Skip to main content

Full text of "Journal of the Arnold Arboretum."

See other formats 


JOURNAL 


OF THE 
ARNOLD ARBORETUM 
HARVARD UNIVERSITY 
C. E. WOOD, JR. 


EDITOR 


L. I. NEVLING, JR. LAZELLA SCHWARTEN 
ASST, EDITOR CIRCULATION 


VOLUME XLII 


CAMBRIDGE, MASS. 
1962 


Reprinted with the permission of the 
Arnold Arboretum of Harvard University 


KRAUS REPRINT CORPORATION 
N or 


1968 


DATES OF ISSUE 
No. 1 (pp. 1-108) issued January 15, 1962. 
No. 2 (pp. 109-221) issued April 12, 1962. 
No. 3 (pp. 222-350) issued July 10, 1962. 
No. 4 (pp. 851-468) issued October 15, 1962. 


Printed in U.S.A. 


TABLE OF CONTENTS 


THE GENERA OF RUTACEAE IN THE SOUTHEASTERN UNITED STATES. 
By George K. Brizicky 
THE LEAF BASE IN PALMS, ae MorpHoLocy AND MECHANICAL 
BroLocy. By P. B. Tomlin on 
BOTANICAL AND OTHER onentene ON REponpA, THE WEST 
InpiEs. By Richard A. Howard 
A Taxonomic REVISION oF Popocarpus, XIII. Section Pory- 
PODIOPSIS IN THE SouTH Paciric. By Netta HE. Gray ............ 
TAXONOMIC AND NOMENCLATURAL NOTES ON ZANTHOXYLUM AND 
Gtycosmis (RuTACEAE). By George K. Brizicky ................ 
THE ae PIFICATION OF DiospyROS EBENUM AND DIOSPYROS EBEN- 
r. By Richard A. Howard and Tycho Norlinda ............ 
a ON THE RELATIONSHIPS OF PINUS MERKUSII.” By Nicholas 
T. Mirov 
STUDIES IN THE GENUS JASMINUM, IJ. THE Species rrRoM NEw 
ALEDONIA AND THE Loyatty IsLanps. By P. S. Green. ...... 
A CyroLocicaL Srupy OF THE GENUS ViBURNUM. By Donald R. 
Egolf 
THE GENERA OF SIMAROUBACEAE AND BURSERACEAE IN THE SourTH- 
EASTERN UnITep States. By George K. Brizicky .........0.0.05. 
CoMPARATIVE ANATOMY OF THE LEAF-BEARING CACTACEAE, IV. 
Tue Fusirorm INITIALS OF THE CAMBIUM AND THE ForM 
AND STRUCTURE OF THEIR Derivatives. By J. W. Bailey and 
Laht M. Srivastava 
ON THE ORIGIN OF CARAGANA SINICA. By Raymond J. Moore .... 
Suruyincuua, A New GENUS OF SCROPHULARIACEAB FROM CHINA 
By J. Pact 
PHENOLOGY or Tropical Pines. By Nicholas T. Mirov .............. 
ON THE Status oF Psiraea (THYMELAEACEAE). By Lorin I. Nevl- 


Ulta te 
JosErPH Horace Fautt, 1870-1961. With portrait. By Anna F. 
au 
COMPARATIVE ANATOMY OF THE LEAF-BEARING CACTACEAE, V. THE 
SeconDARY PuHuoEM. By Lalit M. Srivastava and EW: 
Bailey 
VOLCANISM AND VEGETATION IN THE LESSER ANTILLES. By Rich- 
ard A. Howard 
Tue GENERA OF PAPAVERACEAE AND FUMARIACEAE Pi THE SOUTH- 
EASTERN UNItTep States. By Wallace R. Ernst ........0.000... 
Note oN DapHNOPSIS CRASSIFOLIA fe ON By Lorin 
I. Nevling, Jr. 
Two New Asiatic PANDANACEAE. By Benjamin C. Stone ............ 


EeRNEsT JESSE PauMer, 1875-1962. With portrait. By Clarence 
BE. Kobuski 
THE GENERA OF ANACARDIACEAE IN THE SOUTHEASTERN UNITED 
States. By George K. Brizicky 
COMPARATIVE ANATOMY OF THE LEAF-BEARING Cacracear, VI. THE 
XYLEM OF PERESKIA SACHAROSA AND PERESKIA ACULEATA,. 
By I. W. Bailey 
SOME GUTTIFERAE OF THE LESSER ANTILLES. By Richard A. 
Howard 
DaTEs OF PUBLICATION OF THE JOURNAL LinnakaA. By Robert 
(’. Foster 
A MonocrapH OF THE GENUS PLATYDESMA (RUTACEAE). By 
Benjamin C. Stone 
THE THYMELAFACEAE IN THE SOUTHEASTERN UNITED States. By 
Lorin I. Nevling, Jr. 
THE LEITNERIACEAE IN THE SOUTHEASTERN UNITED States. By 
Rk. B. Channell and C. E. Wood, Jr. 
Tue Direcror’s Report 
BIBLIOGRAPHY OF THE PUBLISHED WRITINGS OF THE STAFF AND 
STUDENTS, JuLY 1, 1961—JUNE 30, 1962 
STAFF OF THE ARNOLD ARBORETUM, 1961-1962 
INDEX TO VoLUME XLIII 


JOURNAL 


OF THE 


ARNOLD ARBORETUM 


VoL. XLIII JANUARY 1962 NUMBER l 


THE GENERA OF RUTACEAE IN THE SOUTHEASTERN 
UNITED STATES * 


Georce K. Brizicky 


RUTACEAE Jussieu, Gen. Pl. 296. 1789. 
(RUE FAMILY) 


Armed or unarmed trees or shrubs [sometimes scandent or xeromor- 
phic], rarely herbs. Leaves alternate or more rarely opposite, simple or 
compound, usually glandular-punctate at least at the margin, exstipulate; 
petioles sometimes winged. Flowers bisexual and/or unisexual, the plants 
monoecious, dioecious, or polygamous, regular [rarely irregular], usually 
3—5-merous, the insertion hypogynous, solitary and axillary or in various 
axillary or terminal, often cymose, inflorescences. Sepals distinct or con- 
nate, very rarely wanting, often glandular-dotted, usually imbricate in 
bud. Petals distinct [rarely connate or wanting], often glandular-dotted, 
imbricate or valvate. Stamens as many as the petals and in 1 series 
(haplostemonous) or twice as many to more numerous and in 2 series 
(diplo- or obdiplostemonous), those of the outer series often shorter than 
those of the inner [or occasionally reduced to staminodes] ; filaments dis- 
tinct or -- connate, often conspicuously dilated [or rarely appendaged | 
at base; anthers versatile, introrse, 2[4]-locular at anthesis, often gland- 
tipped, longitudinally dehiscent. Intrastaminal nectariferous disc ring-, 
cup-, or cushion-like, rarely wanting. Gynoecium of (1)2-5(-several), 


the Arnold Arboretum and the Gray Herbarium of Harvard University which has 
been made possible through the support of George R. Cooley and the National Science 
Foundation. This treatment follows the pattern established in the first paper in the 
series (Jour. Arnold Arb. 39: 296-346. 1958) and continued through those in volumes 
40-42 (1959-1961). It should be repeated that the area covered by this work is 
bounded by and includes North Carolina, Tennessee, Arkansas, and I ouisiana. The 
descriptions are based primarily on the plants of this area, with any supplementary 
material in brackets. References which the author has not seen are marked by an 
asterisk. 

The author is indebted to Dr. Carroll E. Wood, Jr., for his criticism and valuable 
suggestions, and to Mrs. Gordon W. Dillon, for her careful help in the preparation 
of the manuscript. The illustration has been made by Dorothy H. Marsh under the 
direction of C. E. Wood, Jr. 


? JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


sessile or stipitate, distinct or incompletely to completely connate carpels; 
stigmas simple or lobed; styles basal, + lateral or terminal, distinct, con- 
nivent or connate; ovaries 1-locular or 2—5(—several) locular, with axile 
placentae [very rarely 1-locular with parietal placentae|; ovules usually 
anatropous and epitropous [rarely apotropous|, with 2 integuments and a 
thick nucellus, 1 or 2—many in each locule of the ovary. Fruit of (1)2— 
5(-several) follicles or drupes, or a capsule, berry, drupe, samara, or 
schizocarp; pericarp often glandular-pitted to -verruculose. Seeds with 
or without endosperm, sessile or funiculate, 1 or 2-several in a locule; 
embryo relatively large, straight or curved, with plano-convex, some- 
times convolute [rarely plicate] cotyledons and a superior radicle. Typr 
GENUS: Ruta L 


A family of about 150 genera and 1600 species, widely distributed in 
tropical and temperate regions, most abundant in tropical America, South 
Africa, and Australia, extremely scarce in Europe. Eight genera (five 
naturalized) in three subfamilies are represented in our area. 

The presence of secretory cavities (or at least inner multicellular glands) 
containing an aromatic volatile oil in stems (in cortex, rarely in phloem), 
leaves, floral parts, and pericarp of fruits is a characteristic feature of 
the family, distinguishing Rutaceae from their morphologically very simi- 
lar allies, especially Simaroubaceae and Meliaceae. Rutaceae are also 
closely related to Zygophyllaceae, Cneoraceae, and Burseraceae. 

The family, in general, seems to be entomophilous, insects being at- 
tracted by the strong smell of the flowers and/or by usually abundant 
nectar, sometimes also by showy corollas (e.g., Citrus spp.). Ornithophil- 
ly has also been presumed for some genera. Cross-pollination seems 
to be the rule. In some genera self-pollination is prevented by dicliny, 
or (in monoclinous genera) by proterandry (e.g., Ruta, Ravenia, Barosma) 
and/or the position of the stigma in regard to the anthers (e.g., Triphasia, 
Dictamnus). In some genera, however, both cross- and self-pollination 
seem equally possible and effective (e.g., Choisya, Skimmia, Murraya, 
Poncirus, Citrus). Cleistogamy has been recorded in the New Zealand 
alae simplex A. Cunn. Nucellar embryony (apomixis) has been 
shown in several genera (e.g., Zanthoxylum, Esenbeckia, Ptelea(?), Tri- 
phasia, Aegle, Murraya, Poncirus, Citrus), but probably is of even wider 
distribution in the family. Zygotic (gametic) polyembryony seems to be 
very rare, having been recorded or presumed in only a few cases. 

Chromosome counts have been made for about 50 genera and 150 species. 
On the basis of these counts, nine appears to be a basic number for the 
family. (Banerji [1954], however, suggested three as a basic number and 
nine as a result of secondary polyploidy for Citrus grandis (L.) Osbeck.) 
The family, in general, appears to be + euploid, except the Australian 
aneuploid tribe Boronieae (which has chromosome numbers based on 7, 8, 
9, 11, 13, 17, 19) and apparently some genera from other tribes. Poly- 
ploidy is widely distributed in Rutaceae and seems to be of importance 
in the evolutionary development of the family. 


1962 | BRIZICKY, GENERA OF RUTACEAE 3 


The family is of economic importance, notably for a number of im- 
portant fruits (Citrus spp., g.v.), timbers, aromatic oils (e.g., Citrus spp., 
Ruta graveolens L.), various products of medicinal value, and ornamentals. 
REFERENCES: 

CHAKRAVARTBY, R. 5S. Polyembry ony in Murraya Koenigii Spreng. Curr. Sci. 
Bangalore 3: 361, 362. 1935.* [Nucellar embryony. | 

———. Nucellar polyembryony in the Rutaceae. /bid. 5: 202, 203. 1936.* 
| Murraya exotica L. (= M. paniculata (L.) Jack) and Aegle Marmelos 
(i) Corral 

Desat, S. Cytology of Rutaceae and Simarubaceae. Cytologia 25: 28-35. 1960. 
[Includes chromosome counts for 8 genera and 10 spp. of Rutaceae; Ptelea 
trifoliata L., 2n = 42; Zanthoxylum americanum L., 2n = :36.] 

ENGLER, A. Rutaceae. Nat. Pflanzenfam. III. 4: 95-201. 15896 

—. Rutaceae. Nat. Pflanzenfam. ed. 2. 19a: 187-359, 458, “459. 1931. 

———. Studien tiber die Verwandtschaftsverhaltnisse der Rutaceae, Simaru- 
paces und Burseraceae nebst Beitragen zur Anatomie ae Systematik 
dieser Familien. Abh. Naturf. Ges. Halle 13: 111-158. pls. 12, 13. 1874. 

GALLeT, F. Développement et structure anatomique du tégument Te des 
Rutacées. Thesis, 66 pp. Univ. Paris, Ecole Supér. Pharm. 

Harti, D. Struktur und Herkunft des Endokarps der Rutaceen. Beitr. Biol. 
Pfi. 34: 35-49. 195 

Die eee a nnees des Endokarps der Simaroubaceen, Rutaceen 
Leguminosen. /bid. 34: 452-455. 1958 

pace E. Osservazioni sulla struttura dell’ovulo e sulla cariologia di Calo- 
dendron capense Thunb. e Pilocarpus pennatifolius Lem. Ann. Bot. Roma 
24: 438-448. pls. 19-21. 1954. [Includes brief survey of chromosome 
numbers in Rutaceae. | 

Jourr, B. M., & M. R. Anvya. A contribution to the floral morphology and 
embryology of Aegle Marmelos Correa. Phytomorphology 7: 10-24. 1957. 
| Anthers 4- locular at anthesis; nucellar embryony the rule. | 

Mauritzon, J. Uber die Embryologie der Familie Rutaceae. Sv. Bot. Tidskr. 
29: B1902347. 1935. 

Moore, J. A. Floral anatomy and phylogeny in the Rutaceae. New Phytol. 

: 318-322. 1936. [Phylogenetic considerations based on Saunders’ data 
of floral anatomy. | 

Necopr, G. Lineamenti sulla cariologia delle Rutaceae e delle ewes. 
Avo Bot. Forli 13: 93-102. 1937. [Includes Ruta graveolens, 2n 
Ptelea trifoliata, 2n = 36; Citrus Limon and C, simensis, 2n = 18. 

Penzic, O. Studi botanici sugli agrumi e sulle Sh affine. Ann. Agr. Roma 
1887: i-vi, 1-590; 58 pls. [atlas (folio) ]. 

Recorp, S. J., & R. W. Hess. American woods ae as family Rutaceae. Trop. 
Woods 64: 1-28. 1940. 

SaunpeERrS, E. R. On carpel polymorphism. VI. Ann. Bot. 48: 643-692. 1934. 
[Rutaceae, 643-673. | 

Scuuize, H. Beitrage zur Blattanatomie der Rutaceen. Beih. Bot. Centralbl. 
12: 55-98. pls. 1, 2. 1902. 

SMITH-WHITE, 8. Chromosome numbers in the Boronieae (Rutaceae) and 
their bearing on the evolutionary development of the tribe in the Australian 
flora. Austral. Jour. Bot. 2: 287-303. 1954. [Basic numbers 7-19; 69 spp. 
in 11 genera; aneuploidy and polyploidy. | 


4 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLII 


SWINGLE, W. T. A new taxonomic arrangement of the orange subfamily, Auran- 
fo dene: Jour. Wash. Acad. Sci. 28: 530-533. 1938. 

. The botany of Citrus and its wild relatives of the orange subfamily 
(family Rutaceae, subfamily Aurantioideae). Chap. IV, pp. 129-474. In: 
Wesser, H. J., & L. D. BatcHetor, The Citrus Industry. Vol. I. His- 
tory, Botany, and Breeding. xx + 1028 pp., front. Univ. Calif. Press, 
Berkeley & Los Angeles. 1943. 

TANAKA, T. The taxonomy and nomenclature of Rutaceae-Aurantioideae. 
Blumea 2: 101-110. 1936. 

Titison, A. H., & R. Bamrorp. The floral anatomy of the Aurantioideae. Am. 
Jour. Bot. 25: 780-793. 1938. [Vascular system does not offer a definite 
basis for division into the tribes and subtribes; the disc appears to repre- 
sent a third whorl of vestigial stamens. 

Urzan, I. Zur Biologie und Morphologie der Rutaceen. Jahrb. Bot. Gart. Ber- 
lin 2: 366-403. pl. 13. 1883. 

Witson, P. Rutaceae. N. Am. Fl. 25: 173-224. 1911. 


KEY TO THE GENERA OF RUTACEAE 


neral characters: leaves usually pinnate to 3-1-foliolate, alternate, rarel 
ee leaflets a least at margin), perianth parts, and fruits glandular. 
often pellucid-punctat 

Plant a perennial herb or subshrub; leaves 2- to 3-pinnately divided; flowers 

greenish yellow, bisexual, 4—5-merous with 8-10 stamens; fruits 5-lobed, 

WAY SOCCEO CAPSULES. 6 bores oe hi-wd sx PAE RS wie base Peeing 1. Ruta. 

. Plants woody, shrubs or trees. 

B. Flowers unisexual or uni- and bisexual, small, yellowish or yellowish 
white, 3-5-merous, sometimes without calyx; stamens as many as petals; 
Soar apo- or syncarpous, (1)2-5-carpellate; fruits dry; leaves 
alternate. 


> 


i Plants dioecious or monoecious, often prickly; leaves pinnate, usually 

5 or more foliolate; flowers unisexual, with or without sepals; gynoe- 

cium apocarpous, (1)2—S-carpellate; fruits dehiscent, 1-seeded fol- 

WGN Gas yoc4 sees wie ae, us Bares one Rha eeee oe ees 2. Zanthoxylum,. 

C. Plants polygamous, unarmed; leaves usually 3-foliolate; flowers uni 
and bisexual; gynoecium syncarpous, 2-loculate; fruit a flat, inde- 
hiscent, 2-locular, 2-winged samara. .................... 3. Ptelea, 

B. Flowers usually bisexdal small to large, usually white, 3- 5- -merous; 
stamens twice as many as petals, or more numerous; gynoecium syn- 


carpous, 2—18-locular, rarely l-carpellate; fruits fleshy drupes or arene 
leaves alternate or opposite. 

D. Stamens twice as many as petals; flowers relatively small; ovary 1—5- 

loculate; fruits small, pulp without pulp-vesicles; petioles not winged. 

E. Ovary 1-carpellate; fruit a 1-seeded drupe; leaves opposite, 3- 

foliolate; flowers usually 4-merous, paniculate; era: aro- 

matic, resinous shrubs or trees. .................... 4. Amyris. 

E. Ovary 2-5-carpellate; fruit a “ 3-seeded berry; leaves alternate. 

F. Flowers (4)5-merous, in short, spikelike axillary panicles; berry 

subglobular, sometimes depressed at eke and slightly oblique, 

white to pink, 1-3-seeded, edible; unarmed shrubs or trees 

with pinnate, usually 1-3(5)-foliolate piers and large leaflets. 

gee ehh dete ete ee esau tas 2 NG ane baht State 5. Glycosmis. 


1962] BRIZICKY, GENERA OF RUTACEAE 5 


F. Flowers usually 3-merous, solitary or in 2’s or 3’s in the leaf 
axils; berry subglobular, sometimes apiculate, reddish orange to 
crimson, 1-3-seeded, insipid; shrubs with solitary or paired, 
axillary spines; leaves 3-foliolate, leaflets cael: Leena cerrare te tcc eeres 

eh en ee NERD RRS eyecare d or fo caea 6. Triphasia. 

D. Stamens numerous (20-60); flowers relatively large; perianth usually 

5-merous; ovary 6-18-loculate; fruits (hesperidia) greenish yellow, 

yellow, orange to reddish orange, large, pulp formed by pulp vesicles; 
usually thorny shrubs or trees, mostly with winged petioles. 

G. Flowers on the previous year’s branchlets; stamens distinct; ovary 
6—-8-loculate; fruits relatively large, pubescent, pulp with very 
sour and acrid juice; leaves 3-foliolate, deciduous. ... 7. Poncirus. 

_ Flowers on new branchlets; stamens polyadelphous; ovary 8~-18- 
loculate; fruits large, glabrous, pulp with sweet or sour not 
acrid juice; leaves 1-foliolate, persistent. ............ 8. Citrus. 


7p) 


Subfam. RUTOIDEAE Engler 
1. Ruta Linnaeus, Sp. Pl. 1: 383. 1753; Gen. Pl. ed. 5. 180. 1754. 


Heavy-scented perennial herbs, subshrubs [or shrubs]. Leaves alter- 
nate, usually glandular-punctate, compound [or simple], odd-pinnate to 
-bipinnate, with pinnae or pinnules respectively deeply cut (divided) into 
obovate-cuneate to oblanceolate or oblong segments. Flowers bisexual, 
4- or 5-merous, in terminal, panicled cymes with simple or 3-fid bracts. 
Sepals + connate at base, persistent. Petals yellow or greenish, glandular- 
punctate, spatulate-cochleariform with incurved, hooded apex, + clawed, 
denticulate [fimbriate or entire], imbricate in bud. Stamens 8-10, in 2 
series, the outer (antipetalous stamens) usually somewhat shorter than the 
inner whorl (antisepalous stamens) ; filaments filiform, broadened toward 


on an intrastaminal, cushion-like, nectariferous disc; stigma small; style 
central; ovary deeply 4- or 5-lobed, 4- or 5-locular, with [2 to] numerous 
ovules on axile placentae. Capsule glandular-punctate, 4- or 5-lobed, 4- or 
S-locular, [few- to] many-seeded, dehiscent loculicidally inward (adaxi- 
ally) at apex, [or split into indehiscent segments]. Seeds angled, brown, 
tuberculate; endosperm fleshy; embryo slightly curved, cotyledons some- 
times 2-lobed. (Including Haplophyllum A. L. Juss.). LECTOTYPE SPECIES: 
R. graveolens L.; see P. Wilson, N. Am. FI. 25: 212. 1911. (Classical 
Latin name of the plant [since Cicero in literature], related to and ques- 
tionably derived from Greek, rAyte, the name of the plant in Nicander; 
etymology obscure.) — RUE. 


A genus of about 60 species, ranging from Macaronesia eastward 
through the Mediterranean region to central Asia and eastern Siberia. 
Ruta graveolens L., common rue, 2” = 72, 81, native to the Mediter- 
ranean region but widely naturalized in temperate parts of the Old World, 
is’ introduced and more or less naturalized in the eastern United States. 
There are few reliable records from our area and further data regarding 


6 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


the distribution of this species in the southeastern United States are very 
desirable. 

The species is proterandrous. The stamens execute peculiar nutation 
movements during the expansion of the flowers. Pollination agents re- 
corded are small Hymenoptera, Coleoptera, and Diptera. Polyploidy 


tetraploid, and octoploid species are known. The occurrence of a nonaploid 
chromosomal race of the otherwise octoploid Ruta graveolens has also 
been recorded. However, no general conclusions can yet be drawn on the 
basis of the few counts. 

The common rue is of ancient culture as an ornamental, spice, and 
medicinal plant. Because of a volatile oil (“oil of rue”) it was formerly 
used in medicine, but at present, its use is quite limited because of very 
unpleasant secondary effects. The species should be regarded as + poison- 
ous. Contact with fresh leaves of the plant produces a dermatitis in some 
individuals. Poisoning by oil of rue is characterized by gastroenteritis. 
The Mediterranean R. chalapensis L. and some other species have also 
been used locally in much the same way as R. graveolens. The latter 
and a few other species are cultivated as ornamentals. 


REFERENCES: 

See also under family references ENGLER (1931. pp. 243-246), GALLeT (1913, 
pp. 23-27), Necopr (1937, pp. 93-97, 101), UrBan (1883, pp. 372, 373, 401), 
WILSON (1911, p. 212). 

AHLEs, H. E., C. R. Bett, & A. E. RADFORD. Species new to the flora of North 
or South Carolina. Rhodora 60: 10-32, 1958. [R. graveolens, Orange 


ALLEN, N. Beware of garden rue. Horticulture 24: 403. 1946. [Poisoning by 
t 


contact. 
BeRSILLOoN, G. Les inflorescences de Ruta graveolens L. Revue Gén. Bot. 63: 
437-460. 1956. 

Branpt, W. Zur Anatomie und Chemie der Ruta graveolens L. Thesis, 33 pp. 
Berlin. 1915.* [See review in Bot. Jahresb. 43¢1)3 S10, 511, 1913.4 
CAPPELLETT!, C. Sterilita di origine micotica nella Ruta patavina L. Ann. Bot. 

Roma 18: 145-166. pls. 6,7. 1 ; 18; includes the embryology. ] 
Outa, T., & T. Mryazaxr. Furoquinolines. XIII. Alkaloids from the pericarps 
of Kuta graveolens L. (In Japanese.) Jour. Pharm. Soe. Japan 78: 538, 
539. 1958. 
SovEGES, R. Développement de lembryon chez le Ruta graveolens L. Bull. Soc. 
Bot. Fr. 73: 245-260. 1926 


2, Zanthoxylum Linnaeus, Sp. Pl. 1: 270. 1753: Gen. Pl. ed. Oe. LOU, 
1754. 


Deciduous or evergreen trees or shrubs. often armed with prickles 
which sometimes become elevated on broad-conical or -pyramidal corky 
excrescences; bark aromatic. Leaves alternate. odd- or even-pinnate to 
1-foliolate; leaflets opposite or alternate. frequently inequilateral, crenu- 
late or entire, glandular-punctate, at least at the margin; petiole and 


1962 | BRIZICKY, GENERA OF RUTACEAE 7 


rachis winged or wingless, unarmed or prickly. Plants dioecious, monoeci- 
ous, [or polygamous]; flowers small, white to greenish yellow, unisexual 
[and/or bisexual], in axillary short spikes or cymose fascicles or in ter- 
minal, sometimes corymbiform, panicles. Sepals 3-5[10], distinct or 
+ connate, deciduous or persistent, or apparently wanting. Petals 3-5 [8], 
imbricate [or valvate| in bud. Stamens 3-5[8], distinct, alternate with 
the petals, rudimentary (staminodial) or wanting (or sometimes trans- 
formed into carpels) in ¢ flowers; filaments filiform to subulate; anthers 
ovate, elliptic to subcircular in outline. Intrastaminal disc small, often 
pulvinate, or obscure. Gynoecium of (1)2—-5 sessile or stipitate, distinct 
or partially united [very rarely completely connate] carpels, rudimentary 
in g flowers; stigmas capitate, distinct or connate; styles sublateral, dis- 
tinct, connivent or connate toward the summit; ovaries usually 1-carpellate 
and -locular [very rarely ovary compound, 2—5-carpellate and -locular | 
with 2 collateral, pendulous ovules in each carpel [or locule]. Fruits 2- 
valved follicles, distinct or connate at base. stipitate to sessile, firm-walled 
or fleshy, glandular-punctate, with separating (loose) or adherent endo- 
carp, 1(2)-seeded. Seeds obovoid to subglobular [or + lenticular], black 
[blue-black, brown, or dark red], shining, with a crustaceous testa and 
fleshy endosperm, at maturity often hanging from the carpels on slender 
funicles: embryo axial, straight or somewhat curved, with a short radicle 
and flat, almost circular cotyledons. (Including Fagara L., nom. cons., 
type: F. Pterota L.) Type species: Z. americanum Mill. (‘“Xanthoxy- 
lum”) (Z. fraxineum Willd.); see F. R. Fosberg, Taxon 8: 103-105. 
1959. (Name from Greek, xanthos, yellow, and xylon, wood.) — PRICKLY 
ASH. 


A genus of about 215 species, primarily pantropical, extending with 
several species into the Temperate Zone of eastern Asia and North 
America. Two subgenera sometimes regarded as distinct genera are 
recognized in the present treatment.’ 

Subgenus ZANTHOXYLUM, with unisexual flowers with petaloid [or 
sometimes sepaloid], 4—5{10|-merous, “simple” perianth in a single series 
(presumably petals) and 4-5[8| stamens or [1]3-5 carpels, includes 
about 15 species, primarily of the Temperate Zone of eastern Asia and 
North America, but with at least two species in Central America. Zan- 
thoxylum americanum Mill., common or northern prickly ash, 21 = 68, 
136, a shrub or small tree with odd-pinnate leaves, paired pseudostipular 
prickles (rarely prickleless) and yellow-green flowers in sessile, axillary, 
umbellate clusters, expanding before leaves, is the only eastern North 
American species of the subgenus, occurring from Georgia and Alabama 
northward beyond our area to North Dakota, Minnesota, Ontario, and 
western Quebec. Zanthoxylum mazatlanum Sandwith is known from 


>The occurrence of species apparently transitional in the character of the perianth 
between Zanthoxylum and Fagara is ample reason to regard both as components of a 
single genus. See “Taxonomic and Nomenclatural Notes on Zanthoxylum and Glycos- 
mis (Rutaceae)” in the present issue of this Journal. 


8 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


Mexico; Z. Williamsii Standl. and Z. ferrugineum Radlk. occur in Hon- 
duras and Costa Rica respectively. 

Subgenus Facara (L.) Triana & Planch. (Fagara L.), with unisexual 

[or both uni- and bisexual] flowers with 3-5[—8]-merous perianth in two 
series (i.e., sepals and petals), 3-5[—7] stamens and/or 3—5 carpels, in- 
cludes about 200 species, primarily of the tropics of both hemispheres, 
four in our area. Zanthoxylum Clava-Herculis L., southern prickly ash, 
Hercules’ club, 2x = ca. 72, armed trees or shrubs with odd-pinnate leaves 
and usually 5-merous flowers in terminal panicles, occurs on the Coastal 
Plain from southern peninsular Florida to Texas and southeastern Vir- 
ginia, northward to southern Arkansas and Oklahoma, The South Ameri- 
can-West Indian Z. Fagara (L.) Sarg. (Fagara Pterota L.), an ever- 
green shrub or tree usually armed with pseudostipular prickles, with 
odd-pinnate leaves having relatively small leaflets and winged petioles and 
rachises, and 4-merous flowers in short axillary spikes, reaches its north- 
ern limit in central Florida and in southern and southwestern Texas beyond 
our area. The West Indian Z. coriaceum A. Rich., a + prickly tree or 
shrub, usually with even-pinnate, coriaceous leaves and 3-merous flowers 
in terminal panicles, is known from southern peninsular Florida and the 
Florida Keys. The West Indian Z. flavum Vahl, yellow-wood, satin- 
wood, an unarmed tree with odd-pinnate, occasionally 3-1-foliolate leaves 
and S-merous flowers in terminal panicles, occurs on the lower Florida 
Keys. 
Little is known regarding pollination, but bees and various Diptera have 
been recorded as the most frequent visitors of the flowers of Z. americanum. 
Nucellar polyembryony has been found in a few species (e.g., Z. ameri- 
canum, “Z. Bungei Planch.,” Z. alatum Roxb.), but probably is of wider 
distribution within the genus. The few published chromosome counts 
(2n = 32, 64, 68, 70, 72, 136) indicate polyploidy and perhaps aneuploidy. 
No interspecific hybrids have been recorded, A modern monograph is 
highly desirable. 

The dried bark of Zanthoxylum americanum and Z. Clava-Herculis, 
“toothache bark,” “prickly ash bark,” or “xanthoxylum,” has been ap- 
plied as a stimulant, tonic, and sialagogue in the United States, and the 
bark of 2. alatum has been used against fever, dispepsia, diarrhoea, and 
cholera in India. Zanthoxylum americanum and some eastern Asiatic 
species are sometimes cultivated as ornamentals. Various other species 
have been used for timber, spices, and medicines. 


REFERENCES: 
See also under family references Desar (1960, p. 32), ENGLER (1895, pp. 
115-119; 1931, pp. 214-224), and Wrtson (1911, pp. 177-199). 
Barser, C. A. The nature and development of corky excrescences on the stem 
of Zanthoxylum. Ann. Bot. 6: 155-166. pls. 7, 8. 1892. 
BocquILton, H. Etude botanique et pharmacologique des Xanthoxylées. Thesis. 
125 pp., 4 pls. Univ. Paris, Ecole Supér. Pharm. 1901. 
Fosperc, F. R. Typification of Zanthoxylum L. Taxon 8: 103-105. 1959, 


1962] BRIZICKY, GENERA OF RUTACEAE 9 


[Type sp.: Z. fraxineum Willd. (2. americanum Mill.) ; see also Taxon 7: 
94-96. 1958. | 

Goto, J. Studies on the toxic principle of Xanthoxylum piperitum De Can- 
dolle. Jap. Jour. Veterin. Sci. 17: 205-215. 1955.* 

Huanc, C. C. Preliminary study on Chinese Rutaceae (1). (In Chinese.) 
Acta Phytotax. Sinica 6: 1-143. pls. 1-36. 1957. [Zanthoxylum, 6-83, 
pls. 1-19.] 

Lonco, B. La poliembrionia nello Xanthoxylum Bungei Planch. senza feconda- 
zione. Bull. Soc. Bot. Ital. 1908: 113-115. 1908. 

Reever, J. R. Xanthoxylum Miller (1768). Taxon 4: 237. 1955. { Proposed 
for conservation; see also L. H. SHINNERS, Taxon 6: 135-137. 1957, against 
conservation. 

SARGENT, C. S. Xanthoxylum. Sylva N. Am. 1: 65-74. pls. 29-32. 1891; 14: 
97-98. 1902. 

Torro, F. G. The gum of Fagara xanthoxyloides. Nature 180: $64, 865. 1957.* 

WirtH, E. H. Xanthoxylum. Natl. Formulary Bull. 9: 149, 150. 1941.* 


Subfam. TODDALIOIDEAE Engler 
3. Ptelea Linnaeus, Sp. Pl. 1: 118. 1753; Gen. Pl. ed.'5. 54. 1754. 


Unarmed shrubs or small trees, with bitter bark and foliage disagree- 
ably scented when crushed. Leaves alternate, usually 3-foliolate, rarely 
4-S-foliolate, leaflets entire or toothed, glandular-punctate, glabrous to 
densely soft-hairy beneath. Plants usually polygamous, the flowers bi- 
sexual and/or unisexual, greenish or yellowish white, aromatic, in termi- 
nal corymbiform, cymose panicles. Sepals usually 4 or 5, distinct, im- 
bricate. Petals usually 4 or 5, relatively narrow, surpassing the sepals, 
imbricate. Stamens usually 4 or 5, alternating with petals, hypogynous, 
inserted at base of a disc, very short, with imperfect sterile anthers in 
9 flowers; filaments subulate, hairy in the lower half; anthers ovate- 
cordate, introrse. Gynoecium syncarpous, usually 2-carpellate, inserted 
on a low disc in bisexual and ¢ flowers, very small, imperfect (lacking 
style and with rudimentary stigmas), raised on a conspicuous truncate- 
pyramidal to subglobular disc in ¢ flowers; stigma capitate, usually 
2-lobed; style relatively short and slender; ovary compressed, usually 
2-locular and narrowly 2-winged, with 2 superposed ovules (the lower 
usually sterile) on axile placenta in each locule. Fruit a flat, subcircular 
to obovate, glandular-punctate samara with 2 broad, thin, reticulate lat- 
eral wings completely encircling the indehiscent 2-locular and 1(2)-seeded 
body. Seeds laterally compressed, semiovate to semilanceolate in outline, 
acute at apex, rounded at base, dark reddish brown to black, densely 
papillose and glossy on the surface; seed coat thin, leathery; endosperm 
fleshy, thin; embryo large, straight, with oblong to ellipsoid cotyledons 
and short, stoutish superior radicle; germination epigeous. LECTOTYPE 
species: P. trifoliata L.; see P. Wilson, N. Am. FI. 25: 208. 1911. (Classi- 
cal Greek name of elm, Ulmus, transferred by Linnaeus to this genus on 
account of the similar fruit.) — HoP-TREE, SHRUBBY TREFOIL. 


A genus of three (according to P. Wilson) or probably more species, 


10 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


of temperate North America. Ptelea trifoliata L. (including P. micro- 
carpa Small fide Wilson), common hop-tree, stinking ash, 2n = 36, 42, a 
shrub or small tree, very variable in regard to size, shape, and pubescence 
of the leaflets and size and shape of fruits, occurs from northern Florida, 
throughout our area, and beyond to Texas and northern Mexico, Nebraska, 
Towa, Michigan, southern Ontario, New York, and Connecticut. Its vari- 
ant var. mollis Torrey & Gray, regarded by Wilson as P. tomentosa Rai 
with branchlets, lower surface of the leaflets and inflorescences subtomen- 
tose, ranges from Georgia to North Carolina and westward to Oklahoma, 
Arizona, and Mexico. Ptelea serrata Small, an irregularly branched shrub 
with shallowly serrate leaflets, probably belongs with the preceding. 
Ptetea Baldwinii Torrey & Gray, a shrub with leaflets smaller and nar- 
rower than in P. trifoliata, has been recorded only from northeastern 
Florida in our area but is apparently conspecific with P. angustifolia 
Benth., widely distributed in the western United States from southern 
Texas to Colorado, Utah, California, New Mexico, and Arizona, and 
southward into Mexico. 

Vegetative reproduction by root sprouts has been recorded in Ptelea 
trifoliata. e occasional occurrence of 3- or 4-carpellate ovaries, some- 
times accompanied by a few additional antipetalous stamens, is note- 
worthy. Polygamy seems to be a generic character, Although herbarium 
specimens usually are represented either by male or by female inflores- 
cences, sometimes both bisexual and male flowers occur in the same in- 
florescence; more rarely a few female flowers (usually terminal) are 
found in the male inflorescence. Cross-pollination is the rule because of 
the predominance of unisexual flowers, but self-pollination seems to be 
possible in the bisexual flowers. Bees, especially short-tongued species, 
and a few other Hymenoptera and Diptera have been recorded as polli- 
nators. The occurrence of a nucellar embryo has been recorded in P. 
trifoliata (Mauritzon, 1935), but the observations (made on scanty ma- 
terial) need verification. Only two chromosome counts are reported for 


the genus. No hybrids have been recorded, ut, Desai (1960), investi- 
gating the cytology of P. ¢trifoliata grown in England (2 = 42), ob- 


served meiotic irregularities resulting in high sterility and concluded a 
possible hybrid origin for the plant. 

Over 60 species of Ptelea have been described, but these are apparently 
not well understood at present, and the reduction of all of them to two or 
three species is open to question. A modern monograph is urgently needed. 

Ptelea trifoliata is a generally known ornamental plant. Its bitter fruits 
have sometimes been used in brewing as a substitute for hops, hence the 
English name of the genus. The bark is reputed to possess medicinal 
properties as a weak tonic. 


REFERENCES: 

See also under family references Desar (1960, pp. 29, 30), ENGLER (1931, 
pp. 302, 303), Maurirzon (1935, pp. 339, 346), URBAN (1883, pp. 397, 398), 
and WiLson (1911, pp. 208-210). 


1962} BRIZICKY, GENERA OF RUTACEAE 11 


Baitey, V. L. Historical review of Pfelea trifoliata in botanical and medical 
literature. Econ. Bot. 14: 180-188. 1960. 

GREENE, E. L. Some Piéelea ss Torreya 5: 99, 100. 1905. [Descrip- 
tions of 3 new spp., 2 of the se. U. S.] 

. The genus Prelea in a western and southwestern United States and 
Mexico. Contr. U. S. Natl. Herb. 10: 49-78. 1906. [59 spp. recognized, 
including 55 new. | 

Harris, J. A. Teratological fruits of Ptelea. Bull. Torrey Bot. Club 38: 385- 
387. pl. 17. 1911. 

SARGENT, C.S. Ptelea. Sylva N. Am. 1: 75-78. pls. 33, 34. 1891; 14: 98. 1902. 

ScHROEDER, E. M. Germination of fruits of Ptelea species. Contr. Boyce 
Thompson Inst. 8: 355-359. 1937. 

Wixson, P. Notes on Rutaceae — V. Species characters in Pfelea and Taravalia. 
Bull. Torrey Bot. Club 38: 295-297. 1911 


4. Amyris Linnaeus, Syst. Nat. ed. 10. 2: 1000, 1367. 1759. 


Usually glabrous shrubs or trees with resinous, fragrant wood. Leaves 
opposite [subopposite or alternate], odd-pinnate, often 3—-5[rarely 1]- 
foliolate, with glandular-punctate leaflets and unwinged [or winged] 
petioles. Flowers small, bisexual [rarely unisexual], usually 4[rarely 3 
or 5]|-merous, pediceled, in terminal or axillary panicled cymes. Calyx 
cuplike, 4-lobed, glandular-dotted, persistent. Petals 4, white, glandular- 
dotted, imbricate in bud. Stamens 8, in 2 series, inserted at base of a disc 
or of the ovary; filaments filiform; anthers ovate to oblong, introrse, 
2-locular at anthesis. Intrastaminal disc pulvinate, gynophore-like, sup- 
porting gynoecium, or wanting. Gynoecium 1-carpellate; stigma capitate 
to discoid-subcapitate; style very short and stout or wanting; ovary 
1-locular, with 2 collateral ovules suspended from the top of the locule. 
Drupes globular, ellipsoidal to obovate, black [or reddish], often glaucous, 
dotted with glands, aromatic, oily, the endocarp chartaceous, 1-seeded 
Seed pendulous, with thin, membranaceous testa, lacking endosperm; em- 
bryo with plano-convex, fleshy, glandular-dotted cotyledons and a short, 
superior radicle. Lecrorypr species: A. balsamifera L.; see P. Wilson, 
N. Am. Fl. 25: 216. 1911. (Name apparently derived from Greek, a, 
with, abounding in, and myron, balsamic juice, resin, with reference to 
the balsamic properties of the genus.) *— TorcHwoop. 


A genus of about 20 species of the West Indies and especially of cis- 
Amazonian tropical America, extending southward (with one species?) to 
Peru and northward (with a few species) to Texas and Florida, The 
South American-West Indian A. balsamifera L., with 3—5-foliolate leaves, 
leaflets dull underneath, puberulous inflorescences, and puberulous, stipi- 
tate ovary, occurs in southern peninsular Florida and the Florida Keys. 
The primarily West Indian A. elemifera L., differing from the preceding 
especially in the glabrous inflorescences and glabrous, sessile ovaries, is 

* More frequently the Greek prefix a has been used in a reverse sense, without, 
devoid of, not. This has led Little (U.S. Dep. Agr. Handb. 41: 57. 1953) to a differ- 
ent, hardly probable derivation: “not myrrh,” “not true myrrh.” 


12 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


also known from southern peninsular Florida and the Keys, and the very 
similar A. maritima Jacq., with stipitate ovaries, has been recorded from 
Key West. Wilson and Small included this last species in 4. elemifera, 
but Urban (1896, 1920) regarded it as distinct. 

The genus is, in general, very imperfectly known. The species, especially 
the West Indian, are closely related, and their delimitation is often weak, 
sometimes being based on few, questionably specific characters. A modern 
revision based on field observations, experimental cultures, and cytology 
is very desirable. 

The resinous timber, especially that of Amyris balsamifera and A. 
elemifera, is of excellent quality but is scarce and of small size. It is 
used locally for fuel, torches (fragrant when burned), small cabinet work, 
and to a limited extent as a source of an oleoresin, Mexican elemi, which 
is used locally in medicine. 


REFERENCES: 
See also under family references ENGLER (1931, pp. 313, 314) and Wrrson 
(1911, pp. 216-220). 
SARGENT, C. S. Amyris. Sylva N. Am, 1: 83-86. pl. 36. 1891; 14: 98. 1902. 
Ursan, I. Additamenta ad cognitionem florae Indiae occidentalis. Particula III. 
Bot. Jahrb. 21: 514-638. 1896. [Amyris, morphological-taxonomic intro- 
duction, key to and description of the spp., abundant synonymy, 595-612. ] 
———. Amyris L. Symb. Antill. 8: 323. 1920. [Includes synonymy and gen- 
eral distribution of our spp. | 


Subfam. AURANTIOIDEAE Engler 
5. Glycosmis Corréa, Ann. Mus. Hist. Nat. Paris 6: 384. 1805, 


Unarmed, evergreen shrubs or trees. Leaves alternate [rarely opposite], 
odd-pinnate, often 1-3(5)-foliolate; leaflets pellucid-punctate, entire [ser- 
rate or crenulate], + inequilateral at base; petiole articulated with the 
blade in 1-foliolate leaves. Flowers relatively small, fragrant, bisexual, 
(4)5-merous, with very short, stout pedicels, in rusty-villosulous, spike- 
like cymes arranged in axillary panicles. Sepals (4)5, nearly distinct 
[or + connate], triangular-semicircular, + fleshy, imbricate. Petals (4)5, 
elongate, somewhat concave, white, imbricate. Stamens (8)10, the anti- 
petalous shorter than the alternipetalous, filaments flattish [or filiform], 
dilated upward [or downward] and abruptly narrowed into an acuminate 
tip; anthers relatively short, ovate-cordate, often apiculate, gland-tipped. 
Gynoecium 2-—5-carpellate, syncarpous, raised on a cushion-like [or cylin- 
drical] nectariferous disc; stigma broad, cushion-like (convex-disciform 
to semiorbicular), persistent; style often indistinct, short and very stout, 
merging into a subglobular or ellipsoidal 2—5-locular ovary covered with 
glands; ovules pendulous, one in each locule. Fruit a relatively small berry 
with thin pulp, + globular, sometimes depressed at apex and -£ oblique, 
1—3-seeded. Seeds without endosperm, ellipsoid to subglobular, with a 
membranaceous testa; embryo with plano-convex cotyledons and a short 


1962 | BRIZICKY, GENERA OF RUTACEAE 13 


superior radicle. LecroTyPE sPEcIES: G. arborea (Roxb.) DC. (Limonia 
arborea Roxb.); see P. Wilson, N. Am. Fl. 25: 215. 1911.4 (Name from 
Greek, glycys, sweet, and osme, scent, odor, referring to the fragrant flow- 
ers of the genus. 


A genus of about 35 species of southeastern Asia, the East Indies, the 
Philippines, New Guinea, and northeastern Australia. The southeastern 
Asiatic Glycosmis parviflora (Sims) Little (G. citrifolia (Willd.) Lindl.), 
with white or pink subglobular berries, widely cultivated in the warm re- 
gions of both hemispheres, has become naturalized in the hammocks of 
Key West (Small, Manual, 1933; Everett, 1940). Wilson (1911) and 
some other authors have included this Specs in “G. pentaphylla (Retz.) 
DC.” or “G. pentaphylla (Retz.) Corréa.’ 

The present knowledge of the genus is very imperfect, and both taxonomy 
and nomenclature are complex. Tanaka has studied the genus critically 
for many years but has not yet published a monograph. 

The genus is of no economic importance. 


REFERENCES: 


See also under family references ENGLER (1931, pp. 316-318), PeNnzic (1887, 
pp. 194-209, pls. 19-21), SwIncLe (1943, pp. 153-158), and Wixson (1911, 
B25) 


EVERETT, T. H. Glycosmis citrifolia. Addisonia 21: 29, 30. pl. 687. 1940. 

LittLe, E. L., Jr. Notes on nomenclature of trees. Phytologia 2: 457-463. 
1946. [Glycosmis parviflora (Sims) Little, comb. nov., 463. 

NARAYANASWAMI, V. A revision of the Indo-Malayan species of Glycosmis 
Correa. Rec. Bot. Surv. India 14(2): 1-72. 1941. 

TANAKA, T. A note on Retzius’ Limonia pentaphylla. Bot. Not. 1928: 156-160. 
1928. [Records finding a presumed type specimen. | 


6. Triphasia Loureiro, Fl. Cochinch. 152. 1790. 


Evergreen shrubs with paired, or sometimes solitary, axillary spines. 
Leaves 3-foliolate [or simple], occasionally 1- or 2-foliolate; leaflets rela- 
tively small, subsessile, the terminal somewhat larger than the lateral, thick- 
ish, without evident reticulation, crenulate to crenate, glandular-dotted; 
petioles short, puberulous, not articulated with the leaf blade. Flowers 
1-1.6 cm. long, bisexual, 3[5]-merous, fragrant, solitary or in 2- or 3- 
flowered cymes in the leaf axils, pedicels short, minutely 2-bracteolate. 
Calyx cuplike, 3[5]-lobed, persistent. Petals 3[5], linear to lanceolate- 
oblong, imbricate. Stamens 6[10] in 2 series; filaments slender, broadened 
toward the base; anthers small, oblong. Disc ringlike to short-cylindric, en- 
circling the stipelike base of the ovary. Gynoecium 3[5]-carpellate, syn- 
carpous; stigma + capitate, 3[5]-lobed; style slender, deciduous; ovar 
ovoid to ellipsoid, narrowed toward the ends, 3[5]-locular, with a solitary 
ovule in each locule. Berry small, dull reddish orange or crimson, with 


4See also “Taxonomic and Nomenclatural Notes on Zanthoxylum and Glycosmis 


c 
(Rutaceae)” in the present issue of this Journal. 


14 JOURNAL OF THE ARNOLD ARBORETUM | VOL. XLIII 


glandular-dotted exocarp (peel), mucilaginous, pulpy flesh, and 1— —3|-5] 
seeds. Seeds ellipsoid to subglobular, lacking endosperm, with fleshy or 
leathery testa; embryo straight, with plano- convex, sometimes unequal 
cotyledons snd small radicle. (Including Echinocitrus Tanaka.) Typr 
species: 7’. Aurantiola Lour. ( = T. trifolia (Burm. f.) P. Wils.). (Name 
from Greek, ¢riphasios, threefold, triple, referring to the usually trifoliolate 
leaves and trimerous flowers of the type species.) — LIMEBERRY. 


A genus of three species, probably native in southeastern Asia, the East 
Indies, and the Philippines. Triphasia trifolia (Burm. f.) P. Wilson, com- 
mon limeberry, 2x = 18, 36, apparently indigenous to southeastern Asia 
and the East Indies, has been recorded as naturalized on the Coastal Plain 
from Florida to Texas. This species, much cultivated as an ornamental 
and hedge-plant in the warm regions of both hemispheres, is widely natural- 
ized in the tropics. Flowers with 4-merous perianth and gynoecium some- 
times occur in this species. 

Cross-pollination by insects seems to be the rule, spontaneous self- polli- 
nation being prevented by the position of the stigma which considerably 
overtops the anthers. Nucellar polyembryony has been recorded in Tri- 
phasia trifolia. A spontaneous autotetraploid form of this species, with 
leaves thicker and flowers larger than in the diploid, typical form, has been 
found. Fruits of 7. trifolia are edible. 


REFERENCES: 

See under family references especially SwincLE (1943, pp. 236- 240); also 
ENGLER (1931, p. 325), Sac (1935), URBAN (1883, pp. 399, 400, 403), 
and WILson (1911, p. 22 
ALINDADA, R. C., and . G. Gonzales. The graft affinity of Triphasia trifolia 

(Burm. f.) P. Wils. with five varieties of citrus. Philip. Agr. 40: 631, 632 
Boh] 6s 
SWINGLE, W. T. New varieties and new combinations in the genera Clausena, 
Oxanthera, and Triphasia of the orange subfamily Aurantioideae ur, 
. Acad. Sci. 30: 79-83. 1940. [T. trifolia var. tetraploidea Single: 


7. Poncirus Rafinesque, Sylva Tellur. 143. 1838. 


Shrubs or much-branched small trees with green twigs armed with 
stout, axillary thorns often flattened at base; foliage spurs with extremely 
short internodes developing from dormant buds just above the thorns on 
the previous year’s branches. Leaves deciduous, palmately 3- foliolate ; 
leaflets sessile, shallowly crenulate to serrate above the middle; petiole 
articulated with the blade, narrowly winged. Flowers fairly conspicuous, 
3—6 cm. in diameter, Biceaual: subsessile, solitary or in pairs on the preced- 
ing season’s twigs (just above the thorns). Sepals 4—7, usually 5, distinct, 
ovate, reece Petals 4—7, usually 5, spatulate to obovate, with claw- 
like bases, white, soon deciduous. Stamens 20-60; filaments free, un- 
equal in length, slender, broadened toward the base; anthers ovate to 


1962] BRIZICKY, GENERA OF RUTACEAE 15 


ovate-oblong in outline, gland tipped. Intrastaminal disc annular to shal- 
lowly cupular, hairy. Gynoecium 6—8(usually 7)-carpellate, syncarpous; 
stigma capitate; style short and stout; ovary subglobular, hairy, 6-8 
(usually 7)-locular; ovules 4—8 in 2 collateral rows on axillary placentae 
in each locule. Hesperidium subsessile, globular to pyriform, 3—5 cm. in 
diameter, dull lemon-colored, fragrant when ripe, finely and densely 
pubescent, many-seeded; peel (exocarp and mesocarp) soft, 5-10 mm. 
thick, with numerous oil cavities, rather rough; pulp (forming together 
with the inner walls of the locules the endocarp of fruits) consisting of 
elongate, cylindric-conical, slender-stalked vesicles (outgrowths of the 
inner surface of the tangential walls of the locules) filled with a very 
acid juice and droplets of acrid oil in the center, and with minute, lateral, 
irregularly branched appendages (Fic. 1g, h) which presumably secrete a 
viscous fluid. Seeds lacking endosperm, plump, ovoid, the testa leathery; 
embryos often several in a seed, differing much in size, with 2 equal or 
unequal cotyledons and a short radicle; germination hypogeous, the young 
seedlings at first with bractlike cataphylls, then intermediate forms that 
soon merge into normal 3-foliolate leaves. Type species: P. trifoliata 
(L.) Raf. (Citrus trifoliata L.). (From French, poncire, of obscure origin 
but applied earlier to a variety of citron with large, tuberculate fruits 
[C. Medica var. tuberosa Risso], perhaps also to other similar varieties.) 
— TRIFOLIATE ORANGE. 


The single species, Poncirus trifoliata, 2n = 18, 36, native to central 
and nothern China, has become + naturalized in our area on the Coastal 
Plain from Florida to Georgia and Texas. Poncirus shows only a few 
insignificant variations and seems to be the most stable species of the 
group of “citrus fruits” (including also Fortunella, Eremocitrus, Clymenia, 
Microcitrus, and Citrus) which have been cultivated for long periods by 
man (Swingle, 1943). 

The floral biology of the species does not seem different from that of 
Citrus. Both cross- and self-pollination seem to be almost equally effec- 
tive. Nucellar polyembryony, apparently of an induced type, has been 
proved. A small percentage of autotetraploids sometimes occurs among 
nucellar seedlings and those of open pollination. Poncirus hybridizes freely 
with species of Citrus and Fortunella, producing hybrids which usually 
are female-sterile but which occasionally produce some fertile pollen. Be- 
cause of competition with nucellar embryos in a seed (and perhaps from 
other causes) the sexual embryo, when produced, seldom reaches maturity 
in most of the Poncirus > Citrus hybrids. This leads to the development 
of maternal-type seedlings, seriously interfering with the normal segrega- 
tion and recombination necessary in plant breeding (Yarnell, 1942). 
Vigorous, variable trigeneric hybrids, some very complex, have also been 
produced. The citrangedin, involving Citrus, Fortunella and Poncirus, is 
extremely resistant to cold and is notable in that it shows clearly traces 
of all three genera, but is strikingly different from any species of those 
genera. 


16 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


Generic distinctions in the Aurantioideae are weak and are much in 
need of further critical study. Although genetic evidence shows that 
Citrus and Poncirus are closely related, the latter has a number of differ- 


SHOAL 


of pulp vesicle (after Penzig, 1887), x ca. 110; i, seed, X 3; j, k, four of nine 
m m a single seed (nucellar polyembryony), X 4—note mostly un- 
equal cotyledons and different sizes and shapes of embryos 


1962] BRIZICKY, GENERA OF RUTACEAE 17 


ential characters: deciduous 3-foliolate leaves; overwintering flower buds 
covered with bud scales; densely pubescent fruits; pulp vesicles with 
minute lateral appendages: pith with transverse plates of thick-walled 
cells; stomata of the green twigs situated at the bottom of deep, narrow 
pits; seedlings with spirally arranged cataphylls, which merge gradually 
into foliage leaves, as in Eremocitrus; and the presence in immature fruits 
of the glucoside ponciridin, analogous to hesperidin but not found in 
Citrus. (Cf. Swingle, 1943.) Since the generic problem is very complex, 
involving a number of other genera, current usage is followed here, and 
Poncirus and Citrus are maintained as distinct. 

Poncirus has been used more or less extensively in many citrus-pro- 
ducing regions of the world as a rootstock for cultivated citrus fruit trees. 
The somewhat dwarfing effect on Citrus scions is noteworthy. The species is 
commonly grown as an ornamental in Asia, southern and central Europe, 
and North America, and is sometimes used for hedges. The fruits have 
been used in medicine in China. The juice and peel of fruits sometimes 
are used after special treatment for flavoring in baking and confectionary. 


REFERENCES: 

See also under family references ENGLER (1931, pp. 332, 333), PENzIc (1887, 
pp. 132-149, pls. 11, 13, 14, ‘Aegle sepiaria DC.’), and SwincLe (1943, pp. 
366-373). 

BenTON, R. J., . Bowman, L. Fraser, and R. G. Kessy. The significance 
of a ee for citrus rootstock problems. Int. Hort. Congr. 
Rep. 13: 1235-1240. 1952.* 

Cuapot, H. Remarques sur la Se auon des pépins de Poncirus trifoliata 
(Rafin.). Fruits 10: 465-468. 1955.* 

Knorr, L. C. Re- -appraising citrus rootstocks, with particular reference to 
their susceptibility to virus diseases. 1. Trifoliate orange (Poncirus tri- 
foliata (L.) Raf.). Citrus Mag. 19(8): 12, 14, 15, 18, 22, 23. 1957.* 

MUKHERJEE, S. K., and J. W. Cameron. Tree size and Chromosome number 
in a trial of tetraploid trifoliate orange as a citrus rootstock. Proc. Am. 
Soc. Hort. Sci. 72: 267-272. 1958. 

SWINGLE, W. T. Poncirus Raf. In C. S. Sarcent, Pl. Wilsonianae 2: 149-151. 
1914. [Restoration of the genus. ] 

WeatTuers, L. G. The effect of host nutrition on the capac of exocortis 
in Poncirus trifoliata. (Abs.) Phytopathology 50: 87. 19 

Wirtrock, G. L. Poncirus trifoliata. Addisonia 15: 59, 60. a oe 1931. 


8. Citrus Linnaeus, Sp. Pl. 2: 782. 1753; Gen. Pl. ed. 5. 341. 1754. 


Glandular, aromatic shrubs or trees, usually with solitary, axillary 
thorns, the older branches often thornless. Leaves alternate, persistent, 
1-foliolate; leaflet subcoriaceous, glandular-dotted, entire or toothed; 
petiole usually winged and clearly jointed with the blade (except in C. 
Medica). Flowers bisexual, sometimes also unisexual by abortion of the 
gynoecium, usually relatively large, 2-5 cm. in diameter, often fragrant, 
solitary or in pairs in the leaf axils or in short, axillary, corymbiform 
cymes. Calyx shallowly cup-shaped, 4- or 5-lobed. Petals 4-8, usually 5, 


18 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


linear-oblong, thickish, white, pink, or purplish pink, glandular dotted, 
imbricate in bud. Stamens 20-60, polyadelphous [or distinct]; filaments 
linear-lanceolate, subulate upward, white, usually variously connate; an- 
thers oblong or somewhat sagittate. Disc annular to cushion-like, sup- 
porting the gynoecium. Gynoecium syncarpous; stigma -- capitate, some- 
times slightly lobed; style cylindrical, deciduous; ovary ellipsoidal to 
subglobular, 8— 18 (usually 10—14)-locular, with several, usually 4—8, ovules 
(arranged in 2 rows) on an axillary placenta in each locule. Hesperidium 
usually large to very large, ellipsoidal and often mammillate at apex or 
pyriform to subglobular, sometimes depressed at apex; pericarp differen- 
tiated into 3 layers: an outer, yellowish-green to orange, leathery exocarp 
(flavedo) dotted with very numerous oil glands, a middle, thick and 
spongy, white mesocarp (albedo), and an inner, membranaceous endocarp 
with juicy, stalked, fusiform, inner outgrowths. (pulp-vesicles) filling the 
locules (segments of the fruit) and forming the “pulp”; the thin, mem- 
branaceous partitions (radial walls) of the locules often loosely coherent 
and easily separated one from another as well as from the spongy, white 
central column (core) of the fruit. Seeds ellipsoidal to obovoid, plump or 
flattened, sometimes beaked at apex, usually a few in each locule (seg- 
ment), the testa leathery; endosperm lacking; embryos 1-several, green- 
ish to white, with fleshy, plano-convex, often unequal cotyledons and 
short radicle. Lecrorypre species: C. Medica L.; see P. Wilson, N. Am. 
Fl. 25: 221. 1911. (Classical Latin name, atiginally used for the wood of 
Tetraclinis articulata (Vahl.) Mast., tne ‘African sandarac tree, and per- 
haps other conifers, but transferred to the citron in about the first cen- 
tury of the Christian Era; etymology of the word obscure.) 


A polymorphic genus of an uncertain number of species (16-145), of 
southern and southeastern Asia and Malaysia. Several species with numer- 
ous cultivars are widely cultivated and often spontaneous in all warm 
regions of the world. Five species, Citrus Medica L., citron, 2n = 18; 
C. Limon (L.) Burm. f. (C. Limonum Risso), 2n = 18, 36; C. aurantii- 
folia (Christm.) Swingle, lime, 21 = 18, 19-21, 27; C. Aurantium L. 
(C. vulgaris Risso), sour or Seville orange, 2” = 18: and C. sinensis — 
Osbeck (C. Aurantium [var.| dulce Hayne), sweet orange, 2n = 
36, 45, all presumably natives of southern or southeastern Asia, a ee 
recorded as more or less naturalized in our area (primarily southern Florida 
and the Keys). Accurate and recent data, however, are scanty, and per- 
haps only C. Aurantium and C. Limon (‘rough lemon”) can with some 
certainty be regarded as extensively naturalized. Seedlings of all the 
above, as well as of C. paradisi Macf., grapefruit, 27‘= 18, 27, 36, are 
often found along roadsides, at the edges of woods, and in secondary woods 
in Florida. Further data as to the extent to which such seedlings persist 
and reproduce themselves are much needed. 

Bisexual flowers usually occur regularly in Citrus sinensis (also in C. 
aa (L.) Osbeck, pummelo, C. paradisi Macf., grapefruit, and C. 
reticulata Blanco, tangerine). Both bisexual and unisexual (staminate) 


1962 | BRIZICKY, GENERA OF RUTACEAE 19 


flowers are common in C. Medica, C. Limon, C. aurantiifolia, C. Auran- 
tium, and some cultivars of other species (polygamo-monoecious species 
and varieties). Cross-pollination and self-pollination (including pollina- 
tion between individual trees of a clone) seem equally effective in the for- 
mation of embryos and seeds, with some exceptions involving absence or 
defective development of pollen, or self-incompatibility. Self- or cross- 
incompatibility has been recorded in some horticultural varieties of dif- 
ferent species. Thrips, honeybees, bumblebees, and some other insects 
attracted by the conspicuous corollas, fragrance, and abundant nectar 
apparently are responsible for natural cross-pollination and for most self- 
pollination. 

Polyembryony is common in Citrus. Gametic (zygotic) polyembryony 

gamospermy) seems to be rare, but can arise either by embryonic fission 
(cleavage polyembryony) or sometimes by the development of two game- 
tophytes (embryo sacs) in the same ovule. Nucellar (somatic) embryony 
(apomixis, agamospermy) is widely distributed within the cultivated 
species, apparently lacking only in C. grandis. (Some cultivars of C. 
Limon and C. reticulata are also mainly monoembryonic.) The number 
of embryos per seed (1-18) varies with species and variety, and there is no 
general consistency within the species. Often only a single seedling or 
rarely more than two or three seedlings develop from a polyembryonic seed. 
Although nucellar embryos sometimes occur in a fertilized ovule alongside 
a zygotic embryo, only pollination (nct fertilization) seems to be necessary 
for their formation. The occurrence of autonomous nucellar embryony has 
not been positively demonstrated. Nucellar seedlings show increased vigor, 
larger leaves and fruits, and a Juxuriant development of spines (“‘rejuvena- 
tion” or “neophyosis”). Parthenocarpy regularly occurs in some culti- 
vars of C. sinensis (e.g., navel orange, Satsuma), C. aurantiifolia (e.g., 
‘Tahiti lime,’ a triploid), and some others. 

The species of Citrus generally are interfertile, and their hybrids are 
+ fertile. Only a few varieties may be cross-sterile, and many inter- and 
intraspecific hybrids are known. The F, progeny of such hybrids usually 
shows great variability, but less vigor, than nucellar offspring or hybrids 
from intergeneric crosses (e.g., with Poncirus and Fortunella). The great 
variabiltiy of Fy, of inter- and intraspecific hybrids is mainly explainable 
by parental heterozygosity, which is common. Bud mutants are frequent. 

Polyploidy is frequent, triploids (of gametic origin) and tetraploids 
(somatic autotetraploids) having been recorded in various species, e.g., 
C. aurantifolia and C. Aurantium. Penta- and hexaploids seem to be ex- 
tremely rare. Aneuploidy and an unbalanced chromosomal complement, 
2n = 28, have been found in a few hybrids. 

A mycorrhizal association with an endophytic fungus has been recorded 
for the genus, but the association seems to be facultative (perhaps + para- 
sitic), since the formation of root hairs has been observed in young seed- 
lings grown in artificial cultures (Girton, 1927, Hayward & Long, 1942), 
as well as under field conditions (Bartholomew & Reed, 1943). 

There is little unity of opinion on the generic delimitation of Citrus. 


20 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


Although Swingle, who is followed here, regarded Poncirus, Fortunella, 
Eremocitrus, and Microcitrus as distinct genera, some taxonomists include 
Poncirus in Citrus, and Burkill (1931) united all of these. Much greater 
differences exist in the concept of the species. Swingle (1943) recognized 
16, at least one, C. paradisi Macf., being considered a satellite (or doubt- 
ful) species, while Tanaka (1952) accepted 145 species. The difficulties 
in delimiting the species, subspecies, and varieties of Citrus seem to de- 
pend, for the most part, on the absence of sterility barriers between groups 
of related forms and their apparent heterozygosity; but the problem is 
further complicated by nucellar embryony, by rejuvenation by nucellar 
progeny of + senescent varieties long propagated asexually, and by the 
spontaneous production of autotetraploids 

In addition to morphological and anatomical characters, the chemical 
composition, especially the presence of certain glucosides in the fruits 
(e.g., hesperidin and eriodictyol glucoside in C. sinensis, those plus hes- 
peridin chalcone in C. Limon, hesperidin in C. Medica, aurantiamarin, 
naringin (?), and hesperidin in C. Aurantium), seems to be of importance 
in distinguishing the species. The presence or absence, as well as the num- 
ber, distribution, and character of the acrid oil droplets in the pulp vesicles 
of Citrus (and relatives) may be of some taxonomic significance. 

The diverse uses and application of citrus fruits in the food and bev- 
erage industries, in essential oil production and the perfume industry, 
and as sources of vitamin C and “vitamin P,” are well known. The peel 
(flavedo) of sour and sweet oranges and lemons or the volatile oil ex- 
tracted from the fresh peel are used in medicine as stimulants, aromatics, 
and flavoring agents. 


REFERENCES: 


The vast number of references has been reduced here primarily to those either 
of general interest or dealing specifically with the southeastern United States. 
For an extensive list of references see H. J. WEBBER and L. D. BATCHELOR, The 
Citrus Industry. Vol. I. History, botany, and breeding. 1943; Vol. II. The 
production of the crop. 1948. (Univ. Calif. Press, Berkeley & Los Angeles). 
Under family references see especially PENzic (1887, pp. 17-132. pls. 1-10) 
and SWINGLE (1943, pp. 386-445); also ENGLER (1931, pp. 333-346), TILLSoN 
& BAMForpD (1938), and WILSON (1911, pp. 221-224). 

Aut, S., & Musanis-Up-Din. Morphology of the spines of Citrus. (Abs.) Proc. 
Pakictan Sci. Conf. 9(3): 23. 1957.* 

Barn, J. M. Morphological, anatomical, and physiological changes in the de- 
veloping fruit of the Valencia orange, Citrus sinensis (L.) Osbeck. Austral. 
Jour. Bot. 6: 1-24. pls. 1-4. 1958. 

BANERJI, I. Morphological and cytological studies on Citrus grandis Osbeck. 
Phytomorphology 4: 390-396. 1954. 

BARTHOLOMEW, E. T., & H. S. Reep. General morphology, histology and physi- 
ology. Chap. VI, pp. 669- Oe The Citrus Industry. Vol. I. 1943. See 
WEBBER & BATCHELOR abov 

Bove, J., & G. Moret. La silice de tissus de Citrus. Revue Gén. Bot. 64: 
34-39, 1957. 

Burkitt, IJ. H. An enumeration of the species of Paramignya, Atlantia and 


1962 | BRIZICKY, GENERA OF RUTACEAE 21 


Citrus, found in Malaya. Gard. Bull. Straits Settl. 5: 212-223. 1931. 
[ Poncirus, ree Fortunella, and Microcitrus reduced to sections of 
Citrus, 217-223. 

Cuapot, H. Systématique des Citrus en relation avec leur composition chimique. 
Bull. Soc. Sci. Nat. Phys. Maroc 35: 69-73. 1956. 

Ex Azount, M. M., & S. H. CaMEron. Adventitious buds in Citrus. Proc. Am. 
Soc. Hort. Sci. 38: 363-368. 1941. 

Ensicn, M. R. Venation and senescence of polyembryonic Citrus plants. Am. 
Jour. Bot. 6: 311-329. 1919. 

FAUVEL-DECROMBECQUE, J. H. Les variations des Citrus et leurs conséquences 
botaniques, culturales, promologiques et commerciales. Thesis, 156 pp. Univ. 
Alger, Fac. Sci. Alger. Philippeville. 1944. 

FLORIDA DEPARTMENT OF AGRICULTURE. Citrus Industry of Florida. 256 pp. 
Tallahassee. 1955.* 

Frost, H. B. Genetics and breeding. Chap. IX, pp. 817-913. The Citrus In- 
dustry. Vol. I. 1943. See WepBer & BATCHELOR, above. 

. Seed reproduction: development of gametes and embryos. /bid. Chap. 
VIII, pp. 767-815. [See also Jour. Hered. 29: 423-432. 1938; Jour. Wash. 
Acad. Sci. 15: 1-3. 1925; Proc. Natl. Acad. Sci. 11: 535-537. 1925.] 

Furr, J. R., & P. C. REECE. Wentincaion of hybrid and nucellar citrus seed- 
lings by a modification of the rootstock color test. Proc. Am. Soc. Hort. 
Sci. 48: 141-146. 1946. 

. C. Cooper, & P. C. Reece. An investigation of as formation 
in ‘adult and juvenile citrus trees. Am. Jour. Bot. 34: 1-8. 1947. 

Furusato, K., Y. Outs, & K. IsHrpasui. Studies on Se in Citrus, 
Seiken Zihé 8: 40-48. 1957.* 

Girton, R. E. The growth of Citrus seedlings as infeed by environmental 
factors. Univ. Calif. Publ. Agr. Sci. 5: 83-11 

GurcEL, J. T. A., & J. SousiHe. Analysis of ene in Citrus, especially 
pummelos. (aka Portuguese.) Anais Esc. Super. Agr. “Luis de Oucion? 8: 
727-746. 1951.* [English summary. | 

Haywarp, H. E., & E. M. Lonc. The anatomy of the seedlings and roots of 
the velence orange. U. S. Dep. Agr. Tech. Bull. 786: 1-31. 1942. 

Hopcson, R. W., & S. H. Caos: Effects of reproduction by nucellar em- 
Reon on clonal characteristics in Citrus. Jour. Hered. 29: 417-419. 1938. 

Hu, C. C. A review of literature on Czfrus nucellar embryony, and some 
new data on old and young lines of Valencia orange. Mem. Coll. Agr. Natl. 
Taiwan Univ. 3: 20-33. 1953. 

Hume, H. H. Citrus fruits. Rev. ed. 444 pp. Macmillan, New York. 1957.* 

Knorr, L. C. The growing of lemons in Florida: historical, varietal, and cul- 
tural considerations. Proc. Fla. State Hort. Soc. 71: 123-128 

,R. F. Suit, & E. P. DucHarmMeE. Handbook of Citrus diseases in Flor- 
ida. Fla. Agr. Exp. Sta. Bull. 587: 1-157. 1957 

Kruc, C. A. Chromosome numbers in the subfamily Aurantioideae with special 

reference to the genus Citrus. Bot. Gaz. 104: 602-611. 1943. [Addendum 
y H. B. Frost, 609, 610. | 
O. Baccui. Triploid varieties of Citrus. Jour. Hered. 34: 277-283. 


Leroy, = F. La polyembryonie chez les Cztrus. Son intérét dans la culture et 
anielioration: Revue Bot. Appl. Agr. Trop. 27: 483-495. : 

LoncLey, A. E. Polycary, Holvepery, and polyploidy in Citrus and Citrus rela- 
tives. Jour. Wash. Acad. Sci. 15: 347-351. 1925. 


7 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


Loustav, J., A. OrtuNo, & O. Carpena. La estructura de los cloroplastidios 
de hojas de Citrus. I. Limonero. Anal, Edafol. Fisiol. Veg. 17: 473-485 
1958.* 

Mat ack, M. B. Observations on the red color of the blood orange. Pl. Phy- 
siol. a 729, 730. 1931. 

. The juice sac of the orange with some observations on the plastids of 
Citrus. Jour. Wash. Acad. Sci. 21: 437-440. 1931. 

Mrnessy, F. A., & C. A. ScHRoEDER. Pistil development in C7trus flowers. Bot. 
Gaz. 117: 343-347. 1956. 

NAITHANI, S. P., & S. S. RacHuvansnt, Cytogenetical studies in the genus 

Pus. Nature 181: 1406, 1407. 1958. 

Nauriva., J Self- incompatibility in pumelo (Citrus maxima Merr.). Curr. 
Sci. Bangalore 21: 347. 1952. 

Neusauer, H. F. Beobachtungen an Blaittern von Citrus maxima Merr. Osterr. 
Bot. Zeitschr. 106: 556-565. 1959. 

Reep, H. S., & T. Fremont, Etude physiologique de la cellule 4 my corhizes ie 

les racines de Citrus. Revue Cytol. Cytophys. Vég. 4: 327-348. 1935 
Factors that influence the formation and development of my ee 
associations in citrus roots. Phytopathology 25: 645-647. 1935. 

Russo, F., & M. Torrisi. Polyploidism in Citrus autopolyploids and allopoly- 
ploids. (In Italian.) Ann. Sper. Agr. II. 5: 1041-1062. 1951.* [English 


summary 
SHAMEL, A. D. Bud variation and bud selection, Chap. X, pp. 915-952. The 
Citrus Industry. 1943. See above 
SuimoMurRA, H. Pharmacognostical studies on the pericarp of Citrus and re- 
lated genera (1-4). (In Japanese, English summary.) Jour. Jap. Bot. 35: 
129-138, a 218, 282-288, 296-303. 1960. 
SWINGLE, W. T. The batawical name of the lime, Citrus aurantifolia. Jour. 
Wash. Acad. Sci. 3: 463-465. 1913. 
. New taxonomic technique in ap eae wild a - major crop 
plants age by Citrus. (Abs.) Am. Jour. Bot. 35: 1948, 
aes 


BINSON, & E. M. ree New Citrus a. Cire. U.S: 
Dep. Agr. 181: 119; 10 pls. 1931. 

TaNnaKA, T. Species problem in Citrus. A critical study of wild and cultivated 
units of Citrus, based upon field studies in their native homes. (Revisio 
Aurantiacearum IX.) 152 pp., pls. 1-3, chart 5. Tokyo 

A revision of Assam citrus. (Revisic Auranbacearan XI) Bull. Univ. 

Osaka B. 9: 29- oa 1959. 

& T. TANINAKA. A revision of Osmocitrus, a ae of the genus Citrus. 

(Revisio rene NUT.) Jbid. 10: 1960. 

TotKowsky, S. Hesperides. A history of the hie and use of citrus fruits. 
371 pp., 113 pls. London Sie 

VENNING, F. D. Cortical tracheids: a new vascular element from the orange 
sub-family oe Aurantioideae). Quart. Jour. Fla. Acad. Sci. 9: 
O 114. 1946 


ns on accessory vascularization in four species of Citrus and 
their bite application as new taxonomic characters. Jour. Wash. Acad. 
Sci. 37: 210-217. 1947. 
Wo tre, H = Some problems of Citrus nomenclature. Proc. Am. Soc. Hort. 
Sci. Cari ». Region 7: 18-21. 
YARNELL, 5S. a eaf segregation in Citrus-Poncirus hybrids. Proc. Am. Soc. 
Hort. Sci. 40: 259-263. 1942. 


1962] TOMLINSON, LEAF BASE IN PALMS 23 


THE LEAF BASE IN PALMS 
ITS MORPHOLOGY AND MECHANICAL BIOLOGY 


P. B. ToMLINSON 


IN PREPARING A SUMMARY ACCOUNT of the palms for the Anatomy of 
the Monocotyledons (Tomlinson, 1961la) the construction of the leaf base 
was described in a very cursory fashion, since little material had been 
available for detailed study. Also, this anatomical survey of palms had 
been carried out from the standpoint of systematic anatomy, and there 
seemed to be little information of taxonomic significance in the microscopic 
structures of the leaf base. More recently, having had access to abundant 
material from living palms, the problem has been reappraised. It has 
become evident that, although, from the point of view of systematic 
anatomy, the initial attitude may have been justified, the construction and 
behavior of the leaf base in palms is a complicated subject which is of 
considerable morphological interest. It is a problem which can only be 
understood by considering mechanical aspects of leaf development in rela- 
tion to growth of the stem. This subject is virtually unexplored, since 
the mechanical behavior of the palm leaf has no parallel elsewhere in the 
plant kingdom, and the construction and growth of palms is itself little 
understood. 

That there is considerable variety in the structure of the leaf base in 
palms is obvious at once when a large collection of living plants is studied. 
Some palms have smooth trunks because the leaves are early deciduous, 
each abscissing cleanly; other palms have a more dishevelled appearance 
because the leaf base persists on the stem, as fibers, shreds, spines, or solid 
stumps. The distribution of these visible characters is not random, for the 
appearance and behavior of the leaf base is very constant for each species. 
Sometimes a single type may occur throughout an entire genus or larger 
taxon. In this way the structure of the leaf base in palms affords good 
diagnostic features. However, in spite of the diagnostic usefulness of 
these features, they are rarely employed by taxonomists, or, if used, their 
description indicates a sad lack of understanding of the fundamental con- 
struction of the palm leaf on the part of the author.. There are some 
exceptions. The notes given by Wallace (1853), although brief, often in- 
clude sufficient information to allow the recognition of the type of leaf base 
which is described. Bailey (1941) considered characters of the stem sur- 
face to be a basis for the subgeneric division of Acrocomia; these char- 
acters are a result of differences in the behavior of the leaf base in the two 
proposed subgenera. 

On the other hand, it is not surprising that these features are poorly 


24 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


understood since many taxonomic descriptions of palms have been made 
from fragmentary specimens in which the attachment of the leaf base to 
the stem is not visible. In addition, there is no accurate account of the 
morphology and anatomy of the leaf base in palms. But this problem is 
only one of the many aspects of palm structure in which our knowledge 
is sadly deficient. The reason for these deficiences can best be appreciated 
by the tropical botanist; living palms must be studied, and it is difficult 
to obtain material of plants which are scarce or inaccessible or too valuable 
to horticulture to be used for science. The problems are on a scale larger 
than that envisaged by most plant morphologists, and special techniques 
must be devised, for it is difficult to apply orthodox anatomical and mor- 
phological procedures to buds which may be up to half a meter in diameter. 

The present account is intended as a general survey of the problem and 
fills in some of the larger gaps in our understanding without offering any 
comprehensive solution. The problem cannot be solved by casual examina- 
tion of unmolested palms. Buds must be carefully dissected and leaves of 
all ages examined because structures visible in mature leaves may only 
be of mechanical significance in early stages of development. Conversely, 
structures may be visible in the young leaf primordium only to disappear 
as the leaf reaches maturity. Thus, a ligule is far more commonly de- 
veloped by palms than an examination of their mature leaves would sug- 
gest. The ligule is present in the leaf primordium but shrivels, erodes 
away, or shreds as the leaf approaches maturity. 

The only published work on the mechanical features of leaf develop- 
ment in palms is that of Schoute (1915) who described the morphology, 
anatomy, and mechanical behavior of the leaf base in Hyphaene. Schoute’s 
work illustrates how the mechanical demands made on a cylindrical 
sheathing organ which encloses growing tissues are met by the development 
of vertical clefts or sutures which appear in precise regions as a result of 
anatomical modifications. The ultimate configuration of the leaf attach- 
ment is shown to be very efficient and to utilize available tissues in an 
economic manner. 

However, the leaf base in Hyphaene represents only one method by 
which the mechanical problems of the leaf base are overcome. Other 
methods involve either the shredding or partial decomposition of certain 
tissues of the leaf base, or the complete disappearance of certain parts, or 
sometimes the abscission of the whole organ. I have recognized a number 
of distinct biological types and these are described subsequently. Schoute 
himself was well aware that there was much to be added to his own 
account, for he wrote, “Il est cependant sir que le corps tout entier des 
palmiers n’a pas encore été étudié suffisamment sous ce point de vue et 
que l’examen attentif nous pourra révéler encore beaucoup de détails 
intéressants.”’ 

It is emphasized that this present article is only a preliminary essay 
and cannot hope to elucidate all the details. It deals with the problem 
entirely in a qualitative way and aims at providing information of use 
to the taxonomist who may wish to include notes on the diagnostic fea- 


1962 | TOMLINSON, LEAF BASE IN PALMS 25 


tures of the leaf base in his description of palms. Since problems of 
growth are involved, future detailed studies must be based on quantitative 
observations. 

As a preliminary aid, an account of the growth processes taking place 
within the leafy crown of a palm are given in order that the subsequent 
discussion of mechanical adaptations may be understood. These growth 
processes have been summarized elsewhere (Tomlinson, 1961a,b), so 
only a brief outline is presented here. 


THE DEVELOPMENT AND MORPHOLOGY 
OF THE LEAF BASE IN PALMS 


Leaves in a palm always originate within the terminal cluster. Most 
stages of leaf development take place within this leafy crown and are 
not visible without careful dissection of the bud. As in most monocotyle- 
dons, each leaf in the palm originates as a minute cowl-shaped primordium 
at the shoot apex. In its earliest stages, the primordium does not com- 
pletely encircle the stem, but its two margins soon grow rapidly around 
the shoot apex so that the base of the primordium is sheathing almost 
from the first. Eventually the base develops into a closed tube, the 
future leaf sheath. The distal part of the primordium elongates to form 
the petiole and rachis, the blade being segmented by peculiar growth 
processes in the distal part of the primordium (cf. Eames, 1953). Since 
differentiation of organs and tissues within the leaf occurs in a basipetal 
direction, the distal parts mature early and complete their growth and ex- 
pansion while the base is still meristematic. This basipetal order of matura- 
tion should be clearly recognized, since it is of considerable mechanical 
significance. In each successive leaf, at the time when the blade is fully 
differentiated and on the point of unfolding, the leaf base is still actively 
meristematic. Therefore, in its early stages of unfolding and assimila- 
tion the blade is borne upon a delicate leaf base in which cell division 
and cell expansion still continue and which is useless as a rigid sup- 
porting organ. Support for the open blades of the younger leaves is pro- 
vided by older, surrounding leaves with rigid, mature sheaths. 

Elongation of the parts of the leaf may take place neither at a uni- 
form rate nor in the sequence in which the parts mature. For example, 
elongation of the petiole is often very tardy, but rapid once it begins. 
In some cocoid palms a true petiole is absent since there is no naked 
portion of the leaf axis between the insertion of the lowest leaflets on the 
rachis and the mouth of the leaf sheath. In these palms a pseudopetiole 
is developed, because as the leaf matures the marginal tissues of the distal 
part of the leaf sheath disappear. This pseudopetiole may be quite long, 
as in Arikuryroba. In yet other cocoid palms, as in species of Orbignya, 
the petiole is absent. Other palms with no petiole, or with a very short 
petiole include species of Copernicia (e.g., C. torreana) and some of the 
scandent lepidocaryoid palms. 

The basal, sheathing part of the leaf in all palms is always tubular 


26 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


in its early stages of development (Fic. 1). The tube is closed and the 
leaf insertion completely encircles the stem. In many palms the initial 
tubular state of the leaf base can be revealed only by dissecting the bud, 
because, as the leaf matures, decay and dissolution of certain tissues re- 
duces its size considerably, and at maturity it may have only a narrow 
insertion. 

A ligule is a characteristic feature of a number of palms. It is a 
tubular extension (rarely the tube is open as in Rhapidophyllum and 
Zombia) of the distal part of the leaf sheath above the insertion of the 
petiole. It is most conspicuous and persistent in many scandent palms 
of the lepidocaryoid alliance in which it may have a specialized func- 
tion. Thus, in Korthalsia it is commonly occupied by ants. A ligule is 
also present in many, possibly all, palms of the bactroid and caryotoid 
groups, although it is visible only in young leaves, disorganizing as the 
leaf matures. In some arecoid palms, e.g., Ptyvchosperma, and some 
borassoid palms, e.g., Latania, there are small auriculate outgrowths on 
each side of the mouth of the sheath. These structures may be homologous 
with a true ligule. 

In order that subsequent descriptions may be clearly understood, it is 
necessary to define certain terms which are used consistently in later 
sections (Fics. 1-3). The abaxial surface of a tubular leaf sheath is its 
outer surface, the adaxial surface being the inner. In accordance with 
the terminology of Von Mohl (1824), which was also adopted by Schoute 
(1915), the part of the sheath on the side below the petiolar insertion is 
referred to as the dorsal side, the opposite side being the ventral side. 
This corresponds to the same use of these terms in descriptions of the 
carpel, itself a modified leaf. The tissues on each side of the dorsiventral 
plane may be described as lateral. 

Although the leaf sheath in all palms is fundamentally a closed tube, 
it is convenient to recognize two main types of leaf base, according to the 
behavior of the sheath as it ages. In the first group the leaf base may 
be described as obviously tubular since the base persists and matures as 
a visible long, closed tube (Fic. 2). Leaf sheaths of this type are always 
long in proportion to the total length of the leaf. Up to one quarter of 
the leaf axis may be sheath. This type of leaf base occurs predominantly 
in members of the arecoid, chamaedoroid, and iriartoid groups and is also 
characteristic of scandent palms in other groups. In the second category 
the leaf base is not obviously tubular (Fic. 3). Only in the early stages 
of development does the sheath have the form of a closed tube. As the 
leaf base grows certain tissues disorganize and the cylindrical sheath 
disappears, the leaf sheath at maturity being represented by a broad clasp- 
ing base which may or may not encircle the stem. Leaf bases of this 
type are always short and rarely ‘exceed more than one-eighth of the 
total length of the leaf. It should be emphasized that there is no sharp 
distinction between these two classes and that intermediate types of 
leaf base, difficult to categorize, are quite common. 

