JOURNAL

OF THE

ARNOLD ARBORETUM

HARVARD UNIVERSITY

EDITORIAL BOARD

C. E. KOBUSKI, Editor

I. W. BAILEY I. M. JOHNSTON
R. A. HOWARD KARL SAX
C. E. WOOD

VOLUME XXxXV

vi ie ‘ee 3 3)

Ss

SES

JAMAICA PLAIN, MASS.
1954

Reprinted with the permission of the
Arnold Arboretum of Harvard University

KRAUS REPRINT CORPORATION
New York

1968

No
No

A

Jo

No

T1693
VEC 1 21969

Ge

DATES OF ISSUE

. 1 (pp. 1-86) issued January 15, 1954.

. 2 (pp. 87-202) issued April 15, 1954.

- 3 (pp. 203-274) issued July 15, 1954.

. 4 (pp. 275-390) issued October 15, 1954.

Printed in U.S.A.

TABLE OF CONTENTS
STUDIES IN THE BORAGINACEAE, X XVI. FurTHER REVALUATIONS
OF THE GENERA OF THE LITHOSPERMEAE. By Ivan M. Johnston

Some DETAILS OF THE STRUCTURE OF RHODOTHAMNUS CHAMAECIS-
tus. With one plate. By Herbert F. Copeland

Two NOMENCLATURAL CHANGES IN THE CHINESE Fora. By
Albert N. Steward
STUDIES IN THE KUHNIINAE (Eupatorigak) II. With five plates.
By L. O. Gavser
MiscELLANEOUS MataysIAN Notes. With one plate. By E. D.
Merrill
STUDIES IN THE BORAGINACEAE, X XVII. Some GENERAL OBSER-
VATIONS CONCERNING THE LITHOSPERMEAER. By Ivan M. John-
ston

Ecotypic VARIATION OF THE PHOTOPERIODIC RESPONSE IN Popu-
Lus. With eight text-figures. By Scott S. Pauley and Thomas
O. Perry

Notes ON THE Fiora oF Cuina, III. With two plates. By Shiu-
RTI ERE acs Beste goles dnp ies cuxvde'vaclcsa ek tavdes

THE CYPERACEAE COLLECTED IN NEw GUINEA BY L. J. Brass, IV.
With one plate and two text-figures. By S. T. Blake ..................
PREVERNAL LEAFING OF ASPEN IN UtaH Mountains. With two
plates. By Walter P. Cottam
THE ConTROL OF TREE GRrowTH BY PHLOEM Biocks. With one
plate. By Karl Sax

CRYPTOGAMS OF THE 1948 ARCHBOLD CAPE YORK (QUEENSLAND)

ExpepITIon. By P. Bibby

AppITIONAL Nore on NotuHoracus. With one text-figure. By
C.G.G. J. van Steenis
New ZEALAND Conirers. With one plate. By Vivienne Dellow
Cassie

A MOoNoGRAPH OF THE GENUS PHILADELPHUS. With six plates. By
Shiu-ying Hu

PoLyPLOIDY AND APOMIXIS IN CoTONEASTER. By Hally J. Sav ........

THE Director’s REPORT ON THE ARNOLD ARBORETUM DURING THE
FiscaL YEAR ENDED JUNE 30, 1954

BIBLIOGRAPHY OF THE PUBLISHED WRITINGS OF THE STAFF AND
STUDENTS JULY 1, 1953—J UNE 30, 1954

STAFF OF THE ARNOLD ARBORETUM 1953-1954

INDEX TO VoL. XXXV

TITLE-PAGE AND TABLE OF CONTENTS

JOURNAL

OF THE

ARNOLD ARBORETUM

VoL. XXXV JANuARY 1954 NUMBER 1

STUDIES IN THE BORAGINACEAE, XXVI
FURTHER REVALUATIONS OF THE GENERA OF THE
LITHOSPERMEAE

IvAN M. JOHNSTON

PREPARATORY TO A GENERAL DISCUSSION of the Lithospermeae to be
published in the near future, seventeen genera of the tribe are given indi-
vidual treatment in the present paper. These, along with six genera dis-
cussed previously, Jour. Arnold Arb. 34: 258-299 (1953), include all the
genera which can be referred to the tribe if that is to be a homogeneous
division of the Boraginoideae. Of the seventeen genera here discussed only
one, Lithospermum, has representatives native to both America and the Old
World or has direct relations with genera in both regions. Since the dis-
tribution and relationships of all other genera are confined within one or
the other of these major regions, the primary division in my key to the
genera has been deliberately based on geography. For most uses this will
be a convenience. Furthermore, it also has the advantage of permitting
sharper contrasts of immediately related genera. A synopsis of all the genera
of the tribe and technical keys for their identification will be provided in
the following paper of this series.

KEY TO THE GENERA

PLANTS NATIVE TO AMERICA.

Anthers completely exserted from the throat; filaments elongate, 6-70 mm.
long, exserted 1-65 mm. from the corolla mouth; corolla large, 39-90 mm.
long, trumpet-shaped, lobes usually ascending or recurved or reflexed; pollen
ellipsoidal to ovoid or ovoid-oblong, 23-33 K 15-28 w ....1. Macromeria.

Anthers completely included in the throat or only partially exserted from the
corolla mouth; filaments at most 10 mm. long and usually very much shorter,
completely included or exserted less than 1 mm.; corolla smaller, usually
less than 25 mm. long and never more than 50 mm. in total length, tubular
to salverform, lobes erect to spreadin

Flowers precociously sexual, corolla opening and exposing stamens and style
before attaining full size; corolla-lobes erect, sharply acute or acute with
an attenuate tip, very narrowly imbricate in the bud, usually evidently
longer than broad; sinus between the corolla-lobes plicate and inflexed and
thickened at the base; pollen ovoid, 16-24 X 13-22 p....2. Onosmodium.

2 JOURNAL OF THE ARNOLD ARBORETUM __ [vot. xxxv

Flowers not precociously sexual, corolla opening only when 7 abies
corolla-lobes obtuse or rounded, frequ ently a s broad as long or ev
broader than long, broadly abricate in the art sinus between oe
corolla-lobes neither plicate nor thickened nor inflexed at the base.

Filaments about half the total length of the corolla, twice as long as the
anthers, arising low in the corolla, towards the base shaggy with
slender multicellular gland-tipped hairs; anthers borne high in the
corolla-throat with its sterile tip exserted; tip of anther 1-2 mm. long;
pollen ovoid, 25-28 K 21-33 wm. 2.1... .3. Nomosa

Filaments less than a third of the total length of the ‘corolla and usually
much shorter than the anther, usually arising at or above the middle
of the tube, glabrous or bearing only a few inconspicuous stipitate
glands; anthers without a sterile tip or the tip less than 1 mm

Filaments broadening upwards from a narrow attachment; anthers with a
ll but definite sterile tip, with a well-developed sinus at the base.
Corolla lacking faucal appendages, corolla-lobes deltoid; filaments
oblanceolate, almost as long as the anthers; anthers conspicuously
hairy on the back, with an erect tip, thecae without darkened
margins; pollen ovoid, ais 16-20 “; coarse plant with broad
strongly veined leave 4. Lasiarrhenum.
Corolla with evident faucal eas -corolla- lobes rounded; filaments
triangular or obovate-triangular, half as long as the anther: anther
glabrous, with a recurved tip, thecae with darkened margins; pollen
oy oniy 24-25 X 20 »; small slender-stemmed plant with very
ow veinless leaves........................ 5. Perittostema.
ics pie or subulate or unguiculate: anther only very rarely
bearing a sterile tip, base emarginate or roun
Corolla-throat angulate, externally with a small swelling directly below
sinus; corolla-lobes strictly ascending; filaments partially
exserted from the throat; anthers one half to three fourths exserted
from the throat; inside of corolla completely glabrous and bearing
neither faucal appendanges i racial glands; style exserted;
pollen ellipsoid, 23-26 K 16-22 uw. .... 1... 6. Psilolaemus.
Corolla-throat not angulate, not haat externally below each sinus of
the limb; corolla-lobes usually spreading; filaments and anthers
always included in the throat; corolla inside with faucal appendages
or stipitate glands or both ‘and sometimes — style usually
included; pollen diverse as to size and form. ..12. Lithospermum.

PLANTS NATIVE TO THE OLD Wor Lp,

Anthers without a sterile tip or with the tip small and inconspicuous and rising
abruptly from the emarginate, truncate or rounded summit of the anther;
base of anther usually rounded or simply emarginate; base of theca not
pointed, usually rounded; anthers always distinct, never joined together;
pollen inconspicuously if at all colpate, pores in one or sometimes two rows,
6-9 or

Nutlets narrowed upwards into a prolonged beak, conspicuously paras with

the apex hamate; cymes bractless above the base; upper ves very
ample and evidently veined; corolla-throat inside abundantly ae pues
villose-strigose; Japan............... Ancistrocarya.

Nutlets not conspicuously rostrate, apex not hamate; cymes Pie through-

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVI 3

out; upper cauline leaves not conspicuously ample, usually veinless;
corolla-throat inside not villose-strigose.

Corolla-throat inside bearing 5 well-developed elongate vertical guide-lines
for insect visitors; guides consisting of vertical inflexed plaits bearing
hairs or stipitate glands or both, or consisting merely of elongate
vertical bands of crowded hairs and stipitate glands; anthers usually
with small sterile tips; style usually with a prolonged bilobed sterile
apex, always short, always shorter than the mature nutlets............
Dera e tear e agin eee As 5s ssi AE 8. Buglossoides.

Corolla-throat lacking guide-lines, bearing localized faucal appendages or
bearing stipitate glands which are scattered or are in localized congrega-
tions, or sometimes naked; anthers only very rarely bearing a minute
sterile tip; style only very rarely with a prolonged bilobed sterile apex,
short to elongate, usually becoming much longer than the nutlets

.Nutlet attachment not on the true base of the nutlet, borne ventral to
the nutlet base at the lower end of a stipe which is directed downward
from the ventral side of the ascending nutlet- eae pollen with
biseriate pores; Mongolian... .9. Stenosolenium.

Nutlet attachment on the base of the erect ‘nutlet- body.

Pollen bearing 2 rows of pores, one about each end of the elongate
grain, grains oblong or medially oe upper and lower halves
of the same size and configuration; corolla without faucal append-
ages, its inner surfaces prevailingly devoid of stipitate glands, such
glands when present scanty, inconspicuous, and confined to the
corolla mouth; flowers heterostylic or monomorphic; stamens
usually whorled, affixed at unequal heights above the corolla base
only in one monomorphic species; Asia extending into Africa...
Sree es gemmiage ree a Se aR 6-0iky So A ee 10. Arnebia.

Pollen bearing a single row of pores either about the equator or below
the equator about the lower half of the grain; grains globose,
oblong-ellipsoidal or with the upper and lower halves dissimilar in
size and configuration

Corolla inside without faucal appendages or stipitate glands; annulus
absent; stamens borne at very unequal heights on the corolla-
tube; flowers heterostylic; Caucasus, Armenia, and adjacent
BPN ie) 2.5255 ee Es a ee oe so 11. Echioides.

Corolla inside with faucal appendages or stipitate glands or both;
annulus present; stamens whorled, all borne at the same level
above the corolla base; flowers heterostylic or monomorphic;
Eurasia, Africa, and America.............. 12. Lithospermum.

Anthers narrowed into a prolonged evident sterile tip; base of anther usually
an open sinus and hence somewhat sagittate; basal tip of theca usually
pointed or narrowly prolonged; anthers usually joined to form a synandrium;
pollen evidently colpate, pores in a single row.

Nutlets bilocular; calyx-lobes very unequal, strongly imbricate; anthers distinct
or joined by the entangling of the tail-like appendages borne at the base
of the theca; pollen barrel-shaped or ellipsoidal, pores 8, borne in a well-
developed equatorial groove; plant nearly glabrous, leaves with a cordate-
aMIplexicaul base. <. isxcradeee hea eee wae tas wads 13. Cerinthe.

Nutlets unilocular; calyx-lobes equal or practically so, not imbricate; anthers
usually coherent at the base or along the sides or both; pollen ovoid or

4 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxv

globose or cylindric or ellipsoidal, pores 3, suprabasal or equatorial, not in

an equatorial groove; plants evidently hairy, middle or lower leaves never

with a cordate-amplexicaul base.

Nutlets bent 90° below the middle, strongly incurved, attachment small
and substipitate appearing to be lateral but actually basal on the short
erect lower section of the nutlet; gynobase with elevated pulvinate lobes
each bearing a small attachment face; indument on herbage with inter-
mixed slender gland-tipped hairs; bracts ag large and bape
fruiting calyx frequently on decurved pedice 14,

— bdo and erect or nearly so, with a analy broad snd evidently

attachment; gynobase not bearing the attachment faces on elevated

ie lobes; indument containing no gland-tipped hairs; bracts

rarely becoming conspicuous and foliaceous; fruiting calyx borne on
t or ascending pedicels.

Corolla-lobes well developed, as long as or much longer than the tube,
spreading or decurved; anthers coherent only along the margins of
the terminal appendage, the appendage evidently longer than the
theca; filaments very short and usually bearing a hairy basal append-
age; throa t of corolla frequently a southern Arabia, Socotra,
Somaliland to Angola............ Sibteeeas . Cystistemon.

a. Vaupelia.

Corolla-lobes short, commonly about as long as broad, conspicuously much
shorter than the tubular part of the corolla, erect or loosely recurved
(or, in one species, Onosma longilobum, with the lobes elongate,
longer than the tube, but erect); anthers usually coherent at the base
and frequently also along the margins of the thecae and the append-
ages, the appendage almost always shorter than the theca; filaments
usually elongate, not appendaged at the base; throat of ee never
with spreading hairs; North Africa and Europe to eastern Asi

Calyx-lobes narrow and elongate, more or less parallel, eee ta by a
very narrow or closed sinus; corolla having no puffed-out ribs pro-
jecting between the calyx-lobes : filaments within a corolla all
similar; anthers included to completely exserted from the corolla;
nutlets smooth to rough with the surface only very rarely evidently
are or muriculate; pollen ovoid to spheric or transversely
ellips 16. Onosma

Calyx- apes more or less triangular, ascending, separated by an open
triangular sinus; corolla with puffed-out ribs projecting between the
calyx-lobes ; filaments within a corolla differing in the shape of
their base and i in the orientation of their attachment on the corolla;

antly and minutely papillate or muriculate; pollen cylindric or
vertically ellipsoid; Himalaya and mountains of southwest China.
Ss bS bs FASS ee da bee Pea os PEN Gea das ok ek eee 17. Maharanga.
1. Macromeria D. Don, New Edinb. Philos. =i 13: 239 (1832). Based
upon M. longiflora Don and M. exserta
Philonomia DC. in Steud. a ed. 2, 2: 320 ne Meisner, Pl. Vasc.
Gen. 2: 189 (1836-43), in m.
Onosmodium § ce Gray, Synop. Fl. N. Am. 21: 205 (1878).
Type species, Macromeria viridifiora A. DC.

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVI 5
Macromeria § Macromerioides (Gray) Johnston, Contr. Gray Herb. 70: 13
(1924).

Plant perennial. Stems coarse, erect, simple or bearing a few leafy fer-
tile branchlets in the upper axils, hispid or sometimes hispid-villose or
strigose. Leaves well developed, all cauline, lowest ones smaller than the
upper, hispid, velvety or strigose (hairs on upper surface usually with dis-
coid or bulbose bases), with evident midrib and usually two or more pairs of
well-developed evident assurgent veins. Cymes scorpioid, simple or some-
times geminate, terminating the main stems and frequently also arising di-
rectly from the uppermost leaf-axils and sometimes terminating leafy branch-
lets arising from the upper axils, relatively loose, with the flowers not evi-
dently biseriate, after anthesis becoming straight, very loosely flowered,
and very elongate; bracts numerous, conspicuous and foliaceous, usually
somewhat accrescent in age, lanceolate to broadly ovate. Flowers at an-
thesis borne on strict pedicels at the summit of the straightened portion of
the cyme and hence erect, or borne on the still curved portion of the cyme
and directed backwards over the top of the cyme with the abaxial side
uppermost and the corolla accordingly resupinate. Calyx 5-fid, the lobes
elongate, usually evidently unequal, linear to narrowly lanceolate; pedicels
elongating at maturity, strict or ascending. Corolla yellow, yellowish or
greenish, straight and regular or curved and having the throat prolonged
on the two-lobed adaxial side, elongate, somewhat trumpet-shaped, having

a slender tube which gradually or abruptly expands into an elongate, sub-
cylindric or conic-cylindric throat once to twice as long as the tube, the
outer surface always evidently hairy, the inner surface glabrous or rarely
inconspicuously hairy in the tube and on the lobes and along the veins in
the throat, usually glanduliferous in the throat and sometimes on the
lobes; faucal invaginations evident in one species but usually absent or
only very obscurely developed, when present decorated with glands. An-
nulus absent or represented by a very narrow, continuous or interrupted
lineate ridge just above the base of the tube, glabrous, or in one species
with very inconspicuous tufts of minute hairs. Corolla-lobes equal, elliptic
to deltoid, erect to ascending or reflexed at the base or rarely loosely re-
curved, shorter than the throat, imbricate, apex tending to be acute. Fila-
ments equal, terete or distinctly flattened or even strap-shaped, glabrous
or somewhat hairy in one species, arising well above the middle of the
corolla-throat and always exserted from it but in only one species extruded
more than the length of the corolla-lobes, affixed all at the same altitude on
the corolla or at three superimposed levels with the medial adaxial one
highest, the abaxial pair lowest, and the adaxial laterals at an intermediate
level. Anthers straight or sometimes weakly falcate, elongate, oblong or
oblong-linear, several to many times shorter than the filaments, affixed at
or slightly below the middle, commonly in a groove in the connective, strict
and erect or becoming horizontal by a subapical bend of the filament or
sometimes versatile, apex usually rounded or obtusish and with a small
inconspicuous tip formed by the prolongation of the connective tissue, but
in one species emarginate and bearing a subapical gland; thecae parallel

6 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxv

but unjoined for a short distance above the unappendaged, apparently
emarginate base of the anther; connective evident on the back of the
anther, especially above the middle, glabrous or in one species bearing
short stout hairs above the middle. Pollen broadly to narrowly ellipsoidal
or somewhat ovoid or ovoid-oblong, 25-33 > 15-28 p, sometimes con-
stricted near the middle, upper and lower half of the grain equal or more
or less dissimilar, pores borne eight or nine in a single row at the equator
or below it and sometimes very low on the grain just above its rounded
base, grain in polar profile circular or obscurely polyhedral. Style filiform,
glabrous, longer than the corolla and becoming evidently exserted; stigmas
two, very small, juxtaposed and terminal on the tip of the style or sub-
terminal and separated (though scarcely if at all surpassed) by an apical
prolongation of the style. Nutlets smooth, ovoid or ellipsoidal, usually
smooth, lustrous and white, straight, erect or more commonly diverging
from the pyramidal gynobase; ventral keel weak or absent, suture com-
pletely closed or represented (sometimes only on the upper half of the
nutlet body) by a lineate groove; attachment scar broad, basal, flat or
convex, usually bearing the projecting end of the tubular bony funicular
canal. Gynobase somewhat pyramidal, usually broadly so and commonly
terminated by the thickened four-angulate base of the style; attachment
faces nearly as broad as long, at maturity each surrounded by an elevated
cartilaginous margin, usually sloping and the basifixed nutlets borne upon
them usually divergent.

A genus obviously related to Lithospermum and most closely so to its
Mexican species. It is distinguished only by the form and large size of its
corollas and by having filaments elongate and exserted from the corolla-
throat. Its eight species are well marked and form a very natural as-
semblage. They range between northern Guatemala and Arizona and New
Mexico in southwestern United States, with most of them confined to very
restricted areas in the mountains of Mexico.

Macromeria, as originally defined and established by David Don, in-
cluded two species, M. longiflora and M. exserta. The most detailed sub-
sequent treatments of the genus have been by De Candolle, Prodr. 10: 68
(1846) and by Johnston, Contr. Gray Herb. 70: 13 (1924). Of the species
which have been referred to the genus, only M. cinerascens DC. (1846)
is to be excluded, that being a species of Lithospermum. Both Gray, Synop.
Fl. N. Am. 2': 205 (1878), and Macbride, Contr. Gray Herb. 49: 19
(1917), have suggested that the genus should be reduced to one species,
M. exserta, and that the remaining species be treated as a section of Onos-
modium. As previously noted by me, Contr. Gray Herb. 75: 16 (1924),
such procedure would do violence to natural relationships by separating
M. exserta from its evident relative M. hispida. It would also destroy the
homogeneity of Onosmodium. The eight species of Macromeria are obvi-
ously more closely related to one another than any of them are to any
species of Lithospermum ot Onosmodium, From the latter genus they
differ in size and form of the corolla, length of filaments, shape of the
anthers, and tardily exserted style.

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVI 7

The species of this genus are coarse perennials having few to numerous
stems usually over half a meter tall and frequently approaching and rarely
even surpassing a meter in height. The leaves are all cauline, the lower-
most being smaller and proportionately more elongate than those on mid-
dle sections of the stem. The leaf-blades have a strong midrib and also
several well-developed assurgent lateral veins. In foliage, as well as in
general habit, Macromeria is suggestive of Onosmodium and Laszar-
rhenum, and to a somewhat less degree also of such American species of
Lithospermum as L. viride, L. oblongifolium, L. guatemalense, and L.
cinerascens.

The corollas, 35—90 mm. long, are among the largest in the Boraginaceae,
with those of M. exserta the very largest in the family. They are more or
less trumpet-like in form, having an elongate subcylindric or conic-cylindric
throat usually several times longer than thick and commonly about as long
as the slender tubular portion of the corolla supporting it. The equal, rela-
tively short lobes are ellipsoidal, deltoid, or triangular ovate and com-
monly pointed. They may be straight and ascending or loosely recurving
or abruptly reflexed at the base. In most of the species the corolla is regu-
lar or practically so, but in M. hispida and M., exserta it is curved, the
throat is oblique and evidently prolonged on the adaxial side, and the
stamens are affixed at superimposed levels in the throat, with the adaxial
medial member highest. The corolla in these two species having a very
distinct bilateral symmetry is clearly zygomorphic. This condition is usu-
ally most obvious in the bud of the corolla just before it opens. For most
of its length the bud is gracefully curved adaxially, but in the thickened
apical portion it becomes curved in the reverse direction. The outer thick-
ened portion of the bud, formed of the unexpanded lobes, is evidently
swollen on the two-lobed adaxial side. The bud has its tip not central but
closest to its abaxial side. These zygomorphic corollas of M. exserta and
M. hispida have their two-lobed lip and their medial stamen both on the
adaxial side of the flower.

In most species of the genus, both before and after anthesis, the corolla
stands erect, being borne on strict pedicels at the summit of the straightened
portion of the cyme. In M. hispida and M. exserta the corolla at anthesis,
and also as a mature bud, is borne at a relatively higher position on the
scorpioid cyme, mostly developing on the curve between its arched sum-
mit and the point at which it becomes straight and vertical. Since the
pedicels are strict and borne on the curved portion of the cyme, the flowers
are directed ascendingly backwards over the arched top of the cyme. The
backward direction of the flowers in M. exserta is given further accentuation
by the marked adaxial curving of the throat and tube. As a combined re-
sult of all this, at anthesis the outer parts of the corolla achieve a nearly
horizontal position. By leaning backwards over the tip of the cyme the
corolla has become resupinate.

The corolla in all species except M. viridiflora bears at least scattered
stiped glands in the upper parts of the throat and usually also on the ad-
jacent portions of the lobes as well. In M. leonotis the salient mouth of

8 JOURNAL OF THE ARNOLD ARBORETUM __ [vot. xxxv

the corolla is encircled by a band of glands. In species such as M. barbigera,
M. hispida, and M. exserta, the glands are more scattered, less evident, and
may be distributed from above the level of the filament attachments upward
onto the lower half of the corolla-lobes. In M. Pringlei and less clearly in
M. longiflora the glands are restricted to obscure congregations associated
with small, weak, ill-defined circular or elliptic convexities, one located at
or slightly below each of the corolla-lobes. In M. notata the association
of glands and invaginations is much clearer. In that species there are five
inflexed plaits, cuneate in outline, which have their broad end (1—1.5 mm.
wide) about 2 mm. below the summit of the throat, and from thence,
gradually narrowing, extend outward along the midline of each corolla-
lobe with their pointed end about 2 mm. below the lobe apex. Stipitate
glands are abundant on these elongate swollen areas but practically absent
elsewhere on the corolla. A unique feature of M. viridiflora is the presence
of minute hairs on the inner surface of the corolla. The corolla is villulose
inside the slender tube and may be scantily and minutely strigulose below
the middle of the throat along the vein leading from each corolla-lobe.
Though hairy the corolla bears no glands.

The corollas of Macromeria have no annulus or at most only a weakly
developed one ca. 0.5 mm. above the base of the tube. In M. notata the
annulus is represented by an inconspicuous narrow encircling ridge, and
in M. barbigera by an obscurely five-lobed one, but in all other species it
is obscure or absent. Hairs, these very minute and inconspicuous, were
noted in association with the annulus only in M. leonotis.

The stamens are affixed in the corolla-throat and within an individual
flower are all at about the same distance (4-10 mm.) below the base of
the corolla-lobes. In regular corollas the filaments arise at equal distances
above the corolla-base. In zygomorphic corollas, however, the throat is
prolonged on the adaxial side and the mouth of the throat is oblique. Be-
cause the stamen attachments have a fixed relation with the summit of
the throat, those on the prolonged adaxial side of the corolla accordingly
have a position at a greater distance above the corolla-base than those on
the abaxial side. This is the condition in M. exserta and M. hispida.

The filaments are well developed. In M. exserta they may become 70
mm. long, and accordingly are very conspicuously long-exserted from the
corolla. The shortest ones (6-9 mm.) occur in M. notata. In other species,
however, the filaments are mostly 10-20 mm. long. They are clearly ex-
serted from the corolla-throat but do not surpass the corolla-lobes when
the latter are in an erect position. In most species the filaments are slightly
compressed. In M. viridiflora, however, they are strongly so, being strap-
shaped, 1.2-1.5 mm. broad just above their base, and then gradually nar-
rowed upwards to become 0.5—1 mm. wide at the apex. In M. leonotis the
filaments may be sparsely hairy below their middle and sometimes also on
their decurrent bases, but in all other species they are always glabrous.

The anthers are Slongate and have an oblong outline. They are affixed
to the filament at or perceptibly below the middle, usually in an elongate
depression in the connective. Although past authors have stated that they

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVI 9

are usually versatile, that condition exists, if at all, only in M. exserta. In
M. hispida, M. longiflora, M. Pringlei, and probably M. leonotis also, the
anther may assume a position at right angles to the filament, but that
appears to be the result of a ninety-degree bend of the filament just below
its apex. In M. viridiflora and apparently also in M. barbigera the anthers
are strictly and firmly affixed to the filament and are erect. The connective
is glabrous in most species, but in M. viridiflora, in which the structure
reaches its maximum development, it may bear numerous stout hairs on the
back side of the anther above the middle. In most species the connective
is slightly prolonged to form a minute, usually thickish, truncate or subulate
apicule on the broad summit of the anther. In M. viridiflora the connective
is thickened, apparently glandular, and locally very hairy just below the
anther tip. It is not prolonged beyond the thecae. The anthers in this
species, accordingly, are emarginate at the tip.

The connective is not only broadest but also most readily observed in
M. viridiflora, M. Pringlei, and M. longiflora. In these species the thecae,
after shedding pollen and becoming explanate, are displayed obliquely on
the ventral side of the anther. The back side of the anther is nearly plane,
and the broad connective and the filament attachment are fully exposed
to view. In the five other species of the genus, however, the explanate
thecae have reflexed lower halves, and their surfaces become parallel and
face left and right in opposite directions. Such mature anthers are strongly
compressed laterally. Their back appears to be strongly conduplicate with
the narrow connective and the filament attachment hidden in the depth
of the fold.

The style is filiform and sufficiently elongate to bear its stigmas com-
monly about 5 mm. beyond the anthers. The minute stigmas are sub-
terminal, being separated and surpassed by the tip of the style in M.
Pringlei, M. leonotis, and M. notata, but in the other species they are ter-
minal and juxtaposed upon the apex of the style.

As in most other genera of the Lithospermeae, in herbarium specimens
I have found the anthers of Macromeria dehiscent and the pollen shed in
those flower-buds in which the corolla is nearly ready to open. Just previ-
ous to the opening of the corolla, the stigma in seven of the eight species
is pushing against the very top of the bud-cavity, and hence is above the
anthers. The stamens have attained almost full development before the
corolla opens. In M. exserta, however, the very elongate, eventually long-
exserted filaments and style are not completely lengthened before the
corolla expands. The filaments have their middle portion lying pressed
against the curve of the swollen adaxial side of the outer half of the bud.
At the tip of the bud cavity the distal portion of the filaments curves back-
wards for 180° to 360°. The anthers usually lie within the adaxial half
of the most ample part of the bud cavity. The style, shortened by undulate
contortions, bears its stigmas appressed against the tip of the bud cavity
or against the abaxial side of the cavity close to the tip. In Echium, in
which elongate filaments and style are also curved and contorted inside
the flower bud, the stigmas before the corolla opens may have a position

10 JOURNAL OF THE ARNOLD ARBORETUM _ [vot. xxxv

amidst the clustered dehiscent anthers. In the bud of Macromeria, how-
ever, the stigmas have a position above or beyond the anthers and so de-
cidedly less accessible to any pollen set free in the bud. The stigma is not
precociously exserted. It escapes from the flower bud only when the im-
bricate corolla lobes have begun to loosen — just before the corolla opens,
or, in many cases, not until the corolla is almost completely open.

Macromeria has pollen similar to that of Lithospermum, and especially
like that of the Mexican members of that genus. Each species of Macro-
meria has distinctive pollen with the grains recognizable if not by shape
at least by size. Related species agree more closely in the length of their
grains than in the form of them.

The pollen bears eight or nine pores in a single row. Its pores are fre-
quently detectable as minute swellings on the outline of the grains when
the latter are viewed in lateral profile. Frequently they are sufficiently
prominent to give the sides of the grains a somewhat obtusely angled sil-
houette. The polar profile of the grain is usually circular, and only very
rarely are the pores sufficiently evident to produce a vaguely polyhedral
outline. Evidences of possible shallow furrows on the grain have been
detected only in M. barbigera. The pores are equatorial, and the upper
and lower halves of the grain are equal in M. leonotis (grains globose
ellipsoidal, sides angulate, 25-26 & 23 »), M. barbigera (ellipsoidal, sides
rounded or angled, 30-33 25-28 »), M. notata (ellipsoidal, sides tending
to be angled, 28-30 « 23-25 »), and M. Pringlei (ellipsoidal, sides rounded
to nearly parallel, 25-26 15-18 ,). In two species, M. exserta and M.
hispida, the grains bear the pores very slightly but still perceptibly below the
middle. The lower half of the grain has a more evenly and broadly rounded
curve than the upper half. Macromeria exserta has ovoid-ellipsoid grains
(38-41 X 25-28 n), and M. hispida globose-ovoid grains (38-41 & 33—
37 «). The pollen of the two remaining species of the genus is elongate,
perceptibly constricted below the middle, and bears its row of pores above
its broad rounded base, where its diameter is greatest. Constrictions of
the grain begin directly above the row of pores and form “shoulders” of a
type previously noted in grains of a similar type in Lithospermum, cf.
Jour. Arnold Arb. 33: 310 (1952). Such elongate, medially constricted
grains are characteristic of M. longiflora (25-28 & 16-18 p) and M. viridi-
flora (28-33 >< 18-22 y). Of these two species with constricted pollen,
M. viridiflora has no close relatives. Macromeria longiflora, however, is
obviously most closely related to M. Pringlei and species having ellipsoidal
grains with equatorial pores. The only agreement is in the length of the
grains,

The nutlets of Macromeria are smooth, shiny, and usually white, and
are symmetric or nearly so and nearly circular in transverse section. In
general appearance they closely resemble those of most American species
of Lithospermum. The venter of the nutlet is only obscurely if at all
keeled. The ventral suture may be represented by a very narrow lineate
groove (sometimes not half the length of the nutlet) or it may be com-
pletely fused and obliterated. It may be present or absent or vary in

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVI 11

length not only within the species but also among the nutlets produced by
a single plant. The large, nearly circular attachment scar, horizontal or
very slightly oblique, is flat or slightly convex and is always distinctly basal.
In all species towards its ventral margin the scar bears a prominence com-
posed of the slightly protruding broken end of the well-developed bony
tubular funicular canal. In the detached nutlets of M. viridiflora, M. bar-
bigera, and M. notata, the scar also bears another process, this small and
obliquely ascending and apparently representing protrudent tissue about
the base of one of the vascular traces leading to the dorsum of the nutlet
body. This dorsal prominence on the scar appears to be absent on the
nutlets of M. exserta. Since thoroughly mature, self-detached nutlets of
M. longiflora, M. Pringlei, M. hispida, and M. leonotis have not been seen,
the nature of the scar in these species is unknown. The gynobase in all
species is well developed, broadly pyramidal, and terminated by a persist-
ing, more or less thickened four-angulate base of the style. The attach-
ment surfaces of the mature gynobase are usually each encircled by a
coarse thickened and elevated cartilaginous margin. When all four nutlets
are matured, the gynobase has more or less distinctly sloping attachment
faces, and the nutlets they bear, being straight and basifixed, are accord-
ingly divergent.
KEY TO THE SPECIES

Corolla with erect or ascending lobes.
Back of anthers hairy above the middle; filaments strap-shaped; corolla
without ca on the inner surface, villulose in the tube; stigmas terminal
Yh Cr ee oe ab Sake bn ee RO 1. M. viridiflora.
Back of ates glabrous; filaments not strap-shaped; corolla glanduliferous
in the throat, tube glabrous inside; stigmas usually subterminal, separated
by the sterile tip of the style.

ashe bearing 5 evident cuneiform plaits, these glanduliferous and extend-
g from below the corolla-lobes upward upon the net to ste
rite PUGOIE? «<2, 35:4 ee ee wae kk ae ee M. notata.
Corolla without cuneiform glanduliferous plaits, bearing ae rarenaeeiaaeies
small circular or elliptical congregations of glands at or below the base

of each corolla-lobe; upper face of lobes glandless
Upper surface of leaves without evident hairs, suet and very
minutely verrucose or muriculate and hence at most scabrellous;
pore eas below the middle and bearing the pores above the
PUNGOt DASE: vc 5S es ee Roo a ane ed 3. M. longiflora.
aes surface of leaves bearing evident stiff appressed hairs usually
arising from evident pallid dot-like bulbose or discoid bases, surface
scabrous; pollen ellipsoidal, broadest at the middle, pores equatorial.
4. M. Pringlei.

Corolla with lobes reflexed or (in no. 6) loosely recurved.
Mouth of corolla densely glanduliferous in a continuous band; stigmas sub-
terminal, ae and surpassed by the short sterile apical prolongation
OTe EO SEVIS icc 5 i eo I ee cae als 5. M. leon
Mouth of ai not conspicuously glanduliferous; stigmas terminal.
Corolla-lobes loosely recurved, deltoid, about as long as broad; corolla

12 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxv

regular, symmetric in the bud, stamens all affixed at the same distance
above the corolla-base; pollen ellipsoidal, pores equatorial..........
eT OT ee Pe ere ee ee ee . M. barbigera.
Corolla-lobes becoming reflexed, elongate; corolla zygomorphic, asymmetric
in the bud, throat oblique, stamens on adaxial side affixed higher than
those on the opposite side of veges — bearing the row of pores
slightly below me middle of the gra
Filaments 10-15 mm. long; corolla 50-55 mm. long; middle stem leaves
usually lanceolate; pollen globose-ovoid .............. 7. M. hispida.
Filaments 55-70 mm. long, conspicuously long-exserted; corolla very
large, 60-90 mm. long; middle stem leaves oblanceolate: pollen ovoid-
CUIDSOIGAL 56.5.5. dc¢ aay acks s BY sak pawn eg ors eee vi 8. M. exserta.

Macromeria viridiflora DC. Prodr. 10: 68 (1846); Sessé & Moc.
Calq. Fl. Mex. t. 904 (1874); Johnston, — Arnold Arb. 30: 110
(1949). Type from Mexico, Sessé & Moci

Macromeria longiflora sensu Johnston, Contr. ae Herb. 70: 14 (1924).

—

Stems 5—10 dm. tall. Cymes terminal on the stems and frequently gemi-
nate and commonly borne also on short leafy branchlets arising from the
uppermost axils. Calyx becoming 25 mm. long, largest lobe up to 2.5 mm.
broad and the smallest 1-1.5 mm. broad. Corolla greenish yellow, 65-80
(usually 70-75) mm. long, regular or nearly so, outside with abundant
appressed and scattered spreading hairs 0.7—2 mm. long, inside sparsely
and inconspicuously strigulose along the veins below the lobes and villulose
inside the narrow tube, stipitate glands very few or absent, annulus absent
or very imperfectly developed. Corolla-lobes triangular-ovate, acute, as-
cending, 8-10 mm. long, 5—~7 mm. broad above the base, usually greenish
and sometimes strigose along the middle of the upper surface. Filaments
equal, 16-20 mm. long, borne 8-10 mm. below the base of the corolla
sinus, broad and strongly compressed, strap-shaped, broadest (ad 1.5 mm.)
slightly above the base and then very gradually narrowed towards the apex
(ca. 1 mm, wide). Anthers 4—5.5 mm. long, affixed at or slightly below
the middle, strictly erect, apex emarginate, base shallowly lobed; connec-
tive on upper half of dorsum broad and bearing scattered coarse appressed
hairs usually most abundant below the gland at its summit. Pollen elongate,
28-33 > 18-22 pu, somewhat constricted below the middle, bearing the
row of pores where broadest 8-10 » above the base of the grain. Stigmas
two, distinctly terminal. Nutlets 3-3.5 mm. long, with an obscure ventral
keel, suture absent or represented by a lineate groove.

Mountains of northern Mexico, in the states of Chihuahua and adjacent
Durango, Sinaloa, and Sonora, and extending north into the United States
only in the mountains (Chiricahua and Huachuca Mts.) of southeastern
Arizona.

la. Macromeria viridiflora var. Thurberi (Gray), comb. nov.
Onosmodium Thurberi Gray, Synop. Fl, N. Am. 21: 205 (1878), Type from

western New Mexico.
Macromeria Thurberi (Gray) Mack. Bull. Torr. Bot. Cl. 32: 496 (1905).

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVI 13

Flower smaller than in typical M. viridiflora. Calyx becoming 20 mm.
long, lobes 1—1.5 mm. broad; corolla 35—50 (usually 40-45) mm. long; fila-
ments 12-14 mm. long, borne 6-8 mm. below the corolla sinus; anthers
3—4.5 mm. long.

Occurring in the mountains from middle eastern New Mexico west to
central Arizona in an area to the north of that occupied by typical M. viridi-
flora.

A very well marked species readily recognized by its strap-shaped fila-
ments and dorsally hairy anthers, and also by the villulose inner surface
of its corolla-tube.

The typical form of the species has slender, very elongate corollas. Like
many other plants of the Sierra Madre Occidental, true M. viridiflora has
a range that extends north into the Chiricahua and Huachuca mountains,
just north of the United States-Mexican boundary, but no further to the
northward. Some collections from the Chiricahua and Huachuca moun-
tains are indistinguishable from those of Mexican origin, but others have
their flowers somewhat smaller. Even in these latter, however, the corollas
are larger than those on plants referable to var. Thurberi which occur
further northward in New Mexico and central Arizona. The differences in
flower size are geographically correlated and sufficiently striking to merit
nomenclatorial recognition.

2. Macromeria notata, sp. nov.

Planta perennis 4—5 dm. alta e radice valida purpureo-tincta erumpens;
caulibus simplicibus foliosis erectis breviter hispidis basim versus ad 5 mm.
crassis; foliis numerosis lanceolatis evidenter venosis acutis, eis medium
versus caulis gestis majoribus 6—8 cm. longis 16—20 mm. latis, pilos rigidos
adscendentis 0.2-1 mm. longos e basi bulboso vel discoideo minuto non
rariter pallido erumpentibus proferentibus, facie inferiori pallidioribus venis
prominulis ornatis; cyma scorpioidea solitaria caules simplicos terminanti;
calyce 15-18 mm. longo, lobis inaequalibus 1-2.5 mm. latis, pedicellis sub
anthesi 0-5 mm. longis maturitate 10-15 mm. longis; corolla flavescenti
regulariter 50 mm. longa extus hispidula pilis 0.5—-1 mm. longis plus minusve
curvatis ornata, intus glabra et solum in appendiculis faucium glandulifera;
tubo 2—2.5 mm. longo 1—-1.5 mm. crasso; annulo anguste annulato; faucibus
1.5—-2 cm. longis gradatim ampliatis apicem versus 5—6 mm. crassis; ap-
pendiculis faucium depressis elongatis a 2 mm. infra basim loborum corollae
sursum fere ad 2 mm. infra apicem loborum corollae prolongatis cunei-
formibus leviter invaginatis prominulis dense glanduliferis basi 1-1.5 mm.
latis truncatis; lobis corollae triangularibus adscendentibus viridescentibus
ca. 5 mm. longis et latis summum ad apicem rotundis secus medium appen-
diculi faucium glanduliferi decurrenti notatis alibi minute sparseque strigu-
losis; filamentis 6-9 mm. longis ca. 5-7 mm, infra basim sinuum limbi
affixis: antheris 2—2.5 mm. longis rectis apice haud appendiculatis; pol-
linis ellipsoideis 28-30 * 23-25 yp, poris 8 secus equatorem dispositis;
stigmatibus 2, terminalibus; nuculis 4 mm. longis laevibus albis ovoideis
obscure asymmetricis obscure carinatis, late basaliter affixis, cicatrice con-

14 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxv

vexo; gynobase — pulvinis distinctis margine cartilagineis promi-
nulis ‘circumdatis

MEXICO (Nuevo Leon): Ascent of Sierra Infernillo, ca. 15 miles south
of Galeana, 9-10,000 ft., common over small areas just below peak, fl. yellow,
June 16, 1934, C.H. & M.T. Mueller 830 (type, Gray Herb.) ; canyon below Las
Canoas on Cerro Potosi, scattered in dense shade on arroyo bank, July 20, 1935,
C. H. Mueller 2238 (G).

The distinctive features of this species are the weakly invaginate elong-
ate densely glanduliferous plaits which extend from the upper part of the
throat out upon each of the corolla-lobes. These glanduliferous plaits are
the closest approximation in the genus to the localized well-developed faucal
appendages present in many species of Lithospermum. In M. longiflora and
M. Pringlei there are small vague convex areas bearing glands at or below
each lobe of the corolla, but in other congeners of M. notata, even this
suggestion of faucal invagination is lacking. The present species is cer-
tainly a very distinct one. Its closest relations are possibly with M. Pringlei
and M. longiflora.

3. Macromeria longiflora [Sessé & Moc.] D. Don, Edinb. New Philos.
Jour. 13: 239 (1832); Johnston, Contr. Gray Herb. 92: 93 (1930).
Type from Mexico, Sessé & Mocino.

Lithospermum longiflorum Sessé & Moc. ex D. Don, Edinb. New Philos. Jour.
13: 239 (1932), in synonym.

Onosmodium longiflorum (Don) Macbr. Contr. Gray Herb. 49: 21 (1917).

Lithospermum flavum Sessé & Moc. Fl. Mex. 32 (1893): Johnston, Jour.
Arnold Arb. 30: 109 (1949). Type from Michoacan, Mexico, Sessé &

ocino.

Macromeria discolor Benth. Pl. Hartw. 49 (1840). Type from Mexico.

Onosmodium discolor (Benth.) Macbr. Contr. Gray Herb. 49: 20 (1917).

Cymes terminal on the stems and frequently also arising directly from
the uppermost axils. Calyx 13-15 mm. long, lobes slender, 1-1.5 mm
broad, unequal. Corolla 45-55 mm. long, regular or very obscurely pro-
longed on the adaxial side, outside clothed with slender spreading or loosely
appressed hairs up to 1 mm. long, inside bearing inconspicuous elliptic
congregations of glands on obscurely convex areas below each corolla-lobe
and sometimes glands also along the decurrent base of the filaments, an-
nulus not developed. Corolla-lobes elongate, lance-oblong, acutish or the
tips obtusish, 11-15 mm. long, 4—7 mm. broad, ascending, upper face with-
out hairs or glands. Filaments 14-15 mm. long, affixed ca. 4 mm. below
limb-sinus, anthers 2-4 mm. long, affixed at or near the middle, erect,
apex minutely appendaged. Pollen elongate, 25-28 16-18 ,, constricted
below the middle, broadest and bearing the row of pores about 8 » above
the base. Sivas 2, usually subterminal. Nutlets ovoid, erect, symmetric,
4 mm. tall, 3 mm. thi ck.

Mountains of western Mexico from Colima to Oaxaca.

A species probably most closely allied to M. Pringlei, from which it dif-
fers in its more herbaceous stems and its larger leaves devoid of evident

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVI 15

hairs on the upper surface. Although at first glance the upper leaf face
appears to be smooth and glabrous, close examination under the microscope
shows it to be actually dotted with minute mineralized warts or conic
muriculations. When the minute roughenings are pointed, the surface
may be perceptibly scabrellous. The minute mineralized roughenings on
the leaf of M. longiflora are evidently homologous with the very much
coarser bulbose or discoid hair bases present on the upper lea‘ face of M.
Pringlet. The lower face of the leaves in M. longiflora is strigose with an
abundance of short, stiff, closely appressed hairs. It is usually cinereous
and contrasts very strongly with the upper surface, which in most her-
barium specimens dries a chocolate brown.

4. Macromeria Pringlei Greenm. Proc. Am. Acad. 34: 570 (1899).
Type, Sierra de Pachuca, Hidalgo, Pringle 11044.

Onosmodium Pringlei (Greenm.) Macbr. Contr. Gray Herb. 49: 20 (1917).
Macromeria guatemalense Johnston, Jour. Arnold Arb. 29: 232 (1948). Type,
Volcan Tajumulco, dept. of San Marcos, Guatemala, Steyermark 35898.

Cymes terminal on the stems and frequently also terminating short
axillary branches arising from the uppermost axils. Calyx 12-15 mm.
long, lobes slender, 0.5—1.2 mm. wide. Corolla pale yellow-green, 35-53
mm. long, regular, straight, outer surface bearing slender ascending hairs
ca. 1 mm. long, inside glabrous bearing very scattered glands, or these only
in very vague slightly swollen areas at or below the base of the corolla-
lobes. Corolla-lobes triangular, acute, erect or ascending, 9-10 mm. long,
5.5—7.5 mm. broad, apex obtusish, upper face usually without hairs or
glands. Filaments 9-12 mm. long, equal, affixed in the throat 3-4 mm.
below the base of the corolla sinus. Anthers 2—3.5 mm. long, medio-affixed,
apex minutely appendaged. Pollen ellipsoidal, sides rounded or nearly
parallel, 25-26 & 15-18 yw. Stigmas 2, subterminal. Nutlets erect, sym-
metric, pointed, ca. 4 mm, tall, 3—3.5 mm. thick, without a ventral keel.

Mountains of Mexico in the states of Hidalgo, Guerrero, and Oaxaca.

Closely related to M. longiflora but a less vigorous plant with more slen-
der fruticulose stems and smaller, usually more elongate leaves bearing
evident appressed hairs on the upper surface. Macromeria guatemalense of
northern Guatemala appears to differ from Mexican M. Pringlei only in
its more elongate, more decidedly fruticose stems and smaller (to 35 mm.
long) corollas. Possibly it may represent a southern variety of M. Pringlei,
but hardly a species distinct from it.

5. Macromeria leonotis Johnston, Jour. Arnold Arb. 16: 188 (1935).
Type, ascent into Taray, Sierra Madre ca. 15 mi. s.w. of Galeana,
Nuevo Leon, Mueller 754.

Cymes terminal on the stem. Calyx 19-25 mm. long, lobes unequal, very
slender, 1—-1.5 mm. wide. Corolla yellow, 55-75 mm. long, slightly zygo-
morphic, straight or somewhat curved, outside hispidulose-villulose, densely
clad with short curly hairs 0.3-0.7 mm. long, inside glanduliferous on the

16 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxv

lower part of the lobes and over the upper half of the throat and most
abundantly so in a band at the throat summit; annulus a very narrow
interrupted ring bearing five groups of short, very inconspicuous hairs.
Corolla-lobes triangular-oblong, 7-12 mm. long, 5—8 mm. wide, becoming
reflexed, apex rounded, upper surface minutely strigulose or rarely glabrous,
glanduliferous at the base. Filaments 14-20 mm. long, compressed, affixed
7-8 mm. below the corolla sinus, sometimes hairy below the middle and
on the decurrent base. Anthers 2—3.5 mm. long, affixed just below the
middle, apex minutely appendaged. Pollen broadly globose-ellipsoidal, sides
angulate, 25-26 23 mw, pores 8, equatorial. Stigmas apparently sub-
terminal. Nutlets not seen.

Mountains of northeastern Mexico in states of Nuevo Leon and Tamauli-

as.

A species notable for its short soft indument and abundantly glandu-
liferous summit of the corolla-throat. The corolla appears to be slightly
zygomorphic, being somewhat curved and perhaps also slightly prolonged
on the adaxial side. The elongate lobes are strongly reflexed at the base
and the opening into the throat is accordingly the most forward part of the
corolla. The rounded lip surrounding this opening to the throat is densely
glanduliferous. Some glands are present on the lower part of the corolla-
lobes and also inside the throat down at least to the level of the stamen
attachments, but only on the lip about the opening into the throat are
they extremely abundant. No other species of the genus has corollas with
glands so numerous. The species is a very distinct one, but seems to share
more characters with M. longiflora and M. Pringlei than with other con-
geners.

6. Macromeria barbigera Johnston, Jour. Arnold Arb. 16: 189 (1935).
Type, slope of Sierra Tronconal, ca. 15 mi. s.w. of Galeana, Nuevo
Leon, Mueller 741.

Cymes terminal on the stem and also arising from the uppermost leaf
axils. Calyx 15-20 mm. long, lobes very slender, 1—1.5 mm. broad, un-
equal. Corolla (“cream-white” but when dry with a distinctly reddish
throat and somewhat greenish lobes) 45-65 mm. long, regular, with a very
slender tube 15-30 mm. long which abruptly expands into a cylindric
throat that is 20-30 mm. long, 5-10 mm. thick, and sometimes perceptibly
broadest below the middle, outer surface of corolla villose, the hairs slender,
white, and 2~3 mm. long, inner surface glanduliferous above the level of
the filament attachments and especially so on the upper face of the lobes,
annulus an interrupted ring. Corolla-lobes deltoid, 4-5 mm. long, loosely
aie distinctly recurved especially beyond the middle. Filaments 12-15

m. long, affixed 8-10 mm. below the base of the corolla sinus. Anthers

3 mm. long, medio-affixed, apex minutely appendaged. Pollen ellipsoidal,

sides rounded or angled, 30-33 X 25-28 p, pores 9, equatorial. Stigmas
terminal. Nutlets ellipsoidal to ovoid, 3—3.5 mm

Northeastern Mexico in the mountains of Nuevo Leon and Tamaulipas.

A very distinct species notable for its coarse habit, bristly indument, and

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVI 17

broad leaves, as well as for the form of its large corollas. The corollas
have a very slender tube, an abruptly expanded elongate throat, and
loosely recurved deltoid lobes glanduliferous on the upper surface. The
immediate relatives of the species are obscure.

7. Macromeria hispida Mart. & Gal. Bull. Acad. Brux. 11: 339 (1844).
Type, near Morelia, Michoacan, Galeotti 1917.
Macromeria longiflora var. hispida (M. & G.) A. DC. Prodr. 10: 68 (1846).
Onosmodium longiflorum var. hispidum (M. & G.) Macbr. Contr. Gray Herb.
49: 21 (1917).
Macromeria longiflora sensu DC. Prodr. 10: 68 (1846).

Cymes terminal and commonly also arising directly from a number of
the uppermost axils to form an elongate thyrse. Calyx with very unequal
lobes, the abaxial one largest and becoming 18-22 mm. long and 1.5-—2.5
mm. broad. Corolla yellow, 50-55 mm. long, zygomorphic, usually some-
what curved and the throat prolonged on the adaxial side, in the bud
thickly clavate with the outer half swollen and evidently more rounded
on the adaxial side, outer surface of corolla bearing numerous ascending
or loosely appressed hairs 0.5—1 mm. long, inside bearing scattered incon-
spicuous slender gland-tipped hairs in the throat above the stamen attach-
ments and also on the upper face of the lobes; annulus not developed.
Corolla-lobes equal, elongate, ovate-elliptic, 8-10 mm. long, 6—7 mm. broad
below the middle, becoming reflexed, apex rounded, margins revolute.
Filaments equal, 10-15 mm. long, affixed at slightly unequal heights above
the corolla base, the medial adaxial filament ca. 2 mm. higher than the
abaxial pair and ca. 1 mm. above the adaxial pair, all attached 4-5 mm.
below the oblique summit of the throat. Anthers 2-3 mm. long, affixed
slightly below the middle, erect, apex frequently bearing a slender minute
appendage. Pollen globose-ovoid, 38-41 * 33-37 ym, upper and lower
halves of the grain slightly unequal, pores eight in a row and borne slightly
below the middle. Stigma terminal. Nutlets not seen.

A Mexican plant known only from the state of Michoacan, especially
near Morelia and Patzcuaro.

In floral organization this species most resembles M. exserta, but in
general habit it is more suggestive of WM. Pringlei and M. longiflora.

8. Macromeria exserta D. Don, Edinb. New Philos. Jour. 13: 239
(1832); Lindley, Bot. Reg. 33: t. 26 (1847). Type from Mexico,
Sessé & Mocino.

Echium longiflorum Sessé & Moc. Pl. N. Hisp. 20 (1888); Johnston, Jour.
Arnold Arb. 30: 109 (1949).

Macromeria exserta var. imparata Macbr. Contr. Gray Herb. 49: 22 (1917).
Type from Oaxaca, 1842, Ghiesbreght.

Cymes terminal and commonly also arising from the uppermost axils to
form a loose thyrse. Calyx 20-30 mm. long, lobes very unequal, 1-7 mm.
broad. Corolla very large, 60-90 mm. long, yellow, zygomorphic, with the

18 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxv

tube curved and the throat prolenged on the adaxial side, in the bud with
its thickened distal half conspicuously distended on the adaxial side, out-
side of corolla bearing abundant spreading hairs usually ca. 0.5 mm. long,
inner surface of throat and lobes bearing scattered, usually inconspicuous
slender gland-tipped hairs; annulus not developed. Corolla-lobes elongate,
equal, 15-28 mm. long, 5-8 mm. broad, becoming reflexed, apex rounded
or obtusish, margins revolute. Filaments very slender, elongate and long-
exserted, 55-70 mm. long, usually curved, affixed in the corolla-throat
4—5 mm. below its oblique summit at three superimposed levels, the odd
medial stamen 5-8 mm. higher above the corolla base than the abaxial
pair and 3-4 mm. above the adaxial pair. Anthers 3—4 mm. long, frequently
curved, affixed below the middle, becoming versatile, apex usually bearing
a minute appendage. Pollen ovoid-ellipsoidal, 38-41 > 25-28 p, upper
and lower halves of grain slightly dissimilar, pores 8, slightly inframedial.
Stigmas terminal. Nutlets 4—5.5 mm. long, 2.5-4.5 mm. thick, weakly
keeled

Mountains of western Mexico from Nayarit to Oaxaca.

A very distinct species notable for its very coarse habit and extremely
large zygomorphic corollas with excessively long-exserted stamens. Be-
cause of the curved adaxially swollen buds and the eventually long-exserted
curved stamens the corolla has a form and appearance more suggestive of
some of the Verbenaceae, e.g., Clerodendron, than of other members of the
Boraginaceae. In size the corolla of M/. exserta surpasses that of all other
species in the whole of the Boraginaceae.

2. Onosmodium Michx. Fl. Bor. Am. 1: 132, t. 15 (1803). Based upon
O. hispidum Michx. |= O. virginicum L.| and O. molle Michx.

Osmodium Raf. Med. Repos. N. Y. 5: 353 (1808); Merrill, Index Raf. 203
(1949). A substitute name for Onosmodium Michx.

Purshia Spreng. Anleit. ed. 2, 2: 450 (1817); Lehm., Asperif. 2: 382 (1818).
A substitute name for Onosmodium Michx.

Osmidium Walp. Ann. 3: 134 (1853). Apparently a printers’ mistake for
Onosmodium Michx.

Plants perennial, with few to many frequently coarse, erect stems, strigose
or more commonly hispid, glanduliferous in one species only, simple or
bearing ascending leafy fertile branches above the middle. Leaves mostly
cauline, numerous, bearing several pairs of strong assurgent veins; lowest
leaves usually larger and more elongate than the middle and upper ones,
usually shed before the time of flowering, persisting in only a few spécies.
Cymes scorpioid, single or paired, borne terminal on the main stem and
its branches, usually many-flowered, at first dense and coiled and later in
the fruiting condition becoming straight, greatly elongate, and loosely
flowered. Bracts numerous, one adjacent to each flower, lanceolate to

y

lobes attenuate or broadly linear or slightly spathulate, acute to obtuse
or rounded at the apex, usually evidently unequal, after maturity of the

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVI 19

fruit usually disarticulating at the base. Pedicels short to elongate. Corolla
opening (and its style exserted and its anthers mature) while incompletely
developed. Mature completely developed corolla white to cream or yellow,
usually with the lobes more or less green or greenish, subtubular, two to
three times as long as broad, regular, straight; outside hairy, usually
strigose or with appressed hairs; inside glabrous (or a few hairs near the
tip of the lobes in one species), without any glands or true faucal append-
ages; annulus evident, consisting of a narrow flange or ten lobules, gla-
brous; throat usually about twice the thickness of the corolla-base, broadest
(and in lateral silhouette somewhat angulate) a short distance below its
summit, bearing ten externally protrudent gibbose convexities, five below
the base of the corolla-sinus and five lesser ones below the corolla-lobes.
Corolla-lobes nearly one half to nearly one fifth (but most commonly
about one third) the total length of the corolla, erect, more or less elongate,
very narrowly imbricate in the bud, cuneate or triangular or ovate-triangu-
lar with the apex acute or acute with an attenuate tip; sinus between the
lobes narrow, acute, thickened and inflexed at the very base. Stamens
borne at equal heights in the thickest part of the throat, affixed at or dis-
tinctly above the middle of the corolla; filaments short, usually somewhat
unguiculate, one third the length of the anthers or less; anthers oblong-
lanceolate, with their tips reaching up to or slightly beyond the base of
the corolla-sinus, affixed between the base and the middle usually at about
one third of its total length above its base; apex of anther bearing a
minute appendage (appendage semicircular or quadrate or three-lobed or
sometimes attenuate, composed of prolongations of the thecae and the nar-
row connective, with the portion prolonged from the connective frequently
darkened and perhaps glandular); base of anther emarginate or lobed,
reaching downward in the corolla-throat to or distinctly below the point of
attachment of the filaments; connective narrow, glabrous; thecae below
the anther attachment distinct and usually spreading, their basal tips
very obscurely if at all thickened. Pollen small, 16-24 * 13-22 p, ovoid
to globose-ovoid, lower half rounded more broadly and evenly than the
upper half, always longer than broad, though at times only slightly so;
pores inframedial, six or seven in a single row, usually obscure; grains in
polar profile circular or rarely somewhat polygonal. Style filiform, pre-
cociously long-exserted, emerging from the incompletely developed corolla
when the latter has not yet attained half of its eventual size; stigmas two,
minute, terminal on the style. Ovules four but usually only one maturing
and very rarely more than two. Nutlets ovoid or ellipsoidal, white or
tawny, lustrous, smooth or pitted, gradually narrowed to the base or some-
times with a suprabasal constriction, nearly circular in transverse section,
lacking a prominent ventral keel or any evidence of a ventral suture;
attachment-scar flat, basal, nearly as broad as long, towards ventral edge
bearing the slightly prominent broken end of the bony tubular funicular
canal and towards the dorsal edge a small, attenuate, obliquely ascending
prominence representing a thickening of scar tissue about the base of a
vascular strand leading to the dorsum of the nutlet body. Gynobase de-

20 JOURNAL OF THE ARNOLD ARBORETUM __ [VOL. xxxv

pressed, nearly flat, attachment surface plane, distinct, lacking strongly
thickened upturned margins.

A genus confined to middle and eastern sections of the United States
and to northeastern Mexico. Most authors have recognized seven species
within the United States, cf. Mackenzie, Bull. Torr. Bot. Cl. 32: 495-506
(1905); Johnston, Contr. Gray Herb. 70: 17-18 (1924); and Fernald,
Manual Bot. 1200-1201 (1951). Two additional species are endemic to
northeastern Mexico. Of the total only four are sharply definable and
always positively recognizable, viz., O. unicum Macbr. and O. dodrantale
Johnston of Mexico, O. Helleri Small of Texas, and O. virginianum (L.)
DC. of the eastern border of the United States. The other five “species”
intergrade and are incapable of sharp definition. There is O. hispidissimum
Mack., ranging in the Ohio Valley and the Middle Atlantic States, O. occi-
dentale Mack., widespread in the region between the Mississippi River
and the Rocky Mountains, and O. bejariense DC., confined to Texas.
These three are habitally similar and obviously related, and over the
larger portion of the area in which they are found they seem relatively
constant in their characters. Another group of related plants is practically
confined to Tennessee, Kentucky, Arkansas and Missouri. These have
slender stems and an indument usually closely appressed and frequently
scanty. Representative of them are O. molle Michx. and O. subsetosum
Mack. Embarrassing transitional forms between these two groups, as
well as between their members, appear to be frequent in Missouri, Iowa,
and Illinois, where the ranges of the various species converge or even
overlap. Analysis will probably reveal evidence of much hybridization
and intragression affecting O. Aispidissimum, O. occidentale, and O. molle
in that area

The genus has general relations with Macromeria and Lasiarrhenum
but is more closely related to the American and particularly to the Mexi-
can species of Lithospermum and, along with the two genera mentioned,
is probably derived from them. It is a sharply defined and thoroughly
natural group readily recognizable by a number of strong characters. Par-
ticularly noteworthy are its precociously sexual flowers. In these the an-
thers mature and the style becomes long exserted in corollas that have
opened before they have attained half their eventual size. The subtubular
corolla is also distinctive in having very narrowly imbricate, erect, sharply
acute or acuminate corolla-lobes and sinuses between the lobes which are
distinctively thickened and inflexed at the very base. The corolla-throat
may have five to ten gibbose swellings outside, but inside is devoid not
only of faucal appendages but also of any hairs or glands. The included
glabrous anthers are attached between base and middle to very abbreviated
filaments and are appendiculate at the apex and lobulate at the base.

Most of the species commonly referred to Onosmodium are obviously
congeneric. To be excluded, however, are O. revolutum (Robins.) Macbr.,
Contr. Gray Herb. 49: 21 (1917), the type of the genus Psilolaemus, and
Onosmodium strigosum (HBK.) Don, the type of the genus Lasiarrhenum.
Gray, Synop. Fl. Am. 2': 205 (1878), and later Macbride, Contr. Gray

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVI 21

Herb. 49: 19 (1917), have suggested that all species of Macromeria ex-
cept M. exserta should be accommodated in an enlarged Onosmodium.
Justification for this apparently rests solely on general vegetative similari-
ties and in the fact that corolla-lobes of Macromeria may be acute. In
size, form, and behavior of the corolla and in structures inside the corolla
the species of Onosmodium are extremely different from those of Macro-
meria. Indeed, the two genera differ more from each other than they do
individually from Lithospermum. If generic values are not to be lowered,
Onosmodium and Macromeria must be kept distinct.

In the Mexican O. dodrantale, the plant not only has very short stems,
1-3 dm. long, but also has the leaves smallest at the base of the stem
and gradually increasing in size upward along it. In the other species of
the genus, all of them with much longer stems (usually 5-12 dm.), the
leaves at the base of the stem are larger and more elongate than those
on its middle sections. The indications are that the stems elongate from
the center of a winter rosette of leaves and not directly from a bud on the
caudex, as in O. dodrantale and in most species of Lithospermum., ‘The
large lowermost leaves have usually fallen away in herbarium specimens
of most species. Indeed, only in those of O. bejariense and O. Hellert are
they commonly found dried up and still persisting in some numbers
crowded at the base of the stem. In most species the foliage has reached
full size and has become firm before anthesis. This is not the case, how-
ever, in O. Helleri. In that species the plant flowers while it is still growing
vigorously and before it has attained full stature and the upper leaves
have attained their maximum size and mature firmness. Plants of O. Helleri
at anthesis and those maturing fruit have a very different appearance. In
fruiting plants of this species the bracts become unusually large and con-
spicuous.

The cymes in O. dodrantale are weakly developed and hardly more than
three- to six-flowered bracteate glomerules terminating the short stems. In
the other species of the genus, however, they are distinctly scorpioid and
abundantly flowered and eventually become straight, extremely elongate,
and very loose at extreme maturity. The flowers maturing their corollas
are borne crowded on the arched summit of the cyme, those with com-
pletely grown corollas just above the point where the rachis begins to
straighten. Since the pedicels are strict, the corollas incline backwards
towards the summit of the cyme, and those near its summit become nearly
horizontal. After the corolla is shed the rachis straightens, its internodes
elongate, and the abundant bracts, previously inconspicuous, increase in
size and become very conspicuous.

The indument on the herbage may be either strigose or bristly. Onos-
modium subsetosum is unusual in having the stems glabrous or nearly so.
A very distinctive feature of the Mexican O. unicum is the presence of very
slender multicellular, gland-tipped hairs intermixed among the stiff ascend-
ing hairs on the leaves, stems and rachis of the cyme and even on the calyx.
I know of no other American herbaceous borage having generally dis-
tributed hairs of this type.

22 JOURNAL OF THE ARNOLD ARBORETUM _ [VOL. xxxv

The flowers of Onosmodium are precociously sexual. The corolla opens
and its anthers are matured and its style is exserted long before the corolla
has attained full size, commonly when it is less than half its eventual
size or even sufficiently elongate to surpass the calyx. In this behavior of
the corolla the genus differs from all other American Boraginaceae, and
among the Lithospermeae, at least, has a parallel only in the monotypic
Halacsya of Albania and Montenegro, cf. Jour. Arnold Arb. 34: 276
(1953). On the densely flowered arched summit of the scorpioid cyme in
Onosmodium, as in Halacsya, the styles are to be seen projecting not merely
from the fully developed conspicuous corollas but also above the latter
from a series of gradually less well developed corollas on younger and
higher parts of the cyme. The style may first emerge when the corolla
is so small as to be overtopped by its calyx lobes. When the corolla is
sufficiently large to equal or slightly surpass the calyx, the style is long
exserted, the corolla-lobes have unfolded completely, and the anthers have
attained nearly full size and in herbarium specimens are dehiscent. At
this stage, above the level of the filament attachments the corolla is ap-
proaching mature form and has attained about seventy-five per cent of its
eventual size. Below the level of the filament attachments, however,
growth has lagged, for the tube is still very short and scarcely developed.
It is after the corolla opens and the anthers and style are exposed and func-
tioning that the tube elongates and increases in diameter and the corolla
achieves full size and mature proportions. The early emergent, eventually
long exserted style evidently makes self-pollination in this genus prac-
tically impossible. How the corolla may function in pollination during
the period in which it doubles its size after opening and maturing its anthers
is a subject deserving investigation.

The fully developed corolla in this genus may be 8-12 mm. long, as in
O. virginianum and O. hispidissimum, or as much as 12-18 mm., as in
O. occidentale. From a base 1.5—2.5 mm. thick they gradually expand
upwards to just below the summit of the tube where they are broadest,
ca. 4 mm. in O. virginianum and 6—7 mm. in most other species. The lobes
are a third to a fifth (usually about a third) the total length of the corolla.
They are cuneate to triangular or ovate-triangular, and from the base or
just above it contract with rie nla sides to the sharp apex,
which is acute or sometimes (in O. cum) acute with an attenuate tip.
The lobes, except near the very ae (i.e., on all surfaces exposed while
folded in the bud), have their outer face hairy, as elsewhere on the outside
of the corolla. The inner face of the lobes is usually completely glabrous. In-
deed, the only exception is in the corolla of O. dodrantale, which is unique
in having the inner surface of its lobes always inconspicuously hairy just
below their tip. In the bud the lobes are imbricate very narrowly and
only sufficiently to produce a very narrow glabrous margin, narrowing
upward, on the dorsum of each lobe. It is this very narrow imbrication
of the lobes, becoming negligible towards the tip of the bud, that facilitates
the early egress of the precociously elongating style that has pushed
upwards inside the very immature flower bu

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVI 23

The base of the acute sinus between the erect corolla-lobes is distinctly
plicate, inflexed, and thickened. A somewhat similar condition occurs
in the flowers of some species of Heliotropium, but among the Litho-
spermeae in no other genus save Onosmodium is it present or at least so
well developed. In most species the corolla is locally distended outwardly by
convex gibbose swellings, one directly below the plicate base of each sinus.
These gibbosities make the corolla more or less five-angled just below the
summit of the throat where it is broadest. Gibbosities of a similar sort
may also be present below the base of each corolla-lobe and alternate with
those below the sinus, but are usually less conspicuous than the latter.
Inside the corolla-throat, except for the inflexed bases of the sinus, there
are no intrusive appendages or invaginations. Faucal appendages and
stipitate glands, common features in the corolla-throat of other Litho-
spermeae, are completely absent. Pubescence of any sort is also absent
inside the corolla, even on the annulus. The annulus is clearly developed,
either as a narrow thickish continuous flange or as ten lobes.

The anthers, 2.5—-3.5 mm. long, are attached to a very short filament at
about a third of their length above their base, and hence very definitely
below their middle. The filaments, a third the length of the anther or less,
are attached in the corolla commonly about 2 mm. below the summit of
the throat. The anthers, accordingly, usually reach up to about the base
of the sinus above them or at most have their tips only a millimeter beyond.
The base of the anther projects downward to the same level as the attach-
ment of the filament on the walls of the throat or slightly below it. In the
lower quarter of the anther the thecae are not united. They may remain
parallel and the base of the anther appear emarginate, but usually, par-
ticularly in age, they tend to spread, making the anther lobulate at the
base. The thecae are only slightly narrowed above the middle. The anther
is always minutely appendiculate at the apex. The very diminutive ap-
pendages appear to be formed not merely by a prolongation of the narrow
connective but also by apical prolongations of each theca. These parts
may be confluent or distinct. The appendage is variable in form within
the species, and even according to the age of the anther bearing it. It may
be three-pronged, it may be quadrate with either truncate or toothed
summit, or occasionally, in O. dodrantale and O. unicum, subulate or
lanceolate. In O. virginianum the anther summit may be even broadly
emarginate and the thecae each tipped by a minute appendage and distinct
from the third small appendage arising from the depth of the sinus
between them.

The pollen of the various species of Onosmodium differs slightly in size
but very little in shape. It is ovoid, or at times, even in the same species,
very broadly so and even globose-ovoid in form. The grains are always
perceptibly longer than broad with the upper and lower half differing to
some degree in outline. In lateral profile the pores are at most very
weakly protrudent. Usually they are very obscure. They are borne in
a single row perceptibly below the middle of the grain and are usually
six but not infrequently seven in number. In polar profile the grains are

24 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXXV

almost invariably circular in outline. The size of the grain seems to be
roughly correlated with corolla-size. Species with large flowers usually
have grains larger than those in species with small flowers. Small-flowered
O. virginianum produces the smallest pollen, 15-18 & 13-16 p». In O.
subsetosum, O. molle, O. hispidissimum, and O. Helleri the grains measure
16-18 & 15-16 p, in O. unicum, 16-22 15-18 pn, in O. occidentale,
18-22 « 16-20 p, in O. bejariense, 20-22 16-20 p, and in O. dodrantale,
22-24 X 20-27 u.

As in the case of all genera of the Lithospermeae previously examined,
pollen production in Onosmodium was found to be prevailingly normal and
abundant, with imperfect pollen very scanty and infertile anthers rare.
Worthy of note, therefore, is the fact that the reverse condition seems
prevalent in O. occidentale over the northern parts of its range, in the
northern Plains and adjacent Rockies. The species deserves cytological
examination.

The nutlets of the genus are generally similar to those of Lithospermum.
Unlike those of Macromeria their ventral suture is always closed and com-
pletely obliterated. There is no appreciable ventral keel. Although the
ovary is four-ovulate, flowers of the genus seldom mature more than a
single nutlet. Only in O. occidentale are the exceptional fruits, frequently
present in limited numbers, found to be maturing two nutlets. In that
species fruit with three nutlets may be encountered very rarely, but none
with four nutlets have been found. Accordingly I have not seen a sym-
metrically developed gynobase. In those maturing one to three nutlets,
the attachment faces are nearly horizontal and plane or nearly so. The
faces are not concave, nor do they have strongly thickened upturned
margins as in Macromeria. The attachment scar on the nutlet is flat, not
convex as in Macromeria, but, like the scar in that genus, it does bear a
projecting end of the broken tubular funicular canal, and also another
more dorsal projection in the form of an obliquely ascending protuberance.

3. Nomosa, gen. nov. Lithospermeae.

Planta perennis herbacea strigosa. Caules hornotini erecti simplices
foliosi ut videtur e caudice ex caulibus vetustis 3-4 mm. crassis procum-
bentibus laxe ramosis composito orientes. Folia breviter strigosa cinerea vel
plus minusve argentacea supra basim triplinervia. Folia basalia oblanceolata
in fasciculos steriles aggregata. Folia caulina numerosa sessilia a basi
caulis sursum gradatim majora, superiora anguste oblonga vel lanceolato-
oblonga apice acuta vel plus minusve obtusa. Cymae terminales geminatae
laxe scorpioideae pauciflorae tandem rectae racemosae. Bracteae numerosae
foliaceae haud conspicuae calycem haud superantes. Flores sub anthesi
in parte curvato supremo symae gesti erecti vel horizontales vel declinati.
Calyx 5-fidus; lobis evidenter inaequalibus firmis costatis elongatis lin-
earibus vel cuneatis. Pedicelli crassiusculi stricti modice elongati. Corolla
ut videtur alba regularis crasse tubularis, calyce subduplo longior, a basi sur-
sum lente gradatimque ampliata (haud faucibus distinctis donata) in tertia
parte superiore crassissima, triplo longior quam lata, extus dense brev-

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVI rae)

iterque strigosa, intus inter basis decurrentis filamentorum pilis gracilibus
glanduliferis praedita alibi glabra; faucibus nec appendiculas nec glandulas
stipitatas proferentibus; lobis imbricatis parvis quam longitudine corollae
10-plo brevioribus erectis late ovatis infra medium latissimis basi ali-
quantum contractis tam longis quam latis vel paullo latioribus quam
longis; annulo nullo. Filamenta angusta elongata sursum gradatim at-
tenuata crassiuscula, in quarta parte inferiore corollae affixa, supra medium
teretia et glabra, infra medium plus minusve compressa et pilis gracilibus
glanduliferis abundanter donata, basi breviter decurrentia incrassata pilis
glanduliferis vestita. Antherae elongatae lanceolatae, supra basim affixae,
filamentis duplo breviores, in parte suprema tubi corollae gestae, basim
versus latissimae deinde sursum gradatim angustatae, appendiculis steril-
ibus pallidis gradatim attenuatis e tubo corollae exsertis sed lobos corollae
haud superantibus terminatae, dorso connectivum pallidum laeve latum
(latitudine quam antheram triplo angustius) pilis paucis (1-4) rigidis
valde adpressis praeditum proferentes; thecis basim versus distinctis sed
parallelis infra medium apicem versus angustatis, basi imo acutiusculis
apiculo inconspicuo incrassato donatis sed nullo modo appendiculatis.
Granulae pollinis late ovoideae 25-28 & 21-23 » infra medium latissimae
et ibi 7 vel 8 poris obscuris instructae, a latere visae late ovatae ca. 8 mm.
supra basim semicircularem latiores deinde sursum per margines rectos
convergentes in apicem latum abrupte rotundum contractae. Stylus
gracilis glaber tandem evidenter exsertus; stigmatibus 2 distinctis termi-
nalibus. Nuculae ignotae.— Nomen a Onosma litteris interversis de-
sumptum.

Nomosa Rosei, sp. nov.

Planta 25-30 cm. alta; caulibus erectis basim versus ad 3 mm. crassis
pilis rectis antrorsis laxe adpressis ad 1 mm. longis vestitis, internodiis
brevibus ca. 5 mm. longis; foliis strigosis (pilis valde adpressis rectis
0.2-0.6 mm. longis), margine inconspicue ciliolatis (pilis strictis vel ad-
scendentibus 0.5-1 mm. longis), costa 10-20 mm. supra basim nervos 2
assurgentes validos conspicuos perelongatos proferenti donatis; foliis bas-
alibus 40-60 mm. longis supra medium 8-12 mm. latis; foliis caulinis
superioribus 40-55 mm. longis medium versus 8-10 mm. latis; cymis ca.
10-floris maturitate rectis ad 8 cm. longis; lobis calycis strigosis margine
hispido-ciliatis, sub anthesi 8-11 mm. longis tandem 10-15 mm. longis,
lobo majore 1—3 mm, lato quam lobis minoribus 3-5 mm. longiore; pedi-
cello sub anthesi 3-5 mm. longo tandem ad 8 mm. longo recto; corolla

crassa; lobis corollae 2 mm. longis medium versus 2—2.5 mm. latis,. basi
ima 2 mm. latis; filamentis 9.5-10 mm. longis, 5 mm. supra basim corollae
orientibus, basim versus 1 mm. latis, apicem versus ad 0.5 mm. lIatis,
infra medium pilos 0.5 mm. longos proferentibus; basi decurrente filamenti
incrassato 2-3 mm. longo ca. 0.8 mm. lato pilis glanduliferis vestito;
antheris 5 mm. longis ca. 1.4 mm. supra basim affixis, thecis 3.5 mm.

26 JOURNAL OF THE ARNOLD ARBORETUM __ [voL. xxxv

longis basi 3—3.5 mm. infra basim loborum corollae gestis; appendiculo
terminali antherae 1.5 mm. longo imam ad basim ca. 0.5 mm. lato; stylo
23-25 mm. longo.

MEXICO: in Sierra Madre near the southern border of the state of Durango,
Aug 16, 1897, J. N. Rose 2360 (type, Gray Herb.).

This remarkable plant is known only from a collection made over a half-
century ago by J. N. Rose during his first expedition to Mexico. In general
appearance it mimics Onosma to a remarkable degree. The original collec-
tion was distributed with locality data given merely as “Durango.” Ac-
cepted as a member of the large and diverse Old World genus Onosma,
and believed to have been introduced into Mexico, presumably at Durango
City, the plant attracted no careful study, particularly since no plant
similar to it was subsequently found in America. Only recently, when it
was dissected and carefully compared during a study of Onosma and
related genera, were the very many distinctive features of this neglected
Mexican plant fully recognized. Upon investigation it was found that the
specimens were made not at Durango City, but rather in the Sierra Madre,
ca. lat. 22° 15’ N., at the extreme southern end of the state of Durango,
in a wild and infrequented area. It was discovered by Rose when on
horseback and with pack-animals he journeyed through Nayarit across
the southern tip of Durango and on eastward into Zacatecas, crossing the
Sierra Madre in a section not subsequently visited by a botanist. Our
plant is apparently endemic in the mountains of this particular region.

Although extremely suggestive of Onosma in external features, Nomosa
is actually a very close ally of the Mexican Lasiarrhenum. Its affinities
are with such American genera as Lasiarrhenum, Macromeria, and Onos-
modium, which appear to be derivatives of American Lithospermum rather
than of the strictly Old World Onosma. Along with Lasiarrhenum, No-
mosa is distinguished from Onosma by its 7—8-porate pollen, unequal
calyx-lobes, triple-nerved leaves, and free anthers with hairy connectives.
From Lasiarrhenum it differs in having very elongate gradually narrowed
filaments which are attached very low in the corolla and, below the middle,
are densely clothed with an abundance of slender elongate multicellular
gland-tipped hairs. The anthers, only half as long as the filaments, are
carried higher in the corolla and have their elongate terminal appendage
exserted from the mouth of the corolla. Unlike the anthers of Lasiar-
rhenum, which are abundantly and conspicuously hairy on the back, those
of Nomosa bear only a very few closely appressed, much shorter incon-
spicuous hairs on the connective. The corolla of Nomosa, coarsely tubular,
does not swell into a campanulate throat as in Lasiarrhenum. Its lobes
are ovate rather than deltoid, and the inner surface of the throat bears no
hairs nor stiped glands. The annulus, very well developed in Lastarrhenum,
in Nomosa is absent or obscure.

The plant is not bristly. The foliage is closely strigose with the minute
hairs abundant but not extremely crowded nor overlapping. The thin
indument is tidy and smooth and gray or somewhat silvery. The leaves

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVI 27

usually have only two strong elongate veins. These arise from the midrib
1—2 cm, above the leaf-base. They are assurgent and nearly as strong as
the midrib and are frequently prolonged almost to the leaf-tip. Occasion-
ally less well developed veins may also be present, but these are never as
conspicuous as the major veins. The corolla, if not perfectly regular,
departs from that condition only by having its two adaxial lobes perhaps
very slightly larger than the other three. The lobes are involutely curved,
and the upper half of the tube directly below each of them is somewhat
swollen. The upper half of the corolla, accordingly, has five weakly inflated
ribs. The filaments are attached unusually low in the corolla. The posi-
tion of their thickened short decurrent base is marked on the outside of
the corolla by five oblong-elliptic glabrous areas extending 2.5—-5 mm.
above the corolla-base. The slender, elongate, gradually narrowed fila-
ments are glabrous above the middle, but below the middle and on their
thickened decurrent base they bear multicellular gland-tipped hairs in
great abundance. This shaggy indument on the filaments is a distinctive
feature of Nomosa. Some of the multicellular gland-tipped hairs occur also
on the walls of the corolla immediately adjacent to the stamen bases. Else-
where the inner surface of the corolla bears no hairs or stipitate glands.
The anthers are carried high enough in the corolla-tube to have their ter-
minal appendage exserted from the corolla-mouth. From below the middle
the anthers narrow towards an attenuate tip. The terminal 1.5 mm. of
the anther, its appendage, consists of a sterile prolongation of the con-
nective. This is narrowed to a slender point and is frequently curved to
one side. It is the only part of the anther exserted from the corolla-tube.
Although exserted, it does not become conspicuous, since it remains hidden
behind the erect corolla-lobes which overtop it. The back of the anther
has a broad, smooth, weakly convex connective which bears a very few
stout closely appressed hairs. These hairs, unlike those on the anther in
Lasiarrhenum, are few and inconspicuous and must be looked for under
the microscope. At the base of the anther the thecae, though distinct for
0.6 mm. above the base, are not spreading but parallel. The lower end of
each theca is acutish. Its tip never becomes appendaged or distinctly
thickened as in Onosma. The style emerges from the fully developed
corolla as its lobes unfold and soon bears its two stigmas 5—6 mm. beyond
the tips of the erect corolla-lobes. Fruit of the plant is unknown. I am con-
fident, however, that the nutlets will prove to be very similar to those of
Lasiarrhenum.

4. Lasiarrhenum Johnston, Contr. Gray Herb. 70: 15 (1924). Type
species Onosma strigosum HBK,

Plant perennial, prevailingly strigose, with spreading hairs only on the
stem and veins of the leaf. Stems coarse, erect, several or more, simple or
bearing ascending leafy floriferous branches above the middle. Leaves all
cauline, numerous and usually crowded, elongate, triple-ribbed, narrowly
lanceolate or the lower ones oblanceolate, those near the middle of the
stem usually largest, those near the base of the stem small and even

28 JOURNAL OF THE ARNOLD ARBORETUM _ [VoL. xxxv

imperfectly developed. Cymes scorpioid, always terminal, geminate on the
main stem but usually single on the branches, in age straightening and
elongating but remaining moderately crowded. Bracts numerous, linear
to lanceolate, ascending, not very conspicuous even in fruiting inflores-
cences, scarcely if at all overtopping the adjacent fruiting calyx. Calyx
5-fid; lobes unequal with the abaxial one largest, acute, usually slightly
more than half the length of the corolla, moderately accrescent in fruit;
pedicel slender, straight, strictly ascending, half as long to as long as the
calyx. Corolla white, regular, below the middle coarsely tubular and above
the middle swelling to form a somewhat campanulate throat, the outer
surface strigose; lobes erect, relatively small, more or less deltoid with the
apex frequently rounded, as broad or broader than long, imbricate; throat
on inner surface bearing scattered stipitate glands, frequently inconspicu-
ously strigose along lines below the corolla-lobes, without faucal append-
ages; the annulus well developed, glabrous, a thick ridge or narrow collar,
obscurely lobed. Filaments affixed at the base of the throat, nearly as
long as the anther, thickish and somewhat fleshy, broad, dorsiventrally
compressed, narrowly obovate to oblanceolate, usually bearing some stiped
glands especially below the middle, but otherwise glabrous, below the at-
tachment narrowly decurrent and forming thickened ridges on the walls
of the corolla-tube. Anthers lanceolate, deeply included in the corolla-
throat, attached distinctly below the middle, appendaged at the apex;
terminal appendage evident, narrow, attenuate, compressed, base as broad
as the connective of which it is an attenuate prolongation, with a length
not exceeding and usually less than the maximum width of the dehiscent
anther; base of anther lobed; thecae narrowing towards their apex, free
for a short distance above the base, parallel or somewhat spreading at the
lower end, the basal end usually acutish with a thickened tip; venter of
anther glabrous, bearing the thecae closely juxtaposed, with the connec-
tive exposed only at the apex; dorsum of the anther with a broad connec-
tive, conspicuously hispid-villose both on the connective and on the back
of the thecae, the hairs slender, white, loosely and antrorsely appressed.
Pollen broadly but distinctly ovoid, 20-23 16-20 ,, broadest and bear-
ing the pores about 8 » above the broadly rounded base; lateral profile
ovate, above the pores with the converging sides nearly straight and the
apex abruptly rounded; polar profile circular; pores 7 or 8, usually obscure.
Style emerging from the well-developed corolla-bud as the lobes unfold,
shortly but distinctly surpassing the erect corolla-lobes; stigmas evidently
two, diverging from the summit of the style. Nutlets smooth, white, lus-
trous, ovoid, narrowed at the base, all four frequently developing, diver-
gent, on the venter bearing a low, rounded, weakly developed keel but no
evidences of a ventral suture, dorsum convex; attachment scar basal,
horizontal, nearly as broad as long. Gynobase broadly pyramidal, sur-
mounted by the thickened persisting 4-angulate base of the style; attach-
ment faces sloping, distinct, with thickened margins.

A very well marked genus with a single species, endemic to south central
Mexico (Michoacan to Puebla and adjacent Jalisco to Oaxaca).

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVI 29

Lasiarrhenum strigosum (HBK.) Johnston, Contr. Gray Herb. 70: 15
(1924).

Onosma strigosum HBK. Nov. Gen. et Sp. 3: 93 (1818).

Onosmodium strigosum (HBK.) Don, Gen. Syst. 4: 317 (1837).

Onosma trinervium Lehm. Asperif. 2: 378 (1818) and Icones 1: 11, t. 9 (1821).
Lithospermum longifolium Willd. in R. & S. Syst. 4: 742 (1819).

Plant 5—10 dm. tall; stems abundantly short hispid with hairs 1-2 (—3)
mm. long. Leaves at middle of stem 5—10 cm. long, 6-22 (usually 8-15)
mm, broad, paler beneath, triple-ribbed, the strong midrib producing
(about 10 mm. above its base) two evident strong, greatly prolonged
assurgent veins, the veins nearly as strong as the midrib and extending
almost to the leaf tip, evident on both sides of the leaf, hairs on the upper
surface of the leaf provided with minute discoid bases but these seldom
conspicuous. Corolla 15-23 mm. long, below the middle 3—4.5 mm. thick
and subcylindric or slightly ampliate, at or slightly below the middle ex-
panding into a campanulate throat 6-12 mm. broad at the top; corolla-
lobes 3.5—5 mm, broad at the base, 2-3 mm. long. Filaments 3-5 mm.
long, from the narrow attachment gradually expanding to become 1.3—1.5
mm, broad between the middle and the acute apex, arising 5-10 mm.
above the corolla-base. Anthers 4—5.5 mm. long, 1-1.3 mm. broad below
the middle, affixed 1.2-1.5 mm. above the base, reaching up to 1-3
(usually 2) mm. below the base of the corolla-sinus; terminal appendage
0.4-1 mm. long and 0.2-0.3 mm. wide at the base; base of anther
held 1.5-3 mm. above the base of the filament; thecae free 0.5-1 mm.
above the anther-base; dorsum of anther bearing hairs 0.6—-1.3 mm. long,
connective one third of the width of the anther. Style 16-23 mm. long,
surpassing the corolla-lobes 1-5 mm. Nutlets ovoid, usually ca. 3 mm.
long and 2.5 mm. thick, in transverse section with the ventral side broadly
obtuse but elsewhere nearly circular in outline.

This monotype is particularly notable for its broad, fleshy, usually
oblanceolate filaments and for its large lanceolate appendaged anthers
which are coarsely and abundantly white-hairy on the back. In gross
habit the plant is very suggestive of Onosmodium, a genus with which it
was long confused. In floral structure, however, it is extremely different.
The closest relations of Lasiarrhenum are with Nomosa. Details concerning
this relationship are treated in the discussion of the latter genus.

The anthers of Lasiarrhenum have a somewhat less well developed
terminal appendage than those of Nomosa, but otherwise are very similar
as to form and size. Unlike the anthers of Nomosa, which bear only a few
inconspicuous hairs on the connective, the whole of the dorsal surface of
the anthers in Lasiarrhenum bears slender ascending white hairs in con-
spicuous abundance. The presence of hairs on the back of the anther is
no common feature. Among American Lithospermeae they occur only in
Lasiarrhenum, Nomosa, and one species of Macromeria, M. viridiflora DC.

The terminal appendage on the anther of Lasiarrhenum (and Nomosa)
is very suggestive of that in Onosma, and because of this the possibility

30 JOURNAL OF THE ARNOLD ARBORETUM _ [vot. xxxv

has been recognized that our plant might have close affinity with that
genus of the Old World. In structures other than the anthers, however,
Lasiarrhenum and Nomosa have much more in common with Onosmodium
and Macromeria The indications are that all these four genera are Ameri-
can derivatives of either Lithospermum or Lithospermum-like ancestors
and so probably have no direct affinity with Onosma. The well-developed
anther-appendages of Lasiarrhenum and Nomosa appear to be simply the
extreme expression of the same tendency towards prolongation of the
connective that is to be observed less well expressed in Onosmodium and
Macromeria. In Onosmodium the connective is narrow, and the maximum
prolongation of it, as in O. unicum Macbr. and O. dodrantale Johnston,
is slender, weak, and at most 0.5 mm. long. The connective is also narrow
in most species of Macromeria. In the one species of that genus, M.
viridiflora DC., in which the connective approaches in width that of
Lasiarrhenum and Nomosa, its prolongation is very short and stout. In
Macromeria the thecae are equally broad from base to tip, and the anthers,
accordingly, are more or less distinctly oblong in outline and have a broad
summit. In Onosmodium the thecae are slightly narrowed above the
middle and usually have an apiculate tip. The anther tends to have a
lanceolate outline. In Lasiarrhenum and Nomosa the thecae are gradually
narrowed to the apex. The anther as a whole is narrowed to the width of
the broad connective at the level of the anther-tips, and the attenuate
prolongation of the connective provides the pointed tip and thus completes
the lanceolate outline of the whole anther.

5. Perittostema, gen. nov.

Herba parva perennis. Caules graciles erecti simplices strigosi foliosis-
simi. Folia gracilia lineari-subulata medio-costata sed enervata margine
valde revoluta. Cymae scorpioideae solitariae glomeratae terminales. Brac-
teae lineares calyce breviores inconspicuae. Calyx 5-fidus; lobis ut videtur
paullo inaequalibus cuneatis corolla plus quam duplo longioribus. Corolla
lutea extus strigosa; tubo infra medium cylindraceo, supra medium in
fauces subinflatos differentiato summum ad apicem (i.e., infra basim
loborum corollae) paullo constricto; lobis tubi (faucibus inclusis) 5-plo
brevioribus stricte adscendentibus paullo longioribus quam latis rotundis,
in alabastro late imbricatis; faucibus quam parte inferiore cylindrico tubi
subduplo crassioribus aequilongis, intus medium versus appendiculas in-
vaginatas arcuatas glandulis stipitatis obsitas gerentibus; annulo. vix
elevato adpresse villuloso. Stamina in faucibus profunde inclusa. Filamenta
basi faucium affixa, valde compressa, medio-costata, a basi angusta sursum
valde gradatimque expansa, apicem versus latissima deinde abruptissime
contracta, obovata vel deltoidea, subduplo longiora quam lata. Antherae
oblongo-ellipticae filamento subduplo longiores paullo infra medium affixae
apiculo attenuato recurvato terminatae basi bilobatae; thecis atro-margi-
natis quartem partem inferiorem distinctis et ibi sinu acuto separatis; con-
nectivo late glabro laevi. Granulae pollinis ellipsoideae 24-25 20 p, a
latere visae ellipticae lateris angulatae, desuper visae circulares: poris

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVI 31

aequatorialibus 6-8 sed saepissime 7. Stylus gracilis lobos stricte adscend-
ents corollae superans; stigmate terminali parvo obscure bilobo. Fructu
ignoto.— Nomen derivatur a zepirros, insolitus, et ornya, stamen, quia
filamenta formam anomalem habent.

Perittostema pinetorum (Johnston), comb. nov.
Lasiarrhenum pinetorum Johnston, Jour. Arnold Arb. 16: 187 (1935).

Stems 10-15 cm. long, 2.5 mm. thick towards the base, internodes very
short; leaves very numerous, 1-2 mm. broad above the base and very
gradually narrowed towards the apex, linear-subulate, 10-30 mm. long,
becoming gradually smaller upwards along the stem, bearing appressed
stiff pallid hairs 0.3-0.5 mm, long, on lower surface all but the midrib
hidden by the inrolling of the strongly revolute leaf-margins; cymes 3—7-
flowered; calyx 4-4.5 mm. long, lobes ca. 0.7 mm. wide at the base,
gradually narrowed; pedicel about 1 mm. long; corolla ca. 10 mm. long;
tubular portion of corolla ca. 8 mm. long, below the middle cylindric and
2-2.5 mm. thick, above the middle expanding to form a swollen throat
which becomes 3-4 mm. broad near its middle and then contracts to be-
come 2-3 mm. thick at the summit; veins in the cylindric tube rather
prominent; corolla-lobes ca. 2 mm. long and 1.8 mm. broad, rounded;
faucal appendages arcuate, ca. 1 mm. broad, prominent, thickened and
weakly invaginate, bearing some stipitate glands, included, borne 1.5 mm.
below the constricted summit of the throat; anthers ca. 2 mm. long and
1 mm. broad, obtusish summit bearing a strongly compressed narrowly
ligulate and strongly recurving terminal appendage which is 0.2 mm.
long and nearly 0.1 mm. broad at the base; thecae free for 0.4-0.5
mm, above their base, in age somewhat spreading to form an open acute
sinus at the base of the anther; base of anther held ca. 0.3 mm. above the
base of the filament; apex of anther held 0.6-0.9 mm. below the summit
of the corolla-throat, surpassing the faucal appendages 0.6—-0.8 mm.; style
17-18 mm. long, surpassing the strict corolla-lobes ca. 2 mm.

A plant known only from the type collection now preserved in the
herbarium at Paris. The specimen was collected by Ghiesbreght, no. 311,
in September [? 1841] in temperate montane pine forests somewhere in
Mexico. No geographical data were provided by the collector. Until the
plant has been rediscovered, its home must remain subject to conjecture.
I suspect that it is in the mountains of Oaxaca.

Although originally described as a species of Lasiarrhenum, the plant
has no close relationship with that genus. Its closest affinities are with
Lithospermum, and particularly with its Mexican species. It differs from
the latter chiefly in its unusual filaments and in the form of the anther.
From Lasiarrhenum it differs not only in habit and very narrow veinless
leaves, but also in its rounded corolla-lobes, its faucal appendages, its
ellipsoid pollen, and its glabrous anthers with dark-margined thecae and
recurved terminal appendage.

Since the type collection is not a generous one and has only a relatively

32 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxv

few flowers, only two corollas have been available for dissection and close
examination. Changes in the maturing corolla have not been studied, and
the possibility that the corolla may be slightly zygomorphic has not been
eliminated. The corolla-bud just before opening is ellipsoidal and at the
base is prolonged downward into a short cylindric tube. At this early stage
the swollen throat near its middle has a diameter about twice that at the
summit of the throat (i.e., at the base of the broadly overlapping corolla-
lobes), and more than twice that of the tube below the throat. The form
of this corolla-bud is unusual in the Lithospermeae, indeed is approximated
only in Onosma and Maharanga. The mature corollas retain their swollen
throat, but this appears to be less constricted above the middle than pre-
viously. In the dissections available I find some indication that the mature
corolla may possibly be somewhat prolonged on one side and may possibly
have two of its lobes slightly larger than the other three. Inside the throat
there are evident arcuate faucal appendages. These are formed partially
by invagination and partly by thickening of tissue. They are dark in color
and bear numerous but not crowded stipitate glands. It is to be noted that
the appendages are borne not at the mouth of the corolla, at the summit
of the throat, as in Lithospermum, but rather distinctly below the corolla-
mouth, slightly above the middle of the throat, and hence are not exposed
but are distinctly included in the throat.

The filaments of Perittostema are unique. They are firm in texture, dark
in color, and very strongly dorsiventrally compressed. They gradually
broaden upward from the narrow attachment and are broadest just below
their truncate or broadly obtuse summit. In shape they are triangular or
obovate-triangular. They measure usually about 1 mm. long and about
0.6 mm. broad across their summit. On the ventral side they have a small
ridge down their middle. They are attached to the anthers by a small tip
arising from the middle of their broad upper edge. These filaments of
Perittostema are extremely different from those of Lithospermum, for in
the latter genus the filaments are never laterally expanded and never other
than linear, subulate, or unguiculate. The only member of the Litho-
spermeae with filaments even suggestive of those of Perittostema is the
genus Lasiarrhenum, In the latter genus the filaments are compressed and
are broadened upward, but are much larger and more elongate, being
oblanceolate rather than triangular.

The anthers in Perittostema are relatively large for the size of the corolla
and are accommodated within it largely because the throat is somewhat
inflated. Although the anthers project above the level of the faucal ap-
pendages, their tips still fail to closely approach the mouth of the corolla.
They are completely included. The open thecae have a dark-colored
margin, an unusual feature which I have encountered among the Litho-
spermeae only in the genus Halacsya. For the lower quarter of their length
the thecae are not joined together, and in age these tend to diverge, opening
a deep acute sinus at the base of the anther. The broad summit of the
anther is appendaged by a small but definite prolongation of the connec-
tive, which is unusual in being not straight but strongly recurved. In no

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVI 33

species of Lithospermum do the anthers evidently surpass the faucal
appendages, nor do the anthers have a distinctly bilobed base. A few
species of that genus have their anthers terminally appendaged, but when
present the appendage is always straight and erect.

The pollen is ellipsoid, having the upper and lower half equal and the
usually obscure pores, commonly seven in number, equally spaced about
the equator. It is very similar to the pollen produced by species of Litho-
spermum, Macromeria, and Psilolaemus. That of Lasiarrhenum differs
in its ovoid form.

The style is early elongate and before the corolla-bud opens is pressed
upward and decurved against the still tightly imbricate corolla-lobes. It
is promptly exserted and surpasses the corolla-lobes as soon as the latter
unfold, In Lithospermum this behavior is duplicated in only a few species,
all Mexican.

In addition to its floral structures Perittostema is also notable as a small
plant with slender stems. Indeed, in these regards, among the Lithosper-
meae it is surpassed only by members of Buglossoides § Rhytispermum.
Among American members of the tribe its slender stems and very slender
leaves are approximated only by the Mexican Lithospermum strictum.
The habit of Macromeria, Onosmodium, Lasiarrhenum, and most species
of Lithospermum is very coarse and vigorous when compared with that of
Perittostema. The leaves of our plant are extremely narrow and, Laven-
dula-like, have strongly revolute margins that roll back and cover all but
the midrib on their lower surface.

6. Psilolaemus, gen. nov.

Planta perennis strigosa scabrella. Caules foliosi fruticulosi simplices
vel laxe ramosi decumbentes vel adscendentes adpresse hispiduli e radice
valida palari lignosa purpureo-tincta erumpentes. Folia omnino caulina
sessilia firma valde costata enervata vel obscurissime paucissimeque nervata
margine angusto-revoluta, facie superiore lucentia pilis rigidis brevibus
saepe valde adpressis e discis pallidis erumpentibus instructa, facie in-
feriora breviter hispidula pilulis non rariter retrorsis donata, inferioribus
oblanceolatis, superioribus lanceolatis. Cymae scorpioideae multiflorae
solitariae caulibus ramisque foliatis terminatae maturitate rectae valde
elongatae. Bracteae foliatae abundantes foliis supremis gradatim minores
lanceolatae vel lanceolato-ovatae acutae pleraque calyce breviores. Calyx
5-fidus; lobis inaequalibus linearibus vel lanceolato-linearibus erectis, eo
exteriore maximo. Corolla lutea (‘‘armeniacea”’) tubulosa 3-plo longiore
quam lata in parti tertiaria superiore gradatim ampliata alibi cylindracea,
extus pilis minutis sparsis pleraque retrorsis donata, intus glaberrima nec
appendiculis faucialis nec glandulis stipitatis praedita; lobis ovatis stricte
adscendentibus supra basim latissimis quam longitudine corollae 6-plo
brevioribus; sinibus limbi clausis basi nec incrassatis nec plicatis; faucibus
angulatis lobis aequilongis, extus apice infra bases sinuum limbi gibbosis
et infra bases gibbarum areolas depressas verticaliter elongatas apice max-
ime invaginatas gerentibus, inter areolas 5 depressas venis 3 prominulis

34 JOURNAL OF THE ARNOLD ARBORETUM _ [vot. xxxv

donatis; annulo obscuro glabro. Filamenta attenuata apicem versus fau-
cium e plicis invaginatis faucium orientia supra medium exserta basi an-
guste breviterque decurrentia. Antherae oblongae 3—4-plo longiores quam
latae utroque emarginatae nullo modo appendiculatae paulo infra medium
affixae filamentis duplo longiores solum tertiam vel quartam partem in-
feriorem in faucibus inclusae, parte majore exsertae. Granulae pollinis
ellipsoideae vel globoso-ellipsoideae 23-26 % 16-22, desuper visae cir-
culares vel obscurissime polygonales, a latere visae ellipticae margine
saepissime convexae poris obscurissime aequatorialibus (7 —) 8 donatae.
Stylus gracilis evidenter exsertus, stigmatibus 2 distinctis terminalibus.
Nuculae erectae parvae ellipsoideae albo-griseae laeves vel perinconspicue
tuberculatae carinam ventralem versus utrinque profunde punctatae vel
interrupte profundeque sulcatae, dorso convexae, ventre obtuse angulatae
carinam latam vix prominentem proferentes, sutura obscura vel non rariter
praesertim supra medium carinae inconspicue lineato-sulcata donatae. Gyn-
obasis pyramidalis facies 4 planas adscendentes angulis prominentibus pyra-
midis separatas gerens basi incrasso styli terminata. — Nomen derivatur
ab yros, calva, et Aapos, fauces, propterea quod fauces corollae glaberri-
mae et nec elandulis stipitatis nec appendiculis faucialis ornatae sunt.

Psilolaemus revolutus (Robins.) comb. nov.

Lithospermum revolutum Robins. Proc. Amer, Acad. 27: 182 (1892).
Onosmodium revolutum (Robins.) Macbr. Contr. Gray Herb. 49: 21 (1917),

Plant at first simple and erect but becoming branched and spreading in
age; stems pale, 1-3 mm. thick, slightly fruticulose; leaves all cauline,
with a strong conspicuous midrib which is sulcate above and prominent
below; lower leaves largest, 3-7 cm. long, 6-15 mm. broad; cymes elongat-
ing, becoming 10-25 cm. long in age; Talve at anthesis 7-10 mm. long,
weakly accrescent; pedicels ca. 1 mm. long at anthesis, becoming 2-4 mm,
long in age; corolla 9-15 mm. long; tubular portion of corolla 7-12 mm.
long, lower 9-10 mm. cylindric and 2—2.5 mm. thick, upper 2.5-3 mm.
expanding to form a short throat 3-5 mm. thick at the summit; sinus be-
tween lobes closed above the base by the overlapping margins of adjoin-
ing corolla-lobes, neither thickened nor plicate at the base; corolla-throat
outside with rather prominent veins and bearing five swellings and five
depressions and hence somewhat angulate, bearing a small swelling directly
below the base of each sinus of the corolla and bearing a small elongate
depression (the complement of an inflexed plait inside the corolla) located
directly under each swelling; stamens borne 0.4—0.5 mm. below the summit
of the throat, arising from a small plait-like invagination of the siiama’
wall; Haaehte 0.5—-1.1 mm. long, exserted from the throat 0.1—0.5 mm.
anthers 1.4—2 mm. long, 0.4—-0.6 mm. broad, oblong or slightly broader
above the middle, emarginate at both ends, exserted from the throat 0.7—
1.5 mm. but overtopped by the strictly ascending imbricate corolla-lobes
and hence not conspicuous; style surpassing the corolla 1-2 mm.., , frequently
emerging from corollas which have not yet attained maximum size; nut-
lets 2 mm. long and 1.5 mm. thick.

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVI 35

A plant known only from the gypseous saline marshes in southeastern
San Luis Potosi, Mexico. As has been suggested by Macbride, Contr. Gray
Herb. 49: 21 (1917), it shares characters with both Lithospermum and
Onosmodium. Its closest relations, however, are with Lithospermum, with
its Mexican species in particular.

Psilolaemus is distinguished from Lithospermum by the form of the
corolla and the form and behavior of its stamens. The corolla-throat is
angulate as in Onosmodium and as in that genus bears localized external
swellings between the filament-attachments and the base of the corolla-
sinus above. The stamens arise from small invaginations which are com-
plementary to small elongate depressions on the outside of the corolla-
throat directly below the subsinal swellings. In Lithospermum the corolla-
throat bears invaginate appendages or stipitate glands or both, and the
corolla-tube, sometimes hairy inside, always bears a basal annulus mod-
erately to well developed. In Psilolaemus the annulus is very obscure and
the inside of the corolla is completely glabrous and devoid not only of
faucal appendages but also of all stipitate glands. The exserted style and
the nearly erect corolla-lobes of the present genus are features duplicated
in only a few of the many species of Lithospermum.

The plant has flowers resembling those of Onosmodium in being glabrous
inside and devoid of stipitate glands and faucal appendages, as well as by
having erect corolla-lobes, an exserted style, and an angulate throat with
swellings below each corolla-sinus. It differs from Onosmodium in having
elliptic pollen, oblong non-apiculate strongly exserted anthers, longer fila-
ments, broadly and persistently imbricate ovate corolla-lobes, and closed
corolla-sinus not thickened or plicate at the base. The leaves of Onos-
modium are strongly veined. Those of Psilolaemus, as in most species of
Lithospermum, are veinless or practically so. The style may be early ex-
serted, but the flowers of Psilolaemus are not precociously sexual as in
Onosmodium.

A a Maxim. Bull. Acad. St. Petersb. 17: 443 (1872) and
Mel. Biol. 8: 543 (1872). Type species A. japonica Maxim.

Plant perennial, minutely strigose, stems fistulose, erect, simple. Leaves
alternate, oblanceolate, evidently veined, all cauline; on lower half of stem
small and imperfectly developed, deciduous at flowering time; on upper
half of stem becoming very large and ample. Cymes scorpioid, 3—5, loosely
disposed at the top of the stem, lowest one axillary, the others extra-axillary
or terminal, simple, pedunculate, bracteate only towards the base, at ma-
turity becoming stiff, straight elongate and loosely racemose. Calyx 5-
parted, usually evidently pedicellate at least at maturity; lobes linear or
sometimes oblanceolate, acute, erect, weakly costate, subequal or the
abaxial one longest, about half as long as the corolla tube. Corolla laven-
der (becoming orange in drying), minutely strigose outside; tube sub-
cylindric, length greater than the diameter of the limb, abruptly con-
stricted at the base, upper portion not differentiated into a distinct throat,
bearing no faucal appendages or glands, inside abundantly and antrorsely

36 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxv

villose-strigose above the middle; annulus represented by 10 tufts of hairs
just above the base of the tube; lobes ascending, rounded, elongate, ellip-
tic, imbricate in the bud; stamens included; filaments unguiculate, affixed
at equal heights near the middle of the corolla tube, a third to a half the
length of the anthers; anthers linear-oblong, affixed between the base and
the middle, borne in the hairy upper middle third of the corolla tube, ends
emarginate. Pollen small (20 x 12-13 » long), elongate with rounded
ends, constricted at the middle, encircled by a single row of 6 (— 8) incon-
spicuous pores about the broadest part of the broader lower half of the
grain, polar profile circular or sometimes very obscurely polygonal. Style
reaching up to the middle of the corolla-tube; stigmas 2, terminal, erect
and ellipsoidal, juxtaposed and usually connate on the back below their
middle. Ovary at anthesis 4-lobed, the lobes ellipsoidal. Nutlets smooth,
lustrous, gray, elongate, obliquely lanceolate and conspicuously rostrate
with the tip hamate, only one or two maturing, broadest just above the
base and then very gradually attenuate into a very conspicuous slender
laterally flattened strongly out-curving subulate beak with a hooked tip;
venter of seminiferous lower half of nutlet distinctly convex, weakly or
not at all keeled, lacking an evident suture; dorsum obtusish; attachment
scar basal, large, slightly broader than long, flabelliform, bearing a conspicu-
ous pit formed by the open funicular canal and (paralleling the dorsal
edge) an arc of about 6 minor vascular strands. Seeds asymmetric, the
lower end appearing obliquely truncate. Gynobase pyramidal when matur-
ing more than a single nutlet, nearly as high as broad, with cartilaginous
thickenings between the attachment faces and at the apex.

A very distinct monotypic genus of Japan especially notable for its
very ample veined cauline leaves, its nearly bractless cymes, its corollas
with the tube densely villous-strigose inside, and its elongate nutlets which
are gradually attenuate into an elongate putcurved beak hooked at the
apex. It appears to be most closely related to Lithospermum, which it re-
sembles in pollen and in having broadly basifixed nutlets with a lustrous
smooth pericarp.

Ancistrocarya japonica Maxim. Bull. Acad. St. Petersb. 17: 444
(1872) and 20: 471 (1875); Mel. Biol. 8: 544 (1872).

Ancistrocarya japonica var. albiflora Honda, Bot. Mag. Tokyo 49: 790 (1935).

Endemic to middle and southern Japan. The stems, branched only in
the apical inflorescence, are 3-10 dm. tall. The underground parts are
unusual and merit description from fresh material. They apparently con-
sist of a congested sympodial rhizome that forms a crowded cluster of
short, ascending, thickened, bulbose branches, each producing an aerial
stem. No radical leaf-clusters are developed. The stems arise directly
from the rhizome. The leaves are imperfectly developed at the base of
the stem but rapidly increase in size as they are produced higher up along
it and above the middle of the stem become unusually large and ample
for an herbaceous borage, as much as 20 cm. long and 7 cm. broad. The

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVI 37

leaves of Ancistrocarya not only differ from other Lithospermeae in their
large size but also in being conspicuously and loosely veined with the veins
anastomosing.

Among the Lithospermeae the genus is also aberrant in its nearly bract-
less cymes. The lowermost flowers in the scorpioid cyme may be opposed
by a small foliar bract 10-20 mm. long, and the second flower in the cyme
may be subtended by a bract 1-3 mm. long, but the numerous later flowers
are borne on a completely bractless rhachis. The cymes themselves have
differing relations to stem-leaves. When more than two cymes are produced
by the stem, the lowermost one is distinctly axillary and the others either
distinctly supra-axillary or not at all intimately associated with a leaf.
The two terminal cymes are usually about equal in development and may
appear to be geminate. However, since no flower is borne in the fork at
their base, they are simple cymes and not formed by the basal forking
of a terminal cyme.

Superficially the corollas of Ancistrocarya much resemble those of the
American species of Lithospermum, particularly so when seen in the her-
barium. When dried they even assume an orange tonality similar to that
presented by the orange or yellow corollas of the American plants in the
same state. Unlike the American plants, however, Ancistrocarya has
corollas that are lilac or bluish (or rarely white) when fresh, The radially
symmetric corolla (11-14 mm. long in total length) has a well-developed
tube 7-10 mm. long and 2.5-3.5 mm. thick, which is abruptly constricted
to 1-2 mm. thick at the very base. At the summit it abruptly expands
into the ascending corolla-limb. There are no markings nor constrictions of
the tube to delimit the throat. The lower half of the tube is glabrous in-
side except for the ten tufts of hairs representing the undeveloped supra-
basal annulus. The inner wall of the upper half of the corolla tube is
clothed with abundant, antrorsely appressed, slender white hairs 0.5—
1 mm, long. The filaments (0.8 mm. long), all equal and all affixed at
the same altitude on the corolla, arise from amongst the hairs near the
base of the villose-strigose upper portion of the corolla-tube. The anthers
(1.5-2 mm. long) are accordingly mattressed dorsally by the dense indu-
ment on the corolla-walls; a condition paralleled among the Boraginoideae
only in Echiochilon, Sericostoma, and one species of Lithodora. Another
feature distinguishing the corolla from that of most species of Lithosper-
mum is the complete absence of faucal appendages and stipitate glands.

In form, the nutlets of Ancistrocarya are unique, and furthermore the
most bizarre in the family. They have a broad basal attachment and are
broadest (ca. 2.5 mm.) about 1 mm. above their base. Above their broad-
est part they become gradually attenuate and prolonged into a strongly
out-curving, laterally compressed subulate beak bearing a hook at the tip.
Of their total length (7-9 mm.), only the lower half is seminiferous. Al-
though the ovary is 4-lobed, only one or two of the lobes mature into nut-
lets, either a solitary abaxial one or an abaxial one and its adjacent axial
companion. When two are developed they are somewhat divergent, for
they are basifixed on adjacent faces of a pyramidal gynobase. The slender

38 JOURNAL OF THE ARNOLD ARBORETUM _ [vot. xxxv

hooked beak is strongly curved outward (away from the center of the
flower), but the lower lanceolate seminiferous body of the nutlet is only
very slightly so. The ventral side of the nutlet body is more rounded
and swollen than the dorsal side. It develops no longitudinal ventral keel
and furthermore bears no trace of a suture. The elongate, curved, subulate
beak is a sterile apical prolongation of the pericarp, completely without
parallel elsewhere in the Boraginaceae. Hooked at the tip it may function
as an adaptation useful in animal dissemination.

8. Buglossoides Moench, Meth. 418 (1794). Type species B. ramosis-
stma Moench. (= Lithospermum tenuiflorum L.)

Aegonychon S, F. Gray, Nat. Arrang. Brit. Pl. 2: 354 (1821). Based upon
0 species, Lithospermum purpureo-caeruleum L. and L. arvense L.
Khytispermum Link, Handb. 1: 579 (1829). Based upon eight species, of
which the first and third (Lithospermum arvense L. and L. pur pureo-

caeruleum L.) represent the present genus. The other six species mentioned
belong to Neatostema (1 sp.), Lithodora (2 spp.), Alkanna (2 spp.), and
Rochelia (1 sp.).

Margarospermum Opiz in Berchtold & Opiz, Oekon.-techn. Fl. Béhmens
2°: 73 (1839). Although Opiz cites Lithospermum § Margarospermum
Reichenb. (1831) as a synonym, the description of his genus is based solely
upon Lithospermum purpureo-caeruleum L.

Plant annual or perennial, herbaceous or fruticulose. Leaves veinless or
nearly so, Cymes unilateral, simple or geminate or ternate, usually elongate,
conspicuously bracted, usually racemose at maturity. Calyx 5-parted or
5-lobed, lobes narrow, at times united at the base to form a short cupulate
tube, equal or unequal. Corolla blue, bluish, or white, funnelform or hyper-
crateromorph, outside usually bearing some appressed hairs, inside from
the base of the corolla-lobes downward to between the tips of the anthers
bearing five distinct longitudinal bands of hairs and/or glands or five
hairy and/or glanduliferous inflexed plaits, below the insertion of the fila-
ments naked or with five congregations of glands and sometimes with in-
vaginate swellings, lobes spreading or ascending, imbricate; annulus near
the base of the tube consisting of a narrow collar or ten scale-like lobes,
frequently weakly developed and sometimes apparently absent; stamens
borne at or below the middle of the corolla, included; filaments equal,
affixed at equal heights on the corolla, shorter than the anther (usually
about half as long); anthers lance-oblong or oblong, affixed at or slightly
below the middle, base cordate, apex appendaged by a short prolongation
of the connective; pollen small (15-25 » long, 12-16 in diameter), cylin-
dric or somewhat ovoid or rarely slightly elliptic, bearing the inconspicuous
pores in a single row on or distinctly below the equator, in lateral profile
with the sides usually straight or nearly so and either parallel or some-
what convergent towards the upper end; style not much if at all surpass-
ing the anthers, usually much shorter than the calyx; stigmas two, rounded,
borne laterally at the summit of the simple style or at the base on opposing
sides of a short bilobed sterile apex of the style, usually evidently sub-

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVI 39

terminal; nutlets 1-4, erect to strongly divergent, smooth or rough, rounded
or angulate, attachment basal or obliquely basal, ventral suture fused,
obscure, sometimes prominent; gynobase flat or depressed pyramidal.

This genus is closely related to Lithospermum and is distinguished only
by the structure of the corolla. In Lithospermum the corolla-throat may
bear hairy and glanduliferous, invaginate, gibbose appendages or be vari-
ously glanduliferous or glabrous. The distinctive feature of the corolla of
Buglossoides is the well-developed guides for insect visitors in the form
of five distinct vertical bands of glands and stout hairs or five hairy and
glanduliferous inflexed plaits. These guide-lines may be traced from the
base of the corolla-lobes downward along a vein to between the anther
tips or slightly above them. The gibbose invaginate faucal appendages
present in many species of Lithospermum are also a feature of the flowers
in many other genera of the Boraginaceae. The elongate inflexed plaits
or the well-developed vertical bands of hairs and glands occur only in
Buglossoides. Several other developments in the genus distinguish its
species from most members of Lithospermum. These are the prevalence
of blue as a corolla color, the apiculate anthers, the lobed sterile tip of
the style, and the small size of the usually cylindric or ovoid-cylindric
pollen grains. Singly, these latter features may occur here and there in
species of Lithospermum, but never together nor in any species possibly
a close relative of any species of Buglossoides. Floral dimorphy, in the
form of heterostyly or cleistogamy, well developed in Lithospermum, does
not occur in the present genus. Since the group gives every evidence of
being a natural one, and since it can be readily distinguished from Litho-
spermum by well-marked, very unusual features of its corolla, it seems to
merit generic recognition.

The stamens of Buglossoides are deeply included in the tubular portion
of the corolla in all species except B. purpureo-caeruleum. In Lithosper-
mum, stamens are borne at or below the middle of the tube only in species
with a very abbreviated tube or in corollas of long-styled flowers of some
heterostyled species.

The anthers are oblong or narrowly oblong and several times longer
than broad and are borne on filaments half to a third of the anther length,
in all species except B. Gastoni. In the latter the anthers are broadly ob-
long and only about twice as long as broad, and are borne on filaments
nearly as long as the anther body. The largest anthers are those of B.
purpureo-caeruleum and B. Zollingeri. These are 1-1.5 mm. long, twice
the length of the anthers in other species. In all species the connective is
prolonged to form a minute but definite tip on the anther. This tip may
be subulate, cuneate, deltoid, or quadrate. It becomes as much as 0.2 mm.
long in B. purpureo-caeruleum, but in other congeners it is usually only
0.1 mm. or less in length. It is least developed in B. Gastoni, in that species
being stout, quadrate or deltoid, and scarcely projecting beyond the sum-
mits of the thecae. This tip on the anthers in species of Buglossoides is a
character that separates the genus from practically all species of Litho-
spermum. The connective is prolonged apically in only a very few species

40 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxv

of Lithospermum, noticeably only in L. tschimganicum Fedtsch. of Central
Asia and L. papillosum Thunb. and L. cinereum DC. of South Africa.
Some of the Mexican species of Lithospermum have the anthers obscurely
thickened at the apex but never to the degree found in Buglossoides. In
most species of Lithospermum there is not even a vague suggestion of a
terminal appendage.

The style is short. At anthesis it never reaches upward beyond the tips
of the anthers. Rarely it may bear its stigmas between the anthers, but
more commonly they are held below the level of the anther bases. In fruit
the style is never more than half the length of the mature nutlets. In
Lithospermum such short styles occur only in cleistogamic flowers or in
the short-styled flowers of some heterostylic species.

The two stigmas are separated by the sterile tip of the style and are
accordingly lateral and borne at or more commonly obviously below the
style apex. The sterile tip of the style is commonly prolonged above the
stigmas and is usually evidently bilobed. This condition is extremely well
developed in B. purpureo-caeruleum and B. Zollingeri. In these two species
the style tip projects beyond the stigma for a distance equal to or sur-
passing the diameter of the latter, and consists of two attenuate lobes.
In B. calabrum and B. Gastoni the lobes of the style tip are shorter, stouter,
and obtusish, but even so, they evidently project above the stigma. In the
four species named, all members of the section Margarospermum, the stig-
mas are Clearly subapical and well separated. Frequently the two stigmas
are not equally well developed, and not uncommonly one may be borne
slightly lower on the style than its companion.

Among the species of the section Eubuglossoides the style tips show
greater variation, The two stigmas tend to be less sharply defined than in
the section Margarospermum. In typical B. arvense the terminal lobes
of the style may be cylindric or narrowly conic and always evidently sur-
pass the stigmas. In B. incrassatum the lobes are evident, but usually
less well developed than in B. arvense and its allied forms (e.g., Litho-
spermum Sibthorpianum Gris.) in the eastern Mediterranean area. In
B. tenuiflorum, in most allies of B. arvense in northern Africa, as well as
in plants referable to L. minimum Morris, the sterile tip of the style ex-
tends up between the stigmas but projects weakly if at all above them.
The sterile tip of the style in these plants has either an unlobed truncate
apex or an apex that is only obscurely lobed or merely notched.

The prolonged lobed sterile tip of the style in this genus needs investi-
gation. Superficially the stigmas and the tip of the style, taken together,
are very Suggestive of the stigmatic head of the Heliotropioideae. It re-
mains to be determined if the course of the vascularization is similar, cf.
Hanf. Beiheft Bot. Centralbl. 544: 126 (1935). Among the Lithosper-
meae lobed style-tips are best developed in the present genus. In almost
all species of Lithospermum the stigmas are terminal on the style, or if
there is a prolongation of the style-tip between the stigmas it is only
moderate and does not surpass the stigmas. Only in a few Mexican
species (L. Nelsoni, L. Berlandieri, L. mediale, L. oblongifolium, L. calci-

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVI 41

cola, L. sordidum, and L. distichum) does the somewhat lobed style-tip
ever project above the stigmas. However, in most of these species the
style-tip is actually short and becomes prominent only because the oblong
stigmas are widely spreading.

