason 5. Prodanii is put 
forward as a distinct species rather than as a variety of S. alopecuroides. 

Sophora Prodanii undoubtedly originated as a semi-glabrous variety 
on the westward edge of S. alopecuroides. A large number of Balkan 
species represent westward extensions of Asiatic species, or find their 
closest relatives in the Asiatic flora. "It is necessary to bear in mind 
. . . that the Hungarian and Roumanian plains were covered with the 
waters of the Sarmatic and Pontic seas and lakes until relatively recent 
geological times (and) that the Bosphorus is no wider than a broad river 
. . . It follows . . . that migration on a wide front between the low- 
lands of the Balkan Peninsula and those of the north has been possible 
for land plants only since the end of the Tertiary period and must for 
the most part have been in one direction — northwards — as the Sarmatic 
and Pontic waters dried up; that migration along the northern part of 
Asia Minor into the Balkan Peninsula is geographically feasible and 
has been even more so in past geological periods."^ 

The persistence of 5. Prodanii in this one isolated station in the 
Dobrudja is to be explained by the geological history of the Babadag 
mountains. These low mountains (or hills) are of very great age and 
though low in elevation have persisted for a long time as a land mass, 
remaining above the waters of the Sarmatic and Pontic seas and lakes. 
"It is safe to assume that they formed a refuge for relatively old types 
of plants and to this fact is due the richness of the Dobrudja in Tertiary 
relicts." (Turrill.loc.cit.) 

To the question as to whether S. Prodanii evolved its distinctive char- 
acteristics before or after its separation from the sophoras of Asia Minor, 
the present day differentiation within the latter suggests an answer. Not 
only is there a progressive reduction westwards in size and pubescence 
from Central Asia to the Bosphorus but the same tendency can be seen 
within Asia Minor itself. The specimens of 5. alopecuroides which most 
closely resemble S. Prodanii are from northwestern Asia Minor. These 
iTurrill, W. B. The Plant Life of the Balkan Pen. Oxford. 1929. 



80 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

facts suggest that in Miocene times 5. Prodanii was already a well 
marked variety of S. alopecuroides. The Sarmatic and Pontic waters 
(Upper Miocene or Pliocene) destroyed the intervening intermediates 
and reduced 5. Prodanii to a dwindling remnant in the Babadag Moun- 
tains. Within the immediate past at any rate, it has been so reduced in 
numbers as to undergo severe inbreeding and further divergencies from 
the parental type would be expected to have accumulated through the 
random effects of inbreeding on a small population. 

Most of the plants at Babadag seemed to be infected with some gall- 
producing organism. The characteristic "witches brooms" produced in 
this way are very conspicuous in the photograph of the type specimen. 
Similar growths are apparently common in Sophora alopecuroides. Dr. 
W. B. Turrill has very kindly supplied me with the following list of 
specimens in the Kew Herbarium which exhibit the phenomenon: Nes- 
torian Mountains and Gawan, Capt. Garden in 1857; Caucasus, Prescot 
in 1828; Near Tabriz, Persia, Gilliat-Smith in 1926; Pamir and Thian 
Shan Journey, H. Appleton 190 in 1906. 

In Babadag, the seeds of 5. Prodanii were reported to be extremely 
poisonous. While S. alopecuroides has never been listed as poisonous 
so far as I know, there are a number of references to the poisonous seeds 
of other species of Sophora. The seeds of 5. secundijlora Lag. are used 
by Mexican Indians as an intoxicant; one seed is said to be sufficient to 
kill a man and a half a seed produces a stupor lasting two to three days.^ 
S. flavescens Ait. contains poisons which are made use of as insecticides.' 
The seeds of S. tomentosa L. yield a poisonous alkaloid. They are a 
common native remedy in the Philippines for stomach disorders.' 

EXPLANATION OF THE PLATES 



Leaflet of Sophora alopecuroides var. tomentosa (X 7). From Stapf 
s. n., collected at Schiraz, Persia, Aug. 23, 1885. 
Leaflet of Sophora Prodanii ( X 7), From Anderson, no. 85 (type). 



^KewBull. 1892: 216-21; 




SoPHORA Prodanii E. Anders. 



19351 PALMER, SPONTANEOUS FLORA OF ARNOLD ARBORETUM 81 



Since the publication in 1930 of the list of plants growing spon- 
taneously in the Arboretum^ observation and collecting has been con- 
tinued, and as a result so many additions have been made to the flora 
that it now seems desirable to publish a supplementary list. 

In 1931 I had an opportunity for the first time to remain at the 
Arboretum throughout the spring months and to make a thorough ex- 
ploration of the native and introduced plants at that season. As a 
result of this and of subsequent investigation a large number of plants 
not recorded in the first list have been found and additional information 
about some of the rarer species previously recorded has also been 
secured. 

Of the 173 new species and varieties reported in this supplement 94, 
or nearly 55 percent, are plants native in the Boston area and presum- 
ably in the Arboretum, and the remaining 79, or 45 percent, are intro- 
duced. Seven plant families and 43 genera are added to the spon- 
taneous flora in this supplementary list. 

The grasses, sedges, and composites, as might be expected, furnish 
the largest number of additions. The really surprising thing about the 
present list is the relatively large number of native plants that have been 
found. Many of these are now quite scarce or rare in the Arboretum. 
Several of the introduced plants are probably recent introductions. 
Amongst the more interesting discoveries are: Polystichum acrosti 
choides, Aristida dkhotoma, A. gracilis, Carex communis, C. Goode- 
nowii, C. longirostris, Erythronium americanum, Luzula nemorosa 
Cypripedium acaule, XQuercus Rehderi, Anemone quinquejolia, Poten- 
tilla canadensis var. villosissima, Polygala sanguinea, Lechea intermedia 
L. tenuijolia, Viola pedata var. lineariloba, V. sagittata, Pyrola ameri- 
cana, Trientalis americana, Epifagus virginiana, Houstonia caerulea 
Liatris scariosa, Aster acuminatus, Helenium nudiflorum, Senecio aureus 
Hypochaeris radicata, Sonckus arvensis var. glabrescens, Hieracium 
florentinum, and H. vidgatum. 

A single weak plant of the Christmas fern was discovered in 1931 
by Dr. Grant D. Darker on a wooded slope of the North Woods, 
where it had probably survived from a native colony. Later, sev- 



82 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

eral plants were found on the north side of Hemlock Hill. At a 
little lower level in the latter locality "a small group of the stemless 
lady's slipper was found growing under the hemlock and pine trees. The 
star flower is also found here, as well as on the top of the hill and as a 
greater rarity along the base of a gravelly ridge in the North Woods. 
The little bluets or innocence grows sparingly among the laurels and 
other shrubs at the foot of the hill, and a little higher up is found the 
round-leaved wintergreen. As the hemlock grove on this rocky elevation 
is probably the only bit of practically virgin timber left in the Arbore- 
tum and as the native flora has been little disturbed even in a few spots 
along the base of the hill where there is an accumulation of richer soil, 
a number of interesting plants have been able to survive here that at 
present are not found elsewhere. 

The March lily has not yet been seen flowering in the Arboretum, but 
a small colony of sterile plants comes up each year in a moist shady 
spot on the south side of Hemlock Hill. The plants apparently lack 
sufficient vitality to produce blossoms, probably due to the increased 
shade. A colony of plants blooming freely is found just outside the 
Arboretum area on a wooded bank in the grounds of the Adams Nervine 
Hospital, and only a few yards from the division fence. The wood 
anemone survives sparingly at the base of wooded hills and even on 
open banks that were cleared of native trees only in recent years. It 
has been found at the edge of the North Woods, on the slopes of Bussey 
Hill in the oak group, along a bank of Bussey Brook below the junipers, 
and in the South Woods near Peters Hill. The purple milkwort grows 
on an open grassy bank on the east side of Peters Hill, and near the same 
spot a single plant of the bird-foot violet was found. Several plants of 
the latter were also found in open rocky woods on the top of Hemlock 
Hill, but they are likely soon to be exterminated by careless picking and 
trampling. A specimen of the beech-drops was collected on the south 
side of Hemlock Hill, but it has not been observed elsewhere in the 
Arboretum. The hairy bush-clover and the pinweed {Lechea tenui- 
folia) grow together in dry gravelly or rocky soil at the edge of the 
Central Woods, on the north slope of Bussey Hill, and near a small aban- 
doned quarry in the South Woods. Lechea intermedia is also found near 
the top of Peters Hill. 

Amongst the native woody plants that have been added to the list the 
low juniper is one of the rarest. Two or three small plants of this are 
growing on conglomerate outcrops in the Central Woods, where they are 
probably indigenous. The scrub chestnut oak grows sparingly near the 
same spot as well as on top of Hemlock Hill, where a single plant was 
seen. At this locality in the Central Woods was also found the inter- 



1935] PALMER, SPONTANEOUS FLORA OF ARNOLD ARBORETUM 83 

esting natural hybrid between the bear oak and the black oak {Quercus 
Rekderi) growing with both of the parent species. The choke berry is 
another native shrub found in the Central Woods and a few plants of 
it have survived the repeated mowings of the grass on the northeast 
slope of Peters Hill. The spice-bush and high-bush blueberry grow in 
lower and richer ground at a few places on the borders of the woods. 

It is gratifying to find evidence that a second species of thorn was 
native in the Arboretum. Specimens of Crataegus rotundijolia collected 
many years ago by Mr. C. E. Faxon and others in Bussey Wood and on 
Peters Hill were found in the herbarium, and sprouts of this species were 
found to be still growing at the latter locality. Mr. Faxon also col- 
lected specimens of the native blackberries and other plants in the 
Arboretum, some of which are preserved in this herbarium and others 
at the Gray Herbarium or in that of the New England Botanical Club. 

It is interesting to note the introduction of new plants appearing 
spontaneously in the Arboretum and how they succeed and spread or 
fail to establish themselves and disappear after a single season or in a 
few years. Before the publication of the original list a few leaves of 
what appeared to be a sterile plant of Senecio aureus were seen between 
the Linden group and the bridle path. In 1931 a vigorous colony came 
up here and bloomed freely, making a conspicuous show. The following 
year only two or three plants remained and a search during the past 
summer failed to reveal any trace of it. A few plants of this species 
have also been found in the poplar group near Peters Hill. The king 
devil {Hieracium jiorentinum) has also recently appeared there, but in 
greater abundance farther up on the slopes of the hill among the thorns. 
This year it was also noted in the Celtis group near the North Woods. 
Fool's parsley has become more common at two localities, at the foot 
of the hills near the Leitneria group and along the base of Bussey bank 
near the Forsythia planting. A vigorous plant of the blue weed came 
up along the bridle path opposite the Horsechestnut group last year. It 
was blooming freely when cut down by the mowers, but this year no 
trace of it could be found. Two weedy grasses, Eleusine indica (the 
goose grass) and Eragrostis cUianensis have recently appeared on dumps 
and in waste ground at the old quarry along Bussey Street, and the 
latter also in the South Street nursery. The flower-of-an-hour, Japa- 
nese knotweed, four-o-clock, Cyperus esculentus and other cultivated 
and weedy things have also turned up here, and this and the South 
Street tract continue to be the chief sources for plants of this class. At 
the latter place, where a considerable tract of low fertile land surround- 
ing the pond is still unoccupied and grown up with weeds, a real plant 
succession has been taking place. A number of plants that appeared 



JOURNAL OF THE ARNOLD ARBORETUM 



here soon after the construction of the pond have already been crowded 
out by the more aggressive weeds but other immigrants arrive from time 
to time. Last year Boltonia asteroides and Galium asprellum were 
noted here for the first time, and the latter at least has become more 
abundant. Last year a large colony of the smooth form of the perennial 
sow-thistle made a conspicuous show with its large yellow flowers. The 
yellow Canada lily also sent up a number of tall spikes above the other 
weeds at one side of the tract. On and about the rubbish dumps here 
several other weeds as well as escapes from cultivation have appeared. 
Amongst them are the gourd and jimson weed, as well as another species 
of thorn-apple, Datura Metel. 

A number of herbaceous plants, as well as a few shrubs and trees, 
persist in the Arboretum from the old gardens formerly planted here, 
and some of these appear to be holding their own or increasing in num- 
ber. The Virginia spiderwort and ox-eye have spread into the meadow 
near the old Dawson House, and Scilla, tulips and crocuses of various 
colors spring up in the grass each year making a pretty display. A 
bank near the barn of the State Laboratory is also carpeted with the 
brilliant blue of the Scilla blossoms in spring, and it is found more 
sparingly in other localities. The star of Bethlehem, day lily, narcissus 
and European bellflower are all well established in different parts of the 
Arboretum. More restricted are the white-flowered form of Campanula 
persicijolia, which is growing along the lilac border and in the open 
ground on the east side of Bussey Hill, and the English violet, abundant 
but local along a bank near the Jamaica Plain gate. A few plants of a 
small perennial pea {Lathyrus pannonicus var. versicolor) come up and 
bloom each year on the east side of Bussey Hill below the Overlook, and 
Corydalis bulbosa is growing near the top of the bank below the Bussey 
greenhouses, where Dr. Edgar Anderson reports having seen it at least 
ten or twelve years ago. 

Several plants reported in the first list have already disappeared from 
the Arboretum, or at least have not been seen again. Most of these 
were waifs escaped from cultivation, such as the cock's-comb, candytuft, 
sweet alyssum, beef-steak plant, sneeze weed, corn flower and Nicotiana, 
or weeds of chance introduction, such as the jointweed, small bindweed 
and Bassia, but amongst them are also the cardinal flower, blue lobelia, 
wild senna, beard-tongue and Venus' looking-glass. It is quite possible 
that some of these will be introduced again at some time. The European 
smoke-tree, mentioned as having formerly been seen on Hemlock Hill, 
has been rediscovered growing there amongst the rocks, and a specimen 
of the moth mullen was collected during the present summer among the 
lilacs at the foot of Bussey Hill. Hepatica has been reintroduced at 



1935] PALMER, SPONTANEOUS FLORA OF ARNOLD ARBORETUM 85 

the place where it formerly grew near the edge of the North Woods. A 
few plants of both Hepatica americana and of H. acutiloba have been 
set out and it is hoped they will survive. It is probable that it was the 
latter species that was formerly native here and not H. americana as 
reported on the list. 

The presence of certain native plants persisting in places scarcely 
suitable to them at present offers some evidence as to former conditions 
in parts of the Arboretum and of the changes that have taken place, and 
this may have some value as a guide or check in future planting, since 
it affords a clue to both past and present soil and drainage conditions. 
Skunk cabbage continues to come up every year along what appears to 
be now a well-drained bank below the stone foot bridge over Bussey 
Brook and near the bald cypresses, as well as along the bridle path oppo- 
site the lindens, among the Chinese apples near Peters Hill, and at sev- 
eral other places. Sensitive fern, royal fern, and the lance-leaved violet 
coming up in the edge of the maple group, at the foot of the hills near 
the Ilex and Aesculus groups, along the Meadow Road by the laurels, 
and elsewhere, indicate former boggy areas and show that the water 
table even now is very near the surface in wet seasons. Along the edge 
of the path near a planting of Aesculus parviflora the water pennywort 
has even managed to survive and still sometimes to produce fruit. The 
persistence also of certain shade-loving plants in open sunny situations 
where they are gradually being exterminated furnishes evidence, in some 
case no longer available from records, that the protecting woods have 
not long been removed. 

As the Arboretum has developed, the natural drainage has been modi- 
fied or changed in many places. A brook formerly entered the Arbore- 
tum area from the west through a gap in the low hills near the Aesculus 
and Linden groups. A small tributary which drained the ponds near 
the Forest Hills gate joined it as it flowed across the level ground at the 
foot of the hills and into the low ground across the Meadow road. The 
water from this brook is now carried under ground and only a small 
fragment of the course of the smaller stream can be made out in the 
somewhat boggy area where the corkwood is now growing. The course 
of these streams is shown on old maps and their history helps to account 
for the presence here of such native plants as Carex crinita, Scirpus rubro- 
tinctus, Pilea pumila, Callitriche palustris, Ludvigia palustris, Hyperi- 
cum majus, Hydrocotyle americana and Scutellaria lateriflora, as well 
as suggesting the great changes that must have taken place in the char- 
acter of the flora and the many plants that must have disappeared from 
the area since the time when these brooks flowed across the fields and 
into the low meadow and bog. 



86 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

The course of Bussey Brook has also been diverted or straightened at 
several points, and the amount of water that it formerly received from 
seepage and springs has been greatly diminished by the clearing off or 
thinning of the forests on the hills and of thickets along its margins, the 
water now running off rapidly after a rain instead of sinking into the 
humus and soil. The diversion of its permanent water supply has also 
been made almost complete by the construction of ditches and sewers 
along its upper course beyond the Arboretum. It is evident from a 
study of the surviving native plants as well as from the topography that 
a small swamp or bog formerly occupied the low ground a little way 
above the stone foot bridge and between the base of Hemlock Hill and 
the slopes now occupied by the conifer groups. A spring and little 
rivulet carrying water except in the dryest seasons still feed the brook 
on the north side, and small areas are kept wet by seepage water here 
for a considerable part of the year. But even beyond these moist places 
some traces of the palustral flora still remain. Such plants as Onoclea 
sensibilis, Lycopodium complanatum var. flabclliforme, Carex lurida, 
and sprouts of Salix pedicellaris, Alnus incann, Vaccinium corymbosum 
and of an undetermined species of Rhododendron have been found here. 
On similar evidence it can be seen that certain parts of Bussey Hill 
and of Peters Hill were covered with forest until recent years. An early 
map of the Arboretum shows native woods extending over a large part of 
Bussey Hill, and Bussey Woods is mentioned on some of Mr. Faxon's 
plant labels, but I have seen no similar record in regard to Peters Hill. 
This hill was probably at one time covered with forest, but from the pres- 
ent composition of the flora it may be inferred that much of the sur- 
face wa;, cleared and used for pasture or other purposes at an early day, 
but thai remnants of the forest remained along the east and north sides 
until quite recently. Sprouts of a number of characteristic forest trees 
and shrubs continue to come up here in spite of annual mowing, and 
stumps of several large trees are still in evidence. A single large white 
oak survives on the east slope and in its protection a number of plants are 
growing that are not found in the open ground. Others still huddle 
rather pathetically about the decaying stumps or in the meager shade 
afforded by the small thorn trees. Several groups of sprouts of ths 
tremblin,^ aspen and of the large-toothed aspen are found on the hill- 
sides as well as scattered specimens of various species of oak, hickory, 
chestnut, birch, elm, wild-cherry and ash; and among shrubs are the 
bayberry, sweet fern, meadowsweet, dwarf juneberry {Amclanchicr 
oblongijolia), choke-cherry, sheep laurel, panicled dogwood and several 
species of wild rose, blackberry, dewberry, raspberry and blueberry. A 
few depauperate plants of the ground pine still survive in one spot, and 



1935] PALMER, SPONTANEOUS FLORA OF ARNOLD ARBORETUM 87 

a large colony of false Solomon's seal (Smilacina stellata) is growing 
about one of the old stumps, with the wood aster, false lily of the valley 
and other plants that are evident relics of a woodland flora. The com- 
plete removal of the forest or thickets on rather steep slopes has resulted 
in the loss of the humus and in the leaching out and removal of the soil, 
which is reflected in the slow growth and poor condition of some of the 
Crataegus trees on this side of the hill. 

The original native flora has almost entirely disappeared from most 
of the Arboretum, and increasing inroads upon such fragments of it as 
remain will necessarily be made as the planting of cultivated trees and 
shrubs continues and as they come to occupy the ground more fully, 
and as the surface and soil are further modified by drainage, grading, 
and the bringing in of outside soils and fertilizers. Such traces of it as 
still remain have considerable significance in a number of ways, and a 
record of it should be of increasing interest and value in the future. The 
introduction of weeds and other exotic plants from various sources is 
certain to continue, and specimens of them should be collected and rec- 
ords kept as they appear or are discovered, and it may be thought worth 
while to issue another supplement to the Spontaneous Flora at some 
time in the future. 

I wish to express my thanks to the members of the staff and other 
friends who have shown an interest in the native and introduced plants 
of the Arboretum through the contribution of specimens and other 
assistance. Mr. Frederic W. Grigg has examined a number of the 
grasses, sedges, and other plants and has aided in their determination. 
I am also under obligation to Mr. C. A. Weatherby for assistance on 
points of nomenclature, and to Professor J. G. Jack and Professor Alfred 
Rehder of the Arboretum staff for information about early conditions 
in the Arboretum and for other suggestions, as well as to several others 
who have brought in specimens of plants found in the Arboretum. 

ENUMERATION OF THE ADDITIONAL PLANTS COLLECTED^ 
Polystichum acrostichoides (Michx.) Schott. Christmas Fern. One 
plant found by Grant D. Darker on east slope of gravelly ridge, North 
Woods, probably surviving from a former native colony ; also several 
plants on rocky wooded slopes on north side of Hemlock Hill. Nos. 
402 73,42588. Rare. 
Thelypteris spinulosa (O. F. Muell.) Nieuwland var. intermedia (Willd.) 
Nieuwland. Spinulose Shield Fern. Base of wooded hills, near 
Aesculus group. No. 39678. 



88 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

Juniperus communis L. var. depressa Pursh. Low Juniper. Rocky 
ground, Central Woods. No. 36405. Rare. 

Typha latifolia L. Common Cat-tail. Borders of ponds and brooks. 
No. 39605. 

Sagittaria latifolia Willd. f. gracilis (Pursh) Robinson. Muddy mar- 
gins of Pond, South Street tract. No. 40248. 

Potamogeton epihydrus Raf. var. Nuttallii (Cham. & Schlecht.) Fer- 
nald. In shallow water along muddy margins of pond. South Street 
tract. No. 42723. 

Panicum philadelphicum Bernh. Waste and cultivated ground, No. 
39669. Uncommon. 

Panicum dichotomijiorum Michx. Waste and cultivated ground. Nos. 
38229, 39704. 

Panicum depauperatum Muhl. var. psilophyllum Fernald. Rocky slopes 
and ledges. Nos. 25608, 25627. 

Panicum linearijolium Scribn. Rocky ledges, conglomerate outcrops. 
No. 40172. 

Panicum lanuginosum Ell. var. Lindheimeri (Nash) Fernald. (P. 
Lindheimeri Nash). Common in dry open woods and meadows. 
Nos. 39588, 39621, 39638, 42646. 

Panicum lanuginosum var. septcntrionale Fernald. Border of woods. 
No. 39635. 

Panicum commutatum Schultes var. Ashei (Pearson) Fernald. Dry, 
open woods, South Street tract. No. 42675. 

Panicum oligosanthes Schultes var. Scribnerianum (Nash) Fernald (P. 
Scribnerianum Nash). Dry gravelly banks between Shrub Collec- 
tion and Arborway wall. Nos. 39627, 39694. 

Panicum latijolium L. Edge of North Woods, near Celtis group. No. 
39637. Rare. 

*Eckinochloa crusgalli (L.) Beauv. f. longiseta (Trin.) Farwell. Culti- 
vated and waste ground, with the typical form. No. 28102a. 

Echinochloa muricata (Michx.) Fernald. Cultivated and waste ground. 
No. 42212. 

Aristida dichotoma Michx. Poverty Grass. Sterile gravelly banks, 
between Shrub collection and Arborway wall, and also on conglom- 
erate outcrops in Conifer group. Nos. 38190, 39742. 

Aristida gracilis Ell. Sterile gravelly banks, between Shrub Collection 
and Arborway wall. No. 38191. 

*Agrostis canina L. Brown Bent Grass. Dry open ground, slopes of 
Bussey Hill. No. 39578. 

*Eragrostis ciliancnsis (All.) Link ex Lutati. {E. megastachya Link.) 



1935] PALMER, SPONTANEOUS FLORA OF ARNOLD ARBORETUM 89 

Waste ground, old quarry near Bussey Street, and also as a weed in 
South Street nursery. Nos. 38197, 38227. 

Eragrostis pectinacea (Michx.) Steud. Meadows and dry banks. Nos. 
39687, 40234. 

Glyceria septentrionalis Hitchc. Margins of Bussey Brook, near Coni- 
fer group. No. 39661. 

Festuca ovina L. Sheep's Fescue. Gravelly slopes, south side of 
Peters Hill. Nos. 36461, 36506. 

Festuca rubra L. var. commutata Gaud. Open ground, border of Aescu- 
lus group. No. 40199. 

*Cyperus esculentus L. Rich waste ground, old quarry along Bussey 
Street. No. 42202. 

Scirpus rubrotinctus Fernald. Along little brook, near Aesculus group. 
No. 36586. 

Carex Crawfordii Fernald. Open banks and meadows. No. 40162. 

Carex tenera Dewey (C. straminea of Gray's Man.). Dry ground, bor- 
ders of woods and meadows. Nos. 42609, 42624. 

Carex laxiflora Lam. var. gracillima Boott. Moist banks of pond, near 
Forest Hills gate. No. 36501. 

Carex laxiflora var. leptonervia Fernald. Shaded banks and borders of 
woods. Nos. 39589, 42650. 

Carex canescens L. var. disjuncta Fernald. Springy ground, near base 
of Peters Hill. No. 36531. 

*Carex caryophyllea Lat. Dry slopes and gravelly banks, Peters Hill, 
Bussey bank, and near Dawson House. Nos. 36455, 40132, 40165. 

Carex angustior Mackenzie. (C. stellulata Good. var. angustata Carey) . 
Local in boggy ground about spring, along southeast side of Peters 
Hill. Nos. 36532, 40128; also a specimen collected by Mary E. Gil- 
breath, June 6, 1892, in herb, of New England Botanical Club. 

Carex panicea L. Grassy slopes of Peters Hill, in Crataegus group. 
Nos. 36423, 36460, 36529, 36558. 

Carex pennsylvanica Lam. var. lucorum (Willd.) Fernald. Rocky 
banks, near top of Hemlock Hill. No. 36578. 

Carex varia Muhl. Dry rocky ledges, south side of Hemlock Hill, and 
along base of hills. North Woods. Nos. 40029, 402 79a, 40281a. 

Carex Goodcnowii J. Gay. Wet ground about spring, southeast side of 
Peters Hill, and also in low meadows near Administration Building. 
Nos. 40129,40159,40177. 

Carex communis Bailey. Specimen in the herbarium of the New Eng- 
land Botanical Club, collected by C. E. Faxon, May 30, 1878; also 
found on Hemlock Hill. No. 36455. 



90 JOURNAL OF THE ARNOLD ARBORETUM hoL. xvi 

Carex brcvior (Dewey) Mackenzie. Dry open woods, Oak group. No. 
42621. 

Carex longirostris Torr. Shaded ground, at foot of Hemlock Hill, on 
south side. No. 40276. 

Carex crinita Lam. var. gynandra (Schwein.) Schwein. & Torr. Wet 
rocky ground along Bussey Brook at foot of Hemlock Hill. No. 
42622. Rare. 

Carex lupulina Muhl. Margin of small pond west side of Bussey Hill. 
Nos. 40188, 40205. 

*Tradescantia virginiana L. Virginia Spiderwort. Freely escaped 
into meadow, near Dawson House. Nos. 39646, 39675. 

*Luztila ncmorosa (Poll.) Mey. Open grassy border, near Dawson 
nursery. No. 40185. 

Erythronium americanum Ker. Yellow Adder's-tongue. Under trees 
at base of Hemlock Hill, on southeast side, where leaves come up 
each year from a small colony, but it has not been found flowering. 
There is also a colony which flowers freely on a wooded hillside of the 
Adams Nervine Hospital grounds, only about 20 feet from the Arbore- 
tum boundary, where the flowering specimen was collected. Nos. 
36387,36415. 

Allium canadense L. Wild Garlic. Open wooded banks. South Street 
tract. No. 42620. 

*Scilla sibirica Andr. Well established on banks near State Laboratory 
barn, and at top of Bussey Bank. No. 36361. 

Sisyrinchium angustijolium Mill. Dry open slopes of Peters Hill, in 
Crataegus group and on slopes of Bussey Hill. Nos. 36497, 40026, 
40144. 

Cypripedium acaule Ait. Stemless Lady's Slipper. Under hemlocks 
and pines, near the base of Hemlock Hill, on the northeast side. No. 
40272. 

Salix lucida Muhl. Shining Willow. Along small spring brook. Coni- 
fer group. No. 39604. 

Salix pedicellaris Pursh. Bog Willow. In boggy ground about spring, 
southeast side of Peters Hill, and margins of Bussey Brook near 
Conifer group. Nos. 36384, 36418. 

Salix humilis Marsh. Prairie Willow. Open slopes of Peters Hill, in 
Crataegus group. Sprouts coming up after repeated mowing. No. 
39706. 

*Salix alba L. var. calva G. F. W. Mey. Wet ground about spring. Pop- 
lar group. No. 36530. 

*SaUx jragilis L. Crack Willow. Slopes of Peters Hill ; sprouts per- 
sisting after mowing. No. 39718. 



1935] PALMER, SPONTANEOUS FLORA OF ARNOLD ARBORETUM 91 

*Salix pyrijolia Anders. Waste ground near pond, South Street tract. 

No. 40250. 
Populus tremuloides Michx. Quaking Aspen. Several large colonies 

of sprouts persistent after repeated mowings, on Peters Hill, in Cra- 
taegus group. No. 36413. 
Carya ovalis (Wang.) Sarg. Broom Hickory. North Woods. No. 

39651a. 
Quercus prinoides Willd. Scrub Chestnut Oak. Rocky banks near 

top of Hemlock Hill and on conglomerate outcrops, Central Woods. 

Nos. 36456, 39692. Rare as a native plant. 
Quercus Rekderi Trelease. {Q. ilicijoliu X velutina). Rocky slope. 

Central Woods. No. 39682. Rare. 
*Quercus Leana Nutt. {Q. imbrkaria X velutina). Rocky slope, Cen- 
tral Woods; also sprouts that appear to be this hybrid coming up 

spontaneously in Oak group. No. 39683. 
Polygonum Careyi Olney. Waste ground and cultivated beds, near State 

Laboratory barn. Nos. 42689, 42708. 
Polygonum lapathijolium L. Moist waste ground and borders of ponds. 

Nos. 42693, 42709. 
Polygonum punctatum Ell. var. leptostachyuni (Meisn.) Small. No. 

42711. 
*Polygonum Sieboldii De Vriese. Waste ground, old quarry near Bus- 

sey Street. No. 42201. 
*Rumex crispus X obtusifolius. Margins of small spring brook, Conifer 

group. Growing with the parent species. No. 40202. 
*Mirabilis Jalapa L. Four-o-clock. Waste ground, as a waif, old 

quarry near Bussey Street. No. 39703. 
*Aristolochia Kaempjeri Willd. Japanese Birthwort. Open ground 

near Administration Building, and also along Meadow^ Road near rock 

spring. No. 42673. Aristolochia Clematitis included in the original 

list, without collection, was probably based on young sprouts of this 

species, and should therefore be dropped. 
*Spergula arvensis L. Corn Spurry. In cultivated ground among 

laurels and other shrubs, near Hemlock Hill. Nos. 38171, 38190. 
*Sagina decumbens (Ell.) T. & G. Pearlwort. Grassy borders and 

waste ground, old quarry and along Valley Road. Nos. 42633, 42660. 
*Silene antirrhina L. Sleepy Catchfly. Waste and cultivated ground. 

Maple group. No. 32635. 
*SUene Armeria L. Sweet William Catchfly. Grassy borders, near 

Administration Building. No. 42688. 
*Saponaria officinalis L. Bouncing Bet. Meadows and waste ground, 

near Dawson nursery. No. 39684. 



92 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

^Ranunculus bidbosus L. Bulbous Buttercup. Common in meadows. 

Omitted through oversight from first list. Nos. 36438, 36526, 36587. 
Anemone quinquejolia L. Wood Anemone. Local in open woods, 

North Woods, slopes of Bussey Hill, near Oak group, and South 

Woods, near Peters Hill. Nos. 36372, 36398, 36410. 
*Clematis paniculata Thunb. Escaped into meadows, in Tilia group, 

No. 40182. 
*Aquilegia vulgaris L. Garden Columbine. Bussey bank, in partial 

shade. No. 42607. 
*Lirwdendron Tulipifera L. Tulip Tree. There is a large tree of this 

species in the edge of the woods along the base of Hemlock Hill that 

appears to be spontaneous. 
*Berberis Thunbcrgii DC. Established in woods near top of Peters 

Hill, and also on Hemlock Hill. No. 36380. 
Benzoin aestivale (L.) Nees. Spice Bush. Near base of gravelly ridge, 

North Woods. No. 36437. 
*Corydalis bulbosa DC. Shaded bank, near Bussey Greenhouse. No. 

40005. 
*Rorippa sylvestris (L.) Bess. Yellow Cress. Waste ground and 

cultivated borders. Nos. 39562, 39657. 
*Diplotaxis muralis (L.) DC. Waste ground about pond, rich soil. 

South Street tract. No. 38233. 
*Ribcs sativum Syme (R. vulgare Lam.) Red Currant. Wooded 

bank near Forest Hills gate, also in woods near top of Peters Hill. 

Nos. 36394, 36412a. 
*Gillenia trijoliata (L.) Moench. Indian Physic. Open woods, edge 

of Oak group. No. 36602. Rare. 
Aronia arbutijolia (L.) Ell. Chokeberry. Rocky open woods, Cen- 
tral Woods, and also on north slope of Peters Hill. No. 36404. 
*Malus baC€ata Borkh. var. mandshurica Schneider. Northeast side of 

Peters Hill. No. 40017. 
*Malus hupehensis (Pamp.) Rehder (M. theijera Rehder). Northeast 

slopes of Peters Hill. No. 40253. 
Amelanchier stolonijera Wiegand. Open woods, top of Hemlock Hill. 

No. 40265. 
Crataegus rotundijolia Moench. Bussey Woods, C. E. Faxon, June 6, 

1882; Geo. Engelmann, Aug. 27, 1882; Peters Hill, C. E. Faxon, Oct. 

1, 1883, Sept. 21, 1889; /. G. Jack, May 23, 1900. Sprouts of this 

plant still persist in the edge of the woods near the top of Peters Hill. 
Rubus Idaeus L. var. strigosus (Michx.) Maxim. Northeast slope of 

Peters Hill, in Crataegus group. No. 39725. 



1935] PALMER, SPONTANEOUS FLORA OF ARNOLD ARBORETUM 93 

*Rubus parvijolius L. Escaped and well established in Quercus group. 

No. 40203. 
Rubus Jeckylanus Blanchard. C. E. Faxon, June 7, 1913. 
Rubus allegheniensis Porter. Open woods and banks. Nos. 36564, 

40145. 
Rubus flagellaris Willd. Dewberry. Common in open rocky woods 

and on conglomerate outcrops. C. E. Faxon, July 19, 1909. Nos. 

36451, 37689. 
Rubus Randii (Bailey) Rydb. C. E. Faxon, July 21, 1912; July 31, 

1912; July 7, 1913. 
*Potentilla canadensis var. villosissima Fernald. Cultivated ground. 

No. 28014. (See Rhodora, 33: 187. 1931). 
*Prunus Cerasus L. Morello Cherry. Rocky woods on Hemlock 

Hill, and in woods near top of Peters Hill. No. 36493. 
*Prunus pumila L. var. susquehanae Jaeg. Sand Cherry. Open woods, 

Hickory group, and persisting as sprouts after repeated mowing on 

Peters Hill. Nos. 36493, 40259. 
'^Colutea media Willd. Bladder Senna. Op)en banks near Arborway 

wall and on Overlook. W. H. Judd, July, 1931. 
*Amorpha fruticosa L. False Indigo. Borders of pond, near Forest 

Hills gate. No. 38240. 
Lespedeza hirta (L.) Hornem. Hairy Bush-clover. Rocky banks 

and borders of woods. Central Woods, near chestnuts, slopes of 

Bussey Hill and South Woods. Nos. 39653, 39735. 
*Medicago hispida Gaertn. Bur Clover. Waste ground, old quarry 

near Bussey Street. No. 38201. 
Apios americana Med. {A. tuber osa Moench). Ground Nut. Open 

grassy slopes, in Malus group, near foot of Peters Hill. 
*Lathyrus pannonicus (Kramer) Garcke var. versicolor (Gmel.) Maly. 

Open bank near small pond, on slope of Bussey Hill. Nos. 39660, 

40008. 
Oxalis europaea Jord. f. villicaulis Wiegand. Cultivated and waste 

ground, with the typical form. No. 39748. 
Oxalis s trie t a L. Rocky open ground, South Woods. No. 38211. 
Polygala sanguinea L. Purple Milkwort. Open slopes of Peters Hill, 

in Crataegus group. No. 39655. 
Callitriche palustris L. Moist banks and borders of ponds and brooks. 

Further study of more mature specimens show that plants reported in 

original list as Callitriche heterophylla Pursh are this species, and the 

latter should therefore be dropped from the list. 
*Evonymus obovatus Nutt. Trailing Strawberry Bush. Moist 

ground at base of hills, North Woods. No. 36569. 



94 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

*Cclastrus orbkulatus Thunb. Rocky slopes, south side of Hemlock 

Hill. No. 42591. 
*Ampelopsis humidijolia Bunge. Dumps and waste ground. South 

Street tract. No. 42694. 
"^Hibiscus Trionum L. Flo\ver-of-an-hour. Waste ground, old 

quarry near Bussey Street. No. 38199. 
*Malva parviflora L. Waste ground, old quarry, near Bussey Street. 

No. 38200. 
*Malva vertkillata L. var. crispa L. Curled Mallow. Waste ground, 

old quarry south side of Bussey Street. No. 42794. 
*Sida hermaphrodita (L.) Rusby. Virginia Mallow. Rocky open 

ground, near Centre Street gate. No. 25893. This was incorrectly 

identified in original list as Napaea dioica L., a plant which it closely 

resembles. 
Lechea tenuijolia Michx. 

of woods, Central Woods and South Woods. 
Lechea intermedia Leggett. Gravelly banks, near top of Peters Hill. 

No. 42701. 
Viola pedata L. var. lineariloba DC. Bird-foot Violet. Rocky open 

woods, top of Hemlock Hill and a single plant found on east slope of 

Peters Hill. Nos. 36557, 40256. 
Viola sororia Willd. Meadows, Aesculus group. No. 36477. 
Viola latiuscida Greene. Common in meadows and on open banks. 

Nos. 36366, 36388. 
Viola sagittata Ait. Arrow-leaved Violet. Moist grassy ground, 

near Arborway wall, in Maple group. Nos. 40268, 42670. Rare. 
*Viola odorata L. English or Sweet Violet. Shaded bank, near 

Jamaica Plain gate. Nos. 36417, 39597, 40007. 
Viola fimbriatula X papilionacea. Base of hills, near Leitneria group. 

No. 42682. 
*Epilobium hirsutum L. Low ground along brook, near Arborway wall 

and opposite Administration Building, C. H. L. Gebjert. 
*Aralia hispida Vent. Bristly Sarsaparilla. Waste ground, old 

quarry along Bussey Street. 
*Cornus stolonijera Michx. Red Osier. About small abandoned 

quarry, South Woods. Nos. 36513, 36524. 
Pyrola americana Sweet. Round-leaved Wintergreen. Woods, north 

side of Hemlock Hill. Nos. 39565, 39591. 
Vacdnium corymbosum L. High Blueberry. Open woods, Central 

Woods, slopes of Peters Hill, and along small brook near leitnerias. 

Nos. 39585, 39631. 40143. 



1935] PALMER, SPONTANEOUS FLORA OF ARNOLD ARBORETUM 95 

Trientalis americana (Pers.) Pursh. Star Flower. Woods, top and 

north slopes of Hemlock Hill, and base of hills, North Woods. Nos. 

36454, 40030, 40271. 
*Ligustrum vulgare L. Privet. Escaped in thickets and open woods. 

Peters Hill and South Woods. No. 38184. 
*Syringa vulgaris L. Common Lilac. Persistent and spreading from 

cultivation in several places. J. G. Jack. 
*Phlox pankulata L. Garden Phlox. Rich waste ground, South 

Street tract. Nos. 40242, 42676. 
*Echium vulgare L. Blue Weed. Weedy and grassy border along 

bridle path, near Aesculus group. Nos. 40183, 40236. 
Lycopus rubellus Moench. Wet meadow, between Administration 

Building and Arborway wall. No. 39691. 
*Physalis heterophylla Nees var. nyctaginea (Dunal) Rydb. Rich open 

ground. South Street tract. No. 28149. 
*Datura Stramonium L. Jimson Weed. On dump. South Street tract. 

No. 38236. 
*Datura Metel L. Thorn Apple. On dump. South Street tract. No. 

42208. 
*Lycopersicon esculentum Mill. Tomato. Not rare in waste and culti- 
vated ground and sometimes producing fruit and self-seeding. No. 

39732. 
Veronica peregrina L. Neckweed. In waste and cultivated ground. 

Nos. 36442, 36516. 
Epijagus virginiana (L.) Bart. Beech Drops. On superficial roots of 

beech trees, south side of Hemlock Hill. No. 38185. 
Plantago major L. Common Plantain. A common weed in waste 

ground. Nos. 39685, 39749, 40214. 
"Galium asprellum Michx. Rough Bedstraw. Moist weedy ground. 

South Street tract. Nos. 40251, 42678. 
Houstonia caerulea L. Bluets. Among shrubs and laurel bushes, foot 

of Hemlock Hill, northeast side, and also one plant collected amongst 

laurels near South Street gate. Nos. 40260, 40270. 
Diervilla Lonicera Mill. Bush Honeysuckle. Rocky open woods and 

ledges. Central and South Woods and slopes of Hemlock Hill. No. 

36571. 
* Lonicera alpigena L. Rocky ground, south slope of Hemlock Hill. No. 

36452. 
"Lonicera dioica L. Honeysuckle. Open woods. Oak group. No. 

42216. 
"Viburnum trilobum Marsh. High-bush Cranberry. Woods near 

top of Peters Hill. No. 38246. 



96 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

*Lagenaria leucantha (Duch.) Rusby. Gourd. On dump, South Street 
tract. No. 42209. 

Liatris scariosa Willd. Blazing Star. Open woods, low hills. Maple 
group. No. 42204. Rare. 

Solidago ulmijolia Muhl. Open woods, South Street tract. No. 42206. 
There is also a specimen of this species in the herbarium of the New 
England Botanical Club, collected by C. E. Faxon, on "Bussey Moun- 
tain," Sept. 4, 1887. 

Solidago odora Ait. Sweet Golden-rod. Open woods, slopes of Bus- 
sey Hill near Oak group and in South Woods. Nos. 38213, 39736. 
Rare. 

*Boltonia asteroides (L.) L'Her. In low weedy ground, South Street 
tract. No. 42211. 

Aster mulHflorus Ait. Dry open slopes, near top of Peters Hill. No. 
38721. 

Aster Unariijolius L. f. leucactis Benke. Slopes of Peters Hill, in Cra- 
taegus group. A form with smaller heads and white rays, growing 
with the species. No. 29730. 

Aster acuminatus Michx. Under shade of apple trees, on hillside north- 
west of Administration Building. No. 42213. Rare. 

Antennaria plantaginijolia (L.) Richards. Dry grassy slopes and 
meadows. Nos. 36362, 36386, 36444. Common. 

Antennaria canadensis Greene. Grassy slopes, gravelly soil, between 
Arborway wall and Shrub Collection, and also on slopes of Peters Hill. 
Nos. 36393, 40014. 

Antennaria neglecta Greene. Dry open slopes of Peters Hill. Nos. 
40014a, 40014b. 

*Heliopsis helianthoides (L.) Sweet. Ox-eye. Spreading from cultiva- 
tion into open ground, near Dawson House. No. 42681. 

*Helenium nudiftorum Nutt. Sneezeweed. Open grassy ground, near 
South Street gate and also near Administration Building. Nos. 42205, 
42655, 42680. 

'^'Chrysanthemum segetum L. Corn Marigold. Waste ground and 
dump, field near Dawson House. 

Senecio aureus L. Golden Ragwort 
Tilia group and bridle path, and a 
Hill. Nos. 36476, 36511. 

*Senecio viscosus L. Clammy Groundsel. Waste and cultivated 
ground, South Street tract and Conifer group. Nos. 38232, 39750. 

*Hypochaeris radicata L. Cat's-ear. Grassy open ground northwest 
of Administration Building. No. 42672. 



1935] PALMER, SPONTANEOUS FLORA OF ARNOLD ARBORETUM 97 

*Sonckus arvensis L. var. glabrescens Guenth. Wimm. & Grab. Per- 
ennial Sow Thistle. Rich waste ground, near pond, South Street 
tract. Nos. 42665, 42691, 42699. 

*Sonchus asper (L.) Hill. Sow Thistle. Waste and cultivated ground. 
Nos. 35710, 35927. These numbers were listed through error as Son- 
chus oleracea in original list. Both species are found, and the latter 
has been collected under numbers 39739a and 40219. 

*Crepis capillaris (L.) Wallr. Hawk's Beard. Grassy open slopes 
near Administration Building and also along Meadow Road near 
Hemlock Hill. Nos. 42192, 42629, 42671. 

*Lactuca scariola L. Prickly Lettuce. Waste ground. South Street 
tract. No. 40245. 

*Lactu€a sagittijolia Ell. Waste ground, South Street tract. No. 
40244. 

*Lactuca spicata (Lam.) Hitchc. var. m/egn/o/m (Gray) Britton. Open 
woods, Carya group. No. 39648. 

*Hieradum florentinum All. King Devil. Grassy slopes of Peters 
Hill in Crataegus group, near Roslindale Gate and in Celtis group. 
Nos. 36581, 39563, 40170, 40180. 

*Hieracium vidgatum Fries. Grassy open ground near Platanus nurs- 
ery. No. 42795. 



JOURNAL OF THE ARNOLD ARBORETUM 



TABLE OF CONTENTS 

ntroduction 99 

t^elative susceptibility of hosts within genera of the Ponioideae 99 

History 99 

Methods used to determine susceptibihty 

Cultural technique 

Recording of data obtained from inoculations 

Investigations and conclusions with respect to the var 

genera considered 

Crataegus 

obtained by observations 



Presentation of data obtained by serial inoculations. . 
Factors influencing the relative susceptibility of 
Crataegus 

1. The geographical distribution of Crataegus 

2. The role of the foliar cuticle 

3. The degree and the duration of the period of 

susceptibility 

Correlation of the data to classify the groups of 

Crataegus with respect to their relative susceptibility 

Suggestions for the selection of resistant species and 



Relative susceptibility as indicated by 
Sorbus — Relative susceptibility as indicated by 

Malus — Relative susceptibility as indicated by 
inoculations 

Amelanchier, Cydonia, Comptonia, Crataegomespilui 
pilus, Myrica, Photinia, Sorbaronia, Sorbopyrus . 



1935] MacLACHLAN, HOSTS OF GYMNOSPORANGIUM GLOBOSUM 99 

IV. The hosts of Gymnosporangium globosum Farl. 

Hosts for the & 1 stage 12^) 

Hosts for the HI stage 137 

V. Summary 137 

VI. Acknowledgments 139 

VII. Bibliography 139 

VIII. Explanation of Plates 141 

I. INTRODUCTION 
Gymnosporangium globosum Farl., a heteroecious rust, is restricted 
in its telial phase to a limited number of species and varieties of Juni- 
perus. To the aecial phase, however, representatives of at least ten 
genera within the Pomoideae may serve as hosts; and certain of these 
genera, especially Crataegus, include a large number of species and 

In spite of the number of hosts hitherto reported for G. globosum, very 
little information is available regarding the relative susceptibility of the 
hosts. This is a question of considerable importance because of the 
great damage done by the rust. A determination of the immune species 
and the comparative resistance of susceptible species within the various 
relevant host genera constitute the major part of this paper. 

Concurrently with the investigations which led to a determination of 
the relative susceptibility of the hosts, the writer was enabled to compile 
a more nearly complete list of the known hosts, from which it appears 
that, instead of the approximately one hundred hosts hitherto reported, 
the number of hosts should be conservatively estimated at more than 
six hundred. This list constitutes the latter portion of the paper. 

The work on the problems outlined above has been carried on at the 
Arnold Arboretum of Harvard University, where may be found one of 
the finest collections in the world of living representatives of species and 
varieties of Juniperus and of the Pomoideae. 



HISTORY 
The earliest successful attempt to determine pomaceous hosts of G. 
globosum Farl. by means of cultures may be credited to Farlow (1880) 
who, in 1876-7, using teliospores from Juniperus virginiana, obtained 
spermogonia on Crataegus tomentosa. Farlow (1885) also made suc- 
cessful cultures on leaves of Crataegus Douglasii, Crataegus Oxyacantha 



100 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

and apple seedlings; but he obtained spermogonia only, because his ex- 
perimental leaves molded before the aecial stage could develop. Thax- 
ter (1887) obtained spermogonia on Crataegus coccinea, Malus pumila, 
Sorbus americana, and Amelanchier canadensis; and in 1887-8 
(Thaxter, 1889) obtained spermogonia on Sorbus americana and Cy- 
donia oblonga (= Cydonia vulgaris), and both spermogonia and aecia 
on Crataegus coccinea and Malus pumila. In a later report Thaxter 
(1891) confirmed the previous results on Malus pumila and records suc- 
cessful infections of Crataegus crus-galli and Sorbus americana, both 
resulting in aecial development. In 1906, cultures were made by 
Arthur (1907) on Crataegus Pringlei and Sorbus americana resulting 
in spermogonia and aecia, and on Malus coronaria giving spermogonia 
only. In 1908 Arthur (1909) confirmed his results on Crataegus 
Pringlei, and in 1909 (Arthur, 1910) those of Farlow on Crataegus 
coccinea. 

Since Arthur's work more than one hundred suscepts have been added 
to the host list, mostly by observations made in the field. Authors who 
have contributed or made significant reference to this list include Clin- 
ton (1904 and 1934), Stewart (1910), Kern (1911), Stevens and Hall 
(1910), Arthur (1921, 1924, 1926 and 1927), Burnham and Latham 
(1917), Hesler and Whetzel (1917), Jackson (1921), Hunt (1926), 
Anonymous (1930), Thomas and Mills (1930), Sherbakoff (1932), and 
others. Bliss (1931), by culture, obtained abundant spermogonia and 
aecia on Crataegus mollis, but obtained only flecking on nine varieties 
of commercial apples. 

These previous reports, together with the investigations made by the 
writer, warrant the conclusion that the genera involved as suscepts for 
the aecial phase of G. globosum are confined to the sub-family Pomoi- 
deae, and include Amelanchier, Crataegus, Cydonia, Malus, Mespilus, 
Pyrus, Sorbus, and the hybrid genera Crataegomcspilus, Sorbaronia and 
Sorbopyrus. 

METHODS USED TO DETERMINE SUSCEPTIBILITY 
Two methods of approach were utilized in the determination of the 
-various hosts and their relative susceptibility within each genus, namely, 
(1) quantitative observations on natural infection, and (2) serial arti- 
ficial inoculations during the progressive development of the foliage to 
determine both the degree and the duration of the period of suscepti- 
bility. These methods of approach were especially applicable to Cratae- 
gus which is by far the largest genus susceptible to G. globosum. All 
cultures and observations were made on trees in the Arnold Arboretum. 



19351 MacLACHLAN, HOSTS OF GYMNOSPORANGIUM GLOBOSUM 101 
CULTURAL TECHNIQUE 
The cultural technique adopted was similar to that described by 
Crowell (1934). The inoculum was collected either the previous eve- 
ning, or in the morning prior to inoculating, from Juniperus virginiana. 
Galls bearing abundant telial flanges were soaked in water until maxi- 
mum swelling had taken place; then the gelatinous mass was crushed to 
form a thick aqueous suspension of teliospores. Fresh inoculum was 
prepared every two hours during inoculation in order to eliminate any 
possibility of crushing the promycelia emerging from the germinating 
teliospores, since microscopic examination revealed that the latter would 
germinate within that time. All inoculations were carried out in dupli- 
cate. For each test six to ten leaves on a twig were inoculated ; the re- 
mainder of the tree served as a control. The spore suspension was 
painted on both sides of the leaves with a camel's hair brush; then a 
celluloid cylinder was slipped over the twig and the ends of the cylinder 
were plugged with moist sphagnum. Care was taken that the inocula- 
tion should not be exposed to direct rays of the sun ; otherwise burning 
of the leaves within the cylinder might occur. The sphagnum-plugged 
cylinder formed an excellent moist chamber; on removal of the tube 
two days later the sphagnum was always found to be still wet, and both 
the inside of the tube and the surfaces of the leaves were moist. Thus, 
with a heavy sowing of spores, a moist atmosphere in the inoculation 
tube, and a temperature below 25°C. the conditions for optimum spore 
germination and infection exceeded any that might occur in nature. 
Plate 127 fig. 5 illustrates a type set-up. 

RECORDING OF DATA OBTAINED FROM INOCULATIONS 

In recording data the inoculated plants were classified according to 

four categories or degrees of susceptibility, based on the number of sori, 

their relative size, and the pathogenic effect on an average-sized leaf. 

They are designated and defined as follows: 

— Immune; no visible infection obtained. 

1 — Resistant; one to five lesions which are relatively small, which 
cause very little leaf killing and no leaf drop; with or without 
aecia. This is a type of infection which causes no material harm 
to the host. 
2— Moderately Susceptible; five to twenty-five lesions per leaf 
with an intermediate pathogenic effect between categories 1 and 3 ; 
aecia always produced. This is a type of infection which, while 
reducing the photosynthetic leaf area and causing some leaf kill- 
ing, does not result in defoliation. 



102 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

3— Very Susceptible; twenty-five or more lesions which are usually 
large or fuse to form large masses and which cause severe leaf 
killing and leaf drop; abundant aecial horns produced in each 
lesion. This is a type of infection which ruins the foliage. 
While these definitions are, in general, applicable in allotting a suscept 
to any one category, they can not be considered as absolute criteria. 
Within the genus Crataegus, for example, as will be shown later in this 
paper, variation in susceptibility is for the most part not physiological 
but is dependent primarily upon a natural barrier, the cuticle; the 
probability that the basidiospore can produce infection varies inversely 
with the thickness of the foliar cuticle. Again, the amount of leaf kill- 
ing is dependent upon whether infection takes place on main veins or 
elsewhere on the leaf. Consequently, for Crataegus at least, the actual 
number of lesions per average-sized leaf was given major significance. 
In other genera, the type of infection was accorded major consideration. 
In the genus Pyrus, for example, certain species exhibited very small 
lesions which died shortly after spermogonia appeared, while other spe- 
cies of this genus showed larger lesions producing aecia. In general, 
however, the foregoing definitions were employed as the bases for plac- 
ing the various species within the different categories of susceptibility. 



For the sake of convenience the various host genera will be considered 
individually with respect to their relative susceptibility. All the known 
hosts within each genus will be listed at the end of the paper. 
Crataegus 

The Arnold Arboretum with almost one thousand trees comprising 
about five hundred and fifty named species and varieties, spread over 
twenty-four groups, afforded an excellent opportunity to study the 
relative susceptibility of the Crataegi. But, since the species of this 
genus hybridize so freely, and since the specific classification is still in 
an unstable condition, the time and labor involved in making inocula- 
tions for each of those species and varieties (especially in the large very 
susceptible groups where an abundance of natural infection was ob- 
served) would not justify the results that might be obtained; conse- 
quently typical representatives of each of the twenty-four groups were 
selected and the results were used as a basis of comparison by groups 
rather than by species. Likewise the data obtained on all the species 
and varieties by observations on natural infection were treated by 
groups rather than by species. 



19351 MacLACHLAN, HOSTS OF GYMNOSPORANGIUM GLOBOSUM : 



In July, 1932, a general spread of G. globosum was observed through- 
out the entire plantation of Crataegi in the Arnold Arboretum/ De- 
tailed observations were warranted by the fact that, within each group, 
the degree of infection was consistently uniform regardless of where the 
tree happened to be situated ; likewise a sharp line of demarcation could 
be seen between the number of foliar lesions per tree in a relatively 
resistant group, such as the Crus-galli, and the number per tree in a 
more susceptible group, such as the Coccineae or Anomalae. 

The amount of infection on any one tree while uniform was slight 
enough to allow fairly accurate counts to be made of the number of 
lesions per tree. While these data would hardly be adequate to permit 
comparison among species within any one group of Crataegus, they 
were sufficient for comparing the relative degree of susceptibility of the 
various groups represented in the Arboretum. As stated above, about 
one thousand trees were available for examination. 

Observations were made at the spermogonial stage, and again at the 
aecial stage of the rust. In order that the amount of infection per tree 
might be fairly compared the trees were graded as to size, five size- 
classes being used.' Counts were made of the number of foliar lesions 
per tree at both stages of the rust; where the counts exceeded one hun- 
dred per tree, the degree of infection was termed "100+".' A collection 
of herbarium material was assembled for permanent reference. 

In correlating the data obtained a method had to be devised by means 
of which a tree, for example size I, could be fairly compared with a tree, 
for example size V. The Coccineae, a group containing 46 species rep- 
resented by 82 trees, exhibited the highest percentage of infection lesions 
per unit-sized tree. This group was classed as having severe infection, 
and the values obtained for this group were selected as a basis of com- 



104 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

parison for all the other groups. It was found that for the CoCcineae: 

Size I (9 trees) averaged 24.3 lesions per tree. 

Size II {33 trees) averaged 51.7 lesions per tree. 

Size III (35 trees) averaged 75.7 lesions per tree. 

Size IV (5 trees) averaged 120.0 lesions per tree. 

Size V (0 trees). ^ 
If, for the sake of convenience, the ratio of the number of lesions per 
tree be changed from 24.3: 51.7: 75.7: 120.0: — to 25: 50: 75: 125: 
200, for the respective tree sizes, and these values be considered as units 
for classifying a tree as having severe infection, then by taking arbi- 
trary averages for the number of lesions required to class a tree as having 
moderate infection, mild infection, or no infection, the scheme as pre- 
sented in Fig. 1 for classifying the trees of all the groups may be 
formulated. 



From this scheme any tree of any size with any number of lesions may 
be classified according to the relative amount of infection present. On 
a tree size I, for example, one to ten lesions would be classed as mild 
infection, ten to twenty as moderate infection, and more than twenty as 
severe infection. As may be noted in Fig. 1, the ratio of the average 
number of lesions for any sized tree for the four degrees of infection is 
5:3:1:0. If, then, we multiply the number of trees classed as having 
severe infection by 5, moderate infection by 3, mild infection by 1, and 
no infection by 0, take the total of these products and divide by the 
number of trees considered, a unit is obtained by which the relative 
susceptibility of any group may be fairly and quite accurately compared 

iThe CocciNEAE did not include any trees of size V; as a matter of fact there are 
only six trees of this size in the plantation. From actual measurements of the vari- 
ous tree sizes and from the table given above, it was estimated that a tree of size 
V must necessarily have at least 200 lesions to be classed as having severe infection. 



1935] MacLACHLAN, HOSTS OF GYMNOSPORANGIUM GLOBOSUM 105 

with a similarly derived unit for any other group. To illustrate this, 
let us consider a moderately susceptible group, the Macracanthae, and 
a resistant group, the Crus-galli: 
Macracanthae (see Table II): 

Severe infection . . 7 trees X 5 = 35 

Moderate infection . . 10 trees X 3 = 30 

Mild infection . . 78 trees X 1 = 78 

No infection . . 4 trees X = 

Total .. 99 trees 143 



LLi (see Table II): 

Severe infection . . trees 

Moderate infection . . 2 trees 

Mild infection . . 46 trees 

No infection . . 80 trees 

Total . .128 trees 

Susceptibility unit of comparison , 



The groups of Crataegus examined, the number of species examined 
in each group and the number of trees representing these species, the 
numbers of trees classed according to the different degrees of infection, 
and finally the units of comparison, which may now be considered as the 
relative degrees of susceptibility as indicated by natural infection, are 
presented in Table II. These values for the degrees of susceptibility 
have been plotted in Fig. 4. 

Serial artificial inoculations were made at the following stages in the 
foliar development: (a) on April 2i and 24, 1934, at which time very 
little foliage was evident, a few buds had begun to unfurl, the majority 
were just breaking through the winter scales, while in many instances it 
was necessary to part the winter scales and insert the inoculum; (b) on 
May 7 and 8, 1932 and 1934, respectively, at which times (the foliar 
conditions being approximately the same in both years) the leaves in 
practically all cases were in an advanced stage of expansion but were 
still tender, exhibiting relatively little cuticular development; (c) on 
May 22 and 22>, 1933 and 1934, respectively, at which times the leaf 



106 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

cuticle was fairly well developed and most of the trees were in an ad- 
vanced stage of blossom;' (d) on June 28, 1934, when the foliage was, 
for all practical purposes, fully mature and certain of the groups ex- 
hibited a very heavy cuticle on the leaves. - 

The number of species inoculated in each group and the percentage of 
these falling into the different classes of susceptibility for each of the 
four serial inoculations are presented in Table III. The correlation of 
these data will be found under sub-section D. 




1935] MacLACHLAN, HOSTS OF GYMNOSPORANGIUM GLOBOSLM 107 

1. The geographical distribution of Crataegus 
Of the twenty-nine groups as given by Rehder (1927), twenty-three 
are of American origin; the remainder have been introduced from 
Eurasia. With the exception of the Macracanthae, which extend into 
the middle west, and the Douglasianae, which are typically western, the 
American groups, as indicated by the dotted area in Fig. 2, are confined 
to the eastern part of North America. While certain of these groups are 
typically more northern than others they overlap to such an extent that 
no correlation could be made between the distribution and the relative 
susceptibility of the respective American groups. Although none of the 
Eurasian groups proved to be very susceptible, no differences from the 
type of infection produced on American groups could be observed. Con- 
sequently, the distribution of the genus gave no information that proved 
of value in determining the relative susceptibility of the various Cratae- 

2. The role of the foliar cuticle 
By using herbarium material collected in the Arnold Arboretum from 
natural infection in 1932 a detailed comparison was made between one 
of the largest and most resistant groups, Crus-g.alli, and one of the 
largest and very susceptible groups, Tenuifoliae, in an attempt to 
correlate the susceptibility of the host plant with the mechanical struc- 
ture of the leaf. As a check on the results obtained, the Coccineae, 
another very susceptible group, was examined in a similar manner. The 
following observations were made: 

(a) Distribution of lesions on the leal. 

i. Number of lesions primarily associated with mid and main 
lateral veins of the leaf. 

ii. Number of lesions on the chlorenchyma which, lor present 
purposes, may be defined as the leaf area other than that 
occupied by the mid and main lateral veins. 

(b) Spermogonial stage. 

i. Number of spermogonia per lesion, 
ii. Diameter of lesion. 

(c) Aecial stage. 

i. Number of aecial horns per lesion. 

ii. Diameter of lesion producing aecia. 
iii. Length of mature aecial horns, 
iv. Number of lesions actually producing aecia. 

(d) Detailed notes on thickness of foliar cuticle, degree of hyper- 
trophy and amount of leaf-killing. 



JOURNAL OF THE ARNOLD ARBORETUM 



In addition to the above data separate i 

made for chlorenchyma and vein infections in the Coccineae. Table I 

gives the results obtained for these three groups. 

TABLE I 

PRESENTING DATA ON BIOMETRICS AS OBTAINED FROM 

HERBARIUM MATERIAL FOR THREE GROUPS OF 

THE CRATAEGI 



to chlorenchyma ar 
below the pairs. 



A comparison of these data brings out three significant facts: 

( 1 ) Practically all the Crus-galli have thick coriaceous leaves with a 
very heavy cuticle. The Tenuifoliae and Coccineae, on the other 
hand, have thin leaves with little cuticle. This condition was checked 
for all the other groups, and while the thickness of the leaf itself did not 
show consistent correlation with the relative susceptibility of the re- 
spective groups, there was a surprisingly consistent correlation on the 
part of cuticular thickness. Groups that finally fell into the moderately 
susceptible class exhibited an intermediate deposition of cuticle, the 
degree of which varied somewhat in different species within the respec- 
tive groups. All the species within the groups which were classed as 
resistant had consistently heavy cuticle and those classed as very suscep- 
tible had consistently little cuticle. 

(2) The Crus-galli leaves have more than eighty percent of the 
infections on veins, the Tenuifoliae approximately fifty percent and 
the Coccineae about thirty percent. By correlating these data with the 
relative susceptibility of the three groups, it appears that the degree of 



1935] MacLACHLAN, HOSTS OF GYMNOSPORANGIUM GLOBOSUM 109 

susceptibility varies inversely as the percentage of infections primarily 
associated with the main veins. 

(3) Although the Crus-galli exhibit the lowest percentage of trees 
infected, and thus would seem the most resistant, the individual lesions 
on the leaves of this group have the greatest diameter, and the largest 
number of spermogonia and aecia per lesion. 

When these facts are fitted into the picture of the relative suscepti- 
bility of any host tree to the rust, they definitely indicate that the differ- 
ence in susceptibility is purely mechanical, the cuticle being the deciding 
factor. The basidiospores of G. globosum, while able to produce infec- 
tion from the lower surface of the leaf, germinate and gain entrance 
primarily through the upper side. Thus, spores carried by the wind and 
alighting on the smooth waxy surface of the Crus-galli leaf are not so 
liable to adhere, and if they do remain and germinate, a large percentage 
of the germ-tubes die before they can penetrate the heavy cuticle. Many 
instances illustrating this phenomenon occurred during investigations of 
the waxy-leaf types. Within the Crus-galli, for example, a much 
higher degree of susceptibility relative to the groups with non-waxy 
leaves was indicated by artificial inoculation where conditions were 
optimum for the infection process, than by natural infection where the 
basidiospores must necessarily withstand a certain amount of desicca- 
tion before infection can take place. Again, in many cases waxy leaves 
infected by natural inoculation were found on very low branches only, 
that is, branches almost touching the ground. Here the leaves were 
kept cool and moist for longer periods of time by the tall grass that hap- 
pened to be growing around these trees; such an environment afforded 
a better opportunity for spore germination and germ-tube penetration. 

The distribution of lesions on the leaves gives further evidence of the 
cuticle acting as a natural barrier. In the Crus-galli eighty-three per- 
cent of the lesions were primarily associated with the main veins. The 
little grooves over these veins afford lodging places for the basidio- 
spores; moisture tends to remain longer along these areas, rendering a 
more favorable environment for the infection process. When making 
artificial inoculation by painting the leaves with an aqueous suspension 
of basidiospores, it was very difficult to get a film of the suspension to 
lie uniformly over waxy leaves. The water would form into droplets, 
and either roll off the leaf entirely or else remain in the little grooves 
over the veins. One can readily picture the same thing happening when 
the basidiospores are brought naturally. Inoculation usually takes place 
during wet weather, as it is then that the telial flanges on the galls swell 
and the teliospores germinate to produce basidiospores. The latter are 



110 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

then carried aerially, either directly to the Crataegus leaf by the wind, 
or else washed out of the air by falling rain onto the host leaf. Here, as 
in the case of artificial inoculation, the moisture necessary for spore ger- 
mination accumulates in droplets and these either roll off the waxy leaf 
or remain in the grooves over the veins, carrying the basidiospores with 

With a non-waxy leaf we have an altogether different picture. A film 
of water readily spreads over the surface of the leaf in a uniform layer, 
in which case the basidiospores are more apt to remain where they hap- 
pen to alight on the leaf. Here the germinating basidiospores have no 
heavy cuticle with which to contend and can successfully penetrate the 
leaf surface almost as easily at any place over the chlorenchyma as over 
the veins. Since the area occupied by chlorenchyma far exceeds that 
occupied by the main veins, one can readily see why only thirty-one per- 
cent of the lesions on the Coccineae leaves were vein infections as com- 
pared with eighty-three percent on the Crus-galli leaves. 

The fact that within the Crus-galli group the rust flourished even 
better than within the more susceptible groups, producing larger lesions 
with a larger number of spermogonia and aecia per lesion, can also be 
attributed to the relatively high percentage of vein infections. Regard- 
less of leaf type the very large lesions, some seven to ten millimeters 
long, with more than one hundred spermogonia and fifty to one hundred 
aecia per lesion, were vein infections. In the Coccineae measurements 
of vein and chlorenchyma infections were kept separate, in order to ob- 
tain quantitative data on the relative size of the lesions and the number 
of spermogonia and aecia per lesion in the two types of infection. As 
may be seen from the foregoing table, the lesions are much larger in vein 
infections, producing almost twice as many spermogonia and aecia. All 
evidences indicate that G. globosum is capable of establishing a much 
more efficient nutritional regime when in direct contact with one of 
the veins. In the early spermogonial stage of even chlorenchyma infec- 
tions one can see j^ellowish lines of fungal hyphae, radiating out along 
the vascular bundles from the centre of the lesion, as shown in Plate 
125, Fig. 2. Again, in Plate 125, Fig. 1, the infection appears systemic, 
extending the entire length of a lateral vein. Plate 125, Fig. 3 shows a 
main lateral vein infection branching out along one of the sub-lateral 
veins. In fact, in every vein infection observed (eight hundred and 
eighteen), as may be seen in Plate 125, Fig. 4, the lesion was typically 
long and narrow, the long axis corresponding with that of the vein. 

Vein infections appeared to produce aecia later in the season than chlo- 
renchyma infections. ]Many cases were found among the former where 



1935] MacLACHLAN, HOSTS OF GYMNOSPORANGIUM GLOBOSUM 111 

the aecial horns were just emerging or else were very short when the 
leaves were collected, while nearly all the chlorenchyma lesions had fully 
developed aecia, with peridial cells ruptured and aeciospores emerging. 
It would seem, then, that the time of spore production is correlated with 
the availability of food supply. An infection not primarily associated 
with a main vein utilizes all the available nutrient and then produces 
spores. Vein infections, on the other hand, have a greater and more 
lasting nutrient supply from the host, develop more mycelium and, when 
they finally do sporulate, have a greater supply of reserve food to pro- 
duce aecia. Thus chlorenchyma infections produce relatively smaller 
and fewer aecia over a smaller lesion and at an earlier date than vein 
infections. This fact would account for the higher percentage of the 
lesions within the Tenuifoltae and Coccineae actually producing 
aecia at the time the herbarium material was collected. 

Severe leaf killing, where relatively few lesions per leaf were involved, 
was due in practically all cases to infections primarily associated with 
the main veins, the amount of leaf killing depending on how far back 
from the edge of the leaf the vein was attacked. Plate 126, Fig. 2 shows 
one lesion on the mid-vein resulting in the death of over half o'f the leaf. 
On the other hand, in Plate 126, Fig. 1, may be seen a chlorenchyma 
infection where leaf killing extends from the point of infection to the 
margin of the leaf but does not extend beyond the enclosing lateral 
veins. A purely chlorenchyma infection nearer the center of the leaf 
rarely causes killing beyond the area of actual infection. 

If the degree of susceptibility is in any way physiological, one would 
necessarily expect that within the resistant groups the rust would have 
greater difficulty in establishing a satisfactory nutritional regime, and 
if once established would produce small lesions with relatively few fruit- 
ing bodies due to some antagonistic physiological reaction on the part of 
the host. Crowell ( 1934) found such to be the case when he determined 
the relative susceptibility of the genus Malus to Gymnosporangium Juni- 
peri-virginianae Schw. In European species of Malus the lesions were 
very small, in some cases producing a few spermogonia but no aecia. 
Somewhat similar instances were found by the writer in determining the 
relative susceptibility of species of Pyrus to G. globosum. In the Cra- 
taegi a few rare instances were found that might suggest differential 
physiological antagonism on the part of the host. In Plate 126, Fig. 5 
is shown a lesion that produced abundant spermogonia but died be- 
fore any hypertrophy or production of aecia took place; the host tissue 
may have been hypersensitive to the rust mycelium, the latter taking 
such a heavy toll on the nutrient content of the leaf that the host tissue 



112 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

was killed and as a result the fungus died. Plate 126, Fig. 4 illustrates 
a case of leaf killing extending below the area of infection; this suggests 
the existence of a toxic agent secreted by the rust. In a few of the col- 
lections very small lesions not more than a millimeter in diameter that 
never produced even spermogonia were found. In Plate 126, Fig. 3 
may be seen a small lesion that exhibits no hypertrophy and produced 
only one aecial horn. However, such instances as the foregoing were 
rare and not corisistent even on a single host, and may be considered as 
insignificant factors in determining the relative susceptibility of the 
genus Crataegus. Indeed, from examination of the herbarium material 
the writer found the exact opposite to any physiological antagonism on 
the part of the host to be true; G. globosum is apparently able to estab- 
lish itself more satisfactorily in the most resistant groups, due to the 
relatively high percentage of vein infections. This condition would 
indicate that the basis for differences in susceptibility is for the most 
part mechanical, involving primarily the cuticle as the deciding factor. 
The Crus-galli is a difficult group for the rust to invade, except for a 
very short period in the spring before the foliar cuticle has developed to 
any extent. However, once the rust has successfully penetrated this 
cuticle it is just as much at home and can do just as much damage or 
even more in the Crus-galli than it can in the Tenuifoliae, Coccineae 
or any other very susceptible group. 

3. The degree and the duration of the period of susceptibility 

The role of the cuticle also explains the significant phase in the dura- 
tion of the susceptibility of any host. There is a definite duration to 
this period of susceptibility for all the groups, the degree of which rises 
rapidly during the unfurling of the leaves and reaches a maximum dur- 
ing and immediately after the period of leaf expansion, then falls off 
gradually at a rate depending, in part at least, on the rapidity of deposi- 
tion of foliar cuticle. 

In Plate 127, Figs. 1-4 are shown the results obtained from the four 
respective serial inoculations on Crataegus Pringlei. At the time of ini- 
tial inoculation, April 25, 1934, the leaves, approximately one quarter of 
an inch long, had just begun to unfurl and a very small amount of infec- 
tion at the tip of one leaf resulted (Fig. 1 ). The inoculation on May 9, 
after the leaves had fairly well expanded, produced severe infection 
(Fig. 2). Inoculation two weeks later resulted in scattered lesions (Fig. 
3), while the inoculation on June 28 gave negative results (Fig. 4). 

The same phenomenon but from a different approach is evident in 
Plate 128, Figs. 1 and 2, which demonstrate the results obtained from 
inoculations on Crataegus Jonesae on May 7 and June 4 respectively. 



1935] MacLACHLAN, HOSTS OF GYMNOSPORANGIUM GLOBOSUM 113 

All the leaves in both inoculations received approximately the same 
amount of inoculum per unit area of leaf. At the time of the first 
inoculation the five basal leaves were well expanded, while the two upper 
leaves were just beginning to expand. As may be noted in Fig. 1, much 
heavier infection occurred on the older leaves. (The large irregular 
white areas on the younger leaves are holes caused by insects.) In Fig. 
2, showing the results of the later inoculation, the reverse situation is 
seen; on the younger leaves at the end of the twig abundant infection 
was obtained, while the older leaves exhibited only scattered lesions. 

It is quite evident, therefore, that the cuticle cannot be the sole deter- 
mining factor for variation in susceptibility throughout the entire life 
of the foliage; certain physiological factors may also be involved. For 
example, the leaves apparently are not so susceptible during the period 
of emergence from the winter scales until they are in a moderately ad- 
vanced stage of expansion, a period prior to any heavy deposition of 
cuticle. It is possible that the rust is unable to establish itself in the 
very young leaf. However, since this rust is not primarily of a systemic 
nature, probably the dilution effect on the number of lesions resulting 
from the intussusceptional type of foliar growth and consequent expan- 
sion, as well as the relatively small leaf area exposed at the time of 
inoculation, will account for the major part of this phenomenon. Again, 
even the most susceptible groups, for example, Anomalae or Cocci- 
NEAE, are apparently quite resistant to the rust by the latter part of 
June, at which time the leaves have by no means the amount of cuticle 
that is formed on the Crus-galli even in the early part of May. It is 
possible that the rust is unable to establish a nutritional regime in the 
mature leaf as exhibited in the latter part of June, a point in favor of 
assuming a physiological antagonism on the part of the host. The rela- 
tively high temperature may also be a factor, by inhibiting spore 
germination. 

Nevertheless these two periods play an insignificant part in any deter- 
mination of the amount of infection that may accumulate on a host, 
regardless of the group. In the former case the period is relatively short 
and the leaf area exposed to the basidiospores by the unfurling buds 
would be small in comparison with that exposed after the leaves have 
expanded. As for the latter case practically all the teliospores on the 
red cedar have germinated by May 25, and the degree of susceptibility of 
any pomaceous host after the last of May would have no significance in 
determining the amount of infection that might occur. Thus, for prac- 
tical purposes in the field the significant period within which infection 
might take place is between the time the leaves are fairly well expanded 



114 



JOURNAL OF THE ARNOLD ARBORETUM [vol. 



and the end of basidiospore dispersal. During this time the thickness 
and rapidity of deposition of the cuticle are the deciding factors. For 
this reason the inoculations in April and June, respectively, are not con- 
sidered in determining the relative susceptibility of the various groups. 



SCEPTIBILITY PERIOD 



COCCINEAE 

RACANTHAE 
CRUS-GALLl 
ENTIRE GENUS 




DAYS BETV 

^JG THE DeGRE 



EEN INOCULATIONS 






To illustrate further the degree and duration of susceptibility within 
the different groups, values may be obtained for the relative degrees of 
susceptibility of the various groups by taking the sum total of the values 
as expressed by the symbols 0, 1, 2 and 3, and dividing by the number 
of representatives inoculated.^ These were obtained from Table I for 
the CocciNEAE, Macracanthae, and Crus-galli, which are, respec- 
tively, typical of the classes very susceptible, moderately susceptible 

iThe objection arises that such a method of correlation utilizes arbitrary quahta- 
tive symbols to designate quantitative entities. Nevertheless, its usage here is not 
to be considered from a statistical standpoint and it does present a clearer picture 
to illustrate both the degree and the duration of the period of susceptibility within 
any one group of Crataegus. It is interesting to note that if such a method be 
employed in correlating the data obtained from serial inoculations in this genus 



1935] MacLACHLAN, HOSTS OF GYMNOSPORANGIUM GLOBOSUM 115 

and resistant, and have been plotted in Fig. 3. A similar curve (in 
heavy line) is given for all the inoculated representatives of the genus. 
The area enclosed by the respective curves would, to a certain extent, 
be a measure of both the degree of susceptibility and its duration. The 

TABLE II 

PRESENTING DATA ON THE RELATIVE SUSCEPTIBILITY 

OF CRATAEGUS TO G. GLOBOSUM, AS INDICATED 

BY NATURAL INFECTION 









No. 


trees in 


theCasses 






















Sps. 


Trees 


iJ^t. 


^L 


infect 


infect. 


susceptibility 


\nomalae 


1Q 


40 


^ 


1Q 


, 


18 


2 95 


Azaroli 


1 


1 













.00 


Bracteatae 


2 






1 








.50 


Coccineae 


46 


82 




24 


20 


36 


3.22 


Crus-galli 


71 




80 


46 







.41 


Dilatatae 


4 






6 





4 


2 36 


Douglasianae 


8 


19 




16 








.84 


Flavae 


10 




11 











.00 




10 


Jo 









^ 


.27 


Macracanthae 




Microcarpae 




J 













Holies 


37 


86 


10 




11 


25 


2 30 


Nigrae 


2 


2 




2 








1.00 




















2 


4 




4 








1.00 


Pruinosae 


58 




36 


57 


3 




.67 




34 


37 




26 




2 












37 






1.45 


Sa^nguinae 


4 


4 


Q 




Q 





1.00 


Tenuifoliae 

Triflorae 


35 

7. 


57 
2 




42 
104 



8 
32 



4 

32 


1.51 


Uniflorae 


I 


2 













00 




18 


30 


20 




' 












Coccineae, characterized by little foliar cuticle, exhibit a much higher 
degree of susceptibility over a relatively longer period of time than the 
Crus-galli which have consistently heavy cuticle on the leaves, whereas 
the Macracanthae, with an intermediate and varying amount of cuticle, 
assume an intermediate position. 



JOURNAL OF THE ARNOLD ARBORETUM 



•Cra 



Bearing in mind that the thickness of the cuticle and its rapidity of 
deposition on the leaves are the primary factors in determining the rela- 
tive susceptibility of any host, while geographical distribution and 
physiological antagonism on the part of the host play a very minor part, 
if any, it is now possible to evaluate. the data obtained by the two pre- 
viously described methods of approach and determine the relative 
susceptibility of the various groups within the genus Crataegus. 



The relative degrees of susceptibility obtained from observations of 
natural infection, as previously stated, are presented in Table II, and 
have been plotted in Fig. 4. In regard to data obtained by serial inocula- 
tions, it is quite obvious from Table III that inoculations before the 
leaves unfurl, and again late in June, result in very little infection. 
However, as the foregoing discussion on the duration of the period of 



1935] MacLACHLAN, 



iTMNOSPORANGIUM GLOBOSUM 117 



susceptibility demonstrates, such a phenomenon, while interesting, plays 
an insignificant role in determining the amount of infection that might 
take place on any tree. The two significant inoculations are those made 



PRESENTING DATA ON THE RELATIVE SUSCEPTIBILITY 

OE THE GENUS CRATAEGUS TO G. GLOBOSUM AS 

INDICATED BY SERIAL INOCULATIONS 




in May, (b) and (c), and for fifteen of the major groups the percentage 
frequency of occurrence of inoculated representatives falling into the 
respective classes of susceptibility have been plotted in Fig. 5 (p. 118). 
In comparing these tables and figures to make a final classification 
of the groups according to their relative susceptibility, one must remem- 
ber that these results were obtained from two altogether different meth- 



118 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

ods of approach. For those groups the representatives of which have a 
heavy cuticle, a much lower degree of susceptibility would be indicated 
by natural infection than by artificial inoculation where the amount of 
inoculum and the cultural environment are optimum. The number of 
representatives examined in each group, and especially for natural in- 
fection, must also be given consideration. 



:LlJ:LUU:L'k 

"lu":i!i':£i:iLC 



Fig. 5. Relative Susceptibility of Fifteen Groups of the Genus 

The results of two inoculations, (b) and (c) respectively, are presented 
in each sub-graph. The numbers on the abscissae of the sub-graphs refer 
to the classes of susceptibility. The numbers in parentheses refer to the 
number of species (with the exceptions noted in text) inoculated in each 
group. 

By correlating the degree of susceptibility as indicated by natural 
infection, and the frequency of occurrence of inoculated representatives 
falling into the various classes of susceptibility, the groups may be classi- 
fied and arranged within each class according to susceptibility as 
follows:' 
V^ery susceptible— Anomalae, Coccineae, Tenuifoliae, Dilatatae. 



1935] MacLACHLAN, HOSTS OF GYMNOSPORANGIUM GLOBOSUM 119 

Moderately susceptible — Molles, Macracanthae, Rotundifoliae, 

PUNCTATAE, DOUGLASIANAE, SiLVICOLAE, PrUINOSAE, ViRIDES, 

Flavae, Oxyacanthae, Intricatae. 
Resistant — Crus-galli, Bracteatae, *Azaroli, *Microcarpae, 
*NiGRAE, *Pinnatifidae, *Sanguineae, *Triflorae, *Uni- 



Immune — None. 

None of the groups examined proved to be wholly immune. No in- 
fection was obtained on the one representative of the Micrgcarpae, 
namely, C. Phaenopyrum (L. f.) Medic. (= C. cordata Ait.), but this 
species has been previously reported as a host to the rust from both 
Delaware and Tennessee. Of almost five hundred and fifty determined 
species and varieties studied, less than one percent of the artificial in- 
oculations gave negative results and it is indeed possible that, given 
optimum conditions for germ-tube penetration, not a single species 
could be considered totally immune. However, as previously stated, it 
must be remembered that the conditions favorable for infection set up 
by artificial inoculation far exceed any that might occur in nature, and 
many species that are now classed as suscepts would probably never 
exhibit infection under field conditions. 



The best guide in the selection of Crataegus trees to be planted on 
estates where G. globosum is in the vicinity would be the thickness of 
the foliar cuticle. A striking example of this was found on an estate at 
Canton, Massachusetts, where two Crataegus trees, one a Coccineae 
species and the other a Crus-galli species, were planted side by side, 
surrounded by red cedars bearing heavy infections of G. globosum. 
These have been under observation for the past three years, and each 
season the foliage on the Coccineae species has suffered very severe 
infection, resulting in more than eighty percent defoliation by the latter 
part of August. The tree is now in a very weakened condition. The 
Crus-galli species, on the other hand, has been entirely unaffected by 
this rust. 

In choosing from species of American origin one should definitely 
avoid the Anomalae, Coccineae, Tenuifoliae and Dilatatae if G. 
globosum be in the vicinity. Certain of the species within the groups 

iThe small number of representatives in the resistant groups indicated by asterisks 
made it impossible to arrange these groups within the class "Resistant" according 
to susceptibility and they have been arranged alphabetically. 



120 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

classed as moderately susceptible have considerable cuticle on the 
leaves and these may be planted with a relative degree of safety. The 
Crus-galli, however, are very resistant, and offer a wide variety of 
species. They are, as Rehder (1927) states, handsome ornamentals 
with dense, dark green foliage which remains till late in autumn or early 
winter, and are very attractive in bloom, with decorative bright red 
fruits that are persistent during the winter. If one desires the Eurasian 
type, the Pinnatifidae offer a group with lustrous leaves and large 
showy fruit. Some varieties of these are cultivated in northern China 
for the edible fruit. The Oxyacanthae will also withstand severe in- 
fection unless under abnormal proximity to Juniperus rusted by G. 
globosum, with C. Oxyacantha Jacq. including some of the most showy 
garden forms. 

This presentation has been confined to foliar lesions, and while infec- 
tion has been obtained on all parts of the flower as well as the fruit and 
young twigs, such instances were sufficiently rare that they were not 
worthy of consideration at this time and have been set aside for a sec- 
ond publication on the life history of G. globosum Farl. 

No consideration has been given to the possibility of variation in 
virulence within different strains of this rust. Practically all the inocu- 
lum was obtained from two adjacent red cedar trees at Waltham, Massa- 
chusetts. 

One must also bear in mind that the relative susceptibility of groups 
within the genus Crataegus to G. globosum is in no respect correlated 
with their susceptibility to other Gymnosporangium rusts. Crowell 
(unpublished) has found, for example, that the Crus-galli, so resist- 
ant to G. globosum, are quite susceptible to G. clavipes Cke. & Pk. 

Studies on relative susceptibility within the genus Pyrus were con- 
fined to the results obtained from serial inoculations made in 1934. The 
species represented in the Arboretum were artificially inoculated in a 
manner similar to that described for Crataegus: (a) on April 25, at 
which time the condition of the foliage varied from buds just bursting 
through the winter scales to leaves a quarter to a half inch long; (b) on 
May 9, when the leaves were fairly well expanded on all species; (c) on 
May 22 when the foliage was fully expanded; and (d) on June 28. Cer- 
tain of the species which had given negative results in the previous in- 
oculation were omitted in the June inoculation. 

In Table IV are given the species inoculated, their distribution, the 
degree of infection obtained on the respective dates of inoculation, the 



1935] MacLACHLAN, HOSTS OF GYMNOSPORANGIUM GLOBOSUM 121 

stages of the rust produced on the foHage, and finally, a classification of 
their relative susceptibility. 

TABLE IV 

PRESENTING DATA ON THE RELATIVE SUSCEPTIBILITY 

OF SPECIES OF THE GENUS PYRUS TO G. GLOBOSUM, 

AS INDICATED BY SERIAL INOCULATIONS 

Deg. suscept. 

indicated by 



P. Balansae Decne. 


Eurasian 


P. betulaefoiia Bge. 


Eurasian 


P. Bretschneideri Rehd. 


Eurasian 


P. communis L. 


Eurasian 


P. elacaKrifolia Pail. 


Eurasian 


P. Korshinskyi Litv. 


Eurasian 


P. Michauxii Bosc' 


— 


P. Lindlcyi Rehd. 


Eurasian 


P. nivalis Jacq. 


Eurasian 



P. ussuriensis Maxim. Eurasian 1 1 - & 1 1 

^P. Michauxii is a 'hybrid {P. amygdaliformis X P. nivalis). 

No consistent correlation between the relative susceptibility of the 
various species and the type of leaf is evident; all species have consider- 
able cuticle on the foliage, and a few are somewhat tomentose. Nor can 
the differences in susceptibility be correlated with the distribution of the 

The lesions in general were found to be much smaller than those ex- 
hibited on Crataegus, and except in the case of P. betulaefoiia rarely 
measured more than one to two millimeters in diameter. Certain spe- 
cies, designated in the table, showed spermogonia only ; the lesions were 
extremely small, and died before any hypertrophy or aecial formation 
was evident. However, it is possible that with a different strain of the 
rust some of these might produce aecia; P. communis, for example, ex- 
hibited only spermogonia in my inoculations but has been reported pre- 
viously from seven different states. 

As in Crataegus, there is a definite duration to the period of suscepti- 
bility, the degree of which reaches its maximum during and immediately 
after the period of foliar growth and expansion, and then falls off gradu- 
ally so that by the end of June all species examined are immune. 



122 JOURNAL OF THE ARNOLD ARBORETUM I vol. xvi 

Classified according to their relative susceptibility, the species exam- 
ined may be arranged (alphabetically) as follows: 
Very susceptible — P. betulaejoUa Bge. 

Moderately susceptible — P. Balansae Decne., P. serotina Rehd. 
Resistant — P. Bretsckneideri Rehd., P. communis L., P. Korshinskyi 

Litv., X P. Michauxii Bosc, P. Lindleyi Rehd., P. nivalis Jacq., 

P. phaeocarpa Rehd., P. scrrulata Rehd., P. syriaca Boiss., P. ussuri- 

ensis Maxim. 
Immune— P. amygdalijormis X'ill, P. clacagrijolia Pall., P. salicijolia 

Pall. 
Previous reports of Pyrus suscepts are confirmed, for the most part, 
to P. communis, to the Kieffer Pear (P. communis X P- serotina) 
and other varieties used commercially in the orchard. Stevens and Hall 
(1910) report G. globosum as being particularly abundant on the Japa- 
nese strain of pear (P. serotina), while Stewart (1910) reports the Kief- 
fer pear as suffering infection from this rust at Long Island, New York. 
In particular he finds that both the fruit and leaves are attacked, and 
that the diseased fruits are very small and misshapen, usually exhibiting 
circular black areas devoid of aecia, although a few show aecia. On the 
other hand, Stewart (1910), and Hesler and Whetzel (1917) classify 
the Bartlett, Bosc, Duchess, and Worden varieties as being for the most 
part immune, although the fruit of the Worden variety is subject to 
infection. 

While little can be added to the knowledge of the relative suscepti- 
bility of the orchard varieties, one may conclude from the foregoing 
classification that, with the e.xception of P. betulaejoUa, P. Balansae, P. 
serotina, and as indicated from previous reports, P. communis, the re- 
mainder of the species can be safely planted in vicinities where the rust 
is present. This conclusion holds true especially for P. amygdalijormis, 
P. elaeagrijolia, and P. salicijolia. 



Serial artificial inoculations were made in 1934 on species and vari- 
eties of Sorbus available in the Arnold Arboretum: (a) on April 25, at 
which time the foliar buds were just beginning to break open and the 
tiny leaves in many cases exhibited a heavy tomentose covering which 
was removed without injury to the leaf by rubbing the latter between 
the fingers, and the inoculum was placed on the exposed green tissue: 
(b) on May 9, at which time practically all the foliage was going 
through a period of rapid growth and expansion: (c) on May 24, at 
which time the leaves were fully expanded (blossoms where present were 



1935] MacLACHLAN, HOSTS OF GYMNOSPORANGIUM GLOBOSUM 123 

also inoculated) ; (d) on June 28, at which time the leaves for all prac- 
tical purposes were mature. 

The results of these inoculations appear in Table V which presents, 
where positive results were obtained, the species and varieties inocu- 
lated, their native distribution, the degree of infection obtained from 
the respective inoculations, the stages of the rust exhibited, and finally 
the resultant classes of susceptibility. 



OF SPECIES AND VARIETIES OF THE GENUS SORBUS TO 

G. GLOBOSUM, AS INDICATED BY SERIAL 

INOCULATIONS 



Species and varieties 



var. nana Hort 



No infection was obtained on the following (alphabetically arranged) 
species and varieties, which are all of Eurasian origin: Sorbus alnijolia 
K. Koch, S. amurensis Koehne, S. Aria Crantz, 5. Aria var. angustifolia 
Hort., S. Aria var. Decaisneana Rehd., S. Aria var. longijolia Pers., S. 
Aria var. lutescens Hartwig, S. Aria var. magnifica Hort., S. Aria var. 
salicijolia Myrin, S. Aria var. sulphurea Hort., 5. Aria var. theophrasta 
Hort., 5. Aucuparia L., S. Aucuparia var. Dirkenii aurea Hort., 5. 
Aucuparia var. edulis Dieck, 5. Aucuparia var. nana Hort., S. Aucu- 
paria var. xanthocarpa Hartw. & Ruempl., S. commixta Hedl., S. com- 
mixta var. rujo-jerruginea Schneid., 5. discolor Hedl., X 5. hybrida L., 
5. intermedia Pers., X S. latijolia Pers., S. latijolia var. atrovirens Hort., 
S. Matsumurana Koehne, X S. Meinichii Hedl., S. pohuashanensis 
Hedl., S. Zahlbruckneri Schneid. 



124 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

All species of American origin that were inoculated proved to be 
susceptible, with 5. americana as the only species on which the foliage 
was materially injured by the rust. Of the thirty-one inoculated Eura- 
sian types, infection was obtained on only four, and these proved to be 

The lesions in all cases were very small, rarely measuring more than 
one to two millimeters in diameter, with an average of three to five 
aecial horns per sorus. Those species on which spermogonia only were 
obtained (see Table V) exhibited bright yellow lesions until the spermo- 
gonia were mature, following which time no further development took 
place and the infections died. An interesting type of natural infection 
was observed on Mt. Monadnock in New Hampshire; the lesions were 
as large as any ever obtained on Crataegus, some being as much as ten 
to twelve millimeters long, each bearing abundant aecial horns. Whether 
this type of infection results from a more susceptible variety of S. 
americana, or another strain of G. globosum, is not known. 

With the exception of S. americana, no infection was obtained on any 
of the species after the second inoculation, while practically all the sus- 
cepts exhibited some infection from the initial inoculation. It would 
seem, therefore, that the resistant forms at least are most susceptible 
during, and immediately after, the period when the foliar buds are 
unfurling; S. americana, however, reached its maximum degree of sus- 
ceptibility immediately after the leaves had expanded. 

It is extremely doubtful that, with the exception of 5. americana and 
its varieties within the American types, and possibly the hybrid Eura- 
sian type, 5. thuringiaca, any representative of the genus Sorbus would 
be seriously alTected by G. globosum regardless of proximity to the rust. 
This is certainly true for the species of Eurasian origin. 
Malus— Rklative Susceptibility as Indicated by Serial Inocu- 

Serial artificial inoculations were made in 1934, similar to those de- 
scribed for the preceding genera: (a) on April 24, at which time the 
leaves had already unfurled and were undergoing the period of rapid 
expansion; (b) on May 9, at which time the foliage was almost mature 
size, and most of the blossoms were in the pink stage; (c) on May 22, 
at which time most of the petals had dropped. No inoculation was made 
in June. Table VI presents the species on which positive results were 
obtained, the origin of the various species, the results obtained from the 
respective serial inoculations, the stages of the rust obtained, and finally 
the relative degree of susceptibility. 



1935] MacLACHLAN, HOSTS OF GYMNOSPORANGIUM GLOBOSUM 125 

TABLE VI 

PRESENTING DATA ON THE RELATIVE SUSCEPTIBILITY 

OF SPECIES AND VARIETIES OF THE GENUS AIALUS TO 

G. GLOBOSUM, AS INDICATED BY SERIAL 

INOCULATIONS 

Deg. suscept. 

Native inoculations Stages Degree 

Species and varieties distrib. (a) (b) (c) found suscept. 

M. astracanica Dum.-Cours.i Eurasian 10 & 1 1 

M. baccata Borkh. Eurasian 12 & 1 2 

M. coronaria Mill. American 10 1 

M. Dawsoniana Rehd.2 Hybrid Oil & 1 1 

M. ioensis var. plena Rehd. American 12 1 & 1 2 

M. magdeburgensis Schoch^ Eurasian 10 1 

M. Soulardi Britt.* Hybrid - 2 & 1 2 

M. sublobata Rehd.''' Eurasian 10 & 1 1 

^M. astracanica is a hybrid (Af. prunifolia X M. putnila) . 

-M. Dawsoniana is a hybrid {M. fusca X M. putnila). 

■5 AT. magdeburgensis is a hybrid (M. pumila X M. spectabilis) . 

*M. Soulardi is a hybrid {M. ioensis X M. patmla) . 

-'M. sublobata is a hybrid (M. prunifolia X M. Sieboldii). 

The following species, alphabetically arranged according to distri- 
bution, gave negative results: 

American distribution: Malus angustijolia Michx., M. bracteata 
Rehd., M. fusca Schneid., M. glaucescens Rehd., M. ioensis Britt., M. 
lancijolia Rehd., M. platycarpa Rehd. 

Eurasian distribution: X Malus arnoldiana Sarg., M. asiatica Nakai, 
X M. atrosanguinea Schneid., M. brevipes Rehd., M. floren- 
tina Schneid., M. floribunda Sieb., M. Halliana var. Park- 
manii Rehd., X M. Hartwigii Koehne, M. honanensis Rehd., M. 
kansuensis Schneid., M. micromalus Mak., M. hupehensis (Pamp.) 
Rehd. {= M. theijera Rehd.), M. pumila Mill., M. prunifolia Borkh., 
X M. purpurea var. Eleyi Rehd., X M. robusta Rehd., M. Sargenti 
Rehd., M. Sieboldii Rehd., M. sikkimensis Koehne, M. spectabilis 
Borkh., M. sylvestris Mill., M. toringoides Hughes, M. Tschonoskii 
Schneid., M. yunnanensis var. Veitchii Rehd., X M. zumi Rehd. 

A variety of an American species, M. ioensis var. plena, and the hy- 
brid M. Soulardi proved to be moderately susceptible to G. globosum, 
while two species, M. coronaria, and M. glabrata, and the hybrid M. 
Dawsoniana, may be classed as mildly susceptible. On the remainder of 
+he American species inoculated no infection could be observed; never- 
theless, Thaxter (1889) obtained aecia on M. pumila Mill. {= M. 



126 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

Malus Britt.). Of all the Eurasian species inoculated only one proved 
to be moderately susceptible, namely M. baccata, and three hybrids 
between Eurasian species, M. astracanica, M. magdeburgensis and M. 
sublobata, proved to be mildly susceptible. 

Although a higher percentage of the American species proved to be 
susceptible, no outstanding correlation could be observed between rela- 
tive susceptibility and geographic distribution. Nor can susceptibility 
be correlated with the type of leaf or type of infection produced. In all 
cases the lesions were small; they were rarely more than one to two 
millimeters in diameter. 

The serial inoculations indicated a definite duration to the period 
of susceptibility which reaches a maximum about the time the blossoms 
are in the pink stage, and falls off to almost zero within a period of two 

Excluding the species found to be susceptible it is very doubtful that 
any of the remaining species considered would suffer from the rust re- 
gardless of proximity to red cedars infected by G. globosum. 

Previous reports would indicate that the commercial varieties of apple 
are more susceptible than the above ornamental types. Bliss (1931) 
using telial material from Iowa culturally obtained flecking on the vari- 
eties Baldwin, Delicious, Fameuse, Greening, Mcintosh, Tolman, 
Wealthy, Yellow Transparent, and York Imperial. From reports of 
Clinton (1934), Thomas and Mills (1930), Sherbakoff (1932), Miller, 
Stevens and Wood (1933), and others, the relative susceptibility of the 
commercial varieties of apple may be classified as follows: 

Varieties on which moderate to severe infection has been observed: 
Fallawater, Fameuse, Hubbardston, Northwestern Greening, Rhode 
Island Greening, and Wealthy. 

Varieties reported susceptible: Baldwin, Cortland, Esopus, Spitzen- 
burg, Fall Pippin, Gano, Golden Delicious, Jonathan, Mcintosh, New- 
ton, Northern Spy, Pewaukee, Rome Beauty, Russett, Stark, Tolman 
Sweet, Tompkins King, Wagener, Winesap, and York Imperial. 

Resistant variety: Ben Davis. 
Amelanchier^ 

Farlow (1885) obtained spermogonia on leaves of Amelanchier cana- 
densis Med. and Harshberger (1902) lists the same species as a suscept 
to G. globosum, exhibiting both spermogonia and aecia. Stone (1908) 
lists A. alnijolia- as a suscept from Alabama. The following species and 
1 Relative susceptibility in this and the following genera was determined by non- 



1935] MacLACHLAN, HOSTS OF GYMNOSPORANGIUM GLOBOSUM 127 

varieties of Amelanckier were inoculated early in May, 1933: Amelan- 
chier amabilis Wieg., A. asiatica Endl., A. Bartramianu Roem., A. 
Bartramiana X A. laevis, A. canadensis Med., A. florida Lindl., X A. 
grandi flora Rehd., A. humilis Wieg., A. humilis X A- sanguinea, A. 
intermedia Spach, ^. laevis Wieg., A. oblongifolia Roem., A. ovalis Med., 
A. sanguinea DC, A. sera Ashe, A. spicata K. Koch, A. stolonijera Wieg. 
All the inoculations gave negative results. 

No reports can be found indicating that any of the spjecies and vari- 
eties of Amelanckier are very susceptible to G. globosum. 

Cydonia 

Thaxter (1888) by culture obtained spermogonia on Cydonia oblonga 
Mill. {= C. vulgaris Pers.). Cook (1913) reports G. globosum as being 
of common occurrence on quince in New Jersey. Harshberger (1902), 
Clinton (1904), and Gussow (1915) report this rust on quince from 
two other states and from the Niagara Peninsula. Cydonia oblonga, 
inoculated by the writer in early :\Iay, 1933, proved to be moderately 
susceptible to G. globosum, producing both spermogonia and aecia. 
None of the varieties of Cydonia oblonga was inoculated, and no in- 
formation can be given with respect to their relative susceptibility. 

The remaining smaller genera were artificially inoculated and the 
results from these inoculations may be summarized and tabulated as 
follows: 
Comptonia 

Comptonia aspleniijolia Ait.— immune. 

Crataeg-omespilus 

Crataegomespilus grandi flora Bean {Crataegus Oxyacantha X Me- 
spilus germanica)— very suceptible; both spermogonia and aecia ob- 
tained; severe leaf killing resulted. Natural infection was also observed. 

Mespilus 

Mespilus germanica L.— moderately susceptible; both spermogonia 



128 JOURNAL OF THE ARNOLD ARBORETUM 

Sorbus /IWfl)— resistant; exhibited spermogonia only. 
bunda X Sorbus Aucuparia — no infection obtained. 
Sorbopjrrus 

Sorbopyrus auricularis Schneid. {Py} 
resistant; exhibited spermogonia only. 



