JOURNAL S740
OF THE S

ARNOLD ARBORET UM
HARVARD UNIVERSITY

B. G. SCHUBERT
EDITOR

T. G. HARTLEY C. E. WOOD, JR.

D. A. POWELL
CIRCULATION

VOLUME 49

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PUBLISHED BY
THE ARNOLD ARBORETUM OF HARVARD UNIVERSITY
CAMBRIDGE, MASSACHUSETTS
1968

MissouR! Boramicat
GARDEN LIBRARY

DATES OF ISSUE

No. 1 (pp. 1-166) issued January 15, 1968.
No. 2 (pp. 167-290) issued April 22, 1968.
No. 3 (pp. 291-380) issued July 16, 1968.
No. 4 (pp. 381-559) issued October 17, 1968.

TABLE OF CONTENTS

IrvinGc Wiper Barry, 1884-1967. With portrait. Richard A.
Howard
INFLORESCENCE IN NANNORRHOPS RITCHIANA (Patmar). P. B.
Tomlinson and H. E. Moore, Jr.
Harperocatuis, A New GENUS OF THE LILIACEAE FROM FLORIDA.
Sidney M Dita
CYTOTAXONOMY OF ILLICIUM FLORIDANUM AND I. PARVIFLORUM
(InuictAcEaE). Donald E. Stone and Judith L. Freeman .......
FurtHeR MorpHouocicaAL Srupres ON ANASTOMOSES IN THE
Dicuotomous VENATION oF CrrcarasterR. Adriance S. Foster
A SyNopsis OF THE ASIAN SPEcIES OF DELPHINIUM, SENSU
Srricto (Continued). Philip A. Munz
FLowers, Fruirs, AND PHYLOGENY OF ALANGIACEAE. Richard H.
yde
Diverse Nopau Types tn Ruopopenpron. W. R. Philipson and
M Philipson
A New Hyprancrea rrom Mexico. Lorin I. Nevling, Jr., and
Arturo Gémez-Pompa :
A Synopsis OF THE ASIAN SPECIES OF DELPHINIUM, SENSU
Srricto (Concluded). Philip A. Munz
MorpPHoLoGy AND ANATOMY OF CROOMIA PAUCIFLORA (STEMONA-
cEAE). P. B. Tomlinson and E. S. Ayensu
HERISSANTIA, BoGENHARDIA, AND Gayorpes (MALVACEAE). George
Brizicky
JAMAICAN AND OTHER SPECIES OF BUMELIA (SAPOTACEAE). William
T. Stearn
ANATOMY OF THE PaLM RHAPIS EXCELSA, V. INFLORESCENCE.
P. B. Tomlinson and M. H. Zimmermann
ANATOMY OF THE PALM RHAPIS EXCELSA, VI. Roor anp BRANCH
Insertion. P. B. Tomlinson and M. H. Zimmermanzn ............
CompaRATIVE MorpHo.ocicaL Stupres 1N Di.ientacear, III.
Tue Carpets. William C. Dickison
REVISION OF THE GENUS NoOTELAEA (OLEACEAE). P. S. Green. ........
CoMPARATIVE ANATOMY OF THE LEAF-BEARING CacTaceAk, XVII.
PRELIMINARY OBSERVATIONS ON THE PROBLEM OF TRANSI-
TIONS FROM Broap TO TERETE Leaves. J. W. Bailey ................
BoranicaAL NAME OF THE TEA SAGERETIA OF CHINA. Marshall
C. Johnston
THE Eco.iocy or AN ELFIN Forest iN Pugrto Rico, 1. [nrropuc-
TION AND Composition Stupies. Richard A. Howard ............

Ture Ecotocy or AN Erin Forest 1N Purrto Rico, 2. THE
MicrocLuiMATE oF Pico pEL Orste. Harold W. Baynton. ........
Srupres or Paciric Istanp PLants, XIX. THe ARALIACEAE OF
tHE New Hepripves, Fis1, Samoa, AND Tonoa. Albert C.
Smith and Benjamin C. Stone
SoRBUS AND THE PROBLEM OF GENERIC TyYPIFICATION. George K.
Brizicky
EmpryoLocy oF Drimys winterI. N. N. Bhandari and Revathi
Venkataraman
Tue Drrector’s Report
InpEx TO VOLUME 49

fie 6

VOLUME 49 NuMBER 1

JOURNAL

OF THE

ARNOLD ARBORETUM

HARVARD UNIVERSITY

B. G. SCHUBERT
EDITOR

T. G. HARTLEY Cc. E. WOOD, JR.

D. A. POWELL
CIRCULATION

THE JOURNAL OF THE
ARNOLD ARBORETUM

Published quarterly by the Arnold Arboretum of Harvard University.

Subscription price $10.00 per year.

Volumes I-XX, reprinted, and back issues of volumes XXI-XLV are
Soctahaes as the Kraus Reprint Corporation, 16 East 46TH Street,
New York, N.Y. 10017.

Subscriptions and Sapien ay should be addressed to Miss Duucte A.

PowseLL, ARNOLD ARBORETUM, 22 Divinity AVENUE, CAMBRIDGE, Massa-
CHUSETTS 02138.

CONTENTS OF NO. 1

irene Wier Baney, 1884-1967. With evotae Richard A.
Howard 1

IyrLonescence: IN Nina. RITCHIANA — (Pavan). P. B.
To: cs H. E. Ss dt:

HER " Mosenovocrcas, Srupres oN Seca IN THE
US VENATION OF CIRC. a. Adriance S. Foster 52

DELPHINIUM, SEeNsu Stricro

J Pa Munz -

IRVING WIDMER BAILEY

JOURNAL

OF THE

ARNOLD ARBORETUM

VoL. 49 JANUARY 1968 NUMBER 1

IRVING WIDMER BAILEY, 1884-1967

RicHArp A. Howarp

With portrait *

THE SUDDEN DEATH OF Irving Widmer Bailey on May 16th ended the
long and productive career of a brilliant scholar and a son of Harvard in
every sense of the word. His place cannot be filled; his kind of contribu-
tion cannot be repeated; his leadership and inspiration are not to be
duplicated. He was ‘Professor Bailey” to all but a very few, and the
appellation was offered in respect and admiration.

Irving Widmer Bailey was born in Tilton, New Hampshire, August 15,
1884, the only child of Solon Irving Bailey and Ruth Elaine (Poulter)
Bailey. His father, an assistant professor of astronomy at Harvard, was
sent to Peru in 1889 to select a site for a high altitude observatory. He
chose the summit of El Misti at 19,000 feet and, with his family in
residence, supervised the operation of an observatory constructed ‘there.
Irving Bailey wrote later, “My reactions and activities in college and
subsequent to graduation were profoundly influenced by the fact that eight
of the first thirteen years of my boyhood were spent in remote parts of
South America rather than in the environment of New England. Having
no formal and stereotyped education until the age of thirteen, having no
playmates of my own race and age, I was forced to rely upon my own
resources for interests and activities. Much of my time was spent in
hunting, in exploring the Andes at high altitudes, in learning at first hand
the traditions and beliefs of Peruvian Indians, and in observing the
spectacular activities of Catholic Spanish in peace and in revolution. I
developed at an unusually early age, in association with my father and
other astronomers, a keenly analytical interest in natural phenomena and
in the activities and foibles of the human race under differing hereditary
and environmental influences.” :

Irving Bailey entered Harvard College in 1903 and was graduated with
the class of 1907. In his 50th anniversary report he related, “In college
I browsed around in history, chemistry, geology and meteorology, but it
was not until my senior year that speeches of Gifford Pinchot and Presi-
dent Eliot induced me to undertake a career in forestry, particularly owing

* Photograph by John Brook, Boston, Massachusetts, 1955.

2 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

to the appeal of an out-of-door profession.” Bailey entered the graduate
school at Harvard in 1907, and his subsequent career was to parallel many
changes in the administrative organization of botanical science at Harvard,
in which he was to play a continuing and major role. The year 1907
marked the establishment of the School of Applied Science at Harvard
with both Forestry and the Bussey Institution, the latter newly reorganized
as a school for study of problems relating to agriculture and horticulture,
transferred administratively to that School. The Harvard Forest was also
acquired in 1907, and the following year a division of Forestry was
initiated. In 1908 Bailey received his first Harvard appointment as
Assistant in Botany. He was awarded the Master of Forestry degree in
1909 and almost simultaneously an appointment as Instructor of Forestry
in the Graduate School of Applied Science. In 1912 a short-lived School
of Forestry was formed, with Bailey on the staff as an Assistant Professor.
In 1914 the Bussey Institution and the School of Forestry were combined
as the Graduate School of Applied Biology, which was then known inter-
nationally and now historically as the Bussey Institution. Professor Bailey
was promoted to Associate Professor of this Institution in 1920 and became
Professor of Plant Anatomy in 1927. In 1931 further administrative
changes abolished the Bussey Institution as a school and transferred its
instruction and degree-granting privileges to the Division of Biology. At
that time Professor Bailey was transferred to the staff of the Arnold
Arboretum for budgetary purposes, although his office was to be in the
Biological Laboratories in Cambridge until the year before his retirement.
Then, with the completion of the Harvard University Herbaria building in
1954, Bailey moved his office and laboratory along with the wood collection
into the new building. After his retirement in 1955, he continued to use
work space in the quarters of the wood collection and followed a regular
schedule in the laboratory until the heart attack that ended his life. This
building represents the fulfillment of his own plan, presented to the Har-
vard Corporation at their request in 1945, for assembling most of the
botanical institutions at Harvard under one roof.

Although he carried the title of Professor, Irving Bailey was not
primarily a teacher. His students during the years at the Bussey Institu-
tion and later in the Department of Biology were those of graduate level
and advanced instruction. Few students actually received their advanced
degrees under his supervision. Because he was not directly responsible for
student guidance, he was even more effective in giving help when it was
requested and needed. The few courses he taught were organized through-
out in the same meticulous fashion with which he wrote. His superb
photographic techniques produced lantern slides that focused on the exact
point to be made. His lectures were a pleasure to listen to and a student’s
delight to annotate, yet he worried incessantly before and after each
scheduled presentation. His talent for teaching, however, was also evident
in his association with younger people who served as his technicians. The
junior authors of many, many papers, Mary Vestal, Anna Faull, Richard
Howard, Charlotte Nast, B. G. L. Swamy, were assistants who shared the

1968 | HOWARD, IRVING WIDMER BAILEY 3

honor of being included in some of Bailey’s writing and “graduated” to
contributions of their own.

It is his research and his published contributions that are the expression
of the true talents of Irving Bailey. His first paper was published in 1909,
and one remained in manuscript at the time of his death. In the 58 years
of his productive career Bailey published the 142 papers listed in the
bibliography which follows. These cover a variety of fields and show a
return, often years later, to an original subject for further elaboration or,
when needed, a change in direction. Professor Bailey is credited with two
books. In reality one is a small printed report on the status of forestry
education and the other a collection of his papers reprinted in book form.
He was often visited by representatives of book publishing firms pleading
for him to write a book. He was most generous in giving his time to
reading manuscripts of other writers and offering his criticisms or com-
pliments. His own failure to write a book he once explained as his
willingness to let others summarize for, while he could, he preferred to
contribute his original ideas and investigations.

Bailey recalled in the Fiftieth Anniversary Report of the Harvard Class
of 1907, “Upon obtaining my degree of Master of Forestry, I spent several
years studying lumbering and wood-using industries, and became con-
vinced that the existing economic, social and political status in North
America was such as to inhibit for many years an extensive application
of intensive European silvicultural methods. At the same time, ecame
increasingly impressed by the lack of accurate and reliable information
regarding the anatomical structure, physical properties and chemical
composition of wood and the vital processes by which it is formed in
plants. Thus, since joining the staff of the reorganized Bussey Institution
in 1914, I have devoted my attention largely to research in various aspects
of this pioneering field. As data accumulated, they became increasingly
significant in the discussion of theoretical botanical problems of plant
identification, plant classification, plant physiology and evolution, as well
as in the solution of practical problems of wood utilization. Therefore, at
present I am generally regarded both in this country and abroad as a
botanist rather than as a forester.” ;

In reviewing his own publications for reprinting in book form in 1954,
Bailey agreed to eight chapters grouping his contributions into the broad
categories of cytology and ontogeny, biochemistry and biophysics, phylog-
eny, taxonomy, entomology, paleobotany, wood technology, and coopera-
tion in scientific research. These areas truly reflect the scope of Professor
Bailey’s abilities, and each is used often in his writings or returned to in
the course of his research. ie

Every scholar has in his background an individual influential in the
development of his career. Regrettably, there is little acknowledgement
of the role of Edward Charles Jeffrey in the development of the career of
Irving Bailey. Jeffrey was Assistant Professor of Vegetable Histology and
General Morphology at Harvard from 1902 to 1907, when Bailey was
an undergraduate. He became Professor of Plant Anatomy in 1907 and

: JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

retired in 1933. In many ways Irving Bailey can be regarded as his
successor. Bailey’s early publications, often recorded as “Contributions
from the Phanerogamic Laboratories of Harvard University,” acknowl-
edged the valuable assistance of Jeffrey or the courtesies of his laboratory.
Their subject matter certainly followed the same interests of Jeffrey and
in the beginning followed his theories. In 1914 in the early numbers of
a series of six papers titled “Investigations on the Phylogeny of the
Angiosperms,” Bailey and his co-author Edmund Sinnott dissented from
Jeffrey’s theories, the split largely starting with the subject of the
“ageregate ray.” Jeffrey’s book, “The Anatomy of Woody Plants,” pub-
lished in 1917, contained many ideas and descriptions which Bailey did
not accept, and a conflict between them continued in print. Following the
appearance of two papers by Jeffrey and R. E. Torrey in the Botanical
Gazette in 1921, Bailey and Sinnott wrote, “a vigourous attempt is made
to discredit our work. The unfortunate tone of this attack we can afford
to ignore but in view of the misrepresentation of our position which they
have introduced into the discussion we feel that it is wise to restate our
conclusions in the light of all the facts which have been brought forward,
and to endeavor to clarify the real point at issue.” The wounds created
were never healed, yet the training Jeffrey had offered the young Irving
Bailey came to an imaginative mind and an energetic body.

During World War I Bailey’s talents as a wood technologist were used
at Wright Field in Dayton, Ohio. His early interest in the preservation
of wood he also associated with studies of the structure of the stem. At
Wright Field he was Chief of the Wood Section of the Materials Engi-
neering Department of the Bureau of Aircraft Production. The principal
woods used in wing and frame construction of World War I aircraft were
spruces and firs. The methods of cutting, curing, milling and splicing
these occupied Bailey’s interests and were his responsibilities. When the
supply of spruce was threatened by insect attacks in the postwar years
he took part in studies which led to nearly classical papers on the spruce
budworm biocoenose. Bailey tendered his resignation as “Aeronautical
Mechanical Engineer” effective December 23, 1918, and returned to the
Bussey Institution in Jamaica Plain.

The two distinctive types of research which were to occupy Bailey’s
time on his return to academic life were to develop quickly. His war time
studies of wood structure developed into a long series of papers on the
cambium and its derivative tissues. At the Bussey he undertook a special
study of the stem structure of some African myrmecophytes for the
director, William M. Wheeler. This involved a study of the feeding habits
of plant-inhabiting ants, and in 1920, Bailey spent the summer in British
Guiana working at the tropical research station of William Beebe. He
had received a grant of $500 from the AAAS for this field work and during
a portion of the time he was in the company of Wheeler. The observations
they made on species of Cordia, Cecropia, Triplaris, and other ant plants
were not published until 1942, after Wheeler’s death in 1937. The work
in entomology with Wheeler and the expedition to British Guiana were

1968 } HOWARD, IRVING WIDMER BAILEY 5

apparently among the most enjoyable experiences of Bailey’s life, for tales
of these were often repeated in social gatherings.

In the course of the work in British Guiana, Bailey found a new species
of Cecropia which he described in 1922. He was also able to make
observations on other plants including species of Marcgravia. The flowers
of the unusual inflorescences of Marcgravia plants were often considered
to be pollinated by birds. Bailey’s observations on these plants and their
pollination formed a series of papers which also included two new species
of Marcgravia. Bailey did not collect many plant specimens and his
collections for that trip numbered 195 in the figures he cited.

Bailey’s ventures into taxonomy were not many. In addition to the
three species described from South America he is credited as the senior
author of the family Degeneriaceae. This resulted from the cooperative
study with A. C. Smith on plants of the Fiji Islands and was a phase of
the study of the primitive Angiosperms which Bailey insisted on calling
inaccurately the “woody Ranales.”’

Bailey is well remembered for his cooperation with taxonomists, and his
work on the Icacinaceae is representative. Perhaps from his survey work
with Sinnott on the nodal structure of plant families he had retained an
interest in families of flowering plants which showed diverse habits or

director of the Gray Herbarium. Professor Bailey assigned to me the
task of preparing the anatomical slides needed for an investigation of the
remainder of the genera. These materials were then studied and dis-
crepancies were soon obvious in the taxonomic classification of the
specimens. In some cases the specimens were incorrectly named and in
others the species were incorrectly placed. The value of anatomical
characters in taxonomy soon became evident in this family. A series of
papers by Bailey and Howard, by Howard, and by Dahl resulted from
this work. One taxon of the Icacinaceae with distinctive anatomical
characteristics could also be shown to be distinct on the basis of characters
commonly used. The genus then distinguished was named in his honor,
Irvingbaileya. In later years Swamy described a new species of vesselless
angiosperms as Sarcandra irvingbaileyi to honor his early association with
Professor Bailey and Bailey’s long interest in such primitive plants.

Professor Bailey was frequently asked for his opinion on aberrant
species or genera as plant materials came into the Arnold Arboretum for
determination. The taxonomic research published in the Journal of the
Arnold Arboretum often carries a few words of anatomical description
or notes on relationships supplied by Bailey or an acknowledgement of his
assistance in finding the right place for the material.

A report to the Dean of the Faculty of Arts and Sciences prepared by
Professor Bailey and entitled “Botany and its application at Harvard
has been called the “Bailey Plan” or the “Bailey Report.” It was the
subject of an acrimonious controversy which lasted over two decades while
Professor Bailey was active, and after his retirement. Although initially

6 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

submitted as a confidential report in 1945, its recommendations were
accepted by the Dean and an “unclassified” edition was prepared and
published. The report and plan which Professor Bailey himself described
at an annual meeting of the American Association for the Advancement
of Science (1946) created discussion which was to involve the faculty, the
Board of Overseers and its committee to Visit the Arnold Arboretum, the
Corporation, many teams of lawyers, and two decisions of the Supreme
Judicial Court of Massachusetts. In particular the wisdom of the recom-
mendations regarding the Arnold Arboretum was challenged and after
due consideration the Harvard Corporation as trustees of the Arnold
Arboretum voted in 1953 that the “plan contained in the Bailey Report
of June 1945 shall not apply to the Arnold Arboretum or to the principal
or income of its endowment.” The Harvard Corporation approved the
construction of a new building in Cambridge, the implementation of the
Bailey plan for other departments, and the removal of a portion of the
collections of books and specimens of the Arnold Arboretum from Jamaica
Plain to Cambridge. The transfer of these resources was approved with
the understanding that a working herbarium and a library were to remain
in Jamaica Plain. The division created an area of horticultural interest
in Jamaica Plain with the library and herbarium there devoted to the
study of plants under cultivation and associated with the living collections
and the greenhouses. The second unit to be formed in Cambridge would
contain the larger portion of the herbarium and library and would be
specialized in its application to the native floras of the world, and to basic
problems in plant anatomy and morphology.

Eventually, in 1967, the Supreme Court approved the division proposed
but in the many hearings, published accounts, and reports the name of
Professor Bailey was inevitably associated with the objectionable features,
Professor Bailey was deeply disturbed, as he felt many of the attacks were
directed at him personally and that he could not, or should not, reply.
His loyalty to Harvard and his interest in increasing the rdle and the
efficiency of use of botanical collections and botanical training at Harvard
he put honestly before a personal concern for single departments or
individuals.

Bailey’s travels were not many. He did not attend international meet-
ings and only infrequently attended those in the United States. His only
trip to Europe was a survey of forestry schools and training made with
Herman Spoehr. This resulted in the book, ‘The Role of Research in the
Development of Forestry in North America,” published in 1929. Dr.
Spoehr was on the staff of the Carnegie Institution of Washington, and
their cooperation resulted in Bailey’s being appointed a research associate
of the Carnegie Institution. For several summers Bailey regularly traveled
to Arizona to work at the desert laboratory. His published papers during
this period listed his association as either the Arnold Arboretum or Har-
vard University and as a research associate of the Carnegie Institution
of Washington. Typical of Bailey is the fact that his association with a
desert laboratory and studies on members of the Cactaceae in the decade

1968 | HOWARD, IRVING WIDMER BAILEY 7

1930-40 would be revealed again in his studies of the leaf-bearing Cac-
taceae in the 1960’s. Bailey made a trip to Cuba to visit the Atkins
Laboratory at Soledad and to Honduras for a study of forestry problems
associated with the United Fruit Company program in Central America.

In later years his summers were spent at his home in Norwell, where
a physical battle with the brush with scythe and axe and clippers gave
him a healthy skin color and a muscle tone envied by his colleagues.
These were happy moments in his life which he shared with his family.
A by-product of these summers usually was revealed around Christmas
when special friends were presented jars of beach plum jelly which Pro-
fessor Bailey made during the carefree summers, when fruits could be
picked with his adored grandchildren.

Professor Bailey married Helen Diman Harwood on June 15, 1911. He
is survived by his wife and their two sons, Harwood and Solon Irving II,
and five grandchildren.

Bailey was a member of the National Academy of Sciences, being an
Emeritus Member at the time of his death; the American Academy of
Arts and Sciences, having served as vice-president from 1947 through
1949; the American Association for the Advancement of Sciences, of
which he was a Fellow; the American Philosophical Society; the Botanical
Society of America, of which he was treasurer 1921-24, vice-president
1928, and president in 1945; the Society of American Foresters; the
American Society of Naturalists; the American Society of Plant Physiolo-
gists; the Torrey Botanical Club; the Society for the Study of Evolution;
the Royal Swedish Academy of Sciences; the International Association of
Wood Anatomists; the Linnean Society of London; the Ecological Society

of Plant Morphologists, of which he was president from 1960 to 1964,
and an honorary member of the Indian Botanical Society. Professor
Bailey served on the editorial boards of the American Journal of Botany
1915-18, the Proceedings of the Society of American Foresters 1914-16,
the Journal of the Arnold Arboretum 1941-1957. He has been listed in
Who’s Who in America and its various subdivisions, the International
Who’s Who and American Men of Science. In the third edition of Ameri-
can Men of Science, Irving Bailey was listed for the first time along with
his father, Solon Bailey. In 1931 Professor Bailey received an honorary
degree of Doctor of Science from the University of Wisconsin. In 1954
he was honored with the Mary Soper Pope award from the Cranbrook
Institute of Science which stated, “The wide influence of his fundamental
work on the biological processes involved in the formation, growth and
differentiation of cells in woody tissues has deeply affected the growth of
other fields, histological chemistry, taxonomy and evolution among them.
... As a teacher and administrator his inspiration and guidance have
contributed to the growth of many leaders in modern botany.”

In 1956 at the Fiftieth Anniversary meeting of the Botanical Society of
America Professor Bailey was among those honored with certificates
of merit, his stating “plant anatomist and inspiring teacher, for his

8 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

outstanding contributions on the structure of the cell wall and the his-
tology of the cambium and for his application of anatomy and morphology
to problems of evolution of angiosperms.”

The University of Syracuse also awarded him an honorary $.D. in 1961
with the citation: “Your name is synonymous with the role of research in
the development of forestry. Your dedication to truth, zeal in research,
and capacity to see the unexpected are gifts you have used to enhance
and ennoble all the lives you have touched. Your solid and creative
scholarship still guides and challenges forestry research. We honor you as
— ’ pre-eminent wood anatomist and botanist and a truly inspired
teacher

The degree he valued most highly, however, was received from his
Alma Mater at the 319th Commencement of Harvard, in 1955, on the
occasion of his retirement. “Irving Widmer Bailey, Doctor of Science,
today your university salutes you for your direction of botanical study
and for your accomplishment in searching, in the anatomy of plants, for
clues to the miracle of growth.”

BIBLIOGRAPHY *

— 1909 —
The structure of the wood in the Pineae. Bot. Gaz. 48: 47-55.

— 1910 —
Microtechnique for woody structures. Bot. Gaz, 49:

58.
Notes on the wood structure of the Betulaceae and Fagaceae. Forestry Quart.
8: 178-185

Anatomical characters in the evolution of Pinus. Am. Nat. 44: 284-293.
Oxidizing enzymes and their relation to “sap stain” in lumber. Bot. Gaz. 50:
142-147.

Reversionary characters of traumatic oak woods. Jbid. 374-380.

— 1911 —

The epi of the leaf-trace to the formation of compound rays in the lower
tyledons. Ann. Bot. 25: 225-241.

A ce Pityoxylon with marginal tracheids. Ibid. 315-325.

— 1912 —

The evolutionary history of the foliar ray in the wood of the dicotyledons; and
its phylogenetic significance. Ann. Bot. 26: 647-661

— 1913 —
The preservative treatment of wood, I. The validity of certain theories con-

cerning the gona of gases and preservatives into seasoned wood.
Forestry Quart. 11: 5-11.

The preservative ee of wood, II. The structure of the pit membranes in
the tracheids of conifers and their relation to the penetration of gases,

* Compiled in part from the bibliography in we to Plant Anatomy
“aoneys yer from 1955 on, made available by Lazella Schwarte

1968 | HOWARD, IRVING WIDMER BAILEY 9

liquids and finely divided solids into green and seasoned wood. Ibid. 11:
12-20.

— 1914—

Investigations on the phylogeny of the angiosperms, II. Anatomical evidences of

eduction in certain of the Amentiferae. (With E. W. Sinnott.) Bot. Gaz.
: 36-60.

Graded volume tables for Vermont hardwoods. (With P, C. Heald.) Forestry
Quart. 12: 5-23.

Some observations on the variation in length of coniferous fibers. (With H. B.
Shepard.) Proc. Soc. Am. Foresters 9: 522-527.

Primitive characters recalled by the ee bark disease and other stimuli.
(With J. S. Ames.) Science 39: 2

Some technical aids for the maa study of decaying wood. (With E. W.
Sinnott.) Phytopath. 4: 403.

sade isln on the phylogeny of the mg nerey III. Nodal ane and the

hology of stipules. (With E. W. Sinnott.) Am. Jour. Bot. 1: 441-453.

Inv contin on the phylogeny of the asitienbesti IV. The origin and dispersal

of herbaceous angiosperms. (With E. W. Sinnott.) Ann. Bot. 28: 547-600.

— 1915 —

Investigations on the phylogeny of the angiosperms, V. Foliar evidence as to the
ancestry and early climatic pe a of the angiosperms. (With E. W.
Sinnott.) Am. Jour. Bot. 2:

The effect of the structure of ease upon its permeability, < a tracheids of
coniferous timbers. Bull. Am. Railway Eng. Assoc. 174: 853.

The evolution of herbaceous plants and its bearing on certain otis of geology
and climatology. (With E. W. Sinnott.) Jour. Geol. 23: 289-306.

A botanical index of Cretaceous and Tertiary climates. (With E. W. Sinnott.)
Science 41: 831-834.

Sanio’s laws for the variation in size of coniferous tracheids. (With H. B.
Shepard.) Bot. Gaz. 60: 66-71

— 1916 —

The ree distribution of certain types of angiosperm leaves. (With E. W.
Sinno

Are Fetracetron Tj rochodendron and Drimys specialized or primitive types?

W. P. Thompson.) Mem. N. Y. Bot. Gard. 6: 27-32.

The aay of the bordered pits of conifers and its bearing upon the tension
hypothesis of the ascent of sap in plants. Bot. Gaz. 62: 133-142.

The significance of certain variations in the anatomical structure of wood.
(With R. P. Prichard.) Forestry Quart. 14: 662-670.

— 1917 —
The réle of the microscope in the ere iragt and classification of the “timbers
of commerce.” Jour. Forestry 15:

— 1918 —

Size variation in tracheary cells, I. A comparison between the secondary xylems
of vascular cryptograms, gymnosperms we fm eae (With W. W.
Tupper.) Proc. Am. Acad. Arts & Sci. 54: 1

Additional notes upon the angiosperms Konrath ieee and Drimys,

10 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

in which vessels are absent from the wood. (With W. P. Thompson.) Ann.
Bot. 32: 503-512

Report on method and apparatus for mechanically testing wooden parts of air-
planes. Wood Section, Materials Engineering Department, Bureau of Air-
craft Production, Dayton, Ohio.

— 1919 —
Depressed segments of oak stems. Bot. Gaz. 67: 438-441.
Structure, development and distribution of so-called rims or bars of Sanio. /bid.

Phenomena of cell division in the cambium of arborescent gymnosperms and
their cytological significance. Proc. Nat. Acad. Sci. 5: 283-285.

— 1920 —

The formation of the cell plate in the cambium of the higher plants. Proc. Nat.
Acad. Sci. 6: 197-200

The significance of the cambium i in the study of certain physiological problems.
Jour. Gen. Physiol. 2: 519-533.

Some relations between ants and fungi. Ecology 1: 174-189.

The Sarniones and its derivative a II. Size aot of cambial initials in
gymnosperms and angiosperms. Am. Jour. Bot. 7: 3 me:

eae, and binucleate cells. oon Gaz. 70: Pat

The cambium and its derivative tissues, III. A reconnaissance of cytological

phenomena in the cambium. Am. Jour. Bot. 7: 4

The feeding habits of Pseudomyrmine rs other ee geen W. M. Wheeler.)

Trans. Am. Phil. Soc. II. 22: 235-2

— 1922 —
The pollination of Marcgravia: a classical case of ornithophily? Am. Jour. Bot.
0-384.

The anatomy of certain plants from the Belgian Congo, with special reference
to myrmecophytism. Bull. Am. Mus. Nat. Hist. 43: 585-621.

The significance of the “foliar ray” in the evolution of herbaceous angiosperms.
With E. W. Sinnott.) Ann. Bot. 36: 523-533

Notes on neotropical ant-plants, I. Cecropia angulata sp. nov. Bot. Gaz. 74:

— 1923 —
Notes on neotropical ant-plants, II. Tachigalia paniculata Aubl. Bot. Gaz. 75:
27-41

Slime bodies of Robinia Pseudo-Acacia L. Phytopath. 13: 332-333.
The cambium and its derivative tissues, IV. The increase in girth of the cam-
bium. Am. Jour. Bot. 10: 499-509.

— 1924 —
Notes on neotropical ant-plants, III. Cordia nodosa Lam. Bot. Gaz. 77: 32-49.
The problem of identifying the wood of Cretacous and later dicotyledons: Para-
phyllanthoxylon arizonense. Ann. Bot. 38: 439-451.
Abnormalities of ring growth and cell structure. In “studies on the spruce bud-
worm,” Tech. Bull. Canadian Dept. Agric. 37: 58-61.
So-called bars or rims of Sanio. Bot. Gaz. 78: 124, 125.

1968 | HOWARD, IRVING WIDMER BAILEY 11

— 1925 —
Some salient lines of specialization in tracheary pitting, I. Gymnospermae. Ann.
Bot. 39: 587-598
The “spruce budworm” biocoenose, I. Frost rings ie usa of the chronology
of specific biological events. Bot. Gaz, 80: 93-1
The “spruce budworm” biocoenose, II. Structural ea in Abies bal-
samea, Ibid, 300-310.
— 1929 —
The role of research in the ae of forestry in North America. (With
H. A. Spoehr.) New Yor
— 1930 —
The cambium and its derivative tissues, V. A reconnaissance of the vacuome in
living cells. Zeitschr. Zellforch. Mikr. Anat. 10: 651-682.

— 1931 —
The cambium and its derivative tissues, VI. The effects of hydrogen ion con-
centration in vital staining. (With C. Zirkle.) Jour. Gen. Physiol. 14: 363-
383.

— 1932 —
Preliminary notes on cribriform and vestured pits. Tropical Woods 31: 46-48.

— 1933 —
The cambium and its derivative tissues, VII. Problems in identifying the wood
of Mesozoic coniferae. Ann. Bot, 47: 145-157
The cambium and its derivative tissues, VIII. Structure, distribution, and diag-
nostic significance of vestured pits in dicotyledons. Jour. Arnold Arb. 14:
259-273.

— 1934 —
The cambium and its derivative tissues, Ix. Structural variability in the red-

The cambium and its derivative tissues, X. Structure, optical properties and
chemical composition of the so-called middle lamella. (With T. Kerr.)
Ibid, 327-349.

— 1935 —

The visible structure of the secondary wall and its significance in physical and
chemical investigations of tracheary cells and fibers. (With T. Kerr.) Jour.
Arnold Arb. 16: 273-300.

— 1936 —

The problem of differentiating and ae ms Oi — fiber-tracheids and

libriform wood fibers. Tropical Woods 45:

— 1937 —
The orientation of cellulose in the pagina! wall of tracheary cells. (With M.
. Vestal.) Jour. Arnold Arb. 18: 185-1
The significance of certain wood-destroying fungi in the study of the enzymatic
hydrolysis of cellulose. (With M. R. Vestal.) Jbid. 196-205

12 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

The structural variability of the secondary wall as revealed by “lignin” residues.
With T. Kerr.) Jbid. 261-272

— 1938 —
Cell wall structure of rie plants. Ind. Eng. Chem. 30: 40-47.
The occurrence of Cedrus in the auriferous Eas of California. (With E. S.
Barghoorn, Jr.) ria Jour. Bot. 25: 641-647

— 1939 —
The microfibrillar and microcapillary structure of the cell wall. Bull. Torrey Bot.
Club 66: 201-213

— 1940 —
A useful method for the study of pollen in peat. (With E. S. Barghoorn, Jr.)
Ecology 21: 513, 514.
The walls of plant cells. A. A. A. §. Publ. 14: 31-43.

— 1941 —

The comparative morphology of the Icacinaceae, I. Anatomy of the node and
internode. (With R. A. Howard.) Jour. Arnold Arb. 22: 125-132.

The comparative morphology of the Icacinaceae, II. Vessels. (With R. A.
Howard.) Jbid. 171-187.

The comparative morphology of the oe ai Ill. Imperforate tracheary
elements and xylem parenchyma. (With R. A. Howard.) Jbid. 432-442.
The comparative morphology of the [acnacene, IV. Rays of the secondary

xylem. (With R. A. Howard.) Jbid. 556-56
Brossiantha, a new genus of Hippocrateaceae — New Guinea. (With A. C.
mi th.) Ibid. 389-394.

— 1942 —

Seedlings as experimental material in the ay of “redwood” in conifers. (With
H. F. Wershing.) Jour. Forestry 40: 411-414.

Identification and physical condition of the stakes and wattles from the fishweir.
With E. S. Barghoorn, Jr.) Chapter 6 in the Boylston Street Fishweir.
Papers of the Robert S. Peabody Foundation for Archaeology Vol. 2
Philips Academy, Andover, Mass.

Degeneriaceae, a new family of flowering plants from Fiji. (With A. C. Smith.)
Jour. Arnold Arb. 23: 356-365.

The significance of x-rays in studying the orientation of cellulose in the secondary
wall of tracheids. (With E. E. Berkley.) Am. Jour. Bot. 29: 231-241.
pee and illustrations of Triplaris surinamensis, in Wheeler, Wm. M..,
: s of neotropical ant-plants and their ants. Bull, Mus. Comp. Zool.

“ 0-53,

7

= 1043
Some misleading terminologies in the literature of “plant tissue culture.” Science
98: 539, 540.

Arb. 24:

The comparative meee of the Winteraceae, I. Pollen and stamens. (With
C. G. Nast.) Ibid. 340-346.

The family or (With C. G. Nast & A. C. Smith.) Jour. Arnold
190-

1968] HOWARD, IRVING WIDMER BAILEY 13

The comparative morphology of the Winteraceae, II. Carpels. (With C. G.
Nast.) Ibid. 472-481.

— 1944 —

The Rael morphology of the Winteraceae, III. Wood. Jour. Arnold
Arb, 25: 97-103.

The meets morphology of the Winteraceae, IV. Anatomy of the node and
vascularization of the leaf. (With C. G. Nast.) Jbid. 215-221.

The comparative morphology of the Winteraceae, V. Foliar epidermis and
sclerenchyma. (With C. G. Nast.) Jbid. 342-348.

The development of vessels in angiosperms and its significance in morphological
research. Am. Jour. Bot. 31: 421-428.

— 1945 —

The comparative morphology of the Winteraceae, ee mars and conclu-
sions. (With C. G. Nast.) Jour. Arnold Arb. 26:

Morphology and Lee ge of Trochodendron or aieedtone I. Stem,
root and leaf. (With C. G. Nast.) bid. 143-154.

ee and relationships of Trochodendron and Tetracentron, II. Inflo-

cence, flower and fruit. (With C. G. Nast.) bid. 267-27

Botaitiy and its applications at Harvard: A report to the Dean of the Faculty

of Arts and Sciences. I-IV, 1-93. June.

— 1946 —
Morphology of Euptelea and comparison with Trochodendron. (With C. G.
Nast.) Jour. Arnold Arb. 27: 186-192.

— 1948 —
oT and relationships of Jilicitum, Schisandra ae Rats I. Stem
d leaf. (With C. G. Nast.) Jour. Arnold Arb. 29:
Amborl Boe ge Baill. A new type of vesselless eiicion. (With B. G.
my.) Ibid. 215.
Ane Teliiode Baill. A new morphological type of vesselless dicotyledon.
(With B. G. L. Swamy.) Jbid. 245-254.

— 1949 —
Origin of the angiosperms: need for a broadened outlook. Jour. Arnold Arb. 30:
64-70.

The bd gpa = relationships of Cercidiphyllum. (With B. G. L. Swamy.)
Ibid. 1

The morphology and relationships of Austrobaileya. (With B. G. L. Swamy.)
Ibid. 2

— 1950 —

Sarcandra, a vesselless genus of the Chloranthaceae. (With B. G. L. Swamy.)
Jour. Arnold Arb. 31: 117-129.

The morphology and relationships of the Monimiaceae. (With L. L. Money &
B. G. L. Swamy.) Ibid. 372-404.

The administration of Harvard’s endowed botanical institutions. A report to

he Coordinating Committee for the Biological Sciences of the Board of

elon 62 pages. (With P. C. Mangelsdorf.)

14 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

— 1951 —
The use and the abuse of anatomical data in the study of phylogeny and classifi-
cation. Phytomorphology i:
Coéperation versus isolation in botanind research. Chron. Bot. 12: 126-133.
The amps carpel of dicotyledons and its initial trends of nen:
B. G. L. Swamy.) Am. Jour. Bot. 38: 373-379.

— 1952 —
Biological processes in the formation of wood. Science 115: 255-259.

— 1953 —
Evolution of the tracheary tissue of land plants. Am. Jour. Bot. 40: 4-8.
A new Fijian species of Calyptosepalum. (With A. C. Smith. y pce Arnold Arb.
34: 52-64.

The morphology and Seow nae: of Idenburgia and Nouhuysia. (With B. G.
amy.) Ibid. 77-85.
The anatomical approach to the study of genera. Chron. Bot. 14: 121- 125,

— 1954—
Contributions to plant anatomy. Chron. Bot. 15: i-xxvi, 1-262. 23 plates +
port. Waltham.

A fourth type of nodal anatomy in dicotyledons, illustrated by Clerodendron

trichotomum Thunb. (With M. P. F. Marsden.) Jour. Arnold Arb. 36:
1-51.

— 1956 —

Nodal anatomy in retrospect. Jour. Arnold Arb. 37: 269-287.

The relationship between seen el ser of New Caledonia and Nouhuysia of
New Guinea. Jbid. 360-

Aggregations of microfibrils ve their yey a he the secondary wall of
coniferous tracheids. Am. Jour. Bot. 44: 415-4

Die Struktur der Tiipfelmembranen bei den Paces der Koniferen. Holz als
Roh und Werkstoff 15: 210-21

— 1957 —

The ee anatomy and the primary vascular Pras of the Calycanthaceae.
( Fahn.) Jour. Arnold Arb. 38: 107-1

The aolenee and limitations of wood anatomy in ae study of the phylogeny
and classification of angiosperms. Ibid. 243-254.

Additional on the vesselless dicotyledon, Amborella trichopoda Baill. Ibid.

— 1958 —
pre oe : — outlook in cell wall terminologies. Phytomorphology 7:

The sae of tracheids in relation to the movement of liquids, —
and undissolved gases. In: The physiology of forest trees. pp.

— 1960 —
Some useful techniques in the study and
ly and interpretation of
Jour. Arnold Arb. 41: 141-151. - n of pollen morphology.

1968 | HOWARD, IRVING WIDMER BAILEY 15

Comparative anatomy of the leaf-bearing Cactaceae, I. Foliar vasculature of
Pereskia, Pereskiopsis and Quiabentia. Ibid. 341-356.

— 1961 —

Comparative anatomy of the leaf-bearing Cactaceae, II. Structure and distribu-
tion of sclerenchyma in the phloem of Pereskia, Pereskionsis and Quiabentia,
Jour. Arnold Arb. 42: 144-156.

Comparative anatomy of the leaf-bearing Cactaceae, III. Form and distribution
of crystals in Pereskia, Pereskiopsis and Ouiabentia. Ibid, 334-346.

— 1962 —

Comparative anatomy of the leaf-bearing Cactaceae, IV. The fusiform initials
of the cambium and the form and structure of their derivatives. (With L
M. Srivastava.) Jour. Arnold Arb. 43: 187-202.

Comparative anatomy of the leaf-bearing perpen V. The secondary phloem.
(With L. M. Srivastava.) Ibid. 234-2

Comparative anatomy of the leaf-bearing Cacacene, VI. The xylem of Pereskia
sacharosa and Pereskia aculeata, Ibid. 376-

— 1963 —
Comparative anatomy of the leaf-bearing Cactaceae, VII. The xylem of the
Pereskias from Peru and Bolivia. Jour. Arnold Arb. 44: 127-137.
Comparative anatomy of the leaf-bearing Cactaceae, VIII. The xylem of Pere-
skias from southern Mexico and Central Ametica. Ibid. 211-221.
Comparative anatomy of the leaf-bearing Cactaceae, IX. The xylem of Pereskia
grandifolia and Pereskia bleo. Ibid, 222-231
Comparative anatomy of the leaf-bearing Cactaceae, X. The xylem of Pereskia
colombiana, Pereskia guamacho, Pereskia cubensis, and Pereskia portulaci-
folia. Ibid, 390-401

— 1964 —
is er anatomy of the leaf-bearing Cactaceae, He The xylem of Pereski-
and Quiabentia. Jour. Arnold Arb, 45: 140-
Compas anatomy of the leaf-bearing Cactaceae, e Preliminary observa-
tions upon the structure of the epidermis, stomata and cuticle. /bid. 374—
389.

— 1965 —

Caadianidve anatomy of the leaf-bearing Cactaceae, XIII. The occurrence of
water-soluble anisotropic bodies in air-dried and alcohol-dehydrated leaves
of peers and Pereskiopsis. Jour. Arnold Arb. 46: 74-85.

Comparative anatomy of the leaf-bearing Cactaceae, XIV. Preliminary observa-
tions on the vasculature of cotyledons. /bid. 44 .4 SZ;

Comparative anatomy of the leaf-bearing Cactaceae, XV. Some preliminary

observations on the occurrence of “Protein Bodies. ” Ibid. 453-464.

— 1966 —
Comparative anatomy of the leaf-bearing Cactaceae, XVI. The development of
water-soluble crystals in dehydrated leaves of Pereskiopsis. Jour. Arnold
Arb. 47; 273-287.
The signiGcance of the reduction of vessels in the Cactaceae. bid. 288-292.

16 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

INFLORESCENCE IN NANNORRHOPS RITCHIANA (PALMAE)

P. B. Tomiinson AND H. E. Moore, Jr.!

PALM INFLORESCENCES IN HERBARIA are largely represented by small
fragments. Hence it is not surprising that botanical literature gives little
aid to an overall appreciation of flowering processes in palms. Also the
multiplicity of terms used in describing the parts of reproductive branches
of palms — spathe, spadix, spathelet, rachilla, involucel, etc. does little
to reveal the basic pattern of construction. These t terms may have some
use in description but lack of consistency in their application diminishes
their value. As strict morphological terms they have no significance.
There is no published account which clearly outlines the principles involved
in the construction of palm inflorescences.

The present article aims at a clarification of some inflorescence features
in palms by describing the unspecialized type found in Nannorrhops
ritchiana (Griffith) Aitchison,? as it is cultivated in South Florida. A
wide familiarity with inflorescence construction in palms both in cultiva-
tion and in the field shows that Nannorrhops can be used as a model for
a general understanding of other palm inflorescences.

It is necessary first to appreciate that the flowering phase is a physio-
logical state and it was as such that the term “inflorescence”? was applied
by earlier botanists, as Rickett (1944) points out, although the term
subsequently has been adopted largely as a morphological one. A general
discussion of shoot construction in palms and other arborescent mono-
cotyledons (Tomlinson, 1964) has emphasized that each axis passes
through three physiological states during its development, these phases
being recognized by combinations of morphological features. There is a
gradual transition from one phase to another and they may overlap. An
initial juvenile phase, during which establishment growth (Tomlinson &
Zimmermann, 1966) takes place, is followed by an adult vegetative phase,
in turn followed by a reproductive phase, the physiological state of
inflorescence. Each axis may grow from a seed, or from a lateral sucker.

In palms inflorescence is expressed in three distinct ways. Most com-
monly the reproductive parts are borne on lateral branches maturing in

* Research on the taxonomy, morphology, and anatomy of palms is supported by

individual research grants from the National Science Foundation, grant GB 2991
and its ceutan acs GB 5762-X (P.B.T.) and GB 3528 (HE.

*The specific epithet is derived from Chamaerops ritchia na Ww. Griffith but is
often spelled ritchieana. Though Griffith named the species in honor of Dr. Ritchie,
he used the shortened epithet in the original publication (Calcutta Jour. Nat. Hist.
ie 342, 343. 1844) and in the posthumously published Palms of British East India

name was intentional rather than a typographical error, the original spelling is re-
tained. :

1968 | TOMLINSON & MOORE, NANNORRHOPS 7

acropetal order and the flowering process does not inhibit growth of the
axis. Corner (1966, p. 124) applies the term pleonanthic to this method
of flowering. Morphologists describe such palms as having “lateral in-
florescences.”” In a second group the flowering process is hapaxanthic,
vegetative growth is completely substituted by an axis with an acropetally-
developed series of reproductive branches. The axis may elongate to
produce a conspicuous “terminal inflorescence” as is most familiar in
Corypha, Nannorrhops and species of Metroxylon. Here the hapaxanthic
condition is obvious. Otherwise, as in Raphia and some other Lepido-
caryoid palms, growth of the axis ceases with the production of lateral
reproductive branches in the axils of relatively unmodified leaves in such
a way that a conspicuous terminal “inflorescence” is not evident.

Another hapaxanthic type is so distinctive as to merit a third category.
This is represented by and characteristic of the small but very natural
subfamily Caryotoideae (Moore, 1963). Here the reproductive phase
inhibits vegetative growth but in such a way that development of lateral
flowering axes is almost invariably in a basipetal direction from suc-
cessively older dormant axillary buds. In effect the whole vegetative axis
becomes converted into a gigantic “inflorescence’’ with basipetal matura-
tion. At least one exception to this rule is found in the recently described
Arenga retroflorescens (Moore & Meijer, 1965).

Only in single-stemmed palms does hapaxanthic flowering terminate the
existence of the individual, as in Corypha. These individuals may be
described as monocarpic. Otherwise continued substitution of old axes
by new ones arising as basal suckers continues the life of the individual,
as in many Lepidocaryoid and Caryotoid palms. Substitution of axes in
Nannorrhops is distinctive as described below. In these hapaxanthic
palms the individuals are polycarpic.

The present article describes the morphological features found in the
reproductive phase in Nannorrhops. This develops a gigantic terminal

(1944) to describe a much-branched inflorescence. This article com-
plements the detailed accounts of floral anatomy by Gupta (1960) and
Morrow (1965), neither of whom were concerned with overall inflorescence
structure.

MATERIAL AND METHODS

Several clumps of Nannorrhops ritchiana, a member of the fan-leaved
Coryphoideae, were available at Fairchild Tropical Garden and the Plant
Introduction Station, Old Cutler Road, Miami, Florida. Observations
have extended over a period of years and the expansion of several
inflorescences has been watched. Two inflorescences (referred to as shoot
1 and shoot 2) have been cut down and examined in quantitative detail.
Approximate measurements of essential parts on each of these shoots are
presented graphically in Ficures 1 and 2. In these figures, ordinates are
arbitrary leaf numbers counted from the cut. Photographic (Fics. 35-42)

18 . JOURNAL OF THE ARNOLD ARBORETUM [ VoL. 49

and other illustrations are largely from these two shoots. Fluid-fixed
material of shoot 2 has been preserved for future anatomical study.

OBSERVATIONS

General habit. In its native habitat in northwest India and Afghan-
istan, Nannorrhops was described by Blatter (1926) as a low-growing
gregarious shrub with a prostrate “rhizome,” but exceptionally growing
erect, as indicated by Gupta’s photograph. This erect habit is the one
largely adopted by specimens in cultivation in South Florida, no doubt
as a result of the relatively luxuriant conditions for growth. In this
facultative ability it recalls Serenoa repens.

The vegetative axis, whether erect or decumbent, is branched. Branch-
ing of the creeping axes and in the basal part of the erect axes is apparently
axillary, suckers developing in a manner which again bears comparison
with Serenoa. On the other hand distal branching of the aérial axes of
specimens in South Florida involves an initial equal forking which sug-
gests, at least superficially, a true dichotomy. This we have illustrated
elsewhere (Tomlinson & Moore, 1966). This dichotomous-like branching,
previously familiar only in Hyphaene (Schoute, 1909), seems more com-
mon in palms than hitherto suspected (Tomlinson & Moore, 1966;
Tomlinson, 1967). Forking of the axis is initiated equally, but the two
axes behave differently. One branch rapidly proceeds to inflorescence,
producing the terminal panicle. The other branch repeats the forking,
apparently in a plane at right angles to the first fork. Repeated forking

with overtopping of a reproductive shoot by a vegetative shoot may
continue for some time.

Reproductive phase. The onset of the reproductive phase involves
direct transformation of one fork into an “inflorescence.” Early stages
are revealed by the asparagus-like aspect of the crown (Fic. 35). Sub-
sequently, as the axis elongates the three main morphological features
of the inflorescence become evident (Fics. 36-38). These are (a) gradual
modification of leaves (b) a narrowing and initial extension of internodes
(c) development of axillary branches. The first two changes are quite
gradual, the latter change is abrupt. At maturity the terminal panicle,
2-4 meters high, rises conspicuously above the vegetative clump (FIGs.
39, 40, 42).

Gradual modification of leaf shape along the main reproductive axis
for both shoots is indicated in Ficures 1 and 2. A continuous leaf series
from shoot 1 is shown in Figure 41. No change in leaf insertion is
involved, the 2/5 spiral phyllotaxis of the vegetative part of the shoot
is continued into the inflorescence. Normal foliage leaves have a distinct
blade, petiole, and sheath (Fics. 1, 14, 15), but these undergo a pro-
gressive reduction along the reproductive axes; leaves are shortened,
petiole first and then the blade are diminished and finally eliminated until
distal leaves are represented by the sheath alone (Fics. 5-13), the vestigial

1968] TOMLINSON & MOORE, NANNORRHOPS .

160} (Th

120} \
80} \

YIN
4ok NN
O 5 10
<< aes Fologe afd ----y.- artis
Transitional leaves
12
10 } SR
st et a, eh :
no x
| Vv
Al ; Diameter
oe Internode
0 5 10 a) 36
Leaf number @rbitrary)
1G. 1. Nannorrhops. Quantitative details of leaves and main axis in shoot 1.

ABOVE: approximate length of sheath, petiole and blade of successive leaves
along main axis, leaves numbered arbitrarily from level where the shoot was cut.
Stippled area to right indicates branched portion of axis. BeLow: length and
diameter of corresponding internodes.

20 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

petiole plus blade appearing only as a short distal appendage (Fics.
11-13). This minute appendage, which establishes the plane of symmetry,
is found even in the smallest bracts (e.g. Fics. 28-30).

cMs Length
240} | | 160 Hl Ist-order branch
200} 1}

160
120
8O}T
40

U 5 10 5 20

one ee uee Foliage and -----¥-- Bracts ------------
Transitional leaves

14 oN

10 wg Ue mt bee

a é en.

| \,

Sy Diameter ~

4

2 Internode

0 = 10

1S 20 20 30 a
Leaf number (arbitrary)
. 2. Nannorrhops. Details of shoot 2, as for shoot 1. ABOVE RIGH
proximate overall length of first-order branches subtended by leaves indicated
below.

Associated with these changes in leaf shape and size is a gradual and
uniform decrease in internode diameter; internode length, on the other
hand initially increases, subsequently declining uniformly (Fics. 1, below

2, below). Maximum internode length is about the level of exsertion of

1968 | TOMLINSON & MOORE, NANNORRHOPS 21

the first branch but the agreement is not very precise and may be quite
incidental.

The sheath of each reduced leaf remains a closed tube, as in the foliage
leaves. Nannorrhops has the Hyphaene-type of foliage leaf base, in
which a dorsal rhombohedral cleft (Fic. 5, detail to left) accommodates
the mechanical stresses imposed by expansion of internal tissues and
organs (Tomlinson, 1962). Progressive narrowing of the axis as the
inflorescence develops gradually eliminates mechanical stresses on suc-
cessive leaf sheaths. Consequently the dorsal cleft gradually disappears;
in shoot 2, leaf number 10 is the last leaf to possess this cleft (Fic. 6,
detail to right). Rarely a dorsal cleft is regained in some of the lower
bracts, presumably to accommodate expansion induced by developing
axillary branches.

In contrast to the above changes, initiation of lateral branches is abrupt.
Normally there are no aborted buds below the level of first branching.
The first branch is usually longest, subsequent branches being progressively
reduced in proportion to the distal decline (Fic. 2, above right). Details
of these branches may now be examined.

Branches and bracts. The main vegetative axis shows a gradual
transformation as it becomes reproductive, most obviously in the long
transitional series from foliage leaves to bracts. In contrast to the
vegetative axis a series of elaborate branches is initiated on the repro-
ductive axis. One of these branches is shown in Ficure 3. Leaves on
main and subsequent axes which subtend branches are defined as bracts.
First-order bracts, i.e. bracts on the main axis which subtend first-order
branches, are simply part of the transition series illustrated in FicurEs
1 and 2. From this series it is clear that bracts do not differ from foliage
leaves in any fundamental morphological way. They are modified leaves
distinguished only by position and definition.

Branching is straightforward and uniform and may be repeated up to
five times. A simple one to one relationship between bract and branch
is maintained. Each bract (br) subtends an axis (ax) of the next higher
order. The main axis (axo) bears first-order bracts (br;) which subtend
branches of the first order (ax;). These in turn bear second-order i
(br2) which subtend second-order branches (ax2) and so on. On t
proximal parts this may be repeated so that fifth-order bracts (brs) i
tend fifth-order branches (ax;). The diameter and length of successive
orders of branches are progressively reduced. TaBie I is a series of
measurements from a lateral branch (cf. Fic. 3) which illustrates this
reduction. This progressively reduced branch system ends in flower
clusters which are themselves condensed branch systems as described
below. Axes bearing flower clusters may be referred to as rachillae since
this is a convenient and widely understood term. A rachilla is a visible
axis which ends each branch, regardless of its order. This is shown most
clearly in Fic. 17 which represents the end of a second-order branch.
Even the main axis itself (axo) ends in a rachilla. However, rachillae

22 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Table I. S1zE OF BRACTS AND BRANCHES OF SUCCESSIVE ORDERS (CMS.)

OVERALL TOTAL NO. BrACT NUMBER; LENGTH AND DIAMETER
AxIs LENGTH BRACTS PROPHYLL 3 5 10 15 20 25
3ax: (ON aXo) 160 ese
LENGTH 15 21 16 5 2
DIAMETER 3 15 1 0.4 0.1
3ax, (on 3ax,) 85 22
LENGTH 8 9.5
DIAMETER 12 0.3 0.2
3ax, (on 3ax,) 30 12
LENGTH 2 2 0.3
DIAMETER 0.4 0.2 0.1
3ax, (on 3ax,) 10 2 0.5 0.2
0.1 <O.1
RACHILLA ca. 25 bracts, each subtending a flower cluster; distal clusters
2ax, (on 3ax,) aborted; bracts each 2-3 mm. long, diameter of axis <1 mm.

wide at base.

Notes: Each axis referred to is third along the axis of previous highest order
ie. 3ax, on 3ax,_,, except for the ultimate rachilla which is the second of the fifth-
order branches (2ax;) borne on 3ax,. Each axis ends in a rachilla, the approximate
total number of bracts listed for each axis does not include the distal bracts sub-

te er clusters on this rachilla. The third bract on 3ax, is the first on the
terminal rachilla and sub s first flower cluster lon y one axis bracts
c ho light increase in length before the distal decrease, e.g. 3ax he

length of these bracts, however, includes a pronounced terminal appendage (e.g.
Fics, 20-22).

are not strictly the ultimate branch units because the flower clusters have
to be interpreted as such, as is shown below.

Length of successive branches is progressively reduced, but not in the
regular way recorded for the first order branches in Fic. 2, above right.
Proximal branches of any one order tend to be short, presumably because
they have limited space in which to develop. Therefore the largest branches
tend to be in the middle of each axis (Fic. 3). Bra¢ts, unlike branches,
show more constant reduction along each axis and from the axis of one
order to that of the next higher order. Some quantitative details which
illustrate this reduction are indicated in Taste I. Distally, bracts are
narrow, tubular organs with asymmetric mouths and they bear little
obvious relation to foliage fan-leaves, but their ultimate morphology is
evident from the transitional series along main and subsequent axes (Fics.
5-13 along axo; 18-22 along ax;; 23 and 25 on axs).

Although they are progressively reduced in size, bracts overlap in all
but the last stages of development and their value in protecting axillary
units which develop within is quite obvious (e.g. Fic. 16). This protective
function becomes less significant in successively narrower axes. Th

1968 |

\
\
\
‘

\

20 <m

TOMLINSON & MOORE, NANNORRHOPS

NV;

RY DL)
\ Fs ig Say
AS

Y Vex

oN
7 Z He . nt - F
ma hare is LZ LD <

the one subtended

). Fic. 3. Lowest first-order lateral from shoot 2,
ranch (lax.) on the above axis, subtending bract cut off t

First-order lateral branch (ax,
ail of insertion of first-order b

Fics. 3 and 4. Nannorrhops.

by leaf 18 in Fic. 2. Fic. 4. Det

adnate to ax,.

rR
Ge

o show ax.

[voL. 49

JOURNAL OF THE ARNOLD ARBORETUM

24

1968 | TOMLINSON & MOORE, NANNORRHOPS 25

initiation of branches is clearly in an acropetal sequence but their expan-
sion seems to be basipetal, so that distal units of each branch order tend
to expand as soon as they project beyond the mouth of the enveloping
bracts (Fic. 37). Further developmental information has not been sought.

Arrangement of bract and branch on successive units follows a regular
pattern. At the base of each branch, bracts (and consequently axillary
branches) are distichously arranged, but distally there is a gradual change
to a spiral arrangement. This change from distichy to polystichy is most
evident on first order branches. Due to compression the distal spiral
arrangement is often obscure. In fact the distribution of parts on distal
branches tends to conform to the requirements of available space and
any inherent phyllotactic arrangement is suppressed. The plane of
distichy of distal branches (ax;—ax;) is always at right angles to the
dorsiventral plane of their subtending bracts. This change in the plane
of insertion allows greater space for expansion.

The first leaf on each branch is, by definition, a prophyll. In the two
lowest orders of branch (ax; and ax2) this prophyll is empty, subtending
no branch. Rarely on ax; the bract after the prophyll is also empty.
Prophylls show some of the features normal for the leaf in this position
in most monocotyledons. It is apparently inserted in the adaxial (ados-
sierte) position even though the plane of insertion of subsequent leaves
is at right angles to this. It is indistinctly 2-keeled. This is largely shown
by two sub-lateral teeth at the mouth of the bract, rather than by a single
dorsal tooth (Fics. 18, 23). These prophyllar features are least evident
on higher orders of branches. Here the branch subtended by a prophyll
is also in a lateral position so that the change in plane of distichy starts
with the prophyll.

Adnation of branch to main axis is a constant feature of the system
(Fics. 4 and 24). This probably reflects intercalary growth of the inter-
node after a branch primordium has been initiated, together with com-
pression of the developing system. Even beyond the level where a branch
diverges from its parent axis the common interface is flattened. Adnation
is more nearly complete and more pronounced on successively higher
orders of branches. In the lower orders, a branch is adnate to the parent
axis for a little over one-half the length of the internode above (Fic. 4),
and the union is concealed within the enveloping spathe. Distally there
is a progressive shortening of bract in relation to the internode above,
the branch is adnate more or less completely throughout this internode
so that the union is obvious (Fic. 24). Prophyll of branch and the next

Fics. 5-15. Nannorrhops. Transitional leaves and bracts from the main axis
of shoot 2. This series includes every third leaf from shoot 2 starting with leaf
F :

from dorsal side showing cleft). Fic. 7. Leaf number 13. Fics. 8-13, leaves
number 16, 19, 22, 25, 28, and 31, respectively. Fics. 14, 15 (inset). Fic. 15 is
leaf number 7 (Fic. 5) drawn on a smaller scale together with leaf number 1
(Fic. 14) to compare size of foliage leaves.

[ VoL. 49

JOURNAL OF THE ARNOLD ARBORETUM

26

1968 | TOMLINSON & MOORE, NANNORRHOPS 27

leaf above on the parent axis are at about the same level. In distal units,
therefore, branches appear to diverge just below a leaf insertion.

Rachillae and Bracteoles. Flowers are usually in clusters of three
on all terminating axes. The flower clusters (Fic. 28) are in an irregular
spiral but so compressed to fit available space that any regular phyllotactic
arrangement is lost. This regularity might be sought in the early stages
of rachilla development. Each flower cluster is subtended by a scale-like
bract 2-3 mm. high. The dorsal tooth is still distinct and establishes the
plane of symmetry of the bract. This is still a tubular organ, enclosing
the axis at its base but somewhat like an inverted funnel, the mouth
being expanded by the developing flowers. Within the bract subtending
each cluster is a series of bracteoles and flowers. We use the term bracteole
simply to denote bracts associated with flowers. Each bracteole is identical
with the bract subtending the whole cluster, but smaller. The arrangement
of flower buds and bracteoles is indicated by the dissections of a flower
cluster in Figures 29-32, all drawn from the same aspect, in which the
bract and each bracteole and flower is removed in turn. These flowers
and bracteoles are inserted more or less in one plane but compressed

Oo

parts might be suspected, but if present it is obscured by compression.
It can only be demonstrated by anatomical studies which we have yet
to undertake.

The first bracteole, as indicated by its dorsal tooth, is inserted per-
pendicular to the dorsiventral plane of the subtending bract and encloses
all organs within. It is also opposite the largest flower. Enclosing the
base of the two innermost flowers is a second bracteole opposite the
first. The second flower is opposite the first flower. The third bracteole,
opposite the second, encloses the third flower. Between this third flower
and its attendant bracteole is a minute bud (Fics. 31, 32) which
represents the continuing axis of the system and a potential fourth flower
bud. This flower does not seem to develop.

There can be several interpretations of this flower cluster. Lacking
anatomical evidence we tentatively interpret it as a cincinnus. The

16-27. Further details of ~—— me bracts of Nannorrhops. Fic. 16.
r branch fro h

shortly before anthesis of first flowers. This includes branches of next three
higher orders (axs, ax,, and ax;). Fics. 18-22. aa bracts (br.) from a first-
order branch (ax,) as in Fic. 3. Fic. 18. First bract (prophyll) (1 br.). Fre. 19.

Second bract (2 br.). Fic. 20. Third bract (3 br.). Fic. 21. Fourth bract (4 br.).

Fic. 22. Thirteenth bract (13 br») from this branch. Fic. 23. Three lowest bracts
(1-3 br,) from second- arr branch (ax.), the first (1 br;) is a ec slightly
2-keeled adaxially. Fics. 24 and 25. Details of distal branches. Fic Third-
order branches (ax;) ae is second-order branch (ax.). The bract ‘i br;) sub-
tending the uppermost branch is cut off. It is shown in Fic. 25. Fics. 26 and 27.

Flower clusters. Fic. 26, Left-handed flower cluster, flowers unopened. Fic. 27.

Right-handed flower cluster with oldest flower expan nded.

JOURNAL OF THE ARNOLD ARBORETUM

[ VOL. 49

1968 | TOMLINSON & MOORE, NANNORRHOPS 29

processes involved in ramification of proximal parts of the branch system
are continued into its ultimate axes which are, however, much condensed.
Sympodial is now substituted for monopodial branching. The flower
cluster (Fics. 33, 34), therefore, represents a series of condensed branches
of successively higher order, each branch bearing one bract (bracteole)
and terminating in a flower (Fic. 34). Each successive branch unit orig-
inates in the axil of the bracteole of the previous order. As in the
lower orders of branches, adnation and the requirements of close-packing
modify the arrangement of parts. The position of the first unit at right
angles to the plane of insertion of the subtending bract corresponds to
that found proximally in branches of third and higher orders. Close-
packing maintains the second and third units in this plane, but alternately
on opposite sides. From its position each bracteole is a prophyll and more
or less maintains the normal relation between prophyll and parent axis
(Fic. 33). Regarding the flower cluster as a sympodium is the simplest
interpretation which fits the available facts. We appreciate that additional
information from development and anatomy may cause us to modify this
opinion.

Flower clusters may be right- or left-handed, depending on whether
the largest (lowest) flower is exserted to left or right of the dorsiventral
plane of the subtending bract. The flower cluster in Ficures 29-32 is a
right-handed one, a left-handed one like that in Fic. 26 is the mirror
image of this. Right- or left-handed clusters, appear at random along
the rachilla (Fic. 28) and they may owe their distribution entirely to
chance. It is quite possible, on the other hand, that the position of the
first flower in each cluster is determined by a direct requirement of
available space in close packing of the developing rachilla.

Flowers. Flowers (Fic. 27) are perfect and expand in order of their
age. The structure and vascular anatomy of the flower of Nannorrhops
has been dealt with in great detail by Gupta (1960) and Morrow (1965),
and is not within the scope of this article.

DISCUSSION

The terminal inflorescence of Nannorrhops, of seeming complexity, is
shown to be built up by repeated and uniform ramifications of a simple

Fics. 28-34. Rachilla and flower clusters of Nannorrhops. Fic. 28. Details
of rachilla with several flower clusters. Fics. 29-32. Single (right-handed) flower
cluster and its dissection. Fic. 29. Entire flower cluster, the subtending bract
removed. Fic. 30. Bract, lowest flower and its attendant bracteole removed.
Fic. 31. Bract, 2 lowest flowers and their attendant bracteoles removed. Fic. 32
The same, but with uppermost bracteole removed to show continuing axis (or

cluster illustrated in Fics. 29-32. Fic. 33. In cross-section. Fic. 34. Theoretical
2-sided cincinnus in expanded state. This is regarded as the most probable
morphological interpretation of a flower cluster. The parts are drawn on elongated
axes to show their relative positions more clearly.

0 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Fics. 35-38. Inflorescence in Nannorrhops. Fics. 35-37. Developmental fea-
2 nflorescence

tures. Fic. 35. Young inflor October 4th, 1966. Fic. 36. Same, em-
ber Sth, 1 IG. 37 e, one week later; distal branches exposed indicating
their tendency to expand in a basipetal sequence. In Fics. n he e-

tative shoot to the left is the twin of the axis which is flowering, derived by
dichotomy of a single parent axis. Fic. = Details of transition of shoot 2, an
enlarged view of the axis shown in Fic. 4

kind. Progressive distal reduction in diameter of axes and size of bracts
produces a system which may be recognized as fundamental for many
flowering plants (Troll, 1964). In the simple descriptive terms of classical
morphology it would be called a panicle (Rispe of German authors).

1968] TOMLINSON & MOORE, NANNORRHOPS

ter
Leaves from in axis of shoot 1 arra nged “serially to show transition fro
foliage leaves to Bracts. Tubular base of each leaf has been split vertically if
release it from the axis. The dorsal ak Hs lowest (foliage) leaves is evident to
left. Detailed measurements in Fic. 1. . Shoot 1, an old specimen, before
dissection. Detailed measurements in Sn

Troll’s more modern typological terminology it is a “polytelic synflo-
rescence” (Troll, 1964; Weberling, 1964). The elaborate terminology
which this typological éutlook has developed could, no doubt, be applied
to the mammoth inflorescence of Nannorrhops but it would become un-
necessarily cumbersome. Our concept of this inflorescence, in which simple
constructional principles are recognized, does away with the need to apply
names to individual branches. The complexity of an inflorescence depends

32 JOURNAL OF THE ARNOLD ARBORETUM | VOL. 49

on the number of branch orders in which constructional principles are
repeated. It is of interest that the inflorescence in Nannorrhops is
constructed like that in many woody liliflorous plants of which Troll
(1962) illustrates numerous examples. In none of these, however, is the
branch pattern repeated as often as in Nannorrhops.

Using this simplified concept it is possible to make proper comparison
between different palm inflorescences. The lengthy and elaborate ter-
minology developed by earlier students of the palms is not only unneces-
sary but may even be misleading if it suggests to the uninitiated that
“spathe,” “bracteole” or “involucel” are distinct kinds of organs. ““Spathe”
itself is a redundant term. In our description of Nannorrhops we have
found that axis and bract are sufficient terms to describe the branch sys-
tem, rachilla is useful to identify an axis bearing flower clusters, bracteole
to identify bracts associated with (and possibly subtending) flowers,
prophyll to distinguish the first bract on each branch. No other terms
seem necessary.

In many other Coryphoid palms (e.g. Copernicia, Sabal, Washingtonia)
the lateral inflorescence is equivalent to one first order branch unit of
Nannorrhops as shown in Fic. 3. In all these palms it is evident that
exactly the same principles are involved in the construction of reproductive
branches. Sabal, for example, differs from Nannorrhops largely in that
vegetative growth of the axis continues into the reproductive phase. The
“lateral” inflorescence is strictly a partial inflorescence and was recognized
as such by many older taxonomists: (e.g. Blatter, 1926). This concept
tends to make redundant the idea that one type of inflorescence, “‘ter-
minal” or “lateral,” is necessarily more primitive than the other, as
discussed by Holttum (1955) and Corner (1966) for example. The
difference between them is physiological and possibly one of degree only.
Both types do exist together in one genus (Metroxylon) and might exist
together even in one species. An adaptive advantage in the pleonanthic
mode of flowering is suggested by its predominance.

From the elaborate Nannorrhops type the multifarious inflorescence
types of palms can be traced, although this article is not the place to carry
out the analysis which is required. Briefly, within the Palmae several
distinct trends may be noted. Reduction in the amount of branching
produces less elaborate forms, the ultimate expression being the un-
branched axis, an apparently “simple” spike. Recognition of this spike
(“spadix”) as a highly specialized type makes redundant any discussion
of affinities between the Palmae and Araceae. Reduction in the number
of bracts and their restriction to the base of the inflorescence is evident.
There may be few bracts, commonly two, or even only one. These basal
bracts are usually empty. The protective function of this reduced number
of bracts is facilitated by their enlargement and inflation so that one, OF
more, wholly encloses the inflorescence. The stage of development of ‘the
axis at which bracts cease to be protective varies widely in different palms.
The morphological equivalence of these specialized, inflated bracts
(“spathes”) with the sheathing base of a foliage leaf is not as evident as

1968 | TOMLINSON & MOORE, NANNORRHOPS 33

in Nannorrhops. Nevertheless, some vestiges of the foliage leaf may
persist even in the most specialized bracts, as in the woody terminal spine
of many Cocoid bracts which, by analogy with Nannorrhops, occupies the
position of the vestigial petiole and blade.

Adnation of branch to parent axis may be even more pronounced than
in Nannorrhops. The rachilla may be wholly adnate to a parent axis and
this, together with further condensation of ultimate flower clusters, may
produce highly specialized types whose fundamental morphology is diffi-
cult to decipher. Anatomical studies are beginning to reveal this com-
plexity. Aristeyera is a recently described geonomoid type which
exemplifies the degree of reduction and the approach needed in assessing
it (Moore, 1966; Uhl, 1966). Further specialization in palm inflorescences
involves changes to monoecism and dioecism.

Many morphological features of the palm inflorescence still await careful
observation and recording. Developmental and anatomical aspects have
scarcely been investigated. The subject is scope for a whole volume. It
is hoped that this preliminary study of Nannorrhops will be of value in
wider interpretations.

SUMMARY

Hapaxanthic flowering of the palm Nannorrhops ritchiana is described
from specimens in cultivation in South Florida. Erect vegetative axes
branch by an apparent dichotomy, one axis proceeding to inflorescence
with a gradual reduction of leaf size and elimination of petiole and blade.
Bracts are equivalent to the sheathing base of a foliage leaf with a vestigial
blade. Lateral branches from the axils of first-order bracts are subdivided
with up to 5 orders of branches leading to an elaborate paniculate
inflorescence. A simple one to one relationship between bract and branch
is maintained throughout the system. Lateral branches are basally
adnate to their parent axis. The distal decline in bract size can be traced
progressively to terminating branchlets (rachillae) where funnel-shaped
bracts only 2-3 mm. long subtend flower clusters. The arrangement of
bracteoles in each flower cluster suggests a condensed cincinnus. Nan-
norrhops has a basically simple branching pattern, despite its elaborate
ramification, and can be used in the interpretation of other palm in-
florescences.

ACKNOWLEDGEMENTS

We much appreciate the assistance of Miss Priscilla Fawcett who
prepared Ficures 3 to 34, the help with photography given by Dr. M
V. Parthasarathy, and the preparation of typescript by Miss Anne
Bellenger. We gratefully acknowledge the cooperation of staff at the U.S.
Plant Introduction Station, 13601 Old Cutler Road, Miami, Florida, in
providing material.

34 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

LITERATURE CITED

BLATTER, E. Se The palms of British India and Ceylon. Oxford University
ress, Lon

CorNeER, E. J. H 1966. The mee meas of palms. University of California
Press. Berkeley and Los Ange

Gupta, S.C. 1960. Organography ni pie anatomy of Nannorrhops ritchieana
H. Wendl. Jour. Res. Agra Univ. 9: —111

Hotttum, R. E. 1955. Growth-habits e aabcalyeanka — variations on a
theme. Phytomorphology 5: 399-413.

Moore, H. E., Jr. 1960. A new subfamily of palms — the Caryotoideae. Prin-
cipes 4: 102-117.

——. 1966. Aristeyera, a new genus of geonomoid palms. Jour. Arnold Arb.
47: 1-8.

& W. — 1965. A new species of Arenga from Borneo. Principes
9: 100-10.

Morrow, L. a 1965. Floral morphology and anatomy = certain Coryphoideae
(Palmae). Ph.D. Thesis. Cornell University. pp. 37

Rickett, H. W. 1944. The classification of aigk veces Bot. Rev. 10: 187-
231.

SCHOUTE, J. C. 1909. Uber die ‘See bei monokotylen coger? II. Die

era istelung von Hyphaene. Rec. Trav. Bot. Néerland. 6: 211-2

eet P. B. 1962. The leaf iad . Sins, its morphology and ae
biology. Jour. Arnold Arb. 43:

: Troe Stem structure in harder monocotyledons. Jn: M. H. Zim-

mermann, ed. The formation of wood in forest trees. pp. 65-86. Academic
Press, New York.

. 1967. Dichotomous branching in Allagoptera? Principes 11: 7

bi2.
& H. E. Moore. 1966. Dichotomous branching in palms? Principes 10:
agg

. H. ZIMMERMANN. 1966. Anatomy of Park palm Rhapis excelsa,
Ill, ae phase. Jour. Arnold Arb. 47: 301—
TROLL, W. 1962. Uber die “Prolificitat’” von Paspopeviiis comosum. Neue
Hefte Morphologie 4: 9-68.
. 1964. Die Inflorescenzen. 1. Gustav Fischer. Jena.
Unt, N. W. 1966. Morphology and anatomy of the inflorescence axis and
flowers of a new palm, Aristeyera spicata. Jour. Arnold Arb. 47: 9-22.
WEBERLING, F. 1965. Typology of inflorescences. Jour. Linn. Soc. (Bot.) 59:
2 1

FAIRCHILD TROPICAL GARDEN
Miamr, FLoriwa 33156

Casot FOUNDATION
Harvarp UNIVERSITY

and

BAILEY Hortortum
CORNELL UNIVERSITY
IrHaca, NEw York 14850

1968 | McDANIEL, HARPEROCALLIS 35

HARPEROCALLIS, A NEW GENUS OF THE
LILIACEAE FROM FLORIDA

SIDNEY MCDANIEL

AS A RESULT primarily of the botanical explorations of A. W. Chap-
man, the area around the Apalachicola River in western Florida is known
for its endemism and for its exceptional variety of plant species.
Approximately fourteen well-defined species are restricted to this im-
mediate area. Other species such as Baptisia megacarpa Chapm., B.
simplicifolia Croom, Gentiana pennelliana Fern., Hedeoma graveolens
Chapm., and H ypericune chapmanii P. Adams are endénic to a wider area
of the northeastern Gulf Coastal Plain. Within the Apalachicola River
area also occur a number of uncommon, though wide-ranging, species such
as Leitneria floridana Chapm., Croomia pauciflora (Nutt.) Torr., and
Schisandra glabra (Brickell) Rehd. The variety of habitat within the area
is exceptional and ranges from alluvial river bottom to sandy longleaf pine
forests or to Sarracenia bogs. The diversity within the Apalachicola River
area both of species and habitat and the known endemism within the area
help to explain a previously undescribed, apparently endemic genus of the
Liliaceae which has been discovered in Franklin County, Florida.

The relationships of Harperocallis, the remarkable new genus described
below, seem to be with the tribe Tofieldieae Kunth of the Liliaceae. This
tribe as circumscribed by Krause (1930) consists of Tofieldia Hudson,
Pleea Michx., Narthecium Hudson, and Nietneria Klotzsch & Schomb.
Bentham and Hooker (1883) and Hutchinson (1959) delimited the
Narthecieae Benth. & Hook. (synonymous with the Tofieldieae in the
sense here used) to include the genera listed above and several others
morphologically more divergent. Harperocallis is markedly distinct from
all other genera of this tribe and may be distinguished readily by its
solitary flower, tuberculate ovary, and calyculus of three (or four)
separate bracts.

The members of the Tofieldieae, including Harperocallis, are all rhizo-
matous perennials, with linear, mostly basal equitant leaves and terminal
inflorescences on simple stems. The leaves of Harperocallis are almost
identical to those of Narthecium and certain species of Tofieldia. The
simple stem has a few reduced bractlike leaves above and one or more
sheathing leaves near the base. A calyculus of three or four bracts is
immediately beneath the solitary terminal flower. Flowers of certain
species of Tofieldia may have a calyculus, but the segments are united.
In Pleea each individual flower is subtended by a large sheathing bract.
Narthecium and Nietneria have single bracts subtending the individual
flowers along the axis of the inflorescence. The stamens are six in all

36 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Tofieldieae, except Pleea where they may be six to twelve but most com-
monly are nine. Except for the pilose filaments of Narthecium the fila-
ments are glabrous. In the Tofieldieae the ovary is trilocular and often
three lobed. The styles are distinct and suberect or slightly recurved in
Pleea, Tofieldia, and Harperocallis. In Narthecium and Nietneria the
styles are united and erect. The capsule in Harperocallis as in Narthecium
and Nietneria is loculicidal. Pleea and Tofieldia have septicidal capsules.

The unique morphological characteristics of Harperocallis and the
various morphological features held in common with different genera of
the Tofieldieae suggest no particularly close relationship with any other
genus. The presence of the calyculus and other morphological similarities
suggest closest affinity to be with Tofieldia.

At the type locality Harperocallis is most frequent in the vicinity of
isolated individuals of Cliftonia monophylla (Lam.) Britt. ex Sarg. and
Myrica inodora Bartr. in areas slightly drier than the adjacent open bog.
Within the bog Harperocallis is less frequent and is associated most
prominently with Sarracenia flava L., S. psittacina Michx., and Pleea
tenuifolia Michx. On the edge of the bog Harperocallis was noted grow-
ing in the detritus of an old pine log beneath Cliftonia. The open bog
abruptly changes to the west to a dense thicket largely consisting of
Cliftonia and an occasional individual of Pinus serotina. The whole area
is surrounded by sandy, occasionally burned longleaf pine woods. Ini-
tially a few individuals of Harperocallis flava were observed in a roadside
ditch slightly to the south of the type locality.

Although as earlier mentioned the Apalachicola River region is noted
for its endemic species of plants, there are no previously described endemic
genera. I have examined numerous areas from the vicinity of Tallahassee
to extreme western Florida seemingly quite similar to the type locality
without finding any sign of Harperocallis. Probably, however, the genus
occurs elsewhere, for other endemics of the Apalachicola River area, such
“a and Scutellaria floridana Chapm., occut
any Tegion. | Harperocallis may eventually prove to

ave a similar type of distribution.

It is appropriate to associate with this genus of the Lower South the
ng . pm Ne — Merete who greatly contributed
the generic name emphasizes the arate pence tenia Seer :

iveness of the flower. The specific
epithet refers to the color of the perianth segments.

Harperocallis McDaniel, gen. nov.

Lit TACEAE Mirt antrinamear ’T

a

Herba perennis. Rhizoma gracile nodosum, saepe terminante in planta
emergenti. Folia equitantia, lineares, rigidula, striata, quam flos multo
breviora. Inflorescentia constans ex flore solitario ot erecto. Bracteae
subflore 3 vel 4, persistentes, plus minusve aequales, contiguae, deltoideae
apice acutae. Perianthium persistens, flavum interne et patentia per

1968]

McDANIEL, HARPEROCALLIS

Fic. 1. Harperocallis flava, X 7.

38 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

anthesin; segmenta 6, distincta, oblanceolata, acuta, hae subtentae a brac-
teae intimis segmentis longiora. Stamina 6, hypogyna, quam perianthium
breviora; antherae lineari-lanceolatae, basifixae, loculis introrse dehiscen-
tibus, filamentae glabrae, sensim latae basin versus. Ovarium sessile,
trilobum, triloculare, ellipsoideum, valde tuberculatum, ovula in loculis
numerosa; styli 3, suberecti, breves; stigmata capitata. Capsula vix maior
quam ovarium per anthesin, triloba, ellipsoidea, loculicide dehiscens.
Semina parva, anguste fusiformia, recta vel curvata. SPECIES TYPICUS:
Harperocallis flava.

Harperocallis flava McDaniel, sp. nov.

Herba glabra, prope basin fibrillosa. Caulis simplex, 22-55 cm. longus.
Folia 0.2-0.3 cm. lata X 5—21.5 cm. longa. Segmenta perianthii 10-12
nervata, hae subtentae a bracteis 12-15 mm. longa, segmenta interiora
9-12 mm. longa. Stamina 8-9 mm. longa. Ovarium 7—8.5 mm. longum.
Capsula 8-9 mm. longa. Semina 2-3 mm. longa. Hototypus: McDaniel
6205 (A)

Perennial herb, glabrous throughout, basally fibrillose. Rhizome slender,
nodose with scarious scale leaves, often ending in an emergent plant.
Leaves 0.2-0.3 cm. wide * 5—21.5 cm. long at maturity, equitant, linear,
stiff, and striate, much exceeded by the scape; sheath scarious margined.
Stem simple, 22-55 cm. long, with 3-5 much reduced leaves. Bracts 2—3
mm. long, persistent, usually 3 in number, contiguous, deltoid, forming a
calyculus beneath the flower. Perianth persistent, yellow above and
greenish beneath at anthesis, spreading, after anthesis becoming erect,
revolute, externally green and internally purple margined; perianth seg-
ments 6, 10-12 nerved, oblanceolate, acute, those subtended by the bracts
12-15 mm. long, the inner series 9-12 mm. long. Stamens 6, 8-9 mm.
long, hypogynous; anthers ca. 4 mm. long, linear-lanceolate, basifixed,
introrsely longitudinally dehiscent, filaments glabrous, somewhat broad-
ened toward the base. Ovary 7-8.5 mm. long, sessile, 3-lobed, 3-locular,
ellipsoid, strongly tuberculate, ovules numerous; styles 3, suberect, ca.
1 mm. long, stigmas capitate. Capsule 8-9 mm. long, ellipsoid, loculicidal,
tuberculate, almost hidden by the perianth. Seeds 2-3 mm. long, pale
yellow, narrowly fusiform, straight or curved, often twisted.

SPECIMENS EXAMINED: Florida. FRANKLIN Co.: 2.2 mi. south of Sumatra,
open bog surrounded by Cliftonia-Pinus serotina, May 11, 1965, McDaniel 6205
(holotype, A; isotypes, BM, DUKE, FLAS, FSU

, GA, MO, NCU, NY, PH, SMU, US),
July 21, 1965, McDaniel 6542 (a, Psu); 2.5 mi. south of Sumatra, roadside near
pine woods, May 4, 1965, McDaniel 6127 (FSU).

M any persons have aided in the preparation of this paper. I am
especially grateful to Robert K. Godfrey for initial advice and for reading
the manuscript, to Carroll E. Wood, Jr., for confirming the uniqueness of
Harperocallis and for suggestions as to its affinity, and to Barbara N.
Culbertson for her careful preparation of the illustrations.

1968 | McDANIEL, HARPEROCALLIS

Fic. 2. Harperocallis flava: a, flower from below and above, X 214; b, fruit, X
21%; ¢, anthers, showing dehiscence, « 6; d, ovary and calyculus, « 6; e, seeds,
* 6; f, germinating seed with corona of root hairs at base of hypocotyl, x 12.

40 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

LITERATURE CITED

BentHaM, G., & J. D. Hooker. 1883. Genera Plantarum. Vol. 3. London.
{Narthecieae, 825-829. |

Hutcuinson, J. 1959. The Families of Flowering Plants. Vol. Il. Monocoty-
edons. Ed. 2. London. [Liliaceae, 591-612; Narthecieae, 595, 596.]

Krause, K. 1930. Liliaceae. Jn: A. ENGLER, Die Natiirlichen Pflanzenfamilien.
Aufl. 2. 15a: 227-386. Leipzig. [Tofieldieae, 254-256. ]

DEPARTMENT OF BOTANY

MissIssipP1 STATE UNIVERSITY
STATE CoLLEcE, Mississtppr 39762

1968] STONE & FREEMAN, CYTOTAXONOMY OF ILLICIUM 41

CYTOTAXONOMY OF ILLICIUM FLORIDANUM
AND I. PARVIFLORUM (ILLICIACEAE)

Dona tp E. STONE AND JupITH L. FREEMAN 1

RECENT CHROMOSOME CouNTS of the Ranales by Raven and Kyhos
(1965) provide new insight into evolutionary patterns of the most primi-
tive members of extant angiosperms. They concluded that the basic
chromosome number for the primitive angiosperms is probably «=7. It
is argued that numbers such as n=6, 8 (Aristolochiaceae), n=8, 9 (An-
nonaceae), m=10 (Eupomatiaceae), m=11 (Calycanthaceae, Saurura-
ceae), #=12 (Lauraceae), m= 13 (Canellaceae, Drimys sect. TASMANNIA),
etc. are aneuploid derivatives of the base number 7 and its multiple, m= 14.
Stebbins (1958, 1966) has recently reiterated an alternative hypothesis
which seeks to explain the higher numbers such as n= 12 and n= 13 as the
result of “direct polyploidy,” rather than a doubling of 7 followed by
aneuploid reduction. He pictures aneuploid changes in chromosome num-
ber from a basic x=6 or x=7 to have gone both upward (x=9) and
downward (x=5) at an early stage of angiosperm evolution. This period
was followed by the build up of higher basic numbers in woody members
through polyploidy. Once the polyploid levels were reached Stebbins
implied that further evolutionary diversification was relegated to processes
of speciation and generic differentiation at the homoploid level. While
such a hypothesis is highly plausible and, in fact, is generally accepted in
the derivation of »=19 in the Magnoliaceae from unknown m=7 and
n = 12 (found in the related Degeneriaceae and Himantandraceae) ances-
tors (Stebbins, 1950; Darlington, 1956; Raven and Kyhos, 1965); evi-
dence does exist in the Ranales for aneuploid changes at the polyploid
level. The Piperaceae, for example, have n=11, 12, 14, and 16, numbers
which are surely indicative of aneuploid derivation. Stebbins (1966, Table
2) overlooked another example cited by Raven and Kyhos (1965; Stone,
1965) in which /ilicium floridanum Ellis with n= 13 is a probable deriva-
tive of the basic n= 14 common to other members of the Illiciaceae and
the related Schisandraceae. It is the purpose of this paper to examine the
karyotypes of two of the four New World species of Jllicium and to
document the aneuploid reduction from n= 14 to n=

MATERIALS AND METHODS

The collections and chromosome numbers of Jilicium floridanum Ellis
and J. parviflorum Michx. ex Vent. are presented in TABLE 1. The
populations that were sampled do not span the range of each species but
presumably are representative. J. floridanum ranges from northwestern

1 Work done in conjunction with research supported by NSF grants GB- aap
GB-5233X, GY-2484, and the Duke University Council on Research. We wish to
thank Dr. Carroll E. Wood, Jr. for his helpful comments on the manuscript.

42 JOURNAL OF THE ARNOLD ARBORETUM [vor. 49

TABLE 1. Collections and chromosome numbers of
Illicium floridanum and I. parviflorum

SPECIES AND COLLECTION CHROMOSOME
NUMBER NUMBE COLLECTION DATA

n 2n

Illicium floridanum Ellis (sect. ILLictum)

Stone 1357 13 Washington Parish, near War-
nerton, La. (NO)
Stone 1522 26 Hancock Co., near Logtown,

Miss. (DUKE)
Stone 1819 26 Alachua Co., cultivated near
Gainesville, Fla. by A. M.
Laessle. (DUKE)
Illicium parviflorum Michx. ex Vent. (sect. CyMBOSTEMON )
Stone 1422 14 28 Alachua Co., cultivated near
Gainesville, Fla., by A. M.
Laessle. Cuttings originally
obtained from Norwalk, ne.
corner of Ocala National For-
est. (DUKE
Stone 1820 14 28 Orange Co., Chapel Hill, N.C.,
cultivated on Univ. of North
Carolina campus. (DUKE)
Stone 1821 28 Orange Co., Chapel Hill, N.C.,
cultivated on Univ. of North
Carolina campus. (DUKE)

Florida to eastern Louisiana and northward to central Alabama. /. par-
viflorum is restricted to a few localities at the headwaters of the St. Johns
River in northeastern Florida and is quite often marketed by Florida
nurserymen as J, anisatum (pers. comm., A. M. Laessle).

Flower buds suitable for studies of meiosis were fixed in 3 (absolute
ethanol) : 1 (glacial acetic acid) and stored in 70% ethanol until prepared
for examination by the standard acetocarmine squash technique. Kary0-
type analyses of mitotic chromosomes were conducted on the root tips of
young seedlings or cuttings stimulated to produce callus and adventitious
roots with hormone treatment. Most satisfactory preparations were ob-
tained by treating root tips in a 0.1% colchicine solution for 3 to 4 hours
before fixation.

Chromosome squashes deemed satisfactory for analysis were photo-
graphed with a Leitz Ortholux microscope (x 90 objective, N.A. 1.32)
and Kodak Contrast Process Ortho sheet film. Prints of the negatives
were standardized at 3000 for measurement with a millimeter rule.
Measurements were made of the total chromosome length and length of
each arm. The measurements of homologous chromosomes were averaged

1968] STONE & FREEMAN, CYTOTAXONOMY OF ILLICIUM 43

and expressed in percent as a function of the total chromosome length of
the genome (i.e. mean percent length, Martin and Hayman, 1965). This
procedure was an attempt to reduce variation in chromosome length
introduced by variable cell-sizes and squash preparations, while at the
same time expressing the results from different cells on a standard scale.

RESULTS

Karyotype analyses. The data from untreated and colchicine treated
cells were processed separately. The differences due to treatment were
negligible, however, and the samples were subsequently pooled. While
the basic proportions of the karyotype were unaffected by treatment, the
highly contracted colchicine-treated chromosomes were less suitable than
untreated chromosomes for analyses of secondary constrictions. The
results presented in TABLE 2 are based on ten cells each from seedlings
and cuttings of Jllicium parviflorum (Stone 1422, 1820) and a combined
total of eight cells from seedlings and cuttings of J. floridanum (Stone
1522, 1819)

The data from the native Florida (Stone 1422) and cultivated North
Carolina (Stone 1820) plants of Jllicium parviflorum are presented sep-
arately to show the remarkable similarity in karyotypes (Fic. 1). Since
these samples were independently analyzed, we consider the good agree-
ment in results to be ample vindication of the techniques employed.

Illicium floridanum has 2n= 26 (Fic. 3) and a karyotype consisting of
1 v-shaped, 10 J-shaped, and 2 rod chromosomes. J. parviflorum has
2n=28 (Fics. 7, 8) and a karyotype which includes 10 j-shaped and 4
rod chromosomes. On the basis of total size, relative arm lengths, and
secondary constrictions, it has been possible to match 9 3 (#2-10) and
2 rod chromosomes (#11, 12) of the J. floridanum and I. parviflorum
complements (Fic. 1). Chromosomes 1 and 15 of J. floridanum and
chromosomes 1, 13, and 14 of J. parviflorum do not have identical
equivalents. Since J. floridanum (sect. Ittictum) and /. parviflorum (sect.
CYMBOSTEMON) are representative of two clearly demarcated sections of
the genus, direct derivation of one from the other is not possible. However,
if we assume the 14-chromosome karyotype of J. parviflorum is probably
not too dissimilar from other 14-chromosome species, including /. anisatum
of sect. Irticrum, then the simplest explanation is that three parviflorum-
like chromosomes have undergone reciprocal translocations of unequal
chromosome segments to be repatterned into the two of /. floridanum.
The basic scheme proposed by Darlington (1937) shows how the entire
euchromatic arm of a rod-shaped chromosome could be transferred to the
heterochromatic arm of a second rod chromosome. This process, some-
times referred to as centric fusion, followed by loss of one heterochromatic
centromere region, would result in gametes with one less chromosome but
without any loss of genetic material. Of course, proof for this type of
rearrangement must necessarily come from meiotic studies of F; species
hybrids, as Togby (1943) has Sopaerien in Crepis (n = 4 ton = 3)
and Kyhos (1965) in Chaenactis (n = 6 ton = 5).

JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

44

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STONE & FREEMAN, CYTOTAXONOMY OF ILLICIUM 45

1968]

o

floridanum

parviflorum
ee

(Filia) (NC)

a

—

RKRI LRG
KU MQ

RRSSOE
(KG

RRQ
MQ

LLG RRLNE
KEM

OOOOIOS
KKK

AOR RRL
CMW

KKK

RRLKLLLL LL m
KKK

SERRE D>
KKK

ROO MED
Kd QQ
SORRY s+
KCK

os
SOK OOO RY OFD
CCK MCU

RRO KM EN
WMC ME

C#K{’™XK{#

KOK

BSS RRS
KKH OM

RRS SIR SRR
CM XMM CE

SESGEEE GEER

KING IE

KG UAE

KKK), HMKKK KG.
mo

™N
—

1

—

ao

Idiogram of karyo-

1 translocations of unequal chromosomal segments.

gy of Illicium chromosomes.

iproca

.

Fics. 1 and 2. Morpholo
mosomes by rec

type of J.
‘0

oretica

like chro

46 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Fic. 3. Ilictum floridanum. Mitotic metaphase, 2n = 26, Stone 1522, X 1780.

Neither hybrids between these shrubs nor studies of pairing configura-
tions in meiosis have been attempted. However, the simplest explanation
for the derivation of »=13 from n=14 would involve centric fusion.
Fusion of rod chromosomes 13 and 14 (Fic. 2) might have produced
chromosome 15 of J. floridanum. Chromosome 1 of J. floridanum (Fic.
1) is very similar to chromosome 1 of J. parviflorum. Both chromosomes
are the longest of their respective genomes and both have secondary
constrictions located in approximately the same region in each arm. The

1968] STONE & FREEMAN, CYTOTAXONOMY OF ILLICIUM 47

J-shaped morphology of /. parviflorum could have been transformed into
the v-shape of J. floridanum by adding segments to the ends of the arms
(Fic. 2).

Meiotic analyses. Acetocarmine squashes of pollen mother cells were
prepared. Meiosis was found to be normal in both species with a base
number of m=13 in Illicium floridanum (Fics. 4-6) and n=14 in J,
parviflorum (Fics. 9-10). An analysis of 19 cells of J. floridanum
revealed an average chiasma-frequency of 1.31 per bivalent (TABLE 3).

TABLE 3. Chiasma-frequency at diakinesis in Illicium floridanum *

TYPES OF O. OF O, OF

CHIASMATA CHROMOSOMES CHIASMATA % CHIASMA-TYPES
Zero 16 0 6.5
One terminal 69 69 27.9
Two terminals 35 70 14.2
One interstitial 73 73 29.6
Two interstitials 25 50 10.1
One terminal as 50 10.1

— one interstitial
Two terminals 4 1 1.6

— one interstitial
TorTaLs 247 324 100.0

Chiasma-frequency/bivalent = 324/247 = 1.31

* Based on 19 cells

SUMMARY AND CONCLUSIONS

Chromosome counts on the related Schisandraceae (Whitaker, 1933;
Stone, 1965) and two of the three members of the Illiciaceae (Whitaker,
1933), suggest that 14 is the base number for the alliance. Viewed in
this light, »=13 in Jllicium floridanum clearly seems to be the prod-
uct of aneuploid reduction. Though we have made direct comparisons
between the karyotypes of J. floridanum (sect. Ittictum) and J. parvi-
florum (sect. CymBostEMON), it should be re-emphasized that these
species are representatives of the two sections of the family. It would be
most interesting to examine the 14-chromosome karyotype of /. anisatum
(Whitaker, 1933), for this species is one of the twelve Smith placed in
section ILLictum (formerly sect. BApIANA) with /. floridanum. The cytology
of section CyYMBOSTEMON is equally unknown. The n= 14 count of J. par-
viflorum is the only report on the 29 species recognized by Smith (1947).
While it is quite likely that the 14-chromosome species of both sections
have similar karyotypes, it will be necessary to analyze close relatives of
I. floridanum and meiotic pairing configurations formed in fs 1s xX
n = 14 hybrids before the exact nature of aneuploid reduction is estab-
lished.

Centric fusion (Fic. 2) of rod chromosomes 13 and 14 of J. parviflorum

\

JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

we ®
sa »& 3
4 ©

Fics. 4-6. Illicium floridanum. Meiosis in rare mother cells, m = 13, Stone
1357, X 1000. 4. Diakinesis. 5. Metaphase I. Anaphase I.

1968] STONE & FREEMAN, CYTOTAXONOMY OF ILLICIUM 49

FI 7 and 8 icium parviflorum. Mitotic metaphase, 2” = 28, Nae
. North C soli paedlatiiee Stone 1820. 8. Florida saoubsine: Stone 142

50 JOURNAL OF THE ARNOLD ARBORETUM [VvoL. 49

. 9 and 10. Illictum parviflorum. Meiosis in pollen mother cells, 7 = 14,
ins 820, * 1500. 9. Diakinesis. 10. Anaphase I.

could account for number 15 of J. floridanum (Fic. 1). In addition, il
the residuum of number 13 was transferred to the terminal ends of the
arms of chromosome 1 (Fic. 2), a v-shaped chromosome similar to
chromosome 1 of J. floridanum would result.

1968] STONE & FREEMAN, CYTOTAXONOMY OF ILLICIUM 51

We have no information on the significance of the reduction from
n=14 to n=13. Whitaker (1933) originally reported n= 14 for Jllicium
anisatum L. (as I. religiosum) and J. floridanum. As we have shown,
his report for J. floridanum is probably incorrect. Unfortunately, as
Wood (1958) has noted, Whitaker’s counts were not documented with
voucher specimens and there is no way to tell if he miscounted or mis-
identified the material. It does open the question, however, of whether
I. anisatum of the same section might not also have n=13. If not, then
n= 13 may be exclusive to the American species of section ILLictum. For
as Smith (1947) commented, “The SS [Z. floridanum| is clearly
separable from all Old World members of § Badiana [ = § Ittictum] on
the basis of its comparatively long eilicsts, numerous stamens, and
brightly colored perianth segments. Its only close relative is the following
new species |[/. mexicanum].”

LITERATURE CITED

BatLey, I. W., & C. G. Nast. 1948. Morphology and relationships of oe
Schisandra and Kadsura. I. Stem and leaf. Jour. Arnold Arb. 29: 77-89

Daruincton, C. D. 1937. Recent advances in cytology. 2 ed. biaaten.
Philadelphia.

. 1956, ea heen botany. The University Press, Aberdeen

KyHos, D. W. 1965. The independent aneuploid origin of two species of
Chaenactis (Compositae) from a common ancestor. Evolution 19: 26-43.

Martin, P. G., & D. L. Hayman. 1965. A snespragt Ae sips for comparing
the karyotypes of related species. Evolution 19: 15

RAvEN, P. H., & D. W. Kynos. 1965. New evidence oe the original
basic chromosome number of angiosperms. Evolution 19: 244-248

SMITH, ss C. 1947. The families Illiciaceae and Schisandraceae. Sargentia 7:
1-—

Semen G. L., Jr. 1950. Variation and evolution in plants. Columbia Uni-
sealer | Press, New Yor
958. On the hybrid origin of angiosperms. Evolution 12: 267-270.
——-, iene Chromosomal variation and evolution. Science 152: 1463-1469.
Stone, D. E. 1965. Jn: Documented chromosome numbers of plants. Madrofio
18:

125.

5 a press. Cytological and morphological notes on the southeastern
endemic, Schisandra glabra (Schisandraceae). Jour. Elisha Mitchell Soc.
Tocsy, H. A. 1943. A cytological study of Crepis fuliginosa, C. neglecta, and

their F, hybrid, and its bearing on the mechanism of phylogenetic reduction

Wuirtaker, T. W. 1933. Chromosome number and relationship in the Mag-
noliales. Jour. Arnold Arb. 14: 376-385.
Woop, C. E., Jr. 1958. The genera of the woody Ranales in the southeastern
United States. Jour. Arnold Arb. 39: 315-318.
DEPARTMENT OF BOTANY
Duxe UNIVERSITY
DurHam, NortH CAROLINA 27706

52 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

FURTHER MORPHOLOGICAL STUDIES ON ANASTOMOSES
IN THE DICHOTOMOUS VENATION OF CIRCAEASTER

ADRIANCE S. FOSTER

CONTINUED RESEARCH ON DICHOTOMOUS LEAF VENATION in specimens of
Circaeaster agrestis Maxim. grown at the University of California Botan-
ical Garden, Berkeley, during 1965 and 1966 revealed that the three kinds
of anastomoses described in my last paper (Foster 1966) do not include
the full range of variation in types of vein unions which are developed in
this genus. The main purpose of the present article, therefore, is to de-
scribe the additional forms of anastomoses and their associated vein
approximations which have been found, and to compare their morphology
with the types previously considered.

A further objective of this paper is to present a brief account of the
early phases of development of the seedling and the complex organography
of mature flowering and fruiting specimens. This phase of my study is
illustrated by photographs of living plants (Fics. 1-3) which it is believed
will convey a realistic idea of the general organization and development
of the leaf-rosette and the condensed fascicles of flowers characteristic of
this remarkable little dicotyledonous herb.

MATERIALS AND METHODS

The cultivated plants which provided the material for this study were
grown during 1965 and 1966 from the collection of fruits given to me
through the kindness of Dr. M. Anantaswamy Rau, Regional Botanist of
the Botanical Survey of India. He had collected these fruits on October
22, 1963, from the colonies of Circaeaster originally discovered in 1962 by
Bhattacharyya (1964) “growing gregariously on shady and flat mossy
rocks under the shade of Viburnum bushes” near Gangharea, North Garh-
wal, India, “at an altitude of nearly 3300 meters.”

Fruits were stored at a temperature of approximately 40°F until needed
and under such conditions retained for more than three years an ability to
germinate. The seedlings were raised in pots of soil in a growth chamber
illuminated for 16 hours a day and maintained at a constant temperature
of 42° F. Some variation in the time required for germination was note
over the successive years but in general a period of 3 to 4 weeks elapses
before the conspicuously elongated hypocotyl elevates the cotyledons above
the surface of the ground. Young plants were kept in the growth chamber
for about a month during which time a series of leaves and. in some plants,
the first fascicles of flowers were formed. The plants ware then taken to
the Botanical Garden, transplanted into small individual pots and placed

1968 | FOSTER, CIRCAEASTER

GS. 1 and 2. General STEBROETADHY of young cultivated specimens of Cir-
cae aster sakes ton amen from above to show the cotyledons and primary Base ink
1, seedling, cae ons sid six expanded primary leaves.

vO inea 7: older
stage The. smal structures in the center of the rosette are young involute
leaves. Both fi 4

either in a lath house or a cool greenhouse. Mature fruits collected from
potted plants during mid- or late summer of 1966 have proved viable and
some of them this spring (1967) are producing a new crop of seedlings in

54 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

the growth chamber. Seedlings are also now developing in the open from
fruits which had fallen from a group of plants which were grown last year
in an especially prepared bed in the rhododendron dell in the Botanical
Garden. It is hoped that from these plants a “permanent” colony may
become established, comparable to the colonies maintained for a number of
years at the Botanical Gardens at Upsala and Edinburgh (Foster 1963,
p. 302).

When fully mature, the leaf-rosettes of a large number of plants were
either preserved directly in 70 percent alcohol or in some cases were first
killed and fixed in formalin-acetic acid-alcohol before storage in alcohol.
All of the foliar organs of each plant (including the innermost small bracts
of the flower-fascicles) were subsequently removed by means of small scis-
sors, carefully examined for vein approximations and anastomoses, and
the data recorded in tabular form. The leaves selected for venation studies
were Cleared in 5 percent NaOH followed by concentrated chloral hydrate,
dehydrated in the usual alcohol series, stained in a 1 percent solution of
ee in equal parts of absolute alcohol-xylene and mounted in “Pic-
colyte.”

I am extremely grateful to Mrs. Irene Baker, who has undertaken each
year the planting of the fruits and has kept careful records of the begin-
ning and the extent of the period of germination and early development of
the seedlings. The line drawings in this paper represent direct tracings of
photomicrographs and were prepared by Mrs. Emily R. Reid. I wish also
to thank my wife for her assistance in proof-reading the manuscript.

ORGANOGRAPHY OF SEEDLING AND ADULT PLANT

The present description of the germination and general organographic
development of Circaeaster is intended to supplement my previous account
which Was based entirely upon the study of a very limited number of
herbarium specimens (Foster 1963, pp. 304-306).

During the early phases of germination, the curved, elongating hypo-
cotyl, bearing the pair of closely appressed cotyledons at its apex, emerges
from the soil. As a result of differential elongation, the hypocotyl soon
maser erect and the two linear cotyledons reflex and assume a horizontal
ee Rep eo RiaA gE fear green and persist throughout the life

Approximately a week after the cotyledons have reflexed, a series of
small, cuneate primary leaves begins to expand (Fics. 1 and 2). The form
and relatively simple dichotomous venation patterns of these earliest

pain wei already been described (Foster 1963, pp. 308, 309). The
ernatio : : , Pp. ). Th
idea of the primary leaves, like that of the adult type of leaf, is

As elongation of the h

- ypocotyl continues, the terminal rosette becomes

ge and morphologically complex because of i
: the development of addi-
iy crowded adult leaves followed by the rapid bcs casicn of bracts and

eir axillary fascicles of small flowers (Fic. 3). Since there is no clear

1968 | FOSTER, CIRCAEASTER 55

A maturing "ee ated specimen of Circaeaster showing the rosette of
dicho pl veined leaves, the central fascicles of flowers and, at center left,
a maturing fruit with shi aaeaela ie uncinate hairs,

demarcation either in form or venation between foliage leaves and well
developed bracts, the latter were included in the total number of leaves in
each of the 80 plants which were dissected and analysed in the present
study (TABLE 1), Using this admittedly imprecise method of tabulation,
it is nevertheless apparent that the number of leaves varies widely from
plant to plant, ranging from 11 in very small precociously flowering indi-
viduals to as high as 180 in a few very robust plants. The “average leaf
number” for all plants studied is approximately 6

In an earlier paper (Foster 1963, p. 304) the question was raised
whether the compact rosette of leaves and tiny flowers of mature plants is
borne in an upright or prostrate position in nature. Cultivated plants,
rowing under the best horticultural conditions which could be provided,

56 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

throw some light on this question. During the maturing stages of these
plants, many of them became “top heavy” and there was a marked tenden-
cy for the slender hypocotyl to be bent towards the ground. In fact it was
necessary, in most cases, to use a miniature “Carnation-type” plant-sup-
porter to hold each plant in a more or less erect position. In maturing
plants in nature it seems possible that the rosette also may ultimately
become prostrate but this conjecture needs to be tested by extensive field
observations.

The full cycle of development in Circaeaster is relatively short and fol-
lows the general pattern of many dicotyledonous herbs. As accurately as
could be determined, the life span of cultivated plants is about five months
from seedling to complete maturity. This cycle is somewhat longer than
that of the plants growing in their native habitat in Gangharea, North
Garhwal, India, according to notes kindly given to me in a recent letter by
Dr. M. A. Rau. He writes: “Regarding the growth cycle in nature, I do
not have precise information about the time of appearance of seedlings.
At these heights snow remains till the first week of June and it is, therefore,
likely that germination takes place sometime during June. They are in
full bloom by the middle of August and by the end of September the
plants will have fruited. The plants are spent out by October and when I
collected the ripe fruits on October 22 (1963), the fruits were all on the
ground and the plants had collapsed. There may be some little variation
in the dates given above from year to year depending on the environmental
factors but in general you may take the above as the schedule in nature.”

VEIN APPROXIMATIONS AND ANASTOMOSES

Résumé of Previous Studies. Before presenting the results of the
present investigation, it seems highly desirable to review briefly (1) the
classification i.e. “typology” of the anastomoses proposed in my recent
paper (Foster 1966) and (2) to re-emphasize the morphological signifi-
cance of vein approximations i.e. transitions between open dichotomous
venation and true anastomoses. The latter represent the union between two
xylem strands while in a typical vein approximation the tracheary elements
of the adjacent branches of a pair of very closely spaced veins are separated
by parenchyma (see Foster 1966, Figs. 26, 27 for histological details).
The interrelationship between open dichotomous venation, vein approx-
mations and Types I to III anastomoses are represented in Fic. 4. It
should be emphasized that the very symmetrical vein approximation shown
in this figure is not hypothetical but is based upon actual patterns observed
in a number of leaves (Foster 1966, Figs. 3,4, 6, 8). The distally “open”
areole created by this kind of vein approximation is strikingly similar in
form to the “closed” areoles formed by both Type I and Type II anasto-
moses, These two kinds of vein unions differ from each other in that xylem
fusion in Type I is “transient,” i.e. the extent of contact is relatively short
while in Type II, the two veins completely merge to form a single, un-
branched excurrent vein. Both of these forms of anastomoses are funda-

1968 | FOSTER, CIRCAEASTER 57

AM

Type I anastomosis

Yt

Open Type Il anastomosis

dichotomous Vein

venation approximation
Type III anastomosis

. Diagrams representing the morphological relationships between open
dichotomous venation, symmetrical vein approximation and Types I to III
anastomoses. Further explanation in text.

mentally similar however in that the union occurs between the adjacent
branches of two vein-dichotomies (Foster 1966, Figs. 1, 2).

Type III anastomoses, as shown in Fic. 4, are less easily interpreted
because the approximated branches of two vein-dichotomies remain discrete
and are only interconnected by a very short obliquely oriented “commissural

58 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

vein.” The xylem of the latter may consist of one or at most of only a few
tracheary elements (see Foster 1966, Figs. 28, 33a, b). Whether the “com-
missural vein” should be regarded (1) as a much “reduced” branch of a
vein-dichotomy or (2) as a “new formation” is problematical, although my
previous study indicates a reasonable probability in favor of the first
alternative. As I have pointed out, comparable difficulties arise in the
interpretation of the short “commissural veins” in the leaves of certain
ferns (Goebel 1922, pp. 19, 20) and in the petals of Ranunculus (Arnott &
Tucker 1963, Fig. 16, type C’”).

Although multiple anastomoses (i.e. two or more vein unions in one leaf)
were not observed in any phase of my investigations on Circaeaster, sev-
eral leaves were encountered each of which had formed two vein-approx-
imations. A striking illustration is represented in Fic. 5 which shows a
small bract (13 mm. long X 3 mm. wide) with two symmetrical vein
approximations, each forming a very elongate distally open areole. A
venation pattern of this type suggests the strong possibility of the occa-
sional formation of multiple anastomoses in Circaeaster but thus far no
examples have been discovered either in leaves from a wide range of
herbarium specimens or in those of the numerous cultivated plants which
I have studied.

New Observations. Taste 1 provides further evidence for the con-
clusion previously reached that anastomoses are very uncommon in the
leaves of Circaeaster (Foster 1966, p. 592). In more than 5000 leaves
critically examined in the present survey, only 54 or about 1 percent of
them had formed an anastomosis. Vein approximations were slightly more
frequent and occurred in 2 percent of all the leaves studied.

Despite the sporadic distribution of anastomoses in the leaf sample @s @
whole, the percentage of anastomoses in the leaves of certain individual
plants was comparatively high and ranged from 1.8 to 6.6 percent. As shown
in TaBLe 2, the 12 plants in which this higher frequency of anastomoses
was observed, were distinct from all the other plants because of the pres-
ence of solitary vein-unions in two or more leaves of the same individual.
It seems reasonable to postulate that the number of anastomoses per plant
may be under some type of genetic “control” but this is an idea which
requires confirmation by appropriate breeding and experimental techniques.

From a morphological viewpoint, the majority of the vein unions an-
alysed could readily be classified under either Type I or II anastomoses
(Fic. 4). But 22 percent of the anastomoses found in the present survey
are sufficiently distinctive to justify their segregation into new categories
which for convenience are designated as Types IV and V. (Fic. 6).

In both of the new types of anastomoses, the areole is formed by the
union between an unbranched arcuate vein and one of the branches of its
sister vein lying above it (Fic. 14). This pattern is clearly different from
eae represented by Types I to IIT anastomoses in which union occurs
sang two of the branches of a pair of adjacent vein dichotomies (cf.

Ics. 4 and 6). As shown diagrammatically in Fic. 6, Types IV and V

1968 | FOSTER, CIRCAEASTER

Fic. 5. Floral bract showing the rare occurrence in Circaeaster of two vein
approximations, each forming a greatly elongated areole “open” at its distal an,
x 24.

60 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

TABLE 1. Summary of anastomoses and vein approximations in leaves
of cultivated plants of Circaeaster agrestis

PLANTS PLANTS
GROWN GROWN
IN 1965 IN 1966 TOTALS
TOTAL NUMBER
OF PLANTS 40 40 80
TOTAL NUMBER
OF LEAVES 2248 va A 5020
NUMBER OF
LEAVES WITH
ANASTOMOSES 29 25 54
NUMBER OF
PLANTS WITH
ANASTOMOSES IN
2 OF MORE LEAVES 8 4 7
NUMBER OF
LEAVES WITH VEIN
APPROXIMATIONS 45 75 120
NUMBER OF PLANTS
WITH VEIN
APPROXIMATIONS
OR MORE LEAVES 11 18 29
% OF LEAVES WITH
ANASTOMOSES LZ 0.9

TABLE 2. Percent of anastomoses in plants with vein unions in two or more leaves

TOTAL NUMBER NUMBER OF LEAVES % OF LEAVES WITH

PLANT NUMBER LEAVES WITH ANASTOMOSES =ANASTOMOSES

Sa 30 2 6.6
15a 109 4 1.8
loa 93 2 ZA
27a 109 3 2.7
29a 145 3 2.0
33a 80 2 25
34a 45 2 44
35a 45 2 4.4

bi 95 2 21
19b 80 2 25
21b 84 4 47
26b. 60 4 66

TOTALS vie 30

1968] FOSTER, CIRCAEASTER 61

anastomoses are morphologically connected, by corresponding vein approx-
imations, with an asymmetrical form of open dichotomy which commonly
occurs in the leaves of Circaeaster. This pattern is termed ‘‘asymmetrical”
because one of the derivatives of a vein-dichotomy remains unbranched
throughout its course while its sister derivative bifurcates one or more
times before reaching the distal margin of the lamina (Fics. 9, 11, 18, 21,

With the preceding analysis as a general basis, it is now appropriate to
describe in detail the new forms of anastomoses and their interrelationships
with vein approximations,

Type IV Anastomoses. The distinguishing character of this type of
anastomosis is the very short extent of xylem-contact between the arcuate
vein and the branch of a contiguous vein-dichotomy (Fic. 6). After
separation of the two strands, the upper portion of the arcuate vein either
terminates in a marginal tooth or ends blindly in the lateral margin of the
lamina (Fics. 11

The resemblances between vein approximations and Type IV anasto-
moses are indeed striking and support the idea that the two patterns differ
from one another only in degree (compare Fics. 7, 10, 11). In some cases,
a relatively wide strip of parenchyma separates the region of approximation
between the arcuate vein and its contiguous strand (Fic. 8). But very
commonly, the interval is so narrow that it is impossible to draw a distinc-
tion between an anastomosis and a vein approximation until a leaf has
been thoroughly cleared in NaOH and chloral hydrate. Fic. 7 is a good
example of this because it shows the extremely close approximation be-
tween the two veins at the upper left margin of the leaf. Additional illus-
trations of closely approximated veins are evident in Fics. 9, 10, and 12.

Type V Anastomoses. In this type of vein union, the tip of an
arcuate unbranched vein converges toward and becomes confluent with
the branch of a vein-dichotomy situated above it. Two “variations” of
this kind of anastomosis have been discovered and, as is shown in Fic. 6,
each of them is morphologically related to a corresponding type of vein
approximation.

In the simpler of the two “variations,” the arcuate vein joins the outer
branch of a first order vein-dichotomy, creating a conspicuous and rela-
tively symmetrical areole (Fics. 14, 16-19). The exact point of anatom-
ical union between the xylem strands fluctuates. Commonly the two
xylem strands merge well above the level of the adjacent vein dichotomy
(Fics. 14, 16-18) but the union may take place very near the point of bi-
furcation of the vein (Fic. 19). An interesting example of the arrested
development of a Type V anastomosis is shown in Fic. 15. Examination
of this leaf under high magnification reveals that the terminal tracheary
elements of the marginal arcuate vein at the left are not in direct contact
with the xylem of the neighboring vein, and hence that the areole is of
the “open” type

62 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

=i Type IV anastomosis

Vein
approximation
ps Type V anastomosis
Open
dichotomous
venation

y

Vein
approximation
Type V anastomosis
6. Diagrams ap ncegphe: the morphological om ge between open

Fic
dichotomous venation, vein approximations, and Types IV and V anastomoses.
Further explanation i in vat

In the other Type V “variation,” an areole is created by the union of
the arcuate vein with the outer branch of a second order vein-dichotomy
lying above it (Fics. 6, 21-23). Vein approximations which morpholog-

1968 | FOSTER, CIRCAEASTER 63

ically connect such anastomoses with open venation are not common but
the leaf shown in Fic. 20 apparently developed such a transitional form.

When examples of Types IV and V anastomoses were first encountered
in this survey it seemed possible that they might be restricted to that part
of the vein system adjacent to the lateral margins of the lamina. (Fics.
13, 16, 21, 22). Further study however clearly indicated that both of the
new types of anastomoses may also occur in a more central position in the
venation (Fics. 11, 18, 19, 23).

DISCUSSION

The various forms of vein approximations which occur in the leaves of
Circaeaster are regarded as morphological “transitions” which connect the
different types of anastomoses with open dichotomous venation (Fics. 4—
7). This conclusion seems fully justified by the present comprehensive
survey and it is important in this connection to note that “near fusions” of
veins were also observed by Arnott (1959, p. 408) in the foliage leaf of
Ginkgo and by Arnott and Tucker (1963, p. 822) in the petals of Ranun-
culus repens var. pleniflorus. In both of these examples, anastomoses are
considerably more frequent than in Circaeaster but the fundamental pat-
tern of venation is dichotomous. It is of further comparative interest that
anastomoses and vein approximations remarkably similar to those in Cir-
caeaster, also occur in the dichotomous venation of certain living ferns. In
a brief unpublished study on the venation of Anemia herzogti Rosenstock,
I observed that the most common types of anastomoses correspond to
Types I and II (see Fic. 4). In proportion to the relatively high number
of dichotomously branched veins in each pinna however, anastomoses and
their coordinated types of vein approximations in this fern seem just as
sporadic and “unpredictable” in position as in the leaves of Circaeaster.’

These brief comparisons between Circaeaster and other living taxa with
dichotomously veined foliar appendages clearly support my concept that
a parallel evolution of similar types of anastomoses has very probably oc-
curred in morphologically divergent and taxonomically unrelated plants
(see also Foster 1961, p. 974 and 1966, pp. 596, 599). But even with this
idea in mind, the question nevertheless arises as to the original or most
ancient type or types of vein anastomoses in vascular plants as a whole.

Although there appear to be very few published descriptions of the de-
tails of venation patterns in extinct groups of the Pteropsida, Types I and
II anastomoses, as defined in the present paper, seem to represent extremely
ancient trends in the evolution of areolate venation. Both of these types

* According to Mickel’s (1962) study on Anemia, the pinnae of species classified in

brichtii, a hybrid between A. (subg. Coptophyllum) ‘apg and A. g.
Anemia) phyllitidis, the venation is “intermediate” in character, i.e., rather weakly
anastomosed. An intensive study of venation patterns Abana, ‘the genus Anemia is

m
obviously highly desirable.

64 JOURNAL OF THE ARNOLD ARBORETUM | voL. 49

of vein unions were observed in Neuropteris and other fern-like plants of
the Paleozoic by Potonié (1912, pp. 111-117) and Type II anastomoses
evidently created the lax areolate venation characteristic of various species
of Ctenis (Florin 1933, pp. 53-80) and Gangamopteris (Zimmermann 1959,
p. 284 and Fig. 148b). Very recently, my attention has been kindly
directed, by Mr. William Glen of the Department of Paleontology at
Berkeley, to the illustrations of typical Type II anastomoses in Psygmo-
phyllum flabellatum (= Ginkgophyton flabellatum) given by Seward
(1919, p. 83, Fig. 666). In this Paleozoic plant of uncertain affinities,
Seward observed that “the veins spread from the base of the lamina and
are repeatedly forked; they may be very numerous and in some forms
occasionally anastomose, as in P. flabellatum”’ etc.

On the other hand, how should one interpret phylogenetically the ap-
parently extremely simple type of anastomosis in which an areole is
created by the union of the two branches of a single vein-dichotomy? Is
this form of anastomosis, which is the most common type in Ginkgo
(Arnott 1959, Fig. 3), “primitive” or “advanced” in vascular plants as a
whole? As far as I am aware, this kind of anastomosis, designated as
“Type A” by Arnott, has not been recorded in paleobotanical literature.
In living plants aside from Ginkgo, however, “Type A” anastomoses occur
in the dichotomous venation of certain cycads (e.g. Stangeria, Dioon,
Encephalartos, and Zamia), in a few ferns which I have personally exam-
ined such as Stenochlaena palustris (Burm.) Bedd. and Anemia herzogit
Rosenstock, and in the petals of Ranunculus where, however, they are very
infrequent (Arnott & Tucker 1963, p. 827). A single example of a “Type
A” anastomosis was found in my survey of Circaeaster but the leaf in
which it occurred was so damaged in processing that it had to be discarded.
In a few leaves, as illustrated by Fic. 24, a very rare type of vein approx-
imation was noted which might be regarded as “transitional” between a

Type A” anastomosis and a normally bifurcated vein. It should be clear
peg this brief discussion, however, that the “phylogenetic status” of

Type A” anastomoses in the living ferns and seed plants must for the
present remain an entirely open question.

There remains for final consideration the morphology and possible phylo-
genetic significance of Types IV and V anastomoses (Fics. 6, 14). I know
of no record of these forms of anastomoses in extinct tracheophytes but I
have seen excellent examples of Type V unions in the marginal veins of
certain pinnae in the leaves of Anemia herzogii. Among angiosperms, Type
V anastomoses are abundantly formed during the ontogeny of the venation
in the cotyledons of Pulsatilla vulgaris. In this case, the anastomoses ate
interpreted by Héster and Zimmermann (1961, pp. 85 86, p. 93 and Figs.
12-14) as morphologically comparable with the “Goniopteridis” type of
vein union previously recognized by Zimmermann (1959, p. 277, Fig. 142)
as primitive in his phylogenetic series of reticulate venation patterns in

Pteridophylls.” Type V anas omoses are also common in the dichotomous
venation of the petals of Ranunculus. Arnott and Tucker (1963, 1964)
regard these anastomoses as a “variation” of their “Type C” vein union

1968 | FOSTER, CIRCAEASTER 65

(= Type II anastomosis in Circaeaster). On the basis of these few, scat-
tered examples it is clearly impossible, however, to reach a phylogenetic
conclusion, although I suspect that both Types IV and V anastomoses
ultimately may prove to be relatively “primitive” in an evolutionary sense.

In conclusion, I wish to emphasize the need for more detailed attention
by paleobotanists to the diversified trends of specialization of areolate
venation in extinct members of the Tracheophyta. It should be more
widely recognized, in my opinion, that the presence or absence or degree
of development of anastomoses in any given system of dichotomous vena-
tion has phylogenetic significance in addition to its diagnostic importance
for purposes of identification and classification. The same viewpoint is
justified, I believe, with reference to the occurrence of dichotomous vena-
tion and sporadic anastomoses in the leaves of certain living seed plants.
An excellent illustration is provided by Arnott’s (1959) discovery of
various types of anastomoses in the dichotomous venation of such a “living
fossil” as Ginkgo biloba. Napp-Zinn (1966, p. 166) in his review of leaf
anatomy in gymnosperms points out that anastomoses also occur in the
dichotomous venation of two living genera of cycads (e.g. Stangeria and
Macrozamia). In a study of leaf venation in Zamia now in progress, I
have found striking examples of anastomoses entirely comparable, in a
mor phological sense, to the Types I and II vein unions which occur in the
leaves of both Kingdonia (Foster 1959, Foster & Arnott 1960) and Cir-
caeaster (Foster 1966). Whether the venation patterns in these two
remarkable dicotyledons represent (1) the “retention” of a condition which
appeared during the early evolution of angiosperms or (2) are the result
of “reversion” from a more complex vasculature, are questions which can-
not be convincingly answered until future paleobotanical discoveries have
clarified the origin and major trends of phylogeny of the angiosperms as a
whole.

LITERATURE CITED
Arnott, H. J. 1959. Anastomoses in the venation of Ginkgo biloba. Am. Jour.
Bot. 46: 405-411.

. C. Tucker. 1963. Analysis of petal venation in Ranunculus. I.
Anastomoses in R. repens v. pleniflorus Am. Jour. Bot. 50: 821-830.

& 1964. Analysis of petal venation in Ranunculus. 11. Number
and pepe ‘of dichotomies in R. repens var. pleniflorus. Bot. Gaz. 125:
13-2

elo U. C. 1964. Circaeaster agrestis Maxim. a hase a ae
Ww record from North Garhwal Himalaya. Bull. Bot. Surv. India
297, 298,

Fiorin, R. 1933. Studien iiber die Cycadales des Mesozoikums nebst Erérter-
ungen iiber die Spaltéffnungsapparate der Bennettitales. Sv. Vet-akad.
Handl. IIT. 12(5): 1-134. 15

Foster, A. S. 1959. The morphological and taxonomic significance of dichot-
omous venation in shes tts uniflora Balfour f. et W. W. Smith. Notes
Bot. Gard. Edinb.

. 1961. The Die iopenesie significance of a venation in angio-

sperms. Recent Advances in Botany 2: 971-9

66 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

1963. The — and relationships of Circaeaster. Jour. Arnold

Arb. 44: 299-3

1966. a of anastomoses in the dichotomous venation of

Sacra Am. Jour. Bot. 53: 588-599.

H. J. Arnott. 1960. Morphology and ge te vasculature of the
ae of Kingdonia uniflora. Am. Jour. Bot. 75:

GOEBEL, K. 1922. Gesetzmassigkeiten im price og Abhandl. Heft. 1.
Gustav Fischer, Jena.

Hoster, H.-R. & W. ZIMMERMANN. 1961. Die Entwicklung des Leitbiindelsys-
tems im Keimblatt von Pulsatilla vulgaris mit besonderer Beriicksichtigung
der Protoxylem-Differenzierung. Planta 56: 71-96.

MickEL, J. T. 1962. A monographic study of the fern a Anemia, subgenus
Coptophyllum. Iowa State Jour. Sci. 36(4): 349

Napp-ZINN, K. 1966. Anatomie des Blattes. L. ae der Gymnosper-
men. Handb. d. Pflanzenanatomie. Band VIII, Teil 1. Gebriider Born-
traeger, Berlin-Nikolassee.

Potonié, H. 1912. Grundlinien der Pflanzen-Morphologie im Lichte der Palaeon-
tologie. Gustav Fischer, Jena

SEWARD, A. C. 1919. Fossil Plants. Vol. IV. University Press, Cambridge

ZIMMERMANN, W. 1959. Die Phylogenie der Pflanzen. Gustav Fischer, Stutt-
gart.

DEPARTMENT OF BOTANY
UNIVERSITY OF CALIFORNIA
BERKELEY, CALIFORNIA 94720

EXPLANATION OF PLATES

PLATE I
Fic. 7. Cleared leaf showing, at left, the approximation between an un-
branched vein and a b derived from an adjacent oe This
type of vein approximation is represented diagrammatically in Fic. 6, X 1
PLATE II

Fics. 8-13. Venation patterns showing oe between vein approx-
imation and Type IV anastomoses. Fic arginal areole at upper left
orm

by convergence and abrupt Se smace: ape adjacent $
9. Vein approximation similar to one in Fic. 8 but the two veins nearly
confluent, x 3 Very close approximation between two veins forming
a elongated “open” submarginal areole, 7 11. Leaf with a Type I

open” submarginal bre re x ay . 13. Leaf with a ih IV perenf
(lower right margin) formed by tbe transient union betwe n unbranched
blindly terminating vein and the adjacent branch of a vein- alichothe x 4.5.
US, unconnected vascular strand.

PLATE Ill

Fig. 14. Small floral bract showing, near the middle of the lamina, a striking

example of a Type V anastomosis, < 32.

PLATE IV

Fics. 15-19. Fic. An unusual type of an “arrested” anastomosis in which
the tip of the asbraeuet arcuate xylem-strand at the left margin ends a few

1968 | FOSTER, CIRCAEASTER 67

cells below the outer branch of the vein-dichotomy ye above it, X 7. Fics.

16-19. Various examples of Type V anastomoses. Fic. Pri rimary leaf with

anastomosis at right margin of lamina, & 5.5. Fic. 17. "Typical oe igre of
sine leaf showing a conspicuous submarginal areole, x 3.8. Fie. Similar
x

of a vein-dichotomy. Note blind vein-ending between two marginal teeth a
lower left side of lamina, x 5.5. US, unconnected vascular strand.

PLAT

Fics. 20-24. Fic. 20. Leaf illustrating approximation ieoy: ee vein
(at right) and adjacent branch of second order vein-dichot This type of
: aye

x 5.5. Fic. 24. Leaf with very uncommon type of vein approximation (at left
margin) in which the branches of a single vein-dichotomy converge and then
separate without xylem-fusion, X 4.8. rt, rudimentary marginal tooth; US, un-
connected vascular strand.

Jour. ARNOLD Ars. VOL. 49

Pek
a

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ae
ee
oe

Foster, CIRCAEASTER

Jour. ARNOLD ARB. VOL, 49 Pirate II

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Foster, CIRCAEASTER

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Jour. ArNoLD Ars. VoL. 49

, CIRCAEASTER

FOSTER

Jour. ARNOLD Ars. VoL, 49 Piate IV

18 19

Foster, CIRCAEASTER

Jour. ARNoLtD Ars. VOL. 49 PLATE V
20 21 ’
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Foster, CIRCAEASTER

1968] MUNZ, ASIAN SPECIES OF DELPHINIUM ie

A SYNOPSIS OF THE ASIAN SPECIES OF DELPHINIUM,
SENSU STRICTO *

Puitre A. Munz

GROUP VII

Like Group VI, but the leaves all basal; none, or rarely one, cauline;
finely dissected into laciniae mostly 1-3 mm. wide.

Not eee eee to any of the pauls Tien proposed for
the genus

Key TO SPECIES

1. Spur not as long as the blade of the upper sepal; plants to 2.5 dm. t
2. Stem strigulose in upper part; leaf blades 3-6 cm. across; bra deep
violet, minutely strigulose, 2-3 cm. long; spur 15-20 mm. long. Yunnan.
133. D. likiangense.
2. Stem sparsely stiff-hairy; leaf blades 2-3 cm. across; sepals pale blue,
yellow-hairy, ca. 3 cm. long; spur 20-25 mm. ~~. Ws a es
0. D. chrysotrichum.

1. Spur equal to or longer than blade of upper sepal
ant not more than 1.5 dm. tall; flowers 1-3 on a stem (several in D.
souliei); leaf blades 1-2 cm. in diameter.
pur 3 cm. long; sepals ieee purple; upper petals ea yellow.
SIRARB cs inst. iv eariousiaks tee eee ae ee ui.
4. Spur 2-2.4 cm. long; sepals blue to purplish blue; sa bluish.
Szechwan
-F Flowers mostly several in an elongate raceme; pubescence spread-
ee a ee 138. D. souliet.
5. Flowers solitary at end of scape; oaacieere erties Nee wee aan
a eR ore spay eee yeast Ue gol a aol aA ih Ie gee . D. sino-scaposum.
3. Plant taller; flowers often more numerous; leaves bs rger.
6. Axis of inflorescence and the pedicels quite glabrous except for a few
hairs just below the flowers.
7. Sepals about 20 mm. long; spur 25 mm. ges petals bluish.
Yunnan. (Treated under Group VI). ........ . D. au
7. Sepals — mm. long; spur 19-21 mm. ‘aa petals black.
CTI ee a 131. D. erlangshanicum.
6. Axis of sige and the pedicels more generally pubescent.
8. Upper sepal 23-24 mm. long; spur 27 mm. long. Szechwan. .....
136. D. saxatile.

8. Upper sepal 12-20 mm. long.
9. Bracteoles well below the flower.
10. Upper petals entire at the tip; leaf laciniae 2-3 mm. wide.
Vee 135. D. pycnocentrum.

* Continued from volume 48, p. 545.

74 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 49

10. hag petals 2-toothed; leaf laciniae 1 mm. wide. Szech-
OGLE re eee oan eee er ee 138. D. souliet.
Brats almost against the flower; upper ate emarginate
at t

ae "Seiials 11-14 mm. long; spur 19-24 mm. long; petals blue.
12. sea _— emarginate at gy — pubescent.
pie a aires 134. D. pycnocentroides.

if. ae aid entire at tip; follices glabrous. Tibet
PRIS RE ACES rey ea a RD, thibetioums.
11. Sepals 15-18 mm. es spur 14-20 mm. ong, upper petals
ere GOS 8 ca ee . D. taipaicum.

ye

130. Delphinium chrysotrichum (sphalm. chrysotricum) Finet &
Gagnep. Bull. Soc. Bot. France 51: 488. pl. vii, B. 1904. Fic. 12, A.
Delphinium chrysotrichum var. pygmaeum Ostenfeld in Hedin, Southern Tibet
6(3): 79. pls. 1, 2. 1922. (A list of flowering plants from inner Asia col-
lected by Dr. Sven Hedin [compiled by Ostenfeld & Paulsen]).

Low, to 1.5 dm. tall, subprostrate, divaricately branched, sparsely stiff-
pubescent above, subglabrous below, striate; leaves all radical; petioles
ca. 5 cm. long, dilated at base; the hairy blades to 3 cm. in diameter,
roundish, 3-parted into cuneate segments, the 2 lateral 2-lobed, the middle
one 3-lobed, the lobes 3-5-toothed at the apices, obtuse, mucronate,
flowers ‘12 ona branch; bracts pubescent, ca. 1 cm. long, petioled, 3-
parted into narrow lobes; pedicels 2—7 cm. long, pubescent; bracteoles 2,
pubescent, lance-oblong, ‘obtuse, ca. 1 cm. long, placed near the middle
of the pedicel; flowers pale blue, yellow-hairy without; upper sepal
broad-ovate, 25-30 mm. long, obtuse, the spur straight or slightly curved,
20-25 mm. long, 6 mm. wide at base, obtuse; lateral sepals elliptic, 25
mm. by 12 mm., pubescent, emarginate-truncate at tip; lower sepals 25
mm. by 14 mm., obtuse; upper petals with the limb almost straight,
glabrous, emarginate at apex, 15-17 mm. long, the spur 20-22 mm. long;
lower petals oblique, the limb oblong, 7 mm. by 6 mm., 2-lobed over half
its length, bearded over most of lower part, sparsely short ciliate, the
claw 6-7 mm. long; stamens glabrous; follicles 3, bearded.

fied Tibet, prov. Batang, Zambala, Aug., 1903, Soulié 3039 (P);

ILLUSTRATION. Var. pygmaeum OSTE
NF
Fl. Pl. Inner Asia. pl. 1, 2. 1922. ELD in Ostenf. & Paulsen, List

EXAMPL
3905 (p). ES SEEN. Tibet oriental: Yargong, Batang province, Soulié 3904 (P),

131. Petey meio erlangshanicum W. T. Wang, Acta Bot. Sinica 10:

a 3 on vig leaves about 4, basal, long petioled, glabrous; blades
linear, - pee ce OF VaR, 6. 5-9 cm. wide, the ultimate lobules
5 mm. wide, revolute, entire or 1-lobuled: petioles 14-18 cm.

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 75

long, broad-vaginate at base; scapes erect, 3 dm. tall, retrorsely crisped-
pilose and with looser yellow hairs intermixed, racemiferous above the
middle; racemes ca. 6-flowered, lax; lower bracts 22 mm. long, lanceolate,
the upper 13-16 mm. long; pedicels 2-6 cm. long, suberect, slightly in-
curved; bracteoles near the flowers, lanceolate, 10-11 mm. long, 1.6-1.8
mm. wide, puberulent; flowers suberect, blue; sepals ovate or elliptic-
ovate, puberulent, 14-16 mm. long; spur 19-21 mm. long, subulate, 3.5—4
mm. wide at base, strongly decurved; upper petals black, bifid into lance-
olate lobes, loosely ciliate; lower petals black, the limb narrow-elliptic,
bifid to below the middle into narrow lanceolate lobes, silky villous above
the base; stamens glabrous; carpels 3, densely puberulent.

Type: Erlangshan, Tienchuan, Szechwan, W. K. Hu no. 37800 (PE), not
seen; photograph (£) seen.

132. Delphinium hui Chen, Bull. Fan Mem. Inst. Biol. Peiping (n.s.)
42173; 1948.

Stem 3-3.5 cm. long, with long dense hairs; leaves all radical, long
petioled, deeply 3-partite, 1.5 cm. long, 2 cm. wide, the parts 3-lobed,
cuneate, the lobes ovate-acute, densely pubescent above, long-pilose be-
neath; peduncles 3 cm. long, long white-villous; flower horizontal, solitary,
greenish purple; sepals round-ovate, the upper and 2 lower prolonged into
a short tip, the 2 lateral rhombic-rounded, densely pilose without, sparsely
so within, ciliate on margin; spur 3 cm. long, straight or somewhat de-
curved, densely hairy; upper petals sordid yellow, the laminae 6 mm.
long, ovate, acute, the spur almost as long as the sepal spur; lower petals
bluish yellow, the laminae oblong-ovate, apically bifid, pilose and ciliate;
stamens 5 mm. long, the filaments ciliate; carpels 3, glabrous with styles
6 mm. long.

Type: S. Sikang, Konkaling, Chandoye, at 4500 m., T. T. Yu 13101,
Aug. 31, 1937 (pE?), not seen. Described as having the general appear-
ance of a violet-colored pansy.

133. Delphinium likiangense Franchet, Bull. Soc. Philom. Paris VIII.

3: 180. 1893. Fic. 12, sath

Delphinium oliganthum Franchet, Pl. Delavay. 1: 29. pl. 8. 1899; not Boiss.,
1867.

Perennial from a creeping woody rhizome, the stem subglabrous, simple,
1-2.5 (—5) dm. tall, strigulose in upper part; leaves basal, or sometimes
also 1 above; petioles 5—12 cm. long, slender, glabrous, dilated at base;
leaf blades 3-6 cm. wide, subglabrous, thickish in texture, divided nearly
or quite to the base into 5 segments which are in turn deeply divided into
subobtuse linear lobes 2—3 mm. wide; flowers 1-3-5 on long pedicels, the
bractlets lanceolate, commonly 1.5—2 cm. long, 3-5 mm. wide, placed just
below the sepals; sepals deep violet, broad, minutely strigulose; upper
sepal 2-3 cm. long, 1.5-2 cm. wide, broadly ovate, obtuse, the spur

76 JOURNAL OF THE ARNOLD ARBORETUM [voL, 49

troides; blade of upper petal 1 ] :
mm. long, claw 5 mm. ; drawn from Forrest 6643 (P

, Claw 9 mm.; d 18
; blade of upper petal 10 mm. long, spur
5 mm. long, claw 6 mm.; drawn from Soulié 3049 (P).

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 77

straight, horizontal, 15-20 mm. long, ca. 5 mm. wide at the base, narrowed
to an acute point; lateral sepals 18-22 mm. long, 14-15 mm. wide,
rounded at the tip; lower sepals 25-30 mm. long, 15-20 mm. wide,
pointed; upper petals blue, the laminae almost straight, unequally 2-
toothed, often with a few short hairs and ca. 15 mm. long, narrow, the
spur 16 20 mm. long, straight; lower petals with a claw 6-7 mm. long,
the lamina narrowly oblong-ovate, bearded, ca. 10 mm. long, cleft into 2
lance-oblong lobes ca. 5 mm. long; stamens 8-10 mm. long, the filaments
slightly ciliate; anthers dark, 1 mm. long; follicles 3, finely hairy, 12-14
mm. long, 5 mm. wide, the styles an additional 1.5 mm.; seeds gray-
brown, subglobose, 2—2.5 mm. long, longitudinally winged at the angles.

Type: Yunnan, on calcareous hills of Likiang, Suee-chan, at 4000 m.,
Delavay 2484 and 2201. At Paris number 2201 is designated as TYPE
and has the parts dissected and drawn.

ILLustrations. Plantae Delavayanae 1: pl. 8. 1889 as D. oliganthum;
Gardeners’ Chronicle III. 60: 128. 151. 1916.

DistrisuTIon. About 3200 to 4600 m., in Yunnan, apparently largely
in the Likiang or Lichiang Snow Range.

EXAMPLEs. C. Schneider 3251, 2928, 3774; G. Forrest 11360, 11052, 11203,
22291, 16949, 3768, 6634, 6483, 2903, 15322, 15170; Rock 25330, 24920, 24914,
17332, 6111, 10665, 10792, 10409, 11413, 11435, 5242, 9867; Ching 30572.

134. Delphinium pycnocentroides W. T. Wang, Acta Bot. Sinica 10:
163-164. 1962 Fic. 12, D, E.
Delphinium sibiabicsee Finet & Gagnep. ah schizophyllum Hand.-Mazz.
Symb. Sinicae 7: 275. 1931. Based on Handel-Mazzetti 4100 from near
Lidjiang (Lichiang) (w), not seen; but he no, 7671 was also cited, and that

I have seen (£, GH).

Stem 3-8 dm. tall, 2-4.5 mm. thick at base, terete, densely retrorse-
strigulose, sometimes glabrous below, simple or rarely 1-branched; basal
leaves ca. 3, long petioled, the leaf blades suborbicular or round- reniform,
8-12 (-20) cm. wide, cordate, trisect, the middle segment rhombic-obovate,
cuneate at base, petiolulate, twice subpinnately parted, the ultimate lobes
lance-linear, 1 4-5 mm. wide, slightly revolute; lateral segments obliquely
flabellate, twice as wide as the middle one, narrow-cuneate at base or
sometimes petiolulate, unequally 2-parted, the upper parts like those of
the middle segment, but more or less oblique; petioles 8-18 (—24) cm.
long, glabrous, narrow-vaginate; cauline leaf 1, smaller than basal;
cemes, elongate, narrow, 5—many-flowered, lax: pedicels 2—6 cm. sag
divergent, densely strigulose and with few spreading yellow glandular
hairs; bracteoles next to the flower, lanceolate, 6.5-11 mm. long, 1.2-1.6
mm, wide, purplish blue, ciliate; sepals elliptic-ovate or -obovate, purplish
blue, 12-14 mm. long, puberulent: spur 19-24 mm. long, cylindric-sub-
ulate or subulate, 2-3 mm. thick at base, obtuse, straight or strongly
decurved; upper petals blue, emarginate or bilobulate, glabrous; lower
petals blue, the laminae narrow-elliptic, bifid into 2 obtusish lance-deltoid

78 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

lobes below the middle, long-ciliate, yellow bearded at base; filaments
loosely ciliate to glabrous; follicles 3, spreading-pubescent, 12-14 mm.
long, the styles an additional 2.5 mm.; seeds obovate-triquetrous, 1.5—2.5
mm. long, brown, winged on the angles.

Type: From Yunnan, K. M. Feng 9033, Aug. 15, 1942 (PE?), not seen.

Rance. In Yunnan, especially about the Lichiang Range, at elevations
of 3500 to 4500 m.

ExaMpLes. Forrest 6533; Rock 17338, 10657, 17319, 10813, 6135, 3787,
10670; C. Schneider 2278; Handel-Mazzetti 7671; T. T. Vii 13822; K. M. Feng
2125; R. C. Ching 30566.

Wang described also var. latisectum (Acta Bot. Sinica 10: 164) with
hairs of stems spreading; leaf segments broader; based on C. W. Wang
66226, which I have referred to D. tatsienense.

135. Delphinium pycnocentrum Franchet, Bull. Soc. Bot. France 33:
379. 1886. Fic. 12, F.
Delphinium lankongense Franchet, Bull. Soc. Bot. France 33: 378. 1886.
Based on Delavay no. 1844, Oct. 12, 1885, Mt. Hee-chan-men (Lankong)
(P, uc), both seen.
Delphinium pycnocentrum var. lankongense (Franchet) Huth, Bot. Jahrb. 20:
465. 1895.

Perennial, from a rather woody root; stem erect, mostly simple, 2-9
dm. tall, subglabrous below to strigose or spreading-pubescent with rather
long fine hairs, more densely strigose and pubescent above, the spreading
hairs more or less enlarged and yellowish at the base; leaves basal, the
petioles often purplish, 1-3 dm. long, subglabrous to strigulose and spread-
ing-hairy, strongly dilated at the base; blades to 12 cm. in diameter,
rounded-pentagonal in outline, 5-parted to the base, then deeply incised
into numerous lance-linear lobes 2-4 mm. wide, sharply acute, subgla-
brous to ciliate; inflorescence racemose, few to many flowered, secund,
peduncled, to 3 dm. long, strigose and with spreading hairs pustulate at
the base; bracts entire to trifid, erect, 1-4 cm. long, mostly lanceolate;
pedicels strict, appressed, 1-6 cm. long; bracteoles linear to linear-lan-
ceolate, situated well below the flower; calyx bluish purple, pubescent;
upper sepal oblong-obovate, 15 mm. long, 10 mm. wide, ending in an
apiculate tip, the spur 25-28 mm. long, somewhat decurved, 3 mm. wide at
the base, gradually narrowed to a finely pointed tip; lateral sepals rhombic-
ovate, 14 mm. long, 9 mm. wide, obtuse, pubescent along midvein; lower
sepals asymmetrically obovate, 13 mm. long, 6 mm. wide, obtuse, pubes-
cent; upper petals blue, the blade somewhat oblique, 11 mm. long, 4 mm.
wide, rounded, entire at tip, the spur 25 mm. long; lower petals bluish,
the limb obovate-oblong, 7 mm, long, 5 mm. wide, cleft from 2 to 4.5
mm., bearded and ciliate, the claw 5 mm. long; stamens 7—7.5 mm. long,
glabrous; anthers dark, 1.5 mm. long; follicles 3, somewhat pubescent to

subglabrous, erect, 15-18 mm. long, ca. 4 mm. thick, the styles an addi-
tional 3.5 mm. :

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 79

Type: Yunnan, in silvis montis Yang-in-chan, prope Lankong, alt.
3000 m.; Delavay 993, Oct. 8, 1884 (holotype P, isotype K); both seen.

Another collection referable to this species is from Yunnan, “au col de Hu-
chan-men,” Delavay 4220 (P).

136. Delphinium saxatile W. T. Wang, Acta Phytotax. Sinica 6: 367.
1957.

Perennial, scapose, the leaves all basal; leaf blades broadly cordate-
pentagonal in outline, 2.5—7 cm. long, 3.5—7.5 cm. wide, 5-fid into obovate
parts with cuneate base and 3-5 deep lance-linear lobes, these apiculate,
loosely hirsute, ciliate, to ca. 5 mm. wide; petioles 6-15 cm. long, hirsute,
canaliculate below; stem 2-flowered; bracts leaflike, smaller, the petiole ca.
1.5 cm. long; pedicel to 12 cm. long; bracteoles near its middle, linear, 5
mm. long, ciliate; flowers deep blue, pubescent, the sepals acuminate,
ovate-oblong, the upper and lateral 23-24 mm. long, 9-10 mm. wide; spur
27 mm. long, straight or decurved, 3-4 mm. wide at base; lower sepals 28
mm. long, 8.5 mm. wide; upper petals about half as long as the sepals,
the blades oblique-truncate, emarginate, ciliate, the spur 25 mm. long;
lower petals 12.5 mm. long, the blade bifid to below the middle into 2
ovate-triangular lobes rounded at the apex and ciliate, the laminae yellow
bearded near the base, claw ca. 6 mm. long, appendiculate; stamens
glabrous, 7 mm. long; carpels 3, velutinous.

Type: Szechwan, Pao-hsing-hsien; Kuo-pa-yan, at 2100 m., Aug. 8,
1933, T. H. Titi 4366 (PE), not seen; photograph (£), seen.

137. Delphinium sino-scaposum W. T. Wang, Acta Bot. Sinica 10:
165. 1962.
Delphinium scaposum W. T. Wang, Acta Phytotax. Sinica 6: 364. 1957, not
Greene, 1881.

Plant 11-16.5 cm. high; leaves all basal, glabrous, long petioled (4-13
cm.); laminae broadly reniform, 1.5-3.5 cm. long, 2-6 cm. wide, trisect,
the middle segment broadly rhombic-obovate, basally cuneate-attenuate,
3-parted to below the middle, the ultimate divisions to 3.5 mm. wide;
petioles 2-10 cm. long, vaginate; scapes 1-flowered, 9-16 cm. high, re-
trorse-pubescent; bracts sparsely pilose or trifid, 1.5-1.8 cm. long, the
lobes lanceolate or the whole entire; flower purplish blue; pedicel 3-6
cm. long; bracteoles near the flower, lanceolate, 11 mm. long, subglabrous;
sepals broadly elliptic, obtuse, 16.5 mm. long, 12 mm. wide, pubescent;
spur 22-24 mm. long, 3 mm, thick at base, declinate; upper petals slightly
shorter than sepals, upper abruptly narrowed above the middle, emargin-
ate at apex, ciliate, spur as long as sepal spur; lower petals 16 mm. long,
the limb divided to the lower third, loosely silky, as long as the claw which
is appendaged; stamens glabrous, 7 mm. long; carpels 3, sparsely short-
silky.

80 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Type: Szechwan, T. H. Tii 3606 (PE?), not seen; photograph (®£),
seen.

No specimens available. The author of Delphinium sino-scaposum
says it resembles D. likiangense but has 1-flowered scapes; upper petals
emarginate and ciliate, lower not yellow bearded; carpels loosely silky.

138. Delphinium souliei Franchet, Bull. Soc. Philom. Pars Wii. 5:
172. 1893. ¥ig..12,/G, Hi.
Delphinium paludicola Ulbr. Notizbl. Bot. Gart. Berlin 12: 357. 1935. Based

on H. Stevens 87, holotype (F), isotype (Ny), seen; from Szechwan, Tachien-
lu to Sachou, via Mouping.

Stems 1—4 dm. tall, slender, erect, finely hairy or pilose, simple; leaves
in a basal cluster, the petioles very slender, 2-15 cm. long, subglabrous,
vaginate and purplish at base; blades pale green, 1-2 cm. across, un-
equally many partite, the segments 2-3 times divided into linear lobules
1-2 mm. wide with revolute margins and prominent midvein beneath;
flowers mostly 2-15, in a simple elongate raceme; lower bracts frequently
vaginate at base, then with few linear lobes, the upper lanceolate, often
purple, 1-2 cm. long; pedicels erect, pilose, 3-7 cm. long; bracteoles
lance-linear, opposite, about the middle of the pedicel or above; sepals
blue, the upper 15—20 mm. long, acutish, pilose, the spur slightly curved,
20-30 mm. long, ca. 5 mm. wide near the base, very gradually narrowed
to the blunt tip; lateral sepals oblong-ovate, 14-15 mm. long, 7-8 mm.
wide; lower sepals elliptic-ovate, 14-17 mm. long, 7 mm. wide, with an
acutish prolongation at the tip; upper petals pale blue, the blade almost
straight, 14-16 mm. long, deeply notched to shallowly bilobed, the spur
21 mm. long; lower petals tinged with blue, the blade oblong, sparsely
bearded, 6-6.5 mm. long, bifid for ca. 2 mm., the claw broad, 8-9 mm.
long; stamens glabrous, 6-7 mm. long; anthers dark, 1—-1.5 mm. long;
follicles 3, hairy when young, ca. 12 mm. long, reticulate-veined; seeds
fuscous, broadly winged, scarcely 1 mm. long.

‘Type: Szechwan, ‘“‘lieux secs a Tizou, Tché-to-chan, environs de Ta-
tsien-lou,” Sept.—Oct., 1891, R. P. Soulié 364 (Pp); seen.
DisTRIBUTION. In Szechwan and Sikang, 3500 to 4000 m.

REPRESENTATIVE COLLECTIONS. Szechwan: Mousset 27; Soulié 2388; E. H.
Wilson 4689, 3092, 3092a; Harry Smith 4048. Sikang: Harry Smith 11732,

fare Apparently D. paludicola was described for taller plants of Delphinium
souliei.

139, arias taipaicum W. T. Wang, Acta Bot. Sinica 10: 164-
. 1962,

‘ase ca. 22 cm. tall, 3 mm. thick at base, retrorsely silky-pilose or
ommaaeeeny, basal leaves ca. 2, long petioled, the blades pentagonal-reni-
orm, 5-8 cm. wide, trisect, the middle segments sessile, cuneate, trifid

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 81

to middle, the middle lobes obovate-oblanceolate, trilobulate or entire,
the lateral unequally bifid, the lobules ovate, obtusely mucronate, ulti-
mate divisions lanceolate, densely strigulose beneath, loosely villous above:
petioles 7-14 cm. long, loosely retrorse-pilose, narrowly vaginate; racemes
2—4-flowered, 4—7.5 cm. long; lower bracts trisect, short vaginate-petiolate,
the segments trifid or entire, the others oblong-ovate or lanceolate, 14-16
mm. long; pedicels divergent, 1.5-4 cm. long, densely retrorse-pilose:
bracteoles to 2 mm. distant from the sepals, oblong-ovate, 9-12 mm. long,
3—3.5 mm. wide, obtusish, loosely puberulent; sepals purplish, obovate,
15-18 mm. long, loosely puberulent; spur 14-20 mm. long, cylindric-sub-
ulate or subulate, slightly decurved; upper petals yellowish, glabrous,
emarginate; lower petals purplish, the limb yellow bearded, obovate-ellip-
tic, bilobulate to 1/4 its length, loosely ciliate, claw shorter than limb,
appendiculate; stamens glabrous; carpels 3, villous, becoming 15 mm.
long; seeds obovoid-triquetrous, 2 mm. long, dark brown, winged on angles.

Type: Shensi, Tai-pai-shan, Pa-sian-tai, 3900 m., Sept. 15, 1939, T. NV.
Liou & P. C. Tsoong 986 (PE), not seen; photograph (£), seen.

I have had no material of this species which the author differentiates
from Delphinium thibeticum in its trisect leaves and densely villous car-
pels; and from D. pycnocentrum in the wider laciniae of the leaves, densely
retrorse-pilose pedicels, and bilobed lower petals.

140. Delphinium thibeticum Finet & Gagnep. Bull. Soc. Bot. France
51: 489. 1904. Fre. 12, I.
Delphinium thibeticum var. subintegrum Finet & Gagnep. Bull. Soc. Bot.
France 51: 489. 1904. Based on Soulié 3044, E. Tibet, between Zambal &
Yargong, (P); seen.

Perennial, 6-10 dm. tall, from rather woody roots; stem slender, leafless
or with 1 leaf, mostly simple, subglabrous or with scattered retrorse hairs,
these becoming more numerous upward and strigulose in the inflorescence;
leaves 5—7; petioles slender, to 12 or 15 cm. long, dilated at base; blades
sparsely pubescent below, glabrous above, 5-10 cm. in diameter, broader
than long, divided almost to base into narrowly cuneate segments, these
3-parted to about the middle and each part with 2-3 lance-oblong lobules
to ca. 1 cm. long, 3-4 mm. wide, mucronate; inflorescence a simple, rather
lax, many flowered raceme or with 1 or 2 smaller branches; bracts mostly
entire, lance-oblong, 7-10 mm. long; pedicels ascending to suberect,
strigulose, 1.5—5 cm. long; bracteoles subtending the flower, 4-5 mm.
long, lance-oblong; flowers deep blue, rather loosely short-pubescent ;
upper sepals oblong-ovate, 11 mm. long, 7 mm. wide, apiculate, the
spreading-decurved spur 20-22 mm. long, 2.5 mm. wide at the base,
slender; other sepals ca. 11 mm. by 6 mm., elliptic-obovate, obtuse; petals
bluish, the upper lamina rather straight, 100 mm. long, 3 mm. wide, entire
at the glabrous tip, the spur 19-20 mm. long; lower laminae rather nar-
row-oblong, 5 mm. long, 2.5 mm. wide, short bearded from the middle

82 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. 49

out, short-ciliate, bifid 1/3 its length into lanceolate lobes, claw 6 mm.
long, appendiculate; stamens glabrous; carpels 3, becoming ca, 12 mm.
long, the styles an additional 3 mm.

Type: Eastern Tibet, Batang province, Yargong, Soulié 3049 (P); seen,
COLLECTIONS SEEN. Yargong: Soulié 3893 and 3894.

GROUP VIII

Heavy-rooted perennials. Leaves with the ultimate divisions lanceolate
to ovate, mostly over 5 mm. broad. Sepals not persistent and membra-
nous. Petals blue or paler, not almost black. Spur usually not strongly
curved.

This is a purely artificial group, not corresponding to any of the taxo-
nomic divisions of the genus

Key TO SPECIES

L ne obviously longer than the blade of the upper sepal.
Stem and axis of inflorescence glabrous
3. Lamina of upper petal ca. 7 mm. long. Siberia. .. 163. D. laxiflorum.
3. Lamina of upper petal 9-12 mm. long. China.
4. Upper sepal 14-15 mm. long

5. Bracteoles paired near the summit of the pedicel, sometimes a

third one farther down; — of upper petal emarginate; stem

angled, fistulose. Yunn 2. D. georgit.
Bracteoles 2, near the middle of the pedicel; blade of upper
petal erose or entire; stem not angled or fistulose. Szechwan.
6. Stem white-hirsute below; carpels bgt spur often
Meena CUNVOG... 2. ck. . D. laxicymosum.

6. Stem glabrous below; carpels ae iis mostly nearly
RAI ep ees See ge as 169. D. mitzugense.

4. Upper sepal 10-11 mm. long.
7. Bracteoles 3-5 mm. long; ipa glabrous.

8. Lower petal blade ca. 5 mm. wide; racemes few flowered;
petioles 6-9 cm. long. co, Viana...
ee 154. D. glabricaule.

8. Lower petal blade ca. 3 mm. wide; racemes many flowered;
petioles longer. Shansi. 153. D. giraldii.

7. Bracteoles 10-12 mm. long; carpels ae oa (This
species treated in Group VI). .......... 8. D. sutchuense.
2. Stem and axis of inflorescence, or at least the latter, pubescent
9. Pubescence of stem and inflorescence appressed or nearly so
10. Lamina of upper sepal 8-10 mm. long; sepals strigulose: carpels
glabrous.

11. Leaf segments petiolulate; petioles somewhat vaginate at
WONG ee a 183. De

ih, Leal segments sessile; petioles not Vaminate: ....56- 40%
ee ea on 180. D. sungpanense.

wn

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 83

10. Lamina of upper sepal 12-18 mm. long; carpels more or less
b -

pubescen

12. Bracteoles 10-18 mm. long, near the base of the pedicels;

_
Re

149. D. davidii.

sepals yellow-spotted at base on back; spur slightly uncinate
ibet

Bracteoles es 5-10 mm. long; sepals not so spotted; spur

baer at apex

_—
Los)

. Upper petal rounded and entire at apex; flowers pale

blue, the spur 18-22 mm. long. Sikang, Szechwan.
173;.6, pachycentrum.

. Upper petal truncate and minutely bilobed at apex.

14. Flowers ta the spur 25~32 mm. long. North-

west Yunnan te ae: 184. D. wiser
14. cre fntenil blue, the spur 18 mm. long. Bok-
BAYA. Cc hee a 165. D. lipskii.

9. Pubescence of stem ad inflorescence spreading
15. Inflorescence with some hairs yellow and ei
16. Upper sepal blade 18-20 mm, long; lower petal blade quite

abrous. Asi BGG. «2 iy 159. D. knorringianum.