The pollen of Buglossoides is small and shows little variation in size
from species to species, Though sometimes ellipsoidal or ovoid, its usual
form is subcylindric or barrel-shaped, being circular in polar outline and
commonly straight-sided or nearly so in lateral profile. Its pores, always
in a single row, are extremely small and obscure, so much so that I have
been unable to determine their number. In the section Exbuglossoides the
pores are distributed around the grain exactly half-way between the poles,
i.e., on the equator. The upper and lower halves of the grains are always
similar in size and outline. In lateral profile the sides are usually straight
and parallel and only occasionally somewhat convex. With proper light-
ing the grains may show a vague but definite narrow band about the
equator. This equatorial band in which the pores are located has not been
detected in any pollen of the other section of the genus. In the section
Margarospermum the pores are borne perceptibly below the middle of the
grain. The shorter lower portion of the grain (that below the line of pores)
is usually perceptibly broader than the upper portion. In lateral outline
the grains vary from oblong to more or less ovate, even within a single
species. In profile they may have their straight sides practically parallel
or very slightly convergent towards the upper end. Sometimes the sides
are evidently convergent and the grain obviously somewhat ovoid. Fre-
quently the grains show a very localized contraction just above the pores,
with the result that they tend to develop short sloping shoulders which,
though very much less well developed, are still recognizable as similar to
the shoulders developed on the asymmetric pollen of Lithospermum, cf.
Jour. Arnold Arb, 33: 310 (1953). Such grains, narrowed very slightly
in a zone above the pores (to form the shoulders), usually have their upper
half with straight paralleling sides, but this portion is discernibly narrower
than the shorter rounded basal portion. Accentuation of these tendencies
would produce the strongly asymmetric grains of the footprint and hour-
glass type known in Lithospermum. In the genus Buglossoides, however,
this tendency to develop shoulders on the grains is only very weakly ex-
pressed and frequently must be looked for before it is detected.

The distinctive corollas of Buglossoides have been illustrated and com-
pared by Spengler, Oesterr. Bot. Zeitschr. 68: 110 and 116, ff. 1, 2, 23-26
(1919). Synonymy for most of the species has been compiled by Stroh,
Beih. Bot. Centralbl. 58 ®: 203-4 and 206 (1938).

Section Eubuglossoides.

Lithospermum § Rhytispermum (Link) Reichenb. Fl. Germ. Excur. 336
(1831).

Plant annual or biennial, herbaceous; corolla small, 4-9 mm. long, white
to blue, regular, inside bearing 5 vertical bands of glands or hairs but no

42 JOURNAL OF THE ARNOLD ARBORETUM _ [vot. xxxv

strongly inflexed plaits, lacking congregations of glands or invaginations
directly below the stamen attachments; pollen symmetric, bearing the pores
about the equator; style with or without a prolonged bilobed sterile tip;
nutlets rough, usually all four developing, ventral keel prominent.

The corollas of species belonging to this section are more simply organ-
ized than those of the section Margarospermum. In the present section
the walls of the corolla are only very slightly swollen beneath the five
vertical bands of hairs and glands on its inner surface. Complementing
these bands on the outside of the corolla are merely five shallow lineate
grooves. Only an incipient tendency for invagination along the hairy ver-
tical bands is accordingly present. In the section Margarospermum the
invagination is very pronounced, and the hairs and glands clothe well-
developed inflexed plaits that form the five intruding ridges on the inside
of the corolla.

To be included in the section Eubuglossoides is Lithospermum tenui-

orum L.., as well as L. arvense L. and the undetermined number of critical
species all closely related to it.

Buglossoides tenuiflorum (L. f.) comb. nov.
Lithospermum tenuiflorum Linn. f. Suppl. 130 (1781).

A well-marked species which ranges from Greece east to Irak. The nut-
lets are distinctive. They have a very fragile pericarp. They are erect or
slightly incurving and when in situ have their tips proximate and their
ventral keels parallel. These nutlets are also smaller than those of B.
arvense and its allies and are further differentiated by being distinctly
constricted just above their smaller attachment surface. The cymes are
short (even in maturity seldom more than 8 cm. long) and obviously
geminate or ternate at the ends of the branches. The flowers (with a very
small blue corolla) are always crowded and evidently biseriate on the

The calyx does not develop a cupulate tube nor does it become
enlarged abaxially as may be the case in B. arvense and its allies.

Buglossoides arvense (L.) comb. nov.
Lithospermum arvense L. Sp. Pl. 1: 132 (1753).

This species is either exceptionally variable or is a complex of minor
species awaiting analysis by a monographer. The typical form is that with
the largest flowers. Its white corollas are decidedly funnelform and evi-
dently longer than the calyx. It is the form prevalent in middle and north-
ern Europe and thence extends across Asia. In southern Europe and north
Africa there are a number of distinguishable closely related plants, some
of which are certainly geographically correlated and deserve specific recog-
nition. Among these only B. incrassatum can be mentioned in the present
paper. Others seem to merit recognition. Useful in distinguishing them
from true B, arvense are differences not merely in habit but also in size,
form, and color of corolla, size and form of the calyx lobes, degree of de-

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVI 43

velopment of the sterile style tip, and the presence or absence of a band
of scattered short ascending hairs on the inner surface of the corolla just
below the attachment of the stamens.

The fruit of B. arvense and its allies presents a number of very interest-
ing features. The bony, hard, rough nutlets are straight and have a broad
basal attachment which tends to become oblique. In the Boraginaceae
nutlets with oblique attachments ordinarily have the attachment surface
sloping upward towards the center of the flower and hence transgressing
on the ventral side of the nutlet body. In B. arvense and its allies the re-
verse condition is true. If the nutlets are held in a vertical position, it is
to be seen that the scar slopes upward, not towards the ventral but towards
the dorsal side, and that as a consequence the nutlets are shorter on the
dorsal side than on the ventral. Nutlets with an oblique attachment sur-
face of this sort, when affixed to a low-pyramidal or nearly flat gynobase,
are not erect with paralleling ventral keels, but are necessarily strongly
divergent. This strong divergence of the nutlets of B. arvense and its
allies is a development late in ontogeny. When they are young the nut-
lets are erect and parallel. They become noticeably divergent only as they
approach full maturity.

Buglossoides incrassatum (Guss.) comb. nov.

Lithospermum incrassatum Guss. Ind. Sem. Hort. Boccad. yi or
Siculae Prodr. 1: 217 (1827); Fl. Siculae Synop. 1: 217 (18

A close ally of B. arvense notable chiefly because of its remarkable
calyx. After the fall of the corolla, the calyx, as it increases in size and
the ensheathed nutlets mature, gradually becomes greatly modified in form
as a result of excessive abaxial prolongation. At first the calyx is very
similar to that of B. arvense at the same stage of development. It is
affixed centrally on its symmetric base to the pedicel. At this early stage
the central axis of the flower (or for all practical purposes, the style)
points away from the leaf axil and is hence divergent from the stem. In
later development, because of excessive growth on the abaxial side of the
calyx-base, the axis of the flower is gradually shifted in an arc as great as
90° and finally points, not away from, but actually towards the adjacent
stem. In other words the calyx is shifted from a central basal attachment
to one that is distinctly lateral. The central line up the pedicel, if pro-
jected outward, no longer passes up the style but rather across the low
gynobase, where it meets the style at an angle of as much as 90°. The ma-
ture nutlets are accordingly borne on a low gynobase which is now adaxial
and which actually faces the adjacent stem. Because of the dislocation,
the distorted calyx-base faces outward and so becomes the most conspicu-
ous part of the fruiting calyx. The condition is unusual but not unique
among the Boraginaceae. A very similar development is found in the genus
Pectocarya, cf. Jour. Arnold Arb. 20: 400 (1939).

Various authors have dismissed the remarkable fruiting calyces of this
species as teretological. With this I cannot agree. In B. arvense and other

44 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxv

near relatives of B. incrassatum, there is also a tendency for some en-
largement of the fruiting calyx on the abaxial side and also a slight but
definite tendency for the transverse axis of the gynobase to slope down-
wards towards the adjacent stem. In the present species these tendencies,
merely incipient in its relatives, are developed excessively. The wide dis-
tribution of B. incrassatum about the Mediterranean area, where in many
localities it appears to be the only representative of its group, is evidence
that it is not a casual freak.

Section Margarospermum (Reichenb.) comb. nov.

Lithospermum § Margarospermum Reichenb. Fl. Germ. Excur. 337 (1831).
Founded to include four species of which the first listed was Lithospermum
purpureo-caeruleum L.; the other three are representatives of the genera
Lithodora and Moltkia

Plant perennial, herbaceous or fruticulose; corolla larger, 15-19 mm.
long, blue or purple, the limb slightly oblique with the three abaxial lobes
more spreading than the posterior two, inside bearing 5 vertical glandu-
liferous and/or hairy inflexed plaits and below each filament attachment
bearing a congregation of glands and sometimes an invaginate swelling;
pollen slightly asymmetric, the pores borne below the equator; style
always with a bilobed sterile tip; nutlets smooth and lustrous or rugose,
not strongly keeled, usually only one maturin

This section includes only the four extremely well marked species enum-
erated below.

Buglossoides purpureo-caeruleum (L.) comb. nov.
Lithospermum purpureo-caeruleum L. Sp. Pl. 1: 132 (1753).

Ranging from western Europe to Iran. Leaves usually broadest below
the middle. Flowering stems clustered, arising from a densely branched
rhizome, terminated by a pair of leafy racemose cymes. Corolla bear-
ing faucal plaits 2-3 mm. long which originate 6-9 mm. above the base
of the tube; also bearing distinct invaginate swellings 2 mm. long below
each stamen; plaits glanduliferous but not hairy. Filaments attached
5—8 mm. above the base of the corolla; anthers ca. 1.5 mm. long, elongate,
apiculate, twice as long as the filament. Style 8-9 mm. long, the sterile
tips attenuate. Nutlets ellipsoidal, smooth, white, 3.5-4 mm. long, 3-3.5
mm. thick, back convex, venter obtuse

The nutlets of this species are usually slightly larger but otherwise very
similar to those of B. Zollingeri and B. calabrum. In being smooth, white,
and porcelain-like, they much resemble the type of nutlet prevalent in
Lithospermum., Unlike the nutlets of the other species of Buglossoides
those of the present species have a relatively small basal attachment-
surface which is located not at the center of the nutlet-base but rather on
its ventral half. For notes on the habit of growth of this species see
White, Jour. Bot. 22: 74-76 (1884).

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVI 45
Buglossoides Zollingeri (A. DC.) comb. nov.
Lithospermum Zollingeri A. DC. Prodr. 10: 586 (1846).

A plant of Japan, China, and Korea. Leaves usually broadest above the
middle. Flowering stems arising from procumbent stems persisting from
the previous season, terminated by a simple unilateral cyme. Corolla on
inner surface bearing glanduliferous, short-hairy inflexed plaits 4-5 mm.
long which originate 5—6 mm. above the base of the tube, and also provided
with short glanduliferous invaginate plaits below the attachment of each
stamen. Filaments attached 3-4 mm. above the corolla base; anthers 1.5
mm. long, elongate, apiculate, several times as long as the filaments. Style
2-4 mm. long, the sterile tips attenuate. Nutlets ellipsoidal, 3-3.5 mm.
long, smooth, white, back convex, venter obtuse or weakly keeled.

Collectors appear to have been aware of this plant only in its flowering
state. Nearly a hundred collections of the species in American and Euro-
pean herbaria have been examined, but in this large suite only five collec-
tions show the plant in the fruiting state, and none of them bear nutlets
that are completely mature.

Buglossoides calabrum (Tenori) comb. nov.
Lithospermum calabrum Tenori, Fl. Nap. 3: 174 (1824-29).

Endemic in southern Italy. Leaves broadest at the middle. Flowering
stems arising from slender procumbent stems persisting from the previous
season, terminated by a simple racemose cyme. Corolla with a tube evi-
dently much surpassing the calyx, inside bearing hairy, sparingly glandu-
liferous plaits 7-8 mm. long which arise 4-6 mm. above the base of the
tube, below the filament attachments obscurely if at all invaginate. Fila-
ments attached 2 mm. above the base of the tube; anthers almost 1.5 mm.
long, elongate, apiculate, twice as long as the filaments. Style 1-2 mm.
long, the sterile tips stout and obtuse. Nutlets ellipsoidal, smooth, white,
3.5 mm. long, 2.5 mm. thick, back convex, venter obtuse.

Buglossoides Gastoni (Benth.) comb. nov.

Lithospermum Gastoni Benth. ex DC. Prodr. 10: 83 (1846); Bot. Mag. 47:
t. 5926 (1871).

Known only from the French slopes of the western Pyrenees. Leaves
broadest below the middle, lanceolate. Flowering stems clustered, arising
from a densely branched rhizome, terminated by a crowded densely flowered
forked cyme. Corolla with hairy and glanduliferous inflexed plaits 3—4
mm. long which arise 2.5-3 mm. above the base of the tube, bearing no
invaginations below the filament attachments. Filaments borne 1-1.5
mm. above the base of the corolla; anthers oblong, less than 1 mm. long,
weakly apiculate, about as long as the filaments. Style 0.5—1 mm. long,
the sterile tips stout. Nutlets yellowish, punctate and abundantly rugulose,
very stout and plump, 4.5 mm. long and nearly as thick, keeled only about
the sharp apex, attachment broad and basal.

46 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxv

A very distinct species differing from the other members of its section
in having small anthers, very congested forked cymes, and very large,
very plump, short-beaked, yellowish rugulose nutlets.

9. Stenosolenium Turcz. Bull. Soc. Nat. Moscou 13: 253 (1840); DC.
Prodr. 10: 103 (1846). Type species Anchusa saxatilis Pallas.

Plant a hispid annual herb with one to many erect or decumbent stems.
Leaves costate but veinless, lower oblanceolate, the upper ones becoming
lanceolate. Inflorescence scorpioid, elongate, bracted throughout (lower
bracts foliaceous), distinctly racemose in age. Calyx short-pedicellate,
S-fid, hispid; lobes slenderly lanceolate, attenuate, slightly unequal with
the abaxial one usually the longest. Corolla violet or purple, salverform,
outer surface with minute, short, usually spreading hairs, inner surface
with a villulose annulus but otherwise glabrous, bearing no stipitate glands;
limb spreading, breadth less than the length of the tube, rounded lobes
slightly broader than long, imbricate in the bud; tube elongate, about
twice as long as the calyx, subcylindric, upper third to half broadest and
forming an ill-defined throat, above the base bearing a short thickish
collar-like villulose annulus; throat unappendaged. Stamens borne at
unequal heights in the throat, included, the uppermost one usually
abaxial; filaments slender, a half to a third as long as the anther;
anthers elongate, laterally compressed, affixed near the middle. Pollen
elongate with rounded ends, symmetric, constricted at the middle, 38-41
x 22-25 » (16-20 p» thick at the middle), bearing two rows of 8 or 9 in-
conspicuous pores one at each end of the grain, polar profile circular.
Style shorter than the calyx, forked below the apex; stigmas two, distinct,
compressed, obovate-spathulate. Gynobase flat or concave, with four
triangular attachment faces, usually maturing four nutlets. Nutlets tuber-
culate, usually brownish, ascending, supported on a stout, laterally affixed
inframedial vertical stipe; axial edge angulate, vertical, nearly straight,
above the middle formed by the short ventral keel on the nutlet-body and
below the middle by the ventral side of the stipe; seminiferous body of
the nutlet ovoid, inclined at an angle of 40-45°, its pointed apex held
above the nutlet-attachment and its rounded base abaxial to it; stipe stout,
usually partially hollow, expanding abruptly at the base, sides minutely
verrucose; funicular canal ascending vertically inside the stipe and enter-
ing the oblique nutlet-body near the middle of its ventral side; ventral
keel short, supramedial, formed of a usually completely fused ventral
suture. Seed straight, funicular attachment slightly above the middle.

A monotypic genus of eastern Asia. In most characters it shows close
similarities with the Asiatic species of the genus Arnebia and is probably
most closely related to them. Its fruit, however, is very different, not only
from Arnebia but from all other members of the Lithospermeae.

Stenosolenium saxatile (Pallas) Turcz. Bull. Soc. Nat. Moscou 13:
253 (1840).
Anchusa saxatilis Pallas, Reise 2°; 718, t. “F”, f. 1 (1773).

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVI 47

Onosma saxatile (Pallas) Lehm. Asperif. 2: 371 (1818).
Arnebia saxatilis (Pallas) B. & H. ex Forbes & Hemsl. Jour. Linn. Soc. 26:
155 (1890).

Ranging from the mountains near Pekin and northwestward across east-
ern Mongolia towards the Baical region.

The corolla is 12-19 mm. long in total length and has a spreading limb
7-10 mm. broad. The tube is 8-12 mm. long and its lower half or two
thirds averages ca. 1 mm. thick. The upper half or third of the tube, how-
ever, is perceptibly thicker (ca. 1.5 mm.) and so forms an ill-defined cylin-
dric throat. This elongate throat bears no hair nor glands on its inner
surface and no appendages.

The stamens vary as to their distribution in the throat. In some plants
they are affixed all at evidently different altitudes in the throat along most
of its length, while in others they tend to be grouped below the middle
of the throat, most of them with very slight differences in height of attach-
ment and only one obviously higher than the others. When the stamens
are loosely distributed, the uppermost stamen may be affixed 2-3 mm.
above the lowermost one, but when crowded towards the base of the throat,
the maximum difference in height of attachment may be reduced to 0.5—
1 mm. The distribution of the stamens seems best described as spiral.
Except that the uppermost one always appears to be on the abaxial side
of the tube, the relative heights of the stamens seem to have no relation
to any possible plane of symmetry in the corolla. The behavior of the
androecium is suggestive of that in forms of Arnebia decumbens, cf. Jour.
Arnold Arb. 33: 322 (1952). In that species the stamens may have a loose
spiral arrangement in the corolla-throat or be grouped near the summit
of the tube, sometimes with only one stamen attached obviously lower
than the others. This grouping, when present, is at the top of the throat
rather than at the bottom as in Stenosolenium, but the extreme range of
variation in stamen attachment is otherwise very similar.

Although there are differences among plants as to the distribution of the
stamens in the throat, the variation seems to be independent of any dif-
ferences in style length. The forked style is always short, 3-5 mm. long,
and reaches up into only the narrow lower half of the corolla-tube. The
two compressed, somewhat spathulate stigmas are borne, accordingly, al-
ways well below the level of the lowest stamen. There is no evidence that
either heterostyly or cleistogamy are ever present.

The pollen of Stenosolenium is elongate, constricted at the middle, with
the upper and lower halves similar in size, form, and other details. At both
ends, where broadest, it is encircled by a set of eight or nine equally spaced
very conspicuous pores. These elongate, medially constricted, symmetric
grains with biseriate pores closely resemble those of the species of Arnebia.
Indeed, the only striking difference appears to be in the number of pores
in each encircling row, eight or nine in Stenosolenium and four or five in
Arnebia.

The nutlets of Stenosolenium are very distinctive. Viewed laterally,
their seminiferous body (2 mm. long) is inclined about 45° and is supported

48 JOURNAL OF THE ARNOLD ARBORETUM __ [voL. xxxv

ventrally by a stout downwardly directed vertical stipe. This stipe is
affixed obliquely below the middle of the sloping (ventral) under side of
the nutlet-body and is short-columnar with the base ampliate and flaring.
In lateral profile the nutlet has a vertical, nearly straight ventral edge
(2 mm. high) which is formed above the middle by the short vertical
ventral keel and below the middle by the ventral side of the stipe. Because
of this the stipe appears to be a downward projection from the base of the
short supramedial ventral keel. This keel, formed of a prominent fused
ventral suture, ends abruptly. There is no suggestion of a downward pro-
longation onto the stipe below it. The stipe is minutely verrucose on all
sides. Although distinct from the nutlet-body for most of its length, it
remains close to the latter and its lower attachment end is in the same
horizontal plane as the lower end of the body. Superficially, at first
glance, the nutlet seems to have a strongly excentric but still basal attach-
ment. A study of the course of the funicle, however, proves this is not
the case. The funicle in its tubular canal is conducted upward from the
gynobase inside the partially hollow stipe and then through the pericarp
near the middle of the sloping ventral side of the nutlet-body. The im-
portant fact here is that the funicle enters the nutlet-body not at the
base but below its middle on the side. In the strict morphological sense,
the nutlets of Stenosolenium are laterally attached! Were the stipe on
these nutlets suppressed and the attachment scar sessile, the latter would
have a position almost half-way up on the venter of the nutlet. Steno-
solenium, accordingly, has a character of the Eritrichieae.

The position of the attachment on the nutlet-body usually has phylo-
genetic significance and is the most important single character used in
assigning genera to the tribes of the subfamily Boraginoideae. As an in-
dicator of natural relationships it is prevailingly satisfactory, but as with
all single characters, there can be expected to be instances in which its
indications are not acceptable. Such seems to be the case as regards
Stenosolenium. Only in the position of its nutlet attachment does that

Among the Boraginaceae distinctly stipitate nutlets are developed in
only a relatively few genera, in Stenosolenium and Alkanna of the Litho-
spermeae, in Caryolopha of the Anchuseae, and in Plagiobothrys § Echi-
diocarya of the Eritrichieae. Because it is partially hollow and has a dilated
base, the stipe on the nutlets of Stenosolenium obviously differs in struc-

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVI 49

ture and appearance from those of the other genera mentioned. Surpris-
ingly, the closest approximation is found in the obconic obliquely affixed
attachment on the fossilized nutlets of the extinct Prolithospermum John-
stonii Elias, Special Paper Geol. Soc. Amer. 41: 105, t. 15, f. 10 (1942)
and Am. Midland Nat. 36: 374-77, f. 3 (1946). These remarkably pre-
served nutlets, all that is known of the species, are found in middle Plio-
cene deposits of Kansas and Nebraska in middle United States. Thanks
to Prof. Maxim K. Elias of the Nebraska Geological Survey, I have had a
series of them for close study and comparison. They have a smooth, nearly
erect, ovoid, slightly asymmetric body 2—2.5 mm. long, and on their con-
vex side below the middle and above the rounded base bear a short, hol-
low, downward directed, basally expanding appendage with the attachment-
surface on its horizontal base. This appendage is structurally similar to
the stipe on the nutlets of Stenosolenium. Indeed, it differs only in originat-
ing less high on the side of the nutlet, in being shorter, and in being
laterally adnate to the nutlet-body rather than free and slightly divergent
from it. If the appendage on the nutlet of Prolithospermum were free
rather than completely adnate laterally, it would be shorter, but would
otherwise closely resemble the stipe in Stenosolenium. The homologies are
so very clear that I am content to believe that the fossil plant belonged
in a group ancestral to Stenosolenium and possibly even congeneric with it.

The nutlets of the extinct Prolithospermum give us an early stage in the
evolution of the medio-lateral stiped attachment of the nutlets of Steno-
solenium, and furthermore are suggestive of the manner by which it could
have been originally evolved from a truly basal attachment on a Litho-
spermum-like nutlet. The attachment in our plants was probably evolved
from a broad, horizontal, exactly basal attachment by shifting its center
first to the ventral side of the nutlet-base and subsequently to a buttressed
projection on the ventral side of the latter. As the attachment-surface,
still large and still in relatively the same horizontal plane, became more
and more to one side of the nutlet-base, the latter would cease to be trun-
cate and would become more and more convex. The laterally dislocated
attachment-surface could remain in the same relative plane only by but-
tressing its support higher on the ventral side of the nutlet-body. The
combined results would place the attachment at the base of an obliquely
affixed outgrowth to one side of the rounded base of the nutlet-body, pre-
cisely as is to be seen in the nutlets of Prolithospermum. A still further
horizontal shift of the attachment would produce a still higher buttress
on the nutlet venter and eventually even a “flying buttress’ (i.e., the
stipe) on the nutlets of Stenosolenium.

The peculiar feature of this support of the attachment in Stenosolenium
and Prolithospermum is that it is more or less hollow. It is a tube-like
or somewhat funnel-like organ which conducts the funicular canal from
the flat gynobase upward to the point where it enters the nutlet-body.
Accordingly it appears to be appendicular on the pericarp and not formed
by constriction of the pericarp above the attachment or by localized gibbose
prolongations of its walls, as in other stipitate nutlets. In various species

50 JOURNAL OF THE ARNOLD ARBORETUM _ [VvoL. xxxv

of Lithospermum and its relatives the attachment scar is surrounded by a
projecting rim of pericarp. The nutlets of Lithospermum incisum provide
one of the best examples of this. In that species the abscission is about
the edge of the collar-like rim surrounding the centrally depressed scar. By
modification of such a rim attachment the obliquely affixed suprabasal
attachment support in Prolithospermum and the tubular support in Steno-
solenium might have been evolved. If the ancestors of these two genera
had such a rimmed attachment, the attachment, as it was shifted to a posi-
tion off the nutlet base, could remain in the same horizontal plane by in-
creasing the height of the rim, particularly so on the off side. Certainly
the laterally adnate, more or less funnelform support of the attachment
in Prolithospermum is very suggestive of some such origin. If such is the
case, then the tubular support of the attachment in Stenosolenium is only
a more advanced modification by which a rim about an attachment scar
has become transformed into a hollow stipe.

The gynobase of Stenosolenium is flat and horizontal, or its four attach-
ment pads slope slightly towards its center. On this gynobase a basifixed
nutlet would be erect. A nutlet with a sessile lateral attachment would be
oblique or horizontal. The peculiarities of the nutlets of Stenosolenium and
Prolithospermum appear to be those modifications necessary if the large
attachment on an originally basifixed Lithospermum-like nutlet was shifted
from the base to a more and more lateral position while at the same time
the body of the nutlets maintained an erect or nearly erect orientation
and the flat gynobase continued unaltered. As the attachment shifted, the
funicular canal within it would also be displaced and would enter the nutlet
no longer at the base, but laterally. An increasing development of tubular
stipe would be required for conducting the funicular canal upward from
the flat gynobase, for the greater the horizontal shift of the actual attach-
ment, the higher up on the venter of the nutlet body would become the
point at which the funicular canal pierced the pericarp.

Change in position of attachment from basal to lateral, and even apical,
is a progressive evolutionary trend responsible for much of the diversity of
the fruit in the Boraginaceae. The various stages represent increasing de-
grees of specialization and in general are associated with the phylogeny
of the groups in which they are illustrated. The tribe Eritrichieae has nut-
lets usually with lateral attachments. The fact that it has a higher evolu-
tionary position than the Lithospermeae is suggested also by its more
specialized corollas and other floral structures. In the Eritrichieae the
development of lateral nutlet-attachment has been concomitant with the
development of a pyramidal, spire-like or columnar gynobase. Steno-
solenium and Prolithospermum have a funicle that enters the nutlet-body
laterally, and hence, certainly in a morphological sense, have a lateral
attachment. This character, however, appears to have had an origin in-
dependent of that in Eritrichieae, most likely as an aberrant development
in the genus Lithospermum. It is an example of parallel evolution and
not an indicator of relationships in the Eritrichieae. Unlike the condition
in.that tribe, the lateral attachment of Stenosolenium was developed un-

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVI 51

accompanied by compensating alterations of the gynobase, which has con-
tinued primitively flat as in Lithospermum.

10. Arnebia Forsk. Fl. Aegypt.-Arab. 62 (1775). Type species A. tetra-
stigma Forsk.
Dioclea Spreng. Syst. 1: 502 and 556 (1825), not HBK. (1823). Type species
Arnebia hispidissima (Lehm.) DC.
Strobila G. Don, Gen. Syst. 4: 327 (1837). A renaming of Dioclea Spreng.,
t HBK.

no
Meneghinia Endl. Gen. 648 (1839). A renaming of Dioclea Spreng., not
HBK.

Macrotomia DC. in Meisner, Gen. 1: 281 and 2: ae (1840); DC. Prodr.
10: 26 (1846). Type stg o a (Wall.) D

Munbya Boiss. Diag. ser. 1, 11: 114 (1849). Based on oe species, the first
two being forms of See a (Royle) Johnston and the last three
forms of Arnebia densiflora Lede

Toxostigma A. Rich. Tent. Fl. Abyss. 2: 86, t. 77 (1851). Based on T. luteum
A. Rich. and T. purpurascens A. Rich., both forms of Arnebia hispidissima
(Lehm.) DC.

Leptanthe Klotzsch, Ergebn. Reise Prinz Waldemar 95, t. 63 (1862). Type
dale macrostachya Klotz., a synonym of Arnebia Benthami (Wall.)
Johnst

Phan ‘Chiov. Fl. Somala 227, t. 24, f. 1 (1929). Type species Arnebiola
migiurtina Chiov., a form of Arnebia hispidissima (Lehm.

Plants annual or perennial, herbaceous. Stems simple or loosely
branched, arising from a taproot. Leaves all cauline or some in a basal
cluster, numerous, veinless or in a few species with a few well-developed,
greatly prolonged assurgent veins that parallel the midrib. Cymes scor-
pioid, simple or forked, terminal on the main stem and leafy branches and
frequently also arising directly from the upper leaf-axils along the main
stem, few to many, loosely to densely disposed, at times aggregated in dense
corymbose or cylindric thyrsoid clusters terminating the main stems, in
age remaining densely flowered or becoming loosely flowered. Bracts nu-
merous, only rarely surpassing the adjacent calyx. Flowers heterostylic
or monomorphic. Calyx lobed nearly to the base or 5-parted, weakly to
strongly accrescent, in age sometimes developing a short swollen tube en-
closing the ripening fruit, persistent or deciduous when the enclosed fruit
is matured, usually shorter than the corolla-tube; calyx-lobes slender to
coarse, linear, ligulate, subulate or lanceolate, in age sometimes promi-
nently and coarsely veined and crested or papillate towards the base.
Pedicels short, erect. Corolla regular or nearly so, yellow, cream, blue
or purple, sometimes yellow with the limb conspicuously spotted with blue
or black, salverform or rarely tubular, outer surface hairy or at least so
on the outer surface of the limb; tube elongate, gradually ampliate or en-
larging above the middle to form a cylindric throat; limb. usually well
developed, flat or broadly funnelform, narrow to broad;. lobes strongly
imbricate in the bud, usually rounded and about as broad as long, usually
spreading or widely ascending, entire or sometimes with the margins

LY JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxv

lacerate, erose or crisped; throat inside without hairs or faucal appendages
and usually without any stipitate glands (stipitate glands when present
very scattered and few, occurring about the mouth of the corolla, present
only in some heterostylic species and usually only in the short-styled
flowers); corolla-tube glabrous within or strigulose or villulose in two
species only; annulus a papery collar, a thickened ring or completely
absent, usually hairy. Filaments linear or unguiculate, usually very much
shorter than the anther but in the short-styled flower of one heterostylic
species nearly as long as the anther, affixed in the corolla-throat all at
the same level or in one species at different levels. Anthers oblong to
narrowly oblong, affixed at or slightly below the middle, in short-styled
flowers and some monomorphic flowers borne high in the throat and fre-
quently with their tips exserted, in long-styled flowers usually borne low
in the throat and deeply included, apex emarginate or rounded or rarely
acute, base rounded or rarely with the thecae somewhat pointed at the
lower end, thecae usually joined down to the very base, connective narrow
to relatively broad, not sulcate. Pollen elongate, 25-75 x 14-43 un, in
lateral outline oblong with rounded ends, the sides straight or more or
less strongly incurving and the grains medially constricted, encircled by
a row of pores at both the upper and the lower end of the grain, upper
and lower half of the grain equal in size and configuration, polar outline
circular, pores four or five in each row, obscure; in all heterostylic species
the grains of long-styled flowers conspicuously smaller and more con-
stricted medially than those of the short-styled flowers. Style slender,
simple or simply forked or bis-bifid, included or shortly exserted; stigmas
two or sometimes four, capitate, oblong or flabellate or rarely cylindric,
simple or somewhat bilobed, juxtaposed and either strict or divergent at
the apex of the simple style, or solitary and terminating the individual
branches of the style. Nutlets one to four developing, erect, gray, brown,
fulvous, greenish or rubiginous, never white nor porcelain-like, rough,
never perfectly smooth, usually evidently tuberculate, verrucose, rugose
or rugulose, surface usually dull, frequently minutely verruculose, papil-
late or muriculate; body of nutlet usually narrowing and more or less
rostrate above the middle, sometimes conic-ovoid, conic-lanceolate or lance-
ovoid and usually broadest below the middle and longer than broad, or
sometimes more or less dorsiventrally compressed, much broadened below
the middle and as broad or nearly as broad as long and ovate, triangular-
ovate, or even cordate in dorsal outline; venter usually decidedly angulate,
keel well developed or more or less obscure or well developed only above
the middle of the nutlet; suture usually absent, when present usually ob-
scure; dorsum frequently carinate above the middle, below the middle
convex or medially depressed (in one species the dorsum plane or slightly
concave and the venter convex) ; attachment basal, large, usually flabellate
or ovate, rarely lobed, horizontal or somewhat oblique, occasionally pro-
longed upward for a short distance on the venter of the nutlet body,
plane on the truncate base of the nutlet or sometimes margined by the
downwardly prolonged pericarpial walls, rarely convex and protrudent

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVI 53

and visible even when the nutlet is viewed laterally. Gynobase flat or
broadly pyramidal; attachment surfaces separated by lineate grooves, not
margined by prominent cartilaginous tissue, plane or somewhat concave
and sometimes very strongly upcurved ventrally to form a side of a
frustum-like prominence at the center of the gynobase.

Arnebia as here defined includes the first fifteen species keyed and de-
scribed in my recent study of Lithospermum, Jour. Arnold Arb. 33: 315—
316 (1952), as well as four others discussed in a supplementary paper,
op. cit. 34: 10-15 (1953). The well-known garden plant Arnebia Echioides
(L.) DC. is excluded and assigned to the monotypic genus Echioides. As
here accepted, therefore, the genus includes most of the species that have
been traditionally assigned to Arnebia or its segregate Macrotomia. A
few species of Arnebia extend into the drier portions of northern Africa,
but most of the species in the genus are confined to Asia.

Over a year ago Arnebia was considered only in relation to Lithosper-
mum, and after an evalution of its characters it was merged with that
genus. Returning to Arnebia and Lithospermum after having critically
examined all the other genera of the Lithospermeae, I find myself judging
the two genera not merely by the number and decisiveness of characters
useful in distinguishing them, but also according to the degree of phyletic
divergence they represent as compared with that given generic recognition
elsewhere in the Lithospermeae. It is now apparent that the amplified
Lithospermum includes greater morphological extremes and is accordingly
a relatively more comprehensive unit than other genera recognized within
the tribe. Since I believe that genera within a natural circle of relationship
should be roughly equivalent in value, it now seems best to abandon my
broad concept of Lithospermum and reclassify its species under three
smaller, more homogeneous genera (Lithospermum, s. str., Arnebia, and
Echioides) that represent units of evolutionary divergence comparable with
those given generic recognition elsewhere within the tribe.

Arnebia and Lithospermum are very closely related but evidently repre-
sent diverging phyletic lines. The characters they share are more numerous
than their differences. The intimacy of their relationship is evidenced by
unusual morphological features present in precisely the same form in
both genera. Especially notable is the heterostyly in many of their species.
This heterostyly, occurring in diverse species-groups in both genera, is of
the most advanced sort, involving not merely dimorphy in corolla-form,
stamen-attachment, anther-size, style-length, and pollen-size, but also pol-
len-form. The latter feature is unknown in other heterostylic plants and
may be unique. This unusual development as an indicator of affinity be-
comes especially significant when it is recalled that heterostyly is un-
common in the Boraginoideae and that elsewhere in the Lithospermeae
it is developed in only one other genus, in a very simple form in Litho-
dora. A minor feature, also rare in this family and among the Lithosper-
meae, developed only in Arnebia and Lithospermum, is the lobulate,
Jacerate, or strongly crisped margins of the corolla-lobes. A few very
dissimilar species in both genera share this character.

54 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxv

The difficulties in separating Arnebia from Lithospermum have been dis-
cussed at length, Jour. Arnold Arb. 33: 310-315 (1952). As has been
noted, the only decisive character useful in distinguishing them involves
pollen morphology. Other possible differences have occasional exceptions
or are difficult to express. The fruit has evolved differently in the two
genera, but the differences cannot be concisely stated. The corolla-throat
in Arnebia is never decorated with faucal appendages as is frequently the
case in Lithospermum, and seldom, and then only very sparingly, does
it bear stipitate glands comparable to those always present, usually in
abundance, on the corolla-throat of Lithospermum. Some developments
present in only some species of a genus are also indicative. The annulus
inside the corolla, extremely well developed in some species of Arnebia,
is completely lacking in others. In Lithospermum the annulus, weak to
moderately well developed, is always present. The forked style, frequently
present in Arnebia, is unrepresented in Lithospermum. The subterminal
stigmas, frequent in Lithospermum, do not occur in Arnebia. The evanes-
cent dark-colored spot on the corolla limb, developed by various yellow-
flowered species of Arnebdia, occurs also in Echioides but not in Lithosper-
mum, With only one exception, all species of Lithospermum have corollas
that are yellow, orange, or white. In Arnebia the corolla has a greater
range of colors, including not only yellow and orange, but also blue, pink,
and brown, Seven of the eighteen species of Arnebia are very definitely
herbaceous annuals. Of the forty-four species of Lithospermum only two
are plants of short duration, and these may be biennials or very short-lived
perennials.

Since I discovered that Arnebia has very different pollen from that of
Lithospermum, cf. Jour. Arnold Arb. 33: 308-311, f. 1-32 (1952), I have
examined the pollen in all the other genera of the Lithospermeae. I am
now of the opinion that the differences in pollen are much more important
than previously realized. Pollen of a type similar to that of Arnebia has
been encountered only in Stenosolenium. It differs from that in all other
genera of the Lithospermeae in bearing two rows of pores, one at each
end of the grain. The grains are always elongate and are symmetrical
with the upper and lower half similar in size and configuration. In all
other genera of the tribe, including Lithospermum, the pores are in a single
row and the grains vary from symmetrical to very asymmetrical. The lower
half of the grain is frequently much larger and more rounded than the
upper half and hence different in configuration. No evidences of transition
between the two types have been detected. The pollen in Arnebia (and
Stenosolenium) has unusual and distinctive features and is the most
extreme type in the Lithospermeae.

During my study of the Lithospermeae I have become increasingly im-
pressed by the prevailing constancy as to type that is exhibited by the
pollen within the various genera of the tribe. Within a given genus pollen
may be very uniform in all species or it may present a limited number of
variations which are all modifications of a single basic type. Closely re-
lated genera usually show close similarities in pollen. Other than in the

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVI 55

amplified Lithospermum (including Arnebia), now abandoned, I found a
single type of pollen or modifications of a single type in all genera except
Moltkia. In that genus § Echianthus has globose-ellipsoidal grains with
six to eight equatorial pores. In § Eumoltkia, however, the globose grains
appear to have about twenty pores, these not lined up about the equator
but arranged, rather, in a strongly undulate line that crosses the equator
diagonally at four places. The precise description of the pollen of §
Eumoltkia is impossible with my technique, but in any case I am sure that
the pollen in this section differs more from that in § Echianthus than is
customary within the genera of the Lithospermeae. Even so, the dif-
ferences are less fundamental than those between the pollen of Arnebia
and that of Lithospermum. It is therefore not inconsistent to emphasize
the pollen differences separating Arnebia and Lithospermum and to use
them in bolstering the less decisive macroscopic differences in justifying
my present recognition of both these genera.

The eighteen species of Arnebia have been described and discussed, their
synonymy listed, and keys provided for their identification, in two previous
papers of this series, Jour. Arnold Arb. 33: 315-334 (1952) and 34: 7-16
(1953). In the reports mentioned the species were treated as members
of the genus Lithospermum. Their correct names as members of Arnebia
are given below.

Section Euarnebia.

Corolla subtubular; lobes erect, triangular, acute, longer than broad.
Stigmas four, elongate, cylindrical. Nutlets plano-convex with a cordate
base, dorsum plane or very slightly concave, venter broadly convex, lack-
ing a ventral keel; attachment surface three-lobed, the scar of the dorsal
vascular traces located in the sinus of the cordate nutlet-base much more
conspicuous than that of the funicular canal. Plants annual. Corolla with-
out annulus.

1. Arnebia tetrastigma Forsk. Fl. Aegypt.-Arab. 63 (1775); Johnston
op. cit. 321 (1952).
Section Strobilia (G. Don), comb. nov.

Corolla with a well-developed spreading limb, lobes rounded, spreading,
about as long as broad. Stigmas usually two, capitate or oblong or flabel-
late. Nutlets with a rounded back, an angulate keeled venter, and a broad
base, attachment surface not lobed, the scar of the dorsal vascular traces
less conspicuous than that of the funicular canal.

* PLANTS ANNUAL; COROLLA WITH ANNULUS.
2. Arnebia decumbens (Vent.) Coss. & a Bull. Soc. Bot. France
4: 402 (1857); Johnston, op. cit. 322 (1952).
3. Arnebia hispidissima (Lehm.) DC. Prodr. 10: 94 (1846); Johnston,
op. cit. 325 (1952).

56 JOURNAL OF THE ARNOLD ARBORETUM __ [voL. xxxv

. Arnebia Griffithii Boiss. Diag. ser. 2, 3: 135 (1856); Johnston,
op. cit. 326 (1952).

as

un

Arnebia fimbriopetala Stocks in Hook. Jour. Bot. & Kew Miscl.
3: 180, t. 6 (1851); Johnston, op. cit. 7 (1953).

ON

. Arnebia minima Wettst. in Stapf, Denkschr. Acad. Wiss. Wien 50:
30 (1885); Johnston, op. cit. 327 (1952).

~I

Arnebia linearifolia DC. Prodr. 10: 95 (1846); Johnston, op. cit.
328 (1952).

** PLANTS PERENNIAL.
x COROLLA WITH ANNULUS.

oo

Arnebia fimbriata Maxim. Bull. Acad. St. Petersb. ser. 3, 27: 507
(1881); Johnston, op. cit. 328 (1952).

oO

. Arnebia obovata Bunge, Mem. savants étrang. St. Petersb. 7: 407
(1851); Johnston, op. cit. 329 (1952).

_
oO

. Arnebia Szechenyi Kanitz, Pl. Exped. Szecheny 42, t. 5 (1891);
Johnston, op. cit. 329 (1952) and 8 (1953).

—
—s

Arnebia guttata Bunge, Ind. Sem. Hort. Dorpat. p. vii (1840) ;
Johnston, op. cit. 330 (1952).

—
bo

Arnebia Lindbergiana (Rech. f.), comb. nov.
Macrotomia Lindbergiana Rech. f. Ann, Naturhist. Mus. Wien 58: 58 (1951);
Johnston, op. cit. 10 (1953).

<x COROLLA WITHOUT ANNULUS.

13. Arnebia densiflora Ledeb. ex Nordmann, Bull. Acad. St. Petersb.
2: 312 (1837); Johnston, op. cit. 331 (1952).

14. Arnebia inconspicua Hemsl. & Lace, Jour. Linn. Soc. 28: 326
(1891); Johnston, op. cit. 12 (1953).

15. Arnebia euchroma (Royle) Johnston, Contr. Gray Herb. 73: 49
(1924); Johnston, op. cit. 333 (1952).
16. Arnebia Benthami (Well. ex G. Don), comb. nov.

Eichium Benthami Wall. Numerical List no. 931 (1829), nomen; G. Don,
en, Syst. 4: 333 (1838); Johnston, op. cit. 333 (1952).

17. Arnebia speciosa Aitch. & Hemsl. Proc. Linn. Soc. 18: 81 (1880) ;
Johnston, op. cit. 13 (1953).

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVI a7

Arnebia nobilis Rech. f., Ann. Naturhist. Mus. Wien 58: 58
(1951); Johnston, op. cit. 14 (1953)

—_
=

—
—

Echioides Ortega, Tabulae Botanicae 7 (1773); Johnston, Jour.

Arnold Arb. 33: 314 (1952). Type species, Arnebia Echioides (L.)

DC.
Aipyanthus Stevens, Bull. sa Nat. Moscou 24: 599 (1851). Type species

Arnebia Echioides (L.) D
Plant perennial. Stems es erect, simple below the inflorescence.
Leaves borne along the stem and in basal clusters, uppermost ones with
rounded or subcordate sessile bases, basal ones oblanceolate; midrib prom-
inent but veins usually evident only on the basal leaves. Cymes scorpioid,
simple or forked, terminal on the stems, in age elongating, becoming
straight and racemose. Bracts numerous and conspicuous, foliaceous,
usually evidently surpassing the adjacent calyx, lanceolate, asymmetric
with the base rounded or subcordate. Calyx lobed almost to the base;
lobes moderately unequal, cuneate-linear to narrowly lanceolate, shorter
than the corolla-tube, accrescent; pedicels short, stout, erect. Flowers
strongly heterostylic. Corolla yellow, villulose outside, with an elongate
cylindric or weakly ampliate tube which towards its summit abruptly
expands into the short open throat and the spreading limb; limb with a
diameter about equaling the length of the tube, bearing five evanescent
brown or blackish spots, one at the base of each sinus; lobes widely
spreading, rounded, about as broad as long; throat open, shallow, glabrous,
without stipitate glands or faucal appendages; tube villulose inside, with-
out glands; annulus absent. Stamens borne in the tube at several super-
imposed levels, in the long-styled flowers within a zone below the middle
of the tube and in short-styled flowers in a zone just below the summit
of the tube; filaments linear, very short, one fourth to one fifth the length
of the anther; anther narrowly oblong, affixed at or just below the middle,
emarginate at base and apex, dorsum with a relatively broad connective
that is sulcate above the middle; thecae separate for a short distance above
their bases but remaining parallel and juxtaposed. Pollen spherical or
very slightly longer than broad, bearing nine obscure pores about the
equator, 30-35 p» in diameter in short-styled flowers and 40-50 » in long-
styled flowers. Style slender, half as long or as long as the corolla-tube,
somewhat thickened directly below the terminal stigma; stigma deeply
bilobed, the lobes joined at the base and becoming divergent. Nutlets
large, very plump, ovoid or elliptic-ovoid, prominently and narrowly keeled
on the venter but elsewhere convex, tawny and finely mottled with purple,
opaque, smooth or sparsely and broadly low-tuberculate, ventral suture
absent; attachment scar basal, large, plane, usually margined by the
weakly protrudent pericarpial walls. Gynobase broadly pyramidal, the
attachment faces flabellate, plane, sloping, margins not upturned nor
thickened, adjacent faces separated by a groove

A monotypic genus native to Armenia, the Caucasus, and adjoining
northern Iran.

58 JOURNAL OF THE ARNOLD ARBORETUM _ [vot. xxxv
Echioides longiflorum (C. Koch), comb. nov.

Arnebia longiflora C. Koch, Linnaea 20: 640 (1849).

Lycopsis Echtoides L. Sp. Pl. ed. 2, 199 (1762).

Arnebia Echioides (L.) DC. Prodr. 10: 96 (1846).

Aipyanthus Echioides (L.) Stevens, Bull. Soc. Nat. Moscou 24: 600 (1851).
Arnebia cyrousiana Parsa, Kew Bull. 1948: 211 (1948) and Fl. ’Iran 47: 221
(1952).

Lithospermum cyrousianum (Parsa) Johnston, Jour. Arnold Arb, 34: 7 (1953).
Lithospermum Tournefortii Johnston, Jour. Arnold Arb. 34: 7 (1953).

The present plant has been traditionally classified as an Arnebia, this
with considerable justification, for indeed it does have many similarities
with that genus. Its pollen, however, differs extremely from that of
Arnebia, being very similar in type to that characteristic of Lithospermum.
In this regard the plant destroys the universality of the one set of char-
acters that might be used in separating Arnebia and Lithospermum in a
decisive manner. Because decisive characters could not be found, I re-
cently merged Arnebia and Lithospermum, Jour. Arnold Arb. 33: 311-315
(1952). The amplified Lithospermum produced I now recognize is more
heterogeneous than other genera of the Lithospermeae. In order to re-
establish Arnebia and have it separated from Lithospermum by at least
one crucial character, and at the same time have both genera homogeneous,
it has been necessary to exclude the present plant from both of the genera
mentioned. .

Although Echioides shares many characters with Arnebia and Litho-
spermum it differs from both in the attachment of the stamens in its
strongly heterostylic flowers. The stamens within a flower are borne at
several different altitudes above the corolla-base, being irregularly dis-
tributed within a broad zone inside the corolla-tube. This staminiferous
zone is located below the middle of the corolla-tube in long-styled flowers
and just below its summit in the short-styled ones. In Arnebia the stamens
may be borne at unequal distances above the corolla-base in A. decumbens,
but that is not a heterostylic species. In both Arnebia and Lithospermum
the stamens in all heterostylic species are always whorled. In having the
irregularly distributed stamens confined to a zone on the corolla that varies
in position according to whether the individual is long- or short-styled,
the present plant differs from all other members of the Boraginaceae and
may even be unique among all heterostylic plants.

The corolla of Echioides as to general form is more suggestive of species
of Lithospermum than of those of Arnebia. The corolla-tube, in having
hairy inner surfaces, finds its closest parallel among American species of
Lithospermum. In having a naked throat the corolla agrees with that of
Arnebia and is very different from that of Litkospermum, for in the latter
genus the throat is always decorated with faucal appendages or stipitate
glands. The complete absence of the annulus is also suggestive of Arnebia.
The annulus is suppressed in some species of Arnebia but is always present
in some form in all species of Lithospermum. Most suggestive of Arnebia,
however, are the five transitory dark spots decorating the yellow corolla-

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVI 59

limb, a very attractive feature usually present, which Echioides shares
with only four or five species of Arnedia.

The large plump nutlets suggest those of Lithospermum in form but
differ from that genus in their tawny mottled opaque surface. Their
surface features are more like those of nutlets in Arnebia. The pericarp
is not dotted with pits, nor does it bear a row of pits and slots paralleling
each side of the ventral keel. These are features regularly present on the
nutlets of many species of Lithospermum but never developed in Arnebia.
The attachment faces on the gynobase are flat and separated by a groove.
They do not have thickened margins, nor are they separated by prominent
cartilaginous ridges. The gynobase, accordingly, is of a type universal
in Arnebia but relatively uncommon in Lithospermum. The pollen grains
of Echioides are spheric and have nine pores arranged about the equator.
They are, hence, extremely different both as to form and type from the
elongate grains with biseriate pores which are universal in Arnebia. Al-
though much larger than those of any species of Lithospermum, the grains
of Echioides are the same as to type. In form and general organization
they are duplicated by several species of Lithospermum.

12. Lithospermum L. Sp. Pl. 132 (1753) and Gen. Pl. 64 (1754). Type
species L. officinale L.
Batschia Gmel. Syst. 2: 315 (1791); Rehder, Kew Bull. 1935: 396 (1935).
d on Anonymos caroliniensis Walter, Fl. Carol. 91 (1788), which
represents Lithospermum caroliniense (Walt.) MacMill.

Cyphorima Raf. Am. Monthly Magazine 4: 191 and 357 (Jan. and March,
1819) and Jour. de Phys. Chem. Hist. Nat. 89: 98 (Aug. 1918); Merrill,
Ind. Raf. 202 (1949). “. .. le type du genre est le Lithospermum lati-
folium de Linné [L. latifolium Michx.!]. Les Batschia longiflora et
decumbens (Nuttall) doivent peut-étre s’y rapporter?”

Pentalophus A. DC. Prodr. 10: 86 (1846). Based on L. longiflorum Spreng.
and L. mandanense Spreng., both forms of L. incisum Lehm.

Plants distinctly perennial and arising from a strong, frequently dye-
stained taproot or rarely short-lived and springing from a slender biennial
or perhaps even annual root. Stems herbaceous or rarely somewhat frutic-
ulose, simple or more or less branched, hispid, villose or strigose, erect or
spreading, short to elongate. Leaves usually numerous, sometimes pin-
nately veined but more commonly veinless, occasionally in a sterile basal
cluster but usually all cauline. Cymes scorpioid, simple or geminate, few-
to many-flowered, borne terminal on the stems and branches or sometimes
also in the uppermost leaf-axils and becoming aggregated into a cylindrical
thyrse, usually elongating, straightening and becoming racemose in fruit.
Bracts usually numerous and surpassing the adjacent calyx, foliaceous.
Calyx 5-fid, the lobes linear-cuneate or lanceolate, short to elongate,
usually evidently shorter than the corolla-tube, the abaxial one usually
appreciably the largest; pedicels very short to elongate, strict or ascending
and rarely decurved. Flowers monomorphic or strongly heterostylic, rarely
cleistogamic. Corolla regular, yellow, orange, or white (or pinkish or bluish

60 JOURNAL OF THE ARNOLD ARBORETUM _ [VvoL. xxxv

to purple-red in one Chinese species), hairy on the outer surface, salver-
form, funnelform, or cylindric; tube straight, usually elongate, subcylindric
or gradually ampliate or more commonly enlarging at or above the middle
to form a differentiated throat; limb usually spreading or broadly ascend-
ing, small to large, its diameter less than, equal to, or surpassing the length
of the corolla-tube; corolla-lobes equal, strongly imbricate in the bud,
usually orbicular to ovate but sometimes semicircular or obovate, spread-
ing or loosely ascending or rarely strictly ascending and nearly erect,
margins entire or rarely lacerate, crisped or erose; corolla-throat frequently
bearing small invaginate pubescent or glanduliferous appendages and
always bearing some stipitate glands either generally distributed or local-
ized in a band or in congregations; faucal appendages when present weak
to well defined, trapeziform, lunate, gibbose, or merely convex; corolla-
tube glabrous inside or sometimes distinctly hairy; annulus usually evident
and usually hairy, a tumid band, a narrow ring, or represented by ten
quadrate or gibbose lobes or rarely reduced to 5 to 10 tufts of hairs. Sta-
mens arising at or usually well above the middle of the corolla-tube, borne
below the middle of the tube only in long-styled flowers of some hetero-
stylic species. Filaments unguiculate or linear, usually half the length of
the anther or less, longer than the anther in one species only, all affixed
at the same height above the corolla base, usually hidden behind the
anther, never exserted, glabrous and glandless or sometimes bearing a few
glands on the decurrent base. Anthers small, oblong to narrowly oblong,
usually compressed laterally, included or rarely with the tips exposed in
the corolla mouth, rounded at both ends or with the tip obscurely apiculate,
affixed at or near the middle; connective on dorsum usually narrow; thecae
united to the base or free for only a very short distance above the base,
parallel and closely juxtaposed throughout. Pollen isodiametric or more
commonly longer than broad, 13-42 » long and 8—42 ,» broad, spheric or
barrel-shaped or ellipsoid with the upper and lower halves of the grain
equal in size and configuration, or ovoid or medially constricted with the
upper and lower halves very dissimilar in size and configuration, in hetero-
stylic species grains of long- and short-styled flowers differing in size and
frequently also in form; pores 6 to 9 but most commonly 8, borne in a
single row about the equator or about the lower half of the grain where it
is broadest. Nutlets erect, usually ovoid or ellipsoid and plump, broadest
at or below the middle, gradually narrowed to the base or sometimes
suprabasally constricted to form a stout neck just above the broad flaring
base or sometimes with a thickish collar about the base; apex usually obtus-
ish, dorsum convex (flattened below the middle in one species) ; venter con-
vex or somewhat obtuse, its medial keel rarely sharp, usually low and
rounded and sometimes obscure, the ventral suture lineate or obscure; sur-
face of nutlet usually polished and porcelain-like, white or somewhat stained
with brown, rarely verrucose or rugose or tumulose, usually completely
smooth or smooth with few to numerous punctate depressions, the depres-
sions sometimes shallow but usually deep and pit-like and frequently on the
venter of the nutlet elongating to become slot-like, a well-defined row of

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVI 61

slots or pits or both frequently present on both sides of the ventral keel
and parallel to it; attachment of nutlet large, basal, plane, convex or in
one species excavated, horizontal or somewhat oblique, the end of the
broken funicular canal usually evident near the ventral angle, near the
middle of the dorsal half of the attachment the ends of the dorsal vascular
supply marked by a dot, a swelling, or by an erect or ascending subulate
or peg-like process. Style slender, simple, included in the corolla-tube or
sometimes exserted; stigmas 2, usually distinct, globose, hemispheric or
ellipsoid, usually small, terminal and juxtaposed on the tip of the style
or subterminal and separated by the short, somewhat bilobed sterile tip
of the style. Gynobase flat to broadly pyramidal, sometimes surmounted
by a thickened angulate persisting base of the style; attachment faces flat
or slightly concave (or saucer-shaped or cupulate in one species), hori-
zontal or ascending, plane or the margins somewhat thickened and promi-
nent, separated by a groove or by the thickened protrudent angles of the
gynobasic pyramid.

In the past the genus Lithospermum has been very broadly defined. As
one of the relatively few original Linnean genera of the family it early
became a catch-all for a great variety of plants subsequently recognized
as generically separable. During the past century its history has been
one of shrinking limits. As here defined the genus is further shrunk and
I believe at last reduced to homogeneity. Various diverse elements still
included in the genus by recent classifiers I have excluded and treated
under such genera as Buglossoides, Lithodora, Psilolaemus, Moltkia, Molt-
kiopsis, Mairetis, and Neatostema. Also excluded are Echioides and
Arnebia, recently included in the genus in my “Survey of the Genus Litho-
spermum,” Jour. Arnold Arb, 33: 299-363 (1952). My reasons for now
excluding these two latter will be found discussed under Arnebia.

Lithospermum, as here accepted, includes forty-three of the fifty-nine
species treated in my recent paper on the genus. The first fifteen species
in the ‘‘Survey” are now referred to Arnebia, and species no. 17 (L. Tour-
nefortii) to Echioides. 'To be added to the forty-three species keyed and
described in the “Survey” is one other species that was improperly ex-
cluded. This is L. cinerascens of Peru and Ecuador, which will be dis-
cussed presently. The genus Lithospermum as now defined contains,
accordingly, a total of forty-four species.

The genus is especially notable for its floral dimorphism and its di-
versity of pollen forms. For a discussion of these and various other details
the reader is referred to the above-mentioned “Survey.”

Lithospermum, among all the genera of the Lithospermeae, is the only
one with representatives on all continents. Of the forty-four species, four
(L. officinale, L. erythrorhizon, L. tschimganicum, and L. Hancockianum)
are native to Eurasia, one is confined to the highlands of tropical Africa
(L. afromontanum), four occur in South Africa (L. papillosum, L. di-
versifolium, L. cinereum, and L. scabrum), and five are indigenous to the
highlands of northwestern South America (L. cinerascens, L. Macbridei,
L. mediale, L. peruvianum, and L. Gayanum). The remaining thirty

62 JOURNAL OF THE ARNOLD ARBORETUM _ [vot. xxxv

species occur in the United States and Mexico, with the greatest concen-
tration in the latter country. Among the species in the Old World the
most distinct is LZ. Hancockianum of southwestern China. The other
species of the Old World have not only the technical characters, but the
facies of the species in America, and were they natives of the New World
would be accepted in Lithospermum without question. The interrelation
of the species of Europe, Africa, and America is very clear and strong, and
the genus, despite its wide distribution, is a very satisfactory and natural
assemblage

The genus is not only notable for its wide distribution and particularly
for its occurrence in both Old World and New but also for the number of
other genera with which it seems to be closely related. It is the only
genus in the Lithospermeae which has close generic affinities in both
America and the Old World. The genus has affinity with all the genera
of the tribe native in America and is very closely related with Macromeria,
Perittostema, and Psilolaemus. In the Old World its closest affinities are
with Arnebia, Echioides, and Buglossoides, and perhaps also Lithodora.
Its affinities with other Old World genera are only very generalized,
certainly not immediate.

The wide distribution of the genus and the number and distribution of
genera Closely related to it suggests that perhaps Lithospermum may be
a relatively generalized conservative old stock from which many of the
modern Lithospermeae may have been differentiated. Such a conclusion,
however, is difficult to reconcile with the fact that Lithospermum in many
features shows specializations not to be expected in a primitive group.

he modification of the corolla-throat is noteworthy in this regard. In
other tribes of the Boraginoideae, all without doubt containing more highly
evolved plants than the Lithospermeae, faucal appendages, usually in-
vaginate and frequently very elaborate, are features of the corolla in prac-
tically all genera. It is interesting to note, therefore, that among the
twenty-three genera of the Lithospermeae, well-developed faucal append-
ages are developed only in Perittostema and in various species of Litho-
spermum. Elongate plaits in the throat, possibly homologous with the
faucal appendages of other genera, occur only in Buglossoides and Mac-
romeria, both closely related to Lithospermum. Stipitate glands in the
corolla-throat, localized and usually abundant, which appear to have some
relation with the development of faucal appendages in the Lithospermeae,
occur only in Lithospermum and its close relatives (Macromeria, Lasiar-
rhenum, Perittostema, Arnebia, and Lithodora). The corollas of most
genera of the tribe have no faucal appendages and bear no stipitate glands
in the throat. Among genera evidently related to Lithospermum this is
also the case in many or all the species of Macromeria, Onosmodium, Psilo-
laemus, Nomosa, Lithodora, Arnebia, and Echioides. Modifications of
the corolla-throat are features developed only in Lithospermum or to some
degree only in genera evidently related to it. As a complex elaboration
of the corolla, it seems to indicate that the genus is a highly specialized
member of the Lithospermeae and not a generalized conservative one.

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVI 63

A high position in the evolution of the Lithospermeae is also indicated
for Lithospermum by the elaborate development of heterostyly among
various groups of its species. Heterostyly of a precisely similar sort is
also present among certain groups of species in Arnebia and in a slightly
more complicated state in Echioides. Elsewhere in the tribe it is developed
only in Lithodora, and there in only a very simple form. The condition,
accordingly, occurs in the Lithospermeae only in Lithospermum and in
genera evidently related to it. Heterostyly, like the elaborations of the
corolla, gives reason for believing that Lithospermum is an advanced
member of its tribe. .

The nutlets of Lithospermum are prevailingly plump, ovoid or ellipsoid,
and usually obtusely pointed, and have a convex dorsum and a venter that
is merely obtusely angled or obtusely angled with a low rounded medial
keel. Nutlets with a narrow sharp ventral keel occur only in a few species,
e.g., L. strictum and L. Hancockianum. Only in L. strictum is the dorsum
of the nutlet distinctly flattened. The surface of the nutlet is prevailingly
lustrous and porcelain-like, white or somewhat stained with brown. It is
completely smooth or smooth with scattered pits usually most abundant
on the ventral surface. Tumulose, rugose, or verrucose nutlets occur only
in L. papillosum and L. cinereum of South Africa and in L. Parksii and
L. mirabile of Texas and adjacent Mexico. The only species with smooth
nutlets that are not lustrous is L. indecorum.

A distinctive minor feature of the nutlets of Lithospermum, evident to
some degree in most species of the genus, is the two vertical lines of deep
pits or deep slots in the pericarp which parallel the ventral keel. One of
these lines of depressions is usually present along each side of the keel,
close to its lateral base or only a short distance removed. Outside of Litho-
spermum the lines of pits or slots associated with the ventral keel have
been observed only in Psilolaemus and in one species of Onosmodium,
O. virginianum. Possibly they may also occur on the nutlets of Nomosa
or Perittostema, but at present the fruit of these genera is unknown. In
any case they do not occur in Arnebia, Echioides, Macromeria, and Lasiar-
rhenum. In Buglossoides they are not developed in § Margarospermum,
though possibly comparable developments may be present in § Eudbuglos-
soides. Interestingly, they are present in L. Hancockianum, that very
distinct species of unusual habit native to eastern China, which, of all
species of Lithospermum, is most suggestive of Arnebia.

The nutlets usually narrow gradually towards the broad base or towards
a constriction just above the base. In some species the suprabasal con-
striction forms a stout neck just above the flaring base of the nutlet. In
others the suprabasal constriction is weak, but below it the nutlet is
modified and has a more or less collar-like base. The scar is usually broad,
distinctly basal, and horizontal or somewhat oblique. Towards the ventral
edge of the scar the broken end of the funicular canal is usually evident
either as a pit or as a short section of protruding tube. The lower end of
the vascular supply to the dorsum of the nutlet may be undifferentiated
or be marked by a small convexity near the middle of the dorsal half of

64 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXXV

the scar. Commonly, however, the tissue about the ends of the dorsal
traces protrudes to form a small process on the scar. It is usually ascending
and subulate in form but sometimes is peg-like and erect. Although in
various genera related to Lithospermum there is frequently a modest eleva-
tion of tissue about the dorsal traces, when present it is usually less ex-
treme than that to be encountered on the base of the nutlets in many
species of Lithospermum. Indeed, outside of Lithospermum well-developed
processes were noted only on the nutlets of Macromeria viridiflora. The
most extreme elevation of tissue about the dorsal traces occurs on the
nutlet base of L. incisum. In that species the process is relatively coarse,
peg-like, and vertical, and because the surface of the scar collapses it
gains in apparent height and becomes conspicuous in the excavated lower
end of the nutlet. A similar, less extreme development of the peg-like
process also occurs on the nutlet of the related species L. Parksii. The
feature is probably a compensatory development associated with the elabo-
ration of the broad collar-like base distinctive of the nutlet in these species,
although strangely in L. mirabile, the other close relative of L. incisum,
the nutlet base is flat and the process is very weak and short. The well-
developed peg on the nutlet attachment of L. incisum is associated with
another unusual feature of that species. When the nutlet detaches, the
withdrawal of its peg leaves a central depression on the attachment face
of the gynobase. After the fall of the nutlet the attachment face shrinks
and assumes a cupulate form that is unique in this genus. Although the
peg on the nutlets of ZL. Parksii is almost as prominent as that in L.
incisum, its fall leaves no central depression on the attachment faces of the
gynobase. In L. Parksii the faces remain permanently plane and are similar
to those of other species in the genus.

A species deserving of special comment is Lithospermum cinerascens
(DC.) Johnston, Contr. Gray Herb. 75: 40 (1925), originally described
as Macromeria cinerascens DC. Prodr. 10: 69 (1846). This plant, a native
of the mountains of Ecuador and northern Peru, was excluded from Litho-
spermum in my recent study of that genus. I have subsequently had all
the representation of the species preserved at Kew and the British Museum
(collections of Mathews, Jameson, and Lobb) available for close study and
comparison, and am now of the opinion that the species is closely related
to L. guatemalense of northern Guatemala and adjacent Mexico, and con-
generic with it. The South American plant is distinguishable from L.
guatemalense and all other members of its genus by its elongate filaments,
these being once and a half to twice as long as the anthers. The corolla
appears to be rather variable as to size and form. It is 34.5 cm. long.
The elongate tube is gradually expanded for most of its length or it swells
and becomes somewhat cylindric above the middle. Just below its summit
the tube abruptly expands and is there somewhat funnelform. From its
edges arise the short (3.5—-5 mm.) ovate or ovate-triangular corolla-lobes.
The lobes are usually ascending, but in one collection of Lobb there is
indication that they may sometimes be reflexed. The anthers are borne
in the open funnelform mouth of the tube, their tips barely if at all pro-

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVI 65

truded from it. The slender filaments, 3-5 mm. long, arise 4-6 mm. below
the tube summit. The medio-affixed anthers, 2-3 mm. long, are oblong
and bear a pair of minute tips on their rounded summits. The thecae are
joined to the very base. The corolla-tube bears scattered stipitate glands
in a band 2-4 mm. broad just below the tube summit. There are no faucal
appendages. The annulus consists of ten quadrate glabrous lobes. The
lustrous white nutlets are sparsely punctate and bear pits and slots in a
line on both sides of the ventral keel. The stigmas are subterminal and
are separated by the very short and stout bilobed sterile apex of the style.
The pollen is broadly ellipsoid, 25-28 22-24 y, and has eight obscure
pores on the equator. In large-flowered plants the trumpet-shaped corolla
of the species is suggestive in size and form of that of the genus Macro-
meria. Though the filaments are more elongate than in all other species
of Lithospermum, they are definitely included within the corolla-throat,
as is characteristic in that genus. They are not exserted from the corolla-
throat, as are the filaments in all species of Macromeria. The anthers
are also included, or at most have only their tips exserted. The well-
developed annulus in the corolla is that of Lithospermum and not Macro-
meria. The nutlets also have features of Lithospermum. They are punc-
tate and bear pits and slots in a row on either side of the ventral keel,
which is never the case in Macromeria but is frequent in Lithospermum.
The plant gives no evidence of being closely related with any particular
species of Macromeria. It does, however, have an evident ally in Litho-
spermum in L. guatemalense. Between Macromeria and Lithospermum,
the plant clearly belongs in the latter genus.

13. Cerinthe [Tournef.] L. Sp. Pl. 136 (1753) and Gen. Pl. 66 (1754).
Type species C. major L.

Annual, biennial, or perennial herbs, glabrous or nearly so, sometimes
glaucescent. Stems leafy, one to numerous, simple or branched, usually
erect, Leaves all cauline or some in basa! clusters, glabrous or at most
with very scattered appressed hairs or a ciliolate margin, with inconspicu-
ous pinnate veins, upper surface usually dotted with pallid groups of
mineralized cells; lower cauline leaves more or less oblanceolate but usually
broadened at the very base and usually with a somewhat amplexicaul
attachment; middle cauline leaves oblong or ovate, sessile, with a broad
cordate-auriculate amplexicaul base, apex usually obtuse or rounded.
Inflorescence loosely scorpioid, elongating and unilaterally racemose in
age; cymes single or geminate at the ends of the stems and branches;
bracts very well developed, foliaceous, very conspicuous, sometimes pur-
purescent, usually hiding the calyx, base cordate-auriculate and amplexi-
caul. Calyx 5-fid; lobes very unequal, strongly imbricate, foliaceous or
somewhat membranous, glabrous or with margin ciliolate, usually accres-
cent in age, outermost sepal broadest and usually somewhat cordate at
base; pedicel slender, glabrous or short hispidulous, in age elongate, as-
cending or decurved. Corolla yellow or partially purple or violet, com-
pletely glabrous, coarsely tubular, several times longer than broad, broadest

66 JOURNAL OF THE ARNOLD ARBORETUM _ [vot. xxxv

at or above the middle, straight and regular or most prolonged on the two-
lobed adaxial side with the tube ventricose and the limb more or less
distinctly oblique; throat without appendages, hairs or stipitate glands;
annulus evident, a narrow thickish glabrous ring or collar; lobes broadly
to narrowly triangular, very short or as much as two thirds the length of
the tube, more or less recurved or sometimes erect or even connivent, acute
or the tip somewhat rounded. Filaments affixed above the middle of the
corolla-tube, glabrous, usually thickish and narrowed from a broad base,
equal or within the corolla differing slightly in shape (odd medial stamen
adaxial), less than a quarter the length of the anther to nearly as long.
Anther elongate, affixed above the base, included in the throat or more
than half exserted from it, free or joined only by entangling of the tails
on the thecae, terminated by an evident gradually narrowed sterile ap-
pendage; terminal appendage usually with a narrow tip, frequently with
a denticulate margin; thecae with minutely papillate-ciliolate edges, near
the base remaining separate and tending to spread and thus form the more
or less sagittate base of the anther; basal end of theca sometimes with only
a short stout thickened tip but more commonly with the tip prolonged
into a slender flexuous barbellate tail; back of anther pale or frequently
black, more or less coarsely muricate or muricate-papillate, connective
usually smooth and only moderately prominent. Pollen barrel-shaped or
ellipsoid to nearly spheric, 18-37 « 15-33 w, polar profile circular or
somewhat polygonal; pores eight, borne in a well-developed equatorial
groove, rarely prominent; colpi eight, fusiform or linear. Ovary four-lobed,
four-ovulate. Nutlets one or two, each two-celled (one-celled or one-
ovulate only by abortion), smooth, rarely lustrous, usually mottled with
brown or purple, erect or weakly incurved, more or less broadly ovoid,
back convex, apex notched and bicuspid, venter flattened and not keeled;
suture linear-sulcate, evident on the venter of the nutlet and frequently
also down the dorsum and rarely even across the attachment scar; at-
tachment broad, basal, plane. Gynobase strongly depressed, when fully
developed bearing two semicircular to near circular attachment faces which
are horizontal or are slightly inclined away from the base of the style.
Style slender, eventually exserted from the corolla, dorsiventrally com-
pressed at the base; stigmas two, small, terminal, juxtaposed, distinct or
more or less united.

A very distinct and readily recognizable genus with representatives oc-
curring from middle Europe and northwest Africa east to the Caucasus
and northern Iran. No detailed comprehensive treatment of its species
has been published. The genus contains at least three or four major
species and also a goodly number of well-marked geographic varieties that
are accepted as species by many authors. Cerinthe has its greatest con-
centration of forms in the western and northern portions of the Mediter-
ranean area.

Among the Boraginoideae Cerinthe is notable for its practically glabrous
herbage, its usually broad leaves and bracts that are cordate-amplexicaul
at the base, its very strongly unequal broadly imbricate calyx-lobes, and

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVI 67

especially its bilocular two-seeded nutlets. The plant is obviously a mem-
ber of the Lithospermeae, and though very highly specialized and distinct,
probably has its closest affinities with Onosma and related genera.

In general structure and appearance the corolla and androecium are
very similar to those of Onosma and Podonosma. The corolla-lobes may
be short and recurving as prevalent in Onosma or elongate and erect as in
Podonosma. Although the corolla is completely glabrous and thus different
from many of the species in Onosma, corollas that are glabrous or very
scantily pubescent are found in Podonosma and in some species of Onosma.
The stamens of Cerinthe show a prevailing basic similarity with those of
Onosma and its relatives. The elongate anthers are terminated by a sim-
ilar terminal appendage. They are also roughened on the back, and their
thecae are also free and more or less spreading at the base. The lower
end of each theca may have a thickened point as in Onosma or, unlike in
that genus or in any of its allies, may be prolonged into a slender flexuous
barbellate tail. Unlike in the genera mentioned, the anthers are joined
neither by coherence at the base of the thecae nor by coherence along the
margin of the thecae. The anthers of Cerinthe, though regularly free in
some species and varieties (as in Onosma), in most species are held to-
gether at the base — not by growing together, but only by the entangling
of the flexuous tails on the thecae. The synandrium produced, though not
achieved by adnation, functions exactly like that in those species of
Onosma which have only the basal tips of their thecae coherent.

The corolla of Cerinthe may be perfectly regular or have a more or less
clear bilateral symmetry. In C. retorta Sibth. & Sm., and to a less extent
in some of the forms of C. major L. and its allies, the corolla may be more
or less distinctly ventricose, not only above the base on the abaxial side,
but also above the middle of the adaxial side. When this is relatively well
developed, as in C. retorta and C. gymnandra var. macrosiphonia Mutb.,
the unilateral swellings can so modify the form of the corolla that it be-
comes weakly curved in a sigmoid manner. In the two plants mentioned,
the corolla is slightly prolonged on its two-lobed adaxial side, and the
upper portion of the tube is somewhat outcurved and the limb is oblique.
Associated with this zygomorphy of the corolla is also some differentiation
among the filaments inside. In C. gymnandra var. macrosiphonia, further-
more, the exserted cone of connivent anthers is not central and vertical in
the mouth of the corolla, but rather, emerging from the abaxial side of
the mouth, is directed obliquely upward, with the tip over the opposite
side of the corolla-limb. The flowers of C. retorta and C. gymnandra var.
macrosiphonia have an evident bilateral symmetry that has involved not
only the corolla but also the androecium. Among the many species of
Onosma the only parallel to the conditions described in these species of
Cerinthe is that in O. multiramosum H.-M. of China. In that species the
matured corollas in the bud and the exserted synandrium in the open
flowers are prolonged on the adaxial side and are strongly out-curved,
i.e., bent away from the cyne axis in a manner suggestive of the subapical
out-curving of the corolla observable in Cerinthe.

68 JOURNAL OF THE ARNOLD ARBORETUM __ [voL. xxxv

The fruit of Cerinthe differs from that of all other Boraginoideae in
having bilocular two-seeded nutlets and these never more than two in
number, The two carpels of the fruit, which in other Boraginoideae are
each divided to form a pair of distinct one-celled one-seeded nutlets, are un-
lobed in Cerinthe. The nutlets are superficially similar to the two-seeded
nutlets produced by some species of Heliotropium and are comparable also
to the two-seeded pyrenes of the drupaceous fruits of some Ehretioideae.
For the proper interpretation of the fruit of Cerinthe the evolutionary steps
in the development of fruit within the Boraginaceae should be recalled. The
fruit in this family has been elaborated from a two-carpellate, two-celled
ovary bearing four ovules which become four-celled by the development
of a partition within each carpel, and then, subsequently, become two-
lobed and then four-lobed, and finally in the Boraginoideae become four-
parted, with each lobe (the nutlet) unilocular and one-seeded and affixed
independently about the base of the style. Since the two-seeded nutlet
of Cerinthe represents a whole carpel, and the one-seeded nutlets of all
other members of the Boraginoideae only half of a carpel, it is perhaps
not surprising that the fruit of Cerinthe has been interpreted by some
authors as being not only the most simply but also the most primitively
organized within the subfamily, and even as more or less transitional to
the less highly evolved fruits in the Heliotropioideae and Ehretioideae.
With such a conclusion, however, I cannot agree. Along with most au-
thors, I am of the opinion that the two-seeded nutlets of Cerinthe are
simply single-seeded nutlets which somehow have failed to develop sep-
arately during ontogeny. Cerinthe, like other genera of the Lithosper-
meae, no doubt had immediate ancestors with a typical boraginoid fruit
consisting of four discrete single-seeded nutlets. In the present genus, the
union of the four nutlets into two double nutlets, although presenting a
condition suggestive of a primitive one, is actually a late secondary de-
velopment and accordingly a highly specialized one.

The ovary of Cerinthe, viewed shortly after anthesis, is distinctly four-
lobed and is generally similar in appearance to that of most Lithospermeae
at the same stage of development. The pair of lobes (those derived from
one carpel) which will develop into a bilocular nutlet are at this stage
united only below the middle, usually for no more than one third their
total length. The depth of separation between the lobes within each pair
is evidently less than that between the two pairs. Serial transverse sec-
tions through the base of the ovary will show the presence of only a basal
zone in which the ovarial lobes are united in pairs. Only if in its subse-
quent development growth is largely confined to this basal zone can the
ovary of Cerinthe produce bilocular united nutlets. Growth centered above
this zone would produce single-seeded nutlets as in other Lithospermeae.
In Cerinthe growth in the upper zones of the ovary is retarded. The free
tips of the ovarial lobes are represented in the mature nutlets only by the
bicuspid apex.

The nutlets of Cerinthe, when perfectly developed, are two-celled and
two-ovulate. Single-seeded one-celled nutlets may be found, but these

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVI 69

always show some evidence of being imperfectly developed. In some uni-
locular nutlets the undeveloped half may be reduced to a narrow longi-
tudinal ridge, or in others represented only by a discolored spot (i.e., the
attachment-scar of the suppressed half) at the base of the nutlet. The
attachment scar of the two-seeded nutlet is symmetrical and evenly
rounded, but in the one-seeded nutlet it is irregular and somewhat lobed
on the side of the nutlet with the suppressed cell. There are other indica-
tions that the nutlets of Cerinthe are duplex. The tips of the united nutlets
remain free when both are equally developed and form the characteristic
bicuspid tip to the Cerinthe nutlet. The wall separating the two cells within
the nutlet is dense and homogeneous. It bears no cavities nor any abscission
layer, as frequent in comparable partitions in the nutlets of the Helio-
tropioideae and the Ehretioideae. The endocarp of the united nutlets is
completely confluent. There are, however, lines of suture on the exocarp
delimiting the boundaries of the component nutlets. A lineate groove is
always present down the middle of the flattened venter, from between the
two nutlet-tips down to the base. This groove may also continue more or
less clearly down the middle of the convex dorsum and occasionally even
be traced across the middle of the attachment scar. In the attachment
other evidence of the duplex nature of the nutlet is evident, for the two
halves of the nutlet share no vascular supply, both being directly affixed
to the gynobase, and each supplied directly from it.

Cerinthe has pollen very different from that of Onosma and Podonosma,
which is surprising, for the corollas and androecium of the three are very
similar. The symmetric grains of Cerinthe have eight pores borne in a
well-marked equatorial groove and associated with eight colpi. T he strong
equatorial groove distinguishes the pollen from that of all other Litho-
spermeae, Pollen of Onosma and its allies is uniformly three-porate. In
number of pores, accordingly, Cerinthe is in very much closer agreement
with the pollen of Lithospermum and its allies and, interestingly, with
that of the American genera which most resemble Onosma in corolla and
androecium. The three natural species-groups within Cerinthe have each
a distinctive pollen.

The largest grains (33-37 x 30-33 m) are associated with the well-
marked species, C. retorta. These are ellipsoid-sphaeric, almost as broad
as long, and possess a narrow equatorial groove and eight linear (not
fusiform) colpi which extend well towards the polar areas. The pores are
unusually large. In C. major L. and its obvious relatives, the pollen grains
are barrel-shaped. They are distinctly longer than broad and have convex
or nearly parallel sides and abruptly narrowed ends. They measure
24-26 K 16-18 p». The grains have a broad equatorial groove and mod-
erately elongate fusiform colpi. The pores are obscure, but apparently
always eight in number. C. minor L. and C, glabra Mill. and their varieties
all have similar ellipsoid grains, 18-22 K 15-18 p, the smallest in the
genus. In lateral profile the sides are convex or slightly angled. The
eight pores are associated with an equatorial groove and short fusiform
colpi.

70 JOURNAL OF THE ARNOLD ARBORETUM _ [vot. xxxv

14. Podonosma Boiss. Diag. ser. 1, 11: 113 (1849) and FI. Orient. 4:
178 (1875). Type species Onosma syriacum Labill.

Plant perennial, herbaceous, or fruticulose at the base, hispid-villose or
somewhat hispid with integmixed slender gland-tipped hairs. Stems numer-
ous, simple or laxly branched, loosely spreading or decumbent. Leaves
numerous, cauline, lanceolate, acute, with evident midrib, obscurely pin-
nate-veined or veinless. Cymes loosely scorpioid, borne solitary or gemi-
nate, terminal on the stems and leafy branches, in age elongating and
becoming unilaterally racemose. Bracts lanceolate, numerous, becoming
foliaceous and conspicuous in the fruiting inflorescence. Pedicels slender,
elongating in age and usually becoming decurved in fruit. Calyx lobed
almost to the base; lobes lanceolate, nearly as long as the corolla-tube,
equal or nearly so, in fruit accrescent and becoming lance-cuneate with the
tips connivent and the bases reduplicate. Corolla blue, glabrous, regular,
inside without faucal appendages or stipitate glands; tube cylindric,
abruptly narrowed at the base, without a differentiated throat; lobes nar-
rowly triangular, soon reflexed, twice as long as broad, a quarter to a third
the length of the tube, sides straight, apex acute; annulus well developed,
villulose, distinctly ten-lobed. Filaments not half the length of the anther,
affixed in the upper third of the corolla-tube, compressed, narrowing upward
from the base. Anthers elongate, becoming half exserted from the corolla
mouth, laterally coherent below the middle, affixed about one fourth of
their total length above the base, upper half consisting of a somewhat
firm stramineaceous appendage which is narrowed to an acute tip, base
emarginate; thecae spreading only at the very base, their lower end with
a thickened tip, not coherent; back of anther with a broad slightly promi-
nent stramineaceous connective, not evidently papillate nor muricate. Pol-
len globose-ovoid, 18-22 16-20 », broader and more rounded at the
lower end; pores three, associated with fusiform colpi, submedial, borne
about the broadest part of the grain. Style filiform, tardily exserted from
the anther-tube; stigmas united, minute, terminal, Nutlets triangular-
ovoid, tuberculate, abruptly bent 90° below the middle, the small substipi-
tate attachment basal on the erect lower third of the nutlet body; ventral
keel thick and prominent. Gynobase strongly four-lobed; the lobes pale,
cartilaginous, pulvinate, each bearing a small oblique concave attachment
surface.

A well-marked genus containing two species, P. syriacum (Labill.)
Boiss., a plant of cliffs, walls, and other rocky places from Syria to Palestine
and east to Iran, and P. galalense Boiss., said to be a rhizomatous plant of
eastern Egypt. No material of the latter has been seen.

In general habit, glanduliferous herbage, form of calyx, decurved fruit-
ing pedicels, and most important of all, in appearance and structure of
nutlets and gynobase, Podonosma is extremely similar to Alkanna § Eual-
kanna. The corolla and androecium of Podonosma, however, is extremely
different from that of Alkanna. In these floral structures it shows detailed
similarities with those of Onosma. The similarities with both Alkanna and
Onosma combined in Podonosma are so striking and involve so many sig-

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVI 71

nificant details that I am forced to the belief that Podonosma shares a close
relationship not only with A/kanna but also with Onosma, one possible only
through a shared common ancestry.

The corolla and stamens of Podonosma are of the type present in
Onosma, The most obvious difference in the corolla is in the length of
the lobes. In our present genus these are elongate (twice as long as broad)
and reflexed, whereas in Onosma they are practically always short (about
as long as broad) and erect or only loosely recurved. Indeed in only one
species of Onosma, O. longilobum Bunge, are the lobes as elongate as in
Podonosma. The anthers of Podonosma, as in many species of Onosma,
are united into a tube, but unlike the anthers in the latter genus, those of
Podonosma cohere only for a limited time and then only along the margin
of the fertile lower half, and not at the base of the theca nor along the
margin of the terminal appendage. The terminal appendage is conspicu-
ously developed as in Onosma, but differs in texture, being pale, firm,
straw-colored, and opaque. Unlike that in all species of Onosma except
O. longilobum it narrows to a pointed tip. The back of the anther has
a well-developed convex stramineous connective, which, unlike the well-
developed connectives in Onosma, is not evidently papillate nor muricate.

The pollen is globose-ovoid, 18-22 16-20 un, slightly but distinctly
longer than broad and with the lower end more broadly rounded than the
upper. The three pores, in furrows, are usually obscure. They are sub-
medial and borne about the broadest part of the grain. This pollen is simi-
lar to that in species of Onosma which have the anthers united into a tube.
It approaches the isodiametric pollen which prevails among the more
easterly ranging species of that genus. The poller of Alkanna also has
three pores and furrows, but is more elongate and is conic-ovoid rather
than globose-ovoid in form.

The nutlets of Podonosma are very different from those of Onosma, but
agree to a remarkable degree with the very unusual nutlets of Alkanna,
cf. Jour. Arnold Arb. 34: 279 (1953). The nutlets are small, 1—1.5 mm.
long, and below the middle are abruptly bent 90° adaxially. The small
substipitate attachment scar appears to be superbasally lateral, but mor-
phologically speaking is actually basal on the short section of the nutlet
below the bend which remains erect. Dissection of the nutlet reveals that
the seed has a transverse flexure across the cotyledons. In situ within the
nutlet the tip of the cotyledons is adjacent and vertical to the portion of
the pericarp bearing the attachment scar, the proper position in a basifixed
nutlet. As in Alkanna, but very unlike that in Onosma, the gynobase of
Podonosma bears four pale cartilaginous cushion-like lobes, each bearing
a small concave attachment surface on its summit. The attachment sur-
faces are sloping and the nutlets when affixed to them are accordingly
ascending with their tips connivent.

15. Cystistemon Balf. f., Proc. Roy. Soc. Edinb. 12: 82 (1883) and
Trans. Roy. Soc. Edinb. 31: 186, t. 56 (1888). Type species C.
socotranus Balf. f.

15a. Vaupelia Brand, Fedde’s Repert. 13: 82 (1914); Green, Kew Bull.
1935: 528 (1935). Type species V. barbata (Vaupel) Brand.

72 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxv

Material adequate for a survey of Cystistemon and Vaupelia is not
available. Cystistemon I know only from the original description and il-
lustration. Of the eight species of Vaupelia I have only two represented
by specimens suitable for analysis, V. Aispida (Baker & Wright) Brand,
from 40 mi, s.w. of Norok, Kenya, A. G. Curtis 649, and V. barbata
(Vaupel) Brand, from betw. Menongue and R. Chimpompo, Angola, M.
A, Pocock 610. For details concerning the other species of Vaupelia I have
been forced to rely on original descriptions and upon Brand’s synopsis of
the genus, Fedde’s Repert. 13: 82-83 (1914).

Cystistemon has a single species endemic to Socotra and Vaupelia has
eight described species, one in southern Arabia and the others in tropical
Africa, where they range from Somaliland to Angola. The two genera are
immediately and very closely related and probably should be united. Their
affinity with Onosma is clear. There seems to be every reason for believing
that they represent a southern derivative of that genus in which the corolla-
tube has become abbreviated as the corolla-lobes increased greatly in rela-
tive size and importance. In all species of Onosma, with the single excep-
tion of O. longilobum Bunge, the tubular portion of the corolla is many
times longer than the corolla-lobes. In the present genera the lobes, or
at least the well-developed limb, are one to six times as long as the tube,
and furthermore are not erect or slightly recurving but spreading. In
Onosma the throat of the corolla is glabrous or at most only scantily and
inconspicuously strigose. It is never villose as in some of the species in
the present genera.

The anthers are very conspicuous features in the flowers of Cystistemon
and Vaupelia. They are extremely well developed, much longer than the
corolla-tube, and borne on short stout variously modified filaments affixed
near the middle of the short corolla-tube, and are accordingly almost com-
pletely exserted. The terminal appendage is extremely elongate, and unlike
that of Onosma usually greatly surpasses the fertile portion of the anther
in length. Its tip is usually pointed.

As in some of the species of Onosma, the anthers of the present genera
are united, but unlike those in Onosma, they cohere not at the base and
along the sides of the theca but only along the edges of the sterile terminal
appendages. The back of the anthers has a prominent broad connective
and a papillate surface, as is common in Onosma. The base of the anther
is more or less emarginate. The lower end of each theca, unlike in Onosma,
is completely unappendaged or at most has only an_ inconspicuously
thickened tip.

The filaments are of special interest. They are relatively short and stout
and affixed to the anther only a short distance above its base. In V. hispida
they are flattened and ovate-lanceolate in outline and ventrally on their
thickened base bear a small triangular ciliate appendage. In V. barbata
they are narrow but thick and above their base are nearly encircled by
a shelf-like ridge ciliate on the margin. Other members of the two genera
appear also to have comparable appendages associated with the base of

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVI 73

the filaments. The filaments of Cystistemon are described as ‘‘obcordatis
expansis inflatis basi annulo villoso cinctis.” Brand in his diagnosis of
Vaupelia describes the filaments as “brevia basi andropodio globoso piloso
inserta.”’ The hairy appendages associated with the filament in V. hispida
and V. barbata are unparalleled in Onosma. If similar structures are asso-
ciated with the filaments in other members of Vaupelia, they provide a
character of generic importance. The precise morphological nature of these
appendages is uncertain. Those in V. hispida and V. barbata I found very
suggestive of the infrastaminal appendages in Lobostemon, cf. Jour. Arnold
Arb. 34: 297 (1953). It seems unlikely, however, that the appendages in
Vaupelia and Cystistemon can be homologized with displaced lobes of the
annulus as in Lobostemon. The annulus is apparently suppressed or absent
in V. barbata, but in V. hispida it is represented at the base of the corolla-
tube by ten very minute tufts of hairs. In some species of Onosma the
lower portion of the filaments may be decidedly hairy, and the base may
be decurrent and thickened. Possibly from an ancestral condition of this
sort the hairy appendaged base associated with the filaments of modern
Vaupelia could have been elaborated.

The pollen of V. Aispida is sphaeric, or in polar profile, sometimes ob-
scurely three-sided. The grains measure 24-26 » in diameter. The three
pores, in furrows, are equatorial. In V. barbata the grains are globose-
ovoid, as broad as long (20-23 ,) or at most only very slightly longer than
broad. In lateral profile the grain is semicircular below the middle, but
above the middle it is lightly narrowed and tends to be somewhat flattened
at the upper end. The furrows are deep and rather conspicuous. The three
pores are slightly inframedial. The pollen, accordingly, is in general agree-
ment with that of Onosma and particularly with that of its more eastern
species, and especially those with united anthers.

16. Onosma L. Sp. Pl. ed. 2, 196 (1762) and Gen. Pl. ed. 6, 76 (1764).

Type species O. echioides L.
Colsmannia Lehm. Mag. Ges. Naturf. Berlin 8: 92, t. 4 (1818) and Asperif.
2: 356 (1818). Type species C. flava Lehm. [= Onosma sericeum Willd. ].

Perennial or biennial, herbaceous or sometimes fruticulose, coarsely
hispid with the hairs either appressed or spreading or frequently strigose
or sometimes more or less tomentose, stellate hair-clusters sometimes abun-
dant. Stems erect or spreading, simple or with leafy branches, arising from
a taproot or caudex. Leaves with a strong midrib but usually veinless,
veins when evident pinnate and usually weak; basal leaves clustered,
usually oblanceolate; cauline leaves usually decreasing in size upwar
along the stem, usually numerous. Inflorescence simple and consisting of
single or paired scorpioid cymes terminal on the stem and leafy branches,
or a thyrse composed of few to many cymes arising along the stem below
the terminal cyme; cymes distinctly scorpioid or loose and the flowers not
distinctly biseriate, usually not greatly elongate in fruit. Bracts numerous
but conspicuous only in a few species, usually shorter than the adjacent

74 JOURNAL OF THE ARNOLD ARBORETUM __ [VoL. xxxv

calyx, sometimes very small and shorter even than the adjacent pedicel.
Flowers at anthesis borne on strict pedicels on the curved highest portion
of the cyme, held vertically, horizontally or even pendulous according to
position on the curve of the cyme. Flowers regular and radially symmetric
or in one species with the corolla-bud and the anther-tube strongly curved
abaxially. Calyx usually accrescent, lobed to the base or in a few species
with a short tube which becomes distended by the ripening fruit; lobes 5,
equal or practically so, all distinct or rarely two partially united, very
elongate, linear to lanceolate or oblanceolate or ligulate, separated by
narrow Closed sinus, erect and parallel or in fruit more or less connivent,
not imbricate. Pedicels short and stout to slender and elongate, usually
lengthening in age. Corolla yellow, blue, or rarely white or red, tubular or
obconic-tubular or rarely ellipsoidal, usually gradually enlarging upward
from the base or sometimes more strongly ampliate above the middle to
form a more or less campanulate throat, usually broadest a short distance
below the base of the lobes but sometimes almost cylindric or less com-
monly broadest at or near the middle, shorter to much longer than the
calyx, commonly about twice as long, outer surface glabrous or more com-
monly evidently puberulent or strigose or hispidulous. Corolla-lobes small,
as broad or broader than long (or cuneiform in one species), triangular,
acute or with the tip rounded or short-attenuate, erect or with the tip and
margins more or less recurved. Corolla-throat without faucal appendages,
devoid of glands except those sometimes present about the base of the
filaments, glabrous inside or rarely with some appressed hairs along the
veins below each corolla-lobe or on and about the bases of the filaments.
Annulus usually evident, a well-developed ring or collar, usually more or
less lobed and more or less evidently villulose. Filaments affixed below
or near or above the middle of the corolla, short to very elongate, shorter
to longer than the anther, narrowly ligulate or gradually narrowed upward
or sometimes abruptly narrowed above a broadened base, glabrous or
rarely hairy or somewhat glanduliferous particularly near the base; base
frequently decurrent, symmetric or practically so. Anthers elongate, ter-
minated by a well-developed appendage, usually coherent at the base and
frequently joined laterally (usually by interlocking of minute marginal
trichomes) to form a tube, usually affixed distinctly below the middle, in-
cluded in the throat or partially to completely exserted from the corolla;

thecae distinct and usually spreading for a short distance above their ap-
pendaged base, or rarely joined and parallel throughout and their basal
appendages broadly joined and making a subtruncate or emarginate base
to the anther; basal appendage usually short and thick, sometimes hardly
more than a distinctly thickened basal tip of the theca, those of adjacent
anthers usually coherent but those on the same anther usually distinct. Pol-
len usually sphaeric to ovoid and as long or longer than broad, 16-25 « 16—
23 pw, but in one species strongly oblate, transversely elliptic and measuring

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVI 75

13-14 X 16 »; polar profile circular or more or less three-sided; pores
three, associated with fusiform furrows, equatorial to suprabasal. Style
filiform, glabrous or very rarely hairy, usually becoming exserted from the
corolla and surpassing the anthers, commonly emerging from the corolla as
the bud unfolds but sometimes emerging precociously from corolla-buds
which have not yet attained maximum size and in species with united an-
thers sometimes very tardily emerging from the synandrium, usually persist-
ing until the fruit is ripened; stigmas two, distinct or partially to completely
united, usually juxtaposed and terminal on the style, rarely subterminal
and separated by the very short obtusish sterile tip of the style. Nutlets
erect, small to large, smooth or less commonly variously tumulose or ver-
rucose or rugose, lustrous to opaque, gray to olivaceous brown or blackish,
frequently mottled, never white, somewhat ovoid to broadly lanceolate,
frequently somewhat rostrate and in one species with terminal and lateral
cornute appendages; venter usually angled, rarely with a thickened ventral
keel; dorsum convex or sometimes flattened below the middle; ventral
suture fused and obscure; attachment scar basal, usually triangular, as
broad as long, horizontal or somewhat oblique, sometimes green. Gynobase
pyramidal or flat or even depressed at the center, the attachment faces
usually distinct, frequently concave.

A large and variable genus with species occurring from northwest Africa
and Middle Europe east to southern Siberia and to western China and
Burma. Stroh, Beiheft Bot. Centralbl. 59B: 430-454 (1939) in a recent
purely bibliographic listing of the species, enumerates 123 species referable
to the genus as here accepted. Subsequent to Stroh’s census O. sericeum and
its allies have been treated by Levin, Not. Syst. Leningrad 12: 228-41
(1950), and Popov has published a detailed discussion of the genus,
Proalemy botaniki 1: 70-108 (1950) and also, Not. Syst. Leningrad 14:
287-304 (1951) a key in Latin as well as in Russian covering the thirty-
four species accredited to the U.S.S.R. The twenty-nine Chinese and
Indian species: have been keyed and described by Johnston, Jour. Arnold
Arb. 32: 201-356 (1951).

The genus has its closest relatives in Maharanga, Vaupelia, Cystistemon,
and Podonosma. Other close relations within the Lithospermeae are wit
Alkanna, Echium, and Cerinthe. American genera, such as Onosmodium
and Lasiarrhenum, sometimes suggested as relatives of Onosma, appear
to be more closely related to Lithospermum and are mimics of Onosma
rather than close relatives of it.

The corolla is almost always coarsely tubular, from the base moderately
and gradually expanding for most of its length or becoming more abruptly
expanded near the middle and differentiated into a stout tube and a more
or less campanulate throat. Such corollas are broadest a short distance
below the base of the small, commonly deltoid lobes. A noteworthy de-
parture from such conventional forms occurs in a few species. In
pyramidale Hook. of the Himalayas, the corolla has an abbreviated proper
tube and a large inflated throat. It is ellipsoid in form and hence sugges-
tive of the corollas of Maharanga. In O. stenosiphon Boiss. of Iran, the

76 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxv

corolla is scarcely at all ampliate, being very elongate and slenderly cylin-
drical. The most aberrant corolla is that of O. longilobum Bunge, of north-
eastern Iran and adjacent U.S.S.R., the only member of the genus with
the lobes conspicuously longer than broad. It has a gradually ampliate
tube and throat, together ca. 4 mm. long, and is broadest a short distance
below the base of the lobes. The erect lobes, cuneate with an attenuate tip,
are 2 mm. wide at the base and about 6 mm. long, and hence longer than
the tubular portion of the corolla. They are loosely folded longitudinally
(conduplicate) and the throat below them is angulate. The tube has five
elongate invaginations, one below each filament attachment. Inside, the
corolla is orthodox except for the terminal appendage of the anthers, which
gradually taper to a slender point. Because the corolla of O. longiloba
is so aberrant, it is probably inevitable that sooner or later the species
will be given generic recognition. I have refrained from doing so because
the plant, in all details other than corolla-form and pollen, is remarkably
similar to O. stenosiphon Boiss. and O. limitaneum Johnston and appar-
ently closely related to them. To be sure, these two latter species are not
conventional members of Onosma, for both have small, unusually slender
and elongate corollas, and O. limitaneum has somewhat tailed anthers and
usually rounded, rather than deltoid corolla-lobes. They are, however,
readily accommodated in Onosma, and O. longilobum, because of its rela-
tions with them, even though its corolla is aberrant, can be accommodated
in the genus also.

The species of Onosma usually have corollas with perfect radial sym-
metry. Indeed, the only notable departure from this condition is that pre-
viously reported in Jour. Arnold Arb. 32: 223 (1951) in O. multiramosum
Hand.-Mazz. of China. In that species the corolla-buds and the anther-
tube are both conspicuously bent abaxially above the middle. Though
other species with slender corolla-buds may have an occasional bud show-
ing a slight subapical bend, in none of these is it so consistent and extreme
as in O. multiramosa. In the closely related Makaranga the filament-bases
within each corolla are clearly differentiated into three types, and in this
respect the androecium has a manifest bilateral symmetry. This is not
so in Onosma. In the vast majority of species in Onosma the stamens
within the corolla are indistinguishable. If differentiation of the filaments
does exist in Onosma, as perhaps very obscurely in O. Waddellii Duthie,
O. Wardii Johnston, and O. mertensioides Johnston, the difference is very
slight and difficult to detect.

Among the most distinctive features of Onosma are its elongate anthers.
These are almost always coherent, at least at the base, and are always ter-
minated by an evident sterile appendage. Descriptions of the organ by
past authors seem to have been largely if not exclusively based upon its
representation in the species of Europe and the Near East. Among the
Chinese, Indian, and some of the species of the Middle East, the organ
shows more variation than is generally recognized. In western species the
anthers, if joined, are coherent only at the base, but in many of the more
eastern species they unite laterally to form a tubular synandrium. Simi-

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVI 77

larly, western species have a terminal appendage that is strap-shaped and
papery hyaline, whereas among the more easterly ones the appendage on
the anther is frequently much firmer in texture, dark in color, more attenu-
ate, and frequently somewhat rostrate in nature. Among the western plants
the anther averages very much larger than in the east. The connective
on the back of the anther, conspicuously developed and usually broad and
prominent in the west, becomes very narrow and inconspicuous in many
of the eastern species. The back side of the anther usually has its surface
papillate or muriculate. Among eastern species, however, it may be nearly
smooth. In most all species the anther has its thecae distinct and more or
less evidently spreading for a short distance above the base, and the lower
end of each theca is thickened and more or less prolonged to form a short
basal appendage. Only in O. limitaneum are the appendages notably
elongate. In O. multiramosum and to a less extent among other Chinese
species, the thecae remain joined and parallel for their total length, and
their broadened and confluent basal appendages form a truncate or weakly
emarginate base to the anther.

Pollen of forty-five species of Onosma has been examined. The grains
have three pores which are frequently somewhat protrudent and are always
associated with fusiform colpi. In polar profile the grains may be circular
or more or less three-sided with the angles rounded or emarginate. The
pollen varies from species to species in lateral profile. All the many inter-
mediate forms between decidedly ovate and perfectly circular and even
transversely elliptic are represented. The lateral outline, as in O. echioides
L., O. arenarium W. & K., and O. dichroanthum Boiss., can be decidedly
ovate with the length evidently much greater than the breadth (25 X 16—
20 »). In such grains the pores are borne where the grain is broadest, about
a third of the length of the grain above its base. In other species the grains
may still retain a modified ovate outline but show less difference between
length and breadth. In O. confertum W. W. Sm., for example, the grains
are equal or practically so in length and breadth (16 ») but are noticeably
more broadly rounded below the inframedial pores than above them. From
grains such as these there are many transitions to grains with equatorial
pores and equal hemispheres, in which the lateral profile is perfectly cir-
cular or even very slightly broader than long. In O. pyramidale Hook. the
grains have a polar axis which is obviously shorter than the breadth at
the equator (13-14 < 16 »). In lateral profile these grains are distinctly
transverse-elliptic. The pores are equatorial or at most only very slightly
inframedial.

Most of the species of Europe and the Near and Middle East have
pollen of the ovoid type, i.e., longer than broad, 20-25 X 16-20 pn, and
bearing the pores well below the middle, usually ca. 8 » above the base of
the grain. Among the Sino-Indian species, only seven out of the twenty-
nine species have pollen of this type, viz., O. chitralicum Johnston, O.
hispidum Wall., O. khyberianum Johnston, O. barbigerum Johnston, O.
Griffithii Vatke, O. bracteatum Wall., and O. dichroanthum Boiss. Species
with grains isodiametric or nearly so (16-26 ») are most abundant in the

78 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxv

more eastern portions of the range of the genus. There are twenty-two of
the twenty-nine Sino-Indian species with pollen of this type. Among the
species of the middle East I have encountered the type only in species
such as O. stenosiphon Boiss. and O. rostellatum, which appear to be more
closely related to the Indian and Chinese congeners than to the conven-
tional Onosmas of Europe and the Near East. Interestingly, similar pollen
occurs in Vaupelia, a very close relative of Onosma, which has long-exserted
united anthers, as do many of the eastern species of Onosma.

17. Maharanga A.DC. Prodr. 10: 71 (1846). Type species Onosma
Emodi Wall.

Plants herbaceous, perennial or biennial or perhaps sometimes annual.
Stems one to several, erect to decumbent, hispid or hispid-villose, simple
or with strict to ascending leafy floriferous branches, arising from a cluster
of basal leaves or more commonly from among the remnants of a leaf
cluster produced the previous season. Leaves very obscurely to evidently
veined; veins strictly ascending, usually pinnate but frequently with the
lowest pair most elongate and most prominent; basal leaves usually ob-
lanceolate; cauline leaves numerous, the middle and upper ones usually
larger than those near the base of the stem, oblanceolate to lanceolate,
usually hispid or hispid-villose with the hairs frequently appressed, hairs
scanty to abundant and those on the upper leaf-face frequently with dis-
coid bases, stellate hair-clusters present in two species. Cymes usually
geminate, not clearly scorpioid and forming somewhat glomerate clusters
broader than long, terminal on the main stem and sometimes also on the
leafy branches, in fruit loosening to become somewhat corymbose or rarely
loosely racemose. Bracts small, narrow, not conspicuous, none except the
lowermost surpassing the adjacent calyx. Calyx with a broad deeply
saucer-shaped or shallowly cup-shaped tube and ascending lobes; lobes
cuneate or narrowly triangular, gradually narrowed from a broad base,
one and a half to three times as long as broad, separated by open angulate
sinus, tip acute or somewhat attenuate; lobes in fruit becoming loosely
incurved and the two abaxial ones sometimes slightly larger than the other
three; pedicels elongating, in fruit usually slender and as long or longer
than the calyx. Corolla blue, purplish, yellow or white, about twice the
length of the calyx, outside strigulose or villose-hispidulous, ovoid-ellipsoid
or obovoid, contracted at both ends, abruptly expanding from a short tube
into a relatively large inflated throat and at or above the middle contract-
ing to a small mouth, above the middle usually with persisting narrow
inflexed vertical plaits and hence appearing sulcate below each sinus, below
each lobe with an inflated rib that protrudes between the calyx-lobes and
has an abruptly contracted lower end that: rests chin-like in the broad
angulate sinus between the calyx-lobes, outside below the middle and
usually hidden by the calyx-lobes bearing five elliptical glabrous areoles
which are usually slightly depressed and sometimes invaginate near their
upper end; mouth of corolla very small; lobes very small, more or less
deltoid, loosely recurved above the middle, as long as broad or somewhat

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVI 79

longer than broad, apex acute or rounded; inside of corolla glabrous or
more or less hairy about the bases of the filaments, devoid of glands or
faucal appendages; annulus evident, collar-like, more or less _villulose.
Filaments affixed at base of the throat, abruptly narrowed from a broad
base, slightly longer than the anthers or distinctly shorter, two filaments
with a symmetric base, a medial vein and a transverse attachment on the
corolla, the other three with an oblique or vertical attachment and an
asymmetric base with an excentric vein. Anthers lanceolate, always in-
cluded, coherent at the base, affixed below the middle, terminated by a
usually well developed, firm, gradually attenuate sterile appendage which
is usually armed with some stout ascending hairs or cuneiform teeth on
the margin and by a cluster of cuneiform teeth at the apex, base of anther
with the thecae distinct and somewhat spreading; back of anther very
obscurely papillate, the connective narrow and neither prominent nor con-
spicuous. Pollen cylindric or in one species ellipsoidal, pores three, equa-
torial, associated with fusiform colpi; polar profile circular or three-sided
with notches at the corners. Style filiform, glabrous, included or short
exserted, rarely persisting with the fruit; stigmas two, juxtaposed and
terminal on the style, small, distinct or more or less united. Nutlets slightly
incurved, brown or olivaceous, dull, tumulose and coarsely rugose with the
surface very abundantly and densely muriculate or papillate, somewhat
ovoid, with the venter angulate and provided with a coarse prominent
ventral keel; ventral suture absent; attachment scar basal, about as broad
as long. Gynobase broadly pyramidal, the sloping attachment surfaces
separated by low cartilaginous ridges.

A group of nine species occurring in the mountains from the middle
Himalaya east into southwestern China. The genus is obviously related
to Onosma but is readily distinguishable by its calyx and pollen, as well
as by the form and structure of its corolla.

The corolla of Maharanga has a very short proper tube and a relatively
large, well-developed, much inflated throat, usually broadest near the
middle. The mouth of the corolla and the base of the tube usually have
about the same diameter. In Onosma the corolla form of Makaranga is
closely simulated only by the inflated ellipsoid corolla of O. pyramidale
Hook, This lacks, however, the characteristic outward features of the
corolla of Maharanga, such as the narrow, tightly inflexed plaits below
each sinus and, below each corolla-lobe, the distinctly puffed-out ribs which
have an abruptly contracted lower end that rests chin-like in the angle
of the open sinus between the calyx-lobes. Also lacking in Onosma pyra-
midale is the glabrous elliptical areole on the outside of the corolla
which is associated with the very different attachment of the filaments in
Maharanga.

The filaments are abruptly narrowed and become more or less linear
above their broad base. They are affixed directly on the corolla-walls or
arise from the shoulders of gibbose invaginations. The five stamens within
the corolla differ in the shape of the filament, in the course of the vein
within them, and in the orientation of their attachment on the corolla-

80 JOURNAL OF THE ARNOLD ARBORETUM _ [vot. xxxv

walls. Two filaments have an arcuate attachment oriented transversely,
and their vein is medial. Two other filaments are obliquely affixed, and
their asymmetric base has an excentric vein. The fifth stamen (apparently
the medial adaxial one) is vertically affixed and has a strongly excentric
vein and a more prolonged and asymmetric decurrent base. Although no
evidence of it has been detected elsewhere in the corolla, bilateral rather
than radial symmetry is manifest in the filaments of Makaranga. In
Onosma the stamens are all similar, or if there is any differentiation among
them within the corolla it is very obscure.

The anthers of Maharanga in size and general form are similar to those
in some species of Onosma (e.g., O. pyramidale Hook. and O. cingulatum
W. W. Sm.) but differ from those of most species of that genus in their
smaller size, narrow smoother connective, and firmer dark, beak-like ter-
minal appendage. Occasionally in Onosma as in the species just mentioned,
the terminal appendage of the anther may be obscurely papillate on the
margin and so appear very minutely denticulate. In all species of Mahar-
anga, except M. bicolor DC., the lateral margins of the appendage bear
some short, stout, ascending hairs or subulate teeth. The tip of the ap-
pendage in Onosma may be attenuate, truncate, emarginate, or bidentate.
In Maharanga the very tip appears to be somewhat thickened and com-
monly bears a number of cuneiform teeth, sometimes in a more or less
crown-like arrangement.

The pollen of Maharanga is 3-porate and 3-colpate as in Onosma but is
very different from that of Onosma in form. The grains (20-23 % 10-16
».) are elongate and in most species are cylindric. In lateral profile they
have abruptly rounded ends and sides that are nearly straight and parallel
except for a weak constriction at the equator. The pores are equatorial.
In polar profile the grains are circular or somewhat three-sided with the
broad corners somewhat emarginate. The only departure from the cylin-
dric form described occurs in M. bicolor. In that species, aberrant in a
number of other minor details (included style, relatively large united
stigmas, and very elongate laterally unarmed anther appendages), the
pollen grains are distinctly ellipsoid. In lateral profile the sides are rounded
or slightly angled at the equator where they are broadest and bear the pores,

The nutlets of Mahkaranga are similar in size and form to those of many
of the Chinese species of Onosma, and like many of the latter they have
a rough, tumulose, and rugose epicarp. The distinctive feature of the
Maharanga nutlet is the surface of the epidermis, which is very abundantly
and very densely beset with crowded minute papillae or muriculations.
Only in a very few Chinese and Indian species of Onosma are the nutlets
provided with comparable minute epidermal roughenings, and even in these
they are never so well developed and so very abundant as on the nutlets
of Maharanga.

The units of the inflorescence are not of the extreme scorpioid type
prevalent among the species of Onosma. In Maharanga the rhachis of the
cyme remains relatively slender and is not greatly elongate. The flowers,
neither extremely numerous nor evidently biseriate, become slenderly ped-

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVI 81
icellate. Cymes resembling those of Maharanga, however, are developed
in some Chinese species of Onosma such as O. album W. W. Sm., O. con-
fertum W. W. Sm., O. cingulatum W. W. Sm., O. exsertum Hemsl., O.
fistulosum Johnston, and O. paniculatum Bur. & Franch., as well as in
such congeners from the Middle East as O. stenosiphon Boiss., O. longi-
lobum Bunge, and O. polyphylium Ledeb.

The species of Maharanga have been described and distinguished, as
members of the genus Onosma, in my treatment of the “Sino-Indian Species
of Onosma,” Jour. Arnold Arb. 32: 206, 356-67 (1951). With the validity
of the genus Makaranga now recognized, the following new binomials are
required:

Maharanga verruculosa (Johnston), comb. nov.
Onosma verruculosum Johnston, Jour. Arnold Arb. 32: 356 (1951).

Maharanga Borii (Fischer), comb. nov.

Onosma Borii Fischer, Kew Bull. 1940: 38 (1940).
Maharanga lycopsioides (Fischer), comb. nov.

Onosma lycopsioides Fischer, Kew Bull. 1940: 39 (1940).
Maharanga microstoma (Johnston), comb. nov.

Onosma microstoma Johnston, Jour. Arnold Arb, 32: 360 (1951).
Maharanga dumetorum (Johnston), comb. nov.

Onosma dumetorum Johnston, Jour. Arnold Arb. 32: 361 (1951).
Maharanga egregia (Johnston), comb. nov.

Onosma egregium Johnston, Jour. Arnold Arb. 32: 366 (1951).

ARNOLD ARBORETUM,
HARVARD UNIVERSITY.

82 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxv

SOME DETAILS OF THE STRUCTURE OF
RHODOTHAMNUS CHAMAECISTUS

HERBERT F. CoPELAND
With one plate

THIS NOTE is a supplement to my paper (1943) on the genera of the
subfamily Rhododendroideae of the family Ericaceae. In that paper, the
genus Kalmiopsis Rehder (1932) was suppressed: its only species, K.
Leachiana (Henderson) Rehder, an undershrub local on certain summits
in southwestern Oregon, was transferred to Rhodothamnus. This was done
without original study of the only previously recognized species of Rhodo-
thamnus. The latter, R. Chamaecistus (L.) Reichenbach, is local on the
Alps; under the conditions of the times, it had been impossible to obtain
material of it. As soon as communication between Germany and America
again became feasible, Dr. Hermann Sleumer had the kindness to send
material of R. Chamaecistus fixed for histological study. The material is
from cultivated plants ex Museo botanico Berolinensi, and was collected
during and just after flowering, in April and May, 1948. It has been sub-
jected to the remaining steps of routine microtechnique and duly studied.
The results permit further discussion of relationships between Kalmiopsis
and Rhodothamnus.

Rhodothamnus Chamaecistus is an undershrub with crowded small thick
alternate leaves. In spring, it produces a few flowers on long pedicels
arising from between pairs of bracteoles in the axils of leaves near the tip
of the stem. The growth of the vegetative shoots of the year, terminal
and axillary, begins at about the same time.

The flowers (fig. 6) have five narrowly triangular sepals; a pink corolla
with a brief tube and a rotate limb, 5-lobed, with deep sinuses; ten long-
exserted stamens; a depressed-globular ovary with five locules opposite
the lobes of the corolla; and a long style springing from a depression in
the summit of the ovary, the stigma not expanded. The flower is slightly
zygomorphic; the median petal is on the lower or abaxial side. The sepalad
stamens are of greater average length than the petalad stamens, the longest
stamens of both whorls being on the adaxial side. The anthers (fig. c)
are elongate. The two terminal “pores” of each anther are actually brief
slits which gape widely at maturity.

The leaves, but not the bracteoles, are ciliate with stout gland-tipped
bristles. Stems, pedicels, and calyces are beset with similar bristles. There
are simple hairs on stems, pedicels, ovaries, and the bases of the filaments.

The stems are of the same internal structure as those of other Rhodo-
dendroideae, being of the type which lack a cylinder of fibers of pericycle.

The leaves reach a width of about 3 mm. and are about 300 ,» thick

1954] RHODOTHAMNUS CHAMAECISTUS 83

(fig. a). The epidermis is of large cells with a thick cuticle; the cells are
larger, and the cuticle is thicker, on the upper side. The stomata are confined
to the lower epidermis and open at the level of the outer surface. The
guard cells bear a ridge at the outer opening of the stomatal passage, but
not at the inner opening. There are two or three layers of palisade cells
occupying about half of the thickness of the leaf. The veins are not joined
to the epidermis by flanges of differentiated tissue; they are imbedded
between palisade and spongy tissue. Each vein contains a conspicuous
strand of fibers between the xylem and phloem.

The base of the floral receptacle is impressed. The ten main perianth
bundles which arise from the stele in this region (fig. d@) are recurved at
their origin. Theoretically, these bundles are two whorls, but all arise at
approximately the same level. Five of them are the median bundles of
sepals. The five which alternate with these undergo forking, each into
three; the middle branch forks again and supplies a lobe of the corolla;
the lateral branches are the lateral bundles of the adjacent sepals. There
are occasional deviations from the typical pattern thus described. The ten
stamen bundles arise from the upper sides of the main perianth bundles.
The vascular tissue which continues beyond the departure of the main
perianth bundles forms for the most part five bundles, fused pairs of ventral
bundles of adjacent carpels, which ascend the central column of the ovary
in the planes of the septa to about half the height of the ovary. Each of
them divides into two branches which run out into placentae in adjacent
locules. From the bases of the perianth bundles and the carpel ventrals
small bundles originate, in no evident pattern, and run out into the ovary
wall. Some distance above the base of the ovary, five of these small
bundles become recognizable, by their course in the median planes of the
-carpels, as carpel dorsals. The other small bundles fade out; the carpel
dorsals ascend the ovary wall to its summit, dip under the depression
about the base of the ovary, and ascend the style.

The internal structure of the ovary, the moderately elongate ovule
(fig. e), and the embryo sac are quite as in other Rhododendroideae.

Nearly all of the observed characters of Rhodothamnus Chamaecistus
are those which are to be expected of a plant included in subfamily Rho-
dodendroideae and tribe Phyllodoceae. Zygomorphy is somewhat more
evident in the flowers of this plant than in those of other members of the
tribe, and the rotate corolla with deep sinuses is exceptional. The anthers
and the vascular system in the receptacle are quite as in Kalmiopsis Leach-
iana, Kalmia, and Phyllodoce. The anatomy of the leaves is essentially ex-
actly as in Kalmiopsis Leachiana: Breitfeld (1888) was mistaken (so far
as the available material shows) in describing the epidermal cells as small
and the veins as “durchgehend.” Kalmiopsis Leachiana remains different
from Rhodothamnus Chamaecistus in its larger leaves, its more extensive
inflorescence, its shallowly indented campanulate corolla, and particularly
in its peculiar scales, which simulate those of Rhododendron and Ledum
while not being of the same structure.

Judgment as to the expedient limits of taxonomic groups is often in-

84 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxv

escapably subjective. To such judgment it appears, more definitely than
before, that Kalmiopsis is not to be maintained as a genus. The plant of
southwestern Oregon is to be called Rhodothamnus Leachianus. As repre-
sented by the two species here ascribed to it, Rkodothamnus is indeed a
remarkable example of interrupted distribution.

LITERATURE CITED

1. Breirretp, A. Der anatomische Bau der Blatter der Rhododendroideae in
Be eziehung zu ihrer systematischen Gruppierung und zur geographischen
Verbreitung. Engler’s Bot. Jahrb. 9: 319-379. 1888.

2. CopeLanp, H. F. A study, anatomical and taxonomic, of the genera of Rho-
dodendroideae. American Midl. Nat. 30: 533-625. 1943.

3. Renper, A. Kalmiopsis, a new genus of Ericaceae from northwest America.
Jour. Arnold Arb. 13: 30-34. 1932.

SACRAMENTO JUNIOR COLLEGE
SACRAMENTO, CALIFORNIA

Jour. ARNOLD Ars., VoL. XXXV PLATE I

[eyo

OBR

O75)

RHODOTHAMNUS CHAMAECISTUS (L.) Reichenbach

RHODOTHAMNUS CHAMAECISTUS. a, cross section of leaf * 320. b, flower
X 1.6. c, anther X 8. d, vascular system in the floral receptacle X 40. e, ovule
X 320.

86 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxv

TWO NOMENCLATURAL CHANGES IN THE
CHINESE FLORA

ALBERT N. STEWARD

DuRING THE STupyY of herbarium specimens and literature related to the
flora of the Lower Yangtze Valley it has come to the attention of the writer
that there is confusion in the nomenclature of the species described by
Hemsley under the name Diospyros sinensis in Jour. Linn. Soc. Bot. 26:
71 (1889).

It appears that this name was earlier ascribed by Naudin to Blume in
Nouv. Arch. Mus. Paris II. 3: 221 (1880) to cover what is probably a
form of D. Kaki, since Naudin cites D. Roxburghii (Carr. Rev. Hort.
1872, p. 253, fig. 28-29) as a synonym of D. sinensis, and the figures for
D. Roxburghii appear very much like a form of D. Kaki. Perhaps Naudin
had in mind Blume’s D. chinensis (Cat. Gewass. in ’s lands Plantentuin te
Buitenzorg, p. 110. 1823), which is a synonym for D. Kaki Linn. The
name now proposed for this species is:

Diospyros cathayensis nom. nov.

Diospyros sinensis Hemsl. in Jour. Linn, Soc. Bot. 26: 71 (1889), non Blume
ex Naudin in Nouv. Arch. Mus. Paris II. 3: 221 (1880).

This wild persimmon was well described by Hemsley in Index Florae
Sinensis (cited above) and was later figured in Hooker’s Icones Plantarum
29: t. 2804 (1906). It was reported by Rehder and Wilson as “‘a common
small evergreen or subevergreen tree in the dry valleys of western Szechuan”
in Pl. Wils. 2: 591 (1916). The type specimen was collected by Faber at
an elevation of 4000 ft. on Mt. Omei, Szechuan. In addition to other locali-
ties in Szechuan, the species is now known from stations in Hupeh and
Yunnan in Western China, and from Chekiang, Kiangsi, Fukien and
Kwangtung in Central and Southern China.