To our present knowledge of the relative susceptibility of Junipcrus 
little can be added by the writer. From previous reports, including 
those of Adams (1919), Arthur (1926) (1927), Bliss (1933), Claassen 
(1897), Connors (1934), Hunt (1926), Kern (1929), Martin (1922) 
(1925), Stone (1909), and others, and from an examination of the 
material in the Farlow Herbarium and the herbarium of Professor J. H. 
Faull, the host list includes at least six species of Juniperus, and at 
least four varieties of Juniperus virginiana. These have been presented 
in the subsequent host list. 

It may be added here that Martin (1922) lists Larix species as hosts 
to G. globosum from nine states. No infection by this rust has ever 
been observed on Larix in the Arnold Arboretum. 

Juniperus virginiana is the most common telial host throughout the 
eastern and central part of North America, having been reported from 
twenty-five states and from Ontario. Severe infection may occur, as 
exemplified at the Morton Arboretum, Lisle, Illinois and from many 
estates and nurseries surrounding Boston. The writer has observed 
trees that were killed by the abundance of galls present. Other trees, 
while not killed, were disfigured to such an extent that they were no 
longer of ornamental value and had to be removed. Juniperus scopu- 
lorum has also been reported as suffering from infection by G. globosum 
at the Morton Arboretum. 

As far as the eastern and central part of North America are concerned 
no information to date would indicate that any species other than Juni- 
perus virginiana and Juniperus scopulorum and their varieties would 
suffer to any extent from infection by G. globosum. 

IV. THE HOSTS OF GYMNOSPORANGIUM GLOBOSUM FARL. 
The following list includes as far as can be ascertained all the known 
hosts of G. globosum. The hosts have been arranged alphabetically by 
genera and their included species. Within the genus Crataegus the 
species and varieties have been arranged within their respective groups. 



1935] MacLACHLAN, HOSTS OF GYMNOSPORANGIUM GLOBOSUM 129 

Following each host name in parentheses are symbols which may be 
defined as follows: 

a — as obtained by inoculations made by the writer; the inclusion 

of an author's name and reference indicates that this host has 

been determined previously by inoculation, 
n — as determined by observations of natural infection made by 

the writer. 
The inclusion of the abbreviated name of a State implies that 

this species has been reported previously as a host from that 

State. 
All new hosts submitted would necessarily be records for the 

State of Massachusetts, as all studies were made in the 

Arnold Arboretum, Boston. 

HOSTS FOR THE & 1 STAGE 

AMELANCHIER: 

Amelanckier alnijolia Nutt. (Ala.),' A. canadensis Med. (Thaxter 

[1885]; Penn.). 
CRATAEGOMESPILUS : 

Crataegomespilus grandifiora Bean (a; n). 
CRATAEGUS (by groups): 
Anomalae: 

Crataegus affinis Sarg. (a; n), C. asperijolia Sarg. (a; n; Vt.), C. 

Brockwayae Sarg. (a; n), C. Coleae Sarg. (n), C. cyclophylla Sarg. 

(a; n; Vt.), C. Dunbari Sarg. (a; n), C. Egglestonii Sarg. (a; n; 

N. Y., Vt.), C. errata Sarg. (a; n), C. honesta Sarg. (n), C. Ideae 

Sarg. (n), C. improvisa Sarg. (n), C. misella Sarg. (n), C. pinguis 

Sarg. (n; Mich.), C. putata Sarg. (n), C. repulsans Sarg. (n), C. 

Saundersiana Sarg. (n), C. scabrida Sarg. (a; n; Vt.), C. shirley- 

cnsis Sarg. (a; n), C. urbana Sarg. (n). 

AZAROLI: 

Crataegus Heldrekhii Boiss. (a), C. tanacetijolia Pers. (N. Y.). 

Bracteatae: 

Crataegus Ashei Beadle (a; n), C. Harbisonii Beadle (a; Tenn.). 

Coccineae: 

Crataegus acclivis Sarg. (n), C. arcuata Ashe (n; Penn.), C. assur- 
gens Sarg. (a; n), C. aulica Sarg. (n), C. caesa Ashe (n), C. cAi> 
pewaensis Sarg. (n), C. conjiww Sarg. (n), C. cons pec ta Sarg. (n). 



130 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

C. contigua Sarg. (n), C. cristata Ashe (n), C. Dayana Sarg. (n), 
C. delecta Sarg. (n; 111.). C. densiflora Sarg. (n), C. Eamesii Sarg. 
(n; Conn.), C. elongata Sarg. (n), C. fluviatilis Sarg. (a; n), C. 
/re/a//j Sarg. (n; Conn.), C. Hillii Sarg. (n), C. Holmesiana Ashe 
(a; n; Conn., N. Y., Vt.), C. Holmesiana var. tardipes Sarg. (n), 
C. Holmesiana var. 't^///i>e5 Ashe (n), C. /rra5c Sarg. (n), C. /en/o 
Ashe (n), C. lobulata Sarg. (n), C. Macounii Sarg. (n), C. miranda 
Sarg. (n), C. neolondinensis Sarg. (n; Conn.), C. pedicellata Sarg. 
(a; n), C. pedicellata var. gloriosa Sarg. (n), C. perrara Sarg. (n), 
C. /'o/Z^fl Sarg. (n; previously reported, state not given), C. />o//7a 
var. Tatnalliana (Sarg.) Eggl. (Mo., N. Y.), C. Prm^/^; Sarg. (a, 
Arthur [1907]; n; Conn., Ind., N. Y.), C. pura Sarg. (n), C. 
sejuncta Sarg. (n), C. sertata Sarg. (n), C. Thayeri Sarg. (n), C. 
uticaensis Sarg. (n), C. t'n)?Wo Sarg. (n). 
Crus-galli: 

Crataegus algens Beadle (a; n), C. arborca Beadle (a; n), C. crrfw- 
ewwce Sarg. (a; n; Ind.), C. armata Beadle (a), C. or^c Beadle (a), 
C. attenuata Ashe (a; n), C. barbata Sarg. (a), C. barrettiana 
Sarg. (a), C. Bartramiana Sarg. (a), C. 6e///ca Sarg. (a), C. cc/o- 
M^^fl Sarg. (a), C. C(/;76>'/ Sarg. (a; n), C. cerc^ma Sarg. (n), 
C. consueta Sarg. (a; Mo.), C. crus-galli L. (a, Thaxter [1891]; 
n; Ind., Ky., Maine, Mass., Miss., Mo.. N. Car., Ohio, Penn., Tenn., 
Va.), C. crus-galli var. arbutijolia Hort. (a), C. crus-galli var. 
exi^MG (Sarg.) Eggl. (n), C. crus-galli var. pyracanthijolia Ait. 
(a: n), C. crus-galli var. r«6en5 Sarg. (a), C. cjjer^a Sarg. (a), C. 
effulgens Sarg. (a), C. Engelmannii Sarg. (a; n; Mo.), C. erec/c 
Sarg. (a; n), C. Farwellii Sarg. (a; n), C. /ecMnrfa Sarg. (n), C. 
Fontanesiana (Spach) Steud. (a; n), C. geneseensis Sarg. (a), C. 
Aawa^c Sarg. (a), C. /r/r/e//a Sarg. (a), C. w/e5^a Sarg. (a; n), 
C. insignis Sarg. (a), C. jasperensis Sarg. (a), X C. Lavallei 
Herincq (a; n), C. lawrencensis Sarg. (a), C. leptophylla Sarg. (a; 
n), C. livoniana Sarg. (a; n), C. wccra Beadle (a), C. MoAr?'/ 
Beadle (a; n; Ga.), C. munita Sarg. (a), C. pachyphylla Sarg. 
(a), C. Palmeri Sarg. (a; n), C. paradoxa Sarg. (a), C. par ci flora 
Sarg. (a; n), C. Pardee Sarg. (a), C. Penny packeri Sarg. (a; n), 
C. peoriensis Sarg. (n), C. permera Sarg. (a; n), C. persimilis Sarg. 
(n), C. persistens Sarg. (a; n), C. phlebodia Sarg. (a; n), C. />///■- 
/ero Sarg. (a), C. polyclada Sarg. (a), C. rega//^ Beadle (a; n), 
C. Reverchonii Sarg. (Tex.), C. ni^o/w Sarg. (a; n), C. robusta 
Sarg. (a; n), C. ro^wnJc Sarg. (a), C. rubrijolia Sarg. (a; n), C. 
r«G?75 Sarg. (a), C.^etosa Sarg. (a), C. ^et'crc Sarg. (a). C. signata 



1935] MacLACHLAN, HOSTS OF GYMNOSPORANGIUM GLOBOSUM 131 

Beadle (a), C. sinistra Beadle (a), C. sublobtdata Sarg. (a; n), 
C. tardiflora Sarg. (a), C. tetrica Beadle (a; Tenn.), C. trium- 
phalis Sarg. (a; n), C. uniqua Sarg. (a), C. vallicola Sarg. (a; n), 
C. villijiora Sarg. (a), C. Wilkinsoni Ashe (a). 

Dilatatae: 

Crataegus cocanioides Ashe (a; n; Mo.), C. dilatata Sarg. (= C 
coccinioides var. dilatata [Sarg.j Eggl.) (a; Mass., N. Y., Penn., 
Vt.), C. durobrivensis Sarg. (n), C. hudsonica Sarg. (n). 

Douglasianae: 

Crataegus Colorado Ashe (n), C. columbiana Howell (a), C. Doug- 
lasiiUndX. (a, Farlow [1885J ; n), C. Douglasii f. 6arfm Sarg. (n), 
C. Douglasii var. Suksdorfii Sarg. (n), C. erythropoda Ashe (n), 
C. P«>n Britt. (a), C. rivularis Nutt. (n). 

Flavae: 

Crataegus arrogans Beadle (a), C. colonica Beadle (a), C. rfi5/>ar 
Beadle (a; S. Car.), C. e//i/>^icfl Ait. (a), C. frugiferens Beadle (a), 
C. ignava Beadle (a; n), C. impar Beadle (a), C. insidiosa Beadle 
(a), C. linuata Beadle (a), C. visenda Beadle (a). 

Intricatae: 

Crataegus apposita var. Bissellii (Sarg.) Eggl. (a; Conn.), C. 6i/^- 
moreana Beadle (Mo.), C. Boyntonii Beadle (N. Car.), C. 5mc^- 
/e>'i Beadle (a; N. Car.), C. Z>e/o5« Sarg. (a), C. /oe^^f/a Ashe 
(a), C. fortunata Sarg. (a), C. laetifica Sarg. (a; n), C. macilenta 
Beadle (Ala.), C. modesta Sarg. (a), C. neobuskii Sarg. (n), C. 
Painteriana Sarg. (a; n), C. rw6e//a Beadle (a), C. Sargentii 
Beadle (a), C. 5c<26r<z Sarg. (a; n), C. Schweinitziana Sarg. 
(Penn.), C. 5^rflw/«€a Beadle (Penn.), C. ^ec^a Beadle (Ala.), C. 
villicarpa Sarg. 

Crataegus ambrosia Sarg. (n), C. aquilonaris Sarg. (n), C. arrfz^ 
Sarg. (n), C. baccata Sarg. (n), C. Balkwillii Sarg. (n), C. ^ec^i- 
cwfl Sarg. (n), C bristolensis Sarg. (n), C. calpodendron (Ehrh.) 
Medic. (Penn.), C. chadfordiana Sarg. (n), C. Chapmanii (Beadle) 
Ashe (a; n; N. Car.), C. conspecta Sarg. (n), C. conspicua Sarg. 
(n; Vt.), C. corporea Sarg. (n), C. delectabilis Sarg. (Ont.), C. 
Deweyana Sarg. (a; n), C. rfiufrfa Sarg. (n), C. dumicola Sarg. 
(n), C. Emersoniana Sarg. (a; n), C. jerentaria Sarg. (a; n), C. 
/er^fl Sarg. (n), C. jertilis Sarg. (a; n), C. finitima Sarg. (a; n), 
C. jlagrans Sarg. (n), C. fiammea Sarg. (n), C. jrutescens Sarg. 
(n), C. /M/^e«5 Sarg. (a; n), C. /w/gfrfa Sarg. (n), C. Gam/^w Sarg. 



JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

(a; n), C. gemmma Sarg. (n), C. glabrata Sarg. (n), C. globosa 
Sarg. (a; n), C. Halliana Sarg. (n), C. hystricina Ashe (n), C. illi- 
noiensis Ashe (n), C. integriloba Sarg. (n), C. Laneyi Sarg. (a; n), 
C. laurentiana Sarg. (n), C macracantha Lodd. (a; n; Conn., 
N. Y., S. Dak., W. Va., Wis.), C. macracantha var. succulenta 
Rehd. {= C. succulenta Schrad.) (n; Penn., Wis.), C. mem- 
branacea Sarg. (n; Vt.), C. michiganensis Ashe (n), C. micro- 
sperma Sarg. (n), C. missouriensis Ashe (a; n), C. neofiuvialis 
Ashe (n; Penn.), C. wwrfa Sarg. (n), C. ogdensburgensis Sarg. (n), 
C. Peckietta Sarg. (N. Y.), C. pellucidula Sarg. (n), C. peramoena 
Sarg. (n), C. pertomentosa Ashe (Iowa, Kansas), C pisijera Sarg. 
(n; Vt.), C. praeclara Sarg. (a), C. propixa Sarg. (a), C. /^rwrn- 
/o/w (Marsh.) Pers. (a; n), C. /)«rfe«j Sarg. (a; n), C. rhombijolia 
Sarg. (n; Conn., N. Y., Mass., Vt.), C. Robinsonii Sarg. (n), C. 
rupkola Sarg. (a), C. saeva Sarg. (n), C. Seami Sarg. (n), C. 
simulata Sarg. (n), C. spatiosa Sarg. (n), C. spinulosa Sarg. (a; n), 
C. structilis Ashe (n), C. tomentosa L. (a, Thaxter [1880] ; n; 111., 
Iowa, Ky., Maine, Miss., Mo., Ohio, Ont., Que., Wis.), C. truculenta 
Sarg. (n), C. 7;a^a Sarg. (a; n), C. iiege^a Sarg. (a; n), C. venu- 
losa Sarg. (a; n), C. venustula Sarg. (n), C. Wilsonii Sarg. (n). 



Crataegus Handyae Sarg. (n). 
Microcarpae: 

Crataegus Phaenopyrum (L. f.) Medic. (= C. cordata Ait.) (Del., 

MOLLES: 

Crataegus anomala Sarg. (n; Conn., N. Y.), C. arnoldiana Sarg. 
(a; n), C. Berlandieri Sarg. (n), C. canadensis Sarg. (n), C. c/fflm- 
plainensis Sarg. (a; n; N. Y.), C. contorti folia Sarg. (n), C. corw^ca 
Sarg. (111-.), C. rfi^wfl Sarg. (n), C. dw/^ej^a Ashe (a; Mo.), C. dume- 
tosa Sarg. (a; Mo.), C. Ellwangeriana Sarg. (a; n), C. exclusa 
Sarg. (n), C. Fulleriana Sarg. (n), C. Greggiana Eggl. (a), C. 
mc?«/a Sarg. (a), C. invisa Sarg. (n), C. /am^era Sarg. (n), C. 
lanuginosa Sarg. (a; n), C lasiantha Sarg. (a; n; Mo.), C. /awfa 
Sarg. (n), C. limaria Sarg. (a; n), C. macrophylla Sarg. (n), C. 
meridionalis Sarg. (n), C. mollipes Sarg. (n), C. wo//w (Torr. & 
Gr.) Scheele (a. Bliss [1931] ; n; 111., Ind., Iowa, Kan., Ky., Mass., 
Mo., Nebr., Ohio), C. noelensis Sarg. (n), C. nutans Sarg. (n), 
C. pennsylvanica Ashe (n), C. peregrina Sarg. (a; n), C. Robesoni- 
ana Sarg. (n), C. jerc Sarg. (a; n), C. submollis Sarg. (a; n; Vt.), 



1935] MacLACHLAN, HOSTS OF GYMNOSPORANGIUM GLOBOSUM 133 

C. Tatnalliana Sarg. (n), C. Tracyi Ashe (a), C. transmississippi- 
ensis Sarg. (n), C. Treleasei Sarg. (Mo.), C. umbrosa Sarg. (n), 
C. urbica Sarg. (n). 

X Crataegus hiemalis Lge. (n), C. nigra Kit. (n). 

Oxyacanthae: 

Crataegus monogyna Jacq. (a; n; Mass.), C. nwnogyna var. in- 
ermis Rehd. (a), C. monogyna var. laciniata (Stev.) Kegel (a; n), 
C. monogyna var. pteridijolia Rehd. (a; n), C. Oxyacantha L. 
a, Farlow [1885]; n; Maine, Mass., Ont.), C. Oxyacantha var. 
Gireoudii Bean (a), C. Oxyacantha var. leucocarpa Loudon (a), 
C. Oxyacantha var. rw^ra Hort. (a), X C. sorbijoUa Lge. (a; n). 

Pinnatifidae: 

Crataegus pinnatifida Bge. (n), C. pinnatifida var. wcjor N. E. 
Br. (n). 

Pruinosae: 

Crataegus alacris Sarg. (a), C. amoena Sarg. (a), C. arcana Beadle 
(n), C. aridula Sarg. (a), C. a5/'era Sarg. (a; n), C. ater Ashe (a), 
C. ^)eflfa Sarg. (n), C. 6e//M/a Sarg. (n), C. bracteata Sarg. (a), 
C. caerulescens Sarg. (n), C. ce^^'ca Sarg. (a), C. Clintoniana 
Sarg. (n), C. cognata Sarg. (n), C. comata Sarg. (n), C. comparata 
Sarg. (n), C. confragosa Sarg. (n), C. conjuncta Sarg. (a; n; Conn., 
Mass.), C. delawarensis Sarg. (a), C. deltoides Ashe (a; n), C. 
disjuncta Sarg. (a; Mo.), C. divisifolia Sarg. (n), C. exornata 
Sarg. (n), C. Ferrissii Ashe (n), C. /e^fii^a Sarg. (Conn., Vt.), 
C. jormosa Sarg. (a; n), C. /M5ca Sarg. (a), C. georgiana Sarg. 
(a; n), C. glareosa Ashe (n), C. horridula Sarg. (a; n), C. mcwa 
Sarg. (a; n), C. inusitula Sarg. (a; n), C. iracunda Beadle (a; n), 
C yeJw/'M Sarg. (Penn.), C. Kellernmnii Sarg. (a), C. latijrons 
Sarg. (n), C. latisepala Ashe (a; n), C. leiophylla Sarg. (a; n; 
N. Y.), C. /ei'w Sarg. (a; n), C. littoralis Sarg. (a), C. locuples 
Sarg. (a; n), C. «wwero5a Sarg. (a; n), C. oWifa Sarg. (a; n), 
C. Pequotorum Sarg. (a; n; Conn.), C. perampla Sarg. (a; n), 
C. perjucunda Sarg. (a), C. philadelphka Sarg. (a; n), C. /'xVo^a 
Sarg. (n), C. platycarpa Sarg. (a), C. Former/ Britt. (n), C. /?ro- 
cera Sarg. (a; n), C. pruinosa (Wendl.) K. Koch (a; n; Conn., 
Mo., N. Y., Ohio, S. Car., Penn.), C. pruinosa var. latisepala (Ashe) 
Eggl. (Mass., Mich.), C. pulchra Sarg. (a; n), C. quinebaugensis 
Sarg. (Conn.), C. radiata Sarg. (a; n), C. relicta Sarg. (n), C. 
remote Sarg. (n), C. rubicundula Sarg. (a; n), C. 5«7m/c Sarg. 



134 JOURNAL OF THE ARNOLD ARBORETUM 

in), C. Sicca S^rg. (n) 
(n), C. uplandia Sarg. 

Crataegus decorata Sarg. (n; Mo.). 

Pulcherrimae: 

Crataegus ancisa Beadle (Ala.), C. illustris Beadle (a). 

Punctatae: 

Crataegus amnicola Beadle (a; n), C. angustata Sarg. (a), C. bar- 
bara Sarg. (a; n), C. Broumietta Sarg. (n), C. calvescens Sarg. 
(n), C. ce/^a Sarg. (n), C. coZ/mc Chapm. (Ga., Va.), C. com- 
/•ac^fl Sarg. (n), C. Dewingii Sarg. (n), C. Eatoniana Sarg. (n), 
C. Eastmaniana Sarg. (a; n), C. florifera Sarg. (a; n), C. g/a6n- 
/o^ia Sarg. (a; n), C. incerta Sarg. (n), C. Lettermanii Sarg. (a), 
C. macropoda Sarg. (a; n), C. notabilis Sarg. (n), C. pausiaca 
Ashe (a; n), C. porrecta Ashe (n), C. praestans Sarg. (a; n), C. 
pratensis Sarg. (a; n), C. punctata Jacq. (a; n; 111., Ind., Iowa, 
Maine, Mass., Mich., Mo., N. Y., N. Car., Ohio, Ont. Penn., Vt., 
W. Va.), C. punctata var. aurea Ait. (a; n), C. punctata var. 
canescens Britt. (n), C. punctata var. malijormis ? (n), C. />««<:- 
^a^c mutabilis Gruber (a; n), C. ^ec^a Sarg. (a; n), C. sordida 
Sarg. (a), C. suborbkulata Sarg. (a; n), C. succincta Sarg. (a), 
C. JM<:/fl?c Sarg. (Mo.), C. swanensis Sarg. (a; n), C. ^cwax Ashe 
(a; n), C. umbratilis Sarg. (a; n), C. verruculosa Sarg. (n), C. 
z'/dwfl Sarg. (a). 

Rotundipoliae: 

Crataegus Bicknellii Eggl. (n), C. Blanthardii Sarg. (n), C. ^rc?- 
«er(/« Sarg. (a; n; Vt.), C. Brunetiana Sarg. (a), C. calkiglabra 
Schuette (a), C. chrysocarpa Ashe (N. Y.), C. coccinata Sarg. 
(n), C. crassijolia Sarg. (n), C. cupulifera Sarg. (n), C. diver gens 
(Peck) Sarg. (a), C. Dodgei Ashe (n), C. Evansiana Sarg. (a; n), 
C. Faxonii Sarg. (n), C. illuminata Sarg. (n), C. inaudita Sarg. (a), 
C. m5o/e«5 Sarg. (n), C. /ac-t/7 Sarg. (n), C. /owe^ag Sarg. (a; n), 
C. A'ee/>«7 Sarg. (n), C. Kennedyi Sarg. (n), C. kingstonensis Sarg. 
(n), C. lemingtonensis Sarg. (n), C. maligna Sarg. (n), C. mansfield- 
ensis Sarg. (n), C. Margaretta Ashe (n; Iowa, Mo.), C. Margaretta 
f. xanthocarpa Sarg. (n), C. Maribella Sarg. (n), C. Oakesiana Eggl. 
(a), C. praecoqua Sarg. (= C. praecox Sarg.) (n; N. Y.), C. Proc- 
/omwa Sarg. (n), C. propria Sarg. (n), C. rotundata Sarg. (n), 
C. rotundijolia Moench (= C. coccinea L. p. p.) (a, Thaxter 
[1889]; n; Iowa, Mo., N. Y., Ont., Vt.), C. rotundijolia var. 



1935] MacLACHLAN, HOSTS OF GYMNOSPORANGIUM GLOBOSUM 135 

aboriginum Sarg. (n), C. rotundijolia var. pubera Sarg. (n), C. 
rotundijolia f. rubescens Sarg. (n), C. i^amm Sarg. (n), C. M^e/>- 
5^m Sarg. (n), C. Williamsii Eggl. (n). 

Sanguineae: 

Crataegus altaka Lange (n), C. dsungarka Zab. (n), X C. Law- 
bertiana Lge. (n), C. Maximowkzii Schneid. (n), C. sanguinea 
Pall. (Ont.). 

Silvicolae: 

Crataegus aemula Beadle (n), C. a/Zec/a Sarg. (n), C. Barryana 
Sarg. (n), C. blairensis Sarg. (n), C. congesti flora Sarg. (a; n), 
C. crwcfa Sarg. (n), C. delectata Sarg. (n), C. rfi^M5a Sarg. (= C. 
silvkolavar.Beckwithae [Sarg.] Eggl.) (n; CoT\n.,Vt.),C. dissona 
Sarg. (n; Mass., N. H., N. Y.), C. eifero Sarg. (n), C. /j^i/>ej Ashe 
(n), C. joliata Sarg. (n), C. Fre^2« Sarg. (n), C. graz;w Ashe (n), 
C. iterata Sarg. (n), C. /ae^aw5 Sarg. (n), C. Livingstoniana Sarg. 
(n), C. luxuriosa Sarg. (n), C. wcce^-a Sarg. (n), C. Maineana 
Sarg. (n), C. medioxima Sarg. (n), C. o/JM/e«5 Sarg. (n), C. 
promissa Sarg. (a; n), C. />ro«a Ashe (n), C. puta Sarg. (n), C. 
rcrfmc Sarg. (n), C. recordabilis Sarg. (n), C. Robbinsiana Sarg. 
(Vt.), C. rwr/co/fl Sarg. (n), C. stolonijera Sarg. (n), C. j^rigo^a 
Sarg. (n), C. tortuosa Sarg. (n), C. xanthophylla Sarg. (a; n). 

Tenuifoliae: 

Crataegus acuminata Sarg. (a; n), C. acutiloba Sarg. (a; n; N. Y., 
Vt.), C. alnorum Sarg. (n), C. apiomorpha Sarg. (n), C. ascendens 
Sarg. (n), C. asperata Sarg. (n), C. 6a5i/zca Beadle (a), C. 6e//G 
Sarg. (a; n), C. benigna Sarg. (a; n), C. blandita Sarg. (n), C. 
BootMana Sarg. (n), C. co/ora^a Sarg. (a; n; Ont.), C. con- 
jerta Sarg. (n), C. crMrfe/J5 Sarg. (n), C. cyanophylla Sarg. 
(a; n), C. Damei Sarg. (n), C. delucida Sarg. (n; Vt.), C. rfewma 
Sarg. (n; Mass., Vt.), C. dissimilis Sarg. (a; n; Conn., Mass., 
Vt.), C. Edsoni Sarg. (n; N. H., Vt.), C. Eganii Ashe (n), C. /irwc 
Sarg. (n), C. flabellata (Bosc.) K. Koch (a; n), C. /lorea Sarg. 
(n), C. Forbesae Sarg. (a; n; Conn.), C. /mcoja Sarg. (n), C. 
gewJc/« Sarg. (a; n; Vt.), C. glaucophylla Sarg. (a; n; Conn., 
N. Y.), C. gracilipes Sarg. (n), C. Grw^eri Ashe (n), C. Habereri 
Sarg. (n), C. Hadleyana Sarg. (n), C. heidelbergensis Sarg. (n), 
C. m5o/z7a Sarg. (n), C. leptopoda Sarg. (n), C. lucorum Sarg. 
(n), C. luminosa Sarg. (n), C. macros perma Ashe (n; N. Y., 
Penn.), C. marcida Ashe (n), C. wa/wrc Sarg. (n), C. media Sarg. 
(n), C. OTcnVa Sarg. (n), C. miniata Ashe (n), C. worf/ca Sarg. 
Sarg. (n), C. Napaea Sarg. (n), C. ne5CJa Sarg. 



136 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

(n), C. otiosa Ashe (n), C. Paddockeae Sarg. (n), C. Paineana 
Sarg. (n), C. pallidula Sarg. (n), C. parvi flora Sarg. (n), C. pas- 
torum Sarg. (a; n), C. paucispina Sarg. (a), C. pentandra Sarg. 
(a; n; Vt.), C. perlevis Ashe (n), C. populnea Ashe (n), C. /^Mw^Ya 
Sarg. (n), C. retrusa Ashe (n), C. roanensis Ashe (Ky., Vt.), C. 
rubicunda Sarg. (n), C. rubrocarnea Sarg. (n), C. rw^/^ej Ashe 
(n), C. sarniensis Sarg. (n), C. ^a^wra^a Sarg. (n), C. 5ere»a Sarg. 
(n), C. 5er^?Y« Sarg. (n), C. wrferea Sarg. (n), C. Slavini Sarg. (n), 
C. Streeterae Sarg. (n), C. jmcw Sarg. (n), C. taetrka Sarg. (n), 
C. icrrfc Sarg. (n), C. tenella Ashe (n; Conn.), C. tenera Ashe (n), 
C. tenuiloba Sarg. (n), C. trachyphylla Sarg. (n), C. M^er Ashe 
(n), C. viridimontana Sarg. (n), C. vittata Ashe (a). 

Triflorae: 

Crataegus austronwntana Beadle (a). 

Uniflorae: 

Crataegus armentalis Beadle (a), C. Brittonii Eggl. (a). 

ViRIDES: 

Crataegus abbreviata Sarg. (a; n), C. atrorubens Ashe (a; n), C. 

6/anrfa Sarg. (a), C. enucleata Sarg. (a; n), C. lanceolata Sarg. 

(a; n), C. larga Sarg. (a), C. lutensis Sarg. (a), C. wf/ewj Sarg. 

(a), C. mVfrfa (Engelm.) Sarg. (a; n), C. oiifl^a Sarg. (a; n), C. 

penita Beadle (a), C. poliophylla Sarg. (a), C. uvaldensis Sarg. 

(a), C. ■ye/w^ma Sarg. (a), C. z^mrf/5 L. (a; n; Okla.), C. ^^w/^a 

Beadle (a; n). 
CYDONIA: 

Cydonia oblonga Mill. (= C. vulgaris Pers.) (a; Thaxter 1 1889] ; 

Conn., Niagara Peninsula, N. J., Penn.). 
MALUS: 

Malus angustifolia Michx. (S. Car.), X M. astranica Dum.-Cours. 

{3i),M.baccataBorkh. {^), M. coronaria MiW. (a, Arthur [1907]), 

X M. Dawsoniana Rehd. (a), M. glabrata Rehd. (a), M. glau- 

cescens Rehd. (Ind.), M. ioensis var. plena Rehd. (a), X *M. 

magdeburgensis Schoch (a), M. pumila Mill. (= M. Malus |L.] 

Britt.) (Thaxter [1886] ; Conn., Maine, Mass., Mo., N. H., N. J., 

N. Y., Vt.), X M. Soulardi Britt. (a), X M. sublobata Rehd. (a). 
MESPILUS: 

Mespilus germanica L. (a). 
PYRUS: 

Pyru^ Balansae Decne. (a), P. bettdaejolia Bge. (a; n), P. Bret- 

schneideri Rehd. (a), P. communis L. (a; Conn., Ind., Iowa, Mass., 



1935] MacLACHLAN, HOSTS OF GYMNOSPORANGIUM GLOBOSUM 137 

N. Car., N. Y., Penn., R. I.), P. elaeagrijolia Pall, (a), *P. Kor- 
shinskyi Litv. (a), */>. Michauxii Bosc (a), *P. Lindleyi Rehd. (a), 
*P. nivalis Jacq. (a), P. phaeocarpa Rehd. (a), P. salicijolia Pall, 
(a), P. serotina Rehd. (a), *P. serrulata Rehd. (a), P. syriaca 
Boiss. (a), P. ussuriensis Maxim, (a). 

SORBARONIA: 

X *Sorbaronia alpina Schneid. f. superaria Zabel (a). 

SORBOPYRUS: 

X *Sorbopyrus auricularis Schneid. (a). 

SORBUS: 

Sorbus americana Marsh, (a; Thaxter [1887 and 1891]; Maine, 
Mass., N. Y., Penn., Vt.), S. americana var. jructu albo Hort. (a), 
*S. americana var. nana Hort. (a), X *5. arnoldiana Rehd. (a), 
*5. Aucuparia L. var. Backhousei Hort. (a), *S. dumosa Greene 
(a), *5. japonica var. calocarpa Rehd. (a), X S. thuringiaca 
Fritsch (a). 

HOSTS FOR THE HI STAGE 

JUNIPERUS: 

Juniperus lucayana Britt. (= /. barbadensis Auth., not L.) (Ala.), 
/. communis L. (Penn.), J. jragrans Hort. (Ont.), /. korizontalis 
Moench (= /. prostrata Pers.) (N. Dak.), /. scopulorum Sarg. 
(Colo., 111., Iowa, N. Dak.), /. virginiana L. (Ala., Conn., 111., Ind., 
Iowa, Kansas, Ky., La., Mass., Mich., Minn., Miss., Mo., N. H., 
N. Y., N. Car., N. Dak., Ohio, Okla., Ont., Penn., S. Car., Tex., 
Vt., W. Va., Wis.), /. virginiana var. Burkii Hort. (111.), /. virgini- 
ana var. Canaertii Senecl. (111.), /. virginiana var. elcgantissima 
Hochst. (III.), /. virginiana var. glauca Carr. (111.). 

LARIX: 

Larix sp. (Conn., Kan., Minn., Miss., N. Y., Okla., Tex., Va., 
W. Va.). 

V. SUMMARY 

1. At least ten genera, all within the Pomoideae, include hosts on 
which the aecial phase of Gymnosporangium globosum may occur. One 
genus only, Juniperus, is known with certainty to include hosts for the 
telial phase. 

2. Relative susceptibility to G. globosum within the respective 
host genera has been studied by the writer to determine: (1) immune 
species; (2) resistant species which suffer no material harm from this 
rust; (3) moderately susceptible species which may be infected but not 
to the extent of defoliation; and (4) very susceptible species whose 
foliage can be ruined by G. globosum. 



138 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

3. These investigations were carried out by means of artificial inocu- 
lations, substantiated by observations of natural infection where pres- 
ent, in the Arnold Arboretum of Harvard University. 

4. The results of these investigations on relative susceptibility, 
added to those of previous writers, may be summarized as follows: 

A. On host genera for the aecial phase of G. globosum. 

(a) On the genera on which serial inoculations were made. 
Crataegus. A marked variation in susceptibility was found within 

the genus, the degree of which is dependent primarily on the thickness 
and the rapidity of deposition of the foliar cuticle. Due to the large 
number of species and the unstable condition of taxonomy within the 
genus, the classification according to susceptibility to G. globosum was 
made by groups rather than by species. The observations on natural 
infection substantiated the results obtained by artificial inoculation. 
Suggestions have been made for the selection of resistant species and 
varieties within the respective groups. 

Pyrus. Of seventeen species inoculated, one proved to be very 
susceptible, two moderately susceptible, ten resistant, and three 
immune. Certain of the commercial varieties are classified from pre- 
vious reports according to their susceptibility to G. globosum. 

Sorbus. Infection was obtained on all the species and varieties of 
American origin inoculated. Of thirty-one species and varieties of 
Eurasian origin inoculated four are resistant, the remainder are immune. 

Malus. Of seven American species inoculated three proved to be 
susceptible, while infection was obtained on only one species and three 
hybrids of the twenty-seven Eurasian types considered. Infection was 
obtained also on two hybrids between Eurasian and American species. 
Certain of the commercial varieties are classified from previous reports 
according to their susceptibility to G. globosum. 

(b) On the genera otherwise inoculated. 

Amclanchier. Seventeen species and varieties were inoculated; all 
inoculations gave negative results. Nevertheless, the rust has been 
reported on two species, .4. canadensis and A. alnijoUa.^ It is not prob- 
able that any species in this genus would suffer severely from infection 
by G. globosum. 

Cydonia. Gymnosporangium globosum has been reported as occur- 
ring commonly on quince in New Jersey. Cydonia oblonga by culture 
proved to be moderately susceptible to G. globosum. 

Crataegomespilus, Mespilus, Sorbaronia and Sorbopyrus. The re- 



1935] MacLACHLAN, HOSTS OF GYMNOSPORANGIUM GLOBOSUM 139 

suits obtained by inoculations on representatives of these more or less 
susceptible genera have been tabulated on page 127. 

Comptonia, Myrica and Photinia. These genera were found by in- 

B. Host genera for the telial phase of G. globosum. 

Juniperus. No information to date would indicate that any species 
other than /. virginiana and /. scopulorum and their varieties would 
suffer to any extent from infection by G. globosum. 

5. In the genera Crataegus, Mains, Pyrus and Sorbus there is a defi- 
nite duration to the period of susceptibility reaching a maximum during 
or immediately after foliar expansion. 

6. In selecting ornamentals to plant in vicinities where Gymno- 
Sporangium rusts are present, it must be remembered that the relative 
susceptibility of any host to G. globosum is not necessarily correlated 
with its susceptibility to other Gymnosporangium rusts. 

7. No consideration has been given to the possibility of variation 
in virulence within different strains of G. globosum. Such may very 
well occur. 

8. A complete list of all the known hosts of G. globosum is recorded 
in this paper. 

VI. ACKNOWLEDGMENTS 

To Professor J. H. Faull for his generous assistance in making this 
study possible and for his guidance, supervision and other expressions 
of personal interest the writer acknowledges deep obligation. 

To the Arnold Arboretum for permission to use its facilities; to Pro- 
fessor A. Rehder and Mr. E. J. Palmer for their invaluable assistance 
in the taxonomic treatment of the host genera; to Dr. A. E. Navez for 
his careful analysis of data and for helpful advice; and to Dr. Ivan H. 
Crowell for his cooperation and his help in field work the writer also 
expresses gratitude. 

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plants in the United States in 1931. (Plant Dis. Rep. Suppl. 84:50.) 



1935] MacLACHLAN, HOSTS OF GYMNOSPORANGIUM GLOBOSUM 141 

Rehder, A. (1927). Manual of cultivated trees and shrubs. (MacMillan 

and Co., New York, pp. 363-376.) 
Sherbakoff, C. D. (1932). The more important diseases of apples in 

Tennessee. (Univ. of Tenn. Agric. Exper. .Sta. Bull. 145:34-37.) 
Stevens, F. L. and Hall, J. G. (1910). Diseases of economic plants. 

(MacMillan and Co., New York, pp. 106-107.) 
Stewart, F. C. (1910). Notes on New York plant diseases, I. (N. Y. 

Agric. Exper. Sta. Bull. 328:376.) 
Stone, R. E. (1908). Cedar apples and apple leaf rust. (Ala Agric 

Exper. Sta. Cir. 2:5.) 

(1909). Species of Gymnosporangium in southern Alabama. 

(Torreya, 9:116.) 

Thaxter, R. (1887). VII. On certain cultures of Gymnosporangium 
with notes on their Roesteliae. (Proc. Amer. Acad, of Arts and Sci. 
22:263.) 

(1889). Notes on cultures of Gymnosporangium made in 1887 

and 1888. (Bot. Gaz. 14:167-168.) 

(1891). The Connecticut species of Gymnosporangium (cedar 

apples). (Conn. Agric. Exper. Sta. Bull. 107:4.) 

Thomas, H. E. and Mills, W. D. (1930). Rust diseases of the apple. 
(Plant Dis. Rep. 14:214-215.) 



VIII. EXPLANATION OF PLATES 
Plate 125 
rations of the tendency of the mycelium to follow along the veins 
aegus leaves : 
A series of lesions obtained by inoculation on a waxy-type of 
leaf (Crataegus fecunda), giving the appearance of systemic 
infection along the veins. 

A single lesion at the spermogonial stage on Crataegus suavis. 
The rust mycelium concentrates along the vascular strands caus- 
ing the latter to show as bright yellow lines within the lesion. 

I extends along a lateral vein, forking at the junc- 



the lesion correspond- 



- 1. Illustrate 


the relative amount of leaf killing caused by vein in- 


fections. 


and by infections not primarily associated with the 


main veir 




?. 2. A single 


nfection on the mid vein resulting in the death of over 


one-half 


of the leaf. 


^ 3. A very s 


nail type of lesion, exhibiting no hypertrophy and pro- 


ducing a 


single aecial horn. 


r. 4. A single 


v^m infection (indicated by the black spot on the plate) 


killing th 


e leaf behind the lesion along the vein ; suggesting a 


toxic age 


It on the part of the rust. 


-. 5. A large 




appeared 


suggesting hypersensitivity on the part of the host. 



JOURNAL OF THE ARNOLD ARBORETUM 



, 2, 3 and 4, illustrate the relative degree of susceptibility of ( 
gus Pringlei, as indicated by serial inoculations on April 2, 
9, May 23 and June 28, 1934, respectively. 

. The type of chamber used in all the inoculations. (Explai 



Plate 128 
Serial inoculations on Crataegus Jonesac to illustrate the period of s 
ceptibility (explanations in text) : 

Fig. 1 . Inoculated May 7, at which time the two upper leaves were v( 

small, while the five basal leaves were well expanded. As in 

cated by the number of lesions the latter are the more susceptil 

Fig. 2. Inoculated June 8, at which time all leaves were fully expandi 

the two upper (youngest) leaves are now the more susceptible 



' OF Plant Pat 



JoLR. Arnold Ark. Vol. XVl. 






m^^ 



1935] HUNTER, STUDIES OF EUROPEAN SPECIES OF MILESIA 



Lillian M. Hunter 

Although eleven species of Milesia are known to occur in Europe 
(Faull, J. H. Taxonomy and Geographical Distribution of the Genus 
Milesia. Contr. Arnold Arb. Harvard Univ. II. 1932) up to the pres- 
ent the life histories of two only of them have been worked out, namely, 
M. Blechni (Syd.) Arth. (Klebahn, H. Kulturversuche mit Rost- 
pilzen. In Zeitsch. Pflanzenkr. 26: ISl-lll. 1916) and M. Kriegeri- 
ana (Magn.) Arth. from Dryopteris Filix mas (L.) Schott (Mayor, 
EuG. Notes Mycologiques VIII. In Bull. Soc. Neuchat. Sci. Nat. 
58:23-26. 1933). 

Recently it was my privilege to make certain investigations on life 
histories of Milesia rusts in England. Teliosporic material of several 
species was assembled and inoculation experiments were made on vari- 
01^ firs with the results that spermogonia and aecia of the following 
species of Milesia have been obtained for the first time— 

' (1) Milesia Scolopendrii (Fuckel) Arth. (from Scolopendrium vul- 
gare Smith) on Abies alba Mill., and A. concolor Lindl. and Gord. 

(2) Milesia Polypodii B. White (from Polypodium vulgare L.) on 
Abies alba and A. concolor. 

(3) Milesia vogesiaca (Syd.) Faull (from Polystichum angulare 
Presl) on Abies alba. 

(4) Milesia Kriegeriana (Magn.) Arth. (from Dryopteris spinulosa 
[O. F. Miiller] Kuntze) on Abies alba, A. concolor and A. grandis 
Lindl. 

Spermogonia and aecia were also obtained for Milesia Kriegeriana 
(from Dryopteris Filix mas) on Abies alba and on two new hosts, 
namely, A. concolor and A. grandis. 

Aeciospores thus obtained by cultures were used in inoculating vari- 
ous ferns, and uredospores were obtained for the following species — 

(1) Milesia Scolopendrii on Scolopendrium vulgare. 

(2) Milesia Polypodii on Polypodium vulgare. 

(3a) Milesia Kriegeriana (from Dryopteris spinulosa) on Dryop- 
teris Filix mas, D. spintdosa and D. spinulosa var. intermedia (Muhl.) 
Underw. 

(3b) Milesia Kriegeriana (from Dryopteris Filix mas) on Dryop- 
teris Filix mas and D. spinulosa var. dilatata (Hoffm.) Underw. 



JOURNAL 

OF THE 

ARNOLD ARBORETUM 



STUDIES IN THE BORAGINACEAE, XI 

Ivan M. Johnston 



CONTENTS 
Tournefortia and Mes 



2. Notes on Brand's Treatment of Cryptantha. 

3. New or otherwise Noteworthy Species. . . . 



The Species treated here have, in the past, all been referred to the 
genus Tournefortia. I am, however, suggesting that certain of them be 
segregated to form the redefined genus Messerschmidia. During the 
work on this paper I have been privileged to examine almost all the type- 
specimens concerned. This has permitted me to place definitely a large 
number of poorly understood old species that have troubled workers in 
the past. The work has been undertaken as part of a projected study 
of the Boraginaceae-Heliotropioideae. It is the first attempt to treat 
all the Old World species of Tournefortia since the presentation by 
DeCandolle in the ninth volume of the Prodromus in 1845. 

Tournefortia Linnaeus, Sp. PI. 140 (1753) and Gen. PI. ed. 5, 68 
(1754). 

The species of Tournefortia found in the Old World all belong to the 
following: 

Section EuTOURNEFORTiA Johnston, Contr. Gray Herb. 92 : 66 
( 1930) . — type-species, T. hirsutissima L. Tournefortia — Pittoniae 
Humboldt, Bonpland & Kunth, Nov. Gen. et Sp. 3: 80 (1818). — type- 



146 JOURNAL OF THE ARNOLD ARBORETUM hoL.xvi 

species, T. hirsutissima L. Tournejortia sect. Pittonia Don, Gen. Syst. 
4: 366 (1837). — type-species, T. hirsutissima L. Pittonia Plumier ex 
Adanson, Fam. PI. 2: 177 (1763). — type-species, T. hirsutissima L. 
Oskampia Rafinesque, Sylva Tellur. 123 (1838). — type-species, O. 
scandens Raf. & 0. hirsuta Raf. Tournejortia sect. Tetrandra DeCan- 
dolle, Prodr. 9:527 (1845). — type-species, T. tetrandra Blume. 
Tetrandra (DC.) Miquel, Fl. Nederl. Ind. 2:928 (1858). — type- 
species, Tournejortia tetrandra Blume. 

The species of Eutournejortia found in the Old World are remarkable 
for their parallelism of variation. Most of them have corollas with the 
tube either long or short, herbage with the pubescence present or absent, 
as well as leaf-blades that are broad or elongate. The combinations of 
these variations produce forms very diverse in gross appearance so that 
it is not at all surprising that botanists have been impressed by them 
and misled into giving specific names to many of them. A considera- 
tion of all the Old World Eutournejortiae and observation of the recur- 
rent pattern of variation among them, however, lead one to a proper 
estimate of the surprisingly diverse phases which they present. Like- 
wise, a consideration of the facts of distribution leads to a similar end. 
When the variations mentioned are given recognition it is found that 
the resulting numerous ill defined "species" grow together over most of 
a common area of dispersal. When the variations mentioned are dis- 
counted, species may be defined that have a credible geographic range — 
a range that is distinct from that of the closely related species and one 
quite similar and familiar among species of other genera within the 
region. I am accordingly of the opinion that the variations noted deserve 
at best no more than mere formal recognition. Since, however, I do not 
believe that obscure tropical plants should be burdened with numerous 
subspecific names until some evident use for them arises, T have refrained 
from any attempt at formally naming the reoccurring combinations of 
the paralleling intraspecific variations described. 



'ipened fruit breaking up i 
these prominently ribbe 

'ipened fruit breaking up into two carpels which are each com- 
posed of two seminiferous cells and an intervening empty 

Flowers 4-merous 2. T. i 

Flowers 5-merous. 

Continental plants from southern Asia (including the Anda- 



JOHNSTON, STUDIES IN BORAGINACEAE, XI 147 

Calyx-lobes 3-4 mm. long at anthesis, usually sub- 
ulate; leaves drying more or less golden-brown 

beneath ; Sikkim 3. T. Hookcri. 

Calyx-lobes 1-2 mm. long at anthesis, linear or lanceolate. 
Flowers with evident pedicels 1-2 mm. long; Madras 

(cf. no. 5) 4. T. Heyneana. 

, blade more or less 
oval; flowers and fruit usually shortly pedicel- 
late ; Southern Burma and the Andamans 5. T. ovata. 

Leaves short-acuminate and usually not abruptly so ; 
blades oblong to lanceolate; flowers and fruit 

Insular plants. 

Western Pacific Ocean. 

Leaves opposite, flowers sessile; Philippines 7. T. ln::onica. 

Leaves alternate; flowers short-pedicellate; Australia 

and Papua 8. 7. Muellcri. 

Western Indian Ocean. 

Leaves obtuse or rounded at base, 4-11 cm. long. . .9. T. piibenila. 
Leaves acute at base, 10-20 cm. long. 

Stems with minute short closely appressed brownish 
or golden hairs or quite glabrous; calyx very 
sparsely strigose, the lobes cuneate, more or less 

erect ; Reunion 10. T. acuminata. 

Stems with evident abundant loosely appressed hairs 
(usually more or less velvety) ; calyx usually 

Sepals ovate ; Reunion 11. T. arhoresccns. 

Sepals lanceolate ; Mauritius 12. T. Bojcri. 

1. Tournefortia sarmentosa Lamarck, Tab. Encyc. 1: 416 (1791); 
Poiret, Encyc. 5: 357 (1804). Tournefortia orientalis R. Brown, Prod. 
497 (1810); Banks & Solander, Bot. Cook's Voy. 2: 64, tab. 210 (1901). 
Tournefortia tetrandra var. hirsuta Blume, Bijdrag. Fl. Nederl. Ind. 845 
(1826). Tournefortia sarmentosa var. hirsuta Blume ex Miquel, Fl. 
Ind. Batav. 2: 927 ( 1858), lapsus. Tournefortia hirsuta Reinwardt ex 
Boerlage, Hand. Fl. Nederl. Ind. 2^: 487 (1899). Tournefortia Urvil- 
leana Chamisso, Linnaea 4: 465 (1829). Tournefortia frangulae folia 
Zippel ex Spanoghe, Linnaea 15: 334 ( 1841 ?), in synon. Tournefortia 
Horsjieldii Miquel, Fl. Ind. Batav. 2:927 (1858). Tournefortia 
acclinis F. v. Mueller, Frag. 4: 95 (1864). Tournefortia macrophylla 
K. Schumann & Lauterbach, Fl. Deutsch. Schutzgeb. Sudsee 520 (1901). 
Tournefortia sarmentosa var. magnifolia Domin, Bibl. Bot. 22 (Heft 
89^) : 1097 ( 1928). Tournefortia glabrifolia Domin, Bibl. Bot. 22(Heft 
89^): 1098 (1928). 



148 JOURNAL OF THE ARNOLD 

Java to New Guinea, southward into northern Queensland and north- 
ward through the Celebes, Moluccas and Philippines to Formosa. 

A variable plant but readily recognized, even in its most diverse forms, 
by its characteristic fruit. At maturity this breaks up into four equal 
single-seeded nutlets. All the other Old World Tournejortiae have fruits 
with two 2-seeded carpels. Tournejortia sarmentosa has been repeatedly 
confused with continental species and has been the victim of numerous 
unsuccessful attempts at segregation. In T. sarmentosa the corollas 
may be either long or short, the herbage either glabrous or pubescent 
and the leaf-blades either small or large. These characters in various 
combinations have produced a host of forms that are superficially very 
diverse in appearance. These forms, however, agree in fundamental 
fruiting structures. None of them shows any evident geographical cor- 
relation. Grouped together to constitute T. sarmentosa, as here 
accepted, they appear as mere phases in a species which has a range that 
is natural and is of a type quite familiar to any student of the Malaysian 
flora. 

The type of T. sarmentosa, in the Lamarck Herbarium at Paris, is 
labeled "colitur in horto regio insulae Franciae" and "de M. Sonnerat." 
It is a good specimen showing leaves and flowers but no fruit. The 
corolla has a tube ca. 2 mm. long and a limb ca. 2.5 mm. in diameter. The 
calyx is 1.5 mm. long and has broad hairy lobes. The inflorescence is 
velvety with a dense short but somewhat shaggy, tan-colored indument. 
The stems and under surface of the leaves have abundant gray hairs. 
The upper surface of the leaves are- green and only sparsely strigose. 
The petiole is ca. 1 cm. long. The blade is rounded at the base, acute 
at the apex, and is 7.5-10 cm. long and 2.8-4 cm. broad. The plant is 
evidently the small-flowered hairy form of the common Tournejortia of 
the East Indian islands. It is certainly not a native of the Mascarenes! 
Gagnepain, Not. Syst. 3: 32-33 (1914), has discussed this species. His 
notes, except those referring to collections by Spire, Thorel, and Watt, 
all refer to the species as I have taken it. The excluded collections 
are from the Asiatic continent. The species is restricted to the islands 
and is not to be expected from the mainland. 

The type of T. orient alls, at the British Museum, is labeled as col- 
lected in 1770 by Banks and Solander at Endeavor Bay in northern 
Queensland. It is a glabrous plant with ovate to oblong leaves, 7-9 cm. 
long and 3.5-6.5 cm. broad. The corollas are large with a tube ca. 8 9 
mm. long and a limb 3-4 mm. broad. 

The type of T. Urvilleana was collected by Chamisso in Luzon. It 
has corollas 8 mm. long and a limb 3 mm. broad. The leaves are slightly 
less pubescent but otherwise are as in the type of T. sarmentosa. 



1935] JOHNSTON, STUDIES IN BORAGINACEAE, XI 149 

Blume's T. tetrandra var. hirsuta is given as from the Moluccas and 
described as follows: "ramis foliis pedunculisque hirsutissimis." At 
Leiden there is a specimen labeled: "Variet.; Tournefortia hirsuta; 
Manado; T. tetrandra Bl. Variet." The first and the last items are in 
Blume's script. The specimen is a form with elongate corollas and with 
more or less hairy leaves suggesting those of T. Horsfieldii. Menado is 
near the northern tip of the Celebes. Another specimen at Leiden has 
the following label: "1531 Tournefortia hirsuta R.; Habitat in insula 
Celebes ad viam inter Kema et Menado; Oct. 1821." This is associated 
with a printed label reading "Herbarium Reinwardtianum; in Acad. 
Lugduno-Batavia." This second specimen is similar to that first men- 
tioned and both are probably collections made by Reinwardt. They 
are, I believe, the types of T. tetrandra var. hirsuta and T. hirsuta. 

The name T. jrangulaefolia Zippel has appeared only in synonymy. 
At Leiden this name appears on two sheets having a printed label 
bearing: "Herb. Lugd. Batav.; Timor" and one in script reading: "1/6 
Tournefortia frangulaef olia ; Zp." 

Miquel based his T. Horsfieldii upon material cited: "Java, in Pat- 
jitan, Kelak (Horsf.)." I have examined specimens from Horsfield's 
personal herbarium at the British Museum and those from the set he 
made for the East India Company (now kept as a unit) at Kew. He 
made two collections referable to T. sarmentosa, 1: Pajittan (Kalak) 
Horsfield (borage 6) no. 275; and 2: Blambangan, Horsfield (borage 
7) no. 309. The former is evidently the type collection of T. Horsfieldii. 
It is a plant with very large leaves that are grayish velvety beneath. The 
blade becomes 10-14 cm. long and 7-9 cm. broad. The corolla-tube is 
7-8 mm. long and the limb is 3-4 mm. broad. 

Tournefortia acclinis is based upon material from Queensland col- 
lected by Bowman at Broad Sound and Amity Creek, and by Dallachy 
at Edgecombe Bay. A study of the original description and of a dupli- 
cate of Dallachy's material at Kew shows this species to have moderately 
sized leaves (5-10 cm. long and 3.5-6 cm. broad), a coarse appressed 
pubescence, a corolla with a tube 3-5 mm. long, and a corolla-limb 3-4 
mm. broad. It is very similar to T. Horsfieldii, except in leaf-size. 

The type of T. macrophylla was collected by Lauterbach (no. 2003) 
at Erima in eastern New Guinea. It is in fruit. The leaves are similar 
in size and shape to those of typical T. Horsfieldii. In fact the plant 
differs from the type of that species only in the practical absence of 
pubescence. The leaves have only a few weak scattered inconspicuous 

Domin's T. sarmentosa var. magnijolia from northern Queensland 



150 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

{Dietrich 724), to judge from description, seems to be merely a form of 
T. sarmentosa with very large (12-15 cm. long, 6-6.5 cm. broad) hairy 
leaves, and small corollas (corollae tubo breviore). His T. glabri folia is 
another large-leaved (10-13 cm. long and 5-5.5 cm. broad) plant. The 
leaves are glabrous. The corolla-tube is ca. 3-4 mm. long and the limb is 
ca. 2 mm. broad. The plant comes from Harvey's Creek in northeastern 
Queensland. It appears to differ from the type of T. orientalis only in 
its larger leaves and smaller corollas. 

2. Tournefortia tetrandra Blume, Bijdrag. Fl. Nederl. Ind. 845 
(1826). Tournefortia tetragona Blume ex Steudal, Nomencl. ed. 2, 
2:694 (1841). (?)Heliotropium scandens Norona, Verh. Bat. 

Genootsch 5:78 (1827); Hasskarl, Cat. Hort. Bogor. 137 (1844), 
nomen. Tournefortia tetrandra var. glabra Hasskarl, Flora 25- : Beibl., 
p. 27 (1842); Hasskarl, Cat. Hort. Bogor. 137 (1844); Hasskarl, PI. 
Javan. Rariores 492 (1848). Tournefortia glabra (Hassk.) Zollinger 
& Moritzi ex Zollinger, Natuur- en Geneeskundig Archief v. Nederl. Ind. 
2:5 (1845). Tetrandra glabra (Hassk.) Miquel, Fl. Nederl. Ind. 
2: 929 (1858). Tournefortia tetrandra var. longifiora Hasskarl, Cat. 
Hort. Bogor. 137 (1844), nomen; Hasskarl, PI. Javan. Rariores 492 
(1848). Tournefortia Wallichii DeCandolle, Prodr. 9: 527 (1845); 
Ridley, Fl. Malay Penin. 2: 441, fig. 115 (1923). Tetrandra Wallichii 
(DC.) Miquel, Fl. Nederl. Ind. 2: 928 (1858). Tetrandra Zollingeri 
Miquel, Fl. Nederl. Ind. 2: 928 (1858). 

Nicobar Islands, Malay Peninsula, Sumatra, Java, Borneo and 
Celebes. 

This is apparently the most common and best known of the Javan 
species of Eutoumefortia. The Javan plant has received the following 
basic names, Tournefortia tetrandra Blume, Tournefortia tetrandra var. 
glabra Hassk., Tournefortia tetrandra var. longifiora Hassk., and 
Tetrandra Zollingeri Miquel. The differences between these named 
forms are minor and variable ones of corolla-size and of distribution of 
pubescence on the foliage. This variable plant of Java I am quite 
unable to distinguish from Tournefortia Wallichii DC, a species based 
upon material from Singapore and Penang. I have accordingly accepted 
Tournefortia tetrandra as ranging from the Nicobar Islands eastward 
to Java and the Celebes. The leaves of this species are ovate-acuminate 
or lance-ovate and are glabrous or sparsely strigose. The fruit is usually 
subglobose and 4-6 mm. in diameter. The only notable departure from 
this is found among material from northern Borneo where the fruit, of 
several different collections, is narrowly ovoid, 7 mm. long and 4-5 mm. 



1935] JOHNSTON, STUDIES IN BORAGINACEAE, XI 151 

thick. This form may deserve some nomenclatorial recognition. There 
are, however, variations of Tournejortia tetrandra which I believe do 
merit recognition at this time. The characters which set these off from 
typical T. tetrandra may be organized as follows: 
Leaves l>4-25^ times as long as broad, ovate acuminate or 

lance-ovate Tournefortia tetrandra 

Leaves 2i^-354 times as long as broad, more or less lanceolate. 

Leaves dull, east Java var. angustifolia. 

Leaves somewhat glossy, Ceylon var. Waikerae. 

2A. Tournefortia tetrandra Blume var. angustifolia Moritzi, Syst. 
Verzeich. 52 (1845-46). 

Known only from the type-collection in eastern Java. 

This variety is a peculiar plant with very dull thickish leaves that have 
only 3-4 pairs of primary veins evident. The secondary nervation is 
not discernible. I know it only from the type-collection by Zollinger 
(no. 939), made Dec. 17, 1842, "auf den Kalkfelsen von Kuripan." 

2B. Tournefortia tetrandra Blume var. Waikerae (Clarke), comb, 
nov. Tournejortia Waikerae Clarke in Hooker, Fl. Brit. India 4: 147 
(1883); Trimen, Fl. Ceylon 3: 198 (1895). 

Known only from Ceylon. 

This plant is simply a narrow-leaved form of the species that is con- 
fined to Ceylon. The blades are lanceolate but are quite similar to those 
of the species in texture, nervation, etc. The fruit and flowers are simi- 
lar to the common Malaysian plant. 

3. Tournefortia Hookeri Clarke in Hooker, Fl. Brit. India 4: 147 
(1883). Tournejortia Hookeri var. subtropica Clarke in Hooker, Fl. 
Brit. India 4: 147 (1883). 

Known only from the base and lower valleys of the Sikkim Himalayas. 

Specimens Examined : Rangit, May 15, 1876, Clurkc 27953 (K) ; Great 
Rangit, April 1850, Hooker & Thompson (K, type of var. subtropica) ; 
Mangpu, 900 m. aU., May 1905, Meebold 4243 (BD) ; Rangbi, 1500 m., 
May 31, 1870, Clarke 11790 (K, BM) ; Chunbati, 600 m., June 12, 1870, 
Clarke 12024 (K, BM) ; Chunbati, 600 m., April 1876. Gamble 57P (K) ; 
below Punkabaree, Hooker (BD) ; Pancheni, 1875, Gamble 3370 (K) ; 
Siliguri, Jan. 1873, Gamble 3369 (K) ; Dalgaon, mixed forest, April 9, 
1893, Haines 358 (K) ; Sikkim, 1862, Andersson 270 (BD) ; Sikkim, 
March 1871, Clarke 16774 (K) ; Sikkim Terai, Clarke (K) ; Sikkim, 
Griffith 5928 (K). 

Characterized by its slender well developed calyx-lobes and by the 
golden or golden-brown under surfaces of the leaves. These latter are 
nearly glabrous or have only scattered hairs along the dark-colored 



152 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

nerves and veins. The corolla is usually 3-5 mm. long with the tube 
forming half (or even more) of this total length. In the var. subtropica, 
which is merely the large-flowered form of the species, the corollas 
become ca. 8 mm. long and the calyx-lobes only about a third as long as 
the tube. The species is a local one and probably worthy of recognition. 
It is most closely related to the form of T. niontana described as T. 

4. Tournefortia Heyneana Wallich, Num. List no. 910' (1828 29), 
nomen; Don, Gen. Syst. 4:369 (1837); Clarke in Hooker, Fl. Brit. 
India 4: 145 (1883); Gamble, Fl. Madras 893 (1923). Tournefortia 
reticosa Wight, Icones 4-: 16, tab. 1386 (1848); Wight, Spicileg. Neil- 
gherrense2: 83, tab. 189 (1851); Gamble, Fl. Madras 893 (1923). 

Hills of southern peninsular India, about lat. 11°-13° N. and long. 
lt°-ll° E. 

Specimens Examined: Nilgiri Hills, April 1852, herb. Wight 2057 
(K) ; Devala, Nilgiris, 900 m. alt., Nov. 1884, Gamble 15588 (K) ; S. E. 
Wynaad, Nilgiris, 900 m. alt., Nov. 1884, Gamble 15497 (K) ; Wynaad, 
Beddovte 5437 (BM) ; Nadooputtah, June 1846, herb. Wight (K) ; Ana- 
malais, Bcddomc 5438 (BM) ; Carcoor-ghat, Nilgiris, Aug. 1887, flowers 
varying from >^->4 inch according to age, Lawson (Oxford) ; Coorg, 
White (Oxford); Peermade Reav (? spelling), 1350 m. alt, Dec. 1910, 
Meebold 12920 (BD) ; without data, herb. Wight, probable basis of 
Wight's plate and the type of T. reticosa (K) ; without data, c.v herb 
Hcyne, Wallich 910' (herb. Wallich at Kew). 

A study of Wallich's herbarium, now at Kew, shows his number 910 
to consist of two different species from opposite ends of India. The label 
reads: "910 Tournef. Heyneana, Wall.— 1. Herb. Heyn. — 2. Pundua 
F. de S." The Heyne plant represents the species from the Deccan with 
pedicellate flowers, which is the one treated here. The plant from 
Pundua, collected by de Sylva, is accompanied by a large special label 
indicating that is was found in the "Pundouh Hills" in Jan. 1824. 

Clarke describes the flowers of T. Heyneana as 1/8-1/6 inches (3-4 
mm.) long. These measurements are evidently from the duplicate of 
the Wallich collections now in the general herbarium at Kew. The 
Heyne material in the Wallich Herbarium at Kew has corollas 9-10 mm. 
long. The specimens, except for flower-size, are otherwise very similar 
and I believe they represent minor forms of the species. Significant in 
this connection is the note made by Lawson on one of his specimens 
cited above. He states that the corolla varies from 3-12 mm. according 
to age! Though Don makes no mention of the corolla-size in his descrip- 
tion, the first given to the species, we may suppose that it was the large- 
flowered phase since the Wallich Herbarium, then in charge of the 



1935] JOHNSTON, STUDIES IN BORAGINACEAE, XI 153 

Linnean Society, was no doubt consulted by him. In any case, since 
the corolla-size is variable even within the type-collection, the chief 
character whereby Clarke distinguished T. reticosa now disappears. The 
two species, T. Heyneana and T. reticosa, are, I believe, trivial forms 
of one species and quite synonymous. 

DeCandolle, Prodr. 9: 516 (1845), received only the second part of 
Wallich no. 910, and described this as T. Heyneana. His description in 
the Prodromus, consequently, is based upon de Sylva's specimens from 
Pundua. Clarke pointed out this mistake, gave a new name (T. Can- 
dollii) to the de Sylva collection described by DeCandolle, and properly 
restricted the name, T. Heyneana, to the peninsular species collected by 

5. Tournef ortia oval^ Wallich, Num. List no. 908 (1828); Don, 
Gen. Syst. 4: 369 (1837); DeCandolle. Prodr. 9: 516 (1845); Clarke 
in Hooker, Fl. Brit. India 4: 147 (1883). 

Southern Burma and the Andaman Islands. 

Specimens Examined: Rangoon, Aug. 1826, Wallich (no. 15) 908 
(herb. Wallich, type); Rangoon, McClelland (K, three collections); 
Andamans, April 1891, Prain (Cambridge) ; Middle Andaman. Homfray 
Straits, climber, 1915, Parkinson 297 (K) ; Aberden, South Andaman, 
Kurz (K, parasitized; Delessert, normal); Chauldare, South Andaman, 

Characterized by its elliptical abruptly acuminate leaves, its sub- 
pedicellate flowers and its southern occurrence. The corolla becomes 
8 mm. long. The calyx is only 1.5 mm. long at anthesis. The leaves are 
mostly rather firm in texture and are usually brown and glabrous 
beneath. One of McClelland's collections is consequently quite atypical 
in having the leaves not only thin in texture but golden-brown beneath 
as well. Another one of his collections is quite hairy on the lower leaf- 
surface. The pedicels in T. ovata are usually at most only 1 mm. long, 
though in Parkinson's material cited the pedicels become fully 2 mm. 
long and are quite evident. 

6. Tournef ortia montana Loureiro, Fl. Cochinch. 1: 122 (1790). 
Messerschmidia montana (Lour.) Roemer & Schultes, Syst. 4: 544 
(1819). Lithospermum viridifiorum Roxburgh, Hort. Bengal. 13 
(1814), nomen; Lehmann, Asperif. 1: 30 (1818), insynon.; Roxburgh, 
Fl. Indica 2: 4 (1824), description; Roxburgh, Icones ined. Kew. tab. 
2120. Heliotropium viridifiorum (Roxb.) Lehmann, Asperif. 1:30 
(1818). Tournejortia viridiflora (Roxb.) Wallich, Num. List no. 907 
(1828); Clarke in Hooker, Fl. Brit. India 4: 146 (1883). Tournejortia 
Sampsoni Hance, Jour. Bot. 6:330 (1868). Tournejortia Wightii 
Clarke in Hooker, Fl. Brit. India 4: 146 (1883). Tournejortia Rox- 



154 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

burghii Clarke in Hooker, Fl. Brit. India 4: 146 (1883). Tournejortia 
viridiflora var. Grifflthii Clarke in Hooker, Fl. Brit. India 4: 146 ( 1883). 
Tournejortia Candollii Clarke in Hooker, Fl. Brit. India 4: 146 (1883). 
Tournejortia khasiana Clarke in Hooker, Fl. Brit. India 4: 147 (1883). 
Tournejortia Boniana Gagnepain, Not. Syst. 3:33 (1914) and in 
Lecomte, Fl. Gen Indo-Chine 4:217 (1914). Tournejortia Gaudi- 
chaudii Gagnepain, Not. Syst. 3: 34 (1914) and in Lecomte, Fl. Gen. 
Indo-Chine, 4:217 (1914). Tournejortia Heyneana sensu DeCan- 
dolle, Prodr. 9: 516 (1845). 

In the hills, up to 1500 m. alt., in Assam, Upper Burma, northern 
Siam (Payap and Maharat), middle and northern Indo-China (Anam, 
Laos and Tonkin) and southern-most China (Yunnan, Kwangsi and 
Kwangtung). 

This species presents a number of diverse phases resulting from com- 
binations of variations in leaf-size, abundance and distribution of 
pubescence, and size of corolla-tube. These phases have been treated 
as "species" but their variability, their erratic distribution, and their 
occurrence together in various localities lead me to believe they are 
merely further manifestations of the surprising intraspecific variability 
of these structures among the Old World Tournejortiae. After dis- 
counting these variations as mere phases, I am struck with the natural- 
ness of the distribution of the resulting aggregate species. The distri- 
bution is of the pattern found in numerous species of other genera and 
families inhabiting this part of Asia. 

The type of T. montana has not been examined. Its source is not 
given, but the probabilities are that it came from Anam. Dr. E. D. 
Merrill, who has devoted much time to the consideration of Loureiro's 
writings, informs me that he knows no reason for doubting Loureiro's 
generic attribution in the present case. After a study of the description 
I am perfectly content to accept Loureiro's name for this species. The 
leaves are given as ovate-lanceolate and glabrous. Unfortunately, how- 
ever, no information is given as to the shape or size of the corolla-tube. 

The second name applied to our species is Lithospermum viridijlorum. 
It first appears in 1814 as a name in a list of the Calcutta Garden and 
is given as collected by Roxburgh at Chittagong. It was no doubt this 
same garden material that was described in 1824 by Wallich in Rox- 
burgh's Flora and is now represented in Wallich's herbarium (no. 907). 
It is also the plant represented in Roxburgh's unpublished plates (no. 
2120) now preserved at Kew. The first description of the plant, as 
Heliotropium viridijlorum, is that by Lehmann in 1818. His material 
also seems to have come from the Calcutta Garden. Hence, there is 



1935] JOHNSTON, STUDIES IN BORAGINACEAE, XI 155 

every reason for taking the material grown at Calcutta as typical. This 
is a form characterized by distinctly lanceolate leaves that are velvety 
all over beneath and by small strigose corollas. The corolla-tube is 
2-i mm. long, usually densely strigose and commonly only twice the 
length of the calyx or less. This form has not been collected about 
Chittagong. As Clarke has indicated, 1. c. 146, the common form of 
T. montana about Chittagong, particularly in the region in which Rox- 
burgh is known to have collected, is the plant with long corolla-tubes 
described by Clarke as T. Roxburghii. As matters stand, therefore, we 
may either believe that Roxburgh did not collect his plant at Chitta- 
gong, or that having collected the common long-tubed Tournejortia there 
it subsequently became a short-tubed form under garden conditions. I 
have seen material of the type-form of T. viridiflora from Assam, Burma 
and Siam. 

The type of T. viridiflora var. Griffithii is a collection made in the 
Khasia Hills by Griffith. It differs from the type-form of T. viridiflora 
in having the leaves much less hairy or nearly glabrous beneath and 
corollas that are possibly a trifle larger. The type of T. Boniana col- 
lected by Bon (no. 1932) at 0-cach, on the mountain Ma-dong in Indo- 
China, is quite similar. I have seen this glabrescent small-flowered form 
from Assam, Burma and Indo-China. 

In publishing T. Wightii, Clarke gave its source as "Deccan Penin- 
sula, Wight." The type is Wight no. 2056 and is accompanied with one 
of the old printed labels indicating that it was part of the Wight 
materials handled at Kew in 1866-67. The label proper is headed 
"Peninsula Indiae Orientalis." We may accept that no. 2056 was part 
of the Wight Herbarium but as to the collector of the specimen and its 
original source doubt must remain. Since the plant agrees closely with 
plants from Burma I suspect that perhaps it came from that general 
region and may represent material received by Wight from Roxburgh 
or some other collector of that period. Gamble, Fl. Madras, 894 ( 1923 ) , 
reports the species from the Anamalai Hills, Madras. The only Tourne- 
jortia I have seen from that general region is T. Heyneana ! Until un- 
doubted material from Southern India is forthcoming I believe that 
T. Wightii should be accepted as clearly applying to the material east of 
the Ganges here discussed. In the type-form of T. Wightii the leaf- 
surface is velvety beneath much as in typical T. viridiflora. The corolla 
is much larger, however, with the tube 2-4 times as long as the calyx. 
Tournejortia Roxburghii is a form of T. Wightii which has lanceolate 
rather than ovate leaf-blades. It is a rather common form. I have seen 
plants similar to the type-form of T. Wightii and T. Roxburghii from 
throughout the range of T. montana. 



156 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

Tournejortia Candollii is based upon "T. Hcyneana, DC. Prodr. ix. 
516; Wall. Cat. 910, as to the Khasia examples." In the Wallich Cata- 
logue no. 910 consists of two parts, 1. material from Heyne, the type 
of T. Heyneana Wall, and 2. material collected by de Silva at Pundua. 
DeCandolle's specimen of Wallich 910 consists only of the second part 
of the Wallich number, that is to say, the material from Pundua by de 
Silva. This specimen was described by DeCandolle as T. Heyneana. 
Clarke, 1. c. 145, recognizing that the name T. Heyneana was obviously 
to be associated with Heyne's material from southern India, gave a new 
name, T. Candollii, to the plant improperly described as T. Heyneana by 
DeCandolle. The type of T. Candollii is accordingly de Silva's material 
in the DeCandollean Herbarium. The specimen at Geneva is broken 
and poor but has good corollas. These are somewhat constricted at the 
throat and very similar to those found in the type of T. khasiana. The 
leaves are lanceolate, dried brown beneath and nearly black above. They 
are very sparsely strigose above and have only scattered hairs along the 
principal veins beneath. 

I consider T. Candollii to be the form of T. montana with elongate 
corolla-tubes and glabrescent leaves. Belonging with it are several fur- 
ther synonymous forms. The type of T. Satnpsoni is from Sai-chii-shan 
caverns in the province of Kwangtung and is now deposited at the British 
Museum. There is some interesting variation within this collection. 
The corolla-tube is medium to long (5-8 mm.) and the lanceolate leaves 
are either distinctly appressed hairy or are quite glabrous beneath. The 
type of T. khasiana was collected by Clarke (no. 15227) at Nonpriang 
in the Khasia Hills. It is a form of T. Candollii in which the corolla- 
tube is contracted upward toward the throat so that the throat is at 
times almost half the diameter of the base of the tube. The type of 
T. Gaudichaudii is a glabrescent plant with elongate corollas and broadly 
lanceolate leaves. It was collected in Anam (Tourane) by Gaudichaud. 

7. Tournefortia luzonicasp. nov., scandens grisea; ramulis obscure 
tetragonis 2-4 mm. crassis pilis numerosis brevibus divergentibus ves- 
titis; foliis oppositis vel suboppositis; petiolis 5-14 mm. longis; lamina 
folii ovata vel late lanceolata 5-13 cm. longa 2-7 cm. lata apice breviter 
acuminata basi rotunda vel (1-4 mm. profunde) cordata, supra pilis 
rigidulis brevibus ascendentibus plus minusve numerosis vestita, subtus 
pallidiore pilis gracilibus falcatis saepe numerosis vestita nervis 6-9- 
jugatis ornata; inflorescentia hispidula; calycibus sessilibus 1-2.5 mm. 
altis, lobis anguste lanceolatis vel linearibus erectis; corolla virescenti- 
bus, tubo 2-4 (-8) mm. longo, limbo 2-2.5 mm. lato; fructu globoso 



1935] JOHNSTON, STUDIES IN BORAGINACEAE, XI 157 

3-4 mm. diametro albo glaberrimo succoso; nuculis 2 biovulatis laevibus. 

Endemic to the Philippines where it is confined to the mountainous 
regions of northern, east-central and southern Luzon. 

Specimens Examined: vicinity of Penablanca, Cagayan Prov., a vine 
on hillside, fl. green, fruit white, May 3, 1917, M. Adduru 237 (type, herb. 
Arnold Arboretum; isotype, Kew) ; Penablanca, 1926, Ramos & Edafio 
46662 (BM) ; Bangui, Prov. Ilocos Norte, Ramos 27563 (BM) ; Burgos, 
Prov. Ilocos Norte, Ramos 4799 (BD) ; Bocana del Abra, Prov. Ilocos 
Sur, Micholits (K); Mt. Pulog, Mountain Prov., Jan. 1909, Curran, 
Merrill & Zschokke 16103 (BD) ; Benguet, Loher 1541, 1542 (K) ; dist. 
of Lepanto, Mountain Prov., Vidal 3326 (K) ; Baguio, Benguet, Elmer 
8467 (AA, K) ; Mt. Maquilong, Prov. Batangas, Vidal 3327 (K) ; Prov. 
Albay, Cuming 1215 (K, BM, BD). 

7A. Tournefortia luzonica var. sublucens,var. nov., a forma typica 
specie! differt foliis sparse inconspicueque pubescentibus, supra vix 
griseis sed sublucentibus. 

Confined to the mountains of west-central Luzon. 

Specimens Examined : Anuling, Zambales Prov., 1924, Ramos & Edaho 
44553 (TYPE, herb. Arnold Arboretum: isotypes, Kew, Brit. Mus.) ; Zam- 
bales, 1907, Ramos 4799 (BD) ; Lamao, Mt. Mariveles, Bataan Prov., 
Meyer 2844 (K, BD) ; Lamao River, Mt. Mariveles, 350 ft. alt., slender 
vine growing over trees for many yards, Williams 525 (K). 

Among all the Old World species of Tournefortia this species is unique 
in the possession of opposite or subopposite leaves. In the treatments 
of the Philippine Boraginaceae by Robinson, Philip. Journ. Sci., Bot. 
4: 694 (1909), and by Merrill, Enum. Philip. PI. 3: 376 (1923), this 
plant has generally passed as T. Horsfieldii Miquel. That species, 
however, with its alternate leaves and a fruit composed of four uniovu- 
late nutlets is one of the forms of the widely ranging T. sarmentosa. 

The var. sublucens is confined to the mountainous country of west 
central Luzon, prov. Bataan and Zambales, and seems to have a range 
quite distinct from the typical form of T. luzonica which ranges in the 
other parts of the island of Luzon. Essentially a glabrate form of 
T. luzonica, with the upper leaf-surfaces more or less glossy, it is signifi- 
cant and worthy of nomenclatorial recognition only if it has a range 
apart, and is geographically correlated. 

8. Tournefortia Muelleri, nom. nov. Tournefortia mollis F. v. 
Mueller, Frag. 1:59 (1858); Bentham, Fl. Austral. 4:390 (1869); 
Bailey, Queensland Fl. 4: 1041 (1901); not T. mollis Bertol. (1852). 

Northern Australia and Papua. 



158 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

(K) ; Cape York Peninsula Exped., Hann 146 (K) ; shores of Montague 
Sound, W. Australia, 1820, Cunningham 182 (K, BM) and 324 (BM) ; 
erect shrub 1.5-2 m. tall, fringing tidal areas, Kapa Kapa, Papua, Brass 
505 (AA, K) ; Port Moresby, Papua, 1918, White (5 (AA). 

The carpels seem to be more bony than in other Old World species of 
this section. The leaves are usually lanate. 

9. Tournef ortia pubenila Baker. Jour. Linn. Soc. London, 20: 211 
(1883). Tournejortia Mocquerysi A. DeCandolle, Bull. Herb. Boiss. 
ser. 2, 1: 581 (1901). 

Forests of eastern Madagascar and the Seychelles. Possibly intro- 
duced in the latter archipelago. 

Specimens Examined: Madagascar: forests east of Ivohibe, 1000 m. 
alt., fl. white, Nov. 3, 1924, Humbert 3163 (P) ; high valley of the Rienana, 
drainage of the Matitana, 1000-4000 ni. alt., fl. white, Nov. 1924, Humbert 
2523 (P) ; Central Madagascar, Baron 1957 (Kew, type of T. puberula; 
BM, BD, isotypes), 2798 (K, BM, P), 3106 (K, P) and 6991 (K) ; forest 
of Ivohimanitra, Nov. 8, 1894, Forsyth Major 64 (K, BM, BD, P) ; forest 
of Analamazaotra near col d'Amboasary, ca. 950 m. alt., shrub with white 
flowers, Oct. 23, 1912, Viguicrro & Humbert 978 (P) ; forest at head of 
Antongil Bay, a liana with white flowers, Mocqiierys 161 (Deles., type 
of T. Mocquerysi). Seychelles : Mahe, common shrubby climber in hills 
near streams, Sept. 1871, Home 247 (K) ; Mahe, Thomasset (K) ; Mahe, 
Thoma^set 10 (BM) ; Mahe. 1867, Wright (B.M); Terne, Mahe, 1908, 

indefinite, a twining shrub generally on rocks near rivers, May 1902, 
Thomassett 22 (K, BM). 

The types of T. puberula and T. Mocquerysi are quite indistinguish- 
able. The species is a readily recognizable one. The leaves are firm, 
apparently glabrous and the stems are covered with a minute brownish 
puberulence. There is both a short- and a long-corolla form. The plant 
of the Seychelles is certainly identical with that of Madagascar. Pos- 
sibly it represents a horticultural introduction to the islands. In 
accounts of the Seychelles flora, Baker, Fl. Mauritius and Seychelles 202 
(1877), and Summerhayes, Trans. Linn. Soc. London, Zoology, 19: 284 
(1931), the species has consistently been misdetermined as T. sar- 



9A. Tournefortia puberula var. Kirkii, var. nov., a varietate genu- 
ina differt pilis brevibus pallidis adpressis ornatis. 

Islands off the northwest coast of Madagascar. 

Specimens Examined: Mohilla Island, Comoro Archipelago, April 
1861, /. Kirk as '•Tournefortia (3)" (type, herb. Kew) ; Nossi-be, June 
1847, Boivin 2086 (P) ; Nossi-be. 1853. Pcrrillc (P). 

This variety comes from a much more arid region than typical T. 



1935J JOHNSTON, STUDIES IN BORAGINACEAE, XI 159 

puberula and may be only a hairy xerophytic form of that species. In 
typical T. puberula the plant is provided with a minute, frequently some- 
what golden puberulence. In the var. Kirkii the stems have a sparse 
pale short strigosity that tends to disappear with age. The petioles are 
sparsely strigose. The lower surface of the leaf-blades has short white 
closely appressed hairs scattered along the rib and veins. The upper 
surface is somewhat strigose but less abundantly so than below. The 
inflorescence has numerous short ascending pale hairs. 



10. Tournefortia acuminata DeCandolle, Prodr. 9:520 (1845); 
Cordemoy, Fl. Reunion 479 (1895). 

Endemic to the Island of Reunion (Bourbon). 

Specimens Examined: les hauts du Boucan Launav, Boivin 1241 (K, 
BD, DC, Boiss, P) ; Bebour au dessus de la plantation de Quinquinas, 
July 28, 1875, G. de I' Isle 499 bis (K, Coss.) ; Bourbon, arbor, [ ICommcr- 
son^ (herb. Smith) ; chemin que conduit de Sante Rose a Saint Joseph 
avaunt la descente qui conduit au Volcan, 1812, Comnierson (P) ; 'Tile de 
France au bourbon," ex Mus. Paris (type, herb. DC). 

The type of T. acuminata at Geneva is given as distributed from Paris 
in 1821 and as from either Reunion or Mauritius. It represents a form 
in which the stems, petioles and inflorescence are glabrous or only very 
scantily strigose. It is obviously a duplicate of the collection at Paris 
which is labeled as collected by Commerson on the road between Ste. 
Rose and St. Joseph on Reunion. The material which I have cited from 
Boivin, which is widely distributed in European herbaria, is a form in 
which the stems, petioles and inflorescence have a short and evident, 
though not very abundant nor very conspicuous strigosity that becomes 
more or less brownish or golden. This I believe is the common form of 
the species. The leaves in T. acuminata are 12-1.7 cm. long and 3.5-7 
cm. broad, are acute at both ends, and have 10-15 pairs of nerves. The 
calyx is 1.5-2 mm. long at anthesis and has erect, cuneate or more or 
less lanceolate lobes. The corolla-tube is 3.5-7 mm. long. The limb is 

11. Tournefortia arborescens Lamarck, Tab. Encyc. 1:417 
(1791); Poiret, Encyc. 5:357 (1804). Tournefortia velutina Smith 
in Rees, Cyclop. 36: sp. no. 13 (Aug. 1817!), not T. velutina HBK. 
(1818). Tournefortia Bojeri sensu Cordemoy, Fl. Reunion 479 
(1895). 

Endemic to the Island of Reunion (Bourbon). 

Specimens Examined: Grand Bassin, Aug. 6, 1875, G. dc VIslc 454 
(K, P); Gauteuron (spelling ?) du Gol, woods, fl. white, Commerson 
(herb. Smith, type of T. velutina): Reunion, Commerson (herb. Smith, 



JOURNAL OF THE ARNOLD ARBORETUM 



[f Sonnerat] (Paris, type of T. arborescens). 

The type material of T. arborescens> is accompanied by a small label 
reading: "Tournefortia d I'inde." The collector is not indicated but 
both Lamarck and Poiret attribute it to Sonnerat who visited the Masca- 
renes during his voyage to India and Malaysia. The material consists 
of two sheets, one bearing a sterile shoot with entire oblanceolate leaves 
more or less tomentose beneath in the manner common in the spicate 
Cordia species of the section Varronia. The second sheet contains a 
Tournefortia in flower. The latter is a form of the species as here defined, 
having the leaves only very sparsely and obscurely strigose, particularly 
above. The stems bear numerous but not very abundant short appressed 
pale hairs. The calyx-lobes are ovate, acute and sparsely pale strigose. 
The specimen evidently represents the sparsely hairy form of the endemic 
species of Reunion. 

The type of T. velutina is the form of the species with very abundant 
long hairs. It has the leaves pale and silky with a dense indument of 
slender very pale hairs. The calyx lobes are ovate, densely hairy and 
more or less golden tawny. Smith mentioned atypical material of his 
T. velutina from Mauritius, but this, in fact, represents a form of T. 
Bojeri. DeCandolle, Prodr. 9: 514 (1845), incorrectly cited T. velutina 
as a possible synonym of T. argentea. I have cited above a specimen 
given as collected on Mauritius by Bojer. I doubt the accuracy of the 
geographical data and believe that the specimen is really from Reunion. 
Its broad calyx-lobes are ovate or orbicular-ovate and hence similar to 
those found in all material indubitably from that island. 

12. Tournefortia Bojeri A. DeCandolle, Prodr. 9:516 (1845); 
Baker, Fl. Mauritius, 202 (1877). Tournefortia bifida sensu Bojer, 
Hort. Maurit. 234 (1837). 

Endemic to the Island of Mauritius (He de France). 

Specimens Examined: Mauritius, woods, 1837, Bojer as T. bifida 
(TYPE, herb. DC) ; without locality, 1839, Bouton as T. bifida (DC, co- 
type) ; Mauritius, mountains and forest, Bouton (K) ; Mauritius 1854, 
Boivin (K) ; Mauritius, 1811, Hardwick (BM) ; Mauritius, ICommer- 
sonl (herb. Smith) ; Mauritius, Sieber98 (BD) ; Mauritius, herb. Labillar- 
diere (Deles); "Bourbon," 1853, Boivin (Boiss); 

Boivin's collection which it cited above and attributed to Reunion 
is, I believe, mislabeled. Indubitable collections of T. Bojeri come only 
from Mauritius. The species is very closely related to T. arborescens 
of Reunion, differing chiefly in the narrower calyx-lobes. In the DeCan- 
dolle Herbarium there is a branch of T. Bojeri, mounted on a sheet with 



1935] JOHNSTON, STUDIES IN BORAGINACEAE, XI 161 

isotypic material of the Philippine T. UrvUleana. The only label accom- 
panying this mixed sheet is in the script of Chamisso and belongs to the 
Philippine species. This mixed sheet makes comprehensible DeCan- 
dolle's, Prodr. 9: 515, adnot. (1845), strange comparison of T. Bojeri 
and r. UrvUleana. Since Chamisso never visited the Mascarenes it is 
evident that the spray of T. Bojeri has somehow become divorced from 
its proper label. The name "T. cymosa Heyne" seems to be based upon 
material from Mauritius. For a discussion of this nomen see my list of 
doubtful and excluded species on p. 166. 

In T. Bojeri the stems, petioles and inflorescence are more or less 
velvety with a pale ascending or spreading (or very rarely appressed) 
usually abundant hairs. The leaf blade is acute at both ends, more or 
less strigose on both surfaces though usually less so above. It has 10-12 
pairs of veins and is 11-1 7 (-24) cm. long and 2-5 (-6) cm. broad. The 
calyx is 2-2.5 mm. long and has the lobes cut at least ^ way to base. It 
is more or less silky strigose. The lobes are lanceolate to broadly lanceo- 
late or cuneate-lanceolate. The corolla-tube is 2-8 mm. long, and 2-5 
times the length of the calyx. The corolla-limb is 2-3 mm. broad. The 
fruit is ca. 3 mm. in diameter. 

Messerschmidia Linnaeus ex Hebenstreit, Nov. Comment. Acad. Sci. 
Imp. Petrop. 8: 315, tab. 11 (1763); Gmelin, Fl. Sibir. 4: 77 (1769); 
Murray, Syst. Nat. ed. 13, 161 (1774); Linnaeus fil. Suppl. PI. 132 
(1781). — type-species, Tournejortia sibirica Linn. Messersmidia 
Linnaeus, Hort. Upsal. 36 (1748) ; Linnaeus, Mant. 1 : 5 and 42 (1767); 
Linnaeus, Syst. ed. 12,149 (1767); Linnaeus, Mant. 2: 334 (1771).— 
a variant spelling of Messerschmidia, type-species, Tournejortia sibirica 
Linn. Tournejortia sect. Messerschmidia (Linn.) DeCandolle, Prodr. 
9: 528 (1845); as to nomenclatorial type only, not as to the species 
of Heliotropium treated. Argusia Amman, Stirp. Rar. Ruth. 29 (1739). 
Arguzia [Amman] Rafinesque, Sylva Tellur. 167 (1838); Steven, Bull. 
Soc. Nat. Moscow 24> : 558 (1851). — type-species, Tournejortia sibi- 
rica Linn. Tournejortia sect. Arguzia [Amman] DeCandolle, Prodr. 9: 
514 (1845); Ledebour, Fl. Ross. 3:97 (1847-49). — type-species, 
Tournejortia sibirica Linn. Tournejortia sect. Mallota A. DeCandolle, 
Prodr. 9: 514 (1845). — type-species, T. argentea Linn. Tournejortia 
sect. Mallotonia Grisebach, Fl. W. Ind. 483 (1861). — type-species, 
Tournejortia gnaphalodes R. Br. ex R. & S. Mallotonia (Griseb.) 
Britton, Ann. Mo. Bot. Gard. 2:47 (1915). — type-species, Tourne- 
jortia gnaphalodes R. Br. 

Segregated here, as the emended genus Messerschmidia, are three 



162 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

remarkable species that depart widely in appearance from the numerous 
and habitually very uniform species formerly associated with them in 
Tournejortia. As I have redefined and amplified Messerschmidia it con- 
sists of the original Asiatic herb, Tournejortia sibirica, the strand-shrub 
of the Antilles, T. gnaphalodes, and the well known strand-tree of the 
Indian and Pacific oceans, T. argentea. All these species differ widely 
not only in their habit of growth and in their selection of habitat from all 
the other species that have been traditionally placed with them in Tour- 
nejortia, but also in their pronounced development of a corky exocarp 
which sets them off not only from all species of Tournejortia but from 
all other Boraginaceae as well. All three of the species show a marked 
preference for saline conditions. Two of them are tropical strand-plants. 
The third species grows along the ocean in temperate eastern Asia, in 
more or less saline soils along streams and about inland seas in Central 
Asia and eastern Europe. The corky exocarp evidently adapts the three 
species for water dispersal. The nature of the hairy covering of these 
three species is of an essentially similar type, consisting of slender silky 
hairs rather different in texture and appearance from that predominat- 
ing among the species of true Tournejortia. 

The generic name Messerschmidia (also spelled Messersmidia and 
Messerschmidtia) is based upon Tournejortia sibirica Linn., and is a 
synonym of Argusia (or Arguzia). The type-species was first described 
by Amman in 1739 who applied to it the mononomial, Argusia, and gave 
a lengthy description of it based upon notes and specimens made by 
D. G. Messerschmid in 1724 along what is now the northwestern frontier 
of Manchuria. The source of this material is given as "Locus in glareosis 
aridisque apricis Argun fluuii et like Dalai Noor in Dauria." Although 
Amman's mononomial was formed from the "loco natali" of the plant, 
i.e. the Argun River, its author deliberately and repeatedly spelled it 
" Argusia" [ Amman states that seeds from Messerschmid's collection 
germinated and grew in the gardens at St. Petersburg. These same cul- 
tures are probably those described and illustrated by Hebenstreit in 
1763. The plants growing in the Upsala Garden in 1748 and described 
by Linnaeus as Messersmidia were probably derived from those grown 
by Amman. In the Correspondence of Linnaeus, ed. Smith 2: 200 
(1821), there is a letter from Amman, dated Nov. 18, 1740, in which 
questions by Linnaeus concerning Argusia are answered and in which 
it is stated that dried specimens of Argusia were being sent him. When 
he proposed the mononomial "Argusia," Amman justified his use of a 
geographic appellation in forming the name, but added that he had no 
objection if the genus was named after Messerschmid, its original col- 



1935] JOHNSTON, STUDIES IN BORAGINACEAE, XI 163 

lector. Linnaeus seems to have preferred the latter. The collector's 
name was spelled "Messerschmid" by his contemporaries. Linnaeus 
latinized it, "Messersmidia," and was consistent in this usage in all his 
writings. Other writers of the last half of the 18th century, however, 
spelled it "Messerschmidia" and it is so spelled in the paper by Heben- 
streit who was the first to use the generic name subsequent to 1753. 
Writers of the past century tended to spell the generic name "Messer- 
schmidtia." The generic name Messerschmidia has variant spellings in 
"Messersmidia" and "Messerschmidtia." Although clearly based upon, 
in fact named after the original collector of Tournefortia sibirica, the 
generic name Messerschmidia (variously spelled) eventually became 
associated with two other very diverse groups of Boraginaceae. A study 
of the facts here presented, however, makes it evident that the name 
"Messerschmidia" is only very improperly applied either to the Ameri- 
can species of Tournefortia sect. Cyphocyema, or to Canary Island and 
South African species of Heliotropium as has been done in some large 
works. In another paper, Contr. Gray Herb. 92: 73 (1930), I have 
given many facts concerning the misuse of the name "Messerschmidia" 
The name was based upon Tournefortia sibirica and was originally 
applied solely to that plant. The type-species of Messerschmidia is 
obviously and logically the original Siberian species. 



Plant a low herb from rhizomes; inflorescence a loose open 
corymbose cyme; calyx pedicellate, lobes cuneate ; anthers 
several times as long as broad ; fruit pubescent, sunken in at 
apex; carpels embedded in the center of the corky exocarp; 

temperate Eurasia 1- ^- ^tbinca. 

Plant a tree or shrub; inflorescence of scorpioid cymes; calyx 
sessile, lobes orbicular or oblong ; anthers about two times as 
long as broad; fruit glabrous, apex conic or rounded ; carpels 
occupying the apical half of the fruit, the lower half com- 
posed entirely of corky exocarp ; tropical strand plants. 
A tree 1-5 m. tall; leaves broadly oblanceolate or obovate, 
3.5-9 cm. broad; inflorescence a conspicuous stif¥ panicle 
of loosely flowered elongating (up to 8 cm.) scorpioid 
cymes; corolla-lobes merely imbricate (not plicate) in the 
bud; anthers partially exserted from the short corolla- 
tube'; fruit dull, breaking in half ; apex and dorsal surface 
of carpels covered with corky exocarpial tissue ; tropics of 

the Old World 2. M. argcntca. 

A shrub 3-12 dm. tall ; leaves narrowly spathulate-linear, 4-10 
mm. broad; inflorescence consisting of single or paired 
long-peduncled very congested short (1-2 cm.) scorpioid 
cymes ; corolla-lobes distinctly plicate in bud ; anthers well 



164 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

included in the cylindrical corolla-tube ; fruit brown and 
lustrous, not breaking in half; apex and dorsal surface 
of carpels not covered with corky exocarpial tissue ; West 
Indies Z, M. gnaphdodes. 

1. Messerschmidia sibirica Linnaeus, Mant. 2: 334 (1771). Tour- 
nefortia sibirica Linnaeus, Sp. PI. 141 (1753); Kusnezow & Popow, FL 
Caucas. Crit. 4-: 77 (1913). Messersmidia Argusia Linnaeus, Mant. 
1 : 42 (1767). Messerschmidia Arguzia Murray, Syst. Nat. ed. 13, p. 
161 (1774); Linnaeus fil. Suppl. PI. 132 (1781). Messerschmidia 
Argunia Gaertner, Fruct. 2: 130, tab. 109 (1791). Tournejortia 
Arguzia (L.) Roemer & Schultes, Syst. 4: 540 (1819); Ledebour, FL 
Ross. 3:97 (1847-49); Herter, Act. Hort. Petrop. 1:503 (1872). 
Messerschmidia rosmarini folia Willdenow ex Roemer & Schultes, Syst. 
4:544 (1819). Tournejortia rosmarinijolia Willdenow ex Steudel, 
Nomen. ed. 2, 2 : 693 ( 1841 ) . Tournejortia Arguzia var. rosmarinijolia 
(Willd.) Turczaninow, Bull. Soc. Nat. Moscow 23' : 498 (1850). Argu- 
zia rosmarinijolia Steven, Bull. Soc. Nat. Moscow 24': 559 (1851). 
Arguzia repens Rafinesque, Sylva Tellur. 167 (1838). Tournejortia 
Arguzia var. latijolia DeCandolle, Prodr. 9: 514 (1845); Turczaninow, 
Bull. Soc. Nat. Moscow 23': 498 (1850). Tournejortia Arguzia var. 
angustior DeCandolle, Prodr. 9: 514 (1845); Turczaninow, Bull. Soc. 
Nat. Moscow 23': 498 (1850). Tournejortia sibirica var. angustior 
Turczaninow ex Fedtchenko, Consp. Fl. Turkestan 5:39 (1913). 
Tournejortia Arguzia var. cynanchoides Turczaninow ex Steven, Bull. 
Soc. Nat. Moscow 24' : 559 ( 1851 ) , in synon. Arguzia Messerschmidia 
Steven, Bull. Soc. Nat. Moscow 24': 560 (1851). Arguzia cimmerica 
Steven, Bull. Soc. Nat. Moscow 24' : 560 (1851). Heliotropium 
japonicum Gray, Mem. Amer. Acad. ser. 2, 6: 403 (1859). 

From Japan, Amur and northern China across Asia, mostly between 
lat. 40° and 55° N., to Rumania and central Russia; affecting moist 
gravelly, usually saline soils. For more details on distribution see Lede- 
bour, Fl. Ross. 3': 97 (1847-49); Herter, Act. Hort. Petrop. 1: 503 
(1872) and Kusnezow & Popow, Fl. Caucas. Crit. 4-': 78 (1913). 

2. Messerschmidia argentea (Linnaeus), comb. nov. Tournejortia 
argentea Linnaeus 1^1., Suppl. PI. 133 (1781). Tournejortia arborea 
Blanco, Fl. Filip. 129 (1837). 

A strand-tree widely distributed within the tropics, on islands in the 
Indian and Pacific oceans. 

The distribution of this species is worthy of a detailed statement. The 
fruit having a corky exocarp is admirably suited for oceanic dispersal. 
In this it has been very successful. The species is, in fact, one of the 



1935] JOHNSTON, STUDIES IN BORAGINACEAE, XI 165 

characteristic strand-plants of the Old World Tropics. It is, however, 
almost exclusively a plant of island-shores. In the Pacific Ocean it 
ranges from the Paumotas (Ducie Isl.), the Marquesas and Palmyra 
Island, westward to Bonin Island, the Liu Kiu Islands, Formosa, Tizard 
Reef (China Sea), "Annam (Turan)," the Philippines, the Moluccas, 
Timor, tropical Australia, and New Caledonia. In the Indian Ocean it 
extends from northwestern Australia, Timor and Java, Christmas and 
Cocos Keeling islands to the Mascarenes, Madagascar (near south end 
only) and coast of Mozambique (rare), north to Zanzibar, the Sey- 
chelles, the Laccadives (Bitrapar in lat. 11°30' N.), the Maldives, Cey- 
lon, the Andamans (Great Coco Isl. in lat. 14° N.), the Nicobars, the 
islands (Vogel, lat. 7°46' N.; Adang calat, lat. 6°30' N.) off the west 
coast of peninsular Siam, and the northwestern Federated Malay States 
("Kedah"). Miquel, Prodr. Fl. Sumatra, 244 (1855), reports it vaguely 
from Sumatra. I have seen no material from Sumatra, Borneo or the 
Celebes. Except for the record from Indo-China and the vague record 
for the Malay States the species is not known from the Asiatic continent. 
In Africa it is reported only from the Mozambique Coast. The record 
for Amboland (Schinz 757), found in the Flora of Tropical Africa, 4^ : 30 
(1905), is evidently a clerical error for the specimen cited is Helio- 
tr opium tuberculosum! 

The original description of T. argcntea is based upon material col- 
lected on the coast of Ceylon by Konig. In the Linnean Herbarium 
there is a characteristic specimen of this plant accompanied by Konig's 
label reading "habitat ad Littora maris Zeylanica." Accompanying this 
is a label in the script of the younger Linnaeus reading "Konig, 1777." 
The Buglossum lanuginosum of Rumphius, cited by the younger Lin- 
naeus, is evidently conspecific with Konig's material from Ceylon. 
DeCandolle, Prodr. 9: 514 (1845), cites T. velutina Smith as a possible 
synonym of T. argentea. Smith's plant, however, is a very different 
species being a synonym of T. arborescens Lam. of Reunion. 

3. Messerschmidia gnaphalodes (Linnaeus), comb. nov. Helio- 
tropium gnaphalodes Linnaeus, Syst. ed. 10, p. 913 (1759) and Amoen. 
Acad. 5:394 (1760). Tournejortia gnaphalodes (L.) R. Brown ex 
Roemer & Schultes, Syst. 4: 538 (1819). Mallotonia gnaphalodes (L.) 
Britton, Ann. Mo. Bot. Card. 2: 47 (1915). 

A strand-plant widely distributed in the West Indies. 

According to Millspaugh, Publ. Field Mus., Bot. 2: 89 (1900), this 
species grows "On the beach line facing the open sea, [and is] very 
seldom, if ever, found in bays or where partially dry reefs guard the 



166 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

shore." Its range may be stated as follows: Bermuda, the Bahamas 
(north to about lat. 2 7° N.), southern peninsular Florida (to Miami 
region) and the Greater Antilles, southward to Granada (in the Lesser 
Antilles), the islands off Venezuela, the Paraguana Peninsula of north- 
western Venezuela (Medanos Isthmus) and westward to Alacran Reef 
(north of Yucatan), the coasts of Yucatan, Cozumel Island, and Swan 
Island (off Honduras). The species has been reported from the "So- 
ciety Islands" in Polynesia by Hooker & Arnott, Bot. Beechey Voy. 
67 (1832), but as Drake, Fl. Polynes. Frang. 130 (1893), has stated 
this is probably the result of some error. This West Indian species is 
certainly not to be expected in the South Pacific. 

