JOURNAL

OF THE

ARNOLD ARBORETUM

HARVARD UNIVERSITY

EDITORIAL BOARD

KOBUSKI, Editor I. M. JOHNSTON

C.E.
I. W. BAILEY KARL SAX

VOLUME XXXII

JAMAICA PLAIN, MASS.
1952

Reprinted with the permission of the
Arnold Arboretum of Harvard University

KRAUS REPRINT CORPORATION
New York

1968

LA

DATES OF ISSUE
No.1 (pp. 1- 96) issued January 15, 1952.
No. 2 (pp. 97-198) issued April 15, 1952.
No. 3 (pp. 199-298) issued July 15, 1952.
No. 4 (pp. 299-426) issued October 15, 1952.

Printed in U.S.A.

TABLE OF CONTENTS

A Taxonomic REVIEW OF THE GENUS ActTINipIA. By Hui-Lin li...

~ STUDIES IN THE BORAGINACEAE, XXII. NotTeEwortHy SPECIES, CHIEFLY
ASIAN AND SoUTH AMERICAN. By Jvan M. Johnston .................

STUDIES IN THE THEACEAE, XXV. THE GENUS ANNESLEA. By Clarence E.
Pg oon eset A Ga 8 OSE eo 29 PRS S Od Cte ebs HOE oes ee

ERIANDRA, A NEW GENUS OF POLYGALACEAE FROM NEW GUINEA. With one
plate. By P. van Royen and C. G. G. H. van Steenis .................

StupiEs oF PacriFIc ISLAND PLANTS, XI. FURTHER NOTES ON FIJIAN
PLOWSRING PLANTS: Ey A.C. S0Ul 2 ook cs ici aw beh andesetaeeercs

STUDIES OF PaActFIc ISLAND PLANTS, XII. THE CUNONIACEAE OF FIJI AND
PRM Poe Eat eros kk ah oa Ra Oe ee Ree ee eee

NOTES ON XANTHOSTEMON F,, MUELLER AND KJELLBERGIODENDRON BURRET.
By BE: DACP ooo eos ga vc es ss <e b4-2 ay BES Se eee

NOTES ON THE FLorA OF CHINA, II. With two plates. By Shiu-ying Hu...

SPONTANEOUS WHITE PINE Hyprips. With two plates. By Albert G. John-

STUDIES IN THE THEACEAE, XXVI. THE GENuS VISNEA. By Clarence E.
POSER Wena d 4 hk gil Sh apard GBA ER PO Ak EE IE BP ERLE SE OE

THE GENUS AMENTOTAXUS. By Hui-Lin Li ........0. 0.200.000.0002 004.--

WILLIAM JACK’s GENERA AND SPECIES OF MALAYSIAN PLANTS. With one
Pate, Bytes W OU. sos vei eels 60a apo eer teaeserehel eee.

THE COMPARATIVE MORPHOLOGY OF THE ICACINACEAE, VI. THE POLLEN.
With nine plates. By A. Orville Dahl .........0.0..0 0000 eee

ee SILVER Mapes. With one plate. By Scott S. Pauley and Albert
RORMSOW: 5.25 x bd ia ee ka 8 be Rae LEAR

STUDIES IN THE BORAGINACEAE, XXIII. A SuRVEY OF THE GENUS LiITHOS-
PERMUM. With three plates. By Jvan M. Johnston ..................

STUDIES OF lara IsLAND PLANTs, XIII. Notes on FIyIAN EvupnHor-
BIACEAE, Al OC SSTVIR: 3 ste se kt in, ed

THE ARNOLD ARBORETUM DURING THE FISCAL YEAR ENDED JUNE 30, 1952

BIBLIOGRAPHY OF THE PUBLISHED WRITINGS OF THE STAFF AND STUDENTS
JULY 3, 1951-JUNE 30, 1952 Ae es Seok ee ee a ewe os

STAFF OF THE ARNOLD ARBORETUM 1951-1952 .......................

296

299

BS

-
: oe :

Bn

OS

JOURNAL

ARNOLD ARBORETUM
Vou, XXXII JANUARY 1952 Nomeer 1

A TAXONOMIC REVIEW OF THE GENUS ACTINIDIA
Hur-Lin Li

Tur Genus Actinidia was revised by 8. T. Dunn in 1911.* He then
recognized 24 species for the genus, and these were arranged in four
sections. The genus is limited to eastern and southern Asia, with the
main center of development in China. Intensive botanical exploration
in eastern Asia during the last forty years has added 26 supposedly new
species to the genus, in addition to a few new combinations and other
nomenclatural changes. A very much larger series of specimens is now
available than at the time of Dunn’s study. It is the purpose of this
paper to coordinate these later additions with Dunn’s treatment and
to evaluate his system of classification in the light of our fuller know]l-
edge of the genus.

Among the 24 species enumerated by Dunn, he had not seen material
of five, namely A. giraldii Diels, A. holotricha Finet & Gagnep., A.
davidii Franch., A. eriantha Benth., and A. fortunatu Finet & Gagnep.
For most of the other species, the number of specimens examined by
him were few. With our present more abundant material, the nature
of many of the species, including their variation, distribution, and
taxonomic delimitation, can be more fully understood and more ade-
quately interpreted.

An account of the early history of the genus is given in Dunn’s
paper and need not be repeated here. He also presented a discussion
of the affinities of the genus, which together with other related genera,
was then generally included in the Ternstroemiaceae or Dilleniaceae,
and he reached the following conclusion concerning Actinidia and
closely related genera: “Saurauja is inseparable from Ternstroemiaceae,
while the remaining two, Actinidia and Clematoclethra, can be properly
excluded from Dilleniaceae.” A search into this problem is not herein
attempted. It may be mentioned that the prevalent view at present is
to establish the family Actinidiaceae for these three genera. In Gilg

Werd ’s treatment (in Engler & Prantl, Nat. Pflanzenf. ed. 2.

*Duwnn, 8. T. A revision of the genus Actinidia Lindl. Jour. Linn. Soc. Bot.
39: 394-410, map. 1911.

2 JOURNAL OF THE ARNOLD ARBORETUM [VoL. XXXII

21: 36-47. 1925), the three genera are considered as constituting the
family Actinidiaceae, but each represents a distinct subgenus, to which
is added a fourth represented by Sladenia. The last-named genus, how-
ever, is very different from Actinidia and Clematoclethra and should not
be associated with these two. Kobuski (Jour. Arnold Arb. 32: 403.
1951) has retained Sladenia in the Theaceae (Ternstroemiaceae), as
originally proposed.

In Dunn’s paper, full bibliographic citations for each species are
given, and these will not be repeated in the present treatment. Aside
from the original citation for each species, only literature subsequent
to Dunn’s paper is here listed. A few misapplied names of taxonomic
significance are given, but no attempt is made to list all such records.
Nakai (in Bot. Mag. Tokyo 47: 251-259. 1933) has enumerated many
such names in synonymy, particularly with regard to Japanese plants;
I think that, at least in some cases, this contributes little to the proper
understanding of the species concerned.

Dunn’s key to the species is constructed on a very sketchy basis,
with only one character to each item, and in general it is unsatisfactory.
The different species are enumerated but not described, which renders
attempts to use his account for identification purposes rather difficult.
He divides the genus into four sections, giving particular emphasis to
the degree of pubescence, shape of ovary, and presence or absence of
lenticels on the fruit. After reviewing all the species, I find it desirable
to modify his system somewhat. A very important and fundamental
character of the genus, namely, the structure of hairs on the leaves,
was neglected by Dunn. This is far more significant in the differen-
tiation of the species-groups than such relative characters as the de-
grees of pubescence and the shape of the ovary. Both stellate-haired
and simple-haired species were included in his section Vestitae. This
section is here divided into two, Stellatae for species with stellate hairs
and Strigosae for those with simple hairs. For the glabrous or nearly
glabrous species, the section Maculatae of Dunn for species with spotted
fruit is retained. For species with unspotted fruit, Dunn originally
proposed two section, Ampulliferae for those with bottle-shaped ovary,
and Leiocarpae for A. kolomikta Maxim., a species with cylindric
ovary. However, this is also not a fundamental character, as such a
species as A. tetramera Maxim. has the ovary intermediate in shape.
Thus these two sections are now combined into one, Leiocarpae. As a
result there still remain four sections in the genus, bul they are modi-
fied in circumscription as compared with Dunn’s classification.

The present study accounts for 36 species and a number of varieties.
Three species, six varieties, and one form are described as new, and
several new combinations are proposed. Many synonyms are listed for
the first time, and the species and varieties are described.

The species of Actinidia, being all climbing or straggling plants, are,
like most other woody vines, very variable in vegetative structures.
They are dioecious plants, and often there is a slight vegetative dif-

1952] LI, REVIEW OF THE GENUS ACTINIDIA 3

ference between staminate and pistillate individuals, though rarely
of a sharp or striking nature. Such differences, if pronounced, will be
mentioned in the descriptions. It appears that Actinidia is in general
more variable in the individual plant than most other ligneous vines;
shoots from the same plant borne at different seasons often bear very
different leaves, variable in pubescence as well as in shape. The large
number of synonyms indicates the lack of appreciation of this fact on
the part of some of the describing taxonomists.

Dunn happily had the opportunity of collecting and studying
Actinidia in the field, and so his judgments are mostly sound and re-
liable. He calls special attention to the variable nature of this genus,
remarking: “The long arching shoots which appear during the spring
and summer months give rise in the following year to secondary
branches bearing leaves which are often strikingly different in shape
and character from those on the primary stems —a point which will
explain some anomalies in the series of specimens in herbaria.” As a
matter of fact, a number of supposedly new species of later years are
based on single anomalous specimens.

The distribution pattern is instructive in revealing the nature of the
species in Actinidia. Most of the species appear to be polymorphic
and to occur in very wide areas, comprising several varieties, mor-
phologically slightly different and geographically often of contiguous
but distinct areas. The type species of the genus, A. callosa Lindl., is
the most variable and also the widest in distribution. It is one of
the two that extend from temperate Asia to the tropical regions of
Malaysia, the other being A. latifolia (Gardn. & Champ.) Merr., which
is less variable in nature. All other species of the genus are primarily
of the hills or mountains of temperate regions. Those of wider ranges,
extending from Japan through northeastern Asia to western China, are
A. polygama (Sieb. & Zuce.) Maxim., A. arguta (Sieb. & Zucc.)
Planchon, and A. chinensis Planchon. Geographical varieties of slight
morphological differences are recognizable in all these species. The
majority of the species are of less wide range and less variable nature,
occurring in Japan, Taiwan, Tonkin, northeastern India, and in all
temperate and subtropical provinces of mainland China. Most of them
occur in southwestern China, which is the present center of develop-
ment of the genus. The species are generally common plants in the
thickets of the region and occupy fairly broad ranges. Only a few,
as records stand at present, are of very limited ranges, such as A.
rudis Dunn and A. rubricaulis Dunn of southern Yunnan and the three
new species herein proposed, but future explorations may prove that
they are also of wider distribution.

In citing specimens, the general order of north to south is followed.
An attempt has been made to standardize the geographical place names
of China, which are very confusing on field labels as well as in botani-
cal literature. With the exception of a few which cannot be de-
ciphered, which will appear in parentheses, all names follow those

4 JOURNAL OF THE ARNOLD ARBORETUM [VoL. XXXIII

given in “Gazetteer of Chinese Place Names based on the Index to
V. K. Ting Atlas,” June 1944, published by the Army Map Service,
Washington, D. C

Actinidia is of economic importance because of the fruits. Actinidia
chinensis and A. arguta, well known as Yang-tao in China, have long
been used for their edible fruits, which have a greenish pulp of pleasant
acid taste. The fruits are collected from wild plants. Actinidia arguta
is common in northern China while A. chinensis is especially common
along the Yangtze valley. Many other species have similarly edible
fruits. Recent efforts in introducing these species into cultivation and
in improving their products are highly desirable and commendable.
Actinidia chinensis, with its densely hairy leaves and large yellowish
flowers, is also highly ornamental. Actinidia polygama and A. kolo-
mikta have very decorative leaves variegated with white or pinkish
blotches and have become desirable ornamental plants.

The material used in this study has been assembled from the fol-
lowing herbaria, to whose curators the writer is indebted for their
generosity in permitting the use of specimens. Corresponding abbrevia-
tions are used in the citation of specimens. Arnold Arboretum (A) ;
Gray Herbarium, (GH); New York Botanical Garden, (NY); U.S.
National Herbarium, (US).

This study was carried on in the Department of Botany of the U.S.
National Museum, Smithsonian Institution, to whose officials grateful
acknowledgment is made for their kindness in placing the library and
herbarium facilities at my disposal. The writer wishes to express espe-
cially his thanks to Dr. A. C. Smith, Curator of Phanerogams, for his
kindness in reading the manuscript.

ACTINIDIA
Actinidia Lindley, Nat. Syst. ed. 2. 489. 1836; Benth. & Hook., Gen. PI.
1: 177. 1862.

Type species: A. callosa Lindl.

Climbing shrubs, glabrous, strigose, or tomentose, the indumentum
of stellate or simple hairs; pith solid or lamellate; bark often with
linear lengthwise lenticels; winter buds very small, enclosed in the
swollen base of the petiole. Leaves simple, alternate, usually long-
petiolate, serrate or dentate, rarely entire, penninerved, the costa
usually sulcate, the veinlets reticulate, usually in cross bars; stipules
minute, obsolete or absent. Flowers white, yellow, or reddish, poly-
gamous or dioecious, usually 5- or 4-merous, in axillary often pseudo-
umbellate cymes of few or many flowers, sometimes solitary; bracts
generally present, minute, 1 or 2 at the apex of the peduncles. Sepals
5, rarely 24, imbricate, rarely valvate, free or slightly connate at the
base, persistent. Petals 5, rarely 4 or more than 5, convolute, thin.
Stamens numerous, in pistillate flowers usually with shorter filaments
and smaller sterile anthers; filaments slender; anthers versatile, at-

1952] LI, REVIEW OF THE GENUS ACTINIDIA 5

tached at the middle, reflexed in bud, usually divaricate at base,
dehiscing lengthwise, yellow, brown, or purple. Disc absent. Ovary
free, superior, tomentose or glabrous, ovoid, cylindrical, or bottle-
shaped, many-celled; ovules attached on the central axis; styles many
(15-30), free, persistent, radiating, elongating after flowering, the tip
stigmatic, excurrent; rudimentary ovary in staminate flowers very
small, with minute styles. Fruit a berry, glabrous or sometimes hairy,
globose or oblong, spotted with lenticels or not, containing raphides.
Seeds numerous, small, biconvex, oblong, immersed in pulp; testa
cartilaginous, reticulate-pitted, dark when dry; albumen copious;
embryo comparatively large, cylindrical, straight, the cotyledons short.

About 36 species from Sakhalin and eastern Siberia, Japan, and
China to the Himalayas and Malaysia; the majority of the species are
found in western to eastern and southern China.

SYNOPSIS OF THE SECTIONS.

Sectio I. Strigosae sect. nov. (Vestitae Dunn, p. p.)

Caulis et petioli longe strigosi, foliis plus minusve setosis, baccis
maculatis.
Sectio II. Maculatae Dunn.

Caulis et petioli plerumque glabri, foliis glabris, baccis maculatis.
Sectio III. Leiocarpae Dunn (Ampulliferae Dunn).

Caulis et petioli glabri, foliis glabris, baccis emaculatis.
Sectio IV. Stellatae sect. nov.

Planta plus minusve tomentosa; indumento sub folio stellato, baccis
glabris vel hirsutis, maculatis.

Key TO THE SPECIES.

A. Plants glabrous or hairy, the pubescence on the under surface of the leaves
when present simple, not stellate
B. Stems, especially young shoots, and sometimes petioles covered with
yellowish or brownish strigose hairs; leaves more or less setose on one
or both surfaces (Sect. Strigosae).
C. Mature leaves more or less densely setose or strigose above, the bases
rounded to cordate.
D. Leaves cordate; petioles long, 3-7.5 em. in length.

E. Leaves ovate; sepals puberulous without ......... 1. A. rudis.
EE. Leaves obovate; sepals glabrous without . . ..2. A. rubus.
DD. Leaves oblong-lanceolate, not cordate; petioles short, about
do] 5 ont. 1OD8.. ee oe esa aes ee 3. A. melliana.

CC. Mature leaves glabrous or very sparsely hairy above.

D. Leaves ovate to broadly ovate, generally less than 1-1/2 times as
long as broad, the base rounded to cordate, the lower surface usually
strigose, not glaucous, the veins or main veinlets ending distinctly
in the marginal teeth.

E. Petioles long, slender, 5-8 em. long; veinlets subconspicuous.
4. A. holotricha.

JOURNAL OF THE ARNOLD ARBORETUM [VoL. XXXII

EE. Petioles short, 1.5-3 em. long; veinlets conspicuous, strongly
5. A. petelotir.

raised below .............. 00000 e eee eee

DD. Leaves narrowly ovate to oblong-lanceolate, generally two or

more times as long as broad, the base cuneate, rounded to cordate,

the upper surface glabrous, not strigose, the lower surface
glabrous or not, the veins or veinlets strongly anastomosing.

E. Leaves ovate, slightly coriaceous, the base rounded to distinctly

cordate; petioles short, 2-5 ¢

. Leaves dark above, pale beneath, the base distinetly cordate;

petioles very short, to 2.5 cm. long ........... 6. A. henryt

FF. Leaves concolored, the base rounded, not cordate; petioles

hes Ole ONE bio 5 4 os os sek eee an A. strigosa.

EE. Leaves sblonelancaolata, rarely ovate, stadrisneo i: the base
cuneate to rounded, rarely cordate; petioles usually long, over
2 cm. 1 g
F. Leaves dark above, pale and glaucous oo the base cuneate
to narrowly rounded, never cordate (S. W. China).
A. hemeleyana.

FF. on aves ianenlinad: the lower sdtiauk green, not glaucous, the

base broadly rounded to cordate (Taiwan). ... .

a Heed Gao’ A. artsanensis.

BB. Stems and neticles nearly alwa ays s glabrous; leav es s generally neptee
or sometimes sparsely setose along the veins or pubescent in
axils on the lower surface, rarely thinly tomentose beneath ee the
veins.

C. Fruit not spotted with pale lenticels; ovary cylindric to bottle-shaped,

strictly glabrous (Section Leiocarpae)

D. Ovary cylindric; pith brown, lamellate ........10. A. kolomikta.
DD. Ovary sea shaped, rarely cylindric; pith white or brown, solid
or lamellat

E. Pith solid. w an rarely slightly lamellate in the center.
F. Calyx of 5 distinet sepals, imbricate; pith are, solid, white.

ede eo eae peat ebes bake ada . polygama.
FF. Calyx of 2 or 3 beens ie pith a id white, or
rarely slightly lamellate. . 12. A. valvata.
EE. Pith lamellate, brown or sometaue se iiia
F. Leaves small, rarely large, elliptic to oblong-ovate, to 10 em.
long and 5.5 em. broad; petals not brownish at base; anthers
yellow.
G. Flowers 4-merous, rarely 5-merous, white, sometimes tinged
with pink. 0... 13. A. tetramera.
GG. hee 5-merous, white tinged with red especially toward
the margins. | 14. A. maloides.

FF. Leaves small or large, elliptic to ovate; peti ae usually brownish
at base; anthers purple
G. Leaves small, narrower, to 9.5 em. long and 4.5 em. broad.
H. Leaves not glaucous beneath. ...... 15. A. kwangsiensis.
HH. Leaves glaucous beneat
I. Leaves thicker, shorter, and broader, ovate, to 7 cm.
long and 4.5 em. broad; pith brown, lamellate to nearly

1952] LI, REVIEW OF THE GENUS ACTINIDIA 7

solid; Hae usually brownish at base; fruit larger,
about 2.5-0m: across (W. CHA) oc. ices eben desu as
16. A. melanandra.
Leaves thinner, longer, and narrower, ovate to oblong-
lanceolate, to 9.5 em. long and 4.3 em. broad; petals
not brownish at ee fruit smaller, about 1.5 cm. across
Aree 17. A. hypoleuca.
GG. Leaves larger, broader, over 8 cm. long and 4.5 em. broad.
H. Leaves elliptic to elliptic-oblong, the serrations appressed ;
fruit obovoid or oblong, purple; pith white (S. W. China)
18. A. purpurea.
HH. "Leaves broadly ovate, the serrations sharp, spreading ;
fruit ace greenish: pith white or brownish (Japan,
Korea, N. China, Liukiv).............. 19. A. arguta.
CC. Fruit spotted with aie lenticells: ovary slightly to densely pubescent
at first, becoming glabrate or pubescent when mature.
D. ze tke ae narrow, 3 or more times as long as broad; ovary soon
glabra

E. pee cuneate to subrounded at base; flowers whitish. .........
0. A. rubricaulis.

EE. Leaves distinctly cordatulate at base; flowers red...
ee ea ea 21. A. T iaratik
DD. Leaves broader, twice or less as long as broad; ovary pubescent.

E. Flowers red; leaves 1-1/2 times or more as long as broad.

F. Leaves coriaceous, the veins indistinct, the base acute to acumi-

—
—

nate (Szechuan, Kweichow, Yunnan)....... 2. A. coriacea
FF. Leaves membranaceous to chartaceous, the veins distinct, the
base auriculate-cordate (Kwangsi). 23. A. asymmetrica.

EE. Flowers white or yellow; leaves 1- 1/2 times or less as long as

roa
F. Flowers yellow; pith large, white, lamellate; leaf-bases broad-

gh Pi a 2 Bae I RS ales 5 Sora dig 24. A. pilosula.
GG. Leaves thinly chartaceous, glabrous or sparsely setose above,
not pubescent
. Leaves with numerous distinct parallel cross bars in the
veinlets, these strongly raised below, the lower surface not
glaucous. PO cee ee es oe ne eee 20, i venosa.

js

tro oe aC (0) . tric sh coutia,

FF. Flowers hie pith small, brown, jamellate or solid and
orange- -coloned: leaf-base narrowe

G. Pith small, brown, lamellate; flowers small, about 1 cm.

across; leaf-margins subentire to inconspicuously mucronu-

late.
H. Leaves subcoriaceous, larger, over 6 cm. long and 3 ecm.
broad, not glaucous (Kwangsi) .......... 27. A. glabra.

jadak Lekue chartaceous, smaller, scarcely to 6 cm. long and
Rs ec: Oe er 28. <A. sabiaefolia.

8 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXXIII

GG. Pith light orange-colored, solid or rarely shghtly and irregu-
larly lamellate; flowers sareer, about 2 em. across; leaf-
margins serrulate, rarely su re. ...... 29. A. callosa.
AA. Plants glabrous or hairy; leaves’ more or a densely tomentose with
stellate hairs on the lower surface (Section Stellatae).
B. Inflorescence large, 10-flowered or more, the peduncles 2-3 times
branched; fruit glabrate. 9.000220. 30. A. latifolia.
BB. Inflorescence smaller, 1- 5 (rarely to 10-) -flowered, us peduncles
once-branched or not branched; fruit glabrous to ae
. Leaves large, oblong to broadly ovate or orbicular, over 7 em. long and
4 em. broad, rounded or cordate at base; stellate tomentum on under
surface of the leaves not ei fruit large, more than 2.7 cm.
long or across, hairy or glabra
D. Leaves broadly ovate to orbieular, to 1-1/2 times as long as broad;
indumentum ai, fruit large, about 3 cm. across, densely hir-
sute or villos
EK. Young ce villose; leaves coriaceous, acute to acuminate, about
1-1/2 times as long as broad; flowers purplish.

Q

oo
_

A. eriantha.
EE. Young shoots setose; leaves chartaceous, truncate to emarginate,

rarely acute to short-acum inate, usually as long as broa
wers white to yellowish. ................ 32. A. chinensis.

DD. Leaves ovate to ovate-oblong, about 2 times as long as broad;
indumentum brownish; fruit small, about 2 cm. across, gl iiata:
flowers white. ©... .... 88. A. fulvicoma.
CC. Leaves small, ovate- lanceolate, to 7 em. long and 3 em. across,
strongly cuneate at base; stellate tomentum on under surface of

leaves appressed; fruit very small, scarcely 1 em. long, glabrous;
flowers greenish. Mee RESS. AGRO EERE ous OY Ree 34. A. Innsedlats,
Imperfectly known species. .....................0.... 35. A. kiusiana.
bee Aveo e Ls eee eee ... 36. A. longicauda.

1. Actinidia rudis Dunn in Jour. Linn. Soc. Bot. 39: 408. 1911.
Large climbing shrubs; stems and petioles densely rigid-strigose, the
hairs brown to light brown or yellow; pith white, lamellate. Leaves
chartaceous, ovate, 12-15 em. long, 7.5-9.5 em. read. acute to acu-
minate at apex, steel oislatia cordatulate at base, the margins minutely
denticulate, sparsely to densely strigose above, densely strigose along
the costa, veins and veinlets beneath, the upper surface dark, the lower
paler, the costa and veins slender, inconspicuous above, raised and
subconspicuous beneath, the veinlets reticulate, inconspicuous, the
secondary veins about 9-11 per side, straight ascending, anastomosing;
petioles variable in length, 1.5-7 em. long, densely strigose. Inflores-
cence densely fasciculate, 3—-5-flowered, densely ferrugineous-tomentose;
pedicels about 5 mm. long. Flowers white; sepals 5, ovate, about 3 mm.
long and 2 mm. broad, acute at apex, slightly puberulous without;
petals 5, oblong-ovate, 5-6 mm. long, 3-4 mm. broad, rounded at apex;
stamens numerous, the filaments about 2 mm. long, the anthers yellow,
linear, about 1 mm. long, rounded at apex, slightly sagittate at base;
ovary oblong densely villose, the styles about 2 mm. long. Fruit

1952] LI, REVIEW OF THE GENUS ACTINIDIA 9

cylindric to oblong, to 1.7 cm. long and 1 em. across, slightly tomentose
to glabrescent, lenticellate.

Southwestern China, in southern Yunnan only, in mountain forests
and in ravines, at altitudes of about 1200-1650 meters. Flowers white,
May—June.

CHINA: Yunnan: Meng-tzu, A. Henry 11307 (A, NY, US, tsorypgs) ;
P’ing-pien Hsien, H. T. Tsai 55137, 55205, 55429, 6O0073A, 60765, 61052,
61944 (A).

Henry originally cited 2 collections, Henry 11307 and 11335, both
fruiting specimens. These were first included in A. henry: by Dunn.
Subsequently he established the species A. rudis, basing it on these two
collections. Flowering materials are now available in Tsai 55205 and
others. The species is distinct in its yellow to light brown strigose hairs
on the leaves and stems and in the auriculate-cordatulate leaf-bases.

2. Actinidia rubus Lév. in Rep. Sp. Nov. 12: 282. 1913, Cat. PI.
Yiin-Nan 270. 1917; Rehder in Jour. Arnold Arb. 15: 97. 1934.

Climbing shrubs; branches grayish, suleate, densely brownish-
strigose, with pale lenticels; pith white, small, lamellate. Leaves
chartaceous, obovate to obovate-oblong, shortly but distinctly acumi-
nate at apex, broad and cordatulate at base, the margins irregularly
setose-serrulate, the teeth long or short, spreading, the upper surface
dark, sparsely scattered-strigose, the lower surface paler, strigose
along the costa and veins, the costa and veins subconspicuous above,
distinct and elevated beneath; the secondary veins about 7-9 per side,
straight-ascending, the veins or their main branches ending in the longer
marginal teeth, the veinlets reticulate, subconspicuous above, with more
distinct cross bars beneath; petioles slender, 5-6 cm. long, densely
strigose. Flowers solitary or few-fascicled, yellow; pedicels 1-1.3 cm.
long, strigose; sepals 5, ovate, unequal, to 6 mm. long and 3.5 mm.
broad, acute to acuminate at apex, glabrous without; petals 5, obovate,
more or less unequal, 10-11 mm. long, 5-7 mm. broad, rounded at
apex; stamens numerous, the filaments slender, about 4 mm. long, the
stamens yellow, about 1.2 mm. long, acute at apex, sagittate at base;
ovary ovoid, about 2.5 mm. long, densely pubescent, the styles about
3 mm. long. Fruit unknown.

In southwestern China, in northern Yunnan only. Flowers yellow,
June.

CHINA: Yunnan: Chao-t’ung, E. E. Maire s. n. (A, fragments of TYPE).

The material on hand is more or less fragmentary, but no additional
material is available. The species appears to differ from the closely
related A. holotricha Finet & Gagnep. mainly in the obovate leaves
with spreading teeth and in the glabrous sepals.

3. Actinidia melliana Hand.-Mazz. in Anz. Akad. Wiss. Wien, Math.-
Nat. 59: 57. 1922, in Beih. Bot. Centralbl. 48(2): 306. 1931; Merr

10 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXXII

in Lingn. Sci. Jour. 7: 315. 1929; Merr. & Chun in Sunyatsenia 2:
283.1935; Chun in Sunyatsenia 4: 190. 1940.

Climbing shrubs to 10 m.; branches, inflorescences, and_ petioles
densely covered with long rigid brown hispid hairs to 8 mm. in length;
pith white, lamellate. Leaves membranaceous to chartaceous, per-
sistent, oblong to ovate or obovate-oblong, 8-18 em. long, 2.5-8 em.
broad, shortly acuminate at apex, broadly rounded and distinctly cor-
datulate at base, with brown-hirsute hairs on both surfaces when
young, becoming scattered hispid-hirsute above, glabrous throughout
or with hirsute hairs along the costa and veins beneath, often glaucous,
the margins entire, with numerous sharp rigid pointed hairs, the costa
and veins slender, slightly conspicuous above, distinct and raised be-
neath, the secondary veins about 6 or 7 per side, arcuately ascending,
anastomosing near the margins, the veinlets reticulate, subconspicuous
on both surfaces; petioles short, 10-15 mm. long, more or less terete,
densely hirsute. Inflorescences in axillary cymes, about 10-flowered,
subsessile, shortly dichotomously branched, densely brown-hirsute;
pedicels slender, to 12 mm. long; bracts subulate, 4-5 mm. long, elongat-
ing to 6 mm. in fruit. Flowers white: sepals oblong-ovate, obtuse to
acute at apex, 4-5 mm. long, hirsute without; petals white, ovate to
obovate, rounded to acute at apex, 8-9 mm. long, 6-7 mm. wide; stamens
numerous, the filaments 2 mm. long, the anthers 1 mm. long, sagittate-
ovate; ovary subglobose, densely hirsute. Fruit strigose-hirsute to
glabrescent when mature, oblong, 16-22 mm. long, 11-15 mm. across,
strongly verrucose, with small white rounded lenticels; styles persistent,
to 3 mm. long.

Southern China (Kwangsi, Kwangtung, southern Kiangsi, and
Hainan), in forests from 800 to 1350 meters. Flowers white, June.

CHINA: Kiangsi: Lung-nan Hsien, S. K. Lau 4429 (A, US). Kwang-
tung: Lung-tau Mountain, R. Mell 22 (A, 1sorypr), Canton Christ. Coll.
12085 (US), 12175 (NY, US). Hainan: “Fan Yah,” N. K. Chun & C. L.
Tso 44219 (A, NY, US).

This species is a distinct one, strongly characterized by its long
indumentum and the oblong distinctly cordatulate leaves, green or
glaucous beneath. The leaf-margins are entire in general appearance,
the very fine teeth appearing as pointed rigid hairs along the straight
edge of the leaves. The stems as well as the leaves and inflorescences
are covered by very long brown hispid hairs. The hairs on the leaves,
however, vary greatly in density. Sometimes these hairs are present on
both surfaces, distributed all over on the costa, veins, and veinlets.
At other times, they may be present all over the upper surface but
only very sparsely so on the costa and a few veins on the lower surface.
As the leaves advance in age, they apparently become more glabrous
and more glaucous on the under surface.

The species is evidently more closely related to A. hemsleyana Dunn
than to any other. It is, however, readily distinguished by its denser,

1952] LI, REVIEW OF THE GENUS ACTINIDIA 11

longer hairs and its distinctly cordate leaf-base. It also has a more
southerly range.

4. Actinidia Saar Finet & Gagnep. in Bull. Soc. Bot. France 52:
Mém. 4: 18, ¢. 3. 1905 (Contr. Fl. As. Or.); Dunn in Jour. Linn.
Soc. Bot. 39: ie 1911.

Climbing shrubs; branches and petioles covered with sparse brown
hispid hairs; pith white, lamellate. Leaves membranaceous to thinly
chartaceous, broadly oblong-ovate, 9-13.5 em. long, 6—7.5 em. broad,
short- to long-acuminate at apex, rounded to truncate at base, the
margins sharply and finely denticulate, the teeth ascending, sparsely
to densely setose along the costa and veins on both surfaces when
young to nearly glabrous when mature, the lower surfaces slightly
paler, the costa and veins slender, sub-conspicuous above, distinct and
raised beneath, the secondary veins about 7 or 8 per side, straight-
ascending, the veins or main branches ending in the marginal teeth,
the veinlets reticulate. inconspicuous above, subconspicuous beneath;
petioles long, slender, terete, 5-8 cm. long, sparsely strigose. Inflores-
cences glabrate or very sparsely strigose, shortly branched; pedicels
3-10 mm. long, slender. Flowers yellow (?); sepals ovate, obtuse,
5-6 mm. long, more or less puberulous without; petals oblong-obovate,
10-11 mm. long, 7-8 mm. broad, rounded at apex; stamens numerous,
the filaments to 4 mm. long, the anothers yellow, sagittate-ovate; ovary
subglobose, densely hirsute. Fruit unknown.

Southwestern China (Yunnan and southwestern Szechuan), at alti-
tudes of about 1400-2000 meters. Flowers yellow (?), May—June.

CHINA: Szechuan: Ma-pien Hsien, F. 7. ae 23086 (A). Yunnan:
Ping-pien Hsien, H. T Tsai 60483 (A), 62642 (A

This species was based originally on a a collection from
“Outchay,” which was not examined by Dunn. I suspect this type
locality is Hui-tse (Tong-ch’tian), in northeastern Yunnan. The speci-
mens here referred to this species are from nearby localities in Szechuan
and Yunnan. They are more or less fragmentary or sterile specimens.

5. Actinidia petelotii Diels in Notizbl. Bot. Gart. Berlin 11: 213.

Tall climbing shrubs; stems slender, the branches grayish, more or
less striated, the young branchlets long and densely ferrugineous-
strigose ; pith large, white, lamellate. Leaves chartaceous, ovate, about
9-14 cm. long, 6-9 cm. broad, long-acuminate at apex, cordate to sub-
cordate at base, the margins palloeccernulate, the upper surface green,
glabrous or very sparsely setose along the veins, the lower surface
paler, densely to sparsely setose along the costa and main veins, other-
wise glabrous, the costa and nerves slender, subconspicuous’ above, dis-
tinct and raised beneath, the secondary nerves about 6-8 per side,
arcuate-ascending, anastomosing, the branchlets ending in the mar-
ginal teeth, the veinlets reticulate, with more or less parallel cross-bars,

12 JOURNAL OF THE ARNOLD ARBORETUM [voL. XXXIII

distinct beneath; petioles 1.5-2.5 ecm. long, densely long-ferrugineous-
strigose. Inflorescence fasciculate, axillary, or on short branches, densely
ferrugineous-strigose; peduncles about 1.5 cm. long. Flowers not seen;
ovary pilose. Immature fruit oblong, about 1.5 em. long and 7 mm.
across, slightly pubescent or nearly glabrate, brown, with pale lenticels;
styles 2.5-3 mm. long; persistent sepals membranaceous, ovate, about
4 mm. long and 2.5 mm. broad, acute at apex, pubescent without.

Indo-China, known from Chapa, Tonkin, only, at an altitude of 1800
meters.

INDO-CHINA: Chapa, A. Pételot 3829 (A, NY, US, 1sorypss).

Specimens of the type collection are all young fruiting ones. No
additional material is available. Based on this somewhat incomplete
material, this species can be compared with A. strigosa Hook. f. &
Thoms., of India, which has smaller, narrower leaves concolored on
both surfaces and never cordate at the base, and peduncles which are
not strigose.

6. Actinidia henryi Dunn in Kew Bull. 1916: 1. 1916.

Large climbing shrubs; branches slightly striated, more or less hispid,
the young branchlets reddish brown-villose; pith small, lamellate,
whitish. Leaves chartaceous, oblong-ovate, 8-14 em. long, 3-6.5 cm.
broad, acuminate at apex, subcordate to cordate at base, the margins
minutely serrulate, dark above, pale beneath, glabrous on both sur-
faces except the veins, the costa and veins slender, subconspicuous
above, puberulous or setose, the secondary veins about 8-10 per side,
arcuately ascending, anastomosing near the margins, the veinlets reticu-
late, inconspicuous to subconspicuous above, conspicuous beneath;
petioles relatively short, 10-25 mm. long, ferrugineous-pubescent. In-
florescences in axillary cymes, 10-flowered or more, densely reddish-
villose; pedicels to 10 mm. long. Flowers white; sepals orbicular, acute
at apex, about 3 mm. long, pubescent without; petals ovate, the base
cuneate, rounded at apex, about 6 mm. long; stamens numerous, the
filaments about 2 mm. long, the anthers yellow, sagittate-ovate; ovary
subglobose, densely pubescent.

Key To VARIETIES

A. Leaves without rigid pilose hairs..................... a. var. henryt.
AA. Leaves with scattered rigid pilose hairs on the costa we veins on both
OE NOE 5555 eres nie Oss Ne de eee eee: b. var. polyodonta.

6a. Actinidia henryi Dunn var. henryi.
Actinidia henryi Dunn in Kew Bull. 1916: 1(excl. spec. Henry 11307,
13385). 1916, in Jour. Linn. Soc. Bot. 39: 407. 1911.

Leaves chartaceous, oblong-ovate, 10.5-14 em. long, 3-6.5 em. broad,
acuminate at apex, distinctly cordate at base, the margins very minutely
mucronulate-serrulate, glabrous on both surfaces except the veins, the
costa and veins reddish-puberulous below; petioles ferrugineous-pubes-
cent when mature, about 15-25 mm. long.

1952] LI, REVIEW OF THE GENUS ACTINIDIA 13

Southwestern China, in southern Yunnan only, in mountain forests
at altitudes of about 1650-2650 meters. Flowers white.

CHINA: Yunnan: Meng-tzu, A. Henry 10381 (US, tsoryre), 10381a (A
NY, US, tsoryprs); Chien-shui Hien, H.T. Tsai 53331 (A).

Dunn’s original description was based on, in addition to Henry 10381
and 10381a as listed above, also Henry 11307 and 13335, which he
subsequently segregated as representing a distinct species, A. rudis
However, he did not then redescribe the present specis. The above de-
scription was prepared by checking the differences between these speci-
mens and his original description, and also the additional collection
made by Tsai.

This species is readily distinguished from A. rudis by its short
petioles, distinctly cordate leaves, and less copiously strigose stems.
In Dunn’s key, A. henry? is separated from A. hemsleyana Dunn by its
leaves being glaucous beneath, while for A. hemsleyana the leaves are
mentioned as green beneath. Actually the reverse condition is true.

bg

6b. Actinidia henryi Dunn var. polyodonta Hand.-Mazz. Symb. Sin.
: 391. 19381.

Leaves chartaceous, oblong-ovate, 8-11.5 cm. long, 2.3-4.7 em. broad,
acuminate at apex, subcordate at base, the margins minutely but dis-
tinctly serrulate, with scattered rigid setose hairs along the veins on
both surfaces; petioles sparsely pilose, about 10-23 mm. long; i
mature fruits eylindric- oblong, sparsely pubescent.

Southwestern China, in central Yunnan and western Kwangsi, at
altitudes of 1650-2450 meters.

CHINA: Kwangsi: N. Lo-ch’eng, R. C. Ching 5897 (NY).

Handel-Mazzetti’s type, from K’un-ming, north of the type locality
of the typical form of the species, has not been seen. The Ching
specimen cited above, from western Kwangsi, close to the Yunnan
border, agrees with Handel-Mazzetti’s description. Compared with
the typical form of the species, this variety differs in having rigidly
pilose petioles, scattered setose hairs on both surfaces of the leaves, and
more distinctly serrulate leaf-margins. Ching 5897 is a young fruiting
specimen.

7. Actinidia strigosa Hook. f. & Thoms. in Jour. Linn. Soc. Bot. 5:
55. 1861; Thiselton-Dyer in Hook. f. FI. Brit. Ind. 1: 286. 1876;
Dunn in Jour. Linn. Soc. Bot. 39: 407. 1911.

Climbing shrubs; branches reddish brown, with scattered elongate
pale lenticels, hispid, the young branchlets more or less densely fer-
rugineeus-setose; pith large, whitish, lamellate. Leaves chartaceous,
ovate to oblong-ovate, about 7-13 cm. long and 4-7 cm. broad, acumi-
nate to long-acuminate at apex, obtuse to rounded at base, often
obliquely so, the margins callously denticulate, the surfaces concolored,
the upper surface glabrous or slightly puberulous along the costa and
veins, the lower surface nearly glabrous or sparsely setose along the

14 JOURNAL OF THE ARNOLD ARBORETUM [VoL. XXXII

costa and veins, the costa and veins inconspicuous above, distinct and
raised beneath, the secondary nerves about 5-7 per side, arcuate-
ascending, anastomosing, the branchlets ending in the marginal teeth,
the veinlets reticulate, with many parallel cross-bars, subconspicuous
beneath, petioles 2.5-3 cm. long, strigose or puberulous. Inflorescences
in short 2-4-flowered axillary cymes, ferrugineous-pubescent, the flow-
ers sometimes solitary; peduncles to 1 em. long; pedicels to 5-10 mm.
long; bracts minute, linear. Flowers white; sepals 5, ovate, about
4-5 mm. long and 3-4 mm. broad, acute to ‘obtees at apex, glabrate
or very sparsely puberulous without; petals 5, obovate, about 8 mm.
long and 5 mm. broad, rounded at apex; stamens numerous, the fila-
ments 2.3 mm. long, the anthers yellow, 1-1.56 mm. long, obtuse at
apex, sagittate’at base; ovary subglobose, about 1.5 mm. across, densely
villose, the styles about 1.5 mm

ane (Sikkim), at altitudes ss 2100-3300 meters. Flowers white.

INDIA: Sikkim: J. D. Hooker s. n. (GH, 1soryPe); eastern Himalaya,
Griffith 57 (GH).

This species seems to be confined to Sikkim Himalaya at fairly high
altitudes. It is characterized by its concolored, ovate to oblong leaves,
which are sparsely setose along the costa and veins on the lower surface
only.

8. Actinidia hemsleyana Dunn in Jour. Linn. Soc. Bot. 38: 355. 1908,
39: 407. 1911.

Actinidia subglaucifolia Metealf in Lingn. Sci. Jour. 11: 15. 1932. Syn. nov.

Actinidia kengiana Metcalf in op. cit. 16. Syn. nov.

High climbing shrubs to 10 m.; branches dimorphic, some long, arch-
ing, densely strigose-tawny especially when young and bearing narrow
leaves, others lateral, short, strigulosge to hispidulous and bearing both
leaves and flowers; buds densely brownish-hispidulous; pith lamellate,
brownish. Leaves chartaceous, oblong-ovate to lanceolate-oblong,
mostly 8-13 em. long, 2.5-4.5 em. broad, some to 18 em. long and 4 cm.
broad or 20 cm. long and 10 cm. broad, acute to obtuse at apex,
rounded or acute to cuneate at base, usually slightly obliquely so,
the margins serrulate to appressed-serrulate, glabrous on both surfaces
or brownish pubescent along the costa or veins beneath, the upper
surface dark, the lower pale, usually glaucous, the costa and veins
slender, subconspicuous above, distinct and raised beneath, the second-
ary nerves about 7 or 8 per side, arcuately ascending, anastomosing
along the margins, the veinlets reticulate, inconspicuous above, incon-
spicuous to subeonspicuous beneath; petioles 1.54.5 em. long, sparsely
hispid to glabrate. Inflorescences in axillary cymes of about 1-3 flow-
ers; pedicels 1-1.5 em. long, slender, densely brownish tomentose.
Flowers greenish; staminate flowers with 5 sepals, these ovate, sub-
acute, about 5 mm. long, densely brownish-tomentose; petals 5, glabrous,
ovate, about 10 mm. long, rounded at apex; stamens numerous, about
as long as the petals; rudimentary ovary 2 mm. across, tomentose.

1952] LI, REVIEW OF THE GENUS ACTINIDIA 15

Pistillate flowers similar but with sepals connate at base, the lobes
ovate, about 8 mm. long, acute at apex, densely brownish-tomentose at
apex; stamens rudimentary; ovary depressed-globose, about 6 mm.
long, densely brownish-villose. Fruit (immature?) cylindircal oblong-
ovoid, about 2.7 cm. long, and 1.5 em. across, densely brown-villose to
glabrate, not lenticellate, rounded at apex, crowned by the persistent
short styles of about 4 mm. long.

Eastern China (Fukien and southern Chekiang), in thickets on moun-
tain slopes at altitudes of 500-900 meters. Flowers greenish, June.

CHINA: Chekiang: Between P’ing-yang and T’ai-shun, R. C. Ching 2120
(A, US); Cheng-ning Hsien, Y. L. Keng 394 (A, rypr of A. kengiana Metcalf).
Fukien: Nan-p’ing Hsien (Yenping), Hongk. Herb. 2400 (A, ISOTYPE) ;
Shouning Hsien, Y. L. Keng 339 (A, TyPE of A. subglaucifolia Metcalf).

This distinct species has a known range limited to northern Fukien
and southern Chekiang. Two specimens of Dunn’s type collection,
Hongkong Herb. 2400, have been seen. Although Dunn’s key specifi-
cally indicates that the leaves of this species are green beneath instead
of glaucous, actually these two specimens have distinctly glaucous
leaves. The other collections here cited show that the leaves vary from
glaucous to subglaucous, but they are never concolored.

Type specimens of both species of Metcalf, in the Arnold Arboretum
herbarium, have been seen. They are leafy specimens with a few de-
tached young fruits, and I cannot separate them from A. hemsleyana.
Actinidia subglaucifolia Metcalf is based on a specimen with nearly
glabrous stems and leaves. Metcalf describes the plants as glabrous
and the leaves as glabrous above and somewhat glaucous beneath. He
has apparently overlooked the dense brown hairs on the buds and the
sparse strigose blackish hairs on the young stems and petioles. In
this specimen the leaves are glabrous throughout. Actinidia kengiana
Metcalf is represented by a stout twig with a few leaves. Among these
leaves some are of the size described by Dunn, but a few are excep-
tionally large. The young shoots are distinctly and densely strigose.
The leaves are, as described by Metcalf, “glabrous above, glaucous
and sparsely rusty-hairy beneath, especially along the veins.” Metcalf
compared his A. subglaucifolia with A. sabiaefolia Dunn, and his A.
kengiana with A. melliana Hand.-Mazz., but he failed to mention A.
hemsleyana Dunn.

Dunn, who collected this plant in the field calls special attention
to the dimorphism in the stems, and flowers, in this and in many other
members of the genus as well. He says that the habit of this plant seems
to illustrate the usual method of growth in many species of Actinidia
and explains some of the apparent anomalous specimens in the her-
barium. He remarks: “Each new extension of the shrub begins in this
case with a long arching, densely strigose tawny shoot bearing abnormal]
narrow leaves. In the autumn these leaves fall, leaving large thickened
leaf-scars and a bud above each, protected by a tuft of stiff yellow
hairs. These buds develop in the spring into short flower- and leaf-

16 JOURNAL OF THE ARNOLD ARBORETUM [voL. XXXII

bearing shoots, the leaves and the indumentum being quite different to
those primary shoots.

9. Actinidia arisanensis Hayata, Icon. Pl. Formos. 8: 11. 1911;
Sasaki in Trans. Nat. Hist. Soc. Formosa. 19: 480. 1929; Kanehira,
Formos. Trees, rev. ed. 448, f. 405. 1936.

Actinidia rankanensis Hayata, op. cit. 13.

Actinidia remoganensis Hayata, op. cit. 13.

Tall climbing shrubs to 5 m. or more; branches dimorphic, the spring
shoots glabrous to densely strigose, with narrower glabrous or sparsely
setose or more or less densely strigose leaves, the summer shoots
flowering, strigose to glabrous, with generally broader nearly glabrous
leaves; pith very small, brown, lamellate. Leaves chartaceous, ovate-
oblong to ovate, 8-15 cm. long, 3-9 em. broad, acute to acuminate at
apex, cuneate to rounded to subcordate at base, usually unequal, the
margins sparsely and finely denticulate, glabrous or strigose on both
surfaces or sparsely setose above and tomentose along the costa and
sometimes also along the veins beneath, concolored or slightly paler
beneath, the costa and veins inconspicuous to subconspicuous above,
distinct and raised beneath, the secondary veins 5-7 per side, straight-
ascending, the veins or their branches ending in the marginal teeth, the
veinlets reticulate, distinct and raised beneath; petioles usually long,
slender, 1.5-4.5 cm. long, densely or sparsely strigose or glabrous. In-
florescences in axillary cymes, 3- or 4-flowered, the pedicels nearly
glabrous, 5-10 mm. long; bracts minute. Flowers white; sepals 5,
oblong, rounded at apex, about 4 mm. long and 2 mm. broad, glabrous
without, the margins pubescent; petals 5, oblong-ovate to oblong-
lanceolate, about 7 mm. long and 3 mm. broad, rounded at apex,
cuneate and contracted at base; stamens numerous, the filaments
filiform, the anthers yellow, oblong, about 1.3 mm. long, obtuse at
apex, sagittate at base; ovary subglobose, densely tomentose; styles
about 4 mm. long. Fruit subglobose, about 2.3-3.5 cm. across, glabrate,
lenticellate.

China, in Taiwan only, in thickets and forests at altitudes of 1160-
2260 meters. Flowers white, April-May.

CHINA: Taiwan: Suao, £. H. Wilson 11122 (A, US); Seem near Giran,
E. H. Wilson 10255 (A); Arisan, FE. H. Wilson 9671 (A), 1 7 (A): Funkiko,
Arisan, FE. H. Wilson 9658 (A); Keitou, Arisan, F. H. ar at 10862 (A, US);
Arisan to Mt. Morrison, E. H. Wilson 10943 (A, US)

This is a very variable plant, with dimorphic soae vegetative and
summer flowering shoots. The young sterile shoots may be strigose
bearing densely strigose leaves as in Wilson 9658, or with nearly glab-
rous stems and leaves sparsely strigose above only as in Wilson 10225,
or with nearly glabrous leaves as in Wilson 11122. The flowering
shoots bear larger broader leaves, generally strigose all over the upper
surface.

In those specimens of nearly glabrous habit, it is difficult to dis-

1952] LI, REVIEW OF THE GENUS ACTINIDIA 17

tinguish this species from A. callosa var. formosana superficially. How-
ever, upon closer examination, strigose hairs can generally be revealed
from either young or old stems in A. arisanensis. The leaves of A.
arisanensis are also larger and relatively narrower, and they never
become obovate as in A. callosa var. formosana. The fruit of A.
arisanensis is also much larger. This species inhabits only the northern
central mountainous parts of the island, at high altitudes, while A.
callosa var. formosana is of the lower altitudes and is more widespread.

The variable nature of the species can be shown by the fact that
Hayata described at the same time three species which now cannot be
maintained. He distinguishes his A. arisanensis from his A. rankanensis
“by the leaves which are nearly obtuse or slightly cuneate at the base.”
Actinidia remoganensis Hayata is: “Near A. rankanensis, but, distin-
guishable from it in the less serrulate or nearly entire leaves and in the
narrower petals.” Photographs of type specimens of all three are
available. These and the original descriptions prove that Sasaki is
justified in reducing the two additional names.

The relationship of A. arisanensis is clearly with A. hemsleyana
Dunn of the nearby coastal provinces Chekiang and Fukien.

10. Actinidia kolomikta (Maxim. & Rupr.) Maxim. in Mém. Acad.
Sei. St. Pétersb. Sav. Etrang. 9: 63. 1859 (Prim. Fl. Amur.).

Climbing shrubs to 7 m.; branches usually dark, glabrous or the very
young branchlets slightly pubescent; pith brown, lamellate. Leaves
membranaceous, sometimes partly discolored or variegated, with a
large white to pink blotch at the apex often extending to the middle
or beyond, especially in the staminate plant, ovate to oblong-ovate,
6-15 cm. long, 3-12.5 em. broad, acuminate at apex, distinctly cordate
at base, more rarely subcordate to truncate, sometimes unequal, the
margins serrulate, concolored or slightly paler beneath, glabrous on
both surfaces to sparsely setose or pubescent along the costa and veins,
the costa and veins slender, inconspicuous above, distinct and raised
beneath, the secondary veins about 6-8 per side, slightly arcuately
ascending, anastomosing, the veinlets reticulate, inconspicuous on both
surfaces to subconspicuous beneath; petioles slender, 2.5-3.5 cm. long,
glabrous to sparsely pubescent. Flowers 1-3-fascicled; pedicels slen-
der, 6-10 mm. long, glabrous to rusty-tomentose; bracts minute, linear;
sepals, 5, ovate, 5-6 mm. long, 3-4 mm. wide, glabrous, acute at apex,
more or less connate at base; petals 5, oblong, about 10 mm. long and
5 mm. broad, rounded at apex, gradually narrowed at base; stamens
numerous, the filaments slender, 5-6 mm. long, the anthers yellow,
slightly sagittate; ovary cylindric-ovoid, about 3 mm. long and 2 mm.
across, glabrous; styles 3-5 mm. long. Fruit globose, about 2 em. in
diameter, glabrous, not lenticellate.

Actinidia kolomikta is here considered as composed of two varieties,
a typical form in eastern Siberia, Manchuria, Korea, and Japan, and
another variety in western China.

18 JOURNAL OF THE ARNOLD ARBORETUM (VoL. XXXII

Key To THE VARIETIES

A. Leaves generally smaller, slightly narrower, to 10 X 8 em., rarely larger,
with or mostly without scattered setose hairs; spare and_ pedicels
usually glabrate................. a. var. kolomikta.

AA. ete generally larger and broader, to 15 x 12 5 cm., often with scat-

tered setose hairs; penduncles and pedicels usually rusty tomentose

. var. gagnepainit.

10a. Actinidia kolomikta sean & Rupr.) Maxim. var. kolomikta.

Actinidia kolomikta (Maxim. & Rupr.) Maxim. in Mém. Acad. Sci. St.
Pétersb. Sav. Etrang 9: 63. 1859 ee Fl. Amur.) ; Dunn in Jour. Linn.
Soc. Bot. 39: 404. 1911

Prunus? kolomikta ‘Maxim & Rupr. in Bull. Phys. Math. Acad. Sci. St.
Pétersb. 15: 129.

Kolomikta see ‘Regel in Bull. Phys. Math. Acad. Sci. St. Pétersb.
15: 219. 1857.

Trochostigma kolomikta Rupr. in Bull. Phys. Math. Acad. Sci. St. Pétersb.
15: 262. 1857.

Leaves ovate to oblong-ovate, sometimes discolored or variegated,
about 6-11 em. long and 3-8 cm. broad, glabrous on both surfaces to
sparsely or slightly setose above and sparingly pubescent along the
costa and veins beneath; peduncles and pedicels glabrous to sparsely
pubescent.

Eastern Siberia, Sakhalin, Manchuria, Korea, and Japan, in thickets
at altitudes of 150-1600 meters. Flowers white, nner

EASTERN SIBERIA: Amur, Mazimowicz s. n. (G , US), S. #.
Enander s. n. (A), Korzinsky s. n. (A, GH, US), V. pena 1088 (US);
Vladivostok, NV. Palczewsky s.n. (A, NY), D. L. Topping 2168 (A), Golden-
staedt s.n. (GH), C. 8. Sargent s. n. (A).

SAKHALIN: Sakhalin, F. Schmidt s. n. (GH), G. Faurie 469 (A); Ohdo-
mari, K. Uno 19923 (A. US), FH. H. Wilson s. n. (A).

MANCHURIA: Ad fl. Amur, R. Maack s. n. (GH); Er-tao-tien-tzu, P. H.
& D. H. Dorsett 3068 (US); Kao-ling-tzu, P. H. & D. H. Dorsett 5985
(A, N, US), V. Skvortzov s. n. (A); Mifun Station, V. Skvortzov s. n. (A);
Mao-erh-shan Station, V. Skvortzov s. n. (A); Hsing-an, V. Komarov 1088
ne NY); Kirin, O-mu Hsien, H. W. Kung 1897 (NY); coast of Manchuria,

C. Witford s. n. (GH)

KOREA: Taiyudo, prov. N. Heian, 2. H. Wilson 8604 (A, US); Kongo-san,
prov. Kogen, E. H. Wilson 10485 (A, US)

JAPAN: Hokkaido: Shibetsu, K. Miyabe s. n. (A); Hyukogen, K.
Uno 16305 (A); Iburi, S. Hashimoto s. n. (A); Konoma, Mazimowicz s. n.
(GH); Mt. Moiwa, BE. Tokukuchi s. n. (GH); Kushiro, EF. H. Wilson s. n.
(A); Morinan jama FE. H. Wilson s.n. (A); Meakan, T. Tanaka 222 (A, US).
Hondo: Sinano, K. Shiota 3320 (A); Mino, K. Shiota 4904 (A), 9698 (A),
M. Kentaro 3322 (A); Fujiyama, P. H. Dorsett & W. J. Morse 374 (A, US);
Huzisan, 7. Sawada 2230 (A); Juzogatake, G. Faurie 5390 (A); Rikuchu,
E. H. Wilson s. n. (A); Lake Chuzenji, £. H. Wilson s.n. (A), J. G. Jack s. n.
(A); Miyachine, G. Faurie 6912 (A).

1952] LI, REVIEW OF THE GENUS ACTINIDIA 19

10b. Actinidia ae (Rupr. & Maxim.) Maxim. var. gagnepainii
(Nakai), comb. n
Actinidia gagnepainn ne in Bot. Mag. Tokyo 47: 258. 1933.
Actinidia kolomikta sensu Hemsl. in Jour. Linn. Soc. Bot. 23: 79. 1886,
p.p.; Dunn in Jour. Linn. Soc. Bot. 39: 404. 1911, p.p.; Rehder in Sarg.
Pl. Wils. 2: 380. 1916; non Maxim.

Leaves ovate to ovate-oblong, sometimes discolored or variegated,
about 6-15 em. long and 5—12.5 cm. broad, usually with scattered setose
hairs along the veins on both surfaces, sometimes glabrous; peduncles
and pedicels more or less rusty-tomentose.

Western China (Shensi, Hupeh, Szechuan, and Sikang), in thickets
at altitudes of 1800-3600 meters. Flowers white, May—June.

INA: Hupeh: Western Hupeh, A. Henry s. n. (NY), 5622 (NY),
6922A (NY), 8806 (GH). Szechuan: Kuan Hsien, FH. H. Wilson 2008
(A, US), 2009 (A, GH, US), W. P. Fang 2239 (A), F. T. Wang 20591 (A);
Wen-ch’tian Hsien, 2. H. Wilson 1058 (A), ees (A, US), F. T. Wang 21097
(A); O-mei-shan, F. H. Wilson 4761 (A, US), W. P. Fang 2801 (A, NY),
2865 (A), F. T. Wang 23356 (A), W. W. Tae 79 (A), S.C. Sun & K. Chang
930 (A), 1141 (A), 1358 (A), H. C. Chow 8235 (A), 12379 (A), Wa-wu-shan,
E. H. Wilson 854a (A, GH, US); Nan-ch’tian Hsien, W. P. Fang 910 (A,
NY); O-pien Hsien, T. T. Yii 856 (A); Ma-pien Hsien, F. T. Wang 23010
(A), 23093 (A); between Hai-tang and P’ing-yu-p’u, H. Smith 1986 (A);
P’ing-shan Hsien, F. T. Wang 22704 (A); “Tsing-chuan-fin Hsien,” F. 7.
Wang 22350 (A). Sikang: No precise locality, H. A. Pratt 101 p.p. (GH);
K’ang-ting Hsien (Tachienlu), 2. H. Wilson 2005 (A, GH); Yueh-sui Hsien,
T.T. Yii 948 (A).

Nakai separates the western Chinese plant as a distinct species,
A. gagnepainu, basing it on Henry 8806, 8994, Wilson 4761, Pratt
830, and David s. n. According to him, the western Chinese plant
differs from A. kolomikta as follows: ‘The Chinese Actinidia has more
vigorous shoots, broader leaves with more brownish and thicker hairs
on the veins, and more rigid hairs on the surface. The leaves never
become white or crimson like A. kolomikta. Its flowers are similar to
those of A. kolomikta but the pedicels are more robust.”

These characters, as can be easily seen, are all relative and neces-
sarily inconstant. There are no fundamental structural differences
between the plant from northeastern Asia and that of western China.
Rehder, following Dunn, gives A. platyphylla A. Gray ex Miq. as a
synonym of A. kolomikta and remarks: “The plant from western China
does not show any obvious difference from the Japanese plant except
that the leaves are generally larger and broader.”

Leaving out for the time being the problem of typification and
identity of A. platyphylla, which will be discussed in connection with
A. arguta, it is sufficient to say that the western Chinese plant, for
which a much larger series of specimens than at Nakai’s disposal is
now at hand, does not show fundamental differences which warrant
specific separation from the typical form of A. kolomikta. Instead,

20 JOURNAL OF THE ARNOLD ARBORETUM [voL. XXXII

one is impressed by the close similarity and resemblance in all respects
between the plants from the two areas. Recognizing two separate and
distinct species on the basis of such slender grounds would obscure
entirely their close genetic kinship. Nakai mentions that the leaves
of the western Chinese plant “never become white or crimson like A.
kolomikta.” This is an erroneous statement, as in the large series of
specimens now available, many specimens, such as Wilson 854a, 2005,
and 2008, have equally white or crimson blotched leaves. This further
attests the close relationship between these plants. It seems that the
plant from western China deserves only to be recognized as a variety,
as it has a distinct geographical range. This differentiation is also in
line with another widespread species of. the genus, A. polygama, where
the western Chinese form is only slightly different from the plant in
northeastern Asia.

11. Actinidia polygama (Sieb. & Zucc.) Maxim. in Mém. Acad. Sci.
St. Pétersb. Sav. Etrang. 9: 64. 1859 (Prim. Fl. Amur.).

Climbing shrubs, to 5 m., the stem slender; branches glabrous or the
very young shoots sometimes lightly puberulous; pith large, white,
solid. Leaves membranaceous, sometimes the whole leaf white or
yellowish or the upper half variegated with a blotch of white or pale
yellow, ovate, 7-12.5 cm. long, 4.5-8.5 cm. wide, the apex long-acumi-
nate, the base acute to rounded to truncate, very rarely subcordatulate,
the margins finely serrulate, dark green and glabrous or rarely sparsely
setose above, slightly paler beneath and setose along the costa and the
main veins or glabrous, the costa and veins slender, inconspicuous
above, distinct and elevated beneath, the secondary veins 6 or 7 per
side, arcuate-ascending, strongly anastomosing, the veinlets reticulate,
inconspicuous above, subconspicuous beneath; petioles slender, 24.5
em. long, glabrous or sparsely setose. Flowers white, solitary or 2- or
3-fascicled, lateral, the pedicels slender, 6-10 mm. long, more or less
puberulous; sepals 5, ovate, sometimes unequal, about 7 mm. long and
4.5 mm. wide, acute at apex, more or less glabrous, the margins
finely ciliate, the base somewhat cuneate; petals 5, oblong-ovate, 12-13
mm. long, 5.5-10 mm. wide, the apex rounded to obtuse, the base
broad; stamens numerous, the filaments slender, filiform, 5-6 mm. long,
the anthers yellow or brown, 1.5-2 mm. long, the apex pointed, the
base sagittate; rudimentary ovary in staminate flowers very small,
globose; ovary bottle-shaped, 3.54.5 mm. long, about 2 mm. in diam-
eter, glabrous, the styles about 3 mm. long. Fruit globose to ovoid,
about 2.5 em. across, yellow, glabrous, not lenticellate, the apex rostrate.

Nakai separates some of the plants formerly referred to A. polygama
from Szechuan and Hupeh as a distinct species, A. lecomtei. The latter
has also the distinct solid white pith and “white or brownish crisped
hairs on the principal veins like Actinidia polygama, but lacks of rigid
glands. Its anthers are not yellow, but are purplish or purple.” The

1952] LI, REVIEW OF THE GENUS ACTINIDIA 21

specimens quoted by him are Farges 406, type, Farges 79 p. p., Henry
s.n., 1788, 6644 p. p., David s

Nakai at the same time areas A. polygama, of Manchuria, Korea,
and Japan, as also present in western China, citing Wilson 1363 and
1363a from Hupeh. These collections are also available for the present
study. Number 1363 agrees in all respects with the other collections
from western China, especially in the nearly glabrous leaves and the
brown anthers. Number 1363a, apparently from a younger shoot, has
leaves that are very sparsely setose on both surfaces, evidently the
so-called rigid glands of Nakai. The anthers are also brownish.

With a very large series of specimens from China proper, Manchuria,
Korea, and Japan, I am convinced that the widespread species A.
polygama, as currently accepted, can at most be differentiated into two
geographical varieties, one in the east, in Japan, Korea, and Manchuria,
and one in the west, in western China, but not as two species coexisting
in western China. The typical form in the east has yellow anthers and
leaves with scattered setose hairs on the veins on both surfaces. Ex-
tending from Korea inland to Manchuria, the plants become more and
more glabrate. The plant of western China, here considered as repre-
senting a separate variety, has nearly glabrous leaves only occasionally
setose on one or both surfaces or slightly hairy along the veins on the
lower surface only. The anthers are brownish in color, generally of a
darker shade than the typical form in the dried state. In all other
characters, the two varieties seem exactly identical

Actinidia polygama has been much confused in literature and her-
baria with A. kolomikta, but, as Rehder has noted, it is readily dis-
tinguished by the large solid white pith of its branches. In A. kolomikta,
as well as most other related species, the pith is lamellate and mostly
brownish in color. Actinidia kolomikta has also distinctly cordate
leaves, while in A. polygama the leaf-bases are rounded to subacute
but never cordate. Also in A. kolomikta the ovary is cylindric-ovoid
and not bottle-shaped as in A. polygama and other allied species.

Key TO THE VARIETIES

A. Leaves more or less sparsely setose along the veins beneath; anthers

yellow (Manchuria, Korea, Japan).............. lla. var. polygama.
AA. Leaves generally glabrous, rarely sparsely setose beneath; anthers brown
PWESteT RIGRY fo ceca ony OS ee a ie eee 1lb. var. lecomtet.

lla. Actinidia polygama (Sieb. & Zucc.) Maxim. var. polygama.

Actinidia ee (Sieb. & Zuee.) Maxim. in Mém. Acad. Sci. St. Pétersb.
Sav . 9: 64 1859 (Prim. Fl. Amur.); Miq. in Ann. Mus. Bot.
ore cs a 15. 1867 (Prol. Fl. Jap. 203); Dunn in Jour. Linn. Soe.
Bot. 39: 403. 1911; Nakai in Bot. Mag. Tokyo 47: 254. 1913, p. p.

Trochostigma polygama Sieb. & Zuce. in Abh. Phys.-Math. Cl. Akad. Wiss.
Miinch. 3: 727, t. 2. f. 2. 1843.

Trochostigma polnbilis Sieb. & Zuce. in loc. cit.

22 JOURNAL OF THE ARNOLD ARBORETUM [voL. XXXII

Actinidia volubilis (Sieb. & Zucc.) Planch. ex Miq. in Ann. Mus. Bot.
Lugd.-Bat. 3: 15, 1867 (Prol. Fl. Jap. 203).
Leaves glabrous or sometimes sparsely setose above, more or less
sparsely setose along the veins on the lower surface; anthers yellow.
Sakhalin, Manchuria, Korea, Japan. Flowers white.

s. n. (A, GH), £. H. Wilson s. n. (A); Mino prov., Shiota 3317 (A), 8038
(A); Hyogoken, K. Uno 19136 (A); Takao-san, Musashi prov., 2. H. Wilson
s.n. (A), P. H. Dorsett & W. J. Morse 794 (US); Mt. Mitsumune, Musashi
prov., No. coll. (US); Mt. Tanigawa, Niigata, S. Suzuki 398026 é. 5
Nanokawa, Tosa, No coll. (GH, US); Yoshino, Yamato, Tokyo Mus. 74
(US); Asama, G. Faurie 6127 (A). Kiusiu: Fukuoka, 7. Tanaka 206 (A).
Tsushima, C. Wilford s. n. (GH).

KOREA: Keijyo, prov Keiki, 2. H. Wilson 8752 (A, US), 10608 (A, US);
Oo-ryong-too, E. H. Wilson 8501 (A); Tanyudo, prov. Heian, F. H. Wilson
8675 (A); Konga-san, prov. Kogan, H. H. Wilson 10533 (A, US); Quelpaert,
T. Taquet 989 (A), 2681 (A), 2682 (A), 2684 (A), 2689 (A).

CHINA: Manchuria: Bay possiet, Mazimowicz s. n. (GH, NY, US);
Port Deans, Dundas, Mazimowicz s. n. (GH); Mao-erh-shan Station, B. V.
Skvortzov s.n. (A); Mifun Station, B. V. Skvortzov s. n. (A), P. H. Dorsett
£1767 (A, GH, NY, US).

Two species of Trochostigma described by Siebold & Zucearini from
Japan, 7’. polygama, and T. volubilis, have long been recognized as
representing the same species of Actinidia, the earlier name being A.
polygama adopted by Maximowicz (1859) and also by Miquel (1867).
Also see Nakai for his listing for various misapplied names based on
Japanese plants of this species.

Trochostigma repanda Sieb. & Zuce. was included in the synonymy
of A. polygama by Maximowicz (in Bull. Acad. Sci. St. Pétersb. 31: 19.
1887) together with T. polygama and T. volubilis. This was followed
by most subsequent authors and a combination in Actinidia was effected
by Honda as Actinidia repanda (Sieb. & Zuce.) Honda ex Koidz. in
Acta Phytotax. Geob. 9: 97, in clavi, 1940. However, the type of this
species, as indicated by A. C. Smith, represents a species of Schisandra,
actually the same as the well-known S. nigra Maxim. of Japan and
Korea, which is now properly known as Schisandra repanda (Sieb. &
Zuce.) A.C. Smith. See Sargentia 7: 143-146. 1947, for a full discussion.

11b. Actinidia polygama (Sieb. & Zucc.) Maxim. var. lecomtei
(Nakai) comb. nov.
Actinidia lecomtei Nakai in Bot. Mag. Tokyo 47: 253. 1933.
Actinidia melanandra sensu Finet & Gagnep. in Bull. Soc. Bot. France 52:
Mém. 4: 21. 1905, p. p. (Contr. Fl. As. Or.); Dunn in Jour. Linn. Soc.
Bot. 39: 403. 1911, p. p.; non Franch.

1952] LI, REVIEW OF THE GENUS ACTINIDIA 23

Actinidia polygama sensu Finet & Gagnep. l.c., p.p.; Rehder in Sarg. PI.
Wils. 2: 380. 1915; Nakai in Bot. Mag. Tokyo 47: 254. 1933, p. p.; Chun
in Sunyatsenia 1: 273. 1934; non Maxim.

Leaves glabrous or rarely sparsely setose above, generally glabrous

on the lower surface, rarely very sparsely setose; anthers dark brown.

Western China (western Hupeh, Szechuan, and southern Shensi), in
thickets at altitudes of 1350-2100 meters. Flowers white, June—July.

CHINA: Shensi: T’ai-pei-shan, W. Purdom 891 (A, US), G. Fenzel
878 (A). Hupeh: Without precise localities, BE. H. Wilson 5922 (GH),
5922A (GH), 5922B (GH), 7637 (GH); Fang Hsien, £. H. Wilson 2013 p. p.
(A); Hsing-shan Hsien, FE. H. Wilson 2013 p. p. (A, GH, US); Wu-tu-ho,
W. Y. Chun 3641 (A); “Gia-che-yuan,” W. ¥. Chun 4203 (A, US).
Szechuan: Wa-shan, FE. H. Wilson 934A p. p. (A, GH, US), 2010 (A, GH,
US); Pao-hsing Hsien (Muping), EF. H. Wilson 934A p. p. (A, US) K. L.
Chu 8760 (A), 3870 (A); South Wushan, A. Henry 5764 (A, GH), 5922 (US),
5922A (A), E. H. Wilson 1363 (A, NY, US), 1363A (A, NY, US); Ch’eng-k’ou
Hsien, R. P. Farges 406 (NY).

The citation of Chun in literature refers to the collection S. P. Ko
53081, from Kwangtung. I have not been able to consult this collection
and therefore am not sure as to the propriety of referring it to this
variety, as it is geographically distinct from the known range.

Some of the herbarium specimens from eastern China alleged to be
A. polygama are found to represent an entirely different and little-
known species, A. valvata Dunn.

12. Actinidia valvata Dunn in Jour. Linn. Soc. Bot. 39: 404. 1911.

Climbing shrubs to 13 m., the young branches glabrous to sometimes
grayish pubescent, pale; pith small, white, solid or slightly lamellate.
Leaves chartaceous, ovate to ovate-lanceolate, 6-10 cm. long, 3-5 em.
broad, acute to long-acuminate at apex, cordate to acute or rarely
truncate at base, sometimes unequally so, the margins minutely ger-
rulate, concolored or slightly paler beneath, glabrous on both surfaces
or sometimes slightly pubescent in the nerve-axils beneath, the costa
and veins inconspicuous to subconspicuous above, distinct and raised
beneath, the secondary veins 5 or 6 per side, arcuate-ascending, -an-
astomosing, inconspicuous above, conspicuous or subconspicuous_ be-
neath; petioles slender, 1-2 em. long, glabrous. Flowers 1-3-fascicled,
axillary; pedicels very slender, 1—-1.3 em. long, sparsely puberulous:
bracts linear, minute; sepals 2 or 3, concave, strictly valvate, more or
less unequal, ovate, 7-9 mm. long, 4-6 mm. wide, acute at apex,
glabrous or slightly puberulous without; petals 7 or 8, white, imbricate,
oblong, 1-1.2 cm. long, about 6 mm. wide, acute to rounded at apex,
attenuate at base; stamens many, the filaments slender, about 5 mm.
long, the anthers yellow; ovary bottle-shaped, glabrous. Fruit globose
or ovoid, about 2-2.5 em. across, yellow or orange, glabrous, not
lenticellate, with persistent calyx at base.

Eastern China (northern Kiangsi, southern Anhwei, southern Ki-

24 JOURNAL OF THE ARNOLD ARBORETUM [voL. XXXII

angsu, and Chekiang), in thickets at altitudes of 200-800 meters.
Flowers white, May.

Huang-shan, R. C. Ching 3054 (A, US); Ch’ung-yiin, A. N. Steward 7226
(A, US). Kiangsu: I-shing Hsien, Ching & Tso 551 (A), Y. L. Keng 2589
(A); Nanking, W. Y. Chun 2117 (A), 2139 (A). Chekiang: Ch’ang-hua
Hsien, F. N. Meyer 1532 (A, NY), Y. L. Keng 590 (A); Tien-mu-shan, H. H.
Hu 1659 (A), T. Tang & W. Y. Hsia 11? (A); TYien-tai-shan, C. Y. Chiao
14392 (A, US).

Dunn’s type is from Lu-shan, Kiangsi, Bullock 121, which has not
been seen by me. The description is clear and the species distinct, but
since its publication it has not been recorded by any other author.
Actually it appears to be a common plant of the coastal provinces of
eastern China. Specimens in the herbaria are mostly erroneously
identified as A. polygama, a species apparently not present in these
provinces. The species has indeed the general appearance of A. poly-
gama, and the two are genetically close, as evidenced by the presence
of the very distinct, solid white pith in the branches of both. The pith,
however, is much smaller in A. valvata and sometimes it is slightly
lamellate. Actinidia valvata, moreover, is readily distinguished from
A. polygama, and for that matter also from all other species of the
genus, in the unique form of the calyx, which is irregularly split into
2 or 3 concave sepals.

13. Actinidia tetrameraMaxim. in Acta Hort. Petrop. 11: 36. 1890;
Dunn in Jour. Linn. Soc. Bot. 39: 404. 1911; Rehder in Sarg. Pl.
Wils. 2: 381. 1915.

Clematoclethra giraldii Diels in Bot. Jahrb. 29: 472. 1900.
Actinidia rubricaulis sensu Dunn in Jour. Linn. Soc. Bot. 39: 407. 1911,
p. p., (quoad Wilson 4764, 3271), non Dunn in Kew Bull. 1906: 2. 1906.
High climbing shrubs to 13 m., the branches grayish to reddish brown,
glabrous, the very young branchlets sometimes puberulous; pith small,
lamellate, brown. Leaves chartaceous, sometimes variegated with
blotches of white or pink, narrowly oblong-ovate, about 5-10 em. long
and 2.5-4 em. broad, long-acuminate at apex, obliquely cuneate to
truncate or rarely subcordate at base, serrulate at margins, glabrous
on both surfaces to slightly setose on the costa beneath, with tufts of
white hairs in the axils of secondary veins beneath, the two surfaces
concolored, the costa and veins inconspicuous to subconspicuous above,
raised and subeconspicuous beneath, the secondary veins very slender,

6-8 per side, arcuately ascending, anastomosing, the veinlets reticulate,

subconspicuous to inconspicuous beneath; petioles 1.5-3 cm. long,

glabrous or nearly so. Inflorescences glabrous or nearly so, the flowers

2- or 3-fascicled or solitary; pedicels slender, 1-2 em. long; bracts

minute, linear; sepals 4, very rarely 5, ovate, 4-5 mm. long, 2-3 mm.

broad, acute to rounded at apex, the margins ciliate; petals 4, rarely 5,

oblong, about 7 mm. long and 5 mm. broad, rounded at apex; stamens

1952] LI, REVIEW OF THE GENUS ACTINIDIA 25

numerous, the filaments very slender, about 4 mm. long, the anthers
yellow, oblong, about 2 mm. long, rounded at both ends, broader but
not sagittate at base; ovary cylindric to slightly bottle-shaped, gla-
brous, the styles about 3 mm. long. Fruit ovoid, about 1.5-2 em. long,
0.7-1.5 cm. across, brown, glabrous, not lenticellate.

Western China (Kansu, Shensi, western Hupeh, Szechuan, and
Sikang), in thickets at altitudes of 1300-2700 meters. Flowers white,
sometimes tinged with pink, June.

CHINA: Kansu: Tien-shiu Hsien, F. Fenzel s. n. (A); Lower Tebbu
Country, J. F. Rock 14728 (A), 15026 (A). Shensi: T’ai-pei-shan, W.
Purdom 892 (A, US); “Kan-y-san,” J. Giraldi s.n. (A). Hupeh: Without
precise locality, H. H. Wilson 6821 (GH); Fang Hsien, E. H. Wilson 2011
(A, US), 4322 p. p. (A, US); Hsing-shan Hsien, FE. H. Wilson 2096 (A, US).
Sikang: K’ang-ting Hsien, C.Y. Chiao 1635 (A). Szechuan: Sung-p’an
Hsien, E. H. Wilson 4557 (A), Pao-hsing Hsien (Muping), Z. H. Wilson 809a
(A); Kuan Hsien, EZ H. Wilson 2006 (A), 4322 p. p. (A); Ch’eng-k’ou Hsien,
R. P. Farges 530 (NY, tsoryre); O-mei-shan, FE. H. Wilson 4764 (A), W. P
Fang 3063 (A), 6559 (A), 16625 (A), C. L. Sun 2166 (A), S. C. Sun & K.
Chang 292 (A)

This species is related to A. kolomikta but can be readily distin-
guished by its smaller narrower oblong-ovate leaves, bearded in the
axils of the secondary veins beneath but otherwise glabrous or rarely
setose on the costa, and also by the tetramerous flowers. Both species
have similar brown lamellate piths. The leaves of A. tetramera, like
those of A. kolomikta, are frequently variegated. ‘The flowers, especially
the pistillate ones, are occasionally 5-merous.

Clematoclethra giraldii Diels is given in the synonymy by Rehder.
A photograph of the holotype is in the herbarium of the Arnold
Arboretum.

Plants from O-mei-shan, Szechuan province, at the southernmost
limit of the species, have slightly smaller flowers, of generally darker
reddish color, and shorter peduncles, and they may prove to represent
a variety of the species when more material is made available.

14. Actinidia maloides sp. nov.

Frutex scandens, 5-6 m. altus, ramis teretibus longitudinaliter
suleatis, purpureo- snipteseentions vel cinnamomeis, glabris, ramulis
novellie leviter puberulis; medulla parva brunnea lamellata; foliis
chartaceis, interdum variegatis, oblongo-ovatis, circiter 5.5-8.5 cm
longis et 3-3.5 cm. latis, apice acuminatis, basi latis, plus minusve
cordatis interdum rotundatis, valde inaequalibus, margine minute ser-
rulatis, utrinque concoloribus glabris vel supra sparse setosis subtus
parcissime secus venas pubescentibus, costa supra subprominente
subtus prominente, nervis lateralibus utrinsecus 6-8, supra paulo
impressis subtus elevatis prominentibus, arcuato- adscendentibus, prope
marginem anastomosantibus, venis reticulatis subtus subprominentibus;
petiolis 1.5-2.5 cm. longis, glabris vel puberulis; inflorescentiis axil-

26 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXXIII

laribus puberulis, 2-4-, plerumque 3-floris, pedunculis gracilibus, ad
1.5 cm. longis, pedicellis ad 1 cm. longis, bracteis nullis; sepalis 5,
ovatis, purpureo-rubescentibus, glabris vel glabratis, circiter 5 mm
longis et 3 mm. latis, acutis, margine minute ciliatis; petalis 5, obovatis,
circiter 8 mm. longis et 7 mm. latis, rotundatis, pallide roseis, ad
apicem rubescentibus; staminibus numerosis, filamentis gracilibus, cir-
citer 3-4 mm. longis, antheris flavis oblongis circiter 2.5 mm. longis,
rotundatis, basi sagittatis; ovario rudimentario minute globoso, glabro,
stylis numerosis; fructibus brunneis oblongis ad 2 cm. longis et 1.2 cm.
diametro, elabris, elenticellatis; seminibus ovoideis compressis, circiter
2 mm. longis et 1.5 mm. crassis, testa in sicco minute reticulata.

Western China (Sikang), on mountain slopes among thickets at an
altitude of about 2000 meters. Flowers pink, May—June.

CHINA: Sikang: ree -sui Hsien, T. T. Yui 927 (A, TYPE); Han-yuan
Hsien, HL. H. Wilson 854 (A, fruit).

This species is of the ae of A. kolomikta. It has similarly
variegated leaves characteristic of some of the species of this general
croup. It is apparently most closely related to A. tetramera Maxim.,
but can be distinguished by the consistently 5-merous, pink flowers,
with purplish red calyx and sagittate anthers, and the broad leaves
which usually have broad, cordate bases.

The pink petals are red toward the upper margins, like the flowering
crab apple. In flower this clearly is the most showy species of the genus.

15. Actinidia kwangsiensis sp. nov.

Frutex scandens, circiter 3 m. altus, ramulis oppositis, atrobrunneis,
lenticellis parvis pallide dispersis, glabris, ramulis novellis plus minusve
ferrugineo- -pubescentibus; medulla .brunnea lamellata; foliis sub-
coriaceis, ovatis vel oblongo-ovatis, 8-9.5 em. longis, 4-5 em. latis,
apice longe caudato-acuminatis, basi oblique rotundatis, margine
adpresse minute serrulatis, supra atro- viridibus glabris, subtus pallidis,
secus venas leviter ferrugineo gr anuloso til costa supra
prominente, subtus elevata, nervis lateralibus utrinsecus 5-7, arcuato-
adscendentibus, prope marginem anastomosantibus, venulis minute
reticulatis, subtus subprominentibus; petiolis 2-8 cm. longis, plus
minusve ferrugineo-glanduloso-pubescentibus; inflorescentiis sub an-
thesi ignotis; ovario distincte ampullifero, glabro, stylis circiter 2.5 mm.
longis; inflorescentiis post anthesin axillaribus, 1- vel 2- oes plus
minusve ferrugineo-granuloso-] pedunculis 5-7 mm.
longis; pedicellis ad 1.8 em. longis; fructibus immaturis Viridibus,
oblongis, circiter 2 cm. longis et 1 cm. crassis, glabris, apice valde
rostratis.

Southern China, in Kwangsi, in open thickets, at an altitude of 1000
meters. Flowers unknown. |

CHINA: Kwangsi: Yin-tung, Min-shan, N. of Lu-chen, near the
Kweichow border, R. C. Ching 6185 (NY, TYPE).

1952] LI, REVIEW OF THE GENUS ACTINIDIA a7

This species is admittedly established on rather inadequate material,
but it seems clearly to represent an undescribed species. Its glabrous
bottle-shaped ovary indicates distinctly and definitely its alliance witla
either the dark-flowered A. purpurea Rehd. or with the pale flowered
A. melanandra Franch. It can be distinguished from the former by the
smaller, narrower leaves and the brown pith, and from the latter by its
leaves being not glaucous beneath. It is also distinct in having opposite
branches, although the material at hand is not sufficient to indicate
whether or not this is constant.

There is also the possibility that this may be the same as A.
longicauda F. Chun, here classified as an imperfectly known species.
However, as Chun’s species is based on a specimen with staminate
flowers only, while the present species is based on a young fruiting
specimen, it is impossible critically to compare the two.

16. Actinidia melanandra Franch. in Jour. de Bot. 8: 278. 1894;
Dunn in Jour. Linn. Soe. Bot. 39: 402. 1911, p. p.; Rehder in Sarg.
Pl. Wils. 2: 378. 1915; Hand.-Mazz. Symb. Sin. 7: 390. 1931;
Nakai in Bot. Mag. Tokyo 47: 252. 1933.

Actinidia rufa var. parvifolia Dunn in op. eit. 403. 1911.

High climbing shrubs to 7 m.; branches reddish, the very young
shoots sometimes puberulous and slightly glaucous; pith lamellate,
more or less whitish. Leaves chartaceous, elliptic or ovate to oblong-
ovate or oblong-lanceolate, about 6-9.5 em. long and 2.5-4 em. broad,
acuminate at apex, cuneate to rounded or.truncate at base, sometimes
unequal, the margins minutely serrulate, the teeth ascending, glabrous
on both surfaces except for small tufts of rusty hairs in the axils of
the secondary nerves on the lower surface, green above, glaucous
beneath, the costa and veins slender, subconspicuous above, more or
less raised and distinct beneath, the secondary nerves about 4-6 per
side, arcuately ascending, anastomosing, the veinlets reticulate, incon-
spicuous to subconspicuous on both surfaces ; petioles slender, 2.5-3
em. long, glabrous or slightly puberulous. Inflorescence in 3- 5-flowered
cymes or the flowers solitary, the peduncles and pedicels slightly
puberulous, slender; pedicels to 1.8 em. long; bracts linear, minute.
Flowers white; sepals 5, occasionally 4, ovate, 6-7 mm. long, 3-4 mm.
broad, acute at apex, glabrous, the margins sometimes ciliate, the base
more or less connate; petals 5, rarely 4, white, sometimes brownish
toward the base, oblong, 1.1-1.3 em. long, 4-7 mm. broad, rounded at
apex, gradually narrowed at base; stamens numerous, the filaments
slender, about 3 mm. long, the anthers blackish when dry, slender,
about 2 mm. long, acute and pointed at apex and sagittate at base :
ovary bottle-shaped, glabrous, 6-7 mm. long, 2-3 mm. in diameter;
styles 3.4 mm. long. Fruit ellipsoid to ovoid, glabrous, smooth, about
3 cm. long and 2.5 cm. across, not lenticellate.

CHINA: Hupeh: Western Hupeh, HZ. H. Wilson 1068 (A, NY, US)
1068a (

I

A, NY, US), 5988a (GH); Fang Hsien, E. H. Wilson 4459 (A).

28 JOURNAL OF THE ARNOLD ARBORETUM [voL. XXXIII

Szechuan: Kuan Hsien, W. P. Fang 2364 (A, NY); Ch’eng-k’ou Hsien,
R. P. Farges 79 (NY).

e Dunn describes very briefly A. rufa var. parvifolia, basing it on
Henry 5938a, 6644, and 6794, all from Hupeh. This is reduced to A.
melanandra by Rehder. This species is readily distinguished from other
species of this general region by the glaucous under surfaces of the
leaves, which are glabrous except for the tufts of brownish hairs in the
axils of the secondary veins on the lower surface. I cannot verify
Handel-Mazzetti’s record of this species in northeastern Yunnan, based
on Teng 260. In the past A. melanandra has often been erroneously
attributed by various authors to species of other regions, but it seems
that the range of the present species is limited to western China (in
western Hupeh and Szechuan, and possibly also in northern Yunnan).

17. Actinidia hypoleuca Nakai in Bot. Mag. Tokyo 38: 312. 1904,
47: 256. 1933.
Actinidia melanandra sensu Finet & Gagnep. in Bull. Soc. Bot. France 52,
Mém. 4: 21. 1905, p. p. (quoted Japanese plants), non Franch.

Actinidia japonica Nakai in Bot. Mag. Tokyo 28: 311. 1914.

Climbing shrubs; branches grayish, the young branchlets much
darker, glabrous, without lenticels or with small inconspicuous lenticels
on older branches only; pith brown, lamellate, sometimes nearly solid.
Leaves subchartaceous, ovate to ovate-oblong, about 2.5-7 cm. long,
2.5-4.5 em. broad, abruptly acuminate at apex, broadly acute to
rounded or truncate at base, the margins finely serrulate, glabrous and
dark above, glaucous and glabrous beneath except with tufts of brown
pubescence in the nerve-axils, the costa and veins inconspicuous above,
dark above, glaucous and glabrous beneath except with tufts of brown
veins about 4 or 5 per side, straight or slightly arcuate-ascending,
anastomosing, the veinlets finely reticulate, inconspicuous above, sub-
conspicuous beneath; petioles 2-3.5 cm. long, dark purple, glabrous.
Inflorescences in small axillary cymes of 1-5 or more flowers, glabrous
or slightly pubescent. the peduncles slender, to 5 mm. long, the pedicels
slender, to 10 mm. long. Flowers white; sepals 5, ovate-oblong, to
6 mm. long and 3 mm. broad, glabrous or puberulous, acute to obtuse
at apex, the margins often ciliate; petals 5, obovate, to 12 mm. long
and 7 mm. broad, rounded at apex; stamens numerous, the filaments
slender, about 3 mm. long, the anthers purple, oblong, about 2 mm.
long, rounded at apex, slightly sagittate at base; ovary bottle-shaped,
glabrous, about 5 mm. long and 2 mm. across, the styles 3-4 mm. long.
Fruit ovoid, about 1.5 em. across, glabrous, not lenticellate, slightly
rostrate at apex.

Southern and central Japan. Flowers white.

JAPAN: Hondo: Satsuma prov., G. Masamune s. n. (NY); Mino prov.,
K. Shiota 4972 (A), 5722 (A), 6341 (A), 9699 (A); Kai prov., Tasko, K
Sakurai s. n. (A); Nonokawa, Tosa, K. Watanabe s. n. (GH). Sikoku:
Nagasaki, R. Oldham 95 (GH).

1952] LI, REVIEW OF THE GENUS ACTINIDIA 29

This species differs from the closely allied A. arguta in the generally
smaller leaves, which are usually glaucous beneath. Among the her-
barium specimens, various degrees appear in the shade of leaf color,
ranging from green to glaucous in leaves even from the same specimen.
Nakai speaks of A. arguta as “easily discriminated from A. hypoleuca
in the field by its green foliages.”

The name A. japonica was first given by Nakai in Bot. Mag. Tokyo
27: (163). 1913, in his key to Japanese species of Actinidia, and later a
description was given in 1915. This species name is given as a synonym
of A. hypoleuca Nakai (mistakenly as “A. hypoglauca Nakai”) in
Koidzumi’s key (in Acta Phytotax. Geob. 9: 97. 1940). The original de-

scription, though brief, is sufficient to prove that this reduction is
justified. It is strange to note, however, that this name is never men-
tioned by Nakai himself in his subsequent discussions of this genus.

18. Actinidia purpurea Rehder in Sarg. Pl. Wils. 2: 378. 1915, in
Jour. Arnold Arb. 15: 96. 1934; Hand.-Mazz. Symb. Sin. 7: 390.
1931; Nakai in Bot. Mag. Tokyo 47: 253. 1933.

Actinidia melanandra Franch. var. latifolia E. Pritz. ex Diels in Bot. Jarhb.
29: 470. 1900. Syn. nov.

Actinidia rufa var. arguta Dunn a Jour. Linn. Soc. Bot. 39: 402. 1911, p. p.
quoted Henry 11008). Syn. no

ee ae var. typica Bina in loc. cit., p. p. (quoted Henry 5622).
Syn. n

Actinidia ee Hu in Bull. Fan Mem. Inst. Biol. Bot. 10: 128. 1940.
Syn. nov

High climbing shrubs to 20 m.; branches glabrous or the very young
branchlets puberulous, rarely tomentose, grayish brown; pith white,
lamellate. Leaves thickly chartaceous, elliptic to elliptic-ovate or
broad-ovate, 8-12 cm. long, 4.5-6.5 em. broad, acute to acuminate at
apex, acute to rounded or truncate at base, usually oblique, the mar-
gins serrulate, with appressed teeth, glabrous and dull above, nearly
concolored or slightly paler beneath, glabrous or sparsely setose or
tomentose along the costa and veins on the lower surface with or with-
out tufts of whitish or yellowish hairs in the nerve-axils, the costa and
nerves slender, subconspicuous above, distinct and elevated beneath,
the secondary veins about 5 or 6 per side, arcuate-ascending, anas-
tomosing, the veinlets reticulate, inconspicuous above, subconspicuous
beneath; petioles 3-5 cm. long, glabrous or rarely tomentulose. In-
florescences in axillary cymes, puberulous, the staminate cymes often
many-flowered, the pistillate usually 3-flowered. Flowers white; sepals
5, ovate, 4-7 mm. long, more or less unequal, obtuse at apex, often
turning blackish on drying, glabrous or rarely puberulous, the margins
ciliate; petals 5, ovate to oblong-ovate, often unequal, 7-12 mm. long,
4-7 mm. broad; stamens numerous, the filaments slender, 3-4 mm.
long, the anthers oblong, blackish, 2 mm. long, the base divergent;
ovary bottle-shaped, glabrous, about 6 mm. long and 2 mm. across,

30 JOURNAL OF THE ARNOLD ARBORETUM (VoL. XXXIII

the styles about 4 mm. long. Fruit _ or oblong, about 2-2.5 cm.
long, purplish, glabrous, the apex rostra

CHINA: Hupeh: Without precise ai E. H. Wilson 1029a (A),
5622 (G); Nan-t’o, HE. H. Wilson 1165 (A, NY, US). Hunan: Heng-shan,
H. Handel-Mazzetti 12212 (A). Szechuan: Without precise locality, F. H.
Wilson 3269 (A); Pao-hsing Hsien (Mu-ping), F. H. Wilson 1314 p. p. (A,

S, GH, rype:), 7314a (A); Mo-tien-ling, F. T. Wang 22445 (A), 22478 (A);
O-mei-shan, Y. 8. Liu 1230 (A), C. Y. Chiao & C. S. Fan 286 (A), 886 (A),
H.C. Chow 7645 (A), 12179 (A). Sikang: Kan-ting Hsien (Tachienlu),
E. H. Wilson 1314 p. p. (A), C. Y. Chiao 2029 (A), 2040 (A). ane nnan:
Without precise localities, FP. Ducloux 465 eas G. Forrest 14845 (A), 16223
(A), H. T. Tsai 57168 (A), 57569 (A), 5 325 (A); Yangtze- ie divide,
H. Handel-Mazzetti 7873 (A); Mekong-s . divide, G. Forrest 19483 (A,
US), J. F. Rock 22720 (A, NY); Weichat. Hsien, H. T. Tsar 57911 (A), C. W.
Wang 63608 (A), 63701 (A), 64046 (A), 68660 (A), 70423 (A); Chung-tien,
K. M. Feng 8341 (A); south of Red River, Man-mei, A. Henry 9694
(A, NY); Meng-tzu, A. Henry 11008 (A, NY, JS); Lan-p’ing Hsien, H. T.
Tsai 54000 (A); P'ng- -pien Hsien, H. T. Tsai Mee (A); Lan-ts’ang Hsien,
C.W. Wang 76627 (A). Kweichow: San-ho Hsien, Y. Tsiang 6435 (NY):
Tu-shan Hsien, Y. T'siang 6761 (NY)

This species is the counterpart of A. arguta in southwestern China,
and it differs from the latter in the relatively longer, narrower leaves
that are never setose and with appressed serrations close to the mar-
gins, the generally smaller flowers, and the long, dark, purple-colored
fruits. It was originally included in the concept of A. arguta Dunn,
who cited specimens such as Henry 5622 and 11008 which were later
designated as types of Rehder’s A. purpurea. Wilson 1512 from Kiangsi,
a sterile specimen with setose hairs, evidently does not belong here as
originally cited by Rehder but should be referred to A. arguta.

A photograph of the type of A. melanandra Franch. var. latifolia
Pritzel is also in the herbarium of the Arnold Arboretum. It is from
Nan-ch’iian, in Szechuan province, collected by V. Rosthorn in 1891.
The original description is very brief, referring only to the size of the
leaves, “foliis latis 8-9 & 6-7.5 cm.”, but the photograph clearly shows
that not A. melanandra Franch. but A. purpurea Rehder is represented.

Actinidia chartacea Hu, another name to be reduced, is from north-
western Szechuan, ‘Mo-Tien-Ling . . F. T. Wang, no. 2245a (type),
Aug. 31, 1930.” Hu compares his plant with A. kolomikta Maxim. and
also with A. latifolia Merr., the latter being a widely different plant
with stellate tomentum and ‘spotted fruit. Hu’s type is not now avail-
able, but two collections made by the same collector from the type
locality at the same time when the type was collected, F. T. Wang
22445 and 22478, clearly represent the same plant. They are unques-
tionably A. purpurea. As Hu’s description also fits the latter species
perfectly, it is believed desirable also to reduce A. chartacea to syn-
nonymy.

The species A. purpurea, as mentioned above, is very close to A.
arguta, and it remains to be seen whether it will eventually be advisable

1952] LI, REVIEW OF THE GENUS ACTINIDIA 31

to revert to the broader concept of Dunn and treat this as a variety of
A. arguta.

19. Actinidia arguta (Sieb. & Zucc.) Planchon ex Miq. in Ann. Mus.
Bot. Lugd.-Bat. 3: 15. 1867.

Climbing shrubs to 7 m.; branches glabrous or the very young
branchlets puberulous, rarely tomentose, grayish brown, the lenticels
absent on younger branches, small and inconspicuous on mature ones;
pith white to brown, lamellate. Leaves membranaceous to chartace-
ous, elliptic-ovate to broadly ovate, 8-12 em. long, 4.5-7.5 em. broad,
abruptly acuminate at apex, rounded to subcordate at base, rarely
cuneate, usually oblique, the margins sharply serrate, glabrous and
dull above, nearly concolored or slightly paler beneath, glabrous to
rusty-tomentose to setose on one or both surfaces, especially beneath,
with or without tufts of whitish or yellowish hairs in the axils of the
secondary veins beneath, the costa and veins slender, subconspicuous
above, distinct and raised beneath, the secondary veins about 5 or 6 per
side, arcuate-ascending, anastomosing, the veinlets reticulate, incon-
spicuous above, subconspicuous beneath; petioles 3.5-8 em. long, gla-
brous or rusty-tomentose, sometimes setose. Flowers white, in axillary
cymes, puberulous, the staminate cymes often many-flowered, the pistil-
late with 1-3 or more flowers; sepals 5, ovate, 5-7 mm. long, obtuse at
apex, the margins ciliate, glabrous or rarely puberulous without; petals
white; brownish at base, ovate to oblong-ovate, often unequal, 7-12 mm.
long, 4-7 mm. broad; stamens numerous, the filaments slender, 3-4 mm.
long, the anthers blackish when dry, oblong, 2 mm. long sagittate at
base; ovary bottle-shaped, about 6 mm. long and 2 mm. across, glab-
rous, the styles about 4 mm. long. Fruit ovoid or oblong, about 2-2.5
em. long, greenish yellow, glabrous, not lenticellate, the apex rostrate.

Actinidia arguta was broadly defined by Dunn to include forms
ranging from Yunnan to Manchuria and Japan. Subsequently the
species A. purpurea was segregated by Rehder and the species A.
platyphylla A. Gray reinstated by Japanese botanists. The present
study, based on a large series of specimens from all these localities,
suggests that it is advisable to return to the somewhat broad concept
of Dunn. The several varieties here recognized occur in more or less
contiguous areas and pass into each other imperceptibly by inter-
grading forms. Clearly this is another of the very widespread, more
or less polymorphic species characteristic of the genus Actinidia. As a
species in its broad sense, it probably should include also A. purpurea
and A. hypoleuca; it is characterized by the long-petioled, more or less
membranaceous leaves, generally glabrous except for the veins and the
nerve-axils, the petals frequently purplish at base, the purple sagittate
anthers, the glabrous bottle-shaped ovary, and the smooth short-
rostrate fruits.

32 JOURNAL OF THE ARNOLD ARBORETUM [VoL. XXXII

Key To THE VARIETIES
A. Leaves membranaceous to chartaceous, rounded to subcordate at base,
glabrous or setose along the costa benea
B. Leaves membranaceous to subchartaceous; young shoots and leaves
as well as inflorescence glabrate except along the veins of the lower

surfaces of the leaves............... 0.2.0 -.0..0 005. a. var. arguta.

BB. Leaves subchartaceous; young shoots and leaves as well as inflores-
cence usually rusty-tomentose. ................. b. var. rufa.

AA. Leaves chartaceous, relatively shorter, broader, cordate at base, more
setose on the veins beneath....................... c. var. cordifolia.

19a. Actinidia arguta var. arguta.
ee das (Sieb. & Zuce.) Planchon ex Miq. in Ann. Mus. Bot. Lugd.-
Bat 1867; Nakai in Bot. Mag. Tokyo 47: 256. 1933.
oo. arguta Sieb. & Zuce. in Abh. Akad. Wiss. Miinchen 3: 727.
1843.

Actinidia rufa var. arguta Dunn in Jour. Linn. Soc. Bot. 39: 402. 1911.

Actinidia giraldii Diels in Bot. Jahrb. 36: Beibl. 82: 75. 1905; Dunn in Jour.
Linn. Soc. Bot. 39: 403. 1911. Syn. nov.

Actinidia megalocarpa Nakai ex Nakai & ae lg in oe Ist Sei. Exp.
Manch. IV. 1: 9, t. 3. 1933 (Pl. Nov. Jehol.) Syn

Leaves membranaceous to subchartaceous, Nig seaneate to broad-
ovate, 8-12 cm. long, 4.5-7.5 cm. broad, abruptly acuminate at apex,
rounded to subrounded at base, rarely cuneate, usually oblique, glab-
rous or sparsely setose along the costa and veins on both surfaces,
especially beneath, with or without tufts of whitish or yellowish hairs
in the nerve-axils beneath; inflorescences puberulous.

Eastern Siberia, Manchuria, and northern China to Korea and Japan,
in thickets at altitudes of 100-2000 meters. Flowers white, the anthers
purple, June.

CHINA: Manchuria: Hsiao-ling, P. H. & J. H. Dorsett 29 (US), P. H.
Dorsett 4086 (A, NY); Sui-fen-ho, B. V. Skvortzov s. n. (A); Kao-ling-tzu,
B. V. Skvortzov s. n. (A); Mao-erh-shan Station, B. V. Shvortzov s. n. (A);
Mifun Station, B. V. Skvortzov s. n. (A); Port Bruce, Maximowicz s. n.
(GH). Chahar: Yang-chia-p’ing, C. W. Wang 61824 (A); “Pao-feng-ssu,’
C.W. Wang 60813 (A), 60816 (A). Hopei: Without precise locality, C. F.
Li 11167 (NY); Ming-ling, K. M. Liou 618 (NY). Shansi: branch shih
Hsien, T. Tang 904 (NY); Chieh-hsiu Hsien, H. Smith 5444 (A). Shensi:
Kuan-yin-shan, J. Giraldi s.n. (A). Honan: Sung Hsien, J. waa 501 (A),
&67 (A), 1265 (A); Lu-shih Hsien, /. Hers 867 (A), 1143 (A), 1180 (A).
Shantung: Lao-shan, C. Y. Chiao oo re NY, US). Kiangsu :
Tung-hai Hsien (Haichow), J. Hers 645 (A). Anhwei: Wu-yuan Hsien,
R.C. Ching 3254 (A); prea shan, R. C. Ching 3046 (A). Chekiang:
T’ien-mu-shan, H. H. Hu 3 (A).

KOREA: Pyongyang. i R.K. Smith s.n. (US); Taiyudo, prov. Heian,

E. H. Wilson 8642 (A, US), Kongo-san, prov. Kogan, R. K. Beattie 10454
mn US), EF. H. Wilson 10718 (A); Mt. Aiensan, U. Faurie 496 (A).

JAPAN: Hokkaido: Sapporo, S. Takenobu s.n. (GH), S. Armoto s. n.
(GH), Y. Tokubuchi s. n. (GH), FB. Tokubuchi s. n. (GH), C. Wilford s. n.
(GH), Mazimowicz s. n. (GH, US); Abashiri, #. H. Wilson s. n. (A).

1952] LI, REVIEW OF THE GENUS ACTINIDIA 33

Hondo: Yoshino (Yamoto), Tokyo Mus. 76 (US); Yokohama, Mazximowicz
s. n. (US); Hakone, Sengoku, T. Sawada 2218 (A); Mino prov., K. Shiota
7712 (A), 7781 (A); Lake Chuzenji, J. G. Jack s. n. (A), EZ. H. Wilson s. n.
(A), C. S. Sargent s.n. (A); Mt. Tamigawa, Niigata, S. Suzuki 993024 (A).
Shikoku: lyo, Y. Ikkaku 9527 (A).

The specific names A. arguta and A. rufa are effected by Miquel who
credits them to Planchon. They are based on Trochostigma arguta
Sieb. & Zuce. and T. rufa Sieb. & Zucc. respectively. Dunn considers
them as separate varieties of the same species, A. rufa var. typica
and A. rufa var. arguta. Nakai maintains them as two distinct species.
Maximowicz in 1886 was the first to unite the two species, and he
took up the name A. arguta for the aggregate. This view is followed
by most subsequent authors. As noted by Rehder, A. arguta, the name
chosen by the first author to unite the concepts, must be adopted.

Nakai considers as A. arguta the common species in Japan which is
found nearly everywhere, while, A. rufa, considered by him as distinct,
is limited only to the extreme south in Kiusiu as well as the Liukius.
As the two are differentiated by him only in the degree of indument,
they seem hardly to merit specific recognition. Among the available
specimens, it is hard to find constant differences. Thus these two are
retained as varieties as given earlier by Dunn, but the range of A.
arguta var. rufa is found not to be strictly limited to the south as indi-
cated by Nakai. A specimen of Oldham’s collection, probably from
Korea, which bears Nakai’s identification as “A. rufa” clearly belongs
to this same form. Rehder and others treat A. rufa and A. arguta as
exact synonyms.

A photograph of the holotype of A. giraldii Diels, Giraldi 4065, from
southern Shensi, is in the herbarium of the Arnold Arboretum. It
matches exactly a specimen in the same herbarium collected by Giraldi
in 1897 from the same locality and determined by Rehder as A. pur-
purea. The setose leaves as shown by the specimen and described by
Diels, clearly indicate that A. giraldii is referable to A. arguta rather
than A. purpurea.

Nakai’s A. megalocarpa from Jehol, Chang-Shan-Yu, based on “N.
H. K. Sept. 13, 1933,” differs, according to the author, from A. arguta
in the thinner leaves and larger fruits. The latter measure 2 * 4 cm.
Basing one’s opinion on his description and illustration, it seems quite
safe to refer this name to A. arguta.

Among the above cited specimens is a collection made by C. be
Chiao, no. 2856, from Lau Shan, Shantung. This was originally de-
termined as A. polygama, but the lamellate instead of solid pith clearly
eliminates it from that species. Four specimens of this number are
observed, all of them sterile. These specimens, so far as the vegetative
characters can show, belong to A. arguta. I suspect that the record of
A. polygama from Lau Shan, Shantung, as given by Gilg & Loesener
in Bot. Jahrb. 39: Beibl. 75: 52. 1904, based on a Zimmerman collection,
should actually be referred to A. arguta.

34 JOURNAL OF THE ARNOLD ARBORETUM (VOL. XXXII

The pith of A. arguta is sometimes solid and at other times, especially
at the very central part, it becomes compactly lamellate, particularly
in older stems. When the pith is solid, it is small, of very firm texture,
and slightly pinkish colored like the wood; thus it is very different from
the solid, pure white, very large, and spongy pith of A. polygama.

19b. Actinidia arguta var. rufa (Sieb. & Zucc.) Maximowicz in Bull.
Acad. Imp. Sci. St. Pétersb. 31: 1886, in Mel. Biol. 12: 424. 1886.
Trochostigma rufa Sieb. & Zuce. in Abh. Akad. Wiss. Miinchen 3: 727, t. //,
f. d8-d13. 18438.
Actinidia rufa Planchon ex Miq. in Ann. Mus. Bot. Lugd. ert 3: 15. 1876
(Pro. Fl. Jap. 203); Nakai in Bot. Mag. Tokyo 47: 257. 1933.
Actinidia callosa var. rufa Makino in Bot. Mag. Tokyo 16: eee 1901.
Actinidia rufa var. typica Dunn in Jour. Linn. Soc. Bot. 39: 402. 1911.
Leaves subchartaceous, ovate to broad-ovate, 6-10 em. long, 5-10
em. broad, cordate to truncate at base, glabrous except a the nerve-
axils on the under surfaces; inflorescence mostly tomentos
Liukiu, Korea, and Japan. Flowers white, anthers a i June.
JAPAN: Hondo: Kai prov., fe Sasaro, No. coll. (US).
KOREA: “Korea arch ?, Pt. Hamilton,” Oldham 94 (GH).
LIUKIU: Takanosima & Onsima, C. Wright 31 (GH).

19c. Actinidia arguta var. cordifolia (Miq.) Bean, Trees & Shrubs
Brit. Isl. 1: 162. 1914.
Actinidia cordifolia Miq. in Ann. Mus. Bot. Lugd.-Bat. 3: 15. 1876. (Prol.
Fl. Jap

Actinidia platyphylla A. Gray ex Miq. |. ¢.; Nakai in Bot. Mag. Tokyo 47:
258. 1933.

Actinidia rufa var. cordifolia Dunn in Jour, Linn, Soc. Bot. 39: 403. 1911.
Actinidia rufa var. dulcisisma Koidzumi in Bot. Mag. Tokyo 44: 100. 1930.
Syn. nov.

Leaves subchartaceous, broad-ovate, about 4-9 em. long. 5-10 em.
broad, distinctly cordate at base, the costa and veins more setose be-
neath; inflorescence rusty-tomentose.

Japan and Korea, in thickets. Flowers white, anthers purple, June.

JAPAN: Hokkaido: Cape Sangar, J. Small s. n. (GH, Type of A.
platyphylla Gray); between Shojiko & Kofu, Dorsett & Morse 560 (US);
Nugata pref., Mt. Tanigawa, S. Suzuki 398025 (A). Hondo: Misaka Pass,
EF. H. Wilson s. n. (A); Nanokawa, Tosa, K. Watanabe s. n. (GH); Kai,
Gunnai, K. Sakurai s. n. (A)

KOREA: Oo-ryong-too, EF H. Wilson 8542 (A).

There is some confusion with regard to the synonymy of this variety.
Miquel’s A. cordifolia is based on “Unicum exemplar legit Pierot in
sylva ad upain fluvil Asija Gawa prope Kojanosa ins. Kiusiu.’”’? When
Dunn proposed the variety A. rufa var. cordifolia, it was clearly based
on Miquel’s A. cordifolia as the latter is cited in the synonymy. Among
the two specimens he cited is “Cape Sangar, Wright.” This is evidently
the type collection of A. platyphylla A. Gray, quoted by Miquel as a

1952] LI, REVIEW OF THE GENUS ACTINIDIA 35

number of J. Small. The holotype is in the Gray Herbarium, a J.
Small specimen in C. Wright’s collection. Dunn was apparently un-
aware of this fact, as he cited, “fide Bretschneider,” A. platyphylla
A. Gray in the synonymy of A. kolomikta Maxim. This was followed
by Rehder, in Sarg. Pl. Wils. 2: 381. 1915. Nakai has had access to
the type of A. platyphylla A. Gray, as he cited the Wright number
from both Gray and Paris, and relegated A. rufa var. cordifolia Dunn
to the synonymy of A. platyphylla A. Gray. However, the species A.
cordifolia Miq., the basis of Dunn’s variety, was cited by Nakai under
A. arguta Planch. ex Miq. He gives no reason for doing so, and appar-
ently he has not had access to the type of A. cordifolia Miq., a Pierot
number from Kiusiu. In Rehder’s Bibliography (Bibl. Cult. Trees &
Shrubs 459. 1949) both A. cordifolia Miq. and A. platyphylla A. Gray
are listed in the synonymy of A. arguta var. cordifolia (Miq.) Bean.

Koidzumi, in Bot. Mag. Tokyo 44: 100. 1930, considers A. arguta
var. cordifolia Dunn as distinct from A. cordifolia Miq. Actinidia arguta
is recognized by him as a synonym of A. platyphylla A. Gray, of which
he cites no specimen. Actinidia arguta var. cordifolia Dunn is renamed
A. rufa var. dulcissima Koidz., of which no description is given but
for which two collections, Pierot 445 and C. Wright s. n., are cited.
The Pierot collection from Kiusiu is most probably that on which
Dunn based his species. No reason is given by Koidzumi for renaming
Dunn’s variety. As I consider that A. rufa var. cordifolia Dunn, A.
cordifolia Miq., and A. platyphylla A. Gray are all synonymous with '
A. arguta var. cordifolia, Koidzumi’s varietal name is thus also in-
cluded. That this name is superfluous may have been later realized
by Koidzumi himself, as in his key to the Japanese species of Actinidia
(in Acta Phytotax. Geob. 9: 97-99. 1940) A. cordifolia Miq. is again
recognized by him, while his own A. rufa var. dulcissima is not men-
tioned at all.

20. Actinidia rubricaulis Dunn in Kew Bull. 1906: 2. 1906, in Jour.
Linn. Soc. Bot. 39: 407. 1911 p. p. (excluding Wilson 3271 & 4764).
Large climbing shrubs; branches reddish brown to purple, lenticellate,
elabrous to minutely hirsute when young; pith solid, whitish, firm.
Leaves chartaceous, oblong-lanceolate to ovate-oblong, about 8-10.5
em. long and 1.2-3.8 cm. broad, acuminate at apex, cordate to sub-
rounded at base, generally oblique, the margins laxly serulate, the teeth
sometimes glandular, glabrous on both surfaces, the upper surface dark,
the lower much paler, the costa and veins inconspicuous above, distinct
and raised beneath, the secondary veins 6 or 7 per side, distinctly
arcuate-ascending, anastomosing, the veinlets reticulate, inconspicuous
above, subconspicuous beneath; petioles 1-2.5 em. long, glabrous or
minutely hirtellous. Flowers usually scattered on short lateral branches,
the peduncles solitary, the lower ones lateral, the upper axillary or the
flowers in axillary fascicles of up to 5; pedicels or peduncles glabrous,
to 1.3 em. long, with or without a minute linear bract at the middle;

36 JOURNAL OF THE ARNOLD ARBORETUM [vVoL. XX XIII

sepals 5, mostly unequal, oblong, to 4.5 mm. long and 3 mm. broad,
obtuse to rounded at apex, the margins usually ciliate; petals 5, often
unequal, oblong-lanceolate, to 7.5 mm. long and 4.5 mm. broad,
rounded at apex; stamens numerous, the filaments slender, 3.5 mm
long, the anthers ovoid, yellow, about 1.5 mm. long, rounded at apex,
slightly sagittate at base; rudimentary ovary pellucid-pilose. Fruit
ovoid, about 1.6 cm. long and 1.3 cm. across, glabrous, lenticellate;
persistent sepals reflexed, the styles about 2-3 mm. long.

Southwestern China, in southern Yunnan only, in mountain forests
at altitudes of about 1500-2300 meters. Flowers whitish.

CHINA: Yunnan: Feng-ch’un-ling, south of Red River, A. Henry 10696
(A, NY, US, corypes); Meng-tsu, A. Henry 11334 (A, NY, corypgs);
P’ing-pien Hsien, H. T. Tsai 62669 (A).

The two Henry numbers are the two original collections cited by
Dunn. In 1911, Dunn also listed Wilson 3271 & 4764, from O-mei-shan,
as belonging to this species. However, in Rehder’s treatment (in Sarg.
Pl. Wils. 2: 381. 1915), these two numbers are included in A. tetramera
Maxim. I agree with Rehder that these two should not be referred to
A. rubricaulis. The latter species, therefore, is limited in its range to
southern Yunnan.

Dunn describes the plant as wholly glabrous, but upon close exam-
ination the young shoots are found to be minutely hirsute. Henry
10696 is from a male flowering plant with narrow oblong-lanceolate
leaves. Henry 11334 is a fruiting specimen with quite different leaves,
which are ovate-oblong, relatively much shorter and broader. In
other characters the two are clearly of the same species, as especially
evidenced by the firm solid pith, which appears to be a very character-
istic feature of this species, overlooked by Dunn. The field labels
show that the two collections are from two different localities and are
not from the same place, as cited by Dunn. T'sai 62669 is a young fruit-
ing specimen, the only collection of the species since it was described
from the original material.

Only rudimentary ovaries in the staminate flowers are observed.
Dunn originally described these as “praeter lanan circa stylos glab-
rum.” In his later revision of the genus, he characterized the species
as having glabrous ovaries and he accordingly differentiated it in his
key from A. callosa and A. coriacea, which have pubescent ovaries.
This was apparently due to his inclusion of the two Wilson collections
noted above that should properly be placed in A. tetramera, a species
with distinctly glabrous ovaries.

Another characteristic feature of this species which Dunn fails to
note is the presence of flowers usually on short lateral branches, espe-
cially on the lower leafless part. This is true in both staminate and
pistillate specimens. This phenomenon is also found in A. coriacea,
another species with solid pith, and it indicates their close relationship.
The flower color of A. rubricaulis is not recorded by Dunn, but is indi-
cated by the collector on one sheet (NY) of Henry 10696 as whitish.

1952] LI, REVIEW OF THE GENUS ACTINIDIA 37

21. Actinidia fortunatii Finet & Gagnep. in Bull. Soc. Bot. France 53:
574. 1906, as A. fortunati; Dunn in Jour. Linn. Soc. Bot. 39: 409.
1911; Lév. Fl. Kouy-Tcheou 413. 1915; Rehder in Jour. Arnold
Arb. 15: 97. 1934.

Actinidia dielsii Lév. in Rep. Sp. Nov. 13: 175. 1914.

Actinidia glaucophylla F. Chun in Sunyatsenia 7: 11, pl. 3. 1948. Syn. nov.

High climbing shrubs to 10 m.; branches dark reddish brown, glab-
rous, the young branchlets sometimes with densely rusty-pubescent
buds; pith small, lamellate, white. Leaves thin- to thick-chartaceous,
lanceolate to elliptic-lanceolate, rarely ovate-oblong, about 8-14 cm.
long and 1.8-3.5 em. broad, sometimes to 14 em. long and 7.5 em.
broad, long-acuminate to rarely acute at apex, obliquely but distinctly
cordatulate at base, the upper surface generally green, glabrous, rarely
sparsely setose and slightly puberulous along the costa when young,
the lower surface glabrous, mostly glaucous, sometimes puberulous
along the costa and veins when young, the costa and veins subcon-
spicuous above, elevated and distinct beneath, the secondary veins
about 8-10 per side, the veinlets reticulate, inconspicuous above, sub-
conspicuous to conspicuous beneath; petioles 1-2 cm. long, glabrous,
sometimes pubescent. Flowers reddish, the staminate in short, small
cymes, the pistillate often solitary; pedicels slender, to 5 mm. long,
slightly pubescent to glabrate; bracts linear, minute; sepals 5, ovate,
about 4 mm. long and 2.5 mm. broad, often unequal, obtuse at apex,
glabrate; petals 5, obovate, about 7 mm. long and 3.5 mm. broad,
rounded at apex; stamens numerous, the filaments 3-4 mm. long, the
anthers yellow, 1-1.5 mm. long, rounded at apex, slightly sagittate at
base; ovary conical-ovoid, slightly pellucid-pilose, later glabrate. Fruit
cylindric, oblong, to about 2.2 em. long and 2 em. across, glabrous,
lenticellate, blackish.

Southern China (southern Hunan, Kweichow, Kwangsi, and Kwang-
tung), in thickets at altitudes of 400-1300 meters. Flowers reddish or
pinkish, June.

CHINA: Hunan: Hsin-ning Hsien, C. S. Fan & Y. Y. li 497 (A).
Kweichow: Kuei-ting, Y. Tsiang 5480 (NY). Kwangsi: Ling-ch’uan
Hsien, W. T. Tsang 28427 (US); N. Lu-chen, R. C. Ching 5828 (NY),
6085 (NY). Kwangtung: Lo-ch’ang, C. L. Tso 20682 (NY), W. T.
Tsang 20803 (NY); Yao-shan, S. S. Sin 9460 (NY); Ju-yiian Hsien, S. P.
Ko 62844 (A).

This species is not classified by Dunn in his key. He says: “In the
absence of fruit or female flowers it is not possible to decide into which
of the three sections it falls. Its long narrow cordate leaves distinguish
it from all other species here enumerated.”

A photograph of the type, Cavalerie & Fortunat 235, from Kweichow,
is in the herbarium of the Arnold Arboretum. The specimens now avail-
able show that the fruits are oblong, glabrous, and lenticellate. It is
clearly associated with the A. callosa group and especially with A.
coriacea, which has similarly reddish flowers.

38 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXXIII

Actinidia dielsit Lév., as noted by Rehder, is reduced to the synonymy
of A. fortunatii by Léveillé himself. Actinidia glaucophylla F. Chun
is based on S. P. Ko 52835 (type), 52844 and 52886, all from Ju-yuan,
Kwangtung. Of these, a specimen of 52844 1s available. This specimen,
as well as the original description and illustration, shows that without
doubt the species in question belongs to A. fortunatu, a very character-
istic species readily recognized by the narrow elliptic-lanceolate leaves
with a distinct cordatulate base and by the reddish flowers.

The series of specimens now available shows that the species is also
a very variable one. The indumentum and leaves, as well as the flower-
color, show variations. While the plant has a decidedly glabrous ap-
pearance, hairs may be found occasionally on young shoots, buds, and
pedicels. Tsang 20803, a vegetative shoot, has the stem rusty-tomen-
tose and the leaves hairy on the costa on both surfaces. Some of the
leaves are also sparsely setose above. The leaves of the different speci-
mens may be very distinctly glaucous in some and partly glaucous,
slightly glaucous, or not at all glaucous in others. Ching 5828, evidently
a very vigorous flowering branch, has unusually large broad leaves
measuring 14 em. long and 7.5 em. broad. In spite of this great differ-
ence in size, all other characters indicate that it clearly belongs to
this species, which is of polymorphic habit like most other species of
the genus

22.

bo

Actinidia coriacea (Finet & Gagnep.) Dunn in Jour. Linn. Soe.
Bot. 39: 405. 1911; Rehder in Sarg. Pl. Wils. 2: 384. 1915; Stapf
in Bot. Mag. 152: t. 9140. 1928; Hand.-Mazz. Symb. Sin. 7: 390.
1931.

ae et callosa Lindl. var. coriacea Finet & i aa in Bull. Soe. Bot.
France 52: Mém. 4: 20. 1905 (Contr. Fl. As. Or.).

Sautine shrubs to 10 em.; branches reddish brown, glabrous or
nearly so; pith solid, firm, whitish or yellowish. Leaves thickly cori-
aceous, oblong to oblong-ovate, about 10-16 em. long, 3-5 em. broad,
acuminate at apex, shortly acute to cuneate at base, usually oblique,
the margins more or less remotely mucronulate-serrate to sharply ser-
rate toward the apex, the tips of serrations often reddish-glandular,
glabrous on both surfaces, the upper surface green, the lower paler,
the costa suleate above, thick and slightly raised beneath, the secondary
veins about 6 or 7 per side, slender, arcuate-ascending, anastomosing,
inconspicuous above, scarcely raised and subconspicuous beneath, the
veinlets reticulate, inconspicuous on both surfaces; petioles 1.5-2.5 em.
long, glabrous. Flowers reddish, solitary or in 2-4-flowered cymes
arranged along short branches which are leafy above and leafless below,
the lower flowers lateral, the upper axillary; pedicels slender, up to
2.2 cm. long, glabrous; sepals 5, ovate, about 4-5 mm. long and 3.5
mm. broad, obtuse at apex, glabrous without, sometimes white-puberu-
lent inside, ciliate along the margins; petals 5, suborbicular, red with

1952] LI, REVIEW OF THE GENUS ACTINIDIA 39

whitish or yellowish margins, about 7-10 mm. long and 4~7 mm. broad,
the apex rounded, the base narrowed; stamens numerous, the filaments
red, about 3-4 mm. long; anthers yellow; ovary conical, about 2.5 mm.
long and 1.5 mm. across, densely white-pubescent, the styles to 3 mm.
long. Fruit ovoid or globose, to 2 em. long, glabrous, brown, white-
lenticellate.

Southwestern China (from Kweichow and Szechuan to northwestern
Yunnan), in thickets at altitudes of 200-1000 meters. Flowers red,
May—June.

CHINA: Szechuan: Without precise localities, 2. H. Wilson 3272 (A),
3272a (A), E. Faber 72 (NY); Cheng-k’ou Hsien, R. P. Farges 1946 (NY);
Hung-ya Hsien, LE. H. Wilson 932 (A, GH, NY); O-mei-shan, FE. H. Wilson
4760 (A), W. P. Fang 3306 (A), 12756 (A, US), Chiao & Fan 132 (A), Y.S.
Liu 2145 (A); O-pien Hsien, Y. S. Liu 2003 (A); Lu-shan Hsien, K. L. Chu
4044 (A); Nan-ch’uan Hsien, W. P. Fang 796 (A, NY, US), 5654 (A, NY);
Ch’ia-ting Hsien, S. C. Sun & K. Chang 25 (A), 1512 (A), L. Y. Tai 643 (A),
832 (A), 1289 (A), 1541 (A); Ch’i-chiang Hsien, W. P. Fang 1430, (A, NY);
Kuan Hsien, W. P. Fang 2113 (A, NY); Chiang-yu Hsien, W. P. Fang
2293 (A). Sikang: Tien-ch’iian Hsien, L. Y. Ta 4172 (A). Kweichow:
Tsun-i Hsien, Steward, Chiao & Cheo 136 (A, NY, US); Shih-ch’ien Hsien,
Y. Tsiang 4108 (NY); T’ung-tzu Hsien, Y. Tsiang 5010 (NY); Tu-yun Hsien,
Y. Tsiang 5698 (NY); Chiang-k’ou Hsien, Y. Tsiang 7507 (NY); Yin-chiang
Hsien, Y. T'siang 7610 (NY), 7894 (NY).

This distinct species is characterized by the coriaceous, remotely
glandular-serrulate leaves, and by the red flowers borne on separate
leafless branches or along the leafless portion of the shoots. The pith
is characteristically solid, firm, and more or less whitish or yellowish.

23. Actinidia asymmetrica F. Chun in Sunyatsenia 7: 13. 1948.
Climbing shrubs; branches reddish brown, longitudinally sulcate,
glabrous, without lenticels or sometimes with a few pale oblong lenti-
cels; pith white, medium-sized, lamellate. Leaves membranaceous or
chartaceous, ovate-oblong, about 7-10 em. long and 3.5-5.3 em. broad,
acute to acuminate at apex, auriculate-cordate at base, often unequally
so, the margins irregularly callose-denticulate, glabrous on both sur-
faces, paler or even glabrescent on the lower surface, the costa and veins
subconspicuous above, distinct and raised beneath, the secondary veins
about 5-7 per side, arcuate-ascending, ending in the marginal teeth,
the veinlets reticulate, inconspicuous except for a few parallel cross-
bars; petioles 2-2.25 cm. long, glabrous; inflorescences in short 3-5-
flowered axillary cymes, sometimes the flowers solitary; peduncles
1-1.5 em. long, glabrous; pedicels 4-5 mm. long; bracts minute, subu-
late. Flowers pink; sepals 5, subequal, oblong, about 6 mm. long and
3 mm. broad, acute to obtuse at apex, glabrous, the margins slightly
ciliate; petals 5, obovate to obovate-oblong, about 8 mm. long and
6 mm. broad, rounded at apex, contracted at base; stamens numerous,
the filaments about 3 mm. long, the anthers yellow, about 1.8 mm.
long, ovary cylindric-oblong, about 3 mm. long and 2 mm. across,

40 JOURNAL OF THE ARNOLD ARBORETUM [VoL. XXXIII

densely yellowish-villose, the styles about 2 mm. long. Fruit cylindrical
ovoid, about 2.2 cm. long and 8-10 mm. across, brown, with lenticels,
glabrous or slightly pubescent toward the tip; seeds oval, 1.5 mm.
long, 1 mm. broad, foveolate-reticulate.

Southern China, in Kwangsi only, on mountain slopes at an altitude
of about 1360 meters. Flowers reddish.

CHINA: Kwangsi: Ling-yiian Hsien, Steward & Cheo 672 (NY);
Nan-ning Hsien, Shih-wan-ta-shan, R. C. Ching 8362 (NY); Shang-ssu Hsien,
Shih-wan-ta-shan W. 7. Tsang 23833 (NY), 24120 (N

The type, a flowering specimen, is Liu 566 (Sunyatsen Herb. 86950),
from Wu-ming, Ta-ming-shan, Kwangsi. It has not been seen by me.
Among the above cited specimens, the Ching and Tsang numbers are
fruiting specimens. Steward & Cheo 672 has female flowers and young
fruits.

The fruit of this species, not known when it was first described, ap-
pears to be very characteristic in its nodding position and slender shape.
This species is also distinetly characterized by the leaf with auriculate-
cordate base. Chun emphasizes particularly the asymmetrical shape
of the leaf-blade, but this is not an exclusive character for the species.
Most species of the genus have some leaves that are asymmetrical in
shape. It is a common phenomenon in most climbing vines and is
probably due largely to environmental rather than genetic factors.

Sin. 7: 390.
Actinidia callosa Lindl. var. tas Finet & Gagnep. in Bull. Soe. Bot.
France 52: Mém. 4: 19, 20. 1907; Dunn in Jour. Linn. Soc. Bot. 39: 406.

24. Actinidia nea Cae & Gagnep.) Stapf ex Hand.-Mazz.
Symb.

Climbing shrubs to 7 m.; branches reddish brown, with scattered
short ovoid lenticels, glabrous or the young branchlets sparsely whitish-
pubescent; pith more or less large, white, thinly lamellate. Leaves
membranaceous, ovate-oblong, 5-12 em. long, 3-6.5 cm. broad, acumi-
nate at apex, broadly truncate at base, often unequal, the margins
finely mucronulate-serrulate, the teeth more or less glandular, dark
green above, nearly concolored or very slightly paler beneath, slightly
but distinctly white-pubescent along the costa, the costa and veins
subconspicuous above, purplish and conspicuous but not raised beneath,
the secondary veins about 7 or 8 per side, arcuate-ascending, anastomos-
ing, the veinlets reticulate, inconspicuous to subconspicuous beneath,
ending in the marginal teeth; petioles purplish, long, slender, about
2-3.5 em. long, white-pubescent to glabrate. Inflorescence in axillary
cymes of about 5-7 flowers, brownish-tomentose; peduncles 1—1.5 cm.
long; pedicels to 1 em. long; bracts linear, small, 0-3 to a pedicel.
Flowers brownish yellow; sepals 5, ovate, about 4-5 mm. long and 2-2.5
mm. broad, obtuse at apex, glabrous to slightly pubescent, the margins
often ciliate; petals 5, obovate-oblong, about 7-9 mm. long and 4—5 mm.
broad, rounded at apex; stamens numerous, the filaments slender, to

1952] LI, REVIEW OF THE GENUS ACTINIDIA 41

5 mm. long, the anthers ‘yellow, ovoid, about 1 mm. long, rounded
at both ends, the base not divergent; ovary elongate-cylindric, to 5 mm.
long and 2.5 mm. across, pubescent when young, becoming glabrous, the
styles about 2 mm. long. Young fruits oblong-cylindric, glabrous, not
lenticellate, with persistent reflexed sepals.

Southwestern China (Yunnan) and northern Burma, in forest at
altitudes of 2100-3300 meters. Flowers brownish yellow, June.

CHINA: Yunnan: Without precise locality, G. Forrest 6385 (A), 13910
(A), 19245 (A); “Tsekou,” R. P. Soulie 1396 (A, ISOTYPE); Mekong Valley,
J. F. Rock 8948 (A, NY, US), Handel-Mazzetti 8825 (A); Mekong- Yangtze
divide, G. Forrest 19442 (A), J. F. Rock 25067 (A); Yung-shan Hsien, H. T.
Tsai 51000 (A).

BURMA: Northern Burma, Adung Valley, F. K. Ward 9559a (A).

The type is from “Yunnan; Thra-na at Tsekou, 10-20 juin 1893, no
1396 (Soulie)”, of which a duplicate is cited. This species is amply
different from A. callosa in the very thin broadly truncate leaves,
often hairy and setose above, the large thinly lamellate pith, and the
brownish yellow flowers. The degree of pubescence varies considerably.

The specific combination was published by Handel-Mazzetti from
the herbarium name indicated by Stapf. When Handel-Mazzetti pub-
lished it he cited three collections, Forrest 13910 and Handel-Mazzetti
9042 and 9046. Forrest 13910 was once identified by W. W. Smith as
“A. championii var. mollis Dunn.” This number and Handel-Mazzettz
9042 are available, and they clearly prove to be A. callosa var. pubes-
cens and not A. pilosula as Handel-Mazzetti originally indicated. The
two are very different from A. pilosula in leaf and pith, as well as in
floral characters. Clearly A. pilosula should be typified by Finet and
Gagnepain’s type, not by the specimens mistakenly identified by
Handel-Mazzetti (and possibly also by Stapf) when the combination
was effected and published.

The three yellow-flowered species, A. pilosula, A. trichogyna, and
A. venosa are intimately related to each other. They are generally
distinguishable, but at times intermediate forms are found. It may
eventually prove desirable to combine the three as varieties of one
species.

25. Actinidia venosa Rehder in Sarg. Pl. Wils. 2: 385. 1915; Hand.-
Mazz. Symb. Sin. 7: 390. 1931, p. p.

Actinidia callosa forma D. Dunn in Jour. Linn. Soc. Bot. 39: 406. 1911.

Climbing shrubs to 9 em.; branches purplish brown, with oblong
whitish lenticels, the young branchlets puberulous to tomentose, soon
glabrescent; pith large, white, lamellate. Leaves thin-chartaceous,
ovate or ovate-oblong to elliptic and elliptic-oblong, about 5-15 cm.
long, 3-6 em. broad, rarely to 7.5 em. broad, acuminate to long-acumi-
nate at apex, mostly rounded at base and sometimes subcordate, usu-
ally obliquely so, the margins denticulate-serrulate, the upper surface
dark green, glabrous or rarely sparsely setose when young, the lower

42 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXXII

surface slightly paler, more or less tomentose on the veins when young,
soon glabrescent or glabrous, the costa and veins conspicuous above,
strongly raised and prominent beneath, the secondary veins 7-11 per
side, nearly straight or arcuately ascending, anastomosing, the veinlets
reticulate, with numerous closely parallel strongly raised and very
prominent cross-bars; petioles slender, 1.5-4 cm. long, puberulous or
glabrate at first, soon glabrous. Inflorescences in axillary cymes of
1—7 flowers, rusty-tomentose, the peduncles 5-10 mm. long, the pedicels
almost as long; sepals ovate-oblong, about 5 mm. long, obtuse at
apex, rusty-tomentose; petals elliptic-oblong, about 8-10 mm. long
and 5-6 mm. broad, rounded at apex; stamens numerous, the filaments
slender, 5-6 mm. long, the anthers oblong, 2-3 mm. long, rounded at
apex, densely villose, the styles 3-4 mm. long. Fruit ovoid or sub-
globose, about 1.5 em. long, brown, glabrous.

Western China (Sikang, Szechuan, and northern Yunnan) and north-
eastern India (Khasia), in thickets at altitudes of 1000-3650 meters.
Flowers buff-yellow, June—July.

CHINA: Szechuan: Without precise locality, E. H. Wilson 3275 (A);
Kuan Hsien, W. P. Fang 2220 (A, NY), 2258 (A, NY); Mao Hsien, W. P.
Fang 21947 (A), ied (A); Wen-ch’iian Hsien, 2. H. Wilson 888 p. p. (A),
1029 (A, Type); “Yang-ching”’ ae E. H. W see 891 p. p. (A, GH, US);
O-mei-shan, /. H. Wilson 4765 (A), W. P. Fang 12835 (A), F. T. Wang 23348
(A), H.C. iow 8108 (A), 1225 . 12310 (A), L. Y. Tai 34 (A), S. C. Sun
& K. Chang 273 (A); Wa-shan, FE. H. Wilson 888 p. p. (A), 891 p. p. (A);
Wa-wu-shan, 2. H. Wilson 888 p. p. (A); Pao-hsing Hsien, K. L. Chu 3269

A). Sikang: Kan-ting (Tachienlu), A. E. Pratt 101 (GH); E. H. Wilson
1029 (GH, US), C. ¥. Chiao 1650 (A), 2046 (A). Yunnan: Without
precise locality, H. T. Tsai 57375 (A); “Sua-kia,” FE. E. Maire 28 (A);
Chienchuan-Mekong divide, G. Forrest 21521 (A, US), 22278 (A, US),
22283 (A); Yun-lu-shan, Yangtze-Mekong watershed, J. F. Rock 25067 (NY);
Shun-ning, 7’. T. Yii 16219 (A); Chung-tien, K. M. ag! 1786 (A); Li-chiang,
R. C. Ching 20462 (A), 21464 (A); Wei-hsi ete : oF. — 57998 (A),

59692 (A), C. W. Wang 63802 (A), 64031 (A ee (A), 68110 (A);

Lan-p’ing, H. T. Tsai 53741 (A).

INDIA: Khasia, J. D Hooker & T. Thomson s. n. (GH).

This species is related to A. callosa but is distinguished by the
broader leaves with a larger number of secondary veins, by numerous,
distinctly raised, very prominent, more or less parallel cross-bars, and
by the large white pith and the yellow flowers. The specimens cited
belong to the typical form of the species, which may be designated as
forma venosa.

25a. Actinidia venosa Rehder forma pubescens f. nov.
A f. venosa foliis subtus pubescentibus differt.
Western China, in eastern Sikang only, at altitudes of 2700-3200
meters.
CHINA: Sikang: Hui-li Hsien, 7. 7. Yii 1451 (A, Type); “Yenyuen,”
Yalung River, H. Handel-Mazzetti 5406 (A).

1952] LI, REVIEW OF THE GENUS ACTINIDIA 43

A form characterized by the leaves being more or less densely
pubescent below. Only young fruiting specimens are observed. The
fruit is brownish.

96. Actinidia trichogyna Franch. in Jour. de Bot. 8: 278. 1894;
Rehder in Sarg. Pl. Wils. 2: 384. 1915.

Actinidia callosa Lindl var. trichogyna Finet & Gagnep. in Bull. Soc. Bot.
France 52: Mém. 4: 20. 1906 (Contr. Fl. As. Or.); Dunn in Jour. Linn.
Soc. Bot. 39: 406. 1911.

Climbing shrubs to 7 m.; branches dark reddish brown, with oblong
pale lenticels, the young branchlets puberulous to tomentose, soon
glabrescent; pith large, white lamellate. Leaves thin-chartaceous, ovate
to ovate-oblong to elliptic-ovate, about 5.5-8.5 em. long and 3-5 cm.
broad, acute to acuminate at apex, broad-truncate or rounded to sub-
cordate at base, often obliquely so, the margins mucronulate-serrulate,
the upper surface dark green, more or less tomentose along the costa
and veins, soon glabrous, the lower surface glabrous or slightly puberu-
lous along the costa, glaucous, the costa and veins inconspicuous above,
purple and distinct but not raised beneath, the secondary veins about
5 or 6 per side, rarely more, arcuate-ascending, anastomosing, the
veinlets finely reticulate, often in parallel cross-bars, inconspicuous
above, purple and subconspicuous beneath; petioles 2-3 cm. long,
tomentose to glabrate, purple. Inflorescence in axillary cymes of 1-3
flowers, tomentose, the peduncles about 5 mm. long, the pedicels about
the same in length. Flowers yellow; sepals 5, ovate, about 4 mm. long
and 2.5 mm. broad, acute to obtuse at apex, tomentose to glabrate;
petals 5, obovate-oblong, about 8 mm. long and 4 mm. broad, rounded
at apex; stamens numerous, the filaments very slender, about 3-4 mm.
long, the anthers yellow, oval, about 1 mm. long, rounded at apex, the
base not divergent; ovary subglobose, about 2-3 mm. long, densely
villose, the styles about 3 mm. long. Fruit subglobose, about 1 cm.
across, brown, glabrous.

Western China (western Hupeh and eastern Szechuan to northwestern
Yunnan), in scrubs and thickets at altitudes of 2800-38000 meters.
Flowers yellow, tinged rose, June.

CHINA: Hupeh: Western Hupeh, A. Henry 7135 (GH), E. H. Wilson
2204 (A, US). Szechuan: Chen-k’ou Hsien, Farges 370 (NY, isotype).

unnan: Without precise locality, G. Forrest 7906 (A), 17781 (Ay
Chienchuan-Mekong divide, G. Forrest 21522 (A, US), J. F. Rock 8618 (A,
US); Mekong-Yangtze divide, J. F. Rock 9403 (A, US), 10373 (A, US);
Li-chiang, R. C. Ching 20841 (A), 22004 (A); Salwin-Chiuchiang divide,
T.T. Yu 19248 (A).

This species is in close alliance with A. pilosula and A. venosa, both
with broad-based leaves, tomentose inflorescences, large white piths,
and yellow flowers. It can, however, be distinguished particularly by
its glaucous leaves. In this character it resembles A. sabiaefolia, a
species of eastern China, but the latter has smaller, crenate-serrulate

44 JOURNAL OF THE ARNOLD ARBORETUM [VoL, XXXII

leaves and glabrous sepals. In the glaucous leaves, A. trichogyna also
resembles A. melanandra Franch., of western China, but the latter has
purple instead of yellow anthers and also narrower leaf-bases which
are not rounded to subcordate.

27. Actinidia glabra sp. nov.

Frutex scandens, 6-14 m. altus, omnino glaber; ramis purpureo-
nigrescentibus, plus minusve striatis, lenticellis pallidis parvis puncti-
formibus, medulla ampla brunnea lamellata; foliis subcoriaceis,
utrinque glabris, oblongo-ovatis, circiter 6-9 em. longis et 3-4.5 cm.
latis, apice acutis vel acuminatis, interdum obtusis, basi oblique cuneatis
vel subrotundatis. margine integris vel apicem versus sparse et minute
serrulatis, supra atroviridibus nitidis, subtus glaucescentibus, costa
utrinque prominente, nervis lateralibus utrinsecus 6 vel 7, arcuato-
adscendentibus, prope marginem anastomosantibus, venulis reticulatis
utrinque conspicuis; petiolis 1.5-2.2 em. longis; inflorssoentiie ignotis;
fructibus plerumque solitarii, axillaribus, ovoideis, circiter 2.5 em. longis
et 2 cm. latis, flavide brunneis, glabris, lenticellis pallide dispersis, semi-
nibus brunneis ovoideis, leviter compressis, circiter 3 mm. longis et 2
mm. crassis, testa in sicco reticulata

Southern China, in Kwangsi, in thickets, Flowers unknown.

CHINA: Kwangsi: Shih-wan-ta-shan, S. of Nan-ning, R. C. Chin
7875 (NY, Type); Shang-ssu Hsien, Sih-wan-ta- he Teng- lung Village, W. T.
Tsang 24135 (NY).

Although flowers are not known, this seems to be a distinct species
because of its entirely glabrous habit. It is one of the alliance of A. cal-
losa Lindl., from which it can be distinguished by the more coriaceous,
nearly entire leaves, brown lamellate pith, and smaller, brown fruits.

28. Actinidia sabiaefolia Dunn in Jour. Linn. Soc. Bot. 38: 357. 1908;
Chun in Sunyatsenia 4: 190. 1940.

Actinidia callosa Lindl. var. sabiaefolia Dunn in op. cit. 39: 406. 1911.

Climbing shrubs; branches dark reddish, glabrous, sparsely lenticel-
late, the lenticels short, oval, pale; pith small, brown, lamellate. Leaves
chartaceous, ovate, about 5-6 em. long and 2.5-3 em. broad, acute to
sometimes obtuse at apex, rounded at base, sometimes obliquely so, the
margins inconspicuously crenate- mucronulate, glabrous, and very dark
above, much paler and more or less glabrous beneath, the costa‘and veins
inconspicuous above, purplish and distinct but not raised beneath, the
secondary veins about 5 per side, arcuately ascending, anastomosing,
the veinlets reticulate, purple and conspicuous beneath; petioles pur-
plish, about 2 em. long, glabrous, sometimes more or less glaucous. In-
florescences in axillary cymes of 1-3 flowers, glabrous throughout, the
peduncles to 5 mm. long; pedicels to 1 em. long: bracts minute. Flowers
dioecious, white; sepals 5, ovate, 2-3 mm. long, obtuse to rounded at
apex, more or less ciliate wide the margins; petals 5, obovate, 5-6 mm.
long, 3.5-4 mm. broad, rounded at apex; stamens numerous, the fila-

1952] LI, REVIEW OF THE GENUS ACTINIDIA 45

ments slender, about 2 mm. long, the anthers yellow, oval, scarcely 1
mm. long, rounded at apex, not divergent at base; ovary ovoid, about
2 mm. long, densely reddish-tomentose, the styles about 2 mm. long.

Southeastern China (Kwangtung, Kwangsi, and Fukien), in moun-
tain forests. Flowers white.

CHINA: Fukien: Nan-p’ing (Yenping), Hongkong Herb. 1120 (A),
2402 (A, ISOTYPE).

This taxon was first described by Dunn as a distinct species, but sub-
sequently he treated it as a variety of A. callosa. Chun, who records the
plant from Kwangtung and Kwangsi and adds a description of the fruit,
is of the opinion that it should be reinstated as a species. It differs from
A. callosa and its varieties in the consistently smaller, more or less
glaucous leaves, and in the smaller flowers. I have seen no specimens
other than Dunn’s original collections. A larger series of specimens,
from Fukien and the neighbouring provinces of Kwangtung and
Kwangsi, is desirable for study of the variations in this species and its
relationship with A. callosa and its many varieites.

29. Actinidia callosa Lindl. Nat. Syst. ed. 2. 489. 1835.

Climbing shrubs to 7 m.; branches glabrous or the young branchlets
tomentose, dark gray to reddish brown, with distinct conspicuous long
yellowish lenticels; pith solid, firm, light orange colored, or sometimes
slightly and irregularly lamellate in the center. Leaves chartaceous,
obovate to ovate-elliptic or sometimes ovate-lanceolate, about 5-13 em.
long and 2.5-6.5 cm. broad, acuminate at apex, rounded to subcuneate at
base, usually obliquely so, the margins serrulate or sometimes crenately
serrulate or subentire, glabrous or both surfaces tomentose in the nerve-
axils or along the veins on the lower surface, dark and more or less shin-
ing above, pale beneath, the secondary veins about 5-7 per side, more or
less straight or arcuate-ascending, anastomosing, the veins or their
branches ending in the marginal teeth, the veinlets reticulate, incon-
spicuous above, subconspicuous beneath; petioles 2-5.5 cm. long, red-
dish, glabrous or more or less slightly pubescent. Inflorescences in axil-
lary 1-5-flowered cymes, glabrate or slightly tomentose, the peduncles
2.5-15 mm. long, the pedicels 5-20 mm. long, slender. Flowers white;
sepals ovate, about 6 mm. long and 4 mm. broad, obtuse at apex, gla-
brous or tomentose, the margins sometimes ciliate; petals broadly ob-
long-obovate, about 10 mm. long and 7 mm. broad, rounded to obtuse at
apex; stamens numerous, the fllaments slender, 4-6 mm. long, the an-
thers yellow, oval, about 2 mm. long, obtuse at apex, divergent at base;
ovary oblong-ovoid, elliptic, about 3-4 em. long and 2-2.5 em. across,
glabrous, grayish green, with gray or brown lenticels.

Actinidia callosa is a very variable species, covering a wide geographic
area from Taiwan to south-central and western China, and to north-
eastern India and Malaysia. Besides the typical form, which extends
from southern China all the way to Java, there are many other varieties

46 JOURNAL OF THE ARNOLD ARBORETUM [VoL. XXXIII

of more limited geographical ranges that can be recognized. A few other
species, such as A. venosa, A. trichogyna, and A. sabiaefolia, were for-
merly included in A. callosa by Dunn in his very broad concept of the
species. They possess, however, not only definite and more limited
ranges but also several distinct and constant characters, and they are
here treated as distinct entities closely associated with A. callosa.

Key TO THE VARIETIES

A. Pedicels and calyces glabrou
B. Leaves glabrous cnet, not tomentose in the nerve- ~_ beneath.

Bee ong aieig ean gee Ege tay gb pd © ove eae aoa r. callosa.
BB. Leaves white-tomentose in the nerve-axils on the vane surface,
Otherwise GIADTOUS 243 cies 48 eco asec sk tere ses ....-b. var. henriii

AA. Pedicels and calyces tomentose
B. Leaves white-tomentose in the nerve-axils on the —_ phils other-
wise glabrous... 0 ee var. formosana.
BB. Leaves tomentose ‘along the costa and nerves on fie lower oe.
C. Tomentum on leaves rusty-granular, present along veins only. .....
d.

Mane tReREN Gas EAC GL EG TERPS r. indochinensis.
CC. Leaves more or less densely villose- pubescent all over the lower
TTR soo bk 5p ba EER ELS ERS ARS e. var. pubescens.

29a. Actinidia callosa var. callosa.
Actinidia callosa Lindl Nat. Syst. ed. 2. 439. 1885; Dunn in Jour. Linn.
39: 405. 1911; Lév. Fl. Kouy-Tcheou 413. 1915; Rehder in
Sarg. Pl. Wils. 2: 382. 1915; Koord. Exk. Fl. Java 2: 602. 1912, FI.
Tjib. 2: 179. 1923; Baker in Jour. Bot. 62: Suppl. 9. 1924; Van Steenis
in Bull. Bot. Gard. Buitenz. III. 13: 174. 1934, Fl. Males. I. 4: 37. 1948

Leaves chartaceous, obovate to ovate-elliptic, rarely ovate-lanceolate,
about 3-13 cm. long and 2.5-6 em. broad, acuminate at apex, glabrous
on both surfaces; pedicels and calyces glabrous.

Southern Asia, from southern China, Indo-China, and northeastern
India to Malaysia, in forests and thickets, at altitudes of 300-2400
meters. Flowers white, rarely yellowish, April-May

CHINA: Anhwei: Hsiu-ning Hsien, R. C. Ching 3321 (A), Wu-yiian
Hsien, R. C. Ching 3267 (A). Chekiang: Li-shui Hsien, Y. L. Keng 4235
(A), H. H. Hu 464 (A); Ch’ing-yiian, R. C. Ching 2487 (A). Kiangsi:
Lung-nan Hsien, S. K. Lau 4690 (US). Fukien: Ch’ung-an Hsien, H. H.
Hu 1336 (A); Ku-shan, H. H. Chung 6706 (A). Hunan: Ch’ang-ning Hsien,
Fan & Ii 176 (A); I-chang Hsien, W. T. Tsang 23757 (US). Kwangs1:
Ling-yun Hsien, Steward & Cheo 97 (NY), 428 (A, NY), 1306 (A); Yung
Hsien, Steward & Cheo 906 (A, NY); Kwei-lin Hsien, W. 7. Tsang

unnan: without precise localities, G. bie 12057 (A), 158
(A), 17939 (A); Mekong-Salwin divide, west of Wei-hsi, J. F. Rock ne
(A); Shweli-Salwin divide, G. Forrest 24261 (US), 24380 (US) ; hier -ning,
T.T. Yu 15998 (A), 16150 (A), 16613 (A); Mien-ning, T. T. Yii 17780 (A);
south of Red River, A. Henry 10056 (US), 10056A (A, US), 10056B (A, NY);
Meng-tzu, A. Henry 10824 (A, NY), 10824A (A, NY); Pi-tsieh Hsien, H. T.
Tsai 52784 (A); Fo-hai, C. W. Wang 73984 (A), 77266 (A); P’ing-pien Hsien,
H. T. Tsai 62630 (A); Chen hsiung Hsien, H. T. Tsai 52686 (A); Liang- te

1952] LI, REVIEW OF THE GENUS ACTINIDIA 47

Hsien, H. T. Tsai 51342 (A). Kwangtung: Lo-ch’ang, Y. Tsiang 1305 (A),
1470 (A), C. L. Tso 20489 (NY), 20536 (NY); Chao-chou Hsien, R. Mell
3883 (A); Tseng-chen Hsien, W. T. Tsang 20304 (NY, US); Ta-pu Hsien,
W.T. Tsang 21150 (A, NY); Wen-yiian Hsien, S. K. Lau 2117 (A); dieyuau,
S. P. Ko 52555 (A), 52563 (A).

INDIA: Nepal, Wallich 6634 (NY, 1sorypr); Garhwal, Hast India Co. 305
(GH), A. LE. Osmaston 822 (A); Sikkim, J. D. Hooker s.n. (GH); W. Nepal,
Rikbar, Doti district, Bis Ram 400 (A); Dehra Dun, Mulkh Raj Beli s. n.
(A), T. Hisain s. n. (A); Pittoragart, E. Almoan, A. FE. Osmaston 1476 (A).

INDO-CHINA: Tonkin, Chapa, A. Pételot 6343 (A), 6344 (A); Laokay,
Tonkin, A. Pételot 8582 (A).

This typical form of the species covers a wide range and exhibits con-
siderable variation, especially in the shape, size, and margin of leaves.
These variations, however, are not constant and they do not seem to be
associated with either other morphological characters or geographical
ranges. Considering the variable nature of individual plants in this
genus, it is not attempted to further divide these forms taxonomically.

29b. Actinidia callosa var. henryi Maxim. in Acta Hort. Petrop. 11:

36. 1890; Lév. Fl. Kouy-Tcheou 413. 1915; Rehder in Sarg. Pl. Wils.
2: 382. 1915.

saan Sia Dunn in Kew Bull. 1906: 1. 1906; Hand.-Mazz. Symb.
Sin. 7 ok,

Actinidia oe var. typica forma C. Dunn in Jour. Linn. Soc. Bot. 39:
409. 1911

Actinidia callosa sensu Diels in Bot. Jahrb. 29: 470. 1900. non Lindl.

Actinidia venosa sensu Hand.-Mazz. Symb. Sin. 7: 390. 1931, p. p., non
Rehder.

Leaves chartaceous, oblong-lanceolate to ovate, about 6-12 cm. long
and 3-6.5 em. broad, acuminate at apex, glabrous on both surfaces ex-
cept for tufts of white tomentum in the axils of the secondary nerves on
the lower surface; pedicels and calyces more or less glabrous.