Desiring to follow a conservative procedure as to nomenclature in con-
nection with studies on the flora of the Lower Yangtze Valley, it has seemed
best to use Satureja Linn. in the broad sense and not to treat Calamintha
Lam. as a separate genus. Consequently, the following combination be-
comes necessary:

Satureja polycephala (Vaniot) comb. nov.
Calamintha polycephala Vaniot in Bull. Acad. Géogr. Bot. 14: 183 (1904).

DEFARTMENT OF BOTANY,
OREGON STATE COLLEGE.

oe

i
we
?

RICHARD A. HOWARD
ARNOLD PROFESSOR OF BOTANY
DIRECTOR OF THE ARNOLD ARBORETUM

JOURNAL

OF THE

ARNOLD ARBORETUM

VoL. XXXV APRIL 1954 NUMBER 2

STUDIES IN THE KUHNIINAE (EUPATORIEAE) II *
L. O. GAISER

With five plates

INTRODUCTION

Liatris and Brickellia, the two largest genera of a small subtribe of the
Eupatorieae, which Robinson (1913) called the Kuhntinae, were ex-
amined cytologically and found to have the basic chromosome numbers
of 10 and 9 respectively. Brickellia with approximately ninety species
(Robinson 1917) has its greatest distribution in Mexico and southwestern
United States. It extends northward to the Canadian border and south-
ward into Central America and sparingly into Brazil. The range of Liaérts,
with thirty-two species (Gaiser 1946), is confined almost entirely to
southern Canada and the United States, though it just crosses the border
into Mexico. The remaining seven genera are small. However, their dis-
tribution in the American hemisphere is interesting in relation to the two
large genera mentioned. Barroetea is wholly Mexican and Kanimia is
found only in South America. Two others, Trilisa and Carphephorus occur
only in southeastern United States while the monotypic genus Garberia
is limited to Florida. Only the two genera, Kuhnia and Carphochaete
overlap from United States into Mexico.

The comparative growth-form of these genera is also a matter of interest.
Liatris is a genus of perennial herbs with mostly corm-like rootstocks,
though in the series Punctatae, deeply penetrating roots occur, and the
singular L. Garberi of the Spicatae, has a tuberous form. By contrast,
Brickellia consists largely of shrubs or shrubby perennials, of which prob-
ably the most woody is B. argyrolepis, which attains a height of twelve
feet and a stem diameter of two inches (Gaiser 1953). It is difficult to
sharply differentiate between herbs and shrubs in this genus, for the stem
is often only slightly woody at the base or consists of a woody caudex.
Others are perennial herbs with only persistent underground parts which
are fusiform, tuberous or more knobby and rhizomatous. Only one species,
or possibly two, are annuals. Of the seven species of Barroetea (Robinson

* Chromosome Studies in the Kuhniinae (Eupatorieae). I. Brickellia. Rhodora
55: 253-267, 269-288, 297-321, 328-345 (1953).

88 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxv

1911b), one is known to be a perennial herb, somewhat woody at the base,
and three are recognized as annuals. Kuhnia species are comparatively
slender herbs with long somewhat conical roots which have been described
by Shinners (1946) as irregularly divided at the top and becoming more
or less ‘‘soft-woody.” Robinson (1911la) described K. adenolepis as hav-
ing a woody caudex. Carphephorus and Trilisa are true perennial herbs
having clustered, thick, almost tuberous roots and leafy rosettes. The
single species of Garberia is definitely a woody shrub attaining a height
of six to eight feet, while the species of Carphochaete are generally
spoken of as small branching shrubs

In the cytological studies of. Liatris, including all the given species
except L. lancifolia, only the diploid number (2n = 20) was found in
those of nine series, while polyploidy prevailed in the Punctatae (Gaiser
1949, 1950, 1951). In one species only the hexaploid number was found,
in a second, only the tetraploid, in a third, the diploid, while both diploid
and tetraploid numbers occurred in two others. Further, one of these two
latter, L. punctata Hook. has the widest distribution of any species in
the genus, extending the full north-south range from western Canada,
east of the Rocky Mountains, over the Mexican border. In forty-one
species of Brickellia examined, representing half of the undoubtedly dis-
tinct species, and four sections, only the diploid number (2n = 18) was
found

On the basis of previous cytological work, it seemed worth while to de-
termine ? the chromosome numbers in the other genera of this small group
of the Compositae with the hope that new clues of relationship would
become evident. From fossil evidence, the family is believed to be of
recent development. With the two largest genera known to differ by one
in their chromosome number, questions arose: Was there a smaller basic
number? If so would it be found in genera limited to the tropics or to
the more woody forms? Would those that were shrubby have the same
number as Brickellia and the perennial herbs the same as Liatris? Or
would there be still further variations in number and if so what relation-
ship would this bear to the classification?

There was also the interesting problem of polyploidy, since though
lacking in Brickellia, it was found in Liatris, a genus limited to the tem-
perate zone. There have been contributions which fail to confirm but
also some which give support to Hagerup’s original hypothesis (1932) that
polyploidy developed under rigorous environmental conditions. In Steb-
bins’ (1950) discussion on the topic of the polyploid complex and geo-
graphic distribution he stated that no tropical group was known well
enough to be included. When undertaking the study of Brickellia, it was
expected that because of its range into Mexico and Central America it

‘The author gratefully acknowledges a grant from the Canadian Research Council
when a beginning was made. Subsequently the project was assisted by a gra ae —
the American Philosophical Society which permitted the collection of spec of
Barroetea and Kuhnia as well as Brickellia in Mexico, without which this fin teat
could not have been completed.

1954] GAISER, STUDIES IN THE KUHNIINAE II 89

would include tropical forms. Field experience in the collection of ap-
proximately two dozen species from Mexico and Guatemala impressed
upon the author that they were on the plateaus and thus were actually
existing in a temperate climate. The annual B. diffusa is somewhat ex-
ceptional for although it occurs on the plateaus too, it is the only species
to be found throughout the Caribbean. All of the species of Barroetea,
according to Robinson (l.c.) occur on the plateaus in Mexico, and no
species has penetrated northward beyond its central zone. There is no
monograph of Kanimia but the labels on specimens at the Gray Herbarium
indicate that they have come from regions in the Andes from the Colom-
bian border to Peru and the province of Minas Gerais, Brazil. Thus there
are not included in this subtribe, plants of tropical lowlands such as make
up the genus Anthurium of the Araceae. But we know that polyploidy
does appear in five of a total of thirty-nine species of that genus examined
(Gaiser 1927). Stebbins (1950) apparently overlooked this when he made
the statement cited above. In contrast to one large tropical genus, as
Anthurium, the Kuhniinae present a challenging succession of related
forms which appear to have progressed northward from the tropics in
varying degrees.

NOTES ON THE TAXONOMY OF THE GENERA

In his key to the Eupatorieae, Robinson (1913) added an additional
subtribe to those given by Hoffmann (1890), and changed the name of
the Adenostylinae to Kuhniinae because the assumed type genus Adeno-
styles did not really belong to the tribe. With its omission from this sub-
tribe, the other nine genera remain the same.

Of them the genus Kanimia stands alone in having a definite number
of phyllaries. Both in the number of florets and phyllaries it is like
Mikania, from which Hoffmann (l.c.) had stated it was only separated
by its 8-10 ribbed achene. However, Mikania is placed at the end of the
previous subtribe Ageratinae. There is no general treatment of Kanimia.
Thirteen species have been described from Colombia, Peru, Ecuador, and
Brazil. Because it is South American, consists largely of vines or woody
forms, as are included in the much larger genus Mikania and has a definite
number of florets and phyllaries, a strong link is suggested between the
two subtribes. Whether the chromosome number would be of any help in
adjustment of generic segregation, will have to await the procuring of
cytological materials of Kanimia.

It is a matter of great satisfaction that of the remainder, the four largest
genera had been monographed, and two of them by Robinson, whose spe-
cialty was the Eupatorieae. Thus we have the excellent treatment of
Brickellia in 1917, following that of Barroetea in 1911, and the key to
the whole group in 1913. The genus Kuhnia was dealt with by Shinners in
1946, as had been one series of the genus Liatris, the Scariosae, in 1943.
The latter genus was in the process of being revised by Gaiser at that
time and publication followed in 1946. Of the four other genera still

90 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

lacking comprehensive treatments, a brief summation of their status is
given here.

Carphochaete

Next to the singular Kanimia, Carphochaete stands apart from the rest
of these genera in the nature of its pappus, which is scale-like and dilated
at the base, instead of being setose.

There has been no recent treatment of this genus, which was established
by Gray (PI. Fendl. 65, 1849). It was based on a somewhat shrubby
herb from northern Mexico, and named C. Wislizeni for its collector. In
1852, Gray (Pl. Wright. 89) emended the description of the genus as
shrubby or herbaceous but shrubby at the base, when describing two new
species, one Mexican, C. Grahami, and the other from the boundary region
of Mexico and New Mexico, C. Bigelovii. Since then only one other
species has been added by Greenman (Proc. Am. Acad. Arts & Sci. 44: 34,
1904). It also is Mexican and was considered as an herbaceous perennial
having a ligneous base. As the author’s description has distinguished it
from the other Mexican species, chiefly by the leaves and phyllaries, it
is clear that all four species are similar in their shrubby or near shrubby
nature. It was possible to include the American species in this study.

Garberia

Another shrub of this subtribe is found in the genus Garberia. When
cataloguing a collection of plants that had come from east Florida, Nuttall
(Amer. Jour. Sci. 5: 299, 1822) gave a brief Latin description of Liatris
fruticosa. It was the only species of that genus referred to as fruticose by
DeCandolle (Prod. 5: 132, 1836) and became the subdivision Suffruticose
in Torrey and Gray (Fl. N. Am. 2: 76, 1841). This plant, referred to by
Nuttall in 1841 (Trans. Amer. Phil. Soc., ser. 3, 7: 285) as agreeing with
the genus Liatris in its flowers but not in its habit, was transferred by
Gray (Proc. Acad. Nat. Sci. 379, 1879) to the new genus Garberia. Thus
it appears in more recent floras of the South as Garberia fruticosa (Nutt.)
Gray. The restudy of William Bartram’s plants (Bartr. Trav. 164, 1791;
ed. 2 162, 1792) by Merrill (Bartonia 23: 24, 1944) called for a new
combination. After Dr. Francis Harper had covered Bartram’s routes, he
was Satified that what Bartram had described as Cacalia heterophylla was
the same shrub. The plant is now known as Garberia heterophylla (Bartr.)
Merrill & F. Harper. It still is the only representative of the genus.

Carphephorus

The genus Carphephorus was established by Cassini (Bull. Soc. Philom.
Paris p. 198, Dec., 1816) when naming as C. pseudo-liatris, a specimen
without locality, seen in the Jussieu herbarium, which for a time was
thought to be of Siberian origin. In his Dictionnaire, Cassini (Dict. Nat.
Sci. 7: 148, 1817) explained that it belonged to the natural Tribe Eu-
patorieae, “section des Liatridees,” in which he was placing it, near Liatris,
the chaffy receptacle being the chief point of difference. Cassini’s “‘sec-

1954] GAISER, STUDIES IN THE KUHNIINAE II 91

tional’? name must be rejected because under the international code a
section is of infra generic rank, while Cassini applied this term to a group
of genera. Torrey and Gray (1841) found that the character of the re-
ceptacle had been overlooked by some in describing other species and
rightly recognized Carphephorus as an American genus. At that time,
three other species were transferred from the genus Liatris, L. bellidifolia
and L. tomentosa (Michx. FI. Bor. 20: 93, 1903) and L. corymbosa (Nutt.

Gen. 2: 132, 1818). At the same time also the older synonyms were given

for these four species, as they were given by Gray (Syn. Fl. 1(2): 1886);

hence they have not been repeated here. They still remain the four ac-

cepted species of the genus and all have been represented in this study.
Two Californian plants described as Carphephorus junceus Benth. (Bot.

Sulph. 21, 1844) and C. atriplicifolia Gray (Proc. Am. Acad. Arts & Sci.

5: 1591, 1861) were early placed in the Helanthoideae instead of the Eu-

patorieae, and assigned to a new genus Bebbia by Greene (Bull. Cal. Acad.

Sci. 1: 179, 1885). Thus though appearing under the genus Carphephorus

on page 113 in Gray (Syn. Fl. 1(2) 1886), they are given under Bebbia

in the supplement of that volume (p. 453). More recently, B. atriplicifolia

(Gray) Greene has been reduced to varietal rank by I. M. Johnston (Proc.

Cal. Acad. Sci., ser. 4, 12: 1197, 1924) so that this genus includes one

species, B. juncea (Benth.) Greene, its variety atriplicifolia (Gray) John-

ston, and variety aspera Greene (l.c.), all removed from Carphephorus.
Other excluded names and species are:

md ete baicalensis Adams in Mem. Soc. Nat. Moscou 5: 115
(1817); DC. Prod. 5: 132 (1836) = Saussurea ai Ledeb.
ae to Benth. & Hook., Gen. Pl. 2: 249 (18

Carphephorus cordifolius DC. Prod. 7: 267 (1838) = ene cordifolia
Robinson, Proc. Am. Acad. Arts & Sci. 47: 200 (1911).

Pee revolutifolin DC. Prod. 5: 132 (1836) according to Benth.

ook. ae 2: 249 (1873) “certainly from description, expelled
= the genus.”

Carphephorus triangularis (DC.) Gray Pl. Wright. 1: 86 (1852), ex

emsl. Biol. Cent. Am. Bot. 2: 109 (1881); Bulbostylis triangularis
DC. Prod. 7: 268 (1838) = Eupatorium vitifolium (Sch. Bip.)
Robinson, Proc. Am. Acad. Arts & Sci. 51: 537-8 (1917), Hebeclin-
ium vitifolium Sch. Bip. ex Klatt Leopoldina 20: 90 (1884).
Trilisa
For the genus Trilisa, which Cassini (Bull. Soc. Philom. Paris, p. 140,

Sep., 1818) established two years after the genus Carphephorus, the type

was Liatris odoratissima Willd. (Sp. Pl. 3: 1637, 1803). Willdenow had

taken up this name from Walter’s Anonymos odoratissimus Walt. (Flor.

Carol. 198, 1788), as he had also A. paniculatus, these names being de-

clared illegitimate by article 33 of the International Code of Botanical

Nomenclature. In his key of the Eupatorieae, Cassini (Dict. Sci. Nat. 26:

228, 234, 1823) placed this new genus in his section ‘‘Liatridees”’ between

Carphephorus and Suprago, a name which has been retained for the plu-

92 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXXV

mose section of Liatris. However, Cassini’s genus was not accepted at
once. DeCandolle (Prod. 5: 131, 1836) used it as a sectional name of the
genus Liatris, and this was followed by Torrey and Gray (Fl. N. Am. 2:
76, 1841). The latter, however, correctly eliminated unrelated species
and limited it to these two, collecting their older flora- and plate-references
not included here. Since Bentham and Hooker (Gen. Pl. 2: 248, 1873)
referred to the incorrect use of Cassini’s generic name for a section, there
has been no further confusion of this small genus.

One further species has been added. Small (Bull. Torr. Bot. Club 51:
392, 1924) had described a new plant from Florida under a generic name,
an anagram of Trilisa which it resembled somewhat in habit, as Litrisa
carnosa. When Robinson (Contrib. Gray Herb. new ser., 6: 104, p. 49,
1934) closely compared the plant with Trilisa and Carphephorus, he was
led to refer it to the former genus.

MATERIALS AND METHODS

The same methods have been employed as were used in the study of
Brickellia (Gaiser, l.c.). As some of these materials were collected along
with those of Brickellia, frequently preparations of both were carried along
together so that there might be as little variation in treatment as possible.
Whenever seeds were received on herbarium specimens or in packets from
other collectors,* they were germinated and eventually treated by the
same variety of techniques and stains. In the preparation of figures of the
chromosomes, the same photographic and microscopic equipment has been
used at the same magnification, with the intention of making adequate
comparisons of the size and morphology of the chromosomes with those
of Brickellia. Thus not only the same terms are used, but with the same
connotation of size. The same precautions were taken to study a sufficient
number of cells so as to be able to choose similar stages for comparison
of the chromosomes of all species of the different genera. Usually com-
parable stages of each have been photographed, but where the illustrations
represent slight variations of condensation, the purpose of the selection has
been discussed. In this study, use has been made of the photographic lens
to illustrate the trichomes and epidermal figures, both to elucidate and
confirm the drawings used for Brickellia.

In Table I are given the list of accessions of each species received along
with the place and date of collection and the collector’s name and number.*
They have been arranged and discussed in the order of a key prepared for
the genera (see page 124). As it has been impossible so far to obtain seeds
or any cytological material of even a single species of Kanimia from South
America, that genus is not included in this key.

Following the pattern of the investigation made of Brickellia, leaves of
the species of these genera were similarly cleared for the study of trichomes.

uthor again a deep gratitude to all contributors.

ma sh to the numbers of collectors will permit examination of a number of
specimens in various oes Sata ie or received by the author will be
deposited in the Gray Herbari

1954] GAISER, STUDIES IN THE KUHNIINAE II 93

As stated there, presence or absence of trichomes was not considered im-
portant for these studies. Attention was given to their form, as that has
been found to change little, if at all, under varying environmental condi-
tions. The terms applied in the descriptions and figures 61 to 88 of Brick-
ellia, have again been used here. The same principle was adopted of obtain-
ing leaves at the Gray Herbarium of specimens which had been referred
to by authors so that they would approach the typical. As these genera
are comparatively small, it was possible to examine one to several leaves of
almost all the species and their varieties. It must be pointed out that it
does not represent an examination of leaves of numbers of specimens of
any species. The difference in upper- and lower-most cauline leaves were
illustrated for one species, Trilisa paniculata. Changes in the ontogeny of
the species were not generally studied in detail.

OBSERVATIONS
I. Seedling growth

Whenever possible, an effort was made to grow some of the germinating
seeds for a time at least, in the greenhouse. Attempts to bring the plants
into bloom met with varying success. Of three species of Carphephorus,
two of Trilisa and the one Garberia none ever produced flowering axes
under the same conditions which stimulated flowering of more than a dozen
species of Brickellia, three species of Kuhnia, two of Barroetea and one
of Carphochaete. At least several plants of Trilisa paniculata were kept
for more than three years, a period longer than normally would be re-
quired for a biennial plant to come into flower, and above ground they still
had only rosettes of leaves. Time and facilities did not permit further
studies along this line.

The plantings attracted attention because of the striking differences of
the seedling conditions. As has been described and illustrated for eighteen
species of Liatris (Gaiser 1950a), during the first season, while the primary
root is becoming very much thickened, those seedlings produce only radical
leaves. As can be seen in figures 42 to 62, the larger ones with a longer
growing period, have formed quite a rosette. Then in the second season,
usually a flowering spike is produced. This was also illustrated in the
progeny of a natural hybrid (Plate II c, Gaiser 1951). Young plants of
Liatris punctata were here compared with those of the other genera.

In figures B and C are shown seedlings of Carphephorus pseudo-liatris
II and Trilisa paniculata VIII, of approximately sixteen and eighteen
months respectively, in comparison with one of Liatris punctata (Fig. A)
of approximately two years. The latter is old enough to show the begin-
ning of a forking and the subsequent elongation into the deeply penetrating
system which is characteristic of this species. In the Trilisa, the modifica-
tion to the thickened somewhat tuberous roots is already evident and to
a lesser extent in the younger Carphephorus also. At maturity, a similar
thickened fibrous condition would develop, for that is common to all the
species of the genus. This photograph happens to illustrate the aptness

Name

c riled aad
Bigelov

Garberia
heterophylla

Carphephorus
pseudo-liatris

—

—
—

C. bellidifolins

TABLE I
Chromosome Numbers in the KuAniinae
Accession Chromosome
Locality of Collector, Number
County Collection No. & Date n 2n
Pima Co., Ariz. Santa Catalina Mts. K. F. Parker 22
7274;
4/29/50
Volusia Co., Fla. North of Ormond Mrs. H. Butts, 20
Dec., 194 7
Highlands Co., Fla. |e. of § Sebrin R. Garrett 20
Arbuckle Creck 11/21/48
Flagler Co., Fla. i iitns R. B. Miller, 20
2/16/50
Marion Co., Fla. Ocala As — Miss L. E. Arnold 20
near ‘Eur 2/2/50
Putnam Co., Fla. 10 mis. s.e. af iniedachon. W. B. Fox, 20
11/18/51
Liberty Co., Fla. 3 mis. e. of Hosford on Kurz, 20
Hwy. #20 —_{tt/s/49
St. Tammany I mi. fot Slidell on L. Ewan 20
Parish, La. LaCombe Rd. 19236,
11/25/49 7
Wake Co., N.C. 2 mis. s. of Fuquay Springs | R. K. eis & 20
nU.S. #15 A pa
ee

+6

WOLAYOUUVY GIONUV AHL AO TYNUNOL

AXXX “10A |

C. tomentosus Perennial I |Pender Co., N. C. 10 mis. s. of Harrel’s R. K. Godfrey 20
herb store 12/22/51
C. corymbosus Perennial I Alachua Co., Fla. Gainesville Miss L. E. Arnold 20
herb II 11/ 19/46
V_ |Putnam Co., Fla. 10 mis. s.e. of Interlachen |W. B. Fox, 20
11/18/51
Trilisa paniculata Perennial II | Volusia Co., Fla. n. of Ormond Mrs. H. Butts 20
herb Dec., 1947
VIII |Leon Co., Fla. 23 mis. w. of Tallahassee H. Kurz 20
oe 11/5/49 oe
IX |Leon Co., Fla. 23 mis. w. of Tallahassee | H. Kurz 20
er, es Seen 11/5/49
T. odoratissima Perennial Alachua Co., Fla. Gainesville Miss L. E. Arnold 20
herb 11/19/46
IV |Highlands Co., Fla. | Along Jackson Creck, R. Garrett 20
near Se brin 11/10/48
V_ {Leon Co., Fla. 23 mis. w. of Tallahassee |H. Kurz 20
11/5/49
VI |St. Tammany I mi. w. of Slidell L. Ewan 20
Parish, La. 19237
11/25/49
T. carnosa Perennial III |Highlands Co., Fla. |w. of Sebring R. Garrett 20
herb 11/11/48
IV | Highlands Co., Fla. R. Garrett 20

wet pinelands
s. of Sebring

11/4/49

[ps6

II AVNIINHNM AHL NI SAIGALS “WASIVD

TABLE I — continued
Chromosome Numbers in the KuAniinae

Accession Chromosome
Growth Locality of Collector, Number
Name Form No County Collection No. & Date n
Kuhnia eupatorioides\ Perennial I |Kalamazoo Co., Sect. 29, Schoolcraft C. R. Hanes 18
var. pyramidalis herb Mich. Twsp. 9/17/47
II |Clark Co., Ky. 12 mis. e. of Winchester F, T. McFarland 18
10/18/47
III |Brazos Co., Texas 5 mis. s. of College H. B. Parks 18
Station 10/20/47
VII |Kalamazoo Co., Along L.S.M.S. Rwy., C. R. Hanes 18
Mich. Schoolcraft Twsp. 1648
10/5/48
K. eupatorioides Perennial I | Dallas Co., Texas 3100 Block on Rosedale L. H. Shinners 9 18
var. texana erb St., University Park, 11/11/48
Dallas
K. eupatorioides Perennial I |Roosevelt Co., Io mis. n.e. of Poplar M. Ownbey 18
var. corymbulosa herb Mont. 3229
8/30/49
K. Mosteri Perennial I |Alachua Co., Fla. Gainesville R. Garrett 9 18
herb 11/3/48
K. rosmarinifolta Perennial III |Santa Cruz Co., Shaded canyon, Patagonia | D.H. Darrow 9 18
herb Ariz. & Haskell
2317
10/14/44
V_ {Apache Co., Ariz. Near cultivated land, K. F. Parker 18
. John’s 7409
12/3/50

96

WOLAMOTUVY GIONAV AHL AO TWNUAOL

AXXX “T0A]

K. Schaffnert Perennial I Mexico, Mex. Hills of El nw. D. B. Gold & 18
herb of Huchuet F. Matuda
10/30/51
ae pil Perennial I jJalisco, Mex. Along rocky cut near top LO; Cae 9 18
ar. latisqua herb of barranca, Guadalajara 66
10/27/50
B. sessilifolia Annual I |Puebla, Mex. Riene dy slopes o ae ee L. O. Gaiser 9 18
Tlacoctli 6 km of
Izucar de Mec ghers: 10/31/50
II |Morelos, Mex. From the limestone mt. LO. Gilet 18
at Yautepec 7]
eee ee 11/3/50 ae
III | Morelos, Mex. ca. 23 kms. from Cuerna-_ | L. O. Gaiser 9 18
vaca on mt. coe on road |77
to Yaut 11/3/50
IV |Guerrero, Mex. Ep km +: brow Taxco, ae L. O. Gaiser 18
Mexico to Taxco Hwy.
11/4/50
Chromosome Numbers in the Subtribe Ageratinae
Mikania Vine II |Calvert Co., Plum Point S. F. Blake 38
scandens Maryland Autumn
1946
III |Flagler Co., Fla. Haw Creek Region Mrs. H. Butts 38
Autumn, 1946
VII | Highlands Co., Fla. | Rich wet soil, Sebring R. Garrett 38
11/15/48
M. cordifolia Vine I Highlands Co., Fla. | Highlands Hammock, R. Garrett 38
ate 11/23/48

[PS61

Il AVNIINHOAM AHL NI SAIGALS ‘YASIVD

98 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXXV

SEEDLINGS

1954 | GAISER, STUDIES IN THE KUHNIINAE II 99

of the specific name pseudo-liatris, given by Cassini to his type species of
the genus. While narrower, linear leaves are found in a number of species
of Liatris other than L. punctata, this is the only species of Carphephorus
possessing them. It is obvious that regardless of the shape and the struc-
ture of the root, development of a rosette of leaves is common to all and
is of rather prolonged duration. At least under greenhouse conditions, this
same Trilisa seedling has not varied much in appearance in more than three
years.

Figures D to G represent comparatively much younger seedlings of
Carphochaete Bigelovii 11, Barroetea sessilifolia 1, Kuhnia rosmarinifolia
V and Brickellia adenocarpa III. Only records were kept of the time of
sowing the seeds, so that from those dates, the Carphochaete is a little
more than a month old, the Barroetea is less than three months, the Kuhnia
a little more than two months and the Brickellia less than two months.
All have developed slender fibrous roots at this time but already show a
fast growing stem-axis with a number of nodes and internodes. This
similarity i in seedlings cannot be attributed to any likenesses of their actual
growth-form, for while this species of Barroetea is an annual and might be
expected to show very rapid growth, that of the KuAnia is an herbaceous
perennial, that of the Brickellia a shrubby perennial, and the Car phochaete
a small shrub.

The matter of chief interest was that there were two types into which
all of these seedlings can be classified: the rosette and the non-rosette.
II. Chromosome Number and Morphology
CARPHOCHAETE

Cells of Carphochaete Bigelovii 11 were found to have a noticeably
ereater amount of chromatin than any of the other genera studied. This
is due first of all to a larger number of chromosomes (2n = 22) but also
to the presence among them of a majority of larger ones. While the exact
size and form of all the chromosomes is not evident in figure 30, among
the thirteen peripherally arranged units can be distinguished some of the
long and medium classes.1 The V approximately at the center top and
the one almost at the center bottom, are designated medium in length
(Mm). in contrast to the two long V’s along the right (Lm), all of these
being medianly constricted. The latter two in turn are distinguished from

“The same abbreviations as have been used in Gaiser (1953) are here applied:
Using capital letters to express ae long chromosomes with median, submedian
and subterminal constrictions are represented as Lm, Lsm, ene has sp vias
chromosomes of median ee similarly constricted as Mm t;
short chromosomes similarly Sm, Ssm and Sst. The short ene ya became SS:
when medianly contricted ce and when terminally SSt.

EXPLANATION OF FIGURES OF SEEDLINGS
res pee seedlin oe of various accessions of different dates. Figs. A-C_ photo-
eraphed 5/9 /51. Figs. ee 26/51
ris punctata leer B. Carphephorus pseudo- wid : 1/19/50: C. Trilisa
ae VIII 11/22/49 Pare ichacte Bigelovit II 3/20 rroctea aed
I 2/26/51. F. Kuhmia ees V 2/15/51. G. Brickellia por rel Il 3/5/51

100 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXXV

two of similar length but having submedian constriction (Lsm), at right
and left at the top, and a pair along the left side, in slightly lower focus,
which are subterminally constricted (Lst). With the remaining marginal
V at the lower left clearly a long chromosome also (Lm), a total of seven
of the long class have been counted merely among the peripheral ones. As
frequently occurred, toward the center among others, are six shorter units
(Sm) of which two can be distingiushed at lower left. In a similar cell
(Fig. 29) of a Feulgen preparation, only one medium chromosome was
left with these six, all of which are short V’s and medianly constricted, to
fill the center around which fifteen long and medium ones were arranged
marginally. There is a sharper break between the six short ones and the
remaining sixteen than separates the equal number of long and medium
ones into which that number has been divided. In the short ones there was
little doubt that the constriction was median. The eight long chromosomes
were analyzed as to centromere position with less difficulty, than those of
medium length where there was some uncertainty. From the study of
material available of one accession the karyotype has been given in Table
II as: 2 Lm, 4 Lsm, 2 Lst, 4 Mm, 2 Msm, 2 Mst and 6 Sm.

GARBERIA HETEROPHYLLA

This monotypic species, restricted to Florida, was received through five
different collectors from as many different counties. Excellent material
was available for study and no variation was found. In a cell (Fig. 28)
of accession VI from Flagler County, the number of chromosomes can be
counted (2n = 20) and their morphology is also clearly recognizable.
Thus, by beginning as at twelve, and proceeding clockwise around the dial,
the marginal chromosomes represented in succession are as follows: Mst,
Lst, Sst, Lsm, Mm, Msm, Lm, Lsm, and Sm. In the cell there are a pair
of each of these classes except Mm, and of that there are three pairs. A
second cell (Fig. 27) in the same section, shows that two of the shortest
units, lower left and right margins, are heterobrachial. This was more
convincing in cells particularly favorabie for study of accession VII,
though not photographed to much better advantage (Fig. 26, 3rd from
left at top). By comparison with the cells of Carphochacte there is this
difference of the fewer (four rather than six) chromosomes of the short
class, and in their variation from the uniformly similar isobrachial type.
As for the rest, the same classes are present though not to the same num-
bers. With one pair of long of each of the three constriction types, rather
than a total of eight long ones, comes a further reduction in amount of
chromatin. This karyotype has ‘the sare variety: 2 Lm, 2 Lsm, 2 Lst,
6 Mm, 2 Msm, 2 Mst, 2 SM, and 2 Sst

CARPHEPHORUS

Two of the four species of Carphephorus were examined from at least
two accessions, while C. tomentosus and C. bellidifolius were limited to
one each. In all, the 2n chromosome number was found to be 20.

Of C. bellidifolius there were a number of excellent cells for study.

1954] GAISER, STUDIES IN THE KUHNIINAE II 101

From comparisons made with Carphochaete Bigelovii it was observed that
the chromosomes were all more nearly equal in length. Two pairs of me-
dium chromosomes one each with medium, Mm, and submedian, Msm,
constrictions, were almost as long as the long chromosomes. Also the next
to the shortest pair were not sharply different from the medium in length.
Frequently when the karyotype was assembled after an analysis of a cell,
there were odd numbers for these two classes. The slightly more condensed
chromosomes of figure 22 were well spread. Beginning as at seven and
proceeding clockwise to four, the marginal ones are: Mst, Mm, Mn, Lst
(pale, as slightly below focus), Lsm, Mm, Mm, Mst, Lst, Lm, Msm, and
Msm. The two latter and the pair, Mm, directly opposite, closely approach
the long ones. In this cell the shortest units are toward the center.

Preparations from seeds of Carphephorus tomentosus, which had been
received from North Carolina, were more adequate for study than photog-
raphy. As in the cell of figure 23, the chromosomes may be spatially
separated but slight overlapping of the ends confuses the picture. Under
the studies of this species the note had been made “remarkably few short
chromosomes.”

As was customary accompanying the studies of cells, diagrams onc.
corymbosus V were made and the pairs of chromosomes numbered in suc-
cession as to their lengths. With a total of twenty chromosomes, the num-
bers of the shortest pairs were 9 and 10. While the latter was sure to be
of the class Sm, opposite 9 there was frequently placed the question, “short
or medium.” In figure 24 for example, along the right side two isobrachial
chromosomes which are clearly medium in length, alternate with two which
are heterobrachial. These latter were numbered 9 and 8 respectively,
meaning that while the bottom one was one of the characteristic medium
pair with subterminal constriction, Mst, the second from the top was of the
questionable class.

Accessions of C. pseudo-liatris from Florida and Louisiana were com-
pared. Because tips were available from seedlings (see Fig. B) growing
in a pot, as well as from germinating seeds, the most convincing analysis
of the morphology of the chromosomes was made of this species. In figure
25, photographed when one each of the two shortest pairs are in focus,
a number of the longer chromosomes are foreshortened. Almost at center
bottom is a short medianly constricted chromosome, Sm, while the second
from the center top is heterobrachial. Frequent analyses left a question
concerning the members of this pair, just as had been the case in studies
of the other species. Certainly no marked difference was noted for any one.
Therefore one karyotype has been given for the four of them: 2 Lm, 2
Lsm, 2 Lst, 6 Mm, 2 Msm, 4 Mst, and 2 Sm. This modification of the
two last classes was made with hesitation but allowing for the factor of
human error, was accepted on the evidence of the camera.

TRILISA
Of several collections of Trilisa paniculata, made in South Carolina in
October, germination of seeds was very poor, though it was possible to

102 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

CHROMOSOMES
(Caption at bottom of facing page)

1954 | GAISER, STUDIES IN THE KUHNIINAE II 103

obtain the chromosome count, 2n = 20. More somatic divisions were ob-
tained from two accessions from northern Florida in November and a few
seedlings were grown (see Fig. C). While a majority of longer chromo-
somes in a peripheral arrangement was the more usual, in figure 19 some
of the medium and short units are seen in this position. Thus a better idea
of the comparative lengths of all the classes of units for this species is
obtained. At the top left corner, two medium chromosomes, the one sub-
terminally, Mst, and the other medianly, Mm, constricted, lie respectively

elow and above a long submedian one. The mate for the former appears
at the opposite right hand corner between two radiating long chromosomes
of subterminal constriction, Lst. Below these, one each of the longest and
shortest medianly constricted chromosomes are sharply contrasted and
followed by a similar one medium in length. The latter is one of the me-
dium class which approaches the long. Certainly this medium chromosome
and the long one are more nearly equal than it and the short one. In size
and form no noticeable variation from the chromosomes of the species of
Carphephorus was evident. Comparison of this cell with figure 24 of C.
corymbosus brings the camera’s confirmation. The same question arose
as to the length of the pair next to the shortest, but they were considered
to be heterobrachial and not unlike the class Mst.

Preparations were made from seeds of Trilisa odoratissima, known as
the Vanilla Plant, from Florida and Louisiana. The long chromosomes
were more prevalent and of the same form as in the previous species. Also,
there was no evidence that the medium and short ones differed. Because
of the abundance of cytoplasmic inclusions in the cells of this species, fewer
cells were suitable for photography. When the chromosomes were wel
spread, globules were often overlying. It would have been better to have
made all preparations by the Feulgen technique. Figure 20 is of a cell
just back of the meristematic apex and the chromosomes are therefore
more crowded. One karyotype is believed to be common to these two
species and it has been found to be indistinguishable from that of Car-
phephorus.

EXPLANATION OF FIGURES

~7, 9-30, are from near comparable a aa plates in root-tips. Fig. 8 is of an
ae oer ye plate 1, 2, 3. Barroetea sessilifolia 1. 4, 5. B. subuligera var. pps heise’ i
. pyramidal as), 9. (II fro

6-10 = 8. Il fr

tucky), 10. (VII zen Michigan). 11. K. age wens var. fexana T. T2-aKe baie alias
var “corymbuora | 13. K. Mosieri 1. 14-16. K. rosmarinifolia; 14, 15. (of accession IIT),
16. of V. 17, “K. Se haffneri : 19. Trilisa - aniculata VIII. 20. Pai: odoratissima IV.
21. Mikania oe HT +22. Carphephors bellidifolins Il. 23 tomentosus 1. 24.
C. corymbosus V. 25. C. pseudo- eet . 26-28. Garberia Lee 26. of accession
VII, 27, 28. of VI. 29-30. Carphochaete Bigelovi Il.

Preparations made of sere: taken from plants, I in Belling’s at the same time and
stained in toto in Feulgen’s, Figs. 5, 11, 12, 25, 29. Root-tips of seeds after Belling’s and
stained in N.G.V., Figs. 16, 24; after Karpe chen ko’s and stained in Feulgen’s, Figs. 9, 18,
27, 28. All the rest were of seeds fixed in Karpechen to" s and stained in N.G.V. The photo-
micrographs were taken with the use Sie a Zeiss microscope and a Homal IV lens. All ene
2300x. Reduced 1650x in sagt

The prepara = n of the es a — aided by a grant from The Society of Sigma Xi,
which the author early ost ot pena It is a pleasure to acknowledge the ee
of Mr. Paul Brown in the photomicrography.

104 JOURNAL OF THE ARNOLD ARBORETUM _ [vot. xxxv

The rarer species, Trilisa carnosa, limited to Florida, which had been
thought to merit new generic rank by Small, was received from the same
locality near Sebring, three times from October to November, 1948, and
again in November, 1949. The difficulty of germinating seeds of this
species was certainly not overcome during the time and with the material
available. Only a few seeds ever appeared to begin growth and none de-
veloped very far so that an extremely limited number of mitoses were
found. The number (2n = 20) and approximately the same kinds of
chromosomes as found in the other two species were sketched from cells
not adequate for photography. However, lacking sufficient material for
careful studies, the karyotype of this species cannot be included at this
time.

KUHNIA

Three of the four American species given by Shinners (l.c.) have been
available for this study. Accessions had been received of the largest, most
widely distributed species, K. eupatorioides L. from the states of Michigan,
Kentucky, Texas and Montana, which, according to Shinners (l.c.) would
represent all the varieties of that species except ozarkana Shinners. Those
from the first three sources were variety pyramidalis Raf. and from the
last, var. corymbulosa T. & G. while Dr. Shinners himself contributed a
packet of seeds of his variety texana. A glance at figures 6 to 12 of this
polymorphic species, shows a chromosome plate of 2n = 18, in which there
are lacking any of the long chromosomes, so characteristic of the genera
here previously discussed.

K. EUPATORIOIDES variety PYRAMIDALIS. From preparations of par-
ticular clarity, stained with Newton Gentian Violet five years ago, the
sizes of the chromosomes can be shown. At the lower left in figure 6, a
parallel pair of the shortest and longest units, both medianly constricted
can be compared with a single chromosome of intermediate length, lying
almost horizontally above them. The units of this figure are probably all
a little more condensed than those in figure 7, which is of a cell in a
similar preparation. In the latter, the two longest chromosomes are at
outer left and right and are submedianly constricted. From such figures,
it is clear that the longest chromosomes would have to be placed in the
medium class, according to the scale of sizes here used. One might then ex-
pect the shortest chromosomes, since there are units intermediate in length,
to be two classes removed and to fall into a short short class, as adopted
for some species of Brickellia (Gaiser 1953). Considerable effort was made
to decide this question by comparison of various stages and of cells of all
the accessions. Figure 8 is of an anaphase plate of a recent mitosis as can
be judged by the parts of two chromosomes outlying in the cytoplasm at
the right, belonging to the other closely overlying plate. The chromosomes
are more slender but again they do not cover three distinct classes of
length. It is considered that the karyotype includes medium to short
chromosomes, and when there are nine pairs of such closely approximating
lengths there is difficulty in deciding the boundary between the two classes.

1954] GAISER, STUDIES IN THE KUHNIINAE II 105

In figure 8, two chromosomes with very small terminal bodies are seen,
one at center bottom and the other at center left. More frequently cells
were found in which only one such satellite was visible, as is shown at the
end of the curved chromosome at upper right of figure 6 and at the inner
end of the chromosome stretching toward the center from the left of figure
7. It is believed these chromosomes are next in length to the two longest
pairs. By careful analysis of numerous cells of other accessions as well as
this, next in succession of length are: 2 Msm and 2 Mm, which more
closely approach the aforementioned three pairs than do the other 4 Mm.
Whether the latter should be classed as short may be debatable. The four
shortest are considered 2 Sm and 2 Sst. In preparations of accession II
from Kentucky (Fig. 9) as well as accession VII from Michigan (Fig. 10)
this analysis was also favored.

K. EUPATORIOIDES variety TEXANA. In the second accession from Texas,
the figures are very much the same. Cells without any visible satellites
were more frequently found than those showing them even delicately as in
figure 11 (the second chromosome on the upper left) from a Feulgen
preparation.

K. EUPATORIOIDES variety CORYMBULOSA. The western variety was re-
ceived from Montana. Well spread chromosomes in the plate of such a
cell as shown in figure 12 help to confirm the previous analyses. At the
center on the left side is the medium chromosome with a satellite appearing
knob-like at its inner tip and close to its centrally placed mate. Above
them, occur one each of the longest of Msm and Mm respectively. The
curved pair at center bottom are the second pair of so-called Msm while
the two overlapping as a V at top are second in length of Mm. It is
evident from such a preparation that no sharp line separates the classes
of chromosomes. However, on the basis of comparison with those of the
other genera the karyotype is believed to be: 8 Mm, 4 Msm, 2 M with
sat., 2 Sm, 2 Sst

Of the two other American species of Kuhnia included, the first is a
restricted endemic in lower Florida. Two cells of K. Mosieri Small,
which the chromosomes were well separated by microscopic pcre,
could be seen in the field of the microscope at one time. Starting with the
longest chromosomes, successively those next in size were sketched in
different colors and the two cells constantly compared. Unfortunately
they did not photograph as well as it was possible to draw them, to add
to this interesting exercise. Of course there were points of indecision.
However, even before it was recognized that in one of these (Fig. 13,
center left) a chromosome with a satellite was represented, its arrange-
ment in the order of consecutive chromosome lengths coincided with that
given to such in K. eupatorioides. The karyotype of these two species
were found indistinguishable. The second species is commonly known as
K. rosmarinifolia Vent. Shinners (l.c.) considered this name a “nomen
confusum” and put it in synonymy with K. chlorolepis Wooton & Stand-
ley. Blake (1942) did not consider it necessary to adopt this name in his
treatment of the genus for Arizona and so I have not, as both accessions

TABLE II

Karyotypes in Species of Genera of Kuhniinae
Chromosomes
Accessions _— __Number of pairseach
ae Studied | Total M iain
Number ith
Lsm |. Lst} Mm} Msm| Mst | sat. | Sm Sst
Carphochaete Bigelovit II 22 2 I 2 I I 3
Garberia heterophylla IV V VI VII 20 I I 3 I I I I
Carphephorus pseudo-liatris II 20 I I 3 I 2 I 1 Mst = S$?
C. bellidifolius II 20 I I 3 I 2 I
C. tomentosus I 20 I I 3 I 2 I
C. corymbosus IV 20 I I 3 I 2 I
Trilisa paniculata VIII 20 I I 3 I 2 I 1 Mst = S?
T. odoratissima IVV 20 r | 1 3 I 2 I
uhnia eupatorioides and 1 Mm the
var. pyramidalis I I II VII 18 4 2 I I I anes other Msm and
another Mm next in length.
K. eles
var. tex I 18 4 2 I I I
K. aoe
var. corymbulosa I 18 4 2 I I I
K. Mosteri I 18 4 2 I I I
K. rosmarinifolia IlI V 18 4 2 I I I
K. Schaffneri I 18 4 2 I I I
Barroetea ——
var. latisqua I 18 4 2 I I I Ditto
B. sessilifolia I III 18 4 2 = I I I

901

WOLAYOPUV GIONUV AHL AO TYNUNOL

AXXX “10A |

1954 | GAISER, STUDIES IN THE KUHNIINAE II 107

here studied came from that state. Of the preparations of K. rosmarini-
folia III, the same repetition of statements could be made, cells showing
a small satellite less frequently seen (Fig. 15, at the center top), than those
without them (Fig. 14). Of accession V of this species, a plate of particu-
larly condensed chromosomes was photographed (Fig. 16) to show the
narrow gap between the shortest and longest units. Thus, for example,
comparison with figures 25 (Gaiser 1953) of Brickellia microphylla where
the chromosomes were shortened by paradichlorobenzene treatment, or fig-
ure 42 of B. glomerata, where a similar drastic shortening accidentally
happened in one cell of a Feulgen preparation, emphasizes that there is
a greater spread in the species of that genus where the karyotype includes
long chromosomes.

One of the three species, which do not occur north of Mexico, K. Schaff-
neri Gray, was available for this study. Because the type sheet contains
also a root of some legume, Shinners (l.c.) renamed the species K. micro-
phylla, but Blake ® considers this unjustifiable.

In a very clear early metaphase plate (Fig. 17) from a Newton Gentian
Violet preparation, the range in size of the chromosomes is shown. One
of the longest chromosomes, submedianly constricted (Msm) is at the
upper right corner and two of the shortest ones, medianly constricted
(Sm), can be distinguished as smaller V’s among the central group, almost
at center top and bottom. At this focus, one arm of several of the marginal
V’s has been foreshortened. Thus, for example, the chromosome at lower
right is the other of the longest pair (Msm) and in succession the other
two at center bottom represent medium chromosomes of the next category,
Mm and Msm respectively. The straight medium chromosome, which is at
a slight angle above the latter, is terminated toward the center by a small
knob and is the equivalent of the chromosome with a satellite. The same
can be seen in a cell of a Feulgen preparation (Fig. 18) at the tip of the
chromosome coming into the small central upper space.

By comparison of figure 17 with that of K. eupatorioides var. corymbu-
losa (Fig. 12) of a cell similarly prepared, it might appear that the chromo-
somes of this species are generally longer, but this is merely so because
it is a slightly earlier metaphase stage. It is worthy of note that in both of
these cells there is a knob-like body at the end of a pair of the chromo-
somes. That its appearance cannot be attributed merely to this one kind
of preparation is proved by the use of the same kind (e.g., Newton Gentian
Violet) in illustrating K. eupatorioides var. pyramidalis (Figs. 6, 7, 8) and
K. rosmarinifolia (Fig. 15) where it is seen separated from the chromo-
some. Yet the same is also visible in Feulgen preparations (see Figs. 11,
18). In summary, in any kind of preparation of all of the species of
Kuhnia here studied, this very small terminal body, which has been called
a satellite, may be found, whereas in similar preparations of the previous
genera, no evidences of such has ever been seen. No variation from the

5 In litt.

108 JOURNAL OF THE ARNOLD ARBORETUM _ [vol. xxxv

karyotype given for K. eupatorioides has been found in the three other
species studied.

BARROETEA

The one perennial species which is somewhat woody at the base, B.
subuligera (Schauer) Gray, is known from the northern boundary to
Hidalgo, a central state in Mexico. The larger variety latisgquama Green-
man, was described from a specimen collected in Jalisco. A plant of this
was collected in the barranca at Guadalajara and from seeds of it, a
seedling was grown for a time in the greenhouse. From pollen mother cells
as well as root-tips, the chromosome number was found to be the same as
in the species of Kuhnia studied, n = 9 and 2n = 18

A comparatively early metaphase of a root-tip cell shows the form of
sixteen of the chromosomes, but two short ones which were lying one above
the other, resulted in the oval body at the upper right of the figure (Fig.
4). One chromosome of greatest length, submedianly constricted, is seen
at lower right and a second in length, medianly constricted, is plainly
contrasted at center top with one of the shortest (Sm). By the scale of
sizes used for the other genera, neither of the former would exceed the
medium class. Thus the karyotype includes medium to short chromosomes
only. In this cell a number of the heterobrachial chromosomes are found
around the periphery and may appear to be more numerous than in a very
comparable stage of a Kuhnia, e.g., Kuhnia Schaffneri (Fig. 17). How-
ever, their number is not greater and in other cells where the chromosomes
are a little more condensed (Fig. 5) no confirmation is found of this differ-
ence. While no chromosome with a satellite is shown, its equivalent is
probably represented in one of medium length, subterminally constricted,
for the free terminal body was seen in some other cells.

Of the annual species, B. sessilifolia Greenman, seeds were collected
from four different localities of three of the central states of Mexico and
from root-tips of each, the chromosome number was found to be the same
as that of the perennial species, 2n = 18. It was also possible to grow
seedlings (see Fig. E) from two of these accessions and similar meiotic
divisions were obtained, n = 9.

Figure 1 of B. sessilifolia III shows a metaphase stage comparable to
that of figure 4 of B. subuligera and the sizes of the chromosomes are very
similar. On one of the longest chromosomes, at upper left, a satellite is
conspicuous. In cells of slightly more condensed chromosomes the same
is visible (Figs. 2, 3, lower right). Examination of a number of cells of
B. sessilifolia I resulted in the same analysis of medium to short chromo-
somes including one of the former showing a satellite, both in Feulgen
and other preparations, either in cells of root-tips taken from the plant or
from germinating seeds.

From this study of two species of Barroetea, it was found that they were
more similar to, than different from, the species of Kuhnia seen. The
points of likeness consist in having: the same number of chromosomes:
chromosomes which range from medium to short across an indefinite

1954] GAISER, STUDIES IN THE KUHNIINAE II 109

boundary, and, one pair of the medium chromosomes showing a satellite.
Certainly no constant difference in the karyotype was detected. It is given
as: 8 Mm, 4 Msm, 2 M with satellite, 2 Sm and 2 Sst.

SUMMARY ON CHROMOSOMES

As summarized in Table II.

1. Carphochaete is the only genus having 22 chromosomes.

2. All the species of the three genera, Garberia, Carphephorus and Trilisa
have 20 chromosomes, in common with a majority of the species of Liatris.
3. Four of the seven species studied of Kuhnia and two of the species of
Barroetea have 18 chromosomes, the same number that had been found
in all of the examined species of Brickellia.

4. Monotypic Garberia and the one species studied of Carphochaete, are
both distinct in their karyotypes. The latter has a greater proportion of
long chromosomes with three pairs of isobrachial short chromosomes,
while in the former, one of two short pairs are heterobrachial.

5. Carphephorus and Trilisa could not be distinguished by their karyo-
types. Yet they varied from Garberia in a closer approach, if not equality,
of the shortest heterobrachial pair to the medium class.

6. Long chromosomes are absent in the five species of KuAnia studied, the
karyotype consisting of medium to short ones. A pair of medium chromo-
somes have small satellites.

7. A similar closely graded series of short to medium chromosomes, of
which one pair bears satellites, makes up the karyotype in the two species
of Barroetea studied.

III. Trichomes.

As the type of B. Pavonii Gray is in the Boissier Herbarium, only six
of the seven species of Barroetea could be included in this study of the
trichomes of the leaves. Five were found to vary slightly in the abundance
of non-glandular trichomes which are uniseriate (Metcalfe & Chalk 1950)
and of the acuminate type according to the classification of Brickellia
(Gaiser 1953). The size and the thickness of the wall of the trichome
vary very little from species to species. Figure 31 represents that of the
annual B. sessilifolia and fig. 32, generally a slightly larger form, that of
B. subuligera var. latisquama, the perennial considered somewhat woody
at the base. The sixth species B. glutinosa Brandegee, with leaves de-
scribed as densely glandular-puberulent, was found to lack this type of
trichome but to have the same biseriate glandular form described as being
common to a number of species of Brickellia. Along the leaf margin these
might be found from a few to seven or eight cells high (Fig. 33). If seen
edgewise they appeared as a stipitate somewhat globular organ (Fig. 34).
When the flat expanse of the leaf was examined, one saw their apices as
two typical hemispherical cells (Fig. 35) or by focussing a little lower as
rounded cells above the level of the leaf surface. Figure 36 was taken at

110 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxv

a still lower focus where the epidermal pattern, including a stoma, became
evident, but the foreshortened trichome was merely a dark blur. Checking
leaves of the other species for this type of trichome, it was found only
very occasionally on the underside of the midrib near the petiolar attach-
ment. However, the depressed gland, as used also for Brickellia (Gaiser,
l.c.), and the punctate condition on the lower surface was common to all
except B. glutinosa where it was lacking on the lower as well as the upper.
As in Brickellia, this glandular trichome is a longer stalked condition of the
short form arising from the epidermal layer in depressions below what is
normally seen as the leaf surface. When a species characteristically de-
velops the elevated form, none or very few, if any, are seen remaining
sunken below the leaf surfa ce

No evidence was seen of ‘the smaller uniseriate capitate glandular
trichomes in any species of Barroetea.

All the species of Kuhnia, according to Shinners’ classification, were
represented and showed a little more variation in trichomes than those of
Barroetea. On the upper surface of leaves of K. adenolepis they were the
shortest, consisting of no more than two or three thick-walled cells, arising
from a very heavily cutinized epidermis (Fig. 37). Those of K. Mosieri
were comparatively short, but consisted of thin-walled cells, almost uni-
form in diameter, so that the trichome tapered very little (Fig. 38). On
K. Schaffneri (Fig. 39) a more attenuate trichome, of approximately the
same length but with heavier walls, showed a tendency towards the
moniliform type of rounded bead-like cells. In all those having thick walls
this character was prevalent as shown in the uneven-walled form of the
scabroid leaf of K. rosmarinifolia (Fig. 40). All the succeeding longer
ones have been figured at the same magnification (220 ) just as all
the shorter ones had, though they were more enlarged (530 *). While
those of K. eupatorioides var. pyramidalis, taken from our accession from
Kentucky (Fig. 41) resembled that of K. Mosieri in the thinner and
straight walls, those of varieties texana, ozarkana and corymbulosa, with
successively thicker and rougher walls (Figs. 42-44) were attenuate and
more moniliform. Those on the scabroid leaves of K. leptophylla var.
mexicana (Fig. 45) and K. oreithales (Fig. 46) did not differ much from
the latter. In these two species an occasional cell filled with an aggregate
crystal was more commonly seen than in any other species. In all of them
the apical cell is longer than the rest but would not be recognized as a cap
cell, as in some species of Brickellia.

On no leaf preparation examined of any of the species of Kuhnia was
evidence seen of the biseriate glandular trichome. By contrast the de-
pressed gland and punctate condition was common to all species except
K. adenolepis, and to ite surfaces of the leaf, though generally it was
less abundant on the u

What in Brickellia had oe called uniseriate capitate glands, on stalks
only a few cells in height, prevailed in all species. Generally they were
small as in that genus, requiring a larger magnification for observation as
well as photography as shown in Figure 47, 1100 *, of K. Mosieri. An

1954 | GAISER, STUDIES IN THE KUHNIINAE II 111

exception to the general was found in K. adenolepis. This species had been
distinguished by Robinson (1911b) because of its unique glandular cili-
olate phyllaries, the glands being described as subsessile and black. The
leaves had been described as “‘puncticulatis’” on both surfaces. Very short
trichomes have been described for the upper surface (Fig. 37). Examina-
‘tion of a cleared preparation of the type specimen revealed more abun-
dantly on the lower surface, a gland that is a little larger than the previous
(Fig. 48, 530 >) and distinctive for its dark apical cell when the leaves
of all species had been cleared similarly. It is possible that the apical or
secretory cell in this species has a different content, as not only was it
dark but as shown in the figure some of its secretion had plugged and
darkened some of the sunken stomata, from which it had not completely
dissolved. From the very evident secretory function of the apical cell of
this uniseriate small type of trichome, which is so similar in form to
that seen in Brickellia (Fig. 87, 88, Gaiser 1953), I have considered them
glandular trichomes in these two genera as did also G. Fischer (according
to Solereder 1908, original thesis not seen). Vuillemin (1884) had done
likewise in Cynara and Echinopus, though there are no figures of those
for comparison. They are quite unlike the bladder-like trichomes observed
by Volkens (1887) in Zollikoferia nudicaulis, which are considered to be
full of cell sap (Solereder 1908, Fig. 103E). In form, they resemble some-
what more the capitate hairs described by Rosenthaler and Stadler (1908,
Fig. 28) for Cnicus benedictus, consisting of six to twelve cells, with an
elongated terminal cell which were said to differ from the non-glandular
hairs in having a richer content.

Two genera, Garberia and Carphochaete had been established as having
entire leaves. Neither the single species of the first nor any of the four
of the latter had trichomes comparable to those illustrated for the pre-
vious genera. They both had a more slender filamentous form somewhat
prostrate over the surface of the leaf.

When Bartram first described Garberia under the name Cacalia hetero-
phylla (see p. 90), he wrote of the fleshy leaves: ‘“‘of a pale whitish green,
both surfaces being covered with a heavy pubescence and vescicular, that
when pressed feel clammy and emit an agreeable scent.’’ It was no sur-
prise then when both surfaces gave the familiar appearance of the punctate
condition at a focus when stomata were clearly visible (Fig. 49) and of
the paired hemispherical cells of the depressed gland (Fig. 50) by sub-
surface focusing. In addition, very narrow delicate filaments which ta-
pered slightly if at all, and had cross walls at rather long intervals, lay
twisted over the epidermis and proved difficult to follow to their origins.
When the margin of this thick, entire leaf was examined it was possible
by careful focusing to follow the epidermis as it alternately rose and fell
into small pocket-like areas. In each depression was a tiered organ, two
cells wide and approximately six cells high (Figs. 51, 52) familiar as the
glandular trichome seen upon the leaf-surface of Barroetea glutinosa (cf.
Fig. 33). In Garberia these arose in the epidermal layer from smaller
cells which were clearly differentiated by their size. The first cells above

112 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

EXPLANATION OF FIGURES OF TRICHOMES

Figs. 31, 32, 37-46, 44-53, 56, 73. Non-glandular ne aoe Figs. 3s 36, 66. Glandular

bee a Seah s. Figs. 47, 48. Non-glandular uniseriate trichom
31. Barroetea ge ay B. subulger var. 5. latequama 33- 36. B . glutinosa. 33, 34.
Along the leaf ia | surface. 36. At the epidermal surface.
37. path dence pps 38 ert, Scha ffnert. 40. K. rosmarinifolia. 41-44.
K. eupatorioides. 41. of fae byramidalis, 42. 3 var. texana, 43. : var. ozarkana, 44. of
var. beth losa 45. K. leptophylla var. mexicana. 46. K. oreithales. 47. K. Mosteri
ss - ade nolepis. 56. iis a acai corymbosu 66, Trilisa pani 73. T. carnosa.
1 these — crographs were made from ge ae leaves a Zeiss microscope

ee a Homal IV lens, oo. sa 32, which is r10x. Figs. 38— 4 56, 66, ea 85x. Figs. 31,
33-37, 48, 200x. Fig. 47, 4

1954 | GAISER, STUDIES IN THE KUHNIINAE II 113

cane

=

ET ; ,
“i ~.

Se oy Sey 5

@ roe # : fom S y

ay : Five ry snk $3 Be ey io FP 3 Kile ave ro

EXPLANATION OF FIGURES OF DEPRESSED GLANDULAR TRICHOMES

oy Sysell ni V :t

Figs. 49, 53, 57, 61, 64, 67, 69, 70, 74, by focusing at the epidermal ser See 50,
54, 58, 60, 62, 63, 65, 68, 71, 75, by ‘fo ocusing below the epidermal surface. eS E52;
55, 59, 72, lateral view seen along the margin o yi epidermal surface

49-52. Garberia heterophylla. 53-55. Carphoch S Hees ovil. 57-60. mee ephorus to-
biat ins 63-65. C. corymbosus. 61, 62. C. blll 67, 68. Trilise paniculata. 69.

T. odoratissima. 70-72. T. carnosa. 74, 75. Liatris p

“All these arti were made from icine 1 vase by use of a Zeiss microscope
and a Homal IV lens.

114 JOURNAL OF THE ARNOLD ARBORETUM _ [vot. xxxv

these had thicker walls which generally reflected the light more than the
thinner-walled upper cells. Alongside such a trichome frequently a slender
filament could be traced until it pushed up to the leaf surface. In Figure
52 part of such a filament is recognizable. after it emerged above the leaf,
though it is seen less clearly along the right wall of the small depression.
Closer microscopic observation of the surface of the leaf then made in-
telligible what had been baffling in several other genera as well. In addi-
tion to the paired hemispheric cells seen when focusing below the surface,
an additional small round cell (shown at the bottom in figure 50) ac-
counted for the delicate emerging filament seen on the surface. Thus they
are really very delicate uniseriate trichomes of long, narrow, thin-walled
cells growing from the sunken epidermal layer alongside the biseriate
glandular trichomes.

Volkens (1890) described scattered groups of trichomes consisting of
five or six which were uniseriate with whip-like tips, surrounding one cen-
tral unstalked, biseriate, bladder-like gland (Pl. 8, Fig. 1) on the upper
surface of the leaf of a plant labeled “Baccharis Richardifolia.’’ A second
species of the Astereae, Olearia Hookeri, was said to be very similar while
on the other Compositae that he examined, the uniseriate elements were
increasingly fewer. In two genera representing the Eupatorieae, Eupato-
rium and Symphyopappus, these trichomes might occur occasionally but
lacked the characteristic thread at the tip. From his figure 4a, showing
two hemispherical cells of Symphyopappus, it appears that the biseriate
gland was depressed with only the uniseriate emerging above the surface.
Thus while the latter (Fig. 4b) appears more torulate, the grouping might
be similar to that described here for Garberia.

Carphochaete Bigelovii had been described by Gray (see p. 90) as
puberulous and resinous punctate. On first examination of the leaf prepa-
ration with lower magnification, very numerous depressed areas were seen
which were not all equal in size. Closer examination of one of the smaller,
revealed besides the two large appressed hemispherical cells, several
accompanying small round ones as the one seen in Garberia. The deli-
cate filaments, of which the latter are the bases, were not as long and
therefore did not appear as abundant over the surface. The larger areas
might merely show central cells of a greater diameter but frequently they
were the result of the confluence of two pairs (Fig. 54). In this figure
there are in addition, the bases of three filaments, two of which are seen
at the surface in figure 53. Since the leaves of this species are small but
rigid it was possible again by focusing along the margin of a prepared
leaf to obtain a lateral view of the gland (Fig. 55). Only the lowermost
pair of cells were thick-walled in comparison with the several in Garberia.
The walls of the cells immediately above were very thin and made it
difficult to show that there was but one intermediate pair. It was clear
that the uppermost secretory cells were very much expanded, thus con-
tributing to their larger surface-appearance.

Although one of the four known species, C. Wislizeni, had been de-
scribed as “not sprinkled with resinous dots,” a leaf of a Pringle specimen

1954 | GAISER, STUDIES IN THE KUHNIINAE II iS

(#765) from the mountains near Chihuahua, where the type was col-
lected, was found to be very similar to C. Bigelovii. All four species
might differ slightly in the abundance but not in the form and the kind
of contents in the depressions.

The species of Carphephorus may have the small, upper cauline leaves
pubescent when the lower and radicle leaves are nearly glabrous. This
is not a character peculiar to the genus, Solereder (1908) having general-
ized that in many genera only the youngest leaves, branches and leaf
petioles are covered with glandular trichomes. Thus much would depend
upon the choice of leaf for preparation. Of Carphephorus, in common
with the other genera, lower larger leaves were ee leaving to
Trilisa paniculata the comparison of lower and upper ones.

orm, the non-glandular trichomes on the ee and along the
margin of the basal leaves of each of the four species of this genus were
of near uniform type, of broader cells than seen in species of Kwuhnia.
Though they varied in length and to a minor degree in thickness of wall,
being thickest in C. tomentosus, the cylindrical shape of the component
cells was remarkably constant. One of medium length of C. corymbosus
is illustrated (Fig. 56).

It was in the punctate condition of the leaf that a greater difference
was noted, as is reflected in the descriptions given by Torrey and Gray
(1841): C. tomentosus, punctate; C. pseudo-liatris, sparsely punctate;
C. bellidifolius, punctate with scattered impressed dots; C. corymbosus,
sometimes obscurely punctate. From these phrases it was not clear
whether the difference was merely one of abundance. Examination of
these leaf preparations showed a considerable difference in glandular
structures.

On either surface of a lower cauline leaf of C. tomentosus, the paired
cells characteristic of the depressed gland were seen abundantly when fo-
cusing below the surface (Fig. 58) in contrast to the craters seen when
stomata are in view (Fig. 57). As described in Garberia heterophylla, the
two hemispherical cells were accompanied by small marginal circular cells
which had been recognized as the median optical view of delicate filaments.
Frequently two or three were found and since they appear in figure 57
they obviously reach the surface though none were found spreading over
the epidermal cells. Looking along the margin of one of these lower nearly
glabrous leaves which is entire, the lateral view of the biseriate trichome
with accompanying filaments at the side was seen in the depressed areas
(Fig. 59). To this species then, the term punctate, as understood for the
previous genera, could certainly be applied. The depressions were con-
spicuous macroscopically and that may be facilitated by their greater size
for it was found that often two glandular trichomes, each with several
laterally placed filaments were within the margin of one depression (Fig.

In the preparation of the other three species the surface pattern ap-
peared quite different. In a similar leaf of C. bellidifolius there was no
sign of the depressed gland of paired cells. Instead, one circular cell,

TAB
Types of Trichomes and Glands in the Epidermis of Leaves of Genera of Kuhniinae

Name

Barroetea sessilifolia
B. subuligera var. latisquama

. corymb
K. a
. Mosteri

K. Schaffneri
K. oretthales
K. — var. mexicana

adenolepis
C arphochaete Bigelovit
. Wishize
C. Graham
C. Schaffneri
Garberia heterophylla
Carpheph
. corym

orus tomentosus
bosus

C. bellidifolius
Trilisa at
- — bate

Elevated above the surface

Non-Glandular

Depressed (i.¢., punctate)

uniseriate,
trichomes

uNniseriate,
api

e
lower

cuminate

occasional

nearly uniform
attenuate towards
moniliform

ditto

scabroid moniliform
scabroi

slightly moniliform

scabroid moniliform
scabroid moniliform
very short

Xx 2xXX MK

KKK K KKK KX XK X

nearly uniform
nearly uniform

nearly uniform

Biseriate

upper lower
occa

on upper cauline

Glandular
eo is Biseriate er uMiseriate
Biseriate uniseriat er in groups I to 4
upper lower upper lower upper lower upper lower
x
x
x
x
x
x
x x
x x
x x
x x
x x
x x
x x
x x
x x
i. 7 x x
x x
x x
x x
x x
x 4 a
x x
x x -
x x

Depressed
(but not punctate)

secretory,

un
upper

.

iseriate
lower

:

x,
occasional

a as trichomes as sccn by microscopic examination of cleared Icaves from annotated herbarium specimens at the Gray Herbarium. > indicates presence of.

1954] GAISER, STUDIES IN THE KUHNIINAE II 1

larger than those accompanying the gland in C. tomentosus, appeared en-
circled by other leaf cells (Fig. 62). By focusing exactly on the surface,
when stomata are visible, there was again a crater-like appearance in its
place (Fig. 61). This gave confirmation that we were seeing the punctate
equivalent for this species which had been characterized as “compressed
dots,” likely because they are smaller.

In C. corymbosus a similar arrangement of cells was characteristic but
the center was usually composed of more cells. Figure 63 shows a fre-
quent linear arrangement of three. When there were four such cells they
conformed to the arrangement common to four spheres and the surround-
ing leaf cells had a circular form (Fig. 65). At the epidermal level with
stomata visible, this also had the appearance characteristic of a punctate
condition (Fig. 64).

In several preparations of C. pseudo-liatris the little evidence seen of
glandular punctation indicated similarity to C. corymbosus.

Thus all four species are punctate, indicating the presence of a depressed
gland or short glandular trichome. In C. bellidifolius there was a single
or uniseriate condition while in C. corymbosus and C. pseudo-liatris there
might be from one to a group of four of the uniseriate form. In C. toment-
osus there was the familiar biseriate trichome accompanied by the uni-
seriate similar to Garberia.

From the time of Walter’s original descriptions of the two species of
Anonymos (see p. 91), Trilisa paniculata has been distinct from T. odora-
tissima in having a viscidly pubescent stem and upper cauline leaves. For
this reason some of the latter as well as the lower glabrous leaves and in
addition phyllaries of the inflorescence of this species were cleared. On
both the phyllaries and small leaves, typical biseriate glandular trichomes
were found (Fig. 66) very comparable to that figured for Barroetea gluti-
nosa. Examination of either surface of a basal nearly glabrous leaf showed
neither these glandular trichomes nor any trace of the typical depressed
gland of two hemispherical cells. Instead rosette-like arrangements around
a central cell were seen when viewed at a sub-surface focal level (Fig. 68).
This figure does not appear unlike that of Carphephorus bellidifolius (sec
Fig. 62). However, when viewed at the surface the epidermal cells clearly
adjoin the specialized cell (Fig. 67). This is confirmed in figure 69 of
T. odoratissima, photographed at the surface and showing more clearly
the secretory droplets. There is no indication of any marked depression
in the epidermis of either species. This explains the inclusion of “not
punctate with impressed dots” in the description of section Trilisa Cass. of
Liatris by Torrey and Gray (l.c., p. 76) since they included only these
two species at that time. At the surface the specialized cell hardly ap-
pears to be free from the surrounding cells of the epidermis. However,
the fact that a central cell remains in focus from that layer through the
next below, which shows distinct encircling cells, indicates that it is a
secretory organ with a greater depth than one epidermal cell. None of
these distinctive cells was found close to the thin margins of the leaves
of these two species and so could not be studied laterally as was possible

118 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxv

in the preparations of Garberia, Carphochaete and Carphephorus toment-
OSUS.

Trilisa odoratissima, commonly named the Vanilla Plant for the fragrance
of the crushed leaves, has been used in the tobacco industry and is included
in lists of drug plants. Therefore references ® to the study of the structure
of its leaves have been found in pharmaceutical journals.

Paschkis (1879) was probably the first to study the leaves microscop-
ically. He described the upper and lower epidermis as including yellow
spots, .015 mm. in size, which were surrounded by regular small epidermal
cells. From tranverse sections he found these to be glands, in small clefts,
consisting of a basal stalk cell wedged in between the two adjacent epi-
dermal cells, and three to four additional ones. These rise just to the level
of the other epidermal cells. He makes a point of their close envelopment
by the thick cuticle of .008 mm., and as his figures show, the small de-
pression in the epidermis is closely filled. This undoubtedly explains my
failure to see any crater-like opening over the gland in surface view. The
secretion, which is of such a pleasant odor when expressed, is due to
cumarin. The diagram in Higley (1893) is very similar to the above de-
scription as is also the figure in Hanausek (1912) who called it a glandular
trichome. From the relation to the leaf surface it has its origin as a de-
pressed form. Among the multiple types found in these genera it is here
termed uniseriate and not punctate.

In the leaves of the recently added third species, T. carnosa, the fre-
quent appearance of a single central cell was again replaced by the two
characteristic of the depressed biseriate gland (Fig. 71). By focusing on
the surface when stomata are in view, the crater-like appearance of the
punctate condition was again shown (Fig. 70). By carefully following
around the margin of the leaf for any traces of the gland, only near the
apex where it was thicker, a lateral view of one was seen (Fig. 72) giving
evidence of at least an occasional uniseriate gland such as found in the
previous genera. Certainly when examined microscopically the large basal
leaf of Trilisa carnosa appears very different from those of its two con-
geners. However, in a similar way the biseriate glandular condition of
one Carphephorus species, C. tomentosus, distinguished it from the other
three.

No non-glandular trichomes were found on the lower leaves of the first
two species and only an occasional short one near the petiole of T. carnosa.
They were very similar to those of Carphephorus (Fig. 73).

SUMMARY (RE TRICHOMES )

With the help of table IIT, the observations on trichomes and glands on
basal cauline leaves can be summarized.
1. Non-glandular, uniseriate trichomes are lacking in Garberia and Car-
phochaete and show very little variation within three genera, being

°T am indebted to Prof. G. N. Hocking for these references.

1954] GAISER, STUDIES IN THE KUHNIINAE II 119

acuminate in Barroetea and nearly uniform in Carphephorus and Trilisa.
In Kuhnia, the largest of these genera, they may vary from being nearly
filamentous or uniform to slightly tapering or attenuate. The thicker
walled forms of the latter suggested the moniliform.

2. The elevated biseriate glandular trichome, comparable to that forming
the indumentum in a number of species of Brickellia, was found only in
one species of Barroetea.

The same structure sunken below the leaf surface, the depressed bi-
seriate glandular trichome, was common to all the other species of Bar-
roetea and to all species of KuAnia and one of Trilisa.

4. In Garberia and one species of Carphephorus the same biseriate gland
was accompanied in the depression by uniseriate but non-capitate gland-
ular trichomes while in all four species of ances the same grouping
occurred but the biseriate gland was different in

5. In the other species of Carphephorus kee ‘glandular a
were depressed either singly, in one species, or grouped, in two othe

6. The two other species of Trilisa were characterized by amen de-
pressed, uniseriate, secretory glands.

7. The small uniseriate capitate glands were observed only on species of
Kuhnia.

8. As the punctate condition results from a depressed gland or glandular
trichome it was found in all genera and all species except Trilisa odoratis-
sima and T. paniculata.

9. While in representatives of each genus, the depression contained the
biseriate glandular trichomes, in three species of Carphephorus there were
uniseriate trichomes singly or in groups. Thus the punctate condition
accompanies different glandular contents in these genera.

A very reticulate pattern is evident in the distribution of the glandular
trichomes. Though some form is found in every species, the same is not
necessarily common to all those of any genus.

IV. Pappus

In the key to his treatment of this group of genera, Hoffmann (1890)
made use of differences in the length of barbules of the pappus, besides
the singular scale-like nature of that of Carphochaete. Thus Kuhnia was
distinguished by plumose pappus from Brickellia and Barroetea. Illustra-
tions of Kuhnia eupatorioides and Brickellia californica A. Gray (Fig. 81
B & C) were drawn to scale and indicate that for these two species, the
barbules of the former are at least four times as long as the latter.

Robinson made little use of the pappus in the classification of Brickellia
because he found that as well as in barbule-length, the number of setae
varied widely from species to species. As pointed out (Gaiser 1953),
Robinson did include its plumose nature under the sectional heading of
Steviastrum and elsewhere only in the species-descriptions of B. brachy-
phylla and B. monocephala. Lactic acid mounts have confirmed a wide
variation in the length of barbules of more than six times from these two

120 JOURNAL OF THE ARNOLD ARBORETUM

SSS SST SE SSS a ee ees SS
oe 0 epee tne gO

[VOL, XXXV

FIGURES OF PAPPI

1954] GAISER, STUDIES IN THE KUHNIINAE II 121

which are the longest, to the shortest. Generally in any species the bar-
bules become a little longer away from the tip of the seta. However, their
arrangement in this genus seemed distinctive from that seen in Liatris.
The generally flattened appearance along the greater length of the seta
in many species resolved itself as a distichous arrangement of the barbules
along the axis, as shown for B. Nelsonii Robinson (Fig. H). In other
species, e.g., B. Coulteri where the barbules were only about half as long,
their arrangement along the mid-portion of the seta approached the mar-
ginal appearance of the serrulate-paleaceous type of Small (1919, fig. 17H).
However, the central axis in all the specimens seen did not broaden greatly,
showing an increase in diameter of not more than twice with an elongation
of barbules of three times or more than that amount. Brickellia diffusa
and B. filipes, the two annuals, appeared exceptional in having the bar-
bules arranged in three rows or tristichously. Always at the apices, the
barely emerging barbules came in less regular arrangement and for this
reason the illustrations include an apical as well as a median portion of a
seta. In some species having short barbules, the irregularity persisted
further along the axis.

Comparison of similar mounts of species of all of these genera here
studied was therefore included. In contrast to the very large genus Brick-
ellia, the smaller genera showed a greater uniformity in the nature of the
pappus.

In Barroetea the length of the barbules varied little in the six species
examined, being about equal to the shortest in Brickellia. Their arrange-
ment also was uniform so that the seta had the appearance of a four-angled
structure, due apparently to a tetrastichous arrangement of the barbules.
In figure I, the row of barbules in the center below were not included. No
exact equivalent for this was found in Small’s figure 17 of pappus forms in
the Compositae.

The plumose pappus of species of Kuhnia varied distinctly from Bar-
roetea not only in the length of the barbules but also in the longer spaces
between the points of their emergence from the axis and in their flat,
distichous arrangement (Fig. J). In the latter they resembled Brickellia.
In comparison with the plumose pappus of a species of Liatris (Fig. K)
the barbules are slightly more slender and appear much less abundant
because they do not extend from the mid-axis. Although the tips of emerg-
ing cells can be seen over the center region, they do not project laterally.

EXPLANATION OF FIGURES OF PAPPUS

All figures are of pappus from specimens in the Gray Herbarium and will be referred
to by the collector’s number, except Figs. M and N, Teh paniculata and T. carnosa which
were of accessions studied eicloaicaily, from Sebring, Highlands Co., Fla., Ray Garrett,
a

rickellia Nelsoniit, Nelson 4449. Fig. 1. Barroetea brevipes, Nelson 1520. Fig. J.
lee ae aC Pringle 2933. Fig. K. Liatris squarrosa var. glabrata, Rydberg 1505.
Fig. L. Liatris ligulistylis, Nelson 1651. Fig. O. Carphephorus tomentosus var. Walteri,
Fernald and Long ee Fig. P. Garberia heterophylla, Nuttall. Fig. Q. Carphochaete
Schaffneri, Schaffner, 24
The author is pre indebted to Dr. B. G. L. Swamy for these figures, drawn X 45.

122 JOURNAL OF THE ARNOLD ARBORETUM __ [VoL. xxxv

This is indicated in another illustration of Kuhnia eupatorioides (Hoff-
mann 1889, Fig. 60D) where it can also be compared with Liatris pyc-
nostachya Michx. (Fig. 60B), which has a barbellate pappus. At least
five of the seven species of Kuknia had barbules of approximately the
same length, while those of K. leptophylla var. maxima and K. adenolepis
were from half to three quarters as long. In length the barbules of this
majority are somewhat intermediate between that of Liatris squarrosa
(L.) Michx. var. glabrata (Rydb.) Gaiser, belonging to section Euliatris
with plumose pappus and L. ligulistylis (Nels.) K. Sch., of section Suprago
in which it is barbellate.

While it is difficult, when the barbules are as long and as abundant as
in the plumose pappus of L. sqguarrosa (Fig. K) to illustrate their points
of emergence even at the apex, when they are shorter as in the barbellate
type of L. ligulistylis (Fig. L), their general disposition all around the
axis is evident. This is similarly shown in the figure of L. pycnostachya
of Hoffmann (l.c.). The projecting barbules are not limited to two sides
and give no suggestion of a flattened appearance. They emerge somewhat
spirally as shown also for this type (Small fig. 17D). This same arrange-
ment was seen in the mounts of species of Trilisa (Figs. M & N), Car-
phephorus (Fig. O) and Garberia (Fig. P) all of which have barbellate
pappi with barbules shorter than those of Liatris ligulistylis though equal
to those of some other species, e.g., L. gracilis Pursh. Except for slight
variations in the length of the barbules, the similarity of the pappus from
genus to genus, as well as of the species within a genus was too great to
allow any differences of classificatory value for Trilisa, Carphephorus and
Garberia.

Of course, Carphochaete with its paleaceous pappus is quite distinctive
from all the others (Fig. Q). It is dilated and scale-like at the bottom
and for this reason the figure of this form includes a basal portion in addi-
tion to the median and apical. Carphochaete Schaffneri, the species illus-
trated, has the shortest free barbules at the apex of any of the four. The
two other Mexican species are very similar to it, while in C. Bigelovii they
are almost twice as long.

DISCUSSION

This study of six genera indicates a relationship with the two others
previously studied, Brickellia and Liatris. With the exception of Kanimia,
of which it has been impossible to include cytological examination and
therefore discussion has been omitted, there is support for the taxonomic
grouping of the subtribe Kuhniinae. Kanimia will have to be compared
cytologically with Mikania, the genus of the previous subtribe Ageratinae,
with which it may well be a connecting link — of the several technical
characters which the two genera have in comm

Furthermore, some points of relationship ant the subtribe have been

1954] GAISER, STUDIES IN THE KUHNIINAE II 123

added to those noted by earlier authors. These have served as a guide in
the preparation of a key (see p. 124) which attempts to more nearly ap-
proach a natural classification. It has already been pointed out that when
Cassini established each of the two new genera, Carphephorus and Trilisa,
he placed them next to Liatris. Also because of the strong similarity of
the flowers, Nuttall first described Garberia as a shrubby Liatris. With
the additional evidence gained from the study of seedlings, trichomes,
pappi, and chromosome numbers, it is proposed to refer to these four
genera as Group I.

Likewise there is sufficient evidence to warrant a second grouping. When
Gray (1879) established the genus Barroetea, he began the description
“TInvolucrum (15-25) flores KuAniae et Brickelliae.” Hoffmann (1890)
in his introduction to the subtribe also referred to Barroetea as approach-
ing Brickellia in all else except the nature of the fruit. Robinson, who
monographed both these genera, re-emphasized their strong similarity in
habit and involucre, giving the flattened achene and sharply toothed
leaves as the chief points of difference. To separate Kuhnia from Brickel-
lia, the same author referred to its less strongly imbricated phyllaries and
plumose pappus. For Shinners (l.c.) these technical characters were not
completely satisfactory, as was also an additional one, the nature of the
root system. His conclusion was that though separately imperfect they
were “strengthened by the obviously close relationship of the species of
Kuhnia to each other, making a closeknit and recognizably distinct
group.” The association of these three genera will be continued as

differences were noted which distinguish the two groups. Those of species
representing Group I, had a rosette of leaves which persisted without
production of a flowering stem for a longer period than would necessarily
be required for a biennial. In seedlings of Garberia, the one shrub of the
group, this stage did not last as long and then a central leafy stem began
to appear though it never flowered. However, species of Group IT, in-
cluding representatives of all three growth-forms, e.g., annual herb. per-
ennial herb, and shrub, produced elongating central axes almost at once
and these flowered before the end of a year. The seedlings of Carphochaete
Bigelovii resembled the members of this latter group.

It was felt by Shinners that in distinguishing Kuhnia undue reliance
had been placed on the plumose pappus because it might not be the ex-
clusive type of a genus, e.g., Liatris. From an examination of the pappus
of these genera by lactic acid mounts, it was found that the arrangement
of the barbules differed in these two. Further, though the barbules were
longer in Kuhnia than in species of Brickellia, their lateral arrangement
was more nearly like the distichous pattern found in the majority of the
species of that genus. In two species of Brickellia, the three-sided or
tristichous arrangement suggested a mid-way condition between that of
the others and that typical of Barroetea where it was four-sided or tetra-

124 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

A GENERIC KEY TO THE KUHNIINAE *

Anther apex having an ovate or oblong membranaceous appendage, achenes
10-20 (rarely 6-9) ribbed, phyllaries indefinite in number.

(a) Pappus squamiform, dilat ed at the base, small shrubs, leaves opposite,
young plants without rosettes, corolla rose-colored or white, chromosome
number n=11......... i Loe ata ats 6 oq eha ae Carphochaete.

(a) Pappus setose.

(b) a not conspicuously striate, leaves alternate, young plants with
osettes, corolla rose-colored, rarely white, barbules of pappus in-
anne arranged along the axis, chromosome numbern=10.......

OE eT Ee ET CT ET ee ree Seer errr ee UP I
(c) Woody shrubs, leaves alternate obovate, rosette-form not persisting
long, phyllaries herbaceous, pappus barbellate ..... Garberia.

(c) Perennial herbs, leaves linear to lanceolate, ae form persisting.
(d) Receptacle chaffy, phyllaries subequal pappus prhewe -
ET Oe ee Tee ee ee re re eT eee ee arphephorus.
(d) Receptacle naked.
(e) ie arising from thickened ea root-system, phyl-
laries 2-3 seriate se barbellate .......... Trilisa
(e) =e arising from a corm-like or pane penetrating sok
itera phyllaries cae herbaceous or colored or
ous, pappus barbellate or plumose ........ Liatris.
Phyllaries ae gern scarcely herbaceous, leaves alternate or opposite,
young plants without rosettes, corolla ochroleucous to yellow, rarely
rose-colored, barbules of peppis tend to be in linear arrangement along
the axis, chromosome numbern=9.................. GROUP II

Achenes nen or rounded.

(g) Phyllaries imbricate in several series, shrubs and
perennial, rarely annual, herbs, pappus tends to
be arranged distichously, rarely aanre -
Leudala dain ny ace ae sibs ve cele pe aa Brickellia.

(g) Phyllaries 2-3 seriate, perennial herbs from long
conical roots, pappus plumose, barbules arranged

istichously ................... uhnia.
(f) Achenes ee annual and perennial herbs,
barbules of pappus arranged tetrastichously

Side edt Artun ds & Suotahocesduh 4.6 Gok fd(s w Gate ohn ae Barroctea.

Y i,’
o
—

stichous. Thus in the pappus of the two smaller genera, in comparison
with Brickellia, the arrangement of barbules alone differentiates Barroetea,
and arrangement in addition to length, Kuhnia. While interspecifically
Brickellia shows considerable variation in length of barbules, the species
have the same basic linear arrangement which adds support to the place-
ment of these three genera in one group. Similarly the four genera of
Group I resemble one another in a less definite, somewhat spiral arrange-
ment of the short barbules as is shown in figures of setose pappus.

In addition to the characters in prevalent use by taxonomists, it has
been noticeable that in the Eupatorieae, it was necessary to pay consider-

7 Exclusive of Kanimia.

1954] GAISER, STUDIES IN THE KUHNIINAE II 125

able attention to glandular structures. Robinson had established species
as well as varieties on the presence of glands on the phyllaries, e.g., KuAnia
adenolepis, Brickellia adenocarpa var. glandulipes, etc. Likewise Brande-
gee (1908), who described two of the seven species of Barroetea, found
B. glutinosa ‘‘differing from the others of the genus” because of the gland-
ular leaves. Also, the common use in manuals, as well as descriptive
treatises, of the term punctate and puncticulate in some species, left open
the question of general application for all of each genus. As explained
under Brickellia, there was the need for interpretation of the overlapping
use of the various terms applied to the depressed glands and the punctate
condition. A more detailed comparison than had been possible by macro-
scopic examination, has helped to clarify.

From the reticulate pattern of glandular trichome distribution in six
genera, summarized in Table III, and of Brickellia (Table III, Gaiser
1953), it was seen that some form was represented in each species. Liatris
also can come in this inclusive statement. In the introduction to the treat-
ment of that genus, it was stated that small sunken resinous glands were
generally found in the leaves of species. Examination of cleared leaves of
at least several species have given confirmation of the presence of the
depressed biseriate trichome in connection with the punctate condition,
as shown for L. punctata (Figs. 74, 75). A closer analysis of these tables
gives indication of some intergeneric similarities.

In several species of Brickellia, the pubescence consisted almost entirely
of the biseriate glandular trichome. That, of all the species of the other
five genera examined, only puberulence of one species of Barroetea was
composed of this form, is another point to add to the number in which
that genus strongly resembles Brickellia. However, in the thick leaf of
Brickellia glutinosa, a similar shorter biseriate trichome was shown to
make up the depressed gland which gives a surface appearance of only
two hemispherical cells as seen in species of Kuhnia, as well as the other
species of Barroetea and Brickellia. The conclusion was reached from the
range of variation seen in the extent to which the biseriate gland rose
above the surface in different species of Brickellia, that the superficial
glandular trichome was an elevated expression of the depressed gland.
With the additional evidence in the fleshy leaves of the other genera of
the heights it may attain and still remain macroscopically subsurface,
there is no reason to doubt the generality of the statement. It is as under-
standable that the height of this small organ should vary in leaves of
different species as that non-glandular trichomes should vary in length.
More important than the variation from a depressed to an elevated con-
dition may be the uniformity of this biseriate glandular organ. While these
three genera of Group ITI have in common this single, similar content in

seriate, or single uniseriate without correlation to the punctate condition.
Examining this apparent diversity, there is a link with the previous
three genera in Garberia and one species of Carphephorus having this

126 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxv

same biseriate form in the grouping along with the uniseriate. To these,
the other Carphephorus are tied in having only uniseriate trichomes and
Carphochaete, in having uniseriate with a different form of biseriate tri-
chomes. Likewise in the two species of Tvilisa the uniseriate trichome,
though not accompanied by leaf punctation, appears similar in form to the
depressed of Carphephrous bellidifolius, while Trilisa carnosa has the de-
pressed biseriate resembling those of Group II. Thus while there is greater
uniformity in the three genera of Group II, those of the other group seem
to have a similitude in intra-generic variations. The particular form of the
biseriate depressed gland of Carphochaete is individual among them.
While its difference sets it apart from the genera of both groups, the true
significance of this variation may be better understood when further
knowledge is obtained of the forms in the many genera of other subtribes.

Further emphasis on relationships by glandular structures in these
genera was found in the fact that there was a smaller, uniseriate, capitate
trichome, which would not be readily observed without a microscope, along
the veins and under surface of leaves of species of Brickellia. The only
other genus of the subtribe studied to which it has been found common is
Kuhnia. A slightly enlarged form apparently became the specialized type
of K. adenolepis.

From a consideration of the non-glandular trichomes * the largest genus
has the widest variety. Comparable preparations of other genera con-
tributed no distinctly new type, though in Carphephorus, Trilisa, and one
species of Kuhnia, they were more nearly uniform in diameter and tapered
a little less than the attenuate form shown for Brickellia. That all the
types seen in that genus were not represented in the smaller genera could
be explained probably by the smaller total of species. However, in Brickel-
lia considerable variation was found in the size of chromosomes and in
some subsections a correlation with the types of trichomes

Cytological evidence has been obtained of some species of each of the
genera of the Kuhniinae except Kanimia. Though incomplete, it supports
the conclusions obtained from other sources. The basic number of chromo-
somes is 10 in the four genera of Group I, 9 in the three genera of Group IT
and 11 in Carphochaete. While no other genus of the subtribe has the
same number as Carphochaete, the fact that the three basic numbers vary
only by one, may be taken as further indication of a fairly close relation-
ship of the members of this subtribe. This seems more significant when the
number of species examined for the several genera to which the same num-
ber is common, totals forty-seven with n = 9, and thirty-nine with n = 10.
Of course these two chromosome numbers are represented many times in
genera of the Compositae as can be seen in Darlington and Ammal (1945,
p. 220) where the basic numbers of the tribes of the Compositae are given.
When turning to the reports for the tribe Eupatorieae, of which there have
been few so far, the number 11 does not appear. The recent addition by

® Gaiser (1946) in haha to pubescence as being of little use for species differen-
easy in rae ris, had not made microscopic analyses and there has not been time

to do so fax hae sie

1954] GAISER, STUDIES IN THE KUHNIINAE II 127

Grant (1953) of numbers for at least thirty-two species of Eupatorium
from the temperate zone, give 10 and 17 as the lowest numbers, and 9 for
one tropical species of Vernonia. The number 11 does not appear. This
does present a challenge to investigate Kanimia. Also it raises the question
of possibly finding the same number in a genus of another subtribe which
perhaps may show further intertribal relationships with Carphochaete.

Of two species of Mikania, the genus of the previous subtribe A geratinae,
which has characters in common with Kanimia, it was possible to obtain
some seeds (see Table I). The somatic chromosome number of three ac-
cessions of Mikania scandens Willd. and one of M. cordifolia (L.) Willd.
was found to be 2n = 38 (Fig. 21), which is quite different from that in
any of the Kuhniinae investigated. Thus it seems quite probable that
cytology might give the deciding evidence for or against the segregation
of these two genera within the same subtribe.

No lower basic number was found among these genera than had been
found in Brickellia and Liatris, or had been known previously among the
Eupatorieae. These numbers, 9 and 10, are higher than have been found
in a number of the other tribes of the Compositae, e.g., Chichorieae with
3, Heliantheae and Astereae, with 4, /nuleae and Senecioneae with 5, etc.,
but this may find its explanation in the few genera studied. Of the
KuAniinae, the genera of Group II with Mexico as their geographic center
of distribution, have the lower number 9, while those of Group I, which
are to the north of the Mexican boundary have 10. At this time there is
no evidence of the origin of these divergent genetic lines. So far, the most
northerly genus, Liatris, and in that only the Punctatae series, remains
the only one in which polyploidy has been found. The tetraploid, L. punc-
tata, reaches the most northerly latitude of any species of that or any
other genus of the subtribe and so exemplifies Hagerup’s theory. Since
the same species reaches the southern extremity of the range for the genus
along the mountains and the diploid is found on the plains in between it
also provides an example of the polyploid occupying a wider area than the
diploid. Miintzing (1936) believes that the extensive range is the com-
bined result of polyploidy and polymorphism which has enabled adapta-
tion to a wider range of habitats some of which may be unfavorable. Of
all the other genera of the Kukniinae few include any species of verv
extensive range and none which is exactly comparable to L. punctata.
Kuhnia eupatroioides is the only polymorphic species of that genus, occur-

is
introduction, Robinson referred to Brickellia grandiflora and B. californica
as the two most widely distributed and variable species. Both were found
to be diploid in these studies which included a generous representation of
thirteen accessions of the latter species. The former species like two others
studied, B. microphylla and B. oblongifolia reach the northern limit for
that genus in Washington State, but they also were diploid.

Excepting the species of Brickellia in Brazil, the most southerly one
from Costa Rica, B. argyrolepis, was also a diploid. For the discussion

128 JOURNAL OF THE ARNOLD ARBORETUM _ [vor. xxxv

of the polyploid complex and tropical plants, as pointed out in the intro-
duction, this genus is somewhat exceptional since though within the trop-
ical latitudes the species live on higher altitudes. Of Barroetea, which at
similar altitudes does not extend north of Mexico, neither of two species
were polyploid. Also one speices of Kudnia from the southerly limits for
that genus was diploid like the three congeners from the United States.
The conclusion from this discussion still remains that for these closely
related genera, as they have been represented from Central America north-
ward, polyploidy was not prevalent and occurred only in the most northern
genus of Group I. However, variation in the size of the chromosomes was
noticeable and the karyotypes of the shorter units occurred only in the
genera of the more southern group.

This does not mean that we should not expect polyploidy in genera of
the tropics. For comparison, in the tropical genus Anthurium, of which
thirty-seven of four hundred and eighty-six species (Engler 1905) were
examined from plants as they grew mostly in the New York Botanical
Garden, the situation is quite different. While many additional species
have also been added since Engler’s monograph of the genus, there is
indication that polyploidy is not singular and occurs in species of both
limited and wide distribution (Gaiser 1927). With 30 as the lowest so-
matic number found, at least two other numbers of a polyploid series
were represented, two species with ca. 50 and three with ca. 60. The five
polyploids were distributed to as many different sections, some nearly
monotopic and others, the largest of the genus, e.g., Urospadix with ninety-
five species. Anthurium radicans which belongs to the section, Chamae-
repium, comprising two species, is limited to east Brazil, and A. Wallisii
of section Polyneurium of thirty species had been reported as having been
collected only once in Colombia. In comparison with tetraploid A. cras-
sinervium, from Venezuela, Colombia, Panama, and Tobago Island, nine
other diploid species of the second largest section Pachyneurium with
fifty species were found to be fairly restricted. Two species of section
Urospadix, very similar to the polyploid A. digitatum from Venezuela and
Tobago Island, had been collected from comparable areas. However, the
fifth, 4. scandens, is the most variable species in the genus, including five
varieties and is known in one of its forms wherever Anthuriums grow.
With even but fractional representation studied cytologically, all the
complexities of polyploidy in percentage of incidence, different multiples
of numbers and correlation of geographic distribution occur in that genus.
Also, since Grant (l.c.) found fifteen polyploids (almost fifty percent)
among the northern Eupatoriums examined, others might be expected
among the many more tropical species of that large genus.

While the studies of Liatris were of a different time and were not all
accompanied by exactly comparable photographic figures, chromosomes
of the series Spicatae (Gaiser 1949) were considered to be the longest
and to show greater variation than those of the series Graminifoliae (ibid.
1950a) and Punctatae (ibid. 1950b). Reference to the analysis of the
morphology of the chromosomes of L. pycnostachva (1949 p. 127) gave

1954] GAISER, STUDIES IN THE KUHNIINAE II 129

two pairs of long chromosomes in the karyotypes of 2 Lm, 2 Lst, 4 Mm,
4 Msm, 2 Mst, 4 Sm, 2 Sst. In the more recent comparisons the few species
of the smaller genera of Group I varied little if at all from species to
species. Also, the generally similar karyotypes of Trilisa and Carphe-
phorus, including three pairs of long chromosomes, were found to vary
somewhat from Garberia only in the types of the short and medium
chromosomes. By comparison, Carphochaete Bigelovii was the only spe-
cies to have a greater number of long chromosomes, four pairs rather
than three.

Brickellia with the greatest number of species, had a great variation in
chromosome sizes. Of thirty-four species in which the karyotypes were
carefully analyzed, nine had only short and medium chromosomes, while
in all the others there were also long ones. From the analyses of species
of all these genera, only those of KuAnia and Barroetea had such a karyo-
type of the two classes of shorter units. Thus not only have these three
genera the same number in common but like a minority of the species of
Brickellia, the species studied of the two smaller genera, have chromosomes
representing an amount of chromatin less than is found in any of the others.

While too few species of Barroetea have been examined to draw conclu-
sions regarding that genus, more than half of those of KuAnia, including
all but one variety of the polymorphic species K. eupatorioides have been
included. The great homogeneity in the karyotype as well as chromosome
number for all of these, shows a genetic basis for the close relationship of
species of KuAnia to each other, pointed out by Shinners. By contrast the
great diversity in chromosome length found in species of Brickellia accom-
panies a diversity of growth from woody shrubs to herbaceous annuals.
None of the genera of Group I can compare with this. In Liatris, the
largest of them, all the species are herbaceous perennials of a fairly similar
type.

In the discussion of Brickellia, it was pointed out that a minority of the
species examined, nine out of thirty-four, had a complement of shorter
chromosomes. The species at the opposite extreme having the longest
chromosomes, B. monocephala and B. grandiflora, were considered by
reason of their modified underground structure, etc., as belonging to a spe-
cialized group of that genus. Others next to them, also having a greater
total of chromatin material by reason of longer chromosomes than in the
major group, included shrubs and herbs native to Costa Rica, Guatemala
and Mexico. With the most woody species from Costa Rica among these,
and the one annual examined, in the first group, there was some ground
for considering that evolution had gone on in conjunction with reduction
in chromatin in that genus. It is of interest therefore that the only other
genus including annuals, Barroetea, also had short chromosomes in the
two species examined. The genus includes no true shrubs and only one
perennial variety of a species that has been described as being somewhat
woody at the base. Kuhnia, too, is a genus consisting entirely of perennials
and in it there was found a similar karyotype of the two shorter classes of
chromosomes. While it is impossible to trace the steps of evolution from

130 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

one genus to another with certainty, it is indicated from the present evi-
dence that the direction at least, was from Brickellia to the two other
genera through some form, not necessarily extant today, having a com-
plement of shorter chromosomes. Whether this happened in two successive
steps or at one time remains a question also, The fact that both genera still
include few and, for the most part, less variable species indicates that they
are more recent than Brickellia. The repeated reference to the close simi-
larity of Barroetea and Brickellia, especially as seen in both including
annual species, suggests that there are possibly blocks of genes common
to these two and this could have been one mutation. Also the indecision
regarding basic differences between Kuhnia and Brickellia gives weight to
the probability that this might have come as another step. The similarities
of each of the two smaller genera to Brickellia appear closer than the
interdependence of all three. Yet, as has been pointed out, from karyo-
logical studies, the six species of Kuhnia and two of Barroetea had karvo-
types consistently represented by a pair of satellite chromosomes. Only
in three, of about half of the species of Brickellia examined, was this tvpe
of chromosome regularly visible. Two of these were xerophytes, B. incana
and B. Greenei, very restricted in distribution and remarkable for their
large heads and seeds which are distinctive. The third was B. Coulteri,
from Baja California, the only species examined of subsection Brachiatae.
Most of the other seven species have been reported but once and from
widely separated regions in Mexico and certainly should be compared cyto-
logically. Without complete representation of a genus, the species not
studied may be the ones which withhold the most significant truths. Per-
haps the best that can be expected is to gain at least a point of vantage
from which to envision the horizons for further efforts in a project.

It is difficult to extrapolate true phylogenetic relationships within Group
I. In all four genera the karyotype is made up of more closely approxi-
mating units than were seen within the one genus Brickellia. Some varia-
tion has been reported in the species of five series of the genus Liatris °
but is not as striking as has been found in Brickellia. There was no evi-
dence of marked morphological changes or reduction in size of the chromo-
somes. Also there is a general similarity of karyotype in the two smaller
genera Trilisa and Carphephorus, including three pairs of long chromo-
somes, From these, Garberia was found to vary only in the medium and
short chromosomes. Likewise, except for this one shrub Garberia, there
is a greater homogeneity in the growth-form represented by each of the
three genera. They are herbaceous perennials, provided with excellent
modified storage rootstocks, mostly corm-like in Liatris and more tuber-
ous in Trilisa and Carphephorus. From the discussion pertaining to
growth-forms in Brickellia, just such as these were considered to repre-
sent a form of specialization and, these may very well be from a more
primitive type now extinct. In the one living species of Garberia may lie

*A chromosome list will shortly be forthcoming for species of the other series
except the Squarrosae, which Dr. Pauline Snure will contribute.

1954] GAISER, STUDIES IN THE KUHNIINAE II 131

the hint of another more woody progenitor for the other three genera.

The dissimilarity of the karyoytpe of Carphochaete Bigelovii to the
species of Group I lay in the fact that it was the only species to have two
additional chromosomes, four long chromosomes instead of three and three
short ones rather than two. These factors contributed to its having the
greatest content of chromatin in any of the genera studied. Comparison
of it with the unusually long chromosomes of Brickellia monocephala (see
Fig. 30, Gaiser 1953) is favorable except for the extra pair of chromo-
somes. The singularity of its karyotype, supported by the distinct form
of the biseriate depressed gland and the individuality of its pappus, is
sufficient to set it apart from the genera of both Groups I and II. Yet the
prime reason for its inclusion in the subtribe Kuhniinae, the similarity of
its anther and achenes, also gains weight in a proximal chromosome number
and karyotype to that of Group I and in the actual presence of a depressed
biseriate gland.

Without examining other genera having paleaceous pappi for comparison
with Carphochaete, it would be premature to generalize on the evolution
of pappus forms. That this small group of related genera may contribute
to the overall study is indicated by the variety shown in the barbule-ar-
rangement especially in the three genera of Group II. In Barroetea and
Kuhnia there is near uniformity for the species of each genus, while in
the more numerous species of Brickellia there is a variety of almost im-
perceptible changes. The general tendency is for a flattened, lateral ar-
rangement of barbules in two rows. While the significance of their quad-
rate. linear arrangement in Barroetea is not clear, it may be but one of
nature’s experiments and so also the tristichous disposition in Brickellia
diffusa and B. filipes may be a further shuffling of genes. However, since
the latter species are annuals and since Barroetea is the only other genus
including annual species, it would be in agreement with phylogenetic evi-
dence to assume that the change in Brickellia was in the direction away
from the flat or distichous arrangement. And if so, the latter referred to
as bearing a marginal resemblance to the serrulate-paleaceous, could have
been derived from the paleaceous or more foliar by reduction. This would
be in agreement with the conception of Babcock and Stebbins (1937) for
genera of the Cichorieae rather than the alternative hypothesis of Small
(1916), that the paleaceous types result from the fusion of the simple
scabroid setose which constitute the primitive type.

BIBLIOGRAPHY

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BLAKE, S. F. 1942. Compositae. In Flowering Plants and Ferns of Arizona by
T. H. Kearney and R. H. Peebles. U. S. Dept. Agr. Misc. Bull. 423: 869-
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DariincTon, C. D. and JANAK1 AMMAL, E. K. 1945. ee Atlas of
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53-295,

Gatser, L. O. 1927. Chromosome Numbers and Species Characters in An-
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1946. The genus Liatris. Rhod. 48: 165-183, 216-263, 273-326, 331-382,

393-412.
1949. Chromosome Studies in Liatris. I. Spicatae and Pycnostachyae.
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1950a. Chromosome Studies in Liatris. II. Graminifoliae and Pauci-
florae. Amer. Jour. Bot. 37: 414-423.
1950b. sa Studies in Liatris. III. Punctatae. Amer. Jour.
Bot. 37: 763-77
1951. A ae Hybrid of Liatris (< L. Weaveri Shinners). Amer.
Midl. Nat. 45: 750-763.
1953. Chromosome Studies in Kuhniinae (Eupatorieae) I. Brickellia.
Rhod. 55: 253-267, 269-288, 297- 321, 328-345.
Grant, W. F. 1953. A Cytotaxonomic Study i in the Genus Eupatorium. Amer.
Jour. Bot. 40: 729-742.
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134 JOURNAL OF THE ARNOLD ARBORETUM _ [vot. xxxv

MISCELLANEOUS MALAYSIAN NOTES
E. D. MERRILL
With one plate

MORACEAE

Ficus porteana Regel, Gartenfl. 11: 280. ¢. 372. 1862; Gagnep. in Le-
comte, Fl. Gén. Indo-Chine 5: 774. 1928.

Ficus Hacienda Warb. in Perk. Fragm. Fl. Philipp. 196. 1905; aie Leafl.
Phili 1: 192. 1906; Merr. Enum. Philipp. Fl. Pl. 2: 56. 923.

Ficus pare ty Elm. Leafl. Philipp. Bot. 4: 1250. 1911

My attention was called to Ficus porteana Regel by Gagnepain’s refer-
ence of certain Indo-Chinese material to it and by his reduction of Ficus
malunuensis Warb. to it as a synonym. Regel’s species, entirely overlooked
by me when engaged in the preparation of the Philippine enumeration, was
based on specimens cultivated in Moscow, grown from seeds secured by
Mr. Porte in the Philippines (Luzon) in 1861. Regel’s description is based
on sterile material and from his figure, a somewhat juvenile form, for the
large leaves are shown as having a fairly large lateral lobe on each side,
a character that is lost in mature specimens. There is not the slightest
doubt as to the identity of Warburg’s species, also based on Luzon mate-
rial, with the form characterized forty-three years earlier by Regel. There
is some doubt in my mind as to whether or not the Indo-Chinese specimens
referred here by Gagnepain actually represent the same species as the
Philippine form. Its general alliance seems to be with Fictus callosa Willd.

Ficus pyriroia Burm. f, Fl. Ind. 226, 1768 = Pyrus pyrifolia (Burm.
f.) Nakai, Bot. Mag. Tokyo 40: 564. 1926; Rehd. Man. Cult Trees
Shrubs ed. 2, 404. 1940

This reduction was made on the basis of an examination of Burman’s
specimens in the Delessert herbarium at Geneva; there are three sheets,
all sterile, from the vicinity of Nagasaki, collected by Kleinhof; these,
according to Nakai, represent the ordinary Japanese sand pear, which by
various authors has been placed under P. sinensis auctt., P. serotina Rehd.,
and P. montana Nakai. Nakai attempted to define a number of minor
forms of this cultigen, but no botanist has, as yet, clarified its relationships
with the Chinese forms of the sand pear. In my treatment of Burman’s
species, Philipp. Jour. Sci. 19: 346. 1921, working solely from Burman’s
descriptions, I suggested that Ficus pyrifolia Burm. f. might prove to be
the same as Ficus erecta Thunb., the type of which was from Japan, but
this is an error. In recent years the Burman name has been attached to
some Chinese collections of Ficus erecta Thunb.

1954] MERRILL, MISCELLANEOUS MALAYSIAN NOTES =S5

URTICACEAE

Laportea peltata Gaudich. Freyc. Voy. Bot. 498. 1830, nomen nudum, et
ex Decaisne, Herb. Timor. Descr. 162. 1835, descr.; Wedd. Arch.
Mus. Hist. Nat. 9: 126. 1856 (Monog. Urt. 126), et in DC. Prodr.
16(1): 80. 1869, cum syn.; J. J. Sm. ex Koord. & Val. Meded. Dep.
Landb. 10: 678. 1910 (Bijdr. Boomsoort. Java 12: 678); Koord. &
Val. Atlas Baumart. Java 4: fig. 796. 1918.

Urtica peltata Blume, Bidjr. 496. 1825.

MINDANAO: Cotabato, Nutol, Bur. Sci. 84935, 84941, Ramos & Edano,
March, 1932, a tree 8 m. high in rather dry forests at low altitudes. Java, Timor,
and probably in other parts of the Malay Archipelago; new to the Philippines.

Both of the above cited specimens agree very closely with authentically
named Javan specimens collected by Koorders, one having been distrib-
uted as representing the very different Laportea crassifolia C. B. Rob.,
and the other as L. mindanaensis Warb. It seems to be apparent that
Gaudichaud did not know of the published description of Urtica peltata
Blume, Bijdr. 496. 1825, for his original publication of the accepted bino-
mial was simply a nomen nudum and a new name for Urtica atrox Leschen.,
also a nomen nudum. Therefore the authority should not be cited as
“(Blume) Gaudichaud,” but merely as Gaudichaud. The first description
of the species under Laportea was that prepared by Decaisne, and he also
cited only Urtica atrox Lesch. as a synonym, Weddell in 1856 first, and
apparently correctly, associated Blume’s earlier name with that so casually
published by Gaudichaud and later validated by Decaisne.

Laportea elliptica sp. nov.
Laportea peltata sensu Merr. Univ. Calif. Publ. Bot. 15: 50. 1929, non Gaudich.

Species L. peltatae Gaudich. affinis, differt foliis majoribus, ellipticis,
haud molliter pubescentibus sed subscaberulis et secus costam nervosque
pilos urentis numerosos ferentibus, nervis primariis utrinque circiter 15.
Arbor, trunco 30-45 cm. diametro, ramulis ultimis plus minusve incras-
satis, siccis circiter 8 mm. diametro, pubescentibus, cicatricibus ad 8 mm.
diametro notatis, partibus junioribus dense subadpresse hirsutis; foliis
longe (8-13 cm.) petiolatis, perspicue peltatis, basi late rotundatis,
ellipticis vel subobovato-ellipticis, chartaceis, 15-30 cm. longis, 11-15 cm.
latis, apice breviter acuminatis, margine breviter dentatis vel denticulatis,
dentibus inter se 1.5-3 mm. distantibus, siccis olivaceis, supra minute
albido-verruculosis, secus costam pubescentibus et pilos urentis ferentibus,
subtus scabridulis, paullo pallidioribus et nervis reticulisque exceptis
glabris vel subglabris; nervis primariis utrinque circiter 15, perspicuis,
subtus elevatis, curvato-patulis, secus marginem arcuato-anastomosantibus,
reticulis distinctis, elevatis et cum costa nervisque plus minusve pubescenti-
bus et pilos brevis numerosos urentis ferentibus; inflorescentiis longis, laxis,
paniculatis, usque ad 45 cm. longis, plus minusve pubescentibus et pilis

136 JOURNAL OF THE ARNOLD ARBORETUM _ [vot. xxxv

numerosis urentibus instructis, ramis primariis 4-8 cm. longis; floribus
@ sessilibus, in ramulis ultimis brevibus flabellatim dispositis, sessilibus,
capitulis 5—9-floris, calycis segmentis acuminatis, vix 0.5 mm. longis;
acheniis compressis, glabris, subovatis, 2 mm. longis, acuminatis, stylis
persistentibus gracilibus, ad 3 mm. longis, leviter patule hirsutis; floribus
é numerosis, calycis segmentis subellipticis, 1.5-2 mm. longis, filamentis
3 mm. longis.

BRITISH NORTH BORNEO: Tawao, Elmer 21472 (2), 11433 (4), Octo-
ber 1922 to March 1923.

When the Elmer collections were being studied at the University of
California, these specimens were referred, on the basis of published descrip-
tions only, to Laportea peltata Gaudich, and were reported as such: but it
was then noted that they differed from Gaudichaud’s species in certain
striking characters. Now that it is possible to make direct comparisons
with authentically named Javan specimens of the Koorders collections, it
becomes evident that there is really little in common between this Bornean
form and Laportea peltata Gaudich., except in the conspicuously peltate
leaves of both. The indumentum of the two species, as well as the leaf
shape and size, and the more numerous lateral nerves in the present species
are very different. In Gaudichaud’s species the lower surface of the leaves
is very densely and softly cinereous-pubescent, the indumentum entirely
covering the parenchyma; in Laportea elliptica the midrib, and to a limited
degree the nerves, are pubescent, but these and the reticulations bear many
short, stiff, stinging hairs; yet the parenchyma within the ultimate reticula-
tions is glabrous. In the herbarium of the Arnold Arboretum the pistillate
type was found under no, 21172, Madhuca elmeri Merr., apparently some
error having been made by Mr. Elmer in completing the labels, which
makes me suspect that the same situation may exist elsewhere. The correct
number is 21472, for 21172 is the Madhuca and 21472 the Laportea.

LEGUMINOSAE
Crudia cauliflora sp. nov.

Arbor parva, foliis 1-foliolatis et ramulis glabris, fructibus caulinis,
breviter denseque pubescentibus; ramis ramulisque teretibus, ultimis 1
mm, diametro; foliolis subcoriaceis, oblongo-ellipticis, 15-18 cm. longis,
6-7 cm. latis, basi late rotundatis, apice distincte sed obtuse acuminatis,
siccis subtus brunneis, supra subolivaceis, subopacis; nervis primariis
utrinque circiter 7, subtus paullo elevatis, laxis, patulis vel subpatulis,
inter se 1.5—2.5 cm. distantibus, 1-1.5 cm. a margine arcuato-anastomo-
santibus, reticulis primariis laxis, ultimis subconfertis; petiolo cum peti-
olulo circiter 1 cm. longo; racemis caulinis, solitariis, vetustioribus glabris
vel partibus ultimis breviter obscure pubescentibus, saltem 10 cm, longis
(floribus ignotis) ; fructibus solitariis, compressis, oblongo-ellipticis, leviter
inaequilateralibus, 8 cm. longis, 4 cm. latis, extus pallide brunneis et dense
breviterque pubescentibus, seminibus 2 vel 3.

1954 | MERRILL, MISCELLANEOUS MALAYSIAN NOTES 137

PHILIPPINE ISLANDS: Siargao: Ramos & Pascasio, Bur. Sci. 34921,
June 1919, the only known collection, TypE in the herbarium of the Arnold Arbo-
retum.

This was designated many years ago by me as new in the genus Sindora,
but was never described, probably because I later realized that it could
not represent any Sindora, and I had hoped that flowering specimens would
eventually be received. It resembles several Malaysian species, such as
Crudia bantamensis (Hassk.) Benth., C. beccarii Ridl., C. curtisii Prain,
C. reticulata Merr., and even C. subsimplicifolia Merr. From all of these
it is distinguished by its simple leaves being broadly rounded at the base;
and from all described species of the genus it is distinguished by its solitary
cauline racemes. None of De Wit’s descriptions of the 18 Malayan species
which he recognized in 1950 seems to apply to this species.*

Pithecellobium splendens (Miq.) Prain, Jour. As. Soc. Bengal 66(2):
516. 1897 (Novic. Ind. 350. 1905), Pithecolobium in nota.

Albizzia splendens Mia. FI. Ind. Bat. Suppl. 280. 1861.

Pithecolobium confertum Benth. Trans. Lin. Soc. 30: 577. 1875; Bak. in Hook.
f. Fl. Brit. Ind. 2: 204. 1878; Prain in King, Jour. As. Soc. Bengal 66(2):
264. 1897 [Mater. Fl. Malay. Penin. 3 (no. 9): 264], op. cit. 508; Rid.
Fl. Malay Penin. 1: 661. 1922.

Malay Peninsula, Sumatra.

Prain’s new binomial does not appear in Index Kewensis nor in any of
its supplements to date, yet the publication of it is valid. From his own
statement it is clear that he did not intend to replace Bentham’s binomial
by the one based on the earlier Albizzia splendens Migq., as he stated that it
did not appear to him to be necessary or just, to rename Bentham’s species
because Miquel’s description was based on leaf specimens only.

RUTACEAE

ai macrantha Merr. Univ. Calif. Publ. Bot. 15: 114, 1929
(April 10).

Glycosmis oliveri Stapf ex Ridl. Kew Bull. 1930: 80. 1930, syn. nov.

The types involved are Beccari 2595 from Sarawak, for Stapf’s species,
and Elmer 12134, 21456, 21528 from British North Borneo, on which the
earlier published G. macrantha Merr. was based. The only reason for
publishing this note is the fact that Tanaka has added annotations to some
of the Elmer numbers accepting Stapf’s specific name. It is unfortunate
that Stapf did not publish his new species at the time he studied Beccari
2595, for in publishing its first description, Ridley’s work was antedated
nearly a year by the publication of G. macrantha Merr. The two species
are identical.

*De Wit, H. C. D. The genus Crudia Schreb. (Leguminosae) in the Malay
Archipelago south of the Philippines. Bull. Jard. Bot. Buitenz. III. 18: 407-434.
fig. 1-3.

138 JOURNAL OF THE ARNOLD ARBORETUM _ [vot. xxxv

MELIACEAE
Aglaia heterobotrys sp. nov. § Euaglaia.

Species A. shawianae Merr. affinis, differt foliis paullo minoribus, flor-
ibus breviter sed distincte pedicellatis, sepalis oblongis. Frutex vel arbor
parva, foliis stricte 1-foliolatis, ramulis teretibus, pallidis, glabris, novellis
circiter 2 mm. diametro, decidue breviter stellato-pubescentibus, indu-
mento ferrugineo; foliis oblongo-ellipticis, chartaceis, 14-20 cm. longis,
4-7.5 cm. latis, basi late acutis vel subrotundatis, apice tenuiter acumi-
natis, acuminibus 1.5—2.5 cm. longis, obtusis, utrinque glabris, siccis pallide
olivaceis, subnitidis; nervis primariis utrinque circiter 20, utrinque dis-
tinctis, elevatis, ad marginem arcuato-anastomosantibus; petiolo cum
petiolulo 2—2.5 cm. longo, glabro; inflorescentiis in axillis superioribus,
breviter stellato-pubescentibus, indumento ferrugineo, inferioribus sim-
plicibus, ad 16 cm. longis, elongatis, floribus in partibus superioribus
racemose dispositis, superioribus paniculatis, ad 14 cm. longis, peduncu-
latis, ramis primariis inferioribus 2.5 cm. longis, superioribus gradatim
brevioribus, floribus racemose dispositis, pedicellis 0.5-1 mm. longis,
bracteolis anguste lanceolatis, stellato-pubescentibus, ad 1 mm. longis;
sepalis liberis vel subliberis, ad 1 mm. longis, oblongis vel anguste oblongis,
obtusis vel subacutis, stellato-pubescentibus, 1 mm. longis; petalis 5,
glabris, liberis, oblongo-obovatis vel late oblanceolatis, obtusis, 2 mm.
longis, 0.6-1 mm. latis; tubo glabro, libero, 1.8 mm. longo, antheris 5,
0.4 mm. longis, inclusis, ovario dense pubescenti.

SUMATRA: East Coast, Kota Pinang District, Si Mandi Angin on the Soengei
Kanan, topographic sheet 41, southeast corner, Rahmat Si Toroes 4197, April—
May, 1933, with the local name kajoe piran.

The alliance of this species is clearly with that small group of simple-
leaved species of which the Bornean Aglaia triandra Ridl., A. odoardoi
Merr., A. matthewsii Merr., and A. shawiana Merr., as well as the Siamese
A. meliosmoides Craib, are typical. It is distinguished by its vegetative
and other characters, and particularly by its always racemosely arranged,
shortly but distinctly pedicelled flowers. The inflorescences in the lower
leaf axils are greatly elongated simple racemes, the flowers borne only
along the upper 2 to 3 cm., but the uppermost inflorescence is a normal
panicle.

Walsura monophylla Elm. Leafl. Philipp. Bot. 9: 3391. 1937, descr.
angl.

Arbor 5.5 m. alta, inflorescentiis obscure pubescentibus exceptis glabra,
foliis omnibus 1-foliolatis. Ramulis teretibus, glabris, lenticellatis, ultimis
circiter 2 mm. diametro; foliolis oblongo-ellipticis, coriaceis, 11-28 cm.
longis, 4-8 cm. latis, subtus pallidis, brevissime et acute acuminatis vel
acutis, basi plerumque obtusis, nervis primariis utrinque 10-15, subpatulis,
curvatis, arcuato-anastomosantibus, subtus distinctioribus; petiolo 1-2 cm.

1954] MERRILL, MISCELLANEOUS MALAYSIAN NOTES 139

longo; inflorescentiis terminalibus, anguste paniculatis, circiter 7 cm.
longis, obscure pubescentibus, ramis primariis 1-1.5 cm. longis, patulis;
floribus inter majores, flavido-albidis, circiter 4 mm. longis, 5-meris;
sepalis triangulari-ovatis, acutis, obscure pubescentibus; petalis oblongis,
obtusis vel subacutis, glabris, 4 mm. longis, 2 mm. latis; filamentis pubes-
centibus, tubo deorsum utrinque glabro; fructibus junioribus 5-8 mm.
longis, breviter ellipsoideis vel subovoideis, pubescentibus.

A second Philippine collection of this species is Ebalo 556 from Mount
Langogan, near Puerto Princesa, Palawan, Feb. 23, 1940; the type is
Elmer 12903 from Brooks Point, Palawan. The above Latin description
has been prepared to validate Elmer’s binomial. The striking character
of the species is its strictly 1-foliolate leaves, all other known species of
the genus havirig pinnate leaves. It has long been known that Aglaia,
another genus of the family, does contain a fair number of species with
strictly simple leaves, although in the vast majority of its species the leaves
are pinnate, and in all the known species of Vavaea the leaves are strictly
simple. This reduction of pinnate leaves to simple ones in certain species
of Aglaia is now paralleled in Walsura.

EUPHORBIACEAE
Aporosa cardiosperma (Gaertn.) comb. nov.

Croton cardiospermum Gaertn. Fruct. . oe ie 107. fig. [11]. 1791.

Agyneia latifolia Moon, Cat. Pl. Ceyl. 6

Aporosa latifolia (Moon) Thwaites, ails Pl, Ceyl. 288. 1864; Trimen,

Hand-book Fl. Ceyl. 4: 39. 1898; Pax & Hoffm. Pflanzenr. 81 (IV. 147
XV): 96. 1922

A species known only from Ceylon. The identity of Gaertner’s species
with that later described as A. latifolia (Moon) Thwaites was determined
by Hallier f., Rec. Trav. Bot. Néerl. 15: 35. 1918. According to Thwaites
the native name kebella, cited by Gaertner, belongs with the distinctly
different Aporosa lindleyana Baill., but the characters as described by
Gaertner and the details of his figure are those of the Thwaites, not of the
Baillon species.

At the end of volume two of his De fructibus et seminibus plantarum,
Gaertner assembled under a center heading Barbarae nine alphabetically
arranged taxa which he described and figured under their native names
(Ceylon and Java), Fruct. 2: 485-488. pl. 180. 1791. Hallier f., op. cit.,
gave some attention to the identity of these, although the descriptions were
manifestly not intended to represent new genera, nor are any binomials
used. They are merely casual names for those fruits that Gaertner had
which he could not refer to any described genus. A very few of these names
have appeared in taxonomic literature as if genera were intended, and in
two cases binomials are involved, although the latter were not originated
by Gaertner. Hence, the identification of these old Gaertner taxa in terms
of the binomial system is of only slight academic interest, as they scarcely
bear on problems of nomenclature. The entries are as follows:

140 JOURNAL OF THE ARNOLD ARBORETUM _ [vot. xxxv

“Balangue. E. madagasc.”
This is the whole basis of Balangue gaertneri DC. Prodr. 8: 316. 1844.
It is possibly some rhamnaceous plant.