When the leaf base is examined at various stages of development it is 


1962 | TOMLINSON, LEAF BASE IN PALMS od. 


4 2 
ae 
en —— petiole 
| distal 
——— sheath 
A <—s 


ventral dorsal abaxial (outer) 


G J) surface i 


adaxial (inner) 
surface 


lateral 


Fics. 1-3. Types of a bases in palms. 1, Diagrammatic representation of 
young leaf primordium. 2, Mature leaf base of obviously tubular type, with 
representative cross eee on of sheath below. 3, Mature leaf base of not 
obviously tubular type, with representative cross section of sheath below. 


obvious that it can only be described as a uniform cylinder in a very 
approximate sense. The leaf sheath is always somewhat conical up- 


conical outline is a result of the basipetal sequence in maturation. The 
distal part of the sheath matures early, so its mouth is always narrow, 
enclosing younger leaves, the tissues of which are not yet bulky. Sub- 
sequent widening of the mouth of the sheath is to a large extent accom- 


28 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


plished as a result of continued growth and expansion of younger leaves 
which are forced through the mouth by their own basal meristematic 
growth. Whereas the mouth of the sheath is widened mechanically, the 
base widens through activity of its own meristematic tissues. In ligulate 
leaves the loss of the ligule is to a large extent caused by internal expan- 
sion of younger leaves. The ligule, being the most distal part of the 
leaf sheath, is always much narrower than the rest of the leaf base, and 
its disappearance from the leaves of larger palms is essential if younger 
organs which it encloses are to expand without restriction. In fact, the 
mechanical function of a tubular ligule may be to direct and support the 
pointed apex of primordia of younger leaves, maintaining them in a 
vertical position in such a way that they meet no resistance from older 
leaves. 

In the tubular type of leaf base, the walls of the sheath are fairly uni- 
form except for slight thickening on the dorsal side (Fic. 2), a thicken- 
ing which is most obvious in the distal region, close to the insertion of 
the petiole. There may also be slight constrictions in regions in which 
sutures subsequently appear. In the nontubular type of leaf base the dif- 
ferences between dorsal and ventral regions are more conspicuous (Fic. 
3); the dorsal region is always thick, woody, and persistent, whereas, the 
ventral region is thin, membranaceous, and often ephemeral. 


DEVELOPMENT OF THE LEAF BASE IN RELATION 
TO THAT OF THE STEM 


Growth processes in the leafy crown of a palm are not comparable to 
those of a woody dicotyledon (Tomlinson, 1961a,b). The apical meristem 
region is minute and is situated at the base of a shallow depression which 
terminates the axis of the palm. Leaves originate in the apical meristem 
proper, and, as they grow, each in turn comes to occupy the center of 
this depression. Each leaf is, however, displaced from this position to 
allow the development of younger leaves within, its insertion widening at 
the same rate as the stem enlarges. Thickening growth of the stem is 
not induced by the apical meristem itself but by a primary thickening 
meristem, a tissue which forms the surface of the apical depression. Each 
annular leaf insertion therefore continually increases in circumference as 
a result of the thickening of the axis, itself brought about by the primary 
thickening meristem. The rest of the leaf sheath also increases in cir- 
cumference at the same rate of enlargement as the leaf insertion. Initially 
this enlargement of the leaf sheath is effected entirely by cell division 
and cell expansion, but in the later stages cell division ceases. At the 
periphery of the apical depression thickening growth is no longer pre- 
dominant, lengthening growth of the axis occurs, and adjacent leaves 
become separated by internodal elongation. It is in this predominance 
of thickening over elongation growth in the early stages of stem develop- 
ment that palms differ so strikingly from most dicotyledons. 

The stem commonly continues to widen as elongation proceeds, but no 


1962 | TOMLINSON, LEAF BASE IN PALMS 29 


‘longer by activity of the primary thickening meristem. Instead, there is 
continued growth and expansion of the ground parenchyma. This diffuse 
secondary thickening (Tomlinson, 1961a, p. 20) may be long continued 
and obvious, as in Roystonea, the trunk tapering upwards (Fic. 14). 
Otherwise, secondary growth is either limited or not obvious, since it is 
restricted to that part of the stem which is still enclosed by persistent 
leaf bases so that the distal tapering of the stem is not visible without 
removal of the expanded leaves. However, it is this phase of stem growth 
which has such a significant effect on the behavior of the leaf base. 

It should be emphasized that, although growth of the palm stem can be 
envisaged as taking place in two distinct phases (an initial phase of 
thickening growth, followed by elongation growth), there is no marked 
disjunction between the two. Thickening growth continues well into the 
phase of elongation growth. In the smaller, canelike palms, and par- 
ticularly in the scandent palms, elongation growth begins early and may 
be much exaggerated. It should also be emphasized that a true under- 
standing of the growth processes at the stem apex in palms can only come 
from quantitative observations and not merely from a qualitative ap- 
proach employed in this article. Apart from figures quoted by Schoute 
(1915) there are no relevant published measurements. It is hoped that 
these qualitative notes can be amplified by measurements in future work. 

The general basipetal sequence of growth and differentiation has al- 
ready been emphasized. Since growth of the leaf and its associated stem 
internodes is closely correlated, this basipetal sequence is continuous from 
leaf to stem. Consequently, growth of the internode, both in length 
and thickness, continues after the leaf base has ceased to enlarge. The 
mature leaf base thus is seen as a rigid envelope enclosing growing tissues. 
Mechanical stresses are set up in the leaf base. Several types of stress are 
imposed and these are indicated below. 


MECHANICAL STRESSES IMPOSED UPON THE LEAF BASE 


Four main mechanical stresses are exerted upon the leaf base. These 
stresses are not of equal significance, and their magnitude differs in dif- 
ferent palms, largely as a consequence of the great variety in growth habit 
exhibited by palms. 

(a) Stresses due to growth of younger leaves. During the early phases 
of growth the leaf sheath is subjected to internal stresses arising from the 
expansion of younger, enclosed leaves. These stresses are not of equal 
significance throughout a single leaf base because the distal part of the 
leaf sheath matures before the base. Thus, the still meristematic base is 
capable of accommodating internal expansion by cell division and cell 
enlargement. The narrow mouth of the sheath is, however, usually 
widened passively under the internal stress. This widening may involve 
splitting or loss of tissues at the mouth of the sheath, changes which are 
most obvious in the ligule, when present. 

(b) Stresses due to expansion of the stem. The growing palm bud is to 


30 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


some extent in a state of dynamic equilibrium. Much of the stress set up 
by the growth and enlargement of the young stem internodes is resisted 
by the rigidity of the encircling, mature leaf bases. That these stresses 
are considerable is shown when the mature leaves around a palm bud are 
removed so that there is no restriction on internal expansion: the axis 
commonly splits vertically. The structural modifications described below 
are largely a result of changes induced by such internal expansive forces. 

(c) Stresses due to the weight of the leaf. Stresses are imposed upon 
the leaf base by the weight of its own rachis and blade. In view of the 
great weight of the leaves in the larger palms, these stresses are likely 
to be enormous. “A single, entire leaf is a load for a man” (Wallace, 
1853). Such forces can trap and kill a man. They are not constantly 
and uniformly distributed because the palm leaf moves in the win he 
distribution and moments of these forces have been indicated by Schoute 
(1915) for Hyphaene. Schoute considers that the mode of attachment 
of the leaf in Hyphaene is of maximum efficiency in catering to these 
forces. However, Schoute discusses the problem as that of a single leaf 
on the axis. This condition does not occur in palms, and much of the 
stability of the leafy crown must be a result of many leaves deriving 
mutual support as a consequence of their overlapping leaf bases being ar- 
ranged on a spiral with a high phyllotactic fraction. This probably ac- 
counts for the scarcity of a distichous leaf arrangement in palms and for 
the relatively short internodes of the larger palms with persistent leaves. 
Some of the stresses set up in the dorsal part of the leaf axis by the 
weight of the blade sometimes cs visible effects. The dorsal tissues may 
buckle under compression, as in the leaf base in Cocos, or a “geniculum” 
may be developed, as in many on palms. 

(d) Stresses due to the development of the inflorescence. The inflo- 
rescence in palms is usually axillary, and its enlargement places some 
strain on the leaf base. This strain may be minimized in several ways. 
In many of the arecoid palms, for example, the inflorescence does not 
expand until the leaf has abscissed (Fic. 15). Commonly the inflorescence 
elongates considerably, and grows nan the mouth of the leaf sheath 
before it expands. The inflorescence sometimes pierces the leaf sheath, 
or grows through an existing suture. In general, mechanical stresses 
imposed by the expansion of the inflorescence are less severe than those 
imposed during the normal processes of vegetative growth. 

Of these four types of mechanical stress, that imposed by the expansion 
of the stem is undoubtedly the most Sinica for the behavior of the 
leaf base. Thus, the measurements given by Schoute (1915) for Hy phaene 
indicate that at least a 30% increase in diameter of the leaf base occurs 
passively, after its growth has ceased. My own measurements on Sabal 
indicate that the diameter of the leaf base must at least double as a 
result of stem expansion. This increase is accommodated entirely by 
splitting of the leaf sheath. since the sheath widens neither by addition of 
new tissues nor by stretching of old ones. 

Anatomical observations indicate that the leaf sheath has only a limited 


1962 | TOMLINSON, LEAF BASE IN PALMS oF 


capacity for expansion by passive stretching. Initially the leaf base 
widens to accommodate internal enlargement by cell division and cell ex- 
pansion. After cell division ceases, tangential expansion of ground tissue 
cells continues, the cells often widening two- to threefold. This enlarge- 
ment involves the cells of the ground parenchyma only, since the tissues 
of the vascular bundles complete their differentiation early and are in- 
capable of passive expansion. Tangential expansion of ground tissue 
cells is most obvious close to the inner surface of the sheath. Here cells 
which were originally isodiametric may be widened up to tenfold in the 
tangential direction. 

The limit of this passive enlargement is soon reached, however, and 
subsequent widening of the leaf base takes place entirely by means of 
certain mechanical contrivances for which the leaf is adapted in various 
ways. In this way the rigid leaf base is able to persist on a stem which 
is still widening. The main biological types which I have recognized are 
described below. There is no clear limit to these categories, and different 
types intergrade. Some palms may show mechanical features belonging to 
more than one class. 


MORPHOLOGICAL TYPES OF LEAF BASES 


The following observations are entirely my own, made on living palms 
either in cultivation or in their natural habitat. In each of the classes 
which I have recognized, the bud of at least one example has been dis- 
sected carefully. A few preliminary remarks should be made concerning 
observations on leaf base morphology in palms. The larger palms have a 
long period of growth in which only juvenile leaf characters are seen 
(Tomlinson, 1960). These juvenile characters may persist longer in the 
leaf sheath than in other parts of the leaf, and a leaf with the adult char- 
acters of the blade may still have juvenile features in the sheath. In 
Mascarena, for example, the leaves at the base of the plant are long 
persistent and not deciduous like the more distal leaves. In the follow- 
ing notes juvenile characters are ignored. Observations on palms in cul- 
tivation can also be misleading, since such palms are commonly leaf 
pruned at regular intervals and not allowed to show features they would 
develop if left undisturbed. These abnormal aspects also have been 
ignored. 

The types of leaf base listed below are identified by the name of a 
genus in which typical features are clearly shown. These types are listed 
in two groups according to whether the leaf base is obviously tubular or 
not. 

Leaf sheath obviously tubular. Palms in this category can be sub- 
divided into two main types according to whether the leaf base is de- 
ciduous or not. However, these two types intergrade. 

CaLamus TyPE. Leaf base long-persistent, little modified with age. In 
palms of this type each leaf base is long, tubular, and wholly or partly 
covers the slender internode. Stem thickening is slight, and the leaf 


oe JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLUI 


base compensates for internal stresses entirely by division and expansion 
of cells. The leaf sheath is subjected to little mechanical strain and 
shows no mechanical adaptations. Each leaf base dies, its tissues dry and 
shrivel somewhat, but the leaf is not abscissed. It is only lost in extreme 
age, by gradual erosion and decay. Where a ligule is present it also may 
persist, as in most of the scandent palms. Otherwise it may decay, as in 
Bactris species. The only change brought about in such leaf bases by 
mechanical means is a widening of the mouth of the sheath early in de- 
velopment. 

This type of leaf base is found in the scandent palms which belong to 
several unrelated groups: lepidocaryoid palms (several genera), bactroid 
palms (Desmoncus), chamaedoroid palms (Chamaedorea spp.). It is also 
found in many of the smaller palms with erect, rigid but canelike stems as 
in species of Chamaedorea, Geonoma and Pinanga. In these last-men- 
tioned genera, however, there is a tendency for the leaf to absciss in the 
manner described below. 


VEITCHIA TYPE. Leaf deciduous, developing distinct separation layers. 
(Fic. 4). In this type the diameter of the stem continues to increase ap- 
preciably long after the tubular leaf sheath has achieved its maximum 
diameter through growth. The extent of this belated stem expansion 
may be considerable, and in Roystonea it may continue throughout most 
of the life of the palm (Schoute, 1912), although the trunk tapers ob- 
viously only close to the crownshaft (Fic. 14). The leaf sheath is not 
resistant to this expansion, but the whole leaf abscisses cleanly as a unit. 
This results from the development of separation layers in precise ab- 
scission regions. An annular separation layer at the node frees the leaf 
from the stem, but since the sheath is a closed tube, the leaf is not yet 
able to fall away from the stem. Complete release is effected by a vertical 
split on the ventral side through a narrow, but often conspicuous, separa- 
tion layer which extends the full length of the leaf sheath (Fics. 1 ae ee 
In this region the leaf may be slightly constricted; rarely the constriction 
is so deep as to form a conspicuous ventral groove (Fic. 15). These 
separation layers are both anatomically specialized and, at maturity, 
represent weak areas. Under the strains imposed from within, the leaf 
sheath splits from above downwards through the ventral separation layer 
(Fic. 12) abscission then being completed by cleavage at the node (Fic. 
13). The annular leaf scar is always clean and smooth, indicating that 
separation is predetermined and not merely a mechanical tearing and also 
that protective layers are developed before the leaf falls (Fic. 13). Ana- 
tomical aspects of this abscission require further study. 

The ventral line of abscission is always present, although it is more 
conspicuous in some species than in others. It is probable, however, that 
its conspicuousness is a good specific diagnostic feature. It is easily seen 
in Veitchia, for example (Fic. 12), but is not obvious in Roystonea. 
This ventral separation layer apparently has been overlooked in taxonomic 
literature. Schoute (1915) certainly missed it in Roystonea, which was 


1962 | TOMLINSON, LEAF BASE IN PALMS 33 


contrasted with Hyphaene as an example of a palm in which the leaf base 
was split in a haphazard manner. Admittedly this irregular tearing does 
occur in Roystonea, but usually only in young palms, and Schoute’s 
observations may well have been made on young Roystonea plants. In 
adult specimens cleavage is almost invariably regular. 

An additional morphological peculiarity, common in species with the 
Veitchia type of leaf base, is the presence of a small triangular notch at 
the node, the apex continuous upward into the ventral separation layer 
(Fic. 11). This structure must be formed early in leaf development and 
represents the region in the minute leaf primordium where the encircling 
halves of the future leaf sheath meet on the ventral side, touching but not 
fusing completely. This feature is probably a good specific diagnostic 
character, although it has apparently never been mentioned in taxonomic 
writing. For example, it is very conspicuous in some species of Veitchia, 
e.g., V. montgomeryana, but absent from others, e.g., V. joannis. Since 
the notch marks the exact ventral point of the leaf asecuon and persists 
as part of the leaf scar, it is a very useful tag by which the original 
phyllotaxis can be determined on old trunks. 

This type of leaf base, exemplified by Veitchia, is common throughout 
the arecoid, chamaedoroid and iriartoid groups. It is almost universally 
associated with an infrafoliar inflorescence. It does, however, intergrade 
with other types. In many small palms with canelike stems abscission is 
often imprecise and there is a transition to the Calamus type. In many 

palms of the arecoid alliance the inflorescence is either interfoliar or not 
consistently infrafoliar, and abscission layers are not well defined. There 
is thus a transition to other types of leaf base, and further study would 
probably reveal them as a new type. Heterospathe is an example of a 
palm the inflorescence of which is, with fair consistency, infrafoliar, but 
in which abscission of the leaf is irregular. This makes it difficult to cate- 
gorize. 

Leaf sheath not obviously tubular. This includes a number of 
distinct biological classes of which the type examples are quite clear 
cut. Some leaf bases may have characters of more than one type, how- 
ever, and in many genera it is not easy to categorize the type of leaf 
sheath. 

HypHAENE TYPE. Leaf base cleaving widely on the dorsal side. (Fic. 
5.) This type has been described by Schoute (1915), and the present 
summary of his observations adds little new information. In palms of this 
type the leaf base is at first shortly tubular. Allowance for the demands 
of stem expansion is made in two ways. Initially a ventral split (fente 
ventrale of Schoute) appears and its widening is accompanied by the loss 
of some ventral tissue. This split permits expansion of younger leaves 
and also causes the leaf as a whole to fall from a vertical position into 
a more pendant one. The second adaptation is more complex and 
accommodates the more extensive forces of stem thickening. These forces 
are exerted on the thick, woody dorsal tissues of the leaf base, and their 


34 JOURNAL OF THE ARNOLD ARBORETUM [| VOL, XLIII 


effects are indicated by Schoute’s measurements. They show that the 
circumference of the leaf base ceases to increase actively after it has 
reached a diameter of about 20 cm. Subsequent increase, up to a diameter 


4 VEITCHIA 


5 HYPHAENE 


(=> 
a 


lod: 


Tics. 4-6. Behavior of leaf base, illustrated diagrammatically, immature 
to right, mature stage to left, with representative cross sections of bot 
- : 


1962 | TOMLINSON, LEAF BASE IN PALMS 35 


of 38.5 cm. is largely due to the development of a dorsal cleft (fente 
dorsale) which widens at the same rate as the stem thickens and ultimately 
assumes a broad, rhombohedral shape (cf. Fic. 18). This cleft is no 
merely the result of mechanical cleavage. It is marked in the immature 
leaf by a slight constriction, the microscopic examination of which shows 
certain anatomical peculiarities. The cleft is mechanically weak so that 
separation occurs easily within it. In Hyphaene the margins of the cleft 
are clean, but in Sadal and some other palms with this type of leaf base 
there is some fraying of the margin and fibers frequently remain to con- 
nect the newly exposed surfaces. 

This type of leaf base is usually distinct and easily recognized, but 
transitional forms exist in some small palms. Thus, in T/vinax, the dorsal 
cleft is rather indistinct, and the rest of the leaf base shows additional 
mechanical adaptations. 

The overall result of the development of a dorsal cleft is the appearance 
of a broad diamond-shaped hole in the woody leaf base. Mature stems of 
palms with this type of leaf base are thus clothed with a crisscross pat- 
tern of overlapping leaf stumps, each “strand” in the pattern representing 
one half of a split leaf base. Schoute’s mechanical analysis of Hyphaenc 
indicates that the clefts do not weaken the leaf attachment since they 
appear in such a position that they offer mechanical advantages to the 
leaf insertion. In older leaves the distal part usually breaks away by a 
fracture of the petiole. Commonly the residual leaf base persists through- 
out the life of the palm, if undisturbed, as in Hyphaene. In Sabal, the leaf 
base itself soon falls to leave a fairly clean trunk. In Washingtonia, the 
whole leaf persists and the stem of an unmolested palm is a striking 
object, being clothed with a “skirt” of old leaves. In Corypha the extent 
of lateral movement of the leaf base is revealed by horizontal scratches, 
scored in the leaf base by the teeth on the margin of the petiole (Fic. 19). 

This type of leaf base occurs in larger palms of the borassoid and 
sabaloid groups. The reason for the consistent correlation between a 
Hvphaene-type leaf base and a palmate blade is unexplained. 


PHOENIX TvPE. Ventral tissues of leaf base eroding without becoming 
conspicuously fibrous. (Fic. 6.) The leaf base is initially shortly tub- 
ular, but resists stem expansion verv little. The ventral tissues disor- 
ganize, decay, and erode, and only the thickened, woody dorsal part of 
the leaf base persists (Fics. 16, 17). Otherwise, there are no specialized 
anatomical adaptations in this type. It is possible that expansion in the 
nodal region ruptures the ventral leaf traces so that the ventral tissues are 
deprived of food and water and die, subsequent decay being a passive proc- 
ess. The dead tissues show little tendency to persist as a fibrous material. 
The annular leaf scar, which is often rather indistinct, is the only indication 
of the original tubular state of the leaf base. The petiole fractures, and 
only the dorsal woody stump persists, sometimes throughout the life of 
the palm. Otherwise it is abscissed or decays gradually. The persistent 
dorsal part of the leaf base may occasionally show the effects of in- 


36 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


ternal forces of expansion, in numerous small vertical splits, as in the 
larger species of Copernicia (Fic. 17). In the larger species of Phoenix, 
e.g., P. canariensis, in which the internodes are congested, the woody 
overlapping de of the leaf bases form a complete vesture on the 
trunk (Fic. 16). This may have some protective value, like the bark of 
a hardwood tree. In other species of Phoenix, as in P. reclinata, the 
internodes are longer, and the woody base is less persistent. Copernicia 
cerifera is an interesting example of the Phoenix type in which the leaf 
bases are more persistent on the older, lower parts of the stem than on 
the younger, upper part. Hence, mature specimens of this palm have old 
woody leaf stumps at the base of the trunk, below the relatively smooth 
upper trunk 

This type is common throughout the palms in many unrelated genera. 
Its limits are not easy to define and it intergrades imperceptibly with 
other types, particularly with the two next described. In Phoenzx itself, 
for example, there is little tendency to form a persistent fibrous mass. 
In some of the cocoid palms which I have included in the Phoenix type, 
however, the ventral tissues tend to split into strips before they disappear. 
One feature which characterizes many cocoid palms is for a part of these 
strips to persist as blunt spines spaced at regular intervals along the margin 
of the leaf sheath, as, for exam oe in Arikuryroba, Butia, Elaeis, Jubaea, 
and Syagrus. The distal part of the petiole beats spines in Phoenie it- 
self, but these are obviously modified leaflets and not homologous with 
the spines in the cocoid palms, Another diagnostic feature of the cocoid 
palms is the absence of a true petiole, the apparent petiole being the 
persistent thickened dorsal region of the leaf sheath, although this may 
be quite long. In Afttalea, Orbignya, Scheelea, and Maximiliana there is 
neither petiole nor pseudopetiole. Only in Cocos itself is a naked petiole 
present. 


Cocos TypE. Ventral tissues of leaf base long persisting as a continuous 
fibrous network. (Fic. 7.) This class shows a complete intergradation 
with the previous class. The type example, Cocos, is, however, quite dis- 
tinct. The ventral tissue decomposes only partly, and its remains persist 
and serve an important mechanical function. There is some loss of tissue 
and splitting in the distal part of the leaf sheath. The splits generally 
separate the thin, ventral tissue from the thickened dorsal tissue, this 
being necessary for the correct mechanical behavior of the leaf base. The 
ventral tissues undergo decomposition, the soft, parenchymatous ground 
tissue decaying and being lost. The fibrous strands and fibrovascular 
bundles persist and form a material which resembles coarse sacking, not 
only in appearance but, to some extent, in texture (Fic. 20). This ma- 
terial owes its texture to the peculiar disposition of the vascular and 
fibrous bundles. Towards the distal end of the leaf sheath the bundles 


one with another, but form two or more systems (cf. Fics. 23, 24). 
Bundles within each system form parallel hiv st the different systems 


1962 | TOMLINSON, LEAF BASE IN PALMS 37 
are arranged so that they execute a shallow spiral, the spiral in adjacent 


systems running in opposite directions. There are frequent anastomoses 
and irregularities within this arrangement, but the general analysis de- 


MCQOEe@> 


8 TRACHYCARPUS 


Ai 
9 ZOMBIA C) ARI) 
a 


= 


C 
© ® 10 CARYOTA 


Fics. 7-10. Behavior of leaf base, illustrated diagrammatically as in Figs. 
4-6. 7, Cocos type; 8, Trachycarpus type; 9, Zombia type; 10, Caryota type. 


38 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


scribes the system quite well. With the decay of epidermal and ground 
tissues, only this fibrous network persists. Internal expansion pulls the 
tissue laterally, but the peculiar arrangement of this network allows the 
angular divergence of bundles in different systems to be increased, with- 
out affecting the structure or strength of the whole tissue. The tissue thus 
comes to resemble a coarse cloth, but only in a superficial way. There 
is no regular interweaving of warp and woof as in a man-made fabric; the 
palm sacking shows interweaving between fibers in different systems in 
an irregular manner and only sufficient to hold the adjacent systems of 
strands tightly together. 

Examination of this material shows that it can be stretched sideways, 
as the mechanical requirements of the palm bud demand, but it is also 
very elastic. Consequently, while offering no resistance to expansive 
forces, it functions as an elastic tissue, cushioning and damping the 
variable strains imposed upon the leaf base by the movement of the blade 
in the wind. A horizontal widening of this leaf tissue must be compensated 
by a vertical shortening. This probably accounts for the distal cleaving 
of the ventral tissues away from the rigid dorsal part of the leaf base, and 
this cleavage is probably entirely passive. The long, pendulous tongues of 
ventral tissue, each of which represents the remains of the distal part of 
the leaf sheath, are conspicuous objects in the leafy crown of a coconut 
(Fie. 20) 

It should be pointed out that the anatomical peculiarities in the dis- 
tribution of the strengthening tissues in the leaf sheath are not restricted 
to palms of the Cocos type, but that a similar arrangement is to be found 
throughout the whole family. This suggests that this construction has the 
same mechanical significance for all palms, but only in the Cocos type is 
the arrangement of major and obvious mechanical significance. Further 
detailed anatomical studies are needed to analyze this construction cor- 
rectly. 

The Cocos type can be recognized in many genera belonging to unre- 
lated groups, but often only in a modified form. Intergradation with other 
types is frequent. For example, the texture of the elastic tissue varies 
considerably. Thus in species of Coccothrinax the fibrous strands are 
quite woody and the tissue is very rigid (Fic, 24). In Livistona, on the 
other hand, the fibers are fine and the tissue delicate and almost w oolly. 
The leaf sheath of Livistona thus resembles the type of leaf base de- 
scribed as the next class. 

In palms of the Cocos type, the useful life of this elastic mechanical 
device is often short, as in Cocos itself. The ventral tissues are lost and 
the woody, dorsal stump soon abscisses to leave a fairly clean trunk. 
Otherwise, the leaf base may be long persistent, as in Paurotis and many 
species of Coccothrinax. 


ie 


— 


TRAcHYCARPUS TYPE. Leaf-base tissue mostly persisting as fibrous 
material, (Itc. 8.) This class represents only a modification of the 
previous type, and all intermediate stages of transition can be seen. In 


1962 | TOMLINSON, LEAF BASE IN PALMS Sie) 


Trachycarpus all the tissues of the leaf base, except the thickened dorsal 
region, persist as a fibrous mass which represents the fibrovascular system 
released by the decay of the ground parenchyma. This fibrous mass offers 
no resistance to internal expansion. The leaf bases are closely crowded 
on the trunk and persist, if undisturbed, throughout the life of the palm 
as a continuous fibrous mat out of which the dorsal stumps protrude 
(cf. Fic. 25). Trachycarpus itself is of some interest because it will grow 
at much higher latitudes than any other palm, even as far north as 
Scotland. One is tempted to correlate this cold tolerance with the shaggy 
coat, but it should be remembered that Trachycarpus is not a warm-blooded 
animal. Coccothrinax crinita has a similar shaggy coat but is not par- 
ticularly cold-resistant. 

Isolated examples of this class can be found in several unrelated 
genera, but it is often difficult to distinguish this type from the Cocos 
tvpe with which it intergrades. A transition to another type is also 
shown by Arenga species. For example. in 4d. pinnata (FiGs-25)- ube 
fibrous remains of the larger vascular bundles protrude prominently 
from the fibrous mass as long, pliable, but not sharply pointed, spines. 
as in the next class. 


ZomBia typE. Leaf base shredding partly or wholly into protective 
spines. (Fic, 9.) Examples of this class can be considered a modification 
of either the Cocos or the Trachycarpus type. The softer tissues of the 
ventral region decay but the fibrovascular bundles persist, either partly 
or wholly, as spines. These completely invest the stem and form a very 
effective armature (Fic. 23). In Zombia, the distal extremities of all 
those larger vascular bundles which have massive fibrous bundle sheaths 
end in a sharp point. These extremities become reflexed as the leaf 
matures and form the prominent stout spines. Anatomically the arrange- 
ment of the vascular bundles corresponds to the general pattern de- 
scribed for Cocos. hree systems of vascular bundles exist, but only 
the bundles of the outer system have massive fibrous tissues. Distally 
these extend beyond the insertion of the petiole into a long ligule which, 
unlike the ligule in most palms, is open on the dorsal side CHiGe 222). 
As the leaf base matures, the spaces between the vascular network are 
enlarged by internal expansive forces (cf. Fics. 22, 23), while the distal 
extremities of the future spines are freed by disappearance of the marginal 
tissues (Fic. 22). The reflexing of the spines is entirely a mechanical 
process, resulting from the effect of expansion on the peculiar mode of 
attachment of the ligular spines to the leaf sheath. 

In Rhapidophyllum hystrix the mature leaf base has the same biological 
function as in Zombia, but a few morphological differences are obvious. 
A ligule is present, open on the dorsal side, the free margins overlapping 
somewhat. The spines are the larger vascular bundles of the ligule freed 
by the breakdown of the surrounding tissues. These spines do not be- 
come reflexed. In mature palms, the spines protrude from a mat of 
fibrous tissue, which represents the remains of the network of smaller 


40 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


nonspinous vascular bundles. Arenga is very similar to Rhapidophyllum, 
although not of the same class because no rigid spines are developed. 
Trithrinax acanthocoma is a third example of this class, although I have 
not studied it in detail. However, the spines seem to originate as in 
Zomobia. 

It is hoped in the future to make detailed histological studies of the 
leaf base in this peculiar biological group. 


Caryora TyPE. Leaf base splitting ventrally, split surfaces connected 
by persistent fibers. (Ftc. 10.) This type is found only in the smaller, 
caespitose species of Caryota, but seems sufficiently distinct to merit 
separate description. The leaf base is initially long, tubular and ligulate, 
the walls of the sheath being much thicker than is usual in tubular 
sheaths. The ligule shreds into fibers and disappears as the leaf matures. 
A slight constriction on the ventral side is visible externally as a vertical 
separation layer. This region is anatomically specialized. As a result of 
internal expansion, the sheath splits along this line. The two margins 
of the suture do not separate cleanly, being connected by persistent 
fibers which are pulled into a horizontal position as the suture widens. 
These horizontal fibers interweave in a complex and irregular manner 
along the line of the original suture (Fic. 21). 

The leaf base in the Caryota type is quite long and more obviously 
tubular than in related palms. It is, therefore, intermediate between the 
tubular and nontubular type of leaf base. 


DISTRIBUTION OF LEAF TYPES IN THE 
MAJOR GROUPS OF PALMS 


The preceding account represents a summary of incomplete observa- 
tions. The main biological classes I have described are quite distinct, 
but others may exist, and the present ones may be subdivisible. In the 
succeeding paragraphs the distribution of these types throughout the major 
tribes of palms is indicated. The taxonomic groups are those employed in 
Tomlinson (1961a, Table 1), recognized partly on an anatomical basis. 
Only genera are listed, although it is evident that some genera include 
more than one leaf-base type. Where a record is based on personal 
observation, the name is followed by an exclamation mark. Otherwise 
the records are based on published descriptions and illustrations, and 
may not be accurate. The list is very provisional and incomplete. Many 
palm genera are omitted. 

Palms in which the stem is more or less subterranean are not considered, 
since these form a group which has not been studied in detail. Such palms 
commonly show no mechanical response to the little stem expansion which 
occurs. 


Arecoid palms 
(a) Inflorescence infrafoliar 
VEITCHIA TYPE: Acanthophoenix, Actinorhytis!, Archontophoenix!, Arecal, 


1962 | TOMLINSON, LEAF BASE IN PALMS 41 


Bentinckia!, Brassiophoenix!, Chrysalidocarpus spp.!, Clinostigma, Cyrto- 
stachys!, Dictyosperma!, Drymophloeus!, Euterpe spp.!, Geonoma spp.!, 
Gulubia, Hedyscepe, Hydriastele!, Iguanura, Linospadix, ree ees 
veitchia, Nephrosperma!, Normanbya!, Oncosperma!, Pina spp.! 
Ptychandra!, Ptychococcus!, Ptychoraphis!, Ptychosperma!, ee 
Rhopalostylis!, Roscheria spp., Roystonea!, Siphokentia!, Veitchia!, 


In the above genera the inflorescences are always infrafoliar and abscission 
of the leaf takes place unequivocably according to the Veztchia type. 


(b) Inflorescence either interfoliar or inconsistently infrafoliar 

PHOENIX TYPE: Manicaria. 

TRACHYCARPUS TYPE: Oenocarpus spp. 

TYPE OF LEAF BASE OBSCURE: The following palms require detailed study: 
Asterogyne, Balaka, Calyptrocalyx, Calyptronoma, Chrysalidocarpus spp., 
Dypsis, Geonoma spp., Heterospathe, Howeia, Hyospathe, Jessenia, Lino- 
spadix, Neodypsis, Neophloga, Oenocarpus spp.. Phloga, Pholidostachys, 
Pinanga spp., Prestoea, Reinhardtia, Roscheria spp., Stevensonia, Vershaf- 
feltia, Vonttra. 

In these genera the leaf base abscisses fairly early, but often in an irregular 
manner. The inflorescence may be frequently infrafoliar, although the leaf base 
differs somewhat from the Veitchia type, as in Heterospathe. In other ex- 
amples the inflorescence is consistently interfoliar, the leaf base is quite different 
from that of Veitchia and abscission is long delayed, as in Neodypsis. 


Bactroid palms 
CaLamus TYPE: Bactris spp.!, Desmoncus!, 
PHOENIX TYPE: Acrocomia!, Aiphanes!, Astrocaryum, Bactris spp., Guilielma!. 


Borassoid palms 
HypHaENe type: Bismarckia, Borassodendron!, Borassus!, Hyphaene!, 
Latania!, Lodoicea, Medemia 
It should be noted that this group of palms is the only one with the same 
type of leaf base in all species. 


Caryotoid palms 
Cocos TyPE: Wallichia spp.! 
TRACHYCARPUS TYPE: Arenga!, Wallichia spp.! 
CaRYOTA TYPE: Caryota spp.! 


Chamaedoroid palms 

CALAMUS TYPE: Chamaedorea spp. ! 

VEITCHIA TYPE: Chamaedorea spp.!, Mascarena!, Opsiandra!. 

In some Chamaedorea species and in Gaussia and Synechanthus, the in- 
florescence is commonly interfoliar, abscission is imprecise, and these palms 
are difficult to typify. They require further study 


42 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLIIL 
Cocoid palms 


PHOENIX TYPE: Arecastrum!, Arikuryroba!, Attalea!, Butia!, Corozo, Elaeis!, 
Jubaea!, Maximiliana!, Orbignya!l, Rhyticocos, Scheelea!, Syvagrus!. 
Cocos TYPE: Cocos!. 


It should be noted that, although most genera of this alliance have leaf bases 
of the Phoenix type, those of this type can be subdivided into two more or less 
distinct groups. Arikuryroba, Butia, Corozo, Elaeis, Jubaea, and Syagrus have 
a spiny margin to the pseudopetiole, whereas in Arecastrum, Attalea, Maxi- 
miliana, Orbignya and Scheelea the pseudopetiole is not spinous and is often 
absent 


Iriartoid palms 
VEITCHIA TYPE: Cafoblastus, Catostigma, Iriartea spp., Socratea, W ettinii- 
carpus, 
In /riartea species and Jriartella the inflorescence is commonly interfoliar, 


abscission of the leaf base may be imprecise and the palms are difficult to 
categorize. 


Lepidocaryoid palms 


CALAMUS TYPE: Avc Ls dass Calamus !, Calospatha, Ceratolobus, Cor- 
nera, Daemonorop: nospatha!, Korthalsia!, Lepidocaryum  spp., 
Mvyrtale pis !, Pletton omia! “Pleclgeona pus Schizospatha. 


PHOENIX TYPE: Coclococcus!, Mauritia spp. (e.g., M. setigera), Metroxylon!, 
Raphia spp.! 


TRACHYCARPUS TYPE: Mauritia spp. (e.g., MW. carana), Raphia spp. (R. tae- 
digera). 


Sabaloid palms 

HypHAENE TYPE: Corypha!, Cryosophila!, Nannorhops!, Sabal!, Thrinax 
spp.!, Washingtonia!l, 

PHOENIX TYPE: Copernicia!, Erythea spp.!, Pritchardia spp.! 

Cocos TYPE: Coccothrinax spp.!, Colpothrinax, Erythea spp., Licuala spp 
Livistona!, Paurotis!, Pritchardia spp.!, Rhapis!, Serenoa!. 

TRACHYCARPUS TYPE: Chamaerops!, Coccothrinax spp. (e.g., C. crinita!), 
Trachycarpus!. 


ZOMBIA TYPE: Rhapidophyllum!, Trithrinax 
Zomobia!. 


rn 


spp. (e.g., acanthocoma!), 


— 


Genera of uncertain taxonomic position 
VEITCHIA TYPE: Pse udophoe nix! 
Cocos Type: Leopoldinia spp. (e.g., L. pulchra), Pelagodoxa. 


TRACHYCARPUS TYPE: Leopoldinia spp. (e.g., L. piassaba). 


The leaf base in Ceroxylon, Orania, and Sclerosperma needs further study. 


1962 | TOMLINSON, LEAF BASE IN PALMS 43 


DISCUSSION 

In the present era of specialized and sophisticated botany with so much 
emphasis on experimental procedures, the plea for more research involving 
simple morphological observation is likely to go unheard. Descriptive 
botany is regarded as too unscientific and unprofitable a field for startling 
new discoveries. And yet, in the realm of tropical botany there is enormous 
scope for research which requires the simplest apparatus: an axe, a knife 
and a scalpel, a clear eye, and the patience and willingness to settle down 
and observe elemental things. The present article is based entirely on 
simple techniques of this caliber. The observations have been made on 
living palms, mostly in the field or outside the laboratory rather than 
within it. The palms are a tropical family, their morphology is im- 
perfectly understood because it differs from most other plants in many 
peculiar ways. It cannot be understood by regarding palms as aberrant 
trees. They are a unique but neglected group, and their study demands 
an outlook unclouded by a knowledge of how hardwood trees behave 
and preconceived notions as to how palms should behave. In so far as 
the growth of palms can be said to be known at all, its peculiarities raise 
problems more easily comprehended by an engineer than a botanist. 
The palms have discovered many devices by which these mechanical 
problems can be overcome. If we could ascribe reasoning powers to plants, 
pas one certainly be granted a high intelligence quotient, for their 

“ingenuity” soon becomes apparent to a thoughtful observer. The success 
and persistence of palms through a long geological history and their size 
as a taxonomic group and wide present-day distribution bear witness to 
their success in experimenting with original, and often apparently ill- 
adapted, growth forms. 

One of the most difficult tasks which may confront a morphologist is 
to recognize the existence of mechanical problems. The purpose of the 
present article has been to emphasize the question of mechanical be- 
havior of leaf insertion in palms, just as much as it has been to indicate 
ways in which the problem has been overcome. Only by further detailed 
studies using the more elaborate and precise techniques of modern 
botany can more decisive answers be given. 

There are many other aspects of palm morphology, in addition to the 
leaf base, which require study from a functional aspect. For example, 
the growth and expansion of the inflorescence enclosed by a leaf sheath 
needs sympathetic consideration. Some of the answers to these mechanical 
problems seem easy enough. The leaf may fall before the inflorescence 
expands. Usually the inflorescence grows through the open mouth of 
the sheath. Sometimes the inflorescence protrudes through a cavity 
developed for quite a different purpose, as it protrudes through the dorsal 
slit in the leaf sheath of Latania and Lodoicea. Commonly the sheath 
is pierced by the expanding inflorescence. But to what extent this break- 
through is facilitated by anatomical preadaptation, we do not know. Here 
is a profitable field for simple observation. The behavior of the spathe 


44 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


in palms is also a subject deserving careful study. Often the spathes 
are deciduous and conveniently fall so as not to hinder flowering. I know 
of no studies on the abscission of the spathe. The large woody spathe in 
cocoid palms deserves looking at with the eye of an engineer. One of 
its most common and striking features is the system of deep longitudinal 
grooves in its outer surface. They are of obvious mechanical significance 
when the life history of the spathe is considered. 

None of these studies, however elementary, can be accomplished swiftly. 
It takes time to become familiar with the biology of the living palm. 
Palms in botanic gardens are to a large extent inviolate, and the gift 
of a palm bud for dissection may come only at long intervals and then, 
quite literally, as a windfall. However skillful a research botanist may 
be in planning a budget for the expenditure of large sums on laboratory 
equipment, he is usually much less adept at acquiring funds for travel and 
collecting. But a travel fund is an important item of research expenditure 
for the tropical botanist. 

The study of some of these elementary problems in tropical biology 
may have wider implication than is at first apparent. Thus, the per- 
sistence or otherwise of the leaf base in palms may be a significant matter 
to the ecologist. The stems of palms with smooth, self-cleaning trunks 
are a common habitat for lichens and small bryophytes. Palms with 
rough, persistent leaf bases provide an important lodging place for many 
epiphytes — ferns, orchids, bromeliads, figs, and others. The shage 
fibrous coat of a palm stem is an excellent habitat for small animals — 
snakes, scorpions, frogs, beetles, ants, spiders, cockroaches, and a host of 
small insects. In dissecting the buds of palms the botanist should be 
prepared for constant zoological surprises. 


SUMMARY 


The problem of the leaf base in palms is introduced in this essay and 
solutions are offered only in a very general way. The leaf base in all 
palms is initially tubular, although it does not always mature as a closed 
tube. It completes tissue differentiation and matures as a rigid sheath 
long before the stem which it encloses has ceased its thickening growth 
The rigid sheath, especially in the larger palms, is thus subjected to 
internal stresses which it cannot accommodate by further active growth 
and expansion. Instead, this accommodation is effected by various 
mechanical devices for which the sheath of the palm leaf is often re- 
markably preadapted. Several distinct biological groups are recognized, 
although intermediate types are common. The leaf may absciss as a 
unit due to the development of separation layers in clearly prescribed 
regions. Otherwise, expansion of the sheath may be permitted through 
the development of wide clefts. Yet again, tissues may break down and 
disappear before they can offer resistance to stem expansion. In other 
types, these same tissues may persist either as an elastic tissue, or as a 


1962 | TOMLINSON, LEAF BASE IN PALMS 45 


fibrous mass, or sometimes even as protective spines, thus serving a 
secondary, mreidental, but important, function. 

In order to emphaciee the taxonomic value of these morphological 
characters, which are quite constant in each species, a provisional list of 
leaf base types throughout the family Palmae is included 

It is emphasized that there is much scope for this type of simple ob- 
servation, not only in palms, but in other groups of tropical plants. These 
observations are likely to have wide significance. 


BIBLIOGRAPHY 


BarLey, L. H. Acrocomia—preliminary paper. Gent. Herb. 4: 421-476. 1941. 

Eames, A. J. Diese morphology of the palm leaf. Phytomorphology 3: 
172-189. 19 

Mou, H. von. =. structura palmarum. /m: Martius, Historia naturalis pal- 
marum 1: I-LII. 1824. 

ScHoutTe, J. C. Uber das Dickenwachstum der Palmen. Ann. Jard. Bot. Buiten- 
zorg II. 11: 1-209. 1912. 

. Sur la fissure médiane de la gaine foliaire de quelques palmiers. Jbid. 
14: 57-82. 1915. 

ToMLInson, P. B. Seedling leaves in palms and their morphological significance. 
Jour. Nenole Arb. 41: 414-428. 

. Anatomy of the monocotyledons, ed. C. R. Metcalfe. Vol. 2. Palmae. 

Oxford. 196la. 

. Essays on the morphology of palms. VI. The palm stem. Principes 5: 
117-124. 1961b. 

WaALLaAcE, A. R. Palm trees of the Amazon and their uses. London. 1853. 


FAIRCHILD TROPICAL GARDEN, 
MIAMI, FLORIDA 


EXPLANATION OF PLATES 
PLATE I 


cs. 11-13. Leaf bases of the Veitchia type. 11, Ptychandra glauca, leaf 
insertion, ventral aspect, separating layers and notch distinc , Veitchia 
johannis, leaf insertion, ventral aspect, leaf sheath splitting from above down- 
ward along the ventral separation layer, notch not developed. 13, V. johannis, 
abscissing leaf, ventral aspect, the clean leaf scar indicating preformed pro- 
tective layers. 


PLATE II 


Fics. 14-17. Palms with the Veztchia and Phoenix types of leaf base. 14, 
Roystonea regia (Veitchia type), falling leaf abscissed as a unit — note distinct 
tapering of stem below crownshaft indicating future extent of internal diffuse 
secondary growth. 15, Mascarena vershaffeltii (Veitchia type), crownshaft 
from below, separation region of outermost leaf sheath visible as conspicuous 
ventral groove. 16, Phoenix canariensis, stem surface with woody armor of 


46 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


overlapping, gases partly encircling leaf bases. 17, Copernicia curbeloi 
(Phoenix type), vertical splits in leaf base near insertion reflecting extent of 
internal ee forces. 


PLATE III 


Fics. 18-21. Palms with the Hyphaene, Cocos, and Caryota types of leaf 
base. 18, Latania commersonit elle type), dorsal view of mature leaf 
base on young nonflowering specimen —=%in flowering specimens inflorescence 
commonly protrudes through large dorsal cleft. 19, Corypha elata (Hyphaene 
type), leaf bases showing horizontal scratches scored by marginal teeth of petiole. 
20, Cocos nucifera, showing persistent ventral tissues of leaf base visible as 
tongues of material resembling coarse sacking. 21, Caryota mitis, ventral aspect 
of mature leaf sheath with peculiar fibrous configuration. 


PLATE IV 


Fics. 22-25. Palms with leaf bases of the Zombia, Cocos, and Trachycarpus 
types. 22, Zombia antillarum, immature leaf base, with ligule (open on dorsal 
side) beginnin ing to separate into individual vascular bundles. 23, Zombia 
antillarum, stem covered with persistent spinous leaf bases, spines released == 
note separation of larger vascular bundles (as compared os = 22) indi- 
cating extent of expansion, 24, ee a esiag acuminata (Cocos type), showing 
woody and rigid remains of ental tissu 5, Arenga nen (Trachycarpus 
type), showing long, sa spines ae from ligular tissues and initially 
standing vertically er 


PLATE I 


oL. XLII 


7, 
] 


Jour. ARNOLD ARB. \ 


S 


M 


, LEAF BASE IN PAL 


N 


TOMLINSO 


Jour. ArnoLp Ars. VoL. XLIII PLaTE IT 


Jour. ARNOLD Ars. VoL, XLIII Puate III 


& 


TOMLINSON, LEAF BASE IN PALMS 


Jour. ARNOLD Ars. VoL. XLIII PLATE IV. 


ae 
< ahs 
1) 


Mae 


or A ea. 
BP ier ys' 


TOMLINSON, LEAF BASE IN PALMS 


1962 | “HOWARD, OBSERVATIONS ON REDONDA 51 


BOTANICAL AND OTHER OBSERVATIONS ON REDONDA, 
THE WEST INDIES 


RicHARD A. HowaArD 


THERE ARE MANY SMALL ISLANDS in the West Indies, but none is so 
intriguing as Redonda to the traveler in the Leeward Islands. Isolated and 
pinnacle-like, Redonda is a landmark to the sailors who pass it. Since 
deep water surrounds the rock, one can approach it closely, only to learn 
that a landing appears to be difficult, if not impossible. From the east, 
Redonda appears to live up to its descriptive name as a round island, and 
as the descriptions of Redonda have varied little from the original supplied 
by Columbus, who named it, one suspects that few of the recent writers 
have done more than reword an earlier description. At one time, the 
island of Redonda supported a mining operation and had about 130 peo- 
ple living on it, but for the last half a century it has been uninhabited. 
The island, however, is reported to have a “‘king” representing a regency 
now in the second dynasty. There are only meager botanical records from 
Redonda, and only one other biologist, an entomologist, has climbed to the 
crest. 

During several weeks of field work on Montserrat in January, 1961,1 
I saw Redonda daily in the distance. Finally, the temptation to visit it 
became overwhelming, and with the assistance of Mr. Kingsley Howes, my 
wife and I chartered the schooner “Melody” for a day and the trip to Re- 
donda. Departure before the break of dawn allowed a smooth and rapid 
passage to the island, where we landed, climbed to the peak, explored the 
shaft of the phosphate mine, and collected some samples of the vegetation. 
Descending in midafternoon, we set sail and broke out of the lee of the 
island with some difficulty, returning to Montserrat before sunset through 
rougher seas. The diversity of the vegetation, although mostly of weedy 
species, was unexpected. This alone compensated for the physical difficul- 
ties of the trip. It is not to be recommended to a tourist. 

Redonda lies about 25 miles southeast of Nevis, and 15 miles northwest 
of Montserrat. It is clearly visible from both of these larger islands. 
Redonda is recorded as being about 1.5 miles long and 0.3 miles wide, and 
the highest point is given by Martin-Kaye as 975 feet by aneroid reading 
(Reports on the geology of the Leeward and British Virgin Islands, p. 77. 
1959). The island is obviously volcanic in origin, presumably of the post- 
Pliocene period, and a part of the volcanic chain of the Leeward Island 
group including Saba, St. Eustatius, St. Kitts, Nevis, and Montserrat. It 
is a fragment of a volcano, however, for the western two-thirds have been 

‘The financial support of Grant No. G4441 from the National Science Foundation 
towards a Flora of the Lesser Antilles made this trip possible, and is therefore grate- 
fully acknowledged. 


52 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


lost in ocean depths, which reach 600 fathoms a short distance off shore. 
From the sea, the western face of the island is sheer, broken only by a 
mining scar at the northern end and one or two flume-like gullies farther 
south. In places, the vertical face appears to extend the full 975 feet. 
Various colored rocks and multiple layers of lava flows draw one’s eye. 
Sea-eroded cliffs 100 feet high surround the remainder of the island, mak- 
ing the initial assault a formidable feat, impossible except in one place. 
Clearly some cataclysmic event split Redonda in the past to leave the frag- 
ment present today. The local story that this split occurred in the 17th 
century and was observed by a passing sea captain cannot be checked. 
One can only wonder when the discrepancy developed between the early 
descriptions of the island and its present condition. 

Redonda was discovered and named by Columbus on November 11-12, 
1493, on his second voyage. Morrison, in Admiral of the Ocean Sea (p. 
410), relates that “from the northern end of Montserrat, Columbus sighted 
a large island to the northwestward but did not care to beat up to it against 
a trade wind. He named it Santa Maria la Antigua.” 

“Proceeding in a general northwesterly direction the fleet passed a small 
steep and rounded but inaccessible rock less than a mile long, which 
Columbus named Santa Maria la Redonda, St. Mary the Round. Redonda 
retains her name and her importance as a sea mark to this day; but she 
has never been worth inhabiting.” 

Other historians have about the same story. In the Life of Christopher 
Columbus, by his son Ferdinand (p. 125), it is reported that “from here he 
proceeded to Santa Maria la Redonda, to which he gave the name because 
it is so round and smooth that it is impossible to climb its sides without a 
ladder.” Markham (Columbus, p. 152, 1892) reports Redonda to be “a 
round islet [that] was seen to the westward, so steep on all its sides that it 
seemed inaccessible without stairs or ropes thrown from the top.” More 
recently, Sir Frederick Treves, in The Cradle of the Deep (1908), de- 
scribed Redonda (p. 196) as “a smooth pale fabric of stone rising out of 
the sea, like the dome of some immense submarine hall, whose span is a 
mile. It reaches to the height of 1,000 feet. It is bare as a pebble. .. .” 
Ober (A Guide to the West Indies, Bermuda and Panama, p. 343. 1920) 
says of Redonda, ‘It appears scarcely more than a rock pinnacle rising 
above the sea between Nevis and Montserrat, but it is a mile and a half in 
length by a mile in breadth, with an altitude of 1,000 feet. The Spaniards 
called it Redonda, or the Round Island, from its shape.” Thus, the dimen- 
sions suggested by the chroniclers of Columbus do not vary from those 
of the present. It is not clear whether Columbus, viewing the island in the 
late evening, passed to the east and did not notice the steep western face, 
or whether the island has truly split since the time of discovery. 

Although many naturalists passed the island of Redonda in the 17th, 
18th, and 19th centuries, apparently not one of them reached it. Only Sir 
Hans Sloane appears to have noted the island, and even he fails to com- 
ment on its shape in his published writing. G. R. de Beer (Sir Hans 
Sloane and the British Museum, 1953) refers to a letter from Sloane which 


1962 | HOWARD, OBSERVATIONS ON REDONDA 53 


is now in the British Museum. Sloane wrote of his trip north from Bar- 
in September, 1687, “Between Montserrat and Nieves lies a very 
sari Island called Redondo or Rotunda, discovered by Columbus in his 


AxpovE: Redonda as seen from the southeast. BreLow: Redonda from the 
northeast. The wave-cut cliffs and the cirques are from one to several hundred 
feet high. 


54 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLIII 


Second Voyage, who gave it the name Santa Maria Rotonda, from its 
Figure. It consists of one Rock, very Perpendicular and high, looking 
like a Pyramide, and if there were nothing but Rock, but I was inform’d 
by those who have been on it, that there is on its top an Acre or two of 
very good Ground, that it has a very good Landing-Place, and a Well of 
very good fresh Water. It has also a great store of Iquanas of black 
color. Many Boobies, and other Birds that come hither to lay their Eggs 
at proper Seasons.” 

Redonda was not visited by a naturalist until the Smithsonian-Bredin 
expeditions of 1956 and 1958 stopped at the island. Waldo Schmitt re- 
lated in the Smithsonian Report (457-8, pl. 6. 1957; 429-30. pls. 9, 10. 
1959) of his visit to the shore and of the ascent of Redonda by J. F. Gates 
Clarke, who climbed the ridge on the first visit, and on the second made 
the ascent and spent the night collecting insects, while defending himself 
from the rats which infest the rock. Dr. Clarke gathered a few plant 
specimens, but some of these were lost, and others so ruined in the descent 
as to be of little use, and so were discarded. Dr. Clarke’s photographs re- 
main, to the present, the only published photographs of the island and its 
summit. Several comparisons between his photographs and my own or 
between our observations will be made. 

Although currently uninhabited, Redonda has not always been so. At 
some unspecified period prior to 1860, ores containing phosphate were 
discovered on Redonda, and between 1865 and 1912 this material was 
mined. Shephard (Am. Jour. Sci. IT. 47: 428. 1869; 48: 96. 1870) de- 
scribed this mineral as Redondite, a name later shortened to Redonite, and 
currently considered to be variscite. It is included in a porphyritic or 
felsparphyric olivine basalt. In 1891, Hitchcock (Bull. Geol. Soc. Am. 2: 
7. 1891) reported an analysis to show concentrations of P.O; as high as 
42.9%, but a modern analysis of the residue remaining on Redonda yielded 
only 18% PsQs. 

Mr. Fred W. Morse visited Redonda in the company of Prof. Charles 
H. Hitchcock in the summer of 1890, and reported on the trip (Popular 
Science Monthly 46: 78-87. 1894). His colorful description of Redonda 
and the activities thereon is worth repeating. Redonda was reached after 
4 sail of three hours from Montserrat, Mr. Morse related. “As we ap- 
proached the pier, a boat manned by two negroes put off to meet us, with 
a strongly built man with a pleasant face and brown beard and dressed in 
white linen sitting in the stern. The man proved to be Captain H —, the 
superintendent of the mine, who welcomed us to Redonda and transferred 
us with our baggage to the shore. 

“The beach was only a few yards in width, and above us towered the 
cliffs, over five hundred feet high. Groups of men stood on their brink, 
looking down at us and appearing like silhouettes against the clear sky. 
The ascent to the plateau above... was accomplished upon an aerial 
tramway. 

“Two stout, heavy wire cables were stretched up the gorge and firmly 
anchored at both ends. Upon each cable ran a trolley, from which was 


Lert: Western escarpment of Redonda, showing remnants of the cable base and landing. An as- 
cent must be made in the right-hand (southern) gully. Ricur: View down the gully from the saddle. 
Plants of Cephalocereus and Melocactus are visible on the right. Notice the narrow coastal shelf and 
the deep water within a few yards of shore. 


[2961 


VONOdGHea NO SNOILVAYWASHO ‘CYVYMOH 


Sal 


56 JOURNAL OF THE ARNOLD ARBORETUM | VOL. XLII 


suspended a large iron bucket. To each trolley was attached the end of a 
light yet strong wire cable, which passed over a set of heavy pulleys at 
the top of the cliff, thus causing one bucket to ascend as the other de- 
scended. When passengers or freight were to be raised, the bucket at the 
top of the cliff was filled with water from a tank, and the lighter load at 
the bottom was quickly drawn up. The speed was regulated by means of 
brakes applied to the pulleys. 

“The main cables were eight hundred feet long and the load was raised 
to the height of five hundred and twenty-five feet above the beach. In 
places the wires ran at a height of sixty feet above the uneven surface of 
the gorge. 

“We were invited to get into the bucket which was at the foot. Captain 
H — stood on the edge, clinging to the trolley, and we rapidly glided up 
between the steep walls of the gorge, from whose rocky sides peered round 
cactus plants like heads of gnomes and several strange shrubs threw down 
aerial roots as though in a vain effort to reach the thin soil at the bottom. 
On gaining the landing at the top, we were. . . conducted up the slope 
a hundred yards to the superintendent’s house. The dwelling and office 
were really two separate buildings joined together by a wide veranda be- 
tween them and along their front. . . . the buildings had been framed and 
ready for putting together, and were small cottages with two rooms and 
with roofs of corrugated iron. We were met at the house by Mrs. H — 
and her young daughter, Dorothea, who, with the captain, were the sole 
white inhabitants of the island. A small black boy called Chalmers 
showed us to our room, where we prepared for dinner. By this time, the 
short twilight of the tropics had been succeeded by darkness, and when 
we returned to the dining room with its bright light we could hardly be- 
lieve that we were upon an almost inaccessible rock in the Caribbean Sea. 

“The next morning, just before daybreak, while yet dark as night in 
the room, we were awakened by the cries of the sea birds, which made their 
homes by the hundreds in crevices and niches of the cliffs. Very soon a 
bell rang in front of the house to awaken the workmen in the huts below 

s.” 

Later Morse visited the mine. “The path to the mine led us along the 
eastern slope of the island to the northern face of the main peak, where 
a wide and deep ravine separated us from the smaller peak. The distance 
from the house to the mine was about three-fourths of a mile. The path 
was very steep in places as it ascended towards the summit in order to 
avoid a deep gorge, and sometimes so narrow that a misstep would give 
one a bad fall down the slope.” 

Morse reported on the birds and lizards around the house, and com- 
mented that “several sheep and goats, two dogs, some hens, two peacocks, 
and a white cat comprised the domestic animals of this Crusoe-like home. 
From time to time the sheep and goats had become wild and had taken to 
the almost inaccessible parts of the cliffs and gorges, where it was exciting 
sport to pursue them. 

“After lunch, when the sun had begun to descend towards the west, 


HOWARD, OBSERVATIONS ON REDONDA 


1962 | 


SedIe JSIOUL UT punof a19M wWinj0 


‘SYIOI ay} Suoure 
[Sq PUe Dimodagag ‘apIs Ula}sea dy] UO UOTIeAI[a appr ay. Inoqe 


adoys 9219 LHOTY “Aqooq Bursar ayy aaoqe udas SI Sapl]s puke] puke wolsola Aq pewoy anbin y 
apeuutd yor ay} SuULMOYs epUopay Jo avy UIAISAM JO MITA :LETT 


‘puvyst ay} Jo jurod ysaysry ay} 


‘ 


58 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


Captain H — took us down to the plateau below the house to look at 
the quarters occupied by the workmen. The buildings consisted of two 
long sheds with close shutters instead of glass windows, and contained for 
furniture nothing but a tier of bunks or rather shelves, of rough boards 
along the walls... . Near the buildings were ovens where the baking 
was done... . Their fare was simple, consisting of bread and salt beef. 
The foremen and skilled workmen occupied two smaller houses, but lived 
in the same manner. Water for drinking was obtained by catching the 
rain on large inclined surfaces of corrugated iron, and collecting it in 
reservoirs. Such a reservoir was built at each end of the island for the 
use of the men, and the superintendent’s dwelling was provided with 
capacious tanks connected with the roof. 

Tempany (West Indian Bull. 15: 22- 25. 1915) visited Redonda in 
1915, after a disastrous hurricane on August 7, 1899, destroyed the build- 
ings Morse had described. Tempany’s descriptions, while less picturesque, 
give further details on the mining operations and on the living conditions 
of the workmen. Until the time of Tempany’s visit, phosphate rock was 
mined in four places on the island. The rebuilt dormitories now had space 
for 180 men, and the working conditions, including the rations of food 
and water, indicate that life for the laborers from Montserrat was any- 
thing but serene. Breakfast and lunch were served at the mine areas. 
After the cable at the northern end of the island weakened and collapsed, 
all ore was carried over the treacherous trail in basket loads of 84 pounds 
each on the heads of the men to the remaining cable near the manager’s 
house. 

Although the last shipment of ore was made in 1912, ore was stockpiled 
at both ends of the cable until July, 1914. It cannot be definitely estab- 
lished when the island was evacuated, but it is known that a skeleton crew 
was maintained on the island until the hurricane of 1929 blew away the 
remaining buildings, and the company’s lease was finally relinquished in 


Martin-Kaye, in describing the resources of the Leeward Islands, sug- 
gests that “the outlook for Redonda, however, is poor. The reserves are 
not known and to determine them would be expensive. Reports have it 
that the material was becoming more difficult to win. . . . There is the 
possibility that important resources remain, and this is supported to some 
measure by the length of time for which the defunct company held on to 
the lease. If they thought that the island held little further reserves the 
lease would presumably have been abandoned earlier.” 

The story of the “King of Redonda” is worth reporting, for it is well 
known and often repeated by the residents of the nearby islands. The 
details vary a bit with the teller, and two published versions are also 
known. Bradley Smith, in his excellent book Escape to the West Indies, 
gives a straightforward account obtained from Charlesworth Ross, re- 
cently commissioner on Montserrat. Apparently the story began in 1865, 
when Matthew Dowdy Shiel claimed the island of Redonda, and a few 
years later had his son, Matthew Phipps Shiel, later better known as the 


1962 | HOWARD, OBSERVATIONS ON REDONDA 59 


writer M. P. Shiel, proclaimed King Felipe the First. Smith reports that 
the British Colonial office eventually “tacitly admitted” his claim, although 
today the island of Redonda remains under the administration of the gov- 
ernment of Antigua. Eventually the younger Shiel passed his title to John 
Gawsworth through an ancient succession ceremony involving the mingling 
of blood through cuts on the wrist. After Shiel’s death in 1947, Gawsworth 
became King Juan the First, and has since appointed many prominent 
figures Dukes of Redonda. A more glamorous version of this tale appeared 
in a recent issue of a popular magazine (Men Only [London] 75: 58-60. 
1960) under the title of ‘King of all the Seagulls.” The coincidence of a 
disputed regency and a seemingly profitable mining operation makes one 
wish for more of the details of Shiel’s claim. 


THE VEGETATION 

A visit to Redonda is becoming increasingly more difficult. Storms of 
recent years have all but destroyed the loading pier, the remnants of which 
mark the landing spot. Once on the narrow, boulder-strewn “beach,” the 
only direction one can move is up. The ascent of Redonda is made along 
a cleft filled with volcanic gravel, which rises at a 60 degree angle. One 
climbs using all appendages, and the only resting place is the saddle, or 
ridge, fully 500 feet above the sea. In the afternoon, this valley assumes 
all of the characteristics of an oven, and, without a hint of a breeze on 
this, the leeward side, the rocks become too hot to touch. At the ridge, one 
finds a small level plateau on which are the foundations of former build- 
ings and the rusting machinery of the old phosphate mine. A grassy ap- 
pearing slope descends at an angle of 30 degrees to the east and the south. 
Broken water catchments, and the foundation of the manager’s house, are 
on the slope to the north, which is capped with a pinnacle of large boulders. 
A disintegrating foot path follows a circuitous route of varying altitude 
to a mine shaft, a short tunnel, at the northern end of the island. In all 
directions there is a breath-taking view of the northern Caribbean. 

Redonda supports a wild herd of nearly one hundred goats, some of the 
males massive and handsome specimens with beards nearly reaching the 
ground. Their hooves left prints the width of my hand. Myriads of sea 
birds nest on the island. During the peak of the laying season, the men 
from Montserrat visit the island and gather eighty dozen eggs a day for 
sale on adjacent islands. Rats persist on all parts of Redonda, and sleep 
is impossible along the shore or in the ruins of the buildings because of 
the attacks of these animals. Lizards and iguanas are commonly seen, but 
appear to be more agile than usual in avoiding capture. 

The early accounts of the plant life of Redonda are meagre. Morse, 
who visited the island with Charles Hitchcock, reports of their interest 
in the minerals, as well as the flora and the fauna. In addition to the 
“round cactus plants like heads of gnomes” and “several strange shrubs 
which threw down aerial roots” seen on the west wall of the island, he 
mentions the “red and yellow blossoms of the cactus’ around the man- 
ager’s house. He reports that “a few air plants, a species of Tvllandsia, 


60 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


Apove: The grassy slope of Redonda as seen across the plateau ( (saddle) and 
site of the former barracks and mining buildings. The ruins of the manager's 
house are seen toward the upper right. BeLow: A closer view of the rocks 
and vegetation of the sa me slope. Volcanic “bombs” are infrequent. Opuntia 


z 


repens, known locally as “su a is abundant. Agave, Croton, Lantana, and 
an 


W edelia species are domina 


1962 | HOWARD, OBSERVATIONS ON REDONDA 61 


clung to the projections of the rocks and formed almost the sole vegeta- 
tion at the extreme summit.” He was also impressed that ‘“‘one of the 
drawbacks to exploring the island was a variety of cactus which the work- 
men spoke of as ‘suckers.’ It resembles the prickly pear in form and had 
a yellow blossom. Its joints or sections were thickly covered with thorns 
or spines, which were from three-fourths of an inch to an inch and a half 
in length and barbed at the tip. The joints were easily broken off, and 
clung to anything upon which their spines could catch. The animals about 
the place were almost always seen with from one to a half a dozen of these 
‘suckers’ clinging to them. When a barbed spine became imbedded in the 
flesh it produced a sore unless removed at once, and it was usually neces- 
sary to cut it out in order to remove it.” 

Although Morse wrote, ‘The remainder of our stay was spent in col- 
lecting and preparing specimens of the phosphate, and also of the plants 
and animals,” no records of these collections can be located. 

Mr. C. Steffens (Globus 67: 49. 1895) reported that the island appeared 
treeless, but that near the peak were “Gold- und Silberfarne” and under 
other plants ‘‘Tillandsien.” 

In 1915, Trepany (doc. cit. 23) reported, “There is but little vegetation 
on the island, the surface being rock-strewn and barren, with no depth of 
soil. The principal vegetable forms which occur are species of Prickly 
pear (Opuntia) and Cacti, notably Cereus. It is, however, worthy of note 
that the silver fern (Gymnogramme trifoliata) and the gold fern (Gymno- 
gramme chrysophylia) both occur fairly abundantly in places in sheltered 
crevices in the rocks.” 

More recently, Martin-Kaye (loc. cit. 77) states that Redonda possesses 
“scant vegetation beyond some coarse grass and extensive networks of 
particularly pugnaceous varieties of prickly pear. . . .’ However, Clarke 
(Smithson. Rep. 430. 1959) reported, “The inclined plateau forming most 
of the top of the island is covered by coarse grasses, sedges, a slender nar- 
row-leafed agave, several cacti in great abundance, lantana and several 
other scrubby shrubs.” Plates. g and ro associated with Clarke’s report 
illustrate some of the plants I was able to collect. 

Although early observers may have described Redonda to be “as bare 
as a pebble,” plants exist on the island. Some ornamental or useful plants 
are remaining following cultivation. and a large number of “weeds” were 
obviously introduced. The typical strand plants of sandy beach and coral 
rock of the adjacent islands were missing; however, even the steep, wave- 
washed cliffs were abundantly populated with plants appearing to be 
perched in crevices or even on boulders soon to fall to the ocean below. 
The only level ground in the saddle, which formerly was occupied by min- 
ing buildings, had a wealth of grasses of wide-spread geographic distribu- 
tion. The eastern slopes of Redonda were simply piles of boulders, yet 
on and between the stones were many plants so abundant that the footing 
was even more treacherous, as the stones were obscured by the growth 
of Croton flavens, Lantana camara, and Wedelia calycina, the broad- 
leaved species present. 


62 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


By far the most memorable plants of Redonda are the “suckers,” 
Opuntia antillana, Opuntia repens, and Opuntia triacantha. No descrip- 
tion does justice to the audacity or tenacity of Opuntia repens. Truly its 
pads are delicately attached and its spines retrorsely barbed the full 
length. The slightest jar caused a fragmentation of the plant, and the 
pads penetrated with ease the canvas, leather, or heavy rubber footwear 
of various members of our party. On the rocky plateau, the plants were 
abundantly branched, extending in many directions, the segments stretch- 
ing pad on pad over rocks and shrubs. On the cliff faces, plants of the 
same species seemed to develop a single stem structure. The plants hung 
in long festoons from the rocks, and single plants appeared to be as much 
as twenty feet in length. A slight touch with a stick or a falling rock would 
send the pads as individuals cascading down the slope on the unsuspecting 
climber below. If my records of the vegetation are inadequate or in- 
complete, the blame rests on this species alone. 

Clarke reported that the only tree on Redonda was an introduced one, 
but failed to name it. Casuarina equisetifolia is clearly shown in his paper 
(pl. ro, fig. 2) near the ruins of the house of the manager of the mining 
company. In the vicinity are also introduced specimens of Bougainvillea 
spectabilis, its brilliant red flowers clearly visible from the schooner ap- 
proaching the island; Citrus aurantiifolia, the lime; and Annona squa- 
mosa, the sugar apple. Since Clarke’s visit in 1958, seedlings of Ficus 
citrifolia have been introduced by birds, for a nine-foot sapling is now 
flourishing from the apex of the central rock in Clarke’s photo of the 
“very tiptop of Redonda” (pl. 9, fig. 2). Other abundant plants, obviously 
residual from the period of mining operation, are Ricinus communis and 
Nicotiana tabacum. 

A spire-like cactus (Cephalocereus royvenii) grew in profusion on the 
steepest slopes of the western escarpment, but ati could the plants be 
approached with safety in order to make a collectio 

Although Clarke reported a cistern on top to ~ in “good condition,” 
during our visit we found that all of them were dry and so cracked as 
to appear scarcely capable of holding water. No rainfall records are avail- 
able, yet fortuitous showers must provide small pools or at least wet places 
deep in the piles of rock, and many of the scree slopes appeared damp and 
slippery. In damp, shady places on the eastern slopes, I was surprised to 
find several plants of Psilotum nudum, Pityrogramma chrysophylla, Pilea 
microphylla, and Peperomia simplex. Among the rocks, usually in slightly 
protected areas, were many plants of Tillandsia recurvata, but none was 
seen at the summit as Morse reported. 

The following species represent the most complete account, to the pres- 
ent, of the vegetation of Redonda. All species cited by number are sup- 
ported by vouchers deposited in the herbarium of the Arnold Arboretum. 
Regrettably, large collections could not be made, and several sight records 
are given for clearly recognized species which either could not be handled 
in my limited amount of collecting equipment, or just could not be reached 
over the edges of the precipices. 


1962] HOWARD, OBSERVATIONS ON REDONDA 63 


Lichenes 

The determinations of this group were made by Dr. I. M. Lamb, of the 
Farlow Herbarium, where the supporting specimens are deposited. The un- 
numbered collections were some of the more colorful lichen-masses and were 
not separated in the field from the small rocks on which they grew. 
Acarospora chrysops (s.n.) 
Buellia prospersa (s.n.) 
Calopaca sp. (s.1.) 
Heppia bolanderi (s.n.) 
Parmelia sp 
Ramalina ape pe rata (15227) — abundant in restricted areas of large boulders. 
Roccella babingtonii (15228) — abundant, pendant from the underside of large 

boulders. 


Psilotaceae 


Psilotum nudum (15220) —a single clump found in a wet crevice under a large 


Polypodiaceae 
Pityrogramma chrysophylla (15231) —the silver form alone was found in a 
gulley on the east slope, but smaller clumps of both the silver and gold forms 
were growing in cracks on the walls of the cistern above and to the west of 
the ruins of the manager’s house 


Gramineae 
Chloris inflata (15251) 
Digitaria sanguinalis (15249) 
Eragrostis ciliaris ares 15253) 
Panicum maximum (15256) 
Pappophorum pappiferum oe —a clump grass with conspicuous inflores- 


Paspalum laxum (15252) —a large clump grass, the largest and most common 


Setaria setosa (15250, 15257) 

Trichachne insularis (15254) 

Trichachne sp. (15234) —a distinct species which, unfortunately has no avail- 
able name. Specimens are known from a range extending from Mexico to 
Peru, and this collection must have been introduced as a weed. Dr. Jason 
Swallen, who supplied this information, will describe and name the species 
at a later date. 

Tricholaena repens —a colorful and easily recognizable grass, but only locally 
abundant. 


Cyperaceae 

Cyperus ligularis (15255) —a heavy clump sedge essentially limited to the 
higher elevations, the edges of the western precipice, and found occasionally 
on the sheer faces. Only rarely does the plant assume its normal shape, for 
the clumps are the favorite nesting places of the boobies (Sula spp.). The 
ae apparently keep the plants trimmed into a cushion shape, and, at the 
time of our visit, only a few of the lateral shoots bore inflorescences. (See 
Clarke's illustration loc. cit. pl. 10, fig. 1.) 

Cyperus sphacelatus (15230) 


64 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


Bromeliaceae 


Tillandsia recurvata (15221) — certainly not as abundant as Morse and Steffens 
implied. All plants were found growing on rocks at higher elevations. 


Agavaceae 


Agave montserratensis — although it was impractical to collect specimens of 
this large Agave, a complete set of photographs allows this determination. 
Trelease’s monographic treatment may need revision, but his recognition of 
ioe) species in the Leeward Island area seems sound. The Redonda speci- 

appears to be the same as the plants found on Montserrat. Clarke’s 
photograph (pl. 10, fig. 2) shows the characteristic habit and the long narrow 
leaves of this species which it is strange that earlier visitors did not notice. 

Agave a several plants of this species were seen near the ruins of the 
mine headquarters, suggesting that the plant may have been introduced as an 
ornamental or a possible crop plant. 


Casuarinaceae 
Casuarina ssid ts te (15233) —a single tree of this species was io near 
the use and is shown in Clarke’s illustration (pl. 10, fig. 2). The 
specimen 1s cichilate and no seedlings were to be found. The tree is a favorite 
nesting place for boobies, and all branches were heavily covered with guano. 


Piperaceae 
Peperomia simplex (15222) — Dr. T. G. Yuncker kindly identified this material 
and indicated that it is commonly known as P. hamiltoniana Miq., which is 


properly referred to P. simplex Ham. The present plants were abundant in 
damp areas under rocks and on the scree in gullies on the eastern side. 


Moraceae 
Ficus ee (15247) — only three plants, all seedlings, were seen on Redonda. 
he largest grew in a crevice of the top-most boulder of the island, a rock 
ne en guano. Clarke’s photograph of this same rock, taken in 1958, 
does not show this plant. The other specimens were in unapproachable loca- 
tions on the steep western face of the island. Clearly the species has been 
introduced recently by birds. 


Urticaceae 


Pilea microphylla (15219) — this species occurred on rocks and the damp scree 
of the eastern gullies. 


Amaranthaceae 
Amaranthus dubiuns (15245) —a weed near the old ae area. 
Centrostachya indica —a weed along the old path to the m 
Tresine angustifolia (15235) —a rampant herb among ees: iv the summit. 


Nyctaginaceae 
Boerhaavia coccinea — on the plateau and on the western scree slope. 
Bouganvillea aan —a woody vine planted and persisting near the ruins of 
the manager's hous 


1962 | HOWARD, OBSERVATIONS ON REDONDA 65 


Portulacaceae 
Portulaca oleracea — a weed around the ruins and in the gully one ascends. 
Portulaca halimoides (15218) — this, and the following two species, occur 
primarily on the scree slopes of the western face of the island. 
Talinum triangulare (15223) 
Trianthema portulacastrum 


Annonaceae 
ii squamosa —a small shrub persisting following planting near the man- 
ger’s house. 


Capparidaceae 
Cleome viscosa (15241) — a most common weed. 


Leguminosae 
Centrosema virginiana 
Galactia sp. (probably G. stricta) — occurring commonly near the ruins, but all 
of the plants seen were sterile. 
Tephrosia cinerea (15237) —an abundant herb found in many locations. 


Rutaceae 
Citrus aurantiifolia (lime) —a shrub persisting after cultivation near the man- 
ager’s house 


Euphorbiaceae 
Croton flavens (15240) — this species, Wedelia calycina and Lantana camara 
represent the most common broadleaved plants and dominant shrubs on the 
island. All of the plants, however, were of lower stature than the species 
ssumes on other islands. Many of the flat-topped shrubs served as nesting 
Bee for the birds, and these plants appeared to be trimmed around the 


ests. 
craton lobatus (15244) — common herb in the western gully along which one 
cends. 


Euphorbia hirta (15246) 
Euphorbia heterophylla 
Jatropha gossypifolia —a weed around the ruins and in the scree of the western 


Phyllanthus amarus — a weed around the ru 
Ricinus commun Lael iden persisting ae cultivation; common in gullies 
on the west face of the islan 


Malvaceae 
Sida cordifolia (15239) 


Cactaceae 
meas royenii — columnar cacti common on precipitous western escarp- 


eee intortus — found primarily on the sheer western face of the island. 
Opuntia antillana — a stout SAAN! of frequent occurrence. 

Opuntia repens — “the sucker”; extremely abundan 

Opuntia triacantha — less frequent than O. repens, and less easily fragmented. 


66 JOURNAL OF THE ARNOLD ARBORETUM [VOL, XLII 


Plumbaginaceae 
Plumbago scandens (15248) — abundant on the western escarpment. 


Apocynaceae 


Catharanthus roseus — occasional in mining area. 


Asclepiadaceae 
Cynanchum parviflorum (15225) — comm mon vine on the plateau. A leafless 
form of ihis species hung in festoons from many rocks on the western face. 


Verbenaceae 
elisa camara (15236) —an abundant shrubby apes: represented by the 
ange-yellow color form. All plants were spineles 
ne involucrata — found primarily at lower tees above the cliffs on 
the windward side. 
Stachytarpheta jamaicensis — infrequent. 
Labiatae 
Hyptis pectinata (15226, 15242) — frequent in occurrence, but usually browsed 


Leonotis nepetifolia 


Solanaceae 
Nicotiana tabacum — occasional plants presumably persisting after cultivation. 


Scrophulariaceae 
Capraria biflora — a weed near the mine ruins. 


Acanthaceae 
Justicia periplocifolia (15238) —a narrow-leaved form usually browsed into 
abnormal growth forms. Determined by E. C. Leonard. 


Rubiaceae 
Oldenlandia corymbosa — occasional among rocks on the eastern slopes. 


Compositae 
Ageratum deter sleet 
Emilia coccinea (15233A) 
Pterocaulon reali (15232) —a few individuals at the northern end of the 
island 


Wedelia calycina — one of the common shrubs on the island. 


After this article was set in type, I discov ne a eg > Harold Box entitled “A 
Note on the Vegetation of Redonda, B. W. dae . 77, 311-313. 1939. r. 
Box, sailing from Antigua, visited Redonda on rie i ate observations were 
made from the landing place, since he was unable to ce the cliff. A list of 27 species, 
including Talinum paniculatum and Lithophila muscoides which I did not encounter, 
is given as the lithophyte flora of the island. 


1962 | GRAY, REVISION OF PODOCARPUS, XIII 67 


A TAXONOMIC REVISION OF PODOCARPUS, XIII 
SECTION POLYPODIOPSIS IN THE SOUTH PACIFIC 


Netta E. Gray * 


SecTION PoLtypopiopsis was established in 1874 by Bertrand (1) for 
Podocarpus vitiensis Seemann, then known only from Fiji. In 1903, Pilger 
(23) included this species and P. minor Parlatore as species of doubtful 
affinity in sect. NaGetA, chiefly because of their opposite leaves. In 1926, 
he mentioned Bertrand section and species synonyms briefly but did 
not change his interpretation. Florin (8) recognized the significant dif- 
ferences of these species from others in sect. NAGEIA and restored the 
use of sect. Potypopropsis, to which he added Podocarpus rospigliosi 
Pilger, a South American species which had been described by then. This 
section now is firmly established in its use by Orr (22), Wasscher (32), 
and Buchholz & Gray (4). 

The five living species now included in sect. PoLypopiopsis are Podo- 
carpus vitiensis Seemann, of Fiji, New Guinea, New Ireland, and the 
Solomon Islands; P. filicifolius sp. nov., newly described from the Moluc- 
cas; P. comptonii Buchholz and P, minor Parlatore, endemic to New 
Caledonia; and P. rospigliosii Pilger, of the Andes of Venezuela, Colombia, 
and Peru. Two fossil species (from Tertiary deposits) referred to this 
section are P. araucoensis (Berry) Florin, found in Chile, and P. brownei 
Selling, lately described from Tasmania. 

This distribution is striking and becomes very significant in recent 
studies of regional floras (Smith 28), eaeee = (e.g., Van Steenis 30, 
Selling 26), conifer geography (e.g., Florin 9, Li 20), and phylogeny 
(Florin 14). An ancient vast southern eae land mass is empha- 
sized by Van Steenis (30) in his consideration of the angiosperm genus 
Nothofagus, which is not only found both in the South Pacific and South 
America, but which has a subsection of twenty-one species limited to New 
Caledonia and New Guinea. Some oi the land areas here included are 
southeastern Asia, Australia, New Guinea, New Caledonia, Fiji, and New 
Zealand. Since Podocarpus vitiensis was known first from Fiji, the section 
has tended to be associated mainly with those islands, but with the dis- 
covery of the closely related P. comptonii in New Caledonia and of P. 
filicifolius in the Moluccas, the New Guinea area of P. vitiensis becomes 


* The author wishes to express her appreciation to Dr. Rudolf Florin, of the 
Bergianska ie ee Stockholm, Sweden, for critically reading the introduction to 
this paper, and to Drs. Florin and J. Tengnér, for examinaticn and evaluation of the 
wood anatomy of two critical specimens of Podocarpus minor. She also thanks Dr. 

mith, of the Smithsonian Institution, for criticism of her Latin description of 
Bodvcurpus filicifolius. 


68 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLII 


of primary importance, with the Fijian specimens then representing an 
eastern extension of the species. There is a remarkably close similarity 
to the distribution of Nothofagus species. One would expect it to be 
possible to distinguish two separate species of Podocarpus from New 
Guinea and Fiji, but the specimens I have seen from New Guinea seem 
the same, and only ripe seeds are lacking from New Guinea. The close 
relationship of Fiji to these land areas, geologically and botanically, is 
shown especially well in the recent studies of the Fijian flora by Smith 
(28, 29). 

The postulation of a great south land mass, often called Gondwanaland, 
would connect not only the continental islands of the South Pacific, but 
also Antarctica, Australia, Asia, South America and Africa. 

The leaf anatomy of the species of sect. AFROCARPUS, endemic to 
Africa, shows the same twisting of the short petiole that is found in sect. 
PoLypoptopsis, even when the leaves are alternate instead of opposite. 
The species of both sections have amphistomatic leaves and the leaves 
lack accessory transfusion tissue; in the leaf epidermis of species of both 
sections, the subsidiary cells of the stomata show the Florin ring. The 
South American species, Podocarpus rospigliosii, is very like P. vitiensis, 
except that its branches are covered only with foliage leaves and we find 
scale leaves or bracts only on the fertile shoots; the seed usually is some- 
what larger. Other gymnosperm genera with species in both South America 
and Australasia are Araucaria, Dacrydium, and Austrocedrus. 

Florin (8, 11, 14) has concluded that Podocarpus is a southern genus 
which probably originated in the early Mesozoic, and we find fossil evi- 
dence in South America in Podocarpus araucoensis and in Tasmania in 
P. brownet, Florin’s new fossil genus (11) from Australia belongs to sect. 
STACHYCARPUS Of Podocarpus which has living species in South America, 
New Zealand, New Caledonia, and Australia. Buchholz (3) maintained 
the northern origin of all conifers. 

The most significant change in recent gymnosperm systematics has 
been the separation of the taxads from the podocarps, leaving the latter 
with the conifers (Florin 10a,b, 12, 13, 14). Pilger (23), in his classical 
monograph of the Taxaceae, included the podocarps with the taxads. 
There was increasing indication that these should not be kept so close 
together, and, in 1926, Pilger arranged the gymnosperms in seven fami- 
lies with the Taxaceae and Podocarpaceae separated. This treatment was 
followed by Buchholz (2) in 1946, in spite of the growing realization that 
the taxads should not be included with other conifer families. Florin (14) 
compared these two groups morphologically in almost every way in which 
they could be investigated to uphold the elevation of a class Taxineae. 
This class has been based chiefly upon the evidence of the development 
of the female strobilus as found in fossils (Florin 10a,b, 13, 14). Wilde 
(33) limited her lengthy discussion to comparisons of the male and female 
strobili of many species of Podocarpus; she included many data, however, 
which were useful in deriving the interpretations and solutions given by 
Florin. 


1962] GRAY, REVISION OF PODOCARPUS, XIII 69 


Only the two sections, Potypopiopsts and Nace, of Podocarpus 
regularly have opposite leaves; in sect. AFROCARPUS some specimens have 
all or only occasional branches with leaves opposite or subopposite. But 
all three of these sections show the peculiar twisting of the leaf bases and 
stem torsion In both sections Potypopropsis and NacetA, this may result 
in the leaves being spread in a single plane, as in the compound frond 
of a fern. This orientation has been described in detail by Florin (8), 
Orr (22), Gray & Buchholz (17), and Wasscher (32). Orr (22) recog- 
nized further similarities in sections Potypopiopsis and AFROCARPUS in 
the leaf anatomy: the leaves are amphistomatic, with more or less hypo- 
derm, transfusion tissue often extending more than half-way from the 
midrib to the margin of the leaf, no accessory transfusion tissue, and a 
single resin canal in all species except P. rospigliosi. The three resin 
canals below the vascular bundle, with additional ones in the blade of 
the leaf of P. rospigliosii, is an exception discussed by Gray & Buchholz 
(17) in relation to reports in the literature (e.g., Bertrand 1, Stiles 31, 
Mahlert 21) that P. vitiensis has more than a single vascular resin canal. 
The single resin canal which I have found in all transverse leaf sections 
of P. vitiensis is in agreement with the findings of Gibbs (16) and Orr 
(22) who emphasize this fact. Neither have I seen any accessory trans- 
fusion tissue in the mesophyll of the leaf blades such as is shown by 
Bertrand (1, Pl. 6, fig. 12). No mention has been made thus far that the 
notable difference in the leaves in sect. NAGEIA is in the many parallel veins 
extending the full length of the large leaf blades, in contrast to the single 
unbranched midrib of the species of sect. PoLYPODIOPSIS. 