Southwestern China (western Hupeh, Szechuan, and western Kwei-
chow to southwestern Hunan) in thickets, at altitudes of 1700-3500
meters. Flowers white, May—June.

CHINA: Hupeh: I-chang and vicinity, A. Henry 3494 (A, GH, US),
3564 (GH), 3955 (GH, NY, US), 4377 (US) (all four numbers eae o
A. curvidens Dunn), 4377A (GH), 5797 (A, GH, US), 6494 (GH),
Wilson 348 (A, NY, US), 512 (A, GH, US), 2096 (GH), 2204 (NY); pe

Hsien, FE. H. Wilson 2012 (A, GH US); Pa-tung Hsien, H. C. Chow 240
(A, NY), 836 (A, NY); Chica Hsien, H. C. Chow 1633 (A, NY);
Kiian-yin- tang, W.Y. Chun 5232 (A); “Suao-ya- tsze,” W. Y. Chun 3627 (A).
Szechuan: Without precise locality, A. Henry 7243 (GH); Wa-shan,
E. H. Wilson 2016 (A, US); Cheng-k’ou Hsien, R. P. Farges s. n. (NY);
Nan-ch’uan, Bock & Rosthorn 1824 (A), W. P. Fang 827 (A), 1079 (A), 1105
(A), 1113 (A), 1350 (A); Kuan Hsien, F. T. Wang 20812a (A); O-mei-shan,

E. H. Wilson 4762 (A), W. P. Fang O45 (A), 7667 (A), Chiao & Fan 376 (A);
amet Hsien, Ff. T. Wang 23087 (A); O-pien Hsien, 7. T. Yu 738 (A), 846
(A), Y. S. Liu 2083 (A). Kweichow: Kwei-ting, H. Handel-Mazzetti

48 JOURNAL OF THE ARNOLD ARBORETUM [VoL. XXXIII

10635 (A); Fan-ching-shan, Steward, Chiao & Cheo 587 (A, NY); Tung-tzu,
Y. Tsiang 5168 (NY); Tu-yun, Y. Tsiang 5669 (NY), 5748 (NY), 6075 (NY).

Actinidia curvidens Dunn was reduced by Dunn himself to this vari-
ety, which was recognized by him as a form of A. callosa. This taxon
closely resembles the typical variety of the species, but it may be distin-
guished by the presence of hairs in the nerve-axils on the lower surface
of the leaves.

29c. Actinidia callosa var. formosana Finet & Gagnep. in Bull. Soc.
Bot. France 52: Mém. 4: 20. 1905 (Contr. Fl. As. Or.); Dunn in
Jour. Linn. Soe. Bot. 39: 406. 1911; Hayata, Icon. Pl. Formos, 4: 2.
1914.
Actinidia formosana Hayata, Icon. 7 oo 8: 12. 1919; Kanehira,
Formos. Trees rev. ed. 449, f. 407. 1

Leaves chartaceous, inn de to ovate, about 6-14 em. long
and 2.5-6 cm. broad, shortly acuminate at apex, glabrous on both sur-
faces except for tufts of white tomentum in the axils of the secondary
veins on the lower surface; pedicels rusty-tomentose; sepals rusty-
tomentose on both surfaces.

China, in Taiwan only, from the central to the northern part of the
island, in forests at altitudes of 1160-2000 meters. Flowers, April-May.

A: Taiwan: Tamsui, Morse 1388 (A); Taipei, FE. H. Wilson
10195 (A), 10257 (A), 10258 (A), T. Tanaka & Y. Shimada 11010 (NY, US),
S. Sasaki s.n. (A), S. Suzuki s.n. (A); Chiao-pan-shan, Y. Shimada s. n. (A);
Mt. Arisan, E. H. Wilson 10862 (A), 10897 (A); Rengachi, 7. Hayashi
21221 (A).

This Taiwan plant is very close to the mainland forms. It resembles
the variety henry? of western China, in having tufts of hairs in the vein-
axils on the lower leaf-surface. The leaves are usually strongly bicol-
ored, like those of plants of the typical form of the species from the
neighboring provinces of Fukien and Kwangtung.

29d. Actinidia callosa var. indochinensis (Merrill) comb. nov.

Actinidia indochinensis Merrill in Jour. Arnold Arb. 19: 53. 1938.

Young branchlets and inflorescences rusty-granular-tomentose ; leaves
subchartaceous when young, becoming coriaceous when mature, ovate,
6-10 cm. long, 2.5-6 cm. broad, dark and glabrous above, paler and
rusty-granular-tomentose along the costa and veins beneath, the mar-
gins subentire to inconspicuously and remotely crenate-serrulate toward
the apex.

Southwestern China, in Yunnan, and Indo-China, in Tonkin, in for-
ests at altitudes of 1400-1 900 meters. Flowers white, April.

CHINA: Yunnan: Without precise locality, J. C. Liu & C. Wang
85443 (A).

INDO-CHINA: Tonkin, Chapa, A. Pételot s. n. (A), 4406 (US), 5938 (A,
TYPE), 5940 (A); Tonkin, Lao-kay, 2. Poilane 17004 (A), 21688 (A).

1952] LI, REVIEW OF THE GENUS ACTINIDIA 49

This variety is distinguished by the granular indumentum on the
young branchlets, inflorescences, and along the costa and veins of the
lower leaf-surface.

29e. Actinidia callosa var. pubescens Dunn in Jour. Linn. Soc. Bot. 39:
406. 1911; Van Steenis, F]. Males. I. 4: 39. 1948

Saurauia tomentosa Korth. ex Koord. & Val. Bidjr. 3: 280. 1896, nomen.

Actinidia pubescens Ridley in Jour. Fed. Mal. Stat. Mus. 8(4): 18. 1917.

Leaves chartaceous, ovate, about 6.5-12 em. long and 4-8 em. broad,
acute at apex, rounded to truncate at base, thinly tomentose all over the
lower surface; pedicels and calyces rusty-tomentose.

Southwestern China (Yunnan) and Assam to Malaysia (Malay
Peninsula, Sumatra, Java?), in forests at altitudes of 2000-2500 meters.

CHINA: Yunnan: Mekong-Salwin divide, “Alulaka,” T. T. Yu 19107
(A); Salwin valley, “Sekai,” 7. 7. Yii 23001 (A); near “Bahan,” H. Handel-
Mazzetti 9042 (A); Meng-tzu, A. Henry 10780 (A, NY, US); without precise
locality, G. Forrest 18017 (A).

This variety is distinct in its broader leaves, which are conspicuously
and evenly white-tomentose throughout the lower leaf-surface. I have
not seen specimens ae ae and Malaysia. This variety may prove
to merit specific recogni

Forrest 13910, identified by W.W. Smith as A. championii var. mollis
Dunn, actually belongs here. Handel-Mazzetti (Symb. Sin. 7: 390, 391.
1931) refers this plant, as well as his own 9042 and 9046, to A. pilosula
(Finet & Gagnep.) Stapf, a new combination based on A. callosa Lindl.
var. pilosula Finet & Gagnep., with “A. championii var. mollis W. W.
Smith in Not. R. Bot. Gard. Edinbgh., XVII., 305 (1930), non Dunn,”
given in the synonymy. Handel-Mazzetti’s 9042 is available, and this is
clearly the same plant as Smith’s and should also be referred here.
Actinidia pilosula Stapf, based on A. callosa Lindl. var. pilosula Finet &
Gagnep. should be typified by the latter, and it is here considered as a
species distinct from A. callosa.

30. Actinidia latifolia (Gardn. & Champ.) Merr. in Jour. Roy. As. Soc.
Strait. Br. 86: 330. 1922.

Climbing shrubs to 7 m. high; branches reddish brown, with pale
oblong to lanceolate lenticels, rusty-tomentose on young branchlets;
pith solid, whitish, firm, becoming hollow in the center when old. Leaves
UP Ariaacous, broadly ovate or obovate or suborbicular to oblong-ovate
or oblong-obovate, about 5.5-11 em. long and 3-9 em. broad, acute to
acuminate at apex, cuneate, rounded, or truncate to reniform-cordate at
base, the margins subentire with remote minute callose teeth, the upper
surface green, glabrate, puberulous or densely puberulous, the lower
surface pale, covered sparsely or densely by a pale appressed stellate-
tomentum, the costa and veins inconspicuous to subconspicuous above,
distinct and elevated beneath, the secondary veins about 6 or 7 per side,
slightly arcuate-ascending, anastomosing, the veinlets with many dis-

50 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXXII

tinct cross-bars below, the reticulations hidden by the indumentum or
not; petioles 24 cm. long, tomentose. Inflorescences in axillary or lat-
eral, many-flowered, 3 or 4-branched cymes, to 10 cm. long, densely
rusty- -stellate- tomentose; peduncles 1.5-8 em. long, forked at top; pedi-
cels 0.5-1.5 em. long, elongating and enlarging in fruit; bracts minute,
linear. Flowers brownish yellow; sepals 5, ovate, about 4—5 mm. long
and 3-4 mm. broad, acute to obtuse, tomentose without: petals 5, about
6-8 cm. long and 3-4 mm. wide, sie to rounded, pubescent without;
stamens numerous, the filaments slender, 2-3 or sometimes to 5 mm.
long, the anthers linear- -oblong, about 1-2 mm. long, the connective pro-
jected and pointed at top, the base not divergent; ovary globose, about
2 mm. long, densely pilose, the styles 2-3 mm. long, slender. Fruit sub-
globose to ovoid, about 3-4 em. long and 2-3 em. across, brown, lenticel-
late, glabrous when mature or pubescent at the base and top on

A distinct species readily recognized by the more or less has tadeie
leaves and inflorescences. The leaves are generally broadly ovate, with
dense stellate tomentum beneath. The leaf-bases vary considerably i in
shape. The inflorescence is much more elaborate than any other species
of the genus; it is large, generally long-peduncled, much-branched, and
bearing many flowers. There is some variation in the size of the inflores-
cence. In spite of its wide range and variations in leaf-shape and size of
inflorescence, A. latifolia is a relatively uniform species. In addition to
the typical form, two varieties, apparently both of local and rare occur-
rence, may be ney

KEY TO THE VARIETIES
A. Tomentum dense, closed on the lower leaf-surface, the reticulations of
veinlets obscure.
B. Upper surface of leaves very sc alaeed puberulous or glabrate; lower

surface densely stellate-tomentose. .................. a. var. latifolia.

BB. Upper surface of leaves densely noah aieae lower surface more thickly
stellate-tomentose. .......00000000..0..0000000000.. var. mollis.

AA. Tomentum thin, open on the lower leaf-surface, the detidillations of
veinlets distinct. 2 0... c. var. indochinensis.

30a. Actinidia latifolia var. latifolia.
Actinidia latifolia tia & Champ.) Merr. in Jour. Roy. se Boe. Strait.
. ; ; Nakai in Bot. Mag. Tokyo 41: 521. 1927: Sasaki in
Trans. Nat. ie Soc. Formosa 19: 480. 1929; Hand.-Mazz. ata. Sin. 7:
391. 1931; Kanehira, era Trees rev, aa 450. 1936; Van Steenis,
Fl. Males. I. 4: 39, f 1. 1948.
Heptaca ? latifolia G ardn a Champ. in Hook. Jour. Bot. Kew Gard. Mise.
: . 1849.

Kadsura pubescens Miq. Fl. Ind. Bat. Suppl. 620. 1862; Kurz in Jour. As.
Soc. Beng. 45(2): 119. 1876.

Actinidia championi Benth. Fl. Hongk. 26. 1861; Matsum. «& Hayata in
Jour. Coll. Sci. Univ. Tokyo 22: 47. 1906 oo Pl. Formos.); Finet &
Gagnep. in Lécomte, Fl Gén. Indo-Chine 1: 28. 1907; Hayata in Jour.
Coll. Sei. Univ. Tokyo 30(1): 44. 1911 (Mat. FI. Formos.), Icon. Pl.

1952] LI, REVIEW OF THE GENUS ACTINIDIA 51

Formos. 1: 88. 1911; Dunn in Jour. Linn. Soc. Bot. 39: 407. 1911; Ridl.
Fl. Mal. Pen. 1: 206. 1922.

Actinidia miquelii King in Jour. As. Soc. Beng. 59(2): 196. 1896, nomen,
in Ann. Roy. Bot. Gard. Calcutta 5: 145, t. 176. 1896.

Actinidia Soe: Hayata, Icon. Pl. Formos. 9: 97. 1920.

Leaves broadly ovate to obovate or sub-orbicular, about 5.5-11 em.
long and 3-9 cm. wide, acuminate at apex, cuneate or rounded to reni-
form-cordate at base, the upper surface glabrate or puberulous, the
lower surface covered by a pale appressed stellate tomentum obliterat-
ing the veinlet-reticulations.

Southeastern and southern China (Taiwan, Kwangtung, Kwangsi,
Hunan, Fukien, Chekiang), Indo-China, Siam, and Malaysia (Perak,
Jehore, Sumatra), in thickets at altitudes of 350-1500 meters. Flowers
brownish yellow, April.

MALAY PENINSULA: Perak, No. coll. 5232 (A); Penang, Singapore Bot.
Gard. 32620 (A); Kedah, Singapore Bot. Gard. 35014 (A).

aes CHINA: Without precise locality, M. Poilane 8060 ae a onkin:

aokay, A. Pételot 8649 (A); Chapa, A. Pételot 8747 (A); : Bassae,
M. Poilane 15877 wee Annam: Dong-cho, Auang-tni, JZ. hea 10713 (A).

CHINA: Taiwan: Bankinsing, A. Henry 825 (A); Horisha, Nanto,
E. H. Wilson 11173 (A, US); Rengachi, T. Hayashi 21244 (A). Chekiang:
Ch’ung-an Hsien, Y. L. Keng 654 (A). Fukien: Ku-shan, J. B. Norton
1365 (US), 1366 (US), H. iL. Chung 7202 (A, NY); Ku-t’ien Hsien, H. H.
Chung 4019 (A). Kiangsi: P’ing-hsiang Hsien, Wang-Te-Hwi 205 (A);
Huang-shu-lin, between Ning-tu and Ch’ang-t’ling, Wang-Te-Hwi 370 (A);
Yung-hsin Hsien, H. H. Hu 803 (A); Ting-nan Hsien, H. H. Hu 1077 (A);
“Hong San,” G. L. Gressitt 1453 (A); Lung-nan Hsien, S. K. Lau 444 (A
US); Ch’ien-nan Hsien, S. K. Lau 3987 (A, US). Kweichow: Sanhao,
Y. Tsiang 6403 (NY). Kwangsi: N. Lo-chen, R. C. Ching 5709 (NY),
6003 (NY); W. Pai-se, R. C. Ching 7378 (NY); S. Nan-ning, Shih-wan-ta-
shan, R. C. Ching 8178 (NY), 8438 (NY); Shang-ssu Hsien, Shih-wan-tai-
shan, W. T. Tsang 22225 (A), 24543 (NY); Ch’iian Hsien, W. T. Tsang
27694 (US); Kuei-lin, W. T. Tsang 28284 (US). Hongkong: C. Wright
s. n. (GH, US, 1sorypes), Ford, s. n. (A, NY, US), Hongkong Herb. 8017
(A); Bae. Kew 53 (GH, tsorypre of A. champion Benth.). Kwangtung:
“Lung-tau Shan,’ Canton Christ. Coll. 12120 (US); Ta-pu Hsien, W. T.
Tsang 21074 (A, NY), 21715 (A, NY); Ying-te Hsien, T. M. Tswi 565 (NY,
US); Lo-ch’ang Hsien, W. T. Tsang 20843 (NY), C. L. Tso 20965 (NY).
Hainan: Hainan, C. Wang 33505 (NY); Ling-men, J. L. Gressitt 1164
(A); Lien-hua-lin, F. A McClure 8061 (A); Wu-chih-shan (Five Finger Mt.)
F.A. McClure 9487 (NY); Pao-ting Hsien, F. C. How 71995 (A), 72701 (A);
Yai Hsien (Yaichow), H. Y. Liang 62264 (A, NY, US), 62597 (NY); Hung-
mao-shan, Tsang & Fung 291 (A); ine ”N.K. Chun & C. L. Tso 43328
(A, NY), 43496 (A, NY, US); Ling-shui Hsien, H. Fung 20096 (A, NY, US);
Lin-kao Hsien, W. T. Tsang 239 (A, US), 470 (A, NY, US), 723 (A, US);
Ch’eng-mai Hsien, C. J. Lei 619 (NY, US), 907 (NY, US).

The reduction of A. miqueliz King was first made by Dunn, who listed
Heptaca ? latifolia Gardn. & Champ. in the synonymy of A. championt

52 JOURNAL OF THE ARNOLD ARBORETUM [VoL. XXXIII

Benth. This earlier name was subsequently adopted by Merrill. A
photograph of the type specimen, Champion s. n., from Hongkong, is in
the herbarium of the Arnold Arboretum. The reduction of A. gnaphalo-
carpa Hayata was made by Sasaki and followed by Kanehira and
others. A photograph of Hayata’s type (in A, US) and collections of
Taiwan plants indicate that this reduction is necessary.

30b. Actinidia latifolia var. mollis (Dunn) Hand.-Mazz. Symb. Sin. 7:
391. 1931

Actinidia championi Benth. var. mollis Dunn in Jour. Linn. Soc. Bot. 39:
407. 1911.

Leaves ovate to oblong-ovate or oblong-obovate, about 10-12 ecm.
long and 5.5-7 em. wide, acute to acuminate at apex, broadly acute to
rounded or truncate at base, the upper surface more or less densely
puberulous, the lower surface very densely and thickly stellate-tomen-
tose, the tomentum obliterating the veinlet-reticulations.

Southwestern China (in southern Yunnan only), in mountain forests
at altitudes of 1400-1500 meters. Flowers yellow, June.

CHINA: Yunnan: No precise locality, J. C. Liw & C. Wang 83077
(A); Ssu-mao Hsien, A. Henry 12041 (A NY, ISOTYPES); P’ing-pien Hsien,
H. T. Tsai 55411 (A), 60153 (A), 60193 (A), 60310 (A), 61405 (A), 62971
(A).

30c. Actinidia latifolia var. indochinensis (Li) comb. nov.

Actinidia indochinensis Li in Jour. Arnold Arb. 24: 366. 1943.

Leaves ovate to elliptic-ovate, 7-21.5 em. long, 4.5-6.8 em. wide, acute
to shortly acuminate at apex, acute to broadly acutish at base, the upper
surface glabrous, the lower surface scattered stellate-tomentose, the
veinlet-reticulations distinct.

Indo-China, in Tonkin only. Flowers May—June.

INDO-CHINA: Tonkin: Dam-ha, W. 7. Tsang 29907 (A, TyPE).

31. Actinidia eriantha Benth. in Jour. Linn. Soc. Bot. 5: 55. 1861;
Dunn in Jour. Linn. Soc. Bot. 39: 408. 1911; Chun in Sunyatsenia 1:
271, pl. 37. 1934; Rehder in Jour. Arnold Arb. 18: 222. 1937.
Actinidia davidii Franch. in Nouv. Arch. Mus. Paris sér. 2. 5: 57. 1884;
Dunn in loe. cit. 39: 408. 1911; Hand.-Mazz. Symb. Sin. 7: 391. 1931;
Chun in op. cit. 2: 63. 1934.

Actinidia lanata Hemsl in Ann. Bot. 9: 146. 1895; Dunn in op. cit. 38:
355. 1908, 39: 409. 1911; Dunn & Tutcher in Kew Bull. Add. Ser. 10:
44. 1912; Rehder in Jour. Arnold Arb. 15: 97. 1934.

Climbing shrubs to 10 m.; branches grayish to dark grayish, glabrate
or tomentose, with conspicuous oblong concolored lenticels, the young
branches densely white-villose; pith white, medium-sized, lamellate.
Leaves chartaceous, broadly ovate to ovate or oblong-ovate, about 8-16
em. long and 5.5-11 em. broad, obtuse to acute or shortly acuminate at
apex, rounded, truncate to subecordate at base, the margins subentire
and with minute scattered callose teeth or minutely mucronulate-serru-

1952] LI, REVIEW OF THE GENUS ACTINIDIA 53

late, green and hispid-puberulous above to nearly glabrous, usually
pilose along the costa, pale and densely white-stellate-tomentose be-
neath, the costa and veins subconspicous to inconspicuous above, raised
and distinct beneath, the secondary veins about 7 or 8 per side, arcu-
ate-ascending, anastomosing, the reticulations with many cross-bars,
raised and distinct beneath; petioles short, stout, 1.5-2.5 em. long,
densely and thickly white-villose. Inflorescences in short axillary cymes
of 1-4 flowers, densely and thickly white-villose; peduncles scarcely to
1.5 em. long; pedicels very short, 3-5 mm. long. Flowers lanrge, rose
pink; sepals broadly ovate, about 7 mm. long and 5 mm. broad, obtuse,
densely villose without; petals 5, ovate, about 1.5 em. long and 1.1 em.
broad, rounded, the margins often slightly fringed; stamens very nu-
merous, the filaments slender, subequal, 5-7 mm. long, the anthers yel-
low, oblong, about 1 mm. long, pointed at tip, rounded at base; ovary
globose, densely villose, the styles 3-4 mm. long. Fruit ovoid, about
3.5 cm. long and 3 cm. across, densely and thickly pale villose.

Southeastern China (Chekiang, Kiangsi, southern Hunan, Kweichow,
Fukien, Kwangsi, Kwangtung), in thickets, at altitudes of 250-1000
meters. Flowers white or pinkish, June.

CHINA: Chekiang: Yung-chia Hsien (Wenchow), R. C. Ching 1872
(A, US), Y. Tsiang 1401 (A); Lung-ch’iian Hsien, R. C. Ching 2479 (A, US),
H. H. Hu 463 (A). Hunan: Ch’ang-ning Hsien, C. S. Fan & Y. ¥. Li 212
(A). Kweichow: Kuei-ting, Y. Tsiang 5483 (NY). Fukien: “Shiu-kia,”
Hongkong Herb. 2397 (A); Nan-p’ing (Yenping), H. H. Chung 3283 (A),
3365 (A), Chou Kuang Hou 8979 (A); Ku-shan, J. B. Norton 136 (US),
H. H. Chung 6641 (A). Kwangsi: N. Lo-chen, R. C. Ching 6132 (A, US);
Kwei-lin Hsien, W. T. Tsang 28428 (US). Kwangtung: Lien-p’ing Hsien,
R. Mell 604. (A); Lo-ch’ang Hsien, C. L. Tso 20538 (A, NY),20927 (NY);
Chia-ying Hsien (Mei). W. T. Tsang 21384 (A, NY); Yuan Hsien, S. K. Lau
2272 (A); “Lung-tau Mt.”, Canton Christ Coll. 12297 (NY, US).

Dunn recognized A. eriantha Benth., A. davidii Franch., and A. lan-
ata Hemsl. as three distinct species. He apparently did not see Ben-
tham’s or Franchet’s types. He misinterpreted A. davidii as a species
without stellate tomentum, and accordingly this species was misplaced
in his key. Bentham’s species is based on “Lindley, from S. China” and
Franchet’s on a collection made by David, from “Kiang-si oriental.
Automme 1873.” A photograph of Lindley 1836 is reproduced in Sun-
yatsenia 1: 273, t. 37. 1934, by Chun, who considers A. eriantha Benth,
and A. lanata Hemsl. as distinct species. He says that A. eriantha “is
easily distinguished from A. lanata Hemsl. by the densely whitish
woolly indumentum on the under surface of the leaves, inflorescences,
and fruit, and by the filiform, not linear, filaments of the stamens.” He
considers A. lanata as having a fulvous or tawny indumentum.

Rehder, however, in Jour. Arnold Arb. 18: 222. 1937, states that none
of the distinguishing characters given by Chun and others seem to be
dependable and considers the two names to be actually synonymous. I
am tentatively following Rehder’s opinion. Hemsley’s type is from

54 JOURNAL OF THE ARNOLD ARBORETUM [voL. XXXII

“China: Kwangtung along the Northwest river, Mr. Ford’s native col-
lector; 228 of 1890 collection.” This is a young fruiting specimens,
which I have not seen. Hemsley describes the indumentum as “fer-
rugineo-lanatis vel tomentosis,” and compares it with A. fulvicoma
Hance. It is possible that his species is referable to the latter, conceiv-
ably to var. hirsuta.

From the original description, it seems certain that A. davidii Franch.
also belongs to A. ertantha Benth., as noted earlier by Handel-Mazzetti
(Symb. Sin. 7: 391. 1931) and others.

32. Actinidia chinensis Planchon in London Jour. Bot. 6: 303. 1847.

Climbing shrubs to 8 m.; branches reddish brown, with paler oblong
lenticels, the young branchlets brownish-pubescent or setose; pith large,
lamellate, whitish or yellowish. Leaves thin- or thick-chartaceous, those
of the sterile branches broadly ovate to elliptic, very shortly acuminate
to cuspidate at apex, those of flowering branches suborbicular, shortly
cuspidate, rounded or truncate at apex, rounded to more or less cordate
at base, 6-17 em. long, 6-15 em. broad, the margins minutely denticu-
late, the teeth produced by tips of veinlets, the upper surface dark green,
more or less puberulous, more densely so on the costa and nerves, or
densely scabrid-hispid throughout, the lower surface very pale, densely
whitish-stellate-tomentose, the costa and veins subconspicuous above,
raised and distinct beneath, the secondary nerves about 5-8 per side,
strongly patent, straight or arcuate-ascending, anastomosing, the
branchlets ending in the marginal teeth, the veinlets in parallel cross-
bars, more or less conspicous; petioles 3.5-7.5 cm. long, more or less
densely pubescent. Inflorescences in few-flowered cymes, from axils of
fallen leaves, pubescent; peduncles about 1.5 cm. long; pedicels 1-2 cm.
long; bracts minute, linear. Flowers orange-yellow, the staminate
slightly smaller; sepals 5, sometimes 3 or 4, ovate-oblong, about 8-10
mm. long and 6-8 mm. wide, obtuse to acute at apex, brownish-tomen-
tose without; petals 5, broadly obovate, shortly clawed, rounded at top,
about 1.4-1.5 em. long, 1-1.2 cm. broad; stamens very numerous, the
filaments filiform, unequal, about 5-10 mm. long, the anthers oblong,
1.5 mm. long, acute to obtuse at apex, slightly sagittate at base; ovary
subglobose, about 6-7 mm. across, densely brownish-villose, the styles
linear, about 5-6 mm. long. Fruit subglobose to ellipsoid, about 3 cm.
across, densely brownish-hirsute all over; seeds oblong-ellipsoid, 2-3
mm. long, foveolate-reticulate.

This is the common Yang-tao of China, widespread in most parts of
the country but especially common along the Yangtze valley. The leaves
vary from emarginate to truncate to those on young shoots sometimes
shortly acutish or cuspidate. The plants from the island of Taiwan have
the leaves relatively longer and much narrower, always acute to shortly
acuminate, and also relatively thinner when compared with mainland

1952] LI, REVIEW OF THE GENUS ACTINIDIA 55

plants. he upper surfaces of the leaves are more or less densely long
hispid-setose. This insular form, only sterile material being available, is
herein recognized as a variety.

KeEY TO THE VARIETIES
A. Leaves thicker, generally orbicular, truncate to emarginate at apex,
hispid mostly along the veins only; stems and petioles soft- pubescent
when young, glabrous when mature. (Mainland China)....... ......
Peed roe Mae aes Belk, he bce 54-8 a a, Ge a Ri: r. chinensis.
AA. Leaves thinner, generally ovate, acute to shortly ee at apex,
scabrid-hispid above: stems and petioles densely hispid- setose. (Taiwan).
. var. setosa.

32a. Actinidia chinensis var. chinensis.

Actinidia chinensis Se in London Jour. Bot. 6: 303. 1837; Oliv. in
Hook. Icon. Pl. 15: t. 1593. 1887; Dunn in Jour. Linn. Soc. Bot. 39: 408.
1911; Sprouse in Bot. Mag. 140: tt. 8538. 1914; Rehder in Sarg. Pl. Wils.
2: 385. 1915; Merr. & Chun in Sunyatsenia 1: 70. 1930; Hand.-Mazz.
Symb. Sin. 7: 391. 1931

Young branchlets and petioles setose, brownish-pubescent; leaves
chartaceous, mostly suborbicular, about 6-12 em. long and broad,
emarginate to shortly cuspidate at apex, rounded to more or less cordate
at base, the upper surface more or less puberulous, more densely so on
the costa and nerves, the lower surface densely whitish-stellate-
tomentose.

Widely distributed in western, central, eastern, and southern China,
in thickets and forests, on slopes or in ravines, at altitudes of 200-2300
meters. Flowers whitish, changing to buff yellow.

CHINA: Shensi: Tai-pei-shan, W. Purdom 657 (A, US). Honan:
ee J. Hers 422 (A), 464 (A): Chi-kung-shan, A. N. Steward 9772

(A, US). Kiangsu: I-hsing Hsien, Ching & Tso 523 (A). Anhwei:
ce hua-shan, R. C. Ching 2639 (A, ae 2678 (A); Huang-shan, R. C.
Ching 2910 (A), W. C. Cheng 3977 (US). Hupeh: Western Hupeh, A.

Henry 1166 (US), 2076 (GH), 5834 ae US), 5834A (GH, NY), 5834B
(US), EZ. H. Wilson 185 (A, NY, US), 720 (NY); I-ch’ang, E. H. Wilson
8347 p. p. (A); Ch’ang-yang Heed Ei. H. Wilson 993 (A, NY, US); “Lan-
tan,” P. C. Sylvestri 1467 (A); “Jan- lan-sean,” P. C. Cesie 6157 (A);
Wu- tu-ho, W. Y. Chun 3638 (A), 3980 (A, US); Pa-tung Hsien, H. C. Chow
289 (A, NY), 750 (A, NY). Szechuan: S. Wu-shan, A. Aen) s. n. (A);
Wen-ch’iian Hsien, £. H. Wilson 347 p. p. (A, GH); Mo-tien-ling, F.T. Wang
22445 (A); Nan-ch’iian Hsien, C. Bock & K. A. v. Rosthorn 1997 (A), W. P.
Fang 1084 (A), 1096 (A); O-mei-shan, W. P. Fang 2613 (A), C. L. Sun
2042 (A), 2094 (A), 2136 (A), T. C. Lee 2746 (A), 2990 (A), H. C. Chow
7800 (A), Sun & Chang 784 (A); oe -pien Hsien, F. T. Waa 22859 (A);
O-pien Hsien, Y. S. Liu 1355 (A). ang: ours -ting, W. C. Cheng 1728
(NY, US), 1729 (A); Ning-yiian ee C. Schneider 993 (A), Tien-ch’iian
sien, T. C. Tat 5135 (A). Chekiang: Mo-kan-shan, Cheo & Wilson
12709 (GH, NY), F. N. Meyer 1568 (A); Ch’ang-hua Hsien, F. N. Meyer
1537 (A); T’ien-mu Shan, R. C. Ching 5090 (A); S. Yin Hsien (Ningpo),

56 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXXIII

D. Macregor s. n. (A); T’ien-t’ai Shan, R. C. Ching 1438 (A, US); Lung-
ch’tian Hsien, H. H. Hu 480 (A); Chi ing-yuan, Y. L. Keng 360 (A).
Kiangsi: Lu- shan, T. L. Bullock 120D (US), A. N. Steward 2646 (A, US),
H. H. Chung & C. S. Sun 664 (A, way Pigs Y. Tsiang 10085 (NY);

Ch’ien-shan Hsien, C. S. Fan & Y. Y. Li 92 (A). Hunan: Hsing-hua cree
H. Handel-Mazzetti 548 (A); Heng- de H. Handel-Mazzetti 729 (A).
Kweichow: Fan-ching-shan, Steward, Chiao & Cheo 430 (A, NY, US);
T’ung-tzu Hsien, Y. Tsiang 5178 (NY); Tu-yiin Hsien, Y. Tsvang 5730 (NY);
Ta-ting Hsien, Y. Tsiang 8949 (NY). Yunnan: Yung-shan Hsien, H. T.
Tsai 51180 (A). Kwangtung: Lo-ch’ang Hsien, C. L. Tso 20704 (NY).

32b. Actinidia chinensis var. setosa var. nov.

Actinidia chinensis sensu Nemoto, Fl. Jap. Suppl. 474. 1936; Suzuki in
Masamune, Short Fl. Formos. 137. 1936; Kanehira, Formosan Trees.
rev. ed. 449, t. 406. 1936; non Planch.

A typo speciei caulibus petiolisque dense ferrugineis hispido-setosis
differt; foliis tenuibus, late ovatis, 12-17 em. longis, 10-15 em. latis,
apice acutis vel breviter acuminatis, superne plus minusve dense
scabride hispidulis, inferne stellato-tomentosis.

China, in Taiwan only, at altitudes of 1800-2600 meters.

CHINA: Taiwan: Mt Arisan, EF. H. Wilson 10802 (US, Type; A, Iso-
TYPE), Oct. 18, 1918, R. Kanehira 2994 (NY).

33. Actinidia fulvicoma Hance in Jour. Bot. 23: 321. 1885.

Climbing shrubs to 10 m.; branches reddish brown to grayish, with in-
conspicuous lenticels, the young branchlets densely brownish-tomen-
tose, the older branches glabrous or nearly so; pith whitish, lamellate.
Leaves membranaceous or chartaceous to coriaceous, ovate to oblong
to oblong-ovate, about 7-15 em. long, 3-9 em. broad, acuminate to
long-acuminate at apex, rounded to cordate at base, the margins setose-
denticulate, the upper surface green, densely to sparsely hirsute to
scabrid-hirsute especially along the costa, the lower surface very pale
densely yellowish stellate-tomentose, the costa and veins subconspicu-
ous above, raised and distinct beneath, the secondary veins about 8
per side, patent-ascending, anastomosing, the veinlets reticulate, with
numerous cross-bars, obliterated by the indumentum beneath; petioles
1.5-4 em. long, brownish-tomentose. Inflorescences in short few-
flowered axillary cymes or the flowers solitary, densely brownish vil-
lose-pubescent; peduncles to 1 cm. long; pedicels 1 cm. or less long;
bracts minute, linear. Flowers white; sepals 5, ovate, about 6-7 mm.
long and 3-4 mm. broad, acute at apex, densely villose without; petals
5, obovate, 7-10 mm. long, 5-7 mm. wide; stamens numerous, the fila-
ments about 3-4 mm. long, the anthers yellow, triangular-ovoid, about
1 mm. long, the apex acute, the base sagittate; ovary subglobose, about

mm. across, densely villose, the styles about 3 mm. long. Fruit
oblong, to 2.7 em. long and 2 em. across, brown, with pale scattered
lenticels, glabrous or sparsely pubescent when mature, the persistent
calyx erect, not reflexed.

1952] LI, REVIEW OF THE GENUS ACTINIDIA 57

Actinidia fulvicoma Hance is a close relative of A. eriantha Benth.
and has the same general range in southeastern China. It can, however,
be readily distinguished from the latter by its relatively narrower and
longer leaves which are pubescent above, its more hirsute stems and
petioles, its yellowish or brownish indumentum, and its nearly glabrous
mature fruit. In A. ertantha the young stems and petioles are softly
and shortly downy, the indumentum is whitish, and the mature fruit is
densely hairy.

The species is composed of three varieties which have more or less
distinct but contiguous ranges. The typical variety is found in Cheki-
ang, southern Kiangsi, southwestern Hunan, eastern Kweichow, and
northern Kwangtung. Variety Tel lik is found only in Kwane-
tung, and var. mrsuta in Kweichow

KEY TO THE VARIETIES

A. Leaves chartaceous to thinly coriaceous; stems oe 1:3)
pie ee eee SS. boo veda pa eee var. fulvicoma.
AA. Leaves coriaceous; stems with short brownish hirsute hae Sey eee
eR 5. rok en lb ye Rite Oke b. var. pachyphylla.
AAA. ra more or less membranaceous; stems densely sas brownish
Fer TS sex Gos ned awh -b 4 . var. hirsuta.

33a. Actinidia fulvicoma var. fulvicoma.

slit Aegis Hance in Jour. Bot. 23: 321. 1885; Dunn in Jour. Linn.

Soc. Bot. 39: 409. 1911; Hand.-Mazz. Symb. Sin. 7: 391. 1931.

ae ae fad petioles soft-brownish-tomentose; leaves
chartaceous to thinly coriaceous, ovate to oblong-ovate, about 7-14
em. long and 3.5-7 cm. broad, long-acuminate at apex, rounded to
cordate at base, the margins setose-denticulate, the upper surface hir-
sute to scabrid-hirsute especially along the costa, the lower surface
yellowish stellate-tomentose.

Southeastern to southern China (Chekiang, southern Kiangsi, south-
ern Hunan, Kwangsi, Kwangtung), on slopes or in valleys, in shade
or open, at altitudes of 500-800 meters. Flowers white, June.

CHINA: Chekiang: Without eae locality, Barchet 65 (US).
Kiangsi: Ch’ung-i Hsien, H. H. Hu 931 (A); Ta-yii Hsien, H. H. Hu 952
(A); Ch’ien-nan Hsien, S. K. Lau 4050 (US). Hunan: Sing-ning Hsien,
C.S8. Fan & Y.Y. Li 680 (A); Heng-shan, H. cae 12182 (A).
Kweichow: San-ho Hsien, Y. Tsiang 6880 (NY). Kwangsi: Hsing-
yeh Hsien, R. C. Ching 6880 (NY), 7177 (NY); Lo-chen, Hsien, R. C. Ching
5583 (NY), 5678 ae 6152 (NY). Kwangtung: Lo- chang Hsien, C.
L. Tso 20841 (N.Y.)

33b. Actinidia fulvicoma var. pachyphylla (Dunn) comb. nov.
Actinidia pachyphylla Dunn in Jour. Linn. Soc. Bot. 39: 409. 1911.
Young branchlets and petioles brownish short-hirsute; leaves charta-

ceous to coriaceous, oblong to oblong-ovate, about 10-15 cm. long and

3-7 cm. broad, acuminate at apex, rounded to cordate at base, the

58 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXXII

margins setose-denticulate, the upper surface hirsute to scabrid-hirsute
especially along the costa, the lower surface stellate-tomentose.

Southern China, in Kwangtung, in thickets at altitudes of 500-1000
meters. Flowers white, May—June.

CHINA: Kwangtung: Lo-ch’ang Hsien, C. L. Tso 20713 (NY), W. T.
Tsang 20987 (NY); Lien-p’ing, FR. Mell 603 (A); Yang-shan Hsien, 7. M.
Tsu 748 (NY); Pei-shan to Tai-ping, W. Y. Chun 5662 (A); Lung-men
Hsien, W. 7. Tsang 20519 (NY, US).

The type of A. pachphylla Dunn, “Swatow, Fung Wan Shan of
Phoenix Mountain, Hong Kong Herb.”, has not been seen. Dunn’s
species can be distinguished from A. fulvicoma only in the thicker
leaves. The above cited specimens seem to be referable to this concept,
which I recognized as a variety of A. fulvicoma. The leaves are gen-
erally larger and narrower, varying from chartaceous to thickly coriace-
ous, sometimes even in a single collection. The thickness of the leaves
may be influenced to a certain degree by variations in habitat.

33c. Actinidia fulvicoma var. hirsuta Finet & Gagnep. in Bull. Soe.
Bot. France 52: Mem. 4: 18. 1907 (Contr. Fl. As. Or.) ; Hand.-
Mazz. Symb. Sin. 7: 391. 1931.

Young stems and petioles densely brownish long-hirsute; leaves
membranaceous to thinly chartaceous, ovate to oblong-ovate, about
7-14 cm. long and 4-9 em. broad, long-acuminate at apex, rounded to
cordate at base, the margins setose-denticulate, the upper surface
densely setose, the lower surface densely stellate-tomentose.

Southwestern China, in Kweichow and western Kwangsi, at altitudes
of 500-1000 meters. Flower white, June—July.

CHINA: Kweichow: R. P. Bodinier 2427 (NY, tsorype) : Kuel-ting,
Y. Tsiang 5586 (NY). H. Handel-Mazzetti 10571 (A); Tu-yun Hsien, Y
Tsiang 5727 (NY). Kwangsi: Ling-yiin Hsien, A. N. Steward & C. C.
Cheo 656 (A, NY).

Dunn, who examined Bodinier 2427, the type, did not recognize this
variety, as he considered the specimen “apparently from secondary
flowering branch and does not differ from corresponding parts of Hance’s
type.” Handel-Mazzetti, however, reinstated the variety. From the
above cited specimens, it seems that in Kweichow and western Kwangsi
the species has thinner and more setose leaves, as well as more densely
hirsute stems and petioles, indicating the existence of a geographical
variety.

34. Actinidia lanceolata Dunn in Jour. Linn. Soc. Bot. 38: 356. 1908,
39: 408. 1911.

Climbing shrubs to 20 m.; branches reddish brown, glabrous, the
young branchlets densely ferrugineous-puberulous; pith small, brown,
lamellate. Leaves chartaceous, lanceolate to ovate-lanceolate, about
4—7 cm.:long and 2-3 cm. broad, acuminate at apex, cuneate at base,
the margins minutely setose-denticulate, the upper surface dark,

1952] LI, REVIEW OF THE GENUS ACTINIDIA 59

sparsely and minutely puberulous to nearly glabrous, the lower sur-
face glaucescent, appressed with white stellate-tomentum, the costa
and veins scarcely conspicuous above, raised and distinct and covered
with brownish pubescence beneath, the secondary veins about 4 or 5
per side, patent-ascending, anastomosing, the veins or their branches
ending in the marginal teeth, the veinlets in parallel cross-bars, in-
conspicuous; petioles 1-1.5 cm. long, brownish pubescent. Inflores-
cence in axillary cymes, 3—6-flowered, ferrugineous-puberulous; pe-
duncles 3-6 mm. long; pedicels 2-4 mm. long; bracts linear, minute.
Flowers greenish; sepals 5, ovate, about 3-4 mm. long and 1.5 mm.
broad, rounded at apex, ferrugineous-puberulous without; petals 5,
oblong, about 4-5 mm. long and 1.5-2 mm. broad, slightly larger than
the sepals, acute or rounded at apex; stamens numerous, the filaments
2-3 mm. long, the anthers yellow, oblong, small, about 1 mm. long,
rounded at both ends; ovary subglobose, about 1.5 mm. long, densely
ferrugineous-tomentose, the styles about 1.5 mm. long. Fruit very
small, ovoid, about 10 mm. long and 7 mm. across, brown, glabrous,
with pale roundish lenticels, the persistent sepals reflexed.

Southeastern China (southern Chekiang, southern Kiangsi, Fukien,
and northern Kwangtung), on mountain slopes or along river banks, at
altitudes of 200-600 meters. Flowers greenish, May—June.

CHINA: Chekiang: Hsien-chii Hsien, Y. L. Keng 467 (A), R. C. Ching
1595 (A, NY, US), 1714 (A, NY, US); P’ing-yang, R. C. Ching 2074 (A,
NY, US); Chiang-shan, R. C. Ching 2582 (A, NY, US). Kiangsi: Yung-
hsiu, Y. Tsiang 10685 (NY). Fukien: Nan-p’ing (Yenping), Hongkong
Herb. 2399 (A, IsoTyPE). Kwangtung: Ta-pu Usien, W. T. Tsang 21222
(A).

This species was described by Dunn from Fukien, based on a single
collection, Hongkong Herb. 2399, collected on his expedition to central
and western Fukien in 1905. A duplicate of this number is available.
The known range of the species now extends to the neighboring
provinces of Chekiang, Kiangsi, and Kwangtung.

This is a most distinct species, strongly characterized by its small
leaves with appressed whitish stellate hairs on the lower surface, the
small greenish flowers, and the very small fruits. It little suggests
relationships with other species. The stellate-tomentum on the under
surface of the leaves is so fine and appressed that Dunn failed to
note and describe it in his original description. Later, in his revision of
the genus, however, he properly keyed it with other stellate-tomentose
species. It is very distinct in this group of species in the cuneate leaf-
bases, as well as in the characters mentioned.

IMPERFECTLY KNOWN SPECIES

35. Actinidia kiusiana Koidzumi, Pl.’Nov. Amami-Oshim. 9. 1928,
in Bot. Mag. Tokyo 43: (421-422). 1929, in Acta Phytotax. Geob.
9: 98, in clavi. 1940.

60 JOURNAL OF THE ARNOLD ARBORETUM [VoL. XXXII

A species of the Liukiu Islands and Kiusiu, Japan, of which no
specimens are available to me. As the original description, which is
very short, was published in an obscure work, it is reproduced below:

“Actinidia (Vestitae) kiusiana Koidz, nov. sp.

This plant seems to be near to Actinidia strigosa Hook. of Himalaya,
but readily distinguished by the leaves cordate at the base and hispid even
in the upper surface.

Arbor ascendens ramis molliter puberulis strigoso-hispidis. Folia mem-
branacea oblongo-ovata 8, 5-13 cm. longa, 4-6 em. lata, acuta, basi cordata,
mucronato-serrata, utraque latere 7-9-penninervia, laxe hispida in utraque
pagina, petiolis 2-6 em longis hispidis.

Nom. Jap. Nagaba-shirakuchizulu.

Hab. The Loochoo archipelago: insula Tanegashima.

Ranges. _Kiusiu: prov. Hiuga, Minaminakagun, Ichiimura; Prov. Buzen,

Usagun, mount. Gongenzen.

Type specimens: in Herb. Bot. Inst. Kyoto Imp. Univ. Japan.”

As the description is based on vegetative parts only, it is not pos-
sible to ascertain the real identity of the species. I have not seen any
specimens of this genus from the Liukius and Japan that have strigose
stems. The nearest species of this group is A. arisanensis Hayata of
Taiwan, which is a very variable plant. From the brief original de-
scription, A. kiwsiana indeed appears close to A. arisanensis and the
two may prove to be conspecific.

36. Actinidia longicauda F. Chun in Sunyatsenia 7: 14. 1948.

The type, C. S. Chen 81944, collected from “Kwangsi; Chuen Hsien,
Shan-Chuan Hsiang, Ku-Kien An,” has not been seen by me. The
species known only from the original collection. The flowers are red-
dish, and they bloom in June.

Chun compares this species with A. kolomikta and A. venosa, saying
that it “Differs from A. kolomikta Maxim. in white not brownish pith,
long acuminate and elliptic to elliptic-obovate, not broadly ovate-
cordate abruptly caudate leaves and in smaller red flowers. The color
of the flowers and the shape of the leaves also serve to distinguish it
from A. venosa Rehd. which has, in addition a tomentose inflorescence
and an ovoid subglobose ovary.”

Since the fruit is not known, I cannot place it in the proper section
from the description. The ovary, which is described as cylindric, the
styles not being mentioned, may be the rudimentary ovary of the
staminate flowers. The actual shape of the ovary, which is an important
diagnostic character among the relatives of A. kolomikta, remains to
be seen. See also A. kwangsiensis for further comments.

1952] LI, REVIEW OF THE GENUS ACTINIDIA 61

HYBRID

Actinidia fairchildii Rehder in Jour. Arnold Arb. 20: 421. 1939.
Actinidia arguta X chinensis Fairchild in Jour. Hered. 18: 58, f. 7. 1927.
This is the only reported hybrid in the genus. It is a deliberate

cross between a staminate plant of A. arguta and a pistillate plant of

A. chinensis, the resulting hybrid showing characters intermediate

between the two parent species. The hybrid was raised in Washington,

Di.©,

DEPARTMENT OF Botany,
U.S. NationaL Museu
SMITHSONIAN an en.
WASHINGTON, D.

62 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXXIII

STUDIES IN THE BORAGINACEAE, XXII
NOTEWORTHY SPECIES, CHIEFLY ASIAN AND
SOUTH AMERICAN

Ivan M. JOHNSTON

Cordia varronifolia. sp. nov.

Frutex 2 m. alta; ramulis plus minusve villulosis in sicco nigrescenti-
bus; foliis elliptico- oblongis vel ovato-oblongis 4-9 em. longis 2-4 em.
latis, apice obtusiusculis, basi obtusis rotundisve 2-5 mm. longe petio-
latis, margine basim versus excepta saepe crenato-dentatis, supra
abundanter minuteque areolato-bullatis setis mollibus brevibus basi
bulbosis obsitis, subtus elevate reticulato-venosis griseo-villulosis;
cymis paucifloris terminalibus vel rare extra-axillaribus initio glomera-
tis, posterius in cincinnos solitarios vel rare geminatos 1-2 em. longos
excurrentibus; calyce sub anthesi 15-20 mm. longo griseo-villoso 10-
costato, tubo 10-12 mm. longo infra medium crassiore (4-8 mm. dia-
metro) apice 3-4 mm. diametro basi rotundato, lobis subulatis flexuosis
5-9 mm. longis in alabastro liberis, sinibus obtusis latis; calyce fructi-
fero ad 28 mm. longo, tubo 17 mm. longo 8-9 mm. crasso; corolla alba
marcescenti persistente infundibuliformi 3-3.5 em. longa, limbo 2.5-3.5
em. diametro, lobis adscendentibus semi-circularibus 9-11 mm. latis
5-7 mm. longis apice emarginatis, sinibus angustis valde acutis, tubo
2--3 mm. crasso cylindrico ca. 10 mm. longo tubum calycis subaequilongo
intus secus bases lineatas decurrentes filamentorum villuloso alibi
glabro, fauce ampliata apice ad 12 mm. diametro; filamentis 10-12 mm.
longis ca. 10 mm. supra basin tubi corollae orientibus; antheris oblongis
ca. 2 mm. longis; ovario glabro 2.5 mm. longo infra medium 1 mm.
crasso deinde sursum gradatim attenuato, 4-ovulato 4-loculato imam
ad basin nectario annulato inconspicuo cireumdato; stylo gracillimo
villuloso 15-25 mm. longo 12-20 mm. supra basin fureato, lobis 4
stigmatiferis angustissime oblanceolatis 2-3 mm. longis; fruetu exsicco
ovoideo calyce persistenti et tubo corollae persistentis investo supra
basin 4-5 mm. crasso deinde sursum angustato (conico, apice rostulato)
basi rotundato.

PERU: Pion valley of the Maranon, dept. Cajamarca, prov. Cutervo, 1300-
1400 m. alt., shrub 3 m. tall, fl. white, June 1915, A. Weberbauer 7138 (TyYPr,
Gray Herb.)

This shrub of northern Peru has the sacra ele, frequently den-
tate leaves suggestive of members of Cordia § Varronia. Its elongate
persisting ribbed calyx and the large persisting marcescent funnelform
corollas suggest those of members of the section Gerascanthus. Actu-
ally, however, this Peruvian plant has close relations only with C.

y

parvifolia DC. (C. Greggit Torr.) and C. elaeagnoides DC. of Mexico

1952] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXII 63

and so constitutes a third member of the section Rhabdocalyx. In size,
form and organization of the corolla, calyx and fruit it is most similar
to C. parvifolia, a species of the deserts of western and northern Mexico.
Like that species, furthermore, its flowers appear to be heterostylic.
Its fruits have all the distinctive characteristics of the section Rhabdo-
calyx. Indeed, the fruit is almost indistinguishable from that of C.
parvifolia and C. elaeagnoides, being conic-ovoid, dry, and nut-like,
having similar proportions and dimensions and developing completely
ensheathed by the persistent calyx. The endocarp is thin and her-
baceous in texture. It is not fleshy nor colored. The fruit is not a
drupe. Its bony endocarp has walls about a millimeter thick and is
4-celled and apparently 4-seeded at maturity. This is a fruit very
different from that developed by most Cordias and especially by mem-
bers of the sections Gerascanthus and Varronia, ef. Johnston, Jour.
Arnold Arb. 30: 85 (1949) and 31: 179 (1950).

Cordia iguaguana, sp. nov.

Arbor 8-20 m. alta; ramulis juventate tenuiter inconspicueque fulvo-
tomentulosis, maturitate glabratis; foliis maturitate late lanceolatis
10-15 cm. longis 4—5 em. latis glabris vel secus costa et petiolo pilulis
minutis adpressis sparsissime donatis, apice acutis vel paullo attenuatis,
basi obtusis vel rotundis 12-22 mm. longe petiolatis, supra nervis et
nervulis abundantibus laeviter impressis ornatis, subtus pallidioribus
nerviis (utrinque costae 5-7) et costa prominulis donatis; inflorescentia
apice ramulorum hornotinorum foliis juvenilibus gestorum prodita mul-
tiflora paniculata vel corymbosa 12-15 mm. diametro; calyce clavato-
cylindraceo ca. 11 mm. longo (apicem versus ad 3 mm. crasso) promi-
nenter 10-costato, extus inconspicue puberulento saepe pilulis sparsis
ad 0.2 mm. longis adpressis sparse donatis, costis saepe evidenter longi-
tudinaliterque sulcatis, lobis 3-5 saepe inaequalibus obtusis triangulari-
bus ca. 1 mm. longis, sinibus obtusis; corolla alba 23 mm. longa extus
glaberrima, limbo ca. 25 mm. diametro, lobis 6-8 mm. longis 8-10 mm.
latis supra basin latioribus rotundis vel apice plus minusve truncatis
non rariter emarginatis, sinibus angustis acutis, tubo 8-8.5 mm. longo
1—1.5 mm. crasso intus infra medium tomentuloso, faucibus 12-15 mm.
diametro ca. 5 mm. profundis, filamentis glabris ore tubi (8-9 supra
basin tubi) affixis inaequalibus 1-3 mm. longis, antheris 1.5-3 mm.
longis inaequalibus; pistillo (ovario cum stylo) glabro 13-14 mm. longo
basi angusta nectario destituto ca. 10 mm. supra basin furcato parte
ovuliferi fusiforma 1—1.5 mm. supra basin 0.8-1.2 mm. crasso deinde
sursum in stylum gradatim transmutato; fructu ignoto.

PERU: Jaen, dept. Cajamarca, prov. Jean, 700-800 m. alt., tree 8-20 m.
tall, fl. white, “Iguaguana,” April 1912, A. Weberbauer 6213 (Typr, Gray
Herb.).

A very well marked species of northwestern Peru, apparently re-
stricted to the seasonally dry interandean valleys of the upper Marafion
drainage, ef. Weberbauer, Bot. Jahrb. 50: 92-3 (1914). The tree is

64 JOURNAL OF THE ARNOLD ARBORETUM (VoL. XXXII

locally known as “iguaguana.” It is a member of the section Geras-
canthus. Only three other species of this section are known from
western South America. The best known is the widely distributed and
variable C. alliodora (R. & P.) Oken, which is readily distinguished by
its indument of stellate hairs and by the swellings on twigs and in the
inflorescence which serve as ant-domatia. The other members of the
section in the area have simple hairs only and are not myrmecophilous.
All three are local in distribution and very distinct and easily distin-
guished. Cordia Goeldiana Huber, formerly known only from the state
of Para, Brazil, has been recently found in northern Colombia (Pivi-
jay, dept. Magdalena, 1948, R. Romero Castafieda 1106). It has
elongate oblong corolla-lobes which have parallel lateral margins.
Cordia macrantha Chodat is a rare tree of western Ecuador, ef. Little,
Caribbean Forester 9: 269 (1948). Like C. iguaguana it has broad
rounded corolla-lobes. It differs from the Peruvian species in its larger
elliptic leaves scantily though evidently villose-hispidulous on the lower
surface. Its flowers are also much larger. The corolla is 35-43 mm.
long, the limb is 35-40 mm. in diameter, and the sinus between the
lobes are truncate rather than acute. The corolla is glabrous except
for a tuft of hairs at the base of each filament. The ovary, unlike that
of C. iguaguana, is short and broad. It is surrounded at the base by
a well-developed collar-like nectary, and has the style arising abruptly
from its broad summit. The evidence available indicates that C.
macrantha and C. iguaguana are both heterostylic. The type of the
latter is probably the long-style form of the species.

Cordia viridis (Rusby), comb. nov.

Bourreria viridis Rusby, Descr. 300 So. Amer. Pl. 100 (1920).

VENEZUELA: Lower Orinoco, 1896, Rusby & Squires 259 (AA, IsoTYPE) ;
Guayapo, Bajo Caura, Bolivar, 100 m. alt., 1939, L. Williams 11745 (G); El
Toro, La Paragua, Bolivar, 70 m. alt., 1940, Williams 12699 (G).

A species of the lower Orinoco Valley which has been confused with
C. sericicalyx DC. It is readily distinguished once its characters are
recognized. The fruit is much larger and is elongate and arises erect,
not obliquely from the calyx. Its leaves, though having a similar sparse
minute strigosity, differ from those of C. sericicalyx in their more
acuminate apex and more numerous (about 10) pairs of primary veins.

Cordia Bridgesii (Friesen), comb. nov.

Varronia Bridgesi Friesen, Bull. Soc. Bot. Genéve ser. 2, 24: 172, f. 9 (1933).

BOLIVIA: Rio Caine, Cochabamba, 1180 m. alt., sandy soil, shrub 3-5 dm.
tall, fl. white, Jan. 1949, M. Cardenas 4239 (G).

The above cited collection agrees with the original description and
illustration in all significant details. The only exception is the shorter
apical appendages on the calyx-lobes. The original description calls
for free appendages 2 mm. long. The collection made by Cardenas has
free appendicular tips 0.2-0.5 mm. long. The species is a very well

1952] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXII 65

marked one and was based on a specimen unprovided with precise
locality data, collected by Thomas Bridges in Bolivia in 1844 or 45,
probably during his journeys north and east of Cochabamba. It was
accordingly obtained in the general area where Cardenas also found it.

Cordia lutea Lam. III. 1: 421 (1791); Svenson, Am. Jour. Bot. 33: 421
and 478 (1946).

Cordia marchionica Drake, Ill. Fl. Ins. Pacific 240 (1892) and FI. Polynése
Fr. 129 (1893) ; F. B. H. Brown, [Fl. Southeast. Polynesia 3: ] Bull. Bishop
Mus. 130: 243 (1935),— based on material from “Iles Marquises (Mer-
cier!, Jardin 54!).” |

MARQUESAS ISLANDS: Uahuka, 1921-22, EF. H. Quale 1737 (A); Hatutu,
shrub on windward cliff, Sept. 27, 1922, Quale 1556 (A).

It has been a surprise to discover that Cordia marchionica of the
Marquesas is the same as the well-known and very distinct C. lutea
of the Galapagos Islands and the adjacent drier portions of western
Ecuador and northwestern Peru. Brown in his detailed account of the
flora of southeastern Polynesia reports the plant from most of the
islands in the Marquesas group and states that it is common there
in dry exposed situations below 1000 m. alt., in places forming thickets
or even becoming one of the dominant woody plants.

With the recognition of the Marquesas plant as identical with C.
lutea of western South America the question arises as to the origin of
the species in the archipelago. As a Polynesian plant with clear Ameri-
can relationships it can be used by those seeking evidence of direct
floristic affinities between Polynesia and America, cf. A. M. Adamson,
Bull. Bishop Mus. 139: 31 (1936). It is conceivable that birds may
have played a role in the spread of the species within the Marquesas,
but considering the large size of its drupe, any suggestion of direct bird-
transportation from America would be fantastic. That the species
reached the Marquesas from America in ocean-drift seems equally
unlikely. This Cordia is not a strand plant. It is a shrub or small tree
of arid situations inland where the rainfall is not only scant but also
limited to a very few months each year. The seeds of such plants must
germinate promptly when moisture becomes available if the seedling
is to become established during the short rainy season. Their seeds and
any sheathing part of the fruit are necessarily permeable by water and
accordingly unadapted for immersion in sea-water and so for success-
ful ocean dispersal. Furthermore, a recent analysis of ocean-dispersed
strand plants (Johnston, Sargentia 8: 55, 1949) indicates that with
very few exceptions such plants have been unsuccessful in colonization
across the expanse of the eastern Pacific. It is unbelievable that this
Cordia could succeed where so many better adapted plants have failed.
Any theory that Polynesians may have contributed to its presence in
the archipelago is untenable. The plant is said to be used only in
making leis, scarcely an important economic use and hardly one to
make it important to the native inhabitants. An aboriginal introduc-

66 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXXIII

tion of the plant from America might be expected only by way of
Easter Island, that lonely outlier of Polynesia in the eastern Pacific.
Although the environment of Easter Island would seem even more
suitable for the Cordia than the Marquesas, the plant is absent there.
I believe that it is significant that this Cordia was not found in the
Marquesas by the early visitors to the islands. With its great abund-
ance of large yellow flowers it is a plant not easily overlooked. I am
willing to believe that the species was introduced into the Marquesas
by the French during the 19th Century, probably as an ornamental
plant.

Eritrichium laxum, sp. nov.
Perenne humile dense caespitosis griseum pilis 0.2-0.8 mm. longis
saepe adpressis haud abundantibus vestitum; caulibus hornotinis flori-
feris debilibus gracillimis erectis vel plus minusve decumbentibus 1-6
(in statu fructifero rare ad 12) em. longis; foliis hornotinis inferioribus
em. longis spathulato-oblanceolatis 3-5 mm. latis sub. apicem
rotundum obtusumve latioribus deinde deorsum in petiolum gracilem
anguste alatum gradatim attenuatis, subtus costatis sed enervatis;
foliis superioribus paucis ad 1 em. longis: foliis annotinis desiccatis
plus minusve persistentibus saepe (praesertim petiolis) spiraliter tor-
tuosis; inflorescentia laxiflora subracemiformi bracteis foliaceis pauls
pedicellos haud suffulcientibus praedita; pedicellis sub anthesi 1-5 mm.
longis vel eis medium versus vel infra medium ecaulis orientibus 5-15
mim. longis; pedicellis fructiferis 5-20 mm. longis gracillimis adscen-
dentibus vel laxe recurvatis; lobis ealycis anguste oblongis vel oblanceo-
oblongis sparse strigosis ecostatis apice rotundis sub anthesi 1-1.5 mm.
longis 0.4-0.5 mm. latis, maturitate ad 2 mm. longis et 0.5-0.6 mm.
latis; corolla alba vel dilute sic medium versus flava 5-6 mm.
diametro; lobis patentibus rotundis ca. 2.2 mm. longis et 2 mm. latis,
tubo 1.3-1.7 mm. longo a basi 03-0. 5 mm. crasso sursum amplato
apice 1.5-1.7 mm. diametro intus 0.2-0.3 mm. supra basin nectario
lineato inconspicuo glabro donato, appendiculis faucium flavis vix
prominentibus; filamentis 0.15 mm. eee ca. 0.7 mm. supra basin tubi
corollae affixis, antheris 0.8 mm. longis; ovario 4-lobato glabro; stylo
sub anthesi 0.4-0.5 mm. longo tempore fructifero persistenti 0.5-0.7
mm. longo; nuculis a gynobase hemispherica divergentibus dorsi-ven-
traliter compressis evidenter marginatis, sine margine 1.2-1.7 mm.
longis 0.8-1 mm. latis, dorse obovatis plus minusve hispidulis, margine
aculeos 0.3-0.6 mm. longos saepe adscendentes triangulares longe atten-
uatos apice glochidiatos composito saepissime cireumdatis, facie ven-
trali glabris vel praesertim ultra medium muriculatis aliquantulum
ultra medium cicatrice parva donatis ultra cicatricem usque ad apicem
nuculi carina 0.3-0.4 mm. longa donatis alibi convexis.
TIBET: Tse La, Langong, lat. 28° 45’, long. = 00’, 14500 ft. alt., little
tufts in dry cliff crevices, corolla creamy white, eye golden, very fragrant,
F. Ludlow, @. Sherriff & G. Taylor 5619 (vpn, ee Herb.) ; hills north of

1952] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXII 67

Lhasa, 15000 ft., clumps under large boulders, fl. pale blue with yellow eye,
Ludlow & Sherriff 8806 (G); hills north of Lhasa, 14000 ft., under cliffs and
rocks, fl. white to pale blue, Ludlow & Sherriff 9704 (G); Nyenchengtang La,
4 days N. W. of Lhasa, 14000 ft., on grassy cliff-ledges and under rocks, fl.
pale blue, Ludlow & Sherriff 9648 (G).

CHINA: Chungtien Plateau, N. W. Yunnan, plant 1 in. high, on open
stony slope, K. M. Feng 1598 (G); Riutzila, one day from Atuntze, mountains
of Moying, northeast of Yangtze-Mekong watershed, N. W. Yunnan, flower
yellow, J. F. Rock 10332 (G); Kon-ka-ling, Sikang, 7. T. Yu 15005 (G);
Kansu, high rocks, 12—13,000 ft., Aug. 1914, R. Farrer 634 (G).

A very well marked species of southeastern Tibet and adjoining
western China where it appears to be the only representative of the
genus. From a weak taproot and abundant fibrous roots the plant
develops crowded short branched stems abundantly clothed with per-
sisting remnants of old leaves. These hug the soil and form a very
dense low cushiony growth which may become at least a decimeter
in diameter. From it arise the functional basal leaves and the weak
sparingly leafy fertile branches. The herbage is green or grayish green
and not silvery silky as with most other species of the genus. The fruit
is especially distinctive. The nutlets are dorsi-ventrally compressed
and proportionately not so thick as those of other congeners. Further-
more, they have an attachment scar that is slightly, but still very
clearly supramedial, rather than medial or inframedial. As a result
they have a shorter ventral keel and the under face is low convex rather
than boat-shaped or frustum-like. The only fully ripe nutlets studied
are those of the type-collection and these were detached and found
adherent to the foliage. Most of them have triangular marginal
appendages of the type described, but a few have only a thickened
marginal rim bearing a few reduced strongly inflexed (not spreading)
appendages. This latter type of nutlet possibly may be produced from
cleistogamic flowers at the base of the stems, as in Actinocarya (cf.
Johnston, Jour. Arnold Arb. 21: 52. 1940) or be the odd nutlet in a
heteromomrphie fruit. The precise condition can be determined only
when specimens in prime fruiting state become available for study.

Eritrichium elongatum Wight var. Paysoni, var. nov.

A forma genuina differt radice gracillima apice caules rosulasque
foliorum perpaucas emittenti, innovationibus ut videtur biennis et post
tempus fructiferum deciduis; caulibus fertilibus e rosulis foliorum anno-
tinis erumpentibus 2-12 cm. altis rigidis erectis cymas terminales et
laterales Sens nee 2-2.5 mm. diametro; nuculis margine
evidenter dentato don

UTAH: La Motte a ane Mts., Summit Co., alpine meadows, 11500 ft.,
July 19,1916, Z. B & L. B. Payson 5039 (G); Henry Forks Basin, Uinta Mts.,
Summit Co., stony slopes and ridges of open forks in upper tree zone above
Henry Forks Lake, plants 2-10 (-12) em. tall, 10850 ft. alt., Aug. 4, 1936,
Bassett Maguire, Dean Hobson, & Ruth Maguire 14385 (typr, Gray Herb.) ;
Upper Henry Forks Basin, in stony soil north of Lake Blanchard, common

68 JOURNAL OF THE ARNOLD ARBORETUM [voL. XXXII

above timber-line, 11200 ft. alt., plant 3-10 (-12) em. tall, Aug. 4, 1936,
Maguire, Hobson & Maguire 14346 (G); Krebs Basin, Uinta Mts., Duchesne
Co., 11400 ft. alt., southeast slopes, alpine tundra near first of the upper
lakes, July 19, 1933, F J. Hermann 5038 (G); Uinta Mts., 12000 ft., Aug.
1869, S. Watson 849 (G)

Over twenty-five years ago I received a collection of the present plant
from the late Edwin Payson, who wrote me at the time that having
observed it in the field he believed it to be very different from the
Eritrichium elongatum with which he was familiar in the mountains
of Wyoming and Colorado. Despite Payson’s belief that an undescribed
species was involved, his plant was put away as one of the many
minor forms of EF. elongatum. Subsequently other obviously similar
plants have been received. These all came from the Uinta Mts. of
northeastern Utah, where no other representative of the genus is
known. They all have relatively stout erect branched fertile stems and
all appear to be relatively short-lived plants that never develop the
woody caudex or achieve the dense pulvinate growth-form of usual
E. elongatum. I suspect that the plants may live only a few years,
probably only two or three. In any case the stems, along with the
leaves of the rosette that clothe their base, all die back to the ground
after the fruit is matured. There is accordingly no great accumulation
of old leaves as in EF. elongatum. The leafy mass at the base of the
plant becomes only 1-5 em. broad. The nutlets always have a dentate
margin and are accordingly unlike the unarmed nutlets of the most
common and widely distributed forms of EH. elongatum. They are,
however, very similar, in fact indistinguishable from those of the un-
common form of FE. elongatum, the var. argenteum. The plant of the
Uinta Mountains may possibly merit specific rank, but pending further
observations by those who can study it in the field it seems best to
treat it as a very well marked geographic variety. The name of Edwin
Payson is properly associated with it.

Trigonotis ciliolata, sp. nov.

Herba repens; caulibus elongatis 1-1.5 mm. crassis hispidulis pilis
divaricatis 0.5-1.2 mm. longis donatis; foliis alternis ovatis 15-22 mm.
longis 8-16 mm. latis utrinque pilis 0.5-1 mm. longis rigidulis (in
facie superiore laminae adscendentibus, in facie inferiore adscendenti-
bus et erectis) e basi pustulata orientibus obsitis, apice obtusis apicula-
tis, basi obtusis asymmetricis in petiolum 1-2 mm. latum 2-5 mm.
longum abrupte contractis; floribus caulinis solitariis extra-axillaribus
saepe circa insertione petioli orientibus tempore anthesi 2-3 mm. longe
pedicellatis, maturitate 10-15 mm. longe pedicellatis; sepalis lanceolatis
sub anthesi 4 mm. longis supra basin 1-1.5 mm. latis, maturitate ad
4.5 mm. longis et 1.5-2 mm. latis, sparse hispidulis, apice acutis vel
paullo attenuatis; corolla alba, lobis orbicularibus 3-4 mm. diametro;
tubo 2.2 mm. longis a basi ca. 1.5 mm. crassa sursum ampliato apice ad
2.8 mm. diametro, appendiculis faucium trapeziformibus prominulis

1952] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXII 69

puberulentis, filamentis 0.4 mm. longis medio tubi corollae orientibus,
antheris 0.7-0.8 mm. longis oblongis infra medium affixis; stylo maturi-
tate ad 1.3 mm. longo apices nuculorum vix superanti; nuculus 4 tetra-
hedraeis nigris laevibus subnitidis angulatis, angulo adaxillari ca. 1 mm.
longo, faciebus 3 inferioribus subaequalibus planis; facie superiore
nuculae convexa triangulari ca. 2 mm. longa et lata, marginibus ciliolatis
(pilulis 0.1 mm. longis) eis lateralibus valde acutis, margine abaxillari
anguste alato (ala adscendente curvata 0.2-0.3 mm. lata).

DUTCH NEW GUINEA: Angi, creeping on sandy bank along Iray River,
Lake Giji, Arfak Mts., 1900 m. alt., fl. white, April 8, 1940, R. Kanehira & S.
Hatusima 13883 (Type, Arn. Arb.).

Though very different in fruit, the present plant simulates T’.. abata
very closely in gross habit and vegetative characters and is probably
most closely related to it. It comes from western Dutch New Guinea
about 600 km. northwesterly from the high mountain valleys, near
Lake Habbema, where T. abata Johnston (1940) has been collected.
The completely glabrous nutlets of 7. abata are bifacial, having a large
convex back, a broadly angled adaxial face, and a superbasal attach-
ment. The very angulate nutlets of 7. ciliolata are tetrahedral and bear
their attachment at the peak of the equally three-sided pyramid.
Their fourth surface, the uppermost and outer one, is also triangular
but differs in being convex. Its abaxial edge bears an up-curving knife-
like wing. Its lateral edges are merely sharply acute. All three of its
edges are ciliolate, a unique development in the genus. Unlike most of
the repent Malaysian species of Trigonotis, T. ciliolata has distinctly
tetrahedral nutlets generally similar in type to those prevailing in the
genus in other regions. Its discovery lends additional support to the
belief that the Malaysian species, formerly segregated as Zoelleria and
Havilandia, are, indeed, merely aberrant members of the present genus,
ef. Johnston, Jour. Arnold Arb. 21: 58 (1940).

Trigonotis cupulifera, sp. nov.

Herba 1-2 dm. alta sparse strigosa (pilis rectis 0.1-0.5 mm. longis
antrorsis) ; caulibus gracilibus erectis subsimplicibus 1-1.5 mm. crassis;
foliis firmiusculis ovatis vel ellipticis sparse strigosis 1-2.5 cm. longis
8-12 mm. latis, basi saepe rotundis obtusisve in petiolum abrupte
contractis, apice rare acutis plerumque rotundis obtusisve et saepe
minute apiculatis; petiolo folii inferiori 1-2 cm. longo folii superiori
2-10 mm. longo; cymis solitariis caulem terminatis et axillis foliorum
supremis orientibus gracillimis ebracteatis maturitate ad 8 cm. longis
laxifloris; sepalis sparse strigosis sub anthesi 1-1.3 mm. longis 0.6—
0.7 mm. latis acutis ovato-lanceolatis, maturitate ovatis patentibus
2-2.5 mm. longis ad 1.3 mm. latis, 0-3 mm. longe pedicellatis; corolla
cacrulea, limbo ca. 4 mm. diametro, lobis rotundis ca. 1 mm. latis,
tubo cylindrico 1 mm. longo 1.38 mm. crasso; staminibus medio tubi
affixis; nuculis valde angulatis laevibus glabris tetrahedraeis nitidis
0.8-1 mm. longis, basi haud pedicellatis, facie superior! evidenter

70 JOURNAL OF THE ARNOLD ARBORETUM [VOL, XXXIII

marginatis, margine opaco erecto crassiusculo 0.3-0.6 mm. alto eupulum
formant.

KIANGSI: Lin-chuan, 105 m. alt., by river, fl. white, June 20, 1932, Y.
Tsiang 9931 (vTypE, Gray )

HUNAN: Changsha, along Linyang-ho, 35 m. alt., in thickets, April 25,
1918, Handel-Mazzetti 11687 (G).

The collection from Changsha, cited above, is the basis for recent
reports of T. brevipes Maxim. from China, ef. Hand.-Mazz. Symb.
Sin. 77: 820 (1936) and Johnston, Jour. Arnold Arb. 18: 6 (1937). It
consists of plants in flowering state which simulate the Japanese plants
closely in vegetative characters and general aspect. The resemblance,
however, proves to be deceptive. Now that fruit of the Chinese plant
is available for study it is obvious that any resemblances between the
Chinese and Japanese plants is superficial and inconsequential. The
two differ so widely in nutlets that direct relation between them is
unbelievable.

The nutlet of 7. cupulifera has a smooth, lustrous, acutely angled,
distinetly tetrahedral body which is 0.8-1 mm. in length along the
inner angle. The total nutlet, however, is actually longer, for its upper
face is bordered by an upturned erect thickish marginal flange 0.3-
0.6 mm. high. The outer faces of this flange are in the same plane as
the flat sides of the nutlet body directly beneath and may appear at
first sight to be a continuation of them. Nutlet body and flange, how-
ever, are readily distinguished by close examination, since the former
has a lustrous and the latter an opaque surface. The shallowly cup-
shaped superstructure on the nutlet body is a distinctive feature of
our present species

—

Trigonotis disviiusie sp. nov.

Planta herbacea fortasse rhizomate gracillimo oriens; ramis gracili-
-bus laxe decumbentibus 1-5 dm. longis 1-2.5 mm. crassis saepe laxe
longeque ramosis sparse antrorseque strigosis (pilis 0.2-0.8 mm. longis) ;
foliis numerosis caulinis ellipticis vel elliptico-ovatis vel ovato-lanceo-
latis 2-6 (saepe 3-4) em. longis 8-27 (saepe 10-20) mm. latis superi-
oribus quam inferioribus saepe duplo vel triplo minoribus, saepe
apiculatis costatis sed obscurissime nervatis, apice acutis vel obtusis
rotundisve, basi obtusis vel rotundis in petiolum 3-18 mm. longum
alatum ca. 1 mm. latum abrupte contractis, facie superiore pilis sparsis
0.3—1.2 mm. longis adpressis vel ndacendentihus non rare basi bulbosa
vel disciforma orientibus obsitis, facie inferiore antrorse strigosis;
cymis numerosis ebracteatis simplicibus vel fureatis 0-5 em. longe
pedunculatis terminalibus et axillis foliorum superiorum orientibus
maturitate 5-15 em. longis dissitifloris; calyce sub anthesi 1.5 mm.
longo 0-1 mm. longe pedicellato, lobis oblongis vel elliptico-obovatis
1-1.2 mm. longis, 0.2-0.4 mm. latis; calyce fructifero 2-3 mm. longo
2-3 mm. longe graciliterque pedicellato basi incrassato plus minusve
pallido, tubo cupulato, lobis adscendentibus 1.5-2.5 mm. longis ob-

1952] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXII yal

lanceolatis 0.2-0.6 mm. latis basin versus angustatis apice obtusis;
corolla caerulescente 2—-2.5 mm. diametro, tubo 1 mm. longo ecylin-
drico 1.2 mm. crasso, limbo ad 2 mm. diametro, lobis rotundis 0.7-
0.9 mm. longis; staminibus paullo supra medium tubi corollae affixis;
nuculis 4 angulatis tetrahedraeis laevibus glabris nullo modo pedicel-
latis, angulo adaxillari 1 mm. longis, angulo facies superiorem ali-
quantulum concavam circumdato acutissimo prominulis, apice nuculis
inconspicue producto et sursum curvato.

WESTERN SZECHUAN: Mt. soe Fu-hu-sse, roadside, 3 dm. tall, fl.
blue, May 14, 1942, W. P. Fang 18747 (G); Mt. Omei, Tru-dien, May 28,
1941, Fang 16747 (G); Mt. Omei, fore train-ping, 35 em. tall, May 21, 1940,
C. L. Sun he (G); Mt. Omei, Hong- train-ping, fl. blue, July 8, 1940, 7. C.
Lee 2704 (G); Mt. Seas Hung- chun-ping, roadside, 950 m alt. June 16,
1938, H.C. tae 7582 (G); Mt. Omei, Hume-chun-ping, roadside, May 22,
1944, H. C. Chow 11851 (G); Mt. Omei, Kuan-hsin-an, ditch, fl. bluish,
1378 m., July 17, 1939, S. C. Sun & K. Chang 877 (TYPE, Gray Herb: Mt.
Omei, 950 m. alts about thicket, fl. bluish, July 1, 1931, F. T. Wang 23129
(G); Mt. Omei, 850 m., hillside, plant 2.5 dm. tall, “ blue, July 18, 1938,
T.C. Peng 17 (G); without locality, Faber 598 (NY)

KWANGSI: Nan Kan, Ling Yi Hsien, valley hale. fl. bluish, Apr. 13,
1933, Steward & Cheo 184 (G).

This plant, and also JT. lara, were incorrectly identified as T.
omeiensis Matsuda (1919), in my synopsis of the Trigonotis of southern
China, Jour. Arnold Arb. 18: 6 (1937). Matsuda’s species properly
belongs in the synonymy of the very different 7. Cavaleriei (Lev.)
Hand.-Mazz. Our present plant is without a name. Its closest relative
is T. lara. Among its distinctive features are the up-curving corners
of the upper face of the nutlet and the mineralized pallid epidermis
developed on the thickened base of old fruiting calyces.

Trigonotis laxa, sp. nov.

Herba ut videtur perennis; caulibus 2-5 dm. longis 1-2.56 mm.
crassis simplicibus vel sparse ramosis sparse strigosis; foliis caulinis
costatis sed enervatis, lamina elliptica vel ovato-elliptica vel late
lanceolata saepe 2-3 cm. longa et 6-15 mm. lata, apice obtusa vel
rotunda et apiculata vel rare acuta, basi obtusa rotundave in petiolum
3-10 mm. longum alatum 1—2 mm. latum contracta, supra glabra vel
apicem versus sparse strigosa, subtus sparse strigosa (pilis 0.2-0.5 mm.
longis non rare basi incrassata disciformave orientibus) ; inflorescentia
terminali et axillis supremis orientibus; cymis gracillimis ad 1 dm.
longis fureatis 1-8 mm. longe pedunculatis ebracteatis; corolla sub-
caerulea vel subalba, limbo 5 mm. diametro, lobis rotundis 1.5 mm
latis, tubo 1-1.5 mm. longo a basi ca. 1 mm. crasso sursum gradatim
ampliato apice ad 1.8 mm. diametro; staminibus medio tubi affixis;
calyce subanthesi 1.5-2 mm. longo 0.3-1.5 mm. longe pedicellato sparse
strigoso, lobis obovatis 1.7 mm. longis 0.3-0.4 mm. latis apice rotundis;
calyce fructiferi 2 mm. longo ad 6 mm. longe pedicellato, lobis laxe

72 JOURNAL OF THE ARNOLD ARBORETUM [vVoL. XXXII

adscendentibus 0.4-0.6 mm. latis obtusis; nuculis acute angulatis 4
tetrahedraeis haud pedicellatis glabris laevibus, angulo adaxiale ca.
1 mm. longo, facie superiori margine angustissime alato circumdata;
stigmate nuculis breviter sed distincte superanti.

SZECHUAN: Nanchuan Tn roadside, 8000-9000 ft., fl. white, May 25,
1928, W. P. Fang 915 (G); nehuan Baien. roadside, 8000-9000, herb
1-1.5 ft. tall, fl. pale blue, ie 31, 1928, W. P. Fang 1159 (G); roadside,
5000-6000 ft., herb 1 ft. tall, fl. pale blue, June 1, 1928, W. P. Fang 1348
(TYPE, Gray Herb. ).

This plant of southeastern Szechuan is most closely related to T.
floribunda of western Szechuan and western Kwangsi. It differs in
having a larger corolla with an ampliate rather than cylindrical tube,
a protruding style, broader and shorter? calyx-lobes, and different
nutlets. The mature calyx, unlike that of T. floribunda, does not have a
noticeably thickened base covered with mineralized epidermis, nor
does it develop a short but distinct cupulate tube. In the fruit the
upper faces of the nutlets slope away from one another much less
steeply than those of 7. lara. In T. floribunda, but not in T. lazxa,
the corners of the upper nutlet face are characteristically up-curving.
The margin about the upper nutlet face in 7. lara tends to become very
narrowly winged on the side opposite the nutlet apex. In 7. floribunda
the nutlet angles are acute and with a suggestion of a wing, but the
latter is not so pronounced as in T. laxa. The relationship between
the two species is very clear, but the two are certainly distinct.

Microula blepharolepis (Maxim.), comb. nov.
Omphalodes blepharolepis Maxim. Bull. Acad. St. Petersbourgh ser.
3, 27: 504 (1881) and Mel. Biol. 11: 269 (1881) ; Brand, Pflanzenr.
Heft 78; 105 (1921).

Microula diffusa (Maxim.), comb. nov.
Omphalodes diffusa Maxim. Bull. Acad. St. Petersbourgh ser. 3, 27:
504 (1881) and Mel. Biol. 11: 270 (1881); Brand, Pflanzenr.
78: 106 (1921).

I am indebted to Prof. W. Th. Kuprevicz, Director of the Komarov
Botanical Institute, Leningrad, for the great privilege of examining
the types of Omphalodes blepharolepis and O. diffusa. These species,
based on material from western China, in the region south and south-
west of Lake Kokonor, are very definitely members of Microula, a
genus well developed in the area of Lake Kokonor south into Yunnan.
In technical characters and general appearance they are obviously
members of Microula. Both species are well marked and distinct from
any heretofore seen by me.

Cryptantha Weberi, sp. nov.

Planta perennis caespitosa pallida e radice palari oriens caudice
denso humili multicipitali proferens; caulibus numerosis erectis foliosis
10-18 em. longis supra medium floriferis, plus minusve hispidis (pilis

1952] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXII 73

patentibus 1-2 mm. longis) et abundanter hispidulo-villulosis (pilulis
0.2-0.3 mm. longis saepe retrorso-adpressis) ; foliis pallidis numerosis
abundanter hispidulo-villulosis (pilulis adpressis 0.1-0.3 mm. longis)
et hispidis (pilis 1-2.5 mm. longis basi incrassato orientibus adpressis
vel praesertim secus marginem laminae adscendentibus vel rare paten-
tibus) anguste oblanceolatis apicem versus latioribus firmis incon-
spicue costatis margine vix revolutis apice rotundis obtusisve; foltis
basalibus confertis tempore florendi vigentibus 3-8 cm. longis 3-7 mm.
latis; foliis caulinis numerosis medionalibus eis basalibus similibus sed
minoribus 2-3 em. longis 2-4 mm. latis; foliis supra medium caulis
cymis axillaribus breviter pedunculatis suffultis; inflorescentia cylin-
drica infra medium bracteis exsertis donata, juventate densa 1.5-2 cm.
crassa 4-6 em. longa, maturitate plus minusve interrupta 2-2.5 em.
crassa 8-10 cm. longa; cymis abundantibus sub anthesi glomeratis
5-8 mm. longis, fructiferis 10-15 mm. longis; calycibus ad anthesim
3-4 mm. longis ,fructiferis 5-6 mm. longis, 0-8 mm. longe pedicellatis,
lobis basi 0.7-1 mm. latis sursum angustatis, inconspicue costatis sparse
hispidis (pilis ca. 1 mm. longis) et dense hispidulo-villulosis apice
rotundis; corolla alba, tubo cylindrico 3 mm. longo ad 2 mm. crasso,
lobis suborbicularibus 2-2.5 mm. diametro, limbo 4-6 mm. diametro;
staminibus supra medium tubo corollae (0.5-0.7 mm. infra faucem)
affixis; stylo cum gynobasi 3.5-4 mm. longo; stylo 1.5-1.8 mm. longo;
nuculis 2-2.3 mm. longis 1.3-1.8 mm. latis ovatis fere symmetricis
compressis angustissime marginatis, ventre sublaevibus sulcatis (sulco
aperto anguste cuneato ad 0.5 mm. infra apicem nuculae attingenti-
bus), dorse tuberculis et rugis brevibus irregularibus plus minusve
transversis sparse donatis.

COLORADO (Saguache County): along road to Stone Cellar Ranger
Station and Saguache Park, near junction of main highway, 4 miles west of
Cochetopa Pass, voleanic ash deposit, 9700 ft., with Aster coloradensis,
Senecio Hallii and Penstemon secundiflorus, July 28, 1950, William A. Weber
5778 (vrype, Gray Herb.); dry knoll one mile north of Stone Cellar Ranger
Station, 9000 ft., July 7, 1936, Reed C. Rollins 1323 (G); Carnero Ranger
Station, Cochetopa Forest, scattered on lower slopes, 10000 ft., June 20, 1922,
C. E. Taylor 573 (G).

A very well marked species of the mountains of south central
Colorado for which I can suggest no close relative. In Payson’s
revision of Cryptantha § Oreocarya, Ann. Missouri Bot. Gard. 14: 239,
240 (1927), it keys out to C. virgata. In Brand’s treatment, Pflanzenr.
Heft 97: 79 (1931), it keys out to O. rugulosa. The caespitose and
perennial habit, different indument and smaller nutlets with open scar
all readily distinguish it from the very different C. virgata. Cryptantha
rugulosa shows greater similarity with C. Weberi but differs in indu-
ment, in the cylindric thyrse of short cymes, small nutlets, ete.

Two flowering collections of the species have been known for over
ten years. Though recognized as representing what was evidently an
unnamed plant, they have not been used in describing the species, since

74 JOURNAL OF THE ARNOLD ARBORETUM [VoL. XXXII

they lack fruit. For at last receiving material of the plant in mature
state [ am indebted to Professor William A. Weber, of the University
of Colorado, who made a special visit to the mountains of Saguache
County in search of it. It is with great pleasure that I associate his
name with the species.

Craniospermum mongolicum, sp. noy.

Herba humilis multicaulis ut videtur perennis et 5-6 cm. alta;
caulibus hornotinis 6 em. longis infra medium simplicibus supra me-
dium cymas brevipedunculatas axillares dense dispositis proferentibus
sparse hispidis (pilis patentibus vel adscendentibus 1-2 mm. longis) et
hispidulis (pilulis 0.4-0.8 mm. longis retrorse adpressis) ; foliis firmis
enervatis villuloso-hispidulis (pilulis saepe retrorse adpressis 0.3-0.9
mm. longis vix abundantibus) et sparse hispidulis (pilis rigidis e
basibus pallidis incrassatis prominulis erumpentibus facie superiore
et praesertim margine laminae folii gestis) foliis basalibus congestis
tempore florendi marcidis oblanceolatis 1-2 mm. latis 8-11 mm. longis;
foliis caulinis oblanceolatis 15-20 mm. longis 3-4 mm. latis apice
acutis obtusisve; calyce 5-partito hispidulo sub anthesi 5 mm. longo
mox ad 8 mm. longo, lobis subaequalibus oblongo-lanceolatis apice
obtusis pedicello ad 1 mm. longo; corolla ca. 7.5 mm. longa glaberrima,
tubo cylindrico 2.5 mm. crasso, faucibus 2 mm. longis apice 4 mm.
crassis, lobis caerulescentibus adscendentibus nervatis 1.5 mm. longis
a basi ca. 1.5 mm. lato sursum laeviter attenuatis apice rotundis;
staminibus basim versus faucis (ca. 4 mm. supra basin corollae) affixis
e plicis intrusis extus foveolatis orientibus glaberrimis; filamentis 5-6
mm. longis e fauce corollae 3-4 mm. longe exsertis ligulato-linearibus
basin versus aliquantum ampliatis basi imo geniculatis; antheris 0.5-
1.2 mm. longis; stylo filiformi glabro longe exserto; nuculis maturis
homomorphis 3.5 mm. longis cinereis minute verrucosis sparse tubercu-
latis, dorse supra medium foveola conspicua donatis.

OUTER MONGOLJA: Daying Gol, dry hills at 5500 ft. alt., 1925, R. W.
Chaney 195 (Type, Gray Herb.).

From the other members of the genus this well-marked species differs
in having a low compact growth habit, hispid or hispidulous rather
than villous indument, and a branched inflorescence. Furthermore, its
filaments are not entirely linear nor do they arise directly from the
walls of the corolla. They are broadened towards their geniculate base
and each arises from a small protuberance situated low in the corolla
throat. On the outside of the corolla the location of each filament-
bearing protuberance is marked by a small but distinct depression.
The plant appears to have a perennial root. From it arise a number
of short stems about 6 cm. long. These stems bear not only terminal
cymes but also numerous, equally well developed lateral ones from
the four to six axils above its middle. The inflorescence is obviously
compound, It is dense, 2.5-3.5 em. long, and the most conspicuous part
of the plant. In other species the inflorescence consists of a solitary

1952] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXII 75

subeapitate glomerule borne terminal on each stem. These stems are
better developed than those in C. mongolicum and are more elongate
and very much more leafy. In the most recent account of Craniosper-
mum, Pflanzenr. Heft 97: 102-3 (1931), Brand recognizes three
members of the genus, C. canescens DC., C. subvillosum Lehm., and
C. subfloccosum Krylow. All these are closely related, so closely in
fact that they may be no more than forms of a single somewhat
variable species. They grow in southern Siberia along the Mongolian
border from the west of the Altai east to beyond Lake Baical. The
proposed new species is more southerly, occurring along the southern
extension of the Altai in western Outer Mongolia. The type was
obtained at Daying (or Da-Ying) Gol, a stream heading in the Baga
Bogdo Range and flowing north into Tsagan Nor, about long. 101° 30’
and lat. 45° 5’. It was distributed misidentified as “Arnebia guttata.”

Trichodesma calycosum Collett & Hemsl., Jour. Linn. Soc. Bot. 28:
92 (1890); Lacaita, Jour. Linn. Soc. Bot. 43: 476 (1916) — type
from Burma; “Shan hills at 4000 feet.”

Lacaitaea calycosa (Coll. & Hemsl.) Brand, in Fedde Repert. 18: 81 (1914)
and Pflanzenr. Heft 87: 44 (1921).

Trichodesma Hemsleyana Levl. in Fedde, Repert. 9: 327 (1911) and Fl.
Kouy Tcheou 55 (1814); Brand, Pflanzenr. Heft 78: 43 (1921) — type
from “Kouy-Tcheou [Kweichow, China], J. Esquirol 745.”

Trichodesma sinicum Brand in Fedde, Repert. 12: 504 (1913) and Pflanzenr.
Heft 78: 43 (1921)—type from Szemao, southern Yunnan, Henry
10124 D.

Trichodesma calcareum Craib, Kew Bull. 1914: 8 (1914) —type from
northern Siam, “Doi Chieng Dao, crevices of limestone rock, 900 m.,
Kerr 2856.”

Trichodesma khasianum var. calcareum (Craib) Brand, Pflanzenr. Heft 78:
33 (1921).

Octosomatium Kerrii Gagnep. Not. Syst. 14: 23 (1950) — type from Laos,
Indo-China; Muong-Khao, Xieng-Khouang dist., ca. lat. 19° 50’ and
long. 103° 30’, Kerr 20978.

Known from Burma (north of lat. 22°) and from China (western
and southern Yunnan; Kweichow) south to about lat. 19° in northern
Siam and adjoining Indo-China; also in Sikkim.

BURMA: Kanpetlet, Chin Hills, 7500 ft., hillside, fairly common, shrub
15 ft. tall, trunk 3 in. thick breast-high, fl. white, F. G. Dickason 8425 (A);
Webula (Falam), damp ravines and open mountain slope at 4000 ft., fairly
common, shrub 8-10 ft. tall, Dickason 7347.

INDO-CHINA: Muong-Khao, Xieng-Khouang, Laos, Kerr 20978 (Paris,
tyPr of O. Kerrii; fragment received on loan).

CHINA: Yunnan: Salween Valley, lat. 25° 6’, in open seruh at 4000 ft.,
shrub 20 ft. tall, Forrest 13665 (A); Salween Valley, lat. 25° 10’, long. 98° 50’,
open dry situations on margin of scrub, half shrubby plant 3-4 ft. tall,
Forrest 19341 (A); Lan-Tsang Hsien, 1100 m. alt., woody plant on mountain
slope, C. W. Wang 72127 (A); Mengtse, 5000 ft. alt., shrub 3 ft. tall, fl.

76 JOURNAL OF THE ARNOLD ARBORETUM [VoL. XXXIII

white, A. Henry 10124 (A, NY); Lunan, shrub 5 ft. tall, fl. white, Henry
10124A (A, NY); Chu-yuan, shrub 3 ft. tall, Henry 10124B (A, NY); Red
River Valley, Manpan, 4000 ft., shrub 4 ft., fl. white, Henry 10124C (A);
Szemao, 5000 ft., shrub 10 ft. tall, fl. white, om 10124D (A, tsoryre of T.
sinic um). Kweichow: Lo fou, Feb. 1909, J. Cavalerie 3498 (G); without
locality, J. Esquirol 745 (G, photo & frag., type of 7. Hemsleyana).

A study of the recently published Octosomatium Kerrii Gagnep.
reveals it to be a synonym of the present plant, which now has accu-
mulated four different trivial names and has twice been made the type
of a monotypic genus, i.e. Lacaitaea Brand and Octosomatium Gagnep.
It is a large shrubby plant with broad opposite leaves, which grows
1—6 m. tall and has fruit and flowers characteristic of Trichodesma in
all details save only for the 8-10 gibbose intrusions developed in the
corolla throat. Unlike most congeners it is not a desert or steppe plant
but rather an inhabitant of the tropical forest.

The invaginate appendages present in the corolla throat of T.
calycosum are of special interest. No other member of the genus has
similar appendages, and furthermore, no other member of the entire
Boraginaceae has them in so large a number. The corolla of T.
calycosum, though usually pentamerous, is occasionally tetramerous
and bears two gibbose appendages in the throat opposite each corolla-
lobe. In other Boraginaceae the faucal appendages are traversed medi-
ally by the primary vein leading to the corolla-lobe and accordingly
occur singly in the throat opposite the middle of the lobe. In T.
calycosum the two appendages opposite each lobe arise lateral to the
vein which courses between them. The condition, though unique in the
family, is readily homologized. Many of the Boraginaceae with well-
developed faucal invaginations have the invagination more or less
strongly retuse at the summit. The vein leading to the corolla-lobe
is detoured over the summit of the invaginate appendage. In those
appendages that are lobed or emarginate at the summit its course
over the summit lies at the bottom of the apical sinus. Should the
appendage become strongly bilobed or parted, the vein would have
no detour and would pass directly between the halves of the appendage,
which would then appear double. The supernumerary appendages of
T. calycosum probably had some such origin.

It is interesting to speculate why double faucal appendages should
have developed only in this species of Trichodesma. The genus is a
member of the Cynoglosseae, the most highly evolved tribe in the
Boraginaceae, in which faucal appendages are usually well developed.
Their loss in Trichodesma is probably associated with the elaboration
of the androecium, in which the large anthers, held together by twisted
tips, form a large protruding cone that controls all access to the
corolla-tube. Faucal appendages, alternating with the anthers, would
hinder rather than guide the proboscis of an insect seeking the slit
between the closely juxtaposed anthers. This is obviated by the divided

1952] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXII 77

faucal appendages of T. calycosum. In that species they probably
represent a modification and persistence of an ancestral character. As
a general rule, in any large natural group such as T’richodesma, an-
cestral characteristics are more frequently retained by the shrubby
or arborescent than by the herbaceous members. The latter, with a
short life history and more frequent generations, can be expected to
vary more rapidly and so deviate more from the ancestral than do the
woody and arborescent congeners. Significantly, 7’. calycosum is the
most robust member of the genus, and with the possible exception of
T. khasianum Clarke, the only large shrub in the genus. The normal
single faucal appendages of the Cynoglosseae having no function and
being perhaps even disadvantageous in the Trichodesma flower, have
been eliminated in most species of the genus. In T.. calycosum, divided
and so no longer hindering insects seeking the nectaries at the bottom
of the floral cup, the appendages appear to have persisted as non-
functional innocuous structures not yet eliminated in the course of
evolution.

Brand states frankly that 7. calycosum, though admittedly other-
wise typical of Trichodesma, should, because of its faucal appendages,
be segregated to form the monotypic genus Lacaitaea. Gagnepain, in
establishing the synonymous genus Octosomatium, gives no intimation
that he recognized the obvious affinities of the plant with Trichodesma.
He considered the plant remarkable because of its opposite leaves,
the prominent line on the stem joining the attachments of opposing
leaves, the tetramerous flowers (of his specimen), and finally, the
double faucal appendages of the corolla. Of the characters mentioned
by him only the faucal appendages are notable when the plant is
compared with Trichodesma. They alone are distinctive of T. caly-
cosum and they alone constitute the only character of possible generic
value that can possibly justify any proposal that the species be given
special recognition as representing a monotypic genus distinct from
Trichodesma.

In estimating the importance to be placed on the faucal appendages
of T. calycosum, I am influenced by the remarkable similarity existing
between that species and 7. khasianum. The latter ranges just west
of T. calycosum and like it is a shrub of monsoonal forests. Tricho-
desma khasianum has absolutely no faucal appendages, has less hairy
anthers, and usually has more conspicuously white-dotted upper leaf-
surfaces, but except for these differences the two species are extremely
alike, so much so that any doubt as to their extremely close relationship
is inconceivable. These two species belong together in one and the
same genus! To disassociate them generically on the basis of one
character is arbitrary and unnatural. I am, accordingly, content to
assign 7’. calycosum as well as T. khasianuwm to Trichodesma § Fried-
richsthalia, a section containing several African species which also
have ample opposite leaves combined with frutescent habit.

78 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXXIII

Trichodesma calycosum var. formosanum (Matsumura), comb. nov.

Trichodesma formosana Matsumura, Bot. Mag. Tokyo 12: 108 (1898) —
“Hab. Formosa: in montosis ad Taichu (Y. Tashiro, no. 35) ; Tenkachila-
isha, Shu-shu-kai (C. Owatari).”

Trichodesma khasianum sensu auct. Formos.; 8. Sasaki [Cat. Govt. Herb. ]
Dept. Forestry, Taihoku, Formosa, Report 9: 431 (19380); D. Hou,
Taiwania 1: 212 (1950).

FORMOSA: Kinkwaseki, 7. Ito (A, photo); South Cape, A. Henry 286
& 939 (A); Boryo to Kuraru, prov. Koshun, uae Shae ear common shrub
3-6 ft. tall, fl. blue, FE. H. Wilson 11014 (A): v. Nanto, low altitudes,
common bush, 6-12 ft. tall, Wilson 9831 (A); » Bankinsing], Henry 1239
(NY); Taito-cho, Shikano, 19381, Tanaka 10462 (NY).

This plant of Formosa has been reported numerous times as “Tricho-
desma khasianum.” Actually it is scarcely separable from T. caly-
cosum. Indeed, it is distinguishable only by the arrangement of hairs
on the back of the anthers. In the plant of Formosa these hairs are
appressed and have the appearance of having been combed in differ-
ent directions, left and right, from the mid-line of the anther. The
hairs of the anthers of the continental plant are less appressed and do
not have the combed, orderly appearance. The difference is a minor
one, and only because it seems to be geographically correlated does it
merit nomenclatorial recognition.

ARNOLD ARBORETUM,
Harvarp UNIVERSITY.

1952] KOBUSKI, THE GENUS ANNESLEA 79

STUDIES IN THE THEACEAE, XXV
THE GENUS ANNESLEA

CLARENCE FE. KosBusk1I

THE THEACEOUS GENUS Anneslea was first described and illustrated
by Wallich in his Pl. As. Rar. 1: 5, t. 5. 1829. One cannot but be
impressed by the manner in which the genus was presented. The details
offered in the description, along with the excellent plate, produce a
picture hard to duplicate even today.

According to G. Don, the genus was named Anneslea “in honor of
George Annesley, Lord Mountnorris, F. R. and L. §., who collected
many plants on his travels in the north of Africa and the south of
Europe, while Viscount Valentia.” It is fortunate that the name has
been included in the list of nomina conservanda, since other genera had
been described earlier under the same name. One of these, Anneslia
Hooker ex Salisbury (1807), belonging to the Leguminosae, has had
over a hundred species attributed to it; while Anneslea Roxburgh ex
Andr. (1810), belonging to the Nymphaeaceae, has had three species
described.

Anneslea is one of the three genera of the Theaceae recorded as
having an inferior or subinferior ovary. The other two are Visnea
Linnaeus f. (1781) and Symplococarpon Airy-Shaw (1937). The geo-
graphical distribution of these three genera is very interesting in that
they are so far removed from one another. Anneslea grows in tropical
and subtropical Asia, extending from southern China through Burma,
Indo-China and Malaya, into Sumatra. The islands of Formosa and
Hainan have both recorded the genus. The relationship of Anneslea is
clearly with Ternstroemia. Symplococarpon is found in tropical Amer-
ica, and its relationship is with the American species of Cleyera. So
close are the last two mentioned genera in appearance that when in
flower it is difficult to separate them. On the other hand, Visnea is
confined to the Canary Islands and Madeira and has no immediate
generic relative, since it is the only species of Theaceae growing in
this area.

Five synonyms have been recorded for Anneslea during the past
century. In checking the literature, I find that all the synonyms except
Richtera Reichenbach clearly pertain to this genus. Richtera was
recorded in a listing of genera by Reichenbach with no description or
reference to herbarium material. Both Dalla Torre & Harms (1907)
and Melchior (1925) record the name as a synonym of Anneslea. I
feel that I am correct in assuming that a specimen bearing this name
had been seen by these authors and thus associated with the genus.
The name Mountnorrisia, introduced by Szyszylowicz, refers also to

80 JOURNAL OF THE ARNOLD ARBORETUM [voL. XXXIII

George Annesley. The name was cleverly used by the author, since
at the time (1893) he recognized the priority of the other two
Annesleae and could not foresee the eventual inclusion of Wallich’s
genus among the nomina conservanda.

In 1948 Gagnepain described a new genus, Paranneslea, which he
separated from Anneslea by the following characters: (1) the anthers
not mucronate by a prolongation of the connective, but bi-mucronate
by lateral projections of the cells; (2) a five-parted stigma; (3) the
cells of the ovary 2-ovulate; and (4) the peduncles solitary in the
axils of the leaves.

Gagnepain states that it is difficult to designate the genus to which
Paranneslea is most closely related. Except for the four differences
listed above, the plant described would fall into Anneslea fragrans —
and after detailed study of the genus (Anneslea), some of the characters
listed by Gagnepain lose their importance.

Because of Gagnepain’s proposed new genus, I made many more
dissections of both flowers and fruits than I would otherwise have
made. Some interesting results were obtained — results not unexpected
in this family, however. Whereas the genus Anneslea was originally
described as having three cells in the ovary and fruit and a three-
parted stigma, I discovered that two-celled ovaries and fruit were
more prevalent than three-celled and were found in a ratio of two to
one. Also, there is no true relationship between the number of divisions
in the stigma and the cells of the ovary. Three-parted stigmas were
more prevalent even in flowers with two-celled ovaries.

As to the five-parted stigma of Paranneslea, this is not an unusual
variation within genera of this family. The genus Hurya has domi-
nantly a three-parted style. However, it has been discovered that a
five-parted style is not unusual, and occasionally four-parted styles
may be found.

The number of ovules in the cells of the ovary varies considerably
in Anneslea fragrans and cannot be used as a diagnostic character in
the separation of species, to say nothing of genera.

The mucronate anther cells are distinct from all the material of
Anneslea that I have studied. However, it is not unusual to find this
character present or absent in members of other genera of the family.

The fourth point, peduncles solitary in the axils of the leaves, is the
most distinctive character listed by Gagnepain.

The three characters (1) flowers solitary in the axils of the leaves;
(2) five-parted style; and (3) the mucronate anther cells, constitute
points worthy of specific, but hardly of generic delimitation. True
enough, they contribute a remarkable variation from the so-called
other species of Anneslea. However, except for Anneslea donnaiensis
and A. steenisit (Sumatra) I feel that all other known material of the
genus belongs to the single species A. fragrans with five regional
varieties.

1952] KOBUSKI, THE GENUS ANNESLEA 81

Anneslea Wallich, Pl. As. Rar. 1: 5, t. 5. 1829. —G. Don, Gen. Syst.
: 565. 1840. — Endlicher, Gen. Pl. 1018. 1840. — Choisy in Mém.

Soc. Phys. Hist. Nat. Genéve 14: 129 (Mém, Ternstr. 41). 1855. —
Bentham & Hooker, Gen. Pl. 1: 182. 1862 — Baillon, Hist. Pl. 4:
258. 1873. — Kurz, For. Fl. Brit. Burma 1: 98. 1877. — Melchior

in Nat. Pflanzenfam. ed. 2, 21: 143. 1925. — Lemée, Dict. Pl. Phan.

1: 289. 1929. — Keng in Taiwania 1: 254. 1950.— Non Anneslia
Hooker (1807), Leguminosae; nec Anneslea Roxburgh (1810),

Nymphaeaceae.
Richtera Reichenbach, Repert. eur Nomencl. Gen. Pl. 208. 1841, nomen.
—Dalla Torre & Harms, Gen. nonce: umarum Suppl. 622. 1907, as

syn. — Melchior in Nat. eee ed. 2, 21: 148. 1925, as syn

Callosmia Presl, Bot. Bemerk. 103. 1844.

Daydonia Britten in Jour. Bot. 26: 11. 1888.

Mountnorrisia Szyszylowicz in Nat. Pflanzenfam. III. 6: 189. 1893.

Annesleya Post & Kuntze, Lexic. Gen. Phan. 32. 1903.

Paranneslea Gagnepain in Bull. Soc. Bot. France 95: 29. 1948.

Calyx persistent, imbricate, bi-bracteate at the base, deeply parted
into 5 lobes, the lobes unequal. Petals 5, imbricate, connate at the base,
deeply constricted at the middle. Stamens 30-40, distinct, in a single
or in double series, the filaments inserted on the torus, the anthers
linear, usually longeuspidate. Ovary imbedded in the torus, 2- or
3-celled with a few to several ovules in each cell; ovules suspended
from the apex of the placenta, the style entire, persistent, the stigmas
usually 3, occasionally 2 or 5. Fruit inferior, subglobose, subligneous,
crowned by the persistent calyx, 2- or 3-celled, the cells 1-3-seeded.
Seeds pendulous, arillate, the embryo curved.

Trees or shrubs with alternate coriaceous persistent leaves.

Type species: Anneslea fragrans Wallich.

KEY TO THE SPECIES

A. Flowers crowded at the apex in a compact spiral arrangement; stigmas
3 or 2; the connective of the anther projected into an apicule usually
1-2 mm. long.

_ Petals connate at the base for 4-5 mm., deeply constricted on each

side at the middle, AEDST in outline like an hourglass............

1. A. fragrans.

BB. Palate joined sar Tightly at the very base, the sides entire with no

MHGGIE CONSEICHON. ..o555.210 548 aaceee ee Satees bens 2. A. steenisii.

AA. Flowers solitary in a axils of the leaves; stigmas 5; individual anther
cells apiculate, not a single apicule from the projected connective

3. Ax donnaiensis.

ee)

1. Anneslea fragrans Wallich, Pl. Asiat. Rar. 1: 5, t. 5. 1830.—G.
Don, Gen. Syst. 1: 566. 1840. — Griffith, Icon. Pl. As. t. 585, fig. 17,
1854. — Choisy in Mém. Soc. Phys. Hist. Nat. Genéve 14: 129
(Mém. Ternstr. 41). 1855. — Dyer in Hooker f., Fl. Brit. India 1:

82 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXXIII

280. 1872.— Kurz, For. Fl. Brit. Burma 1: 98. 1877. — Mason,
Burma People Prod. (Bot.) 2: 6380. 1883.— Pierre, Fl. For.
Cochinch. 2: pl. 127. 1887.— Brandis, Ind. Trees 58. 1906. —
Pitard in Fl. Gén. Indo-Chine 1: 335. 1910.—Craib, Fl. Siam
Enum. 1: 123. 1925.— Melchior in Nat. Pflanzenfam. ed. 2, 21:
143. 1925. — Gagnepain in FI. Gén. Indo-Chine Suppl. 1: 278. 1943.
Callosmia fragrans (Wallich) Presl, Bot. Bemerk. 103. 1844.
Anneslea fragrans a typica Pierre, Fl. For. Cochineh. 2: t. 127. 1887.
Mountnorrisia fragrans (Wallich) Szyszylowicz in Nat. Pflanzenfam. IIT.
6: 189. 1893
Small tree to 15 m. (occasionally shrubs or very large trees) ; branches
terete, glabrous, gray. Leaves coriaceous, few, disposed near the apex
of the branchlets, glabrous, lanceolate, oblong-lanceolate, rarely obo-
vate, (4.5-) 10-15 (-18) cm. long, (2-) 3-5 (-8) em. wide, acute at
the apex, occasionally obtuse, rarely rounded, usually cuneate at the
base, tapering along and into the petiole (2-3 em. long), punctate
below, the margine entire, subrevolute, with occasional minute glands
(seen only with a lens) ; the midrib canaliculate above, elevated below,
the veins 10-12 pairs elevated below, sometimes quite inconspicuous.
Inflorescence consisting of several to many individual flowers in
a compact spiral arrangement, subapical, terminating the season’s
growth, subtended by or occasionally intermixed with a few (ca. 3)
subverticillately arranged leaves; pedicels glabrous, usually 5-6 em.
long, often 2-3-4 cm. long, narrowest at the base; bracteoles 2, opposite
or nearly so, immediately below the calyx, broadly ovate to deltoid,
occasionally rounded, 4-4.5 mm. long, ca, 3 mm. wide, often carinate,
denticulate; sepals 5, imbricate, unequal, glabrous, ovate or rounded,
1-1.5 em. long, the outer ones smaller, thicker, with a narrow ciliate
membranaceous margin, the inner ones thickened at the center and
base, the margin more widely membranaceous, eciliate; petals 5,
imbricate, membranaceous, glabrous, ca. 20 mm. lo ong, 5-6 mm. wide,
with the over-all form of an hourglass, deeply constricted immediately
below the middle to 2 mm. or less in width, the lower portions of the
petals connate for 5-7 mm. simulating a tube but often separating at
maturity, the uppermost portion of the petals broadly ovate, acute at
the apex, appearing decidedly so because of the tendency to curve
inward; stamens 30-40, uni- or bi-seriate, 12-15 mm. long, the filaments
lightly adnate to the base of the corolla, ca. 5 mm. long, the anthers
basifixed, linear, 5-7 cm. long, the connective projected into a con-
spicuous apicule 2-3 mm. long; ovary subinferior, 2- or 3-celled, with
several ovules suspended from the apex of the placenta in each cell,
the style tenuous 1.5-2 em. long, 2- or 3-lobed at the apex, the stigmas
3. Fruit baccate, subinferior, free only at the apex, otherwise joined
with the calyx-tube, somewhat leathery, usually pustular-dotted when
mature, globular or rounded, ca. 2 em. diameter (sometimes as much
as 4 cm. long and 3 cm. diameter), crowned by persistent sepals, 2— or

1952] KOBUSKI, THE GENUS ANNESLEA 83

3-celled, each cell 1-3-seeded. Seeds with a hard woody testa and
reddish arillae, the embryo horseshoe-shaped.

YUNNAN: Kintung Chai, between Keng Hung and Muang Hing, top of
dry slopes, alt. 1350-1800 m., J. F. Rock 2703 (AA), Mar. 1, 1922 (tree).—
Between Tengyueh and the Burmese border, en route to Sadom, J. F. Rock
7314 (AA), Nov. 1922 (tree 30 ft.; flowers white, the pedicels white). — “Ad
orientem fluminis Dsolin-ho humidis ad vicum Lodse-Magai,” alt. 1900 m.,
H. Handel-Mazzetti 6164 (AA), April 29, 1915.— Kien- ehael Hsien, noo,
alt. 1900 m., H. T. Tsai 53142 (AA), April 1 13, 1933 (tree 20 ft.). — Lung-ling
Hsien, in thickets, alt. 1800 m., H. T. Tsai 55629 (AA), 55768 (AA), 55855
(AA), April 1934 (tree 30 ft., fruit yellow). — Mong-ka, in forest, alt. 1600-
1750 m., H. T. Tsai 56722 (AA), 56840 (AA), 56846 (AA), Jan-Feb. 1934
(tree 20-30 ft.; flowers yellow). — Yuanchiang, alt. 4000-5000 ft., A. Henry
11591 (AA). ., alt. 4000-5000 ft., A. Henry 11591A
(AA), 11591C (AA).—Shun-Ning, Holungtan, among thickets, alt. 1850 m.,
T. T. Yu 16195 (AA), June 10, 1958 (rare ae 4-6 ft.; capsule oale.
brownish yellow).—Shun-Ning Hsien, mountain slope, alt. 2800 m., C. W.
Wang 71812 (AA), Feb. 1936.— Chen-Kang Hsien, in woods, alt. 1600 a.
C.W. Wang 72227 (AA), March 1936.— Tsang-Yuan, north slope, alt. 1600
m., C. W. Wang 73247 (AA), April 1936 (tree 15 m.).— Fo-Hai, alt. 1540 m.,
C. W. Wang 73755 (AA), May 1936 (shrub 15 ft.).—Jah-kuang, Che-li
Hsien, in thickets, alt. 1300 m., C. W. Wang 79106 (AA), Sept. 1936.—
Kuen-ger, Che-li Hsien. in thickets, alt. 1260 m., C. W. Wang 79272 (AA),
Oct. 1936. — Ban-chiou-chian, Che-li Hsien, fhickets: alt. 840 m., C. W. Wang
79763 (AA), 79770 (AA), Oct. 1936.— Meng-la, Jenn-yeh Hsien, mixed
forest, alt. 1000 m., C. W. Wang 80498 (AA), 80599 (AA), Nov. 1936.—
Kuan-yeang, Luh-shuen Hsien, mixed forest, alt. 1180 m., C. W. Wang 81161
(AA), Nov. 1936.— Without precise locality: G. Forrest 9615 (AA), 27783
(AA).

KWEICHOW: Tuh-shan, in dense shade, Y. Tsiang 6980 (AA), Sept. 8,
1930

KWANGSI: Shang-sze District, southeast of Shang-sze, Kwangtung border,
Shap Man Tsai Shan, near Iu Shan village, W. T. Tsang 22227 (AA), May
7, 1933.

KIANGSI: Lungnan District, Oo Chi Shan, near Lam Uk Village, steep
slopes in rocky forest, S K. Lau Yov7 (AA), Oct. 1934 (tree 20 ft.).

UPPER BURMA: Taung-gyi, F. G. Dickason 5880 (AA), May 5, 1933.—
Toungu District, Thandoung, alt. 3200 ft., F. G. Dickason 6791 (AA), Jan.
1938 (shrub 15 ft.).— Haka, semi-open hillside, alt. 6000 ft., ”. G. Dickason
ha (AA), 7695 (AA), Apr. 1938 (tree 30 ft.). — Kachin Hills, Shaik Mohkim

n. (AA), Jan. 1898.— Ken Tung [Kiangtung] Territory, Muang Len Ridge,
7 "V. Rock 2027 (AA), Jan. 24, 1922.— Fort Stedman, Abdul Huk s.n. (AA),
Jan. 1893 (tree 100-150 ft.).— Madre Hills, King’s Coll. 174 (AA), Feb. 20,
1893 (tree 80 ft.).

LOWER BURMA: Amherst District, Kyain, common in scrub eae
F. G. Dickason 6882 (AA), Feb. 1938 (small tree 30 thd:
in Zadi Circle, Jaungbyant forest, Maung Po In 5611 CAD. Feb. D5, 19 906.

SIAM: Phu Krading, Loie (N. E. Siam), common in open forests at summit,
alt. 1000 m., Native Collector DE329 (Roy. For. Dept. 4418) (AA), August
19, 1950.

84 JOURNAL OF THE ARNOLD ARBORETUM [voL. XXXII

INDO-CHINA: Laos: Sam-neua, EF. Poilane 2074, Nov. 14, 1920.

Anneslea fragrans is a very variable species extending from southern
China through Burma and Indo-China into Malaya. Several species
have been described in the genus since Wallich first introduced the
original A. fragrans.

After a careful study of material of this genus from all regions, one
cannot help conclude that, with the exception of the isolated A. don-
naiensis and A. steenisii, only a single species is represented, with
regional variations — and these regional varieties not consistently dis-
tinct. A similar situation may be found in the single Asiatic species of
Cleyera, namely C. japonica.

The center of distribution for the genus, as well as the species A.
fragrans, appears to be Yunnan and Burma. Even in this area con-
siderable variation may be found in leaf size and shape and pedicel
leneth, as well as in number of cells in the ovary and the number of
stigmas. It was on these characters that the species had been separated.

On the fringes of the distribution area may be found the regional
varieties. On Formosa is var. lanceolata. This variety was later given
specific status, but in a treatment by Keng in 1950 it was again
recorded as a variety. In this variety the leaves are consistently
narrower than the species and lanceolate, the peduncle short, and the
fruit reportedly smaller. Narrow-lanceolate leaves may be found in
the Yunnan material, but only occasionally. The same is true of the
pedicel length.

An interesting observation on peduncle length can be found in a
series of specimens colected by Tsai in Yunnan. Tsai’s numbers 55768,
55629, and 53142 were collected on the same day, and the peduncles
consistently measure 5.5-6 cm. in length. His numbers 56840, 56846,
and 56722 were collected in the same locality and in these numbers
the peduncles measure 3.5, 3.5, and 3.5-6 em. in length. Handel-
Mazzetti 6164 has fruiting peduncles measuring 2.5 em. Study shows
that the peduncles of a length of 5 and 6 cm. on the typical material,
although outstanding, are actually less frequently found than those of
the shorter measurements.

In Malaya the specimens are of a much sturdier appearance through-
out. The peduncle is outstanding because of its short, thick character,
as it measures as much as 5 mm. in diameter at the apex. This variety
is well named crassipes.

Two closely allied varieties, hainanensis and ternstroemioides, are
found in Hainan and Tonkin respectively. These are characterized by
having smaller flowers on shorter pedicels. However, var. ternstro-
emioides has also much smaller leaves and shorter petioles. Gagnepain
considered this variety a distinct species because of the two-celled
ovary and two-lobed stigma. These characters are not distinctive. I
have seen Gagnepain’s type of A. ternstroemioides and feel that it is
worthy of varietal status only.

1952] KOBUSKI, THE GENUS ANNESLEA 85

The variety alpina is not a marginal or regional variety. It was
collected in Yunnan and reported by the collector, Yi, as common. In
his use of the word “common” Yii was probably referring to material
typical of the species. Still only a single specimen of typical material
seems to have been collected by him. Anneslea alpina was based on
a very poor specimen. However, its rounded leaves and short petioles
cannot be overlooked. Although specimens from Burma show leaves
similar to var. alpina, the leaves on the Yii specimen are all rounded,
while on the Burmese material the rounded leaves are only occasional.

Three specimens, one each from Kwangsi, Kweichow, and Kiangsi,
are cited under the species. These three specimens increase the specific
range somewhat along the southern area of China. The Kwangsi
specimen, 7'sang 22227, has lanceolate leaves very similar to those of
var. lanceolata.

Anneslea fragrans Wallich _ Sea (Hooker ex Choisy) Pierre,
Flor. For. Cochinch. 2: ¢. . 1887.
Anneslea crassipes Hooker ex sine in Mém. Soc. Phys. Hist. Nat. Genéve
: (Mém. Ternstr. 41). 1855.— Dyer in Hooker f., Fl. Brit. India
1: 280. 1872. — Keng, Mater. Fl. Malay Penins. 1: 127. 1890.— Ridley,
Fl. Malay Penins, 1: 193. 1922.— Melchior in Nat. Pflanzenfam. ed.
2, 21: 143. 1925
Anneslea monticola Kurz in Jour. As. Soc. Bengal 42(2): 59. 1873; For.
Fl. Brit. Burma 1: 98. 1877.—- Mason, Burma People Prod. (Bot.) 2:
630. 1883.
Anneslea fragrans Wallich 8 monticola (Kurz) Pierre, Flor. For. Cochinch.
2: t. 127. 1887.
Daydonia crassipes (Hooker ex Choisy) Britten in Jour. Bot. 26: 11. 1888.
Anneslea phe Hooker ex Choisy var. obovata King, Mater. Fl. Malay
Penins. 1: 127. 1890.
Mountnorrisia rates (Hooker ex Choisy) Szyszylowicz in Nat. Pflanzen-
fam. IIT. 6: . 1893.

MALAYA: ae ng: Cameron Highlands, Rhododendron Hill, alt. ca.
4800 ft., MW. R. Henderson 23321 (AA), April 2, 1930.— Cameron Highlands,
alt. ca. 4000 ft., M. R. Henderson 32671 (AA), April 16, 1937 (48 ft. to first
branch). Penang: Hooker s.n. (G) in 1851. Perak: summit of Gunong
Batu Pateh, alt. 6700 ft., L. Wray 315 (AA). Without precise
locality: Griffith 741 (G) and A. C. Maingay 181 (G).

This variety is characterized by an over-all sturdiness and is well
named crassipes. Most characteristic is the stout sturdy pedicel meas-
uring sometimes only 1.5-2 em. long. In fruiting specimens these
pedicels may measure as much as 5 mm. in diameter (Wray 315). In
Henderson 23321 and 32671, two specimens collected in the same
locality, variation in the pedicel character may be found. The earlier
number has much thicker pedicels, especially at the apex, than 32671.
The latter number appears more closely related to the species.

Anneslea fragrans Wallich var. lanceolata Hayata, Icon. Pl. Formos.
3: 42, t. 5. 1913.— Kanehira, Formos. Trees 55, fig. 1917.—

86 JOURNAL OF THE ARNOLD ARBORETUM [voL. XXXII

Melchior in Nat. Pflanzenfam. ed. 2, 21: 143. 1925.— Makino &
Nemoto, FI. Jap. 736. 1931.— Yamamoto in Sylvia 5: 32. 1934.
— Keng in Taiwania 1: 254. 1950.

Anneslea lanceolata (Hayata) Kanehira, Formos. Trees, ed. rev. 455, fig.

412. 1936.

FORMOSA: Hunchuen Peninsula: Matsuda 16781 (photo, AA);
R. Kanehira 7 (AA); S. Sasaki W10 (AA), Oct. 31, 1920.

This variety is separated from the species by its smaller fruits
(ca. 1 cm. diam.), shorter pedicels, 2.5-3 cm. long, and consistently
lanceolate to oblong-lanceolate leaves 10-13 em. long and ca. 3 cm. wide.

The latest and probably the best treatment of this variety was
furnished by Keng in Taiwania 1: 254. 1950. Unfortunately Keng
compared this variety with material from Hainan only, thus dealing
with A. fragrans var. hainanensis rather than with the species. In the
latter named variety from Hainan, the fruit is larger, as are the leaves.
However, both varieties are characterized by shorter pedicels.

The narrow lanceolate leaves are not distinctive of this variety
alone, since material from Yunnan and Kwangsi show the same type
of leaf. The leaves of Forrest 9615 measure ca. 9 & 2.5 em. and are
all similar. However, on T'sai 53142 there are lanceolate leaves meas-
uring ca. 12 3.5 em., very similar to those of var. lanceolata. On the
same specimen may be found wider leaves, less acute at the apex and
more typical of the species. Both the above specimens were collected
in Yunnan. Tsang 22227, collected in Kwangsi, bears leaves similar
to this variety, measuring 10 * 2.8 em. and 8.5 x 2.2 cm. Others.
however, measure 11 X 4 em. All are acute at the apex.

VERNACULAR NAME: Nagaba-mokkoku (Japanese).

Anneslea fragrans Wallich var. hainanensis, var. nov.

A varietate ternstroemioides differt foliis maioribus, ad 15 em. longis
et 5-6 cm. latis, obovatis, apice rotundatis vel obtusis rare late
acuminatis, petiolis ad 3 cm. longis; filamentis 2-3 mm. longis, antheris
4 mm. longis, acumine ca. 2 mm. longo; ovario 2- vel 3-loculato,
stigmatibus 2 vel 3.

HAINAN: Dung Ka to Wen Fa Shi, in thickets, alt. 2000 ft., NV. K. Chun
& C. L. Tso 43792 (AA), 1932-33 (tree 20 m.; seeds red). — Po-ting, in forest,
alt. 2800 m., F. C. How 72915 (AA), June 16, 1935 (tree 10 m. with gray
bark; leaves dark green above, pale green beneath; fruit yellow-green). —
Without precise locality: H. Y. Liang 64168 & 64223 (AA); C. Wang 35183
(AA)

KWANGTUNG: Wung Yuen District: Fan Shiu Shen, steep slopes of
rocky forest, S. K. Lau 2743 (AA), Nov. 1933 (tree 10 m.).

This variety is characterized by small lanceolate leaves 4.5-7 em.
long, 2-3 em. wide, with the petioles ca. 1 em. long, the pedicels short,
2-3 em. long, the filaments short, 2-3 mm. long, and the ovary usually
two-celled and the stigmas two in number.

Except in the shape and size of the leaves this variety very much

1952] KOBUSKI, THE GENUS ANNESLEA 87

resembles var. ternstroemioides from Indo-China. Both varieties have
smaller flowers on shorter pedicels than those found in typical <A.
fragrans and the other varieties. In the stamens of both varieties the
filaments are very short (not over 2-3 mm.) appearing almost bulbous
in shape, while in other members of the species the filaments are
usually approximately 5 mm. long.

Anneslea fragrans Wallich var. ternstroemioides (Gagnepain), comb.

Annestea ternstroemioides Gagnepain in Notulae Syst. (Paris) 10: 116.
1941; in Fl. Gén. Indo-Chine Suppl. 1: 278. 1943.

hen omen Massif du Tam-dao, alt. ca. 1400 m., A.
Pételot 3869 (vypE of A. ternstroemioides, Paris), Dec. 1930

This variety is characterized by lanceolate leaves far smaller than
those of the species or any of the other varieties, being 4.5—7 em. long,
2-3 em. wide with a petiole seldom measuring over 1 cm. long. The
pedicels are short (2 em. long). The filaments are very short (ca.
1 mm. long) and only one-fifth the length of the anthers. The ovary
is two-celled and the stigmas number two.

It is most closely allied to var. hainanensis, which differs in having
larger leaves (up to 15 em. & 6 cm.) with petioles as much as 3 em.
long. However, in the inflorescence there is close agreement between
the two varieties.

The two-celled ovary and two stigmas of var. ternstroemioides may
not prove to be consistent characters. Only a single specimen, the type,
has been collected to date

Originally described by Gagnepain as A. ternstroemioides, it was
separated from A. fragrans primarily on the two stigma lobes, the
shorter sepals, and the size of the leaf. I find the short bulbous fila-
ments and the short-petioled leaves more distinctive characters, and
it is on the basis of these that I separate it from A. fragrans as a variety.

VERNACULAR NAMES: Brol, Cay la chua, Reung, Ko nang na.

Anneslea fragrans Wallich var. alpina (Li), ON nov.

Anneslea alpina 11 in Jour. Arnold Arb. 25: 307.

YUNNAN: Mhien-ning, Po-shang, common in eh alt. 2700 m., T. T.
Yu 18031 (type of A. alpina, AA), Oct. 11, 1936 (shrub 8-12 ft. high with
pinkish red flowers).

This variety can be separated from the species by its smaller ovate
to suborbicular leaves (4-6 em. long, 3-4 em. wide) with the short
petioles only 5 mm. long.

The specimen cited above, Li’s type for A. alpina, is very poor. No
flowers were attached and all pedicels were broken off. The exact
measurements could not be given. Li’s measurement of 1-1.5 mm. is
a mechanical error, of course. He probably meant to say 1-1.5 em.
Even this is not correct, since one pedicel (in packet) measures over
2 cm. long, and this one is broken off at the base.

88 JOURNAL OF THE ARNOLD ARBORETUM [VoL. XXXII

The small ovate leaves are rather distinct from those of other speci-
mens from this region which have been studied. Dickason 7587 from
Upper Burma has rounded leaves 5 X 2.5 em. with petioles 1 cm. long.
However, on the same specimen are longer leaves, more acute, and
typical of the species. On the Dickason specimen the pedicel measures
over 5 cm. in length.

2. eee steenisii, sp. nov.

Arbor ?; ramulis verticillatis subverticillatisve, brunneis vel griseis,
teretibus, pire foliis coriaceis, paucis ad apicem ramulorum dis-
positis, glabris, oblongo-ellipticis vel obovatis, apice acutis vel obtusis,
basi attenuatis, subtus punctatis, margine subrevolutis, subintegerrimis,
costa supra canaliculata, subtus elevata, venis ca. 8-10 paribus con-
spicuis vel subconspicuis, supra leviter impressis, subtus elevatis,
petiolis 1 em. minusve. Flores ad apicem ramulorum leviter spirali-
terque dispositi; pedicellis crassis, recurvatis, 1.5-2 em. longis, ca.
4 mm. diametro, glabris, teretibus, rare ancipitibus ; bracteolis 2,
oppositis suboppositisve, subaequalibus, crasso-coriaceis, subrotundatis
vel late deltoideis, 3 &* 3 mm., 4 & 4 mm., ad 6 6 mm., glabris;
sepalis 5, imbricatis, inaequalibus, subrotundatis, Regen exterioribus
duobus apaselcriliis. 7-12 mm. longis, 9-12 mm. latis, interioribus
tribus leviter latioribus, ad 15 & 15 mm., erp plus minusve
scariosis; petalis 5, basi leviter connatis, obovatis, 12-13 mm. longis,
7-10 mm. latis, margine integris vel subintegris non constrictis; stami-
nibus ca. 35, 2-seriatis, glabris, apice in apiculum 1 mm. longum
projectis, filamentis basi ad corollam adnatis, crassis 3-4 mm. longis,
antheris elongatis, ca. 4-5 mm. longis; ovario glabro, subplano, 2—(vel
3-) loculari, stylo glabro, attenuato, ca. 1.3 em. longo, apice 2—(vel 3-)
partito, non recurvato. Fructus globosi, glabri, sublepidotis obtecti,
4—5 em. longi, ad 3 em. diametro, apice persistentibus sepalis coronati,
2—(vel 3-) loculares, duobus vel tribus seminibus in quoque loculo;
seminibus ca. 1 em. longo, 3-4 mm. diametro.

SUMATRA: Atjeh: Gajolanden: Poetjoek Angasan, common in the
ridge forest above Penosan, alt. 2300 m., C. G. G. J. van Steenis 8327 (Herb.
Bogor.), Jan. 27, 1937.— Mt. Losir, on watershed between bivouacs 4 and 5,
near stream in the forest, alt. 2700-2800 m., Van Steenis 8493 (Herb. Bogor.),
Jan. 31, 1937.— Mt. Kemiri, in scrub at the summit on east side of Camp
at Aloer, alt. 2850-3300 m., Van Steenis 9685 (Herb. Bogor., type), March
10, 1937. — Near junction of Kapi and Aoenan Rivers near Paja camp, flat
forest ridges near sulphur field, Van Steenis 9964 & 9981 (Herb. Bogor.),
March 21, 1937.

Included in a loan of specimens of Adinandra and Ternstroemia
from Herbarium Bogoriense were the five specimens cited above. The
arrival of this material considerably upset the rest of this paper, which
had already been prepared for publication. Until these specimens came
to light the genus was supposed to occur only as far south as the Malay
Peninsula. From a general examination it was thought at first that

1952] KOBUSKI THE GENUS ANNESLEA 89

the above Sumatran material would prove to be only an extension of
the range of A. fragrans, perhaps another variety with very large fruit.
However, characters discovered in the flowers, as well as those in the
fruit, showed that a new species should be designated.

The petals in A. steenisii, like those of A. donnaiensis, lack the “hour-
glass” shape of A. fragrans but are obovate without the single con-
striction along each side. Also the petals are joined only lightly at the
base rather than connate for 5-7 mm. as in A. fragrans, and measure
only approximately one-half the length of those of the mainland
species. The stamens measure slightly less in length than those of
A. fragrans. In A. steenisit the partitions of the style, although either
two or three in number, are erect rather than spreading as in A.
fragrans.

The fruit of this species is far larger than any previously seen for
the genus, measuring as much as 5 em. in length and 3 em. in diameter.
The walls of the fruit are very thick, measuring 5 mm. or more. The
seeds, one centimeter or more in length, are correspondingly large.

In the mature fruit examined (Van Steenis 9981), the bracteoles
had disappeared, only the scars remaining. The measurement of the
scar showed the bracteoles at full maturity to have measured as much
as 9 mm. across at the base. In general the bracteole measurements
even in the young flowers were considerably more than those found
in A. fragrans. The above description of the fruit was drawn from
Van Steenis 9981.

It is a pleasure to name this species after C. G. G. J. van Steenis, the
collector and my close friend.

VERNACULAR NAME: Kajoe gaboe.

3. Anneslea donnaiensis (Gagnepain), comb. nov.
Paranneslea donnaiensis Gagnepain in Bull. Soc. Bot. France 95: 29. 1948,
as alensis.

Trees 20-30 m.; branchlets terete, subverticillate, glabrous, gray.
Leaves coriaceous, glabrous, subverticillate, few, disposed at the apex
of the branchlets, obovate to oblanceolate, 7-15 em. long, 3-5 em. wide,
obtuse at the apex, bluntly acuminate, attenuate at the base, slightly
decurrent, the margin entire, the midrib canaliculate above, elevated
below, the veins 6-8 pairs inconspicuous on both surfaces, the petiole
2-2.5 em. long. Flowers solitary, axillary, subverticillate; peduncles
terete, glabrous, 3-3.5 cm. long, somewhat thickened at the apex:
bracteoles 2, persistent, opposite, immediately below the calyx, un-
equal, broadly ovate to deltoid, ca. 3 mm. long, 2-3 mm. wide; sepals 5,
imbricate, unequal, glabrous, obovate to rounded, the outer ones 5 mm.
long, the inner ones 15 mm. long; petals 5 [fide Gagnepain], orbicular,
strongly concave, 12 mm. diameter in the bud; stamens very numerous,
free, 7 mm. long, glabrous, the anthers basifixed, 5 mm. long. linear,
bi-mucronate at the apex, 2-celled, the cells unequal, the filaments
short, 2 mm. long, flattened; the ovary immersed at the base in the

90 JOURNAL OF THE ARNOLD ARBORETUM [voL. XXXIII

torus, pyramidal, 5 mm. long, 2-celled, 5-lobed at the apex, the stigmas
[5] obtuse, erect, the ovules 2 to each cell, pendulous from the top of
the placenta. Fruit baccate, subinferior, free only at the apex, other-
wise joined with the calyx-tube, leathery, usually pustular dotted,
globular or rounded, 2 cm. diameter, crowned by the persistent sepals,
2-celled, each cell 2-seeded. Seeds with a woody testa, covered with
(reddish ?) arillae.

INDO-CHINA: Annam: Prov. Ht. Donnai; between Dang-kie and
Yonglé, FH. Poilane 23425 (AA), January 28, 1934.— Braian, near Djiring, alt.
1200 m., H. Poilane 24470 (Paris, synrypPe of Paranneslea donnaiensis), Feb.
22. 1935.

A syntype of Paranneslea donnaiensis was borrowed from the her-
barium of the Paris Museum in the hope of studying the floral struc-
ture. However, since only a single flowering bud was found attached to
the specimen, I decided to rely on Gagnepain’s notes for the above
description of petals, stamens, and ovary, rather than remove the bud.

Gagnepain’s description was obviously drawn from the bud rather
than from open or mature flowers. The petals may be similar in shape
to those of either A. fragrans or A. steenisii. I assume from the de-
scription that “orbicularia . . . in alabastro 12 mm. diam.” refers to
the shape and size of the bud rather than the individual petals. No
reference to a style of any sort was made by Gagnepain. Gagnepain
seems dubious about the number of ovules in each cell, citing two.
This ee may vary, possibly, as it does in A. | Re and A,
steenisi

pee the above cited fruiting specimen (Poilane 23425), there
is no obvious character to remove it from A. fragrans except the soli-
tary arrangement of the fruit in the axils of the leaves. This same
specimen was available to Gagnepain in the Paris Herbarium, since he
earlier cited it (Fl. Gen. Indochine Suppl. 1: 278. 1943) as A. fragrans.

Up to the present time the fruit has not been described. In the above
description, details for the fruit were taken from Poilane 23425, a speci-
men collected close to the type locality.

All the fruits dissected for this study showed two well-developed cells
with two mature seeds in each cell measuring ca. 12 mm. in length and
5-7 mm. across. As mentioned earlier in this paper, the bi-mucronate
anthers, the five pistils, and the solitary axillary pedicels constitute
the principal differences separating this species.

The specific name appeared only once in Gagnepain’s publication
and was printed “donnalensis.” Since the name is obviously derived
from the Province Ht. Donnai, I presume the spelling donnalensis is a
mechanical error, and I have here corrected it to donnaiensis.

ARNOLD ARBORETUM,
Harvarp UNIVERSITY.

1952] VAN ROYEN & VAN STEENIS, ERIANDRA 91

ERIANDRA, A NEW GENUS OF POLYGALACEAE
FROM NEW GUINEA!

P. vAN Royen anp C. G. G. J. vAN STEENIS

With one plate

AMONG THE UNIQUE COLLECTIONS made by Mr. L. J. Brass on the
Archbold Expeditions to New Guinea, we found a most remarkable
plant which had been sent to this herbarium under the provisional
name Siderorylon. An examination of the characters convinced us that
it is closely related to Diclidanthera, Moutabea, and Barnhartia, three
genera of the Polygalaceae known only from South America. From a
plant-geographical standpoint this is most remarkable, but this type
of distribution is by no means unique though still exceptional and
interesting.

The group of American genera has a chequered taxonomical history
which we need not recall here, since it has been done previously by
other authors (O’Donell, Erdtman, Sandwith & Sprague).

In comparing the four genera it appears that they show a reticulate
affinity, i.e., their conformity is different when different characters are
chosen for comparison.

The four genera are either trees (Hriandra) or shrubs, while the
leaves are spirally arranged ? and entire. The inflorescences are either
axillary or terminal. In comparison with the other three genera
Kriandra is pauciflorous.

The leaves of Eriandra have a light greenish yellow colour when
dried, reminiscent of some species of Nanthophyllum, which influenced
the senior author in giving as sight determination: “a Xanthophyllum
with regular flowers.” This yellowish colour is often due to the presence
of a certain amount of aluminium, and since according to Chenery
(1948) the other three genera belong to the aluminium-containing
plants, it is reasonable to assume that Eriandra belongs to the same
group.

In the absence of glands on the leaves, petioles, bracts, and bracteoles,
Eriandra differs from the other three genera.

The calyx in Eriandra is 4- or 5-merous and 5-merous in the other
three genera. The corolla is 5-merous in the four genera, actinomorphic
in Diclidanthera or subactinomorphie in Eriandra, subzygomorphie in
Barnhartia, and zygomorphic in Moutabea. The zygomorphy in

—

* Results of the Richard Archbold Expeditions.

*The term “alternate” as used by Sprague & Sandwith to indicate the spiral
phyllotaxis in Barnhartia seems less preferable in botanical terminology. In all
four genera of the Moutabeae the leaves are spirally: arranged.

92 JOURNAL OF THE ARNOLD ARBORETUM (VoL. XXXII

Moutabea is partly due to the boat-shaped lower petal and the zygo-
morphic androecium. The subzygomorphy in Barnhartia is determined
by the absence of two or three stamens and to the (very weak) union
of four petals in two pairs. Gleason (1926) points out that the slight
connation of the paired petals is facilitated by their approximation on
the margin of the hypanthium away from a normal position alternate
with the sepals; thus one might conclude that a slight zygomorphy is
also found in this shifting of the petals.

The petals in Eriandra, Moutabea, and Diclidanthera are united into
a tube and are free in the apical parts only, but in Barnhartia the five
petals are free, four of them cohering in two pairs. In Moutabea, how-
ever, the tube is deeply incised at the dorsal side.

In Eriandra, Diclidanthera, and Moutabea the eight or ten stamens
are united into a single column which is united with the corolla, but
in Barnhartia the seven or eight stamens are inserted on the petals but
never united into a tube. In Moutabea this tube is open at the dorsal
side.

In the four genera the anthers dehisce with one tangential slit, a
character which is otherwise not found in the Polygalaceae.

On the structure of the pollen we have a report by Dr. G. Erdtman,
Director of the Palynological Laboratory at Stockholm, to whom we
sent some material and who, in 1944, examined the pollen of Diclidan-
thera. He pointed out that the pollen closely resembled that of
Xanthophyllum, Salomonia, and Polygala. We investigated the pollen
grains of Barnhartia floribunda Gleason, and as our drawing shows,
it is closely related to Diclidanthera and to Eriandra. The pollen grains
of Eriandra fragrans are slightly constricted at their equator, and in
the opinion of Dr. Erdtman this character may be seen as an evidence
in favour of the distinction of a new polygalaceous genus. His pollen
diagnosis, which he kindly put at our disposal, runs: “pollen grains
8-9-colporate (zonate), prolate spheroidal (30x 28,) slightly con-
stricted at their equator. Sexine probably thicker than nexine; OL
pattern (faint; can be seen at least near the equator) .”

The ovary of Eriandra is 7- or 8-celled, that of Diclidanthera 5-celled
(fruit 5-7(!)-celled), Moutabea 4- or 5-celled (fruit 2-5-celled), and
that of Barnhartia 2- or 3-celled. In all cases the cells contain one
pendulous ovule.

The styles are densely pubescent in Eriandra, Barnhartia, and
Diclidanthera, but glabrous in Moutabea.

The stigmas are capitate, papillate in Hriandra, Diclidanthera, and
Barnhartia, but subquinquelobate to bilabiate and slightly infundi-
buliform in Moutabea, as is clearly shown in Miquel’s drawing in the
Flora Brasiliensis 7: pl. 5, f. 13 and 14. The authors found a bilabiate
stigma.

According to Chodat (1897) and Oort (1932) a disk seems to be
present in Moutabea. However, neither in Aublet’s description (1775)

1952] VAN ROYEN & VAN STEENIS, ERIANDRA 93

nor in the one given by Miquel (1856) is there a disk represented. We
could not find the slightest indication of this character.

Considering the details given above, the proper place of Eriandra
is in the Polygalaceae in the affinity of Barnhartia, Moutabea, and
Diclidanthera, to the last of which it is most closely related. The four
genera are best placed together in the tribe Moutabeae. Arranged
according to affinity with the tribe Polygaleae Barnhartia seems to
come first, followed by Moutabea, whilst Eriandra and Diclidanthera
show the least relationship. The four genera form a series of genera
running from zygomorphic to actinomorphic and from choripetalous to
gamopetalous flowers. Related to this last character is the uniting of
the stamens, which are free in Barnhartia and form a staminal tube
in Diclidanthera and Eriandra. Moutabea forms an intermediate stage,
as the eight stamens are united into two bundles, though still in one
tube which is open at the dorsal side.

As the characters of the Moutabeae given by Chodat (1897) have
to be emendated to include the four genera, we propose to give the
following definition of this tribe:

Tribe Moutabeae

Calyx and corolla united at the base on a torus. Calyx united.
Corolla free, cohering or united, quincuncial in bud. Stamens 7, 8 or
10, united into a tube or free and inserted on the free petals, sometimes
in two bundles. Anthers dehiscing with one tangential slit. Carpels
2-8, united, with one ovule in each cell. Trees or shrubs with entire,
spirally arranged leaves.

Four genera with ten species, in South America and New Guinea.

KEY TO THE GENERA

1. Stamens 8 or 10, connate in a tube.
2. Calyx 5-merous, quincuncial in bud. Anthers glabrous. Ovary 5-celled.
Base of bracts and base of leaf-margin provided with a gland ......
iclidanthera Martius.
2. Calyx 4-merous, decussate or 5-merous, quincuncial. Anthers densely
pubescent. Ovary 7- or 8-celled. Glandless........ randra nov. gen

1. Stamens 7 or 8, either in 2 bundles or not connate in a tube.

3. Flowers markedly zygomorphic. Corolla tubular, splitting at the dorsal
side. Upper 2 stamens absent, the others in two bundles of 4. -Anthers
transversely ellipsoidal. Ovary 4- or 5- celled, style glabrous, stigma
sub-5-lobed and subinfundibuliform or bilabiate. Leaf-blade and petiole
PIANOS 25555. bees dak eee ey weno Moutabea Aublet.

3. Flowers subzygomorphic. Petals connivent, not united. Stamens inserted
on the petals, not united into a tube. Anthers longitudinally ellipsoidal.
Ovary 2- or 3-celled, style densely pubescent, stigma discoid-capitate.
Apex of the petiole with a gland on either side. .... Barnhartia Gleason.

94 JOURNAL OF THE ARNOLD ARBORETUM [voL. XXXII

Eriandra gen. nov.

Arbor parva, foliis spiraliter ordinatis petiolatis integris; racemis
axillaribus parvis paucifloris; floribus epigynis actinomorphis subzy-
gomorphisve; sepalis 4 vel 5 decussatis vel quincuncialibus, basi con-
natis; petalis 4 vel 5 pro 3/4 longitudinis connatis in tubum calyci
adnatum, apice tantum liberis; partibus liberis decussatis vel imbricatis
orbicularibus; staminibus 8 vel 10 coalitis in tubum corollae adnatum,
nune uno latere paululo exsculptum; antheris transverse ellipsoideis,
fissura communi transversa dehiscentibus; valvis 2, dense longeque
pilosis; pollinis granulis in 8-vel 9-colporatis; ovario globoso glabro,
7-vel 8-loculari; loculis omnibus uniovulatis; stylo dense piloso; stig-
mate discoideo-capitato, papillato; fructu adhuc ignoto.

Typus: E. fragrans.

Eriandra fragrans sp. nov.

Arbor parva corona umbrosa instructa; trunco irregulariter et pro-
funde suleato (truncum spurium Ficorum epiphyticarum nonnullarum
in mentem revocante): ramulis glabris; internodiis 0.5-2.5 cm. longis;
foliis ellipticis oblongisve basi anguste cuneatis apice acutis 15-20 cm.
longis 5-7 em. latis coriaceis, utraque facie subnitentibus et glabris
sed in facie inferiore costae mediae interdum pilis paucis raris con-
spersis; costa media subtus prominente, in facie superiore basi sub-
canaliculata, apice prominente; nervis lateralibus utroque latere costae
mediae 12-16, subtus distincte prominentibus, supra prominulis;
margine revoluta; petiolo supra applanato, parte inferiore rugoso,
1-2.5 em. longo; floribus albis fragrantibus; racemis circ. 1 cm. longis;
pedunculo communi breviter piloso, cire. 6 mm. longo; bracteis brac-
teolisque cymbiformibus obtusis; bracteis dense pilosis circ. 1 mm.
longis; bracteolis subglabris circ. 0.5 mm. longis pedicellis 1-5 mm.
longis, glabris; sepalis spathulatis obovatisve, 4-5.5 mm. longis,
utrimque glabris, exterioribus margine toto, inferioribus parte apicali
tantum fimbriatis; partibus petalorum liberis cire. 3 mm. longis,
praeter margines fimbriatos glabris, in sicco rubiginosis; tubi staminei
membranacei parte libera extus glabra, intus pilosa; antheris circ.
0.8 mm. latis; pollinis granulis plus minusve 30» longis, ca. 28 crassis;
ovario plus minusve 2 mm. diametiente; stylo 3-4 mm. longo.

Type: L. J. Brass 7767 in L, duplicate in A.

PAPUA: Western Division, Lake Daviumbu, Middle Fly River, rain forest,
Brass 7767 (TYPE in L; duplicate in A), Sept. 1936 (common small canopy
tree; whole length of trunk deeply indented Ane flanged, like some strangling
figs. Flowers white, fragrant).

NETHERLANDS NEW GUINEA: Mt. Arfak, Putat, Beccari 9928 (in
herb. Firenze), anno 1872 (loose flowers).

The name Eriandra is chosen on account of the pubescent anthers,
while the specifie epithet is used because of the fragrant flowers.

1952] VAN ROYEN & VAN STEENIS, ERIANDRA 95

BIBLIOGRAPHY

AvBLET, F.: Pl. Guy. Fr. 2: 679-680; 4: 274. 1775.
BAILLON, H.: Hist. Pl. 5: 76-77. 1874.
CHENERY, E. M.: in Kew Bull. 1948: 175. 1948.
CuHopat, R.: in Mém. Soc. Phys. Hist. Nat. Genéve. 1891, 1893.
———: in Engler & Prantl, Nat. Pfl. Fam. 3, 4: 323-345, 1897.
ERDTMAN, G.: in Bot. Not. 1944: 80-S4. 1944.
Gra, E.: in Bot. Jahrb. 40: Beibl. 93: 81. 1908.
———: in rat & Gilg, Syllabus Pfl. Fam. ed. 9/10, 323. 1924.
Gueason, H. A.: in Bull. Torr. Bot. Cl. 53: 297-299. 1926.
HALuirr, ce Ueber Juliania etc. 46: 193. 1908.
: in Meded. Rijksherb. Leiden 1: 36. 1910,
———- in Arch. Neéerl. Sc. Exact. Natur., sér. = B, 1: 73. 1912.
Mravet, F. A. G.: in Mart. Fl. Bras. 7: 11-16, L 4-6. 1856.
O’DonELL, C. A.: in Lilloa 6: 207-212, pl. 1 d: 2. 1941.
Oort, A. J. P.: in Pulle, Fl. Suriname 27: 424-425. 1932.
SOLEREDER, H.: Syst. Anat. Dicotyledonen 110, 587-588. 1899.
SPRAGUE, T A. & SANDWITH, N. Y.: in Hooker's Icon: Pl., ser, 5, 2: ¢. $179,
pp. . 1932.

EXPLANATION OF PLATE
PLATE |

Eniandra fragrans Van Royen & Van Steenis (Brass 7767): 1. habit, X 1/3;
2. bud; 3. part of corolla from inner side, flattened; 4. longitudinal section
of flower (schematic); 5. flower with 2 sepals, corolla and androecium re-
moved; 6. base of flower with cross-section of ovary; 7. longitudinal section
of ovary; 8. stigma; 9. anther; 10. ditto, showing the vertical septum;
11-12. pollen grains; 13. basal part of flower in section; 14. detail of leaf-
nervation, underside; 15. stomatal apparatus. Barnhartia floribunda
Gleason: 16-17. pollen grains. Except in fig. 1 all details enlarged.

RIJKSHERBARIU
LEIDEN, ee any

Puate I

Jour. ARNoLD ArB. VoL. XX XIII

ERIANDRA FRAGRANS VAN ROYEN & VAN STEENIS

JOURNAL

OF THE
ARNOLD ARBORETUM
Vor. XXXII APRIL 1952 Numper 2

STUDIES OF PACIFIC ISLAND PLANTS, XI?
FURTHER NOTES ON FIJIAN FLOWERING PLANTS

A. C. SMITH

HE PRESENT PAPER CONCLUDES from No. VII of this series (in Jour.
Arnold Arb. 31: 288-319. 1950) a discussion of recently collected note-
worthy plants of angiosperm families in Fiji; here are included the families
from Rhizophoraceae to Compositae, in the Engler and Prantl sequence.
The description of a new species of Theaceae, kindly contributed by Dr.
C. E. Kobuski, is also here included. This paper completes the study of
the specimens collected by the writer in 1947,? with the exception of a
few families which are being studied more intensively and which will form
the subject matter of further reports in this series. The place of deposit of
cited specimens is indicated as follows: Arnold Arboretum (A); British
Museum (BM); Gray Herbarium (GH); Royal Botanic Gardens, Kew
(K); New York Botanical Garden (NY); and U. S. National Herbarium
(US),

THEACEAE
By C. E. Kosuskxt

Eurya greenwoodii Kobuski, sp. nov.

Frutex 2—3 m. altus; ramis ramulisque glabris, teretibus rubro-brunneis.
Folia subcoriacea vel submembranacea, glabra (gemmis basi leviter pubes-
centibus), ovalia vel ovata, 3—4.5 cm. longa, 1.5-2 cm. lata, apice brevi-
acuminata, leviter retusa, basi rotundata vel cuneata, margine plana basi
conspicue revoluta, costa supra canaliculata subtus elevata, venis 7 vel 8

*No. X of this series was published as Vol. 30, Part 4, of Contributions from the
U. S. National Herbarium (1952).

? Under the auspices of the Arnold Arboretum of Harvard University and the John
Simon Guggenheim Memorial Foundation, with the aid of grants from the Penrose
Fund of the American Philosophical Society and the Bache Fund of the National
Academy of Sciences.

98 JOURNAL OF THE ARNOLD ARBORETUM _ [VvoL. xxxmI

paribus ad marginem reticulatis, petiolis teretibus 3-4 mm. longis, glabris.
Flores axillares, 2—5-fasciculati erecti vel nonnumquam cernui; ¢ : ignoti;
2: pedicellis teretibus 2—2.5 mm. longis, pubescentibus; bracteolis a
oppositis, persistentibus, pubescentibus rotundatis vel subrotundatis circa
1 mm. longis et 1 mm. latis, apiculatis; sepalis 5, imbricatis, glabris,
inaequalibus, 1.5—-2 mm. longis, 2—2.5 mm. latis, concavis, margine scari-
osis; petalis 5, albis, late oblongis, 3.5-4 mm. longis, 2-2. : mm, oS
apice leviter retusis, basi 1% connatis; staminodiis 0; ov globos
glabro 1.5-2 mm. diametro 4-loculato (raro 3-loculato), loculis multi,
ovulatis; stylis 4, liberis, circiter 0.5 mm. longis, maturitate recurvatis.
Fructus ignotus.

Vitr Levu: Mba: Immediate vicinity of Nandarivatu, alt. 800-900 m., Ma
26, 1947, Smith 4898 (A Type, US) (“samu ni mbati”; compact shrub 2
high, in dry forest; petals white) ; same locality, Greenwood 236A (A) oe
branched shrub to 3 m. high; flowers white), Degener 14799 (A, US) (spreading
shrub abcut 1 m. high, in scrub vegetation).