16. Upper sepal blade 10-15 mm. long; lower petals bearded.

17.

i.

Limb of lower petal 8-11 mm. og oblong, _ to
about 3 or 4mm. Yunnan, Szechwan. ..........:....
PRL Aol as eee Fee eas 150. 2. dilated
Limb of lower petal mostly 4-5 mm. long.
18. Bracteoles ca. 7-13 mm. long.
19. Stem subglabrous; i, pape ny in upper third of
pedicel. Szechwan. ....... 170. D. muliense.
19. Stem spreading-hairy; bracteoles subtending
the Homers FINAISVR) ea,
Coie rn, © pea 1 D. drepanocentrum.
. Bracteoles shorter or, if as lone, then the stems
obviously spreading hai
20. The bracteoles 2-3 mm. long, near or above
the middle of the pedicel; spur 20-22 mm.
long; upper sepal 12 mm. long. Tibet........
a See ee a 177. D. sherriffi.
20. a scene mostly 5~7 mm. long.
s of lower cauline leaves 7—12 cm. in
Socks petioles vaginate at base; brac-
teoles in upper fourth of pedicel.
22. Lower petal laminae divided two-
thirds their sana sepals lilac in
. D. lilacinum.
‘ye Lower petal bie divided one-
fourth their ahr sepals dark blue.
174.. 2. pogonanthum.
. Blades of lower cauline leaves 4-7 cm. in

—
i°,<)

NR
—_

bracteoles mostly lower on the pedicel.
23. Sepals minutely puberulent on both

84

i.

JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

sides; upper petals blue; carpels pu-
berulent. Szechwan. ...........:.:
175. D. pseudo-campylocentrum.

. Sepals loosely puberulent to subgla-
brous on the outside, glabrous within;
upper petals yellowish; carpels gla-
Honan. .. 157. D. honanense.

bh
ios)

br
15. Inflorescence lacking yellow glandular hairs.

24. Sepals glabrous, the upper ca. 12 mm. long, spur ca. 15 mm
RE oes rane 5 182.--d). ternatutn.

i)
-

d a with hairs.
The sepals with very fine puberulence.
26. Bracteoles 6-8 mm. long, subtending the flower;

He

(Treated in Group VII). ..... 140. D. thibeticum.
26. Bracteoles smaller, a little below the flower; leaves
up and down the stem. Yunnan. 181. D. taliense.
The sepals with coarser hairs. Yunnan to Tibet
27. Blade of lower petal 6—7 mm. rages sepal spur about
fo a, ae 174. D. pogonanthum.
27. Blade of lower petal 7-10 mm. long; sepal spur
18-72 mim ne 156. D. hirticaule.

Nm
LL

Spur not or scarcely longer than he blade of the upper sepal.
28. et spur considerably shorter than the blade of the me sepal. Eastern

ee ee epee. 167. D. malabaricum.

I
28. The spur about as long as the blade of the upper sepal.

29.

nN
©

Sepals subglabrous
30. Bracteoles 4-5 mm. long; leaf blades to 20 cm. broad; flowers
in short racemes. Northern Iran. (Treated in Group VI). ...-.
ee Cae ee nos eo 101. D. lanigerum.
30. Bracteoles about 2 mm. lon
. Leaf blades to 15 cm. broad; flowers in es open panicle,

pale bie. -Ciigs... 2. ace 8. 179. D. sparsiflorum.
31. Leaf blades to 5 cm. broad; flowers i . — blue. Iran.
(Treated in Group IV). ......... Wee #3 saniculifolium.

Sepals pubescent.
32. Stem glabrous or nearly so
33. Sepals with cornute appendages at tip 1-2 mm. lon
34. Ultimate leaf segments 5-8 mm. wide; upper "petals
entire, blue. Western China. 146. D. ceratophorum.
34. cee leaf segments 2-4 mm. wide; upper petals
emarginate, yellow. Tibet. .. 145. D. ceratophoroides.
33. Sepals lacking cornute terminal ‘appendages. Siberia. .«<.
FT Ta Rete eatin ie, lm RH HAN UES Rg sp Ae Cle ial ling: gti gi hum.
32. Stem pubescent. ae sams
35. Inflorescence lacking glandular hairs
36. The inflorescence with spreading hairs; upper sepals
ca. 12 mm. long. Caucasus. ..... 172. D. osseticum.
The inflorescence with more or less pets hairs.
37. Upper sepal blade ca. 10 mm. long; lower petal
blade entire, not bifid. Szechwan. .......---->
a RU eG ee a 173. D. pachycentrum.

we
oY

1968]

MUNZ, ASIAN SPECIES OF DELPHINIUM

37. Upper sepal blade longer.
38. Blade of upper sepal 20 mm. long

we
aed

39. Upper sepal ca. 8 mm. wide, acuminate;
petals violet. Szechwan.
chon peer 160. D. ssi ee

39. Upper sepal 10-13 mm. wide,
petals very dark. ams OME ee

148. D. dasycarpum.

Blade of upper sepal mostly 13-16 mm. long,

not acuminate.

40. Ultimate leaf divisions long-acuminate;
blade of lower petal rounded, ca. 12
mm. by 10 mm. Tibet.

141. D. acuminatissimum.

40. sot leaf divisions not long-acu-

inat

41. ane petal lamina entire; srg

plant gray-strigulose; spur more

less uncinate at apex. Ural Mis. am
ia 85. D. uralense.

; Mag petal laminae 2-toothed.

2. Flowers light blue; laminae of
lower petal more or less obo-
vate, with broad lobes; carpels
Sirieone. SIAM. 6s Ss a aes
Mes eee 178. D. siamense.

. Flowers deeper blue to violet
or

BSS
a

>
to

purple
: Bracteoles near the mid-
dle of the pedicel; sepals

>
WwW

densely oe —
blue, 13-15 lon
Nepal, Seen Err Gs
detain 143. D. altissimum.

. Bracteoles near the flower.
44, Sepals minutely stri-

=
WwW

166. "D. Iudlowii.
: Sipe blue-violet to
— a some white.

aS
>

Chi
171. "D, orthocentrum.

35. Inflorescence with some glandular hairs.

45.

Upper petals bidentate at tip.

46. Blade of lower petal ca. 10 mm. long; bracteoles

subtending the flower. Himalaya.
176.

D. scaly" florum.

86 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. 49

45. Upper petals entire at tip.
47. Bracteoles somewhat below the flower.
48. Spur 5 mm. thick near the base, incurved;

48. Spur 2.5 mm. thick; lower petals with obtuse
lobes. Lake Marka-kul. .. 142. D. altaicum.
. Bracteoles against the flower.
49. Limb of lower petal ca. 5 mm. long, very
iort-bearded. Yunnan. ....../.--yier
Se ae ear ie 168. D. micropetalum.
49. Limb of lower petal ca. 10 mm. long, with
long hai
50. Inflorescence a crowded, many-flowered
raceme; bracteoles 5—6 mm. long. Tibet.
i ee ee ts 158. D. kingianum.
Inflorescence lax, with few to several
flowers; bracteoles to 1 cm. long. Kam-
CHARA oS. 144. D. brachycentrum.

+
“I

nr
2S

141. Delphinium acuminatissimum W. T. Wang, Acta Bot. Sinica 10:
138. 1962. Fic. 13, B.

About 1 m. tall, smooth, purplish and glabrous above, simple, equally
and densely leafy; midcauline leaves with limb broad-pentagonal, about
6-8 cm. long, 8-10 cm. wide, cordate at base, tripartite to 3 or 4 mm.
above the base, the middle part broadly rhombic, long-acuminate or sub-
caudate-acuminate, trifid to below the middle, the secondary lobes broadly
lanceolate, few toothed or incised-lobulate; the lateral parts broadly
obliquely flabellate, bifid to beyond the middle, the upper secondary lobes
like those of the middle part, unequally bifid, crisped-puberulent; petioles
as long as the laminae, not vaginate; upper leaves gradually smaller;
racemes 8.5-12 cm. long, 8-10-flowered; lower bracts leaflike, the middle
trifid, upper linear; pedicels 2-6 cm. long, divergent, retrorse-puberulent
toward the tip; bracteoles near the flower, linear, 7-12.5 mm. long, puber-
ulent; sepals blue, the upper ovate-elliptic, others oblong, 14-17 mm. long,
crisped-puberulent ; spur 11—14 mm. long, 4 mm. wide at the base, subulate-
cylindric, obtuse, straight, or slightly decurved; upper petals blue or partly
yellowish, glabrous, oblique, triangular, subobtuse; lower petals blue, the
ae slightly shorter than the claw or subequal, obovate or broadly elliptic,
’ Foie wide, enue at apex, entire to emarginate, short-ciliate, yellow

earded near base; stamens glabrous; carpels 5, glabrous.

Type: Tibet, Lung-tze, Aug. 20, 19 ;
photo. (£), seen. g. 20, 1960, K. N. Fu 1146 (Pe), not seen;

Pig sacra: erations referable here, from Tibet, Raleing, 5000 m.,
Aug. 26, 1924, no collector given (x); Karola, Aug. 28, 1927, F.M.B. ();
Trakan La, Char Chu, Ludlow & Sherriff 1987 is Pee

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 87

142. Delphinium altaicum Nevski in Komarov, FI. U.S.S.R. 7: 161,
426; 1937, Fic. 13, C.
Delphinium dictyocarpum var. pubiflorum Trauty., Bull. Soc. Nat. Mosc. 33:
81. 1860; teste Regel, on sheet at Kew, collected by Ludwig in Siberia
Itaica.
Delphinium elatum L. var. pubiflorum (Trautv.) Huth, Bot. Jahrb. 20: 399.
1895.

Stem 8-10 dm. high, tinged violet below and short-strigulose, somewhat
pubescent and glandular above; leaves glabrous or nearly so above, short-
pubescent on nerves beneath, round-cordate or -reniform, to 2 dm. broad,
sometimes subcuneate at base, palmatifid almost to base into 5 or 7 narrow-
rhombic remote segments which are unequally pinnatifid into rather few
divergent sharply pointed lance-ovate to lanceolate lobules 5-25 mm. long;
petioles to 1 dm. or more long, not vaginate at base; racemes simple,
somewhat dense, 1—2 dm. long, the axis densely short-pilose with spreading
hairs; bracts linear, pubescent, ca. 10 mm. long; pedicels divergent, 1-2
cm. long, densely pubescent; bracteoles linear-filiform, 4-7 mm. long, 0.5
mm. wide, densely glandular-pubescent, located in upper part of pedicel;
sepals bluish, more or less glandular-pubescent, ovate or elliptic-ovate, the
upper sepal ca. 15 mm. long, 10 mm. wide, subacuminate-recurved; spur
1.5—2 cm. long, to 2.5 mm. thick, obtusely acuminate, incurved below the
apex; lateral sepals obovate, 14 mm. long, 8 mm. wide, obtuse; lower
sepals elliptic, 12 mm. long, 7 mm. wide; upper petals blue, the lamina
almost straight, 11 mm. long, 2.5 mm. wide, bearded and ciliate, bifid about
half way into oblong lobes, the claw 6 mm. long, appendiculate; stamens
6-7 mm, long; anthers dark; follicles 3, densely glandular-pilose.

Type: near Lake Marka-kul, Altai Mts., Siberia, B. Keller, Aug. 4,
1908 (LE), seen.

143. Delphinium altissimum Wall. Cat. no. 4718. 1830, nomen;
Plantae Asiaticae Rariores 2: 25. pl. 128. 1831. Fic. 13, D.
Delphinium altissimum subsp. wallichii var. nipalensis Briihl in Bruhl & King,
Ann. Bot. . Calc. 5: 101. 1896. Based on Nepal, Wallich; an 1819
specimen seen at BM. Possibly also var. phallutensis Briihl, Joc. cit., of
which I have not been able to see authentic material.

Perennial from a fleshy branching or simple elongate root; stem 1~1.5
m. tall, terete, strigulose, simple below, somewhat branched above, re-
motely leafy; lower petioles to 3 dm. long, slightly vaginate, the upper
shorter; leaf blades cordate-pentagonal in outline, to 15 cm. across,
palmately 5-partite to 1 or 2 cm. from the base, the segments broadly
cuneate-obovate, 3-lobed or with additional short lobes or large coarse
broad teeth, acute, subglabrous above, pubescent beneath on veins; upper-
most leaves trifid; inflorescence strigose with rather long white hairs, a
many-flowered central rather dense raceme and some additional lax few-
flowered branches below it; bracts lanceolate to linear (or the lowest

88 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Ficure 13. Delphinium, Group VIII. Petals, the upper spurred, the lower
clawed. A. D. sordidecaerulescens of Group VI; blade of upper petal 14 mm.
long, spur 13 mm.; blade of lower petal 7 mm. long, claw 6 mm.; drawn from
Harry Smith 3983 Tyree (ups). B. D. acuminatissimum of Group VIII; blade
of upper petal 14 mm. long, spur 13 mm.; blade of lower petal 9 mm. long, claw
8 mm.; drawn from Ralieng, Tibet (z). C. D. altaicum of Group VII; blade
of upper petal 11 mm. long, spur 15 mm.; blade of lower petal 6.5 mm. long,
claw 6 mm.; drawn from Keller, type (1x). D. D. altissimum; blade of upper
petal 12 mm. long, spur 18 mm.; blade of lower petal 7 mm. long, claw 5 mm.,
drawn from Ludlow & Sherriff 8496 (em). E. D. brachycentrum, plade of
upper petal 12 mm. long, spur 14 mm.; blade of lower petal 10 mm. long, claw
5 mm.; drawn from Eschscholtz type (LE). F. D. ceratophorum, lade of
upper petal 9 mm. long, spur 15 mm.; blade of lower petal 7 mm. long, claw 5
mm.; drawn from Rock 17321 (cH). G. D. cyananthum; blade of upper petal
9 mm. long, spur 13 mm.; blade of lower petal 5 mm. long, claw 5 mm.; drawn

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 89

foliose and 2—3 cm. long, the upper sometimes serrate, pubescent, to ca.
1.5 cm. long; pedicels 1-2(—5) cm. long, divaricate, with 2 linear or lance-
olate bracteoles near the middle, these 4-5 mm. long; sepals deep blue,
densely pubescent; upper sepal ovate, acute, 13-15 mm. long, 5 mm.
wide, the spur 20-22 mm. long, 4 mm. wide at base, slightly curved; lateral
sepals oblong-obovate, 15 mm. long, 10 mm. wide, obtuse; lower sepals
oblong-obovate, 15 mm. long, 8 mm. wide; petals light purple, the upper
blades somewhat oblique, bidentate at apex, 11 mm. long, the spur 17-18
mm. long; lower petals with a blade 6 mm. by 3 mm., deeply bilobed into
lanceolate lobes, the claw 7 mm. long; stamens 8-9 mm. long, glabrous;
anthers rounded-oblong, 1 mm. long; follicles 3, pilose, cylindrical, diver-
gent, 12 mm. by 3 mm., the beak an additional 2.5 mm.; seeds obovoid,
not winged or scaly, round in cross section.

Type: “Crescit in sylvis montanis Nepaliae,” a Wallich specimen from
Nepal seen at BM, probably authentic material.

Rance. Growing in old fields, stony jungle, etc., 1500-2500 m., Bhutan
and Nepal and possibly Assam.

ExAmpLes: Bhutan: Ludlow & Sherriff 3496, 3558; Cooper & Bulley 2556,
2008. Nepal: Wigram 99; Lall Dhwoj in 1928; Colom 282; Ludlow & Sherriff
298. Assam: Kingdon Ward 14015.

144. Delphinium brachycentrum Ledeb. Fl. Rossica 1: ae ne -
Ic. 13, E.

Delphinium stenosepalum Turcz. Bull. Soc. Nat. Mosc. 27(2): 278. 1854.
ype, Tigil, Turczaninow (LE); seen.
Delphinium maydellianum Trautv. Act. Horti Petrop. 6: 7. 1879. Type, Terra
Tschukschorum, G. Maydell Aug. 8, 1869 (LE); seen.
Delphinium cheilanthum Fisch. subsp. brachycentrum (Ledeb.) Huth, Bot.
Jahrb. 20: 345. 1895.

Perennial from fibrous roots, the stem 1.5—7 dm. tall, ashy pubescent to
subglabrous, 4-5 mm. thick, simple or few branched, leafy; lower leaves
on slender petioles to ca. 1 dm. long and more or less vaginate at the base,
the blades mostly wider than long, 5-12 cm. wide, 3-5-parted into sub-
cuneate parts 1-3 or more cm. wide, these in turn coarsely few-toothed or
-lobed, subglabrous to soft-pubescent above, more pubescent, often whitish,

from Schrenck type (LE). H. D. dasycarpum; blade of upper petal 9 mm. long,
spur 10 mm.; blade of lower petal 6 mm. long, claw 4 mm.; drawn from Prescott
I o

petal 7.5 mm. long, claw 5 mm.; drawn from Delavay 74 (Pp). K. D. eae
blade of upper petal 9 mm. long, spur 19 mm.; blade of lower petal 7 mm. long,
5

5m
. g, 7

long, claw 5 mm.; drawn from Dhwoj 0191 (pm). M. D. georget; blade of

upper petal 10 m

90 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

beneath; cauline leaves gradually reduced up the stem; flowers few to
several in rather lax racemes; bracts trifid into lanceolate lobes and to 6
cm. long, or the upper entire, linear, 1-3 cm. long; pedicels ascending-
arcuate, 2-8 cm. long; bracteoles setaceous, to 1 cm. long, a pair near the
flower, often an additional one below; sepals deep blue, with a darker spot
near the apex, villous, subacuminate; upper sepal 15-25 mm. long, 5-8
mm. wide, the spur somewhat curved, 15-20 mm. long, 3—4 mm. wide at
the base, pointed; lateral sepals 16-24 mm. long, 5-9 mm. wide; lower
sepals 18-28 mm. long, 6-8 mm. wide; upper petals bluish at tip, the
lamina almost straight, 12 mm. long, entire, the spur 13-15 mm. long;
lower petals blue, the claw 5 mm. long, the ovate-elliptic blade 10 mm.
long, bearded, divided ca. 3 mm.; stamens glabrous; anthers dark, 1.2
mm. long; follicles 3, hairy.

Type: Kamtschatka, Eschscholtz (LE); seen, very densely pubescent.

Ranging from the Lena-Kolyma district to Chuch Peninsula, south to
Kamtchatka, Bering Island, and Northern Kuriles according to Hultén,
the last area named being in America.

From Kamtchatka I have seen among other specimens: Komarov 3896; Peters
in 1831; Eyerdam 2; Hultén 733, 638.

145. Delphinium ceratophoroides W. T. Wang, Acta Bot. Sinica 10:
158. 1962.

_ Stems erect, ca, 3 dm. tall, 2.5 mm. thick below, smooth, glabrous,
simple or one branched, with ca. 3 leaves; basal or lower cauline leaves
withered by anthesis, the other cauline leaves long petioled and with
broadly pentagonal blades, 2.5-4 cm. long, 3.6-7 cm. wide, cordate, 3-
parted to 2 mm. above the base, the primary median parts broadly obovate-
rhombic, narrow-cuneate at base, 3-parted to middle, the secondary middle
parts broadly rhombic, divided into ultimate lobules 2-4 mm. wide, pri-
mary lateral parts broadly obliquely flabellate, bifid to three-fourths their
length, all densely strigulose beneath, somewhat hirtellous above; petioles
511.5 cm. long; racemes 3-11 cm. long, lax; lower bracts trifid or linear,
the others linear; pedicels 4~5.5 cm. long, divergent, glabrous, bibracteolate
in upper part; bracteoles remote from flower, linear, 5-10 mm. long,
puberulent; sepals blue, elliptic, 18-22 mm. long, strigulose, short-cornic-
ulate, spur 25 mm. long, subulate, 3 mm. thick basally, decurved at right
angle to the obtuse, yellowish, glabrous, emarginate upper petals; lower
petals blue, the limb suboblong, bifid to slightly beyond the middle, the
base yellow bearded, claw ca. as long as limb, appendaged; filaments
loosely ciliate; carpels 3, densely hirtellous.

Type: Tibet, Tsa-wa-rung, Aug., 1935, C. W. Wang 65706 (px), not
seen; photo. (£), seen. No material available for this species. Delphinium
ceratophoroides was described as near D. ceratophorum, but the stems 3-
leaved, glabrous; leaves more deeply divided, the basal ones gone by
anthesis; sepal-tips shorter.

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 91

146. Delphinium ceratophorum Franchet, Bull. Soc. Bot. France 33:

377. 1886. Fic, 13, F.
? Delphinium ceratophorum var. hirsutum W.T. Wang, Acta Bot. Sinica 10:
158. 1962.

Perennial from an elongate rhizome with fibrous coating; stem sub-
scapose, slender, 2~9 dm. tall, mostly glabrous except for the few hairs at
the base and above, simple or slightly branched; leaves largely basal or
nearly so, on slender petioles 5-15 cm. long, these scattered hairy, or
sparsely retrorse-strigose or glabrous, more or less dilated at the base, the
blades roundish in outline, 3-12 cm. across, divided almost to the base
into 3 or 5 cuneate obovate parts which are somewhat hairy and variously
incised into lanceolate or ovate, acute to obtuse divisions 5-8 mm. wide;
cauline leaves few, simpler; racemes 1 to few, lax, mostly 3—7-flowered;
bracts linear to sublanceolate, or 3-parted into linear lobes, 5-15 mm.
long; pedicels arched-ascending, 2~6 cm. long; bracteoles linear or wider,
5—8 mm. long and near the summit of the pedicel; sepals purplish blue,
callous-cornute below the apex, the upper sepal ovate, 18-20 mm. long,
8-10 mm. wide, coarse-pubescent, the spur decurved, 20-22 mm. long, 3.5
mm. wide at the base, coarse-pubescent, attenuate to a narrow tip; lateral
sepals broadly elliptic, 20 mm. by 11 mm., the cornute appendages to 2
mm. long; lower sepals broadly elliptic, 18-20 mm. long; upper petals
blue or bluish at the tip, straight, narrow, the limb 9 mm. long, entire,
the spur ca. 15 mm. long; lower petals blue, the claw 5—5.5 mm. long, the
bearded lamina ca. 7 mm. long, 5 mm. wide, cleft for 2 mm. and with a
prominent appendage at the base; stamens 5-6 mm. long, glabrous;
anthers dark, 1.3 mm. long; carpels 3, hairy.

Tyre: Yunnan, “in monte Hee-Chan-Men; inter dumeta, alt. 3200 m.,”
Delavay 967 (p, K) seen, the K specimen in a type folder. But another
Delavay number, 1845 , from Lao-kouan-tsoui, below San-yn-kay, at 2500
m., is marked Type at (Pp), has dissection drawings, and probably should
be taken as the holotype.

REPRESENTATIVE SPECIMENS. Yunnan: Rock 17321, 4746, 10737, 22852, 23080;
Ching 21392; Schneider 3459; Forrest 10967, 6403, 6568, 2763. S. W. Szechwan:
Rock 23992,

US.5.R. 7:

147. Delphinium cyananthum Nevski in Komarov, FI. ee ite

160. 725. 1937
Stems 7.5-10 dm. tall, glabrous, or slightly pilose at times below the
inflorescence; petioles glabrous except on the ciliate margins or —-
the lower to 15 cm. long; leaves round-cordate, the lower to 15 cm. b sie é
palmatisect almost to the base, the segments rhombic, on capiperetl se
2-3 cm., then biternately incised, glabrous above and ee edema
on veins beneath, ciliate-pilose on margins; raceme branched, dense, many
flowered, the flowers light blue; bracts to 1 cm. or longer, ss sale
pedicels commonly ca. 1 cm. long, adpressed to divaricate, densely pilose;

92 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

bracteoles linear-subulate or sometimes almost linear-lanceolate, densely
pilose, 1.5—5 mm. long, placed just below the flower; sepals strigulose and
pubescent, densely puberulent at tips, ovate, obtuse, 10-11 mm. long, 4-8
mm. wide, spur 11-13 mm. long, to 3 mm. thick near the base; petals pale,
the upper emarginate, almost straight, the lamina 9 mm. long, spur 13 mm.;
lower petal 2-cleft, bearded, the lamina oblong-rounded, 5 mm. long, claw
5 mm, long; stamens ciliate; carpels 3, densely pubescent.

Tyrer: Songoria, Tentek River, Schrenk (LE); seen.

A specimen from Irkutsk: distr. Balagansk, by Kryschtofowitsch in 1908 (uc)
seems to belong here.

148. Delphinium dasycarpum Stev. ex DC. Syst. 1: 547. 1817.
Fic. 13, &
Delphinium speciosum var. dasycarpum Rupr. Flora Caucausi 36, 287. 1869.

Stem 5-10 dm. tall, erect, striate, simple, pubescent; petioles pubescent,
not dilated at base; leaf blades pubescent, 5-lobed, the lobes lanceolate,
subtrifid, incised-dentate at apex, the upper leaves cuneate at base, the
lower subcordate; raceme simple, pubescent, scarcely elongate; flowers a
beautiful blue; axis and pedicels soft-pubescent; bracts one-third as long
as the pedicels, linear-subulate; bracteoles linear-subulate; sepals sub-
villous, 20-25 mm. long, 9-13 mm. wide, oval; spur straight, 18-20 mm.
long; petals dark, the 2 upper with emarginate lamina ca. 9 mm. long, spur

Q mm. or more; the lower deeply bifid, the ovate lamina 6 mm., the claw
4 mm. long; follicles 3, densely hairy,

Type: “hab. circa Nartsanam in Caucaso cisalpino,” Steven (G?), not
seen.

A specimen apparently belonging here is Caucasus, Prescott in 1851 (PHIL).

149. Delphinium davidii Franchet, Bull. Soc. Philom. Paris VIII. 5:
1893.

179. Fic. 13, I.
Delphinium longipes Franchet, Nouv. Arch. Mus. Paris II. 8: 191, 192. 1886,
not Moris, 1837.

Delphinium grandiflorum L. var. davidii (Franchet) Brihl in Brithl & King,

Ann. Bot. Gard. Calc. 5: 98. 1896,

Perennial from a slender rhizome; stem slender, 3-4 dm. tall, setulose
with spreading or appressed hairs, branching from base; basal leaves with
petioles to 2.5 dm. long, slender, sparsely pilose; blades 5-8 cm. in
diameter, pale above, glaucescent beneath, rounded-pentagonal in outline,
tripartite into segments cuneate and entire below, trifid to the middle with
oblong-ovate lobules to 15 mm. long and 5-8 mm. wide; cauline leaves
shorter petioled, the uppermost 3—5-parted into entire linear segments;
flowers solitary to few, corymbose-racemose; bracts like reduced leaves,
to 3 cm. long; pedicels 6-10 cm. long, retrorse-strigulose; bracteoles
divided to entire, 10-18 mm. long, placed well below the middle of the

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 93

pedicel; sepals dilute blue, strigose, with yellow spots at base on back;
the sepals 15—20 mm. long, ovate, 10-12 mm. wide, obtuse to rounded at
the tip, spur linear-subulate, decurved-spreading to slightly uncinate at
apex, 20-26 mm. long; lamina of upper petal somewhat hairy at apex,
slightly fuscous, almost straight, 9 mm. long, entire; that of lower petal
pale blue, oblong-ovate, short-bilobed, bearded, short-ciliate, 6 mm. long,
claw 5.5 mm. long; stamens glabrous; follicles 3, hirsute.

Type: Tibet, “Prov. de Moupin, in lapidosis,”’ David, July, 1869, (P);
seen. The above citation probably is in western Szechwan. A collection
by E. H. Wilson, 4690, from western Szechwan, seems to belong to this
species, which is unusual in having the bracteoles near the base of the
pedicels and was long thought to lack them.

150. Delphinium delavayi Franchet, Bull. Soc. Bot. France 33: 379.
1886. Fic. 13, J. K.

Delphinium delavayi var. acuminatum Franchet, Bull. Soc. Bot. France 33:
380. 1886. From meadows near Mt. Yengtze-hay, near Lankong, 3000 m.,
Delavay 1846 (p, K); seen. Leaves much incised.

Delphinium delavayi forma aureum W. T. Wang, Acta Bot. Sinica 10: 144.
1962. Isotype (A), seen, 7. T. Vii 12548.
Delphinium trichophorum var. brevistylum {. brevungue Lévl. Repert. Sp.

Nov. 7: 102. 1909. Type number from San Chan, Ten 579 (E); seen.

Perennial from a rather woody fibrous root system, the stem 5-10 dm.
tall, usually simple, leafy throughout, spreading-hirsute or -pubescent, or
the hairs more or less retrorse, many in the inflorescence enlarged and
yellow at the base; lower leaves with petioles 3-20 cm. long, dilated at
base, somewhat hairy, the blade divided almost to the base into 5 or 7
segments that are broadly cuneate, slightly incised or to about half way,
then coarsely ovate toothed or sometimes more pointed, the laminae 3-10
cm. wide, not as long, mostly quite pubescent, paler beneath than above;
upper leaves gradually reduced, shorter petioled, the uppermost often
trisect; raceme rather dense, often many flowered; lower bracts foliaceous,
trilobed, to 2 cm. long, the middle ones mostly lanceolate, entire, 1-2 cm.
long, the uppermost linear; pedicels ascending-erect, 1~—3 cm. long, hairy;
bracteoles linear to lanceolate, subopposite near the flower, 4-10 mm. long;
flowers mostly bluish violet to rich purple, pubescent; upper sepal broadly
ovate, 15 mm. by 10 mm., obtusely rounded at apex, the spur decurved,
20 mm. long, 3 mm. wide at base; lateral sepals elliptic-oblong, 15 mm. by
7 mm., truncate rounded at apex; lower sepals 14-16 mm. by 9 mm.,
ovate-oblong, obtuse; upper petals bluish, the lamina almost straight,
10-12 mm. long, entire, spur 16-18 mm. long; lower petals with a claw
6 mm. long, blade sparsely short bearded, elliptic-oblong, 10-11 mm. long,
cleft ca. 3 mm.; stamens subequal, subglabrous, 6-7 mm. long; anthers
dark, 1.5 mm, long; follicles 3, somewhat hairy, reticulate veined, erect
and closely parallel, 2-3 cm. long, 4-5 mm. wide, the style an additional

94 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

3-4 mm.; seeds brownish, 1.5 mm. long, densely covered with lamellate
scales.

Type: Yunnan, San Kong to the north of Tali, Nov. 8, 1883, Delavay
22, holotype (P), isotype (K), photo. (BH), all seen. Two other sheets
at p (Delavay 74 and 1847) are also labeled TYPE.

DisTRIBUTION. About 2700 to 4500 m., Yunnan and Szechwan and
eastern Tibet.

REPRESENTATIVE SPECIMENS SEEN. Yunnan: Maire 6367, 7449; Forrest 2861,
2963, 6319, 28587; Rock 4404, 5786, 5799, 5828, 5030, 4849, 5806, 5298, 5768,
6093, 5936, 6149, 9914, 6177, 9474, 18519, 5167, 10634, 11483; Schneider 2143,
3576: Ten 69; Vii 22461, 14367, 15543; Delavay 4109; Henry 9668A, 9668 ;
Ching 21456; Tsai 53738; Feng 2461, 217, 1697. Szechwan: Rock 18324, 24190;
Fang 2944; Peng 145; E. H. Wilson 3090.

/ 151. Delphinium drepanocentrum (Briihl) Munz, comb. nov.
Fro; 13, 4,
Delphinium altissimum var. drepanocentrum Briihl ex Huth, Bot. Jahrb. 20:
419 5 and subsp. drepanocentrum Brihl in Briihl & King, Ann. Bot.
Gard. Calc. 5: 101. 1896.
Delphinium umbrosum Hand.-Mazz. subsp. drepanocentrum (Briihl) Chowd-
hury ex Mukerjee, Bull. Bot. Surv. India 2: 293-5. 1961

Perennial from rather a woody base, 4-6 dm. high, the stem simple or
nearly so, spreading, and long- and soft-hairy with more or less brownish
hairs throughout, often with more or less purplish epidermis, leafy; lowest
leaves gone by anthesis, cauline with petioles 5-20 cm. long, somewhat
dilated at base, soft-hairy; blades 3- or mostly 5-lobed to 1-2 cm. from
the base, with rather long hairs above and on veins beneath, somewhat
pentagonal in outline, 5-10 cm. broad, with a rather deep basal sinus, the
lobes broad-rhombic, cuneate below, with rather shallow and ovate teeth
2-5 mm. deep and even broader, or sometimes shallowly 3-lobulate, then
toothed, the teeth mucronate; leaves gradually reduced upward, the uppet-
most becoming trifoliolate bracts 2-3 cm. long; inflorescence a secund,
5- to 20-flowered raceme 1-3 dm. long, more or less retrorse-hairy and
apparently glandular; upper bracts linear to lance-elliptic, 1-2 cm. long;
pedicels erect, 1-4 cm. ong; bracteoles oblong-linear, ca. 1 mm. wide,
7-8 mm, long, subtending the flower; sepals mauve to dull purple, with
rather coarse appressed hairs; upper sepal ovate, 12-13 mm. long, 8 mm.
wide, obtuse, the spur curved into a half circle or merely decurved-spread-
ing, 20-22 mm. long, 2.5 mm, wide at base, narrowed into a very slender
tip; lateral sepals elliptic-oblong, 12-13 mm. long, 5—6 mm. wide, rounded-
horror mt lower sepals asymmetrically oblong-elliptic, 10-11 mm.
sep fe wi es rounded at tip; petals bluish, the lamina of the upper
i oe i mm. long, 2.5 mm. wide, glabrous, pointed, subentire,
sae ne os = spur 20 mm. long; lower lamina oblong, 6.5 by 3 mm.,
reed ges ut 4 mm. into 2 linear-oblong, bearded lobes, the claw 5 mm.

8; Stamens ca. 5 mm. long, glabrous; anthers dark, 0.8 mm. long;

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 95

follicles 3, 15-18 mm. long, 3 mm. wide, somewhat hairy, the style 2 mm.
long; seeds obovoid angled, scarcely 1 mm. long, with transverse rows of
brownish scales.

Type: W. Sikkim, near Wallanchun, King’s collectors. Not seen.

RANGE. Growing on open and brushy slopes, 3500 m. to 5500 m., mostly
in Nepal.

REPRESENTATIVE SPECIMENS. Lall Dhwoj 0641, 0191; Stainton, July 27, 1956,
1142, 1144.

152. Delphinium georgei Comber, Notes Bot. Gard. Edinb. 18: 236.
Fic. 13, M.

Stem 9-13 dm. tall, erect, fistulose, angled, glabrous, purplish-glauces-
cent, 6 mm. thick at the base; basal leaves gone by anthesis, cauline
numerous, sparsely pubescent, yellowish above when dry, whitish beneath,
the lower blades to 8 cm. long, 15 cm. wide, 5-fid into rhombic-cuneate
segments, the interior 3-lobulate, the outer 2-lobulate, the lobules unequally
and coarsely crenate-dentate, apiculate; upper leaves smaller; lower cauline
petioles 12-16 cm. long, slightly vaginate; inflorescence lax, few branched
or simple, about 20-flowered and 3 dm. long, the slender axis glabrous,
striate; bracts linear-lanceolate, 5-10 mm. long; pedicels slender, 2-6 cm.
long, divergent; bracteoles 2, subulate, inconspicuous, subopposite, near
the summit of the pedicels, 2-3 mm. long and with a third subulate one
7-8 mm. long toward the base of the pedicel; upper sepal purplish blue,
strigose, 15 mm. long, 10 mm. wide, ovate, acutish, the spur 22 mm. long,
subulate, subfalcate, 2.5-3 mm. thick near the base; lateral sepals oblan-
ceolate, 17 mm. long, 5 mm. wide, pubescent only along the central nerve;
lower sepals subelliptic, 20 mm. long, 6 mm. wide, strigose; upper petals
glabrous, the blade 10 mm. long, slightly oblique, truncate, emarginate,
rarely sparsely setose, spur subfalcate; lower laminae oblique, 5 mm. by
3 mm., bilobed, with divergent lance-oblong lobes 3 mm. long, white-ciliate
at apex, yellow bearded at middle, claw 6 mm. long, sparsely ciliate,
appendaged; filaments 5-6 mm. long; anthers 1.5 mm. long; carpels 3,
glabrous or slightly setose.

Type: Yunnan, on the Tong Shan in the Yangtse Bend, at 3600 to 3900
m., Sept., 1913, G. Forrest 11353 (x), seen.

SPECIMENS SEEN, Kansu: Farrer 159. Yunnan: Yi 14490; Cavalerie 7871;
Maire 2128; K. M. Feng 2272, 2742; Rock 10539.

133. Delphinium giraldii Diels, Bot. Jahrb. 36. Beibl. 82: 3
Hand.-Mazz. Acta Hort. Gothob. 13: 54. 1939. y
Perennial, to 1 m. or so tall; stem glabrous; leaf blades to 22 cm. broad,
3-parted into subpetiolulate parts, the median part 2-3-lobed, the lateral
parts 2-lobed at 1-2.5 cm. above the base into irregularly pinnatifid-incised,
ovate or broadly rhombic-acute lobes, their ultimate divisions broadly to
narrowly ovate or subtriangular, acute, strigose above and beneath; lower

9. 1905;
Ic. 14, A.

96 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

petioles very long; inflorescence pyramidal with lax lateral racemes and
elongate many-flowered terminal raceme, glabrous; bracts and bracteoles
minute, lance-linear, 3-5 mm. long, somewhat strigulose, the bracteoles
near the flower; pedicels divaricate, rather short; sepals puberulous
without, apparently pale blue-violet; upper sepal round-ovate, 10 mm. by
9 mm., subapiculate, the spur straight, ca. 20 mm. long, 4 mm. wide at
base, subacute; lateral sepals elliptic, 9 mm. by 5 mm., rounded-truncate
at apex, puberulent especially along midrib and on margin; lower sepals
asymmetrically elliptic-oblanceolate, obtuse, 10 mm. by 4.5 mm.; petals
pale, the upper lamina scarcely oblique, shallowly 2-toothed, glabrous, 9
mm. long, the spur 20 mm. long; lower laminae oblong, oblique, 5 mm.
by 3 mm., divided over halfway into 2 lance-oblong, ciliate lobes, with a
small patch of beard near base of blade, claw 5 mm. long; stamens 5-6
mm. long, glabrous; anthers dark, oblong, 1 mm. long; follicles 3, glabrous;
seeds squamate.

Type: Shansi, mountains of Tsin-ling Shan, Giraldi (B), not seen. At
uc and at kK (in type cover) is probably authentic material from northern
Shansi (Schensi), Rev. Jos. Giraldi, Aug. 4, 1897, but without collection
number.

154. Delphinium glabricaule W. T. Wang, Acta Bot. Sinica 10: 154.
962. Fic. 14, B.
Stem 7 dm. tall, ca. 8 mm. in diameter at the base, glabrous, branched,
equally foliose; lower cauline leaves long petioled, the blades broadly
pentagonal-cordate, 5-6 cm. long, 7-8 cm. wide, trisect to base or 0.5-l
mm, above, the middle division sessile, broadly rhombic, cuneate, entire
below middle, trifid above into rhombic-ovate middle parts and oblique
lateral ones, ultimately divided into few-toothed lobules, loosely strigulose
on both sides along the nerves; the petioles 6-9 cm. long, glabrous, not
vaginate; racemes ca. 36 cm. long, 8-flowered, lax, with glabrous axis;
lower bracts lanceolate, ca. 4 cm. long, 5.5 mm. wide, others smaller,
linear; pedicels glabrous, 4-9 cm. long, somewhat arcuate curved, diver-
gent, bibracteolate above the middle; bracteoles 4—5 mm. long, linear;
sepals blue, broadly obovate, 11 mm. long, puberulent, the spur 18—20 mm.
long, subulate, spirally decurved at middle or below; upper petals blue,
the laminae glabrous, emarginate, slightly oblique, 11 mm. long; lower
petals blue, the laminae 6 mm. long, bifid into lance-triangular short-ciliate
lobes and yellow bearded at the base, claw 6 mm. long stamens glabrous;
carpels 3, glabrous.

Type: Szechwan, Hsio-chin, July 23, 1958, H. S. Chang & Y. H. Jen
6099 (PE), not seen; photo (£), seen.

I refer here, from Szechwan, Rock 16830 and 18121; from Yunnan, R. C.
Ching 2141.
Wang says that Delphinium glabricaule is near D. sung panense, but has

leaf segments not acuminate, the spur spirally decurved, and the filaments
glabrous.

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 97

155. Delphinium gyalanum Marquand & Airy-Shaw, Jour. Linn. Soc.
Bot. 48: 156. 1929. Fic. 14, C.
? D. dictyocarpum DC. subsp. tibeticum P. Briihl in Briihl & King, Ann. Bot.
Gard. Calc. 5: 100. 1896. Described as from “Southern Tibet, between the
northern frontier of Sikhim and Lhassa,” with two varieties glandulosum
and cimereum, but no collectors named. A specimen loaned me from
Gyangtse, July-Sept., 1904, Capt. H. J. Walton (cat), is labeled “var.
tibeticum” and seems to be glandular.

Stem erect, to 2 or 2.5 m. high, glabrescent, purplish, striate, more
pubescent and glandular in the inflorescence; lower petioles to 5 dm. long,
sparsely pubescent or villous, somewhat vaginate at base; lower laminae
subreniform in outline, 18-30 cm. in diameter, 5- or 7-lobed to the middle
or below, the lobes cuneate, 4-10 cm. wide, 3-lobulate with sharp angular
sinuses, then shallowly divided into acute uneven lanceolate teeth 1-5
mm. long and 1-4 mm. wide; upper leaves palmately incised-dentate,
short petioled, glabrescent except for the rows of short hairs along the veins

eneath; inflorescence loosely panicled, many flowered, to 3 dm. long, the
racemes lax, with spreading yellow glandular hairs and somewhat ap-
pressed, non-glandular white ones; bracts linear to lanceolate, entire, 1—2
cm. long, 1-4 mm. wide, pubescent; pedicels 3-8 cm. long, arched-
ascending; bracteoles near middle of pedicel, linear-oblong, 9-10 mm.
long; sepals deep blue, to violet or pale mauve, pubescent, the upper sepal
ovate, 16-17 mm. long, 9-10 mm. wide, subobtuse, the spur straight or
slightly decurved, 18-22 mm. long, 3 mm. wide at base, obtuse; lateral
sepals obovate, 19 mm. by 12 mm., with a truncate-obtuse prolongation
at apex; lower sepals similar, but more acute; upper petals bluish, the
limb lanceolate, shallowly bidentate, 10 mm. long, 4-5 mm. wide, the spur
15-16 mm. long; lower laminae ovate, 5 mm. by 3 mm., yellow bearded
at base, bifid into lanceolate hairy lobes 3 mm. long, the claw 5 mm. long;
stamens 5-6 mm. long; anthers dark, 1 mm. long; follicles 3, densely pilose
to subglabrous, 8-12 mm. by 4-5 mm., the styles an additional 4 mm.
long; seeds dark, angled-obovoid, squamate, ca. 1.2 mm. long

Type: Southeastern Tibet, in shady thickets on exposed flank of valley,
gorge of the Tsang-po near Gyala, 2700 m., July 21, 1924, F. Kingdon
Ward 5969 (x) seen; isotype (E), seen.

REPRESENTATIVE EXAMPLES: Southeastern Tibet: Ludlow, Sherriff & Taylor
5329, 5824, 5701, 4849, 6250: Kingdon Ward 12031; Ludlow & Sherriff 2777,
8789, 9893, 8781, 1942, 2461, 2421, 9632, 11108, 528; F. Spencer Chapman 187,
201; Hingston 367, 408: H. 7 W ‘alton 106; H. M. Stewart in 1907; L. A. Wad-
dell i in 1904.

156. Delphinium hirticaule Franchet, Jour. Bot. Morot grt he
ic. 14, D.

Delphinium hirticaule var. micranthum Finet & Gagnep. Bull. aati oF
France 51: 485. 1905. Based on Farges 630 bis, pro parte, hi coca
lower petal deeply divided.

98 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

regia

Ficure 14. Delphinium, Group VIII. Petals, the eed spurred, the sd a
clawed. A. D. giraldii; es of upper ose 9 mm. long, spur 20 mm.,; blade ©
lower petal 5 mm. long, w 5 wn from Giraldi | in 1897 (vc).

D.
glabricaule; blade of woper ge il at ong, spur mm.; blade of ee
petal 6.5 mm. long, claw

Be

.
blade of upper petal 11 mm. egg spur ade of lower petal 5 mm. long,
law 5 mm.; drawn from Ludlow, Mer. & Tayl r 5329 (BM). D. harte-
sonst blade 2 upper petal 11 mm. long, s

mm.; blade of lower petal 9

ong, claw 6 mm.; drawn from — 6367 (uc). ;

ot upper etal 14 mm. long, spur 20
ky .

long, claw 6 mm.; drawn from Knorring 368 Type (LE).
blade of upper petal 15 mm. long, spur 19 mm.; blade of lower petal 9 mm. lon

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 99

Delphinium coelestinum Franchet, Jour. Bot. Morot 8: 276. 1894. Type,
Farges 630 ter (Pp) with lower petals deeply divided. Seen.

Delphinium hirticaule var. coelestinum (Franchet) Finet & Gagnep. Bull. Soc.
Bot. France 51: 486. 1905.

Perennial from a fibrous root; stem simple or branched above, 3-10 dm.

ose
slightly dilated at the base; blades round-pentagonal in outline, 2— 8 cm.
in diameter, sparsely setulose on both surfaces, 5-parted to near the base
into cuneate-obovate segments to ca. 3 cm. wide, these 3- or more lobed or
toothed, the ultimate lobes lance-oblong to ovate, rounded, mostly 3-7 mm.
wide; flowers few in an open raceme, or with 1—2 lower branches; bracts
mostly simple, 8-12 mm. long, linear to lance-linear; pedicels subglabrous
or more or less setulose, mostly 1-4 cm. long, ascending; bracteoles linear
to lance-linear, to ca. 1 cm. long and placed near the flower; sepals deep
blue, with short stiff hairs; upper sepal ovate, 15-17 mm. long, 7-9 mm.
wide, subacute, the spur horizontal, somewhat decurved, 18-23 mm. long,
3 mm, wide at the base, gradually narrowed to a subacute point; lateral
sepals oblong-obovate, 15—20 mm. long, 8-10 mm. wide, rounded at apex,
with median line of hair; lower sepals 16-20 mm. long, 7-9 mm. wide,
obtuse, more generally pubescent; upper petals pale, the laminae almost
straight, entire, 11-13 mm. long, more or less setose at tip, the spur 10-20
mm. long; lower petals bluish, the blade oblique, rounded or deeply lobed
at the apex, 7 mm. by 5 mm., with a basal patch of bearding and ciliate
on margin; stamens 7-8 mm. long, glabrous; anthers dark, 1.5 mm. long;
follicles 3, sparsely pilose, 10-17 mm. long, 3 mm. wide, erect, the styles
an n additional 2.5 mm. long; seeds brown, 3-angled, farnellats on all sides,

. 1.8 mm. long.

Type: Szechwan, district of Tchen-kéou-tin, Farges 630 bis (P) seen,
some ae with this number having lower petals more deeply divided
than others; same true of specimens at (K). ;

SPECIMENS SEEN: Szechwan: Rock 18205; T. L. Tai 101803. Yunnan: Feng
2841; Maire 6367; Ching 21945; Rock 5167, 6305, 17317; Schneider 1867.
Much material that has been referred here belongs to D. ceratophorum.

Spur 12 mm.; blade of lower petal 4.5 mm. long, c pas
elie & Sherriff 894 (rz). L. D. malabaricum: blade of upper petal 10 mm.

pur 7 mm.; pane of lower petal 7 long, claw 5 mm.; ba
Santopan 11552 (ut MO). D, seagrass: S blade of upper petal 9 = ng,
spur 12 mm.; blade of ae petal 5 mm. long, claw 6.5 mm.,; drawn from
20783 (k).

100 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

157. Delphinium honanense W. T. Wang, Acta Bot. Sinica 10: 146.
1962.

Stems 5-6 dm. tall, 5-6 mm. thick at base, angled, glabrous, simple,
equally leafy; lower and middle cauline leaves with rather long petioles,
the blades 6—7 cm. long, 7-10 cm. wide, cordate, tripartite rather deeply,
the middle part broadly rhombic, acute or short-acuminate, entire in lower
half, above the middle unequally ovate-dentate or indistinctly 3-lobulate,
the lateral parts bifid to below the middle, the upper lobes much as in the
middle part, loosely strigose beneath or glabrescent, with stiffer hairs
above; petioles 1.5—2 times as long as blades, loosely hirsute, not vaginate;
upper leaves reduced; racemes 8-11 cm. long, ca. 10-flowered; lower bracts
trifid, others lance-linear to linear, 8-17 mm. long; pedicels 1—2.5 cm. long,
obliquely spreading, with white and yellow-glandular hairs; bracteoles
near the middle of the pedicels, linear, 5—8 mm. long, puberulent ; sepals
purple, elliptic- to narrow-ovate, 15-16 mm. long, loosely puberulent to
subglabrous; spur 20-21 mm. long, subulate, 3 mm. wide at base, slightly
decurved; upper petals yellowish, glabrous, emarginate or bilobulate; lower
petals blue, the limb bifid into obtuse triangular lobes to beyond the
middle, ciliate, yellow bearded near base, claw subequal, short-appendic-
ulate; stamens glabrous; carpels 3, glabrous.

Type: Honan, Sichia, May 14, 1959, Honan Exped. no. 534 (PE), not
seen; photo. (£), seen.

Said to be near D. hirticaule, but stems glabrous, parts of leaves less
divided. I have seen no material I could refer here.

158. Delphinium kingianum Briihl ex Huth, Bot. Jahrb. 20: 469.
1895; Brihl, Ann. Bot. Gard. Calc. 5: 108. 1896, pro synon.
Fic. 14, E.
Delphinium pachycentrum ssp. tsangensis vars. dasycarpa and lasiocarpa Brihl,
loc. cit. 108 — according to Wang, Acta Bot. Sinica 10: 138. 1962
Delphinium kingianum var. leiocarpum Brihl ex Huth, loc. cit.
? Delphinium kingianum var. eglandulosum Wang, loc. cit.

From a thick central fibrous root, stems in clumps, 3—4 dm. tall, 3-5
mm. thick, subglabrous below, densely spreading-pubescent upward with
yellow glandular hairs, densely leafy, the cauline leaves 3-10 cm. across,
subglabrous above, paler and more strigose beneath, palmatifid almost to
base into narrow-cuneate segments which are sharply lobed about half
way, then with sharp lanceolate teeth 2-10 mm. long, 1-3 mm. wide;
petioles slender, 2~5 cm, long, scarcely widened at base; inflorescence 4
crowded many flowered raceme to ca. 1 dm. long, densely glandular-
pubescent; lower bracts reduced leaves, upper lance-linear, 6-10 mm. long;
pedicels 1-4 cm. long, arched-ascending; bracteoles lance-linear, acuminate,
5-6 mm. long, subtending the flower; sepals purplish blue, white-villous,
the upper lance-ovate, 20 mm. by 10 mm., rounded-obtuse, the spur 20-22
mm. long, 3 mm. wide at base, quite sharply pointed; lateral sepals oblong,

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 101

22 mm. by 8 mm., rounded at tip; lower sepals oblong, 22 mm. by 7 mm.,
narrower at tip; upper petals bluish, the laminae almost straight, 14 mm.
long, 3 mm. wide, entire, rounded at tip, the spur 19-20 mm. long; lower
petals with an obovate lamina 12 mm. by 8 mm., deeply emarginate,
bearded at base, sparsely so above, claw 5 mm. long, appendiculate:
stamens 6-7 mm. long; carpels 5, pubescent to almost glabrous.

Type: Between the northern frontier of Sikkim and Lhassa, King’s
collector. Not seen.

REPRESENTATIVE SPECIMENS: Tibet: Ludlow & Sherriff 527, 855; B. J. Gould
508; L. A. Waddell, Sept. 5, 1904.

159. Delphinium knorringianum B. Fedtsch. Jour. Roy. Hort. Soc.
61: 196, 197, fig. 43. 1936. Fic. 14, F.

Perennial from an elongate root, the stems ascending to suberect, 3.5—4.5
dm. tall, more or less branched above, with short stiff spreading hairs below
and glandular in the inflorescence; leaves mostly basal, with petioles 8-12
cm. long, slightly dilated below; blades 4-5 cm. broad, more or less pen-
tagonal, 5-parted, the parts cuneate toward base, indistinctly 5-lobed at
apex, the lobes oblong-ovate, more or less rounded at tip, to 7 mm. wide;
stem leaves reduced, the upper subsessile, with entire broadly linear lobes;
inflorescence with one to few spreading branches and bristly glandular-
hairy, each branch subcorymbose, with 3-5 blue-purple flowers; lower
bracts 3—5-parted; pedicels 3—7 cm. long, divaricate; bracteoles 2, linear,
ca. 4 mm. long, near the middle of the pedicel; upper sepal subglabrous to
glandular-pubescent, ovate, 17-18 mm. long, 11 mm. wide, obtuse, the
spur straight, 21-22 mm. long, bilobed at tip, the lobes divergent; lateral
sepals broadly ovate, 22 mm. by 12 mm., pubescent; lower sepals pubes-
cent, 22 mm. by 12 mm., ovate; petals pale, blue purple, the upper with
elongate-elliptic limb, clawed, bidentate, glabrous, ca. 14 mm. long, the
spur 19-20 mm. long; lower petals with elongate-linear limb, rounded at
top, bidentate, 6 mm. long, narrowed gradually into a claw 7 mm. long;
follicles 2-3, densely pubescent.

TYPE: Asia Media, Kirghistan, Osh district, sources of Akhbura River
(Alai Range), July, 1913, O. E. Knorring 368 (LE); seen. Illustrated in
Original description. No other material seen.

160. Delphinium lancisepalum Hand.-Mazz. Acta Horti Gothob. 13:
55. fig. 2a. 1939. Fic. 14, G.
Delphinium pachycentrum Hemsl. var. lancisepalum (Hand.-Mazz. ) W. cE:

Wang, Acta Bot. Sinica 10: 140. 1962 and var. pseudo-lancisepalum, Wang,

loc. cit.

Stem 2-5 dm. tall, 3-4 mm. thick, simple or with few short sterile
branches, white-strigulose, densely and equally leafy to inflorescence;
petioles mostly 3-6 cm. long, slender, more or less strigulose, dilated at

ase; blades cordate-orbicular in outline, mostly 4-9 cm. in diameter,

102 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

minutely strigulose on both the upper green and lower pale surfaces, trifid
almost to base, the 2 lateral segments deeply bifid, the 5 parts thus created
rhombic-lanceolate to -ovate, trilobed to about the middle then deeply
toothed, the teeth lance-oblong to ovate, 2—4 mm. wide, acute; inflorescence
a dense raceme, 5-12-flowered, with dense retrorse pubescence; lower
bracts leafy, upper linear, filiform, 1—1.2 cm. long; pedicels 1.5-5 cm.
long, almost erect; bracteoles filiform, 10-12 mm. long, next to the flower;
sepals blue to pinkish, somewhat strigulose and ciliate, the upper lance-
ovate, 19-20 mm. long, 8 mm. wide, acuminate, the spur almost straight,
20-23 mm, long; lateral sepals lance-ovate, 20-22 mm. long, 8 mm. wide,
acute; lower sepals with about the same dimension, more acuminate; upper
petals deep violet, the laminae almost straight, deeply retuse, 15 mm. long,
glabrous, the spur 19 mm. long; lower petals with laminae 9 mm. long,
7.5 mm. wide, with a small basal patch of bristly hairs and 2 gaping oblong
lobes 4-5 mm. long, claw 6.5 mm, long; stamens 7-8 mm. long, quite broad
in lower part, glabrous or ciliate; anthers dark, oblong, 1.2—1.4 mm. long;
follicles 3, hairy.

Type: Sikang, district Kangting (Tachienlu), Ta-pao-shan, 4500 m.,
Aug. 20, 1934, Harry Smith 11248 (w?), not seen; isotypes (A, BM, MO,
NY), seen.

COLLECTIONS SEEN: Szechwan: Legendre 1563; H. Li 78175.

161. Delphinium lasiocarpum Tamura, Acta Phytotax. & Geobot.
Kyoto 15: 196. 1954

Stem striate, sparsely recurved-pilose or subglabrous; cauline leaves
short petioled, the blades reniform, 6 cm. long, 12 cm. wide, curved-pilose
on veins of both sides, tripartite, the middle segment pinnatifid, the lateral
bifid, the lobes unequally pinnatifid and the lower incised; inflorescence
many flowered, paniculately branched, yellow-villous above; lower bracts
3-parted or trifid, the upper linear; pedicels to 5 cm. long, with opposite
bracteoles above the middle; flowers blue, 2.5 cm. in diameter, the spur
longer than the sepals, to 20 mm. long, 5 mm. wide at base, slightly
curved; upper petals glabrous, 25 mm. long, with incurved spur 7-8 mm.
long; lower petals bifid or lobed, white bearded, the lobes acute or obtuse;

follicles 3, villous, 10 mm. long without the style, this 4-5 mm. long; seeds
smooth, winged. :

Type: Tibet near Lhasa, Panchogan, 103900 (kyo); not seen. I have

been unable to borrow any material of this species, so can offer only a

translation of the original description. The author thinks it related to
Delphinium altissimum

162. Delphinium laxicymosum W. T. Wang, Acta Bot. Sinica 10: 148.
noe Fic. 14, H.

as 5-7.5 dm. tall, ca. 4 mm. thick at base, loosely white-hirsute below,
glabrous above, branched, equally foliose; lower cauline and middle leaves

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 103

longish petioled, the blades broadly pentagonal, 4-8 cm. long, 6-12.5 cm.
wide, cordate, 3-parted almost to base, the middle part rhombic, the lateral
obliquely triangular-ovate, all lobulate and toothed, loosely strigulose on
both surfaces; petioles 1.5 to 2 times the laminae, loosely hirsute; upper
cauline leaves reduced; flowers in 3—5-flowered corymbiform clusters; lower
bracts to 3 cm. long, trifid to entire, upper subfiliform; pedicels 2~7 cm.
long, glabrous, divergent; bracteoles near the middle of the pedicel,
filiform, 2—2.5 mm. long, 0.2 mm. wide; sepals intense blue, elliptic-obovate
or elliptic, 14 mm. long, puberulent, the spur 23-25 mm. long, subulate,
slightly to much decurved; upper petals yellowish, 11 mm. long; lower
blue, 6 mm. long, bifid to slightly below the middle into lance-oblong obtuse
lobes, long-ciliate, yellow bearded near base, claw 6 mm. long, short-
appendiculate; stamens glabrous; carpels 3, glabrous.

Type: Szechwan, north of Wen-chuan Hsien, 3100 m., July 20, 1930,
F. T. Wang 21767 isotype (a), seen.

Another collection probably to be referred here is Ts¢-Kou, Abbé Monbeig

oy .

Wang said his proposed species, Delphinium laxicymosum, is near D.
potaninii, but with stem and petioles hirsute and inflorescence more cymose.

163. Delphinium laxiflorum DC. Syst. 1: 360. 1817. Fic. 14, I.

Delphinium villosum Stev. ex DC. var. laxiflorum (DC.) Huth, Bot. Jahrb.
20: 412. 1895.

? Delphinium elatum var. anomalum Kunth ex Huth, loc. cit. 399. From the
Altai Mts. and with light colored petals, hence not D. elatum.

Delphinium amoenum Stev. ex DC. Syst. 1: 546. 1817. Placed in synonymy
here by Nevski in Komarov, Fl. U.S.S.R. 7: 158. 1937. I have not seen
authentic material.

Said to be vegetatively like Delphinium dyctiocarpum DC. (Group IX),
i.e. 6-15 dm. tall, robust, quite glabrous, large leaved, the lower petioles
to 2 dm. long; leaf blades rounded, pentagonal in outline, the upper ca. 4
cm., the lower 15 cm. broad, 3-, 5-, or 7-parted to within 1 or 2 cm. of the
base, the parts not deeply laciniate; inflorescence a dense raceme or open
panicle, many flowered; bracts mostly entire; pedicels 1—3 cm. long; brac-
teoles linear, 3-7 mm. long, well below the flower; sepals violet-blue,
glabrous, 10-12 mm. long, spur 12-20 mm. long; petals blue, or the upper
whitish, the upper erose or scarcely bidentate, the laminae 7 mm. long,
spur 10 mm.; lower petals pale blue, the laminae 4 mm. long, 2-lobed,
bearded, the claw 4 mm. long; ovary apparently glabrous to pubescent.

Type: “Hab. in Sibiria (hortul.)” (G? not seen). A specimen loaned
from Leningrad as “D. dictyocarpum D. Altai, 1826, Herb. Ledebour” has
blue petals and seems to be authentic material of D. laxiflorum

Some other specimens apparently referable here are all Si
differ from D. dyctiocarpum in not having almost black petals.

Siberia: Semipalatinsk, Krylov & Sergievskaya in 1929 (ny); Karelin & Kiri-

berian and

104 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

loft in 1840 (mo). Altai borealis, Zrastova in 1909 (pm); V. Burdakova in 1931
(ny); Chaffanjon 1019 (P).

164. Delphinium lilacinum Hand.-Mazz. Symb. Sinicae 7: 277. 1931.
Delphinium handelianum W.T. Wang, Acta Phytotax. Sinica 6: 365. 1957.

Rhizome very short, the roots slender; stem 5—6 dm. tall, striate, hairy,
equally leafy, weakly branched almost from base; leaves pentagonal, 7-12
cm, in diameter, 5-fid to three-fourths or seven-eighths their length, the
divisions rhombic-ovate, the margins rarely contiguous, deeply incised, then
coarsely crenate, green on both surfaces, sparsely strigose above, reticulate
veined beneath; petioles one and one-half times as long as the middle
laminae, the upper half as long, more or less vaginate, sparsely hirsute;
racemes terminal on stems and branches, to 14-flowered, densely and
retrorsely white-strigulose and somewhat yellow-hirsute; lower bract
3-parted, others entire, upper subulate-lanceolate, half as long as pedicels
or the uppermost as long; pedicels divergent, 1-5 cm. long; bracteoles in
upper fourth of pedicel, lance-linear, ca. 5 mm. long, ciliate; sepals lilac,
broadly ovate, rounded at apex, sinuate-attenuate, white strigulose and
yellow-hispid, densely ciliolate; spur about twice as long as blade, attenu-
ate; petals about equal to sepals, the upper 3 mm. wide, with 2 linear
lobes 1.5 mm. long; lower petals with lamina equaling claw, 3 mm. wide,
cut into 2 narrow lobes for two-thirds its length, ciliate and long white-
bearded; stamens 6 mm. long; anthers brown, oblong, 1 mm. long; carpels
3, sparsely white-strigose.

ae Beyendjing (Peyentsin), Yunnan, forests near Guti, Aug. 29,
vt eat 1292 (PEP); not seen. The only collection that I have seen
which seems to me possibly representing this species is Cavalierie 2936

(£), also from Yunnan. This proposed species, Delphinium lilacinum, may
be too close to D. pogonanthum.

165. Delphinium lipskii Korsh. Bull. Acad. Pétersb. V. 9: 402. 1898.
: Fic. 14, J.
Rhizome woody, elongate; stems erect, 6-7.5 dm. tall, cinereous-
pubescent below, glabrous in the inflorescence, openly branched above;
leaves grayish pubescent beneath, glabrous above, 4-5 cm. wide, the lower
hae reniform, trisect almost to the base, the segments peeile. the
ae pre hovente to base, all segments 3-lobed into entire or incised
ese oblong, 4-10 mm. long, 3-6 mm. wide); parts of the upper
eaves anceolate, entire; bracts all entire, lance-linear, 4-5 mm. long;
bie glabrous, 3—4.5 cm. long; bracteoles 2-3, sninuite, at middle of
tay sl age upper sepal ca. 14 mm. by 7 mm., apiculate, intense blue,
coc “ ong, 3.5 mm. wide at base, spreading, subcylindric, blunt;
~ sepals oblong-elliptic, 14 mm. by 8 mm., rounded; lower sepals 14
mm. by 6 mm., apiculate; upper petals pale blue, oblique; “the laminae

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 105

ca. 13 mm. by 4 mm., shallowly incised, the spur ca. 19 mm. long; lower
petals blue, the round-obovate laminae 6 mm. long, 4 mm. wide, deeply
incised, white-hairy, the claw 5 mm. long; stamens 7-8 mm. long, ciliate;
anthers yellow, 1 mm. long; follicles 3, hirsute, 14-20 mm. long, with blue
styles an additional 2 mm. long.

Type: Buchara (Bokhara), Darwaz, near Chumbau River, between
Sary-dasch and Kala-i-chum, 1300 to 1600 m., June, 1897, Korshinsky
(LE), seen; photo. (A), seen.

166. Delphinium ludlowii Munz, sp. nov. Fic. 14, K.

Delphinium ludlowianum Chowdhury ex Mukerjee, Bull. Bot. Surv. India 2:
—296. 1960; nomen.

About 1 m. tall, the stems sparingly and minutely strigulose, more
densely so in the inflorescence, rather slender, remotely leafy; cauline
leaves pentagonal-reniform in outline, green above, paler beneath, stri-
gulose on both surfaces, to about 12 cm. broad, 5-fid to about 1 cm. from
base, the segments cuneate-rhombic, incised about halfway to the midrib
into 3 broad lobes, these shallowly and evenly rather coarsely toothed, the
teeth lance-oblong, mostly 3-5 mm. long, 2-3 mm. wide, mucronate;
petioles to ca. 12 cm. long, slender, subglabrous, slightly vaginate at base;
leaves gradually reduced toward the inflorescence into leafy bracts; inflor-
escence mostly a lax 6—12-flowered raceme 1-2.5 dm. long, closely
strigulose; upper bracts petiolate, lanceolate, 1-2.5 cm. long, 2-6 mm.
wide; pedicels arched, 1-5 cm. long; bractlets subtending the flower,
subspatulate, 4-6 mm. long; sepals rich purple to pale maroon, minutely
strigulose; the upper sepal oblong-ovate, 13-15 mm, long, 11-12 mm.
wide, with an extended blunt tip, the spur spreading, 12-15 mm. long,
4.5 mm. wide at base, rather abruptly narrowed to an acute tip; lateral
sepals oblong-obovate, 12 mm. by 8 mm., subtruncate; lower sepals 13
mm. by 6—7 mm., rather pointed; petals bluish, the upper laminae strongly
oblique, 10 mm, long, 2.5 mm. wide, glabrous, bidentate, the spur 1 1~13
mm. long; lower laminae oblique, oblong, 5 mm. by 2.5 mm., divided
almost halfway into 2 oblong lobes, subglabrous except near the central
base, the claw 5 mm, long; stamens 5—6 mm. long, glabrous; anthers dark,
oblong, 1 mm, long; follicles 3, divergent, glabrous, 12-13 mm. by 3 mm.,
the styles an additional 2.5 mm. long; seeds angled. ve

Caules ca. 1 m. alti, minute strigulosi, tenues, remote foliosi; folia
caulium pentagono-reniformia, supra viridia, infra pallida, 6-12 cm. lata,
S-fida, segmentibus cuneato-rhombeis, incisis, dentatis; petioli 6—-12 cm.
longi, tenues, subglabri; inflorescentia cum 6~12 floribus, laxa, 1-2.5 dm.
longa, strigulosa; bracteae superiores petiolatae, lanceolatae, 10-25 mm.
longae, 2~6 mm. latae; pedicelli 1-5 cm. longi; bracteolae subter florem,
subspatulatae, 4-6 mm. longae; sepala purpurea vel purpureo-rubra,
strigulosa, 12-13 mm. longa, 4-7 mm. lata, calcar 12-15 mm. longum,
base 4.5 mm. latum; petala subcaerulea, laminae superiores obliquae, 10

106 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

mm. longae, 2.5 mm. latae, glabrae, bidentatae; laminae inferiores ob-
liquae, oblongae, 5 mm. longae, 2.5 mm. latae, bilobatae, base barbatae,
unguis 5 mm. longus; stamina glabra; antherae oblongae, 1 mm. longae;
folliculi 3, divergentes, glabri, 12-13 mm. longi; styli 2.5 mm. longi; semina
angulata.

Type: Northeastern Bhutan, Trashiyangsi, 3300 m., Aug. 28, 1934,
among stones in dense jungle, Ludlow & Sherriff 894 (BM); isotype, (E).
OTHER NUMBERS SEEN. Bhutan: Singhi Dzong, Khoma Chu, Ludlow & Sher-
riff 470 (pm, £); Me La, Ludlow & Sherriff 368 (pm, ©); Trashiyangsi, Ludlow
& Sherriff 1008 (pm); Singhi Kusted, Cooper for Bulley, 4322 (pm); Bamthang,

Ludlow & Sherriff 16972 (sm); Rudo La, Ludlow & Sherriff 20978 (sm); Dur
Chu, Ludlow & Sherriff 19485 (Bm).

One of my great pleasures when working at the British Museum of
Natural History in 1965 was to become acquainted with Mr. Frank
Ludlow, whose extensive collection from the Himalaya and Tibet have
been of great value in this study, and I am delighted to name this species
for him! Delphinium ludlowii seems close to D. altissimum, but has more
evenly toothed leaves, bracteoles nearer the flower, and sepals more
minutely pubescent.

Instead of using the name Delphinium ludlowianum which Chowdhury
wrote on some herbarium sheets in Great Britain, and which Mukerjee
listed in the Bull. Bot. Surv. India 2: 293-6. 1960, without description,
I feel that I should employ a different epithet. I have had no contact with
Dr. Chowdhury to learn whether he wished his name published, and have
been unable to ascertain his whereabouts. I am, therefore, using the name
Delphinium ludlowii for my new species.

167. Delphinium malabaricum (Huth) Munz, comb. nov. Fic. 14, L.

es sca dasycaulon Fresen. var. malabaricum Huth, Bot. Jahrb. 20: 435.

Perennial from a woody cylindrical rhizome; stem 3-10 dm. tall, rather
slender, glabrous to retrorse-pubescent below, strigose in the inflorescence,
a0 to few branched above, remotely few leaved; lower petioles 1-2-5
- long, glabrous to somewhat pubescent, strongly vaginate at base, the
ew cauline petioles reduced, the upper very short; lower leaf blades more
or less reniform in outline, 3-10 cm. broad, glabrous or nearly so above,
nr few to many stiff hairs on the margin, and along veins beneath, 5- or
sor Bae 3 or 1/2 their length, the broad cuneate-obovate segments

y bifid, then coarsely toothed with broad entire or toothed teeth with
mucronate tips, middle cauline leaves more deeply divided, the segments
entire to 3-toothed or more; inflorescence a series of 1 to few rather dense,
oe to many flowered racemes; lower bracts leafy, trifid, to 2 or 4 cm.
‘ioe upper neat, entire, 6-10 mm. long; pedicels divergent, 5-20 mm.
ong, strigose; bracteoles subulate, near middle or base of pedicel; sepals
bright blue, with a white subterminal spot, strigose to subglabrous: upper

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 107

sepal ovate, 10-14 mm. long, 5—7 mm. wide, the spur subconic, 7-9 mm.
long, 3-4 mm. wide at base, rather obtuse; lateral sepals elliptic-obovate,
truncate-rounded at apex, 13-16 mm. long, 7-9 mm. wide; lower sepals
narrowly obovate, 11-13 mm. long, 5-7 mm. wide; upper petals pale, the
laminae glabrous, slightly oblique, ca. 10 mm. long, with 2 terminal round-
tipped lobes ca. 2.5 mm. long, the spur 7 mm. long; lower laminae blue,
round-oblong, 7 mm. by 5 mm., divided to near the base, short-ciliate and
strigulose, the claw 5 mm. long; stamens 6—7 mm. long, ciliate; anthers
bluish, almost round, less than 1 mm. long; follicles 3, strigulose, divergent,
perhaps 10 mm. long, the styles an additional 3 mm.

Type: The only collection cited by Huth which is, therefore, the type,
is, E. India, Malabar Concan, Stocks, cited from G, B, W, LE . I have seen
none of these, but have seen the collections from GH and Pp. I suggest
the Geneva specimen as the holotype, the others as isotypes.

Apparently a species of the Western Ghats in India and quite distinct
from the Ethiopian Delphinium dasycaulon in its non villous stems, longer
bracts, glabrous upper petals, etc.

MATERIAL SEEN. Khandala, Santapau 11551 and 11552 (mo); Purandhar,
Santapau 13868 and 13869 (mo); Iooner, Deccan, Stocks (K); Bombay, Dalzell
(K); Ghanta (x).

168. Delphinium micropetalum Finet & Gagnep. Bull. Soc. Bot.
France 51: 479. 1905. Fic. 14,

Delphinium micropetalum subsp. confusum Choudhury ex Mukerjee, Bull.
Bot. Surv. India 2: 293-296. 1960; nomen.

Delphinium jugorum Hand.-Mazz. Symb. Sinicae 7: 273. 1931. Based on
Handel-Mazzetti 9712 from Mekong-Salwin Divide, n.w. Yunnan (w). Not
seen, but the published figures are like those of D. micropetalum. Put into
synonymy also by W. T. Wang, Acta Bot. Sinica 10: 142. 1962.

Stem striate, 4.5-10 dm. tall, simple or sparsely branched above, thick,
retrorsely hirtellous, with 3-4 equally spaced leaves, or leafless above;
leaves suborbicular in outline, rather stiff-strigose above and beneath, the
lower cauline the largest, 11-14 cm. in diameter, trifid to 1 cm. from the
base, all parts broadly rhombic-obovate, obtuse, trifid, coarsely and irreg-
ularly narrowly cuneate, dentate and incised, the teeth subovate, 1-5 mm.
long and almost as wide; lower petioles 3-4 times as long as the blades,
vaginate; racemes densely 8—13-flowered, those on the branches reduced,
the axis white-strigose and with yellow glandular hairs; lower bracts
foliaceous, petioled, dilated at base, 3—5-lobed, 1-2 cm. long; the others
oblong-lanceolate, entire; pedicels 1.5-5 cm. long, divergent; bracteoles
puberulent, lance-linear, 10-12 mm. long, just below the flower; sepals
intensely blue, upper sepal ovate, 14 mm. by 8 mm., the spur 12-14 mm.
long, 3.5 mm. wide at the base; lateral sepals oblong, 16 mm. by 8 mm.,
subtruncate; lower sepals asymmetrically obovate, 18 mm. by 12 mm.,
broad at apex; lamina of upper petals yellowish, at right angles to the

108 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

spur, 9 mm. long, 3.5 mm. wide, obtuse or emarginate, the spur 12 mm.
long; lower laminae 5 mm. by 3 mm., oblique, oblong, bifid one-fourth
its length, the lobes obtuse, short hairy below the middle, claw 6.5 mm.
long; stamens 6-7 mm. long, glabrous; anthers dark, oblong; carpels 3,
puberulent, to 10 mm. long, the style an additional 3 mm.
Type: Western Yunnan, Tsé-Kou, April 1895, Soulié 1098 (Pp); seen.
SPECIMENS SEEN. 3300-4400 m. Southeastern Tibet: Rock 22401, 22402, 22373.

Yunnan: Yii 20783, 20639, 20369, 22320; Forrest 221; Tsai 58069.
Chawchi Pass, Farrer 1867, 1156; Forrest 27516, 27225, 25091; Lepcha 865.

169. Delphinium mitzugense Ulbr. Notizbl. Bot. Gart. Berlin 12: 358.

Fic. 15, A.

Delphinium taliense Franch. var. glabrum W. T. Wang, Acta Bot. Sinica 10:
159. 1962. Based on same type as D. mitzugense.

? Delphinium taliense var. pubipes Wang, loc. cit. I have not seen cited
material.

Stem 10-12 dm. high, erect, stout, glabrous and glaucous below with
some red, more or less bluish and glaucous above, branched above, with
few remote leaves; basal leaves gone by anthesis, the lower cauline with
petioles 1.5-2 dm. long, stout, glabrous, vaginate, the upper shorter and
more slender; blades rounded in outline, the lower 10-13 cm. in diameter,
scattered-strigose above and beneath, paler beneath, palmately 7-fid to 1
cm. or less from the base, the segments subcuneate-obovate, mostly 3-lobed
distally, each lobe divided into 2-3 smaller, oblong, acute to obtuse
mucronulate divisions 2-5 mm. wide, the uppermost leaves divided into
5 lanceolate entire segments; inflorescence few branched, paniculate,
glabrous, lax; bracts subulate, 8-15 mm. long, bluish; pedicels slender,
ascending, 2~5 cm. long, blue; bracteoles subulate, 5—8 mm. long, placed
near the middle of the pedicel; sepals blue, somewhat strigulose; the
upper sepal ovate, 13-15 mm. long, 7 mm. wide, obtuse, the spur horizon-
tal-decurved, 20-25 mm. long, 4 mm. wide at base, subcylindric, the tip
cao ES mm. wide; lateral sepals 15 mm. long, 8 mm. wide, rounded-trun-
cate, strigose on median nerve; lower sepals subelliptic, 14 mm. long, 6
mm. wide, obtuse; petals bluish, the upper laminae slightly oblique,
spatulate, erose, 9 mm. long, the spur 20 mm. long; lower laminae oblique,
oblong-obovate, 7-8 mm. long, deeply cleft into lance-oblong lobes, yellow-
bearded at base of lamina, the remainder with long white hairs and ciliate,
claw 5 mm. long; stamens 4-6 mm. long, moderately dilated below,
glabrous; anthers bluish, 1 mm. long; carpels 3, pilose

Type: Southwestern Szechwan, Muli, Mt. Mitzuga, 4300 m., J. F. Rock
18317, holotype (B?), not seen; isotypes (GH, Us) " seen

_Another collection is Szechwan, Huei-li Hsien, T. a. Vii 1598 (GH).
Since Wang reduced D. mitzugense to varietal status under D. taliense, he
must have considered the two very close, but the latter has a much wider
lamina of the lower petal, and an emarginate upper petal.

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 109

170. Delphinium muliense W. T. Wang, Acta Phytotax. Sinica 6:
365.: 3957. Fic. 15, B.
Delphinium pachycentrum Hemsl. var. tenuicaule Chen, Bull. Fan Mem. Inst.
Biol. Peiping (n.s.) 1: 168. 1948. Type, T. 7. Vii 6989, 3300 m., Muli,
Kulu (PE?); not seen. Reduced to synonymy of his new species (D. mu-
liense) by W. T. Wang, Acta Bot. Sinica 10: 149. 1962,
Delphinium muliense var. minutibracteolatum W. T. Wang, Acta Bot. Sinica
10: 149. 1962. Based on T. T. Vii 14371, which I have not seen, but the
separation because of reduced bracteoles hardly seems well founded.

Perennial, 6-7 dm. tall, striate, subglabrous, slender, simple or with
short sterile branches, equably leafy; basal leaves lacking at anthesis;
cauline laminae pentagonal, cordate, 2.5—5.5 cm. long, 4-7 cm. wide, 3-
sect, the central segment rhombic, petiolulate, trifid to its middle, lateral
segments 2-parted, bifid to trifid, all parts incised into dentate lobes,
strigose above, pubescent beneath on nerves; petioles 4-15 cm. long,
glabrous, somewhat vaginate; raceme few to many flowered, 6-20 cm.
long, retrorsely white-hairy and yellow-glandular; lower bracts leaflike,
but smaller, upper lanceolate to linear, 8-14 mm. long; pedicels erect-
spreading, 1-5 cm. long; bracteoles in upper third of pedicel, lanceolate
to linear, 5-13 mm. long, 1-2 mm. wide; flowers bluish-purple to yellow-
ish, the upper sepal obovate, cymbiform, 12-15 mm. long, the spur some-
what decurved, 20-25 mm. long; other sepals oblong-obovate, obtuse,
14-18 mm. long; petals bluish, the upper laminae ca. 8 mm. long, obliquely
truncate, glabrous to pubescent, entire, spur 15-20 mm. long; lower
laminae 5-6 mm. long, ovate, yellow bearded near base, bifid to below
the middle with oblong triangular lobes strigose and ciliate, claw 4.5—5.5
mm. long, appendiculate; stamens glabrous; carpels puberulent on sutures.

Type: Szechwan, Muli, Ha-lin, 3500 m., July 11, 1937, T. T. Yu 6989?,
published as 6988 (PE, A), photo. (A); photo. and isotype seen.

Exampies. Seemingly agreeing with the type, Muli, Rock 16916. Tibet:
Yigrong Gorge, 3300 m., Kingdon Ward 12190; Kongbo Prov., Ludlow, Sherriff
& Taylor 6920; Tsanpo Valley, Ludlow, Sherrif & Taylor 5397; Lochen La,
Kingdon Ward 12240; Shweli-Salwin Divide, Forrest 11792.

171. Delphinium orthocentrum Franchet, Bull. Soc. Philom. Paris

VIII. 5: 178. 1893. Fic. 15, C.

Delphinium szechuanicum Ulbr. Repert. Sp. Nov. 14: 298. 1916. Type, West
Szechwan, E. H. Wilson 1088 (B?) not seen; isotypes (GH, US), seen.

Perennial from a woody root, the stem glabrous below, erect, simple,
leafy throughout, 5-10 dm. tall, grayish in inflorescence with appressed
short hairs; petioles to 1 dm. or more long, somewhat dilated at base ;
laminae of leaves somewhat strigulose on both sides, 1 dm. or more in
diameter, deeply 5-parted, the segments broadly obovate-rhombic, sete
at base, unequally incised into acute lobes, the terminal one elongate;
inflorescence a rather crowded raceme with perhaps 8-20 flowers; bracts

110 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Ficure 15. Delphinium, Group VIII.
; Petals, the upper spu og ce lower
wed. A. D. mitzugense; blade of upper petal 9mm = wate 20 : blade
mm. :

tal 12 mm. long, spur 15 mm.; blade o
, f lower petal 10 mm. long, claw 6 mm.;
ooeete from Ludlow & Sherriff 3342 (BM). H D. sherriffi: sae a upper

1968} MUNZ, ASIAN SPECIES OF DELPHINIUM 111

all foliaceous, lanceolate to linear, entire or few toothed, 1-5 cm. long;
pedicels erect, pubescent, 2-4 cm. long; bracteoles 2-3, linear, placed
from just below the flowers to near the middle of the pedicel, 10-15 mm.
long; sepals blue-violet, pubescent, the upper one oblong-ovate, 15 mm.
long, 6 mm. wide, obtuse, the spur erect, obtuse, 12-16 mm. long, 3.5 mm.
thick at the base; lateral sepals 13 mm. by 7 mm., oblong-elliptic, pubes-
cent along midrib, obtuse; lower sepals 15 mm. by 7 mm., pointed,
pubescent; upper petals bluish, the laminae 10 mm. long, almost straight,
unequally bidentate at apex, the spur thick at base, recurved, narrow at
tip; lower petals bluish, the claw 6 mm. long, the bearded lamina oblong,
7 mm. long, divided ca. 4.5 mm. into lance-oblong segments; stamens 7-8
mm. long; anthers dark, 1 mm. long; follicles 3, sparsely silky.

Type: Szechwan, in western part, Potanin in 1885 (P); seen.

REPRESENTATIVE MATERIAL. Szechwan: H. Smith 4200, 3541; E. H. Wilson
3093; W. P. Fang 4037, 4038, 1083.

172. Delphinium osseticum N. Busch, Bull. Acad. Sci. U.S.S.R. 7:
449, 1931 Fic. 15, D.

Stem ca. 1 m. tall, more or less muricate-roughened to bristly with white
spreading hairs, leafy; petioles thickly hairy, not vaginate at base; leaf
blades to 1.5 dm. in diameter, deeply 5-fid into rather narrow segments,
densely and softly hairy, especially on the upper surface, more or less
appressed underneath where the hairs along the veins are longer, the
blades pinnately shallowly laciniate or serrulate into lanceolate teeth or
lobes 2-5 mm. wide, 4-15 mm. long; inflorescence rather a dense raceme,
2~5 dm. long, branched, with soft pubescence; middle and upper bracts
entire, lanceolate, hairy, bluish, 15-25 mm. long; pedicels ascending or
erect, 1-4 cm. long; bracteoles densely white-hairy on the back and edge,
ovate, ovate-elliptic or ovate-lanceolate, acuminate, usually 7-10 mm.
long, 3-5.5 mm. broad, bluish, located near the flower; sepals blue, the
upper membranous, strongly veined, with long white hairs, strongly cu-

petal 9 mm. long, spur 20 mm.; blade of lower petal 6 mm. long, claw 5 mm.;
drawn from Ludlow & Sherriff 53833 TYPE (BM). IL. D. siamense; blade ~ upper
petal 11 mm. long, spur 18 mm.; blade of lower petal 6 mm. long, claw —
ISOTYPE (BM). J. D. sparsiflorum; blade of upper peta

mm. . :
from Rock 13191 (cH). K. D. taliense; blade of upper petal 9 mm.
22 mm.; blade of lower petal 7 mm. long, claw 5 mm.; dr f

mm.; blade of lower petal 6.5 mm. long, claw 5.5 ;

Hissar (te). M. D. Iriloliolesatts blade of upper petal 10 mm. long, rte oh

mm.; blade of lower petal 7 mm. long, claw 6 mm.; drawn from Henr id blad

(cH). N. D. umbrosum; blade of upper petal 8 mm. long, spur 24 mm.; blade

of lower petal 6 mm. long, claw 5 mm.; drawn from Wang 69052 (a). O. ue

uralense; blade of upper petal 10 mm. long, spur 13 mm.; blade of lower peta
mm. long, claw 5 mm.; drawn from Lifwinow TYPE (LE).

112 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

cullate, 12 mm. long, 7 mm. wide, the spur 13 mm. long, 4 mm. wide at
base, straight; lateral sepals 12 mm. by 8 mm., rounded-obovate ; lower
cucullate, mm. by 5 mm.; petals pale, the upper laminae slightly
oblique, 9 mm. long, emarginate, glabrous, the spur 10 mm. long; lower
laminae slightly oblique, ovate, 6 mm. by 5 mm., bifid halfway into 2
round-oblong ciliate lobes, the claw broad, 4.5 mm. long; stamens 6-7
mm. long, glabrous; anthers dark, 1.5 mm. long; carpels 3, glabrous.

Type: Upper borders of forest, Caucasus, eastern cis-Caucasus (Os-
setia), at 1800 m., July 23, 1930, Busch (LE), seen.
ILLUSTRATION. Bull. Acad. Sci. U.S.S.R. 7: 451. 1931.

173. Delphinium pachycentrum Hemsl. Jour. Linn. Soc. Bot. 29:
301. 1892. Fic. 15, E.

Delphinium pachycentrum subsp. hemsleyi Briihl ex Briihl & King, Ann. Bot.
Gard. Calc. 5: 108. 1896. me

Delphinium pachycentrum var. lobatum W. T. Wang, Acta Bot. Sinica 10:
140. 1962.

Delphinium sertiferum Franchet, Bull. Soc. Philom. Paris VIII. 5: 171. 18953.
Type, Tatsienlu, Szechwan, R. P. Soulié 24 (Pp); seen.

Perennial from a thick woody root, stem simple except at summit, 4-6
mm. thick, 3—4 dm. tall, leafy throughout; petioles slender, slightly dilated
at base, strigulose, 3-10 cm. long; leaf blades divided almost to base into
3 or 5 segments (the 2 lateral then divided almost to base), rounded-
pentagonal in outline, 3—10 cm. in diameter, ultimately shallowly incised
or toothed, the teeth oblong-ovate or deltoid, acute; inflorescence close-
pubescent, somewhat corymbiform, the lower pedicels elongate, solitary
or on short suberect branches, the upper 1-5 cm. long; lower bracts re-
duced foliage leaves, the upper smaller; bractlets linear, 5-8 mm. long,
near the flower; sepals pale blue, densely appressed-pubescent; uppet
sepal 14-16 mm. long, 9-10 mm. wide, ovate, acutish, the spur straight,
138-22 mm. long, 3-3.5 mm. wide at base, only slightly narrowed to a
blunt end; lateral sepals 14-16 mm. long, 6-8 mm. wide, oblong-ovate,
obtuse, pubescent on whole outer surface; lower sepals slightly longer,
oblong; laminae of upper petals whitish yellow, straight, 13-14 mm. long,
rounded and entire at apex, the spur 20-22 mm. long; lower laminae blue,
almost round, 9-10 mm. long, divided 1-2 mm., hirsute at base, sparsely
so otherwise, claw 8 mm. long, ciliate; stamens 7-8 mm. long, glabrous;
anthers dark, 1.4 mm. long; follicles 3, densely yellow-hairy, 10-12 mm.
long, 3-4 mm. wide, erect, the styles an additional 2 mm.: seeds rounded,
hyaline-alate.

Type: Szechwan, near Ta-tsien-lou, Pratt 462 (K), isotypes (BM, CAL,
P); all seen, :

OTHER COLLECTIONS. Tatsienlu, Soulié 2379 (x, Pp); Harry Smith 11051 (A,
MO, NY); Mt. Konka, Konkaling, Rock 16884 (Pp).

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 113

174. Delphinium pogonanthum Hand.-Mazz. Symb. Sinicae 7: 279.
1931. Fic. 15, F.

Stems 1-2, fistulose, 5-8 dm. tall, scattered-bristly with stiff white hairs,
remotely few leaved; lower petioles 1-2 dm. long, bristly hairy, the bases
broadly sheathing, upper petioles short; leaf blades broadly cordate-
rounded in outline, to 1 dm. across, with scattered appressed stiff hairs,
trifid to ca. 1 cm. from the base, the two lateral divisions deeply cut so as
to make 5 divisions, these broadly cuneate-obovate, coarsely and deeply
toothed distally, the teeth 3-8 mm. long, 3-4 mm. wide, subacute; upper
leaves reduced, then passing into trifid foliose bracts 15-25 mm. long,
upper bracts linear-lanceolate, entire, hairy, ca. 10 mm. long; inflores-
cence a lax raceme to 3.5 dm. long, or with 1-2 short racemose branches
below; the axis and pedicels with spreading and retrorse, coarse, hyaline,
white or yellow hairs to almost 1 mm. long; pedicels ascending, I1—5 cm.
long; bracteoles 2, erect, broadly linear, 6-10 mm. long, near to or some-
what below the flower; sepals dark blue, stiff-pubescent; upper sepal
broadly ovate, 13 mm. long, 10 mm. wide, obtuse, the spur horizontal,
slightly decurved, ca. 20 mm. long; 3 mm. wide at the base, narrowed to a
long tapering apex; lateral sepals asymmetrically elliptic-oblong, 12 mm.
by 6 mm., pubescent along the midline, rounded at apex; lower sepals
more pubescent, 12 mm. long, 4 mm. wide, rounded at apex; upper petal
laminae bluish, almost straight, entire, glabrous, 9 mm. long, the spur 19
mm. long; lower laminae oblong, oblique, 7 mm. long, 2 mm. wide,
bearded near base, long-ciliate the apex unequally shallowly 2-lobed;
stamens 6-7 mm. long, glabrous; anthers dark, linear-oblong, 1.3 mm.
long; follicles 3, pubescent, 10 mm, long, the style an additional 3 mm.;
immature seeds with transverse tiers of scales.

Type: N. W. Yunnan, 3600-4025 m., southeast of Dschungdien, be-
tween Peti and Alo, Aug., 1914, no. 4576, Handel-Mazzetti? (w?), not
seen. In the original description there is a figure on p. 281 which is
helpful.

Specimens to be referred here: Yunnan: H. 7. Tsai 57615 (A); Konkaling,
Muli, 3600 m. T. T. Vii 13270 (a, BM, E); Muli, Kulu, 3300 m., Yu 14367 (BM,
E); Haba Snow Range, K. M. Feng 2115 (a); S. Chungtien, Feng 1330 (A).

175. Delphinium pseudo-campylocentrum W. T. Wang, Acta Bot.
Sinica 10: 150. 1962
Delphinium hirticaule Franchet var. mollipes Wang, Acta Bot. Sinica 10: 146.
1962. Based on K. H. Yang 58941, from Wu-hsi, Szechwan, isotype (E),
seen.

Root elongate, 2-5 mm. in diameter; stem ca. 4 dm. high, 4 mm. thick
at base, spreading-hirsute below, glabrous above, simple or few branched
above, equally foliose throughout; lower cauline leaves with broadly pen-
tagonal blades 4-4.5 cm. long, 4.5—6.5 cm. wide, cordate at base, trisect,
the middle segment 3-parted to below the middle, the lateral segments

114 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

broader than the middle, 2-parted for three-fourths their length, all parts
few lobuled, strigulose especially above; petioles 4.5-5.5 cm. long, not
vaginate, hirsute; upper cauline leaves gradually reduced; racemes 4-9-
flowered, lower bracts leaflike, 2-4 cm. long, others linear to subulate, 3-10
mm. long; pedicels 2-4 cm. long, divergent, slightly incurved, bracteolate
above the middle, loosely yellow glandular-puberulent; bracteoles oblong-
linear to subulate, 2-6.5 mm. long, loosely strigulose; sepals deep blue,
narrowly elliptic or broadly lanceolate, 11-15 mm. long, minutely puberu-
lent on both surfaces, the spur 17-22 mm. long, subulate, slightly decurved
at apex; upper petals blue, glabrous; lower petals blue, the limb ovate,
bilobed one-third to one-half the length, ciliate, yellow bearded at base;
stamens glabrous; carpels 3, loosely puberulent.

Type: Szechwan, Tachin, 4400 m., Aug. 5, 1958, H. Li 78358 (PE), not
seen; photo. at (E) seen.

The only specimen I have seen is the one mentioned above as isotype of
D. hirticaule var. mollipes.

176. Delphinium scabriflorum D. Don, Mem. Wernerian Soc. 3: 412.
1821. Fic. 15, G.

Delphinium carela Buch.-Ham. ex D. Don, Prod. Fl. Nepal. 195. 1825. New
name for D. scabriflorum.

Perennial from rather a woody root; stem 6-12 dm. or more tall, rather
slender, terete, more or less spreading-hairy throughout or at least above,
scattered-leafy; leaves pentagonal, the blades of the lower 5-15 cm. wide,
open-cordate, 5-fid to near the base or 1-2 cm. above, the segments sub-
cuneate below, stiff-pubescent on both sides, but especially on nerves
beneath, the segments incised-lobed then coarsely toothed, the lobules
oblong-ovate, 2-5 or more mm. wide; petioles hairy, 1-2 dm. long, scarcely
vaginate at the base; upper leaves gradually reduced; central raceme
open, 8~12-flowered, with fewer-flowered branches from below; inflores-
cence densely short-hairy, some of the hairs glandular; lower bracts
foliose, divided, to 3 cm. long, the upper simple, entire, linear to lanceolate,
1-2 cm. long; pedicels arched-spreading, densely pubescent, 1—5 cm. long;
bracteoles linear, to lanceolate, 5-10 mm. long, subtending the flower;
sepals blue to rich blue-violet, with rather stiff appressed hairs, the upper
sepal round-ovate, 15 mm. long, 10 mm. wide, subacute, the spur 18 mm.
long, 3 mm. wide at base; lateral sepals oblong-obovate, 13 mm. long, 7
mm. wide, obtuse; lower asymmetrically obovate, 15 mm. long, 8 mm.
wide; petals blue, the upper laminae oblique, 12 mm. long, 3 mm. wide,
glabrous, bidentate, the spur 15-16 mm. long; lower laminae oblong, 10
mm, long, 3.5 mm. wide, bifid into 2 unequal lance-oblong lobes, white-
ciliate, the claw 6 mm. long, bearded; stamens 6-7 mm. Jong; anthers
elongate, 1.5 mm. long; follicles 3, subglabrous.

Type: “Alpibus Nepaliae,” D. Wallich; not seen.

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 115

RANGE. Growing on open or wooded, often somewhat rocky hillsides,
mostly in Bhutan.

2500 m., Ludlow & Sherriff 3342 (pm); Yatola ridge, Tongsa, 3200 m., Cooper
G& Bulley 4114 (pm). A specimen from Burma, Chimili woods, R.A., Aug. rae
1919 (£) is near the above.

¥177. Delphinium sherriffii Munz, sp. nov. Fic. 15, H.

Perennial, the stem 10-16 dm. tall, 7-8 mm. thick, with short white
spreading pubescence, simple below, short branched above, rather equably
but not densely leafy; cauline petioles pubescent, slender, 6-10 cm. long,
slightly dilated at the base; cauline leaves to ca. 1 dm. across, 5-fid to 1
cm. or less from the base, pubescent chiefly along the veins, the segments
rather broadly cuneate-rhombic, 2—3-lobed to about the middle, then
sharply incised into few lanceolate or lance-ovate teeth mostly 3-5 mm.
broad, mucronate; inflorescence with several short branches, thus form-
ing a rather diffuse but compact panicle, the axes with short, spreading,
yellow, glandular hairs; lower bracts divided, but mostly simple, filiform,
5-7 mm. long; pedicels spreading, 2-4 cm. long, glandular-pubescent;
bracteoles filiform, 2-3 mm. long, near or above the middle of the pedicel ;
flowers few to several on each branch of the inflorescence, not crowded;
sepals somewhat strigulose, bright bluish purple, the spur dark wine-red ;
upper sepal 12 mm. by 5 mm., broad-ovate, apiculate, the spur spreading
decurved, 20-22 mm. long, 4-5 mm. wide at base, rapidly narrowed; lateral
Sepals 12 mm. long, 4.5 mm. wide, rounded-obtuse at apex; lower sepals
12 mm. long, 4 mm. wide, more apiculate; petals pale, the upper laminae
strongly oblique, 8-9 mm. long, 4 mm. wide, glabrous, emarginate, the
spur 20 mm. long; lower laminae 6 mm. long, divided two-thirds their length
into lance-oblong ciliate lobes, claw appendiculate, 5 mm, long; stamens
5—6 mm. long; anthers dull slaty green; carpels 3, strigulose.

Caulis 10-16 dm. altus, 7-8 mm. crassus, pubescens, supra breve
ramosus, foliosus; petioli caulium 6—10 cm. longi, base dilatati, pubes-
centes; folia caulium 6-10 cm. lata, 5-fida, praesertim in venis pubescentia,
segmenta foliorum cuneato-rhombea, 2—3-lobata, tum incisa lobulis paucis
lanceolatis vel lanceo-ovatis; inflorescentia ramosa, axes aureo-glandulosi ;
bracteae infimae divisae, aliae integrae, filiformes, 5-7 mm. longae; pedi-
celli divaricati, 2-4 cm. longi, glanduloso-pubescentes ; bracteolae fili-
formes, 2-3 mm. longae; flores pauci vel plures; sepala strigulosa, caeruleo-
purpurea, 12 mm. longa, calcar 20-22 mm. longum, base 4-5 mm. crassum;
petala pallida, laminae superiores obliquae, 9 mm. longae, glabrae, emar-
ginatae; laminae inferiores 6 mm. longae cum 2 lobis lanceo-oblongis cilia-
tisque; unguis 5 mm. longus appendiculatus; stamina 6 mm. longa;
antherae subviridia; carpella 3, strigulosa.

Type: Southeastern Tibet, Tsanang La, 3200 m., July 15, 1938, Ludlow

116 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

& Sherriff 5833 (sm), growing in clearings in mixed forest. Characterized
by its small flowers in a branched compact, but not dense, inflorescence.
Mr. Sherriff’s collections, largely made with Mr. Ludlow, have been of
great help in the preparation of this paper.

178. Delphinium siamense (Craib) Munz, comb. nov. Fic. 15,5
Delphinium stapeliosum Brihl var. siamense Craib, Florae Siamensis Enumer-
atio 1 1925.

Stem slender, 3-5 dm. tall, few branched above, scattered-leafy, with a
few spreading white hairs below, strigulose upward, often quite loosely so;
lower petioles slender, 1-2.5 dm. long, vaginate, with scattered white
hairs, the upper petioles shorter; blades mostly round-pentagonal in out-
line, cordate at base, 3-7 cm. long, 4-10 cm. wide, divided halfway or
farther with sharply acute sinuses and broad cuneate-rhombic segments,
sparingly strigose above, glabrous beneath except on the veins, the seg-
ments shallowly but sharply trilobed then incised with lobules or few
teeth triangular-lanceolate or broader (2—6 mm. wide), acute to acuminate,
mucronate; upper leaves reduced; inflorescence of a few slender-stemmed
racemes (the central 4-6—-8-flowered), these loosely strigose; lowest bract
trifid or entire, the upper lance-linear, 10-15 mm. long; pedicels mostly
1-2 cm. long, ascending; bracteoles linear, 5-7 mm. long, at or below the
middle of the pedicel; flowers light blue, the sepals strigose, the upper
round-ovate, 14 mm. long, 9 mm. wide, obtuse, the spur straight, 3.5 mm.
wide at base, 16-18 mm. long, tapered very gradually to a blunt tip;
lateral sepals round-obovate, 13-14 mm. long, 12 mm. wide, rounded at
apex; lower sepals obovate, 14 mm. long, 8 mm. wide, obtuse; petals pale,
the upper laminae somewhat oblique, 11 mm. by 5 mm., glabrous, strongly
bidentate at apex, the spur 17-18 mm. long; lower laminae more or less
obovate, 6 mm. by 4 mm., bearded, deeply bifid into broad oblong lobes,
the claw 6 mm. long; stamens 6-7 mm. long, glabrous, the darkish anthers
1 mm. long; carpels 3, strigose.

Type: Siam, Doi Chiengdao, 2100 m., open rocky ground, Kerr 6612
(K, BM); seen.

THER COLLECTIONS: Doi Chiengdao, Put 337, Herb. A. F. G. Kerr (BM, K),
4454 (BM, K); Doi Pa Mawn, 1510 m., H. B. G. Garrett 452 (x).

179. Delphinium sparsiflorum Maxim. Bull. Acad. Pétersb. 23: 307.
ites Fic. 15, J.
Perennial, the rather coarse stem 10-15 dm. tall, subglabrous, rather
equably leafy in lower half, the petioles scarcely dilated at base, slender,
to ca. 1 dm. long; leaf blades sparsely strigose, paler beneath, to ca. 1.5
dm. in diameter, ternate, then the 2 lateral parts divided nearly or quite to
the base, they and the central one with cuneate base and incisely pinnately
lobed, each lobe with few large, mucronate, lanceolate or ovate teeth or
lobules; flowers scattered in an open diffuse panicle forming the whole uppet

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 117

part of the plant; lower bracts foliaceous, more or less coarsely incisely
lobed, 1-3 cm. long, the upper subulate, entire, 2-4 mm. long; pedicels
filiform, glabrous, 2-4 cm. long; bracteoles alternate or paired, ca. 2 mm.
long; flowers pale blue, the upper sepal broadly ovate, 9 mm. long, 8 mm.
wide, subglabrous except for a very minute puberulence about the obtuse
apex; spur conical, ca. 7 mm. long, 3.5 mm. wide at base; lateral sepals
10-11 mm. long, 6-7 mm. wide, glabrous except at the tip; lower sepals
10-11 mm. long, 4-5 mm. wide, also minutely puberulent at apex; laminae
of upper petals 8-9 mm. long, pale, suboblong, glabrous; lower laminae
pilose, 7 mm. long, the blade bifid, 4 mm. long, the claw ciliate; stamens
ca. 4 mm. long, glabrous; anthers dark, 1 mm. long; follicles 3, sparsely
pilose, to ca. 12 mm. long, 3 mm. wide, ‘the styles an additional 2m mm.

Type: Kansu, China; Terra Tangutorum, Przewalski, 1872 (LE), seen.
Huth says this region is Tangutica at River Tetung, 2600 m.

ILLUSTRATION. Maxrmowicz, Fl. Tangut. pl. 5. 1889.

The only collection I have seen in addition to the type is from southwestern
Kansu, Tao River Basin, at 3000 m., Rock 13191 (cH).

180. Delphinium sungpanense W. T. Wang, Acta Bot. Sinica 10: 149.
1962.

Stem about 9 dm. tall, 2 mm. thick, smooth, glabrous below inflorescence,
equably leafy; middle cauline leaves with rather long petioles, the blades
broadly pentagonal, 6 cm. long, 10 cm. wide, trisect, the middle segment
rhombic, long-acuminate, cuneate at base, trifid to middle, the middle lobe
narrow- Mmmnbic or lance rhombic, with 1 lobule on each side below the
middle, the lateral lobes triangular- ovate or obliquely ovate; lateral seg-
ments broadly rhombic, parted and lobed as in the middle segment; blades
subglabrous below, strigulose above; racemes ca. 15 cm. long, lax, about
8-flowered; lower bracts trisect or linear, 4.5—5 cm. long, others linear,
shorter; pedicels 1.5—5 cm. long, strigulose, bibracteolate above; bracteoles
remote from the flower, linear, 5-7 mm. long, glabrous; sepals purple-blue,
oblong-obovate, 8 mm. long, strigulose, short-corniculate, the spur 19-22
mm. long, 2.5 mm. wide at base, decurved into a right angle; upper petal
laminae yellowish, entire, elabrous: lower blue, subequal to sepals, ovate,
bifid to middle into lance-triangular lobes, ciliate, yellow bearded near
middle, claw shorter than limb, appendiculate; stamens glabrous; carpels
3, glabrous.

Type: Sungpan, Szechwan, W. P. Fang 4038 (PE?); nots

Said to be near D. giraldii, but racemose, the axes and nedicels densely
puberulent, and the spur decurved into a right angle.

181. irene taliense Franchet, Bull. Soc. Philom. — VIL. 2
174. 18 c. 15,

Stem me to 1 m. and more in height, much branched from _

glabrous and Waneous: leaves numerous and finely pubescent, with petioles

118 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

8-12 cm. long and somewhat dilated at base, the blades 5S—10 cm. wide,
deeply divided in the lower leaves into 3 broad segments, these cuneate-
obovate, trifid, incised, then shallowly lobed, the ultimate teeth few,
broadly ovate, mucronate, 5-15 mm. long, 5-12 mm. broad; the divisions
of the upper leaves lanceolate, subentire or few lobed; branches of in-
florescence elongate, arched, with an entire linear leaf at the base; bracts
linear-setaceous, 8-10 mm. long; pedicels 2-4 cm. long; bracteoles seta-
ceous, minute, placed a little below the flower; sepals puberulent, blue
with a whitish band on the back, 15-20 mm. long, obtuse to rounded, spur
straight or slightly curved, 22-25 mm. long, 3 mm. wide at the base; upper
petals bluish to whitish, the laminae slightly oblique, 9 mm. long, glabrous,
deeply emarginate, the spur ca. 21 mm. long; lower laminae sometimes
white, deeply bilobed, yellow bearded near base, ciliate, round-obovate, 7
mm, long, the claw 5 mm. long, appendiculate; follicles 3, glabrous; seeds
black, pyramidal, truncate, somewhat rugulose, winged on angles.

Type: Yunnan, Mt. Pen-gay-tzé, below Tali, Delavay 9 (Pp), seen.

OTHER COLLECTIONS. Yunnan: Bodinier & Ducloux 379 (pe); Delavay, Sept.
22, 1887, at 2000 m. (P).

This species, Delphinium taliense, suggests D. yunnanense in the long
segments of its upper leaves. It has stem leaves like those of D. exaltatum,
and has been confused with D. elatwm, which has the spur equal to the
sepals, and almost black petals.

182. Delphinium ternatum Huth, Bot. Jahrb. 20: 421. 1895.
Fic. 15, L.
Perennial, 5-10 dm. tall, erect, soft-pubescent, the petioles to 1 dm.
long, scarcely dilate at the base, the leaf blades 4-8 cm. broad, glabrescent
on both surfaces, ciliate on margins, ternate, the leaflets deeply tripartite,
the parts entire or sparsely serrate-incised, subsessile, cuneate, the final
lobules lanceolate, somewhat acuminate, to 3 cm. long; inflorescence
branched, the racemes lax, glabrous; bracts linear, 5 mm. long; pedicels
1-2 cm. long in fruit, divergent, shorter in flower; bracteoles linear, at
about the middle of the pedicel, 2-3 mm. long; flowers blue, the sepals
round-ovate, glabrous, the upper 12 mm. by 5 mm., subacute, the spur 15
mm. long; lateral sepals elliptic, rounded-obtuse, 13 mm. by 5 mm.; lower
sepals more pointed; upper petals yellowish with bluish tinge, the laminae
oblique, 10 mm. long, bidentate, the spur 15 mm. long; the lower laminae
bluish, deeply divided, sparsely white-bearded, 6.5 mm. by 4 mm., the
claw 5.5 mm.; anthers yellow; follicles 3, hirsute: seeds squamate. the
scales contiguous.

Type: Huth cited two specimens from Buchara: (1) Darwas, Omar ad
fluvii Pansch ripam sinistram, Regel, 1882 (tr) and (2) distr. Hissar,
mts. near Hakimi in valley of R. Karatag, Regel, 1882 at 1600-2300 m.
(LE). Both seen; I would take the Hissar specimen collected by Regel as
lectotype since it has more leaves intact than his other collection.

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 119

Apparently belonging here is Mt. Kuhitang, eastern Turcomania, July 14,
1931, S. Nevski 790 (x).

183. Delphinium trifoliolatum Finet & Gagnep. Bull. Soc. Bot. France
51: 481. plate 6A, figs. 1-6. 1904. Fic. 15, M.

Perennial from a short thick root; stem slender, subflexuous, striate,
simple or branched above, 3-7 dm. tall, more or less strigulose throughout,
densely so in the inflorescence, leafy throughout, the leaves rather remote;
petioles very slender, 3-10 cm. long, somwhat inflated at the base; the
blades round-pentagonal in outline, strigose to strigulose on both surfaces,
especially along the veins, 4-10 cm. in diameter, trifid to the base, the
lateral segments bifid almost to the base, all 5 segments thus formed more
or less petiolulate, lance-ovate, narrow-cuneate at base, 1-2 cm. wide,
shallowly incised or unequally sharply dentate into acute teeth 2-4 mm.
long; upper leaves somewhat reduced and passing into trifoliolate bracts
5-20 mm. long; racemes lax, rather few flowered, the pedicels strigose,
5—30 mm. long, very slender, divergent; bracteoles lance-oblong, 3-8 mm.
long, near the summit of the pedicel; flowers pale blue, sepals strigulose,
yellowish near the apex; upper sepal ovate, 10 mm. long, 6 mm. wide,
subobtuse, the spur 18-20 mm. long, 2.5 mm. wide at base, somewhat
curved toward the narrow tip; lateral sepals oblong, 10 mm. by 5 mm.,
rounded at apex, pubescent along midline; lower sepals 12 mm. by 7 mm.,
more pointed, more pubescent; laminae of upper petals pale, slightly
oblique, 10 mm. long, bidentate at the acute apex, glabrous, the spur 20
mm. long; lower laminae bluish, oblong, 7 mm. long, 3 mm. wide, sparsely
bearded, cleft to below the middle into lance-oblong lobes; stamens 5 mm.
long, glabrous; anthers dark, 1.5 mm. long; follicles 3, glabrous or nearly
so, 5 mm. long, the styles 1-2 mm. more.

Type: Hupeh, China, E. H. Wilson 2496, published as 3496, July, 1901,
holotype (P) seen; isotypes (E, K, NY) seen.

REFERRED HERE: Hupeh, A. Henry 7448 (BM, GH, K); E. H. Wilson, April,
1900 (us). Szechwan: Nan-chuan, 1580 m., K. F. Li 63169 (£).

184. Delphinium umbrosum Hand.-Mazz. Symb. Sinicae 7: 278. fig.
4,nos. 7-9. 1931. Fic. 15, N.
Rhizome short, thickish, the roots fascicled, long, thick, woody; stem
6~9 dm. tall, simple, sulcate, short- and retrorse-strigose, equably leafy;
leaves pentagonal-reniform in outline, to 12 cm. wide, tripartite almost to
base, the lateral parts less profoundly bifid, all parts narrowly obovate-
rhombic, rather long-acuminate, incised, dark green, sparsely strigose
above, densely so beneath; upper petioles gradually shorter, narrowed into
slender sheaths; racemes lax, to 10-flowered, densely retrorse-hairy, with
normal leaves below and entire lanceolate to linear bracts above; pedicels
1.5—-6 cm. long, suberect or arcuate; bracteoles apical or farther down,
linear, 8-10 mm. long, strigulose; sepals violet, broadly ovate, 12-15 mm.

120 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

long, rounded, the lateral with median nerve dilated at apex, strigulose ;
spur more than twice the length of the blade of the sepal, deflexed, 3 mm.
wide at the base, obtuse; upper petals almost 3 mm. shorter than the
sepals, ca. 3.5 mm. wide, oblique; truncate, minutely and acutely bilobed
at tip; lower laminae shorter, hairy from the claw, 6 mm. by 3 mm., rather
deeply oblong-lobed, the claw 5 mm. long; stamens 6-7 mm. long,
glabrous; anthers brown; carpels 3, sparsely hairy.

Type: Northwest Yunnan, hemlock forests on the Nguka-la between
Dschungdien (Chungtien) and Djitsung on the Djinsche-djiang, 3750-
3800 m., Handel-Mazzetti ? 7809 (w?), not seen. The illustrations are
helpful.

SPECIMENS SEEN. Yunnan: C. W. Wang 69052 (a); H. T. Tsai 52675 (a); K.
M. Feng 1726 (a); A. Henry 9668 (mo, NY), 9668 A (us). Szechwan: T. T. Yu
979 (GH). Upper Burma: Forrest 25052 (us), characterized by its long-acuminate
leaf lobes.

185. Delphinium uralense Nevski in Komarov, Fl. U.S.S.R. 7: 159.
125s; 1937, Fig. 15, 0.
Delphinium dyctiocarpum DC. subsp. uralense (Nevski) B. Pawl, Fragmenta
Floristica & Geobotanica 9: 429. 1963.
Delphinium dyctiocarpum var. canescens Korsh. Tent. Fl. Ross. Or. 18. 1898.

Entire plant grayish-strigulose; stems 5—11 dm. tall, ribbed below, rather
equably and closely leafy; petioles to 1 dm. long, vaginate at base; leaf
blades round-cordate to -reniform, 3.5-7 cm. long, 5.5-12 cm. wide,
palmatisect, the segments commonly ternately divided into terminal lance-
linear lobes 2-4 mm. wide and with 2-3 teeth, densely strigose on both
surfaces, but paler beneath; raceme sometimes branched below, dense,
many flowered, the flowers light blue; bracts lance-subulate, stiff, 5-7 mm.
long; bracteoles strigose, linear-subulate, 3.5-6 mm. long, placed above
the middle of the pedicel which is strigose, suberect, to 1 cm. long; sepals
ovate, obtuse, tomentellous, the upper ovate, 14-15 mm. long, 8 mm. wide,
the tip subacuminate, upcurved; spur cylindrical, 12 mm. long, 2—-2.5 mm.
wide, obtusely acuminate, more or less uncinate at tip; lateral sepals ellip-
tic-obovate, 13-14 mm. by 8-9 mm., obtuse; lower sepals 13 mm. by 6
mm.; petals pale, the upper laminae somewhat oblique, 10 mm. long, 3
mm, wide, entire and glabrous, the spur 13 mm. long; lower laminae very
oblique, rather oblong, 5 mm. by 3 mm., bifid to one-fourth its length into
rounded-oblong lobes, bearded and long-ciliate, the claw 5 mm. long;
stamens 5—6 mm. long; anthers dark, round-oblong, 0.8 mm. long; carpels
3, very densely pubescent.

Type: Prov. Orenburg, near Andreevka, D. Litwinow July 7, 1893
(LE); seen.

Delphinium uralense differs from D. dyctiocarpum which is much less
pubescent and has dark petals.

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 121

GROUP Ix
Leaves rather coarsely lobed as in Group VIII, but the petals very dark,
in most cases almost black. Spurs straight or some decurved.
This group is purely artificial and does not correspond with any tax-
onomic subdivisions that have been proposed.

Key TO SPECIES
i sae about half as long again as the blade of the upper sepal.
Stem and inflorescence axis glabrous or nearly so
3. Sepals mostly glabrous, 10-25 mm. ;
4, aes sepal blade 21-25 mm. fae spur cage mm. long. Szech-
than mips, o teil tet meiats raga amen tam atite ae . D. oxycentrum.
4. tate sepal blade 10-14 mm. long; spur ee mm. long. Altai.
2 GEG oo voy pete a i ee ne 199. D. dyctiocarpum.
3. Sepals not glabrous, 14-16 mm. long. Szechwan. Saree’ | in Group
VATD) s,s at te eee 169. D. mitzugense.
. Stem and/or inflorescence axis pubesc
5. Inflorescence with spreading Re hairs; bracteoles i mm. long.
Shansi. 212. D. leptopogon.
5. Inflorescence lacking glandular hairs; bracteoles largely ” 10 mm. long.
6. Lamina of upper petal ciliate; carpels g page stem pilose.
Basteen SipeHa.04 Jd isaien yan oe 195. D. crassicaule.
6. Lamina of upper petal glabrous; carpels hai
7. Stems more or less retrorse-pubescent; cated lance-linear;
sepals glabrous within. Western China. ...... 191. D. chenii.
7. Stems with long spreading hairs; bracteoles lance-ovate; sepals
strigose on both surfaces. Semipalatinsk. . . 228. D. sauricum.
1. Spur mostly not half as long again as the blade of the upper sepal.
8. Sepals pubescent within.
9. Sepals glabrous on the outer surface.
10. Inner surface of sepals with long tangled hairs at base, or at least
a few hairs near the tip; sepals 12-15 mm. long. Southern
Caucasus, adjacent Turkéy..<-7. 2... 5... 190. D. buschianum.
10. Inner surface of sepals not long-hairy; sepals 18-30 mm. long.
11. Spur 15~25 mm. long; laminae of upper petals glabrous.
China. (Treated in Group VI). .... 81. D. albocoeruleum.
11. Spur 12-15 mm. long; laminae of upper petals bristly.
205. D. formosum.

i)

ae ee ee ek oe
9. Sepals pubescent on the outer surface
12. Inflorescence axis with spreading hairs.
13. Upper sepal 12-15 mm. long, spur 10-15 mm. long.
14. Sepals white with yellow nerves; lamina of upper petal
entire, hairy at tip. Uzbekistan. ..... 219. D. nevskii.
. Sepals blue or purple; lamina rv upper petal emarginate,
often glabrous at tip.
15. eaapaeN at the base of the flower; sepals deep
blue to purple.
16. Stems 1-4 dm. tall; spur 10-12 mm. long;
lower petioles vaginate. Turkestan, Pamir.
(Treated in Group V). .. 62. D. dasyanthum.

rs
+

122 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

16. Stem 6-12 dm. tall; spur 12-15 mm. long;
lower petioles scarcely dilated. Himalaya. ...
oped as Piles 1533 224. D. pyramidale.

15. Bracteoles remote from the flowers; sepals dilute
lue Ra cts whic 221. D. oreophilum.
13. i ge sepal 15-25 mm. long; spur 15-18 mm. long.
Lamina of upper petal entire, 10 mm. long; lamina of
lower petal 5 mm. long. Altai. _. 217. D. mirabile.
17. Lamina of upper petal not entire, 13-14 mm. long;
lamina of lower petal 7-8 mm. lon
18. The lamina of upper petal bidentate; bracteoles ca.
10 mm. long; seeds winged. Himalaya. (Treated in
SEE NK, fies Ge ae 2. 55. D. vestitum.
18. — of upper petal emarginate; seipiares: 4-6
m. long; seeds squamate. Caucasus. ........--
es Baie ee went ss 229. D. speciosum.
12. Inflorescence with appressed hairs.
r sepal slaty to pale blue, | mm. long. Tibet,
China. (Treated in Group VI). .... 81. D. albocoeruleum.
19. Upper sepal deep blue to purplish, 12- os mm. long.
20. Lamina of upper petal rounded at apex; upper sepal 12-
mm. long; spur 11.5-13 mm. long. Western China.
Be ie oe ov es a 210. D. kuanit.
20. Lamina of upper petal emarginate at apex; upper sepal
15-17 mm. long; spur 18-20 mm. long. Iran. ........
ee ce va UE oy 201. D. elbursense.
8. Sepals not pubescent on inner surface.
21. Sepals glabrous on outer surface except sometimes ciliate on margin.
22. Pedicels So hess aaiptiged stem 10-15 a tall, mostly gla-
bE en cee 4, D. pyramidale.
Pal Pedicels not act,
23. Upper sepal 10-16 mm. long.

24. Flowers ca. 10 in a lax raceme; upper sepal 13-14 mm.
long; spur 17-18 mm. long; laminae of upper petal
emarginate. Sinkiang. ....... 213. D. longiciliatum.
Flowers many in a dense raceme.

25, i sepal 10-11 mm. long, the spur 10-13 mm.

nm
aE

nN

ae

heel: |
aa

Leaf blades cordate at base; bracteoles ovate

brous at tip. Sikang. ... 218. D. naviculare.
Leaf blades almost straight across the base;
bracteoles oblanceolate, 5-8 mm. long; upper
petal bristly-hairy at se Sens. (oe
See eer 226...) retropilosum.
. ie: sepal 13-16 mm. long; spur 13-18 m

i)
=

Led
on

27. Lamina of upper petal subtruncate at apex;
leaf blades round-cordate in outline. Central
and eastern Siberia. ....__. 200. D. elatum.

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 123

27. Lamina of upper petal sneered = re leaf
blades broader than long. Caucasus. .......
ode ca ees, Cte eee 204. > arene)

23. Upper sepal 18-30 mm. long.
28. Bracteoles 2~3 mm. wide; upper sepal 18~20 mm. long.
29. The bracteoles well below the flowers; inflorescence
lt be eg stem with scattered long
hairs. Ossetia. ... 188. D. bracteosum.
29. The preieri subtending the flowers; inflores-
cence racemose; stem strigulose peri otherwise
glabrous. Dagestan Mie kay; D. mariae.
28. papers 3~5 mm. wide; upper wa pats longer.

Spur 12-15 mm. long; upper sepal 20-30 mm,

long; inflorescence paniculate. Turkey. .........

Bi aa ae tee eee 205. D. formosum.

30. Spur 18-20 mm. long; upper sc a mm.
long; inflorescence racemose.

31. Lamina of upper petal deply shied up-
per sepal 20 mm. long. . a ruprechtii

31. Lamina of upper petal pan lee h upper
sepal 20-80 tI NWR oe i Sis hee

21. Sepals not glabrous on outer surface
32. Inflorescence with some glandular hairs (yellow, inflated), these
often mixed with various non-glandular hairs.
33. Sepals variegated, pale blue and white. Transcaucasia. ....
, Ems Phe ar Sea Bae 202. D. elisabethae.
33. Sepals not so variegated.
34. Flowers rather large, the sepals mostly 15-22 mm. long.
35. Lamina of upper petal sharply eae of lower

petal long- gente Dagestan. . arcuatum.
35. Lamina of upper petal entire, ot ais pin: not
ciliate. Siberi cou eee 0. D. ochotense.

. Flowers smaller, oe sepals 9-14 mm. ape
36. Upper petal with 2 narrow terminal pointed lob-
ules; flowers white, tinged gare any Sou
Tibet.

Ww
>

thern
. D. hillcoatiae.

Ot ee ee ds

36. Flowers violet, blue or purple.
37. Inflorescence 1-2—(3)-flowered; upper sepal
mm. long; spur 14-16 Fes . long. Bhu-

te a ee 193. D. cooperi.
ous eo reneagt with more numerous flowers.
38. Plant 2.5-4 dm. tall; raceme 6-15-flow-

ered; spur Bact mm. long. Altai. .....
232. D. ukokense.

. Plant mostly taller.
39. Flowers 3-10; upper oa 12-14
mm. long. — Baia.
Su Col esr et rarer 189, D. gj Pape
39. Flowers more numerous; upper sepal

Se
ie)

124

JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

40. Sam of sepals not gland-

ao
wa

"Sepals villous with spread-
ing white hairs, pale pur-
plish blue; carpels hairy.
Afghanistan... ...-..-.4
oiey 3 186. D. afghanicum.
puberulous, pale
blue; carpels glabrous.
Temacacasia. .
25. D. pyramidatum.
. Pubescence of sepals short,
glandular; sepals 10-18 mm.
long; — glabrous. Mon-
Mihe Mts
aa = D. inconspicuum.
32. Inflorescence lacking glandular hairs, the hairs white and slender.
42. Petioles scarcely or not dilated at the base.
43. Stems scarcely taller than the basal leaves; sepals 19-
25 mm. long. Caucasus. (Treated in Group V). .....-
eee as ee 61. D. caucasicum.
43. Stems much taller than the basal leaves.
44. Sepals strigulose.
Raceme many flowered, the sepals deep blue,
17-18 mm. long; spur 17 mm. long. Turke-
BM ee eas 234. D. winklerianum.
45. Raceme 4~-10-flowered, * sepals purplish,
10-15 mm. long. Sikang
Bracteoles 5-7 mm. long; spur 2-2.5
mm. thick at base; sain glabrous
eee cl kes . 203. D. elliptico- -ovatum.
. Bracteoles 8-10 mm. long; spur 4-4.
mm. thick at base; stamens pilose. ....-
ee eee 210. D. kuanit.
44. Sepals villous or hairy.
47. Upper sepal 12-15 mm. long, the spur about
as long.
Raceme lax; stem 10-12 dm. tall; lamina
of upper petal glabrous, entire. Siberia.
Pete eS 233. D. villosum.
. Raceme subcapitate; stem 2-6 dm. tall;
lamina ne upper Lee seeprt setose.
Wilrkestan. 5. D. poltaratzkii.
47. ‘Sia od Ves mm. ee spur 18-20 mm.
ng.

>
«
~
ue)
&
5,
wm

vs
te

as
an

—
Oo

>
ie)
a

>
o2)

49. Upper sepal ca. 18 mm. long. per
Assam to Nepal. .. 230. D. stapeliosum.
49. Upper sepal 20-22 mm. long. Caucasus.
(Treated in Group VIII). ......-.
Co 148. D. dasices pie
42. Petioles, at least the lower, more or less dilated at base.

1968] MUNZ, ASIAN SPECIES OF DELPHINIUM 125

50. Upper petals entire at the apex or nearly so,
51. Inflorescence with some glandular hairs; upper
sepal 14-17 mm. long; spur about as long. Central
1

un
~

Siberia
: Tnflovescence lacking glandular hairs.
5

an
nN

96. D. crassifolium,

2. Hairs of stem mostly spreading; upper sepal

14-16 mm. long; spur 15-20 mm. long. Man-
churia, Korma, =... 214 iaachianinn.

Hairs of stem retrorse to appressed.
53. pase sepal ca. 18 mm. long; spur 16-
. long; flowers more or less yel-
lowish: hairy. Burma. .... 211. D. lacei.

. Upper sepal 14-15 mm. long.

54. Spur ca. 22 mm. long. Sikang......
ict eet 91. D. chenii.
54. Spur ca. 12 mm. long. Nepal.
(Treated in Group v1). at an
- incisum,

UL
w

50. Upper petals emarginate or notched sp

55. Hairs closely appressed; sepals light. ig the
upper 13-15 mm. long; spur 14 mm. long. Trans-
caucasia. (Treated in-Group V1). 3.52, as.

. Hairs more or less spreading on —

91. D. crispulum,

56. Sepals nearly or quite glabrou

amina of lower petal divided halfway.

58. Upper petal with hairs at tip; sepal
spur 15-18 mm. long. Caucasus

204. D. flexuosum.

Upper petal glabrous at tip; sepal

spur 13-14 mm. long. Kolgujev

Island, northern ‘gig hes,

197. D. cryophilum.

57. Lamina of lower petal tannthy less deeply

divided.

un
—

59. Lamina of upper petal broadened
upward, oblique; bracts and brac-
atic rman payors Siberia. . .
Lamina of Mee petal scarcely
broadened upward, almost straight;
bracts and bracteoles linear. Widely
distributed across northern Siberia.

200. D. elatum.

on
—

56. Sepals pubescent.

60. Pedicels 5-15 mm. lon
61. Bracteoles lance-linear. Caucasus,
D

61. Bracteoles sige ovate. Asia
. D. confusum.

Media
60. Pedicels 10-80 mm. a

126 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

62. Flowers few, subumbellate, pale vi0-
let. East Turkestan. ..
oes 194. D. coryebosit:
62. Flowers several to many, racemose.
63. Bracteoles near the middle of
“m3 pedicel.

. Upper sepal 18 mm. long;
spur 20 mm. long. Khasia,
Dhaai
ae 230. D. stapeliosum.

. Upper sepal 12-16 mm.
long; spur 12-16 mm. long.
Northern . Rae
eee Rees . D. elatum.

63. Bracteoles ng near the flower.

65. Upper sepal 17-18 mm.
long; spur often suberect.
Nepal. 207. D. himalayae.

=

an
>

oO
on
oo
Ko)
a,
as)
ca
wm
om
a,
3
2
_
a
—
on
B
B

reniform in outline;
bracteoles slightly be-
low the sepals. Sikang.

ON
n
e
oO
p
ee
n
ae
fo)
&
2
At
©,
i=]
“eB
ig)
7

Turkestan, Tianshan.
_ 198. D. duhmbergit.

186. mee jose afghanicum Rech. f. Anz. Osterr. Akad. spon Math.-
Nat. 91: 74. 1954. 16, A.
Perennial from a woody root, forming clumps, each stem 4-10 dm. tall,
simple or few branched near the middle, rather evenly leafy to below the
inflorescence, loosely villous below, more densely so above with long spread-
ing white nonglandular hairs, mixed with shorter glandular ones in the
inflorescence; lower petioles to about 2 dm. long, villous, vaginate at the
base, the upper gradually shorter; leaf blades to 6 tt. long by 4 cm.
broad, the veins conspicuous beneath, loosely villous, 5-fid about two-
thirds of the way to the base, the segments cuneate-obovate, lobed-incised
into 3 principal parts, these with broad rounded coarse mucronate teeth;
inflorescence racemose or with 1 to few supplementary racemes below
rather laxly flowered; lowest bracts trifid, others lanceolate to ence tinear,
villous, 5-10 mm. long; pedicels divaricate, villous, 1-4 cm. long; brac-
teoles 2, near the flower, lance-linear, 3—4 mm. long ; sepals pale purplish
blue, densely villous, the upper sepal ovate, 12 mm. long, 6 mm. wide,

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 127

subacute, the spur subconic, 7-8 mm. long, 3-4 mm. wide at base ; lateral
sepals elliptic-oblong, 12 mm. long, 5 mm. wide, rounded at apex, hairy
on the whole outer surface; lower sepals elliptic, 13 mm. long, 5.5 mm.
wide, subacute; petals dark, the upper lamina almost straight, broad in
middle, 8.5 mm. long, slightly bidentate at the glabrous apex, the spur
9.5 mm. long; lower lamina almost straight, ovate, 5 mm. by 4 mm., cleft
to near the middle, yellow bearded at base, with scattered white hairs on
rest of the surface and margin, claw 5 mm. long; stamens 6 mm. long,
moderately dilated below, glabrous; anthers brownish, oblong, 1 mm. long;
follicles 3, hairy.

Type: Afghanistan, Minjan Pass, at 3800 m., July 26, 1937, W. Koelz
12719 (us); seen. Delphinium afghanicum differs from D. speciosum in
its longer hairs, entire bracts, and lax inflorescence.

187. Delphinium arcuatum Busch in FI. Cauc. Crit. III. 3: 71. 1902.
Fic. 16, B.

Stem 8-10 dm. tall, densely strigulose, somewhat flexuous, sulcate,
rather densely and evenly leafy; cauline petioles to 1 dm. long, not
vaginate at base, the blades more or less round-reniform in outline, 5-12
cm. broad, sparsely strigose above, densely so beneath, 5-parted to 1-2
cm. from the base, the segments broadly cuneate, 3-lobed distally and with
rather many divergent lance-oblong teeth or lobules 3-7 mm. wide; inflo-
rescence few branched below, the main raceme many flowered, to 3 dm.
long, more or less strigose to partly spreading-pubescent; lower bracts
leafy, palmatifid, others lanceolate, bluish, 1-2 cm. long, 3-4 mm. wide;
pedicels arched, with some glandular hairs, usually 3-4 cm. long, the
upper 1-2 cm., very densely covered with short spreading hairs; bracteoles
just beneath the flower, ovate-lanceolate or ovate, 6.5-12 mm. long, 2.5—
3.5 mm. wide, blunt; sepals violet blue, 16-21 mm. long, 6-12 mm. wide,
pubescent, the border white-ciliate; upper sepals ovate, 18-20 mm. long,
7 mm. wide, subacute, the spur spreading decurved, 16 mm. long, 4 mm.
wide at base; lateral sepals subovate, 20 mm. by 10 mm., rounded-obtuse;
lower sepals 18 mm. by 9 mm., more pointed; petals black, the upper
laminae almost straight, 11-12 mm. long, glabrous, bidentate, the spur 12
mm. long; lower laminae strongly oblique, oblong, 5.5 mm. by 3 mm.,
bearded at base, ciliate, bifid to about the middle into 2 lanceolate diver-
gent lobes, claw 6 mm. long; stamens 6—7 mm. long; anthers dark, over
1 mm. long; carpels 3, sparsely strigose.

Type: E. Caucasus, Dagestan, distr. Samur near Kurusch, 2400 m.,
Aug. 1, 1898, Th. Alexeenko (LE); seen.

188. Delphinium bracteosum Somm. & Levier, Acta Horti Petrop. 13:
25 3808 Fic. 16, C.

Erect, light green, with long scattered spreading hairs, the stem bee

somewhat angled, subglabrous above, ca. 3 dm. tall; leaves palmately

128 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

5-parted, 5-8 cm. in diameter, sparsely hairy on both sides, the sinuses
narrow, acute, the segments broad, the lower divergent, incised, the ulti-
mate divisions few, subobtuse, 6-8 mm. wide, the nerves prominent be-
neath: upper leaves tripartite; petioles not strongly dilated at the base,
those of the midcauline leaves equal to the blade, of the upper shorter to
none; inflorescence pyramidal-paniculate to simple and few flowered;
pedicels glabrous to hairy, the lower longer than the spurs; bracts elliptic
or lanceolate to spatulate, villous, 10-20 mm. long; bracteoles blue, near
the base of the flower, ca. 10 mm. long, 2 mm. wide; sepals intensely blue,
18-22 mm. long, 12-15 mm. wide, ciliate, otherwise glabrous to hairy:
upper sepal 19-22 mm. long, spur straight, 18-24 mm. long; other sepals
18-22 mm. long, 9-10 mm. wide; petals dark, the upper laminae almost
straight, 14 mm. long, subemarginate, glabrous; the lower laminae oblong-
ovate, 9.5 mm. by 7 mm., long-ciliate, yellow bearded at base, divided
about halfway into 2 unequal oblong lobes, claw 6 mm. long, appendicu-
late; stamens glabrous, 7-8 mm. long; follicles glandular; seeds trans-
versely lamellate-cristate.

Type: Zei in Ossetia, central Caucasus, coll. Lojka (LE?) ; not seen.

ILLUSTRATION. DEcHyY, Kaukasus, 3: pl. 8, fig. 1-6. 1907; according to
Index Londinensis.

SPECIMEN SEEN. South Ossetia: E. & N. Busch, Aug. 12, 1935, at 2700 m.
(BM).

189. Delphinium burmaense Munz, sp. nov. Fic. 16, D.

Delphinium stapeliosum Briihl var. shanicum Briihl in Briihl & King, Ann. Bot.
Gard. Calc. 5: 102. 1896. Based on Shan Hills, Collett (K); seen.

Perennial from a slender woody rootstock; stem slender, mostly dark,
with numerous spreading whitish or yellowish hairs ca. 2 mm. long, simple,
2-12 dm. tall, few leaved; basal leaves with slender villous petioles 5-25
cm. long, very vaginate at base, the blades subreniform, rather densely
villous on both surfaces, 3-15 cm. broad, 2-8 cm. long, deeply to shallowly
notched at base, 3—5-segmented about halfway to base, the segments
broad, oblong-ovate to -obovate, sometimes shallowly lobed, but more
usually with rounded-ovate coarse mucronate teeth 2-5 mm. deep, 2-10
mm. wide; upper leaves reduced; inflorescence a few-flowered, somewhat
secund raceme, villous, more or less glandular; lower bracts foliose, others
oblong-linear to lanceolate, 5-12 mm. long; pedicels ascending to erect,
1-5 cm. long; bracteoles subtending the flower, oblong-linear, 4-10 mm.
long; flowers violet, mostly 3-10, villous; upper sepal ovate, 12-14 mm.
long, 8-10 mm. wide, obtuse, the spur 15-20 mm. long, 2.5-3.5 mm. wide
at base; lateral sepals 10-17 mm. long, 7-10 mm. wide, oblong-obovate,
rounded at apex; lower sepals 9-14 mm. long, obtuse; petals dark, the
upper laminae slightly oblique, 8-10 mm. long, bidentate at the somewhat
bristly apex, the spur 15-18 mm. long: lower laminae oblong, bearded,
deeply bifid into lance-oblong lobes, the claw 5-7 mm. long; stamens 6-7
mm. long, glabrous; anthers dark, scarcely 1 mm. long; carpels 3, hairy.

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 129

Caulis tenuis, villosus, simplex, 2-12 dm. altus, cum foliis paucis; petioli
foliorum basilariu um 5-— 25 cm. longi, vaginati; laminae subreniformes, vil-
losae, 3-15 cm. latae, 2-8 cm. longae, 3- —5-partitae, segmentis latis, ob-
longo- ovatis vel -obovatis: inflorescentia racemosa, secunda; bracteae
oblongo-lineares vel lanceolatae, 5~12 mm. longae; pedicelli ascuadantes
vel erecti, 1-5 cm. longi; bracteolae florem subtendentes, oblongo-lineares,
4-10 mm. longae; flores 3-10, villosi, violacei; sepalum superiore ovatum,
12-14 mm. longum, 8-10 mm. latum, obtusum, calcar 15-20 mm. longum,
base 2.5-3.5 mm. crassum; sepala lateralia 10-17 mm. longa, 7-10 mm.
lata, oblongo-obovata; sepala inferiora 9-14 mm. longa, obtusa; laminae
Setslorum superiorum atrae, 8-10 mm. longae, bidentatae, setosae, calcar
15-18 mm. longum; laminae inferiores oblongae, barbatae, bifidae, unguis
5-7 mm. longum; stamina 6—7 mm. longa, glabra; carpella 3, hirsuta.

Type: Nam-Tamai Valley, Burma, ca. 3000 m., thickets in a steep
ae Sept. 6, 1937, F. Kingdon Ward 13170 (pa).

ANGE. Throughout Burma from 1500-4000 m. alt.

OTHER EXAMPLES. Burma: Shen Hills, Collett 964 (x); Shan States, Hilde-
brand (kK); Melville 43 (gsm); Upper Adung Valley, no. of Lung Sa, Kingdon
Ward 10115 (pm); Mindat, Kingdon Ward 22505 (pm, F); Taunggyi, Dickason
9052 (A); Nam Tami Valley, Kingdon Ward 3351 (gm); Burma-Tibet frontier,
Kingdon Ward 9819 (sm, F); Taunggyi crags, Robertson 15 (kK); Sirhoi, King-
don Ward 18120 (a, Bm). A specimen from Assam (Orka La, Kingdon Ward
14308, BM) seems to belong here; it is rather depauperate.

Delphinium burmaense differs from D. stapeliosum from Nepal in its
smaller flowers, presence of glandular hairs in the inflorescence and the
spreading, not retrorse, hairs on the stems.

190. Delphinium buschianum Grossheim, Trud. Azerb. Otdyel. Za-

kavk. Fil. Akad. Nauk. U.S.S.R. Sect. Bot. 1: 51. 1933. Fic. 16, E.

Delphinium linearilobum var. hirticaule Grossheim, Fl. Cauc. 2: 99. 1930, fide
Nevski in Fl. U.S.S.R. 7: 143. 1937.

Stem ca. 7.5 dm. high, somewhat angular below, with some divergent
retrorse simple white hairs 0.75 mm. long, foliose; inflorescence rather
glabrous; petioles scattered hairy, somewhat vaginate; cauline leaves
rounded-cordate to almost round in outline, glabrous on both surfaces or
weakly pubescent on veins beneath; laminae divided into 3 lobes 3-6 mm
from base, the middle lobe rhombic-lanceolate, somewhat acuminate, the
lateral lobes divided into 2 so that there are 5 resultant gaint nina
into acute or acuminate lanceolate teeth or lobules 3-6 mm. 3-15
mm. long; inflorescence several branched, rather compact, the axes purplish,
glabrous, to ca. 2.5 dm. long; bracts purplish, ca. 1 cm. long, lance-linear,
rarely ciliate near the tip; bracteoles linear, glabrous or slightly ciliate,
placed just below the flower, 5-7 mm. long; sepals dark blue or violet-
blue, glabrous on outer surface, with white felt inside in the lower half of
long entangled hairs or at least with a few hairs, sometimes weakly ciliate

130 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Figure 16. ~— Group IX. Petals drawn, the upper rs the
lower clawed. A. D. afghanicum; upper petal blade 8.5 mm. long, spur 9.5

lower blade 5 mm. long, claw 5 mm.; drawn from Koelz 12719 (ws). .
arcuatum ; upper blade 11 mm. long, spur 11 mm.; lower blade 5.5 mm. Pie
cl .) draw fo Mg ke TYPE (L bracteosum; upp

mm
amina 14 mm. long, spur 17 mm.; ae lamina 9.5 mm. long, c
— N. Bus M). D.

D. burmaense ; ia lamina 9 mm. long,
spur ia ey lamina s mm. lon , claw 5.5 mm ; drawn from King-
0 (BM D.

on: upper lamina 12 mm, lon 2
mm.; gets from Abolin 3193, TYPE (L E), :
ees m. lon r 16 mm.; lower lam mina 4 mm. ia ng, claw 5 mm.; drawn from
Contes & Bulley 2294, TYPE (E). I. D. corymbosum; upper lamina 11.5 mm.

long, spur 10.5 mm.; lower lamina 7 mm. long, claw 7 mm.; drawn from TYPE

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 131

ovate, 12-15 mm. long, 7-8 mm. wide, obtuse; spur 10-15 mm. long, often
hooked at apex; petals black, the upper laminae somewhat oblique, 9 mm.
long, bidentate at the glabrous apex, spur ca. 10 mm, long; lower laminae
sparsely bearded, strongly ciliate, very oblique, obovate, 5 mm. by 3 mm.,
bifid over halfway into lance-oblong divergent lobes, claw 5 mm. long;
follicles 3, erect, 10 mm. long, 3 mm. wide, glabrous except along sutures.

Type: Southern Caucasus, shores Lake Cevan, Karjagin & Safiev, Aug.
15, 1931 (Kk, LE), seen.

DISTRIBUTION. The species grows on igneous rocks below 2100 m.,
Soviet and Turkish Armenia, and is represented by Davis 32562, in part,
(K) from Kars Prov., T urkey.

191. Delphinium chenii W. T. Wang, Acta Phytotax. Sinica 6: 369.
1957. Fic. 16, F.

Delphinium albomarginatum Chen, Bull. Fan Mem. Inst. Biol., Peiping (n.s.)
170. 1948; not Simonova, 1924.

Perennial from a slender rhizome; stem 3—4 dm. tall, simple or short
branched above, 3-4 mm. thick, striate, more or less retrorse-pubescent,
equably and fairly densely leafy; petioles slender, 3-5 cm. long, scarcely
dilated at the base, somewhat retrorse-strigose; blades pentagonal-orbicular
in outline, 2-5 cm. in diameter, subglabrous to sparsely pubescent, pallid
beneath, parted to near the base into cuneate-obovate segments, these 3-
toothed or -lobed, or with a few additional teeth, the ultimate parts lance-
ovate, rather short, obtuse to acute; inflorescence racemose or narrow-
paniculate, retrorse-pubescent with white and yellow hairs, some rather
long; flowers several; bracts largely reduced divided leaves, hairy; pedicels
erect, 1-3 cm. long, hairy; bracteoles lance-linear, 5-10 mm. long, pubes-
cent, placed slightly below the flowers; sepals strigose and short-hairy,
blue with white margins (especially on the 2 lower pairs) ; upper sepal
broadly ovate, 15 mm. long, 12 mm. wide, somewhat apiculate, the spur
22 mm. long, 4 mm. wide at the base, spreading decurved, narrowed to a
subcylindric tip; lateral sepals pubescent on midrib, oblong-ovate, 14 mm.
long, 10 mm. wide, rounded at apex; lower sepals rhombic-obovate, 15
mm. long, 9 mm. wide, rounded-obtuse; petals dark, the upper laminae

sent b eee icaule; upper lamina 10 mm. long, spur 15 mm.;
ower antag s ole peo . ao et from Bernhardi Herbarium (mo
1)

a mm. long, Sd 5 mm.; ae Haw
upper lamina 10 mm. long, spur a. , ;
Poa from Pohle in 1902, Type (LE). M. D. duhmber git; te lamina 10
mm. long, spur 16 mm.; lower lamina 4.5 mm. long, claw 5.5 mm.; drawn fro
Regel in 1887 (us). N. D. dyctiocarpum; upper lamina 9 mm
mm.; lower lamina 4.5 mm. long, claw 6 mm.; sg anit
O. D. elatum; upper lamina 10 mm. long, spur 11 mm

.; lower la .
long, claw 5 mm.; drawn from Korshinsky in 1891, type of D. korshinskyanum
(LE).

132 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

straight, 13 mm. long, 3 mm. wide, glabrous, somewhat erose at the
rounded summit, spur 22 mm. long; lower laminae oblique, oblong, 6 mm.
long, 4 mm, wide, cleft halfway into ciliate oblong lobes, with patch of
yellow hairs at base below the cleft, claw 6.5 mm. long; stamens 6 mm.
long, strongly dilated at base, glabrous; anthers dark, 1.2 mm. long;
carpels 3, densely yellow-hairy.

Type: Southern Sikang, Taucheng, Hsien, Wongtula, 3000 m., T. 2s
12809 (PE), not seen; photo. (£), seen. A sheet at A has the same number,
but says “Yunnan” and lacks data.

192. Delphinium confusum Popov in Komarov, Fl. U.S.S.R. 7: 136,
qa. 1937, Fic. 16, G.

Delphinium duhmbergii Huth, Bot. Jahrb. 20: 402. 1895, pro parte.
Delphinium speciosum Huth, loc. cit. 415, pro parte.

Stem 3-6.5 dm. tall, densely pilose, the hairs in the lower part 1.5-2
mm. long, the upper part densely soft-pubescent, not glandular; stem
rather equably but not densely leafy; petioles to 2 dm. long, dilated at
base, long-pilose; leaf blades round-reniform, densely pubescent or more
or less pilose on both sides, trifid, the middle segment oblong-obovate, cut
into 3 lobes each with 3—4 lance-ovate teeth to 1 cm. long and 8 mm. wide;
lateral segments of leaf cut into parts much like those of the middle one;
raceme dense, sometimes branched at the base, many flowered; bracts
broadly lanceolate or lanceolate, entire or rarely the lower trisect, obtusish,
ca. 1 cm. long, densely and short pilose; pedicels 5-15 mm. long, with
short spreading hairs; bracteoles broadly ovate, obtuse or subobtuse, 5.5—8
mm. long, 3-5 mm. wide, pubescent; flowers dark violet; sepals ovate or
obovate, obtuse, 10-15 mm. long, 5-8 mm. wide, densely pubescent with
short hairs without, almost glabrous within; spur spreading horizontally,
incurved near the apex, 12-15 mm. long, 2.5-3 mm. wide at base; petals
black, the upper laminae almost straight, 12 mm. long, emarginate and
bristly at apex, the spur 15 mm. long; lower laminae oblong-obovate, long-
ciliate, cleft almost halfway, ca. 7 mm. long, the claw appendiculate, 5.5
mm. long; carpels 3, densely pubescent.

Type: Asia Media, m. Akbach-tau, Aug. 12, 1921, Abolin 8193 (LE);

seen. This species keys out near D. flecuosum from farther west, but has
much broader bracteoles.

193. Delphinium cooperi Munz, sp. nov. Fic. 16, H.

From a slender woody rootstock, the stem very slender, 1—4.5 dm. tall,
with soft spreading hairs throughout, especially above; leaves few, mostly
near the base, 1-2 reduced ones above; petioles spreading-villous, very
slender, 2-10 cm. long, somewhat vaginate at base; blades pentagonal-
subreniform in outline, mostly 2-5 cm. broad, 3- or 5-fid about 2/3 their
length into broad obovate segments with some scattered long hairs on both

1968} MUNZ, ASIAN SPECIES OF DELPHINIUM 133

surfaces and shallowly 2-3-lobed, each lobe with 2—-few ovate-mucronate
teeth to ca. 5 mm. long and 4 mm. wide; inflorescence open, simple or
1-branched, each with 1-3 flowers; lower bracts compound or trifid, 5-15
mm. long, the upper oblong-elliptic, 5-8 mm. long; pedicels 2~5 cm. long,
spreading-villous and with some inflated glandular hairs; bracteoles ellip-
tic, 4-6 mm. long, situated above the middle of the pedicel; flowers purple-
violet with yellow hairs; upper sepal obovate, rounded above, 12-13 mm.
long, 7-8 mm. wide, the spur decurved, 14-16 mm. long, 3 mm. wide at
the base, rather acute; lateral sepals elliptic-ovate, 12-13 mm. by 9-10
mm., obtuse; lower sepals 13 mm. by 8 mm., subacute; petals black, the
upper laminae quite oblique, entire, glabrous, 9 mm. long, 3 mm. wide,
the spur ca. 13 mm. long; lower laminae oblong-ovate, ca. 4 mm. by 3
mm., bearded and long-ciliate, erose but not bifid, the claw 6 mm. long;
stamens 5—6 mm. long, glabrous; anthers dark, 1 mm. long; follicles 3,
erect, 13-14 mm. by 2.5 mm., spreading-pubescent, the styles 2.5 mm.
long.

Caulis tenuis, 14.5 dm. altus, patulo-hirsutus, supra subglandulosus;
folia pauca, subbasilaria; petioli villosi, valde tenues, 2-10 cm. longi, base
subvaginati; laminae pentagonali-subreniformes, 2-5 cm. latae, 3—5-fid,
segmenta 2—3-lobata, supra et infra sparse hirsuta; inflorescentia laxa,
integra vel 1-ramosa, in quoque parte cum 1-3 floribus; bracteae inferiores
trifidae, 5-15 mm. longae, superiores oblongo-ellipticae, 5-8 mm. longae;
pedicelli 2-5 cm. longi, villosi et glandulosi; bracteolae ellipticae, 4-6 mm.
longae; flores purpureo-violacei cum pilis aureis; sepala 12-13 mm. longa,
7-10 mm. lata, calcar 14-16 mm. longum, base 3 mm. latum; petala atra,
laminae superiores obliquae, glabrae, integrae, 9 mm. longae, calcar 13
mm. longum; laminae inferiores oblongo-ovatae, 4 mm. longae, barbatae,
ciliatae, erosae, unguis 6 mm. longus; stamina glabra; follicula 3, erecta,
13-14 mm. longa.

Type: Bhutan, Tonjsa, Oct. 3, 1914, R. E. Cooper, collector for Bulley
2294 (£) at 2600 m.

OTHER COLLECTIONS. Bhutan: Ridang Angduphorang, Cooper for Bulley 2024
(E); Chendebi, Ludlow & Sherriff 3519 (BM, E), 17065 (BM), 19578 (BM).

A collection from Nepal [Lulo Khola, Polunin, Sykes & Williams 35 18,
(BM)| seems near Delphinium cooperi, but has the lower petals divided.
The proposed species suggests D. stapeliosum, but that species lacks
glandular hairs.

194. Delphinium corymbosum Regel, Acta Horti Petrop. 7: S47.

1880; Gartenflora 1881: 23, 24. pl. 1059. 1881. Fic. 16, I.

? Delphinium corymbosum var. baicalense Huth, Bot. Jahrb. = 409. 1895.

Based on Karelin & Kiriloff 1841 (w) as D. intermedium var? I have not

seen the Vienna specimen, if it is Karelin & Kirilof no. 11 64, it is D. elatum.

Delphinium umbellatum Regel ex Nevski in Komarov, Fl. U.S.S.R. 7: 155.
1937, in synon.

134 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Stem 3-6 dm. tall, retrorse-pilose, branched above forming a dense
corymbose panicle, the stem leafy throughout; lower petioles dilated at the
base, 1-1.5 dm. long, upper ones shorter; blades of leaves hairy on both
sides, palmately 5-lobed, the lobes broadly cuneate, 3-lobulate at apex, the
lobules coarsely crenate-dentate, the teeth callus-tipped; upper cauline
leaves short petioled or sessile, palmately 3- or 5-lobed; branchlets of
inflorescence with linear entire herbaceous bracts 10-15 mm. long; pedicels
3-7 cm. long; bracteoles linear, 10 mm. long; flowers 4-5 on side branch-
lets crowded in subumbellate fashion at the apex of the dense terminal
raceme; sepals subequal, ovate or lance-ovate, 13-17 mm. Jong, 6-10 mm.
wide, pale violet, greenish on back, hirsute, cucullate at apex; spur 8-10
mm. long, 3-4 mm. wide at base, hirsute, more or less incurved at the
obtuse apex; upper petals blackish, the laminae emarginate, ca. 11 mm.
long, glabrous, the spur 10.5 mm. long; lower laminae oblong-ovate, 7 mm.
long, deeply cleft into oblong lobes, bearded below the lobes, long-ciliate,
claw 7 mm. long; follicles 3, densely pubescent.

Type: Grown from seeds from alps of east Turkestan, sent by Dr.
Akegel, June, 1881, ex horto bot. Petropolitano (L£); seen.

ILLUSTRATION. Gartenflora pl. 1059. 1881.

195. Delphinium crassicaule Ledeb. Fl. Rossica 1: 62. 1841. Fic. 16, J.
Delphinium caucasicum var. chinense Regel, Mem. Acad. Pétersb., VII. 4:
10. 1861

. Based on a collection from Chinese Mongolia, Herb. Fischer;
not seen.

Stem to 1 cm. thick in the type, leafy largely below, leafless above,
pilose; leaves round in outline, 5- to 7-parted to well above the base, sub-
glabrous to scattered-pilose, especially beneath, the segments cuneate,
broad, incised to near the middle into lanceolate, acuminate lobes; raceme
rather dense, many flowered; pedicels more or less divaricate, up to more
than twice the length of the flower, more or less pilose; bracts lance-linear,
long-pilose, entire, to 10 mm. long; bracteoles 2-3, lance-linear, somewhat
shorter, located near the base of the flower; sepals blue, subglabrous
without, the upper ovate, 12 mm. by 6 mm., acute, the spur slightly de-
curved, 17 mm. by 3 mm., narrowed rather rapidly to a rather pointed
apex; lateral sepals rhombic-obovate, 12 mm. by 5 mm., the narrow apex
somewhat recurved; petals dark, the upper laminae acute, ciliate, almost
straight, ca. 10 mm. long, the spur ca. 5 mm. long; lower laminae almost
straight, 5 mm. long, 2-toothed apically, hairy in distal part, claw 5 mm.
long; stamens 6—7 mm. long, glabrous; anthers dark, oblong, 1 mm. long;
follicles 3, glabrous.

Type: Eastern Siberia, between Wilnick and Olekminsk, Kruhse (1");
seen. The type specimen now has no flowers. Three sheets of the Bern-
hardi Herbarium (mo) seem to be this species; one labeled Ex Museo
Berolinensi, June, 1818. Nevski (Fl. US.S.R. 7: 154. 1937) cites this

species as a synonym of D. crassifolium, but the leaf lobes are much
sharper.

1968] MUNZ, ASIAN SPECIES OF DELPHINIUM 135

196. Delphinium crassifolium Schrad. ex Spreng. Gesch. Bot. 2: 201.
1818, nomen; Ledeb. Fl. Rossica 1: 62. 1841. (A var. compactum
Malysch was later printed on herbarium labels with a Latin diag-
nosis). Fic. 16, K.

? Delphinium crassifolium var. tangutica Maxim, FI. Tangut. I: 22. 1839;
not known to me.

Perennial from a woody root; stem simple, scapose, slender, with scat-
tered long spreading hairs and shorter glandular ones above, 3-7 dm. tall;
petioles 7-15 cm. long, spreading-hairy, vaginate at base; leaf blades
rounded angled in outline, 5-15 cm. broad, sparsely hairy on both sides,
paler beneath, 5- or 7-parted into broad cuneate trifid segments, these
with more or less oblong-ovate teeth or shallow lobes 4-10 mm. broad;
racemes strict, 2-15 cm. long, several flowered; bracts lanceolate or nar-
rower, to 10 mm. long; pedicels appressed to the axis, slender, pubescent,
1-5 cm. long; bracteoles lance-linear, 3~6 mm. long, from the middle of
the pedicel to near the flower; sepals blue, scattered-pubescent; upper sepal
14-17 mm. long, ovate, obtuse, spur about as long, straight to somewhat
curved; lateral sepals oblong, 14-15 mm. long, rounded at apex; lower
sepals 16-18 mm. long, obtuse; petals black, the upper laminae almost
straight, acute, entire, 10 mm. long, glabrous; lower laminae 5 mm. by
2 mm., cleft about 2 mm., with long white hairs on both sides toward the
apex and on the margin, claw 6 mm. long, broad; stamens 6-7 mm. long,
the base of the filaments much widened, glabrous; anthers dark, | mm.
long; follicles 3, glabrous, reticulate; seeds squamate, 1.5 mm. long.

Type: Krasnijarsk, Turkestan?, Herb. Ledebour (LE); seen.

ILLustration, GMELIN, FI. Sibirica 4: pl. 79. 1769.

REPRESENTATIVE SPECIMENS, In alpibus Sajanensibus, Radde (GH); var.
compactum Malyschev, 2300 m., Mts. Sajanenses, Jugum Tunkenense Aug. 14,
1858 (BM, E, K); var. stubendor fii, Sibiria orientalis, leg. Stubendorf (k, In type
cover) I have no record of publication; crassifolium, Sibir. media, in herbario
Petropolitani (kK); Mts. Sajan, Komarov in 1902 (P); Issyk-Kul, Songaria,
Krasnow in 1886 (P).

[Re 146,

197. Delphinium cryophilum Nevski in Komarov, Fl. U.S.S HAE L

724. 1937
Delphinium elatum var. intermedium f. hirsutum Pohle in herb.

Perennial, the stem 2-6.5 dm. tall, to almost 10 mm. thick below, re-
trorsely soft hairy, the hairs 1—2.5 mm. long, rather equably leafy ; petioles
5-10 cm. long, strongly dilated at base; laminae with long white hairs on
both surfaces, to ca. 1 dm. broad, round-cordate in outline, parted almost
to base into rhombic-cuneate parts which are more or less plainly shallowly
3-lobed in their distal portion, then with few deep teeth or lobules age
oblong, to almost 1 cm. long, 2-4 mm. wide, acute); central raceme 2—
dm. long, thick-stemmed, many flowered, with 1 or 2 few eo
Slender racemes near the base; bracts 1-3 cm. long, narrowly linear to

136 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

broadly lanceolate; pedicels spreading-hairy, erect, 1-4 cm. long; brac-
teoles linear-subulate, 4-5 mm. long, near the flower; sepals violet-blue,
glabrous except for a few hairs near the apices, the upper sepal ca. 12 mm.
by 6 mm., attenuate at tip, the spur spreading, 13-14 mm. long, 3.5-4 mm.
wide at base, obtuse; lateral sepals obovate, 12-13 mm. long, 7-8 mm.
wide, round-obtuse; lower sepals 11-12 mm. by 4-5 mm., attenuate; petals
black, the upper laminae oblique, 10 mm. by 3 mm., short-bidentate,
glabrous, the spur 11-12 mm. long; lower laminae yellow-bearded at base,
4-5 mm. long, bifid halfway into lance-oblong ciliate lobes, claw 5 mm.
long; stamens 5 mm. long; carpels 3, glabrous.

Type: “In decliviis argillosis herbosis insulae Kolgujev, lg. Pohle, Aug.
23, 1902 (LE); seen.

198. Delphinium duhmbergii Huth, Bull. Herb. Boiss. 1: 330. pl. 16
(flower). 1893. Fie. 16, M.
Delphinium iliense Huth, Bot. Jahrb. 20: 402-3. 1895. No type stated. From
LE was loaned as type: “Prope lacum Turnkul, flora iliensis, Arassno in
1886,” originally identified as D. caucasicum var. hirsutum. D. iliense vat.
pubiflorum (cited by Huth) was, Méngoté, Regel, 1879, seen (K); D. iliense
var. angustatum (Huth, p. 403), no cited material seen; D. iliense var.
macrocentrum (Huth, p. 403), a Przewalski, 1880, specimen from Kansu,
China cited, not seen or identified; D. iliense var. hispidum (Huth, p. 403),

no cited material seen.

Delphinium englerianum Huth, Bot. Jahrb. 20: 418. 1895. First specimen
cited was Turkestan, Hissar, “in valle fluminis Jaquob,” Regel, 1883 (LE,
not seen, isotype E, seen). Huth had 3 vars.: var. incisum (p. 418), no type
cited, opinion expressed it might be D. incisum Wall., which it cannot be if
related to D. duhmbergii; var. hoffmeisteri Klotzsch, Reise Wald. 132.
1862; no specimen cited and I do not know the plant; var. simplex Huth
(p. 418) which I cannot place. :

Delphinium turkestanicum Huth, Bot. Jahrb. 20: 420. 1895. If the first of
two specimens cited is taken as type (Aryslyn, 3000 m., Regel, 1879 (LE)),
it belongs here, but has a few short bristles on upper petal lamina. :

Perennial from a woody rhizome, stem slender, 1-8 dm. tall, hirsute

below, tending to be glabrous to sparsely hirsute above, mostly leafy at the
base or in lower half; lower petioles hairy, 5-20 cm. long, dilated at base,
upper much shorter ; leaf blades round-pentagonal in outline, subglabrous
to hairy above, pilose beneath on veins and margin or on whole surface,
3-6 cm. broad, divided to below middle into cuneate, obovate segments
1—2.5 cm. broad, these shallowly lobed or coarsely toothed, the teeth ovate,
len cauline leaves remote, the uppermost reduced, subsessile; in-
‘ : ; :

rive ap erento _ glabrous to pilose; bracts trifid or mostly

’ m. long, ca. 2 mm. wide; pedicels ascending,
1-3.5 mm. long; bracteoles lanceolate, ciliate, membranous, bluish, 5—7
mm. long, subtending the fl : ! ,

g ower; sepals mostly blue, glabrous to pubescent;
— upper sepal ovate, 13-15 mm. long, 8-10 mm. wide, acutish, the spur
almost straight, about as long as sepals, 3 mm. wide at base, narrowed to

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 137

a sharp point; lateral sepals round-ovate, 13-15 mm. long, 10-11 mm. wide,
rounded-obtuse; lower sepals 12 mm. by 7 mm., acute; petals almost
black, the upper laminae oblique, glabrous or with a few short bristles on
the face, bidentate, ca. 10 mm. long, the spur 16-17 mm. long; lower
laminae oblique, ovate, 4.5 mm. by 3 mm., cleft ca. 2.5 mm. into somewhat
erose pra with very long cilia and very long hairs at base of lamina, claw
5.5 mm, long; stamens 6-7 mm. long, glabrous; anthers dark, 1.2 mm
long; follicles 3, glabrous to hirsute, to 10 mm. long; seeds triquetrous,
winged.