Orr (22) used only three species in describing the leaf anatomy of sect. 
Potypopiopsis: Podocarpus vitiensis, P. rospigliosii, and P. minor. The 
external appearance of the foliage of the two former is quite similar, and to 
these we may now add P. filicifolius, which has foliage most like that of 
P. vitiensis. The external appearance of P. minor differs in not showing 
the pinnate arrangement of the leaves in a single plane except in the 
foliage of seedlings and on occasional lower branches; the branches are 
otherwise fully covered with crowded, opposite, decussate, ascending, oval 
or elliptic leaves. The discovery in New Caledonia of P. comptonii, with 
transitional foliage, affirms the judgment that P. minor really belongs to 
this section. Podocarpus comptonii is a large tree, recognized by Compton 
(5) and others. It generally has the foliage which has been described in 
such detail, although the reproductive branches lose the pinnate arrange- 
ment and are covered with decussate, opposite, oval, or elliptic leaves, as 
seen in P. minor. 

The only section of Podocarpus which has both bifacially flattened 
lanceolate leaves and scale leaves in the mature foliage is PoLyPopiopsis. 
The two kinds of leaves in sect. DacrycarPus are scale leaves and the 
needle-like leaves on the pinnate twigs are tetragonal in transverse sec- 
tion. This remarkable dimorphism shown by the foliage on the main 
shoots and leafy branches was described by Florin (8) and Wasscher 
(32) for Podocarpus vitiensis, but it is also evident in the other species. 


70 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


In his consideration of the external morphology of the coniferous leaves, 
De Laubenfels (6) described this section as having only Type II (bi- 
facially flattened) leaves. The scale leaves (De Laubenfels Type IIT) have 
been found only in connection with mature foliage. This combination is 
rare among other gymnosperms, in fact, having been found only in seed- 
lings of a Dacrydium; transitional foliage of Chamaecyparis, Neocalli- 
tropsis, and a Dacrydium; and mature foliage of Athrotaxis, Taxodium, 
and a group of Juniperus. Podocarpus vitiensis and P. filictfolius have 
scale leaves much as described by De Laubenfels, tapering sharply from 
the point of attachment and closely appressed to the stem in P. vitiensis. 
They are rarely crowded, except at the beginning of a growth period, but 
are usually spaced by internodes of about the same length as those between 
the foliage leaves. They are decurrent, and their arrangement is always 
decussate except at the one or two nodes immediately preceding distichous 
foliage leaves. Podocarpus comptonii and P. minor have some scale leaves 
of this kind but they become obtuse and ovate in shape, divaricate, often 
elliptic as they approach true leaves, even being abruptly narrowed at 
the base but not usually becoming more than 4 mm. in length. According 
to De Laubenfels, only in one group of Dacrydium do scale leaves follow 
juvenile leaves of his Type II, and, when scale leaves (Type III) are 
developed, they are never followed by any other leaf type. In sect. Poty- 
Poptopsis, however, I find that all variations are exhibited in the alterna- 
tion between scale leaves and foliage leaves. Main shoots may bear only 
scale leaves; a leafy branch which continues growth may first bear 
foliage leaves, then scale leaves. Leafy branches may have only one 
growth period with a dormant terminal bud or they may have at least 
one other burst of growth, as is evident in the leafy shoots where each 
growth period shows first an increase in leaf length to about the middle 
of the growth period, after which it again uniformly decreases; one to 
several pairs of scale leaves may separate the growth periods. The small- 
est of the leaves may be very similar to scale leaves and a pair of scale 
leaves is usually present in ?. comptonii and P. minor at the first node of 
a leafy branch. Scale leaves have been found only on the special repro- 
ductive branchlets of P. ros pigliosii. 

The leaf anatomy, as seen in transverse sections of the foliage leaves, 
was described by Orr (22) for Podocarpus vitiensis, P. minor, and P, 
rospigliosui. The leaves are amphistomatic, and palisade parenchyma may 
be found on both sides or developed only on the side facing the light. The 
single vascular bundle is flanked by wings of transfusion tissue which 
sometimes extend fully half-way to the margin of the leaf. The extent 
of the transfusion tissue varies too much from one leaf to another for 
it to be used as a diagnostic character, but it is greatest in P. minor and 
P. comptonti. There is no organized accessory transfusion tissue, and iso- 
lated lignified cells with large lumina were detected only very rarely in 
the mesophyll, with none at all present in P. rospigliosii. I agree with 
Orr that P. rospigliosii has the greatest number of hypodermal fibers with 
often a continuous layer at the margin and midrib. In P. vitiensis and 


pl 


— 


1962] GRAY, REVISION OF PODOCARPUS, XIII 71 


P. filicifolius the hypodermal fibers are few, small, and often isolated 
even at the margin and midrib. In P. minor and P. comptonii the hypo- 
dermal fibers are larger and scattered, or grouped together, with the 
fewest in P. minor, except for an almost continuous layer at the margin. 
Vascular fibers are usually large and abundant above the midrib in all 
the species but are absent or rare below the midrib in P. vitiensis, P. 
filicifolius, and P. rospigliosii. The leaves of these latter species are thin, 
usually only between 0.3-0.6 mm. thick, those of P. comptonii are a little 
thicker (especially on fruiting branches), being from 0.5—0.8 mm., while 
P. minor has very thick leaves, from 0.6—-1.2 mm. 

The female strobili are remarkably similar in sections PoLypopropsIs, 
NaceIA, and AFrocarpPus, even to the fertile bract adhering after the seed 
is separated from the axis at maturity. The detailed description of the 
ovule development of Podocarpus vitiensis by Gibbs (16) indicates a 
peduncle covered with imbricate scale leaves, succeeded by 6-10 bracts 
on the strobilus, the terminal one or two fertile. The portion of the pe- 
duncle covered by bracts might be designated as a woody receptacle, the 
uppermost 2—4 bracts subtending the ovule sometimes having definitely 
thickened adhering bases. This was not apparent to Gibbs in P. vitiensts, 
but that three sterile and one fertile bract unite to form a receptacle has 
been recognized in P. minor and P. comptonu. A thickened woody re- 
ceptacle is very apparent on the mature seeds of P. filicifolius, where an 
additional pair of bracts may become involved. 

In her observations on the wood structure of Podocarpus, Kaeiser (19) 
found that of species in sect. Porypoptopsis usually like that of those in 
sections AFROCARPUS and NAGEIA. 

In sect. Potypopiopsis, Hair & Beuzenberg (18) counted the chromo- 
somes of P. vitiensis, P. comptonii, and P. minor and found the 2m num- 
ber to be 20, If one considers ten to be the basic number, this count 
affirms the suggestion that this section and sect. NAGEIA, in which 27 = 2 
in P. blumei, are the oldest groups in the genus. 


Key To SPECIES OF SECT. POLYPODIOPSIS 


A. All foliage leaves arranged pinnately on the twigs. 
Bee Mature. seed Crested «ae, <n ten eee were are) at P. rospigliosit.’ 
B. Mature seed not crested. 
Ce Mature: seeds. pear shaped.¢ 25 yeeros ek tic Ish vitrensts. 
€.- Mature seeds, spherical: 2. is.4 oye eee rn ed x: 2. P. filicifolius. 
A. Some twigs pinnately leaved, but fertile twigs not flattened, and always 
bearing decussate foliage. 


D. Small trees or shrubs; foliage mostly decussate: twigs rarely flattened. 
ae 4, P. minor, 


‘This species treated previously in Jour. Arnold Arb. 29: 118-122. 1948. 


72 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


1. Podocarpus vitiensis Seemann, Bonplandia 10: 366. 1862, Jour. 
Bot. 1: 33. 1863, Fl. Vitiensis 266. 1865-73; Van Tieghem, Bull. 
Soc. Bot. Fr. 38: 169. 1891; Pilger, Pflanzenr. IV. 5(Heft 18): 63. 
1903, Nat. Pflanzenfam. ed. 2. 13: 245. 1926; Gibbs, Jour. Linn. 
Soc. Bot. 39: 182. 1909, Ann. Bot. 26: 533. 1912; Stiles, Ann. Bot. 
26: 455. 1912; Dallimore & Jackson, Handb. Conif. 58. 1923, 1931, 
85. 1948; Florin, Sv. Vet-akad. Handl. III. 10: 275. 1931; Wasscher, 
Blumea 3: 425, 1941. 


A large forest tree up to 43 m. high with trunk often over 1 m. in 
diameter (sometimes buttressed in New Guinea) and a crown of spread- 
ing branches. Young twigs decussate, usually in pairs at distances of 
2.5—4 cm., spreading or erect-spreading, dimorphic; main twigs terete 
or slightly flattened below the ramifications, leafless, alternately bearing 
axillary decussate lateral leafy twigs and decussate, ovate to orbicular, 
deciduous bracts nearly 2 mm. long; lateral leafy twigs slender, usually 
unbranched, with the leaves usually smaller toward each end, decussate 
but turned in one plane, pinnately arranged, divaricate, the twigs up to 
38 cm. long and terminated by small buds which may not develop. Ter- 
minal buds of main twigs globose or ovate, with decussate, ovate, orbicular, 
or obovate, obtuse scales, 1.5-2 mm. long. Leaves spread out in one plane 
by twisting at the base so that the adaxial sides are exposed on one side 
of the twigs and the abaxial sides exposed on the opposite side; leaf pairs 
5-8 mm. apart; leaves sessile, lanceolate, rounded at the base and broadly 
decurrent, gradually narrowing toward the narrowly obtuse apices, am- 
phistomatic, 1.5—3 cm. long by 3—5.5 mm. broad; midrib sometimes promi- 
nent on either surface and with stomata in the epidermis above and below. 
Leaves in transverse section with scattered hypodermal fibers on both 
sides, palisade mesophyll on upper, lower, or both sides, depending on 
the orientation of the leaf; a single median vascular bundle with a single 
resin canal (never 3) abaxial to the phloem and flanked by wings of trans- 
fusion tissue; vascular fibers conspicuous above the vascular bundle and 
no accessory transfusion tissue. Male strobili solitary or clustered 2 or 3, 
terminal on axillary leafy twigs or bracted pedicels, or on bracted main 
shoots often branching once or twice so that strobili or fertile branches 
may arise in bract or leaf axils, strobilus cylindrical, 12-20 mm. long and 
2—2.5 mm. in diameter; microsporophylls triangular, with acute or obtuse 
tips, margins scarious. Female strobili terminal on peduncles in axils of 
bracts or normal leaves; peduncles 2-8 mm. long, sometimes branched, 
covered with 6-10 pairs of imbricate scale leaves; strobilus of 6-10 bracts, 
1 or 2 fertile; receptacle not differentiated. Seed bluish red when ripe, 
to 2 cm. long, obliquely attached to receptacle, obovate, narrowed towards 
the base which may still bear the attached bract when the seed has fallen, 
apex obtuse; hard-coated inner seed terminated at the micropylar end 
by a sharp, sometimes recurved point. 


DistripuTion: In dense mixed forest, Fiji Islands, at altitudes between 
100 and 900 m.; New Guinea, in the western part at altitudes of 1100-1200 


1962 | GRAY, REVISION OF PODOCARPUS, XIII (es 


m. and in the southeastern part at 1650-2000 m.; and Vanikoru, Solomon 
Islands. 


Fiji Islands. Vir1 Levu: Mba, Nandarivatu, Degener 14483 es ISH, K, 
us),” 14485 (mo), 14496 (A, Ny), Mead, Sing. Field No. 1974 (x), Parks Bae 
(BIsH, tuc), Gibbs 674 (BM, K); valley of Singatoka River, Gillespie 3273.2 
(BISH), 3712 (BISH, DS, tUC), 3865 (BISH, K, NY, UC, us), 4308.1, 4402.2 
(BIsH); Tailevu, e. of Wainimbula River, Swih 7076 (us). VANUA LEVU: 
Thakaundrove, Yanawei River, Mt. Kasi, Smith 1796 (BISH, GH, NY, tUC, US). 
WirHovut Locatity: Horne 531 (GH, K), Seemann 576 (Holotype, K; GH), 
Graff (K). New Guinea. NETHERLANDS New GuINEA: Idenburg River, Ber- 
nard Camp, Brass 19534 (+a), 12787 (+a), 12787a (A); Cyclops Mts., Ver- 
steegh BW913 (Laz, +L); Papua: Alola, Carr 14160 (A, +Ny); Lala River, 
Carr 15666 (+A, BM). Solomon Islands. Santa Cruz Group, Vanikoru, near Lemon 
River, Walker B.S.J.P.212 (+a). Cultivated. AUSTRALIA: Bot. Gard. Sydney, 
Boorman, in 1908 (a), Eames, in 1937 (cu 


Even though Podocarpus vitiensis has been recognized as a species since 
1862, an understanding of its relationships within the genus is quite recent. 
On the basis of the single vascular bundle in the leaf, stomata on both 
sides of the leaves, and supposedly two lateral resin canals as well as the 
one below the vascular bundle, Bertrand (1) put this species in a separate 
section, Potypopropsis. The confusion brought about by Bertrand and 
Mahlert (21) in ascribing the three vascular resin canals has been dis- 
cussed previously (17) and referred to P. rospigliosii from South America 
which is now included in the same section. 

Subsequently, Pilger (23) put Podocarpus vitiensis in his sect. NAGEIA 
on the basis of the opposite leaves which show the same characteristic 
turning of the leaf bases so that the lower surface is uppermost on one 
side of the twig and the upper surface on the other side of the twig when 
these decussate leaves orient themselves in the pinnate “fronds.” Pilger’s 
classification was followed by Gibbs (16) and Stiles (31) after Gibbs 
(15) had briefly considered sect. StacHycarpus. Florin (8) recognized 
the validity of Bertrand’s sect. Potypopiovsis for P. vitiensis as a sep- 
parate group and added P. minor. This section is now well established. 

1 of these studies, including Gibbs’ (16) description of the female 


° The following symbols indicate the location of the specimens cited: Arnold Arbo- 
retum, Cambridge (a); Bernice P. Bishop Museum, Honolulu (pIsH); Botanic Mu- 
seum and Herbarium, Brisbane (BRI); British Museum (Natural History), London 
BM); Jardin Botanique de |’Etat, Bruxelles (Br); Herbarium of the University of 
California, Berkeley (uc); Wie gand Herbarium, Cornell University, Ithaca (cu) ; 
Dudley Herbarium, Stanford University, Staniord (ps); Herbarium of the Univer- 
sity of Georgia, Athens (ca); Gray Herbarium, Cambridge (cH) ; University of Illi- 


Rijksherbarium, Leiden (1); Missouri Botanical Garden, Saint Louis (mo); Depart- 
ment of Forests, ee and New Guinea, Lae (Laz); New York Botanical Garden, 
New York eM uséum National aonictoie Naturelle, Paris (Pp); United States 
National Museum, Washington (us); National Herbarium of Melbourne, Melbourne 
(MEL). 
A dagger (+) preceding the abbreviation of an herbarium signifies that the details 
of the ee of this specimen have been examined in transverse section. 


74 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


strobilus, were on Fijian specimens of Podocarpus vitiensis. In 1944, 
Wasscher (32) recognized specimens of this species from New Guinea 
and, for the first time, described the dimorphic foliage which had been 
aveooked in the Fiji material. It is interesting that the early specimens 
(Seemann 576, Graff s.n., and Gibbs 674) all have only terminal male 
cones on leafy twigs, wheueds recent male specimens (Degener 14496, Parks 
20653, Smith 7076) definitely show the dimorphic foliage, main twigs 
with scale leaves, and lateral leafy branches bearing male strobili or fertile 
branches bearing scale leaves and strobili or axillary branches bearing 
strobili. 

New Guinea specimens may still prove to represent a separate species, 
but the half-matured ovules which I have seen are still pear-shaped and 
obliquely attached to the receptacle, as in the Fijian Podocar pus vitiensis. 

Both Seemann (25) and Gibbs (16) have waxed ene on the 
beauties of this species and I must quote from Seemann: “This is one of 
the finest Coniferae I have ever seen. . . It attains sixty feet in en 
has a stem nine feet in circumference, and has drooping, extremely grace- 
ful branches, which would render the species a highly desirable acquisi- 
tion to our living collections.” 


2. Podocarpus filicifolius, sp. nov. 


Arbor 15 m. alta ramulis numerosis, divaricatis, spiralibus vel oppositis; 
alabastris in ramulis principalibus parvis, late ovatis, squamis paucis, late 
triangularibus decussatis, 1.5—2 mm. longis;: alabastris i in ramulis foliiferis 
permutatis, ovatis, plerumque constanter dormientibus; foliis dimorphis; 
squamis in ramulis principalibus, divaricatis, tenuibus; late triangularibus, 

mm, longis, acutis vel obtusis; foliis ad ramulis pinnatis terminalibus 
vel axillaribus, sessilibus, divaricatis, oppositis, 6-24 mm. longis, 3.5—4.5 
mm. latis, nceslane tenuibus, planis, apice late acutis, basi fate rotun- 
datis, decurrentibus, costa non manifesta; strobilis masculis ignotis; stro- 
bilis femineis solitariis (?), pedunculis 10 mm. longis, squamas 4— 6 binas 
decussatas vel cicatrices gerentibus; receptaculo parvo lignoso bracteis 
oppositis 2 late obtusis subtento, 7 mm. longo, 3 mm. lato, bracteis 3 
coalescentibus equalibus, apice liberis composito, bractea unica fertili; 
semine maturo globoso, 1.5—1.8 cm. longo et lato, apice aequaliter rotundo, 
obtuso. 


DISTRIBUTION: Morotai, in the Moluccas. 
Moluccas. Morotai, A. Kostermans, in 1949 (Holotype, +L). 


This tree differs from Podocarpus vitiensis in the spreading scale leaves, 
thinner foliage leaves, the distinct receptacle 7 mm. long supporting the 
seeds, and the spherical seeds. The dissected seed shows a smooth, brown 
outer coat 0.3 mm. thick, a hard, woody, light tan middle layer which is 
pointed at the micropyle, and a thin papery brown inner layer. 

The name refers to the fern-like appearance of the pinnately leaved 
twigs, reminiscent of the royal fern, Osmunda regalis. 


1962 | GRAY, REVISION OF PODOCARPUS, XIII 75 


oy pe uiee Reais aie Buchholz, Bull. Mus. Hist. Nat. Paris II. 
284. Guillaumin, Acta Horti Gothob. 19: 8. 1952; Che- 
he pes Mélanésiennes II. 1: 114. 1956. 


A tree becoming 10-12 m. high with trunk up to 80 cm. or more in 
diameter, bearing few branches below but profusely branched above; bark 
brownish, longitudinally furrowed; young twigs green and showing decur- 
rent leaf bases; branches erect or ascending, usually opposite while still 
young; terminal buds appearing naked but protected by special scales 
which become part of foliage but do not grow into leaves. Foliage di- 
morphic; scale leaves on main twigs decussate, broadly triangular, de- 
current, 2 mm. long, becoming obtuse, then ovate-elliptic, divaricate, 3—4 
mm. long; 1 pair opposite scale leaves usually at first node of leafy twigs; 
leaves decussate, emerging in 4 ranks from the twigs and, on vigorous 
branches, twisted at the base so that the pinnately leaved twigs bear 
opposite leaves, on one side all with the abaxial side exposed and on the 
other side the adaxial; on upper reproductive branches of mature trees, 
leaves held obliquely in 4 vertical ranks; leaves lanceolate to elliptic, 
obtuse, 7-15 mm. long, 2-5 mm. wide, thick, sessile, with decurrent leaf 
bases; single midrib not usually distinct, with the surface becoming dry in 
many longitudinal wrinkles. Usually only foliage leaves on small vigorous 
plants and seedlings becoming 30 mm. long, 6 mm. wide, very thin, acutish, 
and opposite on pinnate twigs. Leaves differentially amphistomatic, vary- 
ing with the orientation; transverse sections showing hypoderm inter- 
rupted by the stomatal rows, single vascular bundle with single resin canal 
below phloem, transfusion tissue well developed, no accessory transfusion 
tissue, vascular sclereids abundant above the bundle with fewer below; 
palisade mesophyll on either or both sides depending on leaf orientation. 
Male strobili sessile and terminal on lateral twigs usually bearing regular 
foliage leaves, subtended by a pair of narrow, short foliage leaves or 
bracts; strobili ovoid, 5-6 mm. long, 3 mm. wide; microsporophylls 
broadly triangular, apices acute, margins thin and somewhat erose. Female 
strobili terminal on lateral twigs which may bear ordinary foliage leaves 
or spreading scale leaves; each strobilus bearing 3 or 4 pairs of small, 
opposite, sterile bracts on an axis 10 mm. or more long, with the two 
upper bracts unequal and forming a nonfleshy receptacle, one of the 
bracts fertile. Seed obovoid, 25 mm. long, 18-20 mm. diameter, fleshy, 
red (?) when ripe; kernel with two ridges to a straight or curved beak 
at the micropyle. 


DISTRIBUTION: In forests on mountain slopes of New Caledonia above 
1000 m. altitude. 


ew Caledonia. Mt. Ignambi, Compton 1524 (+pm), 1587 (+BM); Mt. Mou, 
Vieillard 1275 (+p), 3064 (+cH), 3264 —) Bitiel 1085 Grint f4 52) 
1474, 1578 (1LL), 1684 (Holotype, +1LL), 7697 (1LL), MacDaniels 2323 (cv), 
Virot in 1938 (a), White 2033 (+a, BRI), a Laubenfels P129 (GA), a 
607 (pm); Mt. Humboldt, Schlechter 15331 (+pBr, +BM), Mt. Dzumac, 

Laubenfels 153 (GA), Montagne des Sources, De Laubenfels 123 (Ga); Koe, 


76 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


Balansa 184 (+P); Plain des Lacs, Mons du Mai Forest, Buchholz 1350, 1350a, 

LL); Bai des Piroques, White 2120 (+A, 2 sheets); beside River Blanche, 
Ingle I.66 (+MEL); River Tane & Bourail, Balansa 1385 (x). No specified 
locality, Buchholz 1539a (+mo). 


The twig dimorphism observed and described by Wasscher (32) for 
Podocarpus vitiensis is likewise very apparent in P. comptonii. The plant 
bears several kinds of foliage: definitely acute triangular scale leaves on 
vigorously growing main shoots; ovate or elliptic scale leaves on main 
shoots and lower parts of twigs; true foliage leaves lanceolate and ar- 
ranged pinnately; and elliptic foliage leaves arranged on the twigs de- 
cussately. Variations, such as twigs bearing alternately scale leaves and 
foliage leaves for three growth periods with no branching, or branches 
bearing foliage leaves becoming main branches but subsequently bearing 
only scale leaves of either kind, are numerous. 

This species has long been confused with Podocarpus minor which is 
a small tree along streams and in swamps at lower elevations. Compton 
(5), who strangely saw only the large trees, remarked on the inappro- 
priateness of the name “minor.” All the specimens he assigned to P. 
minor belong here. The specimen White 2120 from the Arnold Arboretum 
was labeled P. vitiensis, showing that it was recognized as being different 
from P. minor, Dr. J. T. Buchholz collected both species abundantly and 
the differences became clear to him. The specimen Compton 1273, listed as 
an undetermined species, belongs here. 


- 


Podocarpus minor (Carriére) Parlatore in DC. Prodr. 16(2): 509. 

1868; Brongn. & Gris, Bull. Soc. Bot. Fr. 16: 326. 1869; Mahlert, 
Bot. Centralbl. 24: 281. 1885; Pilger, Pflanzenr. IV. 5(Heft 18): 
62. 1903, Nat. Pflanzenfam. ed. 2, 13: 245. 1926; Schlechter, Bot. 
Jahrb. 38: 16. 1907; Compton, Jour. Linn. Soc. Bot. 45: 425. 1922; 
Dallimore & Jackson, Handb. Conif. 51. 1923, 1931, 75. 1948; 
White, Wilson & Guillaumin, Jour. Arnold Arb. 7: 78. 1926; Florin, 
Sv. Vet-akad. Handl. III. 10: 278. 1931; Guillaumin, Acta Horti 
Gothob, 19: 8. 1952; Chevalier, Etudes Mélanésiennes II. 1: 114 
LOS i; 


Nageia minor Carriere, Traité Gén. Conifeéres. ed. 2. 641. 1867. 
Podocarpus palustris Buchholz, Bull. es Hist. Nat. Paris II. 21: 284. 1949; 
“hevalier, Etudes Mélanésiennes II. 1: 114. 1957. 


A dwarf tree or shrub, 2—3 m. high, trunk 15-30 cm. in diameter with 
numerous ascending branches; bark rough, brown or dark gray; twigs 
opposite on young stems or scattered, ridged by decurrent leaf bases; 
terminal buds normally small with few opposite scales, frequently abnor- 
mally enlarged due to infestation by insect larvae. Leaves dimorphic; 
scale leaves on main shoots often deciduous, decussate, triangular-obtuse, 
keeled, decurrent, 1 mm. long, becoming divaricate: elliptic, thick, to 4 
mm. ions: and usually 1 pair of elliptic scale leaves at the first node of 
leafy branches; foliage leaves crowded on short branchlets, opposite, de- 


1962] GRAY, REVISION OF PODOCARPUS, XIII Ue 


cussate, ascending and held obliquely in 4 vertical ranks, oval or elliptic, 
obtuse, sessile, 10-20 mm. long. 3.5—-5.5 mm. wide, thick and longitudinally 
wrinkled when dry, with broadly decurrent bases; midrib not evident. 
Leaves differentially amphistomatic with twice as many stomata on the 
adaxial side, stomata easily seen under low magnification on young leaves 
as minute white dots. Transverse sections of leaves showing rare hypo- 
dermal fibers on both sides but abundant at the margins, palisade meso- 
phyll on both sides, a broad midvein due to extension of transfusion tis- 
sue half-way to the margin, single resin canal, usually abundant vascular 
sclereids above and below the vascular bundle. Male strobili terminal on 
short, bracted peduncles, 1-1.5 mm. long, in clusters of 3-5 or more at 
ends of short lateral leafy twigs; 5-8 mm. long, 2.5-3 mm. in diameter; 
microsporophylls broadly hastate with narrowly acute upturned apiculi. 
Female strobili terminal on lateral twigs, axis 4 mm. long; strobili of 3 or 
4 pairs of decussate bracts 1-2 mm. long; immature ovules usually crested, 
long-pyriform; 1 terminal fertile bract (rarely 2), keeled, 2 mm. long 
and spread upon the back of the base of the large seed (away from the 
micropyle) with minute or suppressed apex, the 2 upper bracts fusing 
into a small receptacle 4 mm. long. Seed crested, obliquely conically pyri- 
form, 2.8 cm. long, 1.7 cm. wide, becoming maroon-red when ripe; inner 
woody layer usually obtuse, with a beak formed at the micropyle, usually 
straight. 


DistrIBUTION: Along banks of streams and lakes on Plaine des Lacs 
and near Prony Bay, New Caledonia. 

New Caledonia. Borders of Lac Arnaud, Vieillard 1275 (Holotype of P. minor, 
+P; BM, ¢{NOUMEA), Vieillard or Deslanche 170 (+P); Plaine des Lacs, River 
des Tacs Beas 1719 (tILL), 1729 (+1LL), LeRat 607 (+BM), 22 ent Sta- 
tion, Buchholz 1347 (t1LL), 1348 (1LL), 1421 (Holotype of P. palustris, +IL1; 
P), flooded lake shore, headwaters of Yate R., De Laubenfels P112 (cA), Rio des 
Pirogues, White 2261 (srr, tuc, us); Lac en Huit, De Laubenfels P115 (Ga), 
McKee 3382 (us); river sw. of Grand Lac, Virot 658 (+A), MacDaniels 2544 
(t+cu); Prony, Frank 207 Ser. A (+BM, BRI, tF, UC, US); Baie du Sud, Viedllard 
1275 (a, BM). Without specific locality: Levormand 9171 (+k), Raoul s.n. 
(+p), Petit N138 (+p), LeRat 1752 (+P). 

At some sites these dwarf trees grow very slowly and become very. old, 
their bases often growing in standing water and becoming buttressed. At 
times they may be entirely submerged beneath muddy, silt-laden water, 
reddish from the ferruginous soil carried down from the surrounding moun- 
tains during heavy rains. Dr. Buchholz (ms. data) believed there to be 
a related species, Podocarpus palustris which he described, but the dif- 
ferences he listed do not fall outside the normal range of variation in P. 
minor, and the difference in wood density is no more than expected from 
the slightly different ecological habitats. 


LITERATURE CITED 


1. BerTRAND, C. E. Anatomie comparée des tiges des feuilles les Gnétacées 
et les Coniféres. Ann. Sci. Nat. Bot. V. 20: 5-153. 1874 


wn fo 


a 


“I 


bo 
No 


BRO bo 
ios} 


JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


BucuuHoLz, J. T. Gymnosperms. Taxonomy of Coniferales. Encyclo- 
paedia Bans 11: 22-34. 

: ric and subgeneric distribution of the Coniferales. Bot. Gaz. 

110: 30-91. 1948, 

E. Gray. A taxonomic revision of Podocarpus I, The sec- 

tions of ‘the genus and their eae with special reference to leaf 
anatomy. Jour. Arnold Arb. 29: 49-63. 1948. 

Compton, R. H. A systematic account of the plants collected in New 
Caledonia and the Isle of Pines. Part II. Gymnosperms. Jour. Linn. 
Soc. Bot. 45: 421-434. 1922 

De LAUBENFELS, D. J. The external morphology of coniferous leaves. 
Phytomorphology 3: 1-20. 1953. 

FLortn, R. Untersuchungen Stammesgeschichte der Coniferales und Cor- 
daitales. Part I. Morphologie und Epidermisstruktur der Assimilations- 
organe bei den rezenten Coniferen. Sv. Vet-akad. Handl. III. 10: 1931. 

. Die Koniferen des Oberkarbons u. des unteren Perms. Paleonto- 
ae $5(B1 7): 457-654. 1938-44. 

Tertiary fossil conifers of south Chile and ra phyto- 
_geoeraphical significance. Sv. Vet-akad. Handl. III. 19(2). 19 

———. On the relationships of the Taxaceae. Bot. Gaz. 110: A 39. 1948, 

sant in Cordaites and conifers. Acta Horti Berg. 15: 285- 

388. 1951. 

. On two conifers from the Jurassic of southeastern Australia. 

Paleobotanist 1: 177-182. 1952. 

———. The female reproductive organs of the conifers and taxads. Biol. 
Reviews 29: 367-389. 1954. 

he systematics of the gymnosperms. Pp. 323-403 in A Cen- 
ae a Progress in the Natural Sciences, 1853-1953. Calif. Acad. Sci. 

955 


———. Notes on the systematics of the Podocarpaceae. Acta Horti Berg. 
17: 404-11. 1958. 

Grpps, L. S. A Sage to the montane flora of Fiji. Jour. Linn. Soc. 
Bot. 39: 130-212. 19 

—. On the developmen of the female strobilus in Podocarpus. Ann. 


Gray, N. E. & J. T. a CHHOLZ. A taxonomic revision of Podocarpus, III. 
The American species of Podocarpus: section Polypodiopsis. Jour. Arnold 
Arb. 29: 118-122. 8. 

Harr, J. B. & E. J. Beuzenperc. Chromosomal evolution in the Podo- 
carpaceae. Nature 181: 1584-1586. 58. 

KarIser, M. Microstructure of wood of Podocarpus. Phytomorphology 4: 
39-47, 1954. 

Lr, H. L. Present distribution and habitats of the conifers and taxads. 
Evolution 7: 245-261. 1953. 

Mauntert, A. Beitrage zur Kenntnis der Anatomie des Laubblatter der 
Coniferer mit besonderer Beriicksichtigung des Spalt6ffnungs-Apparates. 
Bot. Centralbl. 24: 278. 1885. 

Orr, M. Y. The leaf anatomy of Podocarpus. Trans. Proc. Bot. Soc. Edin- 
burgh 34: 1-54. 1944. 

PiLcer, R. Taxaceae. Pflanzenr. IV. 5(Heft 18): 1-124. 1903. 

Podocarpaceae, Nat. Pflanzenfam. ed. 2. 13: 211-249, 1926. 


1962 | GRAY, REVISION OF PODOCARPUS, XIII 79 


Sib 
Sy 


SEEMANN, B. Flora Vitiensis 266-7. 1865-73. 
SELLING, O. H. Further studies in Schizaea. Sv. Bot. Tidskr. 41: 431-450. 
1947. 


———. Some Tertiary plants from Australia. /bid. 44: 551-560. 1950. 
Smitu, A. C. The vegetation and flora of Fiji. Sci. Monthly 73: 3-15. 
1951. 


erogam genera with distributions terminating in Fiji. Jour. 
Arnold eee 36; 273-292. 1955. 
STEENIS, C. G. G. J. vAN. Results of the Archbold Expeditions. Papuan 
Nothofagus. Jour. Arnold Arb, 34: 301-374. 1953. 
Strives, W. The Podocarpaceae. Ann. Bot. 26: 442-514. 1912. 
WasscHER, J. The genus Podocarpus in the Netherlands Indies. Blumea 
4: 359-481. 1941. 


. Witz, M. H. A new interpretation of coniferous cones I. Podocarpaceae. 
1944. 


Ann, Bot. IT. 8: 1-41. 


AGNES ScoTT COLLEGE, 


DECATUR, GEORGIA 


80 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


TAXONOMIC AND NOMENCLATURAL NOTES ON 
ZANTHOXYLUM AND GLYCOSMIS (RUTACEAE) ! 


GEORGE K. Brizicky 


IN THE COURSE of surveying the genera of Rutaceae in the southeastern 
United States (see Jour. Arnold Arb. 43: 1-22. 1962), the author en- 
countered several nomenclatural and taxonomic problems in Zanthoxylum 
L. and Glycosmis Corréa which require further comment. These include 
the generic limits of Zanthoxylum; the legitimacy of the name Z. cori- 
aceum A. Rich., in view of the existence of a supposedly earlier homo- 
nym; and the correct name of the type species of Glycosmis. The prob- 
lem of Z. coriaceum has led to further bibliographic research on the dates 
of publication of Achille Richard’s work on the flora of Cuba in Ramon de 
la Sagra’s Histoire Physique, Politique et Naturelle de VIle de Cuba. 
These items are dealt with separately below. 


THE GENERIC LIMITS OF ZANTHOXYLUM 


The nomenclatural confusion concerning Zanthoxylum L. and Fagara 
L. seems to have been cleared up by the typification of the former by 
Z. fraxineum Willd. (= Z. americanum Mill.) by Fosberg (1959). (It 
is notable in this connection that Jussieu (1825, p. 505) and Triana and 
Planchon (1872, p. 310) indicated Z. fraxineum Willd. as the type of 
Zanthoxylum L. (“Zanthoxylum Colden—L. J.— Schreb.’”’).) As the 
matter stands at present, the name Zanthoxylum L. has to be applied 
either sensu stricto to the genus with one perianth whorl (a simple or 
haplochlamydeous perianth, according to Engler, or one composed of 
petals, according to Eichler), or sensu lato to the combined genus, in- 
cluding Fagara L. (type, F. Pterota L.) with two perianth whorls (a 
double or diplo- and heterochlamydeous perianth). The recognition of 
two separate genera or of a single inclusive genus remains a matter of per- 
sonal evaluation of the evidence. For the students of tropical African, 
South American, and West Indian floras the generic status of Fagara 
apparently does not seem questionable, since no species of Zanthoxylum 
L. sensu stricto have been recorded from these regions. But students of 
the floras of eastern and southeastern Asia and North and Central America, 
where species of both these taxa occur, face the problem of the recognition 


* Continuing a series of miscellaneous notes and papers on the flora of the south- 


presses his sincere gratitude to Dr. Carroll E. Ait. d, Jr., for his Gaitical reading of 
the manuscript, valuable suggestions, and advi 


1962 | BRIZICKY, ZANTHOXYLUM & GLYCOSMIS 81 


of Fagara as a segregate genus. Opinions have differed. Thus, Rehder 
(1945, p. 73) in his study of Asiatic species of Zanthoxylum and Fagara 
came to the conclusion that “the two genera are close and none of the 
characters are strong enough for generic separation, so it seems preferable 


A different view is represented by Reeder and Cheo (1951, p. 
68) who say, “After studying numerous specimens in this complex, it 
is our feeling that both these genera are worthy of recognition. Although 
there are no striking vegetative differences, flowering specimens are quite 
distinct. Accordingly we are accepting both Xanthoxylum and Fagara, 
an interpretation which is, we believe, in harmony with that of most 
modern students of the group.” 

Such disagreément in regard to the generic status of Fagara is based 
on differences in views as to the morphological nature of the perianth in 
Zanthoxylum. The earliest view, that formulated by Linnaeus in his generic 
descriptions, that the flowers of Zanthoxylum are apetalous while those 
of Fagara have sepals and petals, was followed by all taxonomists up to 
1878 (e.g., Humboldt, Bonpland & Kunth, 1823; De Candolle, 1824; Jus- 
sieu, 1825; Triana & Planchon, 1872; Engler, 1874). This interpretation 
of the perianth favored considering Fagara a subgenus of Zanthoxylum, 
as was done by Triana and Planchon (1872), as well as by Engler (1874). 
Although maintaining the Linnaean view on the perianth of Zanthoxylum, 
Jussieu (1825, p. 505) noted, “Sepala interdum plura, sex aut etiam 
(teste Kunth) novem. Quorum analogia cum petalis confirmatur situ 
alterno staminibus ovariisque opposito, praetereaque metamorphosi ipsorum 
in stamina non infrequenti (observante C. Richard). These features of 
the flowers spoke against the Linnaean interpretation of the perianth 
leaves of Zanthoxylum as homologous with the sepals, but Jussieu’s note 
was disregarded or overlooked by his contemporaries. 

In 1878, Eichler (p. 323), like Jussieu, taking into consideration the 
alternate position of the stamens and the opposite position of carpels (in 
the case of isomery) in respect to the perianth parts (leaves) in Zan- 
thoxylum americanum (“Z. fraxineum”), concluded that the perianth 
leaves in this species are homologous with the petals and the flowers are 
asepalous. Eichler’s view, accepted and followed by most American taxono- 
mists, also was in favor of the inclusion of Fagara in Zanthoxylum as a 
subgenus or section of the latter. 

Engler (1896, 1931), however, after reconsideration of his previous 
(Linnaean) view, came to the conclusion that the perianth of Zanthoxylum 
is simple (haplochlamydeous), not homologous with the sepals, of a primi- 
tive type which occurs in Rutaceae only in this genus, and that it cannot 
be derived from the double (diplo- and heterochlamydeous) perianth of 
Fagara. Consequently, Zanthoxylum and Fagara should be recognized as 
distinct genera. This view has been adopted by many taxonomists, espe- 
cially in Europe. Engler’s interpretation of the perianth in Zanthoxylum 
is objectionable on at least two bases 

First, there is no evidence that the simple perianth of Zanthoxylum 


82 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLUI 


really represents a primitive condition which is not the result of sim- 
plification. On the contrary, Saunders (1934, p. 660, figs. 37-39, & p. 
661), analyzing the carpellate flowers of Z. planispinum Sieb. & Zucc. 
(= Z. alatum Roxb. var. planispinum Rehd. & Wilson), treated its eight- 
merous perianth as consisting of four sepals and four petals. She also 
noted that ‘“exsertion of the two median sepals is delayed until after that 
of petals.” The latter observation, if confirmed by more abundant ma- 
terial, may also suggest some complexity in ontogenetic development of 
the perianth expressed in delay, or perhaps discontinuation of develop- 
ment, of some or all of the sepals (or their homologues). In his considera- 
tions of phylogeny of the Rutaceae, based on Saunder’s floral anatomical 
data, Moore (1936, p. 321) stated that his type “A” of vascularization 
of the floral parts (including Zanthoxylum fraxineum Willd., and appar- 
ently Z. planispinum Sieb. & Zucc., the species studied in greater detail 
by Saunders) ‘is more than likely the result of reduction.” 

Second, the occurrence in Mexico and Central America (perhaps also in 
South America) of species of Zanthoxylum which appear to be transitional 
to Fagara in their perianth structure supports Eichler’s interpretation of 
the perianth in Zanthoxylum, rather than that of Engler. Thus, the peri- 
anth of carpellate flowers of Z. ferrugineum Radlk. (Donnell Smith 6468) 
from Costa Rica is described by Radlkofer (in Smith, 1897, p. 392) as, 
“nerianthii foliola 9-10, linearia (2-3 mm. longa), alia (plus minus con- 
spicue exteriora et sepalis respondentia) breviora et angustiora, alia (sub- 
interiora, reliquiis subalterna) longiora et paullulo latiora, omnia tenuiter 
membranacea.” The staminate flowers of Hinton et al. 10136 (GH) from 
Mexico (identified at Kew as Z. ferrugineum Radlk., but perhaps repre- 
senting a different species closely allied with the latter) possess five to 
ten, often eight to nine, perianth leaves, all similar in appearance, with 
one to five -+_ exterior and smaller than the others. The four to five 
stamens usually are opposite the smaller and alternate with the larger 
perianth leaves. (One five-merous flower which the author examined cor- 
responded exactly with the Fagara type with only the difference that the 
minute sepals were of the same appearance as the petals.) Almost similar 
conditions were found in staminate flowers of Z. mazatlanum Sandw. (iso- 
type, Gonzales Ortega 5210, GH), from Mexico, with 4—11-merous perianth 
and four to six stamens, and Z. Williamsii Standl. (isotype, A. Molina 
1078, Gu), from Honduras, with four to eight perianth leaves and usually 
four stamens. A few carpellate flowers (fruits) of 7. Williamsi exhibited 
six or seven persistent perianth leaves. It is also notable that Engler 
(1874, p. 180), having described Zanthoxylum ciliatum as a new species 
from Venezuela, remarked, ‘Species valde insignis et cum nulla alia 
Austro-Americana adhuc descripta confundenda. . . . Characteribus suis 
transitum inter Zanthoxvlum et Fagara efformat.”’ The present author’s 
conclusion from the above is that the “simple” pcnanth of Zanthoxylum 
is most likely a secondary condition, derived by reduction from that of 
the Fagara type by abortion of some or all the sepals. The occurrence 
of species of Zanthoxvlum which appear in their perianth structure to 


1962 | BRIZICKY, ZANTHOXYLUM & GLYCOSMIS 83 


be transitional to Fagara not only supports this view, but also is ample 
reason to regard Fagara as a subgenus of Zanthoxylum. 


ZANTHOXYLUM AMERICANUM MILL. VERSUS ZANTHOXYLUM 
FRAXINEUM WILLD. 


Fosberg (1958, 1959), when discussing the problem of typification of 
Zanthoxylum, introduced an element of confusion into the nomenclature 
of the type species through the suggested replacement of Z. americanum 
Mill. (1768) by its later synonym, Z. fraxineum Willd. (1796). He 
reasoned, “Furthermore, according to Miller’s description, his X. ameri- 
canum could not be what is now called that as he says of it ‘the flowers 
grow in loose panicles as on the first sort’ rather than in axillary fascicles 
which is the case with the modern Z. americanum Mill. Lawrence, when 
photographing Miller’s types in the British Museum in 1950, was unable 
to locate a specimen of Xanthoxylum americanum so we have no way of 
knowing what plant this name actually refers to. . . . As pointed out in 
Taxon 7(4): 95. 1958, the earliest available name for this is Zanthoxylum 
fraxineum Willd. (1805).” A number of items argue against Fosberg’s 
proposition, however. 

The general description of Xanthoxylum americanum given by Miller 
and his mentioning both the natural occurrence of the species in Penn- 
sylvania and Maryland and its resistance to cold seem to show clearly 
that Miller’s name referred to the plant of the northeastern United States, 
1.e., to the only species of Zanthoxylum which occurs north of Virginia and 
Arkansas. Furthermore, Aiton (1813, pp. 382, 383) mentioned only three 
species of Zanthoxylum which were introduced and cultivated by Miller 
in the Botanic Gardens at Kew. These were “Z. emarginatum Willden. 

. Cult. before 1739, by Mr. Philip Miller. Mill. dict. vol. 2 addenda, 
Lauro affinis 2.0; “Z. Clava Herculis Willden. . . . Cult. 1739, by Mr. 
Philip Miller”; and “2. fraxineum Willden. . . . Cult. before 1759, by 
Mr. Philip Miller. Mill. dict. ed. 7. n. 2.” The last quotation indicates 
that the plant grown at Kew as Z. fraxineum Willd. was that introduced 
by Miller and described by him in the seventh edition (1759) of his 
Gardeners Dictionary as “Xanthoxylum no. 2.” Since the latter, in turn, 
was the same entity as Xanthoxvlum americanum of the eighth edition 
(1768) of Miller’s work (the corresponding descriptions in both editions 
are identical), the conspecificity of the latter species with Z. fraxineum 
Willd. appears to be unquestionable. The circumstance that the type 
specimen of X. americanum has not been found among Miller’s types but 
that the species was introduced into the Botanic Gardens at Kew prior 
to the appearance of the seventh edition of Gardeners Dictionary makes it 
probable that the description of this species was made from the living 
plant. 

In 1771, Du Roi (pp. 57, 58) mentioned the species as Xanthoxylum 
americanum Mill., and provided it with a rather detailed Latin diagnosis, 
including the correct description of the inflorescence and staminate flowers 


84 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


(which he erroneously took for bisexual). He, however, regarded X. amerit- 
canum as a variety of Zanthoxylum Clava-Herculis saying, “Cl. Millero 
species, mihi varietas Zanthoxyli Clavae Herculis L.” In 1772, Du Roi 
(pp. 511, 512) again treated Miller’s species in the same way, but the 
description was given in German. Wangenheim (1787, p. 116) treated 
Xanthoxylum americanum as a species, noting that perhaps it should be 
considered a variety of Z. Clava-Herculis L. Neither Du Roi nor Wangen- 
heim had any doubts regarding the entity named and described by Miller. 

Willdenow, in describing Zanthoxvlum fraxineum (1796, p. 413), ap- 
parently was aware that his new species was identical with that of Miller, 
since he mentioned ‘“‘Zanthoxylum Clava Herculis Du Roi” and ‘“Zan- 
thoxylum americanum Wangh.” in the synonymy and also noted that 
“Der Herr von Wangenheim und Miller geben dieser Art ungezahnte 
Blattchen.” Schkuhr (1803, pp. 467, 468, pls. 323, 3236), although he 
called the species Z. fraxineum Willd., remarked (p. 466) that this was 
the species which ‘‘auch Miller, Wangenheim und andere schon unter 
Zanthoxylum americanum bemerkt haben.” 

Since the entity described by Miller as Xanthoxylum americanum is 
known, neither the occurrence of an error in the description nor the lack 
of a type specimen makes this binomial illegitimate. Therefore, its re- 
placement by Z. fraxineum Willd., as suggested by Fosberg, appears un- 
justifiable in the light of the present Code of Botanical Nomenclature. 


ZANTHOXYLUM CORIACEUM A. RICHARD AND ITS DATE 
OF PUBLICATION 


The legitimacy of Zanthoxvlum coriaceum A. Richard in Ramon de 
la Sagra (Hist. Phys. Polit. Hist. Nat. Cuba. Bot.-Pl. Vasc. [Essai FI. 
Cuba 1.] 326. pl. 34) is often questioned because of the existence of a 
supposedly earlier homonym, Z. coriaceum (Desv.) Walpers, Repert. 1: 
521. 1842 (“‘coriacea”). The former name refers to the species of south- 
ern Florida and the West Indies (absent from Jamaica and a few other 
islands); the latter is a synonym of the Jamaican Z. spinosum (L.) Sw. 
According to Urban (1894, p. 563) and Kuntze (1898, p. 162) the French 
edition of the Richard work appeared in 1845 and preceded the Spanish 
edition which bears the same year of publication. The publication date of 
the portion of this work that included Z. coriaceum A. Rich. was presumed 
by Wilson (1911, p. 185) to be 1842 and by Little (1953, p. 437) to be 
1842 or 1843. Finally, Van Steenis-Kruseman (1960, p. 741), with refer- 
ence to data given by Grisebach (1841, 1842, 1847, 1852) in connection 
with this work, stated, “Up to and including 1840, 28 parts were issued; 
in 1841 11 others followed. At that time the Ist part of the cellular crypto- 
gams was out and the phanerogams were published up to the end of the 
Thalamiflorae (Cand. Syst.). In 1846 parts 1-54 had been published, 
probably no other parts appeared. The atlas, dated 1845 too, was not 
finished before 1851 (cf. Wiegmann’s Repert. I.c. 1852, 387).” This 
statement, however, is not entirely clear and needs further explanation. 


1962 | BRIZICKY, ZANTHOXYLUM & GLYCOSMIS 85 


The Richard work consists of 42 signatures (printed sheets or Bogen) 
of 16 pages each, except the last with only 7 printed pages. Similarly, 
Montagne’s Plantes Cellulaires, of the De la Sagra work, contained 35 
such signatures, the last one of 15 pages. Usually several signatures formed 
an installment (a fascicle or an issue, Lieferung, livraison). It is un- 
known whether Grisebach’s ‘“Lieferungen” translated as “parts” by Van 
Steenis-Kruseman should be understood as installments (improbable be- 
cause of their high number, e.g., 53 by 1846) or referred to the printed 
signatures (Bogen). 

Richard’s work was most likely published irregularly in relatively 
few, perhaps unequal, installments consisting of at least several signa- 
tures. Thus, a notice in Hooker’s London Journal of Botany (1: 308- 
311), issued in-June or July, 1842, mentioned receipt of ‘“‘several of the 
first numbers” of the Richard work, accompanied with plates. At approxi- 
mately the same time, Walpers apparently had at his disposal 192 pages 
(12 signatures) of this work, including Ranunculaceae through Buttneri- 
aceae, since he included in the first volume of his Repertorium (1842) 
new taxa of this group of families (the last species mentioned being Gua- 
zuma parvifolia). Pages 193-336 (signatures 13-21) became available to 
Walpers sometime later in 1842 or 1843, since Richard’s new taxa of 
Tiliaceae (e.g., Belotia) through Rutaceae (e.g., Zanthoxylum coriaceum) 
appeared in Supplementum 1 which was included in the second volume of 
the Repertorium (1843-1844). Endlicher, however, apparently had at his 
disposal both these portions in 1842, since Richard’s new genera, includ- 
ing Belotia, were mentioned in the Addenda to the 2nd Supplementum to 
the Genera Plantarum (1842, after March). Grisebach (1842), recording 
the publication of Richard’s work during 1841, mentioned Simaroubaceae 
as the last family treated by Richard in that portion. Therefore, one 
should assume that the part of Richard’s work which appeared in 1841 
consisted of pages 1-336 (signatures 1-21), and probably Plates 1-35, 
and included the families Ranunculaceae to Simarubeae (as well as one 
page of Ochnaceae with a portion of the generic description of Gomphia). 

According to Grisebach’s report of 1847, the part of Richard’s work 
published during 1846 included the families Ochnaceae to “Portulacceae” 
up to page 624, Although it is unknown whether the corresponding Plates 
36-44(2) also appeared at that time, there is no ground (at least at 
present) for the belief that the appearance of these plates was delayed. 
A relatively long interval between publication of this and the first part 
of the work is also evident from the Richard ‘“Avant-propos” (p. vii) 
which is dated February 1, 1845. “Les circonstances tout a fait indépen- 
dantes de notre volonté ont singuli¢rment retardé la publication de la 
deuxiéme moitié de ce volume. Nous espérons que désormais cet ouvrage 
marchera avec plus de régularité vers la fin de sa publication.” It is un- 
known whether this second part of Richard’s work was issued in install- 
ments or whether it remained undistributed until the completion of the 
whole volume. One may assume that this part was either rare or not 
available at all to botanists for at least a few years after its publication. 


86 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


Thus, Planchon (1846-1847) did not mention Richard’s new species of 
Gomphia in his review of the genera and species of Ochnaceae; Naudin 
(1849-1852) did not include either Pachyanthus A. Rich. or Naudinia 
A. Rich., which was named after him, in the monograph of Melastoma- 
taceae; andl only one new genus (Belairia, Leguminosae) of the second 
part of Richard’s work was mentioned in the third edition of Lindley’s 
The Vegetable Kingdom (1853). 

Finally, in Grisebach’s report of 1852 on phytogeographical and taxo- 
nomic works which were published during the preceding year, we read 
(p. 375), “Von R. de la Sagra’s Werk tber Cuba wurde der erste Band 
der von Richard bearbeiteten Gefasspflanzen (s. Jahresb. f. 1846. p. 53) 
vollendet 14!). Ubersicht der seitdem bearbeiteten Familien: 5 Phytolac- 
ceen, 3 Cacteen, 4 Umbelliferen, 1 Araliaceae, 1 Hedera, 4 Loranthaceen”’ 
and (p. 387) “141) R. de la Sagra, Histoire physique, politique et naturelle 
de Vile de Cuba. Botanique. Plantes vasculaires, par A. Richard. al. tit. 
Essai d’une flore de Vile de Cuba. T. IT. contenant les Dicotylédones poly- 
petales. 663 pag. 8. Atlas. Paris 1845. (aber jetzt vollendet).” It is evi- 
dent that the last parenthetical phrase referred to both the text and the 
atlas, not to the atlas alone. The circumstance that no new taxa of 
oy tolaccaceae (Trichostigma A. Rich., p. 627; T. rivinoides, p. 628; 
Stegnos perma cubense, p. 632), proposed by Richard § in this last part, were 
mentioned by Moquin (1849) in his monograph of Phytolaccaceae (in- 
cluding “Addenda et Corrigenda”’), seems to support the last Grisebach 
statement. 

In summary, I assume the following probable dates of publication for 
Richard’s work: 


ean 


1841: Part 1 (in a few installments?), pp. 1-336 (signatures 1-21), probably 
ls. 1-35, Ranunculaceae to seers ae. 

1846: Part 2 (in installments?), pp. 337-624 (signatures 22-39), probably pls. 
36-44(2), Ochnaceae to Portulacceae. 

1851: Part 3, pp. 625-663 (signatures 40-42) and i-vili, probably pl. 44(3), 
Phytolacceae to Loranthaceae 


Since 1841 may be accepted as the publication date for Zanthoxylum 
coriaceum A. Rich., the priority of this binomial over Zanthoxylum cori- 
aceum (Desv.) Walpers (1842) is clear and the former is the correct 
name for the species of southern Florida and the West Indies. 


THE TYPE SPECIES OF GLYCOSMIS 

Much nomenclatural confusion, with resulting taxonomic discrepancies, 
surrounds Limonia arborea Roxb. (1798), the type species of Glycosmis 
Corréa. Tanaka considered this species to be identical with Limonia 
pentaphylla Retzius (1788) and applied to it the binomial Glycosmis penta- 
phylla (Retz.) Corréa. Narayanaswami (1941), however, came to the 
conclusion that Limonia arborea Roxb. and Limonia pentaphylla Retzius 
are entirely different species, Glycosmis arborea (Roxb.) Corréa and 


1962 | BRIZICKY, ZANTHOXYLUM & GLYCOSMIS 87 


i pentaphylia (Retz.) Corréa, respectively, A brief history of the genus 
d the basic species toed is necessary for an understanding of the 
aa 

In 1788, Retzius (p. 24) proposed Limonia pentaphylla based on 
Koenig’s specimen from the East Indies. The leaves of this species were 
described as ‘‘Folia plerumque in petiolo quina, alterna, ovata, acuta, 
integra, magnitudine foliorum Citri Medicae.’ Roxburgh (1798, p. 60, 
pl. 84) provided the plant which he believed to be Limonia pentaphylla 
Retzius with a more or less detailed description and drawings, and also 
described and illustrated a new species, Limonia arborea (p. 60, pl. 85). 
According to him, the former species possessed relatively small, entire 
leaflets and staminal filaments conspicuously dilated upwards, while the 
leaflets of the latter species were larger than in the preceding and toothed, 
the staminal filaments + filiform. However, Tanaka (1928a, p. 159) 
pointed out that “Roxburgh unfortunately transposed the figures [of the 
flowers] in the two drawings given in his above mentioned book.” 

In 1805, Corréa founded the genus Glycosmis, basing it on Limonia ar- 
borea and L. pentaphyilla as they were understood by Roxburgh. Corréa, 
however, did not make formal transfers of these species into his genus, 
but only remarked (1805, p. 386), “Le Limonia arborea et le Limonia 
pentaphylla de Roxburgh, pl. Coromand. vol. 1, fig. 85, 86 [sic] ... 
m’ont furni le caractére du genre Glycosmis, qui se distingue aisément du 
reste de la famille.” De Candolle (1824, p. 538) apparently was the first 
to make the formal transfer. Glvcosmis arborea (Roxb.) DC. was based 
on Limonia arborea Roxb., and Glycosmis pentaphylla DC. was based 
on the plant identified and illustrated by Roxburgh as ‘“‘Limonia penta- 
phylia Retzius” and only questionably on Retzius’ species (‘G. penta- 
phylla, ... Limonia pentaphylla Retz. obs. 5. p. 24 ? Roxb. cor. 1. t. 
84.”’).2 Most taxonomists, however, subsequently used the binomial Gly- 
cosmis pentaphylla (Retz.) Corréa for this species. 

Engler (1896, p. 185) introduced a new combination, Glycosmis cochin- 
chinensis (Lour.) Pierre ex Engler, based on Loureiro’s Toluifera cochin- 
chinensis (1790) and including G. pentaphylla (Retz.) Corréa taken in 
a very broad sense, It is unknown for what reason Engler ascribed the 
authorship of this new combination to Pierre. As far as the present author 
has been able to determine, Pierre himself did not make the transfer of 
Toluifera cochinchinensis into Glycosmis. In 1893 (text to pl. 285), 
having described a new species, G. montana, from Cochinchina, he men- 
tioned “le Glycosmis pentaphylla Corr. qui est la méme chose que le Toluzf- 
era cochinchinensis (Lour. Fl. Coch., p. 262) ou cam ruu des Anna- 
mites. . .” From a brief description of G. pentaphylia, included in the 
note, it is clear that Pierre understood this species in a broad sense. 

Tanaka, who studied extensively Glycosmis (as well as the other genera 


?Further evidence that G. pentaphylla DC. was reais based on the nonexistent 
basionym Limonia pentaphylla Roxb. is found in the way in which De Candolle 
ee p. 538) cited oe basionym of his Clausena cee “Limonia pentaphylla 

erb. Lamb. non Rox 


88 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


of Aurantioideae), visited European herbaria in the late 1920’s in a search 
for generic types. In 1928, he reported finding, in the Lund Herbarium, 
a Koenig specimen which presumably was the authentic type specimen 
of Limonia pentaphylla Retzius.* An examination of this specimen showed 
that the true Limonia pentaphylla Retz. was conspecific with L. arborea 
Roxb. and different from the plant identified by Roxburgh as L. penta- 
phylla. Consequently, Tanaka reduced G. arborea (Roxb.) Corréa 
(= G. arborea (Roxb.) DC.) to the synonymy of G. pentaphylla (Retz.) 
Corréa. To Roxburgh’s “Limonia pentaphylla Retz.” Tanaka (1928b) 
applied another name, Glycosmis mauritiana (Lam.) Tanaka, based on 
Limonia mauritiana Lam. (1792). Narayanaswami (1941), apparently 
having overlooked Tanaka’s article on the type of Retzius’ species, came 
to the conclusion, based only on Retzius’ description of Limonia penta- 
pAylla which mentions the entire leaflets, that Tanaka’s interpretation of 
L. pentaphylla was entirely incorrect and that L. pentaphylla Retz. and 
L.. arborea Roxb. are different species. 

Narayanaswami was also of the opinion that Corréa should be assigned 
the authorship of the combination Glycosmis pentaphylla, as well as that 
of G. arborea. “But when we take into consideration the facts regarding 
the origin of the genus Glycosmis and the species G. pentaphylla and their 
acceptance by all botanists up to this day, as having been created by 
Correa, it logically follows that Correa should be assigned the authorship 
of G. arborea also, since Limonia arborea Roxb, formed one of the com- 
ponents of the types of the genus Glycosmis of Correa. There appears 
to be no rule in the botanical nomenclature, that governs such cases where 
an author creates a new genus from two species of another genus, and 
leaves it without making the necessary transfer of the earlier species to 
the new genus. But when a subsequent worker assigns the two species 
in their new status to the author of the new genus, does it not become bind- 
ing on all subsequent botanists to follow this adoption by the first bota- 
nist, subsequent to the publication of the genus” (Narayanaswami, 1941, 
p. 25). Narayanaswami was correct insofar as there apparently was no 
clear rule governing the transfers at that time. However, in 1952 at Paris, 
the Eighth Botanical Congress formulated a rule concerning validly pub- 
lished new combinations and illustrated it with very clear examples (Art. 
32, Int. Code Bot. Nomencl. 1954). In the light of this rule, Corréa’s 
mentioning “Le Limonia arborea et le Limonia pentaphylla de Roxburgh” 


®JIn 1932, Fischer reported on an examination of the Koenig collection of the 
Lund Herbarium, which was sent on loan to Kew. Interestingly, Limonia pentaphvlla 
Retzius was listed neither ran the ee ‘Retzius specimens” examined nor among 33 
species mentioned in Retzius’ _Observationes but not found in the Koenig collection. 
specimen of Citrus decumanus,” quoted by Tanaka 
(1928a) included in either of the two lists. The presence, however, in the Koenig 
collection of a number of other specimens which were recognized as authentic Retzius 
jean by Tanaka (Triphasia trifolia (Burm. f.) P. Wils., Plezospermum oe 
(W. & A.) Swingle, and — limonia (L.) Swingle ionone acidissima of 
Retzius]) makes Tanaka’s conclusion in regard to the authenticity of the specimen 
of Limonia pentaphylla ee very probable. 


1962 | BRIZICKY, ZANTHOXYLUM & GLYCOSMIS 89 


as the species on which the genus Glycosmis was based does not constitute 
publication of the new combinations in this genus. Therefore the use of 
Corréa’s authorship for these combinations is against the rules. De Can- 
dolle (1824) seems to be “‘the first botanist subsequent to the publication 
of the genus” who made formal transfer of the above-mentioned species of 
Limonia into Glycosmis as G. pentaphylla and G. arborea, and his author- 
ship for these binomials is indisputable. 

As was mentioned above, Tanaka (1928a), on the basis of the presumed 
type specimen of Retzius’ species, stated that Limonia pentaphylla Retzius 
and L. arborea Roxb. were conspecific and different from the plant de- 
scribed and illustrated by Roxburgh as Retzius’ species. Then Glycosmis 
pentaphylla DC., based on Roxburgh’s plant, not on that of Retzius, must 
be regarded not as a new combination, but as a new name in Glycosmis 
for the species for which Tanaka later (1928b) created the combination 
G. mauritiana (Lam.) Tanaka (based on Limonia mauritiana Lamarck, 
validly published in 1792, not in 1789 as Tanaka believed). Since G. 
pentaphylla DC. cannot be applied to Retzius’ Limonia pentaphylla, 
the next available name for the latter species is Glycosmis arborea (Roxb.) 


The third species involved in the nomenclatural confusion was Gly- 
cosmis cochinchinensis (Lour.) Pierre ex Engler (1896, p. 185) which 
was nomenclaturally based on Toluifera cochinchinensis Loureiro (1890, 
p. 262). Creation of this combination was of no assistance to our knowl- 
edge of the entity described by Loureiro as Toluifera cochinchinensis, for 
the type of the latter is not extant. Merrill has helped to reveal its iden- 
tity. “Although Engler in taking up Pierre’s transfer [sic! ] of Loureiro’s 
specific name intended it to replace G. pentaphylla Corr. as a collective 
species, it is not the same as Limonia pentaphylla Retz. (Obs. 5 
1789) = Glycosmis pentaphylla Corr. Loureiro’s species is represented 
by Clemens 3363, 4448, from thickets at Hue and Tourane [presumed 
classical localities for most of Loureiro’s species from Cochinchina], and 
de Pirey’s specimen of cam ruou, Chevalier 41186. Guillaumin’s descrip- 
tion of Glycosmis cochinchinensis (Lour.) Pierre applies only in small 
part to Loureiro’s species as he treated it as a collective one, citing 14 
synonyms, most of which have to be excluded with the restriction of 
specific limits to the form actually described by Loureiro. True Glycosmis 
epee (Retz.) Correa does not occur in Indo-China” (Merrill, 1935a, 

21). Merrill commented further, “The Hainan specimens closely 
ian Chevalier 41186, Anamese com ruou (Loureiro’s cay cam ruu) from 
Anam, and I believe these to represent Loureiro’s species. This form closely 
resembles G. citrifolia (Willd.) Lindl., but the leaves are constantly 
simple. Chun 5722 from Hainan, which represents the same form as the 
specimens cited above, has been identified by Tanaka as representing 
Glycosmis citrifolia (Willd.) Lindl. var. obtusa (Miq.) Tanaka” (Mer- 
rill, 1935b, p. 17). The quoted notes suggest that Toluifera cochin- 
chinensis Lour. is closely related to Glycosmis parviflora (Sims) Little 


90 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


(G. citrifolia (Willd.) Lindl.) and perhaps should be included in the 


In conclusion, it is to be said that in the light of the current rules of 
botanical nomenclature and of our present knowledge of botanical bibli- 
ography, the nomenclature of the two basic species of Glycosmis appears 
to be the following: 

Glycosmis arborea (Roxb.) DC. Prodr. 1: 538. 1824. 

Limonia arborea Roxb. Pl. See ie - as 1798. 

Limonia pentaphylla Retz. Obs. Bot. 5: 24. 

Glycosmis pentaphylla sensu Bea - a other authors, not G. penta- 

phylla DC. 1824. 


Glycosmis mauritiana (Lam.) Tanaka, Bull. Soc. Bot. Fr. 75: 708. 


Limonia mauritiana Lam. Encycl. Méth. Bot. 3: 517. 1792. 
Limonia pentaphylla Retz. according to Roxb. Pl. Coromand. 1: 60. pl. 84. 
98, not Retzius, 1788 
Glycosmis pentaphylla DC. Prodr. 1: 538. 1824; incorrectly given as G. 
Ne de Se Corréa_ by ona: Rec. Bot. Surv. India 
14(2): 12. 1941 


SUMMARY 


With regard to the generic limits of Zanthoxylum L., especially in con- 
nection with the difference in perianth structure between Zanthoxylum 
L. sensu stricto and Fagara L., the ‘‘simple” perianth of the Zanthoxylum 
type is presumed to be a secondary condition derived by reduction from 
the double perianth of the Fagara type. The occurrence in Mexico and 
Central America (perhaps also in South America) of species of Zan- 
thoxylum which appear to be transitional to Fagara in their perianth 
structure is considered ample reason to regard Fagara as a subgenus o 
Zanthoxvlum, rather than as a distinct genus. 

A number of lines of evidence show that Miller’s binomial Xanthoxylum 
americanum (1768) referred to the northeastern American species which 
was later described by Willdenow (1796) as Zanthoxylum fraxineum. 
Since neither the occurrence of an error in Miller’s description nor the 

*Tanaka’s interpretation of Limonia pentaphylla Retzius, based on the presumed 
type specimen, seems preferable to the Narayanaswami’s concept of the species. This 
is especially true if ae also takes into consideration that the original description of 
Limonia pentaphylla Retz. is well applicable to L. arborea Roxb., except for the entire 
leaflets of the former species and the serrate to crenate ones of the latter. It should 
be remembered, however, that the serration of leaflets in some specimens of L. arborea 
is perceptible only with a lens. On the other hand, the Retzius’ species differs from 
“Limonia pentaphylla Retz.” of Roxburgh in acute leaflets Mes acute” in the 
latter according to Narayanaswami, 1941, p. 14) and their size, “magnitudine foliorum 
Citri Medicae,” (“small” according to Narayanaswamil, loc. cit., “from two to three 
inches long, and about one and a half broad” according to Roxburgh 1798, p. 60). 
The leaves of Citrus Medica are about 12-20 cm. long, 


1962 | BRIZICKY, ZANTHOXYLUM & GLYCOSMIS 91 


lack of the type specimen makes Miller’s binomial illegitimate, the re- 
placement of Zanthoxylum americanum Mill. by Z. fraxineum Willd., re- 
cently suggested by Fosberg, seems to be unjustifiable in the light of the 
present code of botanical nomenclature. 

In the light of the publication dates of Achille Richard’s work on the 
flora of Cuba (in Ramon de la Sagra), as reported by Grisebach (1842, 
1847, 1852), the priority of Zanthoxylum coriaceum A. Rich. (1841) 
over its homonym Z, coriaceum (Desv.) Walpers ( 1842) appears indis- 
putable. Thus, the former binomial is the legitimate and correct name of 
a well-known species of southern Florida and the West Indies. 

A very complex situation in the nomenclature of the type species of 
Glycosmis Corréa, Limonia arborea Roxb. (1798), is shown, and a brief 
history of this genus and the basic species is given. Tanaka’s assump- 
tion of the conspecificity of Limonia pentaphylla Retzius (1788) with L. 
arborea Roxb., based on the presumed authentic type specimen of the 
former, seems correct. Since G. pentaphylla DC., based on Limonia penta- 
phylla sensu Roxburgh (1798), not of Retzius (1788), cannot be applied 
to Limonia pentaphylla Retz., the next available name for the latter species 
is G. arborea (Roxb.) DC. The correct name for G. pentaphylla DC. is 
G. mauritiana (Lam.) Tanaka. The third species involved in the nomen- 
clatural confusion was G. cochinchinensis (Lour.) Pierre ex Engler, which 
was based on Toluifera cochinchinensis Loureiro (1790). Merrill’s notes 
on the latter species, quoted by the present author, suggest a close re- 
lationship of Loureiro’s species with G. parviflora (Sims) Little (G. 
citrifolia (Willd.) Lindl.), rather than with G. arborea. 


LITERATURE CITED 


Arton, W. T. 1813. Hortus Kewensis. ed. 2. 5: 568 pp. London. 

CANDOLLE, A. P. DE. 1824. Rutaceae. Jn: Prodr. Syst. Nat. Regni Veg. 1: 
709-732. 

CoRREA DE SERRA, J. F. 1805. Observations sur la famille des orangers et 
sur les limites qui la circonscrivent. Ann. Mus. Hist. Nat. Paris 6: 376— 


87. 

Du Ro, I. P. 1771. Dissertatio inauguralis observationes botanicas sistens. 
Thesis, 62 pp. Helmstadii. 

. 1772. Die Harbkesche wilde Baumzucht theils Nordamerikanischer 
und anderer fremder, theils einheimischer Baume, Straéucher und strauch- 
artigen Pflanzen 2: 512 pp. & indices. pls. 4-6. Braunschweig. 

Ercuier, A. W. 1878. Bliithendiagramme. 2. pp. ITI-XX & 575 pp. Leipzig. 

ENDLICHER, S. 1842. Mantissa botanica sistens generum plantarum supple- 
mentum secundum. 114 pp. Vindobonae. 

ENGLER, A. 1874. Rutaceae. Jn: Martius, C. F. P. Fl. Bras. 12(2): 77-196. 
pls, 34-37. 

. 1896. Rutaceae. Nat. Pflanzenfam. IIT. 4: 95-201. 

———. 1931. Rutaceae. Nat. Pflanzenfam. ed. 2. 19a: 187-359. 

FiscHer, C. E. C. 1932. The Koenig collection in the Lund Herbarium. Kew 
Bull. 1932: 49-76, 256. 


92 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLII 


Fosperc, F. R. 1958. Zanthoxylum L., “Xanthoxylum Mill.,” and Thylax Raf. 
Taxon 7: 94-96. 

1959. Typification of Zanthoxylum L. Ibid. 8: 103-105. 

GrisEBACH, A. 1842. Bericht tiber die Forschungen in we ie a 
wahrend des Jahres 1841. Arch. Naturgesch. 8(2): 462. 

. Bericht iiber die Leistungen der Sanaa wahrend 
des Jahres 1846. bid. 13(2): 409-472. 

. 1852, Bericht tiber die Leistungen in d. Ape Nae und systema- 

tischen Botanik wahrend der Jahres 1851. Jbid. 18(2): 308-429. 

Humeotpt, A., A. BoNPLAND, AND C. S. KuntH. 1823, ver Nova genera 
et species plantarum 6: 535 pp., pls. 513-600, 

Jussieu, A. 1825. Mémoire sur le groupe des Rutacées. Mém. Mus. Hist. Nat. 
Paris 12: 384-542. pls. 14-29. 

KunTzE, O. 1898. Revisio generum plantarum 3(2): 1-201. 

Lamarck, J. B. A. P. M. ve. 1792. Encyclopédie méthodique. Botanique. 
3(2): 361-753. 

Littte, E. L., Jr. 1953. Check list of native and naturalized trees of the 
United States (including Alaska). U. S. Forest Serv. Agr. Handb. 41: 
472 pp. Washington, D.C 

Loureiro, J. pe. 1790. Flora cochinchinensis: sistens plantas in regno Co- 
chinchina nascentes 1: 353 pp. Ulyssipone 

MerriLL, E. D. 1935a. A commentary on Loureiro’s “Flora cochinchinensis.” 
Trans. Am. Philos. Soc. II. 24(2): 1-445. 

. 1935b. A sixth supplementary list of Hainan plants. Lingnan Sci. 
Jour. 14: 1-62. 

MiItter, P. 1759. The Gardeners Dictionary. ed. 7. London. 

. 1768. Ibid. ed. 8 

Moore, J. A. 1936. Floral anatomy and phylogeny in the Rutaceae. New 
Phytol. 35: 318-322 

Moaguin—Tanpon, A. C. H. B. 1849. Phytolaccaceae. Jn: DC. Prodr. 13(2): 
—40 


NARAYANASWAMI, V. 1941. A revision of the ae ae an species of Glycos- 
mis Correa. Rec. Bot. Surv. India 14(2): 

NaupIn, C. 1849-1852. Melastomacearum que in museo Parisiensi continentur 
monographicae descriptionis et secundum epee distributionis tenta- 
men. Ann. Sci. Nat. IIT. 12-18. 

Pierre, L. 1893. Flore foresti¢re de la Cochinchine. Fasc. 18. pls. 273-288. 
Paris. 

PLANcHON, J. E. 1846, 1847. Sur le genre Godoya et ses analogues, avec des 
observations sur les limites des Ochnacées, et une revue des genres et 
especes - ce Ce London Jour. Bot. 5: 584-600, 644-656; 6: 1-31. 

REEDER, J. p S.-Y. CHEo. 1951. ee on Xanthoxylum & Fagara in 
China. i face Arb. 32: 67-72. pls. 1, 

REHDER, A. 1945. Notes on some euiltivated es and shrubs. Jour. Arnold 
Arb. 26: 67-78. 

Rerzius, A. I. 1788. Observationum botanicarum fasciculus quintus. 32 pp., 
3 pls. Lipsiae. 

Ricuarp, A. 1841-1851. Botanique.-Plantes vasculaires. (Essai d’une flore 

e Vile de Cuba. 1.) Jn: Sagra, R. de la. Histoire physique, politique et 
naturelle de ile de Cuba. Paris. 


1962 | BRIZICKY, ZANTHOXYLUM & GLYCOSMIS 93 


RoxpurcH, W. 1798. Plants of the coast of Coromandel. 1(4): 57-68. pls. 
76-100. London. 

SaunberS, E. R. 1934. On carpel polymorphism. Ann. Bot. IJ. 48: 643-692. 

ScHkuUHR, C. 1803. Botanisches Handbuch der mehresten theils in Deutsch- 
land wild wachsenden, theils auslandischen, in Deutschland unter freiem 
Himmel ausdauernden Gewachse 3: 564 pp. & Nachtrag und Register, 69 
pp., pls. 213-358. Wittenberg. 

SmitH, J. D. 1897. Undescribed plants from Guatemala and other Central 
American Republics. 19. Bot. Gaz. 24: 389-398. 

mae eer AN, M. J., VAN. 1960. Dates of publication and bibliographical 

Fl. Males. Bull. 15: 730-743. 

ee T. 1928a. A note on Retzius’ Limonia pentaphylla, Bot. Not. 1928: 

156-160. 


. 1928b. Revisio Aurantiacearum I. Bull. Soc. Bot. Fr. 75: 708-715. 

TRIANA, J., AND J. E. PLANCHON. 1872. Prodromus vgs es Granatensis. 
Te mebinthace ye: Rutaceae. Ann. Sci. Nat. Bot. V. 14: 

Ursan, I. 1894. Additamenta ad cognitionem florae eee Cae iE. 
Bot. Jahrb. 19: 562-681. 

Watpers, G. G. 1842. Repertorium botanices systematicae 1: 947 pp. Lipsiae. 

; 184 3. Ibid. 2: 1029 pp. Lipsiae 

WANGENHEIM, F. A. J. 1787. Berag zur teutschen holzgerechten Forstwissen- 
schaft, die Anpflanzung Nordamerikanischer Holzarten, mit Anwendung auf 
teutsche Forste. 124 pp., pls. 1-31. Gottingen. 

WILLDENow, C. L. 1796. Berlinische Baumzucht, oder Beschreibung, der in 

n Garten um Berlin, im Freien ausdauernden Baume und Straucher. 452 

pp., pls. 1-5. Berlin. 

Witson, P, 1911. Rutaceae. N. Am. FI. 25(3): 173-224. 


94 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


THE TYPIFICATION OF 
DIOSPYROS EBENUM AND DIOSPYROS EBENASTER 


RicHArp A. Howarp ANp TycHo NorLINDH 


IN THE LATE PART of the eighteenth century, botanists were uncertain 
of the identity of the trees which produced two valuable commercial tim- 
bers, red sandalwood and true ebony. John Gerhard Koenig is credited 
with the identification of each during a stay in India and Ceylon. Koenig 
was born in Polish Latvia in 1728. He moved to Denmark in 1748 and 
studied with Linnaeus from 1757 to 1759. As a physician, he joined a 
Danish mission to India in 1767 and died in India in 1785. The present 
paper concerns the typification of Diospyros ebenum oa source of the true 

ebony), the proper citation of the name, and its synonymy. 

A paper with the Swedish title, ‘ ‘Diospyros Ebenum fe Akta Ebenholz, 
beskrifvit af John Gerhard Konig” |Diospyros ebenum or true ebony, 
described by John Gerhard Koenig|, was published in the first volume of 
the Lund Physiographiska Salskapets Handlingar, printed in Stockholm 
in 1781." In addition to a Latin description of Diospyros ebenum, the 
article contained in a footnote a discussion of the distribution of the 
plant, its characteristics and its use, the method of formation of the char- 
acteristic black wood, and, finally, an eulogy of Koenig for supplying the 
information. The article, in contrast to others in the Handlingar, does 
not indicate the author. The discussion is written in the third person and 
we shall show that an original article written by Koenig was translated, 
edited and published by A. J. Retzius, the founder and the secretary of 
the Lund Physiographiska Salskapet and its publication. The correct cita- 
tion of the name given the ebony tree should be Diospyros ebenum Koenig 
es Retzus, 

The original description apppears to be a composite one, including uni- 
sexual and hermaphrodite flowers and fruit. It was obviously based on a 
field knowledge of the plant in Ceylon. No specimens were cited, and the 
selection of a lectotype for the species is now necessary 

In the library of the Botanical Museum, in Copenhagen, there are a 
number of long letters from Koenig in India during the period 1777-1783 

"The date of publication of Diospyros ebenum is given in Index Kewensis and by 
most recent authors as 1776, the date on the title page. The first volume of this 
eae was published in four parts. It is significant to note that Parts 1 ee 2, pages 
1-64 and 65-132 respectively, were published in 1776, but that Part 3, pages 133-220, 
ee: the description of Diospyros ebenum, was published in 1781, and Part 
4, pages 221-318, was issued in 1786. Otto Gertz viene! Fysiogr. Sallsk. i Lun 
1772-1940 Historisk Overblick, 16. 1940) reports that “the Secretary of the Society 
announced on May 2nd, 1781, that the printing of ‘Handlingarna’ Part 1:3 was 
finished.” 


1962 | HOWARD & NORLINDH, DIOSPYROS 95 


to Rottboell who was chief of botany in the Chair of Medicine in Copen- 
hagen until 1797. One letter, written on August 22, 1777, is fifteen pages 
long. Another pertinent to the present paper was written on February 5, 
1778, and is eight folio-size pages in length. Koenig’s letters are hand- 
written in an old-style German. The structure of the language he used is 
strange today, as is the spelling of many of the words. We are indebted 
to Mr. Sigurd Molander, of the Library of the Botanical Museum of Lund, 
for his assistance in the interpretation and translation of Koenig’s script. 

From Tranquebar, India, on February 5, 1778, Koenig wrote to Rott- 
boell of his recently completed journey to Ceylon and his discovery of 
the true ebony. On May 10, 1777, the English warship “Seahorse,” under 
Captain James’ command and with Koenig aboard, sailed from Madras 
and arrived at Trincomalee, Ceylon, nine days later. Koenig wrote a brief 


4 hE Elon: Clay Tgawe 


os ben the Parag Llbar aloe 
eile me : “a Bilin gt foe Gl 


alta da we : ee Cathey ges. ee opifal= 
Ub ns nage ad mith § ine eo tad eomavee fur aS, an ‘ jicd 
Lee bh is oposce ; i rLeveds gredhes ater eh, nase axafey rel. 
c oe Ver bigfens, phduls, covet 
: a Fy f eget, Haren Pt ar es VE pe 


2 ‘Pp 
pee ee eet - 
» stflert.fujes eb duceti , re aypee’ed ‘Li VRE T, od : ‘ 
Y i vane L y) — a OTTO. ma oa 
ah fot, f re aes: ina fo bs Tak 
je in Yen an me, lip fiellin, ue a, thik Cum m a 


tune pare, vem one, 
oe fle nip oh 
Forartes ONG, oe. ed prespey evbix'c Bee axe , 
: pls aa fees cone soar Cadib, vinedfefes. 
Pract pe ft gl my Lere bee ores, Pad... 
2 un Yoo! por jbrcbufeon:, Vlora Oriente 


CofE pn ea len hus x a xv yobs mnrille brectcrey: 


ms) 


TextT-Fic. 1. First page of Koenig’s manuscript, “Descriptio Ebeni Classis 
Polygama Dioecia.” 


96 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


description of the Trincomalee Mountains, their geology and vegetation. 
He stated that their slopes were rugged and densely covered by many 
kinds of trees, among others very many ebony trees. The season was not 
well adapted for the collection of flowering material, but Koenig declared 
that he had been able to make a complete description of ebony which 
he enclosed and from which the addressee, Rottboell, could see that the 
ebony was a true Diospyros. The description Koenig sent was titled ‘“De- 
scriptio Ebeni Classis Polygama Dioecia”’ and is preserved today. The 
first page of this classic manuscript is reproduced here as TEXT-FIGURE 1. 
During a six-day stay in the area of Trincomalee, Koenig collected many 
new herbs and even concluded that he should very much like to live there. 
Later in his letter Koenig described another landing at Jaffna where the 
true ebony used to be cut. Here he said there were very tall and well- 
formed trees, many of which bore fruit. However, he was not able to find 
a single flower, though he offered rewards of much gold to the finder and 
many trees were thoroughly searched. In his presence two trees were cut 
down, and in these only the innermost part had turned black, showing 
beyond doubt that it was ebony. A forester on the spot told Koenig that 
holes were cut through the bark into the wood in order to encourage 
the development of pigmentation. He explained the white streaks found in 
ebony as incomplete mummification due to premature cutting and con- 
trasted this with the uniformly colored woods obtained in the French and 
Mauritian islands. Koenig indicated in his letter that he had collected 
as many plants as possible and that these had been sent to Copenhagen 
(‘von welchen allen ich im vOnenten Jahr auch nach Copenhagen zureich- 
ende Exemplar tiberstandt habe. 

Although there is no corespondence to support the conclusion, it seems 
clear to us that Rottboell received the specimens sent by Koenig as well 
as the “Descriptio ebeni.” He kept for himself a fruiting specimen and 
the wood specimen. He sent to Retzius a poorer fruiting specimen, and 
either the original manuscript, ‘‘Descriptio ebeni,”’ which was later returned 
to Copenhagen, or a copy thereof. Retzius then edited Koenig’s manu- 
script, for, when the original and the published documents are compared, 
we find several changes in Koenig’s Latin constructions but no significant 
alteration of the diagnosis. The German text of the original manuscript 
was translated freely into Swedish for publication, and Retzius supplied 
only the footnote. 

The selection of a lectotype, therefore, is a choice between the specimen 
Rottboell received from Koenig and sent to Retzius and the specimen 
which Rottboell retained. The specimen in the Retzius herbarium at 
Lund (PratEe I) bears only two ee “Diospyros ebenum,” in 
Retzius’ handwriting (PLATE Va), and “e coll. Retzii,” which is probably 
the writing of Professor Agardh. There are fruits attached to this speci- 
men but these are not associated with the leaves, which are only at the 
ends of the branches and probably represent a later period of growth. 
The leaves are thin in texture, pointed at the apex, and have dried a 
dark color. 


1962 | HOWARD & NORLINDH, DIOSPYROS 97 


are the annotations ‘“Diospyros ebenum verum” and “habitat in vastis- 
simis sylvis prope Jafnapatnam & ad latere montium Trinquemallensium”’ 
(PLATE Vb). Associated with this sheet is a piece of ebony wood and a 
section of a stem with both wood and bark. 