This species is most closely allied to EF. vitiensis A. Gray, but can be
separated from the latter by the smaller glabrous oval or ovate leaves
(3-4 cm. long), and by the glabrous branchlets and terminal buds. In
E. vitiensis the obovate leaves are longer, measuring up to 7.5 cm. long,
and with pubescent petioles. The very young branchlets and terminal buds
are also pubescent. The three cited specimens of the new species bear only
pistillate flowers.

RHIZOPHORACEAE

Crossostylis pedunculata sp. nov.

Arbor gracilis ad 8 m. alta, ramulis validis, juvenilibus complanatis
pilis circiter 0.3 mm. longis stramineis copiose pilosis mox glabrescentibus,
internodiis 2—2.5 cm. longis; stipulis oblongo-lanceolatis circiter 13 mm.
longis et 4 mm. latis dorso basim versus puberulis ceterum glabris caducis;
petiolis rugulosis subteretibus 15—25 mm. longis ut ramulis decidue pilosis;
laminis in sicco fuscis chartaceis ellipticis, 10-13 cm. longis, 5—8 cm. latis,
basi obtusis et in petiolum longe decurrentibus, apice obtuse cuspidatis
saepe obscure emarginatis, margine basim versus integris superne crenatis
(crenaturis circiter 2 per centimetrum), praeter costam subtus ut petiolum
pilosam utrinque glabris, costa supra subplana subtus prominente, nervis
secundariis utrinsecus 8 vel 9 arcuato-patentibus utrinque leviter elevatis,
rete venularum irregulari utrinque subprominulo vel immerso; inflores-
centiis binis in axillis superpositis pedunculatis, pedunculo crasso subtereti
6-9 mm. longo ut petiolo breviter piloso; floribus ut videtur 2 per inflo-
rescentiam primo involucro (bracteis connatis) ovoideo circiter 9 mm.
longo parce piloso apice cuspidato unilateraliter fisso inclusis; pedicellis
sub anthesi 6-9 mm. longis’ stramineo-hispidulis; calyce carnoso rotato-
cupuliformi 8-10 mm. diametro extus parce strigilloso plerumque 5-lobato,
lobis oblongo-deltoideis circiter 4 mm. longis 2.5-3 mm. latis apice sub-

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XI 99

acutis et hispidulis; petalis mox caducis submembranaceis obovatis circiter
5 mm. longis et 2 mm. latis, basim versus angustatis, apice emarginatis et
inconspicue glanduloso-erosulis, praeter lineam medianam dorsalem hispi-
dulam glabris; disco carnoso perigyno margine libero et obscure hispidulo;
staminibus plerumque 20, filamentis gracilibus teretibus glabris 2—-2.5 mm
longis, antheris oblongis circiter 0.8 mm. longis; ovario subcomplanato
faciei superiore copiose sed minute hispidulo, stylo crasso circiter 1.5 mm.
longo glabro apice radiatim plurilobato, lobis reflexis filiformibus stigma-
tosis plus minusve coadnatis.

Vanua Levu: Thakaundrove: Natewa Bay region, hills west of
Korotasere, alt. 100-300 m., June 8, 1934, Smith 1925 (GH, NY, US 1676416
TYPE, etc.) (slender tree 8 m. high, in forest; calyx-lobes and filaments white).

The collection described above was originally identified by me as C.
seemanni (A. Gray) Schimp., and as such duplicates were distributed.
However, it clearly differs from C. seemanni in its less copiously pubescent
young parts and its glabrous stipules and leaf-blades, by having its flowers
borne at the apex of short stout peduncles, and in its larger flowers, which
furthermore have the sepals and petals only sparsely pilose and the fila-
ments comparatively long. In foliage the new species is suggestive of
C. biflora Forst., of the Society Islands (whether or not this includes the
Samoan form discussed by A. Gray, Bot. U. S. Expl. Exped. 1: 610. pl. 77.
1854, and Christophersen, in Bishop Mus. Bull. 128: 155. 1935), and
C. banksiana Guillaumin, from the New Hebrides. From these, however,
C. pedunculata differs in the pubescence of its young parts and flowers.
It further differs from C. biflora in its smaller flowers, with shorter fila-
ments and style and without staminodes or disk-lobes. Crossostylis bankst-
ana appears to have flowers smaller than those of C. pedunculata and
petals pilose within, but it is perhaps the closest ally of the new species.

Crossostylis pachyantha sp. nov.

Arbor parva ubique praeter flores glabra, ramulis subteretibus (juvenili-
bus subquadrangularibus) ; stipulis in sicco subcoriaceis lanceolatis 10-15
mm. longis; petiolis gracilibus 10-17 mm. longis margine leviter ae
laminis in sicco coriaceis fuscis elliptico-obovatis, 8-12 cm. longis, 3—5 cm.
latis, basi attenuatis et in petiolum longe fecurrenGbus. apice rotundatis
vel obscure emarginatis vel obtusis, margine integris, costa supra in
sulculam elevata subtus prominente, nervis secundariis utrinsecus 7—9
erecto-patentibus marginem versus curvatis et anastomosantibus supra
subplanis subtus leviter elevatis, rete venularum immerso vel subtus laxe
subprominulo; inflorescentiis apices ramulorum versus axillaribus breviter
pedunculatis, pedunculo crasso haud 3 mm. longo apice ut videtur flores
circiter 4 gerente, bracteis non visis; pedicellis validis glabris sub anthesi
5-8 mm. longis in sicco superne leviter angulatis, floribus apice circiter
7-8 mm. diametro; calyce crasso-carnoso rotato-cupuliformi extus glabro,
lobis plerumque 5 interdum 4 oblongo-deltoideis 3-4 mm. longis 2-3 mm.
latis apice subacutis intus pilis 0.1-0.2 mm. longis fulvis dense sericeo-

100 JOURNAL OF THE ARNOLD ARBORETUM _ [VoL. xxxmI

tomentellis; petalis mox caducis carnosis valde carinatis obovatis circiter
3.5 mm. longis et 2 mm. latis, basi valde angustatis, apice emarginatis,
utrinque basim et medium versus puberulis; disco carnoso perigyno glabro
margine haud libero; staminibus 21-26, filamentis crassis 0.5-0.7 mm
longis glabris, antheris oblongis circiter 0.7 mm. longis; gynoecio glabro,
ovario subcomplanato sub anthesi ad 3 mm. diametro inconspicue 10—12-
lobato, ovulis numerosis (ut videtur 20-24) geminatim radiatis; stylo
crasso (ad 1 mm. diametro) tereti circiter 1.5 mm. longo apice radiatim
plurilobato, lobis filiformibus stigmatosis in peltam carnosam ad 2 mm.
diametro coadnatis.

Bile Levu: Naitasiri: Near Tholo-i-suva, alt. about 200 m., Feb. 23
947, J. H. Vaughan 3370 (BM type) (small tree, with dark, shiny foliage and
pone flowers) ; Nasinu, Gillespie 3650 (GH).

In general appearance and leaf-shape, the species here described could
be taken for a form of C. richit (A. Gray) A. C. Sm., although it is more
robust in vegetative parts than any available specimens of C. richit,
which now appears to be the most abundant species of the genus in Fiji.
In inflorescence characters, however, the two species are quite different,
C. pachyantha being at once distinguished by the thick texture of its
much larger calyx, the dense indument of the inner surface of calyx-lobes,
the numerous stamens and ovules, the glabrous ovary with 10-12 radiating
sulcae, and the stout style with numerous stigmatic branches.

COMBRETACEAE

Terminalia vitiensis A. C. Sm. in Sargentia 1: 74. 1942.

ANUA Levu: Mathuata: Southern slopes of Mt. Numbuiloa, east of
Lambasa, alt. 350-500 m., Smith 6441 (A, US) (tree 5 m. high, in thin forest on
rocky slope; calyx, filaments, and style pale yellowish green).

The cited specimen, the second known of the species, agrees excellently
with the type, from the Province of Serua on Viti Levu. The only points of
difference are minor; the Mathuata specimen has the leaf-blades rarely
larger, up to 10 & 4.5 cm., the rachis and pedicels sparsely pilose rather
than glabrous, the pedicels longer (up to 7 mm. long), and the filaments
and style up to 15 mm. in lengt

Terminalia luteola sp. nov.

Arbor multiramosa ad 8 m. alta, partibus novellis pilis aureis 0.6—-1 mm.
longis copiose hispidulo-sericeis, ramulis infra folia apice congesta copiose
cicatricosis glabratis demum cinereis apicem versus ad 8 mm. diametro;
petiolis semiteretibus 0.7—-2 cm. longis ut ramulis juvenilibus copiose
patenti-pilosis, laminis chartaceis siccitate fusco-olivaceis late obovatis,
8-12 cm. longis, 5—8 cm. latis, basi obtusis in petiolum breviter decurrenti-
bus, apice rotundatis vel leviter emarginatis, margine integris, utrinque ut
petiolo subpersistenter pilosis (pilis pallidis costa nervisque magis copiosis) ,
costa supra paullo subtus valde elevata, nervis lateralibus utrinsecus 8—10

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XI 101

erecto-patentibus marginem versus curvatis utrinque elevatis, rete
venularum laxo utrinque prominulo; inflorescentiis axillaribus spicatis
sub anthesi 6-11 cm. longis, pedunculo ad 3 cm. longo et rhachi gracilibus
pilis mollis 0.3-0.6 mm, longis copiose tomentellis; floribus numerosis
congestis sessilibus bracteolatis, bracteolis reflexis oblongo-linearibus 3—5
mm. longis utrinque copiose tomentellis; calyce infundibuliformi 6-8 mm.
longo extus pilis 0.3-0.4 mm. longis copiose et molliter tomentello, tubo
2-3.5 mm. longo 2—2.5 mm. diametro basi truncato, limbo tenuiter carnoso
4-5 mm. longo intus pilis 1-1.3 mm. longis pallidis sericeo, lobis deltoideo-
lanceolatis circiter 2 mm. longis acutis; lobis disci 5 carnosis copiose
hispidulis; staminibus 8-11, filamentis filiformibus sub anthesi 7-9 mm.
longis glabris superne angustatis, antheris oblongo-ellipsoideis circiter
1 mm. longis, loculis ad medium liberis; stylo in floribus § tereti carnoso
8-11 mm. longo glabro superne contracto, in floribus ¢ rudimentario;
ovulis 2 a funiculis gracilibus pendulis.

Vanua Levu: Mathuata: Southern slopes of Mt. Numbuiloa, east of
Lambasa, alt. 350-500 m., Oct. 29, 1947, Smith 6409 (A Type, US) (tree 8 m.
high, in transitional zone between reed-covered slopes and forest; filaments and
style yellowish green; anthers yellow).

Although the new species was collected in the same locality as T.
vitiensis A. C. Sm., noted above, it is not closely related to that species,
being immediately distinguished by its dense indument, larger leaves,
sessile flowers, etc. Terminalia litoralis Seem. has leaves somewhat re-
sembling in shape those of the new species, but they are glabrous or
essentially so and the inflorescence is also glabrous and has small pedicellate
flowers.

MELASTOMATACEAE
Medinilla subviridis sp. nov.

Frutex scandens, partibus novellis et ramulorum apicibus copiosissime
tomentosis, pilis pallide ferrugineis 1-2 mm. longis multicellularibus
lateraliter brevi-calcaratis, ramulis elongatis gracilibus teretibus vel
superne inconspicue sulcatis mox glabratis cinerascentibus, internodiis
2-4 cm. longis; foliis valde disparibus minoribus mox caducis; foliis
majoribus: petiolis leviter canaliculatis 5-15 mm. longis ut ramulis
juvenilibus tomentosis demum glabratis, laminis chartaceis in sicco fusco-
olivaceis oblongo-ellipticis, 10-15 cm. longis, 3.5—7 cm. latis, basi obtusis
vel rotundatis raro subattenuatis vel leviter cordatis, apice obtusis vel
breviter cuspidatis (apice ipso haud 1 cm. longo), 5- vel 7-nerviis,
nervis superioribus cum costa ad 1-2 cm. conjunctis supra planis vel
interdum leviter sulcatis subtus valde elevatis, nervis aliis debilioribus,
venulis transversis subtus prominulis aliis immersis, primo ut ramulis
pilosis demum supra glabratis subtus certe nervis subpersistenter tomen-
tellis; foliis minoribus: petiolis subnullis, laminis suborbicularibus vel

102 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxu1

ellipticis, 1-2.5 cm. longis latisque, basi rotundatis saepe amplexicaulibus,
apice rotundatis vel obtusis, inconspicue 3- vel 5-nerviis, demum glabratis;
inflorescentiis axillaribus vel e ramulis infra folia orientibus solitariis
cymoso-racemosis, 2—5 cm. longis paucifloris (floribus in verticillis 1-3
dispositis); pedunculo brevi (haud 1 cm. longo) et rhachi teretibus
gracilibus pilis pallide brunneis circiter 1 mm. longis copiose tomentosis
demum subglabratis; bracteis 3 vel 4 e nodis papyraceis oblongis, 3-7 mm
longis, 1.5-3 mm. latis, basi et apice obtusis, primo ut rhachi pilosis
dvmum glabratis; floribus 3 vel 4 e nodis, pedicellis gracilibus 3-5 mm.
longis sub anthesi ut rhachi copiose pilosis; bracteolis apice pedicellorum
binis florem obtegentibus submembranaceis deltoideo-ovatis, 12-20 mm.
longis, 9-15 mm. latis, basi et apice rotundatis, utrinque molliter pilosis
(pilis patentibus 0.3-0.7 mm. longis breviter calcaratis) demum subglabra-
tis; calyce sub anthesi circiter 5 mm. longo et apice diametro carnoso,
tubo cupuliformi 2.5-3 mm. longo basi rotundato et supra bracteolas
minute stipitato ut bracteolis dense tomentoso (pilis ad 1 mm. longis),
limbo suberecto membranaceo subintegro vel minute 4-dentato; petalis 4
membranaceis glabris oblongo-obovatis, 7.5-8.5 mm. longis, 6-7 mm.
latis, basi angustatis, apice rotundatis et paullo retusis, obscure nervatis;
staminibus 8, filamentis gracilibus glabris circiter 3 mm. longis, antheris
oblongis 1.7-2 mm. longis, basi trilobulatis (lobis 2 anterioribus haud
projectis, lobo posteriore circiter 0.4 mm. diametro), apice obtusis, poris
confluentibus; stylo carnoso tereti circiter 5 mm. longo, stigmate minuto;
fructibus subglobosis demum glabratis, calycis limbo persistente, pericarpio
tenui, seminibus numerosis.

Vitt Levu: hci : Northern portion of Rairaimatuku Plateau, be-
tween Mt. Tomanivi [Mt. Victoria] and Nasonggo, alt. 870-970 m., Sept. 18,
1947, Smith 6112 A TYPE, US) (liana, in dense forest; bracts and ‘bracteoles
pale greenish white or very faintly pink-tinged; calyx white with pale brown
pubescence; petals, filaments, and style pure white; anthers rich blue with bright
yellow basal lobes; fruit rich purple); same locality, Aug. 21, 1947, Smith 5793
(A, US) (high-climbing liana, in dense forest; bracts and bracteoles thin, very
pale green, with brown puberulence; calyx white, similarly puberulent; corolla
pure white; filaments and style white; anthers rich purple with bright yellow
basal lobes; young fruit white to pale green, then blue-tinged).

The species here described, observed at only one locality, in central Viti
Levu, is without close relatives, at least in Fiji and the adjacent archi-
pelagos. In general type of indument and in bracteole-size the new species
approaches M. longicymosa Gibbs, but differences in foliage, length of
trichomes, color of inflorescence-parts, size and shape of bracts, shape of
bracteoles, and dimensions of petals and stamens are so great that com-
parison of the two species is superfluous. A closer ally of M. subviridis
may be M. rhodochlaena A. Gray, which suggests it in leaf-shape, type of
trichomes, proportions of -bracts and bracteoles, and flower-size, but
M. rhodochlaena has rich pink bracts and bracteoles, the latter being
only 4-11 mm. in diameter.

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XI 103
Astronidium degeneri A. C. Sm. in Sargentia 1: 93. 1942.
Vitt Levu: Mba: Northern portion of Mt. Evans Range, between Mt.

Vatuyanitu and Mt. Natondra, alt. 700-900 m., Smith 4359 (A, US) (tree 13 m.
high, on edge of forest); mountains near Lautoka, alt. about 550 m., Greenwood
(A).

The second and third known collections of this species agree very
closely with the type, which was obtained at lower elevation near the
south coast of Viti Levu.

ARALIACEAE
Plerandra pickeringii A. Gray, Bot. U. S. Expl. Exped. 1: 729. pl 95.
4.

This striking species is the common sole or sole ndina (i. e. the true sole)
of the Fijians. It is readily recognized by its very large leaves (with petioles
up to 1 meter long, the 9-12 leaflets up to 40 x 18 cm. or more) and
robust compound umbels (with 10-15 rays, each as much as 30 cm. long
and with 30-60 flowers). Gray’s original description of the staminate
flowers is more or less correct, except for the reference to “stamens . . . as
many as 60 or 70.” Actually the stamens, in both staminate and perfect
flowers, are between 200 and 300, crowded in 5-7 series; the type specimen
(U. S. Expl. Exped., US 62359) also bears this out. In both staminate
and perfect flowers the ovary seems well developed, with 13-16 locules,
but in the former case the ovules are abortive and the stylopodium is
inconspicuous, truncate, and depressed at the center; in perfect flowers
the stylopodium is elongate (up to 1 cm. long in fully developed fruits)
and composed of styles firmly connate to the apex. Among my recent
collections the species is represented by nos, 4021 and 5803 (staminate
flowers) and 6896 (perfect flowers and fruits). I do not find that this
species has the petals connate into a calyptra, a character emphasized
by Seemann (FI. Vit. 117. 1865); in most specimens the petals tend to
separate before falling, but I do not believe the degree of coherence to be
a reliable character.

Plerandra insolita sp. nov.

Arbor gracilis ad 9 m. alta pauciramosa ubique glabra, ramulis crassis
teretibus fistulosis, foliis apices ramulorum versus aggregatis magnis;
petiolis crassis subteretibus interdum copiose verrucoso-lenticellatis 40-50
cm. longis, basi in vaginam coriaceam 2—4 cm. latam dilatatis, ligula
oblonga 1.5-2 cm. longa; foliolis 7-12 (plerumque 9-11), petiolulis
crassis semiteretibus vel leviter canaliculatis 3.5—5.5 cm. longis, laminis
in sicco subcoriaceis et fusco-olivaceis oblongo- vel anguste obovato-
ellipticis, 20-35 cm. longis, 5—9 cm. latis, basi acutis vel attenuatis, apice
obtusis vel rotundatis, margine integris et anguste recurvatis, costa valida
utrinque prominente, nervis secundariis utrinsecus 15—25 erecto-patentibus
curvatis utrinque elevatis, rete venularum immerso vel subtus prominulo;

104 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxuI

inflorescentia pro genere compacta e ramulis infra folia oriente composito-
pseudoumbellata, radiis 5-7 (4—7 cm. longis) apice pedunculi brevis (1-5
cm. longi) dispositis, pedunculo et radiis inflorescentiae crassis (saepe
1 cm, diametro) conspicue verrucosis vel lenticellatis, radiis infra apicem
incrassatis et ibi fructiferis vel copiose cicatricosis apice vel in centimetrum
distale ie umbellulis sub anthesi circiter 5 cm. diametro, floribus
circiter 15-20 (¢ solis visis) bracteis subtentis, bracteis subcoriaceis
oblongis ad 15 <7 mm. apice rotundatis scarioso-marginatis longe
persistentibus; floribus ¢ sub anthesi subsessilibus vel pedicellis (paullo
post anthesin) ad 12 mm. longis crassis; calyce sub anthesi 7-9 mm.
longo obconico-cupuliformi, tubo tereti vel leviter angulato, limbo sub-
coriaceo erecto-patente lobis 5 vel 6 late deltoideis circiter 1 mm. longis
obtusis inclusis circiter 2 mm. longo; petalis 5 vel 6 (raro 4) in calyptram
a subacutam videtur subconnatis crasso-carnosis deltoideis, 7—8 mm.
longis, 4-6 mm. latis, apice valde incrassatis et subacutis; staminibus
numerosis (circiter 75) disco carnoso annulari 2- vel 3- seriatis, filamentis
paullo ante anthesin ligulatis superne angustatis gracilibus 2-3.5 mm.
longis, antheris oblongis 2—2.5 mm. longis utroque obtusis; stylopodio
breviter conico vel umbonato obtuso carnoso 4-5 mm. diametro margine
lobato sulculis 9-12 radiatis ornato; ovario sterili 7-8 mm. diametro,
muro valde incrassato fistulis numerosis longitudinalibus copiose notato,
loculis 9-12 magnis, ovulis nullis vel sterilibus; fructibus e radiis
inflorescentiae 2-3 cm. infra apicem enatis, pedicellis sub fructu crassis
ad 15 mm. longis, fructibus coriaceis obovoideo-ellipsoideis ad 2.5 cm.
longis et 1.8 cm. diametro conspicue 9—12-sulcatis, calycis limbo persistente
et stylopodio coronatis, stylopodio oblongo-umbraculiformi circiter 1 mm
longo et 3 mm. diametro, stylis firme connatis apice radiatis sed haud
liberis, pyrenis 9-12 coriaceis falcato-obovoideis valde complanatis ad
20 mm. longis et 7 mm. latis, semine forma simili circiter 13 & 5 mm.

Vitr Levu: Mba: Southern slopes of Mt. Ndelainathovu, on the escarpment
west of Nandarivatu, alt. 870-970 m., June 26, 1947, Smith 4922 (A Typr, US)
(“sole lailai”; slender tree 3-4 m. high, in dense forest; leaves terminal, with
9-11 leaflets; inflorescence arising just below leaves, composed of 5—7 umbels;
petals deep purple without, pale green within; filaments white; anthers yellow;
fruit black); Serua: Mbuyombuyo, near Namboutini, Tabualewa 15584 (A,
US, etc.) (‘sole’’; sparingly branched tree 9 m. high, in gine Vatuvilakia,
vicinity of Ngaloa, alt. 0-150 m., Degener 15172 (A, U tc.) (“sole’; un-
branched tree 1-2 m. high, in open forest; petals purplish ae without, green-
ish within).

The species described above is readily distinguished from the other
Fijian Plerandrae by its compact inflorescence, of which the rays are not
strictly umbelliform, the flowers being crowded upon the distal portion of
each ray instead of arising from its actual apex. The three available
collections are essentially identical in floral characters, all the available
flowers being staminate. Fruits are borne from a swollen portion of the
inflorescence-ray somewhat below the apex, indicating the polygamo-

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XI 105

monoecious character of the species; other species of the genus, in my
observation, are polygamo-dioecious. In its large number of ovary-locules
P. insolita suggests P. pickeringii A. Gray and P. grayi Seem., from both
of which it differs in the characters mentioned above, the short-pedicellate
and comparatively small flowers, the very thick ovary-walls traversed by
copious longitudinal canals, etc.

EPACRIDACEAE

Leucopogon cymbulae Labill. Sert. Austro-Caled. 36. pl. 39. 1824;
Seem. in Bonplandia 10: 36. 1862, Fl. Vit. 147. 1866; Guillaumin in
Bull. Soc. Bot. Fr. 74: 699. 1927, in Jour. Arnold Arb. 13: 11. 1932,
in Bull. Soc. Bot. Fr. 82: 350. 1935.

Leucopogon vitiensis A. Gray ex Seem. in Bonplandia 10: 36, nomen. 1862.

Vit1 Levu: Mba: Upper slopes of Mt. Koromba [Pickering Peak], alt. 800—
1075 m., Smith 4700 (A, US) (tree 5 m. high, on forested crest; fruit orange).
Kanpavu: Seemann 285 (GH). Vanua Levu: Mbua: H. B. R. Parham, Jan.
7, 1937 (BM); Mathuata: Wainikoro, Greenwood 703 (A) (shrub about
3 m. high, on dry open hillsides; flowers white; fruit yellow and finally red);
between Ndreketi and Nasorowangga, B. E. Parham 1094 (A); Seanggangga
Plateau, in drainage of Korovuli River, vicinity of Natua, alt. 100-200 m.,
Smith 6667 (A, US) (“seruserumasala”; shrub or tree 2-4 m. high, frequent in
open rolling country; corolla white; fruit dull orange; wood used in making small
hand-drums) ; summit ridge of Mt. Numbuiloa, east of Lambasa, alt. 500-590 m.,
Smith 6493 (A, US) (tree 3-5 m. high, in dense forest, locally abundant; corolla
white; fruit at length orange). Fiji, without definite locality: U. S. Expl. Exped.
(GH, US, source of the name L. vitiensis), Horne 654 (GH), H. B. R. Parham
14 (A).

The only previously published records of the occurrence of this genus
in Fiji appear to be those of Seemann, cited above; Guillaumin has reported
it from several islands in the New Hebrides, and it is frequently mentioned
in literature pertaining to New Caledonia, the type locality. In the sense
of Brongniart & Gris (in Ann. Sci. Nat. Bot. V. 2: 153. 1864), L. cymbulae
is a variable species in New Caledonia. In general, the material from Fiji
and the New Hebrides has the leaves longer and proportionately narrower
than that from New Caledonia, but I find no inflorescence differences of
note. Possibly the Fijian and New Hebrides specimens should receive
varietal recognition, but at present this seems inadvisable and I am
willing to follow Seemann and Guillaumin in their identifications.

As the above citations show, the species is rare in Fiji except in northern
Vanua Levu; in Mathuata it is a very frequent component of the
“talasinga” vegetation, but curiously there appear to be no collections of
it from the similar zone of northern and western Viti Levu. The cited
specimen from Mt. Koromba was obtained in a forested highland which
interrupts the “‘talasinga’”’ of western Viti Levu.

106 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxx

MYRSINACEAE

Maesa stenophylla sp. nov.

Frutex ad 4 m. altus ubique glaber, ramulis gracilibus teretibus fuscis
lenticellis pallidis copiose verrucosis; petiolis gracilibus canaliculatis 8—12
mm. longis, foliorum laminis papyraceis siccitate fusco-viridibus lanceolatis,
7-12 cm. longis, 1.7—2.5 cm. latis, basi attenuatis et in petiolum decurrenti-
bus, apice obtusis, margine integris vel undulatis et anguste revolutis,
costa supra elevata subtus prominente, nervis lateralibus utrinsecus 4—6
valde adscendentibus elongatis supra planis vel prominulis subtus leviter
elevatis, lineis nervilliformibus numerosis irregularibus subtus manifestis
supra obscuris; inflorescentiis axillaribus racemosis vel anguste paniculatis
sub anthesi 2.5—4 cm. longis, pedunculo subnullo et rhachi gracilibus,
bracteis lanceolatis acutis 0.5—1 mm. longis, pedicellis gracilibus 1.5—2 mm
longis; floribus 5-meris circiter 2 mm. longis et apice 2.5 mm. diametro;
prophyllis liberis deltoideo-lanceolatis 0.5-0.8 mm. longis dorso parce
lepidotis; calyce circiter 1.5 mm. longo, tubo obconico minute lepidoto,
lobis deltoideis circiter 0.7 mm. longis subacutis non glanduloso-lineatis;
corolla campanulata circiter 1.5 mm. longa inconspicue glanduloso-lineo-
lata, lobis ovatis circiter 0.9 mm. longis latisque apice rotundatis margine
erosulis; staminibus basim corollae versus insertis, filamentis gracilibus
0.1—0.2 mm. longis, antheris deltoideo-ovoideis 0.3-0.4 mm. longis; ovario
levi semigloboso-conico, stylo brevi crasso tereti 0.2-0.3 mm. longo,
stigmate obscure bilobato, ovulis circiter 10 biseriatis; fructibus immaturis
ovoideo-ellipsoideis inconspicue luteo-glanduloso-lineolatis circiter 2.5 mm,
diametro, calycis lobis et stigmatibus persistentibus.

Vanua Levu: Mathuata: Northwestern slopes of Mt. Numbuiloa, east of
Lambasa, alt. 350 m., Nov. 6, 1947, Smith 6490 (A Type US) (“kolonimbeka”;
shrub 4 m. high, locally frequent in hillside thickets; corolla pale yellow with
salmon-pink markings; filaments pale green; anthers yellow

From its closest ally, M. persicaefolia A. Gray, the new species differs in
its narrower leaves with sharply ascending secondaries, its simpler and less
obviously lepidote inflorescence, and its slightly larger flowers with less
prominent glandular markings. Maesa neriifolia Gillespie seems very close
to M. persicaefolia but has even more conspicuously glandular-lineolate
floral parts.

Tapeinosperma chloranthum sp. nov.

Arbor ad 12 m. alta inflorescentiis exceptis glabra, partibus novellis
minute puberulis mox glabratis, ramulis cinereis plerumque teretibus ad
nodos incrassatis robustis superne cicatricosis, ramulis foliiferis saepe
gracilibus; petiolis sat robustis semiteretibus 10-15 mm. longis, laminis
chartaceis in sicco fusco-viridibus immerso-glandulosis anguste oblongo-
ellipticis, 10-20 cm. longis, 4—7.5 cm. latis, basi in petiolum attenuatis et
longe decurrentibus, apice obtusis vel anguste rotundatis, margine integris

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XI 107

anguste recurvatis, costa valida supra subplana subtus prominente, nervis
lateralibus primariis utrinsecus circiter 18-22 patentibus utrinque cum rete
venularum prominulis; inflorescentiis apices ramulorum versus axillaribus
paniculatis sub anthesi ad 6 cm. longis latisque, pedunculo brevi et rhachi
ramulisque validis subteretibus copiose tomentellis (pilis cinnamomeis
0.2-0.4 mm. longis multicellularibus saepe irregulariter ramulosis eglandu-
losis), bracteis oblongis 2—3 mm. longis subacutis intus glabris; pedicellis
crassis teretibus 2.5-5 mm. longis ut inflorescentiae ramulis cum calyce
dense tomentellis superne incrassatis; calyce cupuliformi 3.5-4.5 mm.
longo et apice circiter 5 mm. diametro textura subcarnoso immerso-nigro-
glanduloso intus glabro profunde lobato, lobis 5 (raro 6) ovatis subacutis
circiter 2 mm. longis latisque; corolla crasso-carnosa infundibulari-rotata
sub anthesi circiter 6 mm. longa et apice diametro, tubo 1-2 mm. longo,
lobis 5 immerso-nigro-glandulosis late imbricatis ovatis circiter 4 & 3 mm
obtusis; staminibus corollae faucibus insertis, filamentis carnosis ad 0.4
mm. longis, antheris deltoideo-ovoideis circiter 1.2 mm. longis obtusis
dorso glandulis inconspicuis ornatis; gynoecio sub anthesi circiter 3 mm.
longo, ovario ovoideo minutissime et pallide glanduloso-puberulo in
stylum gracilem attenuato, stigmate subpeltato circiter 0.5 mm. diametro,
placenta ovoidea apice subacuta, ovulis 3; inflorescentiis sub fructu
majoribus ad 20 cm. longis, pedunculo ad 3.5 cm. longo et ramulis
pedicellisque subpersistenter tomentellis, pedicellis superne valde in-
crassatis, calyce demum rotato ad mm. diametro extus persistenter
tomentello; fructibus subglobosis ad 7 mm. diametro stylo coronatis,
pericarpio coriaceo circiter 1 mm. crasso glandulis magnis atris ovalibus
copiose ornato, semine 1 subgloboso.

Vitt Levu: Mba: Hills between Nggaliwana and Nandala Creeks, south of
Nauwanga, alt. 725-850 m., Aug. 26, 1947, Smith 5820 (A Type, US) (tree 10
m. high, in dense forest; calyx brown; corolla dull green with purplish glands;
mature fruit red) ; slopes of Mt. Nairosa, eastern flank of Mt. Evans Range,
alt. 700-1050 m., Smith 4023 (“kutumirase”; tree 12 m. high, in dense forest) ;
eastern slopes of Mt. Koroyanitu, Mt. Evans Range, alt. 950-1050 m., Smith
4133 (A, US) (tree 10 m. high, in dense low forest)

From its closest ally, T. clavatum Mez, the new species is readily dis-
tinguished by the much more copious and persistent indument of its
inflorescence-branches and calyces and by its larger flowers with thick-
carnose infundibuliform corollas. Tapeinosperma greenwoodii A. C. Sm.,
known only from the Mt. Evans Range, where the new species also occurs,
has similar foliage but a more delicate and small-flowered inflorescence,
the indument being composed of very minute (less than 0.1 mm. long)
simple glandular hairs.

Tapeinosperma ligulifolium sp. nov.

Frutex ad 2 m. altus partibus novellis et inflorescentiis glanduloso-
puberulis exceptis ubique glaber, ramulis cinereis teretibus subvalidis;
foliis apices ramulorum versus confertis, petiolis crassis rugulosis 2-5 mm.

108 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxim

longis, laminis in sicco coriaceis olivaceis elongato-ligulatis, 20-32 cm.
longis, 1-1.5 cm. latis, basi obtusis, apice obtusis vel anguste rotundatis,
margine integris plerumque valde revolutis, utrinque inconspicue glandu-
losis, costa supra valde impressa subtus prominente, nervis lateralibus
primariis numerosis circiter 1 cm. distantibus brevibus patentibus intra
marginem anastomosantibus cum rete venularum inconspicue prominulis;
inflorescentia (unica visa) prope apicem ramuli axillari gracili multiflora
anguste paniculata circiter 12 cm. longa et 6 cm. lata, pedunculo circiter
5 cm. longo et rhachi ramulisque gracilibus subteretibus pilis haud 0.1 mm
longis parce glanduloso-puberulis, bracteis oblongo-ligulatis 1-1.5 mm.
longis obtusis extus glandulosis intus glabris; pedicellis sub anthesi 2—3
mm. longis ut inflorescentiae ramulis cum calyce glandulosis; calyce
rotato circiter 3 mm. diametro intus glabro profunde 5-lobato, lobis
lanceolatis 1—1.2 mm. longis et circiter 0.7 mm. latis subacutis minute
glanduloso-ciliolatis; corolla tenuiter carnosa paullo ante anthesin circiter
3.5 mm. diametro glanduloso-punctata, lobis 5 fere ad basim liberis
ovatis circiter 2.5 mm. longis latisque apice obtusis; staminibus prope
basim corollae insertis, filamentis circiter 0.5 mm. longis, antheris oblongo-
deltoideis circiter 1.2 mm. longis acutis obscure glandulosis; gynoecio
sub anthesi circiter 2 mm. longo, ovario oblongo minute glanduloso in
stylum crassum attenuato, stigmate minute subpeltato, ovulis ut videtur 2.

Vanua Levu: Mathuata: Summit ridge of Mt. Numbuiloa, east of Lam-
basa, alt. 500-590 m., Nov. 6, 1947, Smith 6522 (A TYPE) (shrub 2 m. high, in
dense forest; inflorescence-branches and perianth-parts rich pink).

This extraordinarily distinct species, represented by a single though
reasonably ample specimen, is at once distinguished by its remarkably
long and narrow leaf-blades, which are conspicuously revolute and
essentially sessile; its inflorescence is comparatively delicate, with slender
pedicels and narrow calyx-lobes. The relationship of T. ligulifolium is
probably with such species as T. grande (Seem.) Mez and T. megaphyllum
(Hemsl.) Mez, both with large narrow leaves, which, however, do not
approach in proportions those of the new species.

Discocalyx obtecta sp. nov.

Arbor vel frutex ad 6 m. altus, partibus novellis copiose ferrugineo-
puberulis, ramulis teretibus gracilibus cinereis mox glabratis ad nodos
incrassatis; petiolis subvalidis supra canaliculatis ad 1 cm. longis saepe
fere ad basim anguste alatis mox glabratis; laminis foliorum in sicco
chartaceo-subcoriaceis fuscis oblanceolatis, (S—) 7-12 cm. longis, (2—)
34.5 cm. latis, basi attenuatis et in petiolum longe decurrentibus, apice
obtusis, margine integris et leviter recurvatis, supra glabris, subtus copiose
glanduloso-punctatis et costa saepe pilosis, costa valida supra subplana vel
elevata subtus prominente, nervis lateralibus utrinsecus circiter 8—10
erecto-patentibus subtus prominulis vel interdum utrinque immersis, rete
venularum subimmerso; inflorescentiis apices ramulorum versus axillaribus
amplis paniculatis 6-11 cm. longis multifloris, pedunculo (ad 3.5 cm. longo)

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XI 109

rhachi ramulisque subteretibus striatis copiose puberulis (pilis brunneo-
ferrugineis glandulosis) demum subglabratis, bracteis oblongo-lanceolatis
2-3 mm. longis caducis, bracteolis minoribus; floribus paullo ante anthesin
subsessilibus (pedicello ad 1 mm. longo), calyce et pedicello pilis brunneo-
ferrugineis ad 0.1-0.2 mm. longis densissime glanduloso-tomentello-
puberulis; calyce obconico-pyriformi circiter 2.5 mm. longo et diametro
intus glabro, lobis 5 ovatis 1-1.3 mm. longis latisque acutis ciliis circiter
0.3 mm. longis copiose ornatis; corolla carnosa submatura videtur rotata
3-3.5 mm. diametro profunde 5-lobata, lobis oblongis circiter 1.5 mm
longis apice rotundatis immerso-glandulosis dorso parce pilosis alibi glabris;
staminibus immaturis, filamentis gracilibus haud 0.1 mm. longis, antheris
deltoideo-oblongis circiter 0.5 mm. longis apice obtusis; gynoecio glabro
in stylum gracilem circiter 0.6 mm. longum et stigmatem parvum peltatum
attenuato, ovulis ut videtur paucis; calyce sub fructu rotato circiter 3 mm.
diametro persistenter ciliolato et saltem inconspicue glanduloso-tomentello.
fructu subgloboso 5—6 mm. diametro stylo minuto coronato.

Vitt Levu: Mba: Hills east of Nandala Creek, about 3 miles south of Nan-
darivatu, alt. 870-970 m., Sept. 25, 1947, Smith 6217 (A Type, US) (tree 4 m.
high, in dense forest; flower-buds brown); hills between Nggaliwana and Tum-
beindreketi Creeks, east of the sawmill at Navai, alt. 725-800 m., Smith 6007
(A, US) (tree 6 m. high, in dense forest; young flowers cinnamon- -brown);
southern slope of Mt. Ndelainathovu, on the escarpment west of Nandarivatu,
alt. 870-970 m., Smith 4921 (A, US) (‘‘sila”; simple-stemmed shrub 2 m. high,
in dense forest: fruit red, at length purplish); Mt. Evans Range, alt. about 970
m., Greenwood 1149 (A) (tree 5-6 m. high, in dense forest; flower-buds brown).

Of the four available collections, my no. 4921 bears fruits and the others
immature flowers. The species is readily distinguished by the close and
copious glandular-tomentellous indument of the inflorescence-branches and
calyces. Among the described Fijian species, only D. divaricata Gillespie
has an obviously pilose inflorescence, and from that the new species differs
in its short petioles, its thick obovate leaf-blades narrowed at base, and its
flowers with shorter pedicels and pyriform calyces. Discocalyx divaricata,
a very graceful and attractive species, is quite frequent in upland Viti Levu,
being represented by my nos. 4250, 4526, 4823, and 6216. In foliage D.
obtecta more nearly resembles D. fusca Gibbs, but that species is essen-
tially glabrous throughout.

Discocalyx gillespieana sp. nov.

Arbor gracilis ad 4 m. alta partibus novellis et inflorescentiis obscure
glanduloso-puberulis exceptis ubique glabra, ramulis teretibus gracilibus
fuscis ad nodos incrassatis; petiolis semiteretibus rugulosis 10-17 mm
longis, laminis foliorum in sicco subcoriaceis fusco-olivaceis elliptico-
obovatis, 7—-11.5 cm. longis, 3-4.5 cm. latis, basi acutis et in petiolum
decurrentibus, apice rotundatis vel late obtusis, margine integris anguste
recurvatis, subtus parce glanduloso-punctatis, costa supra subplana subtus
prominente, nervis lateralibus utrinsecus 10-12 subpatentibus utrinque

110 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxx

prominulis vel interdum cum rete venularum obscuris; inflorescentiis apices
ramulorum versus congestis compacte paniculatis ad 3.5 cm. longis latisque,
pedunculo brevi rhachi ramulisque paucis gracilibus teretibus parce glandu-
loso-puberulis mox glabratis, bracteis oblongo-lanceolatis circiter 2 mm.
longis caducis; pedicellis gracilibus sub anthesi 1.5—-2 mm. longis sub-
glabris; calyce rotato sub anthesi 2—2.5 mm. diametro extus obscure
glanduloso-puberulo, lobis 5 (raro 6) deltoideo-oblongis circiter 0.7 mm.
longis obtusis margine ciliis glandulosis circiter 0.2 mm. longis copiose
ornatis; corolla carnosa rotata circiter 5 mm. diametro profunde 5-lobata,
lobis oblongis circiter 1.5 mm. longis latisque rotundatis immerso-glandu-
losis; antheris subsessilibus oblongo-ovoideis 0.7—0.8 mm. longis obscure
glandulosis apice obtusis; ovario sub anthesi conico 0.5—0.8 mm. diametro
inconspicue glanduloso, stigmate subsessili peltato circiter 0.7 mm. diametro
eroso-marginato, ovulis 3 vel 4.

Viti Levu: Nandronga & Navosa: Northern portion of Rairaimatuku
Plateau, between Nandrau and Nanga, alt. 725-825 m., Aug. 7, 1947, Smith 5546
(A typE, US) (“vutuvutu”; slender tree 4 m. high, in dense forest; calyx and
corolla deep purplish red; anthers and stigma pale yellow).

Discocalyx gillespieana is related to D. multiflora Gillespie and D. syl-
vestris A. C. Sm., but both of these have obvious styles, whereas in the new
species the stigma is essentially sessile. From D. multiflora the present
plant also differs in its longer-petiolate and proportionately broader leaves,
more compact inflorescence, smaller and less conspicuously glandular
corolla, and smaller anthers; from D. sylvestris it is further distinguished
by its differently proportioned leaves with more numerous secondaries. The
three species are certainly close relatives, but I believe that combinations
of characters and especially the stylar differences amply differentiate them.

EBENACEAE

Diospyros foliosa (Rich. ex Gray) Bakh. in Bull. Jard. Bot. Buitenzorg
IIT. 15: 447. 1941.

Maba foliosa Rich ex A. Gray in Proc. Amer. Acad. 5: 326. 1862; Seem. FI.
Vit. 152. 1866; Hiern in Trans. Cambr. Phil. Soc. 12: 113. 1873; Drake, III.
Fl. Ins. Mar. Pac. 230. 1892.

Vanua Levu: Mathuata: Southern slopes of Mt. Numbuiloa, east of
Lambasa, alt. 350-500 m., Smith 6439 (A, US) (“ulalo”; tree 8 m. high, in
rocky forest on steep hillside, the trunk 20 cm. diam.; fruit yellow, at length red;
seed edible), 6443 (A, US) (tree 10 m. high, in thin forest on rocky slope; fruit
yellow, at length red). Fiji, without definite locality except “Muthuata and
Ovolau” [i.e. Mathuata coast of Vanua Levu and island of Ovalau], U. S. Expl.
Exped. (GH, US 65907 type),

The three available Fijian collections agree in fundamental characters,
my no. 6439 being especially similar to the type. Number 6443, however,
differs in having its slightly narrower leaves copiously brown-tomentellous

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XI 111

beneath and its fruits shorter in proportion, nearly subglobose rather than
narrowly ellipsoid and about twice as long as broad (as in no. 6439 and
the type). These characters, which in the field I took to be significant,
probably merely indicate some of the variability to be expected in the
species. A few of the older leaves of no. 6443 have become merely canes-
cent-puberulent or nearly glabrous beneath, and some of the fruits vary in
shape toward ovoid-ellipsoid.

SYMPLOCACEAE

Symplocos turrilliana sp. nov.

Arbor ad 10°m. alta, partibus novellis et ramulis apices versus pilis
stramineis 0.5—0.8 mm. longis hispidulo-pilosis, ramulis teretibus rugulosis
brunneis demum glabratis; petiolis gracilibus leviter canaliculatis 1-3 cm.
longis ut ramulis juvenilibus pilosis, laminis foliorum in sicco chartaceis
flavo-viridibus oblongo- vel ovato-ellipticis, 6-11 cm. longis, 3-6 cm. latis,
basi rotundatis vel obtusis et in petiolum breviter decurrentibus, apice
breviter cuspidatis, margine inconspicue crenulatis, supra praecipue costa
pilosis mox glabratis, subtus pilis patentibus castaneis 0.3-0.7 mm. longis
praesertim costa nervisque subpersistenter hispiduloso-pilosis, costa supra
leviter canaliculata subtus prominente, nervis secundariis utrinsecus 5-8
erecto-patentibus curvatis anastomosantibus supra prominulis vel planis
subtus elevatis, rete venularum utrinque prominulo vel supra immerso;
inflorescentiis axillaribus compactis racemosis vel e basi 2- vel 3-ramosis
paucifloris, floribus inclusis haud 1.5 cm. longis, pedunculo brevi ramulisque
pilis stramineis 0.2-0.4 mm. longis dense sericeis, bracteis oblongo-semi-
orbicularibus circiter 1 mm. longis et 1.5 mm. latis rotundatis extus copiose
sericeis intus glabris; floribus subsessilibus, pedicello (haud 1 mm. longo)
et calyce ut inflorescentiae ramulis sericeis; calyce sub anthesi circiter 2.5
mm. longo et apice 4 mm. diametro, tubo cupuliformi circiter 1 mm. longo,
lobis 5 imbricatis, 3 exterioribus slain oblongo-ovatis obtusis 1.5—2
mm. longis latisque intus apic versus saepe sericeo-puberulis alibi
glabris, lobis interioribus ee haee ad 2.5 mm. longis extus mar-
ginem versus glabris; corolla membranacea sub anthesi rotata 7-8 mm
diametro glabra vel lobis extus obscure puberula, fere ad basim 5-lobata,
lobis oblongo-ovatis 3-3.5 mm. longis, 2—2.5 mm. latis apice rotundatis
interdum obscure eroso-ciliolatis; staminibus circiter 40-45, 2- vel 3-seri-
atis, filamentis ligulatis glabris sub anthesi 3-4 mm. longis, antheris sub-
globosis 0.30.4 mm. diametro; stylo tereti circiter 3 mm. longo, stigmate
obscure lobato; ovario infero; fructibus ovoideo-ellipsoideis immaturis ad
12 mm. longis puberulis vel glabratis, superne contractis, calycis lobis
persistentibus.

Vitt Levu: Mba: Summit of Mt. Nanggaranambuluta [Lomalangi], east of
Nandarivatu, alt. 1100-1120 m., June 23, 1947, Smith 4845 (A Type, US) (“kai
namo”; tree 5 m. high, in dense forest; petals and filaments white; anthers he
low) ; summit of Mt. Koroyanitu, high point of Mt. Evans Range, alt. 1165-

112 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxmI

1195 m., in dense ridge forest and thickets, Smith 4206 (A, US) (tree 10 m.
high; young petals white), 4222 (A, US) (tree 8 m. high; flower-buds white;
fruit green).

Turrill (in Jour. Linn. Soc. Bot. 43: 30. 1915), in describing the only
Fijian species of Symplocos as S. leptophylla (designated as a new species
but actually to be construed as a new combination based on S. stawelii var.
leptophylla Brand, both concepts being typified by Seemann 294, from
Kandavu), points out its variable nature. My 1947 collection contains 16
numbers, from Viti Levu and Vanua Levu, which may be referred to this
broad concept. Obvious variations are to be seen in the indument of the
calyx, which ranges from essentially glabrous to copiously sericeous, and
in the size and shape of leaves. I am inclined to consider this specific con-
cept too broad, but perhaps it should not be subdivided without careful
consideration of species from island groups to the west. Turrill has indi-
cated as forma compacta a plant from Nandarivatu with a reduced inflo-
rescence and coriaceous leaves, and probably some of my collections from
upland Viti Levu are referable to this form, although the value of the
mentioned characters is questionable. All the specimens thus far referred
to S. leptophylla have the leaves glabrous or merely minutely strigillose on
the costa beneath. The new species here proposed differs from S. lepto-
phylla in the obvious and persistent indument of its lower leaf-surfaces
and in the densely sericeous inflorescence-branches and calyx. As a rule in
S. leptophylla these inflorescence-parts are merely puberulent or minutely
strigillose and glabrescent, but occasionally the pubescence, in type and
persistence, approaches that of S. turrilliana.

OLEACEAE
Linociera vitiensis A. C. Sm. in Bull. Torrey Club 70: 549. 1943.

Fagraea vitiensis Seem. in Bonplandia 9: 257, nomen. 1861; non Gilg & Bene-
dict (1921).
Olea vitiensis Seem. Fl. Vit. 155. 1866.

Viti Levu: Seemann 307 (GH, type coll. of Olea vitiensis); Nandronga&

avosa: Northern portion of Rairaimatuku Plateau, between Nandrau and
Rewasau, alt. 725-825 m., Smith 5632 (A, US). Vanua Levu: Mathuata:
Southern slopes of Mt. Numbuiloa, east of Lambasa, alt. 100-500 m., Smith
6374 (A, US), 6580 (A, US).

The cited material agrees very well with the type of the species, Smith
864, from Taveuni. It had previously escaped my notice that Seemann’s
Olea vitiensis is actually a species of Linociera, and my inadvertent selec-
tion of the same specific epithet makes impossible a combination based on
Seemann’s earlier one. My more recent collections are from trees 8—20 m.
high, growing in dense or open forest; all have the fruit immature and
orange or dull orange-tinged. The local name teinivia was noted for
no. 5632.

1952 | SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XI 113
Linociera gillespiei A. C. Sm. in Bull. Torrey Club 70: 548. 1943.

Vitrt Levu: Naitasiri: Northern portion of Rairaimatuku Plateau, be-
tween Mt. Tomanivi [Mt. Victoria] and Nasonggo, alt. 870-970 m., Smith 5794
(A, US), 6135 (A, US).

The cited specimens, the second and third of the species known to me,
were collected in dense forest from trees 15-20 m. high, with green but
apparently nearly mature fruit; a local name recorded for no. 6135 is
thaunilawa. These two collections agree very well with the type, Gillespie
4289, from the vicinity of Nandarivatu.

It is curious that all eight collections of Linociera now available from
Fiji are in fruit, no flowering material having been obtained; this fact
makes adequate consideration of the two species impossible. It may be
seriously doubted that more than one species of Limociera occurs in Fiji,
but I cannot make the implied reduction at this time. In general, the
leaves of L. vitiensis are broader (about twice as long as broad) and thicker,
with often immersed veins; the leaves of L. gillespiei incline toward lanceo-
late in shape (about three times as long as broad), have more obvious
venation, and dry somewhat darker in color. The recent collections tend
to break down the distinctions between the species and, unless future col-
lections indicate the existence of stronger characters, the two entities may
be better combined.

LOGANIACEAE
Fagraea gracilipes A. Gray in Proc. Amer. Acad. 4: 323. 1860; Seem.
Fl. Vit. 165. 1866.
Fagraea viridiflora Seem. in Bonplandia 9: 257, nomen. 1861.

Vitt Levu: Serua: Vicinity of Ngaloa, Degener 15101 (A, US, etc.)
(“makamakandora”’; tree 5 m. high, on inner edge of mangrove swamp; tea from
bark and leaves used medicinally; timber considered durable and valuable).
OvaLau: Seemann 306 (source of the name F. viridiflora, GH, K). Vanua Levu:
Mathuata: Seanggangga Plateau, in drainage of Korovuli River, vicinity of
Natua, alt. 100-200 m., Smith 6665 (A, US) (‘mbuambua”; spreading tree to
10 m. high, in patches of forest in open rolling country, locally frequent but
seldom flowering; corolla and filaments cream-white or pale yellow; fruit .waxy
ivory-white; trunks used as houseposts). Fiji, without definite locality: U. S.
Expl. Exped. (GH, US 62265 type), Horne 1124 (GH).

The distribution and habitat of this beautiful endemic tree are more
fully indicated above than by the Exploring Expedition and Seemann
collections from which it has previously been understood. It is a rare plant
in Fiji and was observed by me only once, although in that locality, cited
above, it was a frequent component of the low dry forest that occurs on
certain reddish clay areas of northern Vanua Levu. The type specimen is
accompanied by a detached fruit which obviously represents the other, and
more abundant, species of Fagraea in Fiji, presumable F. berteriana A.
Gray (including F. vitiensis Gilg & Benedict). The mature fruit of
F. gracilipes is waxy white, ellipsoid, not much exceeding 2 by 1.5 cm.

114 JOURNAL OF THE ARNOLD ARBORETUM _[vot. xxxur

APOCYNACEAE

Pagiantha thurstonii (Baker) comb. nov.
Tabernaemontana thurstoni Horne, A Year in Fiji, 268, nomen. 1881; Horne ex
Baker in Jour. Linn. Soc. Bot. 20: 368. 1883; Burkill in Jour. Linn. Soc.
Bot. 35: 46. 1901; Gillespie in Bishop Mus. Bull. 74: 19. fig. 24. 1930.

Vitr Levu: Mba: Western and southern slopes of Mt. Tomanivi [Mt.
Victoria], alt. 850-1150 m., Smith 5089 (A, US) (‘‘nda’alu”; tree 10 m. high,
with abundant white latex, on edge of forest; flowers fragrant; corolla-tube
greenish, the lobes pure white); Nandronga & Navosa: Northern portion
of Rairaimatuku Plateau, between Nandrau and Rewasau, alt. 725-825 m.,
Smith 5638 (A, US) (“kau ndrenga”; tree 15 m. high, with abundant white
latex, in dense forest; pericarp yellow within, the seeds red); Naitasiri:
Nanduna, alt. about 30 m., B. E. Parham 1049 (A) (“tandalo”; tree 30 m.
high, with large white flowers).

The proposed combination (with a change to the termination -ii
as recommended by the International Rules) is necessary if one follows
Markgraf’s revision (in Notizbl. Bot. Gart. Berlin 12: 540-552. 1935)
of the Asiatic genera allied to Tabernaemontana. Pagiantha koroana
Markgraf (in op. cit. 549, 1935, in Bishop Mus. Bull. 141: 129. fig. 65,
h-k. 1936) is almost certainly the same species. Markgraf apparently
overlooked the binomial Tabernaemontana thurstoni, as indicated by his
remark that: “The present specimen is the first collection of a thick-
flowered species of Tabernaemontana from Fiji.” Gillespie’s description
and plate of 7. thurstoni are very adequate; he cites several collections
from Viti Levu and Taveuni, which, added to previously collected material,
indicate that the species is fairly frequent on the large islands of Fiji.

Alyxia linearifolia A. C. Sm. in Sargentia 1: 107. fig. 5. 1942, in Jour.
Arnold Arb. 27: 321. 1946.

Vitt Levu: Mba: Vicinity of Nalotawa, eastern base of Mt. Evans Range,
alt. 550-600 m., Smith 4471 (A, US) (shrub 2-3 m. high, the upper branches
becoming scandent, in forest along stream; flowers and fruits very scarce;
corolla-lobes white; fruit black). Vanua Levu: Mathuata: Southern
slopes of Mt. Numbuiloa, east of Lambasa, alt. 350-500 m., Smith 6563 (A, US)
(shrub 1-2 m. high, becoming scandent, in dense crest forest on summit of
southwestern ridge; fruit at length black).

The cited collections agree excellently with the two already known for
the species, both from Viti Levu. It is noteworthy that the species is
found in Mathuata in the same locality as the following, which is described
as a closely allied novelty. The small-leaved Alyxiae in Fiji are very
puzzling, for which reason I collected all those seen during my 1947 trip.
These are too few to permit an accurate picture of the range of variability,
but for the time being it seems best to limit A. linearifolia to the form with
leaves approximately 10-20 times as long as broad (not more than 3.5 mm.
in breadth) and parallel-margined.

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XI 115

Alyxia amoena sp. nov.

Frutex scandens copiose ramosus ubique partibus florum exceptis glaber,
ramulis gracilibus subteretibus fusco-cinereis; foliis oppositis vel ternatis
raro quaternatis, petiolis minutis canaliculatis ad 2 mm. longis, laminis in
sicco viridibus chartaceis anguste oblongo-ellipticis, (2—) 3-5 cm. longis,
(4—) 5-12 mm. latis, basi obtusis vel acutis et in petiolum decurrentibus,
apice obtusis, margine integris et leviter revolutis, costa supra paullo
impressa subtus elevata, nervis lateralibus numerosis immersis in nervulum
marginalem terminantibus; inflorescentiis axillaribus cymosis 3-—5-floris,
pedunculo gracili 1-4 cm. longo, bracteis anguste oblongis 1-2 mm. longis
obtusis, pedicellis gracilibus (2—) 3-9 mm. longis, bracteolis nullis; calyce
sub anthesi 1.6-1.7 mm. longo, lobis erectis membranaceis oblongo-
deltoideis subacutis, 1.2-1.3 mm. longis, 0.6-0.7 mm. latis, obscure
ciliolatis; corolla submembranacea, tubo urceolato-cylindrico circiter 3.5
mm. longo, superne circiter 1.5 mm. diametro, basi et apice contracto,
intus superne molliter retrorso-piloso, lobis oblongo-ovatis circiter 2 1.8
mm. apice rotundatis superne obscure ciliolatis; staminibus circiter 1 mm.
infra apicem tubi insertis, filamentis ligulatis circiter 0.3 mm. longis,
antheris oblongo-deltoideis acutis circiter 0.8 mm. longis; disco minuto
glabro; carpellis ovoideis distinctis glabris, stylo gracili circiter 1 mm.
longo, stigmate parvo capitato, ovulis in quoque carpello plerumque 4;
fructibus saepe persistenter binis, calyce subpersistente, stipite 2—2.5 mm.
longo; drupa ellipsoidea maturitate 7-11 mm. longa et 6-7 mm. diametro,
basi et apice obtusa, stylo persistente, pericarpio tenui (haud 0.2 mm.
crasso) extus levi vel obscure longitudinaliter sulcato intus ruguloso,
semine unico ellipsoideo rugoso.

Vanua Levu: Mbua: Ramasa Hill, H. B. R. Parham VIII (A) (small
shrub; flowers white); Mathuata: Southern slopes of Mt. Numbuiloa,
east of Lambasa, alt. 100-350 m., Oct. 27, 1947, Smith 6375 (A type, US)
(“vono”; liana, in open forest; corolla-lobes white; fruit at length black);
summit ridge of Mt. Numbuiloa, alt. 500-590 m., Smith 6498 (A, US) (“vono”;
shrub 2 m. high, with subscandent aera in dense forest; fruit becoming
black). OvaALau: Vicinity of Levuka, alt. 125 m., Gillespie 4568 (GH). Fiji,
without definite locality: U. S, Expl. Exped. ae in part, US 65908), Horne
671 (GH).

The new species is apparently most closely related to A. linearifolia
A. C. Sm., like which it has, for the genus, narrow leaves and comparatively
small flowers and fruits. In foliage A. amoena tends to be intermediate
between A. linearifolia and A. stellata (Forst.) R. & S.; in the former the
leaves are essentially linear, not more than 3.5 mm. broad, and with
parallel margins, while in the latter they are elliptic and usually sub-
stantially more than 1 cm. in breadth. The new species also differs from
A. linearifolia in its usually longer peduncles and pedicels, its larger
calyx-lobes, and its glabrous disk. From A. stellata (for laa
discussions of which see Christophersen, in Bishop Mus. Bull. : 184.
1935, and F. B. H. Brown, in op. cit. 130: 230. 1935) the new oe

116 JOURNAL OF THE ARNOLD ARBORETUM _[VoL. xxxuI

differs not only in its smaller leaves, but also in its shorter corollas and
smaller fruits. Some of the Fijian specimens referred to A. stellata (e. g.
Degener 14825, 14841, and 15255) have leaves nearly as small as those
of A. amoena, but their fruits are approximately twice as large. A revision
of the Pacific Alyxiae is much needed, as entities like A. stellata have been
so widely interpreted in herbaria that their actual limits are uncertain.

Fijian specimens intermediate in foliage between A. amoena and A.
linearifolia are: Viti Levu: Mba: Northern slopes of Mt. Namendre,
east of Mt. Koromba [Pickering Peak], alt. 750-900 m., Smith 4513
(A, US) (liana, in crest forest on wind-swept ridge; ripe fruit deep
purple); Nandronga & Navosa: Southern slopes of Nausori Highlands,
above Tumbenasolo, alt. about 450 m., Greenwood 1065A (A) (shrub 1 m.
high with long twining shoots, in forest; flowers white). These two speci-
mens, from adjacent localities in western Viti Levu, have the leaves shorter
and proportionately broader than those of A. linearifolia but yet averaging
smaller than those of A. amoena. The flowers of Greenwood 1065A show
the large calyx-lobes of A. amoena, and I believe this to be the better
position for the two collections, although they are not typical.

BORAGINACEAE

Cynoglossum amabile Stapf & Drummond in Kew Bull. 1906: 202.
1906.

Vitt Levu: Mba: Western slopes of Mt. Nanggaranambuluta [Lomalangi],
east of Nandarivatu, alt. 1000-1100 m., Smith 4809 (A, US) (naturalized along
track in dense forest; corolla bright blue).

Originally described from western Chinese specimens, this species is the
“Chinese forget-me-not” of gardens. In Fiji it was naturalized in the
indicated locality, probably as an escape from a European garden at
Nandarivatu, where, however, it no longer appears to be grown. I am
indebted to Dr. I. M. Johnston for the identification; the genus is not
otherwise recorded from Fiji.

LABIATAE

Salvia splendens Sellow ex Roem. & Schultes, Syst. Mant. 1: 185. 1822;
Epling in Rep. Sp. Nov. Beih. 85: 94. 1935.
Viti Levu: Nandronga & Navosa: Northern portion of Rairaimatuku
Plateau, between Nandrau and Rewasau, alt. 725-825 m., Smith 5398 (A, US)
(shrub 1-2 m. high, along trail in forest-grassland transition; calyx bright red).

This frequently cultivated species appeared to be naturalized in the
cited locality, which is far from any European settlement; it has not
previously been noted from Fiji.

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XI 117

ACANTHACEAE
Graptophyllum sessilifolium sp. nov.

Frutex ad 2 m. altus calycis lobis exceptis ubique glaber, ramulis
elongatis gracilibus novellis subcomplanatis demum teretibus cinerascenti-
bus; foliis subsessilibus interdum subamplexicaulibus, petiolis crassis
canaliculatis 0.5—2 mm. longis, laminis chartaceis vel subcoriaceis in sicco
olivaceo-viridibus oblongis vel anguste ovato-oblongis, (3.5—) 6-9 c
longis, 2-4 cm. latis, basi leviter cordatis, apice rotundatis vel obtusis,
margine integris et leviter revolutis, utrinque cystolithis linearibus 0.2—0.4
mm. longis manifeste ornatis, costa supra leviter subtus valde elevata,
nervis secundariis utrinsecus 4—6 arcuato-adscendentibus utrinque prom-
inulis vel subimmersis, rete venularum obscuro; inflorescentiis apices
ramulorum versus axillaribus cymosis plurifloris ad 5 cm. longis, pedunculo
(circiter 1 cm. longo) et ramulis gracilibus subteretibus, bracteis papyraceis
lanceolato-deltoideis acutis 1.5-2 mm. longis subpersistentibus, bracteolis
similibus 0.5—-1 mm. longis; pedicellis gracilibus sub anthesi et fructu 7—
12 mm. longis superne in calycis tubum gradatim incrassatis; calycis lobis
5 erectis lanceolatis, 1.7-2 mm. longis, 1-1.3 mm. latis, acuminatis, extus
glabris, intus minute puberulis, inconspicue 3- vel 5-nerviis; corolla
membranacea 25-30 mm, longa curvata, tubo circiter 13 mm. longo et
2—2.5 mm. diametro superne in faucem ventricosum sensim ampliato, lobis
5 subaequalibus, 8-9 mm. longis, 3.5-4 mm. latis, apice rotundatis et
obscure puberulis, 2 posticis paullo minoribus et altius connatis; staminibus
2 non exsertis, filamentis gracilibus circiter 7 mm. longis, antheris oblongis
3-3.5 mm. longis obtusis; disco carnoso integro ovario haud latiore;
ovario oblongo sub anthesi circiter 3 mm. longo in stylum filiformem quam
corollam breviorem attenuato, stigmate minuto, ovulis in quoque loculo 2
superpositis ; capsula anguste obovoidea circiter 2 cm. longa, basi stipitata,
apice acuta, seminibus 2 compresso-suborbicularibus conspicue rugulosis.

VaNvuA Levu: Mathuata: Southern slopes of Mt. Numbuiloa, east of
Lambasa, alt. 350-500 m., Nov. 10, 1947, Smith 6566 (A type, US) (shrub 1-2
m. high, with elongate branches, in steep open forest; corolla rich pink).

The closest relative of the new species seems to be G. insularum (A. Gray)
A. C. Sm., fairly abundant in Fiji (for distributional notes see Sargentia
1: 118. 1942), which, however, has distinctly petiolate leaves, of which the
blades are thinner, more obviously nerved, and obtuse to acute at base.
Although the flowers of the two species are essentially similar, G. insularum
often has the inflorescence-branches and pedicels (as well as young
vegetative parts) closely puberulent, and the inflorescence comparatively
contracted and fewer-flowered.

COMPOSITAE

Centipeda minima (L.) A. Br. & Aschers. Ind. Sem. Hort. Berol. App.
6. 1867

118 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxm

VANUA Levu: Mathuata: Seanggangga Plateau, in drainage of Korovuli
River, vicinity of Natua, alt. 100-200 m., Smith 6885 (A, US) (on edge of pond
in open rolling country; corolla-lobes pale green).

Although I find no previous published record of the occurrence of this
plant in Fiji, Mr. William Greenwood has indicated its presence by
including it in an unpublished list. In the cited locality in Mathuata the
small plants were quite common, although very inconspicuous, growing
on moist savanna with Cyperus polystachyos Rottb., Fimbristylis annua
var. diphylla (Retz.) Kiukenth., Echinochloa colonum (L.) Link,
Paspalum orbiculare Forst., Jussiaea suffruticosa var. ligustrifolia (H. B.
K.) Griseb., Limnophila fragrans (Forst.) Seem., and Erechtites valerianae-
folia DC. Although this would seem to be an association of widespread
weeds, I have not seen the Centipeda elsewhere in Fiji.

DEPARTMENT OF BOTANY,

U. S. NATIONAL Museum,
SMITHSONIAN INSTITUTION.

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XII 119

STUDIES OF PACIFIC ISLAND PLANTS, XII
THE CUNONIACEAE OF FIJI AND SAMOA

A. C. SMITH

ALTHOUGH THE CUNONIACEAE, a family of considerable size and
diversity in New Guinea and New Caledonia, are represented eastward in
the Pacific by a sharply decreasing number of members, they are neverthe-
less a puzzling group in the area under consideration, to judge from the
uncertainty of herbarium identifications and the difficulty of analyzing
specific criteria. While attempting to place the specimens of the family
which I collected in 1947 1, it seemed desirable to assemble earlier material,
to prepare keys, and to redescribe the species, the original descriptions
having been based upon too few specimens to show their variability. The
place of deposit of specimens cited in this paper is shown as follows: Arnold
Aboretum (A); Bernice P. Bishop Museum (Bish); British Museum
(BM); Gray Herbarium (GH); New York Botanical Garden (NY):
and U. S. National Herbarium (US). The kindness of the authorities of
these institutions in permitting study of their material is greatly ap-
preciated.

The Cunoniaceae do not occur in Tonga, on the basis of available records,
but in Fiji the family is represented by 14 species and in Samoa by three
species, all being endemic to one or the other archipelago. Of the four
genera occurring in this area, only Weinmannia is widespread, with an
extensive distribution mostly in the Southern Hemisphere. Spiraeanthemum
is found from New Guinea and Australia eastward to Samoa, where its
range is terminated by a single species; Geissois is more limited, occurring
in New Caledonia, Australia, and the New Hebrides, with four species
terminating its range in Fiji. Of special interest is ‘he occurrence in Fiji
of a species of Pullea, a genus previously believed limited to New Guinea.

In this paper three species and three varieties are described as new.
Criteria for the demarcation of species in the Cunoniaceae are often
neither obvious nor constant, but the genera are well marked. The
following key to genera utilizes only characters found in the species of our
region:

Inflorescense racemose, the racemes solitary or 2-4 at apex of a short common
eduncle or arising from inconspicuous glomerules; ovary 2- carpellate, the
capsule septicidally 2-valved; leaves compound or sometimes simple.

Flowers large, the calyx and filaments red, the stamens 8-26, with filaments

mm. long; petals none; disk pulvinate, entire; ovules numerous,

Under the nda the Arnold Arboretum of Harvard University and the John

Simon ae eim Memorial Foundation, with the aid of grants from the Penrose

Fund of the eae Philosophical Society and the Bache Fund of the National
epee of Sciences.

120 JOURNAL OF THE ARNOLD ARBORETUM _[VoL. xxxmII

20-42 per locule, 2-seriate; seeds distally winged; plants with herma-
phrodite flowers and 3-foliolate leaves ................ 1. Geissots.
Flowers small, the petals and filaments white or greenish, the stamens 8,
with filaments up to 4 mm. long; petals 4; disk divided into 8 free
lobes; ovules 3-12 per locule; seeds comate at both ends, not winged ;
plants polygamo-dioecious, the leaves pinnate or 3-foliolate or simple

2. Weinmannia.

Inflorescence paniculate; flowers small, the calyx and filaments white to yellowish
or greenish, the petals none, the stamens (6 —) 8-12; the disk divided into
lobes; leaves simple.

Plants dioecious or possibly polygamo-dioecious; inflorescences solitary; ¢
flowers with (3 —) 4-6 free or loosely connate disk-lobes and no carpels;
Q flowers with (6 —) 8-12 disk-lobes and 4 or 5 (rarely 3 or 6) free

disk-lobes 10 or 12, often coherent in pairs; ovary 2-carpellate, the
ovales A per W6cdle. 3). coin ney eek ere eres sang h es cnet ess 4. Pullea

1. Gertssors Labill.

Geissois, originally based on a species from New Caledonia, is now
considered to include about 17 species; its range centers in New Caledonia,
but species also occur in Australia and the New Hebrides and eastward
to Fiji, beyond which the genus does not extend.

In Fiji the local names vure and vota are usually applicable to Geissois,
and in some localities G. ternata is a frequent and striking component of
the vegetation, conspicuous for its beautiful red-flowered inflorescences.
Criteria for the recognition of species, insofar as they refer to the size
and indument of vegetative parts, are satisfactory within limits. The
stipules in particular provide reliable characters, and in the case of G.
superba the extreme length of the raceme is very obvious. Floral characters
are usually too uniform to be of much taxonomic value, although the
indument of the ovary, on the contrary, is too variable. In both G.
imthurnii and G. ternata the ovary may vary from glabrous to strigillose
or even densely sericeous, and as this variation seems uncorrelated with
other characters I have not emphasized it.

In Fiji four species can be discerned, of which one is described as new.
By far the greater part of the Fijian population of the genus represents
G. ternata, which I here divide into four varieties, three of them new.

KEY TO THE SPECIES
Inflorescence robust, 22—45 cm. long, the stamens 14-26; leaves comparatively
large, the petiole 1.5-8 cm. long, the petiolules 1.5-8 cm. long, the leaflet-
blades usually 24-50 X 10-19 cm., with 13-20 secondary nerves per side;
stipules large, ovate-oblong, comparatively persistent, up to 60 & 45 mm.,
proximally laterally connate. .............-.---.+02005: 1. G. superba.
Inflorescence much smaller, not exceeding 10.5 cm. in length, the stamens 8-15;
leaves smaller, the petiole not exceeding 5 cm. and the petiolules 6 cm.
(usually less than 2.5 cm.) in length, the leaflet-blades not more than

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XII 121

23 X 10.5 cm., usually much smaller, with not more than 15 secondary
nerves per side; stipules usually not persistent after attaining a length of
about 7 mm., if persistent oblong or elliptic, not more than 20 mm. broad,
and laterally free.
Leaflets nearly sessile, the blades rounded to subacute at base, the petiolules
of lateral ones up to 2 mm. (rarely to 3 mm.) and of terminal one up
o 3 mm. (rarely to 6 mm.) long; leaflet-blades hispidulous on both
surfaces, the hairs usually persistent, densest on costa and secondaries ;
branchlets, petioles, and petiolules copiously setulose or strigillose, rarely
subglabrate; inflorescence-rachis and pedicels hispidulous, the calyx-lobes
sparsely strigillose on both sides. ..................2. G. imthurni.
Leaflets obviously petiolulate, the petiolules usually 4 mm. or more long (if
shorter, the leaflet-base attenuate and long-decurrent); leaflet-blades
glabrous or faintly strigillose on costa.

Stipules subpersistent, at length ligulate-oblong, up to 10 x 2 cm., copiously
hispid without with hairs 1.5-2.5 mm. long, glabrous within; branchlets
robust, distally conspicuously flattened, the petioles similarly flattened,
copiously hispidulous or strigillose, the petiolules 1.5-6 cm. long,
the leaflet-blades 11-23 < 6.5-10.5 cm.; calyx-lobes 6.5—7 mm. long.
ess ors heehee ek See 3. G. stipularis.

Stipules usually early caducous, if subpersistent apparently not exceeding
a size of about cm., variously pilose or glabrous on both sides;
branchlets subterete or distally slightly flattened, the petioles semiterete,

1. Geissois superba Gillespie in Bishop Mus. Bull 83: 9. fig. 9. 1931.

Tree 10-13 m. or more in height, the branchlets stout (0.8-1.5 cm. in
diameter toward apex and there subquadrate or slightly flattened), distally
hispidulous or puberulent (hairs 0.1—-0.5 mm. long), the older parts
glabrate and lenticellate; stipules subcoriaceous, ovate-oblong, rapidly
enlarging to about 6 cm. long and 4.5 cm. broad before falling, laterally
connate at base, forming a bilobed cupule, recurved at margin, very densely
velutinous-hispidulous on both sides (hairs 0.3-0.5 mm. long), rarely
glabrate, the scars forming a conspicuous continuous ring; leaves opposite,
3-foliolate, the petioles semiterete, stout, 1.5-8 cm. long, hispidulous
like branchlets, sometimes glabrate, the petiolules stout, shallowly
canaliculate or subterete, 1.5-8 cm. long (terminal slightly the longest),
pilose like petioles; leaflet-blades chartaceous to subcoriaceous, elliptic or
obovate-elliptic, (15—) 24-50 cm. long, (6—) 10-19 cm. broad, acute to
obtuse at base and decurrent on the petiolules, rounded at apex, entire
and narrowly recurved at margin, glabrous above or sparsely strigillose on
costa and secondaries, beneath strigillose on principal nerves and some-
times on surface, the costa elevated or prominent above, very prominent
beneath, the secondary nerves 13-20 per side, erecto-patent, nearly
straight, plane or slightly elevated above, prominent beneath, the veinlet-
reticulation intricate, prominulous on both surfaces; racemes axillary and
borne within the stipule-cupule or lateral below leaves, solitary or paired,

122 JOURNAL OF THE ARNOLD ARBORETUM _ [vot. xxxut

pendent, 22-45 cm. long, the peduncle 3—12 cm. long, stout, terete, curved,
with the conspicuously striate rachis sparsely hispidulous; flower-subtending
bracts lanceolate, 2—2.5 mm. long, dorsally strigillose, soon caducous, the
pedicels strigillose, 7-12 mm. long, articulate near or slightly below middle;
calyx-lobes 4, carnose, becoming subcoriaceous, narrowly deltoid or
lanceolate, 5-6 1.8-2.5 mm., acute, sparsely strigillose without and
more densely hispidulous-tomentellous within (hairs 0.2-0.5 mm. long) ;
stamens 14-26, the filaments 15-20 mm. long, the anthers oblong, about 1
mm. long; disk pulvinate, inconspicuously grooved, 0.5-1 mm. high and
about 2 mm. in diameter; ovary copiously hispidulous-tomentellous or
merely sparsely strigillose (hairs 0.8-1.3 mm. long), the styles 10-12 mm.
long, the ovules about 40 per locule, imbricate, biseriate; receptacle
swollen in fruit, the calyx-lobes soon caducous; capsule cylindric, falcate,
15-22 mm. long, persistently strigillose-hispidulous but the hairs often
sparse, the pericarp coriaceous, the seeds about 3 mm. long.

DIsTRIBUTION: Endemic to Fiji and thus far known only from Viti Levu, at
elevations up to 900 m. The species is a tree up to 13 m. high or perhaps larger,
occurring in forest; the calyx and filaments are crimson and the anthers yellow.
Vure is a reported local name. The type is Gillespie 4274, cited below.

FIJI: Virr Levu: Mba: Between Nandarivatu and Vatuthere, Gillespie
3178 (Bish, GH), 4274 (Bish type, GH, K); Nandronga & Navosa
Northern portion of Rairaimatuku Plateau, between Nandrau and Rewasau,
Smith 5434 (A, US); Serua: Korovisilou, B. E. Parham 1434 (A); Rewa
or Naitasiri: “Central Road, Suva,” Tothill 471 (K); Viti Levu, without
further locality, Tothill 189c (K).

The very distinct G. superba is readily distinguished from its congeners
in Fiji by its robust leaves and elongate inflorescence, and by its char-
acteristically large stipules, which are comparatively persistent and
connate into a bilobed cupule.

2. Geissois imthurnii Turrill in Jour. Linn. Soc. Bot. 43: 19. 1915, in
Hook. Ic. Pl. 31: pl. 3053. 1916.

slightly flattened above, (1—) 1.5-3 cm. long, copiously setulose like
young branchlets, rarely glabrate, the petiolules comparatively incon-
spicuous, copiously setulose, of lateral leaflets 0-2 (-3) mm. long, of
terminal leaflet 1-3 (—6) mm. long; leaflet-blades chartaceous to sub-
coriaceous, oblong- or obovate-elliptic, 6—11.5 (-18.5) cm. long, (2.5—)
3.5—6 (-8.5) cm. broad, broadly obtuse or rounded and inequilateral
(lateral leaflets) or subacute (terminal leaflet) at base, obtusely cuspidate
to broadly obtuse at apex, entire and slightly recurved at margin, hispidu-

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XII 123

lous on both surfaces with spreading hairs 0.3-0.8 mm. long, rarely
subglabrate but usually with persistent indument at least on costa and
secondaries, the costa plane or slightly elevated above, prominent beneath,
the secondary nerves 9-15 per side, subspreading, curved, prominulous
or plane above, strongly elevated beneath, the veinlet-reticulation intricate,
slightly prominulous or plane on both surfaces; racemes borne on defoliate
branchlets, axillary to leaf-scars, solitary or 2 or 3 arising from an incon-
spicuous glomerule, 3-10 cm. long, the peduncle 0.5—3.5 cm. long, with
the rachis sparsely hispidulous (hairs 0.2-0.4 mm. long) ; flower-subtending
bracts deltoid or lanceolate, 0.8-1.6 mm. long, sparsely setulose without,
soon caducous, the pedicels pilose like rachis, 3.5-8 mm. long, articulate
slightly below middle; calyx-lobes 4, papyraceous, deltoid-lanceolate,
4.5-6 < 2-2.5 mm., acute, sparsely strigillose on both sides; stamens
9-12, the filaments 13- 16 mm. long, the anthers oblong, 1—-1.2 mm. long;
disk pulvinate, 0.8-1 mm. high, 2—2.5 mm. in diameter; ovary glabrous or
with a few stiff hairs or setulose-strigose with hairs 0.5—-1 mm. long (some-
times variable on same individual), the styles 9-15 mm. long, the ovules
about 40 per locule, biseriate; capsule linear-oblong, falcate, 18-28 mm.
long, 3-5 mm. in diameter, glabrous or persistently strigillose, the seeds
4—5 mm. long, the nucellus ellipsoid, about 2.5 mm. long, the wing distal,
rounded at apex.

DIstTRIBUTION: Endemic to Fiji and thus far obtained only from a limited area
near Nandarivatu, Viti Levu, at elevations of 750 to 900 m. It is a tree,
recorded as 10-20 m. in height and with a trunk diameter up to 1 m., occurring in
forest and on hillsides, mentioned by some collectors as locally common. The
calyx and filaments are bright red to deep rose-pink. The local name, as for
other species of Geissois, is vure; im Thurn mentions the name vunga, which
is usually applicable to Metrosideros.

FIJI: Virt Levu: Mba: Nandarivatu and immediate vicinity, im Thurn
137 (Bm, K type), Parks 20671 (Bish), Greenwood 886 (A, K), Degener
14265 (A, Bish, K, NY, US), Reay 17 (A, K, US), Vaughan 3432 (BM);
Nukunuku Creek, Vaughan 3401 (BM).

The nearly sessile leaflets and the general pubescence of this plant,
which usually persists on the leaflet-blades, differentiate it without difficulty
from G. ternata. However, transitional forms, in which the petiolules are
comparatively obvious (e. g. Reay 17), indicate that G. imthurnii is not
as isolated a taxon as might be inferred from the type specimen alone.

3. Geissois stipularis sp. nov.

Arbor, ramulis crassis apices versus conspicue complanatis et pilis
0.2—0.5 mm. longis parce strigilloso-puberulis, demum glabratis cinereisque
inconspicue lenticellatis; stipulis papyraceis juventute in gemma compacta
subglobosa cohaerentibus, mox accrescentibus subpersistentibus, demum
ligulato-oblongis 6-10 cm. longis 1.2-2 cm. latis ad basim liberis apice
rotundatis, extus pilis 1.5—2.5 mm. longis copiose hispidis ac etiam minute
puberulis, margine puberulo-tomentellis, intus glabris, cicatricibus con-

124 JOURNAL OF THE ARNOLD ARBORETUM _ [VOL. xxxII

spicuis leviter curvatis; foliis oppositis 3-foliolatis, petiolis crassis valde
complanatis 1.5—5 cm. longis pilis 0.4-0.7 mm. longis copiose hispidulis vel
strigillosis, petiolulis canaliculatis vel semiteretibus 1.5—6 cm. longis ut
petiolis pilosis vel subglabratis; foliolorum laminis coriaceis in sicco
brunnescentibus elliptico- vel obovato-oblongis, 11-23 cm. longis, 6.5—10.5
cm, latis, basi obtusis vel subacutis et in petiolulum decurrentibus, apice
obtuse cuspidatis, margine integris anguste recurvatis, utrinque glabris vel
costa parce strigillosis, costa valida supra leviter elevata subtus prominente,
nervis secundariis utrinsecus 8-12 erecto-patentibus supra subplanis subtus
prominentibus, rete venularum conspicuo intricato utrinque prominulo
vel supra subimmerso; racemis infra folia enatis solitariis 4-8 cm. longis,
pedunculo brevi tereti et rhachi striata gracilibus strigilloso-puberulis;
bracteis caducis, pedicellis sub anthesi 5-7 mm. longis medium versus
articulatis, infra articulationem ut rhachi pilosis superne glabris; calycis
lobis 4 carnosis deltoideo-lanceolatis, 6.5-7 mm. longis, 2.5-3 mm. latis,
extus glabris, intus pilis pallidis 0.4-0.7 mm. longis hispidulis; staminibus
12 vel 13, filamentis ligulatis sub anthesi 12-15 mm. longis, antheris
oblongis 1.2—-1.4 mm. longis utroque emarginatis; disco carnoso pulvinato
0.8-1 mm. alto circiter 2.5 mm. diametro; ovario oblongo-conico glabro,
stylis 10-12 mm. longis, ovulis biseriatis circiter 30 in quoque loculo.

DIsTRIBUTION: Known only from the two collections cited below and
perhaps limited to southeastern Viti Levu, Fiji.

FIJI: Virr Levu: Naitasiri: Tamavua woods, 7 miles from Suva, alt.
150 m., Aug. 9, 1927, Gillespie 2118 (Bish TypE, GH, US); Viti Levu, without
further data, Parks 20940 (Bish).

Although the number cited as the type collection is sterile, it bears
locality data and shows the foliar and stipular characters that are diagnostic
for the species. The Parks specimen is accompanied by’ inflorescences
but is without data; some of his plants were also obtained in southeastern
Viti Levu.

Superficially the new species, in its large and long-petiolulate leaflets,
suggests G. superba, but its leaflets are actually considerably smaller and
fewer-nerved than in that species, and its stipules are entirely different.
In inflorescence it seems closer to G. ternata, but characters pertaining
to the stipules and branchlets differentiate it, while the predominantly
larger leaves and calyx-lobes of G. stipularis are also characteristic.

4. Geissois ternata A. Gray, Bot. U. S. Expl. Exped. 1: 679. pl. 86.
1854

Shrub or tree, up to 25 m. high, the branchlets subterete or distally
flattened, sparsely strigillose with hairs 0.2-0.3 mm. long, soon glabrate,
lenticellate; stipules ovate to oblong or elliptic, usually caducous when
small, copiously setulose or strigillose to glabrous, rarely persisting to a
size of 3 cm. long, free to base, the scars elongate, callose-thickened;
leaves opposite, 3-foliolate, the petioles semiterete, 7-35 (—50) mm. long,
sparsely strigillose when young, glabrate, the petiolules slender, shallowly

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XII 125

canaliculate, Acie ti like petioles and soon glabrate, of lateral leaflets

mm. long, of terminal leaflet to 25 mm. long; leaflet-blades sub-
coriaceous or chartaceous, elliptic or obovate-elliptic, 3-17 (—19) cm.
long, (1.2—-) 1.5-9 (-10.5) cm. broad, obtuse to attenuate at base and
decurrent on the petiolule, rounded to obtusely cuspidate or acuminate at
apex, entire at margin and plane or narrowly recurved (rarely denticulate-
serrulate), glabrous on both surfaces or faintly strigillose on costa beneath,
the costa plane or slightly elevated above, prominent beneath, the second-
ary nerves 5—13 per side, spreading or ascending, slightly curved, plane or
prominulous above, slightly elevated beneath, the veinlet-reticulation
intricate, prominulous on both surfaces or immersed above; racemes
borne on defoliate branchlets, solitary or 2 or 3 arising from a small
glomerule, 2.5-10.5 cm. long, the peduncle subterete, 0.5—-2 cm. long,
with the slightly angled rachis glabrous or faintly strigillose-puberulent;
flower-subtending bracts lanceolate, 1.2-1.5 mm. long, glabrous or very
sparsely strigillose dorsally, soon caducous, the pedicels 4-10 mm. long,
articulate near middle or slightly above middle or sometimes near base,
glabrous; calyx-lobes 4, papyraceous or carnose, lanceolate or narrowly
deltoid, 4.5—6 & 1.4-3 mm., glabrous on both sides or sparsely strigose to
crispate-pilose within; stamens 8-15, the filaments 11-18 mm. long, the
anthers ellipsoid or oblong, 0.8—1.2 mm. long; disk pulvinate, 0.5-1.2 mm.
high, 1.5—2 mm. in diameter; ovary glabrous or sparsely strigillose with
hairs 0.3-0.7 mm. long (rarely densely strigose-sericeous), the styles 8-13
mm. long, the ovules 20—42 per locule, biseriate; capsule cylindric, falcate,
12-27 mm. long, 3-5 mm. in diameter, glabrous or strigillose to setulose-
puberulent, the seeds about 5 mm. long, the nucellus ellipsoid, the wing
distal, rounded at apex.

DISTRIBUTION: Throughout Fiji, endemic, at elevations up to 1050 m. The
species is a shrub or tree, up to 25 m. in height, occurring in a variety of habitats,
including forest, hillside thickets, open places, etc. The calyx, filaments, and
styles are deep red or bright red, the anthers, disk, and ovary yellow, and the
fruit dull yellow or red-tinged, becoming brown. Local names for this common
species are vure and vota, sometimes vurevure, and rarely vunga.

The residual population of Geissois in Fiji, when reasonably well
characterized taxa like G. superba, G. imthurnii, and G. stipularis have
been segregated, may be designated as G. ternata. Superficial acquaint-
ance with this population shows that it is fairly heterogeneous, but detailed
examination does not disclose obvious lines of differentiation. Nevertheless
the available material seems too diverse to be left in a single taxon, and
one is able to discern in it various morphological tendencies that seem
usable for the establishment of infraspecific groups. The four groups
here proposed as varieties are far from satisfactory, but I believe that
their recognition permits a better understanding of G. ternata.

Of the proposed varieties, the best marked is characterized by a reduc-
tion in size of leaves and number of floral parts; the leaflet-blades tend
to be blunter at apex and more attenuate at base, and concomitantly the

126 JOURNAL OF THE ARNOLD ARBORETUM _ [VOL. xxxiII

stamens are reduced in number (8-12), the disk is shorter, and the ovules
are comparatively few (20-34 per locule). The latter character, although
impracticable for general use, suggests that the tendencies here recognized
are not entirely superficial. This variety (var. minor) usually occurs at
high elevations or in exposed places.

A proposed variety (var. serrata) known only from the Yasawa Group
differs from the typical form in its comparatively large leaflets with
serrulate margins (the species otherwise having entire leaflets), and also
in the longer indument of its stipules. The value of this variety can scarcely
be assessed without more material, but it seems inadvisable to include
a form with toothed leaflets with typical material.

The remaining specimens are more homogeneous, although there is
still a great deal of variation in leaf-size. Two types of stipule-indument
are discernible; these organs may be copiously setulose with spreading
hairs or they may be essentially glabrous (with hairs, when present, of a
closely appressed type). This character may not be very consequential,
but it is readily observed and is fairly constant, the apical stipules being
present even on specimens in advanced fruiting stages. The type of the
species falls into the first group, with setulose stipules (var. ternata),
and the other group I propose as var. glabrior, Elsewhere in the
Cunoniaceae the type of stipule-indument is a reliable character and is cor-
related with other criteria; in the present case it seems to be supported by
no other consistent characters.

KEY TO THE VARIETIES

Leaves comparatively large, the petiolules (2—) 4-25 mm. long, the leaflet-blades
sually 5-17 * 3-9 cm., obtuse to acute at base, obtusely cuspidate to
acuminate at apex; inflorescence 4-10.5 cm. long, the stamens 12-15, the
disk 0.8-1.2 mm. high, the ovules 36—42 per locule.
Leaflet-blades entire, usually 5-15 3-7.5 cm., the secondary nerves 5-11

per side.
Stipules copiously setulose with spreading hairs 0.2-1 mm. — syanquag

Pe RLSHS HPESR SL eES ee beh eee AV Nena yaa eee eenen es 4a. r. ternata.

Stipules glabrous on both sides or strigillose with appressed ale 0.1-0.4
mm. long, sometimes puberulent-tomentellous at a pee aae ies

Se Gkis bos 4a boo cele a Peed S bd OR kee RRNA ES ES 4b. var. glabrior.
oe vane denticulate-serrulate at margin, large, ae 9-17 X
_ , the se ondary nerves 9-13 per side; stipules copiously setulose

with ee 1.5- 0 i (AGU cc nnd chet es sees ees 4c. var. serrata.
Leaves comparatively aa the petiolules 1-11 mm. long, the leaflet-blades
usually 3-9.5 1.5-5 cm., attenuate at base, obtuse to rounded at apex,
entire; stipules copiously setulose: inflorescence 2.5—8 cm. long, the stamens
8-12, the disk 0.5-0.6 mm. high, the ovules 20-34 per locule. 4d. var. minor.

4a. Geissois ternata var. ternata.

Geissois ternata A. Gray, Bot. U. S. Expl. ii 1: 679. pl. 86. 1854; Seem.
Fl. Vit. 109. 1865; Pampan. in Ann. di . 2: 58. 1905; Gibbs in Jour.
Linn. Soc. Bot. 39: 144. 1909.

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XII 127

The typical variety, characterized by having stipules copiously setulose
with spreading hairs 0.2-1 mm. long; petiolules (2—) 4-25 mm. long;
leaflet-blades (4—) 5-15 (-19) cm. long, (2—) 3-7.5 (—10.5) cm. broad,
acute to obtuse at base, obtusely cuspidate at apex, entire at margin, the
secondary nerves 5-11 per side; racemes 4-10.5 cm. long, the stamens
12-15, the disk 0.8-1.2 mm. high, the ovules 36—42 per locule.

DISTRIBUTION: Known from several of the islands, at elevations from near
sea-level up to 900 m., and apparently the most abundant variety on Viti Levu.
The type material, obtained by the U. S. Exploring Expedition, comes from at
least two plants, obtained on Ovalau and in the Province of Mathuata, Vanua
Levu.

FIJI: Virt Levu: Graeffe (K), 27 (BM); Mba: Northern portion of Mt.
Evans Range, between Mt. Vatuyanitu and Mt. Natondra, Smith 4271 (A, US);
vicinity of Nandarivatu, Gibbs 591 (BM), Smith 5969 (A, US); Nandronga

avosa: Southern slopes of Nausori Highlands, in drainage of Namosi
Creek above Tumbenasolo, Smith 4605 (A, US); vicinity of Mbelo, near
Vatukarasa, Degener 15274 (A, Bish, K, NY, US); Serua: Mbuyombuyo,
near Namboutini, Tabualewa 15609 (A, Bish, K, NY, US); Thulanuku, near
Ngaloa, Degener 15120 (A, Bish, K, NY, US). KAnpAvu: Seemann 201 (BM,
GH, K); hills above Namalata and Ngaloa Bays, Smith 76 (Bish, GH, K, NY,

US). OvaLau and Vanua Levu: U. S. Expl. Exped. (GH, K, NY, US 47817
and 47818 Type). Fiji, without definite ‘Teealiey: Horne (GH).

4b. Geissois ternata var. glabrior var. nov.

Frutex vel arbor grandis a var. ternata stipulis utrinque glabris vel pilis
adpressis 0.1-0.4 mm. longis strigillosis interdum margine puberulo-
tomentellis differt.

DISTRIBUTION: Recorded from several islands in Fiji, at elevations from
near sea-level up to 500 m.; it seems to have a more easterly distribution within
the group than var. ternate. As type I designate Smith 1590, from Vanua Levu,
a collection with flowers and fruits which also shows the diagnostic stipule
character.

FIJI: Virr Levu: Namosi: Between Namuamua and Laselase, Gillespie
3213 (Bish, GH, K, NY). VANua Levu: Mbua: Upper Ndama River Valley,
Apr. 24, 1934, in dense forest at 100-300 m., Smith 1590 (Bish, GH, K, NY Type,
US); Thakaundrove: Hills south of Nakula Valley, Smith 343 (Bish,
GH, K, NY, US); Valanga, Savu Savu Bay region, Degener & Ordonez 14034
(A). Taveuni: Vicinity of Waiyevo, Gillespie 4699 (Bish, K, NY, US);
western slope, between Somosomo and Wairiki, Smith 847 (Bish, GH, K, NY,
US). Koro: Western slope, Smith 1085 (Bish, GH, K, NY, US). VANUA
MBatLavu: Slopes of highest peak, Bryan 583 (Bish); near Lomaloma, Smith
1424 (Bish, K, NY). Lakempa: Harvey (GH, K

4c. Geissois ternata var. serrata var. nov.

Arbor ad 15 m. alta, stipulis pilis 1.5-2 mm. longis dense setosis, petiolis
(10—) 17-25 mm. longis, petiolulis (5—) 12-25 mm, longis, foliolorum
laminis ovatis vel ellipticis, (6—-) 9-17 * (2.5-) 4-9 cm., apice obtuse
cuspidatis vel acuminatis, margine saltem supra medium denticulato-

128 JOURNAL OF THE ARNOLD ARBORETUM _[VoL. xxxmI

serrulatis (dentibus 1—3 per centimetrum parvis superne calloso-apiculatis) ,
nervis secundariis utrinsecus 9-13; a var. ternata foliolorum laminis
manifeste serrulatis, magnis, apice saepe acuminatis, nervis secundariis
numerosis, stipularum pilis longioribus differt.

DISTRIBUTION: Known only from the type collection, from Waya Island in
the Yasawa Group, northwest of Viti Levu

FIJI: Waya, Yasawa Group: North of Walon _ along re Creek, alt.
120-240 m., July 19, 1937, St. John 18128 (Bish TYPE, US) (‘‘vunga’’; tree
15 m. high, common, the trunk 8-10 cm. in diameter; pee red; gent ie sed
for houses; birds visit the flowers).

4d. Geissois ternata var. minor var. nov.

Frutex vel arbor ad 4 m. alta, stipulis eis var. ternatae similibus;
petiolis 7-25 mm. longis, petiolulis 1-11 mm. longis, foliolorum laminis
anguste ellipticis vel lancantaie. ellipticis, 3-9.5 & (1.2—) 1.5—5 cm., basi
attenuatis, apice obtusis vel rotundatis, nervis secundariis alaaecay 5-8;
racemis 2.5—8 cm. longis, staminibus 8-12, disco 0.5—0.6 mm. alto, ovulis
20-34 in quoque loculo; a var. ternata foliis minoribus, foliolorum laminis
basi attenuatis apice saepe rotundatis, inflorescentia minore, staminibus
et ovulis paucis, disco breviore differt.

DIsTRIBUTION: Known from the two large islands of Fiji, often at com-
paratively high elevations (500-1050 m.), where it occurs in forest, dense
thickets, or in exposed places. It is a comparatively small plant, noted as a
gnarled shrub or a tree 2-4 m. high. The type, a specimen with flowers, young
fruits, and characteristically small leaves, is Smith 679, from Vanua Levu.

FIJI: Vit1 Levu: Mba: Mt. Evans Range, Greenwood 119 (K); Tholo-i-
Nandarivatu, Gillespie 3898 (Bish); Namosi: Summit of Mt. Voma,
Gillespie 2730 (Bish). VANUA Levu: Mathuata: Summit ridge of Mt.
Numbuiloa, east of Lambasa, Smith 6514 (A, US); Thakaundrove:
Summit of Mt. Mbatini, alt. 1030 m., Nov. 29, 1933, Smith 679 (Bish, GH, K,
NY type, US).

2. WEINMANNIA L.

The genus Weinmannia, as here considered, is represented by five species
in Fiji and two in Samoa, being absent from Tonga as far as known. The
Fijian and Samoan species appear to be endemic, records of their occurrence
in more than one archipelago being discussed below. The genus is not a
common component of the vegetation in either group. Criteria for specific
delimitation in Weinmannia are not satisfactory, such characters as
simple vs. pinnate leaves and degree of indument being highly variable.
To a certain extent more dependable characters are found in the shape
of stipules, the persistent or caducous nature of the calyx, and the number
of ovules. In my observation, the Fijian species have the ovules 3-6 per
locule as opposed to 8-12 in the Samoan species. Leaflet-shape is a
usuable character only within very broad limits; one species, here des-
cribed as new, is characterized by very small leaves and compact

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XII 129

inflorescences. All the species of our region are probably polygamo-
dioecious; staminate flowers have comparatively long filaments, short
styles, and sterile carpels, while hermaphrodite flowers have shorter
filaments, longer styles, and readily observed ovules in the carpels.
Superficially the two types of flower are not easily distinguished. It may
be noted that the Fijians seem to have no common name for Weinmannia
which is generic in nature, as they do for Spiraeanthemum and Geissois.

KEY TO THE SPECIES
Leaves simple, rarely 2- or 3-foliolate, the blades up to 11 X 6 cm.; stipules
ovate to elliptic or suborbicular-obovate, entire, often nearly as broad as

ong, the petals 1.6-1.8 mm. long; perianth caducous in fru
Stipules comparatively large, 13-25 * 10-15 mm., fecal a barbellate in
axils, the tufts of hairs often subpersistent ; leaves nearly always simple,
very rarely 2-foliolate, the blades oblong-elliptic, usually 7-11 X 2.5-6
cm., the marginal crenations usually 1 or 2 per centimeter; ovules usually
4 per locule; seeds copiously comate at both ends, the hairs is 1 mm.
1

long, with obvious cross-walls; Fiji . hs ; affinis.
Stipules smaller, 6-12 & 2-9 mm., not ba rbellate in “axils or nu incon-
spicuously so; leaves simple or 3- he ig the blades predominantly
Ee usually 4-10 X 1.3-4 cm., the marginal crenations usually 3

or 4 per centimeter; ovules 10-12 ee Jocule: seeds more sparsely comate
at both ends, the hairs 0.4-0.5 mm. long, the ae Fd ata
Se 0, ae a ee er W. manuana
Leaves compound, 3-9-foliolate, a qaoliolate: ers on known for
) Spy ee ne small, the sepals less than 1.2 mm. long, the petals
less than 1.6 m

eae a eeaty large, only rarely less than 2 X 1 cm., usually much
the marginal crenations only rarely as few as 8 per side; racemes

more than 4 cm. long, often up to 12 cm. or longer
Leaves with the petiole, rachis, and lower leaflet- surfaces hispidulous ae
0 5-1 mm. long) ; eee suborbicular or ovate- oblong, about ~
Pr mm., conspicuously dentate; known only in sterile are

3. J

Fij Ee et a) tee eee ee ; spiracoides.
tear labo or with the petiole, rachis, = costa be lower leaflet-
surfaces puberulent (hairs up to 0.2 mm. long), in the Samoan

ee ae petiole and costa sometimes strigose (hairs 0.5-1 mm.
long); stipules entire.

Stipules suborbicular, very variable in size but usually slightly broader
than long; leaves variable, with (1—-) 3-9 leaflets, these predom-
inantly elliptic or oblong-elliptic; sepals 0.5-0.7 mm. long; petals
1-1.3 mm. long; ovules 4-6 per locule; perianth persistent in fruit;
| | rier ere WU: 2 hater ar ae ee emery” 4. W.. richit.

Stipules oblong or ovate to lanceolate, longer than broad; leaflets pre-

minantly lanceolate or lanceolate-elliptic ; sepals 0.7-1.2 mm. long;
petals 1.1-1.6 mm. long; perianth caducous in fruit.
Ovules usually 4 per locule; leaves (as ee as known) 3-foliolate, all
of the leaflet-blades attenuate at base; .....5. W. vitiensis.
Ovules 8-12 per locule; lateral leaflet- one with the lower basal
margins obtuse or rounded: Samoa.

130 JOURNAL OF THE ARNOLD ARBORETUM | [vot. xxxur

Leaves 3—9-foliolate (very rarely simple), the leaflet-blades 1-2.5 cm.

NONE. ie oe 24-4. 1a 25555 © ede boas 6. W. samoensis.
Leaves sometimes 3-foliolate, usually simple, the blades 1.3-4 cm

Pu dGN Hee Se ed B4E ae boku W. manuana.

Leaflets small, 8-16 mm. _ long, 3-6 mm. broad, with 3-6 marginal crenations

per side; stipules i tena 2-4 mm. in diameter, strongly revolute;

racemes 2-3 cm. long; Fiji. .......................... 7. . exigua

1. Weinmannia affinis A. Gray, Bot. U. S. Expl. Exped. 1: 674. 1854;
C. Muell. in Walp. Ann. Bot. Syst. 5: 30. 1858; Seem. Fl. Vit. 110.
1865; Engl. in Linnaea 36: 648. 1870; Pampan. in Ann. di Bot. 2:
92, 1905; Gibbs in Jour. Linn. Soc. Bot. 39: 145. 1909.

Shrub or small tree, the branchlets glabrous or inconspicuously puberu-
lent distally; stipules chartaceous or subcoriaceous, elliptic or suborbicular-
obovate, entire, 13-25 % 10-15 mm., obtuse at apex, conspicuously
barbellate in axils, the hairs (pale, stiff, 1-1.5 mm. long) often sub-
persistent; leaves glabrous, simple, rarely 2-foliolate, the petioles (4—)
8-17 mm. long (20-25 mm. long in compound leaves, then the leaflets
sessile), the blades subcoriaceous, oblong-elliptic, (3.5—) 7-11 cm. long,
(1.5—) 2.5—-6 cm. broad, acute to cbtiee at base and decurrent, obtuse or
obtusely cuspidate at apex, conspicuously crenate-serrate with 1 or 2
crenations per centimeter, the venation obvious, the secondary nerves 7—14
per side, the veinlet-reticulation usually prominulous on both surfaces;
racemes usually paired or ternate at apices of peduncles (1.5—4 cm. long),
4-9 cm. long, the peduncle, rachis, and pedicels puberulent (hairs 0.1—0.2
mm. long), sometimes glabrate; flowers crowded, sometimes pseudovertic-
illate, the pedicels 1.3-2 mm. long or slightly shorter at anthesis: sepals
essentially glabrous, oblong, 1.2-1.5 0.7-1 mm., rounded at apex; petals
membranaceous, oblong, 1.6-1.8 0.9-1.2 mm., rounded at apex; disk-
lobes oblong-clavate, 0.5-0.7 mm. long; stamens with filiform filaments
up to 4 mm. long (in ¢ flowers) and anthers 0.3-0.4 mm. in diameter;
carpels ovoid, minutely hispidulous-puberulent, the styles less than 1 mm.
long at anthesis, the ovules usually 4 per locule (in % flowers, none or
undeveloped in ¢ flowers); perianth soon caducous; capsule oe
2.5—-4 mm. long, usually persistently puberulent, the styles up to 1.5 .
long; seeds 0.5—0.8 mm. long, copiously and persistently comate at ik
ends, the hairs 0.7—1 mm. long, many-celled, crispate.

DIsTRIBUTION: Endemic to Fiji, thus far known from Viti Levu, Ovalau, and
Taveuni but doubtless to be expected from other high islands, at elevations of
350-1200 m. (as far as recorded). It is a shrub or small tree, up to 7 m. in
height, usually occurring in dry forest or ridge forest or on dry open ridges,
occasionally in wetter localities. The petals and filaments are white and the
capsules red. Recorded local names are vure (Gillespie 2736) and katakata
(Smith 4905), names usually referred to the genera Geissois and Spiraeanthemum
rae The type, cited below, is an Exploring Expedition specimen from
Ovalau

FIJI: Vitt Levu: Seemann 197 (BM, GH, K); Mba: Mountains near
Lautoka, Greenwood 247 (K); vicinity we Nandarivatu, Gibbs 642 (BM, K),

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XII 131

733 (BM), 881 (BM, K), Greenwood 864 (A, K, US), Smith 4905 (A, US),
Vaughan 3405 (BM); Nandronga & Navosa: Southern slopes of
Nausori Highlands, above Tumbenasolo, Greenwood 1188 (A); Namosi:
Mt. Korombasambasanga, B. E. Parham 2201 (A); Mt. Voma, Gillespie 2736
oe B. E. Parham 602 (A), 2598 (A). Ovatau: U. S. Expl. Exped. (GH,

NY. US 48070 type), Graeffe (K). Taveunt: Borders of lake east of
eee Smith 878 (Bish, K, NY); Vuna, Seemann 200 (BM, GH, K). Fiji,
without definite locality: Horne 916 (GH, K), Gillespie 2730 (Bish).

The cited specimens probably give a fair picture of the variation to be
expected in W. affinis, which is without difficulty recognized by its simple
(only very rarely 2-foliolate) leaves with proportionately broad and
coarsely crenate blades. Its flowers are slightly larger than those of other
species of our region, and in general the indument is negligible. The type
material is among the larger in foliage-dimensions, and from it there is a
series of specimens toward such small-leaved forms as that found on
Taveuni (e. g. Smith 878, which also has the inflorescence strictly
glabrous). Gray’s var. 8, with 3-foliolate leaves, may be referred to W.
vitiensis, as suggested by Seemann. The closest ally of W. affints seems
to be the Samoan W. manuana, as noted below.

2. Weinmannia manuana Christophersen in Bishop Mus. Bull. 154:
10. fig. 2. 1938.

Weinmannia affinis sensu Reinecke in Bot. Jahrb. 25: 635. 1898; Chris-
tophersen in Bishop Mus. Bull. 154: 9. 1938; non A. Gray

Shrub or small tree, rarely epiphytic, the branchlets in distal internodes
strigose with hairs up to 1 mm. long or glabrous; stipules ovate or elliptic,
entire, 6-12 2-9 mm., obtuse or subacute at apex, soon glabrate, not
(or very obscurely) barbellate in axils; leaves simple or 3-foliolate, the
petioles 3-30 mm. long, often strigose like young branchlets but soon
glabrate, the blades (sessile in lateral leaflets of compound leaves, with
petiolules 5-10 mm. long and winged in terminal leaflets) chartaceous,
lanceolate, (3—) 4-10 cm. long, (1—) 1.3-4 cm. broad, acute to attenuate
at base and decurrent (or lateral leaflets of compound leaves with the
lower edge of base rounded), obtusely cuspidate or short-acuminate at
apex, crenate with (2—) 3 or 4 crenations per centimeter, the costa often
hirtellous beneath but soon glabrate, the secondary nerves 8—14 per side,
prominulous or nearly plane, the veinlet-reticulation copious, essentially
plane; racemes usually ternate at apices of peduncles (0.8—1.5 cm. long),
4—8 cm. long, the peduncle, rachis, and pedicels pale-puberulent, sub-
glabrate, the subtending bracts lanceolate, about 1 mm. long, caducous;
pedicels 1.5-2 mm. long (or slightly shorter at anthesis); sepals ovate-
oblong, 0.7—1.1 0.7-0.9 mm., rounded at apex; petals py ranean
oblong, 1.3-1.6 « 0.8-1 mm., ae at apex; disk-lobes 0.4-0.5 m
long; stamens with filiform filaments 1-1.5 mm. long (in % eo or
up to 3 mm. long (in ¢ flowers), the anthers 0.3-0.4 mm. in diameter;
carpels ovoid, strigose in bud, the ovules 10-12 (in & flowers, undeveloped

132 JOURNAL OF THE ARNOLD ARBORETUM | [voL. xxxim

in 6 flowers); perianth soon caducous; capsules ellipsoid, 3.5—4 mm. long,
obscurely puberulent or glabrate, the styles 1-1.5 mm. long; seeds 0.6-0.8
mm. long, comate at both ends, the hairs comparatively sparse, 0.4—0.5
mm. long and with inconspicuous cross-walls.

DIsTRIBUTION: Limited to Samoa and apparently to be expected throughout
the group, at elevations of 480-1500 m. It has been recorded as a shrub or
tree 1-7 m. in height, occurring in forest, in wet scrub-forest, on high ridges,
and on open “old lava fields. The type is Garber 1031, from Olosenga, cited
below

SAMOA: Sava: Above Aopo, Christophersen 889 (Bish, NY); Aopo-
Gagamalae, Pca eieiad 3446 (Bish); Matavanu crater, 828 (Bish, US),
2222 (Bish). Tutuma: Le Pioa, at top, Christophersen 1201 (Bish, NY),
3565 (Bish). OLosencA: Piumafua Mt., at top, Garber 1027 (Bish), 1031
(Bish type); Piumafua ridge, Garber 1066 (Bish). Tau: Trail to peak,
Garber 720 (Bish, US). Samoa, without definite locality: U. S. Expl. Exped.
(US 66032).

The specimens which Christophersen referred to W. affinis have been
carefully compared with the type and other Fijian specimens, and also
with W. manuana, and it appears to me that they represent the latter and
that true W. affinis does not occur in Samoa. In proposing W. manuana,
Christophersen separated it from W. affinis on the basis of its densely
hirtellous young branches and inflorescences and the more prominent
crenation of the leaves. While these characters are valid as far as the two
type collections are concerned, examination of other material of ~
relationship shows that the indument is too variable to be very useful;
the Fijian material the young branchlets and inflorescence vary fiom
copiously puberulent to glabrous, and in the Samoan material from
somewhat strigose or merely puberulent to glabrous. Characters pertain-
ing to leaf-shape and marginal crenations are apparent but are also very
variable, while the flowers (sepals, petals, and disk-lpbes) of W. affinis
are slightly larger than the corresponding parts in W. manuana. More
dependable characters in differentiating these species pertain to the stipules,
number of ovules, and seed-indument, but these characters are either
minute or transitory. Nevertheless it seems reasonable to separate the
Fijian and Samoan populations as indicated in my key.

3. Weinmannia spiraeoides A. Gray, Bot. U. S. Expl. Exped. 1: 677.
1854; C. Muell. in Walp. Ann. Bot. Syst. 5: 30. 1858; Seem. FI. Vit.
110. 1865; Engl. in Linnaea 36: 644. 1870.

Small tree, the branchlets, at least distally, copiously setulose with pale
hairs 0.5—0.8 mm. long; stipules suborbicular or ovate-oblong, about 10
7-10 mm., conspicuously dentate with 7-9 teeth, sparsely setulose on both
surfaces; leaves 5-foliolate (as far as known), the petioles 13-27 mm.
long, like the rachis and petiolules very slender, copiously hispidulous with
hairs 0.5—1 mm. long, the petiolules (of lateral leaflets) 1 mm. long or
less, of terminal leaflet 5-10 mm. long, the leaflet-blades chartaceous,

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XII 133

lanceolate-elliptic, (2.5-) 3-6 cm. long, 1—-2.5 cm. broad, acute to at-
tenuate at base, acute or obtusely cuspidate at apex, conspicuously serrate
with 3 or 4 teeth per centimeter, copiously hispidulous beneath especially
on nerves, often subglabrate above except on costa, the secondary nerves
5-9 per side, prominulous or nearly plane like the veinlet-reticulation ;
inflorescences unknown.

DistripuTION: Known only from the type collection, from the island of
Ovalau, Fiji, at about 150 m.
FIJI: Ovarau: U.S. Expl. Exped. (US 48073 Type).

The sterile specimen so optimistically described by Gray as a new
species has not yet been matched among more recent collections, and it
may conceivably represent a distinct species. However, the possibility
cannot be ignored that this specimen may be merely a juvenile form of
some other taxon, perhaps of W. richii. Nevertheless, juvenile forms of
W. richii so far available do not show the dentate stipules or the type of
leaf-indument described for W. spiraeoides, which for the time being is
accepted as a separate, if quite unsatisfactory, entity.

4. Weinmannia richii A. Gray, Bot. U. S. Expl. Exped. 1: 675. pl. 85,
B. 1854; C. Muell. in Walp. Ann. Bot. Syst. 5: 30. 1858; Seem. FI.
Vit. 110. 1865; Engl. in Linnaea 36: 643. 1870.

Weinmannia rhodogyne Gibbs in Jour. Linn. Soc. Bot. 39: 145. 1909;

Turrill in Jour. Linn. Soc. Bot. 43: 20. 1915.

Shrub or small tree, often compact, up to 7 m. high, the branchlets
sparsely to copiously puberulent distally with hairs 0.1-0.2 mm. long,
often glabrate; stipules chartaceous, suborbicular, 1.5-12 X 1.5-16 mm.,
rounded at apex, entire, sericeous-puberulent on both sides but usually
glabrate, strigose-tufted in axils; leaves 3—9-foliolate (very rarely simple),
the petioles 0.7—3 cm. long, puberulent like young branchlets or glabrous,
the rachis similar, narrowly winged or flattened above in distal internode,
the lateral petiolules up to 2 mm. long or essentially none, the terminal
petiolule 3-15 mm. long, distally winged, often puberulent, the blades
subcoriaceous or chartaceous, elliptic or oblong-elliptic, (1.5—-) 3-7 cm.
long, (1—) 1.2-3.8 cm. broad (terminal rarely to 9.5 x 4.5 cm.), acute to
attenuate at base and decurrent, obtusely cuspidate at apex, crenulate
with 2 or 3 crenations per centimeter, glabrous (or puberulent beneath on
costa and rarely on lower part of blades), the secondary nerves 7-9 per
side, with the veinlet-reticulation prominulous on both surfaces, or the
veinlets subimmersed; racemes 2—4 (often ternate) at apices of peduncles
(1-10 mm. long) or sometimes solitary, the peduncle, rachis, and pedicels
puberulent like young branchlets, rarely glabrate, the racemes 4-12
(-14.5) cm. long, the flowers crowded, subfasciculate in groups of 2-8;
pedicels at anthesis 0.7-1.5 mm. long, in fruit up to 2 mm. long; sepals
oblong-ovate, 0.5-0.7 0.4—0.6 mm., obtuse at apex, essentially glabrous
but sometimes sparsely pilose distally or ciliolate; petals membranaceous,

134 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxxm1

obovate-oblong, 1-1.3 0.6-0.8 mm., rounded or obscurely retuse at
apex; disk-lobes 0.2—0.4 mm. long; stamens with filiform filaments up to
2.5 mm. long, the anthers about 0.3 mm. in diameter; carpels ovoid,
crispate-pilose with pale hairs 0.2—0.4 mm. long, the styles up to 1.5 mm.
long. the ovules 4—6 per locule in & flowers; calyx and often petals
usually persistent, even after carpel has shattered; capsules narrowly
ellipsoid, up to 3.5 mm. long, sparsely soft-pilose, eventually subglabrate,
the styles persistent; seeds about 0.7 mm. long, comate at both ends, the
hairs 0.7—-1.5 mm. long, with obscure cross-walls.

DIsTRIBUTION: Limited to Fiji, thus far known from Viti Levu, Vanua Levu,
and Taveuni but doubtless occurring on other islands. Elevations of 100 to 1100
m. have been recorded, as well as a variety of habitats, such as open forest,
thickets, scrub, dry slopes, open country, etc. The species is a shrub or small
tree up to 7 m. in height, with pinkish to dark red peduncles, rachises, and
pedicels, white or greenish white petals and stamens, and white carpels which
become deep red during development and in fruit. A recorded local name is
vota (Smith 6813). The type is an Exploring Expedition specimen from Mbua
Bay [Sandalwood Bay], Vanua Levu, cited below.

FIJI: ae he Mba: Mountains near Lautoka, Greenwood 230 (K),
384 (A, K), 401 (K); vicinity of Nandarivatu, Gibbs 594 (BM type of W.
rhodogyne, ae im Thurn 73 (BM, K), Mead 1989 (K), Gillespie 4035 (Bish,

H. K, NY), 4233 (Bish, kK), Devener & Ordonez 13599 (A, Bish, K, NY),
Degener 14379 (A, Bish, K, NY, US), Smith 5052 (A, US), Vaughan 3228
(BM); slopes and ridges . Mt. Nanggaranambuluta [Lomalangi], east of
Nandarivatu, Gillespie 4333 (Bish, GH, NY), 4071.1 (Bish), Smith 5739 (A,

US); Nandronga & Navosa: Southern slopes of Nausori Highlands, in
drainage of Namosi Creek above Tumbenasolo, Smith 4710 (A, US). VaNnvua
Levu: Mbua: H. B. R. Parham, Jan. 3, 1937 (A, BM); Mbua Bay, JU. S.
Expl. Exped. (GH, K, NY, US 48071 TYPE): Ndama, B. E. Parham & M.
Sealolo 2277 (A); Wairiki, B. E. Parham 1122 (A); Mathuata: Seang-
gangga Plateau, in irinick of Korovuli River, vicinity of Natua, Smith 6813
(A, US). Taveuni: Above Somosomo, Gillespie 4837 (Bish, GH). Fiji, without
definite locality: Storck, set 1883 (BM, K), 25 (GH), Horne 1097 (K).

Although W. richii is the most abundant species of the genus in Fiji,
it does not seem widely dispersed throughout the group, and in my observa-
tion it is nowhere abundant; even at Nandarivatu, where many collectors
have obtained it, it is not a conspicuous element of the vegetation. As
here delimited, W. richii is readily distinguished from its closest relative,
W. vitiensis, by its suborbicular stipules, proportionately broader leaflets,
somewhat smaller flowers, and persistent perianth.

The characters utilized by Gibbs to separate W. rhodogyne from W.
richii are seen to be inconsequential when a series of specimens is examined.
It is true that the branchlets and leaves of the type of W. rhodogyne are
glabrous, whereas in the type of W. richii the young branchlets, as well
as the petiole, leaf-rachis, and costae of the leaflets beneath are setulose-
puberulent. However, every stage between these conditions is to be
found, even among plants from the vicinity of Nandarivatu, the type

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XII 135

locality of W. rhodogyne. Stipules of the two type specimens are similar
in shape, those of W. rhodogyne being much the smaller, but this appears
to be a matter of stage of development, as on other specimens stipules are
to be seen varying in diameter from about 2 to 15 mm., often on the
same plant. Differences in size of sepals and petals are inconsequential,
and filament-length seems to be a matter of state of development of the
flower, The ovary is similarly pilose in both concepts. Number of
leaflets cannot be utilized to divide the series of specimens cited above.
In the type of W. richii the leaflets vary between 3 and 9, whereas they
seem to be always 3 on Gibbs’ type. Other material shows the entire
range in this character, and occasionally even unifoliolate leaves are
found. In general, it can only be stated that the type of W. rhodogyne
and much of the other material from the Nandarivatu region is com-
paratively delicate and inclines to be less pubescent than typical specimens
of W. richii from the drier regions of leeward Vanua Levu. No reasonable
means has been found to separate W. rhodogyne from the older concept,
even varietally.

5. Weinmannia vitiensis Seem. Fl. Vit. 110. 1865; Pampan. in Ann.
di Bot. 2: 93. 1905.

Weinmannia affinis var. B A. Gray, Bot. U. S. Expl. Exped. 1: 674. 1854; C.
Muell. in Walp. Ann. Bot. Syst. 5: 30. 1858; Engl. in Linnaea 36: 649. 1870.
Shrub or small tree, the young parts and branchlets strigose-puberulent
or strigillose with whitish hairs 0.1—0.5 mm. long, soon glabrate; stipules
chartaceous, oblong or narrowly elliptic, 3-10 * 1.5-5 mm., rounded or
obtuse at apex, sparsely puberulent without or glabrate, strigose-tufted
in axils: leaves 3-foliolate (as far as seen), the petioles 4-18 mm. long,
glabrous, often narrowly winged distally, the petiolules winged nearly to
base (in lateral leaflets essentially none or up to 7 mm. long, slightly
longer in terminal leaflets), the blades subcoriaceous, lanceolate or
lanceolate-elliptic, (2.5—) 3—-5.5 cm. long, 1-2 cm. broad (terminal some-
times up to 7.5 2.8 cm.), attenuate at base and decurrent, obtuse or
obtusely cuspidate at apex, crenulate with about 3 crenations per centi-
meter, glabrous on both sides, the secondary nerves 6-12 per side, with
the veinlet-reticulation sharply prominulous on both sides, or the veinlets
nearly plane; racemes ternate at apices of peduncles (these very short
and insignificant or up to 2 cm. long) or perhaps sometimes solitary, the
peduncle, rachis, and pedicels strigose-puberulent like young branchlets, at
length glabrate, the racemes 4-7 cm. long; flowers scattered or sub-
fasciculate in groups of 2-4, subtended by caducous oblong bracts up to
1 mm. long, the pedicels slender, about 1 mm. long at anthesis and up to
2 mm. long in fruit; sepals oblong, 1-1.2 0.7—0.8 mm., rounded at apex,
glabrous; petals oblong, 1.4-1.6 0.8-1 mm., rounded at apex; disk-lobes
0.5-0.6 mm. long; stamens with filiform filaments up to 1.7 mm. long,
the anthers about 0.3 mm. in diameter; carpels ovoid, sparsely strigose,
the styles at anthesis less than 1 mm. long, the ovules 4 per locule in &

136 JOURNAL OF THE ARNOLD ARBORETUM | [vot. xxxut

flowers; calyx and petals caducous in fruit, leaving a flattened receptacle;
capsules narrowly ellipsoid, 3-3.5 mm. long, sparsely puberulent or
soon glabrate, the styles up to 1 mm. long; seeds narrowly ellipsoid, about
0.8 mm. long, comate at both ends, the hairs 0.5—1 mm. long, with obscure
cross-walls.

DIsTRIBUTION: Endemic to Fiji, thus far known from three of the smaller
islands but to be expected elsewhere in the group. The species occurs at eleva-
tions up to 400 m., from the scanty data thus far available, in dense forest or in
open places as a compact shrub or tree up to 8 m. in height. The petals and
filaments are white and oe mature capsules brown. On Moala I recorded the
local name as molau ndam

FIJI: Kanpavu: Sine 199 (GH, K type). Ovatau: U.S, Expl. Exped.
(source of the Dieuienase to W. affinis var. 8, GH, NY). Moata: Summit ridge,
Bryan 317 (Bish); Ndelaimoala, Smith 1354 (Bish, GH, K, NY, US). Fiji,
without definite locality’: Harvey (GH, K).

This apparently uncommon species is distinguished from W. richii by
characters pertaining to stipules, foliage, and perianth, as noted above.
The species is one of the conspicuous elements in the rather dry, low
forest and open scrub on the island of Moala, but I have not personally
observed it elsewhere.

6. Weinmannia samoensis A. Gray, Bot. U. S. Expl. Exped. 1: 677.
1854; C. Muell. in Walp. Ann. Bot. Syst. 5: 30. 1858; Engl. in
Linnaea 36: 647. 1870; Reinecke in Bot. Jahrb. 25: 634. 1898;
Pampan. in Ann. di Bot. 2: 92. 1905; Rechinger in Denkschr. Akad.
Wiss. Wien 85: 286. 1910; Setchell in Carnegie Inst. Publ. 341:
92. 1924; Christophersen in Bishop Mus. Bull. 154: 11. 1938.

Weinmannia samoensis {. glabrescens Pampan. in Ann. di Bot. 2: 92. 1905.

Shrub or small tree, the branchlets hispidulous-puberulent with hairs
0.1-0.7 mm. long, at length glabrate; stipules elliptic or lanceolate-
elliptic, 5-15 3.5-10 mm., entire, rounded or obtuse at apex, sparsely
puberulent and glabrate; leaves 3-7-foliolate (rarely 9-foliolate, very
rarely simple), the petioles (1—) 1.5-4 (rarely to 6) cm. long, at first
hispidulous, soon glabrate, the rachis similar, usually narrowly winged in

distal internode, the petiolules essentially none in lateral leaflets, 3-8

(-13) mm. long in terminal leaflets, winged nearly to base and some-

times hispidulous; leaflet-blades chartaceous, lanceolate, (3—) 4-9 (-12)

cm. long, (0.7—) 1-—2.5 cm. broad, attenuate at base and decurrent (lateral
leaflets with lower basal margin obtuse or narrowly rounded), narrowed to
an acuminate or obtusely cuspidate apex, crenulate with 2 or 3 crenations
per centimeter, glabrous except the costa sometimes hispidulous like
petiole, the secondary nerves 8-14 per side, short, prominulous on both
surfaces, the veinlet-reticulation immersed or faintly prominulous; racemes
usually ternate (sometimes paired) at apex of peduncles, a subsidiary
pair sometimes arising from a lower node of inflorescence-rachis, the
peduncle 1.5—2.5 cm. long, like the rachis and pedicels puberulent, even-

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XII 137

tually glabrate, the racemes 5-8 cm. long; flowers mostly single on the
rachis, on pedicels 1.5—2.3 mm. long; sepals deltoid-oblong, 0.8-1 x 0.5-
0.7 mm., subacute, sometimes faintly puberulent without; petals mem-
branaceous, oblong-ovate, 1.1-1.5 & 0.7-1 mm., rounded at apex; disk-
lobes elongate, 0.4-0.7 mm. long; stamens with filiform filaments
2-3 mm. long (in ¢ flowers) or up to 1.5 mm. long (in & flowers),
the anthers about 0.4 mm. in diameter; carpels ovoid, faintly hispidulous-
puberulent, glabrate, the styles up to 1.2 mm. long (in &% flowers) or
shorter (in ¢ flowers), the ovules 8-10 per locule in &% flowers, un-
developed in ¢ flowers; calyx and petals caducous in fruit, the receptacle
flattened; capsule ellipsoid, 2.5-3 mm. long, puberulent like rachis but
soon glabrate; seeds narrowly ellipsoid, 0.6—-0.7 mm. long, comate at both
ends, the hairs usually about 0.7 mm. long.

DISTRIBUTION: Samoa, recorded from the three large islands at elevations
of 300-1000 m. The species is said to be a shrub or tree 2-6 m. in height; habitat
data are sparse, but Christophersen describes it as growing on lava fields and on
river-banks. The type is an Exploring Expedition specimen from Tutuila, cited
below. Additional collections were cited by Reinecke and Rechinger.

SAMOA: Savair: Central region, Reinecke 538 (BM, K, US); Asana, back
of Sologa, Vaupel 373 (Bish, K, NY, US); Matavanu lava field, Christophersen
& Hume 1943 (Bish); above Sili, oe a (Bish, US).

Above Vailele, Reinecke 567 (Bish). TUILA: U. Expl. Exped. (GH, K
US 48072 type). Samoa, without a. locality: wae 218 (K), Powell
(GH), 323 (K).

Weinmannia samoensis, a species characterized by its narrow lanceolate
leaflet-blades, is distinguished from the preceding, W. vitiensis, by the
greater number of ovules. This character, being observable only in
pistillate flowers, is not very practical but nevertheless seems dependable
in the material at hand. The difference in the base of the lateral leaflets,
utilized in my key, also seems fairly constant. It is not always easy to
separate specimens of W. samoensis and W. manuana, the only other
described Samoan species, when the latter has 3-foliolate leaves. In
general the leaves of W. samoensis are 5—7-foliolate and those of W.
manuana are simple; when 3-foliolate leaves occur on Samoan plants
they are also accompanied by one or the other more characteristic type, in
my observation. Nevertheless this character is not entirely satisfactory
and must be supplemented by the shape of the leaflet-blades, which are
somewhat broader in W. manuana. I find no consistent differences between
the two species in indument or inflorescence.

Pampanini’s forma glabrescens is based upon Reinecke 502, 538, and
567. Two of these numbers have been examined and I do not observe any
reason to separate them from the population as a whole.

7. Weinmannia exigua sp. nov.

Frutex, partibus novellis copiose cinereo-strigoso-puberulis, ramulis
superne subcomplanatis et puberulis demum teretibus glabratis; stipulis

138 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxurr

chartaceis suborbicularibus 2-4 mm. diametro, dorso copiose strigosis,
margine valde revolutis, caducis, basi intus strigoso-barbellatis; foliis
apices ramulorum versus congestis 3- vel 5-foliolatis raro simplicibus
vel 7-foliolatis, petiolis 4-9 mm. longis (vel 2—4 mm. in foliis simplicibus)
superne anguste alatis, rhachi etiam anguste alata ut petiolulis puberula,
petiolulis lateralibus subnullis terminalibus 1-4 mm. longis alatis, laminis
chartaceis in sicco fuscis anguste ellipticis, (S—) 8-16 mm. longis, 3-6 mm.
latis, basi (lateralibus) obtusis vel (terminalibus) attenuatis, apice
obtusis vel obtuse cuspidatis, margine dentibus utrinsecus 3-6 crenatis,
supra glabris, subtus praecipue costa pilis 0.2-0.3 mm. longis strigoso-
puberulis demum subglabratis, costa supra subplana subtus elevata, nervis
secundariis utrinsecus 3—6 inconspicuis subtus prominulis, rete venularum
saepe immerso; racemis summo pedicelli brevis (ad 5 mm. longi) 2 vel
3 vel videtur solitariis 2-3 cm, longis, pedunculo ut rhachi pedicellisque
minute puberulo; pedicellis sub anthesi 1-1.5 mm. longis; sepalis 4
subliberis papyraceis oblongis, 0.7-0.8 mm. longis, 0.4—0.5 mm. latis, apice
obtusis, superne obscure ciliolatis; petalis 4 membranaceis obovatis, 1.2—
1.3 mm. longis, 0.6-0.7 mm. latis, apice rotundatis, basi angustatis; disci
lobis 8 oblongis 0.3—-0.4 mm. higuis apice truncatis; staminibus 8, in
floribus & filamentis filiformibus 1.2—1.5 mm. longis, antheris circiter
0.3 mm. diametro; carpellis ovoideis sub anthesi 1-1.5 mm. longis, pilis
pallidis 0.2-0.3 mm. longis copiose hirtellis, stylis erectis circiter 1.5 mm.
longis, ovulis 3—6 per loculo; perianthio videtur caduco; capsulis ellipsoid-
eis circiter 3 mm. longis glabratis, seminibus ellipsoideis 0.6-0.7 mm.
longis utroque conspicue comatis, pilis stramineis circiter 1.5 mm. longis.

DIsTRIBUTION: Fiji, known only from the type collection.

FIJI: Vanua Levu: Thakaundrove or Mathuata: Between
Waiwai and Lomaloma, May 1878, Horne 632 (K type) (large shrub about 3 m.
high, on top of the mountains).

The very distinct entity here described differs from other species of the
region in its very small leaves and compact inflorescences.

Weinmannia sp.

Weinmannia richii (?) sensu Christophersen in Bishop Mus. Bull. 154:

11. 1938; non A. Gra

SAMOA: Savair: Tuisivi Range, alt. 1600-1700 m., Christophersen 787
(Bish, NY); above Matavanu, alt. about 1600 m., Christophersen 2561 (Bish);
rim of Papafu crater, alt. 1500 m., Christophersen 2735 (Bish

Although the cited specimens are sterile, they apparently do not repre-
sent either known Samoan species, W. samoensis or W. manuana, unless
juvenile forms in this alliance are extremely variable. Although these
specimens bear a general resemblance to W. richii, I see no reason to refer
them here on the basis of present material; the sterile Samoan specimens
are inclined to have crenulate stipules and longer petiole-indument, al-
though they do not agree too well among themselves. These collections

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XII 139

suggest that an undescribed species is present on Savaii, unless they are
extreme juvenile variants.

A sterile specimen from Upolu (above Saluafata, ridge to Maunga Tele,
alt. 830 m., Christophersen 534 [Bish, US]) cannot be placed at present;
it is neither precisely like the Savaii specimens mentioned above nor does
it seem to represent either W. samoensis or W. manuana.

3. SPIRAEANTHEMUM A. Gray

Spiracanthemum was described by Gray (Bot. U.S. Expl. Exped. 1: 666.
1854) on the basis of two species, one Samoan and one Fijian. Although
these two species have been taken as congeneric by subsequent
students, and although in my opinion this is a reasonable interpretation,
it is possible they will eventually be placed in at least different sections
or perhaps subgenera. I do not find that anyone has made the selection of
a genotype, and therefore I should like so to designate S. samoense, Gray’s
first species and the one which in basic characters seems best to agree
with the greater number of subsequently described species of Spiraeanthe-
mum,

Approximately 27 binomials have thus far been proposed in Spiraean-
themum, which has a range from New Guinea, Australia, and New Cale-
donia to Samoa; it is evidently lacking in Tonga. Like so many of the
genera first described from Fiji or Polynesia, Spiraeanthemum proves to
have its center of distribution in New Guinea and New Caledonia; from
the former island eight species are discussed by L. M. Perry (in Jour.
Arnold Arb. 30: 139-143. 1949). In our area five species are discernible,
four from Fiji and one from Samoa. Although individuals of the genus
are seen fairly frequently in Fiji and Samoa, for the most part they occur
singly and do not form a striking feature of the vegetation. A common
name for the genus in Fiji is katakata.

Spiracanthemum vitiense differs markedly from the other species of our
region in its verticillate leaves, short stipule-scars, solitary ovules, and
seeds with a distal wing only; the remaining species have opposite leaves,
elongate and curved stipule-scars, paired ovules, and seeds winged at both
ends. Usable characters to differentiate the species are found in the type
of indument, leaf-margins, stipule-shape and indument, etc. In general
these characters are not strong, but they seem more adequate than those
one is forced to utilize in Weinmannia. All of our species are probably
dioecious, the staminate flowers lacking carpels and the pistillate flowers
having probably sterile anthers; it is possible, however, that these anthers
are sometimes fertile and the species thus occasionally polygamo-dioecious.

KEY TO THE SPECIES
Leaves verticillate; stipules leaving inconspicuous, transversely elliptic, nearly
straight scars; leaf-blades obovate-elliptic, rounded or broadly obtuse at
apex, the secondary nerves 4-6 per side; inflorescence 2-6 cm. long, the
branches usually ternate; carpels 1-ovulate, the seed with a distal wing
ony, the nucellus basal: Fj ci. ea seo eek 1. S. vitiense.

140 JOURNAL OF THE ARNOLD ARBORETUM _[vot. xxx

Leaves opposite; stipules leaving curved, elongate scars; leaf-blades raion
- or elliptic-oblong, acuminate or cuspidate or callose-apiculate
apex, the secondary nerves 5-11 per side; inflorescence 5-17 cm he
the branches opposite or subopposite; carpels 2-ovulate, the seeds with
distal and basal wings, the nucellus median.

Branchlets and petioles glabrous or distally evanescently strigose-puberulent,
he leaf-blades glabrous on both surfaces (rarely sparsely puberulent on
costa when young); Fijian species.

Leaf-blades usually less than twice as long as broad, obtuse at base and
abruptly decurrent on the petiole, entire or inconspicuously serrulate at
margin, the teeth obsolete or 1 or 2 per centimeter; peduncle of in-

orescence usually more than 4 cm. long. ..... 2. S. graeffet.

Leaf-blades usually more than twice as long as broad, attenuate to acute at
base and long-decurrent on the petiole, conspicuously serrate at margin
ie 3 or 4 teeth per centimeter; peduncle of inflorescence 8 than

cm, long. ratum.

Grane and petioles copiously velutinous-puberulent or fea ai
tardily glabrate, the leaf-blades puberulent or oo at least on costa
and secondaries ‘beneath, the indument persis

Indument of branchlets and petioles sh aon je (hairs 0.1-0.15

. long, very dense, long-persistent); st ipules oblong-ovate, up to

m., velutinous-puberulent or sericeous on both surfaces;

leaf-blades entire or inconspicuously denticulate, the costa and second-
aries minutely puberulent beneath (hairs scarcely 0.1 mm. long);

ultimate branchlets of inflorescence (below pedicel- articulation)

sampienanag or to 0.6 mm. long; disk-lobes usually or! setulose

OF BRR: FU. os ksh eGekyedwsdavassreiieawvesveaas . katakata.

Indument of eer and petioles hispidulous (hairs ae 1 mm. long);
stipules lanceolate-oblong, comparatively narrow, up to 35 & 10 mm.,
sericeous or hispidulous without, glabrous within; leaf-blades con-
spicuously serrulate with 2-4 teeth per centimeter, the costa and
secondaries strigose-puberulent beneath (hairs 0.2-0.7 mm. long);
ultimate branchlets of inflorescence (below pedicel-articulation) 0.7—2.5
mm. nai disk-lobes sparsely each or glabrous; Samoa. .. ;

S. samoense.

1. Spiraeanthemum vitiense A. Gray, Bot. U. S. Expl. Exped. 1: 669.
pl. 83, B. 1854, in Ann. Sci. Nat. IV. Bot. 4: 177. 1855, in Proc. Am.
Acad. 3: 128. 1857; C. Muell. in Walp. Ann. Bot. Syst. 5: 24. 1858;
Seem. FI. Vit. 111. 1865; Gibbs in Jour. Linn. Soc. Bot. 39: 144. 1909.

Shrub or small tree, up to 3 m. high, presumably dioecious or possibly
polygamo-dioecious, the branchlets terete, glabrous, the young parts ob-
scurely glandular; very young stipules ovate, glabrous and obscurely
glandular, soon caducous, the scars inconspicuous, transversely elliptic,
nearly straight; leaves verticillate, in threes or fours (rarely in fives), the
petioles shallowly canaliculate or semiterete, 4-20 (—23) mm. long, gla-
brous or obscurely glandular, distally winged; leaf-blades coriaceous, obo-
vate-elliptic, 4-8 (—10.5) cm. long, 1.5-4.5 (—5) cm. broad, acute to at-
tenuate at base and long-decurrent on the petiole, rounded or broadly
obtuse at apex, narrowly recurved and entire at margin, glabrous, the costa

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XII 141

slightly elevated above and prominent beneath, the secondary nerves 4-6
per side, arcuate-ascending, usually nearly plane above and elevated be-
neath, often with inconspicuous domatia in the axils beneath, the veinlet-
reticulation intricate, usually plane or immersed above and prominulous
beneath; inflorescence paniculate, axillary, solitary, compact, many-
flowered, 2-6 cm. long and nearly as broad, the peduncle slender, 7-16 mm.
long, very minutely puberulent and soon glabrate, the branches usually
ternately arranged, more obviously puberulent than peduncle but sub-
glabrate, the bracts lanceolate-oblong or subfoliaceous, up to 7 mm. long
or even approximating leaves in size, soon glabrate, the ultimate bracteoles
minute, 0.2—-0.3 mm. long; é flowers not seen; ? (or perhaps 8 ) flowers
in clusters of 2-6, each actually solitary at apex of a minute (0.1-0.5 mm.
long) ultimate branchlet, the pedicels (above articulation) 0.5-1 mm. long
(to 2.3 mm. long in fruit), minutely puberulent, glabrate; calyx L215
mm. long, glabrous or very sparsely puberulent without, deeply 4-lobed
(rarely 5- or 6-lobed), the lobes ovate, 0.7-1 mm. broad, subacute;
stamens 8 (rarely 10 or 12), the filaments glabrous or very sparsely pale-
pilose, 0.7-1.5 mm. long, the anthers broadly ellipsoid, 0.2-0.4 mm. long,
dubiously functional; disk-lobes 8 (rarely 10 or 12), free or rarely a pair
connate, carnose, angular-obovoid, 0.3—0.4 mm. long, truncate at apex,
glabrous; carpels 4 (rarely 5 or 6), ovoid, faintly sericeous, the style
0.7-1.2 mm. long, the ovule solitary, pendulous from near middle; calyx
and stamens persistent in fruit, some carpels often aborting; mature carpels
ovoid, 2.2-3 mm. long, sparsely hirtellous (hairs 0.1-0.2 mm. long) or
essentially glabrate, the style persistent; seed solitary, oblong, 2-2.5 mm.
long, 0.8-1 mm. broad, the nucellus ellipsoid, exceeded distally by a wing
0.8-1.2 mm. long, this rounded at apex, the basal wing lacking.

Viti Levu and Vanua Levu, at elevations of 450-1200 m. The species has been
noted as a shrub or small tree, up to 3 m. in height, growing in open or ridge-
scrub (Parham) or in a forest clearing (Gibbs). The type is an Exploring
Expedition collection, cited below, apparently obtained from two localities,
Sandalwood Bay [Mbua Bay] and Mathuata, Vanua Levu.

FIJI; Vitr Levu: Graeffe 16 in part (BM, K); Mba: Tholo-i-Nandarivatu,
Gibbs 732 (BM); Namosi: Summit of Mt. Voma, B. E. Parham 1743 (A),
1910 (A). Vanua Levu: Mbua and Mathuata: Mbua Bay (part) and
presumably Mathuata coast (part), U. S. Expl. Exped. (GH, K, NY, US 47621
TYPE). Fiji, without definite locality: Horne 759 (GH, K), 1104 (K), 1113 (K).

As indicated in my key and generic discussion, S. vitiense is a strikingly
distinct species both in vegetative features and in the more fundamental
characters of the ovulation and seed-shape.

2. Spiraeanthemum graeffei Seem. Fl. Vit. 111. 1865; Gibbs in Jour.
Linn. Soc. Bot. 39: 145. 1909.

Shrub or tree up to 6 m. high, dioecious, the branchlets slender, glabrous
or the youngest parts very obscurely strigillose-puberulent; very young

142 JOURNAL OF THE ARNOLD ARBORETUM _[VoL. xxx

stipules ovate and dorsally sericeous, the older ones oblong or ovate-oblong,
up to 22 mm. long and 10 mm. broad, obtuse, very sparsely strigose-
puberulent on both surfaces or glabrate, soon caducous, the scars elongate,
curved; leaves opposite, the petioles semiterete, 1-3 cm. long, glabrous,
distally winged; leaf-blades coriaceous or subcoriaceous, lanceolate or
ovate-elliptic, 5-10 cm. long, (2—) 2.5-6 cm. broad, obtuse at base and
abruptly decurrent on the petiole, acuminate or obtusely cuspidate at apex,
entire or inconspicuously serrulate at margin (teeth minute, callose-glandu-
lar, 1 or 2 per centimeter, or obsolete), glabrous on both surfaces, some-
times with inconspicuous axillary domatia on lower surface, the costa nearly
plane above and prominent beneath, the secondary nerves 5—9 per side,
arcuate-ascending, plane above, elevated beneath, the veinlet-reticulation
intricate, prominulous on both surfaces; inflorescence paniculate, solitary,
axillary, ample, many-flowered, 6-15 cm. long and nearly as broad, the
peduncle slender, glabrous, (1—) 4—7 cm. long, the branches opposite or
subopposite, very sparsely puberulent with hairs about 0.1 mm. long, often
glabrate, the bracts oblong or lanceolate, up to 7 mm. long, essentially
glabrous, the ultimate bracteoles about 0.5 mm. long; flowers solitary at
apices of short ultimate branchlets (0.2—1.5 mm. long), the pedicels (above
articulation) 0.5-1.5 mm. long, essentially glabrous; calyx subcarnose,
1.5—2.2 mm. long, deeply 4-lobed, the lobes 1-1.3 & 0.7—1 mm., subacute;
stamens 8, the filaments glabrous, in ¢ flowers 2.5—-3 mm. long, in ?
flowers 0.8—-1.3 mm. long, the anthers broadly ellipsoid, in ¢ flowers about
0.4 mm. long, in ? flowers minute and apparently sterile; disk-lobes in
$ flowers 4, obovoid-angled, 0.4—-0.6 mm. long, free or loosely connate,
truncate or often emarginate at apex and hispidulous there and ventrally
with hairs 0.3-0.4 mm. long; disk-lobes in ? flowers 8, essentially similar
but sometimes more obscurely setulose or strictly ale bronse carpels in 9
flowers 4, ovoid, copiously sericeous-puberulent (hairs 0.1—0.2 mm. long),
the style 0.3-0.5 mm. long, the ovules 2, collateral, narrowed at both ends;
calyx and stamens persistent in fruit; mature carpels (1 or more some-
times aborted) elongate-ovoid, 3.5—-4 mm. long, 0.7-1 mm, broad, per-
sistently pilose, the style persistent; seeds 2, collateral, 2.5-3 mm. long,
the nucellus ellipsoid, about 1 mm. long, the wings subequal in length, the
distal wing lanceolate, slightly narrower than nucellus, the basal wing
subulate.

DISTRIBUTION: Endemic to Fiji, apparently infrequent, now known from
Viti Levu and Kandavu at elevations of 870-1050 m. The species is reported as
a shrub or tree, up to 6 m. in height, occurring in dense forest or in forest-
clearings; the calyx and filaments are white and the anthers yellow. Recorded
local names are katakata (Smith) and kutakuta (Gillespie). The type, collected by
Graeffe on Kandavu, is cited below

FIJI: Vitr Levu: Mba: Tholo- a ridge, Gibbs 731 (BM);
Naitasiri: Northern portion of Rairaimatuku Plateau, between Mt.
Tomanivi and Nasonggo, Smith 5800 (A, US): N amosi: Summit of Mt.
Voma, Gillespie 2728 (Bish, GH, K, NY, US). Kanpavu: Mt. Mbuke Levu,
Graeffe 16 in part (BM, K Type).

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XII 143

Spiraeanthemum graeffei and S. serratum are readily distinguished from
S. katakata by their usually strictly glabrous branchlets and leaves, these
parts bearing a long-persistent velutinous-puberulent indument in S. kata-
kata. Characters pertaining to the indument seem much more reliable here
than in the genus Weinmannia.

3. Spiraeanthemum serratum Gillespie in Bishop Mus. Bull. 83: 11.
fig. 11. 1931.

Small tree, up to 4 m. high, dioecious, the branchlets slender, glabrous
or distally strigose-puberulent (hairs pale, 0.1-0.2 mm. long); very young
stipules ovate, obtuse, copiously sericeous dorsally, glabrate, the scars
conspicuous, curved; leaves opposite, the petioles semiterete, 1-2 cm. long,
glabrous or faintly puberulent like young branchlets, distally winged;
leaf-blades subcoriaceous, lanceolate- or oblong-ovate, 4-8 cm. long, 1.3-—
3.8 cm. broad, attenuate or acute at base and long-decurrent on the petiole,
gradually acuminate or obtusely cuspidate at apex, conspicuously serrate
at margin (teeth callose-glandular, 3 or 4 per centimeter), glabrous on
both surfaces or sparsely puberulent on costa, sometimes with axillary
domatia on lower surface, the costa slightly elevated above and prominent
beneath, the secondary nerves 6-10 per side, subascending or arcuate,
plane above, slightly elevated beneath, the veinlet-reticulation intricate,
prominulous on both surfaces or subimmersed above; inflorescence panicu-
late, solitary, axillary or pseudoterminal, ample, many-flowered, 6-12 cm.
long and nearly as broad, the peduncle slender, glabrous or fatally pu-
berulent, 2—4 cm. long, the branches opposite or subopposite, the distal
portions strigose-puberulent with hairs 0.1-0.3 mm. long, the bracts
papyraceous, oblong, about 2 mm. long, soon glabrate, the ultimate bracte-
oles about 0.5 mm. long; flowers scattered, not fasciculate, solitary at
apices of short ultimate branchlets (up to 0.8 mm. long or essentially none),
the pedicels (above articulation) 0.6-1 mm. long, glabrous at anthesis;
calyx carnose, glabrous, 1.6—2 mm. long, deeply 4- (rarely 5-) lobed, the
lobes oblong-ovate, 1.2—1.5 >< 0.8-1 mm., obtuse or subacute; stamens 8
(rarely 10), the filaments glabrous, in ¢ flowers 1.2—1.6 mm. long, in @
flowers less than 1 mm. long, the anthers ellipsoid, in 4 flowers 0.3—0.4
mm. long, in @ flowers minute and apparently sterile; disk-lobes in ¢
flowers 4-6, carnose, free or loosely connate within stamens, irregularly
obovoid, 0.3—-0.5 mm. long, truncate or emarginate at apex, glabrous; disk-
lobes in @ flowers 8 (rarely 10), essentially similar, sometimes sparsely
setulose at apex (hairs few, to 0.3 mm. long); carpels in @ flowers 4
(rarely 5), ovoid, copiously sericeous (hairs 0.1-0.2 mm. long), the style
0.5-0.7 mm. long, the ovules 2, collateral, falcate, narrowly winged at both
ends; mature carpels narrowly ellipsoid, up to 3.5 mm. long and 0.8 mm.
broad, sericeous, the style persistent; seeds paired or only 1 developing,
lanceolate, 2.5—2.7 mm. long, 0.3-0.5 mm. broad, the nucellus ellipsoid,
less than 1 mm. long, the distal and basal wings subequal, very narrow,
subacute.

144 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxu

DIsTRIBUTION: Fiji, apparently limited to the summits and upper slopes of
a few high hills, at elevations of 1100-1323 m., on Viti Levu and Taveuni. The
species is recorded as a small tree, up to 4 m. in height, occurring in the dense
thickets of high ridges. It is noteworthy that the localities thus far known are
the four highest mountains in Fiji. The type, Gillespie 4107, is cited below.

FIJI: Virr Levu: Mba: Mt. Evans Range [presumably Mt. Mbotilamu],
Greenwood 364 (K), 457 (K); summit of Mt. Tomanivi [Mt. Victoria],
Gillespie 4107 (Bish type, GH), 4122.1 (Bish, K, NY); Namosi: Summit
ridge of Mt. Kor ombasambasanga, B. E. Parham 2200 (A), TAVEUNI: Summit
of Uluingalau, Smith 891 (Bish, GH, K, NY, US)

On the basis of type collections, it would appear that S. serratum is very
distinct from S. graeffei, but actually the suite of specimens now available
shows that the two are closely related. Differences in the leaf-margin and
the other points mentioned in my key seem to provide adequate grounds
for the maintenance of Gillespie’s species.

4. Spiraeanthemum katakata Seem. in Bonplandia 10: 36, nomen.
1862, Fl. Vit. 111. pl. 17. 1865; Pampan. in Ann. di Bot. 2: 51. 1905;
Gibbs in Jour. Linn. Soc. Bot. 39: 145. 1909.

Spiraeanthemum samoense sensu Gibbs in Jour. Linn. Soc. Bot. 39: 145. 1909;
non A. Gray.

Spiraeanthemum parksu Gillespie in Bishop Mus. Bull. 83: 10. fig. 10. 1931.
Shrub or tree up to 15 m. high, dioecious, the branchlets terete or distally
flattened, copiously velutinous-puberulent toward apices with pale spread-
ing hairs 0.1-0.15 mm. long, the older parts cinereous, glabrate; stipules
oblong-ovate, rapidly enlarging in size, up to 15 mm. long and 10 mm.
broad, obtuse, densely velutinous-puberulent on both surfaces or closely
sericeous, soon caducous, the scars elongate, curved; leaves opposite, the
petioles semiterete, 0.8-4 cm. long, narrowly winged distally, copiously
puberulent like branchlets, tardily glabrate or not; leaf-blades papyrace-
ous or subcoriaceous, ovate to lanceolate- or ovate-elliptic, 4-14 cm. long,
1.5~8.5 cm. broad, rounded to obtuse or rarely acute at base and abruptly
decurrent on the petiole, obtusely cuspidate or short-acuminate at apex,
narrowly revolute and entire at margins or undulate or inconspicuously
denticulate (teeth if present minute, 1-3 per centimeter), obscurely
puberulent on costa above, obviously puberulent on costa and secondaries
beneath, otherwise glabrous on both surfaces, usually with obvious axillary
domatia on lower surface, the costa nearly plane above and prominent be-
neath, the secondary nerves 5—11 per side, arcuate-spreading, usually plane
above and sharply elevated beneath, the veinlet-reticulation intricate, usu-
ally slightly prominulous on both surfaces; inflorescence paniculate, solitary,
axillary or pseudoterminal, ample, many-flowered, 5-13 cm. long, 4-8 cm.
broad, the peduncle slender, 1-4 (—5) cm. long, copiously and persistently
puberulent like young branchlets, the branches opposite or subopposite,
persistently puberulent and sometimes also hispidulous with pale hairs to
mm. long, the bracts oblong, papyraceous, to 2.5 mm. long, rarely

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XII 145

larger and subfoliaceous, puberulent on both sides, the ultimate bracteoles
about 0.5 mm. long; flowers solitary or in fascicles of 2—5, each actually
terminal on a minute ultimate branchlet to 0.6 mm. long or insignificant,
the pedicels (above articulation) 0.5—1.4 mm. long, copiously but minutely
puberulent; calyx 1.3-1.8 mm. long and broad, sparsely puberulent or
glabrate, deeply lobed, the lobes usually 4, rarely 3 or 5, ovate-oblong,
0.7-1.2 & 0.6-1 mm., subacute and minutely cucullate; stamens 8 (rarely
6 or 10), the filaments glabrous, in ¢ flowers 2—2.6 mm. long, in @ flowers
0.6—1.2 mm. long, the anthers ellipsoid, in ¢ flowers 0.3-0.4 mm. long, in
2 flowers minute and apparently sterile; disk-lobes in ¢ flowers usually
4 or 5, rarely 3, free or loosely connate, carnose, obovoid-angled, 0.4—0.7
mm. long, irregularly truncate at apex and setulose or crispate-pilose with
pale hairs 0.3-0.6 mm. long; disk-lobes in @ flowers 8 (rarely 6 or 10),
essentially similar, the apical hairs shorter (0.2-0.3 mm. long) or some-
times lacking; carpels in 9 flowers usually 4, rarely 3 or 5, ovoid, closely
sericeous (hairs about 0.1 mm. long), the style 0.5-1 mm. long, the ovules
2, collateral, attached near middle, narrowed and winged at both ends;
calyx and stamens persistent in fruit; mature carpels (1 or more sometimes
aborted) falcate- or lanceolate-ellipsoid, 2.5-3 mm. long, persistently
puberulent, the style persistent; seeds 2, collateral, 2.2-2.8 mm. long, the
nucellus ellipsoid, about 0.8 mm. long, the wings subequal in length, the
distal wing oblong, the basal wing subulate.