This species was founded by Linnaeus entirely upon an illustration 
and phrase-name given by Plukenet, Phytogr. tab. 193, fig. 5 (1691). 
This basic phrase-name is as follows: "Heliotropium gnaphaloides 
litoreum fruticescens Americanum Sea Lavender, Barbadensibus dic- 
tum." From it we may suppose that Plukenet's material came from 
the Barbados. 

Doubtful and Excluded Names 
Tournefortia angustifolia (Lam.) Roemer & Schultes, Syst. 4: 539 
(1819). — Hcliotropium messersckmidioides Kuntze. 

Tournefortia angulosa Desfontaines, Tab. ed. 2, 85 (1815). — A 
bare name in a garden-list. 

Tournefortia bifida Lamarck, Tab. Encyc. 1:417 (1791); Poiret, 
Encyc. 5: 360 (1804); Poiret, Diet. Sci. Nat. 41: 177 (1826); Smith 
in Rees, Cyclop. 36: sub sp. no. 25 (1819); Baker, Fl. Mauritius, 202 
(1877). — The type of this species was collected on ITle de France by 
Commerson and represents Antirhea jrangtdacea DeCandolle, Prodr. 
4: 460 ( 1830) ! The correct name for this Mauritian species of Rubia- 
cece is, accordingly, Antirhea bifida (Lam.), comb. nov. 

"Tournefortia cjonosa Heyne In Herb. Rottler, not of Linn." ex 
Clarke in Hooker, Fl. Brit. India 4: 145 (1883). — This reference con- 
cerns a small specimen at Kew which may possibly represent T. Bojcri 
of Mauritius. It is certainly not T. i/evneawa as given by Clarke! The 
specimen bears a printed label reading: "Herbarium Rottlerianum; 
Penins. Indiae Orientalis; Presented by the Council of Kings College, 
Feb. 1872." Accompanying this are two labels in script giving the 
determination as T. cymosa Swartz and the collector as Mace. Clarke's 
citation accordingly is merely a reference to a misdetermined specimen 
in the Kew herbarium. 



1935] JOHNSTON, STUDIES IN BORAGINACEAE, XI 167 

Tournefortia Edgeworthii A. DeCandolle, Prodr. 9: 529 (1845). — 
Heliotropium zeylanicum Burman. 

Tournefortia fruticosa (Linn, f.) Ker, Bot. Reg. 6: tab. 464 
(1820). — Heliotropium messerschmidioides Kuntze. 

Tournefortia linearis E. Meyer in Drege, Flora 26- : Beigabe p. 
57 and 226 (1843), nomen. — Heliotropium lineare (E. Meyer) Wright. 

Tournefortia Messerchmidia Sweet, Hort. Suburb. London 31 
(1818), nomen subnudum. — Heliotropium messerschmidioides Kuntze. 

Tournefortia micranthos (Bunge) A. DeCandolle, Prodr. 10: 67 
(1846); Ledebour, Fl. Ross. 3:98 {\M1-A9). — Heliotropium 
micranthos (Bunge) Boissier. 

Tournefortia mollis A. Bertoloni, Misc. Bot. 12:44, tab. 1 
(1852). — Based upon material from Mozambique representing Van- 
gueria tomentosa Hochst. This species of Bertoloni's is not mentioned 
in Robyn's recent monograph of Vangueria, Bull. Jard. Bot. Brux., vol. 
11 (1928). 

Tournefortia mutabilis Ventenat, Choix PI. tab. 3 (1803). — The 
basis of this species was given by Ventenat as follows: "Arbrisseau origi- 
naire de Java, cultive chez Cels et au Museum d'Histoire Naturelle, de 
semences rapportees par La Haye." I have seen Ventenat's original 
material in the Delessert Herbarium at Geneva and duplicates of it at 
Kew, Berlin and Paris. The plant is evidently not a species of the Old 
World, in fact, it appears to be a form of the Mexican T. Hartwegiana 
Steud. Since Ventenat's name is much older than T. Hartwegiana it 
must be taken up in place of the latter. La Haye (or Lahaia) was a 
gardener who travelled to the East Indian Islands collecting seeds and 
plants which were subsequently grown at the garden of J. M. Cels and 
at the Jardin des Plantes at Paris. Since he is not known to have visited 
America it is evident that all of Ventenat's original data are incorrect. 

Tournefortia Royleana DeCandolle, Prodr. 9: 527 {\%AS).~ Helio- 
tropium zeylanicum Burman. 

Tournefortia stenoraca Klotzsch in Peters, Reise Mossamb. 250 
( 1861 ) . — Heliotropium zeylanicum Burman. 

Tournefortia subulata Hochstetter ex DeCandolle, Prodr. 9: 528 
(1844). — Heliotropium zeylanicum Burman. 

Tournefortia tuberculosa Chamisso, Linnaea 4:467 (1829). — 
Heliotropium tuberculosum (Cham.) GUrke. 



168 JOURNAL OF THE ARNOLD ARBORETUM [vol. 

Tournefortia zeylanica (Burman) Wight, Illust. Ind. Bot. 2: 2 
tab. 170 (l&SO). — Heliotropiumzeylanicum Burman. 

Messerschmidia angustifolia Lamarck, Tab. Encyc. 1 : 4 
(1791). — HeUotropium messerschmidioides Kuntze. 

Messerschmidia cancellata d'Asso, Synop. Aragon. 21, tab. 
(1779). — Rochelia species. 

Messerschmidia floribunda Salisbury, Prodr. 112 (1796). — HeUo- 
tropium messerschmidioides Kuntze. 

Messerschmidia fruticosa Linnaeus fil. Suppl. 132 ( 1781 ). — HeUo- 
tropium messerschmidioides Kuntze. 

Messerschmidia hispida Bentham in Royle, 111. Bot. Himal. Mts. 
306 (1836). — HeUotropium zeylanicum Burman. 

HeUotropium pannifolium Burchell ex Hemsley, Voy. Challenger, 
Bot. 2: 78 (1884). — This species from St. Helena is known only from 
Burchell's material. It is now probably exterminated. I have studied 
Burchell's unpublished drawing of the plant and his specimen at Kew. 
The plant is evidently a strong shrub much resembling the Eutourne- 
jortiae of the Andes in foliage and habit of growth. I know of no HeUo- 
tropium that could be recognized as a close relative of it nor one that 
could be said to resemble it in gross habit. Burchell's specimen is in 
the flowering condition only. The corollas, most unfortunately, have 
been almost completely eaten away by insects. There are consequently 
no reproductive structures on the type which might help in definitely 
placing the St. Helena plant generically. Since the plant is no doubt 
extinct and no further specimens are to be expected, the species will 
probably remain one of dubious generic affinities, and since a nomen- 
clatorial transfer would add nothing to our regrettably small knowledge 
of it, I am not giving this obscure plant a new name under Tournefortia. 
However, I do strongly suspect that it belongs in that genus. 

2. NOTES ON BRAND'S TREATMENT OE CRYPTANTHA 
The treatment of Cryptantha by Brand appeared in the second and 
posthumous volume he contributed on the Boraginaceae in "Das Pflan- 
zenreich." It is based almost exclusively upon the material available 
to him in the German herbaria. Having had no field experience in 
Western America and having had no opportunity to examine either the 
very numerous types or the great collections of Cryptantha in American 
herbaria, it is not surprising to find that Brand's treatment of the genus 
contains numerous errors arising from his restricted opportunities for 



1935] JOHNSTON, STUDIES IN BORAGINACEAE, XI 



the study of this large, difficult and characteristic West American genus. 
I have found some of Brand's statements very puzzling. Hence it is 
that during a recent visit in Germany I took the opportunity of study- 
ing the material available to him and, in the light of these studies, made 
copious annotations in copies of his published work on the Boraginaceae. 
The data given here concern Cryptantha and embody the notes correct- 
ing Brand's more serious errors, as well as those clarifying the more 
puzzling details of his work on that genus. In the following discussion, 
as a heading for the pertinent notes, I have given the specific name 
accepted by Brand and have preceded it by the number under which the 
species may be found in his treatment. Following these is an abbrevi- 
ated reference to the page of the Pflanzenreich, iv. 252- [Heft 97] pp. 
28-75 (1931), on which the given species may be found. 

5. Cryptantha macrocalyx (Phil.) Reiche; Brand, Pflanzenr. 30 
(1931). — The specimen cited and described is evidently a duplicate of 
the material to be found in Philippi's Herbarium at Santiago labelled as 
collected by San Roman in Quebrada de Serna. The material is so poor 
that I can add nothing to my previous discussion, Contr. Gray Herb. 
78: 70 (1927), of this peculiar plant. It is most certainly not C. 
macrocalyx. 

6. Cryptantha Buchtienii Brand, Pflanzenr. 30 (1931). — I have 
studied the type-specimen. It is a form of C. glomerata from a locality 
in which it has been repeatedly collected. 

7. Cryptantha phacelioides (Clos) Reiche; Brand, Pflanzenr. 31 
(1931). — Brand cites two specimens as seen. The collection by 
Philippi, from which Brand's description is derived, is an isotype of 
Eritrichium Rengijoanum Phil., which I consider to be a phase of C. 
aprica (Phil.) Reiche. I have not seen the material collected by 
Buchtien, which is cited by Brand, but suspect that it may be C. 
glomerulifera (Phil.) Johnston, which Buchtien obtained at 2400 m. 
alt. near Juncal. Neither the name used by Brand nor any of the sup- 
posed synonyms he lists belong to either of these species I have men- 
tioned. The synonyms he lists belong to three distinct species. 

9. CryTstantha talquina (Phil.) Brand, Pflanzenr. 52 (1931).— 
This species is unquestionably a synonym of C. alyssoides (DC.) Reiche. 
Brand attempted to separate it by stating that basal cleistogenes were 
present in C. talquina and absent in C. alyssoides. This is contrary to 
fact. An isotype of C. alyssoides in the DeCandolle Herbarium at 
Geneva shows a fine display of these cleistogenes. The type at Paris 
has had them all knocked off". 



170 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

12. Cryptantha candelabrum Brand, Pflanzenr. 33 (1931). — Based 
upon three collections, all from Philippi. These are: 1. Santiago, 
"Philippi (sub E. congestum), 2. Santiago, "Philippi dedit 1876" (sub 
E. congestum) and 3. Prov. de Santiago, "Philippi dedit 1876" (sub E. 
lineare; "Dimorphocarpum est"). The first specimen has flowers and 
fruit. The second and third show flowers only. For evident reasons the 
first specimen is selected as type. It is a form of C. linearis (Colla) 
Greene. The other specimens are quite similar and probably represent 
immature C. linearis or perhaps even C. aprica. 

13. Cryptantha fallax (Phil.) Reiche; Brand, Pflanzenr. 33 
(1931). — The specimen from Philippi, cited by Brand, seems to repre- 
sent C. Kingii (Phil.) Reiche. The chasmogamic flowers are in bud 
only. The label associated with the specimen is in the script of Philippi. 
There is a question mark following the locality, "La Serena." 

15. Cryptantha campylotricha Brand, Pflanzenr. 34 (1931).— 
This species is a synonym of C. Kingii (Phil.) Reiche. 

16. Cryptantha diffusa (Phil.) Johnston; Brand, Pflanzenr. 34 
(1931). — The single specimen cited, that collected by Philippi at 
Paihuano, represents C. globulijera (Clos) Reiche. 

17. Cryptantha modesta Brand, Pflanzenr. 35 (1931). — This spe- 
cies is a synonym of C. diplotricha (Phil.) Reiche. 

19. Cryptantha Vidalii (Phil.) Reiche; Brand, Pflanzenr. 35 
(1931). — The only specimen examined by Brand is one grown in the 
Berlin Garden. It seems to be a form of C. glomerata Lehm. It is of 
course not authentic C. VidalU. 

21. Cryptantha CandoUeana Brand, Pflanzenr. 36 (1931). — This 
species is based upon specimens from Macrae, Gay, Besser, and two from 
Philippi. At Berlin there is no specimen of this species collected by 
Gay in "Colchagua," but there is one given as from "Chile." All the 
material of this species cited by Brand represents forms of C. glomerata 
Lehm. 

25. Cryptantha globulif era (Clos) Reiche; Brand, Pflanzenr. 37 
(1931). — The only specimen cited by Brand seems to represent C. 
linearis (Colla) Greene. The specimen is immature. The corollas are 

28. Cryptantha capituliflora (Clos) Reiche, var. compacta Brand, 
Pflanzenr. 38 (1931). — The single specimen cited of this variety rep- 



1935] JOHNSTON, STUDIES IN BORAGINACEAE, XI 

resents a stunted 
does not appear t 

33. Crjrptantha barbigera (Gray) Greene; Brand, Pflanzenr. 39 
(1931). — Among the three specimens cited, those of Jones and of 
Heller represent this species. That collected by Greene represents 
typical C. nevadensis Nels. & Kenn. 

34. Cryptantha nevadensis Nelson & Kennedy; Brand, Pflanzenr. 
39 (1931). — The collection by Rusby, in the Dehra Dun Herbarium, 
which I examined while still on loan at Berlin, represents C. barbigera 
(Gray) Greene. 

38. Cryptantha affinis (Gray) Greene; Brand, Pflanzenr. 42 
(1931). — The material at Berlin of Heller 5882 is good C. Torreyana 
(Gray) Greene and that of Jones 856 is at least in part good C. affinis. 
The data on the latter collection is probably questionable. 

39. Cryptantha microstachys Greene; Brand, Pflanzenr. 42 
(1931). — The single specimen cited by Brand, Jones 3138 from San 
Diego, is C. Clevelandi Greene. 

40. Cryptantha Lyallii Brand, Pflanzenr. 42 (1931). — This is a 
synonym of C. flaccida (Dougl.) Greene, all the cited material falling 
readily into that species. 

49. Cryptantha Hossei Brand, Pflanzenr. 45 (1931). — This is an 
evident synonym of C. diplotricka (Ph.) Reiche. 

63. Cryptantha Famatinae Brand, Pflanzenr. 49 (1931). — The 
type of this species represents C. diffusa (Phil.) Johnston. 

66. Cryptantha parvula (Phil.) Brand, Pflanzenr. 50 (1931).— 
Of the three specimens cited, Philippi 694 is C. diffusa (Phil.) Johnston, 
that from Caldera is a form of C. globulijera (Clos) Reiche, and that 
from San Roman is C. diffusa. 

75. Cryptantha leiocarpa (Fisch. & Mey.) Greene, var. eremo^ 
caryoides Brand, Pflanzenr. 53 (1931). — This is apparently an odd 
form of C. leiocarpa. 

81. Cryptantha confusa Rydberg; Brand, Pflanzenr. 56 (1931).— 
Among the specimens cited, Leiberg 2271, is C. Watsoni (Gray) Greene, 
the remainder represents C. affinis (Gray) Greene. 

83. Cryptantha Fendleri (Gray) Greene; Brand, Pflanzenr. 57 
(1931). — Greene's material from Beaver Creek is C. ambigua (Gray) 



172 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

84. Crjrptantha Torreyana (Gray) Greene, Brand, Pflanzenr. 57 
(1931). — In the Berlin collections Rydberg & Bessey 48S5 and Heller 
9074 represent C. ambigua (Gray) Greene. 

85. Cryptantha Rattanii Greene; Brand, Pflanzenr. 58 (1931).— 
The cited material at Berlin is in flower only. The corolla is 3-5 mm. 
broad. It is probably a form of C. hispidissima Greene. 

86. Cryptantha grandiflora Rydberg, var. anulata Brand, Pflan- 
zenr. 59 (1931). — This is a form of C. Hendersoni (Xels.) Piper. 

91. Cryptantha hispidula Greene ex Brand, Pflanzenr. 60 (1931).— 
The type is Baker 2966 from Napa County. The collections from Elmer 
and Eastwood are C. Clevelandi var. florosa Johnston. 

91. Cryptantha hispidula var. Elmeri Brand, Pflanzenr. 60 
(1931). — The cited material represents one of the forms of C. Hender- 
soni (Nels.) Piper having a single polished nutlet. 

92. Cryptantha flaccida (Dougl.) Greene; Brand, Pflanzenr. 60 
(1931). — The specimen collected by Congdon, no. 72, near Soledad 
represents C. decipiens var. corollata Johnston. 

94. Cryptantha hispida (Phil.) Reiche; Brand, Pflanzenr. 61 
(1931). — The cited specimen is an isotype of the very different C. 
Romanii Johnston. 

96. Cryptantha albida (H. B. K.) Johnston; Brand, Pflanzenr. 63 
(1931). — Among the cited specimens Fendler 635 represents C. Fend- 
leri (Gray) Greene, and the collection by Echegaray represents C. diplo- 
tricha (Phil.) Reiche. 

106. Cryptantha granulosa (Ruiz & Pav.) Johnston; Brand, Pflan- 
zenr. 65 (1931). — Two of the cited collections, Weberbauer 5693 and 
5700, represent C. limensis (A. DC.) Johnston. 

107. Cryptantha Philippiana Brand, Pflanzenr. 66 (1931).— 
This is a form of C. glomerata Lehm. having a developed chasmogamic 
inflorescence. 

108. Cryptantha mirabunda Brand, Pflanzenr. 66 (1931). — I 
consider this species to be a synonym of C. utahensis (Gray) Greene. 

113. Cryptantha ambigua (Gray) Greene, forma robustior Brand, 
Pflanzenr. 69 (1931). — The material cited from California all repre- 
sents C. cchinella Greene. One collection by Howell, no. 48, is C. 
simulans Greene. 



1935] JOHNSTON, STUDIES IN BORAGINACEAE, XI 173 

115. Cryptantha Stuebelii Brand, Pflanzenr. 69 (1931). — The 
type of this species, from Yosemite Valley, is an equal mixture of C. 
murkata var. Jonesit (Gray) Johnston, and C. simulans Greene. Han- 
sen's collection seems to be young C. simtdans. 

121. Cryptantha Hansenii Brand, Pflanzenr. 71 (1931). — This 
represents one of the puzzling forms of C. intermedia (Gray) Greene 
found in the foothills of the central Sierra Nevadas. The variety 
pulckella Brand, is merely an immature specimen of this Sierran form. 



3. NEW OR OTHERWISE NOTEWORTHY SPECIES 

Cordia Weddellii sp. nov., arbuscula 3-4 m. alta laxe ramosa 
pilis malpighiaceis strigosa; ramulis pallide strigosis; foliis ellipticis 
2.5-4 cm. longis 1.5-2.5 cm. latis utrinque strigosis, nervis 7-10-jugis 
rectis parallelibus inconspicue sparseque ramosis, subtus pallidioribus, 
margine integris, apice rotundis, petiolis gracilibus 5-9 mm. longis; 
floribus ad apicem ramulorum in cymulis parvis 3-10-floris breviter 
pedunculatis affixis; calyce ad anthesim ca. 1 mm. longo, apice irregu- 
lariter disrumpente, extus strigoso obscure multisulcato; corolla alba 
ca. 3 cm. longa extus pilosa intus glabra, tubo ca. 1 cm. longo 3-4 mm. 
crasso, faucibus late ampliatis, lobis 5 suborbiculatis ca. 1 cm. longis 
rotundis ascendentibus in alabastro valde plicatis; staminibus 5, supra 
(4 mm.) basem tubi affixis, filamentis inaequalibus 5 et 6 mm. longis 
glabris; antheris oblongis 2-2.5 mm. longis; stylo 4 mm. profunda 
lobato basim versus sparsissime setifero, lobis 1.5 mm. lobulatis, lobulis 
spathulatis ; ovario glabro ; fructu ignoto. 

Bolivia : Prov. of Chiquitos, small shrub 3-4 m. tall at edge of forest, 
fl. white, Sept.-Oct. 1845, Weddell 3454 (type, Paris). 

A very remarkable species of the section Eucordia and related to C. 
aberrans Johnston (C. mucronata Fres.) and C. caruiida Veil. These 
two relatives come from the Brazilian coast near Rio Janeiro. Cordia 
Weddellii was collected in the extreme eastern section of the Dept. of 
Santa Cruz, Bolivia, and is distinguished at once by its malpighiaceous 
hairs. This type of pubescence is extremely rare in Cordia. In the 
present species it is particularly interesting since the mid-section of 
each hair (above where it is attached) is glandular and thickened. 

Cordia aberrans, nom nov. Cordia mucronata Fresenius in Mar- 
tins, Fl. Bras. 8' : 9 (1857); Johnston, Contr. Gray Herb. 92:42 
(1930); not Poiret (1818). 

The existence of an earlier homonym makes it necessary to rename 



174 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

this remarkable species. The type has been examined at Munich. It 
is labeled: "Inter Vittoria et Bahia; S. Princ. Maxim. Vidensis; Martius 
comunic. 1856." 

Cordia taguahyensis Vellozo, Fl. Flum. 98 (1825) and Icones, 2: 
tab. 154 (1827). Cordia amplijolia Mez, Bot. Jahrb. 12: 538 (1890) ; 
Johnston, Contr. Gray Herb. 92: 62 (1930); not A. DeCandolle (1845). 
Lithocardium Mezianum Kuntze, Rev. Gen. 2: 976 (1891). Cordia 
Meziana (Kuntze) Gurke in Engler & Prantl, Nat. Pflanzenf. IV. Abt. 
3a, p. 84 (1893). 

An examination of the type of C. amplijolia, at Munich, proves it to 
be simply a very large-leaved northern form of C. taguahyensis. Blan- 
chet has collected similar luxuriant forms in Bahia. 

Cordia revoluta Hooker fil. Trans. Linn. Soc. London 20: 199 
(1847); Riley, Kew Bull. 1925:225 (1925). Varronia revoluta 
Hooker f^l. ex Andersson, Kung. Svensk. Vet. Akad. Handl. 1853: 204 
(1855); Andersson, Freg. Eugenies Resa, Bot. 84 (1861). Lithocar- 
dium revoluta (Hook, f.) Kuntze, Rev. Gen. 2:977 (1891). Sebes- 
tena revoluta (Hook, f.) von Friesen, Bull. Soc. Bot. Geneve, ser. 2, 
24: 183(1933). Corrffa /meam Hooker fal., Trans. Linn. Soc. London 
20: 199 (1847), not DeCandolle (1845). Varronia linearis Hooker fil. 
ex Andersson, Kung. Svensk. Vet. Akad. Handl. 1853: 204 (1855) and 
Freg. Eugenies Resa, Bot. 84, tab. 11, fig. 4 (1861). Sebestena linearis 
(Hook, f.) von Friesen, Bull. Soc. Bot. Geneve ser. 2, 24: 182 (1933). 
Lithocardium Hookerianum Kuntze, Rev. Gen. 2: 976 (1891). Cordia 
Hookerianum (Kuntze) Gurke in Engler & Prantl, Nat. Pflanzenf. IV, 
Abt. 3a, p. 83 (1893). 

Galapagos Islands: Narborough : Stcivart 3177 ; Snodgrass & Heller 
327. Albemarle : Snodgrass & Heller 28, 155, 196, 272, 897; Stewart 3169, 
3170, 3172, 3173; Baur 213; Macrae (cotype of C. revoluta). James: 
Stewart 3175, 3176; Cheesman 388; Darwin (type of C. linearis). 
Charles: Baur 214; Schimpff 215; Darwin (type of C. revoluta). 

I have had the opportunity of examining the types of Cordia, from 
the Galapagos Islands, described by Hooker and by Andersson. The 
study of this critical material, supplementing a careful examination of 
the large general collections from the islands preserved at the Gray 
Herbarium, has established specific identities which necessitate changes 
in the names currently applied to the island species. The above cited 
species and the three following are the only endemic species of Cordia 
on the islands. All belong to the section Varronia. While it may be 
generally stated that they are most closely related to the species of 



1935] JOHNSTON, STUDIES IN BORAGINACEAE, XI 175 

western Peru and Ecuador, their immediate relationships on the con- 
tinent are quite obscure. The three following species are closely related 
to one another but probably not immediately related to the well marked 
C. revoluta. The following key will aid in distinguishing the four insular 
endemics: 

Corolla elongate, tubular, length of tube 3-4 times the breadth of 
the weakly developed limb; leaves linear; stems and upper 
surfaces of the leaves covered with stout short appressed 

falcate hairs; inflorescence capitate Cordia revoluta Hook f. 

Corolla coarsely funnel form, length of tube less than 2 times 
width of the conspicuous spreading limb ; leaves lanceolate ; 
stems and upper surfaces of leaves with erect or ascending 
(at times minute) hairs; inflorescence tending to elongate; 
flowers short-stipitate at maturity. 
Upper surface of leaves with abundant minute stout forked 

hairs Cordia Anderssoni Giirke 

Upper surface of leaves with stiff slender spreading simple 
hairs. 
Stems and lower surface of leaves bearing a mixture of 
minute appressed forked or stellate hairs and coarser 

stiff erect simple hairs Cordia Scouleri Hook. f. 

Stems and lower surface of leaves with only stiff erect 

simple hairs Cordia leucophlyctis Hook, f. 

Cordia Scouleri Hooker fil., Trans. Linn. Soc. London 20:200 
(1847). Varronia Scouleri Hooker fil. ex Andersson, Kungl. Svensk. 
Vet. Akad. Handl. 1853: 204 (1855) and Freg. Eugenics Resa, Bot. 83 
(1861). Lithocardium Scouleri (Hook, f.) Kuntze, Rev. Gen. 2: 977 
(1891). 

Galapagos Islands: Albemarle: Stewart 3162. James: Baur 209; 
Scolder (type). Indefatigable: Si'enson 7. 

This plant is particularly close to the two following. It appears to 
be rare. The few collections seen, other than the type, are all mis- 
determined as C. galapagensis or C. leucophlyctis. The mixed pubes- 
cence on the leaves and stems decisively separates it from those species. 

Cordia Anderssoni (Kuntze) Gurke in Engler & Prantl, Nat. Pflan- 
zenf. IV. Abt. 3a, p. 83 ( 1893). Lithocardium Anderssonii Kuntze, Rev. 
Gen. 2: 976 (1891). Varronia canescens Andersson, Kungl. Svensk. 
Vet. Akad. Handl. 1853: 203 (1855) and Freg. Eugenics Resa, Bot. 83, 
tab. 11, fig. 2 (ISbl), not Cordia canescens UBK. (1818). 

Galapagos Islands: Albemarle: Stewart 3195. Abingdon: Stewart 
3158. James : Stewart 3157. Duncan : Baur 215. Charles : Lee; Anders- 
son (type). Chatham: Stewart 3165, 3166; Baur 216; Andersson (det. 
V. leucophlyctis). 



176 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

Cordia leucophlyctis Hooker fil. Trans. Linn. Soc. London 20: 199 
(1847). Varronia leucophlyctis Hooker fil. ex Andersson, Kungl. 
Svensk. Vet. Akad. Hand). 1853: 203 (1855) and Freg. Eugenies Resa, 
Bot. 83 (1861). Lithocardium leucophlyctis (Hook, f.) Kuntze, Rev. 
Gen. 2: 977 (1891). Varronia scaberrima Andersson, Kungl. Svensk. 
Vet. Akad. Handl. 1853: 202 (1855) and Freg. Eugenies Resa, Bot. 82, 
tab. 11, fig. 3 (1861), not Cordia scaberrima HBK. (1818). Utho- 
cardium galapagosenum Kuntze, Rev. Gen. 2:976 (1891). Cordia 
galapagensis (? Kuntze) Gurke in Engler & Prantl, Nat. Pflanzenf. IV. 
Abt. 3a,p. 83 (1893). 

Galapagos Islands : Narborough ; Snodgrass & Heller 331, 342. Albe- 
marle : 5"Morf^ra;y5 & Heller 75, 136, 195, 291, 857, 881, 893; Stewart 3159; 
Baur 210, 212; Macrae; Darwin (type of C. leucophlyctis). Inde- 
fatigable: Baur 211; Andersson (type of Varronia scaberrima). Har- 
rington : Stewart 3164. Hood : Stewart 3168. 

The type of C. leucophlyctis and Varronia scaberrima are remarkably 
similar. The plant illustrated as V. leucophlyctis by Andersson, Freg. 
Eugenies Resa, Bot. tab. 11, fig. 1 (1861), appears to be Cordia Anders- 
soni from Chatham Island. 

Cordia setigera, sp. nov., dumosa; ramulis gracilibus strigulosis; 
foliis lanceolatis 3-6.5 cm. longis 12-24 mm. latis tenuibus, apice 
acutis, basi cuneatis, margine evidenter irregulariterque arguto-dentatis, 
dentibus apiculatis, faciebus pilis numerosis 0.5-1 mm. longis ascen- 
dentibus asperatis (basibus pilorum saepe pustulatis) subtus pallidi- 
oribus, nerviis 4-6-jugatis perinconspicue pauceque ramosis, petiolis 
1-5 mm. longis; pedunculis terminalibus gracillimis 1-7 cm. longis; 
inflorescentia congesto-capitatis 7-10 mm. diametro; corolla alba in- 
fundibuliformi 15-18 mm. longa, limbo ascendenti ca. 1 cm. diametro, 
lobis rotundis ca. 3 mm. longis; calycibus ad anthesim ca. 6 mm. longis 
sparse strigosis, tubo 2-2.5 mm. longo et crasso, lobis triangularibus ca. 
1-1.5 mm. longis, apice linearibus 2-?> mm. longis attenuatis; fructu 
ca. 5 mm. longo apice exserto. 

Brazil: near Fazenda de Bom Jardin, Rio Jequitinhonha, in north- 
eastern Minas Geraes, 1817, St. Hilaire B' 1478 (type, Paris). 

A very well marked species which keys out with Cordia grandiflora 
and C. paucidentata in my revision of the Brazilian species of Cordia, cf. 
Contr. Gray Herb. 92: 20 (1930). From both of these species it is 
distinguished by its smaller corollas, its thin leaves, its sparsely setulose 
herbage, and its very sparsely strigose calyces. It is evidently a very 
slender loosely branched bush and hence quite different in habit from 
the much more southerly ranging C. paucidentata. Cordia grandijiora 



1935] JOHNSTON, STUDIES IN BORAGINACEAE, XI 177 

has different pubescence, very much larger differently shaped corollas, 
and comes from the Amazon Valley. The closest relative of the pro- 
posed species is probably C. Neowediana DC, of the forests back of 
Ilheos, Bahia. That species has more finely serrate, more hairy leaves, 
larger corollas, much larger brown-hairy calyces, and only short tips on 
the calyx-lobes. It is a plant of the wet coastal forests, C. setigera is a 
plant of the dry catingas inland. 

Cordia Neowediana DeCandolle, Prodr. 9:498 (1845); Fresenius 
in Martius, Fl. Bras. 8^ : 23 (1857), as C. Neowidiana; Johnston, Contr. 
Gray Herb. 92 : 64 ( 1930) . Varronia macrocephala sensu Nees & Mar- 
tius, Nov. Acta Acad. Caes. Leop.-Carol. Nat. Cur. 11: 78 (1823). 
Lithocardium Neowiedianum (DC.) Kuntze, Rev. Gen. 2: 977 (1891). 

The type of this species is preserved at Brussels. Through the kind- 
ness of Prof. W. Robyns I have had the privilege of borrowing it for 
study. The species is a well marked one and is certainly worthy of 
recognition. The single collection upon which it is based was obtained 
in southern Bahia, in the country back of Ilheos. It is one of the species 
of the section Varronia having the flowers capitately congested. The 
large white corollas, in size and shape, are very suggestive of those 
found in the distantly related C. paucidentata. The stems, calyces and 
both leaf-surfaces are conspicuously bristly. There is no other kind 
of pubescence. The hairs are stiffish, spreading and mostly ca. 2 mm. 
long. Most of them spring from a small pustulate base. The hairs on 
the calyx are brown. The leaves are lanceolate, serrate and about 7 cm. 
long and 2 cm. broad. The calyx is ca. 1 cm. long, bristly, and sparsely 
glandular above the middle. The lobes are nearly 3 mm. long and 
triangular with a short subulate tip 1-1.5 mm. long. In my revision of 
the Brazilian species, 1. c. p. 20, C. Neowediana keys out with C. longi- 
jolia, C. poliophylla and C. leucocephala, which are rather closely re- 
lated to it. From these three species it can be quickly distinguished by 
its very bristly leaves and much larger bristly calyx. 

Cordia Braceliniae, sp. nov., fruticosa diffusa; caulibus ca. 3 dm. 
longis laxe ramosis strigosis; foliis elliptico-obovatis vel oblanceolatis 
2-3 cm. longis 10-17 mm. latis basim versus attenuatis, apice obtusis, 
margine crenatis, nervis 3-5-jugatis rectis ascendentibus vix ramosis 
supra impressis subtus prominentibus, facie laminae superiore sparse 
rigideque strigosis pilis pustulae insidentibus, facie inferiore pallidiori- 
bus strigosis vix pustulatis; pedunculis terminalibus gracilibus 2-3 cm. 
longis strigosis; inflorescentia capitata ca. 8 mm. diametro 20-25-flora; 
calycibus inflatis sparse strigosis ca. 3 mm. longis tubo pallido, lobis 



JOURNAL OF THE ARNOLD ARBORETUM 



I. longis breviter ca. 0.5 mm. longeque 
. longa; fructu irregulariter turbinate 



April 14, 1931, Ynez Mcxia 5617 (type, Gray Herb.; isotype, 
Arn. Arb.). 

A very distinct species perhaps most closely related to C. paucidentata 
of southern Brazil and adjacent regions. It is quickly separated from 
that species by its low branching habit, sparse strigose indument, and 
short-appendaged calyx-lobes. In my treatment of the Brazilian species 
of Cordia, Contr. Gray Herb. 92 : 20 ( 1930), it keys out with C. latijolia, 
C. poliophylla and C. leucocephala, though it does not seem closely 
related to any of them. From these species, however, it is readily dis- 
tinguished by its low spreading habit, small leaves lacking in secondary 
nervation, and different pubescence. The proposed species is strigose on 
the leaves, younger stems, peduncles and calyx. The hairs are abundant 
but do not cover the leaf-surfaces. They are stiff, straight and closely 
appressed. Those on the upper leaf-surface spring from small disks of 
dark mineralized cells. The type material has shrivelled corollas only. 
It has been distributed incorrectly determined as "C. truncata." 

I find it a pleasure to associate with this well marked species the 
name of Mrs. H. P. Bracelin, of California. Her effective handling and 
distribution of the extensive collections of Mrs. Mexia make it fitting 
that her name should be associated with them and that it should be 
remembered by the students of the Brazilian flora. 

Cordia campestris Warming, Kjoeb. Vidensk. Meddel. 1867:12, 
fig. 2 (1868). 

Brazil: Minas Geraes : Lagoa Santa, Pinhoes, in campis, Jan. 28, 1866, 
Wanning (Copenh., type) ; Lagoa Santa, in campis ad Cabejeiras da 
lagoa, March 8, 1864, Wanning (Copenh.) ; Formigas, St. H Hair e sine no. 
(Paris) ; indefinite, Clausscn 221 (G, K, Copenh., Stockh., Paris). Goyaz : 
Formosa, shrub, corolla white, Dec. 24, 1894, Glaziou 21781 (K, ED, 
Paris). 

Warming's species has been treated by me, Contr. Gray Herb. 92: 29 
(1930), as a synonym of C. multispicata Cham, and several of the above 
collections cited under that species. Cordia campestris is very distinct 
from C. multispicata, however, and probably most closely related to C. 
verbenacea DC. and C. chacoensis Chodat, particularly to the latter. 
Among the Brazilian spicate species of the Varronia section, C. multi- 
spicata is readily recognized by having the pedunculate spikes prevail- 



1935] JOHNSTON, STUDIES IN BORAGINACEAE, XI 179 

ingly lateral (and axillary) and the petiole of the subtending leaf evi- 
dently decurrent upon the peduncle, quite in the manner observable in 
C. buddleyoides Rusby, C. axillaris Johnston, and C. gtuizumaejolia 
(Desv.) R. & S. The specimens I have cited above, including the type 
of C. campestris, have terminal spikes and the petioles not decurrent on 
the peduncles. They are quickly separable from C. verbenacea by hav- 
ing the upper leaf-surface abundantly and evidently hairy. Cordia 
campestris is separated from C. chacoensis by its low habit of growth, 
and general coarseness of its parts. It appears to be a small (under 1 m. 
tall), sparsely branched shrub of the open country. The leaves are 
usually 3-5 cm. broad, and the spikes 5-10 cm. long. The flower buds 
are usually apiculate. It ranges in the campo of Minas Geraes and 
Goyaz. Its relative, C. chacoensis, ranges from southern-most Brazil 
into Paraguay and across northern Argentina. 

Cordia guazumaefolia (Desv.) Roemer & Schultes, Syst. 4:463 
(1819). Varronia guasumaejoUa Desvaux, Jour, de Bot. 1:276 
(1809). Lithocardium gmzumijolia (Desv.) Kuntze, Rev. Gen. 3-: 
206 (1898). Cordia axillaris var. gymnocarpa Johnston Contr. Gray 
Herb. 92: 35 (1930). 

In my treatment of the Brazilian species of Cordia, 1. c. p. 30, I cited 
C. guazumaefolia as a synonym of C. corymbosa (L.) Don. A recent 
study of Desvaux's type at Paris, however, has proved this to be quite 
incorrect. Among material from Jussieu's herbarium at Paris I have 
found specimens of this species determined by Desvaux. One of these 
is labeled, "Brasil, envoye de Lisbonne pour M. Vandelli 1790." The 
plant described by Desvaux is evidently that which I treated as C. 
axillaris var. gymnocarpa. This plant should bear the name C. guazu- 
maefolia. It may be added that while there is an evident relation be- 
tween C. axillaris and true C. guazumaefolia, I am now of the opinion 
that their differences warrant specific rather than mere varietal 
separation. 

Cordia insigrnis Chamisso, Linnaea 8: 122 (1833). Cordia Haen- 
keana Mez, Bot. Jahrb. 12: 560 (1890). 

An examination of the type of C. Haenkeana, at Munich, makes it 
evident that it is only a form of C. insignis. The collection is given as 
having been collected by Haenke in Peru. The accuracy of this data, 
however, I greatly doubt. Cordia insignis is known only east and south 
of the Amazon Basin, from eastern Brazil to eastern Bolivia, and is 
certainly not to be expected in Peru. 



180 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

Cordia laevior, sp. nov., arborescens, 6 m. alta; ramulis fusculis, 
juventate pilis brevibus adpressis vel ascendentibus vestitis mox gla- 
brescentibus; foliis homomorphis oblongo-lanceolatis 15-26 cm. longis 
4-10 cm. latis medium versus latioribus, basi acutis, apice longissime 
acuminatis, supra in costa et nervis primariis hirsutulis ceteris glabris, 
subtus pallidioribus, costa et nervis numerosis puberulentibus, nervis 
7-8-jugatis, nervis tertiaribus obscuris, petiolis 5-10 mm. longis; 
cymis in furcis ramulorum ortis laxe ramosis; calyce in alabastro obo- 
vato 4-5 mm. longo 2.5-3 mm. crasso pilis minutis abundantibus vestito 
obscure costato, apice rotundo, lobis 5 plus minusve irregularibus tri- 
angularibus; corolla alba, tubo ca. 4 mm. longo, lobis ca. 2 mm. longis 
et latis, filamentis ca. 3 mm. longis basim versus pilosis; stylo sparse 
piloso; supra medium ovarii evidenter pilosis; fructu ignoto. 

Peru : Pongo de Cainarachi. Rio Cainarachi, tributary of the Huallaga, 
dept. of San Martin, ca. alt. 230 m., tree 6 m. tall, fi. white, Sept.-Oct., 
1932, King 2756 (type. Am. Arb. ; isotype, Gray Herb.). 

This species is related to C. Sprucei Mez, of the Rio Negro and the 
Guianas, from which it differs in having smoother, more elongate, less 
hairy leaves and a more loosely branched inflorescence. The leaves are 
not roughened above by slightly prominent repeatedly branched vein- 
lets. The lower surface is much less hairy. The specimen was dis- 
tributed as C. Vlei Johnston, from which, like C. Sprucei, it differs in 
having a very hairy ovary, finer pubescence on the lower leaf-surfaces, 
and more papery, more hairy, less regularly and sharply lobed calyces. 

Cordia ripicola, sp. nov., arborescens 8-10 m. alta dichotome ramosa; 
ramulis sordidis pilis brevibus rigidulis scabridis; foliis subhomomorphis 
oblongis vel obovato-oblongis vel lanceolatis 8-14 cm. longis 3-7 cm. 
latis medio vel supra medium latioribus, apice acuminatis, basi acutis, 
supra sublucidis sparsissime breviterque strigosis, subtus minute 
rigideque hispidulis, nervis 6-8-jugis, nervis tertiaribus obscuris, petiolis 
2-6 mm. longis; cymis gracilibus 3-10 cm. crassis laxe ramosis; calyce 
strigoso in alabastro obovato ca. 4 mm. longo 2-i mm. crasso, intus 
supra medium strigoso, apice rotundo, ad anthesim in lobos 2-5 irregu- 
lares disrumpente; corolla alba 4-5 mm. longa, lobis ca. 2.5 mm. longis, 
filamentis ca. 3.5 mm. longis basim versus pilosis; ovario glabro vel 
summum ad apicem sparse pubescente ; fructu ignoto. 

Peru : Florida, Rio Putumayo, at mouth of Rio Zubineta, dept. Loreto, 
ca. 180 m. alt., "Chore-ey," forest along river, fl. white, tree 8-10 m. tall, 
May-June 1931, Khig 2262 (type, Arn. Arb.; isotype, Gray Herb.) and 
2277 (AA. GH). 

A species related to C. Sprucei Mez and C. laevior Johnston, from 



1935] JOHNSTON, STUDIES IN BORAGINACEAE, XI 181 

which it differs in having scattered appressed hairs on the upper face 
and abundant minute appressed ones on the lower face of the smaller, 
more oblong leaves. The calyx is more papery in texture and opens more 
irregularly. The style and ovary are sparsely hairy. The character of 
the calyx, the appressed hairs on the leaves and the hairiness of the 
pistil readily separate it from C. Ulei Johnston, the species under which 
the type has been distributed. Cordia Ulei comes from southwestern 
Brazil, at ca. lat. 11°S. The proposed species was collected nearly 
under the Equator. Cordia ucayaliensis, comb. nov. (C. Ulei var. 
ucayaliensis Johnston), readily distinguished by having the upper sur- 
face of the leaves strigose, comes from northeastern Peru. 

Saccellmm brasiliense, spec, nov., gracile; foliis lanceolatis 4-7.5 
cm. longis 18-28 mm. latis medium versus latioribus, margine obscure 
sinuatis vel supra medium sparse denticulatis, supra viridibus pilos 
graciles rectos valde adpressos gerentibus, subtus pallidis distincte seri- 
ceis pilos abundantissimos minutos valde adpressos gerentibus, apice 
basique acutis, nervis 8-10-jugatis, petiolis 3-6 mm. longis strigosis; 
ramulis gracilibus laxe ramosis brunescentibus juventate sordide pubes- 
centibus mox glabrescentibus, lenticellis numerosis orbiculatis pallidis 
punctatis; ramulis fertilibus 3-5 cm. longis ca. 5-foliatis; inflorescentia 
terminali paniculata 2-6 cm. longa folia vix superante; calycibus stri- 
gosis, dentibus laxe recurvatis. 

Brazil: Corumba, JMatto Grosso, Dec. 23, 1902, Malmc 2759 (type, 
Herb. Berol.) ; Corumba, Dec. 22, 1902, Robert 804 (BM, BD). 

The two collections cited are devoid of corollas and are in early fruit- 
ing condition only. A study of the immature calyx and ovary, however, 
leave no doubt as to the generic relations of this interesting plant. The 
species is evidently a relative of the Bolivian 5. Oliveri, but is readily 
separable by its small silky-strigose leaves and generally more compact 
habit of growth. Saccellium brasiliense has been reported from 
Corumba, doubtfully as 5. lanceolatum, by Moore, Jour. Bot. 45: 405 
(1907). Following I give the names of the known species of Saccellium 
and cite all the collections I have examined of these relatively rare 
species. The three known species may be separated as follows: 
Plant glabrous or practically so ; leaves 3-5 cm. broad, broadest 

at or slightly above the middle ^. Olivcrii Britt. 

Plant evidently pubescent ; leaves less than 3 cm. broad, broadest 
at or below the middle. 

Leaves broadest near the middle, beneath silky strigose. lus- 
trous, blade 4-7.5 cm. long; fertile branches 3-5 cm. long, 
bearing about 5 leaves ; old branches brown, with evident 
pale lenticels 5. brasiliense n. sp. 



JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

adest near base, beneath velvety or somewhat 
I, blade 5-12 cm. long; fertile branches 
10-20 cm. long, bearing about 10 leaves; old branches 
gray or only rarely brown, without evident lenticels .... 
.S". lanccolatum H. & B. 

Saccellium Oliverii Britton ex Rusby, Bull. Torr. Bot. CI. 26: 147 
(1899). 

This species is known only from the type-collection made by Rusby, 
no. 2535, in May 1886 at 600 m. alt at Guanai (or Huanay), Bolivia. 
The locality is in the department of La Paz at the confluence of the 
Rio Mapiri and Rio Tipuani at about lat. 15°30' S. and long. 68° W. 
in Amazonian Bolivia. Only fruiting specimens of the species are 
known. In its slender brownish branches, rather evident pale lenticels, 
and general leaf-outline, the species resembles S. brasiliense more than 
it does 5. lanceolatum. The leaves are glabrous except for a few short 
inconspicuous ascending hairs along the midrib and principal veins. A 
similar scanty inconspicuous indument is also found in the inflorescence. 

Saccellium lanceolatum Humboldt & Bonpland, PI. Aequin. 1 : 47, 
tab. 13 (1806); Humboldt, Bonpland & Kunth, Nov. Gen. 7:209 
(1825); Miers, Trans. Linn. Soc. London, Bot. 1: 25, tab. 6 (1875). 

In two widely separated areas, 1. Northern Peru in northern parts 
(prov. Jaen) of the Dept. Cajamarca, lat. 5°-6° S., in the Amazonian 
drainage; 2. mountains of southern Bolivia (prov. Chuquisaca and 
Tarija) southward along the mountains of northern Argentina to Tucu- 
man, ca. lat. 27° S. 

Peru : between Jaen and Bellavista, prov. Jaen, 600-700 m. alt., shrub 
or small tree, common, April 29, 1912, Weherbauer 6209a (BD) ; Valley 
of the Maranon between Bellavista and the mouth of the Rio Chinchipe, 
prov. Jaen, 500 m. alt., small tree 4 m. tall, flowers white; accrescent 
calyces yellow-green, May 1, 1912, IVeberbauer 6226 (G, BD) ; Valley of 
the Maranon at the mouth of the Rio Chinchipe, prov. Jaen, 400-500 m. 
alt., tree 6 m. tall, flowers white, accrescent calyces yellowish green, April 
30, 1912, Weberbaner 6217 (G, BD) ; near Rio Huancabamba, Bonpland 
(type, Paris; fragments, DC, Lindl., Gray). Bolivia: between Atajado 
and Parapiti, 700 m. alt., small tree, Dec. 1910, Hersog 1192 (BD) ; south 
of Rio Pilcomayo, prov. Tarija, Feb. 18, 1916, Steinbach 1776 (BD) ; 
Bolivia, Pampas, evergreen tree 4.5-6 m. tall, woods. May 1864, Pearce 
(BM). Argentina: Islota, Sierra Sta. Barbara, Jujuv, dry open place, 
tree 15-20 m. tall, July 5, 1901, Fries 260 (Munich) ; Sierra Sta. Barbara, 
Salta, Schuel 38 (G) ; Rio Blanco, dept. Oran, Salta, 650 m. alt., flowers 
yellowish, tree 10 m. tall, trunk 5 dm. thick, in high forest, Nov. 19, 1927, 
Venturi 5546 (AA, G, K, BM) ; Abra Grande, dept. Oran, March 1927, 
750 m. alt., tree 5 m. high, flowers vellowish, Venturi 6780 (AA) ; Rio 
Piedras. dept. Oran, Nov. 15, 1911, Rodriguez 85 (G) ; Campo Duran, 



1935] JOHNSTON, STUDIES IN BORAGINACEAE, XI 183 

dept. Oran, a tree common on higher slopes, "Guayabil," Jan. 28, 1930, 
Parodi 9269 (G) ; Tartagal, Salta, a tree, Feb. 1923, Hamnan (G) ; hills 
near the crossing of the Rio Juramente, Salta, tree or shrub up to 6 m. 
tall, Feb. 21, 1873, Hieronynms & Lorentz 295 (BD, Deles) ; Alemania, 
dept. Guachipas, Salta, 1100 m. alt., flowers white, tree 6 m. tall, in high 
forest, trunk 2 dm. thick, Dec. 22, 1929, Venhiri 10005 (G, K, BM) ; El 
Cadillal, dept. Burruyacu, Tucuman, Dec. 20, 1909, Lillo 9823 (Deles) ; 
Tucuman, dept. Capital, alt. 450 m., tree 10 m. tall, flowers white, Dec. 12, 
1907, Lillo 7234 (G) ; Estate of Professor Lillo, dept. Capital, Tucuman, 
460 m. alt., March 1925, Venturi 3816 (AA) ; Tucuman, Dec. 12, 1907, 
Stuckert 18375 (Deles) ; Tucuman, Feb. 10, 1910, Lillo (G). 

The distinctly lanceolate leaves and the leafy, elongate stiffish 
branches readily characterize this species. The range of the tree is 
peculiar for it occurs in two far-separated regions in Peru and Argen- 
tina. Though this behavior suggests that two species or that a species 
and a variety is involved, a careful comparison of copious material has 
failed to produce any differences that would justify the proposal of even 
a new variety. The Peruvian plant differs from that of Argentina only 
in its perhaps somewhat sparser and slightly more slender pubescence 
on the herbage and in its somewhat darker stems. 

In the Plantae Aequinoctiales 1: 47 (1806), the source of the origi- 
nal Humboldt & Bonpland collection of Saccellium is given as "ad rivos 
fluvii Guancabamba." Similar data are on the type-specimen at Paris. 
In the Nova Genera, 7: 208 (1825), the locality is given in more detail 
as follows: "inter Loxam et Tomependam Bracamorensium, ad ripas 
fluminis Guancabambae." The locality, Loja, of course, is in southern 
Ecuador. Tomependa is a ruined village near the junction of the Rio 
Chinchipe and the Rio Marafion. The Rio Huancabamba joins the 
Maraiion about 50 km. above Tomependa. In all probability the type 
was collected in or near the province Jaen, in the region of northern Peru 
in which it has been collected by Weberbauer, Bot. Jahrb. 50: suppl. 
p. 92 (1914). 

Coldenia conspicua, sp. nov., prostrata ut videtur annua; caulibus 
articulatis laxe ramosis 2-15 cm. longis, juventate dense graciliterque 
hispidulis et plus minusve glanduliferis; foliis aggregatis numerosis, 
lamina late lanceolata vel elliptica 5-13 mm. longa 2-5 mm. lata, subtus 
prominenter costata et nervosa (nervis 2-3-jugatis vix conspicuis) pilis 
gracilibus brevibus numerosis erectis asperata, supra pustulosa pilis 
robustioribus longioribus rigidioribus numerosis ascendentibus asperata, 
margine laxe revoluta Integra vel obscurissime sparsissimeque crenata; 
petiolis gracilibus 2-9 mm. longis glanduliferis pilis abundantibus longis 
gracilibus erectis setosis; calyce 5-partito, lobis gracilibus basim versus 



184 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

subinduratis et subnavicularibus praeterea linearibus hispidis glandu- 
liferis ad anthesin ca. 9 mm. longis fructiferis ad 15 mm. longis; corolla 
conspicua coerulea, tubo ca. 9 mm. longo 2.5-3 mm. crasso lobis calycis 
subaequilongo intus glaberrimo, limbo 10-12 mm. lato patenti, lobis 4-5 
mm. diametro, faucibus baud appendiculatis, filamentis 4-5 mm. longis 
glabris apicem versus tubi affixis ca. 2 mm. longe extrusis, antheris 
oblongis medio-affixis 1-1.4 mm. longis; stylo filiformi glabro 15 mm. 
longo 2 mm. profunde bilobato, stigmatibus 2 minutis obscure bilobu- 
latis; nuculis 4 globosis 1.5-2 mm. diametro dense minuteque tessellato- 
tuberculatis per carunculas 1 mm. longas et crassas in apice receptaculi 
basaliter affixis; receptaculo ad anthesin cylindrico, fructifero turbinato. 

Peru: sand flat near Mejia, Dept. Arequipa, 40 m. alt, flowers blue, 
Oct. 26, 1923, Guenther & Buchtien 155 (type, Inst. Bot. Hamburg) ; 
Mejia, July 21, 1923, Guenther & Buchtien 156 (Hamburg); Mollendo, 
Dept. Arequipa, Miss D. Stafford K60 (Kew). 

A very distinct and remarkable species belonging to the Chilean and 
southern Peruvian section Sphaerocarya, Johnston, Contr. Gray Herb. 
70: 57 ( 1924). The nutlets of the new species are quite similar to those 
of this section in size, shape and markings. From the previously de- 
scribed species of the section, however, C. conspicua differs in its ex- 
tremely large corollas, its protruding stamens and its remarkable nutlet- 
attachment. The corollas are at least twice the size of those of any other 
species of Coldenia. The nutlet-attachment is also unique in the genus. 
In the known species of the section Sphaerocarya the immature nutlets 
are attached laterally at the middle of the sides of an erect subcylindrical 
gynobase. This is distorted somewhat by the crowding of the growing 
nutlets and tends to become constricted medially. After the nutlets 
have fallen away it is consequently more or less spool-shaped. In the 
proposed species the immature nutlets are borne laterally, not about the 
middle, but about the summit of the subcylindrical gynobase. By 
growth and the consequent pressure of crowding, the nutlets at ma- 
turity come to be attached basally in the expanded summit of the now 
turbinate gynobase. What is most peculiar is that each nutlet has a well 
developed strophiolate basal plug which is immersed in the gynobasal 
tissue. At maturity the strophioles loosen from the gynobase and with 
their attached nutlets fall away leaving 4 deep more or less united sock- 
ets in the much broadened apex of the gynobase. The mature gyno- 
base, hence, becomes more or less cupulate. 

The species is known only from along the coast in southern Peru in 
the general region of the port of Mollendo. The type has been reported, 
Bruns, Mitt. Inst. Allgem. Bot. Hamburg 8: 67 (1929), as C. dicho- 



1935] JOHNSTON, STUDIES IN BORAGINACEAE, XI 185 

toma, but that species, of course, has small corollas and utterly different 
fruit-structures. The other species of the section Sphaerocarya are 
poorly understood. Since publishing, 1. c, on the South American 
species of Coldenia I have seen the types of Philippi's species. I have 
been unable to separate his C. litoralis, C. atacamensis and C. parvifiora, 
though from geographic considerations one would expect that the plant 
from the coastal region (C litoralis) would be distinct from that of the 
high Puno de Atacama (C. atacamensis and C. parvifiora). The type 
of C. parvifiora is quite distinct from the Peruvian plants of the Arequipa 
region, which I cited under that name in my synopsis of the South 
American species of Coldenia. The correct name for this species is C. 
elongata Rusby! Its elongate leaf-blades, woolly petioles and calyces, 
and usually evidently crenate leaf-margins serve to distinguish it from 
Philippi's species. Coldenia elongata is known only from middle alti- 
tudes east of the coastal deserts of southern Peru and northern-most 
Chile. In Peru only two species of the section Sphaerocarya are known. 
These are C. conspicua which grows along the coast and C. elongata 
which grows along the Cordilleras in the interior. 

Coldenia Nuttallii Hooker, Kew Jour. Bot. 3: 296 (1851); John- 
ston, Contr. Gray Herb. 75: 43 (1925). Coldenia decumbens Hauman, 
Apuntes Hist. Nat. Buenos Aires 1: 55 (1909) and Anal. Soc. Cient. 
Argentina 86: 301 (1918). 

This species so wide-spread in the intermontane area of the western 
United States has been known only from two small areas in the high 
Cordilleras of Argentina, in northwestern San Juan, Johnston, Physis 
9: 316 (1929), and in the Uspallata Pass region in Mendoza, Hauman, 
1. c. The plant was collected around 3000 m. alt. in San Juan and about 
2300 m. alt. in Mendoza. A third locality for the species in South 
America, one much further south and so, not surprisingly, at lower alti- 
tudes, may now be put on record. I have seen a collection of C. Nuttallii 
in the herbarium at Munich which was obtained by Erik Ammann (no. 
5) in Oct.-Nov. 1927, at 700 m. alt. near Cobunco, Neuquen, Argentina. 

Tournefortia brasniensis Poiret, Encyc. 5: 357 (1804); Johnston, 
Contr. Gray Herb. 92: 89 (1930). 

I have studied the type of this doubtful species in the Lamarck 
Herbarium at Paris. It represents a specimen of Vernonia scorpioides 
(Lam.) Pers., with the flowers just beginning to develop. It is remark- 
ably like, and probably a part of the collections by Commerson made at 
Rio Janeiro ("de I'ile aux chats") in July, 1767. Consequently it may 
be a part of the same material as the type of Conyza scorpioides Lamarck, 
Encyc. 2:88 (1790). 



186 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

Heliotropium transalpinum Vellozo, Fl. Flum. 6$ (1825) and 
Icones, 2: tab. 40 ( 1827). Heliotropium tiaridioides var. schizocarpum 
Johnston, Contr. Gray Herb. 81: 7 (1928), where other synonyms are 
cited. 

Vellozo in describing and illustrating his species gave no indication 
as to whether the carpels were dorsally sulcate or not. Suspecting that 
the carpels were sulcate, however, since only plants with such develop- 
ments were known about Rio Janeiro, I provisionally cited the name 
H. transalpinum among the synonyms of my H. tiaridioides var. schizo- 
carpum. Vellozo's name, unhappily, is several years older than H. tiari- 
dioides Cham., the species I then accepted. Subsequent study and con- 
sideration of much South American material of Heliotropium, not avail- 
able when my monograph was written, has left no reasonable doubt as 
to the identity of the plant described and illustrated by Vellozo. The 
scores of specimens examined from Sao Paulo, Rio Janeiro, Minas Geraes 
and northward in Brazil, uniformly have sulcate nutlets, and there seems 
every reason for believing that Velloza's plant had them also. I am 
accordingly taking up H. transalpinum as the correct appellation for 
the plant formerly treated by me as H. tiaridioides var. schizocarpum. 
The southern plant with non-sulcate nutlets, which I treated as H. 
tiaridioides var. genuina must have the new name I am publishing below. 
The type of H. transalpinum was collected in the state of Rio Janeiro 
near Boa Vista, ca. 9 km. up the Rio Parahyba from the town of Para- 
hyba do Sul and beyond the coastal mountains (whence the specific 
name) from the city of Rio Janeiro. 

Heliotropium transandinum var. tiaridioides (Cham.) comb. nov. 
Heliotropium tiaridioides Chamisso, Linnaea 4:453 (1829). Helio- 
tropium tiaridioides var. genuina Johnston, Contr. Gray Herb. 81 : 6 
(1928), where other synonyms are cited. 

Heliotropium angiospermum Murray, Prodr. Stirp. Gottingen 217 
(1770); Johnston, Contr. Gray Herb. 81: 10 (1928). Heliotropium 
humile Lamarck, Tab. Encyc. 1: 393 (1791). 

In my treatment of the South American species of Heliotropium, 
Contr. Gray Herb. 81 : 66 (1928), I cited H. humile Lam. as a doubtful 
synonym of H. jruticosum L. This I now find is incorrect. In the La- 
marck Herbarium at Paris there is only one specimen determined by 
Lamarck as H. humile, this bears a label in his script reading: "heliotr. 
humile lam. illustr," The specimen is small but represents good H. 
angiospermum. The original description of Lamarck's species reads: 
"1757 HELIOTROPIUM humile. H. foliis (.vato-lanceolatis villosis; 



1935] JOHNSTON, STUDIES IN BORAGINACEAE, XI 187 

spicis solitariis lateralibus. Ex ins. Carib. Annuum. H. Diet. no. 6 Quoad 
descr." The reference is apparently to Lamarck's earlier account of 
Heliotropium in vol. 3 of the Encyclopedie, pp. 92-95 (1789), but no 
mention of H. humile is to be found there. Species no. 6 in the work is 
H. jruticosum, described as having linear-lanceolate leaves. Poiret, 
Encyc. Suppl. 3: 25 (1813), was evidently puzzled by Lamarck's de- 
scription of H. humile. He mentions that species under H. ternatum but 
suggests that it might be H. jruticosum. I am content, however, to place 
H. humile among the synonyms of H. angiospermum, for the named 
specimen in Lamarck's herbarium seems authentic and agrees with the 
few words in the original description. 

Lasiarrhenum pinetonim, sp. nov., herba; caulibus erectis simplici- 
bus 10-15 cm. altis gracilibus strigosis foliosis; foliis lineari-subulatis 
1-3 cm. longis 1-1.5 mm. latis sessilibus medio-costatis sed vix nervatis 
apicem versus caulis gradatim reductis margine valde revolutis supra 
sparse strigosis; floribus cymas terminalis 3-7-floris lineari-bracteatas 
aggregatis; pedicellis 2-3 mm. longis strictis strigosis; calycibus 5-lobatis 
ca. 4 mm. longis, lobis lineari-lanceolatis strigosis; corolla flava ca. 10 
mm. longa extus strigosa, tubo ca. 4 mm. longo ca. 1.5 mm. crasso in 
fauces 3.5-4 mm. longas ca. 3 mm. crassas abrupte transmutato intus 
glaberrimo, lobis erectis oblongis 2.5 mm. longis 2 mm. latis apice 
rotundis; antheris glaberrimis 2 mm. longis oblongis basi sagittatis erec- 
tis (vix versatilibus) sub medium affixis; filamentis 4 mm. supra basim 
corollae affixis inclusis 1-1.3 mm. longis late alatis (in ambitu obovatis) 
apicem versus ca. 0.7 mm. latis; stylo 12 mm. longo filiformi longe (ca. 
5 mm.) exsertis; fructu ignoto. 

Mexico : growing in the mountains in pine-forest, very rare, September, 
Ghicsbrcght 311 (type, Paris). 

This is a remarkable species which is placed in Lasiarrhenum chiefly 
because of its broadly winged filaments. From L. strigosum, formerly 
the only known member of its genus, it differs in its very small size, its 
uninerved leaves, its glabrous anthers and its precociously long-exserted 
style. The rounded corolla-lobes and the expanded filaments separate 
L. pinetorum from the genus Onosmodium, while the long-exserted style, 
the erect corolla-lobes, the obovate filament and the sagittate anthers 
distinguish it from Lithospermum. No locality is given for this inter- 
esting plant. Ghiesbreght, however, collected chiefly in southern 
Mexico and mostly in the state of Oaxaca. 



188 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

2-5 dm. altis e radice crasso dense multicepite rumpentibus strigosis 
vel basim versus breviter hispidis; foliis strictis firmis costatis sed vix 
nervatis vel rarissime perinconspicue sparseque nervatis, inferioribus 
oblongo-ellipticis, aliter lanceolatis, sessilibus, apicem versus caulis 
gradatim reductis 1-4 cm. longis 3-8 mm. latis, apice acutis, supra minute 
strigosis et pustulatis, subtus in margine et costa strigosis sed ceterum 
glabris; inflorescentia bracteata scorpioidea terminali solitari vel gemi- 
nata vel ternata 3-10 cm. longa; calyce ad anthesin ca. 6 mm. longo, 
lobis inaequalibus cuneatis, pedicellis 1-3 mm. longis strigosis; corolla 
subcylindrica 15-19 mm. longa ca. 3 mm. crasso ut videtur f^avescente 
intus glaberrima extus adpresse pubescente, lobis minutis ascendentibus 
suborbicularibus ca. 1 mm. diametro, faucibus inconspicue plicato- 
appendiculatis saepe plus minusve constrictis; staminibus 2 mm. sub 
apice tubi coroUae affixis, filamentis ca. 1 mm. longis, antheris oblongis 
ca. 2 mm. longis inclusis; stylo filiformi ad anthesin 1-3 mm. longe 
extrusis; fructu ignoto. 

slope of Sierra Tronconal between Canon de los Charcos and Canon de 
San Miguel, Sierra Madre Oriental, ca. 25 km. s. w. of Galeana, Nuevo 
Leon, 1800-2700 m. alt., June 4, 1934, C. H. & M. T. Mueller 739 (type, 
Gray Herb.). 

A very distinct species of uncertain affinities. Its subtubular corolla, 
frequently with a narrowly constricted ring about the throat, and its 
extremely small round ascending lobes, separate it from L. strictum, the 
only species I am inclined to believe which possibly may be a close 
relative of it. 

Macromeria leontis, sp. nov., perennis erecta ca. 5 dm. alta e radice 
crasso profunde oriens; caulibus subsimplicibus pilis brevibus gracilibus 
erectis vel subretrorsis dense vestitis; foliis lanceolatis medium versus 
caulis grandioribus 4-10 cm. longis 1-2 cm. latis utroque acutis sessilibus 
evidenter nervatis, subtus pilis gracilibus brevibus erectis abundantibus 
vestitis vix pustulatis, supra viridis pustulatis et breviter hispidis; in- 
florescentia terminali evidenter bracteata; calyce 1.5-2 cm. longo, lobis 
linearibus, pedicello 1-5 mm. longo; corolla 5-7 cm. longo intus glaber- 
rimo extus breviter pubescenti, tubo 2-3 cm. longo 1.5-2 mm. crasso 
supra in fauces 2 cm. longos 7-8 mm. latos gradatim ampliato, lobis 
triangularibus ca. 9 mm. longis et 6 mm. latis non rariter plus minusve 
recurvatis; antheris elongatis ca. 3.5 mm. longis; filamentis ca. 4 mm. 
infra apicem faucium corollae affixis 10-15 mm. longe exsertis; stylo 
filiformi tarde exsertis; fructu ignoto. 



1935] JOHNSTON, STUDIES IN BORAGINACEAE, XI 189 

Madre Oriental, ca. 25 km. s. w. of Galeana, Nuevo Leon, ca. 2400 m. 
alt., June 5, 1934, C. H. & M. T. Mueller 754 (type, Gray Herb.) ; scat- 
tered in dense pine and oak woods along the descent into Alamar, Sierra 
Madre Oriental, May 29, 1934, C. H. & M. T. Mueller 594 (G). 

Probably a relative of M. Pringlei, but differing in having a fine slen- 
der spreading indument throughout. In M. Pringlei the more rigid, 
somewhat longer sparser hairs are closely appressed and the upper leaf- 
surfaces are a much clearer green than in M. leontis. The latter species 
has leaves noticeably grayer and duller in color. 

Macromeria barbigera, sp. nov., perennis, setosa, robusta; caulibus 
erectis 5-8 dm. altis saepe simplicibus; foliis lanceolatis vel ovatis evi- 
denter nervatis, inferioribus parvis vix persistentibus, ceteris latioribus 
3-5 cm. latis 5-11 cm. longis subsessilibus basi plus minusve rotundis, 
superioribus elongatioribus et minoribus; floribus terminalibus in cymas 
racemosas bracteatas aggregatis; bracteis foliaceis 2-7 cm. longis 1-4 
cm. latis; pedicellis ca. 5 mm. longis; calyce ad anthesin ca. 18 mm. 
longo, lobis inaequalibus subulato-linearibus erectis; corolla ut videtur 
flavescenti intus glaberrima 5-6 cm. longa recta vel plus minusve curvata, 
tubo 1.5-2 cm. longo 1.5-2 mm. crasso lobis calycis pauUo longiore, 
faucibus e tubo abrupte ampliatis ca. 2 cm. longis 5-6 mm. crassis cylin- 
draceis in alabastro paullo asymmetricis, limbo abrupte dilatato 12-15 
mm. diametro, lobis 5-6 mm. longis acutis ascendentibus apicem versus 
recurvatis; filamentis in faucibus ca. 8 mm. infra sinibus loborum affixis 
inaequalibus 12-15 mm. longis glabris filiformibus exsertis; antheris 
oblongis medio-affixis ; stylo filiformi breviter tardeque extruso; stigmato 
minimo bilobulato ; fructu ignoto. 

Mexico: common in dense oak wood beyond the pine and fir belt, north 
slope of Sierra Tronconal between Canon de San Miguel and Canon 
de los Charcos, 1800-2700 m. alt., Sierra Madre Oriental about 25 km. 
s. w. of Galeana, Nuevo Leon, June 4, 1934, C. H. & M. T. Mueller 741 
(type. Gray Herb.). 

Related to M. Thurberi but quickly separable by its more robust habit, 
larger broader leaves and very different pubescence. The foliage of M. 
Thurberi is copiously and finely strigose with an admixture of coarse 
more or less spreading hairs. In the proposed species the strigosity is 
lacking and the spreading hairs much longer and very conspicuous. The 
corollas of M. Thurberi have a much more abundant and paler indu- 
ment than do those of M. barbigera. The range of the new species is 
to the southeast of the most easterly station of its relative. 

Evidently to be identified with M. barbigera are collections made by 
Mueller in 1933. These specimens have been kindly sent to me from 



190 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

the Field Museum by Mr. P. C. Standley. One of these collections, no. 
174 from the "Trail to Puerto," Nuevo Leon, has leaves becoming 17 
cm. long and 7 cm. broad. Its flowers are immature. The second col- 
lection, no. 173 from Diente Canyon, 21 km. south of Monterey, is evi- 
dently from a very mature plant and consists of the elongated 
inflorescence showing mature bracts and the old pedicels and calyces. 

Among his collections of 1934 Mueller obtained one which may also 
represent a form of M. barbigera. This specimen, no. 830, was collected 
on Sierra Infernillo, about 25 km. s. e. of Galeana, Nuevo Leon, where 
it was common over small areas just below the crest, 2 700-3000 m. alt. 
In leaf-outline and in general habit the plant suggests M. Thurberi, but 
differs in its lack of strigosity and in its very much less hairy flowers. 
The corollas differ from those of M. barbigera. They are somewhat 
smaller. The tube is gradually expanded towards the lobes and not 
abruptly expanded into a well developed cylindrical throat as I have 
indicated in my formal description above. In addition the corolla is 
slightly less hairy and the lobes not so acute. The plant is evidently 
related to M. barbigera and chiefly because of geographical considera- 
tions I am tentatively, at least, referring it to that species as a possible 
ecological form. 

Havilandia opaca, sp. nov., procumbens; caulibus foliosis abundan- 
ter ascendenterque ramosis 1-1.5 mm. crassis in nodis radiculas graciles 
gerentibus pilis brevibus rigidis appressis dense vestitis, internodiis 3-10 
mm. longis; foliis iirmis subcoriaceis costatis sed enervatis numerosis, 
apice rotundis vel obtusis, supra glaberrimis sparsissime pustulatis in 
costa sulcatis, subtus supra medium pustulatis in costa prominente 
strigosis ceteris glabris vel sparsissime strigosis, margine strigoso-ciliatis 
vel basim versus sparse ciliatis; foliis ramorum fertilium ellipticis 4-10 
mm. longis 3-5 mm. latis, basi rotundis et oblique 1-2 mm. lateque 
sessilibus; foliis ramorum sterilium plus minusve oblanceolatis 8-12 mm. 
longis pauUo sub apicem basim versus in petiolum 1 mm. latum ca. 2 mm. 
longum gradatim attenuatis; floribus solitariis numerosis axillaribus; 
corolla alba 4 mm. diametro, tubo ca. 1.2 mm. longo 1 mm. crasso intus 
glaberrimo, limbo patenti, lobis suborbicularibus ca. 1.5 mm. diametro, 
appendiculis faucium 5 intrusis trapeziformibus ; antheris oblongis 
inclusis ca. 0.4 mm. longis, filamentis perbrevibus pauUo supra medium 
tubi affixis; calyci ad anthesin 2 mm. longo, lobis 5 ciliatis latis, pedi- 
cello 0.5-1 mm. longo; nuculis 4 erectis angulate ovoideis 1 mm. longis 
opacis dense minutissimeque papillatis, dorso convexis, ventre angulatis, 
imam ad basim anguli ventralis ad gynobasim planum affixis. 



1935] JOHNSTON, STUDIES IN BORAGINACEAE, XI 191 

British New Guinea : common in open grassland, Murray Pass, Whar- 
ton Range, 2840 m. alt., prostrate herb forming masses 3 dm. broad or 
more, flowers white, June 12, 1933, Brass 4178 (type, Gray Herb.; iso- 
TYI'E, NY). 

A species evidently related to H. papuana Hemsl., from which it 
differs in its stout somewhat ovate, gray, dull, minutely papillate, rather 
than elongate, somewhat lance-lunate, black, lustrous, smooth nutlets. 
The margins of the leaves in H. papuana are evidently ciliate. In H. 
opaca the marginal hairs of the leaves, similar in size, number and posi- 
tion, are not spreading, but antrorsely appressed along the leaf-margin. 
The habit of growth in H. papuana is quite similar to that of H. opaca. 

Havilandia robusta, sp. nov., procumbens; caulibus elongatis sparse 
ramosis; foliis coriaceis oblanceolatis 2-4.5 cm. longis 5-9 mm. latis 
paullo sub apicem basim versus in petiolum vaginatum gradatim attenu- 
atis, apice rotundis vel subemarginatis, margine sparsissime strigosis, 
supra sparse strigosis, subtus glaberrimis vix nervosis, costa prominente 
sparsissime strigosa; floribus axillaribus; calycibus ad anthesin ca. 4 
mm. longis, lobis lanceolatis margine sparsissime strigosis, pedicellis 
5-7 mm. longis; calycibus maturitate ca. 6 mm. longis pedicellis 8-12 
mm. longis; corolla 8-10 mm. diametro; nuculis 4 angulato-ovoideis 
opacis ca. 2 mm. longis dense minutissimeque papillatis, dorse convexis, 

British New Guinea: common about forest borders, Mt. Albert Ed- 
ward, 3680 m. alt., June 1933, Brass 5681 (type, N. Y. Bot. Gard.). 

Evidently related to H. opaca, also of southeastern New Guinea, from 
which it differs only in being much larger in all its parts, and in having 
well developed pedicels and more elongate leaves. The upper surface 
of the leaves is lustrous and distinctly strigose. 

Havilandia papuana Hemsley, Kew Bull. 1899: 107 (1899). 

British New Guinea: thickly massed on shallow soil over rock in 
grasslands, Mt. Albert Edward, 3680 m. alt., flowers white with yellow 
throat, June 18, 1933, Brass 4245 (G, NY). 

This species was briefly, though adequately described by Hemsley 
from material obtained on Mt. Scratchley, 3660 m. alt., and in the 
Wharton Range, 3330 m. alt. It is known only from the high mountains 
of eastern British Now Guinea. 

The genus Havilandia is confined to high altitudes and consists of the 
three above enumerated species from British New Guinea, and H. 
borneensis Stapf from Mt. Kinabalu in British North Borneo. It is 
possible, in addition, that Lithospermum minutum Wernh., described 
from the Mt. Carstensz region in Dutch New Guinea, may also belong 



192 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

to Havilandia. Unfortunately the type and only known collection of 
this puzzling species is so scanty and inadequate that it must remain 
an obscure, troublesome element in the flora of New Guinea until some- 
one recollects it. The type consists of two minuscule snips in flower 
only, a ridiculously inadequate basis for the proposal of any species of 
Boraginaceae and certainly for one whose acquaintance with the genera 
of that family may be judged by his selection of the genus under which 
he essayed to publish the imperfect specimen from Dutch New Guinea. 

Plagiobothrys Scouleri (H. & A.) Johnston, Contr. Gray Herb. 
68:75 (1923) and Contr. Arnold Arb. 3:51 (1932). Myosotis 
Scouleri Hooker & Arnott, Bot. Beechey Voy. 370 (1840). Allocarya 
media Piper, Contr. U. S. Nat. Herb. 22: 107 (1920). Plagiobothrys 
medius (Piper) Johnston, Contr. Arnold Arb. 3: 58 (1932). Allocarya 
divaricata Piper, Contr. U. S. Nat. Herb. 22: 107 (1920). 

The original and only mention of Myosotis Scouleri in the writings of 
Hooker & Arnott appears in the Botany of Capt. Beechey's Voyage in a 
note on a collection of Plagiobothrys Ckorisianus from California. The 
note is as follows: "The flowers here are on pretty long pedicels, while 
the Columbia plant has them shortly pedicellate; the latter presents, 
besides, a different aspect, and may be called M. Scouleri; it appears 
very closely allied to M. califomica, Fisch. et Meyer, but the corolla is 
longer than the calyx." Gray, who apparently never studied the type of 
M. Scouleri, applied the name to an erect-growing plant with geminate 
spikes of conspicuous corollas, which is widely distributed in western 
Oregon and Washington, and all subsequent writers have followed him 
in that identification. A study of the type, however, shows this usage 
to be quite incorrect. 

The specimens evidently the type of M. Scouleri are to be found on 
a mixed sheet, formerly in the Hooker Herbarium, now at Kew. This 
sheet bears three different collections: ( 1 ) the specimen of P. Chorisianus 
mentioned in the Botany of Beechey's Voyage, (2) specimens of P. 
scopulorum (?) or P. cognatus (?) collected by Nuttall, and (3) three 
plants labeled: "N. W. Coast, Dr. Scouler." The latter evidently 
constitute the type of Myosotis Scouleri H. & A. Duplicates of this 
Scouler collection are to be found on a sheet from Bentham's herbarium, 
at Kew, labeled: ''Am. bor. occ. Scouleri, 1828," and in the herbarium at 
Edinburgh labeled: "Columbia, Scouler, 1827, (932)." These collec- 
tions appear to represent a form of the plant I have treated in my mono- 
graph as Plagiobothrys medius (Piper) Johnston. They have the rufous 
calyx-lobes, evident corollas, and the general habit of that species. 



1935] JOHNSTON, STUDIES IN BORAGINACEAE, XI 193 

Scouler is known to have collected about the mouth of the Columbia 
and at many small ports along the coast of Washington and Vancouver 
Island. Plagiobothrys medius is the common species near the coast in 
northwestern Washington and on Vancouver Island, and there is every 
reason that Scouler should have encountered it. Though the nutlets of 
Scouler's collection show certain peculiarities not matched in the avail- 
able material of P. nuedius, I believe that they can be accommodated in 
that concept. The nutlets of the type of M. Scouleri have the rather 
bony pericarp common in P. medius, but the ridge attending the lateral 
scar is very closely appressed to the latter and encloses an areole 
(entirely filled by the scar) scarcely, if at all, broader than long. The 
nutlets of P. medius are, however, very variable and I believe the nutlet- 
variations of M. Scouleri can be admitted without destroying the natural- 
ness of the concept. 

Plagiobothrys hirtus ( Greene ) , comb. nov. Allocarya hirta Greene, 
Pittonia 1:161 (1888). Allocarya Scouleri var. hirta (Greene) Nelson 
& Macbride, Bot. Gaz. 61:36 (1916). Plagiobothrys Scouleri var. 
hirtus (Greene) Johnston, Contr. Arnold Arb. 3: 52 (1932). Allocarya 
calycosa Piper, Contr. U. S. Nat. Herb. 22: 101 (1920). 

I have indicated above that the type of Myosotis Scouleri H. & A. has 
been misinterpreted. The earliest correct name for the plant that has 
been called Krynitzkia, Allocarya and Plagiobothrys Scouleri is Allo- 
carya hirta Greene. It is, however, strictly applied only to a local plant 
of the Umpqua Valley, Oregon, which has evidently spreading rather 
than appressed pubescence. The common form of this species must 
bear the following name: 

Plagiobothrys hirtus var. figuratus (Piper), comb. nov. Allo- 
carya figurata Piper, Contr. U. S. Nat. Herb. 22: 101 (1920). 

This strigose form ranging from Oregon to Vancouver Islands is 

Plagiobothrys hirtus var. corallicarpus (Piper), comb. nov. Allo- 
carya corallicarpa Piper, Proc. Biol. Soc. Wash. 37 : 93 ( 1924) . Plagio- 
bothrys Scouleri var. corallicarpus (Piper) Johnston, Contr. Arnold Arb. 
3: 52 (1932). 

A local form of southern Oregon characterized by its deeply alveolate 

Plagiobothrys calandrinioides (Phil.) Johnston, Contr. Gray Herb. 
78: 91 (1927). Allocarya alternifolia Brand in Fedde, Repert. 26: 169 
(1929). 



H JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

The type of Brand's species has been examined. The lowermost leaves 
:count for the very misleading spe- 
i Patagonian F. calandrinioides . 

Thaumatocaryon dasyanthum var. Sellowianum (Cham.), comb, 
nov. ^«c/rM5fl Se//ow/awa Chamisso, Linnaea 8: 115 (1833). Moritzia 
Sellowiana (Cham.) Fresenius in Martius, Fl. Bras. 8' : 63 (1857). 
Thaumatocaryon Sellowianum (Cham.) Johnston, Contr. Gray Herb. 
70 ; 13 (1924) and 78 : 16 ( 192 7 ) . Moritzia dasyantha var. Sdlounana 
(Cham.) Brand in Fedde, Repert. 27: 148 (1929). 

This plant differs from typical T. dasyantka only in its smaller corollas 
and appressed pubescence. Difficulty with connecting forms has con- 
vinced me that Brand might best be followed in treating Sellowianum as 
a mere variety. A collection of this variety from the state of Rio Janeiro, 
by Glaziou (no. 8731), supplies the basis for Glaziou's astonishing 
report of Cyphomattia lanata in Brazil, Bull. Soc. Bot. France 57: Mem. 
3: 480 (1910). I have examined the specimen at Paris so determined 
by Glaziou. 

Hackelia patens (Nutt.), comb. nov. Rochelia patens Nuttall, 
Jour. Acad. Phila. 7:44 (1834). Lappula cocrulescens Rydberg, 
Mem. X. Y. Bot. Gard. 1 : 328 ( 1900). Lappula subdecumbens coeru- 
lescens (Rydb.) Garrett, Fl. Wasatch Reg. 78 (1911). Hackelia diffusa 
var. caerulescens (Rydb.) Johnston, Contr. Gray Herb. 68: 48 (1923). 
Hackelia caerulescens (Rydb.) Brand, Pflanzenr. [Heft 97] IV. 252-: 
130 (1931). Hackelia Nelsonii Brand in Fedde, Repert. 26: 170 
(1929). Lappula decumbens Nels. ex Brand, Pflanzenr. [Heft 97| 
IV. 252^: 126 (1931), lapsus calami. 

I have examined Nuttall 's type of Rochelia patens at the British 
Museum. The specimen was collected "near the Flat-Head River" on 
June 8, 1833, by N. B. Wyeth. The specimen is a good one and is evi- 
dently conspecific with Lappula caerulescens, a species also based upon 
material from western Montana. The species is known from western 
Montana and Wyoming and westward into Idaho, northern Utah and 
northern Nevada. 

Hackelia grisea (Woot. & Standi.), comb. nov. Lappula grisca 
Wooton & Standley, Contr. U. S. Nat. Herb. 16: 164 (1913). 

A readily recognizable species of New Mexico and adjacent Texas. 
Its relatively small corollas, with ascending lobes, quickly distinguish 
It among the west American annual and biennial species of this genus. 

Lappula echinata Gilibert, Fl. Lituanica, 1:25 (1781). Cryp- 



1935] JOHNSTON, STUDIES IN BORAGINACEAE, XI 195 

tantha Lappula Brand in Fedde, Repert. 24: 56 (1928) and Pflanzenr. 
[Heft 97] IV. 252^: 147 (1931). 

In the Pflanzenreich Brand placed his Cryptantha Lappula among the 
synonyms of Lappula Redowskii (Hornem.) Greene. I believe, however, 
that the species belongs under L. echinata Gilib. The evident corollas 
and the gross aspect of the type are of that species. A microscopic 
study of the (immature) nutlets of C. Lappula seems to show a double 
row of lateral prickles. Finally the type is given as from Concepcion, 
Chile, a locality at which L. Redowskii is certainly not to be expected to 
grow 'naturally, though a busy port at which an aggressive weed, such 
as L. echinata, might be introduced without any cause for surprise. 

Lappula echinata is generally accepted as introduced into North 
America. This seems probable, though it is to be noted that the plant 
was collected in the New World at a very early date. A specimen in 
the DuBois collection at Oxford is labeled "brot from Maryland by Mr. 
Wm. Vernon, 1698." Among Michaux's collections at Paris there is 
one of this species labeled as "Dans ville de Montreal, 1792." The 
Smith collections in London contain a specimen labeled: "North America, 
1817, F. Booth." In the British Museum there is a collection made by 
Douglas, during 1826, "In the valleys of the Rocky Mts.," most likely 
in northeastern Washington. It seems to have been again collected in 
the latter region only within the past ten years, though it has been well 
known in the southern parts of western Canada for at least a generation. 
There are reasons to believe that the railroads may have much aided in 
the distribution north of the International Boundary. The plant has 
exhibited an evident, progressive increase and migration westward across 
the more northern of the western United States. It is now rapidly 
increasing in eastern Washington where it gives every evidence of being 
a recent immigrant. 

Cryptantha circumscissa (H. & A.) Johnston, Contr. Gray Herb. 
68: 55 (1923). 

A few years ago, 1. c. 81: 75 (1928), I reported this characteristic 
plant of western United States from near Zapala, Neuquen, Argentina. 
A second station in Neuquen may now be recorded. At Munich I have 
seen specimens labeled as collected by Erik Ammann (no. 7) at Cerro 
Mesa between Sept. and Nov. 1927. The new station is nearly 90 km. 
southeast of Zapala. 

Cryptantha clandestina (Trev.), comb. nov. Lithospermum clan- 
destinum Treviranus, Del. sem. a 1832 in hort. Bonnensi collect, p. 2 
(1832-3). Cryptantha glomerata Lehmann, Del. Sem. Hamb. 1832: 4 
(1832), nomen nudum; Fischer & Meyer, Ind. Sem. Hort. Petrop. 2: 8 



196 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

and 35 (1836); Johnston, Contr. Gray Herb. 78: 58 (1927). Cryp- 
tantha microcarpa Fischer & Meyer, Ind. Sem. Hort. Petrop. 2:8 and 
35 (1836). 

A study of the original description of Lithospermum clandestinum, 
and of old garden material representing it, has made it clear that it is 
that well known cleistogamic species of Chile, the two forms of which 
have passed as Cryptantha glomerata and C. microcarpa. Fischer & 
Meyer, when describing C. microcarpa, in fact, actually cited L. clan- 
destinum as a synonym. In the Bonn seed-list for 1832, published in 
Dec. 1832 or Jan. 1833, the name Lithospermum clandestinum appears 
in the alphabetic list on the second of the pages of that quarto catalogue. 
A reference leads to a footnote which reads as follows: "Diffusum his- 
pidum; fol. lanceolatis amplexicaulibus; calycibus subsessilibus ven- 
tricosis coroUam excedentibus; seminn. granulatis. Annuum. Corolla 
alba, tubo ventricoso, limbo conniventi. Semina duo plerumque 
abortiunt. T[reviranus] ." The name, L. clandestinum, appears again 
in the Bonn list for 1833, but not in those for 1834 or 1835. 

Cryptantha glomerata Lehm. is the type-species of Cryptantha. Re- 
cently I had the privilege of consulting the extensive collections of old 
seed-catalogues at Berlin and Geneva. I now find it possible to record 
several important references in the history of that genus and species 
which were either unknown or unavailable to me at the time of my work 
on the group. The first mention of Cryptantha glomerata Lehm. and 
of the generic name appears in Lehmann's seed-list of the Hamburg 
Garden for the year 1832. The binomial appears as a mere name on 
page 4, thus: "Cryptantha glomerata Lehm." No description or ex- 
planation of the name is given! The list is dated 1832 and was prob- 
ably published, as was customary with such lists, around the close of the 
year. No mention of the binomial is found in the Hamburg lists for 
1830, 1831 or for 1833 or 1834. In 1835, p. 4, again without descrip- 
tion, appears: "Cryptantha glomerata Lehm. (Del. Sem. 1832)." In 
1836, p. 4, the following two names appear bare of description: "Cryp- 
tantha glomerata Lehm." and "Cryptantha microcarpa F. & M." These 
are repeated in the list for 1837, p. 4. In the list for 1838, p. 4, there 
is merely the name, "Cryptantha microcarpa F. & M." Fischer & Meyer, 
in their St. Petersburg seed-list for 1835, supplied the first descriptions 
of Cryptantha glomerata Lehm. and C. microcarpa F. & M. This Rus- 
sian list bears a censor's date, Dec. 25, 1835, the equivalent of Jan. 5, 
1836 of our present calendar. There is no mention of Cryptantha in 
the St. Petersburg list for 1834 ! Fischer & Meyer, when publishing and 
describing "C. glomerata Lehm." in their list for 1835, attribute the 



1935] JOHNSTON, STUDIES IN BORAGINACEAE, XI 197 

name to "Bernhardi in litt." A study of Bernhardi's seed-lists, Sel. sem. 
hort. Erfurt., shows that the name "C. glomerata Lehm." appears as a 
mere binomial in those for 1833 (Jan. 18, 1834), 1834 (Feb. 24, 1835) 
and 1835. There is no mention of Cryptantha in the Erfurt list for 1832 ! 
From the facts I have given it becomes evident that Cryptantha 
glomerata was in cultivation at Bonn and Hamburg in 1832. Treviranus 
immediately described the Bonn cultures as Lithospermum clandes- 
tinum. Lehmann applied to his Hamburg cultures the name Cryptantha 
glomerata, but did not describe it, that being done for him three years 
later by Fischer & Meyer who based their description on plants grown 
at St. Petersburg. There is no information as to the channels by which 
the species was introduced into cultivation. I suspect, however, that 
the original seed may have been obtained by Bertero, who col- 
lected the plant near the Rio Quillota, Chile, as early as 1828, and that 
seeds from this source may have been distributed from Turin. 

Amsinckia intermedia Fischer & Meyer, Ind. Sem. Hort. Petrop. 
2: 2 and 26 (1836). 

This name appears bare in the alphabetic list on page 2 of the seed- 
list cited above. On page 26 (p. 1 of reprint) the following description 
is found, "A. intermedia. A. corolla fauce glabra nuda, limbo tubo sub- 
breviore; staminibus ad faucem insertis. — Corollae tubus 1>4 lin. 
longus, limbus fere 3 lin. in diametro, saturate aurantiacus maculisque 
5 saturatioribus pictus. — Species intermedia A. lycopsioidem inter et 
A. spectabilem; a priore dignoscitur insertione staminum, a posteriore 
corollis longe minoribus et praesertim corollae tubo non (ut in ilia) ad 
faucem plicis intrusis semiclausa. — Hab. cum sequente specie [A. spcc- 
tabilis] circa coloniam ruthenorum Ross in portu Bodega Novae Cali- 
forniae. Annua." The seed-list in which this description occurs bears 
the printed censor's date, Dec. 25, 1835. This equals Jan. 5, 1836 of 
the present calendar. 

Through the kindness of Prof. B. A. Keller, Director of the Institute 
and Botanic Garden at Leningrad, I have received authentic material 
of Amsinckia intermedia. This consists of an authentic fragment of the 
species, from the herbarium of Meyer, one of the co-authors of the 
species, and a fine specimen from the plantings in the St. Petersburg 
Garden in 1836. The specimens agree with the interpretation of 
A. intermedia given by Suksdorf, Werdenda 1: 88 (1931). The plant 
is a member of that variable and bewildering island species that Mac- 
bride, Contr. Gray Herb. 49: 12 (1917), and Jepson, Man. Fl. PI. Calif. 
844 (1925), have incorrectly called "A. Douglasiana." Greene, Bot. S. 



198 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

Francisco Bay, 262 (1894), and Jepson, Fl. W. Mid. Calif, ed. 2, 350 
(1911), earlier treated it, partly as A. intermedia and partly as A. spec- 
tabilis. The name, A. intermedia F. & M., is properly applied to the 
polymorphous species which is common in California in the interior 
valleys and on hillsides back from the immediate vicinity of the ocean. 

Amsinckia spectabilis Fischer & Meyer, Ind. Sem. Hort. Petrop. 
2: 2 and 26 (1836). 

This species appears on page 2 of the above publication as a bare 
name in an alphabetic list of seeds. On page 26 (p. 1 of reprint) the 
following description is found: "A. spectabilis. A. corolla fauce glabra 
plicis intrusis semiclausa, limbo longitudine tubi; staminibus ad faucem 
insertis. Species pulchritudine florum insignis atque distinctissima. 
Corolla aurea, limbo 6 lin. in diametro, ad faucem plicis 5, squamulas 
simulantibus, aucta. Annua." 

In 1925 through the kindness of Prof. Boris Fedtchenko, with the 
assistance of Miss Olga Enden, I received two generous fragments of 
authentic specimens of this species. The specimens were grown in the 
St. Petersburg botanic garden in 1835-36. They are given as grown 
from seeds collected at Fort Ross, California, by Wiedemann. These 
fragments were examined by Suksdorf, Werdenda 1: 96 (1931). He 
correctly identified them with the coastal plant that Brand, in Fedde's 
Repert. 20: 319 (1924), has described as A. nigricans. Brand's plant. 
Heller 5614, is from the type-locality of A. spectabilis. I have seen 
many specimens of this plant in various herbaria from numerous garden- 
cultures. While evidently conspecific, these specimens rarely have the 
corollas as well developed as that found in the original culture at St. 
Petersburg in 1835. This is not surprising. I have grown Amsinckia 
in a botanic garden and under glass and have in most cases discovered 
remarkable differences in habit of growth and corolla-size between my 
cultures and the wild specimen from which the seed was obtained. 

Macbride, Contr. Gray Herb. 49: 7 (1917), in his monograph of the 
genus, has treated the coastal plant (the true A. spectabilis F. & M.) 
under the name "A. intermedia." Jepson, Man. Fl. PI. Calif. 844 (1925), 
attempted to follow him and has described the coastal plant as "A. inter- 
media." His illustration, however, is the inland species, which just 
happens to be the true A. intermedia F. & M. Previous to Macbride's 
paper in the writings of Gray, of Greene, and of Jepson, the coastal plant 
appears as "A. lycopsoides." Macbride, 1. c. 5, of course, was quite 
incorrect in applying the name "A. spectabilis" to the smooth-fruited 
A. grandi flora Kleeb ex Gray. The name A. spectabilis F. & M. properly 



1935] JOHNSTON, STUDIES IN BORAGINACEAE, XI 199 

belongs to the strictly coastal plant of California that has small dark 
nutlets, acute more or less denticulate leaves, and a pair of the calyx- 
lobes frequently more or less united. Abrams, Fl. Los Angeles, 335 
(1904), seems to have been the only author who has properly applied 
the names A. spectabilis and A. intermedia. 

Amsinckia lycopsoides Lehmann, Del. Sem. Hort. Hamburg 1831 : 
1 and 7 (1831). 

On the first page of the Hamburg seed-list for 1831 appears the name 
"Amsinckia lycopsoides Lehm.'" The exponent refers to a note on page 
7 where the following is found, '"Genus novum e familia Borraginearum, 
praeter alias notas cotyledonibus 4 distinctissimum. Benthamia Lindl. in 
litteris (non Richard Monog. des Orchidees iles de France et de Bour- 
bon pg. 43, t. 7, fig. 2)." In the seed-list for 1833, p. 3, and 1834, p. 3, 
the binomial appears perfectly bare. In 1835, p. 3, it is listed in com- 
pany with /I. an^Mi^z/o/m Lehm. In 1836, p. 3, and 1837, p. 3, it is listed 
as one of four species, A. angustijolia, A. intermedia, A. lycopsoides and 
A. spectabilis. 

In the writings of Fischer & Meyer the binomial, A. lycopsioides 
Lehm., appears as a bare name in company of A. angustijolia Lehm., in 
the first St. Petersburg list, Ind. Sem. Hort. Petrop. 1: 2 (1835). In 
the next list, 2: 2 (Jan. 1836), it appears with A. angustijolia, A. inter- 
media and A. spectabilis and on page 26 (p. 1 of reprint) has the follow- 
ing note concerning it: "Amsinckia lycopsioides. A. corolla fauce 
barbata, limbo tubo triplo breviore; staminibus corollae tubo paulo 
supra basin insertis. — A. lycopsioides Lehm. delect, sem. h. Hamburg. 
1831. — Tubus corollae 3>^ lin. longus; limbus 2 lin. in diametro, vix 
latior." 

The species, Amsinckia lycopsoides Lehmann, is the type of the genus 
Amsinckia. The Hamburg seed-list in which it was first published is 
dated 1831. That it was actually published that year is proved by the 
review of this publication in the Litteratur-Bericht zur Linnaea (vol. 6) 
which bears the title-page date of 1831. The description of the species, 
Amsinckia lycopsoides Lehm., by Fischer & Meyer, appears in a seed- 
list for the year 1835 but this pamphlet bears a printed censor's date, 
Dec. 25, 1835 which is the equivalent of Jan. 5, 1836 in our present 

It is to be noted that when, in 1831, Lehmann published his generic 
name, Amsinckia, that he definitely associates it with Benthamia of 
Lindley. This latter v'tnerK- name was published by Lindley, in the 
same yea.\ but only a> u no:iua nudum, Lindley, Nat. Syst. 241 (1831). 



200 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

It was undoubtedly based upon material collected by Douglas along the 
Columbia River. This is clearly indicated by specimens in herbaria at 
Cambridge, Kew, London and Geneva. In the Lindley Herbarium at 
Cambridge there is only one sheet that has been determined as Benthamia 
by Lindley. This contains Cuming's no. 512 from Valparaiso and a 
specimen labeled "North West Amer. H. H. G. 1827, Douglas." Lindley 
has written in the corner of the sheet "Benthamia lycopsoides Mihi." 
There are various strong reasons for believing that this sheet in Lind- 
ley's herbarium formerly bore only the material from Douglas and that 
the Cuming material was later added to it, probably after Lindley's 
annotation. At Kew there are two significant specimens. One from the 
Bentham Herbarium is labeled "Benthamia lycopsoides Lindl. M. S. 
sem. ex Amer. occid. ex Douglas, Hort. Soc. Hort. London, 6-6-28." A 
similar sheet from the Hooker Herbarium is labeled "Anchusa, fl. yellow, 
Benthamia Lindl. mss. N. W. Am. Douglas, cult." Lindley published 
only the genus name, Benthamia. The binomial "Benthamia lycop- 
soides" seems to have been published first by DeCandolle, Prodr. 10 : 118 
(1846). This reference is clearly based upon a specimen at Geneva 
bearing the following data: "Benthamia lycopsoides Lindl. ined., Hort. 
Societ. horticult. in Chiswick 6 jun. 1828." The name on the label is in 
the script of Lindley. The source is written by DeCandolle. The date 
given is the same as that found on the sheet in Bentham's herbarium and 
falls within the period when A. DeCandolle visited London for work on 
his Campanulaceae. The herbarium of the Horticultural Society was 
sold to the British Museum. There is a specimen from this source at 
South Kensington labeled: "sandy plains of the Columbia, 1825 (accord- 
ing to Lindley a new genus)." Lindley was in charge of identifying the 
plants grown in the gardens of the Horticultural Society at Chiswick. 
All the specimens mentioned are probably from seeds grown at Chis- 
wick. They all represent the plant recently described as A. simplex 
Suksdorf, Werdenda 1: ii and 53 (1927 and 1931). 

There are a number of good reasons for believing that Lehmann's 
genus Amsinckia and his species A. lycopsoides are based upon Lind- 
ley's genus Benthamia and B. lycopsoides. In the first place shortly 
before 1830 Lehmann travelled in England and met various botanists 
there. He was a well known student of the Boragimiceae. Lehmann, 
in any case, was later in correspondence with Lindley, for he cites his 
authority for Benthamia as "Lindl. in litteris," and we may well believe 
that he received seed or specimens of Douglas's curious borage from 
Lindley and grew it in the Hamburg garden. Lehmann devotes about 
half of his short description of Amsinckia to citing Lindley's unpub- 



1935] JOHNSTON, STUDIES IN BORAGINACEAE, XI 201 

lished Benthamia and its earlier published homonym. The specific 
name used by Lehmann is that selected by Lindley. What is most im- 
portant, however, is that an Amsinckia conspeciiic with Douglas's plant 
was in cultivation in various European botanic gardens under the name 
"Amsinckia lycopsoides." There is a specimen at Kew collected by J. 
Gay in the Jardin des Plantes at Paris in June 1833, only a year and a 
half after Lehmann published Amsinckia. This plant was grown under 
Lehmann's binomial and represents the species collected on the Columbia 
by Douglas. In conclusion it may be noted that the short descriptive 
notes concerning A. lycopsoides, given in 1835 by Fischer & Meyer, 
apply to the plant collected by Douglas. 

A study of Douglas's Journal, p. 116 (1914), fortunately reveals some 
information as to the original source of Amsinckia lycopsoides. The 
plant is evidently that mentioned under the date of May 2, 1825, in an 
enumeration of collections made on "Menzies Island, in the Columbia 
river, opposite the Hudson Bay Company's establishment at Point Van- 
couver." According to Piper, Contr. U. S. Nat. Herb. 11: 620 (1906), 
Menzies Island is that now known as "Haydens Island." The notes by 
Douglas are as follows: "(151) Myosotis sp., annual; hirsute, branch- 
ing; leaves long, entire; linear-lanceolate; flowers bright yellow; tube 
long; mouth of the corolla spreading, with a dark spot opposite teeth; 
seeds not yet known; this very interesting species was found on Menzies 
Island in company with Mr. Scouler, who agreed with me to call it 
Myosotis Hookeri [not Myosotis Hookeri Clarke (1883)] after Dr. 
Hooker of Glasgow; scarce, only three specimens of it were found, two 
of which are in my possession. — I have since found it in abundance 
near all the Indian lodges above the Rapids of the Columbia. S [eeds] ." 
From these notes it is evident that seeds were not obtained on Menzies 
Island and that, later, they were obtained somewhere above the Colum- 
bia Rapids. Amsinckia simplex Suksd. is known only from the general 
vicinity of Portland, Oregon (just south of Menzies Island). It is 
scarcely separable from A. arenaria Suksd. which is reported from the 
Columbia Gorge and in eastern Washington. The name Amsinckia 
lycopsoides (Lindley) Lehmann is properly applicable to these 
concepts. 