Type: Altai Mts., Siberia, Dukmberg in 1881 (8), not seen. At Kew
are 3 sheets “teste Huth”: Turkestania, Dschungarischer Alatau, Regel
in 1878, and Dschagartai, Regel in 1877. These I would use as representa-
tive of the species. The third sheet is the Méngét6 collection cited above
under the discussion of the name iliense.

REPRESENTATIVE SPECIMENS. Asia Media: Lipsky 1783 (BM); Turkestan: Regel,
July 18, 1877 (Le); Alatau, Karelin & Kiriloff 1164 (ny, p); Issinsul, Regel,
Aug., 1877 (K); Tianschan, Merzbacher 723 (us); Terskei Alatau, Artemjev
4266 (BM, E, K); Koksu Valley, Littledale (x).

199. Delphinium dyctiocarpum DC. Syst. 1: 360. 1817. Fic. 16, N.

Delphinium dictyocarpum Steud. Nom. ed. II. 1: 488. 18
oot ciliatum Stev. var. dictyocarpum Huth, Bot Jahrb. 20: 404.
895.

Delphinium dyctiocarpum var. glaberrimum Trautv. Bull. Soc. Nat. Mosc. 33:
860. I have not seen type. The variety is reduced to synonymy by
sia in Komarov, Fl. U.S.S.R. 7: 159. 1937.

Perennial from a woody root, the stem 6—15 dm. tall, erect, leafy, robust,
glabrous or nearly so, often purplish, mostly simple; lower petioles to ca.
2 dm. long, scarcely dilated at the base, the upper much shorter; leaf
blades rounded-pentagonal in outline, the upper ca. 4 cm., the lower ca. 15
cm. broad, subglabrous above except for a few appressed hairs on the
veins, strigose or pilose beneath especially on the veins, 3- to 7-parted to
within 1 or 2 cm. of the base, the lower part of the segments subcuneate,
entire for 1-3 cm. then widening into broad parts that are pinnately incised
into lanceolate teeth or lobes to almost 2 cm. long and mostly 2-4 mm.
wide, subacuminate; inflorescence a dense raceme or open panicle of
racemes, many flowered, nearly or quite glabrous; bracts mostly entire
(the lowest trifid), subsetaceous, 5-15 mm. long; pedicels divaricate to
suberect, 1-3 cm. long; bracteoles lance-linear to setaceous, 3-7 m m. long,
placed well below the flower; sepals violet-blue, glabrous or ne
minutely strigulose; upper sepal 10-12 mm. long, ovate, acutish, the spur
straight, 12-20 mm. long, 2.5—3 mm. wide at base, narrowed to a slender
apex; lateral sepals 12 mm. long, 7 mm. wide, elliptic-oblong, with rounded
apex; lower sepals 12-14 mm. long, 6 mm. wide, more pointed; petals
dark, the upper slightly oblique, the laminae 8-9 mm. long, glabrous,
bidentate, the spur 12-24 mm. long; lower laminae strongly oblique, hairy,

138 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

ovate-oblong, 4.5 mm. long, cleft half its length, the claw broad, 6 mm.
long; stamens 5—6 mm. long, glabrous; anthers dark, 1 mm. long; follicles
3, glabrous (ciliate on suture when young), suberect, 10-14 mm. long, 4-5
mm. wide, the styles an additional 3.5 mm.; seeds brown, angled, ca. 2 mm.
long, winged on the angles.

Types: A cultivated specimen from Siberia (G?); not seen.

ExAmpies. Cultivated: Ithaca, N. Y., Dress 8603 (pH); Cambridge, Mass. in
1901 (BH); Kew, Nelmes 804 (BH); ex herbario horti Petropolitani, Karkoraly
in 1862 (mo, Ny). Wild: Altai, Schrenk (GH, K); Altai occident., Titov (NY);
Altai borealis, Burdakova in 1931 (Ny); Songoria, Schrenk, ex herb. Trautv.
July, 1843 (x).

200. Delphinium elatum L. Sp. Pl. 1: 531. 1753. Fic. 16,0; 17, A, B.

Delphinium palmatifidum DC. Syst. 1: 358. 1817, and probably vars. hispidum
and glabellum of same reference and based in part on Gmelin, Fl. Sibirica 4:
pls. 79 and 75, respectively. This reference and the next three are treated

synonyms of D. elatum by Nevski, Fl. U.S.S.R. 7: 147. 1937.

Delphinium glabellum Turcz. Bull. Soc. Nat. Mosc. 10: 56. 1837, nomen.

Delphinium alpinum Huth, Bot. Jahrb. 20: 405. 1895, pro parte, not same as
Waldst. & Kit. Pl. Rar. Hung. 3: 273. 1812, European. Possibly D. alpinum
vars. ajanense Huth and productum Huth, loc. cit. 406.

Delphinium discolor Fisch. ex Sweet, Hort. Brit. ed. II. 9. 1830, nomen.

Delphinium elatum subsp. palmatifidum (DC.) Sergievsk, Animadvers. Syst.
Univ. Tomsk 1930: 3 and vars. glabellum (DC.) and hispidum (DC.), and
glabrum Sergievsk.

? Delphinium elatum subsp. subglabrum Ledeb. Fl. Ross. 1: 64. 1841.

? Delphinium elatum subsp. intermedium (Soland.) Fleisch & Lindm. Bull.
Acad. Polon. Sci. & Lett. Cl. Math. Nat. 1933: 96. 1934.

Delphinium intermedium DC. Prodr. 1: 55. 1824. Based on a European type.

? Delphinium elatum var. longicalcaratum Huth, Bot. Jahrb. 20: 398. 1895.
Type, “Sibiria pr. fl. Jenisei,” Lessing (8); not seen.

? Delphinium atropurpureum Pall. Reise Statth. Russ. Reich. 1: 134. 1771.
Apparently a name used later for cultivated plants only.

Delphinium korshinskyanum Nevski in Komaroy, Fl. U.S.S.R. 7: 153, 724.

937. , “Inter fl. Zeja & Bureja, near Ivavovskoje,” Korshinsky, July
11, 1891 (LE); seen.

? Delphinium elatum var. sericeum W. T. Wang, Acta Bot. Sinica 10: 81.

1962. Said to be glandular puberulous in the inflorescence, from Sinkiang

province, China. I have seen no material and doubt its relationship to the
D. elatum complex.

Perennial from a woody rhizome, stem 5-20 dm. tall, glabrous to
spreading- or retrorse-villous, rather robust, simple below, simple or
branched in the inflorescence, leafy, the leaves only gradually reduced up-
ward; basal petioles to 2 dm. long, membranous-dilated at base, glabrous
to hairy, the cauline shorter and scarcely dilated below; blades mostly
5-15 cm. broad, more or less rounded-pentagonal in outline, subglabrous to
hairy, mostly 5- or 7-parted to within about 1 cm. from the base, the
segments broad, cuneate at base, ovate, incised or deeply toothed to lobed

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 139

with 2~3 coarse terminal teeth 8-20 mm. long, 8-10 mm. wide; inflores-
cence of subglabrous to hairy racemes solitary at end of each stem, the
flowers many, crowded; bracts leaflike and divided or the upper entire,
linear, to ca. 1 cm. long; pedicels ascending, mostly 1-3 cm. long; brac-
teoles linear, 4-6 mm. long, near to flower or above the middle of the
pedicel; flowers pale blue to dark blue, the sepals glabrous or ciliate or
more or less pubescent without; upper sepal ovate, 12-16 mm. long,
subobtuse, the spur about as long or longer, stout, 3~3.5 mm. wide at
base, straight or decurved, blunt; lateral sepals mostly 12-16 mm. long,
8-10 mm. wide, rounded at apex; lower about as long, narrower, obtuse;
petals almost black or of same color as sepals, the upper laminae suberect,
ca. 8 mm. long, bidentate at the glabrous or sparsely bristly apex; lower
laminae oblique, long-bristly, 4 mm. by 2 mm., shallowly bilobed, the claw
6 mm. long; stamens 5 mm. long, glabrous, strongly dilated; anthers dark,
1 mm. long; follicles 3, glabrous, 12-15 mm. long, the styles an additional
3 mm.; seeds dark, winged on angles, 2—3 mm. long.

Type: Specimen in Hort. Cliff. Herb. (Bm) from Silesia; seen.

RANGE. Widely distributed in mountains from central Europe across
northern Siberia.

ItLustrations. Hect, Ill. Fl. Mittel-Eur. 3: pl. 114. 1909; Bot. Reg.
pl. 1963. 1837.

COLLECTIONS SEEN. Siberia orient.: Dr. Merk, Herb. Pallas (pm); Sibiria
subalpina, Jacquin 1773 (BM); W. Siberia: Mamlev 211 (xk, us), Stabindorff (¥);
Lake Baical, Radde (x, Pp); Iter Ircutense, Kusnezow 885 (micH), Irkutsk,
Besser (kK); Transbaicalia, Enander 7-19-1914 (£); Mongolia, Ikonnikov-Galitz-
ky 2915 (mo); Soongaria, Karelin & Kirilof 1168 (BM, K in part, Pp); Altai,
Ledebour (x, p), Borgand (cH); Ochotsk Sea, C. Wright (GH, K, NY, US);
Dahuria, Turczaninow in 1828 (uc), Fischer in 1842 (P); Jenissei, Tolmatchew
174 (cH), Brenner in 1876 (us).

A variable species as here recognized. Pawlowski (Fragmenta Flor. &
Geobot. 9: 429. 1963) recognizes subspecies intermedium and others, but
I do not feel that I understand D. elatum well enough to do so. I have seen
cultivated material under many names: D. formosum, D. caucasicum, D.
decorum, D. bicolor, D. intermedium and D. cuneatum.

201. Delphinium elbursense Rech.f. Anz. Osterr. Akad. Wiss. Math.-
Nat. 88: 223. 1951. Fic. 17, C.
Perennial from a rhizome; stem to 6 dm. tall, simple or short branched
in the inflorescence, sulcate-striate, with long soft spreading hairs below,
deflexed-strigose above, leafy to inflorescence; leaves rather numerous,
petioles more or less villous, the lower to 1 dm. long, somewhat dilated at
the base, the upper shorter; leaf blades sparsely strigose or with looser
hairs on veins, broadly round-cordate, the lower to 12 cm. across, gradually
smaller up the stem, 5-parted to within 1 or 2 cm. from the base into
cuneate-obovate segments, these incised into lance-oblong teeth or lobes
mostly 2-4 mm. wide, obtuse to acute, thickened at apex; inflorescence a

140 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

rather open raceme 1-3 dm. long, with or without supplementary branches
below, densely strigose; bracts trifid or entire and lanceolate, 1—2 cm. long,
pubescent; pedicels 1-3 cm. long, ascending; bracteoles 2, lance-linear,
near the flower; sepals deep blue, crisp-villous without, with some long
loose hairs within; upper sepal ovate, 15-17 mm. long, 10 mm. wide, obtuse,
the spur usually slightly curved, 18-20 mm. long, ca. 4 mm. wide at base,
narrowed to an acute apex; lateral sepals broadly rhombic-obovate, 20
mm. long, 15 mm. wide, rounded-obtuse; lower sepals elliptic, 18 mm. b

10 mm., acute; petals dark, the upper laminae bidentate, slightly oblique,
glabrous, 11 mm. long, the spur 11-12 mm. long; lower laminae somewhat
oblique, 5 mm. long, 4 mm. wide, deeply cleft into 2 oblong-lanceolate
divaricate, bearded and ciliate lobes, claw 6 mm. long, not appendaged ;
stamens 6-7 mm. long, glabrous; anthers dark, 1-2 mm. long; follicles 3,
pubescent.

Type: Iran, prov. Gorgan (Asterabak), Mt. Shahvar near Hadjilang,
2400-2600 m., K. H. & F. Rechinger 6093a, July 27, 1948 (w) not seen;
isotypes (BH, MO, UC, US) seen.

Possibly too near D. speciosum, but perhaps a local entity recognizable
by the pubescence.

Uae. freee. 3. 355. 1937

202. Delphinium elisabethae N. Busch, Acta Inst. Bot. Acad. Sci.
Fic. 17, D

Stem perhaps 1 m. tall, retrorse-strigulose and with some spreading
hairs, these more numerous and mixed with glandular ones in the inflo-
rescence, stem leafy; cauline petioles 1-6 cm. long, scarcely dilated at base;
leaf blades 4-10 cm. broad, green and sparsely strigulose above, paler and

-fid into rather narrow segments (1-2
cm. wide), these laciniate ca. halfway to midrib into lance-linear lobules
and teeth 1-4 mm. wide, 2-15 mm. long; inflorescence densely yellow-
glandular, branched at base, side branches with rather compact, few-
flowered racemes, central raceme lax, to 2 dm. long; lowest bracts foliose,
others lance-linear, pubescent, 5-12 mm. long; pedicels divergent, glan-
dular, 2-5 cm. long; bracteoles ca. 2 mm. long, near the flowers; sepals pale
blue, to whitish striped, more or less glandular and pubescent ; upper
sepal oblong-ovate, 13 mm. long, 5 mm. wide, obtuse, the spur spreading
decurved, 10-14 mm. long, 2.5 mm. broad at base; lateral sepals oblong-
elliptic, 11 mm. by 5 mm., rounded at apex; lower sepals 14 mm. by 4
mm.; petals black, the upper laminae somewhat oblique, 8 mm. long,
glabrous, emarginate, the spur 10 mm. long; lower laminae round-ovate,
5 mm, in diameter, bearded near base, sparsely so on the 2 lobes, claw 5

mm. long; stamens 6—7 mm. long: anthers d
glabrous. : s dark, 1 mm. long; carpels 3,

3
2)
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4,
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5
ig")
n
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oO
3
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ia)
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a
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Type. Transcaucasia (Ossetia Meridionali :
: . near S
1933, Eliz. Busch (iE) seen. ) near Sredneje Sba, Aug. 19,

ANOTHER SPECIMEN. South Ossetia: E. & N. Busch, Aug. 6, 1935 (BM).

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 141

203. Delphinium elliptico-ovatum W. T. Wang, Acta Bot. Sinica 10:
83. 1962.

Stem ca. 7 dm. tall, 3 mm. thick at base, sparsely retrorse-hirsute, few
branched above, equably leafy; median cauline leaves long petioled, blades
as in D. naviculare, ca. 5.7 cm. long, 8 cm. wide, 3-parted to 8 mm, from
the base, the middle part obovate-rhombic, acuminate, 3-lobed to middle,
strigulose on both surfaces; petioles slightly longer than blades, not or
scarcely vaginate at base, with the pubescence of the stems: racemes 7—8-
flowered, rather lax, the axes loosely retrorse-strigulose; basal bracts trifid,
others lanceolate or elliptic-ovate, 8-12 mm. long, 2—2.5 mm. wide; pedicels
1-2.7 cm. long, densely retrorse-strigulose, divergent; bracteoles 1-7 mm.
remote from the flower, 5.5—7 mm. long, 2-3 mm. wide, acuminate, strig-
ulose; sepals blue-purple, 10-13 mm. long, strigulose; spur 12—13.5 mm.
long, cylindro-subulate, 2—-2.5 mm. thick, slightly decurved; petals black,
the upper laminae puberulent at apex, the lower round-ovate, bilobed, with
triangular lobules, long-ciliate at end, yellow- and white-bearded at base,
claw appendiculate, longer than the limb; stamens glabrous; follicles 3,
loosely puberulent to glabrous, 9 mm. long, the styles an additional 2.5
mm.; seeds triquetrous, 1.5 mm. long, scaly-laminate.

Type: Sinkiang, Wen Chuan, K. C. Kuan 3858, Aug. 28, 1957 (pr);
not seen. I have seen no material of this species. It is supposed to be near
to D. retropilosum (Huth) Sambuk, in which the leaves are basally some-
what cuneate, the sepals densely strigulose.

Fic. 17, E.

Delphinium ciliatum Stev. Ind. Sem. Hort. Dorpat (1811) ex Bieb. Fl. Taur.
Cauc. 3: 370. 1819. A specimen from Narzana (Nartsana) labelled D.
ciliatum Stev. (K) has been seen.

? Delphinium flexuosum vars. brevipila Busch and hirsuta Busch in Fl. Cauc.
Crit. III. 3: 67. 1902.

Delphinium elatum var. subciliatum Ledeb. Fl. Rossica 1: 64. 1842, A new
name for D. flexuosum and D. ciliatum.

? Delphinium flexuosum vars. dasyanthum and lasiocarpum Rupr. Fl. Cauc.
34, 35. 1869.

Perennial from a woody rhizome with strong fibrous roots; stem flexuous,
robust, 4-8 mm. thick, striate, reddish, spreading-hairy to subglabrous,
6-10 dm. tall, leafy, simple or branched above; petioles subglabrous to
hairy, dilated at base, the lower to 1 dm. or more long, the upper almost
suppressed; leaf blades pentagonal in outline, broader than long, 3-10 dm.
broad, glabrous to villous on both surfaces, paler beneath than above,
parted to one cm. or so above the base into segments with rather parallel
sides near the base, then more or less ovate and trifid or incised into teeth
or shallow lobes sublanceolate in shape, acute; inflorescence a dense many-
flowered raceme or an open panicle of racemes, glabrous to hairy; lower
bracts leafy, trifid, the upper entire, lance-linear, 5-20 mm. long; pedicels

204. Delphinium flexuosum Marschall Bieberstein, Fl. Taur. Cauc. 2:
2. 1808.

142 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

a<

: :
FIGURE 17. De um, Group IX. Petals, the upper spurred, the lower

Iphini
clawed. A. D. elatum; upper lamina 8 mm. long, spur 8 mm.; lower lamina 3.5
mm. long, claw 6 mm.; dra n from Nilsson i in 1898 (us). B. D. elatum; upper
i ; drawn

L

=

. long, spur 10 mm.;
lower lamina 5 mm. long, cla s wa from — reac in 1933,
TYPE (LE). E. D. flexuosum; upper iaios ur 15 ; lower
lamina 6 mm. long, claw 5 mm.; drawn from: Grossheim & Schiskin 333 dehy

F. D. formosum,; upper lamina 11 mm. long, spur 12 m na 6m

ower lam
long, claw 6 mm.; drawn from Balls 496 (x). G. D. ledsonaad: ibes dian
11 mm. long, spur 17 mm.; lower lamina 4.5 mm. long, claw 5 mm.; drawn from

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 143

divaricate, 5-15 mm. long; bracteoles lance-linear, 4-8 mm. long, sub-
tending the flower; sepals blue, glabrous to villous; upper sepal ovate,
13-15 mm. long, 1 8 mm. wide, acute, the spur almost straight, 15-18
mm. long, 3.5—-4 mm. wide at base; narrowed to an acute tip; lateral sepals
rounded-ovate, 12 mm. by 8 mm., obtuse; lower sepals 13-14 mm. long,
6—7 mm. wide, subacute; petals dark; the upper laminae slightly oblique,
9 mm. long, bidentate, pilose or glabrous, the spur ca. 15 mm. long; lower
laminae only slightly oblique, the blade 6 mm. long, long bearded and
ciliate, the lance-oblong lobes divaricate, claw 5—6 mm. long; stamens 6-7
mm. long, a few hairs along the margin of the filaments; anthers dark, 1
mm. long; follicles 3, glabrous, divergent, 7-8 mm. long, the styles 2.5
mm. long; seeds triquetrous, 1 mm. long, dark, squamate.

Type: No specimen cited, but from LE one was loaned me, from the
Caucasus, Mt. Berchtau, Steven, ex Herbario Marschall Bieberstein,
labelled typE; there are photographs at (a, K). Another specimen in the
Leningrad loan was “In herbidis montosis circa acidulam Nartsana, 3500’,
August 9, 1829,” no collector given, labeled D. flexuosum.

ILLUSTRATIONS. TREVIRANUS, Delph. & Aquileg. Obs. pl. 1. 1817 and
‘Bieberstein, Cent. Pl. Rar. 2, 11:64. 1843, according to Index Londinensis.

REPRESENTATIVE SPECIMENS. Transcaucasia: prov. Tiflis, Grossheim & Schiskin
333 (BM, K, NY); Mt. Kochta, Wissjulina in 1928 (BH); Azerbajdzhan, Karjagen
in 1935 (BH); Funtel flum., Kchetiuscaja, Busch in 1903 (x); in Caucaso iberico
lectum, dedit Steven Jan., 1821 (K).

205. Delphinium formosum Boiss. & Huet, Diagn. sér. IT. 5 ‘ aS. eae
IG.

Delphinium formosum var. centiflorum Huth, Bot. Jahrb. 20: 411. 1895.
Huth cited Balansa from “Vallée alpine de Djimil (Lazistan), 2000 m
this is 1352 (A, K, Pp) from northeast Turkey.

Perennial, 1-2 m. tall, the stem thick, ridged, pilose below, glabrous
above, branched above, equably leafy; leaves subglabrous, 5- or 7-parted
to ca. 1 cm. from base, the divisions broadly cuneate-obovate, incisely
serrate into teeth or short lobes 3-10 mm. wide; cauline petioles to 1 dm.

Shei & Sherriff 2424 (sm). H. D. himalayae; Hear lamina 10 mm. long,
mm.; ees pe ioe long, claw 6 mm.; drawn from rece eg
5

(e : L D. tgs upper lamina 8.5 pad long, ‘oni: ‘19 mm.; lower lamina 4.
m : long, claw és rawn from Lace 6012 (£). J. D. leptopogon; upper
lamina 9 m cae g, spur fs mm.; lower lamina 9 mm. long, cla uf
from Harry Smith 7109 K. D. maackianum; upper lamina 11 mm. long,
spur 16 mm.; lower seein 5 mm. long, claw 5.5 mm.; drawn from James in
1886 (cH). L. D. riae; upper lamina 12 mm. long, mm.; lower
lamina 5 mm. lon mm.: drawn from Alexeenko in 1898, TYPE (LE)
M. D. megalanthum; upper lamina mm. | r 16 mm.; lower lamina
mm, | claw m.; drawn from Lipsky in 1892, TYPE (tz). N. D. mira-

bile: upper lamina 10 mm. long, spur 17 mm.;
6.5 mm.; drawn from Bunge in 1839 (mo).

144 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

long, scarcely dilated at base; upper leaves 3- or 5-parted, reduced, sub-
sessile; inflorescence paniculate, the central raceme densely many flowered,
1-3 dm. long; bracts lanceolate, 1-2 cm. long, entire, or the lowest trifid
and larger; pedicels divaricate, 1-5 cm. long, glabrous; bracteoles opposite,
ovate to lanceolate, 5-12 mm. long, glabrous, reaching the base of the
flower or not; sepals violet-blue, broadly to oblong-ovate, 2-3 cm. long,
pubescent to glabrous without and within, the margins ciliate; spur straight
or slightly curved, 12-15 mm. long, 4 mm. wide at base, rapidly narrowed ;
petals black-purple, the upper laminae somewhat oblique, 11 mm. long,
somewhat pilose at apex, spur 12 mm. long; lower laminae bifid, yellow
bearded, ca. 6 mm. long, ciliate, the claw 6 mm. long; stamens 6-7 mm.
long; anthers bluish, 1 mm, long; young follicles glabrous in type collec-
tion at Kew, sometimes short-pubescent, divergent in maturity, 12-15 mm.
long, 4-5 mm. wide, the styles 3-4 mm. long; seeds brown, almost 2 mm.
long, covered with horizontal rows of scales.

Type: “Cerca Maadenkhan inter Erzeroum et Baibout,” Armenia, Huet
du Pavillon, 1843 (G) not seen, (BM, K) seen; photo. (BH).

RANGE. Said to grow on coarse limestone scree, rocky slopes, 1500-2790
m., Turkey, Georgia.

ItLustraTIONs. FI. des Serres I]. pl. 1185. 1857; Rev. Hort. IV. 8:
528. 1859; Gartenflora 8: pl. 253. 1859

REPRESENTATIVE COLLECTIONS. Turkey: E. K. Balls 1975, 496; Davis 21 iZé;
21211; Sintenis 1590; du Parquet in 1886. Cultivated as D. formosum, D. for-
mosum coelestinum; D. grandiflorum.

Y 206. Delphinium hillcoatiae Munz, sp. nov. Fic. 17, G.

Perennial from a woody root, the stem simple or branched from base,
often more or less purplish, ridged, with short spreading glandular hairs,
quite leafy throughout, 1.5—5 dm. tall, the leaves gradually reduced up-
ward; petioles hairy, 2-8 cm. long, dilated at base; laminae viscid, more
or less cupped, not pressing flat, 2-4 cm. long, 4-10 cm. wide, subcuneate
at base, parted to within 1 cm. from base into subrhomboid segments,
these mostly 3-lobed ca. halfway to the midrib, then each lobe with ca.
3 oblong-ovate teeth 5-6 mm. long and wide, or in upper leaves in turn
divided into smaller narrow lobules; raceme terminal, simple, 1-3 dm.
long, glandular, spreading-pubescent, 4- to 20-flowered; lower bracts
reduced leaves, the upper simple and entire; pedicels divergent, stiff,
straight, 1-5 cm. long; bracteoles subtending the flowers, lance-linear, 5—7
mm. long; sepals glandular, white tinged green and with green tips, the
spur pale green; upper sepal ovate, 18-19 mm. long, 9-11 mm. wide,
apiculate, the spur 20-25 mm. long, 3.5-4 mm. wide at base, spreading
decurved; lateral sepals narrowly oblong-obovate, 14-15 mm. by 4-6 mm.,
apiculate; lower sepals oblong-ovate, 14-15 mm. long, 5-6 mm. wide,
apiculate; petals dull velvety black, the upper laminae somewhat oblique,
10-12 mm. long, 3 mm. wide, with 2 narrow lobules 1—1.5 mm. long, the
spur 17-20 mm. long; lower laminae subovate, 5 mm. by 2.5-3 mm.,

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 145

somewhat bearded from near base to tip, bifid about 1/3-1/2 its length
into lanceolate lobes, the claw 5 mm. long, not appendaged; stamens 6—7
mm. long; anthers dark, 1 mm. long; carpels 3, glabrous.

Perennis de radice lignosa; caulis simplex vel base ramosus, glanduloso-
hirsutus, 1.5-5 dm. altus, foliosus; petioli hirsuti, 2-8 cm. longi, base
dilatati; laminae foliorum viscidae, 2-4 cm. longae, 4-10 cm. latae, base
subcuneatae, profunde partitae; segmenta subrhomboidea, 3-lobata, lobis
dentatis; racemus terminalis simplex, 1-3 dm. longus, glanduloso-pubescens,
cum 4-20 floribus; bracteae simplices, integrae vel inferiores lobatae;
pedicelli divergentes, 1-5 cm. longi; bracteolae lanceo-lineares, 5—7 mm.
longae, florem subtendentes; sepala glandulosa, albida, sepalum superiore
ovatum, 18-19 mm. longum, 9-11 mm. latum, apiculatum, calcar 20-25
mm. longum; sepala lateralia anguste oblongo-obovata, 14-15 mm. longa;
sepala inferiora oblongo-ovata, 14-15 mm. longa; petala subatra; laminae
superiores subobliquae, 10-12 mm. longae, acute lobulatae; laminae in-
feriores subovatae, 5 mm. longae, barbatae, bifidae, unguis 5 mm. longum;
carpella 3, glabra.

TypE: Kap, Chayul Chu, S. Tibet, July 23, 1936, at 11000’, on open
dry stony hillside, Ludlow & Sherriff 2424 (BM).

ADDITIONAL MATERIAL. Tibet: Chayul Dzong, slate screes in Nye Chu Valley,
3700 m., Sept. 21, 1935, Kingdon Ward 12370 (BM).

It is a pleasure to dedicate this species to Miss Dorothy Hillcoat of the
Department of Botany, British Museum (Natural History), whose interest
in the genus Delphinium was most useful to me in my work at that
institution.

207. Delphinium himalayae Munz, sp. nov. Fic. 17, H.

Delphinium himalayense Chowdhury ex Mukerjee, Bull. Bot. Surv. India 2:
293-296. 1960, nomen.

Perennial, the stem 4-6 dm. tall, simple, slender, more or less retrorse-
strigulose throughout, more densely so in the inflorescence; leaves few,
largely near the base; petioles 1-2.5 dm. long, dilated at the base; leaf
blades to 1 dm. broad, minutely strigulose on both surfaces, but more
heavily so beneath, 5-fid to within 1 cm. from the base and the 2 outer
segments immediately divided to make 7, these cuneate-obovate, bisect
or trisect to ca. the middle, then each part lobulate into few lance-oblong
lobules to 1 cm. long and 2-3 mm. wide, mucronate; inflorescence a secund
several-flowered raceme 1-1.5 dm. long; lower bracts foliose, others
lanceolate to oblong, 1-2 cm. long, conspicuous; pedicels erect, 1-3 ee.
long; bracteoles subtending the flower, oblong-linear, 6-10 mm. long;
sepals purplish blue, strigose, the upper broadly ovate, 17-18 mm. long,
15-16 mm. wide, obtuse, the spur often suberect, 13-15 mm. long, 4 =
wide at the base, only gradually narrowed to an obtuse tip; lateral sepa Is
ovate, 14-17 mm. long, 10 mm. wide, subacute; lower sepals asymmetri-

146 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

cally ovate, 15 mm. by 12 mm., subacute; petals black, the upper laminae
10-11 mm. long, 4 mm. wide, slightly oblique, bidentate and somewhat
bristly at the apex, the spur 13-14 mm. long; lower laminae oblong, 8-10
mm. long, bifid about halfway with 2 linear-oblong lobes and more or less
hairy, the claw 6 mm. long, appendiculate; stamens 6-7 mm. long; anthers
dark, oblong, 1.5 mm. long; carpels 3, densely pubescent.

Perennis; caulis 4-6 dm. altus, simplex, tenuis, retrorso-strigulosus ;
folia pauca, vulgo basalia; petioli 1-2.5 dm. longi, base dilatati; laminae
foliorum 5—10 cm. latae, strigulosae, 5-fidae; segmenta cuneato-obovata,
trisecta, tum lobulata; inflorescentia secunda, racemosa, 10-15 cm. longa;
bracteae infimae foliosae, superiores lanceolatae vel oblongae, 1-2 cm.
longae, conspicuae; pedicelli erecti, 1-3 cm. longi; bracteolae oblongo-
lineares, 6-10 mm. longae, florem subtendentes; sepala purpureo-caerulea,
strigosa; sepalum superiore late ovatum, 17-18 mm. longum, 15-16 mm.
Jatum, obtusum, calcar saepe suberectum, 13-16 mm. longum, base 4 mm.
latum; sepala lateralia ovata, 14-17 mm. longa, subacuta; sepala inferiora
ovata, 15 mm. longa, 12 mm. lata; petala atra, laminae superiores 10-11
mm. longae, parum obliquae, bidentatae, leviter setosae, calcar 13-14 mm.
longum, laminae inferiores oblongae, 8-10 mm. longae, bifidae, lobis
lineari-oblongis, villosis, unguis 6 mm. longus, exappendiculatus; stamina
6-7 mm, longa, glabra; carpella 3, pubescentia.

Type: Nepal, Ser Oive Wigram 118, in 1927 (k£).

Rance. The proposed new species, Delphinium himalayae, grows in
grassy places at ca. 3000 to 4500 m. in Nepal.

EXAMPLeEs. Nepal: Langdeng, Sharma 117 (pm, £); Pongsing, Dhwoj 122 (8M,
E); below Mugu, Polunin, Sykes & Williams 5290 (gm); Ghurchi Lekh, Polunin,
Sykes & Williams 5128 (gm); Maharigaon, Polunin, Sykes & Williams 264
(BM) ; Chutta, Polunin, Sykes & Williams 4885 (sm); Balangra Pass, Polunin,
Sykes & Williams 2503 (pm); Langtang Village, O. Polunin 1489 (pm), 552

(pm); Chilime Kharka road, Polunin 1413 se :
1778 (BM). (pm); Kyangjin Ghyang, Polunin

I have had no contact with Dr. Chowdhury and do not know why he
never published Delphinium himalayense, the name which appears on
some herbarium material in Great Britain. Since I do not know what his
species was supposed to include, I do not feel free to use his name, taken
up by Mukerjee, but never validly published. My concept of Delphinium

himalayae 1S entirely my own, as are the description and citation of her-
barium material.

208. Delphinium inconspicuum Sergievsk. Animadvers. Syst. Herb.
Univ. Tomsk 1930. (3-4) : 6. fig. 1930

.

Stems many, 2.5-10 dm. tall, leafy throughout, short glandular-pubes-
cent, more so upward; leaves orbicular-cordate or -reniform, 4.5—-7 cm.
long, 5-14 cm. wide, coarsely toothed, short pilose on both surfaces;
raceme simple or few branched below, dense, 3-17 cm. long; lower bracts

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 147

large, leaflike; bracteoles linear, 0.5 mm. wide; inflorescence pilose and
glandular; sepals blue, 8-18 mm. long, 5-9 mm. wide, spur 11-13 mm.
long, 2.5 mm. wide at base; petals dark, upper laminae bidentate, glabrous;
lower laminae semibifid with unequal oblong lobes, ciliate, the claw hirsute;
carpels 3, glabrous; seeds winged.

Type: Semipalatinsk in jugo Narymense; not seen. The species ap-
parently occurs also in Mongolia. Delphinium inconspicuum seems, from
the meager description, to key out near D. afghanicum of Afghanistan and
D. pyramidatum of Transcaucasia, but is far removed from them geo-
graphically.

209. Delphinium kawaguchii Tamura, Acta Phytotax. & Geobot.
Kyoto 15: 194. fig. 2. 1954.

Leaves radical; petioles 16-20 cm. long, pilose, the blades reniform, 7
cm. long, 9 cm. wide, 3—5-fid, the lobes pinnately incised-lobed, pubescent
on both surfaces on veins and margins; inflorescence racemose, 3—4 dm.
long, 18—23-flowered; lower bracts trifid, upper entire, linear-oblong;
pedicels 2—3.5 cm. long; sepals blue, pubescent, the spur cylindric, 18 mm.
long, surpassing the sepal blades, 3.5 mm. wide at base, incurved; laminae
of upper petals glabrous, 9 mm. long, the spur 16 mm. long; lower laminae
10 mm. long, round-ovate, 2-lobed, the lobes acute, bearded above;
carpels 3, densely villous; styles long, glabrous.

Type: Tibet near Shigatse, Linbun rivulet, July 29, Kitamura 103908
(kyo). I have been unable to see material of this species. The original
description said that it differs from D. speciosum in the inflorescence not
being leafy, and from D. iliense in its villous carpels, scarcely erect pedi-
cels, and bracteoles not next to the flower.

210. Delphinium kuanii W. T. Wang, Acta Bot. Sinica 10: 84. 1962.

Stem 5 dm. tall, 3 mm. thick at base, longitudinally striate, retrorsely
or spreading-hairy, few branched, equally foliose; basal leaves and lower
cauline long petioled; blades shaped as in D. naviculare, broad-reniform,
5.5 cm. long, 10 cm. wide, 3-parted to ca. 1 cm. above the base, middle
part cuneate-obtrapezioid, 3-lobed above, the lobes 1—2-toothed, the teeth
deltoid, laxly hirsute on both surfaces, petioles from twice as long as
laminae to equal to them, narrowly vaginate, with same indument as the
stems; racemes 4—10-flowered, lax, the axes strigulose; lower bracts trifid,
others lanceolate, 8-14 mm. long, 2-3 mm. wide, densely strigulose;
pedicels 1-3 cm. long, densely retrorse-strigulose, ascending; bracteoles
to 15 mm. below the flowers or near them, lance-oblong or lanceolate,
8-10 mm. long, 2—2.5 mm. wide; sepals purplish, elliptic, 12-15 mm, long,
densely strigulose without, glabrous to loosely hairy within, spur 11.5-13
mm. long, saccate-cylindric, 44.5 mm. thick at base, straight, obtuse;
petals black, the upper laminae glabrous, rounded at apex; lower laminae

148 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

broadly rectangular, with 2 deltoid lobules at the rounded apex, few
toothed, yellow bearded at base, claw appendiculate; filaments pilose;
carpels 3, densely strigose; style barely 1 mm. long.

Type: Sikang, Wen Chuan, Aug. 27, 1957, K. C. Kuan 4698 (PE), not
seen. I have had no material of Delphinium kuanii, which the author said
was near D. elliptico-ovatum but with bracteoles lance-oblong to lan-
ceolate; spur saccate-conic; ovary densely strigulose, and styles shorter.

211. Delphinium lacei Munz, sp. nov. Fic. 17,4.

Perennial from thick fleshy roots; stems slender, usually somewhat
branched above, often reddish, slender, retrorse-strigulose or -pubescent,
scarcely if at all glandular, 4-8 dm. tall, with a few scattered leaves; basal
leaves with blades somewhat reniform in outline, 3-6 cm. broad, S- or
7-fid to ca. 3/4 their length, with a scattering of long hairs on both
surfaces, the segments subcuneate, sharply 3-toothed at apex or sometimes
3-lobed, each lobe then sharply toothed, the teeth acute-acuminate,
mucronate, lance-ovate; petioles slender, to 1 dm. long, spreading-villous,
somewhat dilated at base; upper leaves reduced, the uppermost to ca. 1
cm. long, 3-lobed with broad petioles; inflorescence a few-flowered raceme,
with shorter subsidiary branches; bracts mostly lanceolate, 8-15 mm. long,
villous; pedicels ascending, 1-3 cm. long, the bracteoles near the middle or
above, lanceolate, 5—8 mm. long; flowers bluish, with more or less yellowish
hairs throughout; upper sepal ovate, 18 mm. long, 12 mm. wide, obtuse,
the spur somewhat decurved, 16-20 mm. long, 2.5-3 mm. wide at base;
lateral sepals round-obovate, 15-16 mm. long, 12 mm, wide, rounded-
truncate at apex; lower sepals elliptic-obovate, 18 mm. by 10 mm., obtuse;
petals black, the upper laminae somewhat oblique, 8 mm. by 3 mm.,
slightly emarginate at the rounded apex, the spur 15-17 mm. long; lower
laminae oblong-ovate, 4.5 mm. by 4 mm., bearded, ciliate, bifid ca. 1.5
mm. into broad lobes, claw broad, 5 mm. long; stamens 6-7 mm. long,
glabrous; anthers dark; follicles 3, erect, 12-16 mm. long, 4 mm. wide,
pubescent, the styles an additional 3 mm.; seeds black angled scarcely
winged or squamate, almost 2 mm. long.

Planta perennis, ex radicibus lignosis; caules tenues, supra plerumque
ramosi, saepe rubelli, retrorso-pubescentes, vix vel non glandulosi 4-8 dm.
alti, parum foliosi; laminae foliorum basalium subreniformes ver cm.
latae, 5—7-fidae, sparse villosae, segmentibus subcuneatis apice acute
3-dentatis vel -lobatis; petioli tenues, ad 1 dm. longi extenso-villosi,
subvaginati; folia Superiora parva, 3-lobata, late netiolata; inflorescentia
racemosa, cum floribus paucis et ramulis brevibus: beaitene lanceolatae,
8-15 mm, longae, villosae; pedicelli ascendentes, 1-3 cm. longi; bracteolae
in medio pedicelli, lanceolatae, 5-8 mm. longae: flores subcaerulei cum
pilis aureis; sepalum superiore ovatum, 18 mmm. longum, 12 mm atum
obtusum; calcar curvatum, 16-20 mm. longum, base 2 5-3 mm latum;
sepala lateralia rotundo-obovata, 18 mm. longa, 12 mm lata; sepala in-

it b

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 149

feriora elliptico-obovata, 18 mm. longa, 10 mm. lata, obtusa; petala atra,
laminae superiores 8 mm. longae, parum emarginatae, calcar 15-17 mm.
longum; laminae _inferiores oblongo-ovatae, 4.5 mm. longae, hirsutae,
bifidae, unguis latus, 5 mm. longus; stamina 6-7 mm. longa, glabra;
follicula 3, erecta, 12-16 mm. longa, pubescentia; semina atra, angulosa,
fere 2 mm. longa.

Type: Burma, Ruby Mines district, Bernardmyo, 1750 m, Uet. SS,
1912, J. H. Lace 6012 (cat, £ 3 sheets, K); seen. I would designate the
K sheet as type, those at cat and F as isotypes.

A? ER COLLECTION. Burma: Mt. Victoria Ridge, 2700 m., Oct. 26, 1956,
Kingdon Ward 22749 (pm).

212. Delphinium leptopogon Hand.-Mazz. Acta Horti Gothob. 13:
58. 1939. Fic. 17, J.

Perennial, the stem 7-8 dm. tall, branched only above, slender, to 5
mm. thick, glabrous and glaucous below or with some stiff white hairs at
base of petioles, evenly but rather remotely leafy; leaf blades roundish in
outline, the lower 8-13 cm. wide, green and minutely strigulose on both
surfaces, the cauline divided almost to base into 5 divaricate lanceolate,
entire to 2—3-toothed segments 5-8 mm. wide, acuminate; petioles not
dilated at base, about as long as the blades or the uppermost shorter ;
inflorescence subcorymbose at the end of each branch, 4~20-flowered, with
spreading glandular hairs; bracts leaflike and trifid or entire and lanceolate,
1-2 cm. long; pedicels divaricate to erect, 1-4 cm. long; bracteoles in
middle or above, subulate, 3-5 mm. long; sepals violet-blue, densely
strigulose; upper sepal ovate, 14 mm. by 8 mm., acute, the spur almost
straight, 20-22 mm. long, 4-5 mm. wide at the base, gradually narrowed
to a blunt tip; lateral sepals ovate, 17 mm. by 8 mm., acute; lower sepals
15 mm. by 6 mm., subacuminate; petals mostly dark, the upper laminae
quite oblique, 9 mm. long, subacuminate, somewhat emarginate, the spur
19 mm. long; lower laminae sometimes yellowish, strongly oblique, oblong,
with scattered white hairs, entire to shallowly 2-lobed, ca. 9 aie by 3 eae,
the claw 6 mm. long; stamens 6-7 mm. long, the base only slightly widened,
glabrous; anthers dark, linear-oblong, 1.5 mm. long; follicles 3, glabrous
to sericeous; seeds winged on angles.

Type: Shansi, distr. Chiao-ch’eng, Pa-shui-ko, 1900 m., Harry Smith
7109, holotype (ups), not seen; isotype (A), seen.

I would place in this species also Rock 13136 (GH, NY) from southwestern

ansu.

213. Delphinium longiciliatum W. T. Wang, Acta Bot. Sinica 10:
84. 1962

Stem ca. 8 dm. tall, 3 mm. thick at base, loosely spreading- or retrorse-

hirsute, simple, equably leafy; middle cauline leaves longish petioled; leat

blade as in D. naviculare, broadly pentagonal, 6.5 cm. long, 11 cm. wide,

150 JOURNAL OF THE ARNOLD ARBORETUM [vor. 49

3-parted, the middle part rhombic, upper trilobed, the middle lobes
triangular-ovate, with few coarse teeth, the lateral lobes smaller, obliquely
ovate, coarsely 1—2-toothed, loosely strigulose on both surfaces; petioles
ca. as long as blades, not vaginate at base, with same pubescence as on the
stems; raceme ca. 16 cm. long, 10-flowered, rather lax; pedicels 1-3 cm.
long, ascending, glabrous; lower bracts lanceolate, ca. 15 mm. long, 3 mm.
wide, others 7-12 mm. long, 1—1.2 mm. wide, acute, long-ciliate; bracteoles
to 3-5 mm. below the flower, linear-lanceolate to oblanceolate, 7 mm. long,
acute, long-ciliate; sepals purple, the upper ovate, others obovate, 13-14
mm. long, glabrous or slightly ciliate at apex; spur 17-18 mm. long,
straight; petals black, the upper laminae emarginate, loosely pilose; lower
laminae broadly ovate, with 2 obtuse, triangular lobes, yellow bearded at
base, long-ciliate at ends, claw shorter than lamina, appendiculate at base;
stamens glabrous; carpels 3, subglabrous.

Type: Sinkang, at Ili, Aug. 8, 1957, K. C. Kuan 3209 (pe?), not seen.
The author of the species suggested that it is near D. tianshanicum
Wang, but with the sepals, pedicels, and ovaries glabrous.

214. Delphinium maackianum Regel, Mém. Acad. Pétersb. VII. 4 (4):
9. 1861. Fig. 17, 3.
Delphinium maackianum var. ussuriense Regel, loc. cit. “bracts and carpels
glabrous,” apparently based on same material as the species. Var. Jasio-
carpum Regel, based on Wilford from the “Coast of Manchuria” (GH, K, P)
seen

Delphinium maackianum var. cuneatum Regel ex Huth, Bot. Jahrb. 20: 400.
1895. Apparently the type collection is “Ad flum. Amur, montes Burgae”
(Bureja in Huth), Radde (x, Pp) seen. :

Delphinium maackianum Regel var. palmatum (Lévl.) Nakai, Jour. Coll. Sci.
Imp. Univ. Tokyo 31: 433. 1911. D. elatum var. palmatum Lévl., Bull.
Acad. Géogr. Bot. 11: 300. 1902. D. lycoctonifolium Lévl., Repert. Sp.
Nov. 7: 100. 1909. All based on Faurie 26, from mountains Kan-OQuen-To,
Korea (£), isotype (uc); both seen.

Perennial from a woody rhizome, the stem 5-8 dm. tall, glabrous or

pubescent below, pubescent above with short spreading hairs, stem simple
or branched in the inflorescence, ca. 5-6 mm. thick below, leafy, the
petioles glabrous to hairy, dilated at base, the lower to 2 dm. icaag, the
upper much shorter; leaf blades mostly broadly rounded in outline, the
lower to 2 dm. wide, not quite as long, subglabrous to strigose on both
surfaces, 5- or 7-fid to well above the base into broad cuneate-based ovate
segments commonly 2—5 cm. broad and rather shallowly incised or toothed
into ovate or lance-ovate lobes or teeth; upper leaves reduced, often only
3-fid; inflorescence loosely paniculate, pubescent, the lower bracts folia-
ceous, divided, the others ovate or elliptic-ovate, membranous, bluish
glabrous, 5-20 mm, long; pedicels pubescent, ascending, 1-5 cm. foie:
bracteoles lance-ovate, shorter, inserted well below the flower: sepals dull
blue, subglabrous or slightly pubescent, the upper sepal 14-16 mm. long,

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 151

8-10 mm. wide, ovate, subacute, the spur straight or curved or even
uncinate, 15-20 mm. long, 3.5-4.5 mm. wide at base, narrowed to rather
a sharp tip; lateral sepals 12-16 mm. long, 8-10 mm. wide, obtuse; lower
sepals 12-14 mm. long, 6-7 mm. wide, subacute; petals dark, the upper
laminae slightly oblique, glabrous, subacute, entire or nearly so, 10-11
mm. long, the spur 15-16 mm. long; lower laminae ovate, oblique, 5 mm.
long, 3 mm. wide, cleft one half their length, ciliate and sparsely bearded,
with a patch of yellow hairs near the base, claw 5.5 mm. long; stamens
6-7 mm. long, glabrous, slightly dilated below; anthers dark, 1.5 mm.
long; follicles 2-3, glabrous, divergent, 18-20 mm. long, the styles an
additional 3 mm.

Type: Manchuria, on the Sungacha River (tributary of the Ussuri
River), Maack, isotypes (GH, k, P), seen.

DIstRIBUTION. The species occurs in Manchuria and Korea.

EXAMPLES: Manchuria: Komarov 671 (BM, K); Maximowicz in 1860 (BM, GH,
K, NY, P); Litvinov 2026 (Ny), 3013 (NY); F. H. Chen 341 (BM, Gu, NY, P);
Bohnhof 163 (Ny, p). Korea: Chung In-Cho’s collector 9440 (micH), 10347
(MICH), 9391 (miIcH); Ohwi 2808 (BH); Bushnell 470 (BM).

215. Delphinium mariae Busch in FI. Cauc. Crit. III. 3: 69. 1903.
Fic. 17, L.

Perennial, the stem several dm. tall, strigulose below, otherwise glabrous,
angular-sulcate, leafy; cauline leaves slightly strigose on both surfaces,
paler beneath, ca. 1 dm. broad, 5-parted almost to the base, the segments
narrowly cuneate-obovate, deeply laciniate into lance-linear lobes 1.5—2.5
mm. wide, 5-25 mm. long; petioles scarcely vaginate at base; raceme lax,
many flowered, glabrous, to 3.5 dm. long, branched below; bracts linear-
lanceolate, 1-2 cm. long; pedicels suberect, 3-4 cm. long; bracteoles
subtending the flowers, linear-lanceolate, ca. 10 mm. long, 2 mm. wide;
sepals blue-violet, glabrous, slightly ciliate, the upper sepal ovate, 18-20
mm. long, 8 mm. wide, obtuse, the spur spreading decurved, 16-17 mm.
long, 3.5 mm. wide at base; lateral sepals elliptic-obovate, 20 mm. by 12
mm., rounded at apex; lower sepals 18 mm, by 7 mm., rounded-truncate
at tip; petals black, the upper laminae slightly oblique, 12 mm. long,
slightly bidentate at the glabrous apex, the spur 13 mm. long; lower
laminae 5 mm. by 4 mm., oblong-obovate, bearded near base, long-ciliate
on the round-oblong divergent lobes, the claw appendiculate, 6.5 mm. long;
stamens 6-7 mm. long; anthers dark; carpels 3, glabrous.

Type: In meadows at 1850 m., between Tsulkan and Gapschina, Dages-
tan, July 19, 1898, Alexeenko (LE), seen.

I have seen no other examples of this species.

216. Delphinium megalanthum Nevski in Komarovy, TL USSR. 7:
129,422. 1937. Fic. 17, M.
Delphinium flexuosum var. paulovii Akinf. Trav. Soc. Nat. a P'Univ. Imp.
Kharkow XXVIII: 167. 1894. Delphinium speciosum var. paulovii N. Busch

152 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

in Fl. Cauc. Crit. III. 3: 64. 1903. Cited as synonyms in the original de-
scription of D. megalanthum.

Stem 3.5-7.5 dm. tall, ribbed, lightly pilose below, glabrous above,
foliose; petioles scarcely dilated at base, ciliate along margin; leaves
lightly pilose or glabrous above, more pallid and pilose on veins beneath,
ciliate on margins, 5-10 cm. broad, 5-fid, the segments cuneate-obovate,
3-incised to the middle, then coarsely few-lobulate into lanceolate parts
2-5 mm. wide, 2-10 mm. long, acute; racemes erect, branched at base, to
3 dm. long, lax; lowest bracts foliose, others linear-lanceolate, 6—20 mm.
long, 1-2.5 mm. wide, glabrous; pedicels strict or slightly spreading,
glabrous or almost so, 3-7 cm. long; bracteoles ovate-lanceolate or ovate,
7-9 mm. long, 3-4 mm. wide, obtusely acuminate, glabrous, rarely slightly
ciliate, usually violet in color, near or to 0.5 cm. below the flower; sepals
blue, the upper 22 mm. long, 10 mm. wide, obtuse, glabrous, the spur 20
mm, long, 4 mm, wide at base, slightly incurved toward the acute tip;
lateral sepals 23 mm. by 12-14 mm.; lower ca. 22 mm. long; petals black,
the upper laminae slightly oblique, glabrous, emarginate, 13 mm. long, the
spur 16 mm. long; lower laminae perpendicular to the claw, oblong, 9 mm.
by 4 mm., bearded at base and on the 2 oblong lobes, claw 7 mm. long;
stamens 7-8 mm. long; anthers dark, 1.5 mm. long; carpels 3, glabrous.

Type: “Caucasus septentrionalis, prope glaciem Bezengi, July 25, 1892,
Ig. Lipsky” (LE); seen. It is near to D. speciosum but seems different in
its glabrous flowers.

217. Delphinium mirabile Sergievsk. Animadvers. Syst. Herb. Univ.
Tomsk 1930. 3-4: 5. 1930. Fic. 17, N.

. .

Delphinium laxiflorum var. 8 alpinum Bunge, Fl. Altaica Suppl. 33 [orig. 555].
1836. Placed here in synonymy by Nevski in Komarov, Fl. U.S.S.R. 7:
131. 1937. I have seen a collection from “Altai” Herbar. Bunge, FI. orient.
altaica, 1839 (mo) which may represent this taxon.

Stem slender, 2.5—4 dm. tall, leafy below, sparsely pilose above, simple
or nearly so, angled below; petioles to 1 dm. long, slender, somewhat
dilated at base, pilose; leaf blades round-reniform in outline 3-9 cm. wide,
more or less strigulose above, paler and short-hairy ene 3-, 5-, or
7-parted almost to base into cuneate-rhombic segments 5-10 mm. wide
below, 10-15 mm. wide above, these trilobed to about the middle then
sometimes incised into sharp divergent teeth; inflorescence a 5—10-flowered
raceme or panicle 7-10 cm. long, strigose and with some short more
spreading hairs; bracts entire, mostly linear, 5-15 mm. long, sometimes
2—3-parted, pubescent; pedicels pilose, 1-2 cm. long; bracteoles linear
3—4 mm. long, placed just below the flower; sepals sordid blue-violet white
pilose on both outer and inner surfaces; upper sepal ca. 18 mm. by 14 mm
apiculate, the spur spreading decurved, 16-17 mm. long, 3.5 rare. wide an
base, subcylindric, the apex obtuse; lateral sepals broadly elliptic-obovate
18 mm. by 10 mm., broadly obtuse; lower sepals 20 mm. by 8 mm.,

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 153

subacute, asymmetrically lance-obovate; petals blackish, the upper laminae
almost straight, 10 mm. long, 3 mm. wide, rounded and slightly erose at
the summit, spur 17-18 mm. long; lower laminae oblique, oblong, 5 mm.
by 3.5 mm., bifid from near the middle into oblong lobes, scattered-hirsute,
ciliate, the claw 6.5 mm. long; stamens 6-7 mm. long, glabrous; anthers
dark, oblong, 1.3 mm. long; follicles 3, pubescent.

Type: “In montibus Altaicis austro-orientalibus ad fl. Taldura,” July
18, 1903, P. Krylov (TK); not seen.
Drawings and description made from “Altai,” Herbar. Bunge (mo).

218. Delphinium naviculare W. T. Wang, Acta Bot. Sinica 10: 82.
962.

Stem ca. 7 dm. tall, 3.5 mm. thick below, spreading or retrorsely hirsute,
glabrescent below the inflorescence, simple, subequally foliose; basal and
lower cauline leaves long petioled; leaf blades broadly reniform-pentagonal,
3—4.5 cm. long, 5—5.5 cm. wide, cordate, 3-parted, the middle part rhombic,
acute, 3-lobed below the middle, the lateral parts oblique, unequally
2-lobed to the middle, all 5 parts then few toothed, loosely hirsute on both
sides; petioles vaginate at base; racemes ca. 3 dm. long, many flowered,
rather dense; pedicels 1-3 cm, long, erect-spreading, glabrous; lower bracts
14-17 mm. long, trifid, the lobes lanceolate, acute, the other bracts ovate,
7-10 mm. long, 3.8 mm. wide, abruptly acute; bracteoles near the flowers,
ovate to elliptic, 4-5 mm. long, boat-shaped, acuminate, ciliate; sepals
purplish, glabrous or ciliate at apex, the upper broadly elliptic, lateral
round-obovate, lower elliptic, 9-12 mm. long; spur 12—13.5 mm. long,
glabrous, almost straight, conic-subulate; petals black, the upper laminae
glabrous, obliquely truncate, entire; lower laminae broad-oblong, with 2
deltoid-ovate lobes to the middle, yellow bearded at base, long-ciliate, the
claw exappendiculate; stamens glabrous; carpels 3, glabrous.

Type: Sikang, at Chigo-Su, Aug. 11, 1957, K. C. Kuan 3229 (PE), not
seen.

I have not seen this species which Wang says is near D. ruprechtii but
with smaller flowers and bracteoles. Wang’s Delphinium naviculare var.
lasiocarpum (Acta Bot. Sinica 10: 83. 1962), described as having pedicels
and sepals with spreading yellow-glandular hairs, is not known to me and
I am uncertain as to its disposition.

219. Delphinium nevskii Zakirov, Not. Syst. Herb. Inst. Bot. Acad.
och USS Ri IZ: 92.1950. Fic. 18, A.
Perennial from a woody root, 4-6 dm. tall, the stem white-hairy espe-
cially in the inflorescence, 5-7 mm. thick, few branched above, scattered
leafy; lower petioles 12-25 cm. long, sulcate, retrorse-pubescent, somewhat
vaginate at base, the upper much shortened; lower blades roundish in
outline, 6-10 cm. wide, pilose on both surfaces, 3-parted to below the
middle, the broad parts 2-3-lobed in distal portion, then with coarse more

154 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

or less oblong-ovate, mucronulate teeth 4-8 mm. long and almost as broad;
central raceme rather dense, hairy, to 2 dm. long, many flowered; bracts
lanceolate, white-hairy, 6-8 mm. long, 1-2 mm. wide; pedicels spreading-
ascending, 1-2 cm. long, soft-hairy; bracteoles above the middle of the
pedicel, subulate-lanceolate, 3-5 mm. long; sepals white with yellow nerves,
pilose on both surfaces; upper sepal 15 mm. by 7 mm., acutish, the straight
spur 12-15 mm. long, 2.5 mm. wide at base; lateral sepals subelliptic, 13
mm. by 6 mm., obtuse; lower sepals subelliptic, 12 mm. by 6 mm., more
pointed; petals blackish, the upper laminae almost straight, 8 mm. long,
bidentate, stiff-hairy at apex, the spur ca. 12 mm. long; lower laminae
oblong-ovate, oblique, 5 mm. long, bifid about halfway, hairy and white-
ciliate, the claw 5 mm. long; stamens 5-6 mm. long, glabrous; anthers
pale, 1 mm. long; follicles 3, soft-hairy, 10 mm. long, 3 mm. wide, the
styles an additional 3 mm.

Type: Uzbekisten, system of River Zeravschan, Mt. Chazrat-Sulton,
limestone, 3300-3800 m., Aug. 27, 1942, K. Zakirov (LE); seen.

The species is said to be near D. oreophilum, differing in smaller white
flowers and equal pubescence on both surfaces of the leaves.

220. Delphinium ochotense Nevski in Komarov, Fl. U.S.S.R. 7: 146,
724. 1937. Fic. 18, B.
Delphinium elatum var. subglabrum Regel & Tiling, Nouv. Mém. Soc. Nat.

Mosc. 11: 39. 1859, not var. subglabrum Ledeb. 1841. Based on same type
as D. ochotense.

Stem 2-5 dm. tall, with soft short pubescence, slightly glandular above,
densely leafy; leaves round-cordate to -reniform, to 15 cm. broad, slightly
pubescent, palmatisect, the segments commonly biternately divided into
terminal obtuse or subobtuse, broadly elliptic to lanceolate teeth 5-20
mm. long, 4-10 mm. wide; raceme many flowered, the axis and pedicels
densely soft-pubescent; bracts linear, 8-20 mm. long, the lowest lanceolate
or trifid, to 4 cm. long; pedicels erect, 1-3 cm. long; bracteoles subtending
the flower, linear, 4-6 mm. long; sepals blue, ovate to ovate-lanceolate,
obtuse, 1.3—2.2 cm. long, 5-9 mm. wide, with some spreading hairs; spur
spreading, almost straight or somewhat incurved near the apex, 1.2-1.7
cm. long; petals black, the upper laminae somewhat oblique, 10 mm. long,
glabrous, entire, the spur 16-17 mm. long; lower laminae bearded at base,
ovate, 4 mm. long, with triangular lobes 1.5 mm. long, claw 6 mm. long.

Type: Extreme eastern Siberia, near Ajan, legit Tiling (LE); seen.

Delphinium ochotense differs from D. elatum in having some glandular
hairs in the inflorescence and lower petals with broader laminae.

221. Delphinium oreophilum Huth, Bot. Jahrb. 20: 412. 1895.
Hic, 16, ©.

Stem thick, erect, subglabrous, 5-6 dm. tall; leaves well distributed on
the stem; petioles 2-4 dm. long, dilated at the base; the blades roundish,

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 155

8-12 cm. in diameter, divided almost to base into 3 or 5 parts, subglabrous
above, quite pubescent beneath especially on the veins, the parts broad,
then lobed about halfway to base into broad portions which are in turn
divided at tips into lanceolate lobules or teeth 5~15 mm. long; inflorescence
somewhat branched at base, the racemes very dense, the central one rather
many flowered; bracts, except the lowest, entire, hirsute, 5 mm. long;
pedicels hirsute, 1.5—6 cm. long; bracteoles linear, remote from flowers, ca.
2 mm. long; sepals dilute blue, white-hirsute without and within, the upper
12-13 mm. long, 7 mm. wide, the spur straightish or somewhat curved,
12-13 mm. long, 3.5 mm. wide at base, blunt; lateral sepals elliptic-oblong,
16 mm. by 7 mm., rounded at apex; lower sepals 13 mm. by 8 mm.;
petals black-violet, the upper laminae short-bristly on face, oblique, 11
mm. long, bidentate at the glabrous tip, spur 11 mm. long; lower laminae
yellow bearded on face, white-ciliate at tip, oblong-ovate, oblique, 5 mm.
long, bifid to below the middle, claw 6 mm. long; stamens 7-8 mm. long,
glabrous; anthers dark, oblong, 1.5 mm. long; carpels 3, densely hirsute;
seeds squamate, the scales coalesced in transverse rows.

Lectotype: Buchara oriental; inter Talbas and Sagridascht, 3000-3300
m., A. Regel, 1882 (LE); seen.

In 1940 Afanasiev published as a new species Delphinium rotundifolium
(Not. Syst. Herb. Inst. Bot. Acad. Sci. USSR. 8: 108) based on A fanasiev
227 from Turkestan, which he said was near D. oreophilum. So far as I
can judge from the description this is the case. I have seen no material
and therefore am tentatively reducing it to synonymy under D. oreophilum.

222. Delphinium oxycentrum W. T. Wang, Acta Bot. Sinica 10: 73.
1962.

Stems 4.5—7 dm. tall, 3.5—4.5 mm. in diameter, glabrous, simple or with
one branch above, the branch racemiferous; basal leaves ca. 2, long
petioled; leaf blades broadly round-reniform, 4-6 cm. long, 7-11 cm. wide,
open-cordate at base, 3-parted to beyond the middle, the parts imbricate,
the middle part 3-lobed into ovate incised-lobulate or coarsely toothed
lobes, the lateral parts unequally trifid, then lobed and toothed much like
the middle one, loosely strigulose on both surfaces or subglabrous above;
petioles 2-2.5 dm. long, loosely hirtellous, inconspicuously vaginate at
base; cauline leaves 1-2, like the basal but smaller, short petioled or
subsessile; racemes terminal, the main one 9-12-flowered, rather dense;
lower bracts 3-parted, others spatulate- to linear-oblong, 9-12 mm. long,
subglabrous or loosely hairy; pedicels 2—6.5 cm. long, stout, glabrous;
bracteoles apical, lance-linear, glabrous or loosely hairy; flowers erect or
nearly so; sepals subglabrous or loosely ciliate, the upper 21-25 mm. long,
17-22 mm. wide, the subulate spur 26-42 mm. long, sharply acute; lateral
sepals rounded or round-ovate, 14-17 mm. long, 11-14 mm. wide; lower
15-18 mm. long, 6-10 mm. wide; petals blackish brown, glabrous, the

156 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Ficure 18. Delphinium, Group IX. Petals, the upper spurred, the lower
clawed. A. D. nevskyi; upper lamina 8 mm. long, spur 11 mm.; lower la aug

ochotense: upper lamina 10 mm. lon ng, spur 17 mm.; lower lamina ong,
claw 6 mm.; drawn from Tiling near Ajan, Type (LE). C. D. oreophilum; Hace
lamina 10 mm. long, spur 11 mm.; lower lamina 5 mm. | mm.; dra

from Regel in 1382 ‘type (LE). D. D. poltaratzkii; upper lamina 12
i g, claw 6 mm.; drawn from Ludlow 762
l

lam a 5 mm. :

sles wn rp er oe ie Sherri reer ee E), D. pyramidatum; upper lamina 9
long, spur 12.5 mm.; lower lamina sf mm. lon ng, claw ge

“bof 194 (ny). H. D. eeraniicned upper lamina 8 mm. long, spur 9 mm.;

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 157

upper laminae 2-lobulate; lower laminae ovate, bilobed one-third their
length, the lobes triangular, claw 3 to 5 times as long; stamens 10-12 mm.
long; carpels 3, glabrous

Type: Szechwan, Kung-ka-ling, 1000 m., Sept. 1, 1937, Vii 13129 (pr),
not seen, photo. (£), seen

223. Delphinium poltaratzkii Rupr. in Osten-Sacken & Rupr. Sert.
Tian-shan. 37. 1869. (Mém. Acad. Pétersb. VII. 14: 4. 1869).
He, 18; D.

Perennial, the stem 2.5-5.5 dm. tall, robust, spreading-villous, not
glandular, more or less leafy; lower leaves with blades 5-7 cm. long, 6~12
cm. wide, subglabrous above, more or less villous beneath especially on
veins, 3- or 5-fid halfway to base or deeper, into broad-rhombic segments
shallowly lobed or toothed, the teeth ovate, 3-6 mm. long and wide, mostly
acute, mucronate; lower petioles 1-2 dm. long, scarcely dilated at base;
upper leaves gradually reduced; inflorescence branched, the terminal
raceme 4—10-flowered, the lateral 1—4-flowered, or the inflorescence simple,
subcapitate; bracts colored, 2-3 cm. long, the lower trifid, the upper entire,
1-2 cm. long; pedicels 1.5-4 cm. long; bracteoles colored, appressed to
sepals, linear-oblong, to 10 mm. long; sepals purplish mauve to saturated
blue, the upper sepal ovate, 15—20 mm. long, 10 mm. wide, obtuse, the spur
8-12 mm. long, 5 mm. wide at base; lateral sepals broadly obovate, 14-20
mm. long, 11 mm. wide; lower sepals 12-16 mm. by 9 mm.; petals almost
black, the upper laminae almost straight, 12 mm. long, bidentate, the spur
12 mm. long; lower laminae broadly elliptic, 7 mm. long, bearded, bifid
two-thirds its length into broad lobes, the claw 6 mm. long; stamens 6—7
mm, aes glabrous; anthers dark, 1.2 mm. long; carpels 3, somewhat
pubescent

Type: Turkestan, Tian-shan Mts., below Schamsi, Poltaratzki (LE);
seen.

The species apparently grows in open grassland at about 2500 m. It is repre-
sented by Ludlow 762 (Bm, £) from Koksu-Kensu Junction, Tian Shan,

lower lamina 4 mm. long, claw 6 mm.; drawn from Tarchevskaya & ——
ami

es I. D. ruprechtii; upper lamina 12 mm. long, spur 14 mm.; lower
long, claw 6 mm.; drawn from Alexeenko in 1898, TYPE (LE).
speciosum: upper lamina 14 mm . long, spur 20 mm.; lower mm. |
w 6 mm.; drawn from Kamennik in 1933 (Ny). K. hota upper
pe ina 10 mm. long, spur 20 mm.; lower lamina he mm. long, claw 6 m

Fp Clarke 4 44861 (us). L. D. tianshanicum; upper lamina 9 mm. long, spu

mm.; lower la ati “aly long, claw 6 mm.; drawn from Li & Chu 7003 ue
M D. villosum; upper lamina 9 mm. long, spur 12 mm.; lower lamina 3.5 m
long, claw 6 mm.; drawn from Gillett in 1939 (BH). of. winkleranun:
upper male 10 mm. long, spur 14 ee lower lamina 5 mm. long, claw 6 m
drawn from Winkler 7230, TYPE (LE)

158 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

224. Delphinium pyramidale Royle, Illustr. Bot. Himal. 56. 1834. :
Fic. 18, E,

Delphinium speciosum ce var. pyramidale (Royle) Mukerjee, Bull. Bot.
Surv. India 2: 293.

Delphinium chicka ype Wall. Cat. n. 4716. 1830, momen; Hook. f. &
Thoms. Fl. Ind. 51. 1855, based on Wallich 4716, Bhudrinak RB, seen (K).

Delphinium elatum L,. var. leaped all (Wall.) Hook. f. & Thoms. in
Hook. f., Fl. Brit. Ind. 1: 18

Delphinium speciosum Bieb. Sues esilibalapihias (Wall.) Brihl & King,
Ann, Bot. Gard, Calc. 5: 105. 18

Delphinium speciosum var. romunculifolium (Wall.) Huth, Bot. Jahrb. 20:
416. 1895.

Delphinium speciosum var. bruehlianum Huth, Bot. Jahrb. 20: 416. 1895, new
name for D. elatum var. ranunculifolium in Hook. f. Fl. Brit. Ind. 1: 26.

1872.

Delphinium speciosum subsp. ranunculifolium vars. brachycentra, eriantha,

katatricha, lasiantha, ale sang apes pulverulenta and thyrsantha

aie

Brihl in Brith) and King, Ann. Bot. Gard. Calc. 5: 106-7. 1896. These are
uncertain - to me. fie oo material has ae been available
to me on loa

Bie er ah te Chowdhury ex Mukerjee, Bull. Bot. Surv. India
61; nomen, I have seen material from Edinburgh annotated “pul-
Salli? by Chowdhury
? D. hoffmeisteri Klotzsch, Bot. Ergeb. Waldem. Reise 132. 1862. It has
been referred to D. elatum, but since it came from the Himalaya, it seems
south of the range of that species. No material seen.

Perennial from a rhizome, stem 6-12 dm. tall, 4-8 mm. thick, glabrous
to spreading-villous, usually leafy throughout, branched only in the
inflorescence; lower petioles to 4 dm. long, scarcely dilated at base,
glabrous to villous, the upper short; blades rounded-pentagonal in outline,
6-15 cm. wide, glabrous above and beneath, to subtomentose or villous,
especially beneath and along the veins, 3- or 5-partite 2/3 to 4/5 the way
to the base into broad cuneate segments, these cleft and incised-serrate,
the ultimate lobes ovate to lanceolate, mostly 2-7 mm. wide, mucronate;
upper leaves passing into foliose parted bracts, these lanceolate to lance-
ovate, to ca. 15 mm. long; inflorescence racemose or a group of racemes,
many flowered, variously pubescent; pedicels ascending, to ca. 5 cm. long;
bracteoles lanceolate, 3-8 mm. long, subtending the blue or purple flowers;
sepals pubescent without and within; upper sepal 15 mm. by 7 mm., ovate,
obtuse, spur straight or decurved, 12-15 mm. long, 3.5 mm. wide at base,
gradually narrowed to a blunt apex; lateral sepals rhombic-ovate, 17 mm.
by 10 mm., subtruncate at apex; lower sepals 16 mm. by 8 mm., obtuse
narrow-obovate: petals bluish black, the upper laminae slightly “ohliie,
10 mm. long, bidentate at apex, the spur 14 mm. long; lower laminae
elliptic-oblong, 6.5 mm. by 3.5 mm., cleft to the middle into lance-oblong
lobes long-ciliate, eset outside and with patch of long hairs at inner base
of lamina, claw 7 mm. long; stamens 7-8 mm. long, with strongly dilated
glabrous filaments; anthers dark, 1 mm. long; follicles mostly 3, glabrous

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 159

or pubescent, more or less divergent, 12-15 mm. long, 3-4 mm. wide, the
styles an additional 3 mm. long; seeds blackish, obpyramidal, 1.5 mm.
long, densely clothed with laminate scales.

Type: Peer Punjal (Pir Panjal), Royle. Through the very great kind-
ness of Mr. E. F. Greenwood, Keeper of Botany, City of Liverpool
Museums, I have had photographs of the type of Delphinium pyramidale.
This name has to replace the name D. ranunculifolium which was for many
years a nomen nudum.

Rance. Apparently from West Pakistan to Nepal at elevations of about
2000 to 3500 m.

ILLUSTRATION. Briut & Kino, Ann. Bot. Gard. Calc. 5: pl. 127, 1896;
CoventrY, Wild Fl. Kashmir 2: pl. 6. 1927

EXAMPLES SEEN. Swat: R. R. Stewart 29167 (micH). Kumaon: Strachey &
Winterbottom 2 (cH, K). Kashmir: Duthie Aug. 25, 1892 (BM, E); C. B. Clarke
29506 (BM, K); R. R. & I. D. Stewart 18357 (GH, Ny); Stewart 18316 (uc, US),
8079 (NY), 5268 (PHIL), 28781 (E); Duthie 19114 (x); Koelz 9313 (NA, NY);
Jacquemont 908 (Pp). Nepal: Sharma E98 (pm); Dhwoj 234 (e); F. M. Bailey
in 1936 (E). W. Tibet: Falconer 62 (cH, K, P); Polunin, Sykes & Williams 5120
(Bm) ; Polunin 2629 (BM).

225. Delphinium pyramidatum Alboff, Descr. Nouv. Esp. Pl. ( 1891)

II. (Trav. Soc. Hort. Odessa 1891); Busch in FI. Cauc. Crit. III.

3: 71. 1962. Fic. 18, G.

Delphinium elatum L. var. pyramidatum (Alboff) Huth, Bot. Jahrb. 20: 399.
1895

Delphinium flexuosum var. glandulosum O. Kapeller ex Grossh. Fl. Cauc. 2:
00. 1930, according to Nevski in Fl. U.S.S.R. 7: 142. 1937.

Stem 1—1.5 m. tall, robust, leafy; leaves 5- or 7-parted, the parts deeply
incised into narrow-lanceolate acuminate lobes; inflorescence pyramidate-
paniculate to ca. 4 dm. long, glandular-pubescent and strigulose, much
branched and with very numerous crowded flowers; bracts narrow-linear,
to 2 cm. or more long; pedicels 3-5 cm. long, ascending, pubescent; brac-
teoles linear, 10-12 mm. long, placed just above the middle of the pedicels;
calyx pale blue, more or less puberulous; upper sepal ovate, obtuse, 12
mm. long, 8 mm. wide, the spur decurved, 10 mm. long, 3 mm. wide at
base; lateral sepals asymmetric, oblong, 17 mm. by 7 mm., rounded at the
broad apex; lower sepals lance-ovate, 14 mm. by 4 mm., pointed; petals
blackish brown, the upper laminae erect, glabrous, bidentate at apex, 9
mm. long, the spur 12.5 mm. long; lower laminae oblique, 5 mm. long,
cleft to middle, more or less bearded, the claw pubescent, 6 mm. long;
stamens 6.5 mm. long, glabrous; anthers dark, oblong, 1 mm. long; follicles
3, glabrous; seeds winged at angles.

Type: West subalpine Transcaucasia, Abkhasia, 1300-1400 m., Aug.,
1893, Alboff 195, isotype (P); seen.

ADDITIONAL COLLEcTIONS. Alboff 194 (Ny), 196 (Pp), 182 (K).

160 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

226. Delphinium retropilosum (Huth) Sambuk, Soc. Bot. Russ. 14:
428. 1929 Fic. 18, H.

Delphinium duhmbergii Huth var. retropilosum Huth, Bot. Jahrb. 20: 402.
1895

Delphinium pterospermum Turcz., nomen, according to Nevski in FI. YS5R
+ 152. 1937.

Stem 6-8 dm. tall, 6 mm. thick, striate, densely clothed on lower half
with white retrorse-spreading hairs 2-3 mm. long, subglabrous above, few
branched in inflorescence, rather closely leafy; lower leaves gone by
anthesis, main cauline petioles hairy, 5-8 cm. long, dilated at base; blades
roundish-elliptic in outline, almost straight across the base, 4-5 cm. long,
6-8 cm. wide, green and glabrous above, conspicuously veined and paler
and hairy beneath, palmately 5-fid to 1 or 1.5 cm. above the base into
cuneate obovate segments laciniate into 3 main lobes, these toothed or
incised into lanceolate lobules 2-3 mm. wide and up to 10 mm, or more
long; inflorescence a long, many flowered raceme with short branches
below, subglabrous; lower bracts more or less foliose, palmately divided,
15-20 mm. long, with some long bristly hairs; upper bracts lacking;
pedicels erect, 1-2 cm. long, angled; bracteoles oblanceolate, 5-8 mm.
long; strongly ciliate, placed near the base of the flower; sepals blue,
mostly glabrous except for a few marginal hairs; upper sepal ovate, 10
mm. long, 5-6 mm. wide, the spur straight, 1 cm. long, 2.5 mm. wide at
base; lateral sepals more or less elliptic, 8 mm. long, 4 mm. wide, rounded
at apex, with a few marginal hairs; lower sepals 9 mm. by 5 mm., obtuse;
petals dark, the upper laminae slightly oblique, entire, 8 mm. long, with a
few bristly hairs, the spur 9 mm. long; lower laminae oblong, 4 mm. by 2
mm., unequally lobed, with long yellow hairs on the surface and long
white cilia, claw 6 mm. long; stamens 5 mm. long, glabrous, moderately
dilated; anthers dark, 1 mm. long; follicles 3, somewhat pubescent.

Type: Indicated only as coming from Siberia.
ILLusTRATION. Fl. U.S.S.R. 7: pl. 10. 1937, drawing of leaf.

I would place two herbarium specimens that I have seen (both from ny, since
I was unable to borrow any from LE) in this species: North Altai, Tarchevskaya
& Vandysheva, July 10, 1932, and Tomsk, without collector, in 1913.

227. Delphinium ruprechtii Nevski in Komarov, Fl. U.S.S.R. 7: 138,
723. 1937. Fria. 18; 1

Delphinium flexuosum var. bracteolatum N. Busch in Fl. Cauc. Crit. III. 3:
67. 1903, pro parte, according to Nevski.

Stem to | m. tall, ribbed, lightly pilose or almost glabrous, ridged, leafy;
petioles not dilated at base, pilose, to ca. 1 dm. long; leaves like those of
D. flexuosum, scarcely pubescent above, to 12 cm. wide, pilose on nerves
beneath, palmatifid almost to base, the middle segment oblong-rhombic,
divided into 3 lobes, each with 3 to 5 lanceolate teeth, lateral segments

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 161

like the middle one, the ultimate teeth 5-15 mm. long, 3-6 mm. wide;
raceme dense, usually branched, the axis glabrous, many flowered, to 3
dm. long; bracts lanceolate, more or less ciliate on margin, 10-15 mm.
long, 2-4 mm. wide, somewhat bluish; pedicels glabrous, hen 1-4
cm. long; bracteoles elliptic or elliptic-ovate, 5.5-8 mm. long, 4—-4.7 mm
wide, subobtuse, violet tinged, glabrous; sepals glabrous without, ciliolate
within, the margin densely ciliate, blue, the upper oblong-ovate, sabacite,
20 mm. by 10 mm.; spur 18-20 mm. long, 3.5 mm. wide at base; lateral
sepals elliptic, 20 mm. by 10 mm., obtuse; lower sepals somewhat nar-
rower ; petals black, the upper laminae almost straight, 12 mm. long, deeply
bidentate, glabrous, the spur 14 mm. long; lower laminae 7 mm. long,
oblong-ovate, short bearded at base, bifid to middle into oblong, ciliate
lobes, claw appendiculate. 6 mm. long; stamens 7-8 mm. long, glabrous;
carpels 3, glabrous.

Type: ‘“Dagestania, inter pagos Czirch & Rhycza,” leg. Alexeenko,
July 23, 1898 (LE); s

Differs from Delphinium flexuosum and D. bracteosum in its elliptic
or elliptic-ovate glabrous bracteoles.

228. Delphinium sauricum Schischk. Animadvers. Syst. Herb. Univ.
Tomsk 1936(8): 1. 19

Stems solitary, erect, 3-5 dm. tall, from a thick rhizome, with longish
spreading hairs; leaves petioled, these 6-14 cm. long, densely hirsute;
blades round-reniform, deeply cordate at base, 4-6 cm. long, 2.5-4 cm.
wide, deeply divided or coarsely toothed, with long appressed hairs on
both surfaces; inflorescence racemiform, 1—2.5 dm. long, 6—10-flowered;
lower bracts 2—3-lobed to middle, shorter than or subequal to the pedicels,
upper bracts entire, Jance-ovate, violet; bracteoles 2, lance-ovate, to 10
mm. long, 3-4 mm. wide; sepals blue-violet, more or less appressed hairy
on both surfaces and mixed with a few lageniform hairs, broadly ovate,
14-15 mm. long, 10-11 mm. wide, the lower narrower, 15 mm. by 8 mm.;
spur 20-23 mm. long; petals black-fuscous, the upper laminae 10 mm.
long, 4-5 mm. wide, the lower deeply bifid, ciliate, yellow bearded on
surface near the base; carpels 3, strigose.

Type: Montes Saur. In jugo Mus-tau toward head of Ulkun-Obo River,
in alpine meadows, July 26, 1914, W. Saposhnikov (tk); I was unable to
borrow material.

229. Delphinium speciosum Bieb. Fl. Taur. Cauc. 2: 12. 1808.
Fic. 18, J.
spe deanery speciosum var. typicum biebersteinii and speciosum var. ste
and var. trichocarpum Rupr. Fl. Cauc. 35. 1869, fide Nevski in Fl. U.S.S.

7, ee : :
Delphinium ironorum N. Busch, Bull. Acad. Sci. U.S.S.R. 1931: 450. 1931.

162 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Based on a collection by E. & N. Busch, Aug. 24, 1930, at 2750 m., from
Dzhomakh, Mount? or Mountain Pass? Bakh-fandar (LE); seen
Delphinium bracteosum var. macranthum Som. & Levier, and var. albidum
Som. & Levier, Acta Horti Petrop. 13: 25. 1893; according to Nevski.
? Delphinium caucasicum var. bracteosum Schrenk & Trautv. Bull. Soc. Mosc.

? Delphinium speciosum var. bracteosum Huth, Bot. Jahrb. 20: 416. 1895.

? Delphinium boissieri Boiss. & Buhse, Nouv. Mém. Soc. Nat. Mosc. 12: 8.
1860. Delphinium speciosum var. boissieri Buhse, Reise Turcm. u. Pers.
p. 8. “Beim Dorfe Livan auf der Thalsohle,” no. 539, June 18, 1947. Col-
lector? Not seen.

Perennial from a woody root, the stem 3-8 dm. tall, simple or branched
above, villous, striate, rather equably leafy throughout; leaves 3-12 cm.
wide, rounded-pentagonal in outline, more or less villous on both surfaces,
the hairs especially long on the veins of the upper surface, more generally
distributed beneath, blades mostly 5-parted to near the base into broad
cuneate segments, these incised-serrate into lanceolate lobes 2-10 mm.
wide, long pointed; petioles somewhat dilated at base, villous, 3-15 cm.
long; flowers racemose on axis or branches, the inflorescence densely vil-
lous, several to many flowered; lower bracts 3-parted to the base, the upper
entire, lanceolate, villous, 7-15 mm. long; pedicels almost erect, 1—3 cm.
long; bracteoles lanceolate, villous, 4-6 mm. long, placed just beneath the
flower; sepals violet, villous without and somewhat within, upper sepal
ovate, 18-25 mm. long, 8 mm. wide, rather sharply pointed, the spur 17-18
mm. long, 3 mm. wide at base, tapered rather rapidly to the narrow, some-
what decurved apex; lateral sepals 18-25 mm. by 10 mm.; lower sepals 20
mm. by 9 mm.; petals black, the upper laminae straight, 14 mm. long,
deeply emarginate, glabrous, the spur 20 mm. long; lower laminae 8 mm.
long, 4 mm. wide, oblong, turned to one side, with a patch of long yellow
hairs near the base, white-ciliate, cleft ca. 3 mm., the claw broad, 6 mm.
long; stamens glabrous, 6 mm. long; anthers dark, oblong, 1.2 mm. long;

rpels 3, somewhat villous, becoming 15 mm. long; seeds dark, trans-
versely lamellate-squamate, ca. 2 mm. long.

Type: Mt. Kaischaur, Caucasus, Herb. M. a. Bieberst., plant in photo-
graph (K) has 1 leaf and is in fruit. Another specimen labeled Delphin-
tum speciosum, M. Comm. Stephan, ex Herb. Bieberstein, typus (photo.
K

ILLUSTRATIONS. DELESSERT, Icon. Select. 1: pl. 62. 1820; Bot. Reg.
18: pl. 1503. 1832.

DisTRIBUTION. The species seems to range from the Caucasus to Turke-
stan,

ExAMPLes. Hisiar, in valle fl. Jaquob, A. Regel, Iter Turkestanicum (£, K);
Caucasus: Bunge in 1856 (P); South Ossetia, E. & K. Busch in 1935 (BM);
Patcha Pass, Seraischeniar, Komarov in 1892 (x); Circassia, source of the
Lascha, Alboff in 1894 (P).

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 163

230. Delphinium stapeliosum Briihl ex Huth, Bot. Jahrb. 20: 419.
1895 and in Briihl & King, Ann. Bot. Gard. Calc. 5: 102. 1896.
Fic. 18, K.
sernen Stapeliosum var. khasianum Brihl in Brihl & King, Ann. Bot.
ard. Calc. 5: 102. 1896.

Stem erect, slender, 4-15 dm. tall, branched above, retrorse-pubescent,
scattered leafy, the upper leaves reduced: lower petioles to 2.5 dm. long,
slightly dilated at base, pubescent, the upper shorter; leaf blade to 15 cm.
across, palmately 5-fid to 1 cm. or more from base, deep green and sparsely
pubescent above, paler and more pubescent beneath especially on the veins,
the segments broadly cuneate, entire in lower part, trifid above and then
incised-dentate with large lance-ovate or broader teeth, acute; racemes
lax, rather few flowered or the terminal raceme with more; bracts lance-
olate to lance-linear, to ca. 10 mm. long; pedicels ascending, gray-pubes-
cent, 1-8 cm. long: bracteoles lance-linear, 7-10 mm. long, alternate,
near the middle of the pedicel; sepals blue to violet or reddish, hairy
without, the upper sepal ovate, 18 mm. by 10 mm., obtuse, the spur slightly
curved, 20 mm. long, 4 mm. wide at base, the apex fairly slender; lateral
sepals rhombic-ovate, 18 mm. by 15 mm., rounded-truncate at apex; lower
sepals somewhat narrower, 17 mm. by 10 mm., obtuse; petals blue-black,
the upper laminae almost straight, 10 mm. long, glabrous, bidentate at
apex, the spur 2 cm. long; lower laminae oblong, 6 mm. by 3 mm., deeply
cleft, bearded but not ciliate, the claw 6 mm. long; stamens 5—6 mm. long,
glabrous; anthers dark, linear, 1.3 mm. long; follicles 3, pubescent, diver-
gent, slender, 12-18 mm. long, the styles an additional 3.5 mm.; seeds
obpyramidal, winged along the edges, puckered on the sides.

Lectotype: No specimens are cited in Briihl & King, but in Huth there
is cited “Assam: Khasia Hills ad 2000 m. alt.,” Hooker and peeks ae
would select the K specimen as the lectotype and cite also BM, E, GH, NY

DIstTRIBUTION. Eastern Nepal and Khasia, Assam, at elevations ws about
1200 to 3000 m.

REPRESENTATIVE SPECIMENS. E. Nepal: Stainton 1162 (Bm); Stainton, Sykes
& Williams 7867 (pm), 8587 (BM). Khasia: Clarke 16354 (BM, K), 16117 (BM),
pag (E), Had (K), 38970 (us), 40301 (us), 44861 (US); Hooker & Thomson

M, E, GH, K, NY); Griffiths 461 (BM, K), 827 (K); Chaud 7192 (£, MICH),
51880 eas Koel 25579 (MIcH), 31250 (rE, MICH).

254. <3 aaron tianshanicum W. T. Wang, Acta Bot. — AE es
IG.

Lae 5-11 dm. tall, terete, 2-5 mm. thick at base, sparsely retrorse-
hirsute, few branched above, mostly simple, equably foliose; lower leaves
long petioled (to 5 times as long as the blades) ; blades as in D. naviculare,
broadly reniform or pentagonal-reniform, 3-9 cm. long, 5-14 cm. wide,
cordate, 3-parted to 1 or 2 cm. from the base, the middle part rhombic,
acute, 3-lobed above the middle, the lobes somewhat ovate, coarsely 4—-5-

164 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

toothed, the lateral parts smaller, 2-3-toothed, densely strigulose on both
surfaces; racemes 7—14-flowered, rather dense; lower bracts trifid, the
others lance-linear, 8-14 mm. long, 1-1.5 mm. wide, strigulose; pedicels
2-6 cm. long, densely retrorse-strigulose, suberect; bracteoles 1.5-6 mm.
below the flower, linear to lance-linear, 5-9 mm. long, strigulose; sepals
blue-purple, ovate or obovate, 11-15 mm. long, densely strigulose; spur
11-14 mm. long, cylindric-subulate, 3 mm. thick at base, straight or
slightly decurved; petals black, the upper laminae glabrous, emarginate,
9 mm. long; lower laminae subobovate, 5 mm. long, 2-lobed to near the
middle, claw 6 mm. long, short-appendiculate; stamens glabrous; anthers
over 1 mm. long; follicles 9-11 mm. long, densely strigulose, the styles 2
mm. long; seeds obconic, triquetrous 1.5 mm. long, scaly-lamellate.

Type: Sinkiang, T. N. Liou 3620, Aug. 24, 1931 (PE); not seen. Other
specimens cited in the original description are also from Sinkiang, Ho-
shuo, 2300 m., Li & Chu 7003 (£), seen, and Sha, 1700 m., Kuan 2290
(£), seen.

232. Delphinium ukokense Sergievsk. Animadvers. Syst. Herb. Univ.
Tomsk 1954: 9. 1955.

Erect perennial, 2.5-4 dm. tall, 2-4 mm. thick below, leafy almost to
the inflorescence, short-pubescent throughout, with gland-tipped hairs
mixed in; leaves 3.5-6 cm. long, 4-6 cm. wide, round-reniform or rounded,
3- or 5-parted to 4/5 or 5/6 their length, the lobes rhombic, coarsely
toothed above, short-pubescent on both surfaces; raceme 10-12 cm. long,
lax, 6-15-flowered; lower bracts leafy; pedicels 5-10 mm. long; bracteoles
linear; sepals blue, pilose; spur. slightly curved at tip, 14-15 mm. long;
petals brown, the upper laminae glabrous, the lower yellow-pubescent
down to middle, ciliate; carpels 3, glabrous.

Type: Altai, Ukok, Saposhnikov in Herb. Krylov (rk); not available
o me.

The author compared his species, Delphinium ukokense, with D. mira-
bile, but said the stems of the former were glandular-pubescent their
whole length, the petioles shorter than or equal to the laminae; the flowers
smaller; spur 14-15, not 16-20 mm. long; and the carpels glabrous.

233. Delphinium villosum Stev. ex DC. Syst. 1: 546. 1217; Prodr- 1:
55. 1824 Fic. 18, M.
Delphinium villosum var. anomalum Huth, Bot. Jahrb. 20: 412. 1895. Huth
cited “in deserto Tentek,” Schrenk 1840 (LE); not seen.

Delphinium villosum var. winkleri Huth, loc. cit. Huth cited Alexander Mts
Dschilamysch, 1600-2000 m., Fetissow, 1880 (LE): not seen :
Delphinium rha Schipcz. ex Nevski in Komarov, Fl. USSR. 7: 152. 1937
on,

Stem 10-12 dm. tall, leafy, simple, glabrous below; petioles scarcely

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 165

dilated at base, to 15 cm. long, the upper reduced; blades glabrous or
slightly pubescent on veins above, subvillous beneath, the lower truncate-
cordate at base, the upper cuneate, all broader than jong, the lower to 15
cm. broad, 3- or 5-parted to 1 cm. or more from the base, the parts cuneate-
obovate, shallowly incised or coarsely toothed, with acute or acuminate
teeth or lobes; raceme simple or few branched, many flowered, rather lax,
2-4 dm. long, minutely strigulose and with some retrorse-spreading ie
pubescence; bracts mostly linear-subulate, villous, to ca. 1 cm. lon
pedicels erect-divergent, 1-7 cm. long; bracteoles like the bracts, es
slightly below the flowers; sepals deep blue, sparsely villous; upper sepal
12-15 mm. long, 6-8 mm. wide, acute, the spur straight, about as long,
rather slender; lateral sepals oblong-obovate, 12-14 mm. long, 6-8 mm.
wide, villous along midrib, rounded at apex; lower sepals more pointed
and more hairy; petals dark, the upper laminae almost straight, glabrous,
9 mm. long, entire, the spur 12 mm. long; lower laminae oblong-ovate,
3.5 mm. by 2.5 mm., long-bearded, cleft, the claw 6 mm. long; stamens
6—7 mm. long, the filaments ciliate; anthers dark, 1 mm. long; follicles 3,
glabrous except on the ciliate sutures, oblong, 12- 15 mm. long, 4-5 mm
wide, the styles an additional 3 mm.; seeds brown, 2.5 mm. long, slightly
winged along the 3 angles.

Typr: Hort. Dorpat, Steven in 1817 (G?) not seen; photograph (BH),
seen,

EXAMPLES OF THE SPECIES. Cultivated at Ottawa from Tabor Bot. Gard.;
Germany: Gillett, July 7, 1939 (BH); Chinese Turkestan: Mts. Bogdo-ola &
Urumtschi, Merzbacher 1304 (cu).

234. Delphinium winklerianum Huth, Bot. Jahrb. 20: 419. 1896.
Fic. 18, N.
Perennial, 6-8 dm. tall, with scattered stiff white hairs, rather equably
leafy; midcauline leaves with petioles 2-8 cm. long, villous, scarcely
vaginate, the blades ca. 1 dm. across, broadly cordate at base, 3-parted to
a. 1.5 cm. from base into broad-cuneate obovate segments ca. ° cm. by
3-3.5 cm., scattered stiff-hairy on both surfaces, each segment with ca. 3
coarse ovate teeth to 1.5 cm. long and broad, mucronate, the uppermost
leaves reduced; inflorescence with 2-3 basal branches, the central raceme
to ca. 3 dm. fone. laxly ney flowered, somewhat strigulose, the pedicels
suberect, 3-10 cm. long; lower bracts foliose, upper lance-linear, 1—1.5
cm. longs bracteoles cibteading the flower, lanceolat te, 8-10 mm. long,
2—2.5 mm. wide; flowers deep blue, sepals strigulose ‘without, glabrous
within, the upper sepal ovate, 17 mm. by 10 mm., obtuse, the spur spread-
ing decurved, 17 mm. long, 3.5 mm. wide at base, blunt; lateral sepals
broadly elliptic- obovate, 18 mm. long, 14 mm. wide, rounded at apex;
lower sepals elliptic-ovate, 16 mm. by 8 mm., subacute; petals black, the
upper laminae almost straight, 10 mm. long, rounded and bidentate at the
glabrous tip, the spur 13 mm, long; lower laminae strongly oblique, ob-

166 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Jong-obovate, 5 mm. by 3.5 mm., somewhat bearded, bifid about halfway
into rounded-oblong lobes, the claw 6 mm. long; stamens 7-8 mm. long,
the anthers dark, rounded, ca. 1 mm. long; carpels 3, hairy.

Type: June 4, 1881, Kasch, Turkestan, sem. m. Dr. A. Regel, C. Wink-
ler 7230 (LE); seen.

[To be continued |

VoLuME 49 NuMBER 2

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CONTENTS OF NO. 2

3 FLowErs, Fruits, AND PHYLOGENY oF ALANGIACEAB. Richard
H. Eyde

Diverse Nopat Types in Ruopopenpron. W. R. Philipson and
M. N. Philipson

A New Hyprancrea From Mexico. Lorin i. ges Jr. and
Arturo ie Sears aS

A Synopsis oF THE ASIAN SPECIES OF _Daveansro, SENsU
Srricro (Concluded). Piebp'A. ee :

MorPHoLocy AND ANATOMY OF Croomia PAUCIFLORA (STEMONA-
ceaE). P. B. Tomlinson and E. 8. Ayensu

K. Brizicky

A, AND Gavowss epider ale George

JAMATCAN AND orien SPzorEs oF BUMELIA (Saroracean). William
‘iT. Stearn

167

JOURNAL

OF THE

ARNOLD ARBORETUM

VoL. 49 Apri 1968 NUMBER 2

FLOWERS, FRUITS, AND PHYLOGENY OF ALANGIACEAE

RicHaArpD H. Eype

PUBLICATIONS ON THE ALANGIACEAE include an excellent monograph
(Bloembergen, 1939) and a series of carefully documented anatomical
studies (Govindarajalu & Swamy, 1956; Govindarajalu, 1961; 1962 a, b,
c), but details of the flower and fruit have never been adequately scru-
tinized for evolutionary trends or for evidence of extra-familial affinities.
This article will emphasize three neglected features that are known to have
taxonomic value in many groups of plants: the floral vascular system, the
secretory canals, and the stone or endocarp portion of the fruit wall.
Species groups within the Alangiaceae differ greatly with respect to these
structural features, and some of the differences can be interpreted to show
directions of evolutionary change.

Although many older botanical works recognized two genera of Alan-
giaceae, Alangium Lam. and Marlea Roxb., Bloembergen’s (1939) opinion
that Marlea deserves only sectional status is now followed in most treat-
ments. Bloembergen’s monograph lists 18 species of Alangium, all woody,
distributed in eastern Asia, Africa, Australia, and islands of the Pacific
and Indian Oceans. Japanese botanists accept an additional species, A.
premnifolium Ohwi (1938, p. 30, 1965, p. 652), considered doubtful by
Bloembergen (p. 172, 173). Three species have been described since
Bloembergen’s monograph appeared. Two of these, 4. chungii Li (1943,
p. 448) and A. tonkinense Gagnepain (1950, p. 22 ), are based on single
collections, and their circumscriptions differ so little from those of better
known species ! that an anatomical study need not be concerned with them.
The third, A. grisolleoides Capuron (1962) is sufficiently distinct to merit
a special commentary later in this paper.

Bloembergen distributed the species in four sections, using the number
of stamens and characters of style, stigma, and embryo as basis for dis-
tinction. Govindarajalu’s anatomical observations strongly support the
soundness of this distribution. Although I believe the order in which

1 Alangium tonkinense differs from A. salviifolium mainly in that its leaves are
cuspidate and its filaments not dilated. I have examined the type of A. chungii (Chung
82038, 4) and I think it belongs to A. kurzii Craib. The only fertile material labeled
A. premnifolium that I have seen, a flowering specimen from Okinawa (Kanashiro
692, a), also belongs to A. kurzii.

168 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Bloembergen listed the sections obscures their evolutionary relationships,
I follow his delimitation of taxa throughout this paper. I also adhere to
his nomenclature and orthography, with two minor exceptions: (1) section
Ancotum Baill., which includes the type species, is herein called section
ALANGIUM in accordance with Article 22 of the current /nternational Code
of Botanical Nomenclature (1966), and (2) the specific epithet salviifo-
lium is spelled according to Recommendation 73G of the Code.

MATERIAL

Cooperating botanists and foresters (see Acknowledgments) supplied the
following collections in liquid preservative.

Section ConosticMA Bloemb.

A, havilandii Bloemb. SAR S.22767; Sarawak

A. ridleyi King (1) cult., Bogor, VIII. H. 14
(2) cult., Bogor, 7X. A. 20.

A, javanicum (Bl.) Wangerin (1) P.P. Sam, s.n. (SAN 36750).

(2) cult., Bogor, VIII. F. 25.

Section ALANGIUM
A, salviifolium (L.f.) Wangerin (1) Govindarajalu 6840A;
adras, i
(2) cult., Bogor, Indonesia,
IIT. G. 41.
(3) cult., Bogor, XVII. C. 136.
A. longiflorum Merr. (1) B. Hernaez 635; Makiling

Natl. Park, Philippines
(2) without voucher; Makiling
Park

Section MARLEA Baill.
A. platanifolium (S. & Z.) Harms (1) cult., Bot. Gard. Tohoku Univ.
(2) K. Soke; s.n., Shiraishi
Pass, Twate Prof Japan

A. chinense (Lour.) Harms (1) cult., Tjibodas, Indonesia
(2) without voucher; Makiling
Natl. Park
A. rotundifolium (Hassk.) Bloemb. cult., Tjibodas

To verify the identifications, I consulted Bloembergen’s descriptions
and compared the material with herbarium specimens that Bloembergen
annotated or cited. Some of the collections are from individual trees that
Bloembergen mentioned and that still survive in Indonesian gardens. The
infraspecific identity of the specimens seems unimportant to this investiga-
tion; so I have not tried to make subspecific or varietal distinctions,

Unable to obtain pickled material of A. villosum (Bl.) Wangerin, the

1968 | EYDE, ALANGIACEAE 169

only species in Bloembergen’s section RHYTIDANDRA, I removed a few
dried flowers from sheets in the U.S. National Herbarium. (Because of
Dr. A. C. Smith’s field activities, the Herbarium is well supplied with the
Fijian variant of A. villosum.) My remarks on mature fruits of Alangium
are drawn mostly from the examination of dried specimens in the U.S.
National Herbarium, and in the Gray Herbarium and the Herbarium of
the Arnold Arboretum of Harvard University.

Sectioning and staining procedures were routine (Eyde, 1963, 1966),
but clearing of flowers required some experimentation. Good results were
obtained through Fuchs’ (1963) ‘Method No. 1,” with the concentration
of NaOH reduced to 5 percent to prevent maceration of tissues. Another
successful procedure included the following steps: overnight immersion
of the flowers in 5 percent NaOH at 60°C, immersion for several hours in
warm “CLP” (Bersier & Bocquet, 1960), transfer through an ethanol series
to toluene. In most cases, it was necessary to cut the flowers into thick
freehand sections before clearing.

OBSERVATIONS

General. Bloembergen (1939) followed Wangerin (1910) and several
earlier authorities in using the relative number of stamens and petals to
subdivide Alangium. Stamens of Bloembergen’s sections Maries, RHYTI-
DANDRA, and ConosticMA — collectively corresponding to subgenus Mar-
LEA of Wangerin’s treatment — were said to equal the petals in number.
Neither Bloembergen nor Wangerin added that the number of stamens
may be one more or one less than the number of petals. I have recorded
the following counts for corolla (C) and androecium (A). A. havilandii:
C4, A4: C5, AS; C6, AS. A. ridleyi: C5, AS; C6, A6. A. javanicum:
C6, A6: C7, A7; C8, A8. A. grisolleoides: C4, A3; CS, AS. A. platan-
ifolium: C6, A7; C7, A7. A. chinense: C7, A7; C8, A8; C8, A9. A,
rotundifolium: C7, A7; C8, A8. A. villosum: CS, AS: C5, A6; C6, Fe
C6, A6. For section ALANGIUM (“subgenus Eualangium”) Wangerin gave
the number as two to four times the number of petals; Bloembergen's
description says two to six times the number of petals. My observations,
which follow, indicate that Wangerin was more nearly correct. A. salviifo-
lium: C5, A12; C5, A17; C6, Al5; C6, A26; C7, A26; C7, A28; C8, A30.
A, longiflorum: C4, A14; C4, Al6. With limited material, I have not
tried to estimate the frequency with which any of these numbers occur.
Not uncommonly, flowers are found in which one petal is “double ; Le., it
is seen as an abnormally broad single appendage when sectioned near its
base but as two normal appendages in distal sections. This condition is
not restricted to any particular group of species. I also found one flower
of A. grisolleoides (C4, A3) in which a stamen with six anther sacs and two
vascular bundles is clearly a double structure.

Regardless of their number, the stamens are arranged in
around a prominent pulvinate disc. De

latrorse to somewhat introrse. As Bloembergen emphasized, variations 1n

170 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

the form of the filaments and in the length of their trichomes are important
for separating the species. A distinctive character of section CONOSTIGMA
is the papillose epidermis of the anther.

The microspore mother cells in a developing anther are surrounded by
four cell layers: an inner tapetum, an outer epidermis, and two undif-
ferentiated intermediate layers. The subepidermal layer never develops
the “fibrous” thickenings of a typical endothecium. Another unusual fea-
ture, already reported by Gopinath (1945), is the arrangement of micro-
spore mother cells in a single file in each anther lobe; this is caused by the
conversion of primary sporogenous cells into microsporocytes without in-
tervening divisions. Following meiosis, all surrounding layers degenerate
except for the thin-walled epidermis, which usually ruptures before the
corolla opens. In most species the septa between anther-halves also degen-
erate before the pollen is released, but anthers of A. /ongiflorum retain four
locules at anthesis.

The trichomes and crystal inclusions of Alangium flowers are described
in detail by Govindarajalu (1962 b, c). My observations agree with his
except that I have found druses to be plentiful in floral tissues of section
ConosticMA, where he found none. Included in my material is one species,
A. havilandii, that Govindarajalu did not examine: the floral trichomes are

Fic. 1.
1X AL 20, S35.
$.n., X 4.3. Part of each flower has been cut away to show disc

a, flower of Alangium ridleyt (section Conosticma), cult., Bogor,
wer of A. chinense (section Marea), cult., Tjibodas,

1968 | EYDE, ALANGIACEAE 171

of the type that he called “stellate-peltate,” as could have been predicted
from his observations on the other members of section CoNosTIGMA.

Sectional differences in the form of the stigmatic area were stressed by
Bloembergen. He named section Conosticma for the “conical’’ termina-
tion of the style, but the four stigmatic rays (Fic. 1, a; Fic. 2, a) give it
a pyramidal appearance to my eye. In section Marea (Fic. 1, b) the
stigma is capitate and lacks the rays, but is distinctly four-lobed. Section
ALANGIUM has a capitate stigma that is not four-lobed — here I contradict
Bloembergen — but irregularly folded and convoluted (Fic. 2, b). Alan-
gium villosum (sect. RHYTIDANDRA) is distinguished by two styles, each
stigmatic along its ventral surface and slightly bifid at its apex. These are
basally united for about half their length (Bloembergen, 1939, Fig. 7, b, k)
in a stylar column. Microtome sections through the stylar column of A.
villosum reveal a symmetrically four-lobed stylar canal. When cut about
halfway between base and apex, the style in section CoNosTIGMA has a
similar cross section, but in section Marvea the four-lobed appearance is
less symmetrical, and in section ALANGIUM the stylar canal is very irreg-
ular, often three-lobed. Followed downward through serial sections, the
stylar canal of any species loses its lobed appearance, becoming oval or
elliptical in outline at its base (Clarke, 1850, Figs. 4-7). It is always
hollow at anthesis and filled with a mucilaginous exudate.

The ovary, completely inferior throughout the genus, is one-locular in
sections ConosticMA, ALANGIUM, and RHYTIDANDRA. In some species of
section MARLEA two-locular ovaries are common, but the locules are usually
unequal, and one of them may be empty (Fic. 5, b). Except for such
cases, the ovules are solitary in each locule, pendent and anatropous with
the micropyle directed laterally (Fic. 2, c; 5, a).

Vascular system. The most conspicuous sectional differences are in
the number of carpel bundles and in the extent of fusion of calyx bundles
to bundles supplying petals and stamens. The vascular systems of sec-
tions ConosticMa and ALANGIUM differ somewhat (Fic. 4), but the dif-
ferences that separate these two groups from sections Mar-EA and Ruy-
TIDANDRA are much greater. :

Pedicels in all species of Alangium contain a rather compact cylinder
of vascular tissue, difficult to resolve into individual strands. At the base
of the flower this cylinder expands into a circle of 10 or more major bundles
(usually twice the number of petals). In A. javanicum and A. ridleyi two
additional series of bundles diverge radially from the first within the short
distance of a few 15, microtome sections and still below the base of the
ovarian cavity. The inner series, containing up to 30 fine strands, com-
prises the gynoecial vasculature. The middle series supplies stamens and
petals, and the outer series supplies the calyx (Fics. 2, c, d; 3, a). In A.
havilandii the two outer series and the middle series are united for a dis-
tance; their divergence takes place about halfway up the ovary wall. And
in section ALANGIUM (Fic. 3, b-d), these two series remain united almost
to the insertion of petals and stamens.

172 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

cos
Fiewers of Alansium. a, A. javanicum, P.

hee P. Sam, s.n.; cross section

near apex showing 4 stigmatic rays; & 16. b, NL Hernaez 635,
US; cross section near apex, showing asymmetrical convolutions of stigma;

c, javanicum, P. m, s.n., longitudinal section of ovary, showing

attachment of ovule; arrows indicate 3 series of vascular bundles;
javanicum, P. P. Sam, 5.N., CTOSS iis through inferior ovary showing vascular
undles in 3 series (arrows); X 1
In section Conosticma there are some heavy — and radial
strands near the level of insertion of petals and stamen
connect the bundles of the outer and the intermediate series. ia
vascular feature in Alangium javanicum and A. ridleyi is the occurrence
of extra longitudinal bundles between the principal strands of the outer

ese inter-

1968 | EYDE, ALANGIACEAE 173

series, bundles that terminate blindly at their basal end, not connecting
with any other bundles. At their apical end these join with main bundles
of the outer series and with the bundles of the intermediate series in
supplying branches to the disc.

The dorsal supply in section CoNosticma is peculiar in that a discrete

Fic. 3. Gymnoecia, of Alangium in cross section. 4, A. javanicum med
= asin’ Fic. 70). Bosaen region; all bundles of inner ies except ' ose

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174 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

carpellary midvein is absent from the ovary wall. The position where a
single dorsal bundle would be expected is occupied by a group of three
or more fine, anastomosing strands, and on the other side of the ovary
from this group is a second cluster of fine bundles (Fics. 2, d; 3, a). Al-
though these two groups of bundles can be considered a part of the inner
series, they tend to lie a little to the outside of other members of the series.
Followed distally, the members of each dorsal group merge, forming a pair
of well defined stylar bundles (Fic. 4, a). The two stylar bundles bifurcate
a short distance above the base of the style, and profuse ramification of
the resulting four branches supplies the stigma.

Section ALANcium has a similarly complex dorsal supply, but the com-
ponent strands separate from the outer series at a higher level, passing
transversely across the upper part of the ovary to enter the style (Fic.
4, b). Also, the stylar bundles bifurcate closer to the stigmatic region than
they do in section CONOSTIGMA.

Ovules receive a similar vascular supply in both of these sections. The
entire inner series except the dorsals converges at the placenta (Fic. 3, a),
uniting in an ovular bundle that passes down one side of the ovule and up
the other.

Floral vasculature of section Marea and section RHYTIDANDRA differs
from that in the foregoing sections primarily in the fewer gynoecial
bundles. The inner series in section MARLEA usually contains, in addition
to the two dorsals, only two pairs of gynoecial bundles, one pair at each

4. Dorsal vasculature in section Conosticma (a, ba
ed
ridley) and section ri sipen ts (b, based on A. longi “i ) sed on Alangium

1968 | EYDE, ALANGIACEAE 175

side of the ovarian septum (Fic. 2, b). The paired structure is less evident
in some cross sections than others because the two strands of each pair
branch and anastomose somewhat; cross sections through the upper part
of the ovary are best for showing distinct pairs (Eyde, 1968, Fig. 13).
I refer to these pairs as ventral bundles, since that term has been used for
similarly situated pairs in Cornus (Wilkinson, 1944) and Nyssa (Eyde,
1963); however, as I have stated in earlier papers, the homology of these
bundles to the centrally or marginally located ventrals of other families
is uncertain. Paired bundles supplying the ovule or ovules diverge from
both sets of ventrals and pass transversely through the summit of the ovary.
An unusual feature, never observed in Cornus or Nyssa, is the occurrence
of additional transverse (or oblique) bundles lower in the ovarian septum
(Fics. 5, a; 8, r). These branch at intervals from the ventrals and end
blindly in the septum, perhaps indicating the former location of ovules
eliminated in the course of evolution. The ventrals in some flowers end
abruptly where the ovular strands diverge; in other flowers they end in
the base of the style.

A single outer series of prominent bundles supplies the calyx, corolla,
and androecium in section MartEA, as in section ALANGIUM, but in section
Mar-EA the dorsal bundles (Fic. 5, a, b) are free for most of their length.
Furthermore, the dorsal supply in section Martea usually consists of single
bundles,? unlike the dorsal complexes of sections Conosticma and
ALANGIUM. The two dorsals pass through the style and divide into four
strands a little below the stigma.

Dorsal bundles of section RHYTIDANDRA are also discrete, but they are
united with the outer bundles to a greater degree than in section MARLEA
(also, because there is only one locule, the ventral bundles lie more to
one side of the ovary). One of the dorsals diverges a little below the gi
tion of stamens and petals; the other, always the midvein of the “lost
carpel, is independent for about half the length of the ovary (indicated
diagrammatically in Fic. 9). Both dorsals pass into the stylar column as
discrete bundles, but they divide below the separation of the two styles;
thus the free portions of the styles each receive two bundles.

Petals of Alangium species contain three to seven parallel vascular
strands that merge basally in the region of insertion. The most common
number is five. Where petals are thick (section ConosTIGMA), additional
small strands are often located internal to the others. These are “inverted
bundles, oriented with their xylem toward that of the principal petal
bundles. Stamens of Alangium species commonly contain only one bundle,
but two slender lateral strands flank the principal bundle in 4. ridleyi.

vasculature. He stated that “the so-called dorsal ribs are m
vular strands,” even though he correctly portrayed their passag ich sar oabi
(see his Text-fig. 6, G, H). Horne worked with very young flowers, which may
contributed to the misinterpretation.

e into the style

176 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Fic. 5. a, Alangium chinense, Makiling Natl. Park; cross section through

er
folium, cult., Tohoku; pth section through gynoecium showing 1 empty, 1 fer-

1968] EYDE, ALANGIACEAE 177

The filaments of A. rotundifolium and A. chinense also have some thin
“extra”’ bundles.

The nectariferous disc is thoroughly vascularized in all species of Alan-
gium, receiving transverse strands from the bundles that supply calyx,
corolla, and androecium. Although the fine vascular ramification within
the disc is usually composed of phloem only, as is the case in most dicoty-
ledons (Frei, 1955), I have observed xylem in preparations of A. chinense
and A. rotundifolium; its presence may relate to stage of development.
Floral vascular bundles, especially those of the hypanthium and gynoecium,
tend to be amphicribral in all species.

Secretory structures. Sertorius (1893, p. 561) recorded the occur-
rence of “sehr grosse Sekretzellen” in fruits of Alangium, and Faure
(1924, Fig. XV, 1, 7) figured ‘“poches sécretrices,” erroneously providing
them with a secretory epithelium of small cells. Schnarf’s (1922, Figs.
1, 3) drawings of sectioned flowers portray the secretory structures ac-
curately and indicate their proximity to a vascular bundle. None of these
authors realized, however, that the structures are articulated laticifers.

The laticifers begin as longitudinal files of elongating parenchyma cells,
which enlarge to a diameter several times that of adjacent parenchyma
cells and eventually become vessels by the disintegration or rupture of
transverse walls. They are usually arranged in an arc around the phloem
of floral vascular bundles, and where phloem completely surrounds the
xylem of a bundle, a ring of laticifers may surround the phloem. T he latici-
fers do not actually differentiate within the phloem: there are usually
three or more parenchyma cells between a laticifer and the nearest sieve
tube. In this respect the laticifers of Alangium differ from the very similar
structures of Argemone mexicana (Papaveraceae) figured by Kapoor and
Sharma (1963). Laticifers can accompany vascular bundles in any part
of the Alangium flower with the possible exception of the ovule and the
style, but they terminate at the base of the flower or extend only a short
distance into the pedicel. :

The content of the laticifers stains darkly and usually includes the
pellets (Kiigelchen) that Schnarf (1922) observed. Laticifers ofa matur-
ing ovary often seem to be partially empty when examined in cross section,
and the pickled material does not “bleed” when cut, as it should if the
contents are under pressure (Esau, 1965, p. 323).

One would not expect laticifers to be abundant in flowers of Alangium
and to be absent from other parts of the plant. Having at my disposal
some pickled leaves of A. chinense, I prepared simple freehand sections

tile locule; arrows indicate dorsal and ventral bundles; X 27. C; ee wont at)

B. Hernaez 635, us; cross section of fruit wall eigen) DE sieyawarl, Weer
ici . tteniusa nucifera, , "

and empty laticifers (arrow); X 11. d, Me cecium; arrows

indicate dorsal (short) and ventral (long) bundles; X 18. e, A. chinense, Makil-

ing Natl. Park; cross section o

accompanying laticifers (bottom); X 150.

JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

178

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fruit;

inferior part of flower; laticifers
Bogor, V111.F.25;

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1968 | EYDE, ALANGIACEAE 179

of a petiole and found laticifers in abundance around the major vascular
bundles (Fic. 5, e). It is probable that other anatomists have overlooked
the secretory structures in leaves because they were working with dried
material. Govindarajalu had living material of A. salviifolium, but I have
observed that laticifers are inconspicuous or absent in flowers of this
species; perhaps they are equally difficult to detect in other parts of the

Latex vessels are plentiful in fruits of section ALANGIUM, although they
are hard to find in the flowers. In section MarLEA and in section RuyTI-
DANDRA, however, laticifers are conspicuous at all stages of flower and fruit.
Their arrangement in cross sections creates an attractive “scalloped” effect
(Fic. 5, a), which Clarke (1850) showed in his Fig. 8 and 9, even though
he did not understand the reason for it. Laticifers of sections MARLEA and

wall. Section Conosticma is peculiar in that the floral laticifers seem to
disappear as the fruits develop. My longitudinally sectioned flowers of
A, javanicum show many latex vessels of large diameter (Fic. 6, a), some
of them branching and perhaps anastomosing with neighboring vessels.
Fruits of this species, although they contain densely-staining tanniniferous
cells in great numbers (Fic. 6, b), have no laticifers. Some of the tannin
cells are aligned in such a way as to suggest that they have been derived
from laticifers through a secondary development of cross walls; i.e., through
the reversion of coenocytic laticifers to a cellular condition. I offer this
inference cautiously because my flowers and fruits are from different
sources. Moreover, I do not know that anyone has ever documented a
reappearance of the cellular condition in laticifers.

Endocarp. Fruits of most species contain a thick, woody stone, but
the innermost tissue of fruits belonging to section ALANGIUM is thin,
leathery, and easily opened with an ordinary razor blade. It is difficult
to separate the intact endocarp from outer tissues of such fruits, and when
this has been done (by boiling in 5 percent NaOH) the endocarp collapses
readily (Fic. 7, f). The endocarp of A. longiflorum is so little lignified
that satisfactory sections can be prepared by the paraffin method (Fic.

z

ae
The stones of section Conosticma (Fic. 7, a-e; see also Harms, 1906,
Fig. 16) are more massive than those of other sections; they are further
distinguished by the many fine longitudinal ridges on their surfaces. The
ridges may be very faint or fairly conspicuous, straight or Sanpete a
They undoubtedly arise in association with the many fine vascular bane =
of the gynoecial series. This can be verified by examining the apex of the

tangential section showing vascular bundles and adjacent : a a S
laticifers may be caused by reversion from coenocytic to cellular i on (
text); & 54. c, Mastixia cf. pentandra, Kostermans & — on and Se
section through wall of young fruit showing two secretory canals with ep

cells: < 210.

180 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

stone, where the bundles converge and pass into the endocarp. No
matter how carefully the stone is cleaned, remnants of vascular tissue
adhere to the apical end.

Serial sections ae that the fleshy outer tissue of the fruit wall in section

—C,
VHF .2 25, us; 3 views of 1 stone. d, A.
a e, A. grisolle vides, Rarvite Forestier 20056-

Pie salviifolium, Mrs. D. 7. Colline 1. do21, U8. Au, 2

ConosTiGMA has a broad zone of radially elongate cells immediately pe-
ripheral to the stone. In all of my specimens this zone contains large
irregular cavities, apparently resulting from tearing of the cells as the fruit
wall expands.

Fruits of section Mar.Ea are often two-carpellate, and stones from dif-
ferent species can be arranged in a sequence to show the gradual loss of one
of the carpels. A. kurzii (Fic. 8, a-c) commonly has two nearly equal
locules in its fruit; therefore, a plane through the septum of the endocarp
divides it into roughly identical halves. In A. chinense (Fic. 8, p, q), one
half is smaller than the other, but both locules can be fertile notwith:
standing. Gynoecial asymmetry is more pronounced in A. platanifolium
(Fic. 8, n, 0), A. alpinum (Fic. 8, g-i), and A. rotundifolium (Fic. 8,
m). In stones of these species one of the carpels is reduced to a shield-
like prominence, often with a tightly closed abortive locule. A. griffithii
(Fic. 8, s-u) completes the series; its stones bear no trace of a second
carpel. In A. barbatum (Fic. 8, d-f, j-1) different collections show marked
differences in the degree of reduction of the second carpel.

1968 | EYDE, ALANGIACEAE 181

Alangium kurzii, J. & M.S. Clemens 3740,
d-f, A. barbatum, C. S. Fan & Y. Y. Li 315 :
2-carpellate structure. g-i, A. alpinum, G. Forrest 27290, us; 3 views of 1 stone.
jo, A. barbatum, J: & M.S

A. C. Smith 7409, us; 3 views of 1 stone showing large pits in surface, antec
second carpel. y, A. C. Smith 5249, us; orientation 1s intermediate between that
of w and that of x; note absence of second carpel. All X 2 except r, x 4.

182 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

From its shape and lack of sculpturing the stone of A. griffithii might
at first glance be confused with stones belonging to section CONOSTIGMA.
Stones of the other species of section MARLEA are unmistakable. They are
slightly ridged longitudinally and deeply pitted in the region between the
two carpels. When one of the carpels is much smaller than the other,
the pitting is confined to the side with the rudimentary carpel. Most of
the pits occur where vascular bundles pass into the septum from the ventral
carpellary bundles. Breaking open the fertile locule longitudinally to show
the septum in face view exposes a radial pattern formed by these bundles
(Fic. 8, r).

Stones of section RHYTIDANDRA (Fic. 8, v-y) also have a rudimentary
second carpel in most cases, but they are easily distinguished from stones
of section MarteEa by their larger size, more inflated appearance, and more
profuse pitting. Pits are larger than those of section MARLEA, and they

Conostigma Alangium Marlea Rhytidandra
rd os
" x
Style &
stigma ce
; “aN
stamens | |
| ( ev ¥
floral
vasc.
system

<7

@ = BS, a

ZN

vessel
members

Fic. 9, Some sectional differences in Alangium.

1968 | EYDE, ALANGIACEAE 183

cover almost the entire surface of the stone. Most of them are not associ-
ated with vascular bundles, and I have no ontogenetic explanation. for
their presence.

Histologically, the stones of all Alangium species are very much alike.
Macerations show the component sclereids to be more or less equidimen-
sional and irregularly lobed like the pieces of a jig-saw puzzle. To judge
from the position of the cracks (Fic. 7, a, b; Fic. 8, m, 0) that appear in
repeatedly boiled and dried specimens, all of the stones open in the same
manner at germination.

DISCUSSION, CONCLUSIONS

Evolutionary trends within the Alangiaceae. Ficure 9 sum-
marizes five structural features to which I attach evolutionary importance.
The first four rows of the diagram represent sectional differences * de-
scribed in this article. The bottom row, based on observations of other
workers (Metcalfe & Chalk, 1950; Govindarajalu, 1961), shows two
distinctive features in the secondary xylem of section ConosticMa: the
vessel members have scalariform perforation plates, and they are about
twice as large as the vessel members in other sections.

Pollen provides still another set of characters. The information in
TABLE 1, taken from Erdtman (1952) and Cookson (1957, A. javanicum
only), reveals sectional differences in the number of pores, in the presence
and distinctness of colpi (furrows), and in surface markings.

TABLE 1. Sectional differences in the pollen characters of Alangium.*

CONOSTIGMA ALANGIUM MarRLEA HALT TIOANDEA
(2 —) 3 (—4)-col- 3-5-porate; warts 3(-4)-colporate or 4(—5)-colpoido-
porate or colpoid- on surface colpoidorate; sur- rate; warts on
orate; small- face granular, re- surface
meshed surface ticulate, or striate
pattern

* Adapted from descriptions by Erdtman (1952) and Cookson (1957).

Ficure 10 is an evolutionary diagram that will account for the sec-
tional differences. I visualize members of the ancéstral group as having
pluricarpellate gynoecia, isomerous androecia, separate bundles to all floral
appendages, vessels with scalariform perforations, 3-colporate pollen grains,
and multiple styles. Arrows indicate the direction of structural changes,
and the size of the lobes roughly indicates the relative number of species
in each section. :

I assume a direct derivation of section ALANGIUM from section Cono-
STIGMA in order to account for the strong similarities between the two and

* Cytological studies should be undertaken to determine whether each section has
its own chromosome number. I know of only two counts for Alangiaceae: dag m
A. lamarckii (= A. salviifolium; Gopinath, 1945) and 2 = 22 In EAs begomounm
(one or another species of section Marea; Wanscher, 1933).

184 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

vessel perforation
scalariform | simple

floral vasculature
little fusion | much fusion
——

1 carpel
MARLEA

2 carpels

10. Evolutionary relationships in Alangium. Area of each lobe roughly

ri bundles to sepals, petals, stamens; isomerous androecium; 3-colporate
a bf

for the concentration of primitive characters in section CONOSTIGMA.
Sections RHyTIDANDRA and Mar ea also have many characters in com-
mon; explain this by assuming a close common ancestry. A direct
derivation of either one from the other would be unconvincing, because
section RHYTIDANDRA retains the more primitive stylar characters while
some re of section Martea retain the more primitive two-locular
conditio

This kind of diagram is necessarily speculative, but I have tried to
adhere to the most conservative and widely followed concepts of evolution-
ary morphology. The derivation of simple perforations from scalariform
perforations is universally accepted by wood anatomists, and the gradual
union of floral structures (in this case, styles and vascular acline) is

1968 | EYDE, ALANGIACEAE 185

another trend generally recognized as irreversible (Stebbins, 1967, p. 135).
That the two-carpellate gynoecium preceded the one-carpellate (pseu-
domonomerous) gynoecium is unlikely to be challenged because all degrees
of reduction can still be seen in section MarzEA: it would be difficult to
interpret this evolutionary series in the opposite direction without assign-
ing some selective value to the abortive carpels in intermediate forms. My
view of evolutionary changes in the pollen is based on the fact that grains
of the 3-colporate type occur very early in the fossil record of angiosperms
and are retained in many of the less specialized groups of dicotyledons.
From this I infer that 3-colporate pollen grains of Alangium are more
primitive than pollen with four or five pores and with colpi poorly developed
(the colpoidorate condition) or absent.

Erdtman (1954, p. 79) has hinted that Alangium might be made into
two genera on the basis of pollen characters. From his earlier remarks
(1952, p. 40), I gather that he would separate the species with porate
grains from those with colporate grains, thus requiring the recognition of
section ALANGIUM as one genus and the three remaining sections as another
(a reinstatement of Marlea Roxb.).* However, this is not the only arrange-
ment that could be made if only pollen characters were considered: section
ALaNcIuM, lacking colpi, could be allied with section RHYTIDANDRA, In
which colpi are poorly developed, for these two sections share the warty
surface pattern. However, to adopt either of these arrangements would
be to ignore all of the other characters discussed in this paper. It seems,
therefore, that the same changes — development of a warty exine, increase
in the number of pores — have occurred in separate evolutionary lines
within the Alangiaceae. If it were necessary to divide Alangium into two
genera — and I do not see that anything would be gained by doing so —
a consideration of all characters would require that section CONOSTIGMA
and section ALANGIUM be put in one genus, section MARLEA and section
RHyYTIDANDRA in the other. : :

The association of many stamens with advanced characters in section
ALANcIum will seem unusual to some floral morphologists, for we tend to
look for oligomerization of all whorls, including the androecium (Takhta-
jan, 1959, p. 76; 1964, p. 69). Indeed, the traditional view that evolution
favors a change from polystemony to oligostemony was codified a half
century ago as Bessey’s (1915) 24th dictum. Recently, however, Stebbins
(1967, p. 121-123) listed several taxa for which considerations of compara-
tive morphology indicate the opposite trend. The most compelling evidence
for adding Alangiaceae to this list is in the vascular supply to the stamens.
If flowers with many stamens were primitive, each stamen would have a
corresponding vascular strand in the extracarpellary portion of the ovary
wall, as is the case in section CoNostTIGMA. But section ALANGIUM has
far fewer bundles in the inferior part of the flower than section Cono-
d gone
to the editor, explicitly separates the ca of artes “pains

re ‘ ie nding to ;
dae a, Mrs. Mevientan did not consult Bleembergen s mono-
graph before writing her paper.

186 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

sTIcMA, and the extracarpellary strands are arranged in a single series,
each bundle supplying several stamens. Also, the stamens of all Alangium
species are in a single whorl. This is a clue that Alangium is a special
case, since taxa in which gradual oligomerization of the androecium has
occurred often retain transitional stages in the loss of whorls.

Systematic position of A. grisolleoides. By a happy coincidence,
Monsieur R. Capuron was working at the herbarium of the Muséum Na-
tional d’Histoire Naturelle, Paris, when I visited there in 1964, and he
kindly gave me a few flowers and fruits from his collections of A. grisol-
leoides. This Madagascan rarity, discovered only a few years ago,” is
clearly a member of section ConosticMa, as Capuron (1962) surmised.
Although its floral trichomes do not occur in the stellate-peltate pattern,
it has the Conostigma type of endocarp (Fic. 7, e) and the characteristic
papillose epidermis on anthers and style. It is the only dioecious species
of Alangium.’ 1 did not get good serial sections from the flowers (all
staminate) that Monsieur Capuron gave me, but the preparations were
adequate to show a tiny empty locule in the rudimentary ovary. It is note-
worthy that this species, with its reduced number of floral appendages,
has evolved in the same group that gave rise to the polystemonous section
ALANGIUM.

Place of Alangiaceae in the general system. Some of our most in-
fluential systematic works place Alangiaceae near Cornaceae (Némejc,
1956; Hutchinson, 1959, 1967; Melchior, 1964; Takhtajan, 1966; Sod,
1967). This concept has a long history (Harms, 1897, and Horne, 1914,
review the early opinions), and it has been the prevailing one among those
who have worked with the two families. I began my own investigation of
Alangiaceae with the belief that Alangium is related to Cornus, as sug-
gested by embryological studies (Chopra & Kaur, 1965), and to Nyssa,
as suggested by similarities in the general appearance of the plants’ and
their pollen grains (Erdtman, 1952). I expected to find an even closer
resemblance between Alangium and Mastixia, the only cornaceous genus
with secretory canals. In fact, however, the secretory structures of Mas-
tixia are true resin canals with secretory epithelia (Fic. 6, c), not at all
like the laticifers of Alangium. Furthermore, I have come to realize, con-

* The discovery extended the range for section Conosticma well beyond ae area
mapped by Bloembergen (1939, Fig. 8). The known range for section ALAN

(Bloembergen’s Fig. 2) has also as xtended te the sittin: of A. Selatan to
the flera of tropical East Africa (Verdcourt, 1958).

: = Bloembergen (1939, p. 195) mentions specimens of A. griffithii and A. salviifolium
with sterile anthers; these may indicate a tendency toward diocecism in section MARLEA
and section ALANGIUM.

s of Alangium can be confused with V yssa in the herbarium. Thus, “Alan-
gium” shaweliense W.W. Sm. is a synonym of Nyssa sinensis Oliv. and “Nyssa” hollrungii
s not

so
ES

mble Neale none ssp: or almost any other characters —a

les equently, systems that place Alangiaceae and “a daperae
in Myrtales while sce rsp in Umbelliflorae are manifestly in erro

1968 | EYDE, ALANGIACEAE 187

trary to an earlier assumption (Eyde, 1968), that the gynoecial vascula-
ture of Alangium is not exactly like that of Cornus. The difference is that
Alangium has extra (ovular?) bundles ending blindly in the septum. Were
such bundles present in the ancestors of Cornus, or do they indicate an
entirely different origin for the vasculature of Alangium? Atypical arrange-
ment of gynoecial vascular bundles must be studied further in other plant
families before a confident choice can be made between these two alterna-
tives. It might also be informative to examine the vasculature of a three-
locular Alangium flower like the one Clarke (1850) figured (I have never
seen one). In any case, there now seem to be good reasons for doubting
the close affinity of Alangiaceae to Cornaceae and Nyssaceae, and the
opposing views therefore deserve careful consideration.

Several modern works link Alangiaceae with Rhizophoraceae and Com-
bretaceae in the Myrtales (Roederer, 1930; Pulle, 1952; Novak,
1954; Vinogradov, 1959). Therefore, I sectioned some flowers of Com-
bretaceae (Combretum, Terminalia) as a part of this investigation, but
I found no structural resemblance to Alangium. Flowers of Combretaceae
have gamopetalous corollas, biseriate androecia, and unilocular gynoecia
with two or more ovules — all points of difference from Alangium — an
there are corresponding differences in the floral vascular system. Another
dissimilarity is that the ovules of Combretaceae have two integuments:
there is no trace of a second integument in Alangium.* Still another is that
the characteristic rudimentary stipules of the Myrtales (Weberling, 1960)
are lacking in Alangium. Furthermore, the pollen of Alangium and Com-
bretaceae are “+ different’? (Erdtman, 1952, p. 40) and the xylem char-
acters are distinctly different (Janssonius, 1918, p. 707). Finally, the
known alkaloids of Rhizophoraceae are quite unlike those of Alangiaceae
(Hegnauer, personal communication). In short, all lines of evidence weigh
heavily against the alliance of these three families.

Hallier (1905; 1912, p. 221) once considered Olacaceae the closest
family to Alangiaceae, although he later (1918, p. 117) modified this
opinion. H. K. Airy Shaw, in his revision of Willis’ (1966) Dictionary,
cautiously offers a similar idea. Under Alangiaceae, Shaw says “Relation-
ships obscure; perhaps some connection with Olacaceae and Ehretiaceae.
The suggestion of an affinity with Olacaceae is worthy of further investiga-
tion, because many olacaceous genera have non-articulated laticifers (Met-
calfe & Chalk, 1967). However, the ventral carpellary vascular system
differs from that of Alangium in that the bundles rise through the center
of the flower (Agarwal, 1963 a, b). The floral structure of Ehretia (Johri
& Vasil, 1956) is even less like that of Alangium.

Shaw adds Metteniusa to the Alangiaceae, a change that I do not accept.
Metteniusa is a genus of uncertain systematic position (Icacinaceae and
Olacaceae have been suggested) with epipetalous stamens wee mia

. itzon’ mments on Alangium are especially pertinent.
plas crlasoh Pot oe eds of reviewed by Johri (1963, p. 424) and ogee
& Kaur (1965). Hutchinson (1967, p. 41), in attributing two integuments to Alangium,
repeats an error from his 1959 work, despite Takhtajan’s (1959, p. 239) effort to

correct him

188 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

ovary. A sectioned gynoecium (Fic. 5, d), followed serially from base to
style, displays the vascular system and the ventral suture of a single
carpel, more like a Prunus gynoecium than the complex pluricarpellate or
pseudomonomerous gynoecium of Alangium.

Recent discovery of the alkaloids emetine, cephaeline, and psychotrine
in Alangium salviifolium indicates to Hegnauer (1965, p. 245; 1966, p.
219) that the Alangiaceae may be related to the Rubiaceae, the only other
family in which these substances are known to occur. Alkaloids are not
perfect taxonomic indicators, to be sure, but the suggestion is supported by
Alangium-like characters in certain rubiaceous flowers. There are rubiace-
ous taxa (Oxyanthus natalensis, Gardenia resinifera, Mussaenda sp.) in
which all or part of the ovular supply passes through the septum. Guet-
tarda speciosa, like certain species of Alangium, has a great many gynoecial
bundles around its locules. Furthermore, a vascularized disc is of common
occurrence in the Rubiaceae. (Details of floral vasculature are taken from

ao, Ramarethinam, & Iyer, 1964.) Most Rubiaceae are sympetalous, with
epipetalous stamens, but in Synaptantha free stamens accompany a corolla
that is divided almost to its base. Among the many poorly known genera
of this large family may be others that are even more like Alangium than
those I have mentioned.

In conclusion, therefore, I urge other workers to look into the Rubiaceae
for the closest allies of Alangiaceae. It is conceivable, as Hegnauer (1965)
suggested, that Alangium is related both to the Rubiaceae and to the
Cornales (sensu Takhtajan, 1966), forming a link between the two groups;
however, I do not share that opinion at present.

SUMMARY

Principal observations and conclusions resulting from an anatomical
study of flowers and fruits of Alangium (the only genus of Alangiaceae)
can be summarized as follows:

1. The secretory structures of Alangium are non-articulated laticifers.

2. Marked differences in the number of gynoecial vascular bundles
separate Bloembergen’s section CoNosTIGMA and section ALANGIUM from
section Marea and section RHyYTIDANDRA. In addition, section CoNo-
STIGMA differs from section ALANGIUM and section Mar eA from section
RHYTIDANDRA in the degree to which gynoecial bundles are united with
other bundles of the flower.

3. Characters of the endocarp further substantiate Bloembergen’s
concept of four subgeneric sections. Surface features of the woody stone
can be used to separate three of the sections, and the fourth (section
ALANGIUM) is distinguished by its poorly lignified endocarp.

4. In section ALANGIUM stamens are much more numerous than petals,
but their number is variable and not an exact multiple of the petal number
as earlier treatments would lead one to believe. Polystemony is probably
an advanced condition in Alangium, because it is associated with a reduced
gynoecium, an advanced floral vascular system, and advanced characters
of the pollen and wood.

1968 | EYDE, ALANGIACEAE 189

The prevailing opinion that the Alangiaceae belong to the Cornales
is not well supported by anatomical characters of the flower. However, all
indications weigh heavily against a close affinity to Combretaceae or
Rhizophoraceae, and present evidence is insufficient to prove a relationship
with Olacaceae. Similarity in alkaloid content and in certain details of
floral structure suggest that the closest allies of Alangiaceae may be found
in the Rubiaceae.

ACKNOWLEDGMENTS

For Indonesian specimens of Alangium, I thank Professor C.G.G.J. van
Steenis, Leiden, Dr. A. Kostermans and Dr. W. Soegeng Reksodihardjo,
Bogor; for Philippine specimens, Mr. M. Q. Lagrimas and Mr. M. Mon-
salud, University of the Philippines, and Professor W. L. Stern, University
of Maryland. Professor E. Govindarajalu, Madras, and the late Professor P.
Maheshwari sent the Indian material. Other collections of Alangium came
from Dr. W. Meijer, Sandakan, Malaysia, Mr. L. S. V. Murthy, Kuching,
Sarawak, and Dr. K. Sohma, Sendai, Japan. Dr. David Bierhorst, Cornell
University, and Dr. P. B. Tomlinson, Fairchild Gardens, supplied flowers
of Terminalia and Combretum for comparison with Alangium, and Dr.
John J. Wurdack, Smithsonian Institution, gave me the specimens of
Metteniusa nucifera. Credit is due Mrs. Xavier F. Smith for microtech-
nical assistance, Miss Anne Donovan for the drawings, and Victor Krantz
for the photographs. My colleague Harold Robinson patiently listened to
my ideas on phylogeny as they were forming and suggested several im-
provements in the drawings. The Smithsonian Institution provided fi-
nancial support through Research Award $g0625005

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1968 | EYDE, ALANGIACEAE 191

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DEPARTMENT OF BOTANY

SMITHSONIAN INSTITUTION
WasHINGcTON, D.C. 20560

1968 | PHILIPSON & PHILIPSON, RHODODENDRON 193

DIVERSE NODAL TYPES IN RHODODENDRON

W.R. PHILIPSON AND M. N. PHILIPSON

RHODODENDRON Is ONE Of the largest of the angiosperm genera. In spite
of obvious diversity in habit, flower form, and many other characters,
there is little doubt that the group is a natural one. The only serious res-
ervation to this view is that the three species forming the subgenus THERO-
RHODION might be better regarded as a distinct genus (Sleumer, 1949).
Proposals to subdivide the genus in recent years have been based on ex-
pediency in the face of such large numbers rather than on botanical merit
(Copeland, 1943).

It might be expected, therefore, that studies of the range of form shown
by various features of Rhododendron would reveal interesting examples
of diversity within a coherent group. This is the first in a series of in-
vestigations into the anatomy of the genus. Here the anatomy and develop-
ment of the nodes of foliage leaves and cataphylls is considered.

Previous studies by others have brought to light striking examples of
such diversity. Maximowicz (1870) defined subgenera by the relationships
between floral and vegetative buds. Sinclair (1937) demonstrated two
well-defined types of vegetative bud, and Cowan (1950) published a de-
tailed survey of trichome types within the genus. Kingdon-Ward (1947)
makes some further suggestions, especially concerning characters of the
seed. Hayes et al. (1951) thoroughly investigated the anatomy of the
leaf blade and discussed its relation to the subdivisions of the genus.

In 1914 Sinnott demonstrated that three types of node occurred in
dicotyledons, and that in many families one type only was found to be
present. He also noted several families in which more than one type of
node was present, and further examples have been recorded, e.g. Icacinaceae
(Bailey & Howard, 1941), Epacridaceae (Dormer, 1945; Watson, 1967),
Monimiaceae (Money, Bailey & Swamy, 1950), Escalloniaceae (Swamy,
1954). These findings did not seriously affect the importance of the node
in classification, because different nodal types usually were found to cor-
respond to major subdivisions within a family. ae

The need for comparisons to be made between appendages of similar
age was brought out by Swamy (1949), working with Degeneria, who
found that the complexity of the node might increase as successive leaves
formed in the seedling. Canright (1955) found a similar ontogenetic
sequence in members of the Magnoliaceae. Bailey (1956) also compared
the nodes of seedlings and mature shoots and found that while sequences
from the unilacunar to the tri- or multilacunar condition were frequent
the reverse transition was not observed. This limitation 1s generally of
little significance when the foliage of mature woody plants Is being com-
pared, though the work of Kato (1966, 1967) shows that variation may

194 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

occur between successive nodes of some genera. Work by Miiller (1944),
Philipson (1948), and Post (1958) also demonstrated the need for con-
siderable caution in comparing nodal types in herbaceous plants. For
example, in Helleborus foetidus L., Miller found mature foliage leaves
to possess seven to eleven traces, but this number diminished in higher
leaves until bracts with a single trace were reached. Post found uni-, tri-,
and multilacunar nodes in the genus Frasera, with reduction series up the
appendages of the flowering stem. In the related Swertia perennis L. the
multilacunar nodes diminished in complexity upwards, and also down-
wards in the basal cataphylls.

Records of different nodal types within one genus of woody plants
are few. Pellegrin (1908) records tri- and unilacunar nodes in Genista,
and Saha (1952) tri-, bi-, and unilacunar nodes in Citrus. The Aralia-
ceous genus Pseudopanax, with most species multilacunar, has at least
one trilacunar species (Philipson, 1965) and the South American mem-
bers of Griselinia are trilacunar whereas the New Zealand species are
pentalacunar (Philipson, 1967).

The occurrence of the five types of node here described in the mature
foliage of a genus of woody plants is therefore of interest. Their rela-
tionship with other characters which have been used in the classification
of the genus is discussed in the last section of this paper.

SCOPE AND METHODS

This investigation is concerned principally with the pattern of the
vascular supply to mature foliage leaves. The node and petiole have been
examined in 264 species, as listed in the last column of the APPENDIX.
These were selected from all the subdivisions of the genus recognized
in Sleumer’s “Ein System der Gattung Rhododendron, L.” (1949), as
modified by him (Sleumer, 1958 and 1964), as well as from all the series
and subseries recognized in the Rhododendron Handbook, Part I (Synge,
1963).* Subsidiary observations relate to the development of these pat-
terns during the growth of the primordium and also to the acquisition
of the mature pattern in successive leaves of seedlings. The nodal pat-
terns of other lateral appendages (cotyledons, bud scales, and perulae)
have also been examined

A series of hand sections at right angles to the main axis was made to
determine nodal types. For a few selected examples of each nodal type,
tangential and radial series of longitudinal sections were also cut to
clarify the interrelations of the traces. Petioles were sectioned, always at
the base and mid-point, but often also distally. All these sections were
stained in an aqueous solution of aniline sulphate and mounted in glyc-
erine jelly.

* Throughout the text — proposed by Sleumer has been used, but

re

authors have included it in Rhododendron. Authorities ve Sine names of Rhodo-
dendrons are omitted from the text but appear in the App

1968 | PHILIPSON & PHILIPSON, RHODODENDRON 195

THE NODE OF THE MATURE FOLIAGE LEAF
The er of vascular supply to the foliage leaves fall into five types
as follows
I. Simple Unilacunar Node (Fic. 1)
The simplest type of node found consists of a single trace which de-
parts from the stele of the axis leaving a single gap. The vascular supply

oo e

Fic. SIMPLE Cee Nove (semi-diagrammatic). The single leaf
trace ee the bud trace depart from one gap.

196 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

normally continues across the cortex and into the petiole as a single unit
and remains unbranched as far as the base of the lamina. Rarely, lateral
branches may occur as the petiole approaches the lamina. Such branches
have been observed in Rhododendron camelliaeflorum, R. tsangpoense,
R. maddenii, and R. scabrifolium.

This simple type of node, or slight modifications of it presently to be
described, occurs in all the 136 species examined from the groups listed
in Taste 1. These groups comprise approximately 55 percent of the
species in the whole genus.

Rhododendron albiflorum. Leaf base, showing origin of two accessory
hander outside the abscission layer.

TABLE 1. Distribution of the Simple Unilacunar Node

SUBGENERA EQUIVALENT SERIES
(Sleumer’s system) (Handbook, 1963)
PENTANTHERA Azalea
TSUTSUTSI
AZALEASTRUM Albiflorum

(except Sect.

CHONIASTRUM ) Semibarbatum
RHODORASTRUM All the
PsEUDORHODORASTRUM “lepidote”’
RHODODENDRON Series

Preciicknk

1968 | PHILIPSON & PHILIPSON, RHODODENDRON 197

Modifications of this simple type occur in a small proportion of the
species. The variants fall into a number of categories, which are sum-
marized below.

a) The leaf trace branches in the proximal half of the petiole, or near the
abscission layer (Fic. 2). Examples are listed in TABLE
e leaf trace divides into separate strands while still in the cortex of the
axis. These subdivisions reunite after passing the abscission layer. This frag-
mentation of the leaf trace into distinct strands which form an arc is very clearly
developed in R. nuttallii and occurs in some other Jarge-leaved members of
MappentA and VireyA, but not in all. In very large leaves of R. nuttallii one
or two bundles may remain as distinct strands in the petiole for some distance,
even as far as the mid-point, but they eventually merge into the central bundle.
The leaf trace is fragmented into distinct strands in leaves examined of the
following species of VirEYA: R. intranervatum, R. javanicum, R. brookeanum,
R. lowti, R. crassifolium, and R. retivenium. However, some species of this sec-
tion with very large leaves were found to have very little, if any, fragmentation.
Examples are R. superbum and R. leucogigas.

) The accessory branches in the petiole originate from the stele of the axis
independently of the central leaf trace. This condition was observed in one leaf
of R. schlippenbachii and applied to the accessory on one side only. The other
accessory arose, as in other leaves of this a as a branch of the central
trace. This particular node, therefore, was bilacun

d) ore adaxial strands of vascular tissue whine the petiole bundle
(Fic. 3). This was observed in species of the subsection MADDENIA poten
to subseries Ciliicalyx (R. johnstoneanum) and Megacalyx (R. lindleyi, R
megacalyx and R. nuttallii).

1G. 3. Rhododendron megacalyx. Transverse section slg adaxial bundles
in Pes ab, adaxial bundles; f, fibers; p, phloem; x , xylem

198 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

TABLE 2. Simple nodes with accessory bundles

GROUP SPECIES REMARKS
BooTHIA sulfureum branches at base, or mid-
petiole, or unbranched
PHAEOVIREYA superbum branches at base
PSEUDOVIREYA invasiorum branches near base
SOLENOVIREYA jasminiflorum branches near base
EUVIREYA
BUXIFOLIA commonae branches at base
JAVANICA culminicolum branches at base
leucogigas branches at base
CANDIDASTRUM albiflorum branches at base
VISCIDULA nipponicum branches at base
PENTANTHERA luteum branches at base
occidentale branches near base
BRACHYCALYX schlippenbachit branches at base
TSUTSUTSI yedoense branches near base
kaempferi branches at base

unbranched

A review of these modifications indicates that the occurrence of branch

blance is superficial because the lateral bundles in the petiole always
reunite with the central bundle, whereas in the complex type they remain
distinct and enter the lamina

II. Intermediate Node (Fic. 4)

In general there is a clear distinction between the simple node just
described and the more complex type found in Rhododendron subgenus
Hymenanthes. However, the nodes of all the species examined of the
section CHONIASTRUM (series Stamineum) exhibited features of both
these nodal types. Because of the difficulty in placing the CHoNIASTRUM
node, it was felt that its aberrant nature was best emphasized by refer-
ring it to a separate type, here designated the Intermediate Node. The

1968] PHILIPSON & PHILIPSON, RHODODENDRON 199

xrw |

Ic. 4. INTERMEDIATE Nope (semi-diagrammatic). While in the cortex the
leaf traces are closely aggregated into a central cylinder. In the petiole these
ee unite to form a central strand to each side of which is an accessory

undle.

recognition of the peculiarity of this nodal type appears to be justified,
as the section CuonrastruM has been considered aberrant within the
genus on several grounds (see, for example, Kingdon-Ward (1947) on
fruit and seed characters). :
In nodes of this type, the leaf trace, which arises from a single gap in
the stele, is composed of a number of bundles (typically five) at the
level of the abscission layer. These soon unite to form a central arc-
shaped strand within the petiole. Immediately outside the abscission
layer accessory bundles arise from this central strand and follow a
course parallel to the abscission layer before turning along the petiole.
The above description would apply almost as well to the Complex
Unilacunar Node to be described below. However, it is considered that
the CHonrastrum node (invariably present in all species of this section

200 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

examined — see APPENDIX) would be wrongly placed in the complex
type. This opinion is based on the fact that the separate portions of the
central trace do not diverge, as in the typical complex type, but remain
throughout their course as a compact arc. And again, this node differs
from the simple type in the presence of accessories together with the
division of the central trace. It is of interest that this small, very ab-
errant and natural group displays a vascular pattern which does not
conform to any of the major nodal types.

The term Intermediate is used in a descriptive sense, without phylo-
genetic implication.

KRW !

Fic THREE-TRACE ion Nope (semi-diagrammatic). Three strong
traces ‘diverge from a single gi

1968 | PHILIPSON & PHILIPSON, RHODODENDRON 201

III. Three-trace Unilacunar Node (Fic. 5)

The foliage leaves of Rhododendron camtschaticum are borne on lateral
shoots which arise from buds situated below the terminal inflorescence
buds. These terminal buds open to reveal shoots which bear a small num-
ber of flowers. Each flowering branch is subtended by a large leafy bract
somewhat different in shape from the foliage leaves. Both the bracts and
the foliage leaves are characterized by a type of leaf trace distinct from
that of all other species examined. The nodal structure varies in detail,
but the most distinctive form is found in the leafy bracts where three
strands diverge from a single gap in the stele. These traverse the cortex
and enter the petiole separately. The mid-trace is larger than the two
laterals, but these are noticeably stronger than the accessory bundles found
in the petioles of many Rhododendrons. In foliage leaves a single arc di-
verges from the stele before dividing into three separate strands (Fic. 5).
The point at which the laterals diverge varies considerably, and frequently
the two laterals at one node separate from the mid-trace at different
distances from the gap. Indeed, in some leaves one or both laterals may
not separate until the trace has passed beyond the abscission layer.

This type of node, with three strong traces arising from a single gap,
was observed only in the very distinctive species R. camtschaticum. T wo
other closely related species are placed in THERORHODION but no material
of these was available for study.

IV. Complex Unilacunar Node (Fic. 6)

In contrast to the simple node, this is a type with a highly complex
structure. A number of strands (often five) leave the edge of a single
leaf gap. If an axillary bud is present, the upper strands characteristically
diverge from the bud trace and may branch during their passage through
the cortex. As the strands approach the abscission layer they splay out,
often widely, before entering the petiole. Within the petiole the strands
converge, uniting to form a large central arc-shaped or cylindrical
bundle. Immediately outside the abscission layer accessory bundles
arise from this central bundle, as in the Intermediate Node, and follow
a course parallel to the abscission layer before turning along the petiole.
Alternatively, in species with a very widely splayed central complex, the
accessories arise from its lateral extremities without further divergence.

The relationship between the origin of the lateral strands of the central
complex and that of the bud trace varies. In many species the laterals
diverge from the edges of the bud trace (Fic. 7, a); in others they arise
from the stele of the axis, and enclose the bud trace (Fic. 7, 0). These
relationships vary according to the development of the axillary bud and
probably also according to the amount of secondary growth which has
taken place. Developmental studies will be required to evaluate this
feature, but the character of enclosed bud traces was found to be prevalent
in certain subsections, notably LAcrEa and THOMSONIA.

Without exception, this Complex Unilacunar Node has been observed

202 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

i
I
i
!
t
'
\
\
I
!
\
4
|
i
1
|
ef

G. 6. Complex UNILAcuNAR Nope (semi-diagrammatic). Several leaf traces

ir to form the central bundle of the petiole, to each side of which is an ac-
cessory bundle.

in all subdivisions of the subgenus HyMENANTHES. Indeed, all three
previously described nodal types are unknown in this large subgenus.
Some variants of the Complex Unilacunar Node are described below, as
is the basically similar Complex Trilacunar Node, which is the only
other nodal type known in HYMENANTHES.

The species in which trilacunar nodes occur will be considered in the
next section. Here some interesting variations of the Complex Unilacunar
Node will be described. They are confined to the following subsections.

TALIENSIA. In R. pronum (subseries Roxieanum) no accessory bundles were
present in the petiole. However, the widely splayed and fragmented central
strand is very similar to that of R. gymnocarpum which is considered to be
closely related.

NERIIFLO R. forrestii no accessories were observed, though these oc-
curred in the varieties repens and tumescens, In the lowest foliage leaf of a

1968 | PHILIPSON & PHILIPSON, RHODODENDRON 203

Fic. 7. Relation of lateral leaf traces to bud trace. A, Rhododendron arbore-
um, laterals oe to outer edge of bud trace; B, R. fulvum, laterals inserted at
inner edge of bud trace pe: -diagrammatic) a, Ruin layer: b, bud trace;
I, lateral face of leaf.

branch of R. sperabile only one accessory was present, and in R. neriiflorum one
accessory became weak towards the base of the petiole and made no junction
with the central eet In R. haematodes one accessory ephsiad gradually
from the central complex instead of making the usual abrupt depart

CaMPANuLata. In R. tsariense no accessories py present. Ina ‘eat with no
axillary bud the central strand was inserted in the gap without obvious frag-

most nodes were of the ee eto type (see below) but one leaf of this plant
varied in having only one accessory bundle which did not appear until ——,
along the petiole. And again, a leaf of R. succothii with only one accessory
bundle was found to have a bilacunar node.

HOMSONIA, SELENSIA, CAMPYLOCARPA, SOULIEA. All these rants age
exhibited some variants. The accessory bundles either arose de novo outside t
abscission layer or diverged from the central complex in the ase see in

. campylocarpum, R. telopeum, R. selense and R. wardii. In R. dasycla an
and R. wardii i some accessories bias gradually pa alge eh
In s €

hese
complex cust het encloses the bud traces in this series, in which case t

204 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

bundles appear to be inserted by separate gaps in the stele. This feature be-
comes exaggerated as further secondary growth occurs. In two species of this
series, namely R. hookeri and R. stewartianum (both subsection THOMSONIA)
the nodes were undoubtedly trilacunar, at least in the sense employed in this
paper (see next section).

The variation within the species R. williamsianum is so great that it warrants
separate description (Fic. 8). In some leaves the node was of the typical Complex
Unilacunar type; in others no accessory bundles were present in the petiole; in
others a single accessory occurred and was either confined to the distal portion

Fic. 8. Rhododendron williamsianum, diagrammatic representation of four

types of node. A, petiole without accessories; B, petiole with one accessory
which fuses with lateral bundle; C, petiole with one accessory which enters stele
roe eh oe D, petiole with two accessories, each fusing with its lateral
undle.

1968 | PHILIPSON & PHILIPSON, RHODODENDRON 205

only or arose from the central strand at the base of the petiole; while in others
the nodes were bilacunar, the accessory arising from a separate gap in the stele.

BarBaTA. In one petiole of R. bainbridgeanum (subseries Crinigerum) only
one accessory bundle was present and this diverged gradually from the central
complex. Other nodes of the same shoot were normal.

To summarize, the Complex Unilacunar Node is found in all species
of the subgenus HYMENANTHEs except those with the even more elaborate
Complex Trilacunar Node, still to be considered. Considerable variation
is found in a few subsections of this subgenus. When accessories are
not present this Complex Unilacunar Node can be defined by the com-
plexity of the bundles at the abscission layer. Very rarely this feature
may be lacking in leaves without buds, but no species of this subgenus
has been found where this complexity never occurs. However, since
fragmentation of the central trace occurs in R. nuttallii and some species
of VirEYA (see above) no absolute distinction can be maintained between
the nodal type of the lepidote subgenera as compared with those of the
subgenus HyMENANTHES. Nevertheless, these two groups, each with over
400 species can be separated by their nodal structure, almost without
exception. The intermediate nature of the node in the section CHONI-
ASTRUM (series Stamineum) has been discussed previously. In the sub-
genus HyMENANTHES only three species were found to lack accessory
bundles and in them the central strand becomes divided in at least some
nodes. When it is recalled that the node of the subgenera TsuTsvutst,
PENTANTHERA (azaleas) and AZALEASTRUM can also be distinguished
from that of the subgenus HyMENANTHES without exception, the taxo-
nomic importance of the node in this genus becomes evident.

V. The Complex Trilacunar Node (Fic. 9)

In several different taxonomic groups within the subgenus HyMEN-
ANTHES the accessory bundles in the petiole do not diverge from the
central system as in the Complex Unilacunar Node just described. Instead,
they arise from strong bundles which leave the stele by independent gaps
and link up with the central system just outside the abscission layer.
The gaps left in the stele by the origin of these bundles are defined by
primary as well as by secondary xylem. The term trilacunar, therefore,
is applied to these nodes. However, since all the strands of a leaf trace
are included within the same mesh of the original primary pattern of
protoxylem, it is perhaps arguable that the pattern Is still basically
unilacunar. Nevertheless, for comparative purposes this is a most dis-

tions: BarBata (R. argipeplum, R. barbatum, R. erosum, R. sisahedryiei
and R. smithii); FaLconera (R. arizelum, R. basilicum, R. falconeri es
R. hodgsonii); Fortunea (R. grifithianum); LactEAa (R. wightii), an

206 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Tuomsonta (R. hookeri and R. stewartianum). The nodes of other mem-
bers of these five subsections examined (see APPENDIX) were of the com-
plex unilacunar type. In addition, the Complex Trilacunar Node was
observed exceptionally in R. campanulatum.

n interesting consistent feature distinguishes the trilacunar node of
the subsection Barpata. The strong flanking bundles run horizontally

>

ne

CoMPLEX TRILACUNAR NODE (semi-diagrammatic). Bundles leave the
stele af independent gaps and join rat Bice complex in the base of the petiole.
From them the accessory bundles

across the cortex, so that their gaps occur higher in the stele than the
gap of the midrib. In the remaining groups with trilacunar nodes the
flanking bundles arise from gaps some distance below that of the midrib
and ascend sharply while crossing the cortex to the abscission layer. The
only exception found occurred in R. basilicum (FALCONERA) where the
node resembles the BARBATA type.

Such complex nodes are subject to considerable variation of detail. In

1968 | PHILIPSON & PHILIPSON, RHODODENDRON 207

particularly large specimens of R. griffithianum (subsect. ForTUNEA) and

. magnificum (subsect. GRANpiA), the central complex of bundles is
convoluted so that the appearance of a double midrib is produced, and,
in the latter, ventral branches of the laterals join the additional ventral
fold of the midrib. On the other hand a plant of the Trinwald variety of
R. grifithianum grown at the Dunedin Botanic Garden had asymmetrical
nodes which were bilacunar, and the same condition was found in R. mono-
sematum (subsect. MACULIFERA). Shoots of R. grande and R. griffithianum
had successive nodes which were unilacunar (complex), bilacunar and
trilacunar.

In summary, the trilacunar node occurs in some species of five subsec-
tions (BARBATA, FALCONERA, FoRTUNEA, LACTEA, THOMSONIA); 0Cc-
casionally in one species of another (CAMPANULATA); and in all the
species examined (though not all the nodes) of the subsect. GRANDIA.

VI. Nodes of Subgeneric Hybrids

Hybrids between species of the subgenus HyMENANTHES and those of
other subgenera are rare (Martin, 1963). In view of the distinctive
features of the node of HyMENANTHES, the nodal patterns of two such
hybrids, from species belonging to two subgenera, were investigated.
Specimens of Azaleodendron (hybrids between species of HyMENANTHES
and PENTANTHERA) growing at the University of Canterbury and in the
Botanic Garden, Dunedin, were found to have simple nodes similar to
those of the subgenus PENTANTHERA. On the other hand, the hybrid
‘Grierdal’ (R. (subgen. Hymenanthes) griersonianum X R. (subgen.
Rhododendron) dalhousiae), from the Savill Gardens, Windsor Great Park,
had nodes similar to those of the subgenus HYMENANTHES, that is, of the
Complex Unilacunar type.