DISTRIBUTION: Fiji, where it appears to be the most abundant species of the
genus, although it is known from only a few islands and is nowhere an obvious
feature of the vegetation. A wide range has been recorded for altitudinal
occurrence (100-1195 m.) and habitat (dense forest, dry forest, open places,
forest-grassland transition, dense ridge thickets and forest, etc.). The species
is a shrub or tree, sometimes attaining 15 m. in height; the calyx and filaments
are white or greenish white, the styles are white often flushed with pink, and
the mature carpels are dull pink. The most frequently recorded local name is
katakata, but other names, some perhaps questionable, are kutakuta, tandalo,
vurewai, rure, singasinga, and (on Vanua Levu) wakathere. The type collection
is Seemann 196, of which the precise locality is in doubt; in the original descrip-
tion Seemann reports it as Kandavu, but one sheet at Kew with this number is
indicated as being from Port Kinnaird (Ovalau) in part and from Namosi
(Viti Levu) in part.

FIJI: Virr Levu: Milne 69 (K); Mba: Mt. Evans Range, Greenwood 863A
(A, Bish, US), 1220 (US); summit of Mt. Koroyanitu, Mt. Evans Range,

Natondra, Smith 4371 (A, US); Mt. Nairosa, Mt. Evans Range, Smith 4409
(A, US); Nandarivatu and vicinity, Gibbs 673 (BM), Parks 20676 (Bish),
20725 (Bish type of S. parksii), Gillespie 4021 (Bish, GH, K, NY, US), Tothill
777 (K), 778 (K), 778a (K), Sykes 27 (A), Greenwood 863 (A, K, NY, US),
Smith 4904 (A, US), Vaughan 3260 (BM); western slopes of Mt. Nanggaranam-
buluta, Smith 4792 (A, US); valley of Nggaliwana Creek, Smith 5371 (A, US);
western and southern slopes of Mt. Tomanivi, Smith 5222 (A, US); Ra

Numbumakita (about 10 miles east of Mt. Tomanivi), Gibbs 880 (BM);
Nandronga & Navosa: Northern portion of Rairaimatuku Plateau,

146 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxxu1

between Nandrau and Nanga, Smith 5429 (A, US); Namosi: Near Namosi,
Gillespie 2589 (Bish, GH, NY); Naitasiri: Nakatia, Navuakethe District,
B. E. Parham 2738 (A). Ovatau: Milne 52 (K), 267 (K). VAnua Levu:
Thakaundrove-Mathuata boundary: Crest of Korotini Range, Smith
553 (Bish, GH, K, NY, US); Thakaundrove: Natewa Peninsula, hills
south of Natewa, Smith 1967 (Bish, GH, K, NY, US). Fiji, without definite
locality: Seemann 196 (Kandavu, Ovalau, or Viti Levu?) (GH, K type),
Horne 845 (GH, K), 846 (GH, K), 1007 (GH, K).

The very close velutinous-puberulent indument of vegetative parts read-
ily distinguishes S. katakata, from the essentially glabrous S. graeffei and
S. serratum on the one hand, and from the hispidulous- or strigose-
pubescent S. samoense on the other. Gillespie noted the close relationship
of his S. parksit with S. katakata, suggesting as differentiating characters
only the thicker, smaller, and more coriaceous leaves. Among the specimens
cited above are many which show, on a single plant, variations in these
characters covering the extremes of the two type specimens. As I cannot
find any consequential points of difference, either in foliage or inflorescence,
among the cited specimens, Gillespie’s binomial is reduced to synonymy.
The prominence and length of the hairs of the disk-lobes, or even their
presence or absence, are not correlated with other characters and appear
strictly individual in nature.

wn

Spiraeanthemum samoense A. Gray, Bot. U. S. Expl. Exped. 1:
667. pl. 83, A. 1854, in Ann, Sci. Nat. IV. Bot. 4: 177. 1855, in Proc.
Am. Acad. 3: 128. 1857; C. Muell. in Walp. Ann. Bot. Syst. 5: 23.
1858; Reinecke in Bot. Jahrb. 25: 633. 1898; Pampan. in Ann. di
Bot. 2: 51. 1905; Rechinger in Denkschr. Akad. Wiss. Wien 85: 286.
1910; Setchell in Carnegie Inst. Publ. 341: 92. 1924; Christophersen
in Bishop Mus. Bull. 128: 96. 1935.

Shrub or tree up to 8 m. high, dioecious (or possibly sometimes polygamo-
dioecious), the branchlets distally copiously hispidulous with spreading
pale brown hairs 0.3-1 mm. long, at length becoming subglabrate; stipules
ovate when young, rapidly enlarging, at length lanceolate-oblong, up to 35
mm. long and 10 mm. broad, obtuse, densely or sparsely sericeous or
hispidulous without, glabrous within, soon caducous, the scars elongate,
curved; leaves opposite, the petioles semiterete, 1-4 cm. long, copiously
hispidulous like branchlets, at length subglabrate; leaf-blades subcoriaceous
or chartaceous, elliptic- or ovate-oblong, 5-14 (—15) cm. long, 2.5—7
(—7.5) cm. broad, broadly obtuse to acute at base and shorf-decurrent on
the petiole, obtusely cuspidate or callose-apiculate at apex, conspicuously
serrulate at margin with 2—4 callose-glandular teeth per centimeter, sparsely
strigose-puberulent on both sides (hairs grayish, 0.2—0.7 mm. long), at
length glabrate except indument persisting on costa and secondaries be-
neath, usually with small axillary domatia on lower surface, the costa
plane above or elevated in a groove, prominent beneath, the secondary
nerves 6-13 per side, arcuate-spreading, usually plane above and strongly
elevated beneath, the veinlet-reticulation intricate, plane or immersed

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XII 147

above, prominulous beneath; inflorescence paniculate, solitary, axillary or
pseudoterminal, ample, many-flowered, 7-17 cm. long, 3-10 cm. broad,
the peduncle subterete or slightly flattened, 1.5—-6 cm. long, hispidulous-
puberulent with hairs 0.1—0.3 mm. long, the branches opposite, persistently
hispidulous (hairs 0.2-0.5 mm. long), the bracts lanceolate, often sub-
foliaceous, to 15 mm. long, strigose-puberulent on both sides, the ultimate
bracteoles about 0.5 mm. long; flowers solitary at apices of ultimate branch-
lets (these 0.7—2.5 mm. long), the pedicels (above articulation) 0.5—1.8
mm. long, strigose-puberulent, often glabrate; calyx 1.7—2.3 mm. long and
broad, proximally puberulent, usually glabrate, the lobes 4 or 5, rarely 6,
ovate-deltoid, 1-1.5 & 0.7—1.1 mm., acute; stamens 8 or 10 (or 12?), the
filaments glabrous, in ¢ flowers 1.5—3 mm. long, in 9 flowers 1-1.5 mm
long, the anthers broadly ellipsoid, in ¢ flowers 0.2-0.3 mm. long, in ?
flowers minute and probably sterile; disk-lobes in ¢ flowers 4—6, carnose,
loosely coherent or free, oblong, 0.6—1 mm. long, irregularly truncate at
apex, ventrally and apically sparsely hispidulous with hairs 0.2-0.3 mm.
long or glabrous; disk-lobes in ? flowers 8 or 10 (or 12?), similar but
slightly shorter, 0.4-0.7 mm. long; carpels in ? flowers 4 or 5 (or 6?),
ovoid, densely sericeous with stramineous hairs 0.2—0.4 mm. long, the style
0.5—0.7 mm. long, the ovules 2, collateral, pendulous from near middle;
calyx and stamens persistent in fruit; mature carpels narrowly oblong-
ellipsoid, up to 3 mm. long and 1 mm. broad, persistently sericeous, the
style persistent; seeds 2 (1 often smaller or perhaps aborted), 1.5—2 mm.
long, the nucellus ellipsoid, 0.8—-1 mm. long, the wings subequal in length
(0.3-0.5 mm. long), the distal wing oblong, obtuse, the basal wing subulate.

DISTRIBUTION: Endemic to Samoa, where it occurs on at least the larger
islands at altitudes of 500-1700 m., being indicated by Christophersen as common
at middle and high elevations. It is usually noted as a tree 2-8 m. high,
occurring in various types of forest; the flowers are said to be white or yellowish
white and fragrant. Recorded local names are tauli, maota mea, saitamu, and
lau matui, The type is an Exploring Expedition collection, cited below, which
Gray cited with a question as being from Tutuila.

SAMOA: Savatir: Above Letui, Christophersen 780 (Bish, K); above Safune-
Letui, Christophersen 819 (A, Bish, US); above Matavanu, ase cd &
Hume 1992 (A, Bish, US), 2069 (Bish, K, NY, US), 2131 (Bish, US). Upotu:
Laulii River basin, Reinecke 281 (US); above Utumapu, ee 1518 (BM,
K, US); Lanutoo, Funk 113 (BM), Rechinger 706 (BM), 1920 (BM, K, US);
Malololelei- Lanutoo trail, Christophersen 373 (Bish, K, NY), 399 (Bish, US).
TuTuILa: Matafao Ride Collarino (in Setchell) 549 (Bish), Christophersen
1048 (Bish, NY), 1067 (A, Bish, US); Le Pioa, Christophersen 3506 (Bish, K,
NY), 3576 (Bish). Samoa, without definite locality: U. S. Expl. Exped.
(probably from Tutuila) (GH, US 47619 and 47620 type), Whitmee (BM,
GH), 37 (K), 257 (BM), 957 (K), Powell (GH, K, NY), 121 (K).

The single representative of the genus from Samoa is well characterized
by the comparatively long indument of its branchlets, lower surface of leaf-
costa, etc., by its elongate stipules which are glabrous within, by its ser-
rulate leaf- blades, and by the comparatively elongate ultimate inflorescence-

148 JOURNAL OF THE ARNOLD ARBORETUM _[VoL. xxx

branchlets. In this, as in other species of the genus, the pedicels might be
described as ‘jointed,’ but only the portion above the articulation is
strictly pedicellary.

4. PuLLea Schlechter

The genus Pullea, described in 1914 (in Bot. Jahrb. 52: 164. fig. 9), is
now composed of six species, all limited to New Guinea, as discussed by
Perry (in Jour. Arnold Arb. 30: 163-165. 1949). The species here
described as new indicates the occurrence as far east as Fiji of another
Papuasian genus. That so many genera with this distribution have not yet
been reported from the Solomons and the New Hebrides can only indicate
the sparsely collected nature of those archipelagos.

Pullea perryana sp. nov.

Arbor ad 6 m. alta, ramulis teretibus vel superne leviter complanatis,
glabris vel apices versus obscure strigillosis, inconspicue lenticellatis,
nigrescentibus demum cinerascentibus; stipulis subcoriaceis obovato-
suborbicularibus vel obovato-ellipticis, (8—) 10-12 mm. longis, (3—) 8-11
mm. latis, basi angustatis, apice rotundatis, margine valde revolutis,
utrinque obscure strigoso-puberulis glabratis, mox caducis, cicatricibus
brevibus transverse. ellipticis subrectis; foliis oppositis, petiolis crassis
semiteretibus vel leviter canaliculatis superne alatis (7—) 10-25 mm.
longis parce strigoso-puberulis mox glabratis, laminis coriaceis in sicco
brunnescentibus ellipticis vel lanceolato-ellipticis, (7—) 11-18 cm. longis,
(3.5—) 4-9 cm. latis, basi attenuatis et in petiolum longe decurrentibus,
apice obtuse cuspidatis (acumine ipso ad 1 cm. longo rotundato), margine
grosse undulato-crenatis (dentibus circiter 1 per centimetrum) et anguste
revolutis, utrinque glabris, costa valida supra plana vel leviter elevata
subtus prominente, nervis secundariis utrinsecus 7-11 arcuato-adscendenti-
bus supra subplanis subtus valde elevatis, rete venularum intricato supra
subimmerso subtus prominulo vel plano; inflorescentiis axillaribus vel
pseudoterminalibus ample paniculatis multifloris, binis vel ternatis super-
positis, sub anthesi (3—) 6-11 cm. longis et (2—) 3-7 cm. latis, pedunculo
gracili subcomplanato (1.5—) 4-7 cm. longo parce hispidulo (pilis pal-
lidis 0.1-0.2 mm. longis) glabrato, ramulis primariis apice pedunculi 3—7
aggregatis, bracteis primariis subfoliaceis lineari-oblongis ad 9 & 2 mm.
puberulis mox caducis, ramulis ut pedunculo minute hispidulo-puberulis;
floribus hermaphroditis sessilibus confertis 1-bracteolatis, bracteolis obo-
vatis 1-1.2 mm. longis apice rotundatis extus parce strigillosis intus
glabris margine ciliolatis; calyce 2—2.5 mm. longo demum subrotato ad
4 mm. diametro, tubo minute obconico, limbo papyraceo profunde lobato,
lobis 5 vel 6 anguste imbricatis oblongo-ellipticis 1.5-1.7 1-1.2 mm.,
inconspicue nervatis, extus parce strigilloso-puberulis vel glabratis, intus
dense puberulis, margine ciliolatis, apice rotundatis; petalis nullis; stamini-
bus 10 vel 12 demum deciduis, filamentis gracilibus 2—2.3 mm. longis,
antheris ellipsoideis circiter 0.4 mm. longis utroque rotundatis; disci lobis

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XII 149

10 vel 12 carnosis plerumque binatim cohaerentibus oblongo-obovoideis
subquadratis 0.3-0.4 mm. longis glabris, apice complanatis vel rotundatis;
ovario subsupero, basi in calycis tubo leviter immerso, crispato-tomentello
(pilis 0.5-0.7 mm. longis), stylis subulatis curvatis glabris 1.5-2 mm
longis, ovulis in quoque loculo 4 biseriatim pendulis.

DIsTRIBUTION: Fiji, known from only two collections, both obtained in
southeastern Viti Levu; Parham notes his collection as a tree 6 m. high, growing
in ridge forest, with cream-white flowers.

FIJI: Virr Levu: Naitasiri: Tholo-i-Suva, B. E. Parham 1646 (A);
Rewa or Naitasiri: ‘Central Road, Suva,” November, 1928, B. H. Tothill
472 (Bish, K type, US).

This remarkable species, so completely unlike anything yet described
from Fiji, is most suggestive of the New Guinean P. decipiens Perry, from
which it differs in its much larger leaves and inflorescences and in having
the ovary nearly completely superior, immersed in the calyx-tube only at
its base. The New Guinean species seem to have the ovary about half-
inferior, but otherwise no basic characters are discerned for segregation
of the Fijian plant.

It is a pleasure to name this species for Dr. Lily M. Perry, in recognition
of her valuable work on the flora of Papuasia, and with particular reference
to her review of the Cunoniaceae (in Jour. Arnold Arb. 30: 139-165.
1949). As Dr. Perry first suggested that the new species might be sought
in Pullea, the epithet seems particularly fitting.

DEPARTMENT OF BOTANY,

U. S. NATIONAL MusEuM,
SMITHSONIAN INSTITUTION.

150 JOURNAL OF THE ARNOLD ARBORETUM _[vov. xxxur

NOTES ON XANTHOSTEMON F. MUELLER AND
KJELLBERGIODENDRON BURRET

E. D. MerrrILu

GUGERLI’S * MONOGRAPH OF THE GENUS Xanthostemon F. Mueller was
published in Germany in 1940 and did not become available to us
until about a decade later. My interest in this work is not so much in
the New Caledonian species considered, which form the bulk of the
described forms, and the few Australian ones, but rather in his treat-
ment of certain Philippine, eastern Malaysian, and Papuasian species.
He recognized forty-three species and a few subspecies and varieties,
distributed into five newly proposed sections, Vesicaria, Brevistyla,
Cylindrica, Bullata, and Campanulata, the latter subdivided into two
subsections, Multiflora and Pauciflora. I do not criticize these minor
categories. Like other revisions of its type it has the merit of a
proposed system of classification, and of bringing together the widely
scattered published data regarding all the species described up to 1940.
One judges that perhaps certain obvious errors in nomenclature may
be due perhaps more to a lack of critical editorial work on the manu-
script than as wholly chargeable to a beginner who was working with
a peculiarly difficult group of plants. In nomenclature the author was
apparently misled by Pampanini’s erroneous interpretation of the rules
of nomenclature governing the validity of certain published binomials
in 1905. The several cases are discussed under Xanthostemon speciosum
Merr., X. pubescens C. T. White, XY. multiflorum (Montr.) Beauvisage,
and X. gugerli: Merr.

Xanthostemon is a genus of considerable significance from the stand-
point of phytogeography. Its great center of diversification is New
Caledonia, with a total of about thirty-three endemic species. Five
species are recorded from northern and northeastern Australia, three
from New Guinea, four from the Philippines, and one from Celebes
(this Celebesian form also extending to Moena, Ternate and Batjan).
To date no representative of the genus has been reported from any part
of the Sunda or Lesser Sunda Islands, the latter group extending east-
ward from Java. Incidentally Gugerli’s distribution map extends the
range of the genus, within the Philippines, to northern Luzon, about
250 miles beyond the known actual range of the Philippine representa-
tives.

One problem, not solved by Dr. Gugerli, as he did not have access
to certain historical material, was the status of Xanthostemon celebi-
cum Koord. He merely quoted the original distinctly unsatisfactory

*Gugerli, K. Monographie der Myrtaceengattung Xanthostemon. Repert. Sp.
Nov. Beih. 10: 1-49. pl. 1-16. 1940. Reprinted without change in pagination as a
doctorate thesis, University of Zurich, 1940.

1952] MERRILL, NOTES ON XANTHOSTEMON eo |

description and left the species as one of the two unknown to him.
The large fruits, described as 1.8 cm. long and 1.2 cm. in diameter,
might lead one to assume that some genus other than Xanthostemon
was represented, and this proves to be the case. Koorder’s unpublished
dissection notes and sketches clearly show that he knew the fruit to
be indehiscent and 1-seeded, and that the stamens were arranged in
five distinct phalanges; these are not Xanthostemon characters. It is
unfortunate that he did not include these data in his published descrip-
tion. The species proves to be a representative of the very different
Kjellbergiodendron Burret. When I initiated this study I had no
intention of considering Burret’s genus until an examination of material
now available indicated that a representative of this genus was involved.
Dr. van Steenis informs me that Beccari had recognized, named, and
described this striking genus on the basis of his own Celebesian col-
lections at some time previous to 1890. Unfortunately he never pub-
lished his description. Had he done so his name would have antedated
that of Burret by four or five decades and doubtless would have obvi-
ated the later Koodrders errors.

In the course of this study I have been impressed with the excellent
representation of the known species to be found in the herbarium of
the Arnold Arboretum. At least two-thirds of the described species
are represented by from one to many specimens, and it is of interest
to note that many isotypes are to be found here. Most of this material
has been acquired within the last fifteen years. I have accounted for
both of the described species which Dr. Gugerli could not place, X.
celebicum Koord. being transferred to Kjellbergiodendron, and X.
papuanum Lauterb. being reduced to XY. novaguineense Valeton. I add
X. crenatus C. T. White, of New Guinea, described in 1942, and here
describe as new Xanthostemon brassii Merr. from New Guinea and
X. confertiflorum Merr. from Celebes. Certain adjustments in nomen-
clature are also involved for four previously described species, where
Dr. Gugerli erred in selecting wrong specific names.

I am indebted to the officials of the Rijksherbarium, Leiden (L), the
United States National Herbarium (U), and the Gray Herbarium (G),
for the courteous loan of needed material. Except for collections indi-
cated by the above symbolic letters all other material studied is in the
Arnold Arboretum Herbarium (A), although before drafting this paper,
and after its completion, I had seen the material at Kew and the
British Museum.

Xanthostemon F. Mueller
NEW CALEDONIA

Xanthostemon gugerlii nom. nov.

Xanthostemon speciosum (Brongn. & Gris) Pamp. Nuovo Giorn. Bot. Ital.
II. 12: 688. 1905, in obs.; Gugerli, Repert. Sp. Nov. Beih. 120: 97. 1940;
Guillaumin, Fl. Nuov. -Caléd. 233. 1948, non Merr. 1904.

152 JOURNAL OF THE ARNOLD ARBORETUM _[VvoL. xxximI

Fremya speciosa Brongn. & Gris, Bull. Soc. Bot. France 12: 299. 1865.

Gugerli in accepting the validity of this New Caledonian species in
1940 adopted the name Xanthostemon speciosum (Brongn. & Gris)
Niedenzu, but I can find no record of an actual transfer of the specific
name to Xanthostemon antedating Pampanini’s overlooked one of 1905.
Even this, when published, was an unnecessary binomial as it was
antedated by X. speciosum Merr. (1904), which belongs to a very
different Philippine species. The only reference given by Gugerli is to
Zahlbruckner, Ann. Naturhist. Hofmus. Wien 3: 279. 1888, but there
only the binomial Fremya speciosa Brongn. & Gris was used, as is the
case in all other references I have found and checked. Of course Fremya
speciosa Brongn. & Gris (1865) does not invalidate Xanthostemon
speciosum Merr. (1904). The series of curious errors resulting from
Pampanini’s misinterpretation of the rules of nomenclature governing
the validity of binomials, and perpetuated and expanded by Gugerli
in 1940, is further discussed under Xanthostemon speciosum Merr.,
q.v. There is a duplicate of the type collection of this New Caledonian
species, Vieillard 2579, in the Gray Herbarium.

Xanthostemon multiflorum (Montr.) Beauvisage, Ann. Soc. Bot. Lyon
26: 46. 1901; Pamp. Nuovo Giorn. Bot. Ital. II. 12: 673. 1905;
Guillaumin, Bull. Soe. Bot. France 81: 14. 1934.

Draparnaudia multiflora Montr. Mém. Acad. Lyon, 10: 205. 1860, sphalm.

“Drapernandia.”
Fremya pubescens Brongn. & Gris, Bull. Soc. Bot. France 10: 373. 1863.
Xanthostemon pubescens Pampaloni, Nuovo Giorn. Bot. Ital. II. 13: 128.
1906; Gugerl, Repert. Sp. Nov. Beih. 120: 126. 1940; Guillaumin, FI.
Nouv.-Caléd. 234. 1948, non C. T. White (1917).

Gugerli states that Draparnaudia multiflora was, in part, Xanthoste-
mon flavum (Panch.) Schltr. However, Beauvisage clearly states
that there was but one poor specimen in the Lyon herbarium named
by Montrouzier as Draparnaudia multiflora Montr.; the type collection
hence could not have been a mixture. But when Beauvisage drew up
a complete description of Xanthostemon multiflorum (Montr.) Beau-
visage, on the basis of about 20 individual collections, he cited about
seven synonyms including not only Montrouzier’s original Drapar-
naudia, but also Fremya flava Brongn. & Gris, F. deplanchei Brongn.
& Gris, F. pubescens Brongn. & Gris, and F. elegans Brongn. & Gris,
all published in 1863. Incidentally Montrouzier in 1860 did not prepare
and publish an actual species description, other than as such data were
included in his generic description, and as he had only one poor speci-
men there could have been no mixture in his original species concept.
But Beauvisage’s description of 1901 is definitely of a collective species.
Here, then, must be the basis of Gugerli’s statement that Montrouzier’s
species was, in part, Xanthostemon flavum (Panch.) Schltr. Yet Pam-
panini, op. cit. 675, definitely stated that he had seen the Montrouzier
type, and on p. 682 repeated the statement. He concluded that this

1952] MERRILL, NOTES ON XANTHOSTEMON 153

specimen, admittedly a poor one, was the same as Fremya pubescens
Brongn. & Gris and cited Montrouzier’s binomial in the synonymy of
Xanthostemon multiflorum (Montr.) Beauvisage, making it X. multi-
florum (Montr). Beauvisage, var. typicum Pamp., forma pubescens
(Brongn. & Gris) Pamp. Under the circumstances I do not hesitate
to replace Gugerli’s invalid binomial Xanthostemon pubescens (Brongn.
& Gris) Gugerli by the earlier name of Montrouzier. Until by a reex-
amination of the Montrouzier type it can be proved that Pampanini
erred, there seems to be no other choice, unless one wishes to propose
a new specific name which I consider to be uncalled for at present.
The species is known only from New Caledonia.

Xanthostemon myrtifolium (Brongn. & Gris) Pamp. Nuovo Giorn.
Bot. Ital. II. 12: 682. 1905; Gugerli, Repert. Sp. Nov. Beih. 120:
68. 1940.
Fremya i Migr Brongn. & Gris, Bull. Soc. Bot. France 12: 299. 1865,
Ann. Sci. Nat. V. Bot. 3: 227. 1865.
Perrier % integrifokum Baker f. Jour. Linn. Soc. Bot. 45: 311. 1921;
Gugerli, op. cit. 58; Guillaumin, Fl. Nouv.-Caléd. 232. 1948, syn. nov.
Fremya integrifolia Brongn. & Gris ex Baker, f. l.c. in syn., syn. nov.
Gugerli noted, which Baker f. did not, that Fremya integrifolia
Brongn. & Gris was an unpublished herbarium name. But Baker f. in
transferring it to Xanthostemon in 1921 failed to provide a description,
although he did publish Compton’s field note to the effect that it was a
shrub with small hard leaves with thickened margins, white corollas
and pale yellow stamens. This Gugerli accepted as a description and
with misguided confidence placed the species in his section Brevistyla
where it does not belong. His judgment must have been based solely
on the statement that the flowers were white.
The actual specimen on which the Brongniart and Gris and the Baker
f. binomials were based is in the British Museum herbarium, and
Baker f. was correct in referring to it Compton 826, but erred, as did
Gugerli, in so identifying Compton 375 (cited by Gugerli as Baker
375). The latter is a small-leaved form of Metrosideros operculata
Labill. Guillaumin Not. Syst. 1: 109. fig. 5, 1909, concluded that the
proposed varieties of this species should be abandoned because of the
intergrading forms. The British Museum specimen of Fremya integri-
folia carried the collector’s ample note, but no collector’s name or num-
ber. I showed this to Dr. Tardieu-Blot, who was at the British Museum
for a few days in July, 1951, and on her return to Paris she completed
the record. The same field note appears on a New Caledonia collection,
Baudouin 638; this Paris specimen is identical with the London one,
and both match the type of X. myrtifolium (Brongn. & Gris) Pamp.
Thus another minor mystery is solved, for in 1934 (Bull. Soc. Bot.
France 81: 14) Guillaumin had stated that Xanthostemon integrifolium
Bak. f. nomen (Fremya integrifolium Brongn. et Gris) was totally un-
known to him and that it was not represented in the Paris herbarium.

154 JOURNAL OF THE ARNOLD ARBORETUM _[vot. xxxim

He accepted the species in 1948 solely on the authority of Gugerli.
But neither Gugerli, who did not see the British Museum specimen, nor
Guillaumin, had any reason to believe that Baudouin 638, with which
both were familiar, was an isotype of the elusive Yanthostemon integ-
rifolium (Brongn. & Gris) Baker f. which now proves to be the case.

AUSTRALIA

Xanthostemon hues Gugerli, Repert. Sp. Nov. Beih. 120: 83. 1940,
sphalm. whiti

Xanthostemon hi C. T. White, Proc. Roy. Soe. caren 28: 57. 1917;

Queensl. Dept. Agr. Bull. 20: 14. 1918, non Pampaloni, 1906.

The type of this species was from the Atherton and Herberton dis-
tricts, Queensland, Australia. It is well represented by Kajewski 1046
(A), May 24, 1929, from Gadgarra, Atherton, not far from the type
locality. At first sight one infers that a new name was not needed here.
At any rate Gugerli’s reason for publishing the new name was invalid
as Fremya pubescens Brongn. & Gris (1863) did not invalidate Xantho-
stemon pubescens C. T. White (1917) ; and yet it develops that the new
name was needed because of the earlier and still unlisted Xanthostemon
pubescens Pampaloni (1906) which all authors have overlooked.
Gugerli’s new specific name was misspelled, he should not have cited
C. T. White as the parenthetic author, and the one collection cited by
him is not C. T. White 1046, but is S. F. Kajewski 1046; the identifica-
tion was by C. T. White.

NEW GUINEA
Xanthostemon brassii sp. nov. Sect. Campanulata, Multiflora.
Xanthostemon paradorum sensu C. T. White, Jour. Arnold Arb. 23: 83.
1942, non F. Muell.

Arbor usque ad 30 m. alta, decidua, inflorescentiis leviter et brevis-
sime adpresso-pubescentibus exceptis glabra, ramulis ultimis rugosis,
4-5 mm. diametro, cicatricibus distinctis ornatis; foliis alternis, sub-
confertis, coriaceis vel junioribus subchartaceis, ellipticis vel oblongo-
ellipticis, sicco brunneis vel pallide olivaceis, 6-15 cm. longis, 3-7 cm.
latis, apice plerumque late rotundatis, rariter obscure retusis, junioribus
distincte sed adultis obscure glanduloso-punctatis; nervis primariis
utrinque circiter 15, irregulariter dispositis, patulo-curvatis, utrinque
distinctis, leviter elevatis, arcuato-anastomosantibus sed venam intra-
marginalem vix formantibus; petiolo 6-14 mm. longo; inflorescentiis
pseudoterminealibus, singulis in axillis foliorum vel delapsorum disposi-
tis, totis ad 8 em. diametro, leviter adpresse breviter pubescentibus,
sub fructu glaberrimis, singulis 3-4 em. longis, breviter (ca. 1 em.)
pedunculatis, 3-5-floris; floribus 5-meris, flavidis, breviter (5-8 mm.)
pedicellatis, bracteolis haud visis, ut videtur cite deciduis; calycibus
extus leviter pubescentibus, tubo infundibuliforme, circiter 4 mm.
longo, 5-6 mm. diametro, intus glabro, lobis orbiculari-ovatis vel

1952] MERRILL, NOTES ON XANTHOSTEMON 155

reniformi-ovatis, basi 2-3 mm. latis, sursum vix angustatis, 1.5-2.5 mm.
longis, apice late rotundatis; petalis ellipticis, late rotundatis, 4 mm.
longis et 3 mm. latis, in partibus medianis obscure glandulosis; stami-
nibus circiter 20, 1-seriatis, filamentis liberis, 1.5-2 cm. longis; antheris
ellipsoideis, obtusis, 1.8 mm. longis; ovario glabro 3-loculare, subhem-
isphaerico; stylo ad 2.5 cm. longo; capsulis globosis, 1 cm. diametro,
3-locularibus, seminibus numerosis, compressis, ambitu subtriangularis,
3-4 mm. longis latisque.

BRITISH NEW GUINEA: type Brass 7869 (flowers), 7503 (fruits) taken
from the same tree, Lake Daviumbu, Middle Fly River, the flowers Sep-
tember 1, the fruiting specimen August 26 “large tree, 30 m., briefly deciduous,
a crop of flowers appearing a few days before the fall of the leaves, flowers
yellow.” Other specimens are Brass 6556, 5932 (both in fruit), common in the
savannah forest at Dagwa. Oriomo River, and Mabaduan, Western Division,
and Brass 8575, Tatara, Wassi Kussi River, abundant on savannah forest
ridges, entering the rain forest. The full notes are given by C. T. White, l.c.

C. T. White after examining F. Mueller’s apparently not very satis-
factory type, from Arnhem Land [Northern Territory], Australia,
considered that all of these Papuan collections represented Xantho-
stemon paradorum F. Muell., sensu lat. While I have available only
two good specimens representing the Australian species, and one of
these (herb. Gray) is an isotype, there are so many differences that I
feel justified in describing the New Guinea form as a distinct species.
Gugerli, Repert. Sp. Nov. Beih. 120: 81. 1940, provided some additional
descriptive data for the Australian form, citing about nine individual
collections all from the Northern Territory of Australia. These notes
are of such a character that they support my opinion that had he had
access to the Brass collections at the time he studied the group, he
would have recognized this New Guinea form as a distinct species.
As White noted, F. Mueller had two individual collections, these not
quite identical, and he based his description on the characters of both.
In his original description of 1857 Mueller stated: “In collibus petraeis
ad flumina Victoria et Fitzmaurice,” these two rivers in the western
part of what is now the Northern Territory of Australia. In his ampli-
fied description of 1858 Mueller cited only the Victoria River locality.
It is this Victoria River collection, represented at Kew and at the Gray
Herbarium, that I accept as the type. It has distinctly pubescent
leaves, and densely cinereous-pubescent inflorescences, including the
outside of the calyces. In this the densely pubescent bracteoles are
persistent or at least subpersistent. There is in the U. S. National
Herbarium another specimen of this pubescent form merely labelled
“Schomburgk, North Coast”; this is undoubtedly the Port Darwin
collection distributed by Schomburgk, Schulz 356 as cited by Gugerli.
In addition to these two specimens which have been available to me
for comparison, I made notes on the Kew collections of Mueller,
Cunningham, Basedow, Spencer, and Stokes, these also seen by Dr.
Gugerli.

156 JOURNAL OF THE ARNOLD ARBORETUM _[VvoL. xxx

The individual 3- to 5-flowered inflorescences in Xanthostemon
brassii Merr. are associated with mature leaves or often with very
young leaves, or occur in the axils of fallen leaves. Taken together
they give the impression of a terminal many flowered panicle up to
8 cm. long and wide. These individual inflorescences are associated
with the deciduous character of the tree, the flowers apparently com-
mencing to develop with the fall of the old leaves and the almost
simultaneous appearance of the new foliage.

Xanthostemon crenulatum C. T. White, Jour. Arnold Arb. 23: 82.
1942. Sect. Campanulata, subsect. Multiflora.

This addition to the species considered by Dr. Gugerli in 1940 was
based on Brass 5805, 8358, 8473, 8602, all in the Arnold Arboretum
herbarium, collected from various parts of British New Guinea in 1934
and in 1936-37. Aside from its almost strictly opposite leaves (described
as subopposite), which is an anomalous character in Xanthostemon,
another striking feature is its unusually small flowers. These were
described by Mr. Brass as white and as greenish white. The bracts
and bracteoles, not described in the original description, are present
but are deciduous, being present in inflorescences with young buds,
falling as the flowers open. They are linear-lanceolate, pubescent, and
up to 5 mm. long.

This species may better be placed in Nani Adanson (Nania Miquel)
because of its opposite leaves. Adanson’s genus, by common consent,
is placed as a synonym of Metrosideros Banks. Valeton, however, Ic.
Bogor. 1: 63, 67, pl. 98, 99. 1901, accepted Nania Miq. as generically
distinct from Metrosideros Banks, recognizing two species, Nania vera
Miq., and Nania petiolata Valeton, calling attention to the fact that the
fruits of Nania Mig. (= Nani Adanson) are entirely free from the
calyx (superior), and that the placentas and seeds are quite different
from those of Metrosideros Banks; I may add that the valves split to
the very base, while in Metrosideros the fruits are inferior and open by
radiately arranged valves across the truncate tops. I had, at first, in-
cluded certain of these opposite-leaved species in Xanthostemon (New
Guinea and Amboina), which I have eliminated, since I am now con-
vinced that they do not belong in the latter genus, but really represent
species of Nani Adanson. Involved here are Metrosideros vera Lindl.
(1821-24; Roxb. 1832) from Amboina; M. suberosa Roxb. (1814,
1832), Moluccas, Syncarpia vertholenii Teysm. & Binn. (1855) =
Metrosideros vera Lindl., and Nania petiolata Valeton (1900) (prob-
ably from Celebes, not Java). I have not had access to sufficient mate-
rial to settle the various matters involved, both as to generic and as to
specific limits among the taxa above listed.

Xanthostemon novaguineense Valeton, Bull. Dép. Agr. Ind. Neérl.
10: [72]. 1907, Ic. Bogor. 3: sub. pl. 239. 1907; Gugerli, Repert.
Sp. Nov. 120: 85. 1940.

1952] MERRILL, NOTES ON XANTHOSTEMON 157

Xanthostemon paradorum sensu Valeton, Ic. Bogor. 3: 95. pl 239. 1907,
non F. Muell.

Xanthostemon papuanum Lauterb. Nova Guinea 8: 854. 1910; Gugerli, op.
cit. 130, inter sp. dub.; C. T. White, Jour. Arnold Arb. 23: 82. 1942, syn.
nov.

All the collections involved in this case were from the Humboldt Bay
region, north coast of New Guinea. Valeton’s type was from Tobadi,
a village on the inner bay, and Lauterbach’s type was from the lower
slopes of the neighboring Cyclops Mountains. Hollandia is the impor-
tant town here, and the other localities mentioned are near that place.
Sigafoos 42 from near Lake Sentano, exactly matches a duplicate of
Wichtman 125 in the Rijksherbarium, the type collection of Valeton’s
species and also agrees with the excellent illustration and detailed
description of X. novaguineense Val., while Brass 8801 from Hollandia
agrees perfectly with Lauterbach’s description of X. papwanum Lauterb.
and with Gyellerup 488, in the Rijksherbarium, the latter being the
Lauterbach type collection. The only differences I have been able to
detect after a searching comparison are that in the taxon of Valeton
the inflorescences are glabrous, and in that of Lauterbach they are
somewhat pubescent. The Sigafoos note reads, in part, “shrub in the
grass savannah on laterite, common 250 to 400 ft., flowers brilliant red.
A similar plant observed later was a tree 40 to 60 feet high.” The
Brass note is ‘“common tree 15 to 17 m. in old seral rainforest, abundant
as a small tree or shrub on dry forested slopes covered with grass and
ferns, flowers red, alt. 20-100 m.” It should be noted that the Lauter-
bach type from the neighboring Cyclops Mountains at 400 m. was
from alang covered slopes, alang being the coarse grass Imperata.

Here Dr. Gugerli cleared up the nomenclatural difficulties appertain-
ing to Valeton’s erroneous concept of Xanthostemon paradoxum F.
Muell. After Valeton’s detailed description and illustration was in
press he apparently saw authentic material of F. Mueller’s Australian
species, and published his new binomial first in his corrections to his
list of Papuan plants in an unnumbered sheet of the Bulletin in Buiten-
zorg, and a little later in a supplementary unpaged sheet in the Icones
which was printed in Holland. Unfortunately Dr. Gugerli did not see
Lauterbach’s type, and being unable to place the species in his arrange-
ment of them, left it among the few of doubtful status. It is, however,
rather strange that he should have expressed the opinion that a species
of the Australian genus Kunzea Reichb. might be represented. The
description is all of Xanthostemon, not at all of Kunzea, and apparently
no Kunzea has as yet been found in New Guinea. My conclusion is
that the very slight differences between the two supposedly distinct
species, 1.e., glabrous as opposed to somewhat pubescent inflorescences,
are due to local conditions as to exposure, etc., and that but a single
valid species is here represented.

158 JOURNAL OF THE ARNOLD ARBORETUM _ [VOoL. XxXxIII
CELEBES

Xanthostemon confertiflorum sp. nov. Sect. Vesicaria.

Ut videtur arbor vel arbor parva omnino glabra, ramulis ultimis
1.2-2 mm. diametro; foliis numerosis, plus minusve confertis, coriaceis,
vix vel obscure puncticulatis, plerumque obovatis interdum subellipticis
vel subelliptico-obovatis, apice late rotundatis vel rariter subretusis,
basi late acutis, breviter (5-8 mm.) petiolatis, sicco subolivaceo-brun-
neis vel pallide brunneis, opacis vel subnitidis, 4-8 em. longis, 3-5 cm.
latis; nervis primariis utrinque 10-12, gracilibus, vix vel obscure
elevatis, haud perspicuis, subtus dense reticulatis; inflorescentiis termi-
nalibus, sessilibus, circiter 3 cm. diametro, floribus confertis, 5-meris,
breviter (ad 3 mm.) crasseque pedicellatis; bracteolis binis, anguste
oblongis, acutis, coriaceis, glabris, circiter 5 mm. longis et 1.2 mm.
latis; calycibus, lobis inclusis, circiter 7 mm. longis, glabris, tubo
subpatelliformibus, ad 1 cm. diametro, lobis 5, triangulari-ovatis,
coriaceis, deorsum 3-4 mm. latis, sursum angustatis, 3 mm. longis,
acutis vel subacuminatis, sub fructu saepe recurvatis; petalis 5, orbicu-
laribus, late rotundatis, 6 mm. diametro; staminibus circiter 30, 1-seri-
atis, filamentis liberis, immaturis (inflexis) 6 mm., maturis rectis 1.5 cm.
longis; stylo ad 2.5 em. longo; ovario superiore vel semisuperiore,
glabro, depresso-globoso, 3-loculare, cellulis multiovulatis; capsulis
globosis vel subglobosis, 3-loculatis, punctato-glandulosis, circiter 1 cm.
diametro; seminibus numerosis, compressis, obovatis, circiter 3 mm.
longis.

CELEBES: Malili and vicinity, Neth. Ind. For. Serv. Cel. I1I-103 (A, L),
bb. 18018 (L), 18011 (A), 18672 (A), 21782 (A), 22723 (A); Manado,
bb. 19686 (A, L), 31512 (A).

This series of specimens was collected in 1933, 1934, 1935, and 1939.
Only bb. 19636 is sterile, the others having either flowers or just opening
flower buds, or mature fruits. The indicated type is the first cited
specimen, although its flowers are not quite mature. The capsule
characters were taken from bb. 18011, and bb. 31512. Notes regard-
ing the plant are lacking except that the altitude is indicated as from
25 to 500 m.; all but one of the specimens (and that a sterile one) were
apparently dried out from material originally preserved in alcohol,
thus all traces of the flower color are lacking, but the flowers were
probably purplish. It is the first true Xanthostemon to be discovered
in Celebes. I have placed it in the section Vesicaria because of its
shallow calyces, in spite of the fact that it lacks the five protuberances
on the calyx tube, which is one of the characters of that section. It
is distinguished from the Philippine M. speciosum Merr. (M. merrillii
Pamp., M. purpureum Gugerli), not only by lacking the calyx pro-
tuberances but also by its leaves being very obscurely or not at all
glandular-punctate.

1952] MERRILL, NOTES ON XANTHOSTEMON 159

PHILIPPINES
Xanthostemon speciosum Merr. Govt. Lab. Publ. 6: 10. 1904.
Xanthostemon merrillii Pamp. Nuovo Giorn. Bot. Ital. 12: 688. 1905.
Xanthostemon purpureum Gugerli, Repert. Sp. Nov. Beih. 120: 53. pl. 15.
g. a. 1940.

This species, Gugerli’s description of 1940 having been based on
Merrill 682 from Culion (the type), Weber 1551 from Busuanga, and
F. B. 29266 Cenabre from Palawan, is also represented by F. B.
28902 (A) from Culion, and Philip. Nat. Herb. 218 Edano (A), and
12431 Sulit (A) from Palawan, the first from near Puerto Princesa,
the second from the vicinity of Victoria Peak. I have examined speci-
mens of all the numbers cited by Gugerli.

This is the type of section Vesicaria Gugerli. The species is distinct
from X. verdugonianum Naves to which I erroneously reduced it in
1923 (Enum. Philip. Fl. Pl. 3: 183). Gugerli correctly reinstated it as
a species in 1940, even if he erred in redescribing it as new, for already
two other binomials had been published for it. In nomenclature and
in the recognition of species we here have a strange comedy of errors,
for Gugerli cites Merrill 682 from Culion, as the type of X. purpureum
Gugerli (1940). He said, op. cit. 131-132, that the Kew specimen of
this number actually represented X. verdugonianum Naves. I have
reéxamined it and find it, like all other specimens of this number, to
be X. speciosum Merr., and not the Naves species; Mr. H. K. Airy
Shaw later verified this at my request. This now historical Merrill 682,
all specimens taken by me personally from a single tree in the Cogonal
Grande, Culion, Feb. 12, 1902, is thus the basis of X. speciosum Merr.
(1904), X. merrillii Pamp. (1905), and X. purpureum Gugerli (1940).
There is no possibility of a mixture of material under the number dis-
cussed. The holotype was destroyed when the Manila herbarium was
burned near the close of World War II at the time of the reoccupation
of Manila by American troops. I have examined the duplicate types
at Kew, the Gray Herbarium, and the U. S. National Herbarium.

The errors commenced with Pampanini in 1905 who proposed the
unnecessary new binomial X. merrillii Pamp. because, while he correctly
accepted the reduction of Fremya Brongn. & Gris to Xanthostemon
F. Muell., he erroneously concluded that Fremya speciosa Brongn. &
Gris (1863) invalidated Xanthostemon speciosum Merr. (1904), which
is utterly contrary to the rules of botanical nomenclature. I continued
the error when, without checking the details, I accepted Pampanini’s
conclusion in 1923 and added to the confusion, thus probably aiding
Gugerli in some of his misinterpretations, as I then erroneously reduced
X. speciosum Merr. (X. merrillii Pamp.) to X. verdugonianum Naves.
The superficial resemblances of X. verdugonianum Naves and X.
speciosum Merr. (X. merrillii Pamp., X. purpureum Gugerli) are close,
but the calyx characters are very distinct. And finally Gugerli, in 1940,
misled by Pampanini’s misinterpretation of the rules governing the

160 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxxmI

validity of binomials redescribed this already twice named Culion
species as Xanthostemon purpureum Gugerli (p. 53), sect. Vesicaria,
cited its type collection as representing X. verdugomanum Naves
(p. 64), sect. Cylindrica, and finally (p. 131) listed and discussed X.
merrillii Pamp. (1905), with X. speccosum Merr. (1904) as a synonym,
as an excluded species. And all the time X. speciosum Merr. was the
valid name for this now thrice-named species. He even mentioned the
striking calyx characters specified by me in 1904 by which X. speciosum
Merr. was distinguished from X. verdugonianum Naves, which were
the very characters on which he based his section Vesicaria. And so
a page or two of print is now called for to explain the details of this
nomenclatural comedy of errors, in which Merrill, Pampanini, and
Gugerli are involved.

There is no overlapping 1 in the Philippine ranges of the two super-
ficially similar species, X. speciosum Merr. and X. verdugoniamum
Naves. The latter is confined to the central and southern Philippines,
the former to the Calamian-Palawan group in the central western part
of the Philippines. Incidentally these islands all lie on the continental
shelf, and their floras and faunas contain strong Bornean elements
which do not extend into the Philippines proper.

Xanthostemon verdugonianum Naves ex F. Vill. in Blanco FI. Filip.
ed. 3, Novis. App. 82. pl. 300. 1880; Merr. Enum. Philip. Fl. PI.
3: 183. 1923, excl. syn.; Gugerli, Repert. Sp. Nov. Beih. 120: 64.
1940, excl. syn.

To be excluded from Gugerli’s consideration of this endemic Philip-
pine species are the synonyms X. merrillii Pamp. and X. speciosum
Merr., and from the specimens cited, Merrill 682, these being accounted
for under X. speciosum Merr., above. Otherwise, all of the collections
cited by Gugerli manifestly belong with this species of Naves. Addi-
tional collections are F. B. 24422 Miras, Soriano & Mariano (A, U),
from Agusan Province, Mindanao, F. B. 7546 Hutchinson (U), F. B.
22833 Ponce (A,U), F. B. 23023 Razon (G), and 2302 (A), F. B.
28158 Tomeldan (A), and F. B. 29419 Rojas, all from Surigao Province,
Mindanao, F. B. 23942 Cortes & Knapp (U), Panay, and F. B. 19535
José from Sibuyan. The species, type from Surigao no longer extant,
is now known from more than 20 individual collections, its range being
Sibuyan, Panay, Leyte, Dinagat, Tinago, and the Provinces of Agusan
and Surigao in Mindanao.

UNLISTED BINOMIALS

In the course of this little study I have noted several unlisted
binomials, one dating from as early as 1886. These are:

Xanthostemon *myrtifolium [Brongn. & Gris] Pampaloni, Nuovo
Giorn. Bot. Ital. II. 18: 135. 1906 [Fremya myrtifolia Brongn. «&
Gris]; Gugerli, Repert. Sp. Nov. Beih. 120: 68. 1940.

1952] MERRILL, NOTES ON XANTHOSTEMON 161

Doctor Gugerli gave the reference to Pampanini’s paper in the Nuovo
Giorn. Bot. Ital. II. 12: 682. 1905, this being an error; the binomial
does not there appear, nor is it to be found elsewhere in Pampanini’s
paper of 1905. The next year it was published without its name-bring-
ing synonym, as above indicated. Yet in his monograph of 1940 Dr.
Gugerli also erroneously included a second reference to X. myrtifolium
Pamp. [ex Baker f.] in Jour. Linn. Soc. Bot. 45: 34. 1921. But there
Baker f. credited the binomial to Guillaumin. Guillaumin however,
apparently never published such a name, as a check on his various
papers on the New Caledonian flora shows that he correctly credited
the binomial to Pampanini. Pampaloni undoubtedly received his bi-
nomials from Pampanini.

Xanthostemon *pachyspermum F. Muell. & F. M. Bailey, Occ. Pap.
Queensl. Fl. 1: 4. 1886; F. M. Bailey, Queensl. Fl. 2: 642. 1900;
Gugerli, Repert. Sp. Nov. Beih. 120: 132. 1940, inter sp. excl. =
Tristania pachysperma (F. Muell. & F. M. Bailey) Francis,
Queensl. Nat. 14: 56. 1951 (7. odorata C. T. White, 1920).

Doctor Gugerli cited the authority for the binomial as F. M. Bailey,
and gave the reference to the Queensland Flora only; there, however,
the reference is to the earlier (but as yet unlisted) place of publication
as I have above recorded the entry. He was correct in excluding the
species from Xanthostemon, as it was described as having but one or
two large seeds in each cell of the capsule, these seeds 7 to 10 mm. in
diameter. He thought that Bailey might have had a specimen of

Kunzea, sect. Salicia. The mystery is now solved by W. D. Francis’

reexamination of the type in 1950, who finds it to represent the same

species as the Queensland Tristania odorata C. T. White

Xanthostemon *pubescens [Brongn. & Gris] Pampaloni, Nuovo
Giorn. Bot. Ital. II. 13: 128. 1906; Gugerli, Repert. Sp. Nov. Beih.
120: 126. 1940; Guillaumin, Fl. Nuov. Caléd. 234. 1948. New
Caledonia = X. multiflorum (Montr.) Beauvisage, supra.

Pampaloni cited no synonyms and no authorities; he received his
binomials from Pampanini, with whom he was working; but nowhere

did the latter author publish this binomial.

Xanthostemon *speciosum Pamp. Nuovo Giorn. Bot. Ital. II. 12: 688.
1905, in obs. (Fremya speciosa Brongn. & Gris). New Caledonia.

In Dr. Gugerli’s treatment, p. 97, he erroneously credited this bino-
mial to Niedenzu who never published it, overlooking Pampanini’s
obscure entry. The proper name for this New Caledonian species is

X. gugerlia Merr.

EXCLUDED SPECIES

XANTHOSTEMON CELEBICUM Koord. Meded. ’s Lands Plant. 19: 465,
637. 1898; Gugerli, Repert. Sp. Nov. Beih. 120: 130. 1940 =
Kjellbergiodendron celebicum (Koord.) Merr., infra.

162 JOURNAL OF THE ARNOLD ARBORETUM _[VoL. XxxIII

XANTHOSTEMON PACHYSPERMUM F. Muell. & F. M. Bailey, Occ. Pap.
Queensl. Fl. 1: 4. 1886; Gugerli, op. cit. 132 = Tristania pachy-
sperma (F. Muell. & F. M. Bailey) Francis, supra.

Kjellbergiodendron Burret

This genus was described in 1936, with two species, on the basis of
two collections made by Gunnar Kjellberg in Celebes in 1929. The
previously unplaced and inadequately described Xanthostemon celebi-
cum Koord. is now found to belong in Burret’s very distinct genus.
The genus is strongly characterized by its relatively large, more or
less fleshy, 1-celled and 1-seeded, indehiscent fruits, 2-celled ovaries,
stamens arranged in five phalanges, and its alternate leaves. While
the genus is a sharply defined one, this statement apparently does not
apply to its few species, as they impress me as being difficult to dis-
tinguish from each other. It is probable that this difficulty stems
largely from the inadequateness of the available herbarium specimens,
most of these being sterile, or with young flower buds or with very
immature fruits. Certain sterile specimens from Malili, Celebes, dis-
tributed as representing the undescribed T'ristania celebica Koord.,
belong with Kjellbergiodendron hylogeiton Burret. This Koorders bi-
nomial appears as a nomen nudum in Koorders-Schumacher, Syst.
Verzeich. 3: 96. 1914; I have seen three of the four Koorders specimens
listed under this Tristanza, all sterile.

Kjellbergiodendron celebicum (Koord.) comb. nov.

Xanthostemon celebicum Koord. Meded. ’s Lands Plant. 19: 465, 637.
1898; Koord.-Schum. Syst. Verzeich. 3: 96. 1914; Gugerli, Repert. Sp.
Nov. Beih. 120: 130. 1940, inter sp. ign.

In Koorders’ report on the Celebes flora he first listed this species
with brief comments on page 465, and on page 637 he published a short
and inconclusive description, some of the reasons for this perhaps to
be apparent in the following notes. It was not until 1914 that any of
the Koorders Celebes numbers were actually associated with the de-
scription (for Koorders cited no numbers and indicated no type in
1898) when Mrs. Koorders listed eleven numbers, all but four of which
represent sterile specimens. The description was of such a nature that
Dr. Gugerli in 1940, without access to authentically named specimens,
could not place the species in relation to the other described ones in
Xanthostemon.

Five of the Koorders Celebes numbers are sterile, so that in selecting
a type we are limited to four numbers only, 18097, 18544, with not
fully developed flower buds, and 18240, 19302, with fruits. All of these
have the smaller leaves, 10 to 15 em. long, 3.5 to 5.5 em. wide. I arbi-
trarily designate Koorders 18544, three sheets at Buitenzorg, as the
type for flowers, and Koorders 19302 in the same herbarium as the
type for fruits. I have actually seen Koorders 18097, 18964, 18322,

1952] MERRILL, NOTES ON XANTHOSTEMON 163

18960, 18321, all in the Leiden herbarium, but I deliberately ignore
the last three of these as they are sterile and with very large leaves,
up to 33 X 7 and 37 X 9.5 cm.; nos. 18192, 18305 also belong in this
category. Koorders thought that these large leaved forms were from
young plants; one cannot prove this short of intensive field work, but
they may, of course, belong with the species. Numbers 18097, 18240,
18544, 19302 are apparently normal, as these are the specimens with
flower buds or with fruits, the leaves being 10 to 15 cm. long and 3.5 to
5.5 em. wide. They clearly belong to a single species, and 18964,
(sterile) falls in this category.

It is rather curious that Koorders placed this species in Xantho-
stemon, because of its large fruits, which he mentioned as being 1.8 cm.
long and 1.2 em. thick (the largest at Buitenzorg is 1.9 cm. long), and
which he apparently knew, from his dissections and sketches, were
indehiscent and 1-seeded. In his dissection notes on the flower buds,
which he also did not publish, he observed that the stamens were ar-
ranged in five phalanges. These fruit and flower characters are remote
from those of Xanthostemon. A dissection of a flower bud of Koorders
18097 shows that the strictly inferior ovaries are 2-celled, each with
a fair number of ovules. All these characters are those of Kjellbergio-
dendron Burret which was not characterized and published until 1936.
I am indebted to Dr. J. H. Kern of Buitenzorg for copious notes on
the Koorders Buitenzorg specimens and his unpublished dissection data.

In addition to the several Koorders numbers that I am willing to
accept as representing Xanthostemon celebicum Koord. I feel safe in
referring to this species the following collections, unfortunately all
sterile:

CELEBES: Minahassa, Koorders 18097 (L), 18964 (L); Manado, Neth.
Ind. For. Serv. bb. 19646 (4,L); Palopo. bb. 20895 (A,L); Moena Island im-
mediately south of southwestern Celebes, Neth. Ind. For. Serv. bb. 21097
(A,L), 21886 (A), 4187 (L); Moluccas, Ternate and Batjan, Neth. Ind. For.
Serv. bb. 16476 (A,L) (the small islands of Ternate and Batjan are close to
the west coast of Halmahera, across the Molucca Passage from the classical
locality, Minahassa, northeastern Celebes). Because of the small flower buds,
Neth. Ind. For. Serv. Cel. IT. 485 (L) from Malili probably belongs here while
Neth. Ind. For. Serv. bb. 24124, 31513 (A) have puberulent calyces, the buds
somewhat intermediate in size between those of Koorders’ species and K.
limnogeiton Burret.

While it is clear that Xanthostemon celebicum Koord. is a Kjellber-
giodendron, it is not easy to determine its relationships with the two
previously described species of that genus. The almost mature but as
yet unopened flower buds are 6 mm. long, and under a lens the calyces
are densely puberulent. Hence the flowers should be distinctly smaller
than are those of Kjellbergiodendron limnogeiton Burret, which are de-
scribed as glabrous, while the fruits (mature?) are very much smaller
(1.8-1.9 em. * 7-9 mm.), as opposed to 5-6 cm. X 3.5-3.8 em. in
Burret’s species. One may judge by Koorders’ sketches and by the

164 JOURNAL OF THE ARNOLD ARBORETUM _[VvoL. xxxuI

thickness of cotyledons observed by him that his fruits were at least
partly mature. It is suspected that Koorders species is closest to K.
hylogeiton Burret, the fruits (mature?) of the latter described as 2.5
to 3 cm. long and 1.5 em. thick; but its coriaceous leaves are described
as being up to 25 cm. long and 9 cm. wide; they are actually up to
30 em. long and 10 cm. wide.

Some additional descriptive data for this Koorders species are: Leaves
normally 12 to 15 cm. long, 4 to 5 em. wide, firmly chartaceous or sub-
coriaceous, usually olivaceous above, pale brownish beneath and glandu-
lar-punctate when dry; primary nerves up to 15 on each side of the
midrib, slender but distinct and somewhat elevated on the lower surface,
rather irregular, anastomosing and forming a distinct intramarginal
nerve 3 to 5 mm. from the margin, and with a less distinct secondary
marginal nerve close to the edge of the leaf; alternating with the
primary nerves are less conspicuous secondary ones. Flowers appar-
ently small (only unopened buds seen), the calyces rather densely
puberulent, the buds seen not over 6 mm. long. Stamens many, in five
distinct phalanges. Ovary wholly inferior, 2-celled, the cells with many
ovules. Fruit indehiscent, 1-celled, 1-seeded.

Kjellbergiodendron hylogeiton Burret, Notizbl. Bot. Gart. Berlin 13:
103. fig. 5, 4-6. 1936.
Tristania celebica Koord. ex Koord.-Schum. Syst. Verzeich. 3: 96. 1914, nom.
ud., syn. n

OV.

The type is Kjellberg 2016, from Malili, Celebes, at sea level, its
mature flowers not yet known. I have not seen the type, but confidently
refer here Neth. Ind. For. Serv. Cel. 334 (L), two sheets, one with very
immature flower buds, the other with immature fruits, and also Cel.
II-261 and bb. 23269 (A), all from Malili, Celebes. On the sheet with
the immature fruits of Cel. 334, the leaves are up to 30 em. long and
10 cm. wide, the stout infructescences up to 20 cm. long, and the imma-
ture fruits (seeds not formed) 2 to 2.5 em. long. It is, of course, pos-
sible that some of the smaller leaved sterile specimens above referred
by me to K. limnogeiton Burret may belong with K. hylogeiton Bur-
ret. One concludes that it would perhaps be better not to name sterile
specimens to the species in this difficult assemblage. I cannot help
but feel that certain sterile specimens, and one or two with very imma-
ture inflorescences from Malili, partly distributed as representing the
undescribed Tristania celebica Koord., mentioned above, belong with
this Burret species. Some of these are Neth. Ind. For. Serv. Cel. IV-106,
Cel. 106, Cel. 166, Cel. 193, bb. 18800, bb. 26286 (all A).

Kjellbergiodendron limnogeiton Burret, Notizbl. Bot. Gart. Berlin
13: 103. fig. 5, 1-3. 1936.
The type of this, which I have not seen, is Kjellberg 2170 from
Towuti, Celebes, alt. 300 m., on lake margins, its leaves coriaceous,
15-19 em. long, 5-6 em. wide, its fruits 5-6 em. long. Burret’s flower-

1952] MERRILL, NOTES ON XANTHOSTEMON 165

ing material was very scanty and unsatisfactory, yet he described the
flowers as “majusculi,” and, as illustrated, about 2 em. in diameter and
1.2 em. long (he did not actually specify these measurements; his illus-
tration shows only a single attached flower). This is a much larger
flower than one would suspect to develop from the small buds of the
Koorders species, as these, nearly mature, are but 6 mm. long. I am
certain that the following specimens from Malili, Celebes, represent
this Burret species; Neth. Ind. For. Serv. bb. 23544 (A), its one fruit
4.5 em. long, 3 em. thick; bb. 18920 (A,L), its flower buds about 1.4 cm.
long, glabrous, oblong-obovoid, the stout calyx tube 4 mm. thick,
elabrous, narrowed below, rugose, and 4-5 mm. long, the concave im-
bricate petals suborbicular or obovate, rounded, at least 1 cm. long.
Sterile but almost certainly belonging here in spite of their, in general,
smaller leaves are: Neth. Ind. For. Serv. Cel. IT. 240; IT. 241; II. 248
(A,L); Cel. IT. 242, 253, 485 (L); bb. 19820 (A,L); bb. 22730, 23544,
29974 (A); and Boschwezen Mantri bb. 1836, 1854, 1890 (L), from
the Malili region.

ARNOLD ARBORETUM,
HARVARD UNIVERSITY.

166 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxmt

NOTES ON THE FLORA OF CHINA, II
SHIU-YING Hu
With two plates

In 1947 K. M. Feng of the Fan Memorial Institute of Biology made a
botanical expedition in southeastern Yunnan, in the region immediately
east of Mengtze where A. Henry made his classical collections. From
August 5 to December 20 of that year Feng collected 2985 field numbers
in a relatively small area between longitude 104° and 105° East, on the
Tropic of Cancer and a half-degree south. From working over this
collection at the Arnold Arboretum it is apparent that a number of new
entities are included. Below are described a new genus, Styrophyton, of
the Melastomataceae, and several new species of the same family. Critical
notes on other taxa in this family are recorded. The material is arranged
alphabetically according to the genus and species. All the collections cited
belong to the herbarium of the Arnold Arboretum.

Since the system of transliteration employed in Feng’s filed labels differs
from that used in the Gazetteer of Chinese Place Names based on the
Index to the V. K. Ting Atlas compiled by the United States Board on
Geographical Names and from that of the Map of China published in
1945 by the National Geographic Society of America, the equivalent
spellings for the major localities are given below so that the reader may be
helped to find the places on standard maps. After Feng’s transliteration,
those of the other two systems are entered with their abbreviations USB
and NGS following: Ma-kuan, Ma-kuan (USB), Makwan = Anping
(NGS); Mar-li-po, Ma-li-p’o (USB), Malipo (NGS); Si-chour, Hsi-ch’ou
(USB), Sichow (NGS); Wen-shan, Sen-shan (USB), Wenshan = Kaihua
(NGS)

Allomorphia urophylla Diels in Bot. Jahrb. 65: 102. 1932.—Li in
Jour. Arnold Arb. 25: 10. 1944
YUNNAN: Si-chour-hsien, Shiang-pyng-shan, in mixed forest by stream, alt.
1400-1500 m., K. M. Feng 11468 (shrub 5 ft. high, flower pink, common);
Ma-kuan-hsien, Ching-kou (Lao-chiin-shan), in open thickets, alt. 1100-1500 m.,
K. M. Feng 13675 (shrub 5 ft. high, fruit green, common).

As far as we know, Feng 11468 establishes the eastern and northern
limit of the range of the species.
Blastus fengii sp. nov.

Frutex ca. 1.3 m. altus, ramis subquadrangularibus, glabris, tuberculatis,
ramulis junioribus squamuloso- glandulosis, glaberrimis; foliis chartaceis,
junioribus glandulosis, oblongo-ovatis vel ellipticis, 8— 14 cm. longis, 4—7.5
cm. latis, basi obtusis raro rotundatis, apice acuminatis, acumine 2 cm.

1952] HU, NOTES ON THE FLORA OF CHINA, II 167

longo, margine integris setosisque, nervis primariis 5, marginalibus duobus
tenuioribus additis, nervis transversis conspicuis, supra inter nervos sparse
setosis, subtus glabris, petiolo 2-4.5 cm. longo, squamuloso-glanduloso,
glabro; cymis subsessilibus, 3-floris, ex axillis foliorum delapsorum
productis; flores ignoti; pedicellis fructuum 9-12 mm. longis; capsulis
subglobosis, 5 mm. diametro (Tab. II, fig. 1), postatis 8 inconspicuis,
glaberrimis, sepalis persistentibus, semi- rotundatis, 2-3 mm. longis latisque,
glabris; seminibus subovoideis, plus minusve curvatis (Tab. IT, fig. 1),
0.4 mm. longis, tuberculatis, breve rostratis.

YUNNAN: Mar-li-po, Chung-dzia, in open thickets, alt. 1800-2100 m., K. M.
Feng 12726 (type) (herb 3 ft. high, fruit yellowish red, common); same region,
Sze-tai-po, in mixed forest, alt. 1600-2000 m., K. M. Feng 13780 (shrub 4 ft.
high, flower pink, common).

In its axillary inflorescences situated at the mature portion of the stem
where the leaves are often absent, and in its long pedicellate fruits with
persistent roundish sepals, this species appears to be nearest to Blastus
hirsutus Li. The latter species can readily be recognized by its hirsute
branchlets and fruits.

Cyphotheca hispida sp. nov.

Herba 1 m. alta, ramis subteretibus, 2 mm. diametro, hispidis, junioribus
incanis; foliis membranaceis, oblongo-ovatis, 5.5-10 cm. longis, 3-5 cm.
latis, basi rotundatis vel subcordatis, apice acuminatis, acumine 1 cm. longo,
margine integris hispidisque, nervis primariis 5, nervis transversis supra
obscuris, subtus elevatis, supra inter nervos sparse setosis, basi glanduloso-
hirtellis, subtus hispidis nervis praesertim, petiolo 1-5 cm. longo, hispido;
inflorescentiis subumbelliformibus cymis, terminalibus, 3- vel 4-floris,
pedunculis 8 mm. longis; flores 4-meri (Tab. II, fig. 10), pedicellis 9 mm.
longis, receptaculis subcampanulatis, hispidis; sepalis 4, ovatis, hispidis,
2 mm. longis, acutis; petalis 4, rubris, obovatis, 8-10 mm. longis; stamini-
bus 8, inaequalibus, antheris incurvis, apice attenuatis, ee, thecis
basi nonproductis, majoribus 5 mm. longis, minoribus mm. longis,
connectivis postice incrassatis; ovario ad receptaculum ae adnato,
apice collo tubiformis ieiniato laciniis glanduloso-ciliatis ornato, stylo
paulo curvato, glanduloso-hirto; fructibus ignotis.

YUNNAN: Si-chour-hsien, Faa-doou, in mixed forest, alt. 1450-1550 m.,
K. M. Feng 11746 (tyre) (herb 3 ft. high, flower pink, common)

The thickened connectives of the anthers of the smaller stamens, the
glandular-ciliate corona at the apex of the ovary, and the hirtellous style
of this species suggest a relationship with Cyphotheca montana Diels.
Cyphotheca hispida can readily be distinguished from the Diels species
by its hispid stem and leaves, its smaller habit, and the lack of any evident
overgrowth of the thecae. The thecae of the smaller stamens of Cyphotheca
montana are slightly produced at the lower front ends.

It is interesting to note that, although the form of the seeds of the

168 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxxu1

Melastomataceae has been employed as a major character for the delimita-
tion of the genera or sections of the family, the nature of the seeds of
Cyphotheca has been lacking in botanical descriptions. It seems fitting
to add here a note concerning it: Seeds numerous, subovoid-cuneate,
beaked (Plate II, fig. 3), 0.8 mm. long, 0.5 mm. wide at the truncate end,
the testa brown, distinctly tuberculate, the beaked side with a shiny black
aril. This description is prepared from a specimen (T. T. Yu 16627) of
Cyphotheca montana Diels.

Fordiophyton longipetiolatum sp. nov.

Frutex ca. 1.3 m. altus; ramulis glabris; foliis membranaceis, inaequali-
bus, lanceolatis, majoribus 9-14 cm. longis, 3—4 cm. latis, minoribus 3—6.5
cm. longis, 1-2 cm. latis, basi cordatis, apice acuminatis, acumine 1—2.5
cm. longo, margine obsolete serrulatis, nervis principalibus 5, paulo elevatis,
petiolo 1-1.4 cm. longo; inflorescentiis paniculatis; pedunculis 2—3 cm.
longis; ramis strictis patentibus, glanduloso-setulosis, bracteolatis; cymis
singularibus 1- to 3-floris; bracteis cordatis, 2 mm. longis, 2—3 mm. latis,
glabris, persistentibus; floribus pedicellatis, pedicellis 4-6 mm. longis
glabris, receptaculis anguste infundibuliformibus, 6-7 mm. longis, glabris,
rare 1—3 glandulosis setis; sepalis deltoideis, 2 mm. longis, 2 mm. latis,
apice seta glandulosa; petalis rubris, apice in setulam exeuntibus; stamini-
bus 8, inaequalibus, antheris majoribus linearibus, 15 mm. longis, ‘thecarum
basta cornuto-productis, antheris minoribus oblongis, 4 mm. longis,
flavis, thecarum basibus productis; capsulis ignotis.

YUNNAN: Si-chour-hsien, Faa-doou, in mixed forests, alt. 1450-1550 m.,
K. M. Feng 11829 (type) (shrub 4 ft. high, flower pink-rose, rare).

In general appearance this species suggests Fordiophyton strictum Diels
and is probably most closely related to it. The latter species can readily
be distinguished from F. longipetiolatum by its subsessile leaves and
glandular-setose bracts and sepals

Medinilla nana sp. nov.

Frutex humilis, 0.5—1 m. altus; ramis cinereis, rugosis et verruculosis;
ramulis quadrangularibus, brunneis, internodiis 1-3 cm. longis, alatis,
axillis foliorum setis sparse vestitis; foliis coriaceis, obovatis, 1-2.3 cm
longis, 0.7-1.7 cm. latis, paucidenticulatis, ad apicem 2 vel 3 dentibus
minutis, basi obtusis, apice rotundis et retusis, costa supra impressa,
subtus elevata, nervis lateralibus obscuris, petiolo 1-3 mm. longo; in-
florescentiis terminalibus, solitariis vel cymosis simplicibus, cymis 2-floris;
pedunculis 5 mm. longis; bracteis ovatis, glanduloso-erosis; pedicellis 2—3
mm. longis; flores ignoti; baccis oblongo-subglobosis, 8 mm. longis, 6 mm.
diametro; sepalis persistentibus rotundatis, erosis et glandulosis; seminibus
numerosis, minutis, ovoideis, 1 mm. longis, 4-5 mm. diametro, testis
reticulatis.

YUNNAN: Si-chour-hsien, Faa-doou, in mixed forest, alt. 1500-1600 m.,
K. M. Feng 11920 (shrub 1 ft. high, fruit green, common); Mar-li-po, Chung-

1952] HU, NOTES ON THE FLORA OF CHINA, II 169

dzai, in mixed forest, alt. 1600-1800 m., K. M. Feng 12784 (type) (shrub 3 ft.
high, fruit green, common).

I can suggest no close relative for this very distinct species. Its broadly
winged paired branchlets, its small obovate paucidenticulate coriaceous
leaves, and its terminal solitary flowers or simple cymes are unlike any
known species of Medinilla. Nevertheless, the fleshy berries with their thin
calyx tube and exterovarial chambers place the species in Bakhuizen’s
section Hetero-Medinilla of this genus. The small ovoid seeds with their
reticulate testas suggest relationship to Pachycentria fengi S. Y. Hu.

Medinilla petelotii Merr. in Univ. Cal. Publ. Bot. 13: 137. 1926.

YUNNAN: Si-chour-hsien, Ting-mann, in mixed forests, alt. 1300-1500 m.,
K. M. Feng 12334 (scandent shrub 5 ft. high, flowers pink-rose, rare); Mar-li-po,
Huang-jin-in, in mixed forests, alt. 1300-1400 m., K. M. Feng 13063 (shrub
5 ft. high, flowers lavender, rare); same region, Sze-tai-po, Loa-chiin-shan, in
mixed forest, alt. 1300-1500 m., K. M. Feng 13946 (shrub 4 ft. high, fruit green,
common).

Feng 13063 fits Merrill’s description well, and it matches the photograph
of Pételot 1791 (type of Medinilla petelotii Merr.) in every respect. The
fertile portion of Feng 12334 appears to have been a section from the lower
part of a creeping stem. It has many fibrous adventitious roots. The
inflorescences on such a stem are much more ramified and also appear
longer than the typical ones as represented by Pételot 1791 and Feng 13063.
With all other aspects identical, I am inclined to think that the larger
inflorescence is due to the vigor of the plant.

Tsai 60315 from Ping-pien-hsien, Yunnan, appears to belong here.

Melastoma normale D. Don, Prodr. Fl. Nepal. 220, 1825.— Rehd. &
Wils, in Sarg. Pl. Wils. 2: 421. 1915.— Li in Jour. Arnold Arb. 25:
8. 1944.

YUNNAN: Si-chour-hsien, Faa-doou, in open thickets on rock, alt. 1450-
1550 m., K. M. Feng 11870 (shrub 4 ft. high, fruit green-yellow, common);
Mar-li-po, Sze-tai-po (Loa-chiin-shan), in open thickets by stream, alt, 1100-
1300 m., K. M. Feng 14053 (herb 3 ft. high, flowers pink-purple, common).

Feng’s designation of the habit of the plant as represented by his
number 14053 does not seem to be appropriate, for our specimen of this
number exhibits a strong woody third-year stem. In the herbarium this
species is often confused with Melastoma candidum D. Don, and in
botanical literature the diagnostic characters given are rather inadequate,
since they were drawn from the indumentum of the stem and the shape of
the leaf-base. Both of these characters vary with the ecological conditions
under which the plant grows and the age and vigor of the plant. Melastoma
normale LD). Don is generally recorded as, ‘‘stem with long spreading hair,
leaves with round and obtuse base,’’ and Melastoma candidum D. Don
as ‘‘stem with appressed scale-like hairs and leaves with subcordate bases.”
Round and subcordate bases are sometimes very hard to distinguish, and

170 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxuII

the spreading hairs are so coarse and dented that they ae scaly.
Detailed examination of our material seems to reveal two constant
characters that can be employed in the diagnosis of these two ae In
Melastoma normale D. Don the sepals are linear-lanceolate and the
anterior lobes of the connectives of the large stamens are small, round,
and almost undivided, while in Melastoma candidum D. Don the sepals
are broad deltoid and the anterior lobes of the connectives of the large
stamens are pointed and over 2 mm. long. In general Melastoma normale
D. Don has lanceolate leaves and smaller globose fruits, and the leaves
of Melastoma candidum D. Don are ovate and the fruits larger and oblong.

Osbeckia chinensis Linn. Sp. Pl. 345. 1754. — Li in Jour. Arnold Arb.
aot
YUNNAN: Si-chour-hsien, Faa-doou, on open grassy slopes, alt. 1450-1550
m., K. M. Feng 11765 (herb 3-10 in. high, flowers pink, common).

This is probably the most widely distributed species of the Melastoma-
taceae in China. It occurs in the coastal region from Foochow to Hongkong
and thence westward over all the warm temperate region of China. It
can readily be recognized by (1) the small herbaceous habit; (2) the
lanceolate leaves; (3) the terminal simple cymes; (4) the stellate tufted
calyx tubes; (5) the persistent corona of rigid hairs at the apex of the
ovary; and (6) the glabrous urn-shaped fruits. The vernacular name
“ien-hsiang-lu” (heavenly incense burner) is a very good connotation of
the characteristic fruit.

Oxyspora paniculata DC. Prodr. 3: 123. 1828.—Li in Jour. Arnold
Arb, 25: 12. 194

YUNNAN: Wen-shan-hsien, Hwang-tsao-bah, by streams on scrub, alt. 1600
m., K. M. Feng 11011 (herb 2 ft. high, flowers lavender-red, common) ; Si-chour-
hsien, Shiang-pyng-shan, in mixed forests by streams, alt. 1400-1500 m., K. M.
Feng 11472 (shrub 5 ft. high, flower rose-pink, common); Mar-li-po, Sze-tai-po
(Loa-chiin-shan), in open thickets by streams, alt. 1300-1500 m., K. M. Feng
14037 (shrub 3 ft. high, flower rose-pink, common)

Oxyspora paniculata DC. is the type species of the genus. It has very
unique seeds, the form of which is so characteristic that I have not seen
the like in any other Asiatic genus of the Melastomataceae. The embryo
proper is straight, club-shaped, or subovoid. The testa is so produced
that the seed appears cuneate. It is beaked on the aril side of the truncate
end, and opposite the beak there is a distinct spur which is longer than
the beak. In a profile view the seed appears like the head of an adze
(Plate IT, fig. 4).

Pachycentria fengii sp. nov.
Frutex sempervirens, epiphyticus, scandens, ramulis teretibus, glabris,

fusco-rubescentibus, extimis compressis, lenticellis albis, prominentibus;
foliis subcoriaceis, integerrimis, obovatis, 3.5-8.5 cm. longis, 2—3.5 cm

1952] HU, NOTES ON THE FLORA OF CHINA, II Lia

latis, basi cuneatis vel obtusis, apice acuminatis, acumine 5-10 mm. longo,
obtuso, trinerviis, nervis supra planis, subtus elevatis, reticulis obscuris,
petiolo 6-10 mm. longo, glabro; flores ignoti; pe es fructiferis
corymbosis, laxis, terminalibus; pedunculis 1-1.7 cm. longis; pedicellis 5
mm. longis; ae urceolatis, 6-7 mm. longis, 4— 1 mm. diametro (Tab.
II, fig. 5~7); sepalis persistentibus deltoideis, apice postice minutis uncis
instructis; seminibus numerosis, minutis, ovoides, 1 mm. longis, 0.7 mm.
diametro, testis reticulatis (Tab. II, fig.

YUNNAN: Si-chour-hsien, Faa-doou, on trees in mixed forest, alt. 1500-1550
m., K. M. Feng 11789 (type) (shrub 4 ft. high, fruit green, pedicel red, common).

Pachycentria Blume was first discovered in Java. The center of its dis-
tribution is the Malay Archipelago. Hayata in 1912 described a species
from Formosa and Ridley in 1915 published another one from the Malay
Peninsula. Pachycentria fengii S. Y. Hu seems to be the first species
recorded from the Chinese mainland. It sets the northern limit for the
genus. To Blume, Pachycentria differs from Medinilla in: “1) tubo
calycis sursum epathiiocni-dilatati circa ovarium fortius constricto; 2)
petalis acuminatis; 3) staminum structura diversa, quum antherarum
connexivum in Medinillis omnibus ad basin antice auriculas duas et
postice calcar subulatum exserat, hic autem tantummodo postice calcari
parvo crassiusculo muniatur.”’ Bakhuizen f. regarded the degree of union
of the ovary with the calyx tube to be more significant for the generic
delimitation. He maintained that in Pachycentria the ovary is “wholly
concrescent with the calyx tube,” while in Medinilla it is “adnate to the
calyx tube with longitudinal septs.’ The calyx tubes of the Formosan
and the Chinese material are not so much constricted as those of the
Malaysian species. Nevertheless, the connection between the ovary and
the calyx tube is complete. There does not seem to be any doubt that our
material is a species of Pachycentria.

Phyllagathis wenshanensis sp. nov.

Herba, caulibus petiolisque dense longe crispo-hirsutis, rhizomate
prostrato, ramis teretis, mm. diametro; foliis valide membranaceis,
orbicularibus, 5-7 cm. longis latisque, olivaceo-viridibus, basi perspicue
cordatis, apice rotundatis et plerumque emarginatis, nervis primariis 5 vel
7, nervis transversis supra obscuris, subtus perspicuis et dense crispo-
hirsutis, margine integris et ciliatis, supra disperse breve setosis et molliter
crispo-hirtis, subtus hirsutis, nervis dense crispo-hirsutis, petiolo 2.5—4.5
cm. longo; inflorescentiis umbellatis, terminalibus, 4-floris, pedunculis
11 cm. longis, hirtellis; flores ignoti; pedicellis fructuum 1 cm. longis,
hirtellis; calycibus persistentibus deltoideis, acutis; capsulis turbinatis,
hirtellis, 5 mm. longis; apice 5 mm, diametro, placenta supra ramificata,
seminibus ovoideis, 0.8 mm. longis, 0.4 mm. diametro, truncatis, tubercula-
tis, breve rostratis.

YUNNAN: Wen-shan-hsien, Lao-jiun-shan, in mixed forest, alt. 2300 m.,
K. M. Feng 11186 (type) (herb 4 in. high, fruit greenish, rare).

172 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXXIII

The creeping habit and rounded leaves of this species suggest a relation-
ship with Phyllagathis cavaleriei (Lévl. & Van.) Guill., which has glabrous
capsules and leaves with long setose hairs.

Plagiopetalum henryi (Kranzl.) comb. nov.

Barthea cavaleriei Lévl. in Fedde, Rep. Spec. Nov. 8: 61. 1910, pro parte,
quod Esquirol 215.
Sonerila henryi Kranzl. in Viert. Nat. Ges. Ziirich 76: 152. 1931.
Plagiopetalum esquirolii sensu Li, in Jour. Arnold Arb. 25: 10. 1944, non Rehder.
YUNNAN: Wen-shan-hsien, Pyng-bah, moist sandy margin of mixed forest,
alt. 1700 m., K. M. Feng 11028 (shrub 2 ft. high, flower pinkish-red, rare); Mar-
li-po, Pan-chia-chii, roadside, alt. 1800-2100 m., K. M. Feng 12674 (shrub 3 ft.
high, fruit red, common).

Regarding Plagiopetalum Rehder stated, ‘‘In its floral structure it seems
nearest to Sonerila, but that genus is easily distinguished by its trimerous
flowers and symmetrical petals.” If Plagiopetalum is accepted as a genus
at all, then the transfer of Sonerila henryi Krinzl. is justified because of its
tetramerous flowers and asymmetrical petals. Li (1944) interpreted
Sonerila henryi Kranzl. and Plagiopetalum esquirolii (Lévl.) Rehd. to be
conspecific. After detailed examination of the materials at hand, including
Wilson 3261, the type of Plagiopetalum quadrangulum Rehder which, in
turn is the type species of the genus and has been interpreted by both
Diels and Rehder as synonymous with Plagiopetalum esquirolii (Lévl.)
Rehder, I find it hard to accept this view. Plagiopetalum esquirolii (Lévl.)
Rehd., as typified by Wilson 3261, represents a plant devoid of scabrous
hairs on the petioles, peduncles, pedicels and receptacles, while these parts
of the specimens that I designated as Plagiopetalum henryi (Kranzl.) S. Y.
Hu are scabrous.

When Léveillé published Barthea cavaleriei he cited three specimens,
namely Cavalerie 1552 and Esquirol 215 and 1581. He designated no type.
Diels (1932) segregated Cavalerie 1552 and transferred it to Bredia. Here
in our herbarium there is a photograph and a good fragment of Esquirol
215 which is identical with A. Henry 9077, an isotype of Sonerila henryi
Krinzl. Since Barthea cavaleriei Lévl. is a synonym of Bredia cavaleriet
(Lévl.) Diels, the next published specific name, is here adopted.

The seed of Plagiopetalum henryi (Kranzl.) S. Y. Hu has a very unique
form. It is oblong in outline, 1 mm. long, 0.5 mm. in diameter, obscurely
papillate, and appendaged on three sides (Plate I, fig. 11).

Sarcopyramis bodinieri Lévl. & Van. in Mem. Soc. Nat. Sci. Nat.
Cherbourg. 35: 397. 1906; et in Fedde, Rep. Spec. Nov. 4: 95. 1907.
Sarcopyramis nepalensis var. bodinieri (Lévl. & Van.) Lévl., Fl. Kouy-Tchéou

Sarcopyramis nepalensis sensu Diels in Bot. Jahrb. 65: 111. 1932. — Li in Jour.
Arnold Arb. 25: 25. 1944, non Wallich.

1952] HU, NOTES ON THE FLORA OF CHINA, Il 173

YUNNAN: Wen-shan-hsien: Loa-jiun-shan, in mixed forest, alt. 2300 m.,
K. M. Feng 11188 (herb 3 inches high, leaves green above, purple below, fruits
green and purple, common).

Sarcopyramis bodinieri Lévl. & Van. has been misinterpreted by several
authors as Sarcopyramis nepalensis or its variety. Additional material in
the flowering stage reveals certain characters which prove such a concept
to be wrong. The type sheet of Sarcopyramis bodiniert Lévl. & Van. (E.
Bodinier 2393) contains five specimens, four with small colored leaves and
one with larger green leaves. Guillaumin (Bull. Soc. Bot. France 60: 343.
1913) suggested that the plants might represent a smaller form of S. napal-
ensis Wall. Léveillé, without giving any reason, published it (1914) as
such. Diels (1932) actually sank it into the synonymy of Sarcopyramis
nepalensis, and his view has been accepted by later authors. In Feng 11188
we have five plants which match the small plants of the holotype of Sarco-
pyramis bodinieri Lévl. & Van. Careful examination of these specimens as
well as of the fragments of Bodinier 2393 reveals that besides the smaller
size of the plants and the purple color of the foliage there are certain floral
characters which distinguish them as a distinct species. In Sarcopyramis
napalensis Wall. the bracts are ovate, acute, the calyx lobes are truncate
and ciliate, and the anthers are subelliptic with the notched apical ends
slightly pointed. In Sarcopyramis bodinieri Lévl. & Van. the bracts are
spathulate, the calyx lobes are eciliate, each with two horn-like projections,
and the anthers are obcordate, each not much longer than the upward spur
on the posterior side of the connective. The leaves of Sarcopyramis bodiniert
Lévl. & Van. are sparsely setose above and glabrous beneath with an acute
apex, while those of Sarcopyramis napalensis Wall. are elliptic or ovate-
elliptic with an acuminate apex. Judging from the shape, the texture and
the pubescence of the leaves, and the horned calyx lobes, Sarcopyramis
bodinieri Lévl. & Van. is nearer to the Kwangsi specimens which have been
named as Sarcopyramis delicata C. B. Rob. than to Sarcopyramis napalensis
Wall. All specimens of the Kwangsi material have solitary flowers. hee se
Chiao 1304 from Ya-an, Sikang, with small ovate leaves and umbellate
inflorescences, definitely belongs here.

Sarcopyramis napalensis Wall., Tent. Fl. Napal. 32, pl. 23. 1824. —
Li in Jour. Arnold Arb. 25: 25. 1944.
YUNNAN: Mar-li-po, Sze-tai-po (Lao-chiin-shan), in mixed forest, alt. 1300-
1500 m., K. M. Feng 13974 (herb 4 in. high, fruit green, common).
Sonerila cantonensis Stapf in Ann. Bot. 6: 302. 1892.—Li in Jour.
Arnold Arb. 25: 35. 1944.

YUNNAN: Si-chour-hsien, Faa-doou, in mixed forest, alt. 1500-1550 m.,
K. M. Feng 11877 (Herb 1 ft. tall, flower pink-rose).

This species has been recorded from Kwangtung, Kwangsi, and Fukien.
This is a new record for the flora of Yunnan.

174 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxx1I
a Seat J. Jeffrey in Notes Bot. Gard. Edinb. 8: 207.

YUNNAN: Si-chour-hsien, Ting-mann, on cliffs in mixed forests, alt. 1100-
1200 m., K. M. Feng 12349 (herb 4 in. high, fruit green, rare).

Jeffrey described the seeds of this species as ‘“‘seminibus ovoideis minute
punctulatis.” This is true. But on closer examination of Feng 12349 as
well as of Henry 12337, the latter an isotype of the species, a very inter-
esting character is revealed, concerning which it is worth while to add a
note. This character is the presence of a two-lobed bladder at the apical
end of the seed opposite the shiny dark brown aril (Plate II, fig. 9). Seeds
of Sonerila cantonensis Stapf, Sonerila hainanensis Merr., Sonerila picta
Korth., and Sonerila tenera Royle were also examined for comparison.
With the exception of Sonerila tenera Royle, all these species have seeds
with the bladder at the apical end of the aril side. This character correlates
with the attenuated anther and the funnel-shaped fruit. One must conclude
that such a character is at least subgenerically distinct.

Stapfiophyton erectum sp. nov.

Herba erecta, 0.75 m. alta, ramis subteretibus, 5 mm. diametro, cinereis,
junioribus furfuraceis; foliis subcoriaceis, glabris, ellipticis, 10-12 cm
longis, 3.5—5 cm. latis, basi obtusis, apice caudato-acuminatis, acumine
1.5—2 cm. longo, subtus glandulosis, nervis primariis 3, marginalibus duo-
bus tenuioribus additis, nervis transversis supra obsoletis, subtus con-
spicuis, margine integro; petiolo 1.5—2.5 cm. longo, glanduloso, glabro;
inflorescentiis subumbelliformi-cymis, paniculatis, terminalibus, pedunculis
3 cm. longis, cymis 3- vel 4-floris, pedicellis 4-5 mm. longis, glabris; flores
ignoti; capsulis subturbinatis, 3-4 mm. diametro, apice depressis, placenta
supra ramificata, seminibus ad basim adnatis; seminibus ovoideis, 0.7 mm.
longis, 0.4 mm. diametro, rostratis, testa coe

YUNNAN: Mar-li-po, Huang-jin-in, in mixed forest, alt. 1300-1500 m.,
K. M. Feng 13082 (type) (herb 2 ft. high, fruit whitish yellow, common).

By its ramified placenta this species can be placed either in Phyllagathis
Blume or Stapfiophyton Li. It is because of its paniculate cymose inflo-
rescences that the latter is chosen.

Styrophyton gen. nov.

Anerincleistus ? sensu Diels in Bot. Jahrb. 65: 101. 1932, non Korthals.

Allomorphia sensu Li in Jour. Arnold Arb, 25: 11. 1944, non Blume.

Inflorescentiae spicatae (Tab. I, fig. 1), floribus sessilibus, 4-meris;
receptaculis subcampanulatis, strigosis; sepalis brevibus; petalis unguicu-
latis (Tab. I, fig. 5); staminibus 8, subaequalibus, antheris rectis, apice
paulo attenuatis, 1-porosis, thecis antice basi paulo productis, connectivo
postice non aucto, ovario basi receptaculi adnato, stylo attenuato, basi setis
ornati, stigmate capitato. Fructus ovoideo-subglobosus, costis pallidis et
prominentibus, quadrilocularibus; semina numerosa, minuta, cuneata,

1952] HU, NOTES ON THE FLORA OF CHINA, II 175

truncata et rostrata. Frutices, ramis dense tomentosis; foliis magnis,
ovatis, petiolatis.

Type species: Styrophyton caudatum (Diels) S. Y. Hu.

In all recent monographic work on Melastomataceae the inflorescences,
whether they are panicles, umbels, scorpioid cymes, or otherwise, have been
used as key characters for generic demarcations. Diels in 1932 first saw
Henry 10761, a specimen from Yunnan with a spicate inflorescence. Realiz-
ing that it was unique, he described it as new, and probably because of its
superficial resemblance to Anerincleistus Beccarii Cogn., he doubtfully as-
signed it to Anerincleistus Korthals. Li in 1944, on the basis of the promi-
nent striation of the fruits, transferred Diels’ species to Allomorphia
Blume and added a description of the immature fruit and seed. With
mature fruit from Feng’s southeastern Yunnan collection I made careful
dissections of the flower of Henry 10761 (isotype), the young fruit of
Tsai 61591, and the mature fruit of Feng 12291, and compared each with
those of Anerincleistus Korthals and Allomorphia Blume. After a review
of all the literature concerning these two genera and a detailed examina-
tion of all the material in the Arnold Arboretum and the Gray Herbarium,
I feel that the southeastern Yunnan plant is generically distinct. Besides
the simple unramified spicate inflorescence, which at once distinguishes it
from the paniculate type found in Allomorphia Blume, the short calyx tube
is also very distinct. In Allomorphia Blume the calyx tube is tubular, at
least twice as long as wide, and with the ovary attached only at its base.
Consequently, in fruit, the free portion of the persistent calyx tube forms
a tall collar, which is usually constricted at the base. Moreover, at the
apex of the ovary in Allomorphia there is no appendage such as bristles or
corona, and consequently the apical end of a mature fruit is smooth and
concave. In Styrophyton the calyx tube (Plate I, fig. 6) is only slightly
longer than wide, and in fruit it does not form an evident collar (Plate I,
fig. 2). Moreover, the persistent bristles at the apex of the ovary render
the apical end of the mature fruit rough. The cuneate, truncate, and
beaked seed and the striate fruit of Styrophyton indicate a close relation-
ship with Allomorphia Blume on the one hand, and its short calyx tube
and the rough apical end of the fruit show affinity with Anerincleistus.
Korthals on the other hand. These characters, together with the true
spicate inflorescence, constitute my reasons for proposing a new genus
whose natural position lies between Allomorphia Blume and Anerincleistus
Korthals. Species of the latter genus can easily be distinguished by their
pedicellate flowers, paniculate inflorescences, prominent corona at the
apex of the ovary, and smooth capsules with persistent corona exposed at
the apical end of the fruit.

Regarding the fruit and seed characters of this taxon Li (1944) recorded,
“The somewhat immature fruits are... one-celled . . . seeds very
minute, oblong.” This is far from correct. Both the cross- and the longi-
tudinal sections of the mature, as well as the young fruit, appear to have
four cells (Plate I, figs. 3 & 4). The seed of the mature fruit is oblong-

176 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxxim

cuneate, 0.6-0.7 mm. long, 0.3-0.4 mm. in diameter, indistinctly ridged,
truncate and beaked, slightly grooved on the beaked side and appendaged
at the proximal end (Plate I, fig. 7). The testa is brown and smooth.

Styrophyton caudatum (Diels) comb. nov.

Anerincleistus ? caudatus Diels in Bot. Jahrb. 65: 101. 1932.

Allomorphia caudata (Diels) Li in Jour. Arnold Arb. 25: 11. 1944.

YUNNAN: Si-chour-hsien, Ting-mann, in mixed forests, alt. 1200-1300 m.,
K. M. Feng 12291 (shrub 6 ft. high, young fruit green, common). Other speci-
mens that I have examined are: A. Henry 10761 (1sotyPE) from Mengtze, H. T.
Tsai 60437, 60563, 61279, and 61591, all from Ping-pien hsien of Yunnan.

This species is a shrub which grows up to 5 meters high. In south-
eastern Yunnan it occurs in mixed forests at altitudes varying from 1000
to 1500 meters. The distribution is rather limited. So far, specimens have
been collected only from the southeastern region of Yunnan between Long.
103° and 105° E. and Lat. 22°50’ and 23°30’ N. Our material exhibits
very little variation even in the vegetative characters. Compared with the
isotype, the base of some leaves may be subcordate rather than round,
occasionally the upper surface of the leaves may be more or even less ver-
ruculose, the pubescence on the lower surface may be brown rather than
white, and between these hairs there may be fewer glands. It seems that
all these variations may be attributed to the age of the plants, the size of
the leaves, or even to the pressing and drying processes in the preparation
of the specimens.

EXPLANATION OF PLATES

PLaTeE I

Styrophyton ee (Diels) S. Y. Hu: Fic. 1. A fruiting branch (x %),
the indumentum ted. Fic. 2. A fruit (X 5). Fic. 3. The a agra eay
section, lower aah not through the center of the fruit (X 10). . 4. Cross-
section of a young fruit (X 10). Fic. 5. A flower (x 5). Fic. 6. big same with
part of the calyx tube, petals and stamens removed (XX 5). Fic. 7. Some seeds
(X 12).

PiateE II

Fic. 1. Fruit of Blastus fengit S. Y. Hu (xX 3), with seeds in lateral view
(X 10). Fic. 2. Habit sketch of Medinilla nana S. Y. Hu (X 2), with seeds
(X10). Fie. 3. Seed of Cyphotheca montana Diels (X 10), side and front
views. Fic. 4. Seed of Oxyspora paniculata DC. (X12). Fic. 5. Longitudinal
section of the fruit of Pachycentria fengit S. Y. Hu (X 6). Fic. 6. Fruit of the
same (X 5), with persistent calyx, each lobe of which has a small hook on the
back. Fic. 7. Cross-section of the ovary of the same, diagrammatic. Fic. 8. Seeds
of Pachycentria fengii S. Y. Hu (X 5). Fic. 9. Seeds of Sonerila yunnanensis
J. Jeffrey (X< 10). Fic. 10. A flower of Cyphotheca hirsuta S. Y. Hu 3
with 1 petal and 1 smaller stamen removed. Fic. 11. Seed of Plagiopetalum
henryi (Krianzl.) S. Y. Hu (Xx 10).

ARNOLD ARBORETUM,
HARVARD UNIVERSITY.

Jour. ARNOLD Ars. VoL. XXXIII PLaTE I

Hu, Notes ON THE FL Lora oF CuIna, II

PraTeE II

Jour. ARNOLD Ars. VoL. XXXIII

aD

SD

Hu, Notes ON THE FL Lora OF CHINA, II

1952] JOHNSON, SPONTANEOUS WHITE PINE HYBRIDS 179

SPONTANEOUS WHITE PINE HYBRIDS
ALBERT G. JOHNSON

With two plates

A NOTEWORTHY FEATURE of the genus Pinus L. is the relative abundance
of both natural and artificial hybrids. Summaries of known crosses pub-
lished in recent years include hybrids within both of the subgenera
Haploxylon Koehne and Diploxylon Koehne, usually within the series
limits of these subgenera; cf. L. P. V. Johnson (3), Richens (5), Righter
and Duffield (6). The hybrids to be described here are hitherto unreported
in the subgenus Haploxylon.

In the spring of 1949 a five-needled pine growing in the Hunnewell
Arboretum, Wellesley, Massachusetts, was observed to differ markedly
from the familiar native white pine, Pinus strobus L.; and yet it was evi-
dently not one of the known cultivated species of this group of pines. The
tree, about twenty years old, twenty-five feet high, and fourteen inches
d. b. h., is, in the opinion of Mr. Walter Hunnewell, of spontaneous origin,
inasmuch as he has no record nor recollection of it having been planted.
Apparently it is a chance seedling which has grown up since annual mow-
ing was discontinued in the Arboretum about 1930. It has been allowed
to remain because of its unusually attractive blue-green foliage.

The tree is a vigorous and rank grower of loose open habit. This rapid
growth capacity plus the effect of repeated weevil damage has caused it to
develop a broad crown with several leaders. Although generally resembling
our native white pine it differs in the curved and twisted blue-green foliage,
the more massive cones, the larger, purple-brown seeds, and the scaly
brown bark. The flowers at the time of initial observation were seen to
differ from those of white pine in both position and color. The red female
flowers were borne from top to bottom of the tree, while on white pine they
rarely are found except in the topmost branches. The flesh-colored male
flowers were few and borne in elongated clusters instead of in yellow com-
pact bunches as in white pine. Subsequent examination of the pollen
showed it to be 23% defective.

The presence in the Arboretum of P. parviflora Sieb. & Zucc. and of
P. parviflora var. pentaphylla Henry, both of which have flowering habits
similar to the tree in question, suggested the possibility of hybridization
with the native white pine, which is abundant on the grounds. Since the
difference between P. parviflora and its variety pentaphylla is slight and
of such a nature as to be obscured in a hybrid, the latter taxon is not being
considered as distinct for the purposes of this paper. Moreover, the loca-
tion of this varietal form in respect to the other trees in the Arboretum
substantially reduces the probability of it having had a rdle in the induc-
tion of this presumed hybrid or of others discovered later.

180 JOURNAL OF THE ARNOLD ARBORETUM _[VvoL. xxxuI

Careful comparison of the characteristics of the tree in question with
both P. strobus and P. parviflora supports the hybrid hypothesis. To facili-
tate comparison of the parental and hybrid forms a hybrid index has been
prepared, TABLE I, after the manner of Anderson (1). In making up such
an index the characters of diagnostic value of one species are arbitrarily
assigned a value of 0, while the corresponding ones of the second species
are assigned values of 1 or more, according to their relative importance
in separating the two species. Since the genetic bases for most of the char-
acters utilized in this particular instance are still imperfectly understood,
the values assigned to the second species have in most cases been kept
relatively low. Values for the corresponding traits in the putative hybrid
are then determined by interpolation according to the degree of resem-
blance to one parent species or the other. The summation of these indi-
vidual values provides an index number for the particular plant indicative
of its relative resemblance to one parental species or the other. Such
indices are of greatest value, of course, in population studies, where they
can assist in the detection of F» segregates and backcross individuals, but
they are also useful tools for presenting quantitatively conclusions neces-
sarily of a subjective nature, as in the case of the present hybrid, where
the summation of its various index values shows it to resemble P. parvi-
flora slightly more than P. strobus.

The reference to odor in the hybrid index table should perhaps be
elaborated, since it is a useful tool in identifying certain pines and the
probable parentage of certain hybrids. Pinus parviflora branches, when
broken, emit a strong and apparently highly specific odor suggestive to
some people of bruised tomato plants. This odor is present in induced
hybrids of this species with P. strobus and P. monticola growing at the
Arnold Arboretum, and is present in the suspected hybrid under discussion
and in other individuals found later. Mirov (private communication) has
suggested that the terpenes and sesquiterpenes of P. parviflora which he
and his co-workers have recently analyzed (2) may predominate in the
hybrid over the simple pinene mixture found in P. strobus (4). A com-
parable situation exists in the hybrid between P. ponderosa Laws. and
P. jeffreyi Grev. & Balf., in which the terpenes of P. ponderosa predominate
over the simple n-heptane of P. jeffreyi, although the turpentines of both
parental species occur in the hybrid.

With the establishment upon a reasonable basis of the probable hybrid-
ity of the tree, it still remained necessary to determine whether the cross
occurred as P. strobus < parviflora or as the reciprocal cross. Several fac-
tors favor the first assumption. The position of the hybrid in relation to
the probable parent trees suggests that it originated as a wind-borne seed
from a near-by P. strobus. An essentially wingless seed of P. parviflora
would have had to be carried by an animal agency nearly twice that dis-
tance to the site of the hybrid. The later finding of additional individuals
of this hybrid, one of which is almost directly below the probable mother
tree, adds further evidence in favor of this parentage. The comparatively
poorer showing in vigor and growth rate of two reciprocal hybrid indi-

TABLE 1

No. or No. oF
oe 2 PE- LENGTH CONE SEEDS SuM oF
LEAF LEAF STROB- STROB- DUN- SEED SEED SEED SCALES PER INDEX
SPECIES LENGTH SHAPE ILI ILI CLE SEEDS WING COLOR SHELL AVER. Opor GRAM Bark VALUES
(0) (0) 0) 0) (0) (0) (0) (0) (0) (0) (0) (0) (0) (0)
Pinus 6-14 straight yellow cream long 5-7 long light thin 38 pleas- 53.08 rough
strobus cm. clus- or mm. brown nt purple
tered pink brown
(1) (1) (2) (2) (1) (1) (0) (2) (1) (1) (3) (3) 1) (19)
2; 7.5- curved pink red- sub- 7-9 long purple thick 30 dis- 13.88 scaly
hunne- 8.5 slightly elong dish ses- mm row agree- brown
welli cm. twisted clus- sile able
tered
(2) (2) (3) (3) (2) (2) (2) (2) (2) (3) (3) (4) (2) (32)
P. parvi- 2-3 urved pink red ses- 10-12 short purple very 13 dis- 4.58 scaly
ra cm twisted elong sile brown thick agree- brown
clust able

182 JOURNAL OF THE ARNOLD ARBORETUM _ [VoL. xxxmI

viduals at the Arnold Arboretum adds further support to this viewpoint
and suggests that some real difference in the reciprocal hybrids may in
fact exist. Also it is reported to be much easier to induce the hybrid
P. strobus X parviflora than the reciprocal hybrid (J. W. Wright, private
communication ).

As already indicated, additional individuals of the hybrid have been
found. The second was discovered in the spring of 1951 about two hun-
dred feet from the site of the first tree and only a short distance from the
probable female parent, a large white pine thirty inches in diameter. This
hybrid tree, designated for purposes of reference as Hunnewell #2, is
about the same age as the first (Hunnewell +1) but is slightly smaller,
twelve inches d. b. h. and twenty-two feet high. Like #1, it is rank-
growing and multiple-stemmed, again in part because of its inherent vigor
and in part due to repeated weevil damage. In details of foliage and cones
it does not differ materially from hybrid #1.

Following the discovery of this second hybrid a systematic search of
the grounds of the Arboretum was made to see if other hybrid individuals
might exist. Additional specimens were found and numbered in order of
discovery as +3, #4, #5, and #6. Tree #3 was later rejected as a
probable hybrid between P. peuce Griseb. and P. parviflora and will be
dealt with independently. To avoid any confusion, however, the original
numbers assigned to the remaining hybrids were retained.

The fall of 1951 provided the first opportunity to collect any number of
cones from the two largest trees. The three smaller ones did not produce
a sufficient number to warrant collection for seed production analysis. In
each case, however, enough were found to supply herbarium material for
comparative purposes. Although badly hit by cone weevils, Conophthorus
coniperda, the first tree yielded sixty-two sound cones and the second seven-
teen. In neither case did this represent the total production of the tree
but instead the more accessible sound cones. These were collected when
mature and just beginning to open on September 7. The two lots of cones
were kept separate, allowed to dry, the seed extracted, cleaned, counted,
and weighed. The good seed was found to be readily separable from the
empty seeds by water flotation, which allowed the latter to be skimmed
off the surface of the water while the filled seed sank to the bottom of the
container. In both cases the proportion of good seed was low, being but
0.9% and 1.6% respectively in trees #1 and #2. If the percentage of
good seed is calculated on the basis of the potential seed production from
the average number of functional scales per cone the figures are further
reduced to 0.1% and 0.4%, since many scales yielded nothing but empty
wings, possibly due to lack of pollination. The seeds produced were large
and relatively thick-shelled. While resembling P. parviflora in these re-
spects, the seed wing was of the long functional type of P. strobus. The
seeds weighed about four times as much as those of P. strobus and one
third as much as those of P. parviflora.

As was pointed out earlier, 23% of the pollen of hybrid #1 was abortive
upon visual examination. That of tree #2, however, appeared wholly

1952] JOHNSON, SPONTANEOUS WHITE PINE HYBRIDS 183

sound. No germination test was undertaken with either pollen lot, so
probable functionability is not known. In both of these trees the amount
of pollen produced was so low in 1951 as to virtually eliminate the pos-
sibility of any significant amount of self-pollination taking place. In the
case of tree #1 similar low pollen production was observed in 1949 ie
1950. In fact, in 1950 scarcely any pollen could be found despite a

heavy production of female flowers. This phenomenon of eae
female flower production is frequently observed in vigorous young pines
and may in part explain the poor seed yield from the cones of such trees.
This point was emphasized in the experience of the writer in the fall of
1950 while collecting pines in southern Mexico. On one occasion a consider-
able number of cones were collected from a small grove of young and ex-
tremely vigorous trees of P. ayacahuite var. veitchii Shaw. The cones of
this species are massive and while green are the equivalent in size and
weight to quite respectable pieces of stove wood, so that some considerable
effort was involved in transporting the cones down the mountain to the
car. When processed some months later, all were found to be devoid of
good seed.

Inasmuch as this hybrid between the native white pine and the Japanese
white pine has occurred at least five times and apparently is capable of
recurring repeatedly as long as the parent species remain in close prox-
imity to each other, it seems appropriate to designate it with a specific title
in honor of Mr. Hunnewell, a lifelong friend of the Arnold Arboretum and
an ardent horticulturist.

x Pinus hunnewelli hyb. nov. (P. strobus L. « P. parviflora Sieb. &
ZNCC.),

Arbor hybrida fertilis habitu corticeque P. parviflorae similis; a P. parvi-
flora differt foliis longioribus, strobilis longioribus, seminibus minoribus
ala elongata donatus; a P. strobo differt ramulis puberulentis, foliis brevi-
oribus, strobilis subsessilibus, apophysis firmioribus crassioribus glaucis;
seminibus majoribus

PE: a eas oer hybrid #1 deposited in the herbarium of the
Arnold Arboret

This hybrid swarm, although limited to small numbers, illustrates the
breakdown of a geographically imposed isolating barrier to genic exchange
through the chance migration of one species, in this case throuh the agency
of man, into contact with another closely related one. The production of
fertile hybrids exemplifies the first stage of introgression of P. parviflora
genes into P. strobus and provides the tree breeder, incidentally, with the
most important tool of his trade, the F,; hybrid, with which to produce
backcross and Fy» generations. From this point of view the discovery of
these trees is of considerable value, since they can save the geneticist many
years of waiting for artificially induced hybrids of this nature to reach
sexual maturity.

The hybrid vigor so apparent in this hybrid has been observed in two

184 JOURNAL OF THE ARNOLD ARBORETUM _[VvoL. xxx1mr

cases to lead to the forking of the branches at a point between the whorls,
and in one case to the production of a forked cone (PLATE 2). Also appar-
ently associated with this hybrid vigor is a snakelike or looping type of
growth which may be responsible in some degree for the poor form of the
trees. The observed tendency of the lowermost branches of the Hunnewell
hybrids to turn up and develop leaders has also been noted in specimens
of P. monticola < strobus and P. monticola parviflora growing at the
Arnold Arboretum. In each case the tendency appears to be associated
with the other manifestations of hybrid vigor, and unless closely planted
or systematically pruned these trees will inevitably develop multiple and
forked trunks. Such undesirable by-products of the heterotic phenomenon
suggest the need of considerable caution in its direct application to the
problems of forest-tree breeding. Improperly utilized it could quite con-
ceivably discredit considerably the youthful tree-breeding science through
the production of coarse, unmarketable trees, in much the same manner as
the use of exotic species has fallen into disrepute in the United States
largely because of the failure to take into account the need for assessment
of the variability within a species chosen and the necessity of choosing a
variety adapted to the conditions under which it must grow.

Both hybrids #1 and #2 have been propagated by grafting upon young
specimens of P. strobus. Hybrid #1 was first so propagated on a small
scale in 1950 and in 1951 the three surviving grafts of the original five
produced six, eight, and eight female strobili respectively and an unex-
pectedly large number of male flowers. Additional grafts of this plant
were made in 1951. Tree #2 was not discovered until after growth had
started in 1951, and to propagate it without loss of a season soft wood
scions were used upon the new growth of potted white pines. Further
propagation is planned of all five individuals of the Hunnewell pine, as
well as of the P. peuce parviflora.

SUMMARY

A small hybrid swarm of Pinus strobus < parviflora is described and the
type individual designated as & P. hunnewelli. This hybrid is noteworthy
as potential breeding stock for incorporating genes of P. parviflora into
P. strobus. It displays marked heterosis and is promising as an ornamental
because of its attractive foliage. The hybrid is at least partially fertile and
is readily propagated by common grafting techniques. A second hybrid
between P. peuce and P. parviflora is referred to, but the description is
withheld pending confirmation of the female parentage.

LITERATURE CITED

1. ANpDERSON, E. 1936. Hybridization in the American Tradescantias. Ann.
Missouri Bot. Gard. 28: 287-292.

2. Haacen-Smirt, A. J., WANG, T. H., and Mrrov, N. T. 1950. Composition of
gum turpentines of Pinus aristata, P. nie ada P. flexilis, and P. parvt-
flora. Jour. Am. Pharm. Assoc. 39: 254-259.

1952] JOHNSON, SPONTANEOUS WHITE PINE HYBRIDS 185

WwW

na

on

. Jounson, L. P. V. 1939. A descriptive list of natural and artificial inter-

specific hybrids in North American forest-tree genera. Canad. Jour. Res.
17: 411-444.

Mirov, N. T. 1948. The terpenes (in relation to the biology of the genus
Pinus). Ann. Rev. Biochem. 521-54

. RicHEens, R. H. 1945. Forest tree breeding and genetics. Imp. Bur. Pl.
9 pp.

Breeding and ae Cambridge, Imp. For. Bur. Oxford.
RIGHTER, F, I, and DurFiELp, J. W. 1951. Interspecies hybrids in pines.
Jour. Hered. 42: 75-80.

EXPLANATION OF PLATES
PLATE I

Cones, leaves, and seeds of Pinus parviflora Sieb. & Zucc., X P. hunnewelli

hybr. nov., and P. strobus L. Fics. 1, 2, & 3, P. parviflora. Fics. 4, 5, & 6,
x P. hunnewelli, Fics. 7, 8, & 9, P. strobus.

PLaTE II

Abnormalities of cone and branch structure of X Pinus hunnewelli, apparently

associated with the expression of hybrid vigor. Fic. 1, Cone forked near base.
Fic. 2, Branchlet showing forking between nodec.

ARNOLD ARBORETUM

HARVARD UNIVERSITY

Jour. ARNOLD Ars. VoL. XXXIII PLATE I

ON We

JOHNSON, SPONTANEOUS WHITE PINE Hysrips

Jour. ARNOLD Ars. VoL. XXXIII PiaTeE II

" \y a
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‘\ AN a

JOHNSON, SPONTANEOUS WHITE PINE Hysrips

188 JOURNAL OF THE ARNOLD ARBORETUM _ [vot. xxxut

STUDIES IN THE THEACEAE, XXVI
THE GENUS VISNEA *

CLARENCE E. KosBuskKI

THE GENUS Visnea was first described by Linnaeus f. (Suppl. Pl. 36.
1781), and on a later page (251) of the same publication the author
recorded the species V. mocanera. The specific name refers to the mocan
of the Guanches, or ancient inhabitants of the Canary Islands, who made
from the plant a kind of syrup which was much used with their daily
food and in medicines. It is thought that V. mocanera is the plant referred
to as “mocan.’”’ A second generic name, Mocanera, was introduced by
Jussieu (1789), who actually recognized the existence of the previously
described Visnea. However, no serious question of nomenclature ever
evolved, since Mocanera became only a little-used synonym and never
challenged the priority of Visnea. To this day no other species has been
added to the genus.

Originally the species was described from the Canary Islands, and it
has been recorded from most of the group, quite extensively from Gran
Canary, Teneriffe, Gomera, Palma, and Ferro. It was at first thought that
the Canary Islands encompassed the complete range of the species. How-
ever, Johnson (Hooker’s Jour. Bot. 9: 161. 1857) reported the species
as growing in the northwestern portion of the islands of Madeira between
the Ribeira da Janella and the Ribeiro do Inferno. From the works of other
authors one may assume that this small area originally mentioned in
Johnson’s work shows the distribution of the species on Madeira.

It appears that the species is usually found growing in rather wild
and inaccessible places. On Madeira its habit is that of a shrub, while in
the Canary Islands it more often attains the stature of a small tree.

There seem to have been considerable question and also several sugges-
tions regarding the correct position and relationships of the genus.
Endlicher (Gen. Pl. 1018. 1840) first placed it in the Theaceae, but later
in the second supplement of the same publication (Suppl. 2, 81. 1842)
considered it as belonging to the Ebenaceae. Johnson (1857) suggested a
relationship with the genus Clethra and suggested Visnea as “another
link of connection between Ericaceae and Vacciniaceae.” However, by 1885
Johnson appears to have changed his opinion, since he treated the genus
in his Handbook of Madeira under the Theaceae.

In 1859, two years after Johnson’s original article appeared, Schacht
(Denkschr. Bot. Ges. Regensburg 4: 47-60, t. 1, 2. 1859), from material
sent by Johnson, offered a detailed discussion on the status of Visnea.

This genus was studied originally with the oriental genus Anneslea, and a publi-
cation comprising the two genera was planned. However, as the study progressed, it
was decided to treat the two separately.

1952] KOBUSKI, STUDIES IN THE THEACEAE, XXVI 189

This last work was undoubtedly based to a considerable extent on the
previous work of Johnson. No definite conclusions were offered. It is
interesting to note that in the plates, which are very well done as a whole,
the anthers are depicted with almost pore-like openings rather than
longitudinal slits.

Toward the end of the nineteenth century botanists in general accepted
Visnea as belonging to the Theaceae. Szyszylowicz (1895) and Melchior
(1925), in their respective treatments of the family in Die Natiirliche
Pflanzenfamilien, both placed Visnea, without reservation, in the Theaceae.
They placed it in the tribe Ternstroemieae along with Adinandra,
Ternstroemia, and Eurya.

Most botanists, it seems, have interpreted the ovary as inferior or half-
inferior. This interpretation stems from the fact that the persistent calyx-
lobes are joined at the base, and in the development of the fruit this fused
basal portion of the calyx is adnate to the side of the fruit. This appears
very much like an inferior ovary, except that the fruit, when dissected
longitudinally, shows a clean demarkation from the calyx-lobes in the
line and character of the pericarp, which is distinct and not fused with the
calyx.

In the flower, the ovary appears to be placed on the torus formed with
the base of the connate calyx-lobes, but retains its individual identity.
Some authors have described the ovary as projecting “lightly” into the
torus. I could detect no projection into the torus from the dissections
I made and studied. It does appear, however, that in the development of
the fruit and the persistent calyx, the base of the fruit does project
somewhat into the torus, but, as I mentioned above, retains its individuality.

Visnea Linnaeus f., Suppl. Pl. 36. 1781. — Endlicher, Gen. Pl. 1018. 1840;
Suppl. 2, 81. 1842. Webb & Berthelot, Hist. Nat. Iles Canar. 3
(2): 144. 1842.— Choisy in Mém. Soc. Phys. Hist. Nat. Genéve 14:
130 (Mém. Ternstr. 42). 1855.— Johnson in Hookers Jour. Bot. 9:
161. 1857.— Schacht in Denkschr. Bot. Ges. Regensburg 4: 47-60,
t. 1, 2. 1859.— Bentham & Hooker, Gen. Pl. 1: 182. 1862.—
Baillon, Hist. Pl. 4: 257. 1873. — Szyszylowicz in Nat. Pflanzenfam.
III. 6: 190. 1895.— Hubbard in Bailey, Stand. Cyclop. Hort. 6:
3480. 1917. — Engler in Veg. Erde [Pflanzenwelt Afr. 3 (2)] 9: 494.
1921.— Melchior in Nat. Pflanzenfam. ed. 2, 21: 145. 1925.—
Lemée, Dict. Pl. Phan. 6: 877. 193

Mocanera Jussieu, Gen. Pl. 318. 1789.— Lamarck, Encycl. 4: 208. 1797.—
Jussieu in Dict. Sci. Nat. 31: 504. 1824. —- Non Mocanera Blanco, FI. Filip.
446-451, 858. 1837.

Flowers axillary, hermaphroditic. Bracteoles 2. Sepals 5, imbricate,
connate at the base forming a shallow tube adnate to the base of the ovary.
Petals 5, imbricate, connate at the base. Stamens 12 [—21], adhering to
the base of the corolla; filaments free; anthers basifixed, erect. Ovary
3-celled, lightly immersed in the torus; ovules few in each cell, pendant

190 JOURNAL OF THE ARNOLD ARBORETUM _ [VoL. XxXxII

from the apex; styles 3, distinct, persistent. Fruit baccate, indehiscent,
projecting into the torus, nearly enclosed and adjoined near the base by
the persistent calyx, appearing subinferior. Seeds small, pyriform, 3-angled,
the embryo curved, cylindrical, the albumen pulpy.

Evergreen trees with small flowers.

Type species: Visnea mocanera Linn. f.

DistRIBUTION: Canary Islands and Madeira.

Visnea mocanera Linnaeus f., Suppl. Pl. 251. 1781. — Willdenow, Sp.
Pl. 2: 926. 1800.— Bory de St.-Vincent, Essai Iles Fortunées 327,

Iles Canar, 3 (2): 145, t, 69 B. 1842-44.— Choisy in Mem. Soc.
Phys. Hist. Nat. Genéve 14: 130 (Mém. Ternstr. 42). 1855.—
Walpers, Ann. Bot. Syst. 7: 360. 1868. — Johnson, Handb. Madeira
220. 1885. Nicholson, Illustr. Dict. Gard. 4: 185, f. 200, 1886. —
Szyszylowicz in Nat. Pflanzenfam. III. 6: 190. 1895. — Thonner,
Blutenfl. Afr. ¢. 99. 1908; FI. PI. Afr. ¢. 98, 1913. — Pitard & Proust,
Fl. Iles Canar. 134. 1908. — Menezes, Fl. Arch. Madeira 30. 1914. —
Hubbard in Bailey, Stand. Cyclop. Hort. 6: 3480. 1917.— Knoche,
Vagandi Mos (Die Kanarische Ins.) 220, ¢. 17. 1923. — Melchior
in Nat. Pflanzenfam. ed. 2, 21: 145. 1925. — Lindiger, Beitr. Kennt.
Veg. Fl. Kanar. Ins. 278. 1926.

Mocanera canariensis Heynhold, Nom. Bot. Hort. 1: 884. 1840, nom. nud. —
Jaume St.-Hilaire, Expos. Fam. Nat. 2: 371. 1805. — Jussieu in Dict. Sci.
Nat. 31: 505. 1824.

Small tree or shrub, Branches brown or grayish brown, terete, glabrous,
lenticellate, the young branchlets brown, angled, pubescent when very
young. Leaves coriaceous, subelliptic-obovate, 4—7 cm. long, 2—2.5 cm.
wide, glabrous (except when very young), acute at the apex, cuneate at the
base, the margin subrevolute, serrulate along the upper half, a gland
(quickly caducous) to each serration, the veins obscure on both surfaces,
occasionally visible below, the midrib 2-3 (—4) mm. long. Flowers
axillary, solitary or in twos, occasionally in fascicles of three; pedicel
terete, 7-8 mm. long, lightly pubescent at anthesis (lens); bracteoles 2,
ovate or long-deltoid, unequal, 1.25-1.5 mm. long, one bracteole im-
mediately below the calyx, the other disposed along the pedicel away from
the calyx, rarely opposite; calyx-lobes 5, imbricate, coriaceous, persistent,
unequal, glabrous (lightly pubescent at anthesis), 3-5 mm. long, 2.5—2.75
mm. wide, joined at the base for 1.5—2 mm. forming a torus; corolla-lobes
5, imbricate, obtuse, membranaceous, 5—6.5 mm. long, 3.5-4 mm. wide,
joined at the very base; stamens ca. 13 [—21], ca. 4 mm. long, unequal,
the filament ca. 3 mm. long, free, lightly adnate to the base of the corolla,
the anthers long-ovate ca. 1 mm. long, projected into an apicule; ovary
subglobose to conical, lightly imbedded in the torus, ca. 1.5 mm. diameter,

1952] KOBUSKI, STUDIES IN THE THEACEAE, XXVI 191

sulcate and glabrous near the base, densely pubescent above, 3-celled,
the ovules few, the styles 3, filiform, persistent, ca. 3 mm. long, free nearly
to the base, pubescent, the stigmas punctiform. Fruit baccate, indehiscent,
conical, crowned by the persistent styles, during development projecting
more deeply into the torus, the cells often indistinguishable, appearing
one-celled, crowded with pulp, 1—4-seeded, others abortive. [Seeds 3-
angled pyriform, acute, the testa glutinous-granulate].

CANARY ISLANDS: TENERIFFE: above Taganana, in forest, alt. 900 m.,
A, Engler sn. (AA).— “Cruz de Taganana, Cumbre, in rupibus,” alt. 900 m.,
J. Bornmiller 927 (AA), June 14, 1900. — Guinar, river ravine, alt. 700-800 m.,
O. Burchard 62 (AA), Feb. 1904.— Guinar, river ravine, alt. 500 m., J. Born-
miiller 2588 (AA), Sept. 6, 1901.—In woods, C. Bolle sn. (G), in 1851.
PALMA: near Brena Baja, Montagneta, alt. 500 m., J. Bornmiiller 2586 (AA),
May 10, 1901. FERRO: El Golfo, Vueltas above the church, R. T. Lowe H 178
(G), Feb. 18, 1858.—Risco de Jinama, alt. 500-600 m., J. Bornmiiller 2589
(AA), May 17, 1901.

MADEIRA: Ribeiro do Inferno, W. Barbey 908 (G), Dec. 16, 1858.

CULTIVATED: Teneriffe: Oratava, in garden, J. Bornmiiller 925 (AA),
July 1900.— France: Antibes (Alpes-Maritimes), Villa Thuret (AA), Mar.
12 & Apr. 10, 1889.

ARNOLD ARBORETUM,
HARVARD UNIVERSITY.

192 JOURNAL OF THE ARNOLD ARBORETUM _[VvoL. xxxmI

THE GENUS AMENTOTAXUS
Hut-Lin Li

Amentotaxus is a coniferous genus, of isolated position, endemic to
eastern Asia: there are considerable differences of opinion regarding its
proper phylogenetic alliance. The genus has long been considered mono-
typic. Its species, A. argotaenia (Hance) Pilger, was first proposed as a
species of Podocarpus, having been based on sterile material. Later Pilger
transferred it to Cephalotaxus (in Engler, Pflanzenr. IV. 5: 104. 1903),
but it was subsequently referred to a separate genus by him (in Bot. Jahrb.
54: 41. 1916) because of its very distinct long staminate inflorescences.
In Pilger’s system of 1926 (Engler & Prantl, Nat. Pflanzenfam. ed. 2. 13:
267. 1926), Amentotaxus is placed in the Cephalotaxaceae, the only other
genus of which is Cephalotaxus. As Amentotaxus is very different from
Cephalotaxus, Kudo and Yamamoto (in Jour. Soc. Trop. Agr. Formos. 3:
110. 1931) proposed for it the monotypic family Amentotaxaceae. Florin
(Palaeontographica 85, Abt. B: 625-630. 1944; Bot. Gaz. 110: 31-39.
1948), however, is of the opinion that Amentotaxus is not closely related
to the Cephalotaxaceae but rather to the Taxaceae. Accordingly, in the
latest system of classification of the conifers by Janchen (in Sitz. Oest.
Akad. Wiss. Math.-Nat. Kl. Abt. I. 1949(3): 155-162. 1950), it is
placed with Torreya in the tribe Torreyeae of the Taxaceae, while
Cephalotaxus is considered as representing the monotypic family Cephalo-
taxaceae. On the basis of both the vegetative and reproductive structures,
this disposition is probably the most commendable.

The varied opinions regarding its phylogenetic position show that
Amentotaxus is of great morphological interest. Also of considerable
interest is its geographical distribution. First discovered around Hongkong
and in Kwangtung, in southern China, the genus was subsequently reported
to occur in southern Formosa, western Hupeh and Szechuan, southern
Yunnan, and Tonkin. These later records all attribute the plants from
these widely separated localities to a single species. In all cases, the
authors recording these findings have apparently studied only local
material, and a comprehensive review of the genus, with specimens repre-
sentative of all localities, has never been made. Although the number of
specimens from the different localities now available is still rather few
and in some cases inadequate, apparently due to the plants being of rare
occurrence, a study shows that the genus is not monotypic, but is rather
composed of several distinct entities, each possessing distinct morphological
characteristics and an exclusive as well as isolated and restricted range.
In other coniferous genera with similar disjunct ranges, such as Taiwania,
the disjunction is found to be specific in nature. Plants of these remote
locations have long been isolated and have undergone considerable

1952] LI, THE GENUS AMENTOTAXUS 193

differentiation, so that they are recognizable as morphologically and
taxonomically distinct entities, here interpreted as species.

The chief differentiating characters among the species are the size and
shape of leaves and the relative width of the stomatal bands. Apparently
the number of staminate racemes and the size and shape of the mature
seeds are also good taxonomic characters. Unfortunately, as most of the
specimens now available are sterile, these latter important characters are
not revealed in all species. However, the stomatal bands on the under-
surface of the leaves are especially conspicuous and strikingly distinct,
especially in combination with other characters, rendering the differentia-
tion of species very certain even when the locality of a given specimen is
not indicated. The width of the stomatal bands and the proportion of these
to the marginal green bands are very constant among plants of the same
general region, that is, of the same species. While only one collection of
the genus in Yunnan is so far noted, a larger and more complete series of
Formosan specimens is available for study. In all cases, the stomatal
bands are of nearly uniform width among plants of the same general
range, clearly indicating the taxonomic dependability of this character.

As a result of this study, four species are recognized: one, the type
species, occurring in Kwangtung, one in southern Formosa, one in Hupeh-
Szechuan, and one in southern Yunnan and possibly also in northern
Tonkin. These species all occur at medium to high altitudes, varying from
300 to 1600 meters, in ravines and on rocky cliffs, mostly in shady damp
situations or sometimes along borders of streams. They are not only of
very restricted range but also of rare occurrence, probably an indication
that these ancient plants are on the verge of becoming extinct. As the
species are highly ornamental because of their striking foliage, and as they
are also of great botanical interest, it would be desirable to exert more
effort toward bringing them into cultivation.

The material utilized in this study is deposited at the following herbaria,
listed with respective abbreviations used for cited specimens: AA =
Arnold Arboretum; NTU = National Taiwan University, Formosa;
US = U. S. National Herbarium.

I am indebted to the officials of these institutions for permission to
study specimens in their care, and to Dr. A. C. Smith, U. S. National
Museum, for reading the manuscript and offering suggestions.

Amentotaxus Pilger in Bot. Jahrb. 54: 41. 1916

Evergreen shrubs or small trees, glabrous, the branchlets opposite,
terete or more or less quadrangular, patent or ascending. Leaves persistent
toward the upper part of the branchlets, sessile, subsessile, or very shortly
eet opposite, coriaceous, decurrent, Hinear-lanceolate, usually acutish

apex, with a prominent or subprominent costa, green above, with 5
Raa longitudinal bands beneath, the costal and marginal bands pale
or silvery green, alternating with 2 broad whitish stomatal bands. Flowers
dioecious, on the year’s branchlets. Staminate inflorescences of 2-4

194 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxxi

(rarely 1 or 5) spike-like racemes, long and more or less pendulous,
short-pedunculate, in the axils of imbricate bracts, the flowers composed
of fasciculate stamens of peltate or subpeltate short-stipitate scales, the
anther-cells 2-8, ovoid. Ovulate flowers solitary, in the axil of a lateral
bract, on short thick to long slender pedicels, the ovule solitary, erect,
subtended by minute imbricate scales below. Seeds solitary, large, drupe-
like, ovoid or ellipsoid, surrounded by a reddish yellow aril open at the
apex, subtended by several persistent scales at base.

TYPE SPECIES: Amentotaxus argotaenia (Hance) Pilger.

Four species in eastern Asia.

As all previous references pertain to a single species, the synonymy
given below is adjusted on the basis of the specimens cited or locations
attributed. Pilger, the author of the genus, erroneously credited the
species Amentotaxus argotaenia (in Engler & Prantl, Nat. Pflanzenfam.
ed. 2. 13: 268. 1926) to western China only, while actually the a
locality is in southern China. His description, which is rather brief,
based on composite elements from different localities, the Fnriee
characters on the basis of only one collection, Westland from Taimo
Mountains opposite Hongkong, and the ovulate characters on the basis
of a single flower from Wilson 3005 from an unspecified locality in
western China.

KEY TO THE SPECIES

A. Stomatal bands white, as broad as or narrower than the outer green mar-
ginal bands.

B. Leaves comparatively short, 4-7 cm. long, straight, rarely slightly fal-
cate, acute to obtuse at apex; stomatal bands as broad as the green
qiareinal TUN, gas tee ve $a ee hha enews 1. A. argotaenia.

BB. Leaves longer, 6-11 cm. long, mostly falcate, sometimes straight, long-

acuminate at apex; stomatal bands about % as broad as the green mar-

Oia) ONE 65sec vey seien seen csanicse eben 2. A. cathayensis.

AA. Stomatal bands white or brownish, 2 or more times as broad as the outer
green marginal bands.

B. Leaves comparatively long and narrow, 5—8.5 cm. long, 7-9.5 mm. broad,
mostly slightly falcate, the base acute to obtuse; stomatal bands white,
twice as broad as the marginal bands. ............... 3. A. formosana.

BB. Leaves shorter and broader, 3-7 cm. long, 8-11 mm. broad, usually
straight; the base broadly acute to subrounded; a bands brown-
ish or yellowish white, 2.5—-3 times as broad as the green marginal bands.

. yunnanensis.

1. Amentotaxus argotaenia (Hance) Pilger in Bot. Jahrb. 54: 41.
1916, p. p. (quoted plants from Kwangtung and Hongkong).
Podocarpus argotaenia Hance in Jour. Bot. 21: 357. 1883; Masters in Jour.
Linn. Soc. Bot. 26: 547. 1902, p. p.; Dunn & Tutcher in Kew Bull. Misc.
Inf. Add. Ser. 10: 256. 1912.

1952] LI, THE GENUS AMENTOTAXUS 195

Podocarpus insignis Hemsl. in Jour. Bot. 23: 287, 312. 1885.
eta argotaenia ae in Engler, Pflanzenr. IV. 5: 104. 1903; Patschke
n Bot. Jahrb. 48: 629.

Shrub, 2-4 m. high; branchlets spreading or sometimes ascending,
subterete or quadrangular; leaves thickly coriaceous, subsessile to very
shortly petiolate, linear-lanceolate, generally straight, sometimes slightly
falcate toward the tip, 4—7 cm. long 6.5-8.5 mm. broad, acute to obtuse
at apex, cuneate at base, slightly revolute at margins, green above, the
costa prominent and raised above, scarcely raised beneath, i—1.5 mm.
broad, the stomatal bands whitish to silvery whitish, 1.5-1.8 mm. broad,
about as broad as the marginal bands, the marginal bands pale silvery
green, 1.3—2 mm. broad; petioles thick, scarcely 1-1.5 mm. long; staminate
racemes subterminal or lateral, solitary or 2 or 3 together, in the axils of
minute scales, short-pedunculate, more or less pendulous, 5—6.5 cm. long,
the flowers composed of subfasciculate peltate or subpeltate stamens, the
filaments short, the anther-cells 2—5, mostly 3, ovoid; ovulate flowers and
mature seeds unknown

Kwangtung Province at Lofaushan Mountains, and also around Hong-
kong, in wet rocky situations or along edge of streams, at altitudes of
600-1500 meters.

CHINA: Kwangtung: Lofaushan (type locality), Levine & McClure 704
(US), C. O. Levine 1502 (AA, US). Hongkong: Lantoa Island, Hongkong
Herb. 2068 (AA).

The specimens examined are all sterile. The only staminate specimen
known is the type of Podocarpus insignis, Westland s. n. from Taimo
Mountains opposite Hongkong, the basis of descriptions by both Hemsley
and Pilger. The type of Podocarpus argotaenia Hance is a sterile specimen
of E. Faber, Sept. 1882, from Lofaushan, Kwangtung, in the British
Museum (Natural History).

2. Amentotaxus cathayensis sp. nov.

Cephalotaxus argotaenia Pilger in Engler, Pflanzenr. IV. be 104. 1903, p. p.;
Rehder & Wilson in Sarg. Pl. Wils. 2: 6. 1916 (excl. sy

Amentotaxus argotaenia Pilger in Bot. Jahrb. 54: 41. 1916, p. p. a Lai
collected by Wilson), in Engler & Prantl, Nat. Pflanzenfam. ed. 2. 270.
1926.

Podocarpus argotaenia sensu Masters in Jour. Bot. 41: 269. 1903, in Jour.
Linn. Soc. Bot. 37: 414. 1906; non Hance

Frutex vel arbuscula 2—5 m. alta; ramulis patentibus, teretibus vel
quadrangularibus, subcrassis, internodiis ad 1.5 cm. longis; foliis breviter
petiolatis, longe lineari-lanceolatis, gracilibus, distincte falcatis, interdum
rectis, 6-11 cm. longis, 6-8 mm. latis, superne gradatim attenuatis, apice
calloso-acuminatis, basi attenuatis, margine leviter revolutis, costa supra
leviter elevata subtus vix elevata circiter 1.5 mm. lata, striis stomatiferis
albidis circiter 1.2 mm. latis, latitudine circiter 2/3 striarum marginalium

196 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxxu1

aequalibus, striis marginalibus pallidis a minusve argenteo-viridibus,
circiter 2 mm. latis; petiolis crassis 2-3 mm. longis; floribus masculis
ignotis; floribus ovulatis (fide Pilger) in axilla bracteae ad basim ramuli
foliati enatis, breviter crassiuscule pedicellatis, pedicellis apice squamis
imbricatis instructis, ovulo solitario; seminibus maturis ignotis.

Western Hupeh and western Szechuan (?), cliffs of ravines, at altitudes
of 300-1100 meters, rare.

CHINA: Western China: No pene ed (presumably western
Szechuan), E. H. Wilson 3005 (AA, TYPE). Wu
9209 (AA, US). Hupeh: Hsing-shan Lt E. H. Wilson 2107 (AA, US).

Only sterile specimens are available. Rehder and Wilson originally
indicated Wilson 3005 as an ovulate specimen, but the available one does
not bear any flowers. In Pilger’s original description of the genus, the
ovulate flower is described on the basis of a single flower from Wilson 3005,
received through Rehder. In vegetative characters alone, these western
Chinese specimens clearly represent a species distinct from those of other
regions.

A sterile specimen from Dupha Hills, J. L. Lister in 1874 (AA), has
long narrow falcate leaves very similar to those of A. cathayensis, but with
slightly broader stomatal bands, which are as wide as the marginal bands.
This specimen may indicate a more western extension of the range of the
species.

3. Amentotaxus formosana sp. nov.

Podocarpus argotaenia sensu Henry in Trans. Asiat. Soc. Jap. 24. Suppl: 91
1896 (List Pl. Formos.); Masters in Jour. Linn. Soc. Bot. 26: 547. 1902,
p. p.; Matsum. & Hayata in Jour. Coll. Sci. Tokyo 22: 399. 1906 (Enum.
Pl. Formos.) ; non Hance.

Cephalotaxus argotaenia sensu Forbes & Hemsl. in Jour. Linn. Soc. Bot. 26:
547. 1907, p. p., non Pilger.

Amentotaxus argotaenia sensu Yamamoto in Bot. Mag. Tokyo 40: 453. 1926,
Suppl. Icon. Pl. Formos. 3: 1, t. 1. 1927, op. cit. 5: 7, f. 1-2. 1932, in Jour.
Jap. Bot. 8: (64). f. 1-6. 1932; Kanehira in Trans. Nat. Hist. Soc. Formos.
84: 80. 1926, Formos. Trees rev. ed. 33, f. 2. 1936; Kudo in Jour. Soc. Trop.
Agr. Formos. 3: 110. 1931; Chen, Ill. Man. Chin. For. Trees & Shrubs 13,
f. 2. 1937; non Pilger.

Arbor vel arbuscula vel frutex ad 10 m. altus, sparse ramosus; ramulis
oppositis, patentibus, subteretibus vel subquadrangularibus, internodiis
6-12 mm. longis; foliis subsessilibus vel breviter et crasse petiolatis,
lanceolatis, leviter sed distincte falcatis, raro rectis, 5—-8.5 cm. longis,
7-9.5 mm. latis, superne gradatim attenuatis, apice calloso-acuminatis,
basi acutis vel obtusis, margine revolutis, supra atro-viridibus, costa
supra subplana subtus leviter elevata circiter 1.5 mm. lata, striis
stomatiferis albidis circiter 2 mm. latis, quam striis marginalibus duplo
latioribus, striis marginalibus pallide argenteo-viridibus, circiter 1 mm.
latis; petiolis nullis vel crassis vix 1 mm. longis; racemis masculis ad

1952] LI, THE GENUS AMENTOTAXUS 197

apicem ramulorum plerumque 3- vel 4-fasciculatis, raro 5—fasciculatis vel
solitariis, gracilibus, circiter 3 cm. longis, breviter pedunculatis, basi
squamis coriaceis 2—4-seriatim imbricato-obtectis, squamis plerumque 7,
carinatis, superioribus gradatim majoribus, oblongo-lanceolatis, ad 15 mm.
longis et 5 mm. latis, basalibus ovatis ad 5 mm. longis et 3-4 mm. latis,
pedunculis circiter 5 mm. longis; floribus masculis subsessilibus, antheris
subpeltatis brevistipitatis fasciculatis 1.5 mm. crassis, loculis plerumque
5-8 pendulis circiter 0.7 mm. longis et 0.5 mm. latis, filamentis brevibus vix
1 mm. longis; floribus ovulatis solitariis lateralibus, subglobosis, circiter
3 mm. crassis, longe pedicellatis, basi squamis circiter 10 minutis opposite
imbricato-obtectis, squamis 5-seriatis, carinatis, subaequalibus, circiter
3.5 mm. longis vel latis; ovulo 2 mm. longo et 1.5 mm. crasso, disco parvo
cupuliformi carnoso, 2.5 mm. diametro et 1.5 mm. longo; semine maturo
solitario axillari, longe pedicellato, oblongo-ellipsoideo, 20-25 mm. longo,
9-11 mm. crasso, apice mucronulato-rostrato, basi squamis minutis
persistentibus imbricatis instructo, arillo rubro-luteo, demum §atro-
purpureo; pedicello gracili, 1.5—2 cm. longo.

Southern Formosa (Taiwan), in broad-leaved forests in shady places,
near ravines and cliffs, at altitudes of 700-1300 meters, rather scarce.

CHINA: Taiwan: rene Daibu, Taririku, R. Kanehira, May 11, 1924
(AA), R. Kanehira & S. Sasaki, Feb. 19, 1925 (AA), S. Soe Feb. 25, 1925
(NTU, tyre), R. Kanehira, Dee 27, 1925 (AA); Southern Koshun, R. Kanehira
in 1924 (AA).

The Formosan plant is better known than plants from the mainland,
although it is also more or less rare. It has been described in detail by
Yamamoto and others and illustrated by numerous photographs and
drawings. The source of the figure given by Chen is not indicated, but it
is an exact copy of Kanehira’s figure. Chen’s generic description also fits
only the Formosan plant.

4. Amentotaxus yunnanensis sp. nov.
Amentotaxus argotaenia sensu Hu in Bull. Chin. Bot. Soc. 1(1): 8. 1935, non
Pilger.

Arbuscula, ramis crassis, ramulis oppositis, erecto-adscendentibus,
gracilibus, teretibus vel subquadrangularibus, internodiis ad 1.3 cm. longis;
foliis subsessilibus, linearibus, rectis, raro apicem versus leviter falcatis,
3-7 cm. longis, 8-11 mm. latis, apice obtusis vel acuminatis, basi late
acutis vel subrotundatis, margine leviter revolutis, supra viridibus vel
pallide viridibus, costa supra prominula elevata subtus subplana vix elevata
circiter 1.5 mm. lata, striis stomatiferis leviter fuscis vel luteo-albidis
circiter 2.5 mm. latis, quam striis marginalibus 2.5—3-plo latioribus, striis
marginalibus pallide argenteo-viridibus ad 1 mm. latis; floribus masculis
ovulatisque ignotis; semine maturo pedicellato, ovoideo, circiter 2.2 cm.
longo et 1.5 cm. crasso, apice leviter rostrato, arillo rubro-luteo, basi
squamis coriaceis 3-seriatim imbricatis instructo, squamis oppositis, circiter
12, distincte carinatis, superioribus gradatim majoribus ad 4 mm. longis et

198 JOURNAL OF THE ARNOLD ARBORETUM _ [vot. xxxur

5 mm. latis, basalibus ovatis ad 2 mm. longis et latis; pedicello crasso,
circiter 1.4 mm. longo, in sicco plano.

Southern Yunnan, a moss-clad plant on rocky hill at an altitude of 1600
meters.

CHINA: Yunnan: Makwan, H. T. Tsai 51887, March 2, 1932 (US, Type).

Merrill (in Jour. Arnold Arb. 19: 21. 1938) records Amentotaxus
argotaenia from Chapa, Tonkin, basing the record on Pételot 3897. I
have not seen specimens of this collection, but judging from the location
the plant is probably referable to the Yunnan species, although it may
possibly represent A. argotaenia, definitely known only from Kwangtung
and Hongkong.

DEPARTMENT OF BOTANy,
U.S. NATIONAL Museum,
SMITHSONIAN INSTITUTION,
WASHINGTON, D. C.

JOURNAL

OF THE

ARNOLD ARBORETUM

VoL. XXXIII JULY 1952 NUMBER 3

WILLIAM JACK’S GENERA AND SPECIES OF MALAYSIAN PLANTS
E. D. MERRILL
With one plate

THIS PAPER was not planned to consider other than certain phases of
Jack’s botanical work. Hooker published an excellent biographical sketch
of William Jack prefixed to one of the series of papers in which the Jack
plant descriptions of 1820-22 were republished, to which the reader is
referred. Supplementing these data are the remarkably interesting letters
written by Jack to Nathaniel Wallich from Penang, Jan. 14 to May 19,
1819, from Singapore, June 8 to June 18, 1819, and from Pencocten,
Sumatra, Aug. 19, 1819, to Oct. 26, 1821.2 These letters contain a wealth
of information regarding Jack’s experiences and observations as a pioneer
botanist operating in the then botanically unknown and very rich forests
of Penang, Singapore, and Sumatra. For Jack was indeed the pioneer post-
Linnean Malaysian botanist, his work antedating the investigations of
Blume at Buitenzorg, Java, which were initiated, as to publication, in 1823,
a year after Jack’s death. On February 12, 1819, he wrote to Wallich, “I
am overwhelmed with the treasures that pour in upon me; I have been em-
ployed night and day so as not even to leave time for correspondence. I
actually wish for a little remission, as my cough has been teasing me, but
how is it possible! I cannot even now get through all; my specimens are
in piles that are quite alarming and I have not time to look over them. I
must however take a day or two to make a selection for you.” An excellent
summary of the essential data regarding William Jack and his botanical
accomplishments has been included by Mrs. M. J. van Steenis-Kruseman
in her recent publication.*

1 Hooker, W. J. Description of Malayan Plants. By William Jack. With a ee
Memoir of the Author and Extracts from his Correspondence. Comp. Bot. Mag
121-147. 1835.

2 Burkill, I. H. William Jack’s Letters to Nathaniel Wallich, 1819-1821. Jour.
Straits Br. Roy. As. Soc. 73: 148-268. 1916.

® Van Steenis-Kruseman, M. J. Malaysian Plant Collectors and Collections,
Fl. Malesiana I, 1: 256-257. 1950.

200 JOURNAL OF THE ARNOLD ARBORETUM _ [vot. xxxim

Briefly, William Jack was born in Aberdeen, Scotland, Jan. 29, 1795,
and died at Bencoolen, Sumatra, Sept. 15, 1822, of pulmonary tuberculosis,
apparently complicated by malaria contracted on a trip to Moco-moco.
His physical condition was so serious that he had been placed aboard a
ship bound for England, the actual sailing of which was delayed by adverse
weather conditions; but failing very rapidly he was removed to Govern-
ment House, Bencoolen, where the end came. The entry in Pritzel’s
Thesaurus that he died on shipboard near the Cape of Good Hope is
erroneous. He was a very precocious student, excelling in languages and
developing an interest in botany at an early age. Receiving his M.A.
degree from Aberdeen University at the age of sixteen, he then studied
for the M.D. degree, finishing his medical training in London where he
was admitted as a Fellow of the College of Surgeons at the end of January,
1812. Having received an appointment for service in India he left England
in January, 1813. His services with the British East India Company were
in the medical field. It was while actively engaged in the Nepalese war
that he commenced to correspond with Nathaniel Wallich in Calcutta, and
it was in this campaign that he unfortunately contracted pulmonary
tuberculosis, which a few years later was to terminate what promised to be
a most brilliant botanical career. William Jack was unquestionably one of
the most able botanists ever to become associated with the tremendously
rich and the then very little known flora of the Malay Peninsula and
Archipelago.

In November 1818, having been strongly recommended by Nathaniel
Wallich to Sir Stamford Raffles, he was appointed by the latter to serve
as botanist on his staff in an attempt to rehabilitate the British East India
Company’s controlled areas on the west coast of Sumatra, where British
influence had long been dominant. What he was able to accomplish in less
than three years was most remarkable. His publishing activities, com-
mencing in the remote settlement of Bencoolen in 1820, terminated there
in 1822, the year of his untimely death, and finally ceased a year later
with the posthumous appearance of the three papers he had prepared in
Bencoolen and sent to the Linnean Society in London. How well he
developed his knowledge of those parts of Malaysia which he personally
explored is manifest from his published papers. Had not his herbarium,
his manuscript descriptions and notes, and his drawings of many species
been destroyed in the burning of the ‘‘Fame” at the time when the British
East India Company relinquished control of its Sumatra holdings in
1824 to concentrate on the building up of Singapore, Jack’s name would
undoubtedly have been written much larger in the annals of Malaysian
botany than is now the case. As it is, no botanist who has concentrated on
a study of the flora of the region has accomplished so much of lasting
value in such a limited time as did William Jack. And what he published
is of very high order indeed. His usually ample descriptions, as contrasted
with the very short diagnostic data provided by Blume in his early work,
as Griffith noted in 1843, are actually autographs of plants. To write a
technical description is a simple matter, but to include in not overlong

1952] MERRILL, WILLIAM JACK’S GENERA AND SPECIES 201

descriptions, as Jack did, all or most of the essential data needed by a
later monographer to place a species in association with those described
by other authors is an art in which Jack excelled. Thus it is that in such
large and critical genera as Antidesma and Ilex, I do not hesitate in
reducing, from Jack’s descriptions, associated with an examination of
Sumatra specimens collected by others, Antidesma frutescens Jack (1822)
to A. ghaesembilla (Linn.) Gaertn., and Octas spicata Jack (1822)
to Ilex spicata Blume (1826), although up to this date European
monographers have retained the former as a valid species allied to A.
ghaesembilla, while although Octas Jack has been correctly reduced to
Ilex Linn., no author has even hinted that Jack’s species is identical with
that of Blume. In the very much larger and exceedingly critical genus
Ficus Linn. I do not hesitate in replacing F. diversifolia Blume (1825)
by the earlier F. deltoidea Jack (1822), and F. glaberrima Blume by
the earlier F. rigida Jack (1822), although no extant Jack types are
known.

DEVELOPMENT OF INTEREST IN JACK PROBLEMS

In my somewhat more than two decades of residence in the Philippines,
I had access to only a part of the reprinted William Jack plant descriptions.
In building up the botanical library in Manila, starting with nothing in
the way of books, I was never able to acquire a copy of the Companion
to the Botanical Magazine containing many of the republished Jack
descriptions. It was only toward the end of my Philippine career that I
discovered the 1887 Triibner reprint of the Jack papers and acquired a
copy of it. This experience stimulated my interest in problems appertaining
to the Jack species when, in later years, I did have access to certain
records never available in Manila. Perhaps the chief reason for the
preparation of this index to the Jack species was my own difficulty
encountered in earlier years in locating various descriptions needed for
reference — and difficulties continue to face all systematists who seek
original or reprinted Jack descriptions, except those located in a very few
favored centers. This applies particularly to those taxa characterized in
the short-lived and never generally available Malayan Miscellanies
published in Bencoolen, Sumatra, 1820-22. Indices are lacking in the
original Jack papers and in the Hooker reproductions of 1830-36, the
latter still being the most generally available source of the Jack descrip-
tions. In standard works of the Index Kewensis type references included
are for the most part only to the original places of publication of new
names, not to subsequent republications of descriptions.

The original Jack Malayan, Miscellanies papers are exceedingly rare
and are to be found in only a very few of the older botanical libraries.
I know of only two complete sets, one at Kew and one at Calcutta, and
while these are complete for the regular Malayan Miscellanies papers, the
Kew set lacks the “Appendix” of 1820. Two of the three Jack papers
were in the Linnean Society Library, but one of these was unfortunately

202 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxur

lost. I judge that Blume must have been familiar with at least a part of
the Malayan Miscellanies botanical papers when he initiated his botanical
work at Buitenzorg, Java, in 1823. Perhaps Jack took with him to
Buitenzorg copies of the parts then issued when he visited Java in 1821;
see Burkill, Jour. Straits Br. Roy. As. Soc. 73: 198. 1916, footnote 188.
However, no copies of the original Jack papers exist today in either the
Buitenzorg or the Leiden libraries.

Where library facilities are limited, as was the case in Manila — and is
the case in a great many institutions established within the present
century, or for that matter within the past century or more — it occurred
to me that an index to all the Jack taxa (1820-23), with references to all
the reprinted Jack descriptions (1830-36, 1843, 1877), might serve a
useful purpose. The preparation of a mere list would have been a simple
matter, but the decision to inquire into the status of each name in relation
to those proposed by other authors made the task a more complicated and
time-consuming one. Certain data were compiled in 1950, including
basic lists. The rough draft of the index proper was written at El Zamorano,
Honduras, in February and March, 1951, data therein rechecked in Boston
later in the year, and the introductory matter was mostly drafted on the
S.S. “Mauretania” en route from New York to Southampton in June, and
finished in London in July, 1951.

The Jack papers were so significant at the time they were originally
printed that their appearance created a great deal of interest in the work
of that young botanist. The very fact that the technical plant descriptions
of 1820-22 have since been reprinted three times (or four times if we
consider the Griffith separately paged reprint of 1843 to constitute a
distinct publication) speaks for itself. I do not know of a parallel case
in the literature of systematic botany. Jack was the first post-Linnaean
botanist to work in the field on the exceedingly rich Malaysian flora. In
his time, from a botanical standpoint, “all the world was new” in Penang,
in Singapore, and in Sumatra. In the period 1819-22 one may safely
estimate that out of every hundred plant species that Jack actually
observed at least seventy-five were unnamed and undescribed, and in
Jack’s time many of the widely distributed Malaysian genera had not
been named or characterized.

JACK’S ORIGINAL PUBLICATIONS 1820-23

What concerned and still concerns most botanists are the Jack descrip-
tions of 1820—22.4 These were the papers published in remote Bencoolen,
of which few copies were distributed at the time of publication and of
which the undistributed stock was destroyed in the burning of the
“Fame” in 1824. No copy of these papers is to be found in any American
library. For a microfilm of the Kew copy I am indebted to Mr. H. S.
Marshall, Librarian, Royal Botanic Gardens, Kew. In addition to these

‘Jack, W. Descriptions of Malayan Plants. Mal. Miscel. 1 (1): 1-27. 1820;
1 (5): 1-49. 1821; 2 (7): i-iii. 1-96. 1822. Sumatran Mission Press, Bencoolen.

1952] MERRILL, WILLIAM JACK’S GENERA AND SPECIES 203

three papers another was printed but never published which has caused
some confusion and misunderstanding. This® was actually printed in
1820, not in 1823 as Hooker surmised. Its purpose was to assemble
certain descriptive data in form for easy reference and to provide
Nathaniel Wallich in Calcutta with a copy for his criticism of various
proposed new taxa. Actual publication of the document was neither
planned nor consummated. Regarding it Hooker, Comp. Bot. Mag. 1:
259. 1836, states:

“In point of interest, the ‘Third’ Memoir, as it is called, of Mr. Jack,
far exceeds the previous ones... . I have reason to think that the present
Memoir is very little known in this country, as I have never seen it
quoted, nor met with any copy but that which has been kindly lent to
me by the mother of the lamented author. This number of the Malayan
Miscellaney is without date, and only bears the title ‘Appendix, De-
scriptions of Malayan Plants, by William Jack, No.

The only known extant copies of this dueacd appear to be the one
sent by Jack to Wallich and now preserved in the library of the Calcutta
Botanic Garden, and a copy in the library of the Asiatic Society of Bengal,
presented by Major General Hardwick, July 14, 1821. For bibliographic
data regarding this item I am indebted to Dr. K. Biswas, Director of the
Calcutta Botanic Garden. The included taxa, some of which were
published elsewhere by Jack himself and by other authors, date from
the time of effective publication by Jack himself, G. Don (one case),
Griffith (one case), and Hooker. The latter thought, with expressed
doubt, that this Appendix was printed in 1823. But Jack died in 1822,
and by the time of his death publication of the Malayan Miscellanies
had ceased. As Burkill notes, Jour. Straits Br. Roy. As. Soc. 73: 227.
1916, footnote 273, on the basis of evidence supplied by Jack and by
Raffles, the document was printed in 1820, and we now know that a copy
was in the library of the Asiatic Society of Bengal as early as July 14,
1821. Some of the more or less confused references to it which one notes
in botanical literature are: “Mal. Misc. iii. [1823?],” “Descr. Mal. PI.
iii, 12 [1823],” “Mal. Misc. App. Ined. [1823] 21,” etc.