It has been shown that Amsinckia lycopsoides Lehm. is based eventu- 
ally upon material collected by Douglas along the Columbia River. In 
subsequent paragraphs I have shown that Lithospermum lycopsoides 
Lehm. (1830) is based upon collections made by Scouler on the north- 
western coast of Washington. In the writings of A. DeCandolle, Prodr. 
10: 118, adnot. (1846). Gray, Synop. Fl. 2: 198 (1878), Macbride, 



202 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

Contr. Gray Herb. 49: 7 (1917), Suksdorf, Werdenda 1: 101 (1931), 
etc., the binomial Amsinckia lycopsoides has been considered as merely a 
nomenclatorial transfer and as based upon Litkospermum lycopsoides. 
The similarity of the specific epithet is a mere coincidence. There are 
no reasons at all for supposing that these two species are identical. I 
have shown that Amsinckia lycopsoides is a plant from along the Colum- 
bia. Lithospermum lycopsoides is an earlier binomial, but since the 
specific name is preoccupied under Amsinckia it can not be legitimately 
transferred to that genus. A new name for the coastal plant of north- 
western Washington is accordingly needed. 

Lithospermum lycopsoides Lehmann, Pugil. 2:28 (1830); Leh- 
mann in Hooker, Fl. Bor. Am. 2: 89 (1838). 

As was his custom in the Pugillus, Lehmann cited no specimens when 
he originally described L. lycopsoides. In the Flora Boreali-Americana, 
in which he contributed the Boraginaceae, however, he repeated his 
original description verbatim and cited the basic specimen. This latter 
is given as "Straits of de Fuca, N. W. America, Dr. Scolder." At Kew, 
from the herbarium of Hooker, there is a specimen that agrees perfectly 
with Lehmann's description and is labelled "Lith. lycopsioides Lehm. 
De Fuca, N. W. Am. Scouler." I agree with E. L. Greene, who has 
written on this sheet that "This, along with fragments in Herb. Benth. 
constitutes the type of Lithospermum lycopsoides Lehm. It has never 
been in cultivation." The plant is undoubtedly conspecific with that of 
northwestern Washington and adjacent Vancouver Island which has 
passed as "Amsinckia lycopsoides" in Piper's Flora of Washington, 
Contr. U. S. Nat. Herb. 11: 480 (1906), and in the monographs by 
Macbride, Contr. Gray Herb. 49: 7 (1917) and Suksdorf, Werdenda 
1: 101 (1931). It is not the same species as Amsinckia lycopsoides 
Lehm., which is based upon specimens collected by Douglas near the 
Columbia. The present plant, a coastal species related to true A. spec- 
tabilis F. & M. of California, strangely has no synonyms. Since the 
specific name is preoccupied under Amsinckia a new name is needed. 
The plant may be called: 

Amsinckia Scouleri, nom. nov. Lithospermum lycopsoides Leh- 
mann, Pugil. 2: 28 (1830) not A. lycopsoides Lehmann (1831). 

Amsinckia Douglasiana A. DeCandolle, Prodr. 10: 118 (1846). 

I have examined the type of this species in the DeCandoUean Her- 
barium at Geneva. It is clearly a species with tessellate nutlets and 
large showy corollas. I consider it conspecific with A. Lemmonii Mac- 



1935] JOHNSTON, STUDIES IN BORAGINACEAE, XI 203 

bride, Contr. Gray Herb. 48: 50 (1916). Suksdorf, Werdenda, 1: 102 
(1931), who has examined authentic material of A. Douglasiana, pre- 
served at the Gray Herbarium, has considered it closely related to A. 
Lemmonii but separable from it. He places these two species together 
in his monograph. Gray erroneously cited the name A. Douglasiana in 
the synonymy of the common inland species of California. Not having 
seen the type of A. Douglasiana, Macbride, Contr. Gray Herb. 49: 12 
(1917), was misled by Gray's erroneous citation and applied it to the 
common inland species of California. The plant treated as A. Doug- 
lasiana by Macbride, and by Jepson, Man. Fl. PI. Calif. 844 (1925), 
who followed him, is properly identified as true A. intermedia F. & M. 
Amsinckia Douglasiana A. DC. is a relatively rare plant of the South 
Coast Ranges of California and was probably originally collected by 
Douglas in San Luis Obispo or southern Monterey counties during his 
journey from Monterey to Santa Barbara and return. 

Amsinckia parviflora Bernhardi, Selec. Sem. Hort. Erfurt. 1833: 
land 4 (Jan. 1834). 

On the first page of the Erfurt seed-list for 1833 two Amsinckias 
appear in the alphabetic list of names, Amsinckia lycopsoides Lehm. and 
A. parviflora Bernh. A reference to the last, fourth but unnumbered 
page of the seed-list gives the following note concerning A. parviflora 
Bernh. : " ( 1 ) Lithospermum calycinum Moris, cui cotyledones 4, s. potius 

2 bipartitae, speciem Amsinckiae sistit, quam A. parvifloram vocarem. 
An A. angustifolia Lehm. eodem planta ?" The list bears a printed date, 
Jan. 18, 1834. No species of Amsinckia are mentioned in the Erfurt 
seed-lists for the year 1832. Amsinckia parviflora Bernh. appears to 
be no more than a mere renaming of Lithospermum calycinum 
Moris. The two names are, accordingly, exact synonyms and apply to 
Bertero's plant from Rancagua, Chile, described and figured by Moris, 
Mem. Accad. Torino 37: 98 tab. 22 (1834). In 1834 Lehmann cited 
Bernhardi's binomial as a synonym of A. angustifolia Lehm. I arn 
inclined to believe this is correct, for as I shall discuss, I suspect that 
Lehmann's species is also based upon Chilean material. 

Amsinckia angnstifolia Lehmann, Del. Sem. Hort. Hamburg 1832: 

3 (1832), nomen; Fischer & Meyer, Ind. Sem. Hort. Petrop. 2: 26 
(1836), description. 

The above binomial appeared as a bare name in the seed-list of the 
Hamburg garden for 1832. It appeared again as a bare name in the list 
for 1833, p. 3, was omitted in that for 1834, and in the list for 1835, p. 3, 
was cited as follows: "Amsinckia angustifolia Lehm. [A. parvifolia 



204 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

Bernh. Sel. sem. h. Erf. 1833)." This reference was repeated in 1836, 
p. 3. In the list for 1837, p. 3, it again appears as a bare name. 

In the first list from the garden at St. Petersburg, 1: 2 (1835) the 
name Amsinckia angtistifolia also appears bare. In the next list from 
St. Petersburg, 2 : 2 and 26 (1836), the name appears in the list of seeds 
and on page 26 (p. 1 of reprint) the following description and refer- 
ences are published: "A. angustifolia. A. corolla fauce glabra nuda, 
limbo tubo duplo breviore; staminibus ad faucem insertis. A. angusti- 
folia Lehm. delect, sem. h. Hamburg. 1832. A. parviflora Bernhardi 
select, sem. h. Erfurt. 1833. Lithospermum calycinum Moris. Enum. 
sem. h. r. bot. Taurinens. 1831 et in Mem. delta Acad. d. Scienze di 
Torino Tom. XXXVII. p. 108. tab. XXII. — Corollae tubus vix 2 lin. 
longus, limbus vix 2 lin. in diametro. — A praecedente [A. lycopsioides 
Lehm.] floribus parvulis et praesertim staminum insertione diversis- 
sima." All the references cited by Fischer & Meyer trace back to 
material, collected by Bertero in central Chile. What is more all the 
garden material, under the name A. angustifolia, seems best referred to 
the Chilean forms of the genus. All authors have applied A. angustifolia 
to the austral plant. There seems every reason for continuing to do so. 
I suspect that the cultures in European gardens were originally from 
seeds obtained by Bertero at Quillota or Rancagua, Chile, and subse- 
quently distributed from Turin by Morris or Colla. 

Omphalodes erecta, sp. nov., herba perennis erecta e caudice laxe 
ramoso oriens pilis mollibus gracillimis subcinerea; caulibus foliosis 
simplicibus vel supra medium sparsissime fertiliterque stricto-ramosis 
3-6 dm. altis partibus maturis plus minusve glabrescentibus brunnescen- 
tibus 2-4 mm. crassis; foliis lanceolatis vel late lanceolatis 5-11 cm. 
longis 15-30 mm. latis (superioribus non-conspicue reductis) sub 
medium apicem versus in acuminem 1-3 mm. longam gracilem gradatim 
attenuatis, margine integerrimis basi angulatis vel subrotundis 3-6 mm. 
longe petiolatis, supra viridis sparse inconspicueque pubescentibus non 
rariter minute pustulatis, subtus pallidis saepe pilis abundantibus 
longioribus subcinereis; infiorescentia gracili laxe racemosa simplice vel 
basaliter furcata ebracteata 5-15 cm. longa 0-1 cm. longe pedunculata; 
pedicellis ad anthesin 3-6 mm. longis ascendentibus, fructiferis ad 2 cm. 
longis saepe decurvatis vel subcontortis; calyx ad anthesin pallide dense- 
que strigoso, lobis 5 inaequalibus lanceolatis ca. 4 mm. longis; corolla 
coerulea vel medium versus violacea, tubo ca. 3 mm. longo, appendiculis 
faueium 5 trapeziformibus ca. 1.3 mm. longis et latis apice subemargina- 
tis, margine pubescentibus, limbo ca. 13 mm. diametro patente ultra 



1935] JOHNSTON, STUDIES IN BORAGINACEAE, XI 205 

medium lobato, lobis 4-5 mm. longis rotundis, sinibus loborum incon- 
spicue plicatulis; antheris oblongis inclusis medium versus bubo corollae 
affixis; filamentis perbrevis; stylo ad anthesin 2 mm. longo, fructifero 
conspicuis 9 mm. longo; stigmato disciform!; fructu 4-ovulato; nuculo 
solitario (3 abortis) minute appresseque strigoso depresse lateque 
ovoideo, (cum alis) ca. 8 mm. diametro, margine evidenter 1-1.3 mm. 
late alato, ala plana patenti leviter denticulata, dorse nuculae convexo. 

Mexico: common in dense oak-wood along an arroyo near Santa Ana, 
between Alamar and Taray, Sierra Madre Oriental, ca. 25 km. s. w. of 
Galeana, Nuevo Leon, corolla blue with a light violet center, July 3, 1934, 
C. H. & M. T. Mueller 992 (type, Gray Herb.) ; scattered in fields and 
waste places in canyon above Alamar, Sierra Madre Oriental, 1500-1800 
m. ah., June 2, 1934, C. H. & M. T. Mueller 680 (G). 

A remarkable species differing from all its congeners in its coarse erect 
habit of growth. The general habit and appearance of the plant, indeed, 
is more suggestive of Cynoglossum than of Omphalodes. From the 
American species of its genus it is further distinguished by its large 
solitary nutlets which possess a weakly denticulate and spreading wing, 
rather than a strongly toothed upcurved one. The foliage of O. erecta 
is very distinctive. All the American species of the genus have long- 
petioled more or less cordate leaves. The new species has them very 
short-petioled and lanceolate. Only one Mexican species, O. aliena, 
has a similar bractless inflorescence. The plant is a remarkable addition 
to the list of Mexican Boraginaceae. 



JOURNAL OF THE ARNOLD ARBORETUM 



LORANTHACEAE COLLECTED IN THE SOLOMON ISLANDS 

BY L. J. BRASS AND S. F. KAJEWSKI, ON THE ARNOLD 

ARBORETUM EXPEDITION, 1930-1932 

With plate 129 

Amylotheca sp. 

San Cristoval Island: Waimamura, Brass 2849, Sept. 
9, 1932, "common, parasitic on rain forest trees, stout shrub, branches 
smooth and glaucous, leaves very thick and fleshy, perianth of unopened 
flowers lower half pale red, upper greenish-yellow." 

Indeterminable for lack of open flowers, but closely allied to Amylo- 
theca Versteegii (Lauterb.) Danser from New Guinea and New Ire- 
land, differing, however, by longer-pedicelled lateral flowers of the triads, 
more thickish inflorescences and flowers, obtuse bracts and more distinct 
calyx lobes. 

Dactyliophora salomonia, n. sp. Plate 129 a-b 

Glabra, inflorescentiis floribusque iuventute forte parce tomentellis 
exceptis. Rami robusti; internodia foliifera teretia, iuventute apicem 
versus applanata nonnunquam ancipita, nodis dilatatis, postea teretia, 
3-5 mm. crassa, nodis ad sesquiplo crassioribus. Folia opposita vel 
subopposita; petioli 14-24 mm. longi, basi teretes, laminam versus facie 
superiore profundius canaliculati; laminae triangulari-ovatae, plerumque 
10-15 cm. longae, 5.5-7.5 cm. latae, basi rotundatae vel leviter cordatae, 
in petiolum contractae, apicem obtusum versus gradatim attenuatae, 
crassiusculae, fragiles, penninerves, costa facie inferiore basin versus 
magis prominente, costa cetera et nervis lateralibus primariis valde 
incurvatis utrinque distinctis paulum prominentibus. Inflorescentiae 
singulae in axillis foliorum et plures circum nodos defoliatos; pedunculi 
teretes, apice basique paulum incrassati, 15-20 mm. longi, 1-1.25 mm. 
crassi; axes ex internodiis 2 vel 1 compositi, quorum inferius 1-5 mm. 
longum, superius brevissimum; nodi deinceps circiter 8, 6, paucas 
triades ferentes; pedicelli triadum inferiores 8-10 mm. longi, c. 0.3 mm. 
crassi, superiores paucis mm. breviores; pedicelli florum lateralium 2-3 
mm. longi; bracteae bracteolaeque ovatae obtusae 1.25-1.5 mm. longae. 
Calycis tubus obovato-campanulatus, 3.5-4 mm. longus, 1.5-2 mm. 
latus, limbus brevissimus erectus vel nonnihil inflexus. Corolla statu 



1935] DANSER, LORANTHACEAE IN THE SOLOMON ISLANDS 207 

alabastri adulti 25 mm. longa, tertia parte inferiore inflata ad 3 mm. 
lata, tertia parte media gradatim ad 1 mm. attenuata, tertia parte 
superiore in clavam 6-angulam obtusam 1.5-2 mm. crassam incrassata, 
postea divisa in petala 6 sublinearia, a basi c. 1 mm. lata in duabus 
tertiis inferioribus gradatim ad 0.5 mm. angustata, in tertia parte 
superiore angustissime spathulata, apice acutiuscula crassiuscula, latere 
interiore c. 2 mm. supra basin squamula minima. Filamentorum pars 
libera c. 2 mm. longa; antherae c. 4 mm. longae, obtusae. Stylus a basi 
c. 0.6 mm. crassa gradatim angustatus, sub stigmate c. 0.25 mm. crassus; 
stigma obovatum, styli apice vix crassius. Fructus obovato-ellipsoides, 
ad 9 mm. longi^ 6 mm. crassi, superea collo solido 2 mm. longo et lato 
coronati, calycis rudimento vix ullo. 

Guadalcanal Island: Vulolo, Tutuve Mt., 1200 m. alt., 
Kajewski 2497, April 14, 1931 ; ''common, a loranthus growing on trees; 
there are two different coloured flowers, yellow and yellow pink, but 
there is no specific difference; fruit green when ripe, length 1.1 cm., 
diameter 6 mm., with a white flesh inside; fruit with a blunt point at 
end; the leaves are heated and rubbed on sore legs." — Vernacular name 
"Bitorchi." 

The genus Dactyliophora was known, before now, only from New 
Guinea and New Ireland. Dactyliophora salomonia is closely allied to 
the New Guinea D. verticillata (Scheffer) Van Tieghem, and perhaps 
only a variety of it, but it differs by less cordate, longer-petioled leaves, 
axillary inflorescences, longer and less thick peduncles and pedicels, 
more slender calyces and shorter corollas. The peculiar beak on the 
fruit is not described for any other species, but as in most Dactyliophorae 
the fruit are unknown, and those of D. Novae-Guineae (Bailey) Danser 
are figured with a similar prolongation (cfr. BuU. Jard. Bot. Buitenz., 
ser. 3, 11: 359, fig. 14, h), it probably is not a characteristic of D. 
salomonia only. 

Sogerianthe versicolor, n. sp. Plate 129c-e 

Omnis glabra (vel pedicellis, bracteis, calycibusque minute puberulis). 
Rami graciles. parce dichotome ramosi, internodiis foliiferis 1-2.5 (-3) 
mm. crassis, plerumque 3-9 cm. longis, nodis valde incrassatis, duplo vel 
fere triple crassioribus. Folia opposita; petioli 5-10 mm. longi, supra 
applanati praesertim laminam versus, subtus rotundati; laminae oblongo- 
ovatae, plerumque 6-10 cm. longae, 2.5-4.5 cm. latae, sub basi rotundata 
in petiolum contractae vel magis attenuatae, apicem obtusum versus 
magis minusve acuminatae, crassiusculae, fragiles, utrinque opacae (vel 
facie superiore lucidulae), penninerves, costa facie inferiore prope basin 
valde prominente, ceterum costa nervisque lateralibus primariis incurva- 



208 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

tis utraque facie visibilibus vix prominentibus. Flores singuli vel in 
umbellis paucifloris omnino sessilibus in scrobiculis corticis inserti, pauci 
in axillis foliorum vel plures circum nodos foliatos et defoliates; pedi- 
celli teretes, basi saepe paulum clavati, 2-4 mm. longi, 0.3 (-0.5) mm. 
crassi; bracteae bracteolaeque triangulares vel ovatae, 0.5-1 mm. 
plerumque 0.75 mm. longae, obtusae vel acutae, basi nonnihil connatae. 
Calycis tubus campanulatus, subcylindricus, (1.5-)2-2.S mm. longus, 
1.25 mm. latus, limbus erectus paulum cupuliformis, 1(-1.25) mm. 
longus, ore (1.75-)2 mm. lato, integerrimus (vel irregulariter incisus). 
Corolla sympetala, statu alabastri adulti (24-) 30-31 mm. longa, in 
duabus tertiis inferioribus fusiformiter inflata ad 3 mm. lata, in tertia 
parte superiore subcylindrica, c. 1.5 mm. lata, apice obtusa, postea divisa 
usque ad dimidiam longitudinem in lacinias 6 lineares reflexas et volutas 
apice crassiusculas acutiusculas, facie interiore prope basin squamulis 
nuUis. Filamentorum pars libera 3.5-4 mm. longa; anthera basifixa, 
3.5 mm. longa, obtusa. Stylus corollae aequilongus (vel paulo longior), 
a basi ad apicem aequicrassus ; stigma globosum, styli apice circiter 
sesquiplo crassius. Fructus ellipsoides, ad 8 mm. longus, 4 mm. diametro, 
calyce integro erecto paulum aucto coronatus. 

San Cristoval Island: Waimamura, lowlands, on rain 
forest trees. Brass 2676 (type), Aug. 18, 1932, "plentiful, leaves very 
pale, corolla-tube white, segments pink, filaments pink, style green." 
Malaita Island: Quoimonapu, 200 m. alt., rain forest, 
Kajeivski 2355, Dec. 12, 1930, "common, a large loranthus growing on 
the rain forest trees, base of corolla pink, ends of petals white cream." 
Ysabel Island: Tiratona, 600 m. alt., 5rfl55 3227, Nov. 26, 
1932, "common, leaves stiff, margins incurved; perianth reddish, with 
brown lobes." — Vernacular names "Oong" (under no. 2355), and 
"Buraronu" (under no. 3227). 

Description after the type Brass 2676, the dimensions between 
brackets after Kajewski 2355, which mainly differs by shorter corollas 
24-26 mm. long, and puberulous pedicels, bracts and calyces. The 
number Brass 3227 is much like the type, but the leaves are much 
smaller, including the petioles 2-5.5 cm. long and 1-1.6 cm. broad. 

The new species does not show the articulation in the pedicel nor the 
scales at the inside of the corolla tube considered characteristic for the 
genus Sogeriantke till now. As, however, it has a 6-merous long-tubed 
sympetalous corolla and 3 bracts at the base of the flower, and moreover 
agrees with the species already known in general appearance, I do not 
hesitate to place it in the same genus with 5. sogerensis (S. Moore) 
Danser and S. scssiliflora Danser. 



1935] DANSER, LORANTHACEAE IN THE SOLOMON ISLANDS 209 

Dendrophthoe falcata (Linn, fil.) Ettingshausen in Denkschr. 
Akad. Wiss. Wien, Math.-Naturwiss. CI. 32: 53 (1872). — Danser in 
Bull. Jard. Bot. Buitenz. ser. 3, 11: 403 (1931). 

Loranthus jalcatus Linn, fil., Suppl. 211 (1781). 

Guadalcanal Island: Berande River, sea level, Kajew- 
ski 2415, Jan. 6, 1931, "common, a loranthus growing on rain forest 
trees, petals green-cream with orange edges, very showy, the largest fruit 
on specimens, the natives say are pretty full growth, length 1 cm., 
diameter 4 mm.; the natives use this plant superstitiously to stop rain 
by placing twigs upright in the ground." — Vernacular name "Ti-nu- 

Distribution : from tropical southeastern Asia all over the 
Malay Archipelago to tropical Australia, but before now not collected 
farther eastward than the Bismarck Archipelago. 

For the very numerous synonyms cfr. Verh. Kon. Akad. Wetensch. 
Amsterdam, Afd. Natuurk., sect. 2, 29, 6: 44 (1933). 

Notothixos leiophyllus K. Schumann in Schumann & Lauterbach, 
Nachtr. Fl. Deutsch. Schutzgeb. Slidsee, 260 (1905). — Danser in 
Bull. Jard. Bot. Buitenzorg, ser. 3, 11: 456 (1931). 

Ysabel Island: Tataba, 50 m. alt., parasitic on Ranches 
of tall rain forest trees, Brass 3432, Jan. 4, 1933 ; "plentiful, small much 
branched shrub, leaves brittle, underside glaucous in old leaves, indu- 
mentum golden yellow." 

Distribution : Philippine Islands, eastern part of the 
Malay Archipelago, Queensland, but before now not farther eastward 



EXPLANATION OF PLATE 129 
: Da-ctyliophora salomoiu'a (type, Kajewski 2497) ; a, twig 
ind inflorescences in bud, X j^; h, fruit, X 1. Fig. c-e : 
versicolor (type, Brass 2676) ; c, twig with flowers in bud, 



JOURNAL OF THE ARNOLD ARBORETUM 



CHROMOSOME NUMBERS IN THE HAMAMELIDACEAE 
AND THEIR PHYLOGENETIC SIGNIFICANCE 



IVith three text figures 
Reinsch' introduced his morphological survey of the Hamamelida- 
ceae by the observation that they form one of those natural families as 
to whose precise delimitation and relationships there has been much 
difference of opinion. The forty years which have elapsed since the 
appearance of his paper have merely provided further illustrations of 
the justice of his remarks. Though universally conceded to be a natural 
group, the only general agreement as to its phylogenetic position seems 
to be the opinion that it occupies an important one. Because of this 
uniformly recognized phylogenetic significance an effort has been made 
to make as complete a survey of chromosome numbers as possible. The 
living collections of the Arnold Arboretum fortunately include several 
genera such as Sinowilsonia and Parrotiopsis which are very rare in 
cultivation but the work has been hindered by the very great technical 
difficulties involved. The chromosomes are small, there is much sec- 
ondary pairing, the cytoplasm is murky and the chromosomes do not 
stain sharply. In most of these details the family shows cytologically 
a strong resemblance to the Rosaceae, paralleling the morphological 
resemblances which have been commented on by most students of the 



The following chromosome counts have been made. The genera are 
arranged according to the classification of Harms in Engler and Prantl. 
In each case the counts were obtained from aceto-carmine smears. Typi- 
cal meiotic plates are illustrated in Figure 1. 




1935] ANDERSON AND SAX, CHROMOSOMES IN HAMAMELIDACEAE 211 

Meiotic irregularities, accompanied by a high percentage of pollen 
sterility were encountered in Liquidambar Styraciflua. This is some- 
what puzzling since this species exhibits none of the morphological 
peculiarities which are usually associated with irregular meioses. It is 
a "good" species with no closely related forms occurring within the same 
area. Its behavior is more probably to be explained as due to climatic 
influences. It is a southern species and at the Arboretum is being culti- 



•«?/;«: -I 



4 

-; 






3 ••«•' 






•• • » • ' ^J\ 






\» 



Figure 1. Camera lucida drawings (X 3000) of pollen mother cells: 
1. Liquidambar Styraciflua. — 2. Parrotiopsis Jacquemontiana. — 
3. Fothergilla major. — 4. Corylopsis pauciflora. — 5. Sinowilsonia 
Henryi. — 6. Hamamelis vernalis. 

vated somewhat north of its natural range. Whitaker' has demon- 
strated the effect of abnormal temperatures upon meiosis in Cypho- 
mandra. It is possible that the irregular chromosome behavior and con- 
sequent pollen sterility of Liquidambar Styraciflua at the Arnold Arbo- 
retum may have a similar explanation. It would be interesting to know 
if L. Styraciflua is characterized by low percentages of fertile pollen in 

The cytological studies present a number of facts of 
nificance. 1. The Hamamelidoideae are a coherent group ^ 



212 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

mon base number. 2. The count on Liquidambar suggests that the 
Liquidambaroideae may possibly be derived from a different stock than 
the Hamamelidoideae since they apparently have a different base chro- 
mosome number. If this difference in base number should be found to 
persist in the other genus of that sub-family it would indicate that the 
divergence between the two sub-families occurred before the differentia- 
tion of the family as a whole. This is in accordance with the views 
expressed by Harms.' Summarizing the anatomical evidence he states' 
that the Hamamelidoideae are a unified group anatomically while the 
other subfamilies, particularly the Liquidambaroideae, have many dis- 
tinctive peculiarities. Reviewing the entire evidence of relationship' he 
suggests that the Liquidambaroideae are so distinctive that they might 
well be considered a separate family. 

3. Polyploid series have been found in F other gilla and Corylopsis 
and are not to be unexpected in other genera of the family when these are 
investigated more extensively. This discovery is of some taxonomic 
consequence since it indicates that in such genera we may expect phylo- 
genetic relationships between species which will be, in part at least, 
reticulate. That is to say that a complete phylogenetic tree of the genus 
Fothergilla or Corylopsis would show anastamosing branches. It will 
be noted that Fothergilla monticola has 24 pairs of chromosomes and is 
therefore a tetraploid and that F. major with 36 pairs is a hexaploid. 
The phylogenetic relationships within and between these two species, as 
indicated by these chromosome counts, must be intricate. These two 
species are so similar that it is very doubtful if F. monticola deserves 
more than varietal rank.* The cytological evidence would suggest that 
F. monticola is merely a tetraploid variety which arose spontaneously 
from the hexaploid species F. major. Such relationships are not un- 
known in other genera of the flowering plants. Erlanson for instance 
has shown-' that Rosa acicularis var. Sayi (Schw.) Rehder is an octoploid 
race (2n = 56) of the hexaploid species Rosa acicularis (2n = 42). 

To the larger problem of the phylogenetic position of the family itself 
this cytological survey contributes important evidence, though unfor- 
tunately not as decisive as the obscurity of the case requires. Before 
going into details it may be said that on the whole the cytological evi- 
dence favors Hutchinson's interpretation of the phylogenetic position of 

lEngler Prantl Nat. Ptlanzenf. 2. Aull. 18a: 3Co -U5 (iu.^0). 
-loc. cit. p. 307. 

^Anderson in Arnold Arb. Bull. Pop. Inform. >er. IV, 1 : 61-64 (1933). 



1935] ANDERSON AND SAX, CHROMOSOMES IN HAMAMELIDACEAE 213 

the Hamamelidaceae.^ Comparisons of two treatments of the family 
are presented in Figure 2. The numbers in the diagram are the base 
chromosome numbers so far as they have been determined." ^ 

By inference and by actual experimental test two kinds of change of 
chromosome number have been established as occurring in the higher 
plants; (1) the addition of whole sets of chromosomes, that is of two 




F/a^a/?ace<?e £/ 

Figure 2. The phylogenetic position of the Hamamelidaceae, accord- 
ing to Hutchinson and Engler. The numbers are thi 

sets of six to make a 12 or the addition of an eight ; 
a 17; (2) the gradual stepping up or stepping down of the chrc 
number by fusion and fraction of one or two pairs of the chromosomes 
in the previous set, that is, the derivation of an 11 chromosomed species 
from one with 12, etc. The whole subject is still in the experimental 
stage but it is at least far enough advanced to indicate that these two 
processes are among the main forces involved in the separation of genera 
in the higher plants. It will be seen that higher numbers may be derived 
iThe Families of Flowering Plants. I. Dicotyledons. Macmillan and Co., London 



214 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

from lower by either process or by both, but that lower numbers can be 
derived from higher ones only by the second. Everything else being 
equal, therefore, those genera with high base numbers will be farther 
out towards the tips of the phylogenetic net-tree than those with much 
lower numbers. In the present case the cytological evidence favors the 
view that the Hamamelidaceae with their base numbers of 12 and 15 are 
derived jrom the Rosales stock where base numbers of 6, 7, and 8 are 
characteristic. Another cytological fact points in the same direction. 




Figure 3. Phylogenetic relatic 
gested by the cytological evidence, 
numbers. Further explanation in the text. 

There' is very strong secondary pairing throughout the family. As 
shown in figure 1, the chromosomes are not scattered equally over the 
plate but tend to be more or less grouped. This phenomenon which was 
first described by Darlmgton and Moffett^ and which has been exten- 
sively studied by Lawrence- indicates that the chromosome complement 
under observation arose ultimately from the duplication of separate 
complements. 

These facts, together with such other information as bears upon the 
subject, have been utilized in constructing the diagram in Figure 3. It 
ijour. Gen. 22: 129-151 (1930). 



1935] ANDERSON AND SAX, CHROMOSOMES IN HAMAMELIDACEAE 215 

should be emphasized that the diagram is purely speculative. It has 
been worked out for those morphologists who would be interested in 
knowing how a cytologist with such information as is available would 
speculate as to the relationships of the groups involved. It might well 
be used as one of a set of possible working hypotheses by students of 
phylogeny. While the anastamoses of the main trunks of the Rosales 
stock represent supposed true-breeding allo-polyploid hybridizations, 
they do not necessarily indicate a cross between families as such. On 
any evolutionary hypothesis, related families derive, ultimately, from 
forms no more differentiated than present day genera or species. All 
that need be hypothecated for these hybrids is that they are between 
forms as diverse morphologically as certain hybrids which have been 
experimentally obtained, those between Zea and Tripsacum, for in- 
stance.^ The diagram is based upon the evidence from chromosome 
number, secondary association and, in the case of the Pomoideae, from 
breeding experiments. It is much more speculative for the Saxifragaceae 
than for the Rosaceae. The Saxifragaceae, with base numbers of 8, 11, 
13, 14, and 16 show a cytological complexity- paralleling their morpho- 
logical diversity. Only a few of the fossil "dead branches" have been 
indicated. There must certainly have been many more. In this respect 
as in several others the actual details of the diagram are probably in- 
correct. The general conception, however, of a more or less webbed 
net-tree for the Rosales is strongly supported by the cytological evi- 
dence. In some other groups of the flowering plants (the Tubifiorae, for 
instance) the webbing would be so much more complex that one would 
scarcely use the word tree in describing it. In the Cyperaceae, on the 
other hand, there would be few if any anastamosing branches. The 
cyto-genetic evidence shows with increasing force that the actual pattern 
of evolutionary progress has been different in different groups of plants. 
The main point of the diagram in figure 3 is to suggest the general nature 
of the evolutionary pattern of the Rosales. 

SUMMARY 

1. Chromosome counts are given for nine species and six genera of 
the Hamamelidaceae. 

2. The phylogenetic position of the family is discussed in the light 
of these results. 

Harvard University. 
iMangelsdorf & Reeves in Jour. Hered. 22: 329-343 (1931). 



JOURNAL OF THE ARNOLD ARBORETUM 



IVith plates 130 and 131 

Coiled chromonemata have been observed in both mitotic and 
meiotic chromosomes in various species of plants. The degree of coiling 
in mitotic chromosomes may vary from an irregular corrugation or loose 
spiral to a rather compact regular coil. At meiosis in certain plants the 
coils are much larger and can be analyzed in more detail. 

The behavior of the meiotic chromosomes of Rkoeo discolor provides 
additional information regarding the nature of the coiled chromonemata. 
The observations are based on both aceto-carmine preparations and per- 
manent smears fixed in Flemming's solution. In both cases the micro- 
sporocytes, after smearing, were usually pretreated with alcohol and 
ammonia before fixing. Immersion in tap water for a few seconds and 
a brief exposure to ammonia fumes also gave good results. This is 
essentially the method used by Kuwada. The ammonia vapor seems to 
dissolve the chromosome matrix and permit the spiral chromonemata to 

The chromosomes of Rhoeo are arranged in a ring or one or more 
chains at the first meiotic division. At early metaphase each chromo- 
some contains a coiled chromonema consisting of two closely associated 
chromatids. The general appearance of these rings is shown in the 
photographs from aceto-carmine preparations (Figs. 1 and 2), but the 
finer details are best observed in permanent smears ( Figs. 3 and 4 ) . The 
number of coils is four or five per chromosome. At this stage the two 
chromatids are so closely associated that the coils appear to be single, 
but their double nature can be observed at certain loci. The diameter 
of the coil decreases gradually before the separation of the chromatids. 

At late metaphase the coiled chromatids separate. The number of 
coils in each chromatid remains the same, but they are much smaller, 
even though there is little or no elongation of the chromosome as a whole 
(Figs. 5 and 6). At this stage both the terminal chiasmata and the 
fiber constrictions are very conspicuous, so that the order of the indi- 
vidual chromosomes in the ring can be determined. As observed earlier 
(Sax 1931), the order of the twelve chromosomes is always the same, 
thus supporting Belling's suggestion that such rings are the result of 



1935] SAX, CHROMOSOME STRUCTURE IN RHOEO DISCOLOR 217 

segmental interchange. Six of the twelve chromosomes are distinctly 
heterobrachial, and the short arms are paired. 

The meiotic chromosomes are always paired by terminal "chiasmata" 
and, in the chromatids at late metaphase, most of the chiasmata appear 
to be symmetrical. The chromatids are often parallel in several suc- 
cessive chromosomes or even in most of the chromosomes in the ring. 

As the chromosomes pass to the poles at the first meiotic division, the 
chromatids separate except at the fiber constriction and become shorter 
(Fig. 7). During interphase the chromatids elongate considerably. At 
the second meiotic division they are about twice as long as they were 
at the first division. At this time the minor or somatic type of coiling 
can be observed. The coils are only about half as wide as the major 
coils found in the first division, and the number of coils is about 12 per 
chromatid (Fig. 8). These coils appear to be single at metaphase. 
There is some evidence of a split at late anaphase, as indicated by 
narrow regions at certain loci. If the chromatids are split at second 
metaphase, the two halves must be coiled together. At late anaphase 
they may tend to separate, but the slipping apart of the coils is difficult 
to detect except where there is a twist which appears to constrict the 
chromosome at such loci. More definite evidence of split chromatids 
has been obtained from microsporocytes which were subjected to low 
temperatures during development. 

Under normal conditions the microspores receive six chromosomes, 
but occasionally there are seven, owing to irregularities in the first 
meiotic division. About 80 per cent of the microspores fail to develop, 
owing to segmental non-disjunction. The normal fertile microspores 
undergo a single nuclear division, followed by the differentiation of the 
daughter nuclei into the large and more or less degenerate tube nucleus 
and the compact elongated generative nucleus. 

When the plants are kept at a temperature of about 6° C, two kinds 
of abnormalities appear. The chromosomes of the one nucleate micro- 
spore may divide but do not form daughter nuclei. The 12 chromosomes 
pass back into the resting stage and divide regularly at the next division 
to form diploid gametes. The other type of abnormal development 
begins when the low temperature inhibits chromosome pairing at meiosis. 
The twelve univalent chromosomes pass into the resting stage without 
nuclear division. They come out of the resting stage, divide without 
nuclear division and form a giant nucleus with 24 chromosomes, each of 
which consists of two coiled chromatids held together only at the fiber 
constriction (Fig. 9). Throughout this process the chromosomes never 
pass through the contraction characteristic of normal telophase stages. 



218 JOURNAL OF THE ARNOLD 

The chromatid spirals are much looser than those found in the chroma- 
tids of the normal chromosomes at the second meiotic division, and it is 
perfectly clear that many of these chromatids are split. The splits are 
especially clear near the ends of the chromatids. At a somewhat later 
stage the two chromatids become completely separated, the chromatids 
elongate, and their structure is very clear. In the same chromatid one 
can observe the transition from a single coil to two parallel finer coils 
(Fig. 10). The mechanism of separation of daughter chromatids in 
these chromosomes, which are essentially of the somatic type, is similar 
to that found in the coiled chromatids in the first meiotic division. 

The structure of the meiotic chromosomes in the permanent prepara- 
tions was clear enough to permit an analysis of direction of coiling in 
the spiral chromonemata. We were able to determine the direction of 
coiling at all loci in each of the twelve chromosomes in 14 cells with com- 
plete chromosome rings. The classification of right- or left-handed 
spirals is purely arbitrary, since the direction depends on the sequence 
of determination in the ring. There is a strong tendency for the direc- 
tion of coiling to be in the same direction in both arms of a chromosome. 
Of the 168 chromosomes examined, 50 had a right-handed spiral in both 
arms, 52 a left-handed spiral, and 66 showed a reversal of coiling, pre- 
sumably at the fiber attachment. In only two chromosomes was there a 
second change of direction of coiling. Individual chromosomes could not 
be identified consistently, so that the direction of coiling could not be 
established for any one chromosome in all the different cells, but there is 
good evidence that direction of coiling is not a stable character. The 
number of chromosomes with left-handed coils ranged from 1 to 5, with 
reversed coils from 2 to 8, and with right-handed coils from 1 to 7, in 
different cells. The direction of coiling of chromonemata of paired 
adjacent chrtmosome arms is at random, with 85 coiling in the same 
direction and 83 coiling in reverse directions. 

Both rings and chains of chromosomes are found at meiosis. If one 
or more chains are formed, the breaks may occur between either the long 
segments or the short segments. Three of the terminal chiasmata are 
formed between short segments, and eight of the terminal chiasmata are 
formed between long segments. The extra chiasma is between a long 
and a relatively short segment. The position of the breaks, or failure 
of chiasma formation, was obtained for 20 cells containing one or more 
chains. There were 9 breaks between the short segments and 16 between 
the long segments. These results suggest that chiasma formation is some- 
what less likely to occur, or less likely to persist until late metaphase, 
between the short segments. The latter possibility is more probable 



1935] SAX, CHROMOSOME STRUCTURE IN RHOEO DISCOLOR 219 

because we have found chromosome rings in practically ail cells in some 
preparations. 

The lengths of the chromosomes at different stages in meiosis were 
obtained to aid in the analysis of factors involved in chromosome con- 
traction. It was not possible to get an accurate measurement of the 
length of the pachytene spireme in Rhoeo, but the total length is 
approximately 700 microns or an average of somewhat more than 100 
microns per chromosome. The approximate average chromosome length 
at meiotic metaphase is 5-6 microns, and is about 9 microns at the 
second meiotic division. 



The great contraction in chromosome length between pachytene and 
meiotic metaphase stages in Rhoeo is associated with the coiling of the 
chromonema. This coiling may not be the only factor involved. Belling 
(1928) believed that the approximation of chromosomes caused about 
one-third of the contraction in the chromosomes of Lilium, and that the 
coiling of chromonema effected the final shortening to give approximately 
a 10 to 1 reduction in length of the meiotic chromosome. Bridges 
(Alexander, 1928), on the other hand, assumed that coiling is the 
primary factor in chromosome contraction, and that the gene string 
maintains approximately the same length at all stages in the chromosome 
cycle. Another factor in chromosome contraction is the secondary or 
minor coils within the primary or major coils as described in Tradescan- 
tia by Fujii, Kuwada and Nakamura (1933) and found in Sagittaria 
by Shinke (1934). 

We believe that three factors are involved in the great decrease in 
length of the meiotic chromosomes of Rhoeo; first, a linear contraction 
of the gene string; second, the major coiling of the chromonema; and 
third, the formation of minor spirals within the major spiral. The minor 
coils are not clearly differentiated at the first meiotic division in Rhoeo, 
but there is some evidence of loose coiling. The contraction of the 
chromonema and reduction in the width of the major coils between 
early and late metaphase are attributed to the further coiling of the 
minor spirals in each chromatid. A similar reduction in the major 
coils with no increase in chromosome length is found in Secale (Sax, 
1930). 

The coiled chromonema at early metaphase consists of two chroma- 
tids coiled together so that the chromonema often appears as a single 
coil, as is the case in Tradescantia, Secale, Lilium, and Vicia. The free 
separation of coiled chromatids has been explained by Kuwada (1927). 



220 JOURNAL OF THE ARNOLD ARBORETUM 

The reverse twists postulated by Kuwada < 
scantia (Sax and Humphrey, 1934) and 
Smith, 1935). 

The direction of coiling in the chromonema spirals is more or less at 
random in Rhoco and in Tradescantia (Nebel, 1932, Sax and Humphrey, 
1934). According to Huskins and Smith, the paired meiotic chromo- 
somes of Trillium usually coil in opposite directions, although no statis- 
tical evidence is presented, nor is there any adequate explanation for 
such behavior. Within a single meiotic chromosome the direction of 
coiling may change at the spindle fiber point, but is seldom reversed at 
other loci in Rhoeo, Secale, Gasteria (Taylor, 1931), Tradescantia, and 
Sagittaria (Shinke, 1934). Huskins and Smith find frequent changes 
in direction of coiling of anaphase chromosomes in Trillium. These 
changes in direction of coiling are usually associated with chiasmata. 
We have found changes in direction of coiling at chiasmata in the meiotic 
chromosomes of Vicia. If the chromonema coiling is caused by a con- 
traction of the matrix, as suggested by Kuwada, the fiber attachment 
points and the chiasmata would tend to break any continuity of stress 
on the chromonema and changes in direction of coiling would be 
expected to be more or less at random at these points. There is a strong 
tendency, in both Rhoco and Tradescantia, for the direction of coiling 
to be the same on both sides of the spindle fiber attachment, and only 
about one-third of the chromosomes show reversal of coiling at this 
locus. Huskins and Smith find that the reversals in direction of coiling 
between the fiber attachment and the distal ends of the chromatids at 
first anaphase is about twice the chiasma frequency at metaphase. This 
relation would be expected if the direction of coiling in homologous 
chromosomes is at random, and if reversals in coiling occur at random 
at the chiasmata. 

In Rhoco all chiasmata are terminal, and most of them appear to be 
symmetrical. The short chromosome arms are paired almost as fre- 
quently as the long arms. Changes in direction of coiling are rarely 
observed between the fiber and the distal end of the chromosome. These 
observations seem to indicate that chromosome pairing in Rhoeo is not 
dependent on the formation of interstitial chiasmata, but is dependent 
on a terminal association of homologous chromosome segments. This 
terminal association in certain rod bivalents in Tradescantia seems to 
involve the chromosome pellicle or matrix, but in Rhoeo there is evi- 
dence of fine chromatic connecting fibers. 

During interphase the meiotic chromosomes of Rhoeo elongate but 
maintain some evidence of loose coils during the resting stage. At the 



1935] SAX, CHROMOSOME STRUCTURE IN RHOEO DISCOLOR 221 

second ftieiotic division, a new spiral appears which has finer and more 
numerous spirals than the major coils at the first meiotic division. These 
minor spirals have been described in Lilium, Rhoeo, Allium, Tricyrtis, 
Najas, and Hosta by Shinke (1930), and in Tradescantia by Nebel 
(1932), Kuwada and Nakamura (1933) and Sax and Humphrey (1934). 
These coils in Rhoeo are wider than the minor spiral' within the major 
spiral at the first meiotic division and presumably are formed inde- 
pendently during the prophase of the second meiotic division. These 
spirals are much like those found in certain somatic chromosomes. The 
transition from about five major spirals to 20-25 minor spirals in the 
successive meiotic divisions in Tradescantia and a similar behavior in 
Rhoeo is difficult to reconcile with the "heterogonic growth" hypothesis 
of spiralization suggested by Huskins and Smith. 

The anaphase chromosomes at the second meiotic division have been 
described as two-parted in a number of plants (Gasteria, Taylor, 1931; 
Galtonia, Smith, 1932; Tradescantia, Nebel, 1932; Trillium, Huskins 
and Smith, 1935; et al.; Cf. Sharp, 1934). Both Kuwada and Naka- 
mura, and Sax and Humphrey found only single coils in the second ana- 
phase chromosomes of Tradescantia. In Rhoeo there is evidence that 
the anaphase coil is double, but that the two half-chromatids are coiled 
together so intimately that they appear as a single coil at early anaphase. 
As the coils begin to separate the gyres are matched so closely that the 
dual structure is not clear, but where a twist occurs there is a narrow 
region in the chromosome. The abnormal "microspores" of Rhoeo 
show the chromatid splits clearly in various stages of separation. The 
minor coils, characteristic of somatic chromosomes, are similar in struc- 
ture to the major coils; the two chromatids (or half-chromatids) are 
coiled together in parallel in such a manner that they can separate freely 
without entangling. If the split occurs while the chromosome is coiled, 
there must be some lateral polarity so that the division occurs in only 
one plane parallel to the axis of the chromosome, as Nebel (1933) has 
suggested. 

If there is a chromatid split in the anaphase chromosomes of the 
second meiotic division in Tradescantia, the chromatid must behave as 
a single unit until midprophase of the microspore division. Tradescan- 
tia microspores subjected to x-rays show chromatid breaks for about 
two days after raying, but after three or four days only chromosome 
breaks are observed at metaphase (Riley— unpublished). 

We find that either abnormally low or high temperatures will cause 
nuclear irregularities. These include failure of chromosome pairing at 



222 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

meiosis, chromosome division without nuclear division, and failure of 
normal differentiation of nuclei. Since these temperatures are within 
the range occasionally experienced in nature, it is probable that tem- 
perature fluctuations have played an important part in chromosome 
changes in nature (Cf. Randolph, 1932). This work with temperature 
effects has been aided by a grant from the American Academy of Arts 
and Sciences. 

SUMMARY 

The twelve chromosomes of Rhoeo discolor are arranged in a seg- 
mental interchange ring at the first meiotic division. Each chromosome 
contains a spiral chromonema consisting of two chromatids coiled 
together. There is some evidence, both direct and indirect, that there 
is a minor spiral within the chromatids of the major spiral. During 
metaphase the major spirals become smaller, and the two chromatids 
separate. The chromosomes elongate greatly during interphase, but 
there is evidence of a spiral structure during the resting stage. At the 
second meiotic division, new minor spirals are formed which are smaller 
and more numerous than the major spirals of the first division. The 
chromatids at anaphase of the second meiotic division are split, but the 
two half-chromatids are so closely coiled together that they are not easily 
observed. They can be differentiated easily in cells where the normal 
chromosome cycle is disturbed by subjection to low temperatures. 

The reduction in the length of the meiotic chromosomes of Rhoeo, 
between prophase and first metaphase is attributed to three factors:— 
a linear contraction of the gene string, the coiling of the chromonema 
into major coils, and the development of minor coils in the chromatids 
of the major spirals. The direction of coiling in the major spirals 
seems to be at random. In a single chromosome the direction of coiling 
may change at the fiber attachment point, but it is seldom reversed at 
other loci. 

The meiotic chromosomes are paired at the ends, apparently without 
the formation of interstitial chiasmata. 



LITERATURE CITED 
ALEXANDER, J. (1928). Colloid Chemistry. Vol. ii, p. IC 

Chemical Catalogue Co., New York.) 
Selling, J. (1928). Contraction of chromosomes during 

divisions in Lilium and other plants. (Univ. Cal. Pub. Bot. 14:335-343.) 
iusKiNS, C. L., and S. G. Smith (1935). Me" " ' 

ture in Trillium erectum L. (Ann. Bot. 49:119-150.) 



1935] SAX, CHROMOSOME STRUCTURE IN RHOEO DISCOLOR 223 

KuwADA, Y. (1927). On the spiral structure of chromosomes. (Tokyo 
Bot. Mag. 41:100-109.) 

and T. Nakamura (1933). Behavior of chromonemata in 

mitosis. I. Observation of pollen mother cells in Tradescantia reflexa. 
(Mem. Coll. Sci. Kyoto Imp. Univ. 9:129-139.) 

and T. Nakamura (1934). Behavior of chromonemata m 

mitosis. II. Artificial unravelling of coiled chromonemata. (Cytologia, 
5:244-247.) 

Nebel, B. R. (1932). Chromosome structure in Tradescantiae. I. Methods 
and morphology. (Zeit. f. Zellf. u. mik. Anat. 16:251-284.) 

(1932). Chromosome structure in Tradescantiae. II. The 

direction of coiling of the chromonema in Tradescantia reflexa Raf., 
T virginiana L Zebrina pendula Schnizl., and Rhoeo discolor Hance. 
(Zeit. f. Zellf, u. mik. Anat. 16:285-304.) 

(1933). Chromosome structure ir 

history of the chromonemata in mitosis oi 
(Cytologia, 5:1-14.) 

Randolph, L. F. (1932). Some effects of high temperature on polyploidy 
and other variations in maize. (Proc. Nat. Acad. 18:222-229.) 

Sax, Karl (1930). Chromosome structure and the mechanism of cross- 
ing over. (Jour. Arnold Arb. 11:193-220.) 

(1931 ). Chromosome ring formation in Rhoeo discolor. (Cyto- 
logia, 3:36-53.) 

and L. M. Humphrey (1934). Structure of meiotic chromo- 
somes in microsporogenesis of Tradescantia. (Bot. Gaz. 96:353-361.) 

Sharp, L. W. (1934). Introduction to Cytology. (McGraw-Hill Book 
Co., Newr York.) 

Smith, F. H. (1932). The structure of the somatic and meiotic chromo- 
somes of Galtonia candicans. (La Cellule, 41:243-263.) 

Shinke, N. (1930). On the spiral structure of chromosomes in some 
higher plants. (Mem. Coll. Sci. Kyoto Imp. Univ. 5:239-245.) 

(1934). Spiral structure of chromosomes in meiosis in Sagit- 

taria Aginashi. (Mem. Coll. Sci. Kyoto Imp. Univ. 9:367-392.) 

Taylor, W. R. (1931). Chromosome studies on Gasteria. III. Chromo- 
some structure during microsporogenesis and the postmeiotic mitosis. 
(Amer. Jour. Bot. 18:367-386.) 



EXPLANATION OF PLATES 



of Rhoeo discolor. Figs. 3 and 
smears. All others from aceto-carmine preparatic 
Figs. 1 and 2. The coiled chromonemata in t 

Figs. 3 and 4. Coiled chromonemata showing reversal of coiling ; 

duction in width of coiling at metaphase. 
Figs. 5 and 6. Separation of coiled chromatids at late metaphase 



Telopha 
Chromo: 
spirals which appear 



JOURNAL OF THE ARNOLD ARBORETUM [v< 

Plate 131 
from abnormal "microspores" produced by ch 
ication induced by cold treatment. Magnification X 
L giant cell derived from a microsporocyte by chn 
on without nuclear division. Each of the 24 chromosomes 
Dnsists of two chromatids held together at the fiber attachment, 
lany of the chromatids are split. 

hromatids at a somewhat later stage showing the transition 
rom a single coiled chromatid to two coiled daughter 



I Arb. Vol. XVI 
















Jour. Arnold Arb. Vol. X\ 



0*"^^? 






FAULL, ELAIOPLASTS IN IRIS 



ELAIOPLASTS IN IRIS: A MORPHOLOGICAL STUDY 



With plates 132 to 137 
INTRODUCTION 

ELAIOPLASTS are a heterogeneous group of intracellular bodies pre- 
senting the characteristics of fatty substances to a marked degree but 
not recognizable as ordinary types of plastids, chondriosomes or 
vacuoles. There is no general agreement in the literature regarding 
their structure, origin, development, classification or chemical compo- 
sition. They have been variously described as aggregations of lipoid 
globules, as modified or unusual types of plastids or vacuoles, as nuclear 
derivatives, as aggregations of mitochondria-like bodies or as inde- 
pendent structures. They have been linked with various physiological 
processes such as assimilation, excretion or degeneration. 

Much of the confusion regarding elaioplasts is due to the use of 
inadequate techniques and to a consequent lack of accurate information 
about the early developmental stages of these bodies. Many of the 
discrepancies are also due to failure to visualize and interpret correctly 
the full range of morphological variability of chondriosomes and plastids. 

The investigations described in this paper were undertaken to clarify 
our conception of the elaioplasts in Iris and to compare these bodies 
with those in other monocotyledons and in liverworts. By using an 
improved technique critical evidence has been obtained to show the 
early stages in the development of the elaioplasts in Iris and the changes 
which these bodies undergo in different tissues and at different seasons. 

In addition to the morphological study an extraction and preliminary 
analysis of the so-called oil in the elaioplasts of Iris and some physio- 
logical experiments on the metabolism of two types of Iris rhizome, one of 
which contains abundant elaioplasts, have been made. The results of 
these investigations are being published elsewhere. 

HISTORICAL RESUMfi 

Since the middle of the last century papers have appeared from time 

to time describing cytoplasmic bodies associated with oil. The writers 

have used various names for these structures which through usage have 

become more or less interchangeable. Thus they are termed elaioplasts, 



226 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

oleoplasts, oleosomes, olkorper, oil bodies, Zellenblaschen, Zellenkorper, 
fatty bodies, elaiosferer, oelplastids, oleoleucites, elements oleiferes and 
systeme oleifere. Sometimes the terms are restricted in their applica- 
tion. For example olkorper is used only for oily bodies in the liver- 
worts, and elaioplast is kept for those in the monocotyledons. But 
recently with an increasing tendency to consider all of these oily bodies 
essentially similar, one name is often used to designate all of them. 

The earliest references to elaioplasts are found in the writings of 
Mirbel (35) in 1835, of Gottsche (13) in 1843, of Holle (24) in 1857, 
of Hofmeister {25) in 1867 and of Ward (49) in 1883. But the first 
adequate descriptions of oil bodies were published in papers by Pfeffer 
(38) in 1874 and by Wakker (47) in 1888. These, together with a 
paper by Lidforss (32) in 1893, provide a description of the three main 
types of mature oil bodies; from this later authors have diverged little. 
Although often resembling one another, the three main types present 
certain distinct features which are further emphasized by their restric- 
tion to a given group of plants. 

Pfeffer (38) described oil bodies characteristic of the liverworts. 
In common with such bodies in general, they are highly refractive struc- 
tures which stain brilliantly in "fat" dyes, such as alkannin, and which 
are more or less soluble in 95 ',f alcohol and in fat solvents such as ether. 
They are distinguished from other oil bodies by their almost complete 
solubility in alcohol, by a characteristic residual ring left after treat- 
ment with alcohol, by their location commonly in the peripheral cyto- 
plasm but within the chloroplast-bearing layer, by their presence in 
practically every species of the group, by their appearance commonly 
in every cell of a plant and by their permanency as cell structures. The 
Marchantiales present a contrast to other elaioplast-bearing hepatics in 
the restriction of the oil bodies to special cells scattered throughout the 
thallus and in the location of a single large oil body in the center of each 
of these cells. Oil bodies in the liverworts vary in shape from round to 
spindle-shaped as a rule, though some are irregular in form. They 
vary in color from colorless to dark brown and in appearance from 
granular to segmented or homogeneous. 

The oil bodies described by Wakker (47) differ from those in the 
liverworts in their location near the nucleus, in their invariably granular 
appearance, in their often irregularly lobed shape, and in their charac- 
ter of being more or less temporary cell structures. Elaioplasts of this 
type are often yellowish in color and are marked by their reaction with 
some reagents which cause an extrusion of the oil and leave a charac- 
teristic net-like structure. Although reported from most tissues, they 



19351 FAULL, ELAIOPLASTS IN IRIS 227 

are often restricted to certain ones. Raciborski (41) and Beer (4) 
found them only in flower or fruit tissues, while Politis (39) described 
them in these tissues and in those of bulbs. Oil bodies of this type are 
further restricted to a few groups of flowering plants. Lists published 
by Zimmermann (51) and by Politis (39) record them in groups of 
species in the Orchidaceae, Liliaceae, Amaryllidaceae, Iridaceae and 
Malvaceae, while Beer (4) found them in one of the Compositae. 

The third type of oil body described by Lidforss (32) is character- 
ized by its homogeneous appearance, by its spherical shape and by its 
unrestricted location in the cell. It is reported from leaf tissues of 
flowering plants and is of common occurrence in this group. 

Besides these three classes of oil bodies there are isolated descriptions 
of elaioplasts that are not included in any of the types described. Such 
are the reticulate, highly refractive structures saturated with an amber- 
colored oil described by Keene (26, 27) in two molds. Such also are 
the yellow, green or black oil bodies near the nucleus found by Hierony- 
mus (22) in some algae. 

In 1888 Wakker (47) demonstrated by abnormal plasmolysis that 
the oil bodies in the monocotyledons and in the liverworts are located 
in the cytoplasm. He showed that, although these structures often 
protrude into the vacuole, they are never located in it as Pfeffer (38) 
and Rattray (42) had thought. Later investigations have substan- 
tiated Wakker's observations and extended them to include all types 
of elaioplasts. 

There is no general consensus of opinion on the structure of the non- 
homogeneous oil bodies. Pfeffer (38) described them as aggregations 
of homogeneous oil globules, a view expressed in modern times by 
Guilliermond (20), by Meyer (34) and by Kozlowsky (28). Other 
students have described a stroma with embedded oil globules. This 
view was first expressed by Wakker (47). It was elaborated upon by 
Zimmermann (51), who pointed out less refractive inclusions which 
he termed vacuoles or portions not producing oil. Later Beer (4) and 
Politis (39) described the elaioplasts in Gaillardia and in the mono- 
cotyledons as aggregations of smaller bodies, each composed of a stroma 
with included oil globules. A more elaborate structure was postulated 
by Woycicki (50) and by Keene (26, 27). Woycicki described elaio- 
plasts in Vanilla with central oily drops surrounded by a mucilaginous 
layer which in turn was covered by a granular layer. Keene described 
a somewhat similar structure in the oil bodies of Sporodinia which 
showed a denser reticulate center and a coarser reticulate outer portion. 
The presence of an unfixable stroma in the oil bodies of the liverworts in 



228 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

contrast to the tixable one in those of the monocotyledons was pointed 
out by Kiister (29). Later Gargeanne (9) and Dombray (7) attempted 
to show that this unfixable stroma was a fluid or a semi-fluid. 

The question of an enveloping membrane has been raised with refer- 
ence to the oil bodies in the liverworts. Pfeffer (38) inferred the pres- 
ence of a membrane from the characteristic ring left after treatment 
with alcohol. Kiister (29) demonstrated in 1894 that this ring is an 
artefact. Gargeanne (9) repeated the demonstration but maintained 
that, although the ring is an artefact, the bodies possess a true mem- 
brane homologous with the tonoplast of the vacuole. Later writers 
have not agreed with Gargeanne in recognizing a membrane. The pres- 
ence of a membrane about elaioplasts in the monocotyledons has been 
described only by Raciborski (41) who considered the stroma at times 
to be reduced to a surrounding layer. 

The development of the oil bodies is also a disputed point in the 
literature. Pfeffer (38), Rivett (43), Lidforss (32), Chalaud (5), 
Meyer (34) and Guilliermond et al. (20) have considered the formation 
of elaioplasts to be a process of aggregation of small drops in the cyto- 
plasm with more or less fusion. Kozlowsky (28) has further stated that 
the drops are first extruded from the chloroplasts. A second theory 
has been postulated by Wakker (47), by Kiister (29) and by Harper 
(21). They consider that a stroma apF>ears first as a shadowy, wrinkled 
structure in which refractive oil drops appear later. Gargeanne (9^ 
stated that the oil drops are secreted by a surrounding membrane, 
while Dombray (7) described the deposition of substances from the 
cytoplasm and their transformation by the cell sap as a catalyser. An- 
other theory is expressed by Hieronymus (22) and by Beer (4) who 
described elaioplasts formed by the aggregation of degenerating plastids 
with the production of oil. Somewhat similar is Woycicki's (SO) theory 
of the aggregation, partial degeneration and fusion of mitochondria-like 
bodies forming oil. Keene (26, 27) postulates the formation of a reticu- 
late structure in homogeneous bodies with the later fusion of several of 
these bodies. Still another theory by Politis (39) and by Raciborski 
(41) describes the development of elaioplasts by the growth of refrac- 
tive drops and the subsequent fusion of the bodies so formed. 

The division of elaioplasts has been noted in a few instances. Raci- 
borski (41) in 1893 described a fragmenting of the bodies after they 
had passed maturity and a breaking off of bud-like protrusions. Again 
in 1914 Politis (39) described division of the elaioplasts. Politis con- 
sidered division not merely an incidental or degeneration phenomenon, 
but a method of increasing the number of these bodies. Besides the 



1935] FAULL, ELAIOPLASTS IN IRIS 229 

budding already described by Raciborski, Politis described passive 
division of the body by the cell wall during cell division. 

The history of oil bodies after they have reached maturity has been 
studied. In the liverworts they are generally thought to remain un- 
changed even after the death of the cell, although Dombray (7) noted 
a decrease in size, fusion of the oily globules and aggregation of the oil 
bodies before death. Elaioplasts in the monocotyledons are generally 
thought to degenerate some time after reaching maturity. Wakker (47) 
described their disappearance in older tissues of Vanilla. Beer (4) 
and Woycicki (SO) described a resolution of the oil bodies into scattered 
oily spheres. Politis (39), on the other hand, described the disappear- 
ance of the oil first, leaving a vacuolated protein mass which might later 
disappear also. 

Movement has been noted in connection with elaioplasts. In 1895 
Zimmermann (51) first recorded the rotation at times of oil bodies in 
the monocotyledons, a phenomenon observed also by later investigators. 
A second type of motion consisting of Brownian movement of the 
globules within the oil bodies appears in oily structures in the hepatics. 
Gargeanne (9) described this as an injury phenomenon, but recently it 
has been noted by Dombray (7) as a normal condition in the elaioplasts 
of some species. 

The chemical composition of the elaioplasts and particularly of the 
oily portion has received much attention. The theories advanced are 
based chiefly upon microchemical reactions. Dombray (7) has inter- 
preted microscopical observations in the light of analyses of extraction 
products. Two opposing theories regarding the composition of the oil 
have been formulated. In one the oil is said to be chiefly a mixture of 
essential oils. This is the view recently expressed by Popovici (40) 
and by Rivett (4) in her description of the oil as a mixture of essential 
oils with small amounts of protein and fatty oils. Dombray (7) stated 
that the oily substance was a mixture of essential oil and "tannoides." 
The opposing theory considers the oil to be composed chiefly of fatty 
oils. This is the opinion of most investigators. Pfeffer (38) described 
the oil as a mixture of fatty oil with some water and protein and with 
traces of wax and resins. Later Kiister (29) designated the oil in the 
elaioplasts of liverworts as a fatty oil resembling castor oil. Lidforss 
(32) identified the oil in the homogeneous oil bodies of flowering plants 
as a non-drying oil containing fatty acids of the type C" H^n-i O2. The 
stroma, if present, is generally considered to be a protein, a view first 
expressed by Zimmermann (51). 

There is little agreement among investigators concerning the origin 



230 JOURNAL OF THE ARNOLD ARBORETUM [vouxvi 

and identity of oil bodies. Raciborski (41), Kiister (29) and Gavaudan 
( 10, 11, 12) have considered them to be cell systems independent of the 
vacuome, chondriome or plastidome and originating more or less de novo 
in the cytoplasm. Wakker (47) considered them to be special plastids, 
while Beer (43) and Hieronymus {22) described them as degenerating 
plastids. Kozlowsky (28) has stated that they are products of the 
chloroplasts. That they are special or transformed vacuoles has been 
postulated by Keene (26, 27), by Dombray (7), by Gargeanne (9) and 
by Rivett (43). Woycicki (50) and Lundstrom {il>) have described 
oil bodies originating from mitochondria-like bodies. Politis (39) has 
ascribed a nuclear origin to them. 

A relationship between elaioplasts in the monocotyledons and crystal 
formation has been suggested. Wakker (47), Politis (39) and Monte- 
verde (36) stated that there is no connection between the oil bodies and 
the calcium oxalate crystals found in the same plants. But Warlich 
(48) considered them to be interdependent structures, while Woycicki 
(50) in 1929 described crystals forming in some of the elaioplasts in 
Ornithogalum. 

Many writers have ventured theories on the physiological and bio- 
logical significance of the oil bodies. In general they have considered 
those in the liverworts and also the homogeneous ones in the flowering 
plants to be excretions. But those in the monocotyledons they term 
assimilation products, although Raciborski (41) stated them to be 
excretions. Various other theories have been offered. Beer (4) in 
1909 demonstrated that the bodies in Gaillardia are degeneration prod- 
ucts of the plastids with the secondary function of producing color. 
Hieronymus {22) and Lundstrom {ii) suggested that the bodies are 
protective in function, a theory opposed by Dombray (7). 

In concluding the summary of the literature on oil bodies it should 
be noted that these structures do not include the olplasma described by 
Leiner (31) and by others of Tschirch's school, nor do they include the 
oil cells described by Lehmann (30) and others. The former (olplasma) 
deals with oil in the cytoplasm — chiefly of fatty seeds. The subject is 
well summarized in the account by Tunman and Rosenthaler (46). The 
phenomenon of the appearance of oil in special oil cells involves the 
transformation of large portions of the cytoplasm or secretion from the 
modified cell wall rather than the appearance of oily bodies in the 
cytoplasm. 

In addition to the literature on oil bodies, some reference should be 
made to the literature on the structure of Iris cells. The most recent 
and complete studies are those by Guilliermond (15 20) and by Dan- 



1935] FAULL, ELAIOPLASTS IN IRIS 231 

geard (6). They have developed a method of vital observation espe- 
cially adapted to this type of study. They have pointed out the 
presence in Iris cells of vacuoles and their inclusions, of cytoplasm, of 
oil globules, of chondriosomes of various types and of plastids. In par- 
ticular Guilliermond has described the chondriosomes and plastids and 
their developmental stages. He has noted the presence of oil globules 
in most plastids and chondrioconts in Iris. These globules which he 
has found associated more often with young or degenerating types of 
plastids he considers to be lipoids separating out from the plastid sub- 
stance. He has described the development of plastids from mitochon- 
drial types differentiated from other mitochondria by their potentiality 
for plastid formation. He has described the formation of chloroplasts 
from an intermediate chondriocont stage by budding and fragmenting. 
Other phases of studies carried out on Iris include the action of hypo- 
and hypertonic solutions on chondriosomes, observations of the amoe- 
boid movements of chondrioconts, and the identification of an oily body 
in the vacuoles of certain cells as a phenol compound. 

MATERIAL 

The plants used in my studies of elaioplasts included numerous irises, 
some liverworts and a few representative flowering plants. They were 
obtained from several sources. The major part of the study was made 
on colonies of Iris versicolor and of an Iris pallida of hybrid origin 
which grew in abundance near the laboratory. For work on living 
tissues it was desirable to have the plants as close at hand as possible. 
It was also desirable to locate single colonies in a natural habitat for 
the basic study of variations. In this way differences due to season, 
development, etc., were less likely to be confused with those due to 
location, to abnormal habitat or to individual variations. 

As a rule the material was used as soon as it was collected. But in 
some instances it was kept in water or in wet sand in the greenhouse for 
later observations, or it was transplanted to garden beds. In the early 
part of the study a few plants of /. pallida and of /. versicolor were 
transplanted to pots in the greenhouse to supplement the outdoor 
material. Although some interesting observations were made on these 
plants, they grew so poorly that this method of providing material was 
abandoned. Fortunately, it was not necessary to rely on greenhouse or 
garden material at any period. 

The Iris versicolor was taken from a swampy field at the corner of 
Weston St. on the Cambridge-Concord turnpike about an hour's drive 
from the laboratory. The Iris pallida hybrid, a garden plant, grew in 



232 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

beds within a few rods of the laboratory. Both of these species were 
sufficiently near at hand to be obtained as they were needed for 
examination. 

The other irises used as supplementary material and for a general 
survey of the genus were obtained from several places. A group of 
native West Coast species was studied in California. Three of these, 
/. macrosiphon, I. Douglasiana and /. longipetala, grew naturally within 
a few hours' drive of the laboratory. But /. Hartwegii and /. missouri- 
ensis had to be brought to Palo Alto from the eastern part of the state. 
A large number of other species were obtained from the Missouri Botani- 
cal Garden, from the Brooklyn Botanical Garden and from the New 
ssouri Garden were examined 
were brought to Boston for 

For a list of the species of Iris studied see the table on page 246. 

In addition to the Iris plants, a group of rhizomatous plants and a 
number of liverworts were obtained. The former were studied at the 
Missouri Botanical Garden for the most part, although a few were 
collected around Boston. Two species of Vanilla, the plant used by 
Wakker (47) in his classical studies on elaioplasts, were obtained from 
Panama. The liverworts were collected in the New England woods for 
study in Boston or they were sent from Oregon to the California labora- 
tory for use there. The hepatics were kept in the laboratory in moist 
glass containers over a period of weeks. 

A list of the flowering plants studied is given on page 248 and one of 
the hepatics on page 254. 

TECHNIQUE 

In choosing a method for the morphological examination of the elaio- 
plasts one fundamental requirement was kept in mind. It was desir- 
able to observe the bodies in as unaltered a condition as possible in order 
to discover their normal development and variations due to seasonal, 
environmental or specific differences in the plants examined. 

At the present time there are two methods used in the study of cyto- 
plasmic bodies. The first of these is the fixation technique introduced 
in the later decades of the nineteenth century and developed to the 
highest degree in the complicated "mitochondrial techniques" and 
"silver or osmic impregnation methods." Essentially it consists of kill- 
ing and fixing blocks of tissue in reagents that solidify proteins and fats, 
rendering them insoluble in specific fluids, and then staining sections 
differentially. Incidentally the technique involves a rather complicated 
process of embedding and one or more dehydrations. 



1935] FAULL, ELAIOPLASTS IN IRIS 233 

The other method is that of examining untreated tissue either with 
or without the aid of vital dyes. Although untreated tissue was used 
before the introduction of fixatives, it was superseded by them. Recently 
the so-called vital technique has been revived and developed, notably 
by the Dangeards and by Guilliermond in France and by Bailey in 
America. Guilliermond has described a technique for vital staining in 
his studies of the vacuome and has contributed data on various aspects 
of injury and death in his studies of the chondriome. Bailey (1) in 
his investigations of the cambium has tabulated criteria that can be 
used in distinguishing living from dying or dead cells. Bailey and 
Zirkle (2, 3) have clarified the vital staining technique by their investi- 
gation of the toxieity of a large number of dyes, of the most suitable 
media in which to use the stains, of the staining properties of different 
dyes and of the varying reaction of vacuoles to given stains. 

Both of these methods were tried in the study of Iris, but that of fixa- 
tion was eventually discarded because of the difficulties involved. The 
vacuoles in the rhizome were found to contain large quantities of a sub- 
stance that precipitated with fixatives and stained deeply, obscuring 
the sections, while the elaioplasts in the rhizomes of Iris versicolor con- 
tained quantities of "oil" that either was dissolved or was extruded in 
large masses obscuring the cell structure. In the one or two instances 
where this did not occur, a good fixation was obtained in mature but not 
in meristematic cells. The fixation images in sections of rhizome meri- 
stem were not comparable with those obtained in root-tip meristems, nor 
could they, as in the case of the root-tips, be identified with structures 
clearly seen in similar "living" cells. A third difficulty, that might in 
time have been overcome, lay in the persistent plasmolysis of cells in 
the rhizome meristem and in leaf tissue. For these reasons it was felt 
that the fixed material did not give an image of unaltered cells, nor could 
it be relied upon for comparative work. Better results were obtained 
with the "vital" technique where dead and dying cells could be observed 
and where those that survived for some hours without undergoing lethal 
changes seemed to present a more reliable picture of an unaltered condi- 
tion. Consequently after some months of unsuccessful experimenting 
with fixatives and dehydrating reagents and with different hydrogen 
ion concentrations of single fixatives, the method was entirely aban- 
doned and the "vital" technique alone retained. 

Although fixation methods were finally discarded, it should be noted 
that in certain instances satisfactory results were obtained in this way. 
Thus the mitochondrial fixatives and stains proved successful for root- 
tips where they apparently produced little or no alteration in the cell 



234 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

structure. Likewise, since chromic and osmic acids fixed the elaioplast 
"oil," occasional slides were obtained of mature rhizome tissue quite 
comparable with that examined "vitally." Other fixations, although 
they did not give exact images of the cytoplasmic contents of the cell, 
proved useful in determining the structure of the oil bodies. The fixa- 
tives that proved most successful for the occasional rhizome slides were 
0.5% osmic acid solution and Flemming's weak solution followed by 
Flemming's triple stain. The most satisfactory of the mitochondrial 
fixatives was ammonium Erliki solution (25 cc. each of l/r solutions 
of ammonium and potassium bichromates plus 25 cc. of an 8% solu- 
tion of formaldehyde) followed by Milovidov's modification' of Vol- 
konsky's stain. With these fixatives the usual dehydrating and paraffin 
embedding schedules were satisfactory. A third instance of useful 
fixation was found for the mitochondrial fixatives. These, although not 
entirely successful except for root-tips, did fix mitochondria throughout 
the plant sufficiently well for a rough survey of the distribution of these 
elements. 

The "vital" method was preferred and finally used exclusively because 
it presented a more reliable picture of unaltered cell structures. 
Although this was the main consideration, there were other factors that 
made the "vital" technique especially favorable for the study of devel- 
opmental and other changes within the cell. Of primary importance was 
the possibility of observing fluctuating changes of a moment's dura- 
tion, as well as those more permanent ones associated with age or season. 
This was possible only with a technique which left the more or less fluid 
contents of the cell unchanged. The "vital" method provided such a 
technique. Another factor favoring the "vital" method was its practical 
simplicity. Although some skill was required in sectioning, after this 
was obtained the actual preparation required but a few seconds. Not 
only was this a saving of time but it was possible to examine material 
as it was brought in, a method that enabled one to proceed quickly with 
the study. A third factor of importance was the applicability of the 
method without modification to all kinds of material. In a comparative 
study of tissues and plants this was an essential requirement for the 
technique. 

As used in this study the "vital" technique was essentially that de- 
veloped by Bailey (1) for the study of cambium. The material was 



1935] FAULL, ELAIOPLASTS IN IRIS 235 

sectioned, placed in appropriate solutions and examined immediately 
and at intervals. For distinguishing the living from injured or dead 
cells criteria were established based upon comparisons between obvi- 
ously injured cells and those that survived for some hours before show- 
ing signs of injury. The only differences in the technique for Iris lay 
in the details of sectioning and of preparing solutions and in the possi- 
bility of more firmly establishing criteria for living cells by comparisons 
with mounts of thin, unsectioned tissue. 

The sectioning was done with a "Gem" razor blade freehand, or, for 
some rhizomes, with a Thomson-Spence sliding microtome. Although 
the microtome sections were more uniform in thickness and more con- 
venient for mature rhizomes, they were less satisfactory with the other 
tissues. Apparently a thinner blade produced less injury in rhizome 
Awhile it was the simplest means for sectioning leaf, flower or 
The razor blade was used for mature rhizomes also when a 
was not. available. In either case, sections were obtained 
varying from one to several cells in thickness. Measured by the micro- 
tome, sections of mature rhizome varied from 30 n to 50 p or more, 
while those of the smaller-celled meristem were 15 p to 20 [.i or less. 

The solutions in which these sections were immersed consisted of a 
basic solution plus one or more of the "vital" dyes, or merely of the 
basic solution alone. Of the three fluids tried, water, nujol and sucrose 
solution, the sucrose solution in a five to ten per cent concentration, 
proved most satisfactory. 

The dyes most commonly used were Neutral Red, Janus Green BB, 
Chrysoidin Y and Benzene-azo-alpha-naphthylamine. Although Chry- 
soidin Y is the only one of these dyes which stains the elaioplasts, the 
light staining of the vacuole with Neutral Red throws the cytoplasm 
into relief and makes its structures more clearly visible. The other 
dyes in combination with Neutral Red and Chrysoidin Y have a clari- 
fying effect. None of these dyes stain the immature oil bodies, while 
the staining of the mature oil bodies by Chrysoidin Y is but temporary. 
Almost all dyes will stain dead, mature oil bodies. In practice, only 
traces of the dyes were used (one drop of a concentrated aqueous solu- 
tion to 25 cc. of sugar solution). Staining is better and more rapid when 
the sucrose solution is made alkaline with Clark's buffers (pH 8.2 to 8.6) 
which shorten the staining period from an hour or more to fifteen minutes 
or less. Since most stains, even in small amounts, are toxic after a time, 
sections that it was desired to keep were removed to pure sucrose solu- 
tions. In this way cells were kept "living" for twelve hours or more. 

An essential part of the technique was the establishment of criteria 



236 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

for distinguishing living from dead or dying cells. By comparing 
obviously injured cells with those which survive sectioning for some 
hours without sign of injury, such criteria have been established for 
cambial tissue. By the same method criteria were found for Iris cells. 
In addition unsectioned roots, bracts and flower parts of Iris and the 
thin leaves of a Potamogeton were examined. Living cells in Iris, like 
those in cambium, are marked by the following characters: regular 
cyclosis, absence of Brownian movement in the cytoplasm and a stain- 
ing of the vacuole in the presence of Neutral Red. Two additional 
criteria were found for living cells of Iris, namely, a pulsation of the 
cytoplasm in isodiametric cells and the amoeboid movement of the 
chondriosomes. Both of these phenomena are essentially a swelling or 
contraction of parts of the structure involved. The pulsation, for 
example, is the swelling of one part of a protoplasmic thread at the 
expense of another, a phenomenon involved in changes in the concen- 
tration of the substance at a given point. The pulsation of the cyto- 
plasm occurs principally in isodiametric cells where there is no streaming. 
The amoeboid movement of the chondriosomes may occur in any cell. 
Both criteria proved valuable as indications of the condition of the 
cells. Dead cells of Iris, as of the cambium, show one or more of the 
following characteristics: coagulation of the protoplasm, a general 
formation of granules in the cytoplasm, staining of the nucleus and 
cytoplasm in the presence of dyes, increasing opacity of the whole cell 
and Brownian movement in the cytoplasm. Dying cells in Iris were 
found to show the following characters: jerky or irregular streaming 
and Brownian movement within the plastids. Parallel phenomena were 
found in the irregular streaming and degenerating plastids of some epi- 
dermal, bract and flower tissues. 

The validity of these criteria for distinguishing living from dying or 
dead cells should be considered. The possibility of injury lies in the 
sectioning, in the action of the solutions in which the sections are placed, 
in the pressure of the cover glass used in mounting sections and in the 
strong light used for microscopic observations. In establishing criteria, 
the use of unsectioned material eliminated the possibility of injury due 
to sectioning, while the examination of water plants in the water of 
their natural habitat provided a check upon the effects of the solutions 
used in the study of Iris. A similar check upon the effects of pressure 
from the cover glass was provided by removing it. The possibility of 
injury due to strong light alone remains. That strong light will pro- 
duce injury and death is clear, but the effects are slow in appearing and, 
if the light is removed in time, they are temporary. They can be taken 



1935] FAULL, ELAIOPLASTS IN IRIS 237 

into account in establishing criteria for distinguishing living from dying 
or dead cells. That there are undetected, instantaneous changes is im- 
probable, for the reactions in plants are in general slow. The effect of 
the light appears chiefly in the slowing down of streaming, and, if 
exposure is continued, unmistakable signs of death such as coagulation 
of the cytoplasm finally are observed. 

It should be noted that the observation of minute details of cyto- 
plasmic structures can be carried on only with the aid of the best high- 
powered microscopic equipment. For the observation of sections 
mounted in aqueous media a water immersion objective is essential. 
Without such equipment, many of the details of structure described in 
the following section cannot be seen. 

OBSERV7\TIONS 

Elaioplasts occur typically in the parenchyma of the rhizomes of 
Iris versicolor. They appear in every cell as granular, highly refrac- 
tive masses with a decidedly yellowish cast (Fig. 1). The individual 
elaioplasts are almost spherical in shape and seem to be composed of 
closely compacted globules approximately one micron in diameter (Fig. 
2). They are relatively constant in size within a given rhizome, gen- 
erally averaging 10 to 13 microns in diameter. Although in some 
material they may be twice this size, they are never as large as the 
nucleus which has a diameter of the order of 40 to 50 microns. Often 
a hundred or more of these elaioplasts will be found in a single cell, 
aggregated for the most part into one large mass. Sometimes there are 
as few as twenty to a cell, but often they more than half fill the cell 
lumen, obscuring the nucleus and protruding into the huge vacuole. 

All evidence shows that the elaioplasts are located in the cytoplasm. 
Although they protrude into the vacuole, protoplasmic threads are often 
observed to spread at their surface as if to include them (Fig. 1 ). Occa- 
sionally one is seen moving in the streaming protoplasm. The study of 
similar bodies in the root, where they obviously are included in the cyto- 
plasm, substantiates these observations. 

Microchemical tests indicate that the bodies are mainly lipoid in 
character. They stain brilliantly in "fat" dyes such as Sudan III, 
alkannin and nascent indophenol blue.^ They are almost completely 

vivante. Comp. Rend. Soc. Biol. 1923. — Zweibaum, J. and G. Mangenot. Appli- 
vitale et post vitale des graisses de la cellule vegetale. Comp. Rend. Soc. Biol. 1Q23. 



238 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

soluble in lipoid solvents such as ether, chloroform and carbon tetra- 
chloride. They also dissolve largely in 95 /i alcohol, a solvent for some 
oils. They are insoluble in hydrochloric acid, sulphuric acid and potas- 
sium hydroxide, although they are more or less structurally disorganized 
by these reagents. They are not volatile at 100°C., which indicates the 
presence of lipoids rather than essential oils. 

The reaction of the elaioplasts to heat and to many reagents in which 
they are insoluble is marked by the extrusion of the lipoid in drops (Fig. 
5). The reaction occurs relatively slowly so that it can be watched. A 
net-like residue remains which is not distinctly lipoid in character. The 
drops characteristically remain in contact with the net and are flattened 
on their attached side. The reaction occurs with heat, picric acid, dilute 
sulphuric acid. Gram's solution, etc. 

Injury to the cell typically produces active Brownian movement of 
the globules within the limits of the elaioplast which eventually bursts, 
freeing the globules within the cell lumen. A similar phenomenon 
occurs in elaioplasts which escape from cut cells. It can be induced by 
mechanical pressure. 



The structure of the elaioplasts in the mature rhizome is that of a 
matrix with embedded globules. This is best shown in sections of 
fixed material, for the globules in fresh material are so refractive and 
so closely packed that it is difficult to distinguish any structure clearly. 
With osmic acid and some of the chrom-osmic fixatives the globules are 
preserved in situ (Fig. 3b). They clearly show a network of a differ- 
ent substance between them. With other fixatives the globules are 
never preserved, but a net-like structure with lacunae of the approxi- 
mate size of the globules remains (Fig. 3a). This can be seen best by 
the use of mitochondrial or plastid fixatives such as ammonium Erliki 
and an appropriate stain. It is well shown, too, by Wakker's method 
for double staining elaioplasts with anilin blue and alkannin after fixa- 
tion of sections in picric acid. In this case, the extruded globules are 
stained red and the matrix appears as a purple network with blue 

The behavior of the bodies in fresh material supports the observa- 
tions on their structure as seen after fixation. The globules show no 
tendency to fuse, a fact which indicates a separation by the presence of 
at least a surface film. In injured material, they move apparently un- 
changed in a liquid portion of the intact body. Further proof of a 
matrix is found in developmental forms and in homologous oil-bearing 



1935] FAULL, ELAIOPLASTS IN IRIS 239 

bodies in other species. Here the matrix is often so abundant as to be 
clearly visible in untreated material. Such is the case in very young 
cells of the rhizome, in some cells of the root-tip, in rhizomes of Iris 
pallida and of Iris Hartwegii, etc. The matrix is also clearly shown in 
the root during the degeneration of elaioplasts. Here before death the 
refractive globules disappear leaving only a net with lacunae. This net 
is very similar in structural appearance, although not in shape, to the 
net-like image of rhizome elaioplasts in fixed material. 

The globules were identified as the material which gives the elaioplast 
as a whole its lipoid characters. They show the reactions previously 
described for the elaioplasts and additional ones equally characteristic 
of lipoids. They stain in the "fat" dyes. This is apparent in intact 
bodies, but it is more clearly seen with the moving globules in disinte- 
grating ones.. They are highly refractive, a property seen in both intact 
and disintegrated elaioplasts. They disappear from sections treated 
with "fat" solvents such as carbon tetrachloride, ether, etc., but they 
may be preserved in sections treated with "fat" fixatives such as osmic 
acid and chrom-osmic mixtures. They are completely soluble in 
alcohol. This was demonstrated with globules in suspension in alkaline 
water. Upon the addition of 95 7o alcohol a homogeneous fluid resulted 
indicating the complete solution of the globules. 

The matrix was shown to be of a different substance from the globules. 
It appears to be more like the cytoplasm in composition. Unlike the 
globules it requires no special fixative for its preservation. At least a 
portion of it is insoluble in alcohol and lipoid solvents such as carbon 
tetrachloride, for it sometimes remains intact after the use of these 
reagents. It is not refractive, for this character can be seen in young 
tissue and in injured cells to be a property of the globules only. Nor 
is it stained to any extent by the "fat" dyes such as Sudan III, etc. This 
is evident in elaioplasts with globules in Brownian movement where the 
stain is largely confined to the globules. That the matrix is of a plasma 
substance was suggested by the difficulty of staining it differentially 
from the protoplasm. This view was substantiated later by the identi- 
fication of the elaioplasts with the plastidome and chondriome. 

No evidence of a differentiated membrane about the elaioplasts 
could be found. None could be seen in fresh material, nor has any been 
brought out by reagents or fixation techniques. The only observation 
that might be interpreted as indicative of a membrane was the "burst- 
ing" of injured elaioplasts already described. But no fragments of 
membrane remained. It is more probable that the sudden freeing of 
the globules depended upon changes in the matrix which made it 
miscible with the surrounding medium. 



240 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

Seasonal Variations in Elaioplasts in Iris versicolor 

Certain variations in the form and structure of the elaioplasts are due 
to the seasonal appearance of starch. In New England the elaioplasts 
are without starch from early November through June. By July or 
August the starch begins to appear, while by September or early October 
a maximum development has been reached. The disappearance of the 
starch then begins and proceeds rapidly. By early November no traces 
of it can be found. 

The starch can be identified with Gram's solution and polarized light. 
In the former the grains stain a blue to a bluish-black, a reaction typical 
of starch in the presence of iodine. In polarized light they appear as 
bright grains with a black maltese cross on each. 

The type of starch formation in rhizomes of Iris versicolor is charac- 
teristic and constant. Each elaioplast develops several included grains 
(Figs. 4b and c). Counts made in early October showed commonly 
from 8 to 12 grains, with a recorded range of 1 to 16 per elaioplast. 
Although the grains are always grouped more or less centrally within 
the globule-filled portion, they form bulges in the otherwise rounded 
contour of each elaioplast (Fig. 4). The individual starch grains are 
approximately isodiametric. They show the central hilum character- 
istic of this shape of grain when it is included within the plastid. In 
size they are small, generally 6 to 7 microns in diameter as measured 
in material collected in early October. 

Climatic differences in the disappearance of the starch from the grow- 
ing point of the rhizome are indicated. In material from the vicinity of 
Boston and of New York the starch disappears completely in the winter. 
But in plants grown in the Missouri Botanical Garden it may be found 
about the growing point in March, although completely absent from the 
rest of the rhizome. 

The disappearance of the refractive globules of the elaioplasts has 
not been observed. Numerous observations have been made from Sep- 
tember to May, during which time they remain in abundance. They 
are likewise present in the rhizome during June, July and August, 
although a less thorough study has been made of their behavior during 
those months. 



By tracing back to the meristem, the elaioplasts in the rhizome were 
found to develop from mitochondria-like primordia by increase in size, 
in visibility and in the number of contained globules (Fig. 19). In the 
youngest cells there are small, irregular, shadowy proplastids with two 
or three included non-refractive globules. In increasingly older cells 



1935] FAULL, ELAIOPLASTS IN IRIS 241 

these bodies become more distinct and larger with a greater number of 
included globules. At the same time the globules become refractive 
and the whole body even more irregular in contour. Later with further 
increase in size and in the number of included globules, the irregular 
contour is lost. The cells then contain the granular, smoothly rounded, 
mature elaioplasts characteristic of the rhizome. 

The young elaioplasts are distinguished by the following characters. 
They have more matrix in proportion to the number of globules than 
the mature forms. They do not stain after death to any degree in 
Sudan III nor in any other anilin dyes in contrast to the brilliant stain- 
ing of the mature elaioplasts. They are restricted to a small region 
about the growing point, while the youngest stages are found only in the 
cells of the growing point. They are all irregular in contour, but this 
irregularity is emphasized in the intermediate forms which are almost 
nodulose. 

The youngest stages show characters ordinarily associated with 
mitochondria. They are about the size of Iris mitochondria, ranging 
from this up to several times their size. They are indistinctly visible 
like much of the chondriome with a peculiar fading and reappearing 
quality. Thus a period of clear definition of these shadowy forms will 
be followed by a fading and disappearing. This, in turn, after a few 
minutes or after several hours may be succeeded by another period of 
clear definition, and so on. In general, although not always, these forms 
show included non-refractive globules. This is a character shared by 
the rod-shaped mitochondria of the species. In the young elaioplasts 
there is no definite arrangement of the globules which in the rod-like 
mitochondria always form a single row. 

The formation of starch occurs in any of the young or mature forms 
of plastids. It was found during the season of its formation in all of 
them. In the young forms the starch grains ordinarily protrude from 
the globule-filled mass of the elaioplast, in contrast to the completely 
included grains of the mature elaioplast. 

No evidence of increase by division was found in mature or develop- 
mental stages. No division was seen at any time, although material 
was collected from September to June and kept under observation for 
hours at a time. In the rhizome tissue even the "dumb-bell" figures so 
often cited as evidence of division were absent. 

No evidence of degeneration was found in the rhizomes of Iris versi- 
color. Elaioplasts are found unchanged and in abundance even in the 
oldest living cells. 



242 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

Two isolated cases of degeneration of elaioplasts similar to those in 
Iris versicolor have been noted in the rhizomes of other species. One 
of these is an abnormal condition produced in a slowly dying plant. 
The second is a normal phenomenon in otherwise morphologically un- 
changed cells. It is apparently unassociated with the death of the 
cells, for no other signs of degeneration appear. This phenomenon 
occurs consistently in the cortex of the rhizomes and in the epidermis 
and sub-epidermis of the roots of Iris macrosiphon var. calijornica. 

In Iris macrosiphon, elaioplasts in the mature cells of the cortex of 
the rhizome appear as large lipoid spheres (Fig. 14c). These spheres are 
marked by their large size, by their distinct yellow color and by their 
brilliant staining in "fat" dyes, Sudan III, etc. They may be demon- 
strated to be in the cytoplasm by coagulating the surrounding proto- 
plasm with fixatives (Fig. 15). The study of developmental forms 
which can be seen to be carried in the streaming protoplasm offers fur- 
ther proof of their inclusion in the cytoplasm. 

Stages in the formation of the lipoid spheres from mature elaioplasts 
can be seen in cells not far from the growing point. The process con- 
sists of the formation of homogeneous spheres by the fusion of the 
globules and the disintegration of the matrix (Fig. 14). A single elaio- 
plast resolves itself into one or more of these spheres. In older cells still 
further fusion occurs for the spheres in them are larger and fewer. In 
these cells each sphere probably includes the substance of more than 

A similar formation of lipoid spheres can be observed in epidermis and 
sub-epidermis of the root-tip (Fig. 18). The phenomenon is identical 
with that in the rhizome, although starch is present in the root elaio- 
plasts. It shows more clearly than in the rhizome the steps in the 
resolution of the elaioplasts. The fusion of the globules proceeds for 
some time before the apparent structure of the elaioplast is lost. The 
final degeneration products include starch grains as well as lipoid spheres. 
The grains and spheres remain distinct in the cytoplasm, although the 
starch is indiscriminately scattered among the lipoid spheres. 

Proof that the formation of lipoid spheres in Iris macrosiphon is a 
degeneration phenomenon is based on two points. First, the structure 
of the elaioplast characteristic of the functioning body is lost. There is 
no evidence that the lipoid spheres can produce starch as the elaioplasts 
do, or function actively in any way. Secondly, the phenomenon occurs 
in tissue which tends to die and slough away. In the root, the epidermal 
cells in which the spheres form are short-lived. This is less evident in 
the cortex of the rhizome where the cells may live for a season or more 



19351 FAULL, ELAIOPLASTS IN IRIS 243 

after the formation of the spheres. But even in this tissue the outer 
cells die and the formation of lipoid spheres is more marked in the outer 
cells. It is not found in the inner cells of the cortex or elsewhere in the 
rhizome. 

A second case of degeneration was found in rhizome cells of Iris 
tectorum (Fig. 7). In a slowly dying plant the elaioplasts appeared 
closely compacted in each cell into one or two masses. The rounded 
contour of each elaioplast was lost, while the matrix seemed to have 
become more plastic. The identity of each elaioplast was lost in the 
mass which appeared as a single granular body with indistinct partitions 
within it (Fig. 7b). Where starch was present the grains were included 
in the composite mass. This condition has never been found in healthy 



Oil-bearing plastids are found in the rhizomes of practically all 
species of Iris. They show the same fundamental structure and devel- 
opment as those in Iris versicolor just described. But they differ from 
one another in their formation of starch. Two clearly marked types 
based on the mode and time of starch production occur. 

The iirst type is that found typically in Iris versicolor (Figs. 1, 2 and 
4). It has already been described. In contrast to the second type, it 
is marked by the disappearance of starch during the winter dormant 
season, by the formation of several starch grains in each plastid and by 
the inclusion of the starch within the plastid. Plastids of this type vary 
considerably, but they usually show at least two of the general characters. 
In some species of Iris the starch persists more or less throughout the 
winter; in others it may persist one season and not the next, and in still 
others, such as /. versicolor, it always disappears. The inclusion of the 
starch in mature plastids, although not in the younger forms, is com- 
plete in most instances. But in some cases the starch grains tend to 
protrude slightly. This is more often the case in the cortex, although it 
may characterize the whole rhizome. An extreme case accompanied by 
an unusually reduced number of lipoid globules in the plastids (Fig. 
12d) was found in one of two collections of Iris Hartwcgii. 

The second type is characterized by the persistence of the starch 
through the dormant season, by the formation of one, large, asymmetric 
starch grain or sometimes two in each plastid and by a conspicuous pro- 
trusion of the grain from the globule-filled portion of the plastid (Fig. 9). 
Caplike elaioplasts attached to one end or side of the large starch grains 
are typical of these plastids. Often the lipoid globules are larger than 



244 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

in the plastids of /. versicolor, while the matrix is abundant enough to 
be clearly seen between them. A typical example of this type of plastid 
is found in rhizomes of Iris pallida (Fig. 8). 

Although the disappearance of starch is not general in this second 
type, it has been noted in one or two instances. The starch disappears 
from the main part of the rhizome of Iris pumila during flowering (Fig. 
11), although it persists around the growing point. The solution of the 
starch leaves peculiarly cup-shaped elaioplasts (Fig. lie). A second 
case of the disappearance of starch may occur under abnormal condi- 
tions. It was induced in the rhizomes of Iris pallida placed in the green- 
house during the winter. It is accompanied by a lack of vigor and the 
disappearance from the rhizome cell vacuoles of substances ordinarily 
present at that season. The change in the vacuoles is apparent in fixed 
material in a lack of the precipitate characteristically produced in them 
by reagents during the winter. The plastids in the rhizomes of the 
greenhouse plants resembled the spherical ones of winter material of 
Iris versicolor except for their smaller size and fewer numbers. 

The distribution within the genus of the two types of rhizome plastids 
has been found to follow closely the recognized taxonomic grouping. The 
homogeneous and closely related groups show the same type of plastid, 
while a heterogeneous group such as the Apogon shows both types. In 
the latter case aberrancies from the prevailing type in the group are 
often correlated with anomalous taxonomic characters. Sufficient 
material has been examined to show definitely the condition in the two 
largest groups, Pogoniris and Apogon, and in several of the smaller 
groups, Evansia, Regelia and the Pardanthopsis and Gynandiris species. 
An indication of the prevailing type in each of the other groups may be 
found in the notes made on a few representative species. 

Rhizomes have been examined largely during the late winter. In 
late winter the pallida type of plastid shows its characters clearly, while 
the versicolor type is generally without starch at that season. An indi- 
cation of the mode of starch formation in the latter type of species can 
often be obtained from the persistent starch in the plastids about the 
growing point. In this study, such observations have been supple- 
mented by notes made during the starch-forming season. 

The pallida type of rhizome plastid occurs characteristically in Pogon- 
iris, Regelia and Oncocyclus. These are homogeneous groups which 
together with Pseudoregelia form a unit of closely related species. The 
same type has been found in a Juno Iris and in a Xyphium Iris. In the 
latter case it occurs only in tissue about the vascular bundles but not in 
the large parenchyma cells which are filled with starch. It is also found 



1935] FAULL, ELAIOPLASTS IN IRIS 245 

in Pardanthopsis, in the closely related hexagona sub-group of the 
Apogons, in the anomalous Apogon, /. verna, and in one of the variable 
Apogon spuria group, /. spuria ochroleuca. 

The versicolor type of plastid appears in the Evansia group and in the 
majority of the Apogons. In the latter it characterizes the following 
sub-groups: Sibirica, Laevigata, Longipetala, Californian, Tripetalous, 
Spuria and Ensata. Its distribution is more limited than that of the 
pallida type, for the Evansia and Apogon sub-groups include but one- 
third of the species. The pallida type appears to characterize the other 
two-thirds of the genus. Since the Evansia and Apogon sections include 
all of the American irises, the versicolor type is predominantly the type 
found in North American species. The only exceptions are the anoma- 
lous Apogons cited in the preceding paragraph and one to be described 
later. A similar predominance of the versicolor type is to be found in 
the Asiatic species. The American and Asiatic species contrast in this 
respect with the more strictly European and Mediterranean species, 
which belong chiefly to groups showing a pallida type of plastid, notably 
Pogoniris and related groups and the bulbous forms. 

The absence of oil-bearing plastids has been noted in five irises. In 
these cases the rhizome cells are filled with starch. The starch is of 
two types paralleling in distinguishing characters and in distribution the 
two kinds formed by the oil-bearing plastids. One is present as large, 
single, asymmetrical grains similar to those in the pallida type of oil- 
bearing plastid. They characterize the anomalous Apogon, Iris ungui- 
cularis, the Reticulata Iris and Gynandiris. The last of these is not 
always included in the genus; in the formation of starch and no lipoid 
in its corms it resembles the closely related genus, Moraca. The 
Reticulata is a group closely related to the Xyphium, which shows simi- 
lar starch grains and a few oil-bearing plastids of the pallida type. The 
second starch grain type resembles the starch grains of the versicolor 
plastids in their small size, in their isodiametric shape and in their forma- 
tion in groups within a single leucoplast. Like the versicolor type of 
plastid they are found in Apogon Irises, the Japanese Iris and Iris Sin- 
tenisii. The former is a hybrid of /. laevigata, one of a group charac- 
terized by the versicolor type of plastid, and another member of the 
same group. The other Iris belongs to the Spuria sub-section, a group 
of intergrading and variable forms, for which no single characteristic 
type of plastid was found. 

For the type of plastid found in individual species, the reader is 
referred to the table on page 246 and also Figs. 12 and 13. The table also 
includes data on the material, its source, the season of examination, etc. 



T.ABLE OF IRISES E> 


AMI.XKD SHOWIN-G THE TYPE 
Section Group 


OF RHIZOME PLASTID AX 


) THE SOUK 


Co'l'lected 


.ekPl.^I 33, 


I. pallida \.^m.X-^ 
I. pallida variety 
I.Cengialti\mh. 

X/. "Zz.an«m6«r«" Hort. 
/. Xyphium L. 

X/.Aexa«.«aWalt.X? 
/. w«tco/or Small 

I. 'spuria Vz\\.v^.r. ochrokuca 
I. setosa Pall. 

/. Hartwegii Baker 

/! TenaxDingl 
I. longipelala Herb. 


Pogonlris 

Pogoniris 

Pogoniris 

Pogoniris 

Pogoniris 

Regelia 

Regelia 

Oncocyclus 

Oncocyclus 

Juno 

Xyphium 

Ai^gon ""''' 

Apogon 

Apogon 
Al^gon 
Apogon 


Pallida 
Pallida 
Pumila 

Hexagona 

Spuril 

En"aU 
Tripetalous 

Californian 
Californian 
Longipetala 




Bussey Garden 
Mo. Bot. Gard. 

Mo." Bol: Garl' 

Brooklyn Gard. 
Brooklyn Gard. 

MrBot" Gard.' 
Brooklyn Gard. 
f Florist 

Mo.' Bot! gZa. 
(Brooklyn Gard'. 
Brooklyn Gard. 

Brooklyn Gard. 
Brooklyn Gard. 

Brooklyn Gard' 
fBrooklyn Gard. 

/California 

\ Brooklyn Gard. 

California 

Brooklyn Gard. 

Califolnia 


bulb 
bulb 


Aprif 
April 

April 

March 

April 
April 

Aug. 

Sept. 
July 


Qa 

Qf 

Pb 

lb 
Ee 

Db 



TABLE {Continued) 



Species 


Section 


Group 


Type 


Source of Material 


Parts 


Collected 


Fig 13 


I.uninicaL. 


Apogon 


Laevigata 




Mo. Bot. Gard. 


^ , 


March 


Bb 




















Apogon 


Laevigata 




iBo^to^n"'' ^'"''*' 








XI. robusta E.Anders. 


Apogon 


L.cvi...a 




Mo. Bot. Gard. 


V 


March 


Be 


I. pseudacorus L. 


Apogon 


Laevigata 




iMrBoLGard. 


r'l 


March 




/. Kaempferi Sieb. 


Apogon 


Laevigata 




Brooklyn Gard'. 




April 




XI. Wilsoni might X? 








Brooklyn Gard. 










Apogon 


Sibirica 




fDuxbu°ry. mLs. 


r'l 


March 


Ae 
Ae 


I. prismatica Pursh 


Apogon 


Sibirica 




\N. Y. Bot. Gard. 


'; 


April 








Sibirica 




Brooklyn Gard. 




April 




XI. -Quesr Hort.' 








Mo. Bot. Gard. 








/. Clarkei Baker 




Sibirica 












/. ckrysographes Dykes 








Brooklyn Gard. 




April 






Evansia 






fN. Carolina 


r'l 




Ja 


7. gracilipes A. Gray 








N.°Y. Bot. Gard. 




April 


Jb 




Evansia 






\Mo^Bot°G?rd.'^' 






Jc 


I. lacustris Nutt. 


Evansia 






Brooklyn Gard. 




April 


Je 


I.japonicaThunh. 








Brooklyn Gard. 




April 


J'i 


I. unguicularis Poir. 


Apogon 






Brooklyn Gard. 




April 
















April 




/. sisyrinchium L. 


Gynandiris 






Brolklyn'ciard.'^' 








/.5,«/««m»Janka 








Brooklyn Gard. 




April 




/. laevigata Fisch. X? 