“Cucumeroides. Ex Japonia.” = Trichosanthes.

“Edokke zeylonens.” = Chaetocarpus castanocarpus Thwaites.

“Giek zeylonens.” = Odina wodier Roxb. = Lannea coromandelica
(Houtt.) Merrill, Jour. Arnold Arb. 19: 353. 1938, cum syn.

““Zoon zelonens.” = Schleichera oleosa (Lour.) Oken, Allgem. Naturgesch.
3(2): 1341. 1841; Merr. Interpret. Herb. Amb. 337. 1917, Jour.
Arnold Arb. 31: 284. 1950. (Schleichera trijuga Willd.).

This in earlier years, was referred to the menispermaceous Pachygone
ovata (Poir.) Miers, and appears as a synonym of that species in the
latest monographic treatment of the family, Diels, Pflanzenr. 46(IV. 94):
343. 1910, where it does not belong. The binomial K. zeylanicus is cur-
rently credited to Gaertner, but he did not originate it. The earliest
reference to it that I have located is in the synonymy of Miers’ species,
Hooker f. & Thomson in Hook. f., Fl. Ind. 1: 105. 1872.

“More zeylonens.” = Euphorbia longana Lam.
“Pite-heddija javan.” = ?
“Terme javan.” = Acronychia?

“Wal-tiedde & Keipisan Zeylonens.” = ?
Apparently some menispermaceous plant, perhaps Tiliacora, is repre-
sented.

ANACARDIACEAE

Parishia maingayi Hook. f. Fl. Brit. Ind. 2: 30. 1876; King, Jour. As.
Soc. Bengal 65(2): 493. 1896 (Mater. Fl. Malay. Penin. 2: 779);
Rid]. Fl. Malay Penin. 1: 535. 1922.

Parishia elmert Merr. Univ. Calif. Publ. Bot. 15: 168. 1929, syn. nov.

I am now convinced that Parishia elmeri Merr. (1929), type from British
North Borneo (Elmer 21662) belongs with Hooker’s species and it is ac-
cordingly reduced to the latter. This gives its range as the Malay Penin-
sula, Sumatra, Riouw, and Borneo, a very natural one. The Sumatran and
Riouw specimens that I have seen are sterile.

Parishia malabog Merr. Philipp. Jour. Sci. 7: Bot. 281. 1912, Enum.
Philipp. Fl. Pl. 2: 472. 1928.

Spondias romblonensis Elm. Leafl. Philipp. Bot. 10: 3683. 1939, descr. angl.,

syn. nov.

The type of Elmer’s species, now reduced, is a staminate specimen, his
number 12164 from Romblon. See the note at the end of my original de-
scription of 1912. Luzon (Tayabas), Mindoro, Ticao, Masbate, Sibuyan,
Tablas, Romblon, Cebu, and Sibutu Islands. Endemic.

1954] MERRILL, MISCELLANEOUS MALAYSIAN NOTES 141

CELASTRACEAE
Celastrus paniculatus Willd. Sp. Pl. 1: 1125. 1798.

Alsodeia glabra Burgersdyk in Miquel, Pl. Jungh. 122. 1852; Mig. Fl. Ind.
Bat. 1(2): 116. 1858; Oudem. Arch. Néerl. 2: 199. pl. 9. 1867, syn. nov.

Rinorea glabra O. Kuntze, Rev. Gen. Pl. 1: 42. 1891, syn. nov.

The type of Alsodeia glabra Burgersdyk, a fruiting specimen, was col-
lected in Sumatra by Junghuhn. I had suspected from Oudeman’s excellent
illustration, because of the terminal panicle (in fruit) and other characters,
that a Celastrus was represented and not a representative of the violaceous
Alsodeia = Rinorea. Accordingly, while in Leiden I looked up the type
and found that Hallier had already made the transfer to Celastrus in the
herbarium but that he considered that a valid species of that genus was
represented; I can find no record of his having published this conclusion.
To me the Junghuhn specimen represents a form of the widely distributed
Celastrus paniculatus Willd., type from India, the species, as currently
interpreted, extending to Ceylon, Burma, Siam, Indo-China, southeastern
China, the Philippines, Malay Peninsula (and now Sumatra), Java, and
Timor. I have seen no specimens from Borneo, although it is to be ex-
pected there, as well as in Celebes and the Moluccas. Koorders, who in
1909 studied the type of Fliiggea ? serrata Miq. (1858) from Java, cor-
rectly reduced that species to Celastrus, and further to C. paniculatus
Willd. Other synonyms currently placed here are: Celastrus alnifolius
D. Don, C. dependens Wall., C. multiflorus Roxb., C. rothianus Roem. &
Schult., C. metzianus Turcz., C. polybotrys Turcz., Ceanothus paniculatus
Roth., Scutia paniculata G. Don, and Diosma serrata Blanco.

Kurrimia robusta (Roxb.) Kurz, Jour. As. Soc. ee ay 73. 1870;
Pitard in Lecomte, Fl. Gén. Indo-Chine 1: 893.

Celastrus robustus Roxb. Fl. Ind. 2: 395.

Bhesa moja Ham. ex Arn. Edinb. ae Philos. en 16: 315. 1834.

Rhesa moja Walp. Repert. 1: 538. 1

Kurrimia pulcherrima Wall. List no. yer 1830, nom.; Laws. in Hook. f. Fl.

Brit. Ind. 1: 622. 1875, descr.
Nothocnestis sumatrana Mia. FI. Ind. Bat. Suppl. 531. 1862.
Sarcosperma tonkinense H. Lecomte, Bull. Mus. Hist. Nat. Paris 24: 534.
Fl. Gén. Indo-Chine 3: 914. 1930, syn. nov.

In their critical treatment of the Sarcaspermataceae, Lam and Varos-
sieau, Blumea 3: 198. 1939, who had seen Lecomte’s type, a fruiting speci-
men, correctly eliminated Sarcosperma tonkinense H. Lecomte from the
genus and family and concluded that it was probably not even a sapo-
taceous plant; no identification of it was suggested. While in Paris in
August 1950, I examined the type and at once the problem resolved itself.
Manifestly Lecomte’s type, Bon 3974, represents a species of the cela-
straceous Kurrimia, not a Sarcosperma. Direct comparisons then made
showed that it was a fruiting specimen of the rather common and widely

142 JOURNAL OF THE ARNOLD ARBORETUM _ [vot. xxxv

distributed Kurrimia robusta (Roxb.) Kurz, which extends from Khasia
and Silhet to Burma, Siam, and Indo-China southward to Singapore and
Sumatra. I think that Nothocnestis sumatrana Miq. FI. Ind. Bat. Suppl.
531, 1862, is correctly placed as a synonym of Roxburgh’s species, for
although I have not seen its type I take Netherl. Ind. For. Serv. 31688,
32124 from Palembang, Sumatra, the type locality of Miquel’s monotypic
genus, to represent it, and I refer these two modern collections to Kurrimia
robusta (Roxb.) Kurz.

RHAMNACEAE
Ventilago gamblei nom. nov.

Ventilago lanceolata Gamble, Kew Bull. 1916: 134. a Alston in Trimen
Hand-book FI. Ceyl. Suppl. 49. 1931, non Merr. (1915).
A new name is needed for this species of southern India and Ceylon, as
the one selected by Gamble in 1916 had been used by me for a different
Philippine species one year earlier.

ELAEOCARPACEAE

Elaeocarpus tectorius (Lour.) Poir. in Lam. Encycl. Suppl. 2: 704.
1812, excl. fruct.; Merr. Trans. Am. Philos. Soc. II. 24(2) : 256. 1935.

Craspedum tectorium Lour. F]. Cochinch. 336. 1790, i eee 411.1793.

Dicera craspedum J. F. Gmel. ex DC. Prodr. 1: 520.

Elaeocarpus robustus sensu Merr. Jour. Arnold . ee 189. 1951, excl. syn.

plur., non Roxb. § Chascanthus.

This is an endemic species of Indo-China erroneously placed as the
equivalent of Elaeocarpus robustus Roxb., the type of which was from
Silhet, India. In the rather extensive synonymy cited by me in 1951 only
Craspedum tectorium Lour. and Dicera craspedum J. F. Gmel. actually
belong with the Loureiro Elaeocarpus species above cited. Loureiro’s genus
and species were based on specimens from the vicinity of Hue, Indo-China.
Clemens 3688 (in flower) and 4156 (in fruit) were collected near Tourane,
which is about 100 kilometers south of Hue. Clearly most of the Cochin-
chinese species characterized by Loureiro were observed in the vicinity
of Hue, where he lived for many years. Both the Clemens’ numbers are
in the Paris and the University of California herbaria, and the first is also
at Kew and the Arnold Arboretum. In 1951 I very critically compared the
Kew specimen with the Loureiro one at the British Museum, and surely a
single species is represented. Clemens 3688 is with mature flowers: the
Loureiro specimen has very immature buds.

The Loureiro original description is ample and well prepared. He un-
fortunately “guessed” at the fruit characters in his generic description of
Craspedum, and for this reason Corner illogically refused to recognize
Craspedum tectorium Lour. as worthy of consideration. I disagreed with

1954 | MERRILL, MISCELLANEOUS MALAYSIAN NOTES 143

him then as I do now, for all one has to do is to exclude the description
of the fruit in Loureiro’s generic account of Craspedum; the specific de-
scription was wholly based on a flowering specimen.

Dr. Gagnepain in 1943 (Not. Syst. 11: 1-14) increased the number of
Indo-Chinese species of Elaeocarpus by describing twelve new ones. He
did not indicate the sections in accordance with Schlechter’s scheme of
classification, but no less than eight of them belong in Chascanthus by the
indicated number of ovary cells (3) and the number of ovules in each
cell (2).

This note is prepared to correct my error of 1951 when I misinterpreted
Loureiro’s species by reducing it to Elaeocarpus robustus Roxb. I add
brief descriptive data based on the Loureiro specimen in the British Mu-
seum and Clemens 3688, 4156 from near the type locality. Dr. Gagnepain
left the two Clemens collections in the Paris herbarium without comment
under the binomial Elaeocarpus tectorius (Lour.) Poir. which I assigned
to them when the identifications were made by me and the sets of dupli-
cates were distributed in 1927-28. I can only assume he approved of my
identifications. I am indebted to Dr. Tardieu-Blot for checking the
specimens.

Leaves nearly or quite glabrous, 7-13 cm. long and 3—5.5 cm. wide;
lateral nerves 7 to 9 or 10 pairs; petiole 2—3.5 cm. long; flowers about
1 cm. in diameter, the petals 20—-30-laciniate; ovary hairy, 3-celled, cells
2-ovulate. Fruit ellipsoid, sharply apiculate, practically glabrous, the bony
endocarp very rugose. A photograph of the Loureiro specimens, a carbon
rubbing of a full-grown leaf, and extensive notes are in the Arnold Arbo-
retum herbarium; this was a second and later collection by Loureiro, sent
by him as representing his species. The specimen is not the actual type.

DILLENIACEAE
Saurauia costata Reinw. ex de Vriese, P]. Ind. Bat. Or. 56. 1856.

Saurauia warburgii Koord. Meded. Lands Plant. 19: 354, 644. 1898, Suppl.
Fl. Celeb. 2: pl. 80. 1922 (poor), 3: 39. 1922, syn. nov.

Reinwardt’s species was based on material collected by him in October
1821 on Mount Sempo, Celebes, there being three sheets of this collection
in the Rijksherbarium, Leiden. It is suspected that Koorders did not see
these specimens, for otherwise he would scarcely have proposed S. war-
burgiit Koord. as new in 1898. The latter was based on Koorders 18954,
19283, from Minahassa Province, northeastern Celebes. The several sheets
representing the two supposedly different species are so similar that all
might have been taken off the same plant or stand of plants.

De Vriese, in publishing Reinwardt’s description, noted that this Celebes
form resembled a Philippine collection, Cuming 455, which de described,
l.c., as Saurauia exasperata De Vr.; this is, however, Saurauia latibractea
Choisy (1854) as I understand that species, one that clearly is not closely
allied to S. costata Reinw. Koorders stated that his new species was allied

144 JOURNAL OF THE ARNOLD ARBORETUM hyo. xxxv

to the Philippine Saurauia elegans (Choisy) F.-Vill. (S. rugosa Turcz.),
and Stapf had noted on one of the Reinwardt sheets that it was “very near
S. rugosa Turcz.” In the latter species, which occurs in most provinces in
Luzon and in Mindoro, the lax, long-peduncled, many-flowered inflores-
cences are up to 16 cm. long and 10 to 12 cm. wide, while its smaller leaves
are, as Koorders noted, rounded, not cordate, at the base. He described
the inflorescences of FE. warburcii Koord. as only 2.5 cm. long. Although
the vegetative characters of the two species are suggestively similar, I do
not think that they are closely allied.

Saurauia lanceolata DC. Mém. Soc. Phys. Hist. Nat. Genéve 1: 421.
1821, quoad descr., excl. 44 4; DC. Prodr. 1: 526. 1824; De Vriese,
Pi. Ind. Bat. Or. 39, 18

This is a very curious case. It is suspected that most authors have
interpreted De Candolle’s species from his distinctly good plate. His
technical description was based wholly on a Javan specimen, Leschenault
643, the type being in the Paris herbarium. An excellent photograph of
this type, kindly supplied by Madame Tardieu-Blot, is before me, includ-
ing even the supplementary sheet with sketches of the flower and dissection
notes. As will be seen from the photograph, the Leschenault specimen
(no. 643) has only slightly developed inflorescences. This type should be
compared critically with that of Saurauia micrantha Blume from Mt. Gede.
Although no exact locality in Java is given for S. lanceolata DC., it would
have to be from a readily accessible place, such as Mt. Gede.

The strange thing is how De Candolle’s plate became associated with
the Javan species. Zollinger, Syst. Verzeichn. 148. 1854, noted the great
similarity of illustration to the South American S. ruiziana Steud., stating:
“S. lanceolata DC. Mem. t. IV tam similis est S. Ruizianae Steud.
(Ap[ateria] lanceolata D.C) ut nullomodo distincta videatur quamvis
petalis basi coalitis. An de patria error quisdam in herbario Parisii adfuit?”
Madame Tardieu-Blot says: “Le type est glabre (photo) et correspond
a la description. Au contraire, la figure (Pl. IV) est trés differente (velue)
et cadre trés bien avec S. Ruiziana.’’ De Candolle’s specific epithet has
priority over any other designating a plant with which this tvpe can be
matched.

GUTTIFERAE

Calophyllum rotundifolium Ridl. Jour. Fed. Malay States Mus. 5: 22.
1914; Fl. Mal. Penin. 1: 188. 1922

British North Borneo, Mount. Kinabalu, Clemens 30984, 31428, 35038,
40705, 50316. The altitudinal range is indicated on two labels as 4,000
and 5,000 ft. I cannot distinguish this strongly marked species from
Ridley’s type at Kew. The sessile, broadly cordate, suborbicular to broadly
ovate leaves vary in length from 2 to 7 cm., their apices rounded or very
broadly rounded occasionally slightly retuse. Malay Peninsula (Selangor),
alt. 5000 ft. New to Borneo.

1954 | MERRILL, MISCELLANEOUS MALAYSIAN NOTES 145

VIOLACEAE
Rinorea lanceolata (Roxb.) O. Kuntze, Rev. Gen. Pl. 1: 42. 1891.

Vareca lanceolata Roxb. Fl. Ind. 2: 446. 1824, ed. 2, . is 1832.

Celastrus pauciflora Wall. in Roxb. Fl. Ind. 2: 400. 1

Pentaloba lanceolata Wall. List no. 4023. 1830, nom.

slr ll p ie Jack ex Griff. Calcutta a Nat. Hist. 4: 195.
4

rie ae Oudem. Arch. Néerl. 2: 196. pl. 6. 1867; Hook. f. Fl. Brit.
Ind. 1: 188. 1872; King & Gamble, Jour. As. Soc. Bengal 58 Aye 404. 1889
(Mater. FI. Malay. Penin. 1: 48); Ridl. Fl. Malay Penin. 1: 131. 1922.

For a rather sharply characterized species known only from Penang
Island, this has accumulated a considerable synonymy. My attention was
called to it through an attempt to place the generally ignored binomials
Pittosporum ? serrulatum Jack and Celastrus pauciflora Wall., both based
on Penang material. Jack sent a copy of his description to Wallich, who
realized at once that no Pittosporum was represented, and, as he says he
had no specimens from Jack, he apparently surmised from the description
that Celastrus might be the proper place for it and so described it as his
own species, although stating that the description was from Jack. Griffith,
who edited the Calcutta reprint of Jack’s plant descriptions in 1843, says
that his data were from Jack’s MS., yet these are the same as those pub-
lished under Celastrus pauciflora Wall. It is suspected that Jack did send
a specimen to Wallich which the latter failed to associate with Jack’s man-
uscript description. Under Vareca lanceolata Roxb. Flora Ind. 2: 446.
1824, Wallich states: “I have specimens belonging probably to this plant,
which were collected at Pinang by W. Jack, who in a Mss. note says:
‘I am at a loss what to make of this shrub. I thought it might be a Vareca
(according to Roxburgh) but the capsule is one-celled, three-valved, with
parietal placentae.’”’ There is a specimen of Wallich 4023, type collection
of Pentaloba lanceolata Wall. which, as far as the record goes, was not
based on Vareca lanceolata Roxb., and two modern collections of the
species from Penang in the herbarium of the Arnold Arboretum. As far as
the two somewhat generalized descriptions of Jack and of Wallich are
concerned, they agree in all respects with the characters of Rinorea lanceo-
lata (Roxb.) O. Kuntze, the type of which was from Penang.

Rinorea semigyrata (Turcz.) J. J. Sm. in Koord. & Val. Meded. Dep.
Landbouw 18: 73. 1914 (Bijdr. Boomsoort. Java 13: 73).

Pentaloba semigyrata Turcz. Bull. Soc. Nat. Moscou 27 (2): 342. 1854.

Alsodeia disticha Zoll. ex Teysm. & Binn. Cat. Hort. Bogor. 183. 1866, nom.,
et in Miq. Ann. Mus. Bot. Lugd.-Bat. 4: 216. 1869, mom., syn. nov.

Alsodeia semigyrata Turcz. ex Jacks. Ind. Kew. 1: 93. 1893.

Alsodeia paradoxa Blume ex Oudem. a Folge 2: 204. pl. 15. 1867, et in
Miq. Ann. Mus. Bot. Lugd.-Bat. 3: 71. 1867.

Rinorea paradoxa J. J. Sm. in Koord. . Val. op. cit. 67; Van Ooststr. in
Backer Beknopte Fl. Java 4a (1): Fam. 48. 3. 1942.

146 JOURNAL OF THE ARNOLD ARBORETUM _ [vot. xxxv

Turczaninow’s specific name is apparently the oldest valid one for this
rare, or perhaps even extinct Javan species. His type was Zollinger 2979,
and a duplicate of this collection is in the herbarium of the Arnold Arbo-
retum, acquired in 1941 from the Boston Museum of Natural History,
having originally been in the private herbarium of John Amory Lowell.
The original description is short and unsatisfactory, leading J. J. Smith,
who reproduced it in 1914, to think that perhaps some genus other than
Rinorea was represented; yet he actually effected the transfer of the
specific epithet to Rinorea. Van Ooststroom in 1942 apparently saw only
the Blume material at Leiden, there being at least four sheets labeled by
Blume as Alsodeia paradoxa; these specimens came from the Salak and
Boerangrang Mountains, western Java. The region has been very inten-
sively explored in the past century. He noted that the species had ap-
parently not been collected since Blume’s time. I cannot distinguish
Zollinger 2979 from these Blume specimens, its label merely indicating
that it was collected in Java. It also bears an unpublished Zollinger bino-
mial in Jmhofia. This leads me to reduce, without question, the nomen
nudum, Alsodeia disticha Zoll., as this specific epithet is the one that ap-
pears on our Zollinger specimen sub Jmhofia. It is suspected that at first
Zollinger thought that he had a representative of a new genus and then
found that the generic name originally assigned to his number 2979 was
a preoccupied one. Teijsmann and Binnendijk, who first printed the Zol-
linger binomial, merely indicate the species as being from “Ind. or.’’ Some
of the higher numbers of the Zollinger collection distributed under Java
labels were actually from Sumatra.

I note that although Rinorea semigyrata J. J. Sm. was legitimately
published in 1914, it was overlooked by the compilers to the supplements
to Index Kewensis. Further I note that the binomial Alsodeia semigyrata
was not actually published by Turczaninow, Bull. Soc. Nat. Mosc. 36(1):
559. 1863, as currently accepted. All that he stated was that his three
species of Pentaloba (P. corylifolia, P. fasciculata, and P. semigvrata)
represented species of Alsodeia, but he made no actual transfers.

LYTHRACEAE
Lagerstroemia speciosa (Linn.) Pers. (L. flos-reginae Retz.)

I have always been impressed by the apparent reluctance of certain
taxonomists to accept changes in the accepted names of plants on the basis
of the priority rule, no matter what the evidence is, and no matter what
botanist upset the use of a generally accepted binomial strictly in accord-
ance with the Code. A case in point is the very common and widely dis-
tributed Lagerstroemia speciosa (Linn.) Pers. versus the later L. flos-
reginae Retz. When Koehne’s monograph of the Lythraceae appeared
(Pflanzenreich 17 (IV. 216): 1-326. fig. 1-59. 1903) he correctly ac-
cepted the Linnaean specific epithet, and as he was known to be a very
critical worker, I did not hesitate to accept his conclusions. The species

1954] MERRILL, MISCELLANEOUS MALAYSIAN NOTES 147

extends from northern India southward and eastward through Malaysia
to New Guinea and northeastern Australia, and is, I suppose, one of the
most frequently collected of the arborescent species because it is so com-
mon and so conspicuous, when in flower, in most parts of its very wide
range. From Koehne’s references it is manifest that he had checked the
first publication of the name-bringing binomial. I was therefore somewhat
surprised to note that Mr. Corner, Gard. Bull. Straits Settl. 10; 272.
1939, rather curtly rejected ZL. speciosa (Linn.) Pers. and accepted the
later L. flos-reginae Retz. He rested his case on King’s statement, Jour.
As. Soc. Bengal 67(2): 9. 1898, and manifestly did not check the original
documents. King’s whole argument is merely that the acceptance of the
Linnaean specific name of 1771 was inadmissible, as ‘‘he describes
M{unchausia| speciosa as a shrub [which is true, as Linnaeus does say
“arbuscula”; he had a plant grown in a greenhouse in Germany], whereas
this plant [L. flos-reginae Retz.| is a large tree; moreover the rest of the
description would apply to various other species of Lagerstroemia.” The
last part of this statement is true, for the Linnaean description of 1771 is
very short and very unsatisfactory. Yet the species is by no means always
a large tree; I have seen immature plants in full flower on Luzon that
were not more than 2 m. high. But what Koehne did not overlook, as did
both King and Corner, is that the original description of Munchausia
jpeg Linn. was published in Muenchhausen’s Der Hausvater 5: 357.
pl. 2. 1770. This description is not only an ample one, but it is accom-
ne by a really excellent plate; and the plant described and illustrated
is in all respects L. flos-reginae Retz. = L. speciosa (L.) Pers. Yet Corner
would maintain Retzius’ specific name even at this late date. The descrip-
tion. according to Muenchhausen’s own statement, was prepared by Lin-
naeus. in spite of the fact that a year later Linnaeus credited it to Muench-
hausen. The specimens on which the description and the illustration
were based were from a plant cultivated in the Botanical Garden at
Goettingen, the source of it being stated as Java, and at the same time the
common Javanese name dboengoes was listed, the modern boengoer which
is widely used in western Java. Muenchhausen said Java and China, —
Linnaeus only China.

Turning to the Linnaean herbarium, a set of photographs of all the
sheets being available at the Arnold Arboretum, there are three sheets in
the Munchausia cover, none of them actually named by Linnaeus; but two
of these are manifestly this very common Lagerstroemia, and for the most
part the plate illustrating the species might have been drawn from these:
and in Linnaeus’ handwriting on one of these is the name boengoes.

We who attempt to apply the approved rules of nomenclature in the
determination of the oldest valid specific name for this or that species are
always subject to criticism on the part of those who apparently abhor
changes in names of well-known species. Generally speaking it will prob-
ably be admitted that those who make changes try to do the best that
they can with the data which are available to them. We all make mistakes,
but that is no reason why we should be tacitly condemned merely because

148 JOURNAL OF THE ARNOLD ARBORETUM _ [vot. xxxv

we do, on occasion, correctly interpret a Linnaean type. There is no ap-
proved method whereby the oldest specific name can be abandoned in
this case.

MYRSINACEAE
Ardisia oligocarpa nom. nov.

Ardisia oligantha Elm. Leafl. Philipp. Bot. 4: 1496. 1912; Merr. Enum. Philipp.
Fl. Pl. 4: 260. 1923, non Baker (1885), nec Mez (190 2).

The type and only known collection is Elmer 12310 from Sibuyan, the
flowers unknown. The species belongs in the subgenus Pyrgus in the alli-
ance with Ardisia perrottetiana A. DC. and A. serrata Pers., but has much
smaller, differently shaped, entire or nearly entire leaves, and short in-
fructescences with very few fruits and apparently with very few flowers.

Ardisia diversifolia Koord. & Val. Meded. Lands Plant. 33: 249, 1900
(Bijdr. 5: 249).
Ardisia oligantha Mez, Pflanzenr. 9 (IV. 246): 134. 1902, syn. nov.
Ardisia javanica A. DC. var. oligantha Blume ex Scheff. Comm. Myrsin.
Archipel. Ind. 74. 1867.

The Mez binomial was based on a single Reinwardt specimen from
Mount Salatta, Java, in the Rijksherbarium, Leiden. It is not accounted
for in Backer’s recent treatment of the Myrsinaceae of Java, Beknopte
F]. Jav. VIIB Myrsinaceae 1-20. 1948. I do not see how this can be
distinguished from the Koorders and Valeton species, of which Mez saw
no specimens. He placed the latter next to his new one, stating “ex de-
scriptione sequenti [A. oligantha Mez] peraffinis’; and in his key to the
species on page 73 he apparently could discover no characters by which
the two could be distinguished, merely inserting the Javan A. diversifolia
Koord. & Val. between A. verrucosa Presl, a Philippine species. and 4.
oligantha Mez of Java, but indicating no separating characters

Ardisia taytayensis nom. nov.

Ardisia pachyphylla Merr. Philipp. Jour. Sci. 12: Bot. 157. 1917. Enum.
Philipp. Fl. Pl. 3: 261. 1923, non Dunn (1912).
A species known from Palawan and Balabac, Merrill 9188, 9216, Escritor
Bur. Sci. 21613.

Discocalyx papuana (S. Moore) comb. nov.

Embelia papuana S. Moore, Trans. Linn. Soc. Bot. II. 9: 106. 1916.

The type and only known collection is a Boden-Kloss specimen, Wol-
laston expedition from Camp VIb Mt. Carstensz, New Guinea, alt. about
1200 m. This is clearly a small erect shrub, not scandent as are all repre-
sentatives of Embelia; Moore did not indicate its habit, and there are no
notes

1954] MERRILL, MISCELLANEOUS MALAYSIAN NOTES 149
Embelia cotinoides (S. Moore) comb. nov.
Maesa cotinoides S. Moore, Trans. Linn. Soc. Bot. II. 9: 103. 1916.

The type of this in the herbarium of the British Museum is an excellent
specimen collected by C. Boden-Kloss at Camp Vla Mt. Carstensz alt.
940 m. on the Wollaston expedition to New Guinea, 1912-13. It is in all
respects an Embelia of the subgenus Evembelia in the general alliance
with Embelia sarasinorum Mez of Celebes and the more widely distributed
Malayan F. coriacea Wall. but is very distinct from both.

EMBELIA ? LUCIDA Wall. List no. 2315. 1830, nom. nud., et ex A. DC. in
Trans. Linn. Soc. 17: 134. 1834, descr., Prodr. 8: 87. 1844 = Anti-
desma coriaceum Tul. Ann. Sci. Nat. Bot. ITI. 15: 204. 1851, non
Antidesma lucidum Merr. (1906).

When a species is described as new but placed not only in the wrong
genus but in a family remote from the one to which it belongs, it is some-
times worth while to prepare a record when a correct reduction can be
made, Although Mez saw the holotype in the De Candolle herbarium, he
was unable to place the species, merely indicating that it did not belong
in the Myrsinaceae. At my request Mr. Burkill examined Wallich 2315
at Kew, the type collection from Singapore, and reported to me that, al-
though no flowers are left on the specimen (it was apparently pistillate,
judging from De Candolle’s description), Embelia ? lucida Wall. is safely
the same as Antidesma coriaceum Tul. Other synonyms are Antidesma
fallax, Muell.-Arg. (1865) and A. pachyphyllum Merr. (1916). Wallich’s
name cannot be used in Antidesma because of the different Antidesma
lucidum Merr. (1906). Penang, Malay Peninsula, Singapore, Borneo;
planted at Bogor, Java, fide Pax & Hoffmann.

Maesa megaphylla Merr. Philipp. Jour. Sci. 12: Bot. 158. 1917, Enum.
Philipp. Fl. Pl. 3: 255. 1923.

Maesa lobuligera Mez, Repert. Sp. Nov. 16: 310. 1920.

Maesa megalobotrys Merr. op. cit. 20: 422. 1922, syn.

Maesa celebica Koord. ex Koord.-Schum. Syst. Verseich. Herb. Koord. 3:

100. 1914, nom. nud., syn. nov.

The type of Maesa megaphylla Merr. was Wenzel 1510 from Leyte,
that of M. lobuligera Mez, Foxworthy Bur. Sci. 727 from Palawan, that
of M. megalobotrys Merr., Merrill 9176 from Palawan, and that of M.
celebica Koord., Koorders 18174 from Minahassa, northeastern Celebes.
Had Koorders published a description of the latter, then his specific name
would be the accepted one for this rather strongly marked species. The
specimen of Koorders 18174 at the Rijksherbarium, which I have seen,
is fragmentary and sterile, consisting of a branchlet and two large glabrous
detached leaves, these about 15 to 18 cm. long and 9 to 12 cm. wide. It
is suspected that the reason why Koorders published no description is that
he had only sterile material. In such a critical genus as Maesa, where

150 JOURNAL OF THE ARNOLD ARBORETUM _ [vot. xxxv

species are often distinguished by very slight characters, some might
hesitate in reducing a binomial to synonymy where only vegetative parts
are known, but the agreements in this case are so close, and the flora of
northern Celebes is so similar to that of the Philippines, that I do not
hesitate to place Koorders’ name in synonymy. It is perhaps unreason-
able to discuss the reduction of a mere nomen nudum in detail. Actual
specimens now available to me are: Leyte, Wenzel 1275, 1510, 1627 (two
of these distributed under another binomial invalidated by Maesa platy-
phylla Elm.); Bohol, Ramos Bur. Sci. 43324; Dinagat, Ramos & Con-
vocar Bur. Sci. 84657; Celebes, Minahassa, Koorders 18174. No specimen
of Merrill 9176 is at present available to me. From the description I had
surmised that Maesa megalobotrys Merr. was not distinct from M. mega-
phylla Merr., and Dr, E. H. Walker, after examining an isotype of the
former in the U. S. National Herbarium, confirms this reduction of it.
It is also recorded from Palawan and from Mindanao.

OLEACEAE

Ligustrum robustum (Roxb.) Blume, Mus. Bot. Lugd.-Bat. 1: 313.
1850; Mansf. Bot. Jahrb. 59: Beibl. 132: 44. 1924, cum syn.

Phillyrea robusta Roxb. Fl. Ind. 1: 101. 1820.
Olea puberula Ridl. Jour. Straits Br. Roy. As. Soc. 59: 128. 1911, Fl. Malay
Penin. 2: 318. 1923, syn. nov.

In checking certain types of Olea at Kew in 1950 I noted a duplicate
type of Olea puberula Ridl., ie., Ridley 15223 from Perlis, Malay Penin-
sula, and although the specimen is a poorly prepared one, manifestly
Ligustrum rather than Olea is represented; this might be suspected from
Ridley’s description of his species as having terminal panicles. The note
made at the time is to the effect that it was apparently a form of Ligustrum
robustum (Roxb.) Blume with inflorescences narrower than in the typical
form. As Roxburgh’s species is currently interpreted, it extends from
Nepaul, Silhet, Assam, and Bengal to Chittagong, Burma, Siam, Indo-
China, and Sumatra, so that this extension of range to the Malay Penin-
sula is not surprising. For the present I prefer thus to dispose of Ridley’s
species, rather than by transferring his specific name to Ligustrum; in any
case this is the first record of the genus Ligustrum for the Malay Peninsula.

Linociera pachyphylla sp. nov.

Arbor glabra, inflorescentiis parcissime pubescentibus exceptis, ramulis
ultimis subpallidis, teretibus, consperse lenticellatis, 3-4 mm. diametro,
nodis superioribus plus minusve compressis; foliis crasse coriaceis, rigidis,
siccis utrinque brunneis vel castaneis, subconcoloribus vel subtus paullo
pallidioribus, supra nitidis, oblongis vel oblongo-ellipticis, 18-27 cm.
longis, 7-9 cm. latis, breviter subobtuse acuminatis, basi late acutis, nervis
primariis utrinque 10-12, supra impressis, subtus elevatis, perspicuis,
circiter ad marginem valde curvatis, fere anastomosantibus, reticulis pri-

1954] MERRILL, MISCELLANEOUS MALAYSIAN NOTES 151

mariis laxis, obscuris, secundariis obsoletis; petiolo crasso, 1.5-2 cm.
longo; inflorescentiis axillaribus, solitariis, breviter pedunculatis vel e basi
ramosis, 6—7 cm. longis, ramis primariis patulis vel adscendentibus; flor-
ibus numerosis, breviter pedicellatis, bracteis inferioribus crassissime
coriaceis, ovatis, concavis, circiter 3 mm. longis, superioribus brevioribus;
petalis fere liberis, loriformibus, obtusis, 5 mm. longis, basi circiter 1.5 mm
latis, sursum vix 1 mm, latis; antheris 1.1 mm. longis.

BORNEO: Sarawak: Native collector 584, notes lost.

This was distributed as representing Linociera callophylla (Blume)
Knobl., but it proves to be remote from the species that Blume character-
ized. It belongs in the group with Linociera pluriflora Knobl., differing in
its very rigid, thickly coriaceous leaves and shorter inflorescences. Other
species in this assemblage are L. nitens Koord. and L. verruculosa Merr.,
but here again the vegetative characters alone separate this proposed new
species.

Linociera stenura sp. nov.

Arbor glaberrima, ramulis ultimis gracilibus teretibus, 1 mm. diametro;
foliis coriaceis, in sicco pallidis, vix vel obscure nitidis, utrinque sub-
concoloribus, lanceolatis vel oblongo-lanceolatis, basi acutis, 11-15 cm.
longis, 2-4 cm. latis, ab infra medium sursum gradatim angustatis, apice
longe graciliter caudato-acuminatis, acuminibus 1-2 cm. longis, obtusis;
nervis primariis utrinque circiter 12, inter se plerumque 1-— 1.5 cm. dis-
tantibus, patulis, obscuris, 2-3 mm. a margine confiuentibus, reticulis
laxissimis, obscuris vel subobsoletis: petiolo 2.5 mm. longo; inflorescentiis
axillaribus, solitariis, brevibus paucifloris, circiter 8 mm. longis, glabris;
floribus usque ad 9, plerumque 3-5, breviter (1-1.5 mm.) pedicellatis,
4-meris, petalis linearibus, circiter 5 mm. longis, deorsum leviter ampliatis
sed basi vix 0.5 mm. latis.

CELEBES: Malili District, G. Kjellberg 2120, August 19, 1929, a tree with
white flowers on riverbank at Waraoe, altitude 50 m., TYPE in the Stockholm
herbarium, a fragment in the recoil of the Arnold Arboretum.

A species strikingly characterized by its distantly and obscurely nerved
leaves which are very slenderly caudate-acuminate, gradually narrowed
upward from below the middle, the obscure nerves spreading at nearly
right angles and anastomosing with the equally inconspicuous and some-
what arched submarginal nerves 2-3 mm. from the leaf margin, the reticu-
lations very lax, obscure or even subobsolete. Its few-flowered, axillary,
solitary racemes are less than 1 cm, in length, even including the 5 mm.
long very narrow petals.

Linociera ridleyi nom. nov.

Linociera ee Ridl. Jour. Fed. Malay States Mus. 8 (4): 61. 1917,
non Knobl.

152 JOURNAL OF THE ARNOLD ARBORETUM _ [vot. xxxv

The type of Ridley’s species is a specimen collected on Korinchi Peak,
Sumatra, at an altitude of about 7300 ft. He stated that it is allied to
Linociera montana (Blume) DC. of Java.

Linociera sp.

Microtropis ? lanceolata Boerl. & Koord. in Koord.-Schum. Syst. Verzeich.
Herb. Koord. 2: 33. 1911

The type of this species is Koorders 10283 from Sumatra, a fruiting
specimen which in 1940, on the basis of an actual examination of the type
in the Bogor herbarium, we could not place other than as a species of
Linociera, perhaps allied to L. oligantha Merr.; see Merrill & Freeman,
“The Old World species of the celastraceous genus Microtropis Wallich,”’
Proc, Am. Acad. Arts Sci. 73: 307. 1940. It seems as unwise now to pro-
pose a new name for this imperfectly known species in Linociera as it
did to us in 1940, However, to call attention to this misplaced entity, it
has been considered desirable to make a record of it in the family and
genus to which the species manifestly belongs. Linociera lanceolata Knobl.
(1933), the name of a Santo Domingo species, invalidates the use of the
epithet of Boerlage and Koorders in Linociera.

APOCYNACEAE

Micrechites micrantha (Miq.) Hallier f. Jahrb. Hamb. Wissensch.
Anstalt. 17: 156. 1899 (1900), im nota; Bakhuizen v. d. Brink in
Backer, Beknopte Fl. Jav. 7B: Apoc. 32. 1948, Blumea 6: 389. 1950.

Otopetalum micranthum haps Fl. Ind. Bat. 2: 400. 1857, Versl. Meded. Kon.
Akad. Wetensch. 6: 191. 1857.
Ecdysanthera schrieckii v: Huerck & Muell.-Arg. Obs. Bot. 191. 1870
Micrechites schrieckii Rolfe, Jour. Bot. 23: 214. 1885: be Phan. ‘Cuming.
Philipp. 126. 1885; Merr. Enum. Philipp. Fl. Pl. 3: 332.
Micrechites polyantha sensu King & Gamble, Jour. As. ei “Bengal 74 (2):
504. 1907 [ Mater. Fl. Malay. Penin. 4 (no. 19): 714], non Miquel.
Trachelospermum philippinense Elm. Leafl. Philipp. Bot. 2: 488. 1908.
Micrechites furcata Ridl. Jour. Roy. As. Soc. Straits Br. 79: 95. 1918, Fl.
Malay Penin. 2: 368. 1923
This adjustment in the nomenclature of Micrechites schrieckii Rolfe,
long considered to be a Philippine endemic, is necessary. Hallier f. in 1900,
on the basis of a critical examination of its type specimen, determined the
status of Otopetalum micranthum Miq. Miquel’s description of its fruits
s “Bacca corticata, semina intra pulpam fibrosum recepta (adhuc valde
immatura),” and again in the species description as “baccae globosae
ovoideae” is, of course, an impossible one for Micrechites, the fruits of
which are slender follicles; what he actually mistook for immature fruits
were corolla tubes deformed by insects. Boerlage, Handl. Fl. Nederl. Ind.
2: 379-380. 1899, had discussed this case previous to Hallier’s extensive
consideration of it. The latter actually utilized nearly six pages of print

1954 | MERRILL, MISCELLANEOUS MALAYSIAN NOTES 153

in his greatly detailed study of the problem, and even then succeeded in
burying his new binomial in the text (p. 156) as “Micrechites micrantha
m.” where all bibliographers overlooked it for nearly half a century (it
still is not listed in Index Kewensis); and at the same time its name-
bringing synonym is equally buried in the text on page 152. I suspect that
even Bakhuizen van den Brink might have overlooked this strangely
published new binomial but for the fact that Hallier had added a reference
on the type sheet to “Kautsch, Lianen p. 156. 1900”; this I noted, in the
herbarium, and this it was that led me to Bakhuizen van den Brink’s 1948
consideration of the case. I accept the synonymy as given by him, and add
several other binomials.

The species, as I now understand it, extends from Siam and the Malay
Peninsula (Maingay 1081) to Sumatra, Java, and also occurs more or
less throughout the Philippines. There are, however, available for study
only comparatively few collections except for the Philippine area, where
about twenty are available. Among the previously unlisted ones are
McGregor Bur. Sci. 47371 from Tayabas Province, Luzon, Ebalo 940 from
Basilan, and Wenzel 3403 from Surigao Province, Mindanao. There are
in the Leiden herbarium certain Javan collections named by Blume as
representing Tabernaemontana polyantha Blume which actually represent
Micrechites micrantha (Miq.) Hall. f., but there are also other specimens
which represent Micrechites polyantha (Blume) Mig. as currently under-
stood, I follow Bakhuizen van den Brink in his interpretation of M. mz-
crantha (Miq.) Hall. f.

VERBENACEAE
Clerodendron fortunei Hemsl. Jour. Linn. Soc. Bot. 26: 259. 1890.
srcesicls simile Merr. Govt. Lab. Publ. 35: 64. 1906, non Pearson (1901),

syn.
Pe pon 6 ie mindorense Merr. oe Jour. Sci. Bot. 7: 342. 1912, Enum.
Philipp. Fl. Pl. 3: 404. 1923, syn.

Hemsley’s description of thea fortunei was based on two sheets
of a Fortune collection merely indicated as coming from China. I had
seen the type in 1935 and was then impressed with its close resemblance
to certain Philippine forms. In September 1950 I again examined the
type and compared it with specimens representing several Philippine
species. I am now convinced that the Fortune specimens were taken from
cultivated plants in China, for nothing like it has appeared in the very
large collections made in southeastern and eastern China in the past thirty
vears. I do not hesitate to reduce the Philippine C. simile Merr. = C.
mindorense Merr. to C. fortunei Hemsl. Hemsley described the corollas
as 2 in. long; the longest one I observed was 4 cm. in length. Those of
C. mindorense Merr. are 3 to 4 cm. long (original description 3 cm.). An
allied form is C. klemmei Elm. from medium and higher altitudes in
northern Luzon, but its corollas are 6 cm. in th.

Chinese associations with the Philippines have extended over a period

154 JOURNAL OF THE ARNOLD ARBORETUM _ [vot. xxxv

of about 2000 years. It is well known that they introduced into the Archi-
pelago various economic plants, for a considerable number of these exotics
are still known in the Philippines only by their Chinese names. Occasion-
ally they also introduced an ornamental or merely curious species, as
exemplified by Vidal’s collection of the bignoniaceous Markhamia cauda-
felina (Hance) Craib in Albay Province, Luzon; see Sprague in Kew Bull.
1919: 310. 1919. The associations between Amoy and Manila were pecul-
iarly close, and it may be that the Fortune collection came from plants
cultivated at Amoy. On the other hand it is more than possible that they
were taken from plants cultivated in the Fatee Gardens, across the river
from Canton, which Fortune visited in 1843 and described the next year,
Gard. Chron. 1844: 590. 1844. The Fatee Gardens still exist, being a
series of nurseries on Lingnan Island where ornamental plants are prop-
agated and sold. In the Philippines the species occurs at low altitudes,
extending from northern Luzon to Mindoro, Semerara, Negros, and Min-
danao. Additional collections at Kew are Cuming 1475 from Batangas
Province, Luzon, Vidal 3453 from Laguna Province, Luzon, and Loker
4414 from conten Luzon. There is a photograph of Hemsley’s type in the
herbarium of the Arnold Arboretum.

Clerodendron cyrtophyllum Turcz. Bull. Soc. Nat. Moscou 36(1):
222. 1863.
Clerodendron amplius Hance, Ann. Sci. Nat. Bot. V. 5: 233. 1866.
Clerodendron formosanum Maxim. Bull. Acad. Sci. St. Pétersb. 31: 85. 1886,
Mél. Biolog. 12: 519. 1886.

Cordia venosa Hemsl. Jour. Linn. Soc. Bot. 26: 143. 1890.

My interest in looking up Hemsley’s type in 1950 was inspired by the
same motives as was the case with Clerodendron fortunei Hemsl.; that is,
that in the Boraginaceae nothing appeared in the very large modern col-
lections from Chekiang that have passed through my hands in the past
thirty years that even suggested the species Hemsley characterized. The
actual type of Cordia venosa Hemsl. is a fruiting specimen from Ningpo.
A casual examination of the type in August 1950 indicated what the diffi-
culty was, for it represents the very common Clerodendron cyrtophyllum
Turcz., Hemsley having erred in placing his fruiting specimen in the boragi-
naceous genus Cordia; as the type specimen was mounted one would con-
clude from a casual examination that its leaves were alternate; on the con-
trary they are opposite. Rehder had added to the sheet ‘“‘cf. Clerodendron,”
and later, Jour. Arnold Arb. 12: 76. 1931, had actually reduced Hemsley’s
species to Clerodendron cyrtophyllum Turcz. This very common and char-
acteristic species is now represented in the herbarium of the Arnold Arbo-
retum by approximately a hundred individual collections, twenty-five of
which are from Chekiang. Other areas represented are Anhwei, Hunan,
Kiangsu, Kwangsi, Kweichow, Fukien, and Kwangtung provinces, For-
mosa, Hainan, and Indo-China. There is a photograph of Hemsley’s type
in the herbarium of the Arnold Arboretum.

1954] MERRILL, MISCELLANEOUS MALAYSIAN NOTES 155

RUBIACEAE

Canthium tavoyanum (Parker) comb. nov.

Plectronia tavoyana Parker, Kew Bull. 1925: 429. 1925. Burma.

This change is made for the obvious reason that the generic name
Plectronia Linnaeus has been misapplied by many modern botanists. The
Linnaean type is a representative of the oliniaceous genus Olinia Thunberg.
See Merrill, Philipp. Jour. Sci. 35: 7. 1928.

Dentella serpyllifolia Wall. ms. “in sched. in Herb. Wall. sub 6206G.”
1832, nom. nud.; Craib, Fl. Siam Enum. 2: 27. 1932, nom. subnud.;
Airy-Shaw, Kew Bull. 1932: 289. 1932, descr.

LUZON: ae Province, ae Sci. 1398, August 1906, distributed as Dentella
repens J.R . Forst

Curiously Wallich’s sine name does not appear in his lithographed
List (generally cited as Catalogue), but Airy-Shaw apparently found it
on a subsidiary label in the master set at Kew. The entry in the distrib-
uted lithographed List is under Dentella repens Forst., 6206’G? Ripa
Irawaddi ad Henzada — 1826. (F[l]os distincta).” The species is now
known to extend from Assam and northern Bengal to Burma, Siam, Lom-
bok, Java, Mauritius, Luzon, and Guam (Fosberg). Curiously Dentella
repens J. R. & G. Forst. has also been found in Guam, an intermediary
stop, and at Acapulco, the terminus of the Manila-Acapulco galleon line
(1565-1815). Both species are ruderals probably in part distributed by
migratory birds, partly by man. This species is otherwise recorded from
Polynesia only from the Marquesas Islands.

Ophiorrhiza sarawakensis nom. nov.
Ophiorrhiza reticulata Ridl. Sarawak Mus. Jour. 1 (2): 32. 1912, non Korth.
(1851).

A new name is needed for the Bornean species characterized by Ridley
in 1912.

Psychotria polytricha Miq. FI. Ind. Bat. 2: 287. 1857.

Psychotria rufipila Val. Ic. Bogor. 3: 253, pl. 291. 1909, syn. nov.

Psychotria trichophlebia Merr. Mitt. Inst. Bot. Hamburg 7: 295. 1937, syn. nov.

I now think but a single species is represented here. Miquel’s species
was based on Korthals specimens from Sumatra and Borneo; that of Vale-
ton on Bornean specimens, Teysmann 8000 and Jaheri 509; and that of
myself on Winkler 1571, from Borneo. I have seen none of the cited
specimens representing Valeton’s species, but my concept of its limits is
gained from his excellent illustration and ample description. Beccari 822,
from Sumatra, in fruit, is currently referred to this species. There are

156 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XXXV

several old sheets at Leiden where the original records were lost or con-
fused, labeled “Sumatra,” “Moluccas” (this scratched), and “Java” (this
very doubtful). I suspect that these were all from Sumatra or Borneo.

Psychotria sangeana (Miq.) comb. nov.

Chasalia sangeana Mia. FI. Ind. Bat. Suppl. 546.
— rhodocarpa Teijsm. & Binn. Nat. oe Nederl. Ind. 27: 30.
864.

The type of Miquel’s species was a Teysmann specimen from Sumatra.
On the basis of the material in the Rijksherbarium representing two sup-
posedly distinct species in different genera, both sent by Teysmann, I see
no tangible differences and, although fruits are absent, I believe Psychotria
to be its proper generic designation. Apparently at one time Valeton
thought that Uragoga was represented, and later that the plant might be
a small form of Psychotria expansa Blume. Miquel’s older name is
here adopted.

ARNOLD ARBORETUM,
HARVARD UNIVERSITY.

Jour. ARNOLD Ars. VoL. XX XV PLATE I

SAURAUIA LANCEOLATA DC.
(Type, Leschenault 643)

158 JOURNAL OF THE ARNOLD ARBORETUM _ [VvoL. xxxv

STUDIES IN THE BORAGINACEAE, XXVII
SOME GENERAL OBSERVATIONS CONCERNING
THE LITHOSPERMEAE

Ivan M. JOHNSTON

IN THE PREVIOUS PAPERS Of this series the genera composing the tribe
Lithospermeae have been redescribed and individually discussed, Jour.
Arnold Arb. 34: 258-299 (1953) and 35: 1-81 (1954). The tribe having
been surveyed, we are now prepared to present a new key to the twenty-
three genera concerned, and also some general observations concerning the
tribe as a whole. Listed alphabetically the twenty-three genera of the
Lithospermeae are as follows: Alkanna, Ancistrocarya, Arnebia, Buglos-
soides, Cerinthe, Cystistemon (includes Vaupelia), Echioides, Echium,
Halacsya, *Lasiarrhenum, Lithodora, Lithospermum, Lobostemon, *Ma-
cromeria, Maharanga, Moltkia, *Nomosa, Onosma, *Onosmodium, *Perit-
tostema, Podonosma, *Psilolaemus, and Stenosolenium. Six of the twenty-
three genera are endemic to America (those marked with the asterisk),
and sixteen are endemic to the Old World. Only one genus, Lithospermum,
has species present in both the Old and the New World.

To be excluded from the Lithospermeae are a number of genera which
at one time or another have been referred to it. These include Moritzia,
Thaumatocaryon, Antiphytum, Amblynotopsis, Amphibologyne, Sericos-
toma, Echiochilon, Megastoma, Mvyosotis, Mertensia, Trigonotis, Pul-
monaria, Bothriospermum, Moltkiopsis, Mairetis and Neatostema. Of
these genera, Pulmonaria is best referred to the Anchuseae and Bothrio-
spermum to the Cynoglosseae, and the remaining fourteen to the Eri-
tricheae.

As to habit the Lithospermeae are prevailingly herbaceous perennials.
Species having only an annual duration are relatively few and occur only
in Buglossoides, Arnebia, Stenosolenium, Cystistemon (including Vau-
pelia), Echium, and Cerinthe. The cymes are abundantly bracted and
usually conspicuously so in all members of the tribe except Ancistrocarya.
Floral dimorphy in several forms is present in five genera within the tribe.
Heterostyly in a very elaborate form is present in Lithospermum,
Echioides, and Arnebia, and in a simple form in Lithodora. Elsewhere
among the Boraginoideae heterostyly is reported only among species of
Anchusa and Pulmonaria of the Anchuseae, among Mertensia, Crvptantha,
and Amsinckia in the Eritrichieae, and in one species of Paracaryum in
the Cynoglosseae. Gynodioecism is widespread within Echium. Outside
the Lithospermeae it is known only in a few species of Mvyosotis and
Lindelofia. Cleistogamy is developed in some species of Lithospermum and
elsewhere in the subfamily is known only in Neatostema, Cryptantha and
Pectocarya.

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVII 159

Deviations from radial symmetry in the flower are more common and
widespread in the Lithospermeae than in any other group within the
family. The corolla becomes distinctly zygomorphic in Echium, Lobo-
stemon, Halacsya, Alkanna, and Macromeria. In addition, zygomorphy
less evidently developed is also present in corollas of some species of
Cerinthe and Onosma. Deviations from radial symmetry in the androecium
occur in many Lithospermeae. Within the corolla the individual filaments
may differ from one another in length, or in the form of their attachment, or
in the height at which they are affixed to the corolla. Androecia deviating
from radial symmetry occur in Echium, Lobostemon, Alkanna, Moltkia,
Lithodora, Maharanga, Cerinthe, Echioides, Arnebia, Stenosolenium, and
Macromeria. Outside the Lithospermeae decidedly zygomorphic corollas
occur only in Echiochilon and Lycopsis. Differentiation among the mem-
bers of the androecium, within the corolla, occur outside the Lithospermeae
in Echiochilon, Moltkiopsis, Lycopsis, Caccinia (includes Heliocarya) and
Amsinckia.

The frequency of yellow or orange as a corolla-color among the Litho-
spermeae is also noteworthy. Orange, yellow, or decidedly yellowish co-
rollas occur in Moltkia, Halacsya, Alkanna, Onosma, Podonosma, Cerinthe,
Lithospermum, Echioides, Arnebia, Psilolaemus, Perittostema, Onosmo-
dium, and Macromeria. Within the tribe blue or bluish or pink corollas
occur in Echium, Lobostemon, Alkanna, Moltkia, Lithodora, Maharanga,
Onosma, Cystistemon, Podonosma, Cerinthe, Arnebia, Stenosolenium,
Buglossoides, and Ancistrocarya. White corollas are normal in species of
Lithospermum, Buglossoides, Lasiarrhenum, Nomosa, and Onosmodium.
Blue, pinkish, or white corollas predominate in the other tribes of the
Boraginoideae. Orange, yellow or yellowish corollas, however, are de-
veloped in other tribes by only a relatively few species of Nonnea, Sym-
phytum, Anchusa, Neatostema, Cryptantha, Hackelia, Amsinckia, Rindera,
Lindelofia, and Cynoglossum.

The corolla has an unappendaged throat in seventeen of the twenty-
three genera of the Lithospermeae. Within the tribe there are only six
genera in which intrusive faucal appendages are developed, viz. Litho-
spermum, Buglossoides, Macromeria, Perittostema, Halacsya and Alkanna.
In the three other tribes of the Boraginoideae, however, the corolla is
almost always provided with faucal appendages. Indeed, within the
Anchuseae, Eritrichieae and Cynoglosseae there are only eight genera, out
of more than sixty, in which the corolla is devoid of faucal appendages in
all or nearly all of the species, i.e., Moltkiopsis, Mairetis, Neatostema,
Echiochilon, Sericostoma, Amphibologyne, Amsinckia and Trichodesma.
In the other genera in these three tribes the faucal appendages are well
developed or are weak or absent only in a few species in which the corolla
is much reduced in size, e.g., Cryptantha, Plagiobothrys, and Pectocarya.

The corollas of the Lithospermeae, unlike those in other tribes of the
family, may bear stipitate glands on their inner surfaces, particularly so
those in and directly below the corolla-throat. Stiped glands, sometimes
in considerable abundance, are a feature inside the corollas of some or all

160 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXxXv

species of Lithospermum, Macromeria, Lasiarrhenum, Perittostema, Ar-
nebia, Buglossoides, Lithodora, and Alkanna.

The corolla-throat in Ancistrocarya and in one species of Lithodora is
densely strigose, a condition paralleled outside the tribes only in Seri-
costoma and Echiochilon. The corolla-tube (as distinguished from the
corolla-throat) is hairy in species of Lithospermum, Macromeria, Echioides,
and Arnebia. Outside of the Lithospermeae I know of a comparable con-
dition only in the flowers of species of V eatostema, Mertensia, Amsinckia,
and Rochelia.

The stamens in the Lithospermeae show a number of unusual features.
In Cerinthe, Podonosma, Cystistemon, Onosma, and Maharanga, the
anthers are connivent and become joined, in Cerinthe by the entangling of
the caudate bases of the thecae, and in the other genera by lateral cohesion
of the anthers. Outside of the Lithospermeae this condition is most closely
approximated in Borago and Trichodesma. In these two genera the
anthers are connivent, but in Borago they remain distinct, while in Tricho-
desma they become joined by the entangling of their contorted tips.

The anthers are terminated by an elongate sterile appendage in Cerinthe,
Podonosma, Onosma, Cystistemon, Maharanga, and Nomosa, Outside of
the present tribe only in Trichodesma of the Cynoglosseae does a com-
parable very elongate appendage terminate the anther, Small appendages
terminate the anther in Halacsya, Onosmodium, Lastarrhenum, and Bu-
glossoides. Outside the tribe I know of such moderately appendaged
anthers only in Myosotis and Rochelia.

In Lasiarrhenum, Nomosa, and one species of Macromeria the connec-
tive on the dorsum of the anther is hairy, and in Halacsya the margins
of the thecae are densely short-ciliolate. These conditions appear to be
unique in the family.

The filaments within the individual corolla are not whorled but affixed
at different elevations above the corolla-base in species of Lithodora,
Alkanna, Echium, Lobostemon, Macromeria, Stenosolenium, Arnebia, and
Echioides. This unusual behavior is duplicated outside of the Lithosper-
meae only in Echiochilon, Lycopsis, and Amsinckia. The five filaments
within the corolla are of different lengths in species of Arnebia, Moltkia,
Alkanna, Echium, and Lobostemon. Outside of this tribe I know of this
latter condition only in Moltkiopsis, Echiochilon, and Caccinia.

Although the pollen of the Boraginaceae, outside of the Lithospermeae,
has not been systematically studied, there have been enough random ex-
aminations to indicate that the pollen of the Lithospermeae is notable
for a high frequency of asymmetric types. Such pollen has the upper and
lower halves of the grain differing in size and configuration. In lateral
outline the grain may be ovate, conic-ovate, oblong-ovate, or be con-
stricted above the pores and approach the outline of an hour-glass or a
shoe-print. It is encircled by a row of pores, not about the equator, but
below it, where the grain has its maximum diameter. Pollen of this sort is
developed in Alkanna, Echium, Lobostemon, Macromeria, Onosmodium,
Nomosa, Lasiarrhenum, Ancistrocarya, Lithospermum, Podonosma, Cysti-

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVII__ 161

stemon, and Onosma. At present this type of pollen is known only in
genera of the Lithospermeae.

The style in the Lithospermeae may be short to elongate and be either
included in the corolla or exserted from it. In Arnebia it may be simple,
bilobed, forked, or bis-bifid. It is simply forked in Echium and Stenoso-
lenium and frequently bilobed in Lithodora and Alkanna,. The forked or
bilobed style is not duplicated in other tribes of the Boraginoideae. Each
lobe of the forked style is usually terminated by a stigma. When the style
is simple it commonly bears two stigmas juxtaposed and terminal on its
summit. In some genera, however, the two stigmas are separated and
frequently over-topped by a bilobed sterile apex of the stylar column. In
such cases the stigmas assume a subterminal position on the style and may
become evidently lateral on it.

The subterminal stigmas are best developed in species of Buglossoides
and Lithospermum. Unless the peculiar stigma of Myosotis proves to be
similar, subterminal stigmas are well developed in other tribes of the
Boraginoideae only in the genus Sericostoma. In most genera of the
Lithospermeae the style is terminated by a pair of juxtaposed stigmas.
Within the tribe the two stigmas usually remain distinct. In other tribes
of the subfamily they are prevailingly united.

The nutlets of the Lithospermeae are prevailingly straight, erect, and
basifixed, and in form are either ellipsoidal (and usually plump) or they
are ovoid with the venter tending to be angulate and the apex coarsely
beaked. Nutlets that are strongly compressed dorsi-ventrally, though fre-
quent in the Eritrichieae and Cynoglosseae, are rare in the Lithospermeae
and indeed are practically restricted to a few species of Arnebia. The
ventral keel in various degrees may be obscure, broad and rounded, or
narrow and prominent. There may be no evidence of a ventral suture or
the suture may be marked only by a line that may be indistinct or some-
times interrupted. The suture is always tightly closed with its margins
usually fused or firmly joined and never overlapping.

Several genera of the tribe have nutlets sufficiently unusual to be de-
serving of special comment. The 2-celled, 2-seeded nutlets of Cerinthe are
well known. They apparently represent the congenital union of a pair of
single-seeded nutlets. The condition is unique in the Boraginoideae. Also
unique is the form of the nutlets of Ancistrocarya. From just above the
broad base these gradually narrow into a very prolonged, slender, curved,
sword-like beak which is hamate at the apex. The beak, which is nearly
as long as the fertile portion of the nutlet, is a sterile prolongation of the
pericarp-apex.

The bent nutlets of Moltkia, Halacsya, Alkanna, Podonosma, and some
insular species of Echium represent another very unusual form of nutlet.
Outside the tribe it is weakly represented only in Neatostema and some
species of Plagiobothrys. In the Lithospermeae mentioned, the nutlets are
basally affixed to the gynobase but only the lower half of the nutlet is
erect. Above their middle they are bent 30-130° towards the ventral side.
The seed within the nutlet is also bent. The cotyledons are vertical to

162 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXXV

the attachment-end of the nutlet and the tip of the cotyledons is directly
above the nutlet-attachment, the proper relation in a basifixed nutlet. In
Alkanna and Podonosma the nutlet-body is so strongly curved that the
attachment actually has the appearance of being lateral.

The nutlets of the Boraginoideae characteristically have a more or less
convex dorsum and a usually angulate or medially carinate venter. An
extreme departure from this conventional form is found in the nutlets of
Arnebia tetrastigma. In that species, the sole member of Arnebia §
Euarnebia, the nutlet has a plane or even slightly concave dorsum and a
venter that is broadly convex with the ventral keel only very obscurely
developed. The nutlets are also aberrant in having a cordate base and a
T-shaped scar.

The nutlet of Stenosolenium bears its attachment-scar not sessile on the
base of the nutlet-body but rather ventral to the proper base of the nutlet
at the lower end of a downwardly directed hollow stipe originating high
up on the ventral side of the nutlet. In all other genera of the Litho-
spermeae the attachment is on the base of the nutlet-body. It is small,
short and substipitate in Alkanna and Podonosma but is relatively large,
sessile, and commonly flabellate or ovate in other genera, The attachment
is usually restricted to the base of the nutlet. In Arnebia guttata and
A. tetrastigma, however, it has a noticeable prolongation upwards for a
short distance above the base on the nutlet-venter, As a consequence it
becomes somewhat T-shaped, especially in A. tetrastigma.

The attachment-scar is usually not only basal on the nutlet but also
horizontal. When oblique it usually slopes upward only towards the ven-
tral side of the nutlet-body and commonly only slightly so. A basal at-
tachment-scar sloping upward, not towards the venter of the nutlet but
rather towards the dorsum, is present only in Buglossoides § Eubuglos-
soides and perhaps in Ancistrocarya. The attachment of the nutlets in
Lithodora is anomalous. The true base of the nutlet is permanently affixed
to the gynobase, the nutlet being freed along a new plane of abscission
developed a short distance above the morphological base of the nutlet.

The nutlets of the Lithospermeae may be smooth, verrucose, tumulose,
or rugose, and the epidermis lustrous or opaque and smooth and shiny
or minutely muriculate, verruculose, or papillate. The smooth, lustrous,
pallid, porcelain-like nutlets characteristic of most species of Lithosper-
mum occur elsewhere in the tribe only in Macromeria, Onosmodium. Las-
iarrhenum, Psilolaemus, Ancistrocarya, and Buglossoides § Margarosper-
mum. Roughened or at least opaque nutlets prevail in other genera o
this group. In other tribes of the Boraginoideae the nutlets may be armed
with numerous glochidiate subulate appendages, or the back of the nutlet
may bear a coroniform or annulate appendage, or it may be encircled by
an entire, toothed or lobed, spreading or upturned margin. Among the
genera of the Lithospermeae, however, only a few species, in Onosma,
Echium, and Ancistrocarya, have appendaged nutlets. Those of Onosma
tricerospermum Lag., of Spain, bear three conspicuous spreading rigid
subulate spines on the dorsum. The nutlets of all other species of Onosma

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVII 163

are unappendaged. In some of the Canary Island species of Echium (cf.
Webb. & Berth., Phytogr. Canar. t. 146, 1844), the pericarp develops a
few very exaggerated protuberances that give the nutlet the appearance
of being coarsely lobed. The slenderly rostrate nutlets of Ancistrocarya
have already been described. The nutlets of the Lithospermeae (like those
of the Anchuseae) never bear glochidiate appendages nor epidorsal annu-
late or coroniform crests, nor is the back ever encircled by a differentiated
margin.

My observations as to the orientation of the zygomorphic corollas in
the Boraginoideae do not accord with the much quoted classical statements
of the matter by Doll, Fl. Baden 2: 775-6, 778 (1859) and Eichler,
Blithendiagramme 1: 197 (1875). The calyx lobes of the boraginaceous
flower are numbered by Eichler in the following order, 1, 3, 5, 2, and 4.
Lobe no. 2 is identified as the one at the rear of the flower and as having
a position opposite the rachis of the scorpioid cyme. This order appears
to be correct, for in those flowers having very unequal calyx-lobes (e.g.,
Cerinthe) the largest outermost lobe, i.e., no. 1 according to Eichler, is
always one of the two lobes on the shawl or front side of the flower.
With the calyx-lobes recognized as serviceable points of reference, it can
be stated that according to Eichler the axis of the irregular corolla of
Echium passes between the corolla-lobes alternating with calyx-lobe no.
4 and through the middle of the corolla-lobe alternating with calyx-lobes
nos. 3 and 5. This gives the flowers of Echium a very strongly oblique
plane of symmetry and places the 2-lobed lip of the corolla on the adaxial
side of the flower.

In my study of herbarium material, I have found in Echium, Lobo-
stemon, Halacsya, Alkanna, Lycopsis, and Amsinckia, that the axis in the
bilaterally symmetrical corolla is oriented in such a manner that the
2-lobed lip of the corolla is abaxial and the 3-lobed lip adaxial. In these
genera the plane of symmetry is only weakly oblique. It passes through
the middle of the rear corolla-lobe (that alternating with calyx-lobes nos.
4 and 2) and between the corolla-lobes alternating with calyx-lobe no. 3.
This is the same orientation that prevails in the zygomorphic corollas of
the Solanaceae, It represents a deviation of only 36° from a truly medial
orientation, not 72° as called for by Eichler.

Only in Macromeria, Echiochilon, and Heliocarya have I found in the
Boraginaceae zygomorphic corollas with medial orientation, that is to say,
so oriented that the rear (adaxial) lip was 2-lobed and the abaxial lip
3-lobed. In the flowers of these genera the axis apparently passes between
the two rear corolla-lobes (those alternating with calyx-lobe no. 2) and
also through the middle of the forward corolla-lobe, that alternating with
calyx-lobes nos. 1 and 3. Medial orientation similar to this prevails in the
corollas of the Verbenaceae and Labiatae. It represents an angular devia-
tion of 36° from the axis of symmetry in Echium

Enough evidence has accumulated from the study of herbarium specimens
to cast considerable doubt on the accuracy of accounts in the books con-
cerning the orientation of the bilaterally symmetric flowers of the Bora-

164 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

ginoideae. What is now needed is observations of flowers in the fresh state.
The orientation should be determined not merely in the flowers having
evidently zygomorphic corollas, but also in those in which the corolla may
be only very obscurely bilateral or which depart from radial symmetry
only in the androecium. Studies of symmetry are especially needed in
Lithodora, Moltkia, Stenosolenium, Arnebia, Echioides, Cerinthe, Onosma,
Podonosma, and M oltkiopsis. The orientation in the corolla of these
genera needs to be determined not only in relation to the axis of the cyme,
but also in relation to calyx-lobe no. 1, and particularly so when the latter
is distinguishable by its size.

KEY TO THE GENERA OF THE LITHOSPERMEAE

Pollen bearing only 3 pores, grains in polar profile usually distinctly 3-sided
Pollen-grains evidently colpate; thecae not ciliate; flowers not precociously

Nutlets strongly incurved, near the middle bent 90-130°, the attachment
small and substipitate, appearing to be lateral but actually basal on the
short erect lower section of the nutlet; herbage usually glanduliferous.

Corolla-lobes narrowly triangular, acute, soon reflexed: corolla-throat
glabrous and devoid of faucal appendages and stipitate glands;
anthers partially exserted. ........................, odonosma.

Corolla-lobes rounded, spreading; corolla-throat usually bearing stipitate
glands; stamens and faucal appendages borne low in the corolla-throat

ence deeply included. ....................... Alkanna.

Nutlets straight (or rarely bent in Echium); herbage not glanduliferous.

Anther lacking a prolonged sterile tip, remaining distinct; corolla usually
evidently zygomorphic.

Annulus borne 1 mm. or less above the base of corolla-tube, composed
of a minute collar or a ring of 5-10 minute sparingly hairy lobules ;
style almost always 2-lobed. ......................

Annulus developed 1.5-6 mm. above the corolla-base, represented by
5 evident densely villous swellings or 5 squam ose appendages
borne below the attachment of each stamen; style simpl
PEMA T ES Re Rede eee RI IAG ¢ dnd ank REM a aes tle £44 Lo bostemon.

Anther narrowed into a prolonged terminal appendage, commonly adnate
at the base or along the sides to form a synandrium; corolla regular
or practica

Corolla-lobes well developed, as long as or much longer than the corolla-

than the theca; filaments very short and usually bearing a
thickened hairy basal appendage; tropical Arabia and Africa. .

Ab Rae hans OPE Ete hd oA Cystistemon and Vaupelia.

cae lobes short, commonly about as long as broad, conspicuously

uch shorter than the tubular portion of the corolla: anthers

male coherent at the base and frequently also along the margin

of the thecae and even along the margin of the terminal appendage,

appendage usually shorter than - theca; filaments usually

elongate, not appendaged at the bas
Calyx-lobes narrow, elongate, more or less parallel, separated by a

1954] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXVII 165

very narrow usually closed sinus; corolla without ribs projecting
outward between the calyx-lobes; pollen ovoid to sphaeric or
strongly oblate Onosma.
Calyx-lobes more or less triangular, separated by an open triangular
sinus; corolla with puffed-out ribs that project outward between
the calyx- lobes; pollen cylindric or vertically ellipsoid. Himalaya
and southwestern China ...................... Maharanga.
Pollen-grains not at all colpate; anthers with thecae margined by a ciliate
fringe of short stout crowded hairs; flowers precociously sexual, style and
stamens exserted while the corolla is still incompletely developed: nutlets
OIE: aie eee eae o etal ee 465k. 3. 4s EO Bee Halacsya.
ae bearing 6 to many pores; grains circular or somewhat polygonal in polar

e.
Nutlets united in pairs, each nutlet 2-locular - 2-seeded; pollen ana
Ipate; leaves cordate-amplexicaul at base ................
Nutlets ne ‘united, each one 1-celled and 1- oe pollen obscurely if 2 : all
; leaves not cordate-amplexicaul.
Pole ee bearing 2 rows of pores, one at each end of the elongate grain.
Nutlet-attachment at the base of a downwardly directed stipe originating
on the ventral side of the ascending nutlet-body ... Stenosolenium.
Nutlet-attachment on the base of the nutlet-body Arnebia
Pollen bearing only a single encircling row of pores.
utlets circumscissile above the base, their major seminiferous portion
falling away leaving the short basal section persisting as a usually
cupu is te appendage permanently affixed to the gynobase . . Lithodora.
Nutlets detaching at the very base
Throat of corolla glabrous and also devoid of stiped glands and faucal

ppendages.

Nutlets evidently bent at or near the middle; corolla blue or purple,
or (in one species) yellow, filament evidently eee from
corolla-throat except in one species Moltki

Nutlets straight.

Corolla ee the broad spreading yellow limb usually
d ted with 5 evanescent black or purplish spots; stamens
. at unequal — in the corolla-tube; Armenia,
Caucasus and Azerbaijan .................... Echioides.

Corolla tubular, white or oe bearing no dark evanescent spots,
lobes erect or ascending; stamens all arising at the same height
in the corolla-throat ; American:

Filaments about half the total length of the corolla, arising well
below the middle of the corolla, base of filaments ren
with slender multicellular gland- tipped hairs Nom

Filaments about one tenth the length of the corolla or ieee
arising above the middle of the corolla, glabrous sigh

Corolla-lobes narrow, acute or acuminate, very owly
imbricate in the bud, sinus between lobes nee pir i!

a oe at base; pollen ovoid; leaves coarsely

TINCT ons 5 scan 028 Onosmodium.
Corolla- lobes ovate, broadly imbricate in the bud, sinus be-
tween lobes not plicate nor inflexed nor thickened at base;
pollen ellipsoidal; leaves veinless or nearly so

166 JOURNAL OF THE ARNOLD ARBORETUM _ [vot. xxxv

Throat of corolla decorated with appendages, stipitate glands or abun-

ant hairs.
Nutlets gradually narrowed into a very prolonged beak, conspicuously
rostrate with the tip hamate; cymes bractless above e
base; corolla-throat abundantly villose-strigose .. Ancistrocarya.
Nutlets not conspicuously rostrate, not hamate at the apex; cymes
bracted throughout; corolla-throat not conspicuously hairy.
Filaments broadening upwards from the aia above the
middle evidently broader than at the bas
Anthers conspicuously hairy on the back; “ee oblanceolate;
corolla lacking faucal appendages; leaves broad and con-
spicuously veined ...................... Lasiarrhenum.
Anthers glabrous, filaments deltoid to obovoid; corolla with
faucal appendages; leaves very narrow, veinless ..........

Filaments linear or subulate or unguiculate.
Anthers completely exserted from the corolla-throat; corolla
ree, 39-90 mm. long, trumpet-shaped; filaments 6-70 mm
QUE 5.55645 coe nein na cnet iseuee eid Macromeria.
SS completely included or only partially exserted from the
orolla-throat; corolla usually less than 40 mm. long,
oe. funnelform or subtubular; filaments 0.5-5 mm.
lon

ong.
Corolla-throat decorated inside with 5 well-developed shale
lines of hairs and stiped-glands ........ Buglossoides.

Corolla-throat lacking very well developed vertical (es of
hairs and stiped glands, bearing localized faucal append-
ages or bearing scattered or localized ey toons of
stiped glands... ospermum.

ARNOLD ARBORETUM,
ARVARD UNIVERSITY.

1954] PAULEY & PERRY, POPULUS 167

ECOTYPIC VARIATION OF THE PHOTOPERIODIC
RESPONSE IN POPULUS

Scott S. PAULEY ! AND THoMaAs O. PERRY ”
With eight text-figures

INTRODUCTION

DURING THE PAST FEW DECADES, the traditional and complacent notion
that genetic differences within tree species are nonexistent or negligible has
been gradually discarded. Most silviculturists are no longer content with
the assumption that diversity within tree species can be adequately ex-
plained in terms of environment alone. Such a fundamental revision in
biological theory has, quite naturally, produced a marked impact on
silvical research methods and silvicultural practice.

The limitations, for example, of the descriptive method as a means of
analyzing intraspecific diversity have become increasingly apparent. Al-
though field observation and description of phenotypes and environment
provide useful and essential preliminary data for the analysis of observed
diversity, the method provides no means for determining the relative
amounts of environmental fluctuation and genetic variation involved.

The solution of such silvical problems is, however, possible by transplant
methods that have been perfected by various workers (Turesson, 19;
Turrill, 20; Clausen, Keck and Heisey, 2; and others). One application
involves the transplantation of different genetic types into a uniform
environment, Under such conditions the effects of differences in habitat on
the phenotype are eliminated, and the genotypes of different individuals of
the group under investigation may be compared directly. For information
on the rigidity or plasticity of genotypes, wild representatives of the species
may be propagated vegetatively, and the clonal lines thus established may
be planted under different environmental conditions, Both methods have
been used in the photoperiodic studies here reported, and form one of the
approaches to silvical problems currently under investigation by the Cabot
Foundation.

Source of Material

Initial sampling of wild populations in Populus species native to North
America was begun in the fall and winter of 1947-48, and has been con-
tinued annually since that time. During the period 1947-50 principal em-
phasis was placed on obtaining representatives of P. trichocarpa (western
balsam poplar or so-called “northern black cottonwood”’), native of the

2 Lecturer on Forest Genetics and Geneticist to the Maria Moors Cabot Foundation
for Botanical Research, Harvard University, Petersham, Mass.
2 Assistant Professor, School of Forestry, University of Florida, Gainesville, Florida.

168 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXXv

Pacific coast region, and the P. deltoides complex (eastern cottonwood) of
the eastern United States. These collections were all in the form of branch
or stem cuttings taken from what were determined to be average, healthy
specimens growing under natural conditions at the various collection points.
The use of cuttings was designed to insure that the clonal lines represented
were derived from individuals known to be capable of survival in their
native habitat. Since species in the section TACAMAHACA (balsam poplars)
are uniformly good rooters from plants of any age, there was no need to
restrict selected individuals of P. trichocarpa to the younger age classes.
On the other hand, because of the poor rootability of cuttings from old
trees, almost all of the P. deltoides clonal lines established were started
from cuttings taken from plants 2 to 6 years of age.

The photoperiodic studies here reported are primarily concerned with
clones of P. trichocarpa and the P. deltoides complex collected during the
period 1947-50; but reference is also made to clonal line collections of
P. tacamahaca (balsam poplar) made during the same period, and to col-
lections of various other Populus species acquired in the spring of 1950 and
subsequently.

Approximate ranges of P. trichocarpa and P. deltoides and the locality
of origin for the various clonal lines of these species used in the photo-
periodic studies are shown in Ficure 1.

Although a portion of the field collection was done by the senior author,
the bulk of the material was made available through the generous assistance
and cooperation provided by numerous state and federal forestry agencies
in the United States, and similar provincial and dominion organizations in
Canada. Especially extensive contributions have been made by the U. S.
Forest Service, U. S. Soil Conservation Service, Dominion Forest Service,
and many state and provincial conservation departments within the ranges
of the species concerned.

Experimental Areas

The principal propagation and test plantation site utilized by the Cabot
Foundation is located in the town of Weston, Middlesex County, Massa-
chusetts, about 10 miles west of Boston. The area is part of the former
Case Estate, given to the Arnold Arboretum about a decade ago. The
portion of several acres which has been allocated to the Cabot Foundation
was formerly used as a pasture or mowing. The soil is, in general, uniform
and the surface flat. The elevation is 200 feet, and lies somewhat lower
than most of the surrounding area. For this reason temperature inversions
in early fall may result in a somewhat shorter frost-free season than the
average of 148 days for the Weston area as based on records of the Weston
Weather Bureau Station (located ca. 2 miles northwest of the Case Estate
at an elevation of 224 feet).

A portion of the Weston tract has been devoted to the propagation of the
clonal lines assembled since the 1947—48 season, while the remainder of the
area has been set aside for the establishment of permanent test plantations.

1954] PAULEY & PERRY, POPULUS 169

NORTH AMERICA

COLLECTION POINTS
Legend:
Approximate range boundary,
Pet &

+ trichocarp ; ee “I 2
Sy ? :
Approximate range boundary, les
10] ae P, deltoides

i ss e fe
© Collection point (5 - 10 clones)

|
\
|
120 10

.
tet Lomorruoe

Fic. 1. Approximate ranges of P. trichocarpa and P. deltoides and collection
points for clones used in this study.

170 JOURNAL OF THE ARNOLD ARBORETUM _ [VvoL. xxxv

Although two such plantations were established in 1950 and 1951, observa-
tions at Weston on the date of cessation of height growth for the various
clones concerned in this study were all made in the propagation area, here-
after referred to simply as the ‘“‘Weston area.”

The combined experimental facilities of the Arnold Arboretum and the
Bussey Institution, located in Jamaica Plain, Massachusetts, a part of
greater Boston (ca. 10 miles southeast of Weston), have also been used in
this study. The outdoor long-day and normal-day plots used in the 1951
experiments were set out in a portion of the experimental plots used jointly
by these institutions, and located on a flat gravelly knoll in the first range
of hills south of the Boston basin. The soil, thanks to continuous enrich-
ment and good management, is highly fertile and uniform.

PHOTOPERIODIC RESPONSES IN PLANTS

Biologists, as well as most primitive peoples, have long recognized the
indispensable nature of solar energy in the biological scheme. But only
within the past thirty years have botanists recognized the notable signifi-
cance to plant growth and development that is associated with the periodic
manner in which this energy is supplied to the earth’s surface.

Within the tropical and subtropical regions of the world, there is little
change in the length of the daylight period throughout the year; but outside
these regions, marked seasonal differences occur. At the latitude of Boston
(ca. 42° N.), for example, the length of the daylight period on the shortest
day of the year is ca. 9 hours, and on the longest day ca. 15 hours, i.e., a
difference of 6 hours (Fic. 2). With increases in latitude the annual varia-
tion in day-length is greater. Thus, at the latitude of Juneau, Alaska (ca.
58° N.), day length at the time of the winter solstice is ca. 6.5 hours as
compared with ca. 18.5 hours on the day of the summer solstice, a differ-
ence of 12 hours. The above day lengths are based on the time of sunrise
and sunset. Actually, the effective photoperiods at all latitudes are some-
what greater due to the effects of the morning and evening twilight periods
(Fig. 2).

Flowering Response

The response in growth and development exhibited by plants in relation
to the length of the daily light period (i.e., photoperiod) is called photo-
periodism. The earlier and, indeed, most of the later investigations con-
cerned with this phenomenon have centered on the flowering response of
various herbaceous plant species and varieties when exposed to photoperiods
of varying length.

First recognition of the influence of day length on reproductive devel-
opment in plants was made by Garner and Allard (3) in 1920. These
workers observed that Maryland Mammoth tobacco, which normally does
not flower in the field during the summer season at the latitude of Wash-
ington, D. C. did, nevertheless, flower profusely when grown in a green-

PAULEY & PERRY, POPULUS 171

1954]

t t

W3G045C) waONI1d3C asnony
—t —-

Av Wey Hwy huvowea 4 Awan

Bavo i

T Ly T T

$M1d HLONBIAVG

——— on HLONDIAVO
SHOAL

Sd HLINTIAPG

HLIONBIAVG

HLONTIAVG
NI NOLLVINWA TYNOSVaS

Fic. 2. Seasonal variation in day length at various latitudes: equator, 42° 20’

N. and 58° 20’ N.

172 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxv

house at the same latitude during the winter. The dissimilar reproductive
development observed led to the hypothesis that day length was a con-
trolling environmental factor, the relatively short photoperiods of winter
apparently favoring flower production in this variety. Subsequent con-
firmation of this hypothesis was made through the use of carefully con-
trolled experiments by these and numerous other workers.

The significance of the photoperiodic influence on flowering is obviously
a matter of much practical concern in the case of cultivated crops grown
for their fruit or seeds. Many of the grains, for example, require long days
to flower, and for this reason long-day varieties cannot be successfully
moved to southern latitudes. Other long-day plants grown for the food
stored in their vegetative organs, such as spinach, lettuce, and radish,
must be grown as spring or autumn crops in regions where the long days of
summer would otherwise cause them to “bolt.”

In order for such short-day plants as the Biloxi soy bean and numerous
ornamentals (cosmos, aster, chrysanthemum, etc.) to flower, they must be
grown either in the low latitudes or in the short days of spring or autumn
in the higher latitudes. Many short-day ornamentals native of the tropics
are, of course, grown successfully in greenhouses under the short-day
regime of the winter months in northern latitudes.

Plants indeterminate with respect to their photoperiodic requirements
for flowering, such as tomato, cotton, buckwheat, and sunflower, may be
grown successfully over a wide range of latitudes.

Vegetative Response

Although major interest in the photoperiodic studies involving herbaceous
plants has centered on the influence of day length upon reproductive de-
velopment, most workers have also noted marked effects on vegetative
growth as well. Most plants exhibit much better vegetative development
when exposed to long photoperiods than to short ones, regardless of the
fact that they may be classified as “‘short-” or “long-day” types with respect
to flowering. Maryland Mammoth tobacco, for example, when grown
under the long days of summer at the latitude of Washington, D. C., by
Garner and Allard (3), attained heights of 10-15 feet; whereas, when
greenhouse grown in the short days of winter, it did not exceed five feet.

Conversely, movement of long- or short-day flowering types into natural
or artificially created shorter days usually results in marked reduction in
vegetative development

Photoperiodic studies of woody plants have for the most part involved
seedlings, and consequently little is known of the influence of day length
on reproductiveness. These investigations have, however, demonstrated
that variation in day length is a factor of marked importance in the annual
vegetative growth cycle of many woody species, particularly as it relates
to the control exerted over the timing of physiological processes concerned
in the onset of dormancy.

In common with techniques used in the investigation of photoperiodic

1954] PAULEY & PERRY, POPULUS 173

responses in herbs, studies of woody plants have involved the testing of
various species in a day-length regime different from that prevailing in their
native habitat. Whether such tests have involved actual movement of the
plants north or south into a new natural day-length pattern, or if short or
long days have been artificially created at any particular latitude, the re-
sults have been the same. In general, movement from the latitude of the
natural habitat northward (i.e., into longer days) prolongs the active
period of growth; and movement southward (i.e., into shorter days)
shortens it. Such modifications in the length of the active growth period
normally result in marked differences in total seasonal increment and frost
hardiness. Thus, movement of plants into a day-length regime longer than
that of the native habitat characteristically gives increased height growth
accompanied by decreased resistance to early autumnal frosts; whereas
movement into a short-day regime results in dwarfing, associated with in-
creased frost resistance.

Similar photoperiodic growth responses have been, with but few excep-
tions, demonstrated by Moshkov (12, 13, 14) and Bogdanov (1) to charac-
terize the behavior of various species in the following genera: Acer,
Aesculus, Ailanthus, Alnus, Caragana, Corylus, Fraxinus, Juglans, Larix,
Phellodendron, Pinus, Prunus, Rhus, Robinia, and Ulmus; by Kramer (8)
and Jester and Kramer (6) in North American species of the following
genera: Acer, Fraxinus, Fagus, Robinia, Liriodendron, Liquidambar,
Quercus, and Pinus; by Sylvén (18) in the European aspen (P. tremula) ;
by Johnsson (7) in Betula; by Langlet (9) in Scotch pine; and by Olmsted
(15) in sugar maple.

INFLUENCE OF PHOTOPERIOD ON TIME OF HEIGHT GROWTH
CESSATION IN POPULUS

First suggestions of photoperiodic sensitivity among the clones in the
propagation area at Weston appeared during the first (1948) growing
season. Asa result of early height growth cessation and consequent dwarf-
ing, P. trichocarpa ecotypes from high latitudes stood out in sharp contrast
to those of more southerly origin.

Interclonal Responses to a Uniform Day Length

Precise measurements of height increment by the use of auxanometers of
various design have been made by several workers. By their use the time of
terminal growth cessation could doubtless be pinned down to a certain hour,
or at least to a particular day. Such methods are not, however, practical
for the measurement of large numbers of individuals in the field, since the
cost would be prohibitive.

In the present studies total height measurements were taken to the
nearest centimeter at 7-day, 5-day or 3-day intervals by direct measure-
ment with meter sticks or tapes. Although lacking a high degree of pre-
cision, the technique is sufficiently accurate for determining comparative

174 JOURNAL OF THE ARNOLD ARBORETUM _ [vot. xxxv

differences in growth cessation which are spread over a period of several
weeks or months. The error occurs during the period immediately preced-
ing the time of actual height growth cessation. The increments between
measurements during this period may be so small that the actual day of
growth cessation is indeterminable. Depending upon the interval between
measurements and other variables, deviations in the date of recorded
growth cessation probably vary in the order of 3 to 7 days before and after
the actual date of growth cessation. In general this method tends to en-
courage a later recording of growth cessation.

First systematic recording of the time of height growth cessation in the
Populus collections was made during the 1950 season in the propagation
area at Weston. Relatively small samples of tacamahaca and deltoides
clones were included in the study, the majority being trichocarpa. All were
derived from cuttings which had been planted in the spring of 1948 or 1949
and were thus in their second or third growing season. Measurements of
total height for each of the clones included in the study were taken at
weekly intervals from June 27 to November 7

Active terminal growth in the group of tacamahaca clones measured was
observed to occur over a period of about 158 days (April 14 to Septem-
ber 19) ; in the deltoides clones over a period of about 178 days (April 20 to
October 15); and in the trichocarpa group over a period of about 197 days
(April 14 to October 28). Although the first killing frost of autumn at
Weston occurred the night of September 24-25, the growing season was
by no means ended. Plants still in active height growth at that time
(deltoides and trichocarpa of southern origin) suffered some damage to the
young unfolding leaves near the growing point, but continued in active
height growth thereafter. Subsequent light frosts in October (on the 6th,
8th, 16th, and 21st) caused similar damage; but height growth persisted
in some clones until the first heavy frosts of October 26 and 27. These
frosts were apparently of sufficient severity to kill all meristematic tissue
in the growing points of the plants still active at that time.

The recorded dates of height growth cessation for the various taca-
mahaca clonal lines ranged from about June 20! to September 19, a period
of 91 days, representing 58 percent of the growing season. Among the
deltoides group, cessation of height growth occurred between August 15
and October 18, a period of 64 days, or 31 percent of the growing season.
The time of height growth cessation in the trichocarpa clones was consid-
erably more widespread: first terminal growth cessation was recorded
about June 20,1 and the last on October 28, a period of 130 days, or about
two-thirds of the growing season.

Since this considerable intraspecific diversity in time of terminal growth
cessation occurred under the essentially uniform environment prevailing in
the propagation area at Weston, the conclusion is inevitable that the vari-

Measurements were started on June 27. This date proved too late to record the
cessation of height growth of certain tacamahaca and trichocarpa clones of high latitude
origin. Subsequent observations in 1951, 1952, and 1953 have confirmed that height
growth in these clones stops ca. June 2Q

1954] PAULEY & PERRY, POPULUS 175

ous clones within the species observed are genetically diverse in this char-
acter. That this diversity is not randomly distributed throughout the range
of each species, but is inversely correlated with latitudinal distribution, is
revealed by the scatter diagrams that result when latitude of origin for
each clone of each species is plotted over the corresponding date on which
height growth ceased at Weston (Fics. 3, 4 and 5). The correlation co-
efficients for these diagrams are all high (r = —.893, —.706, and —.823
respectively), and all exceed the 1 percent level of significance.

P. TACAMAHACA
bore a Lat. of origin / date of
+ Ht. growth cessation
Weston, Mass., 1950
e
55
50 = . .
. e
5 45
oH e-
bs
|
4Oo
35
r-e 0893
30 } | { ! ! |
June July Aug Sept Oct
DATE
Fic. 3.

The trend of the gradient revealed by these diagrams clearly indicates
a tendency for clones of high latitude (long-day) origin to cease terminal
growth early, and clones of low latitude (short-day) origin to stop terminal
growth late when grown under the mid-latitude day-length regime prevail-
ing at Weston. These results are in agreement with the photoperiodic
growth reactions which characterize the behavior of European aspen
(Sylvén, 18) and species in other genera when subjected to day-length
regimes differing from those in their native habitats. As previously pointed
out, various workers have demonstrated that the movement of plants from
native habitats of long day into short day is normally characterized by
earlier height growth cessation; whereas movement from short day into
long day results in delayed growth cessation. As a working hypothesis it
therefore appears tenable to conclude that the inverse correlation between
latitude of origin and time of terminal growth cessation for the ecotypes

176 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxv
of the Populus species here concerned is the result of similar photoperiodic
reactions between the various genotypes of the ecotypes represented and
the uniform day-length regime to which each was exposed at Weston.

P. DELTOIDES
60
Lat. of origin / date of
Ht. growth cessation
oe Weston, Mass., 1950
50 F-
fof :
Hw e
& °
Re «=
40 + e:*
t
. «7 *%
355 *s
“. 7 : r=- 0706
30 N ! t es 4 i
June July Aug Sept Oct
DATE
Fic. 4.

Although the clinal nature of the variation is sufficiently well-defined in
these diagrams to support the above hypothesis, there is ample evidence to
suggest that the relationship between time of height growth cessation at
Weston and latitude of origin of the various clones is not necessarily simple
and direct, The spread in dates of growth cessation from mid-July to late
October among the trichocarpa clones native of latitudes between 40° and
50° (Fic. 5), for example, indicates that a pronounced local diversity in
photoperiodic response exists, apparently quite independent of latitudinal
origin.

That diversity in photoperiodic response may occur locally within a
species population under essentially uniform day-length conditions leads to
interesting speculation. There can be little doubt, in the first place, that
height growth cessation in Populus signals the onset of initial physiological
processes culminating in winter dormancy. The time during the growing
season at which terminal growth cessation occurs thus assumes a role of
critical survival value. Through the selective pressure exerted by the first
killing frosts of autumn, only those genotypes capable of terminating
height growth at a sufficiently early date to escape such frosts are capable
of survival. Within any uniform day-length zone, therefore, where the
growing season varies considerably in length, due to topography or other
factors, the hypothesis may be made that adaptation to any particular

1954 | PAULEY & PERRY, POPULUS 177
length of growing season is effected through the selection of those geno-
types having a suitable photoperiodic response to the prevailing day-length
regime of that latitude.

P. TRICHOCARPA
60 ab Lat. of origin / date of
e . ° Ht. growth cessation
a . ny e
Weston, Mass., 1950
55 .
°
50 i e
e oe :° e
? - ts te
B 45 e : oh 4 t + 7 $
| - °
e e
J ‘
3
3
rs=e 823
30 I ! | ] ] een
June July Aug Sept Oct Nov
DATE
Fic. 5.

The group of clones originating between 40° and 50° (Fic. 5), referred
to above, represents a case in point. This group is actually composed of a
broad longitudinal sampling of trichocarpa ecotypes, extending from the
Pacific coast to western Montana, and with a vertical dispersion from
near sea level to ca. 5000 feet elevation. Since the length of the frost-free
season in this latitudinal zone is, in fact, known to vary widely within
comparatively short distances, due to elevation, the nature of the topog-
raphy, etc., the considerable diversity in time of height growth cessation
for clones native of the zone might be satisfactorily explained on the basis
of the above hypothesis.

A direct test of the hypothesis may, of course, be made by determining
the degree of correlation, if any, that may exist between the lengths of the
growing season at the points of origin for clones native of a narrower
latitudinal band (i.e., having an essentially uniform seasonal day-length
regime) and the dates of height growth cessation for the same clones as
recorded at Weston.

Although precise data on the average length of the growing season for
the native habitats of each of the clones represented in our collections is
for obvious reasons unavailable, reliable estimates nevertheless can be

178 JOURNAL OF THE ARNOLD ARBORETUM _ [vot. xxxv

made for those native to areas in the vicinity of U.S. Weather Bureau
Stations (21).

FicuRE 6 shows the scatter diagram resulting when the length of
the average growing season for those clones native only of the 2° lati-
tudinal zone (45° to 47°) is plotted over the date of height growth ces-
sation at Weston. The regression path here defined appears sufficiently
uniform in trend (r = +.7891) to confirm further the hypothesis that
trichocarpa populations are capable of adaptation to growing seasons of
varying length within uniform day-length zones by selection of those
types having a suitable photoperiodic response.

P. TRICHOCARPA .
Av. Growing Season of origin /
Date of Ht. Growth Cessation
Clones from 45° - 47° N. Lat.
200 |- Weston, Mass., 1950
os
3
7° s .
J
z
3
vw) o
a
”
4 S
F 150
° *
o
3 . °
§ a
<
100 }
r = + .789
80 i ! +.
July August Sept Oct
DATE
Fic. 6.

Intraclonal Responses to Diverse Day-length Patterns

During the 1950 and 1951 seasons, tests of photoperiodic sensitivity as
revealed by influence on time of height growth cessation were made in about
100 clonal lines within the following species: P. trichocarpa, P. tacamahaca,
P. angustifolia, P. deltoides, and various hybrids of P. tremuloides « P.
tremula. The tests of 1950 were made in a greenhouse at the Arnold
Arboretum with potted propagules of 45 clonal lines grown under short-,
normal- and long-day photoperiods. Results of these observations have
been previously published (Hoffmann, 4).

* Exceeds the 1 percent level of significance.

1954] PAULEY & PERRY, POPULUS 179

Similar tests of the same clonal lines, and about 55 additional, were
made outside during the 1951 season in portions of the Bussey Institution
nursery. Ramets of the various clones in these studies were exposed only
to the natural-day pattern prevailing at the latitude of Boston (ca. 42° 20’
N.) and a long-day regime corresponding to that at Juneau, Alaska
(ca. 58° 20’N.).

Flood lights and reflectors for the long-day plot were so arranged as to
obtain complete coverage of the area with a minimum ground surface light
intensity of 1.5 foot candles. Since the work of Withrow and Benedict
(22), Matzke (10), and others demonstrates that light intensities of less
than 1 foot candle are capable of producing photoperiodic responses, twice
the duration of civil twilight was added to the ‘“‘sunrise to sunset” day
length (Fic. 2). According to Humphreys (5), light intensity at the be-
ginning of civil twilight (morning sun 6° below the horizon) is 0.4 foot
candle with a clear sky, and quickly rises above 1 foot candle.

Supplemental light was supplied the long-day plot by means of an auto-
matic time switch in the morning and evening. Adjustment of the switch
settings was made every third day to conform to the natural day at
Juneau. Use of the lights was discontinued on September 30, when the
photoperiod of Boston became greater than that of Juneau. Thereafter
both experimental plots received the normal day length of Boston.

The variable responses in terms of height growth cessation to different
day-length regimes that were observed added further confirmation of the
presumed photoperiodic sensitivity in Populus, and also demonstrated the
apparent similarity of this response to that exhibited by other tree genera
of the temperate zone.

Detailed tabulation of the data recorded in these studies cannot, in the
interest of brevity, be included in the present paper. However, a sample
of the records on height growth cessation for P. trichocarpa clones of
various origins when grown under the normal Boston-day and the artificial
long-day regimes at the Arnold Arboretum during the 1951 season are
shown in TaBLE 1 (Columns 7 and 8). Differences in time of growth
cessation are noted in Column 9. Included also are recorded dates of height
growth cessation of the same clones under normal-day conditions at
Weston in 1950 and 1951 (Columns 5 and 6).

As these data indicate, diversity of intra-clonal response to normal- and
long-day regimes was most pronounced in those clones (981 and 984) from
high latitude habitats. The net phenotypic disparity in height and leaf
development between ramets of such northern clones when grown in the
long-day regime to which they are adapted and the shorter-day regime of
Boston is most striking (Fic. 7).

Similarly, marked intra-clonal differences in time of height growth
cessation characterize those clones adapted to the high-elevation (short
growing season) habitats in the lower latitudes when grown in the normal-
and long-day regimes (clones 1047, 1485, and 1239, Tasre 1). In con-
trast, intraclonal diversity in response to normal- and long-day photoperiods
demonstrated by those clones originating in the low-elevation (long grow-

180 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXXV
ing season) habitats of the lower latitudes is characteristically small (clones
895, 899, 825, and 957, TABLE 1).

TABLE 1

TIME oF HEIGHT GROWTH CESSATION IN NORMAL- AND LONG-DAY REGIMES
P. trichocarpa

(1) (2) (3) (4) (5) 6) (7) (8) (9)
Origin Date of Ht. Growth Cessation |
Normal Day |Long D al D
ifference
Clone Lat. Long. Elev. Weston —_|Arn. Arb. | Arn. Ar (days)
No.| (N) (W.) (Ft.) | 19501951} 1951 | on (8)—-(7)
981 60-37 149-30 1300 6/25 6/20 8/28 69
984 60-19 149-21 ...2 6/19 = 6/20 8/19 60
1147. 48-30 =.115-55 =1850) §=8/29 8/28 8/28 10/3 36
1151 48-25 114-30 2100 8/29 8/28 9/12 9/27 15
1565 48-10 114-10 8000 8/1 7/31 8/28 9/27 30
1029. 47-30 123-55 250 9/19 9/21 10/5 14
1047. 47-35 115-25 2400 9/12 8/29 9/28 30
895 46-10 122-55 30 9/26 9/18 10/3 10/12 9
1485 46-20 112-15 5000 8/15 8/7 8/10 9/13 34
899 «945-35 =: 122-35 25 9/26 9/18 10/7 10/12 5
1324 45-15 117-45 3400 9/5 8/28 9/18 10/3 15
825 44-30 123-25 200 9/19 9/18 10/5 10/11 6
1239 44-30 114-15 5200 8/29 8/28 8/31 10/3 Be]
957 43-25 123-25 500 9/26 9/18 10/6 10/11 5
1137. «43-35 = =—-:114-25 55300) Ss 8/29) 8/28 —s 8/31 9/27 27

* Height growth cessation occurred prior to June 27, 1950.

A point of considerable physiological interest is suggested by the results
of the normal- and long-day tests. Theoretically, the net superiority in
height shown by ramets of those clones grown in the long-day environment
may be attributable to the combined influence of two causes: (1) the
longer season of growth; and (2) the longer daily light period. Although
the tests were not designed to analyze quantitative growth differences, the
available data suggest that the longer daily light period did not materially
contribute to the net seasonal height increment of plants grown in the
long day. The superiority in the height of plants given the long-day treat-
ment appears, thus, to have resulted solely from their longer growing
season

Other oe Factors Influencing Time of Height
Growth Cessa

In general, a —e was noted in the time of height growth cessa-
tion among ramets of the various clones when grown under the influence of
the same day-length regime during the same or even succeeding seasons

1954 | PAULEY & PERRY, POPULUS 181

(TaBLE 1). Most minor fluctuations (1 to 10 days) may doubtless be
attributed to inaccuracies inherent in the measurement technique em-
ployed. Others, demonstrating fluctuations of a higher order, such as the
21-day difference in date of height growth cessation of clone 1324 (TABLE 1)
as recorded at Weston and the Arnold Arboretum in 1951, suggest the
modifying influence of other factors on the photoperiodic reaction.

~

Fic. 7. Diversity in phenotype demonstrated by ramets of P. trichocarpa
clone No. 984 when grown in the normal day-length regime of Boston (left);
and in an artificially created long-day regime approximating that of its native
Alaskan habitat (right). Height growth ceased in the ramet exposed to Boston
day length on June 20, 1951; that exposed to the long-day regime stopped growth
60 days later (August 19, 1951). Scale is in feet.

That other environmental factors should serve to modify the photo-
periodic response is by no means unusual. Among the factors most suspect
is temperature. Garner and Allard (3) observed modification in the
photoperiodic requirement for soy beans when grown in different tempera-
tures, and similar reports have been made by other workers (Roberts and
Struckmeyer, 17; and Parker and Borthwick, 16). Since most differences
in time of height growth cessation between ramets of the same clone

182 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXXV

grown at Weston and the Arnold Arboretum were due to earlier cessation
of growth at Weston, the uniformly lower temperatures of that locality
may have been responsible. (Differences in the natural day length of the
two localities is negligible, since they are separated by only ca. 2 minutes
of latitude.)

On the basis of experiments currently in progress, there is evidence that
ramets of a clone grown in sod may be induced to stop height growth at an
earlier date than those grown in cleanly cultivated soil.

There is some indication that excesses of available nitrogen in the soil
(Kramer, 8) delay the onset of dormancy in apple trees, and this factor
may thus influence time of height growth cessation.

In addition to the above factors, the quite empirical conclusion may be
drawn that an apparent intraclonal diversity in physiological functions
may frequently develop among ramets of a clone. Such differences appear
to be associated with mechanical injury or attack by disease or insects.

ROLE OF THE PHOTOPERIODIC RESPONSE IN THE
ANNUAL GROWTH CYCLE OF POPULUS SPECIES

Break of Dormancy and Early Seasonal Growth

Although light exerts a profound influence on growth activity during a
large portion of the growing season in Populus, neither light nor its period-
icity appears to be directly concerned in the break of dormancy.

In the propagation area at Weston, marked intra- and interspecific dif-
ferences in time of leaf flushing are annually observable in early spring;
but preliminary results of forcing studies made in 1950, 1951, and 1952
indicated that such differences reflect a diversity in response to prevailing
temperatures, apparently acting quite independently of the light environ-
ment. In order to test more adequately for a possible light influence on
dormancy break, ramets of various clones and sibs of several progenies
representing species in all sections of Populus were grown in continuous
light, natural day, and continuous darkness for a period of ca. three
months in the late winter and early spring of 1953. The three compart-
ments were maintained at as uniform a temperature as the facilities avail-
able permitted (ca. 70° F.)

Significantly, break of dormancy was negotiated without apparent
incident by all plants in the dark compartment.! Within those clones and
seedling families which broke dormancy early, little disparity (0-2 days)
in date of dormancy break between compartments was noted. Within the
late-breaking clones and seedling families, there were, however, marked
divergencies (3-9 days); but there was no indication that complete ab-

"So-called “break of dormancy” in these studies was based on observed separation
of bud scales. As Populus buds swell, the imbricate bud scales separate, revealing the
transparent portion of the underlying scale. Examination of the plants in the dark
compartment was effected by use of weak yellow light which, in addition to the green
portion of the spectrum, is reported to be photoperiodically ineffective, at least in the
flowering response (Miller, 11).

1954 | PAULEY & PERRY, POPULUS 183

sence of light was more inhibiting to the break of dormancy than continuous
light.

Date of appearance of the first unrolled leaf was also recorded in these
studies. In most cases the flattening process was not complete in the plants
subjected to the dark treatment.

Subsequent growth of the plants in darkness gave rise to the typical
morphological modifications associated with etiolation, i.e., reduction in
leaf size, marked elongation of the shoot with the development of a hook
at the end, inhibited diametral growth, and loss of the erect habit (Fic. 8).

Fic. 8. l-year seedlings of P. trichocarpa grown for 5 weeks in normal day
(left): continuous light (center); and continuous darkness (right). All plants
were dormant at the start of the experiment and all broke dormancy 8 to 10
days later.

Results of these studies appear to confirm the conclusion that the photo-
period is ineffective in the break of dormancy or early growth in length of
Populus. Later, however, a stage in the annual growth cycle is reached
when the day length must be greater than a certain minimum in order for
height growth to continue. As has been demonstrated, high latitude clones
of P. trichocarpa, when grown in the natural-day regime at the latitude of
Boston, cease height growth on or about the time of the summer solstice. If,
however, the photoperiod is lengthened by the addition of artificial illumi-
nation, they continue to grow, ceasing only when the day length again
falls below the critical minimum. The role of the photoperiod in the annual
growth cycle of Populus thus appears to be primarily effective in the timing
of phvsiological processes concerned in the onset of dormancy.

184 JOURNAL OF THE ARNOLD ARBORETUM | VOL. XXXV
Diametral Growth Cessation

On the basis of weekly recordings of height and diametral increment
made on representatives of various Populus species in the plantations at
Weston and elsewhere during the 1952 season, it is clear that diametral
growth is always terminated after height growth ceases. Interclonal differ-
ences in the length of time between height and diametral growth cessation
ranged from ca. 10 to 30 days; but there appears to be little evidence from
these data to suggest that diametral growth cessation is independently
controlled by the photoperiod. A more likely explanation seems to be that
cessation of growth in diameter is simply one of the final, and perhaps
more or less automatic, steps in the dormancy process.

INHERITANCE OF THE PHOTOPERIODIC RESPONSE

The broad and continuous range in time of height growth cessation
which characterizes the behavior of the various ecotypes in these studies,
suggests that a large number of genes are involved in the photoperiodic
reaction. Further indications of multiple factor inheritance are contained
in the records on height growth cessation for the offspring and parents of a
north-south cross included in the 1951 studies. Under the influence of
either the normal- or long-day regime, the progeny ceased height growth
at a time intermediate to that of their parents (Table 2

TABLE 2

NORTH-SOUTH Cross
DATE OF HEIGHT GROWTH CESSATION FOR PARENTS (183A XX 118) AND
ROGENY (1599 AND 1600)

Lat. of _ Date of Height Growth Cessation, 1951
Clone No. origin | Normal-day Plot | Long-day Plot
183A 58-30 8/10 9/15
118 46-50 9/12 10/3
1599 8/25 9/21
1600 8/24 9/20

PRACTICAL AND THEORETICAL SIGNIFICANCE
OF THE PHOTOPERIODIC GROWTH RESPONSE

Seed Source

Although it would be unwise on the basis of our present limited knowl-
edge to extend the results of these studies to all ligneous species, several
practical generalizations concerning the aad ooairaa response in trees
and its importance in seed source problems may be no

(1) The utilization of seed from northern long- pins races for planting in

1954} PAULEY & PERRY, POPULUS 185

southern latitudes of long growing season will result in early seasonal
growth cessation and consequent dwarfing. Such seed sources should obvi-
ously be avoided for the establishment of forest plantations from which
maximum production of wood or fiber is desired. On the other hand, such
seed sources might prove suitable for the establishment of protection forests
in short growing season habitats at high elevations in southern latitudes.
The typically small annual increment of northern races when grown in a
short-day environment might even make them suitable for special horti-
cultural uses.

(2) Seed derived from ecotypes native of the short growing season,
high-altitude habitats of mountainous areas in the low latitudes may be
expected to react in a manner similar to northern long-day types and should
therefore be avoided as seed sources for forest plantations at the same or
more southern latitudes. Such sources of seed may, however, prove highly
desirable for introduction into certain long-day environments in northern
latitudes having suitably long growing seasons. Support for this generaliza-
tion is attested by the long history of successful North American tree in-
troductions into the maritime countries of Europe. Conversely, movement
of long-day races of native European species into the United States has not
met with notable success.

(3) Ecotypes native of long growing season habitats in any particular
latitude should be avoided as seed sources for short season habitats at the
same latitude because of their susceptibility to early autumn frost damage.
For the same reason, such seed sources cannot be successfully moved
northward into a long-day environment.

The problem of selecting suitable seed sources of any species for intro-
duction into another environment must obviously take into consideration
numerous other environmental factors and genetic characteristics. The
above recommendations may be found useful, therefore, only to the extent
that they may contribute to limiting the field of search for desirable
genotypes. Final decisions on adaptability of the proposed introduction to
the new habitat must be based on the observed interaction that results be-
tween the introduced genotype and all the factors of its new environment.

Breeding

Because of its fundamental role in the vegetative growth cycle, the
photoperiodic response should obviously be a matter of primary concern
to the tree breeder. As a means of increasing the net yields of wood or
fiber, gene combinations nicely adapted for full utilization of the growing
season in a particular habitat, may well prove to be quite as promising as a
search for heterotic hybrids.

In view of the probably high genetic diversity of the photoperiodic re-
sponse in wild tree populations occupying a wide diversity of habitats,
initial concern should be directed to the selection of suitable parental
materials. This is especially true in F; breeding programs and may, to a
large extent, explain the unsatisfactory results that have been obtained

186 JOURNAL OF THE ARNOLD ARBORETUM _ [vot. xxxv

by various workers through the selection of parental stocks based on their
taxonomic identity and availability, rather than their genetic quality.

Since the photoperiodic response appears to be controlled by a large
number of factors, the production of a wide-ranging, transgressive series
of photoperiodic types may be expected in F. populations. Theoretically,
by the initial use of parental ecotypes which give reactions approaching
the extremes in day-length response, an F» will result which may be ex-

ected to contain gene combinations adaptable to almost any growing
season length at any latitude.

SUMMARY

For the purpose of studying ecotypic variation in Populus, a living
collection representative of the ranges of various species in this genus was
undertaken in 1947 by the Maria Moors Cabot Foundation for Botanical
Research. The studies here reported are concerned primarily with the in-
fluence of the photoperiod on time of height growth cessation as observed
in the various test areas located in the vicinity of Boston, Massachusetts.

Marked variation in the time that height growth stopped was observed
between clonal lines of single species when grown in the same day-length
regime and otherwise uniform environment. Analysis of these data re-
vealed that the time of height growth cessation was inversely correlated
with the latitude of origin of each clone. Among clones native of uniform
day-length zones, the time at which height growth ceased was directly cor-
related with the length of the frost-free season prevailing in the native
habitat of each clone. On the basis of these observations, the conclusion is
made that adaptation of Populus species to various habitats differing in
length of frost-free season is effected by a genetic mechanism which con-
trols the duration of their seasonal period of growth. The photoperiod,
which is the only factor of the environment with a uniform seasonal varia-
tion that is constant from year to year, functions as the timing device for
this mechanism.

Modifications in time of height growth cessation were effected by expos-
ing ramets of various clones to artificially lengthened or shortened photo-
periods in otherwise uniform environments. The time of height growth
cessation was thus demonstrated to be the result of an interaction between
the individual’s genotype and the photoperiod. Also observed were other
intraclonal modifications in time of height growth cessation that could not
be attributed to photoperiodic response. Differences in temperature, avail-
able nitrogen, intensity of cultivation, and various other factors are be-
lieved to exert a modifying influence on the time that height growth ceases.

Hybrids between clones of northern and southern origin gave a photo-
periodic response intermediate to the responses of the parents. These re-
sults, and the widely varying photoperiodic responses shown by the various
ecotypes used in these studies, indicate that the number of genes involved
in the photoperiodic reaction is large.

1954] PAULEY & PERRY, POPULUS 187

Experiments on the break of dormancy in the spring indicated that temp-
erature rather than photoperiod is the major factor controlling the initiation
of new growth after the period of winter dormancy.

Several practical generalizations concerning the photoperiodic response
in trees and its importance in seed source problems and in breeding are
noted.

LITERATURE CITED

1. Bocpanov, P. L. 1931. O fotoperiodisme u drevensnykh prod (On the
photoperiodism of tree species). Trudy po lesnomu opytnomu delu 10:
54-55. Leningrad, State Research Institute for Forestry and Forest
Industry. U.S. ae Service Translation No. 137, 1935.

Ciausen, J., D. D. Keck, AND W. M. Hiesey. 1940, 1945, 1948. Experi-

mental fies on the nature re species. I, II, III. Carnegie Institution of
Washington Pubs. 520, 564, 58

GARNER, W. W. AND H. A. ae 1920. Effect of the relative length of
day and night and other factors of the environment on growth and repro-
duction in plants. Jour. Agric. Res. 18: 553-606

HorFMANN, DretricH. 1953. Die Rolle des Photoperiodismus in der
ahs hare pceanar? Zeitschrift fiir Forstgenetik und Forstpflanzen-
ziichtung 2 (2): 45-47.

HuMPHREYS, W. J. ie. Physics of the Air. McGraw-Hill Book Co. 3rd
ed. 676 pp.

6. JesTER, J. R. AND P. J. KRAMER. 1939. The Effect of Length of Day on the

Height Growth of Certain Forest Tree Seedlings. Jour. Forestry 37:

> Ww i)

on

ae
°
B
ae
2 oo
Oo
Z
a8

1951, Progeny tests with birch: Preliminary results with

young plantings. Svensk Papp. Tidn. 54: 379-393.

8. Kramer, P. J. 1936. Effect of variation of length of day on growth and
dormancy of trees. Plant Phys. 11: 127-137.

9. LANGLET, O. 1942-3. Photoperiodismus bei der gemeinen Kiefer (Pinus
sylvestris L.). Medd. fran Statens Skogsforsdksanst. Stockholm. 33:

10. Matzke, E. B. 1936. Effect of street lights in delaying leaf-fall in certain
trees. Am. Jour. Bot. 23: 446-451.

11. Mruter, E. C. 1938. Plant Physiology. McGraw-Hill Book Co. 2nd Ed.
1201 pp.

12. Mosuxov, B. S. 1930. ‘“Photoperiodicity of Certain Woody Species” (in
Russian with English summary). Bull. App. Bot., Genet. and Pl. Breeding
23 (2): 479-510.

13. ————.. 1932. “Photoperiodicity of trees and its practical importance”
(in Russian). Bull. App. Bot. Genet. and Pl. Breeding, Ser. A, Soc.
Plant Ind. 2: 108-123. U.S. Forest Service Translation No. 195, 1935.

14, ————— . 1935. Photoperiodismus und Frostharte ausdauernder Gewachse.
Planta 23: 774-803.

15. Oxmstep, C. E. 1951. Experiments on photoperiodism: Dormancy and
leaf age and abscission in sugar maple. Bot. Gaz. 112: 365-393.

16. PARKER, M. W., AND H. A. BortHwicx. 1939. The effect of variation in
temperature during photoperiodic induction upon initiation of flower
primordia in Biloxi soy bean. Bot. Gaz. 101: 145-167.

bo
bh

JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxv

. Roperts, R. H., AnD B. E. STRUCKMEYER. 1938. The effects of temperature
and other environmental factors upon the photoperiodic responses of some

of the higher plants. Jour. Agric. Res. 56: 633-677.

. SyLvEN, N. 1940. Lang- och kortdagstyper av de svenska pares

Svensk Papperstidning 43 (17): 317-324; (18): 332-342; (19):

351.

. Turesson, G. 1922. The species and the variety as ecological units.

. TurRRILL, W. B. 1940, Experimental and synthetic plant taxonomy. In
J. Huxley, ed., The New Systematics: 1.

+ WS, Department of Agriculture. 1941. Climate and Man. Yearbook of
Agriculture. U. S. Government Printing Office. 1248 pp.

. WirHrow, R. B. AND H. M. Benepict. 1936. Photoperiodic responses of
certain green annuals as influenced by intensity and wave length of arti-
ficial light used to lengthen the daylight period. Plant Phys. 11: 225-249,
807-819.

1954] HU, NOTES ON THE FLORA OF CHINA, III 189

NOTES ON THE FLORA OF CHINA, III
SHIU-YING Hu
With two plates

CRITICAL NOTES on ambiguous binomials concerning the Chinese flora,
additional information regarding certain Chinese plants, and the proposal
of eight new species, four new varieties, and two new combinations are
presented in this paper. Unless identified by (G), indicating that the
material is deposited in the Gray Herbarium, all the specimens cited are
to be found in the Herbarium of the Arnold Arboretum.

Clematis angustifolia Auct.

Jacquin in 17621 published the species Clematis angustifolia, from the
coast of southern Europe. According to his description the species was a
glabrous vine with pinnate leaves and linear leaflets. Pallas in 1776 de-
scribed Clematis hexapetala, an erect plant with tomentose perianth from
eastern Siberia in the regions of the Argun and Onon Rivers. In 1786
Jacquin, probably on account of the shape of the leaf segments, doubtfully
amalgamated these geographically widely separated and morphologically
manifestly different species. In the discussion of his Clematis angustifolia
he wrote as follows, expressing his doubt: “An haec eadem cum Am-
maniana planta, a Pallasio citata, sit, dubito, quum nostra sit glabra tota,
Ammanius vero dicat caules superius lanuginosos . . .” In the same year,
under the name Clematis angustifolia, he published a colored plate illus-
trating a plant which he described as erect and with pinnate recurved
leaves and linear-lanceolate leaflets. Judging from this illustration and
description, Jacquin’s concept of his own Clematis angustifolia had, by
this time, changed from a climbing glabrous vine growing on the coast
of southern Europe to an erect pubescent plant occurring in the mountains
of central Siberia, for in the Gray Herbarium there is a specimen collected
by Schschukin in Irkutsk which matches Jacquin’s illustration in every
respect. This illustration of Jacquin’s has been very misleading, for in the
last two centuries botanists who named plants by matching them with
illustrations have identified all members of that heterogeneous group of
erect herbaceous perennial Clematis from North China, Manchuria, Korea,
and Eastern Siberia as Clematis angustifolia Jacq. Meanwhile most au-
thors who have published accounts of the flora of eastern and northeastern
Asia have indiscriminately applied such identifications. Now, both in the
herbaria and the literature, we have this oriental taxon carrying an occi-
dental binomial. As Jacquin’s species was first published as a climbing

’ Enumeratio yond ce ita i quae Sponte Crescunt in Agro Vindobonensi,
Montibusque Confinibus

190 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXXV

plant growing on the coast of southern Europe, the application of that
binomial should be limited to plants of that region, in spite of the fact
that Jacquin later changed the concept of his species. A. P. De Candolle
in 1817 (Regni Vegetabilis Systema Naturelle 134, 154) treated Clematis
maritima Linn. as a variety of Clematis flammula Linn. var. maritima (L.)
DC. in one place and later used it again as a synonym of Clematis angusti-
folia Jacq. In so doing, he seems to have given us a hint that Jacquin’s
original Clematis angustifolia might have been a Clematis flammula Linn.
var. maritima DC., since Clematis flammula Linn. is a vine with pin-
natisect leaves, and the variety, C. flammula Linn. var. maritima (L.)
DC., has linear segments. This vine habit and the pinnatisect leaves with
linear segments are distinguishing characters which Jacquin at first ascribed
to his C. angustifolia. In the herbarium of the Arnold Arboretum there is
a collection from Yeste, Spain, labeled C. flammula Linn. var. maritima
(L.) DC. This specimen represents a glabrous vine with pinnatisect leaves.
The linear segments of these leaves appear similar to those of some form
of the Chinese elements. It is very likely that this represents the true C.
angustifolia Jacq., which is very different from the taxon of northeastern
Asia, which should be designated as Clematis hexapetala Pallas.

Hayata in 1913 overlooked Jacquin’s binomial and published a Clematis
angustifolia from Formosa. This species was originally described as
Clematis leschenaultiana DC, var. angustifolia Hayata in Jour. Coll. Sci.
Univ. Tokyo 30: 16. 1911. It represents a vine with densely tomentose
stems and leaves. The leaves are trifoliate with ovate or oblong-lanceolate
leaflets which are remotely serrate. The achenes of this species are fusi-
form. Clematis leschenaultiana DC. was originally described from Java.
It has been recorded from the Philippines, Formosa, and the warmer re-
gions of the Chinese mainland, as well as from the western Malaysian area.
After comparing specimens collected within this wide range I have de-
cided that the Formosan material is not specifically distinct.

Clematis hexapetala Pallas, Reise 3: 735, pl. Q, fig. 2. 1776, et Voy. 4:
701, pl. 14, fig. 3. 1793. — Komarov & Schischkin, Fl. URSS 7: 318.
1937. — Kitagawa, Lin. Fl. Mansh. 217. 1939. — Nakai in Jour. Jap.
Bot. 20: 191. 1944.

Clematis i a Jacq.. Ic. Pl. Rar. 1: 11, pl. 104. 1786 (non Jacq. 1762).
. sy 153. 1817, et Prodr. 1: 7.1824.— Bunge in Mém. Div.

ae Acad. 7 St. Pétersb. 2: 75 (Enum. Pl. Chin. Bot.). 1833. — Ledeb.,
Fl. Ross. 1: 2. 1841.— Maxim. in Mém. Div. Sav. Acad. Sci. St. Pétersb.
9: 468 (Ind, Fl. Pekin.). 1859, et Enum. Pl. Mongol. 2. 1889.— Franch.
in Nouv. Arch. Mus. Paris II. 5: 166 (Pl. David 1: 14). 1882.—Hemsl.

: . . ; ]
Tokyo 26: 11 (FI. Kor. 1: 11). 1909, et 31: 426 (FI. Kor. 2: 426). 1911. —
Cowdry in Jour. Roy. As. Soc. N. China 53: 160 (Pl. Peitaiho). 1922
—Limpricht, Bot. Reis. Hochgeb. Chin. Ost-Tib. 374. 1922.— Rehder
in Jour. Arnold Arb. 4: 185. 1923.— Komarov, [Fl. Manchzh.] 2: 288.
1950

Clematis pallasii Gmel. Syst. 873. 1791.