A second sheet in the Vahl herbarium in Copenhagen bears a specimen 
in flower and an annotation partly in Koenig’s handwriting (PLATE Vc). 
A label of more recent date designates this specimen as the “originalek- 
semplar’”’ and indicates that the specimen was received from Koenig from 
“Trankebar ca. 1780.” We do not know who supplied this particular 
label, but that botanist wrongly indicated its origin as “Trankebar.”’ He 
failed to notice the annotation on the reverse side of the sheet which 
reads, “habitat in sylvis Zeylanicis copiose” (PLATE Vc). A sample of wood 
accompanies this flowering specimen. It is important to notice that this 
specimen has both older, coriaceous, light-colored leaves and younger, 
thinner textured, dark-colored leaves on the same shoot. It thus indicates 
the relationship between the types of foliage represented by the Retzius 
and the Rottboell collections. The more authentic label associated with 
the specimen in the Rottboell herbarium, the presence of two wood sam- 
ples, the fruiting condition of the specimen, and the supporting corre- 
spondence have led us to accept the Rottboell specimen as the lectotype 
and to disregard the “originaleksemplar” label. A similar specimen is in 
the herbarium at Lund (PLATE I 

There is in the Linnaean herbarium in London a fruiting specimen 
with similarly shaped heavy leaves (Prat IIIb). We have not been able 
to trace the origin of the Linnaean specimen in any correspondence from 
Koenig. The sheet is annotated “Konig 1777” and “Ebenum Verum ex 
vastis sylvis Zeylonae. Flores non vidi! an Diospyros?” We suspect 
that Koenig, a former student of Linnaeus, might well have sent a speci- 
men of his important discovery directly to his former teacher. The 
chances are that it arrived after the death of the elder Linnaeus (January 
10, 1778), for Linnaeus filius described Diospyros ebenum in his Sup- 
plementum Plantarum Systematis Vegetabilium, page 440, in 1781,” with 


2On the basis of the year of issue, Diospyros ebenum L. {. appears to compete 
with the same name attributed to Koenig. According to the date given in the preface, 
the Supplementum of Linnaeus was completed by October 1, 1780. Linnaeus states 
that his determination was confirmed by Thunberg, who worked with him at least 
in the early months of 1779 (Karsten, Jour. 5S. Afr. Bot. 5: 103. 1939). Stoever 
(The Life of Sir Charles Linnaeus, 295. 1794) quotes Ehrhart as stating that Linnaeus 
“sent me the manuscript in the autumn of 1779, to be printed. J perused it, set 
down my doubts and observations, and sent them to Linnaeus. A correspondence 
then began between us, which lasted almost the whole of the ensuing winter. After 
this, I had it copied afresh, and began to get it quite ready for the press; I was how- 


98 JOURNAL OF THE ARNOLD ARBORETUM | VOL. XLUI 


the comment, “Hoc est verum Lignum Ebenum, cuius originem detexit 
Konig, confirmavit Thunberg.” Linnaeus filius was obviously unaware 
of the nearly concurrent publication of the same epithet by Retzius in 
the Lund Physiographiska Sdlskapets Handlingar. Certainly the date 
1777 and the annotation on the Linnaean sheet, in further agreement with 
Koenig’s letter to Rottboell, suggest that Koenig did send the specimen 
to Linnaeus after he had first seen ebony but while he was still uncertain 
of its identity as a species of Diospyros. 

After his initial visit in 1777, Koenig returned to Ceylon several times. 
The majority of his visits were for but a few days. However, in 1781, 
he made extensive investigations of the flora. It is probable that Koenig 
continued to collect specimens of ebony, including ample flowering ma- 
terial, which he sent to Rottboell at Copenhagen. Rottboell was obvi- 
ously aware of the two publications of the name Diospyros ebenum by 
Retzius and by Linnaeus, of the incomplete published description, and 
of the lack of an illustration of this important plant. In 1783, he pub- 
lished still another description of the plant, this time with an illustration, 
and changed the name to Diospyros glaberrima (Nye Samling K. Danske 
Videnskabers Selskabs Skrifter 2: 540. tab. 5. 1783). It is clear that 
Rottboell was renaming the plants described earlier. He points out that 
his description is largely compiled from Koenig’s published description 
and from material Koenig sent him. Rottboell’s own contributions, for 
which he takes full responsibility for errors, are based on dissections he 
made of the flowers, thus lending supporting to our conclusions that 
Koenig sent Rottboell additional material. Fina lly, in the planation of 
the figures for the ey he supplied, Rottboell notes, ‘“‘Ramus 
Diospyri glaberrimi sive Eb ’ The upper portion of the illustration 
(PLaTE IIIa) is clearly a from the Koenig specimen ‘“Diospyros 
ebenum verum” in the Rottboell herbarium which we have designated 
as the lectotype (PLatre 11). Rottboell did not annotate the sheet itself, 
but the outer cover is labeled Diospyros glaberrima in Rottboell’s hand. 
The lectotype we have selected for Diospyros ebenum Koenig ex Retzius 
and the holotype of Diospyros glaberrima Rottboell are one and the same 
specimen. We believe this is what Rottboell intended. 

The second specimen of Diospyros ebenum in the Retzius herbarium 
(Prate IVb) at Lund is comparable to the specimens in the Rottboell 
(Prate IT), Vahl (Prater [Va) and Linnaean (Piate IIIb) herbaria. 
The Lund specimen, however, bears a copy of the description published 
by Linnaeus filius and the correct page reference to this work (PLatE Vd). 
ever, prevented, by the botanical tour through the electorate of Hanover, with which 

S BRITTANIC MAJESTY had expressly charged me. I got it ready at last, in the winter 
between 1780 and 1781, The work was to be printed at Hanover, under my immediate 


of two ducats per sheet, which I sent to Linnaeus after his return from England.” 
Thus it appears that Diospyros ebenum Koenig has a priority of two or three months 
over Diospyros ebenum L. {. (See Footnote 1.) 


1962 | HOWARD & NORLINDH, DIOSPYROS 99 


The label also indicates that the specimen was collected by Koenig in 
Ceylon and sent to Vahl in 1781 (‘‘misit Praeclar: Demonstr. D: Vahl, 
anno 1781’). Vahl may have sent this to Retzius at Lund. Vahl was 
attached to the Botanical Garden of Copenhagen as lecturer in 1779 and 
was appointed professor of botany there in 1801. 

We conclude that Retzius had not seen this flowering specimen, which 
bears the name Diospyros ebenum L. f. as well as the description and 
reference, when he published Diospyros ebenum Koenig ex Retzius. We 
do not know whether it was this specimen from Vahl or whether Retzius 
saw the specimen which Linnaeus filius must have had at Uppsala, but 
it must have seemed to him that the heavier and bluntly pointed leaves 
of the two specimens (PLATES IVb & IIIb) characterized a different plant 
from the one he had on hand (Prater I) when he published the Koenig 
manuscript. Therefore, in his Observationum Botanicarum (5: 31. 1789) 
Retzius published the following: 

88. DIOSPYROS Ebenaster foliis ovali-oblongis coriaceis, gemmis glabris. 

Diospyros Ebenum L. Supplem. p. 440. 

Hebenaster Rumph. Amb. III. p. 13. t. 6. 

Habitat in sylvis circa Calcuttam. Konic. 

Obs. Manifeste utrasque Diospyri species confundit Nob. @ LINNE Fil. De- 

scriptio hujus habetur in Suppl. l.c. quacum conferri meretur Rumphius. 
$9. DIOSPYROS Ebenum foliis ovato-lanceolatis acuminatis, gemmis hirtis. 

Diospyros glaberrima Friis Rottb. in Novis Act. Hafn. II. p. 540. tab. 5. 

Diospyros Ebenum Physiogr. Saelsk. Handl. V. I. P. 3. p. 176. 

Habitat in Zeylonae sylvis. 

Descriptionem ab inventore concinatum vide 1. cit. 

Folia circiter tripollicaria, quoad consistentiam tenuia, flexilia, obscure viridia. 


A translation of Retzius’ comment under Diospyros ebenaster appears 
to be the following: “Obviously he (Linnaeus filius) mixes up both species 
of Diospyros. The description of this (D. ebenaster) is found in Suppl. 
l.c. with which place Rumphius deserves to be compared.” 

It seems clear, therefore, that Retzius was substituting a new name, 
Diospyros ebenaster, for D. ebenum L. {. and that this species probably 
was known to him at least by a specimen in his herbarium which he had 
received from Vahl in 1781. Regrettably, we have not been able to locate 
any specimens annotated as D. ebenaster by Retzius. Furthermore, it 
appears that Retzius made two mistakes in the protologue of D. ebenaster. 
The first is the description of the buds as glabrous. All of the specimens 
we have seen are more or less pubescent when viewed under the magnifica- 
tion of the usual hand lens. Mr. George Proctor examined for us the speci- 
men in the Linnaean herbarium and confirms that the buds on that, too, 
are slightly pubescent. The second mistake appears to be Retzius’ error 
in referring this species to a Koenig collection from Calcutta. Certainly 
the present specimen in the Linnaean herbarium is clearly marked from 
Ceylon. While Koenig did collect in India, we have not seen any speci- 
mens of Diospyros ebenum collected by him and reported to be from 
Calcutta. 


100 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLIII 


It should also be noted that the Rumphius reference does not apply 
to specimens from Calcutta. Furthermore this name is pre-Linnaean 
(1743) and technically need not be considered in the synonymy. However, 
Retzius obviously based his new specific name on this mononomial. Bak- 
huizen van den Brink in a later publication described the Rumphius 
species as Diospyros lolin Bakhuizen van den Brink * (Gard. Bull. Straits 
Settl. 7: 175. 1933 and Bull. Jard. Bot. Buitenzorg III. 15: 152. 1937). 

There remains for consideration the difference in leaf shape given by 
Retzius in the protologues of the two species Diospyros ebenum and D. 
ebenaster. We have already indicated that the existing Koenig collections 
cited and illustrated show variation in single specimens comparable to 
ae described by Retzius for the two species. During 1961, Mr. D. M. 

A. Jayaweera, director of the Royal Botanic Garden, Peradeniya, Ceylon, 
kindly obtained for us a suite of specimens of D. ehenurs series ob- 
tained from a single tree clearly indicates that the size, shape. and tex- 
ture of the leaves of the ebony do vary. The apex of the leaf blade may 
be acuminate, acute, obtuse, or emarginate on one branch. The texture of 
the blade in successive flushes of growth may be thin and membranaceous 
and black upon drying or the older leaves of the same branch may be 
coriaceous and of a lighter color after drying. The young buds on all 
the specimens examined were densely pubescent, while the buds in the 
axils of older leaves showed less pubescence. 

We have concluded that Diospyros ebenaster is a synonym of D. ebenum 
and that the correct citation and synonymy should be: 


Diospyros ebenum Koenig ex Retzius, Lund Physiogr. Salsk. Handl. 
P70. 178) 


Diospyros ebenum L. f. Suppl. Pl. Syst. Veg. 440. 1781. 

Diospyros glaberrima Rottboell, Nye Saml. Kong. Dansk. Vidensk. Selsk. Skr. 
2: 540. pl. 5. 1783. 

Diospyros ebenaster Retzius. Obs. Bot. 5: 31. 1789. 


The lectotype selected for Diospyros ebenum Koenig ex Retzius is the 
specimen in the Rottboell herbarium (Copenhagen) 

Two monographs of the Ebenaceae published in the last century dif- 
fer in their treatments of this species and its synonyms. Hiern (Trans. 
Cambr. Phil. Soc. 12(1, 2): 27-300. 1873) recognized Diospyros ebenum 
and D. ebenaster as distinct species. The treatment published by Bak- 
huizen is inconsistent and less than clear (Bull. Jard. Bot. Buit. III. 15: 
1-515. 1936-41). His first reference to the species was “Diospyros 


“Tt should be noted here Diospyros lolin Bakhuizen based on Hebenaster Rumphius 


ebenum. By typifying Diospyros ebenaster Retzius with Diospyros ebenum 1 Oe f. t 
way is clear for the use of Diospyros lolin Bakhuizen. 


1962] HOWARD & NORLINDH, DIOSPYROS 101 


Ebenum L.” in an observation (loc. cit. 5. 1936). In the Addenda et 
Corrigenda (loc. cit. 369. 1941) this is corrected to read “Koen. et L. f.” 
which is incongruous. In the body of the monograph (loc. cit. 216. 1937) 
he uses “Diospyros Ebenum Koen.” and in the literature cited the refer- 
ence “Linn. f., Suppl. Syst. Plant. (1781) 440, partim” is given. There 
is no discussion of this conclusion that Linnaeus filius had a mixed col- 
lection and no reference to the other part of the Linnaean concept can 
be found in the monograph. Bakhuizen also established as new two vari- 
eties for ““D. ebenum Koen.” The first variety “A. var. glaberrima (L.f.) 
Bakh. — D. Ebenum Koen. typica, D. glaberrima Linn. f.” * if accepted 
today must be var. ebenum including as it does Koenig’s type. The second 
variety is “B. var. timoriana (Miq.) Bakh. — ?D. Ebenaster Retz., D. 
reticulata Willd. var. timoriana A. DC., D. timoriana (A. DC.) Migq.” 
In the following discussion Bakhuizen states, “D. Ebenaster is a very 
vaguely described species collected by Koenig in the forests of Calcutta. 
Hence it is most probable that the plant in question is only a form of 
D. Ebenum Koen... . It will be better to consider D. Ebenum Koen. 
and D. Ebenaster Retz. forms of the same species.” The range of D. 
ebenum var. timoriana, however, is given by Bakhuizen as Timor, Celebes, 
and the Malayan Peninsula — and Calcutta, if one believes the Retzius 
reference on which this location is based. We have not examined material 
of Bakhuizen’s D. ebenum var. timoriana, nor have we seen authentic 
specimens of D. reticulata Willd. var. timoriana A. DC. We call attention 
to this problem and leave for others the decision as to whether this is 
truly a variety of D. ebenum or some other taxon. 

The taxon which Hiern called Diospyros ebenaster was reported to 
occur in the Philippines and the Celebes and to be in such “cultivated 
places in tropical America, perhaps introduced” as Mexico, Brazil, Cuba 
and Montserrat. Bakhuizen van der Brink renamed this Diospyros nigra 
(Gmel.) Perrottet. Many contemporary floras continue to use the name 
D. ebenaster Retz. for this widely distributed species. The senior author 
has pointed out in a previous paper (Jour. Arnold Arb. 42: 430-436. 
1961) that Bakhuizen was in error in using the epithet D. migra and 
that two species are represented by Hiern’s D. ebenaster which is Bak- 
huizen’s D. nigra. One is a native of the Lesser Antilles of the West In- 
dies, and this species does not appear to be introduced into cultivation out- 
side of the Western Hemisphere. Its correct name is D. revoluta Vahl, 
the type of which was collected in Montserrat. The other taxon involved 
is correctly named D. digyna Jacquin. This plant is a native of Mexico 
and Central America but has been introduced into many areas of Asia 
and is cultivated in many botanical gardens. 

In addition to the individuals mentioned in the body of this paper, we 
wish to express our appreciation to Dr. A. Skovsted, of Copenhagen; Dr. 


*It should be noted that this reference is in error. Linnaeus filius did not publish 
“Diospyros glaberrima” but Diospyros ebenum. Bakhuizen either misread the i 
naean descriptive phrase (q.v.) or intended to credit Rottboell as the author of the 
basionym. 


102 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 
Frances Jarrett, of Kew; and Mr. T. B. Worthington, of Ceylon, for many 
instances of valuable assistance during the course of this study. 
ARNOLD ARBORETUM, 
HARVARD UNIVERSITY 
AND 


NATURHISTORISKA RIKSMUSEET, 
STOCKHOLM 


EXPLANATION OF PLATES 


PLATE I 
Diospyros ebenum in the Retzius herbarium. This specimen presumabl y re- 
ceived from Rottboell along with Koenig’s original manuscript. 
Fiare If 
Lectotype of Diospyros ebenum Koenig ex Retzius in the Rottboell herbarium, 
Copenhagen. This specimen is also the holotype of D. glaberrima Rottboell. 


Prate IIT 
a, Illustration published by Rottboell for aes hae Rottboell 
(Nye Saml. Kong. Dansk. Vidensk. Selsk. Skr. 2: pl. 83). Notice by 
— with Plate II accuracy with which this Hare was ser b, Type 
specimen of D. ebenum L. f. in the Linnaean herbariu 
PLATE IV 


a, Specimen of Diospyros ebenum in the Vahl herbarium, Copenhagen, anno- 
tated by an unknown hand as “Originaleksemplar.” This specimen, incorrectly 
attributed to “Trankebar,” is in flower. ; Flowering specimen in the soot 
herbarium, Lund. This specimen, bearing the description and reference of 
D. eben L. f. may well represent Reizius’ concept of D. ebenaster Retzius. 


Prats: ¥. 

Annotations from specimens of Diospyros ebenum illustrated in Plates ae he 
and IV. a, Annotation in Retzius’ handwriting from specimen in Plate I. b, 
Annotations from lectotype in the Rottboell herbarium (PLare II). c, Anno- 
tation from specimen in the Vahl herbarium (PLATE IVa). d, Annotation from 
specimen in the Retzius herbarium (PLate IVb), 


PuateE I 


Jour. Arnotp Ars. VoL, XLIII 


von cnpnanne 


mg 


fee 


ect 


Howarb & NoRLINDH, DIOSPYROS 


Jour. ARNOLD Ars. VoL. XLIII Pirate II 
Pe et ar se ret ae Ne Ob Re ee eee 


=e 


Howarpb & NorLINDH, Drospyros 


Jour. ARNOLD Ars. VoL. XLIII PriaTE III 


n 

ie) 

24 

m 

Ou 

n 

je) 

_ 

A 

q 

Q 

$reg “ as i= 
ae : ne ee a ee UP 1B) 19 0 21 | A 4 

oe * | 

[2] 

2 

a 

) 
Fs 2 
a ee 
=< 

i 

a 


Howarp & Noriinpu, Diospyros 


IIITX “1OA ‘day GIONYY ‘Anof 


AT aivig 


Jour. ARNoLD Ars. VoL. XLIII PLATE V 


Seopeyn Mowe’ 


ox 


Tr 
~ HB, ROTTBOLL, 


Cee De - g 


wee 
oe les ‘Conse tov hag r6foana tip! Oibregs oe ae Lb. 
Glabhre dy pige con 


C.Lny oe ok Mii co ER . ae 
a0 Cae pte PIN) he PL Ey LES <o 


re Via a) oruen OL 


ee 


oo Le 
<! |e py wt ae le lb mr Gh yteon.. 


F oe 2 G oe fIte7e 
f a-y off a wo farts" ate ta see LOLS 7 ve 


Bafa Cer 


a 
ey, oe é | 2 


« a 
4 
*. d 


/ Mh non fy Spill rte olin en, | 
| MM , cif = ase 


Lape oe ona WZ | 
a ie fe 2 “9 Lo eee ae in tage IGE | anne 1Gf, 


Howarp & NorLINDH, DiospyRros 


108 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


NOTE ON THE RELATIONSHIPS OF PINUS MERKUSII 


NicHoLas T. Mirov 


In nis Genus Pinus (Publication No. 5 of the Arnold Arboretum, 
1914), George Russell Shaw with some hesitation placed Pinus merkusii 
Jungh. & De Vriese in the group Lariciones. Shaw wrote, “Of the habit 
of this pine I know nothing. . . . In my specimens the pits of the ray- 
cells of the wood are both large [as in the group Lariciones| and small 
[as in the group Insignes]. In this particular it may belong in either of 
two groups of species. Its uniform leaf-hypoderm associates it with this 
group |Lariciones| or with P. halepensis of the Insignes. I have assumed 
ee cone to be dehiscent at maturity and have placed it with the Larici- 
ones, but if further information shows the cone to be serotinous, this 
8 should be transferred to the serotinous group [of the Insignes |. ' 

In February, 1961, I had occasion to observe Pinus merkusii in the 
field in Thailand, Indonesia, and the Philippines. I noticed that normally 
P. merkusti cones dehisce on maturity, shed their shiny ochre-yellow seeds, 
and remain open and attached to the trees. I could not help but admire 
Mr. Shaw’s wisdom, his sound judgment, and intuition that caused him 
to place P. merkusii in the group Lariciones. Perhaps it is true that in 
certain characters, such as the uniform leaf-epidermis, P. merkusii re- 
sembles some pines of the Insignes group, but I think this pine has much 
more in common with the species of the group Lariciones. 

The geographical distribution of the Insignes and the Lariciones pines 
is shown below: 


PINES OF GROUP PINES OF GROUP 
REGION INSIGNES LARICIONES 
Mexico 4 None 
W. America 4or5 None 
E. America 6 2 
Europe 3 (incl. P. brutia) 3 (incl. P. sylvestris) 
SE. Asia None 7 


It is seen from the above tabulation that the Lariciones pines are pre- 
dominantly species of southeastern Asia (from Japan to Sumatra). On 
he other hand, the Insignes pines are not found there. Therefore, the 
occurrence of Pinus merkusii in southeastern Asia, where most of the 
Lariciones pines grow, seems to me to be one more point in favor of 
placing this southernmost pine in the group where Shaw placed it. 

The Insignes are a rather heterogeneous group, perhaps the first one 
of the genus Pinus to be revised in the light of its genetic characteristics. 
But such a revision would chiefly concern pines of the eastern United 
States and would not affect pines of southeastern Asia. All pines of that 
part of the world most likely will remain in Shaw’s group Lariciones. 


Casot FOUNDATION, HARVARD UNIVERSITY 


JOURNAL 


OF THE 


ARNOLD ARBORETUM 


VoL. XLIII APRIL 1962 NUMBER 2 


STUDIES IN THE GENUS JASMINUM, II 
THE SPECIES FROM NEW CALEDONIA 
AND THE LOYALTY ISLANDS * 


P. S. GREEN 


EXAMINATION OF THE EXTENSIVE MATERIAL of the family Oleaceae col- 
lected on the Franco-Swiss Botanical Expedition to New Caledonia, 1950— 
1952, and generously sent on loan from the Botanic Garden and Museum 
of the University of Ziirich, soon indicated the need for a revision of the 
species of Jasminum recorded from New Caledonia. Even though Guil- 
laumin published a key to the species as recently as 1948 in his Flore de 
la Nouvelle-Calédonie (p. 283), it soon became apparent that some species 
required reconsideration; and, in addition, two further species, J. lineart- 
folium and J. velutinum, had been described since 1948. In this revision I 
have attempted to see as wide a range of material as possible, and although 
I have not seen all the specimens cited by Guillaumin in his numerous and 
valuable ‘Contributions to the Flora of New Caledonia,” I have been en- 
abled to examine the types of all the taxa concerned. 

When considering specific delimitation in Jasminum, as in many genera, 
it is important to bear in mind the phenomenon of heterostyly. This was 
recorded in Jasminum as early as 1867 (Kuhn, Bot. Zeit. Leipzig 25: 67), 
but unfortunately it appears to have been overlooked by Guillaumin in 
his studies of the New Caledonian species. Jasminum artense and J. 
dzumacense are separated in the key just mentioned by the relative posi- 
tions of the anthers and style: the former-with the anthers borne in the 
upper part of the corolla tube above the style and the latter with the 
reverse condition. Ignoring this as a specific distinction here, it becomes 
clear there are not two species but only one. The relative positions of the 
anthers and styles are mentioned in two or three other places in the key 
as well, and, in 1914, Guillaumin described a variety of J. elatum which 
he called brevistylis, based as it was upon the short or “thrum” condition 
of the style. 

* Results of the Botanical Expedition to New Caledonia 1950-52 (French-Swiss 


Mission). “Studies in the Genus Jasminum, I: Secticn Alternifolia” was published in 
Notes Bot. Gard. Edinb, 23: 355. 1961 


110 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


The immediate affinities of most, if not of all, species included in this 
revision lie with others outside New Caledonia, so it is not possible to 
arrange them in strict order of their suspected relationships. A rough 
grouping has been attempted, however, and, where two species are thought 
to be close enough in affinity, e.g., Jasin artense and J. linearifolium, 
they have been arranged together. 


5 MM. 


Calyx types and range of variation in the New Caledonian species 
Pa si inum. J. didymum, C; J. leratii, F and I to G, rarely almost to D; J. 
neocaledonicum, G; J. noumeense, H; J. artense, Ct o D; J. linearifolium, cer 
a oe A an 0, occasionally, almost oo. ey A; J. kriegeri, 
E awn from Pancher 316; B, Déniker 2270: C, Vieiwlard 913; D, Ddaniker 
483; E, Vivillard 2057 +5. Danie 3089; G, Schlechter 15500 Paice 315% 
I, Daniker 1333) 


a 


It is perhaps significant that three out of the nine species are only known 
from a single collection. Furthermore, additional exploration may well 
indicate what can only be suspected at present; that Jasminum lineari- 


1962 | GREEN, STUDIES IN JASMINUM, II ale 


folium may not be separable at specific rank from J. artense and that J. 
promunturianum is equally close to J. elatum. The calyces of these species, 
as pairs, appear identical, and in Jasminum the shape and dimensions of 
the calyx and its teeth, although somewhat limited in the possibilities of 
their variation, often give convenient characters for identification and a 
strong indication of affinities. Jasminum kriegeri, on the other hand, 
known only from the one somewhat scrappy specimen collected almost one 
hundred years ago, has a distinctive calyx. Ficure 1 gives a diagrammatic 
representation of the calyces and their range of variation in the species 
under review. 

Two alien species widely grown in the tropics throughout the world 
have been recorded from New Caledonia, and it is suspected that they 
have established themselves as escapes from cultivation. They are Jas- 
minum sambac (L.) Ait. (see Guillaumin, Ann. Mus. Col. Marseille II. 
Dell Ot Bulls Wrus: sist. Nat. Paris:25* 500.1010 Nor, Syst, 
Paris 35.61. as. 1914; and Fl. Nouv.-Caléd. 284. 1948) and J. multi- 
florum (Burm. f.) Andr. (as J. pubescens (Retz.) Willd.; see Guillaumin 
and Virot, Mém. Mus. Hist. Nat. Paris II (B). 4: 48. 1953). 

One or two native names are recorded for various species. From the 
Island of Lifou (Loyalty Islands) the name “queku” is noted for 
Jasminum elatum on the label of Deplanch 82 (p) and the name “wechu” 
is reported by Daniker (Viert. Naturf. Ges. Ziirich 78(Beibl. 19): 386. 
1933) as being given to J. didvmum, J. elatum, and J. leratii (as J. sim- 
plicifolium). From the adjacent Island of Maré, Daniker records the 
name “wekutsch” for J. didymum and J. leratii and Baumann-Bodenheim 
(14770) “wawekuce” for J. didymum. It would seem that these are gen- 
eral names for jasmines as a whole. Guillaumin (Candollea 5: 151. 1932) 
quotes information from Bergeret, who had lived in the Loyalty Islands 
and knew the language, that on Lifou the name “wexu tremanji’” is used 
for J. leratii and ‘“‘wexu foe” for J. elatum. 


I should like to express my grateful thanks and appreciation to the 
directors and curators of the cited herbaria for the loan of material or 
facilities for study. All material cited has been examined, and the re- 
spective herbaria are indicated by the abbreviations published in Jndex 
Herbariorum, Ed. 4, 1959. I should particularly like to mention Dr. 
Alicia Lourteig, of Sas: Dr. H. Hirlimann, of Basle; and D 
Stauffer, of Ziirich, to whom I am much indebted for their kind foe and 
Cope ton and Miss Judith Kroll to whom I owe the drawings of calyces. 


Rey Or ser Cis 


ie ee es trifoliolate; corolla tube up to 12 mm. long and lobes up to 2.5 mm. 

lr as Sap a TO alata in Re chi Sete hes a tate gd ela eT 1. J. didymum. 

ile ee simple; corolla tube more than 12 mm. long (rarely as short as 8 
mm. in J. elatum) and lobes more than 4 n ong. 


2. Calyx teeth distinct, more than 1 mm. ice ae 1, F-I). 


112 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLUI 


3. Teeth of calyx narrowly lanceolate to subulate-filiform, up to 3 mm. 
long, usually less (Fic. 1, F to G or I); corolla tube 10-22 mm. long. 

4. Corolla tube up to 20 mm. long, usually less; leaves usually 
narrowly ovate but a. from narrowly lanceolate to broadly 


ovate, (1—)2.5-5.5(-8) cm. long. ................ 2. J. leratii. 

4. Corolla tube 20-22 mm, Re leaves broadly or very broadly 
ovate, (4-)6-9.5(-11) cm. long. ........ 3. J. neocaledonicum. 

3. Teeth of calyx lanceolate, subfoliaceous, 2-5.5 mm. long (Fic. 1, H); 
corolla tube 24-33 mm. long. ................... 4. J. noumeense. 


2. Calyx teeth short or indistinct, up to 1 mm. long (Fic. 1, A-E) 
5. Leaf length not more than twice the breadth. 

6. Pedicels 5-20 mm. long; leaves usually less than 4 cm. long, 
rarely up to 5.2 cm., ovate to orbicular, more or less coriaceous, 
the margin more or less recurved and thickened. .... 5. J. artense. 

6. Pedicels 0-2 mm. long; leaves usually more than 4 cm. long 
narrowly ovate, or if ovate to broadly ovate then always more 
ah 4 cm. long, more or less chartaceous, the margin not usually 

rved or noticeably thickened. ................ 7. J. elatum. 
of Leaf length more than twice the breadth 
gth of leaf more than 3 cm.; leaves linear to narrowly 
ee apex obtuse or acute. 
8. Shape of leaf ee not more than 6 mm. broad; calyx 


tube 1.25 mm. lon Oy. linearifolium. 

8. Shape of leaf ere lanceolate, as 1 mm. broad; calyx 
UDG: AS TOU. oie ye nn eRe 9. J. brerer: 

7. Length of leaf up to 2.5 cm.; leaves oblanceolate or narrowly 
oblanceolate, apex retuse........... 8. J. promunturianum. 


Section TRIFOLIOLATA DC. 


Jasminum didymum Forster fil., Prodr. 3. 1786; De Candolle, Prodr. 
8: 311. 1844; Drake del Castillo, Ill. Fl. Ins. Maris Pacif. 231. 1892, 
et Fl. Polyn. Franc. 120. 1893; Jeanneney, Nouvelle-Calédonie Agri- 
cole 121. 1894; Guillaumin, Ann. Mus. Col. Marseille II. 9: 191. 
1911, et Bull. Mus. Hist. Nat. Paris 18: 40, 329. 1912; Guillaumin 
& Beauvisage, Ann. Soc. Bot. Lyon 38: 102 (‘ Species Montrou- 
zieranae,’ 28) 1914: Guillaumin, Not. Syst. Paris 3: 61, 65. 1914, et 
Bull. Mus. Hist. Nat. Paris 25: 291, 652. 1919, et in Sarasin & Roux, 
Nova Caledonia, Bot. 1: 207. 1921; Moore, Jour. Linn. Soc. Bot. 
45: 356. 1921; Guillaumin, Bull. Mus. Nat. Hist. Paris II. 4: 701. 
1932; Da niker, Viert. Naturf. Ges. ae 78( Beibl. 19) +365,.1933% 
Guillaumin, Bull. Mus. Hist. Nat. Paris II. 6: 458. 1934, zbid. 10: 
519. 1938, ibid. 14: 455. 1942, et Not. Syst. Paris 11: 55. 1943, et 
Bull. Mus. Hist. Nat. Paris II. 15: 453. 1943, et Fl. Nouv.-Caléd. 
283. 1948, et Mém. Mus. Hist. Nat. Paris II(B). 4: 47. 1953, et Bull. 
Mus. Hist. Nat. Paris IT. 27: 475. 1955, zbid. 31: 179, 1959, et Mém. 
Mus. Hist. Nat. Paris II1(B). 8: 161. 1959. 


NE Raaerk R. Br. Prodr. 521. 1810; Labillardier, Sert. Austr.-Caled. 
27. 1824-25; Endlicher, Wien Mus. Naturg. Ann. 1: 177. 1836; De 


1962 | GREEN, STUDIES IN JASMINUM, II 113 


Candolle, Prodr. 8: 311. 1844; Montrouzier, Mém. Acad. Lyon 10: 232. 

1860; Jeanneney, Nouvelle-Calédonie Agricole, 121. ee Schlechter, Bot. 

Jahrb. 39: 231. 1907; Guillaumin, Not. Syst. Paris 11: 1943. 

I, ales var. stenophyllum Daniker, Viert. Naturf. on Ziirich 78(Beibl. 

19): 1933. 

als ee Knobl. ex Guillaumin, Bull. Mus. Hist. Nat. Paris II. 14: 456. 
1942, nomen pro syn. 


Evergreen climber; stems puberulent, often minutely so, or more rarely 
glabrous. Leaves opposite, trifoliolate; common petiole 7—17(—22) mm. 
long, terminal petiolule (2—)4-12 mm. long, the lateral (1—)2-10 mm. 
long, puberulent, especially in the groove above, or glabrous; lamina 
thickish or more or less chartaceous, leaflets narrowly ovate, to narrowly 
lanceolate (or ovate) or more or less elliptic to narrowly elliptic, the ter- 
minal leaflet (0.8—)2—7(-9) cm. long by (0.4-)0.8-3.5(—7) cm. broad, 
the lateral (1—)1.5-5(-—7) cm. long by (0.3—)0.6—-2.5(—3.5) cm. broad; 
margin entire, sometimes slightly thickened, often slightly recurved; apex 
rounded, obtuse or acute, often with an apiculus or the remains of one; 
base rounded, subcordate or acute, not attenuate into the petiolule; vena- 
tion more or less visible as a reticulation: with 4 or 5 (6) primary veins 
per side. /nfl ces axillary or terminal on side shoots, a contracted 
ternately rancned panicle, with few to many flowers, nubentiea or 
glabrous; bracts triangular-lanceolate or subulate, 0.5-2 mm. long; ped- 
icels 0.5-1.5 (—2) mm. long. Flowers white, fragrant, heterostylous. 
Calyx glabrous, occasionally slightly puberulent towards the base, the 
tube 1.5-2 mm. long with 4-6 often obscure teeth 0-2.5 mm. long (Fic. 
1, C), usually slightly ciliolate. Corolla hypocrateriform, tube 5—8 (—10.5) 
mm. long, the lobes 4—7, ovate or broadly ovate, rounded to acute, 1.75-3 
mm. long. Stamens 2, the anthers 2-3.5 mm. long, on filaments 0.5—1 
mm. long. Ovary 0.75-1 mm. long, with style 2—2.5 mm. long in short- 
styled flowers or about 5 mm. long in long-styled. Fruit not seen in New 
Caledonian material. 


New Caledonia. Gatope, Vieillard 913 (a, E, GH); montagnes de Gomouen 
pres Gatope, Vieillard 2935 (BM, K); M’bée, Vieillard 2936 (kK); superieur de 
Koumac, Balansa 2785 (&, K): im Tale der Koumac, am Weg von Koumac 
nach Quegona, bei den Kalk Klippen, im Géhodlz am Bache, 23 May 1925, 
Ddniker 1717 (z); Baie Banaré, Balansa 3189 (BM, E, K); buissons, La Coulée, 
28 March 1928, Franc 2313 (A, us); bois de Port-Despointes (Nouméa), +50 m. 
alt., bois secs des collines littorales, schistes nummulitiques, 21 Nov. 1942, 
Virot 846 (A), ibid., + 40 m. alt., 6 Dec. 1942, Virot 882 (a); Anse Vata, 
Brousmiche 577 (A); Anse Vata, low calcareous hill, 23 Apr. 1955, McKee 2428 
(HULL, US); Oundjo, 1958, McKee 6512 (HULL); bords de la Dumbéa, 50 m. 
alt., 6 June 1909, Franc 1377 (Ny); trail from Dumbéa towards Mt. Dzumac, 
650 m. alt., mesophilous forest on serpentine soil, 28 Mar. 1951, Hurlimann 1092 
(Reale slope south of Paoué valley (Tipindjé), 500 m. alt., meso-xerophilous 
forest on rocky serpentine soil, 13 Apr. 1951, Hurlimann 1166 (a, upper 


forest on calcareous soil, 27 July 1950, Baumann-Bodenheim 5032 (A, ny 


114 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLII 


17 Sept. 1950, Baumann-Bodenheim 6057 (a, z), and 14 Oct. 1950, Guillaumin 
é& Baumann- Bodenheim 6842 (A, z); Mt. Kouvelée (Dumbéa), forest on 
ea soil, 9 May 1951, Guillaumin & Baumann-Bodenheim 13127 (a, z); 

Kaféaté, 19 May 1951, Guillawmin 13395 (a, z); Mt. Mou, gully forest, 
RNG 450 m. alt., 1914, Compton 455 (pm); bei Hie shéne, Apr. 1921, 
Heim 74 (z): Tiebaghikuppen, am Nordostabhang, auf Serpentin, im Gebtisch 
auf Steinen, oft und verbreitet, 14 Mar. 1924, Daniker 1436 (holotype, var. 
stenophyllum, z). 

F Pines: Le Rat 109 oR near Kuto, on coral soil, 26 May 1951, 
Baumann-Bodenheim 13454 (a, z); Anse Kanumera, on coral soil, 27 May 1957, 
Baumann-Bodenheim 13608 (2); a aterie Island, near Isle of Pines, Arau- 
caria ei Pandanus forest on coral soil, 25 May 1951, Baumann-Bodenheim 
13436 

ee Tslands. Liou: Mou, oft an den felsigen Abhangen der Ostkiiste, 29 
Nov. 1925, Daniker 24635 (z); Natalo, im Buschwalde, 2 Nov. 1925, Déniker 
2465a (z); Képénéé, in Wald, 17 Apr. 1912, Sarasin 730 (z). OuvEA: May 1912, 
Sarasin 890 (z). Mare: Pénélo, on coral soil, 18 July 1951, Bawmann-Boden- 
heim 14770 (A, 2). 


Easily distinguished amongst the jasmines of New Caledonia by its 
trifoliolate leaves, this species is nevertheless very variable. The vari- 
ability is most obvious in leaflet size and breadth but also in corolla size, 
although the corolla tube and lobes are always smaller than in any of the 
other species. In 1933, Daniker described his variety stenophyllum based 
upon a small, narrow-leafletted variant (Daniker 1436) but, to judge from 
the considerable number of specimens I have examined, it falls into the 
range of the apparently continuous variation exhibited by the species. 
It is worth noting, too, that Virot 846, with leaflets similar to and perhaps 
a little narrower than Ddniker 1436, is said, in the field notes on the 
label with the specimen, to be a “forme jeune’, but I have no knowledge 
to what extent leaflet shape and size may vary with maturity. The name 
“Jasminum fitzgeraldii” attached by Knoblauch to the specimen of Daniker 
2465 at Zurich was never validly published, and Guillaumin, when drawing 
attention to the variability of J. didymum (in Bull. Mus. Hist. Nat. Paris 
II. 14: 455. 1942), remarks that it is only an intermediate between the 
linear-leafletted expression, as exemplified in the extreme by LeRat 1610 
(not seen, also mentioned by Guillaumin in ibid. 6: 458. 1934) and the 
almost orbicular-leafletted Vieillard 2935 

Jasminum didymum lies within a complex distributed over the Aus- 
tralasian and Malaysian areas; a complex which calls for investigation 
and treatment as a whole, with geographical distribution as only one of 
the characters taken into consideration. Jasminum didymum was the 
first species described in this complex; in addition to J. divaricatum R. 
Br., described from Australia and included in the synonymy above, the 
other species are: J. dallachit F. Muell. (Australia), J. degeneri Kobuski 
(Fiji), J. domatiigerum Lingelsh. (New Guinea), J. gilgianum K. Schum. 
(New Guinea), J. lineare R. Br. (Australia), J. micranthum R. Br. 
(Australia), J. parviflorum Decne. (Timor), J. racemosum F. Muell. 


1962 | GREEN, STUDIES IN JASMINUM, II FES) 


(Australia), J. rupestre Blume (New Guinea), J. smithianum Kobuski 
(Fiji), and J. triphyllum Merrill (Philippine Is.). 

It appears that Jasminum didymum may exhibit a certain amount of 
androdioecism, for, despite the examination of several flowers, no styles 
have been found on some specimens (for example, Franc 2313 and Vieillard 
913), and the ovary, whilst normal in size, is presumably abortive. The 
anthers however, were normal and full of pollen. In other specimens both 
the style and stamens were normal, although exhibiting heterostyly. 


Section UNtroLioLtata DC. 


2. Jasminum leratii Schlechter, Bot. Jahrb. 40(Beibl. 92): 32. 1908; 
Guillaumin, Ann, Mus. Col. Marseille IT. 9: 191. 1911, et Not. Syst. 
Paris 3: 62. 1914, et Bull. Mus. Hist. Nat. Paris II. 6: 458. 1934, 
ibid. 13: 476. 1941, ibid. 15: 454. 1943, et Fl. Nouv.-Caléd. 284, 
1948, et Mém. Mus. Hist. Nat. Paris II(B). 4: 47. 1953, et Bull. 
Mus: HisteNat,.Pans Il. 272327, 475. 1955. 


J. francii Guillaumin, Bull. Mus. Hist. Nat. Paris IT. 5: 323. 1933. 

J. neocaledonicum Schlechter var. angustifolium Daniker, Viert. Naturf. Ges. 
Zurich 78(Beibl. 19): 366. 1933. 

J. daenikeri Knoblauch, Repert. Sp. Nov. 41: 150. 1936. 

J. paagoumenum Knoblauch, Repert. Sp. Nov. 41: 150. 1936. 

J. absimile L. H. Bailey, Gent. Herb. 4: 346. 1940, Hortus Second 397. 1941, 
et Man. Cult. Pl. ed. 2. 797, 798. 1949: P. Riedal (Calif. Arboretum 
Foundation), Pl. for Extra-Trop. Regions 335 

J. noumeense Schltr. var. microphyllum Guillaumin, Bull. Mus. Nat. Paris 
II. 1 54. 1943 

“J. simplicifolium Forst. f.” sensu Guillaumin, Ann. Mus. Col. Marseille II. 

1911, et Not. Syst. Paris 3: 62, 65. 1914, et Bull. Mus. Hist. Nat. 
Paris 25: 652. 1919, et in Sarasin & Roux, Nova Caledonia, Bot. 1: 207. 
1921; Moore, Jour. Linn. Soc. Bot. 45: 356. 1921; Guillaumin, Candollea 
5: 151. 1932; Daniker, Viert. Naturf. Ges. Ziirich 78(Beibl. 19): 368. 
1933; Guillaumin, Bull. Mus. Hist. Nat. Paris II. 5: 323. 1933; ibid. 6: 
458, 1934, ibid. 15: 454. 1943, et Fl. Nouv.-Caléd. 284. 1948 

“J. australe Pers.” sensu Montrouzier, Mém. Acad. Lyon 10: 231. 1866; 
Guillaumin & Beauvisage, Ann. Soc. Bot. Lyon 38: 123 (‘‘Species Mont- 
rouzleranae,” 49) 1914. 


Evergreen diffuse shrub, climber, or liane of variable length; the stem 
glabrous or minutely puberulent, especially when young. Leaves opposite, 
simple; petioles 1.5-10(—20) mm. long, glabrous or minutely puberulent, 
articulated 14 to ¥% way from base; lamina more or less chartaceous, 
(broad ovate to) ovate to lanceolate (to linear lanceolate), (1—)2.5—5 (8) 
cm, long by (0.4—)1.5-4(—5) cm. broad; margin entire, not or scarcely 
thickened, rarely slightly recurved; apex obtuse or acute, sometimes more 
or less rounded or apiculate; base rounded or obtuse, sometimes more or 
less truncate or acute or even angustate, often slightly attenuate into the 
petiole; venation more or less obscure or reticulate or with primary veins 


116 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


only visible above and below, 3—5(6) per side, the upper 3 or 4 often 
anastomosing towards the margin. /mflorescences terminal on side shoots 
or axillary, ternately paniculate with 3—9(—18) flowers per panicle, gla- 
brous or minutely puberulent; bracts linear-filiform, 1-4 mm. long or 
occasionally approaching foliaceous and slightly longer; pedicels 2—20 
mm. long. Flowers white, feebly to strongly fragrant, heterostylous. 
Calyx glabrous or minutely puberulent, tube (1.5—)2-2.5(—3) mm. long 
with 4—6, triangular-lanceolate teeth, (0.5—)2—3.5(—4) mm. long (Fic. 1, 
ranging from G, through I and F, to rarely almost D), more or less ac- 
crescent in fruit. Corolla hypocrateriform, tube 11-20 mm. long, lobes 5-7, 
lanceolate or narrowly lanceolate, acute, 7-12 mm. long. Stamens 2, 
anthers 3-4.75 mm. long on filaments 0.5-2 mm. long. Ovary 1—1.25 mm. 
long, style 3-6.5(—9) mm. long in short-styled flowers or 11-16 mm. long 
in long-styled. Fruit ellipsoid-ovoid, paired (or single by abortion) 6-8 
mm. long by 5—6 mm. broad. 


ew Caledonia. Magenta, Jan. 1903, Le Rat 172 (isotype, Pp); Noumeéa, 
Franc 1544 (syntype J. francii, p), Franc 1544A (syntype J. francii Pp; isosyn- 
type, A); Port de France (Nouméa), 1855-60, Vieillard 911 (A, G, K); Port 
Despointes, 50 m., bois secs des ee littorales, schistes nummulitiques, 6 
Dec. 1942, Virot 876, 881 (A), ibid., 13 Dec. 1942, Virot 922, 927, 929 (a); 
Prony, coteaux niles: roches ales aise 20 Fe b. 1914, Gi 1906 (A, 2); 
Koné, oft im niederen Gebusch der Kiistenhiigel, 11 Feb. 1925, Daniker 770a, 
3088 (z); an der Miindung des Flusses von Koné, zerstreut im Gebische der 
sumpfigen Flussdeltas, 10 Feb. 1925, Daniker 1171 (holotype J. neocaledonicum 
var. angustifolium & J. daenikeri, z); bei Paagoumen, im Grasbestand auf 
Serpentingebiisch, 22 March 1925, Ddaniker 1518 (holotype J. paagoumenum, 
z); Ile de la Table, bei Koumac, in Acaciengebtisch, 5 March 1925 

1333, 1335 (z); auf der Insel Neba, im Sumpfwaldchen bei dem Teiche, 8 
April 1925, Ddniker 1599 (z); am Anse longue bei Carticaté, St. Vincent, im 
Ufergehélz, 12 Dec. 1924, Daniker 770 (z); Anse Vata, low calcareous hill, 
remnant of dry forest with trees close together, 23 Jan. 1955, McKee 1973 (A, 
HULL, US), 475 and 2342 (HULL); slope of Ouen Toro, 22 March 1955, McKee 
2251, (A, US); Ouen Toro, promenade Pierre Vernier, 10 m. alt., climax littoral 
a Acacia spirorbis, nummulitique, terraine schisto-calcaire, 30 Jan. 1941, Virot 
442 (a); N.W. of Mt. Natégou, beneath the road to Yaté, shrubby forest on 
rocky serpentine ridge, 270 m. alt., 29 Jan. 1951, Hirlimann 769 (A, z); “Bois 
du Sud” (towards the Yaté valle y, Spermolepis forest on serpentine, 29 March 
1951, Guillaumin & Baumann-Bodenheim 11663 (A, z); south of Mt. Poindas 
(Tip SS mesophilous forest on serpentine ridge, 520 m. alt., 21 April 1951, 
Hiurlimann 1290 (a, z); Mt. Koniambo, on serpentine soil, 21 Dec. 1950, 
Guillaumin & Baumann-Bodenheim 9514 (a, z); Mt. Kaféaté, on serpentine 
soil, 22 Dec. 1950, Guillaumin & Baumann-Bodenheim 9665 (A, z), ibid., on cal- 
careous soil, 10 April 1951, Guillaumin & Baumann-Bodenheim 12043 (A, Z); 
Moro Island, on coral soil, 5 & 6 March 1951, Guillaumin & Baumann-Boden- 
heim 11106, 11135, 11161 (a, z); Col de Vulcain, xerophilous forest on 
serpentine soil, 900 m. alt., 12 Nov. 1950, Baumann-Bodenheim 8285 (A, 2); 
Gatope, 1861-67, Vieillard 2938 (a, G, GH, P) and 2935 (holotype J. noumeense 
var. microphyllum, vp); Voh, March 1921, Heim 49 (z). IsLE or PINEs: 


1962] GREEN, STUDIES IN JASMINUM, II 117 


Germain 13 (A); Kuto peninsula, on calcareous soil, 28 May 1951, Baumann- 
Bodenheim 13642 (a, z) 

Loyalty Islands. Lirou: bei Képénéhé, im Buschwalde, 31 July 1925, Daniker 
1886 (z), ibid., 22 Oct. 1925, Daniker 2269 (z), 19 April 1912, Sarasin 185 
(z); in der Umegebung von Natalo, oft im Gebiisch, 2 Nov. 1925, Denver 2388 
(z); zwischen Natalo und Tingeting, in der Savanne zerstreut, 2 Nov. 1925, 
Déaniker 2388a (z); Kiatheng, in der Savanne, 21 Nov. 1925, Daniker 3089 (z). 
Ouvea: May 1912, Sarasin 885 (z); Fayaoué, in den Gebiischen, 4 Aug. 1925, 
Déniker 1909 (z). Maré: uberall auf dem Plateau von Maré, 29 Nov. 1911, 
Sarasin 424 (z); Eneni, oft in der inneren Ebene von Maré, besonders lings 
der Wege, 29 Dec. 1925, Daniker 2626 (z); Tadine, forest on coral soil, 12 
July 1951, Baumann-Bodenheim 14529 (a, z); Rawa, mesophilous forest on 
coral soil, 17 July 1951, Baumann- Bodenheim 14721 (A, z) without locality, 
Jan. 1908, Franc. 1021 (ny ie 


tie 


a rol ° 
TE oe Kee be ce Et 


it 


ese 


i) 
o 


laced ios re) 


AONANDSYS 


5 
LEAF : AVERAGE LENGTH IN CM. 


Fic. 2. Frequency histograms of leaf eee in Jasminum sei (L}, 
volubile (V), and J. simplicifolium (S). LH1 and NH indicate the posi of rf 
specimens examined from Lord Howe eee and the New Hebrides eee ae. 


118 


Cult 


1917, cae 
enone 


(BH. 


Fic. 
A fact 


4 


o. 
ext 


FREQUENCY 
° 


BH ) 


JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


(A); 


San Dieg 


ivated. United States of oe California: Santa Barbara, 27 July 
g 


19, Woodcock 630 (holotype of J. 


; Golden Gate Park. San Francisco, 23 March 1927, Bailey 9623 
-R, Menzie’s place, Mill Valley, 24 July 1927, Walter 65 (A). 


Greatest leaf breadth factor: measurement B divided by measurement 
ae 


LEAF ° 


i 


9 32 35 .38 41 44 47 80 
GREATEST BREADTH FACTOR 


Fic. 4. Frequency histograms for greatest leaf breadth factors in Jasminum 
leratii (L), J. volubile (v), and J. simplicifolium (s). LHI and Nu indicate the 
position of the specimens examined from Lord Howe iad and the New Heb- 
rides respectively. 


1962 | GREEN, STUDIES IN JASMINUM, II 119 


A close relationship exists between this species, Jasminum sim plicifolium 
Forst. f. and J. volubile Jacq. (J. gracile Andr.), and, after considerable 
study of material of all three, I have come to the conclusion that, for the 
present at least, they should be treated as separate species. Without doubt 
all three lie very close together and it is not without significance that J. 
leratti was placed as a doubtful synonym under J. simplicifolium by 
Guillaumin in 1921 (in Sarasin & Roux, Nova Caled. Bot. 1: 207. 1921) 
and later by Daniker (Viert. Naturf. Ges. Ziirich 78(Beibl. 19): 368. 
1933), although the two were maintained as distinct in Guillaumin’s rela- 
tively recent Flore de la Nouvelle-Calédonie 284. 1948. Bailey (Gent. 
Herb. 4: 342. 1940) separates J. volubile (as J. gracile) and J. simplici- 
folium and inférs that the former is confined to Australia, an inference 
supported by this investigation. For this study a careful analysis has 
been made of leaf length, distance from the base to the greatest breadth 
of the leaf, length of calyx teeth, length of pedicel, and length of corolla 
tube; and each has been considered in relation to geographical distribu- 
tion. The results may be expressed as follows: (i) LEAF LENGTH. The 
extra-Novo Caledonian J. simplicifolium (from Fiji & Tonga) has longer 
leaves than either J. leratii (from New Caledonia) or J. volubile (from 
Australia), although, using the average leaf length of a specimen, a slight 
overlap occurs around 5.5-6 cm. (Fic. 2). (ii) GREATEST LEAF BREADTH 
FACTOR. Bailey (doc. cit.) indicated the possible value of the position of 
greatest breadth of the leaf in distinguishing between these species. In 
this analysis a “greatest leaf breadth factor” was calculated by measuring 
the total length of a leaf, and dividing by the distance up the midrib to 
the position of greatest breadth (see Fic. 3; B divided by A). Using this 
method it was found that J. simplicifolium had the lowest factor, less than 
0.3 whilst J. leratii and J. volubile both had factors ranging from 0.3 to 
0.45 (Fic. 4). That is to say, J. simplicifolium has leaves which have 
their greatest breadth in the lower third and the other species between the 
lower third and middle of the leaf. (iii) LencrH OF CALYX TEETH. 
Jasminum volubile is more or less separable from J. leratii and J. simplici- 
folium on this character. In the first species, the teeth range from prac- 
tically nothing to no more than an average of 0.75 mm. in length, usually 
less, whilst in the latter two species they vary from an average of just 
under 0.75 mm. to 2 mm. in J. simplicifolium and to nearly 4 mm. in J. 
leratii, which is very variable in length of calyx teeth (Fic. 5). (iv) 
LENGTH OF PEDICEL. There is no clear cut separation of the taxa on this 
character but only a very slight tendency for J. volubile to have shorter 
pedicels than the other two species. (v) LENGTH OF COROLLA TUBE. 
Jasminum volubile is distinguishable by its shorter corolla tube which, in 
the material examined, ranges from about 8 to 11 mm., whereas in J. 
lerata and J. simplicifolium (which together seem indistinguishable on this 
character) it ranges from 11-20 mm. (Fic. 6). 

These details may be summarized by saying that Jasminum leratii has 
leaves which are shorter and slightly more elliptical than those of J. 


JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


FREQUENCY 


t 
5 | U 


] | T 
: 2 3 4 
CALYX TEETH : AVERAGE LENGTH IN MM. 
. Frequency histograms for length of calyx teeth in Jasminum leratu 
(L), J. volubile (v), and J. simplicifolium (s). Lui and NH indicate the position 
of the specimens from Lord Howe Island and the New Hebrides respectively. 


5 


FREQUENCY 
a 
pit 
WN 


1.0 
COROLLA TUBE : AVERAGE LENGTH IN CM. 


Fic. 6. Frequency histograms for length of corolla tube in Jasminum 
(L), J. volubile (v), and J. simplicifolium (s), Lut indicates the position 
i i d. 


leratit 
of 
specimens examined from Lord Howe Islan 


the 


1962] GREEN, STUDIES IN JASMINUM, II 121 


simplicifolium and generally has a larger corolla tube and calyx teeth 
than J. volubile. 

Whilst not strictly observations on New Caledonian material, it is 
interesting to make two notes on allied material. The only specimen seen 
from the adjacent New Hebrides (Kajewski 404, a, NY) is Jasminum 
lerati. It was originally identified as J. simplicifolium (Guillaumin, Jour. 
Arn. Arb. 13: 17. 1932), but, although it has no corollas on it to examine 
and measure (being in fruit), in leaf dimensions and shape, and in calyx 
type, it falls into the range of J. leratii. The two specimens examined from 
Lord Howe Island in this investigation (McCornish 4 [A] and “Phytologic 
Museum of Victoria, Government Botanist Melbourne” [E]) have leaves 
and calyx teeth which in length lie on the edge or outside the ranges 
of the Australian specimens of J. volubile. Yet in greatest leaf breadth 
factor and length of corolla tube they agree with the rest of the Australian 
material. 

At the same time as Jasminum leratii, J. volubile, and J. simplicifolium 
were being investigated, the opportunity was taken to examine the 
cultivated species J. absimile L. H. Bailey, and a separate note about 
this will appear in Baileya. It is sufficient to point out here that J. absimile 
is conspecific with J. leratii, although in the former there are usually 
more flowers per inflorescence and the leaves are a little larger on the 
average. But both these differences can be accounted for by favorable 
conditions of growth in cultivation. 

Jasminum leratii is very variable in leaf shape and size, and it is sus- 
pected that this may be a reflection of the habitat conditions under 
which the plants are growing. Some of the largest leaves (up to 7 cm.) 
are exhibited by a specimen of Ddniker 1909; yet under this same 
number is a shoot with quite typical leaves half their size. Ddniker 1171 
exhibits leaves which are consistently small and narrow. Because of this 
it was used as the type of both J. daenikeri and J. neocaledonicum var. 
angustifolium, but some of the leaves on Le Rat 172, the type of J. leratii, 
are an exact match, as are those of Guillaumin & Baumann-Bodenheim 
11663, in which they range from broad-ovate to linear, although con- 
siderably larger than in Le Rat 172. Similarly, a small-leaved scrap (Vieil- 
lard 2935) was the basis of the published name J. noumeense var. miicro- 
phyllum (although the type sheet at Paris bears only the name J. nco- 
caledonicum var. microphyllum in Guillaumin’s hand). The leaves are 
shorter than those of Ddaniker 1171 but they can be matched exactly by 
the smallest leaves of several other specimens and in floral characters this 
specimen is identical with the rest of J. leratit. 

Dr. Hirlimann has remarked in correspondence that it is uncommon 
for endemic species in New Caledonia to occur on calcareous as well as 
on serpentine soils, but using dried specimens I have been unable to discern 
any consistent differences between individuals from the two habitats. It 
is possible that there are differences which are masked in the herbarium 
by the limitations of the material and the wide variability of the species, 


122 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLIII 


and an examination of populations in the field with this point in mind 
might well prove profitable. 

Although Guillaumin described Jasminum francii in 1933, he placed it 
in synonymy the following year (see Bull. Mus. Hist. Nat. Paris Il. 6: 
458. 1934) after he had seen authentic material of Schlechter’s species. 

When describing Jasminum kriegert, Guillaumin cited two numbers, 
one of which was Vieillard 2938; however, all the specimens of this 
number which | have seen, including one from Paris, are clearly J. leratii. 
This number is presumably a mixed gathering, for Guillaumin (in Bull. 
Mus. Hist. Nat. Paris II. 15: 453, 454. 1943) referred Vieillard 2938 to 
both J. kriegeri and J. leratii, adding “pro parte” to the citation under the 
latter. 


3. Jasminum neocaledonicum Schlechter, Bot. Jahrb. 39: 231. 1906; 
Guillaumin, Ann. Mus. Col. Marseille II. 9: 191. 1911, Not. Syst. 
Paris 3: 65. 1914, et Bull. Mus. Hist. Nat. Paris 25: 652. 1919, ibid. 
II. 2: 169. 1930; Daniker, Viert. Naturf. Ges. Ziirich 78(Beibl. 19): 
366. 1933; Gullioumin, Bull. Mus. Hist. Nat. Paris II. 13: 127, 
1941, ibid. 20: 371. 1948, et Fl. Nouv.-Caléd. 284. 1948, et Bull. 
Mus. Hist. Nat. Paris IT. 27: 475. 1955, ibid. 30: 397. 1958. 


J. pulchrefoliatum Guillaumin, Not. Syst. Paris 3: 62, 65. 1914, et Bull. Mus. 
Hist. Nat. Paris 25: 500. 1919, et in Sarasin & Roux, Nova Caledonia, 
Bot. 1: 207. 1921, et Bull. Mus. Hist. Nat. Paris 27: 560. 1921, zbid. II. 4: 
701. 1932; Daniker, Viert. Naturf. Ges. Ziirich 78(Beibl. 19): 367. 1933; 
Guillaumin, Bull. Mus. Hist. Nat. Paris IIT. 10: 520. 1938, et Fl. Nouv.- 
Caléd. 284. 1948 


Evergreen liane; stem glabrous, even when young. Leaves opposite, 
simple; petioles 6-35 mm. long, glabrous, articulated 4%—'™% from the 
base, occasionally on the largest leaves an additional articulation toward 
t:e top; lamina broadly ovate to very broadly ovate, (4—)6—9.5(-11) cm. 
long by (3- a —8.5(—10) cm. broad; margin entire, not noticeably thickened 
or recurved; apex acute or acuminate, more rarely obtuse or even retuse: 
base pounced. truncate or subcordate, only slightly attenuate into the 
petiole; venation more or less evident, especially above, primary nerves 
(4)5-7 per side. Juflorescences axillary and terminal on side shoots, 
ternately paniculate, (5—)7—50 flowered, glabrous; bracts more or less 
linear subulate, 2-7 mm. long; pedicels 3-10 mm. long. Flowers, white, 
fragrant, heterostylous. Calyx glabrous, tube 2-3 mm. long with 4-6 
subulate-lanceolate teeth, 1-2.5 mm. long, often slightly recurved (Fic. 
1G). Corolla hypocrateriform, tube 20-22 mm. long, lobes 5—7, lanceolate, 
acute, about 13 mm. long. Stamens 2, anthers 4.75-5 mm. long on fila- 
ments 0.25—0.5 mm. long. Ovary 1.25-1.5 mm. long, style about 10 mm. 
long in short-styled flowers and 20 mm. long in long-styled. Fruits (one 
loose fruit only seen, Bawmann-Bodenheim 5581), ellipsoid, 22 mm. long 
by 9 mm. broad. 


1962 | GREEN, STUDIES IN JASMINUM, II 123 


New Caledonia. Bei Oubatche, in den waldern der Berge, c. 900 m. alt., 27 
ee 1902, Schlechter 15586 rel ea B [destroyed, not ea: isotypes, BM, 

, K, Z); entre Bourail et Houailou, montagnes, c. 400 m. alt., 6 Feb. 1912, 
coe 525 (syntype J. ee iii: P [photograph seen]; isosyntype, z); 
Bourail, Balansa 1299 (Pe); Prony, buissons du littorale, Jan. 1915, Franc 1935 
(P); road to Montagnes des Sources, 650 m. alt., 4 March 1955, McKee 2198 
(us); south of Mt. Kongouaouri, mesophilous forest on serpentine, 300 m. alt., 
10 Sept. 1950, Baumann-Bodenheim 5874, 5881 (A, z); Mé Amméri, 700 m 
alt., on serpentine soil, 28 Nov. 1950, Guillaumin & Baumann-Bodenheim 8883 
(z); Mé Aoui, hygrophilous forest on schistaceous soil, 500 m. alt., 7 & 8 Feb. 
1951, Guillaumin & Baumann-Bodenheim 10301, 10440 (a, z). 


A handsome species with large leaves and usually numerous flowers 
to the inflorescence which must be very becutiful as a living plant. Vegeta- 
tively it is sometimes very similar to large-leaved examples of Jasminum 
elatum, but the two may easily be distinguished by their calyces and, 
to a limited extent, by the position of the petiolar articulation (see under 
J. elatum). To judge from some of the specimens examined, there is also 
a tendency for specimens of J. meocaledonicum to blacken on drying. 

In his protologue for this species Schlechter cites his specimen no. 15585 
but this must be a misprint, for 75586 is the number on all the Schlechter 
specimens examined, and, in the same publication (Bot. Jahrb. 39: 224. 
1906), he cites no. 15585 as the type of another of his species: Leucopogon 
septentrionalis. 


4. Jasminum noumeense Schlechter, Bot. Jahrb. 40(Beibl. 92): 33 
1908; Guillaumin, Ann. Mus. Col. Marseille IT. 9: 191, 1911, et 
Not. Syst. Paris 3: 65. 1914, et Bull. Mus. Hist. Nat. Paris IT. 6: 458. 
1934, ibid. 10: 519. 1938, ibid. 15: 454. 1943, et Not. Syst. Paris 11: 
55. 1943 (sphalm. J. noumeanum), et Fl. Nouv.-Caléd. 284. 1948, 
et Mém. Mus. Hist. Nat. Paris II(B). 4: 48. 1953, et Bull. Mus. 
Hist. Nat. Paris IJ. 27: 475. 1955. 


J. bouquetii Jeanneney, Nouv.-Caléd. eae ie oa nom. nud.; Guil- 
laumin, Not. Syst. Paris 11: 55. 1943, 

ne Hiaeeniae Guillaumin, Bull. Mus. Hist. "Nat Paris TL 2: 169. 1930; zbid. 4: 
697. 


Evergreen liane of variable length; stem puberulent or occasionally glab- 
rous. Leaves opposite, simple; petioles 3-17 mm. long, puberulent or 
occasionally glabrous, articulated near the base; lamina more or less 
chartaceous, ovate to very broadly ovate, (2—)3-5(-—6) cm. long by (1.5—) 
2-3.5(—5) cm. broad; margin entire, not thickened, occasionally very 
slightly recurved; apex rounded to obtuse, or retuse, occasionally acute, 
often with an apiculus; base truncate or occasionally rounded or sub- 
cordate, only slightly attenuate into the petiole; venation with primary 
nerves usually visible, sometimes more or less obscure, 3—5 per side. J/n- 
florescences terminal or axillary on side shoots, ternately subcorymbose- 
paniculate, with up to about 20 flowers per panicle, usually pubescent, 

& 


124 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLIII 


especially towards the base of the inflorescence, occasionally glabrous; 
bracts more or less foliaceous, especially the basal ones, and up to 2 cm. 
long, occasionally the upper ones ranging to subulate, 1-2 mm. long; 
pedicels 3-12 mm. long. Flowers white, fragrant, sometimes strongly so 
(McKee 2728) or inodorous (Ddniker 1846), heterostylous. Calyx pubes- 
cent, occasionally glabrous, the tube 2—3(—4) mm. long, with 4 or 5 sub- 
foliaceous teeth 2—5.5 mm. long, often wavy margined (Fic. 1,H), some- 
what accrescent in fruit. Corolla hypocrateriform; tube 24-33 mm. long, 
lobes 5, lanceolate or broadly lanceolate, acute or very acute, 8.5—15 mm. 
long. Stamens 2; anthers (3—)4—5.5(—6) mm. long, on filaments 0.2-2 mm. 
long. Ovary 1-1.25 mm. long; style 10-11 mm. long in short-styled flowers 
or 20-30 mm. long in long-styled. Fruits (Balansa 2232, Virot 842) bi- 
lobed, united for about 24 length of each half, 8-9 mm. long by 7-8 mm. 
broad, occasionally single sided with a lateral projection 2-3 mm. in 
diameter representing the abortive loculus. 


New Caledonia. Magenta (near Nouméa), June 1903, Le Rat 574 (isosyntype, 
p) also, May 1903, Le Rat 588 (isosyntype, Pp); bois de Port Despointes (Nou- 
méa), +50 m. alt., bois secs des collines littorale, schistes nummulitiques, 21 


(a); Kiistenhiigel der Halbinsel Nouméa, in den Buschwaldchen am Abhang, 24 
July 1925, Daniker 1846 (z); Nouméa, Aug. 1926, Franc 2232 (holotype J. 
magentae, P; isotypes, A, NY, US); Nouméa, foot of Ouen Toro, just above beach, 
6 July 1955, McKee 2728 (A, HULL); Anse Vata, remnant of dry forest on low 
calcareous hill, 27 March 1955, McKee 2298 (a, HULL, us). Without locality, 
Franc 845 (A, BM, E, K, NY, P, US); Vieillard 2936 (BM); Pancher 315 (A) and 
Pancher sn. (labeled J. bouquetii, a). 


One of the most distinctive features of this species is the large, almost 
subfoliose, calyx lobes (Ftc. 1,H). The corolla tube, too, is longer than in 
any other New Caledonian jasmine. Vegetatively it can be very similar to 
Jasminum artense; for vegetative characters which can be used to dis- 
tinguish between them see under that species. 

It was recognized as a distinct species as early as 1894 when, without 
description, Jeanneney used the name Jasminum bouquetii. In 1930, Guil- 
laumin published his J. magentae with a short description in French, and 
two years later (in Bull. Mus. Hist. Nat. Paris II. 4: 697. 1932) he amplhi- 
fied the description and published it in Latin. Later (7bid. 6: 458. 1934), 
after having seen syntype material of Schlechter’s species published in 
1908, he placed it in synonymy under J. noumeense, where it undoubtedly 
belongs. 


5. Jasminum artense Montrouzier, Mém. Acad. Lyon 10: 231. 1860; 
Guillaumin. Ann, Mus. Col. Marseille IT. 9: 191. 1911, et Bull. Mus. 
Hist. Nat. Paris 18: 468. 1912, et Not. Syst. Paris 3: 61, 65. 1914; 
Guillaumin & Beauvisage, Ann. Soc. Bot. Lyon 38: 102 (“Species 
Montrouzieranae,’ 28) 1914; Dianiker, Viert. Naturf. Ges. Zurich 78 


1962 | GREEN, STUDIES IN JASMINUM, II 25 


(Beibl. 19): 365. 1933; Guillaumin, Fl. Nouv.-Caléd. 284. 1948, et 
Mém. Mus. Hist. Nat. Paris II(B). 8: 161. 1959 


J. dzumacense Guillaumin, Not. Syst. Paris 3: 63, 65. 1914; Daniker, Viert. 
Naturf. se Zurich 78(Beibl. 19): 366. 1933; Guillaumin, Fl. Nouv.-Caléd. 
284. 1948, et Mém. Mus. Hist. Nat. Paris II(B). 4: 47. 1953, ibid. 8: 161. 
1959. 


J. velutinum Guillaumin, Mém. Mus. Hist. Nat. Paris II(B). 4: 48. 1953, non 
Kobuski (1941); Virot, Vég. Canaque, 175. 1956. 


Evergreen liane; stem glabrous or puberulent, rarely somewhat velutin- 
ous, especially when young. Leaves opposite, simple; petioles 3-20 mm 
long, glabrous or puberulent, rarely velutinous, articulated 4—'% from the 
base, sometimes on larger leaves showing an additional articulation toward 
the top; lamina more or less coriaceous, glabrous or rarely puberulent, 
ovate to orbicular, rarely narrowly ovate, (1.2—)2—4.5(—5.5) cm. long by 
(1—)1.5-4(—5.2) cm. broad; margin entire, slightly thickened, usually re- 
curved; apex rounded, obtuse or even retuse to acute, often with a small 
apiculus; base rounded or subcordate, rarely obtuse; venation with primary 
veins visible, sometimes obscure above, (2)3 or 4 per side, occasionally 
reticulate. Inflorescences axillary and terminal on side shoots, ternately 
paniculate, with 5—7(—15) flowers per panicle, glabrous or puberulent, 
rarely velutinous; bracts subulate, 1-1.5 (—3) mm. long; pedicels 5—20 
mm. long. Flowers white, fragrant, heterostylous. Calyx glabrous or 
puberulent, rarely velutinous; tube 1.5—2.5 mm. long with 4 or 5 shallowly 
triangular teeth 0.1-0.7 mm. long (Fic. 1, C to D). Corolla hypocrateri- 
form; tube 10-20 mm. long with 5—8 lanceolate or narrowly lanceolate, 
acute (to rounded) lobes 8-13 mm. long. Stamens 2; anthers 2.75—3.5 
(—5) mm. long on filaments 0.5—1 mm. long. Ovary 1-1.25 mm. long; style 
about 7 mm. long in short-styled flowers or 9-17 mm. long in long-styled. 
Fruit (Balansa 3629) ovoid-ellipsoid, paired (or single by abortion) 10-11 
mm. long by 7-8 mm. broad. 


New Caledonia. Valleé du Diahot (Dothio), collines eruptive, Jan. 1872, 
Balansa 3629 (syntype J. dzumacense, P; isosyntypes, BM, E, K); . Dzumac, 
Jan. 1909, Le Rat 165 (syntype, J. amen Pp); bords des creeks desecheés, 
base des versants ouest du Mt. Kaala, + 20 m. alt., maquis serpentineux, 10 
Nov. 1943, Virot 1302 (holotype J. velutinum, Pp); im Tale des vom Mt. Hum- 
boldt kommenden Zuflusses der Kalouéhola und an den Abhangen des Tontou- 
tatales, im Serpentingebiisch, 18 Nov. 1924, Daniker 483 (z) and 10 Nov. 1924, 
Daniker 588 (z); am Mt. Bacon am dusseren Abhang des siidlichen Serpentin- 
massivs gegen die Tontoutaebene, im Buschwalde, 15 Nov. 1924, Daniker 700 
(z); Prony, 1890, Bougier (Ly); Ile Art, in montanis et silvis, Sept. & March, 
Montrouzier 143 (holotype, P); slope west of “Baie des Pirogues,”’ on serpen- 
tine soil, 40 m. alt., 16 Dec. 1950. Hiirlimann 355 (a, z), ibid., meso-xeroph- 
ilous shrubby forest on serpentine soil with rocks, 170 m. alt., 17 Dec. 1950, 
Hiirlimann 381 (A, z); range of Mt. Podchoumié (Dumbéa), on rocky serpentine 
ridge, 180 m. alt., 23 July 1951, Hiirlmann 1625 (a, z); Col de Plum (Mt. 
Dore), on serpentine soil, 200 m. alt., 22 Aug. 1950, Baumann-Bodenheim 5577, 
5593A (A, 2), ibid., 8 Nov. 1950, Guillaumin & Baumann-Bodenheim 7879, 7933 


126 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLIII 


z); Col de Vulcain, serpentine bush, 900 m. alt., 11 Nov., 1950, Bauwmann- 
es ee 8178 (A, 2). IsLe or Pines: south of Pic Nga, mesophilous forest on 
serpentine, 29 May 1951, Baumann-Bodenheim 13699 (a, z), and creek on the 
southwest slope of Pic Nga, mesophilous forest, 30 May 1951, Baumann-Boden- 
heim 13786, 13789, 13816 (A, 2) 


Jasminum artense is a distinct species which may perhaps be confused 
vegetatively with J. noumeense, but the leaves of the latter tend to be 
thinner in texture and the articulation of the petiole appears never to be 
more than 2 mm. from the base, even on the longer petioles. In J. artense 
the articulation is usually at least 2 mm. from the base, except in the 
case of very short petioles, where however, it is found at least between one- 
third and one-half of the way from the base of the petiole. When material 
bears an inflorescence there is no difficulty in distinguishing the two 
species, for the calyx lobes are quite distinct: in J. artense short and broadly 
triangular, in J. noumeense long and almost subfoliaceous. 

Jasminum dzumacense was separated from J. artense on the relative 
lengths and positions of the style and stamens, but I cannot separate them 
as species in view of the frequency of heterostyly in the genus. In 1953, 
Guillaumin also distinguished and published the name of J. velutinum, but 
examination of the type (Virot 1302, not 1303 as cited in the protologue) 
shows that in everything but its velutinous indumentum the plant matches 
J. artense. Furthermore, J. artense has been found to be very variable in 
the amount of its pubescence, and Virot 1302 is only the extreme expression 
of this character. Similar treatments of hairy variants as species have oc- 
curred in other species of Jasminum, e.g. J. angulare, J. humile, J. sub- 
humile, and J. floridum (see Verdoorn, Bothalia 6: 560. 1956, and Green, 
Notes Bot, Gard. Edinb. 23: 355. 1961), and in each case an examination 
of more extensive collections than had been available earlier has shown 
that separation at specific rank cannot be justified. 

In addition to the collections cited above, there is a specimen without 
sign of flowers or fruit (south slope of Ouen Toro, near Nouméa, on cal- 
careous soil, 30 Oct. 1950, Guillaumin 7414, |A, z|) which appears very 
similar to Jasminum artense. However it bears orbicular, retuse leaves 
which are especially small, varying from 0.8 cm. to 2 cm. in diameter, and 
was gathered from a plant growing on calcareous soil. All the specimens 
of J. artense of which there is mention of the soil type in their field notes 
are from serpentine soil and it would be interesting to see more material 
of this plant, especially flowering material. 


6. Jasminum linearifolium Guillaumin, Mém. Mus. Hist. Nat. Paris 
II(B). 4: 47. 1953; Virot, Vég. Canaque, 175s 1956. 


Evergreen liane of variable length or diffuse shrub; stems glabrous, 
slender. Leaves opposite, simple; petioles (2—)3—5 mm. long, glabrous, 
passing imperceptibly into the lamina, articulated at approximately the 
middle; lamina linear or very narrowly lanceolate, (3-)3.5-5(—7) cm. 
long by (0.1—)0.2—-0.4(—0.6) cm. broad; margin entire, slightly thickened; 


1962 | GREEN, STUDIES IN JASMINUM, II Lea 
apex acute; base narrowly attenuate into the petiole; venation obscure 
above and below with 3 or 4 primary veins per side. /nflorescences terminal 
on side shoots, ternately branched with about 5 flowers per inflorescence, 
glabrous; bracts subulate, 0.5-2 mm. long; pedicels 5-10 mm. long. Flow- 
ers white. Calyx glabrous: tube 1-1.25 mm. long with 4 or 5 shallowly 
triangular teeth, 0.1-0.2 mm. long (Fic. 1, C). Corolla hypocrateriform: 
tube 16 mm. long with 4 narrowly lanceolate, acute or obtuse lobes 10 mm. 
long. Stamens 2; anthers 3.75 mm. long on filaments 2.5 mm. long. Ovary 
globose (fide Guillaumin); style 5 mm. long (fide Guillaumin) in a short- 
styled flower. Fruit unknown. 


New Caledonia. Sommet du Dome de la Tiébaghi. +600 m. alt., maquis ser- 
pentineux, 27 Oct. 1943, Virot 1275 (holotype. P). 


The specimen upon which this species is based is unique amongst all the 
material examined in this investigation. However, one gathering of 
Jasminum lerati (Guillaumin & Baumann-Bodenheim 11663) bears some 
shoots and leaves which approach (and in their extreme match) in slender- 
ness and narrowness the stem and leaves of J. dinearifolium. But a glance 
at the calyx shows that this species is not immediately related to J. leratii. 
Rather is the relationship with J. artense and, in fact, except for the leaves, 
the two species are identical, and further collections may perhaps produce 
intermediates. Yet the difference between linear leaves and ovate or or- 
bicular ones is so great that, for the present at least, it is felt best to main- 
tain this species, based though it is, upon a single few-flowered specimen 
with only one corolla. 


Jasminum elatum Pancher ex Guillaumin, Not. Syst. Paris 3: 63, 
65. 1914, et Bull. Mus. Hist. Nat. Paris IJ. 1: 217. 1929 et Candollea 
§: 151. 1932; Daniker, Viert. Naturf. Ges. on 78(Beibl. 19): 366. 
1933; Guillaumin, Fl. Nouv.-Caléd. 284. 


J. elatum var. brevistylis Guillaumin, Not. Syst. Paris 3: 64, 1914, et in Sara- 
sin & Roux, Nova Caledonia, Bot. 1: 207. 1921, et Bull. Mus. Hist. Nat. 
Paris IT. 15: 453. 1943. 


Evergreen climbing shrub; stems glabrous or puberulent, often minutely 
so and especially when young. Leaves opposite, simple; petioles 3-22 mm. 
long, glabrous or puberulent, articulated near the base; lamina more or 
less chartaceous, narrowly ovate to broadly ovate, or even very broadly so, 
(2.5—)4-8(-10) cm. long by (1.2—)2—7(-—7.5) cm. broad, margin entire, 
not usually recurved or noticeably thickened; apex obtuse or acute, or 
more rarely slightly retuse, often with an apiculus; base truncate, rounded, 
obtuse or cuneate, often more or less attenuate into the petiole and rarely 
almost subcordate; venation usually with primary nerves only visible above 
and below, 4-7 per side. /zflorescences axillary or terminal on side shoots, 
dense, ternately subcorymbose, many flowered, glabrous to densely and 
minutely puberulent; bracts subulate, narrowly oblanceolate or elliptic, 


128 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


the lower ones subfoliaceous, 1-6 mm. long; pedicels short, 0-2 mm. long. 
Flowers white, fragrant, heterostylous. Calyx glabrous and ciliolate to 
densely and minutely puberulent; tube 1.5—2.5 mm. long with 4-6 stout, 
blunt, teeth 0.1-1 mm. long, which in the extreme are aan flattened 
laterally (Fic. 1, A & B, occasionally almost to C). Corolla hypocrateri- 
form; tube 8-21 mm. long; lobes 4—6, broadly ovate or ovate, rounded or 
acute, 4—-7.5 mm. long. Stamens 2; anthers (2.5—)3-4 mm. long on fila- 
ments 0.2-0.5 mm. long. Ovary 0.75-1.25 mm. long; style 3-6.5 mm. 
long in short-styled flowers or 12-22 mm. long in long-styled. Fruit 
(Balansa 1298) ovoid, paired (or single by abortion), 9-10 mm. long by 
7-8 mm. broad. 


New Caledonia. Environs de Nouméa, bosquets, Sept. 1868, Balansa 578 (syn- 
type, P; isosyntype, BM, NY); Port de France [=Nouméa], Vieillard 907 (syn- 
type var. brevistylis, P; isosyntypes, A, G); Kanala, Vieillard 908 (syntype var. 
brevistylis, ep); Mt. Canala, 650 m., Nov. 1911, Sarasin 363 (z); au Said de 
Canala, foréts, vers 1,000 m. alt., Nov. 1869, Balansa 1702 (syntype var. 
brevistylis, Pp); prés de Koe, bords de la Dumbéa, 14 Dec. 1869, Balansa 1298 
syntype, P; isosyntype, K); bei La Foa an der Briicke iiber das Fliisschen La 
Foa, im sneer 27 Sept. 1924, Ddniker 131 (z); Gatope, Vieillard 2929 
(BM, kK); northwest slope of Mt. Mou, forest, 15 Oct. 1950, Guillaumin & 
Baumann- Bodbuhem 6958 (A, z); Mé anita, on serpentine soil, 700 m. alt., 
28 & 30 Nov. 1950, Guillaumin & Baumann- Bodenheim 8770, 9066 (A, 2); 
above the “Ermitage” valley (Mt. Koghi), hygrophilous forest on serpentine, 
400 m. alt., 4 July 1951, Baumann-Bodenheim 14479 (a, z); without locality, 
Pancher 316 (=Vieillard 2930) (syntype var. brevistylis, P; isosyntype, BM), 

massifs sur les sols argilo-schisteux, March, Pancher 313 es Vieillard 2941) 
(lectotype, P 

Loyalty islands: Lirou: Képéneéhé, im Buschwald in Kustennahe, 22 Oct. 1925, 
Daniker 2270 (z); Thosip, 10 Nov. 1925, Daniker 2270a (z); without locality 
Deplanche 82 (syntype, P). 


Jasminum elatum is distinguished amongst the simple-leaved New Cale- 
donian species by its sessile or almost sessile flowers. The apparent ‘“‘flat- 
ness” of the inflorescence branches and the size of the upper bracts are 
also distinctive. In length of calyx lobes it is similar to J. artense, but the 
lobes of the two species are nevertheless different: in J. artense they are 
flat, broadly triangular and acute, or at least with an angular apex, but in 
J. elatum they are blunt and thick, or even somewhat flattened laterally in 
those with the largest teeth. In addition, although the corolla tube in 
these two species is the same length, the lobes in J. edatum are shorter than 
those in J. artense. Large-leaved vegetative specimens of J. elatum may be 
confused with J. neocaledonicum, but in the former the petiolar articula- 
tion is usually found more or less towards the base and in the latter well 
up the petiole. Some purely vegetative specimens with large leaves can 
be difficult to identify and‘a good example is Guillaumin & Baumann- 
Bodenheim 8732 (Mé Amméri, 700 m. alt., on Serpentine soil, 28 Nov. 
1950, z) which is perhaps best left as “cf. elat 

With his original description Guillaumin ee five collections; it has 


1962 | GREEN, STUDIES IN JASMINUM, II 129 


been decided to select as lectotype the specimen of Pancher 313 at Paris: 
it is the only Pancher specimen amongst the syntypes, and on the label 
with the Paris specimen he has written ‘Jasminum elatum — de la hau- 
teur de la tige.”’ 


8. Jasminum promunturianum Daniker, Viert. Naturf. Ges. Zurich 
78(Beibl. 19): 367. 1933; Guillaumin, Bull. Mus. Hist. Nat. Paris II. 
14: 456. 1942, ibid. 15: 454. 1943, ibid. 20: 371. 1948, et Fl. Nouv.- 
Caléd. 284, 1948. 


Evergreen shrub; stem puberulent. Leaves opposite, simple; petioles 1-2 
mm. long, puberulent, articulated at the top; lamina thickish, oblanceolate 
or narrowly oblanceolate, (1—)1.5-2.5 cm. long by (0.2—)0.4—0.6(—0.7) 
cm. broad; margin entire, strongly recurved; apex retuse; base very nar- 
rowly cuneate, attenuate to the petiole articulation; venation more or less 
obscure, especially above, with 2 or 3 primary nerves per side. /nflores- 
cences terminal on side shoots, single flowered; peduncle 1—1.5 mm. long, 
glabrous. Flowers white, ? heterostylous. Calyx glabrous; tube 2 = mm. 
long with 5 or 6 stout, blunt, ciliolate teeth, 0.6-1.1 mm. long (Fic. 1, A). 
Corolla hypocrateriform: tube 20 mm. long: lobes 5, broadly lanceolate, 
acute, 7 mm. long. Stamens 2; anthers 4.25 mm. fore on filaments 0.25 
mm. long. Ovary 1 mm. long; style 7 mm. long in a short-styled flower. 
Fruit unknown. 


w Caledonia. Insel Art der Belepgruppe, an den Kiiste, niedrigen Gebtischen 
nee und exponierter Nordoststeilhang, 8 May 1925, Déniker 1681 (holotype, 
z). 


Vegetatively this species, with its small oblanceolate leaves, is very dis- 
tinct. However, the calyx is extremely similar to that of Jasminum elatum, 
which itself is quite distinct from the other New Caledonian species in this 
respect. The field notes which accompany the one and only specimen state 
that it was collected on a rocky and exposed steep northeast slope on the 
coast, and it seems possible that J. promunturianum is really only a habitat 
form of J. elatum. Field observations and additional material may prove 
that the two ought to be united, but, since as yet I have seen no inter- 
mediate material, it seems best to maintain them for the present as separate 
species. 


9, Jasminum kriegeri Guillaumin, Bull. Mus. Hist. Nat. Paris II. 15: 
453. 1943, et Fl. Nouv.-Caléd. 284. 1948. 


Evergreen; stem glabrous. Leaves opposite, simple; petioles 6-12 mm. 
long, glabrous, articulated 14-2 from the base; lamina thickish, narrowly 
lanceolate, 3. ae 5.5 cm. long by 0.6—1.2 cm. broad; margin entire, slightly 
thickened; apex obtuse or acute, subapiculate; base attenuate- obtuse, 
somewhat decurrent onto the petiole: venation clear and raised above and 
below, 2 (or 3) primary veins each side from near the base, the upper two 


iE) JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


prominent, running the length of the lamina and anastomosing, joined by 
3 or 4 (5) slightly less prominent, short veins per side from the midrib, 
occasionally 1 or 2 of these being prominent. Juflorescences terminal on 
side shoots, ternately corymbose-paniculate, 5—12 flowers per inflorescence, 
glabrous; bracts subulate, 1-2 mm. long; pedicels 8-12 mm. long. Calyx 
glabrous, tube 4.5—5 mm. long (at least in dried material with about 20 
longitudinal raised lines), with 5 acute or obtuse teeth, shallowly triangu- 
lar, 0.2-0.5 mm. long (Fic. 1, FE). Corolia hy pocrateriform:; tube (10—)2 
mm. long with 5 or 6 ia. acute, or obtuse lobes 11-13 mm. long. 
Stamens 2; anthers 4.5 mm, long, subsessile. Ovary 1.25 mm. long; style 
18 mm. long in the long-styled flower examined. Fruit unknown. 