Apogon 


Laevigata 






■■ 




Be 






248 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

Oil-bearing Plastids in Rhizomes, Bulbs etc. of other Plants 
No oil-bearing plastids have been found in any other rhizomes or 
bulbs examined. None are present in either of the species of Moraea 
examined, a closely related genus replacing Iris in the southern hemi- 
sphere. Nor are there any in the many Araceae, Bromeliaceae, Com- 
melinaceae, Liliaceae and Scitamineae examined. Rather, all of these 
plants contain large asymmetric starch grains in their storage organs. 

The chloroplasts in all of these plants characteristically contain more 
or less refractive granules. In general, such appears to be the condi- 
tion in all of the monocotyledons and in many of the dicotyledons. 
Indeed it seems to be true even of some of the lower forms such as the 
liverworts and mosses, although in these the granules are often not 

The following is a list of the species of monocotyledons examined. 

The species are grouped according to families. 

Araceae: Acorus Calamus L., Aglaonema sp., Arisaema triphyllum (L.) 
Schott, Diefenbackia sp., Nephthytis sp., PhUodendron Selloum 
C. Koch, PhUodendron cordatum Kunth, Schismatoglottis crispata 
Hook, f., Schismatoglottis rupestris Zoll. and Mor., Spathiphyllum 
sp. 

Bromeliaceae: Ananas macrodontes E. Morr., Billbergia sp., Cryptan- 

Commelinaceae: Palisota sp. 

Iridaceae: Moraea iridioides L., Moraea sp. 

Liliaceae: Allium sp., Hemerocallis sp., Ornitkogalum umbcUatum L., 

Yucca filamentosa L. 
Orchidaceae: Vanilla planijolia Andr., Vanilla pompona Schiede. 
Musaceae: Strelitzia sp. 

Zingiberaceae: Alpinia nutans Rose, Amonum sp., Hedychium sp. 
Marantaceae: Calathea sp. 

The observations in this section apply to any species of Iris unless 
otherwise stated. A careful study has been made of the conditions in 
Iris pallida and in Iris versicolor. Additional notes have been made on 
other species. 

The formation of oil globules is characteristic of plastids throughout 
the tissues of plants of the genus Iris. The globules are not always so 
numerous as those in the rhizome plastids of Iris versicolor where they 
are developed to an unusual degree. An extreme example of a limited 
formation of globules is found in the chloroplasts of the guard cells where 



1935] FAULL, ELAIOPLASTS IN IRIS 249 

the matrix of the plastids is relatively abundant and clearly visible. Nor 
are the lipoid globules usually the only observable product of the plas- 
tids. Ordinarily starch is also present, while in some plastids chloro- 
phyll or a yellow pigment is formed. 

The elaioplast condition described for rhizome plastids of the versi- 
color type may occur in any of the uncolored tissues. It is dependent 
upon the absence of starch and pigment and upon a large production of 
oil. Such conditions are found at times in the rhizome, in the root and 
in uncolored leaf and flower tissues. 

In the rhizome and root elaioplasts occur generally throughout the 
tissues of these organs. They are restricted to certain species and, at 
least for the rhizome, to certain seasons. There is no connection between 
their presence in the rhizome of a species and their appearance in the 
root of the same species. For example elaioplasts were found in the 
rhizome of Iris versicolor but not in the root (Figs. 2 and 35). On the 
other hand, they were found in the root of Iris pallida but not normally 
in the rhizome (Figs. 9 and 30). An example of their formation as a 
seasonal phase of the leucoplasts in the rhizome has already been de- 
scribed for Iris versicolor. Whether or not they also form a seasonal 
phase for leucoplasts in the root has not been investigated. 

In the leaf and flower the elaioplasts are restricted to a few cells. Often 
they are but transitional forms appearing for a very brief time. Such is 
the case in the flower where they may occur in the course of the develop- 
ment of the chroma tophores. Because of their limited occurrence in a 
few cells it is usually easy to identify them with the leucoplasts or 
chromatophores in neighboring cells. In these tissues they do not de- 
velop the brownish color so characteristic of the rhizome elaioplasts in 
Iris versicolor. Instead they remain entirely colorless. 

The development of elaioplasts can be induced under unfavorable 
conditions. An example of this has already been cited in their formation 
in rhizomes of Iris pallida grown in the greenhouse (p. 244). In this 
case they were formed by the dissolution of the starch leaving only the 
oil-bearing plastid. By growing plants in semi-darkness chloroplasts 
can be prevented from forming pigment or starch. They then appear 
as elaioplasts. In neither of these cases is an increase in the number of 
oil globules involved. Nor has the formation of unusual numbers of 
elaioplasts been observed as a result of abnormal conditions. 

The oil-bearing plastids in other parts of the plant show essentially 
the same features as those described for the rhizome. They differ from 
those in the rhizome in minor characters, also in.the absence of a general 
elaioplast phase except in the root and in the formation of pigments. In 



250 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

addition they show in some cases chondriome types as an intermediate 
stage in their development from the proplastids. In some tissues, not- 
ably in the flower and in epidermal tissues, the mature plastids often 
show further changes involving chondriome types not found in the 
rhizome. These points will be taken up separately in the succeeding 
paragraphs. 

Minor differences between the plastids in other parts of the plant and 
the type found in the rhizome and root appear in the lesser production 
of lipoid globules and in their complete lack of color in colorless tissue. 
Correlated with the smaller number of globules is a greater stability. 
This is shown in their greater resistance to injury by mechanical 
pressure and to distortion or destruction by reagents. Their lack of 
color when not pigmented can be seen in colorless leaf tissue in marked 
contrast to the strongly yellowish cast in the equally unpigmented rhi- 
zome plastids. This is particularly well shown in Iris versicolor. 

The formation of pigment, chlorophyll or yellow pigment, occurs 
ordinarily in the young plastids. But there is no specific stage at which 
it is developed. In the leaf chloroplasts it often forms shortly after the 
appearance of starch in the young plastids, although it may not develop 
for some time. In the chromatophores of the root it sometimes appears 
before the formation of starch, for example in the rootcap of Iris versi- 
color. At other times yellow pigment appears in plastids which do not 
form starch, for example in the chromatophores of roots of Iris pallida. 
In many cases yellow pigment is found in chondriocont types of plastids, 
but its formation is quite unconnected with the phenomena producing 
these forms. Proof of this is found in its formation in the approximately 
spherical plastids of the root-tip before they pass into a chondriocont 
state and in those of the rootcap of Iris versicolor where the mature 
plastids retain a more or less spherical state. 

The location of pigment in the refractive globules and also in the 
matrix and its greater solubility in the former was demonstrated. In 
the guard cells of Iris pallida, where there is little chlorophyll, the green 
color can be seen to be confined to the globules, while the matrix remains 
colorless. That it may also be dissolved in the matrix is shown in degen- 
erating chromatophores of the root where color remains in the matrix 
after the disappearance of the refractive globules. 

Intermediate developmental forms of the plastids are found in the 
root-tip (Figs. 20 24). They differ from the small plastids in other 
differentiating tissue by the retention for a longer period of the shadowy 
visibility of the proplastids and by a plasticity amounting in the younger 
stages to an almost fluid character. They resemble other young plastids 



1935] FAULL, ELAIOPLASTS IN IRIS 251 

in their origin in the proplastids, in their development by increase in 
size, in visibility, in the number of included globules and in their final 
development in many cells into the same type of plastid. In their often 
elongated shape they resemble the chondrioconts of many authors. 

The shadowy character of the younger intermediate forms is evident 
in the p)eculiar fading and reappearing already described for the proplas- 
tids (p. 241 ). With the differentiation of the tissue this shadowy quality 
is lost (Figs. 20-24), but the bodies do not become refractive until a late 
stage (Figs. 24 and 25). Often the more or less indistinct forms persist 
for long periods. 

The plastic quality of the intermediate forms is shown in their more 
or less elongated shape and in their movement in the streaming proto- 
plasm. The movement consists of a continuous changing of form (Figs. 
20 23 ) . Both movement and elongation are more marked in the younger 
stages, some of which are almost fluid. In older cells the plastids be- 
come less and less elongated with increasing viscosity until they are more 
or less spindle- or tadpole-shaped. At the same time the motion of the 
plastid becomes reduced to a moving about of the ends. In the fully 
differentiated plastid the shape is roughly spherical and there is no 
movement. Often the plastids remain in the spindle- or tadpole-stage 

The continuous motion of the intermediate types is essentially an 
amoeboid movement of the plastid (Figs. 20-23). This appears to 
some degree in all of the intermediate types. In its most exaggerated 
expression in the youngest stages, it consists of a change in form from 
a filament, through intermediate stages, to a sphere. Another example 
characteristic of the plastids before the globules have become refractive 
is the formation of two swollen ends connected by a thread. In some 
cases the thread becomes invisible, but it always reappears and shortens 
to reunite the two ends. In its least pronounced form in the older 
spindle- and tadpole-shaped forms, the movement is confined to a turn- 
ing from one side to another of the tapered ends. 

That the movement is. not wholly connected with cyclosis, although 
probably aggravated by it, appears likely. In cells where there is no 
cyclosis, the intermediate forms customarily show a pulsating movement 
associated with changes in thickness. An example of this is seen in 
young plastids in the isodiametric cells of the rootcap. 

It is worthy of note that in none of these forms has division of the 
plastids been seen. Many observations have been made at different 
times and over periods of an hour or more. But even plastids which 
appear to be divided are seen shortly to be connected by a thread which 
after a time thickens to reunite the two parts. 



252 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

The liquid character of the youngest forms is shown in the movement 
of the globules within the plastid. This consists of a sloshing about of 
the globules. In older forms this movement does not appear. Any 
rearrangement of the globules in them is due to the amoeboid movements 
of the plastid. 

The intermediate plastid types develop into leucoplasts in the cortex 
of the root or into chromatophores in the rootcap. But in other regions 
of the root and in the elongated cells of the fibrovascular bundles 
throughout the plant they persist as chondriocont types. The shadowy, 
very plastic forms are found in the central cells of the root. In other 
parts of the central cylinder more differentiated forms are found. Simi- 
lar ones appear in the fibrovascular bundles throughout the plant. In 
the inner cortex the tadpole- and spindle-shaped forms with refractive 
globules often remain. The chromatophores of the rootcap often retain 
their chondriocont-like shape and plasticity after the formation of pig- 
ment and oil (Fig. 29b). 

Associated with the persistent developmental forms are shadowy 
leucoplasts and mitochondrial types not ordinarily linked with plastids. 
In Iris, the latter are marked by their gradation into the plastid forms. 
In the same cells with the persistent chondriome-like plastids, they 
appear as long filaments with a single row of globules and a twisting or 
wavy motion in the streaming protoplasm (Figs. 26-27). Some of the 
filaments are shadowy, while the granules in others are refractive. The 
filaments with refractive granules sometimes show thickened ends 
which contain more than one row of granules. 

It may be noted here that in addition to the proplastids and fila- 
mentous chondriosomes, the more usual types of mitochondria, that is 
globular or rod-shaped forms, are found in Iris. They appear in all 
cells but are concentrated about the apical meristems and in tissues of the 
leaf and root. They are less evident in the cells of the rhizome. Two 
observations are worth recording. The so-called spherical mitochondria 
were observed to be more or less fluctuating in form and only approxi- 
mately spherical. The rod-shaped ones were seen to contain globules. 

Changes in plastids after "maturity" are found in epidermal cells and 
in the flower. The changes in both cases are marked by an increasing 
fluidity of the matrix and by the disappearance of the refractive globules. 
In the epidermis of the root and in the flower the changes culminate in 
the death of the cell. But in the leaf epidermis the plastids remain as 
more or less shade wy chondrioconts (Fig. 37). These are quite similar 
in appearance and in motion to the developmental types in the root. 

The changes in the plastids in the epidermal cells of the root have 



1935] FAULL, ELAIOPLASTS IN IRIS 253 

been carefully studied. The more or less spherical leucoplasts or chro- 
matophores first become somewhat elongated (Fig. 31). This is fol- 
lowed in older cells by increasing fluidity of the matrix and the gradual 
disappearance of refractive globules (Figs. 31-33). Where the plastids 
are pigmented some of the pigment remains after the globules are gone, 
but this also tends to disappear. Where these changes progress far, the 
plastids become shadowy nets much elongated in shape (Fig. 3i). In 
the streaming protoplasm, they are often partially drawn out into long 
filaments (Fig. 32). In some cases the attenuated portions are bent 
back upon the rest of the plastid so as to include a small amount of 
protoplasm (Figs. 30, 33, 34). When an oil globule is included with 
protoplasm it is often in Brownian movement. The attenuated forms 
persist until the death of the cell. 

During flowering the leucoplasts of the floral tissue were seen to 
undergo a series of changes similar to those described as occurring in the 
root. These have not been studied in detail, but the following general 
changes have been noted. The leucoplasts become pigmented and chon- 
driocont-like in shape. As the flower opens and fades the chromato- 
phores become more and more fluid. At the same time the refractive 
globules disappear. An advanced stage shows them partially drawn out 
into long filaments (Fig. 34). Unlike the chromatophores of the root 
the pigment is retained. These forms remain until the death of the 
cell. Similar changes occur in the leucoplasts of the bracts. 

In Iris Xyphium the formation of refractive bosses on the pollen 
grains from oil-bearing chromatophores has been demonstrated (Fig. 
17). In unripened anthers the pollen shows no markings except the small 
refractive dots forming a F»art of the wall structure. The grains are 
surrounded by the tapetal fluid in which are numerous oil-bearing chro- 
matophores. In the shed pollen the grains show not only the refractive 
dots but the closely appressed chromatophores which appear as granular, 
yellowish, refractive bosses. No similar observations have been made 
on pollen grains of other species. Although the latter show refractive 
spines or a network of refractive structures, these are in every case asso- 
ciated with wall formation. They are in no way connected with the 
plastidome or chondriome. 

All of the plastids described are found to show the following general 
characters. They tend to aggregate about the nucleus, a character also 
shown by mitochondria. Unless degeneration is involved they retain 
the ability of the proplastids to form the pigments and other products 
differentiating the different types of plastids. They also retain the 
ability to change from the plastid shape into a chondriocont form and 



JOURNAL OF THE ARNOLD ARBORETUM 



Of the plants recorded in the literature as forming elaioplasts the 
following have been examined: Vanilla planijolia Andr., V. Pompona 
Schiede., Marchantia polymorpha L., Luntdaria cruciata (L.) Dum., 
Pellia epiphylla (L.) Corda, Porella sp., Bazzania trilobata (L.) S. F. 
Gray, Scapania nemorosa (L.) Dum., Cephalozia sp,, Trichocolea 
tomentella (Ehrh.) Dum., Plagiochila asplenioides (L.) Dum., Lopho- 
colea heterophylla (Schrad.) Dum., two thallose species of the Junger- 
manniales from Oregon and two leafy species of the Jungermanniales 
from Oregon. 

The two classes of elaioplasts described by Pfeffer, Wakker and later 
writers were examined. These are the oil bodies characterizing the 
liverworts and those in Vanilla, a classical example of elaioplast-bearing 
monocotyledons. In both cases certain of the observations of previous 
writers have been verified and some additional notes made. 

The following observations made by earlier writers for Vanilla have 
been verified. The elaioplasts are present as highly refractive granular 
bodies near the nucleus in cells which contain also leucoplasts and 
chloroplasts. Structurally they consist of globules of refractive oil 
in a protein or plasma matrix. They are marked by their brilliant stain- 
ing in "fat" dyes and by the extrusion of large globules of oil after treat- 
ment with various reagents. 

In addition it has been noted that the elaioplasts are generally dis- 
tributed in all the cells of leaf, stem and root tissues rather than re- 
stricted to particular tissues in certain parts of the plants. 

It has also been observed that the single large elaioplasts are aggre- 
gates of smaller granular bodies (Fig. 45). The aggregation is more 
or less compact. In some cells it is difficult to distinguish the individual 
bodies, while in other cells they are but loosely grouped or freely circu- 
lating in the streaming cytoplasm (Fig. 46). In some cells the smaller 
bodies could be observed to aggregate into one or more groups from 
which individuals were carried away from time to time by the streaming 
protoplasm. 

The development of the smaller bodies from non-refractive granular 
ones can be observed in younger cells of leaf and root. In successively 
older cells the included globules gradually become more and more refrac- 
tive until the bodies assume the highly refractive condition typical of 
mature cells. In the less refractive stages the bodies seldom form com- 
pact aggregations. No specific stage has been noted in which aggrega- 
tion becomes the rule. The formation of compact groups appears 
possible at any time, although more characteristic of mature tissue. 



1935] FAULL, ELAIOPLASTS IN IRIS 255 

The rotary movement of elaioplasts described by Zimmermann and 
others as characteristic of these bodies has been shown to be an injury 
phenomenon. It is observed in cells which soon show unmistakable 
signs of injury followed by death. It is not seen in any cells which 
remain normal in appearance and actively streaming for a period of 
hours. The movement consists of rotation within a liquid vacuole. It 
is followed by Brownian movement of cytoplasmic inclusions and a 
general coagulation or disintegration of the cellular structure, that is 
by unmistakable signs of death. 

In the liverworts the following observations of earlier writers have 
been verified. Bodies included within the cytoplasm and marked by 
their refractivity, by their staining in "fat" dyes and by their solubility 
in alcohol appear generally throughout the group. They are located 
within the ring of chloroplasts, but, unlike those in Vanilla, show no 
particular affinity for the nucleus. They all characteristically leave a 
residual ring in solution with alcohol, etc. They vary in color from 
colorless to dark brown. Two or three classes are distinguishable. The 
first appears as a single large granular mass almost filling the cell lumen 
(Fig. 38). It is located in scattered cells throughout the plant body and 
is characteristic of the Marchantiales. The second and third types are 
found in the Jungermanniales which they characterize. They are smaller 
than those in the Marchantiales and are round, spindle- or disc-shaped 
in form (Figs. 40-44). They grade from a homogeneous type to a very 
granular one. Commonly there are from one to twenty in a single cell, 
located more or less characteristically in the peripheral cytoplasm. In 
this group they are not restricted to particular cells but are found in 
every cell. Unlike the bodies in Vanilla there is little or no tendency for 
them to aggregate. 

The development of the bodies has been observed in the Junger- 
manniales (Fig. 42). In the younger cells the oil-bodies appear as 
shadowy, wrinkled, granular bodies. They develop into the mature 
bodies of older cells by an increase in substance and in the refractivity 
of the granules. By the time the cells are fully mature, the bodies have 
become plump and refractive. There is no indication of a vacuolar 
origin postulated by some writers. 

In addition the following new observations were made. The homo- 
geneous type found in the Jungermanniales are sometimes seen with 
attached granular bodies (Fig. 44). These appear in the younger cells. 

The single bodies in the Marchantiales can be shown to be aggre- 
gations of smaller ones. This is apparent in younger cells where they 
are less refractive and less highly colored (Fig. 39). In older cells the 



256 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

Structure is obscured by the dark color. Likewise in older cells the 
bodies appear to be more closely compacted. 

The Brownian movement described by some as characteristic of the 
bodies in certain species has been shown to be associated with older 
bodies or with injury. It is never seen in younger tissue, even in cells 
with mature oil bodies. It appears in some of the older cells of the 
Marchantiaceae and can be induced in any cell by injury. 

DISCUSSION 
It has been shown in the preceding observations that the oil bodies 
in Iris are a phase of ordinary plastids. In studying the development 
and variations of these plastids, many interesting observations have 
been made which have a bearing upon the status of elaioplasts and upon 
various problems concerning plastids. In particular the observations 
provide further evidence of the plastid character of elaioplasts and of a 
relationship between the various types of oil bodies described in the 
literature. They also clarify our conception of the interrelationships of 
plastids and chondriosomes. 

To identify the anomalous bodies in Iris as a seasonal elaioplast phase 
of plastids adds another instance to the accumulating evidence of the 
plastid character of oil bodies. This substantiates the theories of 
Wakker (47), of Beer (4), of Hieronymus (22) and of Kozlowsky (28) 
who postulate a relationship with plastids rather than with vacuoles or 
with the nucleus. There is no evidence in any of the observations de- 
scribed in this paper of a vacuolar origin or identity. On the contrary, 
the structural, developmental and chemical similarities between vacuoles 
and oil bodies recorded by some authors were not observed in any of 
the material examined. Nor was there any evidence of a nuclear deri- 
vation of the elaioplasts, a theory based upon the similarity in the 
staining properties of the nucleolus and elaioplasts and in the aggrega- 
gation of the elaioplasts about the nucleus. Both of these phenomena 
have been found to be characteristic of plastids in general. The possi- 
bility remains that some elaioplasts may be more or less fused aggrega- 
tions of oil globules which bear no relationship to plastids. The phe- 
nomenon was not observed, but the possibility of its occurrence was 
not disproved. 

It is probable that the granular elaioplasts of the monocotyledons 
types of plastids. They show the same structure as 



1935] FAULL, ELAIOPLASTS IN IRIS 257 

that of the plastids, that is a matrix with embedded globules. That 
the stroma in the liverworts is non-fixable is not significant morpho- 
logically, although it indicates a chemical difference between the oil 
bodies in the liverworts and plastids in general. Further evidence of the 
plastid character of the granular oil bodies in the monocotyledons is 
found in their similarity in appearance and in general characters to 
those found in Iris. A comparison between the elaioplasts in Vanilla, 
as a classic example of the type found in monocotyledons, and those in 
Iris shows the following characters common to both : presence of refrac- 
tive granules, brilliant staining in "fat" dyes, extrusion of oil with picric 
acid, etc., aggregation about the nucleus, yellowish color, plastid struc- 
ture and the absence of the more usual plastid products such as starch 
and pigment. 

That the homogeneous oil bodies in the liverworts may be classed 
like the granular types as plastids is suggested. Heretofore no distinc- 
tion has been made between the two types because of the intergradation 
occurring between the two extremes. The appearance of attached 
granular portions in the younger stages of the homogeneous forms sub- 
stantiates the view that they should be classed with the granular types 
which, as has been suggested, are plastids. 

That elaioplasts are sometimes a phase of functional plastids as well 
as degenerate forms has been brought out in these studies. Heretofore 
they have been considered to be degenerate forms or secretions of plas- 
tids. In Iris they are found as functional plastids, as evidenced in the 
formation of starch and their apparently continuous presence in indi- 
vidual cells from season to season. That elaioplasts sometimes form by 
degeneration of plastids involving the production of oil has been shown 
by Beer (4). There is no evidence that they are ever secretions from 
plastids. 

It is probable that the granular elaioplasts described in the literature 
are sometimes functional plastids and sometimes degenerate forms. 
Those found in such organs as leaves, roots and bulbs or those found 
widely distributed throughout the plant as is the case in Vanilla are 
doubtless active plastids, while those restricted to the more or less 
evanescent floral tissues are probably degenerate plastids. 

The interpretation of the homogeneous oil bodies in the liverworts is 
not clear. They might be degeneration products, but they might also 
be an accumulation of normal plastid products within a plastid. 

It has also been shown in the studies of Iris macrosiphon that elaio- 
plasts of the type described by Lidforss (32) as homogeneous oily 
spheres may form by the degeneration of oil-bearing plastids. A similar 



258 JOURNAL OF THE ARNOLD ARBORETUM rvoL.xvi 

phenomenon has been described by Beer (4) as a final step in the degen- 
eration of plastids in floral tissue of Gaillardia. That the spheres de- 
scribed by Lidforss {1>2) are likewise degeneration products of plastids 
can only be surmised. It is possible that they are more or less fused 
aggregations of oil globules unconnected with plastids. 

Evidence of a relationship between the various types of elaioplasts 
described in the literature has been found in these studies. A structural 
similarity is seen between the oil bodies in the Marchantiaceae and 
those in Vanilla in that they are both aggregations of plastid-like bodies. 
I have found no record in the literature of the aggregation of these bodies 
in the Marchantiaceae, although the phenomenon was noted for elaio- 
plasts in the monocotyledons as early as 1914. In addition to this 
direct evidence of structural similarity the observations upon the devel- 
opment and variations of elaioplasts in Iris have demonstrated that 
these oily plastids show, at one time or another, the widely varying 
phenomena which have heretofore been considered distinctive of differ- 
ent types of oil bodies. It has already been pointed out that elaioplasts 
of the homogeneous type described by Lidforss (32) sometimes result as 
a degeneration product of a granular type of elaioplast. It has also been 
found in Iris that the oily plastids show at one time or another the 
following phenomena described in the literature for oil bodies: aggrega- 
tion and fusion of homogeneous oil globules, aggregation and compact- 
ing of plastid-like bodies, aggregation about the nucleus, unrestricted 
position in the cell, degeneration involving the disappearance of the oil 
and degeneration involving the formation of oily spheres. In brief the 
morphological distinctions between the various classes of oil bodies 
appear to be breaking down, while it is evident that plastids can show 
widely varying phenomena which, considered separately, might be inter- 
preted as bases for the distinction of fundamentally different types. 
Further study on this subject is highly desirable. In particular further 
observations on oil bodies and plastids in Vanilla, Ornithogalum and 
the hepatics are needed, for much of the literature deals with elaioplasts 
found in them. 

An additional point which tends to reduce the number of recorded 
distinctions between the oil bodies in the liverworts and those in the 
monocotyledons appears in the permanent character of the elaioplasts 
in the rhizomes of Iris. Heretofore elaioplasts in monocotyledons have 
been described as temporary structures, while those in the hepatics have 
been thought to be more permanent. It may be noted here that my own 
limited studies made on Vanilla indicate that elaioplasts are not the 
temporary structures even in this classical plant that one would infer 
from the literature. 



1935] FAULL, ELAIOPLASTS IN IRIS 259 

That conditions producing oil bodies are more or less restricted in 
their occurrence in the monocotyledons has again been brought out in 
these studies. Elaioplasts do not appear generally throughout the group, 
although the appearance of oil-bearing chloroplasts is not uncommon. 
This study adds another genus and many species to the published lists 
of monocotyledons in which elaioplasts occur. Although oil-bearing 
plastids occur in the rhizomes of practically all species, it is noteworthy 
that the elaioplast condition is restricted for the most part to the Apogon 
irises of Asia and America. This is the first record that I find of the 
occurrence of oil bodies in rhizomes, although Politis (39) has described 

Evidence of the function and significance of the oil bodies has been 
found. In Iris the bodies are clearly assimilative organs as shown by 
their formation of starch. That the oil itself is a reserve food supply is 
indicated. In certain species it replaces at least morphologically the 
starch stored in the rhizomes of other species. There is no evidence that 
the elaioplasts are ever excretions, although they may be at times 
degeneration products. 

There is no evidence of the division of elaioplasts recorded by a few 
writers. The fragmentation described by Raciborski (41) and Politis 
(39) is but the separating of the aggregated plastid-like bodies. This 
can be seen in Vanilla. That there is ever a passive division of an aggre- 
gated mass of oil bodies by the cell wall is improbable. Neither such 
aggregations nor a great development of oil was found in 
of Iris, Vanilla and the hepatics. 



With the recognition of elaioplasts as plastids, a study of their varia- 
tions became a study of the variations in plastids and chondriosomes. 
No new phenomena have been noted, but significant interpretations of 
those already recorded in the literature^ have been made. 

Most striking of the phenomena observed was the development of 
large quantities of oil globules in plastids. The formation of oil globules 
in plastids has been known for a long time and has recently been empha- 
sized by Guilliermond's (15-20) studies of Iris. But even Guillier- 
mond's extensive investigations have not shown an accumulation of oil in 
plastids comparable to that found in Iris versicolor where the quantity is 



260 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

so great as to obscure the structure of the plastids and render them 
unrecognizable for months at a time. 

The association of oil globules with young or degenerating forms 
more frequently than with mature plastids has been suggested by 
Guilliermond et al. (20). But such is not the case in Iris where the 
largest formation of oil is in the functioning plastids of the rhizomes. 

A second phenomenon noted was the plastic quality of chondriosomes 
and of transitional types of plastids. As evidenced in amoeboid move- 
ments this has often been recorded in the literature, while it has been 
emphasized in the recent studies by Guilliermond and his associates. 
But I have found in the literature no reference to the extreme plasticity 
amounting to fluidity such as occurs in some young leucoplasts where 
the included globules are moved about at random within the plastid. 

The significance of the chondrioconts has been brought out clearly 
in the survey of the variations of plastids and chondriosomes made in 
this study. The chondrioconts are essentially plastids producing at 
times all of the visible products of plastids such as starch, chlorophyll 
or a yellow pigment. They share, too, the plastic qualities of plastids 
which they display to a much greater degree. They occur in restricted 
tissues as transitional stages in the formation of plastids from mito- 
chondria-like primordia or as more or less degenerating forms of plastids. 
Often in the rootcap and in floral tissue they are pigmented, although 
the formation of pigment is not confined to them. It should be noted 
that in some tissues chondrioconts persist without assuming the more 
usual plastid form. 

Chondrioconts should not be interpreted as invariably forming a 
stage in the development of chloroplasts [Guilliermond et al. (20)]. 
On the contrary my studies show that the majority of chloroplasts and 
other plastids develop from mitochondria-like primordia without the 
intervention of a chondriocont stage. Where chondrioconts do form a 
stage in the development of plastids, the whole chondriocont develops 
into a plastid. There is no budding or fragmenting of the chondriocont 
involved. The appearances that have been interpreted as budding in 
chondrioconts or as evidence of fragmentation are but temporary shapes 
of the plastic chondrioconts. 

It may be noted here that the studies of chondrioconts emphasize 
Kassmann's (25) observations that plastids do not divide under normal 
conditions. This is a much debated point in the literature upon plastids. 

There was no evidence of vacuole formation in degenerating plastids 
or chromatophores such as have been described in flowers [Guilliermond 
et al. (20) I . The appearance which has been interpreted as a vacuole is 



1935] FAULL, ELAIOPLASTS IN IRIS 261 

rather the inclusion of a small amount of protoplasm as a result of the 
amoeboid movements of the plastid at this time. 

It is worth emphasizing here that the complete degeneration of the 
plastids may occur without involving the death of the cell. It has 
already been noted by Beer (4) that such a phenomenon occurs in some 
floral organs where the life of the mature cells is comparatively brief. 
I have found no record, however, of the degeneration of the plastids in 
cells which remain alive for months thereafter, a phenomenon found in 
the rhizomes of Iris macrosiphon. 

In general, it may be stated that there is no sharp line of demarcation 
between elaioplasts, plastids, chondrioconts and mitochondria. In Iris 
they have all been observed to form starch and, with the exception of 
mitochondria, chlorophyll, oil and a yellow pigment. In some instances 
several of these products may appear at once, or they may develop in 
succession, or none of them may form. Nor should any of the chon- 
drioconts and plastids be considered end products of a developmental 
series originating from mitochondria-like bodies, for until irreversible 
changes occur such as a resolution into structureless spheres of oil, the 
shapes assumed are reversible. In other words there is no clear dis- 
tinction between amyloplasts, leucoplasts, chloroplasts, chromoplasts 
and elaioplasts; nor is it possible to consider plastids, chondrioconts, pro- 
plastids and mitochondria as unrelated cell structures. Rather it 
appears that these are all forms of the same fundamental cell organ 
differing only in size and in the chemical products being formed at the 



3. SiGXIFICANXE OF THE StUDY OF OlL-BF.ARING PlASTIDS IX h 
FROM A TaXONOMIC VIEWPOINT 

The occurrence of two types of plastids in rhizomes of Iris each i 
or less restricted to certain groups of species appears to be of t 
significance. The consistent appearance of the same type in well defined 
species indicates a character that may be useful in separating species. 
In addition it should be noted that the substitution of compound starch 
grains for the elaioplasts in rhizomes of a known hybrid and in one or 
two questionable species, although not an invariable phenomenon, sug- 
gests a possible means of identifying some plants as of hybrid origin. 



CONCLUSIONS 
1. The anomalous bodies in the rhizomes of Iris ver 
elaioplast phase of leucoplasts persisting throughout the i 
but forming starch throughout the actively growing perio 



JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

Some, if not all, of the so-called "elaioplasts" are plastids in some 

3. Elaioplasts of the plastid type are not necessarily degeneration 
types: in Iris they are functional plastids. 

4. The rotary movement of elaioplasts described in the literature 
is an artefact due to slow death or injury; the Brownian movement de- 
scribed as characteristic of globules in certain liverworts is a degenera- 

>n or injury phenomenon. 

5. The elaioplasts in Lunularia and Vanilla are morphologically 
nilar in that they are aggregations of small plastid-like bodies that 
-m oil. This establishes another link between elaioplasts in the liver- 
worts and those in the monocotyledons. 

6. There is no sharp line of demarcation between the different kinds 
of plastids and chondriosomes each of which is a more or less temporary 
form capable of changing to the other types. 

7. At all times the plastids are more or less plastic but particularly 
so in young tissues, fibrovascular tissue or slowly dying cells. 

8. Leucoplasts, chloroplasts and chromatophores do not go through 
a set series of changes in developing from plastid types characteristic of 
meristematic tissues. They may pass through various series dejDending 
upon the type of mature tissue involved, or they may merely increase 
in size with probable changes in their physico-chemical structure. They 
never form by budding of chondrioconts succeeded by separation of 
the buds so-formed. 

9. Chondrioconts may form an intermediate developmental stage in 
the formation of "mature" plastids, although not necessarily; they may 
persist in some tissue; or they may be an intermediate stage in the 
degeneration of plastids. 

10. Plastids and chondriosomes in Iris all show the structure of a 
matrix with embedded globules. Pigments are more soluble in the 
globules than in the matrix, although they are found in both. 

11. Two types of degeneration of plastids occur involving (a) an 
increasing fluidity and a decreasing refractivity or (b) a complete 
breaking down into large homogeneous spheres of oil. Degeneration of 
the plastids does not necessarily involve the death of the cell. 

12. The formation --of a vacuole with at times an included oil drop 
in degenerating chondrioconts is in reality an inclusion of protoplasm. 

13. Different species of Iris are characterized by distinct types of 
elaio-leucoplasts in their rhizomes. The distribution of types follows 
closely the taxonomic groupings and may be of significance in separating 



1935] FAULL, ELAIOPLASTS IN IRIS 263 

14. The occurrence of such elaioplasts as those in rhizomes of Iris 
versicolor is confined, so far as could be ascertained, to rhizomes of this 
genus. For the most part they are restricted to rhizomes of certain 
species, chiefly Apogons of Asia and America. 

15. Refractive bosses on pollen grains of Iris Xypkium are oil- 
bearing chromatophores adhering from the tapetal fluid. Other mark- 
ings found on pollen grains were part of the wall structure. 

ACKNOWLEDGMENTS 

I am under deep obligations to Professor I. W. Bailey of Harvard 
University for his continuously helpful suggestions and interest through- 
out the course of the investigations recorded in this paper. I also wish 
to thank Dr. George M. Reed of the Brooklyn Botanical Garden and 
Dr. Edgar Anderson, recently of Washington University and the 
Missouri Botanical Garden for the material which they kindly placed 
at my disposal. Dr. Anderson also aided materially in connection with 
pertinent taxonomic matters. I wish to express my appreciation too of 
the advice and material offered by Dr. Conway Zirkle and others. 

Most of the work was done in Professor Bailey's laboratory in Har- 
vard University. I spent one summer in Professor Bailey's laboratory 
at the Stanford Laboratory of the Carnegie Institution of Washington, 
Division of Plant Biology, Palo Alto, California, and several weeks at 
the Missouri Botanical Garden in St. Louis. I am grateful to these 
) for the facilities that they afforded. 



LITERATURE CITED 

Bailey, I. W. The cambium and its derivative tissues. V. A recon- 
naissance of the vacuome in living cells. (Zeitschr. Zell. Mikr. 
Anat. 10:651-682. 1930.) 

and C. Zirkle. The cambium and its derivative tissues. 

VI. The effects of hydrogen ion concentration in vital staining. 
(Jour. Gen. Physiol. 14:363-383. 1931.) 

, C. Zirkle and A. Faull. Investigations on the cam- 
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399-401. 1931.) 

Beer, R. On elaioplasts. (Ann. Bot. 23:63-72. 1909.) 

Chalaud, G. Le cycle evolutif de Fossombronia pusilla Dum. (Rev. 
Gen. Bot. 41:24, 95, 129, 213, 293, 353, 409, 474, 541, 606, 676. 
1929.) 

Dangeard, p. a. Recherches sur la structure de la cellule dans les 
Iris. [Compt. Rend. Acad. Sci. (Paris) 174:1653-1659. 1922.] 

DoMBRAY, P. Contribution a I'etude des corps oleiformes des hepat- 
iques des environs de Nancy. Nancy, 1926. 

Dykes, W. R. The genus Iris. Cambridge, 1913. 

Gargeanne, a. J. W. Die 01k5rper der Jungermanniales. (Flora, 
92:457-482. 1903.) 



JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

des Jungermanniales. [Compt. Rend. Acad. Sci. (Paris) 184: 
1473-1475. 1927.] 

Sur les rapports du vacuome et du systeme oleifere des 

Jungermanniaceae. (Ibid. 186:163-166. 1928.) 

Sur la presence du systeme oleifere dans les organes de 

multiplication des Jungermanniaceae. (Ibid. 186:886-888. 1928.) 

GoTTSCHE, C. M. Anatomisch-physiologische Untersuchungen iiber 

Haplomitrium Hookeri. (Verb. Leop. Carol. Akad. 12(1) :287- 

288. 1843.) 
Grout, A. J. Mosses with a hand lens. 3rd Ed. New York, 1924. 
GuiLLiERMOND, A. Sur I'etude vitale du chondriome de I'epiderme 

des petales d'Iris germanica et son evolution en leuco- et chromo- 

plastes. [Compt. Rend. Soc. Biol. (Paris), 74:1280-1283. 1913.] 
Nouvelles observations vitales sur le chrondiome des cellules 

epidermiques de la fleur d'Iris germanica, I. (Ibid. 78:241-245. 

1915.) 
Nouvelles observat 

epidermiques de la fleur ( 

1915.) 

Sur I'origine et la signification des oleoplastes. [Compt. 

Rend. Soc. Biol. (Lyon), 86:435. 1922.] 

Sur la presence d'un corps d'aspect oleagineux dans les 

cellules epidermiques d'Iris germanica [Compt. Rend. Soc. Biol. 
(Paris), 109:1182. 1932.] 

, O. Mangenot et L. Plantefol. Traite de cytologie vege- 

tale. Paris, 1933. 
Harper, R. A. Nature and functions of plastids, especially elaio- 

plasts. [Proc. Internat. Congr. Plant Sci. (Ithaca), 1:311-316. 

1916.] 
Hieronymus, O. Beitrage zur Morphologic und Biologic der Algen. 

(Cohn's Beitr. 5:467-468. 1889.) 
HoFMEisTER, W. Die Lehre von der Pflanzenzelle. Leipzig, 1867. 
HoLLE, O. Uber die Zellenblaschen der Lebermoose. (Verb. Leop. 

Carol. Akad. Heidlb. 1857, p. 396. 1857.) 
Kassmann, F. Die Entwicklung der Chondriosomen und Chloro- 

plasten von Cabomba aquatica und Cabomba caroliniana auf Grund 

von Dauerbeobachtungen an lebenden Zellen. (Planta, 1:624-656. 

Keene, M. L. Zygospores in Sporodinia grandis. (Ann. Bot. 28: 

455^70. 1914.) 
Studies of zygospore formation in Phycomvces nitens Kunze. 

(Trans. Wis. Acad. Sci. Arts and Letters, 19:1195-1220. 1919.) 
KozLOwsKY, A. Sur I'origine des oleoleucites chez les hepatics a 

feuilles. [Compt. Rend. Acad. Sci. (Paris), 173:497^99. 1921.] 
KusTER, W. Die Olkorper der Lebermoose und ihr Verhaltnis zu 

den Elaioplasten. Basel, 1894. 
Lehmann, C. Studien iiber den Bau und die Entwicklungsgeschichte 

von Olzellen. (Planta, 1 : 343-373. 1925.) 
Leiner, B. Untersuchungen iiber das Olplasma und die Oleoplasten. 

(Jahrb. Phil. Fak. IL Bern, 4:95-113. 1924.) 
Lidforss, B. Studier ofver Elaiosferer i Ortbladens Mesofyll och 

Epidermis. (Kgl. Fvsiogr. Sallsk. i Lund Handl. 4:1-33. 1893.) 



5] FAULL, ELAIOPLASTS IN IRIS 

LuNDSTROM, A. N. Uber 

Bedeutung der Oeltropfi 

Centrbl. 35:177-181. 1888.) 
Meyer, A. Morphologische und physiologische Analyse der Z 

Jena, 1920-1926. 
MiRBEL, C. F. Recherches anatomiques et physiologiques sui 

Marchantia polymorpha. (Mem. Acad. Roy. Sci. Inst. Fra 

13:345. 1835.) 
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Oxalate in der Pflanze. (Bot. Centralbl. 43:327. 1890.) 
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(Ann. Bot. 32:91-112. 1918.) 
Pfeffer, W. Die Olkdrper der Lebermoose. (Flora, 57:2-6, 17-27, 

33-43. 1874.) 
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Bot. Pavia, ser. 2, 14:335-361. 1914.) 
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bei Liliaceen. (Bull. Acad. Sci. Cracovie Compt. Rend. 1893 

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(Trans, and Proc. Bot. Soc. Edin. 16:123-128. 1886.) 
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Schurhoff, p. N. Die Plastiden. Berlin, 1924. 
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(Jahrb. Wiss. Bot. 19:423-496. 1888.) 
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DESCRIPTION OF PLATES 
^igs. 1-4, 6, 8-10, 38, 39, 45, 46 were made with a camera lucida ; the 
gnifications given for these figures are exact. The other figures were 
iwn free-hand; the magnifications given for them are approximate. 

Plate 132 
If. 1 . Iris versicolor L. Living cells of the rhizome from material 

collected in December. X 475. 
X. 2. Individual elaioplasts from cells shown in Fig. 1. X 1600. 
\. 3. Individual elaioplasts from cells shown in Fig. 1 after treatment 

with (a) ammonium Erliki fixative, and erythrosin and cyanin ; 

and (b) 0.5% osmic acid. X 1600. 



266 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

Fig. 4. Iris versicolor h. Individual elaioplasts from rhizomes collected 
in October: (a) surface view; (b) included starch grains; 
(c) diagram to illustrate position, size, number and shape of 
starch grains. X 1600. 

Fig. 5. Iris versicolor L. Individual elaioplasts treated with Gram's 
solution: (a) material from Duxbury, Mass.; (b) material 
from Lincoln, Mass. These were drawn at the same magnifica- 

Fig. 6. Isolated starch grain from elaioplast shown in Fig. 4. X 1600. 

Fig. 7. Iris tec t arum M.2i.y.im. Elaioplasts from young cells of rhizomes 
collected in March: (a) normal plant; (b) dying plant. X 845. 
Plate 133 

Fig. 8. Iris pallida Lam. X ? Living cell of the rhizome from material 
collected in December. X 475. 

Fig. 9. Individual oil-bearing plastids from cell shown in Fig. 8 showing 
plastids: (a) without starch; (b) with starch; (c) is a diagram 
showing the relative positions of plastid and starch. X 1600. 

Fig. 10. Individual starch grains from plastids similar to those shown 
in Fig. 9. X 1600. 

Fig. 11. Iris pumila L. Oil-bearing plastids from living cells of a rhi- 
zome collected at St. Louis in March: (a), (b) and (c) are 
plastids from successively older cells. X 1270. 

Fig. 12. Oil-bearing plastids from living cells of rhizomes of the follow- 
ing California species of Iris: (a) /. missouriensis Nutt. ; (b) /. 
Douglasiana Herb.; (c) /. longipetala Herb.; (d) /. Hartwegii 
Baker. Material collected in California in August. X 1270. 

Fig. 13. Diagram showing the types of oil-bearing plastids found in Iris 
species in March. See table p. 246 for names. 
Plate 134 

Fig. 14. Iris macrosiphon Torr. Elaioplasts in living cells from the 
cortex of a rhizome collected in July: (a), (b) and (c) are 
taken from successively older cells. X 1245. 

Fig. 15. Elaioplasts shown in (a) Fig. 14b and (b) Fig. 14c treated with 
Gram's solution to show the surrounding cytoplasm. X 1245. 

Fig. 16. Iris Xyphium L. Optical section of living pollen grain. X 500. 

Fig. 17. Iris Xyphium L. Untreated pollen grains in surface view: 
(a) in tapetal fluid of immature anther; (b) from a ripened 

Plate 135 
Fig. 18. Iris macrosiphon Torr. Oil-bearing plastids in living epidermal 



Fig. 19. Ins 

from successively older cells. 

Fig. 20. Iris pallida Lam. X ? Plastids from living, elongated, differen- 
tiating cells of root-tip: (a) successive observations on a single 
plastid to show fluctuating variations in form; (b), (c), (d) and 
(e) similar observations on four additional plastids. X 1620. 

Fig. 21 . Plastid similar to those in Fig. 20 but from an older cell. X 1620 

Fig. 22. Plastid similar to that shown in Fig. 21. X 1620. 

Fig. 23. Plastid similar to that shown in Fig. 21. X 1620. 

Fig. 23a. Plastid similar to that shown in Fig. 21. X 1620. 



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FAULL, ELAIOPLASTS IN IRIS 267 

Iris pallida Lam. X ? Elaioplasts from living cells of cortex 

Iris pallida Lam. X ? Elaioplasts from living cells of cortex of 
older root. X 1620. 

Iris pallida Lam. X ? Plastids and chondriosomes from a single 
living cell of the central cylinder of a root-tip. X 1620. 
Plastids and chondriosomes from another cell of the central 
cylinder. X 1620. 

Iris pallida Lam. X ? Oil-bearing plastids from living cells of 
the meristem of a rhizome: (a), (b) and (c) are from suc- 
cessively older cells and show the appearance of starch. X 1620. 
Iris pallida Lam. X ? Chromoplasts from living cells of the 



Iris 


pallida Lam. X ? Chrom 


atophor 


es from living 


cess 


ermis of the root-tip: ( 
vely older cells. X 1650 


0, (b) 


, (c) and (d) 


Ms 


versicolor L. Plastids f 




ng cells of th 


of a 


root-tip. X 1650. 






Iris 


pallida Lam. X ? Chrom 


atophor 


es from living 


epidermis of a root-tip. X 1650. 




Chr 


Dmatophores from living 


ells of 


the epidermis 


of a 


Pogoniris, probably of /. 




taU X 1650 


Iris 


versicolor L. Oil-bearing plastids from living c 



from living parenchyma cells 
b) vv^ith starch. X 1650. 
Ti living cells of the epidermis 



from a mature thallus. X 1080. 
Lunularia cruciata (L.) Dum. Living elaioplas 
from the younger tissue of a mature thallus. X 1 
One of the Jungermanniales. Living cell from ; 



of the Jungermanniales. X 1250. 

One of the Jungermanniales. Oil bodies 

tiating cells of stem: (a), (b) and (c) fr 

One of the Jungermanniales. Oil bodies 

mature plant. X 1250. 

One of the Jungermanniales. Oil bodies 


from living differen 
om successively olde 

from living cells o 

from living cells o 


younger tissue. X 1250. 

Vanilla Pompona Schiede. Elaioplast and chloroplasts fron 

living cell of a leaf. X 915. 

Vanilla Pompona Schiede. Elaioplasts, chloroplast and chondri 

osomes in living cell of cortex of root-tip. X 1720. 



JOURNAL OF THE . 



NOTES ON YUCCA 



Yucca Thornberi, spec. nov. Plate 138 

Trunci 0.75-1.75 m. alti, congest!, infra foliis siccis reflexo-patentibus 
arete obtecti, supra comam magnam satis elongatam foliorum viridium 
gerentes. Folia lineari-lanceolata, 0.30-1 m. vel ad 1.20 m. longa, 
1.5-3.5 cm. lata, a basi vel a medio paullo latiore apicem versus attenuata, 
acuminata, pungentia, concava, utrinque laevia, luteo-viridia, stricta 
vel leviter recurva, flexibilia, initio margine saepe evanescenter denticu- 
lata, mox filifera filis crassiusculis curvatis tarde deciduis, parte basali 
2.5-7.5 cm. longa et 7-12.5 cm. lata. Inflorescentia scapo 22-45 cm. 
longo incluso 1-1.30 cm. alta. angusta, basi et apice attenuata, folia 
quarta parte vel dimidio superantes; ramuli circiter 25, initio erecto- 
ascendentes, demum patentes; bracteae magnae, late triangulares; flores 
campanulati, 7.5-12.5 cm. longa; pistillum 5.5-7.5 cm. longum, ovario 
plerumque oblongo rarius attenuato, 4j/2 ad 6>^ longiore quam lato, stylo 
5-6 mm. longo, stigmatibus sub anthesi erectis vel fere erectis; filamenta 
4-6.5 cm. longa, apice clavato brevi pro parte inferiore longo gracili. 
Fructus 13-17.5 cm. longus, 3-4.5 cm. diam., baccatus, incrassatus, 
apicem versus attenuatus et 2.5-5 cm. sub apice subito constrictus parte 

Arizona. Pima Co.: foothills of the Rincon Mts., slightly 
north of Rincon Creek, a tributary of Pantano Wash, elevation 3600 ft., 
5. D. McKelvey, no. 1627, March 23, 1930 (type; herb. Arnold Arbore- 
tum). Also from the same region are the author's collections nos. 1585, 
2123, 2556, 2557, 2558, 2559, 2561, 2562, 2682, 2684. 

Yucca Thornberi forms large and crowded clumps and produces many 
rather long stems 2-5 ft. in height which are covered below with a thick 
thatch of reflexed-spreading dead leaves and are crowned by large, some- 
what elongated heads of green foliage which are constricted near the 
base and spreading above; the slightly broadened, concave, not conspicu- 
ously angled leaves are commonly smooth on both surfaces, yellow-green 
in color, with acuminate apex and, when young, often evanescently 
denticulate margins which, when the foliage is more mature become 
abundantly filiferous; the fibres are late-deciduous, moderately coarse 



NOTES ON YUCCA 269 

and loosely curled. The inflorescence is for some time rather fleshy and 
brittle, 3-4 ft. in length overall, with a scape 9-18 in. in length; the 
inflorescence proper is long, narrow, tapered at both ends, and extends 
for Ya-Yz its length above the leaves; its branchlets are about 25 in 
number, at first erect-ascending, eventually spreading; its bracts are 
large, fleshy to leathery, broad-triangular in form. The flowers are 
campanulate, large, 3-5 in. in length; the pistil is lY?.-^ in. long with 
a commonly oblong, only rarely tapered, ovary which is 4>^-6>^ times as 
long as broad, the short style is fV-^ in. long and, at anthesis, with erect, 
or nearly erect stigmas; the filaments vary from approximately l>4-2>^ 
in. in length and reach anywhere from slightly below to slightly above 
the shoulders of the ovary; their clavate tip is short in proportion to the 
long, slender, lower portion. The baccate fruit is 5-7 in. in length, 
slightly enlarged and tapered upward for its major lower portion, for 
1-2 in. below the tip much contracted and commonly somewhat 

Yucca Thornberi appears to be most closely related to Y. arizonica and 
to F. baccata Torr., differing conspicuously from the latter in habit of 
growth, from both species it is distinct in form of inflorescence, in char- 
acter of foliage and, though less so, in fruit. 

It is a pleasure to give to this new species the name Yucca Thornberi 
in recognition of the fact that Dr. J. J. Thornber of the University of 
Arizona called the author's attention to the plant and with her spent 
considerable time in its study. 

Yucca brevifolia Engelm. var. Jaegeriana, var. nov. Plate 139 

A typo recedit habitu humiliore vix 3-4 m. excedente, trunco brevi 
circiter 75-90 cm. longo, ramis brevibus fere erectis arete congestis, 

tarn et symmetricam formantibus, inflorescentiis vix 30 cm. longis, scapo 
2.5-5 cm. longo et 2.5-3 cm. crasso incluso, ramulis tantum 2.5-6.5 cm. 

California. San Bernardino Co.; vicinity of the Shadow 
Mts., elevation approximately 4000 ft., S. D. McKelvey, no. 2732, April 
30, 1932 (type; herb. Arnold Arboretum). 

In several of the broad basins and foothill areas of the eastern part of 
the Mohave Desert of California and also in southern Nevada, occurs a 
form of the Joshua-tree which deserves varietal recognition. The plant — 
in appearance a miniature Joshua-tree — was brought to the attention of 
the writer by Mr. Edmund C. Jaeger of Riverside Junior College, River- 
side, California, and is named in appreciation of this fact. 



270 JOURNAL OF THE ARNOLD ARBORETUM [vol.xvi 

This variety is primarily distinguished from the better known plant 
by its dwarfer habit,— the plants not exceeding, except rarely and only 
in extremely old specimens, 10-12 ft. in height; the trunk is short, about 
lYi-l) ft. in length, stout (although, proportionately to that of the 
type of the species, slender); the branches are short, nearly erect and 
form an extremely dense, compact crown; the clusters of green leaves 
are crowded and symmetrical, about 1-2 ft. in length; the leaves are 
short, including the base about 4 in. long, not exceeding 8 in.,— or about 
the length of the shorter leaves of the type of the species; the inflorescence 
scarcely reaches 1 ft. in length; the scape is 1-2 in. long, 1-1^ in. in 
diameter at base, and the flowering portion 9-10 in. in length with stout 
branchlets only l-2>4 in. long. In flower and in fruit characters it is 
very similar to the plant of taller habit. 

Mr. Jaeger states (in litt., Oct. 2, 1934) that the distribution of the 
variety "reaches its greatest density in the vicinity of the New York Mts. 
in California." It has been collected by the writer in southern Nevada 
between the Colorado River and Searchlight (no. 4094), in the Spring 
Mts. (no. 4142) and on the eastern slopes of the Charleston Mts. (nos. 
4097, 4098, 4099, 4100, 4132), in the first and last of which regions it 
occurs in abundance. 

Yucca arizonica, nom. no v. 

Yucca puberula sensu Torrey in Botany, Emory Report, 221 (1859), 

in part, not Haworth. 
Yucca brevifolia Schott ex Torrey, Botany, Emory Report, 221 
(1859), in part, as synonym of Y. puberula Torrey, not Haworth. — 
Engelmann in Trans. Acad. St. Louis, 3:46 (1873), in part, as 
synonym of Y. Schottii Engelm. — Trelease in Rep. Mo. Bot. Gard. 
13: 100, tt. 57-59, 96 (fig. 2, range map) (1902), first appearance 
as a valid species. — Not Y. brevifolia Engelm. (1871). 
Yucca Treleasei MacBride in Contrib. Gray Herb. ser. 3, no. 56: 15 
(1918) ; not y. Treleasii Sprenger (1906). 
The name Y. puberula Haw. was first erroneously applied by Torrey 
to specimens collected by Arthur Schott in regions adjacent to Nogales, 
Arizona. Because of its connotation it is in the main referable to F. 
Schottii Engelm., the inflorescences of which species are commonly 
puberulous. Schott's material represented a complex. For a certain 
portion of this material the name F. brevifolia used by Schott in his notes 
was adopted by Trelease in 1902. This name, as pointed out by Mac- 
Bride in 1918 was antedated by the name F. brevifolia used by Engel- 
mann in 1871 for the Joshua-tree; MacBride in consequence gave to 
Schott's plant the new name F. Treleasei. 



1935] McKELVEY, NOTES ON YUCCA 2 

Unfortunately the name Y. Treleasii was used by Carl Sprenger 
1906 for a hybrid Yucca (See Bull. Soc. Tosc. Ort. 31 : 134. 1906.- 
Molon, Yucche, 192, t. 6. 1914); the plant is without a name and tl 
new name Yttcca arizonka is here adopted for this species. 



JOURNAL 

OF THE 

ARNOLD ARBORETUM 



THE VISIBLE STRUCTURE OF THE SECONDARY WALL AND 

ITS SIGNIFICANCE IN PHYSICAL AND CHEMICAL 

INVESTIGATIONS OF TRACHEARY CELLS 

AND FIBERS 

I. W. Bailey and Thomas Kerr^ 
With plates 140-149 

INTRODUCTION 

The secondary wall of plant cells has long been known to be a 
heterogeneous structure. That it is more or less conspicuously striated 
and laminated was shown by Mirbel, Von Mohl, Valentin, Meyen, Th. 
Hartig, and other pioneer anatomists who demonstrated, in addition, 
that it may be resolved by specific chemical and mechanical treatments 
into lamellae, fibrils, granules, and other visible units of fairly constant 
form and size. This led, during the second half of the last century, to 
prolonged discussions concerning the fundamental structure of cell walls 
in general, and to much speculation regarding the physiological processes 
involved in their formation. 

Although a voluminous literature developed between 1850 and 1900, 
no consensus of opinion was reached concerning the exact physical and 
chemical significance of the visible heterogeneity of the secondary wall. 
Nor is there a general agreement among different groups of investigators 
at the present time. It is true that the study of anisotropy, of rod double 
refraction, of various types of dichroism, and of X-ray diagrams has in 
recent years contributed much toward a clearer understanding of sub- 
microscopic structures, and regarding the orientation of such structures 
in the grosser layers of the secondary wall, but it has not afforded as 
yet an adequate explanation of the finer types of visible heterogeneity. 



274 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

In view of such facts as these, it seemed desirable to the writers to 
undertake a detailed investigation of the secondary wall in an endeavor 
(1) to verify and, if possible, to amplify the observations of previous 
workers; (2) to correlate results obtained by different techniques .and 
by the study of divergent cell types; and (3) to interpret the visible 
heterogeneity of the secondary wall in terms of its sub-microscopic 
structure and of its chemical composition. 

In an investigation of this character one is faced, at the outset, by a 
serious difficulty, upon the solution of which success or failure clearly 
depends. The range of recorded cases in which the details of wall struc- 
ture are even vaguely visible — without resorting to the use of softening 
or hardening processes, of macerating or swelling agents, and of other 
more or less drastic chemical and mechanical treatments — is very lim- 
ited. Severe treatments are capable of yielding extremely useful and 
significant data, but are likely to produce distortions and other artifacts, 
and therefore must be checked by observations on untreated material. 
In other words, an adequate system of controls — or means of accu- 
rately visualizing the normal structure of the secondary wall — is 
indispensable. 

As indicated in the preceding paper of this series (18), it is possible 
to section dense woods and other hard tissues without resorting to the 
use of softening processes which might modify their structure and 
chemical composition. It seemed advisable, accordingly, to make an 
extensive survey of a wide range of gymnosperms and angiosperms in 
search of species that afford clearly defined images of cell wall struc- 
ture in untreated sections. More than 3000 species, representing 160 
families and 40 orders, were examined. It was found that the large- 
celled woods of various tropical dicotyledons provide unusually favor- 
able material for microscopic investigations. These plants are not 
bizarre or unusual forms; nor are they confined to any restricted group 
or genus. They are widely distributed and of not uncommon occur- 
rence in such families as the Theaceae, Monimiaceae, Icacinaceae, 
Rhizophoraceae, Euphorbiaceae, Flacourtiaceae, etc. When thin 
(5-10 m), smoothly-cut sections of the wood are examined in liquids of 
the right index of refraction, using the best modern optical equipment, 
the relatively broad expanse of wall in the fiber-tracheids and libriform 
fibers of certain of these plants reveals finely laminated, striated, and 
reticulated structures in exquisite detail. By using untreated sections 
of such plants as controls, it is possible to determine the exact effects 
upon normal structures of varied chemical and mechanical treatments, 
and thus to extend the scope of investigation to cover a wide range of 
less favorable species and tissues. 



1935] BAILEY AND KERR, STRUCTURE OF THE SECONDARY WALL 275 

The following discussion of tracheary cells and libers is divided into 
two parts. The grosser and more conspicuous types of layering of the 
secondary wall are dealt with in Part I ; structures which more nearly 
approach the limits of microscopic visibility, in Part II. As previously 
stated, considerable is known^ concerning the physical factors involved 
in the differentiation of the former structures, which must be clearly 
visualized and accurately correlated before proceeding to a detailed 
consideration of the finer types of visible heterogeneity. 

TERMINOLOGY 
The terms middle lamella, primary wall, secondary wall, and tertiary 
wall have been employed in several fundamentally different senses and 
to designate entirely different structures. This has led to much con- 
fusion in the literature and to serious discrepancies, not only in descrip- 
tive morphological work, but also in physiological, biophysical, and bio- 
chemical investigations. As a result of our detailed study of the cam- 
bium and its derivatives and of our preliminary investigations of other 
meristems and their derivatives, we attempted, in a former paper (18), 
to clarify the situation by suggesting that (1) the term middle lamella 
be used synonymously with intercellular substance in referring to the 
truly isotropic material which separates the walls of adjoining cells; 
( 2 ) the term primary wall should no longer be applied to the first-formed 
layer of secondary thickening, but should be reserved for the original 
wall of the cell which is formed in the meristematic region and is carried 
over in more or less modified form into the fully differentiated tissues; 
and (3) the term secondary wall be used in referring to the strongly 
anisotropic layers of secondary thickening which are formed after a cell 
has attained its final size and shape. The term tertiary wall is so vari- 
ously used and interpreted and so confusing that its use should be dis- 
continued. We propose to employ our revised terminology in this and 
succeeding papers. 

I. THE PRINCIPAL LAYERS OF THE SECONDARY WALL 
A. Layering Due to Physical Factors 

The secondary wall of normal tracheids, fiber-tracheids, and libri- 
form fibers commonly consists of three layers of different refractive 
character; (1) a relatively narrow outer layer, (2) a narrow inner layer, 
and (3) an intervening layer of variable thickness. When thin, per- 



276 JOURNAL OF THE ARNOLD ARBORETUM Fvol. xvi 

fectly transverse sections of such cells are examined in polarized light 
between crossed nicols, Fig. 3, the inner and outer layers exhibit strong 
double refraction and are brilliant — except in positions of extinction — 
whereas the central layer is dark or noticeably less birefringent. The 
conditions tend to be reversed in longitudinal sections, Fig. 2, in which 
the central layer shows intense double refraction, and the inner and outer 
layers are dark or less conspicuously birefringent. In other words, 
as shown long ago by Dippel (7) and others, the secondary wall con- 
sists of anisotropic layers which are dark or brilliant in polarized light 
depending upon the plane of sectioning of the cell or upon the angle from 
which the wall is viewed. 

Our extensive survey of gymnosperms and angiosperms has demon- 
strated that most tracheids, fiber-tracheids, and libriform fibers are pro- 
vided with a secondary wall of this 3 -layered type. The narrow inner 
and outer layers are of relatively constant thickness, not only in differ- 
ent parts of a given plant but also in plants of different systematic 
affinities. Variations in thickness of the secondary wall are due, there- 
fore, primarily to fluctuations in the width of the central layer. When 
the secondary wall is thin, as in the tracheids of the early wood of many 
conifers, the inner and outer layers are so closely approximated that the 
tenuous intervening central layer is invisible in polarized light, except 
in very thin (3-7 |j), perfectly transverse sections of straight-grained 
tissue. In thicker or obliquely cut sections, the width of the inner and 
outer layers is much exaggerated by the scattering of light from these 
intensely birefringent structures. This fogs and conceals the central 
layer, just as the closely approximated brilliant outer layers of adjacent 
cells commonly obscure the tenuous primary walls and middle lamella 
(compare Figs. 1 and 3). 

Deviations from the normal 3-layered type of secondary wall are of 
not infrequent occurrence. Thus, many thick-walled libriform fibers 
and fiber-tracheids have no clearly differentiated inner layer, whereas 
others have more than three layers of varying width and birefringence, 
Fig. 4. Walls of a multiple-layered, anisotropic type, which are of rela- 
tively sporadic occurrence in the fiber-tracheids and libriform fibers of 
dicotyledons, are characteristic features of the libers of many mono- 
cotyledonous stems. In transverse sections of such fibers, Fig. 6, there 
are narrow brilliant zones in polarized light which alternate regularly 
with broader and conspicuously less birefringent ones. Variations in 
the thickness of the secondary wall of these cells are due largely to 
variations in the number of successively formed layers. 

The optical behavior of the anisotropic layers of the secondary wall 



1935] BAILEY AND KERR, STRUCTURE OF THE SECONDARY WALL 277 

of tracheary cells and fibers is closely correlated with the orientation of 
striations and so-called fibrillar structures, which are visible in cells that 
have been subjected to various chemical and mechanical treatments. 
When the striations and fibrils are arranged parallel, or nearly parallel, 
to the long axis of a tracheary cell or fiber, a layer is dark in sections 
cut at right angles to this axis, but is brilliant in longitudinal sections 
and in surface view — except, of course, in the four positions of extinc- 
tion. The intensity of the birefringence varies in obliquely cut inter- 
vening sections, decreasing as the plane of section approaches that of a 
truly transverse section. On the contrary, where the striations and 
fibrils are arranged approximately at right angles to the long axis of a 
cell, a layer is brilliant in cross sections and in surface view, but is dark 
in thin longitudinal sections, Fig. 2, which transect the fibrillar struc- 
ture. When the striations and fibrils have a helical arrangement and, 
therefore, are obliquely oriented in relation to the major axis of the cell, 
a layer is brilliant in surface view and more or less birefringent in both 
transverse and longitudinal sections. If the helix has a pitch of approxi- 
mately 45°, an oblique section, which is cut parallel to the striations 
and fibrils on one side of the cell, will transect these structures on the 
opposite side of the cell. Thus, in such sections, the layer will exhibit 
both isotropy and strong double refraction; i.e., it will be dark on one 
side of the section and brilliant on the opposite side. Changing the 
fibrillar orientation from a left-handed to a right-handed helix or vice 
versa will not alter the birefringence in transverse or in longitudinal 
sections so long as the angle of obliquity remains constant. 

In the typical 3-layered secondar>' walls of tracheids, fiber-tracheids, 
and libriform fibers, the striations and fibrils of the central layer are 
oriented parallel to the long axis of the cell, or at angles which do not 
deviate excessively from that axis; whereas those of the inner and outer 
layers are arranged more nearly at right angles to the major axis of the 
cell. Thus, the central layer exhibits strong double refraction in longi- 
tudinal sections. Fig. 2, and isotropy or relatively feeble double refrac- 
tion in transverse sections. Figs. 1 and 3; whereas the conditions are 
reversed in the case of the inner and outer layers of the secondary wall. 
In multiple-layered walls of the type illustrated in Fig. 6, the orienta- 
tion alternates regularly from parallelism to the major axis of the cell 
in the broader layers to marked obliquity in the narrower ones. The 
former layers exhibit intense double refraction in longitudinal sections; 
the latter layers, in transverse sections. 

In the case of optical anisotropy, the so-called index-ellipsoid has, 
according to Frey-Wyssling (13), a major axis (Ny) which is oriented 



278 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

parallel to the striations and fibrils, and two minor axes (Na and N^) 
which are placed at right angles to these structures. On the contrary, 
in the case of swelling-anisotropy, the ellipsoid of expansion has two 
major axes which are oriented at right angles to the striations and fibrils, 
and a minor axis which is parallel to these structures. Therefore, the 
dark layers of Figs. 1, 3, and 6, which have longitudinal striations, 
expand laterally, increasing in both width and circumference; whereas 
the strongly birefringent layers, the striations of which are oriented 
more nearly at right angles to the long axis of the cell, are unable to 
do so and expand longitudinally. Where the dark layers are of con- 
siderable width, they tend, by their excessive lateral expansion, to disrupt 
the thin birefringent layers, as indicated in Fig. 7 . 

The strongly anisotropic behavior of the secondary wall suggests that 
its layers are composed of sub-microscopic units which have definite 
planes of orientation, and that there is a close correlation between the 
orientation of these units and of such visible structures as striations and 
fibrils. It was in fact a consideration of these phenomena which led 
Nageli to formulate the Micellar Hypothesis. 

More recently, X-ray analyses and other physico-chemical investiga- 
tions have indicated that native cellulose consists of chains of anhydrous 
glucose residues which are bound together by secondary valences into 
a space lattice of definite dimensions. These chains are arranged parallel 
to each other, and, in the case of the secondary wall of fibers and of 
Valonia, are oriented parallel to the striations and fibrils — as shown by 
Katz ( 1 7 ) and by Astbury and his co-workers ( 1 ) . Furthermore, there 
is much cumulative evidence' from detailed investigations of anisotropy, 
of rod double refraction, of various forms of dichroism, and of X-ray 
analyses which suggests that the cellulose chains are not uniformly dis- 
tributed throughout the secondary wall, but are aggregated into more or 
less vaguely defined anisotropic units the major axis of which is oriented 
parallel to that of the visible striations and fibrils. 

In view of such facts as these, it is evident that layering of the type 
discussed on preceding pages is not due fundamentally to differences in 
chemical composition, but rather to changes in the orientation of aniso- 
tropic units of cellulose in the successively formed layers of the sec- 
ondary wall. 

B. Layering Due to Chemical Factors 

The broad central layers of normal fiber-tracheids and libriform fibers 

frequently have subsidiary layers of varying width which are much 



1935] BAILEY AND KERR, STRUCTURE OF THE SECONDARY WALL 279 

intensified by differential staining, Fig. 8. These subsidiary layers, un- 
like those illustrated in Fig. 4, are not closely correlated with variations 
in the orientation of the anisotropic cellulose, but are due to differences 
in lignification or to variations in the distribution of non-cellulosic con- 
stituents. They may be eliminated by delignification and other 
standard treatments for the purification of cellulose. It should be 
emphasized in this connection that the anisotropic layers of normal 
tracheids, fiber-tracheids, and libriform fibers are coherent even in walls 
that have been treated to remove their non-cellulosic constituents. There 
are evident planes of weakness but no actual discontinuities in the 
cellulosic matrix. 

Conspicuous, discontinuities are, however, of not infrequent occur- 
rence in the peculiar tracheids of "compression wood," in so-called 
gelatinous fibers, in certain types of bast fibers, and in sclereids. They 
are due to narrow layers of truly isotropic material which contain little, 
if any, cellulose. Thus, when sections of unlignified or delignified cells 
are treated with standard solvents of pectic compounds and hemicellu- 
loses, the layers dissolve and liberate the anisotropic layers of cellulose 
which may be slipped apart as shown in Fig. 26. These truly isotropic 
layers may be accentuated by differential staining and are clearly 
visible in ordinary light. Fig. 21. They present some difficulties, how- 
ever, when sections are examined in polarized light between crossed 
nicols. For example, the entire laminated structure in Fig. 21, with the 
exception of the narrow outer layer, is dark in polarized light, owing to 
the fact that the orientation of cellulose in the anisotropic layers is 
parallel to the long axis of the cell. Therefore, the truly isotropic layers 
are concealed in transverse sections, but they are clearly visible in radial 
longitudinal sections and appear as dark lines between the birefringent 
^" imilar tenuous isotropic films in the 

of each narrow anisotropic layer, Fig. 
6. They are masked in both transverse and longitudinal sections, since 
the broader anisotropic layers of cellulose are dark in cross sections, and 
the narrower ones are dark in longitudinal sections. 

It should be noted, before passing to a detailed consideration of the 
finer types of visible structures, that sclereids and other types of non- 
fibrous sclerenchyma have a fundamentally different type of secondary 
wall. The anisotropic layers of such cells — at least in tissues of the 
higher plants that we have examined thus far — show no conspicuous 
striations or fibrillar structures, either in the untreated or in the swollen 



280 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

condition of the cell wall. Furthermore, the anisotropic layers are bril- 
liant in polarized light in all planes of section of the secondary wall, but 
are dark in surface view. The birefringent layers alternate more or less 
regularly with others which are dark in all planes of view, Fig. 5. A 
detailed discussion of these cells and of other non-fibrous types is re- 
served for subsequent papers of this series. 



LiBRiFORM Fibers 

As stated in Part I, variations in thickness of the secondary wall of 
normal tracheids, fiber-tracheids, and libriform libers are due primarily 
to fluctuations in the width of the central layer, which may attain a 
radial breadth of more than 15 pi in the large-celled woods of various 
tropical dicotyledons. Therefore, the central layer provides more favor- 
able material for sectioning and for study at high magnifications than 
either the inner or the outer layers which are so tenuous as to present 
serious optical difficulties. 

Figure 10 is a transverse section of the wood of Siparuna bifida (P. & 
E.) A. DC. cut without preliminary softening or other modifying treat- 
ments. The broad central layer of the secondary wall is strikingly 
heterogeneous and exhibits a complex pattern of anastomosing radial 
striations. The striations are clearly visible in unstained sections 
mounted in water and in other liquids of varying indexes of refraction ; 
and, in white light, are optically of two types, i.e., light and dark. There 
are corresponding light and dark striations in tangential longitudinal 
sections, Fig. 13. It is evident, accordingly, that the central layer of the 
secondary wall in these cells is composed of thin plates or lamellae which 
have a radio-longitudinal or radio-helical orientation. The lighter 
lamellae are strongly birefringent in polarized light. Fig. 13, except in 
positions of extinction and in sections cut at right angles to the longi- 
tudinal axis of the lamellae; whereas the alternating lamellae are dark, 
or at least comparatively isotropic, in all planes of view.' 

The birefringence of the lighter lamellae is not due entirely to rod 
double refraction, as may be determined by examining sections in a 
graded series of liquids of varying indexes of refraction. Nor is the 



; completely masked t 



1935] BAILEY AND KERR, STRUCTURE OF THE SECONDARY WALL 281 

apparent isotropy of the intervening lamellae due solely to the masking 
effects of lignification or to the presence of other non-cellulosic con- 
stituents. The walls of immature unlignified cells show identical pat- 
terns and a similar differentiation into lamellae of two distinct cate- 
gories of birefringence, as do delignified cells that are treated for the 
removal of hemicelluloses and other non-cellulosic constituents. 

By subjecting untreated sections to the action of such swelling agents 
as acids, alkalies, chloro-iodide of zinc or cuprammonium hydroxide, and 
by carefully controlling the reactions, it is possible to expand the cen- 
tral layer and its constituent lamellae without distorting or seriously 
modifying the original structural pattern (compare Figs. 10 and 11). 
As the central layer expands and enlarges under the microscope, suc- 
cessively finer details of structure become visible. The lamellae are not 
discrete homogeneous entities, and are resolved during the expansion of 
the central layer into aggregations of elongated heterogeneous com- 
plexes of varying degrees of fineness which grade down to the limits of 
microscopic visibility. The darker lamellae are compact sheets of rela- 
tively isotropic material which contain a low ratio of birefringent com- 
plexes. On the contrary, the lighter lamellae are aggregations contain- 
ing a high ratio of birefringent complexes and a low ratio of apparently 
isotropic ones. There are no discontinuities in the structural pattern 
which is firmly knit together by lateral anastomoses and interlocking 

After treatment for the removal of non-cellulosic constituents, the 
purified cellulose exhibits a similar structural pattern, which upon swell- 
ing, Fig. 14, is resolved into a complex and firmly coherent matrix, hav- 
ing elongated, intercommunicating interstices of varying degrees of fine- 
ness. The darker and more compact parts of the matrix, which corre- 
spond to the lighter lamellae of Fig. 10, are strongly birefringent in 
longitudinal sections and show conspicuous dichroism when carefully 
stained with congo red or chloro-iodide of zinc ; whereas the lighter and 
more porous parts of the matrix, which correspond to the darker lamellae 
of Fig. 10, are so feebly birefringent that they appear to be compara- 

Conversely, when the central layer is freed of cellulose by treatments 
with 72 % sulphuric acid, the details of the swollen pattern are preserved 
in the so-called "lignin" residue, Fig. 11, which also is a complex and 
firmly coherent structure, having elongated, intercommunicating inter- 
stices of varying degrees of fineness. The lighter, finer residues of the 
originally birefringent lamellae exhibit well defined rod double refraction 
in longitudinal sections; whereas the darker, denser residues of the 
originally isotropic lamellae do not. 



282 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

It is evident from a detailed comparison of Figs. 11 and 14, that the 
denser parts of the "lignin" residue correspond to the more porous parts 
of the matrix of purified cellulose, and that the "lignin" residue may be 
interpolated within the interstices of the swollen cellulose. Furthermore, 
the rod double refraction of the lighter lamellae of the "lignin" residue 
suggests that the two interpenetrating complexes grade downward in size 
far below the limits of microscopic visibility. In other words, each of the 
visible parts of the original structural pattern is heterogeneous and com- 
posed of optically different complexes. Removal of either the "lignin" 
or the cellulose leaves a coherent matrix of varying texture and porosity. 
It is possible to reconstruct the structural pattern of the swollen cellu- 
lose from the "lignin" residue or vice versa, since they are positive and 
negative images of the same pattern. Although swollen sections of 
purified cellulose afford excellent preparations for visual examination, 
they are difficult objects for photographic reproduction. Therefore, a 
majority of our photomicrographs were made from "lignin" residues. 

The structural pattern of the central layer is not a constant; it varies 
greatly not only in different groups of plants, but also at times in 
homologous cells of the same plant, and even within the wall of a single 
cell. For example, in Siparuna bifida, the two optically different com- 
plexes may be segregated into coarsely radial patterns which are clearly 
visible in untreated sections, Fig. 10, or they may be diffused in finer 
radio-reticulate patterns, Fig. 16, the finest of which are invisible in 
unswollen sections of the secondary wall. In such cells, conspicuous 
concentricities usually are due either to abrupt changes in the texture of 
the structural pattern. Fig. 11, or to zones of varying intensities of 
lignification, Fig. 9. The former persist in purified cellulose; the latter 
are eliminated during delignification. 

Structural patterns of a basically concentric type are, however, of 
common occurrence in the normal tracheids of conifers, Fig. 18, and in 
the fiber-tracheids or libriform fibers of such dicotyledons as Poraqueiba 
sericea Tul., Fig. 15. In the central layer of these cells, the optically 
different complexes are segregated into concentric lamellae of varying 
widths and spatial groupings. The lamellae are of two types, i.e., 
strongly birefringent and comparatively isotropic. They are not dis- 
crete homogeneous entities, but may be resolved by treatment with 
swelling agents into complexes of varying degrees of fineness. As in 
the case of Siparum bifida, the darker lamellae are compact aggregates 
of relatively isotropic material. Figs. 15 and 18, and contain a low ratio 
of birefringent cellulose; whereas the alternating lighter lamellae are 
composed largely of birefringent cellulose and contain a low ratio of 



19351 BAILEY AND KERR, STRUCTURE OF THE SECONDARY WALL 283 

isotropic material. The structural pattern persists in delignified sec- 
tions which are treated with standard solvents of hemicelluloses and of 
other non-cellulosic constituents. When the purified cellulose is swollen, 
it appears as a complex and firmly coherent matrix, which exhibits a 
structural differentiation into compact, strongly birefringent and looser, 
comparatively isotropic lamellae. 

It should be emphasized, in this connection, that the concentric struc- 
ture of swollen cotton hairs — to which the work of Balls (2) has 
directed so much attention — appears to be due fundamentally to a 
similar structural pattern. When extremely thin, very smoothly cut 
sections of raw or purified cotton are treated with diluted Schweizer's 
reagent and are examined in polarized light between crossed nicols, the 
lamellae are, during the early stages of swelling, clearly of two optically 
different types, i.e., strongly birefringent and comparatively isotropic. 
During subsequent swelling, Fig. 17, the central layer is resolved into a 
complex and firmly coherent, spongy structure, the conspicuously bi- 
refringent parts of which are denser and obviously contain a higher ratio 
of cellulose than the more porous, intervening parts. In other words, 
the structural patterns of the central layers of cotton hairs, Fig. 17, of 
coniferous tracheids, Fig. 18, and of the fiber-tracheids of Poraqueiba 
sericea, Fig. 15, appear to be of a fundamentally similar type. In cotton 
hairs, as in tracheary cells, the width of the concentric lamellae is not a 
constant, but varies within relatively wide limits. 

The structural pattern of cotton can not be due to a segregation of 
cellulosic and non-cellulosic constituents, since the central layer of cotton 
is composed of practically pure cellulose — the low ratio of non-cellulosic 
constituents in cotton is confined chiefly to the so-called cuticle or pri- 
mary wall and to the lumen of the cell. Nor can the concentricities be 
due merely to inequalities in the penetration or modifying effects of the 
swelling agent, as may be demonstrated by cross-correlating the struc- 
tural patterns of different hairs from the same boll. For example, in 
Fig. 17, in passing outward from the lumen, there is the following 
sequence of lamellae: six narrow alternating light and dark zones, an 
unusually wide light zone, two broad dark zones separated by a narrower 
light zone, two narrow dark zones and three narrow light zones, and six 
broad dark zones separated by narrower light zones. The fact that this 
identical complex of varying concentricities occurs in other hairs from 
the same boll can not be due to purely fortuitous circumstances, but 
niight be due, either directly or indirectly, to the modifying effects of 
environmental factors upon the developing hairs. 

Nor can the structural patterns of tracheids, fiber-tracheids, and libri- 



284 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

form fibers be due to inequalities in the penetration and modifying effects 
of the swelling agents, since the patterns are visible under favorable 
conditions in untreated sections. Thus, the striking similarities in the 
finer visible structures of the central layer of unlignified and delignified 
cells and of "lignin" residues indicate that there are fundamental struc- 
tural differences in the underlying cellulose to which the pattern of lig- 
nification must more or less closely conform. 

Combinations of radial and concentric patterns of varying texture and 
complexity are of common occurrence in the fiber-tracheids and libri- 
form fibers of dicotyledons. ' In such cells there may be abrupt transi- 
tions within the central layer from coarse to fine texture and from 
radio-reticulate to concentric arrangements and vice versa. Fig. 19 is 
a transverse section of the wood of Tetramerista glabra Miq., cut with- 
out preliminary softening or other drastic treatments. It illustrates a 
type of complex radio-concentric structure which is clearly visible in 
unstained sections mounted in water and other liquids of varying indexes 
of refraction. The pattern is complicated, however, as is so often the 
case in cells of this type, by the presence of zones of varying intensities 
of lignification. A radio-concentric pattern of much finer texture is 
illustrated in Fig. 20. 

In the case of the more heavily lignified zones of such central layers, 
Figs. 9 and 20, both the birefringent and the comparatively isotropic 
parts of the structural pattern persist in "lignin" residues; whereas, in 
the less intensely lignified zones, the birefringent parts leave no struc- 
tural residue. It is of interest, in view of the significance that has been 
attached to the work of Freudenberg and his co-workers (12), that in 
longitudinal sections the residues of heavily lignified parts exhibit con- 
spicuous rod double refraction ; whereas the residues of the less intensely 
lignified parts do not.- In other words, there appear to be submicro- 
scopic structural differences in the two optically different complexes of 
the structural pattern which are reflected in their "lignin" residues. 
Furthermore, as previously noted, when delignified sections are stained 
with chloro-iodide of zinc or congo red, the strongly birefringent parts 
of the structural pattern may become markedly dichroic; whereas the 
more nearly isotropic parts do not. 

The observational and experimental data that we have assembled in 



strongly birefringent i: 
dark in polarized light 



1935] BAILEY AND KERR, STRUCTURE OF THE SECONDARY WALL 285 

our extensive survey of a wide range of gymnosperms and angiosperms 
indicate that the central layer of normal tracheids, liber-tracheids, and 
libriform fibers is composed, in all cases, of a complex and firmly 
coherent matrix of cellulose with elongated, intercommunicating inter- 
stices. Within these interstices more or less "lignin" and other non- 
cellulosic constituents may be deposited. The denser and more porous 
parts of the cellulosic matrix exhibit striking contrasts in birefringence, 
which are accentuated by lignification. Where these optically different 
parts are diffused in various patterns of fine texture — as is usually the 
case in the tracheids of conifers and in the fiber-tracheids and libriform 
fibers of many dicotyledons — the structural complexes are invisible in 
untreated sections of the secondary wall, but may be swollen to micro- 
scopically visible dimensions, Figs. 9, 12, 15, 16, 18, and 20. On the 
contrary, where the two optically different parts are segregated into 
coarser structural complexities. Figs. 10, 13, and 19, the patterns are 
clearly visible in unswoUen sections. 

The cellulosic matrix of the central layer is composed, in all cases, of 
anastomosing elongated complexes which are oriented parallel to the 
long axis of the cell or in a helical arrangement. In fact, it is these 
elongated complexes of two optically different types. Fig. 13, which 
give a longitudinally or helically striated appearance to the central layer 
and determine its helical or longitudinal planes of cleavage into so-called 
fibrils. In other words, fibrils are heterogeneous shredded parts of an 
originally continuous and coherent matrix. 

Although there are serious optical difficulties in studying the tenuous 
inner and outer layers of the secondary wall in sectional view, the striated 
appearance of these layers in surface view strongly suggests that they 
have similar structural patterns, the elongated, strongly birefringent 
complexes of which are oriented more nearly at right angles to the longi- 
tudinal axis of the cell. 



The orientation of the elongated complexes of the structural pattern 
may be relatively uniform throughout the central layer of tracheids, 
fiber-tracheids, and libriform fibers, or it may deviate more or less 
in successively formed parts of this layer. Not infrequently, the changes 
in orientation are correlated with fluctuations in the texture of the struc- 
tural pattern. Where the deviations are of considerable magnitude, 
they may be detected in polarized light, as illustrated in Fig. 4. The 
brilliant internal zones resemble the inner and outer layers of the 
secondary wall in having their birefringent complexes oriented more 



286 JOURNAL OF THE ARNOLD ARBORETUM [vol, xvi 

nearly at right angles to the longer axis of the cell, and therefore are 
bright in transverse sections. 

Although there is a superficial similarity between Fig. 4 and Fig. 6, 
the two cell walls are of a fundamentally different type. In the fibers of 
Pandanus, Fig. 6, as in the libriform fibers of various representatives of 
the Flacourtiaceae, Figs. 21 and 26, and in the bast fibers of ramie and 
of other dicotyledons, there are, as previously stated, actual discontinui- 
ties in the cellulosic matrix produced by narrow isotropic films of a non- 
cellulosic character. It should be emphasized, in this connection, how- 
ever, that the individual anisotropic zones of these multiple-layered 
fibers have complex structural patterns of the general types discussed 
on preceding pages. For example. Fig. 27 is a transverse section of the 
unswollen wall of Homalium luzoniense F. Villar. The layers of cellu- 
lose have a radio-reticulate pattern, the finer structural details of which 
are more clearly visible in swollen sections or in "lignin" residues, Fig. 
22. The elongated birefringent complexes of the structural pattern are 
oriented parallel to the long axis of the cell. Therefore the entire com- 
plex of layers is dark in polarized light in transverse sections. Coarsely 
radial patterns of the type illustrated in Fig. 10 are of not uncommon 
occurrence in the individual anisotropic layers of certain bast fibers; 
whereas in Pandanus, Fig. 7, the two optically different aggregates of 
cellulose are diffused in a pattern of unusually fine texture. Where the 
strongly birefringent complexes are oriented closely parallel to the 
longitudinal axis of the cell, the layer is dark in cross sections, Fig. 6, 
and merges with the truly isotropic film of non-cellulosic material; where 
they are oriented more nearly at right angles to the major axis of the cell, 
the layer is brilliant in transverse sections. 

Variations in the orientation of cellulose in successively formed parts 
of the secondary wall have a marked effect upon the swelling of tracheary 
cells and fibers. Owing to its specific anisotropy, the cellulose expands at 
right angles to the so-called fibrillar axis, and, during extensive lateral 
swelling produced by strong chemical reagents, actually contracts in a 
direction parallel to this axis. In the case of isolated, delignified 
tracheary cells and fibers having normal 3-layered secondary walls, the 
laterally expanding central layer frequently splits the tenuous, longi- 
tudinally expanding outer layer into a series of constricting rings and 
helical bands. Fig. 23, and bulges outward between these structures. 
This ring-bead type of swelling occurs in cotton and has received con- 
siderable attention in literature dealing with commercial fibers. Al- 
though the so-called cuticle or primary wall may aid at times in bead 
formation, the controlling factor in cotton hairs, as in tracheary cells 



1935] BAILEY AND KERR, STRUCTURE OF THE SECONDARY WALL 287 

and fibers, appears to be differences in orientation of cellulose in the 
outer and central layers of the secondary wall. There are no trans- 
verse plates of non-cellulosic material in the secondary wall which are 
concerned in ring-bead formation as hypothesized by Liidtke (21). 

In the case of multiple-layered tracheary cells and fibers, it is possible 
to verify conclusions based upon the study of cells of the 3-layered type. 
We have shown that the concentric anisotropic layers of various repre- 
sentatives of the Flacourtiaceae, Figs. 21, 22, and 26, are separated by 
films of non-cellulosic material, and that the orientation of the cellulose 
is constant except in the outermost layer of the secondary wall, where it 
is more nearly at right angles to the longitudinal axis of the cell. When 
such cells are partly or completely delignified and are swollen in cupram- 
monium hydroxide, the internal complex of anisotropic layers expands 
laterally and disrupts the tenuous outer layer into constricting rings, 
F'lg. 27, or helical bands. Fig. 24. The internal layers of cellulose — 
which may be slipped apart as shown in Fig. 26 — expand more or less 
in unison, Figs. 24 and 27, and no subsidiary internal constrictions are 
formed. 

On the contrary, in the multiple-layered fibers of Pandanus and of 
other monocotyledons — which have similar isotropic films of non-cellu- 
losic material, but where the orientation of the cellulose changes in the 
successively formed anisotropic lamellae — each of the narrow aniso- 
tropic layers, Fig. 6, may be disrupted by the lateral expansion of the 
broader layers, Fig. 7, and in the case of entire, delignified fibers, may 
give rise to constricting rings and helical bands. Fig. 25. In other 
words, the fiber behaves as if it were composed of several two-layered 
secondary walls, each of which swells in turn, forming similar ringed 
and beaded structures, Fig. 25. The two outermost layers swell first, 
the expansion working from the ends towards the center of the cell. 
The first formed ringlike constrictions commonly determine the position 
of subsequently formed internal constrictions. 

Multiple-layered fibers of the Pandanus type are of common occur- 
rence in the primary tissues of the stems of many monocotyledons. It is 
evident from Liidtke's (20, 22) figures and descriptions that the fibers 
of bamboo are of this structural type, and that they exhibit similar 
phenomena during their expansion in such swelling agents as cupram- 
monium hydroxide. It is obvious, in addition, that purely physical 
phenomena of swelling have been misinterpreted by Liidtke as evidence 
for the existence of transverse plates (Querelemente) of non-cellulosic 



JOURNAL OF THE ARNOLD ARBORETUM 

DISCUSSION 



The secondary walls of tracheary cells and fibers are extremely com- 
plex and variable structures. Therefore, it is misleading and fruitless 
to attempt to homologize all types of fibers in a single structural model. 
For example, there are five different types of visible concentricities, 

1. The segregation of two optically different aggregates of cellulose 
into concentric patterns. 

2. Abrupt changes in the form or texture of the structural pattern. 

3. Changes in the orientation of the elongated birefringent com- 
plexes of the structural pattern. 

4. Varying intensities of lignification or differences in the distri- 
bution of non-cellulosic constituents within the structural pattern. 

5. Alternation of cellulosic and non-cellulosic layers. 

In so far as we are able to judge from a study of a wide range of 
gymnosperms and angiosperms, most, if not all, tracheary cells and 
fibers exhibit more or less conspicuous concentricities of the third type, 
i.e., those due to changes in the orientation of the elongated birefringent 
complexes of the structural pattern, but the number and magnitude of 
the deviations in orientation are variable. Inability to detect such con- 
centricities appears to be due to inadequate techniques or to errors of 
interpretation. In most cases, the third type of layering occurs in asso- 
ciation with one or more of the other four types of concentricities. Thus, 
in the secondary wall of cotton hairs, it occurs with the first type; in 
the fiber-tracheids of Siparuna bifida, with the second and fourth types ; 
in the fiber-tracheids of Tetramerista glabra, with the first, second, and 
fourth types; in the fibers of Pandanus, with the first and fifth types, etc. 

Variations in the intensity of lignification or in the distribution of 
other non-cellulosic constituents may at times be closely correlated with 
changes in the orientation or the texture of the structural pattern. For 
example, the narrow inner and outer layers of the secondary wall may 
be more heavily lignified than the central layer or vice versa. Similarly, 
the coarser parts of the structural pattern of the central layer may be 
more heavily lignified or contain a higher ratio of hemicelluloses than 
the finer parts or vice versa. It is such fortuitous correlations as these 
which have led, in certain cases, to the unwarrantable conclusion that 
all types of visible heterogeneities in the secondary wall are due primarily 
to differences in chemical composition. 

There are investigators who believe that all fibers are composed of 
concentric lamellae of cellulose which are held together by non-cellulosic 



1935] BAILEY AND KERR, STRUCTURE OF THE SECONDARY WALL 289 

material. Thus, Ludtke (21, 22), who has attempted to homologize 
all types of fibrous cells in a single structural model, is of the opinion 
that the lamellae are separated by a "Fremdsubstanz" which differs from 
both cellulose and lignin in its chemical composition. Ritter (26) argues 
that it is possible to dissect the secondary wall by chemical means into 
concentric lamellae which may be slipped apart as shown in Fig. 26. 
Liidtke's conclusions appear to have been derived largely from a study 
of bamboo fibers; and Ritter's, from investigations of the libriform 
fibers of elm. We have shown that the anisotropic lamellae of mono- 
cotyledonous fibers frequently are separated by films of non-cellulosic 
material. The libriform fibers of elm are commonly of the so-called 
gelatinous type, which also are characterized in many cases by having 
both cellulosic and non-cellulosic lamellae. In such fibers, where there 
are actual discontinuities in the structural pattern of cellulose, the 
anisotropic lamellae may readily be separated by chemical treatments 
and slipped apart. On the contrary, in cotton hairs and in normal 
tracheids, fiber-tracheids, and libriform fibers, the entire matrix of cellu- 
lose is firmly coherent, and can be dissected only by forcibly tearing or 
rupturing the structural pattern. In Siparuna bifida the more obvious 
planes of weakness in the cellulosic matrix are radio-longitudinal or 
radio-helical; whereas in cotton hairs or in Poraqueiba sericea they are 
concentric-longitudinal. 

B. "Fibrils" and other "Units" of Cellulose 
Since the pioneer days of Von Mohl, Valentin, and Th. Hartig, a 
succession of investigators have visualized the secondary wall as com- 
posed of visible units of cellulose — elementary fibrils, dermatosomes, 
etc. — that are held together by non-cellulosic material. It is essential 
to understand the relationship between these units and the visible struc- 
tural patterns produced by different optical aggregates of cellulose. 

We have shown in Part I of this paper that the orientation of the 
cellulose is correlated with that of the so-called fibrillar structure, as 
has been demonstrated by analyses of X-ray diagrams, of anisotropy, 
of dichroism, and of other physical properties of the cell wall. How- 
ever, these physical correlations are concerned only with the orientation 
of the fibrillar structure and afford no conclusive evidence that fibrils 
obtained by chemical or mechanical treatments are discrete entities of 
constant length or cross sectional area. 

Ritter (27) has discussed the length of the so-called fibril and con- 
cludes that it is variable. He states that "although fibril segments of 
only 230 microns in length have been isolated, it seems that some may 



290 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

be as long or longer than the fiber." Liidtke (22), on the contrary, 
claims that the length of fibrils is determined by the presence and spac- 
ing of transverse plates of non-cellulosic material. Jancke, working 
with R. O. Herzog (15), measured the width of fibrils and obtained 
values of some 0.3-0.5 |j. Balls and Hancock (3), proceeding upon the 
assumption that lamellae' are composed of a single concentric row of 
fibrils, inferred that the width of both lamellae and fibrils in cotton is 
0.4 |j. Frey-Wyssling (13) tabulates the dimensions of fibrils as 0.4 X 
0.4 X 100 |j. 

Fibrils may be dissected by relatively drastic treatments with oxidiz- 
ing agents or acids into short segments which are variously designated 
as dermatosomes, spherical units, ellipsoid bodies, etc. According to 
Frey-Wyssling (13), dermatosomes have dimensions of 0.4 X 0.4 X 
0.5 |j; whereas Farr and Sisson (11) state that ellipsoid bodies pre- 
pared from cotton have axes of 1.1 p and 1.5 |.j. Ludtke (20) believes 
that dermatosomes are held together by his "Fremdsubstanz"; whereas 
Farr and Eckerson (9) maintain that the ellipsoid bodies of cotton are 
jacketed by a pectic cement. 

We have demonstrated in Part II that the central layer of tracheary 
cells and fibers is composed of an extremely complex and firmly coherent 
matrix of cellulose and that the details of the structural patterns of this 
matrix grade down to the limits of microscopic visibility. There is no 
evidence, either in untreated or in carefully swollen fibers, of discrete 
entities of cellulose, i.e., of fibrils or dermatosomes, which may be lib- 
erated simply by dissolving non-cellulosic constituents. The matrix of 
cellulose is shredded and disrupted during the production of fibrils and 
dermatosomes, which are heterogeneous fragments of larger size than 
the finer visible complexes of the structural pattern. In cotton, Fig. 17, 
as in Pinus, Fig. 18, Poraqueiba, Fig. 15, and Siparuna, Figs. 10 and 
14, the lamellae obviously are not composed of a single row of adherent 
fibrils, but are alternating layers of varying width, porosity, and bire- 
fringence. The finer, visible, elongated complexes of the lamellae are 
0.1 n or less in thickness. As indicated at (a) in Fig. 17, the cross sec- 
tional area of an ellipsoid body of the size postulated by Farr and Sisson 
covers more than four lamellae and a relatively large number of the 
finer visible complexes. 

■The form and size of the fragments which may be dissected from the 
secondary wall are clearly dependent upon the structural pattern of the 
matrix of cellulose, and upon the type and severity of the chemical and 



1935J BAILEY AND KERR, STRUCTURE OF THE SECONDARY WALL 291 

mechanical treatments to which the material is subjected. Splits or 
cracks develop in the more porous and weaker parts of the matrix, thus 
liberating the denser parts which contain a higher ratio of birefringent 
cellulose. In addition, there are submicroscopic, transverse, or oblique 
planes of cleavage, i.e., "slip planes," to which the work of Von Hohnel 
(16) and of Schwendener (29) has directed so much attention. It is 
these slip planes, rather than Liidtke's hypothetical "Querelemente," 
which facilitate the dissection of the fiber and of the elongated com- 
plexes of its structural pattern into shorter segments. 

after the action of 72 7o sulphuric acid upon longitudinal sections of 
fibers which yield coherent "lignin" residues. By the use of mechanical 
pressure during the initial stages of the action of the acid, the walls of 
tracheary cells, Fig. 12, and fibers may be resolved into long "lignin" 
threads, similar to fibrils. These shreds of the originally coherent frame- 
work of "lignin" may be dissected by more drastic chemical and mechan- 
ical treatments into nearly isodiametric fragments resembling dermato- 
somes. As previously stated, the amorphous non-cellulosic constitu- 
ents are deposited within the elongated, intercommunicating interstices 
of the cellulose matrix, resulting in two continuous, interpenetrating 
systems. Neither system is composed of discrete entities of visible 
dimensions, but each may be disrupted into fragments of varying size 
and form. If there are actual discontinuities in the systems, they must 
occur in the submicroscopic field, e.g., in the realm of micelles or of 
molecular chains. It should be emphasized, in addition, that so-called 
fibrillar structures are not visible in the secondary walls of parenchyma, 
of sclereids, or of other cells which exhibit statistical isotropy in surface 
view. The structural pattern of the cellulose matrix in such walls is of a 
fundamentally different type from that which occurs in fibrous cells. 

Dermatosomes, spherical units, and ellipsoidal particles are difficult 
to homologize, either as regards their size or their form. They are 
obtained by the action of oxidizing agents or of acids which tend to 
modify the cellulose. Neale (24) has summarized the modifying effects 
of oxidation and hydrolysis as follows: "The loss of strength and fall in 
viscosity which accompany the hydrolysis or oxidation of cellulose are 
quite irreversible, and the general term degradation is applied to these 
changes. The degradation of cellulose is accompanied by the appear- 
ance of chemical properties foreign to the original material. The 
hydrolysis of the glucoside-oxygen bridge causes the appearance of 
reducing sugar properties which may be quantitatively, though arbi- 
trarily, expressed as 'copper number' or 'iodine number.' The reducing 



292 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

sugar properties also arise as a result of oxidation and may be accom- 
panied by the development of acidic properties, so that oxidized cellu- 
lose may retain traces of caustic alkali or absorb basic dyes. This 
latter property has been put on the quantitative basis so essential in the 
chemistry of cellulose in the form of the methylene blue absorption test." 

Thus, it may be seen that the action of acids, which are supposed to 
dissolve some cementing substance and to liberate integral units of 
cellulose, may actually result in partial degradation of the cellulose. We 
have found that the staining of hydrocellulose and oxycellulose with 
ruthenium red is similar to the methylene blue absorption values, as 
listed by Doree (8). Ruthenium red behaves, in some respects, as a 
basic dye, and the staining of ellipsoidal particles, obtained by treating 
cotton with relatively strong acid (10), may be interpreted as an indi- 
cation of the degradation of the cellulose rather than as evidence for 
believing that the particles are coated with a pectic cement. Ruthenium 
red is not a specific test for pectic compounds, as botanists have fre- 
quently assumed. It is removed from dilute aqueous solutions by coagu- 
lated protoplasm and other nitrogenous substances, by gums, mucilages, 
hemicelluloses, oxycelluloses, hydrocelluloses, and certain lipoids, as 
well as by pectic compounds. 

Any hypothesis concerning the visible structure of the secondary wall 
must account not only for the varying structural patterns of a wide 
range of cells, but also for well known facts regarding the physical and 
chemical properties of cellulose. In the case of the hairs of the cotton 
plant, the constituents which do not yield glucose upon hydrolysis are 
small in amount, and are confined chiefly to the so-called cuticle or 
primary wall and to the lumen of the cell. There obviously is not a 
sufficient volume of cutinlike substances or of pectic compounds in the 
secondary wall to serve as a cementing substance of the type postulated 
by Liidtke (21) or by Farr and Eckerson (9). Furthermore, when 
cotton is treated with solvents of such constituents, without degrading 
the cellulose, the structural pattern is not affected. It persists as a firmly 
coherent matrix of cellulose. 

It is now generally admitted that the cellulose molecule is a long chain 
of glucose residues bound together by oxygen bridges. Furthermore, 
there is evidence from X-ray analyses, from anisotropy, dichroism, etc., 
to indicate that cellulose is built up of submicroscopic, crystal-like 
aggregates of these chains. The length of the cellulose chain and its 
arrangement within the crystallite are still subjects of dispute. Thus, 
it is uncertain whether the chain is shorter or longer than the crystallite 
or of equivalent length, and whether micelles are discrete and separate 
entities, or merely parts of a continuous system of overlapping chains. 



1935] BAILEY AND KERR, STRUCTURE OF THE SECONDARY WALL 293 

Estimates of the length of cellulose molecules range from 100-3500 
glucose residues. The highest value of 3500 units, i.e., that of Kraemer 
and Lansing (19), is based upon measurements of viscosity. Such 
molecules would have a length of approximately 1.8 |j, and would be 
visible microscopically if they were of sufficient thickness, which they 
obviously are not. Since the cellulose chains are arranged parallel to 
the so-called fibrillar orientation, and since there are no visible struc- 
tures which transect this axis, it is possible to conceive of chains of the 
length postulated by Kraemer and Lansing arranged in an overlapping 
manner along the fiber axis. 