Yet at least one other botanist in Great Britain must have had access
to the document. G. Don probably saw Mrs. Jack’s copy, as he actually
published the technical descriptions of Stagmaria Jack and S. verniciflua
Jack in 1832, four years prior to Hooker’s similar action. Wallich, for
whom, in part, the document was originally printed, as noted above, had
a copy in Calcutta, and Griffith had access to it in 1843.

If referred to at all this Jack “Appendix” should be cited as “ined.”
In this paper I have included references to it in square brackets, thus:
“TApp. Descr. Mal. Pl. . . . 1820]” followed by a reference to the later
validating description in each case. Unfortunately Hooker in reprinting
the Jack descriptions gave no bibliographic references to individual
species.

Jack, W. Appendix. reba of Malayan Plants. No. 3. p. 1-26. [1820].
Sumatran Mission Press, Bencoolen

204 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxim

Jack’s other botanical contributions were limited to the three papers
published in London in 1823,® the year following his untimely death.
The plant descriptions included in these papers have been generally avail-
able to botanists who have at times needed to consult them. They were
reprinted only by Griffith in 1843

Certain other manuscript descriptions were sent by Jack to Nathaniel
Wallich in Calcutta, to whom discretionary powers were extended. Some
of these appear in volume two of the Carey and Wallich edition of
Roxburgh’s Flora Indica (1824), and one, Pittosporum serrulatum Jack,
was first published by Griffith in 1843. The Jack Melastoma and
Cyrtandraceae papers were summarized in Oken Isis 22: 1036-1039,
1176-1181. 1829.

THE HOOKER REPRINTS, 1830-36

These, initiated in 1830 and completed in 1836, include the genera and
species originally published by Jack in the Malayan Miscellanies papers,
plus most of those contained in the unpublished Appendix discussed above.‘
It has been in this series that the Jack descriptions of 1820—22 have been,
and still are, most accessible to botanists, in spite of the later reprintings
of 1843 and 1887. Because these reprinted Jack descriptions are scattered
through four unindexed volumes of three different serials issued over a
period of seven years, one must often search for an individual description
when needed. One of the reasons why Griffith again reprinted the Jack
descriptions in 1843 was the difficulty encountered both in securing
access to sets of the Hooker periodicals and in locating individual Jack
descriptions when needed. Griffith had in mind the need of individuals
located as he was, remote from the botanical centers of Europe. It is of
course to Hooker’s distinct credit that he did rescue the contents of these
— rare Jack papers from practical oblivion.

e learn from Hooker’s own statement, Comp. Bot. Mag. 1: 122.
me that it was Nathaniel Wallich who suggested to him the desirability
of reprinting the Jack papers, and it was for this purpose that Wallich
provided Hooker with the set of the Malayan Miscellanies now on the
library shelves of the Royal Botanic Gardens at Kew. Wallich, of course,
knew the fate of Jack’s herbarium and the undistributed stock of the
Malayan Miscellanies, and he knew that because of the burning of the
“Fame,” Feb. 4, 1824, it would be impossible for future botanists to
acquire copies of this Bencoolen serial. For on the “Fame” were the
Bencoolen records, Jack’s herbarium, manuscript descriptions, notes, draw-
ings, the extensive natural history collections assembled by Sir Stamford
Raffles, and finally, all the reserve stock of the Malayan Miscellanies.

° Jack, W. On the Malayan Species of Melastoma. Trans. Linn. Soc. 14: 1-22.
pl. 1. 1823; On the Cyrtandraceae, a New Natural Order of Plants. Op. cit. 23-45.
pl. 2. 1823; vate of Lansium and Some Other Genera of Malayan Plants. Op. cit.
114-130. pl. 4. 18

"Hooker, W. “7 Descriptions of Malayan _— by William Jack. Hook. Bot.
Misc. 1: 273-290. 1830; 2: 60-89. 1830; Hook. Jour. Bot. 1: 358-380. 1834; Comp.
Bot. Mag. 1: 121-157. 1835; 219-224, 253-272. 1336.

1952] MERRILL, WILLIAM JACK’S GENERA AND SPECIES 205

THE GRIFFITH REPRINTS, 1843

William Griffith, inspired by much the same reasons that in 1830
induced Hooker to commence the reprinting of the original Malayan
Miscellanies descriptions of Jack, republished all of the Jack descriptions
known to him in a series of three papers in the Calcutta Journal of
Natural History in 1843.8 He noted the rarity of the Jack papers published
in 1820-22 and commented on the very inconvenient subdivision of the
parts as reprinted by Hooker.

This Griffith series includes not only those descriptions reprinted by
Hooker (1830-36) from the Malayan Miscellanies, but also all of those
included in the three Jack papers published in London in 1823, most
of those which -had been published by Wallich under Jack’s name in
volume two of the Carey and Wallich edition of Roxburgh’s Flora Indica
(1824), and even one (Pittosporum serrulatum Jack) which appears
only in the Griffith document. Occasionally one finds complete or partial
copies of these Griffith papers with the original Calcutta Journal of Natural
History pagination, as in the case of the libraries of the British Museum
(Natural History) and the Rijksherbarium at Leiden.

THE GRIFFITH SEPARATELY PAGED REPRINT OF 1843

Immediately following the issue of the Jack papers in the Calcutta
Journal of Natural History in 1843, Griffith reprinted them in the form
of a separately paged volume under the same title as that used in the
Journal itself. The pagination is 1-230, i-iii. In the two copies of this
work that I have seen (Linnean Society Library and the Lindley Library,
Royal Horticultural Society) the text covers the descriptions of three
plates, but the plates themselves are missing. There is also a copy of this
reprint in the library of the British Museum (Bloomsbury). Because of
an irregularity in including on pages 135-160 of the volume the data
published in Griffith's own paper on some remarkable plants in the
Calcutta Garden (Calc. Jour. Nat. Hist. 4: 231-256. 1843) it is suspected
that very few copies of this reprint were distributed. The essential data
regarding this separately paged reprint were kindly supplied by Mr. I. H.
Burkill, and I later examined the copies mentioned above.

Of this pages 1-62 are identical with the first paper in the Calcutta
Journal series. On pages 63 to 77 certain adjustments in page contents
are made, but there are no changes in the text. Pages 135 to 160 include
the Griffith paper above mentioned. Then the rest of the Jack descriptions
appear on pages 161 to 227. Pages i-iii consist of an index by families and
genera.

Had this Griffith Calcutta reprint of the Jack descriptions been generally
available, aegis any further consideration of them would have been

® Griffith, W. Descriptions of Malayan Plants. By William Jack. Arranged Ac-
cording to ae Natural Families, etc. Calc. Jour. Nat. Hist. 4: 1-62: 159-231; 305-
347. pl. 14-16. 1843.

206 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxx1I

unnecessary. But sets of the Calcutta Journal of Natural History are
lacking in very many, perhaps most botanical libraries, and the separately
paged reprint was apparently suppressed; at any rate it was never widely
distributed.

THE TRUBNER REPRINT OF 1887

The Triibner Oriental Series consists of four volumes, a first series of
two volumes, 1886, and a second series, also of two volumes, 1887. A
total of fifty-one papers were reprinted from various sources, covering
important contributions to our knowledge of the botany, zoology, geology,
exploration, history, philology, linguistics, anthropology, inscriptions, cli-
mate, minerals, and other subjects appertaining chiefly to the Malay
Peninsula and Archipelago. Many of the original papers appeared in
serial literature that is sometimes not generally available. I judge that
these reissued papers are not well known to botanists and botanical
bibliographers, for this 1887 reprint of the important Jack papers escaped
the attention of Rehder when he compiled the remarkably complete
Bradley Bibliography, published from 1911 to 1918, covering the literature
of the world appertaining to woody plants, appearing before the end of the
nineteenth century. The Triibner® series is well worth a place on the
shelves of all special research libraries devoted to the subjects above
mentioned. The main title is a somewhat unfortunate one as it appears on
the first series, but it was emended in the second series by the addition
of the phrase, following Indo-China “and the Indian Archipelago.” As
a matter of fact, in the republished papers there are very few which
in any way appertain to Indo-China, most of them applying strictly to
the Malay Peninsula and Archipelago. The initiative behind the selec-
tion and republication of this distinctly important series of 51 technical
papers came from the officers and council of the Straits Branch, Royal
Asiatic Society, Singapore.

It is in volume two of the second series that one can now gain the easiest
access to the early Jack descriptions, for the technical names of all species
are included in the index to the volume. This paper occupies pp. 209-295
of volume two of the second series, 1887. Appended to it and occupying
pages 296 to 302 are various botanical references giving the then accepted
names for many of the Jack species prepared by Sir J. D. Hooker, with
many philological notes on the significance of the Malay names listed
by Jack, these prepared by D. F. A. Hervey.

JACK’S HERBARIUM

It is known from the published records that Jack industriously increased
his herbarium wherever he had an opportunity of botanizing, but chiefly
in Penang, Singapore, and on the west coast of Sumatra and its neighbor-

® TRUBNER’s ORIENTAL Series. Miscellaneous Papers Relating to Indo- — Re-

printed for the Straits Branch of the Royal Asiatic Society. 1: i-xii. 1-318; 2: 1-309.
1886. Second Series. 1: 1-viii. 1-307; 2: 1-313. 1887. Triibner & Company. London.

1952] MERRILL, WILLIAM JACK’S GENERA AND SPECIES 207

ing islands. It must have been a large collection, but there are no extant
records as to its actual size. The Jack holotypes were destroyed with all
of his undistributed duplicates, and all of his unstudied or partly
studied material when the “Fame” burned just after sailing from Bencoolen
Feb. 4, 1824. Such Jack material as now exists represents duplicates of
his Penang and Singapore collections which he sent to Wallich and which
were numbered in the Wallich List (‘Catalogue’). There are certain
Jack Sumatra specimens in the Delessert herbarium at Geneva, and others
at the Rijksherbarium, Leiden, but the total number is apparently not
great. Among those at Geneva are specimens representing Aeschynanthus
radicans Jack, Connarus semidecandrus Jack, C. villosus Jack, Cyrtandra
hirsuta Jack, C. macrophylla Jack, Didymocarpus corniculata Jack, and
Melastoma obvelutum Jack.

There are also certain Jack specimens in the herbarium of the British
Museum (Natural History) other than those in the Wallich distribution.
These seem to be entirely duplicates of his Penang and Singapore collec-
tions sent by Jack directly to Robert Brown. No extant list is known. On
March 7, 1819, writing from Penang, Jack notified Brown that accompany-
ing his letter he would receive the box of specimens mentioned in an earlier
letter. The highest number located is “59” for a specimen of a Trichomanes
from Penang. Later Jack sent at least four Nepenthes specimens from
Singapore, where he arrived from Penang, May 31, 1819. The labels on
these fugitive Jack collections are in his handwriting, but on at least some
of them somebody at the Museum later wrote the name “Wallich” as the
collector, which doubtless explains why many of these specimens were not
previously recognized as being actual Jack duplicates.

At Leiden, in 1950, without making an intensive search, I located
Ternstroemia serrata Jack, T. rubiginosa Jack, Salacia . . . Jack (an un-
published binomial), and Lasianthus attenuatus Jack, and Dr. Hoogland
located Tetracera arborescens Jack. These Jack Sumatra specimens bear
his original labels. They are indicated as having been received in 1829, and
they later reached the Rijksherbarium via the Hasskarl private herbarium.
It is suspected that these specimens may have represented a small lot that
was perhaps left behind when the British left Bencoolen in 1824, which was
probably retrieved by some Dutch official and transmitted by him to Hol-
land, where the specimens came into the possession of Hasskarl.

It is evident that Jack sent certain Sumatran material home, but it is
not known what became of some of these collections. Thus in a Jack letter
to Wallich dated at Bencoolen September 9, 1820, there is a very amusing
account of a collection of botanical specimens he selected at the request
of a certain Marchioness for the Edinburgh Museum.” In this letter,
inter alia, he says: ‘““My best specimens are all gone home as you know.”
Mr. Burkill’s attempts to locate the Jack specimens of this particular send-
ing failed, nor in the relatively little time I could spend in the Edinburgh
herbarium in July, 1951, did I succeed in locating any of these fugitive

* Burkill, I. H. Jour. Straits Br. Roy. As. Soc. 73: 215. 1916.

208 JOURNAL OF THE ARNOLD ARBORETUM | [voL. xxxur

specimens. Late in 1951 Mr. Burtt found one specimen, of which Dr. J. M.
Cowan kindly supplied a photograph. Jack had mentioned the poor quality
of the specimens he selected for the Edinburgh Museum, and the type
collection of Didymocarpus crinita Jack verifies his statement. I have in-
troduced the photograph (Plate 1) chiefly because its extended label pro-
vides an excellent specimen of William Jack’s handwriting.

It is recorded that Jack, as well as Raffles, sent botanical material to
Lambert in England. Burkill notes, op. cit. 200, footnote 194, that at the
Lambert sale in 1842 lot no. 111, catalogued as probably from Jack, was
sold for £1 to William Pamplin, a dealer, and that lot 255, listed as from
Raffles and others, was purchased by a Mr. Rich for £3. What may have
become of the lot purchased by Mr. Pamplin is unknown, but the Jack
Sumatra specimens now in the Delessert herbarium in Geneva certainly
came from the Lambert collection. In 1879 Mr. Pamplin, then living at
Llandderfel, North Wales, provided Henry Trimen with the information
that Mr. Rich, and his father before him, were the accredited British agents
of Delessert; see Jour. Bot. 17: 275. 1879, footnote.

In April, 1952, Dr. Van Steenis found in a Gnetum loan from the Barker
Webb Herbarium, Florence, a single Jack specimen from Penang which
proved to be a representative of the genus Petunga. As the label carried
the statement ‘“‘misit amicissime Guilielmus Jack,” it is probable that this
Jack specimen had passed through the hands of Wallich, as the above is
the phrase that Wallich used in his List where he included Jack material.
There may, of course, be other Jack specimens in the Barker Webb
Herbarium.

SOURCES OF JACK’S BOTANICAL MATERIAL

After Jack’s arrival in India, attracted by its luxuriant flora, which was,
of course, entirely new to him, he commenced, during the Nepal campaign,
to correspond with Nathaniel Wallich in Calcutta, sending him a certain
amount of botanical material from northern India. This correspondence
apparently commenced in May, 1815. In July, 1818, after his return to
Calcutta, he called on Wallich at the Calcutta Botanical Garden, and the
latter insisted that Jack remain as his guest while prosecuting his botanical
investigations there. It developed that Jack was ill, and from a letter
written by Sir Stamford Raffles January 1, 1823, we learn that this illness
was pulmonary tuberculosis contracted during his tour of service in the
Nepalese war. This was soon to terminate a short but very productive
botanical career, for Jack died at Bencoolen, Sumatra, in September, 1822.
In November, 1818, after Wallich had introduced him to Sir Stamford
Raffles, his plans were abruptly changed, for he was offered a position on
Sir Stamford’s staff for botanical and other investigations primarily in
western Sumatra, centered at Bencoolen. He sailed with Raffles from
Calcutta December 10, 1818, and reached Penang on December 31. Pos-
sibly on this trip the ship on which he was a passenger stopped at Car
Nicobar, the most northern island of the Nicobar group, where Jack col-

1952] MERRILL, WILLIAM JACK’S GENERA AND SPECIES 209

lected some botanical material for the types of Microcos glabra Jack and
Connarus jackianus Wall. = Lepidopetalum jackianum Radlk.; yet it is
possible that these Car Nicobar specimens were collected when Jack made
a trip from Bencoolen to Calcutta and return later in 1819. Otherwise all
the Jack species were based on specimens collected by him in Penang,
Singapore (one on Pulo Bintang in the neighboring Rhio Archipelago),
and at various places, chiefly Bencoolen, on the west coast of Sumatra and
its neighboring islands.

PENANG AND SINGAPORE

William Jack commenced his field work in Penang, exploring that island
from January 1 to May 21, 1819. Thus about fifty of his published new
species were based wholly on Penang specimens. On May 31 he landed at
Singapore and he remained there until June 28. From the botanical speci-
mens he then prepared sixteen new species were described. For these
Penang and Singapore species supplementary material was mentioned in
a few cases as coming from Malacca, Sumatra, and in one case from Pulo
Bintang in the Rhio Archipelago southeast of Singapore. Most fortunately,
both Penang and Singapore have been intensively explored, and their floras
are very well known. Again Jack sent to Wallich in Calcutta specimens
representing most of his species, which were listed and distributed by
Wallich. Thus it is that there seems to be little or no doubt as to the limits
and relationships of all the Jack taxa based on material originating in these
two islands, for authentically named Jack specimens have been available to
his successors, in addition, of course, to Jack’s excellent published descrip-
tions.

SUMATRA

The longer period spent in Sumatra naturally resulted in much larger
collections of botanical material being made at various places on the west
coast of that large island and on various islands and islets off its west
coast. While Jack was engaged to prosecute botanical investigations, he
could not devote full time to this work, as various tasks quite unrelated to
botany were from time to time assigned to him. On the basis of the
Sumatran collections assembled by him a total of about 125 new species
were described. Of these about fifty are not more closely localized than
being from “Sumatra,” but in most cases it can safely be assumed that
the material on which they were based came largely from the Bencoolen
region. Somewhat over thirty species were definitely from Bencoolen and
its vicinity, including Gunong Bunko or Sugar Loaf Mountain, about
eighteen miles to the northeast of that town. Sixteen species are definitely
indicated as from Tapanuly and Tapanuly Bay, six species were from
Pulo Nias, one of the larger islands off the west coast, five from Natal
on the west coast, and for smaller west coast localities and west coast small
islands one or two species each were indicated as from Salumah, Kataun,

210 JOURNAL OF THE ARNOLD ARBORETUM | [voL. xxxmt

Laye, the Musi country, Moco-moco, Pulo Nica, Pulo Mosella, Pulo
Pegang, and Pulo Bintangor. These for the most part can be located with
little trouble on any of our better maps.

The flora of Sumatra is still inadequately known, particularly when con-
trasted with our knowledge of such areas as the Malay Peninsula (includ-
ing Penang and Singapore) and Java. If comprehensive and adequate
modern collections were available from the west coast of Sumatra and from
certain adjacent islands, the task of matching Jack’s descriptions with
such material would be relatively simple. Until such collections are avail-
able and are intensively studied, there will be a residue of Jack’s species
which will remain known only from his published descriptions. And to a
certain degree Jack has suffered because many of his published descriptions
were not generally available to his successors; and his types had been
destroyed.

JACK’S GENERA

In his relatively short career as a botanist, what Jack actually accom-
plished in descriptive botany is distinctly remarkable. When he reached
Penang the first of January, 1819, he found himself in a very luxuriantly
forested region rich in species regarding which he know nothing, and up
to the end of his most unfortunately short life he was surrounded by a
profusion of unclassified and unnamed plants, a very high percentage of
them quite unknown to the botanists of Europe and of India. There were
then no professional botanists in all of Malaysia, Jack’s chief contact
with the botanical world being by correspondence with Nathaniel Wallich
in Calcutta. He proposed and characterized one new family of plants, the
Cyrtandraceae, now placed as a subdivision of the Gesneriaceae, thirty-
one new genera, and about two hundred new species of plants. But he
published only a part of the descriptions he prepared. Writing from Penang
March 7, 1819, which he had reached just over two months earlier, he
stated that he had then described about 130 plants, of which eighty were
probably new, “besides examining and ascertaining the characters of at
least as many more.”’ Of some he personally prepared drawings, and he
employed a Chinese artist to prepare others. And this for Penang only,
with Singapore to come, and then the richer Sumatran flora which awaited
his attention!

Although at the beginning of the present century only two of Jack’s
new genera remained that had not been placed in their proper families,
these have now been disposed of. Coelopyrum Jack (1822) = Campnos-
perma Thwaites (1854), and Octas Jack (1822) = Jlex Linn. (1753).
Helospora Jack (1823) is identical with the officially conserved Timontus
DC. (1830); Enchidium Jack (1822) is earlier than the universally ac-
cepted Trigonostemon Blume (1825); Psilobium Jack (1822) antedates
the later and identical Acranthera Arnott (1838) by sixteen years; and
Coelopyrum Jack (1822) antedates the universally accepted Campnos-
perma Thwaites (1854) by thirty-two years. Unless officially conserved

1952] MERRILL, WILLIAM JACK’S GENERA AND SPECIES 211

here are three cases where Jack’s earlier generic names should replace the
later equivalents of Blume, Arnott, and Thwaites.

The other actually (and correctly) reduced Jack genera, as generic limits
are currently accepted, are Epithinia Jack (1820) = Scyphiphora Gaertn.
f. (1805); Glaphyria Jack (1823) = Leptospermum Forst. (1776); Pyrr-
hanthus Jack (1822) = Lumnitzera Willd. (1803); Sphalanthus Jack
(1822) = Quisqualis Linn. (1753); Stagmaria Jack (1820; 1832) = Gluta
Linn. (1753); Hedycarpus Jack (1823) = Baccaurea Lour. (1790);
Chionotria Jack (1822) = Glycosmis Corr. (1805); and Monocera Jack
(1820) = Elaeocarpus Linn. (1753). The eighteen remaining genera, all
universally accepted, are characteristic of the Indo-Malaysian floras, some
small in the number of known species, others large or very large.

There are doubtless those who might feel inclined to criticize Jack for
his failure properly to interpret a few previously described genera. Thus
Veratrum Linn. (one species), Pittosporum Banks (one species), Tern-
stroemia Mutis ex Linn. (five species), and Halorrhagis Forst. (one species)
were clearly misinterpreted, but in most other cases he correctly inter-
preted genera proposed by his predecessors. In the Ternstroemia case he
merely followed Roxburgh. One must constantly bear in mind that he did
not have access to herbarium material other than that which he himself
had prepared, that his library facilities were limited, and that conditions
in 1819—22 in the then primitive Penang, Singapore, and in the much more
remote port of Bencoolen, isolated as it was on the west coast of Sumatra,
were not favorable for scientific work. There were then in all Malaysia
no established scientific institutions or reference libraries, for up to that
time strangely little scientific work had been done in any field of biology,
following the pioneer work of Rumphius, who finished his extensive manu-
script in Amboina about 1690. Jack was the pioneer Malaysian botanist
after the binomial system was established, and he doubtless assumed that
if Roxburgh and his contemporaries and immediate successors could prose-
cute descriptive botany to advantage in India in the opening decades of the
nineteenth century, then he could do likewise in Malaysia. Some of us who
entered the field at the beginning of the present century with very limited
(or no) herbarium and library facilities may only hope that our percentages
of error were as small as were those of William Jack nearly a century earlier.
It is one thing to initiate descriptive work with ample herbarium and
library facilities available; it is quite another matter when one starts in
as did William Jack. To a distinctly high degree, with few books, no speci-
mens, and no previous knowledge of the flora, Jack was dependent on his
own efforts in such remote and primitive places as were Penang and
Singapore in 1819, to say nothing of the now almost forgotten Bencoolen.
He had no one to turn to for assistance or advice other than Wallich in
distant Calcutta, and yet he took full advantage of his opportunity. Very
few individuals would have had the courage to initiate descriptive work in
botany under the conditions that William Jack so successfully faced in
the early decades of the last century.

212 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxmI

JACK’S VALIDATION OF CERTAIN ROXBURGHIAN
NOMINA NUDA

Jack actually validated certain nomina nuda proposed by Roxburgh in
his Hortus Bengalensis (1814) by accepting the binomials and associating
technical descriptions with the names from three to twelve years before
Roxburgh’s own validating descriptions were published. Jack had a manu-
script copy of Roxburgh’s Flora Indica for consultation, the dates of pub-
lication of the several volumes (two editions) of Roxburgh’s work being
1820, 1824, and 1832.. In some cases, doubtless, identifications of Jack
specimens with Roxburghian species were made in Calcutta by Wallich.
Cases are Curculigo sumatrana Roxb., Gmelina villosa Roxb., Loranthus
ferrugineus Roxb., Melastoma decemfidum Roxb., Phyteuma begonifolium
Roxb., Rottlera alba Roxb., Sterculia angustifolia Roxb., and Vitex arborea
Roxb. There are a few similar cases in relation to Wallich’s binomials. In
one or two cases it is evident that the species actually described by Jack
under a Roxburghian epithet is not the same as the one to which Roxburgh
assigned the binomial and which Wallich later published; see the case of
Clerodendron nutans Jack (C. penduliflorum Wall.), 1820, not C. nutans
Wall. List 1829, nom. nud., et ex Hook. Bot. Mag. 53: pl. 3049. 1831, descr.
These and various other minor bibliographic adjustments are made in this
paper, and under the priority rule a certain number of new binomials
appear in the index proper which follows this introduction.

The number of changes in names is small, indicating that much time and
attention have been given by various botanists to ascertaining the status
of these early Jack species, even if others, in the absence of types, may
have ignored the Jack species, or at least made no really serious attempt
to interpret them. As a result, a certain number of species proposed and
described by later authors as new are reduced to synonymy. Gradually
the situation clears, for the status and relationships of most of the Jack
species, whether the types be preserved or not, are now clear.

WILLIAM ROXBURGH’S CONCEPT OF THE MOLUCCAS

As one examines the Roxburgh text of his Flora Indica, one notes an
occasional entry, accompanied by a short description, followed by the entry
“Moluccas.” The natural assumption in such cases is that the material
on which these short descriptions came originated in that group of islands
south of the Philippines and east and southeast of Celebes to which the
term Moluccas is now and probably always was correctly limited. But
Roxburgh’s concept of the Moluccas included, at times, also the Malay
Peninsula and the Sunda Islands proper,'! and so it is that various “Moluc-
can” species of Roxburgh unquestionably came from Penang or from vari-
ous localities on the Malay Peninsula or in the Sunda Islands proper.
It is not necessary, as some have done, to suggest that in such a case as

4 Prain, D. A Brief Memoir of William Roxburgh. Ann. Bot. Gard. Calcutta 5:
1-9, portr. 1895 (p. 6).

1952] MERRILL, WILLIAM JACK’S GENERA AND SPECIES 213

Sonerila moluccana Roxb. he perhaps intended to derive his specific name
from Malacca. Examples are: Ardisia divergens Roxb., Melastoma im-
puber Roxb., Sonerila moluccana Roxb., Uvaria pilosa Roxb. (= Uvaria
hirsuta Jack), and others. These species are now known from Penang
and neighboring places, but have never appeared in any Moluccan collec-
tions and are still unknown from any part of eastern Malaysia. Yet Rox-
burgh did indeed have much botanical material from Amboina and from
other parts of the Moluccas proper. Some idea of the importance of Rox-
burgh’s contributions in his Flora Indica to our knowledge of the Malay-
sian flora is indicated by the following data. About 540 Roxburghian
descriptions are to be interpreted from Malaysian material. Of these ap-
proximately 435 were proposed and described as new on the basis of Malay-
sian specimens. Of these 435 ‘‘new species’ 104 were from Penang, 157
indicated as from the Moluccas, plus 54 from Amboina and Honimoa, 56
from Sumatra, 36 from the Malay Archipelago, with a few indicated more
definitely as from Singapore, Malacca, Banda, etc. Doubtless some, per-
haps many, of the ‘““Moluccan” species were from the Moluccas proper,
but one must constantly bear in mind that probably most of these were
from the Sunda region proper, the Malay Peninsula, Penang, Singapore,
and Sumatra, and not from the Moluccas. These Roxburghian Malaysian
species have not been properly studied and an investigation of them in
relation to those described by other authors is highly desirable.

EXPLANATION OF THE SEVERAL CATEGORIES USED IN
THE FOLLOWING LIST APPERTAINING TO THE
REPUBLISHED JACK DESCRIPTIONS

In the following list of the Jack species I have included references to
the original place of publication of each taxon, and also references to those
places where the descriptions were republished. To save repetition of
references the classification I to V is accepted as explained below, of
which I is scarcely used as such, II seldom used (because in these cases
the references are repeated), but III to V are always used if individual
Jack descriptions were included in this or that set of reprinted descriptions.

. The original Jack papers. In each case a reference is given, the cate-

gory indication I not used.
II. The Hooker reprinted descriptions 1830-1836. For details see p. 204.
In each case the complete reference is given rather than merely II.
III. The Griffith Calcutta Journal of Natural History papers, volume four
(1843). For details see p. 205.
IV. The separately paged Griffith reprint of the above. For details see
205

p. :
V. The 1887 reprint in Triibner’s Oriental Series. For details see p. 206.
These technical names preceded by an asterisk, such as Acacia *grav-
eolens Jack still remain unlisted in standard indices. Most of these are
nomina nuda and appear in Jack’s letters to Nathaniel Wallich, which were
published in 1916. Yet although actual descriptions may never have been
published, most of these fugitive binomials are safely identifiable.

214 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxmI

ACACIA Willdenow.

A. *graveolens Jack, Mal. Misc. 2 (7): 78. 1822; reimpr. Hook. Comp. Bot. Mag.
1: 224. 1836; III. 163; IV. 67; V. 285, nom. nud. in obs. [Sumatra] = Parkia
graveolens King, Jour. As. Soc. Beng. 66 (2): 241. 1897 (Mater, Fl. Mal.
Pen. 3: 241) nom. in obs. = Parkia speciosa Hassk. Flora 25 (2): Beibl. 55.
1842 (P. macrocarpa Miq. Fl. Ind. Bat. 1: 53. 1855). Malay Peninsula,
Sumatra ; oe in Java. Acacia graveolens Jack, Parkia graveolens Prain,
and the very much older Acacia gigantea Noronha (1790) are all nomina

nuda, although al are safely identifiable by the cited Malay name pete or

petek.

ACROTREMA Jack, Mal. Misc. Z — 36. 1820; reimpr. Hook. Bot. Misc. 2:
81. 1830; III. 217; IV. 121; V.
A. costatum Jack, |.c.; reimpr. et op. cit. 82; III. 217; IV. 121; V. 240.
enang. Common in the Malay Peninsula; see Ridley, Fl, Malay Penin. iT
7.1922. Also in Borneo and Sumatra

ADINANDRA Jack, Mal. Misc. 2 (7): 49. 1822; reimpr. Hook. Comp. Bot. Mag.
1: 153. 1835; III. 205; IV. 110; V. 271.
A. dumosa Jack, op. cit. 50; reimpr. ll.cc. Sumatra and other Malay Islands.
The type of the genus; common in the Malay Peninsula, Sumatra, Java
A aaa only planted), and Borneo, represented by very ie collec-
; for synonymy see Kobuski, Jour. Arnold Arb. 28: 55.
A. pines Jack, Mal. Misc. 2 (7): iii. 1822; reimpr. Calc. see cm Hist. 4:
8. 1843; IV. 112; V. 295. Western Sumatra at Moco Moco. Not actually
described and cannot be placed from the inadequate data; see Kobuski, op.
cit. 93.

AESCHYNANTHUS Jack, Trans. Linn. Soc, 14: 42. 1823; reimpr. Calc. Jour.
Nat. Hist. 4: 60. 1843; IV. 60, nom. conserv. (Trichosporum G. Don, 1822).

A. radicans Jack, op. cit. 43; reimpr. III. 62; IV. 62. Sumatra, inland from Ben-
coolen. (Trichosporum radicans Nees). Jack’s type is apparently in the
Geneva herbarium; see C. B. Clarke, Monog. Phan. 5: 41. 1883. Sumatra,
Malay Peninsula, oS

A. volubilis Jack, l.c. pl. 2, fig. 3, a-i; reimpr. III. 61. pl. 15, fig. 3; IV. 61.
Sumatra, near Bencoolen (Trichosporum volubile Nees). Definitely known
only from Sumatra, but has been credited to Celebes.

AGLAIA Loureiro.

A. odorata Lour. Fl. Cochinch. 173. 1790; Ele Mal. Misc. 1 see 1821;
reimpr. Hook. Bot. Misc. 2: 79. 1830; III. 192; IV. 96; V. ey " [Malay
Islands; planted]. Widely planted in the Old World, native of southeastern
Asia.

ALPINIA Roxburgh (1810), mom. conserv., non Linnaeus.

A. capitellata Jack, Mal. Misc. 2 (7): 4. 1822; reimpr. Hook. Jour. Bot. 1: 360.
1834; III. 5; IV. 5; V. 248. Inland from Bencoolen, Sumatra. See Holttum,
Gard. Bull. Singapore 13: 143. 1950, who tacitly accepted Ridley’s 1899 inter-
pretation of the species. There is no extant type. Sumatra and the Malay
Peninsula.

A. elatior Jack, Mal. Misc. 2 (7): 2. 1822, reimpr. Hook. Jour. Bot. 1: 359.
1834; III. 4, sphalm, “elatoir”; IV. 4; V. 247. Pulo Nias and Ayer Bangy =

1952] MERRILL, WILLIAM JACK’S GENERA AND SPECIES 215

Nicolaia elatior (Jack) Horan. Monog. Scit. 32. 1862. This case is an illus-
tration of how one a sometimes be led astray by accepting modern inter-
ee hans. even in standard monographic treatises, without checking the
record. Nicolaia ee was validly published in 1862, its type being NV. im-
perialis Horan. Its author knew of the earlier but invalidly published generic
name Phaeomeria Lindl., for he listed it as a synonym. The latter was pub-
lished by Lindley, Nat. Syst. ed. 2, 446. 1836, the entire entry being “Phae-
omeria = Alpinia magnifica Bojer in Bot. Mag. t. 3192.” This does not con-
stitute valid publication under the conditions specified in Article 41 of the Code
of Botanical Nomenclature, in spite of K. Schumann’s acceptance of Lindley’s
generic name in 1904 (who first published a description of Phaeomeria Lind1.,
although Ridley in 1899 had treated it as a section of Hornstedtia Retz.) and
Loesener’s selection of it in preference to Nicolaia Horan. in Engl. & Prantl,
Nat. Pflanzenfam. ed. 2, 15a: 593. 1930. K. Schumann in 1904 had recognized
sixteen species of Phaeomeria Lindl. The code provision is clear in that the
name of a genus “is not validated by mention of included species” (this is all
that Lindley did, for he never published a generic description); nor can
Lindley’s generic name be validated under any of the exceptions to this rule.
Both K. Schumann and Loesener should have accepted Nicolaia Horan., as
this is the proper name for this genus. In 1921 Valeton !? correctly interpreted
the situation, accepting Nicolaia Horan. and critically considering fourteen
species. It is unfortunate that he did not explain why he rejected Phaeomeria
Lindl. (correctly), for such action might have rendered this discussion unnec-
essary. However, this is perhaps an optimistic statement, considering the
conservatism of the average taxonomist and the tendency that some have to
justify the name-selections of their predecessors, regardless of approved rules.
Except for his several new species being properly listed in Index Kewensis,
Valeton’s paper has been rather consistently ignored. It is worthy of note that
although Valeton did not accept Jack’s specific name (which he listed as a
synonym of Nicolaia speciosa Horan.), he did have specimens from the type
locality (Nias Island) and cited other collections from Sumatra. The last to
consider our particular species was Holttum, Gard. Bull. Singapore 13: 181.
1950, who, while providing a nicely detailed description, was not at all im-
pressed by Valeton’s correct selection of Nicolaia Horan. as the correct generic
name; nor was he impressed by the manifest fact that Jack’ s specific name
had five years priority over the one he accepted. Thus it is that Jack’s beau-
tifully described species has been rather consistently ignored, and in the mean-
time it has acquired a rather extensive synonymy, being, I suppose, the most
spectacular species in the Zingiberaceae. The extensive synonymy is due,
in fact, to the reluctance of some taxonomists to interpret species from de-
scriptions alone when the types are lost, even when some of these descriptions,
like those of Jack; are remarkable for their clarity, and further, to the re-
luctance of others to accept what manifestly is the oldest valid name for a
particular species. These synonyms include Elatteria speciosa Blume (1827),
Alpinia magnifica Rosc. (1828), Phaeomeria *imperialis Lindl. ex K. Schum.,
Pflanzenr. 20 (IV. 46): 262. 1904, Alpinia speciosa Dietr. (1839), Nicolaia
imperialis Horan., and N. speciosa Horan, (1862), Phaeomeria magnific
Schum. (1904), Amomun magnificum Benth, (not published until it appeared
in Index Kewensis 1: 108. 1893), Hornstedtia imperialis Ridl. (1899), Phae-

2 Valeton, T. Nicolaia Horan. Description 7 New and Interesting Species. Bull.
Jard. Bot. Buitenz. III. 3: 128-140. pl. 1-5. 192

216 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxim

omeria speciosa Koord. (1911; Merr., 1923), and Alpinia*longiscapa Jack
ex Burkill, Jour. Straits Br. Roy. As. Soc. 73: 225. 1916, nom. No matter how
desirable it may be to retain Lindley’s invalidly published generic name of
1836, I fail to see how this can be done unless one wishes to ignore the code
provisions governing valid publication. In selecting the name Phaeomeria,
Lindley was undoubtedly influenced by Bojer’s suggestion, in the discussion
of the beautiful plate of Alpimia magnifica [Rosc.], Bot. Mag. 59: pl. 3192.
1832, that a new genus might be represented. It is most unfortunate that he
never found time to characterize his suggested new genus; but K. Schumann’s
tardy recognition of the validity of the group as a genus in 1904, and
Loesener’s action in 1930, in an apparent attempt to justify K. Schumann’s
selection of a generic name for the group, do not save the day for Lindley’s
generic name. Nicolaia elatior (Jack) Horan. is widely distributed in Malaysia,
much of its range being due to this strikingly ornamental plant being man-
distributed; it has also been introduced in many other tropical countries in
both hemispheres, but is unquestionably of Malaysian origin.

AMOMUM Linnaeus.

A. biflorum Jack, Mal. Misc. 1 (1): 2. 1820; reimpr. Hook. Bot. Misc. 1: 274.
1830; III. 3; IV. 3; V. 210. Penang. For the best modern consideration of
the species, with synonymy, see Holttum, Gard. Bull. Singapore 13: 199.
1950. Siam to the Malay Peninsula.

gp melo Linnaeus.

. frutescens Jack, Mal. Misc. 2 (7): 91. 1822; reimpr. Hook. Comp. Bot.
ag Es 257; 1836: III. 229; IV. 133; V. 29 2. Bencoolen, Sumatra = Antt-
desma ghoesembilla Gaertn. This was ‘accepted as a valid species by Pax &
Hoffmann, Pflanzenr. 81 (IV. 147. XV): 157. 1922, and was placed by them in
the alliance with Gaertner’s species. An attentive comparison of Jack’s excel-
lent description (there is no extant type) with Gaertner’s species clearly indi-
cates that what Jack described is only a form of the very common and widely
distributed A. ghaesembilla Gaertn., which might have been expected from
Jack’s comparison of his species to A. pubescens Roxb. Rahmat si Toroes
4185 from Sumatra, which is clearly A. ghaesembilla Gaertn., agrees closely
with Jack’s description, as do other Malayan collections. Western India and
the tropical Himalayan region to Ceylon, eastward to southeastern China and
southward a Malaysia, including the Philippines, to New Guinea and
tropical Australia

ARDISIA Swartz.

A. punctata Jack in Roxb. Fl. Ind. 2: 275. 1824. Penang. A species known only
from Penang and of which A. divergens Roxb. Hort. Beng. 85. 1814, nom. nud.,
Fl. Ind. l.c., is a synonym. Roxburgh said that his specimen came from the
Moluccas, but it should be realized that his concept of the Moluccas covered
all of the Malay Peninsula and Archipelago; see Prain, Ann. Bot. Gard. Cal-
cutta 5: 6. 1895. The actual type of A. divergens Roxb. undoubtedly came
from Penang. The Roxburgh aa Ae compared with that which Wallich
prepared for Jack’s aie is very s

Ardisia punctata Jack was il ri by Griffith when he prepared his
1843 paper on the Jack descriptions. This interpretation of the Jack species
necessitates a new specific name for the common Chinese Ardisia punctata
Lindl., Bot. Reg. 10: pl. 827. 1824, as this plate is dated Sept. 1, 1824. The

1952] MERRILL, WILLIAM JACK’S GENERA AND SPECIES 217

introduction to volume two of Roxburgh’s Flora Indica is dated March, 1824.
The proper name for the Chinese Ardisia punctata Lindl., non Jack, is Ardisia
lindleyana D. Dietr. Syn. 1: 617. 1839

ARECA Linnaeus.

A. tigillaria Jack, ‘er se 2 os 88. 1822; reimpr. Hook. Comp. Bot. Mag. 1:
256. 1836; III. LV; 12% 290. Sumat tra and the Malay Islands
Oncos perma ee (Jack) a (O. flamentosum Blume). In gis die
the specific name to Oncosperma Ridley credited the original binomial to
Griffith, who, however, was merely concerned with Jack’s species. Malay
Peninsula, Sumatra, Borneo, and Java.

ARISTOLOCHIA Linnaeus.

A. hastata Jack, Mal. Misc. 2 (7): 6. 1822; reimpr. Hook. Jour. Bot. 1: 362.
1834; III. 358; IV. 214; V. 249, non HBK. (1817). West coast of Sumatra
at Natal = A. jackii Steud. Apparently known only from Jack’s description.

BAUHINIA Linnaeus.

B. bidentata Jack, Mal. Misc. 2 (7): 76. 1822; reimpr. Hook. Comp. Bot. Mag. 1:
223. 1836; III. 160; IV. 63; V. 284. “Native of Malayan Forests.” This species
of the Malay Peninsula and Sumatra is amply described ii King, Jour. As. Soc.
Beng. 66 (2): 187. 1897 (Mater. Fl. Mal. Penin, 3: 18

B. emarginata Jack, op. cit. 76: reimpr. Il.cc., non an (1768). Sumatra =
? Bauhinia lucida Wall. Baker’s interpretation of Jack’s species is possibly
correct, although B. lucida Wall. seems to be definitely recorded only from
Penang and Perak. However, there is in the Gray Herbarium a Marsden speci-
men from Sumatra named by Hooker as B. lucida Wall. which may represent
both it and the form Jack characterized. In any case Jack’s specific name is
an invalid one.

BEGONIA Linnaeus; Jack, Mal. Misc. 2 Se 8. 1822; reimpr. Hook. Jour.
Bot. 1: 363. 1834; TIL. 342; IV. 198; V.

B. bracteata Jack, op. cit. 13; reimpr. II. ie an 346; IV. 202; V. 353. Gunong
Bunko, inland from Pancovlen: Sumatra. A de Candolle’s description, Prodr.
15 (1): 316. 1864, was based on that of Jack; type not extant. Placed in
Diploclinium by Miquel and in Knesebeckia by Hasskarl. But Koorders in
1912 reduced it to Begonia lepida Blume (1827), although Jack’s name is
older.

B. ene Jack, l.c.; reimpr. II. 363; III. 342; IV. 198; V. 250. West coast

matra at Bencoolen. Known only from Jack’s description; see A. de
Ca a ‘le, op. cit. 397. Placed in Diploclinium by Mique

B. fscicitlata Jack, op. cit. 12; reimpr. II. 365; III. 345; IV. 201; V. 252. West
coast of Sumatra at Tapanuly. See A. de ‘Candolle, op. cit. 322. Placed in
Diploclinium by Miquel and in Petermannia by Klotzsch. Known only from
Jack’s description.

B. diaereo Jack, op. cit. 15; reimpr. II. 368; III. 347; IV. 203; V. 253.
Sumatra = B. isoptera Dryand. (1791), fide A. de Candolle, op. cit. 320, the
latter aie in the Malay Peninsula, Borneo, and Java, as the species is
currently interpreted.

B. orbiculata Jack, op. cit. 9; reimpr. II. 364; III. 343; IV. 198; V. 250. West

coast of Sumatra at Bencoolen. Known only from Jack’s description, Placed
in Diploclinium by Miquel; see A. de Candolle, op. cit. 398.

218 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxxuI

B. pilosa Jack, op. cit. 13; reimpr. II. 366; III. 345; IV. 201; V. 252. West
coast of Sumatra inland from Bencoolen. Known only from Jack’s descrip-
tion. By Miquel placed in Diploclinium; see A. de Candolle l.c.

B. racemosa Jack, op. cit. 14; reimpr. II, 367; III. 346; IV. 202; V. 253. West
coast of Sumatra, inland from Bencoolen. Known only from Jack’s descrip-
tion. Placed by Miquel in Diploclinium and by Klotzsch in Petermannia;
see A. de Candolle, op. cit. 322.

B. sublobata Jack, op. cit. 10. 1822; reimpr. II. 364; III. 343; IV. 198; V. 251.
Under moist rocks on Pulo Pegang, west coast of Sumatra. Known only from
eae ee Placed by Miquel in Diploclinium; see A. de Candolle,
op. cit. 354.

CALLA Linnaeus.

C. angustifolia Jack, Mal. Misc. 1 (1): 24. 1820; reimpr. Hook. Bot. Misc. 1:
288. 1830; III. 11; IV. 11; V. 221. Penang = Homalomena humilis (Jack)
Hook. f. var. pumila (Hook. f.) Furtado, Gard. Bull. Straits Settl. 10: 203.
1939, cum syn. Malay Peninsula, Sumatra, Borneo (Chamaecladon angusti-
folium Schott).

C. humilis Jack, op. cit. 22; reimpr. Hook. Bot. Misc, 1: 288. 1830; III. 11;
IV. 11; V. 221. Penang = Homalomena husnilis (Jack) Hook. f. Fl. Brit.
Ind. 6: 533. 1893; Furtado, Gard. Bull. Straits Settl. 10: 199. 1939. Malay
Peninsula, Sumatra (Batu Island), and Anambas Islands.

C. nitida Jack, op. cit. 24; reimpr. Hook., op. cit. 289; III. 12; IV. 12; V. 221.
Penang = Aglaonema nitidum (Jack) Kunth, Enum. 3: 76. 1841 (A. oblongi-
folium (Roxb.) Schott; Engl. Pflanzenr. 64 (IV, 23, Dc.): 13. fig. 4. 1915, cum
syn.). Malay Peninsula, Buru, Borneo, Sumatra. Engler should have adopted
Jack’s specific name, as it was published twelve years earlier than that of
Roxburgh; the two species are clearly identical.

CAREYA Roxburgh.

C. macrostachya Jack, Mal. Misc. 1 (5): 47. 1820; reimpr. Hook. Bot. Misc. 2: 88.
1830; III. 305; IV. 161; V. 245. Penang = Barringtonia macrostachya (Jack)
Kurz, Malay Peninsula, Sumatra, Borneo.

CELASTRUS Linnaeus.

C. bivalvis Jack, Mal. Misc. 1 (5): 19. 1820; reimpr. Hook. Bot. Misc. 2: 71.
1830; III. 196; IV. 100; V. 231. Penang = Microtropis bivalvis (Jack)
Wall. List, no. 4340. 1840; Merr. & Freem. Proc. Am. Acad. Arts Sci. 73:
301. 1940, cum syn. (Paracelastrus bivalvis Miq.). Jack’s original collection
was distributed as a part of Wallich 4340, A species still known only from
Penang.

CELTIS Linnaeus.
C. *attenuata Jack ex Burkill, Jour. Straits Br. Roy. As. Soc. 73: 196. 1916,
nom. nud, Sumatra at Tapanuly, said to be frequent

CHIONOTRIA Jack, Mal. Misc. 2 (7): 53. 1822; reimpr. Hook. Comp. Bot. Mag.
1: 155. 1835; III. 193; IV. 97; V. 273 = Glycosmis Correa (1805).

C. rigida Jack, op. cit. 54: reimpr. Hook. l.c.; III. 193; IV. 97; V. 273. Penang =
Glycosmis rigida (Jack) comb. nov. hs Glycosmis macrophylla Lindl. in Wall.
List no. 6377. 1830, nom. nud.; Ridl. Jour. Straits Br. Roy. As. Soc. 75: 13.
1917, descr., incl. var. macrorachis (King) Ridl. l.c.; G. pentaphylla Corr. var.
macrorachis ing). In the first place Ridley never should have validated

1952] MERRILL, WILLIAM JACK’S GENERA AND SPECIES 219

Lindley’s species, for, as a binomial, it was invalidated by the different G.
macrophylla Miquel; hence, as long as an invalid binomial is currently applied
to this Penang species, I do not hesitate to replace it by the much earlier
specific name published by Jack in 1822. The only character indicated by
Jack that does not conform to Ridley’s description is that he stated that the
leaves were opposite; they are alternate in all species of Glycosmis. Note
particularly Jack’s description of the inflorescences as racemes. Actually the
inflorescences are very narrow panicles, the distant branchlets being often
only 0.1 inch long, varying from 0.1 to 0.4 in. in length, and thus simulating
racemes. When Ridley considered the species in 1917, he stated, “a very
distinct plant peculiar apparently to Penang,” and in his Fl. Mal. Pen. 1: 349.
1922, he had not extended its range. The only other possibility would be G.
malayana Ridl., which occurs also in Penang, but this has pinnate leaves (had
Jack’s specimen had other than simple leaves surely he would have mentioned
it), while its paniculate inflorescences have branches up to one inch long;
Jack never would have characterized such an inflorescence as a raceme.
species characterized essentially by its very narrow raceme-like inflorescences.
still known only from Penang.

CLERODENDRON Linnaeus.

C. divaricatum Jack, Mal. Misc. 1 (5): 48. 1820 (Clerodendrum) ; reimpr. Hook.
Bot. Misc. 2: 89. 1830; III. 40; IV. 40; V. 246. West coast of Sumatra at
Laye = Clerodendron serratum (Linn.) Spreng. India and Ceylon to Mada-
gascar and the Mascarene Islands through Malaysia to the Lesser Sunda
Islands and Celebes.

C. molle Jack, Mal. Misc. 1 (1): 15. 1820; reimpr. Hook. Bot. Misc. 1: 283. 1830;
III. 38; IV. 38; V. 217, non HBK. (1817). Sumatra and Penang = C. villosum
Blume (1826). Another synonym is Clerodendron velutinum Wall. List no.
1797. 1829, nom. nud. India and Burma to the Malay Peninsula and Archi-
pelago, including the Philippines.

C. nutans Jack, Mal. Misc. 1 (1): 17. 1820; reimpr. Hook. Bot. Misc. 1: 284.
1830; III. 39; IV. 39; V. 217, omn. sub C. molle Jack. Penang (C.
penduliflorum Wall. List no. 1795. 1829, nom. nud. et ex Schauer in DC. Prodr.
11: 664. 1847, descr.), non C. nutans Wall. List no. 1793. 1829, nom. nud. et
ex D. Don, Prodr. Fl. Nepal. 103. 1825, descr. Jack’s description, although
short, is excellent. It was based on Penang material, as he thought that which
he had before him represented the as yet undescribed C. nutans Wallich.
When one scans Jack’s graphic description, “paniculis longissimis terminalibus
nutantibus, pedunculis [ramis] remotis paucifloris,” and again “these panicles
or racemes hang gracefully from the extremity of the branches,” it is under-
standable why Jack thought that he had before him a representative of
Wallich’s species. Wallich erred, List no. 1794. 1829, when he renamed what
he supposed to be the form Jack had described as C. jackianus Wall.; this,
as later described by Schauer, based on the actual Wallich specimen, explains
why the very different C. disparifolium Blume, C. laevigatum Blume, and C.
acuminatum Wall. became involved here. Mr. H. K. Airy Shaw reports that
Wallich 1794 from Penang (this was collected by Wallich in 1822, not by
Jack), in his opinion, represents the very different C. disparifolium Blume.
For the binomial as here accepted and applied, that is C. mutans Jack (non,
Wall.), C. penduliflorum Wall. is a synonym, as Wallich’s species is defined
and amply described by Gamble in King and Gamble, Jour. As. Soc. Bengal
74 (2): 830. 1909 (Mater. Fl. Mal. Pen. 4: 1040), and accepted by Ridley.

220 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxx

Its range is apparently Burma, the Nicobar Islands, Penang, and various parts
of the Malay Peninsula. I am confident that a Korthals collection from Mt.
Singalang, Sumatra (a rather poor specimen of which is before me), which
Hallier f., Meded. Rijksherb. 37: 72. 1918, listed as C. mutans Wall., really
represents C. nutans Jack. The Indian form was not introduced into cultiva-
tion in Malaya before 1820; and Ridley is clear, as to this Malay Peninsula
form with pendulous inflorescences, that it occurs here and there in forests —
i.e., that it is a native of the ;

Schauer, in 1847, bea C nutans Wall. (Bengal, Sylhet), C. jackianum
Wall. (Penang), and C. penduliflorum Wall. (Tavoy) as distinct species.
All taxonomists have overlooked the fact that as far as the binomial C. nutans
is concerned, Jack was the first author who associated a description with it,
and that the binomial to be maintained must hence be Clerodendron nutans
Jack (1820). It seems to be clear that the common Indian form, currently
known as Clerodendron nutans Wall., of which at least fifteen individual collec-
tions are available to me from northern India to Burma, as well as specimens
taken from cultivated plants in Cuba and in Australia, has no valid name.
This is unfortunate, because now that species is widely distributed in cultiva-
tion. For this a new binomial is proposed, Clerodendron wallichii nom. nov
(C. nutans Wall. List no. 1793. 1829, nom. nud., et ex D. Don, Prodr. Fl. Nepal.
103. 1825, descr., et auctt. plur., non Jack, 1820). This species was beautifully
illustrated by Hooker, Bot. Mag. 58: pl. 3049. 1831. The species occurs in the
Malay Archipelago only as an introduced and cultivated plant, unless one be
willing to interpret C. nutans Wall. as being identical with C. nutans Jack,
together with C. penduliflorum Wall., a proceeding that I am not willing to
approve. Clerodendron jackianum Wall., as described by Schauer, and C.
acuminatum Wall. are totally different from C. wallichii Merr.

CNESTIS Jussieu.

C. emarginata Jack, Mal. Misc. 2 (7): 42. 1822; reimpr. Hook. Comp. Bot. Mag.
1: 150. 1835; III. 166; IV. 70; V. 267. Sumatra, at Bencoolen = Roureopsis
emarginata (Jack) comb. nov. (Roureopsis javanica Planch. Linnaea 23: 424.
1850, excl. syn. Blume; Schellenb. Pflanzenr. 103 (IV. 127): 113. 1938).
Schellenberg erred, op. cit. 142, when he disposed of Jack’s species as a
synonym of the utterly different Santaloides mimosoides (Vahl) O. Kuntze,
which has numerous small, truncate-emarginate leaflets. Jack clearly states
that the 5 to 7 leaflets of C. emarginata Jack were long-acuminate and
emarginate, the terminal leaflet frequently 7 inches long. His graphic de-
scription agrees perfectly with Roureopsis javanica Planch. Connarus javanica
Blume (1826) = Rourea javanica Blume (1850), which has been confused
here, is a synonym of pa eet ie floridum (Jack) O. Kuntze. There are now
very many collections available from western Sumatra which agree with
Jack’s excellent description, such as Rahmat Si Toroes 3279, 3369, 3421,
3506, 3599, 3746, 3813, 3939, 4095, 4147, 4246, Bartlett 2882, 6894, as
well as those of Planchon and of Schellenberg. The very conspicuously
acuminate leaflets, the distinctly retuse tips of the acumens are characteristic.

C. florida Jack, op. cit. 43; reimpr. Hook. op. cit. 151; III. 167; IV. 70: V. 267.
Sumatra, west coast, and Pulu Nias = Santaloides floridum (Jack) O. Kuntze;
Schellenb. op. cit. 124, cum syn. Malay Peninsula, Sumatra, Java, Borneo,
and Morotai

C. *longifolia on ex _ Jour. Straits Br. Roy. As. Soc. 73: 197, 249. 1916,
nom. nud. Singapor

1952] MERRILL, WILLIAM JACK’S GENERA AND SPECIES 221

C. mimosoides Jack, op. cit. 44; reimpr. Hook. I.c.; III. 167; IV. 71; V. 268.
Sumatra at Tapanuly = Suntaloides mimosoides (Vahl) O. Kuntze; Schellenb.
op. cit. 142, cum syn. (excl. Cnestis emarginata Jack). Jack cited Connarus
mimosoides Vahl as the basis of his binomial and his interpretation of Vahl’s
species was apparently correct. Siam and Indo-China, the Malay Peninsula,
Nicobar Islands, Sumatra, Borneo, and Java.

COELOPYRUM Jack, Mal. Misc. 2 (7): 65. 1822; reimpr. Hook. Comp. Bot.
Mag. 1: 220. 1836; III. 341; IV. 197; V. 279 = Campnosperma Thwaites
(1854)

C. coriaceum Jack, l|.c.; reimpr. ll.cc. West coast of Sumatra near Bencoolen =
Campnosperma coriacea (Jack) Hallier f. ex van Steenis, Fl. Males. Bull. 3:
74. 1948 (C. macrophylla (Blume) Hook. f.). Malay Peninsula, Sumatra,
Borneo,

This genus remained among the unplaced ones until Hallier f., Beih. Bot.
Centralbl. 39 (2): 161, 162. 1921, correctly associated it with Campnosperma
Thwaites. Van Steenis, l.c., has recommended that Thwaites
be officially conserved against Jack’s earlier one. I cannot distinguish C.
macrophylla (Blume) Hook. f. from Jack’s species, Blume’s taxon dating from
1850.

CONNARUS Linnaeus.

C: gue Jack, Mal. Misc. : AeA 37. 1822; reimpr. Hook. Comp. Bot. Mag.
1: 149. 1835; III. 170; IV. 73; V. 264. Penang. Widely erate in the
Malay Pein see ah Pflanzenr. 103 (IV. 127): 258. 19

C. grandis Jack, o . 40; reimpr. Hook. Comp. Bot. Mag. 1: ee — III.
£72: 1V..76; v Sumatra at Tapanuly. The species is now known pe
the Malay Peninsula, Sumatra, Borneo, Java, and the Moluccas; see
Schellenb. op. cit. 257, cum syn.

C. lucidus Jack, op. cit. 41; reimpr. Hook. Comp. Bot. Mag. 1: 150. 1835; IIT.
172; IV. 76; V. 266. Sumatra. See Schellenb. op. cit. 112, who discussed this
species under Roureopsis pubinervis Planch. (1850) of the Malay Peninsula,
Lingga, Bangka, Sipora, and Sumatra; but as Schellenberg noted, Jack’s des-
cription does not wholly agree with the characters of Planchon’s species.

C. semidecandrus Jack, op. cit. 39; reimpr. Hook. Comp. Bot. Mag. 1: 149. 1835
(semidecander) ; III. 171; IV. 75; V. 266. West coast of Sumatra. A species
known only from Sumatra, C. pyrrhocarpus Miq. (1863) being a synonym.
Jack’s type is preserved in the Delessert herbarium at Geneva; see Schellen-
berg, op. cit. 281.

, oF oe Jack, op. cit. 38; reimpr. Hook. Comp. Bot. Mag. 1: 149. 1835; IIT.

IV. 74; V. 265. Sumatra. Also in the Malay Peninsula and Borneo.
rhea to Schellenberg, i cit. 228, Jack’s type is preserved in the
Delessert herbarium at Gen

CURCULIGO Gaertner.

C. sumatrana Roxb. Hort. Beng. 24. 1814; Roxb. ex Jack, ar rs eG Oe ea 2
1820; reimpr. Hook. Bot. Misc. 1: 277. 1830; III. 8; IV. 8; V. 212; Roxb.
Fl. Ind, ed. 2, 2: 146. 1832. Sumatra and Penang = C. ae es (1811),
the type of which was also from Penang. Burma to Indo-China southward
through the Malay Peninsula and Archipelago to the Moluccas. Jnvolucrum
Rumph. Herb. Amb. 6: 114. pl. 53. 1750, actually typifies Roxburgh’s taxon
as published in 1814; it was also cited by Jack. It has been erroneously

222 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxxmr

referred to the different C. recurvata Dry. = C. capitulifera (Lour.) O.
Kuntze. The Singapore form with hirsute leaves, mentioned but not named or
described by Jack, was undoubtedly C. villosa Wall.

CYRTANDRA Forster.

C. aurea Jack, Trans. Linn. Soc. 14: 29. 1823; reimpr. Calc. Jour. Nat. Hist. 4:
50. 1843; IV. 50, At the foot of Gunong Bunko inland from Bencoolen,
Sumatra. Also in Java; see C. B. Clarke, Monog. Phan. 5: 260. 1883.

C. bicolor Jack, op. cit. 27; reimpr. III. 47; IV. 47. Sumatra. There is a Jack
specimen in the Delessert herbarium, fide C. B. Clarke, op. cit. 242. Also in the
Malay Peninsula

C. carnosa Jack, op. cit. 30; reimpr. III. 51; IV. 51. No locality indicated but
probably .from Sumatra. Known only from Jack’s description; see
Clarke, op. cit.

c Feateaens Jack, op. cit. 31; reimpr. III. 51; IV. 51. No locality indicated,
but Jack’s specimen in the Delessert herbarium at Geneva is from Sumatra,
fide C. B. Clarke, op. cit. 205.

C. glabra Jack, op. cit. 28; reimpr. III. 49; IV. 49. Inland from Bencoolen,
Sumatra. Occurs also in Java, fide C. B. Clarke, op. cit. 245.

C. hirsuta Jack, op. cit. 27; reimpr. III. 48; IV. 48. Sumatra. Known only from
the type collection, there being a Jack specimen in the Delessert herbarium
at Geneva, fide C. B. Clarke, op. cit. 246.

C. incompta Jack, op. cit. 29; reimpr. III. 48; IV. 48. Sumatra, no locality
supe Known ee Jack’s description, fide C. B. Clarke, op. cit. 285;

extant specimen

: ee la Jack, o “cit 25. pl. 2, fig. 1, a-g; reimpr. III. 46; IV. 46.
Sumatra, no locality indicated, but Jack’s type (the only known collection)
in the Delessert herbarium is from Selebang, in the jurisdiction of Bencoolen,
Sumatra, fide C. B. Clarke, op. cit. 243.

C. maculata Jack, op. cit. 26; reimpr. III, 47; IV. 47. Sumatra. No definite
wre indicated. Known only from Jack’s description, fide C. B. Clarke, op.

me ot Tack, op. cit. 30; reimpr. III. 50; IV. 50. Sumatra, no definite mae
type not preserved, but represented by various Sumatran collections, fide C. B
Clarke, op. cit. ‘

C. rubiginosa Jack, op. cit. 32; reimpr. III. 52; IV. 52. No locality cited,
probably from Sumatra: type ‘unknown: see C. B. Clarke, op. cit. 285.

DIDYMOCARPUS Wallich.

D. barbata Jack, Trans. Linn. Soc. 14: 38. 1823; reimpr. Calc. Jour. Nat. Hist. 4:
57. 1843; IV. 57. Sumatra = Chirita hors fieldii R. Br. (1838); see C. B. Clarke,
Monog. Phan. 5: 123. 1883. Sumatra, Java. Here Jack’s specific name
should have been accepted by Clarke, but : is now invalidated in Chirita by
the different Chirita barbata eee (190

D. corniculata Jack, Mal. Misc. 1 (5): rir: reimpr. Hook. Bot. Misc. 2: 62.
1830; ITI. 55; IV. 55; V. 224; ic Jack, Trans. Linn. Soc. 14: 36. 1823.
Sumatra at Tapanuly. According to C. B. Clarke, DC. Monog. Phan. 5: 86.
1883, there is a Jack specimen in the Delessert herbarium at Geneva. Known
only from Sumatra.

D. crinita Jack, Mal. Misc. 1 is 1. 1820; reimpr. Hook. Bot. Misc. 2: 60. ea
TH, 53 TV. 533: V. 223%: et Jack, Trans. Linn. Soc. 14: 33. pl. 2, fig. 2
1823. Penang. A valid a Malay Peninsula, Sumatra, with ae in

1952] MERRILL, WILLIAM JACK’S GENERA AND SPECIES 223

Borneo; see C. B. Clarke, - cit. 93. A duplicate of Jack’s type is in the
Edinburgh herbarium; see pl.

D. elongata Jack, Trans. Linn. = 14: 37. 1823; reimpr. III. 56; IV. 56. Pulo
Bintangor, an island off the west coast of Sumatra = Didissandra elongata
(Jack) C. B. Clarke in DC. Monog. — 5: 67. pl. 7. 1883. Also in Borneo.

D. frutescens Jack, Mal. Misc. 1 (5); 5. 1820; reimpr. Hook. Bot. Misc. 2:
63.2350; “TTF, 58; -1V..:583°-V¥.-225; 2 Jack, ‘Trans, Linn. Soc. 14: 39. 1823.
Lae = Didissandra frutescens (Jack) C. B. Clarke, l.c. Malay Peninsula,
Sumatr

D: Sacad Jack ex Burkill, Jour. Straits Br. Roy. As. Soc. 73: 198. fig. 1.
1916, descr. abbr. Sumatra at Tapanuly = D. corniculata Jack, supra.

D. racemosa Jack, Trans. Linn. Soc. 14: 34. 1823; reimpr. III. 54; IV. 54.
West coast of Sumatra at Tapanuly. A species known only from Sumatra; see

B. Clarke, op. cit. 94.

D. reptans Jack, Mal. Misc. 1 (5): 3. 1820; reimpr. Hook. Bot. Misc. 2: 61.
1830; III. 55; IV. 55; V. 224; et Jack, Trans, Linn. Soc. 14: 35,. 1823.
Penang. Reported by C. B. Clarke, op. cit. 95, also from lower Burma and
from Java; widely distributed in the Malay Peninsula.

DRYOBALANOPS Gaertner f.

D. camphora Colebr. As. Res. 12: 535. 1816; Jack [App. Descr. Mal. Pl.
1820]; reimpr. Hook. Comp. Bot. Mag. 1: 264. 1836; III. 213; IV. 117.
West coast of Sumatra at Tapanuly = D. aromatica Gaertn. i (1805). Malay
Peninsula, Sumatra, Lingga, Borneo.

ELAEOCARPUS Linnaeus.

E. nitidus Jack, Mal. Misc. 1 (5): 41. 1820; reimpr. Hook. Bot. Misc. 2: 84.
1830; III. 224; IV. 128; V. 242. Penang. I accept Corner’s interpretation of
this species, Gard. Bull. Straits Settl. 10: 323. 1939, as he clearly demonstrated
that King’s earlier interpretation of 1891 was erroneous. I found in the British
Museum herbarium an unnamed Elaeocarpus labeled in Wallich’s handwriting
“Elaeocarpus e, Penang miscet Wm. Jack, 1819.” This is E. nitidus Jack as
interpreted by Corner and is unquestionably an isotype of Jack’s species.
Common, Malay Peninsula, Sumatra, Borneo; see Merrill, Jour. Arnold Arb.
32: 184. 1951, for synonymy.

ELODEA Jack, Mal. Misc. 2 (7): 21. 1822; reimpr. Hook. Jour. Bot. 1: 371.
1834; III. 208; IV. 112; V. 256, non Elodes Adanson (1763), nec Elodea
Juss. (1789) = Cratoxylon Blume (1825).

E. egyptica Jack, op. cit. 25; reimpr. Hook. Comp. Bot. Mag. 1: 154. 1835; III.
211; IV. 115; V. 272, in obs., sub Jxonanthes = Hypericum aegyptiocum

Linn.

E. formosa ig op. cit. 22; reimpr. Hook. Jour. Bot. 1: 374. 1834; III. 210;
IV. 114; V. 258. Sumatra = = Cratoxylon formosum (Jack) Dyer in Hook. f.
Fl. Brit. Ind. 1: 258. 1874; Corner, Gard. Bull. Straits Settl. 10: 28, 34. 1939,
cum syn. es Peninsula and Sumatra to Java, Borneo, the Philippines,
and the Moluccas. Corner, l.c., has definitely shown that my application of
the crn Cratoxylon pa arcane (Lour.) Blume to this species was
erroneous.

E. sumatrana Jack, op. cit. 22: reimpr. Hook. op. cit. 372; III. 209; IV. 113; V.
257. Pulo Nias, off the west coast of Sumatra = Cratoxylon ees
(Jack) Blume. See Corner, Gard. Bull. Straits Settl. 10: 27. 1939, for a

224 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxxm1

discussion of this species. He suggests that C. racemosum Blume (type from
Java) is its most likely synonym; to be compared, however, is C. clandestinum
Blume (type from Java), if de Voogt 1168 from Bencoolen, Sumatra, was
correctly named.

EMBELIA Burman f.

E. canescens Jack in Roxb, Fl. Ind. 2: 292. 1824, Penang. A well-understood
species now also known from the Malay Peninsula and Sumatra. Overlooked
by Griffith when he compiled the Jack descriptions in 1843.

ENCHIDIUM Jack,!® Mal. Misc. 2 (7): 89. 1822; reimpr. Hook. Comp. Bot.
Mag. 1: 257. 1836; III. 228; IV. 132; V. 291 = Trigonostemon Blume 1827
(Trigostemon Blume, 1825).

E. verticillatum Jack, op. cit. 90; reimpr. ll.cc. ‘‘Sumatra and the Malay Islands”

(T. indicus Muell.-Arg., 1865; Telogyne indica Baill., 1858). Malay Peninsula,
Penang, Sumatra. Jack’s actual type was from Sumatra, and is apparently no
longer extant. His addition ‘‘and the Malay Islands” was apparently made
because he thought that the Moluccan Abor spiculorum Rumph. Herb. Amb.
3: 167. pl. 106. 1743, represented his species; but what Rumphius illustrated
was a sterile specimen of what is clearly an Actinodaphne of the Lauraceae,
and is A. rumphi Blume.

EPITHINIA Jack, Mal. Misc. 1 (5): 12. 1820; reimpr. Hook. a Misc. 2:
67. 1830; III. 24; IV. 24; V. 228 = Scyphiphora Gaertn. f. (18

Ez. malayana Jack, Le; ; reimpr. ll.cc. Singapore = Scyphiphora Facoiiauacee
(Jack) Gaertn. f. (1805). A common and widely distributed species growing
within the influence of salt or brackish water in the Indo-Malaysian region.

EURYCOMA Jack, Mal. Misc. 2 (7): 44. 1822; reimpr. Roxb. Fl. Ind. 2: 307.
1824; reimpr. Hook. Comp. Bot. Mag. 1: 151. 1835; III. 168; IV. 72; V. 268.

E. longifolia Jack, op. cit. 45; reimpr. ll.cc. Tapanuly and Bencoolen, west coast
of Sumatara, and at Singapore. A small genus, this species common in parts
of Sumatra, Borneo, and the Malay Peninsula, extending to Siam and Indo-
China.

EUTHEMIS Jack, Mal. Misc. 1 (5): 15. 1820; reimpr. Roxb. Fl. Ind. 2: 203.
1824; reimpr. Hook. Bot. Misc. 2: 69. 1830; III. 200; IV. 104; V. 230
E. leucocarpa Jack, op. cit. 16: reimpr. lIl.cc. Singapore, Malay Peninsula,
orneo, and probably Sumatra.
E. minor Jack, op. cit. 16; reimpr. Roxb. op. cit. 304; II. 70; III. 201; IV. 105;
V. 231. Singapore. Widely distributed in the Malay Peninsula, Borneo, and
apparently also in Sumatra.

FAGRAEA Thunberg.

F. auriculata Jack, Mal. Misc. 2 (7): 82. 1822; reimpr. Hook. Comp. Bot. Mag.
1: 254. 1836; III. 29; IV. 29; V. 287, omn. sub F. carnosa Jack; Jack ex Roxb.
Fl. Ind. 2: 34. 1824, descr. Singapore and the west coast of Sumatra at

*8 Jack’s generic name has priority, but because about eighty binomials have been
ergot in Trigonostemon and only one in Enchidium, van Steenis, Fl. Males. Bull.
3: 74. 1948, has recommended that Blume’s name be conserved, which is manifestly
desirable.

1952] MERRILL, WILLIAM JACK’S GENERA AND SPECIES Z25

Tapanuly. Now recorded from the Malay Peninsula, Sumatra, Banka Billiton,
Borneo, Java, and Mindanao (F. epiphytica Elm.).

F. carnosa Jack, op. cit. 81 (sphalm, Fagroea); reimpr. Il.cc. Sumatra, near
Bencoolen. Known only from Sumatra; F. monantha Migq. (1857) is a

synonym.

F. racemosa Jack, op. cit. 82; reimpr. ll.cc., omn. mom. sub F. carnosa Jack;
Jack ex Roxb. FI. Ind. 2: 35. 1824, descr. Penang. A common species extend-
ing from Indo-China through the Malay Peninsula and Sumatra, Java, Borneo,
the Philippines southward to New Guinea. Fagraea volubilis Wall. in Roxb.
Fl. Ind. ed. 2, 2: 36. 1824, is sometimes erroneously listed as a Jack species.
It is a synonym of F. racemosa Jack, its type, a fruiting specimen sent to
Wallich by Jack from Bencoolen; but Wallich, who described it, expressed
doubt if it was distinct from F. racemosa Jack.

FICUS Linnaeus.

F. deltoidea Jack, Mal. Misc. 2 (7): 71. 1822; reimpr. Hook. Comp. Bot. Mag. 1:
222. 1836; III. 369; IV. 225; V. 282. Sumatra (F. diversifolia Blume, 1825).
A species with exceedingly variable leaves. Malay Peninsula, Sumatra, Java,
Borneo, and Palawan

F. ovoidea Jack, l.c.; reimpr. lIl.cc. Singapore, west coast of Sumatra, and
neighboring islands. Clearly only a form of F. deltoidea Jack.

F. rigida Jack, op. cit. 72; reimpr. Hook. Comp. Bot. Mag. 1: 222. 1836; III.
369; IV. 225; V. 282. Sumatra, no locality indicated (F. glaberrima Blume,
Bijdr. 457. 1825; King, Ann. Bot. Gard. Calcutta 1: 37. pl. 43. 1887; Koord.
& Val. Atlas Baumart. Java 4: fig. 710. 1916). Northern India to Burma,
Indo-China, southern China and Hainan, the Malay Peninsula, Sumatra
(Yates 741!), and Java. Jack’s description is an excellent one and can apply
only to this widely distributed, well-known species; his specific name is
valid and has priority.

FLACOURTIA Commerson.

F. inermis Roxb. Hort. Beng. 73. 1814, nom. nud., Pl. Coromand. 3: 16. pl. 222.
1819, Fl. Ind. ed. 2, 3: 833. 1832; Jack: Mal. Misc. 1 (1): 25. 1820; reimpr.
Hook. Bot. Misc. 1: 289. 1830; III. 230; IV. 134; V. 221. Jack’s material
was from Sumatra and Penang, and it seems to be evident that he correctly
interpreted Roxburgh’s species. Roxburgh said that his material came from
C

Jard. Bot. Buitenz. III. 7: 373. 1925, has critically considered the species,
and concluded that the tomi tomi or lobi lobi, as this cultivated fruit tree is
widely known, is an introduced species in the Moluccas, as it is in man

other parts of Malaysia, and further cited Reinwardt as recording the fact
that the species was common in the Moluccas in 1820, although it was ap-
parently unknown to Rumphius. He surmised that its introduction in Amboina
might have been between 1700 and 1800. It has been introduced into Ceylon,
India, and other tropical countries, and in Malaysia extends from the Malay
Peninsula to Sumatra, Java, Borneo, Celebes, the Moluccas, and New Guinea,
chiefly in cultivation and largely man-distributed. See Koord. & Val. Atlas
Baumart. Java 2: pl. 335. 1914 and Ochse, Fruits Dutch East Ind. 47. pl. 18.
1931.

226 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxx

GARDENIA Ellis

'G. anisophylla Jack in Roxb. FI. Ind. 2: 561, 1824. Penang, Singapore = Randia
anisophylla (Jack) Hook. f. Fl. Brit. Ind. 3: 114. 1880. This was published by
Roxburgh under Gardenia, not under Randia, as Hooker f. and King indicate.
Malay Peninsula, Borneo.

GLAPHYRIA Jack, Trans. Linn. Soc. 14: 128. 1823; reimpr. Calc. Jour. Nat.
Hist. 4: 306. 1843; IV. 162 = Leptospermum Forster (1776).

G. nitida Jack, l.c.; reimpr. Il.cc. Gunong Bunko or Sugarloaf Mountain, inland
rom Bencoolen, Sumatra = Leptospermum javanicum Blume (1826) (L.
commune Sm. var. javanica King). Widely distributed in Malaysia, Smith’s
species is Australian. Jack’s binomial antedates that of Blume, but his specific
name is payers in = ate lg by the Saati L. nitidum Hook. (1860).

G. sericea — op. cit. 129; reimpr. III. 307; IV. 163. “Found on Pulo singin
[ Pegang ]}, island on pre acu coast - “Sumatra.” Ex descr. =
pene fruticosum Forst., sensu lat. Indo-Malaysia to Polynesia.

GLOBBA Linnaeus.

G. ciliata iat Mal. Misc. 2 (7): 5. 1822; reimpr. Hook. Jour. Bot. 1: 361.
1834; III. 7; IV. 7; V. 248; K. Schum. Pflanzenr, 20 (IV, 46): 143. 1904.
Sauatak, a only from Jack’s description.

GMELINA Linnaeus.

G. villosa Roxb. Hort. Beng. 46. 1814, nom.; Roxb. ex Jack, Mal. Misc. 1 (1):
17. 1820, descr.; reimpr. Hook. Bot. Misc. 1: 284. ree III. 42; IV. 42; V.
218; Roxb. FI. Ind. ed. 2, 3: 86, 1832. Native of Sum — = Gmelina elliptica
Sm. (1810). Burma through Malaysia to the epee and the Moluccas
eastward to Palau.

GOMPHIA Schreber.

G. sumatrana Jack, Mal. Misc. 1 (5): 29. 1820; reimpr. Hook. Bot. Misc. 2: 77.
1830; ITI. 198; IV. 102; V. 237. Sumatra = Ouratea sumatrana (Jack) Gilg =
Ouratea angustifolia (Vahl) Baill. = Ouratea zeylanica (Lam.) Alst. in Trimen
Handb. Fl. Ceyl. 6: 42. 1931. India and Ceylon through Malaysia to the
Philippines and Celebes, represented by very many collections. The oldest
specific name is that of Lamarck which Alston accepted. The particular
Sumatra form is re by Gomphia sumatrana Jack as interpreted by
Planchon in Hook. 8: pl. 712. 1848. There are those who will perhaps not
be satisfied with ne mene designation here accepted, and certainly those who
will not accept the species as thus interpreted, sensu latiore. Thus Ridley, Kew
Bull. 1925: 79, 281. 1925, retained Gomphia as the generic name with G.
sumatrana Jack limited to Sumatra, and the Malay Peninsula form separated
as G. oblongifolia Ridl.

HALORRHAGIS Forster.

H. disticha Jack, Mal. Misc. 2 (7): 19. 1822; reimpr. cna wage Bot. 1: 371.
1834; III. 336; IV. 192; V. 256 (as Palaea ae Sum Singapore, and
shen parts of the Malay Archipelago = Anisophyllea disc (Jack) Baill.
(A. trapezoidalis Baill.). Malay Peninsula, Sumatra, Born

HEDYCARPUS Jack, Trans. Linn. Soc. 14: 118, 1823; reimpr. Calc. Jour. Nat.
Hist. 4: 184. 1843; IV. 88 = Baccaurea Lour. (1790).

1952] MERRILL, WILLIAM JACK’S GENERA AND SPECIES 227

H. malayanus Jack, l.c.; reimpr. III. 185; IV. 89. Sumatra = Baccaurea
malayana (Jack) King, quoad syn. Jack. Corner, Gard. Bull. Straits Settl. 10:
288. 1939, demonstrated rather convincingly that the Malay Peninsula form
referred here does not represent Jack’s species and that the latter stands as a
species known only from Jack’s description. Jack says that the fruit of bera
tampui ranks in point of taste and flavor with the lanséh (Lansium De
cum), which is one of the excellent Malayan fruits. There is an adage to the
effect that botanists never collect specimens from cultivated plants. I a
of no existing herbarium specimens which represent this Sumatran species.

HEDYCHIUM Koenig.

H. sumatranum Jack, Mal. Misc. 2 (7): 1. 1822; reimpr. Hook. Jour. Bot. 1:
358. 1834; III. 6; IV. 6; V. 246. West coast of Sumatra at Salumah. As
yet unplaced, being known only from Jack’s description. Allied to H.
collinum Ridl. of the Malay Peninsula?

HELOSPORA Jack, Trans. Linn. Soc. 14: 127. 1823; reimpr. III. 16; IV. 16 =
Timonius (Rumph.) DC., 1830, nom. conserv.

ens Jack, l.c. pl. 4, fig. 3; reimpr. Il.cc. Sumatra = Timonius flavescens ;

(Jack) Baker, Fl. Maurit. 144. 1877. Malay Peninsula, Sumatra, Borneo,

many collections. Timonius peduncularis Ridl. (1923) is a synonym. See
Boerl. Bull. Dép. Agr. Ind. Néerl. 26: 34. 1909.

HOYA Linnaeus.

H. *gracilis Jack ex Burkill, Jour. Straits Br. Roy. As. Soc, 73: 222, 225, fig. 2.
1916. Pulo Nias. This species was never described.

H. *grandiflora Jack ex Burkill, op. cit. 223, 225. Sumatra, west coast. Never
technically described, but the notes indicating that the flowers are two
inches in diameter, red shading into white, the whole plant hirsute, is
probably sufficient to place the species if one has access to Sumatran material.

e is invalidated by the earlier H. grandiflora Blume. It must be
closely allied to Hoya imperialis Lindl. of the Malay Peninsula and Borneo.

HYDNOPHYTUM Jack, Trans. Linn. Soc. 14: 124. 1823; reimpr. Calc. Jour.
Nat. Hist. 4: 21. 1843; IV. 21.

H. formicarum Jack, l.c.; reimpr. ll.cc. Sumatra; Griffith in 1843 added Malacca.
Its range is now given as Indo-China, Malay Peninsula, Sumatra, Java, Borneo,
and the Philippines.

*HYPSAGYNE Jack ex Burkill, Jour. Straits Br. Roy. As. Soc. 73: 219, 221, 247.
1916, nom. = Salacia Linnaeus

INGA Scopoli.

I. bubalina Jack, Mal. Mise. 2 (7): 77. 1822; reimpr. Hook. Comp. Bot. Mag. 1:
224. 1836; III. 162; IV. 66; V. 285. Sumatra = Pithecellobium (Pithecolo-
bium) bubalinum (Jack) Benth: Malay Peninsula.

. clypearia Jack, op. cit. 78; reimpr. ll.cc. Sumatra, at Bencoolen = Pithecello-
bium (Pithecolobium) clypearia (Jack) Benth. sipaane Peninsula and Sumatra
through Malaysia to the Philippines and the Moluccas

—

IXONANTHES Jack, Mal. Misc. 2 (7): 51. fe reimpr. Hook. Comp. Bot.
Mag. 1: 154. 1835; ITI. 211; IV. 115; V. 2
I. icosandra Jack, op. cit. 53; reimpr. ll.cc. Been: Sumatra. Throughout

228 JOURNAL OF THE ARNOLD ARBORETUM _[vot. xxxu1

the Malay Peninsula; see King, Jour. As. Soc. Beng. 62 (2): 191. 1893
(Mater. Fl. Mal. Pen. 2: 433) for synonymy and an amplified description.
I. reticulata Jack, op. cit. 51; reimpr. Hook. l.c.; III. 211; IV. 115; V
West coast of Sumatra at Tapanuly. See King, op. cit. 192, 434, for an
amplified description and synonymy. Most or all parts of the Malay Peninsula.

IXORA Linnaeus.

I. neriifolia Jack, Mal. Misc. 2 (7): 82. 1822; reimpr. Hook. Comp. Bot. Mag.
1: 254. 1836; III. 26; IV. 26; V. 288. West coast of Sumatra. Bremekamp,
Bull. Jard. Bot. Buitenz. III. 14: 241. 1937, limited the species to Sumatra,
citing various Sumatran collections, and indicated a Korthals specimen as the
lectotype.

I. pendula Jack, op. cit. 1 (5): 11. 1820; reimpr. Hook. Bot. Misc. 2: 66. 1830;
IIT. 25; IV. 25; V. 228. Penang. See Bremek. Bull. Jard. rag Buitenz, ITI.
14: 292. 1937, and Corner, Gard. Bull. Straits Settl. 11: a 1941 (1. opaca
Don, /. montana Ridl., I. candida Ridl., J. pendula Jack var. opaca Ridl.,
I. parkinsoniana Craib). Common in the Malay sk ti extending to Siam
and Sumatra.

JOHNIA Roxburgh = Salacia Linnaeus
J. *sumatrana Jack ex Burkill, Jour. Strails Br. Roy. As. Soc. 73: 221. 1916, nom.
West coast of Sumatra. This name unquestionably belongs with the named

Johnia, The species is very similar to Salacia er (Willd.) DC.

JONESIA Roxburgh.

J. declinata Jack, Mal. Misc. 2 Ste Ai 1822; reimpr. Hook. Comp. Bot. Mag
223. 1836; III. 161; IV. 283. Stim atra = Saraca declinata rth
Miq. Malay Peninsula, a. pees

KNEMA Loureiro.

K. glaucescens Jack, Mal. Misc. 2 (7) : 35. 1822; reimpr. Hook. Comp. Bot. Mag.
1: 148. 1835; III. 357; IV. 213; V. 263. West coast of Sumatra at Bencoolen
(Knema glauca Warb., 1897; Myristica glauca Blume, 1825: M. sumatrana
Blume, 1835). After an attentive comparison of Jack’s description with
Warburg’s excellent description and with herbarium material, I see no reason
for not accepting Jack’s earlier name for this widely distributed species.
Warburg placed Jack’s species as a doubtful synonym of Knema glauca
(Blume) Warb. Nova Acta Acad. Leop.-Carol. Nat. Cur. 68: 594. 1897
(Monog. Myrist. 594), but was apparently loath to displace Blume’s binomial
by the earlier one of Jack. At the end of his treatment he added a compiled
description of Jack’s species, p. 616, under the heading “species negligenda.”
His hesitancy in adopting Jack’s binomial was due to the fact that, as with
many of Jack’s Sumatran species, there is no extant type. But he cited the
following Sumatran collections, Forbes 2466, Beccari 532, Beccari s.n.,
Korthals, Teysmann, Junghukn; there are many more modern Sumatran
collections now available. I do not hesitate in accepting what is manifestly the
oldest binomial for this widely distributed Malaysian species. Malay Peninsula,
the Nicobar and Andaman Islands, Sumatra, Banka, Java, Borneo.

LAGERSTROEMIA Linnaeus.
L. floribunda Jack, Mal. Misc. 1 (5): 38. 1820; reimpr. Hook. Bot. Misc. 2:

1952] MERRILL, WILLIAM JACK’S GENERA AND SPECIES 229

82. 1830; III. 333; IV. 189; V. 241. Penang. Recorded from Burma, Siam,
Indo- China, and the Malay Peninsula.

LANSIUM (Rumph.) Correa, Ann. Mus. Hist. Nat. Paris 10: 157. pl. 10, fig. 1.

L.

1807; Jack, Trans. Linn. Soc. 14: 115, 1823; reimpr. Calc. Jour. Nat. Hist. 4:
187. 1843; IV. 91.

aqueum Jack, op. cit. 116; reimpr. op. cit. 189; IV. 92. The round-fruited
orm of the next species, indicated by Jack as “Var. 8 L. aqueum,” its Malay
name ayer ayer

L. *domesticum Correa, l.c.; Jack. op. cit. 115. pl. 4, fig. 1. 1823; reimpr. Calc.

Jour. Nat. Hist. 4: 188. pl. 4, fig. 2; IV. 92. Malay Islands. The commonly
cultivated fruit tree known as Cncse lanseh, lansone, duku, etc.

The Correa publication of the binomial Lansium domesticum (1807) is not
admitted in Index Kewensis. He depended on Rumphius for his generic
characters, as did Poiret when he accepted Lansium domesticum Correa in
Lam. Encyl. Suppl. 3: 299. 1813. If one wishes an older specific name it is
supplied by the validly published Melia parasitica Osbeck, Dagbok Ostind.
Resa 278. 1757, as his extant type at Stockholm has been examined and it is
an inflorescence of Lansium domesticum Correa.

asia ais Jack, Trans. Linn. Soc. 14: 125. 1823; reimpr. Calc. Jour. Nat.

. 4: 23. 1843; IV. 23.

5 ener Jack, i > reimpr. Il.cc. Inland from Bencoolen, Sumatra. Jack’s

a

ctual type is preserved in the Rijksherbarium, Leiden, this specimen, labeled
in his own handwriting, poten absolutely with his description, having been
acquired by Hasskarl in 1829. It is well matched by Rahmat Si Boea (Toroes)
1369, 6728, 7420 from rete Sumatra. It strongly resembles L. cyanocarpus
Jack (L. inaequalis Blume) Cae in having very different bracts. Its range,
ee than Sumatra, is unce

. cyanocarpus Jack, Le.: ney “iL cc. Tapanuly on the west coast of Sumatra

(L. inaequalis Blume, Bijdr. 996. 1826). This is Bakhuizen van den Brink’s
interpretation of the species, and I agree that the Javan L. inaequalis Blume
cannot be distinguished from the Sumatran one as described a few years
earlier by Jack. Rahmat si Boea 7042, 9466, 9992, 10020, all from Asahan,
Sumatra, agree with Jack’s excellent description. Malay Peninsula, Sumatra,
Java, Borneo
Lasianthus cyanocarpus auctt. plur. (non Jack) is a very different species,
which has been given a range from northern India to southern China and
Formosa, southward through Malaysia and the Philippines to New Guinea.
One of its rather numerous synonyms is L. oculus-cati Mig. Miquel himself
has cleared up the mystery, for he clearly states, Fl. Ind. Bat. 2: 315. 1857,
that Lasianthus oculus-cati Miq. was based on L. cyanocarpus Blume, Bijdr.
996. 1826, non Jack. In other words, Blume merely misinterpreted Jack’s
description and based his description of L. cyanocarpus on Javan material.
r this widely distributed Lasianthus cyanocarpus Blume, non Jack, I
propose Lasianthus hirsutus (Roxb.) comb. nov., typified by Triosteum
hirsutum Roxb. Hort. Beng. 68. 1814, nom. nud., Fl. Ind. 2: 180. 1824, descr.,
ed. 2, 1: 539. 1832, type from Chittagong. One might hesitate to accept this
name because of Roxburgh’s very short description but for the fact that when
ape tea Wight, Calc. Jour. Nat. Hist. 6: 501. 1846, was proposed
e for Triosteum hirsutum Roxb., Wight provided additional
cee ne Furthermore, Roxburgh’s species is represented in the

230 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxxur

British Museum herbarium by two excellent sheets, the type collection.
Britten, Jour. Bot. 47: 43. 1909, cleared up the confusion which had existed
up to that date. I have examined the Roxburgh specimens. They do not
represent Lasianthus cyanocarpus Jack as Britten thought, but rather L.
cyanocarpus sensu Blume et auctt. plur. (Z. oculus-cati Miq.). The character-
istic bracts are larger than in jaca bia the Malay Archipelago, the
largest ones being 5 cm. long and 2.5 wide. Other synonyms are L.
bracteatus Wight (1846), type from the Malay Peninsula, L. eh A cati Miq.
(1857), a new name for L. cyanocarpus sensu Blume, non Jack, type from
Java, L. laevicaulis Kurz (1875), type from the Nicobar Islands, and L
everettii Merr. (1908), type from the Philippines. Eastern India to Indo-
China and Hainan, through the Malay Archipelago and the Philippines to the
Moluccas and New Guinea.

LAURUS Linnaeus.

L. — A Mal. Misc. oo 7 33. 1822; reimpr. Hook. Comp. Bot. Mag.

1: 147. > JIT. 355; IV. 211; V. 262. ‘Su umatra, west coast, at Natal =
nae iocrasata (Jack) a. nov. (Dehaasia microcarpa Blume, Rumphia
1: 162. pl. 44. 1835; Haasia microcarpa Nees; H. incrassata Nees; Persea
incrassata Nees; Machilus incrassatus Nees). Malay Peninsula, Sumatra,
Java, Borneo.
L. parthenoxylon Jack, Mal. Misc. 1 (5): 28. 1820; reimpr. Hook. Bot. Misc. 2:
76. 1830; IIT. 354; IV. 210; V. 236. Sumatra = Cinnamomum parthenoxylon
(Jack) Meisn: in DC. Prods 15 (1): 26. 1864; poling Jour. As. Soc. Beng.
75 (1): 87. 1912 (Mater. Fl. Mal. Pen. 5: 87), cum syn. Burma to south-
eastern China southward through the Malay peer to Sumatra, Java,
Borneo, and Celebes

LECANANTHUS Jack, Mal. Misc. 2 (7): 83. 1822; reimpr. Roxb. FI. Ind. 2:
319. 1824; reimpr. Hook. Comp. Bot. Mag. 1: 254. 1836; III. 28; IV. 28; V.

288.
L. erubescens Jack, l.c.; reimpr. Il.cc. Inland from Bencoolen, Sumatra. Repre-
sented by many collections from the Malay Peninsula, Sumatra, and Borneo.

LEUCONOTIS Jack, te Linn, Soc. 14: 121. 1823; reimpr. Calc. Jour. Nat.
Hist. 4: 30. 1843; Ae
L. anceps Jack, l.c. pl. 4, fo 2; reimpr. ll.cc. pl. 4, fig. 2. Sumatra (L. eugenifolia
. , 1844; L. a Blume, 1849; Melodinus eugenifolius Wall. list no.
1616. 1829, nom. nud.). Malay Peninsula, Sumatra, Borneo. The cited
Sumatra collections are Teysmann 4053, Beccari 844, Curtis 3551, Forbes
1586, 2725, with at least a half dozen additional modern ones from that island.
It is interesting to note that when Griffith reprinted a s pee erie
in 1844, he added Malacca to the range of the species. I do not hesitate in
accepting Jack’s binomial, the type of the genus, . replace a later and
currently used L. eugenifolia A. DC., in spite of the fact that Jack’s type is
apparently not extant.

LEUCOPOGON R. Brown (1810) = es Smith (1793).

L. malayanum Jack, Mal. Misc. 1 (5): 1820; reimpr. Roxb. Fl. Ind. 2: 301.
1824; reimpr. Hook. Bot. Misc. - a 1830; It. 37: IV. 37; “V. 232:
Singapore = = Styphelia malayana (Jack) J. J. Sm. (S. malaica Spreng.; S.

1952] MERRILL, WILLIAM JACK’S GENERA AND SPECIES 231

malayica Druce). Common in various parts of the Malay Peninsula, occurring
also in Banca and Borneo.

gre Linnaeus.

L. *leptostachya Jack ex Wall. List no. 8585. 1848, nom. sub Bennettia R. Br.
et Hook. f. Fl. Brit. Ind. 1: 492. 1875, nom. in syn. Penang = Galearia jackiana
(R. Br.) Mig. = G. fulva (Tul.) Miq. fide Ridley. Jack’s Penang specimen
was distributed as Wallich 8585A. I suspect that Ridley is correct in eg
the binomial Galearia fulva (Tul.) Miq. Cremostachys fulva Tul. was pub-
lished in 1851; Bennettia jackiana R. Br. in 1852. Malay Peninsula, Price,
and Singapore.

LINOCIERA Swartz.

L. odorata Jack, Mal. ae 2 (7): 96. 1822; reimpr. Hook. Comp. Bot. Mag. 1:
759.1836: 111 33; 1V - V. 295. West coast of Sumatra at Natal, and on
Pulo Mosella. Known ca from Jack’s description. One suspects from the
localities cited that this was a low altitude species, perhaps from near the
seashore. Chionanthus litoreus Miq. Fl. Ind. Bot. Suppl. 559. 1862 =
Linociera litorea Knobl. (1894), type a Teysmann specimen from Siboga on
the west coast of Sumatra north of Natal, from its description is almost
certainly Jack’s species; but Miquel had only a fruiting specimen, and Jack's
lucid description was based on a flowering one. It is certainly not Limociera
purpurea Vahl nor L. dichotoma Wall., to both of which it has been reduced.

LORANTHUS Linnaeus.

L. coccineus Jack, Mal. Misc. 1 (1): 8. 1820; reimpr. Roxb. Fl. Ind. 2: 215.
1824; reimpr. Hook. Bot. Misc. 1: 278. pl. 58. 1830; III. 347; IV. 203; V. 213.
Singapore = Helixanthera coccinea (Jack) Danser, Bull. Jard. Bot. Buitenz.
III, 11: 374. 1931, cum syn. Indo-China to Burma, the Malay Peninsula,
Sumatra and Borneo. I suspect that the cited Horsfield “Java” specimen
actually came from Sumatra, as as ein botanized in Sumatra in 1818.
There are no actual Java specimens know

L. cylindricus Jack ex Roxb. Fl]. Ind. 2: 213. ane reimpr. Calc. Jour. Nat. Hist.

349. 1843; IV. 205. Sumatra = Relicanihera cylindrica (Jack) Danser,
op. cit. 377, cum syn. Burma to Indo-China, the Malay Peninsula, Sumatra,
Java, Borneo, and Celebes

L. ferrugineus Roxb. Hort. Beng. 87. 1814, nom.; Roxb. ex Jack, Misc. 1 (1):
9. 1820; reimpr. Roxb. Fl. Ind. 2: 208. 1824; reimpr. Hook. Bot. Misc. 1:
279, pl. 59. 1830; III. 348; IV. 204; V. 213; Roxb. Fl. Ind. 2: 207. 1824.
Roxburgh’s very short description was based on a Penang specimen; Jack's
ample and excellent one, reproduced by Wallich following that of Roxburgh in
1824, was based on Sumatran material. As Danser noted, Jack’s description
antedated that of Roxburgh, but Jack correctly credited the binomial to
Roxburgh = Scurrula ferruginea (Roxb.) Danser, op. cit. 432, cum syn.
Malay Peninsula to Sumatra, Borneo, Java, and Palawan.

L. incarnatus Jack ex Roxb. Fl. Ind. 2: 213. 1824; reimpr. Calc. Jour. Nat. Hist.

50. 1843: IV. 206. Pulo Nias = Dendrophthoé incarnata (Jack) Miq.;
Danser, op. cit. 411, cum syn. Known only from Sumatra and some of the
west coast islands.

L. patulus Jack ex Roxb. op. cit. 214; reimpr. IIT. 351; IV. 207. Inland from
Bencoolen, Sumatra = Macrosolen cochinchinensis (Lour.) Danser, op. cit.
279, cum syn. Northern India to southeastern China southward through the
Malay Peninsula to Sumatra, Java, Borneo, the Philippines, and Celebes.

232 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxxu1

L. retusus Jack ex Roxb. op. cit. 212; reimpr. III. 349; IV. 205. Singapore =
Macrosolen retusus (Jack) Danser, op. cit. 296. Malay Peninsula, Sumatra,
and Borneo. The Java record, based solely on a Lobb collection, is surely
erroneous, as this specimen came from either the Malay Peninsula or Borneo;
see Merrill, Philip. Jour. Sci. 10: Bot. 184, 1915, Enum. Philip. Pl. 4: 76. 1926.

LOXONIA Jack, Trans. Linn. Soc. 14: 40. 1823; reimpr. Calc. Jour. Nat. Hist. 4:
59. 1843; IV. 59

L. discolor Jack, l.c.; reimpr. ll.cc. Inland from Bencoolen, Sumatra (Loxophyl-
lum racemosum Blume, Bijdr. 751. 1826; Loxonia acuminata R. Br. in Benn.
Pl. Jav. Rar. 105. pl, 25, 1838). Sumatra, Java. I note in passing that Robert
Brown’s description and illustration of 1838 were based on material collected
by Horsfield in Sumatra in 1818 when he accompanied Sir Stamford Raffles
on a trip from Padang to the Menangaboo ih The account closes with
the statement: ‘He did not observe it in a.”

L. hirsuta Jack, op. cit. 41; reimpr. III. 60; W. 60. Inland from Bencoolen,
Sumatra = praec., fide C. B. Clarke in DC. Monog. Phan. 5: 158. 1883.
Jack apparently separated this from his L. discolor chiefly by its hirsute leaves
and branched inflorescences.

MANGIFERA Linnaeu

M. — 5 bye in Roxb FL Ind. 2: 441. 1824; reimpr. Calc. Jour. Nat. Hist. 4:
174. ; IV. 78. Sumatra. A species in part man-distributed (Java, Philip-
ee pine a native of the Malay Peninsula, Sumatra, and Borneo,
although in these regions sometimes also planted.

M. foetida Lour.; Jack ex Roxb. l.c.; reimpr. III. 174; IV. 78. Penang and
Singapore. Loureiro’s species was correctly interpreted by Jack, and Wallich
List 8488A is an actual Jack spec This is, in part, a man-distributed
species, now extending from a a Indo- China southward through the
Malay ‘Peninsula to Sumatra, Java, Borneo, Celebes, the Moluccas, and New
Guinea.

M. quadrifida Jack ex Roxb. op. cit. 440; reimpr. III. 173; IV. 77. Sumatra
“‘and other islands of the eastern Archipelago.” Wallich List no. 8489 is a Jack
ai from Penang. Now known from the Malay Peninsula, Sumatra, and

Born
M. *rubicunda Jack ex Burkill, Jour. Straits Br. Roy. As. Soc. 73: 152. 1916,
descr. abbr. Penang = M. foetida Lour., supra.

MELASTOMA Linnaeus.

M. alpestre Jack, Trans. Linn. Soc. 14: 20. pl. 1, fig. 3. 1823 (alpestris) ; tirade
Calc. Jour. Nat. Hist. 4: 330. pl. 1, fig. 3. 1844; IV. 186. On the summit of
Gunong Bunko or Sugarloaf Mountain, inland from Bencoolen, cia =
Medinilla alpestris (Jack) Blume, Flora 14: 514. 1831. Bakhuizen van den
Brink f., Rec. Trav. Bot. Néerl. 40: 182. 1943, placed this as a doubtful
synonym of Medinilla javanensis (Blume) Blume, which dates from 1826.
Should this prove to be correct, and I think it is, then Jack’s specific name will
replace that of Blume. It should be noted that the type of sage verrucosa
Baker f., which Bakhuizen van den Brink f. reduced to Blume’s species
without eatin was from Mount Dempo a short distance south - Gunong
Bunko. Sumatra, various collections; Java, many collections; Bali.

M. bracteatum Jack op. cit. 9 (bracteata); reimpr. IV. 320; V. 176. Penang =

1952] MERRILL, WILLIAM JACK’S GENERA AND SPECIES 233

Dissochaeta ee (Jack) Blume, ei 14: 495. 1831; Bakh. f., Rec. Trav.
Bot. Néerl. 40: . 1943, cum syn. Malay Peninsula, Sumatra, Borneo, and
perhaps Java.

M. decemfidum Roxb. Hort. Beng. 90. 1814, nom. nud.; Roxb. ex Jack, Trans.
Linn. Soc. 14: 6. 1823, descr.; reimpr. III. 317; IV. 173; Roxb. Fl. Ind. ed. 2,
2: 405 [406]. 1832. Penang = Melastoma sanguineum Sims, Bot. Mag. 48: pl.
2241. 1821. Burma to southeastern China southward to the Malay Peninsula,
Sumatra, Borneo, and Java.

M. erectum Jack, op. cit. 5 (erecta); reimpr. III. 316; IV. 172. Sumatra, west
coast at Tapanuly. This has been placed as a doubtful synonym of Melastoma
polyanthum Blume (1831). Bakhuizen van den Brink f., op. cit. 103, left it
as a species unknown to him, perhaps a villose form of M. polyanthum Blume.
Whenever collections from near the type locality become available, it will
probably be possible to place this species. Melastoma polyanthum Blume,
sensu lat., is credited with extending from India to southern China and
throughout Malaysia to northeastern Australia

M. ageloons Jack, op. cit. 10. pl. 1, fig. 2, a-b (anane reimpr. III. 321. pl. 2, fig.

a—b; IV, 177. Penang = Allomorphia exigua (Jack) Blume, Flora 14:
| 1831, See King, Jour. As. Soc. Beng. 69 (2): 10. 1900. Malay Peninsula
and, fide King, also in Sumatra (Forbes 3062); but Bakhuizen van den Brink

, Rec. Trav. Bot. Néerl. 40: 290. 1943, cites Forbes 3062 under A. magnifica
(Mia. ) Guill. (Sonerila magnifica Miq.) of Sumatra, from which one surmises
that the latter may prove to be a synonym of a PbS (Jack)
Blume, as is also Melastoma impuber Roxb. Fl. Ind. ed. : 405. 1832,
according to King. One should not be misled by Siar statement,
“native of Moluccas,” as he applied this term to material from parts of the
Malay Archipelago. Rahmat Si Boea 8620 from Asahan, Sumatra, matches
our Penang specimen (Henderson 35358) rather closely.

M. eximium Jack, op. cit. 17 (eximia); reimpr. III. 327; IV. 183. Gunong
Bunko or Sugarloaf Mountain, inland from Bencoolen = Medinilla eximia

(Jack) Blume, Flora 14: 515. 1831. A species known only from Jack’s distinctly

M. fallax Jack, op. cit. 13: reimpr. III. 323; IV. 179. Sumatra = Omphalopus
fallax (Jack) Naud. Ann. Sci. Nat. III. Da. 15: 277. 1851. Sumatra, Java,
Bali, a variety reported from New Guinea. For its extensive synonymy see

8.

M. glaucum Jack, op. cit. 15 (glauca); reimpr. III. 325; IV. 181. Penang =
Anplectrum glaucum Triana = Melastoma divaricatum Willd. = Anplectrum
divaricatum Triana = Diplectria divaricata (Willd.) O. Kuntze, Rev. Gen. PI.
246. 1891; Bakh. f. Rec. Trav. Bot. Néerl. 40: 200. 1943, cum syn. Siam
Malay Peninsula, Sumatra, Java, Borneo, Celebes, Moluccas, and New Guinea.

M. gracile Jack, op. cit. 18 (gracilis); reimpr. III. 324; IV. 180. Sumatra =
Dissochaeta gracilis (Jack) Blume in Flora 14: 498. 1831 = = Neodissochaeta
gracilis (Jack) Bakh. f. Rec. Trav. Bot. Néerl. 40: 137. 1943, cum syn. Siam
to the Malay Peninsula, Sumatra, Borneo, Java.

M. malabathricum sensu t Jack, op. cit. 4. pl. 1, fig. 1, a-g (malabathrica) ; reimpr.
III. 315, pl. 8, fig. 1; IV. 171, non Linn. “Abundant throughout Sumatra and
the Malay Islands” = - Melastoma polyanthum Blume. As currently interpreted
a collective species extending from India to southern China, through Malaysia
and the Philippines to New Guinea and Australia; see Bakhuizen van den
Brink f., Rec. Trav. Bot. Néerl. 40: 64. 1943, for its extraordinary synonymy.

234 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxu1

M. nemorosum Jack, op. cit. 8 (memorosa); reimpr. III. 319; IV, 175. Sumatra,
Pulo Nias = Marumia nemorosa (Jack) Blume, Flora 14: 505, 1831 =
Macrolenes nemorosa (Jack) Bakh, f. Rec. Trav. Bot. Néerl. 40: 206. 1943,
cum syn. Malay Peninsula, Sumatra, Borneo. Bakhuizen van den Brink f.,
op. cit. 26, 203, correctly, I believe, accepted Macrolenes Naudin as the
proper generic name for this group, because Marumia Blume (1831) is
invalidated by the earlier and totally different Marumia Reinwardt (1823,
1827), a synonym of Saurauia Willdenow.

M. obvolutum Jack, op. cit. 3 (obvoluta); reimpr. III. 314; IV. 170. West coast
of Sumatra at Tapanuly. Cogniaux, in DC. Monog. Phan. 7: 349. 1891, on the
basis of an examination of Jack’s type, which is preserved in the Delessert
Herbarium at Geneva, recognized this as a valid species, extending its range
to the Philippines M, homostegium Naud.). Pending a re-examination of
this extant type, I accept his conclusions rather than Bakhuizen van den
Brink’s reduction of it, op. cit. 80, to a variety of M. malabathricum Linn.
Sumatra, Borneo, Philippine

M. pallidum Jack, op. cit. 12 (pallida) : reimpr, III. 322; IV. 178. Malay Islands,
no definite locality indicated but probably Penang or Singapore = Dissochaeta
pallida (Jack) Blume, Flora 14: 500. 1831. Malay Peninsula, Banca; see
Bakhuizen van den Brink f. op. cit. 229 for synonymy and a complete
description.

M. pulverulentum Jack, op. cit. 19 (pulverulenta); reimpr. III. 329; IV. 185.
Singapore, Sumatra, and islands on the west coast of Sumatra = Pogonathera
pulverulenta (Jack) Blume, Flora 14: 521. 1831; Bakh. f. Rec. Trav. Bot
Néerl. 40: 128. 1943, cum syn. ihageed Peninsula, Sumatra, Java, Borneo,
Philippine Islands, southward to New ;

M. rotundifolium Jack, op. cit. 11 (otal Rig III. 321; IV. 177.
Musi region, inland from Bencoolen, Sum = Phyllagathis rotundifolia
(Jack) Blume, Flora 14: 507. 1831; Bakh. ey op. cit. 267. Siam, Malay
Peninsula, Sumatra.

M. rubicundum Jack, op. cit. 18 (rubicunda); reimpr. III. 328° IV. 184. Singa-
pore = Medinilla rubicunda (Jack) Blume, Flora 14: 512. 1831; Merrill,
Brittonia 4: 128. 1941 (M. hasseltii Blume, op. cit. 513). Maiey Peninsula;
Sumatra, Borneo, Java, Bali, central and southern Philippines, and Celebes.
Bakhuizen van den Brink f. Rec. Trav. Bot. Néerl. 40: 197. 1943, failed to
place Jack’s species (type from Singapore) probably because the species had
been confused with the distinctly different M. erythrophylla (Wall.) Lindl. of
India and Upper Burma. He made Medinilla hasseltii Blume a variety of the
older M. crassifolia (Blume) Blume, which dates from 1826. But Jack’s
specific name is the oldest one for this group.

M. stellulatum Jack, op. cit. 6 (stellulata); reimpr. 4: 318; IV. 174. West coast
of Sumatra, at Saloomah = Marumia stellulata (Jack) Blume, Flora 14: 505.
1831 = Macrolenes stellulata (Jack) Bakh. f. op. cit. 216, cum syn. Sumatra,
Borneo.

M. viminale Jack, Trans. Linn. Soc. 14: 16. 1823 (viminalis); reimpr, III. 327;
IV. 183. Sumatra, no definite locality indicated = Anplectrum viminale (Jack)
Triana, Trans. Linn. Soc. 28: 84. pl. 7, fig. 90 a. 1871 (Aplectrum viminale
Blume; Backeria viminalis Bakh. f. Rec. Trav. Bot. Néerl. 40: 133. 1943,14

“*4 plectrum Blume, Flora 14: 502. 1831, is meager by Aplectrum (Nutt.)
Torr. (1826). For this reason A. Gray, Bot. Wilkes U. S. Explor. Exped. 597, 1854,
correctly proposed the new generic name ec ermateg oe this Malaysian group. He

1952] MERRILL, WILLIAM JACK’S GENERA AND SPECIES 235

cum syn.). Sumatra, Billiton, with a variety, fide Bakhuizen van den Brink
(Anplectrum rostratum Blume), in Mentawi, Sumatra, Malay Peninsula, Java
and Borneo.

MELIA Linnaeus.

M. excelsa Jack, Mal. Misc. 1 (1): 12. 1820; reimpr. Hook. Bot. Misc. 1: 281.
1830; III. 190; IV. 94; V. 215. Penang. A valid species, but one not well
understood until recently. For a critical consideration and a detailed de-
scription see Corner, Gard. Bull. Straits Settl. 10: 263-267. fig. 1-2. 1939. A
duplicate of Jack’s type was distributed under Wallich List no. 1253, the entry
being ‘Penang, b. Jack,” but the specimen at Kew is very fragmentary, the
flowers all fallen, fide Airy Shaw in lit. Corner expressed the opinion that
the old trees observed by him at the Penang cemetery were the very ones
from which Jack’s material was taken, which may well be the case. I agree
with Mr. Corner, and Mr. Airy Shaw confirms this, that Azadirachta
integrifoliola Merr., type from Palawan, is the same as Jack’s species. I
have material from Sumatra (Bencoolen), Neth. Ind. For. Serv. 31664;
Borneo, Neth. Ind. For. Serv. 29263, 29414; and from British Malaya, Penang,
Md. Haniff 7586, Perak, Corner 31642, Selangor, Corner 31698, and various
Palawan collections distributed as Azadirachta integrifoliola Merr. which, I
believe, all represent Jack’s species. The range, Malay Peninsula, Sumatra,
Borneo, Palawan, Basilan, is a natural one. Because of its strictly pinnate
leaves this species impresses me as being anomalous in Melia

MEMECYLON Linnaeus.

M. coeruleum Jack, Mal. Misc. 1 (5): 26. 1820; reimpr. Hook. Bot. Misc. 2: 75.
1830 (caeruleum) ; III. 310; IV. 166; V. 235. Penang. In all or most provinces
of the Malay Peninsula; also in Sumatra. The credited Philippine range (M.
manillanum Naud.) was due to an erroneously localized collection, Naudin’s
type, Cuming 2322, being from Malacca

M. paniculatum Jack, op. cit. 2 (7): 62. 1822; reimpr. Hook. Comp. Bot. Mag. 1:
210, 1836« 211,.312* JV, 167; V. 27 Tapanuly and Pulo Bintangor (just
south of Padang), west coast of e atra. I interpret this, from Jack’s
excellent description (he failed to indicate whether the branchlets were
terete, subterete, or angled) as the same as M. costatum Miq. (1850), as
interpreted by Bakhuizen van den Brink, Rec. Trav. Bot. Néerl. 40: 345. 1943,

m syn., a very common, variable, and widely distributed species extending
from the Malay Peninsula (possibly from Siam) throughout Malaysia to the

gave no generic description, this being unnecessary as long as he cited A plectrum Blume
as the name-bringing synonym, as he did. Hence what Blume actually described fixed

a representative of another genus has no bearing on the case. Bakhuizen van den
Brink f., Rec. Trav. Bot. Néerl. 40: 130-146. 1943, apparently assuming that Gray
rou a new generic description, erroneously decided that the latter’s new generic
name was invalid, and in its place proposed a new generic name Backeria Bakh. f. to

n
A. rostratum Blume, A. viminale Blume, and A. stipulare Blume, also form the entire
basis of yao Bakh. f. At the same time he segregated from Anplectrum a certain
group of species for which he proposed a new generic name Neodissochacta Ba kh. f.
. ae had assigned the new generic name Backeria to this group, it could have been
ntained; as it is he merely added another generic synonym to Anplectrum A. Gray
pa eroerieee Blin ye

236 JOURNAL OF THE ARNOLD ARBORETUM _ [VvoL. xxxmr

Moluccas and represented by very many collections. Bakhuizen van den
Brink merely listed Jack’s species as one unknown to him (there is no extant
type). The Jack description is distinctly definite on as to the a
characters), a plant with oblong-ovate, obtusely acuminate leaves 7 to
inches long, with distinct nerves which unite into a line near the margins. :
we accept the definitely collective species as Bakhuizen van den Brink
interprets it, I see no reason why we should not also accept its oldest published
binomial.

MICROCOS Linnaeus.

M. glabra Jack, Mal. Misc. 1 (1): 14, 1820; reimpr. Hook. Bot. Misc. 1: 282.
1830; III. 222; IV. 126; V. 216. Carnicobar 1° Island = Microcos paniculata
Linn. India to the Nicobar Islands, Siam, Burma, and southern China south-
ward to the Malay Peninsula and Java.

M. tomentosa Smith; Jack, op. cit. 13; reimpr. Hook. op. cit. 28. pl. 60; III. 221;
IV : 216. Penang. Jack apparently interpreted Smith's species
correctly and correctly cited the very inadequately described and later
Grewia paniculata Roxb. as a synonym (type ie eee Siam and
Indo-China to the Malay Peninsula, Sumatra, and

MILLINGTONIA eaperay (1820); Jack, Mal. Misc. 2 (7): 29. 1822; Sone i
Hook, Jour. Bot. 1: . 1834; IIT. 180; IV. 84; V. 260, non ‘inn, f, (1781

M. sumatrana Jack, Mal. Misc. 2 (7): 30. 1822; reimpr. Hook. Jour. Bot. 1:
378. 1834; III. 181; IV. 85; V. 261. Pulo Nias off the west coast of Sumatra
= Meliosma sumatrana (Jack) Walp. (M. nitida Blume). Malay Peninsula,
Sumatra, Java, Borneo. Meliosma nitida Blume, which is currently placed
as a synonym of Jack’s species, may prove to be distinct.

MILNEA Roxburgh = Aglaia Loureiro.

M. montana Jack, Trans. Linn. Soc, 14: 118. 1823; reimpr. Calc. Jour. Nat. Hist.
4: 180. 1843; IV. 94. Near Bencoolen, Sumatra. Clearly an Aglaia, but as
yet not associated with any other described species. Jack’s rather good

description follows his consideration of Lansium, and at the very end his
binomial appears thus: “if admitted as a separate Sai the nes will
constitute a second species .. . . and may be denominated M. montana.’

MIMOSA Linnaeus.

M. jiringa Jack, Mal. Miscel. 1 (1): 14. 1820; reimpr. Hook. Bot. Misc. 1: 282.
1830; III. 161; IV. 66; V. 285 = Pithecellobium (Pithecolobium) *jiringa
apr Prain, Jour. As. Soc. Beng. 66 (2): 267. 1897 (Mater. Fl. Mal. Penin.

267), in obs. Penang, Malacca, Tenasserim and the Malay Peninsula,
pean Borneo, Java (mostly planted). See Merr. Philip. Jour. Sci. 14: 243.
1919; Contr. Arnold Arb. 8: 72. 1934. The still earlier binomials Mimosa
koeringa Roxb. Hort. Beng. 40. 1814 and M. djiringa Roxb. op. cit. 93 are
nomina nuda
18 Car Nicobar is the most northern island of the Nicobar group, north of Sumatra,

which Jack apparently visited either on his voyage from Calcutta to Penang, or on
his one trip from Bencoolen to Calcutta and return. The basis of Connarus ? jackianus

Wall. List no. 8552 = Cupania jackiana Hiern = Lepidopetalum jackianum Radlk. was

latter, being a nomen nudum, does not, or course, invalidate Schellenberg’s later name.

1952] MERRILL, WILLIAM JACK’S GENERA AND SPECIES 237

MONOCERA Jack, Mal. Misc. 1 (5): 42. 1820; reimpr. Hook. Bot. Misc. 2: 85.
1830; III. 225; IV. 129; V. 243 = Elaeocarpus Linn. sect. Monocera (Jack)
Benth.

M. ferruginea Jack, op. cit. 44; reimpr. Hook. op. cit. 86; III. 226; IV. 130; V.
244. Singapore = Elaeocarpus ferrugineus (Jack) Steud. Nomencl. ed. 2,
545. 1840 (E. jackianus Wall.; E. borneensis Knuth). Malay Peninsula,
Borneo. See Corner, Gard. Bull. Straits Settl. 10: 319. 1939; Airy Shaw, Kew
Bull. 1949: 165. 1949; Merr. Jour. Arnold Arb. 32: 180. 1951.

M. petiolata Jack, op. cit. 43; reimpr. Hook. op. cit. 86; III. 226; IV. 130; V.
243. Penang = Elaeocarpus petiolatus (Jack) Wall. List no. 2673. 1829; A.
Gray, Bot. Wilkes U. S. Expl. Exped. 1: 203. 1854. The Wallich List entry
is merely “2673 Elaeocarpus (Monoceros) petiolata Jack — Hb. 1824 Penang
1822,” so there is a possibility of a quibble as to whether or not this constitutes
publication under Elaeocarpus. Malay Peninsula, Sumatra.

MORINDA Linnaeus.

M. polysperma Jack, Mal. Misc. 1 (5): 14. 1820; reimpr. Roxb. Fl. Ind. 2: 204.
1824; reimpr. Hook. Bot. Misc. 2: 68. 1830; III. 20; IV. 20; V. 229. Singapore
= Lucinaea polysperma (Jack) K. Schum. (L. morindae DC. Prodr. 4:
368 (1830). Jack suggested that his new species of Morinda might represent
a separate genus. It is the sole basis of the genus Lucinaea DC., described
ten years later. Malay Peninsula, Sumatra, Borneo.

M. aeons Pigs op. cit. 13; reimpr. Roxb. op. cit. 203; reimpr. Hook. op. cit.
67; III PORN lot 229, Native of the Malay Islands (probably ba
or sae ote) = == Morinda umbellata Linn. sensu lat. India and Ceylon to
southern China, through Malaysia and the Philippines to northeastern Australia
(as a collective species).

MURRAYA Koenig ex Linnaeus, Mant. 2: 558. 1771 (Murraea) ; Murr. Syst. ed.
13, 331. 1774, nom. conserv. halen Linn., 1767).

M. paniculata (Linn.) Jack, Mal. Misc. 1 (5): 3 31. 1820; reimpr. Hook. Bot. Misc.
Z 1830; III. 191; IV. 95; V. 238 (Chalcas paniculata Linn.). Jack’s
description was apparently based on material from Penang or Singapore, he
citing Chalcas paniculata Lour. (which is Chalcas paniculata Linn.) and
Camunium Rumph. Herb. Amb. 5: 26. pl. 17. 1747, which also represents the
Linnaean species. A common, variable, and widely distributed Malaysian
species,

MYRMECODIA Jack, Trans. Linn. Soc. 14: 122. 1823; reimpr. Calc. Jour. Nat.
Hist. 4: 20. 1843; IV. 20.

M. tuberosa Jack, op. cit. 123; reimpr. Il.cc. Pulu Nias, off the west coast of
Sumatra. See Beccari, Malesia 2: 99. pl. 13, 14. 1884. Malay Peninsula, Java,
Borneo.

NEPENTHES Linnaeus; Jack [App. Descr. Mal. Pl. 20, 1820]; reimpr. Jack
ex Hook. Comp. Bot. Mag. 1: 269. 1836; III. 362; IV. 222.

N. ampullaria Jack [App. Descr. Mal. Pl. 23 1820]; reimpr. Jack ex Hook. op.
cit. 271; III. 366; IV. 222. Singapore and on Bintang Island. See Danser,
Bull, Jard. Bot. Buitenz. III. 9: 265-270. 1938. Malay Peninsula, Sumatra,
Borneo, New Guinea, very many collections. Bintang (Bintan) Island is Hie
the Rhio Archipelago, southeast of Singapore. There are two sheets from

238 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxx

Jack in the type collection in the herbarium of the British Museum (Natural
History

N. distiliatoria sensu Jack [App. ee Mal. Pl. 23. 1820]; eee Jack ex
Hook. l.c.; II. 368; IV. 224 n Linn. Singapore, Malacca = N. gracilis
Korth. Malay Peninsula, Suma i gee Celebes; see Sar op. cit. 2
for details. The Linnaean species is confined to Ceylon. At the British
Museum are two fragmentary specimens from Jack, one labeled “mixed with
distillateria,’ which is N. albomarginata Lobb, the other (sterile) ZL. gracilis
Korth.

N. phyllamphora Willd.; Jack [App. Descr. Mal. Pl. 23. 1820]; — Jack
ex Hook. lL.c.; III. 367 > IV. 223. West coast of Sumatra at Benco = 1,
mirabilis (Lour.) Druce, Rep. British Exch. Club 1916: 637. ine ge
Merr. Interpret. Herb. Amb. 242. 1917 (August). Southeastern China and
Indo-China through the Malay Peninsula to Sumatra, Borneo, Java, Mindanao,
Celebes, and the Moluccas to Palau, New Guinea, and northeastern Australia.

N. rafflesiana Jack [App. Descr. Mal. Pl. 21. 1820]; reimpr. Jack ex Hook. op.
cit. 270; III. 364; IV. 220. Singapore. See Danser, op. cit. 357-361. Malay
Peninsula, Sumatra, Borneo, very many collections. There are two Jack sheets
in the type penile British Museum (Nat. Hist.) herbarium, inscribed “No.
3 Nepenthes si nova sit Rafflesiana from Singapore. Dr. Jack.” These Jack
Singapore Nepenthes specimens were manifestly sent by him to Robert Brown
in London, supplementing a larger lot sent to him from Penang previous to
Jack’s departure for Singapore. The highest number noted in this sending of
Singapore plants is four.

tpicnieues Linnaeus.

N. lappaceum Linn. Mant. 2: 566. 1771; Jack, Mal. Misc. 1 (1): 10. 1820;
reimpr. Hook. Bot. Misc. 1: 279. 1830; III. 183; IV. 87; V. 214. “Frequent
throughout the Malay countries and islands.” Jack correctly interpreted the
rambutan, which is the Linnaean species. It is one of the better of the culti-
vated fruit trees of Malaysia

*NEUROPTERIS Jack ex Burkill, Jour. Straits Br. Roy. As. Soc. 73: 216.
1916, nom. = Neuropeltis Wall.

OCTAS Jack, Mal. Misc. 2 (7): 64. 1822; reimpr. Hook. Comp. Bot. Mag. 1:
219. 1836; III. 340; IV. 196; V. 278 = Zlex Linn.; see Hallier f. Rec. Trav.
Bot. Néerl. 15: 66. 1918.

O. spicata Jack, l.c.; reimpr. Il.cc. West coast of Sumatra at Tapanuly = Jlex
spicata Blume, Bijdr. 1149, 1826. Jack and Blume independently selected the
same specific name, for Blume’s binomial in /lex was not based on Jack’s
earlier one. Jack’s ample and lucid description agrees entirely with the
characters of Blume’s species, this being, in //ex, an unusually sharply defined

alay Peninsula, Sumatra, Java, Borneo, New Guinea, many individual
collections available.

OPHIORRHIZA Linnaeus.

O. heterophylla Jack, Mal. Misc. 2 (7): 85. 1822; reimpr. Roxb. Fl. Ind. 2: 546.
1824; reimpr. Hook. Comp. Bot. Mag. 1: 255. 1836; III. 17; IV. 17; V. 289.
Sumatra, igen probably from near Bencoolen. Known only from Jack’s
descriptio

O. tomentosa a lack in Roxb. Fl. Ind. 2: 246. 1824. Penang and Sumatra; see

1952] MERRILL, WILLIAM JACK’S GENERA AND SPECIES 239

King, Jour. As. Soc. Beng. 72 (2): 176. 1903 (Mater. FI. ae ei 4: 66)
for a more ample description. Malay Peninsula, Penang, Sumat

*PATISNA Jack ex Burkill, Jour. Straits Br. Roy. As. Soc. 73: 196, 218, 255,
1916, nom. = Urophyllum Wall. See p. 248.
P. *glabra Jack ex Burkill, l.c. nom. = Urophyllum glabrum Wall.

PERONEMA Jack, Mal. Misc. 2 (7): 46. 1822; reimpr. Hook. Comp. Bot. Mag.
1: 152. 1835; IIT. 41; IV. 41; 269.

P. canescens Jack, op. cit. 47; reimpr. ll.cc. Sumatra. A monotypic genus. Malay
Peninsula, Sumatra, Bored, Java.

PETROCARYA Schreber (1789) = Parinari Aublet!® (1775) (Parinarium
Jussieu, 1789).

P. excelsa Jack, Mal. Misc. 2 (7): 66. 1822; reimpr. Hook. Comp. Bot. Mag. 1:
220. 1836; III. 164; IV. 68; V. 279. No locality cited but probably from
the west coast of Sumatra = = Parinari jackiana Benth. (1849), as Parinarium.
A species apparently known only from Jack’s description.

P. sumatrana Jack. op. cit. 67; reimpr. Hook. op. cit. 221; III. 165; IV. 69; V.
280 [Sumatra] = Parinari sumatrana (Jack) Benth. in Hook. Niger Fl. 335.
1849, as Parinarium. For a very full description based wholly on Sumatra
specimens see Blume, Mus. Bot. Lugd.-Bat. 2: 97. 1856 (P. costatum Blume
ex Miq. Fl. Ind. Bat. 1 (1): 254. 1855). Malay Peninsula, Sumatra, Borneo,
and Java.

PHALERIA Jack, Mal. Misc. 2 (7): 59. 1822; reimpr. Hook. Comp. Bot. Mag.
1: 156. 1835; III. 353; IV. 209; V. 276 (Drymispermum Reinw., 1828).

P. capitata Jack, l.c.; reimpr. Hook. l.c.; III. 354; IV. 209; V. 276. Sumatra.
This is the type of the genus, re phaleria Meisn. in DC. Prodr. 14:
604. 1867, being a synonym. ave no Sumatra specimens which I can
definitely refer to Jack’s species. See ee & Valeton ;
Landb. 18: 41. 1914 (Bijdr. Boomsoort. Java 13: 41) fora detailed description
based largely on Java material.

PHYTEUMA Linnaeus.

P. begonifolium Roxb. Hort. Beng. 85. 1814, nom.; Roxb. ex Jack Malay Misc. 1
(1): 5. 1820; reimpr. Hook. Bot. Misc. 1: 276. pl. 57. 1830; III. 34; IV. 34;
V. 212; Roxb. Fl. Ind. 2: 108. 1824 (descr. Jack reimpr. 109). Penang =
Pentaphragma begonifolium Wall. List no. 1313. 1829, “Pentaphragma
begonifolium Wall. Phyteuma Roxb. Penang. 1822;” G. Don, Gen. Syst. 3:
731. 1834. A well- ee species of the Malay Peninsula occurring also in
Sumatra, Siam, and Mergui. Jack strongly suggested that Roxburgh’s generic
designation was erroneous and that a new genus might be represented, which
proved to be the case

PIERARDIA Roxburgh, Hort. Beng. 28. 1814, nom. nud.; Roxb. ex Jack, Trans.
Linn. Soc. 14: 119. 1823, descr.; reimpr. Calc. Jour. Nat. Hist. 4: 186. 1843;
IV. 90; Roxb. Fl. Ind. ed. 2, 2: 254. 1832 = Baccaurea Loureiro (1790).

P. dulcis Jack, Trans. Linn. Soc. 14: 120. 1823, reimpr. III. 186; IV. 90. Sumatra,
at Bencoolen, where Jack says it was known as bua choopa, and was

** The original Parinari Aublet (1775) must be accepted unless the Latinized form
Parinarium Juss. (1789) be officially conserved.

240 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxxim

abundant. J. J. Smith, Meded. Dep. Landbouw 10: 21. 1910 (Koord. & Val.
Bijdr. Boomsoort. Java 12: 21) states that it is cultivated in Java, but no
more complete description than the original one of Jack seems to have been
published. This is apparently a planted fruit tree, and perhaps the practical
non-existence of herbarium material is but a reflection of the often repeated
statement that botanists (and often emai never bother to prepare
specimens from cultivated plants. Jack expressed the opinion that sean s
excellent figure of rambeh (Hist. Sumatra A. 6; in ed. 3, 1811 it is pl. 8) w

but a variety of his taxon. He noted that at the time (1819-22) the oa
rambeh occurred in the Malay Peninsula, but not in Bencoolen, while choopa
was abundant at Bencoolen but was not found in the Peninsula.

PITTOSPORUM Banks.

P. serrulatum Jack ex Roxb. FI. Ind. ed. 2, 2: 401. 1824, Sia mab serrulata;’
P. ? serrulatum Jack ex Griff, Calc. Jour. Nat. Hist. pr.
IV. 99. Penang = Rinorea lanceolata (Roxb.) O. Kuntze re lanceolata
Roxb.; Celastrus pauciflora Wall. ; Pentaloba lanceolata Wall.; Alsodeia
lanceolata Oudem.). A species still ‘Tose only from Penang.

POSOQUERIA Aublet

P. *anisophylla Tack © ex Burkill, hn Straits Br. Roy. As. Soc. 73: 196, 220.
1916, nom. nud. Penang, Sumat = Gardenia pellet Jack ex Roxb.
supra, p. 226 = Randia anisophylla “(Jack) Hook. f.

PSILOBIUM Jack, Mal. Misc. 2 (7): 84. 1822; reimpr. Roxb. Fl. Ind. 2: 320.
1824; reimpr. Hook. Comp. Bot. Mag. 1: 255. 1836; III. 27; IV. 27; V. 289 =
Acranthera Arnott ex Meissner (1838) .17

P. nutans Jack, op. cit. 84; reimpr. lIl.cc. West coast of Sumatra, inland from
Bencoolen. A species known from Jack’s description, as yet not safely
associated with any described species of Acranthera; Bremekamp suggests
a species of his subgenus Androtropis (R. Br.) Bremek. It may well be that
Brooks 6681 from the vicinity of Bencoolen (not seen) really represents
Jack’s species, as Ridley thought, Kew Bull. 1925: 84. 1925; see Bremekamp,
Jour. Arnold Arb. 28: 263. 1947.

P. tomentosum Jack, Mal. Misc. 2 (7): iii. 1822; reimpr. Roxb. Fl. Ind. 2: 321.
1824; Calc. Jour. Nat. Hist. 4: 28. 1843; IV. 28; V. 295. West coast of
Sumatra at Kataun. As yet not associated with any described species of
Acranthera. In any case Jack’s specific name is preoccupied in that genus by
the different A. tomentosa R. Br

PSYCHOTRIA Linnaeus.

P. malayana Jack, Mal. Misc. 1 (1): 3. 1820; reimpr. Hook. Bot. Misc. 1: 275.
1830 (excl. syn. P. aurantiaca Wall.); III. 26; IV. 26; V. 228. Penang. (P.
stipulacea Wall. ex Roxb. Fl. Ind. 2: 164. 1824). As explained by me in
Webbia 7: 321-324. 1950, what Jack actually described in 1820 was the
species Wallich characterized four years later as P. stipulacea Wall. Psychotria
malayana has been given a very wide range in Malaya by modern authors,
but what is so named in all herbaria and described in all texts is not at all

remekamp, Jour. Arnold Arb. 28: 261-265. 1947, has shown that Psilobium
Jack (1822), hitherto not properly understood, is the same as the later Acranthera
Arnott ex Meissner (1838). As there are now 35 Acranthera species, he suggested that
the latter name be conserved against that of Jack.

1952] MERRILL, WILLIAM JACK’S GENERA AND SPECIES 241

the species Jack characterized. This misinterpretation was due to Wallich’s
original error in reducing P. malayana Jack to P. aurantiaca Wall., which
was unfortunately accepted by subsequent authors. The species is known
from many parts of the Malay Peninsula and apparently occurs in Sumatra.

I RA Jack, Mal. Misc. 2 (7): 60. 1822; reimpr. Hook. Comp. Bot.

Mag. 1: 157. 1835; III. 309; IV. 165; V. 276.

P. capitellata Jack, Mal. Misc. 2 (7): iii. 1822; reimpr. Calc. Jour. Nat. Hist.
4: 310. 1843; IV. 166; V. 295. West coast of Sumatra at Moco Moco; see
Bakhuizen van den Brink, f., Rec. Trav. Bot. Néerl. 40: 326. 1943, who
followed King in making this a variety of P. coerulescens Jack. Malay
Peninsula, Sumatra, Lingga, Borneo, and fide Mansfeld, New Guinea.

Mag. 1: 157. 1835; III. 309; IV. 165; V. 277. Penang. See Bakhuizen van
den Brink f., Rec. Trav. Bot. Néerl. 40: 324. 1943. Burma and the Andaman
Islands to Tndo- China, the Malay igerace, Sumatra, Lingga, and Borneo.
P. echinata Jack, Mal. Misc. 2 (7): iii. 1822; reimpr. Calc. Jour. Nat. Hist. 4:
310. 1843; IV. 166; V. 295. es coast Cc Sumatra at Kataun = Kibessia
echinata (Jack) Cogn. in Pl. Monog. Phan. 7: 1108. 1891. This may or may
not be the same as Kibessia azurea (Blume) DC. of Sumatra and Java. King
in 1900 followed Cogniaux. Jack’s very inadequate description is merely
“pedunculis axillaribus terminalibusque, calycibus ovariisque echinatis. A
large tree found at Kataun. The leaves are 3-nerved in all species.” In a
footnote Griffith in his 1843 reproduction of Jack’s descriptions indicates

Sumatra, the modern Dutch spelling being Ketaoen. I suspect, but can
scarcely prove, that this is Kibessia azurea (Blume) DC. as interpreted
by Bakhuizen van den Brink f. in 1943, this binomial dating from 1826, four
years later than that of Jack.

PYRRHANTHUS Jack, Mal. Misc. 2 (7): 57. 1822; reimpr. Hook. Comp. Bot.
Mag. 1: 156. 1835; III. 337; IV. 193; V. 274 = Lumnitzera Willd. (1803).
P. *flammeus Jack ex Burkill, Jour. Straits Br. Roy. As. Soc. 73: 205. 1916,

nom. eq.

P. littoreus Jack, l.c.; reimpr. Hook. |.c.; III. 337; aE 193; V. 275. Sumatra,
Malay Peninsula = Lumnitzera littorea (Jack) Voigt (L. coccinea Wight &
Arn.). Common within the influence of salt water. “Tndia through Malaysia to
tropical Australia and Polynesia.

QUERCUS Linnaeus.

Q. racemosa Jack, Mal. Misc. 2 (7): 86. 1822; reimpr. Hook. Comp. Bot. Mag. 1:
255. 1836; III. 370; IV. 226; V. 289, non Lam. (1785). Sumatra. Currently
reduced to Quercus spicata Sm. = Lithocarpus spicata Rehd. & Wils. as that
species is now interpreted, and it perhaps belongs here. In any case Jack’s
specific name is an invalid one. Smith’s species, sensu latiore, ranges from
the Himalayan — eastward to southern China and southward to Sumatra,
Borneo, and Jav

Q. Ceara Jack, ee reimpr. Hook. op. cit. 255; III. 371; IV. 227; V. 290.

= = ithncarcus urceolaris (Jac comb. nov. Ouerc Cus olivoneurs
Korth, 1844; Q. eichleri Wenzig, 1886; Synaedrys eichleri Koidz., 1916;
Pasania eichleri Gamble, 1914; Lithocarpus eichleri, A. Camus, 1931, et Les

242 JOURNAL OF THE ARNOLD ARBORETUM _[VoL. xxxiI

Chénes 3: 71. pl. 395. 1948). King mentioned Miquel’s suggestion that this
might be the same as Quercus lamponga Miq., but the descriptions do not
agree. I find, however, that in all essentials the descriptions of Q. urceolaris
Jack and Q. eichleri Wenzig do agree. All types involved were from Sumatra.
The species occurs also in the Malay Peninsula and Borneo.

RAFFLESIA Jack [App. Descr. Mal. Pl. 1. 1820]; Jack ex Hook. Comp. Bot.
Mag. 1: 259. pl. 14. 1836; III. 260; IV. 216 = Rafflesia R. Brown (1821).

R. Noss omen Marsden ex one in Burkill, Jour. Straits Br. Roy. As. Soc. 73:

. 1916, nom. Sumatra = s

R. we Jack [App. Descr. Mal. PL i. 1820]; Jack ex Hook. |.c.; III. 260; IV.
216. Interior of Sumatra, particularly in the forests of Passummah Ulu
Manna = Rafflesia arnoldi R. Br. (1821). A remarkable species known only
from Sumatra. Jack’s description was withdrawn because of R. Brown’s
treatment of the taxon in 1821, both descriptions having been based on the
same collection, which was made by Sir Stamford Raffles.

RAUWOLFIA Linnaeus.

R. sumatrana Jack, Mal. Misc. 1 (5): 22. 1820; Roxb. Fl. Ind. 2: 543, 1824;
reimpr. Hook. Bot. Misc. 2: 73. 1830; III. 31; IV. 31; V. 233. Sumatra,
frequent near Bencoolen. A valid species amply described by King & Gamble,
Jour. As. Soc. Beng. 74 (2): 424. 1907 (Mater. Fl. Mal. Pen. 4: 634). Also
in-the Malay Peninsula; and to be compared is the Philippine-Borneo R.
samarensis Merr.

RHIZOPHORA Linnaeus.

R. *caryophylloides Jack, Mal. Misc. 1 (5): 34. 1820; reimpr. Hook. Bot. Misc. 2:
80. 1830; IIT. 334; IV. 190; V. 239. Singapore, Penang = R. caryophylloides
Burm. f. Fl. Ind. 109. 1768 = Bruguiera cylindrica (Linn.) Blume. In tidal
forests throughout the Indo-Malaysian region. Jack, in proposing his new
binomial in 1820, overlooked Burman’s earlier use of the same name in 1768
to represent the same species. Mangium caryophylloides Rumph, (1743),
cited by both, provided the specific name.

RHODAMNIA Jack, Mal. Misc. 2 (7): 48. 1822; reimpr. Hook. Comp. Bot.
Mag. 1: 153. 1835; III. 307; IV. 163; V. 270
R. cinerea Jack, l.c.; reimpr. ll.cc. West coast of ‘Sumatra, frequent. Rhodamnia
Jack is a small genus, its limits well understood, and probably Jack’s specific
name should be retained for the common Malaysian form (Malay Peninsula,
Sumatra, Borneo, Java). Corner, Gard. Bull. Straits Settl. 10: 272. 1939,
took exception to my acceptance of Myrtus trinervia Lour. (1790) as a
synonym for this species [R. trinervia (Sm.) Bl.], because Loureiro definitely
described the inflorescences as racemose; yet Clemens 3689, from what must
be the type locality, has racemes ye cm. in length. Yet in other characters
this closely approximates our rich series of Sumatran specimens. The oldest
valid name for this is Rhodamnia ee (Lour.) Merr. & Perry, Jour.
Arnold Arb. 19: 195. 1938, of which R. siamensis Craib is a synonym. But
Corner overlooked C. T. White’s conclusions, Blumea, Suppl. 1: 215. 1937,
that the Australian form is specifically distinct from the eae one. As
mith’s species was based wholly on Australian specim Rhodamnia
wiursio (Sm.) Blume is not the proper name for the et that Jack

1952] MERRILL, WILLIAM JACK’S GENERA AND SPECIES 243

characterized; for this common form with axillary solitary or fascicled flowers,
Jack’s specific name is the earliest available one. The group is in need of a
critical revision.

RHODODENDRON Linnaeus.

R. malayanum Jack, Mal. Misc. 2 (7): 17. 1822; reimpr. Hook. Jour. Bot. 1:
369. 1834; III. 36; IV. 36; V. 254. Gunong Bunko or Sugarloaf Mountain
inland from Bencoolen, Sumatra. This is a well-understood species extending
from the Malay Peninsula and Sumatra to Borneo, Java, Ceram, and, including
R. apoanum Stein, the Philippines.

RHOPALA Schreber (1789) = Roupala Aublet (1775 %

R. attenuata Jack, Mal. Misc. 1 (5): 10. 1820; reimpr. Hook. Bot. Misc. 2: 65.
1830; III. 352; IV. 208; V. 227. Penang = Helicia attenuata (Jack) Blume.
Malay Peninsula, Sumatra, and possibly Java. This is correctly interpreted
by the British botanists concerned with the flora of the Malay Peninsula;
many collections available.

R. moluccana sensu Jack, Mal. Misc. 1 (5): 10. 1820; reimpr. Hook. Bot.
Misc. 2: 65. 1830; III. 352; IV. 208; V. 227, non R. Br. Penang (in a
garden) = Helicia petiolaris Benn. The Jack collection forms a part of
Wallich List no. 1041, Bennett’s binomial being strictly a new one for Jack’s
misinterpretation of R. Brown’s Moluccan species. Known only from the
Malay Peninsula.

R. ovata Jack, op. cit. 2 (7): 95. 1822; reimpr. Hook. Comp. Bot. Mag. 1: 259.
1836; Calc. Jour. Nat. Hist. 4: 353. 1843; IV. 209; V. 294. West coast of
Sumatra at Tapanuly = Helicia ovata (Jack) Benn. Known only from
Jack’s description. The large (25 by 15 cm.) entire sessile or subsessile
leaves should render identification a simple matter once material is available
from the west coast of Sumatra.

RONDELETIA Linnaeus.

R. corymbosa Jack, Mal. Misc. 1 (1): 4. 1820; reimpr. Hook. Bot. Misc. 1: 276.
1830; III. 16; IV. 16; V. 211. Penang = Greenea corymbosa (Jack) K.
Schum. (G. jackii Wight & Arn., 1834; Rondeletia spicata Wall. ex Roxb.
1824). Indo-China to the Malay Peninsula and Sumatra.

alae Roxburgh (1798) = Mallotus Loureiro (1790).
R. alba Roxb. Hort. Beng. 73. 1814, nom. nud.; Roxb. ex Jack, Mal. Misc. 1
(1): 26. 1820, descr.; reimpr. Hook. Bot. Misc. 1: 290. 1830; III. 227; IV.
131; V. 222; Roxb. Fl. Ind. ed. 2, 3: 829. 1832. Penang, Sumatra = Mallotus
albus (Roxb.) Muell.-Arg. (sed non M. albus sensu Pax & Hoffm., 1914)
(Mallotus macrostachys (Miq.) Muell.-Arg.). Pax and Hoffmann misin-
terpreted Roxburgh’s species in 1914, limiting it to India, Ceylon, and
Chittagong. They apparently overlooked the fact that Rofttlera alba Roxb.
was based wholly on Penang (Prince of Wales’ Island) specimens from a
plant cultivated at Calcutta. The Roxburgh description is a very inadequate
one; but that supplied by Jack in 1820 is excellent and validates the Roxburgh
binomial twelve years in advance of Roxburgh’s similar action; Jack’s
material was from Penang and Sumatra, and his Penang specimen was
apparently named at Calcutta. Mallotus macrostachys (Miq.) Muell.-Arg.,
type from Sumatra, is in all respects this misunderstood Mallotus albus

244 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxx
(Roxb.) Muell.-Arg.18 Malay Peninsula, Sumatra, Borneo, many collections.

SAGUS Gaertner (1788) = eae Rottboell (1783).

S. laevis Jack [App. Descr. Mal. Pl. 9. 1820]; reimpr. Jack ex Hook. Comp. Bot.
Mag. 1: 266. 1836; III. 13; IV. 13. Sumatra, Malacca (planted) =
as sagus Rottb, (1783). This smooth form is widely distributed in
Malaysia, extending from the Malay Peninsula and Sumatra to New Guinea,
usually planted. It is the source of sago, and man is probably responsible for
much of its present geographic distribution.

SALACIA Linnaeus; Jack, Mal. Misc. 2 (7): 92. 1822;. reimpr. Hook. Comp.
Bot. Mag. 1: 258. 1836; III. 101; IV. 197; V. 293. A general discussion, no
new names involved. There is a specimen named by Jack in the Rijksher-

arium, Leiden, under an unpublished binomial; the a is in the $
prinoides (Willd. : DC. group. See Johnia Samnairaua Jack abov

SAPINDUS Linnaeus.

S. rubiginosus Roxb. Pl. Coromand. 1: 44. pl. 62. 1795; Jack, Mal. Misc. 1 (1):
11. 1820; reimpr. Hook. Bot. Misc. 1: 280. 1830; III. 184; IV. 88; V. 214.
Penang = Erioglossum rubiginosum (Roxb.) Blume. India southward through
Malaysia to New Guinea and tropical Australia.

SONERILA Roxburgh.

S. erecta Jack, Mal. Misc. 1 (5): 7. 1820; reimpr. Hook. Bot. Misc. 2: 63. 1830;
III. 331; IV. 187; V. 225. Penang. Undoubtedly represented by Wallich 4092
and other collections from Penang. Malay Peninsula and reported from as
far north as eastern Bengal.

S. ae Jack, op. cit, 2 ie . 1822; reimpr. Hook. Jour. Bot. 1: 368.

34; III. > IV. 189; V. es (Sonerila pauciflora Blume, Cat. Gew.
ae 42. 1823). West coast of Sumatra at Tapanuly. A duplicate of the
original Jack collection is in the Delessert herbarium at Geneva. Represented
by various collections from Sumatra and Java.

S. moluccana Roxb. FI. Ind. 1: 182. 1820, ed. 2, 1: 178. 1832; Roxb. ex Jack,
Mal. Misc. 1 (2): 7. 1820; reimpr. Hook. Bot. Misc. 2: 64. 1830; III. 332;
IV. 188; V. 226. Penang. Of this Sonerila begoniaefolia Blume, Flora 14: 490.
1831, is a synonym, as is S. paradoxa Naud. (1851). Roxburgh’s description
is very short and unsatisfactory, but that of Jack is excellent in all respects.
The original author’s statement that the species was a native of the Moluccas
has confused the issue, for Roxburgh used the term “‘Moluccas”’ to cover other
parts of the Malay Archipelago. It is certain that his type actually came
from Penang; Jack’s specimen was undoubtedly named at Calcutta, or by
comparison with the description in his copy of Roxburgh’s manuscript. Some
have surmised that Roxburgh intended to write “malaccana” instead of
“moluccana,” but this is an unnecessary assumption. The latest to discuss
the case was Bakhuizen van den Brink f., Rec. Trav. Bot. Néerl. 40: 254. 1843,

% For Mallotus albus sensu Pax & Hoffm. Pflanzenreich 63 (IV. 147. VII): 168

Fl. Ind. ed. 2, 3: 826. 1832. This was Alston’s selection, Trimen, Handb. Fl. Ceyl.
6: 267. 1931, followed by Croizat, Jour. Arnold Arb, 21: 503. 1940. Its type was a
Silhet specimen, and Roxburgh’s description is an ample one

1952] MERRILL, WILLIAM JACK’S GENERA AND SPECIES 245

who preferred to retain Blume’s name of 1831; but it is apparent from his
references that he did not see Jack’s excellent description of 1820. Malay
Peninsula, Sumatra, Banka, Java. I accept Roxburgh’s earlier name with
confidence and reduce Blume’s later one as a synonym of it. The species does
not occur in the Moluccas.

SPHALANTHUS Jack, Mal. Misc. 2 (7): 55. 1822; reimpr. Hook. Comp. Bot.
Mag. 1: 155. a IIT. 339; IV. 195; V. 274 = Quisqualis Linnaeus.

S. confertus Jack, l.c.; reimpr. Il.cc. Sumatra, without definite locality =
Ouisguai ae (Jack) Exell, Jour. Bot. 69: 122, 1931 (Q. densiflora
Wall. List no. 4011. 1831, mom.; Miq. Fl. Ind. Bat. 1 (1): 611. 1955, deser.;
Sphalanibee diversifolius Jack ex ead Nomencl. ed. 2, 2: 621. 1841, nom.).
A specimen of the original Jack — is in the Delessert herbarium at
Geneva. Malay Peninsula, Sumat

SPHENODESME Jack, Mal. Misc. 1 (1): 19. 1820; reimpr. Hook. Bot. Misc. 1:
285. 1830; III. 43; IV. 43; V. 219.

S. pentandra Jack, l.c.; reimpr. ll.cc. Penang. The type of the genus, the species
extending from Assam to southern China, southward through Burma and
Indo-China to the Malay Peninsula and Borneo, Sphenodesme Jack was a
new name for Roscoea Roxb. nom. nud. (1814), non Smith (1804), and S.
pentandra Jack is supposedly the same as Roscoea pentandra Roxb., nom.
nud. (1814); but the first published descriptions are those of Jack.

STAGMARIA Jack [App. Descr. Mal. Pl. 12. 1820]; Jack ex G. Don, Gen. Syst.
2: 76. 1832; Hook. Comp. Bot. Mag. 1: 267. 1836; reimpr. III. 175; IV. 79 =
Gluta Linnaeus.

S. verniciflua Jack, l.c.; reimpr. ll.cc.; G. Don l.c. Sumatra at Natal on the west

ast = Gluta benghas Linn. Siam to the Malay Peninsula, Sumatra, Java,
and Borneo. Engler changed the spelling of the specific name to renghas, as
this is one of the Malay names of the species, and one may assume that
Linnaeus adopted the spelling benghas through some kind of an error.

STERCULIA Linnaeus.

S. angustifolia Roxb. Hort. Beng. 50. 1814, nom. nud.; Roxb. ex Jack, Mal. Misc.
1 (1): 21. 1820; reimpr. Hook. Bot. Misc. 1: 287. 1830; III. 223; IV. 127;
V. 220; Roxb. Fl. Ind. ed. 2, 3: 148. 1832. Penang = S. rubiginosa Vent.
Hort. Malm. 2: sub pl. 91, 1804 (S. jackiana Wall. List no. 1134. 1829, et in
Hook, Bot. Misc. 1: 287. 1830). Roxburgh’s nomen nudum of 1814 and his
very imperfect description of 1832 were based on specimens taken from a
tree cultivated in the Calcutta Garden labeled as having come from Nepal.
King in 1891 stated that Roxburgh’s excellent colored drawing at Calcutta is
unlike any Sterculia known from any part of the outer Himalayan region or
the plain at its base. He concluded that Roxburgh’s statement as to Nepal
was due to a mislabeled tree. He retained Sterculia angustifolia Roxb., so
well described by Jack, as a valid species allied to S. rubiginosa Vent., but
separated by certain style characters which I have not been able to check.
Ridley, however, Fl. Mal. Pen. 1: 371. fig. 27. 1922, reduced S. angustifolia
Roxb. ex Jack to S. rubiginosa Vent. In any case the Jack description validated
Roxburgh’s nomen nudum of 1814, and the species must be interpreted from
this and from Penang specimens. Burma to Indo-China through the Malay
Peninsula to Sumatra, Java, and Borneo.

246 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxuI

S. coccinea Jack, Mal. Misc. 1 (1): 20. 1820; reimpr. Hook. Bot. Misc. 1: 286.
1830; III. 222; IV. 126; V. 219, non Roxb. (1832). Penang (S. laevis Wall.
List no. 1138. 1829; Hook. Bot. Misc. 1: 287. 1830, in obs.). Wallich
definitely published Sterculia laevis in 1829 as a new name for S. coccinea,
as described by Jack, but not S. coccinea Roxb. Hort. Beng. 50, 1814, nom.
nud. (Fl. Ind. ed. 2, 3: 151. 1832, descr.). But Jack’s excellent description of
1820 antedates that of Roxburgh by twelve years. Adelbert in Backer
Beknopte Fl. Java IV. B (II) Sterc. 22, 1944, correctly eased Jack's
species. Burma and Siam, the Malay Peninsula, Sumatra, Borneo, and Java.

It is Sterculia coccinea Roxb. 1814, nom. nud., Fl. Ind. ed. 2, 3: 151.
1832, that needs to be renamed, and for this species, its type from India,
I propose the name Sterculia indica, nom. nov. I have not been able to locate
any other actually published name for this widely distributed species, which
extends from Sikkim, Bhotan, and Assam to Burma and Indo-China (but is
not as yet recorded from Siam). Sterculia bracteata Gagnep. of Indo-China
is apparently a closely allied species.

STROPHANTHUS De Candolle.

S. *plicatus Jack ex Burkill, Jour. Straits Br. Roy. As. Soc, 73: 218. 1916, nom.
nud. Sumatra at Bencoolen, and Penang = Scleranthera dubia (Sims) Pichon,
Not. Syst. 14: 90. 1951. Synonyms are Cameraria dubia Sims (1814),
Wrightia dubia Spreng. (1825), and Strophanthus jackianus Wall. List no.
1643. 1828, nom. nud., Wall. ex G. Don, Gen. Syst. 4: 85. 1838, descr. Some
modern authors place the species in Wrightia, others in Strophanthus. I
believe Pichon is correct in proposing the new generic name Scleranthera to
take this Jack species and an allied, perhaps even an identical one, of Siam
and Indo-China. The species is closer to Wrightia than to Strophanthus, but
I am convinced that it cannot properly be referred to either of these genera.
Malay Peninsula and Sumatra.