New Caledonia. Gatope. leg. M. Krieger, Vieillard 2937 (lectotype, P). 


Jasminum kriegeri, although based on a scrappy specimen collected in 
y an infantry lieutenant named Krieger, was not described until 

1943, and, despite the many collections that have been made in New 
Caledonia during the last forty years, the original, rather poor gathering 
is still the only example of this species in the herbaria I have examined. 

When describing Jasminum kriegeri, Guillaumin cited two specimens, 
Vieillard 2937 and 2938, but all the material of the latter that I have seen, 
including one from Paris, is undoubtedly J. leratii. The specimen of the 
former number at Paris, with the name written in Guillaumin’s hand and 
collected by Krieger, is the only one which the original description fits, and 
it is therefore selected as the lectotype. 

The calyx is perhaps the most noticeably distinct feature of this species. 
In none of the other species is the calyx tube normally longer than 3 mm, 
at the most, although rarely in Jasminum noumeense it reaches 4 mm. but 
with large subfoliaceous lobes on top. Usually in the New eee jas- 
mines it is only 1.5—2.5 mm. long, whereas in J. kriegeri it is 4.5—5 mm. in 
length and contrasts with the very small comparatively obscure ey The 
calyces on the dried specimen which constitutes the type are furrowed 
throughout their length by shallow grooves which presumably correspond 
to the spaces between the veins. They are never so noticeable in the other 
species, and it is suspected that they appear only on drying. 

The other part of this specimen which is noticeably distinct is the 
leaves. In shape they are very similar to a narrow-leaved expression of 
Jasminum leratii, but the venation is more or less unique amongst the New 
Caledonian species. In several species the main lateral and basal nerve 
runs the length of the lamina near the margin and joins up with the other 
lateral nerves given off by the midrib, but in this species two prominent 
nerves run the length of the lamina and the nerves joining them are fewer 
in number and more obscure. Nevertheless, this difference is only one of 
degree, and within the genus a whole range can be found from a regularly 
pinnately nerved leaf to the extreme as exhibited, for example. by J. crassi- 
folium Blume where the only two prominent lateral nerves run the length 
of the leaf, one on each side of the midrib. 


1962] 


GREEN, STUDIES IN JASMINUM, II 


131 


None of the intact flowers on the type specimen are quite mature, and it 
is feared that the measurements of the corolla given in the description 
above may be too short, for, judging from specimens of other species, it is 
believed that the corolla tube, for example, lengthens right up to the 


time of anthesis. 
point.) 


(Observations on living plants would perhaps settle this 


INDEX TO EXSICCATAE 


The list is arranged alphabetically by the last name of the collector. 
Numbers in parentheses refer to the corresponding species in the text. 


Bailey 9623 (2 
Balansa 578, 1298 (7); 1299 (3); 1702 
(7); 2785, 3189 (1); 3629 (5) 
Baumann- Bode nheim 5032 
5593A (5); 5874, 5881 (3); 6057 rae 
8285 (2); 13436, 13454, 
13608 (1); 13642 (2); 13699, 13786, 
13789, 13816 (53); 14479 (7); 14529, 
14721 2 14770 1 


Gi) 
mae 131 ae 483, 588, 700 (5); 770, 
1335 (2); 1436 (1); 
1681 (8); 1717 (1); 


1846 (4); 1886, 1909, 2269 (2); 2270 
B270G Ce pees 2388a (2); 2465, 
2465a (1); 2626, 3088, 3089 (2) 

Deplanche 82 

Eastwood, s.n. (2 

Franc 845 (4); 1021 (2); 1377 (1); 


> 1906 (2); 1935 (3); 
(1) 


Guillaumin 7414 (under 5); 13395 (1) 

Guillaumin & Baumann-Bodenheim 6842 
Gre 6958 CI) 7879, 7933 (5); 8732 
(under 7); 8770 (7); 8883 (3); 9066 


(7) ; 9514, 9665 (2); 10301, 10440 (3); 
11106, 11135, 11161, 11663, 12403 (2); 
13127- i) 

Heim 49 (2); 74 (1) 

Hiirlimann 355, 381 (5); 769 (2); 1092, 
1166 CLV S 1200, Oe 150711) = 1625 
(3 

Le Rat 709 (1); 165 (5); 172 (2); 574 
588 (4) 

McKee 475, 1973 (2); 2198 (3); 2251 
(2) = 2208 (Ay 2342) 2428" (1); 
2728 (4); 6512 (1) 

Montrouzier 143 (5) 

Pancher 373 (7); 315 (4); 316 (7); 

( 


4) 

Sarasin 185 (2); 363 (7); oe Q)525 
(37307 Gl) 85a (2) | 2890 eCL) 

Schlechter 15586 (3) 

Vieillard 907, 908 (7); 911 (2); 913 (1); 


2929, 2930 (7); 2935 (1); 2935 (2); 
2936 (1); 2936 (4); 2937 (9); 2938 
Cyee2047 (7) 


Virot 442 (2); 842 (4); 846 (1); 865 
(Gee 870; 88). (2); 8821). 922, 92 
O20 mn (Oneweire (6): 1302 “(5)) 

Walter 65 (2) 

Woodcock 630 (2) 


~ 


132 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


A CYTOLOGICAL STUDY OF THE GENUS VIBURNUM ! 


DoNALD R. EGoLr 


THE GENUS VIBURNUM includes some 250 species from diverse habitats 
in Asia, Europe, North America, and South America. Though viburnums 
are varied in form, the ideal ornamental that combines fragrance, colored 
flowers, small stature, evergreen or brilliantly colored foliage, and luxuriant 
fruit does not exist. The present cytological study was initiated to estab- 
lish a basis for genetical research that might yield interspecific hybrids 
combining in one plant the ornamental characteristics of several species. 
Such a study of chromosomes is obligatory for the plant breeder to initiate 
and pursue intelligently an interspecific hybridization program. Many 
Viburnum species and varieties involve complexes that have not been 
adequately covered by any comprehensive taxonomic treatment. The data 
obtained from such related disciplines as cytology and genetics when 
combined with taxonomy may help to resolve the species complexes and 
clarify the classification of the genus 

To the extent that this publication is a portion of more extensive cyto- 
genetical and cytotaxonomical studies in progress, only that portion of the 
research concerned with cytology is reported. Although the literature has 
been frequently consulted to verify the identification and relationships of 
the taxa studied, this paper is not intended as a taxonomic study. The 
author has followed the taxonomic nomenclature of Rehder’s Manual of 
Cultivated Trees and Shrubs (30) and Bibliography of Cultivated Trees 
and Shrubs (31). 


REVIEW OF LITERATURE 


All the chromosome counts reported by various authors are incorporated 
under the respective species in Tapie I. The earliest cytological study 
of Viburnum was that of Sax and Kribs (32), who reported that eleven 
species had a gametic chromosome number of nine (7 = 9). The Asiatic 


*This study includes a portion of the research completed for the Ph.D. thesis in 
the Departments of Plant Breeding and Floriculture, Cornell University, Ithaca, N. Y., 
and later work in the United Kingdom and at the U. S. National Arboretum, Washing- 
ton, D.C. 

Th e author is indebted to all who have co-operated in supplying seeds, cuttings, or 


for a Fulbright Scholarship in the United Kingdom, 1956-58. The many courtesies 
extended in the United Kingdom by the Royal Botanic Garden, Edinburgh; the 
British Museum (Natural History), London; the Royal Botanic Gardens, Kew; 
the Royal Horticultural Society’s Gardens, Wisley; the John Innes Institute, Bayford- 
bury; and the University of London are gratefully acknowledged. 


1962] EGOLF, CYTOLOGY OF VIBURNUM 133 


and American species studied included six of the nine taxonomic sections 
of Rehder (30). Sax and Kribs stated that the chromosomes are large, 
have an affinity for chromosomal stains, and consequently provide favor- 
able material for study. A second basic chromosome number of eight 
(n = 8) was reported for V. fragrans by Simonet and Miedzyrzecki (35), 
who also published counts for seven additional species as n = 9. 
gametic and somatic number of V. tinus was determined by Feng (15) to 
be n = 18 and 2m = 36. Sugiura (38) reported 2” = 20 for V. awabuki 
(syn. V. odoratissimum). In 1946, Poucques (27) listed the gametic 
chromosome counts for five species, four of which were previously un- 
published; in a later publication (28) he listed two additional species. 

Janaki Ammal determined the chromosome number 2m = 16 for Vi- 
burnum fragrans and V. grandiflorum, and for V. x bodnantense, a 
hybrid produced from a cross between these species. The chromosomes 
of the two species paired normally in the hybrid, and the pollen fertility 
was as high as one hundred per cent (36). The extensive cytological 
study of Janaki Ammal (18) included thirty-seven determinations, of 
which twenty-one were reported for the first time. Her survey reported 
somatic chromosome numbers for species in cultivation at the Royal Horti- 
cultural Society’s Garden, Wisley, England; the Royal Botanic Gardens, 
Kew, England; and the Jardin des Plantes, Paris, France. Seventeen of 
these counts are at variance with the somatic chromosome numbers of 
the present study; while three are at variance with previous reports. 
These differences are considered in the discussion. She has interpreted 
the 2n = 18 of the hybrid V. & juddii (V. carlesii, 2n = 20, X V 
bitchiuense, 2n = 16) as a synthesis of a plant with » = 9, and in addi- 
tion, proposed that V. carlest (2n = 20) arose as a backcross between a 
chance triploid (2m = 24) and the normal diploid (2m = 16) of V. 
bitchinuense. This was considered to be an example of the possible manner 
of evolution of Viburnum species in nature, which finally resulted in a 
large number of species with the basic number = 9. A chromosome count 
of V. lobophyllum, 2n = 20, determined by Enoch for a plant grown at 
Exbury, Southampton, England, was included in the publication of Janaki 
Ammal (18). 

The cytological study by Thomas (40) included twenty-nine of the 
plants cultivated in the Arnold Arboretum. A few of these counts were 
obtained from root tips, but most were made from pollen mother cells; 
however, only partial designation is given as to which counts were gametic. 
Thomas concurs with the author that the origin of species with m = 9 as 
postulated by Janaki Ammal (18) is questionable. He states that it is 
more likely that species with basic chromosome numbers of eight and 
ten originated from species with a base number of nine by the loss or 
gain of a chromosome. His study indicates that translocations occur 
rather frequently, as evidenced by bridge formations observed in several 
of the species. He noted a relatively high percentage of aberrations in 
Viburnum cassinoides, V. carlesii, V. dentatum, V. plicatum, Ve. xX rhy- 
tidophylloides, V. sieboldii, V. trilobum {. compactum, and V. veitchit. 


134 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


MATERIALS AND METHODS 


The seventy-seven species, sixty-one varieties and forms, thirteen hy- 
brids, and two unidentified accessions of Viburnum investigated are a 
sented with their sources, in Taste TI. Plant material was obtained < 
seed, cuttings, or plants from native habitats, botanic gardens, cho 
and estate gardens throughout the world, exclusive of Central ay South 
America. In all cases an effort was made to secure representatives o 
each taxon from three or more sources to provide a check on identification 
and chromosome counts of each taxon. Since commercial nurseries often 
propagate horticultural forms of Viburnum by grafting, all plant material 
or cytological study has been propagated from cuttings or from seed 
collected when possible in the native habitat of the species. Seed from 
native habitats have been used chiefly because seed from botanical col- 


in the collection. The plants for cytological study were maintained during 
the summer months in frames or in a lath house. The remainder of the 
year the plants were grown under long-day conditions of twenty to 
twenty-two hours of light in a 70° F. greenhouse. Under these conditions 
it was possible to keep the plants actively vegetative and to avoid any 
dormant period. 

In so far as feasible, plants propagated from seed and cuttings, and 
representatives of all sources, will be maintained at the Cornell Planta- 
tions and the U. S. National Arboretum for further study. Herbarium 
specimens, which are identified with the code accession numbers, were 
prepared for each collection that provided sufficient material. These are 
maintained as part of the permanent record and will be supplemented 
with flowering and fruiting material when the plants mature. Original 
descriptions and many of the type specimens have been studied to verify 
identifications. Photographs of type specimens and photostats of pertinent 
taxonomic literature were prepared to provide a basis for cytotaxonomic 
research. The identification of many of the plants previously studied (14 
has been checked since they flowered and fruited. In those cases in which 
a positive determination was made, the alteration has been entered on 
Tas_LE I. However, the documentation numbers have not been altered 
and will be the same as in the previous list (14). 

The root-tip smear technique was used exclusively in this eae 7. 
liminary use of McClintock’s permanent aceto-carmine (19) a La 
Cour’s (20) acetic-orcein stain techniques revealed that the ee gave 
best results. In a portion of the early work the root tips were pretreated 
in aqueous paradichlorobenzene (22), fixed in Baldwin’s modified Carnoy’s 
(2), hydrolized in a solution of equal parts of 95% alcohol and con- 
centrated hydrochloric acid, and smeared in acetic-orcein on the slide. 
After trial of numerous schedules and variations of these procedures a 
modification of La Cour’s (20) technique was employed. Three- to five- 
millimeter-long root tips were pretreated in aqueous paradichlorobenzene 
for one to two hours. The root tips were placed in a watch glass contain- 


1962] EGOLF, CYTOLOGY OF VIBURNUM 135 


ing one part 1.0 N. hydrochloric acid to nine parts 0.5% acetic-orcein 
stain. The watch glass with root tips was passed two or three times over 
the flame of an alcohol lamp to heat the mixture, but great care was taken 
to keep the solution under the boiling point. After the heated watch glass 
had cooled for several minutes, a root tip was placed in a drop of 0.5% 
acetic-orcein on a slide, smeared, and the cover slip applied. 

The Feulgen technique (10) has been used in recent studies with ex- 
cellent results. Root tips were collected and pretreated with 0.1% col- 
chicine for two hours and fixed in La Cour’s 2BD general fixative for 
twelve hours. The root tips were thoroughly washed with distilled water 
before immersion in a peroxide-oxalate bleach consisting of equal parts 
of ammonium oxalate in distilled water and hydrogen peroxide, and placed 
in direct sunlight or under a spot light for five minutes. After the root 
tips were washed again, they were hydrolized in 1.0 N. hydrochloric acid 

t 60° C. for twelve minutes, stained in leuco-basic fuchsin for 2 hours, 
and smeared. The edges of the cover slip were sealed with a mixture of 
gum mastic and paraffin in equal parts. 

Slides were observed immediately or stored in a 40° F. refrigerator. 
After critical examination of the temporary smears was completed, camera 
lucida drawings made, and photomicrographs taken, selected slides were 
made permanent. The method of Conger and Fairchild (6) accomplishes 
the separation of the cover slip from the slide by freezing on dry ice. 
More recently, compressed carbon dioxide has been utilized for freezing 
slides to separate the cover slip from the slide. Immediately before thaw- 
ing, the separated frozen slide and cover slip are placed in 95% alcohol 
which contains 5—-10% acetic acid. After two or three minutes they are 
placed in absolute alcohol for another few minutes before mounting in 
diaphane. The permanent preparations made by this method are almost 
always equal in excellence and clarity to temporary slides and are superior 
for photomicrographs. Permanent slides of virtually all accessions here 
reported have been prepared. 

Critical examination of the preparations was made with a binocular 
pean equipped with 98 & fluorite objectives, N.A. 1.30, and 12.5 
oculars. A minimum of ten countable cells was located before ascertaining 
the ee of chromosomes in the somatic complement. Drawings were 
made with a camera lucida at table level, using 15 % oculars, giving the 
drawing a magnification of approximately 2400 x. In addition to the 
drawings, photomicrographs were taken on 35 mm. microfilm at a mag- 
nification of approximately 1500 x 


RESULTS 


The somatic chromosome counts of the 153 taxa of Viburnum included 
in this investigation are presented in TasiEe I. The species are arranged 
alphabetically within the taxonomic sections, as designated by Rehder (31). 
Under each taxon the chromosome counts published by other authors 
precede those from this study. The general geographic distribution and 


136 JOURNAL OF THE ARNOLD ARBORETUM [VOL, XLIII 


ey «s 


“~ 


Fics. 1-6. Photomicrographs of chromosomes of Viburnum to show varia- 
tions in chromosome complements, approximately & 1100. 1, Viburnum erubes- 
cens, 2n = 32; 2, V. lobophyllum, 2n = 18; 3, V. sieboldii, 2n = 32; 4, V. 


carlesii, 2n = 18; 5, V. bracteatum, 2n = 72; 6, V. scabrellum, 2n = 72. 


138 


JOURNAL OF THE ARNOLD ARBORETUM 


[ VOL. XLIII 


the source of material are given for each accession. A series of photomicro- 
graphs (Ficures 1-15) illustrates variations in the chromosome comple- 
ment. 

In order to make the table more concise, the sources of material have 
been abbreviated as follows: 


AA 


Arnold ma Jamaica Plain 
30, Mas 

Arboretum des 
rance 

Arturo Anealogs Bologna, Italy 


Barres, Loiret, 


Armstrong Nursery Ontanio. 
Calif. 

Botanisches Museum,  Berlin- 
Dahlem, Germany. 


as Botanic Garden, Brook- 
Y. 


lyn 

Birr Cae Birr, County Kings, 
Trela 

Collected near Bedford, Pa. 

Bodnant Garden, Tal y-Cain, 
Denbighshire, N. Wal 


Borde ill, Haywards Hea, 
Sussex, En 

U_S. Botanic cen Washing- 
on, D. 

Collected ak Brighton, Sussex, 
Englanc 

Boyce eae Arboretum, 
Yonkers Y. 

Botanic Gz aven of the University 
of Copenhaget Copenhagen, 
Denmark. 

eee Castle, St. Austell, 
Cornwall, England. 

Col lected near Chattanooga, 
Tenn, 

R. Chenault, Orleans, France. 


W. B. Clarke & Co., San Jose 3, 
Calif. 

Crathes Castle, Crathes, Kincard- 
shire, Scotland. 

Carnell oo Ithaca, N. Y. 

Dominion Arbor Ottawa, 
O 


ntario, ee 
Collected near South Downs, 
Engla “ 
T. Dawson, es tucket, Mass. 
National ene c Gardens, Glas- 
nevin, Dublin, Treland 
Royal oe Ps rden, Edin- 


iques, Geneva, ae land. 


GA 


Gardens of the Blue Ridge, Ash- 
ord, N. C. 

Goteborgs Sear Tradgard, 
Goteborg, Swed 

The Great Park, Windsor, Berk- 
shire, England. 

N. G. Hadden, West Porlock, 


Somerset, Ruckint 
Hattori Botan ical Laboratory, 
Nichinan, Miyasaki, Japan. 


Headford Court, 
Meath, Ireland. 

Henry Foundation for Botanical 
Research, Gladwyne 

Hermann A. Hesse, Weener, Ger- 


Kells, County 


Hong Kong Dent. of Agriculture, 


Highland i. Rochester, N. Y. 
Collingwood Ingram, Cranbrook, 
Kent, Englan 
George Jackman & Son, Woking, 
d. 


Innes Institute, Bayford- 
. Hertfordshire, England. 
Jackson and Perkins Co., New- 

5 Ns ¥ 

Botanie Gardens, Kew, 
\ and. 
Botanic Gardens of Indonesia 
oe Raya), Bogor, Indo- 


nesi 

Teen Kohankie and Son, Paines- 
ville, Ohio. 

iGnesvi lle Nurseries, 
Md. 

Kornik Gardens, Kornik, Poland. 

Collected — Lake 
Cayuga, Ithaca, N. 

V. Lemoine & Fils, aoa France. 


Kingsville, 


Linn agen Iunsevies Center 
Point, 
Los ‘State & County 


a, Calif. 
Lu Shan ii Garden, Kiu- 
kiang, Chin 


EGOLF, CYTOLOGY OF VIBURNUM 139 


Grier peaeueae Wimborne, Dor- 
set, Englan 

W. ies ae Chapel Hill, N.C. 

McLean Bog, McLea 

i ie Philadelphia 


Morton eee Lisle, III. 

Myddelton House, Enfield, Mid- 
dlesex, England. 

Nanking Botanic Garden, Nan- 
king, Kiangsu, China. 

R. C. Notcutt, Ltd., Woodbridge, 
Suffolk, England 

Nymans Gardens, Haywards 
Heath, Sussex, England. 

New York Botanical Garden, 

enV. 
Onarga Nursery Co, Onarga, Ill. 
Muséum d’Histoire 


nce. 
University, 


Pa. 
.S. Plant Tatreduction Garden, 
ee Dale 
Princeton Nurseries, Princeton, 
N 


Unknown Source, Warsaw, Po- 
nd 


and. 
Regel & Kesselring, Rome, Italy. 


Ringwood Forest, Ithaca, New 

York. 

Royal Horticultural Society's 
Eng- 


ere Wisley, Surrey, 
and, 


‘ Ruys, 
Holland 


Ltd., Dedemsvaart, 


Siebenthaler Co., Dayton 5, Ohio. 


ollected South Hill, Ithaca, 
N. Y. 
L. Spath, Berlin, Germany. 
Bergius Botanic Garden, Stock- 


holm, Sweden. 

Swyncombe House, Oxford, Eng- 
land. 

Taiwan Forest eee rae 
Taipei, Taiwan, 

Taiwan Pineapple ae 
Taiwan, China 

Tokyo Uni eae eae Gar- 
dens, Tckvo 

Univ. of California SBemhical 

Garden, Berkeley, Calif. 


Taipei, 


Univ. of Washington Arboretum, 
Seattle, Wash. 

Vaughan Nursery, Chicago 6, 
Illinois. 

Arnold Vogt, Erlenbach-Zurich, 
Switzerland. 

ae Place, Ardingly, Sus- 
Sex and. 


’ 


Wayside Teen Mentor, Ohio. 


Wildlife Research Laboratory, 
ee Sar 
c L. Williams, Exeter, N. H. 


Willows ood Far Gia ctone: 


womans Garden Centers, Inc., 
Framingham, Mass. 

Wyoming Nurseries, Cincinnati 
ike 


140 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


TABLE I. Chromosome Numbers of Viburnum 


SOURCE GENERAL 
AuTHOoR- DocuMEN- OF DISTRIBU- 
SPECIES nm  ITy* TATION MATERIAL ° 
Sect. I. Thyrsosma (Raf.) Rehd. 
V. X bodnantense Aberc. (V. 
fragrans < grandiflorum) 16 9 76E CL cult 
= 9 874E UW539-S0(HL) ¢ ds 
- 16 9 C02E RH as 
7 16 9 1007E K660—48 (BG) Re 
ne 16 9 1102E HL . 
‘Dawn’ 16 6* — —- cult 
a 16 ge  1357E BG “ 
‘Deben’ 16 g* 1937E NN cult. 
V. erubescens Wall. 48 6 — — Himal. 
32 9 972E AA7602-A ‘ 
* 2 9 1149E K ' 

7 32 9* 170K EX is 
var. gracilipes Rehd. 32 9 100K HP2158 Himal. 
e 32 9 502E RU e 
" 32 9 1092K HL et 
““ 32 g* 1594E DU183P er 

V. foetens Decne. 16 6 — = Himal 
as 16 9 441E RH e 
16 9 1061E K as 
M 16 9 1091E HL a 
. 16 Q* 1571E HA n 
V. fragrans Bge. 16 2 — — n. China 
i. 16 6 HI 
“6 16 9 25E CU 7s 
16 9 27E PI-82380 a 
as 16 9 174E AA11588(K) iE 
“ 16 9 597E MT1007-39 
s 16 9 1074E RH a 
“ 16 9*  1154E HA «“ 
var. album Kriiss, 32 6 — — n. China 
fas var. candidissimum) 16 8 AAS55-50-B AAS55-50-B n. China 
16 9 250E 35-38 uf 
7 16 9 598E MT235-38(VN) « 
7 16 9 896E RH fs 
" 16 g* 1419E MY . 


*1, Sax and Kribs (1930); 2, aa and eae (1932); 3, Feng (1934) ; 
Sipiura (1936); 5, Poucques (1946) ; , tbid. (1949); 6, Janaki Renae (1953); 6*, ibid. 
(1950) ; ; ea (1953); 8, Thomas nae 9, Egolf (1956) ; 9*, ibid., reported here for 
the past 

he abe listed in this column are the accession numbers and voucher herbarium 
specimen numbers for the plants studied. Specimens of others are given as “AA” (Arnold 
oo and “K” (Kew). 
. mber or designation ‘after the code letters identifies specific plants. The abbre- 
aaa ye Bt ” signifies that the accession was obtained as seed. 
e of material reference in parentheses refers ie the original source from which 
the pare: or plants were procured. 


4, 


1962] EGOLF, CYTOLOGY OF VIBURNUM 141 


TABLE I (Continued) 


SOURCE GENERAL 
AUTHOR- DocUMEN- OF DISTRIBU- 
SPECIES TATION MATERIAL TION 
var. nanum Boom 16 9 431E UW198-41 cult. 
sf 16 9 504E “ 
te 16 9 600E MT685-50(CL) = 
‘Roseum’ 16 9 438E RH cult. 
a 16 9* 1422E JG cs 
V. grandiflorum Wall. 16 6x — — Himal. 
ae 16 9 1022E K S 
as 16 9 1082E RH 
“ 16 9 1093E HL oi 
« 16 gx  1426E GP ‘i 
“ 16 ox  1427E G 2 
“ 16 gx 1428E E3025 (Cooper) “ 
z 16 9*  1576E IN “ 
V. henryi Hemsl. 48 6 — — c. China 
ef 32 9 984E Cc ry 
ce 32 9 1039E K 5 
“ 32 9 1069E DU a 
te 32 9 1097E HL * 
ce 32 g* 1168E E i 
st 32 9* 1175E ss 
ae 32 g* 1435E BH1398 sf 
es 32 g* 1441E DU10-39 (Henry) ss 
V. X hillieri Stern (V. henryi & 
erubescens) ‘Winton’ 32 gk 1442E HL cult. 
V. odoratissimum Ker-Gawl. (as 
j (0) 4 — — Malaysia 
: 40 6 —_ — i. 
ce 32 9 68E CU (Wash.) " 
fs 32 9 119E FN ‘ 
“ 40 9 293K UW g 
MY 32 9 371E LO < 
“ 40 9 392E NY (BO) ‘: 
a 32 9 427E ‘ 
sf 32 9 919E HL : 
te 32 9 983E © g 
‘ 32 9 1011E K i 
V. photinioides Fashiro 32 9 691E KB Malaysia 
V. sieboldii Miq. 16 6 2454K. RH Japan 
e 32 8 AA616-6-B AA616-6-B ms 
ss 32 9 62E E ne 
: 32 9 105E HP2161 
_ 16 9 270E SN > 
i 32 9 656E MT159-38(KH) " 
. 32 9 718E R2143 % 
ss 32 9 839E AA616-6-B “ 
- 32 9 903E H ne 
- 32 9 1120E P “ 
f. reticulatum Rehd. 32 9 657E MT281-51(AB) Japan 
t 32 9 878E ‘ 
~ 32 9* 1056E K986-36(LE) as 


142 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 
TaBLe I (Continued) 
SOURCE GENERAL 
AuTHOoR- DoCUMEN- OF DIstTRIBU- 
SPECIES 2n TATI MATERIAL TION 
V. suspensum Lindl. (as V. san- 
dankwa) 18 5 — -- Malaysia 
16 6 — —- _ 
" 16 9 289E UC 
16 9 389K NY a 
: 16 9 2 KB uf 
Sect. II. Lantana Spach. 
V. bitchinense Makino 16 6 -- w. Japan 
- 18 9 4k PI-82381 e 
. 18 9 7OE sd. PI-82381 i 
sf 18 9 307E KR,s “ 
18 9 522E VZ, sd. - 
18 9 788E AA (Wilson) _ 
18 9 980E MR3 » 
7 18 9 998E < = 
. 18 9 1078E RH ef 
18 9 1143E CU ia 
18 9* — 1354E . . 
18 g* 1355E E430-29 a 
V. buddleifolium Wright 18 2 = c. China 
“ 20 6 a “ 
aa 18 9 7E PI-111380 - 
18 9 8E CU 
18 9 422E BB 
18 9 423E KN e 
. 18 9 710E MR “ 
a 18 9 808k AA7533 (Veitch) 7 
a 18 9 1049E K « 
. 18 g*  1328E jl 
“ 18 g* 1360E EX 
burejaeticum Rel. & Herd 18 g 278E GB, sd n. China 
18 9 428E O ae 
" 18 9 434K UW 160-46 . 
a 18 9 585E MT546-32(AA,sd) “ 
7 18 9 586K MT475-40(ST, sd.) : 
s 18 9 628E MT1144-40 
ns 18 9 675E KH 7 
18 9 772E AA4942(RE) 
V. & burkwoodii ony & Skip. 
(Vv. cuntese S< utile) 18 8 AA815-41-B AA815-41-B cult 
18 9 1E te ey 
18 9 518E sd. “ 
* 18 9 587E Me 295— 36(SN) 
- 18 9 1006K K1923HK 
18 9* — 1363E i) - 
‘Park Farm Hybric 18 9 923K HL cull 
Vv. &X ee Han Burk. ex. 
Pike (V. carlesti >< macro- 
ce phalunr) 8 AA618-53-A  AA-618-53-A cult 


1962 | EGOLF, CYTOLOGY OF VIBURNUM 143 


TABLE I (Continued) 


SOURCE GENERAL 
AvuTHOR- DocUMEN- OF DISTRIBU- 
SPECIES ITY TATIO} MATERIAL TION 
a 18 9 77E ONE, s 
18 9 433E UW st 
+ 18 9 689E WG es 
i‘ 18 9 1023E K 
V. carlesii Hemsl. 18 D5 ae — Korea 
- 20 6 = a e 
18 8 AA17981-A AA17981-A e 
18 9 112E HP2193 es 
iv 18 9 421E BB : 
e 18 9 521E WZ, sd. ss 
“ 18 9 536E JP, sd. i. 
i 18 9 676E KH e 
a 18 9 785E AA17981 (Gibbs) s 
om 18 9 1009E K972-34(Trickett)  “ 
os 18 9 1144E CU " 
2 20, 22 9 1145E HN, sd.( Korea) n 
u 20, 22 9 1146E TA, sd. : 
if 20, 22 9 1147E HM, sd. (s.China) “ 
18 g* 1369E DU s 
ss 18 gx 1596E E529-38 " 
V. < carlotta Hort. (V. x burk- 
woodii * carlesii) 18 9 78E CL cult. 
V.& chenaultii Chenault 18 9 406E RU cult 
oS 18 9 409E MT741-50 
s 18 9 519E WZ, sd. 2 
“ 18 9 592E MT218-51(SN) ef 
oh 18 9 1057E K227-48( AB) - 
V. cotinifolium D. Don 18 5 — -- Himal. 
: 18 9 819E AA1236—-52(K) ef 
ae 18 9 871E UW17493 L&S 
(Bhutan, sd.) 4 
S 18 g 1037E K56-81 Himal. 
x 18 9 1040E K1067-83 es 
& 18 9 1067E DU ee 
a 18 9 1086E L ‘ 
V. glomeratum Maxim. 18 9 817E AA, W180-5(K, sd.) China 
a 18 9 992E MT406-44 i 
te 18 9 1052E K sf 
V. < juddii Rehd. (V. carlesi 
<< bitchiuense) 18 — — cult. 
¢ 18 9 81E CL * 
oe 18 o* 9O7E MR “ 
7 18 9 606E MT451-48(AA) % 
be 18 9 756E AA1107-27 (Gibbs) ue 
18 g* 823E AA813-49 es 
18 g* 824E AA284-44 . 
a 18 9 Q73E P : 
“e 18 g* 1312E RH 
V.lantana L. 18 1 — AA Eur., w.As. 


144 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 
TABLE I (Continued) 
SOURCE GENERAL 
AuUTHOR- DoCUMEN- OF DISTRIBU- 
SPECIES I TAT MaArtERIAL TION 
7 18 5*,6 — — ae 
. 18 9 31E U ‘ 
- 18 9 239E MT1010-39 6 
- 18 9 301E KR, sd. ff | 
. 18 se) 385E Y as 
- 18 9 607E MT437-37 
(PI-107644) a 
” 18 9 608E MT640-34 nt 
" 18 9 722E BT78-38 - 
' 18 9 996E K 40-33 = 
18 9 1041E K ae 
. 18 9 1142E P Ki 
: 18 g*  1325E DO « 
a 18 9* 1456E R es 
. 18 g* 1467E E(B193-36) s 
‘Aurea Marginata’ 18 9 609E MT838-37(B) cult. 
‘Floribundum’ 18 9 1138E Pp cult. 
‘Lanceolatum’ 18 9 612E MT616-39 cult 
7 18 9 940E P . 
‘Lees’ 18 9 825E AA(K) cult. 
‘Macrophyllum’ 18 9 72E MT cult 
var. rugosum Lange 18 8 AA907-27-A_  AA907-27-A cult. 
> 18 9 9E HP es 
is 18 9 456E MT352-46 a 
“ 18 9 615E MT213-41(KH) i 
7 27 9 679E KH a 
° 18 9* 775E AA907-27(DA, sd.) “ 
f. variegatum (West.) Rehd. 18 9 618E MT1053-41 cult. 
= 18 g* 1470E E " 
V. macrocephalum Fort. f. sterile 
Dipp. 18 9 82E CL cult. 
“t 18 9 424E KN 
ut 18 9 713E MR47-121 et 
ms 18 9 975E P . 
at 18 9 1015E Kk ut 
sf 18 g* 1597E E tf 
7 micro phyllum (Oerst. ) Hemsl. 18 9 624E MT642-36 Mexico 
.m ongolcun: (Pall.) Rehd. 16 6 — ~- e. As. 
18 9 277E GB, sd. - 
ee 18 9 673E MTSOI1-S3 ° 
V. & rhytidocarpum Lemn. (V. 
buddleifolium  rhytidophyl- 
lum) 18 9 418E R cult 
18 9 643E 03-53 “ 
« 18 9 806E AA412-36(LE) ee 
V. rhytidophylloides Suring. (V. 
rhytidophyllum *K lantana) 18 8 AA711-36-A  AA711-36-A cult. 
af 18 9 102E HP “ 
af 18 9 426E KN(WW) “ 


1962] EGOLF, CYTOLOGY OF VIBURNUM 145 
TABLE I (Continued) 
SOURCE GENERAL 
AutTHOR- DocuUMEN- OF DISTRIBU- 
SPECIES TATION MATERIAL TION 
oe 18 9 449E Y e 
3 18 9 671E MT265-37 s 
- 18 9 754E AA711-36(LE) - 
a 18 9  826E AA1481-52(WW) te 
; 18 9 1038E K387-29 Bs 
V. rhytidophyllum Hemsl 18 2 —_ — c. & w. China 
oe 18 6 2451K RH - 
S 18 9 51E CU oT 
a 18 9 52E PI-58813 oy 
ee 18 9 91E MR * 
“ 18 9 304E KR, sd. : 
= 18 9 1048E K 201-07 (Veitch) se 
Hi 18 9* 1203E E401-37 ie 
e 18 g* 1205E E493-36 i 
eS 18 g* 1259E RH “ 
e 18 9* 1278E HI( Wilson) SS 
s 18 9*  1505E E493-36(CR) es 
f. roseum (Gard. Chron.) 
Rehd. 18 9 295E UW(CL) cult 
te 18 9  800E AA310-41(HL) ‘ 
f. Fe doen Boom 18 9 822E AA2 cult. 
V. EAA EED Maxi 18 9 128E AA15570 nw. China 
18 9 3907E NY(HP) 
We 18 9 465E MT1051-401 “ 
ss 18 9 652E coca 35(AA) iy 
H 18 9 726E 
ts 18 9 1014E KUAASG? 30) “ 
wu 18 9 1090E o 
V. stellulatum Hemsl. 18 9 506E a Himal 
Zs 18 9 1017E K131-38(DU) - 
V. utile Hemsl. 18 6 — — c. China 
s 18 9 214E DA, sd. s 
sf 18 9 425E KN : 
ss 18 9 507E DU oe 
i 18 9 694E PI-111380 i 
a 18 9 1008E K Wy 
18 Q* 1528E BH oS 
i 18 Q* 1529E WA ef 
18 g* 1530E E a 
V. veitchii Wright 18 8 AA7198 AA7198 c. China 
: 18 9 113E HP2177d sf 
a 18 9 261E MT1101-36 sf 
18 9 685E KH - 
i 18 9 727E NY67480 “ 
s 18 9 753E AA7198( Veitch) " 
18 9 1005E K101-13 (Veitch) Me 


18 9* 1560E DU1288W 


146 JOURNAL OF THE ARNOLD ARBORETUM 


TaBLe I (Continued) 


[ VOL. XLII 


SOURCE 


GENERAL 
AvutTHOoR- DocUMEN- OF DISTRIBU- 
SPECIES 2n TATION MATERIAL 
Sect. IIT. Pseudotinus Clarke 
V. furcatum Bl. ex Maxim. 18 6 — Japan 
“a 18 9 740E AA17988 (Wilson) ‘ 
7 8 9 1099E HL a 
lantanoides Michx. (as V. 
alnifolium) 18 1 —— AA e.N. Am. 
18 6 — ‘ 
. 18 9 497E RF os 
™ 18 g* 1876E GA ae 
V. svi podiale Graebn. 18 9 451E NY(LS) China 
V. urceolatum S. & Z. 18 8 \A876-51 A A876-51 Japan 
i 18 9*  1645E PI-227284 « 
Sect. IV. Pseudopulus (Dipp.) Rehd. 
V. plicatum Thunb 16 8 AAI8016-1 AA18016-1 e. AS. 
18 ) 92E MR681 m 
: 18 9 103E HP 2165 
= 18 9 157E AA18016-1 
‘ 16 ) S577E PN 
16 9 S78E PN 
‘ 16 9 779E AA933- aa _ 
16 9 10021 K629 
f. eabren (Nakai) Rehd. 16 9 532E TI, a shad Japan 
ea, urt 16 9 436E RH cult, 
16 9 843E AA134-53 a 
f. lanceolatum Rehd. 16 8 AA6122-1 AA6122-1 cult. 
= 16 9 658E MT1204-41 ~ 
16 763 FE AA6122-1(Sargent) 
{. — (Veitch) Ret a is 
ymentosum var. 
oe 18 6 6K RH cult. 
7 18 8 AA870-51-A AA870-51—-A ‘s 
16 9 SOOKE RS 
16 9 762E AA19355(K) a 
. 16 9 1010E K : 
16 9 1079E RH - 
‘Roseum’ 16 9 580E PN cult 
f. roseum (Doney) Rehd. 16 8 AA856-34 AA856-34 cult 
: 16 9 737E rf 
ee 16 9 742E AA856—34( BB) 
‘Rowallane’ 16 9 437E RH cult. 
‘St. Keverne’ 9 S505E RU cult 
f, tomentosum Thunb.) 
Rehd. (as V. tomentosum) 18 1 = NA e. As. 
- 18 6 2452K RH eS 
ed 18 9 43K CU ne 
a 16 9 95E MR1218 . 
“ 16 9 110E HP2195 . 


1962 | EGOLF, CYTOLOGY OF VIBURNUM 147 
TABLE I (Continued) 
SOURCE GENERAL 
AuTHOR- DocuMEN- OF DISTRIBU- 
SPECIES TATION MATERIAL TION 
i 18 9 484E MT176-37, 1204-41 “ 
i. 16 9 688E KH 
: 16 9 846E CU ‘. 
i 16 9 1001E K257-33(NN) : 
ss 18 9 1084E RH . 
Sect. V. Lentago DC. 

V. cassinoides L. 18 8 AA17997 AA17997 e. N. Am. 
ie 18 9 12E C ay 
a 18 9 106E HP2790 mf 
a 18 9 S75E ML, sd. 8 
18 9 590E MT, 570-43 a 
i 18 9 677E KH ee 
- 18 9 1000E K "i 
var. nanum Kriiss. 18 9 1088E HL cult 

; jackii Rehd. (V. len- 

tago xX prunifolium) 18 9 257E MT1037-40(AA) cult. 

‘ 18 9 965E AA17992-1-B - 
V.lentago L. 18 1 oe e.N. Am 
(v3 18 6 __ ae “e 
a 18 9 166E AA18021-A ey 
= 18 9 535E CU . 
sf 18 9 574E 50 - 

f. sphaerocarpum  (Fern.) 
Rehd. 18 9 619E MT892-35 e.N. Am, 
is 18 9 068E AA11316 nt 
V. nudum L. 18 6 — — e.N. Am. 
ye: 18 9 570E MC ies 
= 18 9 681E KH “ 
ms 18 9 988E MT as 
ie 18 9 1059E K961-31(Gibbs) 
ss 18 9 1151E G 
V. obovatum Walt. 18 9* 982E C se. U.S 
es 18 9*  1802E HF(Fla.) fs 
V. prunifolium L. 18 1 — AA e.N. Am. 
ss 18 6 — — a 
a 18 9 46E CU re 
18 9 47E E ss 
if 18 9 156E AA1805-2 of 
«“ 18 9 223E DA, s “ 
A 18 9 960E CU - 
wf 18 9 1020E K . 
V. rufidulum Raf, 18 9 517E WZ se.U.S 
ss 18 9 569E C af 
sf 18 9 645E MT1012-39 se 
a 18 9 1150E E < 


148 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 
TaBLe I (Continued) 
SOURCE GENERAL 
AuTHOR- DOCUMEN- OF DISTRIBU- 
SPECIES 2n TATION MATERIAL TION 
Sect. VI. Tinus Maxim. 
V. atrocyaneum Clarke 18 9 918E HL Himal. 
Re 18 9 1043E K566—-48(HL) of 
" 18 g* 1173E nt 
wt 18 Q* 1267E DU9198K.W. ms 
V. calvum Rehd. 18 9 922E HL w. China 
. 18 9 1054E K(E23-48) s 
a 18 g*  1285E E16-52(HL) . 
V. cinnamomifolium Rehd. 18 6 — — w. China 
a 18 9 85E CL af 
18 9 884E UW163-41(AR) as 
es 18 ce) 895E RH . 
“ 18 gx 1270E DU963W “ 
“ 18 Q* 1276E HC _ 
" 18 Q* 1372E BH 
a 18 Q* 1374E WA i 
V. davidii Franch. 18 6 — _ w. China 
" 18 9 86E ” 
. 18 9 181E E2485, sd es 
“ 18 9 415E RU a 
18 9 877E vo 
a 18 9 885E UW . 
ss 18 9 900E RH 
18 9 1029E © 
18 9*  1390E Ec63W a 
Af 18 9* 1391E A te 
‘Femina’ 18 9 503E cult. 
te 18 9*  1315E DU(MA) « 
V. harryanum Rehd. 18 5 —_ — w. China 
me 18 9 Q17E HL v 
“ 18 g* 1165E EX fe 
18 g* 1194E E56—42 i 
i 18 g* 1275E BC x 
ss 18 g* 1314E DU es 
V. propinquum Hemsl. 18 5 — = w. China 
Zs 18 9 559E K 
“ 18 9 875E VO " 
- 18 9 886E UW79-50 a 
‘Lanceolatum’ 18 9 924E HL cult. 
V. rigidum Vent. 18 9 556E K Canary Is. 
“ 18 9 858E c « 
“ 18 o*  1174E HI & 
V. tinus L. 36 2; 3; 
5,6 — —_— Eur 
_ 36 9 118E FN 
“ 36 9 267E AN, sd 
. 36 9 291E UW 7 
i 36 9 314E C ne 
36 9 1035E K ot 


1962 | EGOLF, CYTOLOGY OF VIBURNUM 149 
TABLE I (Continued) 
SOURCE GENERAL 
AuTHOR- DocuMEN- OF DIstTRIBU- 
SPECIES n ITY TATION MATERIAL TION 

ie 36 9 1064E K a 

a 36 9* 1180E E - 

i 36 gx 1260E RH sf 
‘French White’ 36 9 915E L cult. 
var. hirtulum Ait. 36 9 1028E K22-11-47 Eur 

ce 36 9 1100E HL n 
var. lucidum Ait. 72 9 288E UC Eur 

2 72 9 292E UW ce 

i 72 9 343E DU, sd i 

ct ees 9 997E K pt 

e 72 Q* 1198E E 

‘Variegatum’ 72 9 999E K cult 
‘Purpureum’ 36 9 1046E K1936 cult. 
Var. variegatum 36 9 1096E HL cult 

36 9 1181E E s 

Sect. VII. Megalotinus Maxim. 
V. coriaceum Bl. 18 9 879E vo As. 

3 18 9 1026E KS0-51(E) tf 

i 18 9 1101E HL 6 

ok 18 Q* 1167E EX te 

" 18 9* 1274E C i 

V. sempervirens Koch. 18 9 276E HN, sd. Malaysia 
Sect. VIII. Odontotinus Rehd. 
V. acerifolium L. 18 1 — AA e.N. Am. 
ss 18 6 — = % 

rf 18 9 1E CU s 

a 18 9 2E SO s 

as 18 9 357E DU 5 

be 18 9 581E MT202-38(WM) ss 

Hy 18 9 799E AA19181 “ 

y 18 9 1066E K258-53 (HP) fe 

cs 18 9 1144E LA oe 

V. betulifolium Batal. 18 6 — — c..& w. China 
is 18 9 297E UW(CL) eS 

7 18 9 447E NY xt 

tt 18 9 674E KH sé 

* 18 9 757E AA550-26 

(Rock 13476) ot 

nS 18 9 1003 E K61-08 at 

i 18 Q* 1187E E346F a 

c 18 g* 1247E RH uf 

rs 18 g* 1338E NS ms 

18 9* 1340E WA 7 

oe 18 Q* 1342E HL “ 

oH 18 9* 1347E DU238A-W fs 
‘Aurantiacum’ 18 9* 283E cult. 


150 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


TABLE I (Continued) 


SOURCE GENERAL 
AuTHOR- DOCUMEN- OF DISTRIBU- 
SPECIES 2n ITY TATION MATERIAL TION 
V. bracteatum Rehd. 72 9 455E MTS60-37 se. U.S. 
72 9 552E K is 
V.dasyanthum Rehd. 18 9 294E UW c. China 
ms 18 9 736E s 
os 18 9 1055E K910-39( LE) o 
7 18 9 1386E E£910-39( LE) " 
V. dentatum L. 54 6 — _ e.N. Am. 
a 36 8 AA17985 AAI7985 My 
: 36 9 16E CU e 
af 36 9 107E HP2212 = 
a 36 9 142E AA17985-B a 
ei 36 9 391E NY (HP) “ 
7 36 9 572E SO tf 
- 36 9 593K MT63S-34(0ON) “ 
¢ 36 9 890E kH ue 
a 36 9 1036E K(JG) ae 
7 36 9 1148E RE of 
var. deamit (Rehd.) Fern. 72 9 144E AA100-38-A C. 
72 9 636E MT1006-39 (Deam) ‘ 
var. pubescens Ait. 36 8 AA18008 AA18008 e.N. Am. 
et 36 8g AA18009 AA18009 ” 
- 72 9 49 CU a 
ia 36 9 179E AA18009 ee 
" 36 9 639E MT355-46 
(AA2106-4A) ‘“ 
“ 36 9 743E AA18008( DS) ef 
- 72 9 7T77E AAS5O70-1 7 
a 72 9 974E HP a 
: 72 9 1019E K1938HK . 
V. dilatatum Thunb. 18 8 AA229-46-B AA229-46-B ce. As. 
18 9 19E CU oe 
a 18 9 23E PI-76383 a 
“ 18 2) 30 PI-C3R29 «“ 
~ 18 9 93E MR4 et 
a 18 9 525E TG, sd. (Japan) «“ 
. 18 9 526E TG, sd. (Japan) e 
" 18 9 S64E HB, sd. (Japan) ‘ 
/ 18 9 784E AA7665—-D(CH) 
“ 18 9 832E AA137-52 
(Japan, sd.) “ 
18 9 835E AA138—52 
i 18 9 888E UW248-49(E, sd.) 
" 18 9 1021E K i 
f{. hispidum Nakai 18 8 AA17486-1-A AA17486-1-A e. Asia 
’ 18 9 146E AA17986-1-A te 
7 18 9 386E NY(TG) a 
ie 18 9 731E N¥2311-36(TG, sd.) “ 
18 9 S18E AA647-53 


(Japan, sd.) 


1962 | EGOLF, CYTOLOGY OF VIBURNUM Lod 
TABLE I (Continued) 
SOURCE SNER/ 
AvuTHOR- DOCUMEN- OF DISTRIBU- 
SPECIES 2n ITY TATION MATERIAL TION 
sf 18 9 828E AAS567-53 
(Japan, sd.) ef 
: 18 9 831E AAS569-5 
(Japan, sd.) s 
se 18 9 833E AA139-52 
(Japan, sd.) 
18 9 870E UW91-46(NY,sd.) “ 
f. pilosum (Thunb.) Nakai 18 9 446E NY(TG) e. As. 
s 18 9* 534E TG, sd.( Japan) 
si 18 9* 562E HB, sd.( Japan) se 
“ 18 9 563E es te 
f. xanthocarpum Rehd. 18 8 A A10140 AA10140 cult. 
a 18 9 40E A10140 a 
: 18 9 373E y . 
18 9 S94E MT457-45(KH) = 
os 18 9* 1397E E(IC) _ 
V. ellipticum Hook. 18 9 430E LO w. USS. 
s 18 9 1060E K316-32(EN) "s 
V.erosum Thunb. 18 9 24E PI-4276P e. As. 
< 18 9 89E MR2015 ef 
" 18 9 515E HB, sd. 
s 18 9 595E se SS(AAI1S06 
18 9 734E NY179 
oe 18 9 810E ee aries g 
18 9 881E UW168-S50(NY, sd.) “ 
ot 18 9 1081E 
s 18 g* 1398E E15163 Yii ee 
var. punctatum Franch. 18 9 511E TC(Japan, sd.) e. As. 
as 18 9 512E = 
18 9 527E a sd.(Mt. Tawa) 
var. taquetii Rehd. 18 9* 1401E ~orea 
V. flavescens W.W.Sm 18 9* 202E eo 86, sd. China 
* 18 9* 882E UW274-49(E, sd.) 
18 9* 1407E E w 
V. foetidum Wall. 16 6 — — Himal. 
x 18 9 440E RH e 
2 18 9 1087E HL M 
var. rectangulatunt 
(Graebn.) Rehd. 16 6 — — w. China 
yi 18 Q* 1158E HA _ 
e 18 9* 1163E EX = 
V. hanceanum Maxim. 72 9 477E MT564-39 China 
os 72 9 821E AA1507-51(HL) ae 
a 72 9 1033E K124-26(AA) “ 
i 72 9 1132E P = 
V. hirtulum Rehd. 18 9 883E UW264-49(LS, sd.) China 
cs 18 g* 1784E wy i 
V. hupehense Rehd. 18 1 — AA c. China 
18 2 — — “ 
(as V. hirtulum) 18 8 AA708-37-B AA708-37-B _ 


152 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 
TaBLe I (Continued) 
SOURCE GENERAL 
AUTHOR- DOCUMEN- OF DISTRIBU- 
SPECIES 2n TATION MATERIAL TION 
. 18 9  302E KR, sd. , 
18 9 355E DU, sd. " 
: 18 9 463E MT, 390-39 
e 18 9 678E . 
18 9 T11E MR601A( Wilson) a" 
- 18 9 738E BB " 
us 18 9 765E AA18020 
(Wilson 601) 
“ 18 9 769E AA708-37-B 
(LS, sd.) ‘ 
. 18 9 978E MR601A-5 
‘ 18 9 995E 
(Wilson 601) . 
oe 9 1080E RH be 
V. ichangense (Hemsl.) Rehd. 18 9 OE CL c. & w. China 
‘ 18 9 695E PI-114819 ee 
4 18 9 838E AA(KN) «“ 
ss 18 9 1072E RH et 
a 18 9 1085E HL i 
= 18 Q* 1162E EX 7 
ot 18 g* 1319E BC 
V. japonicum (Thunb.) Spreng. 18 9 560E K apan 
oe 18 9 566E HB, sd. (Japan) - 
ae 18 9 876E VO re 
“ 18 9 887E UW631-50(HL) “ 
ns 18 9 1047E K167-37(DA) “ 
. 18 gx 1159E ld 
V. lobophyllum Graebn. 18 1 —_ AA c. & w. China 
T9 20 6 ae en (<9 
. 22 7 EX ~ 
M 18 9 108E HP 2152 at 
re 18 9 131E AA19494-C a 
= 18 9 351E Du, sd. 
= 27 9 620E MT1303-35 
(NB, sd.) “ 
- 18 9 730E NY67491(LS) a 
ee 18 9 CO7E —! ne 
18 g* 1004E K88-08 (AA238w) se 
7 18 9 1128E P i 
7 18 9* 1472E ts 
V. molle Michx. 36 9 391E NY (HP) c.US 
ee 36 9 625E MT1044—40 * 
ee 36 9 803E A A18294-A 
(Palmer) i 
" 36 9 1016E 104-35(DU) 
f. leiophyllum Rehd. 18 8 AA4643-1-A  AA4643-1-A c.US 
" 36 9 804E AA4643 (Bush) _ 
- 36 9 985K MT 1097-36 - 
V. ovatifolium Rehd. 18 1 -—- AA w. China 
18 8 AA20078A AA200784 as 


1962 | EGOLF, CYTOLOGY OF VIBURNUM 153 
TABLE I (Continued) 
SOURCE GENERAL 
AUTHOR- DOCUMEN- OF DISTRIBU- 
SPECIES 2n TATION MATERIAL TION 
hy 18 9 290E UW (MT, sd.) i 
_ 18 9 764E AA20078 
(Wilson 590) . 
sf 18 9 1025E K1008-34(NN) 
t 18 9 1076E H ty 
V. parvifolium Hayata 18 9 889K RH Formosa 
V. phlebotrichum S. & Z. 36 9 531E TG, sd. (Japan) an 
is 36 9 723E BT37-39 be 
‘“ 36 9 869E UW38-—49(HL) ef 
or 36 9 1095E HL 
V. rafinesquianum 
Roem & Schult. 36 9 365E K e.N. Am 
SS 36 9 573E N10) : 
‘6 36 9 76E LA t 
var. affine (Schneid.) House 20 8 AA4622-2-B AA4622-2-B e.N. Am. 
ie 36 9 71E : 
36 9 768E AA17972 (Bush) tt 
“ 36 fe) 10S51E 78-36 oy 
V. recognitum Fern. 36 9 707E MR54-94 e.N. Am 
V. scabrellum Chapm. 72 9 C6E MR se. U.S 
oy 72 9 396E NY (BT) cs 
oe 72 9 567E MC us 
os 72 9 579E PN(Va.) n 
uf 72 9 714E MR2200 
“ 72 9 745E AA11549-B 
(Harbison) i 
V. setigerum Hance 
(as V. theiferum Rehd.) 18 2 — c. & w. China 
2 36 8 AA20189 AA20189 2 
i 36 9 55E CU s 
ss 18,36 9 57E PI-104128 « 
36 9 88E MR11( Wilson) is 
v 36 9 654E en an es R) s 
ey 36 9 655E MT52 a 
36 9 709E MR218A (Wilson e 
- 36 9 744E AA20 
aes 236) cs 
{. aurantiacum Rehd 36 8 AA812-32 AA812-32 c. & w. China 
a 36 9 SSE PI-023027 ae 
ie 36 9 60E PI-TN-R8 ef 
a 36 9 94E MR12 eee 236)---% 
fe 36 9 815E AA19 
ana 236) sf 
V. wilsonit Rhed. 18 9 1053E K262-23(HL) w. China 
be 18 9 1068E DU a 
a 18 g* 1562E E262-33(HL) s 
V. wrighti Miq. 16 6 2450K RH Japan 
~ 18 9 4E CL = 
18 9 116E HP2202 +f 
i 18 9 286E Cc 


154 JOURNAL OF THE ARNOLD ARBORETUM [| VOL. XLUI 
TABLE I (Continued) 
SOURCE GENERAL 
AuTHOR- DOCUMEN- OF DISTRIBU- 
SPECIES 2n TATION MATERIAL 
. 18 9 S16E HB, sd.(Japan) a 
ne 18 9 667K MT165-37 e 
“ 18 9 686K ‘ se 
" 18 781E AA18015 (Sargent) 
at 18 9 912 RH “ 
var. eglandulosum  (Miq.) 
Nakai 9 565E HB, sd.( Japan) Japan 
var. hessei (Koehne) Rehd. 18 9 402K NY Japan 
18 9 1031E K - 
Sect. IX. Opulus DC. 
V. edule (Michx.) Raf. 18 9 873E UW 1053-50 
(White R.) n.N. Am. 
- 18 g* 1098E HL _ 
V. kansuensis Batal. 18 9 1083E RH w. China 
~ 18 9 1094E HL “ 
nt 18 g* 1453E GP 13248 L&S : 
V. opulus L. 18 1 = AA Eur., n. Afr. 
“ 18 —— — “ 
(as V. sargentii) 20 6 2459K RH 
s 18 9 39E “U 
18 9 101E K2491, sd. is 
“ 18 0) 324K PO 7 
e 18 9 344E DU _ 
= 18 9 375E NY ae 
- 18 9 400E NY (Austria) ag 
18 9 459E MT1189-35 zs 
7 18 9 S45E < 
is 18 9 672E MT49-52(AA) « 
“ 18 9 687K KH 
a 18 9 741E AA * 
18 9 786E AA20736(E, sd.) a 
" 18 9 1013E AA562-30 = 
ag 18 9 1030E K53-46(RH) es 
ff 18 9 1032E K94-29(G) he 
7 18 9 1063F K106-63 . 
‘Aureum’ 18 9 629E MT1046-40(SP) cult 
“ 18 9 842K AA997-52 
™ 18 9 892E RH en 
. 18 9 1012E K “i 
‘Compactum’ 18 9 417E RU cult 
“ 18 9*  1485E JG a 
var, nmanum (David) Zabel. 18 9 524E CU cult 
" 18 9 630E MT118-53( HL) 3 
ae 18 9 981E C ns 
‘Notcutt’ 18 9 670K MT211-51 cult 
- 18 9 747E AA814-38( NN) ° 


1962 | EGOLF, CYTOLOGY OF VIBURNUM 155 
TABLE I (Continued) 
SOURCE GENERAL 
AuTHOoR- DOCUMEN- OF DISTRIBU- 
SPECIES 2n ITY TATION MatTERIAL TION 
f. roseum (L.) Heg. (as V. 
OC var. sterile) 18 6 2463K RH cult. 
18 ) 41E CU ve 
e 18 9 246E MT156-38 - 
Z 18 9 632E MT1098-36( AA) A 
oo 18 9 790E AA26-47-A Z 
i 18 9 1107E a 
i 18 9 1118E P “ 
f, variegatum (West) Zabel 18 9 pee DA cult 
ee 9 33E T528-51(DA) ce 
{. xanthocarpum (Endl.) Rehd. 18 8 AAn258 28-A AA1298-28-A cult. 
iz9 18 9 O4E P2 ae 
oe 18 9 oe U * 
fe 18 9 634E MT1099-36(HP) 
. 18 9) 791E AA1298-28-A ms 
V. orientale Pall. 18 9 175E AA677-33 w. As 
Me 18 g 296K UW(MT, sd.) “ 
# 18 9 837E AA934—52 _ 
V.sargentii Koehne 18 1 — A! ne. As 
st 18 9 74E PI-81798 if 
o 18 9 300E KR mn 
es 18 9 337E PO a 
i 18 9 404E RU - 
. 18 9 794E AA18012-B(Korea) “ 
“ 18 9 827E AA571— 
ee sd.) i 
i 18 9 829E AA 
saan, sd.) = 
- 18 ) 830E AA 
saan, sd.) ee 
18 9 834E AA 
es sd.) a 
* 18 9 COlE RH ss 
f. calvescens (Rehd.) Rehd. 18 8 AA467-26 AA467-26 ne. As. 
i 18 Q* 4C4E AA467-26 es 
: 18 9 797E ee sd.) 
st 18 9 798E 
eae 1QAGEHe at 
a 18 9 1042E K234-38 Marsh o 
f. flavum Rehd 18 9 117E HP2159 cult. 
18 g* 130E AA21419 se 
‘ 18 9 647E MT2195-22(HP) es 
in 18 9 795E AA11037(HP) . 
‘Puberulum’ 18 9 649E MT975-38(GB) cult. 
V. trilobum Marsh 18 1 — AA n.N. Am 
18 6 — — 
4: 18 9 327E PO a 
is 18 9 659E MT1228-38 af 
oe 18 9 816E AA15673 i 
i: 18 9 1058E K205-48(KR) oe 


156 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


TaBLeE I (Continued) 


SOURCE GENERAL 
AuTHOR- DoCUMEN- OF DISTRIBU- 
SPECIES 2n ITY TATION MATERIAL TION 
‘Andrews’ 18 9 248E MT1055-—40 cult. 
18 9 793E AA292-—42(LI) es 
af 18 9* 1663E WLA66-104 ee 
‘Compactum’ 18 8 AA871-51 AA871-51 cult. 
- 18 9 664E MT216-53(KH) _ 
" 18 9 684E KH ne 
‘Hans’ 18 9 661E MTS801-40 cult. 
“ 18 9 789E AA293-42(LI) it 
‘Wentworth’ 18 9 663E MT1057-40 cult. 
a 18 9 783E AA294-42(LI) « 
Unidentified species 18 9 693E PI-22978 
uf 18 9 700E PI-111382 


CONTROVERSIAL CHROMOSOME COUNTS 


Poucques (27, 28) reported 2” = 18 in Viburnum sandankwa, a syno- 
nym of V. suspensum, while a somatic chromosome number of 16 is re- 
ported here. However, since several different evergreen forms have been 
referred to V. sandankwa, it is possible that Poucques’ plant was not the 
same as V. suspensum. Simonet and Miedzyrzecki (35) reported a chromo- 
some number 2” = 18 for V. setigerum (V. theiferum). In the present 
study a single plant of this species was found with a chromosome number 
2n = 18; whereas other determinations, including that of Thomas (40), 
revealed a chromosome number of 2” = 36. 

The somatic chromosome counts reported in this study for thirteen 
species and four varieties differ from those reported by Janaki Ammal 
(18). In order to check Janaki Ammal’s counts, and, if possible, to 
resolve the differences between our studies, an attempt was made to 
secure all the species grown at the Royal Horticultural Society’s Garden, 
Wisley; at the Royal Botanic Gardens, Kew; and in the Jardin des 
Plantes, Paris. It is assumed that plants from these sources are similar 
to the material studied by her, but there is no assurance that the same 
plants were sampled. Since she published no record of the particular 
plants involved, it is uncertain whether she studied the same species in 
all three gardens or in only one. In any case, it seems logical to assume 
that some of the plants included in this study were the same as some of 
those she studied. 

As with most cytological investigations, many of the studies must be 
made with cultivated plants that may be variants of the native species. 
The plant material used or the technique used could result in differences 
of chromosome counts. In a personal letter from Janaki Ammal it was 
stated that she had used the lacmoid leaf-bud technique (10), whereas 


1962] EGOLF, CYTOLOGY OF VIBURNUM Lo? 


in the present study root-tip smears were used. A critical comparison of 
her cytological study with the present one is impossible, for no slides, 
drawings, or photomicrographs and only a limited number of herbarium 
specimens were available. For clarity of discussion the differences between 
her and my counts are grouped into two classes: 1) species differing by 
a few chromosomes per complement and, 2) those differing in the number 
of sets of chromosomes in each complement. 

In the first category, Viburnum bitchiuense, V. foetidum var. foetidum, 
V. foetidum var. rectangulatum, V. mongolicum, and V. wrightii were 
determined to have 2” = 18 chromosomes, two more than reported by 
Janaki Ammal (18). Among the plants studied were V. bitchiuense, from 
Kew, and V. bitchiuense, V. foetidum vars. foetidum and rectangulatum, 
and V. wright from the R.H.S. Garden, Wisley. In the collection at 
Wisley is a plant identified as V. foetidum var. rectangulatum which is 
probably the plant studied by Janaki Ammal. Plant 1084E of this study, 
which was received from the R.H.S. Garden as V. foetidum var. rectangu- 
latum and which has horizontally spreading branches, oblong-ovate leaves, 
and an inflorescence with fertile flowers surrounded by sterile marginal 
flowers, is correctly identified as V. plicatum {. tomentosum. Upon visiting 
the R.H.S. Garden the author further verified the identification of this 
individual plant. 

The present study has revealed that Viburnum carlesi is composed of 
a complex 2” = 18, 20, and 22 chromosome forms. All the plants studied 
that were obtained from cultivation, including a plant from Kew, had 
2n = 18 chromosomes, which agrees with the reports by Poucques (28) 
and by Simonet and Miedzyrzecki (35). Janaki Ammal (18) reported 
this species have 2m = 20 chromosomes. 

The present study agrees with the previously reported counts of 2n 
= 18 in Viburnum buddletfolium (35) and in V. lobophyllum (32), but 
Janaki Ammal (18) reported 2” = 20 and 22 (the count of Enoch) for 
V. lobophyllum. Included in my study was a plant of V. lobophyllum 
from Kew. 

In this study both Viburnum plicatum {. plicatum and f{. tomcntosum 
were determined to have forms with 2n = 16 and 2m = 18. Janaki Ammal 
(18) reported both these and f. mariesii to have 2n = 18. The plants of 
f. mariesii and f. tomentosum from Kew and of f. mariesii from the R.H.S. 
Garden used in this study have chromosome complements of 2n = 16. 
The herbarium specimens deposited at Kew by Janaki Ammal clarify the 
discrepancies, and this documentation has been added to Tasre I. Speci- 
men 2456K of V. plicatum {. mariesii (V. tomentosum mariesu) has an 
annotation note “2m = 16?” which definitely indicates that her published 
count was questionable. The specimen of V. plicatum (V. tomentosum 
sterile), 2463K, collected at the R.H.S. Garden, is identified by me as 
V. opulus £. roseum. Likewise, specimen 2459K of V. sargentii, collected 
at the R.H.S. Garden, is V. opulus. The plants of V. odoratissimum from 
Kew were found to be of the 2m = 32 variation. Janaki Ammal reports 
2n = 40 for this species. This last number, however, was found elsewhere 


158 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


in the V. odoratissimum complex in this study and has likewise been 
reported by Sugiura (38). 

The number of genomes reported by Janaki Ammal (18) in the comple- 
ments of four species and one variety differ from those found in this 
study. Viburnum erubescens, V. henryi, and V. dentatum, including in 
the present study specimens of all three from Kew and of V. dentatum 
from R.H.S., were found to be tetraploid, whereas Janaki Ammal re- 
ported them all to be hexaploid. Janaki Ammal’s theory that the species 
evolved after spontaneous doubling of an unstable triploid obviously is 
not supported by this new evidence. Viburnum sieboldii, reported by 
Janaki Ammal (18) to be a diploid, was determined in this study to have 
both diploid and tetraploid forms, the tetraploid occurring much more 
frequently. The plant from the R.H.S. Garden studied was the tetraploid 
form. Janaki Ammal (18) reported V. fragrans ‘Album’ to be a tetraploid, 
but plants of this variety from R.H.S. Garden, and from other sources 
utilized in this study, were determined to be diploids. Thomas (40) 
also reports this variety to be diploid. 

Among the 536 counts listed by the author (14) were a number of 
plants procured from the Arnold Arboretum. Thirteen of the plants 
studied by Thomas (40) are identical with those from which the author 
acquired material, while another six plants have the same Arnold Arbo- 
retum accession number but are not necessarily the same individual plant. 
The author lists the counts of an additional sixty-six plants from the 
Arnold Arboretum. Two of Thomas’ counts, those for Viburneum rafi- 
nesquianum var. affine (2n = 20) and V. molle f. leiophyllum (2n = 18), 
differ significantly from the present work. Viburnum rafinesquianum var. 
affine was determined to be 2m = 36 for all plants studied. A plant of V. 
molle {, leiophyllum bearing the Arnold Arboretum accession number 
4643 was determined to be 2” = 36, while Thomas reports 2n = 20 for 
plant 4643—-1—A, the latter having been propagated vegetatively from one 
of the original lot. Quite possibly, either a mixed lot or a mistake in 
labeling may be involved. 


VARIATION IN CHROMOSOME NUMBERS 
WITHIN SPECIES AND VARIETIES 


Differences in chromosome number were found within six species and 
five varieties. These differences can be placed in two classes for dis- 
cussion: 1) those species that differ by a few chromosomes and, 2) those 
species that differ by a number of genomes. 

The first category includes Viburnum plicatum and V. carlesii. Morpho- 
logically indistinguishable plants of V. plicatum, with 2n = 16 and 2n — 
18, have been found. Comparison of the chromosome complements of 
these two forms reveals that the form with 2” = 18 has an extra pair of 
metacentric chromosomes. This species will be discussed later in more 
detail. All plants of V. carlesii from cultivation have 2” chromosome 
complements of 18. However, among plants produced from seeds col- 


1962 | EGOLF, CYTOLOGY OF VIBURNUM 159 


lected in Korea, the native habitat of the species, chromosome comple- 
ments of 2” = 18, 20, and 22 have been found. 

Six species that differed in number of genomes were studied. One plant 
of Viburnum lobophyllum (2n = 27) was determined to be a triploid, with 
three cytologically identical genomes. All other specimens of this species 
studied were diploid, with 21 = 18. Since the triploid plant appears iden- 
tical with the diploid, it does not seem likely that this is a hybrid between 
the diploid V. lobophyllum and one of the tetraploid species. It is possi- 
ble that this triploid could have resulted from the fertilization of an un- 
reduced functional gamete by a normal gamete. Likewise, it could have 
originated as a cross between diploid and tetraploid plants of V. lobophyl- 
lum, although this does not seem probable since no tetraploid V. lobophyl- 
lum is known. 

One species, Viburnum odoratissimum, is represented by tetraploid and 
pentaploid forms. These two forms, which have distinct vegetative dif- 
ferences, comprise a taxonomic complex that is given further consideration 
later. The pentaploid form, with 27 = 40, has five similar genomes, each 
of which is porphologically, identical with the genomes of the tetraploid. 
It is unknown whether the four pentaploids and seven tetraploids are 
representative of the variation that occurs in native populations. 

Individual diploid plants of Viburnum sieboldii and V. setigerum were 
discovered in species that otherwise are tetraploid. The diploid plant of V. 
setigerum was isolated from a group of plants grown from seed obtained 
from the U. S. Plant Introduction Garden, Glenn Dale, Maryland. This 
plant probably resulted from parthenogenesis wherein an unreduced gamete 
developed without syngamy. The diploid plant of V. steboldii was secured 
from a commercial nursery and it is not known whether the plant was 
propagated asexually or from seed. The diploid plants of these species are 
still immature, making it impossible to compare them critically with tetra- 
ploid plants. Although at this time they do not appear morphologically 
distinct from the tetraploid, they are obviously somewhat weaker, and 
growth has been slower. This may be partially or entirely due to environ- 
mental conditions, however. 

Only six of the sixty-one varieties observed in this study had chromosome 
complements with numbers different from the species (i.e., the typical 
varieties). Viburnum tinus var. lucidum, V. dentatum var. deamtt, and 
V. dentatum var. pubescens, all 2n = 72, had double the number of gen- 
omes of the typical variety of the species. One of the five collections of 
V. lantana var. rugosum was found to be a triploid, 2x = 27. Though there 
are three cytologically identical genomes in this particular plant, it does 
not appear to be morphologically distinct from the other collections. It is 
conceivable that this plant developed from a chance unreduced gamete that 
was fertilized by a normal gamete. 

In Viburnum plicatum {. tomentosum and V. plicatum f. mariesit, as 
well as in V. plicatum f. plicatum, occur both 2m = 16 and 18 chromo- 
some forms which cannot be distinguished by vegetative characteristics. 
All other taxa of this species which have been examined (V. plicatum f. 


160 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


glabrum, V. plicatum ‘Lanarth,’ V. plicatum f. lanceolatum, V. plicatum f. 
mariesit, V. plicatum ‘Roseum,’ V. plicatum ‘Rowallane,’ and V. plicatum 
‘St. Keverne’) have chromosome complements of 2” = 16. The forms 
with 2m = 18 differ from those with 2n = 16 by an additional pair 
of metacentric chromosomes. Since the 2m = 16 plants produced the most 
abundant fruit, it is to be expected that these forms should have a higher 
rate of survival. The evidence indicates that most of the minor variations 
in Viburnum that have been given varietal rank are the result of genic or 
intrachromosomal changes, rather than the result of changes in chromo- 
some numbers. 


CYTOTAXONOMIC COMPLEXES 


Differences in the chromosome numbers of different collections of V1- 
burnum odoratissimum, V. carlesti, and V. dentatum suggest that each of 
these is a species complex. 


VIBURNUM ODORATISSIMUM. Two distinct forms are evident in V. odora- 
tissimum, the plants of which were secured from nine different sources. The 
one has smooth-barked branches; thin, coriaceous, elliptic-ovate leaves; 
and indistinct axillary buds. The other has stout, lenticular branches; 
thick, coriaceous, elliptic-obovate leaves; and prominent axillary buds. The 
illustrations of V. odoratissimum by Dippel (13) and of V. awabuki by 
Nakai (23) portray two types which are respectively similar to the two 
forms observed in the present study. Nakai (23) recognized V. odoratis- 
simum, V. liukinense, V. awabuki, and V. awabuki var. serratum in this 
complex, but other taxonomists generally have accepted only one species, 
V. odoratissimum. Because all the plants used in this study have not yet 
produced flowers, it is impossible to make a positive identification of these 
variants. Chromosome counts of 2m = 32 (tetraploid) and 2” = 40 (pen- 
taploid) have been observed among both morphological forms of this 
species complex. Two of the three plants with 27 = 40 are from the same 
original source and are of the variant with coriaceous leaves and stout, 
lenticular branches. Sugiura’s (38) report of 27 = 40 in V. awabuki 
also indicates that his V. awabuki may be different from V. odoratissimum. 
Do these represent species or are they variants of one species? Although 
the cultivated material studied probably does not differ from representa- 
tives of the native populations of this species complex, additional material 
from known populations and further study will be necessary to determine 
the relationships within the species. 


VIBURNUM CARLESII. In this study all plants of V. carlesii received from 
cultivation have chromosome counts of 2m = 18. However, seeds sup- 
posedly collected from native populations in China and Korea produced an 
array of plants with 2m = 18, 20, or 22 chromosomes. It is difficult, if not 
impossible, to explain this variation if this is a true species. Of course, 
there is no assurance that this seed was from isolated plants and not from 


1962 | EGOLF, CYTOLOGY OF VIBURNUM 161 


plants growing near other species with which there could be cross-pollina- 
tion. 

As previously noted, Janaki Ammal (18) proposed that Viburnum car- 
lesii arose as a backcross between a chance triploid (2m = 24) and the 
normal diploid (2m = 16) of V. bitchiuense. In view of the present study, 
her explanation appears inadequate, for V. bitchiuense has 2n = 18 chro- 
mosomes, rather than the 16 chromosomes required by her proposal. If 
triploid plants exist in nature, this variation in the chromosome complement 
of seedlings might be the result of self-pollination or cross-pollination be- 
tween the triploid and diploid forms, which would produce progeny with 
additional chromosomes. It is doubtful, even if the triploid does exist, 
that fruit would normally result from self-pollination. 

Most taxonomists consider Viburnum bitchiuense and V. carlesii to be 
closely related. However, few have gone to the extreme of Nakai (24), 
who not only reduced V. bitchiuense to a variety of V. carlesii, but also put 
both of these in a new genus Solenolantana. It is doubtful if this complex 
requires the latter action. Pollinations by the author of V. bitchiuense (2n 
= 18) X V. carlesii (2n = 18) produced no seed, while the reciprocal 
cross between these species produced, from ninety-nine flowers pollinated, 
forty-three seed which have yielded forty-one plants. A meiotic chromo- 
some study of these plants should indicate more clearly the natural rela- 
tionships of these species. Until a sporocyte study is made of authentic 
materials collected from native populations, this remains an unsolved 
cytotaxonomic complex. 


VIBURNUM DENTATUM. Viburnum dentatum var. dentatum and V. denta- 
tum var. pubescens, represented by 2n = 36 and 2n = 72, form the third 
cytotaxonomic complex. According to Rehder (31), V. pubescens is a 
synonym of V. dentatum var. pubescens. In the V. dentatum—pubescens 
complex specific delimitation has been based almost entirely on the pres- 
ence and distribution of pubescence on petioles, leaf surfaces, and inflores- 
cence branches or combinations of these. Blake (4), who examined So- 
lander’s manuscript of Hortus Kewensis, an early treatment of this com- 
plex, and who also studied native material, recognized V. pubescens, V. 
pubescens var. canbyi, and V. pubescens var. longifolium. Rehder (29) de- 
scribed two new varieties from Indiana, V. pubescens var. deamti and V. 
pubescens var. indianense, which have only minor differences from each 
other. 

Svenson (39) commented on the variations of this group, which he 
separated on the basis of leaf shape and pubescence into Viburnum denta- 
tum, V. dentatum var. lucidum, V. dentatum var. pubescens and V. pubes- 
cens var. semitomentosum. He also concluded that V. pubescens var. 
deamii probably is a separable variation. 

Fernald (16) was in disagreement with Svenson’s reduction of the 
glabrous-twigged form of Viburnum dentatum to varietal rank as V. denta- 
tum var. lucidum, and elevated this glabrous variation to the rank of 
species with the specific name V. recognitum. Viburnum recognitum is dis- 


162 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


tinguished from V. dentatum by its glabrous branchlets and cyme, glabrous 
or glabrate foliage, and flowers produced ten days to three weeks earlier. 
Fernald further stated that V. dentatum and its variations and V. recogni- 
tum “are both hopelessly variable in leaf outline and toothing of leaves, 
each of them with blades varying from lance-ovate to ovate-oblong to or- 
bicular, with veins prominent beneath or obscure, with length from 2.5 to 
10 cm. and breadth from 2 to 8 cm.” Viburnum recognitum, V. recognitum 
var. alabamense, V. crenatum, V. dentatum, V. carolinianum, V. carolinta- 
num var. deamii, and V. pubescens are entities recognized by McAtee (21) 
in this species complex. 

The present study follows Rehder’s treatment which reduces this com- 
plex to Viburnum dentatum, V. dentatum var. pubescens, and V. dentatum 
var. deami. The material studied here was received under practically 
every one of the different names applied by past authors to these varia- 
tions. Many of these collections were from cultivated plants, since mate- 
rials from native populations were not available for all species or varieties. 
Therefore, it must be realized that the material studied is not necessarily 
an adequate sampling of variability of the native population, but it does 
give indications which may establish a basis for further research. The 
plants are being maintained for further study and identification. 

Two chromosomal forms, 27 = 36 and 2m = 72, were located among 
plants of Viburnum dentatum var. pubescens. All plants of V. dentatum 
var. deamti had 2n = 72. Are these plants of another variety or polyploid 
forms of the same species? Are these variations within a species the result 
of natural hybridization that has been followed by segregation and selec- 
tion? The variations between V. dentatum var. dentatum, var. pubescens, 
and var. deamii have been described taxonomically but are only of minor 
magnitude. Likewise, V. recognitum is only slightly different from V. 
dentatum, though it has been elevated to specific rank. Are these cases in 
which speciation is resulting from natural hybridization but in which di- 
vergence of types is not yet great enough for complete delimitation? Is the 
glabrous-branched V. recognitum a plant of a segregating hybrid population 
or does it represent another species? 

It has been suggested that Viburnum dentatum may have crossed with 
V. rafinesquianum, another related species, to produce by introgressive 
hybridization a different ecological population. Normally, however, these 
species are isolated by season of bloom, V. dentatum flowering approxi- 
mately ten days later than V. rafinesquianum, and by habitat, V. dentatum 
being located on moist soil and V. rafinesquianum on dry upland. It is 
possible that a few late flowers of V. rafinesquianum may be shedding viable 
pollen when the first V. dentatum flowers open, and the distance between 
plants would not prevent cross-pollination. The offspring produced by 
such crosses could, over a period of time, produce a population of a type 
differing from the original species of the locale. In the particular area of 
South Hill, near Ithaca, N.Y., where plants were used for controlled pol- 
linations in this study, there is no evidence of natural hybridization, but in 
similar situations elsewhere the densely pubescent V. rafinesquianum may 


1962 | EGOLF, CYTOLOGY OF VIBURNUM 163 


have crossed with the ay glabrous V. dentatum to produce a population 
with additional variation 

A further pees arises from the report by Hes Ammal (18) 
that Viburnum dentatum has a chromosome number of 2” = 54. In the 
present study, 2x = 36 has been counted in all cases. It is ‘gical to think 
that her plant was a hybrid, unless the count was incorrectly determined, 
since all of her work was done on cultivated plants growing in close proxim- 
ity and hence subject to crossing. Since it is the custom of many botanic 
gardens to raise plants from seed, not realizing the seed may be from 
cross-pollination and not true for the species type, it is possible that the 
plants studied by her may have originated in this way. 

One approach to the problem is experimental; that is, to reproduce a 
similar plant by controlled hybridization. All the possible pollination 
combinations between Viburnum dentatum var. dentatum, V. dentatum var. 
pubescens, and V. rafinesquianum were made in the present study. From 
386 flowers of V. dentatum pollinated by V. rafinesquianum were produced 
190 seeds from which 123 plants have been grown. From 130 flowers pol- 
linated in the cross V. dentatum var. dentatum * var. pubescens, 49 seed 
and 15 plants were obtained. The only other combination to yield seed 
was V. dentatum var. pubescens * V. rafinesquianum which, from 329 
pollinations, produced five seed that yielded two plants. The other com- 
binations failed to produce seed, but this cannot be attributed necessarily 
to sterility or incompatibility, for climatic conditions and technique may 
have been variable factors. That seeds and plants were procured from 
crosses between these species and varieties indicates a relationship within 
the complex. As these plants attain flowering size, a study of the meiotic 
chromosome configurations of the sporocytes should reveal the inter- 
relationships more definitely. It will be desirable to repeat those crosses 
that produced no seed and to attempt additional crosses with other related 
species from section ODONTOTINUS. 

Because polyploids are found in this complex, a study of the native 
populations will be required to resolve the problem. Such experimental 
studies should not be concentrated within a few isolated populations but 
should cover the distribution range of this species complex so that dif- 
ferences and relationships between populations, as well as within popula- 
tions, can be determined. With the union of the evidence from cytology, 
genetics, and taxonomy the intricate relationships of this complex should 
eventually be clarified further. 

The section ODONTOTINUS of Rehder’s classification includes other species 
that occur in the same geographical areas as members of the Viburnum 
dentatum—pubescens complex. Other taxa related to this complex and war- 
ranting study include V. molle, V. scabrellum, V. bracteatum, and V. rafines- 
gquianum. The Chinese V. Aanceanum appears to be allied closely to this 
complex. When the plants studied have produced fruit and the identifica- 
tion has been checked, a more valid interpretation of V. kanceanum may 
result. 


This cytotaxonomic complex in eastern North America and the two com- 


164 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


plexes in Asia may be representative of those that exist in other species 
groups of this genus. The author is inclined to believe that a similar com- 
plex may exist with V. cassinoides, V. nudum, and V. lentago of eastern 
North America. 


PHYLOGENETIC RELATIONSHIPS 


The phylogenetic relationships within a genus can be accurately formu- 
lated only when the number, morphology, and behavior of chromosomes 
are correlated with anatomy, morphology, and taxonomy of the species. A 
tentative scheme for the species of Viburnum has been constructed utiliz- 
ing all the available data (Fic. 16). 

The genus Viburnum includes polyploid series with the basic numbers of 
eight and nine. Polyploidy can take place effectively only in one direction; 
the diploid must nearly always be the parent of the polyploid (11, 33). 
Stebbins (37) emphasizes the point that diploid members must be older 
than the polyploids, although they are not necessarily more primitive in 
the sense that they are less specialized in structure. Since higher polyploids 
(tetraploids, hexaploids, etc.) usually cannot revert to the diploid without 
abnormalities in the reproductive cycle, it is likely that one of the lower 
gametic numbers, i.e., 8 or 9, is primitive. 

Wilkinson (44) presented additional evidence of the natural relation- 
ships of selected species in the genus. She reported on fourteen species, 
representing all but two of the sections of the genus, which were placed in 
five groups on the basis of their internal morphology and the vascular 
anatomy of their flowers. Viburnum sieboldii (n = 8) is the most primitive 
of those studied, with only two characteristics that might be considered ad- 
vanced: reduction of peripheral bundles to five and the reduction of the 
sepal supply to a single unbranched trace. In no other species included in 
her study were so many primitive characteristics present. A group of 
relatively primitive species includes V. carlesti, V. lantana, and V. denta- 
tum, all n = 9. Another group of less primitive species would include V. 
lantanoides (n = 9), V. plicatum {. tomentosum (n = 9), and V. lentago 
(x = 9). The group of more advanced nine-chromosome species includes 
V. rhytidophyllum, V. nudum, V. cassinoides, V. dilatatum, and V. trilobum. 
Viburnum opulus (n = 9) is considered the most advanced. Her work 
supports the theory that the species with the basic number of eight are 
primitive, while those with a basic number of nine are more advanced. 

From a study of the stem anatomy, De Vos (12) concurred that V. opu- 
lus, V. lentago, and V. cassinoides are the most advanced and that V. pli- 
catum {. tomentosum, V. lantanoides, and V. sieboldii are the most primi- 
tive. 