1954] HU, NOTES ON THE FLORA OF CHINA, III 191

Clematis recta Linn. var. angustifolia (Jacq.) Kuntze in Verh. Bot. Brand.
26: 112. 1884.

Clematis angustifolia Jacq. var. breviloba Freyn in Oesterr. Bot. Zeit. 45:
59. 1895, et 51: 374. 1901.

Clematis angustifolia Jacq. var. dissecta Yabe, Higasi-Moko Syokubutsu
Mokuroku 14. 1917.

Clematis angustifolia Jacq. Spey dissecta (Yabe) Kitagawa in Rep. First
Sci. Exped. Manch. IV. 4: 17, 3

Clematis hexapetala Pallas a dissecta (Yabe) Kitagawa, Lin. Fl. Mansh.

217. 1939.

Clematis er Pallas forma breviloba (Freyn) Nakai in Jour. Jap. Bot.
20: 191. 194

CHINA: Chili [Hopei]: Hsiao-wu-tai shan, T. N. Meyer 92; Peking, S. W.
Williams, Aug. 1876 (G); Hsing-lung shan, J. C. Liu 622.

EASTERN SIBERIA ann MANCHURIA: Kirin, F. H. Chen 182 age
Dairen, Chinchou, P. H. Dorsett & W. J. Morse 5937 (G); Harbin, P. H.
J.H. Dorsett 3292 (G); Nertschinsk [Nerchinsky], F. Karo (Plantae “ee
152c (1soTyPE of Clematis angustifolia Jacq. B breviloba Freyn); Blagowjest-
schensk, F. Karo 113b; Chabarovsk [Khabarovsk], V. Komarov 706; Khingan-
skia, D. Litvinow 326, 1015; [Ta-ho-shang shan], K. Kobayashi, June 19, 1933;
Suifenho, B. V. Skvortzov, ‘July 10, 1926; Greater Khingan, 2 ia BY.
Skvortzov, July 27, 1938. Mukden [Moukden], E. H. Wilson 8

Pallas was the first post-Linnaean botanist who described and illustrated
the northeastern Asian erect herbaceous perennial Clematis and named it
C. hexapetala. The leaf of his plant is ‘per caulem opposita, adscendentia,
pinnata: foliolis duris venosis, marginatis, acutis, imi paris ramoso-quad-
rifidis, superioribus lanceolatis bifidisque, terminali tripartito.” The seg-
ment of the perianth is “oblonga, extus tomentosa.” Plants so charac-
terized had been recorded from Manchuria, northern Korea, and the North
China Highlands extending from Long. 108° E. and Lat. 34° N. north-
eastward to Long. 122° E. and Lat. 40° N. Unfortunately, on account of
Jacquin’s changed concept of his European species of C. angustifolia and
his misleading illustration of 1776, this taxon has appeared as C. angusti-
folia Jacq. in botanical literature, especially in that concerning the flora
of North China.

Clematis hexapetala Pallas is essentially an upland species growing on
grassy mountain slopes. Its distribution is very interesting. So far as our
material and records show, it has never been collected west of Long. 103°
E. nor south of Lat. 34° N. Its range forms a U-shaped area on the map,
with Khabarovsk and northern Korea on the curve, northern Manchuria,
Dahuria, and the Lake Baikal area on the northern arm, and with southern
Manchuria and the North China Highlands on the southern arm.

The material collected within this range exhibits a definite pattern of
variation in the size and shape of the ultimate segment of the pinnatisect
leaves. Some of these are linear, measuring up to 5 cm. long and 5 mm.
wide in the middle, while others are lanceolate, up to 10 cm. long and 10
mm. or more wide in the middle. Since 1895 many trinomials have been

192 JOURNAL OF THE ARNOLD ARBORETUM _ [vot. xxxv

proposed for the different forms of this variation. Although in the original
description the measurements of the ultimate segments of the leaves of
typical C. hexapetala Pallas were not given, and the type material is not
accessible to me for comparison, from Pallas’ illustration I am inclined
to think that he definitely had a narrow-segmented form with segments
2.5—5 cm. long and 3-5 mm. wide at the middle, and acute at the apex.
When Freyn described his varieties, he stated for C. angustifolia Jacq.
a longiloba, “Die Blatter doppelt gefiedert, mit lineal-lanzettlichen Ab-
schnitten; letztere 8-10 mal linger als breit (4-6-10 mm.). Die Bliiten
bis 5 cm. im Durchmesser, die jungen Sepalen aussen dickt filzig wollig.”’
For his C. angustifolia Jacq. B breviloba he gave no measurements but
simply added, “wie vorige, aber die Blattabschnitte bei gleicher Breite viel
kurzer.” Here in the herbarium of the Arnold Arboretum there is an iso-
type, F. Karo 152c, for the latter variety. The size and shape of the ulti-
mate segments of the pinnatisect leaves are identical with those shown in
Pallas’ illustration. Freyn probably did not realize that Nerchinsky, the
locality where Karo 152c was collected, lies between the rivers Argun and
Onon, the type locality of C. hexapetala Pallas. The morphological iden-
tity and the geographical coincidence lead me to decide that Freyn’s C.
angustifolia Jacq. 8 breviloba is identical with the typical C. hexapetala
Pallas. The trinomials proposed by the Japanese botanists have been in-
terpreted as synonyms of C. angustifolia Jacq. B breviloba Freyn by
Nakai. They are here treated as synonyms of C. hexapetala Pallas.
Pritzel in 1900 recorded this species under the name of C. angustifolia
Jacq. from T’ai-pa-shan of southwestern Shensi, as represented by Giraldi
865. If his identification is dependable, this collection marks the southern-
most limit in the range of distribution for the species. As I have not seen
Giraldi’s specimen, I am not sure whether it actually belongs here or not.

Clematis hexapetala Pallas var. longiloba (Freyn) comb. nov.
Clematis angustifolia Jacq. q longiloba Freyn in Oesterr. Bot. Zeit. 45: 59.
1895.

CHINA: Chili [Chihli, Hopei]: Peking, Western Hills, P. H. Dorsett &
W. J. Morse 7035; same locality, Bretschneider in 1881 (ex Herb. F. B. Forbes
1807); Kiang-hsuai ho, T. F. King 171: San-tun-ying, F. N. Meyer 91; Hsiao-
wu-tai shan, F, N. Meyer 1352; same locality, C. W. Wang 61450; Wei-chang,
Wm. Purdom 44; without precise locality, Father Chanet 28. Shansi: cen-
tral part of the province, Lu-yah shan, H. Smith’s collector, Lao Ch’in 8147.

EASTERN SIBERIA orn MANCHURIA: Zejshaja Pristan am Zeaflusse, F.
Karo 334, July 1899; Blagovesczensk, F. Karo, July 16, 1904; Amur, super et
medius, Korskinsky in 1891 (G); Ircutsk [Irkutsk], Schschukin (G); Moukden,
J. Webster in 1887 (G); Dahuria, Turczaninow 8, Nov. 1859 (G).

KOREA: Pyengyan, Mrs. R. K. Smith, July 5, 1937.

This variety is characterized by the larger leaf segments, which are
10-15 mm. wide, attenuated at both ends, acute, rarely obtuse at the apex,
sparsely villose on the principal nerves, especially beneath, glabrescent

1954 | HU, NOTES ON THE FLORA OF CHINA, III 193

later in the season. This variety has been introduced into cultivation. I
have seen material from Jewell Nursery, Lake City, Minnesota, which is
identical with our spontaneous collections. In North China the plant grows
on grassy slopes along the dry hillsides. Its white flowers appear in June
and July. The obovate achenes are 5 mm. long, 3.5 mm. wide, compressed,
villose, and with a persistent densely villose curved style up to 3.5 cm. long.

Clematis hexapetala Pallas var. smithiana var. nov.

Herba erecta perennis, 20-30 raro usque 60 cm. alta; foliis pinnati-
sectis, segmentis ultimis lanceolatis, 1.5-3 cm. longis, 3-7 mm. latis, apice
obtusis apiculatisque, subtus villosis, inflorescentiis subpaniculatis, peri-
anthiis lanatis, segmentis oblanceolatis, 1 cm. longis, 3 mm. latis, antheris
oblongis, 1.5 mm. longis, carpellis albo-villosis.

CHINA: Shansi: Yin-ch’eng, Chung-t’iao shan, H. Smith 6039 (TYPE);
Huo hsien, T. Tang 1010.

This is a dwarf variety which is usually 20-30, rarely up to 60 cm. high
at the flowering stage. It can easily be distinguished from all other vari-
eties of this species by its villose lower leaf-surface. It occurs in southern
Shansi at an altitude of 1850 meters. The white flower appears in July.

Clematis hexapetala Pall. var. tchefouensis (Debeaux), comb. nov.

Clematis angustifolia Jacq. var. tchefouensis Debeaux in Act. Soc. Linn. Bor-

deaux 31: 117 (Fl. Tché-fou 22). 1877.
Clematis angustifolia sensu Faber in Denkschr. Entwickel. Kiautschou 31.
98.— sensu Rehder in Jour. Arnold Arb. 4: 185. 1923, pro parte, non
Jacquin, 1762.

CHINA: Shantung: Tche-fou [Chefoo, Chih-fou, Yen-t’ai], 0. Debeaux
(1soTyPE of Clematis angustifolia Jacq. var. tchefouensis Debeaux); Tsingtao,
Li-chuan, C. Y. Chiao 2979; near coast, A. Jacot, July 1, 1927; Lao shan, 33
miles south of Tsaingtao, C. Y. Chiao 2691.

This variety was originally described on the basis of material collected
from the north shore of the Shantung Peninsula. Additional material has
proved its rather extensive range, extending to the southern end of the
peninsula. It can be distinguished from typical Clematis hexapetala Pall.
by its subglabrous perianth, which is 2—2.5 cm. in diameter at anthesis.

Clematis hexapetala Pall. var. insularis, var. nov.

Herba erecta, caudicibus 1 m. altis, 4 mm. crassis, foliis pinnatisectis,
segmentis lobatis vel partitis, ultimo lanceolato, 3—6 cm. longo, 8-13 mm.
lato, apice obtuso mucronatoque; floribus subpaniculatis, pedicellis 5-8 cm.
longis, perianthiis glabrescentiis, 2.5 cm. diametro.

CHINA: Shantung: Tsingtao, Tsingtao Island, C. Y. Chiao 2521 (TYPE).

This erect, herbaceous, broad-segmented variety is endemic to a small
island on the southern coast of the Shantung Peninsula, where it grows on
sandy slopes at sea level. Its white flowers appear in mid-June. Its leaf-

194 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxv

segments resemble those of C. hexapetala Pall. var. longiloba (Freyn)
S Hu, but the latter taxon can easily be distinguished by its white
lanate perianth and sparsely villose nerves on the lower surface of the
leaves, while the perianth of this variety is glabrescent and the leaves
entirely glabrous,

Clematis hexapetala Pall. var. elliptica var. nov.
Clematis angustifolia sensu Finet & Gagnepain, Contrib. Fl. As. Or. 1: 21.
1905, pro parte. — sensu Loes. in Beih. Bot. Centralbl. 37 (Abt. 2): 112.
1919, non Jacquin, 1862.

Herba erecta perennis; foliis pinnatisectis, praeter nervos subtus glabris,
segmento ultimo lanceo-elliptico, 1.5-3 raro usque 5 cm. longo, 4—5 raro
usque 8 mm. lato, apice obtuso mucronatoque; perianthiis glabrescentiis.

CHINA: Shantung: Tsingtau [Tsingtao], Zimmerman 205 (tvPE), 448.

This variety can be distinguished by its very deeply cut lanceo-elliptic
segments of the leaves which are obtuse at the apex. In general appearance
it resembles C. hexapetala Pall. var. smithiana S. Y. Hu of southern Shansi,
but the latter variety has a lanate perianth and villose lower leaf surfaces.

Freyn in 1895, on the basis of F. Karo 1256 collected from Nertschinsk
| Nerchinsky], described a form which he named forma stenophylla. Ac-
cording to his description, the segments of the leaves are up to 6 cm. long
and only 1 mm. wide. In the Gray Herbarium of Harvard University
there is a fruiting specimen, 7. Y. Cheo & L. Yen 98, collected from Fei-
hsien of Shantung Province, of which the segments of the leaves are 5—9
cm. long, 3 mm. wide. Flowering material of the narrow-segmented form
from the latter region is awaited for its identity.

Tinospora craveniana sp. nov. (Pl. /, fig. 6).
Frutex scandens, ramis striatis sulcatisque, 2-3 mm. diametro, hor-
notinis hirsutis, internodiis 7-10 cm. longis; foliis tenuiter chartaceis,

stitio inter apices auricularum 1.5—2.5 cm. longo, apice acuminatis, acumine
5-8 mm. longo, integris, praeter nervos glabris, nervis primariis 5, pal-
matis, nervis secundariis laxe reticulatis, utrinque prominulis, petiolo 2.5—
3.5 cm. longo, hirsuto; inflorescentiis racemosis; ¢ racemis fasciculatis,
pedunculis gracilibus, 15 mm. longis, hirsutis, rhachibus 4 cm. longis,
bracteis lanceolatis, 1-2 mm. longis, pedicellis 12-30 mm. longis, sparse
hirsutis, prophyllis 1 vel 2; floribus 3-meris, sepalis 6, glabris, extimis
oblongis, 2 mm. longis, intimis lineari- -oblongis, 4—5 mm. longis, 1 mm.
latis, apice acutis; petalis 6, carnosis, apice suborbicularibus, basi cunea-
tis, 2 mm. longis; staminibus 6, 3 mm. longis, thecis oblongis, 1 mm. longis,
unite lateralibus dehiscentibus; ovario rudimentario minuto, globoso; 9
racemis solitariis, pedunculis as cm. longis; floribus ignotis; fructibus
sa gaa 9-10 mm. diametro, endocarpiis osseis, subhemisphaericis,
7-8 mm. diametro, ventro lene dorso lineis obsoletis inconspicuis
cise as

1954] HU, NOTES ON THE FLORA OF CHINA, III 195

CHINA: Kiangsi: Hwang-kong shan, Y. K. Hsiung 6402 (type, fruit).
Szechuan: Mt. Omei, 7. T. Yii 563 (TYPE, staminate flower).

The outline of the anterior portion of the leaf of this species appears
to resemble that of JT. malabarica (Lam.) Miers., but the base of the leaf
of the latter species is cordate, not sagittate. The indumentum on the
nerves of the lower surface and the sagittate leaf-base of T. craveniana
suggest relationship with 7. sagittata (Oliver) Gagnep., but the latter
species has linear-lanceolate leaves, smaller flowers with the sepals measur-
ing only 2.5—3 mm. in length, obtuse or rounded at the apex, and sub-
orbicular anthers.

This species is named in honor of Miss Mary G. Craven, who retired in
January 1953 after forty-eight years of service in the Herbarium of the
Arnold Arboretum of Harvard University. Her skillful handling of the
specimens has established a monument to her care of the material as well
as to her friendship and association with all who have done taxonomic
research in this herbarium, from the days of its founder, Charles S. Sargent,
up to the present.

Tinospora imbricata sp. nov. (PI. /, fig. 2).

shia dik capillipes sensu Chun in Sunyats. 4: 176, fig. 1940. — sensu
moto in Taiwania 1: 32. 1948, pro parte, non Ae ers 1908.

Frutex scandens, ramulis striatis, 2.5 mm. diametro, sparse ee
internodiis 7-13 cm. longis; foliis chartaceis, ovato-oblongis, 12—14
longis, 44.8 cm. latis, basi cordatis, lobis auriculatis, imbricatis, ace
caudatis, acumine 2 cm. longo, supra glabris, subtus praeter nervos glabris,
nervis primariis 7, palmatis, utrinque elevatis, nervis secundariis laxe
reticulatis, utrinque conspicuis; petiolo 7 cm. longo, hirsutis, basi tortile;
floribus ignotis; infructescentiis subracemosis, pedunculis 3.5—9.5 cm.
longis, glabrescentibus, pedicellis 9-12 mm. longis, stipite ovarii 2-5 mm.
longo; fructibus rubris, subglobosis, 8-9 mm. diametro, endocarpiis per-
gameis, 7 mm. longis, 8 mm. latis, ventro excavatis, dorso lineis obscuris,
ceterum inconspicuo-rugosis.

CHINA: Kwangsi: Yao-shan, C. Wang 40521 (TYPE).

This species occurs along the streams in central Kwangsi at altitudes
of about 1300 meters. Its red fruits remain on the vine in December.
Gagnepain in 1908 described J. capillipes on the basis of Balansa’s collec-
tion from Tonkin.

The type of 7. capillipes represents a staminate plant which has pu-
bescent sepals. I have seen no specimen of this species. Judging from
Gagnepain’s illustration (Fl. Gén. Indo-Chine 1: 133. fig. 14, 11. 1908),
that species is characterized by its ovate leaves with sagittate bases. The
basal lobes are rounded and are far apart. Chun in 1940 interpreted C.
Wang 40521 as T. capillipes Gagn., but Wang’s collection has ovate-oblong
leaves with imbricate basal lobes and represents a taxon very different
from Gagnepain’s species. Yamamoto did not see Wang’s collection. In

196 JOURNAL OF THE ARNOLD ARBORETUM _ [vot. xxxv

recording the occurrence of T. capillipes Gagn. in Kwangsi he simply
adopted Chun’s interpretation.

Tinospora intermedia sp. nov. (Pl. J, fig. 5).

Frutex scandens, ramulis vetustioribus longitudinaliter rimulosis, hor-
notinis hirsutis; foliis chartaceis, sagittato-oblongis, raro sagittato-ovatis,
7-11 cm. longis, 3.5-4.5 cm. latis, basi sagittatis, interstitio inter apices
auricularum 3.5 cm. longo, apice acuminatis, acumine 6-8 mm, longo,
apiculato; inflorescentiis staminatis fasciculatis, pedunculis 2-3.5 cm.
longis, bracteis ovatis, ciliatis, 1-1.5 mm. longis, pedicellis 1-1.5 cm.
longis, prophyllis 1 vel 2, ovatis, apice ciliatis, cum pedunculis glabres-
centibus; sepalis 6, glabris, extimis ovatis, 1-2 mm. longis, intimis oblanceo-
latis, 3 mm. longis, petalis 6, carnosis, suborbicularibus, apice truncatis;
staminibus 6, petalis longioribus, antheris subglobosis, 0.5 mm. diametro;
infructescentiis paniculatis, pedunculis 9 cm. longis, fructibus ellipsoideis,
8 mm. longis, 7 mm. diametro, apice apiculatis, endocarpiis pergameneis,
ventro excavatis, dorso ceterum lineis conspicuo-tuberculatis.

CHINA: Szechuan: Mt. Omei, C. Y. Chiao & C. S. Fan 263; same
locality, C. L. Chow 5777; W. P. Fang 16320 Sees, staminate aati 17522
(type, fruit); W. K. Hu 8803 (sterile); T. T. Vii

The leaves of this species appear to resemble those of 7. craveniana
S. Y. Hu in size and texture, but the basal lobes of the latter species point
backward and its fruit has a bony rugose endocarp, while the basal lobes
of this species all point outward and the endocarp is pergameneous and
conspicuously tuberculate. The fruit characters suggest relationship with
T. szechuanensis S. Y. Hu. Occasionally a few leaves of certain specimens,
such as Chiao & Fan 263, are short and broad, appearing sagittate-ovate.
This character suggests some relationship with T. capillipes Gagnepain,
which has pubescent sepals. On account of its resemblance to several
species in different respects, it is here treated as an intermediate species.

Tinospora szechuanensis sp. nov. (PI. /, fig. 1).

Frutex scandens, ramis striatis et sulcatis, 2.5-3 mm. diametro, horno-
tinis hirsutis, internodiis 6-7 cm. longis; foliis subcoriaceis, integris, sagit-
tatis, 10-13 cm. longis, 4—4.5 cm. latis, basi sagittatis, interstitiis inter
apices auricularum 5.5 cm. longis, apice acuminatis, acumine 15-20 mm
longo, cuspidato, supra glabris, subtus praeter nervos glabris, nervis pri-
mariis 5, palmatis, utrinque elevatis, nervis secundariis laxe reticulatis,
supra obscuris, subtus prominulis; petiolo 5.5—6 cm. longo, glabrescente;
floribus ignotis; infructescentiis paniculatis, pedunculis 9 cm. longis, axibus
secundariis 10-15 mm. longis, pedicellis 5-10 mm. longis, cum pedunculis
hirsutis; fructibus ellipsoideis, 11 mm. longis, 7-8 mm. diametro, stigmatis
subapiculatis, endocarpiis pergameneis, subellipsoideis, 8 mm. longis, 7mm.
latis, ventro excavatis, dorso lineis conspicuo-tuberculatis.

1954] HU, NOTES ON THE FLORA OF CHINA, III 197

CHINA: Szechuan: Hung-ya, Wa-wu shan, E. H. Wilson 3528 (TYPE,
fruit).

This species is characterized by its subcoriaceous sagittate leaves with
the auricles pointing outward, paniculate infructescence with peduncles
9 cm. long, and ellipsoid drupes with conspicuously tuberculate parchment-
like endocarp. It is closely related to T. sagittata (Oliver) Gagnepain, but
the leaves of the latter species are linear-lanceolate, sagittate, with the
auricles pointing backward, the infructescences are racemose with the
peduncles 4-6 cm. long; and the fruits are subspherical with bony endo-
carp inconspicuously tuberculate (Pl. /, fig. 3).

Tinospora yunnanensis, sp. nov. (Pl. /, fig. 4).

Frutex scandens, ramulis striatis et sulcatis, vetustioribus tuberculatis,
lenticellis conspicuis, orbiculatis, elevatis, hornotinis hirsutis vel glabres-
centibus; foliis subcoriaceis, sagittato-ovatis, 12-14 cm. longis, 4.5-5.5
cm. latis, basi cordato-sagittatis, auriculis rotundatis, interstitia inter apices
auricularum 1.5-2.5 cm. longa, apice acuminatis, acumine 11-13 mm.
longo, apiculato, utrinque rugosis, praeter nervos subtus glabris, nervis
primariis 7, utrinque evidentibus, reticulatis obscuris; inflorescentiis
staminatis, ramis vetustis positis, racemosis, racemis solitariis vel fascicu-
latis, pedunculis 4-5.5 cm. longis, glabrescentibus, bracteis lanceolatis,
2 mm. longis, apice ciliatis, pedicellis 7-10 mm. longis, pilosis; sepalis 6,
extimis ellipticis, 1.5 mm. longis, 0.75 mm. latis, intimis unguicularibus,
2 mm. longis, 1.5 mm. latis, glabris; petalis 6, suborbicularibus, basi cunea-
tis; staminibus 6, filamentis petalis subaequalibus; ovario rudimentario
globoso; floribus pistillatis fructibusque ignotis.

CHINA: Yunnan: without precise locality, H. T. Tsai 53100 (TyPE).

This species is closely related to T. capillipes Gagnepain, but the latter
species has villose sepals, while those of this species are glabrous.

Chimonanthus salicifolius, sp. nov.

Frutex, ramulis subteretis, puberulis; foliis lineari-lanceolatis, 3-9 cm.
longis, 1-3 cm. latis, basi obtusis, apice obtusis vel acutis, subcoriaceis,
supra glabra, paulum nitidis, subtus opacis, hirsutis, costa utrinque elevata,
nervis lateralibus 5 vel 6 paribus, reticulatis; floribus solitariis, axillaribus,
pedicellis brevissimis, 4 mm. longis, bracteolis imbricatis, ovatis, puberulis;
perianthiis exterioribus rotundatis, puberulis, interioribus ignotis.

CHINA: Kiangsi: Hsiu-shui, Y. K. Hsiung 5489 (TYPE).
This species is closely allied to C. mitens Oliver, but the latter species
has glabrous ovate-elliptic leaves with a long acuminate apex.

Euonymus orgyalis W. W. Smith in Notes Bot. Gard. Edinb. 13: 161.
1921.

INA: Yunnan: Si-chour hsien, Faa-doou, K. M. Feng 12044; same
district, Ma-chia, K. M. Feng 12502; Mar-li-po, Huang-jin-in, K. M. Feng

198 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXv

13067; Mengtze, A. Henry 11404; Ping-pien hsien, H. T. Tsai 55328, 55345,
60197, 60967, 61020, 61395, 61744, 61778, and 62446.

The description of this species was based on a flowering specimen col-
lected by A. Henry (10661) in southeastern Yunnan at an altitude of
2100 meters. The largest leaf of that material measures only 9 cm. long
and is rounded at the base. Additional material from the same general
area exhibits variations in the size and shape of the leaves and in the habit
of the plant. As we now know it, the larger leaves of this species measure up
to 15 cm. long and 7 cm. wide and are oblong-elliptic in shape. Tsai 55328,
also a flowering specimen, even possesses an evergreen habit. In south-
eastern Yunnan the plant occurs at altitudes as low as 1300 meters, in
ravines, on rocky crevices, or along the streams of the mixed forest zone.
It is usually a shrub 2-3 m. high but occasionally appears scandent. The
green flowers appear in late May. The specimen collected in early Sep-
tember has very small young fruit, the one collected in mid-October has
fruit reaching mature size, and that collected in November has fruit de-
hiscent by apical slits. The mature fruit is globose, 12 mm. in diameter,
with a rough and woody pericarp.

Craibiodendron kwangtungense, sp. nov.

Craibiodendron stellatum sensu Merrill in Lingn. Sci. Jour. 7: 319. 1931, non
(Pierre) W. W. Smith 1914.

Arbor sempervirens, 10-12 m. alta, ramulis glabris, lenticellis obscuris;
foliis alternis, coriaceis, olivaceo-brunneis, supra nitidis subtus opacis,
ellipticis vel lanceolatis, 6-8 cm. longis, 1.8—3 cm. latis, integris, utrinque
attenuatis, basi acutis vel cuneatis, apice acutis, obtusis, raro breviter
acuminatis, costa supra impressa, subtus elevata, nervis lateralibus 18-22
paribus, supra evidentibus, subtus prominentibus, margine anastomosanti-
bus, reticulatis distinctis, stipulis obsoletis, petiolo 8-10 mm. longo; in-
florescentiis racemosis simplicibus, axillaribus, rhachibus 4-5 cm. longis,
minute puberulis, bracteis lanceolatis, 2 mm. longis, ciliatis, deciduis;
pedicellis 2-3 mm. longis, furfuraceis puberulisque, prophyllis 2, submedio
instructis; calycibus cyathiformibus, 2-3 mm. diametro, sparse puberulis,
lobis 5, rotundatis, ciliatis; corolla breviter campanulata, glabra; stamini-
bus 10, inclusis, filamentis glabris, antheris basi subsaccatis, dorso muticis;
ovario globoso, 1 mm. longo, 5-loculari, stylo columnari, 2 mm. longo;
capsulis depresso-globosis, 14 mm. longis, 18 mm. diametro, profunde 5-
angulatis, pericarpiis ligneis; seminibus in loculo quoque 12-14, sub-
ovoideis, leviter compressis, 2 mm. longis, 2 mm. latis, alis obliquis, 10 mm.
longis, 8 mm. latis, rugosulo-striatis.

CHINA: Kwangtung: Ting-wu-shan, Y. Tsiang 792 (TYPE, flower),
1533, 1547; Kwangsi: south of Nan-ning, Seh-feng-dar shan, R. C. Ching
§293 (TYPE, fruit).

Craibiodendron kwangtungense is a tree 14 meters high with a trunk
30 cm. in diameter. It occurs in southwestern Kwangtung and southeastern
Kwangsi at an altitude of six hundred meters. By the lanceolate leaves,

1954] HU, NOTES ON THE FLORA OF CHINA, III 199

attenuated at both ends, and the glabrous corolla, it can be distinguished
from C. stellatum (Pierre) W. W. Smith, which has oblong leaves rounded
at the apex, and pubescent corolla. It is also related to C. henryi W. W.
Smith, which has long acuminate leaves, paniculate racemes, smaller fruits
which are only 8 mm. long and 10 mm. in diameter.

Craibiodendron kwangtungense var. frutescens, var. nov.

Frutex 2-3 m. altus, ramulis puberulis; foliis oblongo-ellipticis, 5-6 cm.
longis, 2.5—3.5 cm. latis, apice obtusis vel abrupte brevi-acuminatis, racemis
puberulis simplicibus, axillaribus.

CHINA: Kwangtung: Ting-wu-shan, W. Y. Chun 6363. Kwangsi:
Shang-sze, Shap-man-taai shan, W. T. Tsang 22252 (TYPE).

This variety differs from the typical C. kwangtungense in habit, being
a shrub 2-3 meters high; also it has broader leaves and more pubescent
racemes.

Ligustrum subsessile, sp. nov.

Frutex, ramulis robustis, teretis, cineraceis, triannis 5 mm. diametro,
longitudinaliter minute rimulosis, lenticellis orbicularibus, cicatricibus
foliorum semicircularibus, elevatis, hornotinis 2-3 mm. diametro, inter-
nodiis 2-3 cm. longis, + striatis, in triis sparse pilosis, ceterum glabris, lenti-
cellis conspicuis; foliis subsessilibus, integerrimis, subcoriaceis, ovato-ob-
longis vel oblongis vel raro suborbicularibus, utrinque rotundatis, 4—9 cm.
longis, 34.5 cm. latis, glabris, subtus punctatis, costa supra plana, subtus
leviter elevata, nervis lateralibus 4 usque 9 paribus, supra obscuris, subtus
evidentibus, petiolo 1-2 mm. longo, glabro; floribus ignotis; infructescentiis
paniculatis, paniculis compactis, subcylindraceis, 4-6 cm. longis, 2—4 cm.
diametro, pedunculis 1.5—2 cm. longis, sparse puberulis, rhachibus sub-
quadrangularibus, in striis sparse puberulis, axibus secundariis 5-15 mm.
longis, sparse minute puberulis, pedicellis 1-2 mm. longis, glabris; sepalis
persistentibus 3 mm. diametro, lobis 4, rotundatis, glabris; fructibus
oblongo-subglobosis, 5-7 mm. longis, 4-6 mm. diametro.

CHINA: Kiangsi: [Hsiu-shiu], Hwang-lung shan, Nung-lung temple, Y.
K. Hsiung 5629 (TYPE).

This species has been reported to be a common shrub in thickets along
the streams of the Kiangsi-Hupeh-Hunan border. The specimen cited
above was collected in late August. The fruits are still too young for the
study of the seed characters. I have not been able to match it with any
Ligustrum in our collection. The general appearance of the compact in-
florescences and of some of the smaller leaves resembles that of those
shown in Hooker’s illustration (Bot. Mag. 123: pl. 7519. 1897) for Ligus-
trum coriaceum Carriére, a species published on the basis of cultivated
plants introduced to European gardens by Robert Fortune, reportedly from
Japan. But specimens from European and Japanese gardens (such as those
from Hort. Vilmorin of France, the Royal Botanic Gardens at Kew in Eng-

200 JOURNAL OF THE ARNOLD ARBORETUM _ [vor. xxxv

land, and the Botanical Garden of Tokyo) which match Carriére’s and
Hooker’s descriptions and illustrations, all have smaller leaves, the smallest
ones being 1.5 cm. long, 1 cm. wide, and the largest ones being 4 cm. long,
3 cm. wide. They all have comparatively longer petioles which are 3—7 mm.
long. In comparing them with Hsiung’s collection from Kiangsi, I conclude
that the latter, with its subsessile large leaves, is specifically distinct.

ARNOLD ARBORETUM,
HARVARD UNIVERSITY.

EXPLANATION OF THE PLATES

PLATE I

Fic. 1. A habit sketch of a fruiting branch of Tinospora szechuanensis show-
ing the basal lobes of the leaf pointing outward. Fic. 2. A habit sketch of a fruit-
ing branch of Tinospora imbricata showing the imbricate basal lobes of the leaf.
Fic. 3. A habit sketch of a staminate flowering branch of Tinospora henryi with
a separate flower enlarged 5 times. Fic. 4. A habit sketch of a leafy branch of
Tinospora yunnanensis and a staminate flowering branch showing the inflores-
cences on old growth, with a separate staminate flower and a smaller outer and
a larger inner sepal enlarged 5 times. Fic. 5. A habit sketch of a fruiting branch
of Tinospora intermedia with a separate staminate flower enlarged 5 times.
Fic. 6. A habit sketch of a fruiting branch of Tinospora craveniana with a sep-
arate staminate flower enlarged 5 times.

PLATE II

Fic. 1. A habit sketch of Craibiodendron kwangtungense showing solitary
axillary racemes. Fic, 2. A flower after anthesis (X 10). Fic. 3. Two anthers,
dorsal and sublateral view (x 18). Fic. 4. A fruit (X 1%). Fic. 5. A seed
(X 5).

Jour. ARNOLD Ars. VoL. XXXV PLaTE I

ZX

SPECIES OF TINOSPORA

Jour. ARNoLD Ars. VoL. XXXV Prate II

CRAIBIODENDRON KWANGTUNGENSE Hv

JOURNAL

OF THE

ARNOLD ARBORETUM

VoL. XXXV Jury 1954 NUMBER 3

THE CYPERACEAE COLLECTED IN NEW GUINEA
BY LJ. BRASS, IV."

S. T. BLAKE

With one plate and two text-figures

THIS PAPER COMPLETES the account of the specimens collected by Mr.
L. J. Brass during the Archbold Expeditions between 1933 and 1939. As
in the previous contributions (Jour. Arnold Arb. 28: 99-116, 207-229.
1947; 29: 90-102. 1948), some other specimens are cited. To the ac-
knowledgements made in earlier contributions, I wish to add my thanks to
Prof. H. Humbert, Dr. O. Hagerup, Dr. A. Hassler and Dr. H. J. Lam
for the loan of types and other material from the herbaria of Paris, Copen-
hagen, Lund and Leiden respectively, and to Mr. R. a a for the
loan of all specimens of Scleria in the Sydney Herbar

The genera treated here are Scirpus, Fimbristylis, "Scleria, Diplacrum
and Uncinia, and there are some additional notes on genera previously
treated. The treatment of the different genera is somewhat unequal, par-
ticularly as to the citation of synonyms, but in all cases it is based on
fairly extensive revisional work on the Malaysian and Australasian species.
The delayed appearance of this paper is due partly to the necessity of
examining certain types before some groups of species could be deter-
mined satisfactorily.

Herbaria are indicated by the following abbreviations: Brisbane, BRI;
Canberra, cANB; Copenhagen, c; Leiden, L; Lund, tp; Melbourne, MEL;
Paris, P; Sydney, Nsw.

Scirpus Linnaeus
Scirpus ternatanus Reinw. ex Miq. FI. Ind. Bat. 3: 307. 1859; Kikenth.
Bot. Jahrb. 69: 259. 1938; Ohwi, Bot. Mag. Tokyo 56: 204. 1942
Scirpus chinensis Munro in Seem. Bot. Voy. Herald 423. 1857; Valck.
Suring. Nova Guin. Bot. 8: 705. 1912; Ridl. Trans. Linn. Soc. Il. Bot;. 9:
242. 1926; non Osbeck 1753.

* Botanical oe of the Richard Archbold Expeditions. See Jour. Arnold Arb.
29: 90-102. 194

204 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

NETHERLANDS NEW GUINEA: Bele R., 18 km. NE. of Lake Hab-
bema, Brass 11472, Nov. 1938, alt. 2200 m., large clumps in moist situations
on grassy, formerly cultivated slopes; Balim R., Brass 11677, Dec. 1938, alt.
1600 m., colonizing loose sand and stones from a landslip; 9 km. NE. of
Lake Habbema, Brass 10993, Oct. 1938, alt. 2650 m., plentiful in Equisetum
cover on landslips, large clumps + 75 cm. high; 9 km. NE. of Lake Habbema,
Brass 10883, Oct. 1938, alt. 2650 m., open banks of a stream in forest.

The range of this species extends north and west through Malaysia to
India, China and Japan

? Scirpus strobolinus Roxb. Hort. Beng. 6. 1814, nomen nudum, FI.
Ind. ed. Carey & Wall. 1: 223. 1820, Fl. Ind. ed. Carey 1: 219. 1832.
PUA: Western Division: Gai Lower Fly R. Sag bank), Brass

8304, Nov. 1936, loose sand on open esa, not comm

The specimens are in flower only and the identification is rather uncer-
tain, though they appear to belong here rather than to S. maritimus L.,
S. fluviatilis A. Gray or S. paludosus A. Nels. Scirpus strobolinus is other-
wise known from different parts of Asia, extending south to Assam and
Pegu.

Scirpus mucronatus L. Sp. Pl. 50. 1753; K. Schum. in K. Schum. &
Lauterb. Fl. Deutsch. Schutzgeb. Stidsee 195. 1901; Valck. Suring.
Nova Guin. Bot. 8: 704. 1912; Kiikenth. Bot. Jahrb. 59: 51. 1924,
69: 259. 1938.

NETHERLANDS NEW GUINEA: Balim R., Brass 11804, Dec. 1938,
alt. 1600 m., occasional in ditches and grassy pools.

PAPUA: Western Division: Junction of Black and Palmer Rivers,
Brass 6943, June 1936, common on silt-covered gravel banks in river (det.
Uittien). Central Division: Urunu, Vanapa Valley, Brass 4810, July-
Aug. 1933, alt. 1900 m., plentiful in small swamps on open slopes of valley;
Koitaki, Carr 12283, May 1935, alt. 1500 ft., swamp in open savannah land,
c. 4 ft. tall (herb. Canberra).

A species widely spread over the warmer parts of the Eastern Hemis-
phere.

Scirpus clemensiae (Kukenth.) Ohwi, Bot. Mag. Tokyo 56: 203. 1942.
— clemensiae (Kiikenth.) Kiukenth. Mitteil. Thuring. Bot. Ver. N. F.
1943.

Scirpus ‘sieves L. subsp. clemensii Kiikenth. Bot. Jahrb. 69: 259.

NETHERLANDS NEW GUINEA: Lake Habbema, Brass 9069, August
1938, alt. 3225 m., plentiful in sandy marginal shallows of lake, tufts 40-60
cm. high; Lake Habbema, Brass 9439, August 1938, alt. 3225 m., abundant
in marginal shallows of lake.

Known also from North-East New Guinea, whence it was originally de-

1954] BLAKE, CYPERACEAE COLLECTED IN NEW GUINEA 205

scribed. The binary combination was made independently by Ohwi and
Kiikenthal. Ohwi spelled the epithet Clemensii, as published in the ternary
combination. When Kiikenthal made the binary combination, he cited
“Scirpus Clemensiae Kiikenth., comb. nova. — Sc. mucronatus L. subsp.
Clemensii Kiikenth. in Bot. Jahrb. 69.2. (1938) 259.” It would appear
that the spelling Clemensii in the first paper was “an unintentional ortho-
graphic error” which was corrected in the later paper, as permitted by
Art. 70 of the International Rules, and I have adopted the emended spell-
ing. The plant was evidently dedicated to Mrs. M. S. Clemens.

Scirpus validus Vahl, Enum. 2: 268. 1806.

PAPUA: Western Divison: Gaima, Lower Fly R. (east bank), Brass
8305, Nov. 1936, gregarious on open sandy foreshores.

New for Papua; widely spread in the countries around the Pacific Ocean
and in the Americas generally. Specimens from New Guinea were formerly
identified with the Eurasian S. Labernaemontani Gmel. by Kikenthal in
Engl. Bot. Jahrb, 59: 51. 1924, and by Ohwi in Bot. Mag. Tokyo 56: 203.
1942. I have accepted Beetle’s arrangement of this group of species in
Amer. Jour. Bot. 28: 691-700. 1941.

Scirpus grossus L. f. Suppl. 104. 1781; Valck. Suring. Nova Guin. Bot.
$3 705. 1912:

PAPUA: Western Division: Gaima, Lower Fly R. (east bank), Brass
8311, Nov. 1936, co-dominant with no. 8312 ( = Cyperus malaccensis Lam.)
in extensive sedge communities on open sandy foreshores (det. Uittien).

New for Papua. The species ranges from India to NE. Queensland, and
in New Guinea was previously known only from Netherlands New Guinea.

Scirpus crassiusculus (Hook. f.) Benth. Fl. Austral. 7: 326. 1878;
Kikenth. Bot. Jahrb. 69: 258. 1938.

Isolepis crassiuscula Hook. f. Fl. Tasm. 2: 86, ¢. 143. 1860.

NETHERLANDS NEW GUINEA: Lake Habbema, Brass 9324, Aug.
1938, alt. 3225 m., gregarious on sand bars in grassland stream; Mt. Wil-
helmina, 4 km NE. of top, Brass & Meyer-Drees 9984, Sept. 1938, alt.
3660 m. " submerged green masses in shallows of a lake.

PUA: Central Division: Mt. Albert Edward, Brass 4300, May-July
1933, alt. 3810 m., submerged in large masses on shallows of an alpine lake.

New for both Netherlands New Guinea and Papua. Recorded by Ku-
kenthal, l.c., for North-East New Guinea, but he credited the authorship
of the combination under Scirpus to Hooker f., l.c. Bentham, l.c., also
credited the combination to Hooker. The species is elsewhere known from
SE. Australia (including Tasmania) and New Zealand.

The sheet seen of Brass & Meyer-Drees 9984 has but one spikelet, too
immature for dissection, but the facies of the plant is of this species.

206 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

Scirpus merrillii (Palla) Kikenth. ex Merr. Enum. Philipp. Fl. Pl. 1:
117. 1925; S. T. Blake, Proc. Roy. Soc. Queensl. 58: 38. 1947.
Schoenoplectus merrillii Palla in Kneucker, Cyperaceae (excl. Carices) et
er teiigsy oo 8: mr. 223. 1911, in Allgem. Bot. Zeitschr. 17:
Beil.

NETHERLANDS NEW GUINEA: Lake Habbema, Brass 9238, Aug. 1938,
alt. 3225 m., associated with mosses, etc., on open seepages.

PAPUA: Central Division: . Albert Edward, Brass 4364, May-July
1933, alt. 3680 m., plentiful on wet es of a small alpine stream; Murray Pass,
Wharton Range, Brass 4725, June—Sept. 1933, alt. 2840 m., common, wet banks
of grassland streams. Eastern Division: Mt. Dayman, W. Armit in 1894
(MEL).

New for New Guinea, though it is very likely this species which has
been recorded as S. inundatus (R. Br.) Poir. by Kukenthal, Bot. Jahrb.
69: 258. 1938, from North-East New Guinea and by Ohwi, Bot. Mag.
Tokyo 56: 203. 1942, from Netherlands New Guinea. It differs from the
polymorphic S, inundatus by the constant development of a filiform
branched rhizome, well-developed leaves often overtopping the culm, more
or less emarginate glumes nearly as broad as long and scarcely if at all
mucronate, and with the nut nearly as long as the glume; also it is almost
invariably a much smaller, very slender, more or less mat-like plant with
mostly only one, rarely twe or three spikelets, and the involucral bract is
commonly elongated. I have seen other specimens from the Philippine
Islands including an isotype (Merrill in Kneucker Cyperaceae et Juncaceae
exsiccatae 8, nr. 223), south-east Queensland, New South Wales, Victoria,
Tasmania and New Zealand.

Scirpus clarkei Stapf, Trans. Linn. Soc. II, Bot. 4: 244. 1894.

Scirpus pulogensis Merr. Philipp. Jour. Sci. 5 (C): 333. 1910; syn. nov.

Scirpus pakapakensis Stapf, Jour. Linn. Soc. Bot. 42: 174. 1914; syn. nov.

Scirpus subcapitatus Thw. var. triangularis Kikenth. Bull. Jard. Bot. Buitenz.

sér. III, 16: 301. 1940; syn. nov

Scirpus subcapitetus Thw. forma iia Kikenth., , syn. no

— clarkei asa var. pakapakensis (Stapf) Beetle, Amer. a Bot. 33:

946; syn.

PAPUA: Central Division: Mt. Albert Edward, Brass 4315, May-July
1933, alt. 3680 m., forest glades and grassland slopes, common, few plants fertile;
Mt. Knutsford, W. MacGregor in 1889 (BRI, MEL); summit of the Owen Stanley
Ranges, W. MacGregor in 1889 (MEL).

New for New Guinea; elsewhere known from the Philippine Islands,
Borneo and Sumatra

F. Mueller, sti Roy. Soc. Vict. n.s. 1 (2): 35. 1889, referred Mac-
Gregor’s specimens to S. cespitosus L. (as S. caespitosus) with the remark:
“Should nevertheless this plant, as a variety or perhaps even as a species,
require separation from the genuine S. caespitosus, then the name heleo-
charoides would be an apt one.”’ Some of the specimens are labelled in

1954] BLAKE, CYPERACEAE COLLECTED IN NEW GUINEA 207

Mueller’s handwriting: Scirpus caespitosus Linné var. heleocharoides. This
ternary combination has never been validly published, for the phrase
quoted cannot be taken as constituting publication of any combination.

The specimens from New Guinea do indeed closely resemble specimens
of S. caespitosus L., but the leaf-sheaths are fewer, tighter and more rigid,
the hard culms are more or less trigonous and less furrowed, the two lower-
most glumes are much shorter and more rigid than the others and only
shortly pointed, and the nut is narrower. Occasionally also the inflores-
cence consists of two spikelets. They also resemble some from the Philip-
pine Islands (1soTyPE of S. pulogensis), Borneo (Mt. Kinabalu, the type-
locality of S. clarkei and S. pakapakensis) and Sumatra (general locality
of the types of S. subcapitatus var. triangularis and S. subcapitatus forma
rigidus). Scirpus clarkei will probably prove to be conspecific with S.
subcapitatus Thw. from Ceylon and southern India; of this I have seen
only one sheet, an isotype, but it is in flower only.

Beetle, in the paper quoted above, recognised five taxa in a group which
he described as Scirpus sect. Paucispicatae Beetle, 1.c., 664. He distin-
guished S. clarkei from S. subcapitatus on differences in the number of
spikelets and scabridity of the mucro to the leaf-sheaths and lowermost
glumes. Scirpus pakapakensis was distinguished as a variety of S. clarkei
by the spikelet not solitary and the stems more or less triangular. He also
saw only a single sheet in flower (an isotype) of S. subcapitatus. The num-
ber of spikelets and shape of the culms are certainly variable, but on the
material seen the character of scabridity seems constant. I have not seen
material of the other species admitted by Beetle.

F. Mueller, ].c., remarked that ‘Another Scirpus is contained in the col-
lections, as gathered on Mt. Knutsford and Mount Musgrave; it is an
aged state of fructification, and may perhaps belong to the variety fluvia-
tilis of S. maritimus.”” These specimens are of Mapania Moseleyi C. B.
Clarke.

Fimbristylis Vahl

No satisfactory account of this genus as a whole has ever been pub-
lished. The latest account which has any claim to be considered as a
general treatment of the genus is Boeckeler’s uncritical descriptions of. the
forms represented in the Berlin Herbarium in Linnaea 37: 2—56. 1871,
38: 384-398. 1874. In Kew Bull. Add. Ser. 8: 107-109. 1908 is a list of
species as accepted and arranged by C. B. Clarke, but one of the tragedies
in botany is that his extensive manuscript on the family was never pub-
lished.

The study of the New Guinea collections, the results of which appear
on the following pages, was based chiefly on the revision of the Australian
species which I commenced in 1932, but which is not yet completed, owing
to the numerous difficulties involved, due partly to the need of critically
comparing some Australian forms with others described from other parts
of the world, of which types were not readily accessible. Little was previ-

208 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

ously known of the Fimbristylis-flora of southern New Guinea, to which
Mr. Brass has added a remarkable number of Australian forms.

For convenience, I have arranged the species under the four sections
proposed by Bentham, FI. Austral. 7: 298-9. 1878, an arrangement which
has been fairly generally followed. The type-species of the genus, F.
dichotoma (L.) Vahl, was arranged under Dichelostylis Benth., |.c., 299,
but Boeckeler, op. cit. 3. 1841, had previously proposed the name Exufim-
bristylis for the section containing this species.

Sect. Heleocharoides Benth.

Fimbristylis setacea Benth. Lond, Jour. Bot. 2: 239. 1843; Valck.
Suring. Nova Guin. Bot. 8: 702. 1912.

Fimbristylis acuminata (Retz.) Vahl var. minor Miq. FI. Ind. Bat. 3: 314. 1859.
shell a acuminata (Retz.) Vahl var. setacea (Benth.) Benth. Fl. Austral.
878.

Pimbrisi aged Seen bs var. setacea (Benth.) Kukenth. Bot.
Jahrb. 59: 47. 1924, 69: 257.

Tsolepis Bein Steud. Synops. ae 100. 1855.

PAPUA: Western Division: Lake Daviumbu, Middle Fly R., Brass
7531A, August 1936, savannahs, abundant on hard-pans and swamp margins
(det. Kikenthal as Fimbristylis acuminata (Retz.) Vahl var. setacea Bth.);
Daru Island, Brass 6245, March 1936, abundant in flattened tufts on damp soil
in savannah-forests

New for Papua. The species ranges from Tropical Asia to northern
and north-eastern Australia. Kiikenthal, 1924, l.c., made a new combina-
tion F. acuminata (Retz.) Vahl var. setacea (Benth.) Kiikenth., evidently
overlooking Bentham’s much earlier combination. But if this form is
treated as a variety of F. acuminata, then the legitimate trinomial would
be F. acuminata var. minor Migq., |.c. The species is very close to F. acu-
minata, differing chiefly in being smaller in all its parts.

Fimbristylis nutans (Retz.) Vahl, Enum. 2: 285. 1806.
Scirpus nutans Retz. Observ. 4: 12. 1786.

PAPUA: Western Division: Lake Daviumbu, Middle Fly R., Brass 7843,
Sept. 1936, common on wet grass plains; Wuroi, Oriomo R., Brass 5750, Jan-
March 1934, alt. 10-30 m., very plentiful on open savannah.

Not previously recorded for New Guinea, though the species is known
to extend from northern and north-eastern Australia to Malaya and China.

Fimbristylis tetragona R. _ ta 226. 1810; Kukenth. Mitteil.
Thuring. Bot. Ver. N. F. 50:

Fimbristylis cylindrocarpa Kunth, Enum. 2: 222. 1837.
Fimbristylis arnottii Thw. Enum. Pl. Zeyl. 348. 1864.

1954] BLAKE, CYPERACEAE COLLECTED IN NEW GUINEA 209

Fimbristylis xyroides Arnott ex Thw., l.c., in syn., nomen nudum.

Fimbristylis abjiciens Steud. Synops. Cyper. 107. 1855.

Scirpus tetragonus (R. Br.) Poir. Encycl. Suppl. 5: 98. 1817.

Mischospora efoliata Boeck. Flora 43: 113. 1860.

PAPUA: Western Division: Lake Daviumbu, Middle Fly R., Brass 7844,
Sept. 1936, tufted on wet grass plains, plant bluish (det. Svenson).

New for New Guinea; also in southern and eastern Asia and northern
Australia.

Fimbristylis dictyocolea sp. nov. (Sect. Heleocharoides Benth.). Fic. 1.

Herba perennis, caespitosa, circiter 20-40 cm. alta, fere aphylla. Culmi
stricti, erecti, setacei, pluristriati, obscure quinquangulares, glabri, laeves
vel minute asperuli, basi haud incrassati. Folia basalia perpauca, setacea,
triquetra, lateraliter compressa, glabra, laevia, brevia vel culmum fere adae-
quantia; folia caulina basi culmi inserta, ad vaginas arctas ore oblique
sectas antice late hyalinas tandem reticulatim fissas redacta. Inflorescentia
unispiculata, quasi ebracteata. Spicula erecta, pallida, oblonga vel ellip-

imbricatae, oblongae, apice obtusa rotundatae, muticae, omnino glabrae,
dorso late coriaceae uninerves nec carinatae, lateribus membranaceae cel-
lulis parvis breviter oblongis, marginibus etiam apice hyalinae, 4-5 mm.
longae, 1-3 imae vacuae crassiores. Stamina 3; antherae lineares, promi-
nule apiculatae, circiter 2 mm. longae. Stylus tenuis, complanatus, basi
dilatatus, marginibus minute ciliolatus, circiter 3-3.5 mm. longus; stig-
mata 3, brevia. Nux straminea, jneidule obovoidea, late umbonulata, vix
stipitata, trigona, leviter tricostulata, minute reticulata verrucosaque cellu-
lis extimis minimis distinctis hexagonis, 1 mm. longa, 0.7 mm. lata.

=

4

=

SS

eared
J

ans >

one ape
SoS

—— Se
Seape

—

Fic. 1. Fimbristylis dictyocolea S. T. Blake: a. upper part of culm with
spikelet, X 3; b. middle part, and c. upper part of leaf-sheath, & 3; d. glume,
~ 10: €; ae of side of glume, < 40; f. style, X 10; g. nut, * 10: h. trans-
verse section of nut; i. surface of nut, X 40. Figures from type.

210 JOURNAL OF THE ARNOLD ARBORETUM [| VOL. XXXV

PAPUA: Western Division: Tarara, Wassi Kussa R., Brass 8400, Dec.
1936, abundant on gray soil flat, savannah-forest; Mabaduan, Brass 6553 (TYPE),
April 1936, common in shallow rain-pools in savannah-forests.

Brass 6553 had been determined by Svenson as F. pauciflora R. Br.,
while both it and Brass 8400, which is in flower only, were cited by Kuken-
thal, Mitteil. Thiring. Bot. Ver. N. F. 50: 9. 1943 as F. cardiocarpa F.
Muell. The species is certainly allied to F. pauciflora R. Br., and but for
the larger size resembles it rather closely in the nut and style. It differs,
however, in the coarser habit, the leaf-sheaths disintegrating into fine
reticulate fibres, the larger and relatively much broader spikelet, and the
rather larger, more oblong, more obtuse single-nerved muticous glumes.
It is also allied to the Australian F. simplex S. T. Blake, but the latter has
rather prominently swollen culm-bases, sheaths splitting into straight
fibres, 3—5-nerved brownish glumes, entirely glabrous rather stout style
and more shining nut cuneate at its base. On the other hand F. cardio-
carpa F, Muell. is an entirely different plant, differing in almost every
respect except for the solitary spikelet and three stigmas.

Fimbristylis pauciflora R. Br. tt 225. 1810; Kukenth. Mitteil.
Thuring. Bot. Ver. N. F. 50: 9.
Fimbristylis filiformis (Nees) Kunth, Enum. 2: 221. 1837.
cirpus pauciflorus (R. Br.) Poir. Encycl. Suppl. 98. 1817.

Trichelostylis filiformis Nees in Wight, Contrib. 102. 1834.

PAPUA: Western Division: Lake Daviumbu, Middle Fly R., Brass
7531, August 1936, savannahs, abundant on hard-pans and swamp margins;
Gaima, Lower Fly R. (east bank), Brass 8357, Nov. 1936, covering patches of
sour soil in savannah-forest (det. Uittien); Wuroi, Oriomo R., Brass 5817, Jan—
March 1934, alt. 10 m., plentiful, shaded ground on a clearing in savannah, flat
spreading and rather fleshy.

Brass 7531 was recorded by Kukenthal, l.c., as new for New Guinea;
previously known from northern and north-eastern Australia, other parts
of Malaysia, and southern and eastern Asia.

Brass 5817 represents what appears to be the usual state in which
three stigmas are present. In Brass 7531 some flowers have two stigmas
only, and flowers with two stigmas are the rule in Brass 8357. Nuts pro-
duced from flowers with three stigmas are always finely 3-ribbed and +
trigonous, though at times distinctly compressed. Three-ribbed nuts have
also been observed produced from flowers with two stigmas, though bi-
convex two-ribbed nuts are more usual. Both trigynous and digynous
flowers have been observed on a collection from Johore, Ngadiman in
Singapore Field No. 36784.

Fimbristylis pumila Benth. Lond. Jour. Bot. 2: 239. 1843, from Am-
boina, was later referred by its author in Fl. Austral. 7: 303. 1878, and
by C. B. Clarke in Hook. f. Fl. Brit. Ind. 6: 633. 1893, to F. pauciflora.
It was described as having a glabrous style with two stigmas and one-
nerved oblong glumes, features which do not well accord with F. pauciflora.

1954] BLAKE, CYPERACEAE COLLECTED IN NEW GUINEA 211

Of the latter I have examined twelve good sets of specimens ranging from
south-east Queensland to the Malay Peninsula, and in all these I find
the style minutely ciliate and the glumes three- to seven-nerved and as
much ovate as oblong. Bentham may have overlooked the minute sparse
ciliation of the style, while the lateral nerves are often close to the keel
of the glume.

Fimbristylis pauciflora is rather \easily recognised by its small, un-
usually narrow, relatively few-flowered spikelets.

As a general rule in Fimbristylis, the number of stigmas is very constant
in each species. Fimbristylis pauciflora shares with F. tetragona R. Br.
the peculiarity of having either two or three stigmas. In F. cymosa R. Br.,
three stigmas occur in the lower part of the spikelet, but in the upper
flowers there are often only two.

Fimbristylis recta F. M. Bail. 3rd Suppl. Syn. Ras: Fl. 80. 1890;
S. T. Blake, Proc. Roy. Soc. Queensl. 58: 44
Fimbristylis xyridis R. Br. var. rigidula Benth. Fl. Austral. 7: 307. 1878.
Fimbristylis stricticulmis Domin in Biblioth. Bot. 20 (85): 452. 1915.
Western Division: Tarara, Wassi Kussa R., Brass 8714, Jan.
1937, savannah-forests, common in grass on ridges

A distinctive species known previously only from northern Australia and
Hammond Island in Torres Strait. It was discussed in some detail by
S. T. Blake, l.c.

Sect. Eufimbristylis Boeck.

paar he ferruginea (L.) Vahl, Enum. 2: 291. 1806; K. Schum.
. Bot. Gart. Mus. Berlin 2: 98. 1898; in Schum. & Lauterb.

Fl. oe Schutzgeb. Suidsee 197. 1901; Valck. Suring. Nova Guin.
Bot. 8: 702. 1912; Kiikenth. Bot. Jahrb. 59: 48. 1924, 69: 258. 1938.

Fimbristylis brevifolia R. Br. Prodr. 228. 1810.

Scirpus ferrugineus L. Sp. Pl. 74. 17

Scirpus brevifolius (R. Br.) Poir. Ency cl. Suppl. 5: 99. 1817.

PAPUA: Western Division: Gaima, Lower Fly R. (east bank), Brass
8308, Nov. 1936, common on open sandy foreshores; Gaima, Lower Fly R. (east
bank), Brass 8313, Nov. 1936, common on sandy foreshores; Upper Wassi Kussa
R. (left branch), Brass 8642, Jan. 1937, abundant in brackish swamp; Daru
Island, Brass 6212, Feb. 1936, gregarious in limited pure stands, associated with
Zoysia pungens on saline marshy ground; Daru Island, Brass 6286, March 1936,
scattered in marginal shallows of large swamp. Central District: Kerema,
Brass 1224, March 1926, on banks of tidal creeks; Arva R., Carr 11436, Feb.
1935, sea-level, marshy places near beach, about 18 in. tall (CANB; very young!).

Brass 1224 and Carr 11436 are cited by Kiukenthal, 1938, l.c.; Brass
8313 was received as having been determined by Kukenthal as F. ferru-
ginea Vahl var. tristachya (R. Br.) Domin. It represents a not uncom-
mon state of the species with the inflorescence reduced to one or two
spikelets, commonly seen on young or small plants. It seems pointless to

212 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXXVv

give such reduced states taxonomic status. I doubt very much whether
it truly represents F. tristachya R. Br., and in any case, if this be re-
garded as a variety of F. ferruginea, the legitimate ternary combination
would be F. ferruginea (L.) Vahl var. foliata Benth. Fl. Austral. 7: 312.
1878, the epithet foliata having priority in the required position. The other
specimens were received as having been determined by Uittien. The
species is widely spread in the warmer parts of the wor

Fimbristylis marianna Gaud. in Freyc. Voy. 413. 1826.

Fimbristylis maxima K. Schum. in Hollr. Fl. Kaiser Wilhelmsl. 24. 1889, in
Schum. & Lauterb. Fl. Deutsch. Schutzg an Siuidsee 196. 1901; Valck.
Suring. Nova. Guin. Bot. 8: 702. 1912; syn. n

PAPUA: Western Division: Lake as ule: Middle Fly R., Brass

7847, 7878, Sept. 1936, common on wet grass plains (both det. Kiikenthal) ;
Wuroi, Oriomo R., Brass 5734, Jan—March 1934, alt. 10-30 m., common all
through savannahs.

ORTH-EAST NEW GUINEA: Augusta R., Hollrung 836, in 1877 (MEL;
duplicate TYPE of F. maxima).

New for Papua; elsewhere known from North-East New Guinea, Mari-
anne Islands and Philippine Islands. I have relied on Kiikenthal’s deter-
mination of Brass 7847 and 7878 for my concept of the species and the
consequent reduction of F. maxima to synonymy.

Fimbristylis aestivalis (Retz.) Vahl, Enum. 2: 288. 1806.
Fimbristylis aestivalis (Retz.) Vahl f. glabra Kikenth. Bot. Jahrb. 59: 49.
1924; syn. nov.
Scirpus aestivalis Retz. Obs. 4: 12. 1786.
PAPUA: Western Division: Penzara, between Morehead and Wassi
Kussa Rivers, Brass 8438, Dec. 1936, wet shaded banks of a permanent water-
hole (det. Kiikenthal).

Brass’s plant, representing the usual pubescent state, forms the first
record of the species for Papua. Kukenthal, l.c., records as f. glabra a
glabrous form from North-East New Guinea. Elsewhere the species is
known from Australia to southern and south-eastern Asia.

Fimbristylis annua (All.) R. & S. Syst. 2: 95. 1817.

Scirpus annuus All. Fl. Pedem. 2: 227. 1785.

PAPUA: Western Division: Wuroi, Oriomo R., Brass 6069, Jan—
March 1934, alt. 30 m., uncommon tufted species on savannah; Daru Island,
Brass 6248, March 1936, common on damp soil in savannah-forest (det. Kiiken-
thal as F. diphylla [Retz.] Vahl f. tomentosa [Vahl] Kiikenth.); Daru Island,
Brass 6369, March 1936, plentiful in drainage ditches in savannah-forest.
Central Division: Baroka, Nakeo District, Brass 3732, April 1933.
alt. 30 m., common, damp savannah flats, plant grayish.

For discussion, see under F.. dichotoma.

1954] BLAKE, CYPERACEAE COLLECTED IN NEW GUINEA 2S
Fimbristylis dichotoma (L.) Vahl, Enum. 2: 287. 1806.

Fimbristylis diphylla (Retz.) Vahl, Enum. 2: 289. 1806; K. Schum. in Warb.
Bot. Jahrb. 13: 265. 1891, Notizbl. Bot. Gart. Mus. Berlin 1: 47. 1895,
2: 97. 1898, in Schum. & Lauterb. Fl. Deutsch. Schutzgeb. Siidsee 196.
1901; Valck. Suring. Nova Guin. Bot. 8: 702. 1912; Ridl. Trans. Linn.
Soc. II, Bot. 9: 242. 1916.

Fimbristylis polymorpha see ae 37: 14. 1871, in Engl. Forschungsr.
= azelle 4 (1): 17

Fimbasiytie novae- ae ed Bot. Jahrb. 5: 93. 1884, in Engl. For-
schungsr. S. M. S. Gazelle 4 (1): 11. 1889; K. Schum. Bot. Jahrb. 9: 195.
1888, Notizbl. Bot. Gart. Mus. Berlin 2: 97. 1898, in Schum. & Lauterb. FI.
Deutsch. ee Sudsee 196. 1901; Valck. Suring. Nova Guin. Bot.
8: 703. 1912

ae annua (All.) R. & S. var. diphylla (Retz.) Kiikenth. Bot. Jahrb.
9: 1924, nomen ex C. E. C. Fischer, Fl. Pr. Madras 1658. 1931.

ce ee NEW GUINEA: Balim R., Brass 11731, Dec. 1938, alt.
m., common on grassy deforested slopes (leaves ae hairy!) ;
ate R., Brass 11816, Dec. 1938, alt. 1600 m., common on sandy, long-de-
forested slopes, erect tufts 60-80 cm. high (leaves hairy!) ; Bele R., 18 km.
. of Lake Habbema, Brass 11489, Nov. 1938, alt. 2200 m., common on
grassy, ies: cultivated slopes, small clumps 60-80 cm. high (leaves gla-
brous!

PAPUA: Western Division: Lake Daviumbu, Middle Fly R., Brass
7927, Sept. 1936, savannah, abundant on seepage area (plant almost entirely
glabrous!; det. Uittien as F. annua | All.] R. & S.); Lake Daviumbu, Middle
Fly R., Brass 7521, August 1936, savannah hard-pans (almost entirely glabrous! ;
det. Uittien as F. annua [All.] R. & S.); Gaima, Lower Fly R. (east bank)
Brass 8265, Nov. 1936, open savannah-forest, plentiful in firm-set soil, inflores-
cence dark brown (leaves slightly hairy!; det. Uittien as F. annua [All.] R. &
S.); Gaima, Lower Fly R. (east bank), Brass 8307, Nov. 1936, plentiful on
open sandy Sees (almost glabrous!; det. Uittien as F. annua [All.] R. &
S.). Central Division: Kanosia, Carr 11034, Jan. 1935, sea-level, open
places ae light shade (glabrous!); Huia, Brass 524, October 1925, coast
sand hills (nearly glabrous!); Baroka, Nakeo District, Brass 3729, April 1933,
alt. 30 m., common, damp savannah flats (nearly glabrous!); Mafulu, Brass
5480, Sept.—Nov. 1933, alt. 1250 m., common on roadside (leaves + hairy!);
Mafulu, Brass 5328, Sept—Nov. 1933, alt. 1250 m., grassy seepages on road-
side, uncommon, plant bluish green, inflorescence erect (leaves distinctly hairy!) ;
Mafulu, C. T. White 600, July-August 1918, alt. ca. 1200 m. (leaves distinctly
hairy!).

Fimbristylis annua (All.) R. & S., F. diphylla (Retz.) Vahl and allied
forms have been a fertile source of difficulty since the time of Linnaeus.
According to C. B. Clarke in Thistleton-Dyer, Fl. Trop. Afr. 8: 416. 1902,
“some closely allied plants, esteemed mere forms of F. diphylla by Kunth
and Boeckler, are here regarded as distinct; even thus narrowed down,
our F. diphylla has 140 names. It should, moreover, be understood that
F. diphylia is so close to the preceding F. dichotoma that different cyperol-
ogists sort the material, as between these two, differently.” On the pre-

214 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

ceding page Clarke cites Scirpus annuus All. (which is Fimbristylis annua
[| All.] R. & S.) as a synonym.

According to C. E. C. Fischer, Kew Bull. 1935: 149-50. 1935, the type
of Scirpus diphyllus Retz. and consequently of Fimbristylis diphylla
(Retz.) Vahl is identical with the type of Scirpus dichotomus L., so that
the plant which has been called F. diphylla (Retz.) Vahl must be called
F. dichotoma (L.) Vahl. Another name thus has to be applied to the
plant which has been passing under the name of F. dichotoma (L.) Vahl,
and according to Fischer, l.c., p. 149, the legitimate combination is Fim-
bristylis bisumbellata (Forsk.) Bubani.*

To my mind, F. annua (All.) R. & S. is conspecific with F. depauperata
R. Br., and I have suggested previously, Univ. Queens]. Papers Dept. Biol.
1(13): 3. 1940, that this is specifically distinct from F. dichotoma (F.
diphylla), differing in the constantly annual habit, the softer and softly
hairy leaves and bracts, the more or less hairy culms and rays, the glumes
often minutely ciliolate at the upper edge, the cells composing the glumes
less distinctly oblong (more nearly square) in shape, and in the relatively
shorter and broader style. Fimbristylis dichotoma has sometimes hairy
leaf-blades and more rarely hairy culms, but the hairs are more rigid
and the blades stiffer than in F. annua. In this restricted sense, F. annua
seems to have been previously unknown from New Guinea.

Fimbristylis dichotoma, or allied forms, has been reported under other
names from New Guinea, apart from those recorded in the synonymy
above. Of some of these I am uncertain of the taxonomic status, and some
seem to be invalid names.

Fimbristylis dipsacea (Rottb.) C. B. Clarke in Hook. f. Fl. Brit. Ind.

Scirpus pied Rottb. Descr. et Ic. 56, ¢. 12, fig. 1. 1773; F. Muell. Pap.
Fi: 2

ee dipsaceum (Rottb.) Desv. tie 1: 21, ¢. 1. 1808.

Isolepis dipsacea (Rottb.) R. & S. Syst. 2: 119.

PAPUA: Western Division: Strickland R., Bdaduerlen in 1885 (prt,
MEL).

This collection, the only one recorded for New Guinea, was recorded by
Mueller, l.c., as Scirpus dipsaceus. The formal transfer of the species to
Fimbristylis was made by C. B. Clarke, l.c., though this author there
cited Benth. in Gen. Pl. 3: 1049. Bentham gave reasons why it should
be placed under Fimbristylis, but did not make the transfer.

Sect. Trichelostylis (Lestib.) A. Gray
Fimbristylis signatus nom. nov.

* From Fischer’s data, it is doubtful if this combination was validly published by
Bubani; I have not seen the publication.

1954] BLAKE, CYPERACEAE COLLECTED IN NEW GUINEA 215

Fimbristylis debilis F. Muell. Fragm. Phyt. Austr. 1: 198. 1859, non Steud.
Syn. Cyper. 109. 1855.
Iriha debilis (F. Muell.) O. Ktze. Rev. Gen. Pl. 2: 753. 1891.
PAPUA: Western Division: Daru Island, Brass 6250, March 1936,
frequent on damp soil in res forest.

New for New Guinea; previously known only from northern Australia.

Fimbristylis eragrostis (Nees & Meyen) Hance, Jour. Linn. Soc.
Lond. 13: 132. 1873; Ktikenth. Mitteil. Thiiring. Bot. Ver. N. F. 50:
9.1943.

Abildgaardia eragrostis Nees & Meyen in Wight, Contrib. 95. 1834.
PAPUA: Western Division: Tarara, Wassi Kussa R., Brass 8405,

Dec. 1936, savannah-forest, common on sour gray soil; Wuroi, Oriomo R.,

Brass 5706, Jan—March 1934, alt. 10-30 m., common on lower savannah ridges.

Central Division: Astrolabe Range, W. E. Armit in 1894-5 (MEL).

South-Eastern Division: Sud-est Island, W. MacGregor in 1889 (MEL).

Brass 8405 was cited by Kiikenthal, l.c., as new for New Guinea. The
species extends northwards to China and southwards to Queensland.

Fimbristylis globulosa (Retz.) Kunth, Enum, 2: 231. 1837; Kukenth.
Bot. Jahrb. 59: 49. 1924, 69: 258. 1938; Ohwi, Bot. Mag. Tokyo
S07.20e, .ote:

Scirpus globulosus Retz. Obs. 6: 19. 1791.

NETHERLANDS NEW GUINEA: Bernhard Camp, Idenburg R., Brass
14087, April 1939, alt. 50 m., on thick beds of floating grass (Leersia) in a
lagoon, erect in large clumps about 1.2 m. high

PUA: Western Division: Lake Daviumbu, Middle Fly R., Brass

7604, August 1936, occasional on floating islands of swamps and lagoons.

New for Papua. Previously known from Netherlands New Guinea
(Ohwi, l.c.), North-East New Guinea, New Ireland, Micronesia, Philip-
pine Islands, Malaya and India.

Fimbristylis insignis Thw. Enum. Pl. Zeyl. 349. 1864.

PAPUA: Western Division: Mai Kussa R., W. MacGregor in 1890
(MEL).

New for New Guinea. Originally described from Ceylon and since re-
ported from China, Borneo and Queensland. MacGregor’s specimen was
written up by F. Mueller simply as “Fimbristylis.”

Fimbristylis complanata (Retz.) Link, Hort. Berol. 1: 292. 1827;
Valck. Suring. Nova Guin. Bot. 8: 703. 1912; Palla in Rechinger,
Denkschr. Math.-Naturw. Kais. Akad. Wiss. Wien 89: 498, 1913.

Scirpus complanatus Retz. Obs. 5: 14. 1789.

216 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXXV

Fimbristylis autumnalis (L.) R. & S. var. complanata (Retz.) Kikenth. Bot.
Jahrb. 59: 50. 1924
PAPUA: Central Division: Port Moresby, C. T. White 4, July-
August 1918.

New for Papua, but known from most of the warmer parts of the Old
World. It is probably this species which was recorded as Fimbristylis
autumnalis (L.) R. & S. by Warburg, Engl. Bot. Jahrb. 18: 186. 1893
and by K. Schumann and Lauterbach, Fl. Deutsch. Schutzgeb. Stidsee
196. 1901.

Fimbristylis microcarya F. Muell. Fragm. Phyt. Austr. 1: 200. 1859.
Fimbristylis autumnalis (L.) R. & S. var. microcarya (F. Muell.) Kikenth.
Bot. Jahrb. 69: 258. 1938.
PAPUA: Central Division: Port Moresby, Carr 11847, April 1935,
open savannah land by the sea (CANB).

This collection was cited by Kukenthal, l.c., as F. autumnalis (L.) R.
& S. var. microcarya (F. Muell.) Kukenth., comb. nov. It is the only
specimen I have seen from outside Australia, though according to C. B.
Clarke in Hook. f. Fl. Brit. Ind. 6: 646. 1893 (where it is treated as a
variety of F. complanata |Retz.| Link) the species extends to India. My
reasons for regarding F. microcarya, F. complanata, and F. autumnalis
as distinct species are given in Proc. Roy. Soc. Queensl. 48: 93. 1937.

Fimbristylis salbundia Kunth, Enum, 2: 230. 1837.

PAPUA: Central Division: Urunu, Vanapa Valley, Brass 4805, July-
August 1933, alt. 1900 m., common species in small swamps on grass country.

New for New Guinea; known previously from Assam and Burma. C. B.
Clarke in Hook. f. Fl. Brit. Ind. 6: 647. 1893 and ex Domin, Biblioth.
Bot. 85: 463. 1915, recorded it from Australia. Domin quoted a col-
lection from the Victoria R., F. Mueller, but the specimens belong to F.
trachycarya F. Muell., a species discussed by me in Proc. Roy. Soc.
Queensl. 48: 92. 1937. Brass’s specimen agrees with pieces of Wallich
3526 ex herb. Berlin and ex herb. Kew, cited in the original description
and by C. B. Clarke, l.c.

Fimbristylis miliacea (L.) Vahl, Enum. 2: 287. 1806, quoad basonym.

Scirpus miliaceus L. Syst. Veg. 10: 868. 1759.

Isolepis miliacea (L.) Presl. Rel. Haenk. 1: 188. 1830.

Trichelostylis miliacea (L.) Nees in Wight, ae 1834, quoad basonym.

Triha miliacea (L.) O. Ktze. Rev. Gen. Pl. 2: 7

Scirpus bengalensis Pers. Syn. 1: 68. 1805.

Fimbristylis ? bengalensis (Pers.) R. & S. Syst. 2: 94. =

Scirpus quinquangularis Vahl, Enum, 2: 279. 1806; syn.

oe pic a (Vahl) Nees in Wight, ‘Cone: 104. 1834;
syn. n

1954] BLAKE, CYPERACEAE COLLECTED IN NEW GUINEA “i?

Fimbristylis quinquangularis (Vahl) Kunth, Enum. 2: 229. 1837; syn. nov.
Triha quinquangularis (Vahl) O. Ktze. Rev. Gen. Pl. 2: 752. 1891; syn. nov.
Scirpus pentagonus Roxb. Fl. Ind. 1: 229. 1820; ed. Carey 1: 218 (1831);
syn. nov.

Isolepis ? pentagona R. & S. Syst. Mant. 2: 69. 1824; syn. nov.

Fimbristylis boeckeleri Steud. Syn. Cyp. 113. 1855; syn. nov.

PAPUA: Central Div Baroka, Brass 3733, April 1933, alt. 30

m., common on damp pny vn plant pale green.

A widely spread species not previously recorded for New Guinea. The
application of the name is explained under the following species.

Fimbristylis littoralis Gaud. in Freyc. Voy. Bot. 413. 1826.
Scirpus tetragonus Poir. Encycl. 6: 767. 1804, nec ae Poir. Encycl.
98. 1817, nec Fimbristylis hihi R. Br.
Scirpus tetragonus Roxb. Fl. Ind. 1: 232. 1820; ed. ae i 228. 1832.
Isolepis ? tetragona R. & S. Syst. Mant. 2: a
Fimbristylis tetragona A. Dietr. Sp. Pl. 2: 152. 1833, non R. Br. 1810.
Fimbristylis flaccidula Zoll. Syst. Verz. Ind. Archip. 2: 61. 1854
Fimbristylis flaccida Steud. Syn. Cyp. 113
Fimbristylis quadrangularis A. Dietr. ex Steud. Sk Cyp. 114. 1855.
NETHERLANDS NEW GUINEA: Bernhard Camp, Idenburg R., Brass
13780, April 1939, frequent on logs floating in lagoons and backwaters at 50 m.
alt.

PAPUA: Western Division: Palmer R., 1 mi. above junction with
Black R., Brass 6946, June 1936, abundant on silt-covered gravel banks; Strick-
land R., W. Bauerlen 22, July 1885 (Bri, MEL); Strickland R., W. Bauerlen
529, Nov. 1885, on river banks (MEL); Gaima, Lower Fly R. (east bank),
Brass 8310, Nov. 1936, common on open sandy foreshores: Daru Island, Brass
6042, March 1934, very common on swampy savannahs. Central Divi-
sion: Thu, Vaitata R., Brass 1016, Feb. 1926, in the sago swamps; Laloki
ty eee i White 173, July ~August 1918,

This is the widely spread plant commonly called Fimbristylis miliacec
(L.) Vahl; Brass 6946, 8310 had been so determined by Uittien, and the
species has been recorded for New Guinea under this name by F. Muell.
Pap. Pl. 2: 35. 1886; K. Schum. Notizbl. Bot. Gart. Mus. Berlin 2: 98.
1898, in Schum. & Lauterb. Fl. Deutsch. Schutzgeb. Siidsee 197. 1901:
Valck. Suring. Nova Guin. Bot. 8: 703. 1912; and Kiikenth. Bot. Jahrb.
59: 50. 1924; also-F. miliacea forma tenerrima Valck. Suring., ].c., Kii-
kenth., l.c.

C. B. Clarke, Jour. Linn. Soc. Lond. 30: 312. 1894, stated that the type
of Scirpus miliaceus L. is a plant of Fimbristylis quinquangularis (Vahl)
Kunth. Through the kindness of Dr. E. D. Merrill and Dr. L. M. Perry,
I have seen a small photograph of the Linnaean type, but unfortunately
the photograph is not sufficiently sharp for critical comparison with
specimens. Through the courtesy of the Director of the Royal Botanic
Gardens, Kew, Mr. E. Nelmes kindly compared some Australian speci-
mens with the Linnaean type. Mr. Nelmes reported as follows:

218 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

I have examined the “type” specimen of Scirpus miliaceus in the
Linnaean herbarium and agree with C. B. Clarke that it represents the
species generally known as Fimbristylis quinquangularis (Vahl) Kunth.
It is a good match of Mr. S. T. Blake’s specimen, no. 11319.

Linnaeus has written “‘miliaceus” on the sheet, and I think the speci-
men may be accepted as his type. There is, however, another Linnaean
specimen, placed next after the type, which has some bearing on this
matter, because it is the species which has always been known as
Scirpus miliaceus L., agreeing well with Mr. Blake’s nos. 7818, 8195, and
8784. C. B. Clarke . . . . does not mention this second sheet, probably
because it bears no inscription by Linneaus. .. .

The combination Fimbristylis miliacea based on Linnaeus’ name must
be restricted to the species represented by Linnaeus’ type, i.e., to the plant
commonly called F. quinquangularis; no other course can be taken that
would be in accord with the International Rules of Nomenclature. C. B.
Clarke’s suggestion, l.c., that Linnaeus would surely have esteemed the
two as one species, does not warrant the deliberate continuance of the
misapplication of the name. N. L. Burman, FI. Ind. 22, ¢. 9, fig. 2. 1768,
seems to have been the first to misapply Linnaeus’ name, and since then
the misapplication has been scarcely questioned.

Several names have been applied to one or other of the two species
commonly known as F. miliacea and F. quinquangularis. Of these only
Scirpus miliaceus L., Scirpus bengalensis Pers., Scirpus quinquangularis
Vahl, Scirpus pentagonus Roxb., Fimbristylis boeckeleri Steud., Fimbri-
stylis flaccida Steud., and Fimbristylis flaccidula Zoll. were based on
specimens, and they are the basonyms of the remaining names. I have
seen types of a portion thereof of S. bengalensis (L), S. quinquangularis
(c), F. littoralis (p) and F. flaccida (v). Fimbristylis flaccida and F.
flaccidula were probably based on the same collection, but I have not
been able to verify this. Scirpus tetragonus Poir. and S. tetragonus Roxb.
were described quite independently of each other; from the descriptions
and specimens from the type-localities there seems no doubt as to the in-
terpretations of these names or of F. boeckeleri. 1 am not so sure of S.
pentagonus, though the description applies well enough to the leafless
states of the species commonly called Fimbristylis quinquangularis. Blake
11319, matched with the type of S. miliaceus, also matches the type of
S. quinquangularis.

Another name must be found for the species commonly called Fimbri-
stylis miliacea. The earliest epithet in the required position is in the
combination Scirpus tetragonus Poir. Encycl. 6: 767. 1804, but this is not
available under Fimbristylis because of F. tetragona R. Br. and upon
which Poiret’s later homonym was based in 1817. The next is in Fimbri-
stylis littoralis Gaud., and this appears to be the correct name for the
species. Scirpus bengalensis Pers. has been generally referred to this
species, but Persoon’s description (“‘involucro tetraphyllo spiculis. .. .
Ovatis (minutis): squamis concavo-carinatis mucronatis”’) agrees better

1954] BLAKE, CYPERACEAE COLLECTED IN NEW GUINEA 219

with true Fimbristylis miliacea; a recent examination of the type (L) con-
firms this.

Two other names require mention. ‘“Fimbristylis angularis Steud.”
Syn. Cyp. 116. 1855 has been referred to F. miliacea (F. quinquangularts )
by some. Steudel’s combination was based on /solepis angularis Schrad.
ex R. & S. Syst. Mant. 2: 69. 1824, which appears to have been based on
Fimbristylis angularis Link, Enum. Hort. Berol. 1: 289. 1821, so that
Steudel’s transfer was quite unnecessary. Boeckeler’s description (Lin-
naea 37: 30-31. 1871) of what appears to be Link’s type refers to some
other species, particularly as to the long bracts, relatively large spikelets
and sometimes bifid styles. Fimbristylis trachycarya F. Muell. Fragm.
1: 199, 1859, sometimes referred to F. littoralis, belongs to an endemic
Australian species which has been discussed elsewhere (S. T. Blake, Proc.
Roy. Soc. Queens]. 48: 92. 1937).

Fimbristylis littoralis and F. miliacea do not differ greatly in spikelet
structure, though the more or less globular, very obtuse, scarcely at all
angular spikelets of the former with their muticous and more or less con-
cave glumes are, with a little experience, readily enough distinguished
from the ovoid, less obtuse, more distinctly angular spikelets of the latter
with their rather prominently keeled, more or less distinctly apiculate
glumes. The foliage of the two species is markedly dissimilar, but is some-
times poorly developed. The leaves of F. littoralis are vertically flattened,
finely striate without prominent veins, with rather thin edges, and are
usually borne as distichous, more or less flabellate tufts between the culms.
The leaves of F. miliacea are of the usual Fimbristylis type, dorso-ventrally
flattened with a prominent midrib and rib-like margins. The revised
synonymy of the two species is given above.

Fimbristylis furva R. Br. Prodr. 228. 1810.

PAPUA: Western Division: Mai Kussa R., W. MacGregor in 1890
(MEL).

Previously known only from Queensland, including islands in Torres
Strait

Fimbristylis pycnocephala Hillebr. Fl. Haw. Isl. 473. 1888.
Fimbristylis cymosa R. Br. var. capitato-umbellata Hillebr. Fl. Haw. Isl.
473. 1888; syn. nov.
Fimbristylis cymosa R. Br. var. subcapitata C. B. Clarke ex Hemsl. Jour.
Linn. Soc. Lond. 30: 197. 1894.
Fimbristylis cymosa R. Br. var. aN a (Hillebr.) Kikenth. in Cristo-
phersen, Bull. Bishop Mus. 128: 20.
SOLOMON ISLANDS: Ulawa, Brass 2990, October 1932, ocean foreshore,
common

New for the Solomon Islands; previously known from the Hawaiian
Islands (!), Tonga (!), New Caledonia and Samoa.

220 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXXV

In this species, the spikelets may all be aggregated in a single dense
head or clustered on the rays of an umbel-like inflorescence; all inter-
mediate states occur. Hawaiian plants with the looser inflorescences were
referred by Hillebrand to F. cymosa R. Br. and F. cymosa var. capitato-
umbellata, but all states of F. pycnocephala may be distinguished from the
Australian F. cymosa as follows:

Spikelets + oblong; broad, hyaline margins of glumes loosely reticulate; nut
somewhat shining, smooth or minutely striate only, the epidermal cells minute,
square or nearly so, arranged in fairly regular vertical series but the transverse
arrangement + irregular. .......... F. pycnocephala.

Spikelets ovoid; broad, hyaline margins of glumes not pinenste reticulate,
the cells composing them very small with inconspicuous margins; nut some-
what roughened, rather indistinctly striate vertically and haaraliber the
minute epidermal cells transversely paid sible, a regularly in
both vertical and horizontal series ._.. F. cymosa.

Sect. Abildgaardia (Vahl) Benth.

Fimbristylis cinnamometorum (Vahl) Kunth, Enum. 2: 229. 1837:
Kikenth. Mitteil. Thiiring. Bot. Ver. N. F. 50:

Scirpus cinnamometorum Vahl, Enum. 2: 278. 1806.

Fimbristylis cyperoides R. Br. Prodr. 228. 1810.

PAPUA: Western Division: Tarara, Wassi Kussa R., Brass 8408,
Dec. 1936, savannah-forest, abundant on wet flats; Lake Daviumbu, Middle
Fly R., Brass 7876, Sept. 1936, occasional in small erect tufts on wet grass
plains; Gaima, Lower Fly R. (east bank), Brass 8262, Nov. 1936, plentiful on
hard soil in open savannah-forests.

These are the only collections known from New Guinea, though the
species is known to range from NE. Australia to Ceylon and SE. Asia.

Fimbristylis cinnamometorum and F. cyperoides have sometimes been
treated as distinct species on the grounds that the former (from Ceylon,
Pegu and China) is an annual, while the latter (from Australia to the
Philippine and Caroline Islands) has a prominent rhizome. But the type
of Scirpus cinnamometorum (c) is from a perennial plant and is well
matched by many Australian specimens; the degree of development of a
rhizome depends on the age of the plant and varies greatly in herbarium
material.

Fimbristylis stenochlaena Kiikenth. Mitteil. Thiiring. Bot. Ver. N. F.
50: 11. 1943.

PAPUA: Western Division: Lake Daviumbu, Middle Fly R., Brass
7840, Sept. 1936, very abundant on wet grass plains, leaves of young plants
arranged spirally in a flat rosette (TYPE collection).

The plants seen by me appear to be somewhat larger than those seen by
Kukenthal, 30-50 cm. high, with asperulous many-ribbed rather than
smooth obsoletely pentagonal sulcate culms, with 9 or 10 (not 4-6)

1954] BLAKE, CYPERACEAE COLLECTED IN NEW GUINEA 221

-rayed inflorescences with rather longer and broader mature spikelets
9-11 mm. long and 2—3 mm. wide (described by Kukenthal as 8-9 mm.
long and 1.5-2 mm. wide). I find the glumes acute and more or less
cuspidulate rather than long acuminate, and the nuts (five examined)
obovoid, not at all pyriform, 0.8 mm. long, 0.55 mm. wide.

Fimbristylis fusca (Nees) C. B. Clarke in Hook. f. Fl. Brit. Ind. 6:
649. 1893.

Abildgaardia fusca Nees in Wight, Contrib. Bot. Ind. 95. 1834.

NETHERLANDS NEW GUINEA: Balim R., Brass 11744, Dec. 1938, alt.
1600 m., common among the grass on long-deforested slopes.

New for New Guinea; previously known from India and the Philippine
Islands to Java and the Moluccas. The plants are glabrous.

Clarke, l.c., cites “Benth. in Gen. PI. iii. 1048” as the author of the
combination, but Bentham here merely reduces the genus Abildgaardia
to a section of Fimbristylis. In Index Kewensis, Suppl. 4: 92. 1913 the
author of the combination is given as “Benth. ex C. B. Clarke in Hook. f.
Fl. Brit. Ind. vi. 649 (1893).”

Fimbristylis fimbristyloides (F.,Muell.) Druce, Rep. Bot. Exch. Club
Brit. Isles 1916: 623. 1917
Abildgaardia fimbristyloides F. Muell. Fragm. Phyt. Austr. 8: 273. 1874.

PAPUA: Central Division: Rona, Laloki R., Brass 3576, April 1933,
alt. 450 m., common, wet places on open grassy hillsides.

New for New Guinea; previously known only from northern Queens-
land.

Fimbristylis intonsa sp. nova (Sect. Abildgaardia). Fic. 2.

erba annua, usque ad 16 cm. alta. Culmi fasciculati, setacei, stricti,
erecti, quinquangulares angulis angustis elevatis sursum scabris. Folia
pauca, usque ad 5 cm. longa, , cu Imi te rtiam partem raro adaequantia, 1—2

orificem pubescens, ceterae glabrae; laminae lineares, sursum admodum
angustatae, apice obtusae vel acutae, falcatae vel tortuosae, planae vel
incurvae, coriaceae, haud carinatae, paucinerves, marginibus leviter in-
crassatae sursum scabrae, supra prominule subtus indistincte celluloso-
reticulatae, eae foliorum caulinorum saepe multo breviores angustioresque.
Anthela simplex vel composita, 2—7-radiata, laxa. Bracteae 1—2 infimae
subfoliaceae usque ad 12 mm. longae, vel omnes setaceae, saepe minimae;
bracteolae minimae. Radii setacei, compressi, scabri, usque ad 2.5 cm.
longi; radioli suberecti, usque ad 7 mm. longi. Spiculae oblongae, acutae,
compressae, saepe tortae, 5-8 mm. longae, 1.5—1.8 mm. latae, multi- et
densiflorae; rhachilla excavata alata. Glumae distichae vel sursum specie
laxe spiraliter dispositae, brunneae vel ferrugineae, latissime ovatae,

222 JOURNAL OF THE ARNOLD ARBORETUM [VOL, XXXV

obtusae, mucronatae, l-nerves, carinatae carina curva sursum valida
excurrente, fere omnino albo- -pubescentes, marginibus vix hyalinis ciliatae,
1.7-1.8 mm. longae, cellulis minutissimis; 1—2 infimae vacuae minores.
Stamina 3; antherae lineares, apiculatae, 0.5 mm. longae; filamenta lata.
Stylus pro ratione robustus, triquetrus, basi latiuscule pyramidatus, omnino
glaber, 1 mm. longus; stigmata 3, stylo breviora. Nux albida, obovoidea
fere ellipsoidea, utrinque rotundata apice leviter umbonulata, minime
stipitata, obtuse trigona angulis haud prominulis, lateribus convexulis,
verrucosa, 0.7 mm, longa, 0.5 mm. lata, cellulis extimis hexagonis parvis
indistinctis.

PAPUA: Western Division: Lake Daviumbu, Middle Fly R., Brass
7841, Sept. 1936, very common on wet grassy plains (TYPE).

This collection was labelled ‘“‘Fimbristylis disticha Boeck. (det. Kiiken-
thal).” It is certainly allied to F. disticha Boeck., but to judge from the
piece of the type of this species in herb. Brisbane ex herb. Berlin, it differs
in the broader, more coarsely keeled, prominently mucronate and densely
pubescent glumes and the broader, more ellipsoid nut not cuneate but
rounded at the base, less distinctly ribbed at the angles and with hexagonal
rather than oblong external cells. According to C. B, Clarke in Hook. f.
Fl. Brit. Ind. 6: 651. 1893, the glumes of F. disticha are minutely ciliate
at the margins, but I find them glabrous. Kiikenthal, Bot. Jahrb. 59: 50.
1924, and 69: 258. 1938, has recorded F. disticha from North-East New
Guinea, but I have not seen these specimens.

The species of sect. Abildgaardia discussed may be distinguished as
follows:

Leaves setaceous, at least half as long as culm, more or less distichous; basal
sheaths + horny; glumes scabrous with reddish glands; nut (0.8-1 mm.
long) with the external cells transversely oblong. .... F. cinnamometorum.

Leaves flat, at least 1 mm. wide, less than half and sometimes scarcely one
quarter as long as culm, spirally arranged, often falcate; basal sheaths not
horny, prominently nerved; glumes not (or in F. fusca very rarely) glandu-
lar, usually pubescent (except in F, disticha); nut (0.6-1 mm. long) with
the external cells hexagonal, not or but slightly elongated transversely.

Glumes 3—5 mm. long, the upper ones somewhat spiral; style 3-4 mm. long,

3-6 times as long as the obovoid nut; perennial (? always).

al &

Fic. 2. eee’ — S. T. Blake: a. inflorescence, & 1; b., c. spike-
lets, X 3; d. e, 5; e. flower, X 15; f. nut, X 15; g. transverse section
of nut; h. aes of nut, X 40. Figures from type.

1954] BLAKE, CYPERACEAE COLLECTED IN NEW GUINEA 223

Glumes about half as wide as long, 4-5 mm. long; style 4 mm. long;
culms many-ribbed; leaves 1.5-4 mm. wide; spikelets 2-3 mm. wide.
slate nae Gao eee aaa eds 4s VO eee eee F. stenochlaena.

Glumes more than half as wide as long, 3-4 mm. long; style about 3 mm.
long; culms 5-ribbed; leaves 2 mm. wide; spikelets 1.5-2 mm. wide.
Be ee eran tle Gites ee en id Ss aS Se ere oo F, fusca.

Glumes up to 2.5 mm. long, regularly distichous, but the whole spikelet some-
times twisted; style 1-1.3 mm. long, less than twice as long as nut; an-
nual plants.

Glumes glabrous, muticous or nearly so, acute, the keel slightly curved,
1.6-1.7 mm. long; nut narrowly obovate, cuneate to a very narrow
base, audit 0.6% 0.05 WM, occu dss se dowee rig eee ies F. disticha.

Glumes puberulous or rarely glabrous, acute, mucronulate, the keel straight
or nearly so, 2-2.5 mm. long; nut broadly obovate, somewhat attenu-
ate above the broad truncate base, about 0.75 X 0.55 mm...........
it. 5. p52 4.454 bee k oe pS F. fimbristyloides.

Glumes densely pubescent with ciliolate scarcely hyaline margins, very
road and very obtuse, rather coarsely mucronulate, the keel curved,
1.7-1.8 mm. long; nut obovate, rounded to the base, 0.7 % 0.5 mm.
ee eee aera a eA ek ea ee asad Se eae F. intonsa.

Scleria Bergius

There is no recent general account of the species of Scleria in the Malay-
sian-Australian region. A revision of the Australian species was commenced
some years ago and has been intensified and expanded more recently to
include the Malaysian species. The following account of the Brass collec-
tions is really a preliminary revision of all the species known from New
Guinea. It would seem that a large number of names for alleged new
species have been based on far too scanty material, and that very little
attention has been paid to individual variation. Misapplications of names
have been fairly frequent; it would appear that some early botanists did
not see the types of the names proposed by their predecessors (which are
frequently scrappy in any case), and their often faulty identifications were
blindly accepted by later botanists, possibly for want of better evidence.
I have been fortunate in seeing types or isotypes of a goodly proportion of
the names mentioned in this paper; of others I have seen topotypes —
specimens from the type-locality which agree with the original description.
Although an extensive series of specimens from Australia, Malaysia and
elsewhere has now been examined, study of further material, particularly
from India, may demand modifications of some of the conclusions pub-
lished in this paper.

A curious fact is that there appears to be no endemic species in New
Guinea. Of the thirteen accepted species, all but one (S. polycarpa) occur
elsewhere in Malaysia and all but two or three (S. pergracilis, S. motleyi
and, perhaps, S. tessellata) occur in Australia, some of them extending
further eastward.

224 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

Scleria pergracilis (Nees) Kunth, Enum. Pl. 2: 354. 1837.
Hypoporum pergracile Nees, Edinb. New Philipp. Jour. 17: 267. 1834.

NORTH-EAST NEW GUINEA: Kani Hills near Ongoruna (Unguruna) Vil-
lage, Lane-Poole 635, Feb. 1924 (has as strong a lemon-verbena scent as to make
a whole hill- tg perfumed); Partep, Commonwealth Nutritional Survey N.S. P.
69,N.S.P.9

New for New Guinea; previously known from Tropical Africa, India
and Ceylon.

—o lithosperma (L.) Sw. Prodr. Veg. Ind. Occid. 18. 1788; Boeck.
n Engl. Forschungsr. S. M. S. Gazelle 4 (1): 18. 1889; K. Schum. &
twee Nachtr. Fl. Deutsch. Schutzgeb. Siidsee 60. 1905; Valck.
Suring. Nova Guin. Bot. 8: 711. 1902; Kiikenth. Bot Jahrb. 59: 58.
1924; Ohwi, Bot. Mag. Tokyo 56: 212. 1942.

Scirpus lithospermus L. Sp. Pl. ed. 1: 51. 1753.

Schoenus lithospermus (L.) L. Sp. Pl. 2: 65. 1762.

Scleria tenuis Retz. Obs. 4: 13. 1786.

Scleria capillaris R. Br. Prodr. 240. 1810.

Scleria wightiana Steud. Syn. Cyp. 176. 1855.

Hypoporum lithospermum Nees in Mart. Fl. Brasil. 2 (1): 172. 1842.
Hypoporum capillare (R. Br.) Nees, Linnaea 9: 303. 1834

NORTH-EAST NEW GUINEA: New Britain, Parkinson in 1885 (MEL).

PAPUA: Western Division: Lower Fly R., east bank opposite Sturt
Island, Brass 8059, Oct. 1936, rain-forest, tufts 60-80 cm. high, forming a
scant ground cover on dry ridges: Daru Island, Brass 6259, March 1936, com-
mon along edge of rain-forest. Eastern Division: Lower Mori R., Brass
1634, June 1926, on rain-forest floor.

Cosmotropical. Further synonyms are listed by Core, Brittonia 2: 27-8.
1936. Brass 6259, 8059 had been determined by Kiikenthal.

Scleria roxburghii (C. B. Clarke) Domin, Biblioth. Bot. 85: 487, 1915.

Scleria lithosperma (L.) Sw. var. B Thw. Enum. Pl. Zeyl. 354. 1864.

Scleria lithosperma (L.) Sw. var. roxburghii C. B. Clarke in Hook. f. Fl.
Brit. Ind. 6: 686. 1894.

Scleria lithosperma (L.) Sw. var. linearis Benth. Fl. Austral. 7: 430. 1878.

a roxburghiu (C. B. wr Domin var. australiensis Domin, Biblioth.

5: 487. 1915; syn. n

repec neh roxburghi Nees, ex C. B. Clarke in Hook. f. Fl. Brit. Ind. 6:
686. 1894, i

Hypoporum an Nees ex Boeck. Linnaea 38: 452. 1874, in syn.

PAPUA: Western Division: Tarara, Wassi Kussa R.. Brass 8504,
Dec. 1936, rain-forest, dense tufted ground cover 50-60 cm. high, in partial
shade. Central Division: Baroka, Nakeo district, Brass 3774, April 1933,
alt. 30 m., common in brushy rain-forest and on garden clearings; Port Moresby
to Kalo, MacGregor in 1889 (MEL).

1954] BLAKE, CYPERACEAE COLLECTED IN NEW GUINEA 225

New for New Guinea. Known elsewhere from Queensland, Philippine
Islands, India and Ceylon.

Scleria motleyi C. B. Clarke, Philipp. Jour. Sci. 2 (C): 104. 1907;
Ohwi, Bot. Mag. Tokyo 56: 213. 1942

Scleria motleyi C. B. pie var. densi-spicata C. B. Clarke, Philipp. Jour.
Sci. 2 (C): 104. 1907; syn. n

ae eon Ridl. ae Str. Branch Roy. As. Soc. 46: 228. 1906,

. Fl. Malay. Pen. (Monocot.) 3: 110. 1907; non Steud. 1855.

Scleria ae Wee Philipp. Jour. Sci. 8 (C): 363. 1913.

Scleria gonocarpa Ridl. Fl. Malay Pen. 5: 177. 1925; syn. nov.

Scleria ee Elmer ex Merr. Enum. Philipp. Fl. Pl. 1: 134. 1922, in
obs. .; Elmer, Leafl. Philipp. Bot. 10: 3541. 1938, descr. anglice.

Ses paved Elmer, Leafl. Philipp. Bot. 10: 3542. 1938, in obs., pro

NETHERLANDS NEW GUINEA: Hollandia and vicinity, Brass 8870,
June-July 1938, alt. 100 m., rain-forest, tufted floor plant on sharp ridge
crests.

Previously recorded by Ohwi, l.c., for Netherlands New Guinea; else-
where known from Malaya, Borneo and Philippine Islands.

Scleria tessellata Willd. Sp. Pl. 4: 315. 1805; K. Schum. in Warb. Bot.
Jahrb. 13: 267. 1891; Valck. abe Nova Guin. Bot. 8: 712. 1912.

Scleria parvula Steud. Syn. Cyp. 174. 1855.

Scleria uliginosa Hochst. ex Boeck. de 38: 471. 1874.

PAPUA: Western Division: Lake Daviumbu, Middle Fly R., Brass
7527, August 1936, sporadic on savannahs; Lake Dav viumbu, Middle Fly R.,
Brass 7875, Sept. 1936, sporadic on wet grass plains, erect in small tufts; Lake
Daviumbu, Middle Fly R., Brass 7963, Sept. 1936, rain-forest, common in
semi-shade on shores of lake.

New for Papua; known to extend in an apparently sporadic manner
north and west to India and Ceylon. Some of the records of this species
from Mayalsia belong to the closely allied S. biflora Roxb. The Australian
plants referred to it by Bentham, Fl. Austral. 7: 430. 1878 belong to
other species.

Scleria novae-hollandiae Boeck. Flora 58: 120. 1875.
Scleria merrillii Palla, Allgem. Bot. Zeitschr. 17: Beil. 8. 1911; syn. nov.
PAPUA: Western Division: Wuroi, Oriomo R., Brass 6012, Feb—-

March 1934, alt. 50 m., on a savannah ridge; Daru Island, Brass 6427, April 1936,
gregarious on wet soil in savannah forest.

New for New Guinea; previously known from northern and north-
eastern Australia and the Philippine Islands.

This species was referred by Bentham, FI. Austral. 7: 428. 1878 to
S. laxa R. Br., and in this he was followed by later authors. I have dis-
cussed these species as to their occurrence in Australia in Proc. Roy. Soc.

226 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

Queensl. 58: 48-9. 1947 and 60: 52-3. 1949. I have now seen the type
and an isotype of S. novae-hollandiae, a photograph and a fragment of the
type of S. axa and isotypes of S. merrillii. The species is well distinguished
from other small annual species by the dull white, nearly smooth surface
of the more or less oblong nut and the small thin tightly appressed disc
-with broadly rounded lobes and shallow sinuses. The chalky appearance
of most nuts is very distinctive. Kikenthal determined Brass 6427 as
S. tessellata, which has an ellipsoid, shining, prominently tessellated nut
and a larger, thicker, more deeply divided disc with rather acute lobes and
sinuses. To judge from his remarks on the disc, the specimen upon which
Kiikenthal based his record for New Guinea of S. browni Kunth, Bot.
Jahrb. 70: 464. 1940 may also belong here.

Scleria rugosa R. Br. Prodr. 240. 1810.

Scleria lateriflora Boeck. Linnaea 38: 455. 1874; syn. nov.

Scleria pubigera Makino, Bot. Mag. Tokyo 27: 55. 1913; syn. nov.

PAPUA: Western Division: Lake Daviumbu, Middle Fly R., Brass
7532 A, August 1936, savannahs, gregarious in flat tufts on hard-pans and
swampy margins; Wuroi, Oriomo R., Brass 5868, Jan—March 1934, alt. 30 m.,
rare, gray soil on low savannah ridge; Wuroi, Oriomo R., Brass 6013, Feb.
March 1934, alt. 30 m., damp slopes of a savannah ridge, uncommon.

New for New Guinea; previously known from northern and_north-
eastern Australia and New Caledonia northwards and westwards to Ceylon
and Japan. Specimens from Ceylon (S. lateriflora Boeck.) were referred
to S. zeylanica Poir. by Thwaites and others, but Poiret’s original descrip-
tion refers to a large plant which, from the examination of an excellent
piece of the type loaned from herb. Paris, proves to belong to the species
previously described by Retz as Scleria levis and later described by Nees
as Scleria hebecarpa. Apparently little attention has been paid to the
extensive description in French following the brief Latin diagnosis. The
references to S. zeylanica in Malaysian and New Caledonian literature
really belong to S. rugosa. I have seen excellent isotypes of S. rugosa and
S. lateriflora, but the interpretation of S. pubigera Makino is based chiefly
on specimens so labelled from Taiwan in herb. Melbourne. Brass 7532 A
was determined by Kikenthal as Scleria tessellata Willd. var. debilis
Benth.; the latter is a common Queensland plant differing from S. rugosa
in habit and the shape of the nut and disc.

Scleria levis Retz. Obs. 4: 13. 1786.

Scleria zeylanica Poir. Encycl. Meth. 7: 3. 1806; syn.

Scleria hebecarpa Nees in Wight, Contrib. Bot. Ind. "MT. 1834; Valck. Suring.
Nova Guin. Bot. 8: 712. 1912; Palla in Rechinger, Denkschr. Math.-Naturw.
Kais. Akad. Wiss. Wien 89: 500. 1913; saosingas Bot. Jahrb. 59: 58. 1924;
Ohwi, Bot. Mag. Tokyo 56: 212. 1942; syn.

Ge hebecarpa Nees var. ee (Steud) C. B. Clarke in Hook f. FI.
Brit. Ind. 6: 689. 1894; syn. n

1954] BLAKE, CYPERACEAE COLLECTED IN NEW GUINEA 227
Scleria hebecarpa Nees forma pilosa Valck. Suring., Nova Guin. Bot. 8: 712.
2: nov.

Scleria neesiana Hook. & Arn. Bot. Beechey Voy. 229. 1841; syn. nov.

Scleria pubescens Steud. Syn. Cyp. 168. 1855; syn. nov.

Scleria japonica Steud. Syn. Cyp. 169. 1855; syn. nov.

Scleria vestita Boeck. Linnaea 38: 482. 1874; syn. nov.

Scleria dietrichiae Boeck. Flora 58: 121. 1875; syn. nov

Scleria wichurai Boeck. Bot. Jahrb. 5: 510. 1884; syn. nov.

PAPUA: Western Division: Lake Daviumbu, Middle Fly R., Brass
7877, Sept. 1936, occasional on wet grass plains (clumps 80 cm. high Lake
Daviumbu, Middle Fly R., Brass 7691, Sept. 1936, savannahs, common on
swamp margins; Gaima, Lower Fly R. (east bank), Brass 8264, Nov. 1936,
open savannah-forest, plentiful on hard soils; Tarara, Wassi Kussa River,
Brass 8715, Jan. 1937, common grass associate, ’savannah- forest ridges; Dagwa,
Oriomo R.., a 5996, Feb.—March 1934, alt. 40 m., amongst grasses on a
savannah ri

This ee has not previously been recorded for Papua, though it is
widely spread from Queensland and New Caledonia to India and Japan.
It is the species commonly known as Scleria hebecarpa Nees, but examina-
tion of the type of Scleria levis Retz. (Lp) and of a piece of the type of
Scleria zeylanica Poir. (P) has shown that these three names are synony-
mous. For some inexplicable reason, Poiret’s name has been almost in-
variably associated with S. lateriflora Boeck. (which is S. rugosa R. Br.)
in spite of the fact that Poiret’s description refers to a much larger and
otherwise different plant (see also under S. rugosa). The name Scleria levis
(the epithet of which is often spelled /aevis) has been commonly applied to
specimens of S. terrestris (L.) Fassett on which the fruits are somewhat
depressed due to imperfect development.

Brass 8264 had been determined as Scleria hebecarpa Nees by Kiikenthal.
Brass 7691, 8715 represent the more or less hairy form described as S.
hebecarpa Nees var. pubescens (Steud.) C. B. Clarke (and were determined
as such by Kiikenthal), S. pubescens Steud. and S. vestita Boeck.

Scleria ciliaris Nees in Wight, Contrib. Bot. Ind. 117. 1834.

Scleria chinensis Kunth, Enum. Pl. 2: 357. 1837

Scleria bancana Miq. FI. Ind. Bat. Suppl. 602. 1860; Valck. Suring. Nova
Guin. Bot. 8: 712. 1912; Kanehira, Jour. Dept. Agr. Kyushu Univ. 4: 282.
1935; Kiikenth. Bot. Jahrb. 69: 261. 1938; Ohwi, Bot. Mag. Tokyo 56:
212. 1942; syn. nov.

Scleria malaccensis see se 38: 507. 1874; K. Schum. in Warb. Bot.
Jahrb. 13: 266. 1891:

PAPUA: Western Ree ion: Lake Daviumbu, Middle Fly R., Brass
7670, Sept. 1936, rain-forest, occasional clumps 1.2-1.5 m. high, along matein
of lake (det. Uittien as S. hebecarpa Nees); Gaima, Lower Fly R. (east bank),
Brass 8252, Nov. 1936, common grass associate in savannah forests (det.
Uittien as S. hebecarpa Nees); Wuroi, Oriomo R., Brass 5808, Jan.—March
1934, alt. 10 m., amongst tall grass on edge of rain-forest. Gulf Division:
Kerema, Brass 1207, March 1926, open grassland near coast, in clumps | m.

228 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXXV

high. Central Division: On range between Sogere and Javararie, White
317, July-Aug. 1918; Astrolabe Range, White 218, July-August 1918, in grass-
land; Astrolabe Range, Armit (MEL). Eastern Division: Cloudy Mts.,
Chalmers (MEL).

SOLOMON ISLANDS: Mariga Island: N’Gela, Brass 3486, Jan. 1933,
small sedge on grasslands.

New for Papua and the Solomon Islands; extends from northern Queens-
land northwards and westwards to Ceylon, India and China.

One state of this species has been generally called Scleria chinensis since
the time of Kunth. Kunth proposed this name to replace Scleria ciliaris
Nees “‘because of the earlier name of Michaux.” But the name for the
American species proposed by Michaux is Scleria ciliata. The suffixes -tus
(-ta, -tum = provided with) and -ris (-re = of or belonging to) have
quite distinct meanings, so that the two epithets ciliata and ciliaris (and
consequently the names of which they form part) must be treated as
distinct (International Code of Botanical Nomenclature, Art. 82, and
Rec. 821); they are not orthographic variants. Kunth’s action was there-
fore unjustified and merely created a superfluous name. The name pro-
posed by Nees is the correct one.

Scleria malaccensis is the same form as S. bancana and has usually
been distinguished on the grounds that the leaves of S. ciliaris (S. chinen-
sis) are scattered and have more or less distinctly winged sheaths while
S. bancana has the middle leaves approximated in pairs with usually un-
winged sheaths, even though S. bancana was originally described as having
winged sheaths, The specimens cited above show a continuous series
between the two extremes.

I have not been able to identify Scleria chinensis Kunth var. biauriculata
C. B. Clarke in Hook. f. Fl. Brit. Ind. 6: 690. 1894. The description could
refer to members of the series just mentioned; the synonym cited, S.
exaltata Boeck., belongs to S. terrestris; one of the two collections cited
is also cited as belonging to the typical form of S. chinensis and the other
may or may not be Boeckeler’s type.

Scleria macrophylla Presl has been referred to S. chinensis by some
authors, but this name refers to an American species (Core, Brittonia 2:
37-8. 1936).

Scleria terrestris (L). Fassett, Rhodora 26: 159. 1924.

Zizania terrestris L. Sp. Pl. ed. 1: 991. 175

Diaphora cochinchinensis Lour. FI. Cocltiah. 578. 1790.

Olyra orientalis Lour. Fl. Cochinch. 552. 1790.

Scleria elata Thw. Enum. Pl. Zeyl. 353. 1864; K. Schum. Notizbl. Bot. Gart.
Mus. Berlin 2: 98. 1898; Lam, Nat. Tijds. Nederl. 88: 194, 203, 207. 1928.

Scleria melanostoma Nees ex Boeck. Linnaea 38: 514. 1874

Scleria exaltata Boeck. Bot. Jahrb. 5: 511. 1884; syn. nov

Scleria hasskarliana Boeck. Bot. Jahrb. 5: 511. 1884; syn. nov.

Scleria haematostachys Boeck. Bot. Jahrb. 5: 512. 1884; syn. nov.

Scleria doederleiniana Boeck. Bot. Jahrb. 5: 512. 1884; syn. nov.

1954] BLAKE, CYPERACEAE COLLECTED IN NEW GUINEA 229

Scleria ploemii Boeck. Bot. Jahrb. 5: 513. 1884; syn. nov.
Scleria luzonensis Palla, Allg. Bot. Zeitschr. 13: 49. 1907; syn. n
Scleria cochinchinensis (Lour.) Druce, Rept. Bot. Exch. Club Brit. Isles 4:
646. 1917
Sclerta chinensis Kunth var. luzonensis (Palla) Uitt. Rec. Trav. Bot. Neéer-
land. re 1935 et Meded. Bot. Mus. Herb. Rijksuniv. Utrecht 17: 201.
1935; sy
Scleria pice Kunth var. luzonensis (Palla) Uitt. forma pilosa Uitt. Il.cc.
syn. nov.
NETHERLANDS NEW GUINEA: 4 km. SW. of Bernhard Camp, eames
R., Brass 13480, March 1939, alt. 850 m., one clump on an open landslip
NORTH-EAST NEW GUINEA: Morobe District: Belung R. to Sara-
waket, J. & M.S. Clemens 4875, Jan. 1937.

PAPUA: Central Division: Port Moresby, Lawes in 1884 (MEL);
Sogere, White 308, July-Aug. 1918; Urunu, Vanapa Valley, Brass 4779, July-
Aug. 1933, alt. 1900 m., in clumps amongst tall weed grass on old garden land,
common.

New for Papua; extends from Queensland to India and southern China.

This is the species commonly known as Scleria elata Thw. The name
Scleria terrestris is based on Zizania terrestris L., which is based on Katu-
Tsjolam Rheede, Hort. Mal. 12: 113, ¢. 60. 1703, and this evidently refers
to a Scleria; C. B. Clarke in Hook. f. Fl. Brit. Ind. 6: 685-694. 1894
recorded only S. elata Thw. and S. hebecarpa Nees from the region covered
by Rheede, and Rheede’s figure could only refer to S. elata. Sclerta
cochinchinensis is based on Diaphora cochinchinensis Lour., the type of
which was seen by Merrill (see Trans. Am. Phil. Soc. n.s. 24: 89. 1935).
Olyra orientalis Lour. is referred here on Merrill’s suggestion, l.c., 90. I
have seen syntypes of Scleria elata (MEL), S. hasskarliana (MEL), S.
luzonensis (BRI, NSW, L) and possibly of S. ploemii (L). Thwaites 3031,
cited by Boeckeler as the type of S. exaltata, is represented at Melbourne,
but the specimen differs considerably from Boeckler’s description in being
much smaller and having differently shaped disc-lobes; it belongs to S. levis
Retz. (S. hebecarpa Nees).

Scleria scrobiculata Nees & Meyen in Wight, Contrib. Bot. Ind. 117.

34; K. Schum. & Lauterb. Fl. Deutsch. Schutzgeb. Suidsee 198.

1901 (at most only partly); Valck. Suring. Nova Guin. Bot. 8: 712.

1912 (at most only partly); Kiikenth. Bot. Jahrb, 59: 58. 1924,

partly; ? Kanehira, Jour. Dept. Agr. Kyushu Univ. 4: 282. 1935;
(?) Ohwi, Bot. Mag. Tokyo 56: 212. 1942.

Scleria timorensis Nees, Linnaea 9: 303.

Scleria purpureovaginata Boeck. Bot. Tk 5: 513. 1884; K. Schum. in
Warb. Bot. Jahrb. 13: 266. 1891; Valck. Suring. Nova Guin. Bot. 8: 713.
1912; syn. nov

Scleria keyensis K. Schum. in Warb. Bot. Jahrb. 13: 267. 1891; Valck. Suring.

ova Guin. Bot. 8: 713. 1912.
Scleria suffulta C. B. Clarke, Kew Bull. Add. Ser. 8: 58. 1908; syn. nov.

230 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

PAPUA: Western Division: New Guinea, near Dutch boundary, Mac-
Gregor in 1890 (MEL); Fly R., D’Albertis (MEL); Lake Daviumbu, Middle Fly
R., Brass 7715, Sept. 1936, mixed with grass fringing shore of savannahs, clumps
2-2.5 m. high (det. Kiikenthal); Daru Island, Brass 6244, March 1936, common
with grasses on damp soil of savannah forests (det. Kiikenthal as Scleria chinen-
sis Kunth). Central Division: Port Moresby, Goldie (MEL): towards
Owen soa Range, Goldie in 1878 (MEL); Astrolabe Range, Armit in 1883,
scrubs (MEL)

New for Papua, and perhaps for New Guinea. The species is widely
spread in Malaysia, extending into Queensland and apparently to China.
At least some of the records of this species for New Guinea belong properly
to Scleria polycarpa (see under this species), but I have not seen many
of the specimens on which the records are based.

Scleria polycarpa Boeck. Linnaea 38: 508. 1874.

Scleria deh grea Willd. Sp. Pl. 4: 312. set aes in Gibbs, Phyt. FI.
Arfak Mts. 200. 1917; non Gaertn. 1788;

Scleria graeffeana Boeck. Flora 58: 121. pie K. ae in Warb. Bot. Jahrb.
13: 266. 1891; Valck. Suring. Nova Guin. Bot. 8: 712. 1912; Palla in seb
Denkschr. Math. -Naturw. Kais. Akad. Wien 89: 500. nie cicoey

Scleria ternifolia Domin, Biblioth. Bot. 85: 490. 1915;

NETHERLANDS NEW GUINEA: Bernhard ne Idenburg R., Brass
13937, April 1939, alt. 50 m., abundant in sago and other is swampy
forest of river plain, 2-3 m. high

NORTH-EAST NEW GUINEA: New Ireland: W. coast, Bradtke 125,
May 1917, Om. (Nsw). Duke of York Island, Bradtke 185, May 1917,
secondary bush, grassfields (Nsw). Morobe District: Augustusfluss
(= Sepik R.) Hollrung 815 (mex, 1); Huon Gulf, Lauterbach 1185 (MEL).

PAPUA: Western Division: New Guinea, near Dutch boundary, Mac-
Gregor in 1890 (MEL); Fly R., Baéuerlen 514, Nov. 1885, on red clay banks;
Lake Daviumbu, Middle Fly R.,, Brass 7590, Aug. 1938, forming dense thickets
2.5-3 m. high, in edge of forest along lake shores (det. Kiikenthal as S. scrobi-
culata Nees); Lake Daviumbu, Middle Fly R., Brass 7663, Sept. 1936, savan-
nahs, scattered clumps 1.5 m. high on swamp margins (det. Kiikenthal as Scleria
chinensis Kunth); Lower Fly R., east bank opposite Sturt Island, Brass 8115,
Oct. 1936, tufted in semi-shade on edge of sago =r (det. Kikenthal as
S. scrobiculata Nees). Gulf Division: Ghu, Vaitata R., Brass 938, Feb.
1926, rain-forest borders, small clumps 2-3 ft. high. Central Division:
Boku, Schlenker 9, July 1909; Deva Deva, White 588, alt. about 1200 m.,
July-August 1918 (large sedge about 6 ft. high); Kubuna, Brass 5563, Nov.
1933, alt. 100 m., common amongst ferny ground cover in rain- forest; Astrolabe
Range, White 359; Sogere, White 370, July-August 1918; Bisiatabu, ‘Bras ss 584,
Nov. 1925, alt. 1500 ft., banks of streams; Owen Stanley Range, between Mts.
Brown and Clarence, Brass 1479, May 1926, alt. 4000 ft. Eastern Divi-
sion: Fife Bay, Turner 95, Sept. 1930 (plant about 4 ft. high); South Cape,
Chalmers Sonne “south-east New Guinea,” Chalmers in 1878; Samarai, Fitz-
gerald 7, common (more or less caespitose, 1-3 ft. high); Dixon’s Bay, Rossel
Island, ages "S Bridge, Jan. 1885; islands near the south-east coast, Armit
in 1

1954] BLAKE, CYPERACEAE COLLECTED IN NEW GUINEA 231

SOLOMON ISLANDS: Without definite locality, Wernhem, Jan. 1911
(Nsw).
NEW HEBRIDES: Without definite locality, Haer 65 in 1902 (Nsw).

Widely spread from New Guinea and Queensland through Melanesia
to eastern Polynesia. Not previously recorded from the Solomon Islands.

This species has usually been called Scleria margaritifera Willd., which
is a later homonym of Scleria margaritifera Gaertn., a name that has been
overlooked by most botanists and by the compilers of Index Kewensis.
Gaertner gave a good account and figure of a portion of the inflorescence,
spikelets and fruit. He cited several synonyms, including Carex litho-
sperma L. and Scleria flagellum-nigrorum Berg. Core, Brittonia 2: 87.
1936, referred Gaertner’s name, figure and description to S. flagellum-
nigrorum Berg., which would seem to be its correct disposition; it is
accordingly a superfluous name for this species. Willdenow, Sp. Pl. 4: 312.
1805, cited S. margaritifera Gaertn. as one of the synonyms of “Scleria
flagellum Sw.,” but on the next page deliberately used the same name for a
species which he described as new from a specimen collected by Forster
on the island of Tanna. Hence Scleria margaritifera Gaertn. and Sclerta
margaritifera Willd. are quite distinct names, and both of them are illegiti-
mate under any circumstance.

I have not seen the types of S. polycarpa, S. graeffeana or S. ternifolta,
but I have seen specimens from the type-localities of each that agree with
the original descriptions. The species is closely allied to S. scrobiculata
Nees, differing from it chiefly in the narrower partial panicles with fewer
and less spreading branches, the inconspicuous bracteoles shorter than the
spikelets, the relatively inconspicuous male spikelets, the more evenly dis-
tributed fertile spikelets, the nut more gradually narrowed to the tip and
nearly smooth to slightly rugulose at maturity, and the less deeply divided
disc with more prominently denticulate margins; the nuts are frequently
bright blue with red discs. The leaf-sheaths in both species may be broadly
or narrowly winged or quite wingless on different individuals. Both species
are robust plants with the middle leaves in false-whorls of three, relatively
numerous partial panicles, medium-sized nuts which are also more or less
hirtellous, and rather short discs.