TABERNAEMONTANA Linnaeus.

T. macrocarpa Jack, Mal. Misc. 2 (7): 80. 1822; reimpr. Hook. Comp. Bot. Mag.
1: 253. 1836; III. 32; IV. 32; V. 286. Sumatra, inland from Bencoolen =
Ervatamia macrocarpa (Jack) comb. nov. (Pagiantha macrocarpa Markgr.
Notizbl. Bot. Gart. Berlin 12: 546. 1935). Additional synonyms are:
Orchipeda sumatrana Miq. Fl. Ind. Bat. Suppl. 553. 1863; Hallier f. Ann.
Jard. Bot. Buitenz. 13: 285. 1896, Bull. Herb. Boiss. 6: 615. 1898; Neuburgia
*sumatrana Boerl. Handl. Fl. Nederl. Ind. 2: 392. 1899; Tabernaemontana
monocarpa Steud. Nom. ed. 2, 2: 658. 1841, sphalm; Tabernaemontana
plumeriaefolia Elm. Leafl. Philip. Bot. 1: 333. 1908; Tabernaemontana
plumeriaefolia Merr. Enum. Philip. Fl. Pl. 3: 326. 1923. Sumatra, Borneo,
Philippines, and apparently in the Malay Peninsula and Java. I have seen
Miquel’s type of Orchipeda sumatrana, a Teysmann specimen from Sumatra,
and also Achmad 110 from Simaloer Island near Sumatra, while Krukoff 4190
a sterile specimen, may belong here; also from Borneo Hallier 1363, Wood
927, 1802, Clemens 9809, 21210, 26195, Ramos 1304, Haviland & Hose
3501, and Neth. Ind. For, Serv. 16316; and from the Philippines Elmer 7502,
7754, and Bur. Sci. 33826 Ramos & Edano

I strongly suspect that 7. sphaerocarpa ‘Blume, Bijdr. 1028. 1826, de-
scribed in detail by Koorders . Valeton, Meded. Lands Plant. 11: 103. 1894
(Bijdr. Boomsoort. Java 1: 103), and as well illustrated by them, :.
Baumart. Java 4: pl. 623, 624. 1916 = Pagiantha sphaerocarpa Markgr.

1952] MERRILL, WILLIAM JACK’S GENERA AND SPECIES 247

Notizbl. Bot. Gart. Berlin 12: 546. 1935 = Ervatamia sphaerocarpa Burkill,
Kew Bull. 1935: 317. 1935, may well belong here, and Tabernaemontana
megacarpa Merr. Philip. Jour. Sci. 4: Bot. 318. 1909 = Pagiantha megacarpa
Markgr. |.c., of which no specimens are actually available to me at this time,
may also represent a form of Jack’s species. In view of the discordant
elements placed in Pagiantha, particularly as to the shape, size, and general
characters of the fruits (compare those of T. pandacagui Poir. with those of
T. macrocarpa Jack and T. sphaerocarpa Blume, all supposed to belong in
Pagiantha), I see nothing to be gained by recognizing a distinct genus here.
At the Rijksherbarium, Leiden, where I made numerous notes on various
species in this difficult group, Tabernaemontana plumeriaefolia Elm. had
been referred to Rejoua, where I believe it does not belong, while Tabernae-
montana sumatrana Merr. Contr. Arnold Arb. 8: 139. pl. 13. 1934 had been
placed with one of Miquel’s species. It is not the same as Jack’s species but
is an Ervatamia. Pseudixora sumatrana Miq. Fl. Ind. Bot. 2: 209. 1857 =
Randia sumatrana Miq. Ann. Mus. Bot. Lugd.-Bat. 4: 235. 1869, should be
compared. Note the slender elongated corolla tubes, which are very different
from those of T. macrocarpa Jack.

Under suspicion are various collections from the Malay Peninsula and Java,
distributed as representing Tabernaemontana sphaerocarpa Blume. Two
Johore collections, neither with good flowers, Corner 20696 and Ngadiman
34741, are suspiciously similar to Sumatra specimens which I refer to Jack’s
species. On the whole, the few flowers I have seen of Java amine seem to
be more slender than are those of Jack’s species as I here interpre

Jack described the fruits as being subglobose and as large as ee.
The fruit characters of the various species above reduced conform, as is also
the case with the other characters in Jack’s distinctly good description. I do

and prominently nerved leaves, and fairly large flowers with relatively thick
corolla tubes. Here is a case where it seems to be apparent that an early-
named and well-described species has been rather consistently ignored, largely,
it is suspected, because there is no extant type of Jack’s species. Here it is
not at all difficult, now that modern collections from Sumatra are available,
to match Jack’s original description with some of these collections.

TACCA Linnaeus.

T. cristata lees: Mal. Misc. 1 ae 23. 1820; ets Hook. Bot. Misc. 2: 73.
1830; III. 9; IV. 9; V. 234. Singapore and Penang (T. rafflesiana Jack ex
Wall. List no. 5172. 1832, nom. nud.). A hate well-known species.
Malay Peninsula, Sumatra, Borneo.

TERNSTROEMIA Mutis ex Linnaeus.

T. acuminata Jack, Mal. Misc. 2 (7): 26. 1822; reimpr. Hook. Jour. Bot. 1: 375.
1834; III. 204; IV. 108; V. 259. West coast of Sumatra at Tapanuly =
Saurauia sp. aff. S. tristyla DC. Known only from Jack’s description. Jack
merely accepted Roxburgh’s misinterpretation of the genus.

T. cuspidata Jack, op. cit. 2 (7): 28. 1822; reimpr. Hook. Jour. Bot. 1: 377.
1834; III, 206; IV. 110; V. 260. Sumatra at Salumah, on the west coast =
ern sp. Like T. aaa Jack but with 5-celled ovaries and 5 styles.
Known only from Jack’s description.

T. pentapetala Jack, Mal. Misc. 1 (5): 40. 1820; reimpr. Hook. Bot. Misc. 2:

248 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxur

84. 1830; III. 204; IV. 108; V. 242. Penang = Saurauia sp. Clearly in the
group with S. tristyla DC. Cleyera pentapetala Spreng. Syst. 2: 596. 1825 is
a synonym. Known only from Jack’s Saget ena

T. rubiginosa Jack, Mal. Misc. 1 (5): 38. 1820; reimpr. Hook. op. cit. 83:
III. 203; IV. 107; V. 241. Sumatra = a, rublginosa (Jack) comb. nov.
(Cleyera rubiginosa Spreng. Syst. 2: 596. 1825; Saurauia jackiana Korth.
Verh. Nat. Gesch. Bot. 127. 1842, quoad syn. Jack), All Sumatra specimens
that I have seen named as representing Saurauia jackiana Korth. (Korthals!,
Beccari 669, Lorzing 5689, Biinnemeyer 3463), have pedicels up to 1.5 cm.
long, somewhat appressed scaly below, but not setose, si the sepals are
broadly ovate, coriaceous, glabrous, rugose when d to 1 cm. long.
They do not represent the species that Jack cee mn the Korthals
sheet is an unpublished binomial indicative of his original intention of
describing his specimen as a new species. Fortunately there is an extant type
of Jack’s species in the Rijksherbarium, Leiden. It is a rather poor specimen,
but the pedicels are up to 3 cm. long and rather densely setose, as are the
sepals. This important specimen was filed under S. cauliflora DC., where
it does not belong. The leaves are about 15 X 8 cm. with about 14 pairs of
nerves. I have not succeeded in matching the Jack type with any other
collection.

T. serrata Jack, Mal. Misc. 2 (7): 27. 1822; reimpr. Hook. Jour. Bot. 1: 376.
1834; III. 205; IV. 109; V. 259. Pulo Nias, off the west coast of Sumatra =
Saurauia media Korth. Verh. Nat. Gesch. 125. 1842 (S. camptodonta Miq.
and SS. inflexidens Miq., 1862). A species in the group with S. tristyla DC.,
Jack’s type which I have seen, being preserved in the Rijksherbarium, Leiden.
The synonymy as above indicated is based on unpublished herbarium notes
of C. B. Clarke, who actually examined Jack’s and Miquel’s types. Jack’s
specific name is invalidated in Saurauia by S. serrata DC. (1822). Known
only from Sumatra.

TETRACERA Linnaeus.

T. arborescens Jack, Mal. Misc. 1 (5): 45. 1820; reimpr. Hook. Bot. Misc. 2:
87. 1830; III. 218; IV. 122; V. 244. West coast of Sumatra, Tapanuly Bay.
A Jack specimen is preserved in the Rijksherbarium, Leiden. Tetracera lucida
Wall. List 6631. 1832, nom., validated by Ridley, Fl. Mal. Pen. 1: 5. 1922,
is a synonym, fide Dr. Hoogland (7. euryandra sensu Hook. f. Fl. Brit. Ind. 1:
32. 1872, non Vahl). Malay Peninsula, Sumatra. There are doubtless botanical

because Jack erred in describing this as a tree; it was corrected by him in
one of his letters to Nathaniel Wallich; see Jour. Straits Br. Roy. As. Soc, 73:
229. 1916

TETRANTHERA Jacq. (1797) = Litsea Lam. (1791), nom. conserv

T. cordata Jack, Mal. Misc. 2 (7): 34. 1822; reimpr. Hook. Comp. Bot. Mag. 1:
148. 1835; III. 356; IV. 212; V. 263. West coast of Sumatra = Litsea cordata
(Jack) Hook. f. (L. cordifolia Griff.). Malay Peninsula, Sumatra, Borneo.

UROPHYLLUM Wallich in Roxb. Fl. Ind. 2: 184. 1824; reimpr. Calc. Jour. Nat.
Hist. 4: 17. 1844; IV. 17. This genus was first named Patisna Jack, but never
published; see Burkill, Jour. Straits Br. Roy. As. Soc. 73: 196, footnotes 175,
218, 238, 1916. Jack also proposed the alternate name Wallichia for it, but
Wallich, to whom Jack had extended certain discretionary powers, substituted

1952] MERRILL, WILLIAM JACK’S GENERA AND SPECIES 249

his own name, Urophyllum, apparently being intrigued by Jack’s reference to
“those acuminate gentry the Patisnae.’”’ I therefore do not accept Griffith’s
correction of Urophyllum Wallich to Urophyllum Jack. Wallich did credit
the copy to Jack, as also for the two species, but the published names are
Wallich’s. Burkill’s suggestion that the name Wallichia Reinwardt (1823),
non Roxb. (1819), may have been due to Jack’s having mentioned the name
to Reinwardt when he visited Buitenzorg in 1821, is probably correct.
Blume, in publishing it, apparently thought that Reinwardt originated the

name.

U. glabrum Wall. op. cit. 186; reimpr. Calc. Jour. Nat. Hist. 4: 18. 1843;
IV. 18. Penang = Urophyllum arboreum (Reinw. ex Blume) Korth. This was
Wallich’s own binomial, as he originated the generic name. He copied Jack’s
specific description. The species is common and widely distributed in the
Malay Peninsula and the Sunda Islands; but Wallichia arborea Reinw. (1823)
= Urophyllum arboreum (Reinw. ) Korth. was published one year earlier than
Jack’s species. As I understand the species, some of the synonyms are
Wallichia arborea Reinw., 1823; Axanthes arborea Blume, 1826; Schwenk feldia
glabra Spreng., 1827; Urophyllum repandulum Migq., 1857, and Urophyllum
hexandrum O. Kuntze, 1891. Malay Peninsula, Sumatra, Java, Borneo, very
many collections. The Philippine form referred here is the allied
memecyloides (Presl) Rolfe.

U. villosum Jack ex Wall. in Roxb. op. cit. 185; reimpr. lIl.cc. Penang. Two
exact synonyms are Schwenkfeldia malaccensis Spreng. (1827) and S. villosa
D. Dietr. (1839); others are Axanthes tomentosa Blume ex DC., 1830, and
Urophyllum tomentosum Miq., type of both from Penang. I know this species
only from Penang, Singapore, and various parts of the Malay Peninsula.

UVARIA Linnaeus.

U. hirsuta Jack, Mal. Misc. 1 (5): 46. 1820; reimpr. Hook. Bot. Mise. 2: 87.
1830; III. 220; IV. 125; V. 245. Penang. One of the rather strongly
characterized species of the genus. Malay Peninsula, Sumatra, Java; the
Burma record perhaps doubtful as the species has not been recorded from
Siam. The “Molucca” Uvaria pilosa Roxb. Fl. Ind. ed. 2, 2: 665. 1832 is a
synonym of Jack’s species. Its type was undoubtedly from Penang or Sumatra.
The strongly marked species does not occur in the Moluccas.

VACCINIUM Linnaeus.

V. sumatranum Jack, Mal. Misc. 2 (7): 18. 1822; reimpr. Hook. Jour. Bot. 1:
370. 1834; III. 35; IV. 35; V. 255. Gunong Bunko (Sugarloaf Mountain),
northeast of Bencoolen, Sumatra. Apparently not as yet associated with any
other described species, but clearly in the group with V. ellipticum (Bl.) Miq.
and V. laurifolium (Bl.) Miq. of Java, and probably the same as one of these.

VERATRUM Linnaeus.

V. malayanum Jack, Mal. Misc. 1 (2): 25. 1820; reimpr. Hook. Bot. Misc. 2:
74, 1830; III. 9; IV. 9; V. 234. Penang = Hanguana malayana (Jack) Merr.
Philip. Jour. Sci. 10: Bot. 3. 1915. Synonyms are Hanguana kassintu Blume
(1827), Veratronia malayana Miq. (1859), and Susum malayanum Planch.
(1892). Malay Peninsula, Sumatra, Java, Borneo, and Palawan, Mindoro and
Mindanao in the Philippines. Hanguana Blume dates from 1827, Susum Blume
from 1830, and Veratronia Miq. from 1859

250 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxx

VITEX Linnaeus.

V. arborea Roxb. Hort. Beng. 46. 1814, nom.; Roxb. ex Jack, Mal. Misc. 1
(1): 18. 1820; reimpr. Hook. Bot. Misc. 1: 285. 1830; III. 40; IV. 40; V. 218;
Roxb. Fl. Ind. ed. 2, 3: 73. 1832. Sumatra = V. pubescens Vahl (1794). India
to Indo-China southward through the Malay Peninsula to Sumatra, Borneo,
Java, the Philippines, Celebes, and Timor. Some botanists have followed
Hallier f. in accepting the older V. latifolia Lam. as the proper name for
this species, but Lamarck’s binomial of 1788 is invalidated by the earlier
V. latifolia Mill. (1768).

VITIS Linnaeus.

V. racemifera Jack, Mal. Misc. 2 (7): 94. 1822; reimpr. Hook. Comp. Bot. Mag.
1: 258. 1836; III. 194; IV. 98; V. 294. Sumatra, no definite locality, but
surely from the west coast = Ampelocissus racemifera (Jack) Planch. in DC.
Monog. Phan. 5: 410. 1883. I feel certain that Rahmat si Toroes (Boeea) 107,
1657, 2180, 3875, 7158, 9299, 9548, and Yates 706, all from Sumatra,
represent Jack’s species. Planchon, while recognizing the species and correctly
transferring it to Ampelocissus, based his description entirely on Jack’s
excellent one. At the same time he described A. korthalsiti Planch., type from
Sumatra, as new, but expressed the opinion that it might not be distinct from
Jack’s species. Also to be compared here are the Sumatra collections referred
to A. thyrsiflora (Blume) Planch. and Vitis polystachya Wall.

WALLICHIA Reinwardt ex Blume = Urophyllum Wallich.
W. *glabra Jack ex Burkill, Jour. Straits Br. Roy. As. Soc. 73: 255. 1916, nom.
nud. = Urophyllum glabrum Wall.; see p. 249.

WORMIA Rottboell.

W. excelsa Jack, Mal. Misc. 2 (7): 69. 1822; reimpr. Hook. Comp. Bot. Mag. 1:
221. 1836; III. 219; IV. 123; V. 281. West coast of Sumatra at Bencoolen =
Dillenia excelsa (Jack) Gilg in Engl. & Prantl, Nat. Pflanzenfam. 3 (6):
123. 1893. Hoogland, in his monograph of Dillenia L., Blumea 7: 68. 1952,
lists about 20 synonyms, among them Capellia multiflora Blume (1825),
Wormia oblonga Wall. (1855), W. grandifolia Mig. (1863), and others.
Malay Peninsula, Banka, Sumatra, Java, Borneo, and Balabac in the
Philippines.

W. pulchella Jack, op. cit. 70; reimpr. ll.cc. West coast of Sumatra at Natal =
Dillenia pulchella (Jack) Gilg i in Engl. : ite Le. a Peninsula, Riouw
and Lingga Archipelagos, Banka, Sumatra, and Borneo. Synonyms after the
Hoogland treatment are Dillenia eee Martelli A D. parvifolia Martelli.

ZINGIBER Adanson.

Z. gracile Jack, Mal. Misc. 1 (1): 1. 1820; reimpr. Hook. Bot. Misc. 1: 273.
1830; III. 3; IV. 3; V. 209. Penang. The latest critical consideration of the
species is that of Holttum, Guard. Bull. Singapore 13: 63. 1950. Widely dis-
tributed in the Malay Peninsula.

ARNOLD ARBORETUM,
HARVARD UNIVERSITY.

PLATE I

Jour. ARNOLD Ars. VoL. XXXIII

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PLATE 1. TYPE COLLECTION OF DIDYMOCARPUS CRINITA JACK, THE LABEL IN
WILLIAM JACK’s HANDWRITING (HERB. EDINBURGH)

ya JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxx1r

THE COMPARATIVE MORPHOLOGY OF THE ICACINACEAE, VI.
THE POLLEN

A. ORVILLE DAHL

With nine plates

THE ICACINACEAE Miers is a dicotyledonous family made up of approxi-
mately 60 genera embracing, according to Sleumer (30), about 300 species
largely confined to the tropics of both the old and new worlds. Most of
the species are woody. The affinities of this family, as well as the specific
and generic concepts within the group, have been variously interpreted.
Engler (7) placed the family in the suborder /cacinineae following the
suborder Celastrineae in the order Sapindales while Wettstein (31) and
Hutchinson (24) included the /cacinaceae in the order Celastrales. Hal-
lier (13) interpreted the family as having affinities with the Olacaceae and
included it in the order Santalales. Recently, Howard (16-23) has dis-
cussed the systematic position of the family.

The Jcacinaceae have been the subject of intensive anatomical, morpho-
logical, and taxonomic investigations initiated by I. W. Bailey and R. A.
Howard in 1941 (1). These studies have provided a critical basis for the
interpretation of old as well as recent researches in the family. In the lat-
ter category is the present study of pollen grain morphology of the
Icacinaceae. Studies of comparative pollen morphology have been men-
tioned, not infrequently, as being capable of yielding a distinctive set of

systematics and phylogeny. However, it seems regrettable that detailed,
comparative pollen studies of genera or families are rare in either the con-
temporary or older literature. Citations to extant studies will be found
in Wodehouse (32) and Erdtman (8, 10). When considered along with
data derived from other techniques, i.e. data concerning other plant parts,
the information concerning pollen grain form is a definite aid in arriving
at a more natural arrangement of species. This study of pollen in the
Icacinaceae was pursued from this point of view.

MATERIALS AND METHODS

All observations are based upon pollen grains mounted from herbarium
specimens. In general, two microscopical mounts were made from each
specimen. One mount was prepared in lactic acid (ca. 85%) to present
pollen in the expanded condition and the other in oil to illustrate the
grains in the contracted or air-dry state. The hard, brittle anthers were
softened in 10% alkali (NaOH or KOH) and then transferred to a drop
of lactic acid on a slide. A number of duplicate specimens were prepared
by quickly softening the material in hot 45% acetic acid. After dissecting

1952] DAHL, MORPHOLOGY OF THE ICACINACEAE 253

out the pollen grains excess acetic acid and debris are removed followed
by the addition of lactic acid and cover-glass. With this procedure, the
degree of expansion of the pollen grains is occasionally less than with the
above technique. The preparations were sealed either with paraffin-bees-
wax or ringing cement (Turtox) and stored in a horizontal position. All
observations were made with an apochromatic, oil immersion objective
(90x, N.A. 1.25) and a compensating ocular (12x). In general, data on
size are based upon the measurement, by both camera lucida and ocular
micrometer, of a minimum of 25 suitably oriented grains for each species.
Urandra scorpioides and U. Brassii possess the smallest pollen grains ob-
served in the family — respectively, ca. 7:6 (L) & 14p (D) and ca. 9.4
(L) & 12.3n (D) while Nothapodytes pittosporoides and Demostachys
Vogelii produce the largest grains — respectively, ca. 102 (L) & 89.8p
(D) and ca. 92 (L) & 99.2» (D).

An obvious source of error in this type of investigation is the inade-
quacy of material stemming from immaturity of the specimen and over-
or under-expansion of the grains during preparation. Wherever possible
such errors were reduced by replication of preparations and addition of
specimens. However, in the case of some species of which only limited
material was available, reference to encountered inadequacies will be found
in the text.

All of the drawings are intended to be faithful records of all visible de-
tails. All evident structures were carefully mapped out with the aid of the
camera lucida. Many of the drawings are semi-diagrammatic in the sense
‘that only part of the surface, etc. is completed in essentially photographic
form. The completed portion of the figure of any one grain is intended
to emphasize the essential diagnostic features. Since several focal planes
are presented in each drawing, it is essential to realize that features such
as germinal apertures present in the lower hemisphere (lower side of the
grain) are depicted by broken lines. Those features on the surface (in the
upper hemisphere) are indicated by continuous or stippled lines. Fre-
quently, the relative thickness of the exine (outer layer of pollen grain
wall) as seen in optical section is represented in at least a portion of the
drawing. Where the exine is foveate (‘‘pitted’’),* this is often indicated
in both the surface and optical section views. The foveae (‘‘pits”’) or
lacunae of a foveate (‘“‘pitted’’) or reticulate exine are, in general, indicated
by clear, unstippled areas of a form and size appropriate to the species.
This rule has been abandoned in noted cases where the foveolae (very
small “pits”) are so small that it was necessary to depict the “pits”

final preparation of the great majority of these drawings, we are indebted
to Mrs. John E. Rogerson of Milton, Massachusetts. Pollen material of
some 130 species in 50 genera have been examined in this study.

* Since the terms: — pit, pitting, pitted have special significance in discussions of
somatic cell wall structure (29) it would seem desirable to avoid application of these
terms in a different sense to the pollen grain wall.

254 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxmI

ACKNOWLEDGMENTS

I am deeply grateful to Professor I. W. Bailey who generously provided
the research materials and facilities utilized in this study. Information
concerning the taxonomic status of a number of specimens has been kindly
provided by Dr. R. A. Howard. For research support during the final
phases of this work, I am indebted to the Graduate School of the Univer-
sity of Minnesota.

OBSERVATIONS

From the standpoint of pollen grain morphology, the family /cacinaceae
has proven to be one of great interest because of the marked variation in
form of pollen within the group. Within the family there exists, on the
basis of pollen morphology alone, differentiation between species, genera,
and certain larger categories. As is frequently the case, this differentiation
does not find uniform expression throughout all areas of the family. For
example, a number of genera possess a basically similar pollen morphology
which of course would favor their being grouped together on the basis of
this character. Depending upon the plexus within the family that one is
concerned with, such grouping may culminate in clusters of related species,
e.g. a genus, or in clusters of allied genera. The following is a summary
of the notable pollen grain forms extant in the family together with a
grouping of genera based upon the differentiation finding expression in
pollen grain structure. How this organization of species compares with
that established on other grounds will receive attention in the Discussion.

Data discussed in the present communication pertain largely to the
surface characters of essentially intact pollen grains in the sense that the
preparational technique has preserved both exine and intine. In many
instances, critical interpretation of the minutiae of pollen grain wall struc-
ture must await the analysis of thinly-sectional material (cf. 5).

In general, the pollen grain descriptions have been reduced to their
simplest terms. Data were accumulated under four major headings: (A)
shape of pollen grain; (B) character of germinal apertures; (C) charac-
ter of pollen grain surface (exine); and (D) size of pollen grain.

In recording the size of the pollen grains in this paper the measurements
are approximate and recorded as illustrated by the following: 50 X 35 uh.
The first number always signifies the length of the axis terminated by the
poles and the second number represents the diameter, the axis transverse
with respect to the length. Thus the length is greater than the diameter
in the case of ellipsoidal grains, equal to the diameter in spheroidal types,
and less than the diameter in species having oblate pollen grains. Some-
what more elaborate schemes have been devised [cf. Erdtman (8, 9),
Wodehouse (32), and Faegri and Iversen (12, 25) |] but the above cover
the characters of diagnostic value. In general, a minimum of highly
specialized terms have been utilized. These are briefly defined where they
first appear.

In citing the specimens studied, collector and number (where given)

1952] DAHL, MORPHOLOGY OF THE ICACINACEAE 255

and herbarium (A = Arnold Arboretum, B = Berlin, Museum of Botany,
= University of California, F = Chicago Museum of Natural History

[Field Museum], G = Gray Herbarium, NY = New York Botanical
Gardens, US = United States National Herbarium) are given after the
species name. Grateful acknowledgment is made to the curators of these
herbaria for the contribution of research material.

In viewing the group broadly, one senses two extremes. The first of
these is illustrated by species having essentially smooth (psilate) pollen
grains while the other is seen in species having variously echinate (spinose)
grains. The transitional area between these two extremes is largely popu-
lated by a variety of “pitted” (foveate), “pitted’-reticulate (foveate-
reticulate), reticulate, and papillate types. In setting up a classification
on the basis of pollen characters, the genera can be resolved into some 15
relatively small groups. This classification which represents, essentially,
a table of contents for the family, may be presented as follows:

TYPES OF POLLEN OF THE ICACINACEAE
A. Exine psilate (smooth) to foveate (“pitted”) to reticulate to lophate to
areolate; colpate or, more commonly, colporate (having germinal furrows
with pores).

1. Psilate; tricolporate (having three furrows with germ pores).
1. Emmotum 2. Ottoschulzia 3. Poraqueiba
2. Foveolate (finely ‘‘pitted’”) to foveate to foveate-reticulate; tricolporate.
4. Cassinopsis 9. Citronella 14. Lavigeria
5. Pittosporopsis 10. Dendrobangia 15. Leretia
6. Calatole 11. Platea 16. Pleurisanthes
7. Pennantia 12. Humirianthera 17. Apodytes
8. Oecopetalum 13. Icacina
3. Foveate-reticulate; tricolpate (triaperturate).
17. Apodytes 18. Anisomallon
4. Foveolate with alignment of foveolae (‘‘pits”) in direction of finger-
printing or rugulation; bald “polar” zones, non-committally, triaper-
ate.
19. Leptaulus
5. Reticulate; tricolpate (triaperturate).
20. Rhaphiostylis
6. Lophate; tricolpate.
21. Gonocaryum
7. Foveate-granular, in direction of an areolate exine; tricolpate.

22. Merrilliodendron
8. Areolate; tetracolporate (four furrows with pores, stephanocolporate).

23. Alsodeiopsis

B. Exine psilate to foveate-papillate to papillate to papillate-echinate; colpate
or, more commonly, porate; if psilate not colpate.

256 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxmI
1. Psilate; triporate.
24. Irvingbaileya 25. Gastrolepis

2. Foveolate-papillate to minutely papillate-echinate; triporate.

26. Codiocarpus 28. Medusanthera 30. Cantleya

27. Stemonurus 29. Urandra 31. Lasianthera
3. Foveolate-papillate; stephanoporate or periporate (respectively, 4 or

m es in either equatorial or scattered arrangement); rarely

triporate.

31. Lasianthera 32. Natsiatum

4. Granular-papillate-echinate; tricolpate.
33. Hosiea
C. Exine echinate; commonly with porate germinal apertures usually having
annuli (differentiated pore margins), rarely stephanocolpate.

1. Stephanocolpate; echinate-foveolate.

34, Mappia 35, Desmostachys 36. Nothapodytes
2. Triporate with annuli; echinate.

37. Discophora 40. Polyporandra 43. Polycephalium

38. Mappianthus 41. Miquelia 44. Chlamydocarya

39. TIodes 42. Phytocrene

3. Apparently triporate (or periporate) without annuli; echinate.
45, Rhyticaryum 46. Sarcostigma 47. Pyrenacantha

The various genera will be briefly described in the same sequence seen
in the preceding classification. Where no remarks are appended to the
individual species listing, it is to be inferred that the generic description
covers the diagnostic characteristics of that species. It is perhaps obvious
that the generic descriptions are necessarily based upon the species avail-
able for analysis.

1. EMMOTUM Desv. ex Hamilton

Grains ellipsoidal; tricolporate, the furrows (colpi) elongate, rather
deeply impressed, extending as seen in equatorial view to almost the full
length of the grain; germ-pores well defined, of circular to rectangular out-
line; exine smooth (psilate) or essentially so.

Emmotum nitens (Benth.) Miers. — Fics. 5, 5A.— 20.3 & 16.8 p.
— Burchell 8233 (G).

Emmotum argenteum Gleason. — Fics. 3, 3A.— 27.7 & 25.4 p.—
Tate 564, type (NY).

Conspicuous rectangular germ-pore; the structure of the pollen grain
suggests a close alliance with E. glabrum, a relationship suggested by
Howard (18) on the basis of both species possessing a two-celled ovary.

Emmotum nudum Howard. — Fics. 2, 2A.—26 X 19 p.— Spruce
3541 (G).

The germ-pore is circular to rectangular, with the polar margin slightly

1952] DAHL, MORPHOLOGY OF THE ICACINACEAE 257

thickened. The exine is smooth except for delicate foveation (“‘pitting”’)
faintly evident at the poles.

Emmotum glabrum Benth. ex Miers. — Fics. 4, 4A. — 23.2 & 20.8
p. — Klug 3990 (G).

Well-defined circular germ-pore often showing a slight collar or rim
with a somewhat ragged margin. Except for the shape of the germ-pore,
this species is closely similar to E. argenteum,

Emmotum orbiculatum (Benth.) Miers.—22 18 p.— Ducke
919 (G).
Emmotum affine Miers. — 21.6 & 17.4 ».— Pickel 3125 (G).

Emmotum fagifolium Desv. ex Hamilton.— Fics. 1, 1A.— 26.4
x 20.8 p.— La Cruz 3375 (G); Pinkus 193 (G).

In some grains there is a suggestion of two faint ridges lying parallel to
the equator on either side of the germ-pore. The available material was
only fair. In the specimen of Pinkus, pollen grain shape was commonly
spheroidal with a diameter of ca. 22 p.

2. OTTOSCHULZIA Urban

Grains ellipsoidal; tricolporate; germ-pore rectangular to circular, some-
what irregular in outline, delicate ridges oriented equatorially on either
side of the pore visible in some grains; exine smoot

Ottoschulzia cubensis (Wright) Urban. — Fics. 6, 6A.—31.2
28.4 np. — Wright 2639, IsoTYPE (G).

3. PORAQUEIBA Aublet

Grains ellipsoidal ; tricolporate; furrows usually with ragged and some-
what granular margins; conspicuous germ-pore of rectangular outline; the
pore so elongate equatorially beyond the typical furrow region imparting a
“‘double-belted” appearance (costae pori transversales) to the grain when
equatorially oriented; exine smooth.

Poraqueiba sericea Tulasne.— Fics. 7, 7A.—57.8 & 41 p.—
Ducke 25 (A); Spruce 1748 (G).

It is evident, in expanded grains, that the poral region in the furrow
floor is capable of marked expansion in the form of a circular area sur-
rounded by a ragged collar or rim. The rim is somewhat reminiscent of
that seen in Emmotum glabrum. The pollen of Spruce 1748 was some-
what immature with the grains averaging ca. 41 » in length.

4. CASSINOPSIS Sonder
Grains ellipsoidal to markedly oblate; characteristically tricolporate;
furrows elongate, narrow; germ-pores circular to lenticular; exine smooth
to foveolate (very finely “pitted” a;
Cassinopsis ilicifolia (Hochst.) O. Ktze.— Fics. 10, 10A, 10B. —
20.6 X 19.6 w.— Burchell 5817 (G).

258 JOURNAL OF THE ARNOLD ARBORETUM [voL. xxxim

While most of the material (labelled ‘‘C. capensis Burchell”) was some-
what immature, it is evident that the typically ellipsoidal grains are less
sharply triangular in polar view than those of C. tinifolia. In the majority
of the observed grains, the furrows were constricted at the equator, thus
obscuring the germ pore. The pore is rarely visible as a lens-shaped open-
ing. (Fig. 10A).

mapper tinifolia Harv. — Fics. 11, 11A.—16.5 & 20.2 p.—
Wood 9428 (A).

In polar view, the oblately flattened grains are triangular as in Anisomal-
lon utatlolan, The grains are diamond-shaped in exact equatorial
view. The deeply embossed furrows broaden out markedly at the equator
to form a rectangular area surrounding the circular germ-pore. In ex-
panded grains the pore is bounded by a rim simWar to that found in
Dendrobangia boliviana and Citronella. The exine is foveolate rather
than smooth as in C. ilicifolia.

5. PITTOSPOROPSIS Craib

Grains ellipsoidal; tricolporate; furrows not expanding in the available
material; germ-pore circular; exine in polar regions showing foveolae which
in other regions are so diminished in size that the effect of a finely stippled
surface is obtained.

Pittosporopsis Kerrii Craib.— Fics. 8, 8A.—31.2 & 28.5 p—
Henry 11778 (A)

6. CALATOLA Standley

Grains oblately spheroidal; tricolporate, (one four-furrowed grain
noted); furrows of relatively limited longitudinal extent and, in the ma-
terial studied, do not open widely; the circular germ-pore is rendered dis-
tinctive by its irregularly toothed margin; exine foveolate as in Cassinopsis
tinifolia.

Calatola laevigata Standley.— Fics. 14, 14A.— 24.4 & 27 ».—
Schipp 8-708 (F).

7. PENNANTIA Forst.

Grains ellipsoidal to oblately spheroidal; tricolporate ; furrows elongate,
extending almost to the poles, furrow-margin somewhat irregular; germ-
pore circular to somewhat rectangular with ragged, irregular margin;
frequently fragments of foveolate (‘‘pitted”) exine occur on the poral
membrane; exine nerd foveolate (very finely ‘“‘pitted”), though the
general effect is smoot

Pennantia Ne Forst. — Fics. 12, 12A.— 25.7 & 25.3 ».—
T. Kirk (G); Raoul (as P. Endlicheri) (G).

Pennantia Cunninghamii Miers. — Fics. 13, 13A.— 22.5 « 20.9
p. — Moore (G).

The exine is less foveolate in this species than in the preceding one.

1952 | DAHL, MORPHOLOGY OF THE ICACINACEAE 259

8. OECOPETALUM Greenman & Thompson

Grains ellipsoidal; tricolporate; furrows prominent at equator, gradually
disappearing poleward; germ-pore large and conspicuous, of elliptical out-
line; exine foveolate, the largest of the “pits” occurring at the poles.

Oecopetalum mexicanum Greenm. & Thomps.— Fics. 9, 9A.—
36.3 & 31.8 p.— Purpus 6159, ISOTYPE (G).

In its combination of pollen grain characters (furrow folds, foveola-
tion, conspicuous germ-pore, and size), this species is distinctive within
the group.

9. CITRONELLA D. Don

Grains ellipsoidal to oblate; tricolporate; commonly with germ-pore of
circular outline; exine commonly finely and uniformly foveate (‘‘pitted”)
but, in some species, varying to foveate-reticulate particularly in polar
regions.

Pollen of the investigated species of the Old World is not differentiated
from that of the American species (cf. Figs. 15, 16). The relatively uni-
form pollen morphology throughout the genus supports Howard’s (20)
recent treatment of the complex.

Citronella costaricensis (Donn. Sm.) Howard.— Fics. 17, 17A.
— 26.4 & 31.2 p.— Tonduz 11664 (G); Austin Smith P2394 (A).

The shape of the pollen grain as well as the germ-pores and exine fovea-
tion are essentially as in C. Gongonha.

Citronella mucronata (Ruiz & Pavon) D. Don. — Fic. 18.— 36.7
xX 34.2 p.— Munos & Coronel 1394 (G).

Citronella Gongonha (Mart.) Howard.— Fics. 16, 16A.— 26.4
< 29.8 p.— Hoehne 385 (A).

The oblately spheroidal grains when expanded show large, gaping germ-
pores bounded by low, collar-like rims.

Citronella incarum (Macb.) Howard.— Fics. 21, 21A.—23.4 x
24 ».— Weberbauer 7056 (F).

This species is distinctive in its comparatively small size. The germ-
pore is bounded by a collar-like rim as in C. Gongonha.

Citronella ilicifolia (Sleumer) Howard. — Fics. 19, 19A.— 25.5
« 24.9 ».— Weberbauer 6617 (F).

The pollen of Weberbauer 6617 included a large number of shrivelled
grains.

Citronella sarmentosa (Baill.) Howard. — 32.3 « 34 ,».— Prony
1576—-A (A).

No distinct germ-pores were evident in the uniformly foveate (‘‘pitted’’),
oblately spheroidal grains studied.

Citronella vitiensis Howard. — 29.7 & 25 ».— Degener & Ordonez
13773 (A).

260 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxur

This species is similar to C. SmytAii in having ellipsoidal pollen grains
with luminous granules in many lacunae of the foveate-reticulate exine.

Citronella Smythii (F. v. Muell.) Howard. — Fics. 15, 15A.— 36
x 32.5 p.— Michael 203 (G); Doggrell, 14-12-29 (A); Francis, without
number (A).

Pollen grains ellipsoidal as in C. samoensis; faintly outlined, circular
germ-pore; foveae (“pits”) of exine of variable size, the largest usually
restricted to the poles thus giving that region a foveate-reticulate appear-
ance; luminous granules occur in some of the larger foveae. Some imma-
ture grains were present in the available material.

Citronella philippinensis (Merr.) Howard. — Fics. 20, 20A. — 33.7
xX 32.9 p.— Ramos 33267 (A

The exine is foveate essentially as in C. Smythii. In expanded grains,
the exine bounding the larger foveae, particularly in polar regions, is
fragmented thus imparting a samewhat beaded-reticulate appearance.

Citronella samoensis (A. Gray) Howard. — 29.6 & 27.7 p.— Chris-
tophersen 2721 (A).

The germ-pore is rarely evident as a circular area; the exine is finely
and uniformly foveate. Many shrivelled grains were apparent in the ma-
terial studied.

Citronella lucidula (Sleumer) Howard.— 27.5 , (diameter). —
Franc 1282, ISOTYPE (A).
The spheroidal pollen grains have a foveate-reticulate exine.

10. DENDROBANGIA Rusby

Grains oblately spheroidal to ellipsoidal; tricolporate ; furrows elongate
and smoothly outlined; germ-pore circular, in expanded grains, the pore
bounded peripherally by a rim as in Citronella Gongonha; exine foveolate
(finely “‘pitted”’).

Dendrobangia boliviana Rusby. — Fics. 22, 22A.—24.8 « 30.3
p. — Rusby 1694 (G); Persaud 113 (NY).

11. PLATEA Blume

Grains ellipsoidal to oblately spheroidal; tricolporate; furrows elongate;
germ-pore circular to lenticular; exine rather uniformly foveate except
for immediate margin of the psilate furrow; foveation similar to that seen
in Citronella Gongonha and C. ilicifolia,

Platea philippinensis Merr. — Fics. 23, 23A.— 34.3 & 32.4 p.—
Elmer (Phil. Bur. Sci.) 9777, isotypE (A).

In the preparations studied, the furrows do not open widely but are
constricted (i.e. unopened) in the equatorial region thus obscuring the
germ-pore (see Fig. 23A).

Platea latifolia Bl. — 29 & 30 ».— Clemens 32396 (C).

While the furrows are not markedly constricted at the equator, this

1952] DAHL, MORPHOLOGY OF THE ICACINACEAE 261

specimen resembles P. philippinensis. The exine differs somewhat from
that of the last species in that many of the foveae are aligned thus im-
parting a somewhat striate appearance.

Platea parviflora Koord. & Valet.— 24 X 20.2 p.— Clemens 11163
A).

In the limited material available for study, the mid impressed fur-
rows are constricted at the equator in the germ-pore region

Platea hainanensis Howard.— 24 X 25.8 teas 64940 (A).
In some pollen grains, refractive, lenticular bodies are located in pairs
on either side (either polar or meridional) of the germ-pores.

12. HUMIRIANTHERA Huber

Grains oblately spheroidal to ellipsoidal; tricolporate with germ-pore
circular in outline; exine foveolate

Humirianthera rupestris Ducke. — Fics. 29, 29A.— 30 X 25.7 um.
— Ducke 25288, 342 (US).

The germinal furrows are usually constricted at the equatorial region
as in Jcacina senegalensis. Further, pollen grain shape and character of
exine are as in Jcacina. Not infrequently, luminous fragments are visible
in the furrow regions. The specimen (Ducke 342) belongs here according
to Howard (17) (cf. Figs. 30, 30A). Its pollen grains differ in being
somewhat larger (38 X 36 y»). Abundant starch (diameter of starch
grains up to ca. 4.5 ») also occurred in some pollen grains of this speci-
men.

Humirianthera crispula Howard.—29 x 30 pw.— Haught 2619,
ISOTYPE (G).

The oblately spheroidal grains are foveolate as in H. rupestris. The
available material was inadequate.

13. ICACINA Juss.

Grains ellipsoidal; tricolporate, the furrows elongate; germ-pore large,
circular to elliptical, occasionally bridged by strands of exine (see Fig. 24).
Frequently, that portion of the exine which is directly above the germ-
pore is somewhat depressed (i.e. apparently rests upon the poral mem-
brane) thus contributing to the “shouldering” effect evident at the polar
margins of the germ-pore. Exine foveate to foveate-reticulate.

Icacina Mannii Oliv. — Fic. 24.— 63 & 56 p.— Zenker 475 (G).

The exine is similar to that seen in Lavigeria, Pleurisanthes, and Leretia.
Partial dissociation of such a foveate-reticulate surface would theoretically
yield the surface found in Alsodeiopsis.

Icacina senegalensis A. Juss. — 58.9 & 45.7 ».— Rea 1671 (A).

The material was rather inadequate. Many distorted grains were ob-
served. However, the texture of the exine was essentially as in 7. Mannit.

262 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxim

14. LAVIGERIA Pierre

Grains ellipsoidal; tricolporate; furrows = equatorial germ-
pore with irregular (ragged) margin; exine foveat

Lavigeria salutaris Pierre.— Fics. 25, 25A.—52.5 & 46 ».—
Mildbraed 10536 (A).

15. LERETIA Vellozo

Grains ellipsoidal; tricolporate; furrow elongate with irregular margin;
occasionally the furrow bridged by slender strands of exine; germ-pore
poorly defined, but when evident, circular with a ragged, fimbriate margin;
exine foveate; foveae somewhat enlarged at the polar ends of the grain
where the surface is essentially foveate-reticulate.

Leretia cordata Vell. — Fic. 26.— 60.7 & 52.3 p.— Klug 2967 (G),
641 (F); La Cruz 4235 (G); Blanchet 2347 (F); Spruce 4492 (NY).

The character of the foveation as well as the absence of echination of
the exine renders this species distinct from the genus Mappia.

16. PLEURISANTHES Baillon

Grains ellipsoidal to prolate; tricolporate; furrows elongate with ir-
regular margins, occasionally bridged by strands of exine; germ-pore
either not evident or infrequently, but poorly, defined in the equatorial
region of the furrow; exine rather uniformly foveate.

Pleurisanthes parviflora (Ducke) Howard.— Fic. 27.— 65.2
34.4 p.— Krukoff 6954 (A).
The pollen grains are characteristically prolate in shape.

Pleurisanthes flava Sandwith. — Fic. 28.— 49.3 & 37.5 p.— Sand-
with 590, IsoTYPE (NY).

This species is differentiated from P. parviflora primarily on the basis
of its size and ellipsoidal shape. Shrivelled grains, as well as one abnormally
large grain (ca. 72 » long), were present in the observed material.

17. APODYTES E. Meyer ex Bernh.

Grains oblate, triangular in polar view; three (rarely four, see Fig. 37B)
germinal-pores which also represent the furrows. In this respect, note
the remarks applying to A. yunnanensis and A. dimidiata. The germinal
apertures are circular to lenticular in outline; exine varying from foveate-
reticulate to reticulate.

Apodytes yunnanensis Hu.— 18.4 & 25.8 ».— Wang 74304 (A).

This species and A. dimidiata are of interest in possessing both fur-
rows (colpi) and pores as individually visible structures. In A. yunnanen-
sis the length of the slit-like colpus is somewhat greater than the diameter
of the germ-pore.

Apodytes dimidiata E. Mey.— 22 29.6 u.— Imperial Forestry
Institute 42 (A).

1952] DAHL, MORPHOLOGY OF THE ICACINACEAE 263

In this species the germ-pore is almost, but not quite, coincidental with
the separately visible colpus. Granular, refractive, lenticular boclies, some-
times three in number, are visible near the germinal apertures of occasional
pollen grains.

Apodytes Gardneriana Miers. — Fics. 35, 35A.—27 X 31 p.—
Silva 12 (A).

The foveate reticulation seems relatively massive for a grain of this
size. The germ-pore margins are usually foveolate.

Apodytes cambodiana Pierre. — Fics. 36, 36A.— 25.2 K 33.4 p.—
Clemens 3891 (A

In polar view, one side (representing one “hemisphere’’) is distinctly
more massively reticulate than is the other side (see Fig. 36). Since there
were no tetrads (quartets) present in the preparation, it is not possible to
state which side is distal with reference to the tetrad.

Apodytes brachystylis F. v. Miill.— Fics. 37, 37A, 37B.— 27 X
36.7 p.— Kajewski 1184, 1380 (A).

In the observed material, the germinal aperture as seen in equatorial
view (see Fig. 37A) is rather constricted. The somewhat ragged margin
of the aperture is suggestive of that seen in A. cuminensis. Occasional
grains with four apertures were noted (Fig. 3

podytes beninense Hoch.— Fics. 38, 38A.—33.5 x 39 p—
Elliot 4712 (G).

Apodytes cuminensis Hook. f.— Fics. 39, 39A.— 44.5 & 53.5 p.
— Eala 1930 (F).

This species has the largest pollen grains of the seven examined. It is
comparable in size to Rhaphiostylis cordifolia. The reticulate exine is very
similar to that of A. beninense. Not infrequently, there are bright granules
on the floor of the lacunae.

18. ANISOMALLON Baillon

Grains markedly flattened in equatorial view, distinctly triangular in
polar view; pores three, circular to lenticular, apparently coincidental with
the furrows (i.e. triaperturate); exine foveate-reticulate, the foveation
relatively massive for a grain of this small size.

Anisomallon clusiaefolium Baill. — Fics. 34, 34A.— 13.7 & 22.5
p. — Prony 1731 (A)

Anisomallon is well differentiated from the other genera having scalari-
form perforations in the vessels of the secondary xylem on the basis of its
triangular, flattened, and foveate-reticulate pollen grain.

19. LEPTAULUS Benth.

Grains oblately spheroidal to spheroidal; triaperturate; characteristically,
there are three conspicuous germinal openings (furrow and pore apparently
coincidental). Grains with an additional germinal opening were noted

264 JOURNAL OF THE ARNOLD ARBORETUM | [voL. xxxur

in both L. daphnoides and L. grandifolius; exine varies from foveolate to
foveolate-rugulate with the “polar” regions usually differentiated as es-
sentially bald zones. This genus is rather sharply set off from other genera
of the family in its distinctive pollen grain form.

Leptaulus daphnoides Benth.— Fic. 32.—48 , (diameter). —
Linder 872 (A); Mann 806 (G).

The exine of this species is so finely foveolate that the general effect is
almost smooth. As is evident in Fig. 32, the foveolae are so arranged that
a semblance of finger printing or minute rugulation is attained. The
“nolar” regions are delicately differentiated by a smoother texture and
thinner wall than in the balance of the grain. Infrequently the “polar”
region is visible as a depression.

The three (rarely four) conspicuous germinal apertures have a roughly
circular outline. In one specimen (Mann 806, labelled ZL. species), a fluting
around the apertures derived from thickening plus alignment of foveolae
is somewhat suggestive of Gastrolepis austrocaledonica and certain species
of Stemonurus.

Leptaulus grandifolius Engler. — Fics. 33, 33A.— 42.8 pw (diam-
eter). — Zenker 14 (G).

Approximately 19% of the grains have four germinal apertures. The
aperture is indistinctly defined and while no definite pore is visible the
equatorial region is conspicuously granular-foveolate.

In our preparations, a well-defined bald zone occurs at the “poles”. The
wall of this region lacks the rigidity of the Heavier, foveolate wall. Con-
sequently, the bald area is usually seen as a “polar” depression. In many
grains, exine structure is not uniform, the most massive foveolation and
granulation occurring at the periphery of the bald zone and the germinal
openings. Leptaulus daphnoides is differentiated from this species in its
finer foveolation, the clearly defined germinal apertures, and its less sharply
defined ‘‘polar” zone.

20. RHAPHIOSTYLIS Planch. ex Benth.

Grains oblate; triangular in polar view; tricolpate; three germinal
pores apparently coincidental with the furrows, usually of lenticular
shape; exine reticulate, not infrequently with luminous granules on the
floor of some lacunae, the reticulum beaded when viewed at lower focal
planes.

Rhaphiostylis cordifolia Hutch. & Dalz.— Fic. 40.— 42.3 x S51
pb. — Linder 1218 (A).

Rhaphiostylis ferruginea Engler. — Fics. 41, 41A.— 32.5 « 43
p. — Zenker 65 (C).

Rhaphiostylis latifolia Pierre. — Fics. 42, 42A.—31.2 * 42.5 um.
— Zenker 2242 (A

Rhaphiostylis fusca (Pierre) Pierre.— 25.2 * 36.4 yw.— Klaine
3348 (A).

1952] DAHL, MORPHOLOGY OF THE ICACINACEAE 265

21. GONOCARYUM Miquel

Grains oblately spheroidal; germinal furrows three, rarely four, without
distinct pores, the furrow in at least one species bridged by strands of
substance related to the contiguous reticulum; exine lophate-reticulate,
appearing buttressed or somewhat beaded at sub-surface focal planes;
luminous granules evident in many lacunae.

Gonocaryum longeracemosum King.— Fics. 44, 44A.—-31.7
39.5 p.— Sargent (Singapore Bot. Gard.) (A).

The three (rarely four) germinal furrows are frequently bridged by
strands (see Fig. 44). In polar views (see Fig. 44), the reticulum of the
exine is composed of ca. 97 lacunae.

Gonocaryum cognatum Elmer. — Fics. 45, 45A.— 36.8 X 39.6 py.
— Ramos & Edano 44073 (A).

The reticulum of the exine is more massive than in G. longeracemosum.
In polar view, approximately 81 lacunae are visible in the reticulum. The
usually lenticular furrow-regions are little differentiated from the rest of
the exine and, as a consequence, are frequently difficult to delineate, in
equatorial view.

Gonocaryum calleryanum (Baill.) Becc.— Fics. 46, 46A.— 31.6
33.2 ».— Phil. Bur. Sci. 33 (A).

While the reticulum is less massive than in the previous two species, the
basally buttressed or beaded nature of the reticulum is more evident in
this species than in G. cognatum and G. longeracemosum. In pollen grains
having polar orientation (see Fig. 46) there are approximately 48 lacunae
visible in the exine reticulum. In this species, the lenticular, germinal fur-
rows are quite distinctly differentiated from the adjacent reticulum.

Gonocaryum fuscum Hochr. — 26.4 & 31.3 p.— Sargent (without
number) (A

Approximately 54 lacunae are visible in the reticulum of grains having
polar orientation. The germinal furrow is similar to that in G. calleryanum.

22. MERRILLIODENDRON Kanehira

Grains oblately spheroidal; furrows three without visibly differentiated
germ-pores; islands of foveolate-areolate exine within the furrow region
somewhat as in Alsodeiopsis; exine foveolate to granular-foveolate-areo-
late.

Merrilliodendron rotense Kanehira.—37 43 p.— Kanehira
1775, TYPE (NY); Glassman 240 (A).

23. ALSODEIOPSIS Oliv.

Grains ellipsoidal; tetracolporate; characteristically four furrows which
do not expand widely, the furrow margin irregular, a definite pore difficult
to determine. However, in A. Schumanniit and A. Zenkeri several grains
with more or less rectangular pores were noted. The exine is provided with

266 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxxur

markings that seem best described in terms of either a reticulum or a
foveate surface so incomplete that no intact lacunae are present, i.e.
areolate. It is a surface characterized by interstices separating small areas
of exine. Bounding the furrows is a finely granular-foveolate zone. Lumi-
nous islands of material occur both on the floor and along the margin of
the furrow. Two thickened sub-surface rings (apparently costae aequa-
toriales) in the pollen wall encircle the pollen grain in the equatorial region,
thus imparting an unusual “belted” appearance. In A. Schumannii the
exine varies from foveolate to somewhat areolate.

Alsodeiopsis Schumannii Engler.— 65.9 60.5 ».— Braun 787
(A)

One germinated pollen grain showed the pollen tube emerging from the
equatorial region (potential germ-pore) of the furrow.

Alsodeiopsis Staudtii Engler. — 52.2 & 47.6 ».— Schlechter 13071
A).

The equatorial belting effect characteristic of Alsodeiopsis is ied
prominent in this species. Occasional pentacolporate grains occur

Alsodeiopsis Welwitschii Oliver. — Fics. 43, 43A.— 68.3 X 62.5
p. — Zenker (F, 765310).

Alsodeiopsis Zenkeri Engler. — 68.5 61.5 p.— Zenker 152 (G).

Occasionally, grains with five furrows were observed. The equatorial
region of the furrow, although somewhat irrégularly delineated, apparently
represents the germ-pore.

Alsodeiopsis species. — 63.2 & 56.3 p.— Zenker (F, 765317).

The exine and the furrows of this specimen agree in their characters
with those seen in A. Welwitschii. However, the specimen differs from
this species in having a somewhat smaller pollen grain. Two grains with
five furrows were noted.

24. IRVINGBAILEYA Howard

Grains markedly oblate; typically triangular in polar view; commonly
three pores, although our preparations included four-pored grains to the
extent of 15.7%, the pore surrounded by a foveolate zone of rather limited
extent; exine smooth.

Irvingbaileya australis (White) Howard. — Fics. 60, 60A, 60B. —
15.3 & 23.4 ».— Tardent 259 (A).

This species was previously included in the genus T'ylecarpus Engler, the
components of which are now assigned to the genus Medusanthera. The
pollen grains of Medusanthera are distinct from those of J. australis in
being somewhat smaller and in typically having a minutely papillate exine
with broader, foveolate zones (annuli) bounding the germ-pores.

25. GASTROLEPIS Van Tieghem

Grains oblate, triangular in polar view; three (rarely four) lenticular

1952] DAHL, MORPHOLOGY OF THE ICACINACEAE 267

pores of somewhat irregular outline, a foveolate, thickened zone bounding
the germ-pore; exine essentially smooth, although structurally foveolate.

Gastrolepis austro-caledonica (Baill.) Engler. — Fics. 49, 49A.—
18.2 * 21.2 p.— Franc 1791 (G).

26. CODIOCARPUS Howard

Grains oblately spheroidal; three conspicuous circular germ-pores sur-
rounded by a delicately foveolate-granular zone; exine minutely papillate,
giving the over-all general effect of a smooth surface; papillae rather
widely spaced and occasionally taking on the nature of small, short ne

Codiocarpus Merrittii (Merr.) Howard. — Fics. 47, 47A. —
27 ». — Ramos (Phil. Bur. Sci. 40837, ropotype) (A); Elmer 12622 (6).

Howard’s (21) establishment of a distinct generic concept for this
species is supported by data on pollen grain morphology. From this view-
point the species is distinct from Stemonurus in which genus it was pre-
viously included. Likewise, the pollen grains of the genera Lasianthera,
Gastrolepis, Irvingbaileya, Urandra, and Discophora are distinct from
those of C. Merrittii. In the case of two species of Medusanthera (M.
laxiflora and M. papuana), the exine is minutely papillate and the poral
zones are foveolate, somewhat as in C. Merrittii. However, in each case
the pollen grains are decidedly smaller than in this species.

Codiocarpus andamanicus (Kurz) Howard.— 10.4 * 14.6 p.—
King’s Collector 3-28-94 (A).

Despite some immaturity of the limited material, this species appears
to be closely similar to C. Merritti in its exine and moderately developed
annulus. The difference in size of pollen grain may not be as great in fully
mature specimens.

27. STEMONURUS Blume

Grains markedly oblate; three, very rarely four, pores of circular out-
line, each surrounded by a relatively wide (averaging perhaps 2 yw in
width) foveate zone, the annulus, which in face view resembles a lace
collar; exine structurally foveolate-rugulate-papillate but usually giving
an essentially smooth general effect. In some instances the papillae ap-
pear to be nearly baculiform. Despite this, the outline of the pollen grain
is almost smooth. The highly distinctive rugulate (sense of Iversen and
Troels-Smith, 25) character was most evident in S. Aainanensis although
delicate degrees of rugulation were also noted in other species (e.g. S.
axillaris).

Throughout the complex of species studied there is a characteristic pol-
len morphology particularly with respect to the presence of the con-
spicuously foveate annuli (germ-pore margins). Different opinions are
held with respect to the applications of the name Stemonurus (cf. 16, 28,
23). The generic concept utilized here includes species originally de-
scribed as members of the genus Gomphandra Wallich ex Lindley.

268 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxxmI

Stemonurus affinis Miers. — 15 * 23 ».— Boeea 8028 (A).

The exine, which in general effect is almost smooth, is very minutely
foveolate-rugulate-papillate. The remarkably conspicuous annulus may
have a radius of as great as ca. 5 p.

Stemonurus apoensis Elmer. — 14.3 & 18.4 ».— Elmer 15416 (A).

Stemonurus axillaris (Wall. ex Lindl.) Miers.—19 & 24.2 p.—
Anglade, without number (A).
The exine is minutely papillate-rugulate in this species.

Stemonurus Chingianus Hand.-Mazz.—19 23.6 p.— Ching
5226, ISOTYPE (NY); Pételot 5450 (A).

Stemonurus Cumingianus Miers. — Fics. 50, 50A.—16 19 4.
—Wenzel 696 (A), 350 (A, G).

Stemonurus dolichocarpus (Merr.) Howard.—16 21.3 p.—
Toroes 844 (A).

In this species, the exine is quite smooth (psilate). The annulus is of
comparatively limited extent.

Stemonurus flavicarpus Elmer. — 14 x 20.2 ».— Elmer, Phil. Bur.
Sci. 11330 (A)

Stemonurus hainanensis (Merr.) Hu.— 16 X 23 ».— Henry 10738
(A); How 70388 (A).

The exine, while smooth in general effect, is noteworthy for its distinc-
tive minutely rugulate character (cf. Fig. 1 of Plate II of Faegri and Iver-
sen, 12).

Stemonurus javanicus Bl. (var. lanceolata K. & V.).— 16.3 %& 21.7
p. — Collector unknown 751 (NY).

Stemonurus luzoniensis (Merr.) Howard.—18 27 p».— Elmer
13110 (G).

In grains having polar orientation, the annulus ascends abruptly to the
pore. The exine is very faintly rugulate.

Stemonurus oppositifolius (Pierre ex Gagnep.) Howard. — 15.3 «
23 p.— Clemens 3861 (C).
The annuli are abruptly ascending.

Stemonurus polymorphus Miers. — 17.6 * 25.6 ».— Herb. Wight,
Pen. India Orient., 427 (G).

Stemonurus Yatesii (Merr.) Howard. — 11.2 & 16 ».— Yates 2637,
TYPE (C), 2248

The exine differs from the above species of the genus in being finely
papillate with the diminutive papillae appearing spinose as in Medusan-
thera glabra. ‘There is no suggestion of the distinctive, minutely rugulate
character in this species. On the basis of pollen characters, this species
seems more closely allied to Medusanthera than to other species of Stemon-
UUs,

1952] DAHL, MORPHOLOGY OF THE ICACINACEAE 269

Stemonurus mollis (Merr.) Howard.* — 13.6 19.2 y.-— Pételot
7949, TYPE (A).

Stemonurus Petelotii (Merr.) Howard.** — 21.7 * 25 p.-— Pételot
2647, TYPE (A)

Stemonurus obscurinervis (Merr.) Howard.*** —16 & 23.5 p.—
Pételot 4359, TYPE (A).

28. MEDUSANTHERA Seeman

Grains oblate. Germ-pores more or less circular, typically three, oc-
casionally four, bounded by a foveolate and somewhat thickened zone,
striate in appearance especially when seen in polar view (i.e. optical sec-
tion). In general effect the exine is smooth. Structurally the surface is
beset with minute papillae. Usually the papillae are so diminutive that
it is difficult to determine whether their form is either that of exceedingly
short spines or that of blunt pegs. It appears, however, that such species
as M. samoensis and M. carolinensis have very short spinose papillae.

Medusanthera carolinensis (Kaneh.) Howard.— Fics. 56, 506A.
—11.6 * 15.5 ».— Kanehira 1882 (NY).

Throughout, the pollen grain of this species is essentially identical
with that of M. glabra.

Medusanthera glabra (Merr.) Howard. — Fics. 55, 55A.—9.4 X
15.3 p.— Wenzel 1029 (A).

The pollen grain of this oe has the appearance of being a smaller
edition of that of M. samoen

Medusanthera laxiflora Be re Howard. — Fics. 53, 53A.—17 X
20.2 ». — Brass 2724 (A); Wenzel 3224 (A).

The pollen of the Wenzel specimen (labelled “Stemonurus laxiflorus”),
while agreeing in its morphology with the Brass collection, differs in hav-
ing a somewhat smaller pollen grain (14.6 & 18.3 ,).

Medusanthera papuana (Becc.) Howard. — Fic. 52.— 16.4 & 21.2
p. — Schlechter 17913 (C
The available pollen material was largely immature.

Medusanthera samoensis (Reinecke) Howard. — Fics. 54, 54A.—
14.6 & 19.4 p.— Christophersen 3061 (A).

The exine is minutely roughened by chiefly spinous papillae as in M.
carolinensis.

Medusanthera vitiensis Seeman. — Fics. 51, 51A.— 15 & 19.3 p.—
Smith 578 (G); Storck 877, isotTyPE (G).
The pollen of this species is quite similar to that of M. laxiflora.
temonurus mollis Ses Howard, comb. nov. Gomphandra mollis Merr.,
alse Arnold Arb. 23: 175.
* Stemonurus Bereletl: “Mer Howard, comb. nov. Gomphandra Petelotii
tes. , Jour. Arnold Arb. 23: 176.
wee Stemonurus Mane ate "“(Merr.) sr ede comb. nov. Gomphandra
obscurinervis Merr., Jour. Arnold Arb. 19: 44.

270 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxx

29, URANDRA Thwaites

Grains oblate, more or less triangular in polar view; germinal apertures
three, of circular or lenticular outline; pore margin ornamented by a
thickened, foveolate, often sinuous zone (annulus); exine foveolate-papil-
late, frequently of smooth general effect.

Urandra Brassii Howard. — 9.4 & 12.3 p.— Brass 7421, TYPE (A).
The exine is minutely rugulate. The annulus and pore are typical for
the genus, resembling those of U. secundiflora.

Urandra dolichophylla (Merr.) Howard.—9 X 15.8 j.— Elmer
21415, ISOTYPE (C, G).
The pollen of this species is very similar to that of U. scorpioides.

Urandra Hallieri Merr.—11 15.4 ».— Hallier 497 (A).

The exine is finely papillate, having a smooth general effect (as in U.
dolichophylia).

Urandra scorpioides (Becc.) O. Ktze.— Fics. 58, 58A.—7.6 xX
14 ». — Toroes 4323 (A); Wood 2392 (A).

The exine is so finely foveolate-papillate that in general effect it appears
smooth.

Urandra secundiflora (Blume) O. Ktze.— Fics. 57, 57A.—9 X
14.6 ». — Haviland & Hose 1544 (G).
The exine is so finely foveolate that it seems almost smooth.

Urandra umbellata (Becc.) O. Ktze.— Fics. 59, 59A.— 13.7 xX
17.7 p.— Haviland 1782 (G); Holttum 31267 (A).

The exine is quite distinct from that of the other species of the genus
cited above. The surface is irregularly papillate-rugulate (somewhat like
Fig. 2, Plate II of Faegri and Iversen, 12). The exine of Cantleya cornicu-
Jata is of somewhat similar structure although the papillate foveation is
of a much finer and more delicate order.

30. CANTLEYA Ridley

Grains markedly oblate, triangular in polar view; germ-pores three,
the poral aperture peripherally differentiated by a remarkable foveate
fretwork of rods representing the regions between the foveae in the locally
thickened exine; exine minutely papillate-foveate-rugulate, the foveae so
numerous that the surface appears as if it were faintly papillate.

Cantleya corniculata (Becc.) Howard. — Fics. 61, 61A.—11 17
p. — Ex. Herb. Hort. Bog. 265770 (C).

31. LASIANTHERA P. Beauv.

Grains markedly oblate; pores commonly three or four, rarely five or
six, rendered conspicuous by a thickened peripheral zone (annulus) ; exine
very finely papillate giving an almost smooth general effect.

1952] DAHL, MORPHOLOGY OF THE ICACINACEAE 271

Lasianthera africana Beauv. — Fics. 48, 48A.— 14.6 * 21.2 p.—
Conrau 41 (A, B); Ghesginere 2910 (G).

The distinctive pollen grain of this species makes it clear that “Lasian-
thera austrocaledonica’ of Baillon is best assigned elsewhere. Howard (16)
has already advocated the assignment of this species to the genus Gastro-
lepis on the basis of other characters. Likewise, the pollen grains of Medus-
anthera, Stemonurus, and Discophora are each distinct from those of L.
africana. The dimensions given apply to Conrau 41, the pollen of which
generally has four pores or rarely three, five, or six pores. The Ghesginere
specimen, consistently triporate, differs in having pollen grains which are
about 30% smaller.

32. NATSIATUM Buch.-Ham.

Grains oblately spheroidal; germ-pores circular varying in number from
5 to 7 with 6 being the most frequent. Of these, four are usually located
on the “equator” with the remainder occurring fairly close to the “equa-
tor” in either the upper or lower hemisphere; exine foveolate-papillate.

Natsiatum herpeticum Buch.-Ham. — Fic. 31. — 28.8 » (diameter).
Parkinson 3927 (A); Griffith 828 (G).

33. HOSIEA Hemsley & Wilson

Grains oblately spheroidal; tricolpate, rarely four-furrowed, the fur-
rows elongate (see Fig. 63) with very ragged, torn margins; frequently
with irregular bridges or fragments of exine in the equatorial region;
germ-pore not distinct; exine irregularly ornamented with short spines
and rather blunt papillae, the texture between the projections rather finely
granular-foveolate.

Hosiea sinense (Oliver) Hemsl. — Fic. 63. — 45.5 & 56.2 p.— Wil-
son 960, 960a (A); Yu 1932 (A).

One dwarf pollen grain (diameter ca. 28.8 4) was observed. Hosiea
sinense is the only member of the tribe /odeae having elongate furrows.
A somewhat similar furrow is seen in the tribe Sarcostigmatece.

34. MAPPIA Jacquin

Grains ellipsoidal to oblately spheroidal; furrows three to five clepending
upon the species concerned; margins very ragged and not infrequently
bridged by strands of exine material (see Fig. 64A) with small fragments
of exine in the furrow region; germ-pore indistinct; exine foveate-punc-
tate with very short, rather widely spaced spines, which appear in reduced
light as luminous spots, some spines having the appearance of “reduced”
(or little-developed) papillae.

Mappia angustifolia Griseb. — 55.7 & 51.2 p.— Wright 2638 (G).
This species is characteristically tricolpate.

272 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxxm

Mappia longipes Lundell. — 47 X 44.6 ».— Matuda 4798, TyPE (A).

This species is tetracolpate (stephanocolpate). Many of the massive
“spines” of the exine are blunt or baculiform. The longest of these have a
length of ca. 1.4 p.

Mappia mexicana Robins. and Greenm. — Fics. 64, 64A.— 53.6 X
51 p».— Pringle 5094, 6645 (G).

Of the pollen grains examined, 56% were tetracolpate whereas 44% were
pentacolpate. No tricolpate grains were observed. Despite the outward
differences in the two specimens noted by Howard (17), the equivalent
pollen form of the two collections justifies his assignment of them to a
single species.

Mappia racemosa Jacq. var. brachycarpa Griseb. — 51.5 & 52.3 yp.
— Wright 1389 (G).
The pollen of this species is typically tetracolpate.

35. DESMOSTACHYS Planchon

Grains oblately spheroidal; furrows four to nine, about two-thirds the
length of the pollen grain; germ-pores indistinct; exine, particularly in D.
Preussii, foveolate with a minutely striate character (as seen in optical
section) bearing conspicuous short spines.

Desmostachys Preussii Engler. — Fics. 76, 76A.— 73.3 & 78.4 p.
— Zenker 121 (G), 404 (A).

When closed, the germinal furrows are slit-like apertures bounded by
massive spines. The largest of these are ca. 3.8 » long and 2.8 » wide at
the base. Typically the number of furrows is four.

Desmostachys Vogelii Stapf. — Fic. 77.—92 x 99.2 p.— Linder
1345 (A).

The pollen grains of this species possess 6 to 9 furrows. In our material,
47% of the grains had 7 furrows while 50% had 8 furrows. As seen in
perfectly polar view, the furrows are symmetrically distributed around the
equator and are essentially of the same length. However, there are oc-
casional grains which deviate from the usual pattern in having either 6
or 7 normal furrows and an additional one whose length is only 4% to %
that of the others.

The spines of the exine are of irregular size and distribution and are
commonly shorter than in the previous species. The bald or relatively
smooth zone bounding each furrow (see Fig. 77) is a striking feature of
this pollen grain. Rather massive spines are found at the immediate
margin of the furrow. Luminous flecks of material occur on the floor of
the expanded furrow. The striate nature of the exine is less evident than in
D. Preussit.

The above characters, as well as that of size, render this species quite
distinct from. D. Preussii.

1952] DAHL, MORPHOLOGY OF THE ICACINACEAE 273
36. NOTHAPODYTES Blume

Grains ellipsoidal to spheroidal; tetracolpate or pentacolpate, the fur-
row margin ragged; germ-pore indistinct; exine foveolate, with rather
short massive spines.

Pe a foetida (Wight) Sleumer. — 59 55.6 ».— Gardner

9 (G); Thwaites 492 (G).

e this species, the elongate furrow, of which there are four, is distinc-
tive in possessing a comparatively massive island of exine. This island is
differentiated from the balance of the exine in having a greater frequency
of spines and foveolae. On the basis of furrow structure, this species is
clearly distinct from Mappia mexicana.

Nothapodytes pittosporoides (Oliv.) Sleumer.— 102 X 89.8 p.—
Wilson 3296 (G).

Most of the grains observed were pentacolpate (one tetracolpate grain
was noted). The furrow membrane is papillate-echinate. The exine is
foveolate with scattered very short spines appearing as luminous dots.
This species differs strikingly from its close allies in its relatively gigantic
size. It is also distinct from other species of Nothapodytes in type of hair
in its pubescence (15).

37. DISCOPHORA Miers

Grains oblate; germ-pores circular, typically three, occasionally four,
bounded by a thickened, foveate zone ‘somewhat as in Stemonurus Cumin-
gianus; exine finely echinate, typically with short, luminous spines.

Discophora guinanensis Miers. — 16.7 & 18.8 ».— Klug 3017 (A);
Krukoff 6789 (F

Occasional four-pored grains were observed in the Krukoff collection.

Discophora montana Howard. — Fics. 62, 62A.—15.3 & 19 p.—
Lawrance 535 (G).

The pollen grains of this specimen were infrequently four-pored and,
in addition, showed much variation in size. One giant grain (cliameter,
30.2 ») having nine germ-pores was noted. Pollen grain diameters varied
from 15 to 32.5 p.

Discophora panamensis Standley. — 17.3 & 19 ».— Cooper 12246

The exine of this species with its short, inconspicuous spines is distinct
from that of D. guianensis and D. montana where the pollen grain surfaces
are more evidently spiny.

38. MAPPIANTHUS Hand.-Mazz.

Grains approximately spheroidal; rounded, triangular in polar view;
germ-pores nearly circular, commonly three, with four pores in 15% of
grains observed; the distinctive central aperture surrounded by a thick-
ened, echinate rim which in turn is set off from the surrounding exine, the
floor of the central pore occasionally bearing spines and exine fragments;

274 JOURNAL OF THE ARNOLD ARBORETUM _ [VvoL. xxxilI

exine conspicuously echinate with relatively smooth texture prevailing in
regions between the numerous spines.

Mappianthus iodioides Hand.-Mazz.— Fics. 69, 69A.—30 * 32
p. — Henry 12063 (A).

One large grain, 41.4 » in diameter, was noted.

39. IODES Blume

Grains spheroidal to slightly oblate; germ-pores typically three, more or
less circular (varying from three to five in /. liberica), delineated by a rim
of thickened exine whose inner margin is irregular; exine markedly echinate
with comparatively large massive spines, the surface minutely foveolate
between the spines. Sizes reported for this genus are diameters exclusive
of the spines whose length is reported separately.

Iodes tomentella Miq.— 16.6 » (diameter).— Ex Herb. Miquel,
without number (G).

The longest of the numerous spines of the exine is ca. 1.6 » in length.

Iodes floribunda Merr.— Fic. 65.— 24.8 ,» (diameter).— Toroes
77, ISOTYPE (A).

Iodes africana Welw. — 25 ,» (diameter). — Zenker 303 (G).

The maximum length of the spines is ca. 2.4 p.

Iodes ovalis Blume. — Fic. 66.— 26.2 » (diameter).— Parker 2770

)

This and the above three species are set off from the other investigated
species of the genus by their comparatively small-sized pollen grains.

Iodes kamerunensis Engler. — 34 » (diameter). — Zenker 460 (G).

The conspicuous tapering spines of the exine may attain a length of ca.
7.3 p.

Iodes aed (Hance) Hemsley. — 34.6 ,» (diameter).— Liang
61578 (N

eee ‘6 Howard (personal communication) this species should be
referred to 7. ovalis. However, this specimen is listed separately under the
Hemsley combination since its pollen grains are of larger size and possess
more massive spines than is the case in J. ovalis. In both species, the spines
of the exine are ca. 3.3 p» in length. Until more abundant material is
available it seems best to list these observations separately.

Iodes liberica Stapf. — Fic. 67. — 44.3 y (diameter). — Linder 1102

In this species germ-pore numbers vary from three to five with about
88% of the grains being four-pored. In the single five-pored grain ob-
served, one of the germ-pores was approximately one-half the usual size.
The spines of the exine are massive and relatively long (ca. 5.8 w in
length).

Iodes philippinensis Merr.— Fic. 68.— 53.8 , (diameter). —
Ramos, Phil. Bur. Sci. 42567 (A).

1952} DAHL, MORPHOLOGY OF THE ICACINACEAE zis

One four-pored grain was noted. The massive spines average about 2.8
» in length.

40. POLYPORANDRA Becc.

Grains essentially spheroidal; germ-pores three, similar in form to Jodes;
exine conspicuously echinate with massive, widely spaced spines, granular-
foveolate between the spines.

Polyporandra scandens Becc.— Fic. 70.—24 yw (diameter). —
Kajewski 2206 (A); Schlechter 17835 (C).

The pollen of Polyporandra is similar to that of Jodes floribunda.
differs, however, in its more widely scattered spines and in its somewhat
less conspicuous annulus.

41. MIQUELIA Meissner

Grains spheroidal; germ-pores three, circular with a thickened margin;
exine echinate with prominent massive spines, the exine between the
De minutely foveolate-striate, particularly as seen in optical section.

uelia Cumingii Baill. — Fic. 74.— 30.2 » (diameter). — Phil.
Bur. Sci 49997 (C).

42. PHYTOCRENE Wall.

Grains spheroidal; germ-pores three, of roughly circular outline, the
poral margin only slightly thickened; exine minutely echinate with numer-
ous short spines.

Phytocrene Blancoi (Azaola) Merr. — Fic. 75.— 22 ,» (diameter).
— Loher 13821 (A); Wenzel 44 (G).

Phytocrene bracteata Wall.— 19.5 » (diameter).— Wallich 4947
(G).
In this somewhat immature specimen, the exine was minutely echinate-
foveolate.

Phytocrene dasycarpa Miq.— 18 » (diameter). — Warburg 16683
A).

43. POLYCEPHALIUM Engler

Grains spheroidal to somewhat oblate; germ-pores three, infrequently
four, characteristically circular, the region bounding the pore thickened as
in Miquelia Cumingii, Iodes floribunda, and Chlamydocarya capitata;
exine echinate with numerous diminutive spines.

In this genus, as well as in Chlamydocarya, the exine development is
in the direction of a smooth general effect.

Polycephalium Poggei Engler.— Fic. 72.—20 ,» (diameter). —
Zenker 313 (A).

276 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxxmr

While the pollen of this species is in general similar to that of Phyto-
crene Blancoi, it is distinct in its smaller spines and in its germ-pore with
a more markedly thickened margin

44, CHLAMYDOCARYA Baillon

Grains spheroidal; germ-pores three, circular with margins thickened

as in Polycephalium; exine minutely echinate as in Polycephalium.
Chlamydocarya capitata Baill. — Fics. 73, 73A.— 16.4 p (diameter).
— Linder 1076 (A).
The only material available included a number of immature pollen
grains.

45. RHYTICARYUM Beccari

Grains oblately spheroidal to spheroidal; germinal apertures obscure in
this material, occasional suggestions of porate apertures without annuli;
exine finely echinate with relatively short spines and occasional rather
blunt papillae.

Rhyticaryum elegans Schellenberg. — 27.4 » (diameter). — Schlech-
ter 18681 (C).

The present analysis is provisional in that it is based upon the only
available material, which was poor and immature. Many of the grains
were still adherent in collapsed tetrads.

46. SARCOSTIGMA Wight and Arnott

Grains spheroidal; germinal apertures three (S. Wallichii) ; exine echin-
ate with numerous short to medium length spines, averaging ca. 1 » long,
foveolate between the spines.

The germinal apertures are obscure in the available material of the
first three species. Some grains give the impression of being inaperturate
whereas in others a single lenticular germinal aperture is evident. Rarely
in these species there is a suggestion of three such apertures. However, in
S. Wallichii it is clear that three germinal apertures are present (see be-
low).

Sarcostigma philippinensis Merr.— Fic. 71.—35 m (diameter).
— Wenzel 987 (A).

The numerous spines of the exine are noticeably variable in size. In
length, they vary from ca. 1.7 » down to very short, still rather massive
spines.

Sarcostigma Horsfieldii R. Brown.— 42.2 » diameter (Warburg),
30 » diameter (Sargent). — Warburg 1929 (A); Sargent (without num-
ber) (A).

The spines of the exine are in general shorter, somewhat less conspicuous,
and of more uniform size than in S. philippinensis. In the Sargent speci-
men, spines varied in length from ca. 0.8 to 1.

1952] DAHL, MORPHOLOGY OF THE ICACINACEAE 277

Sarcostigma surigaoénse Elmer ex Merrill. —30 » (diameter). —
Elmer 13732 (G).

Sarcostigma Wallichii H. Brongn.— 28 ,» (diameter). -— King’s
Collector 2-17-94 (A

It is clear from analysis of this specimen that there are three germinal
apertures of lenticular outline. Aperture margins are unspecialized. The
slender spines of the exine attain a length of ca. 2.2 pu.

47. PYRENACANTHA Wight

Grains spheroidal; germinal apertures obscure, varying from three to
six in number; exine minutely papillate-echinate-foveolate, appearing rela-
tively thick.

In P. volubilis, in infrequent pollen grains having apparently equatorial
orientation, the germinal apertures are visible as tightly closed slits ca. 3
pu long, lying parallel to the equator. In consequence, these apertures are
only slightly differentiated when seen in “polar” view. The typical num-
ber of apertures appears to be three in this species, although in the speci-
men examined, 20% were interpreted as having four pores. Germinal
pores were more conspicuous in the specimen of P. vitifolia. Here the cir-
cular pores varied in number from three to six with ca. 56.5% of the
grains having four pores. The pore membranes were flecked with granules.
In the case of P. scandens, the very scanty material suggested a condition
as in P. volubilis.

Pyrenacantha scandens Planch. ex Harv.—39 ,» (diameter). —
Burchell 5214 (G).

Pyrenacantha vitifolia Engler.— 19.7 , (diameter). — Scheffler
345 (A).

Periporate grains are usually ‘“heteropolar.”

Pyrenacantha volubilis Wight. — 27.4 » (diameter) with occasional
(ca. 14%) dwarf pollen grains 17 » in diameter.— Lau 214 (A).

DISCUSSION

Within the Family Jcacinaceae, as presently constituted, there is an im-
pressive array of individually distinct combinations of characters pertain-
ing to the exine (surface) and to the germinal apertures. Fifteen such
combinations have been selected as “pollen types.” It is obvious that
pollen data, together with anatomical and exomorphic data, indicate that
the family is a heterogeneous one. The extent to which genera can be sep-
arated on the basis of pollen morphology is unusual. It is also evident in
some genera that interspecific differentiation with respect to pollen exists.

The list of pollen types of all investigated genera given earlier (pp. 255,
256) will suggest how the pollen characters have been utilized in segregat-
ing and arranging genera. The two main trends which were postulated

278 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxxmt

and followed in this arrangement are given below. With respect to surface
characteristics of the exine, a pattern may be established as follows: —

Echinate Areolate Reticulate Lophate

Papillate Foveate (‘“‘Pitted’’)

7

Psilate (Smooth)

On the basis of germinal apertures the following array can be visualized
within the /cacinaceae: —

Triporate with annuli ——Stephanoporate

Stephanocolpate
Stephanoporate without
annuli

Tricolpate Wie

Triporate without annuli

°

4

Triaperturate

)

Tricolporate

What correlation, if any, exists between a system of genera devised with
reference to these trends of pollen specialization and one based on ana-
tomical characters (Bailey and Howard 1, 2, 3, 4)? An answer may be
sought in the summary graph presented in Text-fig. 1. Anatomical Group
I* is considered to be less specialized than anatomical Groups II and III.
Pollen Type Al* (following the sequence enumerated in the listing on
pp. 255, 256) is assumed, in this family, to be less specialized than Type ox 2
The following conclusions can be drawn from this analysis:
A. The majority of the species having Type A pollen grains ‘eelone to the
less advanced anatomical Group I.
B. Most species of the anatomically intermediate Group II have Type B
pollen
C. The maloriy of species with Type C pollen grains are anatomically
the most specialized in the family (anatomical Group III).
*In an attempt to avoid confusion, Roman numerals and the term ‘ ‘Group” apply
to anatomical aggregations whereas ee letters, Arabic numerals, and the term
“Type” refer to the pollen categories

1952 | DAHL, MORPHOLOGY OF THE ICACINACEAE 279

POLLEN MORPHOLOGY AND WOOD ANATOMY

FAMILY ICACINACEAE

SY

TYPE*
Ww

POLLEN
UBNANO

II IIIA 111B,C,D

ANATOMICAL GROUP (From 1.w. Bailey & R.A. Howard)

(*- LETTERS REFER TO MAJOR TYPE ; ARABIC NUMERALS

REFER TO

INDIVIDUAL EXAMPLES WITHIN THE TYPE. )

280 JOURNAL OF THE ARNOLD ARBORETUM [voL. xxxutr

While the correlation is not perfect, the unusual degree of association
between the pollen and anatomical spectra seems significant. This be-
comes more evident if such association is examined in greater detail. It
is pertinent to point out that each of the so-called major pollen types
represents an attainment series or complex. Further, it should be noted
that species which appear “late” in each pollen type (see Text-fig. 1)
are also anatomically more specialized than those which “initiate” the
type. This is particularly true for pollen types A and C. Pollen type B
proves to be characteristic of genera belonging to anatomical Group II
with the exception of Natsiatum and Hosiea. In the case of type C pollen,
all representatives are anatomically Group II with the exception of Dis-
copnora.

Of particular interest are the pollen spectra of anatomical Group II and
Group III A, the latter of which is made up of unilacunar genera of the
tribe Icacineae. These two groups include the widest range of pollen
grain forms of the four anatomical categories, a condition which suggests
that these groups include forms which are transitional between the less
specialized and the more advanced complexes. This is consistent with the
anatomical observations of Bailey and Howard (1, 2). Leptaulus, Gono-
caryum, and a complex “beginning” with /rvingbaileya and “culminating”
via pollen type B2 in Discophora represent, respectively, what may be
considered to be the three distinct patterns of pollen grain form within
Group II. Anatomically Group H, which is a complex of trilacunar
Icacineae, exhibits a variety of transitional stages in the specialization of
vessels, tracheids, wood parenchyma, and rays. Leptaulus is distinct in
the origin of the simple porous perforations of its vessels while Gonocaryum
is more or less unique in its possession of vessels with scalariform per-
forations of abnormal orientation. It is evident that the two genera just
mentioned are differentiated from their associates in both pollen and
anatomical features.

Group III A is a complex of genera which is anatomically transitional
between the trilacunar /cacineae and the more specialized forms Jodeae,
Sarcostigmateae, and Phytocreneae. In this group some six types of pollen
occur. Genera having pollen of types A2, A5, and C3 also possess vari-
ously shortened, laterally enlarged vessels. This suggests that the trend
to a foveate-reticulate exine in pollen type A2 more or less “culminates”
in the reticulate exine of type AS. Very large, widely-spaced vessels
characterize Merrilliodendron (pollen type A7) while vessels tend to be
radially grouped in Alsodeiopsis (pollen type A8) and in Mappia and
Nothapodytes (pollen type C1). The remaining genus (Desmostachys)
in Group III A is anatomically differentiated in possessing vessels which
tend to occur in tangential groups.

Finally, pollen of the specialized types C2 and C3 occur only in the
remaining unilacunar tribes (Jodeae, Sarcostigmateae, and Phytocreneae)
of anatomical Group III (B, C, D) with the exception of two genera,
Hosiea and Natsiatum which produce, respectively, pollen of types B3
and B4. Hosiea, both in pollen and ray structure, is less specialized than

1952] DAHL, MORPHOLOGY OF THE ICACINACEAE 281

the other components of these tribes. Anatomically, Natsiatum appears
to be similar to Hosiea.

The taxonomic significance of some of these data has already been
suggested. Unfortunately, there have been relatively few attempts to
consider, intensively, the pollen morphology of groups of aggregated species
(cf. Wodehouse, 32; Erdtman, 8, 10; Lindau, 26; Hedberg, 14; et al).
Mrs. Lucy Cranwell Smith (6) has reported briefly on Pennantia of the
Icacinaceae which occurs in New Zealand. In passing, it should be noted
that Sleumer’s (30, p. 337) paragraph on pollen of the /cacinaceae is best
disregarded. In his summary statement, the descriptions of eleven genera
out of the fifteen included are either partially or completely in error.

Engler’s (7) four tribes, Jcacineae, Iodeae, Sarcostigmateae, and Phyto-
creneae are not rendered individually distinct on the basis of either pollen
morphology or anatomy. In the tribe /cacineae, pollen grains with echi-
nate exine and annuli (sem. Jat.) are very infrequent while all but two
genera of the last three tribes have echinate pollen grains with annuli.

In the arrangement of genera with reference to pollen grain type it
is seen that in general those genera which appear early in the taxonomic
sequence also produce less modified (type A) pollen grains while genera
terminating the sequence possess more specialized types of pollen grains
(type C).

Most of the New World genera are characterized by type A pollen.
Only two genera (Discophora and Mappia) of the New World have echi-
nate (type C) pollen grains. In its pollen grain characters, Discophora
finds its closest allies in Old World genera which is also true on the basis
of wood structure and pubescence (15). Mappia resembles certain Old
World genera in possessing cylindrical hairs and radial groups of vessels.

Speculation concerning interfamily relationships seems best delayed
until detailed information derived from intensive studies of the pollen
morphology in other groups is available. Mauritzon (27) and Fagerlind
(11) conclude on the basis of examining some members of the family that
in structure of the gynoecium, including the ovules, the affinities of the
Icacinaceae are with the order Celastrales.

In conclusion, it must be emphasized that from statements concerning
major patterns and trends of pollen grain structure the existence of a
relatively simple, continuous, unidirectional series is not to be inferred.
The striking and seemingly sudden appearance of a notable variety of
pollen types within the family argues against a single, connected evolu-
tionary series. Indeed, in various attempts to organize these genera into
an unrandomized array one cannot escape the impression that the result-
ant scheme is reminiscent of a bird’s-eye view of a shrubbery and not
of just one shrub. In the Jcacinaceae, the aggregations of species into
genera on the basis of largely exomorphic data are on the whole supported
by pollen data. However, clusters of genera represent levels of attainment
(specialization) with respect to the guiding character and are not, in-
variably, indicators of intimately related or recently derived genera. The
oft-repeated remark concerning differential rates of development with

282 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxir

respect to different characters applies here. It follows therefore that the
most dependable clues concerning phylogenetic relationships are to be ex-
pected when information concerning levels of attainment or specialization
for all possible characters is available for comprehensive study. Pollen
characters represent an important element in the total complex of char-
acters which is available for analysis in the very difficult task of achieving
“Phylogeny, resolved.”

SUMMARY

1. A detailed study of pollen of the family /cacinaceae, as presently
constituted, has revealed an impressive assortment of individually distinct
combinations of pollen characters, particularly those which apply to the
exine (surface) and germinal apertures. Fifteen such combinations have
been selected as “pollen types.”

2. Utilization of these types in segregating and arranging genera has
suggested two main trends within the family. First, with respect to sur-
face characteristics of the exine, the following array may be established: —
psilate (smooth) — foveate (“pitted”) — reticulate — lophate — areolate;
psilate — papillate — echinate. Secondly, on the basis of germinal aper-
tures, the following array can be visualized within the /cacinaceae: —
tricolporate — triaperturate — tricolpate — stephanocolpate; _ tricolporate
—triporate without annuli— stephanoporate without annuli — triporate
with annuli — stephanoporate with annuli.

3. In the above text, genera have been arranged with reference to these
trends.

4. A comparison has been made between a system of genera devised
with reference to the above trends of pollen specialization and one devised
independently for the /cacinaceae by I. W. Bailey and R. A. Howard on
anatomical grounds. Such analysis demonstrates that the majority of spe-
cies having the least specialized (psilate, tricolporate) pollen grains belong
to the least advanced anatomical group. Most of the species characterized
by specialized (echinate, porate) types of pollen grains are anatomically
the most specialized in the family.

5. The unusual degree of association between the pollen and anatomical
spectra in this family, made up of approximately 300 species distributed
among 60 genera, is indicative of the significance of intensive morphological
studies of pollen in ultimately phylogenetic problems.

DEPARTMENT OF BOTANY,
UNIVERSITY OF MINNESOTA,

1952] DAHL, MORPHOLOGY OF THE ICACINACEAE 283

i;

mn

own

LITERATURE CITED

BarLey, I. W. & R. A. Howarp. The comparative morphology of the
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22: 125-132. 1941.

. The comparative rig eed of the Icacinaceae.
941

II. Vessels. Jour. Arnold Arb. 22: 171-187.
——— T

~ he comparative morphology of the Icacinaceae.
III. Imperforate tracheary elements and xylem parenchyma. Jour.
Arnold Arb. 22: 432-442. 1941.

———— & —————-.. The comparative morphology of the Icacinaceae.
IV. Rays of the secondary xylem. Jour. Arnold Arb. 22: 556-568. 1941.

CHRISTENSEN, B. B. Om mikrotomsnit af pollenexiner. Medd. Dansk Geol.
Foren. 11: 441-444, 1949.

CRANWELL, L. New Zealand pollen studies I. Key to the pollen grains of
families and genera in the native flora. Rec. Auckland Inst. Mus. 2:
280-308. 1942.

ENGLER, A, Icacinaceae. Nat. Pflanzenfam. III, 5: 233-257. 1893.

ERDTMAN , G. An introduction to pollen analysis. Chronica Botanica Co.,
Waltham, 1943,

———— . Suggestions for the classification of ag a recent pollen
grains and spores. Svensk. Bot. Tidskr. 41: 104-114.

———. , O. HeEpBerc & 7 TERASMAE, Literature on ee XIV.
Geol. Foren. Forhandl. 73: 100-128. 1951.

FAGERLIND, F. Bau des Gynoceums der Samenlage und des Embryosackes
bei einigen Reprasentanten der Familie Icacinaceae. Svensk, Bot. Tidskr.
39: 346-364. 1945.

Faecri, K. & J. IverSEN. Text-book of modern pollen analysis. E.
Munksgaard, fae eras 1950

. HAvirer, Hans. L’ origine et le systeme phylétique des angiospermes.

Archives Néerl. Sci. Exact. et Nat., Sér. III B, 1: 146-234. 1912.

. Hepserc, O. Pollen morphology in the Genus Polygonum L. s. lat. and its

taxonormical significance. Svensk. Bot. Tidsk. 40: 371-404. 1946.

. HEINTZELMAN, C. E. & R. A. Howarp. The comparative morphology of the

Icacinaceae. V. The pubescence and the crystals. Amer. Jour. Bot. 35:
42-52,

. Howarp, R. A. Studies of the Icacinaceae, I. Preliminary taxonomic notes.
940.

Jour. Arnold ag 21: 461-489.

——— udies of the Icacinaceae, II. Humirianthera, Leretia, Mappia,
and coe. valid genera of the Icacineae. Jour. Arnold Arb. 23:
55-78. 1942,

——— . Studies of the Icacinaceae, III. A revision of Emmotum.
Jour. Arnold Arb. 23: 479-494, 1942.

—— tudies of the Icacinaceae, IV. Consideration of the New
World en Contrib. Gray Herb. 142: 1-60. 1942

——— tudies of the Icacinaceae, V. A revision of the genus
Citronella D. Don. Contrib. Gray Herb. 142: 60-92. 1942.

. Studies of the Icacinaceae, VI. Jrvingbaileya and Codiocarpus,

two new eae of the Icacineae. Brittonia 5: 47-57. 1943.
——_ es of the Icacinaceaee VII. A revision of the Genus
Fare ee Seeman. Lloydia 6: 133-143. 1943.

284 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxxim

23. Howarp, R. A. Studies of the Icacinaceae. VIII. Brief notes of some Old
World Genera. Lloydia 6: 144-154. 1943

24. Hutcuinson, J. The families of flowering plants I. Dicotyledons. Mac-
millan, London. 1926

25. IversEN, J. & J. TroeLs-smitH, Pollenmorfolgiske definitioner og typer.
Danm. Geol. Unders. IV, Bd. 3, 8: 1-52.

26. Linpau, G. Acanthaceae. Nat. Pflanzenfam. IV, 3b: 280-287. 1895.

27. Maurirzon, J. Embryologische Angaben iiber Stackhousiaceae, Hippo-
crateaceae und Icacinaceae. Svensk. Bot. Tidskr. 30: 541-550. 1936.

28. Merritt, E. D. Records of Indo-Chinese Plants, III. Jour. Arnold Arb. 23:
176-178. 1942.

29. Recorp, S. J., et al. Glossary of terms used in describing woods. Tropical
Woods, No. 36, 1-12. 1933.

30. SteuMER, H. Icacinaceae. Nat. Pflanzenfam. 20b: 322-396. 1942.

31. WettstTeIN, R. Handbuch der systematischen Botanik. 4. Aufl. Leipsig
und Wien. 1935.

32. Wopenouse, R. P. Pollen grains. McGraw-Hill, New York. 1935.

DESCRIPTION OF PLATES

All figures represent camera lucida drawings of unstained, expanded pollen
grains mounted in lactic acid. The original drawings represent a magnification of
ca. 1800 X except in the cases of Fic. 10B (magnification of ca. 2200 x) and
Fics. 76, 76A, and 77 (magnification of ca. 1000 x). The illustrations are
reduced by approximately one-half in the present publication.

PLATE I

Fic. 1. Emmotum fagifolium (La Cruz 3375, G), polar view. Fic. 1A. ditto,
equatorial view. Fic. 2. E. nudum (Spruce 3541, G), polar view. Fic. 2A. ditto,
equatorial view. Fic. 3. E. argenteum (Tate 564, TYPE, NY), polar view. Fic.
3A. ditto, equatorial view. Fic. 4. E. glabrum (Klug 3990, G), polar view.
Fic. 4A. ditto, equatorial view. Fic. 5. E. nitens (Burchell 8233, G), polar
view. Fic. SA. ditto, equatorial view. Fic. 6. Ottoschulzia cubensis (Wright
2639, corypE, G), polar view. Fic. 6A. O. cubensis, equatorial view. Fic. 7.
Poraqueiba sericea (Ducke 25, A), polar view. Fic. 7A. ditto, equatorial view.
Fic. 8. Pittosporopsis Kerrit (Henry 11778, A), polar view. Fic. 8A. ditto,
equatorial view. Fic. 9. Oecopetalum mexicanum (Purpus 6159, G), polar view.
Fic. 9A. ditto, equatorial view.

PLATE II
Fic. 10. Cassinopsis ilicifolia (Burchell 5817, G), polar view. Fic. 10A.
ditto, equatorial view. Fic. 10B. ditto, equatorial view. Fic. 11. C. tinifolia
(Wood 9428, A), polar view. F 11A. ditto, equatorial view. Fic. 12.

Pennantia corymbosa (Kirk, G), polar view. Fic. 12A. ditto, oblique view.
Fic. 13, Pennantia Cunninghamii (Moore, G), polar view. Fic. 13A. ditto,
equatorial view. Fic. 14. Calatola laevigata (Schipp 8-708, F), polar view.
Fic. 14A. ditto, equatorial view. Fic. 15. Citronella Smythii (Michael 203, G),
polar view. Fic. 15A. ditto, equatorial view. Fic. 16. C. Gongonha (Hoehne 385,
A), polar view. Fic. 16A. ditto, equatorial view. Fic. 17. C. bahviggst
(Tonduz 11664, G), polar view. Fic. 17A. a“ a view. Fic.
mucronata (Munos & Coronel 1394, G), polar v

1952] DAHL, MORPHOLOGY OF THE ICACINACEAE 285

PLATE III

Fic. 19. C. iicifolia (Weberbauer 6617, F), polar view. Fic. 19A. ditto,
equatorial view. Fic. 20. C. philippinensis (Ramos 33267, A), polar view. Fic.
20A. ditto, equatorial view. Fic. 21. C. incarum (Weberbauer 7056, F), polar
view. Fic. 21A. ditto, equatorial view. Fic. 22. Dendrobangia boliviana (Rusby
1694, G), polar view. Fic. 22A. ditto, equatorial view. Fic. 23. Platea
philippinensis gous 9777, ISOTYPE, A), polar view. Fic, 23A. ditto, gaia
view. Fic. 24. Icacina Mannii (Zenker 475, G), equatorial view.
oe salutaris (Mildbraed 10536, A), polar view. Fic. 25A. ditto, ae
view

PLATE IV

Leretia cordata (Klug 2967, G), equatorial view. Fic. 27.
eee parviflora (Krukoff 6954, A), equatorial view. Fic. 28. P. flava
(Sandwith 590, isotype, NY), equatorial view. Fic. 29. Humirianthera rupestris
(Ducke 25288, US), polar view. Fic. 29A. ditto, equatorial view. Fic. 30.
H. rupestris (Ducke 342, US), polar view. Fic. 30A. ditto, equatorial view.
IG. 31. Natsiatum herpeticum (Parkinson 3927, A). Fic. 32. Leptaulus
daphnoides (Linder 872, A), oblique view. Fic. 33. L. grandifolius (Zenker 14,
G), polar view. Fic. 33A. ditto, equatorial view.

PLATE V

Fic. 34, Anisomallon clusiaefolium (Prony 1731, A), polar view. Fic. 34A.
ditto, equatorial view. Fic. 35. Apodytes Gardneriana (Silva 12, A), polar view.
Fic. 35A. ditto, equatorial view. Fic. 36. A. cambodiana (Clemens 3891, A),
polar view. Fic. 36A. ditto, equatorial view. Fic. 37. A. brachystylis (Kajew-
ski 1184, A), polar view. Fic. 37A. ditto, equatorial view. Fic. 37B. ditto,
polar view. Fic. 38. A. beninense (Elliot 4712, G), polar view. Fic, 33A. ditto,
equatorial view. Fic. 39. A. cuminensis (Eala 1930, F), polar view. Fic. 39A.
ditto, equatorial view. Fic. 40. Rhaphiostylis cordifolia (Linder 1218, A), polar
view.

PLATE VI

Fic. 41. Rhaphiostylis ferruginea (Zenker 65, C), polar view. Fic. 41A.
ditto, equatorial view. Fic. 42. R. latifolia (Zenker 2242, A), polar view. Fic.
42A. ditto, equatorial view. Fic. 43. Alsodeiopsis wean (Zenker F,
765310), polar view. Fic, 43A. ditto, equatorial view. Fic. 44. Gonocaryum
longeracemosum (Sargent, polar view. Fic. 44A. ditto, sane view.

PLATE VII
Fic. 45. Gonocaryum cognatum (Ramos and Edano 44073, A), Age view.
Fic. 45A. ditto, equatorial view. Fic. 46. G. calleryanum (Phil. B ur. Sci. 33,

A), polar view. Fic. 46A. ditto, equatorial view. Fic. 47. Codiocarpus Merrittii
(Ramos 40837, TOPOTYPE, A), oblique view. Fic. 47A. ditto, equatorial view.
1G. 48. Lasianthera africana (Conrau 41, A), polar view. Fic. 43A. ditto,
equatorial view. Fic. 49. Gastrolepis austro-caledonica (Franc 1791, G), polar
view. Fic. 49A. ditto, equatorial view. Fic. 50. Stemonurus cumingianus
(Wenzel 350, G), polar view. ae SOA. ditto, equatorial view. Fic. 51.
Medusanthera ary (Smith 578, G), polar view. Fic. 51A. ditto, equatorial
view. Fic. 52. M. papuana ae 17913, C), oblique view. Fic. 53. M.
laxiflora (Brass 2724, A), polar view. Fic. 53A. ditto, equatorial view. Fic. 54.

286 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxx
M. gilda Danses noabag! 3061, A), polar view. Fic. 54A. ditto, ei

view. Fic, M. glabra (Wenzel 1029, A), polar view. to,
equatorial view. a . M. — (Kanehira 1882, NY), polar view.
Fic. 56A. ditto, equatorial view. Fic. 57. Urandra ed hall (Haviland &
Hose 1544, G), polar view. Fic. 57A. an equatorial v
PLATE VIII
Fic. 58. Urandra apt (Toroes 4323, A), polar view. Fic. 58A. ditto,
equatorial view. Fic. 59. U. umbellata (Haviland 1782, G), polar view. Fic.

59A. ditto, nine view. Fic. 60. Irvingbaileya australis (Tardent 259, A),
polar view. Fic. 60A. ditto, equatorial view. Fic. 60B. ditto, polar view. Fic.
61. Cantleya corniculata (ex Herb. Hort. Bog. 265770, oe polar view. Fic. 61A.
ditto, equatorial view. Fic. 62. Discophora montana (Lawrance 535, G), polar
view. Fic. 62A. ditto, equatorial view. Fic. 63. Hosiea sinensis (Wilson 960a,
A), polar view. Fic. 64. Mappia mexicana (Pringle 6645, G), polar view. Fic.

64A. ditto, equatorial view. Fic. 65. /odes floribunda (Toroes 77, ISOTYPE, A).
Fic. 66. I. ovalis (Parker 2770, A). Fic. 67. I. liberica (Linder 1102, A).

PLATE IX

Fic. 68. Jodes philippinensis (Ramos 42567, A). Fic. 69. Mappianthus
iodioides (Henry 12063, A), polar view. Fic. 69A. ditto, equatorial view. Fic.
70. Polyporandra scandens (Kajewski 2206, A). Fic. 71. a e4
pinensis (Wenzel 987, A). Fic. 72. Polycephalium Poggei (Zenker 313, A).
Fic. 73. raga de capitata (Linder 1076, A). Fic. 73A. ditto, pai
apertures in optical section. Fic. 74. Miquelia Cumingii (Phil. Bur. Sci. 49997,

Fic. 75. Phytocrene Blancoi (Loher 13821, A). Fic. 76. Desmostachys
Preuss (Zenker 121, G), polar view. Fic. 76A. ditto, equatorial view. Fic.
. D. Vogel (Linder 1345, A), equatorial view.

Jour. ARNOLD Ars, VoL. XXXIII PLaTE I

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Jour. ARNOLD Ars. VoL. XXXIII

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Jour. ArRNoLD Ars. VoL. XXXIII

DAHL, POLLEN OF THE ICACINACEAE

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Jour. ArNoLp Ars. VoL. XXXIII PraTe VII

DAHL, POLLEN OF THE ICACINACEAE

Jour. ARNOLD Ars. VoL. XXXIII PiaTeE VIII

DAHL, POLLEN OF THE ICACINACEAE

PLATE IX

Jour. ARNoLD Ars. VoL. XXXIII

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DAHL,

296 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxir

ABERRANT SILVER MAPLES
Scott S$. PaAuLEY and ALBERT JOHNSON
With one plate

ABERRANT FORMS OF THE SILVER MAPLE, Acer saccharinum L., have
long been known in Europe and the United States. As early as 1893
Schwerin (1) described five varieties and 23 forms, including most, if not
all, of the forms known today. Most of these forms differ from the normal
species in shape or color of the leaf, but they include also columnar and
pendulous growth habits. Rehder (2) lists the varieties pendulum,
pyramidale, lutescens, tripartitum and laciniatum. The last of these
varieties includes the cultivated “Skinner” maple and “Wier” maple. The
Wier maple has dissected leaves, but the lobes are not abnormally slender,
while Skinner’s maple has leaves with extremely slender lobes. These
extreme forms of the silver maple have been described in the American
Nurseryman (3) with an even more extreme form from the Willis Nursery
Company of Ottawa, Kansas. The Skinner variety seems to have originated
independently in several widely separated areas.

Our experience with silver maple variants began in 1947 when we grafted
a very young, apparently normal, seedling on to a normal one-year old
seedling. The grafted seedling soon began to show what appeared to be a
mosaic infection, with abnormal leaf development and a mottled appear-
ance. Eventually a sucker developed from the rootstock which was normal
in every respect even at the end of two-year’s growth. Grafts between
branches of the scion into shoots from the rootstock showed no apparent
transmission of a virus to the normal stock.

In 1950 seed was collected from an isolated atypical silver maple at the
Arboretum’s Case Estate in Weston, Mass. This tree had abnormal mosaic
leaves, but was not nearly as aberrant as some of its seedlings.. From
approximately one hundred seedlings four distinct types of segregates were
obtained. In addition to the normal segregates there was one resembling
the variety /aciniata, two of which were similar to Wier’s maple, and nine
similar to Skinner’s variety. The latter two types not only had small
leaves with narrow lobes, but were also mottled and deficient in chlorophyll.
Photographs of the four different types are shown in PLATE 1.

The leaf aberrations in these segregates are correlated with growth habit
as is the case in the Wier and Skinner maples. The Jaciniata type of
segregate, with little or no evidence of abnormal chlorophyll development,
is perhaps more spreading in growth habit than the normal silver maple and
is not as vigorous. The Wier type has somewhat pendulous branches and
grows slowly, while the Skinner type of segregate is very spreading in
growth habit and is even less vigorous,

In 1950 and again in 1951 buds from several different normal silver
maples were budded on these mosaic segregates and in no case was there

1952] PAULEY & JOHNSON, ABERRANT SILVER MAPLES 297

any evidence of mosaic transmission. The budding in 1951 was done in
early August, and as soon as the bud had set, the branch above the bud
was cut back, to force the growth of the bud. Thus it was possible to get
several months growth of the normal branch. The buds inserted in 1950
made a complete season’s growth without showing any evidence of mosaic.

The symptoms of these abnormal silver maples certainly suggest virus
infection, but the failure to transmit the mosaic by either budding or
grafting, and the fact that at least two different mosaic types of segregates
were among the progeny of a single tree, throw some doubt on the virus
origin of these variants. In the case of prunus virus (3), however, the
virus transmission by grafting requires two full seasons of growth (4). It
is possible that these aberrant seedlings had a virus which was slow to
develop symptoms when transmitted to a normal host.

Blakeslee (5) has found a mutation in Datura which resembles the effect
of a virus disease. Both the virus infected and the mutant type showed
modified leaves, capsules and flowers. In view of the failure to transmit
the mosaic by budding or grafting, and the segregation of four distinct
types of progeny from a single parental tree, it appears that the aberrant
silver maples described are of genetic origin.

Recently John and Wilson (6) have described a somewhat similar
genetically conditioned leaf character in the cucumber, Cucumis sativus L.
They found what was termed a “ginkgo leaf” variation to be inherited as
a simple Mendelian recessive factor. No variation in degree of expression
such as observed in the Acer mutants was reported.

LITERATURE CITED

1. ScHWERIN, F. G. Die Varietaten der Gattung Acer. Paul Parey, Berlin. 1893.
2. ReHper, A. Manuel of cultivated trees and shrubs. Macmillan Co. New
York,
3. ANONYMOUS. Weeping Silver Maple. American Nurseryman, 68(8):
1938.
4. Smirn, K. Plant Virus Diseases. J. & A. Churchill, Ltd. London. 1937.
5. BLAKESLEE, A. F. Growth patterns in plants. Growth Symposium. 1941.
6. Joun, C. A. & Witson, J. D. A “gingko leafed” mutation in the cucumber.
Jour. Hered., 43: 47-48. 1952.
DESCRIPTION OF PLATE
Typical leaves of silver maple ae from a
single parental tre
1. Leaf of extreme variant similar to those of a variant described by the Willis
Nursery Co. The leaves have extremely narrow lobes and have chlorophyll
deficiencies characteristic 2 a virus infection.
2. Leaf of a normal segregat
3. Leaf of a segregate with ee leaves similar to those of “Wierii.” These
leaves show little or no mosaic characteristics.
4. Leaf of a segregate resembling ‘Skinners’ silver maple. Chlorophyll aberra-

tions resemble those caused by mosaic virus.

PLATE I

Jour. ARNOLD Ars. VoL. XXXIII

N, ABERRANT SILVER MAPLES

O

$

PAULEY AND JOHN

JOURNAL

OF THE

ARNOLD ARBORETUM

VoL. XXXITII OCTOBER 1952 NUMBER 4

STUDIES IN THE BORAGINACEAE, XXIII
A SURVEY OF THE GENUS LITHOSPERMUM

IvAN M. JOHNSTON
With three plates

SoME MONTHS AGO I began a study of the genus Arnedia for the purpose
of determining whether or not the eastern Asiatic species A. Hancockiana
(Oliver) Johnston had been properly assigned to that genus. As the study
progressed questions arose as to precisely how Arnebia could be dis-
tinguished from Lithospermum. As a basis for the solution of this problem
dissections were made of representative species of Lithospermum. This
gave me reasons for doubting the naturalness of that genus as currently
defined. Methodical study of all the species of Lithospermum was then
commenced, and the investigation eventually extended to members of other
obviously related genera. From an examination of a few Asiatic species
my study has gradually widened into a critical re-examination and
redefinition of Lithospermum and all the other genera of the Lithospermeae
characterized by coarsely bracted inflorescence.

The present paper is concerned only with Lithospermum. The species
of the genus are enumerated, their principal synonymy given, and a key for
their identification provided. My chief concern, however, has not been with
the details of species-classification, but rather with the over-all composition
of the genus, and in its units only so far as they reveal morphological
traits pertinent to the definition of the genus and the major grouping of its
species. The observations given concerning the species are primarily those
significant in establishing the relationship and the precise position of the
species in a natural classification. They are mostly concerned with the
inner structures of the corolla, for the most part described only imperfectly,
if not completely ignored, by previous writers. The related genera will be
given similar treatment in the next paper of this series. Their relationships
with Lithospermum will there be discussed at length, and new generic
descriptions, keys, and tabular synopses useful in evaluating and distin-
guishing them will be provided.

As here treated Lithospermum includes Arnebia and Macrctomia, but
excludes Lithodora, Moltkia and Stenosolenium, as well as Lithospermum

300 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxxm1

apulum L., L. miscrospermum Boiss., L. cinerascens (DC.) Johnston, L.
revolutum Robins., and the species allied to L. arvense L. and L. purpureo-
coeruleum L. The genus thus delimited is a very natural one, readily
distinguished from its closest relatives by decisive characters of fundamental
importance. Its species are many and diverse, and, though prevailingly
well marked, exhibit in all degrees of expression and in many combinations
the various evolutionary modifications that distinguish them. As a result of
a wealth of intermediates, and the freedom with which characters are
combined, there are no clear-cut large groupings of the species within the
genus. The genus has great internal coherence. It is exasperatingly lacking
in lines of cleavage. There are no sharply defined natural major divisions.
Even artificial divisions useful in a key are difficult to find. The genus
contains so many well-marked species that even the naming of species-
groups (series) seems impractical, since the majority of them would be
monotypic. Accordingly I have proposed no subgeneric divisions in
Lithospermum. This is not because I have made no attempt to discern
them, but only because I have been forced to the belief, reluctantly, that no
useful, natural, definable ones are present.

The latest comprehensive study of Lithospermum and Arnebia is that
published in 1846 in the 10th volume of De Candolle’s Prodromus. Sub-
sequent publications concerning the genera have been mostly regional,
most of them relating to only a limited number of species. Four papers,
however, have sufficiently broad scope to be especially useful to the student
of Lithospermum. One by Helene Spengler, Oesterr. Bot. Zeitschr. 68:
111-123, f. 1-41 (1919), is a study of the form and internal organization
of the corollas of Lithospermum, Many species were examined and
described, and the dissected corollas (and frequently also the nutlets)
were illustrated. While her observations do not always conform with my
findings, her paper is a stimulating and useful one. The many American
Lithospermums have been treated by Johnston, Contr. Gray Herb. 70: 18—
31 (1924) and 78: 6-11 (1927). The species are keyed and their synon-
ymy given. Incidental to the publication of additional species, another key
to the American species was later published by A. Brand, Fedde Repert,
28: 10-17 (1930). The synonymy of the American species has been rather
completely covered by Johnston and by Brand. Synonymy for the species
of Eurasia and Africa is to be found in the purely bibliographic paper by
G. Stroh, Beih. Bot. Centralbl. 588: 203-211 (1938).

The present study is based primarily on the material in the large and
very representative world-wide collection of the Boraginaceae assembled at
the Gray Herbarium by my efforts during the past twenty-five years.
From that source suites of specimens representative of practically all species
have been available for dissection and repeated examination and com-
parison. Supplementary material of great usefulness has also been received
on loan from other institutions: Mexican and African collections from
the Missouri Botanical Garden, Mexican and South American from the
Chicago Museum of Natural History and from the United States National
Museum; and Asiatic (“Arnebia”’) from the Natural History Museum,

1952] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXIII 301

Stockholm, and from the Botanical Museum of the University of Lund.
The type of Arnebiola was also sent for my examination from the Botanical
Institute, Florence. Through the courtesy of the curators of the institutions
mentioned I have had a very large and representative series of specimens
for study. For this help I wish to express again, now publicly, my grateful
appreciation.

GENERAL OBSERVATIONS ON THE FLOWER AND FRUIT

Most species of Lithospermum have yellow or orange, or at least
yellowish corollas. Albino variants of these, if they exist, are apparently
extremely rare. Pure white corollas, entirely so or yellow only in the throat,
are characteristic of about seventeen species. Blue or pink or brownish
corollas are characteristic of a few other species, mostly Asiatic. In general,
closely related species: agree in corolla-color. Amonz the Asiatic species
formerly placed in Arnebia a number (L. decumbens, L. Griffithii, L.
Szechenyi, L. guttatum and L. Tournefortii) have five black or blue spots
on the yellow or orange corolla-limb. These decorative spots, one adjacent
to the base of each sinus on the limb, are evanescent, being dark and con-
spicuous on the newly expanded corolla but soon fading after prolonged
exposure to sunlight. They occur on the corollas of species sharing general
relationships but no immediate affinity.

The corolla-lobes usually have entire margins. Erose or lobulate or
lacerate margins, however, are characteristic of the Asiatic L. fimbriatum,
L. fimbriopetalum, and L. Bungei, all former members of Arnebia, and
also of the American species L. incisum and L. calycosum.

The throat of the corolla may bear five well-developed invaginate
appendages, or these may be imperfectly developed or entirely lacking.
The faucal invaginations may be trapeziform, gibbose, lunate, or mere
convexities. They may be velvety, velvety with intermixed stipitate glands,
or merely glanduliferous. When present their location is marked on the
outside of the corolla by slits that open into pocket-like recesses or merely
by convex depressions of varying depth. Minute stipitate glands are
common features in and about the throat of the corolla. They may occur
on the faucal appendages, when these are present, but are usually most
abundant on the inner side of the appendages and especially just below
their base. Frequently they are abundant on the walls of the corolla-tube
as far down as the filament-attachments. In many species they decorate
the decurrent base of the filaments. In some of the species lacking faucal
appendages (L. discolor, L. viride, L. californicum) the glands are abundant
and generally distributed in the throat, and in others (L. multiflorum, L.
obovatum, and L. calcicola) they may form sharply circumscribed con-
eregations that take the place of the appendages. The species which have
the corolla-throat unappendaged and sparingly or not at all glanduliferous
are mostly Asiatic. Most of them were formerly placed in Arnebia. Indeed,
lack of such appendages and glands has been used as the crucial character
in attempts to distinguish Arnebia from Lithospermum. It is to be noted,

302 JOURNAL OF THE ARNOLD ARBORETUM _ [VvoL. XxxiIr

however, that among these Asiatic species glands are not completely absent
in the corolla-throat. In species such as L. Hancockianum, L. Sewerzowi,
L. Szechenyi, L. guttatum, L. densiflorum, and L. Griffithii the glands may
occur only in very moderate quantity or be very sparse, but they are present
and are evident when looked for.

The inner wall of the corolla-tube is glabrous in most species or at most
has only very minute hairs on the basal nectary. The tube is distinctly
hairy, however, in L. hispidissimum, L. Tournefortii, and L. discolor, as
well as in the three related species L. cobrense, L. obovatum, and L.
tubuliflorum. The development has diagnostic value but little importance
as an indicator of relationship. The species with hairy corolla-tubes are all
heterostylic. In some the hairs are more abundant in the short-styled
flowers than in the long-styled ones. An extreme condition occurs in
L. obovatum, in which only the long-styled flowers have hairs, the tube of
the short-styled corollas being glabrous.

Another feature of the corolla of Lithospermum is the nectary located
inside the tube 1—2 mm. above the base. This appears in a variety of forms
and has various degrees of development. In some species the nectary
appears to be undifferentiated (L. Tournefortii, L. Benthami, L. euchromon,
L. tetrastigma) or marked only by minute hairs (L. densiflorum, L.
incisum). In other species it is hardly more than a tumid band (ZL.
californicum, L. ruderale) or a lineate ridge (L. fimbriatum). Frequently
it becomes a prominent annulate ridge or flange (L. officinale) or even
collar-like and as much as 1 mm. high (L. guttatum, L. Aucherti). Not
uncommonly it is more or less lobulate. In some species (L. multiflorum,
L. cobrense, L. caroliniense) it is divided into ten proximate quadrate
lobes. Related species tend to have similar nectaries.

The anthers are oblong or narrowly oblong with obtuse or rarely some-
what acute or even apiculate tips. In non-heterostylic flowers they are
borne near the summit of the corolla-tube or below it at a distance not
exceeding their own length. When the tube is short, as in L. officinale,
they may be attached near the middle of the tube, but in flowers with
elongate tubes, such as are present in most species, they are always borne
in the upper quarter of the tube. In corollas with faucal appendages the
tip of the anther commonly reaches up to the appendage-base and is never
more than an anther-length below it. This is the normal condition in the
genus. Most departures from it are associated with heterostyly. In long-
styled flowers the anthers are usually borne at or near the middle of the
tube even when the latter is very elongate. In L. obovatum, L. cobrense,
and L. tubuliflorum they are extremely low, being borne on the lower third
of the tube. In short-styled flowers the anthers are found very high in the
tube and frequently partially exserted from it. In this genus any corolla
bearing anthers near the middle of its tube almost certainly belongs to a
long-styled flower of a heterostylic species.

The filaments are usually short, less than one half and usually only a
third or a quarter of the length of the anther. Two species show notable
departure from this norm. The stamens on the short-styled flower of L.

1952] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXIII 303

hispidissimum have filaments nearly as long as the anther. In the long-
styled flowers, however, the relation of filament to anther is normal. One
of the most ahectant Filathien ts, however, is that present in some of the
plants I have referred to L. tschimganicum. This will be discussed under
that species.

The attachment of the stamens in most species is whorled, i.e., within
the individual corolla all are attached at the same level on the corolla-tube.
In L. Tournefortii, however, although always confined within definite
zones on the tube, the stamens are affixed at several differing levels. In
L. decumbens, western forms have whorled stamens or may have one
stamen borne slightly below the other four. In eastern forms, however,
the stamens within a corolla are frequently borne at several loosely spaced
superimposed levels, with two anthers at the mouth of the tube, another
pair below, and a single anther lower down. In both ZL. Tournefortii and
L. decumbens, accordingly, there is a departure from radial symmetry in
the androecium. Such departures are present in a number of genera closely
related to Lithospermum. Stamens attached at unequal heights in the
tube are developed by species of Stenosolenium, Lithodora, and Alkanna.
In Moltkia the filaments become unequal in length. Zygomorphy associated
with stamens of various length and heights of attachment occurs in Echium
and Halacsya. The sporadic appearance of aberrant stamen-attachments
in Lithospermum is not surprising.

The behavior of the style in the non-heterostylic species deserves some
comment, since its length relative to that of the corolla-tube may change
between the time pollen is shed in the flower-bud just before opening and
the time when the flower is completely expanded. In mature flower-buds
the style is usually sufficiently long to carry the stigmas up between the
dehiscent anthers or just above them. In species such as L. officinale, L.
distichum, and L. ruderale the relative position of stigma and anthers
remains similar in the expanded corolla. In some species there is evident
elongation of the corolla-tube subsequent to anther-dehiscence. When this
is accompanied by comparable elongation of the style, as in L. calycosum,
the relative position of anthers and stigmas is maintained. In L. Nelsonii
and L. strictum, however, the style elongates less, and although the stigmas
are between the anthers at dehiscence, they assume a position below them
when the corolla is completely expanded. In some of the Mexican species
(L. oblongifolium, L. viride, L. Muelleri, and forms of L. calcicola) the
style has become very elongate in the unopened bud. At the time the
anthers dehisce the stigmas are held well above the anthers and the stvle
itself is somewhat contorted in order to accommodate its length in its
cramped quarters. When the corolla eventually opens the style straightens
and the stigmas become exserted from the corolla-tube. Were the corolla-
lobes of these species not strongly imbricate, but valvate as in Onosmodium,
the style could force a way out through the tip of the unopened corolla
and be precociously exserted in the manner characteristic of that related
genus. A very different condition exists among the annual species of the
Old World. In some plants of L. detonsum the stigmas and anthers are

304 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxxmI

juxtaposed at the time pollen is shed, but by the time the corolla is fully
expanded the stamens are carried high above the stigmas, The corolla-tube
has apparently elongated more rapidly than the style. In L. decumbens
and other plants of LZ. detonsum, even when pollen is shed, the stigmas are
well below the anthers. Interestingly, they become even more widely
separated by the time the corolla is fully expanded.

The style usually bears two stigmas; only in L. tetrastigma and L.
decumbens is the number increased to four. Among some of the Asiatic
species, particularly the annual ones, the style is apically forked and the
stigmas terminate the short branches. In most species, however, the stigmas
are sessile on the upper end of a simple style. Though usually terminal, in
some American species they are distinctly subterminal, being affixed
laterally slightly below a convex or conic, and sometimes notched sterile
tip of the style. The latter condition is usually well developed in L. Nelsonit,
L. mediale, and L. oblongifolium. The stigmas are usually distinct, but in
such species as L. strictum, L. Muelleri, L. fimbriatum and L. Szechenyt
they are crowded together on the tip of the style and their juxtaposed bases
may become somewhat joined. Most stigmas are more or less globose, but
in species such as L. guttatum and L. multiflorum they may become
flattened and broadened and even obscurely lobed. Two species have the
style more or less evidently twice forked and frequently bearing a stigma
terminating each of the four ultimate branches. This remarkable condition,
unique among the Boraginoideae, no doubt arose by division of stigmas
borne on.a simply forked style. Indeed, a suggestion of the condition is
not infrequent in L. guttatum. In some individuals of that species the two
stigmas may be very deeply bilobed and even nearly divided. With such
evidence that an incipient stage actually exists, the further evolutionary
steps for the formation of a bis-bifid style bearing four stigmas is not
difficult to imagine. In any case the two species with this extreme develop-
ment give indications in all other structures that they are not immediately
related. The peculiarities they share are probably parallelisms and not the
direct product of shared immediate ancestry. In this regard it is to be noted
that the stigmas of L. tetrastigma are narrow and elongate, a form unique
in this genus

Heterostylic flowers are developed by at least eighteen species of
Lithospermum. The two types of flowers in these species differ not only
in position of stamens and length of style, but also in the size and frequently
even in the shape of the pollen grains. Among most of these species the
anthers in the long-styled flowers are borne near the middle of the corolla-
tube. In L. obovatum and L. tubuliflorum, however, the anthers are in the
lower third of the tube, while in L. fimbriatum, L. densiflorum, and L.
hispidissimum they are borne in its upper third. The style in such flowers
usually reaches the summit of the corolla-tube and may be even slightly
exserted from it. In L. densiflorum long styles may be exserted as much
as 5 mm. The anthers of short-stvled flowers are borne near the summit
of the tube, either entirely included or partially exserted from it. Their
style reaches the middle of the tube or slightly beyond. Very short styles,

1952] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXIII 305

less than a quarter of the length of the tube, are present only in L. discolor
and L. canescens.

Associated with the major differences relating to style-length and stamen-
height, there are also the correlated minor ones usually present in well-
developed heterostyly. The anthers of long-styled flowers are usually
smaller than those of short-styled flowers. In some species, e.g., L.
densiflorum, the corollas with short styles are perceptibly larger than
those with long styles. In most of the species the corolla-tube differs in
form also. It is usually cylindric or very gradually ampliate in short-
styled flowers. In the long-styled flowers there is usually a slight but
abrupt increase in diameter at above the level of the stamen-attachments,
with the result that the upper half of the tube is differentiated into an
elongate cylindric throat. The corollas of L. obovatum present an unusual
difference, perhaps unique among heterostylic flowers. The inner surface
of the tube is hairy in the long-styled flowers but glabrous in those with
short styles. The manifestations of heterostyly in L. Tournefortii are also
unusual and perhaps unique. In that species the stamens in the individual
corolla are not borne at one level, whorled, as is usual in most Lithosper-
mums and other regular flowers. Rather, they are attached at several
different heights on the corolla, and so occupy a zone rather than a single
level on the tube. These staminiferous zones behave as do simple whorls
of stamens. They have very different positions in the two types of flowers,
being located below the middle of the tube in long-styled corollas and near
its summit in the short-styled.

The pollen grains of the eighteen species with well-marked heterostyly
have size-differences correlated with the two types of flower, those of the
short-styled flowers always being the larger (Plates I and IT). In nine of
these species (L. Griffithii, L. Tournefortii, L. densiflorum, L. discolor, L.
canescens, L. multiflorum, L. cobrense, L. obovatum, and L. californicum)
the grains in the two floral types, though obviously different in size, are
otherwise very similar. This is normal and is the condition present in all
genera in which marked heterostyly has been studied. The pollen of the
remaining nine heterostylic species is not of this conventional sort. The
grains in the two types of flower differ not only in size but in shape as
well. This is a most unusual condition. Indeed, it seems to be a type of
pollen dimorphism previously unreported. The flowers of those species of
Lithospermum in which it occurs present heterostyly in one of its most
highly elaborated states. Two of the nine species having dimorphic
pollen are American (L. tubuliflorum and L. caroliniense) and seven are
Asiatic (L. fimbriatum, L. Sewerzowi, L. Szechenyi, L. guttatum, L.
Benthami, L. euchromon, and L. hispidissimum). If classed according to
the intimacy of their relationship, these nine species fall into eight groups
as follows: 1. L. tubuliflorum,; 2. L. caroliniense; 3. L. fimbriatum: 4. L.
Sewerzowi; 5. L. Szechenyi; 6. L. guttatum: 7. L. Benthami and L.
euchromon, and 8. L. hispidissimum. Most of these have their closest
affinities not with each other but with species or species-groups having
pollen little or not at all differentiated in shape. Furthermore, their closest

306 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxx

relations are intra-continental, the American species having a basically
different type of pollen than that of the Asiatic species. Heterostylic
species with dimorphic pollen do not form a natural group.

With heterostyly so strongly developed in some species, it is surprising
that the other members of the genus show so few tendencies of that nature.
Some evidence of incipient heterostyly occurs among some of the many
puzzling forms of L. decumbens. Throughout the wide range of that species
plants within a collection may or may not show differences in style-length,
and moderate differences in the distance below the corolla-lobes at which
the stamens are attached. In some collections of the large-flowered forms
of the species from Central Asia, the differences can be more marked and
even involve the shape of the corolla-tube. In all these forms, however,
pollen shows no variation in size or shape. If tendencies towards
heterostyly do exist, they have only weak and uncertain expression.

Within the Boraginaceae heterostyly is known in Lithodora, Pulmonaria,
Anchusa, Symphytum, Amsinckia, Cryptantha, Oreocarya, and Paracaryum.,
Of these genera only Lithodora is closely related to Lithospermum. Most
of its species have dimorphic flowers differing in style-length and height
of stamen-attachment, and in some, even in the form of the corolla-tube.
Unlike Lithospermum, however, the two types of flowers in Lithodora show
no differences in pollen.

Cleistogamy is present in Lithospermum incisum, L. Parksii, L. confine,
and L. mirabile, all members of a closely intrarelaind group, snd also in L.
calycosum. In a incisum the plant first produces clusters of large, very
attractive chasmogamic flowers, mostly infertile, and subsequently only
large numbers of cleistogamic flowers which fruit prolifically. Chasmogamic
flowers may be completely suppressed in LZ. confine. In the other species the
cymes produce conspicuous chasmogamic flowers until late in the growing
season, and then only minute cleistogamic ones. Both types of flowers appear
to be equally fertile. In all of the five species mentioned the cleistogamic
flowers are very similar. The corolla is calyptrate, usually 1-3 mm. long,
and eventually falls without opening. Faucal appendages and _stiped
glands that may be present in the large open flowers are not developed
in the corollas of the cleistogamic flowers. The pollen produced by open
and closed flowers is indistinguishable, or at most very slightly smaller in
the closed flowers. Since the style of the closed flowers is extremely short,
the fruit produced by each of the two types of flowers is readily distin-
guished by the length of the persisting style associated with it.

It is surprising that cleistogamy is not a more common development in
Lithospermum. Anyone seeking pollen in this genus for examination soon
discovers that the anthers of expanded flowers are nearly always empty.
Although the books report that Lithospermum is proterogynous, without
exception I have found that normal anthers in this genus shed their pollen
before the corolla opens. The pollen is dumped out and adheres in masses
on the inner sides of the faucal appendages and on the glanduliferous wall
of the throat. In some species masses of pollen adhere to the stigma, in the
bud frequently surrounded by the anthers, and these masses are later

1952] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXIII 307

carried up into the mouth of the corolla by stylar elongation. The physical
conditions are extremely favorable for self-pollinization. Indeed, only if
the flowers are self-sterile can it be prevented. Investigation of heterostyly
in many other genera has indicated that the condition is usually accom-
panied by a high degree of self-sterility. Possibly this is also present
in most of the non-heterostylic species of Lithospermum.

The nutlets of most species of Lithospermum are ellipsoidal or ovoid and
usually have a smooth lustrous white surface. There are, however, many
deviations from this common type. The surface may be smooth and
porcelain-like or more or less pitted, or, particularly in Asiatic species,
tuberculate, verrucose, or rugose. It may be white or stained with yellow-
brown or brown, or, in Asiatic species, gray, brownish, or even olivaceous,
Occasionally it is rubiginous (L. tetrastigma) or minutely mottled with
purplish (L. Tournefortii). The prevailing shapes are ovoid or ovoid-
ellipsoidal with rounded dorsum, rounded or obtusely angled venter, and
blunted obtusish apex. Sharply pointed, subrostrate nutlets, however,
occur in L, euchromon and L. Sewerzowi. In L. tetrastigma the nutlets are
compressed, having a broad, flat or slightly concave back and convex
venter. In L. fimbriatum and L. detonsum the broad nutlets have a wide
depression down the middle of the lower half of the dorsum.

American species frequently have a well-developed ventral keel. This
is commonly low and rounded, but in L. strictum it is narrow and acute.
It may extend even over the apex of the nutlet and onto the dorsum.
Among Mexican species the ridge down the middle of the venter is fre-
quently made more prominent by a line of pits or slit-like depressions in
the pericarp on either side of it. In most American species the ventral angle
of the nutlet is traversed from top to bottom by a fine lineate sulcus
representing the ventral suture. Among the Asiatic species other than

Tournefortii and L. tschamganicum there is little or no evidence of
this suture. The venter of their nutlets may be angled but is never
obviously keeled, nor does it bear lines of pits and slits. In most species
the nutlets are gradually narrowed to the base. In some, however, as the
result of a lineate suprabasal constriction, the nutlet may develop a collar-
like base (L. matamorense, L. incisum) and in others, because of a much
broader suprabasal constriction (L. ruderale, L. viride) it may have a short
stout neck just above a flaring base.

The nutlets have a basal attachment surface, commonly rounded and
about as broad as long. It is usually flat or slightly concave or convex. In
L. incisum, however, it is very deeply concave, almost excavated, and
furthermore bears a spur-like projection. In L. guttatum the attachment
has a broad lateral prolongation extending a short but evident distance up
the venter of the nutlet-body and then outward under the truncate base
of the ventral keel. A comparable upward prolongation of the attachment
surface occurs also on the very aberrant nutlets of L. tetrastigma. The
dorso-ventrally compressed nutlets of that species have a cordate base. The
attachment is transversely elongate and depressed at the center and has a
quadrate lobe extending upwards on the ventral side of the nutlet-body.

308 JOURNAL OF THE ARNOLD ARBORETUM _[vot. xxxim

The gynobase in Lithospermum, traditionally described as flat, is, as a
matter of fact, usually depressed pyramidal. When all four nutlets have
been matured and shed, the gynobase usually bears four plane or some-
what concave and upcurved surfaces that slope towards the base of the
style at angles of ten to forty-five degrees. In those species with nutlets
that have a ventral upward extension of their attachment-scar (L.
guttatum, L. tetrastigma, and to a less extent L. Sewerzowi) the gynobase
may even take the form of a strongly truncated pyramid. Among most
American species the surfaces on the gynobase (the scars or pads left by
the fallen nutlets) usually have an upturned or thickened cartilaginous
margin and tend to be joined laterally by cartilaginous tissue. In most
Asiatic species, however, as well as in a very few American (L. Nelsonit, L.
Berlandieri) the attachment pads on the gynobase are unmargined and,
though juxtaposed, remain distinct. In L. oblongifolium and L. strictum,
in which the pyramidal gynobase is about as high as broad, the cartilaginous
tissue between the margined attachment-pads becomes very prominent and
even continues upward to the base of the style as narrow wings along the
angles of the gynobasic pyramid.

POLLEN

In this genus I first examined pollen for the purpose of determining if
there were size-differences associated with heterostyly. When the surpris-
ing variety of forms was revealed, the pollen of Lithospermum and related
genera was methodically examined. It proves to be an extremely good indi-
cator of relationships, generic as well as specific, and hence of great use-
fulness in classification, particularly so in Lithospermum. Since my pur-
pose has been to examine pollen from as many herbarium specimens as
possible (about five hundred were sampled in Lithospermum) my technique
has been a simple one. Dehiscent anthers with pollen, taken from the
flower-bud just before the expansion of the corolla when possible, have been
macerated on a slide in a drop of 85% lactic acid and then protected by
a coverglass. The grains-expand rapidly and reach permanent form in a
minute or two. Labeled, unsealed mounts have been used repeatedly over
a period of six months. Those which dried out have been quickly repaired
by the addition of more lactic acid. Comparison of such old mounts with
freshly made ones shows no difference in pollen size or form. The grains
were studied and measured under 450 magnification. It is obvious that a
refined technique, and especially the use of stains and higher magnifications,
would reveal much more concerning the pollen than the gross features, the
size and form, and the position and number of pores which I have observed.
That, however, is left to a better microscopist than I. In the present
study of Lithosbermum a wide survey of pollen-types is more useful than
one that is limited because it is detailed.

The pollen of Lithospermum is notable in several respects. In the
heterostylic species the grains of long-styled and short-styled flowers differ
not only in size but frequently in shape as well. Differences in size are

1952] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXIII = 309

not unexpected. They have been found associated with heterostyly in

as the ultimate proof that true heterostyly was present. Differences in
shape, however, are a different matter. I have, in fact, been unable to
discover any previous report of such a condition: This pollen dimorphy
which is associated with the heterostyly in some species of Lithospermum
may be unique

The two kinds of pollen produced by the various heterostylic species of
Lithospermum are illustrated in Plates I and II. It is to be seen that size
differences occur in all the species. Dimorphy is extreme in L. tubuliflorum
(fig. 10) and L. caroliniense (fig. 12), moderate in L. Hookeri and L.
euchromon (fig. 3), L. fimbriatum (fig. 4), L. guttatum (fig. 5) and L.
hispidissimum (fig. 7), and only weak or variable in L. californicum (fig.
9), L. cobrense (fig. 15), and L. canescens (fig. 11).

The pollen of five species (L. incisum, L. Parksii, L. confine, L. mirabile,
and L. Tournefortii) is almost nerfectly spherical. In a few others it is
subglobose with the sides slightly angulate when viewed in lateral profile
(L. viride, L. Macbridei, L. strictum, L. Muelleri). In most species,
however, it is distinctly elongate with the length twenty-five to two hundred
per cent greater than the maximum thickness. In polar view and in trans-
verse sections perpendicular to the axis, the grains have a circular outline
or sometimes a slightly polygonal one if prominent pores are included. In
lateral view the profile varies greatly. The sides may be convex or out-
wardly angled, concave or inwardly angled, or straight and parallel or
somewhat convergent. What is most unusual, the outline of the upper
half of the grain may differ greatly from that of the lower half. As a
result, the grains have a wide variety of forms. They may be spherical,
ellipsoidal, barrel-shaped (cylindric with rounded ends), ovoid in various
modifications, or, as a result of a medial or submedial constriction,
achieve a lateral outline suggestive of an hourglass or a shoe-print in the
snow. In size they also vary widely, from 65—72 » in L. densiflorum down
to 13-16 pin L. officinale. As observed by me under 450 magnification, the
grains appear smooth; I have detected no furrows. The pores are small and
usually inconspicuous or even invisible. Only in a few species are they
noticeably protrudent in lactic acid (L. Pringlei, L. indecorum). Their
position and number is usually most readily determined by a study of
shrunken or collapsed grains.

Two fundamentally different types of pollen, best distinguished by the
position of the pores, are recognizable in the genus. One is represented by
fifteen of the species formerly referred to Arnebia and Macrotomia; the
other by the remaining twentv-nine snecies here referred to Lithospermum.
In the latter type the pores are in a single row, equally spaced around the
grain. Their number may be six, seven, eight, or nine, the particular num-
ber being relatively constant in the pollen of a given plant, but usually
varying over two or three numbers among individuals of the species.
When the grain of this type is spheric, ellipsoidal, or barrel-shaped, the
pores are on the equator or at least equidistant from the two poles. There

310 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxur

is, however, in this type of pollen, a tendency for the upper and lower
halves of the grain to differ in size and form. When such asymmetry is
present the band of pores may occur at levels below the middle of the grain.
In the subcylindric grains of LZ. calcicola the pores are only slightly below
the middle. They are more so in the somewhat ovoid grains of L.
tuberosum and very conspicuously so in grains having the hourglass or
shoe-print profiles. In such medially constricted grains the pores are
located where the lower half of the grain is broadest, or, in other words,
at the point where the evenly rounded curve of the base of the grain
terminates and constriction begins. When these grains are viewed in lateral
profile, the constriction just above the pores is by sloping, straight (not
curved) lines. The convergent straight lines of this constriction form the
distinctive shoulders recognizable in many grains of this type. As land-
marks, even in grains of the hourglass form, these sloping shoulders dis-
tinguish the lower from the upper half of the grain and are always an
indicator as to the location of the single band of pores.

The other type of pollen in Lithospermum, that which is characteristic
of most species formerly placed in Arnebia and Macrotomia, is always
perfectly symmetric, with the bottom and top halves alike. It may be
cylindric or nearly so, or weakly to strongly constricted medially and hence
sometimes subcylindric with slightly concave sides or more or less clearly
of the hourglass form. The pores are very indistinct. They appear to be
arranged about the convex ends of the grain, five (or rarely four) in the
row at each end. Possibly there may be additional pores, one at each pole.
In this type of pollen there are no pores on or near the equator. In three
species, L. densiflorum, L. detonsum (fig. 29) and L. Aucherii, the grains
sometimes appear to be serrulate at the constricted equator. When empty,
or dry and shriveled, they have an unusual form, being somewhat basket-
like and consisting of superimposed rings joined by five (or four) equally
spaced ribs. The ribs are usually verrucose at the middle and presumably
form the serrulations previously mentioned. The empty or shrunken
grains of the other species in this type are not ribbed nor basket-like.
Commonly they appear subcyclindric or bag-like. If they burst open, it is
usually at one or more places at the ends, never at or near the middle.
The pollen-walls are evidently thicker and more rigid in these grains than
in the other type of pollen.

The developmental history of the asymmetric pollen grains deserves
study. Prepared sections of the anther at various stages of development
should reveal exactly how the unusual grain-shapes arise and how, for
example, grains with the hourglass form can be efficiently packed together
in the theca. Until the matter is investigated we can only suspect that the
peculiarities in the form of the grains must be a manifestation occurring
late in their ontogeny.

Of the two types of pollen in the genus, the more common one, that with
six to nine pores in a single row, shows the greatest diversity in outline.
The many forms assumed by this pollen type can be homologized, however,
as modifications of a spheric grain with equatorial pores. From a spheric

1952] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXIII 311

grain, by equal enlargement of both hemispheres, ellipsoidal and barrel-
shaped grains can be derived. The asymmetrical grains, those bearing the
row of pores below the middle, are probably the result of hypertrophy of
one hemisphere. In the very asymmetric pollen with the hourglass or
foot-print profile, in which the row of pores is near one end, one half of
the original basic spherical form is to be recognized in the rounded base of
the grain. The major portion of the grain, that above the band of pores,
is the other hemisphere excessively enlarged and greatly modified in outline

7 1Ongess
000008

TEXT-FIG. 1. Relation of the forms in the two types of pollen in Lithosbermum.
Levels at which rows of pores are borne indicated by arrows and broken lines.
Type with two rows of pores (upper figures) always symmetrical, differing in
the degree of medial constriction. Type with one row of pores (lower figures),
probably all modifications of a sphere, becoming ellipsoidal or subcylindric by
equal modifications of the two hemispheres or becoming ovoid or achieving more
asymmetric form by excessive modification of only one hemisphere

What relations this type of pollen may have with that which has the pores
in two rows is obscure. The two different types remain distinct and their
variants give no clues as to how one of the types might have given rise to
the other or how they might both be derived from a common ancestral
form.

THE RELATIONS OF LITHOSPERMUM AND ARNEBIA

In a subsequent paper I will discuss the nature and relations between
Lithospermum and the numerous allied genera and give reasons why certain
species previously classified under Lithospermum should be excluded.
Some justification must be given at this time, however, for the inclusion
within Lithospermum of species which other authors have _ believed
generically separable. The species concerned are those which constituted
the genus Arnebia or its segregate Macrotomia. For convenience in
discussion I have listed below all the species that recent authors have
classified under these two genera. Those genera other than Arnebia under
which the species have been classified are named within brackets.

JOURNAL OF THE ARNOLD ARBORETUM _[VoL. xxxuIr

Ww
_
bo

Arnebia tinctoria

Arnebia decumbens

Arnebia Griffithi

Arnebia hispidissima

Arnebia Bunget

Arnebia fimbriopetala

Arnebia minima

Arnebia linearifolia

Arnebia [Macrotomia] euchroma
Arnebia {Macrotomia; Leptanthe| Benthami
Arnebia |Macrotomia] densiflora
. Arnebia fimbriata

13. Arnebia obovata

14. Arnebia Szechenyt

15. Arnebia guttata

—_—S — —
NF ODO mANIANAWHN

16. Arnebia | Macrotomia] echioides
17. Arnebia [Lithospermum] Hancockiana

Among the seventeen species that botanists have assigned to Arnebia,
the first fifteen in the above list share the greatest number of significant
characters. The two at the end of the list, although giving evidence of
relationship with the others, deviate from them in important details. Below
I have given the characters of “Arnebia,” deriving them only from the first
fifteen species in the list. The aberrant characters of the two species at the
end of the list are not included. For comparison, a characterization of
Lithospermum (exclusive of Arnebia and Macrotomia) is also provided.

ARNEBIA, P.p.

Annual or perennial; flowers frequently heterostyled; corolla yellow,
orange, blue, pink, or brownish, sometimes with an evanescent spot between
the bases of adjacent corolla-lobes; corolla-throat without appendages,
usually also lacking stipitate glands or these sparse if present; corolla-tube
glabrous or rarely hairy; corolla-nectary present or absent, weak or well
developed; style simple, forked or twice forked, stigmas two or four;
stamens whorled in corolla-tube (except in A. decumbens) ; pollen cylindric
or constricted medially, symmetric, bearing a row of four or five pores at
each end; nutlets roughened, more or less tuberculate, verrucose or rugose,
gray, brown, olivaceous or rubiginous, never white. — Asia and north
Africa.

LITHOSPERMUM, P.p.
Perennial or rarely annual; flowers sometimes heterostyled; corolla
yellow, orange, or white, never spotted; corolla-throat with or without
intruding appendages, more or less glanduliferous; corolla-tube glabrous
or rarely hairy; nectary weak or well developed; style simple; stigmas two;
stamens whorled in the corolla-tube; pollen globose, cylindric, ellipsoidal,
more or less ovoid, or more or less constricted medially, bearing six to nine

1952] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXIII 313

pores in one row at or below the middle of the grain or near one end;
nutlets usually smooth, white and lustrous, but sometimes brownish or
pitted, rough only in a few species. — America, Eurasia, and Africa.

Even with its two aberrant species eliminated, Arnebia differs sharply
from Lithospermum only in one character, nature of pollen. The annual
habit is well developed in Arnebia, but a few examples of it exist in
Lithospermum (L. matamorense, L. Pringlei). The corolla of Arnebia,
unlike that of Lithospermum, is sometimes spotted, and may be blue, pink,
or brownish, but it may also be yellow or orange and unspotted as in
Lithospermum. The corolla-throat has no appendages or at most very
weak ones (A. euchroma), but many species of Lithospermum are similarly
lacking in faucal appendages. Most species of Arnebia have no stipitate
gland in the corolla-throat, but A. obovata, A. guttata, A. Szechenyi, and A.
densiflora have them in limited numbers. The style can be forked in
Arnebia, but it can also be simple and consequently similar to that of
Lithospermum. None of the fifteen Arnebias have the smooth, polished,
frequently white nutlets common in Lithospermum. They are brown,
gray, greenish, or reddish, and roughened. Brown or gray roughened
nutlets, however, do occur in some Lithospermums (L. cinereum, L.
mirabile, L. Parksii).

The fifteen species of Arnebia cannot be distinguished from Lithosper-
mum more readily if the practice of some authors be followed and the
coarse perennials (A. euchroma, A. Benthami, and A. densiflora) be
assigned to Macrotomia and the remaining twelve species be left in a
restricted Arnebia. According to some authors Macrotomia is distinguish-

able from Arnebia by its simple style, lack of nectary in the corolla-tube,
and its coarse habit. The authors who have used these characters had not
dissected all the species of Arnebia. Simple styles also occur on A.
fimbriata, A. obovata, and A. Szechenyi, and the nectary is also lacking in
A. tinctoria, the type of the genus Arnebia. Furthermore, the coarseness
of habit supposed to distinguish Macrotomia from Arnebia becomes only
a vague matter of degree when forms of Macrotomia euchroma and
Arnebia guttata are compared. Macrotomia is not a readily definable or
useful concept, furthermore it is unnatural, for Macrotomia euchroma and
M. Benthami are not immediately related to M. densiflora.

The two Arnebias recognized as aberrant, and as yet undiscussed, remain
to be considered. The first of these, A. Hancockiana, combines characters
of Arnebia and Lithospermum. Its blue or pink unappendaged corollas are
more suggestive of Arnebia. In having ellipsoidal pollen with a row of
pores at the equator, a glanduliferous corolla-throat, a simple style with
subterminal stigmas, and smooth lustrous white nutlets, it has the characters
of Lithospermum. The other aberrant species, A. echioides, has a yellow,
spotted corolla devoid of appendages and glands in the throat, which is
very characteristic of Arnebia. So also is the shape of its broadly affixed
nutlets. Its globose pollen, with the pores on the equator, as well as
the nearly smooth surface of its nutlets, are, however, characteristic of

314 JOURNAL OF THE ARNOLD ARBORETUM _[vot. xxxi

Lithospermum. Its hairy corolla-tube is duplicated in both genera, as is
also its simple style. In bearing stamens at unequal heights within the
corolla-tube, it is dissimilar to Lithospermum but suggestive not only of
Arnebia decumbens but also of species in other genera related to Lithosper-
mum, such as Stenosolenium, Lithodora, and Alkanna. The suppressed
nectary is to be compared only with that of A. tinctoria. In most of its
characters the species is more closely allied to Arnedbia than to Lithosper-
mum.

Even when A. echioides and A. Hancockiana are not considered, the dif-
ference between Lithospermum and the fifteen species of Arnebia is weak
and uncertain. When the aberrant species are taken into consideration, the
differences between the supposedly different genera entirely disappear. If
A. echioides be assigned to Arnebia, then there is no longer any difference
in type of pollen. On the other hand, if the species be placed in Lithosper-
mum, then the spotted corolla or the corolla devoid of glands in the throat
is no longer found only in Arnebia. If Arnebia Hancockiana be treated as
an Arnebia, then the two genera again lose their differences in type of
pollen, and furthermore, the smooth, white, polished nutlets are no longer
distinctive of Lithospermum. Should A. Hancockiana be treated as a
Lithospermum that genus no longer has only orange, yellow, or white
corollas, but also pink or blue.

The distinctive developments which occur singly or collectively in species
of Arnebia and can give them an aspect so different from species of Litho-
spermum occur in only some, not all the species of that group. The spotted
corolla-limb of Arnebia occurs in only five of the seventeen species assigned
to that genus. Only six of the seventeen have colors other than yellow or
orange, such as pink, blue, or brown. The forked style is found in only
half the species. Pollen of the type with two rows of pores is found in only
fifteen of the seventeen. Since these characters are not universal in the
group and are frequently combined in species with traits characteristic of
Lithospermum, it seems futile to make further attempts at distinguishing
Arnebia from Lithospermum.

Happily, most of the species of Arnebia retain their familiar specific
epithets when the group is submerged in Lithospermum. The five excep-
tions are as follows: Lithospermum tetrastigma (Arnebia tinctoria), L.
detonsum (A. minima), L. Aucherii (A. linearifolia), L. Sewerzowi (A.
obovata), and L. Tournefortii (A. echioides).

I have not recognized Arnebia because it cannot be sharply defined or
decisively separated from Lithospermum. Another reason for not doing so
involves nomenclature. As pointed out by Rothmaler, Fedde Repert. 49:
56 (1940), the generic name, Arnebia Forsk. Fl. Aeg.-Arab. 62 (1775),
typified by Arnebia tinctoria, is antedated by Echioides {Tourn.] Ortega,
Tabulae Botanicae 7 (1773), typified by Arnebia echioides. If the group
known as Arnebia is to be given generic recognition, Echioides must be
substituted as the correct generic name.

T have seen only the second edition of Ortega’s Tabulae Botanicae (1783),
on page 16 of which the genus Echioides is keyed out in the synopsis

1952] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXIII 315

under the following final item, “Flore infundibuliformi, sed isoperimetro.
Seminibus capitis viberini oa No species, no locality, no reference,
nor other means of further identifying the plant are given. However, Ortega
makes it very clear that he used the Institutiones of Tournefort as the model
of his synopsis. The name ‘“Echioides” appears in Tournefort’s ‘Corol-
larium Institutionum,” page 46 (1703), where it is associated with the fol-
lowing description: ‘‘Echioides est plantae genus, flore monopetalo, infundi-
buliformi, sed isoperimetro, qua nota differt ab Echio. Pistillum enim abit
in fructum ex quatuor seminibus compositum, caput viperinum aemulanti-
bus. — Echioidis speciem unicam novi. — Echioides Orientale, Buglossi
folio, flore luteo, maculis atropurpureis notato. — Echioides quasi planta
ad Echium accedens.” ‘The similarity between Ortega’s short diagnosis
and the longer ene of Tournefort leaves no doubt that the same plant is
intended. The generic name Echioides Ortega (1773), applied to Arnebia
echioides, fulfils all the requirements for legitimate publication. This
is not the case with Echioides J. Agosti, De Re Bot. Tractatus 193
(1770), a name for Nonnea, recently put forward by Schwarz, Mitt.
Thuring. Bot. Ges. 11: 113 (1949). Agosti’s book, also an adaptation
of Tournefort, has descriptions of the genera, but the numerous species
mentioned bear only polynomials. Agosti, unlike Ortega, was not a fol-
lower of Linnaeus nor a user of binomial nomenclature. Because binomial
nomenclature was not adopted, Agosti’s ‘““Echioides” may be ignored. The
name Echioides Ortega remains the acceptable one for the genus formerly
known as Arnebia. Confronted with the choice of transferring all the species
of Arnebia either to Echioides or to Lithospermum, I much prefer the
latter.

KEY TO THE SPECIES

Pollen encircled by 2 rows of pores, one at each end of the grain, the rows each
with 4 or 5 pores, upper and lower half of pollen grain similar in size and

nutlets roughened, rugose or tuberculate or papillate, gray, brown, olivaceous
or rubiginous, never white and porcelain-like; plants mostly Asiatic, with
outliers in Greece and North Africa.

Plants annual.
Stigmas 4, style frequently twice forked.

Mature calyx not developing an enlarged, indurate, pentangular tube;
corolla blue or bluish, lacking a nectary in the tube; stigmas elongate;
nutlets with cordate base, flat dorsum, and convex venter..........
ee OE Dd 0 es nt 2S. Rene 1. L. tetrastigma.

Mature calyx with tube enlarged, indurate per prominently veined and
angulate; corolla orange or yellow, with a membranous collar-like
nectary in tube; stigmas subcapitate; nutlets with a broad nearly flat
base, rounded back and angulate venter.......... 2. L. decumbens.

Stigmas 2, style simply forked

Corolla-tube villulose inside near middle; filaments in short-styled flowers
more than half as long as the anthers: flowers strongly heterostylic;
pollen strongly dimorphic................... 3. L. hispidissimum.

316 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxxmII

Corolla- _ glabrous inside; oo always less than half as long as the
s; pollen monomorphi

Calyx att Aiea accrescent, aie lobes slender, 10-15 mm. long

and m. broad; corolla-limb with 5 evanescent ag spots;

flowers eave heterostylic........... 0.2.00... 4, Griffithi.