Our investigations indicate that the cellulosic matrix of the secondary 
wall is composed of complexes of cellulose of varying birefringence which 
grade down to the limits of microscopic visibility, and that the funda- 
mental units of cellulose are of submicroscopic dimensions. In the case 
of cotton, the available chemical and physical data make it appear im- 
probable that the variations in birefringence are due to differences in 
chemical composition. Correns (5) recognized, more than 40 years ago, 
that cellulose is heterogeneous and attempted to explain the visible 
striations and certain types of lamellae as due to differences in water 
content. This hypothesis originated with Nageli {2?>), who postulated 
water rich and water poor layers as a means of explaining concentrici- 
ties and still permitting growth by intussusception. Differences in water 
content apparently do exist, and may be a factor influencing the inten- 
sity of birefringence in different lamellae. However, it is difficult to 
evaluate such differences by a study pf dried material. Drying the 
walls shrinks the cell so that structures just within the limits of micro- 
scopic visibility when the preparation is in water, may be contracted to 
invisible dimensions. Furthermore, differences in water content must 
be explained in terms of submicroscopic dii^erences in the cellulose which 
permit varying degrees of hydration. The question whether the varia- 
tion in birefringence of different complexes of the cellulosic matrix is due 
to fluctuations in the size, number, or orientation of submicroscopic 
units of cellulose is one which must be attacked from the physical and 
chemical, rather than from the botanical, side. 

C. Significance of Biological Variables in Physical and 
Chemical Investigations 

Our survey of a wide range of gymnosperms and angiosperms indicates 
that the secondary wall is a very complex structure, and that the struc- 
tural pattern of the cellulose matrix varies greatly, not only in different 
groups of plants but also, at times, in homologous cells of the same plant, 



294 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

and even in different parts of the same cell. There is a similar varia- 
bility in the distribution of "lignin" and of other non-cellulosic constitu- 
ents. Therefore, since all types of secondary walls can not be homolo- 
gized in a single structural model, there are grave dangers in generaliz- 
ing from intensive investigations of isolated species, e.g., cotton, spruce, 
bamboo, or ramie. 

Deductions concerning the structure of the cell wall based upon physi- 
cal or chemical analyses, should be checked by microscopic investigations 
and by accurate information concerning the numerous biological vari- 
ables. This is particularly necessary in the interpretation of X-ray 
diagrams, where the investigator of necessity must deal with complex 
aggregates of plant material. Van Iterson (31) has shown that certain 
misconceptions regarding Valonia might have been avoided by an 
acquaintance with the work of Correns (6) and others upon the visible 
structure of the walls of algae. Preston (25) undoubtedly errs in con- 
cluding, from an examination of X-ray diagrams, that there is a single 
plane of orientation of "fibrils" in the secondary wall of the' tracheids 
of Sequoia and of other conifers. Accurate interpretations of X-ray 
diagrams of growing cells and of differentiating tissues are especially 
difficult, and such conclusions regarding structural changes as those of 
Clark and Farr (4) and Ritter and Stillwell (28) must be carefully 
verified from the histological side. 

Although "lignin" residues of thick sections exhibit rod double re- 
fraction, as demonstrated by Freudenberg and his co-workers (12), a 
careful study of the residues of thin sections shows that a considerable 
proportion of the secondary wall "lignin" is isotropic. Similarly, there 
are parts of the cellulosic matrix which do not exhibit a clearly defined 
dichroism when thin sections are stained with chloro-iodide of zinc or 
Congo red. The woods of certain dicotyledons leave no coherent struc- 
tural residue when subjected to standard treatments with 72% sulphuric 
acid, as shown by Harlow (14); whereas others leave compact residues 
such as have been considered to be typical of conifers. In the wood 
of certain plants, the bulk of the "lignin" is confined to the so-called 
middle lamella, as Ritter (27) maintains; whereas in others, there is a 
relatively large proportion in the secondary wall. 

It should be emphasized, in conclusion, that most of our own data 
were obtained from a study of tracheary cells and libers, and that many 
additional types of cells must be investigated before it will be possible 
to visualize the full range of structural variability of the secondary wall. 
In a succeeding paper, we shall discuss methods that have been perfected 
for studying the small-celled, lightly lignified woods of dicotyledons of 
temperate regions. 



1935] BAILEY AND KERR, STRUCTURE OF THE SECONDARY WALL 295 

SUMMARY AND CONCLUSIONS 

1. An extensive survey of a wide range of gymnosperms and angio- 
sperms has shown that the structural pattern of the secondary wall is 
clearly visible in the large fiber-tracheids and libriform fibers of various 
dicotyledons. 

2. By using untreated sections of such cells as controls, it is possible 
to observe the exact effects of specific chemical and mechanical treat- 
ments upon normal structures, and thus to extend the scope of investi- 
gation to cover a wide range of less favorable material. 

3. The cellulosic matrix of the swollen secondary wall of cotton, as of 
normal tracheids, fiber-tracheids, and libriform fibers, is an extremely 
heterogeneous but firmly coherent structure, the finer details of which 
grade down to the limits of microscopic visibility. 

4. There is no reliable evidence to indicate that the matrix is com- 
posed of discrete entities of visible size — e.g., elementary fibrils, derma- 
tosomes, ellipsoidal bodies, etc. — that are bound together by non- 
cellulosic material. On the contrary, our data demonstrate that such 
putative entities actually are heterogeneous fragments that are shredded 
or disrupted from an originally continuous and coherent matrix. If 
there are discontinuities in the structural pattern of the cellulose in 
normal tracheary cells, they are confined to the submicroscopic field, 
e.g., to the realm of micelles or molecular chains. 

5. The visible structural pattern of the cellulosic matrix varies 
greatly in form and texture, not only in different plants, but also in 
homologous cells of the same plant, and even in different parts of the 
same cell. 

6. There are at least two optically different elongated complexes of 
cellulose which may be segregated into radio-helical, radio-longitudinal, 
or concentric-longitudinal lamellae, or into various radio-concentric 
patterns. 

7. The orientation of the elongated complexes of the structural pat- 
tern fluctuates more or less in successively formed parts of the secondary 
wall. In the case of normal tracheids, fiber-tracheids, and libriform 
fibers, there are three layers due to varying orientations: narrow inner 
and outer layers, in which the orientation is more nearly at right angles 
to the longitudinal axis of the cell, and a central layer of varying width, 
in which the orientation is parallel to this axis or does not deviate ex- 
cessively from it. 

8. "Lignin" and other non-cellulosic constituents may be deposited 
in the elongated, intercommunicating interstices of the cellulosic matrix, 
thus resulting in two continuous, interpenetrating systems. In heavily 



296 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

lignified forms, either system may be dissolved without seriously modi- 
fying the structural pattern of the remaining system. The purified 
cellulose and the "lignin" residue reveal positive and negative images of 
the original structural pattern. 

9. Deviations from the typical 3-Iayered type of secondary wall are 
of not infrequent occurrence. Thus, many thick-walled libriform fibers 
and fiber-tracheids have no clearly differentiated inner layer; whereas 
others have more than three layers of varying "fibrillar" orientation. 

10. Conspicuous discontinuities in the structural pattern of the 
cellulose commonly occur in the multiple-layered walls of so-called gela- 
tinous fibers, in certain types of bast fibers, and in sclereids. They are 
due to narrow layers of truly isotropic material which contain little, if 
any, cellulose. 

11. There are five different types of visible concentricities which 
occur in varying combinations, and may be associated at times with 
radio-helical or radio-longitudinal lamellae. Therefore, it is misleading 
and fruitless to attempt to homologize all types of fibers in a single struc- 
tural model. 

LITERATURE CITED 

1 . AsTBURY, W. T., Marwick, T. C, and Bernal, J. D. X-ray analysis 

of the structure of the wall of Valonia ventricosa. (Proc. Roy. 

Soc. London B, 109: 443-450. 1932.) 
Balls, W. L. The existence of daily growth rings in the cell wall 

of cotton. (Proc. Roy. Soc. London B, 90: 542-555. 1919.) 
Balls, W. L. and Hancock, H. A. Further observations on cell 

wall structure as seen in cotton hairs. (Proc. Roy. Soc. London B, 

93:426-440. 1922.) 
Clark, G. L. Cellulose as it is completely revealed by X-rays. (Ind. 

Eng. Chem. 22: 474-487. 1930.) 
Correns, C. Zur Kenntnis der inneren Structur der vegetabilischen 

Zellmembranen. (Jahrb. Wiss. Bot. 23: 254-338. 1892.) 
Zur Kenntnis der inneren Structur einiger Algenmem- 

branen. (Zimmermann's Beitr. Morph. Physiol. Pflanzenzelle, 

1:260-305. 1893.) 
DiPPEL. L. Das Mikroskop und seine Anwendung. (Braunschweig. 

Isted. 1867-69.) 
DoREE, C. The methods of cellulose chemistry. (London, 1933.) 
Farr, W. K. and Eckerson, S. H. Formation of cellulose membranes 

bv microscopic particles of uniform size in linear arrangement. 

(Contrib. Boyce Thompson Inst. 6: 189-203. 1934.) 
Farr, W. K. and Eckerson, S. H. Separation of cellulose particles in 



(Contrib. Boyce Thompson Inst. 6: 309-313. V. 
arr, W. K. and Sisson, W. A. X-ray diffractioi 
lose particles and interpretation of cellulose 
(Contrib. Boyce Thompson Inst. 6:315-321. 1' 



1935] BAILEY AND KERR, STRUCTURE OF THE SECONDARY WALL 297 

'REUDENitERG, K., ZocHER, H., and DuRR, W. Weitcre Versuche mit 
Lignin. XI. Mitteilung iiber Lignin und Cellulose. (Ber. Deutsch. 
Chem. Gesell. 62: 1814-1823. 1929.) 

'Rey-Wyssling, a. Die Stoffausscheidung der hoheren Pflanzen. 
(Berlin, 1935.) 

Iarlow, W. M. Contributions to the chemistry of the plant cell wall. 
II. Lignification in the secondary and tertiary layers of the cell 
walls of wood. (Tech. Pub. N. Y. State Coll. Forestry, no. 24. 



den Einfluss des Rindendruckes auf die 
Beschaffenheit der Bastfasern der Dikotylen. (Jahrb. Wiss. Bot. 
15:311-326. 1884.) 

Katz, J. R. Micellartheorie und Quellung der Zellulose. (In Hess, 
K. Die Chemie der Zellulose und ihrer Begleiter. Leipzig, 1928.) 

Kerr, T. and Bailey, I. W. The cambium and its derivative tissues. 
X. Structure, optical properties, and chemical composition of the 
so-called middle lamella. (Jour. Arnold Arb. 15: 327-349. 1934.) 

Kraemer, E. O. and Lansing, W. D. The molecular weight of cellu- 
loses and cellulose derivatives. (Amer. Chem. Soc, Div. Cellulose 
Chem. N. Y., 1935.) Symposium on the nature of cellulose. 

LiJDTKE, M. Zur Kenntnis der pflanzlichen Zellmembran. I. (Liebig's 
Annalen, 466: 27-57. 1928.) 

Untersuchungen iiber Aufbau und Bildung der pflanzlichen 

Zellmembran und ihrer stofflichen Komponenten. (Biochem. 
Zeitschr. 233: 1-57. 1931.) 

Werden und Organisation der pflanzlichen Zellmembran. 

(Protoplasma, 22: 457-488. 1934.) 

Nageli, C. Ueber den inneren Bau der vegetabilischen Zellenmem- 

branen. (Sitzber. Bay. Akad. Wiss. Miinchen, 1:282-323; 2: 

114-171. 1864.) 
Neale, S. M. The modification of natural cotton cellulose by swelhng 

and degradation. (Trans. Faraday Soc. 29: 228-238. 1933.) 
Preston, R. D. The organization of the cell wall of the conifer 

tracheid. (Phil. Trans. Roy. Soc. London B, 224: 131-174. 1934.) 
RiTTER, G. J. Composition and structure of the cell wall of wood. 

(Ind.Eng. Chem. 20:941-945. 1928.) 

Structure of the cell wall of wood fibers. (Paper Industry, 

16: 178^183. 1934.) 

RiTTER, G. J. and Stillwell, C. W. Rate of formation of the crys- 
talline structure of wood fibers. (Paper Trade Jour. 98(22) : 37-40. 
1934.) 

Schwendener, S. Ueber die Verschiebungen der Bastfasern im 
Sinne Von H5hners. (Ber. Deutsch. Bot. Gesell. 12:234-248. 
1894.) 

Van Iterson, G. Die wording van den plantaardigen celwand. 
(Chem. Weekblad, 24: 166-187. 1927.) 

Biologische inleidung tot het cellulose symposium. (Chem. 

Weekblad, 30:2-19. 1933.) 

Wiesner J. Untersuchungen iiber die Organisation der vegetabili- 
schen Zellhaute. (Sitzber. Akad. Wiss. Wien, 93(Abt. 1): 17-80. 



298 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

DESCRIPTION OF PLATES 
Fi^s. 1-6 and 13 were made from unstained sections and were photo- 
graphed in polarized light between crossed nicols. All the remaining 

photomicrographs were made with an arc-light and Zettnow's filter. Figs. 

1-^ 8, 10, and 21 were made from sections mounted in diaphane (N = 

1.47). 

Plate 140 

Fig. 1 . Myodocarpus simplicifolius Brong. & Oris. Transverse section 
of the xylem, showing a fiber-tracheid and parts of seven adjoin- 
ing ones. The thick secondary walls are composed of three 
distinct layers : a narrow brilliant outer layer, a brilliant narrow 
inner layer, and a wide intervening dark layer. In a section of 
this thickness, 15 n, the isotropic intercellular substance and the 
feebly anisotropic primary walls are more or less completely 
fogged or obscured by the brilliant outer layers of the secondary 
walls (compare Fig. 3 for a section 5 (a in thickness). X 1750. 

Fig. 2. Urandra corniculata Foxw. Radial longitudinal section of the 
xylem, showing the walls of adjacent fiber-tracheids in sectional 
view. The broad central layers of the secondary walls are bril- 
liant. The intercellular substance, the feebly birgfringent 
primary walls, and the inner and outer layers of the secondary 
wall are dark. A bordered pit is shown in the center of the 
photomicrograph. X 1750. 

Fig. 3. Trochodendron arcdioides Sieb. & Zucc. Transverse section of 
the xylem, showing a tracheid and parts of seven adjoining cells. 
In a section of this thickness, 5 fx, the outer brilliant layers of 
the secondary walls of adjacent cells are clearly separated by a 
narrow intervening dark layer, which actually consists of two 
feebly birefringent primary walls and a truly isotropic layer of 
intercellular material. X 1400. 

Fig. 4. Myodocarpus simplicifolius. Transverse section of the xylem, 
showing a fiber-tracheid and parts of seven adjoining ones. The 
thick secondary wall of the central cell consists of a series of 
alternating brilliant and dark layers. X 1750. 

Fig. 5. Urandra corniculata. Thick secondary wall of a sclerenchyma- 
tous cell in sectional view, showing alternating brilliant and 
dark layers. X 1750. 

Plate 141 

lignified fibers, showing secondary walls composed of regularly 
alternating brilliant and dark layers. X 1150. 

Fig. 7. The same. Transverse section of a fiber after standard treat- 
ment with 72% sulphuric acid, staining with Haidenhain's hae- 
matoxylin, and mounting in balsam, showing residue of sec- 
ondary wall. The brilliant layers of Fig. 6 are split and em- 
bossed. X 1300. 

Plate 142 

Fig. 8. Siparuna bifida (P. & E.) A. DC. Transverse section of a 
fiber-tracheid and of parts of several adjoining cells, stained 
with Haidenhain's haematoxylin and safranin, showing zones of 
varying intensities of lignification. X 2000. 

Fig. 9. The same. Transverse section of a fiber-tracheid after standard 
treatment with 72% sulphuric acid, staining with Haidenhain's 



19351 BAILEY AND KERR, STRUCTURE OF THE SECONDARY WALL 299 

haematoxylin and mounting in aniline oil, showing finely radio- 
reticulate pattern and zones due to varying intensities of ligni- 
fication. Dark zones heavily lignified, light zones less intensely 



! secondary wall is radially striated. X 2000. 
e same. Transverse section of a fiber-tracheid after standard 
atment with 72% sulphuric acid, staining with Haidenhain's 
i oil, showing radially stri- 



of lignification. X 1900. 

Plate 144 

Fig. 12. Tetramerista glabra M\q. Tangential longitudinal section of the 
central layer of a fiber-tracheid after treatment with 72% sul- 
phuric acid, staining with Haidenhain's haematoxylin, and 
mounting in aniline oil, showing longitudinal pattern of fine 
anastomosing threadlike components. The longitudinal orienta- 
tion has been somewhat distorted during swelling. X 1900. 

Fig. 13. Siparuna bifida. Tangential longitudinal section through the 
central layer of a fiber-tracheid mounted in water and photo- 
graphed with polarized light between crossed nicols, showing 
alternating birefringent and isotropic striae. X 1900. 

Fig. 14. The same. Transverse section of a delignified fiber-tracheid, 
after treatment with diluted cuprammonium hydroxide and stain- 
ing with Congo red. The denser, darker radii of the purified 
cellulose correspond to the lighter radii of Figs. 10 and 11. 







Plate 145 






Mg. 15. Por 


■aqiieiba se. 


ricea Tul. Transverse se 


ction of the s 


econdary 




1 of a fiber 


-tracheid after standard tr 


■eatment with ! 


72% sul- 




Tie acid. 


staining with Haidenhaii 


a's haematoxv 


'lin, and 


m" 




niline oil, showing concen 


trically lamelk 


ited resi- 


due 


of the ceni 


:ral layer. X 3200. 






Mg. 16. Sip, 


arnna btfid< 


J. Transverse section of t 


he secondary ^ 


kvall of a 


fibe 




ifter standard treatment w 


ith 72% sulpht 


aric acid. 


stai 


nin^wkh ' 


Haidenhain's haematoxylii 




line 


oil, showi 




\l^\dnToi thf '" """' 




lave 


;r. X 3200 


Plate 146 







after swelling with diluted cuprammonium hydroxide and stain- 
ing with Congo red, showing alternating lamellae of varying 
width and porosity in the 'nner part of the secondary wall. A 
particle 1 ^ in diameter in the untreated wall would expand to 
the size of the circle at (a). X 1200. Owing to swelling, the 
original width of the lamellae has been increased 7500 times in 
this photomicrograph. 



300 JOURNAL OF THE ARNOLD ARBORETUM [vou xvi 

Fig. 18. Finns ponderosa Dougl. Transverse section of the secondary 
wall of a tracheid after treatment with 72% sulphuric acid, stain- 
ing with Haidenhain's haematoxylin, and mounting in aniline 
oil, showing concentrically laminated residue of the central layer. 
X 1900. 

Plate 147 

Fig. 19. Tctramerista glabra. Transverse section of a fiber-tracheid and 
of parts of several adjoining cells, mounted in a dilute aqueous 
solution of iodine potassium iodide and photographed with a 
Zeiss 70-water-immersion lens. The broad central layer has a 
coarsely radio-concentric pattern which is complicated by zones 
of varying intensities of lignification. X 2000. 

Fig. 20. The same. Transverse section of a fiber-tracheid after treat- 
ment with 72% sulphuric acid, staining with Haidenhain's 
haematoxylin, and mounting in balsam, showing finely radio- 
concentric pattern and broad zones due to varying intensities of 
lignification. X 1300. 

Plate 148 

Fig. 21 . Homalium lusonietise F. Villar. Transverse section of a libri- 
form fiber stained with Haidenhain's haematoxylin and saf ranin, 
showing alternating broad cellulosic and narrow non-cellulosic 
layers. The radio-reticulate structure of the former layers is 
vaguely visible. X 3200. 

Fig. 22. The same. Transverse section of a libriform fiber after standard 
treatment with 72% sulphuric acid, staining with Haidenhain's 
haematoxylin, and mounting in aniline, showing residue of both 
the cellulosic and the non-cellulosic layers. X 1300. 

Plate 149 

Fig. 23. Rhisophora mangle L. Isolated, delignified, libriform fiber, 
swollen in diluted Schweizer's reagent, showing beadlike swell- 
ing of the central layer of the secondary wall. The outer layer 
of secondary wall is resolved into a series of constricting rings 
and helical bands. X 650. 

Fig. 24. Olmcdiella Betschleriana (Goepp.) Loes. Isolated, delignified, 
libriform fiber, swollen in diluted Schweizer's reagent. The 
outer layer of the secondary wall is resolved into constricting 
helical bands. X 325. 

Fig. 25. Pandanus odoratisstmus. Isolated delignified fiber, swollen in 
diluted Schweizer's reagent, showing that each of the internal 
brilliant layers in Fig. 6 may be resolved into constricting rings 
and helical bands. X 650. 

Fig. 26. Olmediella Betschleriana. Segment of a libriform fiber iso- 
lated from a thick transverse section of the xylem after delig- 
nification and treatment with 50% sulphuric acid. The con- 
centric cylinders of cellulose are slipping apart. X 650. 

Fig. 27. Olmediella Betschleriana. Isolated, delignified, libriform fiber, 
swollen in diluted Schweizer's reagent. The outer layer of the 
secondary wall is resolved into a seri< 
helical bands. X 400. 




Structure of the Secondary Wall 




VisiHLE Structure of the Secondary W 



Jour. Arnold Arb. Vol. XVI 




Jour. Arnold Arb. Vol. XVI 




Visible Stri 



. Arnold Arb. Vol. XVI 











Visible Structui 



Jour. Arnold Arb. Vol. XV] 






•>.-s. 



-^7 



■ -1 
, — *t- 



v^ ^ I 



Li'''', 






Wv 



: Sfxondary VVali 



SAX, TEMPERATURE AND NUCLEAR DIFFERENTIATION 301 



With one text figure and plate 150 

"Heredity is effected by the transmission of a nuclear preformation 
which, in the course of development, finds expression in a process of 
cytoplasmic epigenesis" (Wilson, 1925). The evidence from genetic 
and cytological investigations has proven conclusively that nuclear pre- 
formation is dependent upon the genie constitution of the chromosomes. 
The mechanism of expression in cytoplasmic epigenesis is more obscure. 
The problem is difficult because it is not subject to direct attack. A 
comparison of induced and hereditary effects has provided a method for 
studying certain developmental processes in Drosophila. An analysis of 
the effect of temperature on developing microspores has provided some 
information regarding nuclear cytoplasmic relations in differentiation 
and development, and has some bearing on the problem of genie ex- 
pression. 

Normal microspore development in Tradescantia has been described 
in detail by Sax and Edmonds (1933). The young microspore contains 
a centrally located nucleus surrounded by cytoplasmic granules. The 
granules disappear, and the nucleus migrates to the end of the oval- 
shaped microspore. The cytoplasm is massed around the nucleus, and 
at the other end of the cell there is a large vacuole. There is then a 
migration of cytoplasm and vacuole so that two vacuoles are formed, 
one at each end of the cell. Most of the cytoplasm lies between the vacu- 
oles so that the longer axis of the cytoplasmic mass lies in the short axis 
of the cell. The nucleus at this time lies toward the heavy or dorsal wall 
of the microspore, — originally the inner wall at the time of tetrad 
formation. When the nucleus divides, the daughter nucleus near the 
heavy wall of the microspore is enclosed by a thin temporary wall which 
includes little cytoplasm. This nucleus does not pass into the typical 
resting stage, but retains its chromaticity and finally elongates to form 
the generative nucleus. The other nucleus formed near the center of 
the cytoplasmic mass enlarges to form the inactive tube nucleus. Shortly 
after the division of the microspore nucleus, the vacuoles disappear, and 
the cytoplasm appears to be rather homogeneous. 

The normal development of the microspore of Pseudolarix amabilis 



302 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

resembles that of Pinus, described in detail by Coulter and Chamberlain 
(1901). Soon after the release of the microspore from the wall of the 
microsporocyte, the wings develop rapidly on opposite ends of the spore 
towards the ventral side of the cell, while the dorsal side of the spore, 
which was formed during meiosis, becomes thickened. At the time of 
the first nuclear division, the nucleus lies near the dorsal wall surrounded 
by most of the cytoplasm, and the region towards the wings and the 
ventral side of the spore are more vacuolate. The first division occurs 
across the short axis of the microspore, and the nucleus near the dorsal 
wall is cut off and degenerates. The other nucleus divides again in the 
same axis, and another prothallial cell is cut off. The third division 
produces the generative nucleus and the tube nucleus of the mature 
pollen grain. The generative nucleus lies near the inner or dorsal wall 
of the pollen grain and is cut off by a thin wall which encloses little 
cytoplasm, while the large tube nucleus lies free near the center of the 
cell. 

THE EFFECT OF TEMI'ERATUKE ON NUCLEAR 
DIFFERENTIATION 
Both high and low temperatures are effective in producing abnormal 
development in the microspores of Tradescantia. Plants were placed in 
a constant temperature chamber where the temperature was maintained 
at about 6°C. for the cold treatment, or at about 35°C. for the heat 
treatment. Three days' treatment was sufficient to produce abnormal, 
development at either temperature range, and doubtless a shorter time 
would be effective at the higher temperature. The microspores were 
examined soon after exposure to abnormal temperatures, and for several 
subsequent days after they had been placed in the normal greenhouse 



Three types of abnormalities were produced. Under normal condi- 
tions, only two nuclei are formed in the pollen grain,— the generative 
nucleus and the tube nucleus. The tube nucleus normally does not 
divide and is inactive in further development. The heat treatment 
occasionally causes the tube nucleus to divide in Tradescantia (Figs. 
1 and 2). The subsequent fate of the daughter nuclei is not known. 
Apparently no great deviation from normal environment is required to 
produce a second division, since this behavior was observed by Mr. R. H. 
Goodwin in Tradescantia plants grown in the greenhouse at the Bio- 
logical Institute of Harvard University. 

The second type of abnormality is produced by either cold or heat 
treatment. The polarity of the microspore is disturbed so that the 
nuclear division is no longer oriented across the short axis of the cell. 



1935] SAX, TEMPERATURE AND NUCLEAR DIFFERENTIATION 303 

In extreme cases the division is at right angles to the normal axis, the 
daughter nuclei are not differentiated, and the temporary cell wall is 
formed across the center of the microspore ( Fig. 3 ) . In most cases there 
is partial differentiation of the daughter nuclei, but the more compact 
nucleus does not elongate to form the typical generative nucleus (Fig. 
4). The degree of differentiation of the two nuclei is closely associated 
with the angle of division. With a smaller degree of variation from the 
normal axis of division there is increased differentiation of the genera- 
tive nucleus (Fig. 5). In a single anther all degrees of differentiation 
are found, including the normal condition (Fig. 6). 

In order to determine more accurately the relation between the angle 
of division and the differentiation of the nuclei, a statistical study was 
made. All microspores measured were from a single flower taken from a 
plant which had been kept at a temperature of about 36° C. for three 
days. Camera lucida drawings were made of 163 microspores selected 
at random among those which showed the two nuclei in the same focal 
plane. The angle between the normal axis of division across the short 
diameter of the microspore and the line drawn through the centers of 
the two nuclei was taken as the angle of division. The length of the 
outline of the "generative" nucleus was measured in millimeters. This 
work, as well as the tabulation and analysis of the data, was done by 
my wife, Dr. Hally Jolivette Sax. The relation between the angle of 
division and the length of the generative nucleus is shown in Table 1. 
The high correlation of —.81 ± .02 shows that the degree of nuclear 
differentiation is closely associated with the angle of division. 

A third type of aberrant development was found in microspores which 
began to "germinate" before the division of the nucleus. In some cases 
one of the daughter nuclei was found in the original microspore and the 
other in the newly-formed outgrowth (Figs. 8 and 10). In these cases 
a thin cell wall divided the cytoplasm into appVoximately equal parts, 
and there was no indication of nuclear differentiation. The division may 
occur so that neither daughter nucleus remains in the original micro- 
spore (Figs. 7 and 9). If the division is oriented lengthwise of the cyto- 
plasmic mass, there is no nuclear differentiation, but if it is oriented 
across the short diameter of the outgrowth, so that one daughter nucleus 
is near the cell wall, there is a differentiation which resembles that fol- 
lowing normal diyision in a normal microspore (Fig. 7). 

The precocious growth of the microspores of Tradescantia is unlike 
normal pollen-tube growth. The pollen-tube usually grows from the 
end of the pollen grain adjacent to the heavy dorsal wall, while the 
aberrant outgrowth occurs at the ventral side of the microspore. Per- 



304 



JOURNAL OF THE ARNOLD ARBORETUM 



haps this abnormal growth is the first stage in the development of an 
embryo-sac-like structure such as Stow (1930, 1933) has found in the 
anthers of Hyacinthus. Unfortunately, a study of the further develop- 
ment of the abnormal growth in Tradescantia microspores could not be 
continued because the anthers dehisced and disintegrated so soon. Pos- 
sibly these peculiar microspores could be developed further in a nutrient 

TABLE I. 





OF 


THE Nuc 


.EI IN MiCROb 


PORES OF Trades 


CANTIA 








Angle of 


iivision 






g 

1 







20 30 


40 50 60 


70 8 


) 90 


11 

14 


1 


1 2 
1 3 1 


1 5 3 


3 


\ 9 


1 


17 

23 
26 
29 

35 
38-40 




1 
1 


1 I 1 


^ 


1 



The development and differentiation of the microspores of Pseudo- 
larix amabilis is also affected by environmental conditions. Branches 
containing male flowers were placed in a warm corner of the greenhouse 
for about two weeks. During this time meiosis occurred, and the micro- 
spores developed to maturity. Most of the microspores were normal in 
their development (Figs. 11 and 12), but various types of abnormalities 
were observed. If the first division occurs lengthwise of the cell in the 
axis of the wings, there is no differentiation of the nuclei if each is an 
equal distance from the cell wall (Fig. 13). If, however, one nucleus 
lies near the cell wall, regardless of the orientation of the division 
spindle, this nucleus tends to remain small and form a prothallial cell, 
while the nucleus near the center of the cytoplasmic mass remains large 
and divides again (Figs. 14 and 16). The first division may occur in 
the normal position, but the second division may be aberrant in orienta- 
tion (Figs. 15 and 16). Several years ago Mrs. Sax found a mature 
pollen grain of Picea which contained four undifferentiated nuclei of 



1935] SAX, TEMPERATURE AND NUCLEAR DIFFERENTIATION 305 

approximately equal size. Apparently relatively slight changes in en- 
vironmental conditions can cause abnormal development of conifer 
microspores. Nuclear differentiation in these microspores appears to be 
entirely dependent upon the orientation of the division spindles and the 
position of the nuclei in relation to the cytoplasmic mass. 

Nuclear differentiation in the microspores of Tradescantia and Pscu- 
dolarix appears to be determined by the nuclear cytoplasmic relation- 
ships. In Tradescantia it is possible to observe the relations of nuclei, 
vacuoles, and cytoplasm in the living microspores. Observations at vari- 
ous stages of development show that normal development is dependent 
upon the synchronization of cytoplasmic and nuclear activities. 

In the microspores which develop under normal conditions, the cell 
contents show a gradual shifting in position before the nucleus divides. 
A large vacuole is formed at one end of the microspore, and the nucleus 
and most of the cytoplasm move to the opposite end. The vacuole 
then extends towards the opposite end, near the ventral side of the 
spore, and finally forms two vacuoles, one at either end of the cell. 
Meanwhile the cytoplasmic mass and the nucleus migrate towards the 
center of the cell. The cytoplasm extends between the ventral and 
dorsal walls so that the length of the cytoplasmic mass is across the 
short diameter of the cell. Some cytoplasm extends around the entire 
periphery of the cell. The nucleus lies near the dorsal wall at the time 
of division. After the division the nucleus near the dorsal wall is cut 
off by a thin temporary wall, and then develops into the elongated gen- 
erative nucleus. The other nucleus enlarges, loses its chromaticity, and 
becomes the inactive tube nucleus. The sequence of early development 
of the normal microspore is shown in text figures a, b, c, and d, which 
are camera lucida sketches drawn from living material. 

When the microspore develops at low temperatures, the same cycle of 
development begins, but the nucleus divides before the vacuole, cyto- 
plasm, and nucleus reach their normal positions. (Text figure e.) Since 
the cytoplasm migrates towards the center of the cell along the dorsal 
wall of the microspore, the length of the cytoplasmic mass is at an angle 
to the normal axis of division. The nuclear spindle is oriented in the 
long axis of the cytoplasmic mass, and the daughter nuclei lie towards 
one end of the microspore and do not undergo complete differentiation. 

The exposure to high temperatures for several days appears to accel- 
erate the cytoplasmic movement without causing a corresponding activity 
of the nucleus. At the time the cell constituents are in the position 
usually associated with nuclear division (Text figure c), the nucleus 
may remain inactive. The vacuoles then become smaller or may dis- 



306 JOURNAL OF THE ARNOLD ARBORETUM [vou xvi 

appear entirely before nuclear division. As a result, the long axis of the 
cytoplasmic mass is not oriented in the short iixis of the cell, and the 
division may occur at various angles, depending in part upon the cyto- 
plasmic distribution (Text figures f, g, and h). 

In a single flower, following heat treatment, the angle of division may 
vary from to 90 degrees. Camera lucida drawings were made from a 
random sample of these celK, and the angle of division was determined 
in relation to the di>tribution of cytoplasmic mass. The length of the 




Text Figure. Dexclopincnt of the microspoio under nomial and al)- 
normal conditions. 

All figures are from camera lucida drawings of living microspores. 

Figures a, b, c, and d show the movement of the cell contents during 
early development of the microspore under normal conditions. The vacuole 
becomes extended along the ventral wall and finally forms two vacuoles. 
The nucleus at the time of division becomes oriented near the dorsal or 
heavy wall of the microspore. 

Figure e shows the nuclei formed by nuclear division before normal 
orientation of the cell constituents is attained. This microspore developed 
at a low temperature, which seems to retard cytoplasmic movement with- 
out retarding nuclear division. 

Figures f, g, and h are drawings of microspores which had been sub- 
jected to a high temperature. The vacuoles are small or absent at the time 
of nuclear division, and the axis of division tends to occur in the long axis 
of the cytoplasmic mass. 

cytoplasmic mass was determined for the long axis of the cell, and the 
width was measured across the short axis of the cell. For example, in a 
normal microspore (fig. d) the length of the cytoplasmic mass is the 
distance between the vacuoles, and the width is the distance across the 
microspore in the axis of division, and the angle of division is very small. 
In this cell the length-width ratio is about 0.5. In figure e the length- 
width ratio is about 1.0, and the angle of division is about 45 degrees, 
while in figure h the length-width ratio of the cytoplasmic mass is about 
2.0, and the angle of division is about 80 degrees. The length-width 



1935] SAX, TEMPERATURE AND NUCLEAR DIFFERENTIATION 307 

ratio of the cytoplasmic mass correlated with the angle of division gave 
a value of r = .66 ± .03. It is evident that there is a strong tendency 
for the nucleus to divide in the long axis of the cytoplasmic mass, al- 
though as the volume of cytoplasm increases, there is not a correspond- 
ing tendency for the nucleus to divide in the longer axis. In general, 
however, the direction of division in the. microspore is controlled by the 
distribution of the cytoplasm in accord with Hertwig's rule, and the 
nuclear differentiation is controlled by the position of the daughter 
nuclei in relation to the cytoplasmic mass. 

It is not possible to follow the cytoplasmic movements in the living 
cells of conifers, but judging from the description of normal develop- 
ment (Ferguson, 1904) and the behavior of the nuclei in abnormal micro- 
spores, the failure of normal differentiation is also based on the dis- 
turbed relations of nucleus, cytoplasm, and vacuoles. 

A comparison of nuclear differentiation in Tradescantia and Psetido- 
larix microspores and in the embryo sacs derived from microspores in 
Hyacinthus shows a good deal of similarity in polarity. In both 
Tradescantia and Pseudolarix the center of activity in early microspore 
development is near the dorsal wall which was formed during micro- 
sporogenesis. If a second division occurs in the Tradescantia micro- 
spore, the nucleus nearer the center of the cell divides, as is the case in 
normal microspore development in the conifers. The vegetative nucleus 
is always the one nearer the ventral wall and is surrounded by a large 
amount of cytoplasm, while the generative or sexual nucleus lies near 
the dorsal wall and is enclosed by a thin temporary cell wall which in- 
cludes little cytoplasm. The "embryo sacs" which develop from micro- 
spores of Hyacinthus (Stow, 1933) show the exine of the microspore at 
the egg or sexual end of the embryo sac, while the polar or vegetative 
nuclei lie in the center of the embryo sac, apparently surrounded by a 
relatively large amount of cytoplasm. 

Stow was able to induce embryo sac-like structures in anthers of 
Hyacinthics by subjecting the bulbs to a temperature of 28 °C. for 18 to 
24 hours at the time of planting in the fall. The abnormal development 
observed in the following spring may have been induced either by the 
temperature treatment or by the effects produced by the large number 
of degenerating sterile microspores. At any rate the differentiation of 
the microspore to produce a normal pollen grain or an embryo sac appears 
to depend upon environmental conditions. We are inclined to believe 
that the precocious growth of Tradescantia microspores is the first stage 
in embryo sac formation, and that the complete structure could be de- 
veloped, under temperature control, if the cells could be kept alive over 
a long period of time, as is the case in Hyacinthus. 



308 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

FACTORS IN DEVELOPxMENT AND DIFFERENTIATION 

According to Osterhout (1921), life is dependent upon a series of 
reactions which normally proceed at rates which bear a definite relation 
to each other. If for life we substitute development and differentiation, 
we have an hypothesis which seems to explain development and differen- 
tiation. Certainly the differentiation of the microspore nuclei appears 
to be dependent upon the relative rates of cytoplasmic migration and 
nuclear activity. Less direct evidence indicates that the difference be- 
tween a pollen grain and an embryo sac may be dependent upon the same 
type of timing relationships. If sex can be determined by the timing 
relationships of different reactions, effected either by environmental con- 
ditions or hereditary factors, we have indirect evidence that genie ex- 
pression may be effected by differential reactions. 

More direct evidence regarding the mechanism of genie expression is 
found in the behavior of the chromosomes at meiosis. A failure of 
chromosome pairing, or asynapsis, may be caused by genetic factors or 
by environmental conditions. Genetic asynapsis has been found in 
Drosophila, Zea, Triticum, Rumex and Datura. Induced asynapsis can 
be effected in Rhoeo and Datura by subjecting the plants to low tem- 
peratures for several days, and it has been obtained in Tradcscantia 
following treatment at low and high temperatures. Both the hereditary 
and induced effects are similar in their expression. The chromosomes 
are unpaired at meiosis, the divisions are irregular, diploid gametes may 
be produced, and there is a high degree of pollen sterility in the asynaptic 
plants. Both types of asynapsis may be attributed to the same cause. 
If chromosome development is not synchronized with other cell activi- 
ties, the chromosomes may not be effectively paired before nuclear divi- 
sion is initiated. 

An exceptionally clear case of the timing factor in genie activity was 
found in Aquilegia by Anderson and Abbe (1933). The "compacta" 
mutant of Aquilegia is dependent on a single genetic factor. In the 
mutant type the branches are more erect and numerous, and the flowers 
are upright from the beginning. The dwarf type is caused by the preco- 
cious thickening of the cell walls, and the somatic expression is simply 
the result of disturbed timing relationships, — "the precocious initia- 
tion of a normal feature of normal development." Further aspects of 
the relation between genes and development in Drosophila have been 
discussed by Schultz (1935) and by Goldschmidt (1935), and similar 
work is being done on the cucurbits by Sinnott (unpublished). 

The temperature chambers used in this work were paid for, in part, 
by a grant from the American Academy of Arts and Sciences. 



19351 SAX, TEMPERATURE AND NUCLEAR DIFFERENTIATION 309 

SUMMARY 
The subjection of Tradescantia plants to low and high temperatures 
may produce three kinds of abnormalities in microspore development. 
( 1 ) The tube nucleus, which in normal microspores is inactive and ulti- 
mately degenerates, may divide. (2) The polarity of the cell may be 
disturbed so that the division of the microspore nucleus is not oriented 
in the normal axis. The angle of deviation is closely correlated with 
the differentiation of the daughter nuclei. (3) The microspore may 
become greatly extended on the ventral side, and the nuclear division 
may occur in this new outgrowth. This abnormality may be the first 
step in the transformation of a microspore to an "embryo sac," as found 
in Hyacintkus by Stow. 

When Pseudolarix microspores are developed at a relatively high 
temperature, there is a failure of normal differentiation of the nuclei. 
The differentiation of prothallial cells and generative and tube nuclei is 
dependent upon the nuclear cytoplasmic relationships in the developing 
microspore. 

The normal differentiation in Tradescantia microspores is dependent 
upon the synchronization of cytoplasmic movements and nuclear 
activity. There is some evidence that many differences in development 
and differentiation, induced either by genetic factors or by environ- 
mental conditions, are dependent upon differences in reaction rates of 
different processes. 

LITERATURE CITED 
Anderson, E. and L. B. Abbe (1933). A comparative anatomical study 

of a mutant Aquilegia. (Amer. Nat. 67: 380-384.) 
Coulter, J. M. and C. J. Chamberlain (1901). Morphology of sperma- 

tophytes. (D. Appleton and Co. New York.) 
Ferguson, M. C. (1904). Contributions to the knowledge of the life 
history of Pinus, with special reference to sporogenesis, the develop- 
ment of the gametophytes, and fertilization. (Proc. Wash. Acad. Sci. 
6: 1-202.) 
GoLDSCHMiDT, R. (1935). Gen und Ausseneigenschaft. (Zeit. f. Ind. 

Abstam. u. Vererb. 69: 38-131.) 
Osterhout, W. J. V. (1921). The mechanism of injury and recovery of 

the cell. ( Science N. S. 53 : 352-356. ) 
Sax, Karl and H. W. Edmonds (1933). Development of the male game- 

tophyte in Tradescantia. (Bot. Gaz. 94: 156-163.) 
ScHULTZ, J. (1935). Aspects of the relation between genes and develop- 
ment in Drosophila. (Amer. Nat. 69: 30-54.) 
Stow, I. (1930). Experimental studies on the formation of the embryo- 
sac-like giant pollen grain in the anther of Hyacinthus orientalis. (Cyto- 
logia, 1:417-^39.) 

(1933). On the female tendencies of the embryosac-like giant 

pollen grain of Hyacinthus orientalis. (Cytologia, 5: 88-108.) 
Wilson, E. B. (1925). The cell in development and heredity. (The Mac- 
millan Co. New York.) 



JOURNAL OF THE I 



DESCRIPTION OF PLATE 150 



Photographs of aceto-carmine preparations of abnormal microspores of 
Tradescantia and normal and abnormal microspores of Fscudolarix ama- 
hilis. Figures 1 to 6 inclusive, magnified X 800. The other figures are 



he microspore nucleus has occurred at right 

angles to the normal axis of division, and the daughter nuclei do not 

become differentiated. 
Figures 4, 5, and 6. The nuclear divisions have occurred at various angles 

followed by a corresponding amount of nuclear differentiation. 

These microspores were developed at a high temperature. 
Figures 7, 8, 9, and 10. Abnormal microspores produced by heat and cold 



PSEUDOLARIX 

Figures 11 and 12. Stages in the normal development of the microspore. 
All divisions are across the short axis of the cell, and the prothallial 
cells are always cut off near the heavy dorsal wall. 

Figures 13, 14, 15, and 16. Abnormal development induced by heat treat- 
ment. The nuclei may divide at various angles followed by various 
degrees of differentiation of the daughter nuclei. The prothallial 
cells may be cut off at any point along the cell wall. 



Jour. Arnold Arb. Vol. XVI 



^h 



^ 3 • 



t'., 



f .# .. I • 



(:^^'\'- f A 



-ill--- 



Temperature and Nuclear Difi 



19351 REHDER, LIGNEOUS PLANTS DESCRIBED BY LEVEILLE 



LABIATAE 

Leucosceptnim sinense Hemsley in Jour. Linn. Soc. Bot. 26:310 
(1890). — Leveille, Fl. Kouy-Tcheou, 209 (1914). — Dunn in Not. 
Bot. Card. Edinb. 8: 171 (1913); 6: 192 (1915). 

Elsholtsia Cavaleriei Leveille & Vaniot in Fedde, Rep. Spec. Nov. 

8:424 (1910). 
Leucosceptnim Bodinieri Leveille in op. cit. 9: 224 (1911). 

China. K w e i c h o u : environs de Tsin-gay, au bord d'une 
riviere, E. Bodinier, no. 2709, Sept. 20, 1899 "sous -arbrisseau, fl. roses" 
(syntype of Elsholtzia Cavaleriei; photo, in A. A.); environs de Tou- 
chan, /. Cavalerie in herb. Bodinier, no. 2 710, Sept. 1899 (syntype of 
Elsholtzia Cavaleriei; photo, in A. A.). 

The name Elsholtzia Cavaleriei was changed by Leveille to Leuco- 
sceptrum Bodinieri and later the two type specimens were enumerated 
by him in his Flore du Kouy-Tscheou under L. sinense without citation 
of synonymy, the reduction being based on identifications made by 
Dunn, to whom Leveille had sent material of his Labiatae for revision, 
as it appears from a note in Leveille's Flore du Kouy-Tcheou p. 203 
under Labiacees which reads "(D. Dunn revisit)." 

Leucosceptnim plectranthoideum (Levi.) Marquand in Kew Bull. 
Misc. Inform. 1930: 207. 

Buddlcia plcctranthoidea Leveille, Cat. PI. Yun-Nan, 171 (1916). 

China. Yunnan : patures des montagnes a Pe-long-tsin, 3200 
m., E. E. Maire, Nov. 1912 (holotype of Buddleia plectranthoidea ; 
merotype in A. A.). 

This species seems nearest to L. sinense Hemsl. but can be at once 
distinguished by the shorter inflorescence, the yellowish closer tomentum 
of the calyx and the bracts, and the shorter elliptic or ovate-elliptic to 
oblong-elliptic leaves reticulate beneath and tomentulose above. 

Bot. 14: 



312 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

Leveille, Cat. 111. Seu-Tchouen, 92, pi. 44 (1918). — P'ei, Verben. China 
in Mem. Sci. Soc. China, l(no. 3): 180 (1932). 

Colquhounia elegans Wall. var. pauciflora Prain in Jour. As. Soc. 
Beng. 62:38 (1893). — Dunn in Not. Bot. Gard. Edinb. 6:179 
(1915). 
Caryopteris fluminis Leveille, Sert. Yunn. 3 (1916) ; Cat. PI. Yun- 
Nan, 298 (1917). — P'ei, Verben. China in Mem. Sci. Soc. China, 
l(no. 3): 180 (1932). 
China. K w e i c h o u : environs de Ou-la-gay (Tchin-lin), /. 
Seguin in herb. Bodinier, no. 2237, March 1898 "longues tiges sous-lig- 
neuses, lianeuses" (syntype of C. Seguini; photo, in A. A.); rives du 
fleuve Bleu, alt. 450 m., E. E. Maire, June 1912, "petit arbuste, feuilles 
persistantes, fleurs roses" (holotype of Caryopteris fluminis; photo, in 
A. A.). Yunnan: environs de My-tsao, Fr. Ducloux, no. 110, 
March 4, 1897, "long tiges s'enlagant aux arbres et buissons, fleurs coc- 
cinees, 4 graines ailees" (syntype of C. Seguini; photo, in A. A.). 

By Dunn (I. c.) this species was referred to C. elegans var. pami- 
flora Prain, but as I pointed out in 1916 (1. c.) this variety is best con- 
sidered a distinct species which becomes C. Seguini Vaniot. 

Micromeria biflora Bentham, Labiat. 378 (1834). — Dunn in Not. 
Bot. Gard. Edinb. 6: 157 (1915). — Leveille, Fl. Kouy-Tcheou, 210 
(1914); Cat. PI. Yun-Nan, 138 (1916). 

Thymus Cavaleriei Leveille in Fedde, Rep. S'pec. Nov. 11: 298 (1912). 

China. K w e i c h o u : Tin-Ian, montagnes sablonneuses, J. 
Cavalerie, no. 3778, "blanc-rose" (syntype of Thymus Cavaleriei; photo, 
in A. A.). Yunnan: Pan-pien-kai, paturages des coteaux calcaires, 
alt. 2550 m., E. E. Maire, Sept. 1911 "Thymus vivace, etale, fl. roses" 
(syntype of Thymus Cavaleriei; photo, in A. A.). 

Elsholtzia nigulosa Hemsley in Jour. Linn. Soc. 26: 278 (1890). — 
Leveille, Fl. Kouy-Tcheou, 208 (1914); Cat. PI. Yun-Nan, 138 
(1916). — DunninNot. Bot. Gard. Edinb. 6: 149 (1915). 

Elsholtzia Labordci Vaniot in Bull. Acad. Intern. Geog. Bot. 14: 177 
(1904). 



China. K w e i c h o u : enviro 


ns de Tsin-gay, a Tchao-see, 


abonde dans la mont. et bord de routes, 


/. Laborde in herb. Bodinier, no. 


2711, Sept. 7, 1899 "fleurs d'un bleu tre: 


s pale" (holotype of £. Labordei; 


photo, in A. A.). 





Elsholtzia fruticosa (D. Don) Rehder in Sargent, PI. Wilson. 3: 381 
(1916). 

Elsholtzia polystachya Bentham, Labiat. 116 (1832). — Dunn in Not. 



1935] REHDER, LIGNEOUS PLANTS DESCRIBED BY LEVEILLE 31J 

Bot. Card. Edinb. 8:161 (1913); 6:149 (1915). — Leveille, Fl. 
Kouy-Tcheou, 208 (1914) ; Cat. PI. Yun-nan, 138 (1916). 
Elsholtsia tristis Leveille in Fedde, Rep. Spec. Nov. 8: 424 (1910). 
Elsholtzia Dielsii Leveille in op. cit. 9: 441 (1911). 
Elsholtsia Souliei Leveille in op. cit. 9: 248, non p. 218 (1911). 
China. Szechuan : Ta-tsien-lu, /. A. Soulie, nos. 781 and 
1023, in 1893 (syntypes of E. Dielsii [E. Souliei Levi. p. 248, non p. 
218J ; photos, in A. A.). K vir e i c h o u : environs de Kouy-yang, 
mont du College, E. Bodinier, no. 1944, Nov. 3, 1897, "tige 1 m., fleurs 
blanches" (holotype of E. tristis, photo, in A. A.). 

Elsholtzia ochroleuca Dunn in Not. Bot. Card. Edinb. 8: 161 
(1913). 

Elsholtzia lampradena Leveille in Bull. Geog. Bot. 25: 25 (1915) ; Cat. 
PI. Yun-Nan, 137 (1916).— Synon. nov. 
China. Yunnan : paturages des collines a Tong-tchouan, alt. 
2600 m., E. E. Maire, Sept. 1912 "arbrisseau rameux, haut 0.40 m., fleurs 
blanches en epis dresses" (holotype of E. lampradena ; photo, in A. A.). 
Elsholtzia lampradena has been identified with E. ockroleuca accord- 
ing to a note on the type specimen. 

Pogostemon glaber Bentham in Wallich, PI. As. Rar. 1:31 
(1830). — Leveille, Cat. PI. Yun-Nan, 143 (1916). 

Caryoptcris Esquirolii Leveille in Fedde, Rep. Spec. Nov. 9 : 449 
(1911); Fl. Kouy-Tcheou, 440 (1915). — Synon. nov. 
China. K w e i c h o u : Tchou-ly, alt. 900 m., /. Esquirol, no. 
2053, March 1, 1910, "fl. blanche, labelle rose" (holotype of Caryop- 
teris Esquirolii; merotype in A. A.). 

Pogostemon glaber has not yet been recorded from Kweichou, as far 
as I know, but it is known from Yunnan. 

Plectranthus ternif oHus Don, Prodr. Fl. Nepal. 117 (1825).— 
Leveille, Fl. Kouy-Tcheou 214 (1914); Cat. PI. Yun-Nan 143 
(1916). — Dunn in Not. Bot. Gard. Edinb. 6: 138 (1915). 

Elsholtzia Lychnitis Leveille & Vaniot in Fedde, Rep. Spec. Nov. 

8:425 (1910). 
Teucrium Esquirolii Leveille in Bull. Geog. Bot. 22: 236 (1912). 
China. Kweichou : Tchen-lin-tcheou, route de Lo-pie a Ou- 
la-gay, L. Martin in herb. Bodinier, no. 1937, Oct. 9, 1899, "tige de 1.50 
m. de haut, fleur blanches, ou blanc-bleuatre" (syntype of Elsholtzia 
Lychnitis; photo, in A. A.) ; route de Tou-tcheou a Pien-yang, /. Cava- 
lerie, no. 2573, Nov. 1905 (syntype of E. Lychnitis; photo, in A. A.); 
coteaux de Lo-fou, /. Esquirol, no. 2576, Nov. 1910 (holotype of 
Teucrium Esquirolii; ex Leveille). 



314 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

Of Teucrium Esquirolii I have seen no specimen, but Leveille enu- 
merates it in 1916 (Cat. PI. Yun-Nan, 143) as a synonym of Plec- 
tranthus ternijolius, probably on identification by S. T. Dunn, though 
Dunn does not cite it in 1915. It does not appear in the Flore du Kouy- 
Tcheou. Plectranthus ternijolius should probably not be classed as a 
ligneous plant, though Hooker describes it in his Flora of British India 
(4: 621) as a bush 3-5 ft. high. 

Plectranthus coetsa Hamilton ex Don, Prodr. Fl. Nepal. 117 
(1825). — Leveille, Cat. PI. Yun-Nan, 141 (1916). 

Plectranthus Mairci Leveille, Cat. PI. Yun-Nan. 141 (1916), pro 
synon. P. coetsa Ham. 

China. Yunnan: paturages des mont. derriere Tong-tchouan, 
2700 m., E. E. Maire, Oct. (1912-13), "plante annuelle tres-rameuse, 
fl. rouge vif" (type of P. Mairci; photo, in A. A.) ; haies, plaine de Tche- 
hay, 2550 m., E. E. Mairc, Sept. [ 1912-13 ] , "plante vivace, tumescente, 
en touffes, haut 1.40 m." (in herb. Leveille with P. Mairei; photo, in 
A. A.). 

Plectranthus Mairei is apparently an unpublished name and is cited 
only as a synonym of P. coetsa (1. c). On Mairei's specimen from 
Tong-tchouan the name Plectranthus Mairei appears in Leveille's hand- 
writing; the specimen from Tche-hay is without any name, but placed 
in the cover of P. Mairei. 

The majority of Labiatae described by Leveille are herbaceous and 
most of them have been examined by S. T. Dunn. He published his 
identifications in his Notes on Chinese Labiatae (in Not. Bot. Card. 
Edinb. 8: 153-171. 1913) and in his Key to the Labiatae of China 
(op. cit. 6: 127-208. 1915). The identifications and reductions made 
by Dunn were accepted by Leveille and incorporated in his Flore du 
Kouy-Tcheou (p. 203-217) and in his Catalogue des plantes de Yun- 
Nan (p. 136-149). 

SOLANACEAE 

Solanum aculeatissimum Jacquin, Coll. 1:100 (1786); Ic. Rar. 
1: t.41 (1781-86). — Merrill in Contr. Arnold Arb. 8: 149 (1934). 
Solaumn Bodinicri Leveille & Vaniot in Bull. Soc. Bot. France, 55: 

206 (1908). 
Solanum Cavalcrici Leveille & Vaniot, 1. c. 207 (1908). — Synon. nov. 

China. Hongkong: plage sablonneuse de File Verte, E. 
Bodinier, July 31, 1895 (holotype of 5. Bodinieri; photo, in A. A.). 
K w e i c h o u : Ly-po-hien, /. Cavalerie in herb. Bodinier, no. 2722, 
Aug. 10, 1899 (holotype of S. Cavaleriei; photo, in A. A.). 

Solanum Bodinieri represents a glabrescent form of this very variable 



1935] REHDER, LIGNEOUS PLANTS DESCRIBED BY LEVEILLE 315 

species, while 5. Cavaleriei is much more pubescent throughout. 
Solanum Bodinieri has been identified with 5. aculeatissimum by Mer- 
rill (I.e.). 

SCROPHULARIACEAE 
Brandisia racemosa Hemsley in Kew Bull. Misc. Inform. 1895: 114. 
ncnt::iafuncbris'Liwe\\\i, Sert. Yunn. 1 (1916). — Cat. PI. Yun-Nan, 
296 (1917).-Synon.nov. 
China. K w e i c h o u : rives du fleuve Bleu a Kiang-pien, alt. 
350 m., E. E. Maire, Aug. 1913, "arbuste un peu grimpant; fl. roses" 
(holotype of Deutzia junebris ; merotype in A. A.). 

BIGNONIACEAK 
Incarvillea Delavayi Bureau & Franchet in Jour, de Bot. 5: 138. 
(1891). 

Tecoma Mairei Leveille, Cat. PL Yun-Nan, 20 (1916). — Synon. nov. 
China. Yunnan: rochers sous brousse, mont. de Pe-long-tsin, 
alt. 3200 m., E. E. Maire, May 1911, "plante vivace, fl. roses grandes" 
(holotype of Tecoma Mairei; photo, in A. A.). 

Though this is an herbaceous species, I am including it in this enu- 
meration, because Leveille has described it under the ligneous genus 
Tecoma. 

ACANTHACEAE 
Phlogacanthus pubinervis T. Anderson in Jour. Linn. Soc. Bot. 
9: 508 (1867). — Leveille, Cat. PI. Yun-Nan, 6 (1915). 

Acschynauthus Dunnii Leveille in Fedde, Rep. Spec. Nov. 9:453 

(1911); Fl. Kouy-Tcheou, 180 (1914). — Synon. nov. 
Lonicera Menelii Leveille, El. Kouy-Tcheou, 63 (1914). — Sjmon. 

China. K w e i c h o u : without precise locality, /. Esquirol, no. 
737, "fleur rougeatre" (syntype of Aeschynanthus Dunnii; photo, in 
A. A.); Lo-fou, /. Cavalerie, no. 3475, March 1909 "couleur jaunatre" 
(syntype of Aeschynanthus Dunnii, in fruit; photo, in A. A.); Thing- 
mei, 1100 m., /. Esquirol, no. 3540, Dec. 2, 1913 (holotype of Lonicera 
Menelii; merotype in A. A.). 

Cystacanthus yangtsekiangensis (Levi.), comb. nov. 
Strobilanthes yangtsekiangensis Leveille, Cat. PI. Yun-Nan, 7 (1915). 

China. Yunnan: rives du fleuve Bleu, alt. 400 m., E. E. Maire, 
May 1912, "plante sous-ligneuse en touffes dressees, fl. bleues" (syntype 
of Strobilanthes yangtsekiangensis, photo, in A. A.); rives du fleuve 
Bleu a Ta-tchai, alt. 450 m., "plante vivace, sous-ligneuse, en touff"es, fl. 
roses" (syntype of S. yangtsekiangensis ; photo, in A. A.). 



316 JOURNAL OF THE ARNOLD ARBORETUM [vou xvi 

This species is very near C. yunnanensis W. W. Sm., but is easily dis- 
tinguished by the closer and finer pubescence of the young branchlets, 
the inflorescence and the calyx, and by the numerous lateral 2-6- 
flowered inflorescences along last year's branches. To C. yangtse- 
kiangensis apparently belong Rock 8049 from Yunnan, between Tang- 
yueh and Likiangfu, and Schneider 671 from southern Szechuan, between 
Mo-so-ying and Kung-mu-ying. 

RUBIACEAE 

Oldenlandia Bodmieri(Levl.) Chun in Sunyatsenia, 1: 310 (1934). 

Hedyotis Bodinicri Leveille in Fedde, Rep. Spec. Nov. 11: 64 (1912). 

China. Kwangtung: Tay-mo-chan, sommet de la mont., 

3500 ft., E. Bodinier, no. 1158, May 7, 1895, "petit sous-arbrisseau 

croissant dans les rocailles, fleurs blanches" (holotype; photo, in A. A.). 

I have not been able to identify this plant with any described species. 

Oldenlandia macrostemon (Hook. & Am.) Kuntze, Rev. Gen. 1: 

292 (1891). — Pitard in Lecomte, Fl. Gen. Indo-Chine, 3: 138 (1922). 

Hedyotis macrostemon Hooker & Arnott. Bot. Beechey Voy. 192 

(1841). — Leveille, Cat. PI. Yun-Nan, 245 (1917). 
Hedyotis Esquirolii Leveille in Fedde, Rep. Spec. Nov. 13: 176 

(1914) ; Fl. Kouy-Tcheou, 367 (1915). — Synon. nov. 
Oldenlandia Esquirolii (Levi.) Chun in Sunyatsenia, 1:310 (1934). 
China. K w e i c h o u : without precise locality, /. Esquirol 
(holotype of H. Esquirolii; photo, in A. A.). 
Ophiorrhiza japonica Blume, Bijdr. 978 (1826). 
Ophiorrhiza Cavaleriei Leveille in Fedde, Rep. Spec. Nov. 13: 177 

(1914). 
Ophiorrhiza Labordei Leveille 1. c. (1914); Fl. Kouy-Tcheou, 370 
(1915). — Synon, nov. 
China. K w e i c h o u : district de Tsin-gay, rocailles, bois a 
Kao-tchay, /. Laborde in herb. Bodinier, May 15, 1898 (holotype of O. 
Labordei; photo, in A. A.); without locality, J. Cavalerie (holotype of 
O. Cavaleriei; ex Leveille). 

Ophiorrhiza Cavaleriei is cited by Leveille in his Flore du Kouy- 
Tcheou as a synonym of O. Labordei, but Cavalerie's specimen is not 
cited, only Laborde's specimen from Tsin-gay. Cavalerie's specimen I 

Ophiorrhiza cantoniensis Hance in Ann. Sci. Nat. ser. 4, 18: 222 
(1862). — Leveille, Fl. Kouy-Tcheou, 370 (1915). 

Ophiorrhiza Seguini Leveille in Fedde, Rep. Spec. Nov. 13: 177 
(1914). — Sjmon. nov. 



1935] REHDER, LIGNEOUS PLANTS DESCRIBED BY LEVEILLE 317 

Ophiorrhiza violaceo-flammea Leveille in Bull. Geog. Bot. 25 : 47 
(1915); Cat. PI. Yun-Nan, 247 (1917). — Synon. nov. 

China. K w e i c h o u : environs de Gan-pin, croissant dans les 
rocailles, les trous entre des rochers, E. Bodinier, no. 1549, April 29, 
1897, ^'fleurs blanches" (holotype of O. Seguini, photo, in A. A.). 
Yunnan: vallee de Li-tse-pin, 2700 m., E. E. Maire, April 1912, 
"sous-arbrisseau toujours vert, fl. violettes" (holotype of O. violaceo- 
flammea; photo, in A. A.). 

Ophiorrhiza Seguini is not mentioned by Leveille in his Floie du 
Kouy-Tcheou, but its type is enumerated, together with another speci- 
men, under O. cantoniensis, which shows that Leveille had reduced it 
to O. cantoniensis. The color of the flowers on the specimen of O. 
violaceo-flammea said to be violet does not look at all different from the 
color of flowers of O. japonica. 

Ophiorrhiza cantoniensis is closely related to O. japonica Bl., but it 
may be distinguished by its oblong leaves attenuate at the base and 
quite glabrous, broadest at or above the middle, while O. japonica has 
shorter generally ovate or oblong leaves less attenuate or even nearly 
rounded at base and puberulous on the midrib beneath. 

Wendlandia li^strina Wallich, Num. List. 6272 (1832), in part, 
nom. nud. — Don, Gen. Syst. 2: 518 (1834). — Leveille, Cat. PI. Yun- 
Nan, 242 (1917). — Cowan in Not. Bot. Gard. Edinb. 16: 242 (1932); 
18: 183 (1934). 

Luculia gratissiymi Sw. sensu Leveille, Fl. Kouy-Tcheou, 368 (1915), 
non Sweet (1826). 

China. K w e i c h o u : bords du Hoa-kiang, L. Martin in herb. 
Bodinier, no. 2563, Feb. 18, 1899, "petit arbuste, fleurs blanches" 
(photo, in A. A.). 

This collection extends the range of W. Ugustrina into Kweichou. 
Martin's specimen was identified with W. Ugustrina by J. M. Cowan 
according to a note on the specimen. 

Wendlandia salicifolia Franchet in herb, ex Castello in Jour, de Bot. 
9: 208 (1895). — Cowan in Not. Bot. Gard. Edinb. 16: 244 (1932). 
Ligustrum Thea Leveille & Dunn in Fedde, Rep. Spec. Nov. 10: 147 
(1911). — Leveille, Fl. Kouy-Tcheou, 295 (1914). 

China. Kweichou : without precise locality, J. Esquirol, no. 
327, Dec. 16, 1904, "sous-arbrisseau des bords du fleuve, submerge aux 
grandes eaux; les feuilles donnent une infusion theiforme assez em- 
ployee; fleur blanc-rose" (holotype of Ligustrum Thea; photo, in 
A. A.) ; without precise locality, J. Esquirol, no. 239 (cited in Fl. Kouy- 
Tcheou; photo, in A. A.). 



318 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

Ligustrum Thea was first referred to W. salicijolia by Cowan in his 
"The Genus Wendlandia" (op. cit. 233-316). 

Wendlandia Cavaleriei Leveille in Fedde, Rep. Spec. Nov. 10: 434 
(1912); Fl. Kouy-Tcheou, 373 (1915). — Cowan in Not. Bot. Card. 
Edinb. 16: 263 (1932). 

IVcndlandia Feddci Leveille in Fedde, Rep. Spec. Nov. 10:434 
(1912) ; Fl. Kouy-Tcheou, Z7Z (1915). 

China. K w e i c h o u : Lo-fou, J. Cavalerie, no. 3297, April 
1907, "fleurs blanches" (holotype of W. Cavaleriei; merotype in A. A.) ; 
route de Pin-fa a Lo-fou, /. Cavalerie, no. 2732, April 4, 1906 (holotype 
of W. Feddei; photo, and merotype in A. A.). 

Wendlandia uvariifolia Hance subsp. Dunniana (Levi.) Cowan in 
Not. Bot. Card. Edinb. 16: 287 (1932); 18: 185 (1934). 

Wendlandia Dunniana Leveille in Fedde, Rep. Spec. Nov. 10:434 
(1912) ; Fl. Kouy-Tcheou, 373 (1915). 
China. K w e i c h o u : Lo-fou, /. Cavalerie, no. 3476, March 
1908 (holotype of W, Dunniana; merotype in A. A.). 

Wendlandia long-idens (Hance) Hutchinson in Sargent, PI. Wilson. 
3:392 (1916). — Cowan in Not. Bot. Card. Edinb. 16: 301 (1932). 

China. Yunnan: a mi-mont de Siao-ho, alt. 2800 m., E. E. 
Maire, [1911-14] "arbuste buissonant, fl. roses" (in herb. Leveille sub 
Leptodermis Mairei; duplicate in A. A.). 

In the herbarium Leveille there were in the cover of Leptodermis 
Mairei three specimens collected by Maire, of which two belonged to 
L. Mairei which was identified by Dr. H. Winkler as L. pilosa (Franch.) 
Diels var. glabrescens H. Winkl., while the third was not a Leptodermis, 
but represents the species cited above. From the Szechuan and Hupeh 
specimens before me it differs slightly in the smaller leaves not exceeding 
14 mm. and somewhat more densely pubescent on both sides; the flow- 
ers also are slightly smaller and are rose-colored according to the col- 
lector, while Wilson under his numbers 3756 (Veitch Coll.) and 2359 
gives the color of the flowers as white. Possibly the specimen cited by 
Cowan (1. c.) as Maire, no. 326, without locality, is of the same collec- 
tion as the specimen cited above. 

Emmenopterys Henryi Oliver in Hooker's Icon. 19: t. 1823 (1889). 
Mussacnda Cavaleriei Leveille in Fedde, Rep. Spec. Nov. 13: 178 

(1914); Fl. Kouy-Tcheou, 368 (1915). — Hutchinson in Sargent, 

PI. Wilson. 3: 397 (1916).— Synon. nov. 
Mussaenda Mairei Leveille in Bull. Geo^-. Bot. 25: 47 (1915); Cat. 

PI. Yun-Nan, 247 (1917). — Synon. nov. 



1935] REHDER, LIGNEOUS PLANTS DESCRIBED BY LEVEILLE 319 

China. K w e i c h o u : sur des rochers a Touan-po pres Pin-yue 
(Pin-ue), /. Cavalerie, no. 2481, Aug. 10, 1905, "petit arbre a fl. 
blanches" (holotype of Mussaenda Cavaleriei; merotype in A. A.). 
Yunnan: vallee de Long-ky, 700 m., E. E. Maire, July 1912, 
"arbre moyen, fl. blanches" (holotype of Mussaenda Mairei; merotype 
in A. A.). 

Adina racemosa Miquel, Cat. Mus. Bot. Lugd.-Bat. 1 : 44 (Fl. Jap.) 
(1870). 

Corniis Esquirolii Leveille in Fedde. Rep. Spec. Nov. 13: 257 (1914) ; 
Fl. Kouy-Tcheou, 116 ( 1914). — Synon. nov. 

China. K w e i c h o u : riviere, Tong-tcheou, J. Esquirol, no. 
407, June 1905 (holotype of Cornus Esquirolii; photo, in A. A.). 

Uncaria scandens (Sm.) Hutchinson in Sargent, Fl. Wilson. 3: 406 
(1916). — Leveille, Cat. Pi. Yun-Nan, 248 (1917). 

Cephalanthus Cavaleriei Leveille in Fedde, Rep. Spec. Nov. 10: 434 
(1912); Fl. Kouy-Tcheou, 365 (1915). — Synon. nov. 

China. K w e i c h o u : Ma-jo, J. Cavalerie, no. 3015, May and 
Nov. 1908 (holotype of Cephalanthus Cavaleriei; merotype in A. A.). 

Besides the flowering specimen described by Leveille there is a fruiting 
specimen under the same number in his herbarium which is probably the 
specimen collected in November. This specimen apparently represents 
U. rhynchophylla (Miq.) Miq. which is of wide distribution in Eastern 
Asia, but to my knowledge has not been previously collected in western 



illci Leveille, Fl. Kouy-Tcheou. 365 (1915). 

China. K w e i c h o u : ruisseau qui monte a Kiao-miay, alt. 
800 m., /. Esquirol, no. 3631, June 5, 1913, "arbre, 6 m." (holotype of 
Cephalanthus Navillei; merotype in A. A.). 

This species is very similar to A^. Griffitkii (Hook, f.) Merr. but easily 
distinguished by ternate heads on a rather slender peduncle about 3 cm. 
long, the slender pedicels being 3-4 cm. long with scars of bractlets near 
the middle. The leaves are identical with those of specimens of A". 
Griffithii from Yunnan (Henry, nos. 12676 and 12880) which are in 
flower while Esquirol's specimen is in fruit. 

Mussaenda Esquirolii Leveille, Fl. Kouy-Tcheou, 369 (1915). 
Mussaenda Wilsonii. Hutchinson in Sargent, PI. Wilson. 3:393 
(1916). — Synon. nov. 

China. K w e i c h o u : forets de Tong-tcheou, 1400 m., /. Es- 
quirol, no. 3264, June 22, 1912 "couleur jaune pale" (holotype of M. 



320 JOURNAL OF THE ARN< 

Esquirolii; photo, in A. A.). H u p e h : Chang-lo-hsien, ravines, 650 
m., E. H. Wilson, Arn. Arb. Exp. no. 3265, June 1907 "bracts white, 
flowers yellow" (holotype of M. Wilsonii in A. A.). 

The name Mussaenda Esquirolii does not appear on the original speci- 
men, but an unpublished name under another genus with the same 
specific epithet in Leveille's hand. The specimen diff'ers from the type 
of M. Wilsonii in the more conspicuous pubescence of the veins and 
veinlets of the under side of the leaf, but this may be due to the younger 
state of the leaves. 

Mussaenda pubescens Aiton f., Hort. Kew ed. 2, 1: 372 (1810). 
Mussaenda Bodinieri Leveille in Bull. Soc. Bot. France, 55:59 
(1908) ; Cat. PI. Yun-Nan, 246 (1917). — Hutchinson in Sargent. 
PI. Wilson. 3: 396 (1916). — Synon. nov. 

China. Kwangtung: pied du Tay-mo-chan (Taiman-san), 
commun dans les haies pres des villages, E. Bodinier, no. 1159, May 6, 
1895, "arbrisseau a branches sarmenteuses, fleurs blanches; differs des 
autres Mussaenda de I'absence de grande bractee florale" (holotype of 
M. Bodinieri; photo, in A. A.). 

In Mussaenda pubescens the enlarged calyx-lobe is sometimes much 
reduced in size or entirely absent. I find it entirely absent in Hongkong 
Herb. 2784, J. B. Norton 1475, and H. H. Chung 3391 from Fukien, 
and in R. C. Ching 5433 and 5435 from Kwangsi. The locality given by 
Leveille as Cay-mo-chan is apparently a misprint for Tay-mo-chan as 
it is clearly spelled on Bodinier's label; the same locality appears on 
English maps as Taimau-san. 



. (Hook. & Arn.) Merrill in Philipp. Jour. Sci. 
Bot. 13: 160 (1918). — Metcalf in Jour. Arnold Arb. 13: 29 (1932). 
Ehretia Esquirolii Leveille, Fl. Kouy-Tcheou, 54 (1914), non Leveille 
(1913). 

China. K w e i c h o u : route de Tong-tcheou, 1200 m., /. £5- 
quirol, no. 3775, June 1912 (holotype of Ehretia Esquirolii of 1914: 
photo, in A. A.). 

The name Ehretia Esquirolii (in Fedde, Rep. Spec. Nov. 12:335. 
1913) had been given by Leveille to another specimen collected by 
Esquirol at about the same time and at the same locality and numbered 
3214. This specimen cannot be found in the Leveille herbarium and 
was probably identified by Leveille with another plant and the name 
used again for Esquirol 3775. The number 3775 seems to have been a 
mixture, for Leveille enumerates in his Flore du Kouy-Tcheou the same 
number under Ehretia Dunniana, E. Esquirolii and E. macrophylla; 
under the last named as 3775 p.p. and without locality. 



1935] REHDER, LIGNEOUS PLANTS DESCRIBED BY LEVEILLE 321 

Tarenna incerta Koorders & Valeton in Meded. Lands Plantent. 59 : 
268 (Bijdr. Boomsort. Java, 8) (1902). — Merrill in Philipp. Jour. Sci. 
17:469 (1920). 

Tarenna zeyUinica Koorders & Valeton, 1. c. 82 (1902) ; non Gaertn. 
? Webera pallida Franchet ex Brandis, Ind. Trees, 378 (1906). 
Webera Cavaleriei Leveille in Fedde, Rep. Spec. Nov. 9: 323 (1911) ; 

Fl. Kouy-Tcheou, 372 (1915). 
Webera Henryi Leveille, Sert. Yunnan. 1 (May 1916) ; Cat. PI. Yun- 

Nan, 296 (1917). 
Tarenna pallida (Franch.) Hutchinson in Sargent, PI. Wilson. 3: 410 
(Aug. 1916). 
China. K w e i c h o u : Pin-fa, /. Cavalerie, no. 2342, June 8, 
1905, "petit arbre" (holotype of Webera Cavaleriei; photo, in A. A.). 
Yunnan : Szemao, A. Henry, no. 11923a (holotype of Webera 
Henryi; photo, in A. A.). 

The type of Webera Cavaleriei consists only of a year-old branch with 
a few leaves and a small fragment of an inflorescence with very young 
fruits. It differs somewhat from the type of W. Henryi in the mani- 
festly truncate calyx without any indication of teeth, though in the 
latter specimen perfectly truncate calyces occasionally occur. The type 
of W. Henryi agrees exactly with Henry no. 10686 which was identified 
by Hutchinson with Tarenna pallida together with Henry, nos. 11923, 
11923c and 11923f. 

Tarenna incerta seems to be somewhat variable in the number of 
ovules. Koorders & Valeton state that there are two or sometimes only 
one ovule in each locule and refer to a tree in the garden which had in 
all flowers only one ovule in each cell. In the one ovary of Webera 
Cavaleriei which I examined I also found only one ovule in each cell. 
Merrill (1. c.) states that the usual number of seeds in each fruit is 
apparently two. Brandis (1. c), however, describes the fruit of W. 
pallida as having 4-6 seeds. Two fruits examined of Henry 11923f had 
I and 3 seeds each. It, therefore, seems somewhat doubtful if Webera 
pallida Franch. of which I have not seen the type really belongs to 



Gardenia jasminoides Ellis in Philos. Trans. 51(2): 935, t. 25 
(1761). 

Gardenia florida Linnaeus, Spec. PI. ed. 2, p. 305, 1679 (1762).— 

Leveille, Fl. Kouy-Tcheou, 366 (1915); Cat. PI. Yunnan, 245 

(1917). 

Gardenia Schlechteri Leveille in Fedde, Rep. Spec. Nov. 10: 146 

(1911); Fl. Kouy-Tcheou, 366 (1915). — Synon. nov. 

China. K w e i c h o u : without precise locality, /. Esquirol, no. 



322 JOURNAL OF THE ARNOLD ARBORETUM I vol. xvi 

777, April 1905, "arbre, fl. blanche" (holotype of G. Schlechtcri; mero- 
type in A. A.). 

Esquirol no. 777 represents a rather small-flowered form, but other- 
wise it does not differ from G. jasminoides. 

Varneria augusta L. (in Amoen. Acad. 4: 136, 1759) upon which 
Merrill based the new combination G. augusta, is a nomen nudum. The 
oldest available specific epithet is jasminoides, though florida has been 
generally adopted. 

Ixora Henryi Leveille in Fedde, Rep. Spec. Nov. 13: 178 (1914); 
Fl. Kouy-Tcheou, 367 (1915); Cat. PI. Yun-Nan, 245 (1917). — Pitard 
in Lecomte, Fl. Gen. Indo-Chine, 3:324 (1924). — Chun in Sun- 
yatsenia, 1: 306 (1934). 

China. K w e i c h o u : Lo-fou, J. Cavalerie, no. 3496, March 
1909 (syntype; merotype in A. A.). Yunnan: Szemao, s. moun- 
tain forests, 5000 ft., A. Henry, no. 11637a, "shrub 5 ft., red flowers" 
(syntype; photo, and isotype in A. A.). 

Here also belong Henry nos. 10407 and 10407 a-c from Mengtze and 
11637 and 11637a-d from Szemao, Yunnan. 

Psychotria Henryi Leveille in Fedde, Rep. Spec. Nov. 13:179 
(1914). — Hutchinson in Sargent, PI. Wilson. 3: 415 (1916). 

China. Yunnan : Szemao, s. e. mountains, 4000 ft., A. Henry, 
no. 12146D, "shrub 4 ft., red fruit" (holotype; photo, and isotype in 



Psychotria rubra (Lour.) Poiret, Encycl. Meth. Suppl. 4:597 
(1816). 

Psychotria clliptica Ker in Bot. Reg. 8: t. 607 (1822) ; non II. & H. ex 

Roem. & Schuh. (1819). 
Psychotria Esquirolii Leveille in Fedde, Rep. Spec. Nov. 10:435 
(1912) ; Fl. Kouy-Tcheou, 371 (1915). — Synon. nov. 
China. K w e i c h o u : Ouang-mou, /. Esquirol, no. 119, June 
1904 "fl. blanchatre" (holotype of P. Esquirolii; photo, in A. A.). 

This species has apparently not yet been recorded from western 
China, but seems common in southeastern China west to Kwangsi. 

Psychotria Prainil Leveille in Fedde, Rep. Spec. Nov. 9:324 
(1911): Fl. Kouy-Tcheou, 371 (1915). 

China. K w e i c h o u : Ouang-mou, J. Esquirol, 76 (holotype; 
photo, in A. A.); Heou-hay-tse, /. Esquirol, no. 860, June 1906, "fl. 
blanche" (enumerated in Fl. Kouy-tcheou; photo, and merotype in 



19351 REHDER, LIGNEOUS PLANTS DESCRIBED BY LEVEILLE 323 

This species resembles in its capitate sessile or subsessile inflorescence 
P. morindoides Hutchins., but the inflorescence, branchlets and leaves 
beneath are covered with a ferrugineous pubescence similar to that of 
P. pilijera Hutchins., though shorter; besides it differs from P. pilijera 
in the sessile inflorescence, the smaller more coriaceous leaves glabrous 
above and in the short petioles. 

Lasianthus Hookeri Clarke ex Hooker, f., Fl. Brit. Ind. 3: 184 
(1880).— Leveille, Cat. PI. Yun-Nan, 246 (1917). 

Lasianthus Dunniana Leveille in Fedde, Rep. Spec. Nov. 11:64 
(1912); Fl. Kouy-Tcheou, 368 ( 1915). — Synon. nov. 

China. K w e i c h o u : Lo-fou, /. Cavalerie, no. 3459, Oct. 1908 
(holotype of L. Dunniana; photo, and merotype in A. A.). 

Leveille's description of the species is taken from a note on the type 
specimen which reads "aff. L. trichophlebus Hemsley, sed margine 
foliorum dense ciliata distincta,"' and is signed S. T. D(unn). It agrees 
exactly with Yunnan specimens referred by Hutchinson to L. Hookeri 
(in Sargent, PI. Wilson. 3: 402. 1916). 

Lasianthus Biermanni King ex Hooker f., Fl. Brit. Ind. 3: 190 
(1880). — Leveille, Cat. PI. Yun-Nan, 246 (1917). 

Lasianthus EsquirolU Leveille in Fedde. Rep. Spec. Nov. 11:295 
(1912); Fl. Kouy-Tcheou, 368 (1915).— Synon. nov. 
China. K w e i c h o u : without precise locality, /. Esquirol, no. 
648 (holotype of L. EsquirolU; photo, in A. A.). 

Esquirol's specimen agrees well with Henry no. 11148, identified by 
Hutchinson with L. Biermanni (in Sargent, PI. Wilson. 3: 402. 1916). 

Lasianthus Labordei (Levi.) Rehder in Jour. Arnold Arb. 13: 340 
(1932). 

Canthium Labordei Leveille in Fedde, Rep. Spec. Nov. 13:178 

(1914) ; Fl. Kouy-Tcheou, 384 (1915). 

China. K w e i c h o u : district de Tsin-gay, mont. de Kao-tchay, 

penchant escarpe des montagnes, /. Laborde in herb. Bodinier, no. 2109, 

March 7, 1898, "arbuste" (holotype of Canthium Labordei; photo, in 



Lasianthus Hartii Franchet in Bull. Soc. Bot. France, 46:209 
(1899). 

Canthium Dunnianmn Leveille in Fedde, Rep. Spec. Nov. 9:324 
(1911) ; Fl. Kouy-Tcheou, 364 (1915).— Synon. nov. 
China. K w e i c h o u : without precise locality, /. Esquirol 



(holotype of Canthium Dunnianum; photo. 



324 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

tagnes, /. Cavalerie, no. 3226, May 20, 1907, "1 m. de h., fl. blanches" 
(cited in Fl. Kouy-Tcheou; merotype in A. A.). 

In Flore du Kouy-Tcheou Leveille cites only Cavalerie, no. 3226, 
which is in bloom, while the type, which is not cited, is a fruiting 
specimen. 

This species is apparently related to L. japonicus Miq. from which it 
differs in the glabrous branchlets, glabrous calyx and glabrous or nearly 
glabrous leaves, and to L. longkauda Hook. f. from which it differs in the 
more coriaceous leaves with the veinlets less prominent beneath and in 
the five corolla-lobes being densely bearded inside up to the tip. The 
only flowering specimen of L. longicatida I have seen is Henry no. 10633 
which has a 4-lobed corolla ; this agrees with Hooker's original descrip- 
tion of the flowers as "usually 4-merous." The flowers are also 4-merous 
in Lasiantkus Labordei. 
Lasianthus spec. 

Canthium Cavaleriei Leveille in Fedde, Rep. Spec. Nov. 10:434 
(1912) ; Fl. Kouy-Tcheou, 364 (1915). 
China. K w e i c h o u : Ma-jo, /. Cavalerie, no. 3350 (holotype 
of Canthium Cavaleriei; photo, and merotype in A. A.). 

This species I am unable to identify with any species of Lasianthus 
and in the absence of flowers an exact determination is not possible. It 
can not be a Canthium, since the fruit is a several-seeded berry. 

Paederia scandens (Lour.) Merrill in Contr. Arnold Arb. 8: 163 
(1934). 

Paederia foetida Thunberg in Nov. Act. Soc. Sci. Upsal. 4:32 
(1783); Fl. Jap. 106 (1784). —Leveille, Fl. Kouy-Tcheou, 376 
(1915). — Non Linnaeus (1767). 
Paederia tomentosa Blume, Bijdr. 963 (1826). 
Paederia chinensis Hance in Jour. Bot. 16: 228 (1878). 
Paederia Esqiiirolii Leveille in Fedde, Rep. Spec. Nov. 10: 146 

(1911). 
Paederia Dunniana Leveille, 1. c. (1911). 

Paederia Mairei Leveille in Fedde, Rep. Spec. Nov. 13: 179 (1914). 
Paederia tomentosa Bl. var. Mairei (Levi.) Leveille, Cat. PI Yun- 
Nan, 247 (1917). — Synon. nov. 
China. K w e i c h o u : Ky-che-ten, /. Esquirol, no. 184, Aug. 
1904 (holotype of P. Esquirolii; photo, in A. A.); without precise lo- 
cality, /. Esquirol, no. 775, April 1905, "fleur a gorge rouge" (holotype 
of P. Dunniana; photo, in A. A.). Yunnan; broussailles des col- 
lines a Siao-ou-long, 2550 m., E. E. Maire, July 1911, ''arbuste grim- 
pant, odeur fetide, fl. roses" (holotype of P. Mairei; photo, in A. A.). 
The specimens cited above represent the typical glabrous form of 



1935] REHDER, LIGNEOUS PLANTS DESCRIBED BY LEVEILLE 325 

P. scandens; Paederia Esquirolii and P. Dunniana have narrow generally 
oblong leaves cuneate at base, while P. Mairei has larger and broader 
generally ovate leaves rounded to truncate at base. 

This species which is the most widely distributed of the genus has been 
generally called P. tomentosa Bl. which is described by Blume as having 
the leaves tomentose beneath. I have seen no specimens from the type 
region, but as long as I have no evidence to the contrary, I accept P. 
tomentosa Bl. as a synonym of P. scandens, representing the form with 
leaves pubescent beneath which occasionally occurs also in China. 

Paederia WaUichii Hooker f., Fl. Brit. Ind. 3: 196 (1881). 
Paederia tomentosa Bl. var. purpureo-caerulea Leveille & Vaniot in 

Bull. Soc. Bot. France, 55: 59 (1908). 
Paederia Bodinieri Leveille, Fl. Kouy-Tcheou, 371 (1915); non 

Leveille (1914). — Synon, nov. 
Cynanchum yunnanensc Leveille. Cat. PI. Yun-Nan, 13 (1915).— 

China. K w e i c h o u : environs de Hoang-ko-chou, rochers, etc., 
/. Seguin in herb. Bodinier, no. 2501, Sept. 8, 1898, "liane grimpante, 
fleurs, atro-pourpre, avec bordure bleu-rouge, inodore" (holotype of P. 
tomentosa var. purpweo-coerulea and P. Bodinieri; photo, and merotype 
in A. A.). Yunnan: brousse des montagnes a Tcha-ho, alt. 2800 
m., E. E. Maire, Nov. 1911, "arbuste grimpant, tomenteux, fl. violettes"; 
rives du fleuve Bleu a Siao-ho, 400 m., E. E. Maire, July 1912 "arbuste 
grimpant, feuil. blanches, velues en dessous" (syn types of Cynanchum 
yunnanense; photos, in A. A.). 

Paederia Bodinieri was described by Leveille without reference to his 
earlier P. tomentosa var. purpureo-caerulea, but it is based on the same 
specimen which bears only the name var. purpureo-caendea in Leveille's 
hand. The name is a later homonym of his earlier P. Bodinieri (in 
Fedde, Rep. Spec. Nov. 13: 179. 1914) which he referred the same 
year to Marlea as M. Cavaleriei and which turns out to be identical with 
Gardneria multiflora Mak. (see Jour. Arnold Arb. 15: 309). Cynan- 
chum yunnanense is not different from Seguin's specimen except that it 
has shorter inflorescences. 

The specimens enumerated above are identical with Henry's nos. 9126 
and 12442 and, judging from the description, seemed conspecific with 
P. WaUichii Hook. f. I am indebted to Sir Arthur W. Hill for a com- 
parison of the Henry numbers with the type specimens of P. WaUichii 
in the Kew Herbarium; he writes me that Mr. C. E. C. Fischer reports 
on these specimens as follows: "the only difference between these num- 
bers and the type of P. WaUichii Hook f. that I can see is that the basal 



326 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

lobes of the leaves are rather more rounded and the sinus slightly deeper 
and also the buds rather shorter. I doubt that this would suffice for even 
a variety." With the material at hand P. Wallkhii Hook. f. may be 
characterized by cordate to subcordate leaves densely scabrid above 
with setulose appressed hairs somewhat bulbous at base and densely 
villous-pubescent or tomentose beneath, and by the lateral branches of 
the inflorescence bearing one or several capitate flower clusters. In the 
capitate or subcapitate flowers it resembles P. mkrocephala Pierre from 
which it differs in the short calyx-teeth and in the dense pubescence of 
the leaves. 

Paederia Cavaleriei Leveille in Fedde, Rep. Spec. Nov. 13:179 

Non Blume (1826). 

China. K w e i c h o u : bois des montagnes, J. Cavalcrie, no. 
2058, Aug. 1904, "plante tres puante" (holotype of P. 'Cavaleriei; photo, 
in A. A.). 

This species is similar to the preceding, but differs in the long hirsute 
ferrugineous pubescence of the branches, the petiole and the inflo- 
rescence, in the petiole being 7.5 8 cm. long and in the 1-1.5 mm. long 
fairly straight hairs thinly covering the underside of the leaves, but 
dense on the midrib and veins, also in the glabrous narrower calyx-teeth. 
According to the original description it resembles P. pilijera Hook. f. 
in the long pubescence, but Pitard (in Lecomte, Fl. Gen. Indo-Chine, 
3: 412) describes the leaves as very tomentose beneath, the calyx-tube 
as very tomentose and the petiole as 1-2.5 cm. long. 

A specimen collected in Kweichou at Sanhoa by W. Y. Chun (no. 
6301) agrees in the pubescence of the stem and the leaf and in the thin 
texture of the leaf very well with P. Cavaleriei, but the lateral branches 
of the inflorescence end in cincinnate cymes, as is the rule in P. scandens, 
and not in subcapitate or capitate cymes, characteristic of this and the 
preceding species. 

Leptodennis Potanini Batalin in Act. Hort. Petrop. 14:319 
(1898). — H. Winkler in Fedde, Rep. Spec. Nov. 18: 152 (1922). 

Leptodennis Esquirolii Leveille in Fedde, Rep. Spec. Nov. 9:324 

(1911); 13:179 (1914); Fl. Kouy-Tcheou 368 (1915); Cat. Fl. 

Yun-Nan, 246 (1917). 

China. Kweichou: Hin-y-fou, /. Cavalerie, no. 3930, July 

1912 (cited in Fl. Kouy-Tcheou; duplicate in A. A.). Yunnan : 

Ouan-tse, /. Esquirol, no. 1503, May 22, 1909 "blanche a I'interieur, 

rouge a I'exterieur" (holotype of L. Esquirolii; photo, in A. A.). 



1935] REHDER, LIGNEOUS PLANTS DESCRIBED BY LEVEILLE 327 

Leveille published L. Esquirolii a second time in 1914, but with a 
briefer description, both based on Esquirol no. 1503 which is a flower- 
ing specimen. Cavalerie no. 3930 from Kweichou is a fruiting specimen 
and owing to its fully developed broader leaves looks somewhat different, 
but apparently belongs to this species. Leptodermis Esquirolii was first 
identified with L. Potanini by H. Winkler (1. c). 

Leptodermis Potanini var. gflauca (Diels) H. Winkler in Fedde, 
Rep. Spec. Nov. 18: 153 (1922). 

Leptodermis motsouensis Leveille in Bull. Geog. Bot. 25: 47 (1915) ; 
Cat. PI. Yun-Nan, 246 (1917). — Synon. nov. 
China. Yunnan : collines arides de Mo-tsou, 800 m., E. E. 
Maire, May 1912, "sous-arbrisseau en touffes, fl. blanches" (holotype of 
L. motsouensis ; merotype in A. A.). 

The identification of L. motsouensis with L. Potanini var. glauca was 
communicated to me by Dr. H. Winkler in a recent letter as were the 
identifications of the following species of Leptodermis. 

Leptodermis Potanini var. tomentosa H. Winkler in Fedde, Rep. 
Spec. Nov. 18: 15,3 (1922). 

Leptodermis tongchouancnsis Leveille in Bull. Geog. Bot. 25:47 

(1915); Cat. PI. Yun-Nan, 246 (1917), "tongtchouanensis" — 

Synon. nov. 

China. Yunnan : rochers des coteaux autour de Tong-tchouan, 

2550 m., E. E. Maire, May 1912, "arbrisseau, feuilles velues blan- 

chatres, fl. blanches soyeuses" (holotype of L. tongchotianensis ; mero- 

Leptodermis pilosa (Franch) Diels var. glabrescensH. Winkler in 
Fedde, Rep. Spec. Nov. 18: 160 (1922). 

Leptodermis Mairci Leveille in Fedde, Rep. Spec. Nov. 13: 179 
(1914) ; Cat. PI. Yun-Nan, 246 (1917). — Synon. nov. 

China. Yunnan : plaine de Long-tang, 2500 m., E. E. Maire, 
Aug. 1912, "arbuste non grimpant, en touffes, fl. violet" (holotype of 
L. Mairei; merotype in A. A.) ; haies, plaine de Tong-tchouan, 2500 m., 
E. E. Maire, in 1912 "arbuste buissonant, ecorce blanche, fl. roses" (in 
herb. Leveille in cover of L. Mairei; duplicate in A. A.). 

Only the specimen from Long-tang bears the name L. Mairei in 
Leveille's hand. As Maire no. 21 Winkler (1. c.) enumerates under his 
L. pilosa var. glabrescens a specimen apparently of the same collection 
as Leveille's type of L. Mairei. 

Another specimen "a mi-mont de Siao-ho, 2800 m." in herb. Leveille 
under L. Mairei belongs to Wendlandia longidens (Hance) Hutch, (see 
p318.). 



328 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

Leptodermis oblongaBunge in Mem. Sav. Etr. Acad. Sci. St. Petersb. 
2: 108 (Enum. PI. Chin. Bor. 34) (1833). 

Leptodermis Chancti Leveille in Bull. Geog. Hot. 25:47 (1915). 
China. H o p e i : montagnes de Ping-chan, L. Chanet, no. 538 bis, 
Aug. 1910; without precise locality, L. Chanet, no. 574, June 1904 (syn- 
types of L. Chaneti; merotypes in A. A.). 
Prismatomeris Henr}^ (Levi.), comb. nov. 
Canthium Henryi Leveille in Fedde, Rep. Spec. Nov. 13: 178 (1914) ; 

Cat. PI. Yun-Nan, 245 (1917).— Synon. nov. 
Prismatomeris brevipes Hutchinson in Sargent, PI. Wilson. 3:413 
( 1916) . — Leveille, Cat. PI. Yun-Nan, 247 ( 1917) . — Synon. nov. 
China. Yunnan: Meng-tse, S. E. mountains, 5000 ft., A. 
Henry, no. 9040f, "shrub 6 ft., fruit red" (holotype of Canthium Henryi; 
photo, in A. A. ; paratype of P. brevipes; isotype in A. A.). 

There is also an isotype of the holotype of P. brevipes, Henry 9040e, in 
the herbarium of the Arnold Arboretum and of another paratype, Henry 
9040D. 

CAPRIFOLIACEAE 
Sambucus javanica Bl. var. Argyi (Levi.), var. nov. 
Sambucus Argyi Leveille in Bull. Geog. Bot. 24 (no. 3301): 292 
(1914); in Mem. Acad. Ci. Arts Barcelona, ser. 3, 12: 545 (Cat. 
PI. Kiang-Sou, 5) (1916). 
China. K i a n g s u : Ka-se-dao, trouve venant de Tou-ka-dou, 
Ch. d'Argy [1848-66] (holotype of S. Argyi; photo, in A. A.) ; Se-hom, 
(jardin), Ch. d'Argy [1848-66] "fruit rouge" (with S. Argyi in herb. 
Leveille; photo, in A. A.). 

This variety differs from the type in the shorter and comparatively 
broader more coarsely serrate leaflets, the lateral ones ovate-oblong to 
oblong (5 X 1.8), the terminal elliptic or elliptic-obovate (6 X 3), and 
slightly scaberulous on the veins. 

Sambucus Argyi was first referred to S. javanica by H. K. Airy-Shaw 
according to a note on the specimens. 

Viburnum erubescens Wallich, PI. As. Rar. 2: 29, t. 143 (1830).— 
Leveille, Cat. PI. Yun-Nan, 28 (1916). 

Viburnum botryoidcum Leveille, Cat. PI. Yun-Nan, 28 (1915).— 

China. Y u n 
E. E. Maire, May : 
deum; merotype in A. A.). 

oliganthum Batalin in Act. Hort. Petrop. 13:372 



1935] REHDER, LIGNEOUS PLANTS DESCRIBED BY LEVEILLE 329 

Viburnum Stapfianum Leveille in Fedde, Rep. Spec. Nov. 9:443 
(1911); Fl. Kouy-Tcheou, 66 (1914). — Synon. nov. 
China. K w e i c h o u : Ma-jo, /. Cavalerie, no. 3002, May 1908, 
"petit arbre, fleurs roses" (holotype of V. Stapfianum; merotype in 

A. A.). 

I above is 



Viburnum sympodiale Graebner in Bot. Jahrb. 29: 587 (1901).— 
Rehder in Sargent, Trees & Shrubs, 2: 83, 108, t. 139 (1908). 

Viburnum Martini Leveille in Fedde, Rep. Spec. Nov. 9: 443 (1911) ; 
Fl. Kouy-Tcheou, 66 (1914). — Synon. nov. 

China. K w e i c h o u : Pin-fa, bois, rare, /. Cavalerie, no. 2272, 
April 4, 1905, "fl. blanches" (holotype of V. Martini; merotype in 
A. A.). 

Viburnum Cavaleriei Leveille in Fedde, Rep. Spec. Nov. 9: 442 
(1911); Fl. Kouy-Tcheou, 66 (1914). 

China. K w e i c h o u : Pin-fa, montagnes, /. Cavalerie, no. 977, 
April 13, 1903, "h. 1 a 2 m., fl. blanches odorantes," (holotype; photo. 
and merotype in A. A.). 

This species is closely related to V. jallax Graebn. and V. chinchanense 
Graebn. differing from the former in its coriaceous leaves rugulose and 
stellate pubescent above and from the latter in the glabrous ovaries and 
the leaves being sparingly stellate-pubescent and somewhat scabrid 
above. Both related species have been collected in Kweichou ; V. jallax 
is represented in this herbarium from Kweichou by Steward, Chiao & 
Cheo 11, and V. chinchanense by Y. Tsiang 7419 and 9216 and also by 
another specimen, Tsiang 8424, which approaches V. Rosthornii Graebn. 
by its larger subcordate leaves sparingly stellate above. 

Viburnum congestum Rehder in Sargent, Trees & Shrubs, 2:111 
(1907). — Leveille, Cat. PI. Yun-Nan, 28 (1915). — P'ei in Mem. Sci. 
Soc. China l(no. 3) : 90 (Verben. China) (1932). 

Hedyotis Mairei Leveille in Fedde, Rep. Spec. Nov. 13: 176 (1914) ; 

Cat. PI. Yun-Nan, 245 (1917). - Synon. nov. 
Viburnum Mairei Leveille, Cat. PI. Yun-Nan, 28 (1915). — Synon. 

Premna EsquiroUi Leveille, Sert. Yunnan. 3 (1916) ; Cat. PI. Yun- 
Nan, 298 (1917). 
Oldenlandia Mairei (Levi.) Chun in Sunyatsenia, 1: 314 (1934). 
China. Yunnan : brousse du plateau de Ta-hai-tse, alt. 3200 
m., E. E. Maire, May (1912 or 1913), "grande arbuste, feuilles cadu- 
ques, fl. blanches" (holotype of Hedyotis Mairei, named H. yunnanensis 



330 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

on the label of the type specimen; photo, in A. A.) ; brousse au pied de 
lo-chan, 3200 m., E. E. Maire, May 1912, "arbuste haut 1.30 m.," (holo- 
type of V. Mairei; merotype in A. A.); brousse des montagnes a Mo- 
tsou, 800 m., E. E. Maire, May 1912, "arbuste, feuilles caduques, f^. 
blanches" (holotype of Premna Esquirolii; merotype in A. A.). 

The three specimens cited above have the corolla tube somewhat 
shorter than in the type of V. congestum, the tube being only slightly 
longer than the lobes, but in the otherwise similar V. utile Hemsl. the 
tube is wide-campanulate and much shorter than the lobes. Viburnum 
congestum differs from V. utile also in the less dense grayish tomentum of 
the under side of the leaves, the individual hairs being distinguishable 
and usually have shorter rays, while in V. utile they are matted and the 
tomentum is whitish, brownish on the veins in young leaves. There are, 
however, specimens of V . congestum with a denser tomentum similar to 
that of V. utile. Geographically the two species seem to be well sepa- 
rated ; V. congestum is common in Yunnan and extends into Kweichou 
and western Szechuan, while V. utile occurs in Hupeh and extends to 
eastern Szechuan. 

Viburnum cylindricum Ham. var. crassifolium (Rehd.) Schneider 
in Bot. Gaz. 64: 77 (1917). 

Viburnum crassifolium, Relulcr in Sar^-ent, Trees & Shrul)s, 2: 112 

(1908). — Leveille, Cat. Pi. Yun-Nan, 28 (1915). 
Viburnum pinfaense Leveille in Fedde, Rep. Spec. Nov. 9:442 
(1911) ; pro parte, quoad Cavalerie no. 1483; Fl. Kouy-Tcheou, 66 
(1914). — Syiion.nov. 

China. Kweichou : Pin-fa, bois, /. Cavalerie, no. 1483, Oct. 
12, 1903 (syntype of V. pinfaense; merotype in A. A.). 

This variety has been collected in Kweichou in three different locali- 
ties by Y. Tsiang (nos. 4121, 7558 and 9137). The leaves of Cavalerie's 
specimen are unusually small and narrow. 

Under V. pinfaense Leveille describes two different plants, of which 
the fruiting specimen belongs here, while the flowering one belongs to 
V. sempervircns K. Koch. 