The following references to Scleria scrobiculata belong wholly or in part
to S. polycarpa: K. Schum. & Hollr. Fl. Kaiser Wilhelmsl. 25. 1889; K.
Schum. & Lauterb. Fl. Deutsch. Schutzgeb. Siidsee 198. 1901; F. M. Bail.
Queensl. Agr. Jour. 23: 220. 1909; Valck. Suring. Nova Guin. Bot. 8:
712. 1912; Kiikenth. Bot. Jahrb. 59: 58. 1924.

Scleria poaeformis Retz. Obs. 4: 13. 1786.
Scleria oryzoides Presl, Rel. Haenk. 1: 201. 1830; F. Muell. Pap. Ph. 2-51;

A: Western Division: Fly R., Bauerlen 562, Nov. 1885 (MEL);
Lake Daviumbu, Middle Fly R., Brass 7855, Sept. 1936, in dense pure stands
dominating many large swamps on savannahs; Gaima, Lower Fly R. (east bank),

232 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXXV

Brass 8261, Nov. 1936, swamps in savannah forest area; Wuroi, Oriomo R.,
Brass 5748, Jan—March 1934, alt. 10-30 m., in dense formation in a small
swamp; Daru Island, Brass 6338, March 1936, forming pure stands, 1.5 m. tall,
in shallow open swamp in savannah forest

Bauerlen’s specimens were cited by F. Mueller, l.c.; Brass’s specimens were
received determined by Kikenthal as S. oryzoides Presi. My recent congo
of the type of S. poaeformis (Lp) confirmed the opinion expressed by C. E. C.
Fischer in Kew Bull. 1931: 265. 1931 that this name and S. oryz sites are
synonymous. The species extends from northern Queensland northward and
westward to India

Besides the species mentioned in the foregoing pages, the following have
been recorded for New Guinea, based on specimens that I have not seen:
Scleria hookeriana Boeck.; Kikenth. in Eng. Bot. Jahrb. 59: 59. 1924.

Scleria levis Retz. forma villosa Valck. Suring. Nova Guin. Bot, 8: 712.
912; Kukenth. Bot. Jahrb. 59: 58. 1924; Ohwi, Bot. Mag. Tokyo
56: 212-3. 1942.

Scleria brownii Kunth: Kiikenth. Bot. Jahrb. 70: 464. 1940.

The record of S. brownii is very likely based on specimens of S. novae-
hollandiae (see under this species) ; the other records may also be based on
specimens of species discussed elsewhere in this paper.

The following key will serve to distinguish the species seen from New
Guinea:

Spikelets — at least the fertile ones — androgynous; disc reduced to an indis-
bas

tinct al rim or (in S. motleyi) more or less cupshaped and thin; nut
trigonous.

Annual, lemon-scented; spikelets in small distant clusters along a simple

common axis without prominent bracts... ergracilis.

Perennial; inflorescence of terminal and axillary partial aie with promi-
nent bracts, the partial panicles sometimes spike-li
Disc seh saa nut glabro

Nut quite smooth, even when young.................. S. lithosperma.
Nut rugose, even at extreme maturity.................. S. roxburghii.
Disc cup-shaped; nut hirtellous with ferruginous hairs. oo ..... S. motleyt.

Spikelets unisexual, the females commonly with one or more empty glumes above
the flowers; disc always present, shallowly to deeply 3-lobed; nut not dis-
tinctly angular.

Annual plants inieed up to 50 cm. high; culms about 1 mm. wide or less;

eaves 1-3 mm.

Nut ellipsoid or ‘cylindroid. Ati prominently apiculate; terminal partial
panicle longer than the oth

Disc deeply divided with ely separate ovate to sited more or less

acute lobes; nut deeply tessellate, more or less shining .. _S. tessellata.

Disc small and thin, shallowly lobed with broadly rounded — nut

dull, smooth or somewhat verrucose or somewhat reticulate...._...

1954] BLAKE, CYPERACEAE COLLECTED IN NEW GUINEA 233

Nut globular, A piiopiecaiay apiculate, at first reticulate, finally often smooth
except for a few tubercles on the upper part; disc thick, shallowly lobed;
partial eee SU SEARED oes ha Suk eee 828 ee mee S. rugosa.

Perennials up to 3 m. high or more with prominent rhizomes; culms at least
mm. wide and usually wider; leaves 4-20 mm. wide.

Spikelets mostly in pairs, the lower one of most ats fertile; inflorescence

of 3-8 partial panicles, the lower ones from long, leaf-like bracts; leaves

long-tapering to a narrowly a or filiform tip; rhizome more or less

knotty with approximated culm

Ligule longer than (usually 2-3 oe as long as) wide; bracteoles un-

usually prominent and long exserted from the dense partial panicles

jaris.

Ligule ne (usually much shorter) than wide; bracteoles usually less
t, or setaceous; partial panicles looser- flowered or subspici-

a:
Leaves all about equally distributed along the culm; bracteoles prom-

inent.
Disc-lobes lanceolate, more or less acute, more or less toothed ....
a

Fe eo 5 8 oe ea ob eee es levis.
Disc-lobes broadly rounded, entire ................. S. terrestris.
Leaves clustered in groups of 3 mee 2) at the base of and below the

inflorescence.
Bracteoles wee! as long or as long as the more or less spreading
s of the partial panicles; nuts borne chiefly towards
the fuer of the branches, abruptly rounded below the mucronate
tip, commonly deeply scrobiculate and white... S. scrobiculata.
Bracteoles inconspicuous, much shorter than the more or less erect
b hes of the more or less spiciform partial panicles; nuts
evenly distributed throughout the partial panicles, rather grad-
ually narrowed to the mucronate tip, rugulose smooth at
maturity, often tinged blue; disc often red... ... S. polycarpa.
Spikelets not paired, evenly distributed along the branchlets, the fertile
ones restricted to the lower part of the lower branches; inflorescence
of a pues pedunculate compound compact terminal panicle; leaves
broad, scarcely tapering to the obtusely rounded tips; rhizome elongated
Wilh tater -Cistant CUNMS~ 2.5 so-<s44 4 sas Soumeeee ent S. poaeformis.

Diplacrum R. Brown

Diplacrum caricinum R. Br. Prodr. 241. 1810.

Scleria caricina (R. Br.) Benth. Fl. Austral. 7: 426. 1878; Kukenth. Bot.

Jahrb. 69: 261. 1938.

Scleria axillaris Moon, Catal. Pl. Ceylon 62. 1824, nomen nudum.

Diplacrum tridentatum Brogn. Duperr. Voy. Bot. ¢. 26. 1826.

Diplacrum zeylanicum Nees in Wight, Contrib. Bot. Ind. 119. 1834.

Olyra malaccensis Wall. ex Kunth, Enum. PI. 2: 360. 1837, pro syn.

PAPUA: Western Division: Lake Daviumbu, Middle Fly R., Brass 7842,
Sept. 1936, common on wet plains, also in edge of Tristania forests.

New for Papua; widely distributed in Malaysia, extending into Ceylon,
India, South China and Queensland.

234 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

The limits of Diplacrum and Scleria have been variously circumscribed
and the two groups have often been treated as congeneric. A few American
species have sometimes been referred to Diplacrum, but they are best
treated as a separate genus, Peteroscleria Nees. The more important
characters differentiating the three genera are as follow:

Scleria: Male spikelets (or male portion of androgynous spikelets) borne
above the female; stamens commonly 3, Female spikelet: glumes 3-6
below the flower and usually one or more reduced ones above, 1-
nerved, the keel not winged, entire at the tip, permanently mem-
branous, persistent after the fall of the ripe nut. Nut smooth or
variously reticulate or tuberculate or transversely rugose, the trans-
verse ribbing at least as prominent (usually more prominent) than
the vertical. Outer disc usually present. Leaves in the middle of the
stem much the longest, the uppermost (bracts of partial panicles)
gradually much shorter and narrower, the lower ones gradually re-
duced to bladeless sheaths. Partial inflorescences in the upper part of
the stem, or inflorescence entirely terminal.

Diplacrum: Male spikelets borne below the female: stamen commonly
one. Female spikelet: glumes 2, 3—7-nerved, keeled but the keel not
winged, more or less 3-lobed or 3-toothed, hardened at maturity and
commonly falling with the ripe nut which they enclose; no glumes
above the flower. Nut tending to be heavily ribbed vertically, trans-
verse ribs less pronounced or irregular or absent. Outer disc absent.
Leaves or bracts all very similar, all or nearly all with axillary heads
of spikelets, 1 or 2 basal ones sometimes reduced to sheaths.

Pteroscleria: Male spikelets below the female; stamens commonly 3 in
the lower flowers. Female spikelet: glumes 2, the keel prominently
winged, otherwise nearly nerveless, entire at the tip, not hardened at
maturity (?), persistent (?); no glumes above the flower. Nut tend-
ing to be ribbed vertically, but the ribbing faint or obscure. Outer
disc absent. Leaves dissimilar, only the upper ones with axillary
heads of spikelets.

As thus delimited, Diplacrum comprises a group of about five small
slender annual species of the Old World Tropics. Diplacrum caricinum
is the type-species and is the most widely spread. Endemic species occur
in Malaya (D. reticulatum Holttum), Africa (D. africanum C. B. Clarke)
and Australia (D. pygmaeum |R. Br.| Nees ex Boeck. and another).

Uncinia Persoon

Uncinia riparia R. Br. Prodr. 241. 1810.

NETHERLANDS NEW GUINEA: Mt. Wilhelmina, 7 km. NE. of Wil-
helmina-top, Brass & Meyer-Drees 9847, Sept. 1938, alt. 3560 m., common in
weak clumps on mossy floor of subalpine forest.

1954] BLAKE, CYPERACEAE COLLECTED IN NEW GUINEA 235

PAPUA: Central Division: Mt. Albert Edward, Brass 4415, May-July
1933, alt. 3680 m., sporadic on floor of forest.

New for New Guinea; previously known from New Zealand, Tasmania
and the mountains of the extreme SE. Australian mainland.

The previous records of Uncinia riparia from New Guinea refer to other
species. That of F. Mueller, Trans. Roy. Soc, Vict. 1 (2): 36. 1889 and
of Kikenthal, Pflanzenr. 4 (20), 38: 63. 1909 and Bot. Jahrb. 59: 59.
1924 refer to specimens of U. sclerophylla Nelmes, Kew Bull. 1949: 143.
1949. According to Nelmes, l.c., pp. 142-5, Kiikenthal’s reference in Bot.
Jahrb. 69: 261. 1938 was based on collections in which U. sclerophylla
Nelmes, U. subtrigona Nelmes and perhaps another species are represented.
The Clemens collections are so badly mixed and sometimes so poor that it
is very risky to interpret references by the examination of alleged dupli-
cates

ADDENDA

In a parcel of old specimens from New Guinea received for determination
from the Melbourne Herbarium there were found a specimen of an un-
described species of Hypolytrum and a specimen of a species of Cyperus
previously unrecorded from New Guinea. They are discussed here, to-
gether with other corrections and additions to the previous contributions.

Hypolytrum L. C. Richard
Hypolytrum microcarpum sp. nov. PLATE I.

Culmi e rhizomate brevi caespitosi, circa 50 cm. alti, 2 mm. crassi,
triquetri, lateribus concavis striati, angulis anguste obtusis prope apicem
scaberuli, ceterum laeves. Folia basalia linearia, chartacea, 6—9 mm. lata,
usque 37 cm. longa, basim versus complicata haud vel vix angustata, apice
acutata, prope apicem marginibus nervoque mediano scabra, ceterum
laevia; folia caulina 2, conspicue vaginantia, summum inflorescentiam
superans, basalibus simillima nisi basim versus angustata. Bracteae in-
feriores foliiformes, inflorescentiam superantes. Inflorescentia suboblonga,
subcorymbosa, circa 5 cm. longa lataque, multiflora; axis communis
angulosa scabra; rami divaricati, usque ad 22 mm. longi, compressi, scabri,
prope apicem corymboso-ramosi ramulis nonnullis eodem modo divisis;
ramuli ultimi (pedicelli) usque ad 5 mm. longi, filiformes. Spiculae fusco-
brunneae obovoideae vel oblongae, 3.5—4 mm. longae, sub flore circa 2 mm.
sub fructu circa 3 mm. latae, pauciflorae. Glumae (explanatae) orbiculari-
obovatae, muticae, brunneae, marginibus (apice ipso excepto) subscariosae,
1.3-1.5 mm. longae. Flores 1.3 mm. longi; squamellae more generis 2,
liberae, carina parce ciliatae, explanatae lanceolatae obtusae, circa 1.3 mm.
longae. Nux subglobosa, leviter compressa, circa 1.4—1.5 mm. longa, 1.3—
1.4 mm. lata, brevissime umbonato-rostrata, bicostulata, irregulariter laxe-
que ruguloso-reticulata, dilute flavo-brunnea sed creberrime rubro-
punctata.

Jour. ArNoLp Ars. VoL. XXXV PLATE I

HyYPOLYTRUM MICROCARPUM S, T. BLAKE

1954] BLAKE, CYPERACEAE COLLECTED IN NEW GUINEA VY |

PAPUA: Eastern Division: Cloudy Mountains near South Cape,
Chalmers & Bridge in 1884 (TYPE in MEL).

The description is based on a single specimen consisting of a fruiting
culm and two innovation-shoots all attached to the rhizome. It has the
habit and general appearance of H. Jatifolium L. C. Rich., H. scirpoides
(Presl) Merr. (H. philippense C. B. Clarke), H. scabrum Uitt. and H.
vitiense C. B. Clarke, but it differs from all of these in the much smaller
spikelets, glumes, flowers and nuts and the nearly globular nut with its
beak reduced to a very small umbo. The nearly smooth leaves and stem
further distinguish it from H. scirpoideum and H. scabridum and the
wrinkling on the nut from H. vitiense. It must also be closely allied to
H. minus Ridl.; I have seen no specimen of this species, and although
Ridley’s description leaves much to the imagination, the stress placed on
the scabrous margins of the much broader leaves suggests that it is closer
to H. scirpoideum or identical with this, as suggested by Kukenthal, Bot.
Jahrb. 59: 53. 1924.

Cyperus Linnaeus
Cyperus cinereobrunneus Kiikenth. Mitteil. Thiiring. Bot. Ver. N. F.
50: 3. 1943; Kiikenth. ex S. T. Blake, Jour. Arnold Arb. 28: 216.
1947.
I had not seen Kiikenthal’s paper when I published his name in 1947.

Cyperus globosus All.; S. T. Blake, Jour. Arnold Arb. 28: 220. 1947.
Cyperus globosus All. var. oblonginux Kikenth. Mitteil. Thiiring. Bot. Ver.
N. F. 50: 7. 1943.

I had mentioned (l.c.) that, in the determination on the label, Kikenthal
had distinguished Brass 8309 as a variety; I had not then seen the des-
cription, These plants seem to be no more than an individual variation.

Cyperus fulvus R. Br. var. confusus (C. B. Clarke) Kukenth. Pflan-
zenr. 4 (20), 101: 456. 1936

Mariscus fulvus (R. ne) ie B. Clarke var. confusus C. B. Clarke ex Domin,
Biblioth. Bot. 85: 444.

PAPUA: Central Division: Quaipo, MacGregor in 1889 (MEL).

The species is new for New Guinea, but is common and widely spread

PLATE I

Hypolytrum microcarpum S. T. Blake. Type specimen X about % with
analytical drawings. Fig. 1. Glume. Fig. 2. Nut. Fig. 3. Transverse section of
nut. All figures X about 10. Analytical drawings by S. T. Blake, photography
by G. Cripps, Photographic Section, Department of Agriculture and Stock,
Brisbane.

238 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XXXV

in NE. Australia. Although quite variable, it is doubtful how much im-
portance can be assigned to the varieties that have been described.

Schoenus Linnaeus

Schoenus maschalinus R. & S. Syst. Veg. 2: 77, 1817; S. T. Blake,
Proc. Roy. Soc. Queensl. 60: 47. 1950.

Schoenus foliatus (Hook. f.) S. T. Blake; S. T. Blake, Jour. Arnold Arb. 29:
93. 1948.

Later botanists have overlooked the fact that Roemer and Schultes cor-
rectly gave a new name to Chaetospora axillaris R. Br. when this was
transferred to Schoenus. An extensive and complicated synonymy has be-
come associated with the species, which has been discussed in my papers
cited above.

Rhynchospora Vahl
Rhynchospora triflora Vahl; S. T. Blake, Jour. Arnold Arb. 29: 101.
1948.

Rhynchospora triflora Vahl var. papuana Kikenth. Bot. Jahrb. 74: 427. 1949.

Rhynchospora triflora var. papuana was founded entirely upon Brass
8356; I cannot distinguish our specimen of this from our specimen of
Henderson in Singapore Field no. 24101 which Kikenthal cited under
R. triflora.

Carex Linnaeus

Several emendations to my determinations in Jour. Arnold Arb. 28:
99-116. 1947 appeared in a paper by E. Nelmes in Kew Bull. 1949:
378-386. 1949. This was followed by a revision of the Malaysian species
in Reinwardtia 1: 221-450. 1951. As pointed out in the introduction to
my paper, I had to place considerable reliance on descriptions only; Mr.
Nelmes has had a much wider experience in this genus, and his opinions
should have preference.

(QUEENSLAND HERBARIUM,
BRISBANE, QUEENSLAND, AUSTRALIA.

1954] COTTAM, PREVERNAL LEAFING OF ASPEN 239

PREVERNAL LEAFING OF ASPEN IN UTAH MOUNTAINS
WALTER P. CoTTAM

With two plates

Tue Rocky MounrTAIN ASPEN (Populus tremuloides Michx., var. aurea
Tidestrom) dominates more mountainous terrain in Utah at elevations
between 7,000 and 10,000 feet than any other forest species. Several
features of its growth-form, together with its peculiar autecology and
synecology, make this species a very conspicuous forest type. Unlike all
other prominent forest species with which it may be associated, such as
Douglas fir, white fir, alpine fir, blue spruce, Engelmann spruce, ponderosa
pine and lodgepole pine, the aspen is deciduous in habit.

Often the aspen occurs in almost pure stands, which may vary in size

from a few square rods to several square miles of solid forest. But
whether small or large, the aspen stands are sharply discontinuous, single
aged and usually dense. The shiny green leaves of summer, the multi-
colored shades of yellow in autumn, and the slender, straight, white boles
at all seasons of the year are features of the aspen that set it apart in
contrast to all contiguous association types, whether mountain conifers
or wse.
But the contrast that exists between the aspen stands and other moun-
tain vegetation types is often exceeded, in early spring at least, by striking
differences in the time of leafing within the aspen groves themselves. This
phenomenon is widespread throughout the mountains of Utah and neigh-
boring states. In early spring almost any aspen forest shows sharply
discontinuous colonies of trees that attain full leaf two or three weeks
earlier than the major surrounding stand in which at any other time they
are likely to remain unnoticed except by the critical observer.

At mid-elevations these colonies of prevernal aspen are usually small
in comparison with the forest as a whole, and their outline is variable.
Frequently they assume a circular shape, but they may be seen as a
narrow, serpentine band bisecting large groves. In general, the position
of these prevernal colonies seems to have no correlation with slope, ex-
posure, soil, altitude, or sex of trees, but wherever they occur or whatever
shape they may assume, the line that separates them from the retarded
major population of trees is sharp, and rarely do the two forms intermingle
as individuals.

CONTROLS OF PREVERNAL LEAFING

For years the author has been intrigued with the causes underlying
the prevernal leafing of aspen colonies and surprised at the lack of text-
book reference to this phenomenon. But few facts of aspen ecology

240 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

escaped the critical eye and pen of F. S. Baker, who in 1921 wrote an
excellent description of the prevernal leafing of aspen colonies with special
reference to the Wasatch Plateau of central Utah. His paper (4), en-
titled ‘‘Two Races of Aspen,” points out certain taxonomic differences be-
tween early- and late-leafing stands and concludes: “The writer confesses
an entire inability to explain the causes of these two widely distributed and
closely intermingled races of aspen.” Yet throughout the article Baker
implies that the causes, whatever they may be, are genetic and never
environmental.

There can be little doubt that at least two and probably several races
of aspen exist that show striking differences in time of leafing. Evidence
of these strains is seen in the variable combinations of certain morphologi-
cal characters often associated with the physiological functions that
regulate leafing response. These morphological characters are sufficiently
distinct and constant as to enable one to distinguish the races at the
seasons of the year when leaf color and size no longer offer safe clues to
their identity. For example, Baker pointed out that in the Wasatch
Plateau area boles of the early variety are yellow-green in color in contrast
to the powdery-white appearance of the late-leafing variety. The writer
has found no exception to this phenomenon throughout the northern half
of Utah. This bole color difference is due to the failure of the early-leafing
form to produce the usual copious bloom on the bark exterior character-
istic of the late-leafing variety.

Another character generally useful in distinguishing the leafing strains
of aspen is the pruning habit of the trees. In central and northern Utah,
early-leafing aspens in general fail to prune themselves of the lower, small,
dead branches on the trunk, thereby presenting an appearance of low vigor
and untidiness entirely foreign to the late-leafing strain. (Careful observa-
tion, however, discloses that this lack of vigor is more apparent than real.)

Thus, in central and northern Utah, prevernal leafing of aspen is as-
sociated with the apparently genetic characters of yellow boles and poor
pruning. However, on the Aquarius Plateau of south central Utah, the
early-leafing character is associated with white boles and clean pruning,
while the late-leafing trees are yellow-boled, non-pruning, a complete
reversal of genetic alignment of the northern form.

In distribution within the aspen belt, early-leafing varieties dominate
the higher elevations only. At the upper limits of the belt, aspens exist
mainly as dwarfed thickets, which are always of the early-leafing forms as
distinguished, of course, by the bole color and pruning habit. Conversely,
the lower limits of the aspen belt are dominated by late-leafing varieties.
Thus the contrast between early- and late-leafing aspens is most pro-
nounced in the spring at mid-elevations, where the two forms are often
intermingled as colonies. Here also is found the greatest contrast in
autumn, for the early-leafing forms, as a rule, maintain their green color
several days longer in the fall than do the late-leafing varieties.

For several years the writer has observed a few areas where contiguous
colonies of early- and late-leafing aspen fail to present the morphological

1954 | COTTAM, PREVERNAL LEAFING OF ASPEN 241

differences mentioned above. In all of these exceptions the boles of both
aspen groups are white and well pruned, yet the length of the leafing time
differences is the usual two to three weeks. One such area is located at
the head of South Willow Creek Canyon in the Stansbury Mountains,
approximately fifty miles west of Salt Lake City. Another is found in the
Abajo Mountains west of Monticello in southeastern Utah. Both areas
occur at an elevation of about 7,500 feet.

TEMPERATURE CONTROLS OF PREVERNAL LEAFING

The Stansbury group appears at the bottom of an east-west canyon at
a point where the generally straight canyon forks abruptly. The major
fork heads south and in a circuitous manner drains the summit of the
10,000 foot range. The minor fork proceeds west for approximately one
mile where it ends more or less abruptly, forming a sort of ‘‘boxed”’ seg-
ment rather well protected from air drainage from the summit above.
Each spring in late May, aspens in this box canyon may be seen in almost
full leaf with a sharp line scarcely more than a rod wide bisecting the
grove and separating those in leaf from a half mile area of leafless aspens
lying at and below the confluence of the major south fork.

Here there is an anomalous condition in which a grove of aspens leaf
out considerably before their neighbors two to three hundred feet below
them in elevation. In order to ascertain whether temperature might be a
controlling factor in this phenomenon, a transect of four stations (two
above and two below the leafing line) was established. Recording thermo-
graphs, fastened to the aspen branches ten feet above the ground and
protected by white canvas tents, were used in this preliminary experiment,
as well as in others to be reported below. Readings were begun May 7 and
terminated May 19, 1951. During this period neither group of aspen had
progressed beyond the swollen bud stage. Figure I shows the daily mini-
mum, maximum and mean temperatures in both aspen groups over this
period of 13 days.

The comparative summaries given in Figure I leave little doubt that
these two major stands of aspen, showing marked differences in the time
of leafing, exist under strict temperature controls and are not separate
races. Considering the fact that the early-leafing aspen while occupying
the higher elevations show a daily mean temperature two to three degrees
greater than the late-leafing stands, these temperature differences seem
highly significant. Translated into altitudinal effects, the temperature
difference between these two aspen groups is equivalent to approximately
1000 feet in elevation. Furthermore, observations in this area in early
June, 1951, disclosed several typical prevernal colonies of genetically con-
trolled aspen within the major retarded zone of lower temperature. These
small isolated colonies of yellow-boled aspen came into leaf at approxi-
mately the same time as the large stand of white-boled aspen of the higher
temperature area.

242 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

GENETIC CONTROLS OF PREVERNAL LEAFING

Circumstances prevented the writer from visiting the prevernal leafing
aspen colonies of the Abajo Mountains of southeastern Utah until August,
1951. With the aid of kodachrome pictures taken in May, 1950, the exact
position of these colonies was ascertained. Comparative temperature records
were taken for a period of four days. No significant temperature differ-
ences between the early-leafing and late-leafing stands were found, The
uniformity of the topographical features suggests, moreover, that there
should be none. While considerably more temperature data are needed
for this area, the evidence points to the probability that early-leafing and
late-leafing colonies of aspen here represent different strains un-
associated with observable bole and pruning charact

Continuous temperature data from June 1 to Sa abe 30, 1951,
except for two interruptions of four days each, were secured from con-
tiguous colonies of prevernal and late-leafing aspen at Mt. Timpanogos
located in the Wasatch Range 30 miles southeast of Salt Lake City. These
colonies, occupying mid-altitudinal positions of about 8,000 feet, all
showed the morphological differences described earlier, as well as the

FicurE I
MAY
952 | 7 8 9 10 { UI i2 | 13 | 14} 18 | 16 | 17 | 18 | I9
6
awe DAILY MAXIMUM TEMPER tery
--7T , Zé iY .
55 - ~ | RA
Sy
, a
y 45—< ‘\ 4
S }
Ww I i :
> /
rod DAILY MEAN TEMPERATURES
= & ——
< 50 ae oes 8
a aN aa
% Le IN “
Wl 40}-—>++ = se
oO ay ee ee oad
Ww . Le
Q 30 fa reten
DAILY MINIMUM TEMPERATURES
Ps
40 a Z
| ders, a te
ae: be el
30 Prat ard KL L— | 7 -
iis i. dl as al |
—— EARLY
----- LATE

1G. I. Temperature phenomena of adjacent stands of early- and late- aaa
aspen, eee Mountains, Utah. Solid lines early. Broken lines late

1954] COTTAM, PREVERNAL LEAFING OF ASPEN 243

conspicuous differences in leafing time. Figure II represents weekly sum-
maries of the averages of temperature data for two stations each in a pair
of contrasting colonies selected for their apparent uniformity of soil and
topographical features. Intermittent data from two other pairs of con-
trasting colonies were also secured throughout the summer. These con-
firmed the results shown in Figure II

The data in Figure II reveal that the maximum weekly temperature
averaged slightly higher and the minimum temperature averaged slightly
lower in the late-leafing colony than in the early-leafing stand, although
the mean weekly temperature averages were essentially similar in both.
These slight temperature differences may be accounted for by the fact that
the late-leafing stand, being slightly more open than the early-leafing colony,
allows more sun on the tents by day as well as more effective air circula-
tion by night. However, no aspect of these temperature phenomena is
sufficient to account for the great discrepancy of leafing time for these
colonies, and the conclusion seems justified that the cause of prevernal
leafing lies outside of environmental controls.

Ficure II
JUNE | JUL | AUGUST | SEPTEMBER
| | T
1-8| 8 mie 22| 27-4) 8-15] 15-22) 22-29 29-5 | 5-12!) 12-19 9-2426-2| 2-9 9-16} 16-23) 23~
‘=te)
7 a hos ay, IN Z ~
72 NS oa Fee ey oe ee
60 sta
Ee eo t
re AVERAGE WEEKLY MAXIMUM TEMPERATURES
r eae) |
« ;
= a, ee oa
qt 60 — ">>> y Peek
w len Tg Sco chee i
iW 5D PA |
ce 7 Rey T T
re) ssl | AVERAGE WEEKLY MEAN TEMPERATURES
a 40 |
50 lS re eagle Bre
a ee TI AO
Aa a = hy a
40 2 > as
EZ
aeahe AVERAGE WEEKLY ASI) TEMPERATURES
aA , LAST SPRING FROST JUNE 3 FIRST AUTUMN FROST SEPT. ||
as
=—— EARLY
See CATE

Fic. Il. Temperature phenomena of adjacent stands of early- and late-leafing
aspen, Mt. Timpanogos, Utah. Solid lines early. Broken lines late.

In order to test the hypothesis that the two distinct leafing forms of
aspens possessing recognizable morphological differences at Mt. Timpano-
gos are true genetic races, other methods of approach seemed desirable:
First, the transplanting of aspens from one colony to another, as well as

244 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

to a common habitat; second, investigations on the clonal features of the
aspen; and third, cytological studies.

TRANSPLANTING EXPERIMENTS

Eight aspen sprouts from three separate pairs of colonies were trans-
planted in the fall of 1951, thereby introducing into prevernal colonies
twelve late-leafing saplings and into late-leafing colonies an equal number
of early-leafing saplings. Despite the fact that care was exercised to locate
the transplants on the edge of the colonies where competition for light and
moisture could be reduced to a minimum, none of the aspen sprouts
survived the winter and summer of 1952.

In this experiment it was recognized that several years must elapse
before conclusive results might be expected, assuming successful trans-
plantings were possible, because observations have disclosed the fact that
aspen sprouts must attain considerable height before they acquire either
the morphological characters or the leafing peculiarities of the mature
stand. The retarding of the leafing time of the sprouts in prevernal
colonies is especially noticeable and may be accounted for by the fact that
leafing of the mature trees is generally well advanced before the snow pack
has disappeared from the stand. Obviously the close proximity of the
snow to the sapling buds has a retarding effect on leafin

In addition to these transplantings in the field, four saplings from late-
leafing aspen stands and four saplings from early-leafing colonies were
removed to the University of Utah campus, where previous transplantings
of aspen sprouts have proved successful. These transplants were made in
late May, 1952, to a favorable situation where nearly uniform factors of
soil, water and light were maintained for all saplings.

Three saplings from the late-leafing variety and two from the early sur-
vived the summer of 1952 and the winter of 1953. In the spring of 1953
both early-leafing sprouts came into leaf more than two weeks before the
late-leafing transplants. Thus the leafing character of the two mountain
strains was maintained under totally different environment, and the hy-
pothesis of genetic controls received considerable support.

CLONAL CONNECTIONS OF ASPEN COLONIES

It was assumed that if the present strains of aspen had their origin in
seed mutation, contiguous colonies of early- and late-leafing forms should
show no clonal connections, although such connections might be expected
to exist between trees within the same colony. To determine the nature
and extent of aspen esere connections, at Mt. Timpanogos radioactive
phosphorus was employe

Selected for radioactive a treatment was a white-boled, late-
leafing tree which occupied a position between closely contiguous colonies
of distinct strains of aspen. On one side the branches of this tree inter-

* The writer is indebted to the Research Committee, University of Utah, for
supplying the necessary radioactive phosphorus used in this experiment; to Dr. Robert
C. Pendleton for oe the tree; and to Dr. John D. Spikes for monitoring the
laboratory specimens

1954] COTTAM, PREVERNAL LEAFING OF ASPEN 245

mingled with those of early-leafing trees and on the other with branches
of late-leafing individuals. The nearest bole of the late-leafing variety was
7 feet 6 inches distant, and the nearest bole of the early variety was 6
feet 6 inches. There were six early-leafing trees and seven late-leafing ones
within a radius of 17 feet. Near the tree selected for labeling, a trench
was excavated exposing the roots, one of which was severed about three
feet from the bole. The cut end of this root was inserted into a gallon
jar of water containing 40 millicuries of radioactive phosphorus (P-32).

Within 36 hours most of the radioactive phosphorus solution had been
absorbed by the tree. The tube of a Geiger counter was placed in a
rectangular hole cut breast high in the bark of the labeled tree, thereby
exposing the cambium. The counter showed a radioactivity of 60,000
counts per minute. No other tree of either variety in the vicinity of the
labeled tree showed radioactivity above background. In subsequent days
portions of the roots, cambium, wood and leaves of the labeled tree and
of the early- and late-leafing trees surrounding it were removed to the
laboratory. These specimens were ashed and tested with a standard scaler
provided with a thin-end, window Geiger tube. All ashed samples from
the labeled tree gave more than 50,000 counts per minute. No radio-
activity above background was found in the samples of any other tree.
Excavations around the labeled tree showed that all roots belonging to
the labeled tree were radioactive, but that the radioactive substance had
not passed to the roots of neighboring trees of either strain

The results of this experiment are indeed surprising, for they suggest
complete separation of the aspen sprouts from the parent clone before or
soon after maturity. Considerably more investigation is necessary to
establish the clonal characteristics of the aspen.

CYTOLOGICAL TESTS

The field of cytogenetics should offer an interesting and possibly
a fruitful approach to the problem of the leafing strains of aspen. So
far, however, the writer has encountered perplexing delays in securing
suitable tissue for study. Because of the unreliability of aspen seed pro-
duction, and the major difficulties of securing pollen smears from the
remote and snow-bound experimental area, it was assumed that cuttings
would provide the most feasible source for securing tissue in active cell
division. Numerous greenhouse attempts have been made to root aspen
cuttings without a single positive result to date. Additional effort to
secure root tips for study is currently in progress.

Positive miscroscopic evidence of chromosome aberrations would, of
course, establish the validity of these genetic strains, but negative evi-
dence would not necessarily preclude it, for mutations may be the result
of one or a few unobservable gene changes

DISCUSSION

Whether cytologically demonstrable or not, the fact or the assumption
that these readily distinguishable forms of early- and late-leafing aspen

246 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

represent mutant strains presents problems of origin and subsequent
distribution both interesting and perplexing to the ecologist. For no fact
of aspen ecology, in Utah at least, seems more certain than that aspen
trees in general, despite occasional abundant viable seed production,
reproduce only through vegetative means. They do this because aspen
seeds must germinate within a few weeks after ripening in early June.
Under present climatic conditions precipitation in Utah is invariably
scant and irregular during the summer months. Thus the wide and spotty
distribution of aspen throughout Utah today must have been attained
under climatic patterns of more abundant and more evenly distributed
summer precipitation. Such conditions are postulated by geologists for
the Great Basin region during the pluvials associated with the extensive
Pleistocene glaciations (1, 2 & 3

After the onset of the Postpluvial climate in the Great Basin area, which
Antevs estimates at approximately 8,000 years ago, it is likely that aspen
ceased its reproduction by seeds. Since then migration has been through
clonal reproduction only, and exceedingly slow, but sufficient to effect
the mergence of many previously separated colonies and to extend the
upper altitudinal limits of the aspen belt. Conversely, the increased
temperature and lower summer precipitation of the Postpluvial may have
caused considerable retreating migration from lower limits of the Pluvial
aspen distribution. There are examples of the complete disappearance of
aspen colonies at lower elevations during historical times, but grazing
influence may have been a contributing cause.

Any proposed hypothesis to explain the origin and present-day distribu-
tion of early- and late-leafing aspens must account for the following facts:
1. The predominance of late-leafing forms at low and mid-elevations.

2. The predominance of early-leafing aspen at higher elevations.
3. The widely intermingled colonies of the two forms at mid-elevations,
their sharp discontinuity and the purity of their stands.

Assuming that mutations have occurred within the sex cells and there-
fore have involved successful seedling establishment, it must follow that
the mutant strains date back to Pluvial times. Since the altitudinal
migration of aspen in Postpluvial times has been mainly upward, and
since the late-leafing strain dominates the — aspen elevations today,
the late-leafing form must be the parent ty

It seems obvious that a longer cherie: period (upwards of three
weeks) acquired by the mutant strains would give them distinct physi-
ological survival advantages over the parent form, particularly at higher
elevations where the frost-free period is short, provided, of course, that
they also possessed adaptive resistance to lower temperatures. The ability
of the early-leafing form to withstand frost in spring is apparent because
banks of snow are frequently present beneath the aspen canopy when
these trees come into leaf. Temperature records in the early-leafing experi-
mental colony at Mt. Timpanogos, with no snow on the ground in 1951,
showed night temperatures of 30° F. or below for a period of approximately

1954] COTTAM, PREVERNAL LEAFING OF ASPEN 247

nine hours duration each for June 1, 2 and 3. On the night of June 1,
1951, the minimal temperature of 26° F. prevailed for approximately
three hours. The aspen leaves in the early-leafing colony at this time were
about one half mature size, while the adjacent late-leafing trees were still
in the bud stage.

Whether or not this insensitivity to mild freezing temperature is merely
another expression of the same mutation that produced early leafing
would be difficult to determine, but certainly the prevernal appearance
of aspen leaves would be impossible without this adaptation. That the
same degree of frost resistance is not possessed by the late-leafing form
is suggested by its apparent inability to invade the upper limits of the
aspen belt, by its failure to produce leaves at mid-elevations until the
frost period is over and by its habit of dropping leaves earlier in the fall
than the early-leafing varieties growing in similar habitats.

At mid-elevations it is apparent that both strains of aspen successfully
established seedlings at the close of Pluvial times after these mountain
areas had been cleared of the receding ice. It is probable that at lower
elevations, high temperature and drought were more limiting as factors of
successful aspen establishment than a longer photosynthetic period. Also,
suitable sites for aspen were already occupied by the late-leafing parent
type. Assuming that neither strain had significant advantage at mid-
elevations, the preponderance of seed of the late-leafing strain would
account for the dominance of the late-leafing form at mid-elevations
today. Clonal reproduction over the centuries of Postpluvial time from
trees of these two strains as loci for vegetative migration has, in many
instances, brought about the mergence of these two contrasting leafing
forms into a single forest stand.

A critical test of this hypothesis awaited the experiment using radio-
active phosphorus to trace the clonal relationship of the two strains. It
was postulated that a solution of radioactive phosphorus applied to the
roots of an aspen of either strain, situated at the line of contact of the
diverse colonies, should, according to the hypothesis, pass freely through
the root connections to neighboring trees of its own strain, but never to
the trees of the opposite strain regardless of their proximity to the labeled
tree. The one experiment performed thus far is therefore disappointing
as evidence for or.against the hypothesis in that it suggests that mature
aspen trees in a colony sever their clonal connections.

SUMMARY

1. Almost any aspen forest in early spring in Utah and neighboring states
shows sharply discontinuous colonies of trees that attain full leaf two to
three weeks earlier than the major stand that surrounds them. The colonies
of prevernal aspen are usually small in comparison with the forest as a
whole, and in general they are most pronounced at mid-elevations of the
aspen belt.

2. Extensive data secured with recording thermographs show that one
segment of an aspen forest exhibiting prevernal leafing, is a response to

248 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

temperature controls, but that in general the early-leafing colonies of
aspen represent distinct genetic strains in which temperature is not a
factor.

3. As a rule the leafing habit of the various aspen strains is associated
with morphological characters by which they may be identified at all
seasons.

4. Saplings of early- and late-leafing strains of aspen transplanted to the
University of Utah campus at an elevation of 4,500 feet exhibited the
same time difference in leafing as their parent colonies at 7,800 feet.

5. An hypothesis proposed to explain the origin, present distribution and
nature of prevernal aspen colonies postulated the following:

(a) Sexual mutations of the late-leafing parent type occurred in

Pluvial times.

Migration to and establishment of these strains at higher elevations

followed the disappearance of mountain ice and snow packs.

(c) Early-leafing forms dominate the upper limits of the aspen belt
because of their longer photosynthetic period and their physiolog-
ical adaptations to lower temperatures.

(b

—

(

Qu.
—

Clonal reproduction over the centuries of Postglacial time from
seed-established trees as loci for vegetative migration has produced
the mergence of these contrasting leafing strains into single forest
stands.

UNIVERSITY OF UTAH,
SALT LAKE City, UTAH.

LITERATURE CITED

—

. ANTEvS, ERNST 1948. Climatic oo and pre-whiteman. The Great Basin.
Bull. lini. Utah 38 (20): 168-19

2. 1952. Cenozoic climates ms the Great Basin. Geologischen Rundschau
40 (1): 94-108.

3. 1952. Univ. Calif. Archaeol. Surv. Reports, No. 16, p. 23.

4. BAKER, F. S. 1921. Two Races of Aspen. Jour. Forestry 19 (4): 412-413.

EXPLANATION OF PLATES

PLATE I
Typical stands of early- and late-leafing aspens, mid-elevation (7800 feet).
t. Timpanogos, Utah. In the right mid-ground is seen the protective tent
covering for one of the recording thermographs used to compare atmospheric
temperature phenomena of the contiguous aspen strains.
PLATE II
Late-leafing aspens (foreground) have white, well-pruned boles in northern
Utah, while the early-leafing forms (background) are yellow-boled and poorly
pruned.

Jour. ARNOLD Ars. VoL. XXXV

sree a.
, iept

ees mee ee: pe,

PLATE I

NS

MounrtTAI

NAL LEAFING OF ASPEN IN UTAH

PREVER

PuaTE II

Jour. ARNoLD Ars. VoL. XXXV

“t

ee I RRR I 8: A

AINS

NAL LEAFING OF ASPEN IN UTAH MOUNTS

ER

,

PREV

1954] SAX, CONTROL OF TREE GROWTH 251

THE CONTROL OF TREE GROWTH BY PHLOEM BLOCKS
KARL SAX

With one plate

THE NATURE OF SAP MOVEMENT, nutrient transport and stem polarity
in plants has long been known. Thomas Andrew Knight reporting in the
Transactions of the Horticultural Society of London in 1822 observed
that ‘the vessels of plants are not equally well calculated to carry their
contents in opposite directions” and that ‘‘the true sap of trees is wholly
generated in the leaves, from which it descends through their bark to the
extremities of their roots, depositing in its course the matter which is
successively added to the tree.” Thus Knight recognized polarity in the
transport of elaborated nutrients down the phloem based upon his experi-
ments with inverted cuttings and the effect of girdling the bark of fruit
trees. He observed that “it had long been known to gardeners, that taking
off a portion of bark round the branch of a fruit-tree occasions the pro-
duction of much blossom on every part of that branch in the succeeding
season.”’ He concluded that the effect of dwarfing stocks in promoting
early fruiting and restricting tree growth was similar in nature to the
effect of girdling the bark and attributed both effects to the “obstruction
of the descending sap.”

Phloem blocks induced by killing the stem with steam were used by
Dixon (8) and others in their studies on the ascent of sap. The fact that
the sap passed up the xylem of the dead stem provided evidence for the
“mechanistic” rather than the ‘‘vitalistic’ theory of the ascent of sap.
More recently it has been shown that phloem blocks, induced by killing
a section of the stem with heat, prevent the downward movement of vita-
mins, organic carbohydrates and growth-regulators (Crafts 1951). Bonner
(4) girdled the stems of tomato plants with a jet of superheated steam
which killed all living tissue. In a few days he found an accumulation of
thiamin, pyridoxine, pantothenic acid, riboflavin, sucrose and nitrogen
above the girdle. Rabideau and Burr (11) killed sections of bean stems
with hot wax and used radioactive carbon to trace the flow of photosyn-
thetic products down the stem. Most of the elaborated organic material
accumulated at the dead segment. A phloem block in Phaseolus also
checked the movement of growth-regulators such as 2,4-D as shown by
Weintraub and Brown (15).

It is, however, unnecessary to kill a segment of the stem in order to
induce a phloem block. Christensen (5) found that irradiation of a stem
segment with X-rays was followed by a swelling of the stem above the
irradiated section, and after four or five weeks the swollen area developed
roots if the stems were kept moist. The minimum dosage required for
such a response was 1500 r for Xanthium. Similar results were obtained

252 JOURNAL OF THE ARNOLD ARBORETUM [voL. XXXV

with Nicotiana at 6000 r, Lycopersicum at 24,000 r, and Phaseolus at
16,000 r, but the minimum dosage required to induce a response was not
determined in these genera. The phloem block induced by irradiation
checked the flow of organic materials and auxin down the phloem without
killing the stem segment.

It is also possible to impose a phloem block without killing the stem by
inverting the scion or by inverting a ring of bark. This technique is based
upon the polarity of phloem transport. The botanical studies of stem
polarity date back to the work of Véctung and Sachs with their description
of ‘“root-pole” and ‘‘shoot-pole.”” The role of polarity in grafting was
probably known by horticulturists for hundreds of years. It was de-
scribed by the botanist Strasburger (14) as follows — “Unlike poles of a
plant may readily be induced to grow together, while like poles may only
be brought to do so with difficulty and then do not develop vigorously.”
This conclusion has been confirmed repeatedly. The inversion of the
rootstock has, however, been used to stimulate rooting at the base of the
scion in order to get varieties of apple trees on their own roots. Kerr (10)
found that such an inverted graft would survive long enough to permit
scion rooting above the graft union, due presumably to the accumulation
of auxin at the junction of the ‘‘root-poles” of stock and scion.

The induction of a phloem block by inverting a ring of bark was reported
in 1935, but it is probable that it was also used by the early horticulturists
of Europe. According to Roberts (12) the inverted rings checked the
growth of the tree, but without the deleterious effect resulting from girdling.
Apparently this work was not continued by Roberts.

It is also known that certain incompatible combinations of stock and
scion check tree growth as a result of a phloem block induced by an im-
perfect graft union (2). In some cases, however, the graft union appears
to be normal yet there is considerable overgrowth of the scion accom-
panied by dwarfed growth and precocious fruiting. In these cases per-
haps the stem of the scion can utilize the organic carbohydrates from its
leaves more effectively than can the stem of the rootstock.

There is some evidence that the dwarfing effect of certain Malling apple
rootstocks may be due to a retardation of phloem transport. Dr. F. R.
Tubbs, Director of the East Malling Research Station, writes that: ‘We
do not know of any dwarfing apple rootstock that does not induce the for-
mation of a bulge” (personal communication). This swelling of the stem
of the rootstock or the section of the interstock could be attributed to the
retardation of the downward flow of elaborated organic nutrients and
growth stimulants, as suggested by Knight, thus promoting more rapid
growth of the Malling rootstock or interstock stem. The fact that a long
dwarfing interstock is more effective than a short one, and that certain
rootstocks are more dwarfing if budded high on the stem than if budded
near the ground, would seem to support the above interpretation (13).

It is known that the dwarfing effects of certain rootstocks are due to
factors other than the blocking or retardation of phloem transport. Colby
(6) has suggested that the extreme dwarfing effect of the Malling IX

1954] SAX, CONTROL OF TREE GROWTH 453

apple rootstock may be due to early suberization of the young roots. There
is also evidence that growth can be suppressed, quite independently of the
nature of the graft union, by an interaction between the scion variety and
the root system of the stock (Sax, Proc. Am. Soc. Hort. Sci., in press).

During the past five years a study of the effect of inverting rings of
bark has provided some information on the mechanism of phloem trans-
port and has provided another method for dwarfing fruit and ornamental
trees. We began with the inversion of a single ring of bark about an inch
long on the stems of one- and two-year-old apple trees. The cuts through
the bark were made as parallel as possible, sometimes by using a double
bladed knife — in other cases by using a strip of metal tape as a guide.
The bark was removed, inverted and wrapped tightly with a rubber band,
until the bark had healed onto the stem — a period of a week or ten days.

The inverted bark made little or no growth, but there was a swelling of
the stem above the inverted bark and to a lesser extent at the upper edge
of the inverted section of bark. At the vertical seam of the inverted ring
there was regeneration of tissue and after several months this area had
made considerable growth, and after several years it had so dominated
the inverted area that the dwarfing effect was largely lost. Apparently
there is normal polarity of phloem transport in this regenerated area and
it grows rapidly.

In order to effect a more permanent phloem block it was necessary to
invert two rings of bark, one directly above the other, and orient the
vertical seams on opposite sides of the stem. There is some phloem
regeneration at the seams, but any downward transport through the
seam of the upper inverted ring is checked when it reaches the intact
edge of the lower inverted ring with its vertical seam on the opposite side.
A Cortland apple whip was treated in this way five years ago. It has made
almost no growth in trunk diameter and has increased in height only about
six inches during the past five years, but it has borne several fruits during
each of the past two years and appears to be healthy.

In 1953, a group of Baldwin apple trees, budded on a semi-dwarfing
rootstock, were used for a bark inversion test. A single ring of bark was
inverted on eight two-year-old trees, and above the vertical seam a square
of bark was inverted to prevent transport down the regenerated tissue.
In the eight controls a ring of bark was removed and replaced in the
normal position. In both the bark inversion series and in the controls
there was some early sucker growth below the rings, and the first growth
of suckers were removed. Subsequent sucker growth was limited almost
entirely to the trees with inverted bark. At the end of the growing season
the trunk diameter below the bark ring was measured, together with tree
height and total length of the branches developed below the inverted
bark rings. The results are shown in Table 1, and photographs of a
control and of an inverted bark tree are shown in Figures 1 and 2

The data show that the inversion of the ring of bark reduced the
growth of the trees both in trunk diameter and in height. It also promoted
the growth of buds below the phloem block, indicating that the auxins

254 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

which normally suppress bud development at the base of the trunk had not
passed through the inverted ring of bark in sufficient quantities to suppress
bud growth. Most of the trees with the inverted bark have flower buds
while no flower buds were observed on the control trees. The inverted
bark remains alive, so far as can be determined by its appearance, yet
growth is suppressed and earlier fruiting is induced.

TABLE 1.
EFFECT OF BARK INVERSION ON TREE GROWTH
Baldwin/524/sikkimensis.
Bark inverted June 1, 53. Measured Oct. 6, ’53.

8 Controls 8 Inverted
runk Height Sucker Trunk Height Sucker
Caliper cm. Feet Growth ft. Caliper cm. Feet Growth ft.
2.0 4.5 0 1.5 3.8 4.1
1.7 ee 0 1.1 Xe | 4.2
1.9 4.6 0) 1.4 3.2 4.5
2.1 4.9 0) 1.4 3.2 5.1
1.9 4.5 6) 1.4 3.1 4.9
1.9 4.0 6) 1.6 3.5 2.1
1.6 4.2 0.5 1.5 55 3.2
2.1 4.1 0) 1.5 3.6 4.7
Ave. 1.9 4.5 —Q,1 1.4 3.4 4.1

It should be possible to modify the bark inversion phloem block to
permit the desired amount of growth of the tree. A method developed
several years ago appears to meet this need. Baldwin two-year-old trees,
budded on Malus sikkimensis rootstocks, were used. Two bark inversions
were made with eight-inch strips of normal bark on opposite sides of the
trunk, and the inverted rings were separated by a short segment of normal
bark of half an inch to one inch in length. The downward movement
through the phloem was limited to the slender strip of normally oriented
bark. After passing down the first ring the flow must pass laterally before
it can pass down the normal strip of bark in the second ring. Although
lateral phloem transport is restricted some material does get through as
is evident from the growth shown in Figure 4. It is probable that a short
section of normal bark between the inverted rings would have a greater
restriction on phloem transport than a long one, so that the degree of
dwarfing could be regulated to some extent by varying the length of the
central segment between the inverted rings. Thus by adjusting the width
of the normal strips of bark in the inverted rings and by varying the
length of the normal central segment, it should be possible to obtain any
desired degree of dwarfing.

All of these treated trees flowered and several bore fruit the following

1954 | SAX, CONTROL OF TREE GROWTH 238

year. One of these trees, bearing abundant flower buds as it begins its
fourth season’s growth, is shown in Figure 3. The details of the double
bark inversion are shown in Figure 4. The growth of the normally polar-
ized strips of bark in the inverted rings may eventually result in a direct
line of phloem transport through the normal ring of bark between the
inverted segments, and the dwarfing effect will be reduced and finally lost.
In order to maintain a permanent dwarfing effect the inverted bark tech-
nique may have to be repeated, but when a tree reaches the desired size a
double inversion with only a very narrow ring of bark (or none at all)
between the inverted rings, should insure permanent dwarfing. Since the
bark inversion may have to be repeated, this technique for dwarfing trees
is not likely to be of much value to the commercial nurseryman. The
average back yard horticulturist should, however, have no trouble in using
this technique.

In a few cases we have put double upside-down adjacent rings on

and fruiting. These trees have been treated only for one growing season,
and at least several years must elapse before we can recommend such a
procedure.

The inactivation of the phloem by ionizing radiation is comparable in
several respects to the effects of inverting a ring of bark. The phloem
block is induced without killing the bark, swelling of the stem occurs
above the treated area and bud growth is stimulated below the phloem
block just as it is by inverting the bark. On July 27 the stems of a clonal
line of young poplar trees were irradiated with 2,500, 5,000 and 10,000
roentgen units of X-rays respectively. A lead plate with a 1.5 inch slit was
placed over the stems to limit the radiation to the short stem segment.
After several weeks the swelling above the irradiated areas became evi-
dent particularly at the higher doses. On Oct. 6th the six trees receiving
2500 or 5000 r were all alive, but at 10,000 r four of five treated trees
were dead above, at, and for some distance below the irradiated segment.
This higher dosage may have killed the tissue completely, followed by the
separation of the bark from the xylem as a result of the overgrowth of the
stem above the irradiated area, as shown in Figure 3. The inverted bark
produced a similar overgrowth, but without pulling the bark from the stem
in the treated segment. At the lower doses of X-rays there was evidence
of a phloem block with no death of the stem at the irradiated area.

It is evident that it is possible to produce a phloem block without
killing the stem by inverting a ring of bark or by non-lethal exposure of
a stem segment to X-rays. In both cases the growth of the treated area is
restricted or suppressed. The suppression of growth may be the cause of
the phloem block in these cases. According to Abbe and Crafts (1) —‘‘It
is characteristic of the sieve tubes of all plants, that, after a brief function-
ing period — consisting of from a few days in the case of protophloem
sieve tubes to a single season in most woody plants — the elements col-
lapse, and death occurs.”

256 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXXV

The X-ray induction of a phloem block without killing the tissue is
apparently due to nuclear injury which prevents continued cell division
without killing the cytoplasm. It is known that the cytoplasm is very
much more resistant to injury from X-rays than is the nucleus. Thus the
sieve tubes already formed could continue to function, but no new ones
would be formed to augment and replace those present at the time of
irradiation.

The bark inversion also seems to produce a phloem block by the inhibi-
tion of cell division. The suppression of growth in the inverted bark rings
is apparently due to the inability of nutrients and auxins to move freely
against the reversed polarity. There is usually a slight swelling of the
top of the inverted bark ring, indicating some diffusion of nutrients, but
there is little or no growth of the inverted segment as a whole. Thus the
inversion of the bark may check growth by the failure of nutrients to
move freely through the inverted phloem tissue.

The new phloem produced at the vertical seam in inverted bark segments
appears to be normally polarized since this tissue grows rapidly. Any
new phloem tissue produced within the inverted ring of bark should
eventually also be normally polarized, but the fact that growth does not
occur indicates that few or no sieve tubes are produced, presumably due
to the checking of nutrient flow through the inverted segment. The fact
that bark inversions made early in the growing season do not survive as
well as those made in June and August suggests that some active sieve
tubes are needed even though the inverted polarity checks the passage of
nutrients and auxins down the stem. Since a tree has lived for five years
with a double inverted ring of bark, some material must be transmitted
either through the inverted bark or through some other part of the stem.

SUMMARY

A phloem block can be induced without killing the stem tissues by ex-
posing stem segments to X-rays sufficient to suppress cell division or by
inverting a ring of bark. The phloem block induced by irradiation appears
to be due to the failure of renewal of phloem elements. The reversed
polarity of the inverted ring of bark also prevents renewal of phloem
elements, presumably by preventing the adequate movement of nutrients
and auxins into the inverted phloem cells. The inversion of rings of bark
may be modified to produce the degree of dwarfing desired in fruit and
ornamental trees.

Bussey INSTITUTION AND ARNOLD ARBORETUM,
HARVARD UNIVERSITY

1954] SAX, CONTROL OF TREE GROWTH

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LITERATURE CITED

Aspe, Lucy B. and Crarts, A. S. Phloem of white pine and other conifer-
ous species. Bot. Gaz. 100: 695-722. 1939.

Arctes, G. K. Stock-scion incompatibility in sin trees. Imp. Bur.
Fruit Production. East Malling, Kent, England. 1

BipputpH, O. and Marpte, J. Translocation of oer in the
phloem of the cotton plant. Amer. Jour. Bot. 31: 65-70. 1944.

BoNnNER, JAMES. Accumulation of various substances in girdled stem of
tomato plants. Amer. Jour. Bot. 31: 551-555. 1944.

CHRISTENSEN, Eric. Root production in plants following localized stem
irradiation. Science 119: 127-128. 1954.

Cotsy, H. L. Stock-scion chemistry and the fruiting relationships in
ae trees. Plant Physiol. 10: 483-98. 1935.

Crarts, A. S. Movement of assimilates, viruses, growth regulators, and
chemical indicators in plants. Bot. Rev. 17: 203-284. 1951.

Dixon, H. H. Transpiration and the ascent of sap in plants. Macmillan
& Co. Ltd. London. 1914.

Esau, KATHERINE. Development and structure of phloem tissue. Bot. Rev.
16: 67-114.

Kerr, W. L. A simple method of obtaining fruit trees on their own roots.
Proc. Amer. Soc. Hort. Sci. 33: 355-357. 1935

_ Rapiweau, G. S. and Burr, G. O. The use of C!8 isotope as a tracer for

transport studies in plants. Amer. Jour. Bot. 32: 349-356. 1945.

_ Roserts, R. H. A further trial of ring grafting to produce stock effects.
35

Proc. Amer. Soc. Hort. Sci. 33: 358-359.

. Roperts, R. H. Theoretical aspects of eanage Bot. Rev. 15: 423-463.
1949.

STRASBURGER, E. et al. A Textbook of Botany. Macmillan and Co. Ltd.
London. 1898

Wewaghes, R. L. and Brown, J. W. Translocation of exogenous growth-
regulators in the bean seedling. Plant Physiol. 25: 140-149. 1950.

258 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

DESCRIPTION OF FIGURES

Fic. 1. Baldwin tree, 2 years old, on semi-dwarfing rootstock. A ring of bark
was removed and replaced in the normal position on June 1, 1953. This control
tree has made considerable growth in a single season

Fic. 2. Baldwin tree, 2 years old, on semi-dwarfing rootstock. A ring of bark
was inverted June 1, 1953. Note dwarfing effect in a single season, and the pro-
duction of flower buds. The suckering below the inverted ring of bark indicates
the checking of auxin flow.

Fic. 3. Baldwin on M. sikkimensis, 3 years old, dwarfed by double inverted
ring with a slender strip of normal bark on the opposite side of each of the two
inverted rings. A short section of normal bark was left between the two inverted
rings. This tree bore Stauth in Si third growing season and bears many flower
buds as it begins its fourth y

Fic Details of dea ‘bok inversion show how phloem transport is
checked. “The swelling of the upper end of the inverted ring of bark suggests
some nutrients and auxins may flow into the inverted bark by diffusion, but
active ni transport is blocked by the reversed polarity of the inverted
siege tis

Fic. vi ‘Sie stem with inverted ring of bark, showing swelling of the
stem phe the phloem block imposed by reversed polarity of the phloem.

Fic. 6. A poplar stem exposed to 10,000 r of X-rays. Note the similar
swelling of the stem above, and the suckering below, of the irradiated section of
s The X-rays impose a phloem block by preventing cell division and the
renewal of phloem cells. In this case the bark was killed and is pulled away
from the wood by the expansion of the stem above the irradiated area. but it is
possible to impose a phloem block with X-rays without killing the tissue.

Photographs by Heman Howard.

Jour. ARNoLp Ars. VoL. XXXV PLATE I

CONTROL OF TREE GROWTH BY PHLOEM BLOCKS

260 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

CRYPTOGAMS OF THE 1948 ARCHBOLD CAPE YORK
(QUEENSLAND) EXPEDITION

P. Brissy

MUSCI *
LEMBOPHYLLACEAE

Camptochaete brisbanica (C. M.) Broth., Proc. Linn. Soc. N.S. W. 43:
561.

QUEENSLAND: Mt. Finnegan, Brass 20058, 20155, Sept. 1948, alt. 850 m
on branches in rain-forest undergrowth; Mossman River las Brass 18173,
Mar. 1948, alt. 150 m., plentiful on eranite rocks in rain-fores

Not compared with the type, but agreeing well with material so named
in the National Herbarium of Victoria.
HOOKERIACEAE
Eriopus sp. (aff. E. apiculatus |Hk. f. & W.| Mitt.).

QUEENSLAND: Mt. Finnegan, Brass 20097, Sept. 1948, alt. 1040 m

., trunk
of a tree in high mountain forest.

Very close to and probably identical with E. apiculatus (Hk. f. & W.)
Mitt.

HEPATICAE
PTILIDIACEAE
Mastigophora Nees
Mastigophora diclados (Endl.) Steph., Spec. Hep. 4: 38. 1909.
QUEENSLAND: Leo Creek, Upper Nesbit River, Brass 19957, Aug. 1948,
alt. 420 m., cushioned on trees on bank of creek in rain-forest.
LEPIDOZIACEAE
Bazzania S. F. Gray
Bazzania adnexa (L. & L.) Mont., Voy. au Pole Sud. 243. 1842-45.
QUEENSLAND: Lamb’s Head, Lamb Range, Brass 18227, Mar. 1948, alt.

1000-1100 m., sunny rocks in rain-forest; Mt. Bellenden-Ker, summit of south
peak, Brass 18288, Apr. 1948, alt. 1550 m., on trees of low forest

Bazzania Novae-Zelandiae (Mitt.) Kuntze, Gen. Plant. 832. 1901.

* Determined by J. H. Willis.

1954] BIBBY, CRYPTOGAMS OF CAPE YORK 261
QUEENSLAND: Mt. Finnegan, Brass 20144, Sept. 1948, alt. 1140 m., on
bark of a tree in high mountain scrub.
Lepidozia Dumort.
Lepidozia capilligera (Schwaegr.) Lindenb., Syn. Hep. 204. 1844.
QUEENSLAND: Iron Range, Brass 19051, June 1948, alt.°20 m.. on a
decaying log in floodplain in rain-forest.
HARPANTHACEAE
Chiloscyphus Corda
Chiloscyphus argutus (R. B. & N.) Nees, Syn. Hep. 183. 1845.
QUEENSLAND: Iron Range, Brass 19052, June 1948, alt. 20 m., on decaying
logs in floodplain rain-forest.
JUNGERMANNIACEAE
Anastrophyllum (Spruce) Steph.
Anastrophyllum piligerum (Nees) Spruce, Jour. Bot. 14: 33. 1876.
QUEENSLAND: Mt. Bellenden-Ker, summit of south peak, Brass 18286,
Apr. 1948, alt. 1550 m., abundant on trees of low forest.
Chandonanthus Mitt.
Chandonanthus hirtellus (Web.) Steph., Spec. Hep. 3: 643. 1909.
QUEENSLAND: Mt. Bellenden-Ker, summit of south peak, Brass 18283,
Apr. 1948, alt. 1550 m., on a tree in low forest.
PLAGIOCHILACEAE
Plagiochila Dumort.
Plagiochila abietine * (Nees.) Lindenb., Mon. Hep. Gen. Plag. 134.
1844.

QUEENSLAND: Mt. Finnegan, Brass 20141, Sept. 1948, alt. 1140 m., shrub-
beries of the summit.

This constitutes the first record of this species on the Australian main-
land.
Plagiochilon Hattori
Plagiochilon oppositus (R. B. & N.) Hattori, Biosphaera 1: 7. 1947.
Plagiochila opposita (R. B. & N.) Dum., Rec. d’obs. 15. 1835.

QUEENSLAND: Mt. Finnegan, growing with Sticta, Brass 20095, Sept.
1948, alt. 1100 m., on mossy rocks in high mountain.

Not previously reported on the Australian mainland.

* Determined by Th. Herzog, Jena.

262 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV
SCHISTOCHILACEAE
Schistochila Dumort.
Schistochila cristata Steph., Hedw. 28: 274. 1889.
QUEENSLAND: Mt. Finnegan, Brass 20089, ia 1948, alt. 1100 m., very
abundant on rocks (granite) in high mountain fore
RADULACEAE
Radula Dumort.
Radula acutiloba Steph., Hedw. 28: 271. 1889.

QUEENSLAND: Leo Creek, Upper Nesbit River, Brass 19954, Aug. 1948,
alt. 420 m., on dead twigs in rain-forest undergrowth.

Radula buccinifera (Hk. f & Tayl.) Tayl., Syn. Hep. 261. 1845.
QUEENSLAND: Mt. Finnegan, Brass 20094, Sept. 1948, alt. 1100 m., on
leaves and branchlets of undergrowth in high mountain forest.
Radula reflexa Mont., Ann. Sc. Nat. 19: 255. 1843.
QUEENSLAND: Mossman River Gorge, Brass 18177, Mar. 1948, alt. 150 m.,
moist shady granite rocks in rain-forest.
PLEUROZIACEAE
Pleurozia Dumort.

Pleurozia articulata (Linb.) Schiffn., Engl. Pflanzenfam. I. 3: 115. 1895,
QUEENSLAND: Mt. Finnegan, Brass 20142, Sept. 1948, alt. 1140 m.,
living twigs in shrubberies of summit; Mt. Bellenden- Ker, summit of er
peak, Brass 18287, Apr. 1948, alt. 1550 m., hanging from dead twigs in low

forest.
FRULLANIACEAE
Frullania Raddi
Frullania Johnsonii Steph., Hedw. 33: 163. 1894.
QUEENSLAND: Mt. Bellenden- i summit of south peak, Brass 18289,
Apr. 1948, alt. 1550 m., on dead twig
Frullania sp.
QUEENSLAND: Summit of Mt. Tozer, Tozer Range, sche 19497, July
1948, alt. 540 m., living branches of Casuarina in scrub of summ
LEJEUNEACEAE
Drepanolejeunea (Spruce) Schiffn.
Drepanolejeunea Micholitzii Steph. var. genuina Herz. Ann. Bryol.
7: 80. 1934.
QUEENSLAND: Mt. Finnegan, Brass 20098, Sept. 1948, alt. 1040 m., on
leaves of undergrowth in high mountain forest

1954] BIBBY, CRYPTOGAMS OF CAPE YORK 263

Drepanolejeunea obliqua Steph., Hedw. 35: 82. 1896.
QUEENSLAND: Mt. Finnegan, associated with Brass 20144, Sept. 1948,
alt. 1140 m., on bark of a tree in high mountain scrub.

Drepanolejeunea tenuis (Nees) Schiffn., Consp. Hep. Arch. Ind. 280.

QUEENSLAND: Lamb’s Head, Lamb Range, associated with Brass 18227,
Mar. 1948, alt. 1000-1100 m., sunny rocks in rain-forest

Physocolea Spruce
Physocolea trichomanis (Gott.) Steph., Spec. Hep. 5: 912, 1916.
Cololejeunea trichomanis (Gott.) Steph., Hedw. 28: 168. 1889.

QUEENSLAND: Mt. Finnegan, west slopes, Brass 20054, Sept. 1948, alt.
850 m., on leaves of an undergrowth tree in rain-forest.

LICHENES
SPHAEROPHORACEAE
Sphaerophorus Pers.
Sphaerophorus compressus Ach., Meth. Lich. 135. 1803.

QUEENSLAND: Lamb’s Head, Lamb Range, Brass 18234, Mar. 1948, alt.
950-1000 m., sunny rocks in rain-forest.

COENOGONIACEAE
Coenogonium Ehrenh.
Coenogonium implexum Nyl., Ann. Sc. Nat. 16: 92. 1861.

QUEENSLAND: Speewah, Upper Clohesy River, Brass 18209, Mar. 1948,
alt. 450 m., trunk of a tree in rain-forest.

CLADONIACEAE
Cladonia Hill.
Cladonia aggregata (Sw.) Ach., Vet. Acad. Nya Handl. 16: 68. 1795.

QUEENSLAND: Lamb’s Head, Lamb Range, Brass 18232, Mar. 1948, alt.
950-1000 m., sunny rocks in rain-forest; summit of Mt. Tozer, Tozer Range,
Brass 19495, July 1948, alt. 540 m., in shelter of boulders on exposed granite
rock faces; Mt. Bellenden-Ker, summit of south peak, Brass 18290, April 1948,
alt. 1550 m., on peaty ground of a small clearing.

Cladonia verticillata Hoffm. var. cervicornis (Ach.) Flot., Linnaea 22:
380. 1849.

Seagate Lamb’s Head, Lamb Range, Brass 18233, Mar. 1948, alt.
950-1000 m., sunny rocks in rain-forest.

264 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXXV

PANNARIACEAE

Pannaria Del.
Pannaria myrioloba Muell.-Arg., Bull. d’Herb. Boiss. 4: 92. 1896.

QUEENSLAND: Mt. Bellenden-Ker, summit of south peak, Brass 18291,

April 1948, alt. 1550 m., on bark of living tree in low forest; Mt. Bellenden-Ker,
summit of south peak, Brass 18292, April 1948, alt. 1550 m., on dead twigs in
low forest.
Pannaria sublurida Nyl., Ann. Sc. Nat. 11: 256. 1859.

QUEENSLAND: Mt. Bellenden-Ker, summit hus south peak, Brass 18293,
April 1948, alt. 1550 m., growing on bryophyte

COLLEMACEAE
Leptogium S. F. Gray

Leptogium phyllocarpum (Pers.) Mont. var. isidiosum Nyl., Syn.
Meth. 1: 130. 1858.

QUEENSLAND: Mossman River Gorge, Brass 18168, Mar. 1948, alt.
150 m., on rocks on edge of a rain-forest stream.

STICTACEAE
Sticta Schreb.
Sticta aurata Ach., Meth. Lich. 277. 1803.
QUEENSLAND: Mt. Finnegan, Brass 20359, Sept. 1948, alt. 910 m., on
living bark of an exposed branch in high mountain forest.
Sticta Sayeri Muell.-Arg., Flora 71: 23. 1888.
QUEENSLAND: Lamb’s Head, Lamb Range, Brass 18235, Mar. 1948, alt.
950-1000 m., sunny rocks in rain-forest.
Sticta sulphurea Schaer., Moritz. Verz. 127. 1846.
QUEENSLAND: Mt. Fin ce an slopes, Brass 20059, Sept. 1948, alt.
850 m., on a rotting log in rain-fore
Sticta sp.

QUEENSLAND: Mt. Finnegan, Brass 20095, Sept. 1948, alt. 1100 m.,,
abundant on mossy rocks in high mountain forest.

PARMELIACEAE
Parmelia Ach.
Parmelia latissima Fee forma sorediata Nyl., Syn. Meth. 1: 380. 1858.

QUEENSLAND: Tozer Range, north end, Brass 19369, June 1948, alt. 300
m., exposed granite rocks on summit.

BIBBY, CRYPTOGAMS OF CAPE YORK

Dictyonema irpicinum Mont., Ann. Sc. Nat. 10: 119

HYMENOLICHENES
THELEPHORACEAE

Dictyonema Agardh

420 m., on a mossy log in rain-fores

1848

265

Pra ramimnte os Leo Creek, oan = River, Brass 19958, Aug. 1948,

NATIONAL HERBARIUM OF VICTORIA

SouTH Y.
AUSTRALIA.

ARRA, VICTORI

266 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

ADDITIONAL NOTE ON NOTHOFAGUS
C. G. G. J. VAN STEENIS

With one text-figure

THE PAPER on New Caledonian Nothofagus by Dr. M. Baumann-
Bodenheim alluded to in my revision! was preceded by a preliminary
note.” Also after a personal interview at Leyden, May 12, 1953, when
we again interchanged data, Baumann-Bodenheim still cherished the idea
that the New Caledonian representatives of Nothofagus, as well as some
of those from New Guinea, belong to a distinct genus, Trisyngyne Baill.,
on the single evidence that these species should have “branched caducous
é inflorescences,’ though he later admitted these ‘inflorescences’ were
short-twigs. The confusion arises through the fact that the leaf-blades at
the base of the flush are often reduced and both the ¢ triads and cupules
appear clasped between the stipules. Such a lateral twig in flush may
give the superficial impression of an “inflorescence.” Morphologically it
remains a twig with lateral inflorescences. Whether these short twigs fail
to develop further into persistent branches is entirely irrelevant. In the
New Guinean species the flowering twigs are usually persistent and not
caducous. For these reasons I maintain that the five perfectly good species
described by Baumann-Bodenheim belong to Nothofagus sect. Calusparas-
sus subsect. Bipartitae series Triflorae Steen.

The following transfers are therefore necessary:

Nothofagus
Trisyngyne Baill. Adansonia 11: 136. 1873, syn. nov.
Nothofagus codonandra ( Baill.) comb. nov.
Trisyngyne codonxandra Baill. Adans. l.c.; Baumann-Bodenheim, Bull. Mus.
Hist. Nat. Paris II, 25: 420. 1953.
Nothofagus baumanniae (Baum.-Bod.) comb. nov.
Trisyngyne baumanniae Baum.-Bod. l.c. 420.
Nothofagus balansae ( Baill.) comb. nov.
Trisyngyne balansae Baill. l.c. 137; Baum.-Bod. l.c. 420.
Nothofagus discoidea (Baum.-Bod.) comb. nov.
Trisyngyne discoidea Baum.-Bod. l.c. 420.
Nothofagus aequilateralis (Baum.-Bod.) comb. nov.
Trisyngyne aequilateralis Baum.-Bod. l.c. 421.

‘Van Steenis, Jour. Arnold Arb. 34: 308. 1953.
*Baumann-Bodenheim, Bull. Mus. Hist. Nat. Paris II, 25: 419-421. 1953.

1954] VAN STEENIS, NOTE ON NOTHOFAGUS 267

Mainly through the generous assistance of Dr. F. Kausel, Santiago de
Chile, I obtained some valuable additional material which enabled me to
complete my former paper! with some interesting illustrations of the

Ficure 1. Nothofagus glauca (Philippi) Krasser. a. Cupule with two lateral
nuts, of the central nut only the scar of its attachment is visible. b. A single nut.

* 4 (KauseL 2303).— Nothofagus resinosa Steen. f. Mature nut with re-
duced cupular valve, X 4 (WomersLEY N.G.F. 5134).

cupules of the inadequately known NV. alessandri Espinosa (fig. 1 e) and
N. glauca (Phil.) Krasser (fig. 4 ad).

Another interesting figure is that of the mature nut of NV. resinosa Steen.
from New Guinea, for which I have to thank Mr. J. S. Womersley, Lae.
This shows that the reduced elamellar cupule of this species is not ap-
preciably enlarged in fruit.

1 Van Steenis, Jour. Arnold Arb. 34: 308. 1953.

FLoRA MALESIANA FOUNDATION,
LEYDEN, THE NETHERLANDS.

268 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

NEW ZEALAND CONIFERS
VIVIENNE DELLOW CASSIE

With one plate

ONE OF THE MOST DISTINCTIVE ELEMENTS in the New Zealand flora
is that formed by the native conifers, which are to be found in almost
every scrub and forest community from North Cape to Stewart Island.
They are no longer to be found, however, on the outlying islands. There
are twenty species, all endemic, distributed among three families in five
genera. In size they range from the pygmy pine (Dacrydium laxifolium),
the world’s smallest conifer, which may bear cones at a height of eight
centimeters (cf. Cheeseman 1925), to giant kauris and podocarps well
above thirty metres. A height of nearly sixty metres has been reported
for kahihatea (Podocarpus dacrydioides — Entrican and Reid. 1949),

A detailed discussion of the taxonomic features of each species would
be superfluous here, since a number of adequate descriptions are already
available. These are located in the works of Kirk (1889), Cheeseman
(1925), Allan (1929), Cockayne and Turner (1939), Laing and Black-
well (1940), and Dallimore and Jackson (1948). Instead, the relevant
literature will be briefly reviewed, although this account does not pretend
in any way to be an exhaustive one.

An anatomical basis for taxonomic studies has been adopted by several
workers. Seward and Ford (1906) and Eames (1913) have contributed
to the knowledge of the Araucariaceae in general and of Agathis australis
in particular; Eames from a morphological standpoint, the former authors
making a broader phylogenetic approach. An early investigation of leaf
anatomy of New Zealand conifers was made by Griffin (1907). More
recently, a detailed survey of leaf anatomy in Podocarpus has been
attempted by Buchholz and Gray (1948), and Gray and Buchholz (1951).
These authors place the South Pacific species P. Spicatus and P. ferrugineus
in a new subsection (Euprumnopitys) of the section Stachycarpus. The
new subsection, Euprumnopitys, is distinguished by the absence of
idioblasts (i.e. sclerids) in the mesophyll of leaves. Except for Podocarpus
dacrydioides in the section Dacrycarpus, the remaining New Zealand
species fall into subsection D of Eupodocarpus, based on the absence of
accessory transfusion tissue. Geographic distribution of Podocarpus has
been studied by Foweraker (1934), and later by Buchholz and Gray
(1948). The latter regard the Wegener theory of continental drift as the
only feasible solution to distribution problems in this genus. There is
some evidence from leaf anatomy that migration between Australia-New
Zealand and South-Central America may have taken place in both direc-
tions (Buchholz and Gray, op. cit., p. 61). Eupodocar pus is regarded as the
most advanced section because of the hypostomatic leaves (stomata con-

1954] CASSIE, NEW ZEALAND CONIFERS 269

fined to the under surface), and the usually prominent accessory trans-
fusion tissue. This feature is lacking, however, in subsection D, which
contains the New Zealand species. In the section Stachycarpus, Podocarpus
spicatus has the most primitive known arrangement of female cones.

With a view to practical application, Orman and Reid (1941, 1946)
have investigated wood anatomy in the genus Dacrydium. These workers
have constructed a key for the native Dacrydium species, using diagnostic
features of wood structure. An informative series of bulletins has been
published by the New Zealand Forest Service (Ward and Reid, 1949,
Entrican and Reid, 1949) on properties and uses of six of the major
timber-producing trees in the country (Dacrydium cupressinum, Podocar-
pus ferrugineus, P. spicatus, P. totara, and Agathis australis).

Not the least interesting feature of the New Zealand conifers is the
prevalence of distinct juvenile leaf forms (Cockayne, 1932). These occur
in all species save a few of the podocarps (P. totara, P. hallti, P. nivalis
and P. acutifolius). Epharmony, too, may cause plants of the same species
to have a very dissimilar appearance when grown under different habitat
conditions. For example, Podocarpus nivalis, a medium-sized bushy shrub
in shade and shelter, is reduced to a prostrate shrub with much smaller
leaves in direct sunlight (Cockayne, op. cit.).

Hybrids, although not as common as in angiosperm genera like Hebe
and Coprosma, are known or suspected between the following species: *

Podocarpus acutifolius xX nivalis Dacrydium bidwillii * laxifolium

Podocarpus hallii X nivalis Dacrydium bidwilli > biforme?

Podocarpus hallii < totara (=P. Dacrydium biforme X laxifolium?
loderi Cockn.) Phyllocladus glaucus X_ trichomanot-

Podocarpus hallii X acutifolius? des?

Podocarpus ferrugineus > totara? Libocedrus bidwillii * plumosa?

Podocarpus spicatus X totara?

Ecological studies are more restricted in number, centering mainly
about the kauri (Agathis australis). Cranwell and Moore (1936) drew
attention to modified growth forms of kauris growing above a height of
660 metres on Te Moehau (Coromandel Peninsula), a refuge of both alpine
and subtropical remnants. Further autecological notes have been made by
McKinnon (1937, 1940-41), McKinnon and Dumbleton (1935), Harrison-
Smith (1938), and Foley (1950). Recently Mirams (1948, 1951, unpubl.)
has analysed in detail the environmental factors responsible for growth
and regeneration of the kauri. Root nodules of New Zealand conifers were
investigated by Yeats (1924).

n important contribution is that of Cranwell (1940), who has de-
scribed and figured pollen grains of all the New Zealand species, as well
as formulating distinctive generic and specific characters according to the
peculiarities of each type of pollen grain. Her work indicates that
Dacrydium bidwillii and probably also D. biforme and D. kirkii are rightly

* See Cockayne 1932, Cockayne and Allan 1934, Cockayne and Turner 1939.

270 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

placed in a group apart from Podocarpus, where Sahni and Mitra (1927)
would put them, on the basis of the structure of female cones alone.
Cranwell (1938) had previously attempted a partial solution of post-
glacial vegetation problems through an analysis of pollen from South
Island peat beds. Three main periods are recognised:

1. A grassland period correlated with quite uniformly harsh climatic
conditions;

2. A warmer and wetter podocarp period:

A mosaic of grassland and beech (Nothofagus) forest resulting from

local climatic differentiation.

The role played by conifers in New Zealand’s past geological time has
been clarified still further through the work of Harris, Fergusson and
Couper (1951). It appears that a number of extinct species, including at
least seven podocarps, dominated forest communities between early
Cretaceous and Upper Eocene times. Of the present-day species, Podocar-
pus dacrydioides dates from the Oligocene, and Phyllocladus is known
even earlier from Upper Cretaceous beds.

CLASSIFICATION
The following is an outline classification of the New Zealand species.

A revised key to the species and genera occurring in this country has been
published previously (Hay and Dellow, 1952).

Araucariaceae
Agathis Salisb.

Agathis australis Salisb. Kauri

Cupressaceae
Libocedrus Endl.
Libocedrus plumosa (Don) Sargent ( = L. doniana Endl.) Kawaka
Libocedrus bidwillii Hook. f. Kaikawaka, mountain cedar

Podocarpaceae

Podocarpus L’Hérit.
Section: Stachycarpus
Subsection: Euprumnopitys

Podocarpus spicatus R. Br. Matai, black-pine

Podocarpus ferrugineus D. Don. Miro, brown-pine
Section: Dacrycarpus

Podocarpus dacrydioides A. Rich. (Fic. 1). Kahikatea, white-pine
Section: Eupodocarpus
Subsection: D

Podocarpus totara D. Don. Totara

1954] CASSIE, NEW ZEALAND CONIFERS rap

Podocarpus hallii T. Kirk. Hall’s totara, thin-bark totara
Podocarpus acutifolius 'T. Kirk
Podocarpus nivalis Hook. Alpine totara

Dacrydium Soland.

Dacrydium cupressinum Soland. Rimu, red-pine
Dacrydium kirkii F. Muell. Monoao

Dacrydium biforme (Hook.) Pilger. Pink-pine
Dacrydium bidwillit Hook. f. Bog-pine, mountain pine
Dacrydium colensot Hook. Silver-pine

Dacrydium intermedium T. Kirk. Yellow silver-pine
Dacrydium laxifolium Hook. f. Pygmy pine

Phyllocladus L. C. Rich.

Phyllocladus trichomanoides Don. Tanekaha
Phyllocladus glaucus Carr. Toatoa
Phyllocladus alpinus Hook. f. Mountain toatoa

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Wellington, N. Z., 188 pp., figs. 1-28.

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CHEESEMAN, T. F. 1925. Manual of the New Zealand Flora. 2nd. ed., 1163 pp.
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CocKAyYNE, L. 1932. Polymorphy in New Zealand conifers and its relation to
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Cockayne, L., and H. H. ALian, 1934. An sari list of groups of wild
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CocKAYNE, L., and E. P. TurNER, 1939. The trees a e Zealand, 176 pp., figs.
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CRANWELL, L. M. 1938. Fossil pollens. N. Z. Jour. Sci. & Tech. 19: 628-645.
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Hay, J. A., - U. V. DELLow, 1952. New Zealand Conifers. Tuatara 4: 108-
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293-295.

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native trees. N. Z. Jour. For. 3: 224-225.

MrraMs, R. V. 1948. Some observations on the regeneration of Agathis oe
Salisb. (Kauri). M.Sc. thesis unpubl. in Auckland Univ. Coll. libra

Mrirams, R. V. 1951. A study of some of the factors concerned in ie as
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Auckland Univ. Coll. library.

OrMAN, H.R., and J. S. Rew, 1940-41. Dacrydium. N. Z. Jour. For. 4: 366-367.

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species of Dacrydium. Ann. Bot. 41: 75-89, pl. VII, figs. 1-4.

SewaArp, A. C., and M. F. Forp, 1906. Araucariae, recent and extinct. Trans.
Roy. Soc. London, Ser. B, 198: 305-411, pls. 23 & 24, figs. 1-28.

Warp, W. C., and J. S. Rem, 1949. The properties and uses of Rimu. N. Z.
Forest Serv. Inform., Ser. No. 2. Wellington

Yeats, J. S. 1924. Root nodules of New Zealand pines. N. Z. Jour. Sci. &
Tech. 7: 121-124.

WELLINGTON, NEW ZEALAND.

PLATE I
Podocarpus dacrydioides A. Rich. growing in beech (Nothofagus) forest about
620 metres above sea level, near Lake Waikaremoana, New Zealand.

Jour. ARNOLD ARB. VoL. XXXV PLATE I

PODOCARPUS DACRYDIOIDES A. RICH.

JOURNAL

OF THE

ARNOLD ARBORETUM

VoL. XXXV OCTOBER 1954 NUMBER 4

A MONOGRAPH OF THE GENUS PHILADELPHUS
SHIU-YING HU

With six plates

INTRODUCTION

Philadelphus is geographically one of the most widely distributed
genera in nature, ecologically one of the most varied groups in habitat,
and horticulturally one of the most popular shrubs in our gardens. Its
slightly discontinuous distribution forms a belt girdling the temperate
zone of the northern hemisphere, extending to the higher altitudes in the
tropics in North America. The early inhabitants of Asia Minor, Eastern
Asia and Central America, centers where different culture patterns origi-
nated, have independently discovered its merits and have brought under
cultivation many species for their showy flowers, their enticing fragrance,
or for their medicinal value. In ancient times the Parthians employed the
flowers in the preparation of hair perfume, and the advanced people of
Mexico, and apparently those of Central America, used the flowers for
making garlands, and the leaves for medicine. By the early part of the
eleventh century, the Chinese garden forms had developed to such a
degree of excellence that they were presented by the people of Szechuan
as special tributes to the emperor in Peking. Due to their hardiness, their
ability to tolerate a wide range of environmental conditions, their ease
of propagation, and their flowering late in the spring season, the species of
Philadelphus have become garden favorites, especially in the northern
hemisphere.

By the latter part of the nineteenth century, forms of both Asiatic and
American origin were established in various European gardens. Much
was written about them, and very many binomials based on these exotic
forms were published. Sometimes several or even up to a dozen different
names were assigned to specimens taken from plants of the same origin.
Some of the individual authors had only limited material for purposes
of comparison, and others lacked specimens from the type localities. Thus
it was inevitable that specific names were multiplied and much confusion
resulted. In addition, clerical errors in recording names and in labeling
living plants and preserved material provided further complication. In

276 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

some cases proposed new species were accredited to regions remote from
their actual place of origin. Naturally different names came into use for
species which not even a specialist could distinguish, and at the same time
a single name may have been applied to very different species. Both
botanists and horticulturists have complained of this nomenclatural con-
fusion, yet since 1896 there has been no monographic work for the clari-
fication of the entire group. Only scattered papers treating selected
species or isolated groups from certain areas have been published. Ac-
tually, the lack of uniformity in selecting diagnostic characters by different
authors has created further confusion. Thus it is that a practical scheme
for the classification of the numerous species in the light of modern
taxonomic knowledge and under the regulations of the accepted code of
nomenclature is an urgent need.

Unaware of the difficulties in identifying species of this genus, I tried
to name some unstudied Chinese material in the Arnold Arboretum
herbarium. To my surprise, I was unable to find a workable key which
could enable me to distinguish the Asiatic forms. After studying all the
original descriptions I found myself even more confused for most of them
are so meagre and the characters indicated are so generalized that one
description often applies to any one of a number of obviously different
species. I ttirned to more experienced taxonomists and horticulturists for
advice and was warned that the species of Philadelphus have long been
a botanical problem, and it would be wise for me to leave them alone.
Though inexperienced in the group and perplexed by the confusion, I had
the conviction that the man-made disturbances in the classification of
the group could be solved when a real knowledge of the plants themselves
had been acquired. I thus commenced an intensive study of the genus
involving both living and herbarium material. My hope was that by
careful examination of a large number of herbarium specimens collected
from various centers of the natural range of the species supplemented
with firsthand observations on the living plants cultivated in the Arnold
Arboretum, the general pattern of the evolution of the genus might be
discovered, and a natural order of relationship of the diversified taxa
within the genus might be outlined. Besides the material deposited in
the herbarium of the Arnold Arboretum (A) and the Gray Herbarium
(G) of Harvard University, large loans were obtained from the Bailey
Hortorium of Cornell University (BH), the Chicago Natural History
Museum (F), the Missouri Botanical Garden (MO), the New York
Botanical Garden (NY), the University of Tennessee (TENN), and the
United States National Herbarium (US). In September 1952, after the
annual meeting of the American Society of Plant Taxonomists which
took place at Cornell University, I examined the Philadelphus collection
of the Wiegand Herbarium of that University (CU) and incorporated
the notes in my manuscript. With these specimens, vegetative as well as

studied. Variations in each character were plotted with due allowance

1954] HU, THE GENUS PHILADELPHUS 277

for changes that may be caused by the age and vitality of the plant as
observed in cultivated forms. The more constant characters were distin-
guished from the variable ones, and thus what I believe to be the funda-
mental morphological characters essential to proper specific diagnoses
were ascertained. At the same time due attention was also given to
those characters which might be used to advantage in defining categories
above or below the specific level. By using these categories as standards
of differentiation, various components of the geographically distinct groups
were again investigated. Taxa of the same level were placed in what I
assume to be their proper status, and thus a simple scheme of classifica-
tion expressing the phylogenetic relationship of different groups was
constructed on the basis of morphological characters and geographic
distributions. It is hoped that this scheme will not only illustrate certain
principles in the evolution and distribution of the genus, but also be of
some service to horticulturists and to botanists who are concerned with
the identification of their Philadelphus material, and to cytogeneticists
who are interested in the chromosome numbers and behaviors of the various
forms of this genus.

In this study the determinations of the taxa are made by comparing
the specimens with the types as far as they are available. Since over
40% of the binomials pertaining to this genus were based on cultivated
forms with no type ever designated, and sometimes with no actual speci-
mens preserved, material annotated by their authors or specimens pre-
pared from plants cultivated in the gardens indicated in the original
publication are used to substitute for the types. In cases like P. inodorus
Linn. where the taxon was based entirely on a previously published plate
and a pre-Linnaean description, specimens from the type locality which
best matched the illustration and agreed with the description were selected
for identification purposes.

My thanks are due not only to the curators of the herbaria mentioned
in the foregoing paragraph for the use of the facilities in their herbaria
and libraries or for generous loans of material, but also to Dr. D
Merrill for patiently going over a large part of the manuscript with many
valuable suggestions, to Dr. C. E. Kobuski for helpful criticisms and sug-
gestions, to Dr. I. M. Johnston for assistance in locating literature regard-
ing North American species, to Dr. K. Sax for consultations on cyto-
genetic and hybridization problems, to Dr. D. Wyman for the unselfish
sharing of his profound knowledge of the living plants as well as the
records of the cultivated forms on the grounds of the Arnold Arboretum.
to Miss E. E. Upham for her patience in answering my numerous
questions concerning both English and Latin, and to Mr. H. Howard for
the photographs. I am also indebted to Dr. G. Taylor, Keeper of Botany
at the British Museum and Dr. Charles Baehni, Director of the Con-
servatoire et Jardin Botaniques, Genéve. With the help of Mr. Exell,
the former gave me a detailed description of the nature of the pubescence
on the hypanthium, disk and style of the type of Philadelphus coronarius
Linn. in the Linnaean Herbarium. Through Professor E. D. Merrill, the

278 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

latter sent me the photographs of Schrader’s manuscript and drawings
together with the photocopies of the types of Schrader’s species as pub-
lished in De Candolle’s Prodromus.

HISTORY

The first written account of a plant belonging to Philadelphus is found
in a Chinese poem of the eleventh century describing the fragrance, color
and contrast of the white flower with the green foliage, thus: ‘Herbs for
color, trees for shape; None in fragrance to this [Philadelphus| compare.
Like white foam in a sea of green, Unique among the shrubs it has been.”
Although this passage, like those of Gerarde, Clusius, Bauhin and other
pre-Linnaean authors pertaining to the plant, is nomenclaturally of no
importance to modern taxonomists, it nevertheless indicates the antiquity
of man’s knowledge of the Philadelphus.

By the time of Linnaeus, Philadelphus had already become a widely
cultivated plant in European gardens. In pre-Linnaean literature it ap-
peared as Syringa Tournefort, Frutex coronarius Clusius, or Philadelphus
Athenaci et Rivinii. Linnaeus established the genus in 1737 and validated
it in 1753. He recognized two species, P. coronarius and P. inodorus, and
distinguished them by their leaf margins. Accordingly, the former species
is characterized by its subdentate leaves and the latter by its entire leaves.
The fresh material in the gardens of Clifford and Uppsala provided
Linnaeus firsthand information about P. coronarius. Judging from the
material preserved in the Linnaean Herbarium, he correctly identified it
as representing Clusius’ Frutex coronarius. Of the second species, he saw
no specimen. His binomial was based wholly on the Catesby description
and plate which in turn was based on material observed at the Savannah
River region on the border of South Carolina and Georgia.

From the time of Linnaeus to that of A. P. de Candolle, botanists who
were interested in Philadelphus had their studies limited to the few
species of European and American origin, especially the cultivated forms.
Miller in 1768 added one dwarf species and a variety with variegated
leaves. As he had a larger collection of the living plant belonging to this
genus, he perhaps acquired a more intimate knowledge of the group. He
had seen sterile specimens of P. inodorus Linn. which he raised from cut-
tings sent him by Dale from Carolina. As these plants were killed by
frost when two years old, his records on the height of the plant, the color
of the flower and the size of the fruit were probably abstracted from
Catesby’s accounts. Willdenow in 1809, on the basis of material cultivated
in the Botanical Garden of Berlin, described P. grandiflora, distinguishing
it from P. inodorus Linn, by its prominently toothed leaves. About that
time, P. inodorus Linn. was also in cultivation in a few gardens in England.
Sims in 1812 on the basis of a specimen sent him by Whiltey of the
Fulham Nursery prepared a colored plate and a description for that
species.

During this period, with the exception of Nuttall and Michaux, authors

1954 | HU, THE GENUS PHILADELPHUS 279

on American flora had a very dim concept of the published species of
American Philadelphus. Walter (1788) in the Flora Caroliniana pre-
sented a very general account of P. inodorus Linn. After comparing his
statements about this species with that of Linnaeus, one has a feeling
that he had never seen a specimen of it for he used even the same wording
as Linnaeus. Michaux, being a true field naturalist, had seen the plant
and in his Flora Boreali-Americana published in 1803 he used hitherto
unknown characters about the long acuminate sepals, suboval petals and
elongated undivided style for distinguishing P. inodorus Linn. Pursh in
1814 recognized four species, one of which, P. lewisii, was new. According
to the material he cited, he probably had not seen any P. inodorus Linn.
in America. His description was based on Sims’ illustration and the
cultivated material in England. Elliott in 1821 admitted that he had
seen no specimen of P. inodorus Linn. in the various collections of dried
plants which he had examined. He further stated that the plant was so
rare in nature that no botanists in his day had seen it in the woods.

Nuttall in 1817 described P. hirsutus on the basis of his own collection
from the bank of the French River near Warm Springs, Tennessee. His
description of the species indicates that he was a man of keen and accurate
observations and consequently his treatment gives the truest picture of
the genus up to his time. He recognized four species. As three of them
were American in origin, he was led to conclude that Philadelphus is “a
North American genus, with the exception of P. coronarius.” With our
present knowledge of the genus, there is a great deal of truth in this state-
ment, for not only the largest numbers of species of the genus occur in
North America, but the principal morphological changes of the group
are also found here.

Schrader was the first man who attempted to study the entire genus
extensively. His dissertation on Philadelphus with illustrations which has
been cited by De Candolle and Loudon has never been published. The
manuscript is preserved in the Conservatoire et Jardin Botaniques, Geneve.
Part of it was incorporated by De Candolle (1828) in his Prodromus III.
After Schrader’s death, in the “Reliquiae Schraderianae” published in
Linnaea (1838), an improved form of that dissertation appeared again,
but without illustration. In the manuscript as well as the latter paper he
included nine species and four varieties of Philadelphus. Unfortunately
his material was largely limited to the forms then cultivated in European
gardens. Having no access to specimens representing species already
described by American botanists, he created several synonyms. More-
over, he accepted garden sports or hybrids as representing species and thus
created additional confusion. He divided the then known species into
two sections on the basis of the habit and the inflorescences, but assigned
no sectional names to them. In the section ‘‘Caulibus crassioribus strictis.
floribus racemosis” he recorded five species and three varieties. The rest
were placed in the section “Caulibus tenuioribus virgatis laxis, floribus
solitariis ternisve.” Schrader’s conclusions, whether correct or incorrect,

280 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

were followed by European botanists and horticulturists for over half a
century.

De Candolle in 1828, using Schrader’s manuscript as a standard, pub-
lished a comprehensive treaty on the genus. He recognized eleven species
and three varieties. For the three extra American species not included in
Schrader’s manuscript, he simply annexed them to Schrader’s order and
placed them all in the slender-stemmed and solitary-flowered group. This
represents a very indiscriminate treatment, for closely related species like
P. coronarius Linn. and P. lewisii Pursh, which are alike in their habit
and inflorescences, were placed in separate sections.

Loudon in 1833 in Arboretum et Fructicetum Britannicum gave en
illustrated comprehensive account of the genus. He recognized ten species
and four varieties, placing them in two sections. In the “Stems stiff and
straight. Flowers in Racemes” section he included six species and four
varieties and in the “Stems more slender, rambling, twiggy and loose.
Flowers solitary, or 2 or 3 together” section he had four species. In this
classification what we now interpret as inseparable entities, like P. inodorus
Linn. and P. grandiflorus Willd., were placed in different sections.

Philadelphus from Asia and Central America did not appear in botanical
literature until the eighteen thirties. Wallich was the first person to dis-
cover a species of Philadelphus from Asia. In 1831 he named the speci-
mens procured from Gossainthan, Sirmore and Kamaon as P. tomentosus
and P., triflorus. G. Don in 1832 validated the first binomial by giving it
a description and Royle in 1839 prepared a colored plate for this
Himalayan taxon.

Schlechtendal in 1839 on the basis of Schiede’s collection from Jalapa
and Ehrenberg’s collection from Carmen described P. mexicanus, the first
species of Philadelphus from Central America. About the same time,
Bertoloni (1840) published an illustrated account of a Guatemala species
under the name P. myrtoides, a species which marks the southmost limit
of the spontaneous distribution of the genus.

A. Gray in 1849 on the basis of Fendler’s collection from Santa Fe
Creek, New Mexico, published P. microphyllus, the first known xerophytic
species. The plant was introduced to Europe in 1883 and hybridists there
took advantage of its low habit and very agreeable fragrance and in the
autumn of 1887, the Lemoine Nursery of Nancy, France, put into com-
merce a novelty under the name, P. /emoinei, which was announced to be
the result of a cross between the American small-leaved species, P. micro-
phyllus Gray and the much cultivated European species, P. coronarius.

Lindley and Paxton in 1852 published P. satsumi Sieb., the first known
species of Philadelphus from Eastern Asia, on the basis of both living
material introduced and the herbarium specimens prepared by Siebold
from Japan. Ruprecht in 1856, in a paper of Maximowicz on the im-
portant trees and shrubs of the Amur Region, described P. tenutfolius,
the first known species of Philadelphus from the Eastern Asia mainland.
A year later in a discussion about P. tenuifolius he accidentally published
P. pekinensis which became the first known Chinese species in the genus.

1954] HU, THE GENUS PHILADELPHUS 281

By the middle of the nineteenth century Philadelphus introduced from
America and Asia were cultivated under thirty odd names in European
gardens. Botanists and growers began to be conscious of the confusion
and tried to clarify the nomenclature and to identify the plants. The
general tendency shows a lumping effort. Koch in 1859 in an article
titled, “Notes on the Genus Philadelphus,” first published in Germany
and then translated by De Borre into French, recognized 16 species.
Ten years later, in his Dendrologie he reduced the number to 11 species.
His species concept was rather vague and he clearly erred in making
species of American and Asiatic origin conspecific entities. His work
created confusion rather than clarification. Maximowicz in 1867, in a
“Revisio Hydrangearum Asiae Orientalis,” treated all the species of
Philadelphus that had racemose inflorescences, including those from Eu-
rope, Caucasus, Himalayan Regions, eastern Siberia, northeastern China,
Japan and North America, as varieties of P. coronarius and thus created
a large number of synonyms.

Koehne was the best authority of the group, and he was careful and
keen in his observations. Unfortunately the diagnostic characters he chose
to delimit subsidiary groups between the species and the genus do not at
all well cover the characters of the species involved, and his arrangement
proved to be impracticable. In 1893 he selected the exfoliation of the
bark as the most important character for distinguishing major groups. He
divided the genus into the Corticatae, including those species with closed
bark and the Decorticatae containing those with exfoliate bark. Realizing
this character to be unreliable and the scheme not workable, three years
later, he selected the size of the stigma as the distinguishing character for
dividing major groups. On this basis he proposed two sections, the
Poecilostigma and the Stenostigma. His section Poecilostigma repre-
sents a conglomeration of phylogenetically unrelated species. This section
was subdivided into three subsections, the Gemmati with exposed buds,
the Microphylli containing species with inclosed buds, small leaves and
united and more or less separated stigmas; and the Speciosi including
species with inclosed buds, large leaves and separated broad stigmas. His
section Stenostigma represents species with inclosed buds and separated
narrow stigmas. This was subdivided into four subsections; the Paniculati
with paniculate inflorescences, the Gordoniani with racemose inflores-
cences, decorticate branchlets and late flowering individuals, the Satsu-
mani with racemose inflorescences, decorticate branchlets and early bloom-
ing individuals, and the Coronarii with racemose inflorescences, corticate
branchlets and usually early flowering individuals. In this classification
Koehne failed to give a single character which holds true for the Poeci-
lostigma as a section, and for the demarcation of the Stenostigma he gave
two characters, that is, the narrow stigmas and the inclosed buds. At
first his statement seems to be acceptable. But as one examines the species
that Koehne placed in the Stenostigma section, one finds that this sec-
tion cannot stand as a taxonomic unit, for the first listed species P. calt-
fornicus Benth. has exposed axillary buds, a character which is not

282 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXXV

supposed to exist in the section. Moreover, the differences in the size
and shape of the stigmas between species of this section is just as great as
are those in Poecilostigma. As to the characters he chose for distinguish-
ing subsections they are not strong enough even for delimiting species.
The manner of the peeling of the bark and the time of blooming are sub-
jected so much to the environmental changes that they are not of much
value except for distinguishing closely related garden forms planted in
similar conditions. Although Koehne’s system of classification has been
followed by many authors including Engler in his treatment of the genus
in the second edition of Die Natiirliche Pflanzenfamilien, it is not
adopted in this study, for it represents a more artificial and less
phylogenetic system.

The beginning of the twentieth century seems to mark a new tendency
in the study of Philadelphus. Botanists began to show interest in the
group on a regional basis. Beadle in 1902 concentrated in a study of
Philadelphus of the southeastern United States and proposed three new
species and one new variety from the homeland of P. inodorus. His find-
ings were the basis for the treatment of the genus in Small’s Flora of the
Southeastern United States (1903, 1913) and the Manual of the South-
eastern Flora (1933) of the same author. He introduced the shape of the
base of the capsules for distinguishing species. Thus he characterized
P. inodorus as a species in which the base of the fruit attenuates into the
pedicel and distinguished his new species, P. gloriosus from P. inodorus
by the abruptly contracted fruit base. This character has never been used
by any other botanist in the classification of Philadelphus. As the shape
of the base of the capsule in this genus depends upon the fullness of the
fruit which in turn depends partially upon the environmental conditions
and partially upon the time when the specimens are collected, it has little
value for specific identification.

Rydberg in 1905 prepared a comprehensive treatment of the genus for
North America. The principal diagnostic characters he used were the
presence or the absence of hairs on the style and disk which he called the
upper free part of the ovary, the number of flowers on a flowering branch
and the size of the leaf. He recognized 36 species, 10 of which were new.
In the key, he grouped those species into six subdivisions, the Californici,
Coronarii, Grandiflori, Hirsuti, Microphylli and Mexicani. His work in-
dicates his ability in recognizing differences, and most of his species are
good ones. Although some modern botanists may prefer to place the taxa
he recognized in subspecific rank, that is only a matter of opinion which
does not affect those taxa being distinct entities.

Nakai in 1915 considered the species of Japan and Korea, recognizing
seven species, two of which were new. On the basis of bark characters
of the second year’s growth, whether closed or exfoliate, he raised two of
Koehne’s subsections into sections. In Satsumani he aaa a single
species, P. satsumi and in Coronarii he placed all the other spec

Rehder (1920, 1927, 1940, 1949) considered chiefly the erated
species and varieties. a 1927 he subdivided the genus into six groups:

1954] HU, THE GENUS PHILADELPHUS 283

1. Gordoniani, 2. Sericanthi, 3. Coronarii, 4. Speciosi, 5. Microphylli and
6. Gemmati. Later (1940) he definitely designated these groups as Series.
The characters he employed in the key were the number of flowers on a
flowering branchlet, the pubescence on the hypanthium, the color of the
calyx, etc. He was at times inconsistent when, as an example, he separated
a group of species by the pubescence on their hypanthia, yet under certain
species of this category he recognized varieties with glabrous hypanthia.
as in the case of P. pubescens Loisel. var. intectus (Beadle) Moore. The
diagnostic characters for a series do not always agree with his key char-
acters, as in the case tof P. purpureo-maculatus Rehd. In the text, he
placed this species in series Microphylli which is characteristically a taxon
with hidden buds while in the key it is placed under the category of
exserted buds. His series concept impresses me as rather vague for in
some cases a series consists of species different from each other in many
respects. Thus in Series Gordoniani, he placed P. californicus Benth. with
paniculate inflorescence, glabrous hypanthium and exposed buds as well
as P. pubescens Loisel. with racemose inflorescence, pubescent hypan-
thium and inclosed buds; and in Series Sericanthi he placed P. sericanthus
Koehne with pubescent hypanthium and also P. delavayi L. Henry with
glabrous hypanthium. Moreover, in some cases morphologically insepar-
able and geographically closely affiliated taxa like P. tomentosus Wall and
P. delavayi L. Henry are allotted to different series. Rehder’s identifica-
tion formed the basis for Bangham’s chromosome count of the genus, and
his taxonomic units were adopted by Janaki Ammal in her discussion on
the classification and geographical distribution of Philadelphus. Rehder’s
material was restricted to the cultivated species of the temperate regions.
This geographical limitation combined with the short-comings of his clas-
sification consequently affected the value of the conclusions drawn by the
cytologists.

Hitchcock in 1943 studied the American xerophytic species. His work
contributes materially to an understanding of the group. He considered
four species and thirteen subspecies, all placed in what he called the
ecumeae group. According to him, these small-leaved species, “As a

. are readily distinguished from all other North American species
Be propre yet it is quite apparent that they have been derived
from, and are very closely related to, certain species of the Mexicani.”
I hive carefully examined all the taxa that Hitchcock considered as rep-
resenting a single group, and I can only conclude that this “group”
includes heterogeneous elements. The small leaved character impresses me
as being misleading. The comparative study of the morphological char-
acters and geographical distribution of all the species of the genus
indicates that the elements native of the region considered by Hitchcock
are end products of evolution stemming from two or three directions. It
is true that three of the four species in Hitchcock’s treatment, P. ser-
pyllifoltus Gray, P. purpusii Brandeg. and P. mearnsii Evans are related
to the Mexican elements. But P. microphyllus Gray and its related taxa
are surely distant from them in their phylogenetic relationships. In my

284 JOURNAL OF THE ARNOLD ARBORETUM [| VOL. XXXV

judgement his conclusion on P. microphyllus Gray being a derivative of
P. purpusii Brandeg. is too arbitrary. Before me there are several speci-
mens from the Northern Rocky Mountains, chiefly Montana, with char-
acters intermediate between P. /ewisit Pursh and P. microphyllus Gray.
The size, form and texture of the leaves and the size of the flowers of
these specimens are typical of P. microphyllus Gray, but some, or rather,
most flowering branchlets possess five flowers which is characteristic of
P. lewisii Pursh. These spontaneous intermediate forms are good evidence
of the origin of P. microphyllus Gray and its related species. Nevertheless,
Hitchcock’s paper takes high rank among all those published on the
genus. I believe that it was unfortunate that his studies were limited to
the small-leaved forms, for leaf size is indeed an unreliable character, and
generalizations can scarcely be made except on the basis of a critical
study of all species of the genus.

From the late 1920s up to the present the discoveries of botanists in
fields other than taxonomy have broadened our knowledge of Philadelphus.
Bangham in 1929 ' published the result of his observation on the chromo-
some number of the genus. Janaki Ammal in 1951 ° treated the subject
in greater detail and discovered triploid and aneuploid individuals in
garden forms. These findings will eventually help breeders to obtain better
garden forms, for by suitable mating of parents of known chromosome
compositions, better hybrids with triploid, or perhaps tetraploid or aneu-
ploid chromosome makeups can be induced. As in many ornamental
plants some extra chromosomes often enhance their horticultural merits.

Chaney in 1939 and Condit in 1944* confirmed the occurrence of
fossil Philadelphus in the middle upper Miocene. The fossil P. nevadensis
(Knowlton) Chaney resembles the leaves of P. lewisii Pursh which now
occurs in northwestern United States. This discovery advances our under-
standing of the present distribution of the species in this genus.

EVALUATION OF MORPHOLOGICAL CHARACTERS IN THE
CLASSIFICATION OF THE GENUS

In the historical review of PAiladelphus studies I have noted that the
habit of the plant, the color and exfoliation of the branchlets, the position
of the buds, the size, texture, dentation and indumentum of the leaves,
the inflorescences, and the floral parts have all been used either as major
or minor diagnostic characters in the classification of the species of the
genus. Individual authors have stressed different points and consequently
their systems of classification lack uniformity. In the following discussion
an attempt is made to evaluate characters stressed by others as well as to
add certain fruit and seed data.

The Chromosomes of Some Species of the Genus Philadelphus. Jour. Arnold
Arb. 10: 167-169. 1929.

*Chromosomes and the evolution of Garden Philadelphus. Jour. Royal Hort.
Soc. 76: 269-275. 1951.

. W. Cuaney, Pliocene Floras of California and Oregon, Carnegie Inst. Wash-
ington Publication 553. p. 79. pl. 16. fig. 2. 1944.

1954] HU, THE GENUS PHILADELPHUS 285

Habit: The habit was used as a principal criterion for separating
major groups by early authors like De Candolle and Schrader. As our
knowledge of the genus increased, the importance of the habit in the
classification of Philadelphus gradually decreased. Judging from plants
cultivated in the Arnold Arboretum, there seems to be no correlation
between the habit of the plant and the other morphological characters
such as the position of the buds, or the number of the flowers. Moreover,
the habit is often affected by the environmental conditions and the age
and vitality of the plant. Possibly the habit, such as upright tall stiff
shrub, low widely spreading shrub with twisted branchlets, low compact
shrub with upright branches or moundlike tall shrub with arching branch-
lets, may be used for recognizing special horticultural forms, varieties or
even species, but it should not be used for distinguishing taxa above
these ranks.

Branchlets: In general practice, a few branchlets constitute the only
material a herbarium taxonomist may have for purposes of study and
identification. Regarding the branchlets of Pinlagel pus, my discussion
will be carried on under the headings of the second year’s growth, current
year’s growth, the axillary buds, and the adventitious growth.

The color and exfoliation of the bark of the second year’s growth.
whether closed or exfoliate, has been employed by Koehne, Rehder and
others for distinguishing species or even taxa of higher rank. It is true
that the bark of certain species such as P. pubescens Loisel. and its related
forms are prevailingly gray. But that of the other species is of various
shades of brown, from ash-brown to reddish chestnut. In some species
the bark exfoliates in sheets. In others they slowly wear off. It seems to
be apparent that the size and the age of the shoot, the rainfall of the
growing season, and the amount of snow in the preceding winter are all
contributing factors effecting the exfoliation of the bark. Often different
shoots on the same plant differ in the degree of exfoliation. As the
herbarium specimens represent only very small portions of an individual
plant, the actual selection of specimens may govern the evident exfolia-
tion and bark color very materially. When only a few specimens are avail-
able, one may interpret the bark color and exfoliation of the second
year’s growth as forming a definite criterion for species delimitation, but
when many specimens from the same general area are examined, the
intergradations in color and exfoliation should convince any one that these
characters are of little value in recognizing species.

The current year’s growth of all species of Philadelphus are of two
kinds, the sterile shoots bear leaves only, and the flowering shoots each
of which bears one to three (or very rarely more) pairs of leaves and a
terminal flower or a cluster of flowers. The sterile shoots are more vigor-
ous and their leaves are larger and comparatively more dentate than are
those of the flowering branchlets. The bud position on these shoots is
very constant.

In one group, the buds are hidden in nodal pouches at the ends of the
petioles (Pl. I, fig. 3). When the leaves fall, these buds are covered by

Jour, ARNOLD Ars. VoL. XXXV PLaTE I

Hu, THE GENUS PHILADELPHUS

1954] HU, THE GENUS PHILADELPHUS 287

the abscission layers (PI. I, fig. la, 1b, & 2). As these buds unfold, they
burst open the abscission layers (PI. I, fig. 4) which persist for a long
time at the base of the new branchlets. All the plants with such hidden
axillary buds bear no terminal buds. Their shoots are determinate. All
the Old World species of Philadelphus and many of those of the New
World are characterized by such hidden buds and determinate branchlets.

In contrast, other species bear prominent conical axillary buds on their
sterile and flowering shoots at anthesis (PI. I, fig. 7, 8). In general, nodes
bearing these buds are strongly curved below the insertion of the leaves,
while those with hidden buds are but slightly curved. It is doubtful if
latitude and temperature have any effect on the production and size of
the exposed buds. Thus, P. Airsutus Nutt. is a native of the southern
Appalachian Mountains, but when cultivated in Boston, which is nearly
seven degrees higher in latitude than the original home of the species and
Boston has an average annual minimum temperature 10—15 degrees lower
than Nashville where the plant grows wild, its axillary buds at anthesis
are just as prominent as are those on the Tennessee specimens. Moreover,

PLATE I

1. Shoot apex of the determinate type (x 4, from fresh material collected
in the Arnold Arboretum, Jan. 20, 1952). la. P. coronarius with the lateral
buds completely covered by the abscission layer. 1b. P. microphyllus with
the lateral buds more or less unfolding, the dead terminal portion of the
branchlet is more prominent here. 2. Node of P. delavayi after the leaves
have fallen, showing the lateral buds covered by the abscission layer (x 4,
Rock 16637). 3. Node of P. maculatus with the ends of the petioles attached,
showing prominent nodal pouches containing the buds; the curves below the
insertion of the leaves are gentle (* 4, Mueller 2213). 4. Node of P. delavayi
with the axillary buds unfolding and each abscission layer has burst open (X 3,
Rock 16173). 5. Node of a sucker; A, with two branches B, developed from
normal axillary buds and six branches developed from adventitious buds. Five
of the six are flowering shoots C. and one is a vegetative shoot C’. 6. Shoot
apex of the indeterminate type, P. hirsutus, with a large terminal bud and two
small lateral ones (X 4, from fresh material collected in the Arnold Arboretum
on Jan. 20, 1952). 7. Node of P. texensis with the petioles on, showing the
exposed axillary buds, the curves below the insertion of the leaves are sharp.
(x 4, Reverchon 1523). 8. Node of P. hirsutus after the leaves have fallen,
showing two lateral buds exposed (> 4, fresh material collected in the Arnold
Arboretum, Jan. 20, 1952). 9. Node of P. mearnsii after the buds are unfolded.
The unfolding of the buds does not affect the leaf-scars (* 5, Palmer 11492).
10. Longitudinal section of a fruit of P. inodorus, 1 week after anthesis, show-
ing the attachment of the placentas which are above the insertions of the sepals
(X< 10). 10a, 10b, 10c, and 10d are transverse sections of the above cut through
the points 1, 2, 3, and 4. 10d appears 8-locular. 11. Longitudinal section of a
fruit of P. microphyllus, 2 months after anthesis, showing the attachment of
the placentas which are about the same level as the insertions of the sepals
(xX 10). 12. Longitudinal section of a mature fruit of P. Airsutus showing the
attachment of the placentas which are below the insertions of the sepals ( 10).

288 JOURNAL OF THE ARNOLD ARBORETUM [| VOL. XXXV

such plants often bear strong terminal vegetative buds (PI. I, fig. 6),
therefore their shoots are indeterminate. All the species with exposed
buds occur in the New World.

The presence or absence of exposed buds is not only constant, but the
buds are also very easy to detect. As long as the leaves are mature, no
matter whether the branchlets be sterile or fertile, the buds are evident.
The morphological constancy and the geographical coincidence indicate
that the position of the buds whether hidden or exposed, is a very good
character for separating major groups in Philadelphus. Accordingly, in
this study I take this character as one of the key categories for delimi-
ting the subgenera of the genus.

All species of Philadelphus produce adventitious buds at the basal por-
tions of their principal stems or along the upper parts of their branches.
In the first case, the buds develop into strong sterile shoots which give
rise to flowering branchlets in the following year. In the second case, the
buds may develop into a weaker sterile shoot (PI. I, fig. 5, c’), or usually
they form weak flowering branchlets, (PI. I, fig. 5, c). The leaves of such
shoots are generally smaller and the flowers fewer than are those de-
veloped from normal buds. The presence of a number of such small
flowering branchlets gives the stem somewhat the appearance of a brush.
A branch with multiple branchlets at its nodes should always be taken as
a sign of old age and reduced vitality in the plant.

Leaves: Authors of the first few species of Philadelphus relied chiefly
on leaf characters for distinguishing them. Thus Linnaeus distinguished
P. inodorus from P. coronarius by its entire leaves and Willdenow distin-
guished his P. grandiflorus by its dentate leaves. When only a few speci-
mens are compared, there may be seemingly marked differences in the
size, shape, texture, venation and margin of the leaves. But when a large
number of specimens are available for comparative study, the gradual
changes exhibited in these respects prove that what various authors such
as Schrader, Koch, Maximowicz, and Rydberg accepted as differential
specific characters are not constant. In fact, there is little difference in
the size and shape of the leaves of the species of Philadelphus of any
single area, and also often even between species from distant regions with
similar climatic conditions. The differences that existed in the size, shape,
texture and margin of related species may also occur as modifications
among representative specimens from the same species differing only in
age and vigor. Therefore the size and shape in the leaves can only be
used as supplementary criteria for distinguishing varieties or garden
forms and occasionally even species when it is correlated with characters
furnished by flowers and fruits.

The leaves on the sterile shoots and those on the flowering shoots of
an individual plant differ in size, shape, apex and margin. In general the
leaves on the sterile shoots are larger, more acuminate at the apex, more
coarsely dentate at the margin, more pubescent on the surface and even
broader at the base than are those on the flowering shoots. Yet, leaves
of a flowering shoot developed from an adventitious bud are always much

1954] HU, THE GENUS PHILADELPHUS 289

smaller, more acute, more subentire and more glabrous than are those
developed from normal buds. One judges that the age of the plant, the
vigor of the shoot, the available water supply may be reflected in the
leaves of Philadelphus. For this reason, leaf characters based on these
aspects should not be employed for the delimitation of taxa above varieties
or garden forms.

The nature and the density of the indumentum on the leaf surfaces are
rather constant as to leaves on comparable shoots of a species. They
usually correlate with certain flower characters, therefore may be used
as auxiliary criteria for distinguishing species or categories of even
higher rank, such as section and series.

Inflorescences: Philadelphus as a genus is characterized by determi-
nate terminal inflorescences, each with 1 or 3 flowers at the last node, and
none, one, rarely two single flowers, or very rarely a cyme in the axil of
a bract or a leaf on each side of the succeeding nodes. When the succeeding
node bears no flower, the branchlet has solitary or ternate flowers. When
the succeeding nodes bear flowers the plant has racemose or paniculate
inflorescences, depending on the number of flowers in the leaf axils. In
many species the bracts are small and caducous. Since De Candolle in
1828 used the term raceme to describe P. coronarius Linn. and its related
species, it has been used in reference to those species with a terminal
cluster of flowers in all major treatments of the genus. But actually when
the order of blooming is taken into consideration, none of the Philadelphus
species really bear true racemes. In this genus, the first opened flower
always prohibits the elongation of the flowering shoot and limits the for-
mation of more flower buds. The figures on Plate II actually represent
the number of flowers and their relative developments on the branches of
P. californicus Benth., P. pubescens Loisel., P. lewisii Pursh, P. pekinensis
Rupr., P. inodorus Linn., P. microphyllus Gray, P. falconii Sarg., P. kar-
winskyanus Koehne, P. myrtoides Bertol., P. mexicanus Schlecht., P. hir-
sutus Nutt. and P. serpyllifolius Gray. In each case the flowers at each
node behave as an independent unit. In cases where many flowers are
formed at the end of the flowering branch the terminal one always opens
first. Such inflorescence is a determinate raceme or a panicle.

Philadelphus inflorescences may be roughly arranged in six general
types, the paniculate, the determinate-racemose, the pauciflorous, the
ramiferous, the mexicanus, and hirsutus types. The paniculate type in-
cludes inflorescences composed of simple or more or less compound cymes
(Pl. II, fig. 1). It is represented by P. californicus Benth. and P. cordi-
folius Lange. The determinate-racemose type is the commonest type of
inflorescence in the genus. It is best represented by P. pubescens Loisel.
and P. lewtsiit Pursh in the United States, P. satsumi Sieb. of Japan,
P. tenuifolius Rupr. of the Amur Region, P. pekinensis Rupr. and all the
other Chinese species, P. tomentosus Wall. of the Himalayan Regio and
P. caucasus Koehne and P. coronarius Linn. of Caucasia and southwestern
Europe. It consists of 3, or very rarely 1, or occasionally in P. coronarius
Linn. 5, terminal flowers and several pairs of axillary ones at the suc-

Jour. ARNOLD Ars. VoL. XXXV PiaTeE IT

Hu, THE GENUS PHILADELPHUS

1954] HU. THE GENUS PHILADELPHUS 291

ceeding nodes (PI. II, fig. 2-5). Sometimes, as in P. coronarius Linn., a
pair of flowers may appear in a single leaf axil. The upper axillary flowers
are often subtended by bracts which fall early in the flowering season.
The lower axillary flowers are in general subtended by normal leaves.
Great variations exist in the number of flowers in a cluster, the length
of the central axis and the size of the bracts in specimens representing a
species or sometimes even from a single plant. This is especially true in
P. lewisii Pursh and P. pekinensis Rupr. It seems that the age and vitality
of the shoots which give rise to the flowering branches are the controlling
factors for such modifications. There are certain garden forms character-
ized by a predominantly short flowering axis. In such cases the clusters
appear rather compact (PI. II, fig. 4, 5). There are certain species which
bear loose determinate-racemes. In such cases the lower two pairs of
flowers are borne in the axils of normal leaves (PI. II, fig. 2, 3); the
inflorescences of P. pubescens Loisel. and P. intectus Beadle are typically
of such form.

The pauciflorous type consists of inflorescences with one or three ter-
minal flowers on shoots devoid of exposed axillary buds. This type is best
illustrated by P. inodorus Linn. and P. microphyllus Gray (PI. II, fig. 6,
7) and their related species. In certain garden forms, the lateral flowers
of this type may be replaced by two cymes with elongated peduncles. In
such cases, a true dichasium is formed (PI. I, fig. 8).