To date, this is the extent of comparative morphological and anatomical 
studies of Viburnum species. Viburnum sieboldii (n = 8) is on these 
grounds considered to be the most primitive species of the genus. All other 
species with a basic number of eight have many characteristics in common 
with V. sieboldii, so that it can be assumed that this group Is more primitive 


1962 | EGOLF, CYTOLOGY OF VIBURNUM 165 


*— SPECIES WITH MORE come 
HAN ONE CHROMOSOME 
NUMB 


V. bracteatum 
tu 


2n = 40 


* V. adoratissimum 


srubescens 


" 
<<<<<< 
* ox N 

=. 
a 
2 


* 


2n = 27 
* V. lobophyllum 


x V. lantana In =18 
V. acerifolium * V.lantona 
V. atrocyaneum V. lantanoides 
V. betulifolium V. lentago 
V. bitchivense * V. lobophyll 
V. buddleifolium V. macrocephalum 
V. burejaeticum V. microphyllum 
V. x burkwoodii V. mongolicum 
Vic V. nudum 
Vix “corleephalom V. obovatum 
*V. carlesii V. opulus 
V. x carlotta V. orientale 
V. cassinoides V. ovatifolium 
chenaultii V. parvifolium 
V. cinnamomifolium * V. plicatu 
coria V. propinquu 
V. cotinifolium V. prunifoliu 
yonthum V. x rhytidocarpum 
vidii V. x rhytidophylloides 
V. dilatatum V. chytidophyllum 
pad . edul V. rigidum 
Q@n=18 V. ellipticum V. rufidulu 
V. eros V. saorgent 
V. fHlavescens V. schensianum 
foetidu V. sempervirens 
V. furcatum * V. setigerum 
. glomeratum V. stellulatum 
=16 V. harryanum Vv. sympodial 
Von bodaenteare V. hirtuly V. trilobum 
hupehense V. urceolatum 
V. foetens , ' 
V. ichangense V. utile 
V. fragrans V. x jockii V. veitchii 
V. grandiflorum 
V. joponicum V. wilsonii 
* V. plicatum V. x juddii Vv. ig 
iv sieboldt V. kansvense Unidentified 
V. suspensum 


Fic. 16. Diagram alan on chromosome complements to show interrelation- 
ships of species of Viburn 


166 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLIII 


than those species with a basic number of nine. But the possibility re- 
mains that another species within the m = 8 group may be more primitive 
than V. sieboldii, and likewise that V. opulus need not necessarily be the 
most advanced species of the genus. Only a small sample of species and 
of the variation that exists in this widely distributed genus has been 
evaluated critically. Opinion concerning the relative position of species 
in regard to primitiveness may shift when additional evidence is available, 
but the basic cytological relationships of the various chromosome groups 
appear to be well established. 

On the basis of floral anatomy Wilkinson (43, 45) postulates that there 
are in the Caprifoliaceae two lines of development. Through a form re- 
sembling the prototype postulated for Viburnum one line leads to Viburnum 
and Sambucus; the other stems through Leycesteria and branches sepa- 
rately to the Loniceraeae and Linnaeeae. The evidence at hand is in- 
adequate to hazard an interpretation or conclusion as to what was the 
prototype for the family and from whence Viburnum arose, but the 
present evidence favors the » = 8 forms as the more primitive. 


ALTERATION OF BASIC CHROMOSOME NUMBER 


Evolutionary changes within a genus may be due to polyploidy, to the 
addition or subtraction of one or a few chromosomes of a complement, to 
gross structural rearrangements of the chromosomes, to submicroscopic 
changes, probably involving the chemistry of the chromosomal material, or 
to any combination of these. It can be assumed that all these changes have 
probably functioned in speciation in Viburnum. However, the genus has 
not been studied sufficiently to ascertain the evolutionary significance of 
each. For this reason, this discussion of phylogenetic relationships will be 
centered primarily around the evolutionary significance of the basic 
chromosome numbers of the genus. 

Navashin (25) realized that changes in the basic number must involve 
loss or gain of the existing kinetochore, since kinetochores or kinetochore 
modifications cannot arise de novo. In this light, the increase or decrease 
in chromosome number attributed to “fragmentation” and “fusion” (7) 
could occur only when it involved a gain or loss of the kinetochore. 

Darlington (8) presented a procedure favoring the loss or gain of a 
chromosome by means of an equal translocation between two different 
nonhomologous chromosomes with subterminal kinetochores. An inter- 
change involving the long arm of one and the short arm of the other would 
produce one long metacentric chromosome and one very short chromo- 
some or fragment. It has been pointed out that the consequences of un- 
equal translocation depend on whether the regions about the kinetochore 
are genetically active or inert. An inert centric fragment may be elimi- 
nated, with a consequent reduction in chromosome number. If the fragment 
chromosome is genetically active it may persist as a univalent and be 
passed at meiotic metaphase to the same pole as the other interchange 
chromosome. This will yield, in addition to normal gametes, gametes 


1962] EGOLF, CYTOLOGY OF VIBURNUM 167 


with both or none of the interchange chromosomes. The union of gametes 
with additional chromosomes may yield trisomic or eventually tetrasomic 
plants which evolve into species with permanently increased basic numbers. 
Morphological differences may arise by virtue of either dosage effect or 
by divergent gene mutation in the duplicated chromosomes. Further cyto- 
logical divergence may arise from reciprocal translocation between one 
of the new extra chromosomes and another chromosome of the comple- 
ment (5, 37, 1). 

Tobgy (41) confirmed Darlington’s postulate with the demonstration 
that a Ea: reciprocal translocation between two chromosomes of Crefis neglecta 
gave rise to one chromosome of Crepis fuliginosa. Of the two 
chromosomes resulting from this translocation, the one with a genetically 
inactive region adjacent to the kinetochore was lost. Likewise, Sherman 
(34) obtained evidence that the origin of the Crepis kotschyana (n = 4) 
complement involved reciprocal translocation in the reduction from five to 
four pairs of chromosomes 

Chromosome number can also be increased or decreased by aberrations 
such as the translocations observed by Thomas (40) in the meiotic cycle. 

Asynapsis, desynapsis, nondisjunction, and chromosome lagging may be 
responsible for the production either of gametes with a single extra 

mosome, several extra chromosomes, or with the entire unreduced 
eeicntee or of other gametes with chromosomal deficiencies. The 
union of such gametes may result in individuals deficient in or with addi- 
tional chromosomes. 

Chromosome numbers may be increased by supernumeraries or by mis- 
division of the kinetochore. White (42) considers the formation of super- 
numeraries, fragments produced by deletion or translocation, and frag- 
mentation of the kinetochore to be probably the chief method whereby 
chromosome numbers have become increased in the course of evolution 
in animals. The fragment lacking the kinetochore region is lost in sub- 
sequent divisions unless it is translocated to another chromosome. Thus, 
fragmentation, in association with translocation, provides a mechanism 
or chromosome number increase. According to Darlington and Mather 
(11), misdivision of the kinetochore is the only single change that can 
affect both the number and structure of the chromosomes in a single 
stroke. The kinetochore, rather than dividing lengthwise, divides cross- 
wise, resulting in two telocentric chromosomes which at a later division 
may produce two pairs of unlike isochromosomes (9). Races of Campan- 
ula persicifolia (11) have been found in which two telocentric chromo- 
somes occur instead of a single chromosome and thus add an additional 
chromosome to the haploid number 

The backcrossing of a triploid, produced by a cross between a tetra- 
ploid and diploid plant, to a diploid has experimentally produced a great 
variety of segregant types, from among which have been recovered a small 
proportion of fertile types. Examples of such results have been reported 
in Triticum by O’Mara (26) and in Gossypium by Beasley and Brown 
(3) and have been summarized for Nicotiana by Goodspeed (17). This 


168 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


system of hybridization has artificially changed the basic chromosome 
number but has not been established as occurring in the natural evolution 
of species. 

These examples individually, or in combination, illustrate numerous 
means whereby the basic chromosome number could increase or decrease 
as the taxonomic group evolved. Likewise, the progressive increase or 
decrease in basic number could be followed or accompanied by amphi- 
diploidy to produce a complicated aneuploid series such as occurs in 
Carex, Iris, Sedum, Viola, and other genera. Stebbins’ (37) summary of 
the types of aneuploid series in higher plants shows that the number of 
descending basic series is far greater than either the ascending basic or 
interchange amphidiploid series. 

It is possible that in Viburnum the change in the basic chromosome 
number from eight to nine was adequate to keep the two types isolated 
and to allow each to evolve independently. As the types were exposed to 
new and changing environments the selection pressure further subdivided 
the major groups into minor groups with differential adaptability. It is 
conceivable that under certain circumstances the polyploid was favored 
over the diploid, or in others vice versa, thus increasing the variability. 
During this interval adaptive mutations could arise in certain subgroups. 
As the minor groups became further subdivided and isolated for survival 
in specific environmental niches, the variability within the genus expanded 
until today many Viburnum species have great morphological divergence. 
Although the phylogenetic relationships suggested by the chromosome 
numbers of Viburnum species provide a basic framework from which the 
specific differences evolved, the pathway remains obscure. 


CHROMOSOMES AND THE TAXONOMIC SECTIONS 


The nine taxonomic sections of the genus, based on morphological 
characters, as recognized by Rehder (30) may be correlated with the 
pattern of cytological relationships. Section THyrsosmMa, which includes 
Viburnum sieboldii, is composed entirely of species with a basic chromo- 
some number of eight. This number occurs in only one other section, 
PsEUDOPULUS, in which, however, V. plicatum and V. plicatum {. tomento- 
sum have forms with both » = 8 and m = 9. The question arises as to 
whether V. plicatum and V. plicatum {. tomentosum, both with cytological 
forms that are morphologically indistinguishable, represent the connecting 
link in the evolution of the genus between the basic numbers of eight and 
nine. Wilkinson (43) concluded from the study of floral anatomy and 
morphology that V. plicatum f{. tomentosum was relatively primitive, but 
not as primitive as V. carlesti,n = 9, of sect. LANTANA, and V. dentatum, 
of sect. Opontotinus. This does not support the proposition that V. 
plicatum forms an evolutionary bridge between the groups of species 
with m = 8 and » = 9. It is probable that » = 9 may have evolved more 
than once and in various places. However, with additional study this 
relationship may be clarified. 


1962 | EGOLF, CYTOLOGY OF VIBURNUM 169 


Only diploids are found in sects. PsEUDOPULUS, LENTAGO, MEGALOTINUS, 
Oputus, and LaNnTANA, with the exception of V. carlestt (2n = 18, 20, 
22) and V. lantana var. rugosum (2n = 27 

Tetraploids and higher polyploids are found in sects. THyRsosMa, TINUS, 
and OpontToTINus. The scheme showing the relationship between chromo- 
some complements is presented in Fic. 16, which shows the evolutionary 
trend to be from 2m = 16 to 2m = 32, and from 16 to 18 to 36 to 72. 
In the evolution of the genus the diploids probably have had the highest 
adaptive value and today are represented by the largest number of species. 
At the present time it appears impossible to separate the diploid species 
into taxonomic sections on the basis of chromosome morphology. The 
distinct gross morphological differences used by the taxonomist to divide 
the genus into sections appear to be the result of genic rather than 
structural chromosomal changes. When karyotype analysis has been com- 
pleted for these species, differences of arm length, secondary constrictions, 
kinetochore position, satellites, and size of chromosomes may reveal 
natural relationships between species and sections. 

A study of the chromosome complements of polyploid species reveals 
that the genome is duplicated. These species, preceded by an asterisk in 
Fic. 16, provide additional evolutionary information. For example, in 
sect. ODONTOTINUS the tetraploid V. setigerum has the same genomes 
duplicated that occur in the diploid. In the same section, V. dentatum 
var. pubescens and V. dentatum var. deamit are represented by octoploid 
forms with eight genomes duplicating the four genomes of the tetraploid. 
Viburnum carlesii (sect. LANTANA) and V. plicatum, V. plicatum {. mariesit, 
and V. plicatum f{. tomentosum (sect. PSsuDopULUS) are the only aneu- 
ploid species yet found in Viburnum. The forms with 2n = 20 and 2n = 
22 can be considered to have developed from the 2m = 18 form which 
is the most common. The plants with 20 and 22 chromosomes have, 
respectively, one and two pairs of chromosomes not found in the 2” = 18 
form, but at present the origin and relationship of these additional chromo- 
somes to the usual 18 in V. carlesii is unknown. 

The geographical distribution of polyploids is much more restricted 
than that of diploids. Viburnum species from all the major centers of 
distribution, except Central and South America, are well represented in 
this study. Since the origin of most of the varieties, whether natural or 
by man’s selection, is uncertain, in many cases they cannot be assigned 
to a specific geographic area and are omitted in the following discussion. 
The greatest number of species studied is in the 18-chromosome group, 
and these are distributed over a wide geographic area. The diploid species 
include fifty from Asia, four from Europe, and twelve from North 
America, while the polyploids include eight species from Asia Minor, 
one from Europe, and six from North America. 

From the foregoing it is obvious that the distribution of polyploids in 
Viburnum provides little evidence for one distinct center of origin of 
the genus. It is probable that polyploidy has evolved several to many 
times and in various places: eastern Asia, the Himalayan and Mediter- 


170 JOURNAL OF THE ARNOLD ARBORETUM [VOL, XLIII 


ranean regions and eastern North America. In eastern Asia occur all the 
species with the basic number of eight, along with a large number of 
diploids and a few polyploid forms with the basic number of nine. The 
Mediterranean species are few in number and belong to the group with 
a basic number of nine. The eastern North American species include the 
greatest number of octoploids, possibly indicating that this geographical 
niche has been more favorable for their evolution and establishment. 

Only for certain sections does a relationship exist between the geographi- 
cal distribution and the taxonomic sections of the genus. Sections THyr- 
SOSMA and MEGALOTINUS are entirely of Asiatic distribution. Viburnum 
plicatum and its varieties, composing sect. PSEUDOPULUS, are native only 
to China and Japan. All species of sect. LENTAGO are limited to eastern 
North America. Representative species of sects. Tinus and LANTANA are 
distributed both in Asia and in Europe. Sections Oponrotinus and 
PSEUDOTINUS are represented both by North American and Asiatic species. 
The species of the sect. OpuLtus occur both in Europe and in North 
America. 

At present, the cytological evidence suggests that Rehder’s sectional 
classification of Viburnum corresponds favorably with the natural rela- 
tionships. It is hoped that as this study is continued and expanded a 
more accurate evaluation of the classification can be achieved. 


PROPOSED RESEARCH 


Portions of the preceding discussion are based principally on inference 
which indicates where the problems lie and suggests methods of approach. 
Definite conclusions cannot be drawn until much additional research is 
completed. Therefore, the present study is basic both for plant breeding 
and for cytological studies to be continued in the genus Viburnum. 

To the present, it has been impossible to secure the species native 
to Mexico and Central and South America, but by expedition or other- 
wise, it is hoped that these may become available for future study. Within 
these areas are many species of diverse form which, when secured from 
higher elevations, should prove hardy and noteworthy ornamentals in this 
and other latitudes. These species not only may provide additional 
genetical variability for interspecific hybridization, but also are repre- 
sentatives of one of the centers of diversity in the evolution of the genus. 

Cytological studies, in addition to providing a basis for plant breeding, 
have provided a useful tool for taxonomists in classifying certain plant 
groups, and there is every indication that such information can likewise 
be useful in studies of Viburnum. It is anticipated that a sporocyte study 
and karyotype analysis, associated with genetical and taxonomic studies, 
will aid materially in revealing natural relationships which can be utilized 
in the classification of the genus Viburnum. 


— 


U.S. NationaL ARBORETUM, 
WASHINGTON 25, D.C. 


1962 | EGOLF, CYTOLOGY OF VIBURNUM 171 


BIBLIOGRAPHY 


_ 


Bazscock, E. B. Cytogenetics and speciation in Crepis. ae pee in 
Genetics, pp. 161-206. Academic Press, Inc., New York, 

BaLpwin, J. T., Jr. Kalanchoe: the genus and its Fete Am. 
Jour. Bot. 25: 572-579. 193 

BEASLEY, J. O., & M. S. Brown. ‘The production of plants having an extra 
pair of chromosomes from species hybrids of cotton. Rec. Genet. Soc. 
Am. 12: 43. 1943 

Bake, S. F. On the names of some species of Viburnum. Rhodora 20: 
11- 15. 

CLAUSEN, JENS. Stages in the evolution of plant species. 206 pp. Cornell 
Univ. Press, Ithaca, N.Y., 1951. 

Concer, A. D. & L. M. Farrcuiip. A quick freeze method for making 
smear slides permanent. Stain Tech. 28: 281-283. 1953. 

Dartincton, D. C. Chromosomes and plant breeding. 112 pp. Macmillan 
and Co., London, Z 

. Recent ee in cytology. Ed. 2. 671 pp. J. and A. Churchill, 


bo 


Ww 


al 


On 


oy 


“I 


8. 
Ltd., London, 

9, ———. oe and the genetics of the centromere. Jour. Genet. 37: 
341-364. 1939. 

10: & L. F. La Cour. The handling of chromosomes. Ed. 2. 180 pp. 
George Allen and Unwin, Ltd., London, 1947. 

ila & K. THER. The elements of genetics. 446 pp. Macmillan Co., 


New York, 1949. 

12. DE Vos, F. The stem anatomy of some species of the Caprifoliaceae with 
reference to phylogeny and identification of the species. 81 pp. Ph.D 
thesis, Cornell University, 1951. 

13. Dippet, L. Handbuch der Laubholzkunde Beschreibung der in Deutsch- 
land heimischen und im Freien kultivierten Baiime und Straucher 1: 
172-200. 1889. 

14. Ecotr, D. R. Cytological and interspecific okies ane in the 
genus Viburnum. 131 pp. Ph.D. thesis, Cornell University, 

15. Fenc, Y. A. Observations sur la présence de centrosomes s U'asters 
présidant 4 la caryocinése dans un genre de Caprifoliacées: Lonicera. 
Le Botaniste 26: 3-20. 

16. Fernatp, M. L. Another oilitier of additions to the flora of Virginia. 
Rhodora 43: 635-657. 1941 

17. GoopspEeD, T. H. The genus Nicotiana. 536 pp. Chronica Botanica Co., 

altham, Mass., 1954. 

18. Janaki AmMAL, E. K. Chromosomes and the species problem in the 
genus Viburnum. Curr. Sci. Bangalore 22: 4-6. 1953. 

19. JoHANSEN, D. A. Plant eng 533 pp. McGraw-Hill Book 
Co., New York and London, 

20. La Cour, L. Acetic-orcein a new a fixative for chromosomes. Stain 
Tech. 16: 169-174. 1941. 

21. McAtTer, W. L. A review of the nearctic Viburnum. 125 pp. Chapel Hill, 
North Carolina, 1956. 

22. Mever, J. R. Prefixing with paradichlorobenzene to facilitate chromo- 
some study. Stain Tech. 20: 121-125. 1945. 

Naxal, T. Tentamen systematis Caprifoliacearum japonicarum (System- 


wn 
oO 


bo 
Ww 


JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


atic treatment of sg i guages of Japan), Tokyo Imp. Univ. Jour. 
Coll. Sei. 42> ‘Art. 2; 139 pp. 1. 

Classes, sate familiae, subfamiliae, tribus, genera nova quae 
attinent ad plantas Koreanas. Jour. Jap. Bot. 24: 8- 4: 1949, 


. NAvASHIN, M. The dislocation hypothesis of evolution of chromosomal 


numbers. Zeits. Ind. Abst. Ver. 63: 224-231. 1932. 

O'Mara, J. G. Cytogenetic studies on Tyriticale. I. A method for de- 
termining the effects of individual Secale chromosomes in Triticum. 
Genetics 25: 401-408. 1940. 

Poucques, M. L. pe. Etude caryologique de quelques Viburnum. Compt. 
Rend. Soc. Biol. Paris 141: 183-185. 

———. Recherches caryologiques sur les Rubiales, Revue Gén. Bot. 56: 
5-27, 74-138, 172-188. 1949. 

REHDER, A. New species, varieties and combinations from the herbarium 
and the collections of the Arnold Arboretum. Jour. Arnold Arb. 5: 
49-59. 1924. 

—. Manual of oo trees and shrubs. Ed. 2. 996 pp. Macmillan 

Co., New York, 

A oc of cultivated trees and shrubs. 825 pp. Arnold 

Arboretum, Harvard University, 1949. 

Sax, K. & D. A. Kriss. Chromosomes and phylogeny in Caprifoliaceae. 
coe Arnold Arb. 11: 147-153. 1930, 

SHARP, L. W. Introduction to cytology. Ed. 3. 567 pp. McGraw-Hill Book 
Co., New York and London, 1934. 

SHERMAN, M. Karyotype evolution: a cytogenetic study of seven species 
and six interspecific hybrids of Crepis. Univ. of Calif. Publ. Bot. 18: 
369-408. 1946. 

SIMONET, M., & C. Miepzyrzecxi. Etude caryologique de quelques 
espéces arborescentes ou sarmenteuses d’ornement. Compt. Rend. Soc. 
Biol. Paris 111: 969-973, 1932. 

STEARN, W. T. X Viburnum bodnantense, Caprifoliaceae. Bot. Mag. 167: 
t. 113. 1950. 

STEBBINS, G. L., Jr. Variation and evolution in plants. 643 pp. Columbia 
Univ. Press, New Yor 

Suciura, T. Studies on the chromosome numbers in higher plants with 
special reference to cytokinesis. I. Cytologia 7: 544-595. 1936. 

Svenson, H. K. Plants of the ra United States. I. Viburnum 
dentatum. Rhodora 42: 1-6. 


. Tuomas, J. L. The cytology of some cultivated species of Viburnum. 
1961. 


Jour. Acold Arb. 42: 157-164. 


. Topcy, H. A. A cytological study of Crepis fuliginosa, Crepis neglecta 


and their F, hybrid and its bearing on the mechanism of phylogenetic 
reduction in chromosome number. Jour. ees 45: 67-111. 1943. 


. Wuire, M. J. D. Animal cytology and evolution. 454 pp. Cambridge 


Univ. Press, Cambridge, 1954 


. Wivxinson, A. M. The floral anatomy and morphology of some species 


of Cornus and the Caprifoliaceae. 148 pp. Ph.D. thesis, Cornell 

University, 1945. 

: | anatomy and morphology of some species of the genus 

Viburnum of the Caprifoliaceae. Am. Jour. Bot. 35: 455-465. 1948. 
Floral anatomy and morphology of Triostewm and of the Capri- 

foliacéae in general. /bid. 36: 481-489. 1949. 


1962] BRIZICKY, SIMAROUBACEAE AND BURSERACEAE 173 


THE GENERA OF SIMAROUBACEAE AND BURSERACEAE 
IN THE SOUTHEASTERN UNITED STATES * 


GEORGE K. BRIZICKY 


SIMAROUBACEAE A. P. de Candolle, 
Ann. Mus. Hist. Nat. Paris 17: 422. 1811, “Simarubeae.” 
(QuassIA FAMILY) 


Trees and shrubs [rarely subshrubs], often with bitter bark. Leaves 
usually alternate, never glandular-punctate, pinnately compound or simple 
[rarely rudimentary], with predominantly entire margins, exstipulate [or 
eae Flowers usually small to minute, greenish or variously colored, 

ogynous, regular, bisexual or Griseauel by abortion, or both, with 
biseriate, [3]4— —6[8]-, usually 5-merous perianth, in terminal and/or 
axillary many- or few-flowered cymose panicles or racemes, rarely solitary. 
Sepals distinct or united, imbricate or valvate. Petals distinct, imbricate 
or valvate, rarely wanting, Stamens distinct, as many or twice as many as 
the petals [rarely more numerous], usually inserted at base of an intra- 
staminal disc, rudimentary or wanting in @ flowers; filaments usually 
slender, sometimes appendaged at base on the ventral (adaxial) side; 
anthers usually versatile, 2- or rarely 4-locular at anthesis, introrse, fone 
tudinally dehiscent. Intrastaminal nectariferous disc usually present, an- 
nular, cupular, cushion-like [to columnar], mostly lobed or crenate, rarely 
obscure or wanting. Gynoecium usually inserted on or encircled at base 
by the disc, sessile [or rarely raised on a gynophore], 2-6[8]-carpellate, 
apocarpous to syncarpous, rudimentary or wanting in ¢ flowers; stigmas 
distinct or united; styles basal, lateral or apical, distinct or partially to 
completely united; ovaries | -carpellate and -locular (gynoecium apocar- 
pous) or 2—3[4]- -carpellate and -locular with axile placentae (gynoecium 
syncarpous); ovules anatropous or rarely orthotropous to campylotropous, 
usually epitropous, very rarely apotropous, 2- or rarely 1- integumented, 
with a thick nucellus, 1 or 2 (collateral or superposed) [very rarely more 

1 Prepared for a generic flora of the southeastern United States, a joint project of 
the Arnold Arboretum and the Gray Herbarium of Harvard University which has 
been made possible through the support of Ge eorge R. Cooley and the National 
Science Foundation. This treatment follows the pattern established in the first paper 
in the series (Jour. Arnold Arb. 39: 296-346. 1958) and continued through the 
seventeen subsequent papers in volumes 40-43 (1959-1962). It should be repeated 
that the area covered by this work is bounded by and includes North Carolina, Ten- 
nessee, Arkansas, and Louisiana. The descriptions are based primarily on the plants 
of this area, with any supplementary material in brackets. References which the 


The author is indebted to Dr. Carroll E. Wood, Jr., for his criticism and valuable 
suggestions, and to Mrs. Gordon W. Dillon, for her careful help in the preparation 
of the manuscript. 


174 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLII 


numerous| in a locule, pendulous from the top or ascendant from the 
base of the latter. Fruit apocarpous, of 2—6|8] one-carpellate and -locular 
drupes, [berries] or samaras, or a syncarpous, 2—3|4|-carpellate and 
-locular (occasionally 1-locular by abortion) berry, samaroid capsule 
[drupe or schizocarp| with 1-seeded locules. Seeds with membranaceous 
or leathery testa; endosperm scanty or wanting; embryo rather large, 
straight or rarely curved, with narrow, mostly fleshy, plano-convex or flat 
cotyledons and usually a very short radicle. (Including Surianaceae.) 
Type GENUS: Simarouba Aublet. 


A pantropical family of about 30 genera and 200 species, at least two 
species in temperate eastern Asia. Four of twelve genera indigenous to 
the New World occur in subtropical Florida. One species of Adanthus is 
also naturalized in our area. 

The rather heterogeneous family includes six sharply delimited sub- 
families. Evidence from wood anatomy supports the segregation of at 
east some as distinct families, but, since data from gross morphology 
appear to be inconclusive and those from floral anatomy, embryology, 
and cytogenetics of the family are fragmentary, it seems preferable to 
retain the family in the larger sense. 

Simaroubaceae are closely related to Rutaceae and Burseraceae, differ- 
ing from the former mainly in the absence of multicellular glands (secre- 
tory cavities) with aromatic oils in leaves, axes, and floral parts, and from 
the latter in the absence of schizogenous resin ducts in the bark. 

Wood and/or bark of some species (e.g., Picrasma excelsa (Sw.) Planch., 
“Jamaica Quassia”’; Quassia amara L., “Surinam Quassia”) yield a bitter 
principle employed as tonics and vermifuges, as an insecticide, and some- 
times as a substitute for hops in brewing. Some genera produce timber 
of local importance; a few species are ornamentals. 


REFERENCES: 

Boas, B. Beitrage zur Anatomie und Systematik der Simarubaceen. Thesis, 58 
pp. Dresden. 1912. {Also in Beih. Bot. Centralbl. 29(1): 303-356. 1913.] 

Cronourist, A. Studies in Simaroubaceae—IV. Résumé of the American 
genera. Brittonia 5: 128-147. 1944. [Suriana, Recchia, Alvaradoa. Hola- 
cantha, Picrasma, Picrolemma, and Quassia are treated in detail. | 

DEsat, S. Cy tology of Rutaceae ane Simarubaceae. Cytologia 25: we 35. 1960. 
| Ailanthus altissima, 2n = 80, Quassia amara, 2n = 36. | 

ENGLER, A. Simarubaceae. . Paanzentani. III. 4: 202-230. 1897; ibid. ed. 
2. 19a: 359-405. 1931. 

Harti, D. Die Ubereinstimmungen des Endokarps der Simarubaceen, Rutaceen, 
und Leguminosen. Beitr. Biol. Pil. 34: 453-455. 1958 

Japin, F. Contribution a l'étude des Simarubacées. in. a Nat. Bot. VIII. 13: 
201-304. pl. 1. 1901. 

——-—. Essai de classification des Simarubacées basée sur les caracteres anato- 
miques. Compt. Rend. Assoc. Fr. Avanc. Sci. 30(2): 477-483. 1902.* 
[See review in Bull. Soc. Bot. Fr. 49: 223, 224. 1902. ] 

Narr, N. C., & T. S. Josepu. Floral morphology and embryology of Samadera 
indica. Bot. Gaz. 119: 104-115. 1957. 


1962 | BRIZICKY, SIMAROUBACEAE AND BURSERACEAE 175 


& R. K. JosH1. Floral morphology of some members of the Simarou- 
baceae. Bot. Gaz. 120: 88-99. 1958. [Five spp. of Adanthus, including 
A, altissima ; Picrasma quassioides (D. Don) Benn.. and Brucea sumatrana 
Ox as amarissima (Lour.) Desv. ex Gomes). | 


& N. P. SukUMARAN. Floral morphology and embryology of Brucea 
amarissima. Bot. Gaz. 121: 175— ee 
NARAYANA, L. L. Embryology of two Siniaroubeccae Curr. Sci. Bangalore 26: 


323, 324. 1957.* [Includes ee excelsa. | 
& M. SAYEEDUDDIN. Floral anatomy of Simarubaceae — I. Jour. Indian 
Bot. Soc. 37: 517-522. 1958. gee amara L. and Ailanthus excelsa.| 
Recorp, S. J.. & R. W. Hess. Timbers of the New World. 640 pp. ey 59. 
New Haven. 1943. [Simaroubaceae, re 514; Gane S2o22.4 
SARGENT. C. S. Manual of the trees of North ie (exclusive of Mexico). ed. 
2.897 pp. 1922. [Simaroubaceae. 641- 
SAUNDERS, E. R. Floral morphology. Vol. 2. New York. 1940. [Simaroubaceae. 
195-197. 
SMALL, J. K. Simaroubaceae. N. Am. Fl. 25: 227-239. 1911. [Suriana placed 
in a Separate family. | 
WEBBER, I. E. Sy stematic anatomy of the woods of the Simarubaceae. Am. Jour. 
Bot. 23: 577-587. 1936. 
WEST. Ee. & L. E. ARNOLD. The native trees of Florida. 212 pp. Gainesville. 
1946. | Simaroubaceae, 101-104. | 
Wicer. J. Embrvological studies on the families Buxaceae. Meliaceae. Simaru- 
baceae and Burseraceae. Thesis. Lund. 1935.* [See Mauritzon’s criticism. 
935: 490-502. 1935; Wigers reply. Bot. Not. 1936: 585-539. 
1936. | 


KEY TO THE GENERA OF SIMAROUBACEAE 


General characters: trees or shrubs with alternate. predominantly pinnate 
leaves; flowers small, uni- or eee or both; perianth st usually 5-merous ; 


stamens 5 or 10, distinct; intrastaminal aie usually present; gynoecium apo- 
or synearpous, 2-5- parpelluter ovules 1 or 2 in each eee 


A. Leaves simple. small (not over 5 cm. long) and narrow: flowers bisexual. in 
short. terminal, few-flowered. corvmb-like panicles. rarely solitary; stamens 
10. the 5 antipetalous usually ag or rudimentary: intrastaminal disc 
wanting; carpels distinct; fruit of r fewer achene-like drupelets: strand 
Dlaniwor subtropical Plordac ss exces ee eety Fee eee ace te 1. Suriana,. 

A. Leaves pinnately compound; flowers usually unisexual (occasionally also bi- 
sexual) in large, many-flowered, complex terminal panicles or racemes; in- 
trastaminal disc distinct; carpels connate. sometimes only by the styles. 
rudimentary or wanting in 4 flowers; fruits various, never achene-like 
drupelets 
B. Stamens 10 in 6 flowers. much reduced or wanting in @ flowers; carpels 

4—6, usually 5; styles lateral. connate; ovaries distinct, connivent; fruits 

aggregate. 

C. Leaves usually even-pinnate. persistent; leaflets alternate. + leathery. 
entire; staminal filaments appendaged at base; fruit of 5 or fewer 
olive-shaped drupes about 2 cm. long: hammocks of subtropical 
Florida. 2. Simarouba. 


176 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


C. Leaves odd-pinnate, deciduous; leaflets usually opposite, thin, with a 
few glanduliferous teeth or lobules near the base; staminal filaments 
without appendages; fruit of 5 or fewer oblong samaras 3-5 cm. long, 
each with a flattened seed at the middle of the membranaceous wing; 
northern Florida and northward. .................. 3. Ailanthus. 

B. Stamens 5 and alternate with the sepals in 6 flowers, reduced or wanting 
in 2 flowers; carpels 2 or 3; styles terminal, connate or distinct; ovary 
compound; ieives odd- epinnates hammocks of subtropical Florida. 

D. Leaflets 5-9, 5-12 cm. long; ¢ and @ flowers with linear-lanceolate 
petals, in terminal panicles with slender racemose or spikelike branches; 
stigma 2-lobed; style one, very short and stout; fruit a 1- or 2- joe 
EOE: 4 cyan waewae eee epider ene ee sea e ook 4. Picramnia. 

D. Leaflets 19-51, 1-3 cm. long; 4 flowers with linear-filamentous ae 
the ? flowers without petals, in terminal and axillary racemes; stigmas 
3; styles 3, distinct, subulate, short; fruit a flat, papery, 1- seeded sama- 
roid manele: be kde whe 45 odor hateueees cee pans 5. Alvaradoa. 


Subfam. SURIANOIDEAE Engler 
1. Suriana Linnaeus, Sp. Pl. 1: 284. 1753; Gen. Pl. ed. 5. 137. 1754. 


Shrubs or small trees 1-8 m. high with most parts more or less densely 
pubescent with simple and glandular hairs. Leaves sessile, simple, entire, 
narrow (seldom over 4 cm. long), thickish, with indistinct lateral veins and 
centric mesophyll. Flowers relatively small, bisexual, obdiplostemonous, 
with 2-bracteolate pedicels, in terminal, few-flowered, corymb-like cymose 
panicles (sometimes reduced to a solitary flower). Sepals 5, narrow (about 
6-10 mm. long), connate at base, imbricate, persistent. Petals 5, yellow, 
broad, clawed, about as long as the sepals, imbricate. Stamens 10, distinct, 
the antipetalous usually antherless or rudimentary; filaments subulate, 
hairy below; anthers subcircular in outline, emarginate at both ends, in- 
trorse, 2-locular at anthesis; pollen grains medium sized, suboblate, 3-col- 
pate, finely striate. Intrastaminal disc apparently wanting. Gynoecium 
apocarpous, 5-carpellate; stigmas distinct, capitellate; styles distinct, fili- 
form, nearly basal; ovaries 1-locular, sessile; ovules 2 (collateral), ascend- 
ing from the base of the ovary locule, 1-integumented, orthotropous, be- 
coming campylotropous after fertilization. Fruits 1-seeded achene-like 
drupelets (not over 5 mm. long) with thin flesh and crustaceous endocarp 
(stone), obovoid-subspherical, pubescent. Seeds broadly obovoid, slightly 
flattened laterally, endospermless, with a thin membranaceous testa; em- 
bryo horseshoe shaped, with flat, oblong cotyledons and an elongate radicle 
descendent toward the micropyle which is side by side and in contact with 
the hilum. Type species: S. maritima L. (Named in honor of Joseph 
Donat Surian, a French physician and botanist, “Dioscorides Americanus 
futurus” |[Plumier, Nov. Pl. Am. Gen. 37. 1703], companion and _ col- 
laborator of Plumier.) — BAY CEDAR. 


A monotypic genus represented by Suriana maritima, widely but sporadi- 
cally distributed on the seashores of the New and Old World tropics, but 


1962] BRIZICKY, SIMAROUBACEAE AND BURSERACEAE Liz 


apparently absent from the Pacific coasts of the Americas, the islands of 
the Central Pacific, and the Atlantic coast of Africa. It occurs in the 
coastal sand dunes and hammocks of the Florida Keys and of peninsular 
Florida northward to Brevard and Pinellas counties, and beyond our area 
in Bermuda, the West Indies, and from Yucatan south to Brazil. Mill- 
spaugh (Publ. Field Mus. Bot. 2: 241. 1907) believed the fruits (stones) 
to be carried on the feet of sea birds, but according to Guppy they “could 
readily be carried in the crevices of floating logs, or in the cavities of float- 
ing pumice, such as is stranded on the beaches of tropical regions all over 
the world. But it is on their great floating powers, which fit them for dis- 
persal by currents, that we must mainly rely.” Buoyancy of the stones is 
produced by the unfilled space in the fruit locule (Guppy, Schimper). 
Record & Hess (1943, p. 521), Small (Man. 761. 1933), Wilson (1911), 
and a few others thought the genus to represent a distinct, monotypic 
family Surianaceae; but, if a separate family is recognized, it should also 
include the closely related genera Cadellia F. Muell., Guilfoylia F. Muell., 
and perhaps also Recchia Sessé & Moc. ex DC. (Rigiostachys Planch.). 
The branches exude some kind of ‘“‘manna.” 


REFERENCES: 

See also under family references, Boas, ENGLER (1897, pp. 208, 209; 1931, 
pp. ae 368), JADIN, REcorp & Hess, WEBBER (pp. 577-579, 586, 587), and 
Wi 


IGE 

ee H. B. Plants, seeds, and currents in the West vee and Azores. 531 
pp., 3 maps. bendon: 1917. [Fruit dispersal, 239-24 

Rav, M. A. An embryological study of Suriana maritima a Proc. Indian Acad. 
Sci. B. 11: 100-106. 

cases A. F. W. Die Indo- IMalayische Strandflora. (Botanische Mittheilungen 

us den Tropen. 3.) 204 pp., pls. 1-7. Jena. 1891. [Floating power of 

fe 163, 165. ] 

SOLEREDER, H. Ueber die systematische Stellung der Gattung Rigiostachys, 
zugleich ein Beitrag zur nadheren Kenntnis der Simarubeae-Surianoideae. Jn: 


kanische Gattung Rigiostachys. Verh. Bot. Ver. Brandenburg 47: 41-61. 
1905. [Includes morphology of ovules, fruits, and seeds. | 
WILSON, P. Surianaceae. N. Am. Fl. 25: 225. 1911. 


Subfam. SIMAROUBOIDEAE 
Tribe SIMAROUBEAE 

2. Simarouba Aublet, Hist. Pl. Guiane Fr. 2: 859. 1775.7 
Trees or large shrubs with bitter bark and wood. Leaves even-pinnate 
[or odd-pinnate]|, persistent; leaflets 3-21, mostly 8-16, alternate, rarely 
opposite, entire, -_ coriaceous, shortly petiolulate, Plants polygamo-dioe- 
cious or dioecious Flowers small, unisexual or uni- and bisexual, obdiplo- 
? Aublet’s generic name should be conserved against its earlier homonym, Simaruba 


Boehmer, 1760 [nom. rejic. vs. Bursera Jacquin ex Linnaeus, 1762, nom. cons.]. See 
A. A. Bullock, Taxon 8: 199. 1959. 


178 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLIII 


stemonous, in large terminal and axillary complex panicles. Sepals 4—6, 
usually 5, rather small, connate at least at base, imbricate. Petals 4—6, 
usually 5, yellowish to yellow [or whitish], much longer than the sepals, 
imbricate. Stamens 8-12, usually 10, as long as the petals, reduced, rudi- 
mentary or absent in @ flowers; filaments subulate, broadened toward 
base, with an adaxial, liguliform, pubescent appendage at base; anthers 
oblong in outline, slightly emarginate, versatile, 2-locular at anthesis; pol- 
len grains small, subprolate, 3-colpate. Intrastaminal disc + cushion- 
like, depressed in 6 flowers. Gynoecium (4)5(6)-carpellate, sessile on 
the disc, partially syncarpous, rudimentary or wanting in é flowers; stig- 
mas 4—6, usually 5, long, slender, and divergent in ¢ flowers, short, lobe- 
like in bisexual flowers; styles connate into a short column: ovaries dis- 
tinct, 1-locular, cohering |or perhaps sometimes weakly connate] by their 
ventral sutures and thus resembling a compound deeply lobed ovary; 
ovules solitary in each carpel, pendulous from near the top of the inner 
angle of the locule, anatropous and epitropous, 2-integumented. Drupes 5 
or fewer from each flower, ellipsoid to obovoid, slightly laterally compressed, 
usually 2-ridged, with thin flesh and a crustaceous stone. Seeds endosperm- 
less; testa thin, membranaceous; embryo straight, with fleshy, plano-convex 
cotyledons and a very short, superior radicle macy included between the 
cotyledons (retracted). Typr species: S. amara Aublet. (Name derived 
from the Carib Indian name of the type species in French Guiana.) 


A tropical American genus of about six species. Simarouba glauca DC., 
paradise tree, occurs in coastal hammocks on the Florida Keys and in 
southern peninsular Florida (Dade, Broward, and Palm Beach counties). 
It is widely distributed in the West Indies and in Central America from 
Costa Rica to southern Mexico. The ellipsoidal fruits, about 2 cm. long, 
are scarlet, changing to dark purple or black when ripe. A white-fruited 
form has been reported from El Salvador 

The timber of some species, especially. the South American Simarouba 
amara, is of commercial importance. The bitter bark of roots of S. amara 
and S. glauca is said to be efficient against diarrhea and post-dysenteric 
disorders. Seeds of S. glauca yield about 60 per cent of edible oil and a 
crystalline glycoside, glaucorubin, which reportedly has amoebicidal prop- 
erties and is now being introduced in most of the major tropical areas 
under the trade name of ‘‘Glaumeba.”’ 

The genus is a representative of a very natural group, Simarouboideae— 
Simaroubeae, and is closely related to the tropical South American Simaba 
Aubl. and Quassia L. (monotypic, according to Cronquist), as well as 
to the tropical African Odyendea (Pierre) Engl. 


REFERENCES: 


See also under family references, ENGLER (1897, pp. 211, 213; 1931, pp. 372- 
347), Recorp & Hess (pp. 513, re SARGENT (pp. 643, 644), SMALL (pp. 227— 
229), and West & Rete (p ). 

ArMourR, R. P. spmpeteete on Simarouba glauca DC. in El Salvador. Econ. 


1962 | BRIZICKY, SIMAROUBACEAE AND BURSERACEAE 179 


Bot. 13: 41-66. 1959. [Commercial importance and cultivation as a vege- 
table oil crop. | 

Cronguist, A. Studies in the Simaroubaceae — II. The genus Simarouba. Bull, 
Torrey Bot. Club 71: 226-234. 1944. 

Kryn, J. M. Simarouba, paradise-tree, marupa, Simarouba spp., family: Simarou- 
baceae. U. S. Forest Serv. Forest Prod. Lab. Rep. R. 1956. 10 pp. 1953.* 

KUKACHKA, B. F. Marupa: Simarouba amara Aubl., Simaroubaceae. U. S. Forest 
Serv. Forest Prod. Lab. Rep. 1856. 4 pp. 1960.* 

SARGENT, C. S. Simaruba. Silva N. Am. 1: 89-92. pls. 38, 39. 1891. 

SoLA, F. pE. Notes on the aceituno tree (Simaruba glauca DC.) and its adapta- 
tion as a vegetable oil crop. Ceiba 4: 351-358. 1956. 


Tribe PIcRASMEAE Engler 


3. Ailanthus Desfontaines, Mém. Acad. Sci. Paris 1786: 265. 1788, 
nom, cons 


Trees, sometimes strong smelling. Leaves odd-pinnate, large, deciduous; 
leaflets opposite or alternate, thin, with a few glanduliferous, blunt teeth 
or lobules near base [or entire or rarely coarsely toothed to lobed through- 
out], petiolulate. Plants polygamo-dioecious. Flowers small, uni- and 
bisexual, obdiplostemonous, of unpleasant odor, pediceled, in large, terminal 
panicles. Sepals 5 (6), connate in the lower third or higher, imbricate. 
Petals 5 (6), longer than the sepals, greenish or yellowish, induplicate- 
valvate in aestivation. Stamens inserted at base of an intrastaminal disc, 
10 (12) in 6 flowers, sometimes fewer in bisexual, much reduced and 
sterile or absent in @ flowers; filaments subulate, without appendages; an- 
thers oblong or oblong-ovate in outline, versatile, 2-locular at anthesis, 
subintrorsely dehiscent; pollen grains small to medium sized, subspheroidal, 
3-colpate, reticulate. Intrastaminal disc annular, thick, usually deeply 
5(6)- or 10(12)-lobed, or crenate, of receptacular origin. Gynoecium 
5(6)-carpellate [rarely 3-carpellate], partially syncarpous [or apocar- 
pous|, sessile within the disc, rudimentary or absent in ¢ flowers; stigmas 
distinct, tongue-shaped and divergent or + capitellate; styles lateral, fili- 
form, usually connate [or distinct]; ovaries 1-carpellate and ocular, 
much compressed laterally, distinct but + cohering by their sutures and 
resembling a compound, deeply lobed ovary; ovules anatropous, 2-integu- 
mented, solitary in each carpel, hanging from below the insertion point of 
the style. Fruit of (1)2—5(6) distinct, oblong to oblong-elliptic samaras, 
each with a flattened seed at the middle of the thin, veiny, adaxially emargi- 
nate wing. Seeds lenticular, with thin, membranaceous testa and sparse, 
fleshy endosperm; embryo with flat, obovate to orbicular cotyledons and 
a short, superior radicle. Type species: A. glandulosa Desf. (= A. altis- 
sima (Mill.) Swingle). (Name derived from the Moluccan name for A. 
integrifolia Lam., ailanto, Sea in allusion to the height of the 
trees.) — TREE-OF-HEAVE 


A primarily tropical genus of about 15 species, distributed in eastern 


180 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


and southern Asia, the Philippines, Malaysia, Melanesia (east to the Solo- 
mon Islands), and northeastern Australia, extending into the Temperate 
Zone to northeastern China and Korea (to about 40° N. Lat.). The north- 
ernmost species, Ailanthus altissima (Mill.) Swingle, Chinese tree- of- 
heaven, stinkweed, 2” = 80, introduced into North America in 1784 as an 
ornamental tree, has become naturalized in native woodlands of the south- 
eastern United States as far south as northern Florida. Northward and 
westward beyond our area it seems to be primarily a ‘‘weed” tree of cities, 
but the exact extent of naturalization needs to be more carefully recorded. 
An irregularly branched, rapidly growing tree which reproduces by seeds, 
stump sprouts, and root suckers, it becomes a weed difficult to eradicate. 
However, it has sometimes been used for fastening sterile, sliding declivities 
and for aforestation of bare hill and mountain slopes (Alps, Caucasus) and 
of grasslands (Asia Minor). The leaves contain a substance toxic to seed- 
lings of many species of gymnosperms and angiosperms. “It is possible 
that the toxic substance is washed from the | fallen] leaves by the rain and 
influences the composition of plant communities” (Mergen). 

Small flies and beetles have been recorded as pollinators in Adanthus 
altissima, but bees must participate since it is regarded as an undesirable 
honey plant. Wind pollination is also possible (cf. Wodehouse). Floral 
anatomy has been studied in this and a few other species, embryology only 
in the Indian A. excelsa Roxb., 2n = 62. Apparently only two chromosome 
counts have been recorded. 

Wood of Ailanthus altissima and the other species appears to be of 
relatively little value, being used mostly for fuel, occasionally for cabinet 
work, musical instruments, etc. It also seems to be fit for paper pulp. 
Leaves and bark of A. altissima, and bark of a few other species are used 
locally in Asia as anthelmintics or antidysenterics. 

The genus occupies a somewhat isolated position in the tribe Picrasmeae 
of Simarouboideae, representing a subtribe of its own. 


REFERENCES: 


The large number of references. has been reduced here primarily to those 
either of general interest or dealing > eormaeay| with the southeastern United 
States. Under family references see ENGLER (1931, pp. 390-393), Desat (p. 
39), Narr & JosHI, NARAYANA, NARAYANA & SAYEEDUDDIN, and SMALL (p. 234). 
ANbERSON, E. The tree of heaven Ailanthus altissima 1. A blessing and a 
curse. Missouri Bot. Gard. Bull. 49: 105-107. 1, 

AnpREAE, E. Uber abnorme Wurzelanschwellungen bei Adlanthus glandulosa. 
Thesis, 34 pp., 3 pls. Erlangen. 1894. 

CampREDON, J. Etudes des propriétés physiques et mécaniques de quelques 
bois exotiques. III. Le bois d’Ailante (Ailanthus dees Desf.). Ann. 
Ec. Natl. Eaux Foréts Nancy 5: 211-217. 1934.* 

Davies, P. A. Leaf arrangements in Ailanthus altissima. Am. Jour. Bot. 24: 
401-407. 1937. [See also ibid. 26: 67-74. 9 

; aie glands in Ailanthus altissima. Trans. Ky. Acad. Sci. 11: 12-16. 


: on glands on Ailanthus altissima, Ibid. 12: 31-33. 1945.* 


1962 | BRIZICKY, SIMAROUBACEAE AND BURSERACEAE 181 


& E. Bennett. Abnormal branching in Ailanthus. Jour. Hered. 20: 
348, 349. 1929, 

& E. W. TuHeiss. Factors affecting the method of branching in Ailan- 
thus altissima. Bull. Torrey Bot. Club 64: 229-233. 1937. [See also DAvIEs, 
Bull. Torrey Bot. Club 63: 139-146. pl. 5. 1936.] 

Inyick, J. S., & E. F. Brouse. The Ailanthus tree in Pennsylvania. Penn. Dep 
Forests Waters Bull. 38. 29 pp. 1926. [Includes biology and prospects ie 
the species. | 

Kriz, V., M. CHLEBEK, & M. PEKar. Ailanthus [altissima] from the view- 
point of breeding. (In Czech.) Lesn. Prace 36: 116-118. 1957.* 

MerenpI, A. Ailanthus: cellulose plant. (In Italian.) Gior. Agr. Roma 66: 
137. 1956.* [See also A. pet Lunco, Terra Sole 122: 345-349. 1952.*]| 

MERGEN, F. A toxic principle in the leaves of Ailanthus. Bot. Gaz. 121: 32-36. 
1959. [A. altissima. | 

Miter, R. Zur Anatomie der Ailanthus-Rinden. Pharm. Praxis 7: 261-263. 
1908.* [For review see Bot. Jahresb. 36(1): 473, 474. 1908.] 

Petaj, V. Die extrafloralen Nektarien auf den Bittern as Ailanthus glandu- 
losa. (In Croatian.) Rad Jugosl. Akad. Znan. Umjet. 215: 59-81. 1916.* 
[For review see Bot. Centralbl. 137: 385. 1918. ] 

SayA, I. Ferite su fusto di Adanthus glandulosa Desf. e reazioni di gemme 
dormienti. Nuovo Gior. Bot. Ital. II. 64: 680-682. 1957. [Wounds on 
the stem of A. altissima and reactions of dormant buds. | 

SWINGLE, W. T. The early European history and the botanical name of the 

e of heaven, Ailanthus altissima. Jour. Wash. Acad. Sci. 6: 490-498. 
1916. 

TiEGHEM, P. van. Ailante et Pongéle. Ann. Sci. Nat. Bot. IX. 4: 272-280. 
1906. [Segregation of Pongelion gi from Ailanthus. | 

WopeHousE, R. P. Hayfever plants. xx + 245 pp. Chronica Botanica. Wal- 
tham, Mass. 1945. [Azlanthus en 115, 116.] 


Subfam. PICRAMNIOIDEAE Engler 


4. Picramnia Swartz, Prodr. Veg. Ind. Occ. 2, 27. 1788, nom. cons. 


Trees or shrubs, usually with slender, curving branches, the bark and 
wood often very bitter. Leaves odd-pinnate, persistent; leaflets [3]5—9[21], 
opposite or alternate, entire, chartaceous to + coriaceous, petioluled. 
Plants dioecious [or occasionally polygamous]. Flowers minute, unisexual 
[occasionally also bisexual]. Inflorescences paniculate, terminal [some- 
times opposite the leaves], with slender, raceme- or spikelike branches. 
Sepals 5 [3 or 4], connate from %4 to % of their length, imbricate. Petals 
5 [3 or 4, or wanting|, narrowly linear-lanceolate [or lanceolate], as long 
[or twice as long] as the sepals, imbricate. Stamens as many as and oppo- 
site the petals, inserted below and between the lobes of a low intrastaminal 
disc, reduced to staminodia in @ flowers; filaments subulate; anthers basi- 
fixed, almost globular, with thick connective, introrse, 4-locular at anthesis. 
Gynoecium 2(3)-carpellate, syncarpous, rudimentary in 6 flowers; stigma 
deeply 2-lobed, the lobes thick, divergent; style very short and stout, usu- 
ally inconspicuous; ovary sessile, 2(3)-locular; ovules anatropous, epitro- 
pous, 2-integumented, pendulous from near the top of the carpels, 2 in 


182 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


each locule. Fruit a rather juicy, subglobular, ellipsoidal or obovoid berry, 
(3)2- or (by abortion) 1-locular, with 1-seeded locules. Seed ovoid, often 
plano-convex, filling the cavity of the locule, the testa membranaceous, ad- 
nate to the undifferentiated embryo; endosperm wanting. (Pseudobrasilium 
Adans., 1763; Tariri Aubl., 1775; nom. rejic.) Type species: P. antidesma 
Swartz. (Name derived from Greek, pikros, bitter, and thamnos, shrub, 
in allusion to the bitterness of the vegetative parts.) — BITTERBUSH. 


A genus of about 40 species of tropical American distribution. Picramnia 
pentandra Sw., a shrub or small, slender tree, of the West Indies, Colombia, 
and Venezuela, occurs in our area in the coastal hammocks of southeastern 
peninsular Florida (Dade County) and on the Florida Keys. Berries of 
this species are olive-shaped, 9-15 mm. long, red, turning black when 
fully ripe; the seeds are light brown and lustrous. Leaves, bark, and roots 
of this and other species have been used locally in the tropics as febrifuges. 

The genus, which represents a subfamily of its own, is in need of a re- 
vision. 


REFERENCES: 
Under the family references see ENGLER (1931, pp. 402, 403), Recorp & Hess, 
SARGENT, SMALL, and West & ARNOLD (p. 101). 
RaADLKOFER, L. Ueber die Gliederung der Familie der Sapindaceen. Sitz-ber. 
Akad. Wiss. Miinchen IT. 20: 105-370. 1890. | Picramnia, relationship 
with Alvaradoa, 139-143. | 


Subfam. ALVARADOIDEAE Engler 
5. Alvaradoa Liebmann, Vid. Medd. Nat. For. Kjgbenh. 1853: 100. 


Shrubs or trees up to 15 m. high, with slender, terete, pubescent | or 
glabrous| branchlets and bitter bark. Leaves odd-pinnate, many-foliolate, 
crowded at the end of branches, apparently persistent; leaflets alternate, 
small (not over 5 cm. long), thin and firm [or leathery], entire, petiolu- 
late. Plants dioecious. Flowers minute, unisexual, in slender, many- 
flowered, axillary or terminal racemes. Sepals 5, usually distinct in @ 
flowers and variously connate from near the base to the half of their 
length in 4 flowers, valvate. Petals 5, linear-filamentous, present [or 
absent| in ¢ flowers, wanting in @ flowers. Stamens 5, alternate with 
the sepals, inserted below and between the lobes of the disc, wanting in 9 
flowers; filaments filiform, hairy in the lower part; anthers basifixed, ob- 
long in outline, with a conspicuous, almost orbicular, swollen connective 
and introrse anther-halves, 4-locular at anthesis; pollen grains small, 
prolate- subspherical, 3-colpate. Intrastaminal disc thickish, deeply 5-lobed 
in é flowers, thin and scarcely lobed in @ flowers. Gynoecium 3-carpellate 
(but only 1 carpel fertile), syncarpous, sessile on the disc, wanting in 
é flowers; stigmas small, simple; styles 3, distinct, sibulate, short, re- 
curved; ovary densely villous, flattened, obtusely triangular in cross sec- 


1962 | BRIZICKY, SIMAROUBACEAE AND BURSERACEAE 183 


tion, imperfectly 2- or 3-locular on account of 2 incomplete partitions 
(apparently deeply intruded parietal placentae) demarcating a sole fertile 
locule in the obtuse angle of the ovary; ovules 2 in the fertile locule, basal, 
ascendent, anatropous, apotropous, 2-integumented. Fruit a compressed, 
2|3]-winged, + papery samaroid capsule [or leathery samara], crowned 
by remnants of the styles, appearing 3- or 1-locular, with 1 seed in the 
lower half. Seeds + terete, narrowly ellipsoidal (the shape of rice grains) 
[or rather compressed and broadly elliptical in outline]; testa mem- 
branaceous; endosperm wanting; embryo straight, with plano-convex 
[or flat], fleshy cotyledons and a short, inferior radicle. Typr SPECIES: 
A. amorphoides Liebm. (Named in commemoration of Pedro de Alvarado, 
one of the chief aides of Hernando Cortez in the conquest of Mexico.) 


A genus of about five species, the range disjunct, including southern 
Florida, the West Indies, Mexico, and Central America, Bolivia, and 
Argentina. Alvaradoa amorphoides occurs in a few hammocks in southern 
peninsular Florida (Dade County) and the Florida Keys and in the 
West Indies, Mexico, and Central America. 

Although the genus was originally placed with the Sapindaceae, Radl- 
kofer showed Alvaradoa to be simaroubaceous with the closest relation- 
ship to Picramnia. Engler, however, placed the two in separate unigeneric 
subfamilies. 


REFERENCES: 

See also under family references, Cronqutst (pp. 132-137), ENGLER (1931, 
p. 404), Recorp & Hess (p. 510), SARGENT (pp. 644, 645), SMALL, and WEsT 
& ARNOLD (p. 104). 

RApDLKOFER, L. Ueber die Gliederung der Familie der Sapindaceen. Sitz-ber. 
Akad. Wiss. Munchen II. 20: 105-370. 1890. [Alvaradoa, morphology of 
the flowers and fruits, relationship with Picramnia, 139-143.]| 


BURSERACEAE Kunth, Ann. Sci. Nat. 2: 346. 1824. 
(TorcHwoop FaMILy) 


Trees or shrubs, the inner bark with resin ducts. Leaves alternate, 
usually once pinnate, deciduous [or persistent], usually exstipulate. 
Flowers small, hypogynous, regular, apopetalous, usually unisexual by 
abortion, 3—5-merous, in axillary |or terminal} cymose panicles. Plants 
mostly dioecious. Stamens 6—10, usually distinct, inserted below an intra- 
staminal [rarely extrastaminal] nectariferous disc; anthers versatile, in- 
trorse, longitudinally dehiscent, sterile in @ flowers. Gynoecium 3[2-5]- 
carpellate,. syncarpous, rudimentary or wanting in 4 flowers; ovary 
3|2—5]|-locular, with axile placentae; ovules anatropous, epitropous, 2 in 
each locule. Fruit usually drupaceous, with + dry [or fleshy] exo- and 


without endosperm, solitary in each locule; embryo straight [or curved], 


184 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLIII 


cotyledons contortuplicate [or flat], aes + deeply lobed, radicle 
superior. TyPrE GENUS: Bursera Jacq. ex 


A pantropical family of 15 or 16 genera and about 600 species. Six 
genera (including Bursera and Protium) with about 200 species occur 
in the New World. The family is subdivided into the tribes Protieae 
Engl., Bursereae [H. J. Lam] (Boswellieae Engl.), and Canarieae Engl. 
The subdivisions, although based exclusively upon the structure of the 
fruits, seem to be rather natural ones. 

Burseraceae are closely allied with Rutaceae, Simaroubaceae, and Meli- 
aceae. A close relationship to Anacardiaceae, proposed by some taxono- 
mists (e.g., Radlkofer), is supported by evidence from anatomy and 
palynology. 

Resinous substances obtained from some species are of economic sig- 
nificance. The most important of these are myrrh, from Commiphora 
molmol Engl. and C. abyssinica (Berg) Engl., and olibanum or frankin- 
cense, from species of Boswellia, especially B. Carteri Birdw. Myrrh is 
used in perfumery, and both are used in medicine and as incense. 


REFERENCES: 
See also RADLKOFER under Alvaradoa (Simaroubaceae). 


ENGLER, A. Burseraceae et Anacardiaceae. 71: CANDOLLE, A. & C. pe. Monogr. 
Phaner. 4: 1-573. pls. 1-15. 1883. [Burseraceae, 1-169. pls. 1-3.] 

Burseraceae. Nat. Pflanzenfam. ed. 2. 19a: 405-456. 1931. 

GUILLAUMIN, A. Répartition géographique et biologie des Burséracées. Revue 
Gén. Bot. 20: 321-327. pls. 11-14. 1908 

. Les produits utiles des Burséracées. 73 pp. Paris. 1910. 

. Recherches sur la structure et le développement des Burséracées. Ann. 
Sci. Nat. Bot. IX. 10: 1-302. 1909. 

Heimscu, C., Jr. Comparative anatomy of the secondary xylem in the ‘“Gru- 
inales”’ and “Terebinthales’, of Wettstein with reference to taxonomic 
grouping. Lilloa 8: 83-198. pls. 1-17. 1942. |[Burseraceae, 122-124.] 

Japtn, F. Recherches sur la ees et les affinités des Térébinthacées. Ann. 
Sci. Nat. Bot. VII. 19: 1-51. 1894. |Terebinthaceae divided into tribes 
Anacardieae and Bursereae. | 

Lam, H. J. Beitrage zur Morphologie der Burseraceae insbesondere der Cana- 
rieae. Ann. Jard. Bot. Buitenzorg 42: 97-226. pls. 9-16. 1932. | Pinnate 
leaves of Burseraceae and related families are considered phylogenetically 
reduced shoots; but cf. Stnta, 1938. 

. The Burseraceae of the Malay Archipelago and peninsula, with anno- 
tations concerning extra-Malayan species, especially of Dacryodes, San- 
tiria, and Canarium. Pace Jard. Bot. oo IIT. 12: 281-561. 1932. 
[The “General Part,’ 281-317, contains many important data on mor- 

ology, dispersal, andl phylogeny of vm family. 

ee P. W., C. Katkman, & H. J. Lam. Burseraceae. Jn: C. G. G. J. 

N STEENIS, Fl, Males. ds 5: 209-296. 1955. [Includes notes of general 
nepaien on ecology, dispersal, distribution, wood anatomy, morphology, 
and taxonomy. | 


1962 | BRIZICKY, SIMAROUBACEAE AND BURSERACEAE 185 


MarcuHanp, L. Recherches sur l’organisation des Burséracées. 56 pp., ls. 
1-6. Paris. 1868. 
pane L. L. Microsporogenesis and female gametophyte in Boswellia ser- 
a Roxb. Curr. Sci. Bangalore 28: 77, 78. 1958.* 
———. Studies in Burseraceae. I. Jour. Indian Bot. Soc. 39: 204— -209. 1960. 
[ Boswellia serrata and Garuga pinnata Roxb., floral anatom 
s in Burseraceae. II. Jbid. 402 ~409. 1960. [Bursera serrata 
Colcne (= Protium serratum (Wall. ex Colebr.) Engl.), floral anatomy 
and Ee eee Garuga pinnata, embryology. 
REcorp, S. J., . W. Hess. Timbers of the New World. 640 pp., pls. 1-59. 
New oe ee { Burseraceae, 105-110. ] 
Rose, J. N. Burseraceae. N. Am. Fl. 25: 241-261. 1911. 
oe C. S. Manual of the trees of North America (exclusive of Mexico). 
ed. 2. 897 pp. 1922. [Burseraceae, 645-648. | 
SHUKLA, - D. Studies in the family Burseraceae—I. Floral anatomy of 
Balsamodendron mukul Hook. Agra Univ. Jour. Res. Sci. 4: 567-573. 
L9552% 
. Gametophyte in Balsamodendron mukul. Curr. Sci. Bangalore 23: 333. 
1954.* 


SintA, H. R. Zur Phylogenie der Fiederblatter der Burseraceen und verwandter 
Familien. Ann. Jard. Bot. Buitenzorg 48: 69-100. pls. 13, 14. 1938 
[Against Lam’s theory of the caulomic nature of pinnate leaves of Bur- 
seraceae ae their allies. | 

WeBBER, I. E. Systematic anatomy of the woods of the “Burseraceae.” Lilloa 
6: 441-465. pls. 1-4. 1941. 

WIGER, J. Embryological studies on the families Buxaceae, Meliaceae, Sima- 
rubaceae and Burseraceae. Thesis. Lund. 1935.* 


1. Bursera Jacquin ex Linnaeus, Sp. Pl. ed. 2. 1: 471. 1762; Gen. Pl. 
ed. 6. 440. 1764, nom. cons. 


Trees [or shrubs]. Leaves odd-pinnate [sometimes bipinnate], 3—9[- 
many |-foliolate [rarely 1-foliolate], usually crowded at the end of branch- 
lets, deciduous; leaflets opposite, chartaceous to subcoriaceous [or cori- 
aceous|, entire [or toothed], manifestly petiolulate [to sessile]. Flowers 
very small, unisexual [and/or bisexual], in axillary, raceme-like panicles, 
appearing prior to or with [or after] the leaves. Plants dioecious [or 
polygamous? J. Sepals 3-5, minute, connate at least at base, imbricate 
in bud. Petals 3-5, whitish to creamy, much longer than the sepals, 
spreading and recurved, induplicate-valvate in bud. Stamens 6-10, non- 
functional in @ flowers; filaments subulate; anthers oblong in outline, 
dorsifixed near the base, shorter and without pollen in @ flowers; pollen 
grains medium-sized, 3-colpate, reticulate-striate. Intrastaminal disc an- 
nular, 6-10-lobed, orange or red. Stigma capitate, 3-lobed; style short, 
ovary sessile, ovoid, 3-carpellate and -locular, with 2 collateral. pendu- 
lous ovules in each locule, rudimentary in 4 Aigwers. Drupes eubelobular 
or obliquely ellipsoid, renee triangular, with resinous, fleshy, leathery 
exo- and mesocarp detaching in 3 [or 2] valves when the fruit matures; 
stones (bony endocarp) covered with a thin, membranaceous, light-pink 
coat (probably the innermost layer of mesocarp remaining attached to the 


186 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLIII 


endocarp), usually solitary, attached to the persistent axis of the ovary, 
ovoid-trihedral, essentially 1-locular, but bearing 2 minute, sterile locules 
or rudimentary stones in the upper half on the adaxial (ventral) side, 
usually 1-seeded. Seeds without endosperm; testa membranaceous; em- 
brvo straight Jor curved] with foliaceous, contortuplicate cotyledons and 
a short superior radicle. Germination epigeous, cotyledons 3-fid, the first 
leaf 3-foliolate. (Elaphrium Jacq., 1760; Simaruba Boehmer, 1760; nom. 
rejic.) Typr species: B. gummifera L. (= B. Simaruba (L.) Sarg.). 
(Named in honor of Joachim Burser, 1583-1639, a German physician and 
botanist. ) 


A genus of about 100 species of tropical America. Bursera Simaruba 
(Elaphrium Simaruba (L.) Rose), gumbo-limbo or West Indian birch, 
of the West Indies, Mexico, Central America, and northern South America, 
occurs in coastal hammocks on the Florida Keys and in peninsular Florida 
about as far north as Brevard and Pinellas counties. Easily recognizable 
by its lustrous, smooth, copper-colored bark which exfoliates in thin, 
papery lavers as in some species of Betula, in winter the tree (to 20 m. ) 
is conspicuously leafless, in contrast with its evergreen associates. The 
species is a common “fence-post” tree in tropical America, for pieces of 
the trunk or branches set in the ground quickly develop roots and grow 
into trees. The Mexican B. fagaroides (HBK.) Engl. and B. microphylla 
Gray occur in southern Arizona, the latter in southeastern California as 
we 

Insect pollination has been presumed for the genus, but no data are 
available. Bursera Simaruba apparently is dioecious; records of polygamy 
are in need of verification. Staminate flowers in this species are 5-merous 
(or more rarely 4-merous). while the carpellate appear to be almost in- 
variably 3-merous and only very rarely 4-merous. 

The species of Bursera yield a fragrant glutinous resin which is locally 
applied in domestic medicine. The resin of B. Simaruba is also used as a 
substitute for glue and as cement for mending broken china and glass. 

The genus seems to be most closely related to the paleotropic Boswellia 
Roxb. ex Colebr 


— 


i 


REFERENCES: 
See also under family references. ENGLER (1931, pp. 423-429). Recorp & 
Hess. Rose, and SARGENT. 
Buttock. A. A. Contributions to the flora of tropical America: XNNVII. Notes 
the Mexican species of the genus Bursera. Kew Bull. 1936: 346-387 
ee {Includes kev. economic notes. and references to economic uses and 
anatomy. See also Kew Bull. 1937: 447-457. 1937, and 1938: 163-168 
1938. for further notes. including the identification of Hinton’s Mexican 
collections. | 
SarGENT, C. S. Bursera. Silva N. Am. 1: 95-98. pls. 41, 42. 1891. 
West, 2, 1. 2. ARNOLD. The native trees of Florida, 212 pp. Gainesville. 
1946. [B. Simaruba, 103.] 


1962 | BAILEY & SRIVASTAVA, CACTACEAE, IV 187 


COMPARATIVE ANATOMY OF THE LEAF-BEARING 
CACTACEAE, IV 
THE FUSIFORM INITIALS OF THE CAMBIUM 
AND THE FORM AND STRUCTURE OF THEIR DERIVATIVES 


— 


I. W. BartLey ! AND LAtitT M. SRIVASTAVA ” 


Ir HAS BEEN DEMONSTRATED that in both dicotyledons and mono- 
cotyledons vessels originated by modification of long tracheids having 
scalariform pitting (Bailey & Tupper, 1918; Cheadle, 1942, 1943). In the 
case of the dicotyledons, highly advanced stages in the phylogenetic special- 
ization of vessels occur in plants having short vessel members with simple 
perforation plates throughout both the primary and secondary xylem 
(Bailey, 1944). In such plants scalariform perforation plates are elimi- 
nated. During the evolutionary specialization of vessels there commonly 
tend to be concomitant changes in the ground mass of imperforate 
tracheary cells which, by elimination of the borders of their pits, become 
libriform fibers, which, in turn, may at times retain their living contents, 
become septate, and function in the storage of starch (Bailey, 1936). 

In dicotyledons, the differentiation of sieve tubes in the secondary 
phloem has appeared to afford a phylogenetic parallel to the development 
of vessels in the secondary xylem (Esau, 1953, p. 275; Esau, Cheadle & 
Gifford, 1953). In both tissues, axial translocation seems to be facilita- 
ted by modification in the more or less extensively overlapping ends of 
adjacent cells in axial seriations: in the xylem by loss of pit membranes 
to form perforations and in the phloem by formation of sieve plates 
having larger connecting strands than those in the sieve areas of the 
lateral walls. Some investigators have suggested that there is a direct 
correlation between the degree of evolutionary spcialization of sieve 
plates in the end walls and decrease in conspicuousness of the sieve areas 
in the lateral walls (Cheadle & Whitford, 1941; Cheadle, 1948; Cheadle 
& Uhl, 1948; Esau & Cheadle, 1959). Zahur (1959), however, did not find 
such a correlation. 

It should be strongly emphasized in this connection that huge volumes 
of anatomical data regarding the secondary xylem of dicotyledonous 
families have accumulated during the last half-century. Wood, in general, 
is adequately preserved by simple drying, and industrial pressures have 
stimulated the assembling and study of large collections of wood samples 
by institutions in various parts of the world. No comparable information 
is available at present regarding the secondary phloem of dicotyledons. 

' This investigation was financed in part by a grant from the National Science 
Foundation. I am indebted to the pee Philosophical Society for the loan of a 
Wild microscope. 

? Mercer Fellow of the Arnold Arboretum. 


188 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


Further, as explained by Cheadle (1956), this tissue is difficult to study 
because of its physiological and structural peculiarities. Even the most 
extensive reconnaissance to date (Zahur, 1959) deals with only a rela- 
tively few representatives of the 85 families sampled. 

Furthermore, as one of us (Bailey, 1957) has emphasized: ‘There 
are certain details of the trends of specialization in the xylem which need 
to be more clearly and generally understood in shifting from a consideration 
of the dicotyledons as-a-whole to investigations of the taxonomy of 
individual taxa of restricted size. In revealing salient trends of evolu- 
tionary specialization by analyses of data obtained from the dicotyledons 
as-a-whole, variations due to obtaining specimens from different parts 
of the plant, from plants of different growth rates, from genetically 
different taxa, etc., tend to neutralize one another. In addition, various 
localized, divergent trends of specialization do not obscure or confuse the 
major trends of evolution in the dicotyledons as-a-whole. However, when 
one becomes concerned with taxa of decreasing size, viz. families, sub- 
families, tribes, genera and species, such variations and deviations become 
increasingly significant.” 

For example, data from the dicotyledons as-a-whole clearly reveal 
evolutionary changes from scalariform to simple perforation plates, and 
from scalariform to alternate multiseriate intervascular pitting. In any 
randomly selected minor taxon, one of these trends may be retarded or 
accelerated in relation to the other. Therefore, with present inadequate 
information regarding the secondary phloem in most orders, families, 
and genera, it is not possible to determine with certainty what some of 
the more important trends of phylogenetic specialization in the dicotyledons 
as-a-whole may actually be. This is particularly the case in those paren- 
chymatous cells that are physiologically and ontogenetically related to 
sieve elements. 

During the phylogenetic specialization in the secondary tissue of dicoty- 
ledons, there is a progressive shortening of fusiform cambial initials 
which not infrequently culminates in storied forms of cambia, involving 
longitudinal rather than pseudotransverse anticlinal divisions and the 
elimination of intrusive elongation following such divisions (Bailey, 1923). 

In general, the phylogenetic shortening of fusiform cambial initials is 
most closely paralleled in the secondary xylem and phloem by shortening 
of fusiform parenchymatous derivatives and by changes in the length 
of parenchyma strands (for ordinarily fusiform parenchymatous cells and 
the mother cells of parenchyma strands do not elongate during tissue 
differentiation). As a concomitant of shortening fusiform cambial initials, 
the parenchyma strands commonly tend to be composed of fewer and 
generally of shorter cells. 

Statistical data obtained from the dicotyledons as-a-whole indicate that 
shortening of the fusiform initials likewise is closely reflected in the length 
of vessel members, there being only slight elongation at times during the 
maturation of primitive, long, slender vessel members, and a slight con- 
traction at times during the differentiation of short, very broad, highly 


1962 | BAILEY & SRIVASTAVA, CACTACEAE, IV 189 


-specialized ones. On the contrary, fiber-tracheids and libriform fibers 
become longer than the fusiform cambial initials from which they are 
derived owing to more or less extensive intrusive elongation during tissue 
differentiation. 

In the secondary phloem of such vesselless gymnosperms as the Pinaceae, 
Taxodiaceae, and Cupressaceae, statistical averages indicate that sieve 
cells simulate fusiform cambial initials in length, there being no appre- 
ciable elongation, as occurs at times in tracheids, during their maturation 
from cambial initials. In the phylogenetically specialized dicotyledons 
having very short fusiform cambial initials, particularly those having 
storied cambia, the sieve-tube members in statistical averages closely 
simulate the fusiform initials in length. On the contrary. in some dicoty- 
ledons having less highly specialized cambia, the parallelism in length may 
be modified by more or less numerous divisions of mother cells prior to 
the formation of sieve-tube members, companion cells, and some paren- 
eat cells (cf. Esau & Cheadle, 1955; Cheadle & Esau, 1958; Zahur, 

1959). In such plants, the sieve-tube members tend statistically to be 
considerably shorter than fusiform cambial initials. 

Statements in the literature regarding detailed structure of the xylem 
and phloem of leaf-bearing cacti are casual and fragmentary and are 
based largely upon Pereskia aculeata Mill. The occurrence of septate 
libriform fibers and porous vessels in the wood of this species is recorded 
by Schenck (1893) and Solereder (1899). If the most primitive living 
cacti occur among the leaf-bearing ones, as is generally assumed to be 
the case, it is essential to obtain comprehensive information regarding the 
levels of phylogenetic specialization that they have attained, particularly 
for future use in understanding trends of increasing structural specializa- 
tion that occur in the Opuntieae and Cereeae. 

In all of the putative species of Pereskia, Pereskiopsis, and Quiabentia 
that we have studied (mentioned in previous papers of this series | Bailey. 
1960, 1961a, 1961b]) the fusiform initials of the cambium have attained 
a high level of evolutionary modification. ‘They are palaaaaiy short, 
commonly ranging in length from only 150 to 400 microns. Althou Bh 
they are not consistently in perfect storied or stratified eee’ they 
frequently exhibit a tendency to become storied, at least in some parts 
of a mature plant. Where they approach a perfect storied arrangement, 
they have abruptly tapered ends and a hexagonal form as seen in tangential 
longitudinal sections of the cambium (Fic. 1) and the cells of one stratum 
do not extensively overlap those of higher and lower levels. On the con- 
trary, where stratification is imperfect, the cells have more gradually 
tapered ends, and there is more overlapping of the cells of different levels 
(Fic. 4). It should be noted in this connection that where stratification 
occurs in the leaf-bearing cacti it tends to differ from that which occurs 
in plants of other dicotyledonous families in exhibiting a conspicuous 
tendency for the strata of fusiform initials to have a diagonal, rather 
than a transverse, orientation as seen in tangential longitudinal sections 
of the cambium. 


190 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLII 


In dicotyledons having very short, especially storied, cambial initials, 
where adequately preserved material of the cambium is not available, 
the length and the arrangement of fusiform initials mav be detected both 
in certain of their derivatives in the xylem and in recently formed 
phloem, 

The distribution of wood parenchyma in Pereskia, Pereskiopsis, and 
Quiabentia is of a specialized paratracheal rather than a_ primitive 
apotracheal form, but more or less extensive arcs of thick-walled lignified 
or thin-walled unlignified parenchyma occur at times and may prove to 
be associated in some manner with successive zones of growth in the 
enlargement of stems and roots. The fusiform parenchymatous cells 
and parenchyma strands are short, the latter being composed usually of 
two or three cells only. In tangential longitudinal sections of the second- 
ary xylem, the fusiform parenchyma and parenchyma strands (where not 
deformed and displaced by excessive enlargement of vessels) closely simu- 
late fusiform cambial initials in overall form. Furthermore, where paren- 
chyma is sufficiently abundant, particularly in areas of zonal distribution. 
the arrangement of fusiform cambial initials, whether storied or non- 
storied, is clearly revealed by the arrangement of the parenchymatous 
cells, 

The vessels in stems and roots of the leaf-bearing cacti are of a highly 
advanced evolutionary form having simple perforation plates throughout 
the primary (Fic. 8) and secondary xylem (Fic. 5). In the secondary 
xylem, the vessels vary markedly in diameter from 20 microns to as much 
as 200 microns in certain cases. The vessels occur independently and in 
clusters of varying size and form. The smaller vessels, as seen in trans- 
verse sections, are angular, whereas the larger ones are more nearly cir- 
cular or oval, except where they are modified by compression in clusters. 
Such variations in the vessels occur, not only in different species and dif- 
ferent parts of a single plant, but also in closely oe areas of the sec- 
ondary xylem. The members of the smallest vessels (i.e., those which do 
not expand appreciably in tangential diameter during maturation) re- 
semble fusiform cambial initials in overall form when viewed in tangential 
longitudinal sections of the xylem. In such sections, the vessel members 
may be confused with vascular tracheids, but the perforations in their more 
or less abruptly tapered ends are clearly visible in radial longitudinal sec- 
tions. Where sufficiently abundant, and particularly in association with 
parenchyma, such small vessel members reveal a storied (Fic. 2) or non- 
storied arrangement of fusiform initials in the cambium. 

In the secondary xylem of the leaf-bearing cacti, vessel members of the 
smallest diameter tend to have diagonally oriented perforation plates 
in their terminal radial walls. Larger vessel members, although as short 
or shorter than fusiform cambial initials, vary more or less markedly in 
form owing to excessive lateral enlargement during their maturation. The 
perforation plates, consisting of a circular or oval opening surrounded by 
a clear zone of unpitted wall, may be transversely (Fic. 5) or more or 
less diagonally oriented. Furthermore, where vessels are closely associated 


1962 | BAILEY & SRIVASTAVA, CACTACEAE, IV 191 


in larger clusters and where vessels deviate from a longitudinal course to 
pass diagonally through the broad multiseriate rays (Fics. 6, 7), the vessel 
members may at times have perforation plates in their sides rather than in 
their more or less inclined ends. 

The bordered pitting in the sides of adjoining vessels tends to be of 
the evolutionarily advanced alternate-multiseriate form (Fics. 9, 11), but 
areas of scalariform pitting (Fics. 12, 13) and of opposite-multiseriate pit- 
ting are of not infrequent occurrence. The bordered pits of multiseriate ar- 
rangements vary in size and considerably in form from circular te oval and 
to angular where the pits are closely crowded. (Compare Fics. 9, 11, 13, 
15, 16.) The pit apertures likewise vary from small, nearly circular (Fic. 
9), to oval (Fic. 15), and to transversely slitlike (Frc. 151. Furthermore, 
the borders of the pits exhibit at times reduction in conspicuousness in rela- 
tion to the size of the apertures. (Compare Fics. 11, 15, 16.) In areas of 
scalariform pitting, unconformity in the pitting of the walls of adjacent 
vessels frequently occurs (Fics. 12, 13), i.e.. a transversely extensive pit 
in one wall being related to smaller pits in the adjoining vessel wall. 

The pits of vessels in contact with wood parenchyma and rays exhibit a 
wide range of variability. At times, they closely resemble those of inter- 
vessel contacts in size, form, and arrangement. (Compare Fics. 9, 10; 13, 
14; 16, 18.) However, in the case of the leaf-bearing cacti, there commonly 
is a conspicuous tendency for vessel pits in contact with parenchyma to 
have enlarged apertures and to reduce or eliminate their bordering area 
(Fic. 20). In other words, the pits in the walls of vessels in contact with 
parenchyma become simple pits in conformity with the unbordered pits 
of adjacent parenchymatous cells. Where vessels are completely jacketed 
by wood parenchyma and have large, transversely elongated unbordered 
pits, the vessel members, when isolated by maceration, resemble vessel 
members of those parts of the primary xylem which have reticulate forms 
of wall thickening (Fic. 22). 

The libriform fibers of the leaf-bearing cacti, unlike the vessel mem- 
bers, fusiform wood parenchyma, and parerichyma strands, elongate more 
or less extensively by intrusive elongation during tissue differentiation. 
Therefore, they (Fics. 24-27) become longer than the fusiform cambial 
initials (Fic. 23) from which they are derived. However, the amount of 
elongation varies greatly, not only in different parts of a plant, but also in 
closely adjacent parts of the secondary xylem. In some cases, the elonga- 
tion at one or both ends of the cell is only a few microns, whereas, in other 
cases, the elongation at both ends of the cell is extensive. Thus, libriform 
fibers may be only slightly longer than the fusiform cambial initials from 
which they are derived, i.e., 150-400 microns (Fic. 24) or they may attain 
maximum lengths of as much as 1000-1300 microns (Fics. 26, 27). Some 
libriform fibers are relatively slender and taper gradually and uniformly 
toward their ends (Fic. 26); others are broader in their central part and 
taper abruptly at one or both ends into long narrow tips (Fics. 25, 27). 
In both instances the small oval or slitlike simple pits tend to be concen- 
trated in the central part of the fibers which corresponds roughly in length 


192 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


to that of the fusiform cambial initials. (Compare Fic. 23 with Fics. 24— 
27.) 


The slitlike pits usually are oriented more or less diagonally to the long 
axis of the cell but may at times be oriented parallel to its axis. This in- 
dicates that the orientation of cellulosic microfibrils in the central or Se 
layer of the secondary wall may, at least occasionally, deviate from helical 
to Jongitudinal. 

he fibers in any particular part of the secondary xylem may be pre- 
vailingly septate (usually a single septum to each cell, Fic. 26), nonseptate, 
or in varying mixtures of septate and nonseptate. Both the septate and the 
nonseptate (Fics. 17, 19, 21) fibers retain their protoplasts and nuclei and 
are capable, at least in certain parts of the plant or at certain seasons of the 
year, of storing abundant starch. 

In the members of Pereskia, Pereskiopsis, and Ouiabentia which we in- 
vestigated, the phloem derivatives, viz., sieve elements and parenchyma, as 
seen in tangential longitudinal sections in close proximity to the cambium, 
commonly reflect the form and distribution of the fusiform cambial initials 
(Fics. 3,4). The fusiform parenchyma and parenchyma strands, however, 
are more reliable indicators of the overall form of fusiform initials than are 
the sieve elements. This is because the sieve-tube members commonly dif- 
ferentiate in one or more parts of a cell complex that results after longitudi- 
nal and sometimes transverse divisions in a single phloem mother cell. 
Therefore, although the length of sieve-tube members in statistical averages 
generally corresponds to that of the fusiform cambial initials, their overall 
form frequently does not. The phloem-parenchyma strands are usually 
composed of two or three cells arranged in a vertical file and appear to 
originate by divisions in a single phloem mother cell. 

The sieve-tube members present features of advanced structural speciali- 
zation. The sieve plates commonly occur in the more or less transverse 
(Fic. 28) or slightly oblique end walls (Frcs. 29, 30) but sometimes 
may be present in the lateral walls (Fic. 29). In certain cases, sieve plates 
may not be present at both ends of a sieve-tube element but may occur only 
at one end and in one of the lateral walls (Fic. 29). Usually the pores and 
the callose cylinders are distributed rather evenly throughout the sieve 
plate and the latter may be interpreted as a simple sieve plate. Occasional 
variations in sieve-plate structure occur, however. In some sieve elements 
with somewhat sloping end walls the pores and the callose cylinders may 
be distributed in two or more distinct groups in the sieve plate (Fics. 30, 
31). There is no strict uniformity with regard to the distribution of hese 
two forms of sieve plates. They occur in different parts of the same plant 
and even at the two ends of a single sieve-tube member. These variations 
in sieve-plate structure are related in a general way to the orientation of 
the end walls in the sieve elements which, in turn, seems to be related to 
the character and orientation of end walls in the fusiform cambial initials 
(Fie, 23) 

In addition to sieve plates which usually occur at or near their end walls, 
sieve-tube members have numerous small sieve areas that are scattered 


1962 | BAILEY & SRIVASTAVA, CACTACEAE, IV 193 


throughout both radial and tangential lateral walls (Fic. 32). The pores 
and their callose cylinders in these sieve areas are much smaller than those 
in the sieve plates. (Compare Fics. 31, 32.) The size of these lateral sieve 
areas, as determined by the number of callose cylinders stained with lac- 
moid in one area, varies considerably. In the same sieve-tube member, 
some lateral sieve areas may be very small, composed of one or two strands 
and their callose cylinders only, whereas others may be composed of sev- 
eral strands and their callose cylinders. 

Generally, one or more companion cells occur in association with a 
sieve-tube member (Fics. 28, 29). In addition, one or more members of 
a cell complex, in which sieve-tube elements and companion cells arise, 
may differentiate as parenchymatous elements. The exact relationships 
between the different cells that originate after divisions in a single phloic 
mother cell and the sieve-area connections between them require detailed 
ontogenetic study. 