Calyx serge accrescent, prio lobes lanceolate or 2 ligulate,

10- m. long and 2-5 broad; corolla-limb not spotted;
flowers gear eal a or sopra

Corolla-lobes fimbriate, margins lobulate or lobulate-lacerate _.

5 mbriopetalum.

L. Bunget.

Corolla-lobes with margins entire or practically so.
Mature calyx 20-28 mm. long; leaves with lower surface glabrous
or nearly so; nutlets nearly as broad as long, dorsum with
medio-longitudinal depression below the middle and more or
less definitely carinate above the middle....7. L. detonsum.
Mature calyx 12-17 mm. long; leaves strigose beneath; nutlets
evidently longer than broad, back rounded and neither de-
pressed nor carinate along the medial line... 8. L. Aucherii.

Plants distinctly perennial.
Corolla-lobes with conspicuously fimbriate margins, pink rae to blue;
nectary in corolla-tube a ee lineate ring........ 9. L. fimbriatum.
Corolla-lobes entire or practically
Nectary in corolla-tube a well- Seieel villose collar.
Cerelia BHI. fuckin dr rsayieigests faniessaes4 10. L. Sewerzowt.
Corolla yellow or orange, limb usually bearing 5 black evanescent spots.
Bracts of cymes broad, more or less elliptic, foliaceous, not simulating

the narrow calyx-lobes; style not forked......11. L. Szechenyi.
Bracts narrow, speed the calyx-lobes in size and form; style
usually forked at apex.................... 12. L. guttatum.

Nectary in corolla-tube peti or absent.

Corolla yellow or orange, very large, limb 20-25 mm. broad, tube 20-37
mm. long; stamens borne at or near the mouth of the corolla- tube;
corolla-tube sparsely villulose just above the base inside; pollen
homomorphic, very large; Greece and Turkey. .13. L. densiflorum.

Corolla pink, purple or brownish, limb less than 15 mm. broad, tube
less than 15 mm. long; stamens in long-styled flowers borne near
the middle of the corolla-tube; corolla-tube glabrous inside; pollen
dimorphic; n.w. Himalayas to eastern Iran and north into Central
Asia.

adios cane much longer than broad; calyx-lobes slender,
ngate and flexuous, usually much longer than the —

bbe. ee usually evidently 3—-5-ribbed from the base. .
Sows ab as Pee a ee ey eon La ee gee ee ee ae 14. L. honth onl
Inflorescence subglobular, tending to become broader than long;
alyx-lobes lanceolate, coarse and rigid, usually equalling the
corolla-tube or surpassed by it; leaves usually with only a strong
WU Sc oa cack bas Ae eens eee 15. L. euchromon.

Pollen encircled at the middle, below the middle, or near one end with a single
row of 6-9 pores, upper and lower halves of the grain similar or differing
in outline and size; corolla orange, yellow or white (or exceptionally bluish,

1952] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXIII 317

in L. Hancockianum), with or without evident faucal appendages, usually
bearing abundant minute stipitate glands in and below the throat; nutlets
prevailingly polished and porcelain-like, completely smooth or with scattered
punctate pits, white or more or less stained with brownish (or minutely
mottled in L. Tournefortii; rough and dull only in L. mirabile, L. Parksii,
L. matamorense, L. papillosum, and L. cinereum); plants of America,
Eurasia and Africa.

Corolla orange, yellow, yellowish or greenish, not white
Flowers heterostylic or dimorphic, the two types of flowers differing in the
height at which the stamens are affixed in the corolla-tube.
Pollen broadest at the equator, pores medial.
Flowers not truly heterostylic, pollen (ellipsoidal) of one size, length
of style not correlated with the position of the stamens; Central
sia 16. L. tschimganicum.
Flowers strongly heterostylic, the two types of flowers pales not only
in oes a stamens but also in the length of the style and in the
size of
Stamens affixed at three different levels on the corolla-tube; corolla-
limb bearing 5 evanescent black spots; corolla-tube minutely
hairy inside; pollen globose; basal leaves well developed;
WAS eet hea oa 644+ cewoneed 17. L. Tournefortiz.
Stamens all affixed at the same level on the corolla-tube; corolla-limb
without spots; pollen longer than broad; America.

Stems not arising from a leaf rosette, lower cauline leaves small
and poorly developed, very much smaller than the middle and
upper ones; corolla-tube always glabrous within..........

lB. - dt multiflorum.

Stems. 4 arising from the center of : a ; well developed leaf-rosette;
basal leaves much larger than the middle and upper euline
leaves; corolla-tube hairy inside or at least so in long-styled

rs.
Basal aie large, obovate to elliptic, 15-25 mm. broad, upper
e with coarse appressed hairs only; corolla-tube not
renee ithe in short-styled flowers .... 19. L. obovatum.
Basal leaves narrow, elongate, Bbinecalnte, 5-15 mm. broad,
pper surface bearing long and short hairs; corolla-tube
always hairy inside.
Corolla funnelform, usually almost as broad as long; plant
cinereous, evidently hairy............ 20. . cobrense,
Corolla tubular funnelform, usually about twice as long as
broad; plant green, inconspicuously 6 ee
21. L. tubuliflorum.
Pollen constricted at the middle or evidently broader at one end, bearing
the pores below the middle
Basal sae larger than the middle cauline ones; stems arising from
r of leaf-rosette; pollen si long-styled flowers ovoid, of short-
ee flowers ellipsoid: Wiexse0) ec Oe 3s; 21. L. tubuliflorum.
Basal cauline leaves small and nOnnenS developed, much smaller
than the middle cauline ones; stems arising directly from a bud
on the caudex; pollen never ellipsoidal.

318 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXXIII

Nutlets strongly constricted just above base, 5 mm. long; nectary
rolla-tube a glabrous tumid ring; herbage glaucescent;
fruiting calyx usually recurved; California..................
Seas 6b PEN EAE ARE oppress wrae? 22. L.californicum.
Nutlets not constricted just above base; nectary in corolla-tube
10-lobed, villulose; herbage not glaucescent fruiting calyx re-
maining erect; eastern United States.
Corolla orange- -yellow, 13-25 mm. long, faucal appendages glandu-
iferous ridges not invaginate; mature calyx-lobes 6-13 mm
pli nutlets 3.5-4.5 mm. long......... 23.. i, siralealensé,
Corolla yellow, 10-18 mm. long, faucal appendages —
gibbose; mature calyx-lobes 5-6 mm. long; nu utlets 2-3 mm
Io A ee ere ee eer er earner ee 24. L. canescens.
Flowers not heterostyled, monomorphic.
Stamens borne at middle of corolla-tube; pollen neither ellipsoidal nor
lobose

Stems arising from a cluster of thickened fusiform roots, developing a
basal rosette of leaves which persists at least until anthesis; pollen
distinctly ovoid; southeastern United States...25. L. tuberosum.

Stems arising directly from a strong sparingly branched, dye- stained
tap-root; basal leaves small and imperfectly developed, not forming
a rosette; pollen constricted at middle or rarely a de ~

Stems with slender, loosely appressed or spreading hairs........
gp iatg ad eee habe t PEER Raa HK EHS S 26. J, eee
Stems short-strigose, the hairs closely appressed.
Mature leaves 5-15 (—20) mm. broad, firm, acute, veins 1 or 2
n either side of midrib, not obviously anastomosing nor
branched; calyx at most only shortly surpassing the corolla-
tube, commonly shorter, in fruit 3-5 (—-8) mm. long; nutlets
commonly 3 mm. long; Eurasia.......... 27. L. officinale.
Mature leaves 15-45 mm. broad, thin, frequently somewhat
acuminate, veins 2-4 on either side of midrib, obviously
anastomosing and usually branched; calyx distinctly longer
than the corolla-tube, in fruit 9-13 mm. long; nutlets 4.5—-5
mm. long; America..............:.. 28. L. latifolium.
Stamens borne near summit of corolla tube; pollen ellipsoid to globose or
practically so.

Pollen perfectly globose, 33-42 »; plant always developing some
cleistogamic flowers; chasmogamic flowers with an elongate corolla-
tube and well-developed invaginate faucal appendages.

Nutlets angulate, much punctate and roughened, narrowed to the
ase; attachment-scar flat, without a subulate projection; tap-
root short-lived, fusiform................... 29. L. mirabile.
Nutlets rounded, ellipsoidal-ovoid, usually somewhat constricted just
above base to delimit a basal collar that surrounds the markedly
concave attachment-scar, scar bearing a prominent subulate pro-
jection; taproot not fusiform, usually strong and perennial.
Corolla-lobes erose or fimbriate; fruiting calyx usually nutant or
cernuous; nutlets smooth or somewhat pitted; cleistogamous
flowéts very abundant... ..4.0<sas.e005 30. L. incisum

1952] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXIII 319

Corolla-lobes with entire margins; fruiting calyx erect.
Nutlets roughened, rigid verrucose or rugose; chasrnogamic
flowers abundant, large.................. 31. L. Parksit.
Nutlets smooth and fen chasmogamic flowers few or none,
plant commonly almost completely cleistogamic........
pa eit ah eee antl obi se wcd ae ee ae 32. L. confine.
Pollen Seer if subglobose usually with angulate sides when viewed
al profile; cleistogamic flowers produced in only one species

Corolla small, less than 10 mm. lon
Pollen cylindric, sides straight and parallel in lateral profile; stems
very elongate, 5-10 dm. Nong; Central Africa..............
ea bik. ha bwk® 6 2 33. L. afromontanum.
Pollen ellipsoidal, sides rounded or angulate in lateral profile; stems
merica.

Stems bearing numerous floriferous branchlets from the upper-
most leaf-axils, the inflorescence, hence, an elongate cylin-
drical thyrse, individual cymes even in advanced maturity
usually less than 5 cm. long, obscurely racemose; nutlets
4-6 mm. long, strongly constricted just above base; western
United States and adjacent Canada...... 34. L. ruderale.

Stems loosely branched, the cymes not aggregated into a cylin-
drical thyrse, scattered, usually becoming very elongate and
racemose at maturity; nutlets not strongly constricted just
above base; Mexico.

Margin of corolla-lobes erose-dentate or -lobulate or crisped;
fruiting calyx 8-19 mm. long; corolla-throat with scattered
glands but no invaginate intrusions; corolla very variable
in size, the later ones small and sometimes cleistogamic. .
PS AR as oh 4 ok oa 35. L. calycosum.

Margin “i corolla-lobes entire or practically so; fruiting calyx

—9 mm. long; corolla-throat with gibbous invaginations
or ae congregations of glands.

Leaves evidently veined; corolla-tube 5-9 mm. long, sur-
passing the calyx 1-3 mm., corolla-throat with well-
defined gibbous invaginations; pollen 33-37 & 25-27 uh,

ree PTOININCNE cic lk va va eee eee 36. L. Pringlei.

Leaves obscurely veined or veinless; corolla-tube 3-5.5 mm,
long not surpassing the calyx; corolla-throat with ob-
scure convex areas bearing congregations of glands.

Plant green, hispidulous, hairs spreading or loosely ap-
pressed; larger leaves 6-12 mm. broad; pollen with

prominent pores, 30-33 & 26-28 w..............

sy lilting ot dy petaah ag oo nade ete areas 37. L. indecorum.

Plant with a dense soft gray indument, abundantly ap-

pressed villulose-hispidulous; largest leaves 2-3 mm.

broad; pollen 20 X 16 uw, pores obscure..........

Weliy = painter a eae eats oaca.ai 8 38. L. jimulcense.

Corolla large, 10-40 mm. lon
Throat of corolla bearing 5 alae defined invaginate appendages,
tube usually constricted at the summit.

320 JOURNAL OF THE ARNOLD ARBORETUM _[VoL. xxxmI

Cauline leaves large, 10-30 mm. broad, evidently veined; corolla
0 mm. long; pollen evidently longer than broad... ...
gh aes paket his es vPa GRE Eee Se eS 39. L. oblongifolium.
Cauline leaves 1-8 mm. broad, veinless ; adr 10-20 mm. long;
pollen subglobose, about as long as
Taproot abruptly swollen, fusiform; vale leaf-face more or
less strigose; corolla-lobes evidently longer than broad,
narrowly obovate or oblong; style not exserted .
40. str rictum.
Taproot not abruptly thickened, lower surface of leaves gla-
brous except on midrib; corolla-lobes as broad or broader
than long, broadly attached, not narrowed at the base;
style becoming evidently exserted...... 41. L. Muellert.
Throat of corolla without well-defined invaginate appendages,
glanduliferous with the glands scattered or congregated.
Corolla bluish; leaves silky white-strigose beneath; plant with a
sprawling very loosely branched woody caudex; southwestern
COME cess apinvesih ys 2e4 oetas 42. L. Hancockianum.
Corolla yellow or yellowish; lower surface of leaves not white
nor silky strigose; caudex small and compact; America.
Margin of corolla-lobes erose, denticulate or crisped ; corolla
varying widely in size, becoming reduced in size as the
season — late one small and frequently cleisto-
gamic; Mex 35. L. calycosum.
Margin of ae on entire; flower never cleistogamic; style
becoming tardily exserted.
Corolla-tube gradually expanding, evidently broader above
middle than at the base, lobes ascending; pollen
4 L. guatemalense.
Corolla-tube subcylindric, lobes divaricate or recurved; pol-
len subglobose, about as long as broad.
Cauline leaves 10-35 mm. broad, evidently veined; nutlets
constricted just above the base; northern Mexico

and adjacent United States.......... 44. L. viride.
Cauline leaves 3-7 mm. broad, veinless; nutlets gradually
narrowed at base; Peru......... 45. L. Macbridet.

Corolla white or with only the ae appendages yellow.
Pollen constricted at the mi
Flowers heterostylic; corolla- fee villose inside........ 46. JL. discolor.
Flowers monomorphic; corolla-tube glabrous inside.
Nutlets with a constriction just above the base; America.
Plant esneire nutlets usually conspicuously pitted and more or less
HrOWHIEE «Gc ce ole dR SS ERS E SRA EOE ES 47. L. matamorense.
Plant sad ‘nutlets smooth and white.
Corolla-tube 10-15 mm. nee 2-3 times as long as calyx, limb
48. L.

10-14 mm. in diamet 8. elsonit.
Corolla-tube 2.5-3.5 mm. nae equalling or barely surpassing the
calyx, limb 4-6 mm, broad............-. L. Berlandieri.

Nutlets narrowed to a rounded base; Eura
Stem ili fa the hairs closely ene fruiting calyx 3-5 (-8)
m. long; corolla longer than broad, lobes elongate, ascending
27. L. officinale.

1952] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXIII 321

Stems with spreading or loosely appressed hairs, fruiting calyx 5-10
mm. corolla about as broad as long, lobes broad and
SLCAeIN yy eee ioe ox eens 26. L. erythrorhizon.
Pollen not es at ns middle.
Plants of South Afric
Leaves broadest ae above the base, sparsely appressed hispid, “nutlets
BUC OSG. yg eee ae ie 4k 5d nd Se 50. L. papillosum.
Leaves oa at or above the middle.
Stems 4-8 dm. tall, lower leaves 6-12 cm. long and 15-20 mm. broad;
nutlets with smooth rounded back........ 51. L. diversifolium.
Stems 1-3 dm. tall, largest leaves less than 7 cm. long and 10 mm.
broad.
Herbage with a smooth grayish indument of very abundant short
closely appressed ei nutlets tumulose on back; corolla-tube
surpassing ca 52, ‘L. cinereum.
eae villose-hispid, hairs mostly spreading: nutlets with smooth
ed back; corolla-tube elongate becoming about twice as
Med ns 56-5 5 968 4 6h oe 53. L. scabrum.
Plants of eA America.
Stems very slender, elongate and seas wee igre ee rere
54, . peruvianum.

ig)
=

Stems erect or ascending, not prostrate.
Corolla-throat bearing 5 congregations of glands but lacking definite
ee pollen ellipsoid or more commonly somewhat
BOE pasate 2S itrenimascact nicl sineacnn Gin drerin caasdaace’s 50. calcicola.
Corolla. throat with well-developed oo and velvety more
or less invaginate gibbose appenda

Stems erect, 3-8 dm. tall; pollen ellipsoid. 56. L. mediale.
Stems erect tes loosely decumbent, less than 3 dm. tall; pollen
cylin

Plant ath loosely appressed or spreading hairs; stems erect,
arising from a short-lived thickened fusiform taproot; basal
heaves. DYORG. i tne h54ano cuca ye debs 57. L. sordidum.

Plant closely strigose; stems spreading, numerous, arising from

or less well-developed caudex crowning a
strong perennial taproot.............. 58. L. distichum.
Daesash uisaee Dk a 44 eden ear SGestee. 99 2: Gayanam,

1. Lithospermum popes (Forsk.) Lam. Encyc. 3: 30 (1789),

Tab. Encyc. 1: 398 (1791).

Arnebia tetrastigma Forsk. Fl. Aeg.-Arab. 63 (1775); C. Christ. Dan. Bot.
Arkiv 4°: 17 (1922); Steud. Nom. ed. 2, 1: 56 (1840

Arnebia tinctoria Forsk. F]. Aeg.-Arab. 63 (1775); Coss. & Kral. Bull. Soc. Bot.
Fr. 4: 404 (1857); Boiss. Fl. Orient. 4: 214 (1879).

Lithospermum tinctorium (Forsk.) Vahl, Symb. 2: 33, t. 28 (1791), not L.

Echioides tinctorium (Forsk.) Rothm. Fedde Repert. 49: 56 (1940).
Lithospermum Arnebia Delile, FI. hte Ill. 7 (1813); Lehm. Asperif. 2:
316 (1818); R. & S. Syst. 4: 45 (1819)

mall, compactly branched cinereous herb, 3—9 cm. tall, of Egypt and
northern Arabia. Root annual, slender and dye-stained. Cymes dense,

322 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxu1

becoming 2—5 cm. long at maturity, bracts longer and broader than the
calyx-lobes. Corolla small, 5-8 mm. long, blue or bluish, scarcely longer
than the calyx; limb small, formed of small ascending ovate or cuneate
lobes; tube 4—7 mm. long; throat glabrous, without glands or appendages;
base of tube glabrous inside, nectary completely absent. Stamens borne at
or just below summit of corolla-tube. Pollen (fig. 28) cylindric, in lateral
profile with rounded ends and straight parallel sides, 41-46 « 33 pu. Style
reaching to the height of the stamens, bis-bifid at the apex and bearing 4
minute cylindric stigmas. Nutlets strongly compressed dorsi-ventrally,
rubiginous or brownish, obscurely tuberculate or nearly smooth, somewhat
lustrous, ca. 2 mm. long and nearly 2 mm. broad above the distinctly
emarginate base, cordate in outline, plano-convex in cross-section, with a
decidedly flat or slightly concave dorsum angled at the margin and a low-
convex ventrum with a low broad medial keel. Attachment surface on the
narrow retuse base of the nutlet, depressed and bearing a pit at the middle,
3-lobed, the lobes quadrate, the middle one extending upward on the ventral
face of the nutlet. Seeds cordate. Gynobase with 4 distinct attachment
faces, elevated and truncate at the center.

This very distinct species is the type of the genus Arnebia. It has general
relationships with the other annuals of the Near and Middle East and per-
haps its closest affinities with L. Aucheri. It presents several especially note-
worthy developments. The dorsi-ventrally compressed, plano-convex nut-
lets, with a 3-lobed attachment on the cordate base, are unique. They are
aberrant in this genus and deserve recognition as one of the very extreme
types of nutlets in all the Boraginaceae. In having four stigmas L. tetra-
stigma resembles L. decumbens, but from that and all other congeners it
differs in having elongate stigmas and not capitate, flabellate, or reniform
ones. Another unusual feature of the plant is the complete suppression of
the nectary in the corolla-tube.

2. Lithospermum decumbens Vent. Descr. Pl. Nov. Jard. Cels, t. 37
801).

Arnebia decumbens (Vent.) Coss. & Kralik, Bull. Soc. Bot. Fr. 7 402 (1857).

Echioides decumbens (Vent.) Rothm. Fedde Repert. 49: 56 (1940).

Lithospermum cornutum Ledeb. Fl. Altai. 1: 175 (1 829), Icon. Fl. Ross. 1:
t. 25 (1829) and FI. Ross. 3: 139 (1847).

Arnebia pete cee F. & M. Ind. Sem. Hort. Petrop. 1: 22 (1835) and
Ann. Sci . 2,5: 126 (1836); DC. Prodr. 10: 95 (1846); Boiss. FI.
Orient. 4: is (1879).

Arnebia orientalis [Pallas] Lipsky, Acta Hort. Petrop. 26: 513 (1910).—
Based on a misidentified drawing without analyses; ‘“Onosma orientale”
sensu Pallas, not Linnaeus; cf. Bornm. Beih. Bot. Centralbl. 33": 174 (1915),

The name L. decumbens is here applied to a bewildering assemblage of
very diverse plants occurring from Algeria, across north Africa, and in Asia
from Palestine and the Caspian area eastward to Songoria and Afghanistan.
Only a few of its many synonyms are given above. The group is one that
must be studied in the field before its many forms can be interpreted

1952] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXIII 323

properly. In the present paper I can only circumscribe it, establish its
general relations with other congeners, and indicate some of its more striking
features. The plants with which we are concerned are all annuals, charac-
terized by a combination of unusual features relating to the stigmas, the
attachment of the stamens, and the form and behavior of the fruiting calyx.

The stigmas, four in number, are borne paired terminally on the two
branches of the simply forked style or solitary and terminal on the four
branches of the twice-forked style. They are globose in form. The stamens
are not borne in perfect whorls, or in other words not all at the same level
in the corolla-tube. In some plants, particularly the western ones and
especially those from North Africa, the deviation from a whorled arrange-
ment may be slight, only one anther being slightly though distinctly lower
than the other four. In plants from the region east of Mesopotamia and the
Caspian, however, the anthers are attached at very different heights on the
corolla-tube. Commonly they are borne in three well-spaced superimposed
groups, with a pair in the throat, another pair below, and the fifth anther
lowest down in the tube, 2—4 mm. below the pair in the throat. This arrange-
ment of the anthers at differing heights can be ascertained without dis-
section by simply viewing the corolla with transmitted light.

The calyx is strongly accrescent. At maturity it becomes a tough rigid
bur-like structure which eventually detaches from the rhachis of the in-
florescence with the nutlets still enclosed. As in species of Cryptantha and
M yosotis, the bur-like calyx appears to have some role in dissemination. At
maturity it has thick prominent indurate veins and an enlarged very promi-
nently pentangular tube. The slender lobes are rigid and appressed. Be-
tween the ten indurate ribs the tube becomes membranous or opaquely
vitreous. The five more prominent ribs, at times narrowly cristate, bear
coarse warts or slender papillae, each terminated by a pungent hair.

In the forms of the species growing in Mesopotamia and westward, the
corollas are usually subtubular and 10-15 mm. long and have a limb 2.5—4
mm. broad and a tube 8-12 mm. long. The stamens, as already men-
tioned, are crowded towards the summit of the corolla-tube and frequently
deviate from a perfectly whorled arrangement only by having the base of
one anther projecting below the bases of the other four. In some plants
the stamens are borne in three groups differing in the level at which they
are attached, but unlike the anthers in the plants of Central Asia, the tip
of the lowest anther in the western forms is never below the level of the
base of the anther attached directly above. In other words, the distance
between the attachment-levels of superimposed stamens is always less than
the length of the anther, and frequently only half as long. The stigmas
are always carried distinctly below the level of the lower stamens.

The western plants of Z. decumbens occur in two well-marked forms.
One is found in the whole of the region from Transcaucasia and Mesopo-
tamia to Algeria. It is a small low plant, frequently branching at the base
and decumbent, having corollas ca. 10 mm. long, and mature calyces 7—10
mm. long. It is the form represented by the type of L. decumbens. The
other form of the western plant is restricted to Algeria and Tunis. It is

324 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxxuI

conspicuously more robust, having corollas 12-15 mm. long and mature
calyces 15-20 mm. long, and has been distinguished as Arnebia decumbens
var. macrocalyx Coss. & Kral. Before giving it a close study I had sus-
pected that it might prove to be specifically separable. It has been a sur-
prise to discover that it lacks distinctive structures and that it differs from
true L. decumbens only in the matter of degree.

The forms of the species occurring east of Mesopotamia and the Caspian
Sea differ from the western ones in being commonly much taller (1-6 dm.),
more erect plants with very much larger corollas and very unequal stamen-
attachment. In size the corolla varies widely. At times it is no larger than
in western forms (10-15 mm.), but commonly it approaches 20 mm. in
length and develops a limb 6—8 mm. in diameter. In extreme forms, however,
it can attain as much as 25 mm. in length and may develop a limb 15-20
mm. broad. The corolla-limb seems to be always slightly oblique. It is
yellow or orange (or exceptionally blue or bluish?) and frequently bears
five black evanescent spots. Selected forms of this plant can differ greatly
in size, shape, and coloration of the corolla. In any large representative
series of specimens, however, the corolla-variations are found in all com-
binations and degrees of development. Lipsky, Acta Hort. Petrop. 26:
513-26 (1910), did select out and give varietal names to some of the ex-
treme forms but stated definitely that transition between them was com-
plete and that many specimens were intermediate in character and diffi-
cult to place.

As mentioned previously, the stamens in the large-flowered eastern forms
are usually attached at unequal heights in the corolla-tube. Commonly the
attachments of the superimposed anthers are separated by a distance equiva-
lent to 14-2 anther-lengths. This departure from radial symmetry in the
androecium associated with the slightly oblique corolla-limb allows the
corolla to be described as somewhat zygomorphic. The style in such forms
commonly reaches up to slightly below the lowest stamen or, at times, to
about the same height. Occasionally (e.g. Spiridonow 167 from Golodnaja
Steppe) the flowers seem to be heterostylic, the uppermost anthers on some
plants being 3-4 mm. below the summit of the corolla-tube and surpassed
by the style, while in other plants, as is common in the species, the upper
stamens are just below the summit of the tube and the style is short.
There is, however, no associated difference in size or shape of pollen.

Some of the structures relatively constant and shared by all forms of
L. decumbens remain to be mentioned. The nectary in the corolla-tube is a
membranous collar about 0.5 mm. high. The pollen (fig. 27), despite the
inordinate diversity in size and form of corolla, is relatively constant in
form. It is elongate, in lateral profile has concave sides, and measures
43-61 30-41 p. Within the limits given there is a rough correlation
between pollen-size and size of corolla, particularly as to the smaller sizes.
The nutlets are 2.5-3 mm. long, brownish, and scantily to abundantly
tuberculate. They are roughly ovoid in form and have a rounded back and
angulate venter. The attachment surface is rounded and basal.

1952] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXIII 325

3. Lithospermum hispidissimum [Sieber] Lehm. Icones 1: 23, t. 39
(1821).

Arnebia hispidissima (Lehm.) DC. Prodr. 10: 94 (1846); Wight, Icones 4:
t. 1393 (1848); Jaubert & Spach, Ill. Pl. Orient. 4: 100, t. 363 (1852);
Boiss., Fl. Orient. 4: 213 (1879).

Anchusa hispidissima Sieber ex Lehm. Icones 1: 23 (1821), in synonymy,

Dioclea hispidissima [Sieber] Spreng. Syst. 1: 556 (1825).

Echiochilon hispidissima Tausch, Flora 12°: 643 (1829).

Strobila hispidissima G. Don, Gen. Syst. 4: 327 (1837).

Anchusa asperrima Delile, Fl. Aegypt. Ill. 7 (1813), nomen.

Arnebia asperrima (Del.) Hutch. & Dalz. Fl. W. Afr. 2: 201 and 608, f. 246

931

Echiowdes asperrimum (Del.) Rothm. Fedde Repert. 49: 56 (1940).
Lithospermum vestitum Wall. Numerical List. no. 941 (1829), nomen;
Benth. in Royle, Ill. 305 (1836); Don, Gen. Syst. 4: 325 (1837).
Toxostigma luteum A. Rich., Tent. Fl. Abyss. 2: 86 (1850).
Arnebia lutea (Rich.) Armari in Pirotta Ann. Ist. Bot. Roma 8: 161 (1904).
Toxostigma purpurascens A. Rich., Tent. Fl. Abyss. 2: 87, t. 77 (1850).
Arnebia purpurascens (Rich.) Baker, Fl. Trop. Africa 4°: 56 (1906).
Anchusa polygama Hamilton ex Don, Gen. Syst. 4: 325 (1837), in synonymy.
Arnebiola migiurtina Chiov. Fl. Somala. 227, t. 24, f. 1 (1929),

A hispid annual herb of the deserts of northern Africa and across south-
ern Asia to Pakistan. It grows 5—50 cm. tall and has erect to clecumbent
stems and a slender dye-stained root. Cymes scorpioid, numerous, ter-
minating the main stem and the numerous branches, simple, elongating,
becoming 8-12 cm. long and racemose at maturity. Calyx with very un-
equal lobes, usually much shorter than the corolla-tube, 2.5—10.5 mm. long.
Corolla 10-16 mm. long, retrorsely villulose outside; limb blue, funnelform,
4-7 (-8) mm. broad; tube 7-13 mm. long, inside villulose in 2 band ca.
1 mm. broad near the middle; throat glabrous, without glands or append-
ages, tube-nectary a very narrow lobulate sparingly villulose ring. Flowers
heterostylic. Stamens borne in the upper middle quarter of the tube on
filaments less than half as long as the anthers, or borne at the summit of
the tube on filaments as long or nearly as long as the anthers. Style reach-
ing the middle of the corolla-tube or reaching its summit or even somewhat
exserted, evidently forked, more deeply so on long styles. Stigmas reniform
or in long-styled flowers flabellate-reniform and somewhat lobed. Pollen
(fig. 7) of long-styled flowers constricted at the middle, 25-33 %& 16-25 p.
Pollen of short-styled flowers short cylindric, sides straight and parallel in
lateral profile, 37-46 30-37 ». Nutlets 1-2 mm. long, broadest at base,
ovoid, brown or rubiginous, nearly smooth or more or less ab
tuberculate.

A bristly desert herb varying greatly in habit, apparently in response to
the varying rigors of its habitat. It may be an erect, loosely and ascend-
ingly branched herb nearly 5 dm. tall, but is usually lower and more
compactly branched. The type of Arnebiola is the plant in its most de-
pressed and compact form.

The species has several unusual features. The corolla is villulose inside

undantly

326 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxx

the tube and perhaps has a slightly zygomorphic limb. The filaments of the
short-styled flowers are at least twice as long as those of the long-styled
flowers. In the long-styled flowers, as characteristic in most species of this
genus, the filament is distinctly less than half as long as its anther. In the
short-styled flowers of L. hispidissima, however, the filaments are usually
0.9-1.2 mm. long and the anthers 1.2-1.8 mm. long. The species is well
marked but is not as distinct as the repeated attempts to give it generic
recognition might suggest. Surprisingly, four different generic names have
been proposed for it: Dioclea Spreng., Strobila G. Don, Meneghinia End1.,
and Arnebiola Chiov.

4. Lithospermum Griffithii (Boiss.), comb. nov.

Arnebia Griffithii Boiss. Diag. ser. 2, 3: 135 (1856) and Fl. Orient. 4: 213
1875); Hook. Bot. Mag. 87: t. 5266 (1861); Clarke, Fl. Brit. Ind. 4: 176

1883).
Echioides Griffithii (Boiss.) Rothm. Fedde Repert. 49: 56 (1940).

A slender annual of Pakistan and Afghanistan, having erect hispidulous
simple or ascendingly branched stems 1—3 dm. tall. The very slender root
is only slightly stained with dye. Calyx-lobes slender, linear, hispid-villose,
weakly accrescent, 1 mm. wide or less. Corolla yellow or orange, 20-23
(-30) mm. long; limb spreading, 10-15 (—20) mm. broad, with 5 black
evanescent spots; tube slender, much surpassing the calyx, 15-18 mm.
long; throat glabrous or seattle glanduliferous, without appendages; tube-
nectary a narrow villulose collar. Flowers heterostyled. Stamens borne
either just above middle of corolla-tube or at the summit and almost ex-
serted. Style reaching to above the middle of the tube or reaching the
summit or even shortly exserted, forked at the apex; stigma compressed
reniform or somewhat flabellate, entire or obscurely lobed. Pollen (fig. 6)
somewhat constricted at middle, in lateral profile with concave sides, in
long-styled flowers 33-41 > 20-26 p» and in short-styled flowers 43-49 X
23-28 p. Nutlets 2 mm. long and nearly as broad at the base, brown or
rubiginous, somewhat lustrous, sparsely and coarsely tuberculate, back
somewhat cordate at base.

5. Lithospermum fimbriopetalum (Stocks), comb. nov.
Arnebia fimbriopetala Stocks in Hook. Jour. Bot. & Kew Misc. 3: 180, t. 6
51); Boiss. Fl. Orient. 4: 205 (1879).

Echioides fimbriopetalum (Stocks) Rothm, Fedde Repert. 49: 56 (1940).

A species of eastern Persia and Baluchistan known to me only from the
references cited above. It is most closely related to L. Bungei and perhaps
has a synonym in that species. According to Bornmiller, Beih. Bot. Cen-
tralbl. 33: 176 (1915), Arnebia leptosiphonoides Vatke (1875) is a syn-
onym of the present species.

6. Lithospermum Bungei (Boiss.), comb. nov.

Arnebia Bungei Boiss. Fl. Orient. 4: 215 (1879).
Echioides Bungei (Boiss.) Rothm. Fedde Repert. 49: 56 (1940).

1952] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXIII = 327

Only duplicates of the original collection, from middle eastern Persia,
have been seen. These plants are in full flower but lack fruit. They repre-
sent the long-styled form of the species. The calyx at anthesis is 10-15 mm.
long and has very slender linear lobes 0.5-1 mm. broad. The yellow or
orange corolla becomes 25—28 mm. long and has a spreading limb 10-14
mm. in diameter. The corolla-lobes have distinctly lobulate and hence
more or less fringed margins. The tube is 20-23 mm. long and is glabrous
and devoid of appendages in the throat. Its nectary is a papery collar
ca. 0.5 mm. high. The anthers are borne 15-17 mm. above the base of the
tube. The style reaches the summit of the tube, is shortly forked at the
very summit, and bears 2 simple or bilobed reniform stigmas. The pollen
grains have slightly constricted sides and measure 45-49 *& 28-30 un.

The species is closely related to L. fimbriopetalum of Baluchistan and
eastern Persia and perhaps is not distinct. Boissier stated that his species
differed in its short pubescence, slender less elongated fruiting calyx, and
its crenate rather than fimbriate corolla-lobes. The isotype of L. Bungei
dissected by me, however, has the margin of the corolla-lobes not merely
crenate, but distinctly and narrowly lobulate.

7. Lithospermum detonsum, nom. nov.
Arnebia minima Wettstein in Stapf, Denkscr. Acad. Wiss. Wien 50: 30 (1885);
Bornm. Bull. Herb. Boiss. 7: 783 (1907). Not Lithospermum minimum
Moris. (1827).

A sparingly and loosely branched herb of Transcaucasia and northern
Iran. Its thickish leaves are glabrous or subglabrous on the lower surface.
The slender root is dye-stained. The scorpioid cymes become very elongate
(to 10 cm. long) and very conspicuous at maturity. Calyx conspicuously
accrescent at maturity, 20-28 mm. long, tuberculate and hispid at the base;
lobes ligulate or lanceolate. Corolla yellow or orange, subtubular, 19-25
mm. long; lobes ascending; limb only 4—6 mm. in diameter; tube 17—20
mm. long, usually about 3 mm. longer than the calyx; throat glabrous,
without glands or appendages; nectary a lobed papery collar 0.3-0.8 mm.
high, glabrous. Anthers borne at or just beneath the summit of the tube,
not exserted. Style reaching up to 1-3 mm. below the anthers, short,
forked at apex; stigmas 2, compressed, globular or reniform. Pollen
(fig. 29) with concave sides, usually somewhat dentate at the middle,
49-59 & 30-40 ». Nutlets brownish, 3—3.5 mm. long, ca. 3 mm. broad
near the base, coarsely tuberculate, constricted above the middle, lower
half of dorsum swollen on either side of a medio-longitudinal depression,
upper half usually with a rounded medial keel.

The description given above is based on three collections: Ulja Norashen,
Nakhichevan ASSR, May 9, 1947, Grossheim, Iljinskaja & Kirpitschnikas

G); Transcaucasia, Heideman (Stockholm); and Patschinar, Iran,
Bornmiiller 7711 (G). They appear to represent the short-styled form of
the species. Another collection from Patschinar, Bornmiiller 7708 (G,
Stock.) appears to be the long-styled form. Its coarser corolla is 25 mm.
long and has a limb up to 12 mm. in diameter. Its tube (20 mm. long)

328 JOURNAL OF THE ARNOLD ARBORETUM | [voL. xxxur

does not surpass the calyx. The anthers are borne 5-6 mm. below the
summit of the corolla-tube and the style (surpassing the anthers 2-3 mm.)
reaches up to 3-4 mm. below the summit of the tube. In other respects,
however, and this includes pollen, as well as nectary and nutlets, the plant
is similar to the short-styled form described. If the plant is heterostylic
such dimorphy is unaccompanied by differences in size of pollen.

This plant of Transcaucasia and northern Iran has been confused with
L. Aucherii (Arnebia linearifolia DC.), a related but apparently distinct
species of the deserts of Egypt, Sinai, and southern Palestine. It is a more
northern plant with a more robust habit, scanty indument, larger cymes,
larger calyx, and larger bigibbose nutlets.

8. Lithospermum Aucherii, nom. nov.

Arnebia linearifolia DC. Prodr. 10: 95 (1846); Boiss. Fl. Orient. 4: 214 (1875);
Coss. & Kralik, Bull. Soc. Bot. Fr. 4: 404 (1857). Not Lithospermum
linearifolium Goldie (1822).

Echioides linearifolium (DC.) Rothm. Fedde Repert. 49: 56 (1940).

Arnebia flavescens Boiss. Diagn. 11: 117 (1849), a perer for A. linearifolia
DC. Not Lithospermum flavescens Mey. ex Steud. (1841).

A compactly branched herb 5-10 cm. tall, arising from a slender dye-
stained annual taproot. The plant is known from Egypt and eastward
into northern Arabia and southern Palestine. Its cymes, glomerate at
anthesis, remain short (1—3 cm. long) and dense even in fruit. The calyx
is strongly accrescent; mature lobes lanceolate, 10-17 mm. long, 2—-3.5 mm.
broad, and coarsely tuberculate and hispid at base. The yellow corolla is
subtubular, 10.5-13.5 mm. long, and has a limb of small ascending lobes
2—3 mm. in diameter; tube slender, 9-12 mm. long, usually surpassing the
calyx 2-3 mm.; throat glabrous, without gland or appendages; nectary
a papery lobulate collar ca. 0.5 mm. high. Anthers borne at summit of
corolla-tube, included. Style reaching up to 1-3 mm. below the anthers,
short-forked at apex, bearing 2 compressed reniform stigmas. Pollen
measuring 43-52 33-37 p, with sides slightly concave, frequently seem-
ing to be dentate near the middle. Nutlets 2.5—-3 mm. long, about 2 mm.
broad near the base, rubiginous, sparingly punctate and tuberculate and
sometimes rugose, back rounded.

If this be a heterostylic species, the five collections dissected and de-
scribed all represent the short-styled form of the species. This plant has
its closest relative in L. detonsum of northern Iran and Transcaucasia.

9. Lithospermum fimbriatum (Maxim.), comb. nov.

Arnebia fimbriata Maximovicz, Bull. Acad. St. Petersb. ser. 3, 27: 507 (1881);
Diels in Futterer, Durch Asien 3': 19 and 28, t. 3 (1911).

A low perennial with a purple-stained woody root and a pallid indument
of stiff straight loosely appressed hairs. The plant is known only from
western Outer Mongolia and adjacent Kansu. Leaves all cauline, numer-
ous, acute, mostly oblanceolate and 2-5 mm. broad; those at the base of

1952] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXIII = 329

the stem imperfectly developed. Inflorescence coarsely and loosely glomer-
ate, terminal, bracts narrow and commonly not surpassing the subtended
calyx. Corolla pink changing to dark blue when dry, 16-24 mm. long;
limb 10-17 mm, broad; lobes ovate, spreading, with conspicuously erose
or shallowly lacerate margins; tube 13-17 mm. long, surpassing the calyx
2-8 mm.; throat without appendages or glands; tube-nectary a villulose
linear ring (not a collar!). Flowers heterostylic. Stamens borne low in
the upper third of the corolla-tube (10-12 mm. above the base) or borne
at the summit of the tube and partially exserted. Style reaching to beyond
the middle of the corolla-tube or shortly exserted from it; stigmas 2,
sessile, obovate, strict, compressed, usually somewhat united dorsally.
Pollen in long-styled flowers (fig. 4) measuring 26-31 % 20-23 y, evidently
constricted at middle. Pollen of short-styled flowers (fig. 4) 35-38 X 33 p,
short-cylindric with the sides in lateral profile parallel or nearly so. Nutlets
dorsi-ventrally compressed, 2.5—3 mm. long and below the middle nearly
as broad, 1.5 mm. thick, broadest at base, dull, gray or olivaceous, with
scattered coarse tuberculations, attachment large, nearly flat.

10. Lithospermum Sewerzowi (Regel), comb. nov.
ieee [‘“Amebia”] Sewerzowi Regel, Descr. Pl. Nov. a cl. O. Fedtsch, 57
(1882).

Arnebia obovata Bunge, [Lehmann rel. bot.] Mem. savants étrang. St.
Petersb. 7: 407 (1851): Lipsky, Acta Hort. Petrop. 26: 526 (1910). Not
Lithospermum obovatum Macbr. (1916)

Echioides obovatum (Bunge) Rothm. Fedde Repert. 49: 56 (1940).

Arnebia Olgae Regel, Descr. Pl. Nov. a cl. O. Fedtsch. 57 (1882).

I have seen only two collections possibly referable to this blue-flowered
species of southeastern Turkestan. The two differ to a surprising degree
in shape of leaves, indument, tip of style, and relative length of calyx-lobes
and corolla-tube. One, von Knorring 128 from Namangan dist., Uzbek
S.S.R., has obovate or obovate-oblanceolate obtuse leaves, a hispid-villose
indument with the slender hairs arising from thickened bases, long-styled
flowers with the corolla-tube twice as long as the calyx, and the style with
a forked apex. The other collection, G. Balabajew s.n. (July 12, 1915)
from Iskanderaul, Serawschen dist, Tadzhik S.S.R., has narrow very
elongate acute oblanceolate leaves, an indument of minute retrorse hairs,
short-styled flowers with the elongate calyx-lobes as long or even longer
than the corolla-tube, and a style not forked at the apex. The collection
from Namangan, except as to indument, agrees most closely with the
emended description of Arnebia obovata given by Lipsky, l.c. The elongate
calyx-lobes of Balabajew’s collection appear to be aberrant.

The salient features of the species appear to be as follows: — A perennial
with a woody root and loosely branched stems 1-3 dm. tall. Cymes ter-
minal and very elongate in age; bracts simulating the calyx-lobes in size
and form. Corolla blue, 20-23 mm. long, with a spreading limb 8-10 mm.
broad; tube 14-20 mm. long and usually much surpassing the calyx; throat
not appendaged, glanduliferous in a narrow band at the summit; tube-

330 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxim

nectary well developed, a densely villous collar 0.8-1 mm. high. Stamens
affixed either at the middle of the corolla-tube or at its summit. Style ex-
tending up to the middle of the tube or shortly exserted, sometimes forked
at the summit. The two reniform stigmas are either sessile or terminate
the short branches of the style. Pollen of long-styled flowers measures
28-33 16-20 yn, and is evidently constricted at the middle. The pollen
of short-styled flowers measures 35-41 % 26-30 yp, is subcylindric, in
lateral profile with sides straight and parallel or nearly so. Nutlets longer
than broad, 2.5-3 mm. long, dull, gray, or olivaceous, tuberculate, and
broadest at the base.

11. Lithospermum Szechenyi (Kanitz), comb. nov.

Arnebia Szechenyii A. Kanitz, Pl. Exped. ree 42, t. 5 (1891) and Wiss.
Ergebn. Reise Szechenyi Ostasien 2: 719, t. 5 (1895).

Plant suffrutescent, perennial, 15-35 dm. tall, known only from western
Kansu. Stems erect or loosely spreading, simple or ascendingly branched,
1.5—2.5 mm. thick. Root woody, not stained with dye. Leaves firm, vein-
less, oblanceolate to elliptic, mostly 7-15 mm. broad, slightly cinereous,
bearing hairs of two distinct types, scattered stout bristles (1-1.5 mm.
long) arising from a discoid or bulbous base, and rather abundant minute
slender pallid hairs (0.2—-0.5 mm. long) which tend to be antrorse on the
upper leaf-face and more or less retrorse on the lower face. Lower cauline
leaves smaller than the middle and upper cauline ones. Inflorescence ter-
minal, scorpioid with coarse leafy bracts, in age racemose and 3—7 cm.
long. Corolla yellow, 15-20 mm. long; limb 7-10 mm. broad, with 5
evanescent black spots; lobes minutely puberulent on upper face; tube
11-15 mm. long, shortly but distinctly surpassing the calyx; throat pu-
berulent and also bearing scattered glands, unappendaged; tube-nectary a
well-developed villose collar ca. 0.6 mm, high. Flowers heterostylic. Sta-
mens borne either at the middle of the corolla-tube or at its summit. Style
simple, reaching the middle of the corolla-tube or to its summit; stigmas 2,
cordate-reniform, sessile, distinct nearly to the base or more or less united
along their medio-dorsal line. Pollen of long-styled flower not seen. Pollen
of short-styled flowers measuring 39-44 23-30 , moderately constricted
at the middle. Nutlets broadly attached basally, tuberculate, only im-
mature ones seen

The original description and plate of this species is based on the long-
styled plant from ‘“Tschung-pe-shien in valle Si-ning-fu,’ Kansu. I have
had for study a short-styled plant collected near Hsuin Hwa Hsien, Kansu,
Ching 731. The species is very distinct. Notable among its distinctive
features are the large foliaceous bracts in its inflorescence.

12. Lithospermum guttatum (Bunge), comb. nov.
aah a guttata Bunge, Ind. Sem. Hort. Dorpat. p. vii (1840) and Linnaea 15:
teratur Bericht 85 (1841); Ledeb. Fl. Ross. 3: 139 (1847); Lipsky,
e a Hort. Petrop. 26: 530 (1910); Krylov, Fl. Siberiae Occ. 8: 2282 (1937).
Pe eee guttata Farrer, English Rock Garden 1: 469 (1919)

1952] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXIII 331

Arnebia tibetana Kurz, Jour. Asia. Soc, 43°: 189 (1874); Clarke, Fl. Brit. Ind.
4: 176 (1883).
Arnebia Thomsoni Clarke, Fl. Brit. Ind. 4: 176 (1883).

Ranging from Kashmir north and northeastward in the mountains of
Central Asia to the Mongolian Altai and westernmost Kansu. Plant 1—2.5
dm. tall, flowering the first year but becoming perennial; stems one to
several, simple or bearing short ascending floriferous branches, arising
from a dye-stained taproot. Leaves thickish, oblanceolate, 3-8 mm. broad,
with obtuse or rounded apex; basal leaves well developed but usually dis-
appearing before time of anthesis. Cymes terminal, at first glomerate and
1-2 cm. in diameter, in age elongating and 5-10 cm. long; middle and
upper bracts simulating the calyx-lobes in size and form. Corolla orange
or yellow, 14-19 mm. long, commonly with 5 black evanescent spots on
the limb and occasionally with a purpurescent tube; limb spreading, 8-12
mm. broad; tube 10-15 mm. long, evidently surpassing the calyx; throat
without appendages or glands, or at times (especially in short-styled
flowers) with a narrow transverse band of glands below each corolla-lobe;
tube-nectary a well-developed villose collar. Flowers heterostylic. Stamens
borne just above middle of tube or near its summit and partially exserted.
Style reaching up to middle of tube or short-exserted, usually distinctly
forked with the branches once to several times as long as the stigmas;
stigmas 2, reniform, entire or more or Jess bilobed. Pollen of long-styled
flowers (fig. 5) measuring 25-32 > 13-20 yp, narrowed at the middle.
Pollen of long-styled flowers (fig. 5) measuring 33-37 %& 23-26 p, sub-
cylindric, in lateral profile with sides straight and parallel or slightly
concave. Nutlets 2-3 mm. long, dull, gray or olivaceous, more or less
tuberculate, broadest at the base; attachment surface large, rough, usually
with a quadrate prolongation upward and outward in the recessive angle
beneath the base of the ventral keel of the nutlet.

The species is very distinct. Although to be classed as a perennial, its
root is not as strong nor apparently as long-lived as the other perennials
of this affinity. Among the many collections examined there are frequent
individuals which appear to have flowered when less than a year old. An-
other character suggestive of some relationship with the annual species is
the forked style. That type of style prevails among the related annual
species, but among the perennial species is consistently developed only in
L. guttatum. A tendency for the nutlets to be slightly excised ventrally
at the base and for the attachment-surface to be upwardly prolonged there,
is observable in related perennials, but never to the degree characteristic
of L. guttatum. Indeed, a well-developed ventral upward extension of the
basal attachment is known in only one other species, the annual L.
tetrastigma.

13. Lithospermum densiflorum Ledeb. ex Nordmann, Bull. Acad. St.
Petersb. 2: 312 (1837).

Arnebia densiflora Ledeb. Fl. Ross. 3: 140 (1847).
Munbya densiflora (Ledeb.) Boiss. Diag. 11: 116 (1849).

332 JOURNAL OF THE ARNOLD ARBORETUM _[VvoL. xxxuI

Macrotomia densiflora (Ledeb.) Macbride, Contr. Gray Herb. 48: 56 (1916);
Farrer, English Rock Garden 1: 469 (1919).

Arnebia cephalotes DC. Prodr. 10: 96 (1846).

Munbya cephalotes (DC.) Boiss. Diag. 11: 116 (1849).

Macrotomia cephalotes (DC.) Boiss. Fl. Orient. 4: 612 (1879); Kohler,
Medizinal Pfl. 3: t. 25 (1898); Ball, Gard. Chron. ser. 3, 98: 319, f. 126
(1935).

Munbya conglobata Boiss. Diag. 11: 116 (1849).

? Arnebia macrothyrsa Stapf, Wiener Ill. Gart. Zeitung 16: 128 (1891);
Gard. Chron. ser. 3, 9: 148 and 180 (1891).

A montane plant of Turkey and Greece. The coarse caudex and the
strong perennial root are rich in purple dye. Stems 1-3 dm, tall, 3-6 mm.
thick at the base, simple below the inflorescence. Basal leaves oblanceolate,
10-15 cm. long. Cauline leaves lanceolate, 4-8 cm. long, lower and middle
one about equal in size. Inflorescence globular or corymbose, 6-10 cm.
broad, a dense aggregation of coarse, very short-pedunculate cymes arising
facesiaal on the stem and from its uppermost leaf-axils; bracts simulating
the calyx-lobes in size and form. Corolla yellow or orange, very large and
attractive; limb spreading 20-25 mm. broad; tube 20-37 mm. long, com-
monly ca. 10 mm. longer than the large calyx, without a nectary but
scantily villulose inside just above the base; throat scantily glanduliferous,
without appendages. Style forked at apex, with branches 0.5—1 mm. long;
stigmas 2, rounded, compressed. Flowers heterostylic. Corolla of long-
styled lowers slightly smaller than those with short style; tube 20-27 mm.
long and limb ca. 20 mm. broad; anthers borne slightly below mouth of
tube with their apices 1-2 and their bases 3.5—-4.5 mm. below the mouth;
style 22-32 mm. long, exserted 2-5 mm. from the tube; pollen (fig. 2)
constricted at middle, 46-50 & 26-31 ». Corolla of short-styled flowers
with tube 32-37 mm. long and limb 20-25 mm. broad; anthers borne
partially exserted in the mouth of the tube with their bases 1-2 mm.
below the top of the tube; pollen (fig. 2) constricted at the middle, similar
in form to that of the long-styled flower but much larger, measuring
65-75 38-43 p, in lateral profile frequently appearing serrate at the
middle; style 20-23 mm. long, reaching up to beyond the middle of the
tube. Nutlets 5-6 mm. long, broadest at base, gray, dull, abundantly and
very minutely papillate, coarsely rugose.

This very well marked species is notable for having the largest flowers
and the largest pollen-grains in the genus. Its nutlets, floral organization,
and pollen type are obviously those of the Arnebia group. Although
heterostyly is well developed in the species, being expressed in differences
of corolla-size and -form, length of style and pollen size, the anthers differ
very little as to the level at which they are borne in the very elongate
corolla-tube. In both types of flowers the anthers are high in the tube,
in the short-styled only 2-3 mm. higher than in the long-styled flowers.
In the former they are partially exserted and in the latter they have their
tip only 1-2 mm. below the mouth of the tube.

1952] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXIII = 333

14. Lithospermum Benthami (Wall.), comb. nov.

Echium Benthami Wall. Numerical List. no. 931 (1829), nomen; G. Don, Gen.
Syst. 4: 333 (1838).

Macrotomia Benthami (Wall.) A. DC. Prodr. 10: 27 (1846); Clarke, Fl. Brit.
India 4: 177 (1883); Hooker, Bot. Mag. 114: t. 7003 (1888); Coventry,
Wild Flowers Kashmir 1: t. 37 (1923); Blatter, Beautiful Flowers of
Kashmir 2: t. 45 (1929).

Leptanthe macrostachya Klotzsch, Bot. Reise Prinz. Waldemar. 95, t. 62
(1862).

A coarse perennial of northwestern Himalaya. Its underground parts
contain much purple dye. The flowering stems are 1-8 dm. tall and 5—10
mm. thick, and arise from the center of a leaf-cluster. The lower cauline
leaves are accordingly larger than the middle and upper ones. Leaves
usually 3—5-ribbed from the base. A very distinctive feature of the plant
is its elongate slender long-attenuate bracts and calyx-lobes and its elongate
cylindrical flower cluster. The inflorescence is a dense cylindrical thyrse
4—7 cm. thick and 5—30 cm. long. The calyx, 13-25 mm. long at anthesis,
becomes 25-35 mm. long in fruit. Its very slender flexuous lobes are
0.5-1 mm. broad at the base and very gradually attenuate. Corolla
elongate, pink to purple or maroon, 20-25 mm. long, usually shorter than
the calyx; throat without glands or appendages; tube-nectary not de-
veloped. Flowers heterostyled. Stamens borne either at the middle of the
corolla-tube or partially exserted at its summit. The style reaching up
to the middle of the tube or nearly to its summit; stigmas 2, rounded,
broader than long. Pollen of long-styled flowers strongly constricted at
middle, 30-37 16-20 ». Pollen of short-styled flowers subcylindric, 39
25-26 uy, in lateral profile with sides nearly parallel or somewhat concave.
Nutlets 3-4 mm. long, dull, abundantly and very minutely papillate, and
also coarsely and irregularly rugose and more or less tuberculate, broadest
near middle, the ventral keel continuous over the beaked apex, the attach-
ment scar frequently with 2 distinct dorsal traces.

Previously, Contr. Gray Herb. 73: 48 (1924), I stated that the corollas
of this species were slightly zygomorphic and possessed a ventral longi-
tudinal plication. These observations are entirely false and the result of
misinterpretation of the poor and inadequate material then available for

y.

No material of Arnebia speciosa Aitchison & Hemsl, (1880) from
Afghanistan, or Arnebia inconspicua Hemsl. & Lace (1891) from Baluchi-
stan, has been available for analysis. These two species are close relatives
of L. Benthami, but are probably sufficiently distinct to be worthy of
recognition.

15. Lithospermum euchromon Royle ex Benth. in Royle, Ill. Bot.
Himal. 305 (1836).
Macrotomia euchroma (Royle) Paulsen, Bot. Tidsschr. 27: 216 (1906) and
Studies Veg. Pamir 58, f. 20 (1920); Lipsky, Acta Hort. Petrop. 26: 505
(1910).

334 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxxuI

Arnebia euchroma (Royle) Johnston, Contr. Gray Herb. 73: 49 (1924).

Stenosolenium perenne Schrenk ex Fischer & Meyer, Enum. Pl. Schrenk 1: 34
(1841).

Arnebia perennis (Schrenk) DC. Prodr. 10: 95 (1846).

Munbya perennis (Schrenk) Boiss. Diag. 11: 115 (1849).

Macrotomia perennis (Schrenk) Boiss. Fl. Orient. 4: 212 (1879).

Macrotomia endochroma Hook. & Thom. ex Henders. & Hume, Lahore to
Yarkand 328 (1873); Aitchison, Jour. Linn. Soc. 18: 81 (1880), nomen; cf.
Clarke, Fl. Brit. India 4: 177 (1883).

Macrotomia onosmoides Regel & Smirn. Acta Hort. Petrop. 2: 624 (1878).

Macrotomia euchroma var. subacaulis Lipsky, Acta Hort. Petrop. 26: 510

910)

Macrotomia ugamensis Aca in Baranov, Jour. Turkest. Branch, Russian
Geogr. Soc. 17: 26, t. 3 (1925).

Arnebia tingens A. DC. Prodr. 10: 96 (April, sat

Lithospermum cyanochroum Boiss. Diag. 7: 33 (1846).

Macrotomia grandis Bornm. Oesterr. Bot. Zeit. 47: 289 (1897).

A coarse, hirsute, frequently glanduliferous perennial. In typical form
it ranges from the northwestern Himalaya northward in the mountains of
Central Asia to the Dzungarian Ala-tau. A pungently hirsute form also
occurs in the mountains of eastern Iran. This latter may deserve at least
varietal recognition. The last five names listed above apply to it. The
underground parts of all forms of the species contain an abundance of
purple dye. The flowering stems are 1—5 dm. tall and arise from the axils
of leaves that formed a sterile basal cluster the previous season. The basal
cauline leaves are small and imperfectly developed and much smaller than
the middle cauline ones, The terminal inflorescence is congested and tends
to be broader than long. The corolla is purplish white to purple or brown-
ish, 16-22 mm. long; limb 8-16 mm. broad, with ascending lobes; throat
usually unappendaged and glandless, but, particularly in long-styled
flowers, occasionally developing 5 small weakly invaginate swellings;
nectary not developed. Flowers heterostyled. Anthers borne either at
middle of tube or at its summit. The style reaches up to the middle of
the tube or is short-exserted, and is usually forked at the very summit;
stigmas 2, compressed, rounded, broader than long. Pollen in long-styled
flowers (fig. 3) strongly constricted at middle, 26-30 x 16-20 yw. In
short-styled flowers (fig. 3) pollen somewhat cylindric, in lateral profile
the sides nearly parallel or only slightly concave, 33-43 % 25-28 yw. The
gray or dusky nutlets are 3—4 mm. long, irregularly and coarsely tubercu-
late and more or less rugose, the surface dull, covered with crowded
microscopic papillae. The nutlet is broadest near the middle and has a
well-developed keel on the venter which extends up and over the beaked
apex and continues down the back of the nutlet. The attachment scar is
flat and frequently bears 2 ventral traces.

The species has its closest relative in L. Benthami and agrees with that
species in pollen, fruit, coloration, and general organization of the corolla.

1952] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXIJI = 335

16. Lithospermum tschimganicum B. Fedtsch. Bull. Jard. Bot. St.
Petersb. 5: 42 (1905); Lipsky, Acta Hort. Petrop. 26: 510 (1910).

A very distinct and unusually interesting species of Central Asia, endemic
in the mountains east and southeast of Tashkent. It is a perennial with
stems 15-60 dm. tall. The numerous evidently veined leaves, 1-5 cm.
broad, increase in size upward along the stems, the lowest one being
small and imperfectly developed. The cymes, terminal on the main stem
and on a few upper branches, are conspicuously bracted, and are small in
size, even in fruit being less than 7 cm. long. The nutlets are lance-ovoid,

—5 mm. long, and white. They are completely smooth or only sparingly
punctate. In general appearance and nutlets, and also in pollen, the plant
is more suggestive of the large-flowered American species than of any of
the other species of the Old World. The corollas, however, particularly
in having an unappendaged very sparingly glanduliferous throat, are
perhaps more suggestive of the Asiatic ‘““Arnebias” than of American
species.

Of the five collections available for study, three have provided corollas
for dissection, viz., von Knorring 180 from Osch dist., Kirghiz S.S.R., von
Minkwitz 752 from Kokand dist., Uzbek S.S.R., and von Knorring 359
from Namangan dist., Uzbek S.S.R. Though apparently all representing
one species and though indistinguishable in all other structures, these three
collections show surprising differences inside their corollas. The corollas
from Kokand and Namangan districts are 17 mm. long and have a gradu-
ally ampliate tube ca. 9 mm. long. The anthers (ca. 1.5 mm. long) are
borne on slender filaments 0.2-0.3 mm. long attached 5 mm. above the
base of the corolla. The unappendaged throat is sparingly and inconspicu-
ously glanduliferous. The glands occur very sparingly from the throat
down to the middle of the corolla-tube, with most of them confined to the
principal veins. More occur, locally abundant, just below the attachment
of each filament. The collections from Kokand and Namangan agree very
closely in all details save only the length of the style. In the former the
style is 15-17 mm. long and is shortly exserted from the corolla-tube. In
the collection from Namangan, however, it is only 5.5 mm. long and
reaches upward in the corolla-tube only to the tip of the anthers.

The dissection of the corolla on the plant from Osch, on the other hand,
reveals a very different condition. The corolla is 20 mm. long and is
tubular below the middle and gradually ampliate above. The anthers
(2 mm. long) are borne on uncinate-cuneate filaments 1.7—1.9 mm. long
affixed in the corolla-throat 13.5-14 mm. above the base of the corolla and
only 3 mm. below the base of the corolla-lobes. The filaments are most
unusual, not only in size but also in form. They are 0.5—0.6 mm. broad
at the base, gradually narrowed, curved, very strongly compressed laterally,
and provided with an evident excentric vein. There are no glands clustered
at their base. Glands, however, do occur scattered in the corolla-throat.
The style is 5-6 mm. long. As in the collections from Kokand and
Namangan, the 2 stigmas are obcordate in form and divergent from the

336 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxxmt

tip of the style. In all three collections the nectary in the corolla-tube is
glabrous and obscurely developed, at most only a vaguely defined tumid
annular area just above the corolla-base.

Since the collections from Kokand and Namangan have anthers borne
low in the corolla-tube and that from Osch has slightly larger anthers
borne high in the tube, heterostyly can be suspected. The flowers in the
collections from Kokand and Namangan, however, have low-placed sta-
mens associated with a short, as well as a very long style. Furthermore,
the pollen in all the three collections mentioned is similar in size and form
and shows no correlation with the differences in style-length or stamen-
attachment. In all collections mentioned the grains are ellipsoidal, 20-25
> 16-18 yp, and in lateral profile have rounded or slightly angled sides
(fig. 30). They bear their very obscure pores at the middle and are
broadest there. Heterostyly, certainly of the strongly developed type
characteristic of other congeners, is not present in L. tschimganicum. If
heterostyly is not present, then the collections must represent another type
of floral dimorphism or two distinct species must be represented. I find
it impossible to believe that two species with such floral differences could
be so all-prevailingly similar in all other structures. I am also aware of the
fact that in this genus, among species with elongate corollas, the stamens
are always borne above the middle of the corolla-tube when only mono-
morphic flowers are produced. Stamens borne low in the corolla-tube are
found only in long-styled flowers of heterostylic species. It is not un-
reasonable to believe, therefore, that, though L. tschimganicum may not
now be heterostylic, it may have been derived from such an ancestor.
Though now without correlated differences of pollen and style, it may have
retained only the corolla-dimorphism of its heterostylic ancestry. If so,
the condition is very unusual, and certainly unique in this genus. The
matter should be investigated by someone who can observe the plants
in the field, or at least by one who has access to many more specimens than
have been available to me.

17. Lithospermum Tournefortii, nom. nov.

Lycopsis Echioides L. Sp. Pl. ed. 2, 199 -(1762).— Based on Echioides,
Tourn. Coroll. Inst. 46 (1703) and Buxbaum, Cent. 1: 1, t. 1 (1728). Not
Lithospermum echioides Benth. in Royle (1836).

Arnebia echioides (L.) DC. Prodr. 10: 96 (1846); Bot. Mag. 74: t. 4409
(1848); Gartenfl. 25: 259, t. 877 i 76).

Anchusa echioides (L.) M. v. B. Fl. Taur.-Cauc. 1: 123 (180

Macrotomia echioides ie * Fl. Geek 4: 211 ne Farrer, English
Rock Garden 1: 469 (

Aipyanthus echioides na Bull. Soc. Nat. Moscou 26: 600 (1851).

Arnebia longiflora C. Koch, Linnaea 22: 640 (1849). Not Lithospermum
longiflorum Salisb. (1796), nor Spreng. (1825).

A handsome and very distinct species of Armenia and the Caucasus,
frequently cultivated as a rock-garden plant. The large yellow corolla
becomes 2-3 cm. long and has a tube usually twice as long as the calyx.

1952] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXIII = 337

The tube is villose inside and the broad limb has an evanescent black spot
at the base of each sinus. The throat has neither appendages nor glands
and the tube neither nectary nor rudiments of it. The stamens are affixed
on the corolla at several superimposed levels, in the long-styled flowers in
a zone below the middle of the tube and in the short-styled flowers in a
zone high in the throat. In these antheriferous zones there appear to be
two stamens that are uppermost, an alternating pair that are lowermost,
and a fifth stamen at an intermediate level. Furthermore, very close ex-
amination reveals that in both upper and lower pair one member has a
slight but still perceptibly lower attachment than its companion. Accord-
ingly, in various degrees, the stamens are all attached at differing levels.
The androecium appears to have no plane of symmetry. The style reaches
either to the middle of the corolla-tube or to its summit. The two stigmas
are broad and terminal and tend to be somewhat united, sometimes to form
a single obconic stigmatic mass. The pollen (fig. 8) is globose or very
slightly longer than broad. There are 9 obscure pores spaced about its
equator. In long-styled flowers the grains measure 30-35 p» in diameter
and in the short-styled flowers 40-50 ». The large nutlets are smooth or
only inconspicuously and minutely tuberculate towards the apex. They are
brownish, minutely mottled with purple, and not lustrous. The ventral
side is sharply angled. The attachment surface is broad, rounded, and
basal. The gynobase is very broadly pyramidal.

18. Lithospermum multiflorum Torr. in Gray, Proc. Am. Acad. 10:
51 (1874).
Lithospermum cognatum Greene ex Spengler, Oesterr. Bot. Zeit. 68: 118, f. 31
(1919).

A species of western United States (Colorado and Utah south to western
Texas and Arizona) and adjoining Mexico. Flowers are clearly hetero-
stylic. The corollas have § glanduliferous slightly swollen (but not in-
vaginate) areas in the throat and a well-developed basal nectary composed
of 10 quadrate villulose lobes. The corolla-tube is glabrous within. In
short-styled flowers the stamens are borne in the throat of the corolla and
the style reaches up to the middle of the tube. In long-styled flowers the
style is nearly exserted and the stamens are borne at the middle of the
corolla-tube and the tube above them is glanduliferous. In flowers of both
types the filaments are usually somewhat glanduliferous at and just below
their base. The pollen (fig. 14) is characteristically ellipsoid and broadest
at the equator, but occasionally may become almost cylindric with the
sides nearly parallel. The pores are in a single medial row. In long-styled
flowers the grains measure 16-20 * 8-13 yw and in short-styled flowers
25-33 20-28 w. The two terminal stigmas are somewhat semicircular
or flabellate and frequently bilobulate. They tend to be spreading or even
slightly reflexed. Commonly they are umbonate at the base just above
their attachment to the end of the style. I have had a great number of
specimens of this species for study, among them, surprisingly, are very few
in the fruiting state.

338 JOURNAL OF THE ARNOLD ARBORETUM _[VoL. xxx1II

19. Lithospermum obovatum Macbride, Gontr. Gray Herb. 48: 56
(1916).
Lithospermum gentianoides Brand, Fedde Repert. 28: 15 (1930).

A well-marked species having obvious relations with L. cobrense. It is
known only from the Sierra Madre Occidentale of northern Mexico
(Chihuahua and Durango). The thickish basal leaves are very broad,
obovate to elliptic, and form a conspicuous rosette. They are conspicuously
veined, with the veins not merely prominent beneath but also strongly
sulcate on the upper face. Their indument consists of slender elongate
hairs only, not of a mixture of long and short hairs as in L. cobrense. The
corolla resembles that of L. cobrense in form but differs in a number of
other respects. The glands in the throat of the corolla tend to form small
but evident aggregations below the base of each corolla-lobe. The corolla-
tube inside is hairy only in long-styled flowers. In short-styled flowers the
tube is glabrous. The stamens in the long-styled flowers are borne in the
lower third of the corolla and hence lower proportionately than in L.
cobrense. The pollen (fig. 16) is ellipsoid and broadest at the equator.
It measures 14-18 & 10-13 » in long-styled flowers and 25-28 « 18-23 p»
in the short-styled flowers. The nutlets of this species are unknown.

20. Lithospermum cobrense Greene, Bot. Gaz. 6: 157 (1881).

A species known only from Arizona, New Mexico, and western Texas,
and in the mountains of northern Mexico, in Chihuahua and Durango.
The flowers are strongly heterostylic. The funnelform orange or yellow
corollas have the tube hairy inside and possess a well-developed basal
nectary composed of 10 crowded quadrate lobes. The throat is unappend-
aged but evidently glanduliferous. The glands are numerous and generally
distributed. They are not aggregated as in L. obovatum. In the short-styled
flowers the stamens are borne in the throat and the style is very short,
usually not half the length of the calyx. In the long-styled flowers the
stamens are borne just below the middle of the corolla and the style reaches
up into the glanduliferous.throat. The two stigmas are terminal, spreading,
and semicircular or obovate. The pollen (fig. 15) is ellipsoidal and usually
broadest at the equator. In the two types of flowers it differs only in size
and in the position of the 7-9 pores. In the long-styled flowers the pores
tend to be slightly submedial rather than exactly medial in position. The
grains of the long-styled flowers measure 21-25 12-20 yn, and those of
the short-styled flowers 26-39 « 23-31 up.

21. Lithospermum tubuliflorum Greene, Pittonia 1: 155 (1888).
Lithospermum lasiosiphon Johnston, Contr. Gray Herb. 70: 22 (1924).

A species showing relationship with L. cobrense and L. obovatum. It is
known only from the east base of the Sierra Madre Occidentale in
Chihuahua and Durango in northern Mexico. The foliage is thin, light
green, and inconspicuously hairy. The basal leaves form a rosette and,
like those of L. obovatum, have the veins evident on both surfaces. The

1952] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXIII 339

flowers are strongly heterostylic. The corolla is tubular-funnelform with
ascending lobes and most resembles that of L. multiflorum in general form.
The throat is very sparingly glanduliferous and completely lacking in
appendages. The tube is always hairy inside. The nectary is similar to
that of L. cobrense but is less prominent. The stamens of the long-styled
flowers are borne extremely low, in the lower third of the tube. The pollen
(fig. 10) in the two types of flower differs in shape as well as size. That
of long-styled flowers is broadest and shouldered just below the middle.
It measures 26-30 16-25 yw. The pollen of short-styled flowers measures
33-40 26-32 p» and is ellipsoidal and broadest at the equator. There
are apparently 8 pores. The nutlets resemble those of L. cobrense but are
slightly smaller and perhaps more sharply keeled.

22. Lithospermum californicum Gray, Proc. Am. Acad. 10: 51 (1875).

A species known only from northern California and adjacent Oregon.
Although in many ways very suggestive of L. ruderale, and especially so in
form and organization of corolla and nutlets, L. californicum differs in type
of inflorescence, pollen, and presence of heterostyly. Distinctive features of
the species are its slightly glaucescent herbage and its recurved fruiting
calyx. The elongate yellow corolla is gradually ampliate, 10-15 mm. long,
and has a limb of loosely ascending lobes 5-10 mm. in diameter. The
throat is glanduliferous and sometimes bears very obscure swellings below
each of the corolla-lobes, but is otherwise unappendaged. The stamens
are borne either slightly above the middle of the glabrous corolla-tube or
near its summit. The style reaches up to near the middle of the tube or
is nearly exserted from it. The two stigmas are terminal. The pollen on
the long-styled flowers (fig. 9) measures 25-28 & 10-18 (-22) yp. It
tends to be rather variable in form, sometimes resembling that of the
short-styled flower and sometimes being distinctly constricted at the middle.
The pollen of the short-styled flowers (fig. 9) measures 33 & 20-25 » and
is nearly ovoid or is ovoid with distinct shoulders. The nectary in the
corolla-tube is an obscurely lobed glabrous tumid ring. The smooth white
ovoid nutlets, ca. 5 mm. long, are rather pointed and have a strong con-
striction just above their base.

23. Lithospermum caroliniense (Walt.) MacMill. Metasp. Minn.
Valley 438 (1892).

Anonymos carolintense Walter, Fl. Carolina 91 (1788).

Onosmodium carolinianum (Lam.) A.DC. Prodr. io: 70 (1846).

Batschia Gmelini Michx. Fl]. Bor. Am. 1: 130 (18

Lithospermum Gmelini (Michx.) eer spate fo Manhattan 30 (1894).
Anchusa hirta Muhl. Cat. 19 (1813), no

Lithospermum hirtum (Muhl.) Lehm. hevcen 2: 304 (1818).
Lithospermum strigosum Raf. New FI. No. fea a 18 (1836).
Lithospermum bejariense A. DC. Prodr. 10: 79 (1846).

Lithospermum croceum Fernald, Rhodora 37: oe t. 376 (1935).

340 JOURNAL OF THE ARNOLD ARBORETUM [voL. xxxi

A well-known species, widely distributed in eastern United States. Plant
3-10 dm. tall, arising from a strong dye-stained root, and commonly be-
coming very dark in drying. Flowers heterostyled. Corolla orange-yellow,
13-25 mm. long, with a funnelform limb nearly as broad. Corolla-tube
cylindric, in long-styled flowers 7-8 mm. long, about equalling the calyx;
in short-styled flowers 10-12 mm. long, surpassing the calyx by as much
as 3-4 mm. The corolla-throat is very obscurely if at all invaginate, its
appendages are represented only by 5 arcuate clusters of glands at the
summit of the tube. Inside the tube is sparingly glanduliferous above the
middle but otherwise glabrous. The anthers are borne either above the
middle of the tube or just below its summit. The style reaches almost to
the middle of the tube or almost to the summit. The 2 stigmas are terminal.
The corolla-nectary is 10-lobed and minutely villulose. The pollen (fig. 12)
has 7—9 pores, and in the two types of flowers differs in size as well as
shape. In long-styled flowers it is elongate, constricted at the middle, and
measures 33 >< 13-18 p». In short-styled flowers the pollen is ovoid, with
shoulders, and is broadest at one end. It measures 33-39 & 25-33 yw. It
may be noted in passing that the floral differences used by Fernald to
distinguish L. croceum from L. caroliniense are those which distinguish
the short- and long-styled flowers of the species.

24. Lithospermum canescens (Michx.) Lehm. Asperif. 2: 305 (1818).

Batschia canescens Michx. Fl. Bor. Am. 1: 130, t. 14 (1803).

Anchusa canescens Muhl. Cat. 19 (1813).

Anchusa virginiana L. Sp. Pi. 133 (1753). Not Lithospermum virginianum L.
(1753),

Lithospermum sericeum Lehm. Asperif. 2: 306 (1818).

Batschia sericea (Lehm.) R. & S. Syst. 4: 743 (1819).

Batschia conspicua R. Br. in Richardson, Bot. Append. to Frankl. Jour. 732
(1823).

A native of eastern United States. Stems 1-4 dm. tall, arising from a
strong dye-stained root. When young the plant has a very distinctive
vesture of slender, appressed, somewhat silky hairs. The flowers are
heterostylic. The yellow corolla is 10-18 mm. long and has a funnelform
limb 11-15 mm. in diameter. Its tube is cylindrical, 7-8 mm. long, and
evidently longer than the small calyx. The throat bears weakly invaginate
appendages which are usually gibbose and somewhat velvety as well as
glanduliferous, especially on the sides. The tube is glanduliferous above
the middle, and most abundantly so in long-styled flowers. Its nectary is
10-lobed and minutely villulose. The anthers are borne either just below
the middle of the corolla-tube or near its summit. The style is either very
short, 1-2 mm. long and less than a third of the tube-length, or surpasses
the stamens and becomes 5—9 mm. long. It is terminated by 2 small
stigmas. The pollen (fig. 11) of the long-styled flowers is constricted at
the middle and measures 20-25 & 10-13 yw. That of the short-styled
flowers (fig. 11) is shouldered-ovoid, broadest at one end, and measures
25-33 & 14-23 yp.

1952] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXIII 341
25. Lithospermum tuberosum Rugel ex DC. Prodr. 10: 76 (1946).

A species of southeastern United States readily recognized by its rosette
of basal leaves and clustered fleshy fusiform roots. The flowers resemble
those of L. officinale and allies. Lithospermum tuberosum probably has its
closest relations with those species. The yellow or yellowish corolla is
4.5—-6 mm. long and has a tube 3-4 mm. long. The corolla-lobes, 1.5—-2 mm.
long, are longer than broad and ascending. The throat bears 5 small
intruding velvety trapeziform appendages formed by invagination. Below
the appendages the throat is sparingly glanduliferous. The calyx is shorter
than the corolla-tube or at most equals it in length. The filaments are
attached at the middle of the corolla-tube and have no glands at their base.
The nectary is a narrow glabrous flange. The style is 2-3 mm. long and
bears 2 small terminal stigmas. The pollen (fig. 20) is somewhat ovoid,
being broadest towards one end and commonly measuring 20 X 16 p. The
nutlets are small, 1.5—2.5 mm. long, and may be abundantly punctate.

26. Lithospermum erythrorhizon Sieb. & Zucc. Abh. Bayer, Akad.
Wiss. 42: 149 (1846); Hara, Bot. Mag. Tokyo 51: 50 (1937) and
Enum, Spermatoph. Japon. 1: 176 (1948).

Lithospermum officinale 8 erythrorhizon (Sieb. & Zucc.) Maxim. Bull. Acad.
St. Petersb. 17: 441 (1872).

Lithospermum officinale subsp. erythrorrhizon (Sieb. & Zucc.) Hand.-Mazz.
Symb. Sin. 7*: 817 (1936).

Lithospermum murasaki Siebold, Syn. Pl. Oecon. Jap. 32 (1830), nom. subnud.

Lithospermum officinale var. japonica Miquel, Ann. Mus. Lugd.-Bat. 2: 94

(1865).
Lithospermum albiflorum Vaniot, Monde de Plantes ser. 2, 7: 42 (1905).

This species is a very close relative of L. officinale and occurs beyond
the eastern limit of the latter in China and northward in Korea and Japan.
It differs in having spreading rather than closely appressed hairs on the
herbage, a larger, somewhat differently shaped corolla, and a larger, more
strongly accrescent calyx. Its root appears to contain larger quantities of
purple dye than that of L. officinale. The corolla has been illustrated as
pure white, and collectors have so reported it. Unlike that of ZL. officinale,
the limb of the corolla is spreading and formed of rounded lobes about as
broad as long. Its diameter is commonly about equal to the total length
(5-9 mm.) of the corolla. The corolla-tube, 3-4 mm. long, is usually
equalled or shortly surpassed by the calyx-lobes. The faucal appendages
are trapeziform invaginations which are velvety on the summit and glandu-
liferous on the side. The filaments are affixed at the middle of the tube
and are glanduliferous at the base. The nectary is a glabrous flange. The
pollen, 13-16 X 8-10 uy, is constricted at the middle or rarely has near
straight and parallel sides. The fruiting calyx is commonly 5-10 mm. long
and usually several times longer than the nutlets.

Plants from Afghanistan, Pakistan, and Kashmir agree with those from

342 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxxnr

eastern Asia in having spreading hairs on the stems. In characters of calyx
and corolla, however, they agree with typical L. officinale and probably
deserve to be classed as variants of that species.

27. Lithospermum officinale L. Sp. Pl. 132 (1753).

Native in Europe and east to central Asia. In Afghanistan, Kashmir
and Pakistan it is replaced by a variety with spreading hairs, and in China,
Korea, and Japan by the closely related L. erythrorhizon, Over its wide
range L. officinale remains a reasonably constant species. Its stems are
closely strigose and have no spreading hairs. The yellowish or greenish,
or sometimes nearly white corollas are 4-6 mm. long. The corolla-limb
is at most 4 mm, broad and consists of ascending usually oblong lobes
1-1.5 mm. long. The tube is 2.5-3.5 (or rarely 4) mm. long. The calyx
may be shorter than the corolla-tube or slightly longer, but commonly it
has about the same length. The corolla-throat bears 5 trapeziform, in-
truded, distinctly invaginate appendages which are densely velvety on
top and glanduliferous, particularly on the sides. Below the level of the
faucal appendages glands are scattered. The filaments, attached at the
middle of the corolla-tube, are usually glanduliferous at the base. The
style is 1-2 mm, long and bears 2 small terminal stigmas. The nectary on
the corolla-tube is a glabrous, entire, or obscurely lobed flap. The pollen
(fig. 32) is constricted at the middle and measures 13-16 X 8-10 p. The
nutlets (2.7-3.8 mm. long) are most commonly about 3 mm. in length
and more than half as long as the fruiting calyx.

28. Lithospermum latifolium Michx. Fl. Bor. Am. 1: 131 (1803).
Lithospermum officinale 8 latifolium Lehm. Asperif. 2: 311 (1818).
Cyphorima latifolia (Michx.) Raf. in DC. Prodr. 10: 76 (1846).

Cyphorima lutea Raf. Cat. 13 (1824).
Lithospermum luteum (Raf.) House, Bull. N. Y. State Mus. 243-4: 61 (1923).
Lithospermum lutescens Coleman, Cat. Pl. Grand Rapids 29 (1874).

A species of northeastern United States very closely related to the
Eurasian L. officinale. The pale yellow corolla is 5—7 mm. long and has
ascending lobes nearly as broad as long. Its tube is 2.5—3.5 mm. long and is
evidently shorter than the slender calyx-lobes. The throat has 5 intruded
velvety trapeziform appendages formed by invagination. The inner face of
the appendages is glanduliferous and so also is the throat directly below
them. The filaments arise at the middle of the corolla-tube and usually
bear some glands at their base. The style is very short, usually only 1 mm.
or less long, and bears two small terminal stigmas. The nectary is a gla-
brous flange. The pollen (fig. 25) resembles that of L. officinale but is
slightly larger and less constricted. It measures 16-20 < 8-14 p. The
nutlets, 4.5—5 mm. long, are usually much surpassed by the calyx-lobes.

29. Lithospermum mirabile Small, Fl. Southeast. United States 999
and 1337 (1903).

Lithospermum longiflorum var. mirabile (Small) Brand, Fedde Repert. 28:
14 (1930)

1952] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXIII 343

A very distinct species which is most closely related to L. incisum and
L. Parksii. It is confined to the eastern half of Texas. The plant is a
biennial or a short-lived perennial and has a thickened, dye-stained, fusi-
form taproot. The stems, usually few, bear elongating cymes that produce
only conspicuous chasmogamic flowers or first chasmogamic flowers and
then later cleistogamic ones, Both types of flowers are fertile. The con-
spicuous flowers have corollas similar in form, size, and organization to
those of L. incisum, but differ in their darker coloration and in the entire
margin of their lobes. The nutlets are very distinctive and are readily
separable not only from those of related species but from those of all other
members of the genus. They are brown, dull, abundantly punctate, rough,
and distinctly angulate. They have only a weakly defined collar at the
base and their attachment-scar is flat. The gynobase is depressed pyramidal
or nearly horizontal. The pollen resembles that of L. incisum in form and
size (33-44 ), but unlike that species tends to have its pores slightly
visible. These pores are 7 or more commonly 8 in number and are equally
spaced about the equator.

Meriting publication are some interesting field observations concerning
L. mirabile contained in a letter addressed to me on Aug. 5, 193 7, by Dr.
H. B. Parks, then Chief of the Apicultural Research Laboratory, San
Antonio, Texas. “This is the plant that sent me on the quest to find out
something about this genus. I noted that in travelling through the country
in early spring there were two colors of flowers among the Lithospermums
seen by the roadside. A casual investigation revealed the fact that the
darker yellow one had smooth edges to the corolla, while the light yellow
one was toothed or crisped. On investigation I became convinced that the
yellow flowered one was L. mirabile and now after having raised the plant
and produced the seeds I am sure of this determination. Lithospermum
mirabile does produce a few small cleistogamous flowers which sometimes
develop fruits, however the most of its fruits come from perfect flowers.
This plant contrary to statements is not restricted to the vicinity of San
Antonio. It seems to be restricted to the Eocene plains which stretch
across Texas south of a line from Texarkana to Del Rio. The plants are
generally found in poor, gravelly or clay soils. They commence to bloom
by the latter part of March and will stay in bloom until the first of July.
From a distance, with the exception of color in the flowers, there is little
to distinguish this plant from L. incisum; however, the whole manner of
growth, the shape of the leaves, and the shape of the seed show it is a
good species. I have collected it from Bexar County on the north to
Brooks County on the south and from Medina County west to Anderson
County on the east. In many places it is more abundant than L. incisum,
a thing which I think is due to the difference in soil requirement.”

30. Lithospermum incisum Lehm. Asperif. 2: 303 (1818).
Lithospermum angustifolium Michx. Fl. Bor. Am. 1: 130 (1803), not Forsk

Lithospermum linearifolium Goldie, Edinb: Phil. Jour. 6: 322 (1822),

344 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxxu

Batschia longiflora Nutt. in Pursh, Fl. Sept. Am. 1: 132 (1814).
sa il longiflorum (Pursh) Spreng. Syst. 1: 544 (1825), not Salisb.
(1796).

Batschia decumbens Nutt. Gen. 1: 114 (1818).

grt alg decumbens (Nutt.) Torr. Ann. Lyceum N. Y. 2: 225 (1826),
not Vent iF

evans crypthantiflorum Brand, Fedde Repert. ee 13 (1930).

Lithospermum boreale Brand, Fedde Repert. 28: 13 (1930).

A species widely distributed in the United States, chiefly on the Great
Plains and along the Rocky Mountains, and extending into adjacent
Canada and Mexico. The plant has been given many names. Only the
older and the most recent ones are given above. For additional synonymy
see Johnston, Contr. Gray Herb. 70: 24 (1924).

In the spring the plant produces compact terminal clusters of very
conspicuous yellow flowers. The tube of the corolla is 20-35 mm. long,
2-3 mm. thick, and two to three times as long as the calyx. The limb is
8-15 mm. in diameter, and its broad rounded lobes have erose-fimbriate
margins. There are five evident faucal appendages. These are invaginate,
trapeziform, and moderately glanduliferous. The stamens are always
borne high in the corolla-tube, 1-2 mm. below the base of the appendages.
The style varies considerably in length, from half as long as the corolla-
tube to slightly longer. There is some evidence for believing that the style
may change in length between anthesis and the time the corolla is shed.
The two stigmas are semicircular or ovate and terminal. The nectary at
the base of the corolla-tube is very weakly developed or absent and com-
monly consists only of five very minute tufts of hairs.

The nutlets are elongate, 2.5-3 mm. long, broadest near the middle and
smooth or somewhat punctate. A lineate constriction just above the base
gives the nutlet a more or less well defined basal collar. This collar,
commonly slightly different from the rest of the nutlet in color and surface,
surrounds the concave or excavated basal attachment-scar. The scar is
notable not only for its concavity, but also for bearing a subulate ap-
pendage, ca. 1 mm. long, resulting from a projection of tissue surrounding
the dorsal vascular bundles of the nutlet. The gynobase is distinctly
pyramidal, about one and a half times as broad as high. After the fall of
the nutlet each of its attachment-surfaces has a central pit, the socket
into which the appendage on the nutlet-scar formerly fitted. As a result
of a bend in the pedicels, the fruiting calyces are usually nutant or cernuous.

After the appearance of the bunched large conspicuous vernal flowers,
the plant becomes much branched and then produces cleistogamic flowers
exclusively, and these in very great abundance. The early flowers with
conspicuous corollas mature few fruits, but the later flowers with minute
(1-3 mm. long) closed corollas are extremely fertile. This fact is readily
established by observing the length of the persistent style. Few fruits are
to be found associated with the long (10-30 mm.) style of the conspicuous
vernal flowers. The pollen (fig. 17) of this species is spherical and bears
7 or 8 obscure pores equally spaced about the equator. In chasmogamic

1952] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXIII 345

flowers it measures 33-42 » in diameter. In cleistogamic flowers it is
slightly smaller, 27-35 yu, but otherwise indistinguishable.

31. Lithospermum Parksii, sp. nov.

Herba perennis 2-3 (—5) dm. alta erecta; caulibus pluribus praesertim
supra medium adscendenti-ramosis; foliis costatis sed saepissime enervatis
griseis adpresse villoso-hispidulis, maturitate margine saepe evidenter
revolutis; foliis inferioribus majoribus oblanceolatis 5-10 cm. longis 5-10
mm. latis obtusis; foliis caulinis mediis linearibus obtusis saepe 2—4 cm.
longis 2-4 mm. latis; cymis conspicue bracteatis caules ramulosque ter-
minantibus simplicibus, juventate circinatis, maturitate elongatis racemosis
ad 10 cm. longis, saepe floribus chasmogamicis abundantibus gestis et
solum senescentibus apicem versus flores cleistogamicos paucos gerentibus
vel rare a basi usque ad apicem floribus cleistogamicis donatis; floribus
chasmogamicis fertilibus, corolla more ZL. incisi sed margine loborum
integerrimo, tubo saepe 15 mm. longo 2-3 mm. crasso, limbo 10--12 mm.
diametro, lobis rotundis integerrimis, fauce appendiculis trapaeziformibus
glanduliferis instructo, calyce supra medium tubi corollae attingenti, in
statu fructifero erecto, lobis calycis linearibus margine revolutis quam
nuculis saepe subtriplo longioribus; floribus cleistogamicis fertilibus, corolla
1-3 mm. longa perinconspicua, calyce ei florum chasmogamicorurm simili;
nuculis elongatis opacis densissime verrucosis et punctatis supra basim
aliquantum constrictis, basi tumidis, cicatrice concava.

TEXAS: Edwards County: 25 mi. n.w. of Rocksprings, Cory 24195 (G);
7.25 mi. n.w. of Rocksprings, Cory 38768 (G); Little Hackberry Creek, 14.5 mi.
s.e. of Rocksprings, Cory 42962 (G); Pulliam Creek below Blue Hole, Cory
43779 (G). Kinney County: 23.7 miles north of Brackettville, Cory 645
(G). Sutton County: Substation no. 14, Pasture E, Corey 24189 (G).
Val Verde County: 6.33 miles south of Loma Alta, Cory 41685 (G); Devils
Lake, ca. 20 miles n.-n.w. of Del Rio, McVaugh 7725 (type, Gray Herb.).
Jeff Davis County: Piedra Pinta, 1851, Wright, field no. 110 (G); Brewster
Glass Mountains, 1936, Cory (G); Gage Ranch, Glass Mts., Warnock 553 (G);
Hess Canyon, Glass Mts., Warnock 294 (G); Jim Nichol’s Ranch, Old Blue Mt.,
Warnock 567 (G); Sierra del Norte, ca. 10 mi. s.e. of Alpine, McVaugh 7856 (G).

Lithospermum Parksii var. rugulosum, var. nov.

A varietate genuina differt pilis sparsioribus vestita, radice minus
persistente; nuculis subnitidis dense rugosis haud verrucosis.

MEXICO: Coanuita: near Rancho Encampanada, Sierra Hechiceros, Stewart
206 (G); Muzquiz, Marsh 2108 (G); Caracol Mts., southeast of Monclova,
Palmer 897 (G); Soledad, west of Monclova, Palmer (G). Nuevo Leon:
descent into Alamar, ca. 15 mi. s.w. of Galeana, Mueller 598 (type, Gray Herb.).
TAMAULIPAS: cliffs s.w. of Victoria, Runyon 726 (US).

A species known only from areas of limestone in southwestern Texas
and northeastern Mexico. Obviously a close relative of L. incisum and
native to an area in which that species occurs also. It is, however, certainly

346 JOURNAL OF THE ARNOLD ARBORETUM [VvoL. xxx

distinct! Among the characters distinguishing it from its relative are the
olivaceous rather than gray-green herbage, the loosely appressed hairs of
stem and leaves, the entire margins of the corolla-lobes, and the verrucose
or rugose nutlets. Unlike L. incisum the plant does not become diffusely
branched. Its cymes are all elongate and racemose at maturity.

The chasmogamic flowers are usually fertile. They are the flowers first
developed in the spring and frequently make up the majority of those
developed on the elongating cymes. The last flowers on the cyme, however,
are usually cleistogamic. Cymes developed late in the season may bear
only cleistogamic flowers. Mature, fully elongate cymes, whether producing
open or closed flowers, are similar in size and form. This is very different
from the condition in L. incisum. The chasmogamic corolla of L. Parksti
differs from that of L. incisum only in its paler yellow color and entire
lobes. The appendages, stamens, pollen, and style in the two species are
indistinguishable. The nutlets of the two species, though very different in
surface features, are rather similar in size, form, and attachment. In
L. incisum the nutlets are smooth or merely pitted. In L. Parksii they are
covered with crowded warts or are distinctly rugose. The fruiting calyx
is erect and never nutant or cernuous as prevalent in L. incisum

As here defined L. Parksii included two recognizable forms, the var.
typicum of Texas and the var. rugulosum of near-by Mexico. Possibly
these should be treated as two closely related species. However, until a
larger suite of specimens of the Mexican plant becomes available for com-
parison, and especially until collections showing the fully mature nutlets
can be studied, the present disposition of the two plants seems desirable.
The Texan plant is much more vigorous and obviously has more numerous,
stiffer stems and a stronger and much more persistent root than does the
Mexican. All the plants of Texas give the appearance of having grown in
sunny exposed places, while those from Mexico seem to have come from
partial shade, perhaps from open woodland. The most important difference
between the varieties, however, is in the nutlets. The nutlets of the Texan
plants are coarsely and densely verrucose. The surface is opaque and
covered with very crowded warts which are separated here and there by
pits and deep narrow irregular fissures. On the other hand, the nutlets of
the Mexican plants are slightly lustrous and are roughened only moderately
by crowded irregular ridges.

The species is named for Dr. H. B. Parks, former chief of the Texas
Apicultural Research Laboratory, to whom I am indebted for many
valuable notes concerning the Texan species of Lithospermum, the results
of his cultivation and observation of the species over many years. The
present plant, which he first called to my attention fifteen years ago, is
very fittingly associated with his name.

32. Lithospermum confine, sp. nov.

Planta erecta strigosa perennis e radice palari valida erumpens, 2—4 dm.
alta; caulibus pluribus erectis praesertim supra medium ramosis foliosis;
foliis costatis sed enervatis viridibus utrinque strigosis numerosis obtusis

1952] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXIII 347

2-6 cm. longis 1-10 mm. latis, inferioribus oblanceolatis, superioribus
lanceolatis vel linearibus, margine anguste revolutis; cymis caules et ramos
terminantibus, juventate glomeratis maturitate ad 10 cm. longis racemosis
foliosis distantifloris; inflorescentiis a basi fere ad apicem flores chasmo-
gamicos gerentibus solum apice floribus cleistogamicis donatis vel cymis
omnino flores cleistogamicos proferentibus; floribus chasmogamicis flavis,
tubo 7-10 mm. longo 1-2 mm. crasso calyce subduplo longiori, limbo
5-6 mm. diametro, lobis rotundis margine integris, fauce appendiculis
trapeziformibus invaginatis glanduliferis donato, stylo 5-10 mm. longo;
floribus cleistogamicis inconspicuis 1-3 mm. longis, stylo 1.5-3 mm. longo;
calyce subanthesi 4-5 mm. longo, lobis linearibus statu fructifero saepe
ad 10 mm. longis nuculis duplo longioribus; pedicello fructifero 2-10 mm.
longo erecto; nuculis laevibus nitidis albis supra basim plus minusve
constrictis 3—3.5 mm. longis 2—2.5 mm. crassis.

ARIZONA: Chiricahua Mine, 6500 ft. alt., Oct. 17, 1907, Blumer 1796 (G);
8 m. north of Metcalf, Greenlee Co., June 5, 1935, Maguire et al. 11805 (G).
TEXAS: Smith Canyon, Guadalupe Mts., Culberson Co., 5500 ft., Sept. 15,
1948, Warnock 113 (G); (?) Little Hackberry Creek, 14.5 mi. southeast of
Rocksprings, Edwards Co., Aug. 9, 1943, Cory 42961 (G).

MEXICO: Coauutta: Sierra del Pino, mouth of southern canyon, Aug. 26,
1940, Johnston & Muller 746A (G). Nuevo Leon: Canyon de los Capulines,
above San Enrique, Hacienda San Jose de Raices, Aug. 6, 1935, Mueller 2378
(type, Gray Herb.) and 2379 (G).

The plants above described agree in gross habit and vegetative charac-
ters, in calyx, nutlets and pollen, and in their predominantly cleistogamic
flowering. Annotation on the specimens cited gives evidence of my uncer-
tainty and changing opinions concerning their identity. At one time or
another they have been questionably identified as L. obtusifolium, L.
calycosum, L. multiflorum, L. cobrense, and L. incisum, and even their
possible hybrids, as well. In grouping them in a proposed species I am
aware that they are relatively few in number and that they represent a
wide and erratic geographic distribution on either side of the United States—
Mexican boundary. The specimens come from scattered localities in south-
eastern Arizona, in western Texas and northeastern Mexico, all areas much
visited by botanists. I can only suggest that because the species is pre-
vailingly cleistogamic and accordingly deficient in colorful flowers, it has
not attracted the attention of collectors and so is poorly represented in
herbaria.

In general appearance L. confine closely simulates L. calycosum, a Mexi-
can species also developing cleistogamic flowers. It is, however, readily
separable from that species by its large spherical pollen, basally constricted
nutlets, and chasmogamic corollas with faucal appendages. The same char-
acters also eliminate L. cobrense and L. multiflorum, as does:also the pres-
ence of cleistogamy. Cleistogamy is known in Lithospermum only in L.
calycosum and in the group of species containing L. incisum, L. Parksii,
and L. mirabile. Our present plant obviously belongs in this latter group.

348 JOURNAL OF THE ARNOLD ARBORETUM [voL. xxxmI

It has the distinctive pollen of that group, and also similar chasmogamic
corollas. Its nutlets, furthermore, are very similar in appearance, size,
form, and structure to those of L. incisum and, in all except surface mark-
ius, to those of L. Parksii also. I believe it to be most closely related to
L. incisum. From that species it differs in its infrequent, much smaller
chasmogamic flowers, entire-margined corolla-lobes, erect fruiting pediceis,
elongate cymes, and erect, never diffusely branched stems.

33. Lithospermum afromontanum H. Weim. Bot. Notiser 1940: 65,
f. 7 (1940

A very distinct plant of central Africa formerly confused with the very
different L. officinale. From the latter it is easily distinguished by having
very different pollen, a more accrescent calyx, and stamens borne high in
the more elongate corolla-tube. The stems are very elongate, 5-10 dm.
long, and bear numerous sharply acute lanceolate leaves 4—8 cm. long and
7-30 mm. broad. The yellow corolla has a subcylindric tube 5—7 mm. long,
which usually surpasses the calyx by 1-2 mm. The spreading limb is about
8 mm. in diameter. The faucal appendages are well developed gibbose
invaginations which are minutely hairy and glanduliferous. The nectary is
a thickish collar. The style is 1.5—4 mm. long and terminated by two small
stigmas. The pollen is cylindric with rounded ends and straight paralleling
sides. It measures 20 by

34. Lithospermum ruderale Dougl. ex Lehm. Pug. 2: 28 (1830).
Lithospermum pilosum Nutt. Jour. Acad. Philad. 7: 43 (1834).
Batschia pilosa (Nutt.) G. Don, Gen. Syst. 4: 372 (1838
Lithospermum Torreyi Nutt. he Acad. Philad. 7: 44 (1834),
Batschia Torreyi (Nutt.) G. Don, Gen. Syst. 4: 372 (1838).
sara! opi ruderale var. Torreyi (Nutt.) Macbride, Contr. Gray Herb. 48:
55 (191
oo laxum Greene, Pittonia 3: 263 (1898)
Lithospermum lanceolatum Rydb. Mem. N. Y. Bot. Gard. 1: 333 (1900).
Lithospermum ruderale var. lanceolatum (Rydb.) Nelson, Bot. Gaz. 52: 272
1).

Lithospermum ruderale var. macrospermum Macbride, Contr. Gray Herb. 48:
55 (1916).

Widely distributed in northern portions of western United States and
adjacent Canada. A distinctive feature of the species is its cylindric thyr-
soid inflorescence. Unlike most species of the genus, there is no large
dominant cyme terminating the main stems. Indeed, in L. ruderale the ter-
minal cyme is usually less developed than the numerous small lateral cymes
that arise from the upper leaf-axils. The aggregation of these numerous
small cymes is elongate, very leafy and cylindric. The pale yellow corollas
have a cylindric tube not much longer than the calyx. The limb has ascend-
ing lobes. The throat bears more or less well defined congregations of glands
below each corolla-lobe, but has no intruding appendages. Scattered glands
are numerous in the tube above the level of the filament-attachments and

1952] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXIII = 349

further down, scanty along the principal veins to near the middle of the
tube. The nectary is a somewhat tumid obscurely lobed glabrous ring.
The style reaches up to the level of the stamens and is usually shorter
than the calyx-lobes. The two stigmas are terminal and juxtaposed. The
pollen (fig. 23) is ellipsoidal 25 > 18-20 yp, and in lateral profile has
rounded or weakly angulate sides. It is clearly broadest at the equator.
The pores are weak or obscure. The nutlets, 4-7 mm. long, are ovoid
or globose-ovoid, pointed at the apex, and strongly constricted just above
the broad base.

35. Lithospermum calycosum (Maebride) Johnston, Contr. Gray
Herb. 70: 30 (1924).
Lithospermum strictum var. calycosum Macbride, Contr. Gray Herb. 48: 56
(1916).
Lithospermum obtusifolium Johnston, Contr. Gray Herb. 70: 27 (1924).
Lithospermum Galeottii Brand, Fedde Repert. 28: 17 (1930).

A variable species ranging from northeastern Mexico south into the
mountains of Guatemala. One of its features is its capacity for cevelop-
ing cleistogamic flowers. Cleistogamy is known in the genus otherwise only
in L. incisum and its close relatives. The chasmogamic corollas of ZL.
calycosum vary widely in size, being largest on vigorous plants and espe-
cially in the spring of the year. Late flowers or those on plants in unfavor-
able habitats tend to be small and frequently cleistogamic. The yellow
corollas have a subcylindric tube 5-16 mm. long and 1.5—2.5 mm. thick.
When large the tube may be twice the length of the calyx but when small
it may scarcely surpass the calyx. The corolla-limb varies from 2 to 7 mm.
in diameter. Its lobes are spreading and 1—2 mm. long and are erose or
strongly crisped at the margin. The throat lacks invaginate intrusions.
It is distinctly glanduliferous with the glands tending to congregate below
each of the corolla-lobes. Scattered glands occur down in the tube to about
the level of the stamen-attachment. The nectary in the corolla consists of
10 minutely hairy swellings. The style reaches up to the stamens or just
beyond them and at times can be almost exserted from the tube. The
two stigmas are terminal. The pollen is ellipsoidal and measures 35-41
24-33 », and commonly much resembles that of L. Pringleit and L. inde-
corum. In lateral profile it is evidently broadest at the equator and the sides
are rounded or somewhat angulate. The pores may be either prominent or
obscure.

The corollas of the cleistogamic flowers are 1-3 mm. long. Such flowers
may be produced only at the ends of the cymes late in the growing season,
or occasionally may be the only flowers produced by the plant. The type
of L. obtusifolium appears to represent the latter condition. Since nutlets
produced by cleistogamic flowers are associated with styles only [—3 mm.
long and those from chasmogamic flowers with styles up to 16 mm. long,
the relative fertility of the two types of flowers is readily ascertainable.
The two appear to be equally fertile, but the chasmogamic ones are much
more numerous.

350 JOURNAL OF THE ARNOLD ARBORETUM _[VoL. xxxiIr

Plants from southern Mexico and Guatemala tend to be more robust
than the more northern plants and have coarser, more spreading hairs. The
name L. Galeottii is available for them if they prove worthy of recognition.

36. Lithospermum Pringlei Johnston, Contr. Gray Herb. 70: 22
(1924

Lithospermum Seleri Johnston, Contr. Gray Herb. 70: 28 (1924).

A species of central Mexico, probably most closely related to L. calycosum
and L. indecorum. Most of the specimens seen have a weak root which,
if not annual, is probably no more than biennial in duration. The yellow
corollas have a cylindric tube 5-9 mm. long and 1.5—2 mm. thick, which
surpasses the calyx by 1-3 mm. The limb is 3-7 mm. broad. The rounded
lobes frequently have a somewhat erose or crispulate margin and hence are
suggestive of those of L. calycosum. The throat bears 5 definite gibbose
invaginations which are densely glanduliferous on the inner side and at
times are minutely velvety on the summit. Stiped glands occur only on the
faucal appendages and just below them. The nectary in the tube is 5—10-
lobed and usually minutely villulose. The style reaches up into the corolla-
throat and bears two nearly terminal stigmas, left and right on its truncate
or convex sterile tip. The pollen (fig. 21) is ellipsoid, 33-37 25-27 u,
and has prominent pores. The pores seem to be seven in number.

37. Lithospermum indecorum, sp. nov.

Herba perennis 1-4 dm. alta; caulibus simplicibus vel laxe ramosis
basim versus 1—2.5 mm. crassis hispidulis vel villoso-hispidulis (pilis 0.5—
2.5 mm. longis saepe adpressis) et pilulis inconspicuis 0.1—0.3 mm. longis
saepe incurvatis obsitis, internodiis saepe 1-3 cm. longis; foliis basalibus
tempore florendi delapsis; foliis costatis sed saepissime enervatis, eis infra
medium caulis gestis oblanceolatis 1.5—4 cm. longis 4-10 mm. latis, eis supra
medium caulis gestis plus minusve oblongis vel lanceolatis 3 cm. longis et 4
mm. latis vel minoribus, supra viridibus vix abundante adpresseque hispid-
ulis (pilis gracilibus 1-4 mm. longis et pilulis 1-2 mm. longis praeditis) ,
subtus pallidioribus sparse adpresse hispidulis vel secus costam et marginem
adscendenter hispidulis; floribus inter folia apicem versus caulis et ramu-
lorum dispositis in inflorescentiam distinctam haud collectis; calyce adpresse
hispidulo, lobis lanceolatis tubo corollae aequilongis vel eum breviter super-
antibus maturitate 6-9 mm. longis; corolla flava extus sparse strigosa, limbo
2-3.5 mm. diametro, lobis rotundis 1-1.2 mm. latis, tubo subcylindraceo
4—5.5 mm. longo ad 2 mm. crasso, fauce appendiculis prominulis debiliter
invaginatis sparse glanduliferis praedito; antheris 0.8 mm. longis, fila-
mentis 3—3.5 mm. supra basim tubi corollae affixis; nectario tubi annulato
tumido lobulato, lobulis villulosis; stylo 2.5-4 mm. longis; stigmatis 2
parvis terminalibus vel paulo subterminalibus saepe compressis et divari-
catis; nuculis ovoideis ca. 3.5 mm. longis supra medium subconicis saepe
brunnescentibus et sparse punctatis.

MEXICO: Nuevo Leon: Canyon Los Capulines above San Enrique, Hacienda
San Jose de Raices, shaded situations on slopes, fl. cream-yellow, Mueller 2380

1952] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXIII 351

(type, Gray Herb.; Mo, FM). TAmauttpas: between Marcella and Hermosa,
burned-over area, fl. cream-yellow, Stanford, Lauber & Taylor 2637 and
2649 (G).

Probably most closely related to L. Pringlei, from which it is distin-
guished by smaller veinless or nearly veinless leaves and smaller corollas
that have weak faucal appendages and a tube not surpassing the calyx.

38. Lithospermum jimulcense, sp. nov.

Planta perennis indumento griseo vestita; caulibus erectis pluribus 10-15
cm, longis e rhizomate lignoso orientibus supra medium sparse breviter
ramosis pilis rectis adpressis pallidis 1-2.5 mm. longis obtectis; foliis
numerosissimis, infimis oblanceolatis 5-10 mm. longis 1—2.5 mm. latis
tempore florendi delapsis, ceteris (eis parte medionali caulis majoribus)
anguste oblongis vel lanceo-oblongis 2-4 mm. latis 1-5 mm. distantibus,
apice obtusis, basi abrupte contractis sessilibus, margine anguste revolutis,
in facie superiore pilis gracilibus hispidulo-villosis 1-2.5 mm. longis griseo-
vestitis, in facie inferiore tomentulosis (pilulis contortis 0.1-0.3 mm
longis) et adpresse hispidulo-villosis; floribus parvis inter foliis minoribus
supremis dispositis; calyce 3-4 mm. longo 0.3-1 mm. longe pedicellato,
lobis 0.6-1 mm. latis tubo corollae brevissime longioribus; corolla lutea,
extus supra medium dense adpresseque villulosa, tubo 3.5-4 mm. longo a
basi ca. 1 mm. crasso sursum gradatim ampliato summum ad apicem ali-
quantum constricto, limbo 3-4 mm. diametro, lobis rotundis 1.5 mm. latis
longisque, fauce glandulifero (glandulis dispersis) nullo modo invaginato-
appendiculato; antheris 1.3 mm. longis in tertiam partem superiorem tubi
positis apice mucronulatis; pollina late ellipsoidea 20 16 » a latere
viso rotunda poris uniseriatis obscuris donata; filamentis 2.5 mm. supra
basim tubi affixis; nectario tubi annulato lobulato, lobis inconspicue
minuteque viluloss: stylo 2-4 mm. longo stigmatibus 2 minutis terminali-
bus donato; suculis ignotis.

MEXICO: CoaHvuILa: summit of Mt. Jimulco, 3100 m., thick underbrush
ith oak, pine and juniper, fl. yellow, June 29, 1941, mora: Retherford &
Northeraft 100 (type, Gray Herb.; Mo).

A very distinct species which has a gross habit more suggestive of
Heliotropium than Lithospermum. The short subsimple stems are very
numerous and crowded and arise from a very well developed loosely
branched caudex growing in rock-crevices or among loose rocks. Among its
notable features are its very abundant crowded small elongate leaves and
its rather dense grayish indument of appressed slender hairs. The corolla
is also noteworthy, being not only unusually small, but also very densely
tomentulose-villulose outside. Until the fruit of the plant becomes avail-
able for study, the relations of L. jimulcense will remain questionable.
However, its closest relationship is probably with L. indecorum.

352 JOURNAL OF THE ARNOLD ARBORETUM _[VoL. XxxIII

39. Lithospermum oblongifolium Greenm. Proc. Am. Acad. 32: 300
(1897).

Lithospermum euryphyllum Brand, Fedde Repert. 28: 16 (1930).

This well-marked species probably has its closest affinities among the
large-flowered Mexican species with ellipsoid pollen, and particularly with
L. viride, L. guatemalense, L. strictum, and L. Muelleri. Among its more
distinctive features are its well-developed faucal appendages and its cylin-
dric or somewhat ovoid pollen. The plant is confined to the central plateau
of Mexico. It is a relatively coarse perennial with stems 3-10 dm. tall.
The leaves are large and evidently veined. The greenish yellow corollas
have a gradually expanded tube 15-30 mm. long and a small limb of
broad rounded lobes 5-10 mm. in diameter. The throat has evident in-
vaginate, emarginate, somewhat trapeziform appendages that are short-
hairy and glanduliferous. Inconspicuous, very scattered glands are also
present in the throat below the appendages and frequently also on the
vein below the filament-attachment. The nectary is an ill-defined slightly
swollen glabrous band. The style eventually becomes exserted 3-8 mm.
from the throat and bears its two stigmas terminally or subterminally. In
some plants the semicircular or transversely oblong stigmas arise from
the tip of the style, but in others they are decidedly below and lateral to
the obscurely lobed prolonged sterile tip of the style. The pollen, 25-27
s< 16-20 p, is cylindric with rounded ends or somewhat ovoid. In lateral
outline the sides of the grains are usually straight and accordingly may be
either slightly convergent or parallel. The pollen is not ellipsoidal or
subglobose nor distinctly broadest at the middle, as is that of closely
related species. Nevertheless, as in the latter, its pores are medial. They
are apparently six in number and are evident only in collapsed grains.

40. Lithospermum strictum Lehm. Asperif. 2: 303 (Nov.—Dec. 1818).

Anchusa tuberosa HBK. Nov. Gen. et Sp. 3: 92 (Sept. 1818). Not Lithosper-

mum tuberosum Rugel (1846).
Heliotropium lithospermoides R. & S. Syst. 4: 737 (1819).
Heliotropium mexicanum Sessé & Moc. Pl. N. Hisp. 20 (1888); Johnston, Jour.
Arnold Arb, 30: 109 (1949).
Lithospermum rosmarinifolium Sessé
Boiss. (1879).
Lithospermum a
(1775).

& Moc. Pl. N. Hisp. 20 (1888), not

ngustifolium Sessé & Moc. Fl. Mex. 32 (1893), not Forsk.

nizable species of central Mexico. From a slender tap-
root, which has a conspicuous fusiform swelling just below the surface of
the soil, the plant produces one to many slender stems 2—4 dm. tall. The
d terminate in elongate scorpioid

A readily recog

zygomorphic limb 3-6 mm. in diameter. The upper surface of the obovate
or elliptic ascending lobes is microscopically velvety-puberulent and fre-
quently also coarsely strigose. The slender corolla-tube is very gradually

1952] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXIII 353

ampliate for most of its length but at the very summit is perceptibly con-
stricted. The throat bears five evident faucal appendages. These are weakly
invaginate convexities bearing a prominent arcuate ridge of tissue across
their upper end. Like the corolla-tube below them, they are glabrous and
lustrous. A cluster of glands is usually present at the base of the corolla-
lobes just above each arcuate ridge, but below the ridges glands are very
few and inconspicuous. The nectary in the corolla-tube consists of a 10-
lobed very narrow flap which is very minutely, scantily, and inconspicu-
ously villulose. The style is variable in length, in some plants reaching
only to the middle of the corolla-tube but in others as high as the top of
the anthers. The two stigmas are terminal, strictly ascending, and at times
appear to be joined at the base. The lustrous, very smooth nutlets are dis-
tinctive. They have a very sharp and prominent keel which extends up the
venter over the apex and partially down the back of the nutlet-body. Be-
low the middle of the nutlets the dorsum is noticeably flattened or very
low-convex. The pollen is globose or globose-ellipsoidal, as long as broad
or slightly longer than broad. It measures 33-37 33-35 yp. In lateral
outline the sides are rounded or angulate. The grain is clearly broadest at
the equator. The pores (apparently 8) are very obscure.

41. Lithospermum Muelleri Johnston, Jour. Arnold Arb. 16: 187
(1935).

A very distinct species known only from the mountains of northeastern
Mexico. It seems to be most closely related to L. strictum. The lower sur-
face of the leaves in L. Muelleri bears some appressed hairs along the mid-
rib, but otherwise the surface is glabrous or practically so. This condition
is very uncommon in the genus. The elongating scorpioid cymes, as in
L. strictum, have relatively small bracts and are produced, singly or gemi-
nate, terminal on the simple very leafy stems. The greenish yellow corolla
has an elongate subcylindric tube 15-20 mm. long which is abruptly con-
stricted at the top and bottom. The small limb, 2—3 mm. broad, is com-
posed of broad, short, rounded ascending lobes. The throat has five small
but eledefned faadal appendages. Each consists of a small low-convex
area delimited above by an arcuate or somewhat trapeziform intrusion
formed partially by an invagination and partially by a thickened ridge of
epidermal cells. They are glabrous and somewhat lustrous below the in-
trusion. On the upper side of the intrusion stipitate glands are numerous.
Below the appendages glands are very scattered and tend to be most
numerous along the vein just below the filament-attachment. ‘The style
eventually becomes exserted from the corolla-tube, commonly by 1-3 mm.
The two stigmas are terminal, strict, closely juxtaposed, and frequently
more or less confluent. The nectary in the corolla-tube consists of a thin
ridge bearing minute tufts of hairs. The pollen (fig. 18), 25-30 » broad, is
globose or very slightly longer than broad. In lateral profile the sides are
rounded or somewhat angled. The grain is broadest at the equator. The
pores, apparently 8 in number, are very slightly evident or are obscure.
The fruit has not been seen.

354 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxx

42. Lithospermum Hancockianum Oliver in Hooker Icones 25: t.
2457 (1895); Hand.-Mazz. Naturbilder aus S.W. China 116, t. 2
(1927).

Lithodora Hancockiana (Oliver) Hand.-Mazz. Sym. Sin. 7: 818 (1936).
Arnebia Hancockiana (Oliver) Johnston, Jour. Arnold Arb, 18: 21 (1937).
Lithospermum Mairei Lévl. Fedde Repert. 12: 286 (1913).

A very distinct species of China, known only from limestone ledges in
eastern Yunnan. The functional leaves are borne clustered at the ends of
the trailing branches of a very loose shrubby caudex and directly above a
skirt-like mass of dead reflexed leaves persistent from previous seasons.
They are very elongate and are covered with lustrous white silky appressed
hairs beneath. The inflorescence, at first glomerate, later becomes some-
what racemose but is always shorter than the leaves in the cluster from
which it arises. The salverform corolla is pinkish or bluish to purplish-
red, but usually becomes yellowish in drying. The tube, 2-3 mm. thick, is
cylindric and 18-25 mm. long. The spreading limb is 14—20 (—‘25”) mm.
broad. The throat bears numerous scattered glands above the level of the
filament-attachment. There are no faucal appendages. The anthers are
borne either 2—3 mm. below the summit of the tube or at the summit and
partially exserted from it. The nectary in the tube is a glabrous flange.
The style is exserted 2-5 mm. from the tube. The two stigmas are terminal
or subterminal. The pollen (fig. 31) is subglobose (26-33 » in diameter) to
ellipsoidal (26-30 * 23-26 ,»), as long as broad to evidently longer than
broad. In lateral profile the sides are rounded or angled. The grains are
clearly broadest at the equator. The pores (7, or less commonly 6) may
be evident or obscure. The nutlets (only submature seen) are white,
smooth, bony, and ovoid.

In its non-yellow corolla devoid of faucal appendages, and in its par-
tially exserted stamens, the plant suggests the Asiatic members of the
genus formerly referred to Arnebia. It differs, however, in its smooth white
nutlets and ellipsoid or globose pollen. The plant is one in which heterostyly
might be expected. Evidence of it, however, has not been found in the
five collections dissected. All the plants studied have elongate, shortly
exserted styles, and stamens borne high in the corolla-tube. To be sure
there are slight differences in the level at which the anthers are carried.
On some plants the anthers are borne just low enough to be included in
the corolla-tube, while in others they are borne just enough higher so that
they are half exserted from the tube. Such a small difference in level of
stamen-attachment is associated with heterostyly in L. densiflorum, but in
L. Hancockianum there appears to be no correlation between stamen-
position and a particular length of style or size of pollen.

43. Lithospermum guatemalense Donn. Sm. Bot. Gaz. 27: 436 (1899).

A coarse perennial native in the mountains of northern Guatemala and
adjacent Mexico. The yellow corollas have a gradually ampliate tube

1952] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXIII 355

which is 18-25 mm. long and at least twice the length of the calyx. The
rounded lobes are 2—3 mm. long and ascending. The throat bears slightly
convex, densely glanduliferous areas below each corolla-lobe. There are no
strong invaginations. Glands are scattered over the inner surface of the
tube above the level of the filament-attachments. The style becomes even-
tually exserted, usually as much as 2-3 mm. The two stigmas are terminal.
The nectary in the tube consists of 5-10 sparsely villulose very minute
swellings. The pollen is ellipsoidal and measures 28 & 25 yp. In lateral
profile it has slightly angled sides and is evidently broadest at the equator.
The pores, 7-8 in number, may be either obscure or evident. The nutlets
are ovoid, ca. 4 mm. long, and usually grayish or tawny rather than white.
They are usually very conspicuously sulcate and punctate adjacent to the
ventral keel and generally punctate on the back also.

44. Lithospermum viride Greene, Bot. Gaz. 6: 158 (1881).
Lithospermum Palmeri Wats. Proc. Am. Acad. 18: 122 (1883).

A species of northeastern Mexico and adjoining United States (Arizona
to Texas). Its closest affinities appear to be with ZL. Macbridei of Peru.
The tubular corollas are yellowish or greenish and have a relatively small
limb of divergent or recurved elliptic to broadly ovate lobes. The tube,
15-35 mm. long, is cylindric with a constriction at the summit. Inside, the
throat is devoid of swellings or invaginations, but is abundantly glandu-
liferous. The nectary is represented by 5—10 very minute, usually villulose
swellings. The pollen is subglobose to ellipsoidal and as long or very
slightly longer than broad. It measures 16-23 % 16-22 py. In lateral out-
line it is evidently broadest at the equator and its sides are angulate. There
are 7 or 8 very obscure pores about the equator. The style is tardily ex-
serted 1-5 mm. from the tube. It bears 2 terminal stigmas. The mature
fruiting calyx becomes 10-20 mm. long. Its very narrow and elongate
lobes greatly surpass the nutlets.

45. Lithospermum Macbridei Johnston, Contr. Gray Herb. 78: 8
(1927);

This Peruvian plant seems to be most closely related to L. viride of Mex-
ico, but is readily distinguished by having crowded, very numerous, much
smaller, narrow veinless leaves as well as a neat pallid strigose indument.
The greenish yellow corolla has a conspicuous subcylindric tube 10-14 mm.
long and a small limb of diverging or recurving lobes. The throat has no
intrusions. It is, however, densely glanduliferous with the glands in greatest
concentration below each of the corolla-lobes. The nectary is represented
by 5-10 minute villulose swellings. The style is eventually exserted 2-5 mm.
from the throat and is terminated by 2 stigmas. The pollen is subglobose
or nearly spherical, 16-20 » in diameter. In lateral profile it is as broad
as long, broadest at the equator, and with rounded or slightly angulate
sides. There are 8 obscure pores on the equator. The mature fruiting calyx
is 5-8 mm. long, and at most only twice as long as the nutlets.

356 JOURNAL OF THE ARNOLD ARBORETUM _[VvoL. xxxmI

46. Lithospermum discolor Mart. & Gal. Bull. Acad. Brux. 11: 337
(1844).
Lithospermum discolor a candicans Kuntze Rev. Gen. 2: 439 (1891).
Lithospermum obtusiflorum Sessé & Moc. Fl. Mex. 32 (1893); Johnston, Jour.
Arnold Arb. 30: 109 (1949).
Lithospermum chersinum Macbride, Contr. Gray Herb. 49: 22 (1917).
Lithospermum hypoleucum Johnston, Contr. Gray Herb. 70: 23 (1924).

A plant with erect, subsimple to loosely branched, usually hispid stems
5-15 dm. tall, frequently becoming suffrutescent. It is native to central
and western Mexico. The evidently veined, usually lanceolate leaves are
pale green above and white from a dense appressed indument beneath.
The flowers are strongly heterostylic. The white corolla is 1-2 cm. long
and has a limb 8-15 mm broad. The tube, slightly to conspicuously longer
than the calyx, has the middle three-fifths of its length appressed villose
inside. The throat bears abundant scattered glands but is devoid of ap-
pendages. The nectary is a tumid obscurely lobed glabrous ring. The stamens
are borne below the middle of the corolla-tube or just below its summit.
The style is either a quarter to a third the length of the tube or nearly as
long as the tube. The two stigmas are terminal. The elongate pollen (fig.
13) is much constricted at the middle, most strongly so in the long-styled
flowers. In the latter it measures 22-28 * 10-15 pw. In the short-styled
flowers it is larger, 33-39 % 18-20 ». There are 7-9 pores. The smooth
white nutlets are elliptic-ovoid and are rounded and unkeeled ventrally.

47. Lithospermum matamorense A. DC. Prodr. 10: 76 (1846).
Lithospermum prostratum Buckley, Proc. Acad. Philad. 1861: 462 (1861).

A species known only from northeastern Mexico and adjacent Texas,
which is probably most closely related to L. Berlandieri. It is an annual,
with a juicy taproot and several ascending loosely branched hispidulous
stems. The basal leaves are largest. The corolla has a spreading limb
4-6.5 mm. broad. Outside it is very minutely and sparingly strigulose.
The tube, 1—-1.5 mm. long, is about half the length of the calyx. The throat
is provided with trapeziform invaginate appendages which are velvety on
top and glanduliferous on the sides. Some glands occur also on the throat
below the appendages. The minute anthers are borne on filaments attached
at the middle of the corolla-tube. The nectary is a tumid ring, entire or
5—10-lobulate, which is glabrous or very inconspicuously puberulent. The
pollen is constricted at the middle and measures 16-20 10-13 ». The
nutlets of the species are distinctive. They are 2.5-3 mm. long, usually
brownish and conspicuously punctate. The ventral keel is broad and
rounded and continues up over the apex and down onto the dorsum. The
back of the nutlet is not only pitted but frequently also somewhat tuber-
culate or tumulose, As the.result of a lineate constriction above its base
the nutlet has a more or less well defined basal collar. The attachment-
scar is concave. The funicular canal is evident, but the dorsal traces usually
evident on the attachment-scar of the nutlets in other species are very

1952] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXIII 357

vague or absent in L. matamorense. The gynobase is obscurely pyramidal,
in fact almost plane.

48. Lithospermum Nelsonii Greenm. Proc. Am, Acad. 40: 31 (1904).

A species local to northeastern Mexico. The corolla is large, white, and
salverform. The limb is 10-14 mm. broad. The tube is 12-15 mm. long,
commonly 2-3 times as long as the calyx, and though abruptly expanding
at the very base it is cylindric for most of its length. The throat bears small
but evident faucal appendages which are invaginate, trapeziform, and
glanduliferous. Glands occur also in the throat below the appendages, and
others are present at the base of the filaments. The nectary is a glabrous,
obscurely lobed flange. The pollen (fig. 26) is medially constricted and
measures 15-23 & 10-13 p. It is similar in size and appearance to that of
L. matamorense but larger than that of L. Berlandieri. The white smooth
nutlets are 2.5-3 m. long, and as in L. Berlandieri, are constricted just
above the base. The attachment-scar is concave.

49. Lithospermum Berlandieri, sp. nov.

Herba perennis 2—5 dm. alta; caulibus pluribus erectis hispidulis (pilulis
minutis 0.2—0.3 mm. longis saepe incurvatis et pilis majoribus 0.7--1.2 mm.
longis saepe divaricatis praeditis) supra medium sparse adscendenterque
ramosis, basim versus 2—3 mm. crassis; foliis basalibus et caulinis inferiori-
bus tempore florendi emarcidis; foliis caulinis numerosis saepissime patenti-
bus oblanceolatis (majoribus 5-7 cm. longis et 10-15 mm. latis) supra
medium latioribus deinde deorsum gradatim attenuatis, pilulis 0.2-0.3 mm.
longis saepe adpressis et pilis majoribus 0.7—1.2 mm. longis adscendentibus
vel adpressis vestitis, apice acutis vel saepissime obtusis, subtus pallidiori-
bus saepe venis lateralibus pauce donatis, supra basibus pallidis discoideis
pilorum non rariter ornatis; cymis caules et ramulos terminantibus, maturi-
tate laxe racemosis 5—10 cm. longis bracteis divaricatis oblongis vel lanceo-
latis ad 3 cm. longis ornatis; corolla alba extus parce strigosa, tubo sub-
cylindraceo 2.5—3.5 mm. longo ca. 2 mm. crasso, limbo ad 6 mm. diametro
rotato, lobis rotundatis 2 mm. longis et latis, fauce appendiculis intrusis
gibbosis ornato, appendiculis sae ee apice sparse velutinis latere
abundanter slanduliferss: antheris 0.7-0.8 mm. longis in tertiam partem
superiorem tubi corollae gestis; ete ca, 2 mm. supra basim tubi
affixis basi imo sparse glanduliferis; pollina elongata medie constricta
13 7-10 p»; nectario tubi corollae glabro 10-lobulato; stylo altitudinem
antherarum attingenti 2—2.5 mm. longo e tubo corollae nullo modo exserto;
stigmatibus 2 minutis apice emarginato styli positis; nuculis albis laevibus
non rariter sparse punctatis ovoideis 3—3.5 mm. longis supra basim laeviter
constrictis, cicatrice basali concava; gynobasi latissime pyramidal.

MEXICO: Tamauttpas: oak forests on Jaumave road about 13 mi. southwest
of Ciudad Victoria, 1000 m. alt., abundant, fl. white, May 13, 1949, R. McVaugh
10517 (tyPE, Gray Herb.) ; pié la cuesta de Victoria a Tula, Nov. 1830, Berlandier
s.n. (G); Jaumave, 1932, Rozynski 567 (Chicago); mountains south of Victoria,

358 JOURNAL OF THE ARNOLD ARBORETUM _[vot. xxxu1

O m., scattered, March 1925, Runyon 747 (US); foot of mountains near
Victoria, 400 m., April 1926, Runyon 921 (US).

Closely related to L. Nelsonii, from which it differs in its very much
smaller corollas with proportionately shorter tube and narrower limb. The
present species is known only from the mountains southwest of Victoria,
Tamaulipas, whereas L. Nelsonii is known only from the mountains west
and south of Monterrey, Nuevo Leon.

50. Lithospermum papillosum Thunberg, Prodr. Pl. Cap. 34 (1794);
Thunberg in Schrad. Neues Jour. Bot. 1°: 44 (1806); Lehm. Asperif.
2: 329 (1818); Wright, Fl. Cap. 47: 21 (1904).
? Lithospermum papillosum 8 ambiguum DC. Prodr. 10: 74 (1846).

A well-marked species endemic to South Africa. It has numerous ascend-
ing lanceolate or lance-oblong leaves crowded along erect stems 2—4 dm. tall.
In general appearance it much resembles the American Heliotropium terna-
tum Vahl and its close allies. Especially distinctive of L. papillosum is the
generous development of minute obese hairs on the upper face of the
corolla-lobes, giving the latter a granular or somewhat velutinous appear-
ance. The corolla-tube, 2-3 mm. long, may be as long as the calyx or be
surpassed by it. The faucal appendages are densely glanduliferous, broad,
low-convex invaginations. The throat is usually abundantly glanduliferous
below the appendages and sparsely so at the base of the filaments. The
nectary is a well-developed thickish collar. The style, 1-2.5 mm. long,
bears 2 sessile terminal stigmas, and in some specimens appears to be spar-
ingly and very minutely hairy and glanduliferous below the middle. The
pollen is short-cylindric with rounded ends and measures 20-25 & 13-16 p.
No fruit has been seen. The nutlets, however, have been repeatedly de-
scribed as rugose.

51. Lithospermum diversifolium DC. Prodr. 10: 77 (1846); Wright,
Fl. Cap. 4°: 24 (1904).

A plant of South Africa that somewhat resembles L. officinale and was
formerly confused with it. From L. officinale our plant is readily distin-
guished by the elevated position of the anthers in the corolla-tube, the
broader and petiolate lower cauline leaves, and the very different pollen.
The tube of the small white corolla is subcylindric, ca. 2.5 mm. long, and
about twice the length of the calyx. The faucal appendages are prominent
invaginate gibbosities bearing glands and obese hairs. The throat below
the appendages is densely glanduliferous. Other glands are found at the
base of the filaments. The nectary is a 5—10-lobulate collar. The style is
1.5—2 mm. long and bears 2 sessile terminal stigmas. The pollen is short-
cylindric with rounded ends and measures 20 X 13 ». The pores distributed
about the equator are very obscure.

52. Lithospermum cinereum DC. Prodr. 10: 73 (1846); Wright, FI.
Cap. 4°: 23 (1904).
Lithospermum inornatum DC. Prodr. 10: 73 (1846).

1952] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXIII = 359

A well-marked species from South Africa. Among its notable features are
its smooth closely appressed grayish strigose indument, small thickish vein-
less leaves, very small flowers, and tumulose nutlets. The white corollas are
2.5-3 mm. long. The tube is slightly ampliate, 1.5-2 mm. long, which is
shorter than the calyx. The corolla-lobes, 0.5-1 mm. long, are ascending,
oblong, and evidently veined. The throat bears 5 slightly invaginate
rounded convex areas which are velvety and densely glanduliferous. A few
glands are present on the vein directly below the filament-attachment. The
nectary is 5—10-lobulate. The style is 1-1.5 mm. long and bears 2 terminal
stigmas. The pollen measures 16-18 & 11-14 » and varies from distinctly
ellipsoidal with rounded sides to cylindric-ellipsoidal with nearly straight
sides. The pores are borne at the equator and are very obscure. The
nutlets, 2.5-3 mm. long, are half or two thirds the length of the fruiting
calyx. Dorsally they have a broad low rounded keel, and towards the sides
below the middle are pitted and sparingly but distinctly tumulose.

53. Lithospermum scabrum Thunberg, Prodr. Pl. Cap. 34 (1794);
Thunb. in Schrad. Neues Jour. Bot. 1°: 44 (1806); Lehm. Asperif.
2: 309 (1818); Wright, Fl. Cap. 47: 22 (1904).
Lithospermum hirsutum E. Meyer ex DC. . 10: 77 (1846).
Lithospermum affine DC Prodr. 10: 78 (18

A species of South Africa usually a6 recognized because of its
spreading, short, villose-hispid indument. The white corolla is 8-9 mm.
long. Its tube, 5-6 mm. long, is twice as long as the calyx. For most of its
length, 4—5 mm., it is cylindric, but towards its summit it becomes ampliate.
The corolla-limb becomes 5—6.5 mm. broad. Its ascending lobes are rounded
and ca. 2 mm. broad. The throat bears 5 weak gibbosities which are some-
what velvety at the summit and are glanduliferous on the sides. A few
glands may also be present at the base of the filaments. The nectary is
10-lobulate. The style reaches at least to the apex of the stamens and
may become even slightly exserted from the tube. It has a sterile apex
that may be obscurely and weakly prolonged beyond the attachment points
of the two stigmas. The pollen, 16-18 X 14 uy, is ellipsoidal. It is broadest
at the equator and in lateral profile has rounded sides. The pores are
very obscure and borne about the equator or very slightly below it. The
small nutlets are ovoid, white, and very smooth.

54. Lithospermum peruvianum A. DC. Prodr. 10: 77 (1846).
Lithospermum aequatoriale Brand, Fedde Repert. 28: 14 (1930).

A plant of northern Peru and southern Ecuador which forms prostrate
mats. Stems numerous, very slender, very elongate and hispidulous. The
leaves are very numerous and small, 15-25 mm. long. The small flowers
are borne along the terminal portion of the elongate leafy stems and not
in a sharply defined inflorescence. The small white corolla has a stout tube
1.5-2 mm. long, which barely if at all surpasses the calyx. The limb is
3-4 mm. broad. Its broad rounded lobes nearly equal the length of the

360 JOURNAL OF THE ARNOLD ARBORETUM _ [vot. xxxm

corolla-tube. There are 5 circular glanduliferous and minutely hairy areas
in the throat. These are convex and only very weakly invaginate. The
nectary is villulose. The style is 1-2 mm. long and bears 2 juxtaposed small
terminal stigmas. The pollen varies in form from cylindric to ellipsoidal, in
lateral profile having the sides more or less parallel or slightly angulate
and the body broadest at the equator. It measures 13-20 % 10-14 pe
The pores are usually obscure. The nutlet tends to develop a slightly
swollen rim at the base. .

55. Lithospermum calcicola Robinson, Proc. Am. Acad. 27: 182
(1892).
Lithospermum Conzattii Greenm. Bull. Field Mus., Bot. 2: 339 (1912).
Lithospermum calcicola var. Conzattii (Greenm.) Johnston, Contr. Gray Herb.
70: 27 (1924).

Lithospermum madrense Brand, Fedde Repert. 28: 14 (1930).

Lithospermum hoyasense Brand, Fedde Repert. 28: 15 (1930).

A species of Mexico, ranging from Coahuila and Nuevo Leon south to
Oaxaca. The plant has a distinctive indument that aids in its ready recog-
nition. The leaves on both surfaces bear stiff straight appressed hairs 1—1.5
mm. long. These hairs are loosely spaced (commonly 0.5—1 mm. apart)
and, especially on the upper surface, arise from well-developed, evident,
usually discoid mineralized bases. The corolla and also the style-length are
surprisingly variable as to size and proportions, even among plants from a
single locality. This variation may be correlated with the season and
general vigor of the plant. The tube of the white corollas is subcylindric,
3—7 mm. long, and may equal the calyx in length or surpass it. The limb
is 4—7 mm. broad. The throat bears a congregation of glands below the
base of each corolla-lobe and also in the tube above the anthers. There are
no definite faucal invaginations. The style, 3-11 mm. long, commonly
just surpasses the anthers but is not quite exserted from the throat. In
some plants, however, it becomes tardily exserted and protrudes as much
as 1-4 mm. In other plants it reaches only to the base of the anthers. Fur-
thermore, some very mature plants have nutlets associated with a style
only 1 mm. in length. This latter condition suggests that cleistogamy may
be present. Though searched for, no cleistogamic flowers have been recog-
nized in the species. The two stigmas are either juxtaposed and terminal or
are slightly subterminal and separated by the sterile tissue of the rounded
tip of the style. The latter condition seems to prevail in plants from
Puebla and Oaxaca. The nectary in the corolla-tube is a glabrous flange.
The pollen is ellipsoidal or, more commonly, slightly but perceptibly
broader at one end (fig. 19) and hence somewhat ovoid. The pores are
weak or obscure and usually borne at a slight distance below the middle
of the grain. Between plants, the grains of L. calcicola show more variation
in size than is usual in the genus. They measure 16-25 & 13-16 p.

56. Lithospermum mediale Johnston, Contr. Gray Herb. 70: 28 (1924).

Lithospermum discolor y subviride Kuntze, Rev. Gen. 2: 439 (1891).
Lithospermum colombianum Brand, Fedde Repert. 28: 16 (1930).

1952] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXIII 361

This species is known only from Guatemala, Colombia, and Venezuela.
The white corolla has a subcylindric tube 5-8 mm. long and 2—2.5 mm.
thick, which surpasses the calyx 1-3 mm, The spreading limb is 4-8 mm.
broad. The throat has well-developed invaginate gibbose appendages which
are somewhat velutinous at the summit and densely glanduliferous on the
sides. Glands are also present, not only on and just beneath the faucal
appendages, but also along the vein just beneath the attachment of the
filaments. The style reaches to the throat of the corolla. The two stigmas
are clearly subterminal, being borne below the prolonged bilobed sterile
tip of the style. The nectary is more or less lobulate and villulose. The
pollen (fig. 22) is ellipsoidal and measures 23-27 & 18-20 up. The pores
are very obscure. In lateral profile the sides are rounded or somewhat an-
gulate. In the South American plants the pollen is clearly broadest at the
equator, but not so in all the Guatemalan material. In the latter the
grains sometimes become nearly straight-sided. The fruiting calyx in L.
mediale is usually weakly accrescent. Its lobes are usually only about
twice as long as the nutlets.

57. Lithospermum sordidum Brand, Fedde Repert. 28: 15 (1930).

Closely related to L. distichum, but distinguishable by its short-lived
stout fusiform root and the broad, usually clustered basal leaves, the more
or less evidently spreading indument, and the subsimple erect stems. It is
known only from eastern Mexico (Nuevo Leon, Hidalgo and Puebla),
where it appears to frequent lower altitudes and drier situations than its
relative. In technical characters it agrees with L. distichum.

58. Lithospermum distichum Ortega, Hort. Matr. Dec. 1: & C2797):

Batschia disticha G. Don, Gen. Syst. 4: 326 (1838).

Myosotis grandiflora HBK. Nov. Gen. et Sp. 3: 90, t. 199 (1818).

Heliotropium scorpioides Willd. ex Lehm. Nov. Act. Acad. Caes. Leop. Nat.
Cur. 9: 140 (1818), nomen; R. & S. Syst. 4: 737 (1819), not HBK. (1818).

Myosotis foliosa Lehm. Asperif. 1: 99 (1818).

Lithospermum spathulatum Mart. & Gal. Bull. Acad. Brux. 11: 337 (1844).

Lithospermum laevigatum Sessé & Moc. Fl. Mex. 32 (1893); Johnston, Jour.
Arnold Arb. 30: 109 (1949).

Anchusa mexicana Sessé & Moc. Pl. N. Hisp. 21 (1888); Johnston, Jour.
Arnold Arb. 30: 109 (1949).

Lithospermum approximatum Brand, Fedde Repert. 28: 15 (1930).

A perennial with a strong woody taproot and, usually, a shallowly buried,
loosely branched caudex. A plant of the mountains of Mexico and Guate-
mala. The stems are ascending, usually branched at the base, and com-
monly only 1—2 dm. long. The herbage bears minute appressed hairs and
is smooth in appearance. The white corollas have well-developed gibbose
invaginations in the throat. They are evidently puberulent and are glandu-
liferous on the inner side. Glands are most abundant on the faucal ap-
pendages. A few, however, are usually present at the base of the filaments.
The nectary is a thickish lobed collar. The two stigmas are subterminal

362 JOURNAL OF THE ARNOLD ARBORETUM (vot. xxximt

and borne laterally just below the sterile bilobulate tip of the style. The
pollen (fig. 24) is cylindric with rounded ends. The pores are equatorial.
The grains measure 16—22 & 10-14 ». The plant is variable in corolla-size
and in the relative length of the corolla-tube.

59. Lithospermum Gayanum (Wedd.) Johnston, Contr. Gray Herb.
78: 10 (1927).
Eritrichium Gayanum Weddell, Chor. Andina 2: 88 (1859).
Lithospermum andinum Krause, Engler’s Jahrb. 37: 636 (1906).
Lithospermum Shepardae Johnston, Contr. Gray Herb. 78: 10 (1927).

An Andean species ranging at high altitudes from central Peru south
into northern Bolivia. It is most closely related to the Mexican L. dis-
tichum, but differs in its smaller stature and in details of floral structure.
The faucal appendages are merely puberulent swellings and are not in-
vaginate. Glands are few in the throat and none is present at the base of
the filaments. The pollen is short-cylindric with rounded ends and meas-
ures 16-20 10-14 p. The style is terminated by the two stigmas. It
does not have the bilobate sterile tip developed by the styles of L. distichum
and L. sordidum.

1952] JOHNSTON, STUDIES IN THE BORAGINACEAE, XXIII] = 363

EXPLANATION OF PLATES

PLATE I

Pollen of heterostylic species of Lithospermum. In each pair the larger grain
(left) from short-styled flower, the smaller (right) from long-styled flower. The
arrow indicates the position of ow of pores. Species = Asiatic

Fic. 2. L. densiflorum. Fic. 3. L. euchromon. Fic. 4 L. fimbriatum. Fic. 5.
L. guttatum. Fic, 6. L Ca. Wie, 7. de hispidissimum.

PLATE II

Pollen of heterostylic species of Lithospermum. In each pair the larger grain
(left) from short-styled flower, the smaller (right) from long-styled flower. The
arrow indicates the position of a row of pores. First species Asiatic, the others
American.

Fic. 8. L. Tournefortii. Fic. 9. L. californicum. Fic. 10. L. tubuliflorum. Fic.
11. anescens, Fic. 12. L. caroliniense. Fic. 13. L. discolor. Fis. 14. L.
multiflorum. Fic. 15. L. cobrense. Fic. 16. L. obovatum,

PLATE III

Pollen of representative non-heterostylic species of Lithospermum (American
species, figs. 17-26; Eurasian, figs. 27-32). The arrow indicates the position of
the row of pores.

Fic. 17. LZ. elena Fic. 18. L. a Fic. 19. L. calcicola. Fic. 20.
L. tuberosum. Fic. 21. L. Pringlei. Fic. L. mediale. Fic. 23. L. ruderale.
Fic. 24... cron Fie: 25... Hien Fic. 26. L. Nelsonii. Fic. 27.
L. decumbens, Fic. 28. L. tetrastigma. Fic. 29. L. detonsum. Fic. 30. L. tschim-
ganicum. Fic. 31. L. Hancockianum. Fic. 32. L. officinale.

AARNOLD ARBORETUM,
HARVARD UNIVERSITY

Jour. ARNOLD Ars. VoL. XXXIII PraTeE I

+
Nn

10-20 30 & 6o 70 |
J

0 40 50
MICRON SCALE

JOHNSTON, POLLEN oF LITHOSPERMUM

Jour. ARNOLD Ars. VoL. XXXIII PLATE II

0 Op
Q0 OO 00

10 20 #30 40 «50 60 +7 ~~ ‘4|
MICRON SCALE |

JOHNSTON, POLLEN oF LITHOSPERMUM

Journ. ARNOLD Ars. VoL. XXXIII

Pate III
7 8 19 20 rr
22 23 24 aa 26
>
* +
. ° > ° (|
28
re 29
30 31 32

10 20 #30 +40 5o 60 7 ‘|
MICRON SCALE ]

JOHNSTON, POLLEN OF LITHOSPERMUM

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XIII 367

STUDIES OF PACIFIC ISLAND PLANTS, XIII
NOTES ON FIJIAN EUPHORBIACEAE

A. C. SMITH

THE FAMILY EUPHORBIACEAE is taxonomically one of the most difficult
groups of plants, and studies of it should preferably be undertaken only
by a specialist with many years of experience in the group. However,
the necessity of applying names to the material I collected in Fiji in 19474
has led me to study the Fijian Euphorbiaceae, the results of this study
being incorporated in the present paper. Here are mentioned only the
new or unusual species of my collection, except in the genera Antidesma,
Macaranga, and Acalypha, which could be clarified only by examination
of many Fijian specimens. For these genera keys are here provided and
material is cited from the following herbaria: Arnold Arboretum (A);
Bernice P. Bishop Museum (Bish); British Museum (BM); Gray Her-
barium (GH); Royal Botanic Gardens, Kew (K); New York Botanical
Garden (NY); and U. S. National Herbarium (US). The directors and
curators of these institutions have kindly permitted the study of their
material. Genera are discussed in the order of Pax & Hoffmann’s treatment
in Nat. Pflanzenfam. ed. 2. 19c (1931).

ANTIDESMA L.

The interesting genus Antidesma was singularly overlooked by the earlier
plant-collectors in Fiji; except for an Exploring Expedition specimen (the
type of A. pacificum) and a single collection made by Horne, the genus
was scarcely recorded from Fiji until Gillespie’s trip in 1927. However,
Antidesma is a not infrequent component of the Fijian hill-forest. Gillespie
indicated his several collections as a new species, A. insulare; examination
of these collections and those subsequently gathered demonstrates that
this species has been too broadly interpreted in herbaria. In the present
treatment I recognize five species, of which three are described as new.

KEY TO THE SPECIES

Calyx of 2 flowers rotate, deeply 4- or 5-lobed, the lobes lanceolate-deltoid,
puberulent on both sides; ovary and fruit asymmetrical, the stigmas lateral;
inflorescence-branches and pedicels persistently puberulent; leaf-blades
eerie rounded ee subcordate at base, faintly pilose on costa or in

mils of nerves beneath. «070625 ee ee eae . pacificum.

Git. cupuliform, truncate ‘at apex or inconspicuously lobed, usually glabrous

xcept on margin; ovary and fruit symmetrical, the stigmas terminal;

* Under the auspices of the Arnold Arboretum of Harvard University and the John
Simon Guggenheim Memorial Foundation, with the aid of grants from the Penrose
Fund of the American Philosophical Society and the Bache Fund of the National Acad-
emy of Sciences.

368 JOURNAL OF THE ARNOLD ARBORETUM _[VvoL. xxxIr

inflorescence-branches and pedicels usually glabrous at anthesis; leaf-blades
chartaceous to subcoriaceous, attenuate to obtuse at base.
Leaf-blades glabrous; 6 flowers with the rudimentary ovary copiously
puberulent at least distally.

Petioles 4-15 mm. long; leaf-blades usually 14-21 & 5-11 cm., narrowly
revolute at margin; 2 calyx at anthesis 1-1.2 mm. long and about 1.7 mm.
in diameter, the margin truncate or minutely denticulate, the limb
equalled or exceeded by the disk; disk glabrous on both sides, ciliolate

at apex; ovary narrowed into a short style 0.2-0.3 mm. long, the
stigmas slender, acute, sharply recurved. 2. A. insulare.

Petioles 1-5 mm. long; leaf-blades usually 4-15 a 2-7 cm., plane or
slightly recurved at margin; @ calyx at anthesis 1.5—1.7 mm. long and
usually more than 2 mm. in diameter, the margin 4- or 5-lobed or
dentate, the limb exceeding the disk; stigmas stout, obtuse, spreading
but not recurved.

Leaf-blades usually 8-15 X 3.5-7 cm., the costa stout (0.7-1.5 mm.
near base of blade); disk of 2 flowers pilose on both sides at

least distally as well as ciliolate; ovary narrowed into a short style
about 0.2 mm. long; rudimentary ovary in ¢ flowers oblong-ovoid,
about 0.8 mm. broad. ....................... 3. A. gillespieanum.
a 4-9 & 1.8-4 cm., the costa comparatively slender (0.5-0.8
m. broad near base of ‘blade) : disk of @ flowers essentially glabrous

sca at the ciliolate margin; ovary narrowed into an obvious style
about 1 mm, long; rudimentary ovary in 6 _ oblong-cylindric,
O50 a Wii, BYONG. os 5 inion che cedars A, elassophyllum.
Leaf-blades uniformly and persistently soft-pilose Hen oblong- or ovate-
elliptic, 8-13 cm. long, 4.5-7 cm. broad; young ¢ flowers with the disk
very short, much exceeded by the calyx-limb, the preset ovary
essentially glabrous. te ee ee sere 5. A. trichophyllum.

1. Antidesma (§ Tetrandra) pacificum Muell. Arg. in DC. Prodr. 15
(2): 254. 1866; Seem. FI. Vit. 217. 1867; Pax & Hoffm. in Pflanzenr.
81 [IV. 147. XV]: 150. 1922.

DIstTRIBUTION: Endemic to Fiji, known definitely only from Vanua Levu
and Moala, at low elevations. On Moala I noted it as a tree 5 m. high with a
deep purple fruit, growing in thickets.

Vanua Levu: Mathuata or Thakaundrove: Undu Point, Tothill 44 (K).
Moata: Near Maloku, Smith 1332 (Bish, GH, K, NY, US). Fiji, without definite
locality: U. S. Expl. Exped. (type cott., GH, US), Horne 491 (GH, K).

This very distinct species is quite different from the remaining Fijian
material of the genus; it is more closely related to the Samoan A. sphaero-
carpum Muell. Arg., which also belongs to § Tetrandra but has much larger
leaf-blades that are acute at base. Antidesma sphaerocarpum has been
erroneously reported from Fiji by K. Schumann (in Notizbl. Bot. Gart.
Berlin 2: 130. 1898); it appears to be limited to Samoa, where it is now
represented by numerous collections.

2. Antidesma (§ Montana) insulare Gillespie in Bishop Mus. Bull. 91:
12. fig. 13 (excl. e-g). 1932

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XIII 369

DISTRIBUTION: Endemic and apparently rare, as here circumscribed rep-
resented only by two collections from southeastern Viti Levu at elevations of
150-250 m

Vitt Levu: Rewa: Southeastern slopes of Mt. Korombamba, Gillespie 2292
(A, Bish type, GH, K, NY); Naitasiri: Tamavua woods, Gillespie 2030
(Bish, GH).

In describing this species, Gillespie cited six specimens in addition to the
type; he noted that “The montane specimens tend to have smaller leaves,
approaching lanceolate in shape, with shorter petioles, than those from
lower altitudes, as represented by the type.” A reconsideration of the Fijian
specimens of Antidesma, with the benefit of many more recent collections
in addition to those seen by Gillespie, convinces me that more than one
species of this alliance must be recognized. As represented by the type
and no. 2030, A. insulare differs from the bulk of the Fijian material of
§ Montana not only in the larger leaves and longer petioles, but also in the
narrowly revolute leaf-margin and in characters of the pistillate flower, as
mentioned in my key. Although Gillespie referred his species to § Venosa,
it clearly falls into § Montana as outlined by Pax & Hoffmann (in Pflanzenr.
81 [IV. 147. XV]: 112, 158-165. 1922), being of the general relationship
of the widespread A. bunius (L.) Spreng. Gillespie’s habit sketch and
drawings of the pistillate flower were apparently made from his type, but
the drawings of the staminate flower (figs. e-g) were from unspecified
material and are probably referable to the species I describe below as A.
gillespianium. Since in the present treatment A. insulare is used in a limited
sense, a redescription based on the two cited collections follows:

Small trees, glabrous except for the strigose-puberulent young parts and
some floral parts, the branchlets slender, verrucose-lenticellate; stipules
oblong, 4-5 mm. long, about 1.5 mm. broad, obtuse at apex, caducous;
petioles stout, rugose, shallowly canaliculate, 4-15 mm. long, the leaf-
blades chartaceous, drying brownish, oblong-elliptic, (10—) 14-21 cm. long,
(4—) 5-11 cm. broad, attenuate at base and decurrent on the petiole, short-
acuminate at apex, narrowly revolute at margin, the costa stout, nearly
plane above, prominent beneath, the secondary nerves 7—9 per side, erecto-
patent, slightly curved, obviously anastomosing 7-15 mm. within the mar-
gin, slightly elevated above, sharply elevated beneath, the veinlet-reticula-
tion coarse, immersed above, prominulous or subimmersed beneath; @ in-
florescences racemose, axillary, 3-6 cm. long at anthesis, short-pedunculate,
the rachis angled, the flower-subtending bracts deltoid, acute, 0.5—0.7 mm.
long, caducous, the pedicels 1-2 mm. long at anthesis; calyx thin-carnose,
cupuliform, 1—-1.2 mm. long, about 1.7 mm. in diameter, truncate at apex
or very minutely denticulate, the teeth (not more than 0.05 mm. long)
obscurely tufted-pilose at apex, the limb exceeded by the projecting ciliae
of the disk-margin; disk 0.4—0.5 mm. high, glabrous on both sides, copi-
ously ciliolate at margin with hairs 0.1-0.15 mm. long; ovary ovoid, at
anthesis 2.5—3 mm. long and about 2 mm. broad, narrowed into a short
style 0.20.3 mm. long, the stigmas 4, slender, narrowed to an acute apex,

370 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxx

0.5-0.7 mm. long, sharply recurved; ¢ inflorescences (Gillespie 2030)
immature; disk sparsely pilose without toward apex and copiously ciliolate,
the stamens 4 or 5, the rudimentary ovary oblong-ovoid, copiously tomen-
tellous-puberulent.