The ramiferous types of inflorescences are depauperated panicles. They
may appear as racemes or panicles, but each individual flower is on a short
pedicel jointed to a woody axis. Thus each flower stalk is morphologically
composed of a peduncle and a pedicel. In P. myrtoides Bertol. the pe-
duncles of the lower flowers are as long as those of the upper one and
the inflorescence appears more or less racemose, but the individual pedicels
are jointed (PI. II, fig. 10). In the more ramified form, as represented by
P. karwinskyanus Koehne, the inflorescence is composed of elongated
lower branches (PI. II, fig. 9) and shorter upper ones, and it appears like
a panicle. Some of the lower branches even bear axillary buds on their
basal nodes.

The mexicanus type has one or three terminal flowers on short but
aided a stalks, as found in P, mexicanus Schlecht. Such an inflorescence

PLATE II

TYPES OF INFLORESCENCE IN PHILADELPHUS (all X %). 1. The PANICULATE
type, P. californicus (Abrams 4649). 2-5, The DETERMINATE-RACEMOSE TYPES.
2. P. pubescens (AA 2221). 3. P. lewisii with 3 terminal flowers (Thompson
10522). 4. P. lewisii with 1 terminal flower (Engberg, June 19, 1905). 5. P.
pekinensis (Hers 2515). 6 & 7. The PAUCIFLOROUS TYPE. 6. P. inodorus (AA
4159-1). 7. P. eee (Fendler 266). 8 True picHastum, P. falconert
(Hu, June 14, 1951). 9 & 10, The RAMIFEROUS TYPE. 9. P. karwinskyanus
(FGW 872). 10. P. myrtoides (Carlson 435). 11. The MEXICANUS TYPE,
P. mexicanus (Standley 65124). 12. The H1rsutus TYPE, P. hirsutus (Palmer
35517). 13. The SERPYLLIFOLIUS TYPE, P. serpyllifolins (Cary 9435).

292 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XXXV

is probably a reduced or simplified compound inflorescence with the
portion above the bracts representing the pedicel and the portion below
the joint, the peduncle.

The hirsutus type, as represented by P. Airsutus Nutt. and P. mearnsii
Evans, has inflorescences of solitary or ternate flowers on short flowering
branches with exposed axillary buds (PI. II, fig. 12, 13). Such inflores-
cence often develops from a terminal bud.

The type of inflorescence in Philadelphus is a definite and easily recog-
nizable character. When correlated with the position of the buds, the
form of stigma, the shape of fruit, and the appendages of the seed, the
type of inflorescence can be employed as a very useful device for distin-
guishing sections of the genus as well as for assisting in placing the species
in its phylogenetic position. In using the inflorescence in the classification
of Philadelphus several essential points should be observed. (1) All the
inflorescences of Philadelphus, no matter whether they are composed of
a single flower, or three or five up to 20 flowers in simple or ramified
branches, are borne at the end of current year’s growth. In other words,
they are all derivatives of individual buds which are usually axillary. (2)
All of them are determinate inflorescences; that is, the terminal flower
in each type opens first and they thus limit the elongation of the floral
axis. (3) The type of inflorescence of a species is definite, but the number
of flowers on each flowering branch is variable. A species with pauciflorous
type of inflorescence may have some branches with a single flower, some
with three and others with a true dichasium of seven or nine flowers.
This variation seems to be a reflection of the environmental condition of
the plant as well as the physiological state of the particular flowering
branch. In the cultivated Philadelphus, good soil, sufficient water supply,
proper pruning, vigor of the plant and the bearing year always induce a
large number of flowers on a branch. In case of the determinate-racemose
type of inflorescence the floral axis is always lengthened.

Rickett in 1944 discussed the inflorescences of Philadelphus at length.
He used P. coronarius Linn., P. californicus Benth., P. lewisii Pursh. P.
argenteus Rydb. and P. hirsutus Nutt. to illustrate a hypothesis that the
terminal dichasium is ancestral in Philadelphus. He maintained that this
primitive type can be found in P. Airsutus Nutt., and suggested that the
present forms of the inflorescences of Philadelphus are evolved through
the reduction of the terminal dichasium to a single flower as found in
P. argenteus Rydb. Further reduction of the lateral flowering branches
leads to the production of axillary flowers of P. coronarius Linn. This
condensation is accompanied by a lack of dormancy in the new axillary
buds, which open the same season as the leaves which subtend them.
From flowering branches formed in such a way, derived the floral arrange-
ment as seen in P. /ewisii Pursh and also the thyrse of P. californicus
Benth., the latter represents a condensation of the second order, that is.
not only are a number of the original flowering branches laterally disposed
on a central rachis, but several groups thus constituted are similarly
disposed on the main axis of the thyrse. According to this hypothesis the

1954] HU, THE GENUS PHILADELPHUS 293

evolution of the inflorescences of Philadelphus has been brought about
merely by reduction and condensation of the flowering branches. As I
understand the morphology of Philadelphus, certain assumptions accepted
by Rickett apparently do not conform to what are found in nature. First
he selected the inflorescence of P. Airsutus Nutt. to represent the ancestral
form, but this species is morphologically highly specialized and its low
point of placentation and its ecaudate seeds all indicate its advanced posi-
tion in the evolution of the genus. Secondly, he maintained that the in-
florescence of P. argenteus Rydb. is a derivative of that of P. hérsutus
Nutt., but these two species are phylogenetically very remote. They
belong to different subgenera. Thirdly, the condensation processes that
he assumed to take place between P. argenteus Rydb. and P. coronartus
Linn. and between P. lewisii Pursh and P. californicus Benth. involve
more than a single year’s growth. The plant part he took from P. ar-
genteus Rydb., or P. lewisii Pursh to illustrate his principle represents
two year’s growth with twigs bearing buds on part of old wood and what
he took from P. coronarius Linn. or P. californicus Benth. represent a
single year’s growth, originated from a single winter bud. When he as-
sumed that the branch of P. lewisii Pursh with a number of flowering
branches laterally disposed to be condensed on a central rachis to form
an inflorescence corresponding to those of P. californicus Benth., he over-
looked the age differences of the two elements involved. It is not possible
within the organization of the plant to attain such assumed condensation.
Moreover a shoot of P. lewisii Pursh that bears laterally disposed flower-
ing branches never possesses a terminal bud, so that it is impossible for
that shoot to be condensed on the main axis of the thyrse to form a
paniculate inflorescence of P. californicus Benth. which ends with a ter-
minal flower. Moreover, P. coronarius Linn. and P. lewisii Pursh have
the same determinate-racemose type of inflorescence. When one takes
into consideration all factors, it is unnecessary to invoke imagination to
explain the evolution of the inflorescences of Philadelphus. Rickett was
perhaps right in assuming the terminal dichasium as the ancestral type.
He was partly correct in recognizing reduction and condensation as a
process that brings about the various forms of inflorescences in Philadel-
phus but unfortunately he chose the inflorescence of P. Airsutus Nutt. to
represent the primitive model. The most primitive type of inflorescence
in Philadelphus is the pauciflorous type as found in the true terminal
dichasium of a healthy vigorously growing flowering branch of P. imodorus
Linn. Normally such dichasia may be reduced to three or a single ter-
minal flower. The solitary or ternate terminal flower of P. microphyllus
Gray and all its related species represents a comparatively stable stage
of development of such reduced form. The reduction in the height of the
plant, the size of the leaves, the number of stamens, the length of style
and the size of the stigma all conform with the fact of such reduction in
number of flowers. Multiplication as well as reduction, and some times
a combination of both processes may have taken place in the evolution
of the Philadelphus inflorescences. The multiplication in the number of

294 JOURNAL OF THE ARNOLD ARBORETUM [| VOL. XXXV

flowers which are produced in the leaf-axils of the succeeding nodes below
the terminal one, gives rise to the determinate-racemose type, and further
multiplication which involves the branching of the flowering pedicels
gives rise to the paniculate type. The mexicanus type is another primitive
model which is closely related to the pauciflorous type. The multiplica-
tion in the number of flowers and the ramification of the flowering
branches of this type give rise to the ramiferous type. The hirsutus type
is remote from all other types of Philadelphus inflorescence. Its slender
pedicels with median or super median bracts, its irregular way of branch-
ing, and its appearance remind one of the inflorescence of certain Deutziae.
It is probably derived from some form very remotely related to the mex-
icanus type but the linking form is not known in our present flora.

Pedicels: The length of the pedicels in Philadelphus varies consider-
ably. The general tendency is, in the determinate-racemose type, the
lower ones are longer than the upper ones, or in the pauciflorous types,
the lateral ones longer than the central one. As a criterion for classifica-
tion, this character has little value. Nevertheless, most species have
pubescent pedicels. The nature and density of the trichomes on the
pedicels of closely related species are sometimes employed as auxiliary
characters for recognizing species or varieties.

Hypanthia: Part of the ovary of Philadelphus is fused to the recep-
tacle. In this paper the term hypanthium is used to cover that part called
the calyx tube by some authors. For the visible top portion of the ovary
the term disk is used.

At anthesis, there is no appreciable difference in the shape and size of
the hypanthia of closely related species. But the nature and density of
the indumentum which persist in the fruit are very useful for specific
identifications, or even for the delimitation of series. Most Philadelphus
species have yellowish green hypanthia. Certain Chinese species have
purplish ones. In such case, color may be used as a handy auxiliary
character for identifying species or varieties.

Sepals: The sepals of Philadelphus are ovate, rarely deltoid, 4-20 mm.
long, acuminate (rarely acute) at the apex. The relatively shortest sepals
occur in P. Airsutus and the longest ones in P. inodorus and P. mexicanus.
The length, shape and apex of the sepals have been employed by various
authors for distinguishing morphologically similar and geographically
intermixed species. The study of a large amount of material in this genus
reveals that the natural population of any general area belonging to the
same section exhibits very little variation in the size and shape of the
sepals, Thus, in my opinion, they have no specific significance.

Corolla and petals: Characters of the corolla and the petals have
been little used in the classification of the genus. One Central American
species has been recognized for its pubescent petals, while another species
of northwestern United States has been characterized by its pointed petals.

In shape and size the petals of various species are rather constant. They
may be oblong, elliptic, suborbicular or lanceolate. The variations in the
shape and size of the petals give the different appearance of the flowers.

1954] HU, THE GENUS PHILADELPHUS 295

Thus P. inodorus var. laxus (Schrader) S. Y. Hu has oblong petals, and
its flowers are distinctly cruciform when fully open (PI. III, col. 3). The
petals of P. inodorus Linn. var. grandiflorus (Willd.) Gray are sub-
orbicular, and consequently its flowers appear disciform (PI. III, col. 1).
is an important aspect for distinguishing varieties and garden forms, the
of Philadelphus. It shows a considerable difference in the size, shape and
appearance of the corollas in the genus. As the appearance of flowers
is an important aspect for distinguishing varieties and garden forms, the
size and shape of the petals may eventually have a more important role
in demarcating entities subordinate to the species.

Stamens: There is noticeable variation in the number of stamens not
only between different species or different plants which clearly represent
a single species, but also between different flowers of the same plant. The
forms possessing the most stamens are found in P. inodorus Linn. and
its varieties where as many as 90 stamens have been observed. The
smallest number of stamens observed is in P. mearsnit Evans where as few
as 13 stamens have been counted. But variations in the number of stamens
between different flowers on the same plant do exist. With the first cited
example as few as 60 stamens and with the second species as many as
20 stamens have been observed. For demarcating morphologically closely
related and geographically inseparable species the number of stamens
does not seem to have much significance. But there apparently exists a
general pattern of stamen number between morphologically different and
geographically widely separated groups. Thus the native population of
Philadelphus of the Southern Appalachian region with pauciflorous in-
florescences have the highest stamen count which is between 60 and 90;
the xerophytic plants of the southern Rockies with the pauciflorous type
of inflorescences have an average count of 30 to 40, rarely as low as 24 or
as high as 50 stamens; the native species of Central America with rami-
ferous inflorescences have 40 to 50 stamens and those with the mexicanus
type of inflorescences have 36 to 44 stamens. The Old World species with
racemose inflorescences generally have 25 to 35, rarely to 50 stamens.
Garden hybrids usually have low stamen counts, and many of them are
sterile, with the exception of one form, which has up to 60 stamens, they
have 20 to 29, rarely up to 40 stamens. The species with the hirsutus
type of inflorescences have the lowest stamen count, ranging 13 to 33.
Such a general pattern when correlated with the form of buds, the type
of inflorescences and the shape of fruits and the tails of the seeds, may
serve as auxiliary criteria for characterizing sections or series of the

Hitchcock was the first person who observed the tendency of the union
of the lower portions of the filaments of several adjacent stamens into
bundles in P. argenteus Rydb. (Pl. IV, fig. 8c), and used this character
for distinguishing that xerophytic taxon. Normally the filaments of
Philadelphus are separated. They are always shorter than the petals.
With the exception of the extreme xerophytic dwarf forms like P. ser-
pyllifolius Gray or P. mearnsii Evans, which has very small flowers and

Jour. ARNOLD Ars. VoL. XXXV PLaTE III

Hu, THE GENUS PHILADELPHUS

1954] HU, THE GENUS PHILADELPHUS 297

short stamens, the length of the filaments in Philadelphus is rather con-
stant, with the longest outermost ones being 6-9 mm. long and that of
the shortest innermost ones 2-3 mm. long. In the xerophytic forms, the
filament of the outer stamens measures up to 4 mm. long and the inner
ones 2 mm. long. So far as I know, the longest filament of Philadelphus
is found in a garden form called P. ‘“slavini” hort ex Wymn. with the
inner filaments 7 mm. and the outer ones 12 mm. long. The filaments of
Philadelphus have not and very likely never will furnish outstanding
canis for distinguishing taxa of any rank in the classification of the

gen

The mane of Philadelphus are generally oblong, being 1—-1.2 mm.
long, 0.9 mm. wide. The xerophytic species like P. serpyllifolius ci
has Snleboe anthers, 0.5 mm. in diameter. There are a few natur
populations occurring in Central China and western North America ie
hair-like processes occur on the thecae of the anthers (PI. IV, fig. 8c).
This is probably an expression of an unusual mutation which Poatally
may lead to speciation.

istil: A normal pistil of Philadelphus is composed of four carpels.

Occasionally few 5-carpelled or very rarely some 3-carpelled pistils may
occur in a garden form, Characters furnished by the parts of the pistil,
ovary, style, and stigmas, have been employed for distinguishing taxa
of various rank in Philadelphus. Schrader recognized P. speciosus for its
8-loculed fruit. Pursh distinguished his P. /ewisii for its relatively long
and much divided style. Koehne subdivided the major groups of the
genus on the basis of the size of the stigma. Ruprecht, Koehne and
many others have proposed species because of the presence of hairs on the
style, or disc, or both. Nevertheless, due to our limited knowledge of the
pistil in PAiladelphus unnecessary binomials have been created. A normal
Philadelphus ovary has four locules and an axile placentation. In each
locule there are a pair of more or less fused auricular placentas projecting
from the upper portion of the central column (PI. I, fig. 10-12). The
superior lobes of these placentas are much shorter than the inferior ones,
the latter are so pressed together that they appear as one organ in cross
sections (Pl. I, fig. 10B). In P. inodorus Linn., its varieties, and its re-
lated species, the inner wall of the roof of each locule, which corresponds

PLATE III

FLOWERS OF PHILADELPHUS CULTIVATED IN THE ARNOLD ARBORETUM SHOW-
ING DIFFERENT FORMS OF THE COROLLA, THE READING IS BY THE COLUMNS 1, 2,
3, AND 4, EACH FROM THE TOP TO THE BOTTOM:

1 a og Weenie! ‘e ”P. grandiflorus, P. caucasicus, P. coronarius “spe-
ciosissimus,”’ P. coronarius, var. primulaeflorus, P. coronarius; COLUM
insignis (?), P. incanus, P. falconeri, P. floridus, P. delavayi, P. “Cole aileadtl
CoLtumn 3. P. “Belle Etoile,” P. satsumanus, P. lemoinei, P. laxus, P. ma
dalenae, P. splendens; CoLUMN 4. P. maximus, P. sericanthus, P. tenuifolius,
P. pekinensis, P. nepalensts, P. “bicolor”; CoLuMN 5. P. “Favorite,” P. vir-
ginalus, P. “Norma,” P. zeyheri, P. domentosud, P. purpurascens.

298 JOURNAL OF THE ARNOLD ARBORETUM [VOL, XXXV

to the upper portion of the middle of the carpel, grows down and projects
between the superior lobes of the placentas in such a manner that it
appears like a false septum separating those lobes. Thus the cross sections
made through the upper portion of a normal 4-locular ovary of such species
appear 8-chambered (PI. I, fig. 10D). This was perhaps what Schrader
saw in P. speciosus. The position of the attachment of the placentas on
the central column affects the shape of the fruit and the length of the
tail of the seed. In P. inodorus Linn. the placentas are attached above
the insertion of the sepals. Consequently the fruits are ellipsoid with
circumferential sepals. These fruits have long-tailed seeds (PI. I, fig. 10).
On the other hand, the placentas of P. microphyllus Gray are attached
about the same level as the insertion of the sepals (Pl. I, fig. 11). The
fruits are ellipsoid with supermedian persistent sepals, and the seeds are
short-tailed. In P. hirsutus Nutt. the placentas are attached below the
insertion of the sepals (PI. I, fig. 12). The fruits are obconic with apical
persistent sepals and the seeds are ecaudate. This correlation of the at-
tachment of the placentation, the shape of the fruits and the length of
the tails of the seeds indicate that the position of the placentas can be
used as auxiliary criteria for the delimitation of subgenera or sections
of Philadelphus.

The length and the degree of the union of the style, the number and
shape of the stigma and the indumentum of the style and disc have all
been used in the classification of Philadelphus. Michaux in 1803 in dis-
tinguishing P. inodorus Linn. from P. coronarius Linn. introduced the
relative length of the style and the stamens and the degree of union of
the style. Since then Pursh has characterized P. lewisii as a species with
three stigmas and a 3-parted style as long as the stamens, and Nuttall
has characterized his P. hirsutus as a species with clavate undivided 4-
grooved stigma and a style shorter than the stamens. Fortunately the
species that Michaux, Pursh and Nuttall dealt with each represents a
phylogenetically unrelated group, and incidentally the character holds
true for distinguishing them. Later authors including Rydberg and
Rehder have used the same character for delimiting closely related
species, varieties and garden forms which present varied degrees in the
union of the styles. Philadelphus microphyllus Gray has been treated as
a species with united stigma. But flowers of different collections exhibit
varied degree of the union of the styles. Some are completely united, some
are partially united and others are completely separated (PI. IV, fig. 8a,
b, e, f). This happens also in P. argyrocalyx Wooton (PI. IV, fig. 9 A-b).
So far as I know there is a variation in the degree of the union of the
style in every natural population. For this reason such characters fail to
be of value for distinguishing species or taxa of lower ran

There are certain morphological differences existing in the styles and
stigmas of geographically separated groups, which, when correlated with
characters as the position of buds and the type of inflorescences, may be
used as auxiliary criteria for demarcating series or sections. The styles
of the southern Mexican species may be long or short, but they all have

1954] HU, THE GENUS PHILADELPHUS 299

elongated cristate stigmas with the fertile papillose surfaces situated
largely on the elevated abaxial sides (Pl. IV, fig. 13-15). The styles of
the southern Appalachian species with pauciflorous inflorescences are rela-
tively long. They carry oar-shaped stigmas with adaxial as well as abaxial
fertile papillose surfaces, the latter though definite in outline, are shorter
(Pl. IV, fig. 12). The styles of the Old and the New World species with
determinate-racemose inflorescences are comparatively long with clavate
or linear stigmas. The fertile surfaces of these stigmas, with the exception
of some Yunnan and Japanese species, are longer on the adaxial sides
than on the abaxial sides (Pl. IV, fig. 1-6). The styles of the New
World xerophytic species with pauciflorous inflorescences are relatively
short with linear stigmas, the fertile surfaces of which are largely on the
abaxial sides (Pl. IV, fig. 8-9). The styles of the New World species
with the hirsutus type of inflorescences are columnar with subcapitate
4-grooved stigmas (PI. IV, fig. 16-17).

The presence or absence of on the disc and style is a definite and
easily recognizable character which may safely be used in the identification
of species. Ruprecht in 1857 first used such a character for P. schrenkit,
Koehne later described P. lancifolius from Sikkim, P. karwinskyanus
from Mexico, P. subcanus from Central China and P. caucasicus from
western Asia on the strength of the same character. Rydberg in 1905
employed this character as the principal criterion in his key. But as
abundant material is available for a comparative study of the natural
populations of various regions this character is probably not as important
as earlier taxonomists considered it to be. In nature among closely related
taxa in all regions, there are paired species which differ chiefly in the
absence or presence of hairs on the style, disc or both. For example, there
are the P. coronarius and P. caucasicus in southern Europe and western
Asia, P. tenuifolius and P. schrenkii in northeastern Asia, P. subcanus and
P. incanus in Central China, P. satzumi and P. lasiogynus in Japan, P.
lewisti and P. gordonianus in northwestern United States, P. stramineus
and P. pumilus in southwestern United States, P. pubescens, and P.
gattingii in southeastern United States, and P. coulteri and P. asperi-
folius in Mexico, It is not impossible that such phenomena merely
represent an expression of normal Mendelian inheritance of one char-
acter. But our knowledge of the genetics of these groups is lacking. For
the present, and in the absence of cytogenetic information, the presence
or absence of hairs on the disc, style, or both is retained for the purpose
of distinguishing certain manifestly closely related taxa.

Capsules: The fruit of Philadelphus has not been used to any extent
in the classification of its various taxa. The material that I have ex-
amined apparently presents subgeneric or at least sectional differences in
the shape of the capsules and the position of their persistent sepals. The
southeastern Mexican species all possess ellipsoid or obovoid fruits with

ellipsoid fruits and median persistent sepals while the species of the same

Jour. ARNOLD Ars. VoL. XXXV PLaTE IV

Hu, THE GENUS PHILADELPHUS

1954] HU, THE GENUS PHILADELPHUS 301

area possessing the hirsutus type of inflorescences have obconic fruits with
apical persistent sepals (Pl. IV, fig. 25). The position of the persistent
sepals may be used as a satisfactory auxiliary criterion in distinguishing
morphologically close but geographically remote species. Philadelphus
lewisii Pursh and P. pekinensis Rupr. both have pseudoracemose inflores-
cences and glabrous hypanthia. Besides the geographical separation, the
subapical sepals attached to the fruit of the former and the apical ones

PLATE IV

THE STYLE AND STIGMA OF Philadelphus (all X 2%). 1. P. coronarius, the
stigmas slightly enlarged, the abaxial surface definite, shorter than the adaxial.
2. P. caucasicus-with pubescent style and disk. 3. P. coronarius var. duplex
(Kew 1307); the stigmas broadened. 4. P. tomentosus, the adaxial surface split
into two. 5. P. pekinensis, the abaxial surfaces much shorter than the adaxial.
6. P. kansuensis, the stigmas often 3, the disk with few hairs, the abaxial sur-
face much longer than that of P. pekinensis. 7. P. delavayi, a. and c. from
Yunnan (Feng 1012), b. from Latong, Tibet (Younghusband). 8. a—b and d-f.
P. microphyllus with the styles separated in various degrees, the fertile surface
of the stigma abaxial (a. Ellis 107, b. Eggleston 18654, d. Heller 3792 and f
Alexander-Kellogg 1785). 9. P. argyrocalyx with the stigmas separated or
united (a. Eggleston 14541, b. Wooton in 1895). 10. P. argenteus with the
style slightly separated (Mearns 1617). 11. P. inodorus with oar-shaped
stigmas, the abaxial surfaces broadened, shorter than the narrow adaxial sur-
faces (AA15347). 12. P. “bicolore” with elongated style and enlarged stigmas
(Hu in 1951). 13. P. affinis, the style long and hairy, the stigmas elongated
with few hairs on the sterile portion (Berlandier 333). 14. P. calcicolus with
the stigmas more or less united, broadened and cristate, the style short, glabrous
(Meyer & Roger 2662). 15. P. karwinskyanus with the stigmas cristate and
hairy on the sterile portion, the style short and pubescent. 16. P. hirsutus
with the style comparatively long, the stigmas clavate and undivided, the
stigmas on 4 ridges. 17. P. serpyllifolius with the style short, the stigmas
undivided and subcapitate.

eee . Philadelphus (X 1). 18. P. tenuifolius with the apical end rounded
(Cen. 263). 19. P. tomentosus with the apical end pointed and the persistent
calyx ae circumferential (Parker, in 1919). 20. P. sericanthus with the
apical mn rounded, the lower portion pubescent. (Steward, Chiao & Cheo
609). P. pekinensis with very small fruit (King 626). 22. P. inodorus
with oe fruit. 23. P. argyrocalyx with subglobose fruit, the lower por-
tion lanate (Rehder 315). 24. P. hirsutus with flat apical end (Biltmore
Herb. 4333). 25. P. hitchcockianus with very small top-shaped fruit which is
broader than long (Moore 3477). 26. P. sargentianus with subglobose fruit
(Pringle 2094). 27. P. karwinskyanus with ellipsoid fruit pointed at both ends

Rose & Hough 4412). 28. P. mexicanus with obovoid fruit the lower portion
4-angled. 29. Cross-section of the fruit of P. mexicanus, one third from the base.

SEEDS oF Philadelphus (X 10). 30. The short-caudate seed of P. pekinensis
with round-lobed crown. 31. The long-caudate seed of P. inodorus with pointed-
lobed crown. 32. The short-caudate seed of P. californicus. 33. The ecaudate
seed of P. hirsutus.

So NUSUAL STAMENS OF Philadelphus. 8c. Stamens with fused filaments
(Eggleston 18654). 8g. Stamen with hirtellous anther (Abram 7200).

302 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXXV

to the fruit of the latter are outstanding characters for distinguishing these
species.

Seeds: Seeds are minute in all species of Philadelphus. Each indi-
vidual seed consists of a small ellipsoid body about 1-2 mm. long wrapped
in a thin testa extending towards both ends. The large portion of this
small body is the copious oily endosperm which imbeds the microscopic
embryo. The hypocotyl and the radicle constitute two-thirds of the
embryo proper. The cotyledons and the epicotyl are very minute. The
testa is reticulate, light brown, castaneous, or rarely black. It extends into
an irregular collar-like crown at the proximal end and a tail-like wing
at the distal end. The crowns of the seeds of different species vary in
shape. That of the seed of P. pekinensis Rupr. has more or less rounded
lobes (PI. IV, fig. 30), that of the seed of P. inodorus Linn. has pointed
lobes (PI. IV, fig. 31), and that of the seed of P. hirsutus Nutt. is very
minute and inconspicuous (PI. IV. fig. 33). The length of the seed-tail
varies as well as the shape of the crown. Philadelphus inodorus Linn.
has a long-tailed seed, P. pekinensis Rupr. has a short-tailed seed and
the seed of P. hirsutus Nutt. has no tail. Seed characters have never been
used in the classification of Philadelphus. The shape of the crown and
the length of the tail of the seed correlate with the position of the buds,
the type of the inflorescence, and the form of the stigma. In my opinion
they are characters of subgeneric or sectional importance. In some cases
they can be used to distinguish intricate species. For example, because
of the superficial similarities of the leaves and the corollas of P. lewisii
Pursh and P. californicus Benth. and also because of their overlapping
geographic distribution, some botanists have placed the latter as a variety
of the former. As P. lewisii Pursh has long-tailed seeds and P. californicus
Benth. has short-tailed seeds (PI. IV. fig. 32), and as this character is asso-
ciated with the difference in the inflorescences, P. californicus Benth.
should be considered as a good species.

n summarizing the study of the gross morphology of Philadelphus, it
seems legitimate to conclude that characters such as the dichasial inflores-
cence, large number of stamens, elongated stigma with enlarged papillose
fertile surface, high insertion point in the placentation, ellipsoid fruit with
circumferential calyx lobes, and long-tailed seed can be interpreted as
primitive ones, while in contrast, characters like solitary flower, the
racemose or paniculate inflorescence, the small number of stamens, the
reduced length of the stigma, the low point of insertion in the placentation,
the obconic fruit with apical persistent sepals and the short-tailed seed
can be a as representing more advance stages in the evolution
of the genus.

SUBDIVISIONS OF THE GENUS AND THE PHYLOGENY
OF ITS MAJOR GROUPS

Based on the data found in the position of the buds, in the type of
inflorescences, in the form of stigmas, in the shape and position of the
persistent sepals on the fruits, and the nature of the seeds, I am arranging

1954] HU, THE GENUS PHILADELPHUS 303

the species of Philadelphus into four subgenera and nine sections. The
characters of these taxa are stated in the following synopsis.

SupcENuS I. Gemmatus: This subgenus includes all the species with
exposed buds; depauperate paniculate inflorescences, one or three ter-
minal flowers with jointed pedicels or many flowers in ramified branchlets;
stamens 40—60; semi-inferior ovary, enlarged and often cristate, separated
stigmas; obovoid-ellipsoid or subglobose fruits with circumferential sepals;
and long-caudate seeds. It contains 14 species in two sections.

SecTION 1. Poecilostigma: Inflorescences compound, I- up to many-
owered; stamens 40 to 50; stigmas elongated and cristate;
fruits obovoid-ellipsoid; southeastern Mexico, Guatemala to
Costa Rica.

Section 2. Coulterianus: Inflorescences with solitary flowers, rarely
3-flowered: stamens 30—44; stigmas elongate, not cristate;
fruit subglobose; northeastern Mexico.

SusBcENus II. Euphiladelphus: This subgenus includes all the
species with enclosed buds, determined branches, determinate-racemose or
pauciflorous inflorescences, large or medium number of stamens, clavate
or oar-shaped or linear stigmas, and long- or short-tailed seeds. It com-
prises 41 species in three sections.

Section 3. Pauciflorus: Inflorescences true dichasial cymes, often re-
duced to one or three flowers; stamens 60-90; stigmas oar-
shaped; fruit ellipsoid with circumferential sepals; seeds
long-tailed with pointed lobes on the crown; two species,
native of the Southern Appalachian region of the United
States.

SecTION 4. Stenostigma: Inflorescences determinate-racemose; — sta-
mens 25-35, rarely over 40; stigmas clavate or linear; fruit
ellipsoid with subapical persistent sepals; seeds long- or short-
tailed, the crown with pointed or rounded lobes; 30 species
in both the Old and the New Worlds.

SecTION 5. Microphyllus: Inflorescences pauciflorous; stamens 25—40,
rarely up to 50, stigmas linear; fruit ellipsoid with circum-
ferential or subapical persistent sepals; seeds short-tailed
with rounded lobes on the crown; 11 species, Colorado
Plateau, Mexican Highland, and northern Mexico.

SuBGENUS III. Marcrothyrsus: This subgenus includes species with
exposed buds; determined branches, paniculate inflorescences; medium
number of stamens; clavate stigmas, and short-tailed seeds. It has one
section including three species, native of California.

SecTIon 6. Californicus: Characters as the subgenus.

304 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXXV

SUBGENUS IV. Deutzioides: This subgenus includes all the species
with exposed buds, and indeterminate branches; solitary, rarely ternate
flowers; 13-35 stamens; inferior ovary, and columnar or subcapitate 4-
grooved stigmas; turbinate or subglobose fruits with apical persistent
sepals, and ecaudate seeds. It comprises eight species in three sections.

SECTION 7. Hirsutus: Flowers ternate; mesophytic plants; leaves ser-
rate, hirsute, hairs all straight, style 4 mm. long; two species,
Tennessee, Alabama.

SEcTION 8. Pseudoserpyllifolius: Flowers solitary; xerophytic, dwarf
ants, leaves entire, strigose or strigose-villose, hairs all
straight: style up to 1 mm. long; four species, Texas, New

Mexico, and Northern Mexico.

SECTION 9. Serpyllifolius: Flowers solitary; xerophytic, dwarf plants,
eaves entire, lanate and hirsute beneath, trichomes dimor-
phous; two species, Texas, New Mexico, Arizona, Northern

Mexico.

The interrelationship of the subgenera and sections is indicated graph-
ically in the following figure.

In this classification primitive characters such as enlarged stigmas, high
point of attachment in the placentation and long-tailed seeds are found
both in the section Pauciflorus of the subgenus Euphiladelphus and the
section Poecilostigma of the subgenus Gemmatus. The true dichasial in-
florescences and the very large number of stamens of sect. Pauciflorus
indicate that species in this section retain more primitive characters than
those of sect. Poecilostigma which has apparently reduced compound in-
florescences and moderate number of stamens. Yet the exposed buds of
sect. Poecilostigma and the elongated cristate stigmas of many of its
included species show that this section is more primitive in these respects.
Morphological evidences, obviously suggest that our present species of
Philadelphus seem to express two lines of evolution. They probably have
originated from a common stock with exposed buds, dichasial inflores-
cences, exceedingly large number of stamens, and elongated cristate
stigmas. Such a hypothetical ‘“Protophiladelphus” does not exist in our
present flora. The species of the section Pauciflorus occur in the meso-
phyllous forest of the southern Appalachian region, and those of the sec-
tion Poecilostigma are associated with vegetations constituting the same
type of forest in Mexico.' Apparently species in these two sections have
existed for millions of years with very little changes.* They both represent
very ancient stocks in the development of the genus.

TA. J. Suarp, selena of the Vegetation in Certain Temperate Regions of
Eastern a logy 31: 313-333. 1950
*S: he Ter rtiary Character of ‘the Cove Hardwood Forests of the Great
Smoky sane National Park. Bull. of the Torrey Bot. Club 70: 213-235. 1943;
d A. J. Su ser es relation of the Eocene Wilcox Flora to some Modern Floras.
Evolution 5: 1

305

1954] HU, THE GENUS PHILADELPHUS

Species of the sect. Stenostigma share the character of enclosed buds
with the sect. Pauciflorus. This section with its determinate-racemose in-
florescences, moderate number of stamens and some long-tailed but largely
short-tailed seeds seems to have derived from the sect. Pauciflorus, the
direct link seems to be from the P. inodorus type of plant to the P. lewisi
type of plant. In this process of evolution, the changes involved are: (1)
the suppression of the branching of the rachis of the lateral flowers of the
true dichasia of the sect. Pauciflorus, thus prohibiting the formation of

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HYPOTHETICAL PROTOPHILADELPHUS
e subgenera and sections of Philadelphus as
l

Fic. 1. Relationships of th
by morphological similarities. The solid lines indicate relative close

indicated
relationships while the broken lines indicate lesser affinities.

the lateral cymes, (2) certain genetic modifications which promotes the
differentiation of the tissue in the shoot apex, so that flower primordia are
formed not only at the terminal nodes, but also in the axils of the leaf
primordia, and consequently the determinate-inflorescences of the sect.
Stenostigma are developed, (3) the reduction of the number of stamens,
(4) the narrowing of the stigmatic surfaces, and (5) the lowering of the
point of attachment in the placentation. The shortening of the seed-tails
represents a more advanced stage of evolution in this section. These
changes must have taken place before the Miocene period of geological
time for fossil species of Philadelphus resembling the present small-leaved
forms of P. lewisii Pursh existing in the Upper Miocene have been dis-
covered by Chaney in Oregon and Colorado. It is very probable that
during the Eocene period when North America, northern Asia and north-

306 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXxXV

ern Europe supported a temperate and subtropic flora, the species of the
section Stenostigma were so well established and so widely distributed on
the American and Eurasian continents that they had a holarctic distribu-
tion and a transcontinental range. The present disjuncted Old World and
New World distribution represents its disturbed range and the existing
species of this section may hence be considered as “living fossils’ corre-
sponding to the situation of Sequoia and Metasequoia.

The section Microphyllus culminates the Euphiladelphus line of descent.
Its species share the characters of narrow stigmas and short-tailed seeds
with the section Stenostigma. The majority of them have small leaves
and pubescent hypanthia, characters common to xerophytic species. In
the course of the evolution of these species the changes involved are chiefly
in the development of the dwarf habit of the plant, in the reduction of the
size and the increase in the thickness of the leaves, the formation of dense
indumentum and in the suppression of the formation of the axillary
flowers. These changes must have taken place at or after the end of the
Eocene period when America saw the elevation of the Rocky Mountains.
The consequent aridity necessitated such adaptation for xerophytic living.

The subgenus Macrothyrsus, though small in constitution, has a peculiar
position in the classification and evolution of Philadelphus. On one hand
it shares the moderate number of stamens, the clavate stigmas, the me-
dianly attached placentation and the short-tailed seeds with species in the
Stenostigma section of the subgenus Euphiladelphus but on the other
hand its exposed buds link it to the subgenus Gemmatus. In addition to
this peculiar combination of characters its paniculate inflorescence is
very unique. It is probably most closely related to the subgenus Euphila-
delphus. The general habit, the leaves and the appearance of the indi-
vidual flowers of its included species, P. californicus Benth., resemble
those of P. lewisit Pursh, a species of the section Stenostigma of the
Euphiladelphus, so much that it has tempted botanists to treat it as a
variety of the latter species. Apparently this stock has originated from
the subgenus Euphiladelphus with a tenuous influence of the section
Gemmatus.

The species of the section Coulterianus share the exposed buds and
the long-tailed seeds with the Poecilostigma. Apparently this section
represents a continuation of the Gemmatus line of evolution. In this stock
a general tendency in the adaptation for a more xerophytic habitat is very
prominent. It is shown both in the reduction of the size of the leaves and
in the increased thickness of the indumentum. The stigmas of this section
are elongate but not as a rule cristate, and the fruits are more ur less
globose in outline. In the size of the leaves, the pubescences of the hy-
panthia and in the form of the fruits, some species of this section, such as
P. sargentianus S. Y. Hu express a strong resemblance to P. argyrocalyx
Wooton of the section Microphyllus of the subgenus Euphiladelphus.
These similarities are rather superficial. The gap between these species is
wide for they differ in the fundamental criteria as to the position of the
buds and the nature of the seeds.

1954] HU, THE GENUS PHILADELPHUS 307

Morphologically the subgenus Deutzioides is only remotely related to
Philadelphus. In the low point of attachment of the placentation, the
obconic fruit and the tailless seeds, this subgenus shows a close affinity to
the Asiatic genus Deutzia. In fact, the appearance of a branch of P.
hirsutus Nutt., the type species of the section Hirsutus, resembles that of
a branch of Deutzia grandiflora Bge. of West China (Szechuan and Yun-
nan) more than it does of any American species of Philadelphus. Never-
theless, the species of this subgenus should not be separated from PAztla-
delphus because of the indefinite number of unappendaged stamens and
the simple straight hairs. In Deutzia the stamens are usually 10 and are
always appendaged, that is, the filaments are winged. The trichomes of
Deutzia are all stellate. The point of origin of this line of evolution in
Philadelphus is not clear, but evidently it was not from the stock which
gave rise to the Euphiladelphus species. The exposed buds of this sub-
genus indicate its faint affinity with the subgenus Gemmatus. Yet its
ecaudate seeds, the columnar styles, and the coalescent stigmas imply a
wide gap between them. It seems apparent that in the course of the evolu-
tion of Philadelphus, there has been a tendency towards the production
of Deutzia-like plants. It appears that in the New World the process
stopped in the subgenus Deutzioides, while in the Old World the change
was more abrupt, the resulting genus, Deutzia, once established, diver-
sified more rapidly than Philadelphus, and consequently occupies a wider
area than Philadelphus. This conforms with Professor Sax’s finding in
the chromosome numbers of the two genera.' Accordingly, the basic
chromosome number of Philadelphus and Deutzia are the same, n = 13,
but in Deutzia, many species are polyploids. The xerophytic taxa of the
subgenus Deutzioides may have the appearance of some species of the
section Microphyllus. For example, P. mearnsii Evans of the section
Pseudoserpyllifolius resembles P. pumilus Rydb., and P. texensis S. Y. Hu
of the section Serpyllifolius has the appearance of P. microphyllus Gray.
But these resemblances are superficial. Philadelphus mearnsii and P.
serpyllifolius have exposed axillary buds, columnar styles, and ecaudate
seeds and P. pumilus and P. microphyllus have enclosed buds, divided
styles and short-tailed seeds. These superficially resembled species are
developed from different stocks, the separation of which occurred in re-
mote geological times. The exposed buds and the large serrate leaves of
P. hirsutus Nutt. resemble those of P. mexicanus Schlecht. But the low
point of attachment in the placentation, the columnar styie and the
ecaudate seeds of the former species indicate its advanced position on the
genealogical tree of the genus. It is legitimate to assume that the repre-

into the present forms. The species of the section Pseudoserpyllifolius are
pygmies of this line of development and come about through the reduction
in the size of the leaf to meet changed climatic and topographic require-

“K. Sax, Chromosome numbers in the Ligneous Saxifragaceae. Jour. Arnold Arb.
12: 198-204, 1931

308 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XXXV

ments characteristic of a xerophytic environment. They are intermediate
forms between members of the sections Hirsutus and Serpyllifolius. In
indumentum they resemble the former and in size and general appearance
they resemble the latter. The species of the section Serpyllifolius repre-
sent the climax of this line of evolution.

CYTOLOGY

Investigations of the cytology of PAiladelphus have been undertaken
by W. Bangham in Boston and Janaki Ammal in London. Bangham in
1929 ! examined the pollen mother cells of 37 species, hybrids and varieties
of Philadelphus grown in the Arnold Arboretum and concluded that there
‘is no marked difference in the chromosome groups among those taxa. He
reported that each species had a diploid chromosome count of 26. He
observed that the chromosomes of the hybrids were perfectly compatible
and there was no evidence of lagging and other aberrant behavior. His
observation on the chromosome behavior of a garden form of P. pubescens
Loisel., which was then assumed by Rehder to be a cross between P.
tomentosus Wall. of the Himalayan Region and P. pubescens Loisel. of
southeastern United States, led him to conclude that there must have been
very little change in the chromosome makeup of P. tomentosus Wall. and
P. pubescens Loisel. in the millions of years that they, or their ancestors,
have been separated.

Janaki Ammal in 1951 studied the chromosome number and behavior
of Philadelphus grown at the Royal Botanic Gardens, Kew and the gardens
of the Royal Horticultural Society at Wisley, England. She confirmed
Bangham’s findings that the natural species are diploids with the chromo-
somes 2n = 26. But her observation of the chromosome behavior of the
hybrids differs from that of Bangham. She reported that lagging and
other types of aberrant behavior are generally associated with hybridity
in Philadelphus and consequently give rise to high pollen sterility. In
the case of the trispecific hybrids, P. purpureo-maculatus Rehd., the
chromosome behavior is so irregular that it causes the production of un-
reduced germ cells. By the fertilization of an unreduced egg cell with a
normal pollen, triploid garden forms resulted. She found that P. “Sybille,”
P. “Belle Etoile” and P. “bicolor” are all triploids. She also observed a
newly produced hybrid, P. “Beauclerk,” with tetrasomic diploid chromo-
some composition, 2n = 28. She thus concluded, ‘though differences in
chromosome number do not exist in nature, there are differences in mor-
phology of the chromosome which interfere with pairing in their hybrids
between widely separated species . .. for the first time since Eocene
times polyploidy has been induced in a genus which has remained diploid
for millions of years in nature, by the bringing together under cultivation

*The Chromosomes of Some Species of the Genus Philadelphus. Jour. Arnold Arb.
10: 169. 1929.

*Chromosomes and the evolution of Garden Philadelphus. Jour. Royal Hort.
Soc. or 269-275, 1951.

1954] HU, THE GENUS PHILADELPHUS 309

and the consequent hybridization of races widely separated geographically.”

Bangham worked on plants cultivated in Boston and Janaki Ammal’s
material was grown in London. These two cities are located out of the
natural ranges of the genus Philadelphus. The climate of these cities only
supports the outdoor growth of a few species and their forms. A close
examination of the lists of species studied by Bangham and Janaki Ammal
reveals that they covered very limited portions of the geographically widely
spread and morphologically much diversified genus. Thus it appears to
me that the cytological investigation of the natural species is not inclusive
enough for any cytologist to draw a conclusion as to whether differences
in the chromosome number of Philadelphus do exist in nature or not. A
large scale transplanting of natural species from southeastern and western
United States and Central America into gardens situated in areas warmer
than Boston or London, such as the United States National Arboretum in
Washington, D. C. or the Boyce Thompson Southwestern Arboretum for
the study of the drought resisting species, will doubtlessly provide better
opportunities for more exhaustive cytological studies of the genus and
advance our knowledge of the cytotaxonomy in this group.

GEOGRAPHIC DISTRIBUTION

The subgenus Gemmatus with its exposed axillary buds, simple, reduced
or ramiferous thyrse, high point of attachment in the placenta, broadened
usually cristate stigma and long-tailed seeds, represents morphologically,
I feel, the more primitive elements of the genus. The species are largely
subtropical or tropical in distribution. They are limited to the New
World, with eastern Mexico being its center of concentration. The range
extends ten degrees toward the north and the south of the Tropic of
Cancer. The effect of rainfall, temperature, altitude and latitude on the
distribution of the different species of this subgenus in Mexico is very
evident. Philadelphus karwinskyanus is found only on the Great Cross
Range and hence westward along the Pacific Coast at lower altitudes
where the climate is warmer. In the warm and humid section of the Sierra
Madre Oriental, one finds P. affinis with its subglabrous hypanthium. At
higher altitudes of the same range grows the small-flowered P. asperifolius.
Hence northward into the arid northeastern Mexico, occur various ‘en-
demic species belonging to the Section Coulterianus. In the neighbor-
hood of the Tropic of Cancer at high altitude where the rainfall is low,
P. maculatus, a small-leaved form with reddish purple spot at the base
of each petal, grows.

The true P. mexicanus with its solitary or ternate flowers and pubes-
cent petals occurs only on the Sierra Madre del Sur, the southern area
of the Sierra Madre Oriental, and in Chiapas, hence southwestward to
Guatemala. In southern Mexico and Guatemala, a species with pubescent
petals and ramiferous inflorescences, P. myrtoides, occurs. This species
extends southward to the Province of Chiriqui in Panama, the southern
limit of the range of distribution in the genus.

310 JOURNAL OF THE ARNOLD ARBORETUM [| VOL, XXXV

The subgenus Euphiladelphus includes geographically the more widely
distributed elements. Its representatives occur in Europe, Caucasus, the
Himalayan region, China Proper, Manchuria, Japan, a large part of the
United States, and adjacent southwestern Canada and northeastern Mex-
ico. It is an assemblage of species adapted to the climatic conditions of
the North Temperate Zone, between latitudes 25—50° N. in both the Old
and the New World.

In the Old World, the subgenus is rather stable. Radical changes did
not occur in the course of its evolution. All the Old World species belong
to one section, Stenostigma. In Europe, it is represented by P. coronarius
which occurs, according to Hegi, spontaneously in Romania, Austria and
southern Germany. In the Caucasus Mountains, it is represented by
P. caucasicus, a species which differs from P. coronarius only in its pubes-
cent discs and styles. The eastward distribution of the section is inter-
rupted in Central Asia. No species is met with until it reaches the
Himalayan Mountains. There the species concentrate in Sikkim, Nepal
and eastern Punjab where it is represented by P. tomentosus and its re-
lated forms. In China more marked morphological changes were involved
in the course of the evolution of this group. There the section can be
subdivided into several series. The Tsingling Range seems to provide a
central stage from where the different lines of evolution in the species can
be traced. At the western end of this mountain range occurs P. kansuen-
sis, a species having the leaf character of P. pekinensis, a North China
element, the pubescent disc of P. subcanus, a Central China element and
the pubescent hypanthium of P. Aenryi, a southwestern China element.
At the eastern end of this mountain range, occurs P. sericanthus and its
related forms. South of this range grows P. purpurascens, an interme-
diate form between the Tsingling and the Yunnan species, P. delavayi.
The latter species morphologically and geographically links the Chinese
and the Himalayan elements. North of the Tsingling Range in Shansi,
Shensi and Kansu occurs P. /axiflorus, a species morphologically and geo-
graphically intermediate between P. pekinensis and P. sericanthus. The
northernmost limit of the distribution of this section in the Old World is
the wooded valley of the Amur River and its tributaries, the Sungari and

P. schrenkti. These species also occur in Korea. In Japan this subgenus
is represented by P. satsumi and its related forms.

The distribution of Philadelphus in Eastern Asia presents a very in-
teresting phytogeographic phenomenon, that is, the complete absence of
the genus from the flora of Taiwan (Formosa). It is a well known fact
that Taiwan and Yunnan have very pronounced floristic affinities. Many
species as well as genera that occur in one province are also present in
the other. But in the case of Philadelphus, the genus is abundant in
northwestern Yunnan and is completely absent from Taiwan. This fact
may be taken as an indicative factor for confirming our belief as to the
origin of the genus and for explaining the pattern of its distribution in
Eastern Asia. The genera and species that are common to Taiwan and

1954 | HU. THE GENUS PHILADELPHUS 311

Yunnan are of Old World origin. Philadelphus is obviously of New World
origin, and its distribution in Eastern Asia seems to have been southward
in direction. Apparently neither the Japanese elements nor the continental
Chinese elements of this genus reached Taiwan in their migration.

In the New World, the subgenus Euphiladelphus is much more diver-
sified, and consequently three sections are represented, Two of these three
sections, Pauciflorus and Microphyllus, are endemic to the United States.
Few species of the section Microphyllus also occur at the northern border
of Mexico. The third section, Stenostigma, America shares with Europe
and Asia. In North America the species of the section Stenostigma can
be subdivided into two series the Gordoniani and the Pubescentes. Species
belonging to the Gordoniani series occur along the Snake and Columbia
Rivers, hence southward reaching the mountains of the California Coast
Range. Among the wild population, there seems to be two elements, P.
lewisii with a glabrous hypanthium and P. elleri with a slightly pubescent
hypanthium. Morphologically there seems to be little distinction between
them, other than the presence or absence of the hairs on the hypanthium,
but ecologically there are certain differences. In the natural population
of the general area considered, I have examined 292 individual collections
totaling 361 sheets. Of these, 177 have glabrous hypanthia and 115 have
slightly pubescent ones. After plotting the localities of these collections
on a map and then superposing this figure on Rehder’s map of climatic
zones,! a rather striking fact is revealed. Over 77% of P. lewisii occur
in zones where the average annual minimum temperature is — 10
— 5° F. while over 47% of P. helleri occur in zones where the average
annual minimum temperature falls to — 20° to 10° F. The checking
also indicates that the center of distribution of P. lewisii is in the lower
valley of the Columbia River and in the northern portion of the Pacific
Border Province while P. helleri occurs largely in the intermontane basin
and the deeply dissected mountain uplands of the northern Rocky Moun-
tains. Species belonging to the series Pubescentes occur in the Ozark
Plateau and the Arkansas Valley of the Interior Highlands and the Interior
Low Plateau region of Tennessee, Kentucky, and southern Ohio.

The section Pauciflorus with dichasial inflorescences, very large number
of stamens, enlarged stigmas, and high point of attachment in the placen-
tation and long-tailed seeds, retains all the primitive characters of the
genus. Species of this section concentrate in the mesophyllous forest of the
southern Appalachian region with the periphery of their range reaching
the Coastal Plain region along the borders of Georgia and Alabama in the
south. It seems that species of this section are rather inactive geograph-
ically and stable morphologically, for with all the material I have exam-
ined, there are very slight variations.

The section Microphyllus includes the xerophytic species. Its range
covers the southern Rocky Mountains in Central Colorado, the Colorado
Plateau in western Colorado, Utah, northern New Mexico and Northern
Arizona, the Sonoran Desert region in southern Nevada and southeastern

Man. Cult. Trees, Shrubs pp. xii—xiii, 1940.

Biz JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

California and the Mexican Highland in Eastern Arizona, southwestern
New Mexico and western Texas, and the adjoining Mexico. In this large
semidesert or desert area the populations are localized and isolated. There
are many endemics

The subgenus Deutzioides is strictly a North American taxon. Its three
sections occur in two different floristic provinces. The section Hirsutus,
as represented by P. hirsutus Nutt. and its related forms, concentrates in
the mesophyllous forests of the Southern Appalachian Mountain areas
with the Tennessee River and its tributaries being the center of its range.
The other two sections, Pseudoserpyllifolius and Serpyllifolius, are con-
stituted of xerophytic species which occur in southwestern Texas, New
Mexico, southern Arizona, and northern Mexico. It seems that the Mex-
ican Highland is a meeting ground of different elements, the subgenera
Euphiladelphus from the north, Deutzioides from the east and Gemmatus
from the south.

TAXONOMY

ae Linn. Sp. Pl. 470. 1753; et Gen. Pl. ed. 5, 211. no. 540.
— Miller, Dict. Gard. ed. 8, 834. 1768.— Willd. Sp. Pl. 2:
oe 1800. — Michx. Fl. Bor. re 1: 283. 1803.— Pursh, Fl. Am.
Sept. 1: 329. 1814. — DC. Prodr. 3: 205. 1828. — Schrader in Lin-
naea 12: 388. 1838. — G. Don, Gen. Syst. 2: 807. 1832. — Schlecht.
in Linnaea 13: 418. 1839. — Torr. & Gray, Fl. N. Am. 1: 594. 1840.
— Endl. Gen. Pl. 1187. 1841.— Walp. Rep. 2: 151. 1843. — Koch
in Woch. Gartn. Pflanzenk. 2: 229. 1859.— Rupr. ex Maxim. in
Bull. Phys.—Math. Acad. St. Pétersb. 15: 133. 1856; 15: 365.
1857. — Benth. & Hook. f., Gen. Pl. 1: 642. 1865.— Maxim. in
Mém. Acad, Sci. St. Pétersb, VII. 10(16): 35 (Rev. Hydrang. As.
Or.). 1867. — Koehne, Deutsche Dendr. 178. 1893; in Gartenfl. 45:
450. 1896; et in Mitt. Deutsche Dendr. Ges. 1904(13): 76. 1904. —
Rydb. in N. Am. FI. 22: 162. 1905. — Schneider, Il. Handb. Laubh.
1: 1905. — Syereistchikof, Il. Fl. Mosc. 2: 220. 1907. — Nakai in
Bot. Mag. Tokyo 29: 61. 1915.— Moore in Bailey, Stand. Cycl.
Hort. 5: 2579. 1916.— Rehd., Man. Cult. Trees Shrubs 270. 1927,
ed. 2, 264. 1940; et Bibliogr. Cult. Trees Shrubs 191. 1949. —
Engler, Pflanzenf. III, 2a: 69. fig. 36. A-G. 1891: ed. 2. 18a: 190,
fig. 110. A-G. 1930. — Hitchcock in Madrofio 7: 35. 1944, — Bean,
Trees Shrubs ed. 7, 2: 410. 1950; et in Chitt. Dict. Gard. 3: 1545.
1951.

Syringa Adanson, Fam. Pl. 2: 244. 1763. — Moench, Meth. Pl. 678. 1794, non
Linn. 1753
TYPE SPECIES: P. coronarius Linn.
Flowers solitary, ternate, in determinate racemes or in depauperate
panicles, generally fragrant. Calyx-tubes turbinate or subcampanulate,
adnate to the ovary forming the hypanthia, glabrous or pubescent, the

1954] HU, THE GENUS PHILADELPHUS 313

sepals 4, rarely 5, ovate, acute or acuminate; corolla white, rarely with
purplish center, the petals 4, rarely 5, in cultivated forms often doubled;
stamens 13-90, epigynous, the filaments subulate, free, rarely several
united at the base, the anthers oblong, rarely subglobose, glabrous, rarely
pilose; ovary inferior or semi-inferior, 4-, rarely in anomalous forms 5-
locular, the styles 4, rarely 3 or 5, entirely connate, partially free or rarely
in some cultivated forms entirely free, the stigmas free and linear, clavate,
oar-shaped, cristate, or coherent and columnar or subcapitate; placentas
auriculate, projected from the upper portion of the central axis; ovules
numerous, multiseriate, imbricate, pendulous, Capsules ellipsoid, tur-
binate, hemispherical or subglobose, corticate, the cortex chartaceous, the
pericarp cartilaginous, loculicidal. Seeds oblong-subcylindric, the testa
brown or nigrescent, membranous, reticulate, generally extending ante-
riorly into a fimbriate white crown and posteriorly into an obtuse or
acuminate tail, the funiculus nigrescent, persistent, the embryo minute,
embedded in the carnose and oily endosperm. Shrubs, erect, arching, sub-
scandent or rarely subspinescent; branchlets opposite. Leaves opposite,
deciduous, rarely evergreen, serrate or entire, triplinervate or quintupli-
nervate, glabrous or pubescent, the hairs simple, rarely evergreen, exstipu-
late; axillary buds exposed or enclosed.

The generic name, Philadelphus, is derived from the Greek root ¢:Aos
meaning love, SeAdos meaning brother. It was originally used by the
ancient Greeks, said to be named for Ptolemy Philadelphus, king of
Egypt, 283-247 B. C., for some plant the identity of which is now un-
known. Many pre-Linnaean authors called the plants which we now
ascribe as Philadelphus, “Syringa alba.” Clusius ' named his illustration
of the plant Frutex coronarius. Bauhin was the first man who interpreted
the pre-Linnaean “Syringa alba” as the Philadelphus of the Greeks."
When Linnaeus established the genus, he adopted Bauhin’s concept. The
vernacular names occurring in various literatures are enumerated in the
following list:

English: Syringa; common syringa; mock-oranges (for the
odor of the flower); pipe tree; pipe privet

French: Syringa odorant

German: Pfeifenstrauch; wilder jasmin; falscher jasmin

Dutch: Witte syring

Chinese: T’ai-ping-hua = flower of peace; san-mei-hua =
mountain mume flower; Mi-tsai = rice fuel.

Spanish: Geringuilla

Russian: Tschubuschnik ; pustoryl

Central American: Mosqueta; Acuilotl = water vine; cozticacuilotl;

cozticacuilotl xochitl; azahar

1 Rariorum Plantarum Historia 1: 55. 1601.
* Pinax Theatri Botanici 399. 1623.

314 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXxXVv

KEY TO THE SUBGENERA AND SECTIONS.

A. Axillary buds exposed.
. Seed long-caudate; stigmas free, broadened, usually cristate; flowers
solitary, ternate, or in depauperate panicles: fruit obovoid or subglo-
bose, with circumferential persistent calyx. Subg. I. GEMMATUS.

C. Leaves acuminate, rarely acute; flowers in a depauperate panicle,

- up to many-flowered, each on a pointed pedicel; the stigmas
broadened, cristate; fruits obovoid-ellipsoid

elongate but in general not cristate; fruit subglobose
pa RER GP PRESS EAs PEGA Pee ads Sect. 2. COULTERIANUS.

thyrsus; fruit ellipsoid with subapical persistent calyx.
bbs eee bas oo wees a Subg. III. MACROTHYRSUS
(Sect. 6 CALIFORNICUS )

BBB. Seeds ecaudate; stigmas connate, columnar or subcapitate; flowers
solitary or ternate; fruit turbinate or subglobose, with apical per-
sistent calyx... are Subg. IV. DEUTZIOIDES.

C. Leaves hirsute or strigose, all hairs straight.
D. Style 3-4 mm. long; subscandent shrubs; leaves serrate
ect ey YR ae Em & PATEP ALG a bas 4k Sect. 7. Hirsutus.
DD. Style 1 mm. long; subspinescent shrubs; leaves entire
tb peewee - Sect. 8. PSEUDOSERPYLLIFOLIUS.
CC. Leaves strigose and lanate beneath; style 1-2 mm. long; sub-
spinescent or small slender shrub... Sect. 9. SERPYLLIFOLIUS.

AA. Axillary buds enclosed . ; - Subg. I]. EUPHILADELPHUS.

B. Flowers solitary, ternate or in dichasial cymes.
C. Large arching shrubs; stamens 60-90: stigmas free, oar-shaped;
leaves 4-10 cm. long. a ..... Sect. 3. PAucIFLoRus.
CC. Erect low shrubs of compact growth or spinescent xerophytic
bushes; stamens 35-50: stigmas free or partially connate, linear;
leaves 0.5-2.5, rarely up to 3 cm. ong. . ere . 3
eet Tee ere - Sect. 5. MICROPHYLLUS.

BB. Flowers in determinate racemes __ _ Sect. 4. STENOSTIGMA.

Subgenus I. Gemmatus (Koehne), stat. nov.

Philadelphus subg. I. Gemmatus (Koehne), stat. nov.

Philadelphus Reihe 4. Decorticatae pauciflorae Koehne, Deutsche Dendr. 180,
184. 1893, pro parte.

Philadelphus sect. 1. Poecilostigma subsect. 1. Gemmati Koehne in Gartenfl.
45: 450. 1896, pro parte; et in Mitt. Deutsche Denar. Ges. 1904(13): 77.
1904. — Engler, Pflanzenf. ed. 2, 18a: 191. 1930.

TYPE SPECIES: P. mexicanus Schlechtendal.

Flowers solitary, ternate, or in depauperate panicles; hypanthia ob-
conic, or subglobose, generally dense pubescent: stamens about 40, rarely
more or less; ovary semi-inferior, the stigma distinct, broadened or cris-

1954] HU, THE GENUS PHILADELPHUS 9

tate; capsules obovoid-ellipsoid or subglobose, the persistent calyx cir-
cumferential or subcircumferential; seeds long-caudate, the crowns with
pointed lobes, erect or subscandent shrubs or vines; leaves evergreen or
deciduous, remotely denticulate or serrate, triplinervate or quintuplinerv-
ate; petioles often twisted; axillary buds exposed.

This subgenus is strictly a Central American taxon. In the southern
portion of its range there are several widely spreading species, and in the
northern portion, the species are largely endemic to limited areas. It
contains two sections including fourteen species. Their geographical
distributions are as illustrated in Map 1.

Section 1. POECILOSTIGMA Koehne

Philadelphus subg. I. Gemmatus sect. 1. Poecilostigma Koehne in
Gartenfl. 45: 450. 1896: et in Mitt. Deutsche Dendr. Ges. 1904(13):
77. 1904, pro parte.
Philadelphus sect. Poecilostigma Koehne, ll.cc. — Engler, Pflanzenf. ed.
2, 18a: 191. 1930.

Philadelphus Mexicani Rydb. in N. Am. Fl. 22: 164. 1905, in clavis, s. stat.

rae a ser. Gemmati (Koehne) Rehder, Man. Cult. Trees Shrubs ed.
1940; et Bibliogr. Cult. Trees Shrubs 194. 1949, pro parte.

Type spEciIES: P. mexicanus Schlechtendal.

Arching shrubs or vines, the current year’s growth pubescent and usu-
ally verrucose, the axillary buds exposed; leaves remotely denticulate or
serrate, the base rounded, 3- or 5-nerved, the apex acuminate, rarely acute;
inflorescences depauperate panicles, or flowers solitary, the bracts linear
or lanceolate, foliaceous; hypanthia subglobose or cyathiform, the sepals
foliaceous, (5—) 8-20 mm. long; corolla usually disciform, often pubes-
cent; stamens 40-50; disc conic, style (3—) 4-7 mm. long, the stigma
broadened and cristate; fruit obovoid-ellipsoid; seeds long-caudate. Seven
species, Mexico to Panama.

KEY TO THE SPECIES

A. Flowers many on ramified branches; hypanthia subglobos
B. Inflorescences loose, paniculate, the secondary axis _ unequal length,
4-30 mm. long, the lower ones longer
C. Hypanthia glabrous or with few weak pilose hairs; oo of the hairs
on the stem not thickened; axillary buds subglobos

eee enn eee ET eae eo ee ee a Pe affinis.

Cc, Hypanthia pubescent; base of the hairs on the ‘stem thickened;
conic.

D. Hypanthia lanate, the underneath tissue Nanay: inside of the

petals glabrous P. karwinskyanus.

DD. Hypanthia villose. ‘the underneath ae Ena petals

pubescent on the apical ends 3: P. pueblanus.

BB. hie compact, raceme-like, the “secondary axis equal
length, 2-4 mm. long, the flowers crowded at the end of elongated
leafy branches . buestyak ha dam aoh MSMR 25 $6 48 dans 4. P. myrtoides.

316 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

AA. Flowers solitary or ternate; hypanthia cyathiform or obconic (except P.
glabripetalus )
B. Petals pubescent on both surfaces; hypanthia sparsely villose or pilose.
C. a and disc pubescent; ei ry buds conic; leaves large, 5-11
m. long, the petioles 8-10 mm. long | 5. P. mexicanus.
cc. “Style and disc glabrous; axillary buds aia leaves small,
1-3.5 cm. long. the petioles 2-3 mm. lon
SebeesaWhaekane gaedh ears e245 ‘6. P. austro-mexicanus.
BB. Petals nan at least on the inside: hypanthia lanate
1 #: glabripetalus.

Sect POECILOSTIONA.- — Seek. COULTERIANUS..-...
AR affinis 1P asperifolius

BP. austro-mexicanas t P. caleico/us

o PF glabripefalas ¢ P coulters

a P karwinskyanus —& =P. oblongifolius
e P mexicanus 6a PR osmanthus
4 PR myrtoides x PR pringks

+ P. pueblanus SP. sargentianus

Map 1. Geographical distribution of the species and sections of Philadelphus
in the subgenus Gemmatus

1. Philadelphus affinis Schlecht. in Linnaea 13: ia 1839. — Walp.
Rep. 2: 151. 1843.— Hemsl. in Biol. Centr. Am. 1: 383. 1879. —
Koehne in Gartenfl. 45: 487. 1896; et in Mitt. ace Dendr. Ges.
1904(13): 78. 1904. — Schneider, Il]. Handb. Laubh. 1: 362. 1905.
— Rydb. in N. Am. FI. 22: 171. 1905. — Standl. in Contr. U. S. Nat.
Herb. 23: 511. 1922. — Engler, Pflanzenf. ed. 2. 18a: 191. 1930.

Philadelphus zeyhert sensu Hemsl. in Biol. Centr. Am. 1: 384. 1880, non
Schrader ex DC. 1828

1954] HU, THE GENUS PHILADELPHUS 317
Type: Ehrenberg, Barranca de la Hacienda Del Carmen, S. Mexico.
An arching shrub or vine up to 4 meters high, the flowering branches

slender, 10-20 cm. long, bearing 3 to 5 pairs of leaves, sparsely pilose, the

base of the hairs not thickened; axillary buds subglobose, pubescent.

Leaves ovate rarely ovate-elliptic, 4.5-8 cm. long, 2.5-4.5 cm. wide,

rounded, rarely subcordate at the base, 5-nerved, acuminate at the apex,

remotely serrate, each side with 2 to 5 teeth, subglobose or very sparsely
strigose on both surfaces, more so on the principal nerves beneath, reticu-
lations obscure above, conspicuous beneath, petioles 3-10 mm. long,
pilose. Inflorescences depauperated panicles (Pl. V fig. 1) with 5-, rarely

3-, 1-, or 11-flowers, the woody part of the stalk 1-3 cm. long, pubescent

as are the branchlets, bracts linear, the pedicels 2-3 mm. long, glabrous,

after fruit up to 6 mm. long; hypanthia subglobose, 6 mm. in diameter,

glabrous or with few weak pilose hairs, the sepals ovate, acuminate, 10

mm. long, 4-5 mm. wide, glabrous or very sparsely pilose; corolla sub-

disciform, 3.5—5 cm. across, the petals obovate to suborbicular, 1.3—-1.8

cm. in diameter, glabrous, the apex round; stamens about 50, the longest

three-fourths the length of the petals; disc subconic, glabrous; style 7 mm.

long, undivided or the upper one-fifth separated, hirsute, the stigmas

broadened, cristate, the abaxial surfaces broader and longer than the
adaxial ones, the sterile portion hirsute (Pl. V fig. 1a). Capsules ellipsoid,
the persistent calyx circumferential, the seeds long-tailed.

MEXICO: ae Zacualtipan, P. Maury 5826 (NY), same locality,
H. E. Moore 3238 (BH, G); in a barranca below Trinidad Iron Works, C. G.
Pringle 8833 (F, G, MO. NY, US); Atotonilco, Schiede, June 1830 (A, US);
Bluff above Cuera Humada, A. J. Sharp 4618 (TENN.). Tamaulipas:
Tampico, Real del Monte, M. Berlandier 333 (F, MO, US). Without precise
locality, Herb. Dendrol. Schneider (A).

The type not being available, this interpretation is made on the basis
of Schlechtendal’s original description and Koehne’s supplementary re-
marks. Schlechtendal characterized the inflorescences as 5-flowered ra-
cemes, the peduncles and calyces smooth and glabrous. Unfortunately he
did not mention the pubescence of the style. For this character I rely
upon Koehne who certainly had access to the now destroyed type. Ac-
cording to Koehne the style of this species has loose stiff hairs. I therefore
place the Mexican material with 5-flowered depauperate panicles, more or
less glabrous hypanthia and calyces, and pubescent styles with P. affinis
Schlecht. Hemsley in 1880 doubtfully published Berlandier 333 from
Tampico as P. zeyheri Schrad. In this he mistook the material of a spon-
taneous Mexican species, P. affinis Schlecht., for a garden form cultivated
in European gardens.

The ovate leaves, the subglobose glabrous hypanthia, the large foli-
aceous sepals with acuminate apices, the large number of stamens, the
broadened stigmas and the ellipsoid capsules with circumferential persist-
ent sepals of this species all suggest relationship to P. inodorus Linn. of
the southern Appalachian region of the southeastern United States. As

318 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

the latter species has enclosed buds and this one has exposed buds their
separation must have taken place far back in geological time.

In Mexico this species occurs on the Sierra Madre Oriental at altitudes
of 1852-2000 meters, in regions where the annual rainfall attains at least
80 inches per year. Its white fragrant flowers appear from April to June.
The specimens collected in July are with fruits.

2. Philadelphus karwinskyanus Koehne in Gartenfl. 45: 486. 1896;
et in Mitt. Deutsch. Dendr. Ges. 1904(13): 1904. — Schneider,
Ill. Handb. Laubh. 1: 362. 1905.— Rydb. in N. Am. FI. 22: 170.
1905. — Standley in Contr. U.S. Nat. Herb. 23: 311. 1922.

Philadelphus scandens Moore in Bailey, Stand. Cycl. Hort. 5: 2582. 1916.

Type: Karwinsky, Tolotapa, Oaxaca, Mexico.

A subscandent shrub up to 4 meters high, branchlets slender, the flower-
ing ones over 50 cm. long, much ramified, terete, striate, villose, the base
of the hairs thickened; axillary buds conic, villose. Leaves ovate, 4—7.5
cm. long, 2—3.5 cm. wide, rounded at the base, acuminate at the apex,
5-nerved, reticulations conspicuous beneath, remotely sharp serrate, 5—8
teeth on each side, sparsely strigose on both surfaces; petiole 5-10 mm.
long, attached on cushion-like projections of the node. Inflorescences with
5-31 flowers, in a loose depauperate panicle, the woody portion below the
pedicel 4-20 mm. long, pubescent, the pedicels 2—5 mm. long, thickly
strigose and lanate; hypanthia subglobose, 4 mm. in diameter, strigose-
lanate, the sepals ovate, 5-8 mm. long, densely lanate, the hairs entirely
obscuring the tissue beneath; corolla 2.5—3 cm. across, the petals obovate-
suborbicular 0.8—13 mm. in diameter, sparsely pubescent along the me-
dian dorsal line, glabrous on the ventral side, the apex rounded; stamens
about 45, the longest half as long as the petals; disc subconic, pubescent,
stigmas 2.5-3 mm. long, enlarged cristate, the sterile portion pubescent, the
abaxial surface with 2 papillose ridges. Capsule and seed not known.

MEXICO: Oaxaca: Tepascolula, Loesener 1421(G); Huitzo, L. C. Smith
807(G). Vera Cruz: Orizaba, F. Miiller, Jan. 1853(NY). Sinaloa:
Mazatlan, J. G. Ortega 6770(F). Without precise locality, Ortega 7342(F).

UNITED STATES: Cultivated in California: Altadena, F. W.
Peirson 79(BH); San Diego, F. G. Woodcock 870(F), 871(BH), 872(A,
BH); Santa Monica, L. H. & Ethel Bailey 7804(BH); Los Angeles, L. H. &
Ethel Bailey 9117( BH).

A great altitudinal variation is reported for this species. In Oaxaca it
has been collected 2000 meters above the sea level and in Sinaloa, it has
been reported from humid lowlands almost at sea level. It is an evergreen
shrub, flowering all year around. In California where it is an introduced
and cultivated species, flowering records cover August, October, December,
January, February, March, April and May.

1954] HU, THE GENUS PHILADELPHUS 319
3. Philadelphus pueblanus, sp. nov.

Frutex subscandens, usque 4 m. altus, ramulis floriferis, 20-40 cm.
longis, teretibus, striatis, 2-4 mm. diametro, dense pilosis et sparse stri-
gosis, pilis basi incrassatis, gemmis axillaribus breviter, villosis; foliis
ovatis, 4-7 cm. longis, 2—3.5 cm. latis, basi rotundatis, raro obtusis, quin-
tuplinerviis, apice acuminatis, serratis vel subintegris, dentibus utrinque
1, 2 usque ad 5, utrinque strigosis, petiolo 5-10 mm. longo; inflorescentiis
ramosis, 5- usque 31-floribus, pedunculis 4-20 mm. longis, pedicellis 3—6
mm. longis, villosis, verruculosis, hypanthiis subglobosis, 4 mm. diametro,
villosis; sepalis ovatis, caudatis, 10 mm. longis, 4-5 mm. latis, sparse
villosis; corolla disciformii, 4 cm. diametro, petalis oblongo-suborbicu-
latis, 2 cm. longis, 1.5 cm. latis, partibus medianis extus hirsutis, apice
rotundatis et retusis; staminibus ca. 40; disco subconico, medio hirsuto;
stylis 6-7 mm. longis, columnaribus indivisis; pubescentibus; stigmatibus
cristatis, 2-3 mm. longis, partibus sterilibus hirsutis; capsulis et seminibus
ignotis.

MEXICO: Puebla: vicinity of Puebla, Bro. G. Arséne (Bro. Nicolas)
171 (A, TyPE; G, MO, NY, US isotypes); aoe Bro. G. Arséne
10136(US); Ixtaccihuatl, C. A. Purpus 169(MO, US); Com. Geogr.-Expl.
Rep. WMewcans 102Z20NY) = Evexcala: Tlaxcala, E. A, Balls 4906(A).
SanLuis Potosi: Santa Barbara, Bro. G. Arséne 10093 (US). Mexico:
Par Nicolas, M. Bourgeau 995(US); Temascaltepec, Rincon, G. B. Hinton
2362(A), 5032(BH, NY, US). Michoacan: Zitacuaro, G. B. Hinton
13342(G, NY, US). Morelos: near oe ae J. N. Rose & W. Hough
4412(US).

This species is intermediate between P. affinis Schlecht. which has
glabrous or subglabrous hypanthium and P. karwinskyanus Koehne in
which the hypanthium is thickly lanate with the indumentum entirely
covering the epidermal tissue. In Puebla it occurs at altitudes between
1700 and 2194 meters as semiscandent shrubs in thickets and among trees
in ravines. Its pale yellow sweet-scented flowers appear from March to
October.

—Hemsl. Biol. Centr. Am. 1: 383. 1879. —
ae in Gartenfl. 45: 487. 1896. — Rydb. in N. Am. FI. 22: 170.
1905.— Engler, Pflanzenf. ed. 2, 18a: 192. 1930.— Standley in
Field Mus. Nat. Hist. Bot. Ser. 18: 474 (Fl. Costa Rica). 1937.

eter trichopetalus Kornicke in Gartenfl. 16: 73. 1867.— Koehne in
45: 487. 1869; et in Mitt. Deutsch. Dendr. Ges. 1904(13): 78.
cea in Pittier, Prim. Fl. Cost. II. 1: 90. 1898. — Schneider, III.
Handb. Laubh. 1: 364. 1905.— Engler, Pflanzenfam. ed. 2, 18a: 192. 1930.

4. ace myrtoides Bertol. Fl. Guatim. 21. pl. 7. 1840. — Walp.
Rep. 151. 1843.

ee mexicanus sensu Moore in Bailey, Stand. Cycl. Hort. 5: 2582.
6, pro parte. — Standley & Calderon, Pl. Salv. 88. 1925, non Schlech-
at

320 JOURNAL OF THE ARNOLD ARBORETUM [| VOL, XXXV

Philadelphus sempervirens Hort. ex. Moore in Bailey, Stand. Cycl. Hort. 5:
2582. 1916
Philadelphus matudai Lundell in Contr. Univ. Mich. Herb. 4: 6. 1940.

Type: Botanical Garden of Bologna, Italy.

An arching shrub, 2—4 m. high, flowering branches slender, 21-50 cm.
long, with 14 or more pairs of leaves, longitudinally striate, rather densely
villose, the trichomes slightly enrled. their base more or less thickened;
buds small, rounded, densely brown villose. Leaves ovate, 3-4 cm., rarely
up to 5.5 cm. long, 1-2.8 cm. wide, sparsely strigose above, more so
beneath, the hairs glandular at the base, rounded or rarely obtuse at the
base, acuminate at the apex, subentire, or remotely denticulate, with 2 to
6 sharp teeth projecting outward on each side, triplinervate or quintupli-
nervate, reticulations obscure on both surfaces; petioles 6-9 mm. long,
strigose. Flowers 5, rarely 1, 3, or 7, crowned at the end of long slender
branchlets, the woody portions below the pedicels uniformly short, 2—4
mm. long, the bracts linear, 5-13 mm. long, the pedicels 3-4 mm. long,
white lanate; hypanthia subglobose, 5 mm. in diameter, densely lanate:
sepals ovate, acuminate, 8 mm. long, 4 mm. wide, lanate; corolla disciform,
3—4 cm. in diameter, the petals suborbicular-ovate, 1.2-1.8 cm. long,
1.1-1.7 cm. wide, slightly hirsute along the median longitudinal line on
both surfaces, the apex rounded; stamens ca. 44, the longest half as long
as the petals; disc subconic, pubescent, the style 3-4 mm. long, pubescent,
the stigmas 3 mm. long, enlarged, cristate, the abaxial surface as long as
the adaxial, the sterile portions hirsute; capsules and seeds not known.

MEXICO: Chiapas: V —o Tacana, E. Matuda 2791 (1sotyee of
P. matudai Lundell, A, F, G. NY, U

GUATEMALA: Alta Vera Paz: Vicinity of Coban, P. C. Standley
92452(F). Guatemala: near Finca La Aurora, Ignacio Aguilar 82(F).
Huehuetenango: near Chiantla, P. C. Standley 82517(F). Sacate-
péquez: near Antigua, P. C. Standley 60316(F); S. Hayes (G). Quezal-
tenango: P. C, Standley 86561(F, US). Suchitepéquez: vicinity of
Finca Alvidas, J. A. Steyermark 35458(F).

EL Sag ape ten San Salvador: S. Calderén 687 (F, G, US, MO,
NY); M. C. Carlson 435(F). Ahuachapan: near Ataco, P. C. Standley
o VE. ep 2726(F).

HONDURAS: M orazan: Tegucigalpa, J. V. Rodrigues 3173(F).

COSTA RICA: Cartago, Cervantes, A. Tonduz 10444(US):; San José.
Tonduz 1492(US); Las Céncavas, P. C. Standley 36006(US); Tapisca de
Zaruro, Austin Smith 185(US).

PANAMA: Chiriqui: vicinity of Bajo Mona and Quebrada Chiquero,
R. E. Woodson & R.W. Schery 588(MO).

CULTIVATED: California: Santa Barbara. E. P. Bradbury, Oct. 10,
1915(BH)

The identification of this species is based on Bertoloni’s description and
illustration. In his illustration he characterized P. myrtoides as a plant

1954] HU, THE GENUS PHILADELPHUS 321

with rather small ovate-elliptic leaves on the flowering branches which
terminate with crowded cluster of several flowers each in the axil of a
reduced leaf. Among the available collections of Guatemalan Philadel-
phus, Bertoloni’s description and illustration conforms in all respects to
Standley 60316, 82417 and Steyermark 35458. It is also well represented
by Calderén 687 from El Salvador, Matuda 2791 represents the northern
limit of the range of the species. Bradbury’s collection from Santa
Barbara, in Bailey Hortorium, was identified as P. sempervirens Hort.
This specimen was probably Moore’s material basis for the publication of
his P. sempervirens.

Although 110 years have elapsed since P. myrtoides was published, it
has not been well understood. K6rnicke was probably not aware of
Bertoloni’s species when he published P. trichopetalus from Costa Rica.
According to his description the Costa Rica plant has flowers clustered
at the end of a branchlet, and the petals of the flowers are softly pilose.
I have not seen Kornicke’s type. But judging from the Costa Rica and
Panama collections I have examined and by our present knowledge of the
distribution of the various species of Philadelphus in Central America,
P. trichopetalus Kornicke and P. myrtoides Bertol. are conspecific.

Philadelphus myrtoides is the most southern species of the genus. It
has been collected on the high mountains from Chiapas in Mexico through
Guatemala, El Salvador, Honduras south to Costa Rica and Panama. In
Guatemala it occurs in cypress groves at altitudes ranging from 1200 to
1930 meters. Standley suggested the possibility of it being introduced
and cultivated in Costa Rica. Some of the El] Salvador collections were
from gardens. Apparently this species has been widely cultivated in
Central America where it flowers from January to August. The creamy
white fragrant flowers are sold in the markets for decorative purposes
under the name ‘“‘Mosqueta.”

Philadelphus myrtoides is closely related to P. karwinskyanus Koehne
but the latter species can be distinguished fer its elongated peduncles and
its petals being glabrous on the inner surface.

wn

Philadelphus mexicanus Schlecht. in Linnaea 13: 418. 1839. —
Walp. Rep. 2: 151. 1843.— Hemsley in Biol. Centr. Am. 1: 384.
1879, excl. spec. Seemann, Coulter, Ghiesbreght. —W. G. Smith in

: 753, fig. 123. 1883.— Burbidge in Gard.
Chron. IIT. 34: 218, fig. 89. 1903. — Hook. f. in Bot. Mag. 124: pl.
7600. 1898.— Koehne in Gartenfl. 45: 487. 1896; et in Mitt.
Deutsch. Ges. 1904(13): 78. 1904.— Wittmack in Berl. Gartenz.
1883: 528. fig. 91. 1883. — Dippel, Handb. Laubh. 3: 335. 1893. —
Schneider, Ill. Handb. Laubh. 1: 362, fig. 234 b-b.? 1905. — Rydb.
in N. Am. FI. 22: 170. 1905. — Standley in Contr. U. S. Nat. Herb.
23: 311. 1922.— Rehder, Man. Cult. Trees Shrubs 280. 1927, ed.
2, 275. 1940; et Bibliogr. Cult. Trees Shrubs 195. 1929. — Hansell,
Mexico Pl. 162. 1935.— Bean, Trees Shrubs ed. 7, 2: 419. 1950;
et in Chitt. Dict. Gard. 3: 1546. 1951.

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322 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

A scandent evergreen shrub to 5 meters high with long drooping
branches, the second year’s growth 3-4 mm. in diameter, castaneous,
rugose, the bark closed, longitudinally rimulose, the current year’s growth
1.5—-2 mm. in diameter, hirsute, the trichomes with more or less thickened
bases; axillary buds conic, the first few below the flowers often developing
into virgin shoots. Leaves ovate, 5-11.5 cm. long, 2-5 cm. wide, very
sparsely strigose on both surfaces, rounded or obtuse or subcordate at the
base, 5- or rarely 3-nerved, acuminate at the apex, subentire or with 1 to
6 tooth-like projections on each side, reticulations obscure above, con-
spicuous beneath, the petioles 8-10 mm. long, strigose. Flowers solitary,
yellowish white, very fragrant; pedicels hirsute, 1-3 mm. long, bracts
lanceolate, 1-2 cm. long; hypanthia cyathiform, sparsely villose, the
trichomes slightly curly; sepals foliaceous, ovate, long acuminate or
cordate, 10-20 mm. long, 7 mm. wide, sparsely villose, the hairs tending
to curl slightly; corolla 3—4 cm. across, the petals suborbicular, hirtellous
on both surfaces; stamens ca. 40, the anthers oblong; disc subconic,
hirsute at the center, the styles 3 mm. long, pubescent, the upper half
divided, the stigmas 3 mm. long, cristate, the abaxial surface much longer
than the adaxial one, the sterile portion often hairy. Capsules obovoid-
ellipsoid, the lower portion near the base quadrangular, about 12 mm.
long, 10 mm. in diameter, the persistent calyx subcircumferential. Seeds
long-caudate.

MEXICO: Mexico: Amecameca, G. L. Fisher on July 29, 1924(F, MO,
US); Federal District, A. J. Sharp 445 (A,TENN); Mts. of Toluca, Halstead
(NY). Guerrero: Omiltemé, A. J. Sharp 441551 (A, TENN). Mich-

Cerro de San Felipe, C. Conzatti & V. Gonzalez 471(G); H. Galeotti
2850(US); E. W. Nelson 1398(G,US). Chiapas: Siltepec, E. Matuda
1701(A,MO,US). Vera Cruz: Jalapa, S. Schiede(G.NY, ISOTYPES) ; Orizaba,
Botteri 1105(G, US). Puebla: Cero Guadelupe, Nicolas, June 10, 1909(F).

GUATEMALA: Huehuetenango: San Juan Ixcoy, A. J. Sharp
4616(F); definite locality not given, J. D. Smith, April 9, 1896(G):; Cerro
Canana, Sierra de los a ep J. Az Steyermark 49059(F); Aguacatan,
F.. G, ‘Standley 91218(F). Quezaltenango: Volcan de Santa Maria,
P. C. Standley 83535(F); J. A. Pipa 33985(F); Palestina, P. C. Stand-
ley 84230(F); Santa Maria de Jesus, P. C. Standley 84860(F); above Mujuli,
P. C. Standley 85608(F, US); Zunil, P. C. pa: 85647(F); Volcan Santo
Tomas, J. A. Steyermark 34952 (F). Quic : O. F. Cook 31(US); San
Miquel Uspantan, Heyde & Lux (distributed se J. D. Smith) 3046(G, NY,
US); Chajul, A. J. Sharp 4681(F). Sacatepequez: Volcan de Auga,
P. C. Standley 65124(F). Chimaltenango: Santa Elena, D. S. Johnson,
Aug. 10, 1932(F); A. F. Skutch 335 (A, US). SololaA: Volcan San Pedro,
J. A, Steyermark 47253(F, US); Volcan Atitlan, J. A. Steyermark 47527(F).
San Marcos: Volcan Tajumulco, J. A. Steyermark 35661(F). Without
precise locality, F. C. Lehmann, June 1882(US).

CULTIVATED: California: Berkeley, in the garden of A. Blake, N. F.
Bracelin, 1354( BH)

1954] HU, THE GENUS PHILADELPHUS 323

As the natives of Mexico had cultivated species of Philadelphus from
early times it is inevitable that certain garden forms had been produced
by them. It happened that the first published Mexican species was based
on a double flowered form. Schlechtendal cited three specimens, Schiede
from Jalapa, Mihlenpfordt from Oaxaca and a specimen from Mexico
in Herb. Lehmann. Of these I have seen only the duplicates of Schiede’s
collection, which agree with Schlechtendal’s description. Thus the double
flowered form typifies the species. Apparently this form is found only or
chiefly in the states of Vera Cruz and Puebla where the mean annual
temperature is between 62—67° F. and the mean annual rainfall is 40-80
inches. The Bracelin collection is the only specimen from a cultivated
plant outside of Mexico that I have seen. It has been reported that the
cultivated plants climb to 12 meters and their flowers are yellowish or
deep cream white.

Specimens collected from the wild population all have simple flowers.
Compared with the isotypes of typical P. mexicanus Schlecht., I can
detect no other characters from the above cited specimens to distinguish
them as a form besides their possession of the simple flowers. Like the
double-flowered form, their petals are pubescent on both surfaces, the
disc and style are pubescent, and the hypanthium is sparsely villose with
the trichomes slightly curled. In general, their leaves are larger and more
prominently quintuplinervate than the double flowered form. But this
5-nerved condition is also true with the large leaves of the cultivated
specimens. It seems that with both the wild and the cultivated material,
at the base of each leaf, there is a pair of small nerves originated from the
petiole and running along the basal portion of the margin. When the leaves
are small, these nerves are so closely pressed against the margin that they
appear inconspicuous.

In Mexico this species occurs at altitudes of 2280-3000 meters in
Oaxaca. In Huehuetenango, it has been recorded at altitudes of 1950—
3160 meters on steep limestone slopes of damp oak forests. In Quezal-
tenango it occurs at altitudes of 1500-3000 meters in sunny thickets,
damp sandy hillside forests or in wet brushy ravines as subscandent
shrubs. In Solola it occurs in damp cloud forests.

The vernacular names reported are “Mosqueta” and “Azahar,” the
former more widely applied. Hernandez (1651) in “Nova Plantarum
Mexicanorum Historia” discussed and illustrated a plant with opposite
leaves and a cluster of two simple flowers under the name, Acuilotl or
Volubili Aquatica. He mentioned it as growing in wet places, creeping
on the ground or scrambling up trees. This record has been regarded by
Schlechtendal, Lindley, Hooker and many modern authors as represent-
ing P. mexicanus Schlecht. To the Mexican Indians this species is eco-
nomically rather important. They employ the flowers both in preparing
perfume and for making garlands. The leaves are taken in wine for the
relief of colic, or crushed and applied as a plaster to ease strained members
and dissolve tumors.

The pubescent petals and simple flowers of this species resemble those

324 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

of the flowers of P. myrtoides Bertol. But the latter species can be readily
distinguished by its clustered flowers and densely lanate hypanthium.

6. Philadelphus austro-mexicanus, sp. nov.

Philadelphus mexicanus sensu Decaisne in Rev. Hort. III. 1: 381. fig. 20.
1852. non Schlechtendal. 1839.

Frutex ramosissimus, ramulis floriferis brevibus, ca. 3 cm. longis. 1 mm.
diametro, castaneis, rugoso- strigosis, pilis basi incrassatis, gemmis axil-
laribus subglobosis, pilosis; foliis ovatis vel ovato-lanceolatis, 1-3.5 cm
longis, 0.5—1.5 cm. latis, basi subrotundatis, triplinerviis, apice subacumi-
natis vel acutis, serratis, utrinque 1- usque 5-serratis utrinque sparse
strigosis vel praeter nerviis subglabris, petiolo 2-3 mm. longo: floribus
solitariis, pedicellis 2 mm. longis, strigosis; hypanthiis obconicis, 3 mm.
longis, 4 mm. diametro, sparse pilosis; sepalis ovatis, 9-11 mm. longis,
caudatis, sparse pilosis vel subglabris; corolla disciformi, 3.8-4 cm. di-
ametro, petalis obovatis, 1.5—-1.7 mm. longis, 1.3 mm. latis, utrinque in
partibus medius hirsutis: staminibus ca. 52, disco glabro, stylis 4 mm.
longis, glabris, partibus superioribus liberis, stigmatibus dilatatis, cris-
tatis; capsulis et seminibus ignotis.

MEXICO: Chiapas: Dr. Ghiesbreght 813 (G. type; F, MO, 1sotypEs;
A. fragment of ISOTYPE).

The rugose branchlets, the dentation of the leaves, the large sepals, the
pubescent petals, the numerous stamens, the divided style and the dilate-
cristate stigmas of this species all suggest a close relationship to P. mexi-
canus Schlecht. But it can be readily distinguished by its small leaves,
very sparsely pilose hypanthium and sepals and its glabrous styles.

Decaisne’s figure represents a plant with small leaves. His concept of
Schlechtendal’s species was apparently based on a Ghiesbreght collection,
very likely a part of the same collection of which three specimens are
available to me. All of them have small leaves and glabrous styles.

7. Philadelphus glabripetalus, sp. nov.
Philadelphus mexicanus sensu Lindl. Bot. Reg. een 37. 1840; et 28:
; .— sensu Hemsl. in Biol. Centr. Am. 1: 384. 1879, pro parte,
non Schlechtendal, 1839.

Frutex ramosissimus, ramulis floriferis 5-15 cm. longis, 1-1.5 mm.
diametro, rugoso-hirsutis, pilis basi incrassatis: foliis ovatis vel ovato-
lanceolatis, 1.5—6 cm. longis, 1—2.5 cm. latis, basi rotundatis. triplinerviis,
apice acuminatis, subintegris vel serratis, dentibus utrinque 1 vel 2, raro
4. supra subglabris vel sparse villosis, subtus sparse villosis: petiolo 3—5
mm. longo, villoso; floribus solitariis vel raro ternatis, pedicellis 3—5 mm.
longis, dense villosis, hypanthiis subglobosis, 5 mm. diametro, lanatis:
sepalis ovatis, 10-15 mm. longis, 5 mm. latis, acuminatis: corolla 2.5—3
cm. diametro, petalis obovatis, vel suborbicularibus, 1-1.3 cm. longis,
9-10 mm. Latics utrinque glabris, vel extus hirsutis, staminibus ca. 60;

1954] HU, THE GENUS PHILADELPHUS 325

disco medio hirsuto, stylis glabris vel basi hirsutis, 3-4 mm. longis, stig-
matibus cristatis, 4 mm. longis, supra liberis; capsulis et seminibus ignotis.

MEXICO: Federal District: Sierra de Ajusco, C. G. Pringle 6311(A,
type; G, MO, NY, US, tsotypes). Puebla: Moria, Nicolas, in February
1908(F). Without precise locality, P. Maury 3158(NY), 3822(NY).

The shape of the leaves, the solitary flowers and the glabrous petals
suggest a relationship to P. osmanthus S. Y. Hu, but the latter species
has strigose lower leaf-surfaces and hypanthia, and short styies while the
present species has villose lower leaf-surfaces, lanate hypanthium, and
styles 3-4 mm. long. It differs from P. mexicanus Schlecht. in having
glabrous petals and smaller leaves. Its solitary or ternate flowers also
suggest relationship with P. coulteri S. Wats. but the latter species has
densely villose hypanthia with the epidermis entirely obscured. In the
Federal District, this species occurs at altitudes 2400-2470 meters. It
climbs among shrubs up to 5—7 meters high. Specimens collected from
June to September all bear flowers. This suggests that like many other
Mexican Philadelphus, this species has a very long flowering period; of
course, local climatic conditions may be involved.

Judging by the figures and the descriptions given by Lindley (1842),
his material which was introduced by Hartweg from the Hacienda del
Carmen, could not be the true P. mexicanus Schlecht. for it has small
leaves and a glabrous style while Schlechtendal’s species has large leaves
and a pubescent style. As Lindley described the hypanthium and calyx
of his material as densely pubescent, this eliminates the possibility of it
being P. austro-mexicanus S. Y. Hu which is another small-leaved Mex-
ican species, for the latter has more or less glabrous hypanthium and
calyx, and pubescent style. By the process of elimination, one may log-
ically come to the conclusion that the material Hartweg introduced to
the European gardens belongs here.

Section 2. Coulterianus, sect. nov.
Philadelphus subg. I. Gemmatus sect. Coulterianus, sect. nov.

Type sPEcIEs: P. coulteri Wats.

Frutex erectus vel subscandens, ramulosis hornotinis pubescentibus,
plerumque verrucosis, gemmis axillaribus expositis; foliis integris, raro
denticulatis vel inconspicue serratis, basi rotundatis vel obtusis, raro
acutis, triplinerviis; apice acutis vel obtusis; floribus solitariis vel raro ter-
natis; hypanthiis subglobosis, pubescentibus; sepalis ovatis, 3-8 mm.
longis, corolla cruciformi vel disciformi; staminibus 30 usque ad 44; disco
plano, stylo 1-3 mm. raro 4 mm. longo, stigmatibus noncristatis; capsulis
subglobosis; seminibus longo-caudatis.

Seven species, all in northeastern Mexico.

KEY TO THE SPECIES

A. Lower leaf-surface pilose or strigose; stems rugosely hirsute, the base of the
trichomes thickened.

326 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

B. es pilose, the epidermal tissue visible; the lower pio surface
ROG co o8 so 44.s GA Fahd Khe E ee RA eae EY OAS Gaye BS 8. P. osmanthus.
BB. Scanie densely villose, the epidermal tissue obscure; ae lower
leaf-surface strigos
C. Leaves oblong; peieen 5-8 mm. long; petals aa at the

| + a ee ec er ee 9. oblongifolius.

CC. Leaves ovate: pedicels > 3 mm. long; petals one at the
apex.

D. Style hag center of disc pubescent; leaves on pee shoots

2-3 cm. long ...... 10. P. coultert.

DD. Style and center of disc glabrous; ‘leaves on ee ern

1-2 cm. long. 11. P. asperifolius.

AA. Lower leaf-surface densely villose; stem pilose, the base of the trichomes
not thickened
B. Leaves subcoriaceous, hispid above, densely villose and appearing white

beneath.
C. Style pubescent .....12. P. sargentianus.
CC. Style glabrous... 13. P. pringlet.
BB. Leaves characeous, weakly pilose. above, very sparsely villose be-
neath; style glabrous... ....s.................... 14. P. calcicolus.

8. Philadelphus osmanthus, sp. nov.

Frutex subscandens, 1.3 m. altus, ramosissimus; ramis robustis, 3—5
mm. diametro, fumeis, longitudinaliter rimulosis, bienniis mm. crassis,
castaneis, striatis, rugoso-hirsutis, corticibus siaaa: hornotinis 1-1.5 mm
diametro, pilis basi incrassatis, gemmis conicis, strigosis, pilis luteis; foliis
ovatis, raro ovato-lanceolatis, 2—6 cm. longis, 0.8 cm. latis, basi obtusis,
apice acutis vel obtusis, apiculatis, integris vel utrinque 1 vel 3 serrulatis,

longis, cum hypanthiis et calycibus incano-pilosis, hypanthiis subglobosis,

—7 mm. diametro; sepalis ovatis, acuminatis, 7-8 mm. longis; corolla sub-
cruciformi, 2.5—3.2 cm. diametro, petalis obovatis, apice emarginatis, 1.2—
1.5 cm. longis, 0.7—1.2 cm. latis; staminibus ca. 36; disco medio piloso,
stylo brevissimo, 1 mm. longo, vix hirtello, stigmatibus 3 mm. longis,
cristatis, supra divisis; capsulis ellipsoideis vel subglobosis, 6-8 mm. di-
ametro, calycibus persistentibus supra medium affixis; seminibus longo-
caudatis.

MEXICO: Hidalgo: Jacala, V. H. Chase 7310(NY. type; A, F, G, MO,
ISOTYPES ).

This species occurs in woods in deep ravines at an altitude of 1525
meters. Unlike other species of the genus, the lateral buds on the branch-
lets of this one are suppressed from normal development. Thus its flower-
ing shoots are largely developed from the terminal buds, a very unusual
character in Philadelphus. The thickened base of the epidermal hairs on
the branchlets, the elongate stigmas and the pubescent disc of this species
suggest relationship to P. coulteri Wats. The latter species can be dis-

1954] HU, THE GENUS PHILADELPHUS 327

tinguished by the thickly villose hypanthium. The subentire leaves
sparsely strigose and the long-tailed seeds of this species also suggest a
remote relationship with the southern Appalachian species P. inodorus
Linn. which has enclosed axillary buds. The very short style conforms
to Kornicke’s description of P. asperifolius, a species with glabrous style
and disc.

9. Philadelphus oblongifolius, sp. nov.

Frutex ramosissimus, ramis cinereis, ramulis gracilibus, bienniis 1—1.5
mm. crassis, brunneis, verrucoso-hirsutis, hornotinis 1 mm. iametro, pilis
basi incrassatis: foliis oblongis, raro oblongo-ovatis, 1.3-2.5 cm. longis,
4-8 mm. latis, utrinque obtusis, apice apiculatis, margine integerrimis,
utrinque strigosis, pilis subtus longioribus et densioribus, petiolo 3 mm.
longo; floribus solitariis, pedicellis 6-8 mm. longis, incanis, hypanthiis
calycibusque incanis; sepalis ovatis, 6 mm. longis, 2.5—-3 mm. latis; corolla
cruciformi, 3.5 cm. diametro, petalis obovatis, apice emarginatis, 1.6 cm.
longis, 1.1-1.4 cm. latis; staminibus ca. 30; disco praeter medio glabro,
stylo 5 mm. longo, supra diviso, basi dense hirsuto, stigmatibus 2 mm.
longis. Capsulis ignotis.

MEXICO: San Luis Potosi: Minas de San Rafael, C. A. Purpus
53686 (A, fragment of TYPE; F, Type; MO. NY. ISOTYPES ).

Through its strigose trichomes on the leaves, and pubescent styles this
species is closely related to P. Coulter Wats. It can be distinguished from
P. coulteri by its slender branchlets, oblong leaves, long pedicellate flowers
with emarginate petals and 5 mm. long styles.

10. Philadelphus coulteri Wats. in Proc. Am. Acad. 22: 472. 1887.
Philadelphus purpusii Brandegee in Univ. Calif. Publ. Bot. 4: 270. 1912.

Tyre: Dr. Coulter 77, Zimapan, Mexico (Gray Herbarium).

A subscandent shrub, the branches ferrugineous, the second year’s
growth 2-3 mm. in diameter, the bark rugose, closed, slightly longitudi-
nally rimulose; the current year’s growth rugose and hirsute, the trichomes
with thickened bases. Leaves ovate, or ovate-elliptic, subentire, obtuse
or acute, apiculate, rounded or rarely obtuse at the base, strigose on both
surfaces, more so beneath, those on the flowering branches, 1.5-3 cm.
long, 1-1.5 cm. wide, the petioles 3-4 mm. long. Flowers solitary, rarely
ternate, the pedicels 2-3 mm. long, thickly villose and white like the hy-
panthia and calyx, the sepals ovate, 5-6 mm. long; corolla disciform,
2.5-3 cm. across, the petals orbicular-obovate, rounded at the apex;
stamens 34-38; disc pubescent at the center, the style 2-3 mm. long, the
upper half divided, the stigmas 2—2.5 mm. long, the abaxial surface pap-
pilose, 2 mm. long, the adaxial surface half as long. Capsules and seed
not known.

328 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

MEXICO: Hidalgo: Zimapan, Dr. Coulter 77(G, type); woods above
Minas Viejas between Zimapan and Jacala, H. E. Moore, Jr. 2767(G, BH);
Atotonilco, Schiede, June 1830(NY). San Luis Potosi: Minas de San
Rafael, C. A. Purpus 4910(MO), 5368(F, G, MO, US isotypes of P. purpusii
Brandegee; A, photo and fragment). Tamaulipas: near Frank Harrison’s
“Rancho del Ciel” in Sierra de Guatemala above Gomez Tarias, Sharp, Shank,
Wolfe & Hernandez 52201 (A, TENN), 52060 (A, TENN).

Philadelphus coulteri Wats. was essentially based on Coulter 77 col-
lected from Zimapan. In publishing the species Watson also cited C. S.
Sargent’s collection from Monterey. As the species was named after
Coulter, his number 77 in the Gray Herbarium should be the actual type
of the species. Sargent’s material is specifically distinct. It differs from
the type of P. coulteri in having smooth pilose branchlets, leaves hispid
above and densely long villose beneath and a very short style. In 1888
Watson published an illustrated account of P. coulteri. As Coulter’s collec-
tion is rather fragmentary, that illustration was unfortunately drawn from
Sargent’s material. This publication has an out-reaching misleading effect
in the identification of the true P. coulteri among botanists as well as
horticulturists and consequently cytogeneticists.

Philadelphus coulteri Wats. was first collected in Zimapan, Additional
material extends its range northward to Gomez in the State of Tamaulipas.
At this area, the southeastern end of the Mexican Central Plateau, the
plant grows, according to Moore, as a shrub a little over a meter high and
its flowers are white, and fragrant. The standard reference books on
cultivated shrubs have recorded this species as having purple-centered

owers. This is apparently incorrect.

Watson published this species in 1887 on the basis of a herbarium
specimen. He did not know the flower color, nor did he indicate it in the
description. The next year an illustrated account was published in Garden
and Forest. Here no flower color was mentioned either. Three years
later, Burbidge wrote Hemsley from Dublin, Ireland, saying, “Can you
kindly give me the name of the enclosed? I cannot find it in the books.
It exists in one or two old gardens here, where it is called Rose Syringa.
Its sweet fragrance and purple-centered flowers are remarkable.” What
Hemsley reported we do not know, but this was later referred to P.
coulteri. Meanwhile, Nicholson in the Supplement to the Garden Dic-
tionary incorporated Watson’s review of P. coulteri from the Garden and
Forest. At the end of his summary Nicholson put down 1888, the year
when his source material was published, as the year of the introduction
of the plant into cultivation. Burbidge got this reference and in 1903 he
published a note saying “There is a variety of P. mexicanus called P. M.
Coulteri, introduced, it is said in Nicholson’s Supplement, as recently as
1888; but probably this is a mistake as the shrub has existed for
many years in old Irish gardens, where it is known as the “Rose Sy-
ringa” . . . and it differs from all other species or varieties . . . each of
its four white petals has a purplish blotch at its base.” This is a mis-
interpretation of Watson’s figure and description and also a misidentifica-

1954] HU, THE GENUS PHILADELPHUS 329

tion of the Irish “Rose Syringa.” But since that time in all major works
on cultivated shrubs and even in books dealing principally with the flora
of Mexico, the error in the flower color of P. coulteri Wats. has been
perpetuated.

Claims have been made by hybridizers and cytogeneticists that P.
coulteri Wats. is a grandparent of the triploid garden hybrids, “Belle
Etoile,” “Sybille” and “bicolor” which are segregates of P. purpureo-
maculatis Lemoine.! There seems to be no record that true P. coulteri has
ever been introduced into cultivation, thus it is impossible for it to have
been a parent of P. purpureo-maculatus Lemoine. The above mentioned
claims do not seem to have any material support. The Monterey region is
better known botanically than Zimapan, the type locality of P. coultert.
Consequently the Philadelphus that Sargent collected is much better rep-
resented in American herbaria than is true P. coulteri. It was almost
inevitable that these specimens should have been mistaken for P. coulteri.
When typical P. coulteri Wats. was again collected by Purpus in 1911,
Brandegee was misled into creating a new binomial, for P. pur pustt
Brandegee is the genuine P. coulteri Wats. Purpus 5368, the type number
of P. purpusii Brandegee represents an aggregate of three elements, prob-
ably three separate collections combined under one number of which I
have examined five sheets. The element with glabrous disc and style be-
longs to P. asperifolius Kornicke. The two other elements both have a
pubescent disc and style. One of them has ovate leaves and styles shorter
than the stigmas. This one agrees with Brandegee’s description orf.
purpusii, and is identical with the type of P. coultert Wats. The other
element has small oblong leaves and elongated style which is longer than
the stigmas. It becomes the type of P. oblongifolia S. Y. Hu.

11.

_

Philadelphus asperifolius Kornicke in Gartenfl. 16: 73. 1867. —
Koehne in Gartenfl. 45: 487. 1896; et in Mitt. Deutsche. Dendr.
Ges. 1904(13): 78. 1904. — Schneider, Il. Handb. Laubh. 1: 364.
1905. — Rydb. in N. Am. Fl, 22: 171. 1905.— Standl. in Contr.
U.S. Nat. Herb. 23: 310. 1922.

Type: Karwinsky July 1842, Hacienda Santyaguillo, Mexico (Herb.
Hort. Petropol.).

An elegant shrub, 2-3 meters high, branchlets rigidly hirsute, the
second year’s growth gray, longitudinally rimulose, the bark closed, the
current year’s growth brown, pubescent, the hairs with thickened bases.
Leaves ovate, those on the vegetative shoots up to 3 cm, long, 1.3 cm.
wide, those on the flowering shoots 1.3—1.7 cm. long, 0.5 cm. wide, rounded
at the base, obtuse and apiculate or subacute at the apex, both surfaces
sparsely covered with rigid, straight appressed white hairs, the petioles
1-2 mm. long. Flowers solitary, subsessile, the pedicels 1-3 mm. long,
canescent as are the hypanthia and the calyx, the sepals ovate, 3 mm.
long; corolla disciform, 1.5-2 cm. across, the petals obovate, rounded at

1. K. JANAKI AMMAL in Jour. Roy. Hort. Soc. 76: 272. 1951.

330 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXXV

the apex, twice the length of the sepals; stamens 30-38; disc glabrous,
the style short, about 1.5 mm. long, undivided, glabrous, the stigmas
separated, 2 mm. long, the abaxial surface twice as broad and as long
as the adaxial one. Capsules subglobose, 6-7 mm. long, 7 mm. diameter,
the persistent calyx subapical. Seeds long-tailed.

MEXICO: San Luis Potosi: Minas de San Rafael, C. A. Purpus 4910
(A, F, G, US), 5368a(G, MO, NY, US).

Unable to examine the type, the identification of this species is based
on the original detailed description published by Kornicke and the diag-
nostic characters supplemented by Koehne who probably saw the type
or an isotype. Kornicke described the species as having ovate or broad-
elliptic leaves which are obtuse and apiculate or subacute at the apex,
and about 3 cm. long and 1.5 cm. wide on sterile shoots. He did not
mention the nature and position of the buds and the presence or absence
of hairs on the style. For these characters we have to rely upon Koehne’s
observations. In his latest synopsis of the genus, under the subsection
Gemmati which was characterized as having a smooth almost obsolete
and glabrous style he placed P. asperifolius and P. serpyllifolius Gray.
Judging from the style of the latter species, the type of which is before
me, Koehne’s characters indicate a species in which the styles are short
but definitely present, 0.5—1.5 mm. long. By inference, Kornicke’s de-
scription, which calls for no style and the stigmatic column thick and
subsessile, may also be interpreted to apply to flowers with short but
definite styles. One of the elements of Purpus 5368, which I have in-
dicated as 5368a, has small ovate leaves, 0.5-1.7 cm. long, 0.4—0.7 cm.
wide on the flowering shoots, and glabrous styles. These characters con-
form to Kornicke’s description of the leaves and Koehne’s diagnosis of
the style of P. asperifolius, thus I interpret that element as representing
Kornicke’s species. Purpus 4910 is a poorly selected fruiting specimen.
With the exception of one, all sheets I have examined are sterile, and the
only fertile specimen has just one fruit with a broken top. The branching
is much looser than is that of Purpus 5368a. The leaves seem to fit
Kornicke’s description of those on the vegetative shoots. The small part
of the disc left on the broken fruit appears to be glabrous.

The ovate shape of the strigose leaves, the densely villose hypanthia
and the very short pedicel of the flower of this species suggest a very close
relationship with P. coulteri Wats. Yet it can be readily distinguished
from Watson’s species by its glabrous disc and style and its rather small
leaves on the flowering shoots which are only 1—2 cm. long. Philadelphus
coulteri Wats. has a pubescent disc and style, with the leaves on the
flowering shoot 2—3 cm. long. The fragrant white flowers of this species
appear in June.

12. Philadelphus sargentianus, sp. nov.

Philadelphus coulteri Wats. in Proc. Am. Acad. 22: 472. 1887, pro parte
(quoad col. Sargent, excl. Coulter 77); et in Gard. Forest 1: 232, fig. 40.

1954] HU, THE GENUS PHILADELPHUS aot

1888. — Nicholson, Suppl. Dict. Gard. 594. 1900.— Rydberg in N. Am.
Fl. 22: 170. 1905. — A. H. Moore in Bailey, Stand, Cycl. Hort. 5: 2582,
fig. 2905. 1915.—Standl. in Contr. U.S. Nat. Herb. 23: 311. 1922.—
Rehder, Man. Cult. Trees Shrubs 280. 1927; ed. 2. 275. 1940; et Bibliogr.
Cult. Trees, Shrubs 159. 1949.

Frutex usque 2.3 m. altus, ramis robustis, bienniis 3 mm. diametro,
brunneis, longitudinaliter rimulosis, hornotinis 1.5 mm. diametro, dense
hirsutis, pilis basi haud incrassatis, gemmis axillaribus conicis, strigosis,
pilis luteis; foliis ovatis vel ovato-ellipticis, 2.5—4.5 (raro 5) cm. longis,

latis, supra hispidis, subtus dense argenteo-villosis, 3- vel 5-nerviis,
margine inconspicue serrulatis, dentibus utrinque | usque ad 5, minutis,
petiolo 3-5 mm. longo; floribus solitariis, pedicellis brevissimis, 2—3 mm.
raro ad 6 mm. longis, cum hypanthiis calycibusque dense albo-villosis;
hypanthiis subglobosis, 5 mm. diametro, pilis longioribus et brevioribus
intermixtis; sepalis ovatis, acuminatis, 8 mm. longis, 4-5 mm. latis; corolla
usque 4.5 cm. diametro, petalis obovatis, 2 mm. longis, 1.6 cm. latis, apice
erosis; staminibus ca. 40, antheris 2 mm. longis; disco medio piloso, stylo
brevissimo, 1 mm. longo, stigmatibus 4 mm. longis, liberis; capsulis sub-
globosis, 1 cm. diametro; seminibus longe caudatis.

MEXICO: Nuevo Leon: Monterey, Sierra Madre Mts., Diente Canyon,

_H. & M. T. Mueller 544(A, F); C. G. Pringle 2094(A, NY, US); C. S.
Sargent in April 1887(A Type; G, originally placed under P. coulteri); Cerro
de la Silla, S. S. White 1470(G).

In the publication of P. coulteri, Watson cited two collections, Coulter
77 from Zimapan and Sargent s.n. from Monterey. Watson’s species was
published in 1887. Sargent’s specimen was collected in April of that
year. It is highly possible that Watson had prepared the description of
P. coulteri before he saw the Sargent specimen. When Sargent sent him
his Monterey collection, Watson simply incorporated it in his new species.
As the species was named for Coulter, his collection naturally becomes the
type of P. coulteri Wats. But Sargent’s collection is definitely different
from the type of P. coulteri Wats. which has the rugose branchlets and
strigose leaves, characteristic of species like P. mexicanus Schlechtendal.
Sargent’s material has villose branchlets and leaves hispid above and
densely white villose beneath, a type of pubescence characteristic to the
northern element like P. madraensis Hemsl. Yet Sargent’s specimen is not
Hemsley’s species because of its exposed buds, and it is thus described
as new here.

Philadelphus sargentianus is endemic to Monterey where it grows at
altitudes of 1500-1600 meters, where the mean annual temperature is
62°-67° F. and the mean annual rainfall is limited to 30-40 inches.
There its fragrant white flowers appear from early April to July. As far
as I know this species has never been introduced into cultivation. Un-
fortunately misled by the publication of an illustration drawn on the
basis of the Sargent collection, several authors of standard references on

332 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXXV

the cultivated trees and shrubs had misinterpreted this species as P.
coultert.

13. Philadelphus pringlei, sp. nov.
Philadelphus coulteri sensu Brandegee in Univ. Calif. Publ. Bot. 4: 270. 1912,
non Wats

Frutex, ramis brunneis, bienniis 2 mm. crassis, longitudinaliter rimu-
losis, hornotinis 1.5 mm. diametro, dense hirsutis, pilis basi incrassatis,
gemmis axillaribus conicis, pilosis; foliis ovatis, raro lanceolatis, 2.4—5
cm. longis, 1-1.8 cm. latis, basi rotundatis, obtusis, raro acutis, incon-
spicue triplinerviis, apice acutis, breviter acuminatis vel raro obtusis,
apiculatis, inconspicue serratis, serrulis utrinque 2 usque ad 5, minutis
argutisque, supra hispidis, subtus dense argenteo-villosis, petiolo 4-5 mm.
longo; floribus solitariis, pedicellis 2-4 mm. longis, cum hypanthiis calyci-
busque dense albo-villosis; hypanthiis subglobosis, 5 mm. diametro;
sepalis ovatis, 8 mm. longis, 5 mm. latis, acuminatis; corolla cruciformi,
—5 cm. diametro, petalis obovatis, 2 cm. longis, 1.5 cm. latis, apice ro-
tundatis; staminibus 40 usque ad 44, antheris oblongis, 2 mm. longis;
disco glabro, stylo 4 mm. longo, glabro, supra diviso, stigmatibus 3-3.5
mm. longis; capsulis subglobosis, immaturis 7 mm. diametro.

MEXICO: Nuevo Leon: Sierra Madre, above Monterey, C. G. Pringle
10174 (A, type; F, G, MO, US, tsorypes); Diente Canyon, about 12 miles
south of Monterey, Ci & M. T. Mueller 261(A, F); Waterway below
Alamar, about 15 miles southwest of Galeana, C. H. & M. T. Mueller 656(A,
F, NY).

Geographically this species has the same range as P. sargentianus S. Y.
Hu and morphologically they are very similar except P. sargentianus
has a pubescent disc and style, while this species has glabrous ones. Ac-
cording to the collectors, this taxon is abundant in the open woods and
shrub zones of the Sierra Madre Oriental in the state of Nuevo Leon. Its
white flowers appear in April and May.

14. Philadelphus calcicolus, sp. nov.

Frutex usque ad 5 m. altus, ramulis teretibus, cinereis, bienniis 1.5 mm.
crassis, corticibus clausis, hornotinis brunneis diametro, pilosis,
gemmis terminalibus conicis, strigosis, pris luteis; foliis serratis, lanceo-
latis vel ovatis, 2.5-5 c m. longis, 1—2 cm. latis, basi acutis vel obtusis,
trinerviis apice chicintaae, et apiculatis vel acutis, supra in sicco
nigrescentibus, pilosis, subtus olivaceis, sparsissime villosis, petiolo 5-8

mm. longo; floribus solitariis, pedicellis 3-6 mm. longis, incanis, hypan-
thiis subglobosis, 6 mm. diametro, dense tomentosis, argenteis; sepalis
ovatis, acuminatis, 7 mm. longis, basi 4.5 mm. latis, argenteis; corolla
3 cm. diametro, petalis obovatis, apice emarginatis, 1.5 cm. longis, 1.3 cm.
latis; staminibus ca. 40, antheris sagittatis, 2 mm. longis; disco glabro,
stylo brevissimo, 1.5 mm. longo, stigmatibus 4 mm. longis, cristatis,

3
.5

1954] HU, THE GENUS PHILADELPHUS R RE,

coalescentibus; capsulis ellipsoideis, 8 mm. diametro, sepalis persistentibus
Y% apicem insertis: seminibus breviter caudatis.

MEXICO: Nuevo Leon: Dulces Nombres, F. G. Meyer & D. J. Roger
2662 (TYPE, MO; fragment A); Canyon above Linare toward Galeana, A. J.
Sharp 45787 (TENN. A, sterile). Tamaulipas: Road above Cindad Vic-
toria toward Jaumave, dry canyon slope below 3000 ft., Sharp, Shanks, Wolfe
& Hernandez T3006 (A, TENN).

This species grows on dry limestone cliffs above a dry stream bed at
an altitude of 1300 meters. The white flowers appear in June. It is a very
interesting plant for most of the lateral buds on the branchlets are weak,
as a rule not visible, and do not develop in the following year. Often only
those situated near the apices of the shoots, either the apical alone or the
two axillary ones, develop into flowering or vegetative shoots. In this
respect as well as by its short styles with long cristate stigmas, emargi-
nate petals, and slender petioles this species resembles P. osmanthus S. Y
Hu. The latter species can be distinguished by its pubescent disc, less
canescent hypanthium and calyx, and hairs with thickened bases on the
stem. Its smooth appressed white indumentum on the hypanthium and
calyx and its hairs on the stem not thickened at the base also suggest a re-
lationship with P. argenteus Rydb., in the section Microphyllus of the sub-
genus Euphiladelphus. Rydberg’s species can be distinguished by the en-
closed buds and small entire leaves which are densely white pubescent be-
neath. The coalescent stigmas and the short caudate seeds also suggest
some relationship with the southern Appalachian species P. hirsutus Nutt.,
but the latter species can be distinguished by its prominent axillary buds,
the style longer than the stigmatic column, and the ecaudate seeds.

(To be concluded)

334 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

POLYPLOIDY AND APOMIXIS IN COTONEASTER

Hatiy J. Sax

THE RosAcEAE have drawn the attention of investigators because of
the variety and beauty of their flowers and fruits as well as their horticul-
tural value. They are a large and varied family with a wide distribution.
Many examples of polyploidy and apomixis have been reported in this
amil

Of the four subfamilies of the Rosaceae, the Pomoideae are of special
interest because of their allopolyploid origin. The basic chromosome
numbers of the other subfamilies, the Rosoideae, Prunoideae and Spi-
roideae, are 7, 8, and 9. The basic chromosome number of the Pomoideae
is 17, derived from primitive ancestors by hybridization followed by
chromosome doubling.

The eighteen genera of the Pomoideae are rather closely related, as is
shown by the fact that many generic hybrids occur in nature. Their close
relationships are further supported by the readiness with which most of
the genera can be intergrafted.

Many of the genera of the Pomoideae are characterized by secondary
polyploidy, that is, the chromosome number in many cases has doubled
again, this time probably by autopolyploidy. Secondary polyploidy is
common in Malus and Crataegus.

The genus Cotoneaster is less well known than Malus and Crataegus
of the same subfamily. In his “Manual of Cultivated Trees and Shrubs”
Rehder (56) notes that there are about 50 species of Cotoneaster in the
temperate regions of Europe, N. Africa and Asia (except Japan, where
there are no native species). These are mostly shrubs, rarely small trees,
with some beautiful prostrate forms among them. Their fruit is especially
attractive: red, black, purple and a few orange pomes. They are planted
often because of their brightly colored fruits as well as the flowers, which
are in many cases small, white to pinkish and of many-flowered corymbs.
There are also some evergreen and partly evergreen varieties, both up-
right and prostrate forms, which are very beautiful with the brilliantly
colored fruit.

Plant collectors have brought many species of these Cotoneasters into
cultivation. Their size, variety, and adaptive qualities make them desir-
able as ornamental plants. The relationship of these species is not very
easily determined, and the Cotoneasters have been considered a difficult
genus by the taxonomist.

The characteristics often show differences in size, hairiness, etc., but
mostly intensification of some character predominating over that in an-
other species, and often that is the only difference. In reading over the
descriptions of the many species and varieties, as well as observing them

1954] SAX, POLYPLOIDY AND APOMIXIS IN COTONEASTER 335

in the field, it becomes apparent why Cotoneaster is a difficult genus for
the taxonomist. In describing C. francheti var. sterneana, W. B. Turrill
(74) in 1950 remarks, “While we have not sufficient information to give
full reason for this, since we do not know how far hybridization happens
in the wild, how plastic are individual plants, or how much intraspecific
genetic variations occur, there results frequent difference of opinion be-
tween ‘lumpers’ and ‘splitters.’ Splitters have been carried too much.”’ He
cites Crataegus and appeals to the cytogeneticist to study Cotoneaster.

POLYPLOIDY AND APOMIXIS IN THE ROSACEAE

Doubling of the chromosome number at the time of union of the egg
and male nuclei followed by a reduction to half the number at the time
the spore mother cells divide to form the megaspores and microspores
is general throughout the plant kingdom. Asexual methods, such as root
or stem cuttings, leaf and axillary bulbs, budding, etc., also occur, but
these do not involve a doubling and reduction of the chromosome number;
the offshoot carries the chromosome number of the organ from which it
comes, while in sexual reproduction there is an alternating cycle between
the haploid and the diploid number of chromosomes.

Many instances of changes, modifications of the usual behavior, or
abnormalities have been found to occur in the development of the egg and
male nuclei and in the development of the spores to form the gametophytes
which produce the eggs and the male nuclei. Some of these, such as the
functioning of a cell or cells of the integument of the ovary to form the
embryo, or the direct development of a megaspore mother cell without
reduction to form the spores, the development of an unreduced egg cell.
do not involve a change in the chromosome number, These various sub-
stitutions for fertilization and meiosis are classified under the term apo-
mixis. Parthenogenesis, apospory, apogamy. and vegetative buds are all
included.