During the transition from “functional” to “nonfunctional” phloem * 
the parenchymatous cells of both the axially oriented part and the rays 
retain their living contents, capacity for division, and commonly undergo 
more or less extensive changes in size and form. The most striking changes 
of taxonomic significance are those which occur in the nonfunctional phloem 
of Pereskia, these being absent in Pereskiopsis and Quiabentia. As one of 
us has shown (Bailey, 1961a) three distinct categories of pereskias can be 
segregated upon the basis of differences in the formation of sclereids in 
the nonfunctional part of the secondary phloem. In two of these cate- 
gories of species, sclereid formation involves extraordinary enlargements 
of parenchymatous cells in both diameter and length. In the third cate- 
gory, the changes in size and form are more nearly comparable to those 
that occur in the formation of ordinary sclereids 

Although crystals are commonly present in the ray parenchyma of 
functional phloem in all three genera, they rarely, if ever, occur in axially 
oriented parenchyma. However, they do occur at times in axial parenchyma 
of nonfunctional phloem (Bailey, 1961b). Variations in the presence and 
form of these crystals may prove to be of some taxonomic significance 
when more adequate and extensive collections are available for detailed 
investigation. 


DISCUSSION 


It is evident from our reconnaissance of the secondary vascular tissues of 
Pereskia, Pereskiopsis, and Quiabentia that these tissues have attained 
highly advanced levels of evolutionary specialization. This is shown, for 
example, by the dimensions and form of the cambial fusiform initials and 
their derivatives, by the perforation plates and the lateral pitting in ves- 
sels, by the storage of starch in both septate and nonseptate libriform 
fibers, and by the distribution patterns of vessels and wood parenchyma. 

° Following Esau (1953, p. 299) “nonfunctional” phloem refers to that part of the 
phloem in which sieve elements and companion cells have ceased to function 


194 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLIII 


Such evidence at least suggests that the early leaf-bearing representatives 
of the Cactaceae (those of nearly typical dicotyledonous woody arborescent 
and shrubby form) had attained high levels of internal anatomical speciali- 
zation before increasing succulence and other morphological changes led 
to the differentiation of the Opuntieae and Cereeae 

The ranges of anatomical variability in different parts of a single plant 
and in the same clone when grown under different environmental in- 
fluences are very extensive. Potential diagnostic criteria, such as differ- 
ences in the size, form, and distribution of the constituent cells of xylem, 
in the form and orientation of perforation plates; in pitting between vessels 
and between vessels and parenchyma; in the presence or absence of septa 
in libriform fibers, etc. (which have been utilized so commonly in the 
differentiation of taxa in other dicotyledonous families) appear, in gen- 
eral, in the leaf-bearing Cactaceae to be of a quantitative rather than a 
qualitative character. Likewise, in phloem, the orientation of the sieve 
plates near the ends of sieve-tube members may vary from more or less 
oblique to perfectly transverse, and the number of pore groups (viz., 
sieve areas) on the sieve plate may range from one to three or more. 
The difference between the size of pores and connecting strands in the 
lateral sieve areas versus the sieve plates is very great in all species of the 
three leaf-bearing genera. Therefore, in the case of these cacti, large 
volumes of material must be studied statistically in order to attain results 
of valid taxonomic significance. 

In the past, unjustifiable phylogenetic and taxonomic inferences have 
resulted from comparisons between the frequency of diagonal and trans- 
verse orientations of terminal perforation plates in closely related taxa 
without taking into consideration various factors involved in the statis- 
tical differences. Important influences in this connection are the length 
of fusiform cambial initials, the amount of transverse enlargement of ves- 
sel members during tissue differentiation, the deformation when vessels 
occur in crowded clusters, and the aberrations produced in xylem of dis- 
torted or burly grain. In straight-grained xylem of the leaf-bearing cacti, 
the broader vessel members tend, on an average, to have transversely 
or nearly transversely oriented terminal perforation plates, whereas the 
narrowest vessels commonly have diagonally oriented ones. Where the 
grain of the xylem is distorted, as so frequently happens in stems and 
roots of the Jeaf-bearing cacti (owing in large measure to the frequent 
diagonal dissection of broad rays by conversion of ray initials to fusiform 
ones), steeply diagonal and even laterally placed perforation plates are of 
not infrequent occurrence. Similar aberrations occur in the orientation of 
sieve plates in the phloem. From the point of view of salient trends of 
phylogenetic specialization in the dicotyledons as-a-whole, the occurrence 
of simple perforation plates throughout both the primary and secondary 
xylem is of greater evolutionary significance than are deviations from a 
transverse orientation of perforation plates in the vessels of certain parts 
of the stems and roots of the leaf-bearing cacti. 

The occurrence of septate libriform fibers in various dicotyledonous 


1962 | BAILEY & SRIVASTAVA, CACTACEAE, IV 195 


families has been based largely upon surveys made of dried wood samples. 
It should be recognized, in this connection, that a great many of the 
samples consist of heart-wood. Where specimens of undecayed sapwood 
are available, they are sectioned after boiling in water and frequently 
after softening treatment in hydrofluoric acid. Such sections do not con- 
tain starch, but the assumption is made, probably correctly, that the 
septate fibers in the sapwood retained their living protoplasts at functional 
maturity and were capable, at least for a time, of storing starch. The com- 
mon occurrence of starch in nonseptate libriform fibers of the leaf-bearing 
cacti raises the question whether the storage of starch in nonseptate fibers 
of sapwood is not of much commoner occurrence in dicotyledons than has 
been realized. From physiological as well as phylogenetic points of view, 
the whole problem of retention of living protoplasts and storage of starch 
in nonseptate libriform fibers, whether confined to newly formed sapwood, 
whether capable of seasonal depletion and renewal as in the case of starch 
in ray tissue, etc., needs to be thoroughly investigated, not only in the 
Cactaceae, but also in other families of the dicotyledons. 

In this paper, we have omitted discussion of ray initials in the cambium 
and of their derivatives in the xylem and phloem. The rays in the leaf- 
bearing Cactaceae are obviously of a highly specialized form, being pre- 
vailingly multiseriate, uniseriate rays having been eliminated. It seems 
best to discuss rays in subsequent papers of this series, dealing in greater 
detail with the structure of various putative species of Pereskia, Pereskiop- 
sis and QOuiabentia. 


LITERATURE CITED 


Bartey. I. W. 1923. The cambium and its derivative tissues, IV. The increase in 
girth of the cambium. Am. Jour. Bot. 10: 499-509. 

—_—. 1936, The problem of differentiating and classifying tracheids, fiber 
tracheids and libriform fibers. Tropical Woods 45: 18-253. 

_ 1944. The development of vessels in angiosperms and its significance in 

morphological research. Am. Jour. Bot. 31: 421-428. 

__. 1957. The potentialities and limitations of wood anatomy in the study 

of the phylogeny and classification of angiosperms. Jour. Arnold Arb. 38: 

243-254. 


————. 1960. Comparative anatomy of the leaf-bearing Cactaceae, I. Foliar 
vasculature of Pereskia, Pereskiopsis and Quiabentia. Ibid. 41: 341-349. 
- 1961a. IL. Structure and distribution of sclerenchyma in the phloem of 
Pereskia, Periskiopsis and Quiabentia, Ibid, 42: 144-150. 
- 1961b. IIL. Form and distribution of crystals in Pereskia, Pereskiopsis 
and Quiabentia. Ibid. 334-340. 

___& W. W. Tupper. 1918. Size variations in tracheary cells. I. A com- 
parison between the secondary xylems of vascular cryptogams. gymno- 
sperms and angiosperms. Proc. Am. Acad. Arts Sci. 54: 149-204. 

Cueap_e. V. I. 1942. The occurrence and types of vessels in the various organs 
of the plant in the Monocotyledoneae. Am. Jour. Bot. 29: 441-450. 

_ 1943. The origin and certain trends of specialization of the vessel in 
Monocotyledoneae. Ibid. 30: 11-17 


196 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLII 


———. 1948. Observations on the phloem in the Monocotyledoneae, II. Ad- 
ditional data on the occurrence and rier ee in structure 
of the sieve tubes in the metaphloem. /bid. 35: 131; 
. 1956. Research on xylem and ee in fifty years. Jbid. 
43: 719-731 
& K. Esau. 1958. Secondary phloem of Calycanthaceae. Univ. Calif. 
Publ. Bot. 29: 397-510 
N. W. URL. 1948. “The relation of metaphloem to types of vascular 
bundles in the Monocotyledoneae. Am. Jour. Bot. 35: —583. 

& N. WHITFORD. 1941. Observations on the ae in the Mono- 
cotyledoneae, I. The occurrence and phylogenetic specialization in structure 
of the sieve tubes in the metaphloem. Am. Jour. Bot. 28: 623- soe 

Esau, K. Plant Anatomy. John Wiley & Sons, Inc., New 
CHEADLE. 1955. Significance of cell cighioue in differentiating 
eae phloem. Acta Bot. Neerl. 4: 346-357. 
& 1959. Size of pores and ha contents in sieve elements of 
dicotyledons. Prog. nine Acad. ae Ac —162. 
& E. FFORD, JR. 1953. ‘Compaitive ape and possible 
trends of cea of the cee m. Jour. 40: 9-19 
SCHENCK, H. 1893. ay zur Biologie meu ne ve Lianen. ‘Vol 2 7s 6 
148. Gustav Fischer. a 
se a 1899, aise Anatomie der Dicotyledonen. F. Enke. 
Stuttg 
ZAHUR, M. . 1959. Comparative study of secondary phloem of 423 spec 
of woody oo belonging to 85 families. Mem. N. Y. State College 
Agr. 358: 


EXPLANATION OF PLATES 
PLATE I 


Fics. 1-4. TANGENTIAL LONGITUDINAL SECTIONS OF CAMBIUM. XYLEM 
AND PHLOEM, X 110. 1, Pereskia cubensis Britt. & Rose [Atkins Gard.], storied 
fusiform eta of the cambium. 2, Pereskiopsis blakeana Ortega [Kimnach & 
Moran 82]|, diagonally storied peseel members and wood- -parenchyma strands. 3, 
Pereskia autumnalis (Eichlam) Rose [Moore 8210], storied sieve-tube members; 
terminal sieve plates are black. 4, Pereskia aff. sacharosa Griseb. [ Cardenas |, im- 
perfectly storied cambial Hea initials (left of ray) and phloem derivatives 
(right of ray). 


PLATE II 


Fics. 5-8. OCCURRENCE AND ORIENTATION OF SIMPLE PERFORATION PLATES 

IN VESSEL MEMBERS. 5, Pereskia grandifolia Haw. | Castellanos], transverse sec- 
tion of the secondary xylem showing transversely oriented perforation plates, x 
260. 6, Pereskia colombiana Britt. & Rose | Romero |, radial longitudinal section 
of the secondary xylem showing laterally placed perforation plate, 
Pereskia nicoyana Web. [ Rodriguez 662], tangential longitudinal section of the 
secondary xylem showing divergence of vessel ee through multiseriate 
ray; vessel members at right have perforation plates, as seen in sectional view 
at (X), in their lateral radial walls, X 180. 8, be aguosa (Web.) Britt. 
& Rose [Dressler], radial longitudinal section of the primary xylem showing 
perforation plates in helically thickened vessel members, X 260. 


1962 | BAILEY & SRIVASTAVA, CACTACEAE, IV 197 


PLATE III 


Fics. 9-15. PIrTING BETWEEN VESSELS AND BETWEEN VESSELS AND PAREN- 
CHYMA. 9, Pereskia grandifolia | Castellanos|, crowded alternate-multiseriate 
pitting in walls of contact between vessel members, X 410. 10, Pereskia colom- 
biana [Romero], pitting between vessel and ray parenchyma, X 410. 11, Peres- 
kia en [ Castellanos |, variations in form of bordered pits and their aper- 
tures, X 1130. 12, Pereskia colombiana | Romero |, unconformity i in the pitting 
of vessels; many Oi the transversely elongated pits in the wall of one vessel 
are in contact with rows of shorter pits in the wall of the adjoining vessel, 
x 960. 13, The same, scalariform and transitional pitting in lateral walls of 
adjoining vessels, X 410. 14, The same, scalariform pitting between vessel and 
parenchyma, & 410. 15, Quiabentia aff. chacoensis Backbg. [Tucuman], bor- 
dered pits with large apertures in the lateral walls of adjoining vessels, * 410. 


PLATE IV 


Fics. 16-22. LONGITUDINAL SECTIONAL VIEWS OF VESSELS, PARENCHYMA AND 
NONSEPTATE LIBRIFORM FIBERS. 16, Pereskiopsis blakeana [| Kimnach & Moran 
82], reduction in the borders and increase in size of apertures in intervascular 
pitting, X 410. 17, Pereskia aculeata Mill. { Atkins Gard.], iodine stained starch 
in a nonseptate ineionn fiber, X 750. 18, Pereskia bleo DC. [Atkins Gard.], 
reduction in borders and increase in size of apertures in pitting ne vessel 
and ray parenchyma, X 410. 19, Pereskiopsis chapistle Britt. & Rose [Boke 
B-3], nuclei (black) and unstained starch in nonseptate libriform fibers, 
1000. 20, Pereskiopsis blakeana | Kimnach & Moran 82], pitting between ves- 
sels and parenchyma, * 410. 21, Quiabentia aff. chacoensis |Tucuman], nucleus 
with nucleolus in nonseptate libriform fiber, X 1000. 22, Quiabentia pereziensis 
Backbg. [Cardenas], large unbordered pits between vessel and parenchyma, X 


PLATE V 


Fics. 23-32. DIAGRAMMATIC ILLUSTRATIONS OF CAMBIAL FUSIFORM INITIALS, 
LIBRIFORM FIBERS AND SIEVE-TUBE MEMBERS. 23, Pereskia sacharosa [Tucuman], 
fusiform cambial initials, drawn from tangential longitudinal section of cambium. 
24, Pereskia conzattu Britt. & Rose [Dressler], libriform fiber drawn from 
maceration. 25, Pereskia aculeata [Atkins Gard.], libriform fiber drawn from 
maceration. 26, 27, Pereskia conzatti [Dressler], septate and nonseptate libri- 
form fibers drawn from maceration. 28, Pereskia sacharosa [Tucuman], sieve- 
tube member and companion cells, ein from tangential eon Bee tion 
of phloem. 29, Pereskiopsis aff. chapistle [Boke B-3], sieve-tube member and 
companion cell, drawn from tangential longitudinal section of Set 30, The 
same, sieve-tube member drawn from radial longitudinal section. 31, The same, 
part of a sieve plate, drawn from radial longitudinal section of phloem. 32, The 
same, lateral sieve areas, drawn from radial longitudinal section, companion cell 
on right. Figures 23-30, * 175; figs. 31, 32, & 875. 


PLATE I 


Jour. ARNOLD Ars. VoL. XLIII 


= 


‘ASTAVA, LEAF-BEARING CACTACEAE, IV 


f 


BAILEY & SRIV 


PLATE II 


Jour. ARNOLD Ars. VoL. XLIII 


IV 


BEARING CACTACEAE, 


BAILEY & SRIVASTAVA, LEAF 


Jour. ARNOLD Arps. Vor. XLIII PiaTE III 


ae 


— = i — 


BaILey & Srivastava, LEAF-BEARING CACTACEAE, IV 


Jour. Arnoip Ars. VoL. XLITI PLATE IV 


BAILEY & SRIVASTAVA, LEAF-BEARING CACTACEAE, IV 


Jour. ARNOLD Ars. VoL. XLIII PLATE Y¥ 


rae 26 


BarLey & Srivastava, LEAF-BEARING CACTACEAE, IV 


1962] MOORE, CARAGANA SINICA 203 


ON THE ORIGIN OF CARAGANA SINICA? 


RAYMOND J. Moore 


THE SHRUB LONG KNOWN as Caragana chamlagu Lam. probably was 
introduced to Europe by Father Pierre d’Incarville, who sent seed of 
many plants collected near Peking, China, to the Jardin des Plantes, 
Paris, in the period 1740-1756 (Bretschneider, 1898). Loudon (1844) 
stated that the species was introduced to Great Britain in 1773. Appar- 
ently it has remained since the eighteenth century one oi the less com- 
monly cultivated shrubs. Rehder (1941) has pointed out that the over- 
looked epithet Robinia sinica of Buc’hoz has priority over that of Lamarck 
and, to avoid confusion, the name Caragana sinica (Buc’hoz) Rehder will 
be used exclusively hereafter, although Komarov, Pojarkova, ef al. have 
treated this entity under the name C. chamlagu. Chiefly on the basis of 
flower and calyx size, Pojarkova (1945) has divided C. sinica, sensu lato, 
into C. sinica and C. ussuriensis. 

Caragana sinica, sensu lato, occurs widely in eastern China and far- 
eastern Siberia. It is a shrub 1-2 m. tall, apparently common in dry, 
rocky, or other well-drained sites. The leaves consist of 2 pairs of leaflets, 
usually pinnately arranged, but often so close together as to appear almost 
palmate; the leaflets are obovate, 10-35 mm. long, 5-15 mm. broad, rather 
coriaceous, glaucous above, the base cuneate, the apex retuse, mucronu- 
late. The leaf rachis thickens and develops into a spine up to 25 mm. long 
which persists after the leaflets drop. The persistent rachises and the 
spiny stipules (5 mm.) give the bush its conspicuous spiny character. 
The flowers are large, 20-30 mm. long; the calyx 9-14 mm. long, 5-6 mm. 
broad, with lobes 2-3 mm. long; the pedicel attachment asymmetrical; 
and the calyx almost gibbous. The flowers are borne one or two per node 
on stalks 10-20 mm. long, articulated in the middle between pedicel and 
peduncle. The corolla is pale yellow, usually with a rose or bronze tinge 
which deepens with age. 

It is generally agreed that Caragana frutex and related species comprise 
the only species-group to which C. sinica is clearly related. Komarov 
(1908) placed C. sinica in his series FRUTESCENTES. He recognized its 
unique character and regarded it (oc. cit., p. 370) as the closest claimant 
to the position of generic prototype. He based this view on its leaf char- 
acter, noting that at an early age the leaf is palmate but later becomes 
pinnate. This transition he held to be a phylogenetic recapitulation, ap- 
parently regarding palmate leaves as ancestral to pinnate. Although not 
clearly stated, it appears that Komarov did not refer to the form of early 
seedling leaves but to the varying forms seen on adult plants. He believed 


1 Contribution No. 128 from the Plant Research Institute, Canada Department of 
Agriculture, Ottawa. 


204 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


(p. 371) that different environmental conditions then had resulted in the 
evolution from C. sinica of both palmate-leaved species (FRUTESCEN TES) 
and pinnate-leaved species (ALTAGANAE) and subsequently from these, 
all the other series. Caragana sinica thus assumes special importance as 
the foundation of Komaroy’s phylogenetic scheme. Doubtless as a re- 
sult of this opinion, he designated east Asia (range of C. rosea, C. fruti- 
cosa) as the primary center of species formation. 

To show specific relationships more exactly, Pojarkova (1945) divided 
the series FRUTESCENTES Kom. into three more homogeneous but related 
series: CHAMLAGU, FRUTESCENTES emend., and GRANDIFLORAE. The series 
CHAMLAGU Pojark. consists of C. sinica and C. ussuriensis (Regel) Pojark. 
The latter taxon had been described by Regel as C. frutex var. USSUTIENSIS, 
and, as such, was placed in the synonymy of C. sinica by Komarov (1908). 

Pojarkova separates Caragana ussuriensis and C. sinica chiefly on floral 
characters: flower length, 23-25 mm. vs. 28-30 mm.: and calyx size, to 
9 mm. long X 5 mm. broad vs. 12-14 mm. long & 6 mm. broad. The 
leaflets of C. stnica are said to have a broader, obovate blade and broader 
base, giving a more rounded outline, whereas those of C. ussuriensis are 
narrow with a narrowly cuneate base. Only C. ussuriensis occurs in the 
U.S.S.R. (region of the river Ussuri just north of Manchuria) from whence 
it extends southward into Manchuria and northern China. It is reported 
that C. sinica has a more southerly distribution in China: Hopeh (Chili) 
to Yunnan. The impression is conveyed by Pojarkova that the range of 
C. ussuriensis is more northerly than that of C. sinica and that, although 
they overlap in northern China, C. sinica alone occurs in southern China. 
The two taxa are illustrated in Fics. 1 and 2. 

The illustration accompanying Buc’hoz’s (1779) description of Robinia 
sinica is apparently drawn life-size and shows the larger-flowered plant. 
Mature open flowers are 30 mm. long, the calyx being 11-12 mm. long 
and 6 mm. broad, measurements which fall within the limits indicated 
by Pojarkova for C. sinica. 


MATERIALS AND METHODS 


In the course of a survey of chromosome numbers in Caragana, a special 
attempt has been made to obtain living material of C. sinica. Seeds have 
been received under this name from twelve botanic gardens (eleven Euro- 
pean) but the resulting seedlings proved to be not the desired species but 
typical plants of C. frutex, C. arborescens, C. pygmaea, or C. aurantiaca. 
Some of these species are not closely related to C. sinica; all are readily 
distinguished from it and it is considered that the error is due solely to 
incorrect identification of the seed parent. In two cases, leaflets in the 
seed packet indicated that the source of the seed was C. arborescens and 
C. pygmaea, as were the resulting seedlings. 

Living plants of Caragana sinica have been obtained from only one 
source: F. J. Grootendorst & Sons, Boskoop, Holland. The species has 
been propagated asexually in this nursery. It was received there at some 


1962 | MOORE, CARAGANA SINICA 


L, Baikal 


CHINA -*..-° 


one 
3 ee 
$0 
oe. ai 
: @o0 
e eh 
are soe 
eo? 


U.S,S.R- 


MANCHURIA 


= 


JAPAN 


Y 


ussuriensis 
sinica 


intermediate 


Fics. 1-4. pp eae ae details and distribution of oh iaee simica. 1, ae 


and flower of C. sin 2. Lea 


lato, 


C. ussuriensis. 
plement of C. sinica “(eugene triploid). 


3, Somatic chromosome c 
4, Dcebition of C. sinica, sensu 


206 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


time before 1928, but the source is unknown (H. J. Grootendorst, personal 
communication, 1961). 

Cuttings of the rare species Caragana rosea were received from the 
Arnold Arboretum, Harvard University, where there is a single shrub 
grown from seed collected by William Purdom (seed lot 9a), in 1909, in 
Weichang, Chili (Hopeh) Province, China. This collection was determined 
as C. rosea by Rehder (1926). 

Herbarium material of Caragana sinica from the Arnold Arboretum 
(A) and the Gray Herbarium (cu) of Harvard University and from the 
United States National Museum (us), as well as that in the Herbarium, 
Canada Department of Agriculture (pao) has been examined. Particular 
attention has been paid to the characters which separate C. sinica and 
C. ussuriensis and to the condition and size of the pollen. Pollen was 
mounted in 45% acetic acid and stained with dilute cotton blue. Grains 
with a protoplast which completely filled the cell and stained blue were 
counted as viable. The grains are thin walled and smooth; the diameter 
stated is the measurement to the outer surfaces. 


MORPHOLOGY, POLLEN CONDITION, AND 
NATURAL DISTRIBUTION 


Little variation was found in the size of the flower on any single her- 
barium specimen, and it is believed that these size-characters are reason- 
ably constant for any individual. A similar opinion has been formed 
from observation of numerous living plants of various species of Caragana 
over a number of years, and it is believed that the size-characters used 
by Pojarkova reflect genetic differences and are reliable for systematic 
purposes. 

Following the floral and leaf characters designated by Pojarkova, 34 of 
38 herbarium specimens were referred either to Caragana ussuriensis 
or to C, sinica. The determination of the remaining four, all wild Chinese 
collections, was less certain. Two had the flower size of C. sinica with a 
smaller calyx, and two had the smaller flower of C. ussuriensis with a 
calyx similar in size and appearance to that of C. sinica. 

The localities of the Asiatic collections are plotted on the map of 
eastern Asia (Fic. 4) and are listed below. A few collections were omitted, 
either because of uncertainty about the location or because only a gen- 
eral area was indicated. Symbols were placed in the center of a province 
when data or knowledge were insufficient to place them more accurately. 

It is concluded, in agreement with Pojarkova, that the small-flowered 
plant occurs from northern China to Siberia and that the large-flowered 
plant is dominant in southern China. The small-flowered plant seems to 
extend as far south as approximately 27° N, well into the range of the 
large-flowered entity, and a few intermediates were found in the zone 
of central China in which both entities occur. 

Pollen of 34 herbarium specimens, 27 of these wild Asiatic collections, 
was examined, and in none was the pollen 100% normal. The samples 


1962] MOORE, CARAGANA SINICA 207 


consisted of mixtures of normal grains with full, densely staining proto- 
plasts; grains full but with a thinner, lightly staining content; grains 
only partially filled with a thin protoplast; grains empty and collapsed; 
and empty micro-grains. Difficulty was experienced in classifying some 
grains since the first two classes intergrade. Only grains with a full, 
dense protoplast were classed as normal in the following counts. The 
micro-grains probably were formed from lagging chromosomes excluded 
from the tetrad nuclei and indicate a high degree of meiotic irregularity. 

The maximum percentage of normal pollen found among the herbarium 
specimens was 80, the minimum, 3; 12 lay in the range 0-39%, 14 had 
40-50%; 8 had over 50%. The size of the normal grains ranged from 
18 to 30 » but this range was not found in any single sample. The modal 
range was 20-40 » (13); in 6 samples the lower limit was 18-20 p», and 
the upper limit was lowered correspondingly. The range in 10 samples 
was from 20 p to 25-27 yp, and in 6 the smallest grains were 23 yp, the 
largest 27-30 

Pollen measurements (unpublished) made on other Caragana species 
do not show a correlation between the diploid and tetraploid chromosome 
number and pollen size. The samples are almost invariably 100% normal. 
Pollen of seven diploid species falls in the range 20-27 »; two additional 
diploids measured 23-29 » and 28-30 p». Pollen of one tetraploid species 
(C. frutex) measured 20-27 yp, of another (C. spinosa) 24-34 pw. No 
correlation between pollen size, probable chromosome number, or mor- 
phological features is believed to exist in C. sinica, and the pollen size 
variability is attributed to gene differences between the various popula- 
tions. The significant feature lies in the consistent absence of wholly 
good pollen and the similarity of the condition of all samples. No dif- 
ference was found in the range of size or of fertility between specimens 
classed as C. sinica, C. ussuriensis, or intermediate. Pollen samples of the 
latter group fall in the range of 40 to 60% normal and thus were no more 
“hybrid” than those of the more typical groups. No certain correlation 
was found between the degree of pollen fertility and geographical loca- 
tion, although there may be a tendency for plants with the higher per- 
centages (70% plus) to occur on the extremes of the range (Northern 
China and Yunnan). 


Caragana sinica (Buc’hoz) Rehder, Jour. Arnold Arb. 22: 576. 1941. 
Robinia sinica Buc’hoz, Pl. Nouv. Decouv. 24. ¢. 22. 1799. 
Flowers 27-30 mm. long, calyx 11 mm. or more, leaflets broadly obovate. 


China. CHEKIANG: without location, Barchet 146 (us). FUKIEN: Diongloh 
Hsien, Chung 1239, Apr. 5, 1923 (A). Hope (Chili): Hsi Yu Ssu, Liu 2286, 
June 8, 1929 (A). Hunan: Tschangscha, Handel-Mazzetti 638, Apr. 14, 1918 
(a). Hupeu: W. Hupeh, Wilson s.n., Apr. 1900 (us); W. Hupeh, Wilson 2203, 
May 1907 (cH); without location, Henry 5378 (cH), 3812 (us). Kansu(?): 
Fengwangschan, Forbes 113, Apr. 22, 1877 (A). Kriancsu: without location, 
Tsu 436, Apr. 26, 1920 (a): Yun Dai Shan, Nanking, Tso 79, Apr. 17, 1926 


208 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


(a): without location, Faber s.2. (A). KwetcHow: Kweiyang, Teng 90079, 
Apr. 15, 1936 (A). YUNNAN: East of Tengyueh, Forrest 19343, _ Apr. 1921 
(a, uS); Yunnanfu bei Puchli, Schneider 214, Mar. 7, 1914 (a); zwischen Ssiao 
und Schin lung, Schneider 4046, Mar. 9, 1914 (a, a - without ee Bonatt 
or Maire 7286 (us); Yunnanfu, Smith 1582, 8/3, 1922 (aA) 

Cultivated. Japan. Hondo: Musashi, Teizo a ee y 2, 1904 (A); without 
location or collector, May 13, 1910 (us 1311798); Nagasaki, Maximowicz s.1., 
1863 (cH). FRANCE: Paris, Gay, s.n., 1822 (GH). Bavaria, Scherzer s.1., 
Apr. 1, 1906 (pao). Unrrep States: Washington, D.C., Apr. 24, 1886 (us 
137850); Washington, D.C., 1915 (us 786433); Chico, Calif., Apr. 25, 1922, 
S.P.I. 22981 (originally collected in Soochow, Kiangsu, China) (A). CANADA: 
Royal Botanic Gardens, Hamilton, Ont., Rhodes & Florian 3587, June 11, 1952 
(pA0); Dominion Arboretum and Botanic Garden, Ottawa, Ont. Woore, 53- 
249-1, May 27, 1960 and Moore, 60-231-45, May 29, 1961 (pao); Experi- 
mental Station, Morden, Man., Rhodes & Vitens 4583, Sept. 10, 1953 (DAO). 


Caragana ussuriensis (Regel) Pojark. Flora U.S.S.R. 11: 395. 1945. 


ses frutescens 2 ussuriensis Regel, Mem. Acad. Sci. St. Petersb. ser 
(4): 44. 1861. 


Flowers 23-26 mm., calyx to 9 mm., leaflets narrowly obovate with 
cuneate base. 

China. Hopen (Chili): Western a eee 3 21361, May 7, 1929 
(A); Nankow, Peiping, Chiao 21253, May 11, se Hunan: Yi Chang 
District, Tsang 23430, Mar. 91-29, 1934 ay. ee URIA: Ussurl super, 
Maximowicz s.n., 1860 (us). NortH CHINA: without a. Bunge s.u. (GH). 

Cultivated. JAPAN: without location, Zuccarini s.n., 1842 (GH). 

Specimens intermediate between Caragana sinica and Caragana ussuri- 
ensts. 

China: Hupen: W. Hupeh, Wilson 2203, May 1907 (us); Patung, W7lson 
s.m., Apr. 1907 (A). Kranosr: Kipkiang, Bullock 151, Apr. 20, 1892 (us). TFu- 
KIEN: Diongloh Hsien, Chung 1239, Apr. 5, 1923 (A) 


CULTIVATED CARAGANA SINICA 


The five plants purchased from the Grootendorst nursery are identical 
in appearance, doubtlessly having been propagated asexually from a single 
plant. Flowers are 30 mm. long, with the calyx 11-12 mm. long and 6 
mm. broad. The leaflets are broadly obovate, to 20 mm. long, 9 mm. 
broad, coriaceous, and glaucous. On the basis of flower and calyx size and 
leaflet shape, the plants are to be classed as the larger-flowered species 
C. sinica sensu Pojarkova. In all characters they are sidictineuichable 
from collections of other cultivated and wild Asiatic plants. 


Cytotocy. The chromosome number was determined from leaf squashes 
to be 2n = 24 (Fic. 3). The basic number of the genus is 8: hence this 
number is triploid. Meiosis was studied in one plant growing outdoors 
at Ottawa. The configurations at Metaphase I could be fully analyzed in 


1962 | MOORE, CARAGANA SINICA 209 


only eight cells but these seem to be representative. The minimum pair- 
ing seen was 16 univalents and 4 bivalents; 8 bivalents and 8 trivalents 
were seen in two cells. The trivalent was the largest association found, 
and three trivalents was the maximum number found in a single cell. The 
average pairing for eight cells was 8! + 6.71 4+ 0.86UI 

Approximately 40% of mature pollen grains are normal in appearance. 
Grains judged to be normal have a full, dense protoplast and measure 
20-24 » in outer diameter. Pollen was spread on a mixture of 2% agar 
plus 5% sucrose, and germination of at least 50% of these full grains 
was observed. This agar medium has been found to give good germination 
of pollen of many species of Caragana. 


FLowER Biotocy. At Ottawa, the shrubs flower from late May to 
mid-June, meiosis occurring in buds 5-6 mm. long in the period May 
10-17. Over the past ten years it has been observed that varying weather 
conditions cause little variation in the onset of flowering in Caragana 
species, at most, two to three days. 

Small aborted pistils were noted, but probably not over 10% of the 
flowers are defective. No other abnormality was observed. The style 
elongates markedly and extends approximately 2 mm. beyond the keel 
at anthesis. This feature favors cross-pollination and probably makes 
insect action necessary even for self-pollination. The plants studied were 
located approximately 100 feet from other Caragana bushes (C. arbo- 
rescens, C. frutex, C, aurantiaca) and separated by various other trees 
and shrubs. It is highly probable that the 5 plants of C. sinica were not 
pollinated from other species. Most flowers dropped without ovary en- 
largement; some showed slight enlargement but dropped in two to three 
days. A smaller number, estimated at 8-10%, developed conspicuously 
enlarged ovaries (25-28 mm.) which turned green and for a week ap- 
peared to be forming seed. However, all turned brown and fell. Some 
of the shrubs have been observed for three years and no seed has been 
formed. Plants from the same source grown at the Experimental Farm, 
Morden, Manitoba also are sterile (personal communication). Pollen 
of C. sinica was used to pollinate shrubs of C. arborescens, C. microphylla 
and C. frutex but no seed resulted. Two of 38 flowers of C. frutex volli- 
nated by C. sinica showed slight ovary enlargement before ¢:°:.j:ng. 


OvuLE Histotocy. Ovaries of various sizes, pre- and post-flowering, 
were fixed for histological study of the ovules. Several different types of 
ovule development were observed, but in all cases all ovules within a 
single ovary were similar. 

Ovaries which lacked ovules or contained abnormal and partially de- 
veloped ovules were found. Undeveloped ovules consisted of a small 
nucellus partially enclosed by a tissue two or three cells thick which 
presumably represented the outer, or perhaps both integuments. This 
integument did not enclose the micropylar end of the nucellus. No sporo- 
genous tissue was differentiated and all tissues appeared unhealthy. Such 


210 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


ovules were found in mature open flowers, a stage when mature embryo 
sacs should have been present. 

Structurally normal ovules were found in ovaries of some mature 
flowers. These ovules, approximately 525 » long and 375 » broad con- 
sisted of two integuments and nucellus, all apparently normal. Cell for- 
mations suggestive of aborted embryo sacs were observed in the nucellus. 
A linear formation of three compact cells, the micropylar one of which 
was the largest, was probably the product of meiosis. The innermost of the 
three was the most healthy and normally probably would be the func- 
tional megaspore. A large two-nucleate cell surrounded by disorganized 
tissue seemed to be an aborted early embryo sac. A four-nucleate cell 
(75 w long, 15-20 » broad) also was apparently an immature embryo sac. 
The most advanced structure seen was interpreted as a disintegrating 
mature embryo sac. This structure was 125 » long, 20 » broad and con- 
tained five recognizable nuclei and remnants of at least two antipodal 
cells. In all the above examples both sporophytic and gametophytic tis- 
sues were obviously in an unhealthy condition and incapable of continued 
development. Many ovules were so disorganized that the stage of embryo 
sac development could not be determined. 

The large ovaries mentioned above (to 28 mm.) contained ovules that 
were enlarged by some 50% beyond the size of the most normal mature 
ovules. However, none of these contained embryos. Counts of the cell 
layers and measurements of representative cells indicated that ovule 
enlargement was due solely to cell enlargement in both the integuments 
and nucellus. By the time such ovaries drop, breakdown of the nucellus 
is advanced. A patch or narrow zone of empty collapsed cells in the 
area in which an embryo would normally be found is first seen. Dissolu- 
tion of cell contents and collapse of the walls progresses through the 
middle of the nucellus from the micropylar to the antipodal end. It was 
usually observed that the two-celled peripheral layer of the nucellus at 
its micropylar end remained longest in apparently healthy condition. A 
zone of the inner integument adjoining the micropylar end of the nucellus 
appeared more active than the remainder of the integument endodermis. 
These observations together suggest that the integument cells at this point 
were absorbing the contents of the nucellus. In the final stage of ovule 
collapse only remnants of walls remained of the nucellus. No evidence 
of apomictic seed formation was observed and there is no reason to be- 
lieve that these plants would ever set seed through either sexual or apomic- 
tic processes. 


DISCUSSION 


The occurrence of a major amount of aborted pollen in the living cul- 
tivated (Grootendorst) plant and in all known collections of Caragana 
sinica from the wild suggests that the species is of hybrid origin. Whether 
the species in nature is triploid, like the cultivated plant studied, cannot 
be decided with the available evidence. 


1962 | MOORE, CARAGANA SINICA vans 


Morphology of the species alone suggests a hybrid origin. That the 
leaf, which is pinnate with two pairs of leaflets, frequently appears to 
be palmate due to the lack of elongation of the rachis suggested to 
Komarov that the species was ancestral to both the pinnate and the 
regularly palmate series. An alternative explanation of this phenomenon 
is that C. sinica is a hybrid between a pinnate- and a palmate-leaved spe- 
cies. The variable-leaf type occurs otherwise only in the series SpINOSAE 
and DAsyPHYLLAE, groups of central Asia morphologically very unlike 
C. sinica. It seems obvious that these do not bear on the present problem. 

All authors agree that Caragana sinica is most closely related to C. rosea 
Turcz. The affinity appears in the number, shape and texture of the leaf- 
lets, the persistent spiny leaf rachis, the large flower (more than 2 cm.) 
which is pale yellow with a rosy tinge, and the large calyx which is longer 
than broad. The rose flower color is not known in other species of eastern 
China, Caragana rosea has a range in eastern Asia (Manchuria, Hopeh, 
Honan, Kansu, Chekiang [Rehder, 1926]) very like that of C. sinica. 

If the hybrid nature of Caragana sinica is accepted, C. rosea must be 
proposed as one parent. A pinnate-leaved species of eastern China is re- 
quired as the other parent, but the exact species cannot be named with 
an equal degree of assurance. The large flowers of C. sinica, larger even 
than those of C. rosea, point to another large-flowered species as the 
second parent. Such is to be found in C. microphylla Lam. (flowers 25 
mm., calyx 9-12 mm.) a species of suitable leaf type and range as well. 

It is therefore postulated that Caragana sinica, sensu lato (including 
C. sinica (Buc’hoz) Rehd. and C. ussuriensis (Regel) Pojark.) is a hybrid 
between C. rosea Turcz. and a pinnate-leaved species, probably C. mi- 
crophylla Lam. In the opinion of the author it would be preferable to recog- 
nize the entities simica and ussuriensis at infraspecific level under C. sinica 
(Buc’hoz) Rehd. It seems obvious that they have shared a common origin 
and are separated by characteristics of a minor order, quantitative rather 
than qualitative. The two populations have achieved geographic separa- 
tion, and taxonomic recognition is justified. 

A variation within Caragana rosea in flower and calyx size and leaflet 
shape, parallel to that used by Pojarkova to split C. sinica was noted in 
thirteen specimens (US) examined. These could be divided into large- 
flowered (flower 25-27 mm., calyx 9-11 mm.) and small-flowered (flower 
20-24 mm., calyx 6-8 mm.) plants. The more rounded leaflet shape was 
not invariably associated with the greater flower size, nor was there an 
evident geographical correlation. The range in pollen size in C. rosea was 
20-27 ww. The variation within a single plant was not more than 4 np. 
However, no correlation between larger pollen size and flower size was 
found. The existence in C. rosea of variation of the same type as that 
seen in C. sinica is an additional indication of their close affinity. 

The chromosome number of the single available accession of Caranga 
rosea was found to be 2x = 16. This diploid number is surprising, since 
it might be expected that this species would be tetraploid like the closely 
related C. frutex. The latter was reported to be tetraploid (2n = 32) by 


Zig JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


Tschechow (1930, as C. frutescens), and only this number has been found 
in nine accessions in the present work. Caragana rosea appears to be mor- 
phologically a more advanced species than C. frutex. The persistent 
thickened petiole seen in C. rosea is apparently developed from the de- 
ciduous petiole of C. frutex. Throughout the genus there is an evolu- 
tionary trend toward transformation of the deciduous petiole, first, to a 
persistent but essentially unthickened petiole and, finally, to a persistent 
and much thickened organ. The latter is the stout spine seen in many 
species. This trend is found in both the pinnate- and the palmate-leaved 
series. 

The lower chromosome number of Caragana rosea can be reconciled 
with the view that the species is derived from C. frutex by postulating 
that diploid populations of C. frutex do or did exist. It is, indeed, not 
improbable that the series FRUTESCENTES arose at the diploid level from 
a pinnate-leaved ancestor. All pinnate-leaved species yet examined are 
diploid (Moore, 1958 and unpublished). The series FRUTESCENTES Kom. 
em. Pojark. contains six species which extend from the Black Sea to 
central Mongolia. Three other Chinese species not treated in the Flora 
of the U.S.S.R. but which should doubtless be referred to the series, ex- 
tend the range of the series across northern China and Manchuria to the 
Pacific. Caragana frutex has by far the largest range of any single species 
(Black Sea to northwest Mongolia). Four species of relatively narrow 
distribution are found in Central Asia, south of Lake Balkhash. It seems 
possible that ancestral diploid ‘‘frutex” has spread from central Asia east- 
wards, developing in the most eastern part of the series range into C. 
rosea. The morphologically more primitive “frutex” stock may have 
become autotetraploid in central Asia, and, enjoying an advantage, accom- 
plished the present wide distribution, particularly to the west and north 
from Central Asia. It seems probable that Caragana frutex in cultivation 
has come from the European part of the range, the western extreme, and 
thus is tetraploid. Diploid populations may still exist in Central Asia. 
Unfortunately the chromosome numbers of the more limited species of 
Central Asia are unknown. The range of C. rosea is second only to that 
of C. frutex, suggesting considerable age for the former species. 

The triploid number of the Grootendorst plant of Caragana sinica sug- 
gests a hybrid origin between a diploid and a tetraploid species. Two 
serious difficulties at once arise. The tetraploid number is not known 
in either postulated parent species. A triploid hybrid, if at all like the 
Grootendorst plant, will be seed sterile, and it seems impossible that the 
extensive range of C. sinica was accomplished by any means other than 
by seed dispersal. Moreover, it is known that the species in China does 
set seed. 

These difficulties may be relieved by suggesting that Caragana sinica 
in eastern Asia is diploid, a hybrid between diploid C. rosea and C. mi- 
crophylla. A reduced, but still appreciable, seed fertility will then be pos- 
sible. The triploid condition of the Grootendorst plant may have arisen in 
cultivation. The plants are morphologically indistinguishable from wild 


1962 | MOORE, CARAGANA SINICA 213 


collections, and it is therefore improbable that a cross with a different 
tetraploid species has occurred in cultivation. The triploid condition may, 
owever, have developed from the fertilization of an unreduced egg. In- 
deed, the different climactic conditions of Europe acting on a somewhat 
unstable hybrid genome may have caused the formation of an unreduced 
gamete. By random chromosome segregation, the additional haploid set 
could add an equal number of chromosomes from each parent to the 
diploid hybrid complement, converting some former bivalents into triva- 
lents. The presence of equal numbers from each parental species might 
maintain the gene balance to such a degree that the triploid appears iden- 
tical with the diploid. The absence of a noticeable difference in pollen 
grain size between diploid and triploid may be disregarded since no cor- 
relation between chromosome number and pollen size has been observed 
in the genus, and it is believed that pollen size depends solely on the 
genes governing this character. 

It may be pointed out in conclusion that the widespread occurrence 
of pollen abortion in Caragana sinica is explained better by the hybridity 
hypothesis than by the prototype hypothesis of Komarov. It seems 
improbable that an ancestral stock would have retained a condition of 
abortive pollen, presumably due to meiotic irregularities, which had been 
eliminated from its descendants. 


SUMMARY 


It has been possible to obtain Caragana sinica (Buc’hoz) Rehd. from 
only one source in cultivation. The plant is triploid (2n = 24), pollen 
is 40% normal, no seed is set. All herbarium specimens, of wild and 
cultivated collections, have partially aborted pollen. It is postulated that 
C. sinica is a hybrid between C. rosea Turcz. and probably C. microphylla 
Lam. The hybrid in nature is probably diploid; the triploid cultivated 
clone may have arisen in Europe by the production of an unreduced 
gamete. The chromosome number of C. rosea Turcz. is 2n = 16 


The author wishes to express appreciation to the curators of the herbaria 
from which material was borrowed and particularly to members of the 
staff of the Arnold Arboretum and Gray Herbarium with whom the author 
has corresponded. Dr. J. L. Thomas, formerly of the Arnold Arboretum, 
was most co-operative in making observations and procuring specimens 
and cuttings of the Caragana rosea in the Arboretum. At the author’s 
request, Mr. H. J. Grootendorst looked into the source of the Caragana 
stnica sold by their nursery. Dr. T. Koyama, University of Tokyo, has 
informed the author that collections of C. sinica from Japan are undoubt- 
edly introductions. 


214 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLIII 


BIBLIOGRAPHY 


BRETSCHNEIDER, E. 1898. History of European botanical discoveries in China, 
Vol. 1. London. 

KOMAROV, yi L. 1908. Generis Caraganae monographia. Acta Horti Petrop. 
29(2): 177-388. 

Loupon, J. C. 1844. Arboretum et fruticetum Britannicum. Ed. 2. Vol. 2. 
London. 

Moore, R. J. 1958. The status of Caragana Boisti, Baileya 6: 188-193. 

Poyarkova, A. I. 1945. Caragana. In: Flora of the U.S.S.R. 11: 327-368. 
Moscow and Leningrad. 

Renper, A. 1926. Enumeration of the ligneous plants of northern China III. 
Tour Arnold Arb. 7: 164-65. 

. 1941. New species, varieties and combinations from the collections of 
the Arnold Arboretum. Jour. Arnold Arb. 22: 569-579. [Caragana sinica 
(Buc’hoz), comb. nov., pp. 576, 577.] 

TscHECHOW, WL. 1930. arvelocicel systematische Untersuchung des Tribus 
Galegeae, Fam. Leguminosae (Vorliufige Mitteilung). Planta 9: 673-680. 


1962] PACLT, NEW GENUS OF SCROPHULARIACEAE 215 


SHIUYINGHUA, A NEW GENUS OF SCROPHULARIACEAE 
FROM CHINA 


J. PACLT 


IN HER MONOGRAPH of the genus Paulownia Sieb. & Zucc. (Scrophu- 
lariaceae), Hu (1959, p. 47) excluded P. silvestrii Pampanini & Bonati 
(1911) from that genus and transferred it to Catalpa Scop. (Bignoniaceae). 
I have had the opportunity of examining a photograph of the type speci- 
men, P. C. Silvestri 3286 (PLATE I), which shows characters of a tree 
clearly different from Catalpa. The characters which do not permit one 
to classify this plant as a member of the genus Catalpa may be summarized 
as follows: 

1. The conspicuously dimorphic shape of leaves which are broadly 
ovate (cordate) in the axial position and elliptically lanceolate in the 
abaxial position on flowering branches. 

2. The flowering branches which bear both leaves and almost laterally 
situated (axillary) loose cymes of flowers, as in Paulownia fortunei (Seem.) 
Hemsl. and substantially all other species of Paulownia. In Catalpa, the 
inflorescences are formed terminally and correspond to racemes or true 
panicles (thyrses). 

3. The calyx which is five lobed and patelliform, as in Paulownia but 
not Catalpa in which the calyx is gamosepalous, splitting into two strongly 
convex lobes at anthesis. Also, the flower buds are generally oblong in 
Paulownia silvestrii, instead of showing the typically subglobular shape 
of the bud of Catalpa. 

Although the fruit of Paulownia silvestrii is unknown, the original 
description of this species by Pampanini and Bonati leaves, in my opinion, 
no serious doubts about its correct position in the system. This place is 
to be found in the tribe PAULOWNIEAE Pennell of the family Scrophu- 
lariaceae. 

However, the general shape and consistence of the flower buds of Pau- 
lownia silvestriti alone separate this species sufficiently from all known 
members of Paulownia. This character also seems to be Hu’s basic reason 
for excluding P. silvestrit from Paulownia. In addition, her analysis of 
a flower bud of P. silvestrii revealed another difference concerning the 
structure of the young stigma (Hu 1959, p. 47). Some apparently less 
important differences between P. silvestrii and the other species of Pau- 
lownia may be found in the general appearance of the flowering branches 
and in the size of flowers. Accordingly, Paulownia silvestrii is best inter- 
preted as the type of a distinct new genus which I have the pleasure of 
naming in honor of Dr. Shiu-ying Hu, of the Arnold Arboretum, to whom 
botany is indebted for her contributions to the knowledge of Chinese 
plants. 


Jour. ARNOLD Ars. VoL. XLIII PLATE I 


j 


HoLotyPE OF PAULOWNIA SILVESTRII (Silvestri 3286) 


1962 | PACLT, NEW GENUS OF SCROPHULARIACEAE ral 


Shiuyinghua, gen. nov. 


Plantae lignosae foliis simplicibus oppositis vel etiam ternatim verticilla- 
tis. Inflorescentia axillaria cymosa. Alabastra oblonga. Calyx lobis 5 
membranaceis vel + crassis. Corolla gamopetala quinquefida tubo inflato, 
lobis subaequilongis. Stamina didynama basi tubi inserta. Stigma (in 
alabastro visum) bilamellatum. Fructus ignotus. — Hab.: Asia temperata. 


PUS GENERIS: Shiuyinghua silvestrii (Pamp. et Bonati), comb. nov. Pau- 
oe suvestri Pampanini et Bonati in Pampanini, Nuov. Giorn. Bot. Ital. II. 
18: 177. 1911. Catalpa silvestrii (Pampanini et Bonati) S. Y. Hu, Quart. Jour. 
Taiwan Mus. 12: 47. 1959 

TERRA TypICA: China, provincia Hupeh, praeter ripam fluminis VYang-tze 
Kiang (et praecipue fluvii Han Kiang), alt. 700 m., 20-30 Juni 1907. P. C. 
Silvestri 3286 in hb. Fr (et fragmentum, a). 


The new genus Shiuyinghua becomes now the second known genus of 
the tribe PAULOWNIEAE. However, another genus, Wighktia Wallich, is 
sometimes considered to belong here also. Wightia is likewise a wood 
genus and is represented in the Himalayan, Burmese, Chinese (Yunnan), 
Vietnamese (Tonkin), and Malayan floras. The relatively limited knowl- 
edge of the morphology of Shiuyinghua does not make it possible to 
elaborate for the time being more than the following key to the practical 
identification of the three genera. 


Stamens surpassing the top of corolla; calyx entire, tight-fitting. .... Wightia. 
Stamens not surpassing the top of corolla; calyx with 5 + outstanding lobes. 
Flower buds oblong and slender, with stigma ee bilamellate at this 
stage of immaturity; corolla not longer than 3 cm. ........ Shiuyinghua. 
Flower buds broadly ovate and robust, with a appearing punctiform 
at this stage of immaturity; corolla at least 3 cm. long, mostly much 
|e Nec mm sien A Orn ICR An Ae A a ev _ Paulownia. 


My warmest thanks are due to Professor Richard A. Howard, director 
of the Arnold Arboretum, for his valuable help in sending me a photo- 
graph of the type specimen for study, as well as to Dr. Carroll E. Wood, 
Jr., editor of this journal, for his very kind criticism. 

Poe INSTITUTE, 


K ACADEMY OF SCIENCES, 
on CZECHOSLOVAKIA 


218 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLIII 


PHENOLOGY OF TROPICAL PINES? - 


NicHo.as T. Mirov 


IN THE TEMPERATE ZONE pines shed their pollen during the season 
of the year vaguely designated as spring. At that time pollination occurs, 
and the ovulate strobili begin to develop slowly and, in warmer parts o 
the zone, continue to develop throughout the ensuing winter.* Ovules are 
fertilized the next spring, the cones develop throughout the second grow- 
ing season, and the seeds ripen in the fall. Time elapsed between pollina- 
tion and ripening of the seed thus is equal to about 15 or 16 months and 
embraces two summers and one winter. The farther south, the earlier 
pine pollen flies. Of course, many local environmental and genetic factors 
determine the time of pollen shedding. Pinus radiata D. Don sheds pol- 
len in March both in its naturalvenvironment on the Pacific Coast and in 
low elevations of the Sierra Nevada. It is interesting that when Pinus 
radiata is planted in the southern hemisphere, for example, in New Zealand 
where it is known as Pinus insignis Douglas, it sheds pollen when it is 
spring there, i.e., during August or September. 

It should be noted that photoperiod sensu stricto does not affect flower- 
ing of pines; they are photoperiodically neutral. That is, when a northern 
(long day) pine is moved to a more southern (shorter day) location, its 
flowering pattern is not changed. When a tropical (short day) pine is 
cultivated in a more northern nee? day) latitude, it continues to flower 
as freely as in its southern home.* 

The closer to the Equator, the more distorted is flowering in pines. 
Even in the southern parts of the United States, pines, for instance Pinus 
elliottii Engelm. in southern Texas or in northern Florida, shed their pol- 
len sometimes as early as the end of January. When you go farther south 
to the highlands of Mexico, early ‘flowering’ of pines becomes a wide- 
spread phenomenon, and its relation to the four seasons of the year be- 
comes really distorted. 

I had occasion to observe Pinus oocarpa Schiede at the southernmost 
limits of pine distribution in Nicaragua. It was on a south slope of the 
mountains at an elevation of 4000 feet above sea level. It was the middle 
of February; the trees had just completed blooming (probably at the end 
of January), and numerous female strobili were still pink and tender, just 
having passed their ‘receptive stage.” But the trees also possessed many 
full-sized cones, still green in color but already containing ripe seeds. 

* Regarding aecargt of the term “tropical pines” see my paper on “Some taxonomic 
problems of tropical pines,” Proceedings, 13th Congress of the International Union 
of Forest een Organizations, Vienna, 1961. (In press.) 

* Gifford, Ernest M. Jr., and N. T. Mirov. Initiation and ontogeny of the 
ovulate strobilus in ponderosa pine. Forest Sci. 6: 19-25. 60. 

®Mirov, N. T. Photoperiod and flowering of pines. Forest Sci. 2: 328-332. 1956. 


1962 | MIROV, PHENOLOGY OF TROPICAL PINES 219 


Squirrels were busy cutting the cones and eating the fresh seeds. There 
it was evident that the timetable of events leading to the production of 
seeds was somewhat distorted. 

Springtime in Nicaragua is not an upsurge of life as in the North; 
tropical pines never cease to grow. We cannot make the statement that 
in the mountains of Nicaragua it takes two growing seasons, or two 
calendar summers, for seeds to mature. The ovulate strobili continue 
to develop, apparently without much winter slowing, for there is no win- 
ter; and it takes them only a little over one year to mature. That is why 
a Honduran botanist told me once that in his country it takes only one 
year for Pinus oocarpa to produce seed. 

In Indonesia, late in February and early in March of 1961, I observed 
even more distortion in the flowering of pines. The pine there was Pinus 
merkusit De Vriese, moved from the mountains of northern Sumatra 
(about 3° N. Lat.) to the mountains and lowlands of Java (about 6° S. 
Lat.). I am not familiar with the flowering habits of this pine in Sumatra, 
but I suspect, judging from its performance in the mountains of northern 
Thailand, that it sheds pollen in January. 

In the mountains of Java (elev. 4900 ft.) near Bandung, Pinus mer- 
kusit sheds its pollen twice a year: in January-February, and in July— 
August. Better seeds are obtained from the latter pollination, 

A 20-year-old pine plantation at sea level was visited February 25, 
1961. The forest ranger procured phenological records taken for ecvera! 
years. These records showed that Pinus merkusii pollen had been pro- 
duced and dispersed intermittently all year round; the ovulate strobili 
emerging from the buds, as well as the mature cones, were also recorded 
throughout the year. But Pinus kasya Royle, a pine of Burma and Indo- 
china, growing naturally at elevations higher than those of P. merkusii, 
neither produced pollen nor developed ovulate strobili in the plantation. 
Apparently high air humidity is detrimental to the normal seed produc- 
tion of this pine in the humid and hot lowlands of Java. 

These cursory observations suggest that a more comprehensive study of 
the phenology of tropical pines would be interesting and profitable. Both 
Central America and Indonesia are well suited to such a study. In Central 
America there are several institutions in Honduras, Nicaragua and Guate- 
mala where such work could be done. In Sumatra, where the southern- 
most of all pines, Pinus merkusti grows naturally (about 2° south of the 
Equator) travel is at present hazardous; but in Java there are many 
easily accessible pine plantations where phenological records have been 
diligently kept. There a phenological project could be conducted either 
in the world renowned Herbarium Bogoriense, in Bogor, where the Forest 
Research Institute is also located, or in the Division of Biology of the 
Institute of Technology at Bandung 


CABoT FOUNDATION, 
HaArvarD UNIVERSITY 


220 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


ON THE STATUS OF PSILAEA (THYMELAEACEAE) 


Lorin I. NEVLING, JR. 


THE GENUS PsILAFA was described, with a single species P. dalber- 
gioides, by Miquel in 1861! (FI. Ind. Bat. Suppl. 355). The full com- 
bined generic-specific description was based upon a single Teysmann col- 
lection from “Sumatra occid., ad littus prope Siboga.” Nine years later, 
Kurz (Jour. As. Soc. Beng. 39(2): 83) placed Psilaea dalbergioides as a 
synonym of Linostoma pauciflorum Griffith, and, consequently, Psiaea 
was treated as a generic synonym of Linostoma. In 1922, Psilaea was used 
by Hans Hallier (Med. Rijksherb. 44: 28) to typify his section Psiaea of 
Linostoma. The typification of the section, based on Miquel’s taxon, is 
indicated by parenthetical credit i.e., “Sect. Psilaea (Mig.) m.” Un- 
der section Psilaea, Hallier described two new species, L. leucodipterum 
and L. longiflorum (both synonymous with L. pauciflorum 

Following Kurz’s publication, in which he expressed the belief that 
Linostoma pauciflorum and Psilaea dalbergioides were conspecific, it 
appears as though subsequent authors accepted his determination with- 
out re-examining the ee specimen. In my recent revision of 
Linostoma (Jour. Arnold Arb. 42: 295-320. 1961) I placed P. dalber- 
gioides and the names _ based ae it in a category of undetermined 
status. This procedure was followed because the Teysmann type appar- 
ently never had been re-examined taxonomically in the one hundred years 
since Kurz’s publication; Kurz recognized only a single genus (Linostoma) 
where today we recognize two (Linostoma and Enkleia); recent revision 
of the generic limits of the two genera has produced some shifting of 
species from one genus to the other; and, finally, the genus Enkleta is 
known from several collections from Sumatra, but Linostoma is not (ex- 
cluding the oe type), although it is found on the neighboring 
Malay Peninsula 

To resolve my “hesitancy to accept the validity of Kurz’s determination 
of the Teysmann type required first locating and then examining the 
specimen. This proved more difficult than anticipated. Many of Miquel’s 
types (particularly Teysmann specimens) are deposited currently at 
Utrecht. The specimen was not located at Utrecht or at any of the 
other major American or European herbaria. However, from Kurz’s brief 
remarks on Psilaea it is evident that he had seen authentic Teysmann 
specimens during his curatorship (1864-1878) at the Royal Botanic Gar- 
den, Calcutta (now Indian Botanic Garden); and in addition, Teysmann 
Sumatran material is deposited at Calcutta (see M. J. van Steenis-Kruse- 
man, Flora Malesiana 1: 525. 1950). After considerable effort, it was 
established, through Dr. H. Santapau, S.J., that the Teysmann specimen 


' For date of issue see Steenis, C. G. G. J. van, Flora Malesiana 4: ccii. 1954. 


1962 | NEVLING, STATUS OF PSILAEA 221 


was extant, and a black and white photograph of it has been sent to me 
through the courtesy of Mr. Basu, the specimen itself being judged too 
fragile to be loaned. It is unquestionably the holotype (Teysmann s.n.) 
of Psilaea dalbergioides, and, further, is referable to Linostoma pauciflorum 
Griffith. Thus, Kurz’s identification is confirmed and the faith with which 
subsequent authors have accepted his determination is justified. A photo- 
graph of the holotype has been deposited in the herbarium of the Arnold 
Arboretum. 

The full synonymy of Linostoma and of L. pauciflorum is emended 
to read: 


Linostoma Wall. ex Endl. Gen. 331. 1837 (Type: L. decandrum (Roxb.) 
Wall. ex Endl.). 


Linostoma Wall. Cat. n. 4203. 1831, sine descript. 

Linostoma sect. Eulinostoma Meissn. in Mart. Fl. Bras. 5(1): 72. 1855 
(Type: L. decandrum (Roxb.) Wall. ex Endl.). 

Psilaea Miquel, Fl. Ind. Bat. Suppl. 355. 1861 (Type: P. dalbergioides Miq.). 

Linostoma subg. Nectandra Kurz, Jour. As. Soc. Bengal 39(2): 83. 1870 
(Type: L. decandrum (Roxb.) Wall. ex Endl.). 

Linostoma sect. Psilaea Hallier f. Med. Rijksherb. 44: 28. 1922 (Type: 
Psilaea dalbergioides Miq.). 


Linostoma pauciflorum Griffith, Calcutta Jour. Nat. Hist. 4: 234. 1844 
(TypeE: Griffith 4376). 


Psilaea dalbergioides Miquel, Fl. Ind. Bat. Suppl. 355. 1861 (Type: Teys- 
MN SN. 
Linostoma leucodipterum Hallier f. Med. Rijksherb. 44: 28. 1922 (Type: 


Hallier f. B.2 
Linostoma ee eae Hallier f. ibid. 29 (Type: Haviland 1759). 


JosEpH Horace FAULL 


JOURNAL 


OF THE 


ARNOLD ARBORETUM 


VoL. XLII JuLy 1962 NUMBER 3 


JOSEPH HORACE FAULL, 1870-1961 


ANNA F. FAULL 
With portrait * 


JosEpH Horace Fautt was born in L’Anse, Michigan, on May 3, 1870, 
the eldest son of James and Catherine (Bennetts) Faull. He died June 30, 
1961, at his home in Cambridge, Massachusetts, Professor of Forest Pathol- 
ogy, Emeritus, at the Arnold Arboretum, Harvard University. His studies 
of plant pathogens had led him from the Arctic Circle to the Tropics in 
North America, filled his classrooms with students, and won for him world- 
wide recognition as an authority on forest diseases and in particular the 
rust fungi. 

His early life and training can be summarized quickly. Born in the pine 
forests of Michigan, he grew up as a minister’s son on the nothern shore 
of Lake Ontario. Here he often drove long distances through the farm and 
forest lands with his father talking about Charles Darwin whose theories 
on evolution James Faull was inclined to accept. He began to teach before 
finishing high school, interspersing study with teaching until he had earned 
three Normal School certificates and the first degree in Arts from the Uni- 
versity of Toronto, where he had enrolled at Victoria College with the class 
of 1898. From 1898 to 1900, he taught mathematics in Belleville, On- 
tario, at Albert College, where he had completed his preparation for the 
University some years earlier. 

He married Annie Bell Sargent, of Bellwood, Pennsylvania, late in 1903. 
They had met during the summer in Cold Spring Harbor, Long Island, 
New York, where he replaced Albert F. Blakeslee as assistant to Dr. Dun- 
can S. Johnson, of Johns Hopkins University in the summer botany course. 
Miss Sargent, as a candidate for the doctorate, was studying life-histories 
of spiders. She had obtained her A.B. degree from the University of Penn- 
sylvania, as one of the first small group of young women ever to be admitted 
to the regular courses of study at that institution. Together until her death 
in 1953, she shared with him an unabated interest in natural history, as 
well as a great love for students and all children. Of their own three, Cath- 


* This portrait was taken by J. Horace Faull, Jr., in 1949 at the Pathological Labo- 
ratory of the Arnold Arboretum, Jamaica Plain, Boston, Massachusetts. 


224 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


erine Sargent died at the age of sixteen, an invalid following an attack of 
influenza in early infancy; Anna Forward followed her father into botany; 
while J. Horace, Jr., has made a name for himself in chemistry. 

Dr. Faull’s career in botany began in 1900 when he returned to the Uni- 
versity of Toronto to study with Edward Charles Jeffrey, Lecturer in Biol- 
ogy, whose sectioning of lignites (coal), along with the application of Dar- 
win’s theory of evolution to research on the anatomy of woody plants, was 
already attracting attention. Mr. Faull’s problem was the vascular struc- 
ture of the Osmunda ferns. His first paper was published by the Botanical 
Gazette in 1901. More important, Dr. Jeffrey told him about William 
Gilson Farlow and Roland Thaxter at Harvard, the two American pioneers 
in the new field of mycology. 

Mr. Faull went to Harvard in 1901-1903 as an Austin Fellow to begin 
under Professor Thaxter the studies on the cytology of the Ascomycetes 
which were to be the major subject of his published research until 1913. 
His papers on the formation of the ascus and its spores are still standard 
references in this field. In 1904, he received his doctorate from Harvard. 

In 1903, Dr. Faull had already accepted a lectureship in botany at the 
University of Toronto, the same lectureship (under a changed name) left 
vacant by Dr. Jeffrey’s appointment to Harvard. He remained with his 
wife and family in Toronto until 1928, with sabbatical leave in 1909-1910 
for a brief visit to Harvard and a year’s study abroad, mostly with Robert 
Hartig at the Forst Botanische Institut of the University of Munich, Ger- 
many. Botanically he found the trip abroad somewhat disappointing, al- 
though he profited in prestige, in an increased facility with the German 
language, and in a deepened interest in forest pathology. He had been 
made an associate professor at Toronto in 1907. In 1918, this was changed 
to a full professorship while he continued as head of the department which 
he had created. Before he left, his department taught students in the gen- 
eral (liberal) arts, applied science, household science, pharmacy, and 
forestry; his classrooms were crowded; his graduate students and assistant 
staff were increasing in numbers, and a new building for botany alone was 
being proposed in the Provincial Parliament. 

Summers at the University, with an early closing date in April or May, 
were long and, for Professor Faull, varied. He continued his research with 
publications on the Ascomycetes and other fungi, a natural history of the 
Toronto region and a paper on Charles Darwin. He built a large collection 
of lantern slides, photographs, and preserved material for teaching. He 
attended courses in forestry at the College of Forestry, Syracuse Univer- 
sity, New York. He taught in the summer schools at Cornell University, 
Ithaca, New York (1915-17), and at Harvard (1925). He also taught 
his own students in Toronto. He collected in the Adirondacks with G. F. 
Atkinson, in the Allegheny Mountains of Pennsylvania with his wife 
(1913-1917), in the country a few miles or more from his Toronto home, 
and in the forest lands of Ontario and Quebec (Timagami, Algonquin 
Park, the Rideau Lakes, Georgian Bay, the Laurentians). Wherever he 
went he collected fungi and pathological specimens. These impressive 


1962] JOSEPH HORACE FAULL, 1870-1961 225 


collections of teaching, research, and reference material are one of his 
legacies to botany. 

Apart from the University and at the request of the Canadian Govern- 
ment, Dr. Faull established a department of forest pathology with a field 
laboratory in the coniferous forests on Bear Island in the Lake Timagami 
Forest Reservation of Ontario. Commencing in 1918, much of his summer 
time was spent in the development of this laboratory, in long-range experi- 
ments there, and with one of his students, Wade Watson, in the compilation 
of a check list of the Timagami flora. At the same time his services as a 
consultant were in increasing demand by lumber companies and others 
with forest or tree problems. These requests, although remunerative, were 
to him often annoying interruptions to his major interests at the University 
and in the field laboratory. When he left, the Government laboratory and 
department, as well as a shade-tree laboratory, were as firmly established 
as the department at the University, while his students were beginning to 
appear as responsible men and women in botany and forestry in the uni- 
versities, schools, and government laboratories and departments through- 
out the Dominion. He had indeed earned the citation of “Father of Cana- 
dian Botany” given him in 1959 at Montreal in the opening address at the 
IX International Botanical Congress. 

In 1928, he came to Harvard University as Professor of Forest Pathol- 
ogy to take part in the expanding program of research envisioned by 
Charles Sprague Sargent and Oakes Ames for the Arnold Arboretum. For 
Professor Faull this meant freedom from the increasingly heavy teaching 
and administrative load at Toronto and a chance to devote most of his 
time to writing, research, and the advancement of forest pathology. He 
brought with him his collections of research and reference material, his 
graduate students and his reputation as a consultant. The Arboretum 
provided a laboratory and greenhouse built to his specifications on the 
adjacent Bussey Institution grounds; the Farlow Herbarium gave workin 
space there; and, later, after their completion, the Biological Laboratories 
provided additional laboratory rooms, 

Professor Faull’s conception of the new appointment and of forest path- 
ology can best be stated in his own words taken from letters to Edward C. 
Jeffrey and to Oakes Ames at this time. He wrote: “My mind is set on 
advancing Forest Pathology — completing and writing up accumulated 
researches of which there is a good stock in hand, taking up others in 
mind, organizing the position at the Arboretum on a broad and sound 
basis and perhaps working towards a treatise of the Hartig type — these 
are possibilities within reasonable expectation of attainment.”’ Again, “The 
field is a broad one for it involves the study of the etiology of arboreal dis- 
eases and of the principles of their control. As for etiology only a begin- 
ning has been made in America; there exists a host of diseases the causes 
of which are yet unstudied and unknown, and in several instances that 
come to mind wrongly ascribed. Intelligent control is possible only when 
causes are known, but even then there are principles to be worked out. 
This is particularly true of our untamed and abused American forests 


226 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


where the solution seems to lie along prophylactic lines frequently based 
on a knowledge, mostly not yet acquired but acquirable, of the relation of 
the age of the hosts to susceptibility, of the rate of progress of the patho- 
gens and the relative amount of destruction caused. There is likewise a 
phase in connection with the utilization of immense quantities of waste 
diseased living timber. All these aspects constitute a vast field within 
which researches may be undertaken and if you subscribe to these limits 
your project will be as broadly based as the one in taxonomy which has 
been so long and so successfully developed.” 

Scarcely pausing to settle into the new laboratory, Dr. Faull began at 
once the pregram that he had laid out for himself: the completion of 
earlier research, the study of “immediate pathological problems presented 
by trees in the Arboretum” and “of any or all diseases of all kinds of trees 
wherever found and whether in plantations or in the forests.” The latter 
was to lead him far afield from one end of the continent to the other, while 
his days in Cambridge were filled with writing, research, students, and 
visitors from all over the world. One can trace his collecting and travel 
in his accessions books, 

The first summer found him in Timagami in August checking on the 
experiments he had last seen in June just before coming to Harvard; re- 
turning to old locations at Proulx, Quebec; and extending his range into 
the Gaspé. A month later he was in Maine examining nursery spruce and 
white pine for the Brown Company. His first paper appeared in the Jour- 
nal of the Arnold Arboretum in 1929, ““A Fungous Disease of Conifers Re- 
lated to Snow Cover.” It was based on the work at Proulx, Quebec, begun 
some years earlier, and on observations made at the Brown nurseries. 

From 1929 to 1933, he continued this program of revisiting old locations 
and experiments, adding new ones, advising and co-operating in experi- 
ments with the lumber companies and others. He also gave the undergrad- 
uate half-course in pathology agreed upon. In 1929, he made collections 
at the Arnold Arboretum; at the Kelsey nurseries, in Boxford; at Oqussoc, 
in Maine; and again at Proulx, in Quebec, and at Lake Timagami. He 
extended his trips into Nova Scotia at the request of the Provincial forester 
where he notes finding “Milesina polypodophila (Bell) Faull on Abies and 
Polypodium side by side.” He repeated the June trips in the fall and again 
in 1930. He visited the West and the Pacific Coast in 1931, collecting in 
the Yellowstone National Park, in Wyoming; at the North Priest River 
Forest Station, in Washington; in Oregon; in California; in Idaho; and 
in Illinois. He concentrated on the New England mountains in 1932: Ver- 
mont, the Presidential Range, the Berkshires. In 1932-1934, the Arbore- 
tum published the first of his monographs on rusts, that on Milesia. 

From 1933 to 1940, Dr. Faull included a winter trip to the tropics in his 
regular program for the year. In December and January, 1933-34, he 
visited the mountains of Jamaica, Panama, and the Canal Zone. He re- 
turned to Jamaica in February and March, 1935, and again in 1935-36, 
after a three week stop in Cuba. He wrote in the records of the Harvard 
Botanical Garden at “Soledad,” near Cienfuegos, Cuba: “Arrived Dec, 


1962 | JOSEPH HORACE FAULL, 1870-1961 rs 


17, 1935. Departed Jan. 5, 1936. Activities: 1) Made a pathological re- 
connaissance of the Arboretum and of the forest tree plantations at an 
near Soledad. 2) Examined critically a disease of Marabu [Dichrostachys 
cinerea] between Soledad and San Blas, a disease of Ficus in Soledad and 
several heart rot diseases of trees; assembled some relevant material. 3) 
Collected Pucciniastreae in the Trinidad Mts.” The following winter he 
went to Guatemala where, in addition to rusts, he collected material of an 
Abies growing at 10,000 feet which Rehder later described as A. guatema- 
lensis. In November, 1937, he was in Mexico with Professor Maximino 
Martinez collecting in the remote areas of the states of Michoacan, Mexico, 
Morelos, Hidalgo, Veracruz, and Chiapas, as well as in the Federal District. 
He returned to Mexico in 1938 to visit Oaxaca and in 1939 to collect again 
in Chiapas and Hidalgo, as well as in the northern states of Chihuahua and 
Durango. In the meantime, his summer trips continued: the Gaspé, Maine, 
New Brunswick, Long Island, and Chicago, in 1933; Quebec, Mt. Washing- 
ton and Maine, in 1934; Maine and Metis, Quebec, in 1936; Vermont, 
Maine, and Long Island, in 1937; Vermont and Connecticut, in 1938; 
New Hampshire, in 1939; Massachusetts every year. 

In 1938, the Arboretum published the second of his monographs on 
rusts, that on Uredinopsis. Actually, this was the third of his long papers 
on the rust fungi dealing with the taxonomy, morphology, physiology, 
host relations, etc., which he considered not only knowledge of general 
botanical interest but basic for control of plant diseases specifically or in 
principle. In 1926, two years before his Harvard appointment, he had 
presented a long paper on the Puccineastreae at the International Congress 
of Plant Sciences, Ithaca, New York. While not themselves “treatises of 
the Hartig type” contemplated in his letter to Dr. Jeffrey, they undoubtedly 
were working towards it. One more monograph was intended but never 
written, although he had collected the material and begun the necessary 
studies. Like all of his work, these papers are the scholarly presentations 
of careful study and experiment in field and laboratory by a man with a 
capacity for detail combined with breadth of vision and a keen insight. 

Simultaneously with the field work, Dr. Faull organized the laboratory 
at the Arboretum. By September, 1928, the small building was ready with 
its own library “of several thousand pamphlets,” periodicals, and reference 
books; the important collections of diseased plants (about 1000 speci- 
mens); and the usual apparatus for work with cultures and microscope. 
Addition of the experimental greenhouse in 1929 or 1930 completed the 
small laboratory, while more space for students was provided as needed 
at the nearby Bussey Institution, at the Farlow Herbarium in Cambridge, 
and, later, in the new Biological Laboratories there. On July 1, 1929, Dr. 
Faull reported on a reconnaissance of the living collections of the Arbore- 
tum, in addition to five major projects under active investigation by him- 
self, his assistant (G. D. Darker, from Toronto), and the first of his Har- 
vard graduate students: (1) rusts of fir and spruce, (2) Phacidium blight 
and snow cover, (3) lilac disease, (4) needle-cast diseases of conifers, (5) 
browning of white-pine transplants. The first two were a continuation of 


228 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


work begun at Toronto, in the Timagami forests, and at Proulx, Quebec. 
The last two came from field work with lumber companies in Maine or 
from general collecting. The lilac was an Arboretum problem. 

By 1931, the pathological laboratory envisioned by Oakes Ames and Dr. 
Faull and established in fulfilment of “the expressed wish of the late Di- 
rector, Charles Sprague Sargent, and as part of Dr. Sargent’s conception of 
the Arboretum as an institution for the study of woody plants in all fun- 
damental aspects” was in full swing with the functions of the laboratory 
defined “as comprising interest in the Arboretum’s living collections, ex- 
tension services, instruction and research.” With the propagating houses 
next door and the living collections of the Arboretum itself within hand’s 
reach it was ideally located both for his own researches and for friendly 
co-operation with the rest of the Arboretum group. Contact with the under- 
graduate College was maintained through the half-course in plant patho- 
gens, while graduate students, postgraduates, and scholars continued to 
come in twos and threes and sevens. The United States and foreign govern- 
ments from Canada to Jamaica, estate owners, forest interests, and others 
called with increasing frequency for extension services. This, in turn, of- 
fered not only new problems but often supplementary financial support 
for their study and for the postgraduate work which Dr. Faull encouraged 
his students to do both in his own laboratory and abroad. 

For the next ten years the laboratory continued to operate under Dr. 
Faull’s direction with the same efficiency and devotion. The problems 
brought in by field work, collecting trips, extension services, and the Ar- 
boretum itself were numerous and specific. Many of them were quickly 
answered. But many involved much broader botanical issues: “the host 
of unknown and unstudied” causes of plant disease; ‘the principles of 
control yet to be worked out”; the relationships of host, parasite, and en- 
vironment; morphology; anatomy; physiology; and even some aspects of 
genetics — in short, the entire range of botanical science. At least twenty- 
two such studies were carried through to the publication of significant con- 
clusions, a broadly based beginning in the “vast field” of unstudied ar- 
boreal pathology for which the laboratory had been organized. Of most 
popular interest is the Dutch elm disease, which Dr. Christina Buisman, of 
Holland, found for the first time in the United States at the end of a year’s 
work (1929-30) on American elm-diseases. Early hope of eradication 
faded when some years after its elimination from Ohio it reappeared in 
New Jersey; thereafter, the Arboretum effort was directed at control. 
Equally or more important both botanically and practically were other 
studies: a disease of Fagus, lilac blight (a graft incompatibility leading to 
work on viruses), taxonomy of Ganoderma (initially a question of classi- 
fication), morphology of rust spermogonia as taxonomic criteria, my- 
corrhiza (morphology and physiology involving mineral nutrition of pine), 
wilt diseases of elm (other than Dutch), etc. The record of the laboratory 
can be traced in the annual reports and papers published in the Arbore- 
tum’s Journal or as one of its Contributions, a series inaugurated by two 
papers from the pathology laboratory: Dr. Darker’s Ph.D. thesis on 


1962] JOSEPH HORACE FAULL, 1870-1961 229 


needle-cast fungi and Professor Faull’s monograph on “Milesia.” That the 
laboratory was not continued and his position remained unfilled after his 
retirement was a disappointment to him. 

Professor Faull retired in 1940, a world figure in forest pathology and 
the world’s authority on rust fungi associated with ferns. His students 
have gone on to make names for themselves in the botanical world and to 
serve in responsible teaching, administrative, and research positions both 
in the United States and Canada. The extension services have been taken 
over mostly by government agencies, but the broad and comprehensive 
kind of research begun at the Arboretum seems not to have been continued 
in this country. 

For several years, Dr. Faull himself remained in his laboratory, assisting 
with Arboretum affairs, helping in the fight against Dutch elm disease, 
serving as an associate editor of the Journal, organizing his collections, ad- 
vising at the Farlow Herbarium, and welcoming the scientists who came 
to see him. The visits of two of these gave him particular satisfaction. 
One was that of Professor Maximino Martinez, of Mexico, who had ar- 
ranged for his earlier collecting in the Mexican forests and had accompanied 
him on some of the trips. The other was Dr. Krishnadas Bagchee, from the 
Forest Service in India, who shared Dr. Faull’s love of the forests and his 
interest in fungi. Dr. Bagchee brought with him a collection of Indian 
rusts to my father’s delight. In 1956, the Journal of the Arnold Arboretum 
published his last paper. Shortly thereafter, Dr. Faull gave up his last 
small room in the Cambridge laboratories. 

The proper disposal of his large collections of pathological material occu- 
pied much of his time in these later years. Some, of course, had been left 
at the University in Toronto. But the thousand specimens brought with 
him to Harvard had grown through exchange, communication, and his own 
collecting trips and field work and that of his students and associates to 
nearly fourteen thousand. The small room where the earlier collections 
had at first been kept had become inadequate long before his retirement. 
In 1939, he had placed a thousand duplicates of wood destroying fungi in 
the Farlow Herbarium and three hundred in the Bureau of Plant Industry 
in Washington (later transferred to the National Herbarium). In 1940, 
he noted in his report the deposition of his large collection of polypores at 
the Farlow Herbarium. Finally before his death, he placed a collection of 
fern rusts (type specimens and material documenting his publications) in 
the U.S. National Herbarium, where he hoped it would be both adequately 
cared for and available for reference and future investigations. The large 
remainder of the collections with duplicates has been placed in temporary 
storage at the Farlow Herbarium and the Arnold Arboretum awaiting more 
permanent quarters. A few are still at his home in Cambridge. 

As his health failed, Dr. Faull spent more and more of his time with his 
family, partly in Texas, but most of it at their home in Cambridge. Here 
he enjoyed his garden, his neighbors, a little carpentry, a great deal of 
reading, a little cribbage, and a voluminous correspondence with friends 
and students dating back to those whom he had taught as little children in 


230 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


the Canadian rural schools. As a student at Toronto, he had been the 
gold medalist in his graduating class of 1898. Now he reread the entire 
works of Dickens and Shakespeare, along with his other books. As a young 
man, too, he had played ice hockey on the Victoria College team. Now he 
listened to the sports broadcasts on the radio. He never lost his interest in 
the schools or in the education of women or his sympathy for the under- 
privileged. At his death, a volume of Dickens was near his chair by the 
radio along with a copy of the Cambridge School Report, a box of fern 
rusts near his desk. 

As a teacher and a scientist, Joseph Horace Faull enriched the two coun- 
tries in which he lived. His students continue to do so. 


Professor Faull had been a member of the American Academy of Arts 
and Sciences, the American Association for the Advancement of Science, 
the American Phytopathological Society, the American Society of Natural- 
ists, the Mycological Society of America, the National Agricultural Chemi- 
cals Association, the New England Botanical Club, the Royal Canadian 
Institute, the Royal Society of Canada, the Society of American Foresters, 
the Society of the Sigma Xi, and the Sociedad Botanica de México. 


72 FRESH Ponp LANE, 
CAMBRIDGE, MASSACHUSETTS 


BIBLIOGRAPHY * 


— 1901 — 
The anatomy of the Osmundaceae. Bot. Gaz. 32: 381-420. pl. 14-17, 


— 1905 — 
Development of ascus and spore formation in Ascomycetes. Proc. Boston Soc. 
Nat. Hist. 32: 77-113. pl. 7-11, 
Diseases of timber. Canad. Forestry Jour. 1: 105-108, 3 pl. 


— 1906 — 

Further studies on ascus. (Report of paper read at the annual meeting of the 
Central Botanists held at Ann Arbor, Michigan, December 28, 29, 1905. 
Abstracted by B. M. Davis.) Science 23: 134. 

A preliminary note on ascus and spore formation in the Laboulbeniaceae. Sci- 
ence 25: 157, 153, 


— 1907 — 
Bunt, or the stinking smut of wheat. Canada Dep. Agr. Seed Branch Bull. $3. 
13 pp. 


— 1908 — 
Arceuthobium pusillum Peck. Ottawa Nat, 21: 175. 
Notes on Rondeau Park. Ontario Nat. Sci. Bull. 4: 99-103. 


* Compiled by Lazella Schwarten; with supplementary references added by the 
author from Professor Faull’s own files and from those of the Farlow Herbarium. 


1962] JOSEPH HORACE FAULL, 1870-1961 231 


10002 
The influence of Darwin on botanical science. Ontario Nat. Sci. Bull. 5: 31-37. 
Stele of Osmunda cinnamomea. Trans. Roy. Canad. Inst. 8: 515-534. pl. 4-6. 


— 1911 — 
The cytology of the Laboulbeniales. Ann. Bot. 25: 649-654. 


— 1912 — 
The cytology of Laboulbenia chaetophora and L. gyrinidarum, Ann. Bot. 26: 
325-355. 


— 1913 — 
The natural history of the Toronto region, Ontario, Canada. 1-419. Toronto. 


— 1914 — 
Bark disease of the chestnut in British Columbia. (With G. H. Graham.) 
Forestry Quart. 12: 201-203. 


— 1916 — 
Chondromyces thaxteri, a new myxobacterium. Bot. Gaz. 62: 226-232. Peed, 0. 
Fomes officinalis (Vill.) —a timber destroying fungus. Forestry Quart. 14: 
737-739 
Fomes officinalis (Vill.), a timber destroying fungus. Trans. Roy. Canad. Inst. 
Toronto 11: 185-209. pl. 18-25. 


— 1918 — 
The menace to our white pine. Canad. Forestry Jour. 14: 1685-1687. 
The fight to save our white pine. /bid. 1743-1747. 


— 1919 — 
Forest pathology. Rep. Minister Lands, Forests & Mines. Prov. Ontario 1919: 
11 25. 


Pineapple fungus or enfant de pin or wabadou. Mycologia 11: 267-272. 


— 1920 — 
Forest pathology. Rep. Minister Lands, Forests & Mines. Prov. Ontario 1920: 
224-235. 


— 1921 — 
Plant pathology; its status and its outlook. (Presidential address.) Trans. Roy. 
Soc. Canada III. 14(sect. v): 1-16. 


— 192 
Records for four years on the needle blight of Pinus strobus. Phytopathology 
1] . 


Some problems of forest pathology in Ontario. Needle blight of white pine. 
Jour. Forestry 20: 67-70. 
— 1923 — 
Balsam rusts. Rep. Minister Lands & Forests. Prov. Ontario 1923: 253, 254. 


252 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


— 1924 — 
The aecial stage of Hyalopsora psi (Peck) P. Magnus. (With G. D. 
Darker.) Phytopathology 14: 3 
Stereum sanguinolentum as the cause of ‘“‘sapin rouge” or red heart rot of 
bal (With Irene Mounce.) /bid. 349. 
The treatment of decayed wood in and outside the mill. Pulp & Paper Mag. 
3pp. Feb. 28. 


— 1928 — 
Living cells in heart-wood of trees. Science 67: 296, 297. 


— 1929 — 
A a disease of conifers related to the snow cover. Jour. Arnold Arb. 10: 


The ina biology and ae of the Pucciniastreae. Proc. Int. Congr. 
Pl. Sci. 1926. 2: 1735-17 


— 1930 — 

Blister rust in Nova Scotia. Blister Rust News 14: 63. 

The health of the forests. Illus. Canad. Forest & Outdoors 26: 146-149. (March 
1930. 

Notes on forest diseases in Nova Scotia. Jour. Arnold Arb. 11: 55-58. 

Some general remarks regarding forest pathology in relation to forestry and 
notes on forest diseases in Nova Scotia. Rep. Dep. Lands & Forests Nova 
Scotia 1930; 33-40. 