3. Antidesma (§ Montana) gillespieanum sp. nov.

Pesca dioica ad 12 m. alta partibus juvenilibus cinereo-strigosis et flo-
partibus exceptis ubique glabra, ramulis subteretibus cinereis verru-
epoateeiatl: apices versus 1.5-3 mm. diametro; stipulis chartaceis
elliptico-lanceolatis, 4-10 mm. longis, 1.5—-4 mm. latis: apice obtusis vel
rotundatis, caducis; foliis alternatis, petiolis crassis rugosis leviter canali-
culatis 1.5—5 mm. longis, laminis chartaceis vel subcoriaceis in sicco fusco-
viridibus vel fuscis, elliptico- vel lanceolato-oblongis, (6—) 8-15 cm. longis,
(2.5—) 3.5—7 cm. latis, basi obtusis vel acutis et in petiolum breviter decur-
rentibus, apice acutis vel breviter acuminatis, margine planis vel leviter
recurvatis, costa valida supra elevata vel subplana subtus prominente,
nervis cinage oa utrinsecus 5—9 patentibus paullo curvatis infra marginem
3-10 mm. anastomosantibus supra leviter subtus valde elevatis, rete venu-
larum crasso supra immerso vel paullo prominulo subtus manifeste promi-
nulo; inflorescentiis axillaribus racemosis vel ¢ paniculatis (ramulis 2-5)
2-7 cm. longis multifloris, pedunculo subnullo, rhachi crassa angulata
bracteis sub floribus ovato-deltoideis obtusis 0.5-1 mm. longis interdum
dorso obscure strigosis caducis; floribus @ : pedicellis sub anthesi 1-2 mm.
longis; calyce carnoso distaliter tenui cupuliformi, 1.5—-1.7 mm. longo, 2.5—
2.8 mm. diametro, interdum uno latere fisso, limbo quam disci margine
longiore irregulariter 4- vel 5-lobato, lobis late deltoideis 0.3-0.5 mm.
longis apice minute ciliolatis; disco carnoso cupuliformi 0.5—0.7 mm. alto
utrinque saltem distaliter dense piloso et margine pilis 0.1-0.15 mm. longis
copiose ciliolato; ovario ellipsoideo superne in stylum crassum circiter 0.2
mm. longum angustato, stigmatibus 3 vel 4 crassis obtusis 0.5—0.7 mm.
longis patentibus; floribus ¢: pedicellis sub anthesi 1.5-3.2 mm. longis;
calycis limbo margine integro vel obscure denticulato quam disco saepe
breviore; disco crasse carnoso 0.7—0.9 mm. alto ut 2 copiose piloso; stamini-
bus 4 vel 5 intra discum insertis, filamentis teretibus sub anthesi 1.5—2.5
mm. longis, antheris transverse ellipsoideis circiter 0.5 > 0.8—1 mm., con-
nectivo carnoso, loculis discretis; ovarii rudimento oblongo-ovoideo apice
truncato 1—-1.2 mm. longo circiter 0.8 mm. lato ubique copiose tomentello-
puberulo; pedicellis sub fructu 3-5 mm. longis, calyce demum subrotato;
drupa rhomboideo-ellipsoidea, 15-20 mm. longa, 10-12 mm. lata, utroque
angustata et obtusa, sarcocarpio carnosa in sicco valde contracto, putamine
13-17 mm. longo et 7-9 mm. lato utroque subacuto plerumque subcom-
planato, angulis lateralibus acutis, faciebus obtuse vel acute unicostatis
et medium versus saepe transverse angulatis.

DIsTRIBUTION: Known from scattered localities on Viti Levu, Vanua Levu,
and Kandavu, at elevations of 100-1100 m., but probably infrequently below
about 400 m, It is a forest tree, often slender, with a height of 4-12 m.; the
calyx is greenish or greenish yellow, the filaments are white or pale yellow, the

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XIII S71

anthers yellow, the stigmas greenish white, and the fruits deep red to purple.
Recorded local names are poroporo (Degener 15019) and saukalambuthi
(Smith 1572). As the diagnostic characters in this group are mostly in the
pistillate flowers, I designate my no. 5990, the only available 2 specimen, as

Vit Levu: Mba: Vicinity of Nandarivatu, Gillespie 4330 (Bish), Smith
5050 (A, US); Mt. Nanggaranambuluta [Lomalangi], Gillespie 4071 (Bish, GH);
Nandala, near Nandarivatu, Degener 15019 (A, Bish, K, NY, US); hills east
of Nandala Creek, Smith 6214 (A, US); hills between Negaliwana and
Tumbeindreketi Creeks, east of the sawmill at Navai, alt. 725-800 m., Sept. 12,
1947, Smith 5990 (A type, US), 5878 (A, US); Nandronga & Navosa:
Southern slopes of Nausori Highlands, in drainage of Namosi Creek above
Tumbenasolo, Smith 4719 (A, US); Namosi: Mt. Naitarandamu, Gillespie 3316
(Bish, GH, US); Mt. Vakarongasiu, Gillespie 3255 (Bish, GH, K, NY). VANua
Levu: Mbua: Southern portion of Seatovo Range, Smith 1572 (Bish, GH, K,
NY, US). Kanpavu: Mt. Mbuke Levu, Smith 230 (Bish, GH, K, NY, US).

Antidesma gillespieanum appears to be the most abundant Fijian repre-
sentative of the genus in middle-elevation forest, although it was not ob-
tained by collectors earlier than Gillespie. It differs from the lowland
A. insulare in its shorter-petiolate and smaller leaves with margins that
are not revolute, in its larger and distinctly lobed @ calyx of which the
limb exceeds the disk in length, in its pilose (rather than merely ciliolate)
disk, and in its comparatively stout,’ obtuse and spreading (but not re-
curved) stigmas. The available ¢ flowers of A, insulare are too immature
to permit comparison, but perhaps neither they nor the fruits would show
diagnostic characters. It may be noted that the ¢ flowers of A. gillespie-
anum differ from the @ in having the calyx-limb essentially entire and
the thicker disk projecting at anthesis.

4, Antidesma (§ Montana) elassophyllum sp. nov.

Arbor dioica ad 10 m. alta partibus juvenilibus strigoso-puberulis et
inflorescentiae partibus exceptis glabra, ramulis subteretibus copiose ver-
rucoso-lenticellatis apices versus 1-2 mm. diametro; stipulis papyraceis
oblongo-lanceolatis vel ellipticis, 4-7 mm. longis, 2-3.5 mm. latis, apice
rotundatis vel obtusis, utrinque parce strigosis vel glabris, caducis;
foliis alternatis, petiolis crassis leviter canaliculatis 1-5 mm. longis mox
glabratis, laminis chartaceis vel papyraceis in sicco fusco-viridibus, lanceo-
latis vel lanceolato-oblongis vel obovato-ellipticis, 4-9 cm. longis, 1.8—4 cm.
latis, basi obtusis vel attenuatis et in petiolum decurrentibus, apice obtusis
vel obtuse cuspidatis, margine planis, subtus juventute costa interdum
inconspicue puberulis mox glabratis, costa gracili supra plana subtus pro-
minente, nervis secundariis utrinsecus 4—7 patentibus vel suberectis leviter
curvatis infra marginem 2-5 mm. inconspicue anastomosantibus supra
planis vel immersis subtus prominulis, rete venularum crasso utrinque ob-
scuro vel subtus prominulo; inflorescentiis ¢ et @ axillaribus racemosis
sub anthesi et fructu 2—4.5 cm. longis, comparate paucifloris, pedunculo
brevi, rhachi sub anthesi obscure puberula, bracteis sub floribus deltoideis
subacutis 0.3-1 mm. longis dorso strigosis caducis; pedicellis sub anthesi

372 JOURNAL OF THE ARNOLD ARBORETUM _ [vot. xxxmt

1.2—2 mm. sub fructu ad 3 mm. longis glabris; floribus ?: calyce carnoso
distaliter submembranaceo cupuliformi, 1.5-1.7 mm. longo, 1.8—2.2 m
diametro, limbo quam disci margine valde longiore 4- vel 5-dentato, ‘en.
tibus intis acutis 0.2—-0.3 mm. longis apice obscure ciliolatis; disco subcar-
noso cupuliformi 0.4-0.5 mm. alto extus glabro intus parce piloso margine
pilis 0.2-0.3 mm. longis copioso ciliolato; ovario ellipsoideo superne in
stylum crassum manifestum circiter 1 mm. longum angustato, stigmatibus
3 vel 4 circiter 0.8 mm. longis crassis obtusis patentibus; floribus ¢ : caly-
cis limbo margine truncato vel irregulariter et minute dentato quam disco
interdum paullo breviore; disco crasse carnoso 0.5—0.8 mm. alto utrinque
parce puberulo et margine incrassato puberulo-ciliolato; staminibus 4 vel
5 intra discum insertis, filamentis sub anthesi 1.2-1.5 mm. longis, antheris
deltoideis 0.5—0.6 mm. latis, connectivo carnoso, loculis discretis; ovarii
rudimento oblongo-cylindrico 0.8-1.2 mm. longo 0.3-0.4 mm. lato, apice
truncato, superne copiose puberulo; drupa ei A. gillespieani simili saepe
paullo minore, 14-16 mm. longa, 9-12 mm. lata, putamine 10-15 x 7-9
mm.

DISTRIBUTION: Known only from Viti Levu and Vanua Levu, where it occurs
at elevations of 500-1120 m., characteristically in the mossy forest or dense
crest thickets of high or exposed ridges. It is a shrub or small tree 3-10 m.
high; the calyx and filaments are white and the fruits red to purplish. Recorded
local names are molau (Smith 557) and natha (Smith 656), As the type I
designate my no. 656, from Vanua Levu, the only available specimen with
pistillate flowers.

Vitt Levu: Mba: Mt. Evans Range, Greenwood 951 (A, US), 1073 (A, US),
1263 (US); vicinity of Nandarivatu, Tothill 376 (K), Parks 20538 (Bish); near
summit of Mt. Nanggaranambuluta [Lomalangi], Gillespie 3784 (A, Bish);
hills east of Nandala Creek, south of Nandarivatu, Smith 5941 (A, US). VANUA
Levu: Mathuata: Summit ridge of Mt. Numbuiloa, east of Lambasa, Smith
6465 (A, US); Thakaundrove-Mathuata boundary: Crest of Korotini
Range, between Navitho Pass and Mt. Ndelaikoro, Smith 557 (Bish, GH, K, NY,
US); Thakaundrove: Mt. Mbatini, crest of range, alt. 700-1030 m., Nov. 28,
1933, Smith 656 (Bish, GH, K, NY type, US). Fiji, without definite locality:
Gillespie 4051 (A, Bish).

Antidesma elassophyllum differs from A. insulare in foliage to an even
greater degree than does the above-described A. gillespieanum, to which it
is evidently most closely related. From A. gillespieanum the new species
differs in its smaller leaves; this character is not entirely dependable but
is supplemented by the difference in the robustness of the costa as ex-
pressed in my key. Antidesma elassophyllum is further distinguished by
having the disk of its 2 flowers essentially glabrous except at the ciliolate
margin and by its obvious style; in the ¢ flowers the rudimentary ovary
is comparatively slender. The available specimens of this entire complex
are not sufficiently adequate to indicate whether floral differences are en-
tirely satisfactory, but those differences observed are correlated with the
more obvious foliage characters, so that discontinuities in the Fijian popu-
lation of § Montana are clearly indicated.

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XIII 373

5. Antidesma (§ Montana) trichophyllum sp. nov.

Arbor dioica ad 8 m. alta, partibus juvenilibus pilis stramineis 0.2—-0.3 mm.
longis copiose hispidulo-puberulis mox glabratis, ramulis subteretibus cine-
reis apices versus 2-3 mm. diametro puberulis demum glabratis; stipulis
papyraceis lineari-lanceolatis, 5-8 mm. longis, 1-2 mm. latis, apice acutis,
utrinque parce puberulis mox glabratis; foliis alternatis, petiolis crassis
rugosis semiteretibus 5—8 mm. longis ut ramulis puberulis glabratisque, la-
minis chartaceis vel subcoriaceis in sicco fusco-viridibus, oblongo- vel ovato-
ellipticis, 8-13 cm. longis, 4.5—7 cm. latis, basi obtusis vel subacutis et in
petiolum breviter decurrentibus, apice acutis vel breviter acuminatis, mar-
gine planis, supra costa saepe puberula excepta glabris, subtus pilis patenti-
bus 0.3-0.5 mm. longis uniformiter et persistenter molli-pilosis, costa valida
supra subplana subtus prominente, nervis secundariis utrinsecus 5—8 erecto-
patentibus subcurvatis infra marginem 8-12 mm. anastomosantibus supra
subplanis subtus valde elevatis, rete venularum crasso utrinque immerso
vel prominulo; inflorescentiis ¢ immaturis solis visis axillaribus vel infra
folia enatis, racemosis vel basim versus 2- vel 3-ramosis, 2-4 cm. longis
multifloris, pedunculo brevi, rhachi striata glabra, bracteis sub floribus
papyraceis deltoideis acutis 0.6—0.8 mm. longis dorso parce strigosis cadu-
cis; pedicellis glabris ante anthesin ad 1 mm. longis; floribus eis A. gilles-
pieani subsimilibus; calycis limbo margine obscure dentato ciliolato, disco
in alabastro circiter 0.3 mm. alto quam calycis limbo multo breviore, extus
subglabro, intus parce piloso, margine ciliis, 0.2-0.3 mm. longis ornato;
staminibus 5 intra discum insertis, filamentis glabris, antheris circiter 0.5
< 0.8 mm.; ovarii rudimento oblongo-ovoideo circiter 0.8 & 0.7 mm. apice
truncato ubique glabro vel apice obscure puberulo.

Vitr Levu: Nandronga & Navosa: Northern portion of Rairaimatuku
Plateau, between Nandrau and Nanga, alt. 725-825 m., Aug. 7, 1947, Smith 5573
(A type, US) (molatha; slender tree 8 m. high, in dense forest; flower-buds
yellowish).

Although the new species is described from a single collection bearing
immature staminate inflorescences, it differs markedly from the other Fijian
representatives of the genus in having its leaf-blades uniformly and per-
sistently soft-pilose beneath. No intermediate states between this pubes-
cent form and the usual glabrous-leaved type have been observed. Further
collections are needed to indicate whether the @ flower will provide any
dependable characters and whether the very short disk and the essentially
glabrous rudimentary ovary of the ¢ flower are consequential.

GLOCHIDION J. R. & G. Forst.
Glochidion euryoides sp. nov.

Arbor monoica gracilis ad 4 m. alta multiramosa ubique glabra, ramulis
cinereis teretibus apices versus subflexuosis, internodiis distalibus 3-7 mm.
longis, foliis alternatis congestis, petiolis rugulosis 2-4 mm. longis fere ad
basim anguste alatis, laminis chartaceis vel subcoriaceis in sicco fusco-

374 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxxur

viridibus oblongo-ellipticis, 2-3 cm. longis, 1-1.7 cm. latis, basi obtusis
et in petiolum subito decurrentibus, apice rotundatis et inconspicue glandu-
loso-mucronulatis, margine integris incrassatis, costa supra subplana subtus
valde elevata, nervis secundariis utrinsecus 6—9 patentibus inconspicue anas-
tomosantibus supra planis subtus prominulis, rete venularum immerso
vel subtus obscure prominulo; inflorescentiis axillaribus congestis glomera-
tis, bracteis basalibus 2 papyraceis deltoideis acutis 1-1.3 mm. longis,
bracteolis sub floribus pluribus bracteis similibus sed 0.5—0.8 mm. longis;
floribus ¢ in inflorescentia paucis, pedicellis gracilibus 1-3 mm. longis;
perianthio carnoso, tubo breviter obconico, segmentis 6 late imbricatis
oblongis apice obtusis margine scariosis, 3 exterioribus 1.3-1.6 mm. longis
et 0.8-1.2 mm. latis, 3 interioribus paullo minoribus; columna staminali
ellipsoidea circiter 1 mm. longa, antheris 3 loculis circiter 0.7 mm. longis
connatis, connectivis in apices carnosos deltoideos 0.2—0.3 mm. longos libe-
ros productis; floribus ¢ in inflorescentia (1—) 3—6 sessilibus vel pedicello
haud ad 0.5 mm. longo; perianthio ¢ simili sed segmentis 6 subaequalibus
1.6—2 mm. longis et 1-1.3 mm. latis; ovario depresso-turbinato sub anthesi
0.8-1 mm. diametro, loculis 6, ovulis in quoque loculo 2 collateralibus,
columna stylari carnosa 1.5—2 mm. longa (post anthesin nae stylis
6 apices versus liberis et ventro sulcatis apice emarginatis

Vit1 LEvu: Mba: Upper slopes of Mt. Koromba rPickering Peak ], alt. ee
1075 m., June 3, 1947, Smith 4659 (A type, US) (slender tree 4 m. high,
forest on ridges and spurs; perianth white).

The very distinct new species here described is not closely related to
any species of our region, being characterized by its strictly glabrous habit,
its very small and congested leaves, its sessile pistillate flowers with 6
comparatively large and subequal perianth-segments, its 6-loculate ovary,
and its thick stylar column with distally free styles. It is probably to be
sought in § Hemiglochidion (cf. Pax & Hoffmann in Nat. Pflanzenfam. ed.
2. 19c: 56-58. 1931), in some respects suggesting G. vitiense (Muell. Arg.)
Gillespie. That species, however, has comparatively well spaced and large
leaves and pedicellate ¢? flowers, of which the perianth-segments are much
smaller.

BURAEAVIA Baill.
Buraeavia horneana sp. nov.

Arbor dioica ad 10 m. alta, partibus juvenilibus pilis aureis 0.2—-0.3 mm.
longis strigosis, alioqui inflorescentiis exceptis glabra, ramulis gracilibus
subteretibus rugulosis cinereis; foliis oppositis, petiolis rugulosis leviter
canaliculatis 5-10 mm. longis, laminis chartaceis vel subcoriaceis ow sicco
subviridibus vel fuscis, elliptico- vel obovato-rhomboideis, (3—) 5-8 cm
longis, (1.5—) 2.5—4.5 cm. latis, basi acutis vel attenuatis et in bp
decurrentibus, apice obtusis vel rotundatis, margine integris et leviter in-
crassatis, costa supra leviter elevata vel plana subtus prominente, nervis
lateralibus utrinsecus 4-6 adscendentibus anastomosantibus utrinque pro-
minulis, rete venularum subimmerso; inflorescentiis ¢ axillaribus vel infra

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XIII 375

folia ortis plerumque 2 vel 3 superpositis compacte cymosis plurifloris
ante anthesin ad 7 mm. longis, pedunculo brevi cupulam bracteis oppositis
compositam apice gerente, bracteis subcoriaceis deltoideis subacutis 1—2
mm. longis ut pedunculo fulvo-strigosis, ramulis 3—5 parce strigosis radiatis,
bracteolis oppositis 0.5-1 mm. longis, pedicellis teretibus 1—1.3 mm. longis
subglabris; sepalis 4 membranaceis suborbicularibus, 2 exterioribus circiter
1.5 mm. diametro extus parce strigosis, 2 interioribus minoribus glabris;
staminibus 7-9, filamentis haud 0.3 mm. longis, antheris subgloboso-oblon-
gis circiter 0.5 mm. diametro, disco pulvinato minuto obscuro 4-gono;
floribus @ in axillis foliorum vel e nodis defoliatis 2—4 aggregatis, pedun-
culo ruguloso 1-2 mm. longo apice inconspicue bibracteato, pedicellis paullo
post anthesin circiter 3 mm. longis; sepalis papyraceis glabris, 2 ex-
terioribus ovatis obtusis circiter 1 mm. longis latisque, 2 interioribus sub-
reniformibus circiter 1 X 1.7 mm. margine irregulariter spinuloso-denticula-
tis; disco subcarnoso cupuliformi circiter 0.7 mm. alto inconspicue 4-lobato
margine conspicue ciliolato ovarii basim cincto; ovario ellipsoideo glabro,
stigmate sessili discoideo-pulvinato circiter 1.5 mm. diametro 3-lobato,
lobis leviter sulcatis, loculis 3, ovulis 2 apice carunculae dependentis car-
nosae collateralibus; pedicellis sub fructu 4-7 mm. longis, calyce persisten-
tibus; fructibus maturitate 7-8 mm. diametro, exocarpio in sicco tenui
subcoriaceo ruguloso ab endocarpio in valvas 3 solubili, endocarpio circiter
0.5 mm, crasso in coccos 2-valves mox dissiliente, columella apice dilatata
persistente; seminibus plerumque 2 in coccis descendentibus ellipsoideis
circiter 5 mm. longis et 3 mm. latis, arillo aurantiaco irregulariter laciniato,
testa in sicco rubro-castanea.

VANUA Levu: Mathuata: Seanggangga Plateau, in drainage of Korovuli
River, vicinity of Natua, in patches of forest in open rolling country, alt. 100-
200 m., Smith 6669 (A, US) (tree 6 m. high; fruit dull red), 6872 (A Tver, US)
(Dec. 6, 1947; tree 10 m. high; young inflorescences yellowish green);
Mathuata or Thal drove: Between Waiwai and Lomaloma, alt. about
600 m., Horne 600 (GH, K).

The occurrence of a species of Buraeavia in Fiji is of particular interest,
as otherwise the genus is known only from the two New Caledonian species
originally assigned to it by Baillon (in Adansonia 11: 84. 1873). The ex-
istence of the Fijian species was intimated by Bentham & Hooker (Gen.
Pl. 3: 280. 1880), who briefly discussed the Horne specimen cited above.
Of the three known collections of the new species, the type bears staminate
inflorescences, Horne 600 has pistillate flowers past anthesis and a few
fruits, and my no. 6669 bears mature fruits. In thus extending the range
of another “New Caledonian” genus, one may note in passing that Burae-
avia, like several other genera common to New Caledonia and Fiji, is
known from Vanua Levu but not from Viti Levu (cf. my remarks in Sci.
Monthly 73: 12. 1951).

The Fijian species is allied to B. carunculata Baill., from which it is
readily distinguished by its rhomboid leaf-blades which are gradually nar-
rowed at base and decurrent on the obvious petioles, by its shorter-pedi-
celled staminate flowers with fewer stamens, and by its smaller fruits.

376 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. XxxIII

There is disagreement among students of the family as to the separation
of Buraeavia Baill. (1873) from Longetia Baill. (1866). Pax & Hoffmann
(in Pflanzenr. 81 [IV. 147, XV]: 289. 1922, in Nat. Pflanzenfam. 19c:
75. 1931) combine them under the earlier name, Longetia, but Guillaumin
(Fl. Anal. et Synopt. Nouvelle-Caléd. 175, 181. 1948) retains both genera,
indicating that Longetia lacks the disk that is characteristic of Buraeavia.
The latter viewpoint is here adopted, but if future students consider the
genera not separable our species will need a combination in Longetia.

MACARANGA Thou.

Pax & Hoffmann, in their revision of the vast and complex genus
Macaranga (in Pflanzenr. 63 [IV. 147. VII]: 298-395. 1914), indicate
three main trends of development within the genus, based on the surface
of the developing fruit: (1) Laeves, with the capsule smooth; (2) Tuber-
culatae, having the capsule with short thick tubercles covering the surface
to a greater or lesser degree; and (3) Echinatae, having the capsule orna-
mented with spines often of considerable length. In each of these major
divisions, smaller categories are based upon combinations of such charac-
ters as the presence or absence of patelliform glands on the inflorescence-
bracts, the number of ovary-locules, the number of anther-locules, and the
basic venation pattern (whether palmate or pinnate). As a result of this
selection of characters, Pax & Hoffmann propose 32 sections (36 sections
in a later treatment, in Nat. Pflanzenfam. ed. 2. 19c: 128-134. 1931).
The phylogenetic validity and the usefulness of these sections cannot be
assessed without a study of the entire genus, but if this system is carried
to its logical conclusion certain additional sections will be needed. For
instance, there is no proposed section for species with tuberculate fruits,
patelliform-glandular bracts, and palmately nerved leaves, although two
Fijian species proposed by Pax & Hoffmann (M. vitiensis and M. graeffe-
ana, of which they did not know the fruits) have this combination of
characters. In the present treatment I follow Pax & Hoffmann’s grouping
as to the Fijian species, but the major groups Laeves and Tuberculatae
are not sharply distinct in our region.

Eight species, of which six are endemic, seem to occur in Fiji, on the
basis of material now available; two of these are further subdivided and
one of them is described as new. Since the original descriptions are usually
inadequate I here include brief descriptions emphasizing the salient fea-
tures. All of the Fijian species normally have peltate and palmately
nerved leaves.

KEY TO THE SPECIES
Fruits smooth or tuberculate with oblong-conical processes not more than 1.5
mm. long; inflorescence-bracts (at least the larger ones) patelliform-glan-
dular; styles 2, divaricate, not more than 3 mm. long.

Inflorescence-branches, bracts, pedicels, and at least the 2 calyces persistently

tomentellous or spreading-puberulent; fruits smooth.

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XIII 377

Leaf-blades iti ig pilose on both surfaces with soft persistent hairs
.S-1 mm. long, the hairs of the inflorescence-indument 0.5 mm. lon

or more, the & calyx glabrous; stamens 12-15; stipules, branchlets,
and petioles copiously pilose (hairs 0.4-1.3 mm. long). ............
ee TLL er Tea a ee ere ere ty ee . membranacea.
Leaf-blades glabrous or with the nerves beneath (rarely above) puberulent
or tomentellous, the hairs of the inflorescence-indument minute, rarely
more than 0.3 mm. long, the ¢ calyx with a similar but sparser
indument, at length glabrate; stamens 6-11; stipules, branchlets, and
petioles usually glabrous, sometimes short- pilose. ....2,. M. seemannit,
ama oa branches, bracts, pedicels, and calyces glabrous or soon glabrate
t w hairs, if present, scattered and not forming a uniform indument).
Leaf- rae broadly ovate to deltoid, less than twice as long as broad;
fruits smooth or with processes rarely exceeding 0.5 mm. in length.
ate! his large, the petioles 22-45 cm. long, the blades 34-60 cm.
3-50 cm. broad, broadly peltate (petiole attached 7-10 cm.

ae “ margin), the veinlets strongly elevated on lower surface;
inflorescence often 15-30 cm. long, freely branching; stamens 12- 14;
developing ovary (mature fruits not seen) smooth ....3. M. m magna.
Leaves smaller, the petioles 5-21 cm. long, the blades 8-30 cm. long,
5-23 cm. broad, less broadly peltate (petiole attached 1-6 cm.

from basal margin), the veinlets on lower surface plane or merel
prominulous; inflorescence not exceeding 12 cm. in length; stamens

5-10

Stipules 1-5 cm. long; leaf-blades with 5-8 primary nerves; fruits
comparatively small, 3.5-5 mm. long, 6-8 mm. broad, often
ICUS, os ne ce Pe para eee 4. M. graeffeana.

Stipules 5-7 cm. long, 12-18 mm. broad, glabrous; branchlets and
petioles glabrous, the eng 15-19 cm. long; leaf-blades broadly
ovate, 13-17 K 12-16.5 cm., the petiole attached 3-4.5 cm. from
basal margin, the ees nerves 8-10, the margin closely callose-
crenulate; fruits comparatively large, the mature capsules 6-7 mm.
long, 9-11 mm. broad, smooth or very rarely with a few scattered
conical tubercles 0.1-0.4 mm. long. .......... S. . marikoensis.

Leaf-blades deltoid- or ovate-lanceolate, avetaging 2.5-3 times as long as
broad (13-55 & 4-20 mm.); fruits copiously tuberculate with oblong-
conical processes 0.5—1.5 mm. long; stamens 3-6. ..... 6. M. vitiensis.

Fruits ornamented with somewhat flattened subulate processes 1.5-8 mm. long;
inflorescence-bracts not patelliform- ann inflorescence-branches, bracts,

pedicels, and calyces copiously puberu

Flower-subtending bracts of ¢ A srs 2—4 mm. long, entire; stamens
( 6-9; fruits with processes 3-8 mm. long and with 3 styles, these
4-10 mm. long, ascending to spreading, copiously papillose; branchlets and
petioles glabrous (at least in Fijian specimens) ...... 7. M. harveyana.
Flower-subtending bracts of ¢ inflorescences 4-10 mm. long, at least the
larger ones fimbriate-dentate; stamens 3-5; fruits with processes 1.5-3
mm. long and with 2 styles, these 1.5-2.5 mm. long, divaricate, not or
inconspicuously papillose; branchlets and petioles copiously puberulent.

SRG ic Hehe eGo sits og & ar tse Bete eh te teea Ste eR Rese ahi ok 8, . secunda,.

378 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxx

1. Macaranga (§ Adenoceras) membranacea Muell. Arg. in DC. Prodr.
15 (2): 996. 1866; Seem. FI. Vit. 228. 1867; Pax & Hoffm. in Pflan-
zenr. 63 [IV. 147. VII]: 393. 1914.

Tanarius membranaceus Kuntze, Rev. Gen. Pl. 2: 620. 1891.

Shrub or tree up to 8 m. high, the branchlets and petioles copiously
spreading-pilose with pale hairs 0.5—1.3 mm. long; stipules lanceolate, 1.5—
2.5 mm. long, densely pilose on both sides with hairs 0.4-0.8 mm..: long;
petioles 6-13 cm. long, the blades broadly ovate, 10-24 cm. long, 7-14 cm.
broad, rounded at base and broadly (1.5-4 cm.) peltate, rarely deeply
cordate and epeltate, long-acuminate at apex, the primary nerves 6 or 7,
both surfaces copiously spreading-pilose with soft hairs 0.5—-1 mm. long,
scattered-glandular beneath; inflorescences 5-8 cm. long, the branches,
bracts, and @ calyces densely pilose with spreading hairs 0.5—0.7 mm. long,
the bracts lanceolate or obovate-lanceolate, up to 5 mm. long, at least the ¢
with often obscure patelliform glands; 3 calyx glabrous, glandular distally,
about 1.5 mm. in diameter, the lobes 3, oblong-ovate, rounded; stamens
12-15, the filaments 1-1.2 mm. long; fruits smooth, copiously glandular,
the styles 2, divaricate, 1-2 mm. long.

DisTRIBUTION: Apparently endemic, and known with certainty only from
Vanua Levu; the type, an Exploring Expedition specimen, is without data but
may also be from Mathuata, where the species has been noted at ins up
to 200 m., on edge of forest or in patches of forest in open country. It is said
to bea large shrub or a tree 5-8 m. high, with pale yellow latex that turns on
exposure, and with the calyx and filaments pale greenish yellow. The type is
sterile, but an isotype precisely agrees with the cited specimens. I noted the
local name as mama.

ANUA Levu: H. B. R. Parham 342 (BM); Mathuata: Mathuata coast,
Greenwood 654 (K); Seanggangga Plateau, in drainage of Korovuli River,
vicinity of Natua, Smith 6641 (A, US), 6707 (A, US). Fiji, without definite
locality: U. S. Expl. Exped. (type coLu., GH).

As his material was sterile, Mueller was unable to suggest the alliance
of this patently distinct species. The collection of staminate (Smith) and
fruiting (Greenwood) inflorescences establishes the plant as belonging to §
Adenoceras and as closely related only to M. seemannii, from which it
differs in the longer and more abundant indument (except on the 4 calyx,
which is quite glabrous) and in the increased number of stamens.

2. Macaranga (§ Adenoceras) seemannii (Muell. Arg.) Muell. Arg. in
DC. Prodr. 15 (2): 999, as M. seemanni. 1866.

Macaranga seemannii, a sharply marked species of § Adenoceras,
differs from the species of our region (except the preceding, M. mem-

persisting even on pedicels and calyces of fully mature infructescences.
Although most of the material here considered agrees well with the cotypes
of the species, two variants seem worthy of note. As these have the basic

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XIII 379

characters of M. seemannii, I hardly consider them worthy of specific
recognition, but they are perhaps worth recording as varieties.

KEY TO THE VARIETIES

Stipules oblong-lanceolate, 1.5-4 cm. long, 5-15 mm. broad; petioles 9-27 cm.
long; leaf-blades broadly ovate, usually 13-31 % 8.5-22 cm., rounded at
base, broadly ara (petiole attached 2.5-8 cm. from basal margin);
infructescence 3-12 c

Branchlets and petioles ene or with a few scattered hairs, not uniformly
soft-pilose; stipules 5-10 mm. broad, glabrous or puberulent without

he young; petioles 9-18 cm. long; earn nerves of leaf-blades 6-8;
hairs of inflorescence-indument 0.1—0.2 mm. long ....2a. var. seemannit.
Branchlets and petioles copiously soft- ae stipules 8-15 mm. broad,
puberulent on both sides or glabrate distally; petioles 16-27 cm. long;
primary nerves of leaf-blades 8 or 9; hairs of inflorescence-indument 0.2—

O.4 Wi: TOMY 365553084 ee cus i 2b. var. capillata.
supe ene oblong- -lanceolate, 1- 2 cm. - long, “34 mm. broad, copiously
preading-pilose on both sides; branchlets and petioles densely tomentellous-
Eibaniee the petioles 8-12 cm. long; leaf-blades ovate-deltoid, 12--16 X
7-10 cm., rounded-truncate at base, less broadly peltate (petiole attached
1.5-2.5 cm. from basal margin); infructescence 2-3 cm. long. ............

PO er Te ee eer ee eT 2c. var. delioidea.

2a. Macaranga seemannii var. seemannii

Mappa seemanni Muell. Arg. in Flora 47: 468.

Macaranga seemanni Muell. Arg. in DC. ne i: (2): 999. 1866; Seem. Fl.
Vit. 228. 1867.

Tanarius seemannii Kuntze, Rev. Gen. Pl. 2: 620. 1891.

Macaranga seemannii Muell. Arg. ex Hemsl. in Jour. Linn. Soc. Bot. 30: 192.
1894; Pax & Hoffm. in Pflanzenr. 63 [IV. 147. VII] : 336. 1914.

Tree up to 18 m. high, the branchlets and petioles glabrous or with a
few scattered spreading hairs up to 1 mm. long, not uniformly soft-pilose,
glabrate; stipules oblong-lanceolate, 1.5-4 cm. long, 5-10 mm. broad,
glabrous or sometimes puberulent without when young; petioles 9-13 cm.
long, the blades broadly ovate, (12—) 13-30 cm. long, (7—) 8.5—-22 cm.
broad, rounded at base and broadly (2.5—7.5 cm.) peltate, long-acuminate
at apex, with 6-8 primary nerves, glabrous above, spreading-puberulent on
primary nerves beneath or with occasional longer hairs to 1.5 mm. long,
copiously glandular beneath; inflorescences 3-12 cm. long, the branches,
bracts, pedicels, and calyces tomentellous or spreading-puberulent with
ferrugineous hairs 0.1-0.2 mm. long, the bracts lanceolate or obovate-
lanceolate, up to 5 mm. long, the larger ones obviously patelliform-
glandular; ¢ calyx about 1 mm. in diameter, at length subglabrate, the
lobes 3, oblong-ovate; stamens 6—11, the filaments 0.5—-1 mm. long; fruits
smooth, copiously glandular, the styles 2, divaricate, 1-2 mm. long.

DIsTRIBUTION: Fiji and Tonga; known from several islands in Fiji, at eleva-
tions from near sea-level up to 1100 m., occurring in different types of forest, in
thin woods among reeds, along streams, etc. It is a tree 4-18 m. in height, with a
thin, colorless or yellowish latex. The calyx, filaments, and styles are pale yellow
and the fruit greenish yellow. Local names are mdavo (upland Viti Levu) and

380 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxxu

mama (Mathuata). In the original description Mueller cites three specimens, all
deposited at Kew and cited below, which may be taken as cotypes, these are
Harvey s. n. and Seemann 397 and 419. The occurrence of the species in Tonga
is indicated by U. S. Expl. Exped. (US 66261).

Vitt Levu: “Nakaru,” Parks 20403 (Bish, GH); Mba: Mountains near
Lautoka, Greenwood 285 (K), 286 (K), 1197 (US); slopes of the escarpment
north of Nandarivatu, Smith 6028 (A, US); vicinity of Nandarivatu, Parks
20655 (Bish); valley of Nggaliwana Creek, Smith 5334 (A, US); slopes of Mt.
Tomanivi, Smith 5212 (A, US); Nandronga & Navosa: Northern portion
of Rairaimatuku Plateau, between Nandrau and Rewasau, Smith 5637 (A, US);
Namosi: Vicinity of Namuamua, Gillespie 3012 (Bish, GH), 3049 (Bish, GH,
K, NY, US); vicinity of Namosi, Gillespie 2499 (Bish); Naitasiri: Viria,
Meebold 16900 (Bish, K); vicinity of Nasinu, Gillespie 3654 (Bish); Rewa:
Lami, Tothill 738 (K); near Suva, Tothill 739 (K); Tailevu: Naivithula,
Valentine 18 (Bish). OvaLau: Vicinity of Levuka, Gillespie 4458 (Bish).
Koro: Tothill 697 (K), 698 (K). VANuA Levu: H. B. R. Parham 39 (BM);
Mathuata: Southern slopes of Mt. Numbuiloa, east of Lambasa, Smith 6546
(A, US). TaAveuntr: Vicinity of Wairiki, Gillespie 4716 (Bish, GH, K, NY).
Fiji, without definite locality: Harvey (BM, GH, K cotype), Seemann 397
(GH, K cotyre), 419 (BM, GH, K cotype) [three cotypes indicated as from
Viti Levu in Flora Vitiensis, but Kew sheet of no. 4179 indicated as from
Taveuni], Horne 826 (GH, K).

2b. Macaranga seemannii var. capillata var. nov.

Arbor ad 15 m. alta, ramulis petiolisque copiose pilosis (pilis patentibus
0.2-0.3 mm. vel interdum ad 1 mm. longis); stipulis oblongo-lanceolatis
3-4 cm. longis 8-15 mm. latis utrinque puberulis vel superne glabratis;
petiolis 16-27 cm. longis, laminis late ovatis, 20-31 cm. longis, 16—22 cm.
latis, basi rotundatis et late (5.5—8 cm.) peltatis, apice acuminatis, supra
glabris, subtus praeter nervos saltem basim versus pilosos glabris et
copiose glandulosis, nervis primariis 8 vel 9; inflorescentiis ? et fructiferis
solis visis 5-12 cm. longis ubique pilis 0. 2-0. 4 mm. longis copiose tomen-
tellis, fructibus ut in var. seemannit.

KanpAvu: Hills above Namalata and Ngaloa Bays, alt. 200-400 m., Smith 46
(Bish, GH, K, NY, US) (venua; tree 15 m. high, on edge of forest; fruits green
to black). KAmBaArRA: Lowland forest of central basin, alt. 25-30 m., Aug. 24,
1924, Bryan 500 (Bish TYPE) (venua; tree 10-12 m. high, the trunk 18-25 cm.
diam., with red latex; flowers and fruits green).

The typical variety of M. seemannii is quite uniform, on the basis of
the many collections referred to it above, and from it the new variety
differs in its somewhat longer petioles, large leaf-blades and stipules, the
soft indument of its branchlets and petioles, and the slightly longer hairs
of its inflorescence-indument.

2c. Macaranga seemannii var. deltoidea var. nov.

Arbor ad 5 m. alta, ramulis petiolisque copiose tomentello-puberulis
(pilis 0.2-0.3 mm. longis vel interdum longioribus); stipulis anguste
oblongo-lanceolatis 1-2 cm. longis 3—4 mm. latis utrinque copiose pilosis;
petiolis 8-12 cm. longis, laminis ovato-deltoideis, 12-16 cm. longis, 7-10

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XIII 381

cm. latis, basi rotundato-truncatis et 1.5—-2.5 cm. peltatis, apice graciliter
acuminatis, supra glabris vel primo nervis tomentello-puberulis, subtus
nervis et interdum venulis ut petiolis tomentellis et copiose glandulosis,
nervis primariis 7 vel 8; infructescentiis 2—3 cm. longis pilis ferrugineis
0.1-0.3 mm. longis ubique copiose tomentello-puberulis, bracteis caducis,
calyce sub fructu extus puberulo intus glabrato, fructibus ut in var.
seemannit.

Vitr Levu: Mba: Upper slopes of Mt. Koromba [Pickering Peak], a
800-1075 m., June 3, 1947, Smith 4669 (A Typr, US) (tree 5 m. high, in dense
forest on ridges and spurs).

From the typical variety of M. seemannii, the new variety differs
primarily in its very narrow and copiously pilose stipules and the tomentel-
lous-puberulent character of its branchlets and petioles. The leaf-blades
are somewhat narrower than typical and the infructescence is comparatively
short, but these characters may not be very significant.

3. Macaranga (§Adenoceras) magna Turrill in Kew Bull. 1924: 393.
1924.
Macaranga grandifolia Turrill in Jour. Linn. Soc. Bot. 43: 38. 1915, non
Merr. (1913).

Tree up to 10 m. high, the branchlets glabrous or very soon glabrate;
stipules oblong-lanceolate, 3—5 cm. long, 1.5—2.5 cm. broad, at first with
spreading hairs 0.3-0.5 mm. ibis at length glabrate, sometimes copiously
sessile-glandular without; petioles 22-45 cm. long, copiously spreading-
puberulent with pale hairs 0.2-0.5 mm. long, glabrate (indument often
remaining in irregular patches); leaf-blades ample, ovate, 34-60 cm. long,
23-50 cm. broad, rounded or coarsely undulate-truncate at base and
broadly (7-10 cm.) peltate, acuminate or cuspidate at apex, glabrous
above or sparsely setulose on nerves with hairs 0.3-0.6 mm. long, similar
beneath or soft-spreading-pilose on nerves and copiously glandular, the
primary nerves 6—9, the veinlets strongly elevated beneath; inflorescences
glabrous, the ¢ freely branching, 15-30 cm. long, the @ similar but 6—20
cm. long, the larger bracts lanceolate-obovate, 4-8 mm. long, patelliform-
glandular; ¢ calyx 1.3-1.6 mm. in diameter, eglandular or with few
distal glands, 3-lobed, the lobes broadly ovate; stamens 12-14, the fila-
ments 0.8—1 mm. long: developing ovary smooth, copiously glandular, the
Styles 2, divaricate, 1.2-1.5 mm. long.

ISTRIBUTION: Endemic and apparently limited to Viti Levu, occurring at
elevations of 300-970 m. in forest or on edges of forest. The species is a
slender tree up to 10 m. in height, the inflorescence-parts (branches, bracts,
calyces, and styles) being dark red or crimson. Local names are mdavo and
ndavolutu. The type is im Thurn 134, cited below.

Vitrt Levu: Mba: Nandarivatu and vicinity, im Thurn 134 (K Type),
Tothill 737 (K), Gillespie 3986 (Bish, GH, US); southern slopes of Mt.
Ndelainathovu, on the escarpment west of Nandarivatu, Smith 4951 (A, US);
Nandronga & Navosa: Vicinity of Nandrau, Degener 14911 (A, NY);
Rewa: Mt. Korombamba, Gillespie 2379 (Bish, K, NY); Naitasiri?:

382 JOURNAL OF THE ARNOLD ARBORETUM _[VoL. xxx1I

Prince’s Road, Meebold 21364 (NY). Fiji, without definite locality: Parks
20895 (Bish)

Although mature fruits are not yet available, M. magna, a sharply
marked and spectacular species, seems to fall into § Adenoceras in the Pax
& Hoffmann system, although it might also be sought in § Stipulosae.
The latter hardly seems worth separating from § Adenoceras, if it depends
entirely upon the size of the stipules, which are said to be 5-15 cm. long
in the Samoan M. stipulosa (the only species of § Stipulosae) as opposed
to 1 cm. in § Adenoceras. However, such species as M. seemannii, admitted
into § Adenoceras, have stipules up to 4 cm. in length. The several sheets
of M. stipulosa available to me, including an isotype, unfortunately do not
show stipules, but at any rate M. magna is readily distinguished from the
Samoan plant by its glabrous inflorescences and strongly elevated veinlets
on the lower leaf-surfaces.

4, Eseg ae (§ etn oo Pax & Hoffm. in Notizbl.
Bot. Gart. Berlin 10: 384.

Macaranga graeffeana, described subsequent to Pax & Hoffmann’s
treatment of the genus in the Pflanzenreich, has not been well understood,
practically no specimens in herbaria having been referred to it. The type
is a Graeffe specimen from Viti Levu, described without fruit and
supposedly related to M. seemannii and M. vitiensis. The original
description agrees very closely with several available collections from
southern Viti Levu, these differing from M. seemannii in their essentially
glabrous inflorescences and from M. vitiensis in obvious leaf-proportions.
As Graeffe’s collections come in large part from this same region of Viti
Levu, I have little hesitation in referring Pax & Hoffmann’s name to the
entity here discussed.

The fruit of typical M. graeffeana, as here construed, is usually sparsely
tuberculate with processes 0.2-0.7 mm. long, but occasional fruits are
quite devoid of tubercles. As all intergrades exist between the smooth and
tuberculate states, in plants which are otherwise quite identical, and
sometimes even on the same plant, one cannot feel that the tuberculate
character is entirely reliable. Although the species probably belongs in
§ Adenoceras, it would be excluded from that section by the tuberculate
fruits if Pax & Hoffmann’s system should 8 premise followed.

In reconsidering my own species M. cre I must conclude that it
falls into a reasonable concept of M. akasedeh although its fruits are
smooth nearly without exception, while its leaves are comparatively small
and more obviously toothed. I now refer M. crenata to varietal status and
suggest as a third variety a taxon with unusually large leaves and more
obvious indument than typical.

KEY TO THE VARIETIES

Leaf-blades broadly ovate or deltoid-ovate, (8-) 12-30 X (6-) 8-23 cm.,
the petiole usually attached 2-4 cm. from lower margin, the margin entire
to crenate-undulate; petioles (6—) 8-21 cm. long; fruits sparsely tuberculate,

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XIII 383

the processes 0.2-0.7 mm. long, rarely lacking; plant often with slight
indument on vegetative parts and inflorescence

Stipules 1-3.5 cm. long, 5-10 mm. broad, glabrous or sparsely puberulent

(rarely long-pilose) and usually glabrate: branchlets and petioles glabrous

or soon glabrate; leaf-blades usually 12-24 —K 8-15cm.................

Gear Rae ane eae en eee seca 6 false Re RO eee 4a. var. graeffeana.

Stipules 2.5-5 cm. long, 12-20 mm. broad, copiously puberulent without;

branchlets and petioles soft-pilose, eventually glabrate; leaf-blades 15-30

4 EG tosic he get os oils ae WEE eae eee bas . var, major.

Leaf- oe deltoid, 8-14 & 5-8 cm., narrowly peltate (petiole attached 1-1.5

cm. from lower sie the margin conspicuously glandular-crenate-

ius petioles 5-10 cm. long; fruits smooth, very rarely with a few

minute panical tubercles; Shai glabrous throughout cee 4c. var. crenata.

4a. Macaranga graeffeana var. graeffeana
Macaranga graeffeana Pax & Hoffm. in Notizbl. Bot. Gart. Berlin 10: 384.
1928.

Shrub or tree up to 15 m. high, the branchlets and petioles often
sparsely spreading-pilose or puberulent when young (hairs 0.1-0.3 mm
long, rarely to 2 mm. long), soon glabrate; stipules oblong-lanceolate,
1-3.5 cm. long, 5-10 mm. broad, usually glabrous, sometimes sparsely
puberulent and soon glabrate, rarely copiously spreading-pilose; petioles
(6—) 8-16 (—21) cm. long, the leaf-blades broadly ovate or deltoid-ovate,
(8—) 12-27 cm. long, (6—) 8-20 cm. broad, rounded or rounded-truncate
at base and usually deeply (2-4 cm., rarely 1-6 cm.) peltate, acuminate
at apex, glabrous above, glabrous beneath or with the nerves pilose like
petioles and usually soon glabrate, obviously glandular beneath, the
primary nerves 6-8; inflorescences 3-9 cm. long, glabrous throughout or
the branches sparsely puberulent with hairs 0.1-0.2 mm. long, the larger
bracts obovate-lanceolate, 3-8 mm. long, patelliform-glandular; ¢ calyx
1—-1.2 mm. in diameter, glabrous (rarely with a few hairs when young),
scattered-glandular distally, 3-lobed nearly to middle; stamens 5-10, the
filaments 0.7-1.2 mm. long; 9 calyx glabrous or very sparsely puberulent
and glabrate; fruits 3.5-5 mm. long, 6-8 mm. broad, usually sparsely
tuberculate (processes 0.2-0.7 mm. long), rarely essentially or quite
smooth, copiously glandular, the styles 2, divaricate, 1.5—2.5 mm. long.

DIstRIBUTION: Fiji, thus far known only from Viti Levu and Vanua Levu,
occurring at elevations of 100-1150 m. in dense forest or in forest patches’ or on
open hillsides. The known specimens are from large shrubs or small trees 3-15 m.
high, the young inflorescences being reddish, the flowers yellowish, and the
fruits yellow-green. Recorded local names are: ndavo (upland Viti bes
vouotu (Mathuata), and tavotavo (Thakaundrove). The type is Graeffe 6
from Viti Levu without further data, Aeneid lost in the destruction of a
Berlin pacar no duplicate has been

Vitt Levu: Mba: Vicinity of ke eT Gillespie 3970 (Bish), 3988
(Bish, GH); sputhert slopes of Mt. Ndelainathovu, on the escarpment west of
Nandarivatu, Smith 4928 (A, US); Namosi: Summit of Mt. Naitarandamu,
Gillespie 3234 (Bish, GH), 3293 (Bish, GH, NY); summit of Mt. Vakarongasiu,
Gillespie 3285 (Bish, GH, K, NY, US); vicinity of Namosi, Gillespie 2625

384 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxxmr

(Bish); Naitasiri: Banks of the Wainimala at “Navusa,” Horne (K);
Rewa: Mt. Korombamba, Gillespie 2212 (Bish, GH, K, NY, US), 2367 (Bish,
GH, K, NY). Vanua Levu: Mathuata: Wainikoro, Greenwood 706 (K);
Seanggangga Plateau, in drainage of Korovuli River, vicinity of Natua, Smith
6812 (A, US); Thakaundrove: Mt. Mariko, Smith 421 (Bish, GH, K, NY,
US).

The cited material is quite constant in its basic characters with the
exception of two numbers. Gillespre 3285 is aberrant in having the
primary nerves of the leaf copiously pilose with hairs as much as 1 mm.
long. My no. 421, from Mt. Mariko, obviously comes from two different
plants; the fruiting specimens are glabrous and similar to material from
Viti Levu, but the specimens with staminate inflorescences have the
stipules, petioles, and primary nerves coarsely pilose with scattered
spreading hairs up to 1.5 mm. long. While these two numbers are not
entirely typical of var. graeffeana, they differ from it less markedly than
the two entities treated below as distinct varieties.

4b. Macaranga graeffeana var. major var. nov.

Arbor, ramulis petiolisque copiose puberulis (pilis 0.2-0.5 mm. raro
ad 1 mm. longis) demum subglabratis; stipulis oblongo-lanceolatis 2.5—5
cm. longis 12-20 mm. latis, extus copiose puberulis, intus subglabratis;
petiolis 11-18 cm. longis, foliorum laminis late ovatis, 15-30 cm. longis,
13-23 cm. latis, basi rotundato-truncatis et late (2-4 cm.) peltatis, apice
abrupte acuminatis, margine undulato-crenulatis, supra glabris vel costa
inconspicue puberula, subtus nervis primariis 7 vel 8 ut petiolo pilosis
glabratis, copiose glandulosis; inflorescentiis ? et infructescentiis 3—11 cm.
longis, ramulis inconspicue puberulis glabratis, bracteis dentatis 5-8 mm.
longis patelliformi-glandulosis, calyce haud puberulo pauciglanduloso;
fructibus copiose glandulosis tuberculatis, processis oblongo-conicis obtusis
0.2-0.4 mm. longis, stylis 2 divaricatis 1.5—2.5 mm. longis.

Vitt Levu: Mba: Vuninatambua, near Navai, alt. 750-900 m., March 21,
1941, Degener 14874 (A typr, Bish, K, NY, US) (mdavo; forest tree);
Nauwanga, south of Nandarivatu, alt. 750-900 m., Degener 14809 (A, Bish,
K, NY) (ndavo; forest tree).

The two collections here described as M. graeffeana var. major, both
from north-central Viti Levu, differ from typical material of the species
in their larger and copiously puberulent stipules, their larger leaf-blades,
and the more obvious and more persistent indument of their branchlets
and petioles.

4c. Macaranga graeffeana var. crenata (A. C. Sm.) comb. nov.
Macaranga crenata A. C. Sm. in Bishop Mus. Bull. 141: 86. fig. 44. 1936.
Tree to 10 m. high, glabrous throughout (or the inflorescences with a
few spreading hairs at base of flower-clusters); stipules lanceolate, 1—3
cm. long, 5-8 mm. broad; petioles 4-10 cm. long, the leaf-blades deltoid,
8-14 cm. long, 5-8 cm. broad, rounded-truncate at base and narrowly

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XIII 385

(1-1.5 cm.) peltate, slenderly acuminate at apex, conspicuously glandular-
crenate-undulate at margin, scattered-glandular beneath, the primary
nerves 5—7; 4 inflorescences 3-5 cm. long, the bracts obovate-lanceolate,
short-stipitate, 2.5-6 mm. long, patelliform-glandular, the calyx 1—-1.2 mm.
in diameter, 3-lobed, with scattered or apical sessile glands; stamens 5-7,
the filaments 0.8-1 mm. long; infructescences 1.5—3 cm. long, the fruits
4—4.5 mm. long, about 7 mm. broad, densely glandular, smooth or rarely
with a few (1 or 2 on occasional fruits) minute conical tubercles, the
styles 2, divaricate, 1-1.5 mm. long.

DistRIBUTION: Thus far known from only two collections, from Viti Levu
and Vanua Levu, at elevations of 650-1195 m., occurring in crest thickets. The
plants are trees 9 or 10 m. high; the type bears fruits and no. 4197 staminate
inflorescences, which have a rich pink calyx and white anthers. A local name
(no. 530) was recorded as kitimoku.

Vitt Levu: Mba: Summit of Mt. Koroyanitu, high point of Mt. Evans
Range, Smith 4197 (A, US). Vanua Levu: Thakaundrove-Mathuata
boundary: Crest of Korotini Range, between Navitho Pass and Mt. Ndelaikoro,
Smith 530 (Bish type, GH, K, NY, US).

The staminate specimen cited above agrees very well with the type of
this taxon and indicates that it may be limited to exposed crests and
ridges. Although the entity is fairly well marked by its deltoid, con-
spicuously crenate, and narrowly peltate leaf-blades, its completely
glabrous habit, and its smooth fruits, it does not seem specifically different
from M. graeffeana. Close examination indicates that an occasional fruit
has one or two minute tubercles, this fact providing another point of
similarity with typical M. graeffeana.

5. Macaranga (§ Adenoceras) marikoensis sp. nov.

Arbor gracilis ad 7 m. alta, ramulis, stipulis, petiolisque glabris, ramulis
lenticellatis; stipulis papyraceis vel submembranaceis oblongo-lanceolatis,
5—7 cm. longis, 12-18 mm. latis; petiolis 15—19 cm. longis, foliorum laminis
subcoriaceis in sicco fuscis late ovatis vel suborbicularibus, 13-17 cm
longis, 12—16.5 cm. latis, basi rotundatis vel subtruncatis et late (3—4.5
cm.) peltatis, margine copiose calloso-crenulatis (crenationibus 2 vel 3 per
centimetrum), apice abrupte cuspidatis (apice ipso circiter 5 mm. longo
obtuso), supra glabris, subtus nervis pilis ad 0.1 mm.. longis parce
puberulis mox glabratis et copiose luteo-glandulosis, nervis primariis 8—10,
nervis secundariis numerosis cum primariis supra leviter elevatis subtus
prominentibus, rete venularum intricato utrinque subprominulo; infructes-
centiis 7-12 cm. longis ubique glabris vel pedicellis brevibus parce
puberulis, bracteis caducis; fructibus magnis, 6-7 mm. longis, 9-11 mm.
latis, baie glandulosis, levibus vel interdum tuberculis conicis paucis

mm. longis inconspicue ornatis, stylis 2 divaricatis 2-3 mm. longis.

Vanua Levu: Thakaundrove: Mt. Mariko, alt. 600-866 m., Nov. 14,
1933, Smith 447 (Bish, GH, K, NY, US 1676109 type) (rote; elandee tree 7 m.
high, in dense forest).

386 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxx1m

The entity here described can scarcely be referred to any known species;
its relationship is doubtless with M. graeffeana, which it suggests in its
basic characters and the presence of occasional tubercles on the fruits.
However, M. marikoensis differs from the earlier species in its large stipules
and its much larger fruits, as well as in its suborbicular and short-cuspidate
leaf-blades, of which the nerves are more numerous and more prominent.
Another specimen from Mt. Mariko, Smith 421, is definitely not referable
to the new species; it has the small fruits and ovate leaves of M. graeffeana,
under which I have discussed it as a somewhat atypical representative.

6. Macaranga vitiensis Pax & Hoffm. in Pflanzenr. 63 [IV. 147. VII]:
337, 1914.

Macaranga sanguinea Gillespie in Bishop Mus. Bull. 91: 17. fig. 19. 1932.

Large shrub or small tree, the young branchlets and petioles spreading-
pilose with pale hairs 0.2-0.7 mm. long and usually soon glabrate; stipules
lanceolate, 1.5-7 cm. long, 5-15 mm. broad, copiously spreading-pilose
without (hairs 0.5-1 mm. long) or rarely essentially glabrous, glabrous or
soon glabrate within; petioles 5-23 cm. long, the leaf-blades ovate- or
deltoid-lanceolate, 13-55 cm. long, 4-20 cm. broad, rounded-truncate or
subcordate at base and usually obviously (1-6 cm.) peltate, rarely deeply
cordate and epeltate (probably a juvenile stage), gradually acuminate at
apex, undulate at margin, glabrous on both sides or with a few scattered
hairs 0.3-0.7 mm. long on nerves beneath, scattered-glandular beneath, the
primary nerves 6—8, the secondary nerves arising from costa numerous,
slightly curved; inflorescences slender, 3-10 cm. long, essentially glabrous
(or with a few minute scattered hairs on branches and bracts), the bracts
lanceolate-obovate, stipitate, 3-10 mm. long, patelliform-glandular; ¢
calyx cupuliform, 0.7-1 mm. long, deeply 3-lobed, with copious dorsal
or subapical glands; stamens 3-6, the filaments 0.6—1 mm. long; ? calyx
sometimes minutely pilose but soon glabrate; fruits copiously tuberculate,
the processes conical or oblong, obtuse or subacute, 0.5—1.5 mm. long, the
styles 2, divaricate, 1.5—3 mm. long.

DISTRIBUTION: Endemic, apparently limited to a small area in southern Viti
Levu, where it is often recorded as abundant, at elevations up to 200 m. The
species is said to be a large shrub or few-branched tree up to 5 m. in height,
occurring in forest; the flowers are greenish, the fruits yellow-green with
brown tubercles, and the styles reddish brown. A local name is ndavo.

As type of M. vitiensis, Pax & Hoffmann cite a specimen collected by “Leon”
in Fiji, without further locality. I find no record of a collector with this name
having worked in the region, and the amusing hypothesis occurs to me that
they may have misread the word “Levu.” It can be observed, for instance, that
some of the Graeffe specimens (at least in the British Museum) were distributed
with no data but the hand-written inscription “Viti Levu.” I have examined
several such sheets with the present matter in mind, and the word “Levu” (in
Graeffe’s hand?) can readily be taken for “Leon.” The type of M. vitiensis,
in the Berlin herbarium, has presumably been destroyed, and so my suggestion
cannot be verified, but it seems not unlikely that this type was a Graeffe specimen

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XIII 387

from Viti Levu; he is known to have collected in the southern forests of that
island, where this eee is frequent. The type of M. sanguinea is Gillespie
3625.4, cited belo

Vit1 Levu: Heke Vicinity of Ngaloa, Degener 15068 (A, NY), 15182
(A, Bish, K, NY, US); Naitasiri: Vicinity of Nasinu, Gillespie 3566 (Bish,
GH, US), 3567 (Bish, GH), 3625.4 (Bish Type of M. sanguinea, GH, K, NY),
3642.1 (Bish), Greenwood 1123 (A, US); vicinity of Tamavua, Gillespie 2087
(Bish), 2092 (Bish, GH); Suva Pumping Station, Degener & Ordonez 13764
(A, K, NY); sea Mt. Korombamba, Parks 20143 (Bish), Gillestie 2271
(A, Bish, GH), R. Parham 75 (BM), Vaughan 3324 (BM); Lami,
Meebold 16902 a ae Road, Suva,” Tothill 561 (K), 696 (K), 758 (K);
vicinity of Suva, Meebold 8162 (K), 16901 (Bish). Fiji, without definite
locality: Horne 1044 (GH, K), Yeoward 26 bis (K).

On the basis of the original description of M. vitiensis, I can only
conclude that this name must replace M. sanguinea. The leaf-shape alone
(the blades being usually substantially more than twice as long as broad)
distinguishes this entity from any other Macaranga in Fiji. Gillespie
apparently did not consider M. vitiensis, since it was placed by Pax &
Hoffmann in § Adenoceras, among the species with smooth fruits, whereas
M. sanguinea has conspicuously tuberculate fruits; the type of M/. vitiensis
was a Staminate plant and could not have been accurately placed in the
Pax & Hoffmann system. Gillespie is quite correct in stating that his
species cannot be placed in any existing section; the combination of
tuberculate fruits, patelliform-glandular bracts, and palmate venation is
not accounted for in Pax & Hoffmann’s table (in op. cit. 302). I do not
propose a new section for this species, because sectional criteria in the
genus need careful revision; it may be that § Adenoceras will be extended
to include forms with tuberculate fruits, such as the present species and
M. graeffeana, which above I refer to § Adenoceras on the grounds that its
fruits are sometimes essentially smooth and sometimes tuberculate.

7. Macaranga (§ Eumappa) harveyana (Muell. Arg.) Muell. Arg. in
DC. Prodr. 15 (2): 998. 1866; Seem. FI. Vit. 228. 1867; Drake, FI.
Polyn. Fr. 186. 1893; Hemsl. in Jour. Linn. Soc. Bot. 30: 192. 1894;
Pax in Bot. Jahrb. 25: 646. 1898; Burkill in Jour. Linn. Soc. Bot. 35:
54.1901; Pax & Hoffm. in Pflanzenr. 63 [IV. 147. VII]: 357. 1914.

Mappa harveyana Muell. Arg. in Flora 47: 467. 1864.
Tanarius harveyanus Kuntze, Rev. Gen. Pl. 2: 620. 1891.
Macaranga harveyana var. glabrata Pax & Hoffm. in Pflanzenr. 63 [IV. 147.

VIT]: 357. 1914

Tree to 10 m. high, the branchlets and petioles glabrous (in Fijian
specimens) ; stipules oblong-lanceolate, submembranaceous or papyraceous,
1.5—2.5 cm. long, 5-7 mm. broad, copiously spreading-pilose on both sides
with hairs to 0.3 mm. long, sometimes glabrate; petioles 8-25 cm. long,
the leaf-blades broadly ovate, 9-24 cm. long, 6-19 cm. broad, rounded at
base and deeply (1.5—5 cm.) peltate, slenderly acuminate at apex, in-
conspicuously crenulate at margin, with 7-10 primary nerves, puberulent

388 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxx

on the nerves above with hairs 0.1-0.3 mm. long and sometimes minutely
stellate-puberulent on surface, otherwise glabrous, similarly pilose beneath
or with a few scattered longer hairs, dispersed-glandular; inflorescences
5-14 cm. long, copiously puberulent or tomentellous (on branchlets,
bracts, and calyces) with hairs 0.2—-0.5 mm. long; bracts not patelliform-
glandular, those of 8 inflorescences ovate-deltoid, 2-4 mm. long, entire,
those of @ inflorescences often deltoid-lanceolate, 10-12 mm. long,
copiously fimbriate-dentate; ¢ calyx 0.7-1 mm. long, deeply 3-lobed
nearly to base, with a few distal glands; stamens (4—) 6-9, the filaments
0.5-0.7 mm. long; ¢@ calyx tubular, 2-3 mm. long, with narrow lobes;
fruits with several or many long flattened subulate processes, these
subacute, 3-8 mm. long, minutely puberulent; styles 3, ascending to
spreading, copiously papillose, 3-4 mm. long in flower, up to 10 mm. long
in fruit.

DisTRIBUTION: Fiji, Samoa, Tonga, and apparently eastward to the Society
Islands; the type is Harvey (K, staminate and fruiting sheets), from Vavau or
Lifuka, Tonga. In Fiji the species is definitely known only from Viti Levu and
Taveuni, where it occurs sparingly at low elevations, up to 400 m., as a tree up to
10 m. in height.

Vitt Levu: Serua: Waimbale, near Namboutini, Degener 15476 (A, Bish,
K, NY, US); Namosi: Vicinity of Namosi, Gillespie 2885 (Bish, GH).
TAVEUNI: Vicinity of Waiyevo, Gillespie 4708 (Bish, GH). Fiji, without definite
locality: U.S. Expl. Exped. (GH, US 66263 & 66264), Horne 472 (GH, K).

Macaranga harveyana and M., secunda are the only representatives of §
Eumappa occurring in our region; they are readily distinguished from the
other Fijian species by the presence of long processes on the fruits and by
the absence of patelliform glands on the inflorescence-bracts, The distribu-
tion of M. harveyana seems to extend from Fiji to the Societies, as stated
by Pax and Hoffmann, on the basis of available material. The type of the
species falls into Pax & Hoffmann’s var. glabrata, which is therefore a
superfluous name, to be replaced by var. karveyana if infraspecific taxa
are deemed desirable. The available Fijian material is fairly typical, but
variation within the species cannot be evaluated without examining more
Polynesian specimens. In addition to the type I have seen the following
Polynesian specimens (all US) that seem to belong here: Tonga: U. S.
Expl. Exped.; Samoa: U. S. Expl. Exped., Rechinger 837, Vaupel 259,
Christophersen 585, Christophersen & Hume 2448, Setchell 253; Raro-
tonga, Cook Islands: Parks & Parks 22304; Tahiti: U. S. Expl. Exped.
A specimen from Niue, Yuncker 9618, has the characteristic styles of the
species but has pilose branchlets and petioles, suggestive of those of J.
secunda; this may represent M. harveyana var. puberula Pax & Hoffm.,
typified by a Lister specimen from Tonga that I have not seen

In the Tongan and Samoan specimens with satisfactory ¢ flowers, the
stamens are 6—9 in number, but it should be noted that a Fijian specimen,
Gillespie 2885, usually has only 4 stamens, this reduced number being
more typical of M. secunda.

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XIII 389

8. Macaranga (§ Eumappa) secunda Muell. Arg. in DC. Prodr. 15 (2)*
996. 1866; Seem. FI. Vit. 228. 1867; Pax & Hoffm. in Pflanzenr. 63
[IV. 147. VII]: 354. 1914.

Tanarius secundus Kuntze, Rev. Gen. Pl. 2: 620. 1891.

Tree to 17 m. high, the branchlets and petioles copiously puberulent
with spreading hairs 0.1-0.3 mm. long or occasionally to 1.5 mm. long,
at length subglabrate; stipules papyraceous, lanceolate, 1.5-2.5 crn. long,
4-7 mm. broad, copiously spreading-puberulent on both sides, subglabrate;
petioles 5-18 cm. long, the leaf-blades broadly ovate, 12-20 cm. long, 7-14
cm. broad, rounded at base and deeply (2-5 cm.) peltate, long-acuminate
at apex, obscurely crenulate at margin, with 8-10 primary nerves,
puberulent on nerves on both sides (hairs 0.1-0.2 mm. long) and sometimes
minutely stellate-pilose on surface, the nerves beneath occasionally with
a few longer hairs, the glands scattered, inconspicuous; inflorescences 4-10
cm. long, copiously puberulent (on branchlets, bracts, and calyces) with
hairs 0.1-0.2 mm. long; bracts of ¢ inflorescences ovate-deltoid, 4-10 mm.
long, copiously fimbriate, not patelliform-glandular, the larger ones
stipitate; @ bracts presumably similar but not seen; 8 calyx infundibular,
0.5—0.6 mm. long, 3-lobed nearly to base, eventually subglabrate; stamens
3-5, the filaments 0.5-0.7 mm. long; 2 calyx persistently densely
puberulent; fruits with several or few flattened subulate processes, these
1.5—3 mm. long, puberulent; styles 2, divaricate, 1.5-2.5 mm. long.

DISTRIBUTION: Endemic, and thus far known from a few collections from
Viti Levu, Ovalau, and Vanua Levu, occurring at elevations up to 350 m. in
forest. Sparse data indicate the species as a tree 8-17 m. high, with the local

Viti Levu: Ra: Mataimeravula, vicinity of Rewasa, near Vaileka. Degener
15339 (Bish, GH, K, NY, US). Ovatau: U. S. Expl. Exped. (tyeE COLt..
GH, US 1944716), Seemann 395 (GH [as 359], K). Vanua Levu: Mbu
Southern portion of Seatovo Range, Smith 1519 (Bish, GH, K, NY, U
Fiji, without definite locality: Yeoward (K).

e
ds

390 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxxmI

secunda and has the type of fruit I believe to represent that species; at any
rate such a fruit effectively removes the Seemann specimen from M
harveyana. However, the occurrence of true M. harveyana in Fiji is
indicated by the several specimens I have cited under it, notably Horne
472, which has a fruit similar to that of Harvey’s Tongan type.

NOMEN NUDUM
MACARANGA MAUDSLAYI Horne, A Year in Fiji, 264, nomen. 1881; Baker
in Jour. Linn. Soc. Bot. 20: 371, nomen. 1883; Pax & Hoffm in
Pflanzenr. 63 [IV. 147. VII]: 394, nomen. 1914.

No description has been published for this binomial, and I have not
noted the name on any Horne specimens at Kew; therefore the name cannot
at present be referred to its proper synonymy.

EXCLUDED SPECIES
Mappa leptostachya Muell. Arg. in Linnaea 34: 198. 1865 = Macaranga
leptostachya Muell. Arg. in DC. Prodr. 15 (2): 1007. 1866 =
Tanarius leptostachyus Kuntze, Rev. Gen. Pl. 2: 620. 1891 =
Cleidion leptostachyum (Muell. Arg.) Pax & Hoffm. in Pflanzenr.
63 [IV. 147. VII]: 293. 1914.

The further synonymy of Cleidion leptostachyum, discussed by Croizat
in Occ. Pap. Bishop Mus. 18: 71. 1944, includes Cleidion vieillardi var.
vitiensis Muell. Arg. (in DC. Prodr. 15 [2]: 986. 1866) and Cleidion
degeneri Croizat (in Sargentia 1: 51. 1942).

Mappa macrophylla A. Gray ex Seem. in Bonplandia 9: 258, nomen.
1861 = Macaranga macrophylla Muell. Arg. in DC. Prodr. 15 (2):
1001. 1866 = Tanarius macrophyllus Kuntze, Rev. Gen. Pl. 2: 620.
1891 = Endospermum macrophyllum (Muell. Arg.) Pax & Hoffm.
in Pflanzenr. 63 [IV. 147. VII]: 418. 1914.

ACALYPHA L.

The taxonomic difficulties inherent in Acalypha throughout its vast
range are realized by all who have attempted identifications in the genus;
Croizat (in Occ. Pap. Bishop Mus. 18: 69-71. 1944) has discussed the
Fijian species and has concluded that Mueller and Pax & Hoffmann applied
too narrow a specific concept in their considerations of these species. With
this conclusion one must definitely agree. Croizat acknowledges the specific
identity of A. boehmerioides, A. grandis, A. rivularis, and presumably 4A.
wilkesiana, although the last is not discussed. The remaining six species
(and their varieties) accredited to Fiji are combined by Croizat under A.
insulana, with the implication that varietal categories might later prove
usefu

Examination of a considerable accumulation of herbarium material of
the A. insulana complex convinces me that natural populations, marked
by reasonable discontinuities, exist in Fiji. These populations are certainly
neither very stable nor readily definable, and probably they are inter-fertile,

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XIII 391

but I am loath to identify the various extreme forms as A. insulana without
further stipulation. Characters pertaining to leaf-size and shape and to
inflorescence-details appear too variable to be taxonomically very useful.
However, degree and type of indument are comparatively stable, and on
this basis one can recognize two primary patterns, here accepted as the
species A. insulana and A. repanda. Within each of these, populations of
reasonable constancy can be observed, here designated as varieties. Apart
from the species mentioned above, a very distinct novelty is here
described as A. amplexicaulis.

With the exception of A. boehmerioides, an introduced species, all the
taxa occurring in Fiji fall into Series III.H (Pantogynae-Pleurogynae) of
subgenus Exacalypha (now to be known as subgenus Acalypha) in the
system of Pax & Hoffmann (in Pflanzenr. 85 [IV. 147. XVI]: 13. 1924).
In that treatment one of the Fijian species, A. Jaevifolia, is placed in Series
III.G (Oligogynae), which has bisexual inflorescences. However, type
material of A. laevifolia seems to have unisexual inflorescences, although
rarely (as also in A. amplexicaulis) one or two sterile bracts of a pistillate
type occur toward the base of the staminate inflorescence; this, however,
does not indicate that the inflorescences are bisexual. All of the indigenous
Fijian species appear to have unisexual inflorescences and to be characteris-
tically monoecious; usually both staminate and pistillate inflorescences
occur on the same plant, but in herbarium specimens one frequently receives
the impression that the plants are dioecious. The occurrence or lack of
stalked glands on the pistillate bracts does not seem to be significant in the
Fijian population.

KEY TO THE SPECIES

Inflorescences bisexual, the distal flowers ¢; low annuals, the leaves with
slender petioles 1-6 cm. long and sibmicu ben ee ovate blades usually
TI 0G) ce a ane ar 1. A. boenmerioides.

Inflorescences unisexual; @ spikes cylindric, the bracts dentate; shrubs or trees.

Leaf-blades comparatively broad, less than twice as long as broad, palmate-
nerved, the 2 or 4 basal lateral nerves conspicuous, spreading, the distal
pair of these with several spreading tertiary nerves on the basal side, the
costa with spreading lateral nerves.

Petioles usually 6-12 (rarely 4-25) cm. long, not flattened at the
leaf-blades often cordate at base, sometimes rounded - obtu rns,
sip Et deg ot hen eee doe esi Soe. x wx. aca dw oh ee randis.

Petioles 1-5 cm. long, often flattened and broadened at oe of blade,
the leaf-blades usually broadly obtuse at base (or subcordate in f.
circinata), often reddish- or purple-maculate. ..._.. 3. A. wilkesiana.

Leaf-blades variously shaped, 2 or more times as long as broad, pinnate-
nerved, the basal secondaries not prominent nor with obvious basally
directed tertiary nerves.

Leaves obviously petiolate, not amplexicaul.

Leaf-blades narrowly obovate-lanceolate or subspatulate, 10-31 & 2-5.5
cm. (5—6 times as long as broad), gradually se to an abruptly
truncate or subcordate base, the petioles 0.5-3.5 cm. long. .......
NBGA TSE gaia dpe ok Ben dete oe dew comin etek Os ek a A. rivularis.

392 JOURNAL OF THE ARNOLD ARBORETUM | [voL. xxxur

Leaf- aon lanceolate to ovate or elliptic, 2-4 times as long as broad,
sub ate to acute at base but not gradually narrowed and abruptly
nei the petioles (0.5—) 1-7

Indument of young branchlets sual copious, spreading-pilose, the
hairs 0.2-1.2 mm. long and usually concealing the surface of
distal internodes; petioles ek ‘like young branchlets, the hairs
0.3-1.2 mm. lone: leaf-blades often spreading- pilose on both
surfaces or at least with an obvious indument on costa of lower
surface; rachis of ¢ and @ inflorescence usually copiously
puberulent or spreading-pilose, the ¢ perianth strigose-puberulent ;

? bracts usually spreading-pilose, rarely glabrate, the ovary and
fruit copiously strigose or spreading-pilose with hairs 0.2-0.5 mm
long (essentially glabrous only in var. subvillosa). ..5. A. insulana.
Indument of young branchlets comparatively close, strigose or puberu-
lent, the hairs 0.05—0.2 mm. long, often not obscuring the surface of
distal internodes; petioles pilose like young branchlets, usually
soon glabrate; leaf-blades essentially glabrous above and often

nerve-axils beneath, rachis of ¢ and @ inflorescence often
glabrous, sometimes puberulent or spreading-pilose, the

perianth puberulent or glabrous; ° bracts often glabrous, some-
times strigose, the ovary = fruit sist sales (hairs 0.1-

0.3 mm. long) or glabro A. repanda.
Leaves appearing subsessile, the piles 1-4 mm. Tong, the blades deeply
cordate at base and amplexicaul. 7. A. amplexicaulis.

1. Acalypha boehmerioides Miq. Fl. Ind. Bat. Suppl. 1: 459. 1860;
Muell. Arg. in DC. Prodr. 15 (2): 871. 1866; Seem. Fl. Vit. 226.
1867; Pax & Hoffm. in Pflanzenr. 85 [IV. 147. XVI]: 96. 1924.

Acalypha hispida sensu Benth. in Hook. Lond. Jour. Bot. 2: 232. 1843, non
Burm.

DistRIBUTION: A widespread species throughout parts of the Old World
tropics, growing sparsely in Fiji as a weed. The type was collected on Bangka,
east of Sumatra, presumably by W. S. Kurz under the fictitious name of “J.
Amann” (cf. van Steenis-Kruseman in Fl. Males. I. 1: 305. 1950, hae discussion
of Kurz’s Bangka collection). The Pacific material is referred to var. genuina
in the Pflanzenreich, but this will become var, boehmerioides if infraspecific
taxa are maintained.

Vitt Levu: Seemann 389 (BM, GH, K); Rewa: Nukulau Island, Barclay
(K), 5449 (BM). Fiji, without definite locality: U.S. Expl. Exped. (US 66219).

2. Acalypha grandis Benth. in Hook. Lond. Jour. Bot. 2: 232. 1843;
Seem. FI. Vit. 224. 1867.

Acalypha grandis var. genuina Muell. Arg. in Linnaea 34: 10. 1865, in DC.
Prodr. eae ers _ 1866; Pax & Hoffm. in Pflanzenr. 85 [IV. 147.

XVI]:
— prema Muell. Arg. in DC. Prodr. 15 (2): 807. 1866; Seem. FI.
Vit. 225. 1867.

Ricinocarpus consimilis Kuntze, Rev. Gen. Pl. 2: 617. 1891.
Ricinocarpus grandis Kuntze, Rev. Gen. Pl. 2: 618. 1891.

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XIII 393

DISTRIBUTION: In the Pflanzenreich eae species is assigned a wide distribu-
tion in Malaysia and the Pacific; in Fiji it has been collected on several of the
smaller islands and doubtless it also occurs along the coasts of the large islands.
It is typically found in coastal or lowland thickets, usually near sea-level but
sometimes up to 200 m. elevation, as a shrub or small tree 1.5-5 m. high, with
reddish to lavender styles; a local name of kalatimbuthi was applied to it on
Vanua Mbalavu. Our specimens belong to “var. genuina,” which becomes var.
grandis under present nomenclatural procedure. As cotypes, Bentham cites
Fijian collections of Hinds and Barclay, cited below, and also a Barclay
specimen from Amboina. The type of A. consimilis is an Exploring Expedition
specimen from Fiji, of which duplicates are here cited; the reduction of this
entity to synonymy by Pax & Hoffmann seems unquestionable.

Vitt Levu: Rewa: Nukulau Island, Barclay (BM, K cotype), 3452 (BM),
Tothill 748 (K). VAnua Levu: Mathuata or Thakaundrove: Undu
Point, Tothill 691 (Bish, K). Moata: Bryan 296 (Bish). Matuxu: Bryan 289

Bish). VANUA MBatavu: Central volcanic section, near Lomaloma, Smith
ve (Bish, K, NY). case ars Tothill 705 (Bish, K). LAkemBa: Tothill
704 (K). KAmBara: Moore 4 as Tothill 706 (K). Fiji, without definite
locality: Hinds (K cotype), U. S. Expl. Exped. (TYPE COLL. of A. consimilis,
GH, US 1944713), U.S. Expl. oe (GH, US 66220).

3. Acalypha wilkesiana Muell. Arg. in DC. Prodr. 15 (2): 817. 1866.

DISTRIBUTION: According to the Pflanzenreich treatment, A. wilkesiana has
a wide distribution as a cultivated plant, and apparently it is known only in
cultivation in Fiji. The type is an Exploring Expedition specimen from Fiji, of
which duplicates are here cited. Several horticultural forms have been recog-
nized, of which only f. circinata, in addition to the typical form, occurs in Fiji.
As cotypes of f. circinata, Mueller cites specimens of Seemann and the Exploring
Expedition, but I have not located any Seemann collection representing this
form. In Fiji the species is a shrub, cultivated under the name of kalambuthi
ndamundamu (kalambuthi being generic for Acalypha and the adjective referring
to the reddish leaves). In Ra, according to Degener, both the local forms are
known as ruru and the leaves are used medicinally.

KEY TO THE FORMS

Leaf-blades ovate or ovate-elliptic, (6—) 12-25 cm. long, (4—) 9-19 cm. broad,
broadly obtuse or sometimes rounded at base, obtusely cuspidate at apex.

3a. ilkesiana.

Leaf-blades suborbicular or reniform or broadly ovate, 3-11 cm. long and broad,
subcordate at base, often rounded at apex, somewhat flabellinerved. ......
ee ee ee eS Pho 5; TUR ey 3b. f. circinata.

3a. Acalypha wilkesiana f. wilkesiana
Acalypha wilkesiana Muell. Arg. in DC. Prodr. 15 (2): 817. 1866; Seem. FI.
Vit. 225. pl. 58. 1867; Pax & Hoffm. in eine 85 [IV. 147. XVI]: 153.
1924; Degener, Fl. Bae 2: Fam. 190. fig. 1
Acalypha tricolor Hort. ex Seem. Fl. Vit. 225, as rnin: 1867.
Ricinocarpus wilkesianus Kuntze, Rev. Gen. PI, 2: 618. 1891.
V : Ra: Rewasa, near Vaileka, Degener 15445 (A); Namosi:
Nanggara Island, H. B. R. Parham 276 (BM). Taveunt: Somosomo, Seemann

394 JOURNAL OF THE ARNOLD ARBORETUM | [voL. xxxm

392 in part (BM, GH, K). Fiji, without definite locality: U. S. Expl. Exped.
(TYPE COLL., GH., K, US 1944717 and 1944718).

3b. Acalypha wilkesiana f. circinata Muell. Arg. in DC. Prodr. 15
(2): 817. 1866; Pax & Hoffm. in Pflanzenr. 85 [IV. 147. XVI]:
154. 1924.

Acalypha circinata A. Gray ex Seem. Fl. Vit. 225, as synonym. 1867.

Vitr Levu: Ra: Rewasa, near Vaileka, Degener 15444 (A, NY). OVvALAu,
at least in part: U. S. Expl. Exped. (coTypE coLy., GH, K, US 66221).

4. Acalypha rivularis Seem. in Bonplandia 9: 258, nomen. 1861; Seem.
ex Muell. Arg. in Flora 47: 439. 1864; Muell. Arg. in Linnaea 34:
14. 1865, in DC. Prodr. 15 (2): 817. 1866; Seem. Fl. Vit. 225. 1867;
Pax & Hoffm. in Pflanzenr. 85 [IV. 147. XVI]: 168. 1924.

Ricinocarpus rivularis Kuntze, Rev. Gen. Pl. 2: 618. 1891.

DISTRIBUTION: This sharply marked species is apparently endemic, being
thus far known only from Viti Levu and Vanua Levu, at elevations from near
sea-level up to 850 m. It is a shrub up to 2 m. in height, characteristically
growing on the edges of rivers and streams where the land is inundated during
freshets, sometimes with its branches and leaves trailing in the water. The long
narrow leaves frequently have red nerves; the perianth-segments and stamens of
staminate flowers are greenish white, or the anthers may be reddish, while the
styles of pistillate flowers are usually red. Recorded local names are kandakanda
(Navua and Namosi regions), sotiura or sosotiura (Nandrau and Navai regions),
and sasariwai (in Mbua). The type is Seemann 391, cited below.

Vitt Levu: Near “Nandonga,” Horne 641 (K); Mba: Vicinity of Nandari-
vatu, Degener 14286 (A, Bish, K, NY, US); valley of Nggaliwana Creek, north
of the sawmill at Navai, Smith 5382 (A, US); Nandronga & Navosa
Nandrau, Degener 14921 (A, K, NY); Serua: Navua River at Namata,
Gillespie 3380 (Bish, GH); Serua or Namosi: ‘‘Navua and Namosi Rivers,”
Seemann 391 (BM, GH, K type); Namosi: Near Namuamua, on Wainikoro-
luva River, Gillespie 2960 (Bish, GH); vicinity of Namosi, Gillespie 2521
(Bish, GH, NY), 2874 (Bish, K); Naitasiri: Banks of Rewa River, Milne 292
(K); Viria and vicinity, Parks 20423 (Bish), 20455 (Bish); Tamavua district,

B. R. Parham 282 (BM); Suva — ha Degener & Ordonez 13780
(A, ie K, NY, US). Vanua Levu: Mbu Upper Ndama River valley,
Smith 1591 (Bish, GH, K, NY, US).

5. Acalypha insulana Muell. Arg. in Flora 47: 439. 1864.

Under the original publication of A. insulana, Mueller did not designate
a type, but he proposed three varieties, stipularis, pubescens, and
glabrescens, as varieties a, 8, and y respectively. Later, in the Prodromus,
varieties flavicans and villosa were added and the order of treatment was
changed. It is imperative that one of Mueller’s original varieties be
designated as the type variety of A. insulana. In the present treatment the
varieties stipularis and pubescens are taken as pertaining to the same taxon,
but var. glabrescens is treated as a synonym of A. repanda. Since var.

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XIII 395

stipularis is var. a in Mueller’s original treatment, it may justifiably be
taken as the type variety of his concept; under current nomenclatural
procedure the varietal epithet is to be replaced by repetition of the
specific epithet. Mueller’s first publication of these various narnes (in
Flora 47. 1864) was in a paper entitled “Neue Euphorbiaceen des
Herbarium Hooker in Kew,” and therefore the Kew specimens are to be
taken as the actual types (or cotypes, when more than one collection was
cited).

LocaL NAMES: Kalambuthi and kalatimbuthi are used throughout
Fiji to designate the genus as a whole, but perhaps they are more specifically
applied to A. insulana and A. repanda than to the more obviously
differentiated species.

KEY TO THE VARIETIES

Ovary and fruit obviously pilose; indument of young branchlets and petioles
copious; leaf-blades often pilose on blade of lower surface as well as on

costa.
— surface of leaf-blades sparsely pilose or glabrous, the a less
an on lower surface.......................5a. var. imsulana.
Both ees of leaf-blades very copiously spreading. pilose with ee 0.3-1.2
ong, these especially dense on costa; leaf-blades lanceolate to ovate,
7-25 X 2-10.5 cm., rounded or subcordate at base. ... .5b. var. flavicans.
Ovary and fruit glabrous: indument of young branchlets and petioles com-
paratively sparse, the hairs scattered, 0.5-1.2 mm. long; _leaf-blades
lanceolate, up to 14 & 4.5 cm., the indument usually limited to long hairs
on costa and in nerve-axils of lower surface .......... Sc. var. subvillosa.

5a. Acalypha insulana var. insulana

ere hie insulana Muell. Arg. in Flora 47: 439. 1864, in DC. Prodr. 15 (2):
818. 1866; Seem. Fl. Vit. 225. 1867; Pax & Hoffm. in Pflanzenr. 85 [IV.
147, XVI]: 165. 1924; Croizat in Oce. Pap. Bishop Mus. 18: 70. 1944.

Acalypha insulana var. stipularis Muell. Arg. in Flora 47: 439. 1864, in
Linnaea 34: 14, as A. insula var. s. 1865, in DC. Prodr. 15 (2): 818. 1866;
oar a Vit. 225. 1867; Pax & Hoffm. in Pflanzenr. 85 [IV. 147. XVI]:
166.

ee nen var. pubescens Muell. Arg. in Flora 47: 439. 1864, in
Linnaea 34: 14, as A. insula var. p. 1865, in DC. Prodr. 15 (2): 818. 1866;
Seem. Fl. Vit. 225. 1867; Pax & Hoffm. in Pflanzenr. 85 [IV. 147. XVI]:
166. 1924.

Acalypha insulana var. villosa Muell. Arg. in DC. Prodr. 15 (2): 818. 1866;

em. Fl. Vit. 225. 1867; Pax & Hoffm. in Pflanzenr. 85 [IV. 147. XVI]:
166. 1924.

Acalypha latifolia Muell. Arg. in DC. Prodr. 15 (2): 817. 1866; Seer. Fl. Vit.
225. 1867; Pax & Hoffm. in Pflanzenr. 85 [IV. 147. XVI]: 167. 1924,

Acalypha stipularis Engl. in Bot. Jahrb. 7: 462. 1886.

Ricinocarpus insulanus Kuntze, Rev. Gen. Pl. 2: 618. 1891.

Ricinocarpus latifolius Kuntze, Rev. Gen. Pl. 2: 618. 1891.

DISTRIBUTION: The typical variety of A. imsulana, at least as here interpreted
to include var. pubescens, is accredited to New Guinea as well as to Fiji by Pax

396 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxu1

& Hoffmann. The limits of the range of neither the species nor the variety can
at this time be stated, but it seems likely that A. imsulana in its typical form
occurs throughout the island-groups from Fiji to New Guinea, and perhaps
farther in each direction. Fortunately for nomenclatural purposes, all the type
collections involved are Fijian. In our region this variety is characteristic of
lowland thickets, woods, and dry forests, occurring most commonly near sea-level
but sometimes up to 600 m. elevation. It is a shrub or rarely a tree, 1-7 m
in height, with eeenish yellow staminate flowers and with styles aan are at
first white, becomin

Type material of the three varieties here combined under var. imsulana is
cited below. Two of the specimens involved in this typification are Seemann 392
and 393, which are composed of material from different plants and are therefore
always cited “in part.” Cotype collections of the three varieties were originally
designated as follows: var. stipularis, Harvey, Seemann 392 and 393, both in part;
var. pubescens, Seemann 393 in part, Milne 169 and 265; var. villosa, Vieillard 52
in the herbarium of Lenormand [University of Caen, France], U. S. Exploring
Expedition. The type of A. latifolia is an Exploring Expedition specimen; this
differs from more typical material in its sparser indument and in having its leaf-
blades obovate-elliptic and obtuse to subacute at base, but its basic characters
are those of A. insulana.

Vitr Levu: “Wainiloa River,’ Horne (K); without detailed locality, H. B. R.
Parham 245a (BM), 397 (BM); Mba: General vicinity of Lautoka, Degener &
Ordonez 13722 (A, Bish, K, NY, US), Degener 14989 (A, Bish, K, NY, US),
Greenwood 235A (A): slopes of the escarpment north of Nandarivatu, Smith
6067 (A, US); Nandronga & Navosa: Singatoka, Greenwood (Coll. H.
Phillips) 775 (K). Ovatau: U.S, Expl. Exped. (cotyPe COLL. of A. imsulana var.
villosa, GH, US 66222), U. S. Expl. Exped. in part (GH, US 66223), Milne 265
(K cotyreE of A. insulana var. pubescens); Lovoni Valley, Horne 172 (K); vicin-
ity of Levuka, Parks 20499 (Bish), Gillespie 4534 (Bish). Kanpavu: Namalata
Isthmus region, Smith 27 (Bish, GH, K, NY, US), 190 (Bish, GH, K, NY, US);
hills above Namalata and Ngaloa Bays, Smith 106 (Bish, GH, K, NY, US).
Vanua Levu: U.S. Expl. Exped. (GH); Mathuata: Southern base of Math-
uata Range, north of Natua, Smith 6786 (A, US); Mt. Numbuiloa, east of Lam-
basa, Smith 6538 (A, US); Thakaundrove: Hills south of Nakula Valley,
Smith 335 (Bish, K, NY). Taveunt: Vicinity of Waiyevo, Gillespie 4624 (Bish,
GH, NY); vicinity of Wairiki, Gillespie 4640 (Bish, GH, K). Koro: Eastern
slope of main ridge, Smith 1001 (Bish, K, NY). Ncau: Milne 169 in part (K

YPE of A. insulana var. pubescens). KAMBARA: Limestone formation: Smith
1252 ec GH, K, NY, US). Fiji, without definite locality: Harvey (BM, GH,
K corype of A. insulana var. stipularis), U. S. Expl. Exped. (type COLL. of A.
latifolia, ‘CHD. U.S, Expl. Exped. (US 66224), Seemann 392 in part (BM, GH,

E of A. imsulana var. stipularis), 393 in part (BM, GH, K COTYPE of
sna var. oe) 393 in part (BM, GH, K COTYPE of A. insulana var.
nae. Horne (K), 65 (K).

5b. Acalypha insulana var. flavicans Muell. Arg. in DC. Prodr. 15
(2): 818. 1866; Seem, Fl. Vit. 225. 1867; Pax & Hoffm., in Pflanzenr.

85 [IV. 147. XVI]: 166. 1924.
DIsTRIBUTION: Known from a few scattered localities in Fiji at low elevation
(up to 500 m.), occurring as a shrub in thin forest, with pink styles (Smith 6803,

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XIII 397

the only specimen with data). The type is an Exploring Expedition specimen
from Ovalau.

Vitt Levu: Namosi: Vicinity of Namosi, Gillespie 2830 (Bish). OvALAu:
U. S. Expl. Exped. (type cot. GH, US 66225); vicinity of Levuka, Gillespie
4402 (Bish, GH). Vanua Levu: Mathuata: Southern base of Mathuata
Range, north of Natua, Smith 6803 (A, US).

The very densely long-pilose upper surfaces of the leaf-blades distinguish
this taxon from the typical variety, but its value as a discrete entity is open
to question.

5c. Acalypha insulana var. subvillosa (Muell. Arg.) comb. nov.
Acalypha anisodonta Muell. Arg. in DC. Prodr. 15 (2): 818. 1866; Seem. FI.
Vit. 226. 1867; Pax & Hoffm. in Pflanzenr. 85 [IV. 147. XVI]: 167. 1924.
Acalypha anisodonta var. subvillosa Muell. Arg. in DC. Prodr. 15 (2): 819.
1866; Seem. FI. Vit. 226. 1867; Pax & Hoffm. in Pflanzenr. 85 [IV. 147.
XVI]: 167. 1924
Ricinocarpus anisodontus Kuntze, Rev. Gen. Pl. 2: 617. 1891.
DISTRIBUTION: Known with certainty only from Ovalau, Fiji. In proposing
the species A. anisodonta, Mueller did not designate a type of the binomial, but
his variety q was var. subvillosa, based on an Exploring Expedition specimen
from Ovalau. The disposition of the binomial therefore depends upon the placing
of var. subvillosa. Mueller’s second variety, var. subsericea, appears to me essen-
tially similar to typical specimens of A. repanda, discussed below.
OvaLau: U.S. Expl. Exped. (type coLy., GH), Milne 256 (K).

The variety has the long spreading hairs on the branchlets that are
typical of A. insulana var. insulana, but they are comparatively scattered,
while the indument on other parts of the plant, notably the ovary, is very
sparse or lacking. It seems likely that this taxon represents a hybrid form
between A. insulana var. insulana and A. repanda. Another specimen
that may also belong here, although its indument is even more sparse than
that of the type, is Greenwood 1079 (A, US) (Viti Levu: Mba: Mt. Evans
Range, alt. about 1050 m.).

6. Acalypha repanda Muell. Arg. in Flora 47: 439, 1864.

The entity here recognized as A. repanda differs primarily from A.
insulana in its much closer and usually sparser indument. While the
division of this entire population (A. imsulana sensu Croizat in Occ. Pap.
Bishop Mus. 18: 70. 1944) into two parts may seem arbitrary, there is
seldom doubt as to the position of material. In A. imsulana, as here defined,
the hairs of the young branchlets and petioles are conspicuous and spread-
ing, frequently exceeding 0.5 mm. in length; in A. repanda, on the
contrary, the indument of the young branchlets and petioles, if present at
all, is very close, with often appressed hairs of insignificant length (up to
0.2 mm.). The pubescence of other parts of the plant (leaf-blades, in-
florescences, perianth, ovary, etc.) correspondingly tends to be longer and
denser in A, insulana than in A. repanda. The extremes of this complex

398 JOURNAL OF THE ARNOLD ARBORETUM _[voL. xxxrIr

as to indument, e. g. A. insulana var. flavicans and A. denudata, are so
diverse that one hesitates to place them in the same species if any other
expedient can be found. The treatment here proposed, unsatisfactory as
it is, at least permits the recognition of two major groups of forms.

Acalypha repanda is essentially as variable in leaf-form and size as A.
insulana, but it is here rather arbitrarily divided into two varieties, based
primarily upon leaf-size and degree of pubescence.

Locat NAME: Kalambuthi is used throughout Fiji for this species.

KEY TO THE VARIETIES

Petioles 1-6.5 (-8) cm. long, the blades 9-23 % 3-10 cm. or rarely larger,
usually with 8-12 pairs of secondaries, the apex up to 20 mm. long; young
branchlets and sometimes petioles usually puberulent or very closely tomen-
tellous, the surfaces often obscured by the indument; ovary and young fruits
often copiously strigose with hairs 0.1-0.4 mm. long (fruits at length sub-
mC yt) ae ee eon ae ere oe rere rere er ere ee 6a. var. repanda.

Petioles 0.5-3.5 cm. long, the blades (4-) 5-14 X 1.5-6 cm., with 6-10 pairs of
secondaries, the apex 5-15 mm. long; young branchlets and petioles ob-
scurely puberulent or strigose (hairs not obscuring the surfaces), soon
glabrate; ovary and fruits sparsely strigose-puberulent with hairs 0.1-0.2 mm.
lane Or BIADIOUS «i dic veer re ates enh ap eee crass 6b. var. denudata.

6a. Acalypha repanda var. repanda

Acalypha repanda Muell. Arg. in Flora 47: 439. 1864, in Linnaea 34: 14. 1865,
in DC. Prodr. 15 (2): 819. 1866; Seem. Fl. Vit. 226. 1867; Pax & Hoffm.
in Pflanzenr. 85 [IV. 147. XVI]: 167. 1924.

Acalypha insulana var. glabrescens Muell. Arg. in Flora 47: 439. 1864, in

C. Prodr. 15 (2): 818. 1866; Seem. Fl. Vit. 226. 1867; Pax & Hoffm.
in Pflanzenr. 85 [IV. 147. XVI]: 166. 1924.

Acalypha anisodonta var. subsericea Muell. Arg. in DC. Prodr. 15 (2): 819.
1866; Seem. Fl. Vit. 226. 1867; Pax & Hoffm. in Pflanzenr. 85 [IV. 147.
XVI]: 167. 1924.

Ricinocarpus repandus Kuntze, Rev. Gen. Pl. 2: 618. 1891,

Acalypha insulana var. stipularis sensu Gibbs in Jour. Linn. Soc. Bot. 39: 169.
1909, non Muell. Arg.

Disrrrpution: As here interpreted, the typical variety of A. repanda has a
range of Samoa to New Guinea, or at least this is the range attributed by Pax
& Hoffmann to A. insulana var. glabrescens, which variety as to its nomencla-
turally typical element is here considered a synonym of A. repanda. In Fiji this
variety occurs from sea-level up to an elevation of 1150 m., in thickets, woods,
or various types of forest. The specimens are recorded as shrubs or small trees,
up to 5 m. in height; the staminate perianth is yellowish green to red, and the
styles become red. I have noted individuals growing side by side (my nos.
5132 and 5133) with yellowish and red perianths respectively, although no other

tion), all in herb. Kew; A. anisodonta var. subsericea, U. S. Exploring Expedition
(holotype presumably in De Candolle Herbarium).

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XIII 399

Vitt Levu: Mba: General vicinity of Lautoka, Greenwood 234 (K), 235
(K), 1092 (A), Degener & Ordonez 13721 (A, Bish, K, NY, US); vicinity of
Nandarivatu, Gillespie 4371 (A, Bish, US); Nauwanga, Degener 14482 (A, Bish,
K, NY, US), 14626 (A, Bish, K, NY, US), 14690 (A, Bish, K, NY, US); near
Naval, Gibbs 787 (GM); Ngglaiwana Creek near Nambuyasa, Gillespie 4138 (A,
Bish); western and southern slopes of Mt. Tomanivi, Smith 5132 (A, US),
5133 (A, US), 5746 (A, US); vicinity of ‘Panbenasol, Smith 4613 (A, US);
Nandronga & Navosa: H. B. R. Parham 245b (BM); Singatoka River
at Nude Horne 1001 (K); northern portion of Rairaimatuku Plateau, Smith
5425 (A, US); Serua: Mbuyombuyo, near Namboutini, Tabualewa 15564
(A, Bish, K, NY, US); Namosi: Vicinity of Namosi, Gillespie 2517 (Bish) ;

itasiri: Waindina River basin, MacDaniels 1033 (Bish); Waindra Creek,
igs 728 (K); 9 miles from Suva, sctiris 17037 (K); Kalambo, Tothill

5 (K); Nasinu, Tothill 749 (K). Ova U. S. Expl. Exped. (TYPE COLL.
: A. anisodonta var. subsericea, GH); Vicinity of Levuka, Gillespie 4464.5
(Bish). Kanpavu: Tothill 692 (Kk). Mt. Mbuke Levu, Smith 211 (Bish, GH,
K, NY, US). Vanua Levu: Thakaundrove-Mathuata boundary:
Crest of Korotini Range, Smith 543 (Bish, GH, K, NY, US); Thakaun-
drove: Southern slope of Korotini Range, Smith 516 (Bish, GH, K, NY,
US): Savu Savu Bay region, Degener & Ordonez 13953 (A, NY), 14025 (A,
Bish, K, NY, US). Taveunr: Somosomo, U. S. Expl. Exped. (coTYPE COLL. of
A. insulana var. glabrescens, GH). Natrat: Milne 179 (K cotypeE of A. insulana
var. glabrescens), 182 (K cotypeE of A. insulana var. glabrescens), Ncau: Milne
169 in part (K). Matuxu: Milne 129 (K cotypre of A. insulana var. glabre-
scens). Fiji, without definite locality: Harvey (BM, GH, K type of A. repanda),
Milne 46 (K), 417 (K cotype of A. insulana var. glabrescens), Seemann 393
in part (BM, K corype of A. insulana var. glabrescens), Horne (GH), 90a (K),
145 (K), 331 (K), Yeoward 12 (K), Gillespie 2529 (A, Bish).

Type material of the three entities here combined under A. repanda var.
repanda does not seem significantly to differ. The indument on the
branchlets and petioles of A. insulana var. glabrescens is perhaps slightly
the densest and most persistent, but the hairs composing it are minute as
contrasted with those of A. insulana. Type collections of A. repanda and
A. anisodonta var. subsericea, scarcely distinguishable from one another,
have in contrast a fugacious indument, long petioles, and lanceolate leaf-
blades that are rounded or subcordate at base.

Some of the upland specimens cited above, from the interiors of the large
islands, differ from typical lowland A. repanda in their more robust
branchlets, somewhat shorter petioles, and thicker leaf-blades with more
obvious marginal crenations and a tendency toward being acute at base.
However, no basic characters seem to separate these from typical material,
all the mentioned characters being highly variable.

6b. Acalypha repanda var. denudata (Muell. Arg.) comb. nov.

oe denudata Muell. Arg. in DC. Prodr. 15 (2): 819. 1866; Seem. FI.
Vit. 1867; Pax & Hoffm. in Pflanzenr. bid [IV. 147. XVI]: 167. 1924.
Acalypha “oevifolia Muell. Arg. in DC. Prodr. 15 (2): 853. 1866; Seem. FI.
1867; Pax & Hoffm. in Pflanzenr. 85 [IV. 147. XVI]; 112. 1924,
ee denidotis Kuntze, Rev. Gen. Pl. 2: 617. 1891.
Ricinocarpus laevifolius Kuntze, Rev. Gen. Pl. 2: 618. 1891.

400 JOURNAL OF THE ARNOLD ARBORETUM [voL. xXxxtII

Acalypha insulana var. glabrescens sensu Gibbs in Jour. Linn. Soc. Bot. 39:
169. 1909, non Muell. Arg.
Acalypha repanda sensu Gibbs in Jour. Linn. Soc. Bot. 39: 170. 1909, non

DISTRIBUTION: The smaller-leaved and earlier glabrate variety of A. repanda,
here designated as var. denudata, has been recorded (as the two species con-
cerned in the synonymy) only from Fiji. It occurs on several of the islands at
elevations up to 1120 m., on open hillsides or in woods or various types of forest.
The specimens were taken from compact or slender shrubs or trees 1-5 m. high,
the staminate perianth being greenish white to pale yellow or pink-tinged and
the styles pale pink to red. This variety is characteristic of the comparatively
dry hills of Mathuata Province, and it also occurs freely in the uplands of
Viti Levu.

Two species are concerned in the synonymy listed above, each without varieties
and each typified by a single Exploring Expedition collection. Although Mueller
did not mention localities for these collections, data on the available duplicates
show that they both came from Vanua Levu.

Vitt Levu: Graefe 30 (BM); Mba: Mt. Evans Range, Greenwood 1242
(US); Korovou, east of Tavua, Degener 14955 (A, Bish, K, NY, US); slopes
of the escarpment north of Nandarivatu, Gibbs 709 (BM), Tothill 733 (K),
Smith 6036 (A, US); vicinity of Nandarivatu, Gibbs 589 bis (BM), 590 (BM,
K), 656 (K), Parks 20558 (Bish), Gillespie 3715 (Bish, GH, NY), Tothill 746
(K), Degener & Ordonez 15541 (A, Bish, K, NY, US), Vaughan 3385 (BM);
slopes and summit of Mt. Nanggaranambuluta [Lomalangi], Gillespie 3395
(Bish), 4339 (Bish, GH, K, NY), Degener & Ordonez 15546 (A, Bish, K, NY,
US), Smith 5679 (A, US), 6314 (A, US); ridge between Mt. Nanggaranambuluta
and Mt. Namama, Smith 5005 (A, US); Serua: Navua River at Namata
Rapids, Gillespie 2949 (Bish); Namosi: Mt. Naitarandamu, Gillespie 3365
(Bish); Nanggarawai village on Wainikoroluva River, Gillespie 3224 (Bish,
GH); vicinity of Namosi, Gillespie 2839 (A, Bish, GH), 2868 (A, Bish); be-
tween Namuamua and Namosi, Gillespie 2955 (Bish, GH), 3057 (Bish, GH, K,
US); Rewa: Vicinity of Suva, Meebold 17066 (Bish). Ovatau: Vicinity of
Levuka, Parks 20488 (Bish), 20490 (Bish). WaKAya (?): Milne 324 (K).
Vanua Levu: U. S. Expl. Exped. (tyre cout. of A. laevifolia, GH, US 1944715);
Mathuata: U. S. Expl. Exped. (type cot. of A. denudata, GH, US
1944714); Seanggangga Plateau, vicinity of Natua, Smith 6751 (A, US); slopes
and summit of Mt. Numbuiloa, east of Lambasa, Smith 6371 (A, US), 6528
(A, US), 6544 (A, US), 6547 (A, US).

The two concepts which are here combined as a variety of A. repanda
are separable from one another only in matters of minute degree; A.
denudata is characteristically (as occurring in Mathuata) entirely glabrous
as to its inflorescence-rachis, perianth, and ovary, whereas A. laevifolia
usually has these parts strigose or faintly puberulent. In some cases it is
difficult to separate the cited specimens from var. repanda, but in general
the two varieties I propose to retain under A. repanda are reasonably
recognizable.

7. Acalypha amplexicaulis sp. nov.

Frutex monoicus ad 2 m. altus partibus juvenilibus et interdum bracteis

1952] SMITH, STUDIES OF PACIFIC ISLAND PLANTS, XIII 401

2 parce pilosis exceptis ubique glaber, stipulis lanceolatis 4-5 mm. longis
caducis, ramulis gracilibus teretibus pallide brunneis; foliis valde amplexi-
caulibus, petiolis leviter canaliculatis 1-4 mm. longis, laminis chartaceis
in sicco supra viridibus subtus nervis subrubris, lanceolatis, 7-15 cm.
longis, 2-4 cm. latis, basi sinu ad 1.5 cm. profunde cordatis et auriculatis,
d apicem acutum vel calloso-mucronulatum gradatim angustatis, margine
crenulatis (crenationibus 1 vel 2 per centimetrum saepe incrassatis),
pinnatinerviis, costa supra elevata subtus prominente, nervis secundariis
utrinsecus 8-12 utrinque elevatis, 3-5 paribus infimis e basi laminae
radiatis, aliis patentibus curvatis, rete venularum supra plano vel immerso
subtus prominulo; inflorescentiis unisexualibus axillaribus solitariis gracili-
bus, superne @, inferne ¢ ; inflorescentiis ¢ sub anthesi 7-13 cm. longis,
pedunculo ad 4 cm. longo, rhachi inferne bracteas @ steriles 1 vel 2
interdum gerente, bracteis ¢ deltoideis subacutis 2-3 mm. longis, inter-
nodiis sub anthesi 3—5 mm. longis, floribus ad 10 in fasciculis aggregatis sed
mox caducis, pedicellis gracilibus circiter 0.5 mm. longis; perianthii lobis
deltoideis ad 0.8 mm. longis, staminibus 8, filamentis et antherae loculis
pendulis circiter 0.3 mm. longis; inflorescentiis ? quam @_ paullo
brevioribus longe pedunculatis, internodiis spicarum longis, bracteis
reniformibus 4-6 mm. latis interdum distaliter parce pilosis (pilis ad 0.4
mm. longis) plerumque glabris glandulas stipitatas numerosas margine vel
intus marginem versus saepe gerentibus, 7—11-denticulatis, dentibus
subacutis apice rubro-callosis; sepalis 3 deltoideis acutis 0.7-0.8 mm.
longis, ovario glabro, stylis ad 5 mm. longis pinnatim 8—10-lacinulatis.

Vitt Levu: Mba: Northern portion of Mt. Evans Range, between Mt.
Vatuyanitu and Mt. Natondra, alt. 700-900 m., May 9, 1947, Smith 4300 (A
TYPE, US) (timbothe; shrub 2 m. high, in crest thickets).

The new species is unique in our region for its deeply cordate and
amplexicaul subsessile leaves, and it is further characterized by its
strikingly glabrous habit. In Pax & Hoffmann’s key it seems closest to A.
denudata (here treated as A. repanda var. denudata), which is perhaps its
closest relative, but the distinguishing foliar characters of A. amplexicaulis
are very pronounced.

ENDOSPERMUM Benth.

Endospermum robbieanum A. C. Sm. in Bishop Mus, Bull. 141: 82.
fig. 42. 1936.
Vanua Levu: Mathuata: Seanggangga Plateau, in drainage of Korovuli
River, vicinity of Natua, alt. 100-200 m., Smith 6716 (A, US) (freely branched
tree 10 m. high, in patches of forest in open rolling country).

The second known collection of the species, obtained not far from the
type locality (Wainunu River region near south coast of Vanua Levu),
agrees excellently in foliage with the type; its leaf-blades are inclined to be
rounded or even faintly subcordate at base. Number 6716 bears staminate
inflorescences, which are here described:

402 JOURNAL OF THE ARNOLD ARBORETUM _ [vot. xxx

Staminate inflorescences axillary toward apices of branchlets, paniculate,
at anthesis 6—8 cm. long, the peduncle 1—1.5 cm. long, the lowermost lateral
branches about 1.5 cm. long; indument of peduncle, rachis, and branches
copious, fulvous, the hairs many-branched from base, 0.2-0.4 mm. long;
bracts broadly deltoid, obtuse, the largest about 2.5 & 4 mm., the ultimate
bracteoles about 1 mm. long; flowers subsessile in glomerules of 2—4 or
solitary in axils of bracteoles; calyx submembranaceous, cupuliform, at
anthesis 2—2.5 mm. long and slightly broader, subentire at apex, copiously
and minutely tomentellous without, glabrous within; disk carnose,
irregularly angled; stamens 13-17, minate to column usually in 2 series,
exserted at anthesis, the filaments minute, the anthers 0.7-1 mm, in
diameter.

DEPARTMENT OF BOTANY,

NATIONAL MuSEuM,
SMITHSONIAN INSTITUTION.

1952] THE ARNOLD ARBORETUM DURING FISCAL YEAR 403

THE ARNOLD ARBORETUM DURING THE FISCAL YEAR
ENDED JUNE 30, 1952

Horticulture. — As a result of the trip that the horticulturist made to
Europe last spring, nearly 500 species and varieties of woody plants have
been received from various places visited by him. Some of these are new
to horticulture in North America, others are new to the collections of the
Arnold Arboretum, and some, not new, have disappeared from our col-
lections at some previous time. One of the interesting shipments was of 32
varieties of Ribes, Philadelphus, Deutzia and Weigela from the Proeftuin
te Boskoop in Boskoop, Holland. These clones represent what the Dutch
have finally agreed on as the true named clone for these varieties. They will
be grown here and checked with what we have to see if they agree.

During the spring approximately 255 species and varieties have been
added to living collections at the Arboretum. This includes a spring plant-
ing list of 384 plants. Also a substantial beginning has been made towards
the replanting of Peter’s Hill, of about 55 different varieties of crab apples
which were planted there this spring as the forerunner to making this an
enlarged crab apple planting, 17 of them Dr. Sax’s new hybrids.

A three-hundred foot wisteria arbor was erected from white cedar wood
this last winter, adjacent to the shrub collection and along the Arborway
wall. This was made necessary by the increased land requirements of the
Bussey Institution, on whose land the wisteria arbor was formerly located.

The collections of the Arboretum are now recorded on 106 maps. Of
these 40 were rechecked last year and 18 were redrawn. Approximately
900 display labels were made and 600 machine labels were made.

Over 1,000 black and white photographs were taken, developed and
enlarged to prints 5” x 7”. This includes about 800 taken by the horti-
culturist while in Europe. The remainder were taken by Mr. Howard, who
also was responsible for the developing and printing of the entire lot. He
also made several hundred post cards, pictures of valued specimens here in
the Arboretum. About 300 color pictures were taken by Mr. Howard and
about 800 by the horticulturist in Europe, all of which have increased
our files materially.

Two display cases have been made and equipped with fluorescent light-
ing to show large Echtachrome transparencies of interesting Arboretum
scenes. All the pictures for these cases were taken by Mr. Howard. Each
case has 20-25 large color pictures and it is hoped two more cases will be
made in order to illustrate the four seasons in the Arnold Arboretum. Such
a display is ideal for showing ‘‘uut of season” visitors some of the beauties
of the Arboretum when the plants are at their best.

During the fiscal year 328 shipments of plant materials, including 982
species and varieties, have been sent to cooperating institutions or indi-
viduals in sixteen countries. The Arboretum received 1194 species and

404 JOURNAL OF THE ARNOLD ARBORETUM _[vot. xxxim

varieties. Most of these were obtained from England and Holland as the
result of Dr. Wyman’s trip to Europe last year. Many were also received
as seeds from Argentina and Japan. A total of 12,857 plants were success-
fully propagated; 153 by air layering, 9,906 by cuttings, and 2,447 by
grafting or budding. Well over 600 different numbers or pans of seed were

The regular number of bulletins have been published. The one on air
layering has created wide interest. A résumé of this work was also pub-
lished in the Journal of the Royal Horticultural Society. “Trees for Ameri-
can Gardens,” written by the horticulturist, was published by Macmillan
and Company of New York in November 1951.

Mr. Richard Fillmore left the Arboretum for a more lucrative position
in commercial work, and was succeeded by Mr. Lewis Lipp as chief propa-
gator. One of Mr. Lipp’s first steps was to invite members of the Federated
Garden Clubs to assist in the propagation work and to learn more about
the various methods of propagation. The response was gratifying, and the
program is off to a good start.

Case Estates at Weston.—A small tree demonstration plot was
started last year and now contains 60 different kinds of small trees. The
object is to display here in this one area some of the better small trees
including both those which are generally known and unknown, so they
can be compared closely for growth and habit one with the other. This
has great educational value for street tree superintendents as well as those
interested in planting the home grounds. In the Ground Cover Demonstra-
tion Plot we now have growing 125 different kinds of ground covers.

The two saran cloth houses again proved their worth during the dry
summer of 1951. Plants grown in these, and hardened off properly in the
early fall, seem to do far better than plants grown in the open nursery
where water is difficult to obtain. It is in one of these cloth houses that
we have the 239 plants which are being grown under Post Entry Quaran-
tine regulations with the permit from the U.S.D.A. Also in the same house
are the majority of plants, cuttings and grafts which have been obtained
from Europe this past winter.

Much of the seed germination work of the Arboretum is being done this
year —as last — at Weston. Included in the plants being tried this year
are seeds collected from certain specific places in northern Honshu, Japan
and also from the higher altitudes and colder, drier spots of Argentina.
Some of these undoubtedly will not yield anything new or hardier, but,
because they have been specifically collected in certain areas, some packets
may yield plants of increased hardiness, and if this is the case it will
make the entire project very worth while.

A young orchard of 38 Malus sikkimensis has been set out this year
for the purpose of growing seeds of this rare crab apple to be used in the
production of dwarfing understocks for commercial apple growers. Once
the stock is widely distributéd in this country, this group of trees will un-
doubtedly be discarded.

Approximately 82 clones of the Glenn Dale Hybrid azaleas were growing

1952] THE ARNOLD ARBORETUM DURING FISCAL YEAR 405

in the saran cloth houses last summer, as a test for hardiness. Because of
premature cold spells prior to November 1, 1951, all clones but about a
dozen were actually killed, thus proving that these plants, on the whole,
are not hardy here in New England.

Experimental Horticulture. — The production of polyploid plants
often results in larger flowers and more sturdy growth. Dr. Sax has found
that when the artificially induced tetraploids are crossed with diploids, the
resulting triploids are often even larger and more vigorous than the tetra-
ploids. A new giant triploid Forsythia has been propagated for distribu-
tion, and has been named the “Farrand Forsythia” in honor of our con-
sulting landscape gardener, Mrs. Beatrix Farrand.

Transgressive segregation in the second generation progeny of a Lonicera
species hybrid has produced a promising compact low-growing bush honey-
suckle. More hybrids have been obtained by crossing Malus Sargenti with
other ornamental apple species. Most of these resemble the Sargent Crab
in growth habit, but are more vigorous with larger flowers.

The use of tree lilac rootstocks as rootstocks for Syringa vulgaris seems
to be successful in producing a tree form of the common lilac. Dwarfing
rootstocks to produce small ornamental trees are being developed for
apples, hawthorns, peaches and plums.

Comparative Morphology. —In the Herbarium of the Arnold Ar-
boretum, there are collections of many strange and aberrant genera from
Northern Australia, New Guinea, and adjacent areas northward into China
and Japan. Professor Bailey’s investigations of a succession of these genera
in collaboration with Dr. A. C. Smith and others indicate that they are
relics of an ancient, diversified, woody, dicotyledonous flora. A majority
of them are characterized by their retention of relatively primitive internal
structures. Intensive investigations of adequate collections of them promise
to throw a flood of new light upon the morphological characteristics of.
ancestral dicotyledons and in time upon the great mystery of the origin
of the flowering plants or angiosperms. During the year, Professor Bailey
has made comprehensive studies of Calyptosepalum from Sumatra, Nou-
huysia-and Idenburgia from New Guinea and a new relic tree collected
by Dr. Smith in Fiji. Nouhuysia and Idenburgia prove to be congeneric
and Dr. Smith’s plant appears to be closely related to Calyptosepalum
which clearly does not belong in the Santalaceae. Ing. Domingo Cozzo,
a Guggenheim Fellow from Argentina, is spending a year in residence in
order to work with material in the slide and wood collections. Mr. Chi
Ling Chen, a candidate for the doctorate, has initiated a comprehensive
investigation of accumulated collections of the Sapotaceae. Professor Rhoda
Garrison of Wellesley College is devoting a year to investigation of the
structure and development of buds in Magnolia, Liriodendron, Akebia and
Pterocarya.

The Herbarium. — During the year 6,274 specimens were mounted
and inserted into the herbarium, making a total of 664,989 sheets. In-
coming specimens totalled 16,236, over half of which (8,919) were received
on the basis of exchange. Nearly 5,000 specimens were obtained by pur-

406 JOURNAL OF THE ARNOLD ARBORETUM [vot. xxxmm

chase or subsidy, and close to 2,500 were recorded as gifts. Of the total
number, 6,797 specimens were Asiatic in origin, 5,906 came from the
Pacific Islands and Australia, 1,020 represented European species, 563 were
African, and only 1,850 originated in North America. The larger and
more interesting incoming exchanges included 2,994 Malaysian specimens
from the British Museum, London, 1,714 Indonesian specimens from the
Herbarium Bogoriense, Bogor (Java), and 944 Soviet specimens from the
V.L. Komarov Botanical Institute of the Academy of Sciences, U. S. S. R.,
at Leningrad. By purchase 1,287 Japanese specimens were obtained from
Prof. H. Hara of Tokyo.

Outgoing specimens numbered 41,177, sent mostly as exchange to twenty-
eight institutions. Of these, 5,306 specimens were sent to eleven American
institutions and over 30,000 specimens to seventeen foreign institutions.
Also, 5,257 mounted specimens were transferred to the Gray Herbarium.
Herbarium sheets totalling 3,756 specimens were loaned by the Arboretum
to workers at twenty-three institutions. Of these, twenty-six loans com-
prising 3,531 specimens were sent to sixteen American institutions, while
twelve loans totalling 225 specimens were sent to seven foreign institutions.
The thirty-four incoming loans from fifteen institutions for study by our
staff members totalled 3,245 specimens. Of these, twenty-three loans total-
ling 2,882 specimens came from nine Ametiean institutions, and eleven
loans totalling 363 specimens from six foreign institutions.

A grant of $5,000 from the Guggenheim Foundation enabled Dr. Merrill
to spend six months working in Europe. Most of his time was spent at the
British Museum, where he selected for the Arboretum a large number of
available duplicates of the Carr New Guinea collection. He also visited
the herbaria at Kew, Edinburgh, Leiden, and Brussels, obtaining critical
notes on 1800 types. At Brussels he studied various authentic Roxburgh
specimens in the hitherto little-known collection of that author preserved
in the Martius herbarium. About fifty per cent of the approximately 1,350
specimens turned out to be isotypes. Also during the year he completed
his manuscript, “William Jack’s Genera and Species of Malaysian Plants.”
Prof. Johnston continued his work in the Asiatic Boraginaceae, completing
his study of Lithospermum and its related genera. The pollen of the
family was intensively studied and showed new and interesting characters
which could be used in delimiting species. The last part of the year was
spent working in the British herbaria. Dr. Kobuski continued his work
on the Asiatic Theaceae and initiated work on Adinandra for the Flora
Malesiana. Dr. Perry pursued her studies of the Papuasian flora, complet-
ing work on Elatostema and beginning the study of the Euphorbiaceae.
Dr. Hu contributed two ‘Notes on the Flora of China,’ and continued
her study of the genus Philadelphus.

The Library. — The library now has 48,098 bound volumes on its
shelves, this represents an increase of 370 volumes; some were gifts, others
were purchased or received in exchange for our publications. There were
197 pamphlets catalogued and filed; these were all gifts of the authors.
Our pamphlet collection now numbers 15,064. Four-hundred-twenty cata-

1952] THE ARNOLD ARBORETUM DURING FISCAL YEAR 407

logue cards were prepared, typed and filed in the main catalogue and 2,023
cards were added to the Gray Herbarium species cards.

Many inquiries of a bibliographical or biographical nature were answered
as were many requests for photostats, microfilms and photo prints. About
1100 photographs were added to the collection, these mainly photographs
of plants growing in the Arboretum, but many were taken by our Horti-
culturist on his European travels.

Four-hundred-ninety-seven books were loaned to or borrowed from
other libraries.

Bibliography of the Published Writings of the Staff and Students
July 1, 1951 — June 30, 1952
BarLey, I. W. Biological processes in the formation of wood. Science 115:

255-259. 1952.
Cooperation versus isolation in botanical research. Biologia 2: 126-133.
1

The use and the abuse of anatomical data in the study of phylogeny
and classification. Phytomorphology 1: 67-69. 1951.
FILLMORE, R1cHARD H. Chinese air layerage. Horticulture 29: 297. 1951.
aking summer-wood cuttings. Horticulture 29: 251, 264. 1951.
——— Review of woody plant propagation. Am. Nurseryman 94(11): 7-8, 65,
66. 94(12): 10, 11, 42, 43, 55.
The vegetative propagation of Viburnums. Arboretum Bull. 14: 17-18,
25. 195%
Gowpa, Mart. The genus Pittosporum in the Sino-Indian region. Jour. Arnold
Arb, 32: 263-301, 303-343. 1951.
Hu, SHIU-YING. Notes on the flora of China, I. Jour. Arnold Arb. 32: 390-402.
pl. 1. 1951. II. 33: 166-176. pl. 1, 2. 1952.
saaeieea Ivan M. Studies in the Boraginaceae, XXI. Sino-Indian species of
ma. Jour. Arnold Arb. 32: 201-225, 344-368. 1951.
Stale in the Boraginaceae, XXII. Noteworthy species, chiefly Asian
South American. Jour. Arnold Arb. 33: 62-78. 1952.
osu, CLARENCE E. Studies in the Theaceae, XXIII. The genus Pelliciera.
Jour. Arnold Arb. 32: 256-262. 1951.
— Studies in the Theaceae, XXIV. The genus Sladenia. Jour. Arnold Arb.
32: 403-408. pl. 1. 1951.
Studies in the Theaceae, XXV. The genus Anneslea. Jour. Arnold Arb.
33: 79-90. 1952.
Studies in the Theaceae, XXVI. The genus Visnea. Jour. Arnold Arb.
33: 188-191. 1952
Theaceae. /n: Sm ith, A. C. Studies of Pacific Island plants, XI. Further
notes on Fijian lie: plants. Jour. Arnold Arb. 33: 97-98. 1952.
MerriLL, E. D. Notes on Xanthostemon F. Mueller and Kjellbergiodendron
Burret. Jour. Ae. ‘Arb. 33: 150-165. 1952.
— On the identity of the genus Baranda Llanos. Jour. Arnold Arb. 32:
409-411. 1951.
Readjustment in the nomenclature of Philippine Eugenia species.
Philippine Jour. Sci. 79: 351-430. 1950.

408 JOURNAL OF THE ARNOLD ARBORETUM _ [voL. xxxim

MerrRILL, E. D. & STEENIS, C. G. G. J. VAN. Reductions of two Malaysian genera
of Euphorbiaceae. Webbia 8: 405-406. 1952.

Perry, Lity M. Plantae Papuanae Archboldianae, XX. Jour. Arnold Arb. 32:
369-389. 1951.

SAx, Kart. The Arnold Arboretum during the fiscal year ended June 30, 1951.
Jour. Arnold Arb. 32: 412-416. i;

Photosynthetic energy via agriculture. Proc. Am. Acad. Arts & Sci. 79:

205-211. 1951.

Population problems in world development. Jn: Social progress through

technology 4-6. 1951.

& LurppoLtp, Henry. The effect of fractional X-ray dosage on the
frequency of chromosome aberrations. Heredity 6: 127-131. 1952

SCHWARTEN, LazELia. Index to American botanical literature. Bull. Torrey Bot.
Club 78: 353-362; 431-439; 472-483. 1951. 79: 96-106; 195-203;
273-283. 1952.

translator. Introduction of Chinese ligneous plants into America.
Arboretum Bull. 14(4): 15, 16, 30. 1951.

VERDOORN, FRANS. Foreword to reprint of C. Darwin’s Journal of Researches
(18 39), Pallas 2: i. 1952.

From empirism to applied science in pharmaco-botany, with some remarks

on the need for institutions for certain oe of the history of science.

Am. Jour. Pharm. Educ. 15: 338-348. 1

L’arboretum moderne. Nat. Canad. 189-197. 1952.

On the need for international visitors’ research stations in certain areas

of the tropics. Chron. Bot. 12: 226-230.

Problemen der botanische geschiedschrijving. Vakbl. voor Biologen 31:
201-209. 1951.

ee Donatp. Air layering with polythene film. Arnoldia 11: 49-62. pl. 13,

1951.

ar layering with polythene film. Jour. Royal Hort. Soc. 75: 135-140.
1952.

f. 65, 66.

The Arnold Arboretum. Jour. Royal Hort. Soc. 76: 225-230. f. 107-117.
1951.

The Dove-tree-an unusual flowering tree. Plants & Gardens 7: 107.
1951

Elms grown in America. Arnoldia 11: 79-93. pl. 18-21. 1951.
Five interesting trees. Arnoldia 11: 71-74. pl. 16. 1951.

The flower colors of one hundred hardy Azaleas. Arnoldia 12: 41-44.
1952.

or your garden; ornamental trees and shrubs. House & Garden 100:
204, 206, 208. 1951.

Forty-five of the best trees for Massachusetts gardens. Arnoldia 12:
1-20. pl. 1-6. 1952.
Ground covers. Popular Gardening 3(6): 32-33; 62-63. 1952.
Layering plants in Holland. Am. Nurseryman 95(10): 7, 57.
Make bare spots beauty spots with ground covers. Popular Gardening
3(6): 32, 33, 62, 63. 1952.
Metasequoia brought up to date. Plants & Gardens 7: 265-267. 1951.
National Parks of Western North America. Trans. Worcester County
Hort. Soc. 1951: 49-53. 1951.
Plant trees and shrubs for sequence of bloom. Flower Grower 38: 36,
37, G7.

1952] THE ARNOLD ARBORETUM DURING FISCAL YEAR 409

Wyman, DonaLp. Pruning. Plants & Gardens 7: 106. 1951.
— Simple key to the pines. Arnoldia 11: 63-70. pl. 15. 1951.
Smaller street trees needed. Trees 11(6): 6, 7, 16.
Spring comes to the Arnold Arboretum. Arnoldia Pes 45-46. pl. 12. 1952.
Trees for American gardens. New York, Macmillan Co., 1951. i-vii,
1-376p. illus.

Woody plants used in colonial Williamsburg. Arnoldia 11: 75-78. pl. 17.
951.

Kari SAx
Director,

410 JOURNAL OF THE ARNOLD ARBORETUM [voL. xxxm

Staff of the Arnold Arboretum
1951-1952
Kart Sax, S.D., Professor of Botany and Director.
IvAN Murray JOHNSTON, Ph.D., Associate Professor of Botany and Asso-
ciate Director. Supervisor of the Library and Herbarium.
JosEPpH Horace FAuLL, Ph.D., Professor of Forest Pathology, Emeritus.

ELMER Drew Merrit, S.D., LL.D., Arnold Professor of Botany, Emeri-
tus.

IrnviNG WIDMER BAILEY, S.D., Professor of Plant Anatomy.
RocER Gisss CoGcESHALL, Assistant Propagator.

BEATRIX FARRAND, L.H.D., Consulting Landscape Gardener.
RICHARD HAROLD FILLMorE, M.S., Propagator (Resigned).
ALFRED JAMES ForDHAM, Assistant Superintendent.
CONSTANCE MANSFIELD GILMAN, Business Secretary.
HeMAN ARTHUR Howarp, Assistant Horticulturist.
SHIu-YING Hu, Ph.D., Assistant in the Herbarium.
CLARENCE EMMEREN KosuskI, Ph.D., Curator of the Herbarium.
LEwIs FREDERICK Lipp, Propagator.

Susan DELANO McKeE vey, A. B., Research Associate.

Liry May Perry, Ph.D., Botanist.

LAZELLA SCHWARTEN, Librarian.

FRANS VERDOORN, Ph.D., Research Associate.

RoBERT GEROW WILLIAMS, B.S., Superintendent.

DonaLtp Wyman, Ph.D., Horticulturist.

1952] INDEX

INDEX

Aberrant Silver Maples, 296

— anisodont

— insulana, en 394, 395

— — flavicans, 394, 396

— — glabrescens, 394, 398, 400
—— insulana, 395

— — pubescens, 394, 395

— denudata, 399

Actinidia, A Taxonomic Review of the
Genus, 1

Actinidia, 4

— sect. Leiocarpae, 5

—— arguta

eae
— cordifolia, 34

Actinidia isa rufa, 34
— chinensis, 61

—— iste ak 48
——pilosula, 40
—-— pubescens, 49
——r 3

a,
— — sabiaefolia, 44

| | |
| S|
28 8
Fey. &
moo
~" 3 a9
u~ Ss
Nn’
as
w

—— Ee pe 28

a, 59
Si aisuikte 17,18; 19

412 JOURNAL OF THE ARNOLD ARBORETUM _ [vot. xxximt

Actinidia kolomikta gagnepainii, 19 Aipyanthus echioides, 336
——kolmikta, 18 Allomorphia,
— kwangsiensis, 26 — caudata, 176
—lanata, 52 — exigua, 233
— lanceolata, 58 Weta, sp
aia ees. oi 50 ylla,
indochinensis Alpinia ee 214
Pe 50 —elatior, 214
——mollis, 52 —longiscapa, 216
— lecomtei, 22 — magnifica, 215
— longicauda, 60 — spe , 215
— maloides, 25 Alsodeia rapier 240
— megalocarpa, 32 Alsodeiopsis, 255, 2
—melanandra, 22, 27, 28 — Schumannii, oo
— — latifolia, 29 — Staudtii, 266
—melliana, 9 — dara 266
— miquelii, 51 — Zenkeri, 266
— pachyphylla, 57 Alyxia amoena, 115
— petelotii, — linearifolia, 1
—pilosula, 40 Amentotaxus, The Genus, 192
— platyphylla, 34 Amentotaxus,
— polygama, 20, 21, 23 — argotaenia, — 195, 196, 197
— leco ; — cathayensis,
—— polygama, 21 — formosana, Peo
— pubescens, 49 — yunnanensis, 197
— pu ea, Amomunm, biflorum, 216
— rankanensis, 16 agnificum
— remoganensis, 16 Ampelocissus korthals 250
— rubricaulis, 24, 35 racemifera, 2
— rubus, — thyrsifi er
—rudis, 8 Anchusa saperctiin, 325
— rufa, 34 — canescens,
uta, 29, 3 — echioides, 336
—— cordifolia, 34 — hirta,
— — dulcissima, 34 — hispidissima, 325
— — parvifolia, 27 — mexi ,
—typica, 29, 34 — polygama, 325
— sabiaefolia, 44 — tuberosa, 352
—_ , virginiana, 340
— subglaucifolia, 14 Anerincleistus, 174
— tetramera, — caudatus, 17
—trichogyna, 43 Anisomallon, 255, 263
—valvata, 23 — clusiaef
enosa, 41, 4 Anisophylla disticha, 226
— -— pubescens, 42 trapezoidalis, 226
— volubilis, 22 Anneslea, The Genus. Studies in the
Actinodaphne eae Theaceae, XXV, 79
Adinandra, 2 Anneslea, 81
— dumosa, on Ipi
sylvestris, 214 — crassipes, 85
Aeschynanthus, 214 —— ata, 85
— radicans, 214 —donnaiensis, 89
— volubilis, 214 — fragrans, 81
Aglaia odorata, 214 ——al
Aglaonema nitidum, 218 —— cra

ssipes, 85
— oblongifolium, 218 —hainanensis, 86

1952] INDEX 413

Anneslea fragrans lanceolata, 85 Arnebia Griffithii, 326
monticola, 85 — guttata,
—— 2 ferneeroemioides 87 — Hancockiana, 354
——typica, 82 — hispidissima, 325
—lanceolata, 86 — inconspicua, 333
monticola, 85 Meee ae 326
— steenisii, — linearifolia,
— ternstroemioides, 87 — longiflora, ae
Annesleya, — lutea,
nonymos caroliniense, 339 — macrothyrsa, 332
Anplectrum, 5.239 — minima,
— divaricatum, 233 — obovata, 329
—glaucum, 233 _
—rostratum, 235 — orientalis, 322
vim , 234 —— pe
ntidesma, 216, 367 — purpurascens, 325
—elassophyllum, 371 — Sewerzowi, 32
— frutescens, 216 — speciosa, 333
—ghaesembilla, 216 — Szechenyii, 330
—gillespieanum, 370 — tetrastigma, 321
— insulare, —Thomsoni, 331
— pacificum, 368 — tibetana, 331
— pubescens, 2 — tinctoria, a
—sphaerocarpum, 368 —ti
—trichophyllum, 373 Arnebiola died 325
Aplectrum, 234, 235 Arnold Arboretum during the Fiscal
—rostratum, 235 Year Ended June 30, 1952, The, 403
—stipulare, 235 Astronidium degeneri, 10
—viminale, 234, 235 Axanthes arborea, 24
Apodytes, 255, 262 — tom ig 249
be , 263 Ayer a
—pbrachystylis, 263 Azadirachta rasta 235
cambodiana, 263 Baccaurea, ,
cuminensis, 263 — malayana, 227
— dimidiata, 262 Backeria, 235
— Gardneriana, 263 — viminalis, 2
nnanensis, 262 Barringtonia macrostachya, 218
Ardisia divergens, 216 Barthea cavaleri
nd » 217 Batschia canescens, 340
p , 216, 21 caroliniensis, 33
Areca tigillaria, 217 —conspicua, 3
Aristolochia hastata, 217 — decumbens, 344
5 — disticha, 361
Arnebia, 312 — Gmelini, 339
— asperrima, 325 — longiflora, 344
— Bungei, 326 — pilosa, 348
— cephalotes, 332 — sericea, oe
— cornuta, 322 — Torreyi
— decumbens, 322 Bauhinia its: 217
— acrocalyx, 324 — emargi
— densiflora, 331 — lucida, 217
— echioides, 336 Begonia, 21
— euchroma, 334 — bracteata, 217
— fimbr 2 — caespitosa, 217
— fimbriopetala, 326 — fasciculata, 217

— flavescens, 328 — geniculata, 217

414 JOURNAL OF THE ARNOLD ARBORETUM

Begonia isoptera, 217
7

227
Bibliography the Published Writings
the Staff and Students July 1,
951 — June 30, 1952, 407
Blastus fengii, 166
Boraginaceae, Studies in the, XXII.
Noteworthy Species, Chiefly Asian
ican, 6

aa Studies in the, XXIII. A
Sur the Genus Lithospermum,
ae

Bourreria viridis, 64
rol,
Bruguiera cylindrica, 242
Bua choopa, 239
Buraeavia carunculata, 375

— fragrans 82°

Campnosperma, 2 21

Cantleya, 256,
— corniculata
Capellia peered 250

oe bivalvis, 218

— pau , 240
Celtis attenuata, 218
Centipeda minima, 117
Cephalotaxus eed 195, 196

ta, 237
Chamaecladon angustifolium, 218
China, Notes on the Flora of, II, 166
Chionanthus litoreus, 231
Chionotria, 218

[ VOL, XXXIII

Chionotria — 218
Chirita ba a
— horsfieldi
Salary 256, 276
— —
Choopa, 240
Cinnamomum parthenoxylon, 230
Citronella, 255, 259
— costaricensis, 259

259
Cle idion pane 390
— leptostachyum, 390
— vieillardi vi itiensis, 390

20
— pendaliorum, 219, 220
— serratum,
— arene 219
— villosum, 219
— wallichii, 220
Cleyera seco vgs 248
— rubigino
Cnestis i 220, 221
rida, 220

—coriaceum, 221
ne Morphology of the Icaci-
e, VI. The Pollen, 252
Comarus lin 221
72)

— semidecandrus, 221

1952] INDEX 415

Connarus villosus, 221
Cordia Bridgesii, 64

— marchionica, 65

— varronifolia, 62

— viridis, 64

Craniospermum mongolicum, 74

—su eee
Cremostachys fulva, 231
saison ‘aiiiventia: 99
nculata, 9
Gieolentha igh i
Cunoniaceae of and Samoa, The.
‘i ae of ae ‘Island Plants, i;

Pen jackiana, 236
Curculigo canttuliferd: 222
ral

Zz
Cunosiosat um amabile, 1
Cyphorima latifolia, ns
— lutea, 342
Cyphotheca hispida, 167
Cyrtandra aurea, 222
— bicolor, 222
— carnosa, 222
— frutescens, 222

ipes, 8
Decaspermum fruticosum, 226
Dehaasia i SS ri 230

— microcarpa, 2

eee 255, 260

— bolivia

SU ee festa 231
caine

— Preus

— Voeelii, ae 272

Didissandra elongata, 223
— frutescens, 223
Didymocarpus barbata, 222
—_ corniculata, 222, 223

=
pk
°
QO
=
=
=
8

Draparnaudia multiflora, 152

Drimyspermum phaleria, 239

Drymispermum, 2

an lecrmn aromatic, 223
— camphora, 22

’
Echiochilon hispidissima, 325
Echioides asperrimum, 325

— linearifolium, 328
obov :

— tinctorium, =
Echium Benthami, 333
Elaeocarpus eeu. 237

Embelia canescens, 224
Emmotum, 255, 256

416 JOURNAL OF THE ARNOLD ARBORETUM

Emmotum affine, 257
— argenteum, 256
— fagifolium, 257

_- orbiculatum, 257

Enchidium, 22

— conoid , 224

Endosp ermum mavrophy Tian. 390

— malayana, 224
Eriandra, a New Genus of Polygalaceae
from New Guinea, 91

ans, 94
Erioglossum rubiginosum, 244
Eritrichium elongatum Paysoni, 67

Ervatamia macrocarpa, 246

— sphaerocarpa, 24

Euphorbiaceae, Notes on Fijian. Studies
of Pacific Island Plants, XIJI, 367

Eurya greenwoodii, 97

e Cunoniaceae of.
oo of enna poe Plants, XII,

oe Euphorbiaceae, Notes on. Studies
of Pacific Island Plants, XIII, 36

ai _ Plants, Further Notes

s of Pacific Island Plants,

XL 97

Flacourtia inermis, 225

Fordiophyton onpetiltum, 168

Fremya integrifolia, 1

Fremya ele 153, 160

ee |
ardenia snlaphella, 226, 240
ae ae 256, 2
— austro- caledonica, 267

— malayana, 219

— arenas macrorachis, 218
— rigida,

Gmelina ie 226

— villosa,

Gomphandra mollis, 269

269
pict chlo 226
— sumatra
Prensa 255, 265

— longeracem 265
fa eae sion, 117
wanes corymbos
Grewia paniculata, 236
a,

227
Hedychium Sade 227
—sumatranum, 22

[ VOL. XXXIII

1952] INDEX 417

Helicia attenuata, 243

oe latifolia, 50
mena humilis, 218
mila, 2

Hornstedtia, 215

sce 256

— sinense, 271

Hoya rh 227
grandiflora, 227

ee sere) 227
Hu, Suru-yinc. Notes on the Flora
of China, II, 166
Humincncien. 255, 261
61

— senegalensis, 261

Icacinaceae, The Comparative Morphol-
ogy of the, VI. The Pollen, 252

Tlex, 238

_ philippinensisy 274
— aaa tella

iy
ee 256, 266
2

Ixora candida, 228
— montana, 228
— neriifolia, 228

Ixora opaca, 228

— parkinsoniana, 228

— pendula,

— — opaca, 228

Jack’s Genera and Species of Malaysian
Plants, William, 199

Johnia sumatrana, 228, 244

Jounson, ALBERT G.
White Pine eager 179

Spontaneous

, Ivan M. Studies in the
Boraginaceae, XXIII. A Survey of
the Genus Lithospermum, 299
Jonesia, c
— declinata, 2
Kadsura pubescens, 50
Kai namo,
Kajos
Kalambuthi, nee

Kjellbergiodendron Burret, Notes on
Xanthostemon F. Mueller and, 15¢
ap ae

Knesbeckia, 217

Ko nang na, 87

KosuskI, CLARENCE E. Studies in the
Theaceae, XXV. The Genus Anneslea,
7

OBUSKI, CLARENCE E. Studies in the
ea XXVI. The Genus Visnea,

ca ee sary rae 18
Kolonimbeka, 1

Kutakuta, 142, of
Kutumiras Bele

Lacaitaea a. a, 75
Lagerstroemia floribunda, 228
Langsat

Lanséh, 227; 229

418 JOURNAL OF THE ARNOLD ARBORETUM

Lansium, 229

— aqueum, 229

—— domeaticuts, 227, 229
Lansone, 229
Lasianthera, 256, 270

Lepidopetalum jackianum, 236
Leptanthe macrostachya, 333

Leptospermum, 22
— commune javanica, 226
— javanicum, 226

5
Li, Hur-Lin. The Genus Amentotaxus,
192

Li, Hur-Lin. A Taxonomic Review of

Li hile ech 241
— spicata, 2
— urceolaris, 7
Lithodora da 354

[ VOL. XXXII

Lithospermum, A Survey of the Genus.
Studies in the Boraginaceae, XXIII,

299
Lithospermum, 312
— aequatoriale, 359
— affine, 359
—- nears 348
— meee
—andin
a stim, 343, 352

— Griffithii, 326

1952]

Lithospermum guatemalense, 354
um, 330

— latifolium, 342

— laxum, 348

— linearifolium, 328, 343

—longiflorum, 336, 344
ile, 342

— obtusifolium, 349

— officinale, 342

—-— erythrorhizon, 341
— — erythrorrhizon, 341

— — lanceolatum, 348
—-— macrospermum, 348

INDEX

Lithospermum ruderale Torreyi, 348

— Szechenyi, 3

— poe ae 331
— tinctori

— Tor

Longetia, 376

Lucinaea, 2

ba foliosa,
ga

— — puberula, 388

420 JOURNAL OF THE ARNOLD ARBORETUM

Macaranga leptostachya, 390
— macrophylla, 390

— magna, 381

_- marikoensis, 385

— vitiensis

Macrolenes,

ata, 330
Macrosolen cochinchinensi, 231

— retusus, 2
Macrotomia salbaaek 333
2

334
Maesa tenophsla eh
Makamakandora

Malaysian Plants, rion Jack’s Genera

and Species of, 19

Maple, Wier, 296

— macrophylla, 390
— seemanni, 379
Mappia, 256, 271
— angustifolia, 271

Mappia longipes, 272
—— mexicana, 272

— racemosa brachycarpa, 272
Mappianthus, 256, 2

3
— stellulata, 234
Mbuambua, 113
Medinilla 232
— crassifolia, 2
— erythrophyl 234
— eximia,
— hesseltii,
— javanensis, 232
— nana,
— petelotii, 169

— glabra,

— laxiflora, 269

— papuana, 269

a oe -

— vitiensis,

Melastoma mn 232
— bracteatu

— gracile, 233
oe 234

— impuber

— malabtivicum, 233, 234

— viminale, 234

6
— cnet 236

[VOL. XXXIII

1952] INDEX 421

Melodinus eugenifolius, 230
Memecylon coeruleum, 235

s on Xanthostemon
: Mueller. Oe "Kiellbergiodendron

Burret, 150

Merritt, E. D. William Jack’s Genera
and Species of Malaysian Plants, 199

Mersiliodendeon, 255, 265

— rotense,

ni ay 156

y
manele: 244

man Se 236

— paniculata, 236

— tomentosa, 236

Microtropis bivalvis, 2

Microula blepharolepis, 72

parncy tora 236
umatrana, 236

Milnea ci, 236

Mimosa djiringa, 236

the Icacinaceae, The
Comparative, VI. The Pollen, 252
Mountnorrisia, 81
ee

— conglobata, 332

B 257
— ea dne 237

Myosotis ici 361

— grandiflora, 361

Myristica ae 228
sum 8

co

Nagaba- shirakuchizulu, 60
Nani, 1

Nania, 156

— petiolata, 156

— vera, 156

Natha, 372
Natron 256, 271

— herpeticum, 271

Nda’al

Ndavo, 379, ae 383, 384, 386
Ndavolutu,

Necdigaaees 235

— gracilis, 2

—ampullaria, 237
— distillatoria, 238
— gracilis, 2

Nephelium lappaceum, 238
Neuburghia sumatrana, 246
38

238
ew Genus of Polygalaceae from New
Guinea, Eriandra, A, 91
New Guinea, Eriandra, A aad Genus
of Polygalaceae from,
Nicolaia, 215

— speciosa, 215

Notes on the Flora of China, II, 166

Notes on Xanthostemon F. Mueller and
Kjellbergiodendron Burret, 150

Noteworthy Species, Chiefly Asian and
South American. —— in the Bor-
aginaceae, XXII, 6

ee 256, 7

— foetida,

— pitosproies 2530215

er

238
Octosom ne Kerrii, 75
aap ee 259

mex
Olea ae He

422 JOURNAL OF THE ARNOLD ARBORETUM

~— blepharolepis, 72
usa,

feat aise fallax, 233

Oncosperma filamentosum, 217

— tigillaria, 217

O m carolinianum, 339

Ophiorrhiza heterophylla, 238

miirineertay 255, 257
— cubensis, 257
Ouratea aston 226

tu,
ton ubekic 170
Pachycentria fengii, 170
Pacific Island Plants, Studies of, XI.
Further Notes on Fijian Flowering
sh nts,
on island Plants, Studies of, XII.
e Cunoniaceae of Fiji and Samoa,

te
Pacific Island Plants, Studies of, XIII.
Notes on Fijian Euphorbiaceae, 367
Pagiantha macrocarpa, 246

— macrocarpa, 2

Aberrant ee nee
Pennantia, 255, 2
— corymbosa, 8

Pentaloba ee

ines aa 239
Peronema, 239

— canescens, 239

=i os 230

Pete

S. and — am:

[VoL. XXXII

Petek, 214
Petermannia, 217, 218

—ca a, 239
Phylagathis rotundifolia, 234
— wenshanensis, 171

Phyteuma aad 239
Phytocrene, 256, 2

|

— vane 183
— s < parviflora, 183
Pithecllobim bubalinum, 227
— clypearia, 227
— jirin 36
ne serrulata, 240
Pittosporopsis, 255, 258
— Kerrii, 258
Pittosporum serrulatum, 240
a eer esquirolii, 172

— hen
Plates, 2 Aba
arr 261

61
philippinensis, 260
Plerandra ym 103
picker , 103
Pleurisanthes 255, 262
yee
ieee ae te 262
Podocarps argotaenia, 194, 195, 196
— insignis, 195
Pare ee pulverulenta, 234
Pollen, The. The Comparative Mor-
phology of the Icacinaceae, VI, 252
i cegerny 256,
Pogge

she from hg — Erian-
dra, ew Genus of,

Polvgalaceae tribe “nie 93

Polyporandra, 256, 275

— scandens, 275

Poraqueiba, 255, 257

— sericea,

Poroporo, 371

1952]

Posoqueria anisophylla, 240
Prunus kolomikta, 18
Pseudixora sumatrana, 247
Psilobium, 240

— lamponga, 2

1
-- pceolnne: 241, 242
Quisqualis Sentara: 245

87
Rhaphiost tis . 264
— cordifolia
_— ferrugines, ae
264

Rhododendron apoanum, 243

INDEX 423

Rhododendron malayanum, 243
Rhopala, 243

— attenuata, 243

— ee 243

vata, 243
Rhytiaryum, 256, 276
— elegan
Richtera,
pict anisodontus, 397

Rinorea lanceo slat ata, 240
Rondeletia corymbosa, 243
— spicata, 243

e

Roscoea, 245
— pentandra, 245
Rote

Rourea javanica

Royen, P. van & C. G. G. J. vAN STEENIS.
Eriandra, a New Genus of Polygala-
ceae from New Guinea, 91

Rure, 145

ru, 393

Sagus laevis, 244

Saitamu, 14

Salacia, 227, 228, 244

— prinoides, 228, 244

Salvia splendens, 116

Samoa, The Cunoniaceae of Fiji and.

eer of Pacific Island Plants, XII,

19

u ni mbati, 98
Scatalcuiee Soiduni, 220
— mimosoides, 220, 221

Sapindus rubiginosus, 244

Saraca declinata, 228

Sarcopyramis bodinieri, 172
173

— bo
Saeee 256. 276
— Horsfieldii, 276
~ philippinensis, 276

424 JOURNAL OF THE ARNOLD ARBORETUM

Sarcostigma surigaoénse, 277
— Wallichii, 277

Sasariwai, 394
Saukalambuthi, 371

a rubiginosa, 248

— serrata, 248

— tomentosa, 49

— tristyla, 247, 248

Sax, Kart. The Arnold Arboretum dur-
ing the Fiscal Year Ended June 30,
1952, 403

Schwenkfeldia glabra, 249

— malaccensis, 249

Sicenthers dubia, 246

Scurrula ferrugines, 231

Scyphiphora, 2

- eee 224

Seruserumasala, 105

Sila, 109
Silver Maples, Aberrant, 296
annie
miTH, A. C. Studies of Pacific Island

Plant ts, — chee Notes on Fijian
Flowering Plants,

SmiTH, A. C. i. of Pacific Island
Plants, XII. The Cunoniaceae of Fiji
and Samoa, 119
irH, A. C. Studies of Pacific Island
Plants, XIII. Notes on Fijian Euphor-

ie 245
—confertus, 245
— diversifolius, 245
Sphenodesme, 245

[ VOL. XXXIII

Sphenodesme pentandra, 245
Spiraeanthemum,

e,
Spontaneous ia Pine Hybrids, 179
Staff of the Arnold Arboretum 1951-

1952,

Stagmaria, 245

— verniciflua, 245

Stentinabyben erectum, 174

STEENIS, C.G.G.J. VAN & P. vAN ROYEN.
Eriandra, a New Genus of Polygala-
ceae from New Guinea, 91

Stemonurus, 256, 267

~- mollis, 269

— Petelotii, 26
— polymorphus, 268
— Yatesli, 268

Stenosolenium perenne, 334
Sterculia angustifolia, 245
— bracteata, 246

245
Strobila pene ae ima, 325
Strophanthus Nenrnl 246

Studies in the Boraginaceae, XXII. Not
worthy Species, ey Asian me
South American,

Studies in the Saas XXII.
ie ey of the Genus Lithospermum,

Sr of Pacific Island Plants, XI.
Further Notes on Fijian Flowering
Plants, 97

1952]

sea of Pacific Island Plants, XII.
e Cunoniaceae of Fiji and Samoa,

Seine of Pacific Island Plants, XIII.
Notes on Fijian Euphorbiaceae, 367
Studies in the ae ae, XXV. The

19
Studies in the a cocee. XXVI. The

Genus Visnea, 1
Styphelia malaica, 230

— sphaerocarpa, 246, 247
24

=
oO
n
~

— leptostachyus, 390

Tandalo, 114, 145
ee aT i aaa 106
— ligulifolium

7

Taxonomic ie of the Genus Acti-
nidia, A, 1

Teinivia, 112

Telogyne indica, 224

anne luteola, 100

— vitiensis, 100
Ter cision acuminata, 247
— cuspidata,
— pentapetala, 247
earner 248
— serrata, 24
Tetracera oe 248
—eury ,
— luci

8
Tetranthera cordata, 248
Thaunilawa, 113

INDEX

Theaceae, Studies in the,
enus Anneslea, 79

Theaceae, Studies in the, XX VI.
8

7
eae 227
Tomi tom
eed ee 325
— purpurascens, 32
Trichodesma calcareum, 75

— radicans, 2 14

— volubile, 2

Trigonostemon, 224

— indicus,

— verticillatus, 2

Trigonotis ciliolata, 68
— cupulifera,

— ee 70
Beast

Tegohens , 224

Triosteum hecitans 229

Tristania celebica, 164

— odorata, 161

pachysperma, 161, 162

aeciiicma arguta, 32
— kolomikta, 18

— ey 21

— 34

— volubilis, 21
0

Uvaria hirsuta, 249

XXV.

The.

426 JOURNAL OF THE ARNOLD ARBORETUM

Uvaria pilosa, 249

Vaccinium ellipticum, 249

— laurifolium, 249

— sumatranum, 249

Vareca lanceolata, 240

Varronia Bridgesi, 64

enua,
es 249
alayana, 249

Uncen malayanum

Visnea, The Genus. ane in the Thea-
88

racemifera, 250

Lie 123, 125, 1

1
— rhodogyne, 133

[ VOL. XXXII

Weinmannia richii, 133, 138

5
White Pine Hybrids, sige 179
and Species of

giodendron Burret, Notes on, 150
4

— myrtifolium, 153, ea 161
— novaguineense, 1
— pachyspermum, 161, 162
— papuanum,
— paradoxum, 154, 155, 157
We ince. 152, — 161

— purpu
tara 151, on 159, 161
— verdugonianum, 16
— whitei, 154
Zingiber gracile, 250

n~
an