Two hybrids between the subgenera RHODODENDRON and RHopor-
ASTRUM were provided by the Arnold Arboretum. These were the hybrid
‘P.J.M.’ (R. dauricum var. sempervirens X R. carolinianum) and an-
other unnamed hybrid (R. mucronulatum X R. carolinianum). Both were
found to have nodes of the simple type. This would be expected as both
species belong to subgenera with this type of node.

VARIABILITY IN NODAL PATTERNS

I. Within-plant variation

a. FoLIAGE LEAVES. Since the principal object of this investigation was
to compare the nodes of typical foliage leaves, no systematic study of the
variation over the full range of leaf size was undertaken. However, our
observations were extensive enough to enable us to conclude that little
variation does in fact occur. That is to say, the nodes of plants with
characteristically simple nodes are all simple, as would be expected, though
some variation in minor features was noted. Similarly, species with
complex nodes in general have all their nodes complex, though the greater

208 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

scope for variation possibly leads to more frequent modification. A few
specific instances of within-plant variation follow, though most have
already been mentioned when describing the various nodal types.

R. schlippenbachii. Accessory bundles normally leave the central strand above
the abscission layer, but at one node one accessory left the stele by its own gap
(bilacunar).

R. kaempferi. Accessory bundles are absent from normal pS but in leaves
of strong sucker shoots accessories arose at the base of the peti

grande and R. griffithianum. Successive nodes on the same shoot were
uni-, bi. and trilacunar

R. sperabile and R. succothii. Very small accessory bundles are normally
present at each side of the petiolar strand. In one leaf only one of these occurred.

R. campanulatum. In one plant, accessory bundles normally branched from
the central strand at the base of the petiole, but in one leaf they did not occur
until mid-petiole.

R. sulfureum. In one plant, accessory bundles normally arose at the base of
the petiole, though they did not all persist as far as the blade. In one leaf these
basal accessories were joined by a pair of bundles which branched from the
central strand at mid-petiole.

R. williamsianum. In leaves of the same plant accessory bundles may be
absent or one may be present. The accessory, if present, may leave the central
complex at the base of the petiole or leave the stele by an independent gap.

b. OTHER LATERAL APPENDAGES.

Cotyledons. The cotyledonary node was not investigated in detail.
However, the cotyledons of several species of diverse affinity have been
examined and all were found to receive a single bundle.

Bud scales. The nodes of the scales which enclose the buds of Rhodo-
dendrons are characteristically unilacunar. An exception occurred in a
series cut through the scales of a plant of R. magnificum which had a very
complex pattern of bundles in the nodes of its foliage leaves. The bud
scales of this plant had several veins which united with the midrib at the
base of the scale: only one bundle entered the axis. In one inner scale,
however, a lateral bundle entered the axis independently so that the
node of this scale was bilacunar.

Perulae. A feature of many Rhododendron leafy shoots is that a termi-
nal group of foliage leaves is separated from the scars of the bud scales
by several internodes which bear no foliage leaves. The appendages borne
at these nodes are strap-shaped scales, or perulae, which are usually early
deciduous, leaving a bare axis below the foliage leaves. In some species,
however, these perulae are persistent. The nodes of the perulae were
investigated in many species of diverse affinity, including those with the
most complex foliage traces, and in all they were found to consist of a
single trace. This branches immediately it enters the perula into several
parallel veins. A curious feature of these perula-traces is that they are
frequently decurrent in the cortex, arising from the stele by gaps as much
as 2.5 cm. below the appendage (Fic. 10). The internodes separating
the perulae grade from short at the base (i.e. next to the bud-scale scars)

209

PHILIPSON & PHILIPSON, RHODODENDRON

1968 |

Node of perula. A, B, C, successive
ed on D,

agnificum.

sections at levels indic

WW)

t levels indicat
of perula. Secondary xylem hatched, trace In soll

Ss
A

1 7
hd

Fic. 10. Rhododendron m

lower transverse
of perula; pf, trace

210 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

to a maximum about three-quarters of the way up the shoot. Above this
they decrease in length. The length of the decurrent trace reaches a
maximum in the longest internodes, but no simple relationship exists
between internode length and degree of decurrence.

II. Within-species variation

Several instances of differences between individuals of the same species
were noted. In view of the within-plant variation just described, and
since no extensive surveys were undertaken, little can be concluded about
the range of variation which may exist among members of the same species.
However, the number of species examined, and the great consistency of the
findings, with mainly trivial within-plant variation, make it safe to conclude
that most species exhibit little variation. Some notable exceptions were
encountered, three species standing out as most variable in their nodal
structure. The variations noted in different plants of Rhododendron wil-
liamsianum have already been described and figured (Fic. 8). It is
surprising that a species so distinct and so constant in most morphological
features as R. williamsianum should exhibit such wide variation in nodal
structure. The variation may be found between nodes of the same plant
and between those of different individuals. Almost as great a range of
variation was observed in R. campanulatum and R. campylocarpum.

DEVELOPMENT OF NODAL PATTERNS

I. Development in Seedling Leaves

An examination of a series of seedlings of the subsect. FALCONERA (in
which the mature leaf has a trilacunar node) established that the earliest
leaves, ranging in length from 0.7-1.5 cm. were unilacunar. The trilacunar
state was reached in the node of a leaf only 2.6 & 1.2 cm. and still without
the mature indumentum. However, some longer leaves, between 5 and 7
cm. long and with the mature indumentum were only bilacunar.

One seedling from this same suite of specimens exhibited some inter-
esting nodes intermediate between the simple and the trilacunar state.
The leaves did not form a graded series up the stem because their size
varied considerably and irregularly. Four leaves in ascending sequence
were as follows:

(i) blade 1.8 % 1.25 cm., node bilacunar with one very faint accessory
bundle in the petiole;

(ii) blade 0.9 < 0.75 cm., node simple, with one faint accessory bundle
in the petiole;

(iii) blade 0.8 X 0.55 cm., node simple, with two accessory bundles in
the petiole;

(iv) blade 2.0 % 1.3 cm., node bilacunar with one accessory bundle in
the petiole.

In all these nodes the central (or only) trace was undivided. It is of
interest to note that the bilacunar and trilacunar conditions arise so

1968 | PHILIPSON & PHILIPSON, RHODODENDRON 211

early in the development of the seedling. In contrast to these specimens,
seedlings of R. sinogrande of comparable size had nodes which were of
the Simple Unilacunar type. Leaves with blades 2.0 x 1.0 cm. showed no
sign of accessory bundles in the petiole nor of extra bundles in the cor-
tex. Their only departure from the simple condition was the separation
from the central strand of one or two subsidiary traces, which arose in
the cortex, passed through the abscission layer, and then reunited with
the central strand to form the single bundle of the petiole. In smaller
leaves the central bundle remained entire, although the individual vessels
could be seen to diverge at the abscission layer. This dilation of the
trace as it passes from the stem into the petiole is never observed in typical
simple nodes, and can be regarded as the first sign of the development of
the complex node of the subgenus HYMENANTHES.

II. Development in the Leaf Primordium

Sections were taken through resting vegetative buds in autumn. The
Species investigated (Rhododendron magnificum and R. griersonianum)
have mature nodes with Complex Trilacunar and Complex Unilacunar
nodes respectively. No time-sequence study of primordia was undertaken,
so that the detailed development of the complex node can not be de-
scribed. The foliage-leaf primordia of both species examined were at an
early stage of development with lamina formation only beginning. In
each the primordium contained a single procambial strand, representing
the midrib, and this was traced downwards until it entered the axis. No
rudiment of any other part of the nodal complex was present, thus show-
ing, as might be expected, that the supply of each leaf passes through
a simple stage, no matter how complex its final condition may be.

DISCUSSION

The presence within one genus of such diverse nodal structure is, we
believe, unique among woody plants. Following the pioneer work by
Sinnott (1914), the node was long considered a conservative character,
often stable in all members of large taxonomic groups within the dicoty-
ledons. Latterly, this view has been seen to require modification, since
considerable variation in nodal structure has been reported within fam-
ilies and even genera (Saha, 1952; Post, 1958). However, until the
present time, the unilacunar condition has been regarded as a constant
feature of the Ericaceae, although variation has been recorded in the
closely related Epacridaceae (Dormer, 1945; Watson, 1967).

The variation in nodal structure observed in the genus Rhododendron
has been presented here as falling into five types. One of these occurs
in a group of three species which have been regarded as forming a dis-
tinct genus, Therorhodion (e.g. by Sleumer, 1949). We consider that

212 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

genus Rhododendron, ranging from simple to complex, form a natural
and coherent sequence into which the three strong, divergent traces of
the camtschaticum type have no place.

The remaining four groups fall into two major categories. The Complex
Unilacunar and the Complex Trilacunar nodes are clearly very similar
morphologically and taken together are found in all the species of the
subgenus HyMENANTHES and in no other group. The morphological
similarity evident from a comparison of Fics. 6 and 9 is strengthened by
two other considerations. Firstly, these two nodal types may occur to-
gether in different species of the same taxonomic group, in different
plants of the same species, and even in different nodes of the same shoot.
Secondly, a few species with otherwise typical Complex Unilacunar nodes
have the lateral parts of the leaf trace inserted so widely that they ap-
proach the trilacunar state, or one bundle may clearly enter by its own
gap, giving a bilacunar node.

The Simple Unilacunar Node and the Intermediate Node also appear
to be linked by similar features. These two nodal types are found in all
species of the seven subgenera listed in TasLe I but including CHONT-
ASTRUM. They stand, therefore, in sharp contrast to the eighth subgenus
HyMENANTHES, in which the node is complex. Apart from the section
CHONIASTRUM which will be considered presently, the distinction between
simple and complex nodes presents no real problems. It is true that some
leaf traces of the subsection MaAppENIA divide in the cortex as they pass
into the petiole but accessories are not developed in this group. Similarly,
in some species of section VirEYA the leaf trace divides in the cortex,
and in addition in this section accessory bundles may occur in the petiole.
On the other hand, in the few members of HyMENANTHES which lack
accessory bundles, the leaf trace is divided in the cortex in the manner
typical of that subgenus. These aberrant forms are closely related to
species with nodes of the typical Complex Unilacunar form. That is to
say, most species of the subsection TaLrenst1A, for instance, are complex,
though one of its species lacks accessory bundles. It is concluded, there-
fore, that these instances of approach between the two types are secondary
modifications resulting from convergent evolution.

The Intermediate Node, found in the section CHon1astRuM, is possibly
nearer to the simple than to the complex node since the central strand,
though divided, does not lose the compactness of form characteristic of
the simple type. Therefore, it is probably correct to regard it as a modi-
fication of that form, though sufficiently distinctive to be considered a
separate nodal type.

The other three sections of AZALEASTRUM, namely CANDIDASTRUM,
MUMEAZALEA and EUAZALEASTRUM were found to share certain features
which contrast strongly with those of CHoNIAstRUM. They all bear
minute, often strongly curved, simple hairs on the petiole; the petiolar
strand is fan shaped in section, not incurved; and some species (e.g.
R. semibarbatum and R. ovatum) have serrated leaf margins. It is sug-
gested that the subgenus AZALEASTRUM might well be subdivided in

1968 | PHILIPSON & PHILIPSON, RHODODENDRON 213

order to accentuate the evident close relationship of these three sections
and their equally evident lack of relationship to CHoN1asTRUM.

It is probable that a close study of the histology of the nodes and petioles
of the azaleas (subgenera PENTANTHERA and TsutsutsI) would reveal
characters distinguishing them from the nodes of scale-bearing rhodo-
dendrons. While it was noticed that branches of the central trace were
not infrequent in azaleas but very rare in lepidote species, no diagnostic
feature was recorded.

It will be clear from the foregoing discussion that the node offers a
character of taxonomic importance. The coincidence between nodal types
and the major subdivisions recognized within the genus is striking. That
a completely new set of characters indicates groups that accord with
existing classification is encouraging, and the validity of the groups is
strengthened by this new evidence. The classification of the genus is at
present in a transitional state. On the one hand there is an old-established

?

defined. This system has the grave practical disadvantage that most of
the species, including the majority of interest to horticulturists, fall with-
in two of the major groups. It was for this reason, no doubt, that an
empirical, informal classification into series was adopted in The Species of
Rhododendron (Stevenson, 1930). These series form the most compre-
hensive and discriminating grouping of the temperate species yet devised.
However, the already established major subdivisions of the genus were
not employed, the series being presented in alphabetical sequence. A
much clearer appreciation of the genus results if the systems are united,
giving the advantages of both. This was done by Sleumer in 1949.
While we wish to emphasize this close correspondence between nodal
anatomy and classification, we would also draw attention to the interesting
variation in detail to be found between members of taxonomic groups,
both large and small, between the nodes of different categories of ap-
pendages, and between nodes of the same order within one individual.
Clearly, the adult anatomy of a node may vary, particularly in some
species and some subsections. Nevertheless, the range of variation is
limited so that the definition of the major types remains clear, with very
few exceptions. The interest of these exceptions must not be allowed to
detract from the taxonomic significance of the nodal types which is based

semble subgenus HyMENANTHES in the aestivation of their foliage leaves
(Sinclair, 1937). By employing these three anatomical features in com-

214 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

bination, three major subdivisions can be defined (TABLE 3). However,
the definition of the resulting groups, whether by anatomy or by external
features, such as the position of flower and foliage buds, is complex. For
this reason, and also in view of the long horticultural history of the genus,
there appears to be little advantage in maintaining separate genera.

TABLE 3
SUBGENUS ANATOMICAL FEATURES
PENTANTHERA node simple
TSUTSUTSI scales absent
AZALEASTRUM aestivation revolute
HYMENANTHES node complex

scales absent
aestivation revolute

RHODODENDRON node simple
PSEUDAZALEA scales presen
RHODORASTRUM

aestivation convolute (except in subsect.
PSEUDORHODORASTRUM EDGEWORTHIA)

The prevalence of the unilacunar node in so many genera of the Eri-
caceae suggests that this state is basic for the family and that the com-
paratively few species with trilacunar nodes in Rhododendron are derived
from the simpler type. Indeed, the unilacunar node has come to be regard-
ed as primitive for the dicotyledons as a whole, and there are convincing
instances of the derivation within a family of trilacunar nodes from the
unilacunar condition (Bailey, 1956). This view is supported in Rhodo-
dendron by the occurrence of two forms of trilacunar node, that occur-
ring in the subsection Barpata and that in other groups. Kingdon-Ward
(1947) adopts the view that the lepidote rhododendrons are primitive
and this would imply that the simple node is the original type. On the
other hand, Hutchinson (1946) postulates that the subsection FALCONERA
resembles the ancestral form of the genus. This group is predominantly
trilacunar. Hutchinson’s opinion is based on his belief in a resemblance
between the subsection FaLconera and the Dilleniaceae, a family which
he considered to be ancestral to the Ericaceae. The node in the principal
subfamily of the Dilleniaceae is tri- or pentalacunar (Sinnott, 1914).
In Actinidia and Saurauia, representing the other two subfamilies, the
node is unilacunar. Since the majority of dicotyledonous families are
characteristically trilacunar, the presence of this nodal type in two groups
does not necessarily imply relationship. However, if the Ericaceae are,
in fact, derived from the Dillenioideae, the trilacunar node is likely to be
the original state in the family. In that event, the genus Rhododendron
would be primitive within the Ericaceae, as only in this genus are tri-
lacunar nodes known to occur. It must be emphasized that the opinions
of Kingdon-Ward and Hutchinson are both speculative. While it is
necessary to examine them in the light of the evidence relating to nodal
anatomy, no final conclusion can be drawn.

1968 | PHILIPSON & PHILIPSON, RHODODENDRON 215

Money, Bailey and Swamy (1950) have suggested that single arc-
shaped leaf traces may in fact sometimes represent several concrescent
strands. If the more complex type of node were primitive in Rhododen-
dron, the very prevalent simple leaf trace which is characteristic of other
genera in the family, would be of such a concrescent type.

In this investigation we have observed the node of a little more than
one quarter of the species of the genus. A larger sample would undoubted-
ly have revealed greater variation in some groups. The types described
are based on features of vascular patterns. Future studies will include
histological characters, and preliminary observations suggest that these
additional characters will allow differences between groups with simple
traces to be recognized and also that distinct types of node and petiole
will be found within the subgenus HyMENANTHES.

SUMMARY

1. The anatomy of the nodes of the mature foliage leaf has been in-
vestigated in 264 species of the genus Rhododendron. These are presented
as falling into five types:

(i) Simple Unilacunar: the leaf receives a single trace; rarely, accessory
bundles may be present in the petiole.

(ii) Intermediate: the trace, while in the cortex, consists of a few bundles
closely aggregated into a central cylinder. In the petiole these bundles unite to
form a central strand which is accompanied by two or more accessory bundles.

(iii) Three-trace Unilacunar: a single arc diverges from the stele before
dividing into three separate strands. This division may be in the cortex or Just
outside the abscission layer.

(iv) Complex Unilacunar: several traces arise from a single gap and enter the
petiole, where they unite to form a central strand which is accompanied by two
or more accessory bundles. :

(v) Complex Trilacunar: the pattern of traces is similar to type iv, but ad-
ditional traces leave the stele by separate gaps.

2. Variations are recorded within the framework of these types as they
occur in taxa and individuals.

3. The essential features of the Complex Trilacunar Node were found
to be established very early in the development of seedlings.

4. All types, including the most complex, pass through a stage of
development when only one leaf trace is present. :

5. The nodes of cotyledons, cataphylls and perulae are unilacunar even
in species with trilacunar foliage leaves. : :

6. The significance of these findings is discussed in relation to the
taxonomic subdivision of the genus.

ACKNOWLEDGMENTS

This investigation was possible only with the assistance of ae aps
persons who supplied specimens. All those to whom we appealed re-

216 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

sponded with willing and full co-operation, and to them we tender our
sincere thanks. In particular, we wish to record the generosity of Mr.
R. B. Cooke of Corbridge, Northumberland, England, and Mr. H
Marchant of Stratford, New Zealand, who each contributed a large
number of specimens which together formed the basis of this investigation.
In addition, we are indebted to Mr. T. H. Findlay and the Crown Com-
missioners, Great Park, Windsor; to Mr. J. R. Sealey and the Royal
Botanic Gardens, Kew; to Dr. H. R. Fletcher, Dr. B. L. Burtt, Mr. H.
H. Davidian and Mr. P. J. Woods of the Royal Botanic Garden, Edin-
burgh; to Dr. D. Wyman and the Arnold Arboretum; to Mr. R. W.
Balch and the Botanic Garden, Dunedin; to Mr. C. Puddle and Bod-
nant Gardens, North Wales; to Mr. E. H. Lohbrunner, Victoria, Canada;
to Mr. David G. Leach, Brookville, Pennsylvania, U.S.A.; and to Mrs.
H. A. Hearn, Mr. R. Wilson, and Mrs. N. G. Mitchell of Christchurch,
New Zealand. We are indebted to Dr. R. D. Hoogland of the Depart-
ment of Land Research, C.S.I.R.O., Canberra; Dr. H. M. Burkill of the
Botanic Gardens, Singapore; Dr. H. Sleumer of the Rijksherbarium, Lei-
den; and Dr. R. A. Howard of the Arnold Arboretum, who all generously
contributed herbarium material of otherwise unobtainable species; to
Mr. G. D. Smith of the Northern Horticultural Society, Harlow Car,
Harrogate, England, and Mr. G. B. Petterson of the N. Z. Rhododendron
Association, who kindly supplied series of seedlings for developmental
studies; and to Mr. K. R. West of the Botany Division, D.S.I.R., Christ-
church, New Zealand, who with patience and skill prepared the illustrations
for publication.

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1968 | PHILIPSON & PHILIPSON, RHODODENDRON 217

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ay ; B. Ed. 1930. The species of Rhododendron: Rhododendron
Society, London.
Swamy, B. G. L. 1949. Further a to the morphology of the De-
generiaceae. Jour. Arnold Arb. 30: 10-
1954. Morphotaxonomic notes on ihe ae eee part A, Nodal
and petiolar vasculature. Jour. Madras Univ. B. 24:
SYNGE, P. M. Ed. 1963. Rhododendron Handbook, 1963. Es 1. Rhododendron
species in general cultivation. Roy. Hort. Soc. Londo
Watson, L. 1967. Taxonomic implications of a aannive anatomical study
of Riicidanese. New Phytol. 66: 495-504.

iW. PI [M. N

Botany DEPARTMENT Borany es
UNIVERSITY OF CANTERBURY i acscere OF SCIENTIFIC
CHRISTCHURCH AND INDUSTRIAL RESEARCH
New ZEALAND CHRISTCHURCH

New ZEALAND

APPENDIX: Comparison of Classifications, and Species Examined

SLEUMER, 1949 and 1958

HANDBOOK, 1963

SUBGENUS SECTION SUBSECTION SERIES
(EXCLUDED) — — |Camtschaticum
ENTANTHERA RHODORA — Azalea
VISCIDULA =
PENTANTHERA —
TSUTSUTSI BRACHYCALYX =
TSUTSUTSI —
TSUSIOPSIS —
AZALEASTRUM CANDIDASTRUM — Alb
UMEAZALEA = Leno
CHONIASTRUM — Stamineum
EUAZALEASTRUM — Ovatum
Hy MENANTHES — AURICULATA Auriculatum
Griersonianum
BARBATA Barbatum

MACULIFERA

ARBOREA
ARGYROPHYLLA
FLORIBUNDA
THOMSONIA
SELENSIA

MarTINIANA

CAMPYLOCARPA

SouLIEA

TRRORATA

Arboreum

Thomsonii

Irroratum

SUBSERIES

SPECIES EXAMINED

Canadense
Nipponicum

Schlippenbachii

Tashiroi

Barbatum

Glischrum

Crinigerum

Maculiferum

Arboreum

Argyrophyllum
)

Thomsonii

Cerasinum
Selense

Campylocarpum

Willi .
Trroratum

camtschaticum Pallas
albrechtit Maxim.
nipponicum Matsuma
uteum Sweet
roseum (Loisel.) Rehd.
schlippenbachii Maxim.
kaempferi Planch

moulmainense Hook.
—— Balf. f. &
est

Gas Hemsl.
oe Balf. f. &

ete Balf. f. &
ooper

smithii Nutt. ex Hook.

erosum Cowan

glischroides Tagg &

rres
habrotrichum Balf. f. &

bainbrideeanum Tagg &
Forrest

longesquamatum C.K.

mort Hayata
strigillosum Franch.
arboreum Smith
niveum Hook. f.

argyrophyllum Franch.
oribundum Franch.

cyanocarpum (Franch.)
W.W. Sm.

hoo Nu
thomsoniit Hook. f
cerasinum Tag

£
selense Franch.
eurysiphon Tagg &

orrest
callimorphum Balf. f. &
W.W. Sm.

campylocarpum Hook f.
—— Balf. f. &

pai — Sm.
een m Rehd. &

se taal Cowan
irroratum Franch.

wrayi King & Gamble

vaseyi A. Gray

occidentale A. Gray

yedoense Maxim.

esquiroli Lévl.
henryi Hance
oxyphyllum Franch.
stamineum Franch.

ovatum Planch.

barbatum Wall.

exasperatum Tagg
glischrum Balf. {. &
W.W. Sm.

monosematum Hutch.
pachytrichum Franch.

delavayi Franch.
zeylanicum Hort. ex
d

oud.
insigne Hemsl. & Wils.
hunnewellianum Rehd.

Wils.
eclecteum Balf. f. &
Forres

stewartianum Diels
viscidifolium Davidian
dasycladum Balf. f. &
W.W. Sm.
martinianum Balf. f. &

Forr
caloxanthum Balf. f. &

r
telopeum Balt. f. &

souliet Franch.

eritimum Balf. f. &

W.W. Sm.
pankimense Cowan &
Ward

SLEUMER, 1949 and 1958

SUBGENUS

SECTION

SUBSECTION

PARISHIA

LACTEA

FALCONERA

GRANDIA

FULVA

CAMPANULATA

TALIENSIA

APPENDIX: (Continued )

HANDBOOK, 1963

SERIES

SUBSERIES

SPECIES

EXAMINED

Lacteum

Falconeri

Grande

Fulvum

|Campanulatum

Taliense

Parishii

Adenogynum

cookeianum Davidian
venator Tagg

ashe il Diels

p eo Balf. f.
W.

Fa san Forrest &

pera ee Balt. {.&

coriaceum Franch.

ae
— Balf. f. &
Be seo Watt ex
Balf. £.

sg Balf. f. &
m.
PS tieier Balf. f. &
W.W. Sm

fulvum Balf. {.&
W.W. Sm

campanulatum oh Don
lanatum Hook.
succothii sae
bureavii Franch.

elliottii Watt. ex W.W.
Sm

lacteum Franch.
przewalskii Maxim.

wightii Hook. f.
basilicum Balf. f. &
W.W. Sm

eximeum Nutt.
fictolacteum Balf. f.
rex Lévl.

grande Wight

magnificum Ward

sidereum Balf. f.

uvarifolium Diels

detonsum Balf. f. &
Forrest

NERIIFLORA

PonrTICcA

ForTUNEA

Neriiflorum

Ponticum

Fortunei

Roxieanum
Taliense
sonii

Forrestii
Haematodes
Neriiflorum
Sanguineum
Caucasicum

Ponticum

Calophytum

Fortunei

gymnocarpum Balf. f. &
Ta

ge
roxieanum For
sphaeroblastum ‘Ball, i:
& For

‘iia ak fs
chamae-thomsonii (Tagg
& Forrest) Cowan &

Davidian
chaetomallum Balf. f. &
orrest ;

mallotum Balt. f. &
Ward

neriiflorum Franch.

oe Balf. f. &

sangineum Franch.
pales um Carriere
inot Tage

beronvasielncs Nakai
catawbiense Michaux
ponticum L.

ice Franch.

sutchuenense Franch.
decorum Franch. :
discolor Franch.
eiotesnersma Franch.
biculare Decaisne
grithionum Wight

pronum Tagg & Forrest

rufum Batal.

forrestii Balf, f. ex Diels

haematodes Franch.
sperabile Balf. f. &
Farrer

dichroanthum Diels

tea sea
ungernii Trau

macrophyllum D. Don

praevernum Hutch.

diapre pes Balf. f. &
W.W. Sm.

fortunei Lindl.

oreodoxa Franch.

APPENDIX: (Continued)

SPECIES EXAMINED

SLEUMER, 1949 and 1958 HANDBOOK, 1963
SECTION SUBSECTION SERIES SUBSERIES
_ PSEUDAZALEA sia — Trichocladum — asia Balf. £. & lepidostylum Balf. f. &
orrest Forrest
metnonthm Balf. f. &
a trichocladum Franch.
RHODORASTRUM — 2 Dauricum —_ Pes pe ssi — Kez;
PSEUDORHODORASTRUM TRACHYRHODION ~| Scabrifolium — malian Balf. f. & pube scens B .&
W.W. Sm. orre
bet euitees Franch. spiciferum Franch.
— BOTRYS wal vacemosum Franch.
BDORHODION —- — virgatum Hook. f
RHODODENDRON ears IERUM CAMELLIAEFLORA |Camelliaeflorum -— camelliaeflorum Haak, fi,
BooTHIA Boothii Boothii chrysodoron Tagg ex
Hutch. in ie Franch.
Megeratum leucaspis Tagg me “a Balf.{.&
TEPHROPEPLA Tephropeplum = auritum Tagg wie tes Balf. f.&
Farrer
xanthostephanum Merr.
EpGEWORTHIA Edgeworthii —_ edgeworthii Hook. f. pendulum Hook. f.
GLAUCA |Glaucophyllum Glaucophyllum brachyanthum Franch. charitopes Balf. f. &
Farrer
glaucophyllum Rehd. tsangpoense Hutch. &
Ward
GENESTERIANA me! genesterianum Forrest
MApDENTIA Maddenii Cilii burmanicum Hutch. ciliatum Hook. f.
ciliicalyx Franch. formosum Wall.
johnstoneanum Watt scopulorum Hutch
valentinianum Forrest veitchianum Hook.
Maddenii crassum Franch. maddenii Hook. f.
Megacalyx lindleyi Moore megacalyx Balf. f. &
: Ward

nuttallii Booth taggianum Hutch.
MOowupPINENSIA Moupinense ae moupinense Franch.
SALUENENSIA Saluenense — calostrotum Balt. {. & keleticum Bali. {. &
ard orres
nitens Hutch. radicans Balf. {. &
Forrest
saluenense Franch.
TRIFLORA Triflorum Augustinii augustinii Hemsl.
Hanceanum hanceanum Hemsl.
Triflorum ambiguum Hemsl. in ial Watt ex
Hut
lutescens Franch. cen Hook. f.
Yunnanense concinnum Hemsl. davidsonianum Rehd. &
Wils.
oreotrephes W.W. Sm. yunnanense Franch.
AMPYLOGYNA Campylogynum — campylogynum Franch.
CAROLINIANA Carolinianum — Seblseening re minus Michaux
So A. Gra
FERRUGINEA | Ferrugineum as jerrugine m L. kotschyi Simonk.
HELIOLEPIDA Heliolepis o desquamatum Balf. f. &
Forre heliolepis Franch.
neal Franch.
LApPPoNICcA Lapponicum —_ edgarianum Rehd. &
fastigiatum Franch.
figvliuin Franch. hip otond rae Balf. f.
ate

idoneum Balf. f. &
W.W. Sm. microleucum Hutch.
russatum Balf. f. &

CRANTHA Micranthum —_ micranthum Turcz.
CINNABARINA nabarinum. _— sertge ioe m Hook. f. concatenans Hutch.
reas eysii Nutt. xanthocodon Hutch.
LePmpota Lepidotum lepidotum Wall.
‘BAILEYA Baileyi baileyi Balf. f
UNIFLORA ludlowii Cowan patulum Ward

pemakoense Ward uniflorum Hutch. &
Ward

SUBGENUS

APPENDIX: (Continued)

SLEUMER, 1949 and 1958

SECTION

SUBSECTION

HANDBOOK, 1963

| SERIES

SUBSERIES

SPECIES EXAMINED

PoOGONANTHUM

VIREYA

PSEUDOVIREYA
SIPHONOVIREYA
HAEOVIREYA

MALAYOVIREYA

Stenophylla

Citrina
Buxifolia

Javanica

Anthopogon

Vaccinioides

anthopogon D. Don
sargentianum Rehd. &
Wils.

invasiorum Sleum.

saruwagedicum Foerster

spathulatum Ri
herzogii Warb.
beserinkianu Koord.
superbum Sle

num tack
yelliottii Warb.
jasminiflorum Hook.

gracilentum F. v. Muell.
saxifragoides J.J. Smith

taxifolium Merr.

purpureiflorum J.J.
Smith

citrinum (Hassk.) Hassk.

commonae Foerster

luteosquamatum Sleum.

brookeanum Low ex

javanicum (Bl.) Benn.
locha ]

e F. v. Muell.
macgregoriae F. v.
Muell.

cephalanthum Franch.
trichostomum Franch.
perakense King &

seimundii J.J. Smith
vaccinioides Hook. f.

konori Becc.
vinicolor Sleum.

pleianthum Sleum.

pauciflorum King &
Gamble

culminicolum F. v.
Muell.
intranervatum Sleum.

ong
robinsonii Ridl.

1968] NEVLING & GOMEZ-POMPA, HYDRANGEA 225

A NEW HYDRANGEA FROM MEXICO

Lorin I. NEVLING, JR., AND ARTURO GOMEz-PoMPA

THE GENUS Hydrangea is familar to both layman and scientist primarily
through species which have ornamental value. Since the species most
widely cultivated are shrubby, with the exception of the spectacular climb-
ing H. anomala D. Don, and usually have been selected for inflorescences
bearing large numbers of sterile flowers, one tends to have a distorted
impression of the genus as a whole. In the recent monograph by Elizabeth
McClintock (1957) two sections were recognized: HypRANGEA and Cor-
NIDIA, containing 11 and 12 species respectively. Section HypRANGEA in-
cludes most of the cultivated species, all of which are shrubs, excepting
H. anomala. Two species of this group, H. arborescens L. and H. querci-
folia Bartram, are native to the southeastern United States, the remainder
are Asiatic. In contrast, species of section CorNipIA seem to be more
flexible in growth form with many species occurring both as shrubs and
climbers, depending on circumstances of position and, perhaps, age. The
normal mature form, however, is a root climber. The species of this
section are native to the New World, excepting H. integrifolia Hayata of
Formosa and the Philippines. Thus 13 of 23 species are climbers.

In this paper a new climbing species of section Cornup1a is described
as Hydrangea nebulicola (Fics. 1 and 2). It was discovered in the decid-
uous liquidambar-oak forest of the Sierra de Chiconquiaco north of Jalapa,
Veracruz, México. This zone can be characterized floristically by the
abundance of individuals of Alnus, Clethra, Cyathea, Hedyosmum, Liquid-
ambar, Magnolia, Meliosma, Podocarpus, and Quercus (see Gomez-Pompa,
1966). The climate in this region is temperate due to altitude (ca. 1500
m.) and extremely humid, not only because of high rainfall (250-300 cm.
annually), but also because of the many clouds which sweep through the
forest. It is difficult to determine the temperature in this zone exactly.
However, from data obtained from a climatological station nearby N ao-
linco, Ver.) we are able to give a mean temperature of between “y and
18° C. The coldest month is January with an average of 12 to 14° C, the
extreme minimum temperatures during this period being some degrees
below 0° C. The topography is rather rugged with slopes of 10 to 40
percent. The soil, which is derived from volcanic rock, contains abun-
dant organic matter. The new species is abundant only locally, and it
possesses several characteristics which we believe to be of important evolu-
tionary significance within the genus and which mark it as a very special-
ized member of section Cornip1A. One of the most interesting aspects of
this species is its vegetative reproduction. We believe that this type of
reproduction has an adaptive value sufficient for the species even though
the scarcity of individuals is notable. Several trips have been made without

226 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Hydrangea nebulicola. (Gémez-Pompa 1541). tae branch with

Fic
details of inflorescence and pubescence of lower leaf-surfac

success to search for pistillate specimens. All that we have observed is that
the flowering of the species is rare although it may continue over several
months. With these considerations in mind and the realization that the
species is both rare and dioecious we can understand the importance of
vegetative reproduction and its réle in survival.

In climbing species of this genus, three kinds of shoots are generally
found: juvenile, found only in young plants; the terminal one, which func-
tions as a climbing shoot; and the lateral shoot, which supports the re-
productive functions and the primary photosynthetic activity. When a
tree is the substrate the climbing often is restricted to a single individual.
In this new Hydrangea several deviations from the general ground plan
just mentioned are to be found. Although no juvenile plants were located
in spite of a careful search, a fourth type of shoot bearing a marked re-

1968 | NEVLING & GOMEZ-POMPA, HYDRANGEA 227

Fic. 2; Flowering branch of ee nebulicola showing paired inflores-
cences at anthesis and young sine apie enclosed by cucullate bracts. Pho-
tograph from Nevling & Gémez-Pompa ;

semblance to juvenile shoots was found. These, termed runner-shoots,
were discovered along the forest floor, sometimes covered by leaf-litter.
They permit a single plant to climb several individual trees simultaneously.
Runner-shoots (Fic, 3) were quite common and are characterized morpho-
logically by reduced bract-like deciduous leaves and adventitious roots on
the lower and lateral surfaces, similar in external appearance to those found

few marginal serrations above the middle of the lamina but very soon
become similar to the mature leaf in form and size. The climbing shoot
(Fic. 3) is characterized by a very marked unequal production of secon-
dary xylem, the mass of the new wood being produced in the direction of
the substrate. This apparently permits the continuing production of the ie
ventitious roots necessary for attachment to the substrate. It is believe

that detailed comparison of the wood anatomy of climbing vs. non- regia
species of Hydrangea might prove useful in the determination of evolu-
tionary direction within the genus.

The inflorescences of species of Hydrangea are marked by having cape
monomorphic or dimorphic flowers. These two conditions are setts Dy
students of the genus as fertile and sterile flowers: fertile — a
bisexual with androecium and gynoecium normally developed; sterile

228 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

4

Fic. 3. Young climbing shoot of Hydrangea nebulicola showing side normally
eo substrate. Note adventitious roots and lateral branches, < approx-
imately 3

ere x of runner-shoot of Hydrangea nebulicola showing reduced bract-
like leave es, approximately natural size.

flowers having neither androecium nor gynoecium normally developed but
with the calyx lobes becoming greatly enlarged and showy. An intermedi-
ate type with enlarged calyx lobes and functional reproductive organs is
found in H. scandens (L.f.) Seringe. The inflorescence in this species also
contains “normal” fertile flowers. Inflorescences containing both fertile
and sterile flowers generally are considered more specialized than those
composed of fertile flowers only. The addition of sterile flowers or the
conversion of fertile flowers to sterile ones probably is related to increased
efficiency in terms of insect attraction. Although in some species such

1968 | NEVLING & GOMEZ-POMPA, HYDRANGEA 229

as H. anomala D. Don the number of sterile flowers per inflorescence varies
somewhat from year to year, the presence or absence of sterile flowers
appears to be a relatively reliable taxonomic character. Two notable ex-
ceptions are H. arborescens L. and H. scandens in which considerable vari-
ability occurs.

In Hydrangea nebulicola another floral condition is present: the in-
florescences contain only fertile flowers which appear to be functionally
unisexual. In our field studies one mature plant has been observed to be
staminate through two flowering cycles and for this reason we are inclined
to believe that the plants are dioecious rather than monoecious, Although
McClintock has suggested proterandry for H. serratifolia (Hook. & Arn.)
Phil. f., we do not make a similar interpretation for H. nebulicola. In
the flowers examined, pollen stainability with cotton blue in lactophenol
is approximately 98 percent and all grains appear to be properly formed.
Ovules have not been observed and the sterile placentae are enlarged,
filling the locule, and leaving little room for ovule development. In addi-
tion, no discernible stigmatic surface was found on the styles. Curiously,
the placentation at the base of the ovary is of the axile type noted by
McClintock but the massive central trace divides almost immediately
into two smaller traces which supply the two large parietal placentae,
the ovary being bicarpellate and uniloculate (Fics. 5-12). Near the
apex of the ovary the two placentae divide so that the two carpels
are open and their margins free from one another. At the roof of the
hypanthium the carpels are open (the opening in the form of a linear slit).
It is at such a minute opening that the terminal poricidal dehiscence of
Hydrangea fruits takes place in other species. The two styles reflect the
open condition by being obviously grooved nearly to their apices. Un-
fortunately, it must be recalled that the situation described applies only to
staminate flowers and confirmation must await discovery of pistillate
flowers (although in the bisexually flowered H. arborescens the placental
configuration is identical with that of H. nebulicola). If we are correct in
our interpretation of sexual dimorphism in this species, the pistillate flowers
should have smaller petals, shorter stamens, no stainable pollen, longer
styles, papillate stigmatic surfaces, and many ovules borne on parietal
placentae. :

The evolutionary significance of the “advancement” of bisexual flowers
to unisexual ones is difficult to estimate in Hydrangea because of the few
floral characteristics in which the species vary and the small degree of the
variation. The recognition of sexual differences among fertile flowers may
be helpful in defining evolutionary pathways. Solely on the basis of the
unisexuality of the flowers, H. nebulicola would seem to be a specialized
member of section Cornip1a which is in turn more advanced than section
HYDRANGEA. Fre

A case of sexual dimorphism similar (but not precisely identical because
of the presence of sterile flowers) to that just described appears to be
found in Hydrangea oerstedii Briquet and H. peruviana Moricand. The
characters which McClintock employs in her key to Hydrangea to dis-

230 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

IGs. 5-12. Hydrangea nebulicola ar os & St wie 39). Fic.
Cleared flower showing grooved style, Fic. ransverse — of
flower. Fic. Section at level "ee eee ee note snl 8.
Fic. 7. Section at level of hypanthium roof showing op rpels oa siylar
canals, X ca. 58. Fic. 8. —- of beg seca below Gepantuiins roof
rach free margins of plac ntae, X ca. 25. Fic. 9. Section at mid-ovary show-
g the large, sterile, parietal placentae, 6 ca. 25 ts

Magnification of one
plac nose (inclined 90° from pr

0
evious figure ) showi ing vascularization and absenc
Ss

o x 58. Tie 11. Section . ban of ag acentae. Note fusion a
iinage traces with each ca el becoming closed, & ca. 5 he Section at
base c vary demonstrating bilocular cndian Sad conan) ventral trace,
xX ca Sf.

1968 | NEVLING & GOMEZ-POMPA, HYDRANGEA 231

tinguish these species are (in part) the relative length of stamens and styles
of fertile flowers. In H. oerstedii the stamens are 2—4.5 mm. long and the
styles are 0.3-1 mm. long, while in H. peruviana they are 0.4-1.6 mm.
long and 1-2 mm. long, respectively. McClintock suggests in her discussion
that the two may represent two forms of a dimorphic species. It is our
suggestion that these two species do reflect two sexual forms of the same
species. In addition to the larger stamens in H. oerstedii is accompanying
high pollen stainability (with cotton blue in lactophenol). The short-
styled flowers lack both papillate stigmatic surface and functional ovules.
In H. peruviana the short stamens have no stainable pollen, although the
anthers appear reasonably well formed. The long-styled flowers have a
papillate stigmatic surface and produce many well-developed ovules. The
vegetative morphology and geographic distribution present no great dif-
ficulty in the interpretation of these two taxa as functionally staminate
and functionally pistillate components of the same species.

Hydrangea nebulicola Nevl. & Gémez-Pompa, sp. nov.

Planta fruticosa scandens ad 30 m. alta, sempervirens; ramuli ferruginei,
glabriusculi. Folia simplicia, opposita, elliptica, 7-16 cm. longa, 4-9 cm
lata, apice acuminata, basi cuneata, coriacea, pubescentia pilis stellatis,
6-8-nervia; petioli 1-4 cm. longi. Inflorescentia in umbellis oppositis,

latae, 1.5-3 cm. longae et latae. Flores ¢: hypanthia turbinata, 1-1.25
cm. longa, 1.5 mm. in diametro; petala 4, alba, ovata, 2-2.5 mm. longa,
1.25—2 mm. lata, caduca; stamina 8 vel 9, libera, quam petala longiora;
filamenta 2—2.5 mm. longa; antherae oblongae; pistillum 2-carpellatum,
ovulis 0; stylis effusis, ca. 0.5 mm. longis; stigmatibus 0. Flores 92 et

fructus non visi.

Powerful root climber, evergreen, to 20 cm. in diameter at the base, to
30 m. tall; bark of mature trunk gray, smooth and non-peeling; young
shoots stout, tomentose, the trichomes stellate but often only the basal
stalk persistent and then appearing simple, ferrugineous. Leaves simple,
opposite, coriaceous, broadly elliptic, 7-16 cm. long, 4-9 cm. broad, acumi-
nate at the apex, more or less cuneate at the base, sparsely ferrugineous-
pubescent to glabrescent above, moderately ferrugineous-pubescent be-
neath, the midrib plane above, emersed beneath, the primary lateral veins
6-8 pairs, slightly arcuate, the margin entire; petiole 1-4 cm. long. In-
florescences borne in opposite axillary pairs near the apex of lateral (non-
climbing) shoots, the axillary shoots 3-7 cm. long, ferrugineously felted,
the inflorescences compound, umbelliform, 5—10 cm. in diameter, enclosed
in bud by cucullate bracts, the bracts nearly orbicular, 1.5-3 cm. long and
broad, chartaceous, pubescent, caducous. Sterile flowers absent. Fertile
flowers unisexual by abortion, plants functionally dioecious (?). Staminate
flowers: creamy white, glabrous throughout; pedicel ca. 2.5 mm. long;
hypanthium turbinate, 1-1.25 mm. long, 1.5 mm. in diameter; calyx lobes

232 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

free, very shallowly deltate; petals 4, valvate, free, ovate, 2-2.5 mm. long,
1.25-2 mm. broad, caducous; stamens 8 or 9, free, the filaments folded
in bud becoming erect at anthesis, the filaments 2-2.5 mm. long; anthers
greatly exserted, oblong, ca. 2 mm. long, 1.5 mm. broad, basifixed, de-
hiscing by lateral slits; pistillode 2-carpellate, inferior, ovules 0, the 2
styles free, recurved, ca. 0.5 mm. long, slightly canaliculate, stigmatic sur-
face not apparent. Pistillate flowers and fruit not seen.

México. Veracruz: region de Misantla, Sierra de Chiconquiaco y
Misantla, a 1540 m. de altitud en bosque de encino-liquidambar, 9 de
julio 1966, A. Gémez-Pompa 1541 (holotype MEXxu, isotype GH); R.
Hernandez M. 306 (mexu), Nevling & Gémez-Pompa 39 (GH, MEXU).

Although this new species is placed in section CorNip1A without hesita-
tion, its affinities with other species of the section are still uncertain. Prior
to any speculation concerning affinities, the entire section should be re-
examined in terms of problems of floral sexuality mentioned previously.
In addition, there is at least one, possibly two, undescribed species of
Hydrangea from Chiapas referable to this section. Until these species are
studied, definitive placement seems fruitless.

BIBLIOGRAPHY
Burkett, G. W. Anatomical studies within the genus Hydrangea. Proc. Indiana
Acad. Sci. 41: 83-95. 1932
GOmeEz-Pompa, A. Estidios botanicos en la regién de Misantla, Veracruz. Ed.

irs $99 Méx. de Recursos Naturales Renovables, A.C. México, D.F. pp.
966.

McCuintock, E. The cultivated hydrangeas. Baileya 4: 165-175. 1956.
~———.. Hydrangeas. Natl. Hort. Mag. 36: 270-279. 1957,

. A monograph of the genus Hydrangea. Proc. California Acad. Sci. 29:
147-256. 1957.

edie I., Jk. Climbing hydrangeas and their relatives, Arnoldia 24: 17-

ARNOLD ARBORETUM AND GRAY HERBARIUM
CAMBRIDGE, MASSACHUSETTS
AND
INSTITUTO DE BIOLOGIA

UniversipAp NAcIonAL AUTONOMA DE MExico
Mexico, D.F.

1968] MUNZ, ASIAN SPECIES OF DELPHINIUM 233

A SYNOPSIS OF THE ASIAN SPECIES OF DELPHINIUM,
SENSU STRICTO*
Puitre A. Munz
GROUP X

Leaves rather coarsely lobed as in Groups VIII and IX, but the spurs
strongly curved to make about a half circle.

This group, also, is an artificial one and does not agree with any tax-
onomic subdivision proposed for the genus.

Key TO SPECIES

—

. Sepals with subapical cornute appendages 1-2 mm. long; flowers purplish
blue

2. The sepals 18-20 mm. long; spur 20-22 mm. long. (These two species
treated in Group VI
3. Ultimate leaf-lobules 5-8(—10) mm. wide. Yunnan, etc. ...........
Evaro. ok os a ee . D. ceratophorum.
3. Ultimate leaf-lobules 2-4 mm. wide. Tibet. 145. D. se i ee ea
2. The sepals 15-18 mm, long; spur to 18 mm. long. Szechwan. ...........
Bar Soe. eT Sa ea 236. D. coimplocenirwn
. Sepals lacking stare we cornute appenda
4. Stem and axis of inflorescence largely ee
5. Sepals 10-11 mm. long.
6. Inflorescence racemose; pedicels 3.5-9 cm. long. Szechwan. (This
species treated in Group Vi ee 154. D. glabricaule.
6. Se peso a diffuse panicle; pedicels 1.5—3 cm. long. Yunnan
239. D. polyanthum.

pane

5. Sauls 15-25 mm. long.
7. Bracteoles 7-15 mm. long. Yunnan. ........ 237. D. eriostylum.
7. Bracteoles shorter, 2-7 mm. tin ‘Szec hwan
8. a more or less dark; sepals more or - Jess strigulose, 17—20
mM 1ONR oc: ee eee 240. D. potaninii.
8. Pele bluish: sepals more loosely pubescent, 13-17 mm. long.
244, D. tongolense.
4. Stem and/or axis of geiitercns more or less hairy or pubescent.
9. Sepals 18 mm. or m ong.
10. Sepals abr” soe 17-18 mm. long. Jagger spade in
Group IR) ea mariae.
* Continued from volume 49, p. 166. With this section of the paper the ane series
of stiles by Philip A. Munz on the genera Delphinium and Consolida is completed.
The series was initiated in volume 48 of this Jee ad includes: A synopsis of

species of Consolida da (Ranunculaceae), ibid. 1 159-202; and ie first two portions of the
Present paper, ibid. 249-302, 476-545, which is continued in the first number of the
present volume, 49: 73-166. Ed.

234 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

10. Sepals more or less pubescent; spur 20-25 mm. long. China.
ee ee ar ee 243. D. spirocentrum.
Sepals shorter
+1. pi divisions long-acuminate; spec at middle of pedicels.
a. id ie oe CESS . D. pseudo-tongolense.

2

is Leaf ee not long-acumin
12. Upper petal bidentate < Ate upper sepal largely 15-18
m
13. "Inflorescence racemose; leaves with coarse teeth; brac-
teoles near the flower. .......... 235. D. bulleyanum.
13. Inflorescence panculate leaves = teeth; brac-
teoles near middle of pedicel. ...... 4. D. tongolense.

_
nN

Upper petal entire to janet erose at ante

14. Lamina of lower petal not divided.

_ Limb of the lower petal elongate-linear. Asia
Media. (Treated in Group VIII). ...........---:
pe I ee ee coy jive De knorringianum.

. Limb of the lower petal oblong, 5 mm. wide
Szechwan. (Treated in Group VIII). ..........-
HES a ae a ee ae 156. D. hirticaule.

14. Lamina of lower petal divided.

16. Upper sepal 20-25 mm. long, the spur 16-18 mm.
long. Altai. (Treated in Group IX
re a re ee ie 217. D. mirabile.

16. Upper sepal shorter.

17. Sepals with appressed hairs, spur almost twice
as long as blade. Yunnan. 238. D. hamatum.
17. Sepals with spreading hairs. Szechw Walco.)
eee So os 242,20), sino-witijouien:

—
Leal

235. Delphinium bulleyanum G. Forrest ex Diels, Notes Bot. Gard.
Edinb. 5: 265. 1912. Fic. 19, A.
Delphinium spirocentrum Hand.-Mazz. var. pauciflorum Chen, Bull. Fan Mem.
Inst. Biol. (n.s.) 1: 170. 1943. Based on Vii 15544, (pm), taken on the
east flank of the Lichiang Range, Yunnan, 2800 m.; seen.

Perennial from a woody root, the stem 3-14 dm. tall, glabrous below,
spreading pubescent and more or less glandular in the inflorescence, often
subglaucous and somewhat violet, simple at least below; leaves well dis-
tributed, but gradually reduced up the stem, the lower petioles to 2 dm.
long, glabrous, slightly dilated at the base, the uppermost leaves becoming
foliose bracts; blades rounded-pentagonal in outline, 3-10 cm. broad
glabrous to scattered strigulose on both surfaces, pale beneath, 5-fid isto
subcuneate incised-serrate triangular-ovate segments; inflorescence rather
laxly racemose; upper bracts setose to lanceolate, lower foliose and divided;
pedicels strict, 1-10 cm. long, pubescent; bracteoles setose to widest
usually 5-10 mm. long, located near the flower; sepals rich purple to deen
blue-lavender, pubescent without; upper sepal 10-20 mm. long, 8-12 mm
wide, ovate, obtuse to acute, the spur 18-22 mm. long, 4 mm. wide at base
rather narrow at tip, recurved at the middle to make an almost panei

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 235

circle; lateral sepals oblong, rounded at apex, 12-15 mm. long, 7-9 mm.
wide, glabrous except on the midveins; lower sepals 12-18 mm. long, 8-10
mm. wide, acuminately obtuse, pubescent; petals bluish, the upper laminae
slightly oblique, glabrous, 9-11 mm. long, cleft for 1 or 2 mm., the spur
18-20 mm. long; lower laminae oblong, 5 mm. long, cleft halfway into 2
lance-oblong ciliate lobes, bearded at base, claw 4.5—5 mm. long; stamens
6-7 mm. long, glabrous; anthers oblong, dark, 1.3 mm. long; follicles
3(—4), somewhat divergent toward apex, sparsely pilose, 10-12 mm. long,
the styles an additional 3 mm. long; seeds dark, 1 mm. long, obpyramidal,
covered with minute scales.

Type: Moist rocky places among scrub on eastern flank of the Lichiang
Range, Yunnan, at 3500-3800 m., Aug. 1906, G. Forrest 2694 (£); seen.
Isotype (BM); seen.

Rance. A species of western Yunnan and southwestern Szechwan, at
elevations of 2800 m. to 5100 m.

ExampLes, Yunnan: Rock 9924, 9669, 9766, 9755, 6082, 6085, 5696, 10797,
6091; K. M. Feng 1982, 2001; Forrest 6556, 11130; Kingdon Ward 995. Szech-
wan: Kingdon Ward 4846, 4811.

236. Delphinium campylocentrum Maxim. Act. Horti Petrop. 11:

31. 1889 Fic, 19, 5.

Delphinium tongolense Rehd. & Kobuski, Jour. Arnold Arb. 14: 11. 1933,
pro parte, not Franchet, 1893.

To about 7 dm. tall, the stem simple below, with a few scattered up-
curved branches in the inflorescence, scattered-pilose below, more densely
so above and more or less glandular, leafy, the leaves rather evenly de-
veloped; petioles slender, with somewhat dilated base; blades mostly 3-fid,
the segments subsessile, cuneate, mostly 3~5 cm. long, the 2 lateral bilobed,
then pinnately incisely lobed or coarsely toothed, the central palmately
lobed or toothed, these lobes broadly lanceolate, mucronate, strigulose,
paler beneath than above; inflorescence open, paniculate, several flowered ;
bracts mostly narrowly linear, entire, to ca. 10 mm. long; pedicels 2-5

Type: “Ad limites Kansu et Szetschuan, Valle fl. Heiho, 24 Julii 1885,”
Potanin (LE); seen.

236 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Ficure 19, Delphinium, Group X. Petals, the upper — the lower

clawed. A. D. bulleyanum; upper lami . long, spur 18 mm.; lower
| 5 g, claw 4 mm.; drawn from Rock 992 rs B ne ‘campy-
locentrum gril bare 12 mm. long, spu mm.; lower lamina 6.5 mm. long

lon :
n from Harry Smith 4138 (a). C. D. phe iaeeig upper
lamina 11 mm. tok Ne spur 17 mm.; lower lamina 9 mm. long, claw
from Esquirol 141, TYPE (£). D. D. hamatum; upper lamina 10 mm. Pont
mm.; lower lamina 9 mm. long, claw 5 mm.; ‘drawn from Rock 22852 AS ane

amina 7

(pm). F. D. potaninii; upper lamina 12 mm. long, spur 20 mm.; lower area 7
mm. long, claw 5 mm.; drawn from Farges 630, Decor a “es — bie
D. potaninii; oto lamina 10 mm. long, spur 3m 7 lon 4
claw 5 mm.; drawn from E. H. Wilson 1999 (GH). es D. ‘sinovitifolnm ‘upper
lamina 9 mm. ale spur 23 mm.; lower cae 7m g, claw ; drawn
from E£. H. Wilson 3088, ISOTYPE fe. be seein upper faniina 10

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 237

SPECIMENS SEEN. Kansu and Szechwan: at about 3000 to 3500 m., i
74095 (£); H. Smith 4138 (a, BM); Ching 849 (g£, GH, Ny, US); Rock 13161
(GH, NY), 13111 (GH, NY), 13109 (GH, US).

237. Delphinium eriostylum Lévl. Bull. Herb. Boiss. IT. 6: 505. 1906.
Fre. 19, G,

Apparently tall robust perennial, the stem coarse, smooth, glabrous,
remotely leafy; upper leaves 3—5-fid almost to base, with scattered stiff
hairs on both surfaces but especially on veins beneath, the blades 5—12 cm.
wide, each segment narrow cuneate-obovate, incised ca. halfway to midrib
into lance-ovate lobules or teeth 3-10 mm. long, 1-5 mm. wide, acute to
acuminate; inflorescence a few branched open panicle, each raceme 2- to
6-flowered; bracts stiff-hairy, lance-linear, 2-3 cm. long, 2-3 mm. wide,
long-acuminate; pedicels stiff, 2-7 cm. long, arched-ascending to suberect,
with few to rather many spreading glandular hairs; bracteoles linear-sub-
ulate, 7-15 mm. long, bristly, near the middle of the pedicel; sepals violet,
subglabrous; upper sepal ovate, 14 mm. by 9 mm., obtuse, the strongly
curved spur ca. 20 mm. long; lateral sepals round-ovate, 16-18 mm. by 10
mm., rounded-obtuse; lower sepals elliptic-obovate, 18-20 mm. by 8 mm.,
pointed; petals bluish, the upper laminae slightly oblique, 12 mm. long, 4
mm. wide, glabrous, bidentate, the spur ca. 16 mm. long; lower laminae
perpendicular to the claws, 7 mm. by 3 mm., bifid to beyond middle into
lance-oblong lobes ca. 4.5 mm, long, ciliate and somewhat bearded;
stamens 6-7 mm. long, glabrous; anthers dark, linear-oblong, 1.5 mm.
long; carpels 3, sparsely stiff-hairy.

Type: Kweichow, Ta-tsong, Kouy-hoa, Esquirol 141 (£); seen.

Another specimen came from the same area, 2095 without collector’s
name (£), and one from Yunnan-sen district, Gan-chouen, Cavalerie 4486
(E).

238. Delphinium hamatum Franchet, Pl. Delavay. 28. pl. ri React Pe
G. LY,

Delphinium ceratophorum var. brevicorniculatum W. T. Wang, Acta Bot.
Sinica 10: 157. 1962. Based on T. T. Vii 9598 from Yunnan (A); seen

Perennial from a large root system; stem ascending, to ca. 2 dm. long,
mostly solitary, spreading-hairy, the hairs ca. 1 mm. long; leaves basal,
the petioles purplish, slender, hairy, 3-12 cm. long, thin-vaginate at base;
blades 3- or 5-partite, pentagonal, 1-7 cm. long, somewhat wider, the

spur 20 mm.; lower lamina 7.5 mm., claw 5.5 mm.; drawn from

lon
Baia j ss D.s sees “upper lamina 9
87, TYPE of D. pediforme c) J. pr w 6 mm.; drawn from

ng, spur 16 mm.; lower lamina 7 mm. long, er Nai
Smith 11594 (a 2 ke D. geen eee upper lam m. long, spur 17 mm.,;
lower lamina 5 mm. long, claw 5 mm.; drawn oa Soulié "396 (P).

238 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

lobes obovate-cuneate, strigulose above, more loosely pubescent beneath,
ca. 1 cm. wide and with few coarse triangular teeth 1-6 mm. long; flowers
1 to few, racemose; lower bracts often foliose, the others lance-linear, 5—15
mm, long, acuminate; pedicels arcuate-erect, 1-3 cm. long; bracteoles 2,
linear, to 1 mm. broad, near the flower; sepals purplish blue, strigulose
without, the upper sepal ovate, 13-16 mm. long, 8-10 mm. wide, acutish,
the spur 22-25 mm. long, 3.5 mm. wide at base, tapered gradually to the
hamate apex; lateral sepals ovate-oblong, strigose on midvein, rounded at
apex, 15-17 mm. by 8-10 mm.; lower sepals elliptic-ovate, 16-20 mm.
long, 10-12 mm. wide, obtuse; petals pale blue, the upper laminae almost
straight, 10 mm. long, glabrous or nearly so, entire to emarginate; the spur
22 mm. long; lower laminae bearded, 9 mm. by 8 mm., cleft to ca. 4 mm.,
the claw 5 mm. long; stamens 6-7 mm. long, slightly ciliate; anthers dark,
1.5 mm, long; follicles 3, short-hairy, oblong, 12-15 mm. long, the styles
an additional 3 mm.; seeds brown, 2 mm. long, winged on angles.

Type: Yunnan, “in collibus calcareis ad Fang-yang-tchang supra Mo-
so-yn, alt. 3200 m.,” Oct. 14, 1887, Delavay (P), seen; isotype (K), seen.
Additional sheets with somewhat different data as to elevation are (K, NY).

DistripuTion. A species from Yunnan, largely toward Tibet.

COLLECTIONS SEEN. Forrest 15555 (K); Rock 23457 (NY), 22852 in part (NY,
ee us), Delavay 1676 (K, P), 2676 (pP); T. T. Vii 9832 (a), 12702 (a), 15544

239. Delphinium polyanthum W. T. Wang, Acta Bot. Sinica 10: 145.
1962. Fie.19; 2:
Stem 1.5—2 m. tall, ca. 8 mm. in diameter, glabrous, branched, equally
foliose; the middle cauline leaves with non-vaginate petioles slightly longer
than the blades; the latter broadly pentagonal, 6.5 cm. long 10 cm. wide
cordate at base, tripartite 6/7 of their length, the middle part entire
below, 3-fid to the middle, the lobes further divided or toothed, the lateral
parts unequally bifid to below the middle, loosely strigulose beneath hirtel-
lous on nerves above; racemes 15-20 cm. long, forming a terminal pyra-
midal panicle; peduncles 6-10 cm. long, glabrous, axes with white re-
trorsely appressed hairs above intermixed with yellow glandular hairs;
ai divergent, 1.5—3 cm. long; bracteoles just above middle of pedicel,
eh ait ca xe long, 0.3 mm. wide; sepals blue, elliptic, 10 mm. long,
ee y puberulent; spur 18 mm. long; upper petal laminae yellowish,
glabrous, emarginate; lower laminae bluish, yellow bearded below the
middle, bifid into lance-triangular lobes, ciliate, the claw about as long as
blade; stamens glabrous; carpels 3, glabrous, 11-20 mm. long in imatarit :
seeds obconic-triquetrous, 1.5 mm. long, scaly laminate ‘ Z
Wang 69805 (PE), not seen; photo. (E), wee
Two herbarium sheets (Shansi, Licent 7719 (pm) and W. Kansu, Licent 4279

(BM) agree with the descripti
ak escription and photograph of the type, but have glabrous

Type: Yunnan, Teh-chin, Pai-ma-shan, 3500 m., Sept, 1935, Cy W.

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 239

240. gee potaninii Huth, Bull. Herb. Boiss. 1: 332. pl. 14.
June Fic. 19, F, G.

ae otto var. potaninii Brihl in Briihl & King, Ann. Bot.
Gard. Calc. 5: 896.

? Delphinium pee: var. latibracteolatum W. T. Wang, Acta Bot. Sinica
10: 147. 1962. Based on F. T. Wang 22014 et I have not seen, but
which has bracteoles 10-12 mm. long, 1-1.2 m ide.

— a Franchet, Bull. Soc. Philom. ‘Paris VIII. 5: 164. (Au-
gust?) 1 Type from Szechwan, district Tchen-kéou-tin, Farges 630
(P), seen cae sheets all with lower petal laminae deeply div ide

Deiphiniurn: bonvalotii Franchet, loc. cit. 165. (August?) 1893. Type, Szech-
wan, near Ta-tsien-lou, Prince H. d’Orleans & Bonvalot (P); seen. Lower
petal lamina deeply lobed, gaping.

Perennial from a thick root, the stems several, erect, thick, leafy, sub-
glabrous, tall; petioles glabrous to hairy, the lower 1 dm. or more long,
scarcely dilated at the base, the upper shorter; lower leaf blades broadly
rounded in outline, 8-15 cm. broad, subglabrous or with stiff hairs along
the veins beneath, 5- or 7-parted into broad cuneate-obovate segments,
these incisely lobed or toothed with sharp angles and acute lobes or teeth
8-30 mm. long; upper leaves reduced, 3- or 5-parted; inflorescence an
open panicle, largely glabrous; lower bracts foliaceous, 3-parted into
lanceolate lobes to ca. 3 cm. long, upper bracts entire and lanceolate to
setaceous, then 3-6(—10) mm. long; pedicels arched or straight, more or
less divaricate, slender, 1-5 cm. long; bracteoles setaceous to linear-
lanceolate, 2-6 mm. or more long, near the middle of the pedicel; sepals
bright blue or with some violet, more or less strigulose without; upper sepal
ovate, 10-20 mm. long, 5-12 mm. wide, the spur 20-30 mm. long, 3-4.5
mm. wide at base, strongly curved, with a blunt tip; lateral sepals elliptic,
18-22 mm. long, 10-12 mm. wide, rounded at apex; lower sepals elliptic-
rhombic, ans vii long, 8-9 mm. wide, rounded at tip; petals bluish, the
upper lepatias almost straight, 8-10 mm. long, entire to emarginate,
glabrous; lower —— hirsute at base, ciliate on margins, oblong to
oblong-obovate, 6-8 mm. long, mostly divided into lance-oblong lobes 3-4
mm. long, claw ca. 5 mm. long; stamens glabrous, 6-7 mm. long; anthers
dark, 1.3-1.6 mm. long; follicles 3, divergent, glabrous to pubescent,
10-13 mm. long, the styles an additional 2.5 mm.; seeds brown, ca. 1.3
mm. long, covered with minute scales.

Type: Kansu, China, G. N. Potanin, 1885 (B), not seen; isotype (K),
seen.

Rance. Apparently quite ian: distributed in western China, at eleva-
tions of about 3000 to 4200 m

EXAMPLES, Szech E. H. Wilson 1999 (BM, GH, US), 819 (GH, US), 1088
(BM), 3095 (a), 3007 (Ky. 4702 (xk, P), 1169 (cH); K. L. Chu 3246 (e); Henry
8792 (GH, P); Pratt 344 (K); C. L. Sun 476 (us); Rock 16743 (A, us), 23992
(uc), 16637 (us), 16694 (GH. NY, US), 24166 (GH, UC); W. P. Fang 1480 (vs);
Harry Smith 4072 (a); T. K. Wang & T. S. Wen 581 (a); W. Hupeh: W ilson
1323 (NY). Yunnan: Ducloux 5153 (P).

240 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

241. Delphinium pseudo-tongolense W. T. Wang, Acta Bot. Sinica
10::153.. 1962.

Stem 6-10 dm. high, 5 mm. in diameter at base, glabrous, often branched,
equally foliose; median cauline leaves with petioles 7-8 cm. long, not
vaginate; the blades broadly pentagonal, ca. 7 cm. long, 11-12 cm. wide,
cordate at base, 3-parted to 6-10 mm. above the base, the middle part
rhombic-ovate, acuminate, 3-lobed to below the middle, the lobes with
lobules acutely dentate; lateral parts bifid and with secondary or tertiary
lobules loosely strigulose beneath, loosely hirtellous along veins on upper
surface; racemes 5-18 cm. long, axillary and terminal, forming an open
panicle; pedicels 1-3 cm. long, ascending; bracteoles 4.5-7 mm. long, near
middle of pedicel, with spreading yellow glandular hairs and retrorsely
appressed white ones; sepals bluish purple, oblong to oblong-obovate, 12—
19 mm, long, puberulent; spur about as long as sepals, subulate, spirally
decurved at middle; upper petal laminae blue or yellowish, glabrous,
rounded or emarginate at apex; lower laminae blue, ovate, 2-lobed to
middle, yellow bearded at base, long-ciliate, claw appendiculate, as long as
limb; stamens glabrous; carpels 3, glabrous or puberulent above.

Type: Szechwan, Pao-Lsing, T. P. Soong 38915 (PE), not seen; photo.
(£), seen.

Said to be near D. tongolense, but with long-acuminate leaf parts, brac-
teoles at middle of pedicel, and spur spirally decurved.

242. Delphinium sino-vitifolium W. T. Wang, Acta Bot. Sinica 10:
152; 1963. Fic. 19, H.
Bhi vitifolium Finet & Gagnep. Bull. Soc. Bot. France 53: 126. 1906,

t Willd.

Stem slender with soft spreading hairs, 3.5-4 dm. high; cauline leaves
few, pilose on both surfaces, slightly cordate, 3—5 cm. in diameter, 3-parted
less than halfway into lobes broader than long, these lobes in turn divided
into semi-ovate mucronate-acute lobules 3-5 mm. long, somewhat broader;
petioles 3-5 cm. long, hirtellous; upper leaves smaller, 1—-1.5 cm. long,
sessile; raceme lax, few flowered, simple; lower bracts foliose, to 1.5 cm.
long, the upper subfiliform, pilose, ca. 8 mm. long; pedicels hirtellous,
slender, 3-4 cm, long; bracteoles setiform, pilose, 5-6 mm. long, removed
as much as 1 cm. from the flower; sepals blue, pubescent, the upper ca. 10
mm. long with a spur 25 mm. long and hamate at apex; other sepals
deltoid-ovate, ca. 12 mm. long; petals bluish, the upper laminae oblique
entire, rounded at apex, 9 mm. long, the spur 22-24 mm. long; lower
laminae oblique, oblong-ovate, 7 mm. long, bearded yellow white-ciliate
divided one third its length, claw hirtellous, appendiculate 5-6 mm. long
anthers dark, elliptic; carpels 3, glabrous. é

Type: Szechwan, 3200-4300 m., Sept., 1904, Wilson 3088 (Pp, A); seen.

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 241

243. Delphinium spirocentrum Hand.-Mazz. Symb. Sinicae Pt. 7:
280. 1931. Fie. 19, I, J.
ees spirocentrum var. hirsutum Chen, Bull. Fan Mem. Inst. Biol.
ping (n.s.) 1: 169. 1948. Isotype, T. T. Vii 7769 (a) from Yunnan;

seen.

Delphinium spirocentrum var. grandibracteolatum W. T. Wang, Acta Bot.

Sinica 10: 153. 1962. Isotype, Yunnan, 7. T. Vii 9018 (a); seen.
2S habla pediforme Comber, Notes Bot. Gard. Edinb. 18: 237. 1934. Type,
a pea divide, Salwin Valley, Forrest 17187 (£), seen;
ud (P),s
Delphinium Seamer var. pediforme (Comber) W. T. Wang, Acta Bot.
Sinica 10: 152. 1962.

Stems usually more than one, simple, 1-9 dm. tall, rather slender,
densely retrorse-villous, scattered leafy; lower petioles to ca. 3 dm. long,
villous, more or less dilated at base, the upper cauline much shorter;
leaves rounded-pentagonal in outline, 3-14 cm. in diameter, green and
strigose above, paler and with longer stiffer hairs beneath, 5-fid to within
1 cm. of the base, the segments cuneate-obovate, rather shallowly pin-
natifid into few lance-oblong teeth or lobules 2-5 mm. wide and rounded
to obtuse at apex; racemes simple, rather few flowered, to ca. 2 dm. long;
pedicels erect, 2-8 cm. long, with spreading or somewhat retrorse hairs;
bracteoles linear, 8-10 mm. long, hairy, placed near the middle of the
pedicel or above or below; sepals purplish blue, ovate, strigose without,
the upper 15-18 mm. by 7-10 mm., obtuse, the spur sometimes coiled in
an almost complete circle, ca. 2 cm. long, 5 mm. wide at base; lateral
sepals oblong, 20 mm. by 8 mm.; lower sepals ca. 22 mm. long, 10 mm.
wide, obtuse; petals blue, the upper laminae straight, 9-11 mm. long, en-
tire to emarginate, the spur 16-20 mm. long; lower laminae elliptic-obo-
vate, ciliate, bearded at base, 7-11 mm. long, shallowly to deeply bifid,
the claw 5-6 mm. long, not appendaged; stamens 6~7 mm. long, glabrous;
anthers blue, linear, 2 mm. long; follicles 3, hairy, 15-30 mm. long; seeds
brown, 1.3 mm. long, squamate.

Type: Szechwan, west side of Mt. — near Dschunglien, Handel-
eae 4656 (w), not seen; isotype (E),s

DISTRIBUTION. Occurring at elevations of "2800 to 4800 m., Szechwan
and Yunnan.

EXAMPLES. Szechwan: Handel-Mazzetti 7302 (GH, NY); Rock 18015 (us),
16694 (GH, NY, US), 16637 (US), 24166 (GH, UC), 24589 (NY, UC), 16760 (P,
US), 23979 (£, GH, NY), 16922 (NY, Be), 16929 (GH, UC, US), 18294 (US), 13979
(GH, Ny, UC), 16814 (F, US), 16398 (GH, US), 16694 (GH, NY, US); Forrest
22184 (us); Kingdon Ward 4375 (£). Yunnan: Rock 24771a (cH, K, NY, UC c),

244. Delphinium tongolense Franchet, Bull. Soc. Philom. Paris VIII.
5: 166. 1893. Fig. 19, K, L.

242 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Perennial from woody base; stems 1 or more, to 1 m. tall, glabrous or
with long white or yellowish hairs, (some with enlarged base), few branched
above, leafy almost to the summit; petioles slender, the lower 2~3 dm.
Jong, not much widened at base, the upper shorter; blades palmately 3-
or 5-lobed to middle or below, the lobes cuneate-obovate, coarsely rather
few toothed or lobed, blade as a whole 5-10 cm. wide and long, mostly
subglabrous; inflorescence largely paniculate, the branches few, lax, few to
several flowered; lower bracts foliaceous, trifid, to 2.5 cm. long, the upper
entire, linear or broader, 5—15 mm. long; pedicels slender, ascending, sub-
glabrous to spreading-pubescent, 2—5 cm. long; bracteoles linear-setaceous,
4-7 mm. long, near the middle of the pedicel; flowers deep to pale blue;
upper sepal ovate, 15-17 mm. long, 8-10 mm. wide, pubescent, obtuse, the
spur 2.5 cm. long, 4 mm. wide at base, pointed; lateral sepals 13 mm. by
10 mm., rhombic-ovate, rounded at ends; lower sepals 15 mm. by 10 mm.,
more narrowed at base; petals bluish, the upper laminae oblique, 8 mm.
long, entire to bidentate, the curved spur 18-20 mm. long; lower laminae
bearded, 5-7 mm. long, with 2 divergent lance-oblong lobes 3.5—4 mm
long, claw broad, not appendaged, 5 mm. long; stamens 6—7 mm. long,
glabrous; anthers dark, 1.6 mm. long; follicles 3, subglabrous, divergent,
1-2 cm. long, the styles an additional 2 mm.; seeds many, angular, cov-
ered with lamellose papillae

Type: Western China, Szechwan, Tongolo, valley of Olong-chen, R. P.
Soulié 396 (Pp); seen. Soulié 2383 is in a type cover at (K

Rance. Western China and adjacent Tibet at perhaps 3500 to 4500 m.

Examptes. Szechwan: E. H. Wilson 3089 (x, P); ies 2383 (K, NY, P);

Stevens 454 (¥); McLaren’s collectors 258 (£, UC), 206 (£). Yunnan: Big oo
4600 (Pp). Sikang: H. Smith 11340 (a), 11339 (mo, ) 11594 (a). Tibet:
Batang, Soulie 3890 (P), 3889 (Pp), 3888 (P), 3050 (P).

REPRESENTATIVE EXSICCATAE

(Unfortunately many important specimens, especially of early — in Siberia,
ve is eng mb ne To cite Regel collections, for instance, is almost useless; often

ral made in the same year represent different speci ee .
is not definitive.) ae sy “Reg es

Abolin 8193, D. confusum Arnold 104, 213, D. brunonianum: 12,
Aitchison 957, D. brunonianum: 1004, kamaonense —

serie 671, semibarbatum; 862, Artemjev 4266, D. duhmbergii

Suave; 559, 575, uncinatum Atki mat

, tkin j

Alboff 194, 195, 196, 182, D. pyramida- aria yee nN tag

tum; 190, schmalhausenii; 15, spe- oe

um

ciost

Alexeenko 2620, D. crispulum Bacon, Eliz. 73, D. suav

Ali, Shaukat 53, 54, D. brunonianum Bailey, F. M. 100, D. v um
Anderson, T. 345, D. viscosum Balansa 859, D. a. 1352,
Appleton in 1906, D. dasyanthum ; formosum; 1353, longebracteatum;

3862, suave 728, peregrinum; 858, venulosum

1968 |

Balls, E. K. 496, 1975, D. formosum;
1207, 2406, peregrinum
Balls & Gourlay 1977, 1992, D. dasy-
Stachyum; 1977a, longebracteatum
Banerji 1231, D. vestitum
Barbey 295, D. fissum ssp. anatolicum
Beauvais 409, D. delavayi; 1074, mo-
soynense
ae Al 20. Dd.
3, D. ochroleucum
eae 2253, D. denudatum
Bodinier & Ducloux 370, ee taliense
Bohnhof 163, = maackian
4) Sag June , D. cinereum; in
ihaburense April 1846, pere-

turkmemum; in

cialis
Bor in rei: vey cashmerianur m

6062, aquilegifolium; 1529, 2674,
venulosum
Bourgeau 9, D. peregrinum
Brandis 731, D. vestitum
E. 804, D. pretties:

a Scifolium var.
; 10125, cates e var.

Burtt & Nairn 586, D. suave
Bushnell 470, D. maackianum

Callery 6, 51, D. anthriscifolium var.
calleryi

Calvert & Zohrab 1347, D. albiflorum

Capus 50, 51, ett and June 12, 1886,
D. ochtolen

Carmichael ees "D. chefoense

Cavalierie 2344, 2957, D. anthrisci-
folium; 2169, 2822, anthriscifolium
var, calleryi: 2095, 4486, eriostylum;
7871, georgei

Cave, G. H. 6, D. caeruleum; 177, and
In 1910, in 1917, in 1919, D. vis-
cosum

Chaffanjon 804, D. biternatum
hand, T. R. 72A, 588, 406, 887, D.
brunonianum; 576, cashmerianum;
881, densiflorum; 3449, 7192, 31250,
stapeliosum ; 1190, vestitum

ee 136, 467, 1104, D. grandi-

re & Jen 6099, D. glabricaule

MUNZ, ASIAN SPECIES OF DELPHINIUM

243

Chapman, F. S. 35, D. caeruleum; 167,
201, gyalanum

Chen, F. H. 341, D. maackianum

Chiao 1758, D. hirticaule

Chien 5877, D. anthriscifolium var.
calleryi

Ching, R. C. 1363, 8442, D. anthris-
cifolium var. calleryi; 849, campylo-
centrum; 21392, 21616, 30509, cera-
tophorum; 21456, ey 21418,
glabricaule ; 22022 30535, hirti-

861, trichophorum:

Chu, K. a 3246, D. potaninii; 3606,
Scapos
ae re anthriscifolium var. cal-

ler
Clarke, C. B. 29095B,. D. roylet;
29506, pyramidale; 16117, 6168,
16354, 16381, 38970, 40301, 44861,

burmaense;
5404A, vestitum
Colom 282, D. altissim
Conrath 1889, D. shea
Cooper, R. E. 760,
5309, 5648, cashmerianum:
2294, cooperi; 4322, ludlowii; 4791,
vestitum ;

u
Cooper & Bulley 2008, 2556, D. altis-
simum;, 4114, 4223, scabriflorum
Coventry 226, D. pyramidale
Crookshank, Helen 324, 366, D. cen-
keine
Cunni
sah es

R. potaninii;
3S. : sc el

Davis, P. H. 30593, 31305, D. albi-
florum; 32562, buschianum in part;
22801, 22593, 23061, 23325, 23349,

23327, 32443, ;
31669, 31999, dasystachyum ; 3

davisii; 13140, 14703, fissum ssp.
anatolicum; 21122, 21211, formo-
sum; 38373, ilgazense; 4608 &
4608a, 4894, ithaburense; 31496,
hurdicum; 33853, 33907, ochroleu-

244 JOURNAL OF THE ARNOLD ARBORETUM

cum; 4801, 9913, 13473, —
15421, 31567, 38840, peregrinum
32782, 32788, 32813, oenulosum:
35701, virgatum

Delavay, Abbé 967, 1845, D. cerato-
phorum; 22, 74, 1846, 1847, 4109,
delavayi; 1676, 2676, hamatum,
2484, likiangense;, 8717, mosoyense ,
993, 1844, 4220, pycnocentrum; 9,
taliense; 967 bis, 6515, 6670, yun-

nanense
Dhwoj, Lall 49, 0253, D. altissimum ;
556, caeruleum; 0191, 0641, drepan-

253, a 92, 128, 0258

“

oo. 369, D. brunonianum; 9052,

d’Orléans, Prince, = ect 7-29-
1890, D. tatsien

Dorsett - Dorsett ae. 3367, D. gran-
difloru

Dorsett & Morse 7051, D. grandiflo-

ru

Douglas, W. O. 161, 236, D. aquilegifo-
nee ea ithaburense

Drum d, J. R. distributions 1313,
4341, "4346, 4348, D. brunonianum ;
8328, denudatum; 14811, —
4345, 4349, 6285, vestitu

Ducloux 597, D. c sdvalbbbaies: ats
delavayi; 579, hirticaule; 3875, m
soynense ; 5153, potaninii: 596, 4
sienense; 4600, tongolense; 370,
2874, 5866, 7329, yunnanense

Duthie, J. F. 745, 5301, in 1896, 7-25-
1899, 7-24-1899, 13370, 13751,
13921, D. brunonianum; 794, 3804,
5302, cashmerianum; 2677, 5303,

l ; 5, 5299, kamao-

nense; 13490, 14123, pyramidale;

13440, 14091, roylei; 5304, 19119,

vestitum

Egorova, Tzelev, & Czerepanov 418,
D. pallidiflorum
Emme, L. 451, D. dasyanthum

[ VoL. 49

a ne IAG Vey 8 anthriscifolium
calleryi & var. savatieri
stich 141, eriostylum
Eyerdam, W. 2, 7-27-28, 7-14-28, 7-
15-28, 8-25-28, D. brachycentrum

Faber, 725, D. anthriscifolium
Falconer 66, D. onianum;, 65,
cashmerianum,; 60, ae ae
im

orthocentrum ; 41,
4038, sungpanense;

Farges 1146, 1440, 1441, D. anthrisci-
folium; 630 bis, 630 ter, hirticaule ;
630, potaninii

Farrer 159, ec georget;
micropeta

Farrer & coca 242, 243, D. albo-
caerulem; 237, beesianum; 578, 611,
grandiflorum ; 253, monanthum: 710,

1156, 1867,

pylzowii; 798, 800, 803, trichopho-
rum
Faurie 26, D. maackianum

Fedtschenko in 1901, D. dasyanthum

Feng, K . 2395, D. beesianum;
1982, 2001, bulleyanum; 2714,
chefoense; 217, 97, delavayt;

2742, 2272, georgei; 2207, mairet,
2514, mosoynense; 1836, 2115, po-
gonanthum: 2257, Te 2841,
tenii; 1726, umbrosum: 89, yun-
nanense

Field & Lazar 730, 850, D. peregrinum

Fleming 5-23-51, D. uncinatum

Forrest, G. 6694, 11276, 6364, 14857,
18961, oe 20572, 22967, 22577,
D. beesianum; 2694, 6556, 11130,
Silavetia: 6403, 6568, 2763, 10967,
ceratophorum; 3035, 6690, 11363,
15173, 15198, 16942, 22471, 22483,

28587, 2604, 6319, delavayi;
georget; 15555, hamatum;
6540, 2903, 11360, 11052, 11203,
16949, 15322, 15170, 16949, likiang-
ensé: 221, 27516, 27225, 25091,
micropetalum; 20780, monanthum,

1968 |

2866, 6533, 15233, mosoynense;
6538, 10968, 11449, 15112, 16951,
23185, 22378, arp
14929, 17187, 19031, 30044, 22184,
spirocentrum; 6494, 65
euse= 6083, 3030, °11305,. 16023.
15194, 22347, 22603, 22966, tricho-
phorum: 14665, 18962, 20060,
20979, tsarongense; 25052, umbro-
sum; 2977, 7216, 15536, yunnanense

F ee 328, D. vanense

Fu, K. N. 1146, D. acuminatissimum

Furse 3654, 3785, D. albiflorum; 2627,
aquilegifolium; 3061, cyphoplec-
trum; 3732, dasystachyum; 3404,
macrostachyum

Furse & Synge 329, 330, D. aquilegi-
folium; 784, laxiusculum

Gaillardot 52, 367, D. peregrinum

Garrett 452, D. siamense

Giles 6-24-86, D. kohatense

Gill 151, D. brunonianum

Gillett, J. B. 9501, 12473, D. micran-
thum; 11782, pallidiflorum

Gillett & Rawi 7223, 7404, 11206, D.

icranthum
Gilliatt-Smith 2514, D. crispulum
Giraldi 4819, 4820, 4823, 4825, 4826,
D. giraldii
Gontchazov 8-22-1928, D. dasyanthum
Gould, B. J. 1412, D. glaciale; 508,
Ringianum; 1300, viscosum; 1598,

wardii
Griffith 45, D. centeteroides; 43, ko-
hatense ; 42A, 461, 827, D. stapelio-

Grossheim, A. 305, D. crispulum: 37,
ochroleucum
Grossheim & Schischkin 333, D. flex-

m
Guest, E. V. 7-28-33, D. kurdicum
Guest & Ludlow-Hewitt 2691, D.

micranthum
Gurvitsch in 1937, D. laxiusculum
Guthrie, J. in 1947, D. gyalanum

Haines, R. W. 416, D. kurdicum; 513,
macrostachyum; 1320, micranthum;
712, quercetorum

MUNZ, ASIAN SPECIES OF DELPHINIUM

245

Hanbury-Tracy 143, D. brunonianum;
195, pseudograndiflorum

Handel-Mazzetti 5495, D. autumnale;
11802, anthriscifolium; 8128, beesi-
anum; 4105, coleopodum; 9712,
micropetalum- 5103, mosoynense;
4576, 7819, 7424, ‘ pogonanthum:
4656, 7302, spirocentrum; 53515,
irichophorum: 7934, tsarongense;
7809, umbrosum

Haradjian 552, 571, 2537, D. fissum
ssp. anatolicum: 1525, 2158, pere-
grinum

Harriss, S$. A. 15837, D. suave

Hansekaceht, April 1868, D. cypho-
plectrum; in as bse
June 1867, auercetorum: July 1868
saniculifoli:

Hay 373, D. psc es

Hazara, Kazmi
pyramidale

Heideman 5-27-1932, D. ~~

Heideman & Aliev 7-2~1932, D.

1266,.D.

szowitsianum

Hemsley 7, D. anthriscifolium var.
calleryi

Hen 3 1966, 5862, D
thriscifolium; 7263, anthriscifolium
var. calleryi; 200, anthriscifolium

potanini; 7448, trifoliolatum: 9272,
9272A, 13440, yunnanens
Hingston 367, 408, D. gyalanum
Hohenacker in 1834, in 1835, D. lax-
iusculum; in 1838, ochroleucum in
si nik ia July 1838, szowit-

Hooker, oh D. 2293, D. glacial

Horner, S. 152, 153, D. cyphoplectrum

Hosie in 1904, me trichophorum; in
1904, ¢sarongen.

Hsia 3159, 2065, D. grandiflorum

Huggins, P. 39, D. densiflorum

Hultén 733, 638, D. brachycentrum

Ikonnikov-Galitzky 365, 746, 3335,
3428, D. cheilanthum; 785, dissec-
tum: 2915, elatum; 210, 314a, 802,
grandiflorum; 82, triste

Jacquemont 668, 2025, D. brunonia-

246

num; 720, 1896, — 671,
1005, cashmerianum: 721, , de-
nudatum: 673/791, 791, a ‘roy
lei, 672, uncinatum, 2134, vestitum
Jain & Akbar 2197, D. centeteroides
Jaschke 2495, D. lacostei

Kabir 14811, D. aquilegifolium
Karelin & Kiriloff 1169, D. dasyan-
— 1164, duhmbergu; 1168,

elat
Karjagin & Jsoev 8-25-1933, D. foe-
idum

th

Karjagin & FF 150, D. buschianum

Karo, F. 1 BD: cheilanthum : 207,
iit

Kass & Nuri 27607, D. quercetorum

Keng 1311, D. anthriscifolium var.
calleryi; 880, anthriscifolium var.
age

Kerr, A. ¥..G. 337, 4454, 6612, D.
siamense

Kingdon-Ward; see War

Kitamura 8-2-1955, D. nuristamicum

18257, aquilegifolium; 1000A, 1233,
2400d, 2407, 2508, 2804, 2835a, 3026,
5562, 6516, 6622a, 6703, 6841, 7006,
0551, 10112, brunonianum: A38,
1277, 850, 588, 1313, 2787, 2803a,
2973, 5803, 6026, 6927, 10028,
10041, cashmerianum : 21082, densi
florum; 1848, 3163, 4841, 4638,
4963, 8344, 8659, 8328, 21113, denu-
datum: 4757, 4921, itis koelzii;
18164, lanigerum: minjan-
ense; 18015, palidiforum: 9313,
9261,

pum;
plectrum; 18110a, tuberosum; 3077,
21952, vestitum

Kolenati 1590, D. ochroleucum

Komarov, V. L 6-26-93, D. biterna-
tum; 7-18-1908, 3896, D. Srieny:
centrum; 6-26-92 dasyanthum
670, prandiflorum: 671, maackianum

Korolkov 76, D. dasyanthu

Korshinsky 80, D. ——

Koslov, 240, D. grandifloru

Kotschy 239, 239b, D. aiiideaiinieat

JOURNAL OF THE ARNOLD ARBORETUM

[voL. 49

153, 393a, 400, May 1842, cypho-
plectrum; "427, 503, kurdicum; 430,

lanigerum; 7. 54 ; longebracteatum:
, 610, micranthum; 1, 8-1-41,
peregrinum; 536, saniculifolium

i ae hakewics 7- 22-78, D. dasyanthum
Kusnezow 875, 885, D. elatum

Lace, J. H. 3862, D. centeteroides;
164, 734, 1523, 1924, D. denuda-
tum; 6012, lacei; 2094, lacostet;
467, ey roylei; 3326, 3351, un-
cinat

ern s D. molle

Ladygin, Oct. 1900, D. trichophorum

Lamont, J. 1065, D. —

Lance 18, D. brunonianum 21, -

merianum

Ledebour 809, D. grandiflorum

Legendre 1563, D. lancisepalum

Lepcha 269, D. caeruleum; 865, micro-
petalum

Li, C. F. 10707, D. grandiflorum

Li, H. 74095, D.. campylocentrum,
78175, lancisepalum; 78358, pseudo-
campylocentru m; pseudo-
grandiflorum ; 74723, souliei: 78699,
trichophorum

Li, K. F. 63169, D. trifoliolatum

Li & Chu 7003, D. tianshanicum

Licent, R. P. 1305, 1999, 7665, D.
grandiflorum; 2760, 4544, henryt;
4279, 7719, polyanthum; 4686, pul-
cherrimum; 5655, cary tiie

Ling, K. 9143, D. grondiflo ru

Liou, K. 1405, D. pt Pola
5093, henryi

Liou, T. N. & P. C. Tsoong 986, D.
taipaicum

Lipschitz 685, D. batalinii; 718, min-
janense

Liu, > oe 1968, 12689, D. grandi-

flor
eras 7685, 2.

batalinii; 1783,
duhmbergii; 8-10-1896, propin-
quum; 1887, semiclavatum; 3070,

5-8-1912, turkmemum

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 247

Litchfield in 1890, D. densiflorum
Litwinow 490, 989, D. grandiflorum;
26, 3013, maackianum; 443, 444,

semibarbatum; 447, 448, turkme-
mum

Loftus 5-3-1852, D. BBA ts

Lomakin in 1895, D. foetid

Lowndes, D. 1283, 1323, D. Brawidé.
num; 1229, 1324, kamaonense;
1347, 1505, nepalense

Ludlow, F. 433, 838, D. brunonianum ;
532, candelabrum : 864, cas agi!
mune 715, 181. hancicens
poltaratskii; x aati Ae:
rum; 868, 7

Ludlow & Sherriff 1987, acuminatissi-
mum; 523, 6899, D. albocaeruleum ;
298, 349 6, 3558, altissimum; 5,
6912, 8504, 8930, 8561, 9010, 9925,
brunonianum: 873, 887, 1076,
19686, bhutanicum; 779, 1968, 1977,
2400, 9751, 9952, 11067, caeruleum ;
1881, 2814, 7914, cashmerianum;

an
578, cooperi; 7671, 8106, 9106,
denudatum; 16786, 17128, 19724,
glaciale; 9073, 9814, 11114, grandi-
florum; 154, 528, 2421, 2461, 1942,
8781, 8780, 2/77, -° 98935: 11108;
gyalanum: 2424, hilicoating 527,
855, kingianum; 894, 470, 368, 1008,
16972, 19485, 20978, 21303, lud-
lowii; 3537, muscosum; 17128,
19724, nepalense - 687, 499, 2595,
pseudograndiflorum: 1958,..-1977,
pulcherrimum; pyramidale ;
7560, 8213, 8238, 9376, roylei; 219,
3342, scabriflorum; 5833, sherriffii;
474, 2563, 2096, 2721, 864, 2515,
9958, 10069, 17151, 19702, 19770,
19787, 21057, viscosum
Ludlow, Sherriff & Elliot 15683, D.
pseudograndiflorum
Ludlow, Sherriff & Hicks 17476,
caeruleum; 16825, 17102, 19328,

Ludlow, Sherriff & Taylor 6921, D. al-
bocaeruleum: 6379, beesianum;
5329, 5320A, 5824, 5701, 6250,
4849, gyalanum; 5397, 6920, muli-
ense; 5162, 6093, 13920, pseundo-
grandiflorum : 6066, 6403, viscosum

Ludlow, Stainton & Williams 15621, D.
ikiangense

Lyon, Bowes 25, D. aquilegifolium;
1083, brunonianum

Maack, R. 673, D. cheilanthum
Macmillan, H. F. 7, 8, 9, D. tubero-
sum

Maire, E. E. 2292, 3292, 6367, 7419,
D. delavayi; 2128, D. georgei; 2669,

dbstead va ati 389, 1196, yunnan-

Maciley 211, D. elatum

Manissadjian 6 675, a Perna ssp. ana-
tolicum; 26, pereg

Martin & Bodinier ds D. yunnan-

ense
Matveeva, E. 555, D. laxiusculum
Maydell, G. 8-8-1869, D. brachycen-

trum
McLarens collectors 159, D. bulleya-

tatsienense; 206, 258, tongolense ;
AB79, F68, V88, yunnanense

McNeill, 731, D. aigcoesaet

Melville, 43, D. bur

Merton 3935, - doen 3888,
semibarbatu

— a D. duhmbergii; 1304,
villos

nee - Chang 797, D. anthriscifo-
lium

Metz 52, D. vestitum

Meyer, C. A. 1727, D. caucasicum

Meyer, F. N. 1057, D. grandiflorum ;
626, semibarbatum; 532, turkme-
mum

Meyer, F. S. B4864, D. ithaburense ;
99, virgatum

Mikelson, A. 2279, D. biternatum

Mozheeba 239, D. elatum

Mussot (Moussot) 27, D. souliei; 28,
29, trichophorum

248 JOURNAL OF THE ARNOLD ARBORETUM

Nasir, E. 28459, D. centeteroides
Nath, M. 387, 437, D. vestitum

Norton, J. 369, D. nortonii

Ohwi, J. 10662, D. grandiflorum;
2808, maackianum
Osmaston 75, D. roylei

Parmanand, N. 588a, D. brunonianum ;
1139a, vestitum
Pavillon 1843, D. formosum
Pavlov 336, D. dissectum
Pei 2646, D. anthriscifolium var.
i

Persnin 170, D. cinereum

Pichler in 1882, D. tuberosum

Pinard in 1843, D. cinereum

Pinfold 164, D. denudatum; 329,
pyramidale: 210, 224, roylei

Polunin, O. 2599, 6250, D. brunonia-
num; 56/175, 56/597, 56/647, cash-
merianum, 1440, 1247, densiflorum;
1905, 1790, 1949, glacicle: 555.
1413, 1489, 1778, Sepataane: 872,
1576, kamaonense 208, lacostei;
56/699, pein. 56/330, 56/602,
roylei; 56/283, 56/803, vestitum

Polunin, Sykes & Williams 3518, D.
cooperi; 264, 2503, 4885, 5128, 5290,
himalayae; 3533, nepalense; 2629,
5120, pyramidale; 386, 5025, 5441,
vestitum; 1907, 3916, williamsii

5-20-71, 6-30-82, 8-22-84, D.
peregrinum

Potanin in 1885, D. Po tie sgs
pylzowi and sutchuenens

— 4 Ratcliffe ie. 265, D. pere-
grint

Pratt 4, D. pachycentrum; 344, po-

tan.
EASON 43, D. macropogon
Przewalski in 1872, D. albocaeruleum,
pylzowiit and sparsiflorum: in 1873
and 1874, przewalskii; in 1880,
pylzowii and trichophorum
Ptazycki 373, D. elatum
Purdom 126, D. grandiflorum; 142
trichophorum

>

[ VoL. 49

Radde, G. 212, D.
semibarbatum

Rajkova 129, D. dasyanthum

Ram Bahsli 14811, D. suave

Ram Bahsh or Bakeh 8351, D. cash-
merianum; 4342, 4344, roylet

Rami, Hoshen & Nuri 29481, D. mac-
rostachyum

Ratcliffe 759, D. venulosum

Rawi, 24731, D. micranthum; 12213,
pallidiflorum; 22200, 22706, 23431,
quercetorum

Rawi & Serhang 20208, D. foetidum

Read 746, D. grandiflorum

Sa bas 3838, D. aquilegifolium;

1416, biternatum: 6093a, elbur-

gorganicum; 3364,
saniculifolium; 1698, 4934, semi-
barbatum; 6083, ursinum

hee A. June, 1883, D. biternatum;

in 1882, dasyanthum; 286
tod May, 1876, longipedunculatum:;
Aug. 1876 and 5-17-1880, semibar-
batum

Ren 6290, D. ceratophorum

Ribu & Rhomov 5234, D. glaciale

Ridley 20, D. grandiflorum

Robertson, W. A. 15, D. burmaense

Rock, J. 13194, 14459, 14467, 14501,
14505, D. albocaeruleum; 5421,
7763, 10776, 10778, 10817, 11276,
13398, 13092, 14518, 14520, 14502,
14523, 14640, 17248, 18810, 20204,
22852, 24738, 24914?, beesianum;
6082, 6085, 5696, 6091, 9669, 9755,
9766, 9769, 9924, 10797, 16938, bul-
leyanum; 4746, 5006, 5669, 5696,
5888, 5774, 6948, 10737, 17321,
22852, 23080, 23992, ceratophorum;
6092, 6110, 7761, 7765, 10669, 10764,
10633, 10777, 11435, 11436, 11676,
24738, 24945, coleopodum; 4404,
4849, 5030, 5298, 5768, 5799, 5936,
6093, 6149, 6177, 9474, 9914, 5786,
5828, 10634, 11483, 18324, 18519,
24190, delavayi; 10539, georget;
16830, 18121, glabricaule; 13361,
grandiflorum; 16884, 22852 in part,
3457, hamatum; 6305, 17317,
22373, hirticaule; 13136, leptopo-
gon; 5242, 6111, 6423, 9867, 10409,
10665, 10792, 11413, 11435, 15242,

crispulum; 545,

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 249

17332, 24914, 24920, 25330, likiang-
ense; 18327, 23243, 24272, maximo-
wiczit; 22373, 22401, 23086, 24190,
micropetalum:: 18317, mitzugense ;
16916, muliense; 13024, 13185,
monanthum ; 13730, 17338, mosoy-
nense; 16884, pachycentrum: 16743,
23992, potaninii; 5906, 6171, 6193,
10747, 14538, pseudograndifioras:.
14460, pulcherrimum; 16886, 18316,
pumilum; 6135, 5787, 10670, 10813,
10657, 17319, 17338, pycnocentro-
ides; 13023, 13041, 13086, 13088,
13093, 14504, 14625, 14482, 14483,
14513, 14641, pylzowii: 13191
sparsiflorum: 9755, 9766, 9669,
16760, 16345, 16767, 16814, 16797,
16398, 16637, 16694, 18015, 18294

trum; 6292, 7770, 10686, 11485,
11545, 18485, 23243, tatsienense;
18205, 23457, tenii; 13154, 13193,
14547, tsoongii; 10779, 10860, 11696,
13740, 16824, 18237, 23090, 23444,
24938, trichophorum; 22267, wardit;
16930, wrightii; 6551, 6902, yunna-

nense
Roerich 60 J, 336 I, D. grandiflorum
Russanov 119, D. inopinatum

Sampson 10125, D. ra
pon rs, D. F. 8-2 ordha-

rai 11551, 11552, 13868, 13869,
. malabaricum
Sarchen & Bajkan 1169, D. dasyan-
thum
Schelkownikow 65, D. foetidum
Schindler 78, D. grandiflorum
Schlagintweit 1711, 1985, 6306, 9004,
. brunonianum; 4824, 5171, 5484,
9007, cashmerianum; 9180, 9552,

Schneider, C. 2382, D. beesianum;
2245, 2983, 3459, ceratophorum;
2143, 2285, 3576, delavayi; 2928,
3251, 3774, likiangense; 2430, mati-
rei; 2278, 3250, pseudograndiflorum ;
2278, pycnocentroides; 3249, spiro-
centrum; 2121, 1510, tenii

Schock 276, D. mosoynense

Seringe 1489c, D. vestitum

Sharma, H. N. £295, D. brunonianum;
E482, glaciale; 117, himalayae; E98,
pyramidale; 55, vestitum; 54, 54/
4, E140, viscosum

Sherriff 7310, D. denudatum

Siddiqui & Rahman 26868, D. suave;
26865, kohatense

Sintenis 4883 D. albiflorum; 500, 850,

um; 473, 1207, peregrinum; 383,
semibarbatum; 1672, 1672b, turk-
memum; 4627, venulosum
Skorobogdtko 302, D. biternatum
Smith, H. 2985, 4243, 4326, 11712,
12619, 12515, 11939, 12421, D. bees-
ianum; 4138, campylocentrum,
5741, chefoense; 173, 468, 11063,
4660, 6526, 7139, 8150, grandiflo-

4421, trichophorum; 11285, i132,
11459, tsarongense
Smith, W. W. 4128, D. viscosum
a & Cave 1935, 2131, D. caeru-

leu

suai 292, D. biternatum

Sokolov 97, D. grandiflorum

Soong, T. P. 38839, 38912, 38915, D.
pseudotongolense

Sorokin 1357, D. speciosum

Soulié, R. P. 3040, 3041, "3042, 3042
bis, 3043, 3898, 3899, 3907, D. ba-

cropetalum; 2387, 2389, mosoyn-
ense; 24, 2379, pachycentrum; 3903,
3911, 3914, pylzowii; 364, 399, 2388,
souliei; 1097, spirocentrum ; 25, 438,
525, 2027, 2381, 2382, 2481, 2489,

250

3529, tatsienense; 3049, 3893, 3894,
3044, thibeticum; 396, 2383, 3889,
3890, 3880, 3050, tongolense; 23, 64,
359, 2029, 2384, 2386, 3038, 3909,
trichophorum

Sovetkina & Czansova 4267, D. semi-
barbatum

Stainton 2846, D. brunonianum; 1 142,

1144, drepanocentrum; 1025, glaci-

S

2
ense; 1162, SS aia ba 1327, 1650,
1709, VISCOS
Stainton, ie & Williams 3707, D.

3029 9, 8058, viscosum
Stapf 2300, 2793, ee cole area
Steane 60, D. ves
Steven 8711, D. iced ucum
Stevens, H. 457, D. delavayi; 88, 458,
pseudograndifiorum: 89, 114, tatsien-
nse; 454, tongolense
Steward 2071, D. anthriscifolium var.
calleryi
Stewart, J. L. = ” D. kohatense
Stewart. H. M. 7-1-07, D. gyalanum
Stewart, R. R. 9819a, 9849, 20176,
20698, 22132, 22294, D. brunonian.

53268, 8079, 5714, 9225. TRIT 16,
18357, 21882, 28781, 29167, pyra-
midale ; 5694, 6530, 7272, 8102a,
10546, 19239, 19586, 19693. roylei:
579, suave; 7002, 1268 oO, ef tee,
uncinatum

Stewart & Rahman 252295 D.
gifolium; 25125, tenuipes

Stocks 994, D. uncinatum

aquile-

vestitum

JOURNAL OF THE ARNOLD ARBORETUM

[VvoL. 49

Stukow 1358, D. grandiflorum

San, © i 3765. dD. anthriscifolium
var. calleryi; ninit

Swann R

Swigh 11, D. denudatum

Szowitz 168, D. laxiusculum; 439, szo-
witsianum

Tang ae cen ie Slate
Ten, S. delavayi; 1292,
sania - sens: 134, yunnan-

53, D. anthriscifolium var.
Savatieri ; a cabhrisetok ia

Timins 203, D. pyramidale

Tolmatchew, 174, D. elatum

Toppin 769, D. vestitum

Tsai, H. T. 53738, 59749, D. delavayi;
57754, hirticaule; 57615, pogonan-
thum; — 56240, tenii; 526735,
umbrosu

Tsiang, Y. 8571, DD ese eget e!

Tu, T. H. 4366, D. sa

Purcrsncwicn 929, D. pe ]

Tyson, J. B. 95, D. kamaonense

Uvarov 29, D. peregrinum: 98, venu-
losum

Vnorrili 43, D. longipedunculatum

Wallich 1832, D. brunonianum: 4717,

sigas 4716, pyramidale; 4715,
& B, D. beetles

Wain 1. 7 and July, 1904,
D. Claim ae 1904, pseudo-
grandifiloru

Wang, C. W. 20450, D. sein
var. calleryi; 65706, cerato

ides; 71376, delavayi: oe hat
- matum : 71576, mairei; 69805, poly-
anthum: 64594, 68543, 69805, 68635,
68390, 68590, Spirocentrum; 66226

] 69397, 647

“

21767, D. laxicymo
ies, H. 41742, D. bis ante.

Wace +. F. 9075. D. ee
Wang & Wen 581 Wy ip
Ward, F. Kingdon- 6123, gi te "12317,

1968 |

12224, D. albocaeruleum; 14015, al-
eesianum; 10900,

811, 4846, bul-
leyanum; 3351, 9819, 10115, 13170,
14308, 18121, 22505, burmaense;

10621,
a 12402, pseudograndiflorum;
4375, 4604, 4617,
trum; 4677, 4932, 4956, trichophor-
um; 5390, tsarongense;
12337, 14329, viscosum

Watt, G. 612 af denudatum

Webster & Naser 6100, 6192, 6560, D.

>
brunonianum ; 6524, ee

4690, davidii:

3095, 3097, 4702, potaninii; 3088,
sinovitifolium; 3092, 3092a, ¥,
souliei; 4691, 3094?, tanguticum;

enudatum
Wollaston 18, D. caeruleum

MUNZ, ASIAN SPECIES OF DELPHINIUM

251

Woronow, G. 195, D. laxiusculum;
990, paednclne
2, D. pumilum
wecen Boe D. puniceum

Yang, K. H. 58311, D. anthriscifolium;
65383, henryi; 58041, pseudocampy-
locentrum

Yaroshenka & ws 7-23-1945,
D. szowitsianu

Veo, me 1282 oe D. aconitioides;

13140, 13649, beesianum; 15544,
bulleyanum: 12809, chenii: 12548,
13824, 13543, 22461, delavayi:
14008, dolichocentroides: 14490,

4022, hirticaule: 13101, hut;

14642, trichophorum:; 979, umbro-
sum; 7797, wrightii; 12490, 10547,
13852, 14613, yuanum

Zimmerman 192, D. chefoense; 561,
hirticaule

INDEX TO NAMES

This is an alphabetical list of names that have been proposed for the Asian
species of Delphinium. Subspecific and varietal epithets are indented under the

appropriate specific epithet
paper. Names

t. The numbers refer to the species number in this
rinted in italics are names of taxa reduced to synonymy here.

Species referred to Consolida have been omitted.

aconitioides Chen, 58
acuminatissimum W. T. Wang, 141

var. pumilum Huth, 81

albomarginatum Chen, 191
albomarginatum Simonova, 11
alpinum Huth, 200

var. ajanense Huth, 200

var. productum Huth, 200
altaicum Nevski, 142
altissimum Wall., 143

ssp. drepanocentrum Brihl, 151

ssp. wallichii Brithl & King, 143

var. nipalensis Brihl, 143

252

var. phallutensis Brihl, 143
se ya Wall., 14
r. drepanocentrum Brihl ex Huth,

var. ais Brihl ex Huth, 143
amani Post ex Hu 2
amoenum Steven ex DC., 163
anthriscifolium Hance, ;

var. anthriscifolium,

var. callerii (Franchet) Finet &
Gagnep., 9

var. majus Pamp., 9

var. ramosum Pam

var. savatieri (Franchet) Munz, 9
apetalum Huth, is Aconitum

apetalum Fedts sch
aquilegifolium (Boiss. ) Bornm., 12

bf

var. ean ata Botnm: 12
var. violaceum Bornm., 12
araraticum Busch, 91

©
>
3
5
awd
=
™
3
=
=
rg
-
=
do

00
aureopilosum Chowdh. ex Mukerjee,
iii

autumnale Hand. eek 82
batalinii Huth, 1
batangense he & Gagnep., 83
beesianum W. W. Smit h, 84
forma Galea (W. W. Smith) W.
. Wan

ng,
var. beesianum f. calcicola W. T.
Wan
var. latisectum W. T. Wang. 84

. radiatifolium Hand.-Mazz., 84
var. beeing: (Hand.-Mazz.) W.
T. Wa

amosum ie T. Wang, 84

Bheticaer sles 85

gistigr es Huth, 14
var. leiocarpum Freyn & Sint., 48

bithynicum Davis, 6

boissieri pie me

bonatii Lévl.

bonvalotii Franche, 240

bovei Deca

brachy sneer Lede, 144

bracteosum Som. & Levier, 188
var. albidum Som. & Levier, 229
var. macranthum Som. & Levier, 229

JOURNAL OF THE ARNOLD ARBORETUM

[voL. 49

brunonianum Royle, 59
var. aitchisonit Huth, 59
var. densa Maxim., 63
var. jacquemontianum Huth, 59
var. schlagintweitii Huth, 59

89
buschianum Grossheim, 190
caerulescens Freyn ex Stapf, 53
caeruleum Jacquem. ex Camb., 86
var. caeruleum f{. album W.T. Wang,
86

. corymbosum Brihl ex Huth, 86
. macranthum Finet, 114
var. magnificum Huth, 86
var. majus W. T. Wan 6
. obtusilobum Briihl he Huth, 86
. tenuicaule Briihl ex Huth, 86
calcicola W. W. Smith, 84
callerit Franchet, 9
camaonense Huth, 98
campylocentrum Maxim., 236
candelabrum Ostenf. in Hedin, 87
var. monanthum (Hand.-Mazz.) W.
Wang, 105
carduchorum Chowdh. & Davis, 16
carela Buch.-Ham. ex D. Don, 176
cashmerianum Royle, 60
var. walkeri Hook. f., 60
caucasicum C. A. Meyer, 61
var. bracteosum Schrenk, 229

var.
cavaleriense Lévl. & Vaniot, 9
centeteroides (Briihl) Munz, 17
ceratophoroides W. T. Wanir: 145
ceratophorum Franchet, 146
var. brevicorniculatum W. T. Wang,
238

var. hirsutum W. T. Wang, 146
cerefolium Lévl, & Vaniot, 9

uth,
ssp. middendor fii (Trauty.). ‘Bri,
89

ssp. schizophyllum Briihl, 89
ssp. vera Bri 9
var. albiflora Trautv., 89

1968]

var. oo (Stev. ex DC.)
Hut
var. Lee Trautv.
var. laxiflorum (Trautv.) Huth, 89
var. parviflorum (Turcz.) Huth, 89
var. subbifidum Winkler ex Huth,
nomen, 89
var. subglabrum ee 89
cheilanthum Franchet, 7
chenii W. T. Wang, [Bes
aiehonsé Fisch., 95
chodatii Oppenheimer, 18

r. pygmaeum Ostenf., 130
shiniin Steven, 204
var. dict ee tote DC. in Huth, 199
cinereum Boiss
coelestinum Pedal 156
coeruleum Jacquem., 86
coleopodum 4 “Mazz, 90
confusum M. , 192
conocentrum pe alae 77
cooperi Munz, 193
Mee ee Rege 1, 194
r. baicalense Huth, 194
Oaicah Ledeb., 195
crassifolium Schrad. ex Spreng., 196
var. tangutica Maxim., 196

cyphoplectrum Boiss., i

var. micranthum Bol a

var. pallidiflorum =e vie 41

var. stenophyllum Boiss.,

var. vanense (Rech. f.) Davis, 57
dahuricum Steven ex DC., 89
dasyanthum Kar. & Kir. 62

var. angustisectum Huth, 62

2

, 148
dasycaulon var. malabovicwm Huth,
67

dasystachyum Boiss. & Bal., 20

-

davuricum Besser, 89
delavayi Franchet, 150

MUNZ, ASIAN SPECIES OF DELPHINIUM 253

var. acuminatum Franchet, 150

forma aureum W.'T. Wang, 150
densiflorum Duthie ex Huth, 63

var. platycentra Briihl, 63

var. stenocentra Bruhl, 63
denudatum Hook. f. &

Thoms.

nanense Franchet, 128

ssp. proprium Brihl, 199
ssp. tibeticum Brihl, 199
var. cinereum Briihl, 199
var. glandulosum Briihl, 199
5

var. glaberrimum Trautv., 199

var. pubiflorum Trautv., 142
discolor Fisch. ex Sweet, 200
dissectum Huth, 93
dolichocentroides W. T. Wang, 94
dolichocentrum W. T. Wang, 120
dolichostachyum Chowdh. & Davis, 21
drepanocentrum (Brihl) Munz, 151
duhmbergii Huth, 198

var. retropilosum Huth, 226
dyctiocarpum DC., 199

var. glaberrimum Trautv., 199
elatum L., 200
ssp. intermedium Fleisch. & Lindm.,

200

ssp. intermedium Sergievsk., 200
var. gracile Sergievsk., 2
var. grandiflorum Sergievsk., 200
ee ‘plata (DC.) Sergievsk.,

var. Y baliod DC; 200
var. glabrum Sergievsk., 200
var. hispidum DC., 200
elatum L.
var. anomalum Kunth, 163
var. incisum Hook. f. & Thoms., 97
var. longicalcaratum ee 200
var. palmatum Lévl.,
ie pubiflorum kee) Huth,

var. pvinddatien Huth, 225
var. ranunculifolium Hook. f. &
Thoms., 224

var. sericeum W. T. Wang, 200
var. subciliatum Ledeb., 204
var. subglabrum Ledeb., 200

254

var. subglabrum ree . oe 220
villosum Ledeb.,

2
elliptico-ovatum W. T. Wang, 203
englerianum Huth, 198
var. hoffmeisteri Klotzsch, 198

eriocarpum (Boiss.) Halacsy, 1
eriostylum Leévl., 237
erlangshanicum W. T. Wang, 131
esquirolii Lév\. & Vaniot, 128
exiguum Pritzel ex Diels, 9
fangshanense W. T. Wang, 95
fargesii Franchet, 240
fengit . Wang, 67
fissum Waldst. & Kit., 22

ssp. anatolicum Chowdh. & Davis,

2

flexuosum Bieber., 204
var. bracteolata Busch, 227

204
. glandulosum Kapeller ex
Grossh., 25

var. hirsuta Busch, 204

var. lasiocarpum Rupr., 204

formosum Boiss. & Huet, 205
var. centiflorum Huth, 205

freynii Conrath ex Freyn, 47

georgei Comber, 152

gilgianum ager . Gilg & Loesen., 88
giraldii Diels,

glabellum ae 200

glabricaule W. T. Wang, 154

var. acaule Turcz., 95

. bhotanica Briihl, 95

. chinense Fisch. ex DC., 95

. davidii (Franchet) Briihl, 149
gilgianum (Pilger) Finet &
Gagnep., 88

var. glandulosum W. s nee 95
var. gmeliniit Reichb.,

JOURNAL OF THE ARNOLD ARBORETUM

[voL. 49
var. kumaonensis Huth ex Brihl &

. kunawarensis Bruhl, 86

. latisecta Maxim., 103

var. leiocarpum W. T. Wang, 95

. innaei Briihl & King, 95

var. majus W. T. Wang, 95
mosoynense Huth, 106

. obtusiloba Brihl, 86

. pallidum Huth, 95

. paradoxa Brihl, 95

A potaninit Brihl, 240

var. setchwanensis Brihl, 95

var. mea. Briuhl, 95

var. tigridium Kitagawa, 95

var. tsangensis Brihl, 95
grandiflorum are $
griseum Gilli,
gyalanum Marquand & Airy-Shaw, 155
halophilum Huth, 4

halteratum ‘Sibth. a Sm. var. venulo-

f. concolor W. T. Wang, 96
hillcoatiae Munz, 206
himalayae Munz, 207
himalayense Chowdh. ex Mukerjee,

207
hirticaule Franchet, 156
r. coelestinum Finet & Gagnep.,
6

var. micranthum Finet & Gagnep.,
156

var. mollipes W. T. Wang, 175
hoffmeisteri Klotzsch, 224
honanen Wang, 157
hui Chen, 132
hybridum Steph. ex Willd.,

ssp. Jaxiusculum var.

usch, 44

ssp. ochroleucum Busch, 39

ssp. puniceum Busch, 43

ssp. schmalhausenii Busch, 47
hybridum

var. albiflorum (DC.) Koch, 10
. dasyanthum =~ a 51
. debilius Ledeb.,

. gracillimum Hae 32

arn

var. hirtula Trautv., 32
var. laxiusculum Boiss., 32
var. ochroleucum Boiss., 39
var. pilosula Trautv., 54
var. puniceum Claus, 43
var. sulphureum Regel, 48
var. szowitsiana Trautv., 51

ilgazense Davi

iliense Huth, 198
var. angustatum Huth, 198
var. hispidum Huth, 198
var. macrocentrum Huth, 198
var. pubiflorum Huth, 198

incanum Royle, 45

incisum Wall.,

inconspicuum Sergievsk., 208

ai
jacquemontionum. ‘Camb. in Jacquem.,
59

jugorum Hand.-Mazz., 168
junceum DC.,

var. subveluéinun Ledeb., 1
kamaonense Huth, 98
kantzeénse W. T. Wang, 99
karataviense Pavlov, 11

var. eglandulosum W. T. Wang, 158
r. leiocarpum Brihl ex Huth, 158

eh Ta Fedtsch., 159
koelzii Munz,
kohatense (Briihl) Munz, 29
korshinskyanum Nevski, 200
kuanii W. T. Wang, 210
kumaonense, 98
kurdicum Boiss, & Hohen., 30

var, elongatum Freyn ors 30
kweichowense W. T. Wan
Bie baa: Ulbr. ex Rehd. _ Kobuski,

114

lacei Munz, 211

lacostei Danguy

lancisepalum me -Mazz., 160
lanigerum Boiss. & Hohen., 101
lankongense Franchet, 135
lasiocarpum Tamura, 161
latesquamatum Gilli, 31

MUNZ, ASIAN SPECIES OF DELPHINIUM 255

laxicymosum W. in Wang, 162
pees: DC.

r. alpinum aoe 217
tie ee Shae Rouy, 32
leiocarpum Hut
lepidum aes & fo Lall., 129

s 212

var. hirticaule Grossh., 190
var. crispulum Busch, 91
lipskii Korsh., 165
longebracteatum age ire 33
longiciliatum W. T. Wan
longipedunculatum ‘Sethty & ae
4

longipes Franchet, 149
ludlowianum Chowdh. ex Mukerjee,
16

ludlowii Munz, 166
lycoceomttolleen Lévl., ae
maackianum Regel, 2
var. cuneatum “peel ex Huth, 214
var. lasiocarpum Huth, 21
var. palmatum (Lévl.) Nakai, 214
var. ussuriense Regel, ue 14
macropogon Prokhanov
macros ee Boiss. ex Hak, 36
mairei Ulbr.,
var. viride ee W. T. Wang, 67
malabaricum (Huth) Munz, 167
malacophyllum Hand.-Mazz., 102
mariae Busch, 215
maximowiczii Franchet, 103

W. T. Wan
- ii rt ex Mukerjee,

micropetalum Finet & Beta 168
f. album 8

wiuktctork Trautv., 89
minjanense Rech. f., 68
minutum Lévl. & Vaniot, 9
mirabile Sergievsk., 217
mitzugense Ulbr., 169
molle Danguy, 69

moschatum Munro ex ‘Hook. i &
Thoms., 59

256

mosoynense Franchet, 106
muliense W. T. Wang, 170
var. minuiibracteolatum er Ls
Wang,
muscosum Exell & Hillcoat, 107
nachiczevanicum Tzvelev, 41
nanum 5
naviculare W. T. Wang, 218
var. lasiocarpum W. T. Wang, 218
nepalense Kitamura & Tamura, 70
nevskii Zakirov,
nordhagenii Wendelbo, 71
nortonii Dunn, 108

DC. 39
var. sulphureum Regel, not Boiss.,
39

oliganthum Franchet, 133
oreophilum Huth, 221
orthocentrum Franchet, 171
osseticum Busch,

8

var. peeve (Hand.-Mazz.) W.

T. Wang, 1

var. lobatum a T. Wang, 173

var. laegeo -lancisepalum W. T.
Wang, 1

var. Jems Chen, 170

oyle, 92
pauciflorum Reichb. be Ledeb., 89
pediforme Comber,
pellucidum Busch, 1
penicillatum Boiss., 42
var. macroplectrum Buser in Boiss.,
5

penicillatum Hook. f. & Thoms., 55
peregrinum L

JOURNAL OF THE ARNOLD ARBORETUM

[voL. 49

ssp. sa ty Batt.,
ssp. nanum (DC.) siaebake 3
j boissieri Huth, 3

. subsaccatum Huth, 3
; subvelutinum De. 1
virgatum nee 2
pitntuitil Fedtsch.,
pogonanthum Hand. ‘mele: 174
poltaratzkii Rupr., 223
polyanthum W. T. Wang, 239
ponticum Hees & Bornm., 10
potaninii Huth, 240
var. latibractéolatum W. T. Wang,
240

3
fi

propinquum Nevski, 72

przewalskii Huth, 109

pseudo-caeruleum W. T. Wang, 110

pseudo-campylocentrum W. T. Wang,
175

pseudograndiflorum W. T. Wang, 111
var. glabrescens W. T. Wang, 111
var. lobatum W. T. Wang, 111

pseudo-tongolense W. T. Wang, 241

h, 95
pulchellum Chowdh. ex Mukerjee 224
pulcherrimum W. T. Wang, 1
pumilum W. T. Wang, 113
puniceum Pallas, 43
purdomit Craib, 74
pyenocentroides W. T. Wang, 134
r. latisectum 2 ees 119

Parente Fran

var. lankongense “CE facts Huth,

tos

pylzowii Maxim.,

var. trigynum W. a Wang, 114
pyramidale Royle, 224
pyramidatum Alboff, 225
quercetorum Boiss. & Hausskn., 44
ranunculifolium Wall., 224

var. incisum ae i & Thoms., 97
rectivenium Roy
retropilosum (Huth) So Sciam 226
rha Schipez. ex Nevski, 2
robertianum Lévl, & oe 8)
rockii Munz, 115

1968]

rotundifolium, 221
roylei Munz,
ruprechtii Nevski, 227
salmoneum Mouterde, 27
coos Boiss., 46
ssp. centeteroides Brihl, 17
var. alacialia Boiss., 12
var. gilgitense Brihl ex Huth, 29
var. kohatense Brihl ex Huth, 29
var. kurramensis Brihl, 50
var. refracta Brihl & King, 46
sauricum Schischk., 228

Sscaposum W. T. Wang, Lor
schmalhausenii Alboff, 47
schroedingerianum Hand.-Mazz., 19
semibarbatum Bienert ex Boiss., 48
var. hoeltzeri Huth, 48
semiclavatum Nevski, 49
sertiferum Franchet, 173
sherriffi Munz, 177
siamense (Craib) Munz, 178
sinense Fisch. ex Link, 95
sino-scaposum W. T. Wang, 137
sino- vitifolium W. T. Wang, 242

. speciosum Bieberstein, 229
ssp. ranunculifolium Brihl, 224
var. brachycentra Brihl, 224

var. marriensis Brihl, 224
var. oinophylia Brihl, 224
var. pulverulenta Brihl, 224
var. thyrsantha Brihl, 224
var. boissieri Buhse, 229
var, bracteosum Huth, 229
var. brevicalcaratum cages! 64
var. bruehlianum age
var. caucasicum Hut
var. dasycarpum ae haan a 148
var. gymnopus Rupr., 229

MUNZ, ASIAN SPECIES OF DELPHINIUM 257

var. linearilobum Trautv., 91
ar. paulovii Busch, 2
hae Bhim ce (Royle) Mukerijee,

var. manne (Wall.) Huth,
224

var. sphenolobum Brihl ex Huth, 76
var. trichocarpum Rupr., 229
var. typicum biebersteini Rupr., 229
speciosum Huth,
spirocentrum Han
pomlibvetealaint we a.

43
var. hirsutum Chen, 243
var. pauciflorum Ch en, 235
var. sag vile (Comber) We,

ae Briihl ex Huth, 230
var. khasiana Brihl, 230
var. shanica Bruhl, 189
var. siamense Craib, 178
staphisagria
stenosepalum Taree, 144
suave Huth,
subvelutinum ead ex Boiss., 1
sulcatum Reichb. ex Hoffmanns., a
sungpanense W. T. Wang, 180
sutchuenense Franchet, 118
sylvaticum Turcz., 89

m Boi
ar. i ates (Boiss.) Huth,
ae

taipaicum W. T. Wang, 139
taliense Franchet, 181

tatsienense pian 119
tauricum Pallas, 4

p., 140
var. schizophyllum Hand.-Mazz., 134
eG sags oe Finet & Gagnep.,

cicaaulele W. T. Wang, 231

258

tomentellum Busch, 91
var. araraticum Busch, 91
var. angustibracteatum Busch, 91

forma breviungue Lévl.,
. subglaberrimum Hand.-Mazz.,
4

var, tibeticum Lévl., 74
trifoliolatum Finet & Gagnep., 183
trilobatum Huth, 77
trisectum W. T. Wang, 122
triste Fisch. ex DC., 123

var. linearilobum Huth, 91
tsarongense Hand.-Mazz., 75

var. patentipilum W. T. Wang, 75
tsoongii W. T. Wang, 124
tuberosum Aucher ex Boiss., 53

var. leiocalycinum Bornm., 53

ukokense Sergievsk., 232
umbellatum Regel ex Nevski, 194
umbrosum Hand.-Mazz

var. aitchisonii Boiss. ex es 55
var. pseudovestita Brihl,
var. vicaryi Brihl, 55

uralense Nevski, 185

JOURNAL OF THE ARNOLD ARBORETUM

[voL. 49

ursinum Rech. f., 56
vanense Rech. ‘. 57
venulosum Boiss., 5
vestitum Boiss.,
vestitum Wall.
var. praia a. ere & Rupr.,
62

var. pumilum Huth, 76
var. sphenolobum Briihl ex Huth, 76

uth, 233
var. ie eee DC.) ‘ill 163
var. winkleri ees
virgatum Poir.,
virgatum Jacq. 5 ex Spreng., 95
viride Chen, 67
viscosum Hook. f. & Thoms., 77

var. chrysotrichum net ex Huth,
77

var. gigantobracteum Brihl ex Huth,
iz

vitifolium Finet & Gagnep., 242
wardii Marquand & = Shaw, 78

williamsii ee 125

wilsonii Munz, 126

winklerianum Huth, 234

Vv
zalil Aitch. & Hemsl., 48

Staphisagria macrosperma, 8

ADDENDUM

Chienia honanensis W. T. Wang. Dr. Lothar Benzing of the Institut fiir All-
gemeine Botanik at Mainz, West Germany, has very kindly called my attention
to, and sent me a xeroxed copy of an article in Acta Phytotaxonomica Sinica 9:
104-107. pls. VI, VIT. 1964, in which W. T. Wang proposed a new genus oe
closely related to Consolida and Delphinium. As the Royal Botanic Garden a
Edinburgh had received photographs, and even isotypes, of many of Wan re
earlier species of Delphinium, I asked Dr. Peter Davis whether they had any
material of the Chienia, but word has just come that he and Mr. Lauener have

none.

Wang’s excellent illustration of Chienia honanensis W. T. Wang and the de-
tailed Latin description ions of the genus and species certainly suggest Delphinium
grandiflorum L. in habit, vesture, leaf dissection, flower color, and size. The

1968 | MUNZ, ASIAN SPECIES OF DELPHINIUM 259

plant comes from Honan, China, which is far enough north to be in the range
of the Delphinium, from which it differs in the shorter spur on the upper sepal
and the lack of spurs on the upper petals SS of Wang), so that the
upper and lower petals are quite alike and clawed.

In the course of examining perhaps 6000 phere of the Consolida- Delphin-
ium complex I have met a number of cases of spurlessness. For example, in the
discussion of Comsolida rugulosa (Boiss.) Schréd. it was pointed out (Jour.
Arnold Arb. 48: 188, 189. 1967) that C. paradoxa (Bunge) Nevski [ Delphinium
paradoxum Bunge] was apparently a spurless form of C. rugulosa. In Delphin-
tum anthriscifolium Hance a spurless collection was seen, H. B. Morse (Ny). In
D. macrocentron Oliver, Bally 1205 (kK) has spurless petals. I remember seeing
a sheet of another species in which spurred and spurless flowers occurred on the
same plant, but unfortunately I have misplaced my record of it.

Without seeing material of the proposed Chienia honanensis, I am inclined
to suspect that Wang, whose comprehensive treatment of the Chinese species of
Delphinium (Acta Bot. Sinica 10: 59-89, 137-165, 164-284. 1962) has given him
a much better knowledge of those species than I have, has dealt with an abnormal
plant of Delphinium grandiflorum L.

RANCHO SANTA ANA Botanic GARDEN

1500 NortH CoLLecE AVENUE
CLAREMONT, CALIFORNIA 91711

260 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

MORPHOLOGY AND ANATOMY OF
CROOMIA PAUCIFLORA (STEMONACEAE)

P. B. ToMLINSON |
AND
E. S. AYENSU

Tur STEMONACEAE (Roxburghiaceae) comprises a small and taxonomi-
cally isolated family of monocotyledons with its main distribution in the
Far East. Stemona ranges from the Himalayas to southern Australia and
Stichoneuron is found in Indonesia. The remaining genus Croomia (com-
memorating the pioneer Florida botanist H. B. Croom), has a widely dis-
junct range with two species in Japan and one (C. pauciflora Torr.) in the
southeastern United States. This family therefore provides a further
example of distribution demonstrating the ancient floristic affinity between
eastern North America and eastern Asia to which Asa Gray drew attention.

There is little agreement amongst taxonomists as to where in a natural
sequence the Stemonaceae should be situated. Burkill (1960) provides
a useful survey of the taxonomic history of the family. Its wide and dis-
continuous distribution has led to the assumption that it is ancient
(Hutchinson, 1959) but there is much evidence that the taxon is not very
homogeneous. Any information likely to shed light on the systematics
of the family is therefore of value, especially in view of its possible rela-
tionship with the economically important Dioscoreaceae. Recently one of
us (P. B. T.) had the opportunity to collect Croomia pauciflora in north-
ern Florida and to investigate its morphology and anatomy with ample
fresh material at hand. The other (E. S. A.) has investigated the vegeta-
tive anatomy of the three genera and the total contribution permits a dis-
cussion of their systematic affinities. Only limited herbarium material
of Stichoneuron and Stemona has been available for microscopic investi-
gation, and observations on these two genera are still very tentative. The
vegetative anatomy of Croomia pauciflora was investigated previously by
Holm (1905) with some of the information repeated in a later paper
(Holm, 1927), but it is quite clear that many fundamental morphological
features have hitherto remained undescribed. They are presented in this
paper, together with a detailed re-examination of anatomy. In addition
the anatomy of the flower has been studied in order to compare it with
that of Stemona about which there is a good deal more information.

In the United States Croomia grows on richly wooded bluffs along the
Apalachicola River from Florida into Alabama (Harper, 1942). Plants
form small, very localized populations in dense shade on steep, well-drained

* Research on the systematic anatomy of
aloe y monocotyledons supported by N.S-F.

1968 | TOMLINSON & AYENSU, CROOMIA 261

slopes amongst loose litter. According to Harper they remain green all
summer. Populations seem to persist and spread largely by rhizomes
which are, however, shallow and easily dug up. We have not seen any
seedlings. Harper spoke of plants being “more or less connected by a
network of rootstocks.” This is substantiated by the abrupt boundary
to the populations and the high density of leafy shoots. Rhizome mor-
phology described below accounts for this method of vegetative survival
but also suggests that Croomia would be vulnerable to any disturbance
of its habitat.

MATERIAL AND METHODS

Living plants came from Torreya State Park, Bristol, Florida. We are
indebted to Dr. S. McDaniels and Dr. A. F. Clewell of Florida State
University for the arrangements made so that specimens could be collected.
Plants were examined largely in a fresh condition. Microscopic details
were studied in unstained freehand sections cut with a razor blade, sup-
plemented by observations on preparations made by routine clearing,
staining, and maceration methods. Details of the vascular system were
examined in serial sections of paraffin-embedded material stained in
safranin and Delafield’s haematoxylin. The vascular anatomy of open
flowers and mature flower buds was studied in serial paraffin sections and
in whole flowers cleared and stained in safranin. A number of foliage
leaves were cleared and stained in Heidenhain’s haematoxylin.

Material of Stichoneuron and Stemona from the Herbarium, Royal
Botanic Gardens, Kew, as follows, was supplied through the generosity of
Dr. C. R. Metcalfe, Keeper of the Jodrell Laboratory: Stemona curtisi
Hook. f. (Haniff & Nur 4368); S. kerrii Craib (Kerr 707); Stichoneuron
caudatum Ridley (Corner 28716 & 37056).

VEGETATIVE MORPHOLOGY

GrowrtH HABIT (Fics. 35-39). Growth of shoots is sympodial with a
remarkably precise and constant arrangement of parts on each unit of the
sympodium. Each of these units consists of an underground rhizomatous
portion terminating in an erect leafy shoot which bears the flowers (Fic.
35). At the time of flowering, in March to April, the renewal shoot has
already been initiated (Fic. 37) for the next unit which will culminate
in the following year’s erect shoot. Leafy shoots are, therefore, annual but
the slender rhizomatous part of the sympodium is long persistent and
marked by the scars of each erect shoot. These scars therefore represent
the limit of each annual growth increment. Long rhizomes with many
scars can be dug up easily. :

Each unit of the sympodium begins with a series of scale leaves. Their
position and number are so constant from year to year that they may
be identified by letters for convenience (Fics. 35, 37). Leaves are ar-
ranged distichously (Fic. 38). The plane of distichy is horizontal »
though this is interrupted once in a very significant way. The following

262 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

sequence is standard for each renewal shoot. The first leaf (A) is a scale
leaf at the end of a long internode; it subtends no axillary bud. Leaf
B is a scale leaf opposite leaf A and also at the end of a long internode.
It subtends a conspicuous bud which is, however, strongly inhibited (dor-
mant bud, Fic. 39). Leaf C is a scale leaf at the end of a short internode
and inserted on the dorsal (lower) side of the rhizome and _ therefore
perpendicular to the plane of distichy of leaves A and B. Leaf C subtends
the bud which grows out, without inhibition, as the renewal shoot (renewal
bud). Beyond leaf C the axis turns erect (Fic. 37). Three (sometimes
only two) overlapping scale leaves (D, E or F) separated by short inter-
nodes are inserted at the base of the erect shoot. These re-establish the
earlier distichy so that D is always on the same side of the shoot as B.
The last scale leaf (E or F) is often somewhat remote from the other
two and its apex is usually above ground. The extent to which the inter-
node below the last scale leaf is elongated and whether two or three scale
leaves occur beyond leaf C may be determined by the depth to which
the rhizome is buried. These 2 or 3 ultimate scale leaves subtend no
conspicuous buds although microscopical examination of sections through
their nodes sometimes reveals vestigial buds with little or no vascular con-
nection to the main vascular system. These may be described as non-
functional buds since they never seem to expand and are in sharp contrast
to the conspicuous functional buds at nodes B and C which have a strong
connection to the main vascular system of the rhizome.

Beyond the last scale leaf (E or F) is the long internode, up to 30 cm.
tall, which supports the leafy crown. This internode is white below
ground but exposed parts are red below and green above. The exposed
apex of the distal scale is also red. Foliage leaves from a terminal, irregu-
lar, obliquely ascending series spreading backwards more or less in one
plane and apparently continuing the distichy of the scale leaves although
they are irregularly spaced and often subopposite (Fic. 36). The axis of
the whole sympodium is apparently terminated by a leaf or pseudo-pair
of leaves. Vigorous shoots bear as many as nine leaves, from five to seven
are the most common numbers. Small shoots may have as few as two or
three foliage leaves but we have seen no shoot with one leaf. Flowering
branches are in the axils of leaves. In non-flowering shoots there may be
vestigial axillary buds.

From the above description it is evident that branching is very re-
stricted. Each sympodial unit bears only two functional buds (Fic. 38),
one (renewal bud) is necessary to maintain the sympodium, leaving only
one (dormant bud) to effect multiplication by branching. At the time
of anthesis the renewal bud has usually commenced growth, piercing the
back of its subtending scale leaf (Fic. 37). At this stage the primordia
of the first two scale leaves of next year’s increment are already present
but they so firmly enclose the bud that its apex appears to be naked and
leafless. Examination of herbarium specimens collected in the fall suggest
that the rhizome overwinters with a full complement of leaves initiated.

In contrast, the dormant bud is strongly inhibited because many rhi-

1968 | TOMLINSON & AYENSU, CROOMIA 263

zomes remain undivided even after several years’ growth. However,
divided rhizomes, in which the dormant bud has grown out to initiate a
new sympodium, are quite common. Normally this branch axis repeats the
arrangement of parts shown on the parent axis and produces a vigorous
but flowerless leafy shoot in the first year. Commonly, however, the lateral
shoot is not vigorous and the first two internodes remain unextended so
that a short, depauperate leafy shoot with few (2-3) foliage leaves is
erected close to the parent axis. Although it is reasonable to assume that
damage to a vigorous renewal shoot will release inhibition of dormant buds,
most branched rhizomes appear undamaged. Therefore, it is not easy to
account for the outgrowth of dormant buds which otherwise are strongly
suppressed.

Roots are restricted to the nodes at which leaves B and C are inserted,
from three to five growing out adventitiously on the lower side of the
rhizome (Fics. 35, 37). Roots are somewhat fleshy and produce a fine
system of first-order branches but without conspicuous root-hairs.

Leaves. Scale leaves (Fics, 12-16) borne on underground parts are
readily distinguishable from foliage leaves borne on aérial parts. In many
monocotyledons the morphological nature of the scale leaf, which is
equivalent to the leaf base of a foliage leaf, is clear from a series of
transitional leaves. In Croomia, without transitional leaves, this obvious
relationship is lost. The two types of leaf seem to be very dissimilar
developmentally because the foliage leaf has no sheathing base.

Scale leaves are open, opposite margins scarcely overwrapping. Leaves
A and B are usually shortest with fewest (usually 3-4) veins (e.g. Fic.
13) whereas distal scales are longest and almost invariably have five veins
(e.g. Fics. 14-16). A regular progression in size along the rhizome is not
always maintained, however (e.g. Fics. 12-16), perhaps because growth of
the renewal shoot is not continuous. Veins are parallel, converge and
unite at the apex but are unconnected by transverse veins. A conspicuous
mid-vein is not developed. ;

Foliage leaves (Fics. 35, 36; 9) are petiolate but without a sheathing,
encircling insertion. The blade has five major veins, prominent abaxially,
the veins entering the blade from the petiole independently of each orner.
The outermost vein on each side gives off a marginal and two submarginal
veins into the cordate base of the lamina. The veins unite in turn with the
marginal commissure which itself joins the median vein at the apex (Fic.
9). Longitudinal veins are connected by an irregular series of transverse
veins. Many minor transverse veins end blindly (Frc. 10), an unusual
feature for a monocotyledon, not found in Stichoneuron and very rare in
Stemona.

VEGETATIVE ANATOMY

LAMINA OF FOLIAGE LEAVES. Dorsiventral. Hairs absent. Cuticle thin
on both surfaces. Epidermis moderately shallow, more or less uniformly
thin walled, outer wall scarcely thickened. Adaxial epidermis (Fic. 1)

264 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Fics. 1-16, Croomia pauciflora. Leaf anatomy. Figs. 1 and 2. Surface view
of epidermis, X 100. Fic. ial view. Fic. 2. Abaxial view. Fic. 3.
Transverse section of stoma, < 470, from abaxial epidermis of lamina Fics.
4 and 5. Mesophyll in paradermal view from cleared whole mount of lamina, x

1968] TOMLINSON & AYENSU, CROOMIA 265

uniform in surface view, cells more or less isodiametric with markedly
sinuous anticlinal walls; costal cells elongated, narrow and with less ob-
viously sinuous but somewhat thicker walls than intercostal cells, the
costal cells most distinct above and below large veins. Abaxial similar to
adaxial epidermis except for numerous stomata (Fic. 2). Stomata
anomocytic, 27-36 » long, the stomatal pore apparently orientated at ran-
dom. Each guard cell with a wide lumen, somewhat thickened towards
the pore below the two more or less equal ledges (Fic. 3). Guard cells
including numerous small chloroplasts. Marginal epidermal cells papillose,
the outer wall distinctly striate (Fics. 8, 11).

Mesophyll uniformly chlorenchymatous, 3—5-layered; mostly 3-lay-
ered at margin and becoming several-layered towards midrib and main
veins. Mesophyll cells much lobed; adaxial layer (Fic. 4) relatively com-
pact, but cells not anticlinally extended to form a distinct palisade; abaxial
layer (Fic. 5) loose, much lobed, the intermediate mesophyll layers inter-
mediate in extent of lobing; differences between successive mesophyll
layers most evident in paradermal view (Fics. 4 and 5). Chloroplasts
appearing somewhat denser in adaxial as compared with abaxial mesophyll
layers. Veins (vascular bundles) all alike apart from range in size, from
small blind-ending veins to midrib (e.g. Fics. 6 and 7); all separated
from each epidermis by at least one layer of chlorenchyma. Largest veins
(Fic. 7) resembling vascular bundles of petiole and with inconspicuously
collenchymatous subepidermal layers below; small veins with reduced
vascular tissues (Fic. 6). Veins inconspicuously sheathed by a single
layer of elongated chlorenchymatous cells, without thick-walled sheathing
layers except for cells below phloem with slightly thickened walls.

PeTIoLe (Fics. 17 and 18). Epidermis without sinuous walls, resem-
bling that of aérial stem, with somewhat thicker walls than epidermal cells
of lamina. Cuticle conspicuously striate. Stomata common, Outermost
layer of chlorenchymatous ground parenchyma somewhat collenchymatous
but collenchyma most conspicuous in abaxial hypodermal layer and in
petiole margin. Vascular bundles collateral, usually 5 throughout
greater part of petiole (Fic. 17), but 7 according to Holm; each vascular
bundle (Fic. 18) with many narrow adaxial tracheal elements, the nar-
rowest, adaxial (protoxylem) elements often partly occluded. Phloem
with irregularly arranged, narrow sieve tubes. Bundle sheath cells usually
distinctly collenchymatous.

ScALE LEAF. Epidermal cells thin walled, more or less rectangular

35; scales laid flat, & 1.

266 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49
in surface view. Mesophyll of uniform colorless parenchyma. Veins
resembling major longitudinal veins of foliage leaf.

ARIAL stem (Fics. 19-22). Stem more or less circular in transverse
section, including up to 13 but usually 10 vascular bundles arranged in a

: od).
‘26 Ss
once

\y

Pi

,
(-

fi
x YY

ae
Ly
ay YX
Aa Cry
ses
al

Not OAK
5 VERO eX

% ae > “ 7
Oe
oy

Fics. 17-22, Croomia pauciflora. Anatomy of petiole a éri Fics.
17 and 18. Petiole. Fic. 17, Transverse pe ioe xX 10. oe pas
section of single collateral vascular bundle from petiole, x 180. ‘Fics. 19-22.
Aérial stem. Fic. 19. Transverse section, X 10. Fic. 20. Epidermis " surface
view, X 100. Fic. 2 1. Surface layers of stem in transverse section, < 100. FIG.
22. Transverse section of single amphivasal vascular bundle, < 180.

1968 | TOMLINSON & AYENSU, CROOMIA 267

circle (Fic. 19), the number of bundles increasing distally towards the
insertion of foliage leaves. Stomata absent. Cuticle thin. Epidermis
somewhat thick walled; epidermal cells longitudinally extended, more or
less rectangular in surface view but commonly with oblique end walls
(Fic. 20), normally colorless but basal cells with red (presumably antho-
cyanin-containing) vacuoles. Cortex with outermost 2-3 layers small
celled and distinctly collenchymatous (Fic. 21), the collenchyma most
pronounced distally; ground parenchyma otherwise uniformly large celled,
thin walled, and colorless except for chloroplasts in outermost cells distally.
Medullary ground parenchyma cells widest and with some tendency to
collapse in a lysigenous manner. Each of largest vascular bundles (Fic.
22) amphivasal, including a continuous cylinder of tracheal elements,
uniseriate except in region of narrow, often obliterated protoxylem ele-
ments on the inner side. Narrower vascular bundles collateral, the xylem
discontinuous on the outer side and without conspicuous protoxylem.
Phloem with inconspicuous sieve tubes surrounded by xylem and in turn
surrounding a central strand of somewhat inflated cells (F1c. 22) with ap-
parent wall thickenings which may represent obliterated protophloem.
Vascular bundles sheathed by 2-3 layers of narrow, slightly thick-walled
fibers or prosenchymatous cells, the sheath in large bundles usually in-
complete next to protoxylem.

RuHIzoME (Fics. 23-26). Epidermis uniform, thin walled and closely
resembling that of aérial stem. Cortex wide, uniformly parenchymatous
and with abundant starch except for somewhat narrower but scarcely
collenchymatous peripheral cells (Fic. 25). Stele angular in outline
(Fics. 23 and 24), delimited from cortex by a more or less continuous
but indistinct endodermis of cells with suberized walls but without obvious

tissue resembling cortical ground tissue.

Roor (Fics. 28-32). Epidermis persistent but often collapsing some-
what; cells in surface view more or less rectangular, somewhat longitudi-
nally extended (Fic. 32). Root-hairs uncommon, arising as outgrowths of
otherwise unmodified cells (Fic. 31). Exodermis a compact layer of nar-
row, suberized, but scarcely thick-walled cells (Fic. 30). Cortex very
wide, of uniformly starch-filled parenchyma with well-developed intercel-

268 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

[.) ‘yy pd
= (Ee ER I |
= RESIN WA rer
— Bl! a 6-6 a 32
a e :
=> 6
——

neice,
KARR
SRS

baer a Croomia paucifiora. _Anatomy of rhizome and root. Fics. 23-27.
re IG. 23, Transverse section, X 10, stele cross-hatched. Fic. 24. Dia-
g o show insertion of vascular tissues of lateral organs on rhizome. Leaf

1968 | TOMLINSON & AYENSU, CROOMIA 269

lular space system; innermost cells somewhat wider than outermost.
Endodermis inconspicuously differentiated from innermost cortical layer
but small celled, thin walled and with conspicuous Casparian strips (Fic.
29). Pericycle uniseriate, thin walled, inconspicuous but usually dis-
tinguished from tissues within by its larger cells. Stele narrow (Fic. 28),
polyarch with few (usually 6-7) xylem poles, the xylem arms irregular.
Tracheal elements, together with central, thick-walled medulla, form-
ing a fairly regular stellate core, the narrow, thin-walled phloem strands
occupying the angles between the protoxylem poles (Fic. 29). Lateral
roots are diarch with a more conspicuous endodermis according to Holm
(1905).

CONDUCTING, STORAGE AND SECRETORY ELEMENTS. Vessels apparently
restricted to metaxylem of roots (Fics. 33 and 34); elements average 2
mm. long and 33 » wide, each with long scalariform perforation plates on
very oblique end walls up to 300 p» long, the thickening bars between the
perforations (Fic. 34) scarcely thinner than those on very similar im-
perforated end walls of tracheids in other organs; lateral walls with scalari-
form pitting in areas of contact between adjacent vessels, otherwise walls
with diffuse, narrow pits. Tracheal elements in rhizome often much
longer than those in root; end walls scalariformly pitted but imperforate.
Tracheids in aérial stem and leaves without definite end walls and with
annular or spiral, rarely scalariform-reticulate wall thickenings, the thick-
ening usually wound in one gyre. Sieve tubes in leaf, aérial stem and
rhizome mostly with simple sieve plates on transverse to oblique but often
rather indistinct end walls; sieve plates in sieve tubes of root compound,
on oblique end walls. :

Starch grains abundant in ground parenchyma of rhizome (Fic. 27)
and root; grains up to 18 » wide, ellipsoidal, flattened; hilum inconspicu-
ous but not obviously excentric, Crystalline or other secretory products
not observed.

COURSE OF VASCULAR BUNDLES

Early authors had included Croomia in the Berberidaceae because the
morphology of its embryo was uncertain and the rhizome had been re-
garded as having a closed cylinder of wood. This was corrected by Gray

: : h trace
traces to subtending scale leaf C, solid black. Vascular masses of branct ;
stippled. Vascular bundles of rhizome stele cross-hatched. Traces to two roots

wn
o
2.
94]
ou
o
nm
°
=
°o
i=)
Oo
—
2
Lan |
aa
oO
2
3
oO
=
—_
<
9
w
2
—
o
=
i
oo

dle,
400. Fics. 28-34. Root. Fic. 28. Transverse section, X 10. Fic.
section of stele, X 180. Fic. 30. Transverse section of we fide rate
ae 31. Longitudinal section of root hair, X 100. Fic. 32. Epiderm x
branch root, surface view, . Fic. 33. . ,
extent of perforation indicated by solid black. Fic. 34. Details of vessel element,
180, lower limit of a perforation plate.

270 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

(1859). The vascular bundles which constitute the vascular system of
the rhizome are certainly not obviously discrete but they are best re-
vealed by tracing the changes in anatomy at nodes B and C where scale
leaves, roots, and functional buds are inserted. The basic pattern is
indicated diagrammatically in Fic. 24 which is constructed from serial
sections through node C. The vascular system of the rhizome consists of an
irregular series of larger strands alternating with smaller strands which
are the traces to the scale leaves. These traces appear well below their
exsertion and pass gradually into the scale leaving a distinct “gap” in the
main rhizome system. Two are usually conspicuous in a lateral position
as indicated in the description of the rhizome. The bud trace system is
a series of numerous isolated xylem and phloem strands which are aggregat-
ed into two large masses, each mass attached to a main rhizome strand.
The middle trace of the subtending scale leaf passes between these two
masses of bud trace tissue. Root traces may be attached directly to the
branch system, others directly to large strands on the upper side of the
rhizome. Beyond this complex nodal system there is a rapid change to
the ring of discrete vascular bundles found in the erect stem, these bundles
initially giving off traces to scales D, E and F. At the distal end of the
aérial stem traces to foliage leaves and branches diverge shortly below
the leafy shoot. There are no complicated nodal anastomoses such as
have been described for Stemona by Lachner-Sandoval (1892).

REPRODUCTIVE MORPHOLOGY

GENERAL MORPHOLOGY (Fics. 40-46). Flowers (Fics. 40-44) are
borne in long-stalked, few (1—3)-flowered cincinni (Fic. 46) in the axils
of foliage leaves on vigorous shoots. Each flower is terminal on an
axis bearing a single scale leaf (bracteole) which subtends the next flower-
bearing axis. Each bracteole appears to be at right angles to the preceding
one. Each pedicel has a distinct joint beyond the bracteole, representing
a line of abscission. There are 4 green tepals in 2 pairs at right angles,
the outer incompletely enclosing the inner (Fics. 40-44). All tepals are
somewhat reflexed in the open flower but in bud the apex of each, especially
the inner pair, is distinctly hooded. The four stamens, each opposite a
tepal, constitute a single whorl. Their bases are widened and united. In-
Stemona there have been attempts to interpret the stamens as forming two
series so that they would correspond to the tepals, e.g. by Swamy (1964)
who describes the stamen traces in Stemona tuberosa as diverging in pairs
at different heights; but in this species Baillon (1860) showed the stamen
primordia to arise simultaneously, as did Lachner-Sandoval (1892) in
Stemona javanica. Each stamen (Fics. 40-43) has a short fleshy filament,
enlarged above, with two lateral 2-lobed thecae inserted somewhat
obliquely and with latrorse dehiscence. Pollen grains are more or less spheri-
cal and with a finely reticulate surface (Fic. 45). The ovary is globular,
unilocular, and appears to be unicarpellate. It is grooved on one side only
(the presumed ventral side (Fic. 42)). The placentation is unusual with

1968 | TOMLINSON & AYENSU, CROOMIA 271

2-5 (usually 4) ovules all pendulous from an apical placenta below the
short and inconspicuously 2-lobed stigma. However, the bitegmic ovules
are irregularly anatropous so that the micropyle is directed upwards. The
funiculus varies in length so that the ovules hang at varying heights in the
loculus. A conspicuous feature of the funiculus is the mat of short, uni-
seriate hairs which later develop into the filamentous aril described and
illustrated by Gray (1859). There are no nectaries but the epidermis on
the inner surface of the tepals and of the filament and exposed part of the
connective is papillose, the enlarged cells being densely cytoplasmic and
filled with anthocyanin (Fic. 42).

VASCULAR ANATOMY (Figs. 47, 48). The vascular system of the flower
is very simple (Fic. 47). Each tepal has five vascular bundles, a median
and two pairs of laterals. All tepal bundles end blindly in a distal direction
and never converge or anastomose although they may produce short, blind-
ending branches. Each stamen has a single vascular bundle and the carpel
has two. The vascular system of the flower arises from an irregular ring
of narrow vascular strands, which may be regarded as a very reduced
version of the system in the aérial axis. The way in which vascular strands
initially diverge is not very uniform. The tepal and stamen traces arise
first, the remaining central bundles aggregating to form the 2 bundles which
irrigate the carpel. Tepal traces are small and collateral, each pair of
laterals diverging from a common bundle; the median usually originates
independently but sometimes diverges from the common bundle of one of its
lateral pairs. The stamen trace most commonly diverges from the median
trace to the corresponding tepal, less commonly it originates directly from
the pedicel system, independently of the tepal traces. It may, however,
diverge from the common bundle which produces 2 lateral tepal traces.
The stamen trace is irregularly collateral with an admixture of phloem
and xylem elements, most pronounced in the connective where the bundle
is enlarged. There is no indication of the stamen appendage which is such
a conspicuous feature of the flower of Stemona.

There is no difficulty in interpreting the vascular system of the ovary
(Fic. 48) according to the classical theory of carpel morphology and its
modern counterpart, which interprets the carpel as a conduplicate struc-
ture. One strand (the smaller) can be regarded as the dorsal bundle. It
runs independently up one wall of the ovary into the stigma where it
divides into 2 short procambial strands. In the opposite wall is the ventral
bundle, beneath the distinct ventral groove. It bifurcates at the apex of the
ovary, the two bundles constituting the vascular supply to the apical
placenta. Each ovule is supplied by a branch from one half of this placental
supply. The ovule trace ends in the funiculus, there being no vascular
supply to the integuments, at least at the time of anthesis. There is no
anatomical indication that either the dorsal or the ventral carpel bundle
is a dual structure; in fact xylem and phloem are rather diffuse. The gen-
eral configuration of this ovary recalls that of Sarcandra and other Chlor-
anthaceae described by Swamy and Bailey (1950), except that in Sarcandra

272 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

the ovary has one ovule and the dorsal and ventral traces are most com-
monly double.

DISCUSSION

GrowTH HABIT. Our analysis of Croomia demonstrates that each unit
of the sympodium has only two functional buds (the renewal bud and the
dormant bud), the position of which is quite fixed. The rotation of one
scale so that the renewal bud it subtends finds a position most favorable
for continued growth is noteworthy. Of the two buds, one maintains the
sympodium leaving only one to propagate the rhizome by branching. Even
so, this is strongly inhibited and thereby the potential for vegetative prop-
agation is reduced, It must be admitted that minute buds can be demon-
strated, by microscopical methods, in the axils of other scale leaves. Their
feeble vascular connection with the main vascular system of the rhizome,
compared with that which connects functional buds to the main rhizome
system, suggests that these obscure buds are vestigial. If propagation by
seed is not vigorous the survival potential of Croomia must be low. This
may well account for the relict status of the genus as indicated by its re-
stricted and markedly disjunct distribution.

FLower. The morphology of the ovary calls for discussion since there
seems to be lack of agreement in interpreting its structure in the related
Stemona (cf. Baillon (1860) and Lachner-Sandoval (1892) on the one
hand with Swamy (1964) on the other). It might be unwise to compare
the ovary of Croomia, which has apical placentation, with Stemona which
has basal placentation, but there are some notable anatomical similarities.
It is unfortunate that Stichoneuron (with apical placentation) has to be
excluded from this discussion but we have no comparable information
about this genus.

Both Swamy and Baillon studied Stemona tuberosa Lour., Baillon using
the name Roxburghia gloriosoides by which it was earlier known (Burkill,
1960). Lachner-Sandoval studied mainly Stemona javanica (as Rox-
burghia javanica), which has much smaller flowers, but he also investigated
Stemona tuberosa (referred to as Roxburghia gloriosa) and does not com-
ment on any major differences. The two early authors interpreted the
carpel of Stemona as unicarpellate largely on developmental evidence. It
originates as a single crescent-shaped protuberance opposite the first-
formed tepal. This cushion of tissue grows in such a manner as to enclose
the basal placenta. Swamy unfortunately had no access to this earlier
information so his analysis based simply on the anatomy of the mature
ovary is entirely original. He interpreted the ovary of Stemona tuberosa
as bicarpellate because the ovules (up to 10) are in 2 rows. There are two
identical vascular bundles in the ovary wall, one opposite each of the outer
tepals. These Swamy regards as the dorsal bundles of 2 carpels, the ventral
bundles being represented by the vascular supply direct to the basal ovules.
In Stemona javanica according to Lachner-Sandoval, the two bundles in the

1968 | TOMLINSON & AYENSU, CROOMIA 273

ovary wall are dissimilar, one bundle being double. This difference is most
obvious in the dehiscing capsule, the line of dehiscence passing through
the single bundle but between the double one. Croomia corresponds to
Stemona in having 2 ovary bundles, one opposite each outer tepal. How-
ever, there is no indication of a bicarpellate condition since there is no
reason to interpret these bundles other than as a dorsal and a ventral. Also
the way in which the ovary is closed in the stigmatic region suggests a
manner of development comparable to that described for Stemona by
Baillon, although this speculation has not yet been verified. However, the
great difficulty in comparing the ovaries of these two genera is the marked
difference in placentation and consequently in placental vascular supply.
In the absence of extensive comparative and developmental studies we must
confess that the interpretation of the ovary of the Stemonaceae still re-
mains open.

TABLE 1. Comparison of the three genera of Stemonaceae

CROOMIA STEMONA STICHONEURON
Hasit
General erect herb scandent low shrub
Leaf arrangement distichous distichous, distichous
spiral or whorled
Longitudinal veins diverging from base diverging from some diverging
of lamina base from midvein
Free vein-endings _ present rarely present absent
in lamina
Rhizome extensive not extensive ? not extensive ?
ANATOMY or
Hairs absent absent uniseriate,
thick-walled
Cauline and common not distinct distinct distinct

bundles in stem
internode

Crystals no raphides raphides and raphides
styloids
Vessels in stem absent present present r.
Sieve tubes in stem _ sieve plates sieve plates sieve plates
simple compoun

REPRODUCTIVE PARTS 4
Cincinni few, many-flowered many-flowere:

few-flowered cette
(if a cincinnus)

or flow
olitary
Stamens not appendaged appendaged not ——
Pollen sculpture smooth eer :
vary unicarpellate disputed pee
Placentation apica basal apical :
Aril filamentous vesicular i

274 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 49

Taxonomy. These morphological problems are compounded because
in dealing with the three genera of the Stemonaceae it is not evident that
they form a natural assemblage. Similarities between the genera are few
but differences are many. The following features are common to them and
diagnose the Stemonaceae. Flowers are somewhat perigynous and have: —
two pairs of sepaloid tepals, the outer pair at right angles to the inner; four
stamens in a single whorl, one stamen opposite each tepal; a unilocular
ovary with a short stigma. The ovules are anatropous, few to several, each
with two integuments. The seeds are endospermous, ribbed and conspic-
uously arillate. The fruit is a 2-valved capsule. Leaves have a narrow
insertion (not broadly encircling the stem) and a fairly characteristic
cordate base. Stomata are anomocytic. The epidermis of the lamina has
sinuous anticlinal walls. Some of the vascular bundles in the axis are
amphivasal.

More detailed examination reveals marked differences which are set
out, insofar as they are available, in Taste 1. Even with this limited
evidence it is clear that we are not dealing with a homogeneous assemblage.
At this stage it would be presumptuous, therefore, to make any statements
about the phylogenetic position of the Stemonaceae if it is suspected that
they are an artificial group. A prior requisite to a correct understanding
of their taxonomic status is the accumulation of a great deal more factual
knowledge not only about them, but about their putative relatives.

SUMMARY

Shoots in Croomia are made up of units which branch sympodially.
Each unit consists of a long-persistent rhizomatous portion and an annual
erect, leafy shoot. The number of organs and their sequence, at least on
the underground parts of the axis, is fixed within narrow limits. Only two
functional vegetative buds are developed on the rhizome, a renewal bud
which grows out immediately and a dormant bud which provides the only
method of branching for the rhizome, although it is initially strongly in-
hibited. The anatomy of the vegetative organs and the flower is described.
The simple vasculature of the ovary suggests that it is a single carpel
despite its peculiar apical placentation. A summary of anatomical and
morphological evidence suggests that Croomia, Stemona, and Stichoneuron
constitute a rather heterogeneous assemblage. )

ACKNOWLEDGMENTS

We are indebted to Miss Priscilla Fawcett, Botanical Illustrator at Fair-
child Tropical Garden for the preparation of the morphological illustra-
tions which make Plates I and II. Miss Mary Gregory, Bibliographer at
the Jodrell Laboratory, Royal Botanic Gardens, Kew, was kind enough to
supply us with bibliographic information about the Stemonaceae. Miss
Anne Bellenger is thanked for the series of preparations on which the
analysis of vascular systems is based.