It, Trees & Shrubs, 2: 37, 112 



Viburnum Chaffanfoni Leveille in Fedde, Rep. Spec. Nov. 9:443 
(1911) ; Fl. Kouy-Tcheou, 66 (1914).— Synon. nov. 
China. Kweichou : Pin-fa, bois presque a pic, /. Cavalerie, 
o. 3093, July 2, 1907 (holotype of V. Chaffanjoni; merotype in A. A.). 
This very distinct species, differing in its ternate leaves from all other 
pecies, was known to me before only from Szechuan: banks of Min 



1935] REHDER, LIGNEOUS PLANTS DESCRIBED BY LEVEILLE 331 

River (Wilson 3736), Mt. Omei (W. P. Fang 2461, 2631, 3309, 3355 
and F. T. Wang 23138), and Kuan-hsien (W. P. Fang 2021). 

Viburnum Schneiderianum Handel-Mazzetti in Akad. Wiss. Wien 
Anzeig. 1925: 66 (PI. Nov. Sin. Forts. 33: 4) (1925). 

China. Yunnan: rochers de lo-chan, alt. 3200 m., E. E. Maire, 
May (1911-13), "arbrisseau rampant, toujours vert," (in herb. Leveille 
sub Gaidtheria crenulata; duplicate in A. A.). 

The specimen cited above was referred by Leveille to Gaidtheria 
crenulata Kurz and represents, at least partly, the plant enumerated 
under that name in his Cat. PI. Yun-Nan, 86 (1916). 

Viburnum sempervirens K. Koch, Hort. Dendr. 300 (1853).— 
Rehder in Sargent, Trees & Shrubs, 2: 95, 113, t. 145 (1908). 

Viburnum pinfaense Leveille in Fedde, Rep. Spec. Nov. 9:442 
(1911), pro parte, quoad specim. no. 1056; Fl. Kouy-Tcheou, 66 
(1914). — Synon.nov. 
China. K w e i c h o u : Pin-fa, montagnes, /. Cavalerie, no. 1056, 
June 11, 1903 (syntype of V. pinfaense; merotype in A. A.). 

This species seems to be rare in vi^estern China. I have seen it only 
from Pin-fa, Kweichou (Cavalerie 1056 and Y. Tsiang 6385), and from 
Szemao, Yunnan (Henry 12753). 

Viburnum foetidum Wallich, PI. As. Rar. 1 : 49, t. 61 (1830).— 
Leveille, Cat. PI. Yun-Nan, 28 (1915). — P'ei in Mem. Sci. Soc. China, 
1, no. 3: 90 (Verben. China) (1932). 

Viburnum ajugifolium Leveille in Fedde, Rep. Spec. Nov. 9:441 

(1911) ; Fl. Kouy-Tcheou, 65 (1914). — Synon. nov. 
Premna Vdbrayi Leveille, Sert. Yunnan. 4 (1916); Cat. PI. Yun- 
Nan, 299 (1917). 
China. Kweichou : environs de Kouy-yang, mont du College, 
c. dans les haies, bords des ruisseaux, E. Bodinier, no. 2231, May 18, 
1898, "fl. blanches" (holotype of V. ajugifolium; merotype in A. A.). 
Yunnan: haies et brousses des montagnes, a Tong-tchouan, 2500- 
2700 m., E. E. Maire, July 1912, "arbuste grele, feuill. caduques" (holo- 
type of Premna Valbrayi; merotype in A. A.). 

The two specimens cited above are similar to the form described as 
V. ceanothoides C. H. Wright. 

Viburnum foetidum var. rectangulatum (Graebn.) Rehder in Sar- 
gent, Trees & Shrubs, 2: 114 (1908), "rectangulum." 

Viburnum Touchancnsc Leveille in Fedde, Rep. Spec. Nov. 9:442 
(1911) ; Fl. Kouy-Tcheou, 66 (1914).— Synon.nov. 



332 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

Hedyotis yunnanensis Leveille in Fedde, Rep. Spec. Nov. 13: 176 

(1914); Cat. PI. Yun-Nan, 245 (1917). ~ Synon. nov. 
Old enlandia yunnanensis (Levi.) Chun in Sunyatsenia, 1: 310 (1934). 
China. K w e i c h o u : environs de Tou-chan, bord de la route, 
/. Cavalerie, no. 2192, July 5, 1897, "arbrisseau aux fleurs odorantes" 
(holotype of V. Touchanense ; merotype in A. A.). Yunnan : brous- 
sailles des collines a Long-ky, alt. 700 m., E. E. Maire, June 1911, 
"arbuste a feuilles caduques, fl. blanches" (holotype of Hedyotis yunnan- 
ensis; photo, in A. A.). 

Viburnum setigerum Hance in Jour. Bot. 20 : 261 ( 1882 ) . — Reh- 
der in Jour. Arnold Arb. 12 : 77 ( 193 1 ) . 

Viburnum theiferum Rehder in Sargent, Trees & Shrubs, 2:45, 113, 

t. 121 (1907). 
Viburnum Bodinieri Leveille in Fedde, Rep. Spec. Nov. 9:442 
(1911); Fl. Kouy-Tcheou, 65 (1914). 
China. K w e i c h o u : environs de Kouy-yang, bois de Kin-lin- 
chan, E. Bodinier, no. 2193, April 14, 1898, "arbuste, fl. blanches" (syn- 
type of V. Bodinieri; merotype in A. A.) ; Pin-fa, bois ombreux, /. Cava- 
lerie, no. 1285, May, 1903, "fl. blanches, odorantes" (syntype of V. 
Bodinieri; photo, in A. A.). 

Viburnum Bodinieri was identified with V. setigerum by the writer 
and the identification published in 1931 (1. c). The species has been 
collected in Kweichou also by Y. Tsiang near Tsunyi (no. 5318) and on 
the Yun-fu-shan near Pin-fa (no. 5510) and near Tuyun (5942). 

Viburnum corylifolium Hooker f. & Thomson in Jour. Linn. Soc. 
2: 174 (1858). 

Viburnum Dunnianum Leveille in Fedde, Rep. Spec. Nov. 9:442 

(1911); Fl. Kouy-Tcheou, 66 (1914). — Synon. nov. 
Viburnum barbigerum Leveille, Fl. Kouy-Tcheou, 65 (1914).— 
Synon. nov. 
China. Kweichou: route de Pin-yue a Kouy-yang, bords 
d'une riviere, L. Martin in herb. Bodinier, no. 2598, May 13, 1899, 
"grand arbuste, fl. blanches"; environs de Kouy-yang, mont du College, 
/. Chaffanjon, May 1, 1898, "arbuste, fl. blanches"; route de Pin-fa a 
Oug-lan, J. Cavalerie, Aug. 1908 "fruite rouge" (syntypes of V. Dunn- 
ianum; photos, of Martin's and Cavalerie's specimens, merotype of 
Chaffanjon's specimen in A. A.); Pin-fa, bois de hautes montagnes, /. 
Cavalerie, no. 1742, Aug. 1904 (holotype of V. barbigerum; merotype in 

5 characters with the other speci- 
^ different in being densely cov- 
however, that this development 



1935] REHDER, LIGNEOUS PLANTS DESCRIBED BY LEVEILLE 333 

of hairs is abnormal, since I have found a few other specimens of 
Viburnum namely R. C. Ching nos. 2826 and 2952 of V. ichangense 
(Hemsl.) Rehd. from Anhwei with some of the fruits densely covered 
with similar, though somewhat shorter, hairs, while the rest of the fruits 
was perfectly normal and glabrous. 

Viburnum corylijolium is perhaps only a variety of V. dilatatum Thbg. 
differing chiefly in the long spreading hairs of the young branchlets, in- 
florescence and petioles, while in V. dilatatum these parts are covered by 
a short and close stellate tomentum. Viburnum corylijolium has been 
collected in Kweichou also by Y. Tsiang (no. 5779) near Tu-yun and by 
Steward, Chiao and Cheo, (no. 583) on Niu-tu-shan; V. dilatatum was 
collected by Y. Tsiang (no. 6270) on Yao-ren-shan, Sanhoa. 



I erosum Thbg. var. Taquetii (Levi.) Rehder in Sargent, 
PI. Wilson. 1:311 (1912). — Nakai in Nakai & Koidzumi, Trees & 
Shrubs Jap. ed. 2, 1: 609 (1927). — Makino & Nemoto, Fl. Jap. ed. 2, 
p. 1146 (1931). 

Viburnum Taquetii Leveille in Fedde, Rep. Spec. Nov. 9: 443 (1911). 

Viburnum erosum var. punctatum Franchet & Savatier ex Nakai, Fl. 

Sylv. Kor. 11 : 42, t. 12 c. d (1921) quoad synon. V: Taquetii Levi., 

vix Franch. & Sav. 

China. Korea: Quelpaert, in silvis Yengsil, 1000 m., E. Taquet, 

no. 4281, Aug. 12, 1910 (holotype of V. Taquetii; photo, and isotype in 

A. A.). 

This peculiar variety chiefly characterized by the narrow leaves partly 
with two basal lobes near the base has been collected in Quelpaert also 
by E. H. Wilson (no. 9406). Nakai in 1921 (1. c.) referred it to V. 
erosum var. punctatum Franch. & Sav., but that variety represents 
apparently the plant with broader leaves densely stellate-pubescent 
above which seems to be the most widely distributed form. The gla- 
brous or glabrescent form, var. laeve Franch. & Sav., which seems much 
rarer must be considered the typical form, since Thunberg (Fl. Jap. 
124) describes the leaves as glabrous. 

Dipelta jninnanensis Franchet in Rev. Hort. 1891: 246, fig. 62.— 
Leveille, Cat. PI. Yunnan, 27 (1915). 

Cavaieriella Dunniana Leveille, Fl. Kouy-Tcheou, 61 (1914).— 

China. Kweichou : hautes montagnes, Long-ly, J. Cavalerie, 
no. 3023, May 1908 (holotype of Cavaieriella Dunniana; merotype in 

A. A.). 



334 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

Abelia verticillata Leveille, Fl. Kouy-Tcheou, 61 (1914). 

China. K w e i c h o u : Pin-fa, ruisseau du sud, /. Cavalcrie, no. 
497, Sept. 1912, "fl. blanche-violette-pourpree" (holotype; photo, in 
A. A.). 

This species appears to be closely related to A. uniflora R. Br. and A. 
Graebneriana Rehd., but differs from both in the pilose branches and in 
the ovary being pilose with rather long white hairs; it also differs from 
the former in the ciliate leaves and from the latter in the subcoriaceous 
leaves not bearded in the axils and without hairs alongside the midrib 
and the base of the lateral veins. On one of the branches of the type 
specimen the leaves are in whorls of threes, from which the specific 
epithet is derived, but the other branch has opposite leaves; branches 
with ternate leaves are also occasionally found in A. uniflora and in other 
species, e. g. in Wilson no. 747 of A. parvijolia Hemsl. 

Abelia Schumaimii (Graebn.) Rehder in Sargent, PI. Wilson. 1:121 
(1911). — Leveille, Cat. Pi. Yun-Nan, 26 (1915). 

Strobilanthopsis dcutziaefolius Leveille in Fedde, Rep. Spec. Nov. 

12:20 (1913). 
Abelia deutziae folia (Levi.) Leveille, Fl. Kouy-Tcheou, 60 (1914).— 

Synon. nov. 
Strobilanthes deutziacfolia, Leveille, 1. c. (1914), pro synon. 

Abcliae deutziae foliac. — Ind. Kew. Suppl. 4: 252 (1921). 
Abelia Mairei Leveille. Cat. PI. Yun-Nan, 26 ( 1915). — Synon. nov. 
China. K w e i c h o u : sous bois, /. Esquirol, no. 466, June 1905, 
"sous-arbrisseau, fl. blanches, panachees de rose," (holotype of Stro- 
bilanthopsis deutziaejolius ; merotype in A. A.). Yunnan: rochers 
des collines a Siao-ou-long, 2250 m., E. E. Maire, June 1912, "arbuste 
delicat, buissonant, haut 0.80 m., fl. roses (holotype of A. Mairei; mero- 
type in A. A.). 

The leaves of Esquirol's no. 466 are rather large being up to 3.5 cm. 
long and resemble somewhat those of A. Graebneriana Rehd., though 
scarcely acuminate, but the branchlets are pubescent. 

Abelia myrtilloides Rehder in Sargent, PI. Wilson. 1 : 120 (1911). 
Strobilanthopsis hvpericifolius Leveille in Fedde, Rep. Spec. Ni 

12:20 (1913), "'hypercifolia." 
Abelia Boditiieri Leveille, Fl. Kouy-Tcheou, 61 (1914) pro syn( 

A. pan'ifoliae. 
Abelia pannfolia Hemsl. sec. Leveille, Fl. Kouy-Tcheou, 61 (191^ 

saltern quoad specim. Bodinier, no. 1607, vix Hemsley. 
Strobilanthes hypericifolia Leveille, 1. c. (1914), pro svnon. 

parz'ifoliae. — \m\. Kew. Suppl. 5:252 (1921). 
China. K w e i c h o u : mont de Lou-tsong-koan, 1500 m., Kif 



1935] REHDER, LIGNEOUS PLANTS DESCRIBED BY LEVEILLE 335 

lin-shan qa at la dans les montagnes rocailleuses, E. Bodinier, no. 1607, 
June 1, 1897 and June 19, 1899, "petit arbuste a jolies fleurs roses" 
(syntypes of Strobilanthopsis hypericifolius ; photo, and merotype in 
A. A.). 

Leveille cites in his Flore du Kouy-Tcheou (1. c.) under A. parvijolia 
as synonyms besides Strobilanthes hypericifolia the unpublished name 
A. Bodinieri; both names appear on the labels of the type sheet of 
Bodinier no. 1607. The two specimens on the type sheet are somewhat 
intermediate between A. myrtilloides and A. parvijolia Hemsl., but in 
the oblong-elliptic or oblong-ovate shape of the leaves glabrous above 
and nearly so beneath they seem closer to the former, only in the 
glandular under surface they approach A. parvijolia which typically has 
ovate leaves of thicker texture pilose and glandular on both surfaces. A 
form very similar to Bodinier's specimen was collected near Kwei-yang, 
Kweichou, by Handel-Mazzetti, (no. 10477) who determined it as "A. 
parvijolia Hemsl. trans, ad A. myrtilloidem Rehd." 

Abelia Cavaleriei Leveille, Fl. Kouy-Tcheou, 60 (1914). 

China. Kweichou: sud de Tin-fan, mont. rocheuses, J. Cava- 
lerie, no. 1909, Oct. 1904, "fl. blanches" (holotype; merotype in A. A.). 

This is a very distinct species on account of its subcoriaceous leaves 
which recall those of Ligustrum strongylophyllum Hemsl. The species 
is apparently nearest A. chinensis R. Br., but is readily distinguished by 
the subcoriaceous perfectly glabrous quite entire leaves broadly ovate to 
orbicular-ovate, 1-2 cm. long, rounded or broadly cuneate at base, 
obtuse or rounded and apiculate at the apex. The branchlets and the 
many flowered terminal inflorescence are minutely puberulous. The 
specimen is in fruit but according to the collector the flowers are white. 

Lonicera tangutica Maximowicz in Bull. Acad. Sci. St. Petersb. 24: 
48 (1877); in Mel. Biol. 10: 75 (1877). 

Lonicera Rocheri Leveille in Bull. Geog. Bot. 24 (no. 301): 289 
(1914); Cat. PI. Yun-Nan, 27 (1915). — Synon. nov. 

China. Yunnan: brousse de Lan-mou-kiao, 3000 m., E. E. 
Maire, May 1912, "arbuste en touffes; fl. jaunes" (holotype of L. 
Rocheri; merotype in A. A.). 

The species cited above differs somewhat from typical L. tangutica in 
the linear-lanceolate somewhat leafy bracts about twice as long as 
ovary, in the anthers being exserted about one-half and in the less 
slender corolla-tube, but in its other characters it agrees with this species. 

Lonicera lig^strina Wallich in Roxburgh, Fl. Ind. ed. 2, 2: 179 
(1824). — Leveille, Cat. PI. Yun-Nan, 27 (1915). 



336 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

Lomccra missionis Leveille, Fl. Kouy-Tcheou. 63 (1914), pro parte, 
quoad specim. "Esquirol (May 10, 1906). Chaffanjon no. 2215"; 
Cat. PI. Yun-Nan, 27 (1915). 
China. K w e i c h o u : environs de Kouy-yang, mont. du Col- 
L la cascade, /. Chaffanjon in herb. Bodinier, no. 2215, April 14, 
"arbuste" (syntype of L. missionis; photo, in A. A.); mont du 
College, grotte, /. Esquirol, May 10, 1906 "fl. blanches" (syntype of 
L. missionis; merotype in A. A.). Yunnan: sous bois de couteaux 
Long-ky, 700 m., E. E. Maire, "arbuste toujours vert, fl. blanches," 
in herb. Leveille under L. missionis; duplicate in A. A.). 
This species has been collected in Kweichou also by W. Tsiang (nos. 
4580, 5972, 7640) and by Steward, Chiao & Cheo (no. 244). 

Lonicera pileata Oliver in Hooker, Icon. PI. 16: t. 1585 (1887). — 
Leveille, Cat. PI. Yun-Nan, 27 (1915). 

Lonicera missionis Leveille, Fl. Kouy-Tcheou, 63 (1914) pro parte, 
quoad specimen "Laborde 2502"; Cat. PI. Yun-Nan, 27 (1915).— 
Synon. nov. 
Lonicera huxifolia Leveille, Fl. Kouy-Tcheou, 63 (1914); Cat. 111. 
Seu-Tchouen, t. 11 (1918) MS. — Ssmon. nov. 
China. Kweichou : environs de Tsin-gay a Kia-la-tchong, J. 
Laborde in herb. Bodinier, no. 2502, Nov. 1898 "les fruits sont des jolies 
perles bleu-tendre, pulpeuses" (syntype of L. missionis; photo, in 
A. A.) ; environs de Kouy-yang, mont du College, rochers de la cascade, 
au bords de I'eau, /. Chaffonjon in herb. Bodinier, no. 2169, April 12, 
1898, "fl. jaunatres" (syntype of L. buxijolia; photo, in A. A.); grotte 
du College, 1350 m., /. Esquirol, no. 2069, April 1910, "blanche" (syn- 
type of L. buxijolia; photo, in A. A.); Pin-fa, ruisseaux, /. Cavalerie, 
no. 1319, April 9, 1902, "fl. blanche" (in herb. Leveille under L. buxi- 
jolia; photo, in A. A.). 

This species has been collected in Kweichou also by Y. Tsiang (nos. 
4529, 4562, 7937) and by Steward, Chiao & Cheo (no. 803); the last 
named specimen is approaching in the shape of its leaves f . linearis Rehd. 
The specimens named L. buxijolia by Leveille differ from typical L. 
pileata in their rather small leaves. The two species, L. ligustrina and 
L. pileata, are closely related and connected by intermediate forms in 
regard to shape of the corolla and of the leaves and to pubescence. 
Without flowers L. ligustrina may be distinguished by the leaves being 
generally ovate, rounded at base, acuminate, and with the midrib more 
or less impressed above at least toward the base and strigose, while L. 
pileata has generally elliptic to oblong leaves, narrowed at base, obtuse 
to acute at apex, with the midrib distinctly elevated above and glabrous. 
In regard to shape and pubescence of the leaves L. nitida Wils. seems 



1935] REHDER, LIGNEOUS PLANTS DESCRIBED BY LEVEILLE 337 

intermediate between the two, but the leaves are much smaller and 
usually broader, generally ovate, but not acuminate. Lonicera virgul- 
torum W. W. Sm. is very close to L. ligustrina and chiefly distinguished 
by the shape of the corolla. 

Lonicera fragilis Leveille in Fedde, Rep. Spec. Nov. 13:33 7 
(1914); Cat. PI. Yun-Nan, 37 (1915). 

China. Yunnan: vallee de Li-tse-pin, 2800 m., E. E. Maire, 
April 1913, "arbuste cassant, haut de 1.20 m., fl. roses" (holotype; mero- 
type in A. A.). 

Frutex metralis ramis hornotinis sparse setosis vel glabris; gemma 
terminalis interdum evoluta perulis duabus exterioribus et 4-6 interiori- 
bus. Folia nondum plane evoluta, elliptico-oblonga, acuminata, basi 
cuneata, utrinque hirsuta, glandulis sparsis intermixtis, margine ciliata 
et stipitato-glandulosa. Flores praecoces in axillis bractearum ad basin 
ramulorum; pedunculi brevissimi glabri; bracteae late ovatae, 8-10 mm. 
longae, irregulariter eroso-denticulatae, basin versus ciliatae et sparsis- 
sime stipitato-glandulosae, apicem versus glabrae, ceterum extus intusque 
glaberrimae; ovaria subglobosa, glabra; calyx ovario circiter duplo 
longior, latus et plicatus, dentibus carnosulis inaequalibus 1.5-3 mm. 
longis, late ovatis apice rotundatis margine irregulariter erosulis glabris; 
corolla rosea (ex collectore), infundibuliformis, tubo 7-8 mm. longo 
basi manifeste gibboso supra paullo ampliato extus basi excepta sparse 
setoso-hirsuta, intus a medio ad faucem villoso-hirsuto, lobis late ovatis 
apice rotundatis 3 mm. longis glabris; stamina medio tubo affixa, an- 
theris 2.5 mm. longis faucem non attingentibus, filamentis glabris 
brevissimis; stylus medium tubum non superans, glaber. 

As Leveille's description is very brief and inaccurate particularly in 
regard to the calyx which he describes "calyce ciliato," apparently taking 
the bracts for the calyx, I have given above a more complete description. 
The species seems nearest to L. nubigena Rehd., from which it chiefly 
differs in the bracts being quite glabrous except ciliate toward the base, 
in the large calyx, in the corolla being sparingly setose-hirsute outside, 
not short-pubescent and glandular, in the hirsute pubescence at the 
mouth with the anthers much below the mouth, not just reaching the 
mouth as in L. nubigena, and in the glabrous style. 

Lonicera lanceolata Wall ich in Roxburgh, Fl. Ind. ed. 2, 2: 177 
(1824). — Leveille, Cat. PI. Yun-Nan, 27 (1915). 

Lonicera acrophila Leveille in Bull. Geog. Bot. 24 (no. 301): 289 
(1914); Cat. PI. Yun-Nan, 27 (1915).— Synon. nov. 

China. Yunnan : haut plateau de Je-ma-tchouan, 3200 m.. 



338 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

E. E. Maire, July 1912, "arbre moyen, fl. roses" (holotype of L. acro- 
phila; merotyp>e in A. A.). 

According to Maire this is a medium-sized tree, but by most collectors 
it is described as a shrub, 4-8 ft. tall. 

Lonicera Koehneana Rehder in Sargent, Trees & Shrubs, 1:41, t. 
21 (1902). — Leveille, Cat. PI. Yun-Nan, 27 (1915). 

Lonicera gynopogon Leveille in Bull. Geo^. Bot. 24(no. 301): 289 
(1914) ; Cat. PI. Yun-Nan, 27 (1915). — Synon. nov. 

China. Yunnan: brousse derriere Tong-tchouan, alt. 2550 m., 
E. E. Maire, May 1912, "Lonicera non grimpant, rameaux courts et 
greles, fl. blanc-jaune" (holotype of L. gynopogon; photo, in A. A.); 
haies, plaine de Tong-tchouan, alt. 2500 m., E. E. Maire, May [1912], 
"fl. de Lonicera, mi-blanche, mi-jaune" (in herb. Leveille under L. gyno- 
pogon; photo in A. A.); haies de Tcheou-kia-tse-tang, alt. 2500 m., 
E. E. Maire, "arbuste buissonante, haut 2 m., feuilles molles, velues et 
blanches en dessous, fl. mi-blanches, mi-jaunes, inodores" (in herb. 
Leveille under L. gynopogon; duplicate in A. A.). 

Specimens from the same locality and partly apparently of the same 
collection have been distributed by the Arnold Arboretum under Maire, 
no. 142 and no. 286. 

Lonicera Pampaninii Leveille in Fedde, Rep. Spec. Nov. 10: 145 
(1911); Fl. Kouy-Tcheou, 64 (1914); Cat. PI. Yun-Nan, 27 (1915). 
Lonicera Hcnr\<i var. sctuligcra W. W. Smith in Not. Bot. Card. 
Edinb. 10:47 (1917). 

China. K w e i c h o u : mont de Lou-tsong-koan, Tsin-gay, ro- 
cailles a Che-tiou-tchay, Gan-pin, buissons et rochers de la montagne, 
L. Martin in herb. Bodinier, no. 1623, June 10, 1897 and June 27, 1899, 
"fleurs jaunes" (syntypes; merotype in A. A.). 

This species is similar to L. Henryi HemsL, but is easily distinguished 
by the slenderer corolla-tube densely clothed with reflexed yellowish 
hairs; in the subsessile or sessile flowers with subulate pilose bracts 
exceeding the pilose calyx-lobes; the leaves which closely resemble those 
of L. Henryi are pilose on the midrib above and below otherwise glabrous 
even on the margin. 

This species has been collected in Kweichou also by Y. Tsiang near 
Tsun-yi and Pin-fa; nos. 5277 and 5377; also Steward, Chiao & Cheo 
no. 271 from Tsun-yi is probably the same, but it has no flowers. 

Lonicera macrantha Sprengel, Syst. Veg. 4- : 82 (1827). — Leveille, 
Fl. Kouy-Tcheou, 63 (1915). 

Lonicera Guilloni Leveille & Vaniot, in Bull. Soc. Bot. France. 51: 
cxliv (1904). 



1935] REHDER, LIGNEOUS PLANTS DESCRIBED BY LEVEILLE 339 

China. K w e i c h o u : Pin-fa, /. Cavalerie, no. 1015, May 28, 
1903, "fl. blanches et jaunes au veillisant, sans odeur" (holotype of L. 
Guilloni; photo, in A. A.). 

Cavalerie no. 1015 is cited by Leveille in his Flore du Kouy-Tcheou 
under L. macrantha (1. c), but the name L. Guilloni is not mentioned. 
The specimen differs somewhat from typical L. macrantha in the shorter 
and slighter pubescence of the branches and in the scarcely ciliate leaves. 

Lonicera Esquirolii Leveille, Fl. Kouy-Tcheou, 63 (1914). 

China. K w e i c h o u : without locality, /. Esquirol, no. 889, 
June 1903, ''il. jaunes apres floraison" (holotype; photo, and merotype 
in A. A.). 

This species seems most nearly related to L. ferruginea Rehd., but is 
easily distinguished by the shorter, not hirsute pubescence and the 
glabrous ovary. From L. inodora W. W. Sm. it differs in the glabrous 
style, the glandular pubescence of the corolla, the sessile or subsessile 
inflorescence and in the setulose pubescence extending over the whole 
under surface of the leaf. 

Lonicera japonica Thunberg, Fl. Jap. i 
PI. Yun-Nan, 27 (1915); in Mem. Acad. 
12: 545 (Cat. PI. Kiang-Sou, 5) (1916). 

Lonicera Fauriei Leveille & Vaniot in Fedde, Rep. Spec. Nov. 5: 100 
(1908).— Synon. nov. 

Japan. Nippon: in littore Shiogama, U. Faurie, no. 6823, Oct. 
1905 (holotype of L. Fauriei; photo, and merotype in A. A.). 

Leveille compares his species with L. bracteolaris Boiss. & Buhse and 
describes the fruit as having 3 persistent hairy styles; he apparently 
mistook for styles the sepals which in one of the fruits appear to be only 
three, the other two not being clearly visible. 

Lonicera joinnanensis Franchet in Jour, de Bot. 10: 310 (1896). 
Lonicera Mairci Leveille in Bull. Bot. Geog. 24:289 (1914).— 

China. Yunnan : collines herbeuses autour de Tong-tchouan, 
alt. 2550 m., E. E. Matte, fl. blanc-jaunatre" (holotype of L. Mairei; 
merotype in A. A.). 

The branches of Maire's specimen are apparently from different 
plants; one has the leaves quite glabrous beneath as in the type of 
L. yunnanensis, while in the other they are slightly pubescent beneath 
and are referable to var. tenuis Rehd., but there is no difference in the 
size of the leaves. 



340 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

COMPOSLTAE 

Pertya Bodinieri Vaniot in Bull. Acad. Intern. Geog. Bot. 12: 116 
(1903). — Beauverd in Bull. Soc. Bot. Geneve, ser. 2, 1:386, fig. 6 
(1909). — Leveille, Cat. PI. Yun-Nan, 47 (1915). 

China. Yunnan: environs de Yunnan-fou, dans les ravines de 
la montagne, E. Bodinier, no. 10, Jan. 27, 1897, "tiges sous-ligneuses, 
de 0.6-1 m., fl. roses" (holotype; merotype [from herb. Leveille] and 
photo, of isotype [in herb. Paris] in A. A.). 

Though this is not one of Leveille's species I have included it here, 
since the type is in the herb. Leveille. 

Pertya Esquirolii Leveille (in Bull. Geog. Bot. 24: 251 (1914); Fl. 

Kouy-Tcheou, 100 (1914) from Kweichou, based on Esquirol no. 3633 

is an herbaceous plant and belongs to Ainslea; it seems very near or 

identical with A. rubrijolia Franch. which I have not seen. 

(To be continued) 



REHDER, HUODENDRON 



HUODENDRON, A NEW GENUS OF STYRACACEAE 



With plates 151 and 152 and one text figure. 

Huodendron, gen. nov. 

Flores hermaphroditi, actinomorphi, pentameri; calycis tubus ovario 
adnatus, dentibus 5 triangularibus vel ovatis circiter dimidium tubum 
aequantibus; petala 5, initio basi coherentia, demum libera, lineari- 
oblonga, anguste imbricata vel valvata, sub anthesi revoluta; stamina 
7-10, uniserialia, libera, petalis subaequilonga, sed ob petala revoluta 
valde exserta, filamentis complanatis linearibus, antheris anguste ob- 
longis introrsis, loculis distinctis, connectivo cum filamento continue et 
supra antheras in appendicem conspicuum tri- vel rarius bidentatum 
elongate; ovarium inferum, triloculare; styli 3, triente inferiore vel fere 
ad apicem connati, stigmatibus capitellatis; ovula in quoque loculo 
numerosa, axi centrali affixa, erecta. Fructus capsularis, ovoideus, 
parva, triente infra apicem sepalis circumcincta, trilocularis, loculicide 
dehiscens, valvis interdum demum septicidis, endocarpio crustaceo, exo- 
carpio tenui; semina numerosa, scobiformia, minuta, oblonga vel ellip- 
tico-oblonga, leviter complanata, testa tenui reticulata, basi et apice 
fimbriata et saepius ad marginem sparse breviterque fimbriata, albumi- 
nosa, embryo centralis, rectus. — Arbor vel frutex ramis gracilibus, 
gemmis parvis nudis pubescentibus; folia decidua, altema, petiolata 
estipulata, ovato-elliptica vel ovato-oblonga, acuminata, basi cuneata, 
integra vel remote minuteque denticulata, glabra vel fere glabra, penni- 
nervia, nervis curvatis anastomosantibus; inflorescentiae terminales et 
axillares, paniculatae vel subcorymbosae, ebracteatae et ebracteolatae, 
floribus satis parvis albis graciliter pedicellatis; capsula parva, pedicello 

Ab aliis Styracacearum generibus, petiolis et staminibus liberis vel 
fere liberis, filamentis supra antheram in appendicem 3 -vel 2-dentatum 
elongatis, stylo 3-fido, capsula valvis 3 dehiscente, seminibus scobi- 
formibus numerosis bene distincta. Ob semina numerosa Alniphyllo 
affinis videtur, sed petalis et staminibus liberis, stylo trifido, connectiva 
appendiculato, capsula 3-loculari subinfera, seminibus scobiformibus 
circiter 1 mm. longis facile distinguitur. 

Type Species: Huodendron tibeticum (Anthony) Rehd. 

Distribution : The genus is restricted to southern China and extends 



342 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

northwest across the border into southeastern Tibet and northeastern 
Burma and south into northern Tonkin, where it occurs near Lao-kay, 
about 150 km. southeast of Mengtze. Within China it ranges from 
western Yunnan through southern Kweichou, to Kwangsi and Kwang- 
tung. Of the two species H. tibetkum is restricted to southeastern 
Tibet, about N. Lat. 29°, while H. biaristatum ranges from northeastern 
Burma to Kwangtung and extends south into Tonkin; it does not seem 
to occur north of N. Lat. 25°. 

The two species now known of the new genus were originally both 
referred to the genus Styrax to which the flowers bear a great resem- 
blance, but the fruit is entirely different. In Styrax the fruit is inde- 
hiscent or irregularly dehiscent and contains only one or two rather large 
subglobose or ellipsoid seeds, while the fruit of Huodendron resembles 
strongly that of some Saxifragaceae-Hydrangeae, as Deutzia and Hy- 
drangea, in shape and size and dehiscence of the capsule and in the 
numerous scobiform seeds; also the divided style recalls Saxifragaceae, 
and in some species of Deutzia the flattened filaments are elongated 
beyond the anther or are dentate at the apex. The petals and stamens 
fall off separately after anthesis, though in bud they are cohering at the 
very base; in Styracaceae free stamens and petals are very rare. Any 
doubt, however, one might have in regard to the affinity of Huodendron, 
is convincingly set at rest by the nodal structure of the stem, which 
shows the unilacunar nodes characteristic of all Ebenales, while the 
Rosales have trilacunar or quinquelacunar nodes, as pointed out by Dr. 
I. W. Bailey to whom I am indebted for the examination of the stem. 

As type of the genus I have selected Huodendron tibetkum, because 
this species represents the distinctive characters from Styrax and other 
allied genera in a more pronounced degree, particularly by the deeply 
divided style and by the absence of stellate or fascicled pubescence and 
also in the distinctly corymbose inflorescence. 

For the loan of additional specimens supplementing the material in 
the herbarium of the Arnold Arboretum (A. A.), I am indebted to Dr. 
E. D. Merrill of the New York Botanical Garden (N. Y.), Dr. H. L. 
Mason of the University of California (U. Calif.) and to Sir William 
Wright Smith of the Royal Botanic Garden of Edinburgh (Edinb.). 

I take pleasure in associating with this new genus the name of Dr. 
H. H. Hu, director of the Fan Memorial Institute of Peiping, one of the 
foremost and active Chinese botanists, who has contributed and is still 
contributing extensively to our knowledge of the flora of China. 

Huodendron tibeticum (Anthony), comb. nov. 
Styrax tibcticus Anthony in Not. Bot. Gard. Edinb. 15: 245 (1927). 



19351 



REHDER, HUODENDRON 



Arbor vel frutex 6-25 m. altus, ramis gracilibus teretibus vel apicem 
versus leviter complanatis glabris; folia alterna, sed interdum apicem 
ramulorum versus subopposita, decidua, papyracea, elliptico-ovata vel 
oblongo-ovata vel ovato-lanceolata, 6-11.5 cm. longa et 2.5-4 cm. lata, 
longe acuminata apice mucronulata, basi late cuneata, Integra, nervis 
utrinque leviter elevatis, costa apicem versus supra leviter 




1, subtus manifeste elevata ; petioli glabri, 5-10 
mm. longi, supra leviter canaliculati. Inflorescentia glabra, corymboso- 
paniculata, terminalis 5-7 cm., lata, laterales cum pedunculo 1.5-3 cm. 
longo 4-8 cm. longa et 2.5-5 cm. lata; pedicelli graciles, 3-5 mm. longi, 
ut ramuli glanduloso-verruculosi; calycis tubus cupuliformis, glanduloso- 
verruculosus, 1 mm. longus, dentibus triangulari-ovatis dimidium tubum 



344 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

subaequantibus ciliolatis; petala valvata, lineari-oblonga, 6-7 mm. longa 
et 1-1.5 mm., lata, obtusiuscula, extus tomentosula, intus fere glabra, 
sub anthesi revoluta; stamina petiolis subaequalonga, filamentis 4-5 
mm. longis intus triente inferiore excepto villosis extus glabris, antherae 
1.25-1.5 mm. longae, glabrae, apice tridentato circiter 1 mm. longo, 
dente medio lateralibus plerumque breviore, styli in triente inferiore vel 
ad medium connati, graciles, glabri; discus glaber. Capsula pedicello 
plus minusve recurvo suffulta, ovoidea, 3 mm. longa, fusco-brunnea, 
subinfera; semina brunnea, circiter 1 mm., longa. 

Southeastern Tibet. T s a r o n g : Salween and Kiu-chiang 
divide, northwest of Si-chi-to, Lat. 28^ 35' N., Long. 98° 30' E., alt. 
10-11000 ft., G. Forrest, no. 21648, June 1922, "shrubby, 20-30 ft., 
flowers fragrant, white, in open thickets by streams" (holotype in herb. 
Edinb.); same locality, G. Forrest, no. 22882, Oct. 1922 (paratype in 
herb. Edinb.); Salween and Irrawaddi divide, near banks of Salween 
at Champutong, forests, alt. 7000 ft., /. F. Rock, no. 22020, May-July 
1932, tree 70-80 ft. tall, flowers white (A. A., N. Y., U. Calif.) ; moun- 
tains west of Champutong, forests of upper Salween River, alt. 9000 ft., 
/. F. Rock, no. 22474, Oct. 1932 (A. A., N. Y., U. Calif.). 

This species has a very restricted distribution and is apparently con- 
fined to the mountains of extreme southeastern Tibet between the head- 
waters of the Irrawaddi and Salween Rivers. In some of its characters, 
particularly by the deeply divided style and by the absence of stellate 
or fascicled pubescence is it farther removed from other styraceous 
genera than the more widely distributed H. biaristatum. The fruiting 
branch of this species has some resemblance to certain species of Deutzia. 

Huodendron biaristatum (W. W. Sm.), comb. nov. 
Styrax biaristatus W. W. Smith in Not. Bot. Card. Edinb. 12:233 
(1920).— C. E. C. Fischer in Kew Bull. Misc. Inform. 1933:365. 

Frutex vel arbor 6-12 m. altus, ramis gracilibus hornotinis initio 
tomentosulis demum glabrescentibus, vetustioribus flavido-cinereis vel 
fusco-cinereis cortice demum rimoso vel fibroso vestitis. Folia alterna, 
papyracea, oblonga vel elliptico-oblonga vel obovato-oblonga, 8-17 cm. 
longa et 2.5-6 cm. lata, acuminata, basi cuneata, margine minute et 
remote denticulate vel Integra, supra luteo-viridia, opaca, costa fascicu- 
lato-pilosula excepta glabra, subtus vix pallidiora, axillis saepe barbulatis 
exceptis glabra, costa supra leviter impressa subtus elevata, nervis utrin- 
secus 5-9 arcuatis piargine anastomosantibus supra vix infra manifeste 
elevatis, venulis subtus elevatis; petioli 6-15 mm. longi, supra tantum 
vel undique fasciculato-pilosi. Inflorescentiae terminales et axillares, 
paniculatae, multil^orae, 3-10 cm. longae, ebracteolatae, cinereo- 



1935] REHDER, HUODENDRON 34S 

tomentellae; pedicelli 2-5 mm. longi; calyx cupuliformis, tomentellus, 
tubo 1-1.5 mm. longus, dentibus late triangularibus acutiusculis tubo 
brevioribus; petala imbricata, anguste oblonga, 6-9 mm. longa et 2-2.5 
mm. lata, utrinque tomentella; stamina petalis subaequilonga, filamen- 
tis compressis utrinque dense pilosulis circiter 3 mm. longis, antheris 
glabris 2 mm. longis connective dorse puberulo in appendicem triden- 
tatum vel rarius bidentatum elongata dentibus lanceolatis acutis medio 
plerumque minore ; stylus staminibus paullo longior, crassus dense pilo- 
sulus, apice 3-lobata; ovarium semisuperum. Capsula ovoidea, resu- 
pinata, 4-5 mm. longa, cinereo-tomentella, in triente superiore sepalis 
persistentibus cincta; semina 1-1.25 mm. longa, flavo-fusca. 

China. Yunnan: in thickets in ravines on the western flank 
of the Shweli-Salween divide, Lat. 25° 40' N., alt. 9000 ft., G. Forrest, 
no. 18020, May 1919, "shrub 20-30 ft., flowers fragrant, creamy-yellow" 
(syntype in herb. Edinb.); side valleys of the Shweli-Salween divide, 
Lat. 25° N., alt. 8000 ft., G. Forrest, no. 17894, June 1919, "shrub 
10-20 ft., flowers immature" (Edinb., A. A.) ; N'Maikha-Salween divide, 
at Ho-tou, in thickets and open forests, Lat. 25° 55' N., alt. 7-8000 ft., 
G. Forrest, no. 18400, Aug. 1919, "shrub 12-18 ft., in fruit" (syntype 
in herb. Edinb.) ; same locality, G. Forrest, no. 18833, Nov. 1919 (syn- 
type in herb. Edinb.); Mengtze, S. E. mountain forests, 6000 ft., A. 
Henry, no. 10764 "tree 15 ft." (syntype in herb. Edinb., A. A., N. Y.) ; 
Mengtze, A. Henry, no. 13662a, "shrub 10 ft." (syntype in herb. Edinb.; 
A. A., N. Y.) ; south of Red River, A. Henry, no. 13662, "tree 40 ft." 
(syntype in herb. Edinb.; A. A.); Shweli-Salween divide, Lat. 25° 10' 
N., Long. 98° 50' E., alt. 9000 ft., in open thickets and forests, G. For- 
rest, no. 26108, Dec. 1924, "tree 30-40 ft." (Edinb., N. Y.); without 
precise locality, G. Forrest, no. 26108, 1924-25 (Edinb., N. Y.). 
K w e i c h o u : Waichai, Tuh-shan, near border of Kwangsi, alt. 330 
m., in densely shaded ravine, Y. Tsiang, no. 6686, Aug. 25, 1930, "tree 
6 m., diam. of trunk 12 cm., bark pale gray" (A. A.). Kwangsi: 
Chin-fong, Lin-yuin-hsien, valley forest, alt. 1300 m., Steward & Cheo, 
no. 336, May 6, 1933, "tree 7 m., flowers white, fragrant" (A. A., N. Y.) ; 
Ta-tse-shan, Yung-hsien, forest, alt. 540 m., Steward & Cheo, no. 843, 
Aug. 21, 1933, "tree 9 m., fruit gray" (A. A., N. Y.). 

Burma: Myitkyina Distr.,Htangan, 3100 ft., Sukoe per C. E. Parkin- 
son, no. 9197 ; Pyet Pass, 7200 ft., Sukoe per C. E. Parkinson, no. 10115 
(exC. E. C. Fischer,!, c). 

Tonkin: route de Lao-kay a Chapa, alt. 1500 m., /I. Petelot, no. 3803, 
Aug. 1930 (N. Y.) ; massif du Fan-tsi-pou, chemin du col de Lo-qui-ho, 
environs de Chapa, alt. 1400 m., A. Petelot, no. 4373, Sept. 1931 (N. Y.). 



346 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

This species is readily distin^ished from H. tibeticum by the pu- 
bescent inflorescence, the thicker texture of the leaves, the pubescent 
stout style 3-lobed only at the apex, the shorter filaments pubescent on 
both sides, the broader narrowly imbricate petals pubescent on both 
sides and the tomentulose capsules. The fact that the petals in one 
species of this genus are valvate and in the other imbricate is not un- 
usual in Styracaceae, for both kinds of aestivation are found in Styrax. 
The stamens are mostly 3-toothed at the apex, but the middle one is 
often shorter than the lateral ones; two teeth, as implied by the specific 
epithet, are only occasionally found. 

The leaves of H. biaristatum show some variation in dentation, tex- 
ture, pubescence and in the number of veins. The Forrest specimens 
have remotely denticulate leaves and are of rather thin texture, the 
leaves of the Henry specimens are occasionally furnished with minute 
denticulations reduced to a mucro, but are mostly entire like the other 
specimens and like those are of thicker chartaceous or subcoriaceous 
texture. The midrib is usually impressed and puberulous like the 
petiole, but in Petelot 4373 from Tonkin the midrib is glabrous except 
slightly puberulous toward the base and slightly elevated and quite 
glabrous toward the apex, also the lateral veins are slightly elevated and 
number about 5 pairs, while the leaves of the other specimens have 
mostly 6 or 9 pairs; by these characters this Petelot specimen approaches 
the following variety and connects it with the typical form. 

Huodendron biaristatum var. parviflorum (Merrill), comb. nov. 
Styrax parviflora Merrill in Jour. Arnold Arb. 8: 15 (1927). 

A typo recedit praecipue ramulis foliis petiolisque glabris foliis magis 
coriaceis integris nervis utrinsecus 4-6, costa media nervisque supra 
glabris et elevatis, venulis subtus minus conspicuis. 

China. Kwangtung : Lung-t'au Mountain, near lu, in 
forest, Canton Christian College, nos. 12070 (holotype in hb. N. Y.; 
A. A.) and 12349 (paratype in hb. N. Y.; A. A.). 

The flowers and fruits of the Kwangtung specimens, as far as can be 
judged from the rather poor material, are identical with those of typical 
H. biaristatum and the difference in the leaves does not seem sufficient to 
separate the Kwangtung form as a distinct species, considering the fact 
that the leaves of H. biaristatum show considerable variation and transi- 



KOBUSKI, STUDIES IN THEACEAE, ] 



STUDIES IN THEACEAE, I 
EURYA SUBGEN. TERNSTROEMIOPSIS 



With plate 153 

During the past year the author has been making a survey of the 
Old World Theaceae, starting with a critical study of the genus Eurya. 

The genus as a whole has presented some rather difficult problems in 
specific delimitation and in synonymy which cannot be settled until 
more ample material or photographs of types deposited in various for- 
eign herbaria can be had. 

In Szyszylowicz's treatment,' Eurya comprises three sections: 
Cleyera (DC), Freziera (Sw.) and Proteurya Szysz. In 1896, Urban- 
separated E. sandwicensis from Proteurya and made it the type of the 
new genus, Ternstroemiopsis. The following year Engler'' united 
Ternstroemiopsis with Eurya as a new subgenus, elevating, at the same 
time, the three sections of Szyszylowicz to subgeneric rank. At present, 
however, Eurya is generally considered as containing but two subgenera, 
Proteurya and Ternstroemiopsis, while Cleyera and Freziera represent 
distinct genera, the former Asiatic and American, the latter exclusively 

In this paper, the subgenus Ternstroemiopsis is considered. This is 
distinguished from the subgenus Proteurya by the spiral arrangement 
of its leaves, the thick glandular sepals, fleshy petals and stamens whose 
anthers are twice as long as the filaments. Proteurya is characterized 
by two-ranked leaves, petals more or less membranous and anthers as 
long as or shorter than the filaments. Geographically also Ternstroemi- 
opsis is distinct being confined solely to the Hawaiian Islands while Pro- 
teurya, although found in nearly all the Pacific islands and Asia, does 
not invade the Hawaiian group with a single species. 

The institutions from which material for this study was borrowed 
along with the abbreviations used in this paper, are as follows: herbarium 
of the Arnold Arboretum of Harvard University (AA), herbarium of Otto 
Degener (D), Gray Herbarium of Harvard University (Gr), herbarium 
of the New York Botanical Garden (NY). 

iSzyszylowicz in Engler & Prantl, Nat. Pflanzenfam. III. 6: 189 (1893). 

2Urban in Ber. Deutsch. Bot. Gesell. 14: 49 (1896). 

3Engler in Engler & Prantl, Nat. Pflanzenfam. Nachtr. 1: 247 (1897). 



B. Small trees or erect shrubs 1. E. sandwiccnsis 

BB. Prostrate shrubs la. E. sandwiccnsis var. prostrata 

AA. Leaves cuneate at base, acute at apex 

C. Leaves 3.5-4.7 cm. long, 1.2-1.7 cm. wide 2. E. Dcgcncri 

CC Leaves either larger or smaller than C 

D. Leaves 5.3-8.7 cm. long, 2.5-3.5 cm. wide 

2a. E. Dcgcncri f. qraudifoUa 
DD. Leaves 3.0-4.5 cm. long, 0.7-1.2 cm. wide 

2b. E. Dcgcncri i. stcnophylla 
L Eurya sandwiccnsis A. Gray, Bot. U. S. Expl. Exped. 1838- 
1842, 1: 209 (1854). — H. Mann in Proc. Amer. Acad. Arts Sci. 7: 156 
(Enum. Hawaiian PI.) (1867); Mem. Boston Soc. Nat. Hist. 1: 534, 
539 (1869).--Hillebrand, Fl. Haw. Isl. 41 (1888). — Drake del 
Castillo, 111. Fl. Ins. Maris Pacif. 117 (1890). — Szyszylowicz in Engler 
& Prantl, Nat. Pflanzenfam. iii, 6: 190 (1893). — A. A. Heller in Minn. 
Bot. Studies, 1:856 (1897). — Rock, Indig. Trees Haw. Isl. 308 
(1913). — Melchior in Engler & Prantl, Nat. Pflanzenfam. Ed. 2, 
21: 147 (1925). 

Eiirya sandwiccnsis A. Gray var. scssilifolia A. A. Heller in Minn. 

Bot. Studies, 1: 856 (1897), as a synonym. 
Tcrustrocmiopsis sandwiccnsis Urban in Ber. Deutsch. Bot. Ges. 
14:49 (1896). 
Small trees, 5-6 m. in height, occasionally shrubby in higher altitudes, 
2-3 m.; branches crowded with leaves, ultimate branchlets strigose; 
leaves oblong, elliptical or obovate, coriaceous, glabrous, occasionally 
strigosely hairy on midrib, 4.5-9.0 cm. long, 1.5-3.7 cm. wide, on short 
petioles 2-3 mm. long, sometimes subsessile, obtuse or rounded at the 
apex, more or less cordate, occasionally truncate at the base, closely 
serrulate with inflexed mucronulate teeth, veins and veinlets finely 
reticulate beneath, reddish brown in color; flowers solitary, occasionally 
two in axils, nodding, ebracteolate, pedicels approximately 5 mm. long; 
calyx purplish brown, quite coriaceous, subtended by two small unequal 
bracts; sepals five, unequal, 3-4 mm. long, persistent, suborbicular, thick 
in central portion, membranous, lighter in color and slightly glandular 
on margin, occasional strigose hairs on external surface; corolla pale 
yellow or cream-color, imbricated; petals five, obovate, 5-6 mm. long, 
united at base, somewhat fleshy in central portion; stamens in staminate 
flowers 10-15, slightly adnate to base of corolla, filaments distinct, half 
as long as the oblong mucronate anthers; staminodia in pistillate flowers, 
five sometimes six, 2-3 mm. long; pistil having three or occasionally 



1935] KOBUSKI, STUDIES IN THEACEAE, I 349 

four styles, sometimes connate nearly to stigma, usually divided; stig- 
mas three (or four); ovary glabrous, 3-celled, axial placentation ; fruit 
a globose berry, 7-10 mm. across, dark blue-black, many-seeded; 
mature persistent, subcordate calyx-lobes 8 mm. long, 7.5 mm. across at 
widest portion, lobes at base lighter in color and more membranous. 
Specimens Examined: 

Hawaiian Islands. O a h u : Nuuanu-Pali, U. Faurie, no. 284, 
October 1909 (AA) ; on mountains behind town of Honolulu, Ww Rich, 
collected in 1840 (type) (Gr, NY); exact locality lacking, M. J. 
Remy, no. 562, collected 1851-1855 (Gr) ; data lacking, M. J. Remy 
(NY) ; exact locality lacking, H. Mann & W. T. Brigham, no. 524, col- 
lected 1864-1865 (Gr, NY); exact data lacking, W. Hillebrand (Gr); 
precise data lacking, C. Gaudkhaud, collected probably 1836 (Gr); in 
rain-forest from Kahana church up ridge to summit of mountain south- 
east of Kahana Bay, 0. Degener, no. 8680, July 3, 1932 (AA, D); 
Waipio, Waiawa Ridge, on Dicranopteris-covered ridge, O. Degener & 
Dr. C. L. Shear, no. 9838, March 5, 1928 (tree 15 ft.; fruit inky blue- 
black) (AA, D); open forest in Dicranopteris tangle, Manana Gulch 
ridge, O. Degener, W. Bush & K. K. Park, no. 8679, October 2, 1932 
(AA, D) ; on and near the summit of Konahuanui, A. A. Heller, no. 2240, 
May 2, 1895 (NY); lower slopes of Konahuanui, above Manoa, A. A. 
Heller, no. 2311, May 13, 1895 (AA, NY, Gr) ; ridge west of Kalihi 
valley, C. A^. Forbes, no. 1483.0, March 17, 1910 (NY) ; ridge between 
Pololo and Waialue iki, C. N. Forbes, no. 2408.O, January 30, 1917 
(NY); Koolau Mts., Pumaluu, /. F. Rock, no. 627, December 3-10, 

1908 (Gr); Pumaluu, /. F. Rock, no. 843, December 1908 (NY); pre- 
cise data lacking; /. F. Rock, collected 1910 (Gr). Kauai: Mt. 
Waialeale, alt. 5200 ft., /. F. Rock, no. 8864, October 20, 1911 (Gr, 
NY) ; along stream-beds, Kaholuamano, /. F, Rock, no. 5499, Septem- 
ber 1909 (Gr, NY); Kaholuamano, /. F. Rock, collected March 3-10, 

1909 (NY); Hanapepe, U. Faurie, no. 286, December 1909 (AA). 
M a u a i : Honakahau Drainage Basin, C. N. Forbes, no. 421.M, 
September 25-October 17, 1917 (NY). Hawaii : Kilauea, near 
fern-forest, O. Degener, no. 8678, November 10, 1929 (AA, D). 

In the whole genus this species is probably the most outstanding. 
Although confined to the Hawaiian group, it has been found in nearly 
all the islands from which material has been collected. The flowers and 
fruit are nearly twice the size of any other species. Along with this size 
character can be mentioned the distinct reddish reticulate veining of the 
lower surface of the leaf and the subcordate or truncate base of the leaf. 

The collections of Otto Degener made during the last few years on 



350 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

the various islands of the group had great influence in the decision 
finally to place the majority of Hawaiian specimens in this species. His 
specimens were so ample that it was possible to make two and even three 
sheets of each for the herbarium of the Arnold Arboretum. These sets 
of material show great gradation in leaf-size — a character which might 
cause some, especially in this genus, to describe new species. 

Faurie's specimen, no. 286, according to the collector, was made at 
Hanapepe, Oahu. This probably is a mechanical error made in trans- 
ferring the field notes to the herbarium label. Hanapepe is on the 
island of Kauai and the collection date of no. 286 agrees with other 
material collected by Faurie on Kauai. 

la. Eurya sandwicensis A. Gray var. prostrata, var. nov. 

A typo recedit habitu prostrato et foliis remotis. 

Hawaiian Islands. M o 1 o k a i : At edge of windswept forested 
pali, Ohialele Pali, O. Degener, no. 8676 (type, k\) May 10, 1928 
(more or less trailing along ground with branches sometimes eight feet 
long; flowers yellow, difficult to distinguish plant from Vaccinium) (AA, 
D); Pelekunu trail, C. N. Forbes, no. 249.Mo, July 1912 (NY). 

A quotation from a recent letter from Otto Degener, the collector of 
the cited type throws considerable light on this variety and the species 
E. sandwicensis : — "Eurya, as I have found it on Oahu, grows as an 
erect small tree with very dense foliage. It is rare, and where found, 
usually grows in openings in the lower forest, covered over with Glei- 
chenia. Rainfall would be moderate." 

"The Molokai specimen I have found nowhere except in a typical 
dense extremely rainy rain-forest, and curiously enough, not anywhere 
in that region but only on the brink of a cliff extending for several miles. 
In short, it grows on the very "backbone" of Molokai where the rain 
and fog drive violently over the mountain crest. I collected five months 
on Molokai and do not remember seeing any Eurya except in that one 
type of locality. The rain-forest reaches up to this cliff and it is among 
the shrubs and small trees immediately overlooking the cliff that the 
Eurya is to be found. The plant sprawls rather than creeps, producing 
slender branches of unusual length— possibly 12 feet— with its leaves 
spaced far apart." 

The collector remarked further that at first on seeing sterile plants, 
he thought this variety to be a low-growing Vaccinium. However, later 
on finding flowering material, he discovered it to be an Eurya. He 
suggested it as a variety or possible new species. 

The second cited specimen, Forbes no. 249. Mo, resembles the type 



KOBUSKI, STUDIES IN THEACEAE, I 351 

i of leaves. However the habit of the plant was not 
given by the collector, but it appears to be prostrate. 

2. Eurya Degeneri, spec, nov.; a E. sandwicensi A. Gray foliis 
ellipticis 3.5-4.7 (2.5-6.5) cm. longis, 1.2-1.7 (1.2-2.2) cm. latis, apice 
acutatis et emarginatis, basi cuneatis recedit. 

Branches covered with leaves especially at ends, ultimate branchlets 
sparsely strigose; leaves elliptic, coriaceous, 3.5-4.7 (2.5-6.5) cm. long, 
1.2-1.7 (1.2-2.2) cm. wide, acute at the apex, emarginate, cuneate at 
the base, closely serrulate with inflexed mucronulate teeth, conspicu- 
ously reticulate on under surface, veins and veinlets reddish brown in 
color, especially near base of leaf where color spreads into the leaf; 
petiole 3 mm. long; mature flowers unknown, bud resembling E. sand- 
wicensis sufficiently in coriaceous character and color of calyx with 
occasional strigose hairs, character and number of stamens and corolla 
to show it to be typical of the genus and closely related to E. sandwi- 
censis; berry (probably not mature) blue-black, 5 mm. across with the 
persistent styles separate to near the base, 3-celled, axial placentation, 
many-seeded. 
Specimens Examined: 

Hawaiian Islands. Kauai: open forest, Waineke swamp 
Kokee, O. Degener, no. 8675 (type AA) July 1, 1926 (AA, U); high 
plateau of Waimea, Halemanu to Kaholuamano, J. August Kusche, nos. 
28, 139, 140, collected in 1919 (AA); Kilauea, U. Fawie, no. 285, 
January 1910 (AA); west side Waimea Drainage Basin, Kanaikinaua, 
C. iV. Forbes, no. 1016.K, July 3-August 18 (1917) (AA, NY). 

This species is very closely allied to E. sandwicensis. The leaf char- 
acters are most distinctive between the two species. Eurya Degeneri has 
elliptical leaves, cuneate at the base, acute and emarginate at the apex. 
On the other hand, Ettrya sandwicensis has leaves which are oblong or 
elliptic, subcordate or truncate at base and rounded or obtuse at the 
apex. Eurya Degeneri and its varieties are confined to the island of 
Kauai, while Eurya sandwicensis is found on nearly all the islands in- 
cluding Kauai. 

Otherwise these two species belonging to this distinctive section of 
the genus are very similar. This is especially true in flower and fruit 
characters. Although the mature flowers and fruit were not available 
in E. Degeneri, the material such as it is shows conclusively that there 
is a great resemblance. 

It is a pleasure to dedicate this species to Otto Degener of Hawaii, 
whose recent collections from the islands are extremely fine and whose 
material of Eurya aided tremendously in clearing up this section. 



JOURNAL OF TFIE ARNOLD ARBORETUM [vol. xvi 

Eurya Degeneri Kobuski forma grandifolia (Wawra), comb. 



41 (1888). — Drake del Castillo, 111. Fl. Ins. Maris Pacif. 117 

Eurya sandwicensis A. Gray var. grandifolia Wawra in Flora. 56: 168 
(1873).— J. F. Rock, Indig. Trees Haw. Isl. 308 (1913). 

A typo recedit foliis amplioribus, 5.3-8.7 cm. longis, 2.5-3.5 cm. latis. 
Specimen Examined: 

Hawaiian Islands: Kauai: Wainiha, U. Fauric, no. 298, Janu- 
ary 1910 (AA). 

This large-leaved form has seemed rather evasive to most collectors. 
Hillebrand, Del Castillo, Wawra and Rock have made reference to it in 
literature. The first three in their treatments were working with a single 
specimen, that of Wawra collected at Kealia on the island of Kauai. 
Incidentally we have only the single specimen collected by Faurie (no. 
298) collected at Wainiha. The Faurie specimen was collected on the 
north coast of Kauai while Wawra made his collection on the west coast. 

Rock, although having collected considerably on the islands, never 
encountered this large-leaved form. However, he collected the narrow- 
leafed form cited next. 

These two forms like the species have distinctly cuneate leaf-bases 
and acute apices. Their variation from the species lies chiefly in the 
leaf size. Again, like the species, they are found only on the island of 



2b. Eurya Degeneri Kobuski forma stenophylla, forma nov. 

A typo recedit foliis minoribus angustioribusque, 3.0-4.5 cm. longis, 
0.7-1.2 cm. latis. 
Specimen Examined: 

Hawaiian Islands. Kauai : precise locality and date of col- 
lection lacking, /. F. Rock, no. 17274 (type) (AA). 

Unfortunately, the Rock specimen cited above is sterile and was 
placed in this genus under E. sandwicensis with some hesitation by an 
earlier student. At first, I was quite dismayed because I felt that it 
belonged to a species other than E. sandwicensis, but because of the lack 
of flowers or fruit I hesitated to describe it as new. It was not until 
more material came to my attention that its true affinity with E. Degen- 
eri was discovered. 



;^. 




EuRYA Degeneri Kobuski 



PALMER, TWO NEW SPECIES OF CRATAEGUS 



TWO NEW SPECIES OF CRATAEGUS FROM MISSOURI 



Crataegus hannibalensis, sp. nov. Arbor 6-8 m. alta vel frutex 
arborescens 4-6 m. altus, ramulis annotinis gracilibus vel paulo validis 
glabris fusco-viridibus, spinis numerosis. Folia ovata vel obovata 
vel oblongo-ovata, serrata, apice acuta, basi cuneata in petiolum 8-12 
mm. longum attenuata, 2.5-4 cm. longa, l-l cm. lata, surculorum vali- 
dorum ad 5-6 cm. longa, 4-5 cm. lata, matura firma, crassa, glabra, 
dentata, nervis superne manifeste impressis. Inflorescentiae glabrae, 
laxae, plerumque S-16-florae; flores 14-16 mm. lati, staminibus circiter 
10, stylis 1-3 plerumque 2 ; sepalis lanceolatis integris vel sparse denta- 
tis. Fructus ovoideus, obovoidea-oblongus vel rare subglobosus, 8-10 
mm. longus, 7-8 mm. latus, firmus, viridis denique rubro-luteus raro 
pruinosus, seminibus 2-3 plerumque 2 ovalibus dorso sulcatis. 

A tree 6-8 m, tall, or sometimes an arborescent shrub 4-6 m. tall, with 
intricate ascending or horizontal branches and slender to stoutish flexu- 
ous branchlets, glabrous and olive-green or olive-brown at the end of 
the first season, usually armed with numerous stout, straight or curved 
purplish thorns 3-6 cm. long. Bark gray or pale brown, slightly scaly. 
Leaves obovate, oblong-obovate or oval, acutely pointed, short-acumi- 
nate or rarely rounded at apex, cuneate at the base and attenuate into 
the short 8-12 mm. long petioles, sharply serrate usually nearly to the 
base, glabrous, firm to subcoriaceous at maturity, yellowish-green above 
and slightly paler beneath, with slender but prominent mid-rib and 
5-7 pairs of parallel veins elevated on the under surface and conspicu- 
ously impressed above, those of the fruiting branches mostly 2.5-4 cm. 
long and 2-3 cm. broad, on vigorous sterile shoots often 5-6 cm. long 
and 4-5 cm. broad, and with margins coarsely serrate or dentate. Flow- 
ers in loose glabrous compound 5-16-flowered corymbs, 14-16 mm. in 
diameter; pedicels slender, often glandular, 1-2 cm. long; stamens 
about 10; anthers in specimens examined pale yellow; styles 1-3, usu- 
ally 2 ; calyx-lobes lanceolate, entire or slightly serrate towards the base, 
glabrous without and glabrous or slightly villous within. Fruit oval, 
oblong-obovoid or rarely nearly globose, 8-10 mm. long, 7-8 mm. thick, 
hard and green until late in the season, turning dull red or orange-red 
and becoming mellow when fully ripe late in September, rarely with a 



354 JOURNAL OF THE ARNOLD ARBORETUM I vol. xvi 

slight pruinose bloom. Fruiting calyx sessile or slightly elevated, with 
a broad shallow cavity; calyx-lobes often persistent and appressed, 
tlesh thin; nutlets 1-3 but usually 2, relatively large, oval or elliptic in 
outline, blunt or rounded at the ends, and with broad shallow ridges and 
grooves on the dorsal surface. 




Thickets and borders of woods, in fertile soil, on limestone hills or 
often along bluffs and banks of streams. Crataegus hannibalensis is 
rather abundant in northern Missouri and southeastern Iowa, and it is 
probably more widely distributed. A specimen collected in western 
Ohio seems to belong here. 

This species is conspicuous and easily distinguishable on account of 
its rather large (for the group) yellowish-green leaves with deeply im- 
pressed veins, the pale olive branchlets and comparatively small oval or 
oblong fruit. In their deeply impressed veins the leaves resemble some- 
what certain species of the Punctatae group, but the characters of the 
fruit and flowers and the entire absence of pubescence seem to place it 



1935] PALMER, TWO NEW SPECIES OF CRATAEGUS 355 

clearly in the Crus-galli group. The type specimen is in the herbarium 
of the Arnold Arboretum. 

Missouri: Hannibal (Marion Co.), John Davis, no. 177, Oct. 
6, 1911, May 14, Oct. 10, 1912, Oct. 13, \9U]'E. J . Palmer, no. 20381, 
Sept. 7, 1921; no. 20382 (type), Sept. 7, 1921; no. 20405, Sept. 8, 1921; 
no. 22337, Oct. 24, 1922; south of Hannibal (Ralls Co.), John Davis, 
no. 1645, Oct. 4, 1916; Eolia Pike Co., John Davis, no. 25, Sept. 30, 
1912; no. 2147, Sept. 20, 1913; no. 2149, Sept. 22, 1913; no. 2153, Sept. 
21, 1913; Dumas, Clark Co., B. F. Bush, no. 10139, July 28, 1923; be- 
tween Renick and Clark, Macon Co., E. J. Palmer, no. 35943, May 21, 
1929; between Lancaster and Downing, Schuyler Co., Palmer & Steyer- 
mark, no. 40970, June 30, 1933; Mill Grove, Mercer Co., no. 41270, 
July 4, 1933 ; Shelbina, Shelby Co., no. 40865, June 28, 1933 ; Eagleville, 
Harrison Co., no. 41340, July 6, 1933; St. Francois Co., C. S. Sargent, 
Oct. 5, 1899. Ohio: Springfield, R. E. Horsey, no. 338, May 17, 
Oct. 25, 1915. Iowa: Keokuk, Lee Co., E. J. Palmer, no. 21829, 
Sept. 6, 1922; no. 21831, Sept. 6, 1922; no. 40595, June 25, 1933; Bur- 
lington, Des Moines Co., E. J. Palmer, no. 21800, Sept. 6, 1922. 

Crataegus Danielsii, sp. nov. Arbor 6-7 m. alta vel frutex arbor- 
escens 4-6 m. altus. Folia oblongo-ovata, elliptica vel rhombica, grosse 
serrata, saepe supra medium obscure inaequaliter incisa, apice acuta vel 
i cuneata in petiolum gracillimum 8-15 mm. longum 
ura papyracea sed firma, superne glabra, infra paulo 
villosa, 2.5-4 cm. longa, 1-2.5 cm. lata, ramulorum sterilium ad 5-6 cm. 
longa 3-5 cm. lata. Inflorescentiae laxae, ramosae, paulo villosae, 6-15- 
florae, bracteis linearibus glanduloso-serratis. Flores 14-16 mm. lati, 
staminibus circa 12-15, antheris rubicundis, stylis 2-4, plerumque 3, 
sepalis lineari-lanceolatis integris vel paulo glanduloso-serratis. Fructus 
subglobosus, 8-12 mm. latus, maturus rubicundus; seminibus 2-3 dorso 

A tree 6-7 m. tall, or sometimes an arborescent shrub 4-6 m. tall, 
with erect or ascending intricate branches and slender branchlets, more 
or less villous when young in the typical form, and armed with slender 
thorns 2-3 cm. long. Leaves oblong-elliptic, oblong-obovate or narrowly 
rhombic in outline, sharply and irregularly serrate, often obscurely 
incised above the middle with one or more pairs of shallow lobes or 
unsymmetrical with one or more odd lobes, pointed or acuminate at the 
apex, attenuate at the base into the slender 1-1.5 cm. long petioles, usu- 
ally red as they unfold, and then villous on both surfaces, thin but firm 
at maturity, glabrous above and more or less villous along the veins 
beneath, those of the fruiting branches mostly 2.5-4 cm. long and 1 2.5 



356 



JOURNAL OF THE ARNOLD ARBORETUM 



cm. broad, and up to 5-b cm. long and 3-5 cm. wide on vigorous sterile 
shoots; p)etioles usually sparsely villous, sometimes with a few scat- 
tered glands. Flowers 14-16 mm. in diameter, in loose slightly villous 
compound corymbs; pedicels slender, 8-15 mm. long, glabrous or 
sparsely villous; bracts numerous and conspicuous, narrowly linear, 
finely glandular-serrate on the margins; stamens usually 12-15; anthers 
pink or rose-color in specimens examined; styles 3 4, usually 3; calyx- 
lobes linear-lanceolate, entire or somewhat glandular-serrate towards 




Figure : 



i Danii 



; E. J. Palmer. X 2li 



the base. Fruit subglobose, 8-12 mm. in diameter, pruir 
dull crimson when ripe in late September or October, flesh thin and 
hard; nutlets 2-i^ usually 3, oblong, rounded at the ends and with 
broad shallow grooves and ridges on the dorsal surface. 

Limestone glades and hillsides in the vicinity of Columbia, Missouri. 

Several trees referable to this species have been found, all so far as 
known within a few miles of the type locality. The extremely local 
distribution, the variable and often asymmetrical outline of the leaves 
and the sparse and variable pubescence, nearly or quite absent in some 
specimens, all suggest the possibility of a hybrid origin, and it may have 



1935] PALMER, TWO NEW SPECIES OF CRATAEGUS 357 

originated as a cross between Crataegus crus-galli and C. verruculosa, 
both of which are growing in the immediate vicinity. The specific name 
is for Dr. Francis Daniels, author of a Flora of Columbia, Missouri, and 
vicinity, who first collected it there. 

Missouri: near Columbia, Boone Co., Hawthorn glades, north 
of Columbia, Francis Daniels, Sept. 26, 1902; May 3, 1903; W. H. 
Rickett, no. 8 (Crat. #8), 50 yds. west of Balanced Rock, May 3, 1931 ; 
no. 36 (Crat. #35), north side of Walnut St., west of highway 63, May 6, 
1931; Francis Drouet (Crat. #8, W. H. R. #107), 50 yds. west of Bal- 
anced Rock, Oct. 4, 1931; (Crat. #8, W. H. R. #69), west of Balanced 
Rock, Sept. 15, 1931; (Crat. #35, W. H. R. #71), north side of Walnut 
St., west of highway 63, Sept. 22, 1931; E. J. Palmer, no. 39265 (type). 
May 4, 1931; near Hinton, Boone Co., W. H. Rickett, no. 40 (Crat. 
#39), 4.4 miles north of Hinton, May 17, 1931; no. 43 (Crat. #42), 1.3 
miles south of Hinton, May 17, 1931; 84 (Crat. #54), 3.6 miles north 
of Hinton, Sept. 31, 1931; no. 86 (Crat. #56), north of Hinton, Sept. 
30, 1931; no. 88 (Crat. 42), 1.3 miles south of Hinton, Sept. 30, 1931; 
no'. 91 (Crat. #43), 2.2> miles south of Hinton, Sept. 30, 1931. Type in 
the herbarium of the Arnold Arboretum. All other specimens examined 
are in the herbarium of the University of Missouri. 

In a few specimens examined the young foliage, branches, and in- 
florescence are quite glabrous and in others there is only the slightest 
trace of pubescence in the form of a few scattered hairs on either the 
pedicels, petioles or veins of the leaves. This may be distinguished as 
Crataegus Danielsii f. glabra, f. nov.^ 

Thickets, limestone hills and glades, Boone County, Mo. With the 

W. H. Rickett, no. 39 (Crat. #38), 4.4 miles north of Hinton, Mo., 
May 17, 1931, in the herbarium of the University of Missouri, may be 
taken as the type of this form. 



JOURNAL OF THE ARNOLD ARBORETUM 



NEW HYBRIDS FROM THE ARNOLD ARBORETUM 



X Akebia pentaphylla (Mak.) Makino in Tokyo Bot. Mag. 16: 30 
(1902) = A. quinata Dene. X trifoliata (Thbg.) Koidz. 

Akebia trifoliata var. pentaphylla Makino in Tokyo Bot. Mag. 5: 329 
(1891). 

Artificial hybrids between Akebia trifoliata Koidz. and A. quinata 
Dene, were produced at the Arnold Arboretum in 1932 by Dr. Karl Sax.^ 

Though they have not yet flowered the hybrid seedlings have now 
reached a stage where their intermediate character is clearly evident and 
is in close agreement with Makino's description (1. c.) of X A. penta- 
phylla, a putative hybrid widely distributed in Japan. The hybrids, on 
the whole, resemble A. trifoliata somewhat more closely than they do 
A. quinata. As yet many of the leaves are three-foliolate, although 
leaves with four and five leaflets have been produced. It is of interest 
that Makino originally considered X A. pentaphylla as a variety of A. 
trifoliata (1. c). In this connection one might speculate as to the 
origin of A. trifoliata Koidz. var. australis (Diels) Rehd. Diels" in 
describing the variety commented on its extreme variability, and it occurs 
in a region where both A. trifoliata and A. quinata are native. Further- 
more, it is intermediate between the two species in both leaf and flower, 
though resembling A. trifoliata more closely. It seems not impossible 
that it has resulted through extensive hybridization between A. trifoliata 
and A. quinata in a region where the former is relatively more abundant. 

It should be remembered that the consequences of hybridization be- 
tween two species may be quite different in different parts of their 
ranges, depending upon the relative frequency of the two species, the 
presence of polyploid races within either parent, the adaptability of the 
hybrid to local conditions, etc. Such matters are usually highly specu- 
lative. The production of an artificial hybrid will make it possible, ulti- 
mately, to study such questions experimentally in the genus Akebia. 
E. A. 

Pninus Juddii E. Anderson, hybr. nov. = P. Sargcntii Rehd. 9 X 
yedoensis Mats. $ . 



1935] ANDERSON AND REHDER, NEW HYBRIDS 359 

Intermedia inter parentes, et ab utroque differt praecipue inflorescen- 
tiis 2-6-floris breviter racemosis et breviter pedunculatis, calycis lobis 
sparse et leviter glanduloso-serratis, stylo basi sparse piloso. 

Growing in the Arnold Arboretum under no. 22489 and type specimens 
collected May 5 and 10 and June 3, 1935, are preserved in the herbarium. 

An upright tree with spreading branches. Branchlets glabrous. 
Leaves ovate, acuminate, doubly serrate, dull brownish green when un- 
folding, glabrous throughout. Flowers before the leaves in very short- 
peduncled racemes of two to six, subtended by greenish bracts. Pedi- 
cels with weak scattered hairs at the base. Petals oblong, white or 
whitish, flushed with deep rose pink (Ridgway). Calyx-tube cylindric 
to sub-urceolale, glabrous. Calyx-lobes weakly and irregularly 
glandular-serrate. Style with scattered hairs at the base. Fruit black. 

Among the seedlings of Prunus Sargentii Rehd. (Prunus serrulata 
Lindl. var. sachalinensis [F. Schmidt] Mak.) which have been raised 
from the original trees at the Arnold Arboretum were certain plants 
which are evidently hybrids between that species and other cherries 
which were flowering at about the same time. In the case of one of these 
specimens the evidence for its exact parentage is so clear and the hybrid 
tree promises to be of such horticultural importance for New England 
that it seemed desirable to provide the hybrid with a scientific name. 

I take pleasure in naming the hybrid after the propagator for the 
Arnold Arboretum, Mr. W. H. Judd, whose precise record of the 
material which has passed through his department is of great scientific 
importance. 

The hybrid originated in 1914 at the Arnold Arboretum as a seedling 
of one of the original trees of Prunus Sargentii raised from seed sent from 
Japan by Dr. W. S. Bigelow in 1890. Prunus yedoensis was acquired 
in 1902 and for many years a large specimen stood adjacent to Prunus 
Sargentii, no. 5777. Since their flowering dates usually overlapped it is 
not at all surprising that cross-fertilization should have taken place. Mr. 
Edwin L. Hillier of the West Hill Nurseries, Winchester, England, writes 
me that he has obtained similar hybrids from seed sent him from the 
Arnold Arboretum. Since seed of both Prunus yedoensis and P. Sargentii 
have been distributed very widely for a number of years by the Arnold 
Arboretum, it is quite possible that the hybrid may have turned up in a 
number of nurseries and gardens. 

X Prunus Juddii has proved hardy during the last two phenomenally 
cold winters though it is planted at the edge of one of the coldest spots 
in the Arnold Arboretum. It furthermore holds its flowers longer than 
does P. Sargentii and is a thrifty quick-growing tree. From P. Sargentii 



360 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

it can most easily be distinguished by the greener young leaves, by the 
scattered hairs at the bases of the style and the pedicel, and by the 
glandular serrations of the calyx. From P. yedoensis it can be distin- 
guished by its brighter flowers and by its glabrous calyces and leaves. 
A more complete comparison of the hybrid and the parental species is 
given in Table I. 

TABLE L COMPARISON OF X PRUNUS JUDDII WITH ITS PARENTS 



branches spreading to branches spreading branches upright 

horizontal 

leaves greenish when leaves dull brownish leaves bright bron 

unfolding green when unfolding green when unfold 

flowers in 2-6-flowered flowers in 2-4- flowered flowers in sessile c 



pedicels glabrous 



petals broadly oblong, petals oblong, flushed petals narrowly oblonp 

nearly white with rose pink typically bright rose p: 

calyx tube urceolate- calyx tube sub-urceolate, calyx tube cylindric- 

cylindric, finely pubescent glabrous campanulate, glabrous 

calyx-lobes strongly calyx-lobes weakly and calyx-lobes entire 

glandular-serrate irregularly glandular 



Since it has not been found wild, Prunus yedoensis has itself been 
thought to be a hybrid between Prunus Lannesiana and Prunus sub- 
hirtella} The fact that it comes true from seed^ makes this hypothesis 
less likely, though such true-breeding hybrids are not unknown in the 
genus Prunus.^ 

X Viburnum Juddii Rehder, hybr. nov. = V. Carlesii Hemsl. 9 X 
bitchiuense Mak. $ . 

A Viburno Carlesii praecipue differt foliis supra minus dense pilosis, 
petiolis paullo brevioribus, corymbo laxiore magis multifloro, corolla 
extus magis roseo suffusa graciliore, limbo paullo minore, lobis angusti- 
oribus filamentis quam antherae longioribus; A V. bitchiuensi differt 
praecipue foliis supra magis pilosis, petiolis paullo longioribus, 5-7 mm. 
longis, corymbo 6-7 cm. diam. magis florifero, corollis majoribus tubo 



^Wilson, E. H. The Cherries of Japan, p. 19. Ca 
2Russell, Paul. The Oriental Flowering Cherries, i 
«C. D. Darlington in Jour. Genet. 19: 213-256 (1 



1935] ANDERSON AND REHDER, NEW HYBRIDS 361 

9-10 mm. longo, limbo 14-15 mm. diam., lobis paullo latioribus circiter 
S mm. latis, staminibus medio tubo affixis antheris faucem attingentibus. 

Growing in the Arnold Arboretum under no. 447-20; type specimens 
collected May 14, 1929, May 9, 1930, May 14, 1931 and May 14, 1935. 

This hybrid is in almost all characters intermediate between the parent 
species which are closely related and very similar, the chief difference 
being in the stamens which in V. bitchiuense are inserted in the lower 
fourth or third of the corolla-tube with the filaments about twice as long 
as the anthers and the tips of the latter 1.5-2 mm. below the mouth of 
the corolla-tube, while in V. Carlesii the stamens are inserted above the 
middle with the filaments as long or shorter than the anthers which reach 
the mouth of the corolla-tube. Table II shows the chief characters by 
which the hybrid may be distinguished from the parent. 

TABLE II. COMPARISON OF VIBURNUM JUDDII WITH ITS PARENTS 



or elliptic, sparingly or elliptic, furcate- rather densely furcate- 
furcate-pilose above, pilose above, bright pilose and grayish green 
slightly lustrous above green, not rugose when young, not rugose 



escence loose, rays 7-12 mm. loose, rays about l.S pact, rays S-l 

long, slender cm. long, slender long, stout 

Corolla pink outside, tube 7-8 pink outside, tube 9-10 corolla faintly 



mm. broad 5 mm. broad mm. across, lobes 5-6 

Filaments inserted in the lower inserted about or slight- inserted above to near 
third of the corolla ly beloW the middle, the middle as long or 
tube, about twice as about 1-^^ as long as slightly longer than 

Anthers tips 1.5-2 mm. below tips reaching the mouth tips reaching the mouth 

As shown by the table above, the hybrid holds the middle between the 
two parent species except in the size of the inflorescence and the length 
of the corolla-tube, in which it exceeds both parents. In its general 
appearance it resembles more V. bitchiuense on account of its looser 
habit and the looser inflorescence and more brightly pink flowers. As 
an ornamental plant it is superior to either parent. 

Viburnum Juddii was raised in 1920 by Mr. William H. Judd of the 
Arnold Arboretum staff from seed of V. Carlesii. The largest plant of 
the hybrid is now 2 m. tall and flowered for the first time in 1929. Like 
the parent species it has stood the severe cold of the last two winters 



without injury to its flower-buds. 



A. R. 



362 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

X Syringa diversifolia Rehder, hybr. nov. = Syringa pinnatijolia 
Hemsl. 9 X ohlata Lindl. var. Giraldii (Lemoine) Rehd. 6 . 

A Syringa pinnatijolia differt praecipue foliis partim integris, partim 
basi pinnatifidis pinnis 1-4 ovato-oblongis vel anguste ovatis 2-i cm. 
longis acuminatis basi anguste decurrentibus leviter ciliolatis ceterum 
glabris, foliolo terminali ovato-oblongo sensim acuminate 3.5-5 cm. 
longo, foliis integris ovato-oblongis, 3.5-5 cm. longis, 1.4-2.2 cm. latis, 
basi rotundatis, inferioribus interdum fere ovatis, inflorescentia ad 12 
cm. longa et laxiore, corolla coeruleo-lilacina, tubo circiter 8 mm. longo, 
limbo circ. 1 cm. diam., lobis apice leviter cucuUatis, antheris faucem 
paene attingentibus; a S. oblata var. Giraldii recedit praecipue foliis 
partim pinnatifidis minoribus et angustioribus, gemma terminali 
ramorum evoluta et ramum foliiferum emittente, inflorescentia minore, 
corollae tubo breviore et limbo angustiore, antheris faucem attigentibus. 

Growing in the Arnold Arboretum under no. 148-30; type specimens 
collected May 17 and 21, 1935, preserved in the herbarium. 

A comparison of the chief characters by which the hybrid differs from 
its parents are given in Table III. 

TABLE IH. COMPARISON OF SYRINGA DIVERSIFOLIA 



11 leaf- partly entire and partly always 
ig, leaf- pinnatifid with 3-5 ovate 4- 
broad, leaflets 4-6 cm. long, and ?>-t 



4-7 cm. long, usually 
branches, sessile 


to 11 cm. long, 
one pair at e 
branches, sessile 


white, usually tinged 


whitish or bluisl 



lobes oval, slightly i 



This hybrid was raised in 1929 from seed collected in 1929 from 5. 
pinnatijolia Hemsl. the flowers of which were apparently pollinated by a 
plant of 5. oblata var. Giraldii (Lemoine) Rehd. not very far from 
S. pinnatijolia. In the same year. Dr. K. Sax fertilized S. pinnatijolia 



1935] ANDERSON AND REHDER, NEW HYBRIDS 363 

with pollen of S. oblata var. Giraldii and plants were raised from this 
pollination; these plants have not yet flowered, but in their vegetative 
characters agree with the plant described above. The pollen of S. pin- 
natijoUa is defective, at least that of our plant, and self-pollinated flow- 
ers produce no seeds. The hybrid is clearly intermediate between these 
two species, readily distinguished from both species by the partly pinna- 
tifid and partly entire leaves. In the partly entire and partly pinnatifid 
leaves the hybrid resembles 5. persica L. var. laciniata, which can be 
distinguished by the broadly decurrent often obtusish lobes of the leaves 
and by the narrower and generally smaller entire leaves, by the smaller 
panicles usually in several to many pairs along the branches, the absence 
of the terminal leaf-bud, and by the anthers not reaching the mouth. 



JOURNAL OF THE 



ll-ith plate 154 

Five species of Hypodermataceae have been reported on conifers in 
Japan by Shirai and Hara (1927). Only one species, Lophodermium 
pinastri (Schrad. ex Fr.) Chev., was listed as occurring on pines. The 
present paper describes a new Japanese species of Hypodermataceae of 
unusual interest which was encountered during a hasty examination of 
the Hypodermataceae in the Mycological and Pathological Herbarium 
of the United States Department of Agriculture in Washington, D. C. 
Grateful acknowledgment is made to Dr. C. L. Shear and Mr. John A. 
Stevenson for the privileges extended to the writer. 

Hypodermella Hiratsukae, sp. nov. 

Hysterotheciis in uno ordine epiphyllis oblongis ellipticisque atroni- 
tidis, 0.54-1.30 X 0.26-Q.34 mm., longitudinali incisura aperientibus; 
hysterotheciis in transversali sectione in medio subcuticularibus sed ad 
marginem subepidermalibus, 0.16-0.22 mm. profundis; basilari plecten- 
chymate achroo 20-35 |j crasso; tegente strato atri pseudoparenchy- 
matis 28-34 \\ crasso; hymenio 100-110 p crasso. Ascis latis fusi- 
formibusque octosporis 87-102 X 18-24 p. Paraphysibus 100-110 X 
1 [J simplicibus filiformibus muco involutis. Ascosporis clavatis fusi- 
formibusque ad basin attenuatis hyalinis 35-56 X 3.5-5.0 p, muco 8 \x 
crasso involutis. 

In foliis Pini pumilae Regel, in monte Kuro-dake, provinciae Ishikari 
Japoniae, mense Augusto, 1927, Naohide Hiratsuka legit. 

Hysterothecia in a more or less continuous row, epiphyllous, oblong 
and elliptical, shining black, 0.54-1.30 X 0.26-0.34 mm., opening by a 
longitudinal fissure; hysterothecia in cross section subcuticular in the 
middle but subepidermal at margins, 0.16-0.22 mm. deep (closed); 
basal layer colorless, plectenchymatous, 20-35 ^j thick; covering layer 
of dark pseudoparenchyma 28-34 p thick; hymenium 100-110 p thick. 
Asci broad, somewhat fusiform, truncate to rounded at maturity at tip, 
8-spored, 87-102 X 18-24 p. Paraphyses 100-110 X 1 M, simple, 

'CO.VTRIBUTION FROM THE CrVPTOGAMIC LaBORATORU-S AND THE FaRLOW HERBARIUM, 



Jour. Arnold Arb. Vol. X^ 




1935] DARKER, HYPODERMELLA HIRATSUKAE 365 

filiform, surrounded by a delicate gelatinous sheath. Ascospores clavate 
fusiform, tapering towards the base, hyaline, 36-56 X 3.5-5.0 |j, sur- 
rounded by a conspicuous gelatinous sheath up to 8 p thick. 

On needles of Pinus pumila Regel, Mt. Kuro-dake, Province Ishikari, 
Japan, August 12, 1927, collected by Naokide Hiratsuka. 

Hypodermella Hiratsukae is of special interest because of certain 
morphological resemblances to Hypodermella Laricis v. Tub., the type 
species of the genus. As previously pointed out by the writer in 1932, 
the species of Hypodermella fall readily into four easily recognized 
groups named after the first described species in each as follows: (a) H. 
Laricis group, (b) H. anupla group, (c) H. nervisequia group, and (d) H. 
sulcigena group. Of nineteen species recognized in the genus, H. Hirat- 
sukae approaches most closely H. Laricis, hitherto the only species in 
that group. The linear arrangement of the hysterothecia, the broad 
clavate asci and ascospores and the absence of a slit band along which 
the hysterothecium ruptures are common to both species. The position 
of the fruiting body of H. Laricis in the host tissue is difficult to deter- 
mine even in microtome sections but is considered to be subcuticular by 
the writer. In the new species the hysterothecia are subcuticular in the 
centre and subepidermal at the margins as in Lophodermium pinastri. 
A prominently developed slit band, however, is characteristic of L. 
pinastri. Pycnidia with spores of the microconidial or spermatial type 
which are conspicuous and abundantly formed in the life cycle of H. 
Laricis are unknown in the case of H. Hiratsukae although in the 
material examined there are present certain small blister-like areas 
between the hysterothecia which may represent the remains of pycnidia. 



(3rd edition, pp. 448.) 
Darker. Grant Docks (1932). The Hvpodei 
(Contr. Arnold Arb. 1: 1-131). 



Pimis pumila Regel. 
Portion of needle with hysterothecia (X 17). 
Ascus and paraphyses (X 500). 
Ascospores (X 500). 
Hysterothecium in cross-sectional view (X 270 approx.). 



JOURNAL 

OF THE 

ARNOLD ARBORETUM 

^E XVI. OCTOBER, 1935 Num 

THE HOSTS, LIFE HISTORY AND CONTROL OF 
GYMNOSPORANGIUM CLAVIPES C. AND P. 

Ivan H. Crowell 

With plates 155-160 

TABLE OF CONTENTS 



P., Their Taxoi 



J Hosts of Gy 



The Juniperus 
AND P., The 



■ OF THE Diseases Caused by Gym?; 



1. On Pomaceous Hosts. 

a. Morphological Symptomatology 

b. Histological Symptomatology . . 

2. On Juniperus Hosts. 

a. Morphological Symptomatology 

b. Histological Symptomatology . . 



. ON Juniperus Hosts 396 






JOURNAL OF THE ARNOLD ARBORETUM 



I. INTRODUCTION 

Gymnosporangium CLAViPES C. and P., commonly referred to as the 
quince rust fungus, was one of the earliest species of the genus to be 
defined in North America. Schweinitz (1832) described it from the 
aecial phase occurring on Crataegus sp. as Caeomu (Peridermium) 
germinale. Later, Cooke and Peck (1871) described the telial phase of 
a rust on Juniperus virginiana L. as Podisoma (Gymnosporangium) 
davipes. It was not until 1886, however, that Thaxter (1887) demon- 
strated by means of controlled cultures that these were but two phases 
of the same organism. Throughout the literature both specific names 
have been used. Some authors have followed Kern (1911) in calling 
the species G. germinale (Schw.) Kern. But Sydow (1915), Arthur 
(1934) and other mycologists adhere to the International Rules of 
Nomenclature and so designate the species G. davipes C. & P. 

The prevalence and destructiveness of the diseases caused by G. 
davipes on ornamental pomaceous hosts and on many orchard varieties 
of apples, as well as on red cedars, have occasioned numerous inquiries 
regarding the pathogenicity and control of this rust. The information 
pertainable to these matters has been so meagre that comprehensive 
studies on the causal organism and the diseases produced by it were 
begun four years ago. The results obtained are presented in this paper. 
Certain phases of the investigations not yet completed are being 
continued. 

The main lines of my investigations are as follows: 

1. A determination by means of cultures and field observations of the 
pomaceous hosts of G. davipes together with a discussion of their taxo- 
nomic position and geographic range. 

2. Similar determinations and discussions of the Juniperus hosts of 



G. davipes. 




3. Thesymptor 


natology of the diseases caused by G. davipes. 


4. A thorough e 


xamination of the life history of G. davipes on poma- 



eous and Juniperus hosts. 

5. The practicability of fungicidal and 
leasures of the diseases caused by G. davipes. 



1935] CROWELL, GYMNOSPORANGIUM CLAVIPES 369 

II. THE POMACEOUS HOSTS OF GYMNOSPORANGIUM 

CLAVIPES C. AND P., THEIR TAXONOMIC POSITIONS 

AND THEIR GEOGRAPHIC RANGE 

Contributions to our knowledge of the pomaceous hosts of G. clavipes 
as determined by artificial cultures are due chiefly to the work of Thax- 
ter (1887), Arthur (1910, 1912), Thomas and Mills (1929) and Miller 
( 1 93 2 ) . Numerous other investigators, by their field observations, have 
added several species to the list of pomaceous hosts. The total number 
previously reported is about thirty-six species in seven genera distributed 
as follows: Amelanchier (8), Aronia (3), Chaenomeles (1), Crataegus 
i20),Cydonia (l),Malus {2),Pyrus (1). 

In my own studies on the pomaceous hosts, 701 species and varieties 
in 15 genera were inoculated, following essentially the same procedure 
described in a previous article (1934). These genera and the number 
of species of each inoculated are — Amelanchier (18), Amelosorbus 
(1), Aronia (4), Chaenomeles (2), CrataegomespUus (1), Crataegus 
(588), Cydonia (1), Malus (44), Photinia (1), Pyrus (19), Sorbaronia 
( 1 ) , Sorbopyrus ( 1 ) , and Sorbus (17). One species of Comptonia and 
two of Myrica were also inoculated. From the results obtained the 
inoculated plants were placed in two groups. Those plants which de- 
veloped no evidence of infection were classed as immune; while those 
plants on which infection was evident were classed as susceptible. The 
number of hosts determined by artificial inoculations was augmented by 
field observations in the Arnold Arboretum, on private estates about 
Boston and from the reports of former investigators. 

The results showed that more than 480 species and varieties of poma- 
ceous plants (including 48 varieties of orchard apples) scattered among 
eleven genera are susceptible. These hosts are presented in table 1. 



TABLE I. POMACEOUS HOSTS OF GYMNOSPORANGIUM CLAVIPES 
C. AND P.i 

Amelanchier alnijolia Nutt., A. Bartramiana Roem. {A. oligocarpa 
[Michx.] Roem.), A. Bartramiana X laevis, A. Bartramiana X oblongi- 
jolia, A. canadensis Med., A. canadensis nana, A. erect a Blanch., A. 
florida Lindl, A. intermedia Spach., A. laevis Wieg., A. laevis X humilis, 
A. oblongijolia (Torr. and Gray) Roem., A. sanguinea DC, A. spicata 
K. Koch, A. stalonifera Wieg. 

Amelosorbus Jackii Rehd. 



For the taxonomy 


of the genus Crataegus, Palmt 


;r (192S) was used. Mr. Palmer 




; hosts of G. clavipes in the ge 




unpublished cata 


logue of Crataegi. For the 


taxonomy of the other genera, 



Rehder's Manual ( 



370 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

Aronia arbutijolia (L.) Ell., A. jioribunda Spach {A. arbutijolia atro- 
pttrpurea [Britt.] B. L. Robinson), A. melanocarpa Ell., A. melanocarpa 
elata Rehder, A. monstrosa Zabel. 

Ckaenomeles japonica Lindl., C. lagenaria Koidz., C. lagenaria marmo- 
rata, C. lagenaria foliis rubris, C. lagenaria sanguinea semiplena. 

Crataegomespilus grandiflora Bean. 

Anomalae: C. affinis Sarg., C. asperijolia Sarg., C. Brockwayae 
Sarg., C. Coleae Sarg., C. cydopkylla Sarg., C. Dunbari Sarg., C. Eggle- 
stonii Sarg., C. e; 'a^a Sarg., C. honesta Sarg., C. /rfeae Sarg., C. /w/>ro- 
;;wa Sarg., C. misella Sarg., C. pinguis Sarg., C. />Mfa^fl Sarg., C. repul- 
sans Sarg., C. Saundersiana Sarg., C. scabrida Sarg., C. shirleyensis 
Sarg., C. Mricwc Sarg. 

Coccineae: C. accUvis Sarg., C. arcuata Ashe, C. assurgens Sarg., 
C. aw/^cfl Sarg., C. cae^a Ashe, C. chippewaensis Sarg., C. confinis Sarg., 
C. conspecta Sarg., C. cristata Ashe, C. delecta Sarg., C. densiflora Sarg., 
C. elongata Sarg., C. fluviatUis Sarg., C. /re^a^/^ Sarg., C. /////m Sarg., C. 
Holmesiana Ashe, C. Holmesiana tardipes Sarg., C. ^ew^a Ashe, C. 
Macounii Sarg., C. miranda Sarg., C. neolondinensis Sarg., C. pedicellata 
Sarg., C. perrara Sarg., C. /'o/z7c Sarg., C. Pringlei Sarg., C. /JMrc Sarg., 
C. sejuncta Sarg., C ^er/a/c Sarg., C. Thayeri Sarg., C. i;fz'irfa Sarg. 

Crus-galli: C. arborea Beadle, C arduennae Sarg.-, C. armata 
Sarg., C. attenuata Sarg., C. barrettiana Sarg., C. Bartramiana Sarg., 
C. 6eZ/Jca Sarg., C. calopkylla Sarg., C. Cc«6>'i Sarg., C. certwrna Sarg., 
C. crus-galli L., C. crus-galli arbutijolia Hort. ex Nicholson, C. crus-galli 
exigua Sarg., C crus-galli pyrac ant hi folia Ait., C. crus-galli rubens Sarg., 
C. criis-galli splendens Ait., C. effulgens Sarg., C. erecfc Sarg., C. Far- 
we/Zw Sarg., C. Fontanesiana (Spach) Steudel, C. geneseensis Sarg., C. 
insignis Sarg., C. Lavallei Herincq, C. lawrencensis Sarg., C. leptophylla 
Sarg., C. livoniana Sarg., C. mccra Beadle, C. pachyphylla Sarg., C. 
Palmeri Sarg., C. parciflora Sarg., C. Pennypackeri Sarg., C. peoriensis 
Sarg., C persimilis Sarg., C. phlebodia Sarg., C. Reverckonii Sarg., C. 
ma/w Sarg., C. robusta Sarg., C. rubrifolia Sarg., C. rwrf^i Sarg., C. 
triumphalis Sarg. 

Dilatatae: C. coccinoides Ashe, C. dilatata Sarg. 

Douglasianae: C Colorado^ Ashe, C. columbiana Howell, C. 
Douglasii Lindl., C. Douglasii f. 6a(/ia Sarg., C. Douglasii Suksdorfii 
Sarg., C. erytkropoda Ashe, C. P/>en Britt., C. rimdaris Nutt. apud Torr. 
& Gray. 



1935] CROWELL, GYMNOSPORANGIUM CLAVIPES 371 

Flavae: C. colonica Beadle, C. dispar Beadle, C. flava Ait., C. 
ignava Beadle. 

Intricatae: C. Delosii Sarg., C. flavida Sarg., C. nwdesta Sarg., 
C. nemoralis Sarg., C. neobushii Sarg., C. scabra Sarg., C straminea 
Beadle. 

Macracanthae: C admiranda Sarg., C. aquilanaris Sarg., C 
crrfwa Sarg., C. baccata Sarg., C. Beckiana Sarg., C bristolensis Sarg., 
C. calpodendron (Ehrh.) Med., C Calvinii Sarg., C. chadfordiana Sarg., 
C. corporea Sarg., C Deweyana Sarg., C rfiz;?Wa Sarg., C. Emersoniana 
Sarg., C. jerentaria Sarg., C. /er^a Sarg., C. jertilis Sarg., C finitima 
Sarg., C. flagrans Sarg., C. jidgens Sarg., C. gemmosa Sarg., C. glabrata 
Sarg., C. Handyae Sarg., C. hystricina Ashe, C. laxijiora Sarg., C. wccra- 
cantha Lodd., C. membranacea Sarg., C microsperma Sarg., C. missouri- 
ensis Ashe, C. neofluvialis Ashe, C. occidentalis Britt., C. ogdenburgensis 
Sarg., C. peramoena Sarg., C. prunijolia (Marsh.) Pers., C. radiosa Sarg., 
C. rhombijolia Sarg., C. Robinsonii Sarg., C. Searsii Sarg., C. spatiosa 
Sarg., C. spinidosa Sarg., C. strtictilis Ashe, C. succulenta Schrader, C. 
: L., C. truculenta Sarg., C. zje^e^a Sarg. 

: C. Phaenopyrum (L. f.) Med., C. spathulata Michx. 

MoLLEs: C. anomala Sarg., C. arnoldiana Sarg., C champlainensis 
Sarg., C. contortifolia Sarg., C. rf/gwa Sarg., C. rf/^/^ewa Ashe, C. £/Z- 
wangeriana Sarg., C. exclusa Sarg., C Fidleriana Sarg., C. induta Sarg., 
C. mi>?5a Sarg., C. lanuginosa Sarg., C. lasiantha Sarg., C. /aw^a Sarg., 
C. /iwam Sarg., C. mo^fo (Torr. & Gray) Scheele, C. wm^cwj Sarg., C. 
pennsylvanka Ashe, C. peregrina Sarg., C. Robesoniana Sarg., C. .yera 
Sarg., C. submollis Sarg.,' C. Tatnalliana Sarg., C. urbica Sarg. 

Oxyacanthae: C. altaica Lange, C. fdemalis Lange, C. Heldreichii 
Boiss., C. intermedia, C. Maximowiczii Schneider, C. monogyna Jacq., 
C. monogyna alba-plena Schneider, C monogyna inermis Rehd., C. 
monogyna laciniata Loud., C. monogyna pteridijolia Rehd., C m^wo- 
g>'nc spectabilis, C. monogyna stricta Loud., C. monogyna versicolor, 
C. Oxyacantha L., C. Oxyacantha Gireoudi Bean, C. Oxyacantha alba 
West., C. Oxyacantha rubra Schneider, C. pinnatifida Bunge, C. 50^1?)/- 
/o/ia Lange, C. Wilsonii Sarg. 

Pruinosae: C. arcana Beadle, C. aridtda Sarg., C. as per a Sarg., C. 
austera Sarg., C. 6ea/a Sarg., C bellula Sarg., C. brachypoda Sarg., 
C. bracteata Sarg., C. cestrica Sarg., C cognata Sarg., C. comparata 
Sarg., C. confragosa Sarg., C. delawarensis Sarg., C. disjuncta Sarg., 
C. dissona Sarg., C. divisijolia Sarg., C. Ferrissii Ashe, C. jormosa Sarg., 



372 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

C. jusca Sarg., C. georgian<i Sarg., C. glareosa Ashe, C. horridula Sarg., 
C. incisa Sarg., C. Jesupii Sarg., C. Kellermanii Sarg., C. latijrons Sarg., 
C. levis Sarg., C. macrocalyx Sarg., C. «wwero5a Sarg., C. />fl^rMw Sarg., 
C. pequotorum Sarg., C. perampla Sarg., C. perjucunda Sarg., C. />Mc- 
delphica Sarg., C. platycarpa Sarg., C. pruinosa (Wendl.) K. Koch, C. 
pulchra Sarg., C. relicta Sarg., C. remota Sarg., C. rubicundula Sarg., 
C ^/ccfl Sarg., C. tribulosa Sarg. 

Prunifoliae: C. decorata Sarg. 