No attempt will be made to review all the literature on the subject of
apomixis. The earlier work has been reviewed by several investigators,
Ernst, 1918 (16) and others including Rosenberg (58) in 1930, who gave
a summary of the work up to that time. Steil in 1939 (70) reviewed the
literature on ferns. Then Stebbins (68) brought the review up to 1941
in his discussion on ‘““Apomixis in the Angiosperms.” The subject again
underwent a thorough review by Gustafsson (31) in 1947. Subsequently
many instances of apomixis have been reported, especially in connection
with polyploidy. This work, as well as the extensive literature on poly-
ploidy, will be discussed here only in sail with the work on the
Rosaceae, of which Cotoneaster is a mem

The role of polyploidy and apomixis in ees evolution and geo-
graphical distribution is well illustrated in many genera of the Rosaceae.
In general there is much variation in the appearance of apomixis and a
relatively high frequency of polyploidy.

Much of the early work on polyploidy in relation to speciation was done

336 JOURNAL OF THE ARNOLD ARBORETUM [VOL, XXXV

with the genus Rosa by Blackburn and Harrison (4) in 1921, Tackhohn
(71) in 1922, and Hurst (36, 37, 38) in 1925, 1928, and 1932. A wide
range of polyploidy was found, and Hurst attempted to classify the species
of Rosa into five basic genomes. Crosses between the basic diploid species
gave rise to the complexities of the polyploid species. Later cytological
studies of this same group by Gustafsson and Hakensson (32) and Gus-
tafsson (29) gave further information indicating that the cytogenetic rela-
tionships of the Rosa species are more complex with both auto- and alloploid
as well as intermediate complexes. Fagerlind (18, 20) goes even further
and suggests that segmental interchange between the chromosomes of the
basic genomes leads to complex pairing relationships in triploids. Both
Gustafsson and Fagerlind concluded that apomixis in Rosa was a species
trait present in diploid species, and that the present association between
polyploidy and apomixis is secondary.

Many of the North American species of Rubus were found to hybridize
by Brainerd and Peiterson (5) in 1920 and Peiterson (52) in 1921. In
the experiments, hybrids between different sexual species of Rubus
(Peiterson, 52) and other genera have in no instances shown any clear
indication of apomictic reproduction, even though the parental species are
closely related to apomictic forms. Longley (43) in 1924 suggested that
apomictic forms existed. In 1930 Gustafsson (25) reported unreduced
pseudogamy in a species of Rubus. In some cases of pseudogamy the
number of apomictic and sexual offspring vary according to the chromo-
some number of the pollen parent. Darrow and Waldo (11) 1933, re-
ported that the fertilization of a tetraploid species of Rubus with pollen
from a diploid species yielded a majority of sexual offspring; but it and
other tetraploid apomicts produced few or none with pollen from tetraploid
species. Crane (7) obtained similar results when using pollen from a
diploid form of R. idaeus on an octaploid species of R. vitifolius: only
pentaploid hybrids were produced. When he used pollen from the tetra-
ploid form of the same parent, hybrids and matriclinous octoploid offspring
resulted. Pollen from the hexaploid R. loganobaccus (2 n = 42) pro-
duced only hybrids. Petrov (53), using pollen of the hexaploid R. logan-
obaccus on a triploid with unreduced eggs, obtained hexaploid hybrids;
but using pollen of a triploid, R. idaeus, only triploid pseudogamous
progeny result.

Although polyploidy is common in the genus Rubus, Gustafsson (27,
28) found apomixis was confined to only one section. As in Rosa, many of
the species are facultative apomicts.

Many of the facultative apomicts are heterozygous, and Haskell (34),
1953, has suggested that another factor in the variation of Rubus mav be
crossing over at meiosis in the production of an unreduced egg cell. In
Rubus crosses between two facultative apomicts give sexual progeny.

The genus Potentilla has also provided evidence regarding the role of
polyploidy in evolution and speciation (Claussen, Keck and Hiesey 6),
1940, and the role of apomixis. Potentilla species (Miintzing 46 and 47)
showed variation in the development of the microspore mother cell, Meio-

1954] SAX, POLYPLOIDY AND APOMIXIS IN COTONEASTER 337

sis was regular in the microspore mother cell of P. argentea but irregular
in P. collina and P. hirta. Potentilla argentea is apomictic through faculta-
tive apospory. The embryo may develop autonomously. Similar results
were reported by Popoff (55), Gentcheff (23) and Gentcheff and Gus-
tafsson (24). The genetic studies of Mintzing and Miintzing (49) on
reproduction between sexual and apomictic forms indicate that apomixis
is controlled by multiple factors.

Among the Prunoideae the genus Prunus has been studied extensively.
Meurman (45) in 1929 reported a high polyploid species, Prunus lau-
rocerasus L. with eighty-eight chromosomes. Polyploidy is common in
Prunus, and the allopolyploid origin of P. domestica was determined by
Rybin (59) in 1936. According to Almeida (1), P. lusitanica is an octo-
polyploid with 2 n = 64, an allopolyploid of ancient origin behaving like
a diploid. Cytological work by Schelhorn (65) showed great irregularity
in the nuclear divisions and tetrad formation in a triploid of P. avium,
which the author believes arose by the fertilization of an unreduced
sexual cell of P. avium with a sexual cell, and not as a cross of P. avium
and P. cerasus.

The basic diploids show considerable stability of the genome, as is
evidenced by the regular meiosis and high fertility of the hybrid between
P. tomentosa from China and P. besseyi from Central North America
(Sax unpublished). Apomixis has not been found in this subfamily.

Little cytological work has been done on the Spiroideae, but both
autopolyploidy and allopolyploidy have been found in Spiraea, Sax (64).
Polyploidy appears to be related to geographical distribution, since the
Old-World species are largely diploid while most of the American species
are late-flowering tetraploids.

The Pomoideae are unique in that the subfamily is of allopolyploid
origin. Secondary polyploidy is common in most of the genera, and trip-
loids play an important role in both natural and cultivated species.
Polyploidy has complicated the genetic and taxonomic variation of most
of the genera and in a few cases is related to geographic distribution.

The first extensive work on the genera of Pomoideae was done by
Longley (42) in 1924. He determined the chromosome number in eighty
species of Crataegus from the collection in the Arnold Arboretum. He
found thirteen diploids, sixty-seven triploids, and ten tetraploids.

The chromosome relationship in the Pomoideae was studied by Sax
(62 and 63). In the Rosaceae the subfamilies, the Rosoideae, Prunoideae
and Spiroideae, have basic chromosome numbers 7, 8 and 9, while the
Pomoideae have 17 as the basic chromosome number. The close rela-
tionship among them suggests that they originated as a cross between
primitive ancestors of the other Rosaceae.

Dermen (12) in 1936 discovered in Malus hupehensis a delayed de-
velopment of the embryo from the unreduced egg after the flower opened.

Johansson (39) suggested that there were four places of fruit origin:
the Caucasian region, the Turkestan region, the East Asiatic region in-

338 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XXXV

cluding East Siberia, and North America. The new varieties arose through
crossing, mutation, and doubling of the chromosome number.

Polyploidy in Malus has been studied by several workers, Einset (13,
14) found spontaneous polyploids among apple seedlings. Einset and
Inhofa (15) also described periclinal chimeras; partly diploid, partly
tetraploid, in apples. Hemming (35) discusses the origin of apples and
the relation of diploid, triploid, and tetraploid species.

Some correlation has been shown between geographical distribution and
polyploidy. Hagerup (33) in 1931 postulated an increase in the frequency
of polyploidy with the increase in latitude. The results of studies by
Tischler (73) of the frequency of polyploids at different latitudes in Sicily,
Schleswig-Holstein, the Faeroes and Iceland supported that hypothesis.
Further corroboration came from Flovik (22) in his investigations of the
flora of Spitzbergen, all glacial survivors.

Gustafsson (26), Babcock and Stebbins (2), and Stebbins and Bab-
cock (67) have found apomicts to be excellent material for studying the
effect of environment on the genotype and the tracing of plant migrations.
The distributional center of the American Crepis complex was found in
northeastern California. Fernald (21), followed by Babcock and Stebbins
(2), also considered the Appalachians to hold the ancient American An-
tennaria complex. Gustafsson (28) has made similar studies of Taraxa-
cum and Rubus in Scandinavia. Curtis (9) also used apomicts in Taraxa-
cum in England for the study of the effect of environment on the genotype.

Other literature on the subject of plant distribution was reviewed by
Love and Love (44) in their work on “The Geographical Significance of
Polyploidy.” They conclude that polyploidy increases with the increase
in latitude or the extremeness of the Pleistocene or post-glacial climate.
The estimated frequency of polyploidy in the temperate zone is 30 per
cent or lower.

COTONEASTER

In the present work the chromosome numbers of most of the species
in the genus Cotoneaster were studied. It was hoped that this would show
something concerning the relationships of some of the species, whether or
not the species were polyploid, and give evidence as to whether there
existed in this genus some of the irregularities in reproduction found in
other genera of the Rosaceae as in Malus and Crataegus.

The presence of apomixis was also tested. Each season the flowers on
some of the branches of a few species were destyled and emasculated
while in the bud on some of the plants to see if they would produce fruit
without fertilization; and if, as has been found in the apples, the develop-
ment of the embryo could take place independently of fertilization by
development from some part of the ovule. There are only a few species
in which this work was done. Since only positive results were conclusive
because in a few cases no fruit was set on the controls or occasionally
on the entire plant, these results will be mentioned in connection with the
species.

1954] SAX, POLYPLOIDY AND APOMIXIS IN COTONEASTER 339

Seed from some of the species were planted to compare the progeny
from the same seed source, to observe their resemblance to the parent
plant, and to determine the proportion of hybrid plants occurring in
natural populations.

The Cotoneasters in the Arnold Arboretum are all introduced species
from Europe, Asia, and Africa. A few of these species were obtained from
crosses made in cultivation; others were collected and brought into cul-
tivation. The source of the species was usually known, especially the im-
mediate place from which it was introduced. In most cases the plants
came directly from their place of origin. The Arnold Arboretum’s own
collector, Ernest Wilson (75) brought seeds and plants from Asia directly
to the Arnold Arboretum. Many of them bear the authority of Rehder
and Wilson. In other cases seeds have come from well-known plant ex-
plorers in England, France, Germany, Holland, China, and India, and
other equally familiar sources elsewhere and in the United States.

The material for the present work was collected during the spring and
summer of the years 1950 through 1954 and was confined to the species
available in the Arnold Arboretum. This included most of the well-known
species, as may be noted from the list given in the table (Tab‘e I), al-
though not all the varieties of each species were studied. There were a few
plants labeled “C, sp.” denoting the difficulty in naming them, and as
mistakes in labeling may occur in the best collections, the number on the
metal label on each specimen was copied for further tracing if necessary.
All were checked with the specimens in the herbarium of the Arnold
Arboretum.

The Cotoneasters blossom over a fairly long period in the spring, but
it is necessary to obtain the dividing microspore mother cells at the right
stages in division. During a particularly warm day most of the buds on
a plant might pass through the division stages. In some species where new
buds are produced over a longer period, material is more easily obtained.
The difficulties of fixing such a large number of varieties in a short time
meant that some were missed some years. In some rare cases the plants
were not in flower every year or they were newly planted. The size of
the flower bud when the divisions occurred differed in the various species.
However, most of the available species, though not all, were obtained
when the divisions were in progress each season, and the results were
determined and checked. All those reported were checked at least two
seasons, and many were studied four or five seasons.

The buds were fixed in alcohol acetic solution twenty-four hours, then
changed to 95% alcohol. These were left under refrigeration until aceto-
carmine smears of the pollen mother cells could be made and studied.

The chromosomes in the Cotoneasters are small, but there was an
abundance of material and in practically all cases it was possible to ob-
tain well-fixed material at all stages.

In the study of the chromosome numbers it was noticeable that the
chromosome pairing was quite characteristic for any given species. In
many of the species there was a tendency for the sets of chromosomes to

340 JOURNAL OF THE ARNOLD ARBORETUM [| VOL. XXXV

adhere strongly in the late prophase and metaphase, and at these stages
in many cases the count would be 17 or thereabout even when the form
was triploid or tetraploid except for a few univalents. The chromosomes
in these cases would appear unusually large for Cotoneaster. However,
as the chromosomes separated and were counted in the anaphase stage,
it was apparent that the species were often triploid or even tetraploid.

The diploids are very regular in their divisions. The 17 small chromo-
somes pass to the poles and are easily counted. The polyploids were not
so easily counted in all cases, but they had a very characteristic appear-
ance. The chromosome complex was larger and the pollen mother cells
were usually so much larger that they suggested polyploidy after one be-
came familiar with them. This did vary somewhat, but in all the diploids
studied the pollen mother cells were smaller than in most polyploids;
exceptions were noted in the polyploids.

Univalents were very often present in the dividing nuclei of the poly-
ploids. The triploids showed chromosome bridges and other irregularities.
The chromosome count varied according to the closeness of adherence of
homologous pairs. Often pieces of chromatin, even whole chromosome
bridges were left outside in the cytoplasm. As many as twelve such chro-
mosome remnants were counted, but usually two to six were found outside
in the cytoplasm. These were visible after the daughter nuclei had rounded
up in the later stages. Tetraploids were less irregular, although univalents
were often present. The number was obviously larger, and when they
separated the counts showed the tetraploid number. At time some of the
univalents were lost in the cytoplasm.

The counting of the chromosomes in the polyploids is complicated by
an early loosening of the arms of the V-shaped chromosomes as they near
the poles, which makes it appear as a larger number of chromosomes if
the two arms are counted separately.

In every species the counts were checked from the same slides by
Professor Karl Sax, to whom I wish to express my thanks.

The results of these studies are given in Table I. The species are listed
in alphabetical order.

TABLE I
COTONEASTER
Chromosome

Species and Variety Habitat Zone Number
C. acuminata Lindl. 226-39 * Himalayas 5 2n
C. acutifolia Turcz. 15686 N. China 4 2n
C. acutifolia ? 2291 Tao Basin 3n

C. acutifolia var. villosula Rehd.
& Wils. 13165B Cent. & W. China 5 4n

* Numbers refer to Arnold Arboretum accession numbers.

1954] SAX, POLYPLOIDY AND APOMIXIS IN COTONEASTER 341

Chromosome
Species and Variety Habitat Zone Number
C. adpressa Bois 7951 W. China 4 3n
C. adpressa hessei 813-41 3n
C. adpressa var. praecox (Vilm.)

Bois & Berthault. 134-18 3n
C. affinis var. bacillaris (Lindl.)

Schneid. 17806-A & B Himalayas 7 4n
C. ambigua Rehd. & Wils. 134-22 W. China 5 3n
C. apiculata Rehd. & Wils. 7275 W. China 4 3n
C. bullata Bois 861-—32A W. China rs) 3n
C. bullata f. floribunda (Stapf)

Rehd. & Wils. 6685-2 W. China 3n
C. bullata var. macrophylla Rehd.

& Wils. 13426 W. China 3n
C. conspicua Marquand 1019-36C W. China i? 2n
C. dammeri Schneid. 137-51 Cent. China dP 2n
C. dielsiana Pritz. 686-33 Cent. & W. China 5 3n
C. dielsiana Pritz. 134-28A & B 3n
C. divaricata Rehd. & Wils. 6587C Cent. & W. China 5 3n
C. foveolata Rehd. & Wils.

13431A & B Cent. Chin 4 3n
C. francheti Bois 130-32D W. China 6? 4n
C. frigida ? Lindl. 191-40 Himalayas 7 2n
C. froebelli Vilmorin 757-30B Cult. 3n
C. glabrata Rehd. & Wils. W. China | 2n
C. glaucophylla Franch. 571-36 W. China 7? 3n
C. henryana Rehd. & Wils.

223-07 Cent. China 7? 2n
C. horizontalis Decne. 45-34 W. China 4 3n
C. horizontalis var. perpusilla

Schneid. 7157A W. China 3n
C. horizontalis var. prostrata

1070-38 3n
C. integerrima Med. 1776-3,

1766B Europe, N. Asia to

Altai 5 3n
C. lindleyi Steud. 372-37-A Himalayas 6? 3n
C. lucida Schlecht. 3284A & B N. China, Mongolia

Altai Mts. 4 oi
C. melanocarpa Lodd. 6679-1 Europe to Cent. &

N.E. Asia 4 4n
C. melanocarpa var. commixta

Schneid. 656-33 4n?
C. melanocarpa var. laxiflora

(Lindl.) Schneid. 13490-1A Cent. Asia 3n
C. microphylla Lindl. 22906 Himalayas 5 2n

. moupinensis Franch. 13497A

& B W. China 6? 3n

C. multiflora Bge. 21976, 14916 W. China 5 4n

342 JOURNAL OF THE ARNOLD ARBORETUM [| VOL. XXXV
Chromosome
Species and Variety Habitat Zone Number
Cc. eta var. calocarpa Rehd.
& Wils. 6679-1 & 2 W. China 3n?

C. multiflora var. granatensis

(Boiss.) Wenz. Spain 6 3n
C. nitens Rehd. & Wils. 6681 W. China 4 3n
C. obscura Rehd. & Wils. 6686-1 W. China 5 3n
C. obscura var. cornifolia Rehd.

& Wils. 11261A & B W. China 3n
C. racemiflora var. desfontaint

(Reg.) Zab. 389-34 S. Eur., N. Afr., W.

Asia to Himalayas
and Turkestan 4 3n

C. racemiflora var. soongorica
(Reg. & Herd.) Schneid. 16428.
21846 W. China 3 3n, 4n

C. racemiflora var. veitchii Rehd.

ils. 20075 Cent. China 3n
C. rosea Edgew. 44939 N.W. Himalayas,

Afghanistan 5 3n

C. rotundifolia Lindl. 405-36A Himalayas 6? 3n
C. rubens W. W. Sm. 21991B W. China 6? 3n
C. salicifolia Franch. 434-33A W. China 6?
C. salicifolia var. rugosa (Pritz.)

Rehd. & Wils. 458-36-A Cent. China 5 3n
C. schneideri 574-38-B 4n
C. simonsiit Bak. 596-33 N.W. India, Khasia 5 3n
C. sp. H. Hesse 13492 3n
c. D524. 30A & B 3n
C. sp. 114-36B China (Lu Shan Arb.) 3n
C. tenuipes Rehd. & Wils.

7276A-—C W. China 5 3n
C. tomentosa Se Lindl.

23-42-4A, 13507-1 S.E. Europe, W. Asia 4 3n
C. wardi W. W. aa 659-33 S.E. Tibet 7? 3n
C. zabeli Schneid. 7019B Cent. China 4 3n
C. zabeli var. miniata Rehd. &

Wils. 7343D, 156-85 China 3n

Triploid and tetraploid Cotoneasters are apt to be more vigorous than
the diploids — but there is much variation among these; some are less
vigorous. Often the main differences given in Rehder’s Manual between
some of the species and varieties is in vigor, intensity of color, size of

tions existing between known diploid and triploid or polyploid species
derived from them.

Cotoneaster acuminata Wallich is one of the first species described
(Flora, 1823). It is a diploid species from Zone 5 in Nepalia in the

1954] SAX, POLYPLOIDY AND APOMIXIS IN COTONEASTER 343

Himalayas (Table I.) A natural hybrid between this species and C.
acuminata was collected by R. N. Parker. Cotoneaster acuminata is found
along the Himalayas to Calcutta.

According to the table (Table I) the chromosome count for Cotoneaster
acutifolia Turcz. is 17. The pollen mother cells were small. It is a diploid
and its natural habitat is N. China. It has been found on a mountain
near Peking by Bretschneider. Specimens in the herbarium from various
localities in northwest China at high altitudes are labeled C. acutifolia
Turcz. Wilson (75) says there is no typical acutifolia in China proper,
but in Mongolia, but that there are two well-marked varieties in central
and west China.

Another plant labeled C. acutifolia (?), carrying a different number,
was grown from seed collected by Joseph F. Rock in the Tao River Basin,
Kansu Province, China. It is obviously a different variety with a triploid
chromosome count and lagging chromosomes. This is a triploid probably
derived from C. acutifolia Turcz.

One of the varieties, C. acutifolia var. villosula Rehd. & Wils., is found
in Central and West China. This is a polyploid and was recognized as a
new variety by Rehder and Wilson (56). It is described as ‘“‘densely
villous beneath, somewhat larger; calyx tube more densely villous; fruit
thinly pubescent.” From the chromosome counts this was considered a
tetraploid, although there was some irregularity.

Closely related to C. acutifolia Turcz. is C. ambigua Rehd. & Wils.
from West China which, according to Rehder’s Manual of Cultivated
Trees and Shrubs (56), “differs chiefly in its slightly pubescent or nearly
glabrous calyx tube and subglobose fruit with three to four nutlets.”
Cotoneaster ambigua is a coarser plant in general and more rigid in ap-
pearance than C. acutifolia Turcz. and bears a closer resemblance to C.
acutifolia var. villosula Rehd. & Wils., but on comparison of the two
plants side by side in the field they appeared sufficiently different to be
separate species. In Plantae Wilsonianae, Wilson (75) makes the remark,
“In speaking of ambigua, it is noted that all acutifolia like specimens are
grown in the arboretum and later it may be possible to determine their
relation to each other.” Cotoneaster ambigua is a triploid. The divisions
of the nucleus in the pollen mother cells are irregular with bridges still
holding at anaphase and scattered univalents. The homologues show
some tendency to adhere. The pollen mother cells often did not finish
dividing but decomposed at some stage in their development. There was
also much variation in the stages in an anther and much decomposition in
the pollen mother cells. Cotoneaster ambigua produced fruit. The speci-
men in the Arnold Arboretum was heavily fruited in 1953, suggesting
apomixis in such an irregular triploid.

Cotoneaster tenuipes Rehd. & Wils. is also noted as a species closely
related to C. acutifolia Turcz. It has a very graceful fine slender develop-
ment throughout and would not be mistaken for either C. acutifolia var.
villosula or C. ambigua. It is less coarse even than the diploid species C.
acutifolia Turcz. Cotoneaster tenuipes is a triploid form which has smaller

344 JOURNAL OF THE ARNOLD ARBORETUM [| VOL. XXXV

chromosomes and pollen mother cells like those of the diploid, but the
chromosome number is higher than the basic number for the genus. There
were chromatin bridges and fragments in the cytoplasm at anaphase.
Although the pollen grains were smaller, like those of a diploid, it was
obviously a triploid or aneuploid. There was little mature fruit on the
plant in the years 1950-1952, but it was well fruited in 1953. It is
probably apomictic.

Cotoneaster tenuipes Rehd. & Wils. grew in West China, as did C.
ambigua Rehd. & Wils. and C. acutifolia var. villosula Rehd. & Wils.,
the latter extending into central China. The two former are triploids, the
latter a tetraploid. They grew in Zone 5 (i.e. Rehder’s Map of Climatic
Zones), while C. acutifolia Turcz. was found in Zone 4 and in North
China.

There must be some relationship between them, as they have similarities
that the systematist recognizes. There is the possibility that the triploids
just mentioned came from a cross in which C. acutifolia was one of the
parents, or that the chromosome number of C. acutifolia had doubled in
number, forming a tetraploid (var. villosula) and then back-crossed with
C. acutifolia or another diploid, giving triploids. Their distribution would
support this assumption.

Cotoneaster lucida Schlecht. was once labeled C. acutifolia Lindl. (not
Turcz.). It came from the Altai Mountains in Mongolia, Zone 4 (North
China). It is quite different from C. acutifolia Turcz. Cotoneaster lucida
has larger glossy and lustrous leaves. It is a triploid. It may be that this
species has some relationship to C. melanocarpa as well as C. acutifolia.

Cotoneaster foveolata Rehd. & Wils. is represented by two vigorous
specimens in the Arboretum grown from the same collection of seeds.
They are alike. Cotoneaster foveolata is a large stiff shrub with large
leaves. It is a triploid with much irregularity in the nuclear divisions of
the pollen mother cells. Pieces of chromatin are left out in the cytoplasm
after the dividing nuclei are formed. Cotoneaster foveolata comes from
central China, Zone 4.

Cotoneaster moupinensis Franch. is a large stiff shrub which, like C.
foveolata, has a rigid appearance. Rehder describes it as “similar to
C. bullata but with black fruit.” It comes from western Szechuan, West
China, at 1300-2000 m. altitude. It is common in woods and thickets. It
is a triploid. Wilson comments about the series as follows: “It must be
confessed, however, that there is a great similarity between all these
black fruited Cotoneasters from China.” He noted that there were only
slight variations between these acutifolia-like species and that they could
be arranged in a gradual series. He believed that since they are all

affinities and find the relations of the acutifolia-like ones.

Cotoneaster bullata Bois and its variety C. bullata var. macrophylla
Rehd. & Wils. are among the most attractive shrubs in the Arboretum in
the autumn because of their large clusters of abundant brilliant red fruits
and very healthy dark green leaves. The macrophylla variety is especially

1954] SAX, POLYPLOIDY AND APOMIXIS IN COTONEASTER 345

striking, having a larger leaf and inflorescence. Wilson says that C. bullata
is a comparatively rare plant, though scattered over wide areas in
Szechuan. It is found in open conifer forests in southeastern Tibet at
9000-10000 feet, inhabiting Zone 5—a zone adjacent to that of C.
moupinensis, which is in Zone 6 in that same area. Cotoneaster bullata
Bois is a triploid. The division of the nuclei of the pollen mother cells
was very irregular, with twenty-two to twenty-eight chromosomes, de-
pending on the looseness of pairing. There were chromosome bridges and
many univalents, up to as many as twelve in one anaphase.

The variety C. bullata var. macrophylla Rehd. & Wils. is also a triploid;
univalents, bivalents, and trivalents are present. It is very irregular, with
chromosome bridges at anaphase.

Another form, C. bullata f. floribunda (Stapf) Rehd. & Wils. is very
attractive. It also has a triploid number with univalents and lagging
chromosomes, but its homologues seem to be more closely paired. Under
his remarks Rehder includes C. moupinensis var. (Stapf) as a synonym.
It seems significant that this form C. bullata f. floribunda was described
as C. moupinensis var. Stapf. It has bright red berries. Cotoneaster bullata
f. floribunda is also found in the thickets of Szechuan, China. It is possible
that C. bullata and varieties came from similar parental ancestry as C.
mou pinensis.

Very bright red berries with intensely green foliage and an attractive
low habit characterize C. apiculata Rehd. & Wils., which grows in West
China. This is a triploid with chromosome bridges and other irregularities
in the reduction divisions in the anthers. In the late anaphase there are
chromosome fragments left in the cytoplasm. Rehder gives C. apiculata
as nearly related to C. disticha Lange, which ranges from the Himalayas
to southwest China. (The latter was not in the Arboretum collection ex-
cept for small cuttings, and it is not included in the counts. Counts on
leaf-tip smears show that it is not a diploid.) There were three plants of
C. apiculata grown from seed collected from the type plant by Wilson in
China. They were alike. George Graves raised twenty-five plants from
seed collected in the Arboretum. All were alike. The same was true of
twenty-nine seedlings grown in the greenhouse, making a total of fifty-
seven seedlings from the same original source.

Cotoneaster Lindleyi Steud. is a triploid. It produced no fruit 1950—
1952, although it had some flowers on it in 1950 and 1951. In 1953 it
flowered abundantly and produced fruit. This fruit became a deep lav-
ender or bluish purple as it developed (much like that of C. affinis var.
bacillaris) and gradually turned a purple brown and finally black —a
word used by Rehder. Rehder includes C. Lindleyi as a species closely
related to C. racemiflora (Desf.) K. Koch. It is a conspicuous species
with its broad oval to broad ovate leaves rounded at the base. It came
from the Himalayas (Rehder, Zone 6). It was very susceptible to fire
blight.

Cotoneaster affinis var. bacillaris (Lindl.) Schneid. is probably a tetra-
ploid. There were seventeen large tetravalent chromosomes in early equa-

346 JOURNAL OF THE ARNOLD ARBORETUM [| VOL. XXXV

torial plate stage and double that number at anaphase. Like C. affinis
Lindl., which is not in the Arboretum, it is said by Rehder (56) to be
closely related to C. frigida.

Cotoneaster frigida Lindl. is a diploid having seventeen chromosomes.
Our specimen of C. frigida is quite different from the upright type de-
scribed as typical. It lies flat on the ground with branches very stiff,
like the one collected at Darjeeling. Some fruit set on the two destyled
branches. The fruit is large and of a bright red color quite unlike thai
of C. affints bacillaris. The latter is found in the Himalayas, as is C.
frigida. They grow in Zone 7 (Rehder).

It is probable that C. affinis var. bacillaris came from the doubling of
the chromosome number in C. frigida or a cross between C. frigida and
some other form. There is quite a noticeable similarity between C. affinis
var. bacillaris and C. lindleyi, which is a triploid. Cotoneaster lindleyi
may have come as a backcross from C. affinis var. bacillaris and a diploid.
There are several diploids in that region.

In writing of the variety C. dielsiana var. elegans Rehd. & Wils., Wilson
remarks on the herbarium sheet that “this western plant shows marked
differences from the type . . . in several respects approaching francheti
Bois. It also has affinity with C. Zabeli Schneider. Its thinner yet more
persistent leaves, smaller pendulous fruit, brick and orange red, distin-
guish this variety from the typical form.” It is found in thickets 3500—
4000 feet on the eastern slope of Likiang snow range, Yangtze Watershed,
Szechuan. These observations are supported by a study of the chromo-
somes.

There are three specimens of C. dielsiana Pritz. in the Arboretum.
One (C. dielsiana 686-33) was from the United States Department of
Agriculture. After C. dielsiana was fixed two years in succession, a differ-
ence in the behavior of the chromosomes was noted for the two years. A
check of the numbers showed that the material had come from the other
two plants the second year. These two plants were grown from seeds
labeled C. dielsiana 134—28-A & B, which were collected by Wilson in
China, and they are alike. The fruit of this specimen is red and waxy
and the leaves have color in the fall. It fits the description by Rehder.
The fruit of the U. S. D. A. specimen is a reddish yellow fruit like that
described for the variety elegans. Cotoneaster dielsiana (686-33) varied
in the chromosome counts, depending on the looseness of pairing of the
homologues. There were univalents, bivalents, and trivalents. Bridges
were present in the anaphase stage. This is undoubtedly a triploid.

The chromosome counts from the specimens grown from the seed from
China (134-28) appear to be 17 and up to about 21 in equatorial plate
stage, depending on looseness of adherence. In the first anaphase the
count appears to be higher than it appears at metaphase, and univalents
and stragglers are apparent. The second anaphase counts are 17 and
higher with irregularities. This may be an aneuploid.

George Graves grew a thousand seedlings from seed collected in the
Arnold Arboretum. They were uniform in habit and growth like the 673

1954] SAX. POLYPLOIDY AND APOMIXIS IN COTONEASTER 347

seedlings from 134-28 grown in the greenhouse. They are undoubtedly
apomicts, as there were many kinds of Cotoneasters growing near

There were five specimens of C. divaricata Rehd. & Wils. growing from
a collection of seeds from China. There were no discernible differences
between these. George Graves grew two hundred seedlings of C. divaricata.
They showed no variation.

Cotoneaster divaricata is a triploid species with lagging chromosomes
and univalents. It is very probably an apomictic triploid.

There are several varietal forms of C. racemiflora (Desf.) K. Koch
(in the Arnold Arboretum) which are quite widely distributed in their
origins and show much variation. They are found in southern Europe,
North Africa, western Asia to the Himalayas and Turkestan. Cotoneaster
racemiflora (Desf.) K. Koch is described by Rehder (56) as ‘‘a variable
species, the typical form variety C. racemiflora var. desfontaini (Reg.)
Zab. (var. typica Schneid.) has generally elliptic acutish leaves, while
variety nummularia Dipp. has broader usually obtuse leaves.”’

The divisions of the nucleus of the pollen mother cells in C. racemiflora
var. desfontaini (Reg.) Zab. are fairly stable in appearance, but there are
some univalents and bridges in both divisions and some chromatin left out
after the daughter nuclei are formed. The pollen mother cells and the
chromosomes are large for the genus and the chromosome count shows
that it is polyploid, probably a triploid, the homologues pairing loosely.

Some fruit set on the destyled pistils in C. racemiflora var. desfontaint,
but only on some of the destyled branches. However, when set, the fruit
was abundant on the destvled branches as well as on the controls.

Cotoneaster racemiflora var. soongorica (Reg. & Herd.) Schneid. from
West China is represented in the Arboretum by two plants from different
sources. They are growing side by side. The fruit of one, which came
from the Framingham Nurseries, is a little darker in color, maturing a
little later than the other, which was grown from seed (Hort. Judd).
They both appear to be triploids, but there are some differences in their
chromosome behavior. In the plant raised from seed there appeared to
be a higher chromosome number. Perhaps the homologues were more
easily separated. One of the obvious phenomena noticed in both cases —
but especially in the plant from the nursery — was the large number of
examples of the stages where the homologues were pulled apart along the
plate but still adhering. The whole complex would be in that condition
with several univalents and bivalents less regularly placed.

Cotoneaster racemiflora var. veitchii Rehd. & Wils. from Central China
possessed the number of chromosomes and the irregularities of a triploid.

Cotoneaster rosea, another triploid, has attractive small pink flowers
in glabrous three- to nine-flowered cymes. It comes from the northwestern
Himalayas and Afghanistan. It is probably a facultative apomict, as it
hybridizes at times with C. acuminata, according to Parker’s notes on
the herbarium sheet. Its branches are slender. It is supposed to be re-
lated to C. racemiflora, as shown especially by its fruit.

Cotoneaster multiflora Bge. is a very graceful shrub from western

348 JOURNAL OF THE ARNOLD ARBORETUM | VOL. XXXV

Szechuan. It is handsome in bloom and it has attractive red fruits. This
is a tetraploid. The counts are around 34, Late anaphase shows a larger
count, which is characteristic, as the two arms of the chromosome separate
further in the reorganization of the daughter nuclei. The divisions are
fairly regular. Although C. multiflora grows in West China, the Arnold
Arboretum Herbarium has also a specimen from Kashmir. This may be
a variety of C. multiflora or of C. racemiflora.

Cotoneaster multiflora var. calocarpa Rehd. & Wils., with rosy-colored
blossoms, shows much irregularity. It is probably a triploid, although at
times it appeared to have too many chromosomes. They may be the
homologues prematurely separated

Cotoneaster multiflora var. granatensis (Bois.) Wenz. is a triploid, the
chromosome counts varying with many univalents—as many as fifteen
in one case. The homologues varied in the ease with which they separated.
In some cases the division went no further than the pollen mother cells.
There was much sterility. The pollen mother cells, as well as the chromo-
somes, were large.

Among the unidentified species Cotoneaster sp. Hesse 13492G is a
triploid. The two unnamed specimens collected by Dr. Rock are also
triploids. They are alike and came from the same collection of seeds.

It was found that two collections labeled C. obscura and collected at
different times yielded different results as far as behavior of chromosomes
was concerned. One of these gave results similar to those obtained from
the material labeled C. obscura var. cornifolia. A survey of these speci-
mens in the field showed the two latter to be similar, and a checking of
the numbers carried by these shrubs against the cards in the files disclosed
that the plants had come from the same collection of seeds! (The large
label on the questionable plant had omitted the varietal name.) Coton-
easter obscura var. cornifolia Rehd. & Wils. is a triploid, as it is very
irregular with univalents, lagging chromosomes, and irregular pollen size.
Cotoneaster obscura is also a triploid, but it has fewer chromosomes and
is somewhat less irregular in its divisions. The homologues may adhere
more closely or it may be an aneuploid. It is not as coarse in general,
having smaller leaves, more delicate branches, and dark red fruit, while
the variety cornifolia has a purple-black fruit. These are both from West
China. They are probably apomictic triploids related to C. acutifolia or a
cross between C. acutifolia var. villosula and some diploid.

Cotoneaster francheti Bois and C. schneideri appear to be identical or
nearly so. The appearance and behavior of the chromosomes first called
my attention to the similarity. When the plants were checked in the field,
they were almost identical. The red fruit of C. francheti Bois was a bit
more deeply colored than that in C. schneideri which was orange-red.
The latter, however, although located near by, is more shaded by other
plants. It could, of course, be an apomict which is a “clonal variety” with
very slight differences. Rehder (56) does not mention C. schneideri. The
plant in the Arnold Arboretum came as seed from California. Cotoneaster
francheti is a tetraploid species. It comes from West China.

1954] SAX, POLYPLOIDY AND APOMIXIS IN COTONEASTER 349

Rehder comments on the similarity between C. francheti Bois. and C.
wardit W. W. Sm. The leaves of C. wardii are larger and in general the
bush is more vigorous. It looks quite different. The fruit is larger and
red without the orange-red tint noticeable in the red of C. francheti. The
chromosome behavior is different. Cotoneaster wardii is a triploid, prob-
ably with C. francheti as a parent. This may be a case where the triploid
is more pronounced in various ways than the tetraploid.

Two plants of C. simonsii in the Arboretum come from different sources;
one from seed from Wageningen, Holland, the other as a plant from a
nursery. They are not exactly alike in chromosome behavior, one being

more irregular with long bridges stretching from pole to pole in the ana-
phase and with many univalents. The other is a little less irregular, but
the chromosome count is the same. Both are triploids according to chro-
mosome count and behavior. The plants vary somewhat, too. The fruits
of one are almost always solitary or in a small cyme; the other fruits
are in a cyme. The one with solitary fruits is much like the variety
newryensis, nearly related to C. francheti. A total of 384 seedlings of
C. simonsti gave three variants.

Cotoneaster zabeli Schneid. is represented by four specimens — all from
the same collection of seeds. They are alike. Cotoneaster zabeli is a trip-
loid. In the third year of the experiment fruit was borne on the destyled
branch

Cotoneaster zabeli var. miniata Rehd. & Wils. is a smaller, more delicate
variety in habit and flower, with a light orange-scarlet fruit instead of
the bright red borne by C. zabeli. It is also a triploid. This, like C.
tenuipes, was expected to be a diploid. It may be an aneuploid, as the
count is low for a triploid.

Cotoneaster tomentosa (Ait.) Lindl. is represented by two plants in
the Arboretum. They are alike. Each year the flowers on several branches
were destyled while in the bud. These set fruit in about the same pro-
portion as on the controls. Apomixis is very definitely shown here. These
plants continue to flower all summer up to frost time; only a few flowers
are in blossom on each branch at a time. Cotoneaster tomentosa is a
triploid.

Rehder (56) speaks of C. tomentosa being similar to C. integerrima,
but larger in every part and more pubescent. Cotoneaster tomentosa is
distributed throughout Europe and West Asia in Zone 4 (Rehder).

One specimen is reported from Kansu, but it may differ and should be
compared, as no other report of that kind is made. Two new specime
from a nursery labeled C. tomentosa are not like the older C. tomentosa
planted here, but are more like C. integerrima, but larger.

It was noted above that two plants of C. obscura var. cornifolia Rehd.
& Wils. coming from the same seed source were alike in every way, the
chromosomes being similar in number and appearance. Several other
groups of two or more specimens from the same collection of seeds from a
species were growing in the Arboretum. In all these cases plants from
the same seed sources were alike. The number was not large enough in

350 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

each case to be certain if this was always true for these species, but
Cotoneasters in general were considered to come true from seeds.

In the behavior of the chromosomes definite groupings are suggested,
and a few of these species are very close together. For instance, C.
francheti and C. schneideri are similar, as are the two so closely related
soongoricas. A note with one specimen of C. francheti in the cultivated
group describes the fruit as orange-scarlet. Schneider collected an orange-
scarlet-fruited specimen at Gotha. Perhaps this is where the name C.
schneideri became attached to and substituted for C. francheti for the
California plant, as C. schneideri does show a slightly more orange tint.
At any rate, C. francheti is a tetraploid which shows a deep orange red
as it is ripening, and only a slight variation could be the cause of the
differences in both cases. Cotoneaster wardii, a related triploid, lacks
the orange tint altogether. Cotoneaster francheti is a tetraploid, which
explains to some extent the “splitters” that Turrill mentions in connection
with the variety “‘sterniana” and which turns up in C. schneideri. Crosses
with other species may help to produce the triploids found in cases like
C. ward, C. nitens, C. divaricata, and others.

REPRODUCTION IN COTONEASTER

Most of the Cotoneasters have long been known to breed true from
seeds. This has been shown by the uniformity in the seedlings grown
from the collections of Wilson and others, and frequent mention of the
fact that certain species of Cotoneaster could be propagated by seeds.
Saunders (60), in Refugium Botanicum, mentions that C. buxifolia comes
true from seeds, and gives a list of Cotoneasters that may be propagated
by seeds. Conrad Loddiges & Sons (41) describe C. affinis and C. melano-
carpa as raised from seeds. According to Stapf’s (66) description of C.
bullata in the Botanical Magazine, 1909, it produces an abundant crop
of seeds to increase the plant

In the literature on Cotoneaster several cases of natural crosses have
been reported. Cotoneaster frigida Lindl. is of interest in this respect. It
has been mentioned as one of the parents in several instances.

Exell (17) collected seeds from a plant of C. frigida growing in close
proximity to other species of Cotoneaster and planted them. Among the
resulting seedlings there were six hybrids; the remaining plants were like
C. frigida. Cotoneaster watereri Exell was considered an_ interesting
hybrid between C. frigida Lindl. and C. henryana Rehd. & Wils. (C. rugosa
Pritzel and Diels.). Cotoneaster frigida Lindl. and C. pannosa Franch.
gave rise to C. crispit Exell. The others were not of horticultural interest.

A. A. Pettigrew (54) of Cardiff obtained seeds of a C. frigida fructi-
luteo plant from Stevenstone, Devonshire. This plant differed from the
type in color of the fruit only, which was yellowish or creamy white
instead of red. After several years of genetic experiments, on which the
final report was not available, the author found red to be dominant, and

1954] SAX, POLYPLOIDY AND APOMIXIS IN COTONEASTER oot

he believed that the individual plants of C. frigida were self-sterile and
that they were fertilized by a red-fruited variety growing in the vicinity.

Conrad Loddiges & Sons (41), in describing C. frigida, mention that
it should be budded on white thorn stock.

Another instance of the tendency of C. frigida to hybridize is the an-
nouncement, in Gardeners’ Chronicle 132: 243, of C. cornubia —a hybrid
between C. frigida and an unrecorded species — as a vigorous plant by
Mr. Lionel de Rothschild in 1933. This received an award of merit.

Others report C. frigida as breeding true. A. T. Johnson (40) describes
C. frigida as naturally a free and robust tree “so readily raised from
seeds.” He also mentions the distinct form fructo-luteo. Four seedlings
of C. frigida montana grown from seed obtained by Donald Wyman from
North Africa were alike.

On a note with a specimen from the Flora of Chumbi in the Arnold
Arboretum Herbarium, mention is made of the fact that seedlings of C.
frigida Lindl. are alike; also that seedlings of C. aldenhamensis are all
alike but not so broad-leaved as those of frigida.

R_N. Parker, while collecting in Khadrula India Sikkum, Himalayas.
at an altitude of 2700 meters, June 18, 1928, found in a thicket a specimen
which he considered a cross (C. acuminata Lindl. and C. rosea Edgew.).
There is a specimen of this in the Arnold Arboretum Herbarium. This
has the habit of C. acuminata and the pink flowers of C. rosea. A plant
of C. rosea was growing near by the thicket of C. acuminata where this
hybrid plant was found.

Caution against raising plants from seeds comes from several sources,
concerning both the microphylla and salicifolia groups. Of the probable
hybridization in the several species allied to C. microphylla, C. congesta
(Syn. C. microphylla var. gracilis), and C. thymifolia Arthur Osborn (51)
writes, “It is best to propagate these distinct forms by cuttings or layers.
They do not come true. At least that is our experience at Kew, though
ig may be due to cross pollination as all are growing in close proximity.’

_ J. Bean (3) made similar observations, as was noted earlier. The
Meee of species, all natives of the upper and middle basin of the Yangtze
River, in which are included C. salicifolia and its varieties, C. glabrata,
C. henryana, and C. rhytidophylla, have also been reported (66) to hy-
bridize easily. Cotoneaster watereri is supposed to have one parent in
this group.

The instances of hybridization are few and limited to species or groups
where few specific data are available. In an isolated population a species
would be expected to breed true, but when numerous species are growing
in close proximity variation would be expected. Some cases have been
cited where a small number of offspring from the same seed source are
alike when species overlap and hybridization could result. The most
critical test, however, is the study of the progeny of species grown in a
botanical garden where many species are grown in close proximity.

In the Arnold Arboretum most of the species are grown in close prox-
imity., and the time of flowering is essentially the same in some species

352 JOURNAL OF THE ARNOLD ARBORETUM [| VOL. XXXV

and overlapping in others. There is ample opportunity for cross-pollina-
tion, since the bees at blossoming time are so abundant that it is difficult
to work with the plants.

In order to determine whether most of the species breed true from
seeds where there was so much chance of cross-pollination, seeds were
collected from the following species, stratified, and grown in the green-
house: C. acuminata, C. acutifolia var. villosula, C. adpressa, C. adpressa
hessei, C. adpressa var. praecox, C. ambigua, C. apiculata, C. bullata, C.
bullata {. floribunda, C. dielsiana, C. divaricata, C. foveolata, C. frigida,
C. horizontalis, C. integerrima, C. lindleyi, C. lucida, C. microphylla, C.
moupinensis, C. nitens, C. obscura cornifolia, C. racemiflora var. desfontaini,
C. racemiflora var. soongorica, C, racemiflora var. veitchii, C. rosea, C
simonsii, C. s. var. newryensis, C. tomentosa and C. wardii.

In connection with this part of the work I wish to thank Dr. Karl Sax,
who helped collect the seeds, and Mr. Lewis Lipp, Jeanette Renshaw and
Dorothy Thorndyke for growing and caring for the seedlings. I thank
George Graves, who grew additional seedlings from three of the same
seed sources and furnished the data that are also included in the table
(Table II). These include 25 seedlings of C. apiculata, 1000 of C.
dielsiana, and 200 of C. divaricata.

TABLE II
APOMIXIS IN COTONEASTER

Chromosome Number of | Number of Number of

Species Number Seedlings Maternals Variants
C. acuminata 2n 11 7
C. acutifolia var. villosula 4n 14 12 2
C. adpressa 3n 6 6 0
C. adpressa hessei 3n 10 10 0
C. adpressa var. praecox 3n 4 4 0
C. ambigua 3n 50 50 0
C. apiculata 3n 54 54 0
C. bullata 3n 14 14 0)
C. bullata f. floribunda 3n 34 34 0
C. dielsiana 3n 1673 1673 0
C. divaricata 3n 200 200 0)
C. frigida 2n 2 0 2
C. horizontalis 3n 5 5 0
C. integerrima 3n 44 44 0
C. lin i 3n 16 16 0
C. moupinensis 3n 21 21 0
C. multiflora 4n 57 57 0
C. ni 3n 26 26 0
C. obscura var. cornifolia 3n 16 16 0
C. racemiflora var. veitchii 3n 13 13 0
C. rosea 3n 27 27 0
C. simonsii 3n 384 381 3
C. wardii 3n 36 35 1

1954] SAX, POLYPLOIDY AND APOMIXIS IN COTONEASTER ae

The data for this work are given in Table II. In most instances the
number of seedlings from each species was not large, but most of the
offspring were like the seed parent; most were maternals.

Cotoneaster acuminata and C. frigida showed a tendency to hybridize.
They showed considerable variation in their respective progeny. Both are
diploids.

The large majority of the Cotoneasters breed true. Several others did
show a small percentage of variants.

Among the triploids, three of those propagated threw a small percentage
of variants (Table II). Of the 26 seedlings of C. mitens, three were
different and varying among themselves; the others were alike. One of
the thirty-six seedlings of C. wardii varied from the type. Of 384 seedlings
of C. simonsti all were uniform, with three exceptions. As was also true
of the aberrants in C. nitens and C. wardit, the three seedlings of C.
simonsti were conspicuously different from the others of the same species.
They were striking among an otherwise uniform population. They had
broader leaves which were thicker and tomentose and flattened into a
rosette.

This may indicate that although C. nitens, C. simonsii, and C. wardii
reproduced apomictically, the egg is fertilized at least at times, and that
they may be crossed with other species. They are probably facultative
apomictic species, as Malus sargenti is among the a ;

On the triploid C. tomentosa (Ait.) Lindl. destyled ovaries developed
in about the same proportion as on the controls on other branches of the
same shrub. This was also true of the destyled ovaries of the triploids
C. dielsiana Pritz., C. nitens Rehd. & Wils., C. froebellii—a plant from
Vilmorin — and C. zabeli Schneid., but further work along this line would
be necessary before settling this point on these grounds, except for a few
species. Many species so treated gave negative results, but most of the
negative results are not conclusive, either because not enough work was
done, or because injury in the process might cause the fruit to drop (as
happened most often the first year) or because no fruit was set on the
controls at the same time.

Apomixis must be very general among the triploid Cotoneasters. Many
species grown in the same vicinity are in bloom at the same time. The bees
and other insects are attracted to them in such large numbers that cross-
pollination should be the rule. Although the divisions in the pollen mother
cells were very irregular in the triploids, and often with the loss of
chromatin, there was an abundant production of fruit and a great uni-
formity among the progeny where they were tested. This seems to
establish rather definitely the presence of apomixis in the triploids.

The tetraploids were more regular in the divisions in their pollen mother
cells. Seedlings of only two tetraploids, C. acutifolia var. villosula and
C. multiflora, were grown. Among fourteen seedlings of the former two
variants appeared. All were uniform in C. multiflora. ee fruit set
on the plant, which was in the midst of many other s

There is no doubt about apomixis in the triploid Co aeatiars and it

354 JOURNAL OF THE ARNOLD ARBORETUM | VOL. XXXV

is possible that it may exist in some of the diploids and tetraploids. There
is also some evidence that some of the triploids are facultative apomicts,
since three produced a small percentage of variant seedlings. It is possible
that variants in other triploid species might be found if larger numbers of
progeny were grown or if the triploids were hand-pollinated with pollen
from triploids.

The meagre evidence of facultative apomixis in Cotoneaster is sup-
ported by the breeding behavior of the closely related genus Malus. Malus
sargenti is found in diploid, triploid, and tetraploid forms (K. Sax un-
published). When open pollinated all produce only maternal progeny,
even though they are surrounded by other species. When artificially pol-
linated with pollen from other diploid species, the triploids and tetraploid
M. sargenti forms do produce some sexual progeny, but seldom more than
twenty per cent. In some cases only maternal types of progeny are pro-
duced, and in all cases these types are predominant. The mechanism of
this facultative apomixis is unknown. A few triploid species of Malus
do produce an occasional variant when open pollinated, even though the
great majority of the progeny are maternals. It is possible that the same
type of apomixis is present in Cotoneaster.

DISTRIBUTION AND RELATIONSHIPS

The geographical distribution of Cotoneaster in relation to the 2 n.
3 n, and 4 n chromosome numbers may offer suggestions as to the origin
of some of the species. Eight of the species studied are diploids. There
may be other diploids among those in which the chromosome number has
not been determined. The diploids C. frigida, C. microphylla, and C.
acuminata are found in the Himalayas; C. conspicua in Tibet: C. glabrata
and C. henryana in the Yangtze basin in western and central China: C.
dammeri in West Hupeh in central China: and only one, C. acutifolia
Turcz., in North China and Mongolia.

The diploids have a limited geographical distribution, and fifty per
cent of them are found in Zone 7 (Rehder); only one, C, acutifolia Turcz.,
inhabits Zone 4. Wilson (75) comments that C. acutifolia Turcz. was
originally described from Chinese Mongolia, that it was introduced by
Dr. Bretschneider from mountains near Peking, and that the typical form
is absent from central and western China, but varieties are found growing
in these regions.

There were a number of instances in which specimens were labeled C.
acutifolia or a variety thereof. Some of these have been considered as
natural variations. Some relatives of C. acutifolia Turcz. have been
recognized, but the situation was complex, and much has been left to be
settled in the future.

A variety, C. acutifolia var. villosula Rehd. & Wils., was recognized
as a near relative. A study of the chromosome number proved it to be a
tetraploid form. It is a hardy shrub occupying Zone 5 in central and
western China. It grows in thickets in West Hupeh, West Szechuan, and

1954] SAX, POLYPLOIDY AND APOMIXIS IN COTONEASTER 355

in the Shensi mountains, the latter bordering on the habitat of C. acuti-
folia Turcz. It may have originated through the doubling of the chromo-
some number of C. acutifolia under circumstances in the past conducive
to the disturbance of the chromosome balance. Two specimens labeled
Cotoneaster sp. grown in the Arnold Arboretum from seed collected by
Dr. Rock in the Tao River Basin in western China were triploids.

A large number of species which are considered as belonging to the
“acutifolia group” are triploids or aneuploids with approximately the
3 n number. Included in this category are C. ambigua Rehd. & Wils..
which the authors considered closely allied to C. acutifolia Turcz., and
C. obscura Rehd. & Wils., which they believed resembled C. acuminata
which grows in the Himalayas in India. They thought C. obscura also
resembled C. foveolata Rehd. & Wils. of West Hupeh, which seems more
probable, although there may have been earlier connections with the
acuminata ancestors. Cotoneaster foveolata Rehd. & Wils. in turn closely
resembles C. moupinensis Franch., common in the thickets of West
Szechuan. These were black-fruited. Cotoneaster bullata Bois. and its
varieties are red-fruited, and when not in fruit were mistaken for C.
moupinensis. Further comments show close external resemblance. Coton-
easter bullata Bois., according to Wilson, is a relatively rare plant but
is widely scattered along the edge of the zone occupied by C. moupinensis
in West Szechuan.

At first Rehder and Wilson considered C. tenuipes Rehd. & Wils. as
nearly related to C. racemiflora Koch (which is on the other side of the
mountains), but they lacked the flowers. Later Rehder placed it near
C. acutifolia Turcz., which seems more probable, as these forms are all
found in West China.

These triploids that resemble C. acutifolia Turcz. may have come from
the diploid C. acutifolia and the tetraploid C. villosula. There is also the
possibility of intercrossing of C. villosula and its descendants with other
diploids of the region, as is obvious from the geographical locations and
the fact that C. acutifolia var. villosula is a facultative apomict.

Cotoneaster acutifolia Turcz. may also have contributed some of its
characteristics to the flora of the North. Cotoneaster lucida Schlecht.
was once named C. acutifolia Lind]. It is not easily confused with C.
acutifolia Turcz. It has dark green leaves which are very glossy and
lustrous. It has also been found to be a triploid. It grows in the Altai
mountains in Mongolia in Zone 4 (Rehder), in thickets at 2000-3000 m.,
having a limited distribution. This is probably related to C. acutifolia
Turcz, and may have another relative in C. melanocarpa Lodd.

Cotoneaster melanocarpa Lodd. is a tetraploid species which ranges
from northern and eastern Europe across Siberia to central and northeast
Asia. It is probable that it is another tetraploid form from C. acutifolia
Turcz. (or even C. acuminata, which I doubt) which, having less competi-
tion to the north as well as the necessary vigor and better adaptability,
has occupied more space than most Cotoneasters. It inhabits Zone 4.

Varietal forms of C. melanocarpa Lodd. occupy more restricted areas.

356 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

The variety C. melanocarpa var. commixta Schneid. is a 4 n species and
C. melanocarpa var. laxiflora (Lindl.) Schneid. is a 3 n species from cen-
tral Asia. It may have been a back cross of C. melanocarpa with C.
acutifolia or a cross with C. acutifolia var. villosula.

Other diploids have contributed variation to the flora around them.
Cotoneaster acuminata Lindl. is a diploid found in Zone 5 in the Hima-
layas and on the side away from China. According to Wilson (75) C.
acuminata Pritzel in Bot. Jahrb. 29: 385. 1900 (pro parte, non Lindley)
is under C. acutifolia var. villosula as a synonym. There are different as to
chromosome number and they are geographically separated.

Although they are distinct species, there is some resemblance and a
fairly close relationship taxonomically between C. acutifolia Turcz. and
C. acuminata, They may have some common or similar ancestry in the
past; at present there are very definite geographical barriers between the
species.

Cotoneaster frigida Lindl., a diploid, grows in the Himalayas and is
cultivated in Zone 7. Rehder (56) considers C. affinis and its variety
C. affinis var, bacillaris, a tetraploid cultivated from Zone 7 (?) to be
closely related to C. frigida. Bean (3), in his ‘“‘Trees and Shrubs,” re-
marks that C. affinis var. bacillaris and C. frigida are connected by one
or two intermediates. In 1899, W. J. Bean writes in Vol. 55 of The Gar-
den that C. affinis is between C. bacillaris and C. frigida. The tetraploid
C. affinis var. bacillaris may have been formed by a cross between two
species, one of which was C. frigida, or by the doubling of the chromosome
number of C. frigida.

Cotoneaster lindleyi, a triploid, also shows affinity for this group,
especially C. racemiflora. It is an inhabitant of the Himalayas, and is
cultivated in Zone 6 (?).

Cotoneaster racemiflora (Desf.) K. Koch is considered by Rehder a
very variable species. It is a triploid which ranges in Zone 4 from south-
ern Europe, North Africa, and western Asia to the Himalayas and Turk-
estan. It has several varieties, var. desfontaini (Reg.) Zab. being the
typical one, and it is a triploid. It is not found in China, but some of its
varieties are found there and are more limited in distribution. Cotoneaster
racemiflora var. soongorica (Reg. & Herd.) Schneid. inhabits West China,
Zone 3, and C. racemiflora var. veitchii central China. These are triploids.

If the species in China are closely related to the species to the west,
then a common ancestor must have given rise to those on both sides of
the mountains. Its wide range makes C. racemiflora look like an older
species. It may have come from C. frigida, at least as one of the parents,
in which there was a doubling of the chromosomes, as is probable in C.
affinis var. bacillaris, or it may have come from a cross of C. frigida with
C. acuminata or an unknown or extinct species. It is apparent that there
is a decided difference between the Chinese racemiflora species and the
typical form.

That the C. racemiflora triploids which were grown in close proximity
with other Cotoneasters breed true shows that apomixis is fairly common.

1954] SAX, POLYPLOIDY AND APOMIXIS IN COTONEASTER 557

The seed set is very good. The wide range of these triploid apomicts
shows ability to adapt to a variety of locations. Cotoneaster rosea shows
resemblances to C. racemiflora. It may show both C. acuminata and C.
frigida traits. It is a triploid

Cotoneaster simonsti, first collected in the Khasia Mts. in northern
India, shows some resemblance to C. acuminata and to C. rotundifolia.
There is a possibility that in the past it had some common ancestry with
the francheti group — to which it is often likened.

Several of the most attractive species of Cotoneaster which show various
degrees of similarity inhabit West and Central China. They differ from
the acutifolia group. Among these are C. francheti, a tetraploid, and the
triploids C. dielstana, C. dielsiana var. elegans, C. divaricata, and C. nitens.
Cotoneaster wardii, a triploid facultative apomict inhabiting Tibet, is
very closely related. Cotoneaster dielsiana is an apomict. Cotoneaster
zabeli, which inhabits central China, appears to have some similar traits.
It and its variety are both triploids

The origin of this group is not clear. They are all related to the tetra-
ploid C. francheti Bois. The diploid C. dammeri grows in central China,
C. conspicua in W. China. There are also possibilities of having earlier
species mixed with ancestors of species from the southern Himalayan
groups. Cotoneaster microphylla extends into Yunnan. Further study of
the francheti group is necessary in order to determine its origin.

The salicifolia-like group of plants from the Yangtze Basin have been
a problem to the systematist. Cotoneaster salicifolia Franch., its varieties
floccosa (Pritz.) Rehd. & Wils. and rugosa Rehd. & Wils., C. rhytidophylla
Rehd. & Wils., C. glabrata Rehd. & Wils., and C. henryana (Schneid.)
Rehd. & Wils., are all evergreen or half evergreen species with prominently
veined, elliptic oblong to ovate lanceolate leaves. They have attractive
flowers and fruit.

Exell (17) remarks, ‘I am following the conclusions of Dr. Stapf (Bot.
Mag. ¢t. 8999) that C. henryana Rehd. & Wils. and the earlier C. rugosa
Pritz. ex Diels are synonymous.’ These plants do resemble each other
closely, but Rehder & Wilson (56) recognized some differences, and the
study of the chromosome numbers indicates that they were correct.
Cotoneaster glabrata and C. henryana are diploid species (Rehder Zone
7 2). Cotoneaster salicifolia comes from Zone 6 (?). Its chromosome
numbers have not been determined with certainty. Its variety C. salictfolia
var. rugosa is a triploid from Zone 5.

It may be noted from the zones in which these species are found that
they are fairly tender plants. Except C. rhytidophylla Rehd. & Wils. they
are all growing in the Arnold Arboretum (in fairly protected areas) in
Zone 4.

It is not surprising that Thomas (72) remarks, “If these plants are
raised from seeds variation results.” There are some diploids, C. glabrata
and C. henryana, and some triploids. There may be facultative apomicts
in some of this group.

Included by Rehder as a nearly related species is C. glaucophylla

358 JOURNAL OF THE ARNOLD ARBORETUM [| VOL. XXXV

Franch. inhabiting West China (Zone 7) which is also a triploid. He calls
attention to some resemblance to C. zabeli. Cotoneaster zabeli is a trip-
loid from central China. A variety, C. zabeli var. miniata Rehd. & Wils..
is a triploid from central China.

The microphylla group, mostly prostrate plants, inhabits the Himalayas.
E. H. Wilson (75) remarks that it is not seen in China, although it has
been reported from Yunnan, These plants are reported to show variation
in their seedlings. C. microphylla is a diploid. There are several micro-
phylla varieties with closely related species, as C. congesta Bak. (Hima-
lavas). Rehder mentions C. conspicua Marquand, a diploid, as a closely
related species. It inhabits West China and Tibet.

It may be noted from the foregoing that the triploids vary more as to
range and habitat. Some of them are as limited as the diploids, being
reported from only one vicinity. Nineteen out of about forty triploids
are reported from West China, four from the Himalayas, and six from
Central and West China. Four more are described from Central China.
two from North China, and one, C. multiflora granatensis, from Spain.
All of these show a narrow range.

There are a few more widely ranging triploid species. Cotoneaster in-
tegerrima Med., an early recognized Cotoneaster, is a triploid which is
common to Zone 5 in Europe, in the western Himalayas, and in northern
Asia to the Altai Mountains in Mongolia. It has been reported from
southeast Tibet and Shantung, far from any others that have been re-
ported. Rehder collected it for the Arnold Arboretum in the Savoyan
Alps in France, and Anderson collected it in the mountains in Bulgaria.
Rehder considers C. uniflora Bge., growing in the Altai Mountains, as a
closely related species, but comments that it may be only a variety of
C. integerrima. Cotoneaster sylvestris of central China is also mentioned
as a Closely related species. Cotoneaster zabeli of western China shows
some characteristics of C. integerrima.

Cotoneaster tomentosa (Ait.) Lindl. is somewhat more limited in its
distribution. It grows in northern and eastern Europe and western Asia.
It has been reported from Kansu. It is considered related to C. integer-
rima, but it is larger and more intense in every detail. It is a vigorous
apomictic triploid. Cotoneaster tomentosa leaves are more like those of
the affinis group but heavier and more tomentose; these species differ in
fruit and inflorescence.

Another widely distributed triploid, C. racemiflora, is found growing
from North Africa throughout southern Europe and western Asia to the
Himalayas and Turkestan. It is reported at Tomsk in Siberia. J. F.
Rock remarks that the typical C. racemiflora (Desf.) K. Koch is not
found in China, but well-marked varieties are found in Hupeh and
Szechuan. The McClaren collectors found C. racemiflora var. veitchii in
the Purple Mountains near Nanking. The variety C. racemiflora var.
soongorica has been collected in West China, Shantung, and Shansi.

Of the large number of triploids only two, C. glaucophylla and C. wardit,
are found in Zone 7 (both questioned) ; six are in Zone 6, and C. racemi-

1954] SAX, POLYPLOIDY AND APOMIXIS IN COTONEASTER 359

flora var, soongorica is in Zone 3. The remaining triploids in which the
aa numbers were determined were either varieties or unknown
speci

The tetraploids, which are few in number, also vary in the breadth of
their distribution. Cotoneaster acutifolia var. villosula is a tetraploid
from central and western China in Zone 4. Cotoneaster affinis var. bacil-
laris inhabits Nepal (Zone 7), C. francheti western China (Zone 6 ?).
Cotoneaster melanocarpa ranges in Zone 4 from northern and eastern
Europe and the Caucasus to central Asia (having been reported from
Tomsk), from Turkestan through Siberia to northeast Asia in Mongolia,
Manchuria, Chili and Kansu. The variety C. melanocarpa var. commixta
has a more limited range. Cotoneaster multiflora is found in West China.
Cotoneaster melanocarpa has the widest range of the Cotoneasters except
the triploid C. racemiflora.

Some of the polyploids show a tendency to adapt themselves over a
wider range geographically. The diploids are limited in their range. There
is also a slight tendency for the diploids to be more limited zonally, about
half being in Zone 7.

All European species so far studied are polyploid. They probably
originate from the Himalayan species. No diploids are found in Europe,
and all European species show resemblances to those of central and
northern Asia. Cotoneaster multiflora var. granatensis from Spain is sup-
posed to be related to C. multiflora from western China. It might.
however, be related to C. racemiflora, which is not so far apart from C,
multiflora in distinguishing characteristics. Its leaf pubescence is char-

acteristic of C. racemiflora, being one of the characters separating the
two species.

When the chromosome number and the tendency to be evergreen were
considered at the time of leaf drop there was found to be a slight tendency
on the part of the 2 n species to be more evergreen and therefore more
subtropical. There were deciduous diploids, C. acutifolia and C. acumi-
nata. There were many polyploid evergreens. Some of the plants, both
diploid and polyploid, were partially evergreen.

VARIATION AND EVOLUTION

Cotoneaster has long been recognized as a taxonomically difficult genus.
Wilson (75) in discussing the similarities of the black-fruited Cotoneasters
from China wrote “with C. acutifolia Turcz. at one end of the chain and
moupinensis Franchet at the other, it is almost possible with the material
before us to connect the series.”? As in Hieracium, Crataegus and Rubus,
the species and varieties are often separated by minute, but constant
differences.

Hybridization is still a prominent factor in the variation and evolution
of the Cotoneasters. There is clear evidence of hybridization of the di-
ploid species, both in nature and under cultivation. There is some indica-

360 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXXV

tion of crossing between a few of the tetraploids. It is also probable that
many, if not most, of the triploids were derived from crosses between dip-
loids and tetraploids. Even some of the apomictic triploids may occa-
sionally hybridize with diploids, as is known to be the case in Malus.

Although the great majority of the species of Cotoneaster are apomictic
triploids, there are some variants among the progeny of some species.
Such variation could be caused by occasional sexual reproduction, or it
could be the result of internal segregation if the progeny are from un-
reduced egg cells, as Haskell (34) has suggested for apomictic species of
Rubus. The meiotic irregularity and loss or non-disjunction of chromo-
somes could also lead to variant aneuploids, as has been suggested by
Stebbins (68). There is some evidence of aneuploidy in a few Coton-
easters, although exact chromosome counts were not possible.

In the triploids the meiotic divisions are irregular, as may be expected.
but some species are more irregular than others, with bridges found at
anaphase. Although the division process is typical for each species or
variety, there is much variation between the speices, and due to various
degrees of adherence of the homologues, the exact chromosome number is
sometimes uncertain. Some of the speices may be aneuploid with slight
differences in chromosome number; some of the chromosomes may be
lacking, but this has not been established.

Perhaps these more irregular triploids are from crosses between dis-
tantly related diploids and tetraploids which differ in chromosome struc-
ture and result in inversion bridges in the triploid hybrids. The more
regular triploids may be autopolyploids between closely related forms.
At any rate, the triploids do seem to fall into several rather distinct groups
in respect to meiotic behavior.

Polyploidy within the species would also account for some variation
since it is known that induced tetraploids are often of a stiffer habit and
have larger and more deeply colored flowers than their diploid ancestors.
It is just this type of variation that differentiates many of the species and
varieties of Cotoneaster.

The variation produced by hybridization and polyploidy is “fixed” by
apomixis. Although, as Miintzing and others have found, apomixis is
not confined to polyploids, it does permit cytologically and genetically
heterozygous species to be perpetuated. Slight variations can be main-
tained in “Clonal Species,” a term proposed by Darlington (10).

Apomixis in polyploids permits the constant reproduction of a hybrid
which would normally be sterile and would have little survival value.
Yet it is known that many artificially produced triploids which must be
propagated by grafts or cuttings are superior to either the diploid or tetra-
ploid parent in vigor and in horticultural value (K. Sax unpublished).
The fact that most of the species of Cotoneaster are apomictic triploids
shows that they have high survival values and can be easily propagated.

The polyploid species of Cotoneaster do have a wider range of distribu-
tion than the diploids. The diploids are largely confined to the Himalayas,
West and Central China, while some of the triploids and tetraploids are
widely distributed. The triploids C. racemiflora, C. integerrima, and C.

1954] SAX, POLYPLOIDY AND APOMIXIS IN COTONEASTER 361

tomentosa have occupied a wide territory. Cotoneaster lucida, although
limited in distribution, has adapted itself to a colder zone. The tetra-
ploid C. melanocarpa has spread over large areas from northern and east-
ern Europe through Siberia and Manchuria. These widely distributed
species are polyploid apomicts.

Excessive uniformity brought about by apomixis might be a handicap,
even though under original conditions the triploid or hybrid apomictic
species may have definite advantages and greater adaptability. As Stebbins
(68) has suggested, the uniformity imposed by apomixis would be a
handicap in a changing environment, and the lack of variation would be
a block to further evolution.

In Cotoneaster, however, there is occasional variation in the progeny
of apomictic species. These species still have some plasticity to meet a
changing environment, either through occasional sexual progeny, internal
segregation, or chromosome irregularity. Thus they have the ability to
perpetuate an unbalanced cytological and genetic complex of adaptive
value and yet have the capacity to produce variants to meet new environ-
mental conditions.

Polyploidy and hybridization provide most of the variation in Coton-
easter. Often the differences are slight, yet they are fixed by apomixis,
providing distinct variants which reproduce true to type. Some of these
may have a wide distribution. It is this complex of variation due to
polyploidy and hybridization, combined with apomixis, that makes the
genus Cotoneaster a difficult one for the taxonomist. For the horticulturist
apomixis is an advantage, since most of the ornamental species and
varieties breed true from seed.

SUMMARY

The chromosome numbers were determined in forty-one species and
eighteen varieties of Cotoneaster. Of these Cotoneasters eight taxa were
diploid, forty-three triploid, six tetraploid, and two polyploids could not
be determined. Some aneuploidy may be present, as several triploids
seem to vary.

Progeny from twenty-three species show that two diploids reproduced
sexually. The rest reproduced apomictically, an occasional variant ap-
pearing among three of the nineteen triploids: C. nitens, C. simonsii, and
C. wardii. One tetraploid was a facultative apomict.

The diploids are limited in their geographical distribution, as are most
of the triploids and tetraploids. Among the triploids three species, C.
integerrima, C. racemiflora, and C. tomentosa, show wide distribution.
One tetraploid, C. melanocarpa, is found over wide areas. The diploids
showed a slight tendency toward zonal, as well as geographical limitation.

Apomixis stabilizes a species, but the genus Cotoneaster is given flexi-
bility through sexual reproduction and through facultative apomixis.
There are probably some obligate apomicts, i.e., C. dielsiana.

Apomixis and polyploidy are responsible for the taxonomic complexity
of the genus Cotoneaster.

362

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JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

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JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxv

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~I
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1954] SAX, POLYPLOIDY AND APOMIXIS IN COTONEASTER 365

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Charles Sprague Sargent.

THE DIRECTOR’S REPORT ON THE ARNOLD ARBORETUM
DURING THE FISCAL YEAR ENDED JUNE 30, 1954

The fiscal year of 1953-54 was a momentous one for the Arnold Arbo-
retum, marking as it did the solution of long-standing problems of space,
housing and care of collections which have for over a decade hindered
the work of the Arnold Arboretum.

On January 19, 1953, the President and Fellows of Harvard College
(the trustee of the Arnold Arboretum), in a detailed and careful vote,
stated that “In the exercise of its duties as trustee of the endowment funds
held in trust for the purposes of the Arnold Arboretum, the Corporation
determines that it is in the best interests of the Arboretum and will
promote the purposes of its endowment to remove to a central building
in Cambridge the main body of the library and herbarium of the Arbo-
retum related to research, and to retain in Jamaica Plain such books and
specimens as may be required to provide there a working library and
herbarium.” This proposal had long been advocated as a solution to the
problems of the Arboretum which previous directors have stated in their
annual reports. (See Annual Reports 1936, 1938, 1940, 1941, 1942 and
following.) The Administration Building in the Arboretum had become
overcrowded. Floor space was no longer available for the addition of steel
cases, and as temporary expedients, display cases and cardboard herba-
rium boxes were used to store approximately 20% of the mounted her-
barium and all of the material awaiting determination and mounting.
The library was equally crowded to the point where it was becoming
difficult to care for the valuable books properly. This latter condition
had been alleviated by transferring, for use and storage, many of the
Arboretum books to the Harvard Forest and to Cambridge. However,
even with these measures, additional shelf space could not be found in
the Administration Building. Working space for staff members had be-
come restricted by the growth of the collections and the library. To solve
these and other problems, the Corporation vote was welcomed by a
majority of the staff of the Arboretum.

The vote of the Harvard Corporation called for the appointment of
an Arnold Professor, a post unoccupied since the retirement in 1946 of
Dr. E. D. Merrill. In October the Corporation, with the approval of the
Board of Overseers, appointed Richard Alden Howard as Arnold Professor
and Director of the Arboretum. Dr. Howard was also appointed Professor
of Dendrology within Harvard College. All appointments were effective
February 1, 1954.

The Harvard Corporation had voted that a portion of the herbarium
and library was to be moved to a new building in Cambridge known as
the Harvard University Herbarium Building. One million dollars was as-
signed from unrestricted funds of the University for the construction and
equipping of this building. The Arboretum collections and library would
thereby receive adequate housing and the staff working equipment at no

368 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXXV

capital cost to the Arnold Arboretum endowment. Construction of the
building was completed in February 1954.

The Arnold Professor, by vote of the Corporation, was to implement
the move. Serious considcration was then given by the director and the
staff of the Arboretum to the selection of books and herbarium specimens
to be retained in Jamaica Plain. When tentative plans were formulated,
Dr. Howard, in accordance with the vote of the Corporation, consulted
with Dr. William J. Robbins, Director of the New York Botanical Gar-
den, regarding what specimens were to be left in Jamaica Plain as a work-
ing herbarium. He also consulted with Dr. Keyes Metcalf, Director of
the Harvard University Libraries, regarding what books were to be re-
tained in Jamaica Plain. Each of these conferences produced a final plan
which determined the nature of the Arboretum’s collections of specimens
and books in each of the two locations. Dr. Robbins and Dr. Howard
determined that the vote of the Harvard Corporation could be best im-
plemented by establishing in Jamaica Plain a working herbarium devoted
to the study and advancement of knowledge of horticultural plants. To
that end they made seven recommendations as follows:

=

That all cultivated specimens in the herbarium of the Arnold Arboretum
representing taxa now under cultivation in the Arboretum and elsewhere
be retained in Jamaica Plain.

That sufficient additional specimens of taxa related to a given cultivated
group be retained to insure working control of that group.

That spontaneous or native specimens be used to supplement poorly
represented cultivated taxa.

bo

ww

as

That representative specimens of all taxa of certain major cultivated groups
be retained. For example, it is proposed to have in Jamaica Plain all
material of the difficult genus Crataegus, and to have representatives of all
taxa of the Conifers, Rosa, Rhododendron, Cotoneaster, Quercus, Syringa,
Viburnum, Lonicera, Fraxinus and Ulmus.

mn

. That additional specimens be returned to the herbarium at Jamaica Plain
as their need becomes known; i.e., new introductions, new cultigens, or
additional material found necessary for determination.

That this herbarium be maintained as an active herbarium and that addi-
tional cultivated material, e.g. herbaceous specimens, be added in future
years to make the herbarium at Jamaica Plain a complete working herba-
rium for determination and study of all groups of cultivated plants.

~

That all type specimens of cultivated taxa remain in the Jamaica Plain
herbarium and be clearly marked and properly housed.

The specimens to be retained in Jamaica Plain number approximately
100,000 specimens and are housed in 96 steel herbarium cases, creating
in itself one of the outstanding horticultural herbaria of the wor

A similar set of proposals was worked out by Dr. Metcalf and Dr.
Howard regarding the library books to be retained in Jamaica Plain.
Again, in accordance with the vote of the Harvard Corporation and for
the best interests of the Arboretum, Dr. Metcalf and Professor Howard

1954] THE DIRECTOR’S REPORT 369
recommended the establishment of a working library devoted to the study
and advancement of knowledge of horticultural plants. To that end they
recommended to the Corporation the following proposals:

_

_ That the library in Jamaica Plain retain a complete set of all publications
of the Arnold Arboretum; i.e., the serial publications Arnoldia, the Journal
of the Arnold Arboretum and Sargentia, as well as the complete sets of
publications by staff members. All Sargent, Wilson and Rehder publications
will be kept in Jamaica Plain.

_ That reference books for horticultural study and research are to be re-
tained. These will include basic textbooks, floras, garden dictionaries, horti-
cultural encyclopedias, and all books which normally fall in the library
classification of horticulture, botanic gardens, etc.

_ That a selection of books on dendrology, ecology, physiology, pathology,

economic botany, entomology, biography and natural history, as well as

books on color will remain in Jamaica Plain.

To coordinate the interests of the herbarium with those of the library,

there will be in Jamaica Plain all books dealing with roses, rhododendrons,

lilacs, and similar groups, as well as monographs of Crataegus, Quercus,
and the Conifers, and other horticultural books and monographs which will
strengthen the value of the horticultural herbarium.

ia)

Ww

a

On

Additional periodicals of particular value to horticultural research and
reference. such as Curtis’s Botanical Magazine, Horticulture, Gartenschon-
heit, La Belgique Horticole, Revue Horticole, Gardener’s Chronicle, Journal
Royal Horticultural Society, etc., will be retained.

S

Folio volumes useful to horticulture are to be left in Jamaica Plain.

~I

The large collection of Wilson photographs, photos of botanical gardens,
cultivated plants, etc., are to be considered library material and retained
in Jamaica Plain. The Rehder card catalogue considered essential in the
use of the cultivated plants in the herbarium also will remain in Jamaica
Plain.

_ All books, manuscripts and comparable reference material are to be cata-
logued on cards available in Jamaica Plain, and suitable designation of the
location of these books is to be made in the Union Catalogue at Widener
Library.

oo

ad

Additional books or similar reference material that may be found necessary
in Jamaica Plain will be returned for permanent deposit in the library of the
Administration Building.

—
oO

. The library of the Arnold Arboretum in Jamaica Plain and in Cambridge
is to be considered in the future, as it is now, a reference and non-circulating
library.

These proposals for a library dealing with cultivated plants consider
approximately 7000 volumes of the total Arboretum library, including
1800 periodicals and 100 folio volumes.

Further consideration was taken by the staff as to the proper housing
and location of books and specimens in the Administration Building in
Jamaica Plain. Tentative plans were drawn for a redesigning of the in-
terior of the Administration Building for horticultural activities. It was

370 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

proposed to house the library, offices, and herbarium on the first floor of
the building. With the additional space available, a lecture-demonstration
room will be possible, also on the first floor. Adequate storage and ex-
pansion for both the library and herbarium and sufficient working space
for the staff is again possible in the building.

After all aspects of the proposals and plans were reviewed by the staff
of the Arboretum, Dr. Howard submitted a report to the Corporation
stating his recommendations for the implementation of the Corporation
vote of January 19, 1953. The Corporation at their meeting on May 17,
1954, invited Dr. Howard to appear and to describe the procedures fol-
lowed in preparing his recommendations. Later in the same meeting, the
Corporation voted that the several recommendations presented by the
Arnold Professor and Director of the Arnold Arboretum were in accord-
ance with the statement of policy in the resolution of the Corporation
of January 19th. The Corporation accepted Dr. Howard’s recommenda-
tions and authorized him to proceed in accordance with those recom-
mendations to carry out plans for the removal from the Administration
Building in Jamaica Plain the herbarium specimens and books selected
for location in the University Herbarium in Cambridge.

With this authorization the move began on June 7th and was in progress
at the end of the fiscal year. The herbarium staff proceeded to draw out
the herbarium specimens to be retained in the cultivated herbarium and
to arrange these in the steel cases on the first floor of the Administration
Building. The librarian, with additional assistance, selected the books
to be retained in Jamaica Plain and these were arranged temporarily in
the library reading room on the second floor until their permanent quarters
on the first floor have been completed. The shifting of both the her-
barium and the library was coordinated in such a way that there was no
interruption of horticultural work or of service determinations. By July
Ist the herbarium of cultivated plants in the Administration Building was
completely organized and in use. Casey and Hayes Company, movers,

were charged with the task of moving the herbarium cases with the speci-
mens intact to the new building. The library to be moved was so organ-
ized that the books were taken from shelves in Jamaica Plain and placed
on previously designated shelves in the new library in such a fashion that
the books were always in order. Only the highest praise can be suggested
for care afforded both the books and the specimens in the course of the
move. The Art Metal Company, which originally supplied the steel her-
barium cases of the Arnold Arboretum, was responsible for disassembling
the cases in Jamaica Plain and reassembling them in Cambridge. That
work is in progress. All cases are to be checked and new felts applied
where necessary to assure that the old cases in their new location will be
bug-proof. A large number of herbarium specimens had to be transported
from Jamaica Plain to Cambridge in the cardboard herbarium boxes which
had been their location for a.decade. Eventually these specimens will be
arranged in proper sequence in the regular herbarium and all will be stored
in steel cases. The appropriation by the Corporation from University

1954] THE DIRECTOR’S REPORT 371

funds allowed the purchase of 430 additional steel herbarium cases. The
majority of these will be used to house the Arboretum’s collections. Within
the next year the Arboretum specimens for the first time in many years will
all be housed properly in steel cases.

The Director can only speak with praise of the yeoman’s service
rendered by the Curator, the Librarian and many members of the staff
who worked so hard to implement this move. A major goal in the care
of the Arboretum’s collections is near attainment, and while many months
of hard work remain, the best interests of the Arnold Arboretum have
been fully considered in the activities of the past year involving the move.

Horticulture:

Living collections of plants such as those that comprise the Arboretum’s
collection require constant care. The excellent condition of the grounds
throughout the year and the outstanding floral display of the spring of
1954 are indicative of the activities of the horticultural staff.

During the past year approximately 150 species and varieties of new
plants were added to the collections in Jamaica Plain. Many others were
received either as seeds or living plants. These have been grown or propa-
gated and are now being tested in our nurseries. A total of 152 shipments
including 790 species and varieties was received. Most of these came from
arboreta, gardens and nurseries in the United States and Japan, but ship-
ments originated also in eight other countries. Over 50 species and varieties
of Post Entry Quarantine plants were released by the U.S. Department
of Agriculture, Bureau of Plant Quarantine, and these are being propa-
gated for distribution as they show ornamental value. The propagation
department prepared 7396 plants for our collections or for exchange or
other distribution. These plants were propagated: cuttings, 4164; grafts,
2717; seeds, 485, and layering, 30. During the past year 144 shipments
of plant materials were sent to cooperating institutions in eleven countries.
Five small tree species, either new or extremely rare in America, were
propagated and distributed to 24 cooperating nurseries which requested
materials from a previously circulated list. Nearly 1000 individual plants
were distributed to “Friends of the Arboretum” throughout the year. Sev-
eral large yews and other shrubs were given to the Department of Build-
ings and Grounds of Harvard University for planting about the University
Herbarium Building and other college buildings.

Pruning, fertilizing, spraying and planting operations continue through-
out most of the year at the Arboretum. In the past year the base of
Hemlock Hill was cleaned of weed growth, opening an area for a new
hemlock planting at some future date. Many weed plants were pulled
from the azalea bank on South Street and in the Kalmia collection on
Hemlock Hill. The Taxus collection and the large pines along Walter
Street were given a much-needed thinning, as were the azaleas on Bussey
Hill.

Thirty-one cords of manure were spread on collections needing fertilizer.

372 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

These collections included the plantings of Carpinus, Juniperus, Chamae-
cyparis, Viburnum, Rhododendron and some of the Rhododendron ob-
tusum kaempferi in the woods near the ponds. The Rhododendron collec-
tion had a splendid display of bloom this year as a result of repeated
fertilizer applications of past years. Castor pumice has been used ex-
tensively on these penne. Summer watering has also improved the
condition of these plan

The bank cone a aaa tracks was sown to grass in the fall and
several groups of conspicuous blooming trees were planted there this
spring. Within a few years we hope to develop this bank into a colorful
display, primarily for the benefit of the commuters on the Dedham branch
of the New York, New Haven and Hartford Railroad.

A new “Bean” 400-gallon hydraulic sprayer was purchased to be drawn
by tractor. It has become increasingly evident that control of all insects
and diseases in the Arboretum merely by mist spraying is impossible.
Both mist and hydraulic spraying are used now with better coverage and
control that was formerly possible. Spray equipment was used on 24
different days last year. Particular attention to sprays and techniques of
spraying for use in arboreta and parks is being given under the direction
of the Superintendent and his staff. Fireblight, which in the past few
years has become a serious threat to the large crab apple collection, was
apparently held in check this year by the experimental use of “Agrimy-
cin,” one of the new antibiotics. A supply of this material for testing was
donated by the Chas. Pfizer Company of New York. Weed control by
the use of chemicals on a rigid spraying schedule has made poison ivy
much less evident this year than before. The plant still grows in the
Arboretum, however.

The locations of all cultivated plants on the 265 acres of the Arboretum
are recorded on 74 standard maps scaled at 20 feet to the inch and 34
additional enlargements covering the congested areas. Of these 108 maps,
approximately one-third are carefully field-checked each year. The con-
dition of the plant is noted so that it can be propagated, fertilized, sprayed,
labeled or given any treatment deemed necessary. Each year it is neces-
sary to replace from 1500 to 2000 display labels which have disappeared
or are illegible. At the same time identifications are checked and kept up
to date with current taxonomic revisions. Last spring all the lilac collec-
tion was checked during the flowering period. A few plants were dis-
carded as not being true to color and the remainder are now considered
correct according to the most recent survey of ‘‘Lilacs for America,’’ pub-
lished by the Lilac Survey Committee of the American Association of
Botanical Gardens and Arboretums. To further enhance the value of this
collection to the public, selected plants were labeled “‘best of the variety”
during the flowering season.

The collection of photographs owned by the Arboretum is used con-
stantly for reference by the staff. Prints of these photographs are used
by the staff to illustrate articles for publication. Photographs from our

1954] THE DIRECTOR’S REPORT Cae:

files were also used in several national garden magazines and in news-
papers. In recent years a collection of 35 mm. Kodachrome slides and
5 & 7 Ektachromes has been built up. These are used to illustrate lec-
tures and publications. This collection is increased yearly and its use
grows. Many of the photographs in these collections are made in private
gardens in the vicinity of Greater Boston. The splendid cooperation of
home owners who have unusual or interesting plant materials being used
to advantage and who have given Arboretum staff members permission
to photograph them is greatly appreciated. Within the past year post-
cards have been made from some of the kodachromes taken in the Arbo-
retum. These are on sale at the Administration Building and in Boston
and have proven popular among visitors.

The staff of the Arboretum remains active in the field of plant propaga-
tion. The use and effect of various plastic films in vegetative propagation
is receiving considerable attention. Species of Magnolia, Ilex and Acer
which are normally difficult to root are being tried under polyethylene
plastic. The effect of polyethylene plastic as a tie material over graft
unions is being tested on species of Picea, Pinus and Abies. A polyethylene
grafting case has been used in propagation of species of Ulmus, Quercus
and Betula. The same plastic film is being tested as a means of prevent-
ing the drying of seeds during stratification.

Acer griseum, one of the unusual maples in the Arboretum collection,
is being tested for the effect of different root stocks on compatibility.
Various concentrations of hormones on the rooting of Rhododendron
species and varieties is being undertaken. Artificial pretreatment of seeds
requiring a high temperature period followed by a low temperature is
being applied to species of Viburnum, Cotoneaster and Acer. Reports will
be published on the results of these experimental projects as the work
progresses.

The horticultural staff has also demonstrated the popularity of the
Arboretum collections through a program of adult education. Informal
classes were held in the greenhouses, and the Fall and Spring Field Classes
conducted by Dr. Wyman have attracted a record attendance in the past
year. Conducted tours of the grounds are available to large groups re-
questing such service in advance. Many garden clubs took advantage of
this service during the past year. The Massachusetts Horticultural So-
ciety had its regular field day at the Arboretum on May 15th, and four
busloads of visitors attending the National Convention of the American
Society of Landscape Architects were conducted through the grounds by
members of the staff.

The ground cover and small tree display plots at the Case Estates in
Weston are proving to be of increasing interest to visitors. Classes from
four different colleges in New England made trips to Weston to study
these collections. An experiment in the use of maleic hydracide and Crag
herbicide is under way at the Case Estates. Preliminary results indicate
that these materials may be valuable in reducing the hand labor required
in the Arboretum as well.

374 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

Cytogenetics:

A number of new hybrid ornamental apples produced at the Arnold
Arboretum have proved to be outstanding and are being propagated for
distribution. The hybrids with Malus Sargenti are of special interest
because of their dwarf habit and brilliant foliage during the spring and
early summer. The hybrids, like the M. Sargenti parent, are facultatively
apomictic and tend to breed true from seed.

Species hybridization is common in the subfamily Prunoideae (Rosa-
ceae), and a number of spontaneous hybrids between Prunus Besseyi and
other species have been found. An excellent example of the stability of
the genus is shown in the artificial cross between P. Besseyi of central
North America and P. tomentosa from eastern Asia. Although these
species differ in many characters and have been separated for hundreds
of thousands of years, they cross readily and the F1 hybrid is fertile.

The “Merrill” Magnolia, a hybrid produced fifteen years ago, has
proved to be vigorous and early-flowering. It became available to the
horticultural public from commercial sources in the spring of 1954.

Dwarfing rootstocks and bark inversion are being used for the develop-
ment of dwarf ornamental trees. The bark inversion technique is being
used to induce early flowering in ornamental trees and vines which are
reluctant to produce flowers.

Hally J. Sax (Mrs. Karl Sax), utilizing the extensive living and her-
barium collections of the Arboretum, completed a cytotaxonomic study
of the genus Cotoneaster. Most of the species were found to be apomictic
triploids, There is some evidence of facultative apomixis in these triploid
species. Thus the species is able to perpetuate hybrid complexes through
apomixis, yet retain some genetic plasticity to meet new environmental
conditions by occasional hybridization.

Dexter Sampson, a graduate student, continued his work on the cyto-
logical analysis of the progeny from triploid hybrids in Philadelphus and
Forsythia. Bradford Johnson, another graduate student, began a study
of the cytology of facultative apomixis in Malus.

Comparative Morphology:

Professor Bailey has completed his task of preparing material for pub-
lication in book form by the Chronica Botanica Company. This volume
which deals with salient trends of anatomical research during the period
1909-1953 will be released for sale in the near future. Miss Margery P. F.
Marsden has completed her intensive investigation of Clerodendron tri-
chotomum Thunb. and received the doctorate from Radcliffe College in
June. Mr. Chi Ling Chen developed symptoms of tuberculosis early in
the fall and is having to spend at least a year at the State Sanatorium in
Rutland. This was extremely unfortunate, not only in postponing com-
pletion of his work on the Sapotaceae for the doctorate, but also in de-
priving Professor Bailey of his expert assistance in preparing the wood
collection for transfer to its quarters in the new botanical building.

1954] THE DIRECTOR’S REPORT 375

The wood collection was initiated by Professor Sargent during the early
years of his administration of the Arnold Arboretum. It has subsequently
been enlarged and strengthened by additional accessions at the Arnold
Arboretum and by successive contributions made by the Forestry School,
the Bussey Institute and the Department of Biology. Since 1936, it has
been housed, as have staff members of the Arnold Arboretum concerned
in its use, in the Biological Laboratories without cost to the Arboretum.
For a number of years the Department of Biology provided financial
assistance in the making of catalogues and microscopic slides. Further-
more, Professor Wetmore and his associates contributed much time and
effort to expanding and improving the collection. In transferring the
much enlarged collection, new steel cases and many new trays have been
provided by the University. The Biological Laboratories also generously
contributed a large number of the trays in which the collection was for-
merly stored. In its new quarters, the collection is housed in close prox-
imity to the collections of fruits and seeds and slides of pollen, leaves and
flowers. Thus, any taxonomist working with herbarium specimens now
has efficient and convenient access to supplementary data contained in
these extensive and very valuable collections.

The Herbarium:

The activities of the herbarium staff during the past year were involved
in the preparations to implement the vote of the Harvard Corporation.
The Curator, Dr. Kobuski, and Miss Perry devoted regular time and many
extra hours to the designation and separation of the horticultural her-
barium and the preparation of the non-horticultural material for the move.
In spite of these steps of preparation, the regular activities continued in
the herbarium. During the past year 3560 specimens were mounted and
added to the herbarium, which now contains 675,119 sheets. A total of
12,421 specimens were sent out in exchange, 11,121 to foreign institu-
tions and the remainder to American herbaria. New collections received
at the Arboretum number 27,758 specimens. Of these, 12,000 specimens
came as gifts or in exchange, while approximately 16,000 specimens were
received from expeditions or collectors sponsored completely or in part by
the Arnold Arboretum. Nearly 85% of the incoming materials were re-
ceived from Malaysia and Asia, forming a significant addition to the
valuable collections already in the Arboretum from this area. Among
the noteworthy accessions are 14,000 specimens collected by L. J. Brass
on his most recent trip to New Guinea; Richard Schultes’ collection of
3,500 specimens from the Amazon; 2,000 specimens from the Philippine
Bureau of Science collected in oriental Mindoro, and 3,500 specimens
from Japan, the Ryukyu Islands and other accessible areas of Asia. An
active exchange continues between the Bogor Herbarium in Java and the
Arnold Arboretum. During the past year the Arboretum received 2,400
specimens from the Bogor Herbarium collected in various regions of
Indonesia.

The number of requests for loans of specimens from the Arboretum

376 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXXV

collections continues high. A total of 5,457 specimens were sent out on loan
during the year to twenty-two different institutions. Foreign requests for
loans numbered eleven.

During the year, staff members conducted research on plant families
and areas of their specialties. Dr. Howard was appointed Senior Botanist
on a project sponsored by the Institute of Jamaica in Kingston, Jamaica,
British West Indies, to survey the native vegetation on bauxite soils on
that island. The project hopes in the course of this work, to determine
what plants of potential economic value can be grown on mined-out
bauxite areas. Dr. Howard continued his work on the vegetation of the
Caribbean Islands and made some progress on monographic studies of the
genus Coccoloba.

Dr. Kobuski continued his work on the Asiatic members of the Theaceae
and specifically towards a treatment of this family for the Flora Malesiana.
He also identified several large collections of this family for various
herbaria.

Dr. Johnston concentrated on his studies of the Boraginaceae, with a
special interest in the herbaceous genera found in Asia and the Middle

Dr, Perry continued her work on the New Guinea flora, giving special
attention to the recently received collections of L. J. Brass

Dr. Hu is completing a monographic study of the ornamental genus
Philadelphus and supervises the initial stages of the preparation of a list
of the species of flowering plants reported from China.

Dr, Merrill is now receiving the proofs on his book regarding the botany
of Captain Cook’s two voyages in the Pacific. It is expected that this
major contribution will be published in the fall.

The Library:

The preparation and implementing of plans to organize a horticultural
library and the separation of those books to be moved to Cambridge
formed a major portion of the work of the librarian and the library staff
during the past year. The Corporation requested in its vote that all
Arboretum books to be housed in Cambridge be distinctly marked. Thus,
within the past year all books which are part of the Arboretum library
have been checked and marked where need be, with a stamp or sticker
on the back, an embossed imprint within the pages, or a bookplate, or a
combination of these. An excellent selection of books of horticultural
use was made with the cooperation of Dr. Wyman and other members of
the staff. These books to remain in Jamaica Plain will eventually be
located in a newly designed library and reading room on the first floor
of the Administration Building, with room for expansion or for little used
volumes on the second floor of the herbarium wing.

Additions to the library by gift, purchase or binding totaled 271 vol-
umes, The library now contains 48,673 bound volumes on the shelves.
Pamphlets numbering 206 were added to the pamphlet collection, bring-
ing that collection to a total of 15,410 items. Five hundred and four

1954] THE DIRECTOR’S REPORT 377

catalogue cards were added to the main catalogue and 1206 cards were
added to the Gray Herbarium index of American species. Inter-library
loans showed an increase over past years, although many additional re-
quests were answered by typing short descriptions, or through the use
of microfilm and photostat copies.

Exhibits and Displays:

The Larz Anderson Collection of Japanese Dwarf Trees was featured
at the exhibit of the Arnold Arboretum at the Spring Flower Show of the
Massachusetts Horticultural Society. The plants were exhibited in a
setting of a Japanese garden, An interesting bamboo shelter was erected
to shelter these plants in a fashion comparable to that used in Japan.
Three excellent Kurume azaleas loaned by Mr. John Ames added color
to the display. A first prize and a gold medal were won by this display.

Several additional displays of living plant materials were furnished for
other shows of the Massachusetts Horticultural Society during the year.
An Arboretum display of sprays of flowers, tree photographs and Ekta-
chrome display panels occupied a featured location at the National Con-
vention of the American Society of Landscape Architects at the Hotel
Somerset. This exhibit attracted much attention and was mentioned in
the programs and news releases of the meetings.

Publications:

Four issues of Volumes 34 and 35 of the Journal of the Arnold Arbo-
retum were issued quarterly totaling 522 pages. Dr. Kobuski serves as
editor of the Journal, assisted by an editorial board of other members of
the Arboretum staff. Twelve numbers of Arnoldia were issued under the
editorship of Dr. Wyman and were distributed to the Friends of the
Arnold Arboretum and additional subscribers so that the total distribu-
tion now numbers 2,200. The subscription price of Arnoldia was raised
to two dollars per year. This represented the first increase in price since
this publication first appeared. A demand for back numbers of Arnoldia
increased to the point where it seemed desirable to accept an offer of the
D. Van Nostrand Company of New York to publish some of these as
“The Arnold Arboretum Garden Book.” Some forty-five issues were
selected and Dr. Wyman wrote an introduction and foreword for this
book. Proof of the book is now being received and publication is
expected in the fall of 1954.

Gifts and Grants:

The Arboretum is fortunate to receive a large number of gifts from
individuals interested in the work of the organization and the staff. Most
of these are received as ‘‘memberships” in the “Friends of the Arnold
Arboretum.” During the past year 431 memberships were received. Such
gifts are assigned, unless otherwise designated, to a fund called “gifts for
cultural purposes” and are used exclusively to sponsor additional work

378 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV

on the grounds, such as developing new collections, additional care for the
plantings or for research and work leading to the development of new
hybrids, dwarf plants and similar projects. Additional gifts were received
designated for specific purposes. One gift from a Friend of the Arboretum
was to be used to support foreign collectors and thereby obtain for the
Arboretum additional material of horticultural value. It is hoped that this
meritorious fund can be increased in future years. Another large gift was
placed at the discretion and joint use of Dr. Reed Rollins, Director of
the Gray Herbarium and Dr. Howard, Director of the Arboretum, to be
used for research in systematic botany for the benefit of both the Arnold
Arboretum and the Gray Herbarium.

The Arboretum received a bequest from the estate of Emery Holden
Greenough which was assigned to the endowment of the Arboretum.
Through the generosity of Mrs. Martha Peters, her home and the land
thereabout on South Street was given to the Arboretum. It is hoped
that eventually when funds are available the setting of this old house
can be used as a demonstration landscape area to show plants for small
homes in a good setting. The adjacent land when reconditioned will be
of great value to the Arboretum as a nursery area.

Numerous Friends have given books on plants and flowers to the Arbo-
retum. The value of these books is determined by the librarian and
reported to the donor for income tax purposes. Many of the books are
welcome additions to the library, while others serve as replacements for
worn books long in use.

Bibliography of the Published Writings of the Staff and Students
July 1, 1953 — June 30, 1954

BarLey, I. W. The anatomical approach to the study of genera. Chron. Bot.
14: 121-125. 1953.

COGGESHALL, ROGER G. oo some rarer plants from seed. Trees Mag.

, 21. Mar.—Apr. 1

Howarp, RICHARD A. A ha of the Botanic Garden of St. Vincent, British
West Indies. Geographical Review 44: 381-393. 1954.

Botanical Gardens in West Indies History. Garden Journal 3: 255-257.

53.

anual de la Flora de los Alrededores de Buenos Aires (Review).
Bull. pay Bot. Club 81: 255-257. 1954.
Progress at the Arnold Arboretum. Newsletter Amer. Assoc. Bot. Gar-
dens and Arboretums 18: 13-15. 1954.
Hu, Sutu-yinc. Notes on the flora of China, III. Jour. Arnold Arb. 35:
189-200, pl. 1, 2. 1954.
Jounston, Ivan M. Boraginaceae. In: Botanical Exploration in Venezuela-III.
Fieldiana Botany 28: 511, 512. 1953.
Polemoniales (Pars) Family 2 Boraginaceae. In: Flora of Trinidad
and Tobago 2: 191-209. 1953.
Studies in the Boraginaceae, XXV. A revaluation of some genera of
the Lithospermeae. Jour. Arnold Arb. 34: 258-299. 1953.

1954] THE DIRECTOR’S REPORT 379

JounstTon, Ivan M. Studies in Boraginaceae, XXVI. Further revaluations of the

genera of the Lithospermeae. 35: 1-81.
Studies in the Boraginaceae, XXVII. Some general observations con-

cerning the Lithospermeae. 35: 158-166. 1954.

Kosusk1, C. E. Camellia Soe known in cultivation. Am. Camellia Year-
book 1953: 1-9, figs. 3.

Lipp, Lewis F. New Boe in ) plant propagation. Garden Journal 4: 70, 71.
1954.

Merrill, E. D. Bibliographic notes on G. Forster’s “De ee esculentis
insularum oceani australis (1786)”. Pacific bis 8: 35-4

Biographical memoir of Merritt Lyndon Fernald ane ocd Nat.

Acad. Sci. Biog. Mem. 28 (3rd Memoir): 45-64. portrait, 1954.

Miscellaneous Malaysian notes. Jour. Arnold Arb. 35: 134-156, pl. 1.

1954.

Perry, Liry M. Plantae Papuanae same eca ria The Papuan species
of Macaranga. Jour. Arnold Arb. 34: 191-257.

Sax, Kart. The Arnold Arboretum during the se Be ended June 30, 1953.
Jour. Arnold Arb. 34: 412-416. 1953.

Enough for all? Jour. Heredity 44: 203, 204. 1953.

Interstock effects in dwarfing fruit trees. Proc. Am. Soc. Hort. Sci.
62: 201-209.

SCHWARTEN, LAZELLA. Index to American Botanical Literature. Bull. Torrey
Bot. Club 80: 355-364, 432-443, 518-529. 1953; 159-167, 259-269.

& KATHERINE [FERNALD] Lounes. Bibliography : M. L. Fernald.
Nat. Acad. Sci. Biog. Mem. 28 (3rd Memoir): 66-98. 1954.

VERDOORN, FRANS. Botanical gardens and arboretums of ne past and their
reconstruction (Colloque Int. Jardins Botan.). Année Biol. 29: 274-282.
1953

L’arboretum moderne (Colloque Int. Jardins Botan.). Année Biol. 29:
421-427.
Current organization and activities of the botanical section, subsections,
commissions & committees of the International Union of Biological Sci-
ences. I.U.B.S. Bot. Section Leaflet 4: 1-8. 1953.

Chronica Botanica library and archives. Chron. Bot. Keepsake 6: 1.
953

The discovery of unicellular life. Chron. Bot. Keepsake 7: 1-16. 1953.
Editor’s foreword to Plant Genera, their nature and definition. Chron.
Bot. 14: 93-101. 1953.

The international plant science congresses, their history and aims. Pro-
ceedings 7th Int. Bot. Congress, pp. 42-56. pl. v-viii. 1953 (ie., 1954).
Wyman, Donatp. Botanical Gardens and Arboretums in America. Suppl.

Dictionary of Gardening, Royal Horticultural Society. 1954.

Care of ornamental vines. Plants & Gardens 10: 23-27. 1954.
The Catawba Hybrid— Rhododendrons. Horticulture 32: 218-231.

illust. 1954.

— Famous British Gardens. Trans. Worcester County Hort. Soc. 1952:
70-75. 1953.

Be of the better ground covers. Arnoldia 14: 21-24. 1954.

Flowering displays in the Arnold Arboretum. Arnoldia 14: 9, 10. 1954.

Japanese dwarfed trees. Arnoldia 14: 1-7, illust. 1954.

380 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXV
Wyman, Donacp. A new Magnolia. Horticulture 32: 144, 145, illust. 1954.
Plant for permanence — Rhododendron vaseyi. Flower Grower 41: 10,
11, illust. 1954
Plant for permanence -— The Hemlocks. Flower Grower 40: 15, 1954.
Planting the modern home. Am. Nurseryman 98*: 25, 144-146. 1953.
The propagation and registration of woody plants in the United States:
Report of the Thirteenth Horticultural Congress (1952) 1: 529-536. 1953.
Seeds of Woody Plants. Arnoldia 13: 41-60, illust. 1953.
Rhododendron vaseyi. Flower Grower 41: 10, 11, illust. 1954.

Some shrubs and trees with colored foliage growing in the Arnold
Arboretum. Arnoldia 14: 13-19, illust. 1954.
Vines as ground covers. Plants & iat 10: 58-62. 1954.
Vines for every purpose. Plants & Gardens 10: 4-14. 1954.
Vines with both sexes separate. Plants & Gardens 10: 68, 69. 1954.
Vines for winter beauty. Plants & Gardens 10: 46-50. 1954.
(With Harold O. Perkins) Flowering in Wisteria. Plants & Gardens

10: 70, 71. 1954

RicHarp A, Howarp,
Director.

1954] THE DIRECTOR’S REPORT 381

Staff of the Arnold Arboretum
1953-1954

RicHarp ALDEN Howarp, Ph.D., Arnold Professor of Botany, Professor
of Dendrology, and Director.

JosepH Horace Fautt, Ph.D., Professor of Forest Pathology, Emeritus.

Ermer Drew Merritt, $.D., LL.D., Arnold Professor of Botany,
Emeritus.

IRVING WipMER BAILey, S.D., Professor of Plant Anatomy.

Rocer Gipps CoGcESHALL, Assistant Propagator.

BEATRIX FARRAND, L.H.D., Consulting Landscape Gardener.

ALFRED JAMES ForpHAM, Assistant Superintendent.

HeEMAN ARTHUR Howarp, Assistant Horticulturist.

Suiu-y1nc Hu, Ph.D., Herbarium Assistant.

IvaAN Murray JouNsTON, Ph.D., Associate Professor of Botany.

CLARENCE EMMEREN KopuskI, Ph.D., Curator.

LEWIS FREDERICK Lipp, Propagator.

SusAN Detano McKeE vey, A.B., Research Associate.

Lity May Perry, Ph.D., Botanist.

Kari Sax, S.D., Professor of Botany.

LAZELLA SCHWARTEN, Librarian.

ETHEL ELIZABETH UpuHam, A.B., Herbarium Assistant.

FrANS VERDOORN, Ph.D., Research Associate.

STELLA MABEL WHITEHOUSE, Business Secretary.

Rosert GERow WILLIAMs, B.S., Superintendent.

DonaLp Wyman, Ph.D., Horticulturist.

382

JOURNAL OF THE ARNOLD ARBORETUM

[VOL. XXXV

INDEX

Abildgaardia eragrostis, 215
oT des, 221

— fus

pri 1g

Additional fore on Nothofagus, 266
Aegonychon, 38

Agathis aust tralis, 270

Aglaia heterobotrys, 138

Agyneia latifolia, 139

—

Albizia ‘plendens, 137

Alkanna
Alsodela, Pre 145

— semigyrata,

Anonymos caroliniensis, 59
Antidesma coriaceum

, 149
Apomixis in Cotoneaster, Polyploidy and,

334
Aporosa ae rere 139
— latifolia,
sper oh York (Queensland) Ex-
i » ACE ryptoga ms of the 1948, 260

b
—- rata 56

Arnebia _e 56
uttata,
— hispidissima, 51, 55
56

— migiurtina, 5
Aspen in Uiah Mountains,
Leafing of,

Prevernal

Balangue gaertneri, 140
Barroetea, 108, 124

i soce i ra 106, 116
— setosa
Sharing latisquama, 97, 106, 116
59

ac-Zelandiae, a

Bhesa moja, 141

Brissy, P. Cryptogams of the 1948 Arch-
bold Cape York (Queensland) Expe-

dition, 260
Brake, S. T. The i gees ae
in New eee by L. J. Brass, IV, 203
the XXVI.

Boraginaceae, Studies in
Further aici of the Genera of
the Lithosperm

Boraginaceae, Studies in the, XXVII.

Some General Observations Concern-
g the oon 158

Brickellia, 24

eer 3; 166

— sect. Estuslosies 41

— sec rospermum, 44

— arvense
— calabrum, 45
— Gastoni, 45

1954]

sl cane incrassatum, 43
— purpureo-caeruleum

—ramosissima, 38

—tenuiflorum, 42

— Zollingeri, 45

Calamintha polycephala, 86
Calophyllum rotundifolium, 144
Camptochaete brisbanica, 260
Canthium tavoyanum, 155
Carphephorus, 90, 100, 124

— bellidifolius, 94, 106, 116

— corymbosus, 95, LG
— pseudo-liatris, 94, 106, 116
—tomentosus, 95, 16

Carphochaete, 90, 99, 124

— Bigelovii, 94, 106, 116

— Grahami,

CassIz, VIVIENNE DELLOW. New Zealand
Cane ers, 268

Ceanothus eae 141

Celastrus alnifoliu

Cerinthe, 3, 65, 165

major, 65
Circa: castanocarpus, 140

China, Notes on the Flora of, III, 1
Chinese Flora, menclatural
Changes in the

— tchefouensis, 193
— pallasii, 190

INDEX

383

Clematis recta angustifolia, 191
Clerodendron amplius, 1

— fortunei, 153
sundocale 153
—simile, 153
Coenogonium implex 263

Comlene: are ener 263
Colsmannia,

73
Conifers, New Zealand, 268

Control Tree Growth by Phloem
Blocks, pedi
COPELAND, T F. Some Details of

ER
the Structure of anid Mahadincs cham-
aecistus

Cordia venosa, 154
Cotonck (an Polyploidy and Apomixis
in, 334
Cotoneaster acuminata, 340, 342, 352,
354, 356, 35
— acutifolia, 340, 343, 354, 356, 359
352, 354, 356,

—— villosula, 340, 343,
59

—adpressa, 341, 352
——hessei, 341, 352
—— praecox, 341, 352
— affinis, 3

— — bacillaris, 341, 345, ae 356, 359

—bullata, 341, 344, 352, 355

— — floribunda, 341, 345, 352
—-—macrophylla, 341, 344, 345
— conges

358

Thee 341, 354, 357, 358
—crispii, 350

—dammeri, 341, 354, 357
—dielsiana, 341, 346, 352, 357
—— elegans, 346, 35
—divaricata, 341, 347, 352, 357
— foveolata, 341, 344, 355
— francheti, 341, 348, 357, 359
——newryensis, 34
— frigida, 341, 346, 350, 352, 354, 356
— — fructi-luteo, 350
— froebelli, 341
— glabrata, 341, 354, 357
—slaucophylla, 341, 357

— integerrima, 341,
— lindleyi, 341,

349, 352, 358
345, 346, 352, 356

384 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXXV

Cotoneaster lucida, 341, 344, 355

—melanocarpa, 341, 355, 359
ommixta, 341, 356, 359

— — laxiflora

—microphylla, 341, 354, 357

— moupinensis, 341, 344, 352, 355

— multiflora, 341, 347, 352, 359

—w—-calocarpa, 342, 348

—— granatensis, 342, 348, 358, 359

—nitens, 342, 352, 357

— obscura, 342, 348, 355

—— cornifolia, 342, 348, 349, 352

— racemiflora, 358

— — desfontaini, 342, 347, a

—— soongorica, 342, 347, 356,

—— veitchii, 342, 347, 352, ah en

— rotundifolia,
—rubens, —

— rugosa

a. 342, 357
—- ccosa, 357

wn
°
3
a
me
=
w
sé
BSS
We
>
so
Ww
on
N
Ww
un
~I

s a
—tenuipes, 342, 343, 355
— sa

o
°
B
= |
o
°
we
>
N
Ww
>»
eo)
wW
mn
ao

029
— wardii, yn 349, 352, 357
— watereri, 50

—zabeli, 342, 349, 357, 358

yore hcemeee P. Prevernal ia
of n Utah abies

Craibiodendron Kwangtungense, rh

99

136
Cryptogams of the 1948 Archbold Cape
York (Queensland) Expedition, 260
Cyperaceae Collected in New Guinea by

— socotranus,

Dacrydium bidwillii, 271
—— biforme, 269
——*x lxifotium, 269
— biforme,

——x lxifolium, 269

Diaphora cochinchinensis, 228

Dicera craspedum, 14
ictyonema irpicinum, 265

Dioclea,

Diosma serra 141

Diospyro os cathayensis 86

Director’s Report on the Arnold Arbo-
retum during the Fiscal Year Ended
6

Drepanolejeunea Micholitzii genuina, 262
— obliqua,
— tenuis, 263

Ecdysanthera schrieckii, 152
Echinolytru um dipsaceum, 214
65

Ecotypic Wunik of the Photoperiodic
Response in Populus, 167

Elaeocarpus oon 142
—tectorius, 14

Embelia i: 149

— lucida, ,

apuan
Eriopus eal 260
—sp., 260
Euonymus orgyalis, 197
Euphorbia longana, 140

Ficus cordatifolia, 134
] ensis 13

— Malunu
ea

rifolia, 134
Fimbristsis sect. ge ai 220
t. Eufimbristylis, 2

1954]

Fimbristylis sect. Laoag 208
—sect. Trichelostylis,

——glabra, 212

1
—cinnamometorum, 220, 222
—complanata, 21
—cylindrocarpa, 208
—cymosa,

— — capitato-umbellata, 219
2

214
acorns. 220, 223

INDEX 385

Fimbristylis stricticulmis, 211
— tetragona, 208, 21

— xyridis rigidula, 211

— xyroides,

Frullania Johnsonii, 262

— SPp.,

GalIser, L. O. ge in the Kuhniinae
(Eupatorieae) it;

Garberia,

ee an a 106, 116

Glycosmis macrantha, 137

— oliveri, 137

Halacsya, 165
WARD, RicHArD A. The Director’s Re-
port on the Arnold Arboretum during
the Fiscal Year Ended June 30, 1954,
7

Hv, Suru-yinc. A Monograph of the
5

Hypolytrum ‘nierocaouae: 235
Hypoporum capillare, 224

x
— roxburghii,

Iriha debilis, 215
iliacea, 216

— quinquangularis, 217

Isolepis cochleata, 208

i ¢)

—tetragona, 217

Jounston, Ivan M. Studies in the Bo-
raginaceae, XXVI. Further ye
tions of the Genera of the Litho-

meae, 1
OHNSTON, IvaAN M. Studies in the Bo
raginaceae, X XVII. Some General Ob-
servations on the Lithospermeae, 158

al

Kuhnia, 104, 124
—adenolepsis, 116
—eupatorioides corymbulosa, 96, 105,
106,
—— ozarkana, 116
ie hare, 96, 104, 106,116
96, 105, 106, 116
ee maak 116
— Mosieri, 96, 106, 1

386

wabeecioen Rater ve rig 116
ee Pr 106, 116

— Schafineri, , 106, 116
Kurrimia “lhe, 141
— robust

Lagerstroemia flos-reginae, 146
— speciosa, 1

Lannea coromandelica, 140
Laportea elliptica, 135

3
i
i)
ene =
ae
o
tS)
oO
=

Leptogium phyllocarpum isidiosum, 264
Liatris, 124

Libocedrus bidwillii, 270
oom heet OC

—doniana, 2

— plumo )

— ‘pobuntan, 150
— subsessile, 199

Linociera cuspidata, 151
ened 150

— ridleyi,

—sp., 1 -

—stenura, 151

Lithodora, 38, 44, 165

Lithospermeae, Studies in the Boragi-

aceae, XXV Further Revaluations
the Genera of the,
Lithospermeae, Studies the Boragi-

Some General Ob-

ngiflorum, 14, 59
— longifolium,
—mandanense, 59
— officinale, 59
— purpureo-caeruleum, 38, 44
34

— Tournefortii, 58

JOURNAL OF THE ARNOLD ARBORETUM

[ VOL. XXxv

oe -riidtia 45
Lobostemon
Lycopsis a, 58

Macromeria, 1, 166
Macromerioides: 5

ea

Maharanga, 4, 78, 165
— Borii, 81
— dumetorum, 81

— verruculosa, 81

Malaysian Notes, Miscellaneous, 134

Margarospermum

Mariscus fulvus confusus, 237

“patie avg diclados, 260

Meneghinia, 51

MERRILL, E D. Miscellaneous Malaysian
34

Notes
Micrechites furcata, 152
r 52

Microtropis lanceolata, 152

Mikania cordifolia

— scandens,

Miscellaneous repel Notes, 134

Mischospora efoliata,

Moltkia, 44, 165

Monograph of the Genus Philadelphus,
A, 275

Munbya, 51

Neatostema, 38

1954]

New Guinea J. Brass, IV, The
03

Notes on the Flora of ee III, 189

Nothocnestis sumatrana

ete Additional ne on, 266
266

— resinoee: 267

Odina wodier, 140

ginicum, 18
Ophiorrhiza reticulata, 155
— sarawakensis, 155
Osmidium, 1
Osmodium, 18
Otopetalum micranthum, 152

Pachygone ovata, 140

INDEX

387

Pannaria oe 264
— sublurida,
Parishia ae 140
eT
malabog,
Pabieila isa sorediata, 264
PAuLEy, Scot THomas O. PERRY
Ecotypic vara of the Photopen:
0 esponse in Populus, 167
Pentaloba lanceolata, 145
semigyrata, 14
Pen penta phus, 59
Perittostema, 2, 30, 166

Perry, Tuomas O. & Scott S. PAULE
Ecotypic Variation of the phetorent
odic Response in Populus, 167

ser sci A Mon birkph of the
Genus, 275

Phiadeps, 312

. Californicus, 303,

Bae Coulterianus, 303, aa 325
—sect. Hirsutus, 304,
—sect. Microphyllus, 303, 314

sect. Pauciflorus, 30. 1
—sect. Poecilostigma, 303, 15
—sect. Pseudoserpyllifolius, 304, 314

— sect. Serpyllifolius, 304, 4 4

— sect. Stenostigma, 303, 314

—ser. Decorticatae auctor 314
ati, 315

’ Deutzioides, 304, 314

; Buphiladelphus, 303, 314
. Gemmatus, 303, 3 ri

. Macrothyrsus, 303, 314
. Gemmati, 314

—affinis, 316

ss aspentnie as

—au stro-mexicanus 324

— coulteri, 325, 32 330, 332
—- glabripetalus, 3
— karw sereie oa

20

31
— mexicanus, a $315, 319, 321,324
— myrtoides
Ba ah
—osmanthus, 326

—trichopetalus, 319

388

Philadelphus zeyheri, 316
Phillyrea Tobusta, 150
The Control of Tree
, 251
Photoperiodic Response’ in
pic Variation of the, 1
Sa alpinus, 271
— glaucus

Populus,
7

richomanoides, 269
— trichomanoides

Physocolea preeinent 263
Pithecellobium splendens, 137

— abietine, 261
— oppo

ze car ned oppositus, 261
Plectronia rps Senay 155
Pleurozia articulata, 262
Podocarpus acutifolius, 271

— ferrugineus, 270
otara, 269

, 70, 164
Polyploidy ad Avoutes in Cotoneaster,
334

Populus, Ecotypic Variation of the
Photoperiodic Response in, 167

Populus tremuloides aurea, 2

Prevernal mie ng of Aspen in Utah
Mountains, 239

bi ee 2, 33, 165
— revolutus,

Psychotria nolytvichs: 155

— rhodo 56

—rufipila, 1

— sangea

— trichophlebia, 155
salen 234
Purshia,

Pyrus tie 134

Radula aa 262
— buccini
— reflexa,
Report on i Arnold Arboretum during

JOURNAL OF THE ARNOLD ARBORETUM

[ VOL. XXXV

the Fiscal Year Ended June 30, 1954,
The Director’s, 36
Rhesa moja, 141
Rhodothamnus chamaecistus, Some De-
tails of the Stru see of, 82
Rhynchospora mri 238
p

Rochelia, 38

Sarcosperma tonkinense, 141
Satureja polycephala, 86

Sax, HALiy bs ce and Apomixis
in Cotoneaster, 33

SAX, est Lt. The eimgeirs of Tree Growth
by P oem Blocks,

Ai aes ne

Schleichera oleo 140

—trijuga, 140

Schoenoplectus merrillii, 206

38

a nus, 2
Scirpus aestivalis, 212
— annuus, 212
— bengalensis, 216

1

— gilnuinne wane 216
— strobolinus, 204

1954]

aa subcapitatus rigidus, 206
riangularis,

eimiiee 203

— tetragonus, 209, 217

— ciliaris, 23

— cochinchinensis, 229
— dietrichiae, 227

— doederleiniana, 228
—elata 8

illosa, 232
a —ithosperma, 224, 232

—_—— ecears 224

— densi- ea 225
— neesiana
— novae- pallndie 225.232
— hes
—ersrcts 224, 232
— eens 231%) 235

— rugosa, 226, 233

INDEX 389

Scleria scrobiculata, 229, 233
is, 225

STEENIS, C. G. J. van. Additional
Note on Nothofagus, 266
Stenosolenium, 3, 46,
—saxatile, 46
STEWARD, ALBERT Two Nomencla-
tural Changes in Ce Flora,
me aurata, 264
— Sayeri, 264
—sulphurea, 264
Strobila, 51
Studies in the Boraginaceae, XXXVI.
Further Paints of the Genera

7
Studies in the Kuhniinae (Eupatorieae)
Suetaua S12

Tiliacora, 140
inospora oa 195
94

— yunnanensis
Toxostigm 1
luteum, 51

purpur

Si
Trachelospermum philippinense, 152
eee ae by Phloem Blocks, The

rol of, 251
Teichlostis filiformis, 210
—miliacea, 216

390 JOURNAL OF THE ARNOLD ARBORETUM

Trichelostylis quinquangularis, 216
Trichosanthes,

Two Nomenclatural Changes _ in
Chinese Flora, 86

the

Uncinia riparia, 234
Urtica peltata, 135

Vareca lanceolata, 145

hearer 4, 71, 164

— barbata, 71

Ventilago “ramble, 142
— lanceolata, 1

Walsura monophylla, 138

Zizania terrestris, 228

[VOL. XXXV