1968 | TOMLINSON & AYENSU, CROOMIA 275

LITERATURE CITED
Barton, H. 1860. Organogénie florale des Roxburghiées. Adansonia 1: 245-
250.

appre I. H. 1960. Organography and speeds of Dioscoreaceae, the family
the e yams. Jour. Linn. Soc. Bot. 56: 319-412.
Gas. A. 1859. On the genus Croomia, ne its place in the natural system.
Mem. Am. Acad. IT. 6: 453-457.
Harper, R. os 1942. Croomia a member of the Appalachian flora. Castanea
7: 109-
Hom, T. el Croomia pauciflora Torr. An anatomical study. Am. Jour. Sci.
IV. 20: 50-54.
1927. Sciaphilous plant-types. Beih. Bot. Centralbl. 44: 1-8
Hutcuinson, J. 1959. Families of flowering plants. vol. 2. Mon
E Oxford University Press, Oxfor
LACHNER-SANDOVAL, V. 1892. Beitrag zur Kenntniss der Gattung Roxburghia.
se Centralbl. 50: 65-70, 97-104, 129-135.
Swamy, B. G. L. 1964. Observations on the floral morphology and embryology
ae Stemasa ee Lour. Phytomorphology 14: 458-468.
WV. Battey. 1950. Sarcandra, a vesselless genus of the Chlorantha-
ceae. Jour: Arnold Arb. 31: 117-129.

FAIRCHILD TROPICAL GARDEN SMITHSONIAN INSTITUTION
10901 OLp CuTLER Roap DIVISION oF PLANT ANATOMY
MramtI, Fioriwa 33156 WasHIncTON, D.C. 20560

AND

HARVARD UNIVERSITY
CasBot FOUNDATION
PETERSHAM, MASSACHUSETTS 01366

EXPLANATION OF PLATES
PLATE i

’ hizome to
a plant in ere Apel %< 1/2. Scar of previous year’s erect shoot peat
left. Fic. 36. Leafy shoot from above, X 1/2. Fic. 37. Details of base of erect

tional buds, cross doiched: Fic. 39. Details of node B with scale-leaf a
to show dormant bud, x 3. Letters A-F indicate successive nodes on early pa
of growth unit.

PLATE II : dif
Fics, 40-48, Croomia pauciflora, flowers. Fics. 40-43. Open flowers in dif-
ferent aspects, x 4. oh . 40. Obliquely from sieve Fic. 41. Lateral bo
. 42. Same, with one stamen cut off to show ovary. Insets: surface a
lea age aa adusial — of tepal (right), to show ge ap Le Floeal
Fic. 43. Median section at 45° srg plane of ovary ae a
dupes. Fic. 45. Pollen ae oe hogs 46. 2-flowered cincinnus, pane
Fics. 47 and 48. Diagrams of ke s of flowers. Fic. 47 rome
section of flower constructed from pron tos at different ajeae ears
bundles in solid black. Fic. 48. Ovary as seen in cleared flower, only ¢ ovu
show

Jour. ARNOLD Ars. VOL. 49 PiaTE I

TOMLINSON & AYENSU, CROOMIA

Jour. ARNOLD Ars. VoL. 49 Prate IT

* WO
Ree
Yerctcro €

47

Tomiinson & AYENSU, CROOMIA

278 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

HERISSANTIA, BOGENHARDIA, AND GAYOIDES (MALVACEAE)

GerorcE K. Brizicky

In 1903, J. K. Small raised the section [“subgenus”| Gayoides A. Gray
of Adutilon Mill. to the rank of genus, with Gayoides crispum (L.) Small
(Abutilon crispum (L.) Medic., Sida crispa L.) as the type (FI. Southeast.
U.S. 764, 1335). Subsequently, Gayoides has been adopted and used by
many authors for this segregate of Abutilon. In 1954, Kearney checked
the availability of two older names, Herissantia Medic. and Bogenhardia
Reichenb., for this genus. He found (Leafl. West. Bot. 7: 119, 120. 1954)
Herissontia (Medicus, Philosophische Botanik 1: 90. 1789) to be a nomen

nudum, but Bogenhardia (Reichenbach, Repertorium Herbarii 1: 200.
1841; 2: 48. 1841) to be validly published and hence much earlier than
Gayoides. Consequently, Kearney transferred Gayoides crispum into Bo-
genhardia as B. crispa (L.) Kearney.

There is, however, an earlier work by Medicus, “Kurzer Umriss einer
systematischen Beschreibung der mannigfaltigen Umhiillungen der Saa-
men” (Vorles. Churpfalz. Phys.-Oekon. Ges. 4(1): 244. 1788), in which
Herissantia was published with the following description: “224. HERIS-
SANTIA (Sida crispa. Cavanil. Dissertatio prima, num. 65). Dem aussern
Ansehen nach sieht die Saamenkapsel ginzlich jener von Abutilon gleich,
unterscheidet sich aber von derselben merkwiirdig dadurch, dass bei ganz-
licher Zeitigung keine einzelne Saamenkapseln da sind, sondern alle Scha-
alen bis unten ginzlich gedffnet, und nur wo sie in der Mitte gemeinschaft-
lich zusammenstossen, miteindander verwachsen sind.” Although the de-
scription is rather brief and only refers to the fruit, it indicates the main
character distinguishing Herissantia from Abutilon and certainly merits
recognition as a formal generic description no less than Endlicher’s diag-
nosis (Gen. Pl. 986. 1840) of the subdivision Gayoides of Bastardia HBK.
on which Reichenbach’s Bogenhardia was based. Moreover, an additional
Latin diagnosis of Herissantia is found interpolated in Medicus’s system of
or key to the fruit types (loc. cit. 369), namely: “‘G. [Capsulae] plures,
receptaculo communi affixae, suntque ... a. 3. multivalves 224. Heris-
santia.’’ Thus, Herissantia appears to be a validly published name which
is to be taken up in place of the later Bogenhardia (1841). Since this neo-
tropical genus of at most three species (see H. Monteiro, Anais Soc. Bot.
Brasil 5: 438-444. 1956) is economically unimportant, there is no reason
at all to propose for conservation either Gayoides or Bogenhardia. For
convenience I present below the correct name for this segregate of Abutilon,
with its synonyms, the type species common to all, and the necessary new
combinations.

1968 | BRIZICKY, MALVACEAE 279
Herissantia Medic. Vorles. Churpfilz. Phys.-Oekon. Ges. 4(1): 244

Bastardia HBK. b. Gayoides Endl. Gen. Pl. 986. 1840

Bogenhardia Reichenb, Repert. Herb. 1: 200. 1841; 3 48. oa nom, illeg.
Abutilon sect. Gayoides (Endl.) A. Gray, Gen. Pl. U. 5. 2:6 1849,

Gayoides Small, Fl. Southeast. U.S. 764, 1335. 1903, nom. ty
Pseudobastardia Hassler, Bull. Soc. Bot. Genéve II. 1: 209, 1909, nom. illeg.

Type species: “Herissantia crispa (L.) Brizicky, comb. nov. (Sida
crispa L. Sp. Pl. 2: 685. 1753; Abutilon cryspum (L.) Medic. Kiinstl.
Geschl. Malven-Fam. 29. 1787; Gayoides crispum (L.) Small, Fl. South-
east. U.S. 764. 1903; Bogenhardia crispa (L.) Kearney, Leafl. West. Bot.
7: 120. 1954).

The correct names for two Brazilian species recognized by Monteiro
(loc. cit.) are the following:

Herissantia tiubae (K. Schum.) Brizicky, comb. nov. (Abutilon tiubae
K. Schum. im Mart. Fl. Brasil. 12(3): 381. 1891; Bogenhardia tiubae H.
Monteiro, Anais Soc. Bot. Brasil 5: 440. 1956).

Herissantia nemoralis (St.-Hil. et al.) Brizicky, comb, nov. (Bas-
tardia nemoralis St.-Hil. et al., Fl. Brasil. Merid. 1: 195. 1827; Bogen-
hardia nemoralis (A. Juss.) H. Monteiro, loc. cit. 442.

Derivation of the generic name was not indicated by Medicus. There is
little doubt, however, that the name commemorates Louis Antoine Pro-
spére Hérissant, 1745-1769, a French physician, naturalist, and poet, the
author of the Bibliotheque physique de la France ou Liste de tous les
ouvrages, tant imprimés que manuscrits, qui traitent de EBistotre nat-
urelle de ce Royaume; avec des notes critiques & historiques (8°. 496
pp. Paris), published posthumously in 1771. Besides this rather well-
known bibliographic work, Heérissant’s other, apparently very rare, pub-
lication, Jardin des curieux, ou catalogue raisonné des plantes les plus
belles et les plus rare avec leur culture (12°. Faris) appeared in the same
year. (Cf.: A. Haller, Bibliot. Bot. 2: 611. 1772; G. R. Boehmer, Bibliot.
Script. Hist. Nat. Oekon. 3(1) [III, a] : 719. 1787. )

280 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

JAMAICAN AND OTHER SPECIES OF BUMELIA (SAPOTACEAE)

WILLIAM T. STEARN

Tue GENUS Bumelia, with which Dipholis is here united, comprises about
forty species, all American, of which nine occur in Jamaica. They have
been the subject of a careful, detailed, very useful revision by Cronquist
in the Journal of the Arnold Arboretum 26: 435-471. 1945, which provides
keys, descriptions, and synonymy, the last very extensive owing to insuf-
ficient allowance by many authors for variation within speci

The following notes arising out of a study of the ees for the
Flora of Jamaica supplement Cronquist’s work.

In Jamaica these trees are often known under the name of “Bullet,”
formerly “Bully-tree,” the ripe fruits of some species recalling to the early
colon‘sts those of the European bullace (Prunus instititia).

THE IDENTITY OF MAUROCENIA AMERICANA

In 1754, Philip Miller maintained the genus Maurocenia (in his Garden-
ers Dictionary Abridged, 4th ed.) as distinct from Cassine, with which
Linnaeus had united it, observing that “in this he is mistaken,” and thus
gave the name Maurocenia valid post-1753 publication. In 1768 (in the
8th edition of the Gardeners Dictionary), Miller included four species in
the genus. The first, Maurocenia frangula Miller (Cassine maurocenia L..,
1753), the lectotype of the genus, belongs to the Celastraceae. This and
Miller’s two other species, M. phillyrea Miller (i.e., Cassine peragua L.
sensu stricto, syn. C. capensis L.) and M. cerasus * Miller (possibly an
Euclea) were all from South Africa. His fourth species, M. americana,
however, came from Jamaica and, although not hitherto placed, it is
evidently not congeneric with the African members of the genus. Miller’s
account is as follows:

4. MAUROCENIA Beagles Pra obverse ovatis emarginatis, floribus
solitariis alaribus. Maurocenia bverse oval leaves which are indented at
the edges, and flowers growing oe from the id of the branches. Frangula
folio subrotundo rigido subtus ferrugineo. Houst. MSS. gee Alder
with a rounded stiff leaf, which is of an iron Stee on the under s

The fourth sort was discovered by the late Dr. Houstoun, ciein naturally
at the Palisadoes in Jamaica, from whence he sent the seeds to Europe; this
rises with a woody stalk from fifteen to twenty feet high, covered with a rough
brown bark, and divides into many branches, which are garnished with stiff
leaves, placed alternately; they are about an inch and a half long, and a little
more in breadth, indented at the top, with a stiff reflexed border, of a gray
colour on their upper side, but of a rusty iron colour on their under, standing
upon short foot-stalks. The flowers come out singly along the side of the

1968 | STEARN, BUMELIA 281

Type specimen of Maurocenia americana

«Fae! 4,
(Miller) Stearn) in herb. British Museum (Nat. Hist.).

Miller (Bumelia americana
t.)

Ss permanent. The germen afterwards turns to a round berry, with one or two
cells, each having one oblong seed.

282 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

The Palisadoes is a coastal strip of land enclosing Kingston Harbour
with Port Royal at its tip. Houstoun’s specimen (Fic. 1) from the Palisa-
does is in Miller’s herbarium, now incorporated in the general herbarium
of the British Museum (Natural History), London; it lacks flowers but
undoubtedly belongs to Bumelia retusa Swartz, the lectotype of the genus
Bumelia (Sapotaceae), which is common on the coasts of the Bahamas,
Jamaica, and Cuba. This varies much in the size, shape, pubescence, and
veining of the leaves. The leaves of Houstoun’s specimen agree essentially
with those of B. retusa from southern Jamaica, e.g., Proctor 17427 from
Little Goat Island, St. Catherine, as well as from elsewhere, e.g., Stearn
314 from St. Ann, Proctor & Stearn 11762 from Trelawny. Although the
fruits are usually solitary in the axils, the flowers are rarely so and are
usually in sessile axillary fascicles of 2 to 15. In consequence of the
identity of Miller’s species with Swartz’s, Miller’s epithet must be trans-
ferred to Bumelia and replace Swartz’s.

Bumelia americana (Miller) Stearn, comb. nov.
Maurocenia americana Miller, Gard. Dict. ed. 8, art. Maurocenia no. 4. 1768.

Bumelia retusa Swartz, Prodr. 49. 1788; Fl. Ind. Occ. 1: 490. 1797. Cronquist,
Jour. Arnold Arb. 26: 460. 1945.

For further synonymy, see Cronquist, loc. cit.

THE JAMAICAN SPECIES OF BUMELIA

After stating that Bumelia Swartz (1788) and Dipholis A. DC. (1884),
showed “‘no real diversity in structure or habit” and that the presence of
albumen in Dipholis and its absence in Bumelia have no practical impor-
tance owing to the rarity of fruiting specimens, Hartog in 1879 (Jour. Bot.
(London) 17: 356) accordingly proposed to abolish Dipholis even as a
section of Bumelia. Nevertheless most other authors have continued to
maintain them, by attributing scanty or no albumen and fleshy cotyledons
to Bumelia and abundant albumen and thin cotyledons to Dipholis.

Aubréville (Adansonia, Mém. 1: 78. 1965), for example, keys them out
as:

Graine sans albumehi: (2 0 es Bumelia
Graine avec AIDUINOR a. Diphoks

The seeds are, however, unknown for many species of this group, and it
is thus uncertain whether such species really possess the seed characters of
the genera in which they have been placed. Despite certain tendencies
noted by Cronquist (Jour. Arnold Arb. 26: 445. 1945), e.g. “plant com-
monly spiny or thorny” in Bumelia and “plant unarmed” in Dipholis,
“ovary usually hairy, occasionally glabrous” in Bumelia and ‘“‘ovary nearly
always glabrous” in Dipholis, the “fruit commonly ... rounded . . .
at the apex” in Bumelia and the “fruit commonly abruptly tapering to the
style” in Dipholis, there are no constant correlations between these char-

1968] STEARN, BUMELIA 283

acters and those of the seed. Hence it is difficult and arbitrary to recog-
nize two groups. Baehni in his first memoir on the Sapotaceae (Candollea
7: 434. 1938) referred Dipholis to Bumelia. In his posthumous survey of
the genera (Boissiera 11: 133-136. 1965) he concluded that “en somme
leur séparation aurait plutét comme base une croyance, un mythe,” the
amount of albumen and thickness of the cotyledons being a relative and
not absolute distinction. No good purpose is served in trying to keep them
separate.

The Jamaican species of Bumelia, including those formerly placed in
Dipholis, are as follows:
1. Calyx ca. 2-2.5(-3) mm. long. Flowers usually many (3-15) in a cluster.

2. Petiole 15-32 mm. long. Leaf-blades often more than 8 cm. long and 4
cm. broad; lateral veins clearly evident. Fruits large (12-15 mm. in di-
ameter), subgl ODO8B e665 Fo nanan gee ines eee B. nigra.

2. Petiole 2-15 mm. long. Leaf-blades rarely up to 8 cm. long and 4 cm.
broad; lateral veins inconspicuous. Fruits smaller (4-10 mm. in diameter).
3 Leaf- blades acute or shortly acuminate, mostly narrowly elliptic.

Fruits. ca:S sani domme! econ Fae a a B. salicifolia.

3. Leaf-blades obtuse or rounded, sometimes emarginate at apex, mostly
almost orbicular, rotund or broadly elliptic, varying to obovate or
oblanceolate. Fruits ca. 8-10 mm. long.

4. Young branchlets, pedicels, and sepals glabrous. Leaves green
beneath when young. Fruits oblong-ellipsoid, ca. 4-5 mm. in
diametét,” 0 aS Se ee B. rotundifolia.

4. Young branchlets, pedicels, oy sepals minutely hairy. Leaves
brown-tomentose beneath when young. Fruits subglobose to broad-
ly ellipsoid, ca. 10 mm. in fhesice: Ife ee B. americana.

1, Calyx (3—)3.5—4 mm. long. Flowers usually few (1-8) in a cluster.
5. Sepals 8-9, the outer as well as the inner brown-tomentose.
persistently hairy, Central Jamaica: 0. 22.20. B. octos
5. Sepals 5-8, the outer 2 glabrous or sparsely hairy, the inner tomentose or
sericeous. Pedicels glabrous or sparingly —
6. Leaf-blades not more than 5 cm. broa
7. Leaves without raised network of veins beneath. ele ona s
rf Leaves with raised network of veins beneath. — Cruz vty;
sp. nov. I.
SHS ek wean eee
6. Leaf-blades some or all more than 5 cm. “spar aoe raised ‘network
of veins beneath immediately visible under
8. Petiole 1-10 mm. long. Leaf-blades Coreg or broadly es
Basteti: Jaisaica: 23 hk ee a ee B. bullata,
8. Petiole 10-20 mm. long. Leaf-blades elliptic-oblong. oa
SSI Oe ee Bs :

Bumelia americana (Miller) Stearn, comb. nov. See above.
B. americana subsp. americana Nie. 1+ Fic, 2, G-K; Exc. a2 1,
Bumelia retusa subsp. typica Cronquist, Jour. Arnold Arb. 26: 460. 1945.
Bahamas, Cuba, Jamaica, Navassa. See Map 1.

284 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Ses eve ater"

Nv.

Miles
$2 518 tg 30

30" = 30" 77° 30) ee

Map 1. Distribution of Bumelia americana in Jamaica.

B. americana subsp. neglecta (Cronquist) Stearn, comb. nov.
Bumelia retusa subsp. neglecta Cronquist, Jour. Arnold Arb. 26: 461. 1945.
Mexico, British Honduras.

Bumelia bullata (Howard & Proctor) Stearn, comb. nov.
Dipholis bullata Howard & Proctor, Jour. Arnold Arb. 39: 103. 1958.
Jamaica: Parish of Portland.

ee montana Swartz, Prodr. 49. 1788; Fl. Ind. Occ. 1: 493. ¢. 8,
fig. e-n. 1797. Fic. 3, A-D, J-
Dipl montana (Swartz) Griseb. Fl. Brit. W. Indian Is. 401. 1861. Pierre
an in Urban, Symb. Antill. 5: 135. 1904. Cronquist, Jour. Arnold Arb.

26: 442. 1945

Jamaica: Parishes of St. Andrew, Portland, St. Thomas.

Bumelia nigra Swartz, Prodr. 49. 1788; Fl. Ind. Occ. 1: 487. ¢. 8, fig.
a—d. 1797. Fic. 3, F-H.
Dipholis nigra (Swartz) Griseb. Fl. Brit. W. Indian Is. 400. 1861. Pierre &
Urban in Urban, Symb. Antill. 5: 137. 1904. Cronquist, Jour. Arnold Arb.
26: 439. 1945.
Jamaica: Parishes of bpritanities Adapted. St. Elizabeth, Man-
chester, Clarendon, St. Ann, St. Andre

Bumelia octosepala (Urban) Stearn, comb. nov. Fic.3, ©.
sn} octosepala ie Baa Antill. 7: 324. 1912. Cronquist, Jour.
rnold Arb. 26: 442

a Parishes of Gace Clarendon, St. Ann, St. Catherine.

Bumelia rotundifolia Swartz, Prodr. 50. 1788; Fl. Ind. Occ. 1: 495.

1968] STEARN, BUMELIA 285

1797. Pierre & Urban in Urban, seks Antill. 5: 143. adic g Cron-

quist, Jour. Arnold Arb, 26: 459, 1945. Fic. 2, A-F.

Jamaica: Parishes of Hanover, ee Manchester, St. me Claren-
don, St. Catherine, St. Andrew.

Bumelia salicifolia (L.) Swartz, Prodr. 50. 1788; Fl. Ind. Occ. 1: 491.
1797
Achras salicifolia L. Sp. Pl. ed. 2. 1: 470. 1762.
Dipholis salicifolia (L.) A. DC. in DC. Prodr. 8: 188. 1844. Sargent, Silva
. Am. 5: 179. t. 250, 1893. Pierre & Urban in Urban, Symb. Antill. 5: 138.
1904. incite Jour. Arnold Arb. 26: 440. 1945.

Florida, Bahamas, Jamaica (Parishes of Westmoreland, St. James, St.
Elizabeth, Trelawney, Clarendon, St. Ann, St. Catherine, St. Mary, St.
Andrew), Cuba, Hispaniola, Puerto Rico, Virgin Islands, Guadaloupe,
Barbados, Central America.

Bumelia, sp. nov. I.
Jamaica: Parish of St. Elizabeth; Santa Cruz Mountains, Malvern,
2,200’, ix. 1907, Harris 9742; same locality, 2,100’, ix. 1907, Harris 9803.
This is closely related to Bumelia montana of the Blue Mountains,
eastern Jamaica, but seems distinct in the more evident veining of the
leaves and slightly smaller calyx. Complete flowers and fruits are unknown.

Bumelia, sp. nov. II.

Jamaica: Parish of Hanover; interior summit slopes of Dolphin Head,
limestone ridge forest, 1,100’ — 1,700’, iv. 1955, Proctor 10039.

This species is notable for its large leaves with elliptic blades 5-18 cm.
long, 2.5—8 cm. broad. Complete flowers and fruits are unknown.

OTHER SPECIES

The following species distinguished by Cronquist or by Standley and
Williams under Dipholis do not occur in Jamaica

Bumelia bellonis (Urban) Stearn, comb. nov.
Dipholis bellonis Urban, Symb. Antill. 5: 137. 1904. Cronquist, Jour. Arnold
Arb. 26: 4

Puerto Rico.

Bumelia cubensis Griseb. Cat. Pl. Cub. 164. 1866.
sascenel cubensis (Griseb.) Pierre in ot Symb. Antill. 5: 140. 1904.
ist, Jour. Arnold Arb. 26: 443.

See

286 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

yy)
GLI LTP

SS

ICME if

Fic. 2. A-F, Bumelia rotundifolia, A, flowering shoot (Norman 158) X 3/3;
B, fruiting shoot (Webster & Wilson 4928), X 3/5; C, flower (Norman 158),
X 6; D, dissected corolla (Norman 158), X 6; E, gynoecium (Norman 158),

1968 | STEARN, BUMELIA 287

Bumelia durifolia (Standley) Stearn, comb. nov.
i ee Standley, Carnegie Inst. Wash. Misc. vee iit = Maya
ea 4): 78. 1935. Cronquist, Jour. Arnold Arb. 26: 441.

British osaneas

Bumelia ferruginea (Ekman & O. C. Schmidt) Stearn, comb. nov.
me an ferruginea Ekman & O. C. Schmidt, oe Repert. Sp. Nov. 32: 94.
1933. Cronquist, Jour. Arnold Arb. 26: 444. 1945.

Hispaniola: Dominican Republic.

Bumelia jubilla (Urban) Stearn, comb. nov.
Dipholis jubilla Ekman ex Urban, Symb. Antill. 9: 415. 1925. Cronquist,
Jour. Arnold Arb. 26: 439. 1945.

Cuba.

Bumelia matudae (C. Lundell) Stearn, comb. nov.
Sideroxylon matudae C. Lundell, Phytologia 1: 221. 1937 (as matudai).
Sideroxylon steyermarkii Standley, Field Mus. Publ. Bot. 22: 368. 1940.
Dipholis matudae (C. Lundell) C. Lundell, Contr. Univ. Michigan Herb. 7:
1942 (as matudai); Standley & L. O. Williams in Fieldiana, Bot. 24:
220. 1967 (as matudae).

Mexico, Guatemala.

Cronquist included this in Bumelia minutiflora, from which it differs,
according to Standley and Williams, in having much larger and differently
veined leaves. The genitive of Matuda being matudae, like balansae from
Balansa (cf. Int. Code Bot. Nomencl. 1966, Art. 73, Rec. 73C), Lundell’s
original spelling matudai is to be treated as an orthographic error.

Bumelia minutiflora (Pittier) Baehni, Boissiera 11: 135. 1965.
Dipholis minutiflora Pittier, Contr. U.S. Natl. Herb. 13: 464. 1912. Cronquist,
Jour. Arnold Arb. 26: 438. 1945
Mexico, Guatemala, Honduras, Costa Rica, Panama.

Bumelia sp.
oe dishes or Standley, fet Mus. Publ. Bot. 18: 909. 1938. Cronquist,
old Arb. 26: 442.
es of Standley’s ha parities to Bumelia is precluded by

; [F] unlettered figure in lower right corner, leaf and inflorescence gy Tip tati
11 O40, isotype of B. peckhamensis), X 3/5. G-K, Bumelia americana

, X 3. Drawing by aaa a C. Webb, from specimens in British Museum
(lena Eaory), Londo

288

JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

ICME

Bumelia montana. A, flowering shoot (Proctor 9538), X

D5,
55 oy ‘se (Proctor 9538), X 4; C, dissected corolla (Proctor 9538), X

, Zynoecium in section (Proctor 9538): J, fruiting shoot (Harris 10118),

1968 | STEARN, BUMELIA 289

Bumelia parvifolia A. DC. in DC. Prodr. 8: 190. 1844. Since this species
was described by Standley from specimens without flowers, it would seem
prudent to await more complete material from the type-locality, Los
eg prov. Guanacaste, Costa Rica, before providing it with a new
name

Bumelia repens (Urban & Ekman) Stearn, comb. nov.
Dipholis repens Urban & Ekman, Ark. Bot. 22A. 17: 70. 1929. Cronquist,
Jour. Arnold Arb. 26: 444. 1945.

Hispaniola: Dominican Republic.

Bumelia sericea (Cronquist) Stearn, comb. nov.
Dipholis sericea Cronquist, Jour. Arnold Arb. 26: 444. 1945.

Hispaniola: Dominican Republic.

Bumelia stevensonii (Standley) Stearn, comb. nov.
Dipholis stevensonii Standley, Trop. Woods 11: 21. 1927. Cronquist, Jour.
Arnold Arb. 26: 438. 1945.
British Honduras.

DEPARTMENT OF BOTANY
BritisH Museum (NATURAL History)
CROMWELL Roap
Lonpon, S.W. 7, ENGLAND

xX 2 fs. 4,
nigra. F, fruits oe. 20591), 3/5; G, seed (Proctor 20591), x 15.8,
fruit in transverse section showing Speen embedded in P

and scanty pulp, X 3/5. I, Bumelia americana, fruit = sea ect - ow-
ing fleshy cotyledons, without endosperm, and abundan

Joanna C. Webb, from speci in British Museum (Neturad History), “London,

VoLuME 49 at ' NuMBER 3

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OF THE

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CONTENTS OF NO. 3

ANATOMY OF THE PALM RHAPIS EXCELSA, V. INFLORESCENCE. P. B.
Tomlinson and M. H. Zimmermann

ANATOMY OF THE PaLM RuaPis ExceLsa, VI. Roor anp BRANCH
Insertion. P. B. Tomlinson and M. H. Zimmermann ............
Comparative MorpHotocicaL Srupres 1n Driuentaceas, III.
‘Tue Carpets. William C. Dickison

REVISION OF THE GENUS NoreLara (Onzacean). P. S. Green ..

CoMPARATIVE Anatomy OF THE LEAF-BEARING CacTACEAR, XVII.
PRELIMINARY OBSERVATIONS ON THE PROBLEM OF TRANSITIONS
From Broap To TeRere Leaves. I. W. Bailey

BoranicaL NaME oF ‘THE tae pee or Cuina. Marshall
C. Johnston ...

Volume 49, wi 2, includi |

g pages 167-290,

‘was s issued = 1968

291

377

JOURNAL

OF THE
ARNOLD ARBORETUM

VoL. 49 Jury 1968 NUMBER 3

ANATOMY OF THE PALM RHAPIS EXCELSA, V.
INFLORESCENCE

P. B. ToMLINSON ! AND M. H. ZIMMERMANN

PREVIOUS ARTICLES IN THIS SERIES on the anatomy of Rhapis have de-
scribed the construction of aérial and underground (rhizomatous) axes and
something of the developmental principles underlying this construction
(Zimmermann & Tomlinson, 1965, 1967a; Tomlinson & Zimmermann,
1966a & b). The vascular system thus revealed has also demonstrated a
continuous pathway for long-distance transport. Further, the organiza-
tion of newly developed vascular strands in the apex shows how new leaves
are continuously attached to the axial system.

For further appreciation of vascular continuity an understanding of the
method of attachment of lateral organs to the main axis has been sought.
The present article describes the attachment of the inflorescence axis to
the aérial stem and something of the attachment of lateral branches of
various orders to the main inflorescence axis. A description of the attach-
ment of flowers to rachilla is reserved for a future article dealing with floral
anatomy.

To complete a full understanding of the vascular pathways throughout
Rhapis the attachment of roots and vegetative branches (rhizomatous
suckers) still remains to be described. This will be done in another paper
of this series whereupon a general discussion of developmental principles
involved in branch-axis union in monocotyledons will be permitted. It is
remarkable how little this aspect of monocotyledonous anatomy has
attracted the recent attention of botanists despite its obvious and funda-
mental physiological significance. To the question “Is there, in palms, any
vascular connection between the lateral inflorescence and the foliage leaf
which subtends it?” there has hitherto been no answer, although it is of
obvious significance to tropical agronomists. This present article gives
some indication of a satisfactory answer.

The only previous work on this topic is by the indefatigable von Mohl
(1824) who illustrated various kinds of union without showing anatomical
detail. Micheels (1892) has described the anatomy of the ultimate, fruit-

? Research on the anatomy of palms by P. B. Tomlinson is supported by National
Science Foundation Grant GB-2991 and its continuation GB-5762-X.

292 JOURNAL OF THE ARNOLD ARBORETUM [ VoL. 49

bearing axis of the inflorescence in 32 species from 19 genera of palms with
a discussion restricted to the mechanical significance of the tissues he
observed. Otherwise the anatomy of the inflorescence axis in palms seems
never to have been examined.

The general features of flowering in palms is a subject scarcely con-
sidered in botanical textbooks, Palm inflorescences are seemingly complex
organs and they are not normally available in herbaria, except as fragments.
Constructional principles, however, are straightforward even though the
multiplicity and size of branches tends to obscure this. The recent account
by Tomlinson and Moore (1968) of inflorescence in Nannorrhops has
indicated some of the essential features of the reproductive axis in palms
and it is against this background of understanding that the inflorescence in
Rhapis is described.

MATERIAL AND METHODS

Microscopic analyses of vascular systems are based on the cinemato-
graphic methods which have been described in detail in previous articles
(Zimmermann & Tomlinson, 1965, 1966, 1967b). Inflorescence insertion
was studied in several continuous series of sections cut through nodes
bearing fully developed inflorescences. For the branching of the inflores-
cence itself, serial sections along its main axis were studied together with
a similar series along one first-order branch.

INFLORESCENCE MORPHOLOGY

Development of lateral inflorescences in the reproductive phase is irreg-
ular in Rhapis. Potentially there is an inflorescence bud in the axil of each

cences at intervening nodes are present as small, scale-like flaps of tissue
in the leaf axils. The vascular system of “satellite” bundles associated

(Zimmermann & Tomlinson, 1965). The presence of a vestigial vascular
supply to aborted inflorescences is of considerable developmental signifi-

inflorescences by their more numerous, densely crowded, smaller flowers
on more slender and more elaborately branched axes (cf. TaBLe 1 and
Fics. 11 & 12). It is not known if an erect aérial shoot continues to bear
inflorescences of one kind.

omparative study of inflorescences in Coryphoid palms shows that

1968]

FIG. | , \

FIRST-ORDER BRANCH

BRACT

FIRST-ORDER
BRANCH

op
Ye’
v c/
INFLORESCENCE P
o@ “SRACHILLA

/ ()
yy
canker < Q2ONNND785
JSS SSsS se
y <i
ONS Te
~ hy
Vg,

PROPHYLL

SECOND-ORDER BRANCH

We
J, —

SUBTENDING FOLIAGE LEAF
_Fic. 1. Diagram of construction of Rhapis inflor
discontinuous lines i

escence. Branches ending in
are not completed. Adnation between branch and parent
axis is shown with dotted lines.

les of inflorescence construction in palms, as
revealed by the detailed analysis of Nannorrhops by Tomlinson and Moore
(1968), are clearly demonstrated in Rhapis

TOMLINSON & ZIMMERMANN, RHAPIS EXCELSA

294 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

\ Gs

inflorescence. Fic. 4. Pro opt “(Grst ate hee dorsal side. Fic Same
from ventral side. Fic. 6. Second bract. Fic. 7. Third bract. Fic. 8. " ‘Xbruptly
reduced fourth bract.

pronounced Segregation of the sexes, Rhapis obviously differs from Nan-
in

Rhapis is strictly a partial inflorescence, equivalent to one first-order branch
unit in Nannorrhops but with fewer degrees of branching. A further im-
portant difference is that ultimate branch units in Nannorrhops are con-

1968 | TOMLINSON & ZIMMERMANN, RHAPIS EXCELSA 295

densed few-flowered cincinni whereas in Rhapis branching is racemose
throughout the inflorescence, the flowers being borne singly (Fics. 11 &
12). Otherwise Rhapis retains the paniculate construction of Nannorrhops
with the consistent one-to-one relation between bract and branch which is
its essential feature, although adnation of branch to parent axis tends to
obscure this. The distal reduction in bract size is abrupt in Rhapis rather
than gradual as in Nannorrhops.

The morphology of the Rhapis inflorescence is shown diagrammatically
in Fic. 1, with aspect and essential features in Fics. 2-12. Further quanti-
tative information is presented in Taste 1 which also shows differences
between male and female inflorescences. The inflorescence base has a
broad attachment encircling more than one half of the circumference of
the node (Fic. 15). Until final expansion it is wholly enclosed within the
tubular sheath of the subtending leaf and further protected by the three to
five basal bracts (Fics. 4-7). These are papery but fairly rigid, although
they tend to show a fibrous breakdown with age (e.g. Fic. 6). This is
understandable in comparative terms, because bracts are equivalent to the
sheath of a foliage leaf which characteristically shreds into its constituent
fibers with age. Rhapis has the Cocos-type of leaf base, according to their
classification by Tomlinson (1962). This morphological equivalence be-
tween bract and leaf base, so striking in Nannorrhops (Tomlinson & Moore,
1968), is not apparent in Rhapis because it lacks transitional forms.

The first bract (Fics. 4, 5) is a 2-keeled prophyll in the adaxial (ados-
sierte) position normal for the first leaf on a branch in monocotyledons.
However, the branching of the inflorescence axis itself does not continue
this rule because there is no bract in the prophyllar position on subsequent
branch orders. The first bract is situated above a short (sub-prophyllar)
internode. Unlike all subsequent bracts it subtends no branch. It en-
closes the whole of the developing inflorescence until just before final
expansion. The main axis continues with a succession of bracts separated
by long internodes, each bract subtending a branch. Two or three of the
lowest bracts are well-developed, tubular, and approach or even exceed
the prophyll in size (Fics. 6, 7). They are clearly of some initial protec-
tive value but later are torn irregularly at the mouth as the inflorescence
expands. Beyond the last of these conspicuous bracts there is an abrupt
decrease in the size of bracts (cf. Fics. 7 & 8) and length of inter-
nodes (Fic. 3) so that distally bracts are narrow scale-like structures and
branches are more crowded. Although these distal bracts are vestigial and
somewhat dissociated from their branch, careful search will always reveal
the bract corresponding to each branch. Flowers in turn are each directly
subtended by a minute bract, only conspicuous below the lowermost flowers
of the rachilla (Fic. 10). ; :

Branches of up to three orders occur (axi—s (Fic. 1) disregarding the
flower as an ultimate order of branch), although third-order branches are
few and found only on the proximal part of the inflorescence. Third-order
branches are more numerous on male than female inflorescences (TaBLE 1).
The lowest first-order branch is often so well developed as to equal the

296 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Fics. 9-12. Details of inflorescence in Rhapis excelsa. Fic. 9. Main axis, X
2/3, with bracts removed to show extent of adnation between axis and corre-
sponding branch. Fic. 10. Detail of distal part of main axis, X 2, showing scale-
like bracts subtending branches and flowers, adnate branches and basally inserted
second-order branches. Fics. 11 and 12. Rachillae, X 3. Fic. 11, Rachilla with
female flowers. Fic. 12. Rachilla with male flowers.

whole of the remainder of the inflorescence, as in Fic. 2. Adnation between
branch and parent axis in all but the smallest axes obscures the bract-
branch relation. The lowest first-order branches, for example, are adnate
to the main axis throughout the whole of the internode above, diverging

1968 | TOMLINSON & ZIMMERMANN, RHAPIS EXCELSA 297

shortly below the insertion of the next bract (Fic. 9). This may introduce
seeming irregularities where a morphologically lower branch diverges above
the next (morphologically higher) branch inserted on the opposite side.
In this way a branch may diverge beyond the first flower on a rachilla.
In addition a branch may be inserted so close to the base of its parent
axis that two unequal axes seem to diverge at one level (Fic. 10). Bracts
in a prophyllar (adaxial) position do not occur at the base of branches.

Bracts and corresponding branches are arranged in no obvious, regular
phyllotactic order. Probably any primordial order is early lost because
of the requirements of close packing within the protective envelope of
leaf base and lower bracts. The lowest bracts are apparently inserted
distichously at right angles to the plane of insertion of the prophyll, and
subsequent branches seem to prefer this plane, possibly for the simple
reason that it is the one in which there is most room for expansion. The
irregularity is further emphasized by the variable spacing of branches
along a parent axis and in spacing and arrangement of flowers along and
around the rachillae (Fics. 11, 12). “Rachilla” is a convenient term for
the flower-bearing ends of axes of all branch orders (cf. Tomlinson &
Moore, 1968).

INFLORESCENCE ATTACHMENT

In the first article of this series, in which we described the vascular
system of the mature aérial axis in its vegetative phase, it was shown how
each of the larger bundles of the trace system to every leaf was associated
with a cluster of derivative bundles appearing like a “halo” around the
leaf trace in transverse section. Although all of these derivative bundles
diverged successively from the leaf trace in the same way, and all were
anatomically identical, three kinds could be distinguished in each leaf
trace complex, according to their ultimate behavior. The first, and usually
the largest, was the continuing vertical bundle; a number of subsequent
ones functioned as short bridges linking with nearby peripheral vertical
bundles; others (termed by us “satellites” because of their longer associa-
tion with the leaf trace) were traces to the vestigial inflorescence commonly
found in the lower parts of the stem before it has fully entered the re-
productive phase of growth. Despite these vestigial Inflorescences these
nodes may be described as “sterile.” Examination of higher sean
nodes (with fully developed inflorescence branches) shows that the — i
lites are more pronounced and can be referred to more accurately as “In-
florescence traces.”” However, a second kind of vascular connection between
inflorescence and main axis, not found in association with aborted inflores-
cences, now becomes evident. We can distinguish the first an tele
florescence trace by following the vascular system at a node from : ow
upwards, the second by following it from above downwards. H is is
essentially the way in which the two kinds of traces were recognized 1n we
analytical films since there is otherwise no obvious anatomical way 0

298 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

FIG. 13

BRACT INSERTED LOWER DOWN

INF LORESCENCE

VERTICAL BUNDLE

LEAF SSS :

TRACE BRIDGE
-~

INDIRECTLY ATTACHED \ SA

INFLORESCENCE TRACE i

(SATELLITE) ‘

DIRECTLY ATTACHED \
INFLORESCENCE TRACE :

ki

. 13. Diagram illustrating vascular connection between inflorescence and
main axis (below); method of attachment of inflorescence branch to inflores-
cence axis (above). Bundles in longitudinal view are represented by solid lines
for clarity except for directly attached inflorescence traces (double lines). In
the upper figure the bract which subtends the branch is inserted at a much lower
level (cf. Fic. 9).

differentiating them in single sections. The difference between these two
bundles is shown diagrammatically in the lower diagram of Fic. 13.
Following a major leaf trace up, derivative bundles first begin to split
off at about three internodes below the node at which the trace makes its
exit from the stem. The first is almost invariably the large vertical bundle
which remains associated with the leaf trace complex until it turns upwards

1968 | TOMLINSON & ZIMMERMANN, RHAPIS EXCELSA 299

near the periphery of the central cylinder. The production of derivative
bundles then continues from each side of the leaf trace but on a much
larger scale than is characteristic of traces to a sterile node (Fic. 14).
Several of these make bridging union with nearby vertical bundles, but
most (up to 15) remain to form a semicircle on the inner face of the leaf
trace (Fics. 14, 19, 20). The xylem of all these derivative bundles is con-
tinuous with the metaxylem of the leaf trace in such a way that only pro-
toxylem remains in the leaf trace beyond the level of departure of the
last derivative bundle. At the node (Fics. 15, 21) the leaf trace passes
fairly abruptly into the leaf base whereas the inflorescence traces, often
anastomosing amongst themselves in an irregular manner, pass into the
base of the inflorescence axis. The majority of inflorescence traces are
derived directly from leaf traces in this way.

If the vascular bundles in the irregularly anastomosing system in the
base of the inflorescence axis are followed downwards, it is evident that
many attach directly to vertical bundles at the periphery of the central
cylinder (Fic. 13, directly attached inflorescence traces). These directly
attached bundles cannot be distinguished from satellites in a single trans-
verse section. They form an indistinct series on the adaxial (inner) side
of the inflorescence base although many penetrate much deeper. Both
types of bundle are alike in possessing a few narrow metaxylem elements
and a single phloem strand, the whole bundle surrounded by a narrow
Sheath of thin-walled fibers (Fics. 19-21).

The physiological significance of this method of attachment of inflores-
cence to main axis is obvious. The inflorescence not only taps the main
vascular system via the directly attached bundles but is also in direct
vascular contact with the subtending leaf via the satellite bundles and
dorsal leaf traces.

ANATOMY OF INFLORESCENCE AXIS

Shortly above its level of insertion (i.e. within the sub-prophyllar inter-
node) the arrangement of vascular bundles typical of the inflorescence
axis is established. Irregular anastomoses between inflorescence traces are
thus restricted to the very base of the axis. Morphologically the inflores-
cence axis is equivalent to the main stem but there are many pane
differences. The following description of general anatomy refers to a leve
about halfway up the second internode of the main axis (Fic. 16).

section as a result of flattening against main

Outline elliptical in transverse
i i on adaxial surface; each scale more or less

including numerous strands of thin-walled fibers, the larger — iain
narrow phloem strand and usually one (rarely more) narrow and inconsp

300 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Fics. 14-18. Inflorescence anatomy in Rhapis excelsa. Fics. 14 and 15.
Transverse section of aérial stem. Fic. 14. Shortly below node, leaf traces ae

associated satellites conspicuous (arrows). Fic. 15,
florescence base almost free of stem (cf. GS
. Fic. 16. Halfway between base and 2 on of sec

e. Fic. 17. At level of departure of first branch. Fic. 18. Distally,
showing trace systems to two branches. B— branch

Shortly above node
F 6-18

1968] TOMLINSON & ZIMMERMANN, RHAPIS EXCELSA 301

2i,

Fics. 19-21. Details - inflorescence Nevigioo on main stem in Rhapis spe
1

e (5.7 mm. above Fic. 19).

1 inflorescence traces free of leaf traces se
in base of inflorescence. LT — leaf trace; S —satellite; VB — vertical bundle

tracheal element. Central cylinder not separated from cortex by any limiting
layer but by crowded peripheral bundles; central vascular
uniformly scattered. Centra

302 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

central bundles, without protoxylem, the phloem undivided; fibrous sheath
usually continuous but not well developed. Ground parenchyma uniform, compact
in cortex but with well-developed intercellular spaces in central cylinder. Steg-
mata abundant next to all bundles. Vessel elements of metaxylem usually with
more numerous thickening bars than corresponding elements in aérial stem.

In general, the inflorescence axis differs from the aérial axis in the
smaller vascular bundles which do not show a marked peripheral crowd-
ing or pronounced mechanical development of sheathing fibers.

COURSE OF BUNDLES IN INFLORESCENCE

Bract attachment. Bracts are supplied in essentially the same way
as foliage leaves with which they are homologous, differences being largely
quantitative and related to the reduced size of the bract. Structural (and
very probably developmental) principles which govern the distribution of
traces to leaves also govern the distribution of traces to bracts. Each
bract is supplied by relatively few traces which diverge from the central
cylinder, but many more bract traces are continuous with the cortical
system. The lowest 3-5 well-developed bracts are most like foliage leaves.
The few traces originating from the very center of the central cylinder
correspond to major leaf traces of foliage leaves. The somewhat more
numerous ones diverging from intermediate and peripheral levels of the
central cylinder correspond to intermediate and minor bundles of foliage
leaves. Traces derived from the cortex, on the other hand, are very numer-
ous and correspond to cortical bundles of foliage leaves. Bract traces orig-
inating in the central cylinder do not have the “double curve” of von Mohl
which is such a pronounced feature of the trace to foliage leaves. Followed
downwards from the bract node, they show no tendency to approach the
periphery nor is there any helical curve in the center of the axis. In these
respects bract traces are similar to scale traces in the rhizome (cf. Tomlin-
son & Zimmermann, 1966a). In the nodal region the outgoing bract trace
also behaves like a scale-leaf trace in that it gives off a continuing vertical
bundle near the center, followed by one or more short bridge bundles which
link with nearby vertical bundles. As in the rhizome bract traces have no
associated satellites; this is related to the very different method of branch
supply described below.

Corresponding to the decline in length of bracts distally along the in-
florescence axis there is a distal decline in number of bract traces (both
those derived from the central cylinder, as well as cortical traces). The
number of centrally derived traces is reduced much more drastically than
the number of cortical traces. Quite clearly distal, scale-like bracts are
connected to the axial system via cortical bundles alone or at the most
with very few central traces.

Branch attachment. Each branch is morphologically fused with the
main axis but there is no anatomical indication of this in the adnate part
of the internode. At the level of divergence of the branch a portion of the
axis is then simply “pinched off,” the vascular system of the main axis
being correspondingly diminished (Fic. 17). This is repeated at each level

1968 ] TOMLINSON & ZIMMERMANN, RHAPIS EXCELSA 303

(Fic. 18), and in this way the total number of axial bundles diminishes
distally in proportion to the number of branches. There is some slight
tendency to restore the number by occasional splitting of vertical bundles
but this process is too infrequent to effect the distal diminution. It should
be noted that elsewhere in the axis of Rhapis splitting of vertical bundles
is equally rare and insignificant except in the seedling (Tomlinson &
Zimmermann, 1966b).

DISCUSSION

The vascular contact between an inflorescence and its subtending leaf
has not been described previously in palms. This is the shortest possible
pathway for assimilates to move from leaf to inflorescence. Since it is
such a remarkably direct physiological adaptation it may be of value to
know if the arrangement described for Riapis is typical of palms generally,
Our study of vascular continuity has not been restricted to Rhapis. Two
approaches to a comparative survey have been made. The simplest is to
examine single transverse sections of a wide variety of palms and look
for the characteristic well-developed “halo” of satellite bundles aggregated
around the larger leaf traces. This arrangement is common and evidently
a uniform property of many palms. For example, inflorescence traces are
recognizable in some of von Mohl’s illustrations such is their accuracy
and clarity. These observations suggest a vascular supply to the inflores-
cence derived, at least in part, from the leaf trace system as in Rhapis.
It must be remembered, however, that many palms undergo a long period
of vegetative growth without producing any kind of lateral branch. Sec-
tions taken from the lower part of a palm stem commonly show no satel-
lites, although they may be evident in the upper part of the same stem.
Thus the change from vegetative to reproductive phase of growth in the
axis (Tomlinson, 1964) may be reflected anatomically, This in turn means
that satellite bundles in a palm stem may not be revealed by examining
one level.

The only certain method of demonstrating that a palm has the Rhapis-

The practical value of an accurate knowledge of — conan
inflorescence and foliage leaf can be readily grasped. Our observa

TABLE I. Dimensions of ee: male and female inflorescences
|

apis exceisa
MALE FEMALE
ength ength of No. ax,f Total no. Length Lengt No. Total no,
of bract internode corresponding axs on all of bract oepeloat concede wline axs on
Bract (mm. ) below (mm.) tant (ax1) ax2 (mm.) below (mm.) branch (axi) all axe
Prophyll (1) 120 16 2 7 108 20 Ay i
Z 115 61 19 6 118 55 10 2
95 48 16 5 80 80 10 1
4 78 48 14 1 75 68 6 ine
5 50* 47 6 simple 40* 62 2 (rachillae)
6 <7 32 (rachillae) 5 33 f L
; <2 (0) 3 4 simple
& <2 9 4 13 (rachillae)
9 <2 7 simple <2 7
10 <2 9 (rachillae) <2 7
11 <2 :
12 <2 4 rachilla
1 <2 5
14 mod 6
ds in
rachilla
jax: = axis of first-order branch; axe = axis of second-order branch; axs = axis of third-order branch.

Beets marked with an asterisk are lowest which do not encircle the axis completely.

toe

WOLAYOdYY GIONUV AHL AO TVNUNOf

6 “T0A]

1968 | TOMLINSON & ZIMMERMANN, RHAPIS EXCELSA 305

suggest that an inflorescence may be approached directly through its sub-
tending leaf. It might under these circumstances be possible to influence
the development of an inflorescence at an early stage. Leaf and associated
inflorescence do not mature at the same rate. The apex of the subtending
leaf is exposed many plastochrones before the inflorescence itself. In the
Date palm, for example, the youngest leaf which is exposed may subtend
an inflorescence bud only 5 to 7 millimeters in length, but which will not
expand for a further 22 plastochrones (Hilgeman, 1954).

Of particular interest to the morphologist is the contrast between the
method of vascular attachment of inflorescence to main axis on the one
hand, compared with attachment of lateral to main inflorescence axis, on
the other (cf. upper and lower diagrams of Fic. 13). Morphologically the
two are equivalent but developmentally they are dissimilar. We cannot
comment on this difference until the development of root and vegetative
branch insertion has been analyzed. For the moment, it is interesting to
speculate that the reapportionment of vascular bundles between parent
and branch axis in the inflorescence may well account for the apparent
dichotomy of the vegetative axis in a number of other palms, long familiar
in Hyphaene (Schoute, 1909) and more recently demonstrated in other
palms (e.g. Tomlinson & Moore, 1966). Satisfactory answers to such
speculative questions depend on a thorough understanding of develop-
mental processes in palms.

SUMMARY

In Rhapis there is a direct vascular link between the inflorescence and
the leaf which subtends it. Inflorescence traces are mostly attached to
dorsal traces which supply the subtending leaf. Some inflorescence traces
are attached to peripheral vertical bundles of the main axis. Branching
within the inflorescence is racemose throughout but with marked adnation
between branch and parent axis. Distal bracts are reduced to minute
scales. Inflorescence anatomy is essentially like that of the main vegeta-
tive axis. The cortical system is quite well developed, especially in distal
regions where bracts largely make vascular contact with the cortex only.
Bract traces are otherwise derived from the axial system in the same
manner as traces to rhizome scales. At each branch of the inflorescence
part of the vascular system of the parent axis passes directly into the
branch without reference to the corresponding bract. Different kinds of
vascular supply to morphologically equivalent organs indicates Sia
methods of development. Distal reduction in the size of bract and branc
is paralleled by a corresponding reduction in vascular supply.

ACKNOWLEDGMENTS

We are indebted to Miss Priscilla Fawcett, Botanical pecan .
Fairchild Garden for Figures 1-12, and to Miss Lesley Jackson for the

306 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

preparation of the serial sections on which this study is based. Mrs. B.
M. McCurda’s clerical assistance is much appreciated.

LITERATURE CITED

HitceMan, R. H. 1954. The vain pret nine eee and anatomy of the
axillary bud, inflorescence and offs in the date palm. Rep. Date
Growers’ Inst. (California) 31: 4 pp.

MicuHeets, H. 1892. Recherches d’anatomie ame sur les axes fructiferes
des Palmiers. Bull. Acad. Roy. Belg. 53: 1-4

Mout, H. von. 1824. De palmarum structura. a K. F. P. von Martius, His-
toria naturalis palmarum 1: I-LII. 16 pls.

ScnoutTe, J. C. 1909. Uber die Verastelung ie monokotylen Baumen. II. Die
Veriastelung von Hyphaene. Rec. Trav. Bot. Néerland. 6: 211-232.

Tomutnson, P. B. 1962. The leaf eh in Palms — its morphology and mechanical
biology. Jour. Arnold Arb. 43: 2

——. 1964. Stem structure in Ae monocotyledons. Jn: M. H.
ZIMMERMANN, ed. ee formation of wood in forest trees. pp. 65-86. Aca-
demic Press, New

—— &H.E.

Moore, i 1966. Dichotomous branching in palms? Principes
10: Bs 29.

. 1968. Inflorescence in Nannorrhops ritchiana (Palmae). Jour.
pS Arb. 49: 16-34.
& M. H. ZIMMERMANN. 1966a. sai ad of the palm Rhapis excelsa,
Il. Rhizome. Jour. Arnold Arb. 47: 248-26
1966b. Anatomy of the ake Rhapis excelsa, 111. Juvenile
Pe Ibid. ee
ZIMMERMANN, M. H., & P. B. Tomitnson. 1965. Anatomy of the palm Rhapis
Saree, i Mature vegetative axis. Jour. Arnold Arb. 46: 160-180.
1966. Analysis of one vascular systems in ode Optical
Sutte aeohed Science 152: 72-7
1967a. Anatomy ar ‘the palm Rhapis excelsa, IV. Vascular

development in apex of vegetative aérial axis and rhizome. Jour. Arnold
es” 48: oe

A method for the analysis of “9 course of vessels in
Ret Bull. a Asien Wood Anatomists. 1967(1): 2
FAIRCHILD TROPICAL GARDEN
MIAMI, FLoripa 33156
a

n
HARVARD UNIVERSITY
CasotT FOUNDATION, PETERSHAM
MASSACHUSETTS 01366

1968] TOMLINSON & ZIMMERMANN, RHAPIS EXCELSA 307

ANATOMY OF THE PALM RHAPIS EXCELSA, VI.
ROOT AND BRANCH INSERTION

P. B. ToMLINson ! AND M. H, ZIMMERMANN

IN PALMS, as in all woody monocotyledons without secondary vascular
tissues, there are limitations to the method by which a lateral organ can
be attached to the parent axis so as to make an effective vascular union.
Schoute has discussed this problem at a morphological level in a series
of papers on branching in Pandanaceae, Palmae, and woody Liliiflorae
(e.g. Schoute, 1903; 1918; see also Tomlinson, 1964). Anatomical details,
however, still remain undescribed. In the preceding paper in this series
(Tomlinson & Zimmermann, 1968) two distinctive methods of branch-
axis union were described in relation to inflorescence. On the one hand,
the inflorescence axis is basally supplied from the main axis by numerous
bundles which diverge from dorsal leaf traces, together with bundles
which are attached directly to peripheral vertical bundles. On the other
hand, branching within the inflorescence itself involves a simple redistribu-
tion of existing vascular bundles between parent axis and branch. It is
obvious that although these two types of branch are morphologically
equivalent, they must develop differently. However, we have not yet
investigated inflorescence development. To complete the study of vascular
continuity the present article describes the anatomy of root and branch
insertion.

Although these constructional aspects of palms are of fundamental
physiological significance, previous investigation has been limited. Von
Mohl (1824) and Karsten (1847) made some general observations without
describing anatomical details. Mangin (1882) in a notable paper, de-
scribed various methods of attachment of roots to parent axis in monocoty-
ledons, including a number of palms. Drabble (1904) dealt specifically
with the same problem in palms in his lengthy survey of the general
anatomy of palm roots. Our own observations on Rhapis have confirmed
the findings of these earlier botanists.

METHODS

Attachment of root and vegetative branch has been revealed to a large
extent incidentally during analyses of rhizome and seedling axis. Methods
of serial analysis using microcinematography have been described any
in several previous articles (Zimmermann & Tomlinson, 1965; 1966;
1967b). In addition, continuous series of sections through root insertions

‘Research by P. B. Tomlinson on the anatomy of palms is supported by National
Science Foundation Grant GB-5762-X.

308 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

(in both longitudinal and transverse planes) and the insertion of a branch
at the base of an erect shoot were specially cut and analyzed by the same
methods. Photographs which form Fics. 1-9 were made from these addi-
tional series of sections.

MORPHOLOGY

The distribution of adventitious roots and lateral vegetative branches
on seedlings, rhizomes, and at the base of the aérial stem has been de-
scribed and illustrated in the second and third articles of this series (Tom-
linson & Zimmermann, 1966a, b). Briefly, roots are abundant and closely
crowded on the seedling axis, widely and irregularly spaced on the rhizome
and again numerous at the base of erect shoots; vegetative branches de-
velop either from buds in the axils of transitional leaves on the juvenile
axis or at the base of erect shoots. Buds may grow out (usually in order
of their age) into rhizomes but many are strongly inhibited, persisting as
reduced structures within the protective, somewhat woody prophyll. The
subsequent account refers largely to the attachment of roots on rhizomes
and of branches to the base of erect shoots. Their attachment to seedlings
is briefly commented upon (cf. also Tomlinson & Zimmermann, 1966b).

ROOT ATTACHMENT

Roots originate endogenously, rupturing the surface of the stem and
modifying its peripheral tissues (Fic. 2). There is a discontinuity between
surface tissues of root and stem. The epidermis and wide sclerotic layers
of the outer cortex of the root penetrate deeply into (more strictly, dif-
ferentiate first within) the stem cortex. Cortex of root and stem, however,
are continuous. The root endodermis ends blindly in the outer cortex of
the stem. Root development also locally stimulates expansion and some
division of ground parenchyma cells in the outer part of the central
cylinder in such a way that peripheral vascular bundles bulge outwards
(Fic. 2):

In a transverse section of the root, close to its insertion (Fic. 1), a
peripheral system made up of the normal polyarch xylem and phloem
strands can be distinguished from an irregular series of medullary strands
which include metaxylem only. These two systems are attached somewhat
differently and may be described separately. It must be emphasized in-
itially that no root traces end blindly.

Peripheral system. Peripheral xylem and phloem strands of the root
may become attached independently of each other. More commonly the
two tissues form discrete vascular strands (root traces) uniformly sheathed
by short, thin-walled fibers with a common attachment to one stem bundle
(Fics. 3 & 4). However, root traces commonly divide into two strands,
each with its own xylem and phloem. The vascular strands of this periph-
eral root trace complex fan out tangentially and make contact with
peripheral bundles of the stem. For a vigorous root this tangential spread

1968 | TOMLIN

and 2. Root ena in aes excelsa. Fic. 1. Transverse section
F

Transverse section of arate SS

Fr :
of oct abe to its insertio IG. RT t
race;

2a
root insertion, in approsimate median longitudinal section.
VB — vertical bundle of rhizome.

310 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

may be up to one-half the total circumference of the central cylinder
(Fic. 2). Close to the root insertion some of the root traces (or branches
of root traces) penetrate shortly into the central cylinder to make their
attachment. Otherwise traces run in the boundary between cortex and
central cylinder. They have been recognized as frequent “girdling traces”
in the rhizome (Tomlinson & Zimmermann, 1966a, p. 255), remote from
the root attachment.

Medullary system. Each strand consists of a single wide metaxylem
vessel surrounded by a fibrous sheath. The strands penetrate directly into
the central cylinder of the rhizome and are attached to peripheral vertical
bundles (Fics. 2; 5 & 6).

Vascular connection. It is not usually possible to see direct con-
tinuity between xylem of root trace and stem bundle in a single section,
but only in a continuous series of thin sections. The xylem vessel of the
root trace breaks up into an aggregate of narrower elements which run
parallel to the stem bundle (Fics. 3 & 5). The distal elements of this
cluster are applied to the metaxylem vessel of the stem bundle. This
terminal cluster of root trace elements frequently spreads, often nearly
180 degrees, around the circumference of the stem vessel, so making a very
wide contact. The metaxylem of the stem bundle is not usually itself
modified. This contrasts with vessel-to-vessel contact in the leaf-trace
system of the stem, e.g. between bridges and vertical bundles, where bridge
attachment induces a characteristic break-up of the metaxylem into
several narrow elements. This is the probable result of a difference in the
relative time of differentiation of vessels in the two contrasted types of
contact.

Phloem to phloem contact in the peripheral system is usually more direct
and can often be seen in a single section (e.g. Fic. 3).

In the attachment of a mature root, traces are uniformly and fully dif-
ferentiated throughout their length. Many roots, however, abort or are
suppressed. Traces from these root buds connect with the vascular system
of the stem. These traces are fully differentiated proximally (at their
attachment to the stem) but remain in a procambial state distally (at
their attachment to the undeveloped root).

The deep penetration of traces was described by Mangin (1882) as a
characteristic feature of root attachment in Palmae and Pandanaceae.
Drabble (1904) confirmed these observations on the palms and noted that
in many species with larger roots separate strands or aggregates of separate
strands may persist into the root well beyond its insertion (see also Tom-
linson, 1961, pp. 49-50). In Rhapis the deep penetration of central root
traces is particularly noticeable in the seedling. Here they contribute ex-
tensively to the anastomosing bridge system which characterizes this part
of the stem (Tomlinson & Zimmermann, 1966b). Undoubtedly, this deep
penetration is the result of early root development.

We have not traced the course of the medullary xylem bundles into the
root in Rhapis but the observations of Drabble (1904) suggest that these
may unite with the peripheral metaxylem strands of the polyarch stele.

1968 | TOMLINSON & ZIMMERMANN, RHAPIS EXCELSA 311

Fics. 3-6. Details of root insertion in rhizome of Rhapis excelsa; attachment
of root traces to rhizome bundles. Fic. 3. Attachment of aoe
e bun

n
rhizome bundle. «Oy —
elements here free of vessel in rhizome bundle but continuous into root trace.
The root trace in the lower half of both figures shows part of a vessel. PC —
phloem continuity; XC — xylem continuity; X — xylem.

312 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

However, medullary vessels remain as a feature of the distal parts of
Rhapis roots (Tomlinson, 1961).

BRANCH ATTACHMENT

Vegetative axillary buds which produce branch rhizomes differ from
roots in having an exogenous origin. The rhizome itself has a vascular
system of discrete strands which are inserted into the parent axis very
much like root traces (Fic. 7). Individual branch traces are all alike, each
with both xylem and phloem; there are no traces with xylem only, as in
the root. The strands run from the branch and anastomose irregularly
and at random with the bundles of the parent stem (Fic. 8), often forking
distally (Fic. 9). None of them end blindly. Central branch traces pene-
trate deeply into the stem to make their attachment; peripheral bundles
spread markedly around the circumference of the stem forming a distinct
layer of vascular bundles between the cortex and the central cylinder
(Fic. 7). Vessel to vessel attachment between branch trace and stem
bundle is often much more direct than that between root trace and stem
bundle. This indicates approximately simultaneous development of vessels
in the two strands. However, the contact between branch trace and stem
bundle sometimes involves an intermediate plexus of terminal elements
as in the root, indicating later development of the branch trace (e.g. Fic.

Many vegetative buds are inhibited but the suppressed primordia still
make extensive vascular contacts with the parent stem via branch traces
which remain at a procambial stage of development in the bud itself, al-
though the same traces in the axis are fully differentiated.

DISCUSSION

Physiological significance of vascular continuity. It is obvious
that the discrete vascular bundles represent the pathway for translocation
of water and nutrients throughout the palm. The present article completes
a description of the continuous vascular system in Rhapis. The channels
for long distance conduction throughout the palm are now quite evident.
Water from the roots passes into the stem via the numerous linkages at
the root insertion. A continuous channel along the rhizome and up the
aérial stem is afforded by the axial bundle system which links directly
with each leaf via leaf traces. Cross-linkage is effected by the frequent
bridge bundles associated with outgoing leaf traces. In the aérial stem
the helical path of the central bundles further promotes lateral and more
uniform distribution of water. Vascular continuity into the inflorescence
is provided by inflorescence traces which diverge either directly from the
peripheral vertical bundles or as “satellites” from the major leaf traces.
In this way the inflorescence links with both peripheral and central bundles.
New vegetative branches are also connected directly to the axial system

1968 | TOMLINSON & ZIMMERMANN, RHAPIS EXCELSA 313

of the parent stem through which they must derive their water supply
until such time as they produce an independent root system. ae

The pathway from foliage leaves for products of photosynthesis is
equally continuous. At the node the downward-descending nutrients ene
be redistributed in three possible ways. First, they may pass from the lea

314 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

trace to the vertical bundle, or via bridges to other vertical bundles and
thence upwards to the shoot apex. This pathway is clearly demonstrated
in our illustration of the developing vascular system in the crown (Fig. 2,
Zimmermann & Tomlinson, 1967a). Second, assimilates may move di-
rectly to an axillary inflorescence via satellites. This pathway is ver
significant and is discussed in further detail below. Third, assimilates
may move downwards into the stem. Much of this material is stored as
starch in ground parenchyma, particularly in the rhizome, but some must
be available for developing roots and additional rhizomes.

Vascular continuity in developmental terms. An important aspect
of the analysis of the insertion of lateral organs carried out in this article
and the previous one in the Rhapis series (Tomlinson & Zimmermann,
1968) is the demonstration of dissimilar kinds of vascular union in the
attachment of morphologically equivalent kinds of branch attachment and
similar kinds of union in the attachment of dissimilar organs. Any idea
that morphologically equivalent parts must necessarily have a similar
vascular supply must be abandoned. Roots and vegetative branches are
attached in essentially the same way. The attachment of vegetative
branch, inflorescence axis, and inflorescence branch are all dissimilar
although in morphological terms these organs are all equivalent. An ex-
planation for these apparent discrepancies must be sought in develop-
mental terms.

We have suggested in a paper in this series in which development of the
axial system was described (Zimmermann & Tomlinson, 1967a), that the
distribution of procambial strands in the developing crown can best be
understood in terms of competing centers of growth, in which the shoot
apex proper, leaf primordia, and inflorescence primordia each represent
a growth center. The time of appearance of one growth center in relation
to another is of major significance. Centers all receive a vascular supply
by which they are fed. Those that appear early receive a procambial
supply which is continuous with centers of supply of assimilates. These
may be termed “supply” bundles. Centers which appear relatively late
may initially lack continuity with existing bundles. This is established
by bundles originating in the belated growth center and making a late
contact with relatively mature bundles. These may be termed ‘‘demand”’
bundles. It is probably more useful in a physiological sense to speak of
developing vascular traces in terms of “supply” and “demand” rather
than of traces which grow “towards” or “away from” a growth center (or
upwards, or downwards) since this implies a direction of differentiation
which may not exist.

Differing types of organ are supplied by one or other of these types
of bundle, sometimes by both in combination. Growth centers may,
therefore, be categorized developmentally according to their type of vascu-
lar attachment to the parent axis. Root primordia and branch primordia
arise late in relation to the maturation of the axis. They are clearly con-
nected to the main axis by “demand” bundles which seemingly grow
towards existing vascular strands. The vascular tissue of these existing

1968 | TOMLINSON & ZIMMERMANN, RHAPIS EXCELSA 315

strands is at a relatively late stage of maturation and is little influenced
by the late attachment of a trace. We have shown how vessel-to-vessel
contact from root trace to stem bundle involves a plexus of narrow ele-
ments around the vessel of the bundle which otherwise remains little
altered. This reflects a late attachment of the trace.

Root and branch primordia on seedlings develop earlier than their
counterparts on mature axes. This is indicated by the deeper penetration
of root and branch traces into the seedling axis.

Branch and root primordia whose development is inhibited complete a
vascular union with stem bundles in such a way that the fully differentiated
attached end of each trace is immature distally. It is important to realize
that this sequence of maturation is by no means an indication of a “direc-
tion” of differentiation.

The shoot apex proper and primordia of branches on the inflorescence
axis May serve as examples of growth centers wholly vasculated by
“supply” bundles. The latter example is particularly instructive (see Fig.
13, Tomlinson & Zimmermann, 1968), part of the axial system of “supply”
bundles of the inflorescence is simply channeled towards the new growth
center represented by the branch primordium.

The insertion of inflorescence on the main axis serves as an example of
a union which involves both kinds of vascular attachment. The inflores-
cence primordium is initiated early. Contact with the developing stem
system is established via “supply” bundles in the form of “satellites
which diverge from major leaf traces towards the growth center repre-
sented by the inflorescence primordium. Commonly at this stage the in-
florescence aborts and remains as a vestigial axillary structure a few
millimeters long. The satellite bundles, however, persist as a vascular
system out of all proportion to the organ they supply. However, should
normal development of the inflorescence continue, the vascular ee sad
necessary and moreover is supplemented by an additional series of de-
mand” bundles visible as the inflorescence traces directly attached to
peripheral stem bundles. These must appear late in inflorescence develop-
ment because no evidence of them is found in aborted inflorescences.

SUMMARY

Vascular connection between either a root or a rhizomatous branch, on
the one hand, and the main axis, on the other, is effected in essentially
same way. Root or branch traces are attached directly to ax _ hive al
bundles, often penetrating deeply into the stem. An extensive ogee rn
attachment via girdling traces is also present. No bundles en atin ‘de
The relative times of maturation of main and lateral organ determine A
extent and type of connection. This is in turn reflected in the way in
which vessel-to-vessel continuity is established, either directly or via =
intermediate plexus of tracheal elements. Comparison with ; Len
branching in the inflorescence suggests that the vascular siebee e€ sp
main axis and lateral organ must all be understood in developme

316 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

terms. For this reason morphologically equivalent organs may have dif-
ferent attachments if they develop differently and dissimilar organs may
have similar attachments if their development is the same.

ACKNOWLEDGMENTS

The assistance of Miss Lesley Jackson who prepared all the series of
sections on which this study is based is gratefully acknowledged.

LITERATURE CITED

DraBBLe, E. 1904. On ne anatomy of the roots of palms. Trans. Linn. Soc.
Lond. ser. 2. 6: 427-4

Karsten, H. 1847. Die ae ieee der Palmen. Abh. Akad. Berlin
Physik. 1847: 73-236

Manarn, L, 1882. Origine et insertion des racines adventives et modifications
corrélatives de la tige chez les Monocotylédones. Ann. Sci. Nat. Bot. ser.
6. 14: 216-363

Mont, H. von. 1824. De palmarum structura. Jn: K. F. P. von Martius, His-
toria naturalis palmarum 1: I-LII. 16 pls.

ieee an C. 1903. Die Stammesbildung der Monokotylen. Flora 92: 32-48.

See . Uber die Verastelung bei monokotylen Baumen. III. Die Vera-
ae einiger baumartigen Liliaceen. Rec. Trav. Bot. Néerland. 15:
263-335.

Tomutnson, P. B. 1961. Palmae. Jn: C. R. Metcatre, ed. Anatomy of the
Monocotyledons 2. xv + 453 pp. Clarendon Press, Oxford.
Stem structure in arborescent monocotyledons. In: M. H.
ZIMMERMANN, ed. The formation of wood in forest trees. pp. 65-86.
Academic Press, New York.
_ ZIMMERMANN. 1966 a. peaviad of the palm Rhapis excelsa,
II. Rhizome. pha Arnold Arb. 47: 248-2
966b. Anatomy of the rat “Rhapis excelsa, III. Juvenile
phase. Ibid. S012,
1968. Anatomy of the palm Rhapis excelsa, V. Inflorescence.
Soin 49: 291-306.
ZIMMERMANN, M. H., & P. B. Tomuinson. 1965. Anatomy of the palm Rhapis
— sa, 1. Mature vegetative axis. Jour. Arnold Arb. 46: 160-180.
966. Analysis of oem vascular systems in plants: Optical
Shute a Fle Science 152; 72—
67a. Anatomy : ‘the palm Rhapis excelsa, IV. Vascular

development - of vegetative aérial axis and rhizome. Jour. Arnold
Arb. 48: 1

&

A method for the analysis of a course of vessels in
wood. Bull. Int. Anise Wood Anatomists. 1967(1): 2
FAIRCHILD TROPICAL GARDEN
MIaMI, FLorina 33156
AND

HARVARD UNIVERSITY
CaBoT FOUNDATION, PETERSHAM,
MASSACHUSETTS 01366

1968] DICKISON, DILLENIACEAE, III 317

COMPARATIVE MORPHOLOGICAL STUDIES IN
DILLENIACEAE, III. THE CARPELS *

WitiraM C. DickIson

INFORMATION REGARDING floral morphology and anatomy of the Dil-
leniaceae is surprisingly scarce. Floral studies on Dillenia, Hibbertia, and
Doliocarpus by Wilson (1937), Ozenda (1949), and Sastri (1958) provide
only incomplete data. Recently, Professor C. L. Wilson has undertaken an
extensive investigation of the floral anatomy of the entire family. The
first contribution toward this goal (Wilson, 1965) deals with the genus
Hibbertia.

In the present study, no attempt was made to delve into the area of
floral anatomy. Instead, concentration was focused on a comparative
morphological survey of the dilleniaceous carpel.

comprehensive investigations of all orders and families of angiosperms
(Bailey & Swamy, loc. cit.).

Since the combination of characters encountered in the Dilleniaceae
has suggested a position among other presumed primitive taxa (see
Dickison, 1967a), the family is thought to occupy a key position in
angiosperm phylogeny. With this in mind, the present investigation was
carried out with a twofold objective: (1) to attempt to uncover relation-
ships and trends of specialization within the family, and (2) to provide
additional evidence to help clarify the phylogenetic affinities of this
assemblage.

MATERIALS AND METHODS

Carpels of representatives of all ten genera presently considered to
compose the Dilleniaceae were examined. Material was obtained largely
from the author’s personal collection of preserved and dried specimens.
This was augmented only when necessary by flowers derived from herbar-
ium sheets. a

The most convenient method of studying carpel vascularization was
through the use of cleared specimens. Successful clearing was ag
plished with 5 to 8 per cent hot NaOH (Bailey & Nast, 1943) —-
period ranging from twelve hours to five days. Sclerified hairs on the

*I wish to thank Dr. James E. Canright for reading the manuscript.

318 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

highly pubescent carpels of Schumacheria and C uratella had to be meticu-
lously removed prior to clearing, before details of vascularization could
be observed. When phlobaphene pigments obscured the vascular traces,
it was often necessary to use an overnight treatment of Stockwell’s bleach
(Johansen, 1940). After clearing, the carpels were washed and stained
with saturated aqueous basic fuchsin.

Serial sections of floral buds were obtained by the standard paraffin
methods, Dried specimens were first re-expanded in 2.5 per cent hot
NaOH, after which they were washed, fixed in FAA, and treated as pre-
served material. Staining was accomplished with safranin or a safranin-
fast green combination.

OBSERVATIONS

Dillenia. The gynoecium of Dillenia consists of from four to twenty
carpels arranged in a whorl around a conical receptacle. The carpels are
sessile, cylindrically ovate in shape and have elongate, free, characteristi-
cally abaxially recurved styles. The gynoecium is glabrous or occasionally
sparsely covered with unicellular unbranched trichomes. Calcium oxalate
crystals in the form of raphides or crystal sand, and enlarged secretory
cells are common in the wall tissue of some species.

The stigmatic region in those species investigated was indistinct and
restricted to the terminal portion of the style. Hoogland (1952) reports,
however, that two species (Dillenia serrata and D. celebica) have distinct,
knob-like stigmas. The ovular number varies from four to about 80. The
ovules are bitegmic, anatropous or apotropous and generally borne in
single or double submarginal rows (Fic. 12). In D. parviflora it was ob-
served that the micropylar openings in the two integuments do not always
correspond. This interesting feature was first brought to my attention by
Dr. G. L. Stebbins (personal communication), although it has been re-
ported and illustrated in Acrotrema (Swamy & Periasamy, 1955) and
Hibbertia (Sastri, 1958), and has now also been observed in Davilla and
Curatella.

The conduplicate carpels exhibit various syncarpous tendencies. Young
carpels have their ventral margins closely appressed but retain a distinct
ventral suture. At maturity, the ventral suture opens but the carpel re-
mains closed by the adnation of the free ventral margins to the conical
receptacle (Fic. 12). Carpellary fusion between the ventral margin and
the torus in the lower three-fourths to one-half of the gynoecium results
in a solid core of tissue and isolated locules of individual carpels. There
may also occur, simultaneously, varying degrees of lateral concrescence
between carpels. The extent and type of syncarpy changes with carpel
age and level of examination in the gynoecium. A small ventral notch is
all that is evident of conduplicate folding in the stylar region of the mature
carpel.

The carpels of all seven species of Dillenia examined were uniformly
vascularized by three bundles which depart from a eustele in the receptacle

1968 | DICKISON, DILLENIACEAE, III 319

(Fic. 1). The dorsal trace extends through the style and terminates in
the stigmatic region, whereas both ventrals end within the top of the
ovary. Each ovule is supplied by a vein originating from the corresponding
ventral trace. When there are double submarginal rows (as in D. indica),
the original trace bifurcates to supply both ovules. In some species (€.g.,
D. suffruticosa), the ovular trace may become considerably branched
within the ovule. Lateral veins depart at regular intervals from the dorsal
bundle to vascularize the ovary wall.

Acrotrema. From approximately ten known species of this rare, semi-
herbaceous genus, only flowers of a collection by Thwaites (us 1576875)
were available for examination. The gynoecium is composed of three
rather small (ca. 3 mm. in length) glabrous carpels. Basally, the open
conduplicate carpels are laterally concrescent along their inner surfaces,
resulting in a single internal cavity. At the level of ovular insertion, the
Synoecium becomes completely apocarpous with the suture of each in-
dividual carpel closed. A slender, adaxially recurved style is terminated
by an indistinct stigmatic surface.

The number of ovules in the genus ranges from two to six to ten to
twenty (Hoogland, 1951). Swamy and Periasamy (1955) report that the
ovules of Acrotrema arnottianum are typically anatropous, but following
fertilization become completely amphitropous. A comparable situation
occurs in Hibbertia (Sastri, 1958). :

The vascular pattern in the single species investigated (Fics. 2, 13) is
similar to that described for Dillenia, with the exceptions that in Acro-
trema the ventral bundles approximate the dorsal in length, and there is a
noticeable absence of lateral carpellary wall veins.

Hibbertia. The bisexual flowers of the largest dilleniaceous genus have
one, two, three, five or rarely (in H. crenata), ten carpels (Wilson, 1965).
This genus has often attracted attention because it is thought to contain
members with the least specialized floral structure in the family. In some
species the gynoecium is entirely apocarpous but frequently it displays
varying degrees of basal syncarpy. The carpels are differentiated into a
swollen ovary which is projected distally into a narrow, elongated, often
abaxially orientated style; at the tip of the style is an indistinct, or slightly
enlarged, stigmatic surface. The carpels vary from glabrous to densely
pubescent and may contain numerous raphides crystals (eg., H. exutia-
cies). In addition, the presence of a single druse crystal in the ovules of
certain species (e.g., H. tetrandra and H. cuneiformis) is very —
One to 25 bitegmic, anatropous, submarginal ovules are present in eac
ovary. As has been noted previously, the micropylar openings 1n the two
integuments do not always correspond. ae

Wilson (loc. cit.) has described two distinct types of ier wit 2
the bicarpellate species. He arbitrarily designates these as eo =
“oblique” forms. In an “opposite” gynoecium, the carpels normally ct
their dorsal bundles lying directly opposite each other and the ventra

320 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

sutures remain centrally located throughout the length of the carpel. In
“oblique” carpels, however, the dorsal bundles depart at an angle, with
a subsequent displacement of the ventral traces and sutures to one side
of the carpel. The occurrence of these two morphologically dissimilar
gynoecial types is clearly observable. Wilson considers the oblique condi-
tion to be specialized and suggests it may have arisen more than once
within the genus.

All specimens studied were vascularized by three traces: a dorsal and
two ventrals. In material of such species as Hibbertia scandens (Fic. 3),
the dorsal bundle supplies numerous lateral veins to the ovary wall before
ending at the style apex. The vascular supply to the ovules is derived
from the ventral bundles.

A well documented trend of reduction in carpellary vascularization is
present within the genus. For example, in H. tetrandra (Fic. 4) only two
lateral wall veins originate from the dorsal, and the ventral bundles are
much reduced. The culmination of this specialization can be found in the
greatly reduced carpel of the genus Pachynema (see Fic. 5).

Pachynema. Two species of this Australian genus were available for
investigation. The bisexual flowers contain two glabrous carpels with only
slight basal fusion. Each carpel is differentiated into an enlarged ovary
and a narrow, slightly reflexed, vascularized style. A very localized, apical
stigmatic surface is quite indistinct. Secretory cells are very common in
the ovary wall.

Reduction in the sterile ventral borders results in the conduplicate

nature of these carpels becoming obscured. The bicarpellate gynoecium
is best described as opposite, although this appears to be a rather unstable
character with tendencies toward obliqueness being observed. The ventral
sutures are typically open at the level of ovular placentation, but closed
below and above this location. Variations in the degree of closure reflect
differences in carpel maturity.
The reduced nature of the gynoecium in this genus is very pronounced
in the pattern of carpel vascularization (Fic. 5). One dorsal and two
ventral bundles enter the base of the carpel. Four major lateral veins are
derived from the dorsal bundle and supply the ovary wall. The status of
the ventral traces is considerably reduced with each one serving as the
vascular supply to one of the two basal ovules.

Didesmandra. The gynoecium of the single species, Didesmandra
aspera, is bicarpellate with only slight basal connation present. The
carpels are very characteristically differentiated into a large, swollen,
glabrous ovary and an elongate (15-20 mm.), filiform style. A stigmatic
region is restricted to the style apex. Numerous enlarged secretory cells
are scattered throughout the ovary wall.

The two conduplicate carpels are oblique in their orientation (Fic. 14).
Even though the oblique condition is clear, it is not as pronounced as in
some hibbertias. The conduplicate nature of the carpels is somewhat

1968 | DICKISON, DILLENIACEAE, III 321

obscured by the reduction and fusion of the ventral margins rendering the
gynoecium permanently closed along its entire length.

The vascular system (Fic. 7) at the base of each carpel is eustelic. A
dorsal bundle departs initially from the stele, closely followed by a pair
of ventral traces. Following the departure of the dorsal and ventral carpel
bundles, the stele contracts to a pithless condition and continues distally
to vascularize the single, comparatively large, basal ovule. The ovular
trace terminates in a series of short branches. The ovary wall is supplied
by lateral veins from both dorsal and ventral bundles,

Schumacheria. All observations on this endemic Ceylonese genus were
made from S. castaneifolia. Although the flowers are often described as
possessing three carpels, bicarpellate flowers are quite common. In either
case, the gynoecium is densely covered with sclerified hairs. The presence
of calcium oxalate crystals in the form of either raphides or crystal sand,
in addition to large secretory cells, is also characteristic for this species.
The ovary, containing a single basal ovule (Fic. 16), projects distally into
an elongate, abaxially recurved style which narrows at its tip to form a
terminal stigma.

In bicarpellate gynoecia, the carpels are distinctly oblique in orientation.
In such cases, syncarpy occurs between carpels throughout the length of
the ovary by fusion of only one of their ventral borders. Also, one carpel
of the gynoecium may open at maturity while the other remains closed.

In tricarpellate flowers, one carpel was found to develop at a higher
level than the other two which are initially somewhat oblique. As the third
carpel develops, the “obliqueness” evident basally is lost. This observa-
tion suggests that in Schumacheria the third carpel is in the process of
being lost and the oblique condition is a result of this process. A de-
velopmental study of the flower of Schumacheria would perhaps aid in
clarifying the origins of the oblique gynoecium.

The vascularization of Schumacheria carpels is quite complex but re-
sembles Didesmandra in that following the departure of the dorsal and
ventral carpel traces, the stele becomes pithless and forms a thick, stout
ovular trace which “fans out” in the ovule into numerous small veinlets
(Fic. 6). Three traces leave the stele at approximately the same level.
These bundles branch and bifurcate almost immediately, however, so the
carpel, for all practical purposes, is vascularized by more than three major
traces. The two ventral traces do not contribute to the vascularization of
the ovule, The ventrals extend to the summit of the style with the dorsal
trace. Numerous lateral veins anastomose throughout the ovary wall.

The suggested close relationship of Schumacheria and ianar entity
(Hoogland, 1951) is supported by carpel morphology, pollen morphology,
and wood anatomy.

Tetracera. This genus is particularly noteworthy in ae 1s
unisexual and bisexual flowers in an otherwise entirely bisexual family.

It is interesting in this regard that Hoogland (1953) refers to the flowers

322 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

of the Indo-Malayan tetraceras as bisexual, whereas Hunter (1966) calls
the flowers of the Panamanian forms unisexual. The small number o
African specimens I examined possessed bisexual flowers; thus, unisexual-
ity appears to be confined to the New World. However, additional collec-
tions will have to be studied to support such a basic distinction.

The apocarpous gynoecium is composed of one to five pyriform carpels.
The carpels vary from glabrous to densely pubescent, and possess from
one to twenty anatropous, bitegmented ovules arranged in two to six
submarginal rows. An erect, occasionally abaxially directed, style is termi-
nated with an undifferentiated or slightly peltate stigmatic surface. Cal-
cium oxalate crystals in the form of raphides or crystal sand and large
secretory cells are quite common in the wall tissue of some species (€.g.,
Tetracera indica).

The conduplicate nature of the carpels is apparent in both the ovary
and the solid style where a persistent ventral fold remains. Characteristi-
cally, the carpels are completely closed and fused basally. Progressing
distally, the ventral suture becomes merce more pronounced with
the ventral region becoming open in some spec

Carpel vascularization is as follows (Fic. A following the departure

large, isolated bundles corresponding in number to the carpels. Thus,
each carpel is supplied by a single, large toral trace. Upon entering the
carpel base, the dorsal bundle departs first, closely followed just below
the initiation of the locule by the bifurcation of the remaining toral trace
into two ventral bundles. Many carpels observed had accessory carpel
bundles originating from the toral bundle. These accessory veins are
usually quite reduced and either end blindly or connect with other lateral
wall veins, The occurrence of numerous, highly anastomosing, lateral wall
veinlets between the ventral and dorsal bundles is well developed in the
genus. The ovules are supplied by veins branching from the ventral traces.

Curatella. The flowers of the monotypic Curatella americana are
normally bicarpellate, although Hunter (1966) reports that the number
of carpels may rarely be one to five. The gynoecium is very densely pubes-
cent. Syncarpy occurs between carpels by complete fusion of the adaxial
ovary surfaces. The erect styles are typically free and terminated by a
peltate stigmatic surface. Raphides crystals and large secretory cells are
present throughout the gynoecium

In addition to fusion between carpels, the ventral sutures of both carpels
are also closed basally (Fic. 15). The ventral suture becomes more distinct
at the level of placentation and may even become completely open. In the
solid style a notch remains in the ventral region. The orientation of the bi-
carpellate gynoecium is strictly opposite.

As seen in Fic. 10, the carpel of Curatella is vascularized by five major
bundles: a dorsal, two ventrals, and two large accessory traces. Occasion-
ally, additional smaller supernumerary traces are distinguishable so that
seven carpel traces are present. Each carpel trace arises independently

1968 | DICKISON, DILLENIACEAE, III 323

in the toral stele; however, in some gynoecia the ventrals were observed to
diverge from a common sympodium. At the base of the carpel, complex
anastomosing and branching occurs in the stelar system before the major
carpel traces emerge. A similar situation has been reported in the Winter-
aceae (Nast, 1944; Tucker & Gifford, 1964). The dorsal bundle, which
bifurcates at the base of the style, and the two ventrals terminate just
below the stigma. The accessory bundles usually connect with other lateral
wall traces departing from the dorsal. Each ventral trace supplies the
vascularization to one of the two, bitegmented, basal ovules. The micro-
pylar openings in the two integuments do not always correspond.

Davilla. This genus is uniformly characterized by monocarpellate
flowers. The glabrous gynoecium is distinguished by a slightly clavate
style and a flattened, peltate, stigmatic surface. Two basal, bitegmic
ovules are present in each carpel, along with the rather frequent occurrence
of raphides crystals. The micropylar openings in the two integuments do
not always coincide.

In the Dilleniaceae, Davilla appears to be unique in that the ventral
carpel margins are involute and not conduplicately folded. At the carpel
base, the ventral region begins to open along the outer ventral surface in
what would be considered a conduplicate manner. Slightly higher, how-
ever, the ventral margins become distinctly involute (Fic. 11 b), with the
outer carpel surfaces being brought inside. At this level the carpel may
either be closed or open. Above the point of placentation the involuted
margin begins to fuse internally and eventually all evidences of the ventral
suture and folding are lost (Fic. 11 c).

The vascularization of Davilla aspera is rather simple (Fic. 11 a) when
compared to other New World genera. Three carpel traces are present at
the base of the locule. The dorsal bundle extends unbranched along the
length of the style, except for the departure of two lateral wall veins. Both
ventrals are reduced but give rise to the vascular supply to the ovules.

Doliocarpus. With the exception of the bicarpellate D. coriaceus, es
genus is also unicarpellate (Hunter, 1966, reports the Panamanian species
may rarely possess two carpels), All species have carpels with well-dif-
ferentiated ovaries and extended styles terminated by peltate stigmatic
surfaces of varying size. The gynoecium is either glabrous or densely
pubescent (e.g., D. major). Similar to many other Dilleniaceae, the carpel
wall frequently contains large secretory cells. Doliocarpus dentatus ot
9) is readily distinguished from other species by the presence of raphides
crystals, which were often so numerous they obscured details of VRSCA:
ization in cleared carpels.

The carpels of Doliocarpus are conduplicate although the ventral fare

rgins of D. dentatus and D.
1 of locule formation begin to
the level of ovular

degree of carpel closure. The ventral ma
guianensis are closed basally, but at the leve ot I
open internally until they become completely distinct at

324 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

insertion. Above the level of placentation, the carpel begins to close
centripetally. Doliocarpus major, by contrast, remains open along the
entire ovary length.

Doliocarpus coriaceus shows complete adnation between the two op-
posite carpels. At the level of ovular placentation, the ventral suture
of each carpel opens, creating a continuous internal cavity within the
gynoecium, Immediately above this point, the ventral region fuses solidly
leaving no evidence of sutural structure. Separation of the style is initiated
by an internal cleft which proceeds outward.

Each carpel of Doliocarpus coriaceus is supplied by three major bundles,
a dorsal and two ventrals. The dorsal trace is well pronounced and vas-
cularizes the style. The weaker ventrals terminate within the ovary after
initiating traces to the two basal ovules. Small lateral wall veins occasion-
ally originate from and anastomose between the ventral and dorsal bundles,
as well as from the toral region.

All the monocarpellate species examined were vascularized by more
than three major traces departing from the stele. Generally a distinct
dorsal bundle is recognizable, which, upon entering the style, undergoes a
series of bifurcations resulting in a rich vascularization of the style. Al-
though ventral traces can often be discerned, they are not at all pronounced
or distinguished from the other four to six accessory bundles. The pair
of ovules is supplied by veins originating directly from the receptacle.
Numerous lateral wall veins are present throughout the ovary. The com-
plex basal branching and anastomosing observed in Curatella is also
present in Doliocar pus.

DISCUSSION

Among the various species of Dilleniaceae examined, the following
primitive gynoecial features were observed: (1) complete or nearly com-
plete apocarpy; (2) visible conduplicate folding; (3) open ventral su-
tures; and (4) large numbers of ovules. It is clear, nevertheless, that no
extant dilleniaceous carpel is closely comparable to the primitive ranalian
megasporophyll.

On the other hand, the Dilleniaceae are more readily characterized by
a larger number of moderately to highly advanced phylogenetic specializa-
tions which parallel those that occur in angiosperms as a whole: (1) reduc-
tion in number of carpels to one; (2) differentiation and elongation of a
narrow, vascularized style; (3) formation of a restricted, terminal, often
peltate stigmatic surface; (4) reduction in number of ovules to two or
one; (5) varying degrees of basal adnation and/or lateral concrescence;
(6) complete fusion of the ventral suture; (7) reduction or amplification
of the basic three-veined carpel vascularization: and (8) tendency toward
involute closure. The occurrence of the oblique gynoecium in many bi-
carpellate Old World species is an intriguing feature of possible phylo-
genetic significance

The least specialized carpels in the family are found among the apo-

1968 } DICKISON, DILLENIACEAE, III 325

carpous species of Hibbertia. When considered as a whole, the subfamily
Dillenioideae is less specialized in carpel morphology than the Tetraceroi-
deae. This information may be correlated with that obtained from other
lines of evidence, e.g., wood anatomy. The fact that direct correlations
do not always hold true can be demonstrated in Didesmandra and Schu-
macheria, which have very primitive wood but rather specialized gynoecia,
and Tetracera, with more advanced wood but comparatively primitive
carpels. Most frequently, dilleniaceous species have various combinations
of advanced and primitive characters present simultaneously. For this
reason, no one subfamily or genus retains exclusively primitive or advanced
features. The opinion of Hoogland (1951) that the gynoecium of Dillenia
is the most advanced in the family is not substantiated.

According to the classical concept, the primitive carpel is a three-trace
organ (Eames, 1931). The primitive nature of the three-trace condition
was generally accepted in light of studies on the woody Ranales (e.g.,
Bailey & Swamy, 1951). From the three-trace condition, a multitrace
carpel would be derived by amplification, whereas a two-trace organ would
result from loss of one of the ventrals.

With the discovery of multitrace carpels in some otherwise primitive
families — Degeneriaceae, Lauraceae, Winteraceae, Myristicaceae, and
Annonaceae among others, the primitive nature of the three-trace carpel
was questioned. Some authors, e.g., Swamy (1949), suggested that the
presence of additional bundles in the monocarpellate gynoecia of Degen-
eria was indicative of a multicarpellate ancestry with the extra bundles
being considered remnants of missing carpels. In reviewing the vascular-
ization in ranalian carpels, Sastri (1959) has specifically warned against
the assumption that the three-trace carpel is primitive for all angiosperms.
Tucker and Gifford (1966b) are of the opinion that the three-trace
concept is not essential to an understanding of carpel vascularization, and
point to Sterling’s (1953) conclusion that the basic pattern in the Rosa-
ceae is five-trace. :

When the vascular pattern in the Dilleniaceae is correlated with other
indices of advancement such as closure, ovule number, degree of syncarpy,
etc., there is little question that the three-trace condition is primitive.
Specializations in vascularization have subsequently occurred toward re-
duction (particularly in the ventral bundles) and amplification. It appears,
accordingly, that there exists a fundamental distinction in carpel vascular-
ization between the Dilleniaceae and the Rosaceae, a family often derived

326 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

contrast, the unicarpellate hibbertias have three prominent traces (Wilson,
1965), whereas the New World genus Davilla retains the three-trace con-
dition in a considerably reduced form.

My conclusions (Dickison, 1967a & b), based on wood anatomy and
pollen morphology, that previous magnoliaceous ranalian affinities ascribed
to the Dilleniaceae are not well founded also gain support from the vas-
cular anatomy of the carpels. In some Magnoliaceae and Annonaceae, the
ovules are vascularized by traces from the dorsal bundle (Canright, 1960;
Periasamy & Swamy, 1956). Although there is no uniform method of
ovular vascularization in the Dilleniaceae, ovular traces never originate
from the dorsal bundles. The cortical system characteristic of many
Ranales is also lacking in the dillenias. Likewise, the compound gynoecia
and axile placentation of the Theaceae show little similarity to the Dil-
leniaceae.

A more comparable situation is found in Paeonia. The gynoecium of
Paeonia is apocarpous with each carpel vascularized by three principal
bundles, a dorsal and two ventrals, accompanied by a variable number of
smaller secondary veins. Each of the two ventral traces bifurcates at the
base of the style resulting in a richly supplied stigma. It is particularly
noteworthy that Brouland (1935) diagrammed a bicarpellate gynoecium
of P. delavayi as being oblique. Since this species is more commonly tri-
carpellate, it again indicates that the oblique condition probably results
from reduction within the gynoecium. Also, the occurrence of the oblique
gynoecium in Paeonia suggests that this character might have phylo-
genetic significance in the Dilleniaceae and allies.

Unfortunately, the scanty, or nonexistent information on the floral
morphology and anatomy of many families putatively related to the Dil-
leniaceae prevents a critical comparison.

MATERIAL EXAMINED 1

Acrotrema sp. CEYLON: Thwaites CP pes (us 1576875).
Curatella americana L. Braziu: Irwin (NY
Davilla aspera (Aubl.) Benoist. bea NT, Silva 16.
Davilla sp. BRaziL: Jrwin 5570
Didesmandra aspera Stapf. pa Burtt & Woods B.2540 (£); SAR 18297.
Doliocarpus coriaceus (Mart. & Zucc.) Gilg. Cotompra: Haught 1655 (us).
sete dentatus (Aubl.) Standl. Brit. Honpuras: Gentle 1418 (Mo);

vIA: Krukoff 10407(wuc).
bistigcrie guianensis (Aubl.) Gilg. Costa Rica: Skutch 3980 (us).

‘Material studied was obtained from the State Herbarium of South faarye ae
Adelaide (aD); The Forest Herbarium, Bangko
b ;

( uis (Mo); Animal Industry Branch, Northern
Eines Penang Alice Springs (nr) ;
Sar

e
» Kuching (sar); Botanic Gardens, Singapore (sinc); University
rs Caio geen nee and the United States Nadioual Herbarium, Washington

1968] DICKISON, DILLENIACEAE, III 327

Doliocarpus major Gmel. PANAMA: siete et al, 223 (us).

Dillenia aurea Sm. THAILAND: BKF

Dillenia indica L. AusTRALIA: Cult. BRI Sn.

Dillenia ovata Wall. ex Hook. f. & Thoms. Stncapore: Cult. SIN

Dillenia papuana Martelli. New Guinea: Darbyshire & Hooelind 7030 (CANB).
Dillenia parviflora Griff. THAILAND: BKF s.n.

Dillenia philippinensis Rolfe. PuHirppines: J. V. Pancho s.n.

Dillenia suffruticosa (Griff.) Martelli. inet Cult. SING s.n.
Hibbertia cuneiformis (Labill.) Gilg. Cult.

Hibbertia dentata R.Br. Cult. K s.n

Hibbertia exutiacies Wakefield. AUSTRALIA: cgi 17965 (aD).

Hibbertia scandens (Willd.) Dryand. Cult. K

Hibbertia tetrandra (Lindl.) Gilg. Cult. E C3544

Pachynema dilatatum Benth. Austratia. Northern Territory: NT 6129.
Pachynema junceum Benth. AustrRaLiA. Northern Territory: NT 6750.
Schumacheria castaneifolia Vahl. CEvLON: Abeywickrama s.n.

Tetracera akara (Burm. f.) Merr. Borneo: UC 290527

Tetracera indica (Houtt. ex Christm. & Panz.) Merr., SINGAPORE: Keng s.n.
Tetracera podotricha Gilg. ANGOLA: Coops 9135 (US).

Tetracera scandens (L.) Merr. MaLaya:

Tetracera volubilis L. Pancua: Allen re (mo).

LITERATURE CITED

Battey, I. W., & C. G. Nast. 1943. The Sage ones morphology of the Winter-
aceae, II. Carpels. Jour. Arnold Arb. 24: 472-4
& B. G. L. Swamy. 1951. The ace carpe . rete and
its initial trends of specialization. Am. Jour. Bot. 38:
BrouLanp, M. 1935. Recherches sur l’anatomie florale Aes ieee Le
Botaniste 27: 1-280. a
Canricut, J. E. 1960. The comparative morphology ee relationships of the
Magnoliaceae. III. Carpels. Am. Jour. Bot. 47: 145-15 f
Dickison, W. C. 1967a. Comparative rata me aes in Dilleniaceae,
Wood anatomy. Jour. Arnold Arb. 48: 1-29. 1
1967b. iis sepa pea studies in Dilleniaceae, II. e
pollen. Ibid. 231-240.
Eames, A. J. 1931. The vascular anatomy of the ren cs refutation of the
theory of carpel polymorphism. Am. Jour. Bot. 1
Hoocianp, R. D. 1951. Dilleniaceae. Jn: C. G. G. f, ve see Fl. Males.
I. 4: 141-174. on
. 1952. A revision of the genus Dillenia. Blumea 7: 1- ae
The genus Tetracera (Dilleniaceae) in the eastern Ol or
Reinwardtia 2: 185-225.
Hunter, G. E. 1966. oo (Part VI, Flora of Panama). Ann. Missouri
Bot. Gard. 52: 579-
i York.
Microtechnique. the Hill. New
JoHANSEN, D. A. 1940, madi icr Fae Wikicscase VU Vex
: 454-466.
Publ. Lab.

‘Ecole Normal Supérieure, Ser. Biol. Fasc. II. Paris.

328 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

oe K., & B. G. L. Swamy. 1956. The conduplicate carpel of Cananga
dorata. Jour. Arnold Arb. 37: 366-372.
Sie R. L. N. 1958. Floral morphology and embryology of some Dilleniaceae.
Bot. Not. 111: 495-511.
. 1959. The vascularization of the carpel in some Ranales. New Phytol.
58: 306-309.
STERLING, C, 1953. Developmental “ee of the fruit of Prunus domestica
L. Bull. Torrey Bot. Club 80: 457-4
Swamy, B. G. L. 1949. Further contin to the morphology of the Degen-
eriaceae. Jour. Arnold Arb. 30: 1
& PERIASAMY. 1955. eee to the embryology of Acro-
dale arnottianum. Phytomorphology 5: 301 ‘
1964. The concept of the sbidtaplicate carpel. Phytomor-
tte 14: 319-327.
Tucker, S. C., & E. M. Grrrorp, Jr. 1964. Carpel vascularization of Drimys
lanceolata. Phytomorphology 14: 197-203.
= aps in the carpellate flower of Drimys
lanceolata. Am. Jour. Bot. 53: 433-442.

‘a

& . 1966b. Carpel Saas in Drimys lanceolata. Ibid. 671-

678.
WILSON, be es 1937. The phylogeny of the stamen. Am. Jour. Bot. 24: 686-699.
The floral Peril of the Dilleniaceae. I. Hibbertia Andr. Phyto-
evans 15: 248-2

DEPARTMENT OF BIOLOGY
VircIniA PoLyTECHNIC INSTITUTE
BLACKSBURG, VIRGINIA 24061

EXPLANATION OF PLATES

PLATE I
Explanation of symbols: c., druse crystal; d.b., dorsal bundle; e.p., stigmatic
surface; Lb., lateral Suid, ae ovular bundle; vk. ovular trace; t., trichome;

v.b., ventral Bandi.

Vascularization of dilleniaceous carpels; only one of the ventral
bind a half the lateral veins are shown. 1, Dillenia parvifl BKF 5.1.)
from Thailand. 2, Ac ma sp.

rotre A
K “he 4, Hibbe rtia tetrandra (Cult. E C3544). oF SS ma dilatatum So

PLATE II
Explanation of symbols: a.b., accessory bundle; d.b., dorsal bundle; e.p., dagarts
surface; 1.b., lateral roe ov., ovule; ov.b., ovilar bundle; r., raphi ides; t., tri-

ome; v.b., ventral bundle

Vascularization of dilleniaceous carpels, continued: only one each

of the ‘venta bundles and accessory origi and half the lateral ore are

shown Dads sales aspera (SAR 18297). 8, Tetracera akara (uC 527).

9, pokvewpus dentatus (Krukoff 1

wea 10, Curatella americana (Irwin 5470).
il.

r
11 a, b, c, Davilla aspera
N.T. Silva 16), from Belém, Braz:

1968 | DICKISON, DILLENIACEAE, III 329

PLATE III

Explanation of symbols: d.b., dorsal bundle; ov., ovule; ov.t., ovular trace; s.t.,
stamen trace; v.b., ventral bundle; v.s., ventral suture.

Fics. 12-16. Dilleniaceous carpel morphology. 12, Dillenia rg tasleppe
(J. V. Pancho s.n., from Makiling, Philippines), transverse section 0 ae!
a2. -43, Acrotrema sp. (us 1576875), cleared and stained flower pha prare
of vascularization; one carpel has been removed, X 30. 14, Didesmandra aspera
(SAR 18297), transverse section of gynoecium showing large, solitary ovule, en-

larged secretory cells, and oblique orientation, a5 , Curatella americana
UIrwi id nla transverse section of gynoecium in a of rr we show-
ing c tise of the two oppose carpels, umacheria
cstonifolia (Abeywickrama s.n., rr es aga, Denton ioratteiaal section of

carpel showing sola, basal ovule, 2

I
Jour. ARNOLD Ars. VOL. 49 PLATE

Dicktson, DILLENIACEAE, III

Jour. ARNOLD Ars. VoL. 49
Piate II

~+ێ.p.

1
Dick1son, Ditientaceak, III

Jour. ARNOLD ArB. VOL. 49 PLaTE IIT

DICKIsON, DILLENIACEAE. ITI

1968 | GREEN, REVISION OF NOTELAEA 333

REVISION OF THE GENUS NOTELAEA (OLEACEAE)
P. S. GREEN

THE GENUS Notelaea is endemic to eastern Australia and Tasmania,
although it was first described from a plant under cultivation in France,
introduced from what is known today as New South Wales. As treated
here the genus contains nine species only; those described from outside
Australia being excluded and disposed of in other genera.

Seven species are very closely related and much confusion has existed
in their recognition and differentiation. A critical redetermination of many
misnamed specimens has helped to clarify the species’ limits and the ex-
amination of a large range of herbarium material, together with a very
limited, but invaluable, experience of some of the plants in the field, has
led me to believe that the recognition of these seven taxa is justified at
the rank of species. However, experimental and field studies, if and when
they are carried out, may modify the rank at which the taxa are finally
classified. The other two species, although related, are sufficiently distinct
to warrant the establishment of two new sections for their accommodation ;
their differences from the seven species mentioned seem insufficient to
justify generic separation.

Hybridization between closely related species has not been proved
experimentally but the combination of characters exhibited by a very few
specimens can only be explained on the assumption that this has occurred.
Under Notelaea microcarpa mention is made of a putative hybrid with
N. ovata, and N. microcarpa is thought to have introgressed with NV.
longifolia on occasion, as shown, for example, by the variation in a relict
population near Dorrigo, New South Wales (represented by the range of
specimens in Green 1519 a-1) cited below under N. longifolia f. intermedia.

Field experience of two or three species was made possible for me by
funds from the Arnold Arboretum (where I had the honor of being a
Staff member at the time) together with National Science Foundation
Grant No. GB1545, and most grateful acknowledgment is made for this
generous support. I should also like to express my sincere thanks to Mr.
L. A. S. Johnson of the Royal Botanic Gardens, Sydney, for much informa-
tion gleaned from his knowledge of the species in the field and sss
for the many valuable annotations he attached to the herbarium materia
when it was sent on loan from Sydney; to Messrs. A. G. Floyd and H. C.
Hayes of the Forestry Commission of New South Wales, Coff’s Harbour,
for their generous help with time, transport, and local knowledge of a
species in northern New South Wales; and to Miss Mary Grierson an
Miss Judy Appenzeller for the maps and figures. Finally, I coon as :
record my grateful appreciation to the directors and curators of the ci An
herbaria for the loan of material or facilities for study. All the materia

334 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

cited has been examined, and the respective herbaria are indicated by the
abbreviations published in the Jndex Herbariorum, Ed. 5, 1964

Notelaea Ventenat, Choix de Plantes 25. ¢.25 1804; R. Brown, Prodr.
523. 1810; Endlicher, Gen. Pl. 572. 1838; A. DC. in DC. Prodr. 8:
290. 1844; Bentham, FI. Austral. 4: 298. 1868 & in Bentham &
Hooker f. Gen. Pl. 2: 678. 1876; C. Moore, Handb. Fl. New S. Wales
320. 1893; Knoblauch in Engler & Prantl, Nat. Pflanzenfam. 4(2):
10. 1892; F. M. Bailey, Queensl. Fl. 3: 972. 1900; Johnson, Contr.
New S. Wales Natl. Herb. 2: 411. 1957.

Rhysospermum Gaertner f. Fruct. 3: 232. ¢.224. fig. 2. 1807.
Postuera Rafinesque, Sylv. Tellur. 10. 1838.

Evergreen shrubs or small trees. Leaves opposite, simple, glabrous to
velutinous, lamina thickish or coriaceous, entire or crenulate, venation
obscure or clearly reticulate. Jnflorescence axillary, decussate with terminal
flower, from 1—3 superposed buds, bracts broadly ovate to narrowly lance-
olate, early deciduous. Flowers hermaphrodite. Calyx 0.3-1.2 mm. long
with 4 more or less triangular lobes 0.1—-1 mm. long. Corolla of 4 petals
joined in two pairs at the base of the stamens, the pairs free, or only just
joined at the base (in N. ligustrina) , lobes 1.2-2.5 mm. long, broadly ovate,
rounded or more or less acute (in N. ligustrina), induplicate-valvate or
slightly imbricate in the bud (in N. ligustrina), concave and enclosing
the stamens, closely so and raised on a common “stalk” at maturity in
N. johnsonii. Stamens 2, anthers 0.8-1.5 mm. long (0.3 mm. in NV. john-
sonii and closely invested by the corolla), filaments short 0.1-0.5 mm. long.
Ovary triangular flask-shaped, 0.8—2 mm. long without well differentiated
style, terminal stigma more or less bilobed 0.1-0.5 mm. long. Drupe ellip-
soid to more or less globose, 5-20 mm. long by 4-16 mm. broad; mesocarp
soft when ripe and purplish- ee but white, pink or red in some species;
endocarp hard, 0.1-0.6 mm. ick.

Type species: Notelaea soe Vent.

KEY TO SECTIONS

. Corolla lobes induplicate-valvate in the bud (Fic. 1, A & B
2. Paired corolla lobes and stamens sessile at maturity, the bases of ad-
jacent pairs touching or almost touching, and surrounding the ovary;
anther lobes about the same size as the connective (Fi. 2, BS C+
flowers yellow, creat op white: 545 NoTELAEA.
. Paired corolla lobes and stamens stalked at maturity, me bases of ad-
jacent pairs separate and not surrounding the ovary, even in bud; anther
lobes aie in proportion to the connective (Fic. 4, D & E); flowers
purplish-pink. oe MISCHOPETALA.
Sp Se hia | one LIGUSTRINA.

—

No

~
©
°
oa
mee,
S
—
°
om &
is*)
mn
_.
Be
ie
a
iw]
Q
_—
fe]
5°
&
o
ou
Cc
a
o_
~
o
w
iss)

KEY TO THE SPECIES OF NOTELAEA

. Venation of leaf clear and more or less reticulate above.

—

1968]

GREEN, REVISION OF NOTELAEA 335

mately equal-sized marginal thickening to the lamina, ripe frui
broadly ovoid to subglobular, only a little longer than bro
4. Lamina narrowly ovate to narrowly lanceolate, (3.5- ~)6-10(- -16)

=

cm. long, margin usually entire, except for juvenile or sucker
growth, base acute or obtuse and long acuminate into the decur-
rent base, reticulation below usually at least a little less prominent
than that above, with primary veins usually curving forward
slightly towards the leaf apex before dividing, even in the broadest
leaves; inflorescence usually borne in the axil of the subtending
leaf, the lowest pedicels arising in the lower half of en inflores-
cence, often near the base; shrub or small tree to 7 m.
ol, SERS A eee ee ee ngifolia.
Lamina broadly ovate or ovate to, occasionally, itn: ovate
(rarely lanceolate or elliptic), (2.5-)3-9(-14) cm. long, margin
more or less crenulate, base rounded, although some of the largest
and narrowest leaves with acute leaf bases, but even then with
distinct if sometimes shallow and irregular crenulations, reticula-
tion below usually as prominent as that above with primary veins
more or less straight until they divide, not curving forward towards
the leaf apex; inflorescence usually a little above the axil of the
sear leaf, pedicels often arising only in the upper half of

e inflorescence: low shrub to 1 m. high. .......... 2. N. ovata.

3: Veins, after the primary ones, all more or less evenly, ae and
prominently reticulate, secondary ones not clearly merging with the
marginal thickening to the lamina; ripe fruit ellipsoid- sara clearly

longer than broad. 3.) ic is 2 eee

venosa.

2. Leaf venation below eee! more obscure than above, unless masked
by a velutinous indumen
5. Outline of leaves aneaaiaie or elliptic, (1-)1.5-5 cm. broad.

6.

i

= 0ae )0.3-1.5(-3) cm. broa

as

Reticulation of vein more or less prominent above,
the surface minutely but prominently punctate below; inflores-
cence 1-3 cm. long; paired corolla lobes, with stamens, sessile at

maturity; fruit 7-9 mm. long; central and northern east ain
land, - punctata

below not geese a punctate; inflorescence 4-8 cm ae
corolla lobes, with stamens, stalked at maturity (Fic. vy D iE:
fruit 18-20 mm. long; limited area of northeasternmost New

heasternmost Queensland. ........
South Wales and southe Q aos

Ree rr ee hee ee Cas get aa
Venation of upper leaf surface more or less clear and beg the

primary veins making an angle of 15 °_30° with the ls a
(3-)5-15(-30) mm. broad; Queensland and the central and north-
ern western-slope country of New South Wales, .....--------+-

336

JOURNAL OF THE ARNOLD ARBORETUM [ VoL. 49

slightly raised primary veins which make an angle of about 40°
with the midrib; lamina (2—)3-8(-14) mm. broad; — table-
nds country of mew South Wales... ......aca.. . N. neglecta.

i — of leaf obscure above.

”

ves linear to narrowly oem more than 7, usually at least 10

Baa as long as broad; ripe fruit 5—7 mm. long.

9. Breadth of leaf (1. 5—)2-5(- A. mm., venation totally obscure; in-
oo 4-10 mm. long; southern Queensland and northern table-
lands country of New South Wales. ................ 6. N. linearis.

9. Sd of leaf (2—)3-8(-14) mm., venation obscurely visible: in-
florescence 10-20 mm. long; central ‘tablelands of New South Wales.
ee ae RU eaters oe aa ee ae ee Oe 7. N. neglecta.

8. Leaves broadly = spies lanceolate, up to 7 times as long as broad;

ripe fruit 8-20 m
10. Primary ie phon veins obscure above and below, leaves (2—)3-

acute (Fic. 3, A); fruit 8-12 mm. long, varicolored; Tasmania,
Victoria, and southeasternmost New South Wales. ..............
Ieee Fre tie Pelee ee ha Oe hs ee a kg 8. N. ligustrina.

10. Primary veins just visible below, leaves (4~-)7—10(-12) cm. long;
inflorescence 4-8 cm. long; petals rounded, enveloping the stamens
(Fic. 4, D & E); fruit 18-20 mm. long, siceaprsansene2 apie
most New South Wales and southeasternmost —
es tye RS ik ee a Se OES Ew Veh nes SIE a johnsonit.

Notelaea sect. Notelaea
Sect. Eunotelaea Knoblauch in Engler & Prantl, Nat. Pflanzenfam. 4(2): 10.
1892.

The largest and most widespread section, characterized by the indu-
plicate-valvate aestivation of the corolla. It is closely related to sect.
Licustrina which, however, exhibits imbrication of the corolla in the bud.
In most species the ripe fruit i is invariably purplish-black in color but in
Notelaea punctata it has been recorded as “varicolored,” in NV. linearis as

“white and translucent” or “rich blue” and in N. punctata as white and
red, as well as dark purple to blackish.

I.

Notelaea longifolia Ventenat, Choix de Pl. 25. #.25. 1804; Desfon-
taines, Hist. Arbr. 1: 120, 1809; R. Brown, Prodr. 523. 1810; Sprengel,
Syst. Veg. 1: 35. 1824; A. DC. in DC, Prodr. 8: 291. 1844; Ettingshau-
sen, Blatt-Skel. Dikot. 69. ¢.26, fig. 9. 1861; Reichenbach f. Neuholl.
Pfl. Amelia Dietrich 9. 1866; Bentham, Fl. Austral. 4: 229. 1868;
F. Mueller, Fragm. Phytogr. Austral. 8: 43. 1873, & Syst. Census
Austral. Pl. 92. 1882; F. M. Bailey, Synop. Queensl. Fl. 303. 1883;
F. Mueller, Key Syst. Vict. Pl. 2: 39.1885, & 1: 357: 1887-8; F. M.
Bailey, Queensl. Woods, 85. 1888 & 97. 1889: F. Mueller, Second Syst.
Census Austral. Pl. 156. 1889; Maiden, Useful Native Pl. Austral. 579.
1889; F. M. Bailey, Cat. Pl. Queens]. 29. 1890; C. Moore, Handb. Fl.
New S. Wales 320. 1893; F. M. Bailey, Queensl. Fl. 3: 973. 1900;

1968 | GREEN, REVISION OF NOTELAEA 337

Dixon, Pl. New S. Wales 214. 1906; Guilfoyle, Austral. Pl. 40 [1911];
F. M. Bailey, Comprehensive Cat. Queens]. Pl. 311, 319. fig. 292.
[1913]; Maiden & Betche, Census New S. Wales Pl. 172. 1916; Ewart,
Handb. Forest Trees Vict. Foresters 407. 1925; Francis, Austral. Rain-
Forest Trees 327. 1929; Domin, Bibliot. Bot. 22(89) (Beitr. Fl.
Pflanzengeogr, Austral. 1(3): 512): 1066. 1929; Ewart, Fl. Vict. 941.
1930; Audas, Native Trees Austral. 242 [1935]; Anderson, Trees of
New S. Wales, ed. 2. 132, 271, 338. 1947; Bean in Chittenden, Dict.
Gard. 3: 1379. 1951; Beadle, Evans & Carolin, Handb. Vasc. Pl.
Sydney Distr. & Blue Mts. 348. 1963.

Evergreen shrub or small tree up to 7 m. tall (sometimes to 14 m. fide
Anderson, loc. cit.), bark finely fissured, finely pustular to fairly smooth,
mid to dark gray-brown, blaze granular; young stems densely puberulent
or velutinous to glabrous. Leaves glabrous, glabrate, puberulous or velu-
tinous above or below; petioles glabrous to velutinous, (2~)5—-15(-25)
mm. long; lamina lanceolate, narrowly ovate or elliptic, occasionally nar-
rowly lanceolate or ovate, (3.5—-)6-10(-16) cm. long by (1—)2-3.5(-5.5)
cm. broad; margin somewhat thickened, entire or obscurely and somewhat
irregularly crenulate; apex acute or long acute, finely pointed if undam-
aged; base acute to obtuse, long acuminate into the petiole; venation
reticulate and raised above and below, except when obscured if densely
velutinous, veins below usually a little less prominent than those above,
(5—)6-8(-10) primary veins per side arising at an angle of 40°-70° with
the midrib, usually curving forwards slightly towards the apex before
branching, even on the broadest leaves, basal pair of primary veins usually
arising at an acute angle in the decurrent base of the lamina. In florescence
axillary, 1(—2) per axil, decussate, lowest pedicels usually in the lower
half, often near the base, glabrous or puberulous, often minutely so and
scattered, 1-2.5 cm. long, lengthening after anthesis, (5—)7~11(—13)-
flowered; upper bracts 1-2 mm. long, thin, ovate to lanceolate, puberulous
or glabrous and minutely ciliolate, early deciduous; basal pair thick, pubes-
cent to glabrous (in northeastern plants), 1-3 mm. long, acute-acuminate,
deciduous except for the persistent fused basal portion. Flowers hermaph-
rodite, yellow or pale yellow; pedicels 1-4 mm. long, accrescent in fruit.
Calyx glabrous, 0.5-1 mm. long with 4 more or less triangular lobes, often
acutely pointed, 0.5-0.8 mm. long. Corolla induplicate-valvate, lobes 4,
1.5-2.5 mm. long, in pairs joined in the lower half with the filament,
concave. Stamens 2, 1-2 mm. long, more or less enclosed within the con-
cave petals; anthers 1-1.5 mm. long with a distinct terminal appendage
0.2-0.3 mm. long; filaments short, 0.5 mm. long. Ovary flask-shaped,
1.2-1.5 mm. long, tapering into a style about 0.5 mm. long with 2 short
stigmatic lobes 0.2 mm. long. Drupe ovoid, dark purple or bluish ney.
1-1.6 cm. long by 0.8-1.2 cm. broad; endocarp hard, pointed ovoid, 8-1
mm. long by 5-8 mm. broad, 0.3—0.5 mm. thick.

Notelaea longifolia is the best known and most widespread species of the

338 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

genus, with a distribution stretching from the area of Bateman’s Bay in
southern New South Wales to the southern part of the Wide Bay district
of Queensland. A number of formal taxonomic varieties have been de-
scribed but the most striking, var. velutina, is here transferred to N.
microcarpa (see below under that species). When this is done, and a large
number of specimens are critically redetermined, especially after the care-
ful delimitation of N. ovata, N. venosa, N. punctata and N. microcarpa,
one is left with a far more homogeneous species in which the only obvious
variation is that in degree of pubescence. When the localities for the speci-
mens are plotted on a map, together with a symbol to indicate their
hairiness in three simple categories of pubescent, glabrate or glabrous, it
is found that N. longifolia occupies a coastal strip and exhibits a cline in
the degree of pubescence, with velutinous or pubescent plants in the south
and glabrous plants in the north (Map 1). The southernmost specimens
are most densely hairy, often velutinous, while around Sydney the amount
of pubescence varies greatly; often densely velutinous and only slightly
velvety specimens being found in the same area (e.g. Johnson & Constable
NSW 52347 & 52348 from Little Forest, 7 miles WNW. of Milton, 10 Sept.
1960). In northernmost New South Wales and in Queensland, NV. longi-
folia is quite glabrous, and in the intervening area the leaves of the speci-
mens examined are all glabrate with a fine puberulence on the young stems
and bud scales only. Considerable thought has been given to the question
of whether this cline should be divided taxonomically; clearly any division
will be artificial, but it is believed that names would be useful, for fre-
quently it may be convenient to refer to a plant exhibiting one of the
extremes of pubescence. This then raises the question of the taxonomic
rank to be employed. None of the formal infraspecific ranks are partic-
ularly suitable yet the appropriate biosystematic term of clinodeme ex-
pressly excludes a nomenclatural connotation. Diagnosis, in this case, is
dependent on variation in one character only, that of degree of pubescence,
so although a distinct distributional pattern in this character exists, the
ranks of subspecies and variety seem of too great a consequence so the
conventional rank of form is chosen and appropriate names are published
below. (Incidentally, the use of variety would compel the employment of
Domin’s highly inappropriate epithet of decomposita for the glabrous plant
from the northern end of the range — see below.

In annotations on the Sydney herbarium covers L. A. S. Johnson has
summarized the habitat occupied by Notelaea longifolia: ‘Sometimes
found within rainforest and often on rainforest margins but also common-
ly in eucalypt forest, though not on the poorest siliceous soils. In the poor
Hawkesbury sandstone country it is found chiefly in the gullies or near
salt-water estuaries (often on rocky slopes here) where there is some
enrichment of the soil nutrients.” Also, “in gullies in wet sclerophyll forest
— sometimes in patches of rainforest.”’

In addition to being the most widespread species, Notelaea longifolia
also appears to be central in its relationship to the others. Perhaps it
represents the descendants of the basic stock from which the other species

1968} GREEN, REVISION OF NOTELAEA

339

QUEENSLAND

alia
% = ~ + eo ae
‘ is — 2

NEW SOUTH WALES

iN a ae 4 ; 8
‘ NOTELAEA LONGIFOL
5 @ - LONGIFOLIA
< © fF INTERMEDIA

VICTORIA : © £ GLABRA

JR SUPPOSED HYBRID WITH
N. MICROCARPA

Map 1. Distribution of Notelaea longifolia and forms.

340 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

have evolved, for each of the other species of sect. NoTELAEA appears very
closely related. Where they meet, hybridization seems to take place oc-
casionally with V. microcarpa, a species characteristic of the western- slope
country. The distribution of those specimens of possible hybrid origin
examined in this revision is indicated on Map 1.

Forma longifolia
“Olea apetala” Andrews, Bot. Rep. 5: ¢.316. 1803, up hoes (1794).
2.

Notelaea rigida Desfontaines, Hist. Arbr. 1: 120. 1809.

Postuera longifolia (Vent.) Rafinesque, Sylv. reals 10. 1838.

Notelaea “ovata” Endlicher, Icon. a Pl. xi. ¢.55. 1838, non R.Br. (1810);
Lindley, Veg. Kingdom 616. fig. 1

N. nervosa Se aan f.) Steudel, as ed. 2. 2: 198. 1841; A. DC. in
DC. Prodr. 8: 291. ;

? N. laurifolia Kunth, rae Sem. Hort. Berol. 12. 1846, & in Walpers, Repert.
Bot. Syst. 6: 742. 1847.

N. longifolia Vent. var. candolleana Domin, Bibliot. Bot. 22(89) (Beitr. Fl.
Pflanzengeogr. Austral. 1(3): 514): 1068. 1929

N. longifolia Vent. var. rigida (Desf.) Domin, Joc. ‘it. 1066. 1929.

N. longifolia Vent. var. typica Domin, loc. cit. 1929.

Young stems, petioles, leaves (especially below) and inflorescence
rachides more or less densely pubescent to velutinous. The pubescence
of the leaf sometimes short and inconspicuous but more or less persistent,
at least below.

SELECTED ExsIccaTAE. New South Wales. Bateman’s Bay, June 1906, Boor-
man (NSW 33491); Boyne State Forest, NE. of Nelligen approx. 180 m. alt.,
22 Nov. 1966, Pullen 4203 (cANB 161997-9); Currawon State Forest, near
Bateman’s Bay, June 1959, LIW. & J.G.T. 3663 (prt 35461); Lake Burrill,
S. of Milton, 12 Jan. 1937, Hadley 2404 (Rodway 6239) (kK, Nsw 33416); near
Canjola, South Coast, 6 Dec. 1937, Hadley 2620 (Rodway 6236) (K, NSW
33412); Comerong Is., Shoalhaven R., 18 Sept. 1932, Rodway 880 (6231) (K,
nsw 33408); Cabbage-tree Creek, against the road between Nowra & Currarong,
22 May 1964, Willis (MEL 19628); gully near Barber’s Creek (Tallong), Forsyth
& Hamilton (Nsw 33483); Illawara (‘Sydney woods”), Macarthur 174 (x,
syntype of var. candolleana; BM, isosyntype); Cronulla, Oct. 1919, Cheel (NSW
33528); Sydney, Clowes (kK, syntype of var. candolleana); Port Jackson, 1803,
R. Brown (“2849”) (pM, K); Manley Beach, Nov. 1863, Wilhelmi (LE, P); The
Spit, Middle Harbour, Sydney, 14 Nov. 1949, Constable (x, Nsw 11316); Horns-
by, May 1917, Blakely (a, Bri 7934, c, Nsw 33453); near Harry’s R., central
Cox's. R., 5. of Blue Mts., 23 Dec. 1951, Whaite 1096 (K, NSW 22139); Little
Wheeny Creek to Kurrajong Heights, about 350 m. alt., 10 Aug. 1953, Johnson
(K, NSW 24963); Pulbah Island, Lake Macquarie, 27 Oct. 1941, Messmer, Bryce

Rupp (Nsw 33447); 6 miles N. by W. of Raymond Terrace, 1 May 1960,
Story 7275 (CANB 80051): Flyer’s Creek, near Buladelah, Apr. 1924, Rupp 21
(nsw 33555); Crawford R., near Buladelah, Oct. 1902, Cheel (Nsw 33442);
Wallamba State Forest, near Na biac, 12 Aug. 1 955, M cDonald (nsw 33516);
Waukivory Creek, Gloucester, Sept. 1897, Maiden (Nsw 33500); 16 miles W.

1968] GREEN, REVISION OF NOTELAEA 341

of Dungog, 5 May 1960, Story 7346 (canB 80364): 10 miles NNE. of Ravens-
worth, 23 March 1960, Story 7145 (cANB 80379). Without locality: 1770,
Banks & Solander (BM, £, ? isotypes of Rhisospermum nervosum); Hb. Horne-
mann, ded. Desfontaines (c, ? isotype of N. rigida); Caley (BM, G).

Cultivated. AustraLia: Botanic Gardens, Sydney, Feb. 1917, Blakely (Nsw
33549); Botanic Gardens, Adelaide, 20 Dec. 1930, J. F. Bailey (K); Botanic
Garden, Hobart, 31 Jan. 1932, White 8246 (a, prt 7944); Botanic Gardens, Mel-
bourne, F. Mueller (BM, MEL 19623). FRANCE: ex hort. Cels, ex herb. Ventenat
(c, lectotype) & ex herb. Desfontaines (¥1, ? isolectotype); Hort. Paris, Orange-
rie, 23 Oct. 1819, ex herb. J. Gay (x, topotype of and labelled as N. rigida
Desf. Arb.); Jardin des Plantes de Paris, 1818, Perrottet (Gc). GERMANY: Hort.
Berlin, without collector (c). Great Britain: Hort. Kew, 1794, without collec-
tor (Bm); Isles of Scilly, Tresco Abbey, 18 Sept. 1953, Dorrien-Smith (x).

This form is distinguished from the others by the more or less densely
pubescent or even velutinous indumentum on the young stems, leaves, and
inflorescence rachides. The type, a cultivated specimen, is distinctly hairy
(although less pubescent than much of the wild material) and, at the turn
of the 18th century, was almost certainly introduced from the Port Jackson
area, from which most of the early introductions of Australian plants were
made.

Typification has not been straightforward. In all probability Notelaea
longifolia was originally described by Ventenat from a living plant grown
in the garden of M. Cels and it is possible that no type specimens, as we
know them today, were prepared. However, there is no doubt about the
identity of the species for the protologue was beautifully illustrated and,
thanks to the help of M. Gilbert Bocquet of Geneva, to whom I express my
thanks, I learned that a specimen from M. Cels’ garden, which had been in
the Ventenat herbarium, is today in the Geneva herbarium. This I have
examined on loan and designated as lectotype. ‘

Notelaea rigida was similarly described from a cultivated plant, this
time from the Orangery in the Jardin des Plantes in Paris and a specimen,
although collected some nine years later, has been seen in the Kew Her-
barium with this origin and this name attached. The genus Rhisospermum,
described by C. F. Gaertner, was based on fruit from the collection of Sir
Joseph Banks, who accompanied Capt. Cook on the first of his famous
voyages; no authentic type has been seen but there is a Banks & Solander
collection at the British Museum (Natural History), with a duplicate in
Edinburgh, unfortunately without fruit in both cases. oe

Notelaea laurifolia, described by Kunth in 1846 from a plant cultivat
in the Berlin Botanic Garden, was attributed to New Zealand, but, as
far as is known, none of the related New Zealand plants had been in-
troduced at that time and the admittedly somewhat inadequate description
fits V. longifolia. The type in the Berlin Herbarium was presumably
stroyed during the war, so that the exact identity of Kunth s plant ie
remain uncertain until a duplicate or other authentic specimen is perhap
found elsewhere. For the present the best disposition of the name seems
to be under N. longifolia.

342 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

The typification of the varieties described as new by Domin was more
difficult at first sight but, on learning that the specimens upon which he
had based his new taxa, other than those which were in his own collection,
are often at Kew, and usually without annotation, the types were easily
located by comparison of the labels with his protologues. NV. longifolia var.
decomposita is dealt with under f. glabra below. Var. candolleana is a hairy
form with narrowish leaves. The reference to “Sydney woods” on the label
of the Macarthur syntype seems to have been taken by Domin as a locality
whereas the reference is to a timber or wood specimen, No. 174, sent from
Sydney to the Paris Exhibition of 1854. Finally, var. pedicellaris was
found to belong to NV. venosa, under which species it is now included.

A specimen, Cambage 2702, collected on 15 May 1911, between Baerami
and Dunman in the region of the Upper Hunter in New South Wales
(Nsw 33540) appears of hybrid origin with NV. microcarpa. The leaf shape
is like that of N. microcarpa but the fruit, which is immature, is too large
for this species and looks as though it would have developed to the size
normal for NV. longifolia.

The extent to which the foliage of juvenile, sucker or other strongly
growing shoots differs from that of adult plants cannot be settled with
certainty from herbarium collections, unless these are specially made, but
in Notelaea longifolia they appear to be larger and broader than usual,
more markedly reticulate and with the margin crenulate. Pullen 4203
from Boyne State Forest, New South Wales, is represented in the Canberra
Herbarium by three sheets, two of which show normal adult foliage, one
with fruit, while the third (cANB 161998) has large broadly ovate-lanceo-
late leaves, 10-13 cm. long by 4.5—5.5 cm. broad with rounded base and
short petioles 2~3 mm. long. The field notes say “leaves of sterile branch
larger than those of adult branches.” Specimens in the Sydney Herbarium
annotated as juvenile consist of unattached leaves 18-25 cm. long by
8.5-9.2 cm. broad with petioles 10-15 mm. long which, although not
particularly rounded at the base, are strongly reticulate and more crenu-
late than usual. These specimens are: Patonga, N. bank of Hawkesbury
R. near the mouth, N.S.W., Sept. 1936, Blakely (Nsw 33449) and Bul-
bararing Lake, near Kincumber, N.S.W., 17 June 1905, Maiden (NSW
33448). :

Forma intermedia P. S. Green, f. nov.

Caulibus junioribus et bracteis externis gemmarum minute puberulis;
inflorescentiarum rachidibus minute et dissite puberulis; foliis junioribus
et petiolis glabratis dinoscenda.

SELECTED ExsIccAaTAE: New South Wales. Wattagan State Forest, W. of Dora
Creek, 15 Sept. 1954, Constable (x, Nsw 30732); Port Stephens, Aug. 1911,
Boorman (nsw 33445); Dungog to Clarencetown, 29 May 1934, Fraser (NSW
33443); Clarencetown, 7 Nov. 1936, Fraser & Vickery (NSW 33444): 16 miles
ENE. of Singleton, 26 March 1960, Story 7195 (cans 78754, K); Port Mac-
quarie, Oct. 1892, Maiden (a); Ellenborough to Long Flat, 19 Oct. 1953, John-
son (K, NSW 26257); Ralf's Riverlet, Nov. 1897, Maiden (nsw 33441); Carrai
Carrai State Forest, 30 miles approx. W. of Kempsey, 11 June 1958, Constable

1968 | GREEN, REVISION OF NOTELAEA 343

(kK, holotype; Bm, Nsw 46234, isotypes); Nambucca Heads, June 1910, Boorman
(Nsw 33440); Bellingen, Dec. 1938, Hewitt (Nsw 33437); Dorrigo State Forest,
about 750 m. alt., 4 Oct. 1930, White 7513 (BRI 7937); Coff’s Creek, Coff’s
Harbour, 4 Nov. 1963, Hayes in Green 1515 (a, K); Glenugie Peak, 6 Nov.
1963, Green 1525 (A); Brickmaker’s Creek, 4 miles SE. of Ramornie Crossing,
July 1922, Blakely & Shiress (Nsw 33476); Smith’s Creek, 7 miles NE. of
Ramornie Crossing, July 1922, Blakely & Shiress (Nsw 33473); Coal Ridge, 10
miles N. of Ramornie Crossing, July 1922, Blakely & Shiress (Nsw 33472);
Harwood Is., Clarence R., 10 Feb. 1922, Helms 458 (a, c); Ballina, Sept. 1892,
Bauerlen 853 (a, Nsw 33464, z); Dunoon Road, 7 Nov. 1963, Hayes & Floyd
1 (A); Casino, Dec. 1916, McAuliff (Nsw 33468-33470); Koyogle, Dec. 1917,
McLean (c, nsw 33467); Drake, Oct. 1901, Boorman (Nsw 44087); Acacia
Creek, via Killarney, May 1905, Dunn 112 (Nsw 44088).

Cultivated. AusTrALIA: Garden Palace Grounds, Sydney, Apr. 1898, Cam-
field (Nsw 44057); Botanic Gardens, Melbourne, F. Mueller (MEL 19624).

Introgression between Notelaea longifolia f. intermedia and N. micro-
carpa var. velutina appears evident from the small population of V otelaea
I examined on 5 Nov. 1963 adjacent to the Nymboida River near Dorrigo,
New South Wales. Specimens were collected from each of the small trees
seen, Green 1519 at (A), which exhibit a considerable degree of inter-
mediate variation in leaf shape, veining, and pubescence. It is suggested
that in the not too distant past, with the felling of the forest and estab-
lishment of agriculture, there has been considerable ecological upheaval in
this area and that NV. longifolia and N. microcarpa, which were formerly
separated, came into contact. Hybridization between f. intermedia and
N. microcarpa seems likely for a few other intermediate collections from
New South Wales which are cited here.

Paddy’s Land State Forest, about 900 m. alt., 12 Apr. 1955, Turner 39 (NSW
33535); Unumgar State Forest, 13 March 1963, Jones 2370 (cANB 136002); &
near Mt. Lindsay, Dec. 1943, Jones (BRI 7924); and from the Moreton District
of Queensland: the Ranch, foot of Wilson’s Peak, 17 Jan. 1933, Michael 1983
(BRI 7920, Nsw 33532).

Forma glabra P. S. Green, f. nov.

Caulibus junioribus, foliis, bracteis externis gemmarum et inflorescen-
tiarum rachidibus manifeste glabris.

Notelaea reticulata A. DC. in DC. Prodr. 8: 291. 1844; Ettingshausen, Blatt-

Skel. Dikot, 69. ¢.22, fig. 5. 1861.
N. longifolia Vent. hay eae Fs Domin, Repert. Sp. Nov. 12: 96. ete
& Bibliot. Bot. 22(89) (Beitr. Fl. Pflanzengeogr. Austral. 1(3): 514):
<a Queensland. MorETON District: Mouth of Currumbin
Creek, 7 Nov. 1929, White 6509 (srt 7931, holotype; A, isotype) ; Tamborine,

344 JOURNAL OF THE ARNOLD ARBORETUM [voL., 49

K); Ithaca Creek, Brisbane, 30 Oct. 1949, White 13149 (BRI 42314); between
Brisbane and Redcliffe, bank of South Pine R., 16 Dec. 1930, Hubbard 5499
(prt 7922; K); Moreton Bay, Sept. 1829, A. Cunningham (Gc, type

reticulata; K, isotype) & islands, Moreton Bay, Sept. 1829, A. Cunningham 8
A, BM, K, ? isotypes of N. reticulata); at the creek of Mr. Archer’s Station

E. of Kilcoy), 15 Sept. 1843, Leichardt (nsw 33459); Eumundi, Nov. 1892,
Simmonds (a); Noosa Heads, 26 May 1951, Johnson (NSW 33460). Wipe Bay
District: Lake Cootharaba, Keys (srt 7910 & 7930); Imbil, March 1918,
Weatherhead 393 (pri 7933).

New South Wales. Dunoon, 7 Nov. 1963, Green 1527 (a, K); mouth of
Brunswick R. by Pacific Highway, Brunswick Heads, 30 May 1964, Schodde
(& Hayes) 3549 (canB 135634-5); Tweed Heads, Nov. 1903, Maiden &
Boorman (Nsw 33463).

Cultivated. AusTRALIA: Botanic Gardens, Sydney, Dec. 1895 & Feb. 1898,
Camfield (Nsw 33557) & Dec. 1925, Blakely & Anderson (NSW 33558).

At the northern end of its range Notelaea longifolia f. glabra is not
always easy to distinguish from N. punctata. The distributions of the
two overlap slightly, as represented, for example, by the collections of
Keys from Lake Cootharaba (Bri 7910 & 7930), Francis from Cootharaba
(prt 7932) and Weatherhead from Imbil (srt 7933 & 9697), all in the
Wide Bay District of Queensland. Where there is fruit there is little dif-
ficulty in identification: BRrt 7932 & 9697 have small fruits characteristic
of NV. punctata, whereas prt 7933 has the larger ones typical of NV. longi-
folia. Vegetatively the prominence of the secondary veins below appears
to be the best character for distinguishing most material, at least in the
dried condition, for in both species the leaves are punctate. Habit appears
significant and herbarium studies alone are inadequate to settle satisfac-
torily the problem of specific differentiation in this instance. Careful field
observations, coupled, perhaps, with biosystematic studies, are called for.

Examination of the type of Notelaea reticulata DC. from Moreton Bay
showed it to be the glabrous expression of the species, as might be expected
from the locality from which it was collected. Similarly, the type of
Domin’s var. decomposita, from the Brisbane River, is the glabrous form
and was found to be based in part on a very floriferous specimen (actually
with the flowering stage just past) from the flowering shoots of which the
leaves had just dropped, so that the inflorescences might, at first glance,
be interpreted as compound.

2. Notelaea ovata R. Brown, Prodr. 524. 1810; Sprengel, Syst. Veg. 1:
oo. 1824; A. DC. in DC. Prodr. 8: 291. 1844; Ettingshausen, Blatt-
Skel. Dikot. 69. ¢.26, figs. 7 & 8. 1861; Bentham, Fl. Austral. 4: 299.
1868; F. Mueller, Fragm. Phytogr. Austral. 8: 43. 1873, & Syst. Census
Austral. Pl. 92. 1882; F. M. Bailey, Synop. Queensl. Fl. 303. 1883; F.
Mueller, Second Syst. Census Austral. Pl. 156. 1889: Maiden, Useful
Native Pl. Austral. 580. 1889; F. M. Bailey, Cat. Pl. Queensl. 29.
1890; C. Moore, Handb. Fl. New S. Wales, 320. 1893: F. M. Bailey,
Queensl. Fl. 3: 972. 1900; Dixon, Pl. New S. Wales, 214. 1906; F. M.

1968 | GREEN, REVISION OF NOTELAEA 345

Bailey, Comprehensive Cat. Queens]. Pl. 311. 1913; Maiden & Betche,
Census New S. Wales Pl. 172. 1916; Audas, Native Trees Austral. 242,
[1935]; Bean in Chittenden, Dict. Gard. 3: 1379, 1951; er Evans
& Carolin, Handb. Vasc. Pl. — Distr. & Blue Mts. "348. 1 963.

N. longifolia Ventenat var. ovata (R. Br.) Domin in Bibliot. Bot. 22(89)
(Beitr. Fl. ghenisg iar Adsl, 1 (3): 513): 1067. 1929, quoad syn., excl.
fig. et spec.

Evergreen shrub 0.5-1 m. tall, usually with several stems from an old
common stock; young stems densely puberulous, sometimes minutely so.
Leaves glabrous or minutely puberulous above and below when young,
especially on the midrib and veins, rarely subvelutinous; petioles puberu-
lous, sometimes minutely so, 1-6(-10) mm. long; lamina thickish, more

er MM 2-

fe eR

Fic. 1. Notelaea ovata. A, flower; B, young bud; C, ovary. (Drawn from
Green. 1522a).

or less coriaceous, punctate, very broadly ovate to ovate to narrowly ovate,
more rarely lanceolate or broadly elliptic to elliptic (or even narrowly
elliptic), (2.5~)3-9(-14) cm. long by (1.2—)1.8-3(-6) cm. broad; margin
crenulate, sometimes irregularly so, rarely almost entire, somewhat thick-
ened; apex acute, sometimes obtuse, occasionally slightly acuminate, tip
often minutely apiculate; base rounded (to rarely subcordate, or, on the
narrowest leaves, more or less acute), attenuate into the petiole; venation
reticulate and raised above and below, (5~)6—-7(-11) primary veins per
side. Inflorescence axillary, 1(—2) per axil, in the case of a single inflores-
cence up to 5 mm. above the axil with an unopened bud in between, decus-
sate, lowest pedicels often in the upper half, puberulous or scattered and
minutely puberulous, especially towards the base, (1—-)1.5-3.5 cm. long,
(3—) 5-9(—11)-flowered; occasionally freakish, up to 6 cm. long with
secondary branching and 15 or more flowers per inflorescence; upper bracts

346 JOURNAL OF THE ARNOLD ARBORETUM [ VoL. 49

1-3 mm. long, lanceolate, sometimes almost subfoliaceous, minutely puber-
ulous, early deciduous; basal bracts, thickish puberulous, but most often
raised above the axil 3-10 mm. on the inflorescence axis. Flowers hermaph-
rodite (Fic. 1), pale yellow; pedicels 1.5-6 mm. long. Calyx glabrous
0.5-1 mm. long, with 4 more or less triangular lobes, slightly erose or some-
times minutely ciliolate, 0.5-0.7 mm. long. Corolla induplicate-valvate,
lobes 4, 1.5-2.5 mm. long, in pairs joined in the lower half, concave.
Stamens 2, 1-1.75 mm. long more or less enclosed within the concave
petals, with a slight terminal appendage; filaments short 0.5 mm. long.
Ovary flask-shaped, 1—-1.5 mm. long tapering into a style 0.3-0.5 mm. long
with 2 short stigmatic lobes 0.2 mm. long. Drupe ovoid with whitish
semitransparent sarcocarp (fide B. S. Cole) 1—1.25 cm. long (?) by 0.8-1
cm. broad @); endocarp hard, pointed ovoid, 7 mm. long by 5 mm. broad,
0.3 mm, thic

SELECTED ExSICCATAE. Queensland. Moreton District: Coolum, April 1945,
Clemens (Bri 9704); Mt. Coolum, 24 March 1945, Clemens (kK); Beerwah, Sept.
1919, White (prt 9718); Elimbah, 4 Dec. 1961, McKee 9731 (cANB 120610);
top of Mt. Tunbubudla, Glass House Mountains, about 300 m. alt., 10 Aug.
1930, Hubbard 3606 (a, K); Kedron, near Brisbane, Shirley (prt 9720); Din-
more Pottery, 19 Oct. 1964, L. S. Smith 12171 (srt 64521, K); Taylor Range,
near Brisbane, about 200 m. alt., 24 Aug. 1930, Hubbard 3757 (a, K); Brisbane
River, 1863-1865, Dietrich (sm); hills near Mt. Gravatt, near Brisbane, 11
July 1930, Hubbard 3317 (a, Bri 9708, K); about 5 miles SE. of Ipswich, 16
June 1957, Pedley (x); Russell Island, Sept. 1913, White (prt 9709, NSW
44035); Stradbrooke Island, Scortechini & Bailey (prt 9710); Mount Maroon,
NE, gorge, about 840 m. alt., 10 March 1962, Everist 7111A (BRI 38264); Coo-
mera R., near Canungra, 9 Oct. 1949, White 13124 (Bri 42315).

New South Wales. Trial Bay, Feb. 1882, Betche (Nsw 33551); Barcoogere
State Forest, about 6 miles N. of Corindi, 6 June 1957, Johnson & Constable
(Nsw 47501) & 6 Nov. 1963, Green 1522 a-f (a); Hastings River, Beckler (K);
Kendall, Sept. 1932, Bailey (a, Nsw 44036); Nelson’s Bay, Port Stephens, 6
Feb. 1937, Vickery (LE, Nsw 44037); Wallsend, Oct. 1899, Boorman (NSW
44039); Killara district, Oct. 1948, Price (Nsw 44042);
1803, Caley (pm); Lindfield, 23 Oct. 1927, White 5139 (prt 9717, K); Port
Jackson, R. Brown (BM, holotype: E, K, P, isotypes); Hurstville, April 1901,
Camfield (Nsw 44050); Bargo River, Dec. 1902, Boorman (a, Nsw 44052);
Huskisson, Jarvis Bay, Dec. 1925, Rodway (x).

Although the epithet borne by this species is very appropriate for most
individuals and specimens, the leaves are fairly variable, both in size and
in the proportions of length and breadth, so that on occasion the outline
closely resembles that of Notelaea longifolia. However, the species may
be separated by the more regular and prominent reticulation of the veins
in NV. ovata and the more definitely crenulate margin
Map 2 shows that Notelaea ovata is a coastal species, where it is found
in Eucalypt forest on sandy soils or, in the Sydney district, according to

. A. S. Johnson, on soils derived from argillaceous shale, and i in Queens-
land, to judge from the field notes with herbarium specimens, on sandstone
hills and ridges. The tallest growth recorded on the labels of all the speci-

19

68 | GREEN, REVISION OF NOTELAEA 347

b
&

it
@-NOTELAEA LINEARIS |
D-NOTELAEA OVATA

\

QUEENSLAND QUEENSLAND

NEW SOUTH WALES

V-NOTELAEA NEGLECTA
a- NOTELAEA JOHNSONII

sO

Map 2. Distribution of Notelaea ovata, N. linearis, N. neglecta and N. john-
nit,

348 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

mens examined is 4 feet (1.2 m.) but whether the species ever attains
higher growth than this is not recorded. There is no doubt, both from
personal observation in the field and from the notes and comments on
labels, that plants are often cut back by fire and then regenerate from a
woody basal stock, often of considerable age. Plants flower when only
a foot or so tall but, to judge from the specimens examined, rarely fruit.
In fact the range in fruit size and color is somewhat uncertain. A specimen
in the Sydney herbarium collected at Medowie by R. S. Cole (Nsw 44038)
has fruits in an envelope which appear as though they were fleshy when
ripe; the mesocarp has shrivelled and it is difficult to be sure of the exact
dimensions when they were fresh, so that the sizes given in the description
above are only an estimate. The fruits on this specimen are said to have
been whitish with a semitransparent sarcocarp, although purplish-black
is the color one might expect from related species, and the field notes with
a specimen collected by Pedley near Ipswich, Queensland (x) state that
the fruits are purple. Unfortunately, there are no fruits still attached to
the specimen of this collection at Kew, so that it is impossible to compare
them with those of the Cole collection and impossible to be certain of the
correctness of the statement in Bentham’s Flora Australiensis (4: 299)
that this species has the “fruit of NV. longifolia.’ Just how variable the
fruit color and size can be is not known but it is worth bearing in mind
that the fruit of NV. ligustrina may vary from purple through red and pink
to white. Observations on fresh fruiting material are badly needed.

In Barcoongere State Forest, New South Wales, I collected shoots from
six different plants in a population scattered over a small area in order
to record the variation in leaf size (Green 1522 a—-f). Lanceolate-elliptic
leaves 11 cm. long by 3.5 cm. broad, broadly ovate leaves 6.5 by 5 cm.,
and diminutive but ovate leaves 3.5 by 2 cm. were all represented, each
from a different individual. Bud material was also fixed for cytological
examination and given to Dr. Barbara Briggs of Sydney. From this she
obtained a chromosome count of n = ca. 23. I am most grateful to Dr.
Briggs for permission to publish this count.

The species of sect. NoTELaga are closely related but it is not known
to what extent they are interfertile. Chromosome numbers, such as they
are known, appear to be uniform, 2m = 46, but only one collection has
been seen which appears to be a hybrid of this species. This was collected
by L. S. Smith (no. 12180 a) at Dinmore Pottery, Moreton Bay District,
Queensland, on 19 Oct. 1964 in Eucalyptus fibrosa-E. maculata forest
on hillside (Brt 64516, K). This specimen has the leaf shape of a large-
leaved N. microcarpa but the margin and venation is that of NV. ovata.
L. 3. Smith collected undoubted NV. ovata at the same locality on the same
day (his no. 12171) and to judge from the distribution of N. microcarpa
this species is to be expected in this area as well. The hybrid specimen
bears a few very immature fruits and it is quite possible that the two
species are interfertile.

3. Notelaea venosa F. Mueller in Trans. Vict. Inst. 1854-55: 131

1968 | GREEN, REVISION OF NOTELAEA 349

(“Definitions of Rare or Hitherto Undescribed Australian Plants” 50).
1855, & in Hooker’s Jour. Bot. & Kew Gard. Misc. 8: 163. 1856;
Anderson, Trees New S. Wales, ed. 2. 132. 1947; Beadle, Evans &
Carolin, Handb. Vasc. Pl. Sydney Distr. & Blue Mts. 349. 1963.

N. longifolia Vent. var. pedicellaris Domin, Repert. Sp. Nov. 12: 96. 1913 &
Bibliot. Bot. 22(89) (Beitr. Fl. Pflanzengeogr. Austral. 1 (3): 514): 1068.
929.

Evergreen bushy shrub or small tree, 1.5-6(-9) m. tall; bark grayish,
smooth or eventually rough, flaky gray-brown (fide Schodde 3502); stem
glabrous or puberulous, especially when young, sometimes minutely so.
Leaves glabrous or minutely puberulous at first above and below; petioles
glabrous or minutely puberulous at first, 5-15(—20) mm. long (0.5-3(-8)
mm. long in northern plants); lamina thickish, more or less coriaceous,
obscurely punctate, lanceolate to occasionally narrowly ovate or elliptic,
(4—)6.5-12.5(-17) cm. long by (1—)2—4(—-6) cm. broad, in semi-juvenile
shade leaves (Nsw 44065 & MEL 19621) petiole 20-30 mm. long, lamina
ovate, 15-21 cm. long by 7.5—9.5 cm. broad; margin slightly thickened,
entire or very shallowly and finely crenulate, especially in the upper half,
sometimes extremely shallowly so; apex acute, sometimes slightly acumi-
nate; base acute, attenuate into the petiole, to obtuse (rounded or sub-
cordate in northern plants); venation raised, finely and evenly reticulate,
especially below, (6—)7—10(-—11) primary veins per side. Inflorescence
axillary, 1-2(—3) per axil with a bud or buds between it and the axil but
usually not more than 1.5 mm. above the axil, decussate, 1-3 (—7) cm.
long, 7-11 (—17)-flowered, often fairly dense and occasionally the terminal
5(—7) flowers subumbellate, glabrous; bracts 1.5-3 mm. long, broadly
ovate, ciliolate, early deciduous, basal bracts 1-2.5 mm. long, blunt, gla-
brous, somewhat persistent or the fused bases persistent. Flowers hermaph-
rodite, yellow or greenish-white, pedicels glabrous, 2—5(-10) mm. long.
Calyx 0.5 mm. long with 4 somewhat irregular, more or less triangular,
lobes. Corolla induplicate-valvate, lobes 4, 1.5-2.5 mm. long, joined in
pairs for about 0.5-1 mm., thickish, concave. Stamens 2, 1-1.75 mm. long
with a broad filament 0.3 mm. long and a slight terminal appendage. Ovary
broad flask shaped, 1-1.5 mm. long, tapering to an imperceptible style
with bilobed stigma 0.2 mm. long. Drupe dark purple to blackish Piha
and red, fide E. Gippsland 165-66, no collector, MEL 19626), slightly
glaucous, ellipsoid-ovoid 1.5-2 cm. long by 1-1.4 em. broad; epee
ovoid, pointed 1.1-1.5 cm. long by 0.8-1 cm. broad, 0.4-0.6 mm. thick.

SELECTED ExsiccaTAE. New South Wales. River Hastings, Fraser (K) : vs
liams River, 7 Jan. 1934, Fraser & Vickery (NSW 44063); Mt. poe so
miles E. of Aberdeen, 11 Oct. 1960, Story 7497 (CANB 85286, NSW 5385 ise 4
Royal, 25 miles E. of Aberdeen, 25 Mar. 1960, Story fire (ane gt es ‘
NSW 67494); ridge between Chichester R. and Kerripit R.., ard - ici di
1 Apr. 1954, McDonald (Nsw 43030); O’Sullivan’s Gap, 10 miles » : - -
delah, 30 Nov. 1946, Gilbert (Nsw 6498); Mt. Warrowolong, yee od me rs
24 Aug. 1949, Constable (Nsw 11517); Wandabyne, Nov. 1922, ly

350 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

oan (nsw 44064); track between Victoria Falls Lookout and Falls, about
2 miles E. of Mt. Victoria, 6 May 1959, Constable (Kk, Nsw 47326); Springwood,
5 es 1953, McKee 847 (k, Nsw 26323); Cedar Creek, Central Kowmung Ba
26 Mar. 1948, Johnson 348/28 (NSW 5025); Blue Mountains, March 1910,
Domin 7757 (pr); near King’s Fall, Oct. 1819, A. Cunningham 94/1819 (x);
Mt. Jellore, 16 Nov. 1912, Cheel (Nsw 44071); INawara, Oct. 1818, A. Cunning-
ham (Kk), 1855 Macarthur (pm) & Oldfield (K); near Robertson, Nov. 1918,
Dunn 27 (nsw 44073); Jamberoo-Carrington Falls road, 22 Jan. 1955, Con-
stable (BM, G, K, NSW 32245); Nowra-Kangaroo Valley, near top of pass on
Nowra side, about 350 m. alt., 14 Jan. 1936, Rodway 2113 (6242) (A, BRI 7929,
K, Nsw 33417); Milton, 8 Nov. 1914, Cambage 4121 (Nsw 44078); Sugar Loaf
Mt., Monga, near Braidwood, Jan. 1915, Boorman (Nsw 44079); Eurobodalla,
1 Jan. 1950, Whaite 480 (Nsw 44080); Ohlson’s Creek, near Narooma, against
the road to Tilba Tilba, 23 May 1964, Willis (x, MEL 19629); Quaama, Dec.
1915, Dunn (Nsw 44082); near Tanja (Bega distr.), 1948, Phillips (Nsw 44083) ;
Eden to Pambula, Nov. 1901, Maiden (Nsw 44085); Twofold Bay, Sept. 1860,
F. Mueller (x, holotype of N. longifolia var. oo Womboyn Lake, 135
m. me 12 Oct. 1954, Constable (a, K, NSW 36595).

ctoria. Genoa River, 1880, Reader (MEL 19607); Mallacoota, Hart (MEL
19609 & 19610); Mt. Drummer, 11 June 1959, L. J. W. & J. G. T. 3619 (Bri
34850); sources of Delegate River, 1887, Merrah (MeL 19611); arate
River, Jan. 1855, F. Mueller (mE 19620, holotype; K, MEL 19632, isotypes) ;
Orbost, 20 Oct. 1903, Grove (Gc, K, LE); North Arm, Lakes Entrance, 17 Aug.
1944, Willis (MEL 19621)

This species is closely related to Notelaea longifolia and N. ovata and
has often been misidentified as the former, although its main distribution
is more southerly (Map 3). However, it may readily be distinguished by
the fine and even reticulation of the leaves, especially below. This species
may also be recognized by the fruit which is relatively large and slightly
longer in proportion to its width. When ripe the fruit is quite fleshy and
bluish-black with a slight glaucous bloom, although the label with MEL
19626 from E. Gippsland says “white and red.” According to annotations
by L. A. S. Johnson it is characteristic of sandstone soils in or around rain-
forest, though often of the lower, poorer type, as in mountain gullies, etc.,
but sometimes in more open habitats or in patches of undergrowth of
rainforest species in wet sclerophyllous forest.

Var. A.

Leaves subsessile or with very short petioles, 1.5—3 mm. long; leaf base
subcordate or broadly rounded.

Queensland. Moreton District: top of Mt. Mitchell, about 1050 m. alt., 19
July 1930, White 6886 (Bri 9692); Wilson’s Peak, Peterson (BRI 9706); top of
Mt. Lindsay, about 1200 m. alt., 10 Aug. 1932, Whitehouse (BRI 9693).

New South Wales. Mount Ma ckenzie, Tenterfield, 4 Nov. 1963, Goddard
(BRI 42938); near Hyland State Forest, 1375 m. alt., 5 Apr. 1955, Turner
(Nsw 44058); Orara West State Forest, E. Dorrigo, May 1946, W. T. Jones 5
(BRI 42305-6); Bellangry State Forest, about 20 miles NW. of Wauchope, 2
Jan. 1957, Cousins 180 (Nsw 44059); Comboyne, Jan. 1934, Chisholme (NSW

1968 |

GREEN, REVISION OF NOTELAEA

351

NOTELAEA MICROCARPA
© VAR. MICROCAR PA
@ VAR. VELUTINA

we NOTELAEA PUNCTATA
a NOTELAEA VENOSA
a

VENOSA VAR A

oe cet me

2° o90 #
Pat f

8
3p fete)

90
° 09 Boo

Map 3. Distribution of Notelaea venosa, N. punctata and N. mcr OCT Pe.

352 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

44060) & Fraser (Nsw 44061); The Corker, Upper Williams River to Barrington
Tops, 26 May 1934, Fraser 19 (Nsw 44062).

When studying the material of this species, certain specimens with
subcordate leaf bases and very short petioles (1.5-3 mm. long) came to be
put on one side as constituting a distinct entity. This seemed more prob-
able when it was noted that one or two came from relatively high altitudes
and that when their distribution was plotted on a map they were found
only amongst the northernmost specimens (see Map 3). However, with
the limited material available there seem to be no other distinguishing
characters and one of the southernmost specimens, Miss L. Fraser from
Comboyne (Nsw 44061), exhibits both types of leaf, in this case with the
unusual short-stalked leaves together with inflorescences on the youngest
shoots, and the typical leaves towards the base. Also, a specimen at Kew
collected by Charles Fraser during the last century from about the same
area, “native of the banks of the River Hastings,” bears leaves somewhat
intermediate in leaf base and petiole length (6-8 mm.). A comment, writ-
ten upon another sheet, suggests that the unusual leaves are juvenile, but
several of the specimens exhibiting them bear young fruit or the remains
of flowers, while other undoubtedly juvenile collections bear leaves with
typical bases and petioles, while a seedling collected on Mt. Drummer
(BRI 34850), although with short petioles, has the typical acute leaf base.
Further collections and field observations from the populations in north-
west New South Wales and southernmost Queensland are desirable and
may prove that they do merit nomenclatural differentiation after all.

4. Notelaea punctata R. Brown, Prodr. 524. 1810; Sprengel, Syst. Veg.
1: 35.1824; A. DC. in DC. Prodr. 8: 291. 1844; Reichenb. f. Neuholl.
Pfl. Amelia Dietrich 9. 1866; Bentham, Fl. Austral. 4: 300. 1868; F.
Mueller, Syst. Census Austral, Pl. 92. 1882; F. M. Bailey, Synop.
Queensl. Fl. 303. 1883; F. Mueller, Second Syst. Census Austral. PI.
156. 1889; F. M. Bailey, Cat. Pl. Queensl. 29. 1890; C. Moore, Handb.
Fl. New S. Wales, 320. 1893; F. M. Bailey, Queensl. Fl. 3: 973. 1900;
Dixon, Pl. New S. Wales 214. 1906; F. M. Bailey, Comprehensive
Cat. Queensl. Pl. 311. 1913; Domin, Bibliot. Bot. 22(89) (Beitr. Fl.
Pflanzengeogr. Austral. 1(3): 514): 1068. 1929: Bean in Chittenden,
Dict. Gard. 3: 1379. 1951.