Punctatae: C. amnkola Beadle, C. angustata Sarg., C bar bar a 
Sarg., C. Brownietta Sarg., C. cahescens Sarg., C. ce/^a Sarg., C. cow- 
/»a<:^a Sarg., C. desueta Sarg., C. florijera Sarg., C. glabrijolia Sarg., 
C. incaedua Sarg., C Lettermanii Sarg., C. notabilis Sarg., C. pausiaca 
Ashe, C; porrecta Ashe, C. praestans Sarg., C. pratensis Sarg., C. /)mmc- 
^a^c Jacq., C. punctata aurea Ait., C. punctata canescens Britt., C. Wgen5 
Beadle, C. suborbiculata Sarg., C. succincta Sarg., C. ^ewcjc Ashe, C. 
verruculosa Sarg. 

RoTUNDiFOLiAE : C. BicknellU Eggl., C. Blanchardii Sarg., C. 
Brainerdii Sarg., C. Brunetiana Sarg., C. chrysocarpa Ashe, C. coccinata 
Sarg., C. cupilijera Sarg., C. Dodgei Ashe, C. inaudita Sarg., C. Jonesae 
Sarg., C. Kennedy! Sarg., C. kingstonensis Sarg., C. maligna Sarg., C. 
Margaretta Ashe, C. Margaretta xanthocarpa Sarg., C. Maribella Sarg., 
C. Oakesiana Eggl., C. praecoqua Sarg., C. Proctoriana Sarg., C. propria 
Sarg., C. rotundata Sarg., C. rotundifalia Moench, C. rotundijoUa pubera 
Sarg., C. rotundijoUa f. rubcscens Sarg., C. varians Sarg., C. Webstcri 
Sarg. 

Silvicolae: C. allecta Sarg., C Barryana Sarg., C. blairensis Sarg., 
C. compta Sarg., C. delectata Sarg., C. diffusa Sarg., C. dissona Sarg., 
C. e^erfl Sarg., C. /i/i>e5 Ashe, C. /o/ifl^<z Sarg., C. Fre/2« Sarg., C /ra- 
CM«(^a Beadle, C. jVerc/a Sarg., C. Livtngstoniana Sarg., C. luxuriosa 
Sarg., C. medioxima Sarg., C. opulens Sarg., C. promissa Sarg., C. /rowa 
Ashe, C. ra(/mc Sarg., C. recordabilis Sarg., C. ruricola Sarg., C. stoloni- 
jera Sarg., C. strigosa Sarg., C. tortuosa Sarg., C. xanthocarpa Sarg. 

Tenuifoliae: C. acuminata Sarg., C. acutiloba Sarg., C. alnorum 
Sarg., C. apiomorpha Sarg., C. ascendens Sarg., C. asperata Sarg., C. 
6d/a Sarg., C. benigna Sarg., C. blandita Sarg., C. Boothiana Sarg., 
C. colorata Sarg., C. conjerta Sarg., C. crudelis Sarg., C. cyanophylla 
Sarg., C. Z)amd Sarg., C. delucida Sarg., C. demissa Sarg., C. E^/^ont 
Sarg., C. £gan« Ashe, C. /irwa Sarg., C. flabellata (Bosc.) K. Koch, 



1935] CROWELL, GYMNOSPORANGIUM CLAVIPES 373 

C. florea Sarg., C. Forbesae Sarg., C. jiLCosa Sarg., C. genialis Sarg., C. 
glaucophylla Sarg.. C. gracilipes Sarg., C. Grw^m Ashe, C. Habereri 
Sarg., C. Hadleyana Sarg., C. heidelburgensis Sarg., C. i'n^a/^Va Sarg., 
C. leptopoda Sarg., C. lucorum Sarg., C. luminosa Sarg., C. macrosperma 
Ashe, C. warc^'rfa Ashe, C. mc^Mra Sarg., C. werfxa Sarg., C. wenVa Sarg., 
C. miniata Ashe, C. worf/cc Sarg., C. monstrata Sarg., C. Napaea Sarg., 
C. «e5cm Sarg., C. o^w^a Ashe, C. Paddockeae Sarg., C. Paineana Sarg., 
C. palUdula Sarg., C. parviflora Sarg., C. pastorum Sarg., C. paucispina 
Sarg., C. pentandra Sarg., C. perlevis Ashe, C. populnea Ashe, C. />wwi/<z 
Sarg., C. re/rz/5a Ashe, C. roanensis Ashe, C. rubicunda Sarg., C. rw^Jro- 
c<zr«ea Sarg., C. rM^/>e5 Ashe, C. sarniensis Sarg., C. Serena Sarg., C. 5e;c- 
/^7w Sarg., C. S/at^mi Sarg., C. Streeterae Sarg., C. ^mcuw Sarg., C. 
/ae^r/cfl Sarg., C. /arrfc Sarg., C. ?e«e//a Sarg., C. tenera Sarg., C. ^enw/- 
/o6a Sarg., C. trachyphylla Sarg., C. viridimontana Sarg., C. u/^a^c Ashe. 

Triflorae: C conjimgens Sarg. 

Uniflorae: C. Mwiyiora Moench. 

ViRiDEs: C. abbreviata Sarg., C. ^/owa?a Sarg., C. penita Beadle, 
C. velutina Sarg., C. viridis L., C. vulsa Beadle. 

Cydonia oblonga Mill. 

Malus angustijolia Michx., M. jioribunda Sieb., M. ioensis plena 
Rehd., M. pumila Mill., M. spectabilis Borkh., M. sylvestris Mill. 

Photinia villosa DC. 

Pyrwj cowmwwf^ L., P. 5me««-i Lindl. 

5or6w^ americana Marsh, 5. dumosa Greene. 



A complete enumeration of all species and varieties of inoculated 
plants on which the results were negative is as follows : 

Amelanchier amabilis Wieg., A. asiatica Endl., A. grandiflora Rehd., 
A. humilis Wieg., A. humilis X sanguinea, A. ovalis Med., A. sera Ashe. 

Malus arnoldiana Sarg., M. asiatica Nakai, M. atrosanguinea Schneid., 
M. baccata Borkh., M. baccata costata Hort., M. baccata gracilis Rehd., 
M. baccata /cc^k Rehd., M. baccata mandshurica Schneid., M. baccata 
microcarpa Regel., M. baccata pendula Hort., M. brevipes Rehd., M. 
fiexilis Hort., M. florentina Schneid., M. Halliana Parkmanii 
Rehd., M. Halliana spontanea Rehd., M. Hartwigii Koehne, M. hona- 
nensis Rehd., M. hupehensis Rehd., M. kansuensis Schneid., M. spec. 
(Pyrus Malus laurifolia Gibbs), M. spec. (Pyrus Lemoinei Hort.), M. 
magdeburgensis Schoch, M. micromalus Mak., M. orthocarpa Lavall., 
M. Prattii Schneid., M. pumila Niedzwetzkyana Schneid., M. purpurea 
Rehd., M. purpurea aldenhamensis Rehd., M. purpurea Eleyi Rehd., 



374 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

M. robusta perskijolia Rehd., M. Sargenti Rehd., M. Scheideckeri 
Zabel, M. Scheideckeri "Excellenz Thiel," M. sikkimensis Koehne, M. 
spectabilis Riversii Nash, M. sublobata Rehd., M. toringoides Hughes, 
M. trilobata Schneid., M. Tschonoskii Schneid., M. yunnanensis 
Schneid., M. yunnanensis Veitchii Rehd., M. zumi Rehd., M. zumi calo- 
carpa Rehd. 

Pyrus amygdalijormis Vill., P. Balansae Decne., P. betulijolia Bge., 
P. Bretschneideri Rehd., P. dentictdata Hort. Angl. ex Dum.-Cours., 
P. elaeagrijolia Pall., P. Korshinskyi Litv., P. Lindleyi Rehd., P. longipes 
Coss. & Dur., P. Mickauxii Bosc, P. nivalis Jacq., P. paskia Buch.-Ham., 
P. phaeocarpa Rehd., P. salvijolia DC, P. serotina Rehd., P. serrulata 
Rehd., P. syriaca Boiss., P. ussuriensis Maxim. 

Sorbaronia alpina super aria, S. spec. 

Sorbopyrus auricularis bulbijormis Schneid. 

Sorbus Aria Crantz, S. amoldiana Rehd., 5. Aucuparia L., 5. com- 
tnixta Hedl., 5. discolor Hedl., S. hybrida L., S. intermedia Pers., S. 
japonica calocarpa Rehd., 5. Matsumurana Koehne, 5. Meini- 
chii Hedl., 5. pohuashanensis Hedl., 5. rotundijolia Hedl., S. subpinnata 
Hedl., 5. thuringiaca Fritsch (5. decurrens Hedl.). 

li/a/ic Ait., Myrica 

In addition to the pomaceous hosts just reported, several varieties of 
orchard apples have been found by former investigators to be susceptible. 
A compilation of these is presented in table 2. Mills (1929) gives the 
following account of the occurrence of G. clavipcs on orchard apples in 
New York: "Counts in 14 orchards in 4 counties showed fruit infection 
on Delicious (3 counts) 60 per cent; Fameuse (1 count) 21 per cent; 
Hubbardston (1 count) 28 per cent; Mcintosh (15 counts) 18 per cent 
average; Winesap (2 counts) 74 per cent; Yellow Transparent (1 count) 
84 per cent. Specimens were from 6 or 7 counties. Not found on foliage 
or twigs." This account and the fact that many other varieties of 
orchard apples are susceptible serve to stress the economic importance 
of G. clavipes to orchardists. In several orchards visited in Massachu- 
setts the disease caused by G. clavipes was found to be particularly abun- 
dant on the Delicious variety. As high as 90 per cent of the fruits were 
attacked. This disease, one of the most severe on the Delicious apple, 
is of much concern to orchardists because this variety is being grown in 
greater quantities to meet the steadily increasing demands for it on the 



CROWELL, GYMNOSPORANGIUM CLAVIPES 



Alexander 
Baldwin 


N. Y. 

Ind., Me., N. Y., N. S. 




Bechtel's Crab 
Bellflower 
Ben Davis 


Mass. 
Me. 

N. Y. 


Ind. 


Bishop 
Black Twig 


N.S. 

Va., W. Va. 




Cortland 
Crimson Beauty 
Delicious 


Me., N. Y. 

N.S. 

Ind., Me., Mass., N. Y., 

Tenn., Va., W. Va. 




Duchess 

Early Red Mcintosh 


Me., N. Y. 
Me. 




Fameuse 


N. Y., N. S. 




Family 

Gideon 

Golden Delicious 

Gravenstein 


N.S. 
Ind. 
Me. 
Me., N. Y., N. S. 




Grimes 


Ind.. Tenn. 




Hubbardston 


N. Y. 




Jonathan 


Ind., Me., N. Y. 


Tenn. 


King David 


Ind. 




Maiden Blush 
Mcintosh 


Me., N. Y., N. S. 


Ind. 


Northern Spy 
Northwestern Greening 


N. Y. 
Ind., N. Y. 




Red Delicious 
Red Winesap 
R. I. Greening 
Ribston 
Rome 


Me.. N. Y., Tenn. 
Tenn. 

N. Y., N. S. 

N.S. 

Md., N. Y., Tenn., Vt.. Va. 


Ind. 


Roxbury 
Russett 


N. Y., N. S. 





Stark 

Starking 
Stayman 



Ind., N. Y., Tenn., Va., W. Va. 



JOURNAL OF THE ARNOLD ARBORETUM 



TABLE n. {Continued) 



Stayman Winesap 


Tenn. 


Sweet Winesap 


N. Y. 


Tolman 


Me., N. Y. 


Tompkin's King 


N. S. 


Twenty Ounce 


Me., Mass., N. Y. 


Wagener 


Me., N. S. 


Wealthy 


Ind., Mass., Me., N. Y., Tenn. 


Winesap 


Ind., Me., N. Y., Tenn., Vt., 




Va., W. Va. 


Winter Banana 


Ind., Me., N. Y. 


Wolf River 


N. Y. 


Yellow Bellflower 


N. S. 


Yellow Transparent 


Md., N. Y.. Tenn., N. S. 



The species and genera of pomaceous hosts recorded above show no 
simple relationship or correlation by which one can formulate a rule to 
encompass them and set them apart from related non-susceptible plants. 
Nevertheless, they possess several distinctive features chiefly with respect 
to their taxonomy and their geographic range. Pomaceous hosts of 
G. clavipes are found in eleven genera. So far as I am able to learn no 
other species of Gymnosporangium is known to have hosts in so large a 
range of genera. All of the host genera of G. clavipes are closely related, 
however, and confined within the family Rosaceae. The native geo- 
graphic range of these hosts is more extensive than for those of any 
other species of Gymnosporangium known to me. Pomaceous hosts of 
G. clavipes are found throughout the whole of the temperate portion of 
the northern hemisphere. 

One of the most outstanding of the introduced foreign host species to 
become parasitised is Cydonia oblonga, the quince. This plant is native 
over the greater portion of Asia but has been introduced into North 
America over a portion only of the range of the rust. In the genus 
Chaenomeles, also native to Asia, two of the three species listed in 
Rehder's Manual are attacked. Several varieties and forms of the Japa- 
nese quince {Ghaenomelcs spp.) are also parasitised. One European and 
one Asiatic species of the genus Pyrus are attacked, while all other species 
(Eurasian) have so far proved to be immune. Some native and some 
foreign species of Mains as well as many orchard varieties of apples are 



1935] 



CROWELL, GYMNOSPORANGIUM CLAVIPES 



hosts to G. clavipes. In the genus Sorbus the two North American 
species are susceptible while all of the foreign ones inoculated remained 
immune. The hybrid genus Amelosorbus has but one species susceptible 
to G. clavipes. All of the species of the genus Aronia, native to North 
America, are hosts. With the exception of three or four species the genus 
Amelanchier , as represented in North America, is attacked by G. clavipes. 
A single species of the genus Photinia, the Eurasian species P. villosa, is 
liable to infection. 



I I ^ f 



JJ_ 



t 1 i 



111 



Crataegus Susceptible 

The susceptibility of each group of the genus Crataegus is presented as 
a graph in figure 1 . The data for this graph were obtained by determin- 
ing the percentage relationship between the number of species and vari- 
eties tested and the number that proved to be susceptible. While little 
significance can be attached to the results obtained from groups with a 
small number of species and varietit^s, nevertheless, reliable deductions 
can be made from those with large numbers as well as from the genus as 
a whole. 

Susceptible species were found in all groups except the Bracteatae. 
Unfortunately this group was represented in the Arnold Arboretum by 
but a single tree. In these investigations, 79 per cent of the species that 
produced fruit when the tests were made proved to be susceptible. Many 
species and varieties have not yet been tested, either because they were 
not available or because they did not produce fruits in the years of my 



378 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

investigations. Therefore, the host list is far from complete. All of the 
foreign species and varieties available for testing (placed in the group 
Oxyacanthae) proved to be susceptible. Of the native Crataegi, the 
percentage of species susceptible in the several groups varied widely. In 
some groups, for example, the Anomalae and Douglasianae, all of the 
species and varieties tested were susceptible while in other groups, for 
example, the Molles, Virides and Flavae, the percentage of susceptible 
members was less. It is interesting to note that the relative susceptibility 
of species and varieties and of the groups of the genus is not the same 
for hosts of G. clavipes as MacLachlan (1935) found for hosts of G. 
globosum, though hosts for both of these fungi were in most cases deter- 
mined from the same individual trees. 

III. PRELIMINARY STUDIES ON THE PERIOD OF 

SUSCEPTIBILITY OF POMACEOUS HOSTS 

OF GYMNOSPORANGIUM CLAVIPES 

It has been shown by several investigators that the leaves of poma- 
ceous hosts of certain species of Gymno sporangium are susceptible during 
a limited period in their youth only. This interim is known as the period 
of susceptibility. Thomas (1933) showed that leaves of Crataegus spp. 
were susceptible in their youth only to the attack of G. clavipes. My pre- 
liminary investigations, with respect to this phenomenon on fruits of 
pomaceous hosts of G. clavipes, also indicate that their period of suscep- 
tibility is brief. In this connection inoculations were made every two 
days for a period of four weeks on different fruit clusters of two species 
of Crataegus, C. tomentosa and C jertilis. Experimentation began when 
the flower buds of each species were opening. On C. tomentosa the 
ovaries, calyces, petals, pedicels, peduncles and twigs were attacked 
before the flowers opened. All but the ovaries and young fruit became 
immune within ten days. The fruits became decreasingly susceptible 
and by the time the petals fell they could no longer be infected. On C. 
jertilis, the young fruits only became infected. The flowers were immune 
up to the time the buds were opening and the petals began to expand ; 
they then entered a brief period of susceptibility extending until the time 
when the petals began to fall, after which they again became immune. 

A measure of susceptibility of the hosts of G. clavipes is here suggested. 
Those hosts that are susceptible for a longer period may be considered 
to be more susceptible than those that can be infected for a shorter 
period. No consistent difference was observed with respect to the 
abundance of aecia produced on susceptible as compared with resistant 

The possibility of any relationship between immunity and fertilization 
was not investigated. 



1935] CROWELL, GYMNOSPORANGIUM CLAVIPES 379 

IV. THE JUNIPERUS HOSTS OF GYMNOSPORANGIUM 

CLAVIPES C. AND P., THEIR TAXONOMIC POSITIONS 

AND THEIR GEOGRAPHIC RANGE 

Cultural studies for the purpose of determining the Juniperus hosts of 
G. clavipes have been very limited indeed. Arthur (1912) cultured G. 
davipes on /. communis depressa Pursh (/. sibirica Burgsd.) and Dodge 
(1918) obtained heavy infection from sowing aeciospores of G. clavipes 
on /. virginiana L. Several other investigators have substantiated these 
reports and added other hosts from their field observations. In my own 
work I have repeatedly cultured this rust on the red cedar, /. virginiana 
L. The technique for this work followed essentially the same procedure 
as for culture work on pomaceous plants. A complete enumeration (to 
1934) of the Juniperus hosts of G. clavipes together with data on their 
taxonomic position, as given by Rehder (1927), and their geographic 
range are presented in table 3. I have examined the rust on all of these 
species and varieties. 

TABLE III. JUNIPERUS HOSTS OF G. CLAVIPES 

T,.nin.r„. ho.t. Taxonomic Geographic 



J. horizontalis Moench Sabina " " 

J. Sabina L. " Europe 

J. scopulorum Sarg. " Western North Amer. 

J. virginiana L. " Eastern North Amer. 

In 1933 each individual cedar in the collections at the Arnold Arbore- 
tum of Harvard University was examined for infection and the species 
and varieties were recorded in immune and susceptible groups. The 
species and varieties of Juniperus on which no infection was observed 
are as follows: 

Juniperus chinensis L., /. chinensis globosa Hornibrook, /. ckinensis 
japonica Lav., /. ckinensis rmis Gord,, /. ckinensis pendula Franch., /. 
chinensis Pfitzeriana Spaeth, /. ckinensis plumosa Hornibrook, /. cki- 
nensis plumosa aurea Hornibrook, /. ckinensis pyramidalis Beiss., /. 
chinensis Sargentii Rehd., /. ckinensis Water eri Hort., /. communis 
Ashjordii Hort., /. communis aurea Nichols., /. communis aureo-spica 
Rehd., /. communis compressa Carr., /. communis cracovica Hort., /. 
r oblongo-pendtda Sudw., J . communis oblonga Loud., J. com- 



380 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

munis pyramidalis Hort., /. communis suecica Ait., /. conjerta Pari., 
/. jormssana Hayata, /. horizontalis alpina Rehd., /. korizontalis Doug- 
lasii Rehd., /. horizontalis glomerata Rehd., /. horizontalis plumosa 
Rehd., /. horizontalis variegata Hort., /. procumbens Sieb., /. rigida Sieb. 
& Zucc, /. Sabina cupressijolia Ait., /. Sabina pyramidalis Hort., J. 
Sabina tamariscijolia Ait., /. Sabina variegata Carr., /. scopulorum hori- 
zontalis D. Hill, /. scopulorum viridifolia Hort., /. squamata Buch.- 
Ham., /. squamata Fargesii Rehd. & Wils., /. squamata Meyeri Rehd., 
/. squamata Wilsonii Rehd., /. virginiana aurea Hort., /. virginiana 
Burkii, J. virginiana Canaertii Senecl., /. virginiana Chamberlaynii Carr., 
/. virginiana cinerascens Hort., /. virginiana elegantissima Hochst., J. vir- 
giniana fastigiata Hort., /. virginiana filijera D. Hill, /. virginiana glauca 
Carr., /. virginiana globosa Beiss., J. virginiana Hillii Hort., /. virginiana 
Kosteri Beiss., /, virginiana pendula Carr., /. virginiana polymorpka 
Hort., /. virginiana pyramidalis Carr., /. virginiana pyramidalis glauca 
Hort., /. virginiana plunwsa Rehd., /. virginiana reptans Beiss., /. 
virginiana Schottii Gord., /. virginiana tripartita R. Smith and /. vir- 
giniana venusta Rehd. 

Usually the Juniperus hosts of species of Gymnosporangium are con- 
fined to a single section of the genus. Those of G. clavipes are excep- 
tional in that they are classified in two sections of the genus, Sabina and 
Oxycedrus. Geographically the Juniperus hosts of G. clavipes are found 
throughout the greater portion of the temperate region of the northern 
hemisphere, an unusually wide distribution for telial hosts of any one 
species of Gymnosporangium. 



1. On Pomaceous Ho.sts 
(A) Morphological symptomatology 

Morphological symptoms of disease caused by G. clavipes were first 
described from infected fruits on pomaceous hosts. Schweinitz (1832) 
gave a brief description of the gross morphological symptoms of the 
disease on the fruit of Crataegus sp. Farlow (1880) stated that G. 
clavipes (Roestelia aurantiaca) was "by far the most beautiful species of 
the genus which we have, at once attracting the px)pular eye by its bril- 
liant orange or almost cinnabar colored spores and shining white 
peridium. It is generally found on young fruit, though it is occasionally 
found on the stems and petioles, but I do not recollect having seen aecidia 
on the leaves. — One sometimes sees a quince two inches in diameter 



1935] CROWELL, GYMNOSPORANGIUM CLAVIPES 381 

more than half covered by the bright orange aecidia and occasionally 
small apples are affected in a similar way. Roestelia aurantiaca is gen- 
erally found in midsummer. I have, however, seen it on C. crus-galli as 
late as October." 

Weimer (1917) stated that "the veins alone (of quince leaves) are 
attacked and often become swollen to double their normal size." The 
swellings, he noted, caused the leaves to curl but the infected areas were 
not discolored. No aecia were found on the quince leaves. Adams 
( 1921 ) reported that branches and buds of hawthorns were very severely 
injured by this rust. He stated that "the aecia on the branches always 
precede the appearance of aecia on the fruit of hawthorns." Thomas 
(1933) reported that symptoms on apple foliage appeared 10 to 18 days 
after inoculating and that on apple fruits the symptoms were predomi- 
nately necrotic or hypoplastic. On artificially inoculated leaves of 
Crataegus he obtained numerous spermogonia "but aecia were produced 
only sparingly along the larger veins." 

In the present study, data for the symptomatology of the disease 
caused by the aecial phase of G. clavipes were obtained from observations 
made on more than 400 pomaceous hosts. Data on the relationship 
between elapsed time and progressive stages of development of symptoms 
and signs were recorded from plants inoculated at various stages of 
development of the flowers, twigs and fruits. Comparisons of the symp- 
toms and signs resulting from artificial inoculations made possible a more 
thorough understanding of the phenomena that occur in nature. 

Gymnosporangium clavipes attacks primarily the fruits, less fre- 
quently the twigs and buds and rarely the leaves of its pomaceous hosts. 
The earliest observable symptom of disease on ovaries and young fruits 
is a pale yellowish green discoloration. This symptom was seen on cer- 
tain species even before the petals fell. Occasionally the petals become 
infected and when they do they usually remain attached for the greater 
part of the season. In my experience, fruits are susceptible when in early 
stages of development only. After the petals drop the fruits of most 
species are no longer subject to infection. On the average, from 6 to 10 
days after inoculating the diseased fruits begin to show evidence of 
infection by slightly pale swellings. From 4 to 5 days later the hyper- 
trophied zone becomes dotted in its central portion with numerous, tiny, 
deep-reddish points — the developing spermogonia of the fungus. Within 
1 to 5 more days, the first formed spermogonia begin to exude a pale-red, 
sweetish liquid. During further development the diseased area con- 
tinues to increase in all dimensions and finally involves but a portion of, 
or, in many cases, the whole fruit. Occasionally, the infection spreads 
to the pedicels and even extends into the peduncles and twigs. Spermo- 



382 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

gonia are matured progressively over a large portion of the diseased 
tissue, and oozing of the young spermogonia continues for several days. 
The older spermogonia die and turn black. 

Several irregularities or anomalies have been observed in the sympto- 
matology of the disease during the life span of the spermogonia. Very 
frequently the infected tissues do not become hypertrophied, but quite 
cease development. During this time the adjacent tissues continue to 
expand, resulting in invaginated areas. This is particularly common in 
orchard apples where infection occurs most frequently at the blossom 
end. In many instances the spermogonia are few in number and do not 
reach full maturity. Tissues with this type of infection are usually green 
and firm. Fruits of hawthorns, shadbushes and chokeberries are found 
in which the swelling and discoloration involve the whole fruit, but in 
such cases no or few spermogonia are produced. Many of the fructi- 
fications exhibit various stages of abortion. Symptoms of this type were 
obtained from inoculation tests on Crataegus spp. and Amelanchier 
oblongijolia toward the close of the period of susceptibility of the frurts. 
Of course, in some instances it is possible that more than one kind of 
parasite is involved and that as a result the normal course of the Gymno- 
sporangium disease is altered. 

In 20 to 40 days after inoculating the second fructifications of the 
fungus, the aecia, begin to make their appearance within the diseased 
area. They are usually produced peripheral to, or to one side of, the 
spermogonia; but they are often found among them. Aecia, in pro- 
gressive stages of development, are easily observed during their early 
appearance as is shown in plate 155, fig. 1. Often no aecia penetrate the 
surface of the infected fruits, yet internal ones are frequently formed. 
This is particularly true of orchard apples but has also been observed in 
many other hosts. Fruits that become infected late in the period of 
susceptibility are commonly observed to exhibit this phenomenon. Fail- 
ure to fully develop aecia may be physiological — possibly a type of 
hypersensitivity — or it may be due to the development of a cuticle so 
tough as to prevent aecia from breaking through. 

Many fruits were observed in which the lesions occupied by G. davipes 
were browned or blackened and quite rotted. From such decayed areas 
imperfect fungi were repeatedly obtained by culture. It seems, therefore, 
that these are the real cause of the discoloration and decay noted. Some 
have ascribed such phenomena to G. davipes, but it seems erroneously 
so. The fact that relatively few areas infected by G. davipes become 
decayed strengthens this conclusion. A photograph of a rust-infected 
area that was parasitised by an imperfect fungus is in plate 155, fig. 2. 



1935] CROWELL, GYMNOSPORANGIUM CLAVIPES 383 

An anomolous symptom was frequently observed on fruits of Cratae- 
gus monogyna infected with G. clavipes. The whole of the fruits and 
the pedicels were infected, but from the blossom ends of these fruits 
numerous petals and abortive structures resembling stamens developed. 
Here again, more than one parasitic organism may have been active in 
causing these symptoms. 

Twigs, including thorns, infected with G. clavipes are commonly found 
on certain species of hosts (plate 156). In my experience twigs of the 
current season only have become infected. The early symptoms of 
disease on twigs are pale, yellowish green, elongated areas occurring on 
the young bark. The infection spreads rapidly, however, usually girdling 
the stem and extending up to 3 inches longitudinally. The infected por- 
tion of the twig becomes hypertrophied. Frequently a fusiform swelling 
results, but irregularly swollen and cankered twigs are also common. 
Rarely, however, rotund galls are formed on the twigs. Thorns are also 
subject to infection and manifest similar symptoms. Spermogonia on 
infected twigs usually do not reach maturity until 9 to 12 days after 
inoculation. They follow the same course of development as on fruits. 
Aecia of the rust are produced among the spermogonia as well as outside 
the area occupied by them. Very often they are sparsely produced, but, 
on the other hand, twigs are sometimes found in which the aecia are very 
abundant. The iirst aecia reach maturity on the twigs and thorns in 
30 to 40 days; others are produced progressively, as was described in the 
case of infected fruits, for a period of about a week. 

In the longitudinal advancement of the fungus in a twig, it frequently 
encounters a terminal or lateral bud, which in turn usually becomes 
infected. Subsequently, the buds are forced to develop beyond the 
resting stage normal for the current season. Similar phenomena were 
observed on buds of ornamental apples infected with G. Juniperi-vir- 
ginianae (Crowell, 1934). Forced growth of buds caused by G. clavipes 
has been observed on a large number of hosts. The early stages of in- 
fection, evidenced by a yellow discoloration and swellings of the buds, 
are not observable until late in June. The abnormal development of 
stunted twigs and leaves results. The photographs in plate 156 show 
some of these symptoms in forced buds. Spermogonia in all stages of 
development are produced along the deformed petioles and veins of the 
leaves. Rarely, however, are aecia produced on the forced buds. On 
C. Phaenopyrum several twigs were found in which the infection devel- 
oped systematically, as shown in plate 156, fig. 5. It is possible that 
infection occurred on these twigs while they were in the stage of rapid 
elongation. That infection was confined largely to these malformations 
and did not extend into the main twig substantiates this supposition. 



384 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

Many twigs infected with G. davipes in the spring of 1934 were ex- 
amined in the spring of 1935 to determine whether or not the fungus over- 
wintered in its aecial hosts. No instance of overwintering was found. 
Thomas (1933), however, reported having observed the overwintering 
of this rust in pomaceous hosts. Dodge (1918) reported that the aecial 
phase of G. biseptatum Ellis {G. Botryapites [Schw.j Kern) and of G. 
jraternum Kern (G. trans jormans [Ellis] Kern) were observed by him 
to be perennial. Tubeuf (1906, 1907) also noted this phenomenon on 
pears infected with G. Sabinae (Dicks) Wint. 

The leaves of the pomaceous hosts of G. clavipes are rarely infected. 
When such occurs the lesions are always small and few spermogonia 
develop on them. The spermogonia on leaf lesions are late in appearing, 
and on many hosts they never reach the oozing stage. Spermogonia 
have been observed on leaves of Cydonia oblonga, Amelanchier spp., 
Craetegus spp., and Aronia spp. Amelanchier oblongijolia is the only 
host, however, in which I have seen aecia of G. clavipes on affected leaves. 

On its more susceptible pomaceous hosts G. clavipes is very destruc- 
tive. The diseased fruits become misshapen and discolored, and often 
fall prematurely. In the case of quince and apples the loss is primarily 
a commercial one and frequently very great. Indeed, growing of quinces 
was impossible in many sections of eastern North America due largely to 
the ravages of this rust. Many varieties of orchard apples are also very 
susceptible. Thomas and Mills ( 1930) report instances in which as high 
as 95 % of the fruits of the Delicious variety in New York were attacked. 
Ornamental plants, such as certain species of Crataegus, Amelanchier, 
etc., whose beauty and usefulness depend in a large measure upon an 
abundant production of colored fruits that persist long into the winter, 
sometime becomes worthless because so many of their fruits are spoiled 
by G. clavipes. Twigs killed beyond the infected portion are also un- 
sightly and they tend to materially deform the trees and shrubs because 
of sequent prolific sprout growth. Hawthorns have been seen in which 
so many twigs were killed by the rust that death resulted. 
( B ) Histological symptomatology 

Tissues of pomaceous hosts infected with G. clavipes are usually 
hypertrophied and the diseased portions of fruits and twigs are often 
greatly enlarged; infected parts of leaves, however, are changed but little. 
In the fruits chiefly the outer or cortical tissues are affected. They are 
often greatly enlarged and many of them are more or less filled with a 
deep-staining material while others, fully as hypertrophied. are often 
quite devoid of contents. Cells towards the center of the fruit, although 
they show no evident hypertrophy, contain much of the deeply staining 



1935] CROWELL, GYMNOSPORANGIUM CLAVIPES 385 

material observed in the cortical cells. Exceptions to the hypertrophy 
of the fruit cells as noted have been observed in the infections of many 
orchard varieties of apples. In such infections the earliest stages of the 
disease are manifested as an hypertrophy of a small number of cells. 
The disease soon ceases to be an enlargement; indeed, the infected region 
becomes inhibited in its development. The surrounding uninfected 
tissues, however, continue to expand in their development while the 
diseased area remains stunted, depressed and usually greenish. 

In twigs, mainly the cortical tissues are affected. The effects are simi- 
lar to those in fruits. Occasionally, however, and always in the greatly 
enlarged twigs, the phloem and xylem are hypertrophied. Many of their 
cells contain much material that stains intensely with haematoxylin. 

The mycelium of the aecial phase of G. clavipes is found in the cortical 
tissues of the fruits and twigs and in the palisade and mesophyll tissues 
of the leaves. Haustoria are abundantly formed in the cells of the fruits 
and twigs but are seldom seen in cells of the leaves. Frequently, haus- 
toria are found in close association with the nucleus of its host cell. 

The peridial cells and the aeciospores of G. clavipes are salient fea- 
tures for determination of the species. Gymnosporangium clavipes can 
also readily be identified from median longitudinal sections of the spermo- 
gonium. In my investigations of the spermogonia of the genus (to be 
reported in a separate article) three dimensions were averaged, namely, 
total width, total height and the depth to which the fructification is 
sunken in the host tissue. The measurements for G. clavipes were 
found to be 203 |j X 207 |j X 163 |j respectively. The spermogonia of 
G. clavipes are conspicuous because of their large size and their rotund 
form and they are almost completely sunken in the mesophyll. A 
photomicrograph of spermogonia of G. clavipes is shown in plate 156, 
figure 7. 

Aecia of G. clavipes are found most abundantly in the fruits and twigs 
and are seldom produced in the leaves. Aecia develop in the outer cortex 
of fruits and twigs and in the mesophyll of leaves. They are sunken to a 
depth of approximately one millimeter and are of greater diameter near 
the base than at the apex. Fresh aeciospores of G. clavipes vary in color 
from bittersweet-orange to flame-scarlet (Ridgway, 1912). They are 
irregularly rotund, verruculose and measure 32.2 p X 34.4 |j. Peridial 
cells are almost white in color; the inner wall is maked by coarse ridges 
forming an irregular mosaic pattern. The form of peridial cells varies 
widely, but on an average they measure 14.5 p in width X 55.6 ^i in 
length. They usually adhere in large numbers in water mounts. 



386 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

2. On JuNii'ERus Hosts 
(A) Morpholog^ical symptomatology 

Cooke and Peck (1873) gave a description of the symptoms of G. 
clavipes on Juniperus virginiana; they stated: "the younger branches 
are slightly swollen where attacked by this fungus and the bark is scaly." 
For many years, however, the fungus and the disease caused on its 
Juniperus hosts seem not to have been understood. Thaxter (1891) 
after an extensive series of cultural experiments clearly identified two 
species that were formerly confused with G. clavipes, namely G. clavipes 
proper and a new species, G. Nidus-avis. Kern ( 191 1 ) , in an account of 
the symptomatology of G. clavipes, stated that the telia were "caulicolus, 
appearing on slightly fusiform swellings, usually aggregated, roundish, 
one to four millimeters, often confulent, hemispheric — teliospores two- 
celled ellipsoidal 18 to 26 p X 35 to 51 p — pedicels carotiform. Dodge 
(1918) stated that small witches-brooms are sometimes formed on twigs 
of red cedars infected with G. clavipes. Dodge (1922) stated that 
needles as well as the main trunk are also infected, and that the bark 
over the infected portions of the trunk becomes much thickened and 
blackened. 

The pertinent observations on the symptomatology of G. clavipes re- 
viewed in the foregoing give a clear foundation for an appreciation of the 
disease caused by G. clavipes on its Juniperus hosts. My own findings 
and interpretations, while they add little to what is already known will, 
nevertheless, trace in sequence the development of the disease and its 
relative importance in the various organs attacked. 

On its Juniperus hosts, particularly /. virginiana L., the disease in- 
duced by G. clavipes is perhaps one of the most destructive caused by 
any species of the genus Gymnosporangium. The needles, twigs, 
branches and even the main trunk are attacked. Very frequently the 
disease occurs on the needles but infected needles are relatively incon- 
spicuous and are often overlooked. Usually but a single crop of telio- 
spores is produced on needles, after which they die. From the needles, 
however, the fungus often migrates to the twigs and it is on these that 
the disease is most frequently found. Twigs may also become infected 
directly. On twigs the disease appears as slightly fusiform swellings 
covered by a flaky, darkened bark. Usually by the end of four to six 
years most of the infected twigs die. Occasionally, however, the infected 
twigs survive for a longer period and the larger limbs are distorted and 
covered with a heavy, cracked and blackened bark. If the diseased por- 
tion of a branch is near the main trunk the latter is liable to infection by 
the fungus advancing along the cortex. Infections of G. clavipes on the 



1935] CROWELL, GYMNOSPORANGIUM CLAVIPES 387 

main trunk of red cedars are easily mistaken for one of the diseases caused 
by other species of the genus, because vertically elongated, often irregu- 
lar, blackened, heavy-barked bulges are typical for diseases caused by 
several species of Gymno sporangium. Fructifications are, in my ex- 
perience, necessary to identify the causal organism on trunk lesions. 

That the disease caused by G. davipes is very destructive to its telial 
host is evident since all organs attacked are killed usually in a few years. 
Trunk infections of many years standing are not uncommon, to be sure; 
but trees bearing such burdens show evident symptoms of poor health. 
Recently, I came upon a very striking demonstration of destruction of 
red cedars infected with G. davipes. Several years ago, an estate owner 
cleared a natural grove of cedars of all other trees and shrubs. Gymno- 
sporangium davipes was very abundant on the twigs, branches and main 
trunks of certain of these cedars. Following the unusual cold of the 
winter of 1933-34 every tree that was heavily infected with G. davipes 
died entirely or in large part, while all of the trees in this group that were 
not infected survived. An examination of the cedars in the surrounding 
uncleared lands revealed complete destruction of trees that were heavily 
infected with G. davipes. As the only variable seemed to be the relative 
abundance of infections the loss seemed clearly attributable to the infec- 
tion of G. davipes. 
(B) Histological symptomatology 

Dodge (1922) gave a very complete account of the histological symp- 
tomatology of G. davipes. A review of his paper is presented here. 
Dodge stated that "infection takes place on the proaxial side of the 
young leaf, or directly on the young stem at the base of the leai — after 
entering the leaf, the mycelium invades the region between the cuticle 
and the cellulose walls of the epidermis on the proaxial side. The effect 
of the fungus on this part of the cell wall is usually marked by consider- 
able swelling and the disorganized substances take the stains very 
readily — the fungus explores and feeds in the cuticularized layer, but 
it may go deeper and invade the palisade-mesophyll tissue. Haustoria 
are found in the epidermal cells, sometimes even in the guard cells of the 

Young stems are "susceptible to infection, either directly or through 
the invasion of the fungus by way of the leaf axils." The mycelium occu- 
pies a cancellate portion of the periphery of the young cortex. The 
mycelial "strands actually interlace, weaving in and out around the veins 
[leaf traces] of the leaves and forming a closed network system in the 
cortical region of the young stem." After the leaves are shed the fungus, 
closing the gaps, may be found in the cortex around the entire circum- 



388 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

ference of every section. In the main trunk, infections are confined to a 
portion only of the circumference. 

My findings with respect to the histological symptomatology agreed 
in detail with those of Dodge. My examination included the leaves and 
twigs of Juniperus virginiana as well as infected twigs of all other Juni- 
perus hosts. Certain additional remarks with respect to the fructifica- 
tions are also recorded. 

The telia of G. clavipes arise at irregular intervals from aggregated 
masses of the mycelium on the phellogen. The telia first appear early in 
April in Massachusetts. In youth they are a deep-reddish color, expand- 
ing upon gelatinization to regular pulvinate sori. After two or three 
gelatinizations, however, the sori change their deep red color to a yellow- 
ish red and their shape is very irregular. Five to six gelatinizations occur 
during the season, after which the telia drop from their host. 



The life history of Gymnosporangium clavipes is essentially the same 
as that of other species of the genus. It differs in certain details only 
from the life history of the more generally known species G. Juniperi- 
virginianae. A review of the extensive literature dealing with the life- 
history of G. clavipes shall be confined to the more pertinent reports of 
former investigators. 

Schweinitz (1832) described the aecial phase of a rust which he found 
occurring "rarissime in germinibus Rosae" as Caeotm (Peridermium) 
germinale. Kern (1911) reported the determination of the host of this 
rust to be an error for a species of Crataegus. Cooke and Peck (1873) 
gave an account of the telial phase of a rust which they called G. clavipes 
occurring on Juniperus virginiana L. These two rusts were considered 
to be distinct species until Thaxter (1887) showed from the results of 
controlled cultural experiments that the spermogonial and aecial stages 
on pomaceous hosts were in reality genetically connected with the telia 
stage on red cedars. Following this basic step many observations have 
been reported with respect to details in the life history of G. clavipes. 

In the development of the rust on pomaceous hosts Farlow ( 1886) and 
Thaxter (1887) found that 10 and 11 days respectively elapsed between 
the date of inoculating and the first appearance of spermogonia. Thomas 
( 1933) stated that the first symptoms of disease on "relatively resistant" 
apple foliage were observed 10-18 days after inoculating but that spermo- 
gonia were never formed on the lesions. In contrast he found that symp- 
toms appeared in 4 6 days and spermogonia in 13-15 days after 



1935] CROWELL, GYMNOSPORANGIUM CLAVIPES 389 

inoculating "susceptible" foliage of Crataegus sp. and further that aecia 
were formed sparingly along the larger veins only of the infected leaves. 
Thaxter (1887) determined from his cultures that approximately 30 
days were required for the maturation of aecia on young shoots of 
Amelanckier canadensis. Miller ( 1932 ) and other authors observed that 
spermogonia were frequently abortive in orchard apples and that aecia 
rarely matured in the fruits. 

These observations while they differ greatly when considered sepa- 
rately are nevertheless in harmony when reviewed in the light of more 
complete knowledge of the behavior of the rust. From my cultural ex- 
periments on hosts in several genera, the same variations as reported in 
the foregoing were observed. These variations seemed to be correlated 
with two phenomena, — first, the relative susceptibility of the host 
plants and second, the stage of development of the diseased parts at the 
time of inoculating. The rust developed more rapidly on the more 
susceptible hosts and more slowly on less susceptible ones. In fact, on 
very resistant hosts, as certain orchard apples or on resistant organs as, 
for example, the leaves of nearly all hosts, longer time was required for 
the maturation of the fructifications; not infrequently these reached an 
imperfectly developed stage only. 

The average time required for the maturation of spermogonia of G. 
clavipes on fruits and twigs was from 7 to 10 days. The beginning of 
spermogonial exudation was taken as the criterion of maturity. On indi- 
vidual lesions spermogonia continued to be produced for a period up to 
3 weeks. The production of exudate by any one spermogonium contin- 
ued up to about 7 days, after which the spermogonium became filled with 
long filaments, and soon died, usually turning black. Others in the 
lesion followed these developments. Spermogonia of G. clavipes were 
sub-epidermal in origin and position; they were reddish in color and 
among the largest of the genus. Details of average measurements of 
their size have been given on page 385. 

The aecial primordium of G. clavipes is located deep in the cortical 
tissues of fruits and twigs and in the mesophyll of leaves. It is typical 
for the genus as is also the mycelium and haustoria. From thirty to forty 
days after inoculating, aecia of G. clavipes reached maturity and pene- 
trated the epidermis of the host among or close by the spermogonia. 
Aecia were usually developed progressively over a period of a week or 
more. Upon their first appearance, about June 1 in Massachusetts, they 
had the form of short, blunt, white cylinders. One or two days later 
the white peridia were ruptured irregularly, usually with the loss of the 
cap cells, exposing and releasing the enclosed reddish aeciospores. Vari- 



390 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

ous stages of this phenomenon are shown in plate 1 5 5, fig. 1 . The aecium 
of G, clavipes is broader at about midway between the hymenium and 
surface of the host than at the surface of the host itself. It is evident, 
therefore, that a crowding of the aeciospores occurs at the zone of con- 
striction. Dodge (1924) associates these phenomena with a mechanism 
for the forcible discharge of aeciospores. 

The peridial cells of G. clavipes are broadly rectangular and measure 
14.5 M X 55.6 jj with extremes of 33.4 ^j to 83.5 p X 10.4 u to 23.4 |j. 
Their inner and thicker walls are ornamented with low interconnecting 
prominences forming irregular mosaic patterns. Peridial cells are of 
much diagnostic value as Fischer (1891) and Kern (1910) demon- 
strated. Those of G. clavipes may be easily identified by their markings, 
their size and their form. Peridial cells of G. clavipes usually remain 
flat in water mounts and adhere forming a large sheet of cells. 

Aeciospores of G. clavipes are exceptional for the genus in their re- 
markably intense color. By comparisons of fresh aeciospores the color 
was determined as varying from bittersweet-orange to flame-scarlet 
according to Ridgway's (1912) color standards. With increase of age 
of the mature spores, however, their color gradually changes to orange 
and yellows. In old herbarium material they are frequently almost 
colorless. In many of the better preserved specimens the reddish color 
still is conspicuous. Aeciospores of G. clavipes are among the largest of 
the genus. They measure 32.2 p X 34.4 |j with extremes of 28.4 \x to 
42.3 |j X 26.7 |j to 37.4 p. Their outer surfaces are ornamented with 
numerous, tiny, low papillae. 

The problem of germinating aeciospores of species of Gymnosporan- 
gium has been given a great deal of attention. Difficulty has been ex- 
perienced in germination of aeciospores of many species. Several 
investigators have shown that a period of rest at low temperature con- 
tributed greatly to the germinability of aeciospores of certain species of 
Gymnosporangium (Fukushi, 1925; Miller, 1932). Thomas and Mills 
(1929) reported moderate germination of aeciospores of G. clavipes 
stored for twelve weeks at 3° C. Thomas (1933) tested the germina- 
bility of aeciospores of G. clavipes that were precooled for various lengths 
of time at various temperatures as well as aeciospores that were not pre- 
cooled. The highest germination was obtained from spores that "were 
mounted at 18° C. without precooling." Thomas also demonstrated that 
aeciospores kept dry at 3° C. rapidly lost the property of germination. 
It was found, however, that a small number of aeciospores remained 
viable in aecia in fruits throughout the winter. It should also be re- 
marked that Professor J. C. Arthur observed internal aecia of G. clavipes 



1935] CROWELL, GYMNOSPORANGIUM CLAVIPES 

in fruits of orchard apples. Dr. Arthur stated in a letter to Dr. 
metz, who forwarded the material to him, that aeciospores remove^ 
internal sori germinated in the usual manner. 

In my studies of the germination of aeciospores of G. clavipes 
found that they germinated readily in a moist atmosphere at roor 



r C. and ] 



perature at any time during the summer and fall seasons. Careful studif 
of germination revealed that they germinated over essentially the sam 
range as did most species of the genus studied to date. Optimum cor 
ditions for germination in distilled water on glass slides were reached a 
about 15° C. The results of the tests are shown in figure 2 and i 
table 4. 



JOURNAL OF THE ARNOLD ARBORETUM [vol. 

TABLE IV. GERMINATION OF AECIOSPORES OF G. CLAVIPES 



0°C. 



Percentage c 



1000 



Apart from the irregularities in the aecial phase that were discussed 
under the heading of symptomatology, an unusual development was ob- 
served in the aecia on fruits of Crataegus sp. collected by Prof. J. H. Faull 
in Pennsylvania. Some of these aecia and aeciospores appeared normal 
in all respects. Certain of the aecia were internal with the hymenia 
oriented in various directions. All of these latter and many of the aecia 
that developed in a normal position were filled with irregularly produced 
aeciosfX)res and aeciospore chains. Cytologically either one or two 
nuclei were present in the cells. The cell contents were irregularly inter- 
spersed with vacuoles and deep-staining materials. Camera lucida draw- 
ings of two aeciospore chains are shown in plate 157, figure 2. 

Aeciospores of G. davipes are primarily wind borne. They are dis- 
tributed throughout the growing seasons and probably germinate and 
infect the alternate hosts shortly after inoculation. 

Previous to 1910, however, little investigative work had been con- 
ducted on the telial phase of G. davipes. Arthur (1912) sowed aecio- 
spores of G. davipes on the common juniper and reported successful cul- 
tures. Dodge (1918) traced the life cycle of G. davipes under controlled 
conditions from the telial phase on red cedars to the aecial phase on 
Crataegus Oxyacantha, thence back to the telial phase on red cedars. 
Dodge found that few tella were produced in the spring following inocu- 
lation but that many developed in the second spring. Inoculations made 
during the present investigations substantiated these findings. In addi- 
tion it was shown that inoculations made on July 1, August 10 and Octo- 
ber 3 all resulted in abundant infection, indicating that in nature aecio- 
spores are a menace throughout their entire pei.od of production. 

Infection of the telial host was first described by Dodge (1922). 



1935] CROWELL, GYMNOSPORANGIUM CLAVIPES 393 

Dodge stated that entrance of the germ tube was gained through the 
adaxial surface of the leaves and tender epidermis of the twigs. In the 
leaves of its Juniperus hosts the mycelium of G. clavipes is confined al- 
most exclusively to the epidermal cells. In twigs it is restricted to the 
phellogen cells — a most unusual limitation for the mycelium of a species 
of Gymnosporangium. Within the infected cells characteristic, binucle- 
ate, sac-like haustoria are formed. They are abundant, usually occurring 
singly but frequently in twos or threes. Telial sori of G. clavipes arise 
from masses of the mycelium. In leaves the mycelium usually dies after 
the production of one crop of spores, but in twigs the fungus is perennial 
for several years. After each successive crop of teliospores a new phello- 
gen layer is formed immediately beneath the sorus. The surrounding 
vegetative mycelium then grows over this new tissue. It is from the 
mycelium on the older phellogen that the telium for the ensuing spring 
is produced. In certain microscopic sections it is possible to observe 
progressively (a) the dead mycelium on partially sloughed-off phello- 
gens, (b) sori of the present season as well as (c) the primordia of sori 
for two seasons to come. The camera lucida drawing in plate 160, fig. 1 
was made from such a section. 

Telia of G. clavipes are produced on leaves and on the bark of various- 
sized branches and even the main trunk of red cedars. They were never 
observed on branches of red cedars more than one-half inch in 
diameter. Upon their early appearance telia of G. clavipes are aggre- 
gated, pulvinate in form, 2 to 5 mm. across and are distinctly bright 
reddish in color. During rains in the spring the telia swell to regular 
gelatinous forms as is shown in plate 159. After three or four gelatiniza- 
tions the telia lose their regular form and deep-red color, becoming 
shapeless yellowish-red masses. After 6-8 gelatinizations the telia drop 
from the infected parts. 

The development of teliospores of G. clavipes is essentially the same 
as Dodge (1918, 1922) reported for this and other species of the genus. 
Camera lucida drawings made during these investigations of teliospores 
in various stages of development are shown in plate 160. 

Teliospores of G. clavipes are at once distinguished by the swollen 
pedicels near their bases. No other species of the genus in eastern North 
America has this characteristic. Certain data in regard to teliospores 
of G. clavipes are of interest. Both one- and two-celled teliospores are 
produced. In a count of 1000 spores, 94.8 % were found to be two-celled 
while 5.2 % were one-celled. Two-celled teliospores have one germ pore 
in each cell. In the upper cell the germ pore is apical while in the basal 
cell the germ pore is located near the pedicel. In other respects one- 



394 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

celled teliospores resemble the two-celled ones in all but the septum. 
One-celled teliospores have the germ pore at the apex. In the telium of 
G. davipes both thick- and thin-walled spores were found. Thick-walled 
spores are more numerous and are always produced on the outer surface 
of the telium. Almost invariably one-celled teliospores are thick-walled. 
Thin-walled teliospores are located within or beneath the layer of thick- 
walled spores. Measurements were made of the lengths of the upper 
and basal cells, the total length and the width of teliospores of G. davipes. 
The upper cell measured 23.0 p with extremes of 16.5 to 33.0 p, the basal 
cell 21.2 ^J with extremes of 14.9 to 33.0 p, the total length was 44.5 \y 
with extremes of 33.0 to 57.8 ^i and the width 22.7 \x with extremes of 
16.5 to 33.0 |j. Single-celled spn^res measured 19.7 p in width by 33.4 \\ 
in length with extremes of 26.4 to 51.2 p X 14.9 to 29.7 \i. 

When teliospores of G. davipes first break through the cortical cover- 
ing layer of the host, they expand but little when wetted and the telio- 
spores do not germinate. From 1-3 weeks after their first appearance, 
however, the telia expands fully to a regular pulvinate form and the 
teliospores germinate in great abundance. 

Several workers have reported the results of their investigations on 
germination tests of teliospores of G. davipes. Weimer (1917) formu- 
lated a general curve of germination percentages obtained at various 
temperatures and stated that it applied to the germination of teliospores 
of G. davipes as well as teliospores of other species of Gymno sporangium. 
The extreme temperatures found by Weimer were 7° C. and 29° C. and 
the optimum temperature was between 22° C. and 25° C. Miller (1932) 
found the extreme temperatures of germination to be 4° C. and 2>2° C. 
and the optimum germination at 25° C. He also investigated the phe- 
nomenon of the maximum rate of germination and found that when telia 
of G. davipes were immersed in water for 25 minutes and removed to a 
moist atmosphere they discharged basidiospores in considerable num- 
bers within two hours. It was also found that when telia were mounted 
in moist cotton and kept at 25° C. an abundant germination of the telio- 
spores and the beginning of basidiospore formation occurred after an 
interval of eight hours. 

Thomas also investigated the time required for and the rate of germi- 
nation of basidiospores. He stated that basidiospores submerged in 
water at room temperature developed germ tubes equal in length to the 
spore in two hours and from four to six times their diameter in ten hours. 
Basidiospores were also mounted on a moistened leaf of Crataegus and 
held at 25° C. with the result that germ tubes reached a length of five to 
seven times their diameter in eight hours. 



1935] CROWELL, GYMNOSPORANGIUM CLAVIPES 395 

Farlow (1886) reported the production of secondary basidiospores 
from primary basidiospores of G. clavipes, a phenomenon that has fre- 
quently been reported for other species of the genus. 

Beyond observing the usual germination of the teliospores of G. 
clavipes, a process common to other sj^ecies, no further studies were made 
on the phenomenon. It was observed, however, that germination 
occurred in the field when the telia remained gelatinized for periods of 



\ 'illll '• 


^j-7"' 


f^ \ 




^' 1- 


\ '^(^ 


Ml 




1 'iiiin 




iM^^^Miriiiil 


1 t w^ «w 




li^MS 


:|fc 




S : : Ifm 1 li rr nr J m ^^S 


Jfflj^^ 




^ fir ' ' ' ' n Tul lllnmirn 


^y 




; ^HtlltlffllOI Jn 


pr 




^^ 


fe:; 



Figure 3. Geog^raphical Distribution in North America ol 
and its Hosts. The pomaceous hosts of G. clavipes shown as v 
Tlie Juniperus hosts of G. clavipes shown as horizontal Hnes. 
G. clavipes shown as dots. Note the extreme northern limit 
Alberta, Canada, and the extreme southern limit near Mexico C 



two hours or longer. Four to six gelatinizations were usual during the 
season and basidiospore dissemination occurred at most, if not at all, of 

It is essential to the production of infection that the pomaceous hosts 
are within their period of susceptibility at the time of inoculation. As 
the period is very short in many species, the failure for the coincidence of 
basidiospore dissemination within this period may be held responsible 
for the variable abundance of spermogonia and aecia on certain poma- 
ceous hosts in different years. Crowell (1935) reported unusual de- 



396 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

velopments of the aecial phase of this and other species of Gytnnospo- 
rangium in the very dry spring of 1935. In the spring of 1935, no rains 
sufficient to cause gelatinization of telia of G. clavipes occurred during 
the flowering period of most pomaceous hosts; most fruits becoming 
immune before inoculation took place. This resulted in a very limited 
number of infected fruits and the late appearance of the rust generally. 

Although the host relationships for the perpetuation of G. clavipes are 
found over the greater part of the temperate northern hemisphere, never- 
theless, so far as I am able to learn, this rust is not known outside of 
North America. Reported stations for this rust are most abundant in the 
eastern part of this continent as shown by dots on the outline map of 
North America in lig. 3. The rust has been collected, however, in widely 
separated stations outside the region of greatest concentration, an un- 
usual feature in the distribution of any species of this genus. 

Gymnosporangium clavipes has been reported from southern New- 
foundland, from all the provinces of Canada except Prince Edward 
Island, from all the states of the United States except Arizona. Cali- 
fornia, Idaho, Kansas, Minnesota, Nevada, New Mexico, North 
Dakota, Oregon, South Dakota and Washington. It is also reported 
from Mexico State in Mexico. No other species of Gymnosporangium is 
known to have a longitudinal range extending from central Mexico State 
to northern Alberta in Canada. An even greater range is possible on the 
basis of host distribution. It would be very interesting as well as of much 
practical value to determine the phenomena responsible for the distribu- 
tion of a rust within a portion of the territory occupied by both host 
groups. 

VII. CONTROL MEASURES APPLICABLE TO GYMNO- 
SPORANGIUM CLAVIPES ON POMACEOUS AND 
ON JUNIPERUS HOSTS 
Prophylactic measures to control G. clavipes have been largely an 
adaptation of those practiced for the control of the cedar-apple rust 
diseases caused by G. Juniperi-virginianae. The results obtained in 
control work have been essentially parallel for both of these diseases, 
namely, fungicidal control as practiced was found unsatisfactory, while 
eradication of Juniperus hosts gave excellent protection to pomaceous 

Halsted (1893) reported the destruction of quince, apple, hawthorn 
and shadbush fruit by G. clavipes in New Jersey and recommended 
remedial m?asures. Concerning the disease Halsted wrote: "an enemy 
is beyond the fence; therefore, go out and slay him with an axe." This 



1935] CROWELL, GYMNOSPORANGIUM CLAVIPES 397 

was the earliest record that I found regarding the pathenogenicity and 
control of this rust. Bailey (1894) also recommended the destruction 
of red cedars as a control measure. He gave evidence to show that 
spraying thoroughly (with Bordeaux ?) was of considerable value. 
Many other authors stated that eradication of red cedars offered the 
most satisfactory solution to the control of this rust. 

My own investigations on control measures applicable to G. clavipes 
were carried out simultaneously with those for the control of the cedar- 
apple rust fungus, G. Juniperi-virginianae (Crowell, 1934). Studies in 
this problem included an exploratory investigation of numerous fungi- 
cides with respect to their control value both on pomaceous hosts and on 
red cedars. The most promising of these fungicides were then tested on 
an extensive experimental scale on numerous trees under various weather 
conditions. The fungicides were applied to red cedars (a) to prevent 
germination of the teliospores, (b) to protect them from infection by 
aeciospores, and (c) to pomaceous hosts as a protection against basidio- 
spore infection. 

( A ) Fungicidal applications on red cedars to prevent germination 
of teliospores 

On the Lyman estate, Canton, Massachusetts, Gymno sporangium 
clavipes was in very great abundance and exploratory tests with several 
fungicides were made there. In table V are enumerated the sprays and 
dusts that were used in these tests. 



SPRAY AND DUST 


MATERiIlS U^D IN 


EXPLORATORY TESTS 


Bordeaux 
Linco colloidal su 
Lime-sulfur 
Soluble palustrex^ 


fur^ 


3:3:50,4:4:50,6:6:50 

y4%, y2%, \%, 2%. 3% 

1 :30, 1 :40, 1 :50, 1 :70 

1%, 2%, 3%, 4% 



Sunoco oil'^ 

50% Sunoco oil and 50% soluble palustrex 

50% Sunoco oil and 50% soluble palustrex B. 

80% Sunoco oil and 20% copper resinate 

80% Sunoco oil and 20% soluble palustrex 

Kolo base 

Kolo dust 

Pomo green 

Sulfur dust 
^Obtained from Linder and Co., 296 North Beacon St., Bostoi 
^Obtained from E. W. Coolidge, Jacksonville, Florida. 
■^Obtained from Sun Oil Co., Boston, Mass. 



398 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

All of these spray and dust materials were first applied to potted red 
cedars in the greenhouse in the spring. None of them caused burning of 
the young foliage. In the field the sprays were applied to twig lesions as 
follows: (1) before the telia had emerged, (2) just after the telia had 
emerged, and (3) after one, two or three gelatinizations of the telia, but 
always when the telia were dry. Telia to which the sprays were applied 
were brought into the laboratory for examination; smear slides were 
made of the spores and examination was completed shortly after their 
arrival. It was not the purpose of this examination to determine the 
relative value of each spray, rather the purpose was to determine which 
ones would prevent germination of the teliospores. The most satisfac- 
tory sprays were soluble palustrex B. at 4% and Linco colloidal sulfur 
at 1 % , 2 % and 3 % . In testing these sprays further it was found that 
this colloidal sulfur at 2 7o and 3 % was the most constant in its reactions. 
Colloidal sulfur was, therefore, chosen for the experimental work that 
followed. It should be added, however, that colloidal sulfur at 1 /o 
greatly reduced the amount of germination ; colloidal sulfur at 2 % com- 
pletely prevented germination of the teliospores of G. clavipes. Colloidal 
sulfur at 3%, therefore, was unnecessary. 

In April 1933, many telia of G. clavipes were sprayed upon their first 
appearance with colloidal sulfur at 1 % and 2 % . The effect of the spray 
was determined by gathering several sprayed telia three days after the 
application and also unsprayed telia at the same time for controls, 
thoroughly wetting and keeping them in a moist chamber over night. 
When germination was abundant a spore print resulted, but when no 
spore print was formed a smear slide was made and the teliospores ex- 
amined under the microscope. The unsprayed teliospores always germi- 
nated in abundance. Teliospores sprayed with colloidal sulfur at 1 % 
strength germinated to some extent but few basidiospores germinated. 
Telia sprayed with 2% colloidal sulfur showed no germination of the 

After the rain following each spray application, telia were again 
gathered and immediately tested. The controls germinated in abun- 
dance. The telia sprayed with colloidal sulfur at 1 % and 2 % germi- 
nated to a slight extent. Some change, therefore, was called for in order 
to control this small amount of germination. Certain substances were 
used to lower the surface tension of the spray material which might aid 
its penetration into the telium. Spreaders were tried but with no suc- 
cess. Calcium casienate, a combined spreader and sticker, was next 
used in a series of experiments. A new lot of telia was sprayed with 
colloidal sulfur at the strength of 2 7f' plus 2 pounds of calcium c 



1935] CROWELL, GYMNOSPORANGIUM CLAVIPES 399 

per hundred gallons of water. Telia tested in the laboratory showed a 
high percentage of germination for those that were unsprayed and no 
germination for those that were sprayed. In subsequent rains none of 
the teliospores were observed to have germinated nor did the telia at any 
time regain the property of gelatinizing fully, although they did gela- 
tinize to a slight extent. 

( B ) Fungficidal applications on red cedars as a protection against 
infection by aeciospores 

Fungicidal protection to red cedars against infection by aeciospores of 
G. clavipes has been demonstrated on potted red cedars in greenhouse 
trials. In these tests Linco colloidal sulfur at the strength of ^ % or 
6 pounds per 100 gallons of water only was used. Twenty-five red cedars 
twelve to eighteen inches high were sprayed with the fungicide. These 
and twenty-five unsprayed red cedars were thoroughly wetted with a 
strong stream from a hose the following day. They were then heavily 
inoculated with fresh aeciospores of G. clavipes and kept in a moist 
chamber for five days. In the spring of the second year after inoculating, 
each of the unsprayed plants produced an abundance of sori — a total of 
more than one thousand separate infections — while but a single infec- 
tion was found on the sprayed plants. 

In field experimentation, three applications (one each in July, August 
and September) were made to a group of red cedar trees near heavily 
infected hawthorns. Examination of the twigs the second spring after 
spraying showed a very marked reduction in the number of lesions pro- 
duced. It was estimated after comparing the amount of infection on 
unsprayed red cedars in the vicinity that about 75% control was 
obtained. 

The use of fungicidal means of protecting red cedars in practice 
should be guided, along with other considerations, by two important 
factors, namely, (1) the date of maturity of the aecia and (2) the 
duration of aeciospore production. 

The first of these will vary with the season and with the time of blos- 
soming of the host. In Massachusetts aecia reach maturity early in June. 
The second and the more variable of the two is the period of active aecio- 
spore production. The duration of this period is dependent upon the 
date of maturity of the fruit of the associated pomaceous hosts. As the 
fruit of Amelanchier ripen and drop about the middle of July in Massa- 
chusetts and aeciospore production ceases at this time, protective sprays, 
therefore, need be applied for but a brief period. Twig infections, if 
present on the shadbushes, should be carefully removed, because aecio- 
spore production will continue on them throughout the entire growing 



400 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

season. On the other hand fruits of Crataegi and most other pomaceous 
hosts do not reach maturity until late fall and aeciospore production 
usually continues during this period. 

(C) Fungicidal applications on pomaceous hosts as protection 
against infection by basidiospores 

Fungicidal protection of pomaceous hosts from infection by G. 
clavipes has been conducted through field experiments only. Individuals 
of Amelanchier oblongijolia were sprayed with Linco colloidal sulfur at 
the strength of >4 7o or 6 pounds per 100 gallons of water after each of 
three rains during the early development of the flowers in the spring. 
Spraying was begun just before the first rain after the flower clusters 
began to unfold; the last application was made when about three-fourths 
of the petals had fallen. The protection afforded these plants was ex- 
cellent. Counts of infected fruits showed that 98 /r of them remained 
free from infection, while 95 'a of the fruits on unsprayed plants nearby 
were infected. Certain other experiments with this same material on 
hawthorns and apples have not met with the same success. Excellent 
protection was afforded certain of the tested trees while practically none 
was obtained on others. It is believed that this irregularity can be over- 
come. It is not yet known just when infection occurs nor the limits of 
the period of susceptibility of many of the host species. A knowledge 
of these is necessary to satisfactory control. The value of thorough 
spraying cannot be over-emphasized. 

Gymnosporangium clavipes is largely a fruit parasite on its pomaceous 
hosts, and a difficulty arises with respect to spraying while the hosts are 
in flower. Sprays may be applied at any time except for a few days imme- 
diately after the unfolding of the petals. It is at this time that pollination 
occurs. The problem of controlling the disease caused by G. clavipes on 
orchard apples has not as yet been given attention. A project to deter- 
mine the limits of the period of susceptibility and modifications, if any, 
of the spray schedule now generally used in apple orchards is planned. 

In addition to protective spray applications for the control of G. 
clavipes, other means of attack may be employed. In ornamental plant- 
ings of red or common cedars judicious pruning of infected branches and 
twigs is very efTective. Not only will pruning remove the disease from 
infected trees but will afford a degree of protection to neighboring poma- 
ceous hosts as well. The work is best done in the spring when infected 
parts are clearly marked by the presence of telial sori. Small twigs may 
be cut off below the nearest uninfected shoot, but it seems advisable that 
larger branches be removed well below visible lesions ; in many instances 
this may be back to the main trunk. 



19351 CROWELL, GYMNOSPORANGIUM CLAVIPES 401 

The possibility of freeing infected trunks of red cedars from disease 
has been the subject of an, as yet, incomplete investigation. It will be 
recalled that the mycelium of G. clavipes is localized on the phellogen 
layer only of red cedars. Experimentation was conducted for the pur- 
pose of investigating the possibility of removal of the fungus by remov- 
ing the outer bark. For this purpose a coarse wood rasp was used, and 
the bark, over and for about an inch around the lesion, was scraped off 
and painted with shellac and later with an antiseptic tree dressing. The 
work was done in March and no telia developed in the spring nor did the 
trees show visible symptoms of injury the following growing season. 
Only one season has passed since the undertaking and thus far the 



Eradication of Juniperus and of pomaceous hosts, while often limited 
in practice, is*, nevertheless, a very effective adjunct or under certain 
conditions the most effective means of controlling this rust. The low 
growing types of junipers, such as /. communis and varieties and /. hori- 
zontalis, are frequently weed plants. In some localities in the Annapolis 
valley of Nova Scotia wild junipers have been eradicated for one-quarter 
to one-half mile around commercial orchards with the result of almost 
complete protection to the susceptible varieties of apples. From their 
observations in an apple orchard in Maine, Steinmetz and Hilborn 
(1934) state: "the approximate shortest distance between the infected 
junipers and infected apple trees is 3900 feet. The extreme distance 
between infected hosts is over 4500 feet." Various environmental fac- 
tors, however, may influence the distance that the contagion will travel 
as I have already discussed in another paper (Crowell, 1934 pp. 202-209) 
and these should be considered in plans for eradication. As an expedi- 
ent to the protection of either pomaceous or Juniperus hosts, all nearby 
hosts that can be dispensed with should be removed. Elaborate protec- 
tive measures may be largely upset by a single weed-host plant located 
near valued plantings. 

VIII. RECOMMENDATIONS 
The problem of controlling the diseases caused by G. clavipes on poma- 
ceous and on Juniperus hosts usually varies with the relative value of the 
infected plants and with the interest of the owner. The problem, how- 
ever merits wider attention. The moral obligation of consideration for 
a neighbor's earnest endeavors to improve a serious situation is too often 
thought of very lightly. Several practical control measures have been 
demonstrated. These are as follows: 



402 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

1. Selection oj immune or highly resistant species and varieties for 
planting. Attention should be given not only to avoiding hosts of G. 
clavipes but to avoiding hosts of other species of Gymno sporangium 
native in the vicinity. Host lists for all of the species of Gymnosporan- 
gium in eastern North America are now rapidly nearing completion. 
MacLachlan (1935) has shown that practically the whole of the genus 
Crataegus is more or less susceptible to attack by G. globosum. Many 
species and varieties of hawthorn are also susceptible to G. clavipes as 
shown in the present paper and to another widespread species common 
in this region, namely, G. clavariaejorme. (The presentation of results 
of investigations on this species is planned for an early publication.) 
Species and varieties of Crataegus, either native or foreign, cannot be 
planted in the vicinity of Juniperus hosts in eastern North America with 
expectation of their remaining entirely free from infection by one or 
more species of Gymnosporangium. In the genus Malus, Crowell 
(1934) has shown that all of the native species and varieties and one 
foreign species {M. sylvestris) can harbor and reproduce the cedar-apple 
rust fungus G. Juniperi-virginianae. The present paper and that of 
MacLachlan (1935) give but few additional species and varieties of 
Malus that are hosts to G. clavipes and G. globosum. All other Eurasian 
species and varieties do not harbor and reproduce the rusts native to 
this region. In the genus Amelanchier one species only, namely, A. ama- 
bilis, has proved to be highly resistant to all of the Gymnosporangia in 
this region. Other species and varieties are subject to infection by one 
or more species of Gymnosporangium. 

Few hosts for the native Gymnosporangia were found in the genera 
Pyrus and Sorbus while practically all of the species and varieties tested 
in the genera Aronia, Crataegomespilus, Cydonia and Photinia were 
susceptible. Information as to the hosts in other pomaceous genera is 
too meagre for general recommendations at this time. For sp>ecific host 
lists together with the relative data of various hosts attention is directed 
to the present and the following publications now available. For hosts 
of G. globosum, (MacLachlan 1935). For hosts of G. Juniperi-virgini- 
anae, on ornamental apples (Crowell 1934) ; on orchard apples (Crowell 
1935). 

Of the telial hosts, with the exception of Juniperus virginiana, J. hori- 
zontalis, J. scopulorum and several of their varieties among the native 
species and /. communis and /. Sabina and several of their varieties 
among the foreign species, all other species and varieties tested (see p. 
379) may be considered as desirable for plantings in this locality. 



1935] CROWELL, GYMNOSPORANGIUM CLAVIPES 403 

2. Planting a screen of tall non-susceptible trees about groups of 
alternate host plants. Groups of alternate host plants in close proximity 
may be effectively protected from infection by surrounding them with a 
screen of tall trees. Densely branching trees, as many of the conifers, 
are particularly effective. The corollary is also true. Groups of alter- 
nate host plants riiay be grown in close proximity if planted among taller 
non-susceptible trees. These phenomena have frequently been observed 
in nature. A fuller discussion is given on page 204 of an account of the 
cedar-apple rust disease by Crowell (1934). 

3. Eradication of pomaceous or Juniperus hosts. To be most effec- 
tive eradication of either pomaceous or Juniperus host plants should be 
complete over a radius of at least one-half mile. Even though eradication 
is carried out over this area, complete protection is not assured. Thus, 
the direction of winds during the time of spore production, continued 
humidity and the location of the source of inoculum are factors that may 
tend to offset or vary the results. A high degree of protection may be 
expected from eradication, however. If eradication cannot be complete, 
partial eradication of host plants will reduce the amount of inoculum and 
will therefore aid in controlling the rust. All wild or scrubby pomaceous 
as well as Juniperus hosts that can be dispensed with should be removed. 

4. Removal of infected parts of host plants. As infections in Juni- 
perus hosts are perennial for several years removal of diseased twigs and 
branches will contribute materially to control measures. Diseased twigs 
should be removed well below visible lesions. In the case of infected 
branches it may be necessary to remove them back to the main trunk. 
Similarly, on pomaceous hosts diseased fruits and twigs may be removed 
as an aid to control measures. Unless diseased parts are few in number 
and can be easily and thoroughly picked by hand, the undertaking is not 
recommended. 

5. Removal of the fungus from infected trunks of red cedars. In- 
complete experimentation has shown that infections on the main trunk of 
red cedars may be satisfactorily removed. This is done by rasping off 
the outer bark down to the living outer tissues, painting the wound with 
shellac and later with an antiseptic tree dressing. The practice is limited 
to trunk lesions and is not generally recommended. Removal of diseased 
parts or the disease from parts of Juniperus hosts is best accomplished 
when telia are present on infected areas to guide one in the work. 

6. Protective spray applications. Protective spray applications to 
pomaceous plants for protection against infection by basidiospores of G. 
clavipes are of value during the early stages of development of flowers 



404 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

and fruits. Linco colloidal sulfur was the only fungicide tested for this 
purpose. Applications at Yi % strength or 6 lbs. per 100 gallons of water 
plus a casein sticker are recommended. The first application should be 
made before the first expected rain as the cluster buds are breaking. Sub- 
sequent applications of the same strength should be made at 7-10 day 
intervals until most of the petals have dropped. More frequent applica- 
tions may be necessary if rain is unusually heavy or prolonged, or if 
flowers have expanded with unusual rapidity. 

Spray applications to Juniperus hosts should be confined to the period 
of aeciospore production on neighboring pomaceous hosts, and should be 
made after each two or three rains. In Massachusetts aeciospores are 
first liberated about June first. In the case of most species of Amelan- 
chier the fruits are ripened and dropped by the middle of July. There- 
fore, spray applications need not be continued longer than this time, pro- 
vided that Amelanckkr species are the only hosts in the vicinity. Care 
should be taken, however, to remove all infected twigs from these plants 
for aeciospore production on them continues throughout the entire grow- 
ing season. On most other hosts, aeciospores continue to be liberated 
during the entire growing season. Under such conditions spray applica- 
tions should be continued until the end of the growing season. 

The telia of G. clavipes are fully exposed one to two weeks before 
susceptible parts of pomaceous hosts are released from their buds. This 
affords an opportunity to attack the telia before it is possible for them 
to cause infection. Two spray applications of colloidal sulfur at the 
strength of one percent or 10-12 pounds per one hundred gallons of water 
are recommended. The applications should be made before buds of 
pomaceous hosts burst. Each application should be made just after a 
rain in which the telia are fully expanded and are beginning to dry. At 
this time they expose the greatest surface and absorb water with much 
avidity. It should not be expected that the fungus in the twigs and 
branches will be killed by this means, the telia of the present season only 
will be destroyed. 

The relative merits of each of the foregoing means of control for G. 
clavipes will vary with individual situations. Single or a combination of 
methods of control may be employed. Selective planting methods will 
doubtless give the most permanent results but are limited in their adapta- 
tion. Methods of eradication (3,4 and 5) may be employed where the 
plants are of such high value, or are few, or the rust sufficient sparse as 
to make hand labor practical. Spray applications are perhaps most wide 
in applicability since the practice of spraying is so general. 



19351 CROWELL, GYMNOSPORANGIUM CLAVIPES 405 

IX. SUMMARY 

1. Inoculations with Gymnosporangium davipes C. and P. and ex- 
aminations for infection were made on approximately seven hundred 
species and varieties in thirteen genera of pomaceous hosts. These gen- 
era were AmelancMer, Amelosorbus, Aronia, Chaenomeles, Crataego- 
mespilus, Crataegus, Cydonia, Malus, Photinia, Pyrus, Sorbaronia, 
Sorbopyrus and Sorbus. The genera Comptonia and Myrica as repre- 
sented in the Arnold Arboretum were also inoculated. The results 
show that hosts were distributed in eleven of these genera, namely, 
Amelanchier, Amelosorbus, Aronia, Chaenomeles, Crataegomespilus, 
Crataegus, Cydonia, Malus, Photinia, Pyrus and Sorbus. Although 
pomaceous hosts of G. clavipes are found over the entire temperate region 
of the northern hemisphere, the fungus is confined to North America. 

2. Investigations made on the period of susceptibility of flowers and 
fruits of certain pomaceous hosts showed that the flowers and fruits were 
susceptible after they were released from their buds for a brief period 
only. The more susceptible hosts were susceptible for a longer period 
than less susceptible hosts. 

3. Inoculations and examinations for infection on the genus Juni- 
perus in the Arnold Arboretum and accounts in the literature showed that 
a total of eight species and varieties were susceptible to G. clavipes. 
Hosts were found in two sections of the genus Juniperus. These hosts 
occur over essentially the same geographical range as do the pomaceous 

4. The disease caused by G. clavipes on pomaceous hosts was found 
to occur most frequently on fruits, less frequently on twigs and buds and 
but rarely on leaves. It was most severe on fruits, twigs and buds, usu- 
ally causing marked hyperplastic distortion. Infected buds were not 
only swollen but were forced to develop beyond the usual for the current 
season. On certain fruits, particularly varieties of orchard apples the 
disease produced was limited to small hypoplastic lesions usually at the 
blossom end. On leaves the disease was limited to small, usually partially 

5. On its Juniperus hosts the disease was most abundant on twigs 
from one to five years old but was also found on leaves, branches and the 
main trunk. Diseased leaves were discolored and slightly swollen. They 
were usually killed in one or two years. The disease was perennial for 
several years on twigs and branches. They were usually girdled and cov- 
ered with a thick, flaky or furrowed blackened bark. On the main trunk 
the disease lived for many years but was usually confined to elongated 
swollen patches covered with deeply furrowed and blackened bark. 



406 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

6. The life history of the aecial phase of G. davipes was essentially 
the same on fruits and twigs. It was slower in its development in 
leaves and in fruits of very resistant hosts, in fact in the latter it was 
often aborted. In forced buds the mycelium was essentially systemic 
and developed spermogonia, rarely aecia, progressively "as the buds 
elongated. 

7. The mycelium of the telial phase of G. davipes was confined to 
the epidermis of leaves and to the phellogen of twigs, branches and the 
main trunk of its Juniperus hosts. It remained in leaves for but one, 
occasionally for two years. It was perennial for several years in twigs, 
branches and the main trunk. Telia were produced annually on infected 

8. Several means have been demonstrated for the control of G. 
davipes on pomaceous and on Juniperus hosts. Especial attention has 
been given to finding satisfactory fungicides and formulating practical 
spray programs. Of the fungicides tested Linco colloidal sulfur gave 
very promising results. It was the only one used in field experimentation. 

9. Recommendations with resf)ect to the control of this rust have 
been discussed under the headings of: selective plantings, eradication of 
hosts, removing infected parts from pomaceous and Juniperus hosts, re- 
moving infections from trunks of red cedars and spray applications on 
pomaceous and Juniperus hosts. 

BIBLIOGRAPHY 
Adams, J. F. (1916). Internal Uredineae. (Mycologia, 8: 181-182.) 

(1919). Rusts on conifers in Pennsylvania — I. (Pennsylvania 

Agric. Exper. Sta. Bull. 160: 1-30.) 

(1921). Observations on the infection of Crataegus by Gymno- 

sporangium. (Mycologia, 13: 45^9.) 

Arthur, J. C. (1901). Generic nomenclature of cedar apples. (Proc. 
Indiana Acad. Sci. 1900: 131-136.) 

(1909). Cultures of Uredineae in 1908. (Mvcoloeia, 1: 

225-256.) 

(1929). Plant rusts. (John Wiley and Sons, New York, 

446pp.) 

(1934). Manual of the rusts in United States and Canada 

(Purdue Research Foundation, Lafayette, Indiana, 438pp.) 

Bailey, L. H. (1894). Rust in The Quince in western New York. 

(Cornell Univ. Agric. Exper. Sta. Bull. 80: 625-627.) 
Beach, S. A. (1905). The apples of New York. (New York Agric. 

Exper. Sta. Rep. 1903, Vols. I and II.) 
Cooke, M. C. and C. H. Peck ( 1871 ) . Note on Podisoma. (Jour. Quekett 

Micr. Club, 2:255-268.) 
CooNs, G. H. (1912). Some investigations of the cedar rust fungus, 

Gymnosporangium Juniperi-virginianae Schw. (Ann. Rep. Nebraska 

Agri ^ ^ - --^ 



1935] CROWELL, GYMNOSPORANGIUM CLAVIPES 407 

Crabill, C. H. (1913). Production of secondary sporidia by Gymno- 
sporangium. ( Phytopath. 3 : 282-284. ) 

Crowell, I. H. (1934). The hosts, life history and control of the cedar- 
apple Tust fungus Gymnosporangium Juniperi-virginianae Schw. (Jour. 
Arnold Arb. 15:163-232.) 

(1935). Compilation of reports on the relative susceptibility of 

orchard varieties of apples to the cedar-apple rust disease. (Proc. Amer. 
Soc. Hort. Sci. for 1934. 32: 261-272.) 

(1935). Unusual developments of cedar rusts on pomaceous 

hosts in 1935. (Plant Disease Reporter, 19: 197-198.) 

Dodge, B. O. (1918). Studies in the genus Gymnosporangium — I. Notes 
on the distribution of the mycelium, buffer cells and the germination of 
the aecidiospores. (Brooklyn Bot. Gard. Mem. 1: 128-140.) 

(1918). Studies in the genus Gymnosporangium — II. Report 

on cultures made in 1915 and 1916. (Bull. Torrey Bot. Club, 45: 
287-300.) 

(1918). Studies in the genus Gymnosporangium — III. The 

origin of the teleutospore. (Mycologia, 10: 182-193.) 

(1922). Studies in the genus Gymnosporangium — IV. Distri- 
bution of the mycelium and the subcuticular origin of the telium in G. 
clavipes. (Amer. Jour. Bot. 9: 354-365.) 

(1924). Aecidiospore discharge as related to the character of 

the spore wall. (Jour. Agric. Res. 27: 749-756.) 

(1924). Expulsion of aecidiospores by the May apple-rust. 

Puccinia podophylli Schw. (Jour. Agric. Res. 28: 923-926.) 

(1931). Studies in the genus Gymnosporangium — V. A de- 
structive red cedar rust disease. (Jour. New York Bot. Gard. 32: 

(1933). The orange rust of hawthorn and quince invades the 

trunk of red cedar. (Jour. New York Bot. Gard. 34: 233-237.) 

Farlow, W. G. (1880). The Gymnosporangia or cedar-apples of the 
United States. (Anniversary Memoirs Boston Soc. Nat. Hist. [Sci. 
Papers, 4], 2pl.) 

(1881). Notes on Gymnosporangia. (Bull. Torrey Bot. Club, 

8:85-^7.) 

(1885). Notes on some species of Gymnosporangium and Chry- 



269-307.) 

Hockey, J. F. (1926). 
pathology, Kentville, 
38.) 

Kern, F. D. (1910). The morphology of the peridial cells of the Roes- 
teliae. (Bot. Gaz. 49: 445-452.) 

(1911). A biologic and taxonomic study of the genus Gymno- 
sporangium. (Bull. New York Bot. Gard. 7: 391^83.) 

MacLachlan, J. D. (1935). The hosts of Gymnosporangium globosum 
Farl. and their relative susceptibility. (Jour. Arnold Arb. 16: 98-142.) 

Miller, P. R. (1932). The pathogenicity of three red cedar rusts that 
occur on apple. (Phytopath. 22: 723-740.) 

Mills, W. D. (1929). Cedar rusts, Gymnosporangium Juniperi-virgini- 
anae, G. globosum and G. germinale. (Plant Disease Reporter, Supp. 
70: 188.) 



408 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

Paddock, W. (1902). Quince rust. (Plant diseases of 1901. Colorado 

Agric. Exper. Sta. Bull. 69: 18-20.) 
Palmer, E. J. (1925). Synopsis of North American Crataegi. (Jour. 

Arnold Arb. 6:5-128.) 
Pammel, L. H. ( 1905). Cedar apple fungi and apple rust in Iowa. (Iowa 

Agric. Exper. Sta. Bull. 84: 1-36.) 
Peck, C. H. (1873). Report of the Botanist. (Ann. Rep. New York 

State Mus. Nat. Hist. 25[1871] : 57-123.) 
Ragan, W. H. (1905). Nomenclature of the apple. (U. S. Dept. Agric. 

Bull. 56, issued 1926, 1-396.) 
Rehder, A. (1927). Manual of cultivated trees and shrubs. (Macmillan 

and Co., New York, 930pp.) 
ScHWEiNiTZ, L. D. (1832). Synopsis fungorum in America Boreali media 

degentium. (Trans. Amer. Phil. Soc. II. 4: 141-316.) 
Steinmetz, F. H. and M. T. Hilborn (1934). Observations on Gymno- 

sporangium germinale. (Phytopath. 24:833-834.) 
Stone, G. E. and R. E. Smith (1898). The quince rust. (Rep. Massa- 
chusetts Agric. Exper. Sta. 10:61-63.) 
Stone, R. E. (1909). Studies of Gymnosporangium in southern Alabama. 

(Torreya, 9: 114-117.) 
Sydow, P. and H. (1915). Monographia Uredinearum. III. 
Thaxter, R. (1887). On certain cultures of Gymnosporangium witli 

notes on their Roesteliae. (Proc. Amer. Acad. Arts and Sci. 22: 

259-269.) 

(1889). Notes on cultures of Gymnosporangium made in 1887 

and 1888. (Bot. Gaz. 14: 163-172.) 

(1891). The Connecticut species of Gymnosporangium (cedar- 
apples). (Connecticut Agric. Exper. Sta. Bull. 107: 1-6.) 

Thomas, H. E. (1933). The quince rust disease caused by Gymno- 
sporangium germinale. (Phytopath. 23: 546-553.) 

and W. D. Mills (1929). Three rust diseases of the apple. 

(Cornell Agric. Exper. Sta. Mem. 123: 1-21.) 

(1930). Rust diseases of the apple. (Plant Disease 

Reporter, 14:214-215.) 
Tubeuf, K. von (1906). Uberwinterung des 



(Nat. Zeits. Land Forstw. 5: 217-219.) 
Veimer, J. L. (1917). Three cedar rust fungi, thei 
diseases they produce. (Cornell Agric. Exper. 



EXPLANATION OF THE PLATES 



ressive stages are shown i 
fructifications of Gymnos 



dark colored lesion was caused by 
al fructifications of G. clavipes are 



1935] CROWELL, GYMNOSPORANGIUM CLAVIPES 409 

Fig-. 4. This shoot of A^nelanchier ohlongifolia was inoculated with 
tehospores of G. clavipes. Infection occurred, however, on the 
fruits and pedicels only. 

Fig. 5. Yrmis oi Crataegus mollis m{^ci^dw\i\\ G. clavipes. 

Fig. 6. Fruits of Malus floribunda infected with G. clavipes. 

Plate 156 
Fig. 1 . Twig and thorns of Crataegus sp. infected with G. clavipes. 
Fig. 2. Forced growth of infected buds of Crataegus mollis. The 

diseased buds on the twigs of this species have enlarged greatly. 

Compare with the normal buds on the twig shown on the extreme 

Fig. 3. Another manifestation of the forced growth of infected buds on 
Crataegus sp. On this species the buds and stems have enlarged 
but little while the leaves, though small, have taken on features of 
normal maturity. 

Fig. 4. Another manifestation of the forced growth of infected buds on 
Crataegus sp. On this host the buds have swollen considerably 
and the young stems have elongated but the leaves have remained 
quite stunted in their growth. 

Fig. 5. Infected shoot of Cra/a^5fM.$- P/ia£'no/)3'n<m. This type of symp- 
tom was also found on the English hawthorn, C. Oxyacantha. 

Fig. 6. A globose gall-like swelling of an infected twig of C. Oxyacantha. 

Fig. 7. Spermogonium of G. c/az'i/>e.y. Note its deep, sunken location and 
rotund form. 

Plate 157 

Fig. 1 . Normal aeciospore chain of G. clavipes. Note the regular occur- 
rence of aeciospores and intercalary cells. 

Fig. 2. Abnormal aeciospore chain. The aeciospores are irregular in 
shape as well as in their arrangement with respect to the inter- 
calary cells. 

Fig. 3. Outline drawings of aeciospore 

Fig. 4. Camera lucida drawings of gen 

Fig. 5. Face and side views of peridial cells of G. clavipes. 

Fig. 6. Outline drawings of peridial cells of G. clavipes. 

Fig. 7. Camera lucida drawings of haustoria of G. c/az'i/'c.s as s< 

cells of fruits and twigs of various pomaceous hosts. . 

shown may be found in the same organ of any specific 1 

Plate 158 



Plate 159 
Trunk lesion caused by G. clavipi 
typically oval in outlin 
than normal bark. No 

Several trunk infections on a red cedar tree. Note that the trur 
lesions are found in connection with lateral branches ; many < 
these have died and been removed. 

Telia of G. clavipes in their last stages of gelatinization. At th 
time they are almost shapeless masses. 



JOURNAL OF THE ARNOLD ARBORETUM 



Pla 
Fig. 1 . Shows diagrammatically tl 

of extension of the myccHi; 

its host. This is a camera lucida drawing. 
Fig. 2. The mycelium of G. claznpes in an early stage of extension o^ 

the new phellogen that has recently connected with the existi 

one. Note that the mycelium is found on the phellogen only 

Fig. 3. Formation of the telial sorus. Buffer cells are in various staj 

of development. It will be observed here also that the myceli' 

does not penetrate beneath the phellogen. 
Fig. 4. Early stages in the development of teliospores. The teliosp< 

initials are elongating into the buffer cells. 
Fig. 5. Stages in the maturation of teliospores. These stages are simi 

to those for the maturation of teliospores of G. Junipc 

Fig. 6. Haustoria in the phellogen cells of red cedar. They stand < 
clearly in prepared sections. Two and three haustoria are co 
monly found in a single host cell. 



Jour. Arnold Arb, Vol. XVI 




.. XVI Plate 156 








DIG 



On 



\^ M'l 



>Aeb. Vol. XVI 







Tour. Arnold Arb. Vol. XVI 



■••■^•^ ^/f,^^ 




1935] MacLACHLAN, SPORE DISPERSAL OF GYMNOSPORANGIUM < 



INTRODUCTION 

The basidiospores of Gymnosporangium rusts are thin-walled and so 
are subject to rapid dessication. It would seem then that their effective 
range would be limited to a mile or less but instances have come to the 
attention of the writer which indicate that, under certain circumstances, 
the range may be as great as seven or eight miles. Three of such in- 
stances may be cited. The first was an observation made by Professor 
Roland Thaxter (1887) on basidiospores of G. biseptatum Ellis. He 
found infection on an island off the coast of Maine, eight miles distant 
from the only known source of inoculum. The second instance occurred 
at Lake Wentworth, New Hampshire, and was brought to the attention 
of the writer in the summer of 1933 by Mr. L. S. Mayo. In this locality 
infection by G. davipes Cke. & Pk. and G. globosum Farl. on Amelan- 
chier and Crataegus, respectively, was found on the northeastern side of 
the lake, while the nearest source of inoculum was from a stand of cedars 
on the opposite side of the same lake, more than six miles distant. The 
prevailing winds were from the southwest and had presumably carried 
the basidiospores across the lake. The third instance occurred in the 
Arnold Arboretum, where infection by G. Juniperi-virginianae Schw. and 
G. globosum Farl. on species of Malus and Crataegus, respectively, has 
been observed during the past three years. On both hosts the infection, 
while slight, was markedly uniform over the entire plantations, indicat- 
ing that the sources of inoculum were considerably removed. A detailed 
scrutiny of the surrounding country revealed that no source of inoculum 
sufficient to produce this infection existed within a radius of more than 
six or seven miles. 

In an effort to explain the occurrence of instances such as those that 
have been described, an airplane collection of basidiospores was made at 
different altitudes over infected cedar areas immediately following a 
rainy period in May, 1934; this was followed by laboratory tests on the 
duration of the viability of basidiospores of G. Juniperi-virginianae when 
subjected to varying temperature and humidity conditions. 



JOURNAL OF THE ARNOLD ARBORETUM 



Two areas were selected in which an abundance of basidiospores were 
being discharged, namely, Waltham, Massachusetts, where heavy but 
restricted infections by G. Juniperi-virginianae and G. globosum existed, 
the other about seven miles distant from Waltham at Cochituate where 
there were approximately three acres of red cedars among which were 
scattered apple trees highly susceptible to G. Juniperi-virginianae ; prac- 
tically every cedar tree in the latter area was heavily loaded with galls of 
G. Juniperi-virginianae. 

On May 5, 1934, ideal conditions prevailed for making such a collec- 
tion. Prior to this date there had been intermittent rain, fog, and sat- 
urated humidity for approximately sixty hours; examination of the in- 
fected cedars showed that basidiospores were being released almost con- 
tinually during this period and no high winds had occurred that would 
scatter the spores beyond the possibility of collection. As the plates re- 
vealed after the flight, the air had been washed relatively free of dust and 
smoke so that little trouble was experienced from such contaminations. 
At the time of flight, namely, 8.30 A. M., the clouds had just broken but 
the air was still saturated with moisture, evidenced by the fact that as 
we left the airport water was precipitated out of the air by the propellor 
and thrown on the windshield of the plane. It happened that a south- 
west wind was blowing directly over the area at Cochituate towards the 
Waltham area. Thus, optimum conditions existed for making the 
collection. 

The spores were collected on petri-plates, each plate containing a thin 
layer of 2% pure bacto-agar. To prevent growths by other fungi no 
nutrients were added. Previous tests showed that the basidiospore 
would germinate freely on the agar alone and since they do not grow on 
artificial media, germination was taken as sufficient evidence that the 
basidiospores were viable. Crude but efficient holders for the plates were 
made by using circular lids, whose diameters were slightly greater than 
that of a petri-plate, attached to the ends of wooden rods two and one- 
half feet in length. Wire clamps that could be opened readily served to 
hold the plates in place while the exposures were being made. 

Continuous exposures of one minute each, one overlapping the next 
by fifteen seconds, were made by removing the cover, clamping the open 
plate in the holder and immediately thrusting the latter out through the 
window the length of the handle so that the holder rested on a wind strut. 
Alternate exposures were made from each side of the plane on numbered 
plates. At the same time the course of the flight was outlined on a map 



1935] MacLACHLAN, SPORE DISPERSAL OF GYMNOSPORANGIUM 413 

and by marking the position of the plane at the end of each minute it 
was not difficult to determine afterwards the exact area over which the 
plate had been exposed. 

Exposures were made from within about one mile of the first source 
of spores (the Waltham area) three times at an altitude of 200 feet, then 
directly into the wind at an altitude of 500 feet for seven miles to the 
Cochituate area. Over the latter area four flights were made at 100 ft. 
altitude, then two flights each at altitudes of 500 ft., 1000 ft., 1500 ft., 
and 2000 ft., respectively, finally directly with the wind at an altitude of 
2000 ft. for ten miles, bringing us back again over the Waltham area. 
The average speed during the flight was about 85 miles per hour. Figure 
1 illustrates the course of the flight. 






Note: Each line represents a single exposure; the number in parentheses, below 
the plate number, indicates the number of spores caught on 20 sq. cm. of the plate. 
The topography of the land is shown in profile (from U. S. Geological Survey maps). 

Five hours after the flight the plates were placed in the refrigerator at 
0° C. and spore counts begun. An area enclosing twenty square centi- 
meters was ruled off on the lower side of each plate prior to making the 
count. Within the area fine parallel lines were ruled, the distance be- 
tween any two adjacent lines being slightly less than twice the diameter 
of the field covered by the low power of the microscope. The plates were 
examined from the upper side (lids removed). By moving the plate on 
the microscope stage so that a line (visible though out of focus) was 
just perceptible on one side of the field, and returning across the plate 
so that the next line was just in view on the other side of the field, the 
area enclosed was completely covered without the possibility of either 
overlapping or omitting any of the plate surface to be examined. 

The basidiospores were distinguished from other spores caught on the 
plates by (1) the characteristic yellowish color of their protoplasm as 
seen under strong light, (2) the size and shape of the spores and (3) the 
characteristic germ tubes when present. As a means of comparison 



414 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

basidiospores of G. Juniperi-virginianae and of G. globosum were allowed 
to drop on similarly prepared plates from gelatinized telial masses; these 
two preparations were used for constant reference during the examina- 
tion of the plates. 

Table I gives, for the respective plates, the region over which each 
collection was made, the plate number, the number of spores found on 
twenty square centimeters of the plate and iinally the number of spores 
exhibiting germ tubes. 

TABLE I 

DATA OBTAINED BY MEANS OF AN AIRPLANE COLLECTION OF 

BASIDIOSPORES OF THE GYMNOSPORANGIUM RUSTS 

Altitude Plate Total No. spores 



2000 

Between 2000 

Cochituate 

and Waltham 



is interesting to n 


ote, however, 


2 return flight (g( 


3ing with the 


or so from this a 


rea (Fig. 1). 


the Waltham ar( 


13. was much 


a; this coincided ^ 


vith a smaller 



19351 MacLACHLAN, SPORE DISPERSAL OF GYMNOSPORANGIUM 415 

The count included only those basidiospores of unquestionable iden- 
tity. Spores that were embedded in the agar from the force of impact 
and could not be identified accurately due to the air spaces around them, 
as well as the spores which had burst when they hit the agar, were not 
considered. Certain plates exposed during turning or gaining altitude 
over the Cochituate area were omitted due to the difficulty in determin- 
ing the exact location over which these plates were exposed (see Fig. 1). 

As may be seen from Table I or Fig. 1, basidiospores were picked up 
almost continuously during the entire flight. Approximately one-third 
of them exhibited germ tubes at the time of examination. A sharp in- 
crease in the number of spores collected was evident as the large area at 
Cochituate was approached. Over this area there was a rapid decrease 
in the number of spores collected with increase in altitude, no spores 
being collected at 1500 ft. altitude; this may be accounted for by the fact 
that the wind was carrying the spores away from the immediate cedar 
area below a ceiling of 500 to 1000 ft. 
that spores were again picked up on the i 
wind) at an altitude of 2000 ft., a mile ( 
The number of spores picked up over 1 
smaller than that over the Cochituate area 
amount of infection in the former area. 

Before drawing any conclusion fro 
viable basidiospores may travel, the a 
was carefully examined to determine how much infection existed between 
the two main areas of infection. The survey revealed that scattered in- 
fection loci existed within less than one-half mile of each other over the 
entire area. Thus, although viable basidiospores were found over the 
entire flight it was impossible to determine the exact location of their 
source of dissemination and consequently the distance they had travelled. 

DURATION OF THE VIABILITY OF BASIDIOSPORES OF G. 

JUNIPERI-VIRGINIANAE WITH RELATION TO THE 

FACTORS OF TIME, TEMPERATURE AND 

HUMIDITY 

A knowledge of the length of time during which the basidiospores will 
live when they are subjected to varying temperature and humidity con- 
ditions should, indirectly, give an indication of the distance that the 
basidiospores may travel and still have the potential ability to produce 
infection. Reed and Crabill (1915) state that five to ten days is the life 
limit of basidiospores of G. Juniperi-virginianae in an air-dry condition. 
They also state that in direct sunlight the spores are killed within two to 
five hours; sunlight, then, may be a limiting factor in determining the 



416 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi 

longevity of the basidiospores. However, as has been stated previously, 
dispersal takes place during rainy weather, under which circumstances 
the basidiospores might travel many miles under the cloud line. 

Laboratory tests were conducted in which fresh basidiospores were 
subjected to temperatures ranging from 0° to 35° C. at five degree inter- 
vals and humidities ranging from O'/r to 100/^ at twenty-live percent 
intervals. After varying lengths of time samples of the spores were re- 
moved from the respective environmental combinations and tested for 
germination. 




Such an experiment required eight constant temperature chambers. 
Refrigerators kept in a warm room and equipped with thermostats that 
kept the temperature constant within 0.5° C. served for the 0, 5, and 
10 degree chambers; temperatures of 15, 20, 25, 30 and 35 degrees were 
maintained in De Khotinsky ovens kept in a cold room at 4° C. 

For humidity chambers Kolle culture flasks proved to be very efficient; 
they were of convenient size, exposed a large surface of the humidity con- 
trolling agent with respect to the volume of the flask and could be opened 
without altering the humidity within the flask to any great extent. 
Figure 2 illustrates a typical chamber. 



1935] MacLACHLAN, SPORE DISPERSAL OF GYMNOSPORANGIUM 417 

Zero percent humidity was obtained by placing a layer of dry CaCls 
in the bottom of the flask. Humidities of 25%, 50% and 75% were 
obtained by using aqueous solutions containing, respectively, 55.1%, 
42.8%) and 29.9% of H2SO4. These values were taken from data given 
by Wilson (1921) and are approximately half way between the values 
given for the respective humidities at temperatures of 0° C. and 25° C. 
The greatest difference in percentage of H,S04 to maintain any of these 
humidities over a temperature range of 0° C. to 25° C. is 1.6, which 
would be insignificant when considering such wide humidity intervals. 
Enough H.SO^ solution was poured into each flask to cover the bottom 
to a depth of about one centimeter. The lip between the flask proper 
and the neck served to prevent the H^S04 solution from running out. 
The mouths of the flasks were closed with covers made by cutting gum- 
rubber tubing of the appropriate diameter into four-inch lengths and 
sealing one end with rubber cement. Such flasks could not be used for 
100% humidity as water of condensation would form inside the flask on 
the flat upper surface and drop on the spores. To eliminate this diffi- 
culty, small dome-shaped bell-jars were placed inside moist chambers 
containing distilled water; this type of chamber allowed any water of 
condensation to run down the sides of the dome. 

Small trays of a size that two would conveniently slip into each flask 
and hold the cover glasses bearing the spores well above the humidity 
controlling agents were made out of wire screening. These were coated 
with paraffin (melting point 56° to 58° C.) to prevent the water or 
H.SO^ from coming in contact with the spores as well as prevent corrosion 
of the trays by the HoSO^. 

A complete set of humidity chambers from 0% to 100% was placed 
in each of the respective temperature chambers twenty-four hours before 
the expe