Bushy shrub to 1(-2) m. high (? more), branches glabrous or very
minutely puberulous when young. Leaves glabrous; petiole 2—6(—10) mm.
long, glabrous; lamina thickish, punctate above and below, especially
below, narrowly ovate to lanceolate or elliptic to narrowly elliptic, (3.5—)
5-9(-13) cm. long by (1—)1.5-3(—5) cm. broad; margin entire or occa-
sionally slightly, very shallowly and irregularly crenulate, slightly thick-
ened; apex acute or very slightly acuminate, tip acute; base acute or
cuneate (rarely obtuse and rounded), attenuate or somewhat decurrent
onto the petiole; venation raised and clearly reticulate above, usually

1968 | GREEN, REVISION OF NOTELAEA 553

less clearly so below, often obscure or visible as lighter lines or only
slightly raised, 6-8(—10) primary veins per side. Inflorescence axillary,
often up to 3 mm. above the axil with 1-2 dormant buds in between,
1(—2) per axil, decussate, (1—)1.5-3 cm. long, (7—)9-13-flowered, gla-
brous, except occasionally for a few minute scattered hairs towards the base;
upper bracts 1—1.5 mm. long, thin, lanceolate to ovate, acute, early decidu-
ous, ciliolate, basal pair thick, 1-2 mm. long, acute or acuminate, glabrous
or with a few minute hairs towards the apex, persistent or at least the
fused bases persistent. Flowers hermaphrodite, creamy white or pale
yellow; pedicels 2—5(—8) mm. long. Calyx 0.5—1 mm. long, lobes 4, broad-
ly and irregularly triangular, 0.3-0.6 mm. long, glabrous or ciliolate, some-
times subapiculate. Corolla induplicate-valvate, lobes 4, 2-3 mm. long,
concave, joined in pairs above the base of the filaments for about 0.4 mm.
Stamens 2, broadly ellipsoid, 1.5—2.5 mm. long, more or less enclosed within
the concave petals, filaments 0.3-0.5 mm. long, with a terminal appendage
about 0.1-0.3 mm. long, dehiscence lateral. Ovary 1-1.5 mm. long with 2
stigmatic lobes 0.2-0.3 mm. long on a short style about 0.2 mm. long.
Drupe varicolored (fide McLaughlin 22C), pale (fide note with Biddulph
63), ovoid, 7-9 mm. long by 5-6 mm. broad; endocarp hard, 0.2 mm.
thick.

Queensland. Coox District: Middle Creek, about 6 miles SE. of Cairns,
30 Oct. 1949, Flecker in N. Queensland Nat. Club 13331 (prt 42304). NortH
KENNEDY District: SE. slopes of Gt. Dividing Range, about 4 miles NNE. of
Herberton, about 1060 m. alt., 7 Aug. 1948, L. S. Smith 3729 (BRI 42313) &
about 6 miles NNE. of Herberton, about 1100 m. alt., 26 Aug. 1954, Smith
5301 (BRI 42302 & 42303); Herberton, 1918, Michael 624 (BRI 7911); W. of
Herberton, crest of Gt. Dividing Range, 1050 m. alt., 22 Aug. 1963, Blake 22107
(srt 42318); Rockingham Bay, Dallachy (BM, K, LE, NSW 44056). Port CuRTIS
District: Shoalwater Bay, 1802, R. Brown “2846” (BM, holotype; E, K. MEL
19631 isotypes). LeIcHHARDT District: Blackdown, about 12 miles SSE. of
Bluff, Aug. 1961, Gittins 389 (prt 30808); Aug. 1964, Gittins 887 (BRI 56856)
& 887/2 (BRI 56855); Blackdown Tableland, south of Dingo, Sept. 1937, Sim-
mons (BRI 7917); Rockland Spring, 30 miles ENE. of Springsure Township, 5
Sept. 1961, Lazarides & Story 121 (CANB 111988, K, NSW 67495); Expedition

ange, Aug. 1960, Gittins 360 (BRI 25421); Mt. Playfair, Springsure district,

K); Isla Gorge between Theodore and Taroom, 1 Sept. 1963, Hockings (BRI
42317); 16 miles SSW. of Cracow Township, 10 July 1963, Lazarides 6950
(BRI 51428—-9, CANB 123629). BURNETT DISTRICT: Eidsvold, 1919, Bancroft (BRI
9700). Wwe Bay District: Fraser Island, 6 Sept. 1916 & Dec. 1918, Ses
32 (BRI 9698 & 7927) & Oct. 1921, White (Bri 7936); Cootharaba, Dec. 1917,
Francis (A, BRI 7932); Imbil, 27 March 1918, Weatherhead (prt 9697, K).

Although in its typical state, with an obscure venation on the undersur-
face of the leaf and small fruits, Notelaea punctata 1s easily distinguished
from typical N. longifolia (in which the leaves are clearly reticulate
below with raised veins and the fruits are larger), the two species tend
to merge where their areas of distribution meet (Maps 1 & 3). The

354 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

dotting of the leaf, so abundant and obvious on the type, and thus taken
by Robert Brown as the basis for his specific epithet, is also found in
N. longifolia (and other species) but usually much less obviously and
abundantly so. Using leaf characters alone, it appears that there may
be an area of transition, but too few specimens are in fruit and none from
the intermediate areas. More collecting and further field observations are
badly needed, especially in the Burnett and Wide Bay Districts of Queens-
land. This species is insufficiently well known and it may turn out that
it would be better treated as a subspecies of NV. longifolia.

C. Moore in his Handbook of the Flora of New South Wales of 1893
(p. 320), records Notelaea punctata from the northern coast district of
New South Wales, but this is clearly an error and must have been based
on a misidentification

In habit Notelaea punctata would seem, from the limited field notes
available, to resemble V. ovata, usually growing as a low shrub, with many
erect branches, from a rootstock which presumably survives bushfires. The
only notes giving details of habitat indicate that N. punctata is found
under Eucalyptus spp. on sandy soils on sandstone plateaux and ranges.

unt

. Notelaea microcarpa R. Brown, Prodr. 524. 1810; Sprengel, Syst.
Veg. 1: 35. 1824; A. DC. in DC. Prodr. 8: 291. 1844; Bentham, FI.
Austral. 4: 300. 1868; F. Mueller, Fragm. Phytogr. Austral. 8: 43.
1873, & Syst. Census Austral. Pl. 92. 1882; Bailey, Synop. Queensl.
Fl. 303. 1883, & Queens]. Woods 89. 1888 & 97. 1889; F. Mueller,
Second Census Austral. Pl. 156. 1889; Maiden, Useful Native PI.
Austral. 580. 1889; C. Moore, Handb. Fl. New S. Wales 320. 1893;
F. M. Bailey, Cat. Pl. Queensl. 29. 1890, & Queensl. Fl. 3: 973. 1900;
Dixon, Pl. New S. Wales 214. 1906; Guilfoyle, Austral. Pl. 100.
[1911]; F. M. Bailey, Comprehensive Cat. Queensl. Pl. 311. 1913;
Maiden & Betche, Census New S. Wales Pl. 172. 1916; Domin, Bibliot.
Bot 22(89) (Beitr. Fl. Pflanzengeogr. Austral. 1(3): 514): 1068.
1929; Anderson, Trees New S. Wales, ed. 2. 69, 132, 338. 1947; M.
Gray, Contr. New S. Wales Natl. Herb. 3: 59. 1961.

Evergreen shrub or small tree to 10 m., usually round headed; bark
gray, tessellated; branches puberulous when young, sometimes only mi-
nutely so. Leaves glabrous or scattered puberulous above and below, be-
coming glabrous, except occasionally near the midrib, or densely velutinous,
especially below; petiole (1—)2—5(—10) mm. long, glabrous or puberulous,
becoming glabrous; lamina thickish, scattered punctate, (lanceolate—)
narrowly or very narrowly lanceolate (—linear), (2—)3-8(—15) cm. long by
(0.3)0.5-1.5(—3) cm. broad; margin entire, scarcely thickened or occa-
sionally somewhat thickened, usually flat; apex acute or obtuse-apiculate;
base very narrowly cuneate, decurrent onto the petiole; venation usually
raised and reticulate, especially above, less so, and sometimes almost ob-
scure, below, 5—7 primary veins per side making an angle of about 15°—30°
with the midrib and eventually running almost parallel to the margin.

1968 | GREEN, REVISION OF NOTELAEA 355

Inflorescence axillary, 1-2 per axil, decussate (rarely and freakishly, ter-
minal and then perhaps thyrsoid), 0.5-2 cm. long, (7—)9-13(—15)-flowered
or rarely (more or less freakishly?) paniculate and many flowered
(Nsw 33512 & 33606), puberulous, especially towards the base, to remotely
scattered puberulous, rarely almost glabrous with a few hairs towards the
base; upper bracts 1.5—2 mm. long, thin, more or less ovate or lanceolate,
early deciduous, ciliolate, basal pair thick, their joined bases persistent,
puberulous, rarely glabrous (in northernmost specimens), 1—-1.5 mm. long,
acute. Flowers hermaphrodite, greenish-yellow, yellow or cream; pedicels
0.5-4(-6) mm. long. Calyx 0.3-0.6 mm. long, lobes 4, usually with 2 long
and 2 short (the longer at the base of the corolla pair), irregularly triangu-
lar, 0.1-0.5 mm. long, glabrous or margin bearing a few hairs. Corolla
induplicate-valvate, lobes 4, 1.4-2 mm. long, concave, joined in pairs
above base of filament for about 0.7-0.8 mm. Stamens 2, ellipsoid, (0.8—)
1—1.5 mm. long, more or less enclosed within the concave petals, subsessile
or on filaments 0.1-0.2 mm. long, with blunt terminal appendage about
0.1-0.2 mm. long, lateral dehiscence. Ovary 1-1.5 mm. long, with 2
stigmatic lobes 0.2—0.3 mm. long on short style about 0.2 mm. long. Drupe
dark blue or blackish (becoming soft and purple when ripe, fide Johnson),
ovoid, (S—)7-10 mm. long by (4-)5-8 mm. broad (smallest dimensions
immature?) ; endocarp hard, 0.2-0.4 mm. thick.

Var. microcarpa
Leaves glabrous or with only scattered pubescence above and below.

SELECTED ExsIccATAE: Queensland. Coox District: Stannery Hills, Apr.
1909, Bancroft (Bri 8272, K) & Dec. 1908, Bancroft 259 (prt 8250, K); Lynd
Scrub, approx. 40 miles SW. to SSW. of Mt. Garnet, 24 Aug. 1949, L. S. Smith
3859 & 3869 (BRI 42307-8). NortH KENNEDY District: St. Roman’s Station,
Mt. Garnet, 29 Apr. 1960, Myers (Bri 28627); Dickson Hill, Mt. Fox, 27 Sept.
1949, Clemens (kK); Dalrymple Road, 12 miles N. of Charters Towers, Flecker
(Rodway 7933) (NSW 33422). SouTH KENNEDY DistTRIcT: 3 miles W.
“Collin Downs” Station, 11 Aug. 1964, Adams 1261 (cANB 143194, K). Port

°
a

); Orallo, Oct. 1920, Dunlop (BRI
, NSW 33574) & 25 Oct.
16

Blake 10935 (prt 8257 & 8261, CANB 38903, : nh
8274, k, Nsw 33572); Roma, July 1915, Bick (BRI cat,
1933, White 9510 (A, BRt 8255); Honeymah, about 40 miles SE. of Bollon,

356 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

July 1948, Everist 3478 (prt 8273, CANB 25566, K). BuRNETT DIstRIcT: Eids-
vold, Bancroft (prt 8278, K); Mt. Perry, Keys 269 (srt 9696) ; Dallarnil, 28 Dec.
1939, L. S. Smith 665 (A, Brkt 9699); Haley Creek Road, 21 Oct. 1947, Mitchell
3050 (prt 7915). Wipe Bay District: Theebine, Nov. 1921, White (A, BRI 8265,
K); Wide Bay, 1854, Moore (K). Moreton District: Forestry Reservation,
Yarraman, approx. 400 m. alt., Aug. 1944, Clemens (a, BRI 42312); Laidly Hills,
25 Jan. 1944, Clemens 434760 (a); Pine Mountain, N. of Ipswich, Oct. 1946,
Everist (prt 9575, K); 1 mile N. of Boonah, on Kalbar Road, 24 Nov. 1946,
Everist & Webb 1415 (prt 9715, CANB 16733, K); mouth of Coomera R., 21
Jan. 1927, White 3381 (A, BRI 7912, K). Dartinc Downs District: Glen-
morgan, 19 July 1949, Gordon 63 (par 42316); 2 miles W. of Condamine, 24
Aug. 1956, Everist 5795 (Bri, CANB 100241, Kk); N. of Miles, 14 Aug. 1951,
Webb 1500 (cans 30648); Chinchilla, 8 July 1912, Bensley 1 (BRI 8258, K);
Warwick, Aug. 1913, Shirley (Bri 8254, K, NSW 33575); Silverwood, Sept. 1922,
White 1763 (a, BRt 18276, K, NSW 33576).

New South Wales. 20 miles N. of Torrington, 12 Dec. 1951, Jessup & Gray
23 (CANB 126269, NSw 42868); Mt. Mitchell, near Warialda, 28 July 1922, de
Beuzeville 12 (Nsw 33580); Wallangra, Sept. 1929, Rodway 6225 (kK, NSW
33421); Moree, 29 Sept. 1912, Brennan (Nsw 33582); Narrabri West, Aug.
1907, Boorman (cH, NSw 33590, z); Warrumbungle Mountains, 640 m. alt.,
26 May 1948, Constable (pM, K, NSW 6475); Little Sugar Loaf Mountain, near
Gunnedah, 352 m. alt., 1 Nov. 1954, Johnson & Constable (K, NSw 32028);
“Lowestoft,” 27 miles S. of Tamworth, near highway, 684 m. alt., 18 Apr. 1956,
ening 1658 (A, K, NSW 37333); Wellington, 20 Oct. 1880, Betche (NSW

3592); “Merrot,” Curriecabark, 30 miles NW. of Barrington, Feb. 1949, Hyem
ae 33594); 6 miles SW. of Scone, 16 March 1960, Story 7071 (cANB 80160,
kK); Mt. Wambo, 5 miles NW. of Bulga, 600 m. alt., 30 Aug. 1957, Constable

)-
Cultivated. Botanic Gardens, Brisbane, 31 Oct. 1930, Hubbard 4740 (a, K)
& 8 Nov. 1963, Green 1530 (a).

Notelaea microcarpa is one of the two most widespread species in this
genus (Map 3). To judge from its distribution, and from the annotations
on the herbarium covers at Sydney, it is characteristic of an area of mod-
erately dry, distinctly continental climate with a summer maximum
rainfall. It is an “inland” species of the “western slopes country” and
although it approaches the coast in parts of Queensland, e.g. around Rock-
hampton, it does so in comparatively dry areas. At its southern limit it
stops ‘‘at the Wellington-Hunter Valley latitude, i.e. mene the summer
rainfall zone begins to pass into the transitional zon

At its southern limit the distribution of Notelaea i ae approaches
most closely to NV. neglecta, perhaps the most nearly related species, and
the one under which their differences and affinities are discussed. J.
neglecta is found to the south and east with a damper and often cooler
climate.

Notelaea microcarpa, as its name suggests, is characterized by small
fruits, a character it shares with NV. neglecta and N. linearis, both of which
have narrow leaves. From N. longifolia it may be distinguished by the
relative obscurity of the veins on the underside of the leaf (reticulate in
N. longifolia) and also by the narrow angle at which the primary veins

1968 | GREEN, REVISION OF NOTELAEA 357

join the midrib, approximately 15°-30° in WN, microcarpa and 40°-70° in
N. longifolia.

However, exceptionally narrow leaves may occur rarely on adult plants,
for example, a specimen from Sarina (Port Curtis District) Queensland
(CANB 136113) exhibits linear leaves, (3—)4.5—7 (—9) cm. long by (0.25-)
0.35—0.6(—0.8) cm. broad; the notes on the label say “very narrow leaf
form, extreme dry conditions.” The venation is obscure as well and it
might easily be mistaken for NV. neglecta were it not from far outside the
geographical range of that species.

Using herbarium material alone it is very difficult to evaluate juvenile
characters but it is suspected that four herbarium sheets bearing specimens
with much larger leaves than usual were gathered from juvenile plants.
Three of these are from the Yarraman F orestry Reservation, Moreton Dis-
trict, Queensland, Aug. 1944, coll. M. S. Clemens (a). Two sheets have
notably juvenile, long narrow leaves 10-27 cm. long and 0.3-1.2 cm. broad
exhibiting up to about 20 primary veins on each side of the midrib. The
third sheet, annotated as a slender sapling, is in flower and has broader
leaves than usual, up to 11 cm. long and 3.5 cm. broad. The fourth, from
St. Roman’s Station, Mt. Garnet, N. Kennedy District of Queensland,
has leaves 20 cm. long and 2 cm. wide. Large juvenile foliage is similarly
found in Notelaea longifolia and N. ovata and particularly in the more
distantly related New Zealand species of Nestegis. a9

Notelaea microcarpa is believed to hybridize with both N. longifolia
and J. ovata and more detailed mention of this has been made under these
species.

Var. velutina (F. M. Bailey) P. S. Green, comb. nov.

N. longifolia Vent. var. velutina F. M. Bailey, Comprehensive Cat. Queensl.
Pl. 311, 319 & 839. fig. 293. 1913; White, Queensl. Agric. Jour. II. 22: as
bl. 77, 78. 1924 & op. cit. 23: 435, 436. pls. 80, 81. 1925; Domin, Bibliot.
Bot, 22(89) (Beitr. Fl. Pflanzengeogr. Austral. 1(3): 514): 1068. 1929.

Leaves densely velutinous, especially on the undersurface.

Queensland. Dartinc Downs District: Silverwood, Sept. 1922, geomesy
(A, BRI 7941, K, Nsw 33530); Killarney, Bailey (? Wedd) Ses baie a. Re
K, isotype); Wilson’s Peak, Michael 1983 p.p. (NSW 33532); Stant ae 4
1904, Boorman (Nsw 33531), about 1000 m. alt., June 1924, hace B me pre
K) & Perkins (pri 7943, K); Eabey, via Stanthorpe, Nov. has (x)
7923); Granite National Park, Wyberba, Nov. 1944, Clemens . a sass);

New South Wales. Macphemos anes == pater sir ae Gorge,

2852 (NSW ; tas
Rivertree, 5 Sept. 1911, Cambage River, near Dorrigo, 5 Nov. 1963,
28 Sept. 1912, Stopford
45 (pm); Castle Doyle Road, about 7 miles SE. of Armidale, Feb. 1928, McKie
93 (NSW 33536); Wollomombi, 4 March 1933, \. 7
Falls, about 900 m. alt., 25 May 1957, Johnson (Nsw lame peal
Walcha), Dec. 1898, Betche (Nsw 33538) & Oct. 1900, Forsy

33539).

358 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Although this variety was attributed by F. M. Bailey to Notelaea longi-
folia, it really belongs to NV. microcarpa. Most specimens are in flower
or immature fruit but the collection from Stanthorpe, Queensland, by F. A.
Perkins in June 1924 has mature fruits of the typical small size, 8-9 mm.
long by 7-8 mm. broad. Furthermore, the angle the primary veins make
with the midrib of the leaf, the relative obscurity of the veins beneath
(although this is masked by the velutinous covering), the general propor-
tions of the leaf and the length of the individual inflorescences, all indicate
N. microcarpa. Velutinous forms of N. longifolia exist but are found in
the southern parts of range for that species, so that all V. /ongifolia in the
northern areas of New South Wales or in Queensland are glabrous, or, at
the most, glabrate. Cambage 2702 from the central tablelands region of
N.S.W. (from Baerami to Denman, 15 May 1911, Nsw 33540) was ex-
amined carefully because of its narrow and velutinous foliage and it is
believed that it represents the result of hybridization between this species
and JN. longifolia {. longifolia.

The distribution of this variety is restricted to the southeastern part
of the Darling Downs district of Queensland and the Northern Tablelands
of New South Wales. I have seen material from the northernmost areas of
New South Wales and again in the southern parts of the New England
Ranges but it may well occur in between.

Typical Notelaea microcarpa occurs near at hand in parts of the variety’s
range, e.g. White 1763 from Silverwood, Queensland, which was pre-
sumably collected near to White 1764 a specimen of the densely velu-
tinous variety, while collections with slight pubescence are found scattered
throughout the range of the species. In November 1963 I was fortunate
to be taken to a population of Notelaea near Dorrigo through the kind-
ness of Alex G. Floyd and Harold C. Hayes, and there took samples from
12 bushes and small trees scattered over a small area near the Nymboida
River (P. S. Green 1519 a). At the time I was struck by the variation
in the breadth and pubescence of the leaves, but it is now apparent to
me that this variation is due to introgression which has occurred between
N. longifolia and N. microcarpa var. velutina. The area, like much of that
part of New South Wales, is one where considerable felling has taken
place and the velutinous upland plant may well have come into contact
with the more lowland N. longifolia due to man’s activities. In the citation
of the population sample I have placed those parts which seem nearer
to N. microcarpa var. velutina above and those nearer to N. longifolia
under that species. Turner 29, cited under N. longifolia, from Paddys
Land State Forest, a little distance to the northwest, also shows inter-
mediate characteristics and some of the gatherings cited above as var.
velutina possess leaves which are slightly wider in proportion to their
length than is usual for the species, and may perhaps show the influence
of N. longifolia, for example the Clemens collection from the head of
Wylie Gorge, Liston, New South Wales.

6. Notelaea linearis Bentham, Fl. Austral. 4: 300. 1868; F. Mueller,

1968 | GREEN, REVISION OF NOTELAEA 359

Syst. Census Austral. Pl. 92, 1882; F. M. Bailey, Synop. Queensl. FI.
303. 1883 & Cat. Queens]. Fl. 29. 1890; C. Moore, Handb. Fl. New
S. Wales, 321. 1893; F. M. Bailey, Queensl. Fl. 3: 973. 1900; Dixon,
Pl. New S. Wales 215. 1906; F. M. Bailey, Comprehensive Cat.
Queensl. Pl. 311. 1913; Maiden & Betche, Census of New S. Wales PI.
172, 1916; Anderson, Trees New S. Wales ed. 2. 132. 1947; M. Gray,
Contr. New S. Wales Natl. Herb. 3: 59. 1961.

Evergreen shrub 1-2 m. high, branches glabrous or minutely puberulous
at first, soon glabrous. Leaves glabrous; petiole 1-3(-4) mm. long,
glabrous or minutely puberulous above when young; lamina thick, linear
or occasionally very narrowly lanceolate or elliptic (? juvenile condition),
(2—)3-6(-11.5) cm. long by (0.15—)0.2—0.5(-0.7) cm. broad (? up to 1
cm. broad in juvenile state), punctate, especially below, margin entire,
somewhat thickened; apex very acute; base narrowly attenuate into the
petiole; venation completely obscure above and below, or rarely (?
juvenile state) with primary veins just visible above and below, about 7
per side joining to form a submarginal nerve. /nflorescence axillary, 1 per
axil, decussate or subumbellate, 0.4-1 cm. long, (3—)5(-7)-flowered,
glabrous or puberulous, often only very slightly so; upper bracts 0.5—1
mm. long, thin, triangular-ovate, early deciduous, basal pair thick, per-
sistent, glabrous or puberulous-ciliolate, 1 mm. long, acute. Flowers her-
maphrodite (Fic. 2), pale yellow (fide White & Williams); pedicels 1-4
mm. long. Calyx 0.5-0.7(—1) mm. long, lobes 4, irregularly triangular,
0.2-0.5 mm. long, glabrous, margins sometimes slightly ciliolate. Corolla
induplicate-valvate, lobes 4, (1.5-)2 mm. long, concave, in pairs, joined
above the base of filaments for 0.3-0.6 mm. Stamens 2, (1—)1.5 mm. long,
more or less enclosed within the concave petals, on filaments 0.2-0.3 mm.
long with a broad connective and well developed blunt terminal appendage
0.2-0.3 mm. long. Ovary 1-1.4 mm. long with 2 stigmatic lobes 0.2-0.4
mm. long on style 0.2 mm. long. Drupe ovoid, 5—7 mm. long by 4-5 mm.

men. (Drawn from Hubbard

0
Fic. 2. Notelaea linearis. A, flower; B & C, sta
3961).

360 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

broad, “white and translucent” (fide Galbraith, Bri 49151) and “rich blue
with a thin bloom” (fide Galbraith, Bri 49152); endocarp hard, 0.2 mm.
thick.

Queensland. Moreton District: Crow’s Nest, Sept. 1920,Kenny (BRI 7904),
Oct. 1921, White (a, Nsw 33423); Pearson’s Falls, Helidon, Bailey (prt 7905,
K). Dartinc Downs District: Thulimbah, Schindler 3 (srt 7901); Messines,
near Stanthorpe, approx. 900 m. alt., 13 Sept. 1930, Hubbard 3961 (Bri 7906,
K); Stanthorpe, Bailey (srt 7903) & July 1904, Boorman (Nsw 33424); Glen
Aplin, 9 Sept. 1933, White 9253 (A, BRI 7902); Wyberba, Feb. 1964, Galbraith
(pri 49151-2); Wallangarra, Nov. 1904 & Jan. 1906, Boorman (Nsw 33427 &
33428).

South Wales. Wallangarra area, 16 March 1954, Jessup & Gray 2554
(CANB 126268); Richmond River, Beckler (z); Wilson’s Downfall, 3 Sept. 1911,
Cambage 2809 (Nsw 33425); Boonoo Boonoo, Nov. 1904, Boorman (NSW
33426); Jennings, Oct. 1901, Boorman (Nsw 33429); near Tenterfield, Stuart
(k, syntype); Torrington, Oct. 1911, Boorman (Nsw 33431); Bismuth, via
Deepwater, Aug. 1913, McNutt (Nsw 33430); Gibraltar Range, NE. of Glen
Innes, Apr. 1958, Williams (Nsw 44089); 27 miles E. of Glen Innes, 1000 m.
alt., 4 Oct. 1958, Williams 625 (Nsw 46450); Mount Mitchell, Beckler (x,
lectotype); top of range near Backwater, 30 Oct. 1929, Blakely, McKie oe
Youman (nsw 33432). Without locality: New England, F. Mueller (BM).

Apart from Notelaea neglecta, described below, N. linearis can be dis-
tinguished at a glance from all other species in the genus by its extremely
narrow leaves. In juvenile plants it is suspected that they are a little
broader perhaps, but one cannot be certain without adequate observations
based on living plants. In N. neglecta the leaves are a little broader but
the two species differ from one another not only by their leaf venation,
which is obscure in NV. linearis, but in the inflorescence length, which is
longer in N. neglecta (in fact the inflorescence is shorter than in any other
species except possibly some representatives of N. microcarpa). N
linearis is found on siliceous soils in eucalypt forest and is ecologically
distinct from N. neglecta. N. linearis is confined to an area in north-
ern New South Wales and southernmost Queensland, with N. neglecta
in an even more limited area about 250 miles further south in the Blue
Mountains in central New South Wales (Map 2).

From the type material in the Kew Herbarium I have selected the speci-
men collected by Beckler and sent to Kew by von Mueller in 1868 as the
lectotype because it exhibits both flowers and fruit, whereas the other
collection cited in the protologue, the Stuart specimen from near Tenter-
field, whilst mounted on the same sheet, bears only one or two young
flower buds.

7. Notelaea neglecta P. S. Green, sp. nov.

Species foliis angustis (2.5—)4—7(-11) cm. longis, (0.2—)0.3-0.8(-1.4)
cm. latis, venatione obscura sed manifesta, inflorescentiis 1-2 cm. longis,
drupis parvis 5-6 mm. 4—5 mm. distincta. A N. lineari foliis plerumque
longioribus et latioribus, inflorescentiis longioribus (10-20 mm. longis)

1968 | GREEN, REVISION OF NOTELAEA 361

differt. A N. microcarpa foliis angustis, venatione minore prominente,
venis primariis sub angulo circiter 40° e nervo mediano exortis, inflores-
centlis glabris differt.

Evergreen, probably a shrub; stem minutely puberulous when young,
sometimes becoming glabrous. Leaves thickish, glabrous, or minutely
puberulous above and below at first when young, finely punctate; petiole
1-5(—8) mm. long, minutely puberulent, especially above, often becoming
glabrous; lamina very narrowly elliptic or narrowly lanceolate to linear,
(2.5-)4-7(-11) cm. long by (0.2—)0.3-0.8 (-1.4) cm. broad; margin entire,
somewhat thickened; apex elongated, very acute; base acute, long attenu-
ate into the petiole; venation more or less obscure except for the slightly
visible primary veins, especially above, often not visible below, (6—) 7-10
(—11) per side, making an angle of about 40° with the midrib. Jnflorescence
axillary, 1-2 per axil, decussate, glabrous; 1-2 cm. long, (3-)7—9-flowered;
upper bracts early deciduous, broadly ovate to lanceolate, pointed, 1-2 mm.
long, somewhat puberulent, especially towards the tip; basal bracts thick-
ish, 1-2 mm. long, acute or acuminate, puberulent, fused bases persistent.
Flowers hermaphrodite, yellow (?); pedicels (1-)3-5 mm. long. Calyx
0.5-1 mm. long with 4 irregularly triangular, more or less erose lobes,
0.3-0.6 mm. long. Corolla induplicate-valvate, 1.5—2.25 mm. long, 4 lobes
joined in pairs at the filament 0.3-0.5 mm. from their base, concave. Sta-
mens 2, 1.25-1.5 mm. long, included within the concave corolla lobes;
filament short and broad, 0.2 mm. long. Ovary flask-shaped, 1-1.5 mm.
long, tapering into an imperceptible style with a bifid stigma 0.2 mm. long.
Drupe ovoid, globular, 5—6 mm. long by 4-5 mm. broad (Nsw 33635).

New South Wales. Capertee, Sept. 1915, Boorman (NSW 33618); Jenolan
Caves, Sept. 1899, Blakely (Nsw 33620, holotype; A, K, & Nsw 33621, isotypes),
5 Sept. 1923, Cheel (Nsw 33623), 29 Oct. 1940, Blakely & Ludowici (A, K, NSW
33619); Colong-Yerranderie, 5 Oct. 1909, Cambage 2268 (Nsw 33624); Wom-
beyan Caves, Oct. 1905, Maiden (a, BRI 8263, K, NSW 33629-31), 1 Oct. 1905,
Cambage 1345 (Nsw 33632); Bullio to Wombeyan, Oct. 1905, Maiden (Nsw
33628); Bowral to Wombeyan, 30 Sept. 1905, Cambage 1330 (CANB 7610, nt
33627); Mittagong to Bullio, 28 Nov. 1919, Cheel (Nsw 33625); Berrima, July
1906, Boorman (Nsw 33626); Devil’s Hole, Tallong, Oct. 1917, Rumsey Soi
33636); Wingello-Tallong, Oct. 1938, Murphy (Nsw 33633); Tallong (Barber's
Creek), Dec. 1897, Maiden (Nsw 33635), Oct. 1899, Rumsey (nsw 33634).

The epithet chosen for this species is intended to reflect the fact that
it has been overlooked for many years. L. A. S. Johnson had already
recognized it as distinct and segregated the material in the ona ap
barium, before it was sent on loan, and I am greatly indebted to him for
a : - .

Todas fies the names already on the herbarium sheets, this species
has previously been confused with Notelaea linearis and N ch
presumably because of the general characters of the leaf, whic “6 roug ms
between the two. The typical leaf is neither as narrow as that o : A
linearis nor as broad as that of N. microcarpa. The venation is somewhat

362 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

intermediate as well, more distinct than that in the former and more
obscure than the latter. All three are small fruited and undoubtedly closely
allied but, although it is possible that they arose from a common stock,
or that at some stage NV. neglecta developed as a stabilized hybrid between
the other two, they are distinct today.

Geographically each has its own area of distribution and, to judge from
field notes, their edaphic and climatic adaptations are different. Notelaea
neglecta grows on limestone, and possibly a variety of rocks, but never
on the most siliceous. NV. /inearis, on the other hand, is a species of siliceous
soils, including “acid granite.” N. microcarpa occurs on a variety of rocks
and, like NV. neglecta, never on the most highly siliceous, but climatically
it is restricted to the ‘western slopes country” although, at its southern
limit, the two species approach one another (Maps 2 & 3).

Notelaea sect. Ligustrina P. S. Green, sect. nov.

A sectione Notelaea corollae aestivatione imbricata differt. Flores parvi,
hermaphroditi; corollae lobi 4, leviter imbricati, per paria connati vel ope
baseos filamenti conjuncti; corollae tubus perbrevis vel nullus. Drupa
parva, vel nigra vel purpurea vel rubra vel rosea vel alba.

Type species: Notelaea ligustrina Vent.

The separation of this section and the relationships of its single and
type species, Notelaea ligustrina, are discussed below under that species.

8. Notelaea ligustrina Ventenat, Choix de Pl. sub ¢.25. 1804; R.
Brown, Prodr. 524. 1810; Sprengel, cage Veg. 1: 35. 1824; Loudon,
Arb. Frut. Brit. 4: 2579. 1838; A. DC. in DC. Prodr. 8: 291. 1844;
Hooker f. Fl. Tasman, 1: 268. 1857: eat: Bush Friends Tasman.
t.2. 1860; F. Mueller, Pl. Colony Vict. Lithograms, ¢.54. 1864-65;
Bentham, Fl. Austral. 4: 300. 1868; F. Mueller, Fragm. Phytogr.
Austral. 8: 43. 1873 & 142. 1874; Spicer, Handb. Pl. Tasman. 125.
1878; F. Mueller, Syst. Census Austral. Pl. 92. 1882, Key Syst. Vict.
Pl. 2: 39. fig. 100. 1885 & 1: 357. 1887-8, & Second Census Austral.
Pl. 156. 1889; Maiden, Useful Native Pl. Austral. 579. 1889; C. Moore,
Handb. Fl. New S. Wales 321. 1893; Rodway, Tasman. Fl. 128. 1903;

. Dixon, Pl. New S, Wales 215. 1906; Maiden & Betche, Census New S.
Wales Pl. 172. 1916; Ewart, Handb. Forest Trees Vict. Foresters 406.
1925, & FI. Vict. 941. 1930; Anderson, Trees New S. Wales ed. 2.
132, 338. 1947; Audas, Native Trees of Austral. 242 [1935]; Bean
in Chittenden, Dict. Gard. 3: 1379. 1951; Curtis, Students’ Fl. Tas-
man. 470. fig. 108. 1967.

ayy ee ligustrina (Vent.) L. Johnson, Contr. New S. Wales Natl. Herb.
1957.

N. aie cer aie Nae, L. Johnson in Degener, New III. Fl. Hawaiian Is.
300, Nestegis

Tall evergreen shrub or small tree to 6 or even 12 m. high; young stems

1968 | GREEN, REVISION OF NOTELAEA 363

minutely puberulous, often densely so, at least when young, or rarely
almost glabrous. Leaves glabrous, petiole 2—7(-12) mm. long, glabrous
or minutely puberulous when young, especially above; lamina thick or
thickish, coriaceous, narrowly lanceolate or lanceolate (rarely narrowly
ovate) (2—)3—6(-8.5) cm. long by (0.4—)0.7-1.2(-2.5) cm. broad, scat-
tered punctate above and below; margin entire, flat; apex acute, tip acute
or obtuse; base acute or subcuneate, attenuate into the petiole; venation
obscure above and below, rarely with (4—-)5(-6) primary veins per side
just visible. Inflorescence axillary, 1-2(-3) per axil, decussate, 1.5—2.5
(—3.5) cm. long, (7—)9-11(—13)-flowered (rarely 3-4 flowers at each
node), glabrous, sometimes minutely and scattered puberulous, especially
towards the base; upper bracts narrowly lanceolate, early deciduous, 1—
1.3(—2) mm. long, ciliolate; basal bracts thickish, 1—1.5 mm. long, glabrous
or minutely puberulous, acute, relatively early deciduous, fused bases of
basal pair persistent. Flowers small, hermaphrodite (Fic. 3), yellowish to
greenish-white; pedicels 2-6(—7) mm. long (4-10 mm. long in fruit). Calyx
glabrous, very short tube 0.1-0.4 mm. long with 4 acute, triangular or
ovate-triangular lobes 0.4-1 mm. long, usually entire. Corolla with 4
ovate, acute, slightly imbricate lobes, tube very short or nil, lobes paired,
bases joined for 0.5—0.7 mm. at base of stamen, joined or adnate for 0-0.2
mm. between pairs, free portion of lobes 0.8-1.5 mm. long. Stamens 2,
anthers 1.3-1.5 mm. long, elliptic or broadly elliptic with scarcely dis-
cernible terminal appendage; filament 0.1-0.4 mm. long. Ovary bottle-
shaped, 0.8—1.5(—2) mm. long including gradually tapered style, 0.5 mm.
long, with 2 narrow stigmatic lobes 0.5 mm. long (elongating to 1 mm.
after corolla and stamens shed ?). Drupe fleshy, black, purple, red, pink
or white, ovoid-spherical, 8-12 mm. long by 6-10 mm. diameter; endocarp
crustaceous, 0.1 mm. thick.

SELECTED ExstccaTag, New South Wales. Tuross River, Apr. 1933, Arnold
(Nsw 33398); 8 miles W. of Pericoe, WSW. of Eden, about 900 m. alt., 28 May
1952, Floyd (Nsw 33399).

Views Se cated River, F. Mueller (Kk); Delegate Mt., May ce
Costin (Nsw 33400); Bonang, Feb. 1899, Bauerlen (nsw 33402); about 31 ae,
NNE. of Buchan, Gippsland, 21 Jan. 1953, Melville 3043 (x); Mt. oe a
subalpine, under 1200 m. alt., 11 Jan. 1950, Stewart (BRI 7909); 16 Mile pra
near its confluence with Howqua River, about 730 m. alt., 11 a ee

MM 2

SAN SK

Fic. 3. Notelaea ligustrina. A, flower;

- B, bud; C, ovary. (Drawn from
Comber 2189).

364 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

744 (A, K); Upper Yarra, Apr. 1911, Staer (Nsw 33404); Sherbrooke Forest,
Dandenong Ranges, Dec. 1932, Willis (kK); Sealer’s Cove, Walters (GH); Lutitt
[Loutit] Bay, F. he (pm); Cape Otway, F. Mueller (cu); Curdie’s River,
Nov. 1898, Walte

Tasmania. rs ae River near Waratah, edge of river, 180 m. alt., 3 March
1930, Comber 2189 (£, K); Port Dalrymple, R. Brown (Bm); Cataract, Launces-
ton, 20 Oct. 1880, Simson 551 (Br 7908, LE); Launceston, 22 Jan. 1842, Gunn
(pM, K); Ironstone, Dec. 1899, Rodway 6224 (Nsw 33434); Mt. Field, Nov.
1895, Rodway 6223 (Nsw 33433); Glen Leith, 14 Sept. 1840, J. D. Hooker
(Kk) & 18 Dec. 1840, Gunn (cH, nsw 41468); River Derwent, 1804, R. Brown
(pm, E); “ad ripas fluvium, in clima d’Entracasteaux,” [1801], Leschenault,
ex Herb. Jussieu & pe (c, holotype). Without locality: [1804], R. Brown

, G, K, LE, NSW 3340

Calin, ft Franklin Square, Hobart, 11 Apr. 1931, Rodway 6220
& 6221 (NSW 33436). New ZEALAND. Botanic Garden, Christchurch, Oct. 1962,
Sykes 151/62 (a, cHR 125955).

This species has the southernmost distribution (see Map 4) and, to
judge from various field notes, grows in open sclerophyll forest or scrub
dominated by species of Eucalyptus and, in Tasmania at least, on river
banks throughout the State

L, A. S. Johnson in his review of the family Oleaceae (Contr. New S.
Wales Natl. Herb. 2: 395-418. 1957) transferred this species to the res-
urrected genus Gymnelaea (more correctly Nestegis) along with the native
“olives” of New Zealand and species of Osmanthus from New Caledonia

on .

r-

? NEW SOUTH WALES
ee

VICTORIA

7S ema ve
~

TASMANIA

| @ NOTELAEA LIGUSTRINA |

Map 4. Distribution of Notelaea ligustrina.

1968 } GREEN, REVISION OF NOTELAEA 365

and Hawaii. The reason for doing this appears to have been the fact that
the aestivation of the corolla in this species is different from that of the
rest of those from Australia and not typical of Notelaea. The corolla in
N. longifolia, the type species, and in the other species too, is induplicate-
valvate in the bud (Fic. 1, A & B), but in NV. ligustrina the lobes are
imbricate (Fic. 3, B). However, the imbrication is not very strong, at
least in herbarium specimens, where there appears to be a tendency to
induplication, and some adjacent petals are distinctly valvate. Further-
more, when one considers the rest of the corolla structure it is seen to re-
semble, not that of Osmanthus, but other species of Notelaea. In Osman-
thus there is a distinct corolla tube of varying length; in the New Caledo-
nian species the corolla is campanulate with a tube at least 1.2 mm, but
usually 1.5 mm. or more long. In NV. ligustrina the tube is extremely short
or nonexistent. At the base of the staminal filaments the adjacent corolla
lobes are united for about 0.5 mm. but in the alternate position, where
there is no stamen, they may, in some cases, be joined at the base by only
0.2 mm., in other cases they appear to be merely adnate, and in yet others
they seem quite separate. This situation, with the petals in separate pairs,
joined by the base of the filament, is comparable with that in the rest of
Notelaea, and, in the family, otherwise found only in Linociera and
Chionanthus.

There is an undoubted strong phytogeographic affinity between Tas-
mania (and the southeasternmost mainland of ustralia) and New Zea-
land; and it is interesting to speculate how closely the native New Zealand
“olives” which, following Johnson (J.c.), I have treated in a separate genus
Nestegis (Jour. Arnold Arb. 44: 377-389. 1963), are related to N otelaea
ligustrina. Characters of the corolla are of primary value in the Tribe
Oleeae yet the New Zealand species are apetalous. However, there are

on Norfolk Island) and it is interesting to realize that those of Notelaea
ligustrina are recorded as red, pink, and white, as well as purple and black
(presumably very dark bluish purple), the usual color in the tribe. Never-
theless, the ripe fruits in N. venosa have been reported to be red or
white on occasion and those of V. punctata as “varicolored. :

It is not without significance, perhaps, to consider the sexuality of
flowers in Notelaea, Nestegis, and Osmanthus (the genus Linociera as
not been studied in detail yet but it is suspected that in this respect it 1s
comparable to Notelaea). In Osmanthus sects. OsMANTHUS and SrPHOS-
MANTHUs (from temperate and warm temperate Asia) and NotosMAN-
tHUs (from New Caledonia) the flowers are androdioecious. In sect.
LEIoLEA (tropical and warm temperate
warm temperate and tropical Asia and I
hermaphrodite. In Note/aea, inte N. eh ee
consistently hermaphrodite, but in Nestegzs at Is
species ) ee are eed or hermaphrodite. I am inclined to believe that

366 JOURNAL OF THE ARNOLD ARBORETUM [vot. 49

this latter condition has evolved separately from the basic hermaphrodite
condition independently in Osmanthus sect. Letocea and Nestegis. | also
think that Notelaea is of Australasian origin and is related on the one hand
to Linociera and on the other to Osmanthus and that just as unisexuality
has appeared in sect. LEroLEa so it has developed from the Australasian
stock in the genus Nestegis (which has also become apetalous). I feel
that Notelaea ligustrina stands somewhat between Notelaea proper and
Nestegis, and as a reflection of this have proposed the separate section for
its accommodation.

Notelaea ligustrina was described by Ventenat, along with his description
of the genus Notelaea and its type species NV. longifolia. His description
for N. ligustrina was contained in an “observatio” and from it one can
assume that the type is a specimen collected by Leschenault along the
margins of rivers which open into the “golfe d’Entrecasteaux, prés la
terre de Dieman” and is to be found in the Jussieu Herbarium. The type
might thus be expected to be in Paris but I have seen a specimen, ex herb.
Jussieu, from Herb. Ventenat in the Geneva Herbarium, which I believe
to be the holotype.

Notelaea sect. Mischopetala, P. S. Green, sect. nov.

A sectione Norerara corollae et androecii structura differt. Corolla
stamina etiam maturitate valde involvens; antherae parvae, sed con-
nectivum et filamenta magna; corollae lobi geminati et maturitate cum
staminibus stipitati.

TYPE sPEcIES: Notelaea johnsonii P. S. Green.

The remarkable structure of the corolla and androecium of Notelaea
johnsonii, which is the basis for establishing this monotypic section, is
discussed more fully under the species. It appears to represent an in-

dependent development from the basic stock, as represented by sect.
NOTELAEA.

9. Notelaea johnsonii P. S. Green, sp. nov.

Species inflorescentia elongata, staminibus parvis cum anthera parva
sed connectivo filamentoque latis, corollae lobis etiam maturitate valde
involutis et stipitatis, ab omnibus ceteris speciebus differt.

Evergreen shrub or small tree to 6 m. high; branchlets minutely
puberulous to glabrate. Leaves glabrous, sometimes more or less punctate;
petiole minutely puberulous or glabrate, 2-12 mm. long; lamina thickish,
coriaceous, elliptic to narrowly elliptic, sometimes slightly lanceolate,
(4-)7-10(-12) cm. long by (1—)1.8-3(—5) cm. broad; margin somewhat
thickened, very slightly and extremely shallowly crenulate or entire, some-
times somewhat undulate; apex acute, occasionall

dary veins usually obscure, sometimes almost reticulate above and below,

1968 | GREEN, REVISION OF NOTELAEA 367

Fic. 4. Notelaea johnsonii. A & B, buds; C, flower; D, paired corolla lobes
and stamen, dorsal view; E, paired corolla lobes and stamen, ventral view.
(Drawn from nsw 42340).

(6—)7-12 primary veins per side. Inflorescence axillary, 1(—2) per axil,
decussate. (4-)5-8 cm. long, 5~11-flowered, occasionally with 3(-4)
flowers per node, rachis purplish pink (fide Nsw 42340), slender, very
minutely and scattered puberulous or with a very few small hairs, at
least when young; upper bracts deciduous, unknown; basal pair of bracts
thickish 2-3 mm. long, bluntish acute to long acute, glabrous or minutely
scattered puberulent, more or less persistent during flowering then decidu-
ous, except for the persistent fused bases. Flowers hermaphrodite (Fic.
4), bluish-black (fide nsw 42340); pedicels 3-15 mm. long. Calyx 0.75-1
mm. long, with 4 thin, more or less triangular lobes, 0.4-0.5 mm. long,
minutely ciliolate, deciduous. Corolla of four, fleshy, rounded lobes, joined
at the base in pairs 1.25-1.5 mm. long at first, concave and enclosing the
stamens, eventually 2 mm. long on a “‘stalk” 0.5—0.75 mm. long. Stamens 2,
purplish-pink (fide Nsw 42340), 1.25 mm. long, with broad fleshly filament
and two small rounded anther lobes 0.3 mm. long borne at the top, with
lateral dehiscence. Ovary purplish-pink (fide Nsw 42340), conical, 1.25-—
1.5 mm. high, sometimes minutely and scattered puberulent, with a barely
perceptible style and very slightly bilobed stigma, 0.1 mm, long. Drupe
bluish-black, fleshy, obliquely ellipsoid, 1.8-2 cm. long by 0.9-1 cm.
diameter.

ateau, Lamington National Park,

Queensland. Moreton District: Robert’s Pl
(prt 7926, K), & 28 May 1929,

900-1200 m. alt., Feb. 1920, Tryon & White

368 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

White 6039 (srt 7916, K); National Park, Macpherson Range, Jan. 1919, White
(BRI 9694).

New South Wales. Whian Whian State Forest, N. of Lismore, about 600 m.
alt., 15 Jan. 1953, Constable (Nsw 22217); Goonengerry, near Mullumbimby, 7
Nov. 1963, Hayes in Green 1529 (A, K); Bungalow-Byron Bay road, 16 Nov.
1964, Williams K18 (a, Nsw 72725); Wyangarie State Forest, N. of Kyogle,
about 900 m. alt., 21 Nov. 1963, Hayes & Floyd 9 (a); Lismore, Aug. 1906,
Rothwell (Nsw 44053); Wollongbar Experimental Farm, east of Lismore, about
150 m. alt., 11 June 1957, Johnson & Constable (Nsw 42340, holotype; K, iso-
type) ; Rous, 21 July 1925, Cheel (Nsw 44055); Clarence River, 1868, Beckler
(BM, K).

This most distinctive species, distinctive at least as far as the flower
structure is concerned, is restricted to a relatively small area in northeast
New South Wales and adjacent Queensland (Map 2). Confined to rain-
forest, and to judge from the limited field notes to basaltic and basic soils,
it is often found surviving today in small patches of rain-forest species left
isolated by the spread of cultivation.

The oldest specimen seen for this revision is that gathered by Beckler in
the area of the Clarence River and sent to England by Ferdinand voa
Mueller where it was labelled Notelaea venosa. However, most of the
material had been determined previously as N. longifolia, from which it
can be quickly identified by the rather obscure venation of both the upper
and lower surfaces of the leaves.

Detailed observations on fresh flowering material would be valuable,
but the dried flowers, especially when softened in hot water, show the
unusual corolla and stamens. Throughout the other species the petals are
joined together in pairs, touch each other laterally and surround the ovary.
In this species the pairs are separate even in young flowers (Fic. 4, A & B)
and, as they reach maturity, they come to stand on either side of the
ovary on a stalk-like structure which is possibly a development of the
filament (Fic. 4, C-E). The individual stamens are distinctive too. The
anthers are small and completely enclosed in the permanently concave and
apparently fleshy corolla, except for the top of the connective and the two
small and lateral openings of the pollen sacs (Fic. 4, D & E). Further
observations and more collections may well improve our knowledge of this
plant and its flower and perhaps show that this species should be distin-
guished as a separate genus; but for the present its sonata in the
genus Notelaea is indicated by the provision of a ram secti

It is a pleasure to name this plant after Mr. L. A. S ohieon ‘of the
National Herbarium of New South Wales, who has made important con-
tributions to the study of the Oleaceae, has kindly made extensive notes
and first-hand field observations on the genus Notelaea available to me,
and was joint collector of the type material.

EXCLUDED NAMES

Notelaea cap Picconia ie Knobl. in Engler & Prantl. Nat. Pflanzenfam.
4(2): 10. 1892 = Picconia D

1968 | GREEN, REVISION OF NOTELAEA 369

Notelaea austro-caledonica Vieill. Bull. Soc. Linn. Normand. 9: 345, 1865 =
Osmanthus austro-caledonicus (Vieill.) Knobl.

N. azorica Tutin, Jour. Bot. 71: 101. 1933 = Picconia azorica (Tutin) Knobl.

N. badula Vieill. ex Panch. & Sebert, Not. Bois Nouv.-Caléd. 184. 1874 =
Osmanthus austro-caledonicus (Vieill.) Knobl.

N. brachystachys Schltr. Bot. Jahrb. 39: 228. 1906 = Linociera brachystachys
(Schltr.) P. S. Green.

N. collina Schltr. Bot. Jahrb. 39: 229. 1906 = Osmanthus austro-caledonicus
(Vieill.) Knobl.

N. cymosus Guillaum. shee — Bot. France 89: 232. 1943 = Osmanthus cymos-
us (Guillaum.) P. S. G

N. eucleoides ee Bot. “Jahrb, 39: 229. 1906 = Osmanthus austro-caledonicus
(Vieill.) Knob

N. excelsa taey Webb & Bert. Hist. Nat. Il. Canar. “at (Phyt. Canar. sect.
3): 163. 1845 & #.186. 1848 = Picconia excelsa (Ait.) D

N. Bavicares (Vahl) A. Dietr., Linn. Sp. Pl. ed. 6, 1: 246. ay D. Dietr. Synop.
igh 1839 = Noroahix emarginata (Lam.) Thouars ex Hook.

N. Vranes Gaiiiauh: Bull. Mus. Hist. Nat. Paris 28: 198. 1922 = Linociera
brachystachys (Schlitr.) P. S. Green.

N. glandulosa eae ex Guillaum. Ann, Mus. Col. Marseille II. 9: 192. 1911,
nom syn. = Olea paniculata R. Br. (0. thozetii pe & Sebert).

N. lanceolata Teijem, & Binn. Nat. Tijdschr. Ned. Ind, 27: 3 he = Linociera
montana ( . Don. Type: Java, M. Binnendyck rei

N. monticola Schltr, Bot. Jahrb. 39: 229. 1906 = Osmanthus ae (Schitr.)

Knobl.

N. ? paniculata Guillaum. oe vie Bot. France 89: 233. 1943 = Linociera
eae (Guillaum.) P. $.G
N. posua D. Don. Prodr. Fl. Nepal. 107. 1825 = Osmanthus fragrans Lour.
WN: fags Rates (F. Muell.) Hemsl. Ann. Bot. 10: 243. 1896 = Linociera
quadristaminea (F. Muell.) Knobl. @
N. rostrata Teijsm. & Binn. Cat. Hort. Bogor, pie Oy, nom. Seat g Linociera
montana (BI.) G. Don. Type: Java, M. Binnenaycr (B06,
N. rostrata Teijsm. & Binn. var. latifolia Teijsm. . et aoe nud. =
i . Type: Java, 7nnen
Linociera montana (BI.) G. Don. Ty Ja Tei ein Le 10h =?
nociera montana (Bl.) G. Don. TyPE: no se
N. cao Schltr. ge Jahrb. 39: 550. fs. 22. 190
caledonicus (Vieill.) K
N. zeylanica Gardner ex "Thwaite
Olea polygama Wight. j 864 =
N. zollingeriana Teijsm. & Binn. Nat. Tijdschr. Ned. ak 1
Olea maritima Wall. ex G. Don. Type: Java, M. Binnendy ori

Osmanthus austro-

s, Enum. Pl. Zeyl. 188. 1860, momen in syn. =

RoyaL Botanic GARDENS, KEw
RICHMOND, SURREY
GREAT BRITAIN

370 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

COMPARATIVE ANATOMY OF THE LEAF-BEARING
CACTACEAE, XVII.

PRELIMINARY OBSERVATIONS ON THE PROBLEM OF
TRANSITIONS FROM BROAD TO TERETE LEAVES *

+1. W. Batrey ”

Two OF THE Most significant phenomena in adaptations of the Cacta-
ceae for survival in arid environments are (1) the retention of moisture
in abundantly occurring “mucilage” and (2) the reduction of leaves in
Opuntioideae and their elimination in Cereoideae with concomitant trans-
fer of ellen inSoe activity to green parts of stems. Both phenomena

Siv
ecological, nlivuologial and biochemical points of view than they have
received thus far

A ia. eT and phylogenetically important aspect of the second
phenomenon is the question, how many transitions from ancestral broad
leaves to terete ones have developed in the Opuntioideae? In my ex-
tensive collections of Pereskia, Pereskiopsis, and Quiabentia 1 have not
encountered such transitions up to now. However, they do occur in certain
representatives of Ne Opuntioideae, particularly in Opuntia subulata
(Muehlenpf.) Enge

In my collection at this plant [Moran 7268] the leaves are relatively
large and persistent. The vasculature in the shorter, broader, less fre-
quently occurring ones closely resembles that which occurs in leaves of
comparable forms and sizes of Quiabentia pereziensis Backbg. (compare
Fics, 1 and 2). In both cases the venation of the appendages is palmate
as in Pereskiopsis and Quiabentia, rather than dominantly pinnate as in
the large, thin leaves of such more primitive pereskias as P. sacharosa
Griseb., P. grandifolia Haw., P. bleo DC., and related species. For dis-
cussions of variations in the venation of leaves and cotyledons and their
phylogenetic significance, see 1960 and 19653

In Opuntia subulata (Austrocylindropuntia subulata (Muehlenpf.)
Backbg.) none of the abundant more elongated leaves, although relatively
slender, are truly terete, i.e., circular in cross section. All of them exhibit

+ This paper was completed by Professor Bailey shortly before his death on May

16, 1967. He had planned the pertsene 2 and discussed them with the artist, Elmer
W. Smith, who completed them subseque

y.
‘ Terete is defined in standard peneeneevk as cylindrical and tapering with circular
cross section,

* This aga was supported in part by a grant from the National Science
Foundation, GB-29

* Previous tee a this series are listed by their dates of publication.

1968 | BAILEY, LEAF-BEARING CACTACEAE, XVII 371

more or less extensive broadening laterally. The two systems of veinlets
in opposite sides of the lamina are derived largely from independent vas-
cular strands extending outwardly from the base of palmately veined ap-
pendages, except in the upper central parts of the leaves where more or
less numerous veinlets may be related to the conspicuous paired vascular
traces in the center of the lamina. In such leaves, the two systems of
veinlets tend to curve more or less extensively in opposite sides of the
lamina toward the abaxial surface (Fic. 4). As they do so, more or less
characteristic palisade tissue becomes concomitantly extended laterally,
but does not jacket the abaxial surface.

In contrast to this, the evanescent truly terete leaves of such opuntias as
Opuntia ficus-indica (L.) Mill. and O. cylindrica (Lam.) DC., particularly
during earlier stages of their development, are jacketed by palisade tissue
(Fic. 5). In such tiny terete leaves, the vasculature is much modified.
The paired traces in the center of the lamina commonly are massive in
appearance and exhibit frequently occurring fusions. Independent vas-
cular strands in the basal part of the lamina tend to be eliminated. The
two systems of veinlets in opposite sides of the lamina, although reduced
and modified, commonly tend to be conspicuously coarse and related to
the two more or less fused mid-veins in the center of the lamina rather
than to independently occurring lateral vascular strands as in the large
leaves of O. subulata.

In my collections of Opuntia ficus-indica preserved in FAA the leaves
vary from circular to oval and exhibit conspicuous adaxially oriented
curvatures which may have been intensified during fixation and dehydra-
ion. In some of them the two systems of coarse veinlets coalesce in the
abaxial side of the lamina (Fic. 5), whereas in others they fail to do so.
According to Dr, Boke (1944, and personal communication) the two sys-
tems of veinlets in O. cylindrica do not coalesce in the abaxial side of
matured leaves. ;

Thus, in dealing with transitions from broad palmately veined leaves
to truly terete ones, there are three trends of phylogenetic specialization
which merit detailed consideration. These are (1) changes in the occur-
rence, structure, and distribution of palisade tissue, (2) alterations in the
principal vascular strands that extend outward from the base of palmately
veined appendages, and (3) reduction and modification of two systems of
veinlets in opposite sides of the lamina. ae

The large, thin, pinnately vascularized leaves of primitive pereskias do
not form typical palisade tissue on their adaxial side. However, there is
a more or less conspicuous tendency to do so on the adaxial side in the
case of the palmately veined leaves of Pereskiopsis and Quiabentia (Fic.
3). It is evident in the limited collections available to me that the second
and third trends of morphological specializations in terete leaves are not
consistently similar in minor details of structure, even in different leaves
from the same plant. Therefore, it is essential to obtain and compare leaves
from as many representatives of the Opuntioideae as possible in a com-

ee

JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

——s
28

EIS
Sy
an
SS

Se;

LOPS
STN Ee
Bor

LS

55)

esr

a

KeZ

3

So
Fs
as teu

IE,
FE ESE

ie

Fz

=.
SS

oe

zs
Ss ie

LO 8 2 ABS OG ey

cere

Fics. 1-5. Vascularization of leaves of Cactaceae. 1. Leaf of Opuntia subulata
(the shorter, broader type of leaf occurring in this species). 2. Leaf of Quia-
bentia pereziensis. 3. Diagrammatic cross section of a leaf similar to that in
FIG. 1 or to a typical palmately veined leaf of Pereskiopsis or Quiabentia.
Palisade tissue on the adaxial side is indicated by stippling. 4. Diagrammatic
cross section of one of the more abundant elongate leaves of Opuntia subulata.

1968 | BAILEY, LEAF-BEARING CACTACEAE, XVII 373

prehensive search for additional transitional stages in the development of
terete leaves. Such an extensive investigation is being undertaken by Pro-
fessor Norman Boke and myself.

In comparing the form and vasculature of modified leaves, it is neces-
sary to avoid utilizing the appendages that occur on the uppermost external
surface of the invaginated tori of flowering shoots. Such modified leaves
in the Pereskioideae and Opuntioideae frequently exhibit conspicuous
broadening at their base and patterns of vasculature which simulate those
that occur in the first-formed tepals on the inner surface of the invaginated
tori. These foliar modifications obviously present an independent phy-
logenetic problem and one that merits detailed investigation in the future.
Therefore, in searching for transitions from broad leaves to terete ones,
I have focused attention upon the foliar appendages of vegetative shoots
and have avoided those occurring on the uppermost invaginated part of
flowering ones.

Although possible variations in the form and major details of vascula-
ture in the leaves of additional representatives of the Opuntioideae merit
investigation, there is, even in limited collections now available, cogent
evidence of the derivation of terete leaves from broad, palmately veined
ones. It should be emphasized in this connection, that without knowledge
of significant transitional stages occurring in such opuntias as Opuntia
subulata, it might be erroneously assumed a priori that the highly modified
vasculature of terete leaves is developed by transitional modifications of
broad pinnately veined appendages.

The only cactus that I know of, exclusive of the Opuntioideae, which
forms truly terete leaves is the genus Maihuenia. Three collections of M.
poeppigit (Otto) Weber, obtained from Chile by Hutchison in 1951, are
under cultivation at the University of California Botanical Garden in
Berkeley, California. These plants have grown well in the open, flowering
and fruiting annually, In material kindly sent to me [Hutchison 306],
the numerous short, slender branchlets in the upper part of a plant bear
compact clusters of terete leaves terminally. In such branchlets there is
a more copious production of ‘‘mucilage” than I have observed in any
other cactus. Furthermore, there is a very precocious development of
encasing periderm in parts of stems immediately subtending the cluster
of terminal terete leaves. Photosynthetic activity appears to be largely
confined to these long-persistent appendages. The terete leaves at matu-
rity are jacketed on all sides by very conspicuous palisade tissue which con-
tains much enlarged and elongated mucilage cells. As in terete leaves of
opuntias, there is a similar tendency for fusion of the paired traces in
the center of the lamina, and the two systems of coarse veinlets coalesce
in the abaxial side of the appendages.

Palisade tissue extends laterally but does not cover abaxial surface. 5. Diagram-
matic cross section of truly terete leaf of Opuntia ficus-indica. Palisade tissue
extends all around. (Figures not drawn to scale.

374 JOURNAL OF THE ARNOLD ARBORETUM [ VoL. 49

DISCUSSION

In previous papers of this series, I have emphasized that the Cactaceae
provide unusually comprehensive evidence in the study of highly diver-
gent trends of morphological specializations. These are so bizarre and so
closely correlated from ecological and physiological points of view with
adaptations for survival in excessively arid environments as to leave few
uncertainties regarding the major directions in which phylogenetic changes
are progressing.

e most primitive surviving representatives of the Cactaceae from
both reproductive and vegetative points of view are considered to occur
in the genus Pereskia. Such better known species as P. sacharosa Griseb.
(1962), P. grandifolia Haw. and P. bleo DC. (1963c), with their large,
thin leaves have long been recognized by taxonomists as having a habit of
growth and absence of conspicuous succulence characteristic of many
typically woody dicotyledonous trees and large shrubs. The leaves of
these species have dominantly pinnate vasculature, and photosynthetic
activity is largely confined to them. In contrast to this, the relatively
thick leaves of Pereskiopsis and Quiabentia (1960) have prevailingly
pseudo-palmate and palmate vasculature. Photosynthetic activity occurs
both in leaves and in green parts of succulent elongating shoots (1964a).

Incipient stages of transition from pinnate to pseudo-palmate venation
occur in some of the morphologically more specialized species of Pereskia,
particularly in P. pititache Karw. and allied taxa (1960). Summations
of evidence from leaves and cotyledons demonstrate that the transitions
in vasculature of the Cactaceae are from pinnate to palmate, rather than
vice versa (1965). Furthermore, incipient modifications of that dense,
strongly lignified secondary xylem of such primitive pereskias as P. sacha-
rosa and P. grandifolia (1963c) occur in various morphologically more
specialized species (1962, 1963a, 1963b, 1963d).

However, Pereskiopsis and Quiabentia differ significantly from Pereskia
in the absence of sclereids in their secondary phloem (1961a) and in the
structure of the outer tissues of their succulent stems. In young green

stomata and intercellular spaces in the chlorenchyma being through special
channels in the collenchyma which are jacketed by thick unpitted walls
and which tend to have a tenuous inner lining of cutin.

Although a crystalliferous hypodermis is absent in pereskias, and a

1968 | BAILEY, LEAF-BEARING CACTACEAE, XVII 375

structurally homologous form of hypodermis does not occur in the Cereoi-
deae, it develops in characteristic diagnostic form in Opuntia and related
genera of the Opuntioideae, being present in representatives having flat-
tened green stems, e.g. O. ficus-indica, as well as those with cylindrical
green shoots, e.g. O. cylindrica. It provides significant evidence of rela-
tively close genetic relationships and supports taxonomists (Backeberg,
1966) who include the broad-leaved genera Pereskiopsis and Quiabentia
in the subfamily Opuntioideae. It strengthens my conclusion that the
vasculature of terete leaves in this subfamily develops by modifications of
palmately veined appendages.

At present the taxonomic affinities of Maihuenia with its terete leaves
are obscure and uncertain. To place this genus with its highly specialized
stems and xylem in the primitive subfamily Pereskioideae (Backeberg,
1966) appears to be premature and discordant. To include it in the Opunt-
ioideae merely on the basis of its terete leaves would be equally unreliable.
Similarities in the vasculature of these terete leaves may be due to parallel
or convergent evolution rather than indicative of close genetic relationship.
Similarities in its pollen and in the highly modified structure of its sec-
ondary xylem are suggestive of possible relationship to the Cereoideae,
but other morphological features of the genus raise doubts about such a
relationship. It is evident that summations of evidence from all organs
of the plant and from all species of the genus must be obtained and
harmonized in order to determine, if possible, its taxonomic relationships.

LITERATURE CITED

BACKEBERG, C. 1966. Das Kakteenlexikon. Gustav Fischer. Stuttgart. Clas-
sification Paper 17—

Battey, I. W. 1960. Comparative anatomy of the leaf-bearing Cactaceae. I.
Foliar vasculature of Pereskia, Pereskiopsis and Quiabentia. Jour. Arnold
Arb. 41: 341-356.

. 1961a. II. Structure and distribution of a geenden in the phloem
of Pereskia, Pereskiopsis and Quiabentia. Ibid. 42: -156.
. 1961b. III, Form and distribution of crystals in aa Pereskiopsis

and Quiabentia. Ibid. 334-346.

. 1962. VI. The xylem of Pereskia sacharosa and Pereskia aculeata. Ibid.

43: 376-388.

. 1963a. VII. The xylem of pereskias from Peru and Bolivia. /bid.

44: 127-137.

. 1963b. VIII. nei Sadie, of pereskias from southern Mexico and Central

America. Ibid. 21

, A063c.. TX. ig . of Pereskia grandifolia and Pereskia bleo. Ibid.

222-231.

. 1963d. X. The xylem of Pereskia shegge vy eet Pereskia guamacho,

Pereskia cubensis, and Pereskia portulacifolia. Ibid. 3

1964a. XI. The xylem of Pereskiopsis and Sebo. Ibid. 45: 140-

376

JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

1964b. XII. Preliminary observations upon the structure of the epider-
mis, stomata, and cuticle. /bid. 374-389.
. 1965. XIV. Preliminary observations on the vasculature of cotyledons
Ibid. yi 445-4
944. Histogenesis of the leaf and areole in- Opuntia cylindrica.
Am, Jour. Bot. 31: 299-31

1968] JOHNSTON, THE TEA SAGERETIA OF CHINA 377

BOTANICAL NAME OF THE TEA SAGERETIA OF CHINA

MARSHALL C. JOHNSTON

THE PAUPER’S TEA of China was first described by Linnaeus’ one-time
student and collector, Pehr Osbeck, on page 232 of his ‘‘Dagbok 6fwer en
Ostindisk Resa . . .”’ published in 1757. The short descriptive paragraph,
freely translated fa the Swedish, reads as follows:

Rhamnus Thea, or Pauper’s Tea, is a fathom-high bush, the leaves of which
are similar to those of common tea; but the flowers belong to the first Ordo of
the fifth Class, are quite small and are fastened to the far ends of the branches

which are again branched. Used as a tea by the poor, but in this place [the
Danish Island] it served as a hedge. Called “Tja” Th e., tea] in Chinese.

Linnaeus himself included some of Osbeck’s Chinese plants in the 1753
edition of the Species plantarum (William T. Stearn, Introduction to the
Ray Society facsimile edition, 1957), but not the pauper’s tea. He did not
notice the species in print until 1771 when, on page 207 of the Mantissa,
he re-named it Rhamnus theezans. Immediately under his phrase-name,
Linnaeus cited as a synonym the “Rhamnus Thea Osb. it. 232,” and he
followed this with an unusually full description. Linnaeus’ epithet has
been used for the same species ever since 1771. In 1826 it was combined
with the generic name Sageretia A. Brongn. The name Sageretia theezans
has survived in recent years because of the faith placed in the careful
bibliographic work of C. Schneider (Pl. Wilson. 2: 227. 1914) and A.
Rehder (Bibliogr. of Cult. Trees and Shrubs 436. 1949). sehneider and
Rehder both deemed Osbeck’s name a “nomen nudum” or a “nomen.”
They presumably considered Osbeck’s few comments insufficient to con-
stitute a description.

The question must be raised again, however, whether Linnaeus’ name
can stand under the present code of nomenclature. If Osbeck’s name can
be clearly typified, Linnaeus’ name must be considered a superfluous, un-
usable synonym, despite the scantiness of Osbeck’s description.

Osbeck’s collections were turned over to Linnaeus. As far as I can
determine through search in various herbaria and through correspondence,
the only surviving Osbeck collections from China are those in the Linnaean
Herbarium in London. I was able to examine and photograph parts of the
Linnaean Herbarium in April, 1967. Two specimens are pertinent here,
sheets 262.7 and 262.8 as numbered by Savage (Cat. Linn. Herb. 212.
1945). The latter, 262.8, is a mere fragment pinned to the other. Sheet
262.7 is an excellent flowering specimen, annotated “Rhamnus theezans’
by Linnaeus himself. I can find no evidence that Linnaeus received speci-
mens of this species from any other source than Osbeck. In the absence

378 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

of any information to the contrary, it may be concluded reasonably that
262.7 was prepared in China by Osbeck. I here designate it as the type
of both Rkamnus Thea and Rhamnus theezans. This lectotypification
serves to put Linnaeus’ synonymy on a firm basis and to make quite clear
that his epithet is superfluous and must give way to the prior one of
Osbeck. Below, I give the necessary combination and the known and
suggested synonymy.

Sageretia Thea (Osbeck) M. C. Johnst., comb. nov.

Rhamnus Thea Osbeck, Dagb. 6fwer Ostind. Resa 232. 1757; P. Osbeck, Reise
nach Ostindien und China, translated by J. G. Georgi, 210, 302. 1765; P.
Osbeck, A voyage to China and the East art transl. and with Faunula
and Flora Sinensis by J. R. Forster, 1: 375. 1771; 2: 345. 1771

Rhamnus theezans L. Mant. 207. 1771; J. F. na Syst. 2: 398. 1791; Poir.
in Lam. Encycl. 4: 464. 1796; Willd. Sp. Pl. 1: 1094. 1797; R. & S. Syst
5: 281. 1819; HBK. Nov. Gen. et Sp. 7: 54. 1824; Spreng. Syst. 1: 767.
1825: DC. Frode 2: 76. 182

tangs theezans (L.) Brongn. Mém. Fam. Rhamn. 53. 1826; G. Don, Gen.
S 8. 1832; Benth. in Hook. Jour. Bot. 4: 42. 1852; Benth. Fl. Hong-
kong. 68. 1861; Maxim. Mém. Acad. St.-Pétersb. ser. 6. 10: 20. 1866; Mig.
Ann. Mus. Bot. Lugd. Bat. 3: 32. 1867 (as “theesans’’); Lawson in Hook. i.
Fl. Brit. Ind. I. 1: 641. 1875; Hemsl. Jour. Linn. Soc. 23: 131. 1886; Mat-
samura, Bot. Mag. Tokyo 12: 22. 1898; C. Schneider, Pl. Wils. 2: 227.
1914; Rehder, Jour. Arnold Arb. 8: 166. 1927; Handel-Mazzetti, Symb.
Sinicae, 7; 674. 1933; Rehder, Jour. Arnold Arb. 15; 13. 1934; Kanehira,
Formosan Trees, rev. ed. 428. 1936; Rehder, oe Cult. Trees and
Shrubs 436. 1949: Li, Woody FI. of Taiwan 516-518

Ampeloplis chinensis Raf. Sylva Telluriana 33. 1838.

Sageretia theezans var. integra Dum.-Cours. ex G. Don, Gen. Syst. 2: 28. 1832,
the generic name spelled ‘“Segeretia.”

Sageretia theezans var. diospyrifolia Lawson in Hook. f. Fl. Brit. Ind. I. 1:
641. 1875, as “diospiryfolia.”

Berchemia Chaneti Léveillé, Repert. Sp. Nov. 10: 433. 1912

Sageretia theezans var. tomentosa C. Schneider, Pl. Wils. 2: 228. 1914.

Sageretia Chanetii (Léveillé) C. Schneider, loc. cit., a provisional combination.

Sageretia Hayatae Kanehira, Formosan Trees rev. ed. 426. 1936; this synony-
my suggested by Li, Woody Fl. of Taiwan 516-518. 1963.

spd taiwaniana Hosokawa ex Masamune, Trans hg an Soc. For-

osa, 28: 286. 1936; this synonymy suggested by ae loc

sSuretet stkayoensis Masamune, loc. cit.; this synonymy he by Li,

OC. Cth.
?Sageretia theezans var. microphylla Suessenguth ex Suessenguth & Overkott,
Repert. Sp. Nov. 50: 329. 1941; tentative synonymy suggested by Li, /oc.

Of course, in a genus such as this one, so badly in need of monographic
work, the synonymy is tentative at best and subject to much revision. The
taxonomy of the Chinese species is particularly chaotic.

It is a pleasure to acknowledge the help of Dr. [ka Kukkonen in trans-
lating Osbeck’s paragraph, and of Carroll Wood and Bernice G. Schubert

1968 | JOHNSTON, THE TEA SAGERETIA OF CHINA 379

in going over the preliminary manuscript and making helpful suggestions.
The opportunity to study specimens in the Linnaean herbarium was pro-
vided through National Science Foundation grant GB—3138 to the Texas
Research Foundation.

DEPARTMENT OF BOTAN

THE UNIVERSITY OF TEXAS
AUSTIN, TEXAS 78712

VoLUME 49 NUMBER 4

JOURNAL

OF THE

ARNOLD ARBORETUM

HARVARD UNIVERSITY

B. G. SCHUBERT
EDITOR

T. G. HARTLEY Cc. E. WOOD, JR.

D, A. POWELL
CIRCULATION

- = |
mol a. ,.. OCTOBER, 1968

THE ARNOLD ARBORETUM OF HARVARD UNIVERST
{BRIDGE. ‘MASSACHUSETTS

THE JOURNAL OF THE
ARNOLD ARBORETUM

Published quarterly by the Arnold Arboretum of Harvard University.

Subscription price $10.00 per year.

Volumes I-XX, reprinted, and back issues of volumes XXI-XLV are
available from the Kraus Reprint Corporation, 16 East 46TH SrRezt,
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Subscriptions and remittances should be addressed to Miss Dunct A.
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HUSETTS 02138.

CONTENTS OF NO. 4

Tus Ecoioey or AN ELFIN Forest In Pusrto Rico, 1. Intropuc-
TION AND CoMPosITION Srupises. Richard A. How owared ...........- 381

Tue Ecotocy or sn Exrin Forest in Pusrto Rico, 2. THE
Microciimate oF Pico peu Orste. Harold W. Baynton sie 419

Srupres se le IsutanD Piants, XIX. Tum ARALIACEAE
THE Hee ag Samoa, AND monae Albert C.
Smith ist Benjamin C. Stone 43

SorBUs AND THE ProsLeM or GENERIC aes, aghast K.
Brizicky

Emsryotocy or Drimys winter. N. N. Bhandari and Revath
Venkataraman

Tue Direcror’s Reporr
INDEX TO Votume 49 .

*

Volume 49, No. 8, including pages 291-380
was issued July 16,1968.

JOURNAL

OF THE
ARNOLD ARBORETUM

VoL. 49 OcTOBER 1968 NUMBER 4

THE ECOLOGY OF AN ELFIN FOREST IN PUERTO RICO,
1. INTRODUCTION AND COMPOSITION STUDIES

RicuHarp A. HOWARD

ON THE SUMMITs of mountains in many of the islands of the Antillean
archipelago are stunted and dwarfed forests composed of dense stands
of contorted shrubs and small trees. The forest floor as well as the stems,
branches, and exposed bases of the components of the forest are covered
with festoons of bryophytes, almost lined with vascular epiphytes, and
intertwined or overburdened with woody and herbaceous twiners. The
short vegetation on these mountain tops seems always windswept and
bathed in mist and clouds, so that an impression of a wet, cold, nearly
impenetrable tangle has led to the appellation of elfin-, mist- or cloud-
forest, -woodland, or -thicket. The inaccessibility of these zones, which
in most cases involves difficult and uncomfortable travel, has restricted
visits by botanists and has resulted in few studies of the biology of such
areas,

In the Luquillo Mountains of eastern Puerto Rico, trails and roads to
the summit of Pico del Este and El Yunque have made several examples
of the dwarfed forest easily accessible. In 1965 a road was constructed
to Pico del Este, the easternmost peak of the Sierra del Luquillo where
radio and radar facilities were established by the United States govern-
ment. Access to this road was granted to the Arnold Arboretum staff by
personnel of the Institute of Tropical Forestry and the U.S. Navy, and
a study site was selected on the undisturbed summit of Pico del Oeste
(West Peak) at an elevation of 1050 meters above sea level (Fics. 1, 2, 3,
4). This peak, on the border of Municipio de Ceiba and Municipio de
Naguabo, is located at 18° 16’ 47” and 65° 45’ 47”; it is but two feet
lower than Pico del Este, 50 feet less than El Yunque, and is the third
highest peak in the Luquillo range, and fourth highest in Puerto Rico. A
small flat-roofed shelter was constructed on the summit to protect instru-
ments, equipment, and personnel and to serve as an observation platform.
A study area was marked along a very old and unused foot trail which
crossed the summit through the forest. Instruments were secure in this
isolated area and individual plants could be marked and studied over an
extended time period. The roof of the shelter at the level of the canopy

382 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Fic. 1 (Aspove). A view of the NE slope of Pico del Oeste from the parking
area. ‘ache to the trail is made in the grassy area special by the figure and
arrow. va mt is to the right of the cri of Prestoea

ms. 2 { ow). A view of the south side of Pico del ae showing the
Petter summit. The summit and the observation hut are near the left shoulder
on the top. The 10 0-yard-square transect area is at the right shoulder. The
mountain palm appears only in the valleys

1968 | HOWARD, ELFIN FOREST, 1 383

offered a wonderful view of eastern Puerto Rico on rare occasions and
permitted close observation of plant and animal activity immediately
above the forest. Observations and collections of material were made over
a period of nearly three years to determine the answers to three basic
but deceptively simple questions: what was present; under what con-
ditions did it live; and what was it doing?

This is the first of a proposed series of papers, to : epee by the
many collaborators, which will describe this dwarfed

This study was ‘supported generously by a grant ea the National
Science Foundation (GB: 3975) for which I am most deeply indebted.
The use of the site within the Luquillo National Forest was made possible
by a codperative agreement with the Forest Service, United States De-
partment of Agriculture, with the endorsement of Dr. Frank Wadsworth,
Director of the Institute of Tropical Forestry, Rio Piedras. All partici-
pants in the field study share my deep and personal indebtedness to Mr.
Joseph B. Martinson, who made available a modern house at a lower
elevation which supplied most pleasant living accommodations and a
laboratory area for the duration of this project. The continuity of obser-
vations was possible only through participation of many members of the
staff of the Arnold Arboretum and pro tempore associates. I list these
individually, for their many contributions are anonymous in the data to
follow: Judy Appenzeller, Michael Canoso, Dr. and Mrs. Gordon DeWolf,
Jr., Mr. and Mrs, Henry Draper, Mr. and Mrs. Lawrence Evans, Madelon
Gauer, William Gensel, William Grime, Thomas Hartley, William McClain,
Luis Maldonado, Dulcie Powell, Stephanne Sutton, Robert Wadsworth,
Dr. and Mrs. Richard Wagner, Nathaniel Whittier, Carroll E. Wood, Jr.,
Roy Woodbury, Dr. and Mrs. Donald Wyman.

A special word of appreciation is due to members of my family, my wife,
Elizabeth Howard, and my children, Jean, Barbara, and Bruce, for the
tedious work of collection and measurement of components of the forest
during two summers.

The professional collaborators who took part in this study directly or
used material collected by others will present their own observations in-
dividually or jointly. Dr. Harold Baynton and his associates at the Na-
tional Center for Atmospheric Research, Boulder, Colorado, assumed
responsibility for the meteorological data compiled from their own equip-
ment installed in the study area. Malcolm Gill and Walter Lyford of the
Harvard Forest studied the roots of the plants and the soils, respectively.
Margaret Fulford and students from the University of Cincinnati collected
the liverworts, which dominate the non-vascular epiphytic flora. John
Foerster of the University of Connecticut will report on the algal com-
ponents of the area. Henry Imshaug and his students have studied the
lichen flora. Harvey A. Miller of Miami University and the University of
Washington, and his associates, collected mosses and other bryophytes.
Georgia Persinos of the Massachusetts College of Pharmacy supervised
the analysis of much material for alkaloid and other chemical constituents.
Additional studies on the biologically active constituents of certain spe-

384 JOURNAL OF THE ARNOLD ARBORETUM [voL, 49

Fic ( ee OVE ) The ridge of the Luquillo Mountains extending in a south-
easte tly oe tion. The radar dome on Pico del Es te is evident on the cloudy day.
Pico del Sete although a bit lower, dominates this picture and is marked by an

Ww.
Fic. 4 (BELtow). A view of the uniform ror yice - the zrarbyp Les spre “i
Oeste. The observation hut and instrument shelter o the rig alum
num tower holding the wind giana Tabebuia rivide and eed palronne
dominate the canopy vegetatio

1968 | HOWARD, ELFIN FOREST, 1 385

cies have been undertaken by the National Institutes of Health and the
laboratory of Dr. Norman Farnsworth of the University of Pittsburgh.
Dr. Richard Wagner conducted many studies of the phytochemistry of
individual species of the vegetation, and with Mrs. Wagner, was respon-
sible for drying and shipping many hundreds of pounds and numerous
packages of study materials. Rolla Tryon of the Gray Herbarium identi-
fied the pteridophytes collected by others on Pico del Oeste. David Gates
of the Missouri Botanical Garden contributed to our knowledge of the
physics and environment of the dwarfed forest. William Theobald of
Occidental College is investigating the anatomical structure of the woody
species. J. L. Lappuck and his associates at the Waltham Laboratories
completed the analysis of a series of water samples. Alfred Fordham,
propagator of the Arnold Arboretum, has investigated the germination and
rooting of many plants of the area and cared for the many species grown
at the greenhouses of the Arnold Arboretum. My colleague and com-
panion on much of the work, Lorin I. Nevling, Jr., has been responsible
for studies of the floral biology and phenology, the distribution of airborne
particles, and the cytological investigations.

PREVIOUS AND COMPARABLE STUDIES

Mist or cloud forests in which the component plants support an exten-
sive layer of non-vascular epiphytes and therefore could be called a mossy
forest have been reported for many tropical countries in Asia, Africa,
Australia, Central and South America, and in the West Indies. P. W
Richards’ excellent summary The Tropical Rain Forest (1964) offers
many references to basic papers applying to such areas which should be
consulted for detailed descriptions. In a chapter on the tropical rain
forest at its altitudinal and latitudinal limits, Richards stated that the
tropical rain forest with altitudinal ascent “gives way to a formation-type
which it is proposed to call the Submontane Rain forest; this at a still
higher altitude is succeeded by another formation-type, the Montane Rain
forest. In many places, especially on exposed ridges and isolated peaks,
the Montane Rain forest consists of dwarfed crooked trees smothered
with an overwhelming abundance of epiphytes, especially hepaticae and
mosses; this very characteristic type of vegetation is often called Mossy
forest or Elfin. woodland.”

Asia

Seifriz (1923) describes a mossy forest on Mount Gedeh in Java at
5,500 ft. and near the summit at 8,000 ft. where tree species are few and
the plants crooked, “moss-bedecked” and laden with vascular epiphytes.

Van Steenis (1935) recognized in Malaysia five zones of vegetation in
which the critical altitudinal limits proved to be 1,000, 2,400, 4,000, and
4,500 meters. His montane zone of 1,000 to 2,400 meters has the forma-
tion-type of vegetation of the montane rain forest including the mossy
forest or elfin woodland.

Lane-Poole (1925) described a mossy forest for Papua and New Guinea

386 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

which occurred at an average altitude of 7,500 ft. His description in-
cludes some vivid expressions of the area. “I know of no region in the
tropics so hard to traverse . . . it is the saturated condition of the atmos-
phere and the persistent driving Scotch mist that makes life so desperately
miserably cold.

“Asa rule the wet clouds are down all the daytime and the clinging
fog and absence of sunlight combine to depress one unutterably.

“There is only one story of trees and that is about 20 feet high. The
trunks are grotesquely large in appearance, but this is due to a cloak of
moss which converts a 12 inch diameter which is about the maximum
over bark to 24 inches or more.

“What is more striking yet is the spring mattress effect of the ground.
The trees put out aerial roots, and these lie well above the soil, but are so
covered with moss as to deceive one at first, and when walked over the
whole network of roots gives to the foot a springy effect which is rather
weird. The species are few and are crabbed, gnarled, crooked and bent.
They are blanketted in moss, tufted with a thread-leafed orchid, and they
put out great elbows of aerial roots.”

The classic work of William H. Brown, Vegetation of Philippine Moun-
tains, subtitled “The relation between the environment and physical types
at different altitudes” contains a description of the northernmost mossy
forest at the top of Mt. Maquiling, Luzon, in the Philippines. Brown
found the mossy forest zone between 900 meters and the summit at 1,140
meters. His description parallels that for other mossy forests indicating
a forest of a single story, many individuals but few species, abundant mist
and reduced sunlight, and the overwhelming abundance of epiphytes.
Many comparisons with his biological studies will be possible with the
data presented in this and later papers concerning Pico del Oeste.

Australia
Ley, Webb (1959) reports the limited occurrence of a similar forma-
tion-type which he called a “microphyll mossy thicket.” Its characteristics
he noted were a level canopy with stunted uneven trees averaging between
6 and 15 meters in height, abundant epiphytes and the persistent mists.

Africa

There are few comparable reports for a mossy forest formation-type in
Africa. Richards quotes J. Lebrun (1936) in a paper I have not been
able to consult. Lebrun recognized three zones to a montane rain forest
with the forest of the upper zone having something of the character of an
elfin woodland. The trees did not exceed 10 to 15 meters in height and
their trunks and branches were twisted and irregular. Few species and
plants have compound leaves in this forest and the leaves in general are
small, coriaceous, and entire. A. W. Exell (1944) mentions a mist forest
region at 1,400 to 2,024 meters altitude on SAo Tomé in which the trees
are smaller than in lower zones and the epiphytes even more abundant.
Richards concludes that the dwarf mountain forest of East Africa seems

1968 | HOWARD, ELFIN FOREST, 1 387

in many respects comparable with the dwarf mossy type of montane rain
forest in Malaysia.

South America

Within continental tropical America the recent work of P. J. Grubb,
J. R. Lloyd, T. D. Pennington, and T. C. Whitmore (1963) concerns a
montane rain forest in Ecuador. The bibliography indicates the paucity
of detailed studies of mist- and mossy forests in this area. The authors
suggest a division of Richards’ classification of montane rain forest into
two units, a “lower montane rain forest” and an ‘“‘upper montane rain
forest.” The latter consists of one stratum of trees, 4.5 to 18 meters tall,
lacking buttresses, and with leaf size predominantly microphyllous, climb-
ers uncommon, vascular epiphytes common and non-vascular epiphytes
abundant. Although they did not study such a forest type they suggest
it “certainly occurs in the Andes, as is apparent from the general descrip-
tions of Weberbauer (1911) and Cuatrecasas (1958).”

The Lesser Antilles

John Beard as a resident forester in the Lesser Antilles during the years
of World War II visited most of the islands of the Antilles and published
a number of papers on the forest- and vegetation-types which culminated
in three outstanding works: Forestry in the Windward Islands (1944);
The Natural Vegetation of the Windward and Leeward Islands (1949);
and The Classification of Tropical American Vegetation-Types (1955).
Grubb and his co-workers find Beard’s proposed vegetation classification
“unsatisfactory because too great emphasis is placed on the criterion of
stratification while the criterion of leaf size is not used at all.”

Above the montane rain forest, Beard recognizes woody vegetation types
as “elfin woodland” and “montane thicket.” The elfin woodland according
to Beard is a single storied impenetrable tangle whose height varies from
10 meters down to 1 meter in which the trees are predominantly me-
sophyllous. Grubb and his associates note that elfin woodland is a term
which has been used extensively for forests of similar general aspect in
the Old World but that these are predominantly microphyllous. They
suggest that “depending on locality it may be best to regard the Elfin
woodland as a stunted facies of either the Lower or Upper Montane Rain
forest,” a classification they propose to indicate a more appropriate re-
classification of Richards’ montane rain forest. The same authors define
their “Lower Montane Rain forest” as consisting of two strata of trees
which may have buttresses uncommon or small, the Mesophyll leaf size
class, the canopy at 15 to 27 meters; the photophytic climbers uncom-
mon and the skiophytic climbers common, and both vascular and non-
vascular epiphytes abundant. Beard (1955) previously stated that the
terminology of “Elfin Woodland or Mossy Forest” is not precise for the
type of vegetation found on the mountain peaks in tropical America and
suggested “we should correctly speak of montane woodland and elfin
thicket; ‘elfin woodland’ is however an old established name.”

388 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Grubb and his co-authors objected to Richards’ classification “because
there is nothing particularly ‘Submontane’ about Lower Montane Rain
forest.” As no one has objected to the fact that mosses may be rare in
“mossy forests” where hepaticae abound or that “elves” have not been
seen recently in “elfin forests” both terms may still be applied as they
have been for years to the unusual, dwarfed, high altitude, mist shrouded,
vegetation type of the Caribbean.

Beard (1946) found the elfin formation in Trinidad to be represented
by a consociation of stilt-rooted trees, Clusia intertexta, only on the sum-
mit of El Aripo near 3,085 feet. In addition to this dominant tree there
were only six other dicotyledonous trees and a total woody flora of but
eleven species.

In Grenada the elfin formation is of “low, gnarled growth of small trees
and shrubs 4 to 20 feet high, windswept and festooned with moss.” The
principal species are Didymopanax attenuatum, Charianthus coccineus,
Ilex sideroxyloides, Byrsonima martinicensis, Ternstroemia peduncularis,
and Stylogyne lateriflora . Such areas however are few.

On St. Vincent the trees are from 8 to 12 feet tall on some of the higher
ridges and peaks, covered with moss and epiphytes. Stunted and gnarled
and windswept formations are comprised of Charianthus, Didymopanax,
Ilex, Ternstroemia along with Prestoea montana, Ficus, and Clusia species.

St. Lucia has an elfin woodland capping the principal pitons and this
is composed of Clusia venosa, various Araliaceae, and the same species of
Ternstroemia and Charianthus.

Stehlé (1937) wrote mostly of the vegetation of Guadeloupe where he
was employed. He indicates that at 1,000 to 1,484 meters and derived
from dense forest one encounters various associations which occur due to
the intensity of ecological factors. These he notes to be abundant rain,
accentuated slopes of the soil, rigorous temperature and past and present
volcanic action. The forest is transitional from dominance of Clusia
venosa and Cyrilla racemiflora to the semi-arboreal savannas of Lobelia
guadalupensis and Didymopanax attenuatum. On Martinique, outside the
influence of past volcanic action, he reports Clusia plukenetii plays a
similar role.

Dominica has been studied intensively by Hodge (1943, 1954). He
stated “The vegetation of the elfin forest is perhaps more interesting than
that of any other region of the island, but because of its inaccessibility
plant collections in the past have been few and incomplete. The flora is
distinct. Few of its members occur in other associations on the island;
more endemics and species of restricted range are found here. Near the
mountain summits the dwarf trees grow close together, forming a single
tight canopy; in fact, the crowns are so enmeshed that it seems almost
possible to walk across them.”

The dominant species of the mossy forest in Dominica is Clusia venosa
which can form pure thickets in places. Associated in the formation may
be Charianthus, Didymopanax, Ilex, Hibiscus tulipiflorus, Prestoea mon-

1968 | HOWARD, ELFIN FOREST, 1 389

tana, Freziera, and Endlicheria sericea. Vines are few but a thicket-form-
ing bamboo and a scandent Mikania are familiar.

Only the islands of St. Kitts and Nevis within the Leeward islands
have peaks of sufficient altitude to develop the cloud or mist cover neces-
sary for the development of a mossy forest. However, a fully developed
mossy forest is not found. A concurrent but less detailed study of St. Kitts
will be reported in a later paper in this series. A small amount of mossy
forest type of vegetation is found on the higher peaks of Mt. Misery,
Camp’s Mountain, and Verchild’s Mountain. Here the major components
are the woody genera Didymopanax, W einmannia, Coccoloba, Ilex, Chari-
anthus, Bumelia, Cyrilla, and Podocarpus. A comparable Aa is present
in restricted areas at the summit of the principal peak on Nevi

The Greater Antilles

The work of Forrest Shreve on A Montane Rain-forest in Jamaica
(1914) describes the forest vegetation of the central range. His studies of
forest type, soils, rainfall and cloud cover, transpiration, and light charac-
teristics are cited very frequently. Shreve noted that the montane rain
forest gave way to thickets at the summit of Blue Mountain Peak and
others. He reported on the frequency of the well-known temperate zone
genera, Vaccinium, Hypericum, Clethra, Lycopodium, and Viburnum which
in reality have Andean affinities. Throughout Jamaica as well as in
Cuba and Hispaniola the ‘‘mossy” forest, in contrast to the area being
studied in Puerto Rico, is more open and taller and where low the vege-
tation is less mossy and more frequently interrupted and discontinuous.
Asprey and Robbins (1953) in their survey of vegetation types in Ja-
maica regarded the elfin woodland at the exposed summits of the north-
ern ridges of the Blue Mountains at 5,000 feet and over as a faciation
of the montane mist forest. They noted that the branching was rambling
and distorted and frequently began close to the ground. The crowns
were sheared and sloping due to wind exposure and little emphasis was
made of the mossy characteristic. Asprey and Robbins agreed with
Shreve that rain forest was a misnomer and suggested the use of the
name montane mist forest since the vegetation in such areas is con-
trolled not by rain but by mist and clouds

Seifriz (1943) related that the summit of Pico Turquino in eastern
Cuba “is credited with an altitude of 6,580 ft. This is not high enough
for an alpine meadow in the tropics; its place on Turquino is taken by
an alpine thicket. Alpine thickets have been referred to but little owing
to their absence on temperate mountains and the paucity of ecological
studies in the tropics, but they are very characteristic of mountain tops
around the equator. . . The thicket which caps Turquino is a tall one,
the slender trees averaging 12 to 15 feet in height. The association is very
dense, a machete being necessary to cut one’s way through.”

Seifriz stated “Whether it is in New England, on the summit of Pan-
gerango in Java, or of Turquino in Cuba, the genera are the same: Vac-
cinium, Myrica, Viburnum, Smilax, Clematis, Rubus, . . . Hypericum,

390 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Vernonia, Pilea, Solanum, Eupatorium, Bidens and the sedge Rhyncho-
spora.” However he found “Associated with these familiar northern
genera on Turquino are many tropical ones, of which several are en-
demic. The most abundant tree is Cyrilla racemiflora . . . A typical
minute-leaved alpine thicket tree is Rapanea microphylla (Myrsinaceae).
But that family of trees which, more than any other, is certain to be
found high on tropical American mountains is the Melastomaceae, and
among them the genus Miconia predominates . . .

“Though the summit vegetation is a very heterogeneous mixture, several
of the trees in the Turquino alpine thicket are far more abundant than
others; one of these is Haenianthus salicifolius.” Seifriz gives a list of 81
species of trees, shrubs, vines, epiphytes, and terrestrial herbs and ferns
clearly indicating the heterogeneous nature of the alpine thicket in Cuba.

Ecological studies of Hispaniola are also few in number. The central
mountain range is dominated by Pinus occidentalis which does not be-
come involved in thicket formation. Even at higher altitudes the occur-
rence of tropical species of the Myrtaceae, Melastomataceae, Myrsin-
aceae and similar families is limited to moist valleys and they are not in
any formation which could be compared with a “mossy forest.” In the
Sierra de Bahoruco range a few peaks possess a mossy forest comparable
to that of the Blue Mountains of Jamaica where species of Weinmannia,
Hedyosmum, Miconia, Turpinia, Coccoloba, Vitex, Clidemia, Guettarda,
Brunellia and Gyrotaenia form the woody components of contorted thick-
ets and are laden with epiphytes both vascular and non-vascular.

The most detailed study of the mossy forest in Puerto Rico is that of
H. A. Gleason and M. T. Cook (1927) as part of their ecological survey
of the island. They recognized on the slopes of the Luquillo Mountains
a succession from a rain forest to a palm forest, to a mossy forest in cor-
relation with the increase in altitude. The area of El Yunque which they
visited is today part of an insular national park and is included in the
Luquillo Experimental Forest. Although visited by many people who
walk the easier trails the area has been little changed in the 40 years
which have passed and extracts from their descriptions remain pertinent.

“In the Luquillo Mountains, the habitat of the mossy forest probably
receives the highest rainfall and certainly has the highest average at-
mospheric humidity. Throughout most of the year, even in the dry sea-
sons, the peaks are wrapped in fog or cloud, raising the humidity nearly
or quite to the saturation point, causing rainfall almost every day, and
reducing the sunshine to short intervals. ... As a result of the high
precipitation and humidity, the soil of the mossy forest is generally wa-
ter-soaked and frequently quite boggy, and the vegetation is almost
constantly dripping with moisture. . . . the trail traverses a sharp ridge,
so narrow that one can look down a steep slope on both sides. Even here,
with every opportunity for rapid drainage, the soil is muddy and the pe-
destrian sinks deeply into it at every step. The soil is also usually thin
and in many places the forest floor consists merely of irregular rocks sep-

1968 | HOWARD, ELFIN FOREST, 1 391

arated by masses of wet muck — with fallen leaves, dead twigs,
superficial roots and prostrate stem

“The trade winds blow almost oe over the peaks of the Lu-
quillo Mountains. They are so moist that they probably do not affect
the water-relations of the vegetation seriously but they do have con-
siderable mechanical effect on the plants, so that over the most exposed
portions the vegetation is limited to a height of two to three meters
(6-9 feet), and the tops of the plants are shorn to a perfectly uniform
level

“There are, then, four environmental conditions which may be of con-
siderable importance in determining the peculiar vegetation of the mossy
forest, low temperature, great wind exposure, a water-soaked soil, and
an atmosphere of high humidity, frequent fog and mist, and heavy
precipitation. Of these, it seems probable that wind is the important
factor in differentiating the palm forest from the mossy forest, since the
former is regularly limited to the most sheltered sites and the latter to
the exposed ridges and peaks; wind also determines the general physio-
gnomy of the mossy forest and induces the development of dwarf trees
so characteristic of high altitudes. The other three conditions, but more
especially the nature of the soil and the humid atmosphere, seem pri-
marily concerned in preventing the development of the ila rain forest
and thereby encouraging the dominance of the mossy fore

“Four species constitute the bulk of the arborescent exin W ein-
mannia pinnata L., Ocotea spathulata Mez, Eugenia borinquensis Britton,
and Tabebuia rigida Urban, none of which occurs in the rain forest, ex-
cept possibly as rare and widely isolated individuals. ... these...
together constitute probably 75 per cent. of the dominant trees. Asso-
ciated with them, but of secondary importance are Ravenia Urbani
Engler, Calycogonium squamulosum Cogn., Miconia pycnoneura Urban,
Miconia pachyphylla Cogn., Miconia foveolata Cogn., Ceratostemma
portoricensis (Urban) Hoerold [Gonocalyx portoricensis|, Petesioides
yunquense (Urban) Britton [Wallenia yunquensis], Thibaudia Krugii
Urban and Hoerold [Hornemannia racemosa], Ilex Sintenisii (Urban)
Britton, Clusia Krugiana Urban [Clusia grisebachiana], Grammadenia
Sintenisii (Urban) Mez, Mecranium amygdalinum (Desr.) C. Wright,
and Cordia borinquensis Urban. . . . Marcgravia Sintenisii Urban and
Marcgravia rectiflora Tr. & Pl., both common in the palm forest and to
a lesser extent also in the rain forest, are also very abundant and con-
stitute a striking feature of the vegetation.’

At a lower elevation “. . . these species grow into trees eight to twelve
meters (25-40 feet) high. They differ from the trees of the rain forest
in aspect, having usually crooked trunks and more branches, and even at
this altitude the abundance of mosses, Selaginella, and epiphytes is strik-
ing. With increase in altitude and — with full exposure to the
wind, the stature of the trees is reduced.

“Toward their lower limits, the larger size and spreading branches of
the trees leads to their rather open spacing. At higher altitudes and with

392 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

decreased stature they are more closely crowded, until at the summit

. they form a dense thicket of crooked, gnarly, interlaced stems,
through which vision can seldom penetrate farther than three to five
meters (10-15 feet).

The trees are covered to a height of three or four meters (10-12
feet) with a thin layer of moss, while mosses carpet a considerable por-
tion of the forest floor. The density of this growth increases toward the

. until the ground is completely covered with dense mats of
Selaginella Krugii Hieron., covering the irregular surface of the ground,
hiding the rocks and creeping up the tree trunks to a height of a meter.
Above the Selaginella, the trunks of the trees are entirely covered with
a dense coat of several species of mosses and hepatics, in which the pen-
dent stems are 10-15 cm. long and give the trunk an apparent diameter
far greater than its actual size. This covering extends up the trees and
out to the smaller twigs, completely obscuring the bark and making all

differences between terrestrial and epiphytic habitats become negligible
and most species of secondary plants grow indiscriminately either on the
ground or on the side of the trees.”

In a more recent study of the area by Pierre Dansereau (1966) de-
scriptive material is supplemented by many tabular presentations includ-
ing pictorial charts.

Dansereau selected several sites in the Luquillo Mountains and his
categories, numbering 88 with sub-units, include for El Yunque: a sierra
broadleaf forest of Clusia minor, Calycogonium squamulosum, and Oco-
tea leucoxylon; a sierra palm forest of Euterpe globosa (Prestoea mon-
tana); a sierra moss forest of Tabebuia rigida, Podocarpus coriaceus, and
mosses; an elfin forest of Micropholis garciniifolia, Clusia krugiana,
Tabebuia haemantha, and Rapanea ferruginea; and a high montane scrub
of Eugenia, Tabebuia, Thibaudia krugii (Hornemannia racemosa), Ilex
sintenisii, Ocotea spathulata, and Icacorea luquillensis (Ardisia luquillen-
sis
Dansereau stated, “At the highest levels, below the outcropping and
isolated rocks, the palm drops out almost completely and so-called elfin
forest develops . . . This formation is less than eight meters high —
therefore a scrub by my present standard .. . Like the elfin forest —
or the elfin woodland of Beard (1944) — it may have a moss-line at mid-
height. The most sarmentose trees are twisted into a very complex net-
work.

“It is tempting to consider this montane vegetation as temperate in-
stead of tropical. The simplicity of its composition, the well-marked
dominance of a few species, the importance of mosses, all liken it to
woody formation in Madeira or New Zealand. Several of the genera are
widespread temperate rain-forest characteristics: Podocarpus, Ardisia,
Ocotea, Alsophila, Cyathea, Hemitelia, Myrica, Magnolia, Ilex, Sym-
plocos . . . Another feature is the aortonulatien of humus. One ai these

HOWARD, ELFIN FOREST, 1

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394 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

stands ... hada great depth (35 cm. or so) of fibrous organic matter,
a dry peat of sorts.”

The forest at the summit of Pico del Oeste showed many of the charac-
teristics included in the beautiful descriptions of Gleason and Cook, yet
detailed examination revealed unexpected information relating to the
peculiar forest form. I have difficulty applying Dansereau’s pictographs
and much of his information to Pico del Oeste. The several studies which
follow will, I believe, support some previously published ideas of the
cloud environment and will also offer alternate explanations of the form
and activity of the component species of the forest. Overall it is apparent
that the summit dwarfed forests of the Caribbean islands, being small in
extent and occurring on small land masses, are different from the mossy
forest, the cloud forests, the upper montane rain forests and similar for-
mations previously described in other parts of the world. Their charac-
teristics, however, are similar and the idea of a selection of species with-
in a microclimate is an appropriate one. This is indeed a special ecological
niche. The formation-type on Pico del Oeste does differ slightly from
that on the summit of El] Yunque as well as from those on islands to the
south. The material which follows is descriptive of one variation of the
elfin forest of Beard.

THE SUMMIT FOREST OF PICO DEL OESTE (West PEAK)

The area a for study on West Peak occurs along a ridge and
across the summit of the peak which is flattened into a small plateau,
perhaps an acre in extent (Fics. 1, 2, 3, 4). The shrubs and trees on
the easternmost, windswept face form a very dense tangle often less than
2 meters in height or in valleys or protected lees becoming 4 to 5 meters
tall (Fics. 5, 6). On the summit plateau the canopy is 2.5 to 4 meters
above ground level and on the lee side of the ridge rock slides form
cirques where Prestoea, Micropholis and Magnolia reach 8 to 10 meters
in height. Non-vascular cryptogams, ferns, and filmy ferns and Selaginella
species cover the forest floor throughout. Herbs are scattered individuals
or small seed groupings except in the palm glades where ferns and her-
baceous dicotyledons form solid stands of vegetation. The exposed roots,
the prop roots, the rhizomes and trailing or leaning branches form ridges
or cones or peculiarly shaped mounds where the mosses, liverworts and
selaginellas occur in thick layers as an epiphytic growth, The trunks
and branches are festooned with hanging liverworts and often exhibit
shiny masses of gelatinous colonies of algae. Vascular flowering plants
occur as epiphytes in this zone, and nearly every species of flowering
plant of the forest is found as an epiphyte at least in the immature stage.
Herbaceous and woody climbers such as [pomoea, Rajania, Hornemannia,
and Gonocalyx scramble and climb throughout the shrubs and often form
dominating masses as part of the canopy. Long trailing branches of Marc-
gravia make reticulate patterns on the forest floor. The young climbing

1968 | HOWARD, ELFIN FOREST, 1 395

&

. 6. A view eal the more Aa vegetation on the windward ae ie Pico
del Oc este. Dr. Howard is 6 feet 5 inches tall. Plants Tabebuia rigida, Ilex sin-
ite Micon packyphatla pare borinquensis, and Calyptranthes krugi

n be dis

396 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

shoots are flattened and have adpressed leaves in marked contrast to the

florescences at their apices. Wallenia yunquensis and Gesneria sintenisti
exhibit rhizomatous green growth, from which upright es develop
and form a sub-layer abundant in places and missing in othe

The branching of many of the taller shrubs is Rennie presenting
one characteristic appearance, while other species display a peculiar
growth pattern, horizontal or candelabra-like in nature due to the oc-
currence of terminal short shoots and lateral long and often repeated
shoots, during their development. Slender stem growth is exhibited by
Ilex sintenisii, Trichilia pallida, and Symplocos micrantha so that many
specimens appear to be scrambling and dependent for support on the
surrounding vegetation. The innumerable rosette-forming epiphytes of
the Bromeliaceae, Vriesia and Guzmania, occur in all size classes and ex-
hibit both red photophytic and green skiophytic growth forms. A cane-
like orchid, Dilomilis is nearly equally abundant, with long erect stems
subtended by pendent masses of whitish, green-tipped roots. Frequently,
due to insecure attachment or to top-heaviness, these plants are angled
or pendent from the normal upright position. Aérial roots are conspicuous
on many species as prop roots or adventitious branched or unbranched
growths from horizontal stems, or clustered when developed from broken
and injured stems. Especially conspicuous are the glistening gelatinous
sheaths on the aérial roots of several species

open areas along the trail where light reaches the forest floor the
grasses Ichnanthus and Isachne invade and form dense green mats. Scleria
secans, presumably another invading species, forms tangles of scrambling
branches soon noticed for the sharp marginal serrations on the leaves.
Clumps of Carex polystachya occur on the forest floor, and tangles of the
low creeping bamboo-like grass Arthrostylidium create barriers in other
areas. In moist open places along the trail rosettes of Eleocharis and
masses of Sphagnum are present.

A survey of the vegetation on Pico del Oeste revealed the species of
vascular plants in the following list. The taxa cited and preceded by an
asterisk are the common components of the mossy forest. The analyses
we have made of these will be presented in subsequent papers. The un-
marked species were of single occurrence or quite rare, excepting only
the ferns, which did not lend themselves to phenological studies or to
comparison in structural characteristics to the vascular flowering plants.

Lycopodiophyta
LYCOPODIACEAE
Lycopodium cernuum L. Terrestrial or erage plant. Locally dominant.
Lycopodium taxifolium Sw. Infrequent epiphyte
SELAGINELLACEAE
Selaginella krugii Hieron. Most common species with long naked rhizomes; in
masses or epiphytic and hanging in festoons.
Selaginella substipitata Spring. Occasional, erect, semi-rigid, terrestrial plants.

1968 | HOWARD, ELFIN FOREST, 1 397

Polypodiophyta
CYATHEACEAE
Cyathea borinqguena (Maxon) Domin. Rare, small terrestrial tree fern with
erect stem less than one foot tall.
Cyathea pubescens Mett. ex Kuhn. Most common small tree fern in elfin wood-
land and in palm glades. Terrestrial but young plants commonly epiphytic.

tip.
Lophosoria quadripinnata (Gmel.) C. Chr. Trunk > 2 feet tall, base of fronds
characteristically blue-black. Rare in palm gla

POLYPODIACEAE
Blechnum divergens (Kunze) Mett. Small, erect-stemmed plant of open for-
est, to 2 feet tall. Common.
Blechnum lineatum (Sw.) Hieron. Erect plant in more open areas. Abundant.
Blechnum underwoodianum (Broa C. Chr. Climbing or erect plant in open
areas forming stem to 4 feet long. Occasional.
Cochlidium minus (Jenm.) Maxon. Occasional epiphytic fern in forest area.
Cochlidium seminudum (Willd.) Maxon. Occasional epiphyte in forest are
Diplazium centripetale (Baker) Maxon. Small tree fern with trunk to 1 ne
tall, in palm glades
Diplasium grandifolium Sw. Occasional small tree fern in palm glade, trunk to
3 feet tall.
Diplazium Vherminieri Hieron. Small tree fern with trunk to 1 foot tall in palm
ades

Diplsiom ee (L.) Presl. Terrestrial plant rarely developing a short
rect stem
Pla piloaes ae (Mett.) Urban. Epiphytic plant occurring mostly in
juvenile form
Grammitis hessii (Maxon) Alain. Epiphytic, very small plant locally eigen
Grammitis serrulata (Sw.) Sw. Very small epiphyte, locally abundan

Lindsaea stricta (Sw.) Dryand. Upright terrestrial or epiphytic plant. Rare.

Oleandra articulata (Sw.) Presl. Infrequent terrestrial herb.

Peltapteris peltata (Sw.) Morton, Epiphyte, locally abundant but seen only on
trunks of Prestoea montana

Polybotrya cervina (L.) Kaulf. Terrestrial plant mostly in palm glades. In-
frequent.

Polypodium chnoodes Spreng. Epiphytic, sometimes climbing plant with pubes-
cent rhizome. Occasional.

Polypodium loriceum L. Epiphyte with characteristic green or silvery ascend-
ing rhizomes. Occasional.

Thelypteris decussata (L.) Proctor. Large terrestrial fern of palm glades, young

crozier characteristically surrounded by a gelatinous mass.

HYMENOPHYLLACEA

Hymenophyllum jostaie Sw. Abundant epiphyte.

Hymenophyllum macrothecum Fée. Abundant epiphyte

Trichomanes crispum L. Abundant as a terrestrial or epiphytic plant.

Trichomanes rigidum Se. Abundant plant, epiphytic or more commonly ter-
restrial.

398 JOURNAL OF THE ARNOLD ARBORETUM [ VoL, 49

Magnoliophyta — Liliatae
GRAMINEAE
Andropogon bicornis L. A aS clump of this adventive species was exterm-
inated by travel on the t
*Arthrostylidium sarmentosm Pilger. A trailing terrestrial plant with arching
stem; forest floor. Com
*Ichnanthus pallens (Sw.) ore. Terrestrial herb sometimes forming dense

stands.

*Jsachne angustifolia Nash. Scrambling terrestrial herb.

CYPERA

*Carex _pbsstacky Sw. ex Wahlenb. Rosette or clump plant within forest,
abun

“aera yunquensis Britton. Rosette clump plant in trail area. Common.
*Scleria secans (L.) Urban. Possible adventive into breaks, scrambling. Oc-
casional.

Fimbristylis dichotoma (L.) Vahl. Adventive in area cut over during road con-
struction.

*Prestoea montana (R. Grah.) Nichols. Tree of valleys, common.

ARACEAE

*Anthurium dominicense Schott. Epiphyte or terrestrial, herb, common.

BROMELIACEAE

*Guzmania berteroniana (R. & S.) Mez. Epiphyte, rosette, rare on research
site, common at lower altitudes

*Vriesia sintenisii (Baker) Smith & Pittend. Epiphyte, rosette, common.

DIOSCOREACE

*Rajania cates L. Herbaceous twiner, occasional.

ZINGIBERAC

AE
*Renealmia antillarum (R. & S.) Gagnep. Herbaceous, terrestrial, rare and per-
haps an invader

ORCHIDACEAE

*Brachionidium parvum Cogn. Epiphyte, rare.

*Dilomilis montana (Sw.) Summerhayes. Epiphyte or terrestrial, common.
Epidendrum nocturnum Jacq. Epiphyte, seen outside transect or trail areas.
Epidendrum pallidiflorum Hook. Epiphyte, seen outside transect and trail areas.

Magnoliophyta — Magnoliatae
PIPERACEAE
a emarginella (Sw.) DC. Delicate, herbaceous, root-climbing epiphyte,
mmon
*Peperomia sap cg (Vahl) A. Dietr. Succulent herbaceous root-climb-
ing epiphyte, comm
CHLORANTHACEAE
*Hedyosmum arborescens Sw. Small shrub never reaching canopy, occasional.
MoraceEA

E
*Cecropia peltata L. Tree of disturbed areas; invader of trail.

1968 | HOWARD, ELFIN FOREST, 1 399

URTICACEAE

*Pilea krugii Urban. Terrestrial herb, common.

Pilea obtusata Liebm. Terrestrial herb in palm glades, co

*Pilea yunquensis (Urb.) Britt. & Wils. Terrestrial or fe 2h epiphytic herb,
common.

LAURA

*Ocotea spathulata Mez. Dominant tree in canopy.

MAGNOLIACEA

Magnolia splendens Urban. Common tree of lower elevations, single specimen
found in transect area on lee side.

CUNONIACEAE

Weinmannia pinnata L. Occasional tree in dwarfed forests on El Yunque, single
specimen found on Pico del Oeste but not in transect or trail area.

MELIACEAE

*Trichilia pallida Sw. Very slender scrambling tree on Pico del Oeste, infrequent.

AQUIFOLIACEAE

‘Ten pene (Urb.) Britton. Scrambling shrub reaching canopy; occasional.

CELASTRACEAE

*Torralbasia cuneifolia (C. Wr. ex Griseb.) Krug & Urban ex Segui. Slender
upright shrub with branches reaching canopy in places; occasional.

OCHNACEAE

*Sauvagesia erecta L. Herbaceous terrestrial, early invader of trail area; com-
mon roadside weed at lower elevations.

MARCGRAVIACEAE

*Marcgravia sintenisii Urban. Abundant heterophyllous woody root-climber ter-
restrial and reaching canopy.

THEACEAE

*Cleyera albopunctata (Griseb.) Krug & Urban. Rare as a forest tree, oc-
casional as slender shrub with scrambling branches.

GuTtT ;

*Clusia iinet (Planchon & Triana) Alain. Occasional stalk-rooted
scramblin

Clusia pie Ta Stahl. Rare plant in the forest not encountered in transects.

BEGONIACEAE

*Begonia decandra Pav. ex A. DC. Herb of forest floor; locally abundant.

THYMELAEACEAE

Daphnopsis philippiana Krug & Urban. Rare shrub not encountered in transects;
abundant at slightly lower elevations.

YRTACEAE
*Calyptranthes krugii Kiaersk. Abundant shrub occasionally reaching canopy.
Eugenia borinquensis Britton. Common tree of canopy.

MELASTOMATACEAE
*Calycogonium squamulosum Cogn. Common tree of canopy.
neat amygdalinum (Desr.) C. Wright ex Sauv. Oana shrub, never

can
*Miconia Tee Cogn. Common shrub, rarely in canopy.

400 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

stone pachyphylla Cogn. Common shrub, occasionally in canopy.

*Miconia pycnoneura Urban. Infrequent shrub, never in canopy.

er nities sintenisii Cogn. Common shrub at oer elevation, not encountered in
transects.

ERICA

Sono chin peed! (Urb.) A. C. Sm. Woody root-climber forming masses
in canopy,

<E craeimains racemosa Vahl. Woody climber forming masses in canopy; com-

Myrsti
“Aisi sieaitivete (Britton) Alain. Tree forming portion of canopy, occasional.
‘times sintenisii (Urb.) Mez. Slender shrub, never reaching canopy;

*Wallenia yunquensis (Urb.) Mez. Low shrub, rhizomatous; abundant.

SAPOTACEAE

*Micropholis garciniaefolia Pierre. Tree forming a portion of canopy, 0c-
casional.

SYMPLOCACEAE

*Symplocos micrantha Krug & Urban. Slender shrub usually scrambling or with
scrambling branches; rare.

LEACEAE

*Haenianthus salicifolius ii var. obovatus (Krug & Urb.) Knobl. Tree form-
ing portion of canopy; r

CONVOLVULACE

*I pomoea hati Jacq. Herbaceous twiner; rare.

BORAGINACEAE

— borinquensis Urban. Found only outside the transect area as a small
ree.

BIGNONIACEAE

*Tabebuia rigida Urban. Dominant tree.

GESNERIACEAE

*Alloplectus ambiguus Urban. Epiphytic or terrestrial fleshy herb; occasional.
esneria sintenisii Urban. Terrestrial wandlike shrub, never reaching canopy;
occasional.

ACANTHACEAE

Justicia martinsoniana Howard.’ Occasional herb of forest floor.

‘Justicia martinsoniana Howard, nom. . Fic.
Rhytiglossa verticillaris Nees in DC. P a 354. 1847; type Plée a ‘®)
Justicia verticillaris (Nees) Urb. Sym % ae ill. 4: 579. 1911, non L.

Ste oh hes verticillaris (Nees) Britton, Sci. Surv. Porto Rico Virgin tr. E: aie

When Urban transferred the species described ap Nees to the genus Justicia he
made note of the existence of the identical name used by Linnaeus filius for a differ-
ent plant. ew name is required unless one maintains the species in the poorly de-
fined genus Stethoma, as Britton did.

It is a pleasure to name this attractive though small-flowered endemic herb for
Joseph B. Martinson whose codperation and support through the duration of this
project made our work possible and our life in Puerto Rico more enjoyable.

1968 | HOWARD, ELFIN FOREST, 1

Fic. 7, a-d. Justicia martinsoniana drawn from fresh material; a, habit,
x Ys: b, flower, side view, X 5; c, flower, face view, X 5; d, stamen, x 15.

RUBIACEAE

*Hillia parasitica Jacq. Epiphytic shrub; occasional immature specimens only.

*Psychotria berteriana DC. Shrub, possible adventive in breaks or landslide

areas. :

*Psychotria guadalupensis (DC.) Howard. Small semi-succulent epiphyte, com-
mon.

Rondeletia portoricensis Krug & Urban, Occasional at lower elevations; seen
only as seedling in transect area.

CAMPANULACE

*Lobelia horton eee (Vatke) Urban. Herb of forest floor, occasional in opea
areas.

402 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Fic. 8. Map of the study trail on Pico del Oeste.

COMPOSITAE
Clibadium erosum (Sw.) DC. Shrub of lower elevations; juvenile plants only

s adventive weeds of trail.
*Mikania pachyphylla Urban. Herbaceous twiner; occasional.

The composition of the vegetation on the top of Pico del Oeste is un-
usual for the complete lack of Cyrilla racemiflora found commonly at
lower elevations. The absence of members of the families of Leguminosae,
Rosaceae, Solanaceae, and Verbenaceae also is worthy of note.

1968 | HOWARD, ELFIN FOREST, 1 403

The dominant flowering plants selected for further study represent eight
families of 14 species of Monocotyledoneae comprising one palm, seven

ilies. There are 27 families of Dicotyledoneae in the study selection
represented by 40 species. Twenty-four species are trees or shrubs;
twelve herbs or herbaceous vines; four woody climbers. Four species oc-
cur most commonly as epiphytes. The most common family is the Me-
lastomataceae with three genera and five species, with the genus Miconia
having three species.

GEOGRAPHIC DISTRIBUTION OF THE VASCULAR FLORA

Many patterns of distribution have been discerned within the flora of
the Antilles. The vascular flora of Pico del Oeste consists of 35 principal
species which were considered in this study, 31 species of ferns, 14 species
of Monocotyledoneae and 40 species of Dicotyledoneae. Of these, 27 or
32% are endemic to Puerto Rico; 17 or 20% are restricted to the Greater
Antilles in various patterns of distribution; 17 or 20% are restricted to
the Greater Antilles and the Lesser Antilles in various patterns of dis-
tribution; 17 or 20% are of widespread distribution in tropical America,
occurring in the Greater Antilles, the Lesser Antilles, Mexico and Central
America, and in South America; 4 or nearly 5% are found in the An-
tilles and northern South America but do not occur in Central America ;
2 or roughly 2% occur in the Greater Antilles, Central and South America
but do not occur in the Lesser Antilles; and one species has a distribution
of the Greater and Lesser Antilles and Central America but has not been
reported from South America.

Within the systematic classification of the component taxa 55 7%o of
the Dicotyledoneae are endemic to Puerto Rico and 87% are restricted
to the Greater and Lesser Antilles; 14% of the Monocotyledoneae are
endemic to Puerto Rico and 79% restricted to the Antilles, while in the
ferns only 9% are endemic to Puerto Rico; 497% are restricted to the
Antilles but 38% are known to have a wide distribution in tropical

The distribution of species within Puerto Rico remains an unworked
problem. Britton and Wilson, in their reports on the vascular plants of
Puerto Rico, indicated a very high percentage of endemic species (18%)
are found in the Luquillo Mountains. A large number of these are now
known to have wider distribution in Puerto Rico through the extensive
but unpublished and unreported collection records of resident botanists,
while many other taxa have been reduced to synonymy by monographers.

Within the various growth forms of the component species 61% of the
epiphytes are restricted to the Antilles but only 5% are endemic to Puerto
Rico. All six of the climbing species are restricted to the Antilles. Of
the 58 terrestrial species 39% are endemic to Puerto Rico and 69% are
restricted in their distribution to the Antilles.

404 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Fic. 9. Profile of the study trail on Pico del Oeste.

THE TRAIL

Once the physical site for the research problem was chosen, a marked
trail was laid out so that reference points could be established against
which all information could be correlated. A minimum of disturbance in
establishing the trail was necessary because it followed the vague outline
of an old abandoned trail which led to Pico del Este. The trail is roughly
in the shape of an inverted Y with a much abbreviated base. The juncture
of trails was designated as point 0 and it was at this point that a tower
to hold weather instrumentation was later erected. A small observation
structure was constructed at the end of the base of the Y (and at the
summit of Pico del Oeste). The two large trails were marked (Fics. 8,
9) at ten yard intervals with numbering beginning at O and running to
160 yards on the one arm and 150 yards on the other (the latter is desig-
nated as the A trail and reference numbers all bear an A). The two arms
face roughly in a northeasterly direction which proved to be of con-
siderable significance in regard to general wind direction. As primary

along the trail could be considered as the first wind contacts on Puerto
Rico. An additional advantage was that windward and lee positions
could be found at many places along the trail. The trail was marked by
two-by-four inch X 2 foot posts driven every ten yards with code num-
bers attached. The upper part of each post was painted bright red for
ease of locating.

Along the trail individual specimens of each species were marked and
a variety of labels were tested to determine ease of application, durability
of label, and durability of the legend applied. The most satisfactory la-
bel proved to be a wooden one of the type also used at the Arnold Arbore-
tum (Arnoldia 24: 9-12. 1964) made of pine strip 114” « 4” « 3%” which
had been painted with two coats of white enamel. The name of the
plant (or pertinent data) was embossed on a plastic tape and through its
adhesive affixed to the wood. A copper wire was used to fasten the label
to the plant. The paint and the plastic proved to be resistant to the ef-
fects of weather, the algae, and spores of hepaticae. The tags were placed
on the various species at the beginning of the trail as well as on particu-
lar plants selected for long-term observation along the trail and in the
canopy of the forest surrounding the observation hut. This method of
labeling assured a continuity of observation by the personnel who manned
the project and permitted a rapid mastery of the unfamiliar species by
visitors to the site. In retrospect, we believe that this labeling of plants

Distribution of Species

DISTRIBUTION Monocots Dicots FERNS EPIPHYTES TERRESTRIAL

M D F M D F
Endemic to Puerto Rico 2 22 3 1 19 2
Greater Antilles 3 1 6 3 1 3
Greater Antilles &

Lesser Antilles 2 4 4 2 2 2
Puerto Rico & Guadeloupe 7 1 1
Puerto Rico to Martinique 1
Lesser Antilles to

uerto Rico 1 1 1
Greater Antilles except

Jamaica 1 3 3
Puerto Rico & Cuba 3 3
Antilles, N. South America -

but not Central America 3 1 1 1
Jamaica, Puerto Rico &

Lesser Antilles 1 1
Antilles, Central &

outh America 3 2 12 5 2 7
Greater Antilles, Central &
outh America, not Lesser

Antilles 2 2
Antilles & Central America

but not South America 1 1

14 40 ol 13 32 18

[S961

I ‘LSauOd NIATA ‘AUYVMOH

SOP

406 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

of markers on the West Peak (Pico del Oeste) trail. The vertical lines indicate
the location of the trail itself. The left half of the transect at marker 4, and the
right half of transects at markers 7, 9, 8A and 12A progress along the steep bank
of a land slide zone.

1968 | HOWARD, ELFIN FOREST, 1 407

and species contributed materially to the success of the project. Instruc-
tions could be given to replacement observers for the location of specific
plants between trail markers on a particular side of the trail. Although
several types of labels are made of material on which a stylus or pencil
can make an impression these proved to be less durable and certainly less
satisfactory.

COMPOSITION STUDIES

The vegetation at the sides of the old trail used as a base line had been
affected by the former use of the trail, and in attempts to record the
frequency of component species it was soon apparent that a false im-
pression was being obtained. For that reason transect lines 50 feet long
were established at right angles and in both directions from each of the
10-yard markers along the base trail. Even this short distance, involving
the study of an area 3 feet on each side of the line, proved to be a phys-
ically difficult and exhausting survey (Fic. 10). The plants within 63
such areas were counted as accurately as possible and at the same time
a survey was made to obtain the elevations of each line as well as the base
trail, and thus construct the profile diagram of the hill top (Fic. 9). The
lines were irl ee) leeward and windward plots of the vegetation of
Pico del Oes

The oe of the vegetation in many of the areas was due (1) to
fallen trees from which lateral branches had grown upright or adventi-
tious shoots had developed; (2) to the rhizomatous habit of some of the
undershrubs, where a single rooted plant might produce six to twelve
upright shoots with or without secondary rooting; and (3) to the tangle
of vines which crossed and recrossed the study area. In all cases we at-
tempted to record only individual plants clearly rooted within the study
area, hence even the large figures given in the composition studies do
not reveal accurately the tangle and thickets that characterize the vege-
tation. The very many young plants of Vriesia sintenisii also offered a
minor problem. The wind-blown seeds of this epiphytic bromeliad found
Suitable media for germination in the heavy layers of liverworts which
formed on all trunks, branches, and on the ground. The number of ma-
ture flowering specimens of Vriesia suggested a very high mortality of the
seedlings, hence only plants 4” or more in diameter were included in this
Survey. In one estimate we concluded these represented less than 10%
of the total number of plants of that particular species on a single isolated
tree. For most of the woody species we tried to record separately the
seedlings to which the cotyledons were still attached. Another problem
was encountered in Marcgravia sintenisii, which formed a tangle of stems
on the ground, epiphytically on the trunks of many trees, and in the
canopy. Lateral branches of this woody climber could have either the
juvenile foliage form or the mature foliage or a successive combination
of both foliage types when the stems changed angles or were attached
by adventitious roots to other upright stems. The figures given repre-
sent individual plants as accurately as possible.

408 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

The ten most common species represented in these transect studies
were the following, in order of frequency. The second figure given rep-
resents the number of seedling plants.

1. Pilea krugii 1376 Terrestrial herb
2. Wallenia a 1120-40 Semi-woody, terrestrial
3. Calycogoniu
quamulosu 1063-224 Woody, canopy plant
4. Vriesia sintenisii 978 Epiphyte
5. Ocotea spathulata 904-297 Woody, canopy ae
6. Calyptranthes krugii 807-140 Woody, canopy plan
7. Pilea obtusata Herb restricted to nee glades
8. Dilomilis montana BY Epiphyte
9. Miconia pachyphylla 433-25 Woody, canopy plant
10. Cyathea pubescens 8 Woody, sub-canopy plant

The total distribution is the following:

HERBACEOUS
Pilea krugii 1376 Ichnanthus pallens 110
Pilea obtusata 731 Arthrostylidium sarmentosum 32
Pilea yunquensis 297 Scleria secans 22
Begonia decandra 256 Renealmia antillarum 5
Carex polystachya 168 Justicia martinsoniana 1

EPIPHYTES
Vriesia sintenisii 978 Alloplectus ambiguus ll
Dilomilis montana 717 Peperomia emarginella 10
Psychotria guadalupensis 36-2 Peperomia hernandiifolia 10
Anthurium dominicense 13 Hillia parasitica 9-5
Guzmania berteroniana 11

Woopy EMERGENTS
aia ee Picasa 1063-224 Ardisia luquillensis 40
Ocotea spath 904-297 Clusia grisebachiana 38-5
Colyhivaathees pea! 807-140 Miconia pycnoneura aa
Miconia pachyphylla 433-25 Prestoea montan 23-192
Tabebuia rigida 294-2 as kee garciniae folia 24
Eugenia borinquensis 185-387 Grammadenia sintenisii 3
Torralbasia cuneifolia 104 Symplocos eesti 19-3
Ilex sintenisi 98 Cleyera albopunctata 11
Miconia foveolata 91-37 Haenianthus salicifolius 4
Trichilia palli 59 Magnolia splendens 1
Hedyosmum arborescens 58-1 Rondeletia portoricensis 1
Mecranium amygdalinum 46-34 Miconia pycnoneura 1
UNDERSHRUBS

Wallenia yunquensis 1120-40 = Diplazium os 48
Cyathea pubescens 386 Psychotria berteriana 35-2
Gesneria sintenisii 163-26 Lobelia pa al agi 12

1968 | HOWARD, ELFIN FOREST, 1 409

CLIMBERS
Marcgravia sintenisii 154-117 Mikania pachyphylla 73
Ipomoea repanda 82 Rajania cordata 21
Gonocalyx portoricensis 82 Hornemannia racemosa 19

The total area represented by these 63 transects was 18,900 square
feet, approximately 0.4 acres or 0.17 hectares. Within this area there
were by count 11,328 mature or adult plants and 1,536 juvenile plants
or seedlings, for a conservative total of 12,864 individuals.

The base trail is curved and the transects were run on both the wind-
ward and leeward sides of a ridge as well as on the flatter summit area.
Lines from trail markers 13, 14, and 15 on the leeward side dropped very
sharply and extended across a slight talus slope where Prestoea montana
was the dominant plant. It was only in this area that Diplazium grandi-
folium and Pilea obtusata grew. The palms were all mature plants or
individuals over 5 feet tall. Seedling palms all less than a foot tall were
encountered in the windward transects but none of the plants appeared
to be vigorous. A total of 57 taxa as recorded in the 63 transects, with
a high frequency of 28, a low of 17, and an average of 21 taxa on the
windward plots, while a high of 27 taxa, a low of 15, and an average of
21 were found in the leeward plots.

A second transect study area was selected on the small plateau (lee-
ward side and between markers 3 and 2) near the summit of Pico del
Oeste (Fics. 11, 12). The site was chosen to be as comparable as pos-
sible to the area on El Yunque diagrammed by Dansereau.

The area selected was 10 yards square. Lines a yard apart were laid
through the forest to form a grid. Each square yard was then mapped
to show the origin of the plants and their place of rooting. The path of
each stem was marked, as was the area covered by the crown of each
plant. The transect of 900 square feet contained 1,419 mature plants
and 153 juvenile specimens for a total of 1,670 individuals with 35 taxa.
On a comparative basis with the line transects, the number of individual
plants was 2.7 times as many in this plateau study plot. The ten most
abundant plants were the following:

1. Dilomilis montana 332 Epiphytic herb
2. Vriesia sintenisii 329 Epiphytic herb
3. Wallenia yunquensis 187 Semi-woody terrestrial shrub
4. Ocotea spathulata 106+76 Woody plant of canopy
5. Calyptranthes krugii 75+6 Woody plant of canopy
6. Calycogoniu

Squamulosum 74+19 Woody plant of canopy
7. Carex polystachya $3 Terrestrial herb
8. Gonocalyx portoricensis 51 Woody liana
9. Ipomoea repanda 4112 Leon gag!
10. Marcgravia cane 38 Woody lia

Fic. 11. A representation of the “sal aera of the vegetation a0 the
study trail between markers 2 and 3 pe

sion lines represent one foot, i.e. the
highest tree, an Ocotea, near no, 12 2 ie 12) is 9 feet tall and og dia agram
(Fic. 11 and Fic. 12 pre 5 ‘e e€ linear transect. Component species
represented are: 1, Miconia pachyphylla; 2 Vriesia Sintenisii: 3 rece via sin-

tenisii; 4, Ocotea iy ernie! 5, Dilomilis montana: 6. Ra jani ordata

vf
Calyptranthe S krugii; 8, Cyathea ibescous: 9, Carex poliaieekye: 10, Bieows
borinquensis.

‘trail e i.e.
study trail between marker, ti d 3. Dimension lines aed on

the highest tree, an ar ai 12 is 9 "Tet tall pine each ran
: 5 ct. Com
resents 15 "i f linear transe
pond eee , Tabebuia Bas pp 12, Wallenia Pi 13, lex

tenisii.

412 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Woopy COMPONENTS OF CANOPY

Ocotea spathulata 106+76 Tabebuia rigida 21
Calycogonium squamulosum 75+19 Ardisia luquillensis 21
Calyptranthes krugii 75+6 Ilex sintenisii 6
Eugenia borinquensis 21+9 Hedyosmum arborescens 4
CLIMBERS AND SCRAMBLERS, MOSTLY IN CANOPY
Gonocalyx portoricensis 51 Mikania pachyphylla 32+2
Ipomoea repanda 41+2 Rajania cordata 2
Marcgravia sintenisii 8 Hornemannia racemosa 2
EPIPHYTES
Dilomilis montana 332 Peperomia emarginella 1
Vriesia sintenisii 187 Anthurium dominicense 1
plus numerous juveniles) Alloplectus ambiguus 1
Psychotria guadalupensis 5
TERRESTRIALS
Wallenia yunquensis 187 Miconia foveolata 4
Carex polystachya 53 Miconia pycnoneura 3
Blech 42 Clusia (seedlings) 3
Hymenophyllum Prestoea (seedlings) 2
macrothecum 32 Pilea yunquensis 1
Miconia pachyphylla 26+4 Symplocos micrantha
Cyathea pubescens 10+16 (seedlin 1
Trichilia pallida 8

g
iscellaneous juvenile
Pilea krugii 5 ferns
A SINGLE LINE TRANSECT OR A PROFILE THROUGH THIS SAME PLOT

WOULD SHOW THE FOLLOWING COMPOSITION
Vriesia sintenisii 15 Cyathea pubescens
Ocotea spathulata 11
Calyptranthes krugii
Wallenia yunquensis

a

t=]

—

i=

3

~

ue
a.
i) .
S

a ae a ee

Tabebuia rigida Ilex sintenisii

Dansereau’s pictogram showed 24 individuals representing 12 taxa.
The profile (Fics. 11, 12) constructed from this line through the plateau
plot contains 60 individuals representing 14 taxa.

The roof of the shelter constructed on the summit of Pico del Oeste
permitted observations of the canopy from above. It was obvious that
the vegetation as seen from above presented a different aspect from that
represented by the numerical surveys or by the profile diagrams (Fic.
13). Herbs and undershrubs and some of the epiphytes were not seen in
the canopy, so consideration was given to methods of mapping the forest
from above. I am grateful to friends and pilots from the U.S. Nav
based at Roosevelt Roads for making several attempts to take aérial
photographs of the study area on Pico del Oeste. Their photographs, al-
though not of quality for species analysis, did allow additional studies

HOWARD, ELFIN FOREST, 1

An aérial view of the study site on

s and a win rection indic ator.
visible and were used ee correlate ground and canopy studies of the —
Photo. reproduced from a color Sepaie couiteny of the U.S. Navy

of vegetation on the peak by suggesting areas worthy of further study
(Fic. 13). Pvcauualy we resorted to the procedure of attaching the
camera to a long pole and extending this from the vertical axis of a ladder
fastened to the side of the shelter, from the instrument tower built by
the staff of NCAR, and from a ladder braced upright in the middle of the

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416 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

10-yard-square transect study at markers 3-2. Photographs were taken
in Ektachrome and monochrome film, with a long cable release used to
make the exposure. The patience of my son and daughter, Bruce and
Barbara, in assisting in this study are again acknowledged, for the com-
bination of bad weather, low clouds or mist, and sporadic winds made
this project an extremely exhausting physical experience.

The photographs were used to compare and to verify the mapping
done from the ground by looking skyward from each of the yard-square
quadrats. A diagrammatic map of the area as revealed by these studies is
reproduced as Ficure 14.

The following species formed the canopy and represented the per-
centages given here.

NO. OF PLANTS

IN CANOPY % OF CANOPY
Tabebuia rigida 16 40.7%
Ocotea spathulata 46 30 %
Calyptranthes krugii 26 10.7%
Calycogonium squamulosum is %
Marcgravia sintenisii 7 4.2%
Eugenia borinquensis 7 4.2%
lex sintenisii 2 AG
Ardisia luquillensis z
Gonocalyx portoricensis -
Hedyosmum arborescens 1 3.9%
Mikania pachyphylla 1
Psychotria guadalupensis 1

In comparison with the numerical count of specimens within the same
plot, it is interesting to note that Tabebuia, which would have ranked 16th
in frequency, and Ocotea spathulata which ranked 4th, dominated the
canopy. As viewed from above 46 specimens were seen of the 106 Ocotea
plants counted, while 16 of the 21 Tabebuia plants reached the canopy.

CHANGES IN THE SITE DURING THE STUDY

Although as little disturbance of the vegetation as possible was the
working objective of all personnel during the study of Pico del Oeste, some
changes did take place. Nearly every participant in the study learned
that plants of Cyathea pubescens were shallowly rooted and could not

1968 } HOWARD, ELFIN FOREST, 1 417

be depended upon for support in climbing or descending some of the
transect slopes. Many of these plants were uprooted accidentally.
During the initial visit to the study area one clump of Andropogon
bicornis L. was seen on the trail and a specimen collected from the clump
for identification. Subsequently that single clump was trampled into the

tered by repeated crossings of the trail, and the moss also disappeared.

In an open area of trail where it was most narrow and the sides steepest
(13), seedlings of Sauvagesia erecta appeared within the first year. This
pan-tropical weed is common on the roadsides at lower elevations and it
it probable that staff members brought in seeds in the mud on their
shoes. The plants proved to be perennials which persisted in growth
and increased considerably in number during the course of the study.
After a period of two years seedlings of Clibadium erosum were noticed
in the same area. This species, too, occurs along the roadside and was
presumably introduced by personnel, on muddy shoes.

At the end of the trail and outside of the study area plants of Fim-
bristylis dichotoma (L.) Vahl appeared in a disturbed area. These do
not appear to have been introduced by project personnel.

No major change in the number of specimens along the trail was seen
in any species except Eleocharis yunquensis Britton. The colonies original-
ly consisted of widely spaced clumps. Near the end of the study the
species formed solid mats in areas where the foot traffic was heavy. The
plant appears to flourish under adverse conditions of disturbance.

LITERATURE CITED

Asprey, G. F., & R. G. Rospins. The Vegetation of Jamaica. Ecol. Monogr.
23: 359-412. 1953.

Bearp, J. S. Forestry in the Windward Islands. Development and Welfare W.
Indies Bull. 11. 183 pp. Trinidad. 1944.

———.. The Natural Vegetation of Trinidad. Oxford Forestry Mem. 20. 152
pp. + map. Clarendon Press, Oxford. 1946.

. The Natural Vegetation of the Windward and Leeward Islands. Oxford
Forestry Mem. 21. 192 pp. Clarendon Press, Oxford.

—.. The Classification of Tropical American Vegetation-Types. Ecology
36: 89-100. 1955.

Britton, N. L., & P. Witson. Descriptive Flora. Sci. Surv. Porto Rico Virgin
Is. ce 1-626. 1923-1924; 6: 1-663. 1925-1930.

Brown, W. H. Vegetation of Philippine Mountains. Bureau of Sci. Publ. 13.
434 pp. + 41 pls. Manila.

DANSEREAU, P. Studies on the vegetation of Puerto Rico. 1. Description and
integration of the plant-communities. Inst. Caribbean Sci. Spec. Publ. 1

pp. 1-45. Mayagiiez.

EXELL, A. W. Catalogue of the Vascular Plants of S. Tomé. 428 pp. British
Museum, London.

Gieason, H. A., & M. T. Coox. Plant Ecology of hae one Pts. 1 and 2.
Sci: Surv. Porto Rico Virgin Is. 7: 1-173 + 50 pls. 19

418 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Gruss, P. J., J. R. Luoyp, T. D. PennincTon, & T. C. WHITMorRE. A com-
parison of montane and lowland rain forest in Ecuador. I. The forest
structure, physiognomy and floristics. Jour. Ecol. 51: 567-601. 1963.

Hopce, W. H. The Vegetation of Dominica. Geogr. Rev. 33: 349-375. 1943.

. Flora of Dominica, B.W.I. Pt. 1. Lloydia 17: 1-238.

LANE- Poote, C. E. The Forests of Papua and New Guinea. Empire Forestry
Jour. 4: 206-234. 1925.

Lesrun, J. Répartition de la forét équatoriale et des formations végétales
limitrophes. Brussels. 1936.

Ricuarps, P. W. The Tropical Rain Forest. 450 pp. Cambridge Univ. Press.
1964 (second photocopy of 1952 edition).

Serrriz, W. The altitudinal distribution of _ on Mt. Gedeh, Java. Bull.
Torrey Bot. Club 50: 283-306. pls. 15-17. 1923.

. The Plant Life of Cuba. Ecol. Monogr. 13: 375-426. 1943.

SHREVE, F. A Montane Rain-forest. Carnegie Inst. Publ. 199. 110 pp. Wash-

in
STEENIS, C. G. G. J. vAN. On the origin of the Malaysian mountain flora. Pt. 2.
Altitudinal zones, general considerations and renewed statement of the
problem. Bull. Fon ard. Bot. Buitenzorg. III. 13: 289-417. 1935.
STEHLE, H. Les associations végétales de la Guadeloupe et leur intérét dans
la Vitothiatioit baie Revue Bot. Appl. Agr. Trop. 187. pp. 188-195.
1937.

Wess, L. J. physiognomic classification of Australian Rain forests. Jour.
Ecol. 47: — 570. pls. 14-17. 1959.

ARNOLD ARBORETUM
Harvarp UNIVERSITY

1968 | BAYNTON, ELFIN FOREST, 2 419

THE ECOLOGY OF AN ELFIN FOREST IN PUERTO RICO,
2. THE MICROCLIMATE OF PICO DEL OESTE

Harotp W. BAayNnTon

MOUNTAINS PENETRATE the base of trade-wind cumulus on many Carib-
bean Islands. Topping these mountains is a mossy, bushlike, elfin forest.
One such forest is situated on 1050-meter Pico del Oeste in the Luquillo
Mountains of eastern Puerto Rico. This forest has recently been the
subject of a joint ecological investigation by the Arnold Arboretum and
the National Center for Atmospheric Research (NCAR). The objective
was to determine the environmental factors responsible for the dwarfed
nature of the forest.

One year of meteorological data collected on Pico del Oeste is sum-
marized in this report. In the tropics, as elsewhere, the variability of
weather is such that one year does not define a climate. Nevertheless, the
microclimate associated with the cloud forest is sufficiently unique that
even a single year’s data suggest many characteristic weather features.
Moreover, more extensive climatologies of Puerto Rico (Briscoe, 1966;
Pic6, 1950; Wadsworth, 1948) provide a useful standard for evaluating
the representativeness of the year reported herein.

Pico del Oeste is at the eastern (i.e. windward) end of Puerto Rico. It
is far enough inland to be affected by convective showers produced by
the sun’s heating of the ground, yet near enough to the coast to receive
the full influence of the moist tropical trade winds. Rising to 1050 meters,
Pico del Oeste naturally owes much of its rain to the upslope effect. The
Glossary of Meteorology (1959) specifies 600 to 750 meters as the char-
acteristic base of trade-wind cumulus. It is not surprising that Pico del
Oeste is shrouded in cloud most of the time.

Late in September of 1965 a meteorological survey trip was made to
Pico del Oeste to aid in determining the variables that should be measured
and the instruments that should be used. During February, 1966, the
first of the equipment was installed and routine collection of data began.
Wind sensors were added during the summer of 1966. Within a year
corrosion and electrical failure of equipment had become a chronic prob-
lem and later data were too intermittent and unreliable to be included
in a climatological survey. :

In the following sections the meteorological instruments are described
and weather data observed immediately above the canopy are tabulated.

INSTRUMENTATION

The variables that were measured include temperature, humidity, solar
radiation, rainfall, wind speed and direction, and soil temperature. A

420 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

ABOVE: Fic. 1. Thermometer shelter just above cano
ELOW: Fic. 2. The 20-foot tower showing tipping- bucket rain gage, 3-cup
anemometer, wind vane, and wind-powered generator on top

1968 | BAYNTON, ELFIN FOREST, 2 421

relative measure of the amount of water extracted from the impinging
cloud was also made. However, since the interpretation to be placed on
this measurement is not self evident, the subject will be deferred to a
later report.

emperature and humidity were measured inside a thermometer shelter
designed to protect the instrument from direct rain and sun, while assur-
ing adequate ventilation by its louvered structure. The shelter was
mounted just above the canopy on the roof of the hut as shown in Fic.
1. Solar radiation was measured on top of the thermometer shelter. Rain-
fall was measured just at the top of the canopy on a 20-foot tower shown
in Fic. 2 and below the canopy. Wind speed and direction were measured
at the top of the tower about 8 feet above the canopy, and wind speed
was measured below the canopy also.

Because of the lack of commercial electric power, instruments that re-
quired little or no power were used. A limited amount of 12-volt direct
current was supplied by a wind-powered generator shown atop the tower
in Fic. 2

A standard hygrothermograph, which requires no power, provided con-
tinuous records of temperature and relative humidity. The instrument is
shown inside the thermometer shelter in Fic. 3. A Bourdon tube and a
bundle of hairs were the sensing elements for temperature and humidity

Sedat niece pe thermometer shelter. Cloud-water
oof.

Lert: Fic
éallectie: in backeiueel ol-a-m
Ricut: Fic. 4. Close-up of Sol-a a- cmten:

422 JOURNAL OF THE ARNOLD ARBORETUM [ VoL. 49

respectively. Periodically the calibration of the instrument was checked

y a wet and dry bulb psychrometer. A spring-wound 8-day clock per-
mitted weekly chart changes. Fic. 5 is an example of a typical chart.
These charts gave daily values of the maximum and minimum temperature,
the minimum relative humidity and the duration of less-than-100 percent
relative humidity. Experience showed that the latter humidity condition
was diagnostic of an absence of cloud on the peak.

Solar radiation was measured by a Sol-a-meter shown in Fic. 4. Four
parallel-connected silicon cells generate a current proportional to the
incident solar radiation. The output of the cells is connected through a
milliammeter that indicates the instantaneous insolation rate, and to an
ampere-hour meter that gives a readout of the total solar radiation during
any desired period of time. A dessicant of silica gel is changed when a
change in its color indicates an accumulation of moisture inside the glass
cover. Readings near the beginning and end of the month were used
to give an average value of the daily accumulation of solar radiation. The
Sol-a-meter also required no power.

The rainfall detector was a tipping-bucket gage, shown in Fic. 2;
rain from the collector is funnelled to the tipping-bucket mechanism
shown in Fic. 6. Each bucket-tip, resulting from 1/100 inch of rain,
closed a magnetic reed switch connected to the 12-volt supply. The
switch-closure deflected a pen in an Esterline Angus 20-pen event re-
corder, Thus the amount of rain in hundredths of an inch is equal to
the number of pen deflections. Rate of rainfall was obtained from the
elapsed time between bucket-tips as read from the chart, on which 6 inches
of chart-travel represented 1 hour.

The event recorder was also used to record wind speed and direction.
Each mile of wind passage closed a reed-switch inside a 3-cup anemometer,
deflecting another pen in the recorder. Fic. 7 shows the below-canopy
anemometer. Hourly average winds and the fastest mile of wind were
readily obtained. The wind vane, shown in Fic. 8, is attached to a ver-
tical shaft that activates a contacting assembly consisting of eight wiper-
type contacts spaced at 22.5 degree intervals. Only one contact is closed
at a time. A separate event pen was assigned to each of the eight cardinal
directions. By means of a cam switch built into the recorder, once every
3 minutes the circuit was closed and a wind direction was recorded.

Soil temperature at a depth of 8 centimeters was measured by a mercury
thermometer. The thermometer was read during the regular visits of the
observer.

RESULTS

TasBLe | contains a summary of mean monthly temperatures, winds,
humidity, solar radiation, and total rainfall. Gaps in the record occur
when equipment was not operating.

The dominance of the northeast trade winds was almost complete. For
the seven months of record the frequency of winds from the north, north-

NDAY TUESDAY

WEDNESDAY

[=] wnememameeed Sn ee AY
=f tous ce sole woo oMes ee oles seaWess soles se eWe +s a elke +50 xls epeMess gels ce cWieres oll re eoMe ce

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ia HE: gee FEATeIESEEE spss: issezsrs2csss2cer25 EFEsEEoETEEEDESEEE
{Bee seseeesusgeee=25=5=5526=522c22rersessae=2222255° 5555552 szsszsfsPaeeeer arridte geneeias
Sese8 (7 sSeeeeseen ne SrEESSeeES|
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3

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HYGROTHERMOGRAPH TRACE AT PICO DEL OESTE , JANUARY 2-9 ,1967
Fic. 5. Hygrothermograph chart for Pico del Oeste, January 2—9, 1967.

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424 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

east, and east totalled 81 percent. The fastest mile of wind was 32 mi./hr.
on 26 August as a hurricane passed just south of the island.

Rainfall totalling 453 cm. for the year was probably not abnormal.
Picéd (1950) gives 478 cm. as the 10-year average at La Mina, located
about 5 km. northwest of Pico del Oeste at an elevation of 700 m. Ac-
cording to the U.S. Weather Bureau (1965) 402 cm. is the normal annual
rainfall at Upper Rio Blanco less than 3 km. southwest of the peak at
an elevation of 450 m. The heaviest rate of rainfall during the year

Fic. 6. Tipping bucket mechanism housed inside rain-gage directly under fun-

nel from collector. When one compartment fills bucket tips, closing switch and
simultaneously positioning other compartment under funnel.

occurred in the forenoon of 15 April 1966 when 2.5 mm. fell in just
under a minute, a rate of just over 15 cm

Solar radiation at Pico del Oeste was eenahaia! to Tampa, Florida,
and Swan Island, the two nearest stations for which radiation data are
available. Normally Swan Island gets about 130 cm. of rainfall annually
compared to 153 cm. at San Juan, and since it is at nearly the same
latitude as Pico del Oeste it is comparable to the coastal plains of eastern
Puerto Rico. Tampa probably gets somewhat less radiation than the
coastal plains of Puerto Rico because it is further north. For similar
(although not identical) periods in 1966 solar radiation averaged 478
Langleys/day at Tampa and 518 at Swan Island compared to 262 at
Pico del Oeste.

This suggests that over 40 percent of the incident radiation is at-
tenuated by the cloud, a plausible value since the peak is enveloped in
cloud for about 60 percent of the daylight hours. Based on two years
data Briscoe (op. cit.) found that solar radiation on a nearby mountain,

1968 | BAYNTON, ELFIN FOREST, 2 425

1065-meter El] Yunque, was 60 percent of that measured at Cape San
Juan on the coastal plain.

Moreover, as pointed out by Szeicz (1966), since the spectral com-
position of light changes very little with cloudiness, the light energy
available for photosynthesis is also attenuated by over 40 percen

ences are readily accounted for by their 1 kilometer (1000 m.) difference in
height. During the warmest part of the day temperatures in the first few
hundred meters above ground usually decrease at the rate 9.8° C/km. (the
dry adiabatic lapse rate). Maximum temperatures were observed to dif-
fer by just about this amount. Rainfall at the peak includes a substantial
contribution from the orographic process, namely the forced rising, cool-
ing, and resultant condensation within the moist northeast trade winds.

The climate on Pico del Oeste is described further in TABLE 3 in terms
of occurrences of meteorological events. The first two columns are events
or attributes of unpleasant weather. The last three columns are favorable
weather attributes. There are two seasons of favorable weather and two sea-
sons of unfavorable weather. Favorable weather in March and April co-
incides with the normal dry season over the Caribbean; the favorable
weather of October and November may have been an anomaly since these
are normally among the wettest months. It is noteworthy that measurable
rain fell during 350 days in the year.

Fic. 7. 3-cup anemometer used for measuring wind speed below the canopy.

426 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

The longest spell of fine weather extended from November 8 to 28
or a total of 3 weeks. During this remarkable period there was an average
of 15 hours per day of cloud-free weather and radiation averaged 305
Langleys/day. Maximum temperatures averaged 21°C and _ relative
humidity averaged 91 percent. Even so, rain totalled 20 cm.

There were also spells distinguished by their lack of rain. The best
example occurred during the normal dry season that generally includes
February, March, and April. In the 23 days from March 20 to April 11,
1966, the total rainfall was 6.5 cm., there was an average of 9 hours/day
of cloud-free conditions, humidity averaged 95 percent, maximum tem-
peratures averaged 20.5° C, and radiation averaged 360 Langley/day.

There were also two notable spells during which cloud enveloped the
peak almost continuously. In the 16 days from June 10 to 25, 1966, there
were only 6 hours in which the cloud broke from the peak. On 12 of the
days the relative humidity never fell below 100 percent. Maximum tem-
peratures averaged 20.5° C and rainfall totalled 24 cm.

In the 15 days from January 14 to 28, 1967, there were only 7 hours
in which the cloud cleared from the peak and on 12 of the days it did not
clear at all. This was a cool spell in which maximum temperatures ave-
raged 17.2° C and rainfall totalled 19 cm.

Some additional noteworthy features may be mentioned. The high-
est and lowest observed screen temperatures during the year were 26° C
and 13° C. The diurnal temperature variation is very small. As shown
in Fic. 5, it is well defined only on days when the cloud lifts from the
peak (i.e. when the humidity falls below 100 percent) for several hours.
The peak was enveloped in cloud close to 100 percent of the night hours
and 60 percent of the day hours.

SUMMARY

Garnier (1961) defines the humid tropics as “the area where (1) the
mean monthly temperature for at least eight months of the year equals or
exceeds 68° F (20° C); (2) the vapor pressure and relative humidity
for at least six months of the year average at least 20 millibars and 65
percent respectively; and (3) the mean annual rainfall totals at least
40 inches (1000 millimeters), and for at least six months precipitation is
3 inches (75 millimeters) each month.” By this definition, Pico del
Oeste is clearly not in the humid tropics. During the year reported herein
rainfall met the criterion. Although relative humidity more than met
the criterion, average vapor pressure exceeded 20 mb. in only 6 of the
months, Temperatures were clearly too cold. However the presence of
an elfin forest rather than a rain forest tells even more eloquently that
Pico del Oeste is not in the humid tropics.

There has been much speculation as to the causes of elfin forests. In
discussing this region in Puerto Rico, Cook and Gleason (1928) asserted
that high wind exposure is the most important environmental factor in
differentiating the mossy forest from the palm forest, although low tem-

1968 | BAYNTON, ELFIN FOREST, 2 427

perature, water-soaked soil, and high humidity were also considered im-
portant. They added that on rocky summits of peaks the trees are bush-
like on account of the strong winds and thin and rocky soils. Grubb and
Whitmore (1966) suggest that the most important factor in the develop-
ment of tropical montane forests is the frequency of fog, although they
concede that additional exposure to winds and leaching of soils on ridges
may contribute to stunting. The first year of data on Pico del Oeste offers
little support to the belief that strong winds account for the stunting.
Leaching of soils is probably important. The most unusual features of
the weather appear to be the high fog incidence and the 40 percent at-
tenuation of solar radiation, tending to support the suggestions of Grubb
and Whitmore.

Fic. 8. Wind vane on top of tower.

ACKNOWLEDGMENTS

The contributions of the following individuals are gratefully ac-
knowledged. Mr. Donald Eklund made the original installation of
meteorological equipment including the tower and wind-powered generator,
and designed and assembled the trough rain-gage for measuring rainfall
under the canopy. Later in the year special maintenance and further re-
finements to the installations were carried out by Dr. Julian Pike. Var-
ious members of the staff of the Arnold Arboretum performed the regular
chart changes and other routine duties that are essential to a program of
continuous data collection.

TABLE 1. Mean monthly values of temperature, wind, humidity, rainfall and

solar radiation, Pico del Oeste, 1 March 1966 to 28 February 1967

git SoIL Solar
TEMPERATURE WIND ae

TEMP. Radiation

Daily Daily Daily 8 cm. Prev. Mean Rel Rain- Langleys'

Max. Min. Mean Depth Dir. Speed Hum fall per day
Marcu ’66 19:2°C 16.5°C 179°C 16.92 — — 98 %G 34.8 cm. 245
APRI 20.0° 16.77 18.4° 17.0% — — 96 38.5 o22
AY 20.0° 18.4° 19.2° 183° _ — 99.9 45.1 203
JUNE 20.4° 18.8° 19.6° 18.9° — —_— 99.8 36.4 260
JULY 20.77 18.9° 19.8° 19.22 — 11.0 kt 99.7 43.9 249
AUGUST 4 OF hag 18.5° 19.6° 19:3" NE 11.1 99.8 41.8 266
SEPTEMBER 210 18.4° 19.8° 19.7? NE 9.6 98 39.9 290
CTOBER 216° 18.6° 204" 19.4° NE 8.0 97 25.0 271
NOVEMBER 21,0" 16.8° 19.0° 18.72 N — 94 27.4 297
DECEMBER 13.3” 16.3° eg 178° NE 9.3 99 62.0 £12
JANUARY ’67 17.5 15.6° 16.6° 17.0° NE 10.9 99.4 30.3 a
FEBRUARY 7.6" 15,67 16.6° 17.0° NE 9.2 99.3 27.6 —
ANNUAL 19.9° 17.4° 18.6° 13.37 — —_ 98.5 453.3 262

“One Langley is 1 calorie/square cm.

82r

WOLAXOPAV GIONUV AHL AO TYWNUAOL

6b “T0A]

TABLE o Pega ions of monthly mean temperatures and rainfall at Pico del

ste and Roosevelt Roads, 1 March 1966 to 28 February 1967

Max. Temp.

Min. TEMP. MEAN TEMP. RAINFALL

MontTH Pico RR Pico RR Pico RR Pico RR
Marcu °66 192°C PAs 165°C 23.0°C Lee baa Be 20°1°C 34.8 cm 7.5 em
APRIL 20.0° 29.6° 16.72 73.0° 18.4° 26.4° 38.5 110

AY 20.0° 29.8° 18,4° 24.1° 19;2° 26.9° 45.1 9.3
JUNE 20.4° 30.9° 18.8° 251° 19.6° 28.0° 36.4 10.0
JuLy 207° 31535 18.9° 253" 19.8° 255° 43.9 10.5
AuGuSs 20:7° 5g oe 18.5° 25.4° 19.6° 28:2° 41.8 10.8
SEPTEMBER 21:1" 313° 18.4° CAO le 19.8° 232° 39.9 17,0
OcTOBER 21.6° WED 18.6° 23.9° 20:12 205° 25.6 18.2
NOVEMBER 21-0" 30.0° 16.8° 218° 19.0° 25.95 27.4 W747
DECEMB 18.3° 28.4° 16:3" 224° 3° 25.4° 62.0 12.9
JANUARY ’67 175° 27.8° 15:67 215° 16.67 24.7° 30.2 10.0
FEBRUARY 17:6" 25.3" 156° 7 Ree 16.6° 25:0" 27.6 3.9

[S961

Z ‘LISAYOA NIATA ‘NOLNAVG

62+

430 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Particular mention must be made of the kindness of Mr. J. B. Martin-
son in providing excellent accommodation close to Pico del Oeste. Both
the scope and quality of the research have benefited greatly from this
generous support.

TABLE 3. Occurrences of significant meteorological events by months,
Pico del Oeste, 1 March 1966 to 28 February

NuMBER oF Days

Incloud Raini Max.temp. 8 or more hours
for or more 22°C or with Rel. Humid. No
m

24 hrs. cm ore less than 100% Rain

Marcu ’66 9 12 3 7 3
APRIL 3 9 4 12 5

AY 20 17. 0 1 0)
JUNE ae. 14 0) 1 0
JuLy 14 12 1 2 0
AUGUST 18 13 3 2 0
SEPTEMBER 8 12 4 i 1

CTOBER 8 7 7 11 1
NOVEMBER 3 9 7 19 -
DECEMBER 10 16 2 8 1
JANUARY 67 15 9 0 3 0
FEBRUARY 13 7 0) 6 0
ANNUAL 138 137 31 79 15

REFERENCES

Briscog, C, B. Weather in the Luquillo Mountains of Puerto Rico. Forest
ervice Res. Pap. 1TF-3, Inst. Trop. Forestry, Rio Piedras, Puerto Rico.

Cook, M. T., & H. A. GLEason. Ecological survey of the flora of Porto Rico.
. Jour. “eee i pie Porto Rico 12 (1 & 2). 1928,
ARNIER, b. J. Mapping the humid tropics: climatic criteria. one
serie re er, p atic criteria. Geogr. Rev
GLossary oF METEOROLOGY, American Meteorologi i
. gical Society. Boston. 1959.
Gruss, P. J.. & T. C. Wuirmore. A com arison of montane and lowland rain

US. WEATHER Bureau. Climatography of the United States no. 86-45. Cli-
matic summary of the United States. Supplement for 1951 through 1960.
Puerto Rico and United States Virgin Islands. Supt. Doc. U.S. Govt

Printing Office, Washington. 1965.

WapswortH, F. H. The Climate of the Luquillo Mountains and its significance

to the people of Puerto Rico. Carib. Forest. 9(4): 321-344. 1948.

NATIONAL CENTER FOR ATMOSPHERIC RESEARCH
BouLDER, CoLorapo 80302

1968] SMITH & STONE, ARALIACEAE 431

STUDIES OF PACIFIC ISLAND PLANTS, XIX.
THE ARALIACEAE OF THE NEW HEBRIDES, FIJI, SAMOA,
AND TONGA |!

ALBERT C. SMITH AND BENJAMIN C. STONE

THIS REGIONAL TREATMENT of the Araliaceae is primarily directed to-
ward an adequate understanding of the species native to Fiji. Although
only three genera (Polyscias, Plerandra, and Sche flera) occur indigenously
in that archipelago, an examination of the family in the nearby island
groups has proved instructive. In the area of our present concern species
of Polyscias, Brassaia, and Dizygotheca occur as cultivated or adventive
plants, and for convenience we also include these records.

Generic limits in the Araliaceae have presented problems to students of
the group, and we are well aware that a limited and regional treatment
can add little to their solution. In his recent attempt to bring taxonomic
order into the family, Hutchinson (Gen. FI. Pl. 2: 52-81. 1967) has
recognized 84 genera, rearranging them in seven tribes. Harms’ treat-
ment of 1894 (in E. & P. Nat. Pflanzenfam. 3(8): 1-62) had accepted
51 genera in three tribes. Hutchinson has proposed two new tribes (Cus-
sonieae and Anomopanaceae) for taxa that have their flowers singly dis-
posed rather than in umbels. His first 12 genera, removed from the rest
of the family on that basis, seem to form an unnatural aggregation. As
a single example, the genus Polyscias is not amenable to this tribal cate-
gorization; its species are usually umbelliferous, but some of them have
the flowers individually borne on the ultimate inflorescence-branches, and
other species are transitional in this character. We find Polyscias to be
a satisfactory and coherent genus, as noted in our generic consideration,
below, and in our notes under P. joskei and P. reineckei. By combining
Anomopanax Harms (the basis of Hutchinson’s tribe Anomopanaceae)
with Mackinlaya F. v. Muell. (the basis of the tribe Mackinlayeae of both
Harms and Hutchinson), Philipson (in Bull. Brit. Mus. (Nat. Hist.) Bot.
1: 3-9. 1951) has also indicated his lack of confidence in the value of
umbellate or single flowers as a generic character. Our comments below
under Plerandra insolita will further illustrate the difficulties of utilizing
this character for generic delimitation.

In connection with the present paper the herbarium material of several

* No. XVIII of pe series, by the first author, was published in Contr. U.S. Natl.
Herb. 37: 69-106. 1967. The present paper is based on research partially supported
by a grant from the National Science Foundation. Our work on the Araliaceae was
initiated while both authors were staff members of the Smithsonian Institution, and

we take this opportunity of thanking Dr. Wallace R. Ernst, of that org anization ; ior
his interest in the problem, his advice, and his aid in handling borrowed mater. ‘al.

432 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

institutions has been examined and is cited, with the indicated abbrevia-
tions: Arnold Arboretum of Harvard University (A); Bernice P. Bishop
Museum (BIsH); British Museum (sm); Gray Herbarium of Harvard
University (GH); Royal Botanic Gardens, Kew (k); New York Botanical
Garden (Ny); Department of Agriculture, Suva, Fiji (suva); University
of California, Berkeley (uc); and U.S. National Herbarium (us). We
are indebted to the administrators of these herbaria for the privilege of
examining the cited specimens. The accompanying illustrations were pre-
pared by the second author.

KEY TO THE GENERA

Petals imbricate in bud (tribe Aralieae) ; leaves imparipinnate; flowers in umbels,

the pedicels conspicuously articulate at apex: petals and stamens 5; styles

d ovary-locules 2 (rarely 3); fruits ellipsoid or obovoid. .. 1. Delarbrea.
Petals valvate, broad at base (tribe Schefflereae ).

Flowers densely congested in heads, unisexual; fruits sessile, free to firmly

concrescent; leaves simple, obovate to oblanceolate. ........ 2. Meryta.

Flowers pedicellate (or, if capitate, the leaves compound); fruits not con-

Leaves pinnately compound or unifoliolate.

Pedicels articulate at apex; petals and stamens (in our species) 5-8 (rare-
y 9); styles and ovary-locules (in our species) 2-5; fruits
often laterally compressed (sometimes subglobose), the styles often
conspicuous and persistent (but sometimes stigmas sessile on a
ROM CVI ee ey . Polyscias.

Pedicels not articulate; petals and stamens 8-12; styles and ovary-lo-
cules 8-24; fruits subglobose, surmounte by a truncate-conical
stylar column with numerous radiating sessile stigmas.............
es 4. Reynoldsia.

Leaves palmately lobed or compound.
Leaves digitately compound, with petiolulate leaflets.
Stamens numerous, 15-500, exceeding the petals in number; petals
4-6; styles, stigmas, and ovary-locules 5-17. .... 5. Plerandra.
Stamens 4-6, similar to the petals in number; styles, stigmas, and
ovary-locules 5—12
Anthers 2-locular.
Flowers. pedielath cc ee ee 6. Schefflera.
Flowers sessile in heads borne along the radiating branches of
large panicles. 7, hs
Anthers 4-locular.

1. Delarbrea Vieill. in Bull. Soc. Linn. Norm. 9: 342. 1865; Harms in

E. & P. Nat. Pflanzenfam. 3(8): 61. 1894; Hutchinson, Gen. FI. Pl.
2: 63. 1967.

This small genus, of about seven species, is centered in New Caledonia

1968 | SMITH & STONE, ARALIACEAE 433

but extends from Timor and New Guinea through the Solomon Islands
into the New Hebrides; it is typified by D. collina, the only species oc-
curring in our area. Our specific description is based on New Hebridean
and New Caledonian material. Both Harms and Hutchinson place Delar-
brea in the tribe Aralieae, characterized by having the petals broad-based
and imbricate in bud.

1. Delarbrea collina Vieill. in Bull. Soc. Linn. Norm. 9: 342. 1865;
Guillaumin in Bull. Soc. Bot. France 66: 270. 1919, in op. cit. 74:
698. 1927, in Jour. Arnold Afb. 12": 262, 1931, in Jour. Linn, Soc.
Bot. 51: 553. 1938; Philipson in Bull. Brit. Mus. (Nat. Hist.) Bot.
£5.48: 1951; PL. III, Fics. 10-13.

A tree, glabrous throughout; leaves imparipinnate, large, up to 85 cm.
long, the petiole and rachis subterete, the petiole of mature leaves 12-15
cm. long, swollen to a clasping base and obviously alate in the proximal
1.5—2 cm., the leaflets 11-19, the petiolules slender, of lateral leaflets 3-11
and of terminal leaflet to 40 mm. long, the blades chartaceous, oblong or
oblong-lanceolate, 12-20 cm. long and 4—7 cm. broad (basal ones some-
times smaller), unequally subcordate to broadly obtuse at base, acute to
sharply short-acuminate at apex, entire at margin, the costa raised above,
prominent beneath, the secondary nerves usually 7-10 per side, curved-
ascending, with the intricate veinlet-reticulation prominulous on both
surfaces; inflorescences racemose-paniculate, up to 45 cm. long and broad,
the primary branches to 25 cm. long, all the inflorescence-axes transversely
griseo-corticate-lenticillate, the primary bracts oblong, 0.5—1 cm. long or
more, caducous, the umbel- and flower-subtending bracts and bracteoles
1.5—-3 mm. long, obtuse or subacute; flowers borne in umbels on slender
ultimate peduncles (5—) 10-30 mm. long, 15-25 per umbel, the pedicels
1-3 mm. long at anthesis, slightly elongating in fruit, swollen and con-
spicuously articulate at apex; flowers 3~3.5 mm. long at anthesis, with a
narrowly turbinate calyx 2—2.6 mm. long, the tube obconical, the limb
subcampanulate, 1.5—2 mm. in apical diameter, the lobes 5, obvious, orbi-
cular-ovate, 0.5—0.7 mm. long, narrowly imbricate in bud; petals 5, oblong-
ovate, broad-based, about 1.5 mm. long, imbricate in bud; stamens 5, the
filaments slender, eventually about 1.5 mm. long, the anthers oblong,
about 0.8 mm. long, versatile; stylopodium hemispheric, 0.5—0.7 mm. high,
surmounted by 2 (rarely 3) styles free nearly to base, these elongating
o 1.5 mm., eventually recurved and subclavate, with decurrent stigmas;
fruits ellipsoid or obovoid, 10-15 mm. long, only slightly compressed
laterally, 2- or sometimes 3-locular, each locule containing a marginal
pyrene angular-lunulate in cross-section, the fruit thus with an empty
central cavity, the sepals persistent, the styles usually caducous.

Type Locatity: New Caledonia; the type is Vieillard 625, from the
vicinity of Wagap (isotype at A).

DistrisuTIon. New Caledonia, Loyalty Islands, Solomon Islands, and
New Hebrides, apparently occurring in forest from sea-level to an alti-

434 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

tude of 200 meters. Trees up to 12 meters high have been noted, and
the fruit is said to be black when mature.

Locat NAMES. Reported names in the New Hebrides are morshmor-
shalkara and névarkrab (Espiritu Santo), napouiri (Efate), and nungine-
tum (Eromanga).

New Hebrides. BANKS Group: VANUA Lava: Kajewski 412 (A, BISH). Espiritu
Santo: Hog harbour, /. & Z. Baker 72 (pm), 295 (pm). Eromanca: Dillon Bay,
Kajewski 269 (a, us).

The New Caledonian species, with the exception of this one widespread
one, appear to be endemic. Among them (according to Guillaumin, Fl.
Anal. et Syn. Nouv.-Caléd. 251. 1948), D. collina is distinguished by its
acute, not falciform, leaflet-blades, and its relatively small flowers and
fruits.

2. Meryta J. R. & G. Forst. Char. Gen. Pl. 119. 1776; Seem. in Bon-
plandia 10: 294. 1862, Fl. Vit. 118. 1865; Harms in E. & P. Nat.
Pflanzenfam. 3(8): 34. 1894, in Notizbl. Bot. Gart. Berlin 14: 315.
1938; Hutchinson, Gen. Fl. Pl. 2: 74. 1967.

Botryodendrum Endl. Prodr. Fl. Norfolk. 62. 1833.
Neara Sol. ex Seem. Fl. Vit. 118, as synonym. 1865.
Strobilopanax Viguier in Ann. Sci, Nat. IX. Bot. 4: 148. 1906.

The interesting genus Meryta, typified by M. lanceolata J. R. & G.
Forst., of the Society Islands (cf. Forst. f. Fl. Ins. Austr. Prodr. 92. 1786),
is readily distinguished by its large, simple leaves, of which the blades are
usually obovate or oblanceolate, and its unisexual, sessile flowers crowded
in capitate-paniculate inflorescences. The fruits are either free or firmly
concrescent, thus indicating that the character upon which Strobilopanax
was based is not dependable at the generic level.

Meryta is indicated by Harms (in 1938) to include about 30 species,
by Hutchinson (in 1967) to include 16 species. It has its greatest con-
centration in New Caledonia, but it is now known to extend from New
Zealand, Norfolk Island, New Guinea, and the Palau and Marianas
Islands eastward through Polynesia to the Austral Islands, the Tuamotus,
and the Marquesas. In this vast area, the only major archipelago lacking
the genus is Fiji, a very curious fact but one that must now be taken at
face value, since Fiji is better known than most Pacific areas and since the
genus can scarcely be confused with any other.

The genus is greatly in need of a complete review, since specific charac-
ters are evasive. In our area at least one species occurs in the New He-
brides and one or two in Samoa and Tonga. We are able to record little
new information, but the following key and pertinent bibliographic and
herbarium records will summarize an unsatisfactory situation.

KEY TO THE SPECIES

eager heads composed of fruits rer concrescent except in the distal portion,
r at least united above the middle

1968 | SMITH & STONE, ARALIACEAE 435

Petioles 1~1.5 cm. long; leaf-blades 28-32 cm. long, obtuse at st fruiting
heads at apparent maturity about 4 cm. in diameter. .. 1. o-ebudica.
Petioles 2-16 cm. long; leaf-blades 40-63 cm. long, acute to ices at apex;
fruiting heads at apparent maturity 2-3.5 cm. in diameter. . 2. M. capitata.
Fruiting heads composed of fruits nearly free or united only below the middle.
Petioles up to 23 cm. Bead leaf-blades 40-75 cm. long, cuspidate to short-

Oe Rae ee ee . M. macrophylla.
Petioles 4-8 cm. ae Jeaf- blades 45-120 cm. long, obtuse to acute at apex.
4. M. denhamii.

1, Meryta neo-ebudica (Guillaumin) Harms in Notizbl. Bot. Gart.
Berlin 14: 321. 1938.

Strobilopanax neo-ebudicus Guillaumin in Jour. Arnold Arb. 12: 263. 1931.

As we have seen no additional material of this species, the original
description based on the type collection cannot be significantly amplified.

Type LocALity: Aneityum, New Hebrides; the type is Kajewski 980
(coll. J. P. Wilson), cited below.

DiIsTRIBUTION. Endemic to the New Hebrides and known definitely
only from the type collection, obtained in forest at an altitude of 325 m.
and noted as a small tree with white flowers and yellow fruits.

Locat NAME. Nabrouto was noted by the collector.

New Hebrides. ANEITyYumM: Anelgauhat Bay, September 1929, Kajewski 980
(coll. J. P. Wilson) (a type).

Guillaumin notes his novelty as the first record of Strobilopanax in the
New Hebrides, but he has also indicated the presence of Meryta in that
archipelago. Earlier (in Bull. Soc. Bot. France 66: 270. 1919, in op. cit.
74: 698. 1927) he had reported the genus from Efate, with the local name
nafil gnas; and more recently (in Jour. Linn. Soc. Bot. 51: 554. 1938) he
has reported it from Espiritu Santo, with the local name mvalval. The
specimens supporting these records have not been examined by us.

2. Meryta capitata Christophersen in Bishop Mus. Bull. 128: 161. fig.
23. 1935; Yuncker in Bishop Mus. Bull. 184: 55. 1945.

Meryta sp. Christophersen in Bishop Mus. Bull. 128: 164. 1935.

Meryta aff. macrophylla sensu Yuncker in Bishop Mus. Bull. 184: 56. 1945.

In view of our uncertainty of the actual limits of this species or, indeed,
of its real distinctness from M. macrophylla (see comments below), we
refrain from amplifying the original description, which Christophersen
seems to have based entirely upon fruiting specimens — his type and his
no. 2839

Type tocatity: Near Vaipouli, Savaii, Samoa; the type is Christo-
phersen & Hume 1913, cited below.

DistripuTion. Endemic to Samoa, and definitely known from Savaii
and the Manua Islands, where it has been collected in forest from near
sea-level to 500 m. altitude (with one collection from about 1,400 m.
above Matavanu, Savaii) and noted as a tree 4-6 m. high.

436 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Loca, NAMES. Fagu fagu (i.e. fangu fangu) is a general name; it has
also been recorded on Savaii as Jauma ulu ulu, and on Olosega as lau tetali.
moa. SAVAII: steed Christophersen 2802 (s1sH); Manese plantation,
Pie cna & Hume 2369 (BIsH); ec Avao, "Vaupel 135 (BtsH); back
of Vaipouli, Chatsiothorien & Hume 1913 (A, BIsH type, Us); near Vaipouli,
Christophersen & Hume 1837 (a, BISH); above Matavanu, yaad cia &
Hume 2199 (sis); Taga, Christophersen 2839 (BISH, US). : Above Ofu
village, Yuncker 9545 (BtisH). OLoseca: Oloseguata, ee lee Garber
1079 (sisH). Tau: Amouli trail, Garber 622 (a, BISH); south of Siufaga, Yunc-
ker 9162 (BISH)

It may be seriously doubted that more than one species of Meryta oc-
curs in Samoa and Tonga. As characters to differentiate his M. capitata
from M. macrophylla Christophersen mentions the subglobose heads of
united fruits and the broader leaves. The latter of these is surely of little
consequence, as there is substantial variation in leaf-breadth and also in
petiole-length in plants that otherwise seem essentially identical.

Fruiting specimens referable to M. macrophylla (U.S. Expl. Exped.,
the type, Setchell 15667, Bryan 99, Christophersen 1253, and Yuncker
16181) have the fruits united only in the basal portion or up to the mid-
dle. Fruiting specimens referable to M. capitata (Christophersen & Hume
1913, the type; Vaupel 135, Christophersen 2839, and Yuncker 9162)
have the fruits concrescent except in the uppermost portion, which is
rounded and obtusely costate when dried. The dividing line between the
two conditions is not entirely clear, and indeed is approached in Vaupel
135. By matching intangible foliage characters of staminate and sterile
specimens with the fruiting collections we have grouped the available
material into two taxa, but without much conviction that species are
represented. Staminate flowers are quite uniform throughout, as are
stylar characters; the petiole seems totally variable in length and diameter
and probably reflects conditions of age or habitat.

3. Meryta macrophylla (Rich ex A. Gray) Seem. in Bonplandia 10:
294. 1862, Fl. Vit. 119. 1865; Drake, Ill. Fl. Ins. Mar. Pac. 182.
1890; Hemsl. in Jour. Linn. Soc. Bot. 30: 180. 1894; Burkill in
Jour. Linn, Soc. Bot. 35: 40. 1901; Christophersen in Bishop Mus.
Bull. 128: 163. 1935; Yuncker in Bishop Mus. Bull. 220: 206. 1959.

Botryodendrum ig ime Rich ex A. Gray, Bot. U.S. Expl. Exped. 1:
732. 1854, Atlas pl. 97. - seed

A redescription of this species should await a detailed revision of the
genus; hopefully a specialist will reach a satisfactory conclusion as to the
limits of taxa in Samoa and Tonga.

TYPE LOCALITY: Samoa, without oo locality; the U.S. Exploring
Expedition specimen cited below is the
DISTRIBUTION. Samoa (known aes from Upolu and Tutuila) and

Tonga, where it occurs in forest at elevations from near sea-level to 700
m. and has been noted as a tree 2.5—8 m. high

1968 | SMITH & STONE, ARALIACEAE 437

Loca NAMES. Fagu fagu is the general name in Samoa and kulukulu
in Tonga; further notations indicate leva va’u (Upolu), lau fagu fagu
(Tutuila), and kulukulufa (Vava’u).

Samoa. Upotu: Malololelei, Christophersen 304 (sisu), Setchell 15667
(BisH); above Malololelei, Christophersen 236 BISH); Vaea Mt., Bryan 99
(BIsH). TutuILA: Pago Pago and vicinity, Christophersen 1253 (BIsH), 1254
(BISH), Garber 929 (BisH); Alava Ridge, Christophersen 1130 (pisH, vs);
Papatele Ridge, Christophersen 1006 (BIsH, US). SAMOA, WITHOUT FURTHER
Locality: U.S. Expl. Exped. (GH, us 62360 & 73912 type).

Tonga. Vava’u: Northwestern side of island, Yuncker 16181 (BIsH, GH, US);
Talau hill, MacDaniels 1095 (s1sH); above Anovai Lake, Yuncker 16050 (BIsuH,
GH, Us). Eva: Above Fuai, Vuncker 15688 (BISH, US).

As an example of the difficulty of specific delimitation in Meryta one
may note that Harms (in Notizbl. Bot. Gart. Berlin 14: 320. 1938) has
referred a collection from the Tuamotus to M. macrophylla. In spite of a
similarity in fruit, this particular specimen (St. John 14263, BisH, from
Anaa) has leaf-blades rounded at the apex, and we confidently exclude
it from the species of Samoa and Tonga.

4. Meryta denhamii Seem. in Bonplandia 10: 295. 1862; Harms in
Notizbl. Bot. Gart. Berlin 14: 318. 1938.
Meryta denhami Seem. FI. Vit. 119. 1865; Hemsl. in Bot. Mag. 129: pl. 7927.
1903.

As we have not seen material of this species, we cannot add to the
above-cited descriptions.

TYPE LOocALITy: New Caledonia: Isle of Pines; the type is from a
specimen cultivated at Kew, grown from material obtained by W. G.
Milne on Captain Denham’s Expedition.

DistrIBUTION. In addition to the Isle of Pines, the species is attributed
by Hemsley (as cited above) to the Banks Islands, New Hebrides, by the
statement that “. . . what appears to be the same species has since been
collected by Archdeacon Comins. . .”’ on Santa Maria (i.e. Gaua Island).

The species is listed merely because of Hemsley’s note, but its occur-
rence in the New Hebrides is certainly open to question without re-study
of the material and the several species attributed to New Caledonia. The
excellent figure cited above clearly indicates the nature of the pistillate
plant, which at any rate seems quite distinct from M. neo-ebudica, the
only species of the genus positively recorded from the New Hebrides to
date,

3. Polyscias J. R. & G. Forst. Char. Gen. Pl. 63. 1776; Seem. in Jour,
Bot. 3: 179. 1865; Harms in E. & P. Nat. Pflanzenfam. 3(8): 43.
1894, in Bot. Jahrb. 56: 409. 1921; Hutchinson, Gen. FI. Pl. 2: 75.
1967.

Nothopanax Miq. in Bonplandia 4: 139. May, 1856, Fl. Ind. Bat. 1(1): 765.

438 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Sept. 1856; Seem. Fl. Vit. 113. 1865, in Jour. Bot. 4: 293, p. p. 1866. Non
sensu Harms in E. & P. Nat. Pflanzenfam. 3(8): 47. 1894, nec > itched
Gen. Fl. Pl. 2:.76. 1967.

In view of diverse concepts of the delimitation of the genera Polyscias
and Nothopanax, it may be well once again to summarize their typification.
Polyscias, being based entirely on P. pinnata J. R. & G. Forst., presents
no problem; its type species has pinnately compound leaves (rarely uni-
foliolate) and 4 or 5 styles, as understood by the Forsters. Only three
species may be considered as typifying Nothopanax in Miquel’s ees
sense, These are N. fruticosum (based on Panax fruticosum L.)*, N
cochleatum (based on Aralia cochleata Lam.), and N. obtusum (based on
Panax obtusum Bl.). Two other species (based on Panax pinnatum Lam.
and Panax anisum DC.) were tentatively referred here, prefaced by inter-
rogation marks, and these may be dismissed from consideration as type
species.

In his original circumscription of the genus Miquel stated that the
styles and ovary-locules were two or three. The only one of the original
three species of which the flowers were definitely known to Miquel was
Nothopanax fruticosum, and it would thus appear imperative to accept
that as his type species (cf. Merr. in Philip. Jour. Sci. Bot. 12: 241. 1912).
However, Sprague and Green (in Kew Bull. 1933: 155. 1933) indicated
N. cochleatum as Miquel’s type species, and in this they have been fol-
lowed by Fosberg (in Univ. Haw. Occ. Pap. 46: 9. 1948), Hutchinson
(Gen. Fl. Pl, 2: 76. 1967), and doubtless by other authors.

A misinterpretation of the identity of Nothopanax cochleatum Miq.
(i.e. Aralia cochleata Lam.) has been responsible for the use of the name
N othopanax for certain species with digitately compound leaves. These
Species, as convincingly shown by Philipson (in New Zeal. Jour. Bot. 3:
333. 1965), should be placed in Pseudopanax C. Koch, which appears to
have a distribution from China to Tasmania, New Zealand, New Caledonia,
and to southern South America. The persistent error of including digitate-
leaved species in Nothopanax may have originated with Seemann (in Jour.
Bot. 4: 293-296. 1866) and been continued by Harms (in E. & P. Nat
Pflanzenfam. 3(8): 47. 1894), but it is unfortunate that Hutchinson (Gen.
Fl. Pl. 2: 76. 1967) prolongs it and even attributes the generic name to See-
mann.

As a matter of fact, indication of NV. cochleatum as the type species of
Nothopanax, even if it were acceptable, would not permit confusion of
that genus and Pseudopanax. Aralia cochleata Lam. is based entirely on
Scutellaria prima Rumph., as pointed out by Merrill (Interpret. Rumph.
Herb. Amb. 409, 1917), and hence is a direct synonym of Polyscias scutel-
laria (Burm. f.) Fosberg. Below we point out the specific identity of this
concept with P. pinnata, the type species of Polyscias. Therefore, those
who wish to follow Sprague and Green in the typification of Nothopanax
must equate the genus directly with Polyscias.

"We the neuter termination for these epithets only to retain at usage of
original sathisis in this discussion, although the correct usage is masculin

1968 | SMITH & STONE, ARALIACEAE 439

Acceptance of Panax fruticosum L. as the type species of Nothopanax
Mig. does raise the question of whether a real discontinuity exists between
the species of this complex with 2 (rarely 3) styles and ovary-locules
(Nothopanax) and those species with 3-5 (rarely 2) styles and ovary-
locules (Polyscias). That this character is highly unsatisfactory has been
indicated by the second author (in Taxon 14: 284. 1965), and our present
study merely supports the conclusion that only one genus is to be main-
tained, a viewpoint apparently adopted by some other current students
of the family (cf. Philipson in Bull. Brit. Mus. (Nat. Hist.) Bot. 1: 9.
1951).

Another genus of this relationship is Tieghemopanax Viguier, which
Hutchinson (Gen. Fl. Pl. 2: 75. 1967) maintains as distinct on the basis
of its 4-locular anthers. We are unable to review the New Caledonian and
Australian species placed in Tieghemopanax, but it seems apparent that
the New Hebridean species so referred are properly placed in Polyscias,
as suggested by the second author in 1965 (in Taxon 14: 285) and as
indicated in the following treatment. pale we refrain from listing
this generic name in the synonymy abov

In the area of the present study, Botecite is represented by 12 species,
of which 9 are clearly indigenous and an additional one (P. scutellaria in-
cluding P. pinnata) is probably indigenous in the New Hebrides. The
species of Polyscias yield a characteristic acrid fragrance, especially when
the leaves are cut or bruised, this being particularly obvious in such
cultivated species as P. guilfoylei.

KEY TO THE SPECIES

Leaves once pinnate (sometimes unifoliolate in no. 10).
Styles or stigmas and ovary-locules 2 (very rarely 3); leaves usually with
numerous leaflets, only occasionally as few as 7 or 9, very rarely as few
as 5; indigenous species.
Flowers borne singly, alternate or loosely whorled on Soles agree
branches; leaflet-blades not more than 11 X
Flowers in umbels (or pseudo-umbels in no. 9) or els leaflet- fide ce
cept in nos. 3 and 4) consistently larger.
Inflorescences paniculately capitate, the flowers sessile; leaflets up to
< 8 cm., strongly asymmetric and cordate at base. ............
Oe pei gee raraiaca Ui aoe dea wry a a aor ie oe eiear ee a rar arg 2. P. nusedhul.
Inflorescences racemose-paniculate, the flowers pedicellate, in umbels
(but pedicels sometimes very short in nos. nd 9).

Petioles slightly swollen at base but not alate; leaflet-blades asymmetric,
obviously falcate, usually 4-9 & 1.8-3.5 cm., unequally acute to
rounded at base; young vegetative parts and inflorescence-axes
with a sparse (and sometimes evanescent) indument of minute 1-
several-celled hairs.

Leaflets (5—) 9-13, the blades coarsely crenulate at margin; petioles

8-14 cm. long on mature leaves; flowers usually 5~12 per umbel,

the stylopodium conical, tapering into the slender stylar column.

3. P. neo-ebudarum.

440 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Leaflets 19-23, the blades entire at margin; petioles 5-9 cm. long on
mature leaves; flowers usually 11-32 per umbel, the disk con-

cave, the stylar column slender. .......... . culminicola.
Petioles alate at base, the wings often obvious; leaflet- blades con-
spicuously larger, usually much more than 9 & 4 cm. (basal ones

occasionally smaller); plants glabrous throughout.
Leaflet-blades ovate-falciform, cordate-auriculate at base. ........
SS ig oie HG, ee NOR CA as erie ge a e :
Leaflet- rier not falcate, subequally obtuse to rude’ or ¢ subcor-
d but not auriculate) at base.
Disk < concave to slightly elevated, the styles obvious, divaricate
in fruit, 1-1.5 mm
garees (umbelliferous ) inflorescence-branches very numerous,
ttered, irregularly arranged; pedicels comparatively
ont. not more than 6 mm. long; flowers with a cupuliform
calyx not more than 2.5 mm g.
Petioles inconspicuously alate in the proximal 3-5 cm.; in-
orescence-axes griseo-corticate-lenticellate; flowers borne
in Some on ultimate peduncles 10-20 mm. long, 5-15
per umbel, the pedicels 2-6 mm. "th ee = P. corticata.
Petioles ines alate in the proximal 7— 20 cm.; inflores-
cence-axes smooth, dark; flowers borne in umbels 0
ultimate peduncles 1-6 mm. long (or sient ssel),
(1—) 2-7 per umbel, the pedicels less than lo
an naline Ther Gg <2 eh ¥ > wdsiraee.
Ultimate Se sateen inflorescence-branches often aggre-
oose whorls at 2-4 nodes; flowers borne in umbels
on lginate peduncles 12-80 mm. long, 8-15 per umbel,
the pedicels conspicuous, 7-15 mm. long (up to 25 mm. in
fruit); flowers comparatively long and narrow, the calyx
of perfect flowers urceolate, 3-3.5 mm. long. .........-.-
ee er ee ee, 8. P. samoensts.
Disk elevated into a broadly conical stylopodium surmounted by
two sessile laterally contiguous fasta the fruits similarly
lacking styles; pedicels short, (0.5—) 1-5 mm. long. .......-
oe uta egiae Sule Ea skh Seles oe 9. P. reineckei.
Styles and ovary-locules 3-5 (rarely 2); leaves with 1-9 leaflets; cultivated
species (no. 10 probably indigenous in the New Hebrides ).

Leaflets 1-5, the blades broadly elliptic or orbicular, usually 8-22 cm. long
an road, cordate or broadly concave (rarely truncate) at base,
rounded at apex, entire or coarsely crenulate at margin. .........---
I re rr ee en ge iy ay 10. P. scutellaria.

Leaflets 5-9, the blades usually elliptic to oblong, usually 6-12 & 4-8 cm.
(sometimes larger), obtuse to acute at base and apex riage spinu-
lose-dentate at marge i. guilfoylet.

Leaves irregularly pinnate- compound, usually 2 or 3 times div tee bt rarely

simply pinnate (and then with some leaflets pinnatifid or laciniate); culti-
vated species.

Styles and ovary-locules 3-5; leaflets very variable in shape, some usually

elliptic to oblong, the marginal teeth usually 1-3 mm. long. .........---

ib is Sel be bey Ae IE Keg Cat Lib... guilfoylei var. laciniata.

1968 | SMITH & STONE, ARALIACEAE 441

Styles and ovary-locules 2 (rarely 3); ultimate leaflet-divisions usually lan-
ceolate and 3—6 times as long as broad, the marginal sti aie some
: nay vouslly 5-10 mu long so ao eee P. fruticosa.

1, Polyscias joskei Gibbs in Jour. Linn. Soc. Bot. 39: 148. 1909; J. W.
Parham, PI. Fiji Isl. 86. 1964. PL. I, Fics. 1-5.

Botryopanax joskei Hutchinson, Gen. Fl. Pl. 2: 57. 1967.

A tree, glabrous throughout, the branchlets stout (10-15 mm. in dia-
meter toward apices), copiously cicatricose with large obdeltoid petiolar
scars, the leaves and inflorescences congested toward ends of branchlets;
leaves imparipinnate, up to 60 25 cm., the petiole and rachis com-
paratively slender, shallowly canaliculate, the petiole 10-18 cm. long
on mature leaves, slightly swollen at base, the rachis inconspicuously con-
tracted at bases of petiolules, the leaflets 7-15, the petiolules slender,
(2—) 5-20 mm. long, the blades chartaceous to thin-coriaceous, ovate- to
elliptic-oblong, 5—11 cm. long, (2—) 3—5 cm. broad (basal ones somewhat
shorter and broader than middle ones), unequally obtuse to rounded at
base, obtuse to short-acuminate at apex, crenulate at margin, the costa
raised on both surfaces, the secondary nerves 8-18 per side, subspread-
ing, usually prominulous on both surfaces, the veinlet-reticulation in-
conspicuous; inflorescences broadly compound-racemose-paniculate, up
to 20 cm. long and 40 cm. broad, the peduncle less than 3 cm. long, the
primary branches about 8—12, alternate to subopposite or whorled, each
bearing numerous secondary branches 3-8 cm. long, the bracts soon
caducous; flowers borne singly, alternate or loosely whorled, subtended
by lanceolate-subulate bracteoles 0.8-1.2 mm. long, these soon caducous,
the pedicels slender, articulate at apex, scarcely 1 mm. long (or flowers
subsessile) at anthesis, about 2 mm. long in fruit; staminate flowers about
3 mm. long at anthesis, the calyx conspicuously stipitate at base, cupuli-
form distally, about 1 mm. in diameter, obscurely (4- or) 5-lobed;
petals (4 or) 5, oblong, 2—2.5 mm. long, unguiculate at apex; stamens (4
or) 5, the filaments slender, about 0.6 mm. long, the anthers oblong, 1.5—
1.7 mm. long, versatile; stigmas 2, contiguous, forming a minute conical
protuberance on the flat disk; perfect flowers not seen; fruits laterally
compressed, bilocular, transversely oblong-ellipsoid, 4-5 mm. long, 5-6

m. broad, surmounted by the persistent calycular rim and 2 conspicu-
ously divergent styles, these about 1.5 mm. long, the stigmas long-decur-
rent.

TyPE LocaLity: Nandarivatu, Mba Province, Viti Levu, Fiji; the type
is Gibbs 748, cited below. Curiously, this number is not cited by Gibbs,
although the label notes of the British Museum specimen agree with the
data given in the type description, and although the specimen is labelled
by Gibbs and bears a printed label “type specimen.” The numbers cited
in the description are 893 (staminate) and 750 (fruit). We conclude that
these specimens were combined and assigned a new number, 748; this sheet

442 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

has both flowering and fruiting branchlets and may be indicated as the
holotype.

DistTRIBUTION. Endemic to Fiji, and thus far known from Viti Levu,
Ovalau, and Vanua Levu, occurring infrequently at elevations of 575-
1,150 meters in forest or in the dense thickets of ridges and crests. It
has been noted as a tree 3-10 meters high, often compact, with a thin,
pale latex. The distal parts of inflorescence-branches and pedicels are
purple, the petals are yellow, and the fruits are purple with yellow styles.

Locat names. Sole yalewa and nausasa have been noted on Viti Levu,
and ndanindani on Vanua Levu.

i. Virt Levu: Ma: Mt. Evans Range, Greenwood 1249 (A, BISH, UC, US);
nai tacivatic Gibbs 748 (pM type, photo at us), Degener & Ordonez 13578 (A,
BISH, SUVA); slopes of Mt. Nanggaranambuluta, Gillespie 3197 (BISH, GH, NY,
us), 3914 (sisH); ridge between Mt. Nanggaranambuluta and Mt. Namama,
Smith 4975 (a, BISH, K, NY, US); between Nandala and Nukunuku Creeks, along
trail from Nandarivatu toward Lewa, Smith 6161 (A, BISH, US). NAMosI: Sum-
mit of Mt. Naitarandamu, Gillespie 3151 (BisH, GH, UC, Us); Mt. Voma, Fiji

and adjacent ridge, Smith ae eee us). Vanua Levu: MatHuaTA-THA-
KAUNDROVE BOUNDARY: Crest of Korotini Range, between Navitho Pass and
Mt. Ndelaikoro, Smith 568 (BISH, GH, K, NY, UC, US

Polyscias joskei is clearly different from the other species of our area
in having its flowers borne singly on the ultimate inflorescence-branches
rather than in umbels. It may be noted, however, that P. multijuga and
P. reineckei sometimes have a reduced number of flowers per umbel and
rarely only one, but there the single-flowered condition is sporadic and
never consistent for an entire inflorescence. The other characters of these
two species are strikingly different, but the point noted above may suggest
the flexibility of the single-flowered condition.

A closer relative of P. joskei may be P. tahitensis (Nad.) Harms, of the
Society Islands, but the leaflet-blades of the latter are oblong-lanceolate,
proportionately narrower, and entire. The flowers of P. tahitensis appear
to be borne in compact, few-flowered umbels; when these are one-flowered,
the appearance of racemosely borne flowers is simulated.

In his 1967 review of the family, Hutchinson has proposed a new com-
bination for P. joskei in the genus Botryopanax Miq., unaccountably ac-
crediting our species to the Philippine Islands, even though the generic
distribution of Botryopanax is stated as Mauritius and the Mascarenes.
We cannot express an opinion as to the limits of Botryopanax, but, since
it is stated to have 13-16 petals and 8-12 ovary-locules, the reference to
it of a species with 5 petals and 2 ovary-locules is in error. If those species
of Polyscias with solitary (rather than umbellate) flowers are to be segre-
gated, Hutchinson’s new genus Gelibia (in Gen. Fl. Pl. 2: 57. 1967) seems
better suited for them, whether the leaves are simply pinnate or bipinnate
seeming of little consequence. Since Polyscias includes at least one species
with decompound leaves (P. fruticosa), the two species of Gelibia are

1968 | SMITH & STONE, ARALIACEAE 443

hardly separable on that basis. Actually, it is quite evident that P. joskei
in all its basic characters is well placed in Polyscias, and transitional stages
between single flowers and few-flowered umbels are apparent in the genus,
as noted above.

2. Polyscias nusedhul (Guillaumin) B. C. Stone in Taxon 14: 285.
1965

Tieghemopanax nusedhul Guillaumin in Jour. Linn. Soc. Bot. 51: 554. 1938.

Having seen no material of this species, we cannot add to the original
description.

Typr Locatity: Hog harbour, Espiritu Santo, New Hebrides; two
specimens, J. & Z. Baker 13a and 71a, are cited without designation of a
type. These cotypes are deposited in the herbarium of the British Mu-
seum.

Loca NAME. Niisedhul is apparently used generically in this part of the
New Hebrides, since it was also recorded for the species we discuss as
P. excelsa.

On the basis of the description one may feel sure that a species of
Polyscias is here concerned; Guillaumin’s concept of Tieghemopanax in-
cludes T. neo-ebudarum, which is discussed as our next species of Polyscias,
as well as T. fruticosus, a synonym of P. fruticosa. The description indi-
cates that P. nusedhul is related to the common and variable P. multijuga,
of Fiji, Tonga, and Niue, but the New Hebridean plant would appear to
have more obviously inequilateral leaflets that are consistently cordate at
base, shorter petiolules, a less expansive inflorescence, and strictly sessile
flowers about 10 per head. The flowers of P. multijuga are 2-7 per umbel
(rarely solitary) and short-pedicellate (but rarely essentially sessile).

3. Polyscias neo- prey nt ates aaa B. C. Stone in Taxon 14:
285, as P. neo-ebudara. 1 PL. I, Fics. 6-8.

Tieghemopanax neo-ebudarum Guillaumin in Jour. Arnold Arb. 12: 264. 1931.

A tree, glabrous throughout or the young parts and inflorescence-axes
with a sparse indument of 1—3-celled spreading hairs scarcely 0.1
long and evanescent, the branchlets 6-8 mm. in diameter toward apices;
leaves imparipinnate, up to 40 cm. long, the petiole and rachis slender,
subterete, the petiole 8-14 cm. long on mature leaves, slightly swollen at
ase, the rachis obscurely contracted at bases of petiolules, the leaflets
(5— a 9-13, the petiolules slender, 5-10 mm. long, the blades papyraceous,
asymmetric and subfalcate (lateral ones), ovate-lanceolate, 4-9 cm. long,
1.8-3.5 cm. broad, unequally acute at base, acute to obtusely acuminate
at apex, coarsely crenulate at margin, the costa raised on both surfaces,
the secondary nerves 5-8 per side, subascending, prominulous or plane on
both surfaces, the veinlets usually obscure; inflorescences racemose-pani-
culate, up to 30 cm. long, the peduncle very short, the primary branches

444 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

few, subascending, the bracts small, caducous; flowers borne in umbels
on ultimate peduncles 2-10 mm. long, usually 5—12 per umbel, the flower-
subtending bracteoles lanceolate, about 0.5 mm. long, evanescent, the pedi-
cels 1-4 mm. long, articulate at apex; perfect flowers with a narrowly
cupuliform calyx about 2 mm. long, the teeth minute (calyx of staminate
flowers stipitate at base and smaller); petals 5, oblong, 3-3.5 mm. long,
unguiculate at apex; stamens 5, the filaments slender, 1.5-1.7 mm. long,
the anthers ovate-oblong, about 2 mm. long, versatile; stylopodium conical,
about 2 mm. long including the slender stylar column, this minutely bifid
at apex; fruits laterally compressed, bilocular, transversely oblong-ellip-
soid, 3-4 mm. long, 4—4.5 mm. broad, surmounted by the narrow calycular
rim and the stylar column 0.8—1 mm. long, this bifid at apex with 2 stig-
matiferous lobes less than 0.5 mm. long.

Typr LocaLity: New Hebrides. Three specimens were cited by Guillau-
min without indication of a type, and therefore we herewith indicate
Kajewski 291, from Eromanga, as the lectotype; the holotype is pre-
sumably the specimen in the herbarium of the Arnold Arboretum.

DistTRIBUTION. Endemic to the New Hebrides, and thus far known from
Eromanga and Aneityum, where it is indicated as relatively common in
rain-forest at elevations of 60-600 meters. Specimens were taken from
trees 8-12 meters high; the flowers are noted as purple-brown and the
fruit as black at maturity.

Loca NAMES AND USES. Narse has been noted on Eromanga, and
kiviano on Aneityum; on the former island the leaves are used as an ap-
plication for sore ears.

New Hebrides. ERoMANGA: Dillon Bay, May 23, 1928, Kajewski 291 (a type,
BISH). ANeEITYUM: Anelgauhat Bay, Kajewski 749 (a, BisH, us), 977 (coll.
J. P. Wilson) (a, BIsH, Us).

T his species clearly falls into our concept of Polyscias, in which genus
it is closely related only to the Fijian P. culminicola.

4. Polyscias culminicola A. C. Sm. in Contr. U.S. Natl. Herb. 37:
85. 1967. Pi. I, Fics. 9-12.
Although no material additional to the type collection is available,
examination of isotypes permit a slight amplification of the description.
Plant apparently polygamo-monoecious, the central flowers of umbels
sometimes perfect or functionally pistillate (self-fertile?), the ovary en-
larging before the corolla opens; immature fruit 2-locular, compressed-
oblong-ellipsoid, up to 7 X 5.5 mm.; outer (or sometimes all) flowers of
umbels staminate, as originally described.
: Type Locatity: Mba Province, Viti Levu, Fiji; the type is Smith 4514,
cited below.
DISTRIBUTION, Endemic to Fiji, and thus far known from a single lo-
cality in western Viti Levu, where it was found in crest forest on a wind-
swept ridge at an elevation of 750-900 meters. The material is from a tree

1968 | SMITH & STONE, ARALIACEAE 445

8 meters high, with leaves tufted at ends of branchlets and with the in-
florescence terminal on branchlets.

Loca, NAME. Sawira was recorded for the type collection, but this
usually refers to the family Meliaceae and may be incorrectly applied to
a Polyscias.

Fiji. Virt Levu: Ma: Northern slopes of Mt. Namendre, east of Mt. Koro-
mba (Pickering Peak), Smith 4514 (a, BISH, K, US 1965344 type).

Polyscias culminicola is less closely related to P. joskei, with which it
was originally compared, than to the New Hebridean P. neo-ebudarum,
with which it agrees in its umbellate flowers, its comparatively small, fal-
cate leaflets, the indument of its young vegetative parts and inflorescence-
axes (although this is evanescent in P. neo-ebudarum), and its petioles
lacking the basal wings so characteristic of most species of Polyscias (but
also lacking in P. joskei). Foliar and inflorescence characters noted in our
key clearly differentiate P. culminicola and P. neo-ebudarum.

5. Polyscias excelsa (Guillaumin) B. C. Stone in Taxon 14: 285. 1965.
Tieghemopanax excelsa Guillaumin in Jour. Linn. Soc. Bot. 51: 554, 1938.

As we have not seen material of this species, we cannot add to the
original description; however, in view of the other dimensions given it
seems probable that “. . . floribus . . . 30 mm. longis” is an error for 3

Type Locatity: Hog harbour, Espiritu Santo, New Hebrides; the type
is 1, & Z. Baker 13, deposited in the herbarium of the British Museum.
Loca NAME. Niisedhul is recorded with the original description.

It may be noted that one of the Baker numbers referred to P. nusedhul
is 13a, collected at the same locality as no. 13 but on a different date. On
the basis of the descriptions, the two species are amply distinct and are
both clearly referable to Polyscias. Polyscias excelsa also would appear
to be of the relationship of P. multijuga, with which it agrees in its large
inflorescences and few-flowered, compactly arranged umbels. The flowers
of P. excelsa are noted as 6-10 per umbel, whereas in the Fijian species
they are 2~7 or sometimes single. The ovate-falciform, cordate-auriculate
leaflet-blades, from the description, seem to provide a further differentiat-
ing character for P. excelsa.

6. Polyscias corticata Gibbs in Jour. Linn. Soc. Bot.-39: 149. pl. 13,

figs. 14-17. 1909; J. W. Parham, PI. Fiji Isl. 86. 1964.
Pt. II, Fics. 10-14.

A tree or shrub, glabrous throughout; leaves imparipinnate, large, up
to 100 cm. long, the petiole and rachis comparatively stout, subterete or
obscurely canaliculate, the petiole of mature leaves 20-35 cm. long, con-
spicuously swollen to a clasping, lenticellate base and inconspicuously
alate in the proximal 3-5 cm., the rachis geniculate at bases of petiolules,

446 JOURNAL OF THE ARNOLD ARBORETUM [ VoL. 49

the leaflets (9-) 13-17, the petiolules slender, diverse in length, of lateral
leaflets 2-25 mm. and of terminal leaflet to 45 mm. long, the blades
papyraceous, oblong or ovate-elliptic, (9—) 15-30 cm. long, (4—) 6-10 cm.
broad, subequally obtuse to rounded or subcordate at base, abruptly cau-
date-acuminate at apex (acumen 1-2.5 cm. long but fragile), entire or
with a few callose-denticulate crenations, the costa sharply raised above,
prominent beneath, the secondary nerves 8-18 per side, spreading, usually
plane above and prominulous beneath, the veinlet-reticulation coarse,
prominulous beneath; inflorescences racemose-paniculate, up to at least
50 cm. long, the primary branches diverse in length, up to 40 cm. long,
all the inflorescence-axes griseo-corticate-lenticellate, the primary bracts
oblong-lanceolate, 1-2 cm. long (those at base of inflorescence to 4 cm.
long), at length caducous, the umbel- and flower-subtending bracts and
bracteoles oblong, 1-3 mm. long; flowers borne in umbels on ultimate
peduncles 10-20 mm. long, usually 5-15 per umbel, the pedicels 2-6 mm.
long, articulate at apex; perfect flowers 5-6 mm. long at anthesis, with
a cupuliform calyx 2—2.5 mm. long, the rim undulate or obscurely 5-denticu-
late; petals 5, deltoid-oblong, 2.5-3.5 mm. long, unguiculate at apex;
stamens 5, the filaments slender, about 1 mm. long, the anthers oblong,
1.5—2 mm. long; disk concave or flat, the styles 2, free nearly to base,
erect or subspreading, 1—1.5 mm. long, with decurrent stigmas; staminate
flowers similar but somewhat smaller; fruits laterally compressed, bilocular,
transversely ellipsoid, about 4 * 6.5 mm., substipitate at base, sur-
nine y the persistent calycular rim and the obviously divaricate
styles.

Type LocaLiry: Mba Province, Viti Levu, Fiji; the type is Gibbs 769,
cited below.
DistrIBUTION, Endemic to Fiji, and thus far known from the largest
islands, Viti Levu and Vanua Levu, where it occurs in usually dense
forest at elevations of 300-900 meters. Specimens have been indicated
as slender trees or shrubs, unbranched or few-branched, and 1.5—4 meters
high; the inflorescences are axillary among leaves congested toward apices
of branchlets. The calyx is purplish or reddish; the petals green to white,
but becoming rich purple without at anthesis; the anthers pale yellow
to white; and the styles yellowish.

Locat NAMES. Sole and ndravi have been noted on Viti Levu, and
ndanindani on Vanua Levu.

Fiji. ViT1 Levu: Mpa: Nandarivatu and vicinity, Gibbs 769 (pm type, photo
at us), Degener 14458 (A, BISH, K, NY, UC, US), 14806 (A, BISH, K, NY, UC, US),
14834 (A, BISH, K, NY, UC, US); between Nggaliwana and Nandala ‘Creeks, south
of Nauwanga, Smith 5804 (a, BISH, K, NY, US). VANUA LEvVU: THAKAUNDROVE:
Southern slopes of Korotini Range, below Navitho Pass, Smith 510 (BISH, NY);
southwestern slope of Mt. Mbatini, Smith 611 (BIsH, GH, K, NY, UC, Us).

The Fijian P. corticata and P. multijuga and the Samoan P. samoensis
form a group of related species, but differences among them are obvious.
Polyscias corticata is readily distinguished by having its inflorescence-axes

1968 | SMITH & STONE, ARALIACEAE 447

pale and with congested transverse lenticels that provide a curiously flaky
appearance. In type of inflorescence P. corticata is closer to P. multijuga
than to P. samoensis, but in length of pedicels and of umbel-peduncle it
is intermediate between them. The petiolar wings of P. corticata are less
conspicuous than in either of its two relatives.

7. Polyscias multijuga (A. Gray) Harms in E. & P. Nat. Pflanzenfam.
3(8): 45. 1894; Yuncker in Bishop Mus. Bull. 178: 92. 1943, in op.
est. 220: 207. 1959. PL, . FIGS. 1-9.

Paratropia multijuga A. Gray, Bot. U.S. Expl. Exped. 1: 722.

ie co multijugum Seem. Fl. Vit. 115. pl. 18, 19. 1865, in agian Bot. 4:
295.

Panax nya B. & H. f. Gen. Pl. 1: 938. 1867.

Panax multijugum B. & H. f. ex Drake, Ill. Fl. Ins. Mar. Pac. 181. 1890;
Hemsl. in Jour. Linn, Soc. Bot. 30: 180. 1894.

A tree, glabrous throughout; leaves imparipinnate, large, up to 100 cm.
long, the petiole and rachis stout, subterete or slightly flattened above,
the petiole of mature leaves 10-35 cm. long, conspicuously swollen to a
clasping, often lenticellate base and obviously alate in the proximal 7—20
cm. with subcoriaceous wings, the rachis geniculate at bases of petiolules,
the leaflets 13-25 (—29), the petiolules stout, diverse in length, of lateral
leaflets 5-50 and of terminal leaflet to 60 mm. long, the blades papyrace-
ous to thin-coriaceous, variable in size and shape, oblong to oblong- or
ovate-elliptic, (9—) 12-27 cm. long, (3—-) 4-12.5 cm. broad, subequally
cordate to rounded or obtuse at base, subacute to cuspidate or acuminate
at apex (acumen to 1.5 cm. long), entire and narrowly recurved at margin,
the costa sharply raised above, prominent beneath, the secondary nerves
6-18 per side, spreading, prominulous on both surfaces or plane above,
the veinlets usually plane on both surfaces; inflorescences compound-
racemose-paniculate, up to 150 cm. long, composed of 3-8 panicles arising
from a stout central rachis, the primary branches stout, bearing numerous
irregularly disposed (alternate or opposite or whorled) secondary branches
5-18 cm. long at maturity, the primary bracts often subcoriaceous, ovate-
oblong, 4-20 mm. long, obtuse or acute, the umbel- and flower-subtending
bracts and bracteoles 0.5—2 mm. long; flowers borne in umbels on very
short ultimate peduncles 1-6 mm. long (or umbels essentially sessile),
(1-) 2-7 per umbel, the pedicels 0.5—1 mm. long (or flowers sometimes
essentially sessile) ; perfect (or functionally pistillate) and staminate
flowers sometimes occurring in the same umbel, differing only in the
slightly smaller calyx of the latter; perfect flowers with a cupuliform calyx
1-1.3 mm. long, the rim undulate or obscurely 5-denticulate; petals 5,
oblong-ovate, 2-3 mm. long, unguiculate at apex; stamens 5, the filaments
slender, minute, the anthers oblong, 0.8—1.2 mm. long; disk slightly con-
cave, the styles 2 (rarely 3?), suberect and contiguous in flower, usually
1—1.5 mm. long, divergent at least distally in fruit, with decurrent stigmas;
fruits laterally compressed, bilocular (rarely trilocular ex Seemann),

448 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

transversely ellipsoid, up to 5.5 X 6 mm., surmounted by the persistent
calycular rim and styles, shallowly costate when dried.

Type tocatity: Mbua Bay (“Sandalwood Bay’), Mbua Province,
Vanua Levu, Fiji; the type is the U.S. Exploring Expedition specimen
low.

DistRIBUTION. Fiji, Tonga, and Niue, occurring very abundantly at
least in Fiji at elevations from near sea-level to 1,100 meters, and re-
ported as a component of many types of forest or second-growth habitats.
Specimens have been obtained from trees 2—15 meters high, usually noted
as slender, sometimes unbranched, and with trunks up to 20 centimeters
in diameter; the inflorescence is terminally borne on branchlets with a
cluster of leaves. Color notes indicate the buds to be dark red, the petals
and stamens greenish white to pale yellow, and the fruit purple at maturity.

Locat Names. In Fiji the names ndanindani and sole are general for

wala nimbernggua. In Tonga the names tanetane and kulukulu have been
recorded, and on Niue tanetane and tanetane vao.

Fiji (because of abundance only island and province are cited). YASAWAS:
Waya: St. John 18104 (sisH, us). Virt Levu: Mpa: Parks 20532 (BisH, UC),
Gillespie 4295 (BisH), Greenwood 297 (Kk), 297a (A, uc), 935 (A, BISH), Smith
4065 (A, BISH, K, US), 4837 (A, BISH, US). arenas & Navosa: Degener
15264 (A, BISH, K, NY, UC, US), Smith 5480 (A, BISH, K, NY, US). SERUA: Degener
15137 (A, BISH, K, Ny, UC, US), Smith 9178 (BIsH, sou ‘Fiji Dept. Agr. 12453
(BISH, sUva). NaMmost: Seemann 205 (BM, GH, K), Gillespie 2860 (BISH, UC),
2898 (BISH), 3120 (BISH, GH, UC) Smith 8509 (BISH, us), Fiji Dept. Agr. 5902
(suva). Ra: Degener 15499 (a, BISH, K, NY, UC, US). Natrasiri: Gillespie 3428
(BISH, GH, UC), St. John 18246 (BISH, Us), Fiji Dept. Agr. 10640 (suvA). REWA:
Gillespie 2210 (BISH, GH, NY). KANDAVU: Smith 122 (BISH, GH, K, NY, UC, US).
Ovatau: Graeff » pie Smith 937 aiid GH, K, ee us). NGAU:

ssi
OncEA Levu: Bryan 427 (pisH, us). Fiji WITHOUT
OTHER LOCALITY: Harvey (GH, K), Horne 189 (x).

Tonga. Toncatapu: Near caves of Anahulu, Setchell & Parks 15278 (UC);
near Hufagalupe, Setchell & Parks 15625 (uc); near Kologa Point, Setchell &
Parks 15385 (BISH, NY, UC); near Houma, Yuncker 16227 (BISH, GH, US). "EUA:
Lister (kK, photos at BIsH, Us), Hotta 5434 (BISH).

Niue: Near Alofi, Yuncker 9791 (BISH), 10062 (BISH, uc); near Mutalau,
Yuncker 9717 (A, BISH),

Polyscias multijuga is one of the most frequent small trees of the Fijian
forests, and at one time it was probably equally abundant in Tonga and
Niu

Niue. It is characterized by large, coarse leaves and inflorescences, of
which the umbels are few-flowered and short-pedunculate. The stout

1968 | SMITH & STONE, ARALIACEAE 449

petiole of P. multijuga bears obvious, subcoriaceous wings that extend up-
ward from the base 7~—20 centimeters.

8. Polyscias samoensis (A. Gray) Harms in E. & P. Nat. Pflanzenfam.
3(8): 45. 1894, in Bot. Jahrb, 25: 663. 1898; Rechinger in Denkschr.
Akad. Wiss. Wien 85: 323. 1910; Setchell in Carnegie Inst. Publ.
341: 61. 1924; Christophersen in Bishop Mus. Bull. 128: 165. 1935.

PL. I. Fics. 15-17.

Panax samoense A. Gray, mies U.S. Expl. Exped. 1: 717. 1854; Drake, Iil.
Fl. Ins. Mar. Pac. 182.

Nothopanax Samoense ck mr es 116. 1865, in Jour. Bot. 4: 295. 1866;
Powell in Jour. Bot. 6: 366.

A shrub or tree, glabrous ae leaves imparipinnate, large, per-
haps up to 100 cm. long, the petiole and rachis comparatively slender,
subterete, the petiole of mature leaves up to 20 cm. pan swollen to a
clasping base and narrowly alate in the proximal 5-8 cm., the rachis
geniculate at bases of petiolules, the leaflets 11-17, the atiolites slender,
of lateral leaflets 8-30 mm. and of terminal leaflet to 35 mm. long, the
blades papyraceous, oblong- to ovate-elliptic, (7—) 9-21 cm. long, 4-8.5
cm. broad, subequally subcordate to rounded or obtuse at base, gradually
attenuate to a slender acumen 1-2 cm. long, entire or undulate and re-
curved at margin, the costa slightly raised above, prominent beneath, the
secondary nerves 6-10 per side, curved-ascending, with the intricate vein-
let-reticulation prominulous on both surfaces; inflorescences compound-
paniculate, ample, up to at least 50 * 25 cm., the axes comparatively
slender, the peduncle up to 18 cm. long, the primary branches numerous,
spreading, irregularly disposed on the rachis, sometimes alternate but
more often aggregated in loose whorls at 2—4 nodes including a terminal
one, each primary branch bearing umbels at apex or again divided like
the rachis, the primary bracts caducous (presumably small), the umbel-
and flower-subtending bracts and bracteoles lanceolate, 1-3 mm. long;
flowers borne in umbels on conspicuous slender ultimate peduncles 12-
80 mm. long, 8-15 per umbel (sometimes mixed with additional umbels),
the pedicels slender, 7-15 mm. long at anthesis and inconspicuously ar-
ticulate at apex, elongating to 25 mm. in fruit; flowers (apparently uni-
formly hermaphrodite) slender, 6-7 mm. long a anthesis, the calyx urceo-
late, 3-3.5 mm. long including the obvious undulate or 5-denticulate rim;
petals 5, lanceolate-ovate, 3-3.5 mm. long, unguiculate at apex; stamens
5, the filaments slender, less than 0.5 mm. long, the anthers oblong, 1.5—
1.7 mm. long; disk flat, the styles 2, erect and contiguous in flower,
SPECAGIA nearly from base in fruit and strongly recurved, elongating to

1.5 mm., with decurrent stigmas; fruits laterally compressed, bilocular,
suborbicular, 7-8 mm. long and geal Na coed nerved, surmounted
by the obvious calycular rim and sty

TYPE LocaALity: Samoa, indicated as Savaii and Tutuila; the type is
the U.S. Exploring Expedition collection cited below.

450 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

DistrIsuTION. Endemic to Samoa, and known from the three largest
islands, occurring at elevations up to about 400 meters, presumably in
forest. Specimens have been taken from trees or shrubs up to 7 meters
high; the flowers and fruits are reported as dark purple in color, and the
fruits have conspicuous reddish or purplish veins.

LocaL NAMES AND USES. Tanitani is probably the name in general use,
but Setchell notes afia as the name on Tutuila and indicates that oil from
the plant is used on the hair. It is interesting to note that Powell (in
Jour. Bot. 6: 278. 1868) lists afia as the name for Ascarina lanceolata,
but indicates afia-vao for an unidentified shrub that, from his brief note,
could represent Polyscias.

Samoa. “Savaml AND Tutuma”: U.S. Expl. Exped. (GH, us 47923 type).
Upotu: (No material seen but reported by Harms and Rechinger). TUTumLA:
Leone, Wilder 87 (nist); Pago Pago, Wilder 44 (or 244) (BisH); above Pago
Pago, near wireless station, Setchell 255 (coll. Sutupe) (GH, uc, US); between
Fagaalu and Fatimafutu, Setchell 374 (sisH, uc). SAMOA WITHOUT OTHER LO-
CALITY: Powell 21 (kK), 36 (K), 47 (kK, photos at BISH, US), 159 (K, photos at
BISH, US), 265 (K, photos at BISH, Us), s.n. (K, photos at BISH, US), Whitmee
97 («K), 197 (K, photos at BISH, US).

We have also seen a specimen from Aneityum, New Hebrides, obtained
by an unknown collector in February 1859 (Bm), which suggests this
species. However, it is a very incomplete specimen, suggesting P. samoen-
sis in its flowers but with smaller and strongly cordate leaflet-blades. It
is noted here as a possibly undescribed relative of P. samoensis, which
may safely be inferred to be a Samoan endemic.

From its relatives, P. corticata and P. multijuga, P. samoensis differs
in its much more lax and open inflorescences, of which the comparatively
few branches are whorled at a few nodes and umbellately aggregated
apically. The flowers, borne on conspicuous pedicels in long-pedunculate
umbels, are longer than those of the related species, with a conspicuously
urceolate calyx, and the fruits are larger and very conspicuously nerved.

9. Polyscias reineckei Harms in Bot. Jahrb. 25: 663. 1898; Rechinger
in Denkschr. Akad. Wiss. Wien 85: 323. 1910; Christophersen, Bishop
Mus. Bull. 128: 165. 1935;(?) Yuncker in Bishop Mus. Bull. 184:
56. 1945. Pt. ISI, rics. 1-6.

A tree, glabrous throughout; leaves imparipinnate, large, perhaps up to

100 cm. long, the petiole comparatively stout, of mature leaves 15-25 cm.

long, swollen to a clasping base and narrowly alate in the proximal 10—

14 cm., the rachis more slender, subterete, geniculate at bases of petiolules,

the leaflets (9—) 11 or 13, the petiolules slender, of lateral leaflets 5—40

and of terminal leaflet to 60 mm. long, the blades papyraceous, oblong-

to ovate-elliptic, (S—) 10-25 cm. long, (3—) 4-11 cm. broad unequally
rounded to obtuse at base, gradually attenuate to a slender sharp acumen

1-2 cm. long, entire or remotely spinulose-denticulate and narrowly re-

curved at margin, the costa plane or shallowly canaliculate above, prom-

inent beneath, the secondary nerves 7—10 per side, curved-ascending,

1968 | SMITH & STONE, ARALIACEAE 451

prominulous on both surfaces, the veinlet-reticulation intricate, plane or
prominulous; inflorescences compound- -paniculate, ample, sometimes ex-
ceeding 100 cm. in length, composed of a few (2-4?) elongate primary
branches arising from a comparatively short peduncle and rachis, the
branches pendent, long-pedunculate, each bearing about 6-12 elongate
alternate lax secondary branches, the primary bracts lanceolate, 15-30
mm. long, soon caducous, the secondary bracts 3-10 mm. long, the umbel-
and flower-subtending bracts and bracteoles 0.5—2 mm. long; flowers borne
in (sometimes imperfect) umbels on slender ultimate peduncles (of di-
verse length) 1-20 (—40) mm. long, (1—) 3-8 per umbel, the pedicels
slender, (0.5—) 1-3 mm. long at anthesis and inconspicuously articulate
at apex, scarcely elongating (1-5 mm.) in fruit; flowers (apparently
uniformly hermaphrodite) 4-4.5 mm. long at anthesis, the calyx cupuli-
form, 1.5—2 mm. long, the rim short, inconspicuously 5-denticulate; petals
5, deltoid-ovate, 2-3 mm. long, unguiculate at apex; stamens 5, the fila-
ments ligulate, less than 1 mm. long, the anthers oblong, about 1.5 mm.
long; stylopodium broadly conical, surmounted by 2 sessile laterally con-
tiguous stigmas; fruits laterally compressed (or triquetrous?), bilocular
(sometimes trilocular ex Harms), suborbicular or transversely ellipsoid,
5-6 6-8 mm., surmounted by the calycular rim, the stylopodium pro-
jecting less than 1 mm. and terminated by strictly sessile stigmas.

TYPE Locatity: Letogo Ridge, Upolu, Samoa; the type is Reinecke
136, collected in February 1894, and presumably destroyed in the Berlin
herbarium.

DIstRIBUTION. Endemic to Samoa, and thus far known definitely only
from Savaii and Upolu, where it occurs at elevations of near sea-level to
700 meters in open forest, coastal forest, etc. Plants of the species are
reported as trees 4-12 meters high; no color notes are available. Four
additional specimens from Upolu were listed by Rechinger in 1910.

Locat NAME. Afia has been noted on Savaii.

Samoa. Savatr: Salailua, Christophersen & Hume 2616 (A, BISH, US); be-
tween Salailua and Lataitai, Christophersen 3010 (A, BISH); between Falelima
and Samata, Christophersen 2764 (A, BISH); Matavanu, near crater, Christo-
phersen & Hume 1951 (BISH, NY, US); above sili, Christophersen 3280 (Bish,
us). Upotu: Graeffe (BM).

An additional record of P. reineckei, on the island of Tau, Manua Is-
lands, was noted by Yuncker in 1945, but this record was based on an
observation by Paul L. Guest and is not supported by herbarium material.
In view of the superficial similarity of this species and P. samoensis, the
record must be questioned, especially as the latter is known from nearby
Tutuila and the former is not.

Although in foliage P. reineckei is not easily distinguished from P.
Samoensis, there are excellent inflorescence-differences between these two
Samoan endemics. The primary inflorescence-branches of P. reineckei
bear alternate, elongate branches with racemosely disposed umbels, where-
as the inflorescence-branches of P. samoensis are usually whorled and often

452 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

again branched in a pseudo-umbellate manner. In both species the pe-
duncles of the ultimate umbels are very diverse in length, but in P. rein-
eckei they are usually the shorter. The flowers of P. reineckei are uni-
formly short- pedicellate, the pedicels not exceeding 3 mm. in length at
anthesis and 5 mm. in fruit, whereas P. samoensis has pedicels 7-25 mm.
long. The flowers of P. samoensis are notably longer, and the stylar dif-
ference is particularly noteworthy, P. samoensis having obvious styles
strongly recurved in fruit, and P. reineckei having its stigmas sessile on a
broadly conical stylopodium even in mature fruit.

It may be noted that in P. reineckei (e. g. in Christophersen & Hume
2616) the flowers may sometimes be only pseudo-umbellate, actually
racemosely arranged toward the apices of the ultimate peduncles, which
are freely bracteolate along much of their length. Thus we have another
type of transition between umbellate and individually borne flowers, indi-
cating that this character is not dependable even at the specific level
with exactitude, and certainly not at the generic or tribal level.

10. foe var scutellaria (Burm. f.) Fosberg in Univ. Haw. Occ. Pap.
46: 9. 1948; B.C. Stone in Taxon 14: 284. 1965.
Pr: T, ries. 15, 14%:

Scutellaria prima Rumph. Herb. Amb. 4: ee - 31. 1743.

Crassula scutellaria Burm. f. Fl. Ind. 78.

Polyscias pinnata J. R. & G. Forst. he ae Pl. 64. pl. 32. 1776; Forst.
f. Fl. Ins. Austr. Prodr. 90, 1786; Seem. a Jour. Bot. 3: 180. 1865; Guil-
laumin in Bull. Soc. Bot. France 66: 270. 1919; Harms in Bot. Jahrb. 56:
409. 1921; Guillaumin in Bull. Soc. Bot. France 74: 698. 1927, in Bull.
ae Hist. Nat. II. 9: 289. 1937, in Jour. Linn. Soc. Bot. 51: 554. 1938;

B. C. Stone in Taxon 14: 282. 1965.

Aralia cochleata Lam. Encycl. Méth. 1: 224. 1783.

Panax conchifolium Roxb. Hort. Beng. 21, nomen. 1814, Fl. Ind. ed. 2. 2:
832.

Panax scutellaroides Reinw. in Bl. ee ne 1826.

Panax cochleatum DC, Prodr. 4: 253.

Panax heyneanum Wall. Cat. n. 127, nomen. 1832; G. Don, Gen. Syst. 3:
385. 1835; Walp. Rep. 2: 429. 1843.

Panax forsteri Dec. & Pl. in Rev. eee IV. 3: 105. 1854.

Nothopanax Pn ags Mig. in Bonplandia 4: 139. May, 1856, Fl. Ind. Bat.
1(1): 766. Sept.

Nothopanax passers Mia. FI. ae sen Suppl. 1: 340, 1861; Merr. In-
terpret. Rumph. Herb. Amb. 409, 19

Panax rumphii Hassk. in Abh. Naturf. Halle 9: 220. 1866

Panax pinnata Baill. Hist. Pl. 7: 197. fig. 205. 1879. Non Lam. (1788) nec
A. Rich. (1847).

Nothopanax scutellarium Merr. Interpret. Rumph. Herb. Amb. 409. 1917.

Polyscias tricochleata Fosberg in Phytologia 5: 290, 1955

A shrub or small tree, glabrous throughout; leaves imparipinnate (often
unifoliolate), the petiole usually 10-30 cm. long, swollen to a clasping
base and narrowly alate in the proximal 1-5 cm., the rachis geniculate at

1968 | SMITH & STONE, ARALIACEAE 453

bases of petiolules (or usually jointed in unifoliolate leaves), the leaflets
1,3, or 5 (less often 2 or 4), the petiolules slender, of lateral leaflets (0.5—)
1—5 and of terminal leaflet to 6 cm. long, the blades papyraceous to sub-
coriaceous, normally green but often with white margins or areoles,
broadly elliptic or orbicular, usually 8-22 (—26) cm. long and broad (but
often smaller in cultivated plants), cordate or broadly concave (rarely
truncate) at base, rounded at apex, subentire or coarsely crenulate at
margin (crenations 1—2 cm. distant and sometimes conspicuously spinulose,
or the leaflets sometimes subpalmately lobed), the costa raised above and
prominent beneath, the secondary nerves 5-8 per side, spreading, prominu-
lous on both surfaces or prominent beneath, the veinlet-reticulation intri-
cate, plane or slightly prominulous on both surfaces; inflorescences com-
pound-paniculate, up to 60 cm. long, the peduncle short, to 3 cm. long, the
primary branches numerous, up to 45 cm, long, alternate toward base of
inflorescence but often opposite or verticillate distally, the secondary
branches similarly arranged, the primary bracts ovate, acute, to 1 cm.
long, soon caducous, the umbel- and flower-subtending bracts and brac-
teoles ovate to lanceolate, 0.5-1.5 mm. long; flowers borne in umbels
on slender ultimate peduncles 4-15 mm. long, 8-26 per umbel, the pedicels
slender, 1.5-6 mm. long, articulate at apex; flowers 3-3.5 mm. long at
anthesis, the calyx cupuliform, 1-1.5 mm. long and about 2 mm. in
diameter at anthesis, the rim short, inconspicuously denticulate with 5-8
(rarely 9) teeth; petals 5-7 (sometimes 8, rarely 4 or 9), narrowly ovate-
elliptic, 1.8-2.5 mm. long, unguiculate at apex; stamens as many as petals,
the filaments slender, 0.5—1 mm. long, the anthers oblong, 1.3-1.7 mm.
long; disk concave, the styles 3-5 (rarely 2), contiguous in an oblong-
subconical column but free, 0.4-0.8 mm. long, rounded at apex, ventrally
sulcate; fruits subglobose, costate, up to 5 mm. long and broad, sur-
mounted by the persistent calycular rim and the eventually divaricate
styles.

TYPE Ltocatity: Amboina; the basis for Crassula scutellaria Burm. f.
is the Rumphian pl. 37. Merrill in his 1917 work points out that this
was cited as pl. 30 by Burman, as it also is by Fosberg in 1948. Of the
complex synonymy cited above, most combinations are based either on
Burman’s name or on Aralia cochleata Lam. (also based on the Rumphian
plate) and have been elucidated by Merrill, Fosberg, and Stone. The
other basic synonym is Polyscias pinnata J. R. & G. Forst., typified by ma-
terial collected in Tanna, New Hebrides, on Cook’s second voyage. This
material was labelled as collected by Forster and by Anderson, although
the specimens may well have come from the same plant. Since the
specimen with Anderson’s name bears flowers and immature fruits as
well as leaves, it is herewith indicated as the lectotype and is so cited
below

DistriBUTION. Widespread in cultivation, but presumably indigenous
in the New Hebrides and the Solomon Islands, where it occurs in forest
from near sea-level to about 300 meters. The available specimens are
from shrubs or trees up to 5 meters high, but in cultivation the plants

“

454 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

are often pruned into hedges, in which condition they flower infrequently.
Such flowers as are available are indicated as pale yellow-green.

LocaL NAMES AND USES. On Espiritu Santo, New Hebrides, the names
néwesi and niiesi have been noted, and on Efate nanlass, in Samoa tagi-
tagi, and in Tonga tanitani. It is recorded that on Malekula, New He-
brides, the leaves of this plant are used to rub the gums with the intent
of numbing the nerves prior to the extraction of teeth.

New Hebrides. Esprritu Santo: Hog harbour, J. & Z. Baker 64 (BM), 261
(sm). MALeKuLA: Northwestern part, Cheesman 27 (xk, photos at BISH, US);
without further locality, Herre 69b (Ny). EraTe: Onesua, Stone 2251 (BISH).
Tanna: Anderson (BM lectotype, photo at us), Forster (K, photos at BISH, US).

Samoa. UpoLu: Mulinu’u, near Apia, McKee 3024 (BIsH).

In 1965 (in Taxon 14: 281-285) the second author discussed the
typification of Polyscias and the identity of its type species, P. pinnata
J. R. & G. Forst., indicating that several widely distributed horticultural
forms are closely related to and perhaps derived from P. pinnata. One
of these, P. scutellaria, was discussed in detail because of the tenuous
nature of characters that separate it from P. pinnata. It was concluded
that P. pinnata could be maintained as distinct on the basis of having its
leaves generally 3- or 5-foliolate rather than 1-foliolate, its petals and
stamens 7 or 8 rather than 4 or 5, and its styles 4 or 5 (whereas the styles
of P. scutellaria are sometimes only 3, but also often 4 or 5). A factor
in a decision to maintain both concepts as species was the fact that P.
scutellaria, based on a binomial of 1768, would have priority if the taxa
were merged.

However, on reconsideration and examination of more ample material,
we are unable to recognize any morphological discontinuities that are in
any sense usable to divide this complex into species. Individual plants
often have the leaflets either 1, 3, or 5, and the correlation of number
of petals and stamens with number of styles is haphazard. The most
common numbers of petals and stamens are 5, 6, and 7, and of styles 3
or 4. One specimen from Bougainville, Solomon Islands (Kajewski 1965,
A, BISH), has the petals and stamens 5 or 6 and the styles only 2; its
leaves are 1- or 3-foliolate. The Kajewski specimen came from a rain-
forest tree and was apparently indigenous, and only its reduced number
of styles separate it from the presumably indigenous New Hebridean
specimens cited above.

fe must conclude, therefore, that the forest species of the Solomons
and New Hebrides, P. pinnata, is not separable from the widely culti-
vated P. scutellaria at any rank. It is unfortunate that the epithet of
the type species of the genus is the later one.

The above description is drawn from available specimens, both from
within and outside our area. It may be noted that specimens cultivated
in Samoa and Hawaii are usually inclined to resemble the wild form sug-
gested by the type of P. pinnata, with 3 or 5 leaflets; Micronesian and
Philippine specimens usually resemble the typical and presumably cul-

1968] SMITH & STONE, ARALIACEAE 455

tivated form of P. scutellaria, with unifoliolate leaves. Fosberg, however,
in making the appropriate combination i in 1948, pointed out that various
specimens cultivated in Hawaii show all variations from pinnate to uni-
foliolate leaves.

In his consideration of the Micronesian species of Polyscias in 1965 (in
Micronesica 2: 51-59), the second author again stressed the close re-
lationship of P. pinnata and P. scutellaria. In the same paper he de-
scribed a chimaeric variation of P. tricochleata that is essentially in-
distinguishable from P. pinnata, and therefore concluded that Nothopanax
tricochleatum Miq. is merely a cultivated form of P. pinnata. This
conclusion is herewith reinforced, but in view of our reduction of the
Forsters’ binomial to synonymy the correct designation of this particular
form is P. scutellaria cv. Tricochleata. The cultivar is frequent in Micro-
nesia and doubtless other tropical areas and is characterized by having
leaves smaller than those of the indigenous form (of P. pinnata) or the
normal cultivated form (of P. scutellaria), usually variegated and 3- or
5-foliolate, each leaflet being often further 3-divided and with coarsely
crenate or spinulose-serrate margins. In our area this cultivar was
reported by Yuncker (in Bishop Mus. Bull. 178: 92. 1943, in op. cit.
184: 56. 1945) as a presumable variety of P. guilfoylei. We note P.
scutellaria cv. Trichochleata from our area as follows:

Fiji. Vitt Levu: Rewa: Vicinity of Suva, Degener & Ordonez 13542 (A).

Samoa. TuTurLa: Pago Pago, Bryan 1002 (sisH). Tau: Siufaga Village,
Yuncker 9212 (sisu).

Niue. Alofi, oie 10051 (BIsH).

11. gig guilfoylei (Bull) L. H. Bailey in Rhodora 18: 153.

A fc or small tree, glabrous throughout, usually with suberect vir-
gate branches; sai imparipinnate, up to 50 cm. long, the petiole
slender, 10-17 ¢ m. long on mature leaves, clasping at base and incon-
spicuously alate in the proximal 1-2 cm., the rachis often shallowly
canaliculate, geniculate at bases of petiolules, the leaflets 5-9, the petio-
lules slender, of various length, the blades papyraceous, green or varie-
gated with white margins or areolae, variously shaped and incised ; in-
florescences compound-paniculate to verticillate-umbellate, the primary
branches 5-10, up to 60 cm. long, spreading from the very short pe-
duncle and rachis, the secondary branches numerous, alternate or oppo-
site or often verticillate, the primary bracts deltoid-lanceolate, acute,
about 1 cm. long, soon caducous, the umbel- and flower-subtending
bracts and bracteoles ovate, 0.5—2 mm. long; flowers borne in umbels
on slender ultimate peduncles 5—50 mm. long (these often bracteolate
and geniculate at 1 or 2 nodes), 10-25 per umbel, the pedicels slender,
3-10 mm. long, articulate at apex; flowers with a cupuliform calyx 1.2-
2 mm. long and 2—2.5 mm. in diameter at apex, the rim short, undulate,
denticulate with 5 or 6 inconspicuous teeth; petals 5 or 6, deltoid-oblong,
2-2.5 mm. long, unguiculate at apex; stamens as many as petals, the

456 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

filaments slender, 0.5—-1 mm. long, the anthers oblong, 1-1.3 mm. long;
disk concave, the styles 3-5, at first contiguous in a conspicuous col-
umn 0.8-1 mm. long, soon spreading and jong to 1.5 mm., the
stigmas shortly decurrent; mature fruits not seen

Our description of the species and of the varieties is based on many

available specimens, from both within and outside our area. The im-
mediate relationship of P. gwilfoylei is open to conjecture, and indeed it
may have originated as a sport. Among Pacific species it is suggestive
of P. scutellaria (including P. pinnata) in inflorescence characters, but
the differences in foliage are striking and are summarized in our key
to species.
Polyscias ert je is perhaps the most widely cultivated taxon in the
genus, as plants may readily be pruned into vigorous hedges. Several
horticultural ‘athe have been discussed, characterized by degree of
leaf-division, variegation, etc. Three such forms are treated as botanical
varieties by L. H. Bailey (Stand. Cycl. Hort. 2747-2748. 1925 and
later editions). It is doubtful whether these forms merit any designa-
tion other than “cultivar,” but a definitive review of Polyscias in culti-
vation is outside the scope of this treatment. An additional cultivated
form was recently discussed but not named by the second author (in
Micronesica 2: 57. fig. 4. 1965); this has dark green, crumpled, bullate,
nearly orbicular leaflets. Specimens of it have been noted from Hawaii
as well as Micronesia, but not in the area of the present treatment.

For the time being we retain Bailey’s varietal concept for a form often
grown in the Pacific and elsewhere, with deeply laciniate leaflets. The
two forms occurring in our area may be thus distinguished:

Leaflets elliptic to oblong, spinulose-dentate at margin. ........ r. guilfoylei.
Leaflets deeply and irregularly divided or decompound, the oltnente parts
variously incised: and meniae (2 var. laciniata.

lla. Polyscias guilfoylei var. guiloylei.
— guilfoylei Bull, Cat. 1873; Cogn. & March. Pl. Ornament. 2: pl. 58.
874.

Nothopanaz guilfoylei Merr. in Philip. Jour. Sci. Bot. 7: 242. 1912; Christo-
phersen in Bishop Mus. Bull. 128: 165. 1935; Yuncker in Bisho op Mus.
Bull. 220: 207. 1959; J. W. Parham, PI. Fiji id, 83. 1964.

Polyscias guilfoylei L. H. Bailey in Rhodora 18: 153. 1916; Setchell in Car-
— Inst. Publ. 341: 61. 1924; Yuncker in Bishop Mus. Bull. 178: 91

43.

The type-including variety; leaflets with petiolules (6—-) 10-20 mm.
long (up to 50 mm. on terminal leaflet), the blades variable in shape but
most often elliptic to oblong, (5—) 6-12 (-17) cm. long, (2.5—) 4-8
(—14) cm. broad, obtuse to acute at base, obtuse to mucronately acute
at apex, conspicuously and often irregularly spinulose-denate at margin
(teeth 1-3 per centimeter), the costa sharply raised on both surfaces, the
secondary nerves 5—9 per side, subascending, nearly straight, prominulous

1968 | SMITH & STONE, ARALIACEAE 457

on both sides, the veinlet-reticulation intricate, prominulous on both sur-
faces or plane abo

TYPE LocaLiry: Unknown, but indicated in Bull’s Catalogue as “South
Sea Islands.” It is quite possible that Bull received his material from
W. R. Guilfoyle, whose interesting account of “A botanical tour among
the South Sea Islands” (in Jour. Bot. 7: 117-136. 1869) indicates that
Samoa, Tonga, Fiji, the New Hebrides, and New Caledonia were visited
by Guilfoyle on H.M.S. Challenger in 1868. It is mentioned that culti-
vated plants of “Aralia” were observed in Samoa (p. 119) and the New
Hebrides (p. 133). That plants of horticultural interest were brought
back is indicated by Guilfoyle’s final sentence: ‘‘. . . I have been success-
ful enough to bring with me in good condition, I have no hesitation in
saying, the largest collection of choice and beautiful plants ever yet col-
lected in the islands of the South Pacific.” It is doubtful that any type
specimen has been preserved.

DistripuTion. Widely cultivated throughout the Pacific and American
tropics, and probably in all warmer areas. Plants grow very readily from
cuttings. Although comparatively few flowering specimens are available,
inflorescences will probably mature if the plants are not severely pruned.
When not cut back, plants develop into shrubs or smal] trees with ascend-
ing virgate branches, up to 6 or 7 meters in height. It is probable that
the species has become naturalized in thickets near settlements in parts
of its cultivated range.

LocaAL NAMES AND USES, Tanitani and many variant spellings have
been recorded in Samoa, Niue, and Tonga, and ndanindani in Fiji; these
names are more or less generic but apply especially to the cultivated
forms. As elsewhere noted, this species is widely cultivated as a hedge
plant, the abundance of the typical variety not being suggested by the few
available collections.

Samoa. TutumLa: Setchell 297 (coll. Sutupe) (uc).

Niue: Near Alofi, Yuncker 9933 (BISH).

11b. Polyscias guilfoylei var. laciniata (Hort.) L. H. Bailey in
Rhodora 18: 153. 1916; Christophersen in Bishop Mus. Bull. 128:
165. 1935; Yuncker in Bishop Mus. Bull. 178: 92. 1943.

enh laciniatus Hort. in Gard. Chron. 1876: 735. 1876, in op. cit. 1880: 759.

A variety differing from var. guilfoylei in its more finely divided leaves,
the petiolules often to 60 mm. long, some of the blades deeply lobed or
decompound into irregular divisions, the ultimate leaflets variously in-
cised and laciniate with numerous teeth up to 5 mm. long.

Type Locatity: Unknown; the original material brought into culti-
vation may have come from the Pacific area together with that of the
typical form.

DistrIBuTION. Widespread in cultivation; in our area it seems to be
more abundant than var. guilfoylei.

458 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

LocaL NAMES AND USES. Not distinguished from the typical form in
these respects. Seemann (FI. Vit. 115. 1865) indicates that in Fiji a
juice from the bark was used as a remedy for ulcerated tongue and
throat.

Fiji. Without wana ss Seemann 204 (BM, GH, K, photos at BISH, US),
Horne 588 (K, photos at BISH, US).

Wallis Islands. pe E. Home in 1866 (BM

Samoa. AUNUU ISLAND (east of Tutuila): Diefenderfer 1239 (BISH). SAMOA,
without further locality: U.S. Expl. Exped. (GH); Powell 36 (xk, photos at BISH,

(BISH).
Tonga. ToNGATAPU: Setchell & ‘earls 15418 (uc).

It should be noted that Seemann cited his own and the Home specimen
listed above as Nothopanax fruticosum; his other comments, however, ap-
pear to refer to the following species in our treatment, where his refer-
ence is listed in the synonymy.

Although in inflorescence and basic foliage characters this variety
clearly belongs in P. guilfoylei, we have keyed it with P. fruticosa because
its variable leaves are often decompound. Dissected leaflets usually oc-
cur on the same plants (or even in the same leaves) as simple leaflets,
and the texture and margins of the leaflets are obviously suggestive of
those of typical P. guilfoylei.

12. Polyscias fruticosa (L.) Harms in E. & P. Nat. Pflanzenfam. 3(8) :
45. 1894; Rechinger in Denkschr. Akad. Wiss. Wien 85: 323. 1910;
Harms in Nova Guinea Bot. 8: 275. 1910, in Bot. Jahrb. 56: 412.
1921. Pu. III, Fics. 7-9.

Scutellaria tertia Rumph. Herb. Amb. 4: 78. pl. 33. 1743

Panax fruticosum L. Sp. Pl. ed. 2. 1513. 1763; A. Gray, Bot. U.S. Expl. Exped.
1; 716. 1854; Drake, Ill. Fl. Ins. Mar. Pec 181. 1890; Warburg in Bot.
Jahrb. 13: 396. 1891; Hemsl. in Jour. Linn. Soc. Bot. 30: 180. 1894; Guil-
laumin in Bull. Soc. Bot. France 66: 270. 1919, in op. cit. 74: 698. 1927.

Nothopanax fruticosum Mig. in Bonplandia 4: 139. May, 1856, Fl. Ind. Bat.
1(1): 765. Sept. 1856; Seem. FI. Vit. 115. 1865, in Jour. Bot. 4: 294. 1866;
Merr. Interpret. Rumph. Herb. Amb. 410. 1917: Yuncker in Bishop Mus.
Bull. 220: 207. 1959; J. W. Parham, Pl. Fiji Isl. 83. 1964.

Tieghemopanax fruticosus Viguier in Ann. Sci. Nat. IX. Bot. 4: 61. 1906;
Guillaumin in Jour. Arnold Arb. 12: 263. 1931, in Jour. Linn. Soc. Bot. 51:
554. 1938; J. W. Parham, Pl. Fiji Isl. 86. 1964.

A shrub or perhaps a small tree, glabrous throughout; leaves irregularly
pinnate-compound, usually 2 or 3 times divided but sometimes simply
pinnate, up to 50 cm. long, the petiole slender, 5—14 cm. long on mature
leaves, clasping at base and inconspicuously alate in the proximal 1-3 cm.,
the rachis subterete, obscurely geniculate at bases of petiolules, the pri-
mary leaflets usually 9-13, with slender petiolules 1-6 cm. long and
usually imparipinnately once or twice divided, rarely merely pinnatifid
or laciniate or deeply serrate, the ultimate leaflet-divisions papyraceous,

1968 | SMITH & STONE, ARALIACEAE 459

usually lanceolate and 3-11 x 0.5-3 cm. (rarely to 20 X 5 cm.), nar-
rowed to an attenuate base, long-acuminate at apex, irregularly lobed or
laciniate or spinulose- dentate at margin, the costa prominulous on both
surfaces, the secondary nerves short, spreading, with the veinlet-reticula-
tion essentially plane; inflorescences compound-paniculate to verticillate-
umbellate, the primary branches 5-12, up to 30 cm. long, spreading or as-
cending from a short, stout peduncle and rachis, the secondary and tertiary
branches often irregularly verticillate, the primary bracts lanceolate-ob-
long, 0.5—4 cm. long, soon caducous, the umbel-subtending bracts ovate,
1-2 mm. long, the flower-subtending bracteoles minute; flowers borne in
umbels on slender ultimate peduncles 10-20 mm. long, 8-30 (—40) per
umbel, the pedicels slender, 1-5 mm. long, articulate at apex; flowers
3—4 mm. long at anthesis, the calyx cupuliform, 1-1.7 mm. long and
1.5—2 mm. in diameter at the minutely 5-denticulate rim; petals 5, obo-
vate-oblong, 2-3 mm. long, acute; stamens 5, the filaments slender, 1—1.2
mm. long, the anthers oblong, 1.5—2 mm. long, versatile; disk concave,
the styles 2 (rarely 3), at first contiguous into a slender conical column
1-1.2 mm. long; fruits compressed and bilocular (rarely trigonous and
trilocular), 4-5 x 5-6 mm., surmounted by the obscure calycular rim
and divergent styles, these to 1.5 mm. long and with long-decurrent
stigmas

Type LocaLity: Amboina; the basis for Panax fruticosum is the Rum-
phian p/. 33, doubtless drawn from a cultivated plant

DistTRIBUTION. Widely cultivated throughout the Pacific and Ameri-
can tropics, and probably elsewhere; also popular for greenhouse culti-
vation. The species is commonly grown as an ornamental shrub or in
hedges, attaining a height of 1-3 meters if not pruned. It flowers fre-
quently and has white petals and styles. It may be noted that Kajewski
931, cited below, from the New Hebrides, is said to have been taken
from a rain-forest tree 8 meters high; this could possibly be an indica-
tion of a native habitat, or it could merely represent a naturalization,
since the species is freely cultivated in the New Hebrides.

LocAL NAMES AND USES. Tanitani or a variant is applied in Polynesian
areas; ndanindani in Fiji. Names noted in the New Hebrides are niiluh
(Espiritu Santo) and taconerecott (Efate, ex Guillaumin). Kajewski
notes that the leaves are boiled and eaten on Aneityum. The species is,
of course, a popular ornamental.

Hebrides. Esprriru Santo: Tungwi, J. & Z. Baker 156 (BM). TANNA:
ae gegen 63 (A, NY). ANEITYUM: Anelgauhat Bay, Kajewski 931 (a,
BISH, Us).

FG Os Levu: sine Tumbenasolo, valley of Namosi Creek, Smith 4724
(A, assole US). Au: Lovoni village, Smith 7496 wae US).

Ton a. TONGATAPU: ae 1518 (K, photos at BISH, US

ae description is based on many specimens from other parts of the
cultivated range as well as those cited. We have not seen the Samoan
specimens cited by Rechinger. It should be noted that a form with
much smaller leaves, not flowering in our observation, has been collected

460 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

in Micronesia and in Hawaii, but not within our area. This form and
others doubtless have horticultural designations.

The true relationship of P. fruticosa is not obvious. It is quite probably
a horticultural variant of one of the species of Polyscias with two styles
and ovary-locules, but its derivation from any such species in our area
(species 1 to 9 in the key) seems very unlikely. Although P. fruticosa is
not immediately related to P. guilfoylei, having 2 (rarely 3) rather than
3-5 styles and ovary-locules, the finely divided forms of P. guilfoylet,
such as var. laciniata, may not be readily distinguished from P. fruticosa
in sterile condition. As a rule, the leaflets of P. guilfoylei var. laciniata
are even more variable and irregular than those of P. fruticosa, but they
are usually accompanied by at least a few leaflets suggestive of the typical
form (elliptic to oblong), and the marginal teeth are comparatively short.

4. Reynoldsia A. Gray, Bot. U.S. Expl. Exped. 1: 723. 1854, in Ann.
Sci. Nat. IV. Bot. 4: 177. 1855, in Proc. Am. Acad. Arts Sci. 3: 128.
1857; Seem. in Jour. Bot. 2: 244. 1864; Harms in E. & P. Nat.
Pflanzenfam. 3(8): 30. 1894; Hutchinson, Gen. Fl. Pl. 2: 58. 1967.

Reynoldsia was based on two original species, R. sandwicensis A. Gray,
of Hawaii, and R. pleiosperma A. Gray, of Samoa. Both species are illus-
trated and the generic description would appear to have been based on
both. We are not aware of an earlier designation of a lectotype species
than that of Hutchinson (in 1967), who so denotes R. sandwicensis.
Reynoldsia is a genus of at least 7 and perhaps 17 or more species, oc-
curring in Samoa, the Societies, the Marquesas, and Hawaii. In the lat-
ter archipelago Sherff (Bot. Leafl. 6: 7-19. 1952) recognizes 8 species,
which we do not attempt to evaluate.

In Samoa three species were recognized by Christophersen in his 1935
treatment, and we herewith add a fourth. The species are probably
endemic to various islands: to Savaii 2, to Upolu 1, and to the Manua
Islands 1. It must be acknowledged that none of the Samoan species
are adequately known and that comparably mature parts are not avail-
able for all of them. Nevertheless, good differentiating characters are
evident in the shape and margin of leaflet-blades, the inflorescence-type
(as to complexity of branching and arrangement of flowers), the length
of umbel-peduncles and pedicels, and the size of inflorescence-bracts.

KEY TO THE SPECIES

Inflorescences paniculate, not umbelliferous, the ultimate branches bearing flow-
ers (and fruits) irregularly or in whorls of 2-5 in a few clusters and apically;
leaflet-blades lanceolate, subentire, inconspicuously bau? denticulate at
MATBIN, ee ee ee . R. pleiosperma.

Inflorescences umbelliferous, the flowers 5-12 in umbels, not eae singly.

Leaflet-blades conspicuously dentate at margin with widely spaced callose-
obtuse teeth; umbel-peduncles 1.5—3 cm. long, oe by bracts 2-4
mm. long, the pedicels 7-9 mm. long. . R. lanutoensis.

1968 | SMITH & STONE, ARALIACEAE 461

Leaflet-blades inconspicuously undulate and rarely remotely and minutely
callose-denticulate at margin.

Inflorescences racemose-umbellate, the umbels few, borne singly on the up-
per part of the inflorescence, the umbel-peduncles about 7 cm. long,
simply umbelliferous at apex, the pedicels 15-20 mm. long; leaflet-
blades broadly ovate, 12-14.5 & 6-8 cm. ............ . R. grayana,

Inflorescences racemose-verticillate, the umbels laxly verticillate at 2 or 3
nodes, the umbel-peduncles, 1.5-3 cm. long, sometimes with a secondary
whorl of flowers below the ultimate umbel; umbel-subtending bracts of
proximal nodes conspicuous, 5-15 mm. long; leaflet-blades ovate-
lanceolate,.7=12 6 3-5 CBs eis ae os ee 4. R. tauensis.

1. Reynoldsia pleiosperma A. Gray, Bot. U.S. Expl. Exped. 1: 725.
1854, Atlas pl. 93, pro parte fructifera. 1857, in Ann. Sci. Nat. IV.
Bot. 4: 178. 1855, in Proc. Am. Acad. Arts Sci. 3: 129. 1857; Seem.
in Jour. Bot. 2: 245. 1864; Harms in Bot. Jahrb. 25: 664. 1898;
Christophersen in Bishop Mus. Bull. 128: 160. 1935.

Eschweileria pleiosperma Dur. ex Drake, Ill. Fl. Ins. Mar. Pac. 183. 1890.
Trevesia pleiosperma B. & H. f. ex Jackson, Ind. Kew. 2: 1100. 1895.

A large tree, glabrous throughout; leaves imparipinnate (or perhaps
sometimes even-pinnate by loss of the terminal leaflet), large, up to 90
cm. long, the petiole and rachis subterete, the petiole of mature leaves
20-40 cm. long, swollen to a clasping base and angled in the proximal
1.5-3 cm. (if alate, the wings soon lost), the leaflets 10-15, the petiolules
slender, 5-20 mm. long, the blades papyraceous or subcoriaceous, lan-
ceolate, 8-17 cm. long, 3-6 cm. broad, unequally rounded to broadly
obtuse at base, gradually acuminate at apex (acumen 7-15 mm. long,
slender, obtuse, marginally recurved), undulate-denticulate at margin
(teeth, if present, inconspicuous, about 1 mm. long, callose-recurved),
the costa raised above, prominent beneath, the secondary nerves 10-15
per side, spreading, prominulous on both surfaces, the veinlet-reticulation
intricate, plane on both surfaces; inflorescences paniculate, up to 35 cm
long, not umbelliferous, the peduncle and rachis subterete, stout (3-5
mm. in diameter), the peduncle 13-17 cm. long, the secondary branches
12-17, alternate or irregularly whorled (with 4-6 borne apically), 4-13
cm. long, bearing flowers irregularly or 2—5 loosely whorled at 2 or 3 nodes
and apically, the primary bracts (subtending secondary branches ) oblong-
lanceolate, 2-5 mm. long, caducous, the flower-subtending bracteoles
about 1 mm. long, evanescent, the pedicels 5-10 mm. long ( to 15 mm. in
fruit); calyx cupuliform, 4-5 mm. long and in diameter, with a flaring
undulate or irregularly lobed rim about 1 mm. long; petals (?8-) 10 or
11, narrowly oblong-lanceolate, at anthesis 6-7 mm. long and 1-2 mm.
broad, unguiculate at apex, at length strongly reflexed and introrsely
costate; stamens (?8—) 10 or 11, the filaments carnose, at length 4-5
mm. long, the anthers oblong, about 3 mm. long, versatile; stylopodium
laterally flattened, about 1 mm. high, with (?16-) 20 or 22 congested
radiating stigmas; fruits compressed-subglobose, sulcate, about 5 mm.

462 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

long and 6 mm. broad, surmounted by the calycular rim and the truncate-
conical stylar column 1-1.5 mm. high bearing linear stigmas radiating
on its slightly concave apex, the pyrenes (?16—) 20-22.

TYPE LocaLity: Savaii, Samoa; the type is the U.S. Exploring Expedi-
tion collection cited below. Christophersen (in Bishop Mus. Bull. 128:
161. 1935) has pointed out that Gray’s original material came from two
species, which were kept apart in the description but combined in the
original plate; he has designated the fruiting specimen as the type, a
choice here followed.

DistRIBUTION. Samoa, and thus far known with certainty only from
Savaii, where Christophersen indicates it to be a common tree of con-
siderable size in forests at 700-1,700 meters altitude. It attains a height
of more than 10 meters and its trunk has a diameter of 70—200 centimeters.

LocAL NAME AND USES. Vi vao is the name recorded by Christophersen,
who states that the fleshy fruits make it a favorite tree for pigeons and
consequently well known to natives.

Samoa. Savatt: Le To, above Salailua, Christophersen 2935 (BisH); above
Letui, Christophersen 772 (a, BISH); Olo, above Safotu, Christophersen &
Hume 2319 (BisH, us); Mt. Maugaloa, Vaupel 463 (BISH, US); Savaii, without
further locality, U.S. Expl. Exped. (us 62432 type, GH fragm.). Samoa, with-
out other locality: Whitmee 97 (xk, photos at BISH, US).

Reynoldsia pleiosperma remained the only Samoan species of the genus
recognized until 1925, when Hochreutiner described R. lanutoensis from
Upolu, redescribed and discussed below. In 1935 Christophersen clarified
the situation caused by the mixed materials in the original Exploring Ex-
pedition collection, indicating the non-type portion as a new species, R.
grayana. We agree that this separation is warranted, and below we pro-
vide a first description for Christophersen’s species and indicate points of
difference between it and R. pleiosperma.

a Reynoldsia lanutoensis Hochreutiner in Candollea 2: 482. 1925;
Christophersen in Bishop Mus. Bull. 128: 160. 1935.

A tree, glabrous throughout; leaves imparipinnate or sometimes even-
pinnate, large, up to 55 cm. long, the petiole and rachis slender, sub-
terete, the petiole of mature leaves 16-22 cm. long, swollen to a clasping
base and narrowly alate in the proximal 1-2 cm. (wings free in the distal
1-2 mm.), the leaflets 10-13, the petiolules slender, of lateral leaflets
3-13 mm. and of terminal leaflet to 23 mm. long, the blades papyraceous,
ovate- or oblong-lanceolate, (4.5—) 6-12 cm. long, (2—) 2.5—5.5 cm. broad,
rounded to broadly obtuse at base, acuminate at apex (acumen 5-10
mm. long, obtuse, marginally recurved), conspicuously dentate at margin
(teeth widely spaced, 3-5 per side, callose-obtuse and marginally re-
curved), the costa plane or canaliculate above, prominent beneath, the
secondary nerves 10-15 per side, spreading, with the intricate veinlet-
reticulation usually plane above, prominulous beneath; inflorescences
paniculate-umbellate, up to 20 cm. long, the primary axis 7-14 cm. long,

1968 | SMITH & STONE, ARALIACEAE 463

the secondary branches few, to 12 cm. long, bearing umbels whorled at
several nodes (or umbels sometimes solitary), the umbel-subtending
bracts lanceolate, 2-4 mm. long, entire or with a few marginal teeth, the
ultimate peduncles 1.5-3 cm. long; flowers 5-10 per umbel, subtended
by ovate-deltoid bracteoles about 1 mm. long, the pedicels slender, 7-9
mm. long, not articulated; calyx about 3 mm. long and 4 mm. broad,
with a subtruncate or undulate rim; petals (?10—) 12, 4.5-5 mm. long;
stamens (?10—) 12, the filaments short, the anthers oblong, about 2 mm.
long, versatile; stylopodium laterally flattened, 1-1.5 mm. high, with
20-24 congested radiating stigmas; fruits compressed-globose, sulcate,
4-5 mm. long, 5—7 mm. broad, surmounted by the truncate-conical, bi-
lobed, flattened stylar column bearing linear stigmas, the pyrenes 20-24.

Type Locatity: Lake Lanuto, Upolu, Samoa; the type is Hochreutiner
3270 (deposited at Geneva), collected March 22, 1905, in forest at an
altitude of about 670 meters. We have not seen this collection, but the
original description leaves no doubt of its identity.

DIsTRIBUTION. Samoa, and thus far known only from Upolu, where
it occurs in forest or open grassland at altitudes of 650-750 meters. It has
been noted as a tree 6-10 meters high, with green fruit.

Locat NAME. Vi vao has been recorded by Christophersen.

. Upotu: Near Malololelei, Christophersen 951 (A, BISH, US); above
Malololelei, ridge to Mt. Vaitou, Christophersen 262 (BIsH, US); summit of Mt.
Fao, Christophersen 551 (BISH).

From the other three species now known from Samoa, R. lanutoensis
differs in having its leaflet-blades conspicuously dentate with widely
spaced teeth. Although the actual teeth are small, callose-obtuse, and
marginally recurved (and not very different from those of R. pleiosperma
and the other Samoan species), they surmount undulations some 2—5 mm.
deep, the blades thus appearing more coarsely dentate than in the related
species. While the flowers of R. lanutoensis are clearly umbellate, a sec-
ond whorl of flowers is sometimes borne on the peduncle just below the
apical umbel, suggesting that the mode of flower-bearing in this species
and in R. pleiosperma is not basically very diverse. Nevertheless the two
species are readily separable, at least on the basis of the collections cur-
rently available.

3. Reynoldsia grayana Christophersen in Bishop Mus. Bull. 128:
161. 1935.
Reynoldsia pleiosperma A. Gray, Bot. US. Expl. Exped. 775. Dp. 1654,
Atlas pl. 93, pro parte alabastro et foliacea. 1857.

A tree, glabrous throughout: leaves imparipinnate, large, probably more
than 40 cm. long, the petiole and rachis subterete, the leaflets at least 7,
the petiolules slender, of lateral leaflets 8-15 and of terminal leaflet to
20 mm. long, the blades broadly ovate, (6—-) 12-14.5 cm. long, (3-—)
6-8 cm. broad, rounded or truncate-obtuse at base, acute to gradually

464 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

acuminate at apex, undulate or obscurely denticulate at margin (teeth,
if present, 2 or 3 per margin, callose-obtuse, scarcely 1 mm. long), the
costa raised above, prominent beneath, the secondary nerves 8-15 per
side, spreading, prominulous on both surfaces, the veinlet-reticulation
intricate, immersed or impressed on both surfaces; inflorescences race-
mose-umbellate, up to 22 cm. long, the peduncle at least 10 cm. long, the
secondary branches few (about 5), about 7 cm. long, simply umbelliferous
at apex; flowers 5 or 6 per umbel, the pedicels comparatively stout (1-2
mm, in diameter), 15-20 mm. long, gradually swollen distally, not articu-
lated; calyx obconical, 3-4 mm. long and in diameter, with an erect un-
dulate rim; petals 8—10, narrowly oblong-lanceolate, about 1.5 mm. broad,
4-5 mm. long and calyptrate in bud but doubtless free at anthesis, unguicu-
late at apex; stamens 8—10, the filaments carnose, about 2 mm. long in bud,
the anthers oblong, 2—2.5 mm. long in bud, versatile; stylopodium truncate-
conical, laterally flattened, about 2 mm. in diameter at the slightly con-
cave apex, the stigmas about 18, linear, radiating.

TYPE LocaALity: Savaii, Samoa; the type is the U.S. Exploring Expedi-
tion collection cited below.

DIsTRIBUTION. Samoa, known only from the type material. It is indi-
cated by Pickering (Geogr. Distr. Animals and Plants 2: 294. 1876) that
the Exploring Expedition Samoan material included only one collection
referable to Reynoldsia. For that reason one may assume that both parts
of the mixture treated by Gray and illustrated in the Atlas were indeed
Rees by the forest-king from Interior Savaii,’ to use Pickering’s
words.

Samoa. Savaii, without further locality: U.S. Expl. Exped. (us 73913 type,
GH fragm.).

As discussed under R. pleiosperma, we follow Christophersen in his
choice of the fruiting material available to Gray as the type of that
species, the balance serving as the type of P. grayana. The latter species
has not previously been described, but Christophersen referred to a por-
tion of Gray’s earlier comments and to part of an illustration, thus seem-
ing to have met the letter of the International Code. The Exploring Ex-
pedition specimen deposited as R. pleiosperma in the U.S. National Her-
barium includes poorly preserved material which quite clearly came from
two different plants. We have now separated this material as two her-
barium sheets, one of which may serve as the holotype of Gray’s species
and the other as the holotype of Christophersen’s.

Reynoldsia grayana differs from R. pleiosperma in having its inflores-
cences clearly umbelliferous, the umbels being long-pedunculate and the
pedicels much longer. The leaflet-blades of R. grayana are broadly ovate
and about twice as long as broad, those of R. pleiosperma being lanceo-
late and about three times as long as broad. From R. lanutoensis, the
present species differs very obviously in the shape and margins of its
leaf-blades, in its simpler inflorescences, long umbel-peduncles, and long
pedicels. On the basis of material now available, R. grayana seems to

1968 | SMITH & STONE, ARALIACEAE 465

have fewer petals and stamens than either R. pleiosperma or R. lanutoensis,
but we consider this variation of minor significance.

4. Reynoldsia tauensis A. C. Smith & B. C. Stone, sp. nov.
Sth Sais sp. Christophersen in Bishop Mus. Bull. 128: 161. 1935; Yuncker
Bishop Mus. Bull. 184: 55. 1945.

Arbor ubique glabra; foliis ad 60 cm. longis videtur paripinnatis, petiolo
et rhachidi subteretibus, petiolo 22-25 cm. longo ad basim amplectentem
incrassato et 1-2 cm. anguste alato, rhachidi petiolulorum basi contracta,
foliolis 10-12, petiolulis gracilibus 5-15 mm. longis, laminis papyraceis
ovato-lanceolatis, 7-12 cm. longis, 3-5 cm. latis, basi subaequaliter ro-
tundatis, apice in acuminem obtusum margine recurvatum breviter cuspi-
datis, margine anguste recurvatis cartilagineis inconspicue undulatis et
dentibus callosis haud 1 mm. longis raro et remote denticulatis, costa
supra paullo elevata subtus prominente, nervis secundariis utrinsecus 10—
15 supra planis subtus prominulis, rete venularum intricato supra im-
merso subtus conspicuo plano; inflorescentiis apices ramulorum versus
infra folia congestis, juvenilibus ad 15 cm. longis racemoso-verticillatis,
pedunculo gracili ad 9 cm. longo, umbellulis 5-8 in nodis 2 vel 3 laxe
verticillatis, bracteis umbelliferis papyraceis anguste lanceolatis 5—15 mm.
longis (superioribus interdum 2-3 mm. longis) margine minute serrulatis,
umbellularum pedunculis gracilibus 1.5—3 cm. longis interdum verticillum
secundarium gerentibus; floribus 8-12 per umbellulam pedicellatis, pedi-
cellis in alabastro 2-3 mm, longis non articulatis, bracteolis floriferis
oblongo-deltoideis 1-2 mm. longis acutis; calyce turbinato circiter 2.5 mm.
longo et lato, limbo erecto ad 1 mm, longo undulato et irregulariter 3-5-
denticulato; petalis 8-10 in alabastro oblongo-deltoideis valvatis apice
unguiculatis; staminibus 8-10; stylopodio in alabastro vadose concavo,
stigmatibus 16—18 sessilibus fadiatis,

TYPE Locaity: Tau, Manua Islands, Samoa; the type is Garber 752.

DistrIBUTION. Known only from the type collection and probably
endemic to the Manua Islands; at least Paul L. Guest indicated to
Yuncker (oc, cit. 1945) that the species is abundant on all the (Manua)
islands. The only documented collection was taken from a tree about
3 meters high at an altitude of 685 meters. ;

Locat NAME. No name was indicated on the original label, but Yuncker
(loc. cit. 1945) has listed vi vao, which apparently is applied to the genus
in Samoa.

‘Au, Manua Islands: Top of peak back of Tau, January 28, 1922,
Garber 752 (BIsH type).
Both Christophersen and Yuncker implied that the cited Tau specimen

represents an undescribed species; even though it lacks mature flowers
In having the inflorescence com-

fering in its essentially entire rather than conspicuously dentate leaflet-

466 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

blades and in its comparatively large umbel-subtending bracts. From R.
grayana the new species differs in its more complex inflorescence, its
short umbel-peduncles, its presumably shorter pedicels, and its ovate-
lanceolate, proportionately narrower leaflet-blades.

wn

. Plerandra A. Gray, Bot. U.S. Expl. Exped. 1: 729. 1854, in Ann.
Sci. Nat. IV. Bot. 4: 178. 1855, in Proc. Am. Acad. Arts Sci. 3: 129.
1857; Seem. in Jour. Bot. 2: 241. 1864, Fl. Vit. 117. 1865; Harms
in E. & P. Nat. Pflanzenfam. 3(8): 28. 1894; A. C. Sm. in Bishop
Mus. Bull. 141: 116. nas in mts Arnold Arb. 36: 286. 1955;
Hutchinson, Gen. Fl. Pl. 2:

Bakeria Seem. in Jour. Bot. 2: 248. 1864, Fl. Vit. 117. 1865, oe cit. 429. 1873.
Nesopanax Seem. in Jour. Bot. 2: 249. 1864, Fl. Vit. 116.

Plerandra, typified by the Fijian P. pickeringii, is believed to include
about 14 species occurring in New Guinea, the Solomons, and Fiji; it has
not yet been discovered in the New Hebrides. Seven endemic species
terminate its range in Fiji

The Fijian species of Plerandra have been puzzling, but the reasonably
adequate herbarium material now available indicates that they are well
differentiated. They are probably all polygamo-dioecious, or perhaps
sometimes polygamo-monoecious, and consequently there have been mis-
interpretations of floral dimensions and of the shape of the stylopodium,
since these differ in staminate and perfect flowers. There is remarkable
variation in the number of stamens from 15 in P. bakeriana to as many
as 500 in P. pickeringii, and in the number of styles, stigmas, and ovary-
locules from 5 to 17. While such numbers do not always permit rigid
specific delimitation, they are usable within limits. Seemann’s genus
Bakeria, based on the species now known as P. bakeriana, does not con-
sistently have 15 stamens and 5 ovary-locules as originally supposed by
him; the stamens are actually 15-20 and the ovary-locules and stigmas
are 5-8, indicating that even this distinct species cannot be separated
from Plerandra. As a matter of fact, in 1873 Seemann himself emended
Bakeria to note that its stamens could be as many as 23, but we have not
observed more than 20. Seemann’s genus Nesopanax, based on the species
currently known as P. vitiensis, was said to differ from Plerandra in having
its petals free rather than calyptrate, and by having 5—7 ovary-locules
and free styles. In fact, however, in P. vitiensis the ovary-locules vary
from 5 to 11; all the Plerandrae have eventually separating petals,
whether or not they are connate in bud; and the styles of P. vitiensis

ecome proximally connate in the stylopodium of the fruit.

The most distinct species is perhaps P. imsolita, with short-pedicellate
flowers congested on the distal portion of short rays and scarcely umbel-
late. Also very distinct is P. pickeringii, with the largest number of sta-
mens (200-500) and with a very conspicuously elongated stylopodium of
firmly connate styles. Plerandra bakeriana is readily distinguished by its
reduced number of stamens, but otherwise it is not strikingly different

1968 | SMITH & STONE, ARALIACEAE 467

from several other of our species. Plerandra grayi is likely to be confused
with P. vitiensis but is actually reasonably distinct on the basis of its
numerous stamens (as many as 250), its large flowers, and its broad
leaflets with obviously rounded apices. The remaining three species form
a related group, but characters referring to stamen-number, fruit shape,
stylopodium, and number and size of leaflets are useful in recognizing
them.

KEY TO THE SPECIES

Inflorescences composed of umbels, the flowers borne at the apices of elongate
rays rarely less than 5 cm. long and usually much longer.

Stylopodium inconspicuous, in fruit 1-2.5 mm. long, the styles sometimes
completely or partially separate, sometimes connate with marginally pro-
jecting stigmas.

Styles and ovary-locules 5~11; stamens not more than 150; flowers com-
paratively small, the petals 4-7 mm. long; leaflet-blades rarely more
than 10 cm. broad.

Stamens usually 1-seriate, 15-20; styles and ovary-locules usually 5 or
6 (rarely 7 or 8); flowers small, the petals about 4 mm. long; fruits
10-12 * 8-10 mm., the stigmas projecting marginally from the trun-
cate-concave apex of the stylopodium; leaflets 5-8 (-10)...........

ong.
Leaflets 3-6 (-7); fruits oblong-ellipsoid, about 20 xX 10 mm., the
stigmas obviously separate on the truncate-concave apex of the
inconspicuous stylopodium; ovary-locules 5-8; stamens 50-75. ue

2 grandiflora.

mee af
Leaflets 5-10 (rarely 4-12); fruits ellipsoid, not much longer than
b m

composed of connate styles and minutely concave at apex; le
lets usually 5-7, the blades 8-15 X 3.5-7 cm. .. 3. P. victoriae.
Stamens 3-—5-seriate, 75-155; ovary-locules usually 7-10 (some-
times 5-11); stylopodium in fruit composed of styles connate
proximally but free at the spreading or erecto-patent apices;

leaflets usually 7-10, the blades usually 13-23 X 4-9 cm. ....
P. vitiensis.

11-20 10-15 mm., the stylopodium com
connate into a centrally concave ring; leaflet-blades usually ne
6-14.5 cm., broadly rounded at apex. ..... sees wethee : :
Stylopodium obvious, in fruit 5-10 mm. long, conical-cylindric, composed of
9-17 firmly connate styles with stigmas marginal on the truncate apex;
stamens numerous, 200-500; flowers large, the petals “

16 mm. long;
P. pickeringii.

f
Inflorescences lacking true umbels, the
cm. long) in a strobilus-like pseudo-

75; flowers comparatively large, the petals 7-8 mm. long; fruits large, 25-

468 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

-30 mm., with a persistent calycular rim and a a (1-2 m
long) foes the 9-12 styles firmly connate. ........ Be ak Bo

1. Plerandra bakeriana A. C. Sm. in Bishop Mus. Bull. 141: 118.
1936; J. W. Parham, PI. Fiji Isl. 84. fig. 34. 1964.
Pu. Vil, Fies.3;-4

Bakeria — Seem. in Jour. Bot. 2: 249. fig. (p. 248). 1864, FI. Vit. 117.

pl. 21. ; Anon. in Gartenfl. 36: 71. 1887, in Kew Bull. 1888: 95. 1888.

BS Satara pricks B. & H. f. ex Drake, Ill. Fl. Ins. Mar. Pac. 183. 1890;
rm E. & P. Nat. Pflanzenfam. 3(8): 29. 1894; non Baill. (1880).

A small tree, glabrous throughout; leaves clustered near ends of branch-
lets, digitately compound, the petioles subterete, up to 25 cm. long, ex-
panded at base into a coriaceous sheath 10-15 mm. broad, the ligule
broadly ovate, as much as 7 mm. long, thinner at margin, the leaflets 5—8
(—10), the petiolules 1-4 (—6) cm. long, the blades subcoriaceous, elliptic-
obovate, usually 8-17 cm. long and 4-7 cm. broad, acute to attenuate at
base, subacute to broadly obtuse at apex, the costa prominent beneath,
the secondary nerves usually 7-12 per side, inconspicuously raised on
both surfaces, the veinlet-reticulation immersed; inflorescences umbellate,
composed of 8-15 umbels radiating from a stout peduncle 1—2.5 cm. long,
the rays 4.5-23 cm. long, the subtending bracts soon caducous, the
flowers often 15-40 (but sometimes as few as 10) per umbel, on pedicels
1,5-3 cm. long; staminate flowers with a short calyx 2.5-3 mm. long,
flaring and sinuate at margin; calyx slightly larger in perfect flowers,
petals 5, thick, deltoid-ovate, acute, unguiculate, about 4 mm. long; sta-
mens usually 1-seriate and often 15, sometimes as many as 20, about 2
mm. long, the anthers oblong, obtuse, about 1.5 mm. long; disk concave;
stigmas minute, like the locules usually 5 or 6, sometimes 7, rarely 8;
fruits ellipsoid to subglobose, inconspicuously costate when dried, usually
10-12 >< 8-10 mm., the stylopodium 1-2 mm. long, the stigmas project-
ing as obtuse, inconspicuous, marginal protuberances on the truncate-
concave apex, the pyrenes lunulate.

TYPE LocaLity: Namosi Province, Viti Levu, Fiji; the type is Seemann
209, cited below.

DisTRIBUTION. Known only from the island of Viti Levu, Fiji, and
there infrequently collected. Insofar as data are available, the species
occurs in ridge and crest thickets at 900—1,100 meters, but the Seemann
and MacGillivray & Milne specimens may come from lower elevations.
Field notes are inadequate, but the plant is recorded as a small tree about
5 meters high, the fruit being black at maturity.

Locat NAME. Sole has been noted by Parham, but this is essentially a
generic name in Fiji.

Fiji. Vit1 Levu: Mba: Mt. Evans Range, Greenwood 382 (K, photos at BISH,

us). NaMosi: Mt. Naitarandamu, Gillespie 3153 (BIsH, GH, NY, UC); Kor
mbasambasanga Range, Fiji Dept. Agr. 2197 (A, suvA); Mt. Voma, Gillespie 2712

(BISH), Fiji Dept. Agr. 1723 (A, BISH, SUVA); probably near Namosi Village, See-

1968 | SMITH & STONE, ARALIACEAE 469

mann 209 (K type, photos at BISH, US; GH isotype) ; Viti Levu without locality,
MacGillivray & Milne 99 (x, photos at BIsH, Us).

This species, with a comparatively small number of stamens and car-
pels, is readily distinguished from other Fijian Plerandrae on that basis,
but its floral parts are not as consistent in number as believed by Seemann.
It is the sole species of the genus Bakeria Seem., which subsequent authors
have merged with Plerandra.

2. Plerandra grandiflora A. C. Sm. in Bishop Mus. Bull. 141: 117. fig.
61. 1936; J. W. Parham, Pl. Fiji Isl. 85. 1964.

Shrub or slender tree, glabrous throughout, the leaves and inflorescences
aggregated toward ends of branchlets, the branchlets terete, sparsely len-
ticellate; leaves digitately compound, the petioles subterete, 6-20 cm.
long, expanded at base into a sheath 15—20 mm. broad, the ligule coriaceous,
subentire, 5—9 mm. long, the leaflets 3-6, rarely 7, the petiolules subterete,
1-5 cm. long, the blades subcoriaceous, elliptic, 7-18 cm. long, 3.5—8 cm.
broad, attenuate at base, obtuse or rounded at apex, narrowly revolute
at margins, the costa prominent on both surfaces, the secondary nerves
7-16 per side, slightly raised on both surfaces or subimmersed, the veinlet-
reticulation immersed; inflorescences umbellate, with several or numerous
umbels radiating from a very short peduncle, the rays usually 11-17 cm.
long, the subtending bracts caducous, the flowers (11-) 20-25, per umbel,
on stout pedicels 1-2 cm. long; calyx smooth, coriaceous, cylindric-ellip-
soid, at anthesis 6-8 mm. long and about 5 mm. in diameter, truncate at
margin; petals 5, carnose, deltoid-ovate, 5-6 mm. long, mm. broad,
thickened and slightly unguiculate at apex; stamens 2- or 3-seriate, 50—
75, the filaments filiform, usually about 4 mm. long, the anthers oblong,
about 1.5 mm. long, obtuse; styles 5-8, separate, short and obscure, forming
a small ring of papillae on the flattened or rounded summit of the gynoe-
cium, the locules 5-8, with thick walls; fruits oblong-ellipsoid, carnose
when fresh and smooth or inconspicuously ridged when dry, about 2 < 1
cm. at apparent maturity, the stylopodium projecting about 1 mm., the
stigmas obviously separate on the truncate-concave apex, the pyrenes
lunulate, about 15 < 4 mm., with straight inner margins.

Type Locatity: Mt. Kasi, Yanawai River region, Thakaundrove Prov-
ince, Vanua Levu, Fiji; the type is Smith 1777, cited below.

DistriBuTION. Known only from two collections made on the island
of Vanua Levu, Fiji, where it occurs at elevations of 300—-1,030 meters in
dense crest thickets. Field notes indicate the plant as a shrub or slender
tree about 3 meters high; the petals and stamens are pale yellow and
the fruit black. The available specimens were ape in May and No-
vember h bearing flowers and the type fruits as well.

foot ae Naseadon was recorded for no. 1777, although this
name otherwise seems to refer to the genus Polyscias in Fiji.

Fiji. Vanua Levu: THAKAUNDROVE: Mt. Kasi, Yanawai River region, Smith

470 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

1777 (sIsH type, GH, K, NY, UC, US); summit of Mt. Mbatini, Smith 680 (BISH,
GH, K, NY, UC, US).

In stamen number P. grandiflora is intermediate between P. bakeriana
and P. victoriae on the one hand and P. vitiensis on the other. On the basis
of the type collection the reduced number of leaflets (to three) is note-
worthy. The second collection cited above was in 1936 (in Bishop Mus.
Bull. 141: 117) referred to P. victoriae by the first author, but on re-
consideration we prefer to place it in P. grandiflora. The number of leaf-
lets seems of secondary importance in comparison with such characters
as the comparatively elongate fruit and the intermediate number of sta-
mens.

3. Plerandra victoriae Gibbs in Jour. Linn. Soc. Bot. 39: 150. 1909;
A. C. Sm. in Bishop Mus. Bull. 141: 117, quoad abet 1936; J. W.
Parham, PI. Fiji Isl. 86. 1964. L. VII, Fic. 5.

Plerandra sp. A. C. Sm. in Bishop Mus. Bull. 141: 118. 1936.

A glabrous shrub or small tree; leaves digitately compound, Sa petioles
up to 23 cm. long, expanded at base into a coriaceous sheath 2-3 cm
broad, the ligule deltoid-lanceolate, as much as 2 cm. long, acute, dhinner
at margin, the leaflets usually 5—7 (rarely 4 or 8), the petiolules (1—) 2-
4 cm. long, the blades subcoriaceous, elliptic, usually 8-15 cm. long and
3.5-7 cm. broad, obtuse to attenuate at base, rounded or broadly obtuse
at apex, narrowly revolute at margin, the costa prominent on both surfaces,
the secondary nerves usually 10-14 per side, prominulous on both sur-
faces, the veinlet-reticulation immersed; inflorescences umbellate, com-
posed of 5-12 umbels borne near the apex of a variable peduncle 2-12 cm
long, the rays 4~9 cm. long, the subtending bracts soon caducous, the
flowers usually 12-16 per umbel, on pedicels 5-12 mm. long; calyx about
6 mm. long in perfect flowers but shorter in staminate flowers, sinuate-
truncate at margin; petals 5, deltoid-ovate, usually 6-7 « 3-4 mm
unguiculate; stamens usually biseriate and 25-35, the filaments slender,
about 2.5 mm. long, the anthers 2-3 mm. long, obtuse; stigmas small,
on subconnate short styles, like the locules 5—7, rarely 8; fruits ellipsoid,
smooth or inconspicuously costate, 8-12 x 7-9 mm., the stylopodium
1-2.5 mm. long, truncate-conical, composed of the short connate styles
and minutely concave at apex

Type LocaLity: Mt. Tomanivi (Mt. Victoria), Mba Province, Viti
Levu, Fiji; the type is Gibbs 784, cited below.

DIstRIBUTION. Endemic to Fiji and known only from the two largest
islands at elevations of 1,000-1,323 meters (on Viti Levu) and 700 meters
(on Vanua Levu). The species occurs infrequently in the dense mossy
forest and thickets of ridges and summits, as a compact or slender tree
or shrub 2-3 meters high; the petals are blackish without and white
within, the stamens are yellow, and the fruits are deep purple, doubtless
becoming black at maturity.

LocaL NAME. Sole, as for other Fijian Plerandrae.

1968 | SMITH & STONE, ARALIACEAE 471

Fiji. Virt Levu: Ma: Summit ridge of Mt. Tomanivi, Gibbs 784 (BM type,
photo at us), Gillespie 4112 (BisH), Smith 5145 (a, BISH, Us), VANUA LEVU:
THAKAUNDROVE: Mt. Ndikeva, eastern buttress, Smith 1890 (BISH, NY, US).

In 1936 the first author took his no. 680, from Vanua Levu, to repre-
sent P. victoriae, but in our reconsideration we believe this specimen
better placed in P. grandiflora, as noted above. Smith 1890, in 1936 sug-
gested as a possibly undescribed relative of P. bakeriana, is now seen to
have no basic characters separating it from P. victoriae, otherwise not
recorded from Vanua Levu.

The closest relative of P. victoriae is probably P. vitiensis, but its sta-
mens are notably fewer, its ovary-locules are usually (but not invariably)
fewer, its styles are firmly connate in the stylopodium of the fruit rather
than distally free, and its usually fewer leaflet-blades are quite consistently
smaller. Both of these species differ from P. grandiflora most obviously
in fruit shape, but also in stamen number and in (usually) more numerous
leaflets. The three species form a series in various characters but seem
readily distinguishable.

4. Plerandra vitiensis (Seem.) Baill. Hist. Pl. 7: 169. fig. 221. 1879;
A. C. Sm. in Bishop Mus. Bull. 141: 116. 1936; J. W. Parham, PI.
Fiji Isl. 86. 1964. Pi. Vil, yi08: 1, 2.
Nesopanax vitiensis Seem. in Jour. Bot. 2: 249. fig. 1864, Fl. Vit. 117, pl. 20.
1865; Anon. in Gartenfl. 36: 71. 1887, in Kew Bull. 1888: 110. 1888.
Plerandra seemanni B. & H. f. ex Drake, Ill. Fl. Ins. Mar. Pac. 183. 1890.
Plerandra nesopanax Harms in E. & P. Nat. Pflanzenfam. 3(8): 29. 1894.

A tree usually 5-20 m. high, glabrous throughout; leaves digitately
compound, the petiole often stout, up to 60 cm. long, expanded at base
into a coriaceous sheath often 3 cm. broad, the ligule broadly ovate-del-
toid, to 1 cm. long, the leaflets 6-12 (usually 7-10), the petiolules usually
2-5 (sometimes 1-7) cm. long, the blades thin-coriaceous, oblong-obovate
to oblanceolate or elliptic, as much as 27 X 10.5 cm, but usually 13-23
cm. long and 4—9 cm. broad, acute to attenuate at base, rounded to obtuse
at apex, narrowly recurved at margin, the costa raised above, prominent be-
neath, the secondary nerves usually 10-17 per side and prominulous on both
surfaces, the veinlet-reticulation immersed; inflorescences umbellate, com-
posed of 9-35 umbels radiating from a stout peduncle usually 2—3 cm. long,
the rays diverse in length, 6-26 cm. long, the subtending bracts deltoid-lan-
ceolate, to 1.5 cm. long, caducous, the flowers (3—) 10-40 per umbel, on
pedicels usually 2-4 cm. long; calyx conical, 5-6 mm. long (staminate
somewhat smaller), truncate-sinuate at margin; petals 5; deltoid, 5—6 si
long; stamens usually 3—5-seriate and 100-120 (sometimes 75-155), the
filaments slender, 2—2.5 mm. long, the anthers narrowly oblong, obtuse,
1.5-2 mm. long; disk flat or slightly convex; styles usually 7-10 (some-
times 6 or 11, possibly rarely 5), short, separate, with terminal ee
the locules rarely as many as 12; fruits ellipsoid, inconspicuously ae e
when dried, at maturity 8-14 X 9-11 mm., the stylopodium 1—2.5 mm.

472 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

long, the styles connate proximally, with spreading or erecto-patent apices
and often pale terminal stigmas.

Typr Locatity: Southern Ovalau, Fiji; the type is Seemann 207, cited
below. Seemann indicated his specimen as from “Port Kinnaird,’ a name
used at the time of his 1860 visit for the waters between Ovalau and Mo-
turiki sheltered by the Yanutha Islands. In his book Viti (pp. 67-68.
1862), Seemann discusses the possibility of Port Kinnaird being developed
into the capital of Fiji, an unrealized hope of a few settlers

DisTRIBUTION. Endemic to Fiji, and thus far known from the islands
of Viti Levu, Ovalau, and Vanua Levu, occurring with some frequency
in dense or open forest at elevations of 100—-1,150 meters. It is reported
as a tree 5-20 meters high, often slender; the petals at anthesis are rich
purple without and pale green within, the anthers are pale yellow, the
stylopodium and styles are white, and the fruits are dark purple to black
at maturity.

Locat namgs. Sole is generally referred to this species, for which the
first author has also recorded on Vanua Levu the names kaikai and ndani-
ndani; both of these are doubtful, and the last generally denotes the genus
Polyscias.

Fiji. Vir1 Levu: Mpa: Slopes of Mt. Nairosa, Mt. Evans Range, Smith 4060
A, BISH, K, NY, US); vicinity of ee Parks 20749 (isu, UC, US),
Degener 14380 (a, BISH, K, NY, UC, US); M t. Nanggaranambuluta, near Nandari-
vatu, Smith 4756 (A, BISH, K, NY, US), Greenwood 870 (A); east of Nandala
Creek, south of Nandarivatu, Smith 5943 (A, BISH, K, NY, US); Mt. Tomanivi,
Smith 5124 (a, BISH, K, NY, US). NANDRONGA & Navosa: Northern portion of
Rairaimatuku Plateau, between Nandrau and Rewasau, Smith #58 (A, eae K,
NY, US); between Nandrau and Nanga, Smith 5492 (a, BISH, K, NY, US). SER
North of Komave, St. John 18965 (stsH, us). Namost: Mt. Wakaronesi

Korolevu, Smith 7515 (BIsH, Us); east of Lovoni Valley, Smith 7275 (BIsH, us);
vicinity of Levuka, Gillespie 4533 (BISH, GH, UC); southern Ovalau, Seemann
207 (GH, K type, photos at BISH, es VANUA Levu: Msva: Southern slope of
Mt. Seatura, Smith 1621 (BIsH, GH, K, NY, UC, US). MatuHuata: Southern base
of Mathuata Range, north of Natua, Smith 6773 (a, BISH, K, US). THAKAU-
NDROVE: Mt. Mariko, Smith 412 (BIsH, GH, K, NY, UC, US). Fryj1, without other
locality: Tothill 224 (xk, photos at BISH, US

This fairly variable species is distinguished from the first three species in
this treatment most readily by the greater number of stamens, which are
usually 100-120. Its fruit is also characteristic, the styles being obviously
free in the apical part of the stylopodium. In comparison with our first
three species P. vitiensis usually has more numerous and larger leaflets,
although foliage characters in Plerandra are by no means dependable.
The following species, P. grayi, will superficially often be mistaken for P.
vitiensis.

5. Plerandra grayi Seem. Viti 437, nomen. 1862, in Jour. Bot. 2: 242.
fig. (p. 241). 1864, FI. Vit. 117, pl. 22. 1865; Drake, Ill. Fl. Ins. Mar.

1968 | SMITH & STONE, ARALIACEAE 473

Pac. 183. 1890; A. C. Sm. in Bishop Mus. Bull. 141: 116. 1936; J.
W. Parham, PI. Fiji Isl. 85. fig. 35. 1964. PL. VII, Fics. 6-9.
Plerandra sp. A. Gray in Proc. Am. Acad. Arts Sci. 5: 318. 1862, in Bon-
plandia 10: 36. 1862.
Plerandra graeffei Anon. in Gartenfl. 36: 71. 1887, in Kew Bull. 1888: 117.
1888.

A small tree, glabrous throughout; leaves digitately compound, the
petiole subterete, stout, up to 60 cm. long, expanded at base into a coria-
ceous sheath 2-3 cm. broad, the ligule broadly ovate, 1-2 cm. long, the
leaflets 4-10 (usually 6-9), the petiolules 2.5-7 cm. long, the blades sub-
coriaceous, usually broadly obovate, less frequently oblanceolate or lan-
ceolate, (11-) 14-30 cm. long, (5—) 6-14.5 cm. broad, acute and decur-
rent at base, usually broadly rounded at apex, recurved or nearly plane
at margin, the costa flattened above, prominent beneath, the secondary
nerves usually 9-16 per side, prominulous on both sides or plane above,
the veinlet-reticulation immersed; inflorescences umbellate, composed of
7-30 umbels radiating from a variable peduncle up to 10 cm. long, the
rays (5—-) 12-30 cm. long, the subtending bracts oblong-deltoid, sub-
coriaceous, 1—2 cm. long, caducous, the flowers 15-40 per umbel, on pedi-
cels 1.53 cm. long; calyx of perfect flowers broadly cupuliform, 6-8 mm.
long and 5—7 mm. broad, truncate-sinuate at margin; petals 5, carnose,
deltoid, 8-12 mm. long, 5—7 mm. broad, unguiculate; stamens usually
3-5-seriate, (120-) 175-250, the filaments slender, diverse but usually
2-3 mm. long, the anthers oblong, obtuse, 2-2.5 mm. long; disk slightly
concave at center, flat marginally; stigmas minute, essentially sessile,
like the locules 11-15; fruits ellipsoid or nearly subglobose, usually ob-
viously costate, at apparent maturity 11-20 x 10-15 mm., subtruncate at

Type Locauity: Viti Levu, Fiji; the type is Seemann 208, cited below.
Although Seemann indicated no detailed locality, the holotype bears the
note “July 1860.” In the early part of that month Seemann was 1n south-
ern Viti Levu between the Navua River and the present town of Ngaloa
(Viti, pp. 95-117. 1862); during the rest of July he seems to have ath
in Ovalau, Mbau, and Tailevu not primarily engaged in botanical ate
Therefore we may assume his no. 208 to have been collected along the
Serua coast.

DistrIBpuTION. Endemi
Levu, where it occurs apparently infrequently
meters in forested areas. It is indicated as a usuall hf
meters high; the petals are greenish with copious rich purple mar ~G
without, or at least obviously purple-tinged, and greenish white within;
the stamens are cream white to greenish white; and the fruits are black
at maturity.

Loca NAME. The generic name sole is co

c to Fiji and thus far known only from Viti
at elevations up to 400

mmonly used for P. grayi.

474 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Fiji. Vit1 Levu: Serva: Vatutavathe, vicinity of Ngaloa, Degener 15201 (A,
BISH, K, NY, UC, US). NAmost: Hills bordering Wainavindrau Creek, vicinity of
Wainimakutu, Smith 8523 (BISH, US), 8568 (US); east of Wainikoroiluva River,

near Namuamua, Smith 8902 (BIsH, US), 9046 (BISH, US). NaTITAsIRI: Tamavua,
Gillespie 2464 (BIsH, UC); Prince’s Road, Meebold 16558 (k, photos at BISH, US),
Fiji Dept. Agr. 196 (SUVA). TAILEVU: am of Wainimbuka River, vicinity of
paar bis Smith 7140 (BISH, US). : Mt. Korombamba, Parks 20118

H, Uc); “vicinity of Suva,” peer 16555 (pisH), VitT1 Levu without other
tocaltee (probably Serua Province): Seemann 208 (K type, photos at BISH, US;
isotypes at BM, GH, erroneously labelled 209, photo at us).

Although P. grayi and P. vitiensis are superficially similar, the broader
and more obviously rounded leaflet-blades of the former are characteristic,
but this is not completely dependable. The petals and fruits of P. grayt
are larger than those of P. vitiensis, and the stamens are also more numer-
ous. The ovary-locules of P. erayi are 11-15, those of P. vitiensis being
5-11. The very short stylopodium of the fruit of P. grayi bears very short
styles connate into a centrally concave ring, whereas the stylopodium
of the fruit of P. vitiensis is comparatively obvious, the styles being
definitely free distally.

ol

Plerandra pickeringii A. Gray, Bot. U.S. Expl. Exped. 1: 729. 1854,
Atlas pl. 95. 1857, in Ann. Sci. Nat. IV. Bot. 4: 178. 1855, in Proc.
Am. Acad. Arts Sci. 3: 129. 1857; Seem. in Jour. Bot. 2: 242. 1864,
FI. Vit. 117. 1865; Drake, Ill. Fl. Ins. Mar. Pac. 183. 1890; A. C.
Sm. in Bishop Mus. Bull. 141: 116. 1936, in Jour. Arnold Arb. 33:
103. 1952; J. W. Parham, Pl. Fiji Isl. 86. 1964

Px. VII, Fics: 1, 2.
A tree to 15 m. high or shrub, glabrous throughout; leaves digitately
compound, the petiole stout (to 1.5 cm. in diameter), subterete, up to 100

m. long, sometimes verrucose-lenticellate, expanded at base into a cori-

aceous sheath 2-4 cm. broad, the ligule oblong-deltoid, 1-2 cm. long, the

leaflets 6-12 (usually 7-11, rarely as many as 16), the petiolules stout,

(1.5—) 3-13 cm. long, the blades subcoriaceous, obovate or oblong-obovate

or oblanceolate, usually 16-40 cm. long and 6-22 cm. broad, occasionally
up to 47 24 cm., acute to attenuate at base, rounded to obtuse or broad-
ly acute at apex, usually plane at margin, the costa often prominent on
both surfaces, the secondary nerves usually 10-20 per side, raised on both
surfaces or plane above, the veinlet-reticulation prominulous to immersed;
inflorescences umbellate, composed of 7-15 umbels radiating from a stout
short peduncle usually 2-3 cm. long, the rays 10-50 cm. long, subtended
by lanceolate-deltoid bracts 1.5-2 cm. long, these soon caducous, the
flowers 20-60 per umbel, on pedicels 3—5.5 cm. long; flowers borne termin-
ally or closely aggregated on the clavate ultimate 3 cm. of the ray, sub-
tended by coriaceous oblong bracts 1—1.5 cm. long, these soon caducous;
perfect flowers with an obconical-cupuliform calyx 8-12 mm. long and
7-10 mm. broad at the sinuate-undulate margin, the calyx of staminate
flowers usually 5-7 mm. long and broad; petals usually 5, rarely 4-6,

1968 | SMITH & STONE, ARALIACEAE 475

appearing calyptrate in bud but eventually free, carnose, oblong-deltoid,
9-16 mm. long, 6-10 mm. broad, subacute, inconspicuously unguiculate;
stamens usually 5~—7-seriate, numerous, usually 400-500 in staminate
flowers and 200-350 in pistillate flowers, the filaments slender, very vari-
able (2-10 mm.) in length, the anthers narrowly oblong, obtuse, similarly
variable (2-5 mm.) in length; disk flattened or slightly convex, the sty-
lopodium at anthesis bluntly conical, about 1 mm. long, composed of
firmly connate styles and minute terminal stigmas; styles usually 11-16
(rarely 9-17), the locules similar in number; fruits ellipsoid or subglobose,
at maturity strongly costate, 20-25 (—40) mm. long (excl. stylopodium),
17-20 (—25) mm. broad, the stylopodium conspicuous, broadly conical at
base, cyclindric above, 5-10 mm. long, ridged, with extrorse stigmas mar-
ginal on the truncate apex, the pyrenes lunulate.

TYPE Locatity: Ovalau, Fiji; the type is a U.S. Exploring Expedition
specimen, cited below.

DistrIBUTION. Endemic to Fiji and doubtless to be expected on most
of the high islands from near sea-level to 1,050 meters. It occurs in many
types of forest as a slender tree 3-15 meters high (rarely noted as a shrub),
with the leaves and the large terminal inflorescences clustered at apices
of branchlets. The calyx and petals are white to green or yellowish, usual-
ly purple-tinged; the filaments and anthers are white to pale yellow; the
stylopodium in flower is yellowish white with pale green stigmas; and the
fruits are black at maturity.

LocaL NAMES AND USES. This species is most commonly known as sole
or sole ndina (i.e., the true sole). In interior Viti Levu the name sole ngua
has been reported, and also the use of trunks as posts for fish weirs. The
local name vola has been reported from Vanua Levu, and also ndanindani,
both of these being of questionable application to Plerandra.

Fiji. Vitt Levu: Mpa: Mt. Nairosa, Mt. Evans Range, Smith 4021 (a, BISH,
K, NY, US); vicinity of Nandarivatu, Gillespie 3707 (BIsH); between Nggaliwana
and Nandala Creeks, south of Nauwanga, Smith 5803 (A, BISH, US). SERUA:
Between Navua River and Wainiyavu Creek, near Namuamua, Smith 8978 (BISH,
us). Ra: Mataimeravula, near Vaileka, Degener 15440 (A, BISH, K, NY, UC, US).
Narrasir1: Matawailevu, Wainamo Creek, St. John 18223 (BISH, US); Tamavua,
Gillespie 2156 (BisH, GH, NY); Nasinu, Gillespie 3432 (BISH, GH); Kalambo,
Tothill 222 (a, psu, K), Fiji Dept. Agr. 11242 (suva). Rewa: Namboro, Fiji
Dept. Agr. 5916 p. p. (BISH, suvA). Kanpavu: Mt. Mbuke Levu, Smith 205
(BISH, GH, K, NY, UC, US); without locality, Fiji Dept. Agr. 11943 (svva).
Ovatau: U.S. Expl. Exped. (us 62359 type; fragmentary isotype at GH). Koro:
Eastern slope of main ridge, Smith 936 (BISH, GH, K, NY, UC, US). NGAU: ae
of Herald Bay, inland from Sawaieke, Smith 7751 (pis, us). VANUA Levu:
Msva: Southern portion of Seatovo Range, Smith 1524 (BISH, GH, K, NY, UC,
us). Maruuata: Seanggangga Plateau, vicinity of Natua, Smith 6896 (A, BISH,
NY HAKAUNDROVE: Maravu, near Salt Lake, Degener & Ordonez
14274 (A, BISH, K, NY, UC, US). TAVEUNI: Western slope near Waiyevo and
Wairiki, Gillespie 4678 (BISH, ny, uc). Moata: Near Naroi, Smith 1309 (Bish,
GH, K, NY, UC, US).

The type species of Plerandra, P. pickeringit, is remarkably robust in

476 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

its foliage and inflorescence dimensions, but its distinctness among our
species rests on its characteristic fruits, which have the stylopodium con-
spicuous (5-10 mm. long) and composed of completely connate styles.
The fruit is also comparatively large; when smaller than about 20 X 17
mm. one may suspect it to be immature. Nevertheless certain specimens,
such as Smith 6896 (the subject of our PLare VIII, ric. 1), have com-
paratively small and globose fruits that are not entirely characteristic. The
functionally staminate flowers have the extraordinary number of 400-500
stamens, while the perfect flowers seem to have 200-350 stamens, remark-
ably high for the genus.

7. Plerandra insolita A. C. Sm. in Jour. Arnold Arb. 33: 103. 1952;
J. W. Parham, Pl. Fiji Isl. 86. 1964. PL. VIII, Fics. 3-7.

A slender tree to 9 m. high, glabrous throughout; leaves digitately com-
pound, the petiole stout, subterete, 40-70 cm. long, often verrucose-lenti-
cellate, expanded at base into a coriaceous sheath 2-4 cm. broad, the
ligule oblong, 1.53 cm. long, the leaflets 5-12 (usually 8-11), the petiolu-
les stout, 2-7 (—10) cm. long, the blades subcoriaceous, oblong- or obovate-
elliptic or oblanceolate, 20-41 cm. long, 5—22 cm. broad, acute to attenuate
at base, obtuse to rounded at apex, narrowly recurved at margin, the costa
prominent on both surfaces, the secondary nerves 12-25 on each side,
raised on both surfaces, the veinlet-reticulation prominulous beneath or
immersed; inflorescences compound-pseudo-umbellate, arising from branch-
lets below leaves, composed of 5—7 rays borne at or near the apex of a
short, stout peduncle 1-5 cm. long, the rays 4-8 cm. long, conspicuously
swollen in the distal 1-5 cm. and copiously cicatricose, the flowers 15-35
or perhaps more per ray, congested in a strobilus-like pseudo-umbel, the
pedicels very short at anthesis, elongating to 6-18 mm. in fruit, subtended
by subcoriaceous oblong bracts about 1.5 cm. long, these persistent through
anthesis; calyx of perfect flowers obconical-cupuliform, 7-9 mm. long an
5-7 mm. broad, irregularly lobed on the margin of the short spreading
limb; calyx of staminate (and usually distal) flowers 3—5 mm. long; petals
5 or 6, rarely 4, carnose, subconnate into a conical calyptra, deltoid, 7-8

4—6 mm., inconspicuously unguiculate; stamens 2- or 3-seriate, 50-75,
the filaments slender, 1-3.5 mm. long, the anthers oblong, obtuse, 2-3
mm. long; disk flattened, the stylopodium short-conical or umbonate,
sulcate, composed of firmly connate styles with minute stigmas; styles and
locules 9-12; fruits ellipsoid or subglobose, carnose and smooth when
fresh, when dried coriaceous and strongly costate, 25-36 mm. long, 18-30
mm. broad, the calycular rim apically persistent, the stylopodium stout,
1-2 mm. long and 3-5 mm. in diameter, with grooved stigmas conspicu-
ota ea: marginally on the truncate apex, the pyrenes falcate-
obovoid.

TYPE LOCALITY: Southern slopes of Mt. Ndelainathovu, on the escarp-
ment west of Nandarivatu, Mba, Viti Levu, Fiji; the type is Smith 4922,
cited below.

1968 | SMITH & STONE, ARALIACEAE 477

DistRIBUTION. Endemic to Fiji and thus far known only from Viti
Levu, where it occurs infrequently at elevations from near sea-level to
1,160 meters, It has been found in dense or open forest as a slender tree,
unbranched or sparingly branched, with a sticky colorless latex. The petals
are purple without and pale green or white within; the filaments are white
or greenish and the anthers yellow; the disk is pale green at anthesis; and
the fruits are black at maturity.

Locat NAMES. While sole is the general name for this species, it has
also been recorded as sole lailai and sole ngua in interior Viti Levu.

us); hills near Navua River, Greenwood 1048 (A, BISH, NY). NAITASIRI:
Wainisavulevu-Numbulolo divide, St. John 18325 (BIsH, US); Viria, Meebold
16557 (sisH); Central Road, MacDaniels 1149 (sisH); Suva Ditch trail, Bryan
376 (BIsH); near Nanduna Village, Fiji Dept. Agr. 12593 (BISH, SUVA); Tamavua,
Gillespie 2465 (BISH, GH, US). Rewa: Namboro, Fiji Dept. Agr. 5916 p. p.
(suva). Vitt Levu without further data: Seemann 206 (BM, GH, K, photos at
BISH, US) (July 1860; probably from the Serua coast; cf. comment on type
locality of P. grayi, above).

A redescription of this striking species seems desirable in view of the
number of available collections not at hand in 1952; these collections
permit a few emendations but only serve to strengthen the status of the
species. Plerandra insolita suggests P. pickeringii in the robustness of its
foliage parts, but it is the most distinct of our species because of its con-
tracted inflorescence, of which the flowers are not borne in umbels but
rather in congested spirals on the clavate or cylindric distal portion of
each ray. A further differentiating character is the comparatively large
and persistent calycular rim. The fruits are as large as those of P. picker-
ingii but very different in characters of the stylopodium; and of course the
stamens are not comparable in number to those off: pickeringii. While
a rigid concept of Plerandra might incline students of the family to segre-
gate P. insolita generically, we feel that the circumscription of the genus
is not greatly disturbed by the inclusion of this remarkably distinct species.
In Hutchinson’s 1967 system P. insolita would fall into the tribe Cus-
sonieae, far removed from Plerandra, and the concept would probably re-
quire a new genus.

6. Schefflera J. R. & G. Forst. Char. Gen. Pl. 45, nomen conservandum.

1776; Seem. in Jour. Bot. 3: 175. 1865, Fl. Vit. 116. 1865; Harms in

E. & P. Nat. Pflanzenfam. 3(8): 35. 1894, in Bot. Jahrb. 56: 385.
1920; Hutchinson, Gen. Fl. Pl. 2: 69. 1967.

Schefflera, typified by the New Zealand S. digitata J. R. & G. Forst.,

478 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

is a widespread tropical and subtropical genus, but its limits are still open
to interpretation. In his 1894 review of the family, Harms used Schefflera
in a broad sense, including Heptapleurum Gaertn. and Agalma Mig. On
the other hand, Hutchinson in 1967 reinstated Agalma (at least in the
sense of Miquel’s type species and some 22 others) and also combined
Cephaloscheflera Merr. (Schefflera sect. Cephaloscheflera Harms) with
a greatly expanded Brassaia Endl.

As the family is under further consideration by specialists, we cannot
suggest reasonable limits for Schefflera on the basis of our restricted re-

view. It is evident that our first five species, with paniculate-racemose
inflorescences and free styles, belong in Schefflera in even the limited
sense. The remaining four species, with compound-umbellate inflores-
cences, would fall into Harms’ concept of his subsection Agalma (but
this concept may not be strictly identical with Miquel’s genus in a narrow
sense). Of our species, S. actinostigma has free styles, whereas both this
and S. tannae show transitional stages between paniculate-racemose and
compound-umbellate inflorescences. It would seem that even among our
few species inflorescence-characters of a satisfactory generic nature are
evasive.

For present purposes we adopt Scheffiera in the more inclusive sense,
retaining in it species with strictly compound-umbellate inflorescences as
well as with connate styles, but we refrain from suggesting its synonymy.

KEY TO THE SPECIES

ee paniculate-racemose, with numerous lateral branches bearing
umbels racemosely arranged; fruits comparatively small, usually
less shee 6 X 6 mm. at maturity, with 5-10 locules; leaflet-blades with 7-18
secondary nerves per side

Indument (on young parts, some foliage parts, and inflorescence-axes) com-
posed of conspicuous, coarse, scale-like, many-celled hairs 2-10 mm. long;
primary inflorescence-bracts conspicuous, 25-40 mm. long, the umbel-
subtending bracts 5-10 mm. long. .................. 1. S. euthytricha.
Indument lacking or composed of minute, par goriigs hairs (in nos. 2 and
3 the hairs sometimes scale-like and 0.5-3.5 mm. long, but sparse and
ae Pibeptse primary Spal iter comparatively small,

3-15 mm. long, the umbel-subtending bracts 1-6 mm. long.
Leaflet-blades of normal adult leaves copiously PELE EN serrate at margin
ities spinulose, 1-7 per centimeter); styles and ovary-locules often

more

Styles and oar doce (6—) 7-10, the styles in fruit oe 1-1.2
mm. long; flowers comparatively small, with petals 1.3-1.5 mm. long;
leaflets with petiolules 5-15 mm. long and blade-margins ines den-
ticulate, the teeth irregular, 2~7 per centimeter. ..............-.
sy Ha An x lien Ja Bioce oR gay ae) Se eae . S. samoensis.
Styles and ovary-locules 5 an * the styles in fruit ne mm. long;
owers with petals abou mm. long; leaflets with conspicuous
petiolules 20-40 mm. ie td Slice ene coarsely denticulate,
the teeth 1-3 per centimeter: .. 22.6 2) os. 3. S. neo-ebudica.

1968 | SMITH & STONE, ARALIACEAE 479

hare blades of normal adult leaves entire or crenulate at margin (teeth

inulose or evanescently so, rarely as many as 2 per centimeter);

ae apparently consistently 5-merous, with 5 styles and ovary-
locules.

Inflorescences comparatively compact, the lateral branches 3- a cm. long;
leaflets 9-13, the blades rounded to broadly obtuse at a
ee a eer ee ee aan NE " tannae.

canine comparatively ample, the lateral branches usually 15-40

m. long; leaflets (5—) 7-9, the blades acute to acuminate at apex. ..

A Pea is BR eee Fe ee ee ee ee ee vitiensis.

Inflorescences compound-umbellate, with comparatively few branches, 1-3 times

divided, the flowering umbels aggregated at apices of rays; fruits com-

paratively large, usually more than 6 X 6 mm. (sometimes to 13 X 11 mm.)

at maturity, with 5-12 locules; leaflet-blades with 20-30 secondary nerves
per side, these often interspersed with others nearly as obvious

Fruits truncate at apex, surmounted by the subpersistent calycular rim, the

stylopodium flattened, the styles 5-7, divergent, 1-1.5 mm. long; in-

florescences racemose-umbellate (tertiary branches whorled at a median

node and also aggregated at ray-apex); leaves with the petiole 36-50 cm.

the petiolules 3-5 cm. long, the blades 11-22 X 6-9 cm., rounded

roadiy retiwe at Spe®e ccs as has a 6,5. actinostigma.

Fruits with an obvious stylopodium projecting beyond the sane ay

calycular rim or scar, the styles connate into an obvious column 1.5-2 m

long; inflorescences strictly compound- gerund oT with the settele

less than 22 cm. long, the petiolules 1-3.5 cm

Leaves with a conspicuous petiolar ligule free in ‘the distal 3-4 cm., the
leaflet-blades conspicuously acuminate at apex; fruits with 8-12 costae
“ locules, the stylar column stout, conical, about 1.5 mm. long and
1.5mm, in apical Glameter 2. ee . S. costata.

pom (not known for no. 9) with a petiolar ligule free for less than 1 cm.,
the leaflet-blades rounded or obtusely short-acuminate at apex; fruits

with 5-9 costae and locules, the stylar column slender, about 2 mm

long.
Infructescences comparatively compact, the peduncles of rays 2-9 cm.
long; fruits 11-13 X 9-11 mm., 5-7-costate, the stylopodium

conical, projecting (at base of stylar column) about 2 mm. above
the calycular rit. 2.45060 eee "s seemanniana.

Infructescences ample, the peduncles of rays 4-12 cm. long; fruits 7-9
8-10 mm., 6-9-costate, the stylopodium rounded, sanigtete
base of stylar column) 2-3 mm. above the calycular rim. ..

1. pti euthytricha A. C. Sm, in Contr. U.S. Natl. Herb. bl et
967. :

As no additional specimens have been located since the recent ae
te it is not necessary to supplement that, other than by the
illustratio

Type tocatity: Namosi Province, Viti Levu, Fiji; the type is Smith
8908, cited below

Distr1But TION. " Endemic to Fiji, and thus far known from only two
localities in the vicinity of the junction of the Wainikoroiluva and Navua

480 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Rivers, south-central Viti Levu, where it occurs in dense forest at an eleva-
tion of 50-200 meters. The available specimens are from slender trees
5—6 meters high, with a conspicuous, dull brown inflorescence-tomentum;
the calyx, petals, and filaments are greenish white, and the anthers white.
Both known specimens, in flowering condition, were obtained in October
1953.

Loca NAME. The type collection was indicated as sole tangane.

Fiji. Virr Levu: Serva: Hills between Navua River and Wainiyavu Cree

near Namuamua, Smith 8983 (BISH, US). NaAmosr: Hills east of yeas
River, near Namuamua, Smith 8908 (BIsH, Us 2192113 type).

As indicated in connection with the original description, S. euthytricha
agrees with its only close ally in Fiji, S. vitiensis, in basic inflorescence-
characters, differing obviously in its striking indument and large inflores-
cence-bracts. In respect to its indument, S. euthytricha is approached by
S. samoensis and S. neo-ebudica, endemic to Samoa and the New Hebrides
respectively, but the three species are sharply distinct in the several
characters utilized in our key

2. Schefflera samoensis (A. Gray) Harms in E. & P. Nat. Pflanzenfam.
3(8): 39. 1894; Reinecke in Bot. Jahrb. 25: 663. 1898; Rechinger in
Denkschr. Akad. Wiss. Wien 85: 325. 1910; Christophersen in Bishop
Mus. Bull. 128: 164. 1935. Pt. IV, Fics. 3-5.

Paratropia samoensis A. Gray, Bot. U.S. Expl. Sean 1: 722. 1854.
Cheirodendron samoense Seem. in Jour. Bot. 5: 237. i
Heptapleurum samoense B. & H. f. ex Drake, Ill. Fl. inh Mar. Pac. 183. 1890.

A smal] tree or shrub, the young branches, petioles, petiolules, and
sometimes leaflet-costae beneath densely or sparsely hirsute with pale
linear-lanceolate many-celled hair-like scales 1-3.5 mm. long, soon glab-
rate; leaves digitately compound, the petiole slender, 10—25 cm. long, ex-
panded to an inconspicuously clasping sheath, the ligule free in the distal
3-5 mm., obscurely bifid at apex, the leaflets (5—) 6-9, the petiolules
slender, 5-15 mm. long, the blades papyraceous, oblanceolate to narrowly
obovate, (4—-) 8-20 cm. long, (1.5—-) 2.5-7.5 cm. broad, gradually at-
tenuate at base, usually conspicuously acuminate at apex, strongly dentic-
ulate-serrulate at margins (teeth spinulose, irregular, 2—7 per centimeter),
the costa elevated on both surfaces, the secondary nerves 7—13 per side,
arcuate-ascending, with the veinlet-reticulation usually plane above and
prominulous beneath; inflorescences paniculate-racemose, the axes irreg-
ularly p‘lose but sometimes sparsely hirsute like vegetative parts, soon
glabrate, the main rachis stout, up to 30 cm. long, the primary branches
subopposite or irregularly disposed, 13-25, up to 25 cm. long, the umbels
borne on slender peduncles (1—) 5-18 mm. long at anthesis and up to 30
mm. long in fruit, the flowers usually 3-10 per umbel, the pedicels 1-5 mm.
long (to 7 mm. in fruit), the primary inflorescence-bracts oblong-deltoid,
3—6 mm. long, the umbel-subtending bracts 1-3 mm. long; flowers 1.5—2
mm. long at anthesis, the calyx broadly cupuliform, 0.5—1 mm. long, with

1968 | SMITH & STONE, ARALIACEAE 481

a slightly flaring rim of 4-6 minute deltoid lobes; petals 4-6, oblong-
deltoid, 1.3-1.5 mm. long, 0.7-1.2 mm. broad, unguiculate-thickened at
apex; stamens 4-6, the filaments slender, 1-1.5 mm. long, the anthers
oblong, rounded, versatile, 0.5-0.6 mm, long; ovary-apex rounded, the
stigmas more numerous than petals, (6-) 7-10 (like ovary-locules), the
styles minute, about 0.2 mm. long at anthesis, appressed in a small um-
bonate projection; fruits broadly subglobose, 2-3 mm. long and 3-3.5
mm. broad, conspicuously (6—-) 7-10-costate, broadly conical at apex,
the styles 1-1.2 mm. long, radiating or reflexed, clavate at apex and with
oblique stigmas.

Type LocaLity: Upolu, Samoa; the type is the U.S. Exploring Expedi-
tion specimen cited below.

DIstTRIBUTION. Endemic to Samoa, and definitely known only from the
islands of Savaii and Upolu, where it occurs with some frequency in
forests and ridge thickets at elevations of 700-1,500 meters (according to
Christophersen’s notes). It is indicated as a tree or shrub 3—5 meters high,
with green or yellowish petals, and with dark brown or perhaps black
fruits.

Loca NAME. The only name we find recorded is maialeale (Christopher-
sen 234).

Samoa. Savair: Above Matavanu Crater, Christophersen 634 (BISH); inner
slopes of Matavanu Crater, Christophersen & Hume 2225 (A, BISH, US); near
Olo, above Safotu, Christophersen & Hume 2270 (A, BISH, Us); “Mataana,
Vaupel 351 (BisH, Ny, Us). Upotu: Above Malololelei, Christophersen 234
(BISH, NY); without definite locality: U.S. Expl. Exped. (us 47926 type).
SAMOA, without definite locality: Whitmee 56 (xk, photos at BISH, Us), S. n. |
GH). We have not seen the following numbers cited by Rechinger: Rechinger
1610 and 1753 from Savaii, 1847 and 1919 from Upolu.

Schefflera samoensis is sharply characterized by its finely denticulate
leaflet-margins and by its small flowers and fruits. Whereas the other
species of this relationship have the ovary-locules and styles similar in
number to the other flower-parts, S. samoensis has usually 7-10 locules
and styles, although the petals and stamens are 4-6 in number. The con-
spicuous and radiating styles in fruit further distinguish this very distinct
Samoan endemic.

3. Schefflera neo-ebudica Guillaumin in Bull. Mus. Hist. Nat. II. 9:
289. 1937, in Jour. Linn. Soc. Bot. 51: 554. 1938. PL. IV, Fic. 6.

Schefflera sp. Guillaumin in Jour. Arnold Arb. 12: 263. 1931.

A small tree, glabrous throughout or with a few coarse many-celled eas
like scales 0.5-1 mm. long at base of petioles and on leaf-sheaths (and
rarely with a few obscure, minute, stellate hairs on si apr ecapinnane a
leaves digitately compound, the petiole stout, 15-27 cm. long, expan :
at base into a coriaceous clasping sheath, the ligule rounded, free in the
distal 1 cm., the leaflets 7~10, the petiolules 2-4 cm. long, the blades papy-

482 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

raceous, narrowly elliptic to oblanceolate, 12-25 cm. long, 4~7 cm. broad,
attenuate at base, caudate-acuminate (but easily broken) at apex with an
obtuse tip 1.5—2 cm. long, coarsely dentate at margin (teeth spinulose,
1-3 per centimeter), the costa slightly raised above and prominent beneath,
the secondary nerves usually 8-12 per side, erecto-patent, with the veinlet-
reticulation plane above and prominulous beneath; inflorescences panicu-
late-racemose, up to 40 cm. in length, pedunculate, the main rachis up to
28 cm. long, the primary branches 10 or more in number, usually 12-17
cm. long, the umbels borne on slender peduncles 4-8 mm. long, the flowers
usually 5—7 per umbel, the pedicels 2-5 mm. long, the primary inflores-
cence-bracts oblong-deltoid, 4-6 mm. long, the umbel-subtending bracts
2-3 mm. long; flowers with a small cupuliform calyx with 5 or 6 minute
deltoid lobes; petals 5 (or 6?), deltoid, about 2 mm. long; stamens 5 (or
6?), the filaments about as long as the anthers; styles (and ovary-locules)
5 or 6, the styles minute in flower; fruits broadly subglobose, 3—3.5 mm.
long, 3.5—4 mm. broad, and strongly 5- or 6-costate when dried (up to 6
>< 5 mm. when fresh, ex Kajewski), the stylopodium extending about 1
mm. above the calycular scar and sharply angled, the styles 0.5-0.7 mm.
long, free and spreading or recurved from a shortly connate basal portion,
the stigmas minute.

TYPE LOCALITY: New Hebrides; Guillaumin does not indicate a type
but cites three specimens: de /a Riie from Ambrym, de la Riie from
Pentecost, and Kajewski 114 from Lenakel, Tanna. We have seen dup-
licates of the latter two, and the last is an excellent fruiting specimen with
duplicates in several herbaria. Therefore, we herewith designate Kajewski
114 (holotype at Paris) as the lectotype. This was one of the two Kajew-
ski specimens cited without an epithet by Guillaumin in 1931.

DIstTRIBUTION. Endemic to the New Hebrides, and thus far known from
the islands of Espiritu Santo, Pentecost, Ambrym, Tanna, and Aneityum,
at elevations of 200-1,000 meters in forested areas. It is recorded as a
tree 5—7 meters high; the only color notes (on Kajewski 845) indicate
the fruit as cream-colored.

Loca, NAMES. The name narku is recorded from Espiritu Santo, and
kassaskelimbueret from Pentecost.

New Hebrides. Esprriru Santo: Between Ladhogh and Turworsoksok, J. &
Z. Baker 138 (gM). PENTEcost: Between Kumre and Lasup, de Ja Riie (A).
AmBryM: Mt. Toiio, de Ja Riie (not seen in this study). TANNA: Lenakel,
March 6, 1928, Kajewski 114 (isotypes at a, K, NY, photos at BISH, US). ANEI-
TyuM: Anelgauhat Bay, Kajewski 845 (a, BISH, K, NY, photos at BISH, US).

In 1931 Guillaumin cited Kajewski 114 and 845 as presumably repre-
senting two separate species, but they seem without doubt to be con-
specific. Among the species of our area, S. neo-ebudica suggests only S.
samoensis, clearly differing in its leaves with conspicuous petiolules and
more coarsely denticulate blade-margins, the larger flowers, and the normal
number (5 or 6) of ovary-locules and styles, the latter being comparatively
short, although also free, in fruit.

1968 | SMITH & STONE, ARALIACEAE 483

4. Schefflera tannae A. C. Smith & B. C. Stone, sp. nov. FL. Y.
Schefflera sp. Guillaumin in Jour. Arnold Arb. 12: 263. 1931.

Arbor ad 15 m. alta ubique mox glabra, partibus novellis forsan parce
puberulis; foliis digitatis, petiolis subteretibus 15-30 cm. longis, basi in
vaginam coriaceam parce lenticellatam incrassatis, ligula late rotundata
4-6 mm. longa; foliolis 9-13, petiolulis gracilibus 1-3 cm. longis, superne
anguste alatis, laminis papyraceis oblanceolatis vel obovatis, 8-22 cm.
longis, 3.5—8 cm. latis (lateralibus saepe minoribus), basi attenuatis et in
petiolulum longe decurrentibus, apice rotundatis vel late obtusis, margine
anguste revolutis et subintegris vel obscure crenatis (dentibus haud 1 per
centimetrum), costa supra elevata subtus prominente, nervis secundariis
utrinsecus 8-12 adscendentibus subtus prominulis, rete venularum plano
vel utrinque prominulo; inflorescentiis (immaturis solis visis) paniculato-
racemosis ad 22 cm. longis, rhachide crassa et pedunculo ad 4 cm. longis,
ramulis secundariis erecto-patentibus circiter 5 bracteis papyraceis ad 15
mm. longis mox caducis subtentis, ramulis tertiariis brevibus (3-5 cm.
longis) oppositis vel aggregatis, bracteis mox caducis; umbellularum
pedunculis gracilibus 3-6 mm. longis, floribus 6-12 per umbellulam, pedi-
cellis in alabastro circiter 1 mm. longis bracteolis deltoideo-rotundatis haud
0.5 mm. longis subtentis; calyce cupuliformi haud 0.5 mm. longo, lobis 5
deltoideis minutis; petalis 5 oblongo-deltoideis in alabastro circiter 1 mm.
longis apice cucullatis; disco subplano, staminibus 5, stylis in alabastro
minutis erectis contiguis; fructibus non visis.

Type Locariry: Lenakel, Tanna, New Hebrides; the type is Kajewski

DistripuTION. Known only from the type collection and presumably
endemic to the New Hebrides. The species is indicated by the collector
to be common in rain-forest at an elevation of 200 meters; the available
material is from a tree about 15 meters high, with a trunk 50 centimeters
in diameter.

New Hebrides. TANNA: Lenakel, March 7, 1928, Kajewski 131 (a type, BISH,
K, NY).

Although the single collection here described does not bear fully mature
inflorescences, we venture to describe it because the foliage- and aye
cence-characters are strikingly distinct from those of o neo-ebudica, whic
is known from the same locality. From the earlier species our novelty
differs sharply in its essentially entire leaflet-blades with rounded —
apices. It is also characterized by more compact inflorescences with fewer
and shorter lateral branches, which appear to be aggregated at two d
three nodes along the rachis, suggesting a transition toward a a -
umbellate condition that is further emphasized in S. actinostigma, dis-
cussed below.

5. Schefflera vitiensis (A. Gray) Seem. in Jour. Bot. 3: pace 2
Vit. 116. 1865; Drake, Ill. Fl. Ins. Mar. Pac. 182. 1890; Harms

484 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

E. & P. Nat. Pflanzenfam. 3(8): 39. 1894; Gibbs in Jour. Linn. Soc.
Bot. 39: 148. 1909; J. W. Parham, PI. Fiji Isl. 86. 1964.
Pt. IV, Fics. 1, 2.

Aralia vitiensis A. Gray, Bot. U.S. Expl. Exped. 1: 715. 1854, Atlas pl. 89.
1857.

A tree, glabrous throughout at maturity, or with a close, scaly, minute
indument subpersistent on lower surfaces of leaflets, inflorescence-axes,
etc.; leaves digitately compound, the petiole 12-42 cm. long, expanded
to a coriaceous sheathing base, the ligule ovate-oblong, subacute and some-
times bifid at apex, free in the distal 1-3 cm., the leaflets (5—) 7-9, the
petiolules (0.5—) 2-10 cm. long, the blades papyraceous, oblanceolate or
narrowly obovate or elliptic, (5—) 10-34 cm. long, (2.5—) 412 cm. broad,
attenuate to acute at base, acute to acuminate at apex, usually entire or
crenulate at margin on mature leaves but sometimes sparsely denticulate
(teeth 1 or 2 per centimeter, evanescently spinulose), the costa slightly
raised above, prominent beneath, the secondary nerves 8-18 per side,
erecto-patent, often prominulous above and sharply raised beneath, the
veinlet-reticulation immersed or prominulous beneath; inflorescences later-
al on branchlets below terminal clusters of leaves, paniculate-racemose,
glabrous to scurfy-pilose on axes, up to 100 cm. in length, short-peduncu-
late, the main rachis often stout and subflexuose, the primary branches
usually 15-25 in number, alternate or subopposite, usually 15—40 cm. long,
the primary bracts papyraceous to subcoriaceous, oblong, subacute, 3-8
mm. long (or longer on peduncular portion); umbels borne on slender
peduncles 4-7 mm. long at anthesis and up to 15 mm. long in fruit, the
flowers usually 5-10 per umbel, the pedicels 2-4 mm. long, subtended by
minute bracteoles, the umbel-subtending bracteoles up to 5 mm. long;
flowers 2—3.5 mm. long at anthesis, the calyx cupuliform, 1-2 mm. long,
with 5 minute deltoid lobes; petals 5, oblong-deltoid, 1.2—1.7 mm. long,
unguiculate at the acute apex; stamens 5, the filaments slender, at length
about 1 mm. long, the anthers oblong, obtuse, 0.5—0.8 mm. long; ovary-
apex flattened or rounded, the stigmas 5, sessile in flower, papilliform, soon
elongating; fruits depressed-subglobose, at maturity 4-6 mm. long and
broad, conspicuously 5-costate when dried, broadly conical at apex, the
styles free except at base, sharply reflexed, 0.4-0.6 mm. long.

; Type Locatity: Ovalau Fiji; the type is the U.S. Exploring Expedi-
tion specimen cited below.

_ DISTRIBUTION. Endemic to Fiji and probably occurring on most of the
high islands, being common at elevations of about 50—1,130 meters in
various types of forest, crest-thickets, etc. It is usually noted as a slender,
freely branched, or gnarled tree 3-18 meters high, with a trunk up to 20
centimeters in diameter, and with characteristically congested leaves at
the apices of branchlets and profuse inflorescences lateral just below the
leaves. The inflorescence-axes and calyces are purple or purple-tinged; the
petals and filaments are pale green to pale yellow, with nearly white

1963] SMITH & STONE, ARALIACEAE 485

anthers; the disk is green in flower; and the mature fruit is dull purple
to black

LocaL NAMES AND USES. Frequently recorded names on Viti Levu are
sole, sole lewa, sole ngga, and sangole. On Vanua Levu the names ndani-
ndani, kaikai, and kai-i-voli have been noted, and on Ovalau ndai-
ndainga. In Serua Province a decoction of the leaves has been recorded as
having been drunk for lung trouble, and in upland Naitasiri Province the
plant is indicated as an antidote for poison.

Fiji (because of abundance only island and province are cited): Vit1 Levu:
Mpa: Gibbs 580 (pM), Tothill 221 (BIsH, K), 225 (BISH), K), Greenwood 337
(K), 865 (a), Parks 20536 (isu, uc), Gillespie 3714 (BISH, GH, UC), 4348 (BISH,
GH, Ny, UC), Degener 14627 (A, BISH, K, NY, UC, US), Smith 4082 (A, BISH, K,
US), 4095 (A, BISH, K, US), 4875 (A, US), 4952 (A), 5894 (A, BISH, K, NY, us),
6177 (A, BISH, K, NY, US), Fiji Dept. Agr. 2325 (K), 2327 (x, suvA), 7136 (SUVA).
Nanpronca & Navosa: Degener 14901 (A, NY), Smith 5508 (A, BISH, K, NY, US).
SeRUA: St. John 18966 (BIsH, Us), Degener 15147 (A, BISH, K, NY, US), Smith
9190 (us). Namost: Gillespie 2869 (BISH), 3137 (BISH, GH, uc), Smith 8508
(BISH, US), 8627 (BISH, US), Fiji Dept. Agr. 11660 (suvA). Ra: Degener 15354
(A, BISH, K, NY, US), 15439 (A, BISH, K, NY, US), 15510 (A, BISH, K, NY, UC, US).
Narrastri: Jeoward 87 (kK), Bryan 369 (A, BISH, UC, US), Tothill 223 (x), Parks
20075 (BIsH, UC), Gillespie 3449 (BISH, GH, UC), St. John 18286 (BISH, US),
18333 (BISH, US), Smith 6132 (A, BISH, K, NY, US), Fiji Dept. Agr. 119 (suv),
12786 (suva), TatLevu: Smith 7106 (BIsH, US). REWA: Meebold 16554 (x),
Fiji Dept. Agr. 1179 (suva). Kanpavu: Smith 264 (BISH, GH, K, NY, UC, US).
Ovarau: U.S. Expl. Exped. (us 47685 type), Gillespie 4435 (Bish), Smith 7372
(sisH, Us), 7513 (BISH, US). Ncav: Smith 7791 (BISH, US). VANUA Levu:
Meva: Smith 1562 (BISH, GH, K, NY, U
(BISH, GH, K, NY, UC, US), 513 (BISH, GH, K, NY, UC, US), TAVEUNT:
203 (cH, K), Gillespie 4650 (Bisu, UC), Smith 748 (BISH, GH, K, NY, UC, vs),
8237 (BisH, Us). Fry1, without other locality: Harvey (K), Horne 68 (cH, kK),
136 (k), Jeoward 36 (K).

The cited material of this locally abundant species is reasonably uni-
form, but variations in the degree of indument are noteworthy. Most of
the specimens, as to the lower surfaces of leaflets and the inflorescences,
are glabrous, or evanescently puberulent with pale, minute trichomes.

branches (and umbel-peduncles, calyces, and of
on or near the costa and secondaries) more or less persistently scurfy
with ferruginous, irregular, subfimbriate scales 0.1-0.15 mm. in diameter ;
or sometimes such indument is composed of simple, several-celled trichomes
up to 0.2 mm. Jong. As all intermediate degrees of occurrence and persis-
tence of indument are noted, we believe this feature lacks nomenclatural
significance. The following specimens, cited above, have obvious indument:
Gillespie 3137, St. John 18966, Smith 371, 513, 748, 4952, 5894, 8508.
Another feature not included in our description deals with the margin
f many species with com-

pound leaves) have deeply pinnatifid leaflets, of which the lobes may be

rounded, 5~8 per side, and copious

486 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Lobes and teeth are normally lacking from mature leaflets, but rarely the
denticulations persist on younger leaves.

Schefflera vitiensis seems to have the floral parts uniformly five, agree-
ing in that respect with S. euthytricha and S. tannae. From the former it
shows obvious differences in indument and inflorescence-bracts; from the
latter it differs in its ample, spreading inflorescences, and in its leaves with
fewer leaflets, these being obviously acute to acuminate at apex.

6. Schefflera actinostigma A. C. Smith & B. C. Stone, sp. nov.
Pt. IV, Fic. 7.
Schefflera sp. Guillaumin in Jour. Arnold Arb. 12: 263. 1931.

Arbor ad 10 m. alta ubique glabra; foliis digitatis, petiolis crassis 36-50
cm. longis basi incrassatis (vagina et ligula non visis); foliolis 7-10,
petiolulis comparative gracilibus 3—5 cm. longis, laminis subcoriaceis ob-
ovatis vel ellipticis, 11-22 cm. longis, 6-9 cm. latis, basi attenuatis et

utrinque prominulis, rete venularum subimmerso; inflorescentiis sub fructu
racemoso-umbellatis 35 cm. vel ultra longis, pedunculo crasso integro non
viso, radiis primariis paucis divaricatis ad 30 cm. longis pedunculo sub-
complanato ad 13 cm. longo incluso, ramulis tertiariis circiter 10 e verticil-
lo mediano paucis orientibus et apice radii aliis aggregatis graciliter ped-
unculatis (5-10 cm. longis) et medium versus inconspicue geniculatis;
fructibus (forsan immaturis) plerumque 3-6 per umbellulam pedicellis
7-9 mm. longis enatis, subgloboso-urceolatis, 6-8 mm. longis latisque, ob-
tuse et valde 5—7-costatis, 5—7-locularibus, apice truncatis et marginem
calycinum undulatum subpersistentem circiter 1 mm. altum gerentibus;
stylis gracilibus basi divergentibus rotatis 1-1.5 mm. longis.

TyPE Locaity: Anelgauhat Bay, Aneityum, New Hebrides; the type is
Kajewski 758.

DIsTRIBUTION. Known only from the type collection and presumably
endemic to the New Hebrides. The available specimens were taken from
a small tree about 10 meters high, said to be common in rain-forest at an
elevation of about 60 meters; the fruit was stated to be black when
mature but only about half grown when collected.

New Hebrides. ANEITyuM: Anelgauhat Bay, February 13, 1929, Kajewski
758 (A type, K, NY, photos at BISH, US).

Although to be placed among the Sche flerae of our area with compound-
umbellate inflorescences, this new species is noteworthy as being transi-
tional in character. Its primary inflorescence-rays, instead of radiating
exclusively from the apex of the peduncle, are additionally clustered at a
mid-point of the rachis; they are also jointed near the middle, thus suggest-
ing further an intermediate stage between the paniculate-racemose an
strictly umbellate inflorescences of Schefflera. The new species further

1968 | SMITH & STONE, ARALIACEAE 487

differs from its Fijian allies, S. costata and S, seemanniana, in having its
styles free in fruit, radiating on the flattened stylopodium that is sur-
mounted by an obvious calycular rim.

7. Schefflera costata A. C. Sm. in Bishop Mus. Bull. 141: 119. fig. 62.
1936; J. W. Parham, PI. Fiji Isl. 86. 1964. PL. IV, Fic. 8.

As this species is still known only from the type collection, the original
description requires supplementation only by our illustration and the
following few notes:

Petioles expanded at base into a clasping sheath, the ligule prominent,
oblong, free in the distal 3-4 cm., suberose at the obtuse apex, at length
caducous and leaving a conspicuous scar; leaflets usually 4-9 (sometimes
only 2), the blade up to 12 % 7 cm.; infructescence twice-divided, the
peduncles of primary rays 3-4 cm. long, of secondary rays 1.5—2 cm. long;
costae and locules of fruit 8-12, the stylopodium broadly conical into a
stout stylar column about 1.5 mm. long and 1—1.5 mm. in apical diameter,
the stigmas conspicuous, about 0.5 mm. long and laterally decurrent on the
column.

TYPE LOCALITY: Taveuni, Fiji; the type is Smith 886, cited below.

DistrisuTion. Apparently endemic to Fiji and known only from the
type collection; noted as a tree 5 meters high in forest at an elevation of
600-900 meters.

Fiji. TAvEuNI: Western slope between Somosomo and Wairiki, Smith 886
(BISH type, GH, K, NY, UC, US).

It is surprising that this very distinct species has been collected only
once, as its Taveuni locale has been visited by several collectors. From its
only close relative, S. seemanniana, it differs in its conspicuous foliar ligule,
its long-acuminate leaflet-apices, its increased number of ovary-locules,
and its conical, rather than slenderly cylindric, column of connate styles.

8. Schefflera seemanniana A, C. Sm. in Bishop Mus. Bull. 141: 118.
1936; J. W. Parham, PI. Fiji Isl. 86. 1964. PL. IV, Fics. 9, 10.
Agalma vitiensis Seem. Fl. Vit. 116. 1865; Anon. in Kew Bull. 1888: 91. 1888.
Non Scheflera vitiensis Seem. (1865).
Agalina vitiensis Anon. in Gartenfl. 36: 71. 1887.
Heptapleurum vitiense Seem. ex Drake, Ill. Fl. Ins. Mar. Pac. 183. 1890;
B. & H. f. ex Drake, op. cit. 409. 1892; Gillespie in Bishop Mus. Bull. 91:
24. fig. 27. 1932.

A tree, glabrous throughout; leaves digitately compound, the eed
slender, (5—) 10-22 cm. long, expanded to a coriaceous clasping sheath,
the ligule rounded, often erose toward apex, free in the distal 3-10 a
the leaflets 4-8 (rarely fewer), the petiolules 1-3.5 cm. long, narrowly
alate distally, the blades papyraceous to subcoriaceous, elliptic to os ga
(3.5-) 6-17 cm. long, (2—) 3.5-8 cm. broad, acute to attenuate at oH
and decurrent on the petiolule, rounded to obtusely short-acuminate a

488 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

apex, entire and narrowly recurved at margin, the costa slightly raised
above and prominent beneath, the secondary nerves numerous, 20—30 per
side and interspersed with others nearly as obvious, spreading, prominu-
lous on both sides or plane above, the veinlet-reticulation usually im-
mersed above and prominulous beneath; inflorescence compound-umbel-
late, usually 2 (sometimes 1 or 3) times divided, the peduncle usually
stout, 1-6 cm. long, bearing at apex 3—7 primary rays, these with compara-
tively slender peduncles 3-9 cm. long, the secondary rays (if present) with
peduncles 2-6 cm. long; ultimate umbels with 5—12 flowers (primary rays
sometimes terminated by an umbel of perfect flowers and also bearing
secondary rays); flowers perfect or staminate, sometimes mixed in the
same umbel, the pedicels usually 5-12 mm. long at anthesis (to 20 mm.
in fruit), the ray- and flower-subtending bracts subcoriaceous, orbicular-
ovate, inconspicuous, 1-2 mm. long, laterally contiguous, the pedicels also
subtended by lanceolate evanescent bracteoles about 1 mm. long; perfect
flowers about 1 cm. long at anthesis, the calyx narrowly urceolate, 4-7 mm.
long and 4-5 mm. in diameter at the short, flaring, obscurely sinuate rim;
petals 5 (or 6), subpersistently calyptrate, oblong-deltoid, 4-5 « 3-4 mm.,
apically thickened and unguiculate; stamens 5 (or 6), the filaments slight-
ly flattened, about 2 mm. long, the anthers versatile, oblong, obtuse, 1.5—
2.5 mm. long; ovary-apex conical, produced into a conspicuous stylopodi-
um of firmly connate styles about 1 mm. long at anthesis; staminate flowers
similar but the calyx much smaller, subrotate, the inferior portion of the
ovary obviously sterile; fruits ovoid-subglobose, at apparent maturity
(but dried) 11-13 mm. long (excluding stylar column) and 9-11 mm
broad, sharply 5—7-costate, the calycular rim obvious about 2 mm. below
base of stylar column, the stylopodium conical, terminated by a slender
stylar column about 2 mm. long

Type LOCALITY: Viti Levu, Fiji; the type is Graeffe 38, deposited in
the National Herbarium of Victoria at Melbourne. We have not seen the
type, but an isotype is cited below.

DistRIBUTION. Endemic to Fiji and perhaps to be expected on most of
the high islands, but at present noted infrequently at elevations of 150—
900 meters in dense forest or in crest-thickets. Specimens have been ob-
tained from trees 6-25 meters high, with a trunk up to 70 centimeters in
diameter; the calyx is greenish or bright green; the petals and filaments
greenish yellow or pale green; the anthers pale yellow; and the fruit black
at maturity.

LocaL NAME. Sole has been noted for a single specimen from Ovalau.

Fiji. Virt Levu: (without further locality) Graeffe 38 (Mm isotype, us photo).
Mpa: Mt. Nandende Levu, Fiji Dept. Agr. 14058 (sisH, suva). Namost: North
of Wainavindrau Creek, between Korombasambasanga Range and Mt. Naitara-
ndamu, Smith 8480 (BisH, us); hills bordering Wainavindrau Creek, vicinity of
Wainimakutu, Smith 8543 (BisH, us); Mt. Voma, Fiji Dept. Agr. 1717 (A, SUVA),
Gillespie 2668 (ptsH). Natrasirt: Central Road, Tothill 220 (x, photos at BISH,
US). Rewa: Near summit of Mt. Korombamba, Gillespie 2357 (BIsH, GH).
Kanpavu: Mt. Mbuke Levu, Smith 242 (Bis, GH, K, NY, UC, US). OVALAU:

1968 | SMITH & STONE, ARALIACEAE 489

U.S. Expl. Exped. (us 73841); summit of Mt. Tana Lailai and adjacent ridge,
Smith 7703 (BisH, us). Ncav: Hills east of Herald Bay, inland from Sawaieke,
Smith 7852 (BISH, US).

Schefflera seemanniana is variable in its inflorescences, although basical-
ly these are strictly compound-umbellate. The primary rays usually bear
flowering umbels at their apices, but sometimes they are directly florifer-
ous, and sometimes they also are compound-umbellate (i.e. the inflores-
cence may be once, twice, or three times compound). Additionally, a
flowering umbel may sometimes bear complete umbels lateral to its own
flowers. It is probable that the central flowers of each umbel tend to be
perfect and the peripheral flowers staminate, but this condition is not
uniform, Leaf- and fruit-characters amply distinguish this species from
the allied S. costata.

9. Schefflera sp. Px. IV, Fic. 11.

A tree known only from the infructescence, this compound-umbellate,
3-times divided, the peduncle 5~10 cm. long, bearing at apex 5 or 6 primary
rays, these with slender peduncles 6-12 cm. long, the secondary rays
(6-8 in number) with peduncles 4-9 cm. long; umbels with 4-11 fruits
(on pedicels 11-18 mm. long), these depressed-subglobose, 7-9 mm. long
(excluding stylar column), 8-10 mm. broad, sharply 6—9-costate, the
minute calycular rim evident 1-3 mm. below the rounded apex, the styles
firmly connate in a column about 2 mm. long.

Fiji. Mea: Nauwanga, vicinity of Nandarivatu, alt. 750-900 m., Degener
14540a (A, BISH, US) (as to infructescence only, excl. leaves; sole).

The infructescence here described apparently represents an undescribed
species, but it was collected from the forest floor together with fallen
leaves of a Plerandra, and the material is not a suitable basis for a new
species. From the allied S. seemanniana, this inadequately known taxon
differs in its more ample and spreading infructescence and smaller fruits,
which have more numerous costae and pyrenes, and of which the stylopo-
dium is rounded rather than conical below the stylar column.

7. Brassaia Endl. Nov. Stirp. Dec. 89. 1839; Hutchinson, Gen. Fl. Pl.
2: 73, 622. 1967.

As greatly expanded by Hutchinson in 1967, Brassaia has — 64
species and is said to have a range from India and western China tl te
Malaysia to the Philippines, New Guinea, and northeastern Australia (t :
reference to Hawaii being only to a cultivated plant). However, it mus
be noted that the long list of new combinations presented by Hutchinson
in his 1967 work includes eleven from South America, which is not in-
dicated as part of the range. It seems very unlikely that ee: es ue
family will accept this disposition of species previously heey to oa
flera (including Sciadophyllum, Heptapleurum, Cephalosc nee, : Hf
We believe it more likely that Brassaia will eventually be submerged 1

490 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Schefflera, as indeed suggested by Harms (under his section Cephaloschef-
flera) and Airy Shaw (in Willis, Dict. Fl. Pl. & Ferns ed. 7. 155. 1966).
No indigenous species of this complex occur in our area, but as a matter
of convenience we retain Brassaia for its widely cultivated type species.

1. Brassaia actinophylla Endl. Nov. Stirp. Dec. 89. 1839; J. W. Par-
ham, PI. Fiji Isl. 83. 1964.

Schefflera actinophylla Harms in E. & P. Nat. Pflanzenfam. 3(8): 36. 1894.

TYPE LOCALITY: Queensland, Australia.

DIstTRIBUTION, Widely cultivated in tropical areas, and doubtless often
naturalized. It commonly attains a height of 15 meters or more.

LocaL NAMES. Queensland umbrella tree, umbrella tree, and octopus
tree are used for this striking plant as cultivated.

Fiji. Vitt Levu: Natrastrt: Nasinu, Fiji Dept. Agr. 11241 (BISH, SUVA).

The species is certainly more widely grown in our area than the above
record would indicate. In Fiji, as in Hawaii, it is a frequent street tree
or garden tree and it is also becoming naturalized. The cited specimen
is from a naturalized plant, and the species is now commonly seen in
second growth areas in southeastern Viti Levu.

8. Dizygotheca N. E. Br. in Kew Bull. 1892: 197. 1892; Harms in E.
& P. Nat. Pflanzenfam. 3(8): 31. 1894; Hutchinson, Gen. Fl. Pl. 2:
70. 1967.

Both Harms and Hutchinson maintain Dizygotheca as a genus of the
general affinity of Schefflera; it is typified by D. vieillardii (Baill.) Vig-
uier (D. nilssonii N. E. Br.) and includes 12 to 17 species, usually indi-
cated as New Caledonian. However, Guillaumin has noted the apparently
indigenous occurrence of D. elegantissima in the New Hebrides, and this
species has also been noted as cultivated in Fiji.

Dizygotheca is characterized by having digitately compound leaves,
pedicellate flowers, 5 petals and stamens, and a 10-locular ovary with
free styles. All of these characters also occur in Schefflera, where 10 or
more ovary-locules are sometimes found. Perhaps the only consistent
character is the 4—locular anther of Dizygotheca (cf. Harms in op. cit. 9.
fig. 2, A-C), contrasting with the the 2-locular anther of Schefflera.

1. Dizygotheca elegantissima (Hort.) Viguier & Guillaumin in Not.
Syst, 2: 258. 1912; Guillaumin in Bull. Mus. Hist. Nat. II. 9: 289,
as Dyzygotheca e. 1937; J. W. Parham, PI. Fiji Isl. 83. 1964.
Aralia elegantissima Hort. in Gard. Chron. 1873: 782. 1873.
We have seen no material of this species from our area, but include
the record on the basis of the mention by Guillaumin and Parham.

Type Locatity: The type collection was a cultivated plant often as-

1968 ] SMITH & STONE, ARALIACEAE 491

sumed to have been introduced from New Caledonia, but quite possibly
it is indigenous to the New Hebrides.

DIsTRIBUTION. Widely cultivated in tropical areas and in temperate
greenhouses; the original distribution is conjectural.

Locat Namgs. Certain horticultural plants known as aralia are referable
to this and perhaps other species of Dizygotheca. Neal (Gard. Hawaii
654. 1965) lists the plant cultivated in Hawaii as D. elegantissima as
false aralia.

New Hebrides. AmpryM: (Not seen, but a specimen collected by de La Riie, at
an altitude of 650-800 m., apparently indigenous, is referred here by Guillau-
min

in).
Fiji. (Not seen, but noted by Parham as introduced and commonly cultivated).

The species of Dizygotheca will doubtless remain confused, both in cir-
cumscription and in origin, until a detailed study of the cultivated variants
is made in connection with a new search for wild relatives in at least
New Caledonia and the New Hebrides. A second species sometimes indi-
cated as having come from the latter archipelago is D. reginae (Hort.)
Hemsl. L. H. Bailey (Stand. Cycl. Hort. 1062, 2746. 1925 and later edi-
tions) has presented an informative discussion of this group of horticul-
tural ‘“‘aralias.” A second Dizygotheca from Ambrym, New Hebrides, is
listed as a young form by Guillaumin in his 1937 paper on de La Riie’s col-
lections.

9. Boerlagiodendron Harms in E. & P. Nat. Pflanzenfam. 3(8): 31.
1894, in Bot. Jahrb. 56: 377. 1920; Hutchinson, Gen. FI. Pl. 2: 72.
1967.

Eschweileria Zipp. ex Boerl. in Ann. Jard. Bot. Buitenzorg. 6: 112. 1887, Non

Eschweilera Mart. (1828).

Boerlagiodendron, typified by B. palmatum (Zipp.) Harms, is a genus
of about 40 species extending from Formosa, the Philippines, the Marianas
and Palau Islands, and Malaysia through New Guinea and the Solomons
to the New Hebrides, where its range is terminated by a single species.
It is well represented in New Guinea, from which at least 17 species have

been described.

1. Boerlagiodendron orientale Guillaumin in Jour. Linn. Soc. Bot.
51: 554. 1938.

Having seen no other material of this species than the type, we refrain
from adding to the original description.

Type tocatity: Espiritu Santo, New Hebrides; the type 1s I, & 2,
wpe ial it pete only from the type collection. Recorded as oc-
curring in rain-forest and as having large, long-petioled leaves and a com-
pound-umbellate inflorescence 30 centimeters in diameter.

Locat name. Varakii was recorded by the collectors.

492 JOURNAL OF THE ARNOLD ARBORETUM [voL, 49
New Hebrides. Esprritu Santo: Hog harbour, January 25, 1934, J. & Z.
Baker 258 (BM type).
The species is noted by Guillaumin as a relative of B. novo-guineense
(Scheff.) Harms, with which it agrees in having pinnatifid leaves and
pedicellate flowers; it differs in having only 5 ovary-locules.

: BCs:
UNIVERSITY OF HAwall UNIVERSITY OF MALAYA
HonoLutvu, HAwati KvuaLa LUMPUR
MALAYSIA

EXPLANATION OF PLATES

PLATE I

Fics. 1-5. Polyscias joskei: 1-4 from Smith 4975, 5 from Smith 6161. 1, leaf-
let, X Fal 1 ultimate axis of inflorescence, x i. and 4, flowers, X 5; 5, fruit,
: . 6-8. Polyscias neo-ebudarum: all "from Kajewski 291. 6, leaflet,
<4: 7, ree ett 82.8, Feuit,-< "5, Fica. 9-12. Polyscias eect oa al ae om
Smith 4514. 9, leaflet, x iy; 10, umbel with one attached flower, X a , de-
tails of flowers, X 5; 12 immature fruit with persistent neti xX 5 Fic i 14.
Ti begs seutellaria: from Anderson (lectotype of P. pinnata). 13, details of
, X 5; 14, detailed view of two of the four styles of one flower, —

silage Fic a. 17. Polyscias samoensis: all from Setchell 374. 15, bud just

before ahem 5; 16, flower with some parts removed, x 5; 17, fruit, X 5.

PLATE II
Fics. 1-9. Polyscias multijuga: 1, 2, and i from Smith 122, 3-6 from See-
mann 505, after Fl. Vit. 1, leaflet, x %; 2, ultimate axis of inflorescence, X
1; 3 and 4, staminate flower, ae 5. 5 and 6, tee me flower, X 5; 8, yo

rui ew 5: :

carticate: all from Smith 5804. 10, leaflet, X %; 11, part of inflorescence, X
%4; 12, ultimate branch of inflorescence with terminal ’and lateral umbels, X 1;
13, nerect flower, X 5; 14, fruit, x 5.

PLATE III
Fics. 1-6. Polyscias reineckei: all from Christophersen & Hume 2616. 1,
flower, X 5; 2, stamens, X 5; 3, upper part of calyx and stylopodium, X 5; 4,
pay east top view of stigmas; 5, top view of fruit, X 5; 6, fruit, X 5. Fics. 7-9.
Polyscias fruticosa: 7 from ‘Smith 7496, 8 and 9 from Judd s. n. (Hawaii). 7,
oe age: of bay X 5; 8, top view of trilocular fruit, X 5; 9, bilocular fruit,
ics. ;

view of fruit, X 5; 13, fruit,

PLATE IV
F Ics. 1 and 2. Schefflera vitiensis: 1 from Smith 8627, 2 ~ Parks 20075.
1, details of flowers, left figures X 2%, right figure X 5; 2, fruit, x 214. Fics.
3-5. Schefflera samoensis : ; and 4 from Vaupel 351, 5 from. Christophersen 234.
3, details of flowers, upper left figures X ee others : 4; prit sd ange
; it

tata: fruit, from Smith 886, X 2%. Fics. 9 and 10. Scheflera seemannia
from Smith 7852 (26) and 7703 (8), 10 from Smith 242. 9, details of pe

1968 | SMITH & STONE, ARALIACEAE 493
. ed 10, fruit, X 2%. Fic. 11, Schefflera sp.: fruit, from Degener 14540a,
2.

hefflera tannae: from Kajewski 131. Habit and inflorescence, X 14; portion
of hii Li xX 2.
PLATE VI
Schefflera euthytricha: from Smith 8908. Tip of ae and leaf, and de-
tached inflorescence, the latter partly diagrammatic, x 4,
PLATE VII
Fics. 1 and 2. Plerandra vitiensis. 1, ~ from Seemann 207 (type) ; 4, Ste
th x 2%. Fics. 3 and 4

:
Plerandra victoriae, fruit, from Gillespie 4122, X 2%. Fics. 6-9. Plerandra
grayi: all from Smith 8523. 6, fruit, X 214; 7, apex of fruit, X 2%; 8, stamen-
scars, enlarged; 9, pistillate flower, X 2u.
PLATE VIII
and 2. Plerandra Pek both from Smith 6896. 1, immature
, e 1. Fics. 3-7. Plerandra insolita: 3-5 fro
Smith 9166, 6 and 7 from vito 8912. 3, fruit, X 2%; 4, ray of inflorescence,
x 1; 5, pistillate flower, X 214; 6, staminate flower, x2 4: 7, 7, staminate flower
with petals and most stamens detached, X 2%.

PLATE I

Jour. ARNOLD Ars. VOL. 49

Smi1TH & STONE, ARALIACEAE

Jour. ARNOLD Ars. VoL. 49 Prate II

Smitu & STONE, ARALIACEAE

Jour. ARNOLD Ars. VOL. 49 Pate Ill

SmitH & Stone, ARALIACEAE

Jour. ARNOLD Ars. VoL. 49 PLATE IV

SmitH & STONE, ARALIACEAE

Jour. ARNOLD Ars. VOL. 49 PLATE V

ze. \ ss . |
of *¥o>S SN |] /f

as

“a
Te
= 4

A

SmitH & Stone, ARALIACEAE

Jour. ARNOLD Ars. VoL. 49

PuatTe VI

Smitu & STONE, ARALIACEAE

Jour. Arnotp Ars. VoL. 49 Prate VII

SMITH & STONE, ARALIACEAE

Pate VIII

Jour. ARNOLD ARB. VoL. 49

SmituH & STONE, ARALIACEAE

502 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

SORBUS AND THE PROBLEM OF GENERIC TYPIFICATION

+ GEORGE K. BrizIcKy

HAVING BY CHANCE opened volume two of the Ray Society’s facsimile
(1959) of Linnaeus’s Species Plantarum at page xi of W. T. Stearn’s ex-
cellent preface I read: “The printing of an index to the Species Plantarum
has thus provided a convenient opportunity for summarizing opinions on
the typification of most Linnaean genera. The need for this is emphasized
by Article 22 of the /nternational Code of Botanical Nomenclature 1956
which rules that ‘the subgenus or section including the type species of the
correct name of the genus to which it is assigned repeats that name un-
altered as its epithet’ . . . Its effect may be exemplified by the nomencla-
ture of the sections of the genus Sorbus. Thus, if Sorbus domestica L. is
accepted as the lectotype of Sorbus, as, following Hitchcock and Green,
it should be, then section Cormus (Spach) C. K. Schneider to which it
belongs must be renamed section Sorbus and section Sorbus Pers. (1806)
becomes section Sorbaria Schauer. In genera, such as Sorbus, for which
more than one lectotype has been proposed, it is undesirable to disturb
established nemenclatiire until general agreement has been reached on a
suitable lectoty

Ithough I value most highly Stearn’s “summarizing opinions on the
typification of most Linnaean genera,” I can not agree with his conclud-
ing sentence in the quotation above for a number of reasons: 1) following
Stearn’s suggestion not to disturb established nomenclature until general
agreement has been reached on a suitable lectotype in genera for which
more than one lectotype has been proposed means to admit and tolerate
two or three different type species and,-eventually, two or three different
typical subgenera and/or sections in about 113 Linnaean genera; 2) of
two or three lectotypes proposed for a generic name only one is correct,
i.e., chosen in conformity with the International Code of Botanical Nomen-
clature: 3) when a correctly chosen lectotype is found, this choice must
be followed, and no general agreement is necessary for its adoption; and
4) with regard to the importance of typification of genera, verifying the
lectotype choices and/or typifications of the genera should be encouraged,
not discouraged. For convenience in illustrating these points I shall use the
same example as that given by Stearn, i.e., the typification of Sorbus L.

In his Species Plantarum (1: 477. 1753), Linnaeus established the genus
Sorbus with two species, S. aucuparia and S. domestica. In 1908, Britton
(N. Am. Trees 427) designated Sorbus domestica L. as type of the generic

¥ Dr. Brizicky “ne completed the manuscript for this paper some time before his
death on June 15, 1

1968 | BRIZICKY, SORBUS 503

name. This choice of lectotype, made without explanation, was adopted
by Britton and Brown in 1913 (Illus. Fl. No. US. 2: 287). In 1929,
Hitchcock & Green (Int. Bot. Congr. Cambridge 1930. Nomencl. Propos.
Brit. Bot. p. 158) independently proposed S. domestica L. as the lectotype
species of Sorbus L., with the following argumentation: “The choice of
standard-species lies between S. Aucuparia and S. domestica. Sorbus was
restricted to S. domestica in 1789 by Medicus, Phil. Bot. 138, who treated
S. Aucuparia as the type of an independent genus Aucuparia Rivin, The
type of Tournefort’s Sorbus was evidently S. domestica judging from the
vernacular name “‘Sorbier” and from citations given by him. Therefore,
S. domestica is chosen as the standard-species.”’

Britton’s choice of S. domestica as the lectotype of Sorbus, supported by
Hitchcock & Green’s argumentation, has generally been adopted, except by
Rehder, who designated S. aucuparia L. as the lectotype species of Sorbus,
but without explanation of this choice (Bibliogr. Cult. Trees Shrubs 252.
1949). The general adoption of Britton’s choice of S. domestica does not
necessarily mean that his selection of lectotype was correct from the
standpoint of the International Code, and, since it was challenged by that
of Rehder, the typification of Sorbus must be reviewed.

The genus Sorbus of Linnaeus, originally founded by him in 1737 (Gen.
Pl. 144), was described from a living plant as having flowers with three
Styles and twenty stamens inserted on the calyx. In the same year Lin-
naeus (Hort. Cliffort. 188) also established the single species “Sorbus
foliis pinnatis” (with the synonyms: “Sorbus sylvestris. Dalech. hist. 532.
Sorbus sylvestris alpina. Lod. hist. 544. Sorbus sylvestris, foliis domesticae
similis, Bauh. pin. 415. Sorbus aucuparia. Bauh. hist. 1. p. 62. Boerh.
lugdb. 2. p. 248. Aucuparia rivini. Rupp. jen. 112.”). There was appended
[var.] a. “Sorbus sativa. Bauh. pin. 415.” (with the synonyms: “Sorbus
domestica. Lob. hist. 544. Sorbus legitima. Clus. hist. 1. p. 10. Sorbus.
Bauh. hist. 1. p. 59. Dalech. hist. 330. Dod. pempt. 803.”). Although a
detailed description of the species was lacking, the references to the syn-
onyms characterized the species proper and its variety a quite well. In
addition, the ranges were given separately for the species proper and the
[var.] a: “Crescit in Lapponia, Norvegia, F inlandia, Suecia, Dania,
Germania, Helvetia, Anglia, Gallia & locis umbrosis & spongiosis, at (a)
in Italia, Germania, Helvetia.” In Species Plantarum (1: 477. 1753) the
“Sorbus foliis pinnatis” of Hortus Cliffortianus received the name Sorbus
aucuparia, and its [var.] a was raised to the rank of species with the name
Sorbus domestica. :

The occurrence of three styles (“Styli tres . . .”), i.e., of three carpels,
in Sorbus was considered by Linnaeus to be a generic character, on the
basis of which Sorbus was placed by him in the special order Trigynia of
the Class Icosandria (Gen. PI. ed. 5. p. xxiii). In 1777, in his sgeonieie
systematis sexualis Linnaei, John | John] Miller gave a rather detai ve
description of Sorbus (based on S. aucuparia) as the representative 0
Icosandria-Trigynia, and this was accompanied by an excellent detailed

Synia,

504 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

drawing of S. aucuparia. The pistil was described and illustrated as having
three styles and the cross section of fruit showed three locules.

In 1793, Medicus (Geschichte der Botanik, 86, 87) divided Sorbus L.
into two distinct genera, Aucuparia, a new genus, based on S. aucuparia
L. (Aucuparia silvestris Medic.) and Sorbus, based on S. domestica L.
The two genera differed in the following characters: Aucuparia, ‘““Zahl der

Griffel drei bis fiinf . . . Drei bis fiinf Pericarpien . . . In jedem Gefache
zwei Saamen”; Sorbus, “Fiinf Griffel, selten weniger. Innerhalb dem
Fleische sitzt ein fiinffacherichtes Pericarpium . . . In jedem Gefache ein
aame.”’

In 1834, Spach (Hist. Nat. Vég. 2: 91-98) also divided the original
Sorbus L. into two genera: Sorbus L., including S. aucuparia, S. hybrida
L., S. lanuginosa Kit., S. foliosa Wall., and S. americana Pursh; and
Cormus Spach, a new genus, based on Sorbus domestica L. The distinguish-
ing characters were: Sorbus, “Ovaire adhérent, 4 3 (quelquefois 4 2 ou
4) loges biovulées. Styles en méme nombre que les loges de lVovaire . . .
Pyridion globuleux ou subturbiné, ombiliqué aux deux bouts, 4 2—4 loges
l-spermes; endocarpe mince, crustacé”’; Cormus, ‘““Ovaire adhérent, a 5
loges biovulées. Styles 5, filiformes . . . Pyridion pyriforme ou subglo-
buleux, subquinquéloculaire; endocarpe membraneux.

Neither Medicus’s nor Spach’s segregation has generally been adopted,
and Sorbus, including both original species, has usually been treated either
as a subgenus of Pyrus L. or as a distinct genus (subdivided into two to
four subgenera and/or sections). Although some authors have placed both
of the original Linnaean species in the same subgenus (e.g., Persoon, Synop.
Pl. 2: 38. 1806; Koch, Hort. Dendrol. 177. 1853) or in the same section
(e.g., Dumortier, Florula Belg. 93. 1827; Neilreich, Fl, Nieder-Oesterr.
886. 1858), others have separated the two and placed each of them in a
different subgenus (e.g., Duchartre in D’Orbigny, Dict. Univ. Hist. Nat.
11: 685. 1848) or in a different section (e.g., Boissier, Fl. Orient. 2: 657.
1872). Only Decaisne (Nouv. Arch. Mus. Hist. Nat. Paris 10: 156, 157.
1874) and Koehne (Gartenflora 40: 35, 39. 1891 and Deutsch. Dendrol.
246, 254. 1893) followed Spach in recognizing Sorbus L. and Cormus
Spach as distinct genera (which Koehne even placed in different subdivi-
sions of the family, Sorboideae and Maloideae, respectively).

After this brief review of the establishment of the genus and some facts
important for its typification, it is pertinent to return to Hitchcock and
Green’s argument in support of the choice of Sorbus domestica as the
lectotype species of the genus. Their main points were these: 1) “Sorbus
was restricted to S. domestica in 1789 by Medicus, Phil. Bot. 138, who
treated S. Aucuparia as the type of an independent genus Aucuparia
Rivin.” and 2) “The type of Tournefort’s Sorbus was evidently S. domes-
tica.”. The second point may be dismissed here as irrelevant, because one
must consider the typification of Sorbus of Linnaeus, 1753, and not of that
of Tournefort. The first point, however, deserves a detailed discussion.

“One of the first things to do in selecting a type is to exclude from con-

1968 | BRIZICKY, SORBUS 505

sideration those species that definitely disagree with the generic descrip-
tion,” wrote Hitchcock (Am. Jour. Bot. 10: 511. 1923) a few years before
his and Green’s proposal of S. domestica as the standard-species of Sorbus.
Later, in the preface to their “Standard-species of Linnaean genera of
Phanerogamae (1753-54) (Int. Bot. Congr. Cambridge 1930. Nomencl.
Propos. Brit. Bot. 112. 1929) Hitchcock and Green mentioned this and
another criterion for the typification of genera accepted in the Type-basis
Code, “[Art.] (6c) species which definitely disagree with the generic
description (provided others agree) are to be excluded from consideration;
(7a 1) the type-species is often indicated by closer agreement with the
generic description.” Regrettably, these very sound criteria were not util-
ized by Hitchcock and Green in their choice of the lectotype species of
Sorbus, for they unquestioningly accepted Medicus’s segregation of Au-
cuparia as ‘‘typification by elimination.”’ The most recent /nternational
Code of Botanical Nomenclature (1966) is, however, very explicit in
regard to the typification of taxa, providing a “Guide for determination of
types” (pp. 71, 72), and Sorbus must be typified as outlined below.

There can be hardly a doubt that Sorbus L., of 1737 (Gen. Pl. 144), was
based on and its generic description drawn from the “Sorbus foliis pin-
natis” proper (Hort. Cliffort. 188) and not from its variety a. Since the
generic description of Sorbus in the fifth edition of Linnaeus’s Genera
Plantarum (1754, p. 213) was identical with that of the first, we may
assume that the typical species must have remained the same, i.e., “Sorbus
foliis pinnatis” of Hortus Cliffortianus, 1737 = Sorbus aucuparia L. of
Species Plantarum, 1753.

Moreover, after describing Sorbus as having three styles, Linnaeus
placed it in the order Trigynia of Icosandria. Miller (Illus. Syst. Sexual.
Linnaei 1777) exemplified this order by Sorbus aucuparia, the pistil of
which was described and illustrated as having three styles. Medicus
(Gesch. Bot. 1793; see above) mentioned three to five styles and three to
five pericarpia in S. aucuparia, and five, rarely fewer, styles and a five-
locular pericarpium in S. domestica. His observations, except the occur-
rence of five styles and five pericarpia in S, aucuparia, have been con-
firmed by numerous taxonomists. Thus, Spach (Hist. Nat. Veg. 2: 91-98.
1834) mentioned the three-, sometimes two- or four-locular ovary in
Sorbus, as represented by S. aucuparia, and the five-locular ovary in
Cormus, based on S. domestica. Diill (Die Sorbus-Arten und ihre Bastarde
in Bayern und Thiiringen. Ber. Bayer. Bot. Ges. a4: 11-65. 1961) speaks
of “mostly 5 carpels and styles” in S. domestica and of “ovary with 3,
rarely 4 carpels” in S. aucuparia [translation supplied}. Kovanda (F nei
and fruit morphology of Sorbus in correlation to the taxonomy of the
genus. Preslia 33: 1-16. 1961) reports, “Most often 3 carpels, rarely 4,
in singular cases 2” in S. aucuparia, while in S. domestica There are
always 5 carpels. Their number is absolutely constant and is not subject
to any fluctuations.” eu ‘ge

These old and recent data regarding the gynoecium in the two original
species of Sorbus clearly demonstrate that S. aucuparia definitely agrees

506 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

and S. domestica disagrees with the generic description of Sorbus L. (1754)
in the characters of the gynoecium. Consequently, S. domestica can not
be the type of Sorbus, and Medicus’s Aucuparia, based on the same type
as Sorbus L. (i.e., on S. aucuparia), is illegitimate and must be reduced
to synonymy with the latter. Sorbus Medic. then becomes a later hom-
onym of Sorbus L. In fact, Miller (1777) had already indicated the typifi-
cation of this genus by S. aucuparia. Spach (1834), too, correctly based
Sorbus on S. aucuparia in segregating S. domestica as a distinct genus,
Cormus. Also Duchartre (in D’Orbigny, Dict. Univ. Hist. Nat. 11: 685.
1848), Beck von Mannagetta (in Reichenbach, Ic. Fl. Germ. Helvet.
25(1): 35, 36. 1910?), and Boissier (Fl. Orient. 2: 657. 1872) correctly
based the typical subdivision of Sorbus on S. aucuparia with “ovarium
triloculare (raro 2—4-loculare)” (Boissier), while S. domestica was placed
in the subgenus (Duchartre, Beck) or section (Boissier) Cormus with
“ovarium quinqueloculare, styli 5” (Boissier). Bentham & Hooker (Gen.
Pl. 1: 626. 1865) regarded three styles as a distinctive character of Sorbus,
and Beck von Mannagetta (FI. Nieder-Osterr. 708, footnote. 1892) stated
that “Linnés Gattung Sorbus begreift nach der Beschreibung (Styli 3,
bacca globosa) wohl nur S. aucuparia.”’

Thus, the situation regarding the typification of Sorbus was clear and
unquestionable until 1908, when Britton, contrary to both the then cur-
rent usage and Linnaeus’s protologue, completely reversed it and created
the basis for confusion by his choice of Sorbus domestica as the lectotype
of Sorbus. According to the International Code (1966, Guide for the
determination of types, p. 72, point 4f), “The first choice of a lectotype
must be followed by subsequent workers (Art. 8) unless the original ma-
terial is rediscovered, or unless it can be shown that the choice was based
upon a misinterpretation of the protologue.” It is clear that the first
formal choice of the lectotype species of Sorbus, made by Britton, can not
be followed because it was based upon “a misinterpretation of the proto-
logue,” ie., upon disregard of an important part of the original generic
description. Thus, as already shown, Sorbus domestica disagrees with the
statement of the generic description “Styli 3” and with the position of the
genus in /cosandria-Trigynia. Rehder’s choice (Bibliogr. Cult. Trees
Shrubs 252. 1949) of Sorbus aucuparia as the lectotype of Sorbus, although
tardy and unexplained by him, turns out to be correct since S. aucuparia,
with its usually three-carpellate gynoecium with three styles, entirely fits
the generic description and the position of the genus in Jcosandria-Tri-
gynia. The circumstance that Medicus segregated S. aucuparia as a dis-
tinct genus can not affect the typification, since “If it can be shown that
the element best fitting the protologue has been removed, it should be
se and treated as the lectotype” (Int. Code Bot. Nomencl. 72, point
e

It is rather unfortunate that the authors of such important reference
works as Hutchinson’s Genera of Flowering Plants (vol. 1, 1964) and
Schultze-Motel’s Verzeichnis forstlich kultivierter Pflanzenarten (Kultur-
pflanze Beih. 4. 1966), and some recent monographers of the genus, such

1968 | BRIZICKY, SORBUS 507

as Kovanda (Preslia 33: 1-16. 1961) and Diill (Ber. Bayer. Bot. Ges.
34: 11-65. 1961), have adopted Britton’s (and Hitchcock and Green’s )
choice of Sorbus domestica as the lectotype of Sorbus without verification.
(For the typification of Melochia L., another genus for which two lecto-
types have been proposed, see Brizicky, Jour. Arnold Arb. 47: i. ily
footnote 4. 1966.)

I hope that from this rather detailed account of the typification of
Sorbus it will be evident why I disagree with the opinions expressed by
my esteemed colleague, Dr. Stearn, and why I think that it is essential,
in the interests of nomenclatural stability, to verify immediately and to
establish clearly the typification of al/ generic names, especially those for
which more than one lectotype species has been chosen.

INTRAGENERIC CLASSIFICATION OF SORBUS

There has been much discrepancy in the citation of the names of the
original authors of the subgeneric and sectional names in Sorbus, and,
consequently, many unnecessary new combinations in these categories
have been made. It seems expedient, therefore, to present as an addendum
the intrageneric classification of Sorbus with the correct authorship of the
subgeneric and sectional names. Only the most important synonyms are
included.

Sorbus Linnaeus, Sp. Pl. 1: 477. 1753; Gen. Pl. ed. 5. 213. 1754. LECTOTYPE

SPECIES: S. aucuparia L.; see A. Rehder, Bibliogr. Cult. Trees Shrubs 252.

1949

SUBGENERA

Subg. Sorbus [Sorbus subg. Sorbus Pers. Synop. Pl. 2: 38. 1806, emend. Du-
chartre in D’Orbigny, Dict. Univ. Hist. Nat. 11: 685. 1848.] (Aucuparia
Medic. Gesch. Bot, 86. 1793, nom. illegit. Sorbus subg. Aucuparia Kovanda,
Preslia 33: 14. 1961; Dill, Ber. Bayer. Bot. Ges. 34: 23. 1961.

Subg. Cormus (Spach) Duchartre in D’Orbigny, Dict. Univ. Hist. Nat. 11: wee
1849. (Cormus Spach, Hist. Nat. Vég. 2: 96. 1834. Sorbus Medic. Gesch.
Bot. 87. 1793, non L, Gen. Pl. ed. 5. 213. 1754. Sorbus subg. Sorbus Kovanda,
Preslia 33: 14. 1961; Diill, Ber. Bayer. Bot. Ges. 34: 15. 1961.) TYPE SPECIES:

. domestica L.
rigs Sones. Synop. Pl. 2: 38. 1806. hia SPECIES: S, aria (L.) Crantz;
cf. Int. Code Bot. Nomencl. Art. 22. 1966.

Subg. Be Ma (Reichenb.) Koch, Hort. Dendrol. 178. 1853. (Pyrus L. =
Torminaria (DC.) Reichenb. Consp. Reg. Veg. 1: 168. 1828.) TYPE SPECIES:
S. torminalis (L.) Crantz. ;

Subg. peacnamenl ie (Reichenb.) Koch, Hort. Dendrol. 176. ae : nate
subg. Chamaemespilus (Lindley) Reichenb. Consp. Reg. Veg. 1: : :
Type species: S. chamaemespilus (L.) Crantz.

SECTIONS
Sect. Sorbus [Sect. Eusorbus Boiss. Fl. Orient. 2: 657. 1872.]

508 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Sect. Cormus (Spach) Boiss. Fl. Orient. 2: 657. 1872. (Cormus Spach, Hist.
Nat. Vég. 2: 96. 1834.) Type species: S. domestica L.

Sect. Aria (DC.) Dumortier, Florula Belg. 93. 1827. (Pyrus L. sect. Aria DC.
Prodr, 2: 635. 1825.) Type species: S. aria (L.) Crantz; cf. Int. Code Bot.
Nomencl. Art. 22. 1966.

Sect. Torminaria (DC.) Dumortier, Florula Belg. 93. 1827. (Pyrus L. sect.
r eid DC. Prodr. 2: 636. 1825.) Type species: §S. torminalis (L.)
Crantz.

Sect. Poiidasecailis (Lindley) Schauer, Uebers. Arbeit. Verander. Schles. Ges.
Vaterl. Kult. 1847: 295. 1848. (Pyrus L. sect. Chamaemespilus Lindley, Trans.
Linn. Soc. London 13: 98. 1822.) Type species: S. chamaemespilus (L.)
Crantz.

1968] BHANDARI & VENKATARAMAN, DRIMYS WINTERI 509

EMBRYOLOGY OF DRIMYS WINTERI

N. N. BHANDARI AND REVATHI VENKATARAMAN

DrIMyYs Is ONE OF SIX genera of the vesselless family, Winteraceae. The
genus includes nearly 40 species of which 36 occur in the Old World and
the rest are found in the Americas. The family is of interest because of
the presence of numerous primitive characters such as the undifferentiated
stamens in some members; a complete series from an open conduplicate
to a closed carpel which may be sessile or stipitate; stigmatic crest; lami-
nar placentation, and primitive xylem without vessels.

Bailey (1944), Bailey and Nast (1943a, b; 1944a, b; 1945), and Nast
(1944) have dealt extensively with the morphology and vegetative ana-
tomy of various members of the Winteraceae. However, literature on its
embryology is rather meager. Maheshwari (1950), while discussing the
relationships of angiosperms with other groups, has remarked “Regarding
the relationship of the angiosperms with other groups, we are at present
entirely in the dark, It is possible that a study of morphology and em-
bryology of Degeneriaceae, Winteraceae, Trochodendraceae etc., may
throw some light on the problem.’”’ So a study of the embryology of the
Winteraceae was taken up (see Bhandari, 1963) and the present investi-
gation is a continuation of that project.

PREVIOUS WORK

Strasburger (1905) made some preliminary observations on the ovule
and embryo sac of Drimys winteri var. winteri and reported normal em-
bryo sac and nuclear endosperm. Bhagawathi Kutti Amma (1938)
studied the details of microsporogenesis in Drimys and recorded amoe-
boid anther tapetum. On the other hand, Swamy (1952) observed secre-
tory tapetum, normal embryo sac, and cellular endosperm in Zygogynum
baillonii. In Pseudowintera colorata (Bhandari, 1963) the tapetum is
amoeboid but in P. axillaris (Sampson, 1963) it is secretory. In both
species the embryo sac is of the Polygonum type, and endosperm is cellu-
lar. Post-fertilization stages in the ovules and pericarp have been fol-
lowed only in P. colorata (Bhandari, 1963).

MATERIAL AND METHODS

The buds, flowers, and fruits of Drimys winteri were obtained by the
late Professor P. Maheshwari through the courtesy of Dr. V. Garcia of
Argentina. The collections were fixed on the spot in FAA ‘ay nat
stored in 70 percent ethanol. The material was passed through alcohol-

Fic. 1. Drimys winteri, microsporangium. A, I, transection of anther show-
ing archesporium; B, J, transection of anther (J is an enlargement of a portion
of B showing some archesporial cells in division, sporogenous tissues, and a
parietal layer); C, K, cross section of anther at the stage of organization of
wall layers (wall comprises an epidermis, two parietal layers, and central

Sporogenous tissue; C is diagrammatic); D, a diagrammatic sketch for L; L,
representation of a locule enlarged to show a dividing cell of inner parietal layer
which contributes to the tapetum and middle layer II; E, M, transection of
anther, one locule showing the epidermis, endothecium, 2 middle layers, tapetum,
and the sporogenous tissue; N, the same, note that the tapetum is well devel-

512 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Fic. 2. Drimys abet
microspore wathen

‘ cel
form polyploid masses in M and N; P, a degenerating tapetal cell; Q, a tetra-
hedral tetrad showing microspores with exine and i , the intine muc

g thr
spores at different stages of division; V, a decussate tetrad wh 2- eaied pa
grains; W, polar view of a pollen grain showing the pattern of the exine, germ

1968] BHANDARI & VENKATARAMAN, DRIMYS WINTERI 513

xylene series and later embedded in paraffin, The seeds (mature seeds
without seed coat) were run through the above or through the tertiary
butyl alcohol series. Prior to sectioning, the embedded material (es-
pecially the seeds) was soaked in water or in 30 percent glycerine at
room temperature for two weeks to facilitate microtomy. The sections
were cut 6 to 14 microns thick and the preparations were stained with
Safranin-fast green, but those of seed development stained better with
hematoxylin-eosin combination. Hand sections of mature seeds mounted
in 15 percent glycerine were also examined to study the seed structure.

OBSERVATIONS

Microsporangium. The young anther is histologically undifferentiat-
ed with a well defined epidermis enclosing a homogeneous mass of cells.
As the anther enlarges, in each of the four corners, a group of four or
five hypodermal archesporial cells differentiates (Fics. 1A, I). The hypo-
dermal cells divide periclinally to form the primary parietal layer and
the sporogenous layer, while the subhypodermal ones directly fovm the
sporogenous tissue (Fics. 1B, J). They divide mitotically to add to the
sporogenous tissue (Fic. IJ). The primary parietal layer undergoes
a periclinal division to form two layers (Fics. 1C, D, K, L). The outer
forms the endothecium and middle layer I, while the inner gives rise
to the tapetum and middle layer II (Fics. 1E, M, N).

The epidermal cells become tangentially compressed and highly vacuo-
lated; their nuclei start degenerating but they stay as a thin layer of
collapsed cells at the time of anther dehiscence (Fics. 1G, H, P, Q). During
meiosis, the encothecial cells enlarge radially, develop strong fibrous
thickenings (Fics. 1F-H, O-Q), and are persistent in the mature anther
(Fics. 1H, Q). A few cells of the connective also develop such eee
(Fic. 1Q). The middle layers remain healthy until the beginning o
Meiosis but soon after their cells become flattened. Consequently, the
tapetum becomes detached and lies as an isolated layer inside the anther
locule, surrounding the dividing microspore mother cells (Fics. IF a
The tapetal cells show high metabolic activity, become binucleate ( rg
2K, L) and their nuclei fuse and form large polyploid masses (Fics. v3 a
O). Soon the walls separating the adjacent tapetal cells become 8 He
(Fics. 10, 2N), and after the formation of microspore —— the ce :
show signs of degeneration in situ (Fic. 2P). At a time when the exine 1
differentiating the tapetum gets absorbed completely.

Microsporogenesis and Male Gametophyte. Meiosis of ie 3k
mother cells is non-synchronous. During heterotypic division (Fics.

pore, and the dumbbell-shaped protrusion of intine:; sige oo tae of roa
larged to show the protruding intine and the collar ed pol ‘ rains in a tetra-
protrusion due to thickening of the intine; Y, two-celle wey at xX 780: W, X
hedral tetrad; Z, pollen tetrad at the shedding stage. A-V, a Teil Mal

r
both x 935; Y and Z, X 780

514 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

. Drimys winteri, megasporogenesis and female gametophyte (ant,
6, male nucleus). A, longisection of ovular pri-

Fic, 3
antipodals; e, epidermis;
mordium showing a single-celled archesporium and primordia of two integu-
ments; B, longisection of nucellus showing megaspore mother cell and two
parietal cells; C, the same, megaspore mother cell and a 3-layered parietal tis-

sue formed by the parietal cell; D, the same, megaspore mother cell at meta-
phase; E, longisection of nucellus, two dyad cells at metaphase II, the parietal

1968] BHANDARI & VENKATARAMAN, DRIMYS WINTERI 515

E) an evanescent cell plate is sometimes formed at Telophase I (Fic. 2E).
At the end of homotypic division (Fics. 2F—J) simultaneous quadriparti-
tioning occurs by furrowing. The microspores remain in permanent tetrads
which may be tetrahedral or decussate (Fics. 2Q, U, V). At the beginning
of gametogenesis, the nucleus of the microspore moves towards the proxi-
mal pole of the tetrad where it divides (Fics. 2R-T ). The four microspores
in a tetrad may show different stages in the formation of the generative
cell (Fic. 2U). The wall between the two cells degenerates so that the
generative cell, surrounded by a hyaline cytoplasmic sheath, comes to lie
free within the cytoplasm of the vegetative cell (Fic, 2Z).

The pollen grains are monoporate and the pore is developed towards the
distal end. The intine protrudes through the pore and shows maximum
thickening in this region (Fics. 2Q-Z). As stated by Bailey and Nast
(1943a) for Drimys section WINTERA, here also a collar is present at the
base of protruding intine (Fics. 2W, X). The pollen grains are shed in
tetrads at the 2-celled stage (Fic. 2Z).

Megasporangium, Megasporogenesis and Female Gametophyte.
The young carpel resembles a conduplicately folded leaf. The two approx-
imated margins of the carpel are firmly concrescent except at the stigmatic
region where the epidermal cells interlock. There are 15 or 16 anatropous,
bitegmic and crassinucellate ovules. A single hypodermal archesporial
cell differentiates in the nucellus (Fic. 3A) and divides transversely to
form a primary sporogenous cell and a parietal cell. The latter undergoes
repeated periclinal and anticlinal divisions to form a 2- or 3-layered parietal
tissue (Fics. 3B—E). Meiosis I in the megaspore mother cell (Fics. 3C,
D) results in a dyad, of which the micropylar cell is smaller (Fics. SE, F).
Both of the dyad cells divide simultaneously to form a linear or a T-
shaped tetrad (Fics. 3E, F). The chalazal megaspore functions and the
other three megaspores degenerate (Fic. 3G). Three successive mitoses
result in a 2- (Fic. 3H) 4- (Fic. 31) and 8-nucleate embryo sac (Fics. 3J-
L). Thus, the development conforms to the Polygonum type. Occasional
instances of twin embryo sacs were also observed (Fic. 3L).

Fertilization and Endosperm. Syngamy and triple fusion have been
observed (Fic. 4A). The pollen tube is sometimes persistent during early
stages of the endosperm development. The endosperm is ab initio pas
The primary endosperm nucleus migrates towards the center of the em i
sac before its division (Fics. 4B, C) which is followed by the formation o
a transverse wall (Fic. 4D). The next division in the two unequal cham-

, two dyad cells showing telophase spindles ind
T-shaped tetrad; G, longisection of nucellus showing
e oth

ipodal cells; K, micropylar end ¢
hooked synergids; L, twin
x 61

516 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Fic Drimys winteri, endosperm id sala ee cells; cc, chalazal chamber;
ds, degenerated ible rgid; hap endos , male nu ucleus: mc, micropylar
chamber; pemb, proembry en eae ta caoue ae nucleus; pt, pollen tube;
sn, second ry Pt te , embryo on sage triple aye Bae
the persistent remnants of pollen B, the same, sh
oe nucleus, zygote, neon of degenerated synergid oe aotipoda eels;

ryo sac with the primary e repent uch two
ieee. and four-celled ue stages, the chided chaiber his divided into

two cells of which the lower cell is a Reese G, H, longisections of mas-

1968] BHANDARI & VENKATARAMAN, DRIMYS WINTERI 517

bers is transverse but the division in the chalazal chamber precedes that of
the micropylar (Fic. 4E). The two cells formed in the micropylar cham-
ber are again unequal, the smaller cell being towards the chalazal end
(Fic. 4F). It appears that the chalazal chamber does not contribute much
to the formation of the endosperm proper.

The endosperm soon becomes massive and at the 10-celled stage of the
embryo it is made up of cells having prominent nuclei and dense cytoplasm.
Later, the endosperm cells become packed with small round or oval brown-
ish particles of unknown nature which did not give any positive test for
starch, oil or proteins (Fics. 4G, I). In mature endosperm the cells are
large with little cytoplasm but are packed with bodies which stain darkly
with hematoxylin (Fics. 4H, J).

Embryogeny. The earlier stages observed were a zygote (Fic. 5A)
and a nearly 20-celled proembryo which is rather irregularly organized
(Fic. 5B). Perhaps the divisions do not correspond to any of the known
types of embryogeny. From the globular stage onwards the embryo devel-
opment is regular (Fics. 5C—G) with a prominent suspensor and well de-
marcated protoderm. During late embryogeny periclinal divisions in the
6th or 7th subprotodermal layer begin the demarcation of the embryonal
pith, procambium, and cortex in a globular proembryo (Fics. 5H, 1).
The suspensor cells are filled with starch grains. In the oldest seed that
we examined the embryo was at a pre-heart-shaped stage (Fics. 5H-I).
No more details could be obtained due to paucity of material.

Seed coat. At the megaspore mother cell stage the outer and inner
integuments are 3-layered for the greater part of their length (Fic. 6A).
During megasporogenesis brownish substances accumulate in some cells
of both the integuments. Just after fertilization the cells of the outer
integument and those of the inner epidermis of the inner integument :
the micropylar region become packed with substances which stain an
red with safranin. Subsequently, the space between the outer and the
inner integument becomes almost obliterated (Fic. 6 : :

In a mature seed the testa is formed by the outer integument. The cells
of the outer epidermis elongate radially, become thick walled, a
hard, and packed with brownish contents ( Fics. 6C-E). sa . . oh
two layers of cells filled with tannin and the inner epidermis, whic - na
thin-walled with small nuclei and large vacuoles (Fic. 6C). The inner
integument becomes almost crushed or is i aS by one or two
layers of compressed thin-walled parenchymatous ;

ae mee feature is that in the mature seed the outer surface of

sive endosperm at globular and post-globular stages of the oe 0 ti ba
I, part I in G enlarged to show a few cells of the pe aa) ‘a aa |
cytoplasmic with prominent nuclei and filled with oval or glo u = P highiy
portion J in H enlarged to show a few endosperm Seog oF ea AC.
vacuolate and contain dark squarish particles of reserve foo : ;
X< 550; D-F, x 440; G, H, X 30; I, J, X 360

518 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

SS)

OLE Cee

y) at N
URE ASS (9
e Sel Ba Io/ox9)
“QUES

a) On Y
EE)

Fic. 5. Drimys winteri, embryogeny. A, zygote; B, an irregularly organized
nearly 20-celled embryo; C-G, stages in the development of a globular embryo
with a conspicuous suspensor; H, I, an undifferentiated heart-shaped embryo
contained in a seed obtained from a mature fruit. A, B, x 600; C-G, 325;

the seed coat is thrown into three or four ridges. The endosperm, however,
is not ruminate (Fics. 6D, E).

Pericarp. At the megaspore mother cell stage the ovary wall consists
of ten to twelve layers of parenchymatous cells (Fic. 6F). Two or three

1968] BHANDARI & VENKATARAMAN, DRIMYS WINTERI 519

inner and outer layers are filled with brownish phenolic compounds, Num-
erous ethereal oil cells having a vacuolated cytoplasm and prominent
nuclei are interspersed within the pericarp (Fic. 6F). During post-fertil-
ization stages the wall becomes 20- to 25-layered. The number of ethereal
oil cells and tannin cells also increases (Fic. 6G). When the embryo is
at the globular or post-globular stage the pericarp becomes nearly 30-
layered and the outer and inner epidermis are greatly flattened. The num-
ber of tannin cells increases and they become completely filled with such
substances. The parenchymatous cells become greatly stretched and break
down (Fic. 6H). No meristematic activity of the pericarp as has been
observed in Pseudowintera (Bhandari, 1963) is seen here.

DISCUSSION

Embryological observations on the Winteraceae are rather meager and
are confined to Drimys, Pseudowintera, and Zygogynum. These are dis-
cussed in the light of the present work on Drimys winteri.

The anther tapetum is glandular in Pseudowintera axillaris (Sampson,
1963) and Zygogynum (Swamy, 1952) and amoeboid in P. colorata
(Bhandari, 1963). Bhagavathi Kutti Amma’s (1938) report of amoeboid
tapetum in Drimys is doubtful since the present investigation has clearly
established its glandular nature. In Zygogynum the nuclei in the tapetal
cells divide and fuse to become polyploid but in Pseudowintera they re-
main binucleate. In Drimys, however, the tapetum becomes irregularly
bi-layered with binucleate cells which often become polyploid.

In Zygogynum baillonii (Swamy, 1952) a centripetal furrow is formed
after Meiosis I. Rarely an incipient equatorial cell-plate is also laid down

(Swamy, 1952) and Pseudowintera colorata (Bhandari, yal: cg eted

of the microspore nucleus in a tetrad are simultaneous. In rs nies eal

axillaris (Sampson, 1963) and Drimys winteri (present repor

spores in the same tetrad show different stages o en enera. The
The embryo sac is of the Polygonum type in all se tt in Zygogy-

synergids and antipodal cells degenerate soon after rine - aon

num (Swamy, 1952) but in Pseudowintera colorata r aap ape oF ie

Drimys winteri (present work) they persist for a yd interi_ (present

endosperm. A twin embryo sac has been observed in 2.

work),

520 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Fic. 6. Drimys winteri, seed coat a pericarp og gets oil cells; end,
endosperm; ii, inner integument; ie, inner epidermis: nu, nucellus: oi,
integument; per, pericarp; tc, tannin cally. A, lisence ian OE ovule at mega-

1968] BHANDARI & VENKATARAMAN, DRIMYS WINTERI 521

Triple fusion precedes syngamy in Pseudowintera colorata (Bhandari,
1963) and Drimys winteri (present work). In Zygogynum_ baillonii

be transverse or longitudinal in P. colorata (Bhandari, 1963). In: B:
winteri, Strasburger (1905) reported nuclear endosperm. A reinvestigation
(present observations) of this species, however, has shown this to be er-
roneous. The endosperm, therefore, is ab initio cellular in all the genera
so far investigated.

The present investigation is the only record of embryogeny indicating
that the earlier divisions of the developing embryo are irregular. How-
ever, from the globular stage onwards the development is regular. In
Pseudowisitera colorata (Bhandari, 1963) the embryo is dicotyledonous
but in Drimys winteri it remains undifferentiated.

In Pseudowintera colorata (Bhandari, 1963) and Drimys (present
work) the seed coat is formed by elongated tannin-filled cells of the outer
epidermis, along with two or three hypodermal layers of thick-walled cells
of the outer integument. In Drimys, however, ae work) the outer
epidermis of the outer integument is variously ri

The pericarp is composed of 20 to 25 layers ot parenchymatous cells
with interspersed ethereal oil cells in both Pseudowintera colorata (Bhan-
dari, 1963) and Drimys (present work). In P. colorata the unilocular

meristematic activity of pericarp between the ovules. No such meristematic
activity has been observed in any other winteraceous member

Systematic position. Bentham and Hooker (1862-67) included
Drimys together with //licium in the tribe Wintereae of the Magnoliaceae.

pers ae cell stare to show the two integuments which are 3- or 4- layered;
t the

C, a magnified part of the seed coat at the preglobular stage of the nya
the outer integument has elongated cells completely filled with tannin, the inner-
most layer consists of large parenchymatou Se the inner peta us ing
as a thin layer of compressed parenc vinataie ells; D, transection of mature
seed to show the ridges on the hard seed coat ( asd se ection) these ridges are
restricted only to the outer epidermis of the outer integument; E, a part of D

agnified to show the details of the seed coat; F, tra ice of a carpel at the

are compressed, between these are 20 to 25 layers of piale <asurea cells
with numerous tannin cells and empty iced oil ing A, x aes, Bx 100:
C, x 110: D, 100: E, < 200; F, x 435; G, X 140; H, xX 3

522 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Van Tieghem (1900) grouped all the members with vesselless xylem under
Homoxylées. He placed five winteraceous genera (Drimys, Bubbia, Bellio-
lum, Exospermum and Zygogynum) in this group whereas Dandy (1933)
later added the sixth genus, Pseudowintera, from New Zealand. Barkley
(1966) has recognized three more genera, namely, Tetrathalamus, Wintera,
and Lassonia in this family but includes Belliolum with Drimys. Hutchin-
son (1959), too, considers Tetrathalamus a distinct genus of the Winter-
aceae although Smith (1943) merges this genus with Bubbia as B.
montana, Embryological information on Tetrathalamus, Lassonia (= Mag-
nolia, see Willis, 1966) and Wéintera is wanting and any consideration
regarding their taxonomic position must await such data

Hutchinson (1959) also believes that Degeneria is related to Exosperm-
um and Zygogynum and should, therefore, be included in the Winteraceae.
Bhandari (1963) has compared the embryological features of the Winter-
aceae and Degeneria and supported the conclusions of Bailey and Smith
(1942) that Degeneria be assigned to a separate family, the Degeneriaceae.
However, no light has yet been thrown, embryologically, on the relation-
ship of Winteraceae with the Illiciaceae. The morphological and embry-
ological characters of these families are compared in the following table:

Table 1

WINTERACEAE

ILLICIACEAE

Hasit
*WoopD ANATOMY

*NODAL ANATOMY

FLOWERS

*CALYX

*COROLLA

*ANTHER

*ENDOTHECIUM

Trees and shrubs.

arate heterogeneous ray
ctu racheary pitting
perenne

Trilacunar.

Actinomorphic to zygomorphic,

bisexual or unisexual, hypo-
gynous.

Calyptrate; sepals bract-like,
onnate.

Petaloid to scale-like, reduction
from many to 2

Microsporangia subterminal,
or lateral, protuberant or
embedded.

Endothecium oO towards
the connective regio

* Points of difference.

Shrubs and trees.
Vessels present,
multiseriate ray struc-
ture. Tracheary pitting
scalariform to opposite.

Unilacunar, single-
traced

Actinomorphic, bisexual,
h us.

No differentiation into
calyx and corolla;
tepals numerous.

Microsporangia lateral,
protuberant.

Endothecium does not
extend towards con-
nective region.

1968] BHANDARI & VENKATARAMAN, DRIMYS WINTERI 523

Table 1.- (Continued)

*TAPETUM Amoeboid or secretory, cells Glandular, irregularly
binucleate, parietal in origin. 2-layered, cells binuc-
leate, originates from
sporogenous tissue.
TETRADS pacers tetragonal, or Tetrahedral.
decu
*POLLEN In permanent tetrads, mono- Individual, tricolpate,
GRAINS porate, generative cell cut off isopolar.
at proximal end, intine comes
out of the germ pore
*CARPEL Open or closed, en Closed, sessile.
sessile or stipitat
OvULE ai de cras- Anatropous, bitegmic,
sinucellat crassinucellate
FEMALE ir et type, antipodals Polygonum type, anti-
GAMETOPHYTE degenerate after fertilization. podals ephemeral
ENDOSPERM Cellular. Cellular or nuclear?
*EMBRYOGENY Irregular. Asterad type.

* Points of difference.

It is clear from the above Table that /ilicium differs from Drimys and
allied genera in numerous embryological and morphological features which
negate its relationship with the Winteraceae and support its inclusion in
a separate family, the Illiciaceae (cf. Eames, 1961; Hayashi, 1960, 1963;
Kapil & Bhandari, 1964; Smith, 1947). However, at the same time it has
certain features which indicate its ranalian affinities.

We are grateful to the late Professor P. Maheshwari F.R.S. for suggest-
ing the problem and procuring the material for us, to Professor B. M.
Johri for his keen interest, and to Dr. A. C. Smith for reading the manu-
script.

LITERATURE CITED

Battey, I. W. The comparative morphology of the Winteraceae, III. Wood.
Jour. Arnold Arb. 25: 97-103. 1944

Cc Nast. The comparative morphology of the Winteraceae, I.
Pollen and stamens. Jour. Arnold Arb. 24: 340-346. 1943a.
The comparative morphology of the Winteraceae, II. Car-
pels. Ibid. 472-481. 1943b.
& The comparative morphology of the Winteraceae, IV.
a eal of the node “a vascularization of the leaf. Jbid. 25: 215-221.
1944a.

a comparative morphology of the Winteraceae, V. Foliar
epidermis and eaten Ibid. 342-348. 1944b.

524 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

& ————. The comparative morphology of the Winteraceae, VII. Sum-
mary and conclusions. /bid. 26: 37-47. 1945
A. C. SMITH. oe a new family of flowering plants from
Fiji. Jour. Arnold Arb. 23: 356-365. 1942.
Bark ey, F. A. A list of the orders and families of Anthophyta (Angiosper-
mae) with generic examples. Hopkins Press, Providence. 1966.
Bentuam, G., & J. D. Hooker. Genera Plantarum. Vol. I. London. 1862-
1867.

BuacAvatHt Kuttr Amma, P. R. Microsporogenesis in Drimys. Presidency
College Bot. Mag. Madras 5: 22-25. 1938.

Buanpart, N. N. Embryology of Pseudowintera colorata — A vesselless dicoty-
ledon. Phytomorphology 13: 303-316. 1963.

Danpy, J. E. The Winteraceae of New Zealand. Jour. Bot. Lond. 71: 119-122.
1933.

Eames, A. J. Morphology of the Angiosperms. New York. 196

se ¢. ae Ass division by furrowing in Magnolia. A a ‘Bot. 5: 379-

aun = ae the gies and pollen morphology in the family
Magnoliaceae. Sci. Rep. Tohoku Univ. (Biol.) 27: 45- = 1960.

The embryology of the Magnoliaceae sens. lat. I. Megasporogenesis,

female gametophyte and embryology of Jllicium anisatum. Ibid. 29: 27-33.

963.

1
Hutcuinson, J. The families of flowering plants. Vol. I. Dicotyledons. Lon-
don, 1959

Karin, R. N., ‘& N. N. BHANDARI. Morphology and embryology of Magnolia
Dill. ex Linn. Proc. Natl. Inst. India. 30B: 245-262. 1964.
ee P. An introduction to the embryology of angiosperms. New
ork
Nast, C. G. The comparative morphology of the Winteraceae, VI. Vascular
anatomy of the flowering shoot. Jour. Arnold Arb. 25: 454-466. 1944.
Sampson, F. B. The floral morphology of Pseudowintera, the New Zealand
mber of the vesselless Winteraceae. Phytomorphology 13: 403-422.

1963.

SmituH, A. C. The American species of Drimys. Jour. Arnold Arb, 24: 119-164.
1943.

———. The families Illiciaceae and Schisandraceae. Sargentia 7: 1-224. 1947.

STRASBURGER, E. Die Samenanlagen von Drimys winteri und die Endosperm-
bildung bei Angiospermen. Flora [Jena] 95: 215-231. 1905.

Swamy, B. G. L. Some aspects in the embryology of Zygogynum baillonii V.
Tiegh. Proc. Natl. Inst. India 18B: 399-406. 1952.

TIEGHEM, P. vAN. Sur les dicotylédones du groupe des Homoxylées. Jour.
Bot. Paris 14: 259-297, 330-361. 1900.

WILLIs, a C. A dictionary of the flowering plants and ferns. Ed. 7, revised by

K. Airy Shaw. Cambridge. 1966.

DEPARTMENT OF BOTANY

UNIVERSITY OF DELHI
DELHI 7, INDIA

1968 | THE DIRECTOR’S REPORT 525

THE DIRECTOR’S REPORT

THE ARNOLD ARBORETUM DuRING THE FISCAL YEAR ENDED
JUNE 30, 1968

N UNUSUAL early and temperate spring season produced a spectacular
flowering display, and the living collections were particularly beautiful,
drawing larger numbers of visitors than have been seen in the experience
of the current staff. Spring, 1968, can be considered one of those rare and
enjoyable spring seasons in New England which permit the public to
enjoy the grounds to their fullest and the staff to make observations
through long periods of flowering. The lilacs, for example, were at their
best for two weeks from May 12 to 25, while last year the short and
belated display period occurred between June 3 and 10. In contrast was
the unexplained severity of winter damage to many plants in the nursery
areas. Severe losses of young plants occurred in January when a normally
adequate snow cover was on the ground. The same pattern was also seen
in commercial nurseries and on private grounds throughout the north-
eastern part of the United States, with the most severe losses of plant
materials occurring in areas on Cape Co

An item of major construction completed during the year was the new
garage and storage building for mechanical equipment used on the grounds.
Ever since the construction of the herbarium addition to the Administra-
tion Building in 1905, mechanical equipment and tools had been kept in
the basement, with access through a narrow ramp. Although in earlier
days the grass in the Arboretum was permitted to mature and was, in fact,
cut by scythe and stacked in mounds, in recent years mowing has been a
continuous operation through the growing season, and the cut grass has
been retained as a mulch. The tractor-drawn mowers and similar power
devices to cut grass or to care for the living collections fitted like pieces
of a jig-saw puzzle in the basement. With funds accumulated from the
bequest of Martha Dana Mercer a new garage was planned and con-
structed and was officially accepted on March 1, 1968. Built into a bank
area, the one-story structure, 44 by 105 feet, is of reinforced concrete
faced with brick. Adequate storage at ground level for all vehicles and
grounds equipment is supplemented with special storage rooms for parts,
tools, fertilizers, and sprays. Four electrically operated overhead metal
doors permit easier access. The $145,000 budget for this building in-
cluded a grease pit, overhead lift, 1000-gallon gasoline storage tank, and
additional essential equipment, such as a hydraulic press, steam cleaner,
air compressor, electric welder, and electric grinder. A paved parking
area for twenty-one staff cars removes them from the front of the admin-
istration building and permits easier parking for visitors. The whole

526 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

The newly peas? garage and vehicle-servicing facility ates adja-
cent to the Administration Building. Climbing plants have been planted along
the end walls of pee building and near the re and these ‘eal 3 soon et th
lines of the additio

garage area is enclosed by a chain-link fence topped by barbed wire.
Nineteen different kinds of climbing plants have been started near the
garage and along the fence to beautify the area.

taff :
oak this year the Arboretum staff lost by death one of its valuable
m Dr. Georg

botanist on the Generic Flora of the Southeastern United States. His
careful bibliographic and taxonomic work is exemplified in more than 80
scientific papers. His unusual career will ee reviewed in a biographical
sketch planned for a later issue of the Jou

Two staff members retired at the end of ‘e year: Mrs. Margaret C.
Lefavour, as herbarium secretary, and Mr. Henry Draper, as superintend-
ent of the Case Estates in Weston. New appointees will take their places
but it will be difficult to equal their loyal contributions to the development
of the Mehercinie: during the years they have served on the staff

Resignations were received from Mr. Andrey pangs Herbarium
Assist ee who will do botanical consulting, an Harrison Flint
Associate Horticulturist, who will join the faculty fe tie Department of
Horticulture of Purdue University.

New appointments to the staff included Dr. Paul Davidsen Sorensen,

1968 | THE DIRECTOR’S REPORT 527

as Assistant Horticultural Taxonomist, and Mr. George Howard Pride as
Associate Horticulturist. Dr. Beryl Simpson Vuilleumier was appointed as
Botanist on the Flora of the Southeastern United States project. Mr.
Thomas Matthew Kinahan, of the Arboretum grounds staff, was promoted
to be Superintendent of the Case Estates.

The members of the staff serve the broad fields of systematic botany
and horticulture in many ways, both nationally and internationally. These
are often through various committees, and new assignments accepted dur-
ing the year include the appointment of Dr. DeWolf to an advisory panel
of the National Science Foundation; Mr. Fordham as chairman of the

and
Wyman to the Board of Directors of the Roxbury-Dorchester Ben tiiee
tion Project.

Dr. Howard was elected an honorary foreign member of the Royal
Danish Academy of Science and Letters and the Honorary President-at-
large of the Herb Society of America. He also received the distinguished
citizen citation for 1968 issued by the Metropolitan Area Association of
Warren, Ohio.

Horticulture:
The activities of the staff in the field of horticulture involve the main-
tenance and care of the living collections of woody plants; work in plant

The several pani t terraces for genetic dwarf conifers below the slat house
holding the bonsai collection. The Dana greenhouses appear in the background.

528 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

propagation; and the operation of nursery areas and display plantings
established at the Case Estates in Weston.

In Jamaica Plain, work continued on the old road bed along Centre
Street recently acquired from the City of Boston. The road surfacing
material was removed and replaced with soil and mulching materials.
Grass was sown and some coniferous species planted. This area, although
small, will permit an expansion of the pinetum and the development of
attractive plantings of crab apples visible from the heavily travelled U:S.
Route 1.

With construction of the new garage adjacent to the administration
building, new entrances were constructed to the basement and to the first
floor of the herbarium, the latter in association with a loading platform
which will facilitate delivery of supplies. The ugly ramp to the basement
was replaced with a bulkhead. The lawn in front of the building, which
was ruined by the installation of water pipes and by the construction
equipment, was completely renovated; it will soon look better than
before.

The collection of genetic dwarf conifers was enlarged by the addition
of 177 taxa planted on two additional levels of the terraces below the
bonsai house. Around the nursery, the collection of ground cover plants
especially suited for bank planting was increased by 24 taxa. During the
fall and spring 375 taxa were added to the general collections of living
plants.

Shortages of water during past years have had accompanying city re-
strictions on the outdoor use of water, even in our nursery areas. To
counter this problem a well 115 feet deep was drilled this spring, with
the pleasing result of a supply of 50 gallons per minute. This quantity
will be sufficient to supply five sprinklers in the greenhouse area through
the use of an electric pump and will be adequate for greenhouse and
nursery needs during periods of future restrictions on city water. A new
overhead irrigation sprinkler system was also installed by the staff for the
nursery area. Included in the system is an automatic timing device per-
mitting a sequence of watering in various areas and an intake system so
liquid fertilizers can be applied through the water flow.

The work of the greenhouse staff includes a consideration of needed
additicns to our collections through seeds and the propagation of plants
on the grounds needing replacements. During the year 401 taxa were
propagated to prepare replacements for specimens which appeared to be
failing or which were insufficiently represented within our collections. One
hundred fifty-five shipments of plant materials, comprising 649 taxa,
were received from the United States and nine other countries. Seeds of
339 taxa either for germination and addition to the collections. or for
experimental studies in germination, or for staff research were included in
86 shipments from the United States and 25 other countries. Requests
for materials, seeds, or propagating material from our collection were filled
when possible. Although material which can be procured from commercial
sources is rarely distributed in response to nonprofessional requests, legiti-

THE DIRECTOR’S REPORT 5

1968]

me

=
oy

ae

we

er
Rix
_—

ie

i

eS
ta

if Ga

3

oe a none eet eR lee
ween eee “te ~

i a
~

5
S

solve some watering problem

enhouses struck water at less

A flow of 50 gallons per minute will

The well drilling operation adjacent to the gre

than 100 feet.

in the nursery area.

530 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

mate requests for documented material are honored, and during the year
214 shipments of plant materials comprising 1146 taxa were sent to co-
éperating scientific institutions or nurseries or for individual research
projects in 17 countries.

The program of distribution in a plant hardiness test begun last year
by Dr. Flint was expanded this year with the codperation of Mr. Fordham
and the plant propagation staff. Twenty-five codperating stations from
western Massachusetts to Presque Isle, Maine, and areas in northern New
Hampshire and Vermont were selected, and more than 1500 container-
grown woody plants of 41 taxa were distributed. Uniform methods of
culture have been suggested for these plants, as well as systematic ob-
servations which can be correlated to increase our knowledge of hardiness
of species and of the nature of winter damage.

Surplus plants from the Arboretum nursery areas were given as usual
to the Department of Buildings and Grounds at Harvard and to neighbor-
ing institutions in adjoining states. These materials correctly identified
and properly labelled and documented increase the value of campus plant-
ings as teaching aids to departments of botany and horticulture. We have
also codperated with the City of Boston in programs for beautification
of depressed areas. Two truck loads of plants were given to the Roxbury-
Dorchester Beautification Project, and over one thousand rhizomes of
surplus Iris were made available for distribution.

Activity in the registration of new cultivars of woody plants continued
during the year. A registration list of cultivars in Lantana is nearing
completion, and progress is being made on one for Philadelphus. An issue
of Arnoldia was devoted to the publication of new names submitted for
registration. From our own collections Albizia julibrissin cultivar ERNEST
WILSON, introduced from Korea by E. H. Wilson in 1918 and now con-
sidered to be sufficiently distinctive to merit formal recognition, was
described.

The staff is also codperating with a national committee in re-evaluating
the species and cultivars of lilacs grown in the United States. After a
three-year study the committee plans to reissue its analysis of “Lilacs for
America,” which was first published in 1953.

A mail survey of American colleges was conducted to determine the
number of campus floras and the frequency of their documentation. Local
lists prepared by taxonomists and properly documented by herbarium
specimens can be used for information on the distribution and hardiness
of cultivated taxa with greater reliability than can data obtained from
nursery catalogues.

The staff is also cooperating with the American Horticultural Society
in the establishment of a Plant Records Center. Dr. Howard serves as
chairman of a committee which will coordinate the gathering of informa-
tion on the holdings of various botanical gardens and arboreta. An ulti-
mate goal is a data processing system permitting quick recovery of in-
formation on the source and location of important plant materials. No
up-to-date system for the location of particular plants needed for scien-

1968]

THE DIRECTOR’S REPORT 531

rug ete libris
ardiness in the
ae st oon the more southern

‘Ernest Wilson’, a new cultivar status for an
Nev

vw England climate and the spreading habit distinguish
plants.

532 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

tific studies exists today. A grant to the American Horticultural Society
from the Longwood Foundation will initiate this work. Administrative
procedures are being worked out at the present time, with compilation
of information to begin as soon as possible.

A program of collecting herbarium specimens from plants in the Arbo-
retum collections which are the sources of type material is under way.
Although such specimens exist in our own herbarium, the increased interest
on the part of others in vouchers and the possibility of obtaining equally
valuable material from other institutions makes this a project of particular
value, When the size of the plant permits, twenty specimens are collected
for future exchanges.

The research program of Dr. Owen Rogers, of the University of New
Hampshire, as a Mercer Fellow during the past year is of special interest.
His work during the spring involved controlled crosses of many taxa of
Syringa using plants in the living collections and those from his plots in
New Hampshire. Time is required for an analysis or evaluation of the
seeds obtained and seedlings developed. It is of interest to record that,
although many seeds were obtained from crosses of Syringa velutina and
S. meyeri, only albino seedlings developed, a result similar to that obtained
earlier by Karl Sax from crosses involving Syringa pubescens.

Case Estates:

The Case Estates, comprising 110 acres of land in the town of Weston,
serve both as the nursery area for the Arboretum and for display plantings
open to the public. The land also provides room for specimens which
cannot be accommodated in the plantings in Jamaica Plain.

A generous gift of 575 Rhododendron seedlings and over 100 rooted
cuttings of cultivars of Jlex was made by Mr. Josiah K. Lilly, III, of
Sandwich, the owner of the Dexter Estate. These plants will be grown
to larger size, tested for hardiness, and evaluated at the Case Estates.

Through the efforts of Mr. George Pride and members of the New
England Wild Flower Society a collection of wild flowers also is being
developed. Since the natural areas of the Case Estates are used by the
public schools of Weston and are visited by many interested organizations,
a small collection of wild flowers will enhance the educational value of this
area.

The extensive collection of crab apples planted on the Newton Street
tract flowered profusely this year, drawing the attention of commuters
and visitors. An additional roadside planting on Wellesley street of 32
varieties of Rhododendron will add to the attractiveness of this land in
the center of Weston.

Herbarium:

The problem of adequate space continues in the herbarium of spontane-
ous plants housed in Cambridge. The University is codperating in a study
of means of obtaining increased space either by the addition of a fifth
floor to the present building or by the addition of a wing. Studies of space

1968 ] THE DIRECTOR’S REPORT 533

needs for laboratories, staff offices, and herbarium storage, and methods
of financing construction are ahead.

A total of 31,750 specimens was mounted during the year and added
to the herbarium, bringing the holdings to 881,730, of which 132,750
specimens are housed in Jamaica Plain and are from plants in cultivation.

These additions are from collections on hand from past years, and only
in small part from those received during the current one. During 1967—
1968, 13,234 specimens were received in exchange, 562 as gifts, 4,886 from
the subsidy of expeditions, 4,118 from staff collections and 1,363 in ex-
change for identifications. These ranked in order of magnitude from
Papuasia, the West Indies, Africa, Mexico and Central America, Western
Malaysia, and the United States, with lesser numbers from the rest of
the world. Our holdings from Africa are inadequate for reference and
teaching purposes, and we value the codperation of colleagues in Berlin
and South Africa who have supplied selected African material. A total of
8,867 specimens was sent as exchange, with 112 additional ones sent for
identification or as special gifts.

The combined herbaria of the Arnold Arboretum and the Gray Herbari-
um sent 18,050 specimens on loan for scientific studies. There were 168
loans to 84 institutions, 54 in the United States and 30 in 16 other
countries, For our staff and students we borrowed 13,139 specimens in
80 loans from 35 institutions, 29 in the New World and 5 in the Old.

Herbarium specimens are borrowed for monographic studies or to aid
floristic work. These represent the primary scientific studies of the sys-
tematic botanists on the Arboretum staff which are possible only with the
use of a major herbarium and its supporting library. The resident staff
and some of the Mercer Fellows have the advantage of these important
collections in their work. Our materials are also consulted by numerous
visiting scholars. Unfortunately, working space for these scientists is
minimal, and additional work space and mamittg area for specimens and
books must be provided in the very near future

Although the published bibliography iaidicutes the completed scientific
contribution much significant work is in progress. The Southeastern
Flora project under the supervision of Dr. Carroll Wood is financed by
a grant from the National Science Foundation. Dr. Beryl Vuilleumier and
the late Dr. George Brizicky have codperated with him in the studies of
plant families in this flora. Visiting collaborators appointed as Mercer
Research Fellows during the past year have included Dr. John Thieret,
of the University of Southwestern Louisiana, working on the Scrophulari-
aceae and Orobanchaceae; Dr. Robert Long, of the University of South
Florida, the Acanthaceae; and Sister Mary Victoria Hayden, of Catherine
Spalding College, genera of the Rubiaceae. An important paper on the
genera of the Euphorbiaceae by Dr. Grady L. Webster, University of
California, Davis, was published in the Journal of the Arnold Arboretum.
Reflecting Dr. Webster’s long-standing interest in this complex family,
this account deals with many problems far beyond the geographic borders
of the Generic Flora. Ten additional families are currently under study,

534 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

by various botanists. Mention should also be made of the work of the
artists associated with this project. Mrs. Sydney B. DeVore, who re-
placed Miss Rachel A, Wheeler during the year, is continuing the critical
illustrations of genera. Each illustration is based on living or recently
preserved material and represents an original study. Many genera are
being illustrated for the first time. Errors in fact and interpretation which
have been long perpetuated are revealed and corrected in the course of the
art work. Detailed illustrations have been prepared for 205 genera of
105 families.

The staff wishes to express a special word of appreciation for the work
of Dr. George Brizicky who died during the year. His fluency in Slavic
languages was a particular asset to the project, for he was most generous
with his time and assistance. At the time of his death Dr. Brizicky was
working on the genera of the Malvaceae, and he had completed a manu-
script on the general problem of subgeneric and sectional names, their
starting points and early sources, subjects which had interested him for
a number of years.

Dr. Thomas Hartley has continued his work in the identification of
personal collections of plants made in Papua and New Guinea between
1961 and 1965. The 3700 numbers represent one of the largest recent
collections from the area, and they are specially significant in supporting
phytochemical studies. The final study will be a check list published in
collaboration with chemists of the Commonwealth Scientific and Industrial
Research Organization in Melbourne, Australia, documenting the chemical
analyses which came from field and laboratory studies. Dr. Hartley’s
monographic studies in the family Rutaceae are continuing with an in-
vestigation of the genus Flindersia,

Dr. Howard completed the field studies of an elfin forest on the top
of Pico del Oeste, in the Luquillo Mountains of eastern Puerto Rico.
This project, supported by a grant from the National Science Foundation,
was an ecological study of a peculiar environment and forest type. The
collaboration of many scientists from other institutions made possible
studies on the weather, soils, and root and anatomical characteristics of the
vascular plants, as well as on the non-vascular plants. The study docu-
ments the unusually high and persistent humidity associated with a nearly
continuous cloud cover and places greater significance on the air-borne
moisture of clouds than on the prevailing winds. Studies of composition
of the forest compared the frequency of species with the occupancy of the
canopy surface and revealed that species infrequent in number in the total
flora dominated the canopy. The unusual habits of constituent plants
included the development of prop roots, as well as branched and un-
branched aérial roots, the high frequency of long-shoot-short-shoot growth
patterns, and a candelabra branching system, all of which contribute to
the contorted stem structure and low growth aspect of the elfin forest.
Since liverworts proved to be more abundant than mosses, the commonly
used term ‘mossy forest” is a misnomer for this area. Algal studies by a
collaborator showed an unexpectedly high number of species, not only on

1968 | THE DIRECTOR’S REPORT 535

the ground, but on the trunks of the trees, in the gelatinous sheaths of
adventitious roots, and in the liquid held by epiphytic bromeliads.

Chemical studies showed a low percentage of plants with alkaloids and
related chemicals but an unexpectedly high number of plants containing
chemicals showing biological activity in relation to animal tumors. The
results of the study will be published in subsequent issues of this Journal.

Dr. Hu spent four months in the territory of Hong Kong developing
her studies of the flora of that area. Her field work has greatly increased
the collections of specimens including plants under cultivation in Hong
Kong in our herbarium. The courtesies extended to her by the staffs of
Chung Chi College and the Herbarium of the Urban Council and Urban
Services Department, Hong Kong, are greatly appreciated.

Dr. Nevling, representing the staffs of both the Arnold Arboretum and
Gray Herbarium, is participating actively in investigations on the vegeta-
tion of the state of Veracruz with Dr. Arturo Gémez-Pompa and members
of the Instituto de Biologia, Universidad Nacional Aut6énoma de México.
Two field trips during the year permitted quick surveys and selected col-
lecting of native and cultivated plants. The staff of the Instituto de Bio-
logia is continuing collecting efforts subsidized in part by the Arboretum.
Dr. Nevling is assisting in the identification of specimens and in research
necessarily based at Harvard’s more complete libraries. This study con-
tinues the traditional interest of Harvard’s botanists in the flora of Mexi-
co. Dr. Nevling’s monographic interests continue in the family Thyme-
laeaceae.

Dr. DeWolf and Mr. David Hall, a Mercer Fellow, are collaborating on
a study of the genus Ficus in Venezuela for a national Flora of that
country being prepared by the staff of the Instituto Botanico in Caracas.
A similar treatment of Ficus in Suriname prepared by Dr. DeWolf will be
published by the State University of Utrecht, Netherlands. Detailed
studies of the misunderstood Ficus dicranostyla, a species of western and
central Africa, have shown that this belongs to a small group of species
the nearest member of which is restricted to a small area of northern
Madagascar. This is an unusual distribution pattern for species of Ficus,
but is another example of the phytogeographic patterns previously re-
ported for other genera.

Dr. Schubert has worked towards completing floristic treatments for
Desmodium and Dioscorea. Requests for accounts of Desmodium in the
Galapagos islands, Texas, Panama, and tropical East Africa are all as-
sociated with floras in preparation at other institutions. Her field work
in Mexico has permitted a special investigation of several small-statured
species of Dioscorea which have not been adequately represented by
herbarium material.

Dr. Sorensen has continued his broadly based study on the genus
Dahlia and is currently tracing the introduction of species into cultiva-
tion and their subsequent selection and breeding. Cytological work on
plants grown at the Arnold Arboretum has yielded original chromosome
counts for 16 species and hybrids. Coéperative phytochemical and ana-

536 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

tomical studies of Dahlia materials are being undertaken by scientists

historical studies of plant introductions to and from the West Indies.
With the collaboration of Mrs. Brigid Sturrock, resident in St. Lucia, a
renewed study of the plants of that island is under way. Studies of the
phenology of plants from St. Lucia can be correlated with ecological
studies based in Puerto Rico and St. Kitts.

Each of these staff projects demonstrates the unique contribution
possible from a major herbarium and library of world wide concern.

Library:

The problem of available space for growth of collections referred to
in connection with the herbarium is also of concern within the library.
Studies for an addition to the Harvard University Herbaria building in
Cambridge will involve the nature of growth and the increased use of
the library collections. Currently, 90 per cent of the available shelf
space is filled, which is beyond the recommended maximum of 70 to 85
per cent, the remaining unoccupied space being needed to avoid unneces-
sary shifting of books.

The library of the Arnold Arboretum grew by the addition of 491 vol-
umes and 219 pamphlets and reprints to totals of 54,267 and 21,098, re-
spectively, or a total holding of bound books and periodicals and un-
bound reprints and pamphlets of 75,365.

The regularly issued additions of the Gray Herbarium Index, the Index
Nominum Genericorum, and the Torrey Index were filed. Thirty-six
hundred sixty new catalog cards were added during the year, and the
Rehder Index of important information on cultivated plants maintained
in Jamaica Plain received an additional 625 cards. Until the end of last
year reports of our holdings as duplicates of catalogue cards were sent
to the National Union Catalog at the Library of Congress and to Widen-
er’s Union Catalog at Harvard. Now an additional record will be sent to
the new undergraduate science library at Harvard, which will also hold
general card indices

Miss Sutton continues her work on the biography of Charles Sargent
and on the cermin of manuscripts and letters of historical interest at
the Arnold Arboretum

During the year an inventory was made of the publications of the
Arboretum, including the accumulations of reprints of articles by former
staff members. The reprints were arranged in sets, and a list of these
and available publications was prepared and distributed to libraries and
members of the American Society of Plant Taxonomists. The collections
of reprints offered at cost were quickly sold and the number of publica-
tions on hand significantly reduced, not only making the volumes avail-
able to interested persons, but freeing useful space.

Comparative Morphology:
The wood collection received normal curatorial care during the year.

1968 | THE DIRECTOR’S REPORT oa7

Additional specimens of wood and slides were received, including sets
of prepared slides associated with Project 1, School of Forest Resources of
the North Carolina State University, a study of the woods of the United
States. Requests for wood samples or for the loan of slides were handled
when possible. Active use of the collections involved the work of Dr. Wil-
liam Theobald and Mrs. Helen Roca-Garcia in investigations on the woods,
stems, and leaves of plants of the dwarfed forest on Pico del Oeste,
Puerto Rico, while Dr. Alfred Linn Bogle continued his studies of the
Hamamelidaceae and allied families.

Education:

No formal courses were taught in the Department of Biology by mem-
bers of the Arboretum staff during the year, but active instruction to
graduate students and participation in seminar programs continued as
usual. Dr. Schubert conducted the seminar program in systematic botany
during the fall term. Dr. Hartley again offered the Harvard University
Extension Commission course in general botany.

Field courses were offered to Friends of the Arboretum during the fall
by Wyman and by Mr. Pride, in Jamaica Plain and Weston, re-
spectively. Mr. Fordham offered a discussion of methods of collecting
and treating seeds of woody plants. During the winter a series of lectures
on Caribbean botany was given at the Case Estates by Dr. Howard. The
regular spring class in ornamental plants was offered by Dr. Wyman in
Jamaica Plain, and Dr. Owen Rogers taught a class on hybridization and
breeding of ornamental plants. Mr. Pride presented a course entitled
“Beginning Botany for Gardeners” at Weston during the sp

Staff members are frequently requested to speak to various ss esags
tions on the work of the Arnold Arboretum or on subjects of their research
interests. A new brochure facilitates the handling of these requests. In
addition, the staff participates in seminars and similar programs of other
scientific organizations. Dr. Harrison, Mr. Fordham, and Mr. Williams
all participated in the lecture discussions sponsored by the Massachusetts
Horticultural Society during the annual spring flower show. Mr. Fordham
also talked on the Members’ Monday programs of that organization as well
as at the Regional Meeting of the American Rhododendron Society, the
Connecticut Nurserymen’s Short Course, and the annual meeting of the
International Plant Propagators Society. Dr. Flint spoke at the same
meeting of the Plant Propagators Society and at the annual meeting of
the Vermont Plantsmen’s Association. Mr. Pride was on the winter pro-
gram of the Worchester County Horticultural Society.

Dr. Howard was sponsored by the American Institute of Biological
Sciences at the University of Maine for two talks. He appeared on the
regular programs of the Pennsylvania Horticultural Society, the Barnes
Arboretum, Connecticut Nurserymen’s Short Course at the University of

Clark University. He was the banquet speaker at the Clara B. Ford

538 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Garden Forum in Michigan and the annual meeting of the American
Association of Botanical Gardens and Arboreta at Hamilton, Ontario.

Dr. Wood participated in a two-day Science Seminar as part of the
125th anniversary celebration of Roanoke College in Salem, Virginia. He
represented the field of biology on a panel including two chemists, a
physicist and a mathematician. His talk on the subject of ‘Biology or
Oblivion” commented on the population crisis and man’s destruction of
his environment.

Dr. Wyman spoke at meetings of the Garden Club Federation of Maine
and the Massachusetts Garden Club Federation. He took part in the
Longwood Gardens Symposium held jointly with the University of Dela-
ware. He was on the program of the New York Botanical Garden and
of the University of Georgia short course for nurserymen.

Again this year the Arboretum staff cooperated with the science teaching
program in the elementary schools in Weston. The students were en-
couraged to study the plants and plantings of the Case Estates in Weston
and to prepare competitive reports. The participants were divided into
two groups: first prizes were awarded to Kenneth Leghorn and John
Algird, with second and third prizes also cash awards; as honorable men-
tion recognition, etchings were presented to each group. The prize papers
concerned descriptions of the grounds, the plants or the activities there,
an historical study of the Case family and the property, a special study
of crab apples, one of the bristle-cone pine, and a report including a tape
recording of the behavior of a family of grackles. The interest of the
students and their parents resulted in larger attendance on the grounds
aay perhaps greater awareness and appreciation of the activities of the
staff.

cate, the student’s selection of books of a stated value, and a gift from
the Arboretum of a tree or shrub from the available duplicate plants within
our nurseries. The first award was made to George B. Ransom, a senior
in the Agricultural Department at the Jamaica Plain High School.

The Arboretum donated 10 units of plant material for the annual
auction held by Channel 2, WGBH, Boston’s educational television

mentators who offered the plants to the wide New England audience did
ts

The staff codperated with the Worcester County Horticultural Society
and the Weston Garden Club in preparing and presenting two displays at
the fall and spring flower shows of each organization. A display of mulch
materials and berried or fruiting plants was prepared for Worcester, and

1968 | e HE DIRECTOR’S REPORT 539

an exhibit of methods of plant propagation and a collection of edible and
poisonous wild plants of New England were offered at the Weston shows.

Travel and Exploration:

Members of the staff traveled less than usual during the year, although
one trip was of extended duration. Dr. Hu spent four months in Hong
Kong and offered lectures in plant taxonomy at Chung Chi College. She
collected where possible in the territory from the coastal mangrove area
at Tai Po Kau to the mountain vegetation at Ma An Shan. Dr. Nevling
and Dr. Schubert made independent trips to Mexico, the former in asso-
ciation with the American Institute of Biological Sciences meetings in
Texas and in cooperation with the University of Mexico. Mr. Pride visited
Morocco, obtaining photographs of the vegetation, as well as seeds and
specimens, and continued his trip to Teneriffe, Madeira, and ‘“‘Les Cédres”
at Cap Ferrat, France. He visited gardens and established contacts for
the Arboretum in each area. We are grateful for the many kindnesses and
the hospitality extended to him during this trip by so many people, par-
ticularly Dr. Andres Garcia Cabezon, Director, Jardin de Aclimatacion
de la Orotava; Mr. and Mrs. P. Graham Blandy; and Mr. and Mrs. Julien
Marnier-Lapostolle.

The field study of the dwarfed mountain top forest in Puerto Rico en-
abled many staff members to take part for short periods of time and to
learn of the tropical vegetation in the process or continue their own studies
of taxa represented in that flora. Misses Powell and Sutton and Messrs.
DeWolf, Nevling, Wyman, and Howard participated. Drs. Howard and
Nevling visited a comparable site on the island of St. Kitts, where studies
were made with the permission of the governor, Sir Fred Philipps, and the
coéperation of Mr. Campbell Evelyn. En route from St. Kitts to Puerto
Rico, they spent a short time on St. Thomas for observations on the dry-
land vegetation.

Dr. Flint visited research stations in Minnesota, North Dakota, and
Manitoba in connection with his studies of plant hardiness.

The Arboretum has also been able to support the work of other bota-
nists collecting in areas of particular interest to the staff or of value to
our collections. The field work of Dr. L. Bernardi, of the Conservatoire
Botanique, Geneva, Switzerland, extended from Madagascar through Ma-
laysia and to New Caledonia where he is making botanical collections to
be shared with us. More recently, the threat of the complete destruction
of the vegetation on the inadequately known island of Anegada in the
West Indies was called to our attention. A special gift permitted support
of collecting in this area with the specimens to be identified and distributed
by the Arboretum staff. As a comparable example, Dr. Wood collected
material of Drosera in a unique bog on the Gaspé Peninsula in August,
1967. The bog has since been destroyed but the plants now growing in the
Arboretum greenhouses represent at least three species, one of which ap-
pears to have originated by hybridization of the other two followed by
chromosomal doubling. There are also back-cross hybrids with both

540 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

parental species. Regrettably the botanist cannot act with sufficient speed
and interest to stay ahead of the world wide destruction of essentially
scientifically unknown plant material.

Gifts and Grants:

The many friends of the Arboretum renewed their support during the
year with most gifts being without restrictions for use in the general work
of the Arboretum. We were pleased to accept memorial gifts in honor of
Mrs. John S. Ames and Mrs. Alice Hoagland and several gifts specified for
the care of the bonsai and/or the genetic dwarf conifers.

Many persons have been generous in donating special plant materials,
and we particularly value the collection of Rosa species and ecotypic ma-
terial of Rosa wichuraiana from Mr. Seizo Suzuki and of Rhododendron
and //ex selections from Mr, Josiah K. Lilly, IIT.

A display case in the entrance hall of the Arnold Arboretum holds a
collection of indian artifacts gathered from the grounds of the Arnold
Arboretum by the late Ernest J. Palmer and by Alfred Fordham. Mr.
Palmer bequeathed the collection to Mr. Fordham who has now officially
given the display to the Arboretum. We are grateful for these evidences
of former use of the land occupied by our collections.

Through the kindness of Dr. George T. Jones, of Oberlin College, the
Arboretum received residual materials from the estate of Franklin P.
Metcalfe, formerly of our staff. These included field books and numerical
listings of his and other collections from China, as well as a smaller num-
ber of books of unusual historical interest.

Publications:

Dr. Bernice Schubert continued to edit the Journal of the Arnold
Arboretum during the past year when four numbers, totalling 675 pages
and including 20 articles, were issued. Dr. Harrison Flint served as editor
of Arnoldia, which comprised 9 issues, totalling 91 pages.

The early volumes of the Journal of the Arnold Arboretum have long
been out of print, as have several numbers of more recent dates. The
Kraus Reprint Company, of New York, has been authorized to reproduce
back issues and now volumes 1 to 20 of the Journal of the Arnold Arbo-
retum are available in the reprint edition and volumes 21—40 are being
prepared for reprinting.

Mercer Fellows:

The appointment of Mercer Fellows to the staff of the Arnold Arbore-
tum was begun in 1961 following Mrs. Martha Dana Mercer’s bequest
to the Arboretum. Individuals are selected from candidates who request
consideration for support of their program of work or study of the
Arboretum collections or with the staff. As the awards have been made
in the past no academic credit is given, but Fellows may audit Harvard
classes with permission of the particular instructor and have full access
to libraries, collections, and receive other benefits. Recipients have been

1968 | THE DIRECTOR’S REPORT 541

at all academic levels, from those without college background to post-
doctoral scholars and those on sabbatical leaves from their home institu-
tions. Mercer Fellowships have been awarded to scholars from Argentina,
Belgium, England, Finland, Jamaica, Mexico, the Philippines, South
Africa, and the United States.

The largest number of short term appointments to date was made during
1967-1968. Recipients of Mercer Fellowships for all or part of the past
year and their fields of interest are indicated below.

ARTHUR CHARLES Gipson, Miami University (Systematic botany).

MAXINE GLICKMAN, Stockbridge School, University of Massachusetts (Horti-
culture).

Davip W. Hatt, University of Florida (Systematic botany).

StisteER Mary Victoria Haypen, Catherine Spalding College (Systematic
botany).

Syep MoHaMMED ANWaARD KazmtI, Peshawar University, Pakistan (Systematic

otany ).
Rosert WILLIAM Lonc, University of South Florida (Systematic botany and

oristics).

Joun O’Connor, Wolverhampton, Technical Teachers College, England (Plant
propagation ).

FREDERICK E. Roperts, University of Connecticut (Horticulture).

OWEN ee Rocers, University of New Hampshire (Cytology and cyto-
gene

ARLON ee ce Michigan State University (Horticulture).

Joun Wru1am Tureret, University of Southwestern Louisiana (Systematic
otany ).

Bibliography of the Published Writings of the Staff and Students
July 1, 1967 — June 30, 1968

BarANov, ANpREY I. On the taxonomic status of varieties of Sorbaria sorbifolia.
Advanc. Front. Pl. Sci. 19: 5-16. 1

Brizicky, GEorGE K. AHerissantia, Bogenhardia, and Gayoides (Malvaceae).
Jour. Arnold Arb. 49: 278, 279. 1968.

DeWotr, Gorpon P., Jr. Ficus, subgenus Pharmacosycea, section Pharmacosy-
cea. Hooker’s Te: Pi. V; 7(2): 1-22. tab. 3634-3636. 1967.

. Identification of unknown plants. Arnoldia 27: 51, 52. 19
. Albizia julibrissin and its cultivar ‘Ernest Wilson’. aes 28: 29-35.
68

. Notes on Corydalis bulbosa. Rhodora 70: 296-298. 1968.

———. The Red Maple. Gard. Chron. 163(23): 3, 4. 1968.

Firnt, Harrison L. September bloom. Arnoldia 27: 69, 70. 1967.

———. Country cousins. Arnoldia 28: 1-7. 1968.

. Some occa meee activities of Justin one Morrill. Arnoldia 28: 41-

ForpHam, ALFRED J. Seed dispersal by birds and animals in the Arnold Arbo-
retum, Arnoldia 27: 73-84. 1967.

. Woody plants difficult to root. Int. Pl. Prop. Soc. Comb, Proc. 16:

190-193. 1967.

542 JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

. Dwarf conifers derived from witches’-brooms. Plants & Gardens 23:

22-25. 1968.
———. Propagation of Albizia julibrissin. Arnoldia 28: 36-40. 1968.
Hartley, THomas A revision of the genus Lunasia (Rutaceae). Jour.

Arnold Arb. 48: 460-475. 1967.
Howarp, RicHarp A. The Director’s Report. The Arnold Arboretum during
the fiscal year ended June 30, 1967. Jour. Arnold Arb. 48: 546-567. 1967.
. The horticultural herbarium at the Arnold Arboretum. Arb. Bot. Gard.
Bull. 1: 22-24. 1967.
. Polygonaceae. Jn: J. A. Steyermark, Flora del Auyan-tepui. Acta Bot.
Venez. 2: 208. 1967.
. Problems in nomenclature and registration. Proc. XVII Int. Hort. Congr.
2: 239-244. 1967.
. Irving Widmer Bailey, 1884-1967. Jour. Arnold Arb. 49: 1-16. 1968.
. The image and role of the modern botanical garden. Arb. Bot. Gard.
Bull. 2: 22-29. 1968.
———. Money matters at the Arnold Arboretum. Arb. Bot. Gard. Bull. 2:
14-16. 1968.
& PowELL, Dutcre A. Clerodendrum philippinum Schauer replaces
“Clerodendrum fragrans.” Taxon 17: 53-55. 1968.
, BarcHoorn, E. S., MANGELsporr, P. C., Rotiins, R. C., WeTMoRE,
R. H. Irving Widmer Bailey. Harvard Univ. Gazette 63: 196 [March 23].
19

8.
Hu, Surtu-Yinc. The Compositae of China (V). Quart. Jour. Taiwan Mus.
20: 1-77. 1967.
. The economic botany of Dragon tongue. Econ. Bot. 21: 288-292. 1967.
. The evolution and distribution of the species of Aquifoliaceae in the
Pacific area, I-II. Jour. Jap. Bot. 42: 13-27, 49-58. 1967.
. Araceae — Studies in the Flora of Thailand 41. Dansk Bot. Ark. 23:
411-457. 1968.
- the genus Barclaya (Nymphaeaceae) — Studies in the Flora of Thai-
land 48. Ibid. 535-540. 1968.
———. An early history of daylily. Am. Hort. Mag. 47: 51-71, 81-85. 1968.
. The species of Hemerocallis. Ibid. 86-111. 1968.
—————.. Uses of daylily as food and in medicine. Jbid. 214-218, 1968.
NEVLING, Lorin I., Jr. Comprehensive collecting and taxa of uncertain position
—a means of conservation. Atas do Simpésio sébre a Biota Amazonica 7:
195-199. 1967.
. Thymelaeaceae. Jn: Steyermark & Maguire, Botany of the Chimanta
Massif — II. Mem. N.Y. Bot. Gard. 17: 452-455. 1967
& GOmeEz-Pompa, Arturo. A new Hydrangea from Mexico. Jour. Arnold
Arb. 49: 225-232. 1968.
PowELL, Dutcte A. (with Howarp, RICHARD A.). Clerodendrum philippinum
Schauer replaces “Clerodendrum fragrans.” Taxon 17: 53-55. 1968.
SANDERS, Mary E., & FRANZKE, C, J. Recovery of a parent type from colchicine-
treated F, sorghum seedlings. Jour. Heredity 58: 291-295. 1967.
YMAN, DonaLp. Add a Viburnum to your garden. Horticulture 45: 38-39,
41. 1967.
. The best of the dogwoods. Am. Hort. Mag. 46: 165-174. 1967.
. The best trees and shrubs for the colorful fruits of fall. Home Garden
hey 2d. VOGT:
. Evergreen groundcovers, Horticulture 45: 14-17, 31. 1967.

wi
>

1968 | THE DIRECTOR’S REPORT 543

. Ground covers. Gardens on Parade. 2: 4, 11. 1967.
. Hardiness zones. Plants & Gardens. 23: 31-33. 1967.

. The introduction of plants from Europe to America. Jbid. 23: 12-16.
67.

———. More plant registrations. Arnoldia 27: 61-66. 1967.
. The new dwarf conifer collection. /bid. 28: 9-27. 1968.

RICHARD A. Howarp, Director

The Board of Overseers of Harvard College
Committee to Visit the Arnold Arboretum

AucusTIN H. Parker, Jr., Chairman, Boston, Massachusetts.

GEORGE R. CLarK, Vice-Chairman, Philadelphia, Pennsylvania.

Mrs. GeorcE L. BATCHELDER, JR., Beverly, Massachusetts.

Mrs, Paut C. Casot, Needham, Massachusetts.

Henry F. Du Pont, Winterthur, Delaware.

WILLIAM FLEMeR, III, Princeton, New Jersey.

Henry B. Hosmer, Boston, Massachusetts.

Mitrorp R. LAwreENcE, Falmouth, Massachusetts.

Mrs. Joun E. Lockwoop, Bedford, New York.

Harorp E. Moore, Jr., Ithaca, New York.

R. Henry Norwes, Jr., Mentor, Ohio.

Mrs. RIcHARD W. Pratt, Chestnut Hill, Massachusetts.

Francis W. SARGENT, Boston, Massachusetts.

Mrs. Homer N. SWEET, Weston, Massachusetts.

GEORGE TAYLOR, Kew, England.

Joun J. TrERNEY, JR., Commissioner, City of Boston, Department of Parks and
Recreation.

Mrs. CuHar.es D. WessTER, New York, New York.

RicHarp P, Wuite, Washington, D.C.

NATHANIEL WHITTIER, Medfield, Massachusetts.

Mrs. Joun G. WitttaMs, Gladwyne, Pennsylvania.

544 | JOURNAL OF THE ARNOLD ARBORETUM [voL. 49

Staff of the Arnold Arboretum
1967-1968

RicHarp ALDEN Howarp, Ph.D., Arnold Professor of Botany, Professor of
Dendrology, and Director.

Kart Sax, S.D., Professor of Botany, Emeritus.

GrorcE KonsTANTIN Brizicky, R.N.Dr., Botanist, Southeastern Flora Project. +
MicHaEL ANTHONY CANoso, M.S., Senior Curatorial Assistant. *
GorRDON PARKER DEWOLr?, Jr., Ph.D., Horticultural Taxonomist.
Harrison LEIGH Fiint, Ph.D., Associate Horticulturist.
ALFRED JAMES ForDHAM, Propagator.
WitttaM Ep Grime, B.A., Curatorial Assistant. *
THomAs GorDON HartTLey, Ph.D., Associate Curator of Pacific Botany.
HemMAN ArTHUR Howarp, Assistant Horticulturist.
SuHr1vu-Yinec Hv, Ph.D., Botanist.
THoMAS MATTHEW KINAHAN, Superintendent, Case Estates.
MARGARET CATHERINE LEFAVOUR, Herbarium Secretary. **
VicTOR FERENC Marx, M.Libr., Librarian. *
Lorin he SENG, Jr., Ph.D., Associate Curator and Supervisor of the
Herbar

DULCIE ree PowELL, M.A., Botanist.
GrEorGE Howarp Prive, M.A., Associate Horticulturist.
BERNICE GipUz SCHUBERT, Ph.D., Associate Curator and Editor.
PAUL DAVIDSEN SORENSEN, Ph.D., Assistant Horticultural Taxonomist.
STEPHANNE Barry Sutton, A.B., Archivist.
BERYL SIMPSON VUILLEUMIER, Ph.D., Botanist, Southeastern Flora Project.
Rosert GERow WILLIAMS, B.S., Superintendent.
CARROLL Emory Woop, Jr., Ph.D., Associate Curator.
Donatp WvyMAN, Ph.D., Horticulturist.

¥ Died June 15, 1968.

* Appointed jointly with the Gray Herbarium.
** Retired June 30, 1968.

1968]

INDEX

INDEX

ee sect. Gayoides, 279
pum, 278

Seine Flowers, Fruits, and Phy-
logeny

Mie 167-192

sane , 167-192
sect. Ala angium, ee A 171, 174, 175,
179, 183, 185,

— sect. Angolum

— sect. Conostizma, 168-175, 179, 180,

182-186,

— sect. ites 168, 169,
179, 180, 182-186, 188
—sect. Rhytidandra, 168,
175, 179, 182-185, 188

171, 174, 175,

169, 171, 174,

— begonifolium
— chinense, 168, nae 177, 180
— chungii, 167
— griffithii, 180, 182, 186
— grisolleoides, 167, ee 186
vilandii, 168-171
— javanicum, Se 179, 183, 186

175
— rotundifolium, 168, 169, 177, 180
—salviifolium, 167-169, 179, 183, 186,
188
— shweliense, 186
— tonkinense, 167
— villosum, 168, 169, 171
Albizia julibrissin ‘Ernest Wilson’, 530
Alloplectus ambiguus, 400, 408, 412

Dichotomous Vena-
tion of Circaeaster, Further Morphol-
ogical Studies on,

Anatomy of Croomia pauciflora (Stemon-
aceae), Morphology and, 26

Anatomy of the Leaf- bearing Cactaceae,

. Anem

Mee rina etek 63
— herzogii, 6

— phyllitidis, 63

63
Auicnial pre ee 398, 408, 412
bac 399

ae,
Aral ae 452

— vitiensis, 484
Araliaceae of the New Hebrides, Fiji,
a, and Tonga, The. pes nt of
Pacific Island Plants, XIX,
Araliaceae, 431-501
—tribe Anomopanaceae, 431
—tribe Aralieae, 433
— tribe Cussonieae, 431, 477
—tribe Mackinlayea
Ardisia luquillensis, 400, 408, 412, 416

Arthrostylidium sarmeritosith, 398, 408

Asian Species of Delphi mie: age
Stricto, A Synopsis of the, 73,

Aucuparia, 507

Ayensu, E. §., and pias B. ToMLINsON

Morphology and Anatomy - Croomia
pauciflora PETE
Azaleastrum, 2
Azaleodendron, 207

Bailey, 1884-1967, Irving Widmer
mparative pee of
the Leal- bearing Cactaceae, XVII. Pre-
liminary Observations on the Problem
of Transitions from Broad to Terete

Leaves, 3
Bakeria, mS
— vitiensis
Bastardia b. ates 279
nemoralis, gies
BayntTon, Harotp W. The me of =
lfin anarig in Puerto Rico, 2

Microclimate of Pico del Oeste, oe

546

Be onia oo 399, 408

N. N., and RevatuHi baderd
Embryology of Drim
09

—underwoodianum, 397

Boeriagiodendron, 432, 491

— orientale, 491, 4

Bogenhardia, ca Gayoides (Malvaceae),
Herissantia, 278

Bogenhardia, 278, 279

, 400

Botanical Name of the Tea Sageretia of
China, 3

Meranda hao 434

Branch ager Root and. Anatomy of
the P ya cc VI, 307

. Herissantia, Bogen-

ative Anatomy o
the Leaf-bearing Cactaceae, XVII, 370
Bromeliaceae, 39
agri (Sapotaceae) ,
er Species of, 280
pice 280-289
— americana, 282,
see phate To 283
subsp. neglecta, 284
5

Jamaican and

E.
a

—nigra, 283, 284
— octosepala, 283, 284
— repens, 289

— retusa, 282

JOURNAL OF THE ARNOLD ARBORETUM

[voL. 49
Bumelia retusa subsp. neglecta, 284

— — subsp. typica

— rotundifolia, 283, 284

= salicifolia, 283, 285

— sericea,

— stevensonii, 289

Cactaceae, Comparative Anatomy of the
Preliminary Ob-

tions from rere to Terete Leaves, 370
euler, 6, 7, 370-3
oe: 370, 375
ean Opuntioideae, 370, Sly S13,

37
— subfam. oe, 3435 319
Calycogonium mulosum, 399, 408,

409, 412, 41
Calyptranthes krugii, 399, 408, 409, 412,

416

Campanulac

Carex pltachya, 398, 408, 409, 412

Carpels, The. arative Morpholog-
ical Studies in Dilcaictic II, 317

Celastraceae, 399

Chamaerops ritchiana, 16

— ritchieana, 16

Cheirodendron samoense, 480

Chienia, 258
— honanensis, 258, 259

China, Botanical Waine of the Tea Sag-
eretia of, 3

Chloranthaceae, 3

Circaeaster, Pies Morphological Stud-

ies - ee in the Dichotomous

Venation of,
Ciena, 52-72
agre

, 60
Clea alhopanciat, 399, 408
Clibadium erosum, 402
Clusia, 412
— gri ae 399, 408
— gundlachii, 39
Cheha 4 minus, 397
— seminudum, 397
Comparative Anatomy of the Leaf-bear-
ing Cactaceae, XVII. Preliminary Ob-

parative
Dilleniaceae, III. The Carpels, 317
Compositae, 402
Composition Studies, Introduction and.
e Ecology of an Elfin Forest in
Puerto Rico, 1, 381
Consolida, 233, 258, 259

1968 | INDEX 547

pai a 400 Delphinium caucasicum chinense, 134
Cordia — ceratophoroides, 84, 33
—boringuensi oo —ceratophorum, 84, 91, 233
ates 507, — — brevicorniculatum, 237
Cor 186 — — hirsutum
Crmek scutellaria, 4 — cheilanthum —o Haagechstinainne 89
Croomia pauciflora Stomomacn), Mor- -—chenii, 121, 125,
pholgy and Anatomy of, 260 _- chrysotrichum, a8 -
Guns 260, 261, 265, 269, 272-274 —— pygmaeum, 74
— paucifiora, 260-27 7 — chrysotricum, 74
unoniaceae, 399 — ciliatum,
Curatella, ol — —dictyocarpum, 137
— america — coelestinum,

for)
o
=|
Lona)
&
n
c
Cy
_
iS)
Un
os
WwW
Nm

poeo
Cyathea : aon 397
— pubescens, 397, 408, 412
397

|
i]
°
i)
i
S
—y
bo
=
i
Bb

Cyatheaceae, 3 —corymbosum, 126, 133
Cyperaceae, 39 —— baicalense, 13
Cytotaxonomy of Illicium floridanum and —crassicaule, 121, 134

I. parviflorum (lIlliciaceae), 41 —crassifolium, 125, 135

— — tanguti 3

Sagres ee 399 —crispulum, 1
Davilla, —cryophilum, 125, 135
ries ie — cyananthum, 84, 91
Degeneriaceae, 5, 522 — dasyanthum, a
Delarbrea, 432, 433 — dasycarpum, 85, 92, 124
—collina, 433, 434 — dasycaulon malabaricum, 106
Delphinium, Sensu Stricto, A Synopsis of — davidii, 83,

the Asian Species of, 73, 233 — delavayi, 83, i
Delphinium, 73-166, 233-259 ———aureum, 93

— acuminatissimum, 85, 86 minatum, 93
—afghanicum, 124, 126 — psig eee
—albocoeruleum, 121, 122 ubsp. tibeticum, 97
— albomarginatum, 131 ———cinereum,
— alpinum ajanense, 138 —— — glandulosum, 97
—— productum, 13 — pubiflorum,
—altaicum, 86, 87 — discolor, 138
—altissimum, 85, 87 — drepanocentrum, 83, 94
— — subsp. drepanocentrum , 94 — duhmbergii, 125, 126, 132, 136
— — subsp. wallichii nipalensis, 87 —— retropilosum, 160
—altissimum drepanocentrum, 94 — dyctiocarpum, 121, 137

— phallutensis, 87 a ubsp. uralense, 120
— amoenum — — canescens,
—arcuatum, 123, 127 — — glaberrimum,
— atropurpureum, 138 —elatum, 122, 125, 126, 138
— autumnale, — — subsp. intermedi
— boissieri, 162 —-— subsp. palmatifidum, 138

bonvalotii, 23 ——— glabell
—brachycentrum, 86, 89 ——— glabrum, 138
—bracteosum, 123, 127 ——— hispidum, 138

—albidum, 162 — —subsp. subglabrum, 138
— — macranthum, 162 — nomalum, 103
— bulleyanum, 234 — —intermedium hirsutum, 135
— burmaense, 123, 128 — — longicalcaratum, 13
— buschianum, 12 —— pubiflorum, 87
—campylocentrum, 233, 235 —— pyramidatum, 15
— carela, —— ranunculifolium, 158
baie caucasicum, —_—— sericeum, 138

124
— bracteosum, 162 —— subciliatum, 141

548 JOURNAL OF THE ARNOLD ARBORETUM

Delphinium elatum subglabrum, 154
1

a glabellum, 138

|
>
;
=|
2.
2
“5
ie

— incis

— laxicymosum, 82, 102

Delphinium laxiflorum, 82, 103
B alpi 52

104
~ lineartobum ‘hirticauke, 129
— lipskii, 10
Scent 122, 149
— longipes, 9
—J]udlowianum, 105
— ludlowii, 85, 105
sinigaeklanastil. 425,-150

3
a
=
+ @
5
wn
&
iS)

me, 241
— pogo néhum, 83, 84, 113
— wisn 124, 157

pterospermum,
— pulchellum,
— pycnocentroides, 74, 77
— pycnocentrum, 73, 7
——lankongense, 78

[voL. 49

1968] INDEX 549

Delphinium ebiaageaeiah 122, 158 Delphinium tongolense, 233-235, 241
— pyramidatum, 5 OD —trichophorum brevistylum brevutigue,
—ranunculifolium, 158
ee 122, 160 —trifoliolatum, 82, 119
— rha —turkestanicum, 136
enh 123, 160 — ukokense, 123, 164
— saniculifolium, 84 —umbellatum, 133
— sauricum, 121, 161 tia tgs 83, 11
— saxatile, a p. drepanocentrum, 94
— scabriflorum, 85, 114 beac 85, 120
— scaposu — vestitum,
— sertiferum, 112 — villosum, 124, 164
— sherriffii, 83, 115 ——anomalum, 164
~—— slamense, 85, 116 — — laxiflorum, 103
— sino-scapos 13; 79 ——winkleri, 164
— sino-vitifolium, 234, 240 — vitifolium, 240
— souliei, 73, 74, 80 —‘winkterianum, 124, 165
— sparsiflorum, 84, 116 Dichotomous Venation of Circaeaster,
Biccqieeniaed: 1225 59392,2161 Further Morphological Studies on An-
— subsp. iaviineblifolun, 158 astomoses in
aR pt sree 158 Dicxison, Wr11am C. Comparative
— — — eriantha, 158 Morphological acne in Daliataene,
— — — katatricha, 158 III. The Carpels,
— -—— lasiantha, 158 Didesmandra, 3
— —— marriensis, 158 — aspera, 326
— —— oinophylla, 158 Dillenia, 318
—— — pulverulenta, 158 — aurea,
— -— —thyrsantha, 158 — indica, 327
— — bracteosum, 162 — ovata, 327
— — bruehlianum, 158 —papuana, 327
—— dasycarpum, 92 — parvi 3
—— nopus, 16 — philippinensis, 327
—— pyram 1 — suffruticosa,
——ranunculifolium, 158 Dilleniaceae, Comparative a
— — trichocar : Studies in, III. The Carpels,
——typicum biebersteinii, 161 poesia 317-
— spirocentrum, 234, 241 m. Dillenioideae, 325
— — grandibracteolatum, 241 — su a. Tetraceroideae, 325
— — hirsu , 241 Dilomilis montana, 398, 408, 409, 412
— — pauciflorum, 234 Dioscoreaceae,
— — pedifor Dipholis, 282
— stapeliosum, 124, 126, 163 — bellonis, 285
ha , 163 —bullata, 284
—-—shanicum, 128 — cubensis, 285
—-siamense, 116 — durifolia, 287
— stenose , 89 — ferruginea, 287
— sungpanense, 82, 117 — jubilla, 287
— sutchuense, —ma 8
— szechuanicum, 109 —minutiflora, 287
—taipaicum, 74, 8 — montana,
—taliense, 84, 117 — nigra
—— glabrum, 108 —octosepala, 284
— pubi 108 — parvifolia, 287
—ternatum, 8 18 a ns,
—thibeticum, 74, 81, 84 — salicifolia, 285
—— schizophyllum, 77 — sericea 3 *
—_— — subint a 81 —stevensonil, 28
126, 163 Diplazium pahisinetins: 397

550

a ae oe 397, 408
— Vher

— striat 97

Bhecors me See , The. The Arnold Ar-
boretum During the Fiscal Year Ended
June 30, 1968, 525

Diverse Nodal Types in Rhododendron,

Dizygotheca, 432, 490
—elegantissima, 490, 491
3

— dentatus, 326

— guianensis, 326

— major, 327

Drimys winteri, Embryology of, 509

Gentes: 490

Ecclegy of an Elfin Forest in Puerto Rico,
The, 1. Introduction and Composition
Studies, 381; 2. The Microclimate of

Eleocharis yunque nsis
Elfin Forest in Puerto en The Ecology
"Seca and Com Osi-

Embryology
Epidendru
nae nn
_ iflo , 398

Ericaceae, 211, 214, 400

Eschweilera, 491

Eschweileria, 491

— pleiosperma, 461

Eugenia OE ees 399, 408, 412, 416

Eype, Ricnarp H. Flowers, Fruits, and
Phylogeny of Alangiaceae, 16

Ferns, 412
ie Same, and Tonga, The Araliaceae
the eleringornse Te of Pacific
Ts land pias ay, >. Gs
Fimbristylis peters xen
Florida, ea ConTi” a New Genus of
the Liliaceae from,
Flowers, eri and Pipiaces of Alang-
iaceae,
sy fairs S. Further —
Studies on Anastom
Dichotomous Venation of gr stereo

Pune Jupr
STONE

L., and Don E.
ee ae ee Illicium: aan

JOURNAL OF THE ARNOLD ARBORETUM

[voL. 49
danum and I. parviflorum (lIlliciaceae),

Further Morphological Studies on Anas-
tomoses in the Dichotomous Venation
of Circaeaster, 52

Gayoides (Malvaceae), Herissantia, Bog-
enhardia, ~ : : 8

Gayoides, 278,
— crisp

— “Fypiiation, Sorbus and the
Pro

Genus saciacs Een Revision of

he Liliaeceae from Florida,

00
MPA, —— and Lorin
Nevin, on A New Hydrangea from
Mexi
Gonoeiyx ae 400, 409, 412,

iene 398
rammadenia sintenisii, 400, 408
Sandals hessii, 397

7
GREEN, P. S. Revision of the Genus No-
telaea Anew 333
Guttiferae, 399
Guzmania berteroniana, 398, 408
Gymnelaea ligustrina, 362

Haenianthus salicifolius, 408

— — obovatus,

Harperocallis, a New Genus of the Lilia-
ceae from Florida, 35

Harperocallis, 35-40

ava,
Hedyosmum arborescens, 398, 408, 412,
4l

6

He Gor cies samoense, 480

— vitien: 7

ronan Bogenhardia, and Gayoides
(Malvaceae), 278

Herissantia, 278, 279

— cri 279

andra, 327
Hila parasitica, 401, 408
annia racemosa, 400, 409, 412

1968]

Howarp, RicHarp A. The Director’s Re-
port. The Arnold Arboretum During
the Fiscal Year Ended June 30, 1968,
525

Howarp, RicHarp A. The Ecology of an
Elfin Forest in Puerto Rico, 1. Intro-

Widme er

Hydrangea from Mexico, A New, 225
Hydrangea, 225-23
— sect. Cornidia, oe 229, 232

— macrothecum, 397, 412

Icacinaceae,

Ichna nthas: ae 398, 408

Tlex sintenisii, 399, 408, 412, 416

Illiciaceae, 41-51, 522, 523

llicium floridan um and I. evrieriions
(Illiciaceae), Cytotaxonomy of,
Ilicium parviflorum (lIlliciaceae), con
taxonomy of Illicium floridanum and,

mere 41, $21, 523
Badia

ana,
— aad Cymbostemon, 42, 43, 47
—sect. Illicium, 42, 4 51
— floridanum,
arviflorum, 41-5

Anatomy of the Palm
Rhapis excelsa, V, 29
Inflorescence in Nannorrhops ritchiana
(Palmae), 16
Introduction and enorme rua
an

e Ecology of Elfin rest in
Puerto Rico, 1, pad
Ipomoea repanda, 400, 409,
rving Widmer nae cee 1
Irvingbailey

ba
Tsachne angustifolia, 398

Jamaican and Other Species of Bumelia
Sapotaceae), 280
Jounston, MarsHatt C. Botanical Name
of the Tea Sageretia of China, 377
Justicia, 400
—martinsoniana, 400, 408
— verticillaris, 400

Lauraceae, 399

Comparative
reliminary

Observations on the Problem

tions from Broad to Terete Leaves, 370

INDEX

551

Leguminosae, 402

Liliaceae from Florida, Harperocallis, a
New Genus of the, 35

Liliaceae, 35-40
— subfam. Melanthoideae, 36

— tribe phorigapet 35,.36

Liliatae

er ad aHee 397

Linociera brachystachys, 369

Lobelia err Mh ne 408

Lophosoria quadripinnata, 397
Lycopodiaceae, 396

Lycopodiophyta, 396

Lycopodium cernuum, 396

— taxifolium, 396

Magnolia splendens, 399, 408

Magnoliatae, 398-402
ee” 398-402

Marc re

= arn 399, 409, 412, 416

rocallis, a
New Genus of the Liliaceae ois Flori-
da, 35
Mecranium amygdalinum, 399, 408

9

Mexico, A New dae from, 225
Miconia foveolata, 399, age -
—pachyphylla, 400, 408,

— pycnoneura, 400, 408, He

— sintenisii,

gitar - Pico del Oeste, The.
Ecology n Elfin Forest in ge era
Rico, ate

Microph i garciniaef

olia, 400, 408
Mikania pachyphyl 402, 409, 412, 416
Moore, H. E. Jr., r. OMLIN
ron ase in lara ritchiana
(Palmae), 1
Moraceae, eg

552
Morphological Studies in Dilleniaceae,
o

Morphological Studies
in the Dichotomous Venation of Circa-
52

?
Anatomy . Croomia
pauciflora (Stemonaceae), 26
Munz, Putte A. A Synopsis A the Asia
Species of eae Sensu i.
73, 233

Myrsinaceae, 400
Myrtaceae, 399

ame of the Tea Sageretia of China,
Botanical, 37

(Palmae), In-
Nannorrhops, 16-18, 29, 33

—ritchiana, 16-34

leo 35; 36

Nea
Nesopansy, 466

— vitiensis, 471
seis Tustin, 362
VLING, I, 7 . and ArTuRO

New Hydrangea

New Hebrides, Fiji, Samoa, and Tonga,
The Araliaceae of the. Studies of Pa-
cific Island Plants, XTX, 431

New ae goon, from Mexico, A, 225

Nietneria, 35, 3

Nodal Types in Sk etc Diverse,
19

Noronhia emarginata, 369

Notelaea (Oleaceae), Revision of the

7330

—- sect. Ligustrina, 334, 336, 362-366
— sect. Mischopetala, 334, 366-368
— sect. Notelaea, 334, 336-362, 366
— sect. Picconia, 368
— austro-caledonica, 369
— azorica, Me
— badula,
—brachstachys, 369
— collina
Been poe
— eucleoides, 369
—excelsa, 369
— oe 369
— si
— glandulosa,

johnsonii, ae 336, 366-368
ph par 369
— laurifoli
— Heustsina, 336, 362-366

JOURNAL OF THE ARNOLD ARBORETUM

[voL. 49

Notelaea hy one 358-360
a)

—— glabra, 343, 344
— ee 342, 343
—— ovata, 345

— peices 349
a, 340

—— microcarpa, 355-357

cola, 36
Se at. 336, 360-362
— nervosa, 3
— ovata, 335, 340, 344-348
— eae 369

ie 335, 352-354

— tricochleatum, 452
Nyssa, 186

— hollrungii, 186

— sinensis, 186

Observations on the Problem of Transi-
ene

Prelim
the leat de Leen XVII, 370
Ochnaceae, 399
Ocotea spathulata 399, 408, 409, 412, 416
Olea apetala,

leeae, 365
Oleandra suena: 397
Orchidaceae, 398

1968]

Osmanthus, 364

— monticola, 369

Pachynema, 320
— oe 327

327
Paci arte aan — of XIX.
ew Hebrides,
ie Samoa, and te: nga, a
Palm Rhapis excelsa, Anatomy of the, V.
Inflorescence, 291; V Root and
Branch Insertion, 307
Palmae, 16-34, 398
ene pais 17
5

samoense, 449
— scutellaroides, 452
Paratropi ae 447
—sa

P

Pepe peta a, 397

roel 398, 408, 412
0

— 8
Puitieson, M. N., and W. R. PHizirson.
pene Nodal Types in Rhododendron,

nN, W. R., and M. N. Puizirson.
Diverse Nodal Types i in Rhododendron,

pis seme of Alangiaceae, Flowers, Fruits,
nd, 16

celsa, 369
Pico del Oeste, The Microclimate of. The
Ecology of an Elfin Forest in Puerto

52; 419
Pilea krugii, Lain 408, 412
— obtusata,
ae celine 408, 412
Piperaceae,
Pleea, 35, 36

INDEX

553

Plerandra, 432, 466, 467, 470, 473
ieee a 467-469

— graeffei

Rapaorsg St pic 469, 470
— grayi, 467, -474

— insolita, per 477

— nesopanax, 471
— pickeringii, 467, 474-476
— seemanni, 471

—victoriae, 467, 470, 471

— vitiensis, 467, 468, 471, 472
Polybotrya cervina, 397

—loriceum, 39

—44
— multijuga, si: 447-449
— neo-ebudar
— neo- ae 439, 443, 444
ape era 439, 443
— pin 452
ere 440, 450-452
—samoensis, 440, 449, 450
— scutellaria, 440, 452-455

Comparative Anat
of the Leaf- bearing Cocsaiiene: XVII,

370
Prestoea, 412
— montana, 397, 398, 408
Pseudobastardia, 279
Psychotria berteriana, 400, 408

Puerto Rico, The ology of an Elfin
Forest in, 1. neo oeprne and Co
positi nas Z; e Micro-

deere 507
mabe: Torminaria, 507

Rajania cordata, 398, 409, 412

554
Ranunculaceae, 233

m,

— ae pore Ar-
boretum During the Fiscal Year Ended
June 30, 1968, 525

—— of the Genus Notelaea (Ole-

ae), 333
Resnotsia, 432, 460, 465
na, 461, 463-465
—lanutoenss 460, 462, 463

ans, 378
Rhapis exenian, Anatomy of the Palm, V.
Inflorescence, 291; VI. Root and

Rhapis excelsa, 291-316
Rhododendron, Diverse Nodal Types in,
193

Rhododendron, 193-224
—sect. Brachycalyx, 198, 218

— sect. Candidastrum, 198, 212, 218

— sect. Choniastrum, 196, 198, 199, 205,
242,

= sect. Ruszalesatrans, 212,218

— sect. Lepiph pee

— sect.
— sect.
— sect.
— sect.
— sect.
— sect.
— sect. Trachythodion 222
—sect. Tsusiopsis

seats co

— sect. 218

—sect. Vire , 197, ie Si 212, 224
— sect. Viscidula, :

—ser. Albiflorum, 196, anal

—ser. An
— ser, Ar 19
— ser.
— ser.
— ser.
— ser.
— ser.
— ser.
— ser.
— ser.
— ser.
— ser.
— ser.
— ser.
— ser.
— ser.
— ser.
— ser. F
—ser. Fortunei, 221

JOURNAL OF THE ARNOLD ARBORETUM

[voL. 49

Rhododendron ser. Fulvum, 220
— ser. Glaucophyllum, 222

— ser,
— ser, Gri
— ser.
— ser.
— ser.
— ser.
—ser. L
— ser.
— ser.
— ser.
— ser.

wn
oO
ni
°
:o
2.
=]
oO
=]
$
RS
bo
w

— ser.
— ser,
— ser.
—ser. Saluenense, 223

— ser. Saxifragoides, 224

. Scabrifolium, 2

—ser. Semibarbatum, 196, 218

— ser. Stamineum a 205, 218
— ser. Stenophylla, 2

— ser. Taliense, 220

— ser. ifoli
— ser.
— ser.
— ser.
r. Uniflorum, 223
— ser.
— ser. peed

— Se eerie 196;:205,: 212, :214,

n
oO
oa

wn
oO

Be Hymenanthes, 198, 202, 205, 207,
211-21
as Pentanthera, 196;-205,° 207, 213;

; gy 196, 214, 2

. Pseudorhodorastrum, te 214,
Rhododendron, 196, 207, 214,
Rhodorastrum, 196, 207, 214,

. Therorhodion, 193, 194, 201
. Tsutsutsi, 196, one 213, 214, 218

. Cam
— subsect. Caroliniana, 223

1968]

INDEX

Rhododendron subsect. Cinnabarina, 223
b d 222

— subsect. E

— subsect.
— subsect.
220

— subsect.

subsect.

— subsect.

219
— sees

— subser,

, 224
Falconera, 205~ 207,-.210;-214.

Ferruginea, —

. Floribunda, 2

. Fortunea, - ee 221
Fulva, 220
proeccinioty 222

. Glauc

ct. Gra ast oe —- 220

seen Cagsar es
Trrorata,
- Lactea, a 205, 207, 220

. Maculifera, 207, 219

Maddenia, 197, 198, 212, 222

eeo
t. Taliensia, 202, = 220

Tephropepla,
. Thomsonia, oe ge 204, 206,

Triflora, 223

Boothii, 222
alophytum,

Campylocarpum, 219
anadense,

_ Crinigerum, 205, 218

. Davidii, 221
oe 221
. Fortun

sebonic gi anal 222
Glaucophyllum, 222

Rhododendron subser. Glischrum, 218
bs ri i 221

subser. Maculiferant, 219
subser. Maddenii, 222
subser. Megacalyx, 197, 222
subser. Megeratum, 222
subser. Neriiflorum, 221

subser. Selense, 219

pamnnean re ie 218
— barbatum, 205,

— basilicum, 205, eg 220
— bauhiniiflorum, 223

—calostrotum, 223

555

556 JOURNAL OF THE ARNOLD ARBORETUM

Rhododendron caloxanthum, 203, 219

—camelliaeflorum, 196, 2

—campanulatum, 203, i soe 210, 220
219

— Se 203,

— campylogynu 23

— camtschaticum, 201, 211, 218
3

nocarp’ 219
Bie leita 203, 219
— dauric

— — sempervirens , carolinianum, 207

— davidsonianum, 2
21

— eurysiphon,

— exasperatum, an 218
— eximeum, 22

— falconeri, 205, 220

— fargesii, 2

— ferrugineum, 223

Rhododendron igure 220

al’, 2
— aveeeearann a 218

— herzo
— hippophaeoides, 223
— hodgsonii, 205, 220
— hookeri, 204, 206, 219
— hunnewellianum, 219
— hyperythrum, 221
—idoneum, 223
Piariew yh 221

— insigne, 219
—- intrahieevatans, 197, 224
—invasiorum, 198, 224
— irroratum
os jasminifioram, 198, 224
— javanicum, 197, 224
— johnstoneanum, 197,222
— kaempferi, 198, 208, 218

2g

—laetum,
—lanatu 2

— latoucheae, 21
— lepidostylum, 222

S, 222
weit 197, 198, 224
— lindleyi, 197, 222
— litiense, 219

[voL. 49

1968]

Rhododendron lochae, 224
— longesquamatum, 219

— microleucum, 223

— minus, 223

— mollicomum

— mon ececiaoe on 219

Ben ores 198, 218
— rf

— nuttalli, py 205; 223

— perakense, 224

—‘P.J.M.’,
— dlanetums, 21
— pleianthum, 224

— pubescens, 222

INDEX 557

Rhododendron purpureiflorum, 224
2

_s fetivendaan, 197
— rex, 22

i pobineondd, 224
— roseum, 218
— roxieanum, 221
rubicncsudi, 223

— spathulatum, 224

~- spliaéroblastum, 2 21

— superbum, 197, 198, 224

— vale sol ror 222
— 218

Mt ona ote 222
— venator, 220

558

Rhododendron vernicosum, 221
— vinicolor, 224

, 220
— williamsianum, 204, 208, 210, 219
— wrayl, 21
— nae ese ten

Roytalossa verti
Rondeletia port iciiais. oe 408

cot and B A pee Insertion. Anatomy of

‘ie Palm Rhapis excelsa, VI, 307

Rosaceae, 402, 5

subfam. Maloideae, 504
Pinon . Sorboideae, 504
Roxburghiaceae, 260-277
Rubiaceae, 401

Sageretia of China, Botanical Name of

— taiwaniana, 378

— Thea, 378

— theerat s, 37

— toapriolia, 378

Samoa, and Tonga, The Araliaceae of the
New Hebrides, Fiji. Studies of Pacific
Island Plants, XIX, 431

Sapotaceae, 280-289, 400

Sarcandra irvingbaileyi, 5

Sauvagesia erecta,

— 432, 477-479, 481, 483, 486,

— pa Cephaloschefflera, 478, 490
7

wn
r=
ma
8
ie)
a
>
q
;

la,
— actinostiema, 7, 486, 487
— costata, 479,
— euthytricha, pm
— neo-ebudica, 478, 481, 482

— seemanniana, 479, 487-489

JOURNAL OF THE ARNOLD ARBORETUM

[voL. 49

Schefflera tannae, 479, 483
— vitiensis, 479, 483-486

Scutellaria ree 452
— tertia,
Selaginella wri 396

— substipitata, 396
Selagnlacene, 396
Sida c
pared tee mata, 287
— steyermarkii,

MITH, opiate a and BENJAMIN C.
STONE. BS gra wf Pacific Island Plants,
XIX. The Araliaceae of the New Heb-

rides, a Samoa, and Tonga, 431
Solanaceae, 402
Sorbus and the Problem of Generic Typ-

— sect. Cormus,

— aria, 5 8

— aucuparia, 507

— chamaemespilus, ey 508
— domestica, 507

Species of Bumelia he atic’ Jamai-
can and Other, 2

Species of Delph ae Sensu Stricto, A
Synopsis of ihe Asian, 23

Sphagnum, 4 oe

STE EARN, Wi1aM T. Jamaican and Oth
Species a Sonu (Sapotaceae), es

70-274

— tube y 270, 272
Stemonaceae, 260-277
Stet , 400

laris, 4

1o 2)
4
2)
g

LBERT C,
Situ. Studies of Pacific Island Plants.
XIX. he Araliaceae of the New
Hebrides, Fiji, Saena. and Tonga, 431

1968]

Stone, Donatp E., and Jupity L. FReeE-
Cytotaxonomy of Ilicium flori-
danum and I. parviflorum (Illiciaceae),

Strobilopanax, 434

— neo-ebudicus, 435

Studies in Dilleniaceae, Comparative
Morphological, III. The Carpels, 317

Studies of Pacific Island Plants, XIX. The

New Hebrides, Fiji,

1
Studies pie Anéiomsven in the Dichotom-
Venation of Circaeaster, Further

e, 400
Symplocos assebonggerk 400, 408, 412
Synopsis of the n Species of Del-
phinium, Sensu aos, A, 73, 233

Tabebuia a gee 400, 408, 412, 416

Tea Sageretia of China, Botanical Name
of the 307

Tea Sageretia, 377-379

Leaves, Preliminary Observations

Tetracera, 32

3
Tofieldia, 35, 36

INDEX

559
Tomurnson, P. B., d E. S. AYENsv.
Morpholo and Fase oof Croomia

pauciflora (Stemonaceae), 2
TomLinson, P. B., and H. E. odes Jr.
Inflorescence in Nannorrhops ritchiana

Tomuinson, P. B., and M. H. ZImMMEr-
MANN. Anatomy of the Palm Rhapis

excelsa, V. Inflorescence, 291; VI. Root
and Branch Insertion, 307

Tonga, The Araliaceae of the New Heb-
rides, Fiji, Samoa, and. Studies of
Pacific Island Plants, XIX, 431

Torralbasia cuneifolia, 399, 408

Trevesia pleiosperma, 461

Trichilia pallida, 399, 408, 412

Trichomanes crispum, 397

oo

Triplari

Typist, ee and the Problem of
Gener

Urticaceae, 399

Venation of Circaeaster, Further Mor-
on Anastomoses in

AN, REVATHI, and
Embevolony of Drinys

winterl, : 9
Verbenaceae, 402
Vriesia ak 398, 408, 409, 412

tts ogee aia 400, 408, 409, 412
mannia pinn
Wisteiome, 509, 1, 522525

ZIMMERMANN, M. H. d P. B. Tom
Linson. Anatomy of the Palm Rharis
ep V. Inflorescence, 291; VI.

t and Branch Insertion, 307

Ph abe 398