The spread and the control - the Phacidium blight in spruce plantations. Jour. 
Arnold Arb. 11: 136-14 


— 1931 — 
Milesina rusts on Aspidium braunii Spenner. Jour. Arnold Arb. 12: 218,719; 


— 1932 — 
“Hubert, E. E., An outline of forest pathology.” (Review.) Phytopathology 
22: 393-395, 
Taxonomy and a ar distribution of the genus Milesia. Contr. Arnold 
Arb, 2: 1-138. pl. 1 
231984 <3 


Arthur Bliss Seymour (1859-1933). Proc. Am. Acad. Sci. 69: 543, 544. 

The biology of Milesian rusts. Jour. Arnold Arb. 15: 50-86. pl. 84-86, 

Blister rust of Pinus longifolia Roxb. Ibid. 154-157. 

A remarkable spruce rust, Peridermium parksianum n. sp. Ibid. 86, 87. 

Winter hardiness of trees and shrubs growing in the Arnold Achavetuni, (With 

. G. Jack, W. H. Judd, and L. V. Schmitt.) Arnold Arb. Bull. Pop. Inf. 

IV, 2: 29-47, 53-60. 

Wehmyer’s “The genus oo Nitschke and its segregates.” (Review.) 
Jour. Arnold Arb. 15: 157— 


— 1935 — 
Can we eradicate the Dutch elm disease? Address before the Annual Meeting 


of the Massachusetts Forest and Park Association, January 31, 1935. 4 pp., 
illustr, Mass. Forest and Park Assoc., Boston 


1962] JOSEPH HORACE FAULL, 1870-1961 233 


— 1936 — 
Pathological studies on beech at the Arnold Arboretum. Proc. Natl. Shade Tree 
Conf. 12: 21-29 
Two spruce-infecting rusts, Chrysomyxa piperiana and Chrysomyxa chiogenis. 
Jour. Arnold Arb. 17: 109-114 
The viewpoint of the Arnold Arboretum on the Dutch elm disease. Arnold 
rb. Bull. Pop. Inf. IV. 4: 15-20. 1 pl 


— 1937 — 
Chrysomyxa empetri—a spruce-infecting rust. Jour. Arnold Arb. 18: 141-148. 
pl. 202, 203. 


New England i in autumn array. Yankee 3(10): 26, 27. 


— 1938 — 
The biology of rusts of the genus Uredinopsis. Jour. Arnold Arb. 19: 402— 436. 
The Dutch elm disease situation in the United ee at the close of 1938. 
rnold Arb. Bull. Pop. Inf. IV. 6: 75-78. pl. 7 
Pucciniasirem on Epilobium and Abies. Jour. Rerold Arb. 19: 163-173. 
my and geographical wee of the genus Uredinopsis. Contr. Ar- 
ane Arb. 11: 1-120. pl. 1 
— 1939 — 

A review and extension of our knowledge of Calyptospora goeppertiana Kuehn. 
Jour. Arnold Arb. 20: 104-113. 

— 1942 — 

Report of Northeastern Committee on Dutch elm disease. (With C. C. Hamil- 
ton, D. S. Welch, H. H. York, and J. S. Boyce.) 4 pp. Mass. Forest and 
Park Assoc., Boston. 

— 1947 — 
Tropical fern hosts of rust fungi. Jour. Arnold Arb. 28: 309-319. 


— 1956 — 
A rust on Woodwardia fimbriata. Jour. Arnold Arb. 37: 314-316. 


234 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


COMPARATIVE ANATOMY OF THE LEAF-BEARING 
CACTACEAE, V 


THE SECONDARY PHLOEM 
Lauit M. Srivastava! ano I. W. BarLey 2 


IN AN EARLIER PAPER (Bailey and Srivastava, 1962), it was indicated 
that the vascular cambium, tracheary tissue, and sieve elements in the leaf- 
bearing Cactaceae show structural features that suggest an advanced level 
of evolutionary specialization. The arrangement of fusiform initials in the 
cambium and their derivatives in xylem and phloem, and the structure of 
xylem tissue were described in detail in this paper. In contrast, the account 
of phloem was cursory and restricted mainly to the form of sieve elements 
and their sieve plates and sieve areas. In order to understand the structure 
of phloem tissue, particularly the relationship between sieve elements and 
parenchyma cells associated with them, an ontogenetic study of the tissue 
was considered essential. This study forms the subject of the present paper. 

Three species of leaf-bearing Cactaceae, Pereskia sacharosa Griseb, | Tu- 
cuman ], Pereskiopsis aff. chapistle Britt. & Rose [Boke B-3], and Quiaben- 
tia aff. chacoensis Backbg. [Tucuman] were selected for detailed onto- 
genetic work. The material was killed in FAA. Small pieces from the stems 
of these species were embedded in paraffin and sectioned on a rotary micro- 
tome. Some additional material was embedded in celloidin and sectioned 
on a sliding microtome. Serial transverse, radial, and tangential sections 
were obtained and, later, stained by a combination of tannic se iron 
chloride, and lacmoid as described by Cheadle, Gifford & Esau (19 

In order to determine the origin of phloem elements, several te (that 
is, radial files of derivatives of single cambial initials) were studied in de- 
tail for each of the three species. Tiers were drawn from serial cross sec- 
tions with the use of a camera lucida attachment. The cells in each tier 
were reconstructed from these drawings. The entire height of a tier was 
examined in order to determine the exact relationship between sieve ele- 
ments and the parenchymatous cells associated with them. The results 
obtained from a study of cross sections were later substantiated by a study 
of tiers from serial tangential sections. Radial sections were only of lim- 
ited use in the present study, partly because of the difficulty of obtaining 
good radial sections and partly because of the irregular planes of divisions 
in the phloic initials and the small size of some of the phloem derivatives. 

As is well known, sieve elements and the associated companion cells 

* Mercer Fellow of the Arnold Arboretum. 

* This investigation was financed in part by a grant from the National Science 
Foundation. I am indebted to the American Philosophical eres for the loan of a 
Wild microscope. 


1962] SRIVASTAVA & BAILEY, CACTACEAE, V 235 


usually function for a limited period only. They die in old phloem and, in 
most cases, are eventually crushed by the expanding parenchyma cells and 
the pressures resulting from new growth (cf. Esau, 1953, pp. 299, 300). As 
a result, the arrangement of phloem derivatives, typical of young phloem, is 
distorted and the study of tiers cannot be pursued with accuracy in very 
old phloem. The tiers drawn in the present study, therefore, represent the 
functional phloem and only as much of nonfunctional phloem * as had not 
yet been distorted. 

The terms used here are common in literature on phloem (cf. Esau, 
1950, 1953; Esau & Cheadle, 1955; Cheadle & Esau, 1958), but some of 
them are explained again for clarity. A phloic initial is the daughter cell 
formed towards the phloem after a periclinal division in the cambial in- 
itial. The phloic initials either directly, that is, without any further 
divisions, or after a few divisions produce the various cell types in the 
phloem. For instance, a phloem-parenchyma strand is formed after one 
or more horizontal divisions in the phloic initial. The term precursor 
refers to a cell that would either differentiate directly as a definitive 
phloem element or in which further divisions would occur; in any case, 
it denotes an undifferentiated cell in which further changes are going to 
occur. Thus a phloic initial may behave as a precursor of a fusiform 
phloem-parenchyma cell, or one of the daughter cells after a division in 
the phloic initial may behave as a precursor of the sieve-tube mother cell 
and a parenchyma cell. The term sieve-tube mother cell refers to a cell 
in which one or more divisions occur and within the confines of which 
a sieve element and its companion cells are formed. Occasionally a sieve- 
tube mother cell may differentiate directly as a sieve element and no 
companion cells may be formed. The terms sieve element and sieve-tube 
member are used interchangeably. Various kinds of parenchyma cells 
occur in phloem. In the present paper, the terms used to describe them 
have an ontogenetic implication. Thus, a fusiform phloem-parenchyma 
cell is derived directly from a phloic initial; and a phloem-parenchyma 
strand is formed after one or more horizonta! divisions in the phloic 
initial. Some other parenchyma cells and companion cells are formed in 
association with a sieve element after divisions in a single phloic initial. 
For convenience of description, these cells ontogenetically related to the 
sieve elements are often referred to as parenchymatous cells or elements. 


ORIGIN OF PHLOEM ELEMENTS IN PERESKIA 
Analysis of Tiers 
A cross section through the phloem of Pereskia sacharosa is shown in 
Fic. 32, in which the cell types present in the tissue and their general 
arrangement are seen. Another cross section is drawn in Fic. 1, but the 
individual tiers are separated tangentially for a better illustration of 


° Nonfunctional, in contrast to functional, phloem refers to that part of phloem in 
which sieve elements are no longer conducting (cf. Esau, 1953, p. 299 


236 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLIII 


Fic, 1. Pereskia sacharosa, transection of cambial zone (below) and phloem, 
< 495. Tiers separated from one another to show radial extent of each; sieve 
elements unmarked; companion cells stippled; parenchyma cells with nuclei; 
ray cells on extreme right without nuclei. Tiers A-E analyzed in detail in Fics. 
2-6 and in text. 


their radial extent. A study of similar drawings from serial cross sections 
reveals the axial extent of each tier and the cell types that compose it. 
Tiers A, B, C, D, and E in Fic. 1 are analyzed in detail. The cross sec- 
tion drawn in Fic. 1 did not pass through the middle (in terms of axial 
extent) of all the five tiers selected for study. The cross sectional views 
at different levels in the axial extent of these tiers are illustrated in Fics. 
2-6, a—e; the levels at which these cross sections were taken are indicated 
along the margins of Fics. 2—6, f. In Fics. 2-6, a-e, sieve elements are un- 
marked except for sieve plates which, when present, are indicated by 
hatched areas; companion cells are stippled; and parenchyma cells onto- 
genetically associated with sieve elements are shown with nuclei. Fusi- 


1962 | SRIVASTAVA & BAILEY, CACTACEAE, V 237 


form phloem-parenchyma cells and phloem-parenchyma strands are shown 
with a circle drawn with a heavy pen. Cells internal to the cell complex 
in which sieve element 1 occurs are not considered — they are assumed 
to be undifferentiated cells. Some of them may have differentiated as 
fusiform phloem-parenchyma cells or phloem-parenchyma strands, but 
we were not certain about this. They are shown without nuclei. FIGURES 
2-6, f, represent schematically the axial extent of the sieve elements and 
the parenchymatous elements ontogenetically associated with them in 
tiers A-E; the sieve elements are drawn with numbered solid lines, the 
companion cells with dotted lines, and parenchyma cells with broken 
lines. Lines representing companion cells and parenchyma cells are placed 
arbitrarily on the left and right, respectively, of the lines representing 
sieve elements. If a strand of companion or parenchyma cells is formed 
in association with a sieve element, it is represented by a single line; but 
the individual cells in the strand are demarcated by short oblique lines 
intersecting the vertical line that represents the strand. All cells that 
are ontogenetically related and derived from a single phloic initial are 
included within pairs of horizontal lines drawn at the upper and lower 
limits of the complex of cells. The numbers 0-225 or —300 at left in 
each drawing represent the length in microns. The purpose of this 
schematic representation is twofold. First, it shows at a glance how many 
and what kinds of cells are ontogenetically related with one another. 
Second, the combined length of the cells in a complex of cells, barring 
the overlap of cells, reflects the length of the phloic initial (and, hence, 
that of the fusiform cambial initial). The lengths of different phloic 
initials in a tier represent approximately the axial extent, or the height, 
of the tier. Fusiform phloem-parenchyma cells and phloem-parenchyma 
strands are not considered in Fics. 2-6, f. 


Tier A: Seven sieve elements, indicated by arabic numerals, are pres- 
ent in tier A (Fic. 2, c, f). Sieve element 1 is associated with two com- 
panion cells and one parenchyma cell, sieve element 2 with one com- 
panion and one parenchyma cell, sieve element 3 with three companion 
and two parenchyma cells, sieve element 4 with two companion and two 
parenchyma cells, sieve element 5 with one companion and one paren- 
chyma cell, and sieve elements 6 and 7 with one companion cell each. 
Two fusiform phloem-parenchyma cells occur in the tier and were de- 
rived directly from their phloic initials. 

Each of the sieve elements 1-5 with their associated parenchymatous 
cells and the sieve elements 6 and 7 with their companion cells originated 
by divisions within the confines of a single phloic initial. The origin of 
these elements is considered in detail. 

The phloic initial within the confines of which sieve element 1 is 
present divided obliquely and longitudinally. Of the two daughter cells 
formed, the one towards cambium was the precursor of a parenchyma 
cell, the other away from the cambium was the mother cell for sieve 
element 1 and its companion cells. In this mother cell a longitudinal, 


238 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


é' 
es ee ee 
a es es nt oom ba 
ae: oe ee | ee ee 
b ca eee Gee | ee 
“| {1 "| a yy : 
c 1 ae ee ee oe ee 
| no | real ; 
d at ae r an i | >a 
e iT | bho pe 
| | | 
= \" 
I 
oO —L f 


Fic. 2. Pereskia sacharosa, Tier A of Fic. 1: a-e, cross sections of tier at 
levels indicated in schematic representation of certain members of tier (f) by 
arrowheads opposite a—a, b—b, etc., respectively; a-e, X 495. 

For details of drawings in Fic. 2 and in Fics. 3-6, 10-14, 21-25, see text, p. 
280; 


more or less radial division separated a narrow cell from a larger cell. 
The narrow cell divided horizontally and formed a strand of two cells that 
differentiated as companion cells; the larger cell matured as sieve ele- 
ment 

The sequence of divisions in the phloic initials responsible for sieve ele- 
ments 2 and 5 and their parenchymatous cells was similar to that in the 


1962] SRIVASTAVA & BAILEY, CACTACEAE, V 239 


phloic initial for sieve element 1—the first oblique longitudinal wall 
separated the precursor of a parenchyma cell from the sieve-tube mother 
cell, and a subsequent division in the sieve-tube mother cell produced a 
small and a large cell; the small cell differentiated as a companion cell, 
the large cell as a sieve-tube member. 

The order of divisions in the phloic initials within the confines of 
which sieve elements 3 and 4 occur is more complicated. The cell com- 
plex with sieve element 3 is analyzed as follows. A more or less tangential 
division extending through most of the length of the phloic initial re- 
sulted in two daughter cells, one away from the cambium was the pre- 
cursor of a parenchyma cell (Fic. 2, a—e), the other towards the cambium 
was the precursor of a second parenchyma cell and the mother cell of 
sieve element 3. An oblique longitudinal division in the lower half of 
this latter precursor formed two daughter cells: one was the precursor 
of the second parenchyma cell (Fic. 2, d-e), the other, a larger cell, was 
the sieve-tube mother cell for element 3. A longitudinal division along 
the right radial and outer tangential walls of the sieve-tube mother cell 
produced a narrow precursor that divided horizontally to form a‘ strand 
of two companion cells (Fic. 2, a-e); another longitudinal division along 
the inner tangential wall, but confined to the upper half of the sieve- 
tube mother cell, produced a third companion cell (Fic. 2, a-c). The 
larger cell left after these divisions in the sieve-tube mother cell differ- 
entiated as sieve element 3. The following sequence of divisions is 
visualized for the origin of sieve element 4. Two successive oblique 
longitudinal divisions in the phloic initial, one along left radial and the 
other along right radial wall, separated two precursors of parenchyma cells 
from the sieve-tube mother cell. A more or less radial longitudinal di- 
vision in the sieve-tube mother cell produced a narrow cell, which di- 
vided horizontally and formed two companion cells (Fics. 2, a—-d), and 
a larger cell that differentiated as sieve element 4. 

Sieve elements 6 and 7 are ontogenetically related. A more or less radial 
longitudinal division in the phloic initial separated two sieve-tube mother 
cells, each of which divided subsequently and formed a sieve-tube element 
and a companion cell. 


Tier B: Twelve sieve elements (indicated by arabic numerals) with 
their associated parenchymatous cells and one fusiform phloem-paren- 
chyma cell occur in tier B (Fic. 3, c, f). The divisions in phloic initials 
responsible for sieve elements 3, 7, and 12 are least complicated and, 
hence, are analyzed first. A tangential longitudinal wall laid down in the 
phloic initial of sieve element 3 separated two daughter cells, one (to- 
wards cambium) was the precursor of a parenchyma strand, the other 
(away from cambium) was the sieve-tube mother cell. A radio-longi- 
tudinal wall divided the sieve-tube mother cell into a narrow precursor, 


visions in the phloic initials of sieve elements 7 and 12 is similar except 


240 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLIII 


that the precursor of companion cells did not divide horizontally. In 
the cell complex including sieve element 12, the precursor of the paren- 
chyma cell also did not divide horizontally to form a strand of paren- 
chyma cells (Fic. 3, f). 


2257 | 2 3 4 8 6 7 89 10 I 2 
ze 7 = a 
| Io: : ee a ae oe | 

o4 oat Pfr port tbe 

Bea Ee) a de ee 

Oe fees eed ed dL 

: : a is 1: a 

d eV a) a Sign lS) oa ps 

“| at GV ey ah 

e | a aT bof) i} sft be 
ees | es eens) aes 

re) et mes ee 4 —_J 


f 


Fic. 3. Pereskia sacharosa, TIER B of Fic. 1: a-e, cross sections of tier at 
levels indicated in schematic representation (f), as in Fic. 2; a-e, & 495. Oblique 
line connecting sieve elements 8 and 9 in diagram (f) shows that the two 
originated after division in a single mother cell. For details of drawings, see 
text 


1962] SRIVASTAVA & BAILEY, CACTACEAE, V 241 


Sieve elements 1 and 2; 4, 5, and 6; 8 and 9; and 10 and 11 are onto- 
genetically related and were formed with their associated parenchymatous 
cells after divisions in single phloic initials. One parenchyma cell occurs 
in common association each with sieve elements 1 and 2, 8 and 9, and 10 
and 11. Sieve elements 4, 5, and 6 do not seem to have any parenchyma 
cell. The sequence of divisions in the phloic initials in all these in- 
stances is more complicated than in the case of sieve elements 3, 7, and 
12. It appears that in the phloic initial responsible for the cell complex 
in which sieve elements 1 and 2 occur, an oblique anticlinal wall was 
laid down. Of the two resulting daughter cells, one towards the cambium, 
the precursor of a parenchyma cell and the mother cell of sieve element 
1, divided obliquely, but the dividing wall did not extend to the cell 
tips (Fic. 3, f). The longer cell formed after this division was the 
precursor that matured as the parenchyma cell; the shorter was the 
sieve-tube mother cell for element 1. In this sieve-tube mother cell an 
oblique wall was laid down that separated a narrow precursor (left and 
above), which divided horizontally and formed two companion cells, 
from a larger cell that matured as sieve element 1. The cell away from 
the cambium, formed after the first anticlinal division in the phloic initial, 
was the mother cell for sieve element 2 and its companion cell. In this 
mother cell a more or less radial division separated the precursor of a 
companion cell from a precursor that matured as sieve element 2. In the 
interpretation of the origin of this cell complex, it has been assumed 
that the parenchyma cell was more closely related to sieve element 1 than 
to sieve element 2. It is possible, however, that the first division in the 
phloic initial of this cell complex was not the one that separated the 
sieve-tube mother cell of element 2 from the precursor of the mother cell 
of element 1 and the parenchyma cell, but it was one that separated the 
parenchyma cell from a precursor in which later divisions formed sieve 
elements 1 and 2. In the latter interpretation the parenchyma cell would 
be equally related to the two sieve elements. 

In the analysis of the cell complex in which sieve elements 4, 5, and 6 
occur, it appears that a more or less tangential longitudinal division of 
the phloic initial resulted in two daughter cells. The daughter cell to- 
wards the cambium was the sieve-tube mother cell of element 4; a radial 
longitudinal division in this mother cell followed by two transverse di- 
visions in the narrower of the two cells resulted in three companion cells 
and the sieve-tube member 4. In the daughter cell away from the 
cambium an oblique division extending through the upper half of the 
cell set off the sieve-tube mother cells of elements 5 and 6. The mother 
cell of element 6 was shorter than that of element 5 (cf. Fic. 3, a-f). A 
longitudinal division in the mother cell of element 5 produced a long, 
narrow companion cell and the sieve-tube member 5. Two successive 
divisions occurred in the mother cell of element 6 and formed two pre- 
cursors of companion cells and a precursor that matured as sieve element 
6. In one of the precursors of companion cells a horizontal division re- 
sulted in two companion cells; the other precursor matured directly as 


242 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLII 


a companion cell. As a result, sieve element 6 is associated with three 
companion cells. 

In the phloic initial that formed the cell complex including sieve ele- 
ments 8 and 9, an oblique radio-longitudinal division formed two daughter 
cells. One was the precursor of a parenchyma cell; the other divided 
obliquely near its lower end and produced two mother cells, one each 
for sieve elements 8 (Fic. 3, a-d) and 9 (Fic. 3, d-e). An oblique di- 
vision in the mother cell of element 8 formed a narrow cell that differ- 
entiated as a companion cell and a larger cell that matured as the sieve 
element 8. The precursor of sieve element 9 differentiated as a sieve-tube 
member without any divisions. The sequence of divisions in the phloic 
initial within whose confines elements 10 and 11 appear is similar to 
that in the phloic initial of elements 1 and 2, 


Tier C: Nine sieve elements with their associated parenchymatous 
cells and one fusiform phloem-parenchyma cell occur in tier C (Fic. 4, 
c, f). The origin of the cell complexes including sieve elements 5 and 
9 is simple and followed the pattern described for sieve elements 1, 2, 
and 5$ in tier A, and 3, 7, and 12 in tier B. A strand of four companion 
cells was formed in association with sieve element 5S. 

Sieve elements 1-4 and 6-8 with their associated parenchymatous cells 
were derived from single phloic initials. A new feature is seen in the 
origin of sieve elements 1-4. A more or less transverse division in the 
phloic initial formed two daughter cells. An oblique, more or less tan- 
gential division in the upper daughter cell formed two precursors, the one 
away from the cambium matured without any further divisions as sieve 
element 2, the other towards the cambium divided radially and formed a 
parenchyma cell and sieve element 1 (Fic. 4, a~-b). No companion cells 
were formed in association with either sieve element 1 or 2. In the lower 
daughter cell (formed after the first, transverse division in the phloic 
initial) an oblique longitudinal division separated the mother cell of 
sieve element 4 from the precursor of the mother cell of sieve element 
3 and a parenchyma cell (Fic. 4, c-e). An oblique division in the mother 
cell of element 4 separated a narrow cell that differentiated as a com- 
panion cell (lower right corner) and a larger cell that matured as the 
sieve element 4. In the precursor of the mother cell of sieve element 3 
and the parenchyma cell, a tangential division produced two daughter 
cells: one towards the cambium was the parenchyma cell, the other away 
from the cambium was the sieve-tube mother cell of element 3. In this 
mother cell a tangential longitudinal division separated a precursor of two 
companion cells from a precursor that differentiated as the sieve element 
3 (Fic. 4, c-e). In the origin of sieve elements 6—8, two successive 
oblique divisions, one somewhat anticlinal and separating the phloic 
initial into an upper and a lower cell (cf. Fic. 4, a-e), the other some- 
what tangential and oblique and separating the lower cell into an outer 
and an inner daughter cell (Fic. 4, d-e), resulted in three mother cells, 
one each for elements 6, 7, and 8. The mother cell of element 6 was the 


1962] SRIVASTAVA & BAILEY, CACTACEAE, V 243 


TTL EPR 


a 
b 
c 
d 
Oc. 
mca: OC oz 
| 23 4 § 6 7 8 9 
210 2 =] Sa ae ee 
| ; | 
| aoe | 
a | qm | Pa 
| a on 
os 
| a a aL 
‘ | |! a e 
Pg day int 
d ah <I! | : 
Si alert a 
; : if E 


f 


Fic. 4. Pereskia sacharosa, T1ER C of Fic. 1: a-e, cross sections of tier at 
levels indicated in schematic representation (f), as in Fic. 2; a-e, X 495. Details 
as given in text. 


longest and extended from the top end of the phloic initial to well below 
its middle, that of element 7 was in the lower half of the phloic initial but 
did not extend to its lowermost tip, and that of element 8 extended 
through lower one-third of the phloic initial (Fic. 4, f). Two successive 
longitudinal divisions along the outer tangential wall of the mother cell 
of element 6 produced two precursors of companion cells and a precursor 
that matured as sieve element 6. In one of the precursors of companion 


244 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


cells a horizontal division yielded two cells. As a result, three com- 
panion cells were formed in association with sieve element 6. In the 
mother cell of element 7 a longitudinal division along the outer tangential 
wall produced a narrow precursor of two companion cells and a large 
cell that matured as sieve element 7. In the mother cell of element 8 
two longitudinal divisions, one along the radial wall on left and the 
other along the outer tangential wall, produced two precursors of com- 
panion cells and a precursor that matured as sieve element 8. The pre- 
cursor of companion cells along the outer tangential wall divided hori- 
zontally, Eventually three companion cells were formed. 

That the sieve elements 1—4 originated by divisions in a single phloic 
initial is shown clearly in Fic. 4, f, where the combined lengths of the 
parenchyma cell associated with sieve element 1 and the sieve element 3 
(which are the longest derivatives of the upper and lower daughter cells 
formed after the first horizontal division in the phloic initial) approxi- 
mately equal the length of other phloic initials in the tier. Likewise, the 
combined lengths of elements 6, 7, and 8, barring the overlap, is the same 
as that of the phloic initials responsible for the complexes of cells in- 
cluding elements 5 and 9. 


Tier D: Six sieve elements with their associated parenchymatous cells 
and one fusiform phloem-parenchyma cell occur in tier D (Fic. 5, c, f). 
Sieve elements 3 and 6 and their associated companion and parenchyma 
cells originated in the usual manner described for sieve elements 1, 2, and 
5 in tier A; 3, 7, and 12 in tier B; and 5, and 9 in tier C, and are not 
considered. Sieve elements 1 and 2, and 4 and 5 are ontogenetically re- 
lated. In the phloic initial within the confines of which sieve elements 
1 and 2 occur, a longitudinal wall was laid down. This wall has a some- 
what radial orientation in the upper half of the phloic initial (Fic. 5, 
a—c), but in the lower half it is somewhat tangential (Fic. 5, d) and 
finally oblique (Fic. 5, e). (It would appear that the resultant daughter 
cells were placed somewhat radially in relation to one another near their 
upper end and somewhat tangentially near their lower end. Such ar- 
rangements were commonly seen in tangential sections, cf. Fics. 7-9.) 
Of the two cells thus formed, one (below and right) was the mother cell 
of element 2. A longitudinal division along the outer tangential wall of 
this mother cell separated a narrow cell from a larger cell; the former 
divided horizontally thrice and formed a strand of four companion cells, 
the latter matured as the sieve element 2. In the other cell (above and 
left) was laid down a wall which had a tangential orientation in the upper 
end of the cell (Fic. 5, a-c) and a more or less radial orientation in 
the lower end (Fic. 5, d-e) and which separated the mother cell of 
sieve element 1 from a precursor of a parenchyma cell. A longitudinal 
division along the outer tangential wall of the sieve-tube mother cell of 
element 1 produced a short, narrow cell, which divided horizontally and 
formed two companion cells, and a larger cell that differentiated as sieve 


1962] SRIVASTAVA & BAILEY, CACTACEAE, V 245 


q 
b 
c 
d 
e 
| 2 3 4 5 6 
ale es ee | 
* 150 Bln il Sfr 2 a a ae 
Sole: Se ~ eae an b 
2 ee “| ‘tt aT > 
¢ ae 2 ah alee lt be 
: TI 
d ae |! ae lf bd 
S : é : | 
e : 2 | | I De 
pi Ream | NG meni | 
re) Lae —l_. 


5. Pereskia sacharosa, TIER D of Fic. 1: a-e, cross sections of tier at 
levels indicated in schematic representation (f), as in Fic. 2; a-e, X 495. De- 
tails as given in text. 


element 1. Sieve elements 4 and 5 and their associated parenchymatous 
cells originated in a similar manner. 


Trier E: In tier E there are eight sieve elements with their associated 
companion and parenchyma cells (Fic. 6, c, f). Sieve elements 1, 4, 5, 
and 8 originated in the usual manner —a tangential or a radial longi- 
tudinal division in the phloic initial separated the precursor of a paren- 


246 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


TI cw - 
{I a | TB ge 
a ly ly Gf OM - 2K 2 pa 
oa ‘| tt yh 2 
| : 
b br tt] | a il ee i 
eit at | :|! ili ‘ 
: i \ : : 
c TI ee | a be 
SJE sf s i 3 eS 
e at :| I | a ee bd 
ai ai. | Ae ie 4 
: “TI Z| I | | 2 a be 
|] a ail cat ; i 
0 hi i: rene 


Fic. 6. Pereskia sacharosa, TrER E of Fic. 1: a-e, cross sections of tier at 
levels indicated in schematic representation (f), as in Fic. 2; a-e, X 495. Details 
as given in text. 


chyma cell (or strand) from the sieve-tube mother cell. In the sieve-tube 
mother cell itself a longitudinal division separated the precursor of one 
or more companion cells from the precursor of the sieve-tube member. 
Sieve elements 2 and 3, and 6 and 7 are ontogenetically related and 
originated in the manner common to that of sieve elements 1 and 2, and 
10 and 11 in tier B; and 1 and 2, and 4 and S in tier D. 


1962 | SRIVASTAVA & BAILEY, CACTACEAE, V 247 


Discussion 


The five tiers studied from cross sections reveal some patterns about 
the method of origin of sieve elements and related parenchymatous cells 
in Pereskia sacharosa. In the following, this information is summarized 
and confirmed from a study of tangential sections. 

In the five tiers examined, a total of 32 phloic initials was studied. 
Cells internal to the cell complex in which sieve element 1 occurred are 
excluded from this discussion. As mentioned earlier, they may have been 
fusiform phloem-parenchyma cells or phloem-parenchyma strands, but 
we were not certain that these cells had differentiated. Of the 32 phloic 
initials, 5 had matured directly as fusiform phloem-parenchyma cells: 
the remaining 27 phloic initials had divided in various planes and had 
formed cell complexes in which one or more members had differentiated 
as sieve elements. Most commonly, the first division in the phloic initial 
is longitudinal and radial, oblique, or tangential. The resultant daughter 
cells may be unequal in size, but one is the precursor of a parenchyma 
cell, the other the precursor in which the sieve element and its companion 
cells arise (sieve-tube mother cell). This method of the origin of paren- 
chyma cells, sieve elements and companion cells was encountered in 14 
of the 27 silat initials studied (cf. Fics. 2—6, 

Less commonly, the first division in the Snleie: initial, which again is 
longitudinal, but may vary from radial, oblique, to tangential, separates 
two daughter cells, one of which behaves like the phloic initial, the other 
like the sieve-tube mother cell mentioned in the previous paragraph. As a 
result, two sieve elements, at least one parenchyma cell (or strand), and 
some companion cells are formed within the confines of a single phloic 
initial (cf. Fics. 2-6, f). As mentioned during the analysis of the cell 
complex including sieve elements 1 and 2 in tier B, the sequence of the 
first two divisions is not very clear in such instances. 

Occasionally, two successive oblique divisions in the phloic initial sep- 
arate three cells. The dividing walls may be oriented in different planes 
but usually one is somewhat radial, the other somewhat tangential. Also, 
one or both walls may not extend to the tips of the phloic initial, thus 
producing cells shorter than the initial. (Compare the lengths of sieve 
elements 3 and 4 and their associated parenchyma cells in Fic. 2, f; sieve 
elements 4, 5, and 6 in Fic. 3, f; sieve elements 6, 7, and 8 in Fic. 4, f.) 
Of the three cells that are formed, all three may behave as sieve-tube 
mother cells or one may behave as a sieve-tube mother cell and the other 
two as precursors of parenchyma cells, or strands. As a result, three 
sieve elements with their companion cells (sieve elements 4—6, Fic. 3, f; 
sieve elements 6-8, Fic. 4, f), or one sieve element with its companion 
cells and two parenchyma cells (sieve elements 3 and 4, Fic. 2, f) are 
formed within the confines of a single initial. 

Three other patterns of divisions in phloic initials were recorded. A 
longitudinal division in the phloic initial may separate two cells. Each 
daughter cell may then behave as a sieve-tube mother cell (sieve elements 


248 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


6 and 7, Fic. 2, f), or one may behave as a precursor of a parenchyma cell 
(or strand) and the other as the precursor of two sieve-tube mother cells 
(sieve elements 8 and 9, Fic. 3, f). Finally, a phloic initial may be di- 
vided almost horizontally into two daughter cells. The upper and lower 
daughter cells may then behave as more or less “independent” phloic 
initials producing sieve elements and parenchymatous cells in manners 
described above (sieve elements 1-4, Fic. 4, 

In order to confirm some of the conclusions drawn from a study of 
cross sections, several tiers were also studied by use of tangential sections. 
As is clear from Fics. 2-6, a—e, the planes of divisions in the phloic initials 
and their daughter cells are often irregular, with the result that, even in 
almost perfect tangential sections, the outlines of cells appear at different 
levels of focus in the same section and even in different sections. An- 
other difficulty with tangential sections is that the tangential walls of 
derivatives (and, hence, the tangential limits of phloic initials) are not 
easy to determine. Because of these complications serial sections must 
be studied. 

Three representative tiers are drawn in Fics. 7-9. Although partly 
corrected, these figures show the outlines of cells at different levels of 
focus as truly as was necessary for a clear understanding of the planes 
of divisions. Only a few derivatives in each tier are shown; the last 
cell in each set of drawings represents the fusiform cambial initial. The 
sieve cells are left unmarked, except for the sieve plates which are 
hatched; companion cells are stippled; parenchyma cells ontogenetically 
related to sieve elements are shown with nuclei; and parenchyma cells 
ontogenetically unrelated to sieve elements are shown with a circle drawn 
with a heavy pen. 

A phloem-parenchyma strand occurs in Fic. 7, a. It was derived by 
a single transverse division in the phloic initial. The derivatives shown 
in Fic. 7, b-c, d; and at a—b, and c-d in Fics. 8 and 9, are ontogenetically 
related and originated after divisions within the confines of a single phloic 
initial. One parenchyma cell, a sieve element and a strand of two com- 
panion cells occur in the cell complexes shown in Fic. 7, d; Fic. 8, c-d; 
and Fic. 9, a—b, c-d. In the cell complex shown in Fic. 7, b-c, three 
sieve elements and their companion cells were formed; whereas in the 
cell complex in Fic. 8, a-b, two parenchyma cells were formed in associa- 
tion with the sieve element. The planes of divisions leading to the forma- 
tion of cell complexes shown in Fic. 7, d, Fics. 8 and 9, c-d, are easily 
understood. Others are somewhat more complicated but can be explained 
on the basis of our information obtained from the study of cross sections. 

The wall separating the sieve element from the strand of two compan- 
ion cells in Fic. 9, a-b, appears radial at the lower end, tangential at the 
upper end. This is shown by the fact that these elements are placed 
radially to each other at their lower end but appear in different planes of 
focus near their upper end. Similar orientations of derivatives were en- 
countered in the study of cross sections (cf. cell complex including sieve 
elements 1 and 2 in Fic. 5, a-e). It would appear from these illustra- 


d e 


8 9 


Fics. 7-9. Pereskia sacharosa: a-e, in each figure, successive phloem derivatives from outer to inner phloem (a-d) and fusi- 
form cambial initial of each tier (e), in tangential view, X 325. Derivatives at (a), (b-c), and (d) in Fic. 7, and those at 
(a—b) and (c-d) in Fics. 8 and 9 are ontogenetically related. Broken lines indicate cell outline seen more clearly at another fo- 
cal level. Details of figures and of Fics. 15-20, 26-31 given in text, p. 248. 


A ‘AVAOVLOVO ‘AAD TIVE ® VAVLSVAIYS [Z961 


6b~ 


250 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLIII 


tions that, during cytokinesis in phloic initials and their daughter cells, 
the cell plates may, in their upper and lower extensions, intersect different 
walls of the mother cell. Possibly, also, the cells formed after divisions 
within the confines of a phloic initial expand differentially in mutual 
adjustment with the related cells derived from the same phloic initial 
and with the cells in neighboring tiers, so that the shape of the mature 
cells and the orientations of the walls that separate them are different 
in final stages from those in early stages of ontogeny. These factors may 
account for the peculiar configurations of cells seen in Fic. 7, 

and 9, a—b, and for the lateral intrusion of cells in one tier into neighbor- 
ing tiers (Fic. 1). 


ORIGIN OF PHLOEM ELEMENTS IN 
PERESKIOPSIS AND QUIABENTIA 


Tiers in Pereskiopsis aff. chapistle and Quiabentia aff. chacoensis were 
studied from serial cross sections in the same manner as in Pereskia 
sacharosa. Five tiers from each of these species are analyzed in detail in 
Fics. 10-14, a—d, and Fics. 21-25, a-d. The cross sectional views of the 
tiers and the schematic representation of sieve elements and _ related 
parenchymatous cells in them are drawn in the same manner as in 
Pereskia sacharosa. However, the cross sections are drawn only at three 
levels in the axial extent of each tier, because the difference in the struc- 
ture of the tier from one level to another was not very marked. These 
figures are mostly self-explanatory and do not need much comment. In 
the following, therefore, only the important points concerning the oc- 
currence of fusiform phloem-parenchyma cells and phloem-parenchyma 
strands, and the number and planes of divisions in phloic initials (and 
their daughter cells) preceding the formation of sieve elements and re- 
lated parenchymatous cells are mentioned. 

Altogether, 18 phloic initials in Pereskiopsis aff. chapistle and 35 in 
Ouiabentia aff. chacoensis were studied in the five tiers examined for each 
species. (Cells internal to the cell complex including sieve element 1 in 
all tiers, except that in Fic. 24, are omitted from this consideration. In 


considered as fusiform phloem-parenchyma cells, because the cells in 
neighboring tiers at that distance from the cambium appeared mature.) 
Two of the 18 phloic initials in Pereskiopsis aff. chapistle had given rise 
to one fusiform phloem-parenchyma cell (Fic. 13, a-c) and one phloem- 
parenchyma strand (Fic. 14, a-c), and 15 of the 35 phloic initials in 
Ouiabentia aff. chacoensis had matured as fusiform phloem-parenchyma 
cells (Frcs. 21-25, a-c). The remaining 16 phloic initials in Pereskiopsis 
aff. chapistle and 20 in Ouiabentia aff. chacoensis had formed cell com- 
plexes with at least one cell in each case differentiating as a sieve element. 

As in Pereskia sacharosa, the most common mode of origin of sieve 
elements and related parenchymatous cells is one in which the phloic 
initial divides longitudinally and radially, obliquely, or tangentially. Of 


— 


1962 | SRIVASTAVA & BAILEY, CACTACEAE, \V 251 


fe) d 
Ke) 

300 l 2 3 4 
Aue? ald mane 
al 5 eee 
ti Tk 
alt ota fee 
ac aaa |. 

i al ot 
b 4 lh ly :{/ De 
aT Ne 
| Seek 
an elie. a 

| ml ase 
ae 

a ee ge 
ip 2 

aa 


6) d 
II 
Fics. 10, 11. Pereskiopsis aff. chapistle: a-c, cross sectional views of two 
tiers at levels shown in diagrams (d), schematic representations of some mem- 
bers of the two tiers, as in Fic. 2; a-c, * 390. See explanation in text. 


252 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


the two resultant daughter cells, one cell behaves as a precursor of a 
parenchyma cell, or strand, the other cell as a sieve-tube mother cell. 
Such an origin of sieve elements and related parenchymatous cells was 
recorded in 8 of the 16 phloic initials in Pereskiopsis aff. chapistle (Fics. 
10-14, d) and 15 of the 20 phloic initials in Quiabentia aff. chacoensis 
(Fics. 21-25, d). 


~~ ] |e 


b < bb 
c 4 4 A 
0 d 
l2 
I 2 3 
2257 el mM 
a |! be 
| af 
| 1] 
J a} all 
b 4 ah ti pb 
af ar 
“| I “| 
ats TY 
c 4 i 2 ‘|i be 
=e an 
o! at 
d 


Re) 
Fics. 12, 13. Pereskiopsis aff. chapistle: a—c, cross sectional views of two tiers 
at levels shown in diagrams (d), schematic representations of some members of 
the two tiers, as in Frc. 2; a-c, & 390. For details, see text. 


1962] SRIVASTAVA & BAILEY, CACTACEAE, V 253 


an i 2 3 
qytoqo7 
a 4 | ‘ | pa 
| -[I 
| le 
tel lt 
|! 1 
b 4 a | pb 
1 at 
et ae 
| TI 
ate an 
Eta | 
c 4 oll ma: 
PA, gel ! 
oe il SB. 
ja d 
fs) 
O 
a b Cc d e f h 
I5 2 


Fics. 14, 15. Pereskiopsis aff. chapistle. Fic. 14, a-c, cross sectional views 
of a tier at levels indicated in diagram (d), a schematic representation of some 
members of tier, X 390. For details, see text. Fic, 15, successive phloem deriv- 
atives (a-g) and the fusiform cambial initial (h) of a tier in tan ential view, 

220. Derivatives at (a), (b-c), (d-e), (f), and (g) are ontogenetically re- 
lated. Symbols explained in text. 


254 JOURNAL OF THE ARNOLD ARBORETUM | VOL. XLIII 


iss Coe sayy tee ee ee ee 


16 
9 
Fics. 16-20. Pereskiopsis att. chapistle: representative sieve elements and 
despciated sacar adege cells, drawn from tangential sections, 220. De 
rivatives in each figure originated after divisions in a single phloic initial; some 
derivatives separat ted in Fics. 17-20 for better i sieve elements 


in Frc. 18 with a sieve plate on their common lateral wall. Broken lines represent 
outlines of cells seen more clearly at a different focal level; other symbols ex- 
plained in text. 


Two other patterns of divisions in phloic initials recorded in Ouiabentia 
aff. chacoensis were similar to those seen in Pereskia sacharosa. In one 
method two successive, oblique longitudinal divisions in a phloic initial 
resulted in three daughter cells, one of which matured as a sieve element, 
the other two as parenchyma cells (sieve elements 3 and 4, Fic. 22, a-d). 
Such an origin of sieve elements and parenchyma cells seems to be a 
variation of the same method as was recorded for sieve elements 3 and 4 
in Fic. 2, f, except that in this instance the sieve-tube mother cell differ- 
entiated directly as the sieve element without forming any companion 
cells. Sometimes the phloic initial divided obliquely and the resulting 
daughter cells behaved as sieve-tube mother cells of two elements (sieve 
elements 2 and 3, Fic. 23, a-d; cf. also sieve elements 6 and 7, Fic. 2, f). 

In some other patterns recorded in Pereskiopsis aff. chapistle and Quia- 
bentia aff. chacoensis the number of divisions in phloic initials preceding 
the differentiation of sieve elements and parenchymatous cells was much 
Sometimes the sieve-tube mother cell, formed after the first longi- 


less. 


i 4 
O ama d 
1G. 21. Quiabentia aff. chacoensis: a-c, cross sectional views of a tier at 
levels indicated along the margins of diagram (d), a schematic representation of 
some members of the tier, as in Fic. 2; a-c, X 390. For details, see text. 


tudinal division in the phloic initial, matured directly into a sieve element. 
A parenchyma cell (or strand) was formed in association with the sieve 
element but no companion cells were formed (sieve element 2, Fic. 10, d). 
In a few instances, the phloic initial behaved as the sieve-tube mother 
cell, the first division setting off a precursor of one or more companion 
cells and a precursor that matured as a sieve element. No parenchyma 
cells were formed in these cell combinations (sieve elements 1 and 3, Fic. 
10, d; sieve element 2, Fics. 11-14, d; sieve element 4, Fic. 21, d; and 
sieve element 5, Fic. 22, d). In still other cases, the phloic initial did not 
divide at all and matured directly into a sieve element (sieve element 3, 
Fic. 12 


? 


256 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLIII 


Pb 


De 


L 
[ 


J | 


Fic. 22. Quiabentia aff. chacoensis: a-c, cross sectional views of a tier at 
levels indicated along the margins of diagram (d), a schematic representation, as 
in Itc. 2; sieve element 6 and associated companion cell partly crushed; a-c, X 
390. For details, see text. 


Several tiers in Pereskiopsis aff. chapistle and Quiabentia aff. cha- 
coensis were studied from serial tangential sections. The results obtained 
from this study generally confirmed the observations made from cross 
sections (cf. Fics. 15, 26). Some cell complexes including sieve elements 
and ontogenetically related parenchymatous cells selected from different 
tiers of these two species are shown in Fics. 16-20 and 27-31. These 
figures emphasize the variations in number and planes of divisions in 
different phloic initials, and also give an indication of the number of 
sieve elements that may be formed within the confines of a single phloic 
initial. 


Fic. 23. Quiabentia aff. chacoensis: a-c, cross sectional views of a tier at 
levels indicated along the margins of diagram (d), a schematic representation of 
some members of tier, as in Fic. 2; a-c, X 390. For details, see text 


t may appear from our analysis of tiers that in Pereskiopsis aff. 
chapistle and Ouiabentia aff. chacoensis, as compared to Pereskia sacha- 
rosa, relatively few divisions occur in the phloic initials and relatively few 
parenchymatous cells are formed in association with the sieve elements. It 
would also seem that the planes of divisions are lesss irregular and confus- 
ing in the first two species than in the last. It must be stated, however, 
that 2 or 3 sieve elements and related parenchymatous cells are frequently 
formed from a single phloic initial in Pereskiopsis aff. chapistle and Quia- 
bentia aff. chacoensis (Fics. 17-20, 30), and sieve elements with no onto- 
genetically related parenchymatous cells may be formed in Pereskia sacha- 
rosa, particularly in the neighborhood of rays (Fic. 35). Also, the planes of 
divisions in the phloic initials and their daughter cells in Pereskiopsis aff. 
chapistle and Ouiabentia aff. chacoensis (Fics. 17, 18, 20, 30) are often 
as irregular and confusing as in Pereskia sacharosa. If quantitative dif- 
ferences exist between these species, only a wider sampling of tiers and 
material would reveal them and make the results statistically significant. 


258 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLIII 


oC 


nm 
nN 
on 
Ss fe, Sa ee ee ey ees een ee Sees ees i 


TSO Oe TE 
“Ty eee Fe re] 


ce) 


Pic. 24. Quiabentia aff. chacoensis: a-c, cross sectional views of a tier at 
levels indicated along margins of diagram (d), a schematic representation of 
some members of tier, as in Fic. 2; a-c, * 390. Three cells internal to cell com- 
plex including sieve element 1 considered to be fusiform phloem-parenchyma 
cells. Details of drawings in text. 


1962 | SRIVASTAVA & BAILEY, CACTACEAE, V 259 


| 

| 

| 

| 

| 

| 

| 

| 

| 

: abl 

: | 

| 

: : : sie 

| ral : oh 
O =i ea a 


1G. 25. Quiabentia aff. chacoensis: a-c, cross sectional views of a tier at 
levels indicated along margins of diagram (d), a schematic representation of 
some members of tier, as in Fic. 2; a-c, X 390. Details of drawings in text. 


d 


260 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLII 


@ 


f g h 


Vic, 26, Quiabentia aff, chacoensis: successive phloem derivatives (ah) and 
the fusiform cambial initial (i) of a tier in tangential view, < 220. Derivatives 
at (a), (b-c), (d-e), (f-g), and (h) derived from single phloic initials. Symbols 
explained in text. 


1962] SRIVASTAVA & BAILEY, CACTACEAE, V 261 


2? 28 


Fics. 27-31. Quiabentia aff. chacoensis: representative sieve elements and 
associated parenchymatous cells in tangential view, x 200. Derivatives in each 
figure originated after divisions in a single phloic initial. In Fics. 27 and 28, 
parenchyma cell is seen both above and below sieve element with which it is 
associated. Sieve elements in Fic. 30, b, are connected by a lateral sieve plate. 
Symbols explained in text. 


262 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


SIEVE ELEMENTS 


Sieve elements are considered to be the principal conduits through 
which carbohydrates and other food materials are transported within a 
plant. Their structural peculiarities are important, therefore, not only 
to physiologists concerned with problems of translocation, but also to 
morphologists interested in the evolutionary aspects of the phloem tissue as 
a whole. Esau, Cheadle, and Gifford (1953) suggested some possible trends 
of specialization in the phloem. In recent years, some new information, 
particularly about the length of sieve elements in relation to that of the 
phloic initials from which they are derived and about the association of 
sieve elements with parenchymatous cells, has been added through de- 
tailed ontogenetic work (Esau & Cheadle, 1955; Cheadle & Esau, 1958: 
Evert, 1960). In the following paragraphs some structural features of 
the sieve elements in leaf-bearing Cactaceae, their contents, wall struc- 
ture, sieve areas, and length, are emphasized for a better understanding 
of their physiologic and evolutionary specialization. 

As is typical of this tvpe of cell, the sieve elements in the leaf-bearing 
Cactaceae are enucleate in their functionally mature state. The fixa- 
tive and the staining procedures used by us were not conducive to a 
critical cytological study. However, numerous, doughnut-: shaped bodies, 
probably carbohydrate granules, were seen in the sieve elements. These 
granules aggregated in large numbers near the sieve plates and_ sieve 
areas (Fics. 39, 41). In addition, the material described in the litera- 
ture as “slime” was often present in copious quantities. It was usually 
deposited on one side of a sieve plate in the form of a plug and extended 
like a strand through most of the cell lumen (Figs. ody 61), Jn 1onpi- 
tudinal sections, this feature was often useful in determining the extent 
of a particular sieve element. It was rather scanty in Pereskiopsis aff, 
chapistle (Fic. 39). 

Nacreous walls, reported in the sieve elements of various dicotyledons 
by Esau and Cheadle (1958), were uniformly lacking in the material 
studied by us (cf. Figs. 32, 38, 40). In permanent slides the walls of 
sieve elements appeared rather thin, but they were birefringent under 
polarized light 

The common walls between two sieve elements bear numerous sieve 
areas, some of which are more specialized than others and are termed sieve 
plates (cf. Cheadle & Whitford, 1941; Esau, 1950). Sieve areas occur 
on radial as well as tangential walls (Fics. 44, 48): and their size, as 
determined by the number of callose cylinders stained by lacmoid, varies 
considerably (Fic. 51). The difference in size of connecting strands and 
pores in sieve areas that occur in the sieve plates on the one hand and 
those that occur elsewhere on the walls is very marked (Fics. 50, 51 
The sieve plates are generally of the simple type with a single sieve area, 
but sieve plates with two or more sieve areas are present also (Fic. 50 
As a rule, every sieve element has two sieve plates (cf. Fics. 7-9, 15— 20, 
26-31); but some sieve elements may have as many as 3 or 4 sieve plates 


eu 


a 


1962] SRIVASTAVA & BAILEY, CACTACEAE, V 263 


(Fics. 18, b; 30, b) others may have only one (Fics. 7, b, c; 18, a; 20, a). 
The position of sieve plates in the sieve elements varies likewise. Com- 
monly, the sieve plates occur at or near the ends of the sieve elements, 
but sieve plates on lateral walls are not unusual (Fics. 7, b; 8, b; 18, a, b; 
30,b). The final shape, size, and orientation of the mature sieve elements 
is often different from that of the phloic initials from which they are de- 
rived. Some sieve elements may be rather grotesque in shape (Fics. 7, b; 
8, b; 17, b); and the long axis of some may be placed slantingly in rela- 
tion to the vertical axis of the phloic initial and, hence, that of the stem 
(Fics. 9, b; 20, a; 28; 29, b; 41). Such slantingly placed sieve elements 
are often connected through sieve plates with sieve elements in laterally 
adjacent (rather than vertically adjacent) tiers (Fic. +1). Often, two 
sieve elements derived from the same phloic initial jorm parts of two 
different sieve tubes, one with a vertical, the other with a diagonal 
orientation in relation to the long axis of the stem. Laterally placed 
sieve plates are encountered rather frequently in areas where diagonally 
placed sieve elements abut on the lateral walls of adjacent sieve elements. 

The length of a sieve element depends not only on the length of the 
phloic initial from which it arises, but also on the number and planes of 
divisions in the phloic initial and its daughter cells and on the number and 
planes of divisions in the sieve-tube mother cell.t The length of sieve 
elements as measured from transections (Fics. 2—6, f; 10-14, d; 21-25, d) 
is subject to an error which must be mentioned. When the sieve plates 
occur near but not at the end of the sieve element (e.g., Fics. 7, d; 17, a; 
30, b), the tapering end of the cell may be missed from calculations. The 
following statements about the lengths of sieve elements and the phloic 
initials from which they are derived are, therefore, liable to some error. 

Since the phloic initials in a tier are usually derived by symmetric 
periclinal divisions in a fusiform cambial initial, one normally expects 
them to be of approximately equal length (e.g., Fics. 4, f; 10, d). The 
fluctuations in the lengths of phloic initials in several tiers (e.g.. Fics. 
2,f: 6, f; 25, d) are, therefore, somewhat unusual. A possible explanation 
for the different lengths of the phloic initials in a tier would be that the 
individual phloic initials elongated intrusively in varying degrees. But 
this explanation is contrary to the commonly held belief that the phloic 
initials, in contrast to fusiform cambial initials, do not elongate intru- 
sively. It appears that in the leaf-bearing @ictaceae the periclinal di- 
visions in the fusiform cambial initials are often not strictly longitudinal 
and the dividing walls fall short of one or both tips of the mother 
initial. The daughter cells formed after such an asymmetric periclinal 
division may be equal or unequal in length, but they are vertically dis- 
placed (at one or both ends) in relation to one another. If a number of 
such asymmetric periclinal divisions occur in the daughter cell that be- 


‘ This reasoning is based on the assumption that the sieve clements do not undergo 
intrusive elongation during their differentiation from the phloic initials or their daughter 
cells. Several studies on the phloem of gymnosperms and dicotyledons support this 
assumption (Bannan, 1950; Cheadle & Esau, 1958; Evert, 1960; Srivastava, 1962). 


264 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


haves as the fusiform cambial initial after each division and if such di- 
visions are interpolated among others that are symmetric, the net result 
as seen in mature tissue would be comparable to that shown in Fics. 
2,f; 6,f; 15; 25.d; and 26. In its radial extent, the tier would show 
fluctuations in height that would be independent of the intrusive growth 
of the fusiform cambial initial. The intrusive growth of the fusiform 
cambial initial, if it occurred after a periclinal division, would further com- 
plicate the pattern. 

In instances where the phloic initial matures directly as a sieve element 
and no parenchymatous cells are formed (Fic. 12,d, sieve element 3); 
the length of the sieve element is approximately the same as that of the 
phloic initial. Also, if the phloic initial behaves as a sieve-tube mother 
cell, that is, one or more precursors of companion cells are formed as 
sister cells of the precursor that matures as the sieve element but no 
parenchyma cells are formed (Fics. 10,d, sieve elements 1 an 
sieve element 2; 21,d, sieve element 4; 22,d, sieve element 5), the length 
of the sieve element closely reflects the length of the phloic initial. In 
other instances, the length of the sieve element bears no predictable rela- 
tionship to that of the phloic initial —it may be the same (cf. Fics. 

, C; 15, d-e; 26, f-g) or it may be much less (cf. Fics. 7, b; 8, b, c; 9, b: 
15, b-c, g; 26, a). 

In concluding this section on sieve elements, a few remarks must be 
made about the relative lengths of the sieve-tube members and the paren- 
chymatous elements ontogenetically associated with them. The length of 
parenchyma cells (or strands) may be equal to, or more than, or less than 
that of the sieve element with which they are associated (cf. Fics. 2—6,f: 
10-14,d; 21-25,d). The divisions that separate the precursor of the 
companion cell (or strand) from that of the sieve element are usually 
longitudinal with reference to the long axis of the sieve-tube mother cell, 
but the dividing walls are laid down in such a manner that the precursor 
of the companion cell (or strand) may be almost as long as that of the 
sieve element or it may be shorter (cf. Fics. 7-9; 15-20; 26-31). Oc- 
casionally, the dividing wall is laid obliquely and short of the cell tips 
at either end; as a result, the precursor of the companion cell (or cells) 
at one end projects a little beyond the end of the precursor that matures 
as the sieve element (e.g., Fics. 7,c; 9,a; 19,a; 30, b; 33). Exceptionally, 
the precursor of the companion cell (or cells) may be a little longer 
than the precursor of the sieve element (e.g., Fics. 5, f, cell complex in- 
cluding sieve element 2; 9,c). 


PARENCHYMA CELLS 


Although the parenchyma cells in the phloem of leaf-bearing Cactaceae 
differ in their ontogeny, we were not able to discern any significant 
cytological differences between them with the fixing and staining pro- 
cedures that we used. The fusiform phloem-parenchyma cells and the 
individual cells in phloem-parenchyma strands, which in other plants 


1962] SRIVASTAVA & BAILEY, CACTACEAE, V 265 


commonly store starch, oil, tannins, etc. (cf. Esau, 1953, p. 284), did not 
show any such inclusions in the functional phloem. The parenchymatous 
cells ontogenetically related to the sieve elements also did not show any 
of these inclusions. It is commonly held that companion cells have a 
denser cytoplasm and, in general, stain more deeply than other paren- 
chyma cells in phloem (cf. Esau, 1953, p. 283). We utilized this feature 
in identifying companion cells and were partially successful. But it was 
not always possible to use this criterion because the deeply staining 
appearance of a companion-cell protoplast depends partly on the size of 
the companion cells and partly on the plane of the section with reference 
to the cell lumen. For instance, if a longitudinal section passes close to a 
lateral wall rather than through the middle of the cell, the protoplast may 
appear lightly stained. Because of these features it was difficult for us 
to be certain about the identity of individual parenchymatous cells in 
isolated cross and tangential sections. With the use of serial sections one 
can normally distinguish between fusiform phloem-parenchyma cells and 
phloem-parenchyma strands, on the one hand, and the parenchymatous 
cells ontogenetically related to the sieve elements, on the other. It is also 
possible in most cases to distinguish between companion cells and paren- 
chyma cells ontogenetically associated with the sieve elements, because 
the length of the companion cells (or strand of companion cells) usually 
does not exceed that of the sieve element, whereas that of the parenchyma 
cells (or strands) frequently may. Still, in several cell configurations 
(e.g., those including sieve elements 3 and 4, Fic. 22, a—d) it was almost 
impossible to tell whether one was dealing with companion cells or paren- 
chyma cells or both. (In this instance, the cells were interpreted as 
parenchyma cells.) 

Sieve elements are usually connected with their associated companion 
cells through one-sided sieve areas. Such sieve areas have densely stain- 
ing connecting strands and callose cylinders only in the wall belonging to 
the sieve element; in the companion cell wail there is only a primary pit 
field. Cheadle and Esau (1958) and Evert (1960) reported such one- 
sided sieve-area connections between sieve elements and the parenchyma 
cells ontogenetically associated with them also. In the material of leaf- 
bearing Cactaceae that we have examined, sieve-area connections were 
common between sieve elements and companion cells and between sieve 
elements and ontogenetically associated parenchyma cells (Figs. 45, 46, 
49). Although no counts were made, our general impression is that the 
number of sieve-area connections between sieve elements and companion 
cells is usually higher than that between sieve elements and ontogenetically 
associated parenchyma cells. Cheadle and Esau (1958) made a similar 
observation. One notable feature about the sieve-area connections between 
sieve elements and ontogenetically related parenchyma cells was that the 
connections occurred in those portions of the common wall that bulged 
somewhat in the lumen of the parenchyma cell (Fics. 46, 52). In contrast, 
the common wall between the sieve elements and their companion cells, 
although covered with numerous sieve areas, was usually straight (Fic. 


266 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLHI 


45). Sieve-area connections usually occur between sieve elements and 
parenchymatous cells derived from the same phloic initials; but occasion- 
ally sieve elements derived from one phloic initial may be connected to a 
parenchymatous cell that forms part of a cell complex derived from a 
different phloic initial. Such connections were recorded especially in those 
parts of a parenchymatous cell that projected beyond the end of the 
sieve element to which it was ontogenetically related. 

Sieve-area connections were also observed between sieve elements and 
some fusiform phloem-parenchyma cells (Fic. 47). The sieve areas oc- 
curred in those portions of the common wall that bulged into the lumen of 
the parenchyma cell. The fusiform phloem-parenchyma cells that were so 
connected with sieve elements occurred in the same tier as the sieve element 
or in neighboring tiers, but their number was relatively very small and rela- 
tively few sieve-area connections occurred on the common wall. Fusiform 
i eee cells in such intimate contact with the sieve elements 
were more common in Pereskiopsis aff. chapistle and OQuiabentia aff. cha- 
coensis than in Pereskia sacharosa. 

In the nonfunctional phloem, sieve elements and companion a lose 
their living contents, collapse, and sooner or later are crushed (Fics. 36, 

37). It is our impression that the parenchyma cells ontogenetically a 
to the sieve elements survive for a longer time than the related sieve ele- 
ments and companion cells, but eventually they also lose their protoplast, 
collapse, and are crushed. The fusiform phloem-parenchyma cells an 
the individual cells of phloem-parenchyma strands seem to survive and 
sometimes store starch and even accumulate druses of calcium oxalate. 
However, in the absence of any definite cytological differences between 
the parenchyma cells ontogenetically related to sieve elements and those 
that are not so related, and because of a general distortion of the tissue, 
it is difficult to determine in nonfunctional phloem which cells or strands 
may or may not have been ontogenetically related to the sieve elements. 
Consequently, we are not sure whether all fusiform phloem-parenchyma 
cells and phloem-parenchyma strands continue to live in the nonfunctional 
phloem; we are also not sure whether @// parenchyma cells and strands 
ontogenetically related to sieve elements die and collapse in the nonfunc- 
tional phloem. 


mes 


DISCUSSION AND CONCLUSIONS 

Our studies on the phloem of leaf-bearing Cactaceae were restricted 
in scope, partly because of the difficulty of obtaining and sectioning the 
material of cacti and partly because of the fixative used to kill it. Only 
those species were selected for a detailed study which could be sectioned 
in a satisfactory manner, and only those aspects of cellular detail were 
followed which were not directly related to seasonal changes and the 
elucidation of which did not require elaborate cytological techniques. 
For these reasons we did not concern ourselves with such aspects of 


1962 | SRIVASTAVA & BAILEY, CACTACEAE, V 267 


phloem study as the development and maturation of sieve elements, the 
cytological inclusions in companion cells and parenchyma cells onto- 
genetically related and unrelated to sieve elements, and the longevity 
of sieve elements. We also did not study the changes from functional to 
nonfunctional phloem in any detail, and our investigation of ray tissue 
was only cursory. 

The secondary phloem of the three species that we investigated is rela- 
tively simple. Sieve elements and parenchyma cells of various types were 
the only constituents in the axial tissue. Oil cells, mucilage cells, fibers, 
etc., were absent from the functional phloem of these species. In the 
nonfunctional phloem several parenchyma cells accumulated druses of 
calcium oxalate and, in Pereskia sacharosa, some were modied as sclereids 
(Bailey, 1961a, b). Intercellular spaces were commonly seen in functional 
and nonmineliondl phloem (Tics. 32, 38, 40). The rays are typically high 
and multiseriate and include, especially near the ray margins, some upright 
cells that may be almost as long as some of the fusiform derivatives. High 
and broad rays are often dissected by a conversion of ray initials into 
fusiform initials. In the early stages of such conversion the sieve elements 
that are formed are often of the same size as other ray cells and frequently 
lack companion cells (Fic. 35). Fusiform initials are converted into ray 
initials also, with accompanying changes in the nature of their derivatives. 

Ontogenetic studies of phloem yield valuable data about the origin 
and interrelationships of different phloem elements. Our analysis of tiers 
in the phloem of leaf-bearing Cactaceae revealed several possible ways in 
which sieve elements and parenchymatous cells may arise from phloic 
initials. Most of these methods of origin have been discussed in detail 
elsewhere in this paper. In the following some of the main conclusions 
are summarize 

The three representatives of the leaf-bearing Cactaceae studied, Pereskia 
sacharosa, Pereskiopsis aff. chapistle, and Quiabentia aff. chacoensis, do 
not seem to differ basically in the method of origin of phloem elements. 
Fusiform phloem-parenchyma cells and phivem-parenchyma strands arise 
either directly from or after a few horizontal divisions in the phloic ini- 
tials. In contrast, the differentiation of sieve elements and parenchyma- 
tous cells ontogenetically related to them is usually preceded by several 
divisions in the phloic initials and their daughter cells. Evidence of such 
divisions was seen not only in mature tissue (cf. Frcs. 33, 34), but also in 
immature tissue close to the cambium (cf. Fics. 42, 43). Commonly, a 
parenchyma cell (or strand) is formed in association with a sieve ele- 
ment and its companion cells; but sometimes the companion cell, or the 
parenchyma cell, or both are lacking, and sometimes several parenchyma 
cells, sieve elements, and companion cells are formed within the confines 
of a single phloic initial. Apparently, the phloic initials and their daughter 
cells show varying degrees of mitotic activity before cells are differenti- 
ated. Such variations among different phloic initials may be encountered, 
not only among different tiers of the same species, but also within the 


268 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


same tier. Also, one tier may be composed predominantly of fusiform 
phloem-parenchyma cells and phloem-parenchyma strands and may have 
very few sieve elements (Fic. 24, a—c), whereas a neighboring tier may 
have numerous sieve elements and ontogenetically related parenchymatous 
cells and relatively few fusiform phloem-parenchyma cells and phloem- 
parenchyma strands (Fic. 25, a-c). We do not know what factor or fac- 
tors may be responsible for these variations in the behavior of phloic 
initials in the same tier and in neighboring tiers. 

Although the final number of parenchyma cells, sieve elements, and 
companion cells arising within the confines of a phloic initial varies con- 
siderably, the sequence of divisions seems to be such that the precursors 
of parenchyma cells arise as sister cells of precursors that behave as sieve- 
tube mother cells, and the precursors of companion cells arise as sister 
cells of the precursor that matures as a sieve element. Two observations 
support this assumption. First, in cross sections, the wall separating the 
parenchyma cell from the complex of sieve element and companion cells 
seems to be the first wall laid down in the phloic initial (or its daughter 
cell) and the wall separating the sieve element from the companion cell 
seems to be laid down in one of the daughter cells formed after the first 
division (Fic. 40). Second, the length of the associated parenchyma cells 
may frequently exceed that of the sieve element, but that of the companion 
cells rarely does (Fic. 41). If our interpretation of the sequence of divi- 
sions within the confines of a phloic initial is correct, the parenchyma cells 
in their ontogenetic relationship would be somewhat more distantly re- 
lated to the sieve elements than the companion cells. But this interpreta- 
tion is applicable only if parenchyma cells, as well as companion cells, are 
formed in association with a sieve element. 

It is significant that the sieve elements in the leaf-bearing Cactaceae 
have sieve-area connections, not only with ontogenetically related, but 
also with ontogenetically ee parenchyma cells. These connections 
are numerous between sieve elements and companion cells, less numerous 
between sieve elements and ontogenetically related parenchyma cells, and 
are only infrequently observed between sieve elements and the parenchyma 
cells that are derived from independent phloic initials. Apparently, the 
varying degrees in which the parenchyma cells are ontogenetically related 
to sieve elements are paralleled by the degree of their physiological affinity 
to sieve elements: those with the closest ontogenetic association are also 
those that have the most intimate physiological association. For future 
research, it will be of extreme interest to determine the cytological features 
of these parenchyma cells and to see in what important ways they differ 
from one another 

From an evolutionary viewpoint, the parenchyma cells physiologically 
associated with the sieve elements present a complicated picture. The 
albuminous cells of gymnosperms, although physiologically comparable to 
companion cells of angiosperms, generally have no ontogenetic association 
with the sieve elements (cf. Strasburger, 1891; Srivastava, 1962). Com- 


1962] SRIVASTAVA & BAILEY, CACTACEAE, V 269 


panion cells are lacking in Austrobaileya scandens, a primitive dicotyle- 
don (Bailey & Swamy, 1949); and it is possible that some parenchyma 
cells derived from independent phloic initials serve as companion cells in 
this plant. Holdheide (1951) reported some parenchyma cells in the 
secondary phloem of dicotyledons that he compared to albuminous cells 
of gymnosperms, thus implying that these cells originated from independ- 
ent phloic initials. Other reports include varying degrees of specialization 
among companion cells themselves (Resch, 1954). In view of these 
findings and the ontogenetic studies carried out by Cheadle and Esau 
(1958), Evert (1960), and in the present work, it may seem that the 
evolution of the parenchyma cells physiologically associated with the 
sieve elements has progressed in the direction of a closer ontogenetic asso- 
ciation with the sieve elements. But the presence of sieve-area connections 
between sieve elements and parenchyma cells ontogenetically unrelated 
to them in a family as highly specialized as Cactaceae presents serious im- 
pediments in the acceptance of this idea. It is clear that more and detailed 
ontogenetic and cytological studies of the phloem of vascular plants need 
to be carried out before any valid generalizations about the evolution of 
parenchyma cells physiologically associated with the sieve elements can 
be made. 

In the preceding paper of this series (Bailey & Srivastava, 1962), 
based upon the study of numerous putative species of Pereskia, Peres- 
kiopsis and Quiabentia, we demonstrated that the fusiform initials of the 
cambium and their derivatives in phloem have attained a high level of 
phylogenetic specialization. This is shown by their short length and their 
tendency to occur in stratified arrangements as seen in tangential longi- 
tudinal sections. The well-developed structure of the sieve plates in the 
sieve elements of the leaf-bearing Cactaceae and the marked difference in 
the size of pores and connecting strands in the sieve plates, on the one 
hand, and the lateral sieve areas, on the other (Fics. 50, 51), also suggest 
an advanced degree of evolutionary specialization. 

In this paper we have been concerned primarily with the phenomenon 
of cell divisions in phloic initials and their daughter cells and the inter- 
relationships between sieve elements and different kinds of parenchyma 
cells. It is evident from our studies that the ontogenetic changes in the 
differentiation of sieve elements, companion cells and parenchyma cells 
are diversified and variable. They obviously are qualitatively similar in 
the three genera that we examined. Although there are some evidences of 
possible quantitative differences in these forms, it is clear that a great 
deal of material must be critically studied before valid taxonomic conclu- 
sions can be drawn. This is in marked contrast to the structure of non- 
functional phloem, where, as one of us (Bailey, 1961a) has shown, 
Pereskia differs from Pereskiopsis and Quiabentia by the presence of 
sclereids, and where three groups of species of Pereskia may be differenti- 
ated upon the basis of form and distribution of such sclerenchymatous 
elements. 


270 JOURNAL OF THE ARNOLD ARBORETUM [VOL LTT 


LITERATURE CITED 
BaILey, I. W. Comparative anatomy of the leaf-bearing Cactaceae, I. Struc- 
ture and distribution of sclerenchyma in the phloem of Pereskia, Pereskiop- 
sis and Quiabentia. Jour. Arnold Arb, 42: 144-156. 1961a 


Comparative anatomy of the leaf-bearing Cactaceae, HI. Form oe 
distribution of crystals in Pereskia, Pereskiopsis and Oia Ibid. 3 
346. 1961b. 


& L. M. Srivastava. Comparative anatomy of the leaf-bearing Cacta- 
ceae, IV. The fusiform initials of the cambium and the form and structure 
of their derivatives. /bid. 43: 187-202. 1962. 

B.G wamy. The morphology and relationships of Austrobaileya. 


ibid. 30: 211-226. 49, 

BANNAN, M. W. The frequency : anticlinal divisions in fusiform cambial cells 
of ae ee Amer. Jour. Bot. 37: 511-519. 1950. 

CHEADLE, V. I. & K. Esau. cae phloem of Calycanthaceae. Univ. Calif. 
Publ Bot. - 397-510. 58. 

. WuitForp. Observations on the phloem in the Monocotyle- 
doneae. - The occurrence and phylogenetic raga in structure of 
the sieve tubes in the metaphloem. Amer. Jour. Bot. 28: 623-627. 1941. 

, E. M. Girrorp, Jr., & K. Esau. A ee ‘combination for phloem and 

contiguous tissues. Stain Tech. 28: 49-53. 1953, 

Esau, K. a ee and structure of the phloem tissue. II. Bot. Rev. 16: 
67-114. 1950. 


ee Anatomy. John Wiley and Sons, N.Y., 1953. 
V. I. CHEADLE. Significance of cell divisions in differentiating sec- 

ondary phloem. Acta Bot. Neerl. 4: 346-357. 

Wall ghickenine in sieve elements. Proc, Natl. cicad, Sc. 445 
546-553. 1958. 

—______—— & E. M. Girrorp, Jr. Comparative structure and possible trends 
of specialization of the phloem. Amer. Jour. Bot. 40: 9-19. 

Evert, R. F. Phloem structure in Pyrus communis L. and its seasonal changes. 
Univ. Calif. Publ. Bot. 32: 127-194. 

HotpHEIDE, W. Anatomie mitteleuropdischer Gehdlzrinden (mit mikrophoto- 
graphischem Atlas). Jv: H. Freund’s Handbuch der Mikroskopie in der 
a Vol. 5, Part 1, p. 193-367. Umschau Verlag, Frankfurt-am-Main, 


a . Beitrage zur Cytologie des Phloems. Entwicklungsgeschichte der 
Siebrdhrenglieder und Geleitzellen bei Vicia faba L. Planta 44: 75-98. 
954. 


one ae L. M. Secondary phloem in the Abietineae. Ph.D. Thesis (unpubl.), 
Univ. California, Davis. 

Pen E. Ueber den Bau und die Verrichtungen der Leitungsbahnen in 
den Pflanzen. Histologische Beitrage, Vol. 3. Gustav Fischer, Jena 1891. 


1962] SRIVASTAVA & BAILEY, CACTACEAE, V DHA 


EXPLANATION OF PLATES 


The following symbols have been used consistently in all the plates: C, cam- 
bial zone; CS, companion cell or strand; FP, functional phloem; Gr, carbo- 
hydrate granules; IS, intercellular space; NP, nonfunctional phloem; P, paren- 
chyma cell or strand ontogenetically related to sieve elements; Par, fusiform 
phloem-parenchyma cell or phloem-parenchyma strand; R, ray; S, sieve ele- 
ment; SA, sieve areas; Sl, slime; SP, sieve plate. 


PLATE I 


32-34. Transverse and tangential views of phloem and cambium of 
Poe sacharosa. 32, Transverse section, X 455. 33, Tangential section of 
phloem showing a strand of parenchyma cells (P) and companion cells (CS) in 
ontogenetic association with a sieve element (S), * 220. A strand of companion 
cells (indicated by arrow) projects below the end of the sieve element with 
which it is associated. Fusiform phloem-parenchyma cells (Par) give an idea 
about the length and size of phloic initials. 34, Tangential section of the cam- 
bium, X 220 


PLATE II 


Fics. 35-37. Tangential and radial views of phloem of Pereskia sacharosa. 

, Tangential section through functional phloem. 36, Radial section of func- 

Ce (right) and nonfunctional eae phloem. 37, Tangential section of non- 
functional phloem. All figures & 45 


PLATE III 


Fics. 38-40. Transverse and ee views of phloem. 38, Pereskiopsis 
aff. chapistle, transverse section of phloem and cambium. 39, The same, saa 
tial section of phloem. 40, ee “ail chacoensis, transverse section of 
phloem and cambium. Cell complexes including a Daren chviia oe 1), sieve 
element (S) and companion cell (CS) are shown. All figures & 45 


PLATE IV 


Fics. 41-43. Tangential sections of phloem and cambium of Quwiabentia aff. 
chacoensis. 41, section through mature phloem. The sieve element on lower 
eft is slantingly placed in relation to the parenchyma cell which occurs in 
association with it. 42, section through immature phloem showing divisions 
in phloic initials marked with oy 43, section through cambium. Fics. 42 and 
43 are photographs of successive sections in a tangential series; the cambial 
initials that produced the phloic na in Fic. 42 marked with (x) are similarly 
marked in Fic, 43, All figures & 2 


PLATE V 
Fics. 44-47. Tangential sections of phloem showing sieve areas between sieve 
related parenchyma cells, and sieve elements and ontogenetically unrelated 


parenchyma cells. 44, Pereskia sacharosa, * 910. 45-47, Quiabentia aff. 
chacoensis, X 910. 


272 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


PLATE VI 

Fics. 48-52. Sieve plates, lateral sieve areas and sieve-area connections be- 
tween sieve elements and related parenchymatous cells. 48, Pereskia sacharosa, 
transverse section showing a sieve area (indicated by arrow) on the tangential 
wall between two sieve elements, X 1140. These sieve elements formed part 
of a cell complex that had originated from a single phloic initial. 49, Quiabentia 
aff. chacoensis, transverse section showing one-sided sieve-area connections (in- 
dicated by arrows) between a sieve element a the companion and parenchyma 
cells that are associated with - < 910. 50, Pereskiopsis aff. chapistle, sieve 
plate in radial view, * 1140. Dien all. chacoensts, lateral sieve areas 
between two sieve elements in renee view, X 1140. 52, The same, tangen- 
tial section showing ongesiged sieve-area connections between a sieve element 
and related parenchyma cell, he sieve areas occur in those portions 
of the common wall that ae into the lumen of the parenchyma cell. 


— 


Jour. ARNOLD Arg. VoL. XLIII 


PLATE I 


V 


BEARING CACTACEAE, 


LEAF- 


) 


SRIVASTAVA & BAILEY 


Jour. 


ARNOLD Ars. VoL. XLIII 


PLaTE II 


SRIVASTAVA & BAILEY, LEAF-BEARING CACTACEAE, V 


' 


Ga Gy 


SRIVASTAVA & BarLey, LEAF-BEARING CACTACEAE, V 


IIITX ‘10A ‘aay dTonay ‘anof 


III SLV1d 


Jour. ARNOLD Arp. VoL, XLIII 


PLATE IV 


SrIvASTAVA & BAILEY, LEAF-BEARING CACTACEAE, V 


Jour. ARNOLD Ars. VoL. XLIII PLATE V 


Jour. ARNoLpD Ars. VoL. XLIII PLaTE VI 


SRIVASTAVA & BAILEY, LEAF-BEARING CACTACEAE, V 


1962] HOWARD, VOLCANISM AND VEGETATION 279 


VOLCANISM AND VEGETATION IN THE 
LESSER ANTILLES 


KicHarp A. Howarp 


THE MAJORITY OF THE ISLANDS of the Lesser Antilles in the Caribbean 
archipelago are volcanic in origin. Two of the islands have active vol- 
canoes which currently are dormant. However, Mt. Pelée on Martinique 
erupted in 1902 and 1930, and the Soufriére on St. Vincent erupted in 
1902. Nine of the islands from Grenada, in the south, to St. Kitts, in the 
north, have active fumaroles or soufriéres, indicating residual volcanic 
activity. Of the few volcanic islands without historic volcanic activity, 
Saba, Redonda, Union, and others show the classic forms of their vol- 
canic origin and prehistoric volcanic activity (11, 13, 48, 60). 

The eruptions of the twentieth century on Martinique and St. Vincent 
have been well observed, studied, and recorded. An extensive descriptive 
literature is available for the volcanic and seismic activities of the area. 
The literature regarding the geological phenomenon of fumaroles in the 
Lesser Antilles is less complete, although a specialized interest in utilizing 
the thermal power of one, the Qualibou soufriére in St. Lucia, has been 
recorded recently (8, 49, 50). 

In nearly all of the geologic studies, as well as in many floristic, phyto- 
geographical and ecological papers on the area, some mention is made of 
the effects of volcanoes and soufriéres on the vegetation. These range 
from Perret’s mention of the attempted use of the sensitive plant, Mimosa 
pudica, to record earthquakes, to the papers by Stehlé and Beard con- 
sidering the progressive changes in the regrowth of vegetation on devas- 
tated volcanic slopes. The present paper, a survey of the nature of vol- 
canic activity and its effects on the vegetation in the Lesser Antilles, is 
based on observations made in 1950 during a field trip which began in 
Trinidad, extended to the northern and western islands, and ended in 
Jamaica. Data and specimens were collected in and around many sites 
of past and present volcanic activity. More recent visits to the Lesser An- 
tilles, particularly to Montserrat in 1961, have allowed comparative ob- 
servations after an elapsed time, as well as the gathering of specific in- 
formation. For comparison with the West Indies, I have had the oppor- 
tunity of studying the effects of the recent eruptions of Kilauea and 
Kilauea-iki and of the various fumaroles on Mauna Loa, in Hawaii. 

These trips, observations, and collections form part of a continuing study 
of the vegetation of the Lesser Antilles. I acknowledge with gratitude the 
support of the American Philosophical Society during the 1950 trip and 
that of the National Science Foundation for grants which have aided more 
recent work. Many people in the Lesser Antilles have aided my work in 


280 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLII 


various ways. I mention in particular the assistance of Hugh McConnie, 

f St. Vincent; James Ross, of Grenada; Frank Delisle, now of Antigua; 
Kingsley Howes, of Montserrat; Harold Simmonds, of St. Lucia; John 
Knowlton, formerly of Dominica; and the late Malcolm Smith, of St 
Kitts. 


HISTORY OF VOLCANISM IN THE LESSER ANTILLES 
Perret (46) has recorded the volcanological history of the Lesser An- 
tilles. A chronology of the important eruptions is the following: 


1692 — earthquakes followed by eruption on Mt. Misery, St. Kitts 
1694 — earthquakes followed by eruption on the Soufriere, Guadeloupe. 
1718 — first recorded eruption of the Soufriére, St. Vincent. 
1765 — earthquakes and gas emission on ae 
1766 — eruption of the Qualibou soufriére, St. Luc 
1798 — earthquakes and eruption of the Soufriere, ‘eunsebupe 
1812 — great eruption of the Soufriére, St. Vincent. 
1838 — eruption of the Soufri¢re, Guadeloupe. 
1851 — earthquake and lateral outbreak of Mt. Pelée, Martinique. 
1880 —ash eruption on Dominica. 
1898 — start of three years of earthquakes ~ es emission on Montserrat. 
1902 — most destructive eruptions of Mt. Pelée, Martinique, and the Sou- 
friére, St. Vincent 
1929 - mnt of Mt. Pe lee, Martinique. 
1934 —start of four years of earthquakes and gas emission on Montserrat. 


The occurrence of earthquakes and actual eruptions is usually well re- 
corded. Nevertheless, some of the earlier dates cited above have been 
questioned. Anderson (2), for example, cites both the picturesque report 
of the 1718 eruption of the Soufriére on St. Vincent, as given by Defoe 
(37), and the subsequent questions raised by the report. To the present 
day, the 1718 eruption is a questionable one. However, the geologists 
apparently have overlooked the work by Rev. Mr. Smith (64) who, while 
on Nevis, “heard six or seven dull bounces of noise resembling those of 
Cannon at a great distance pretty quickly following each other at the 
exact time of this Explosion: as the Sky was quite clear in the eye of 
the Wind, and as none of my acquaintance there took the same notice 
of the thing, I durst not venture to insist much upon hearing those dull 
bounces till I had seen Mr. Boyd.” Mr. Boyd was previously identified 
as the captain of a merchant ship en route from St. Kitts to Barbados 
who noted that the sky grew dark and a horrible noise “far surpassing 
the loudest thunder” and a “falling likewise instantaneously so thick a 
Shower of Ashes, that the Sloop’s Deck was covered two or three inches 


ar) 


deep with them. ... They in fright enough turned back homewards 
[and]... it was soon after found out, That a large Mountain in the 
Island of Saint Vincent . . . abounding in Veins of Sulphur and Brim- 


stone blew up at once, viz. Woods, Rocks &c. all together, which must be 
allowed to cause a most dreadful Explosion.” By contrast, the 19Q2 erup- 


HOWARD, VOLCANISM AND VEGETATION 


1962] 
CF 
ao H 
@ SABA st. EUSTATIUS 
ST. KITTS 
@ NEVIS bys ANTIQUA 
» REDONDA 
j MONTSERRAT 
7 
GUADELOUPE 
@ MARIE GALANTE 
DOMINICA 
Mi PEEE 
MARTINIQUE 
ST. LUGIA 
, BARBADOS 
SOUFRIERE 
Z| ST VINCENT 
o 
“THE GRENADINES 
G GRENADA 


Diagonal lines indicate islands 
and the Soutriére (St. Vincent) 
Solid black areas 


VOLCANIC AREAS OF THE LESSER ANTILLES. 
with active fumaroles. Mt. Pelée Seen 
are currently dormant but have erupted in the present century. 


are volcanic in origin. Unmarked areas are nonvolcanic islands. 


282 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


tions of Mt. Pelée and the St. Vincent Soufriére were observed by parties 
of scientists from eight different countries, and, today, modern seismo- 
logical stations record the minute signs of activity of these two dormant 
volcanoes. 

The occurrence and duration of fumarole activity have been less accu- 
rately reported. Nugent’s account of 1810 (42) is regarded as the original 
scientific observations for soufriéres on Montserrat and other islands. 
Earlier accounts of hot springs do appear, with the emphasis generally 
placed on their curative medical properties or the ability to coagulate the 
protein of eggs. Pére Labat (35), a French missionary to the Antilles 
between 1694 and 1705, recorded in six volumes his recollections of the 
Lesser Antilles. In 1696, he visited the Ance de Goyave in Guadeloupe and 
described the boiling fountains (fontaines boiillantes) in the bay, as well 
as in a neighboring swamp. He reported the water within a few feet of 
the coast to be warm enough to cook an egg held in his handkerchief. He 
attempted to determine the source of the heat and reported that, although 
the surface of the beach sand was without heat, when he dug to the depth 
of a foot he encountered sand and water too hot for his hand. This layer, 
he reported, smoked ‘‘comme on voit fumer la terre qui couvre le bois 
dont on fait le charbon.” The fumes of sulphur were nearly unbearable. 
In a neighboring swamp he found a lake approximately 45 feet in diameter 
which boiled at the edges and also more strongly but less frequently in 
the center. The quiet periods, Labat reported, were of a duration suffi- 
cient to allow one to say both a Pater and an Ave. Odors of sulphur were 
strong here, too, and a sulphurous taste was present in the water. Even 
today, previously unknown areas of prehistoric soufriére activity are dis- 
covered in remote places on most of the Lesser Antilles, and new out- 
breaks of fumarole activity have been recorded in the last decade on Nevis 
and Montserrat. In general, the most active areas are well known, and 
the locations of the principal soufriéres are to be indicated in summary 
for the various islands. 


TYPES OF VOLCANIC ACTIVITY 


Two basic types of volcanic activity occur in the Lesser Antilles. The 
most spectacular is the eruptive form so well recorded for Martinique 
and St. Vincent. Lava eruptions on these and other islands apparently 
occurred only in prehistoric times. The historic eruptions have been 
characterized by the forming of muées ardentes, plus ash fall and mud 
flows. Hill (20) has described Mt. Pelée as an ash pile, a description 
which applies equally well to the Soufriére on St. Vincent. 

Less spectacular, but of longer continuous duration in activity, is the 
fumarole. The fumarole is generally defined as a hole or vent (in or near 
a volcano) from which fumes are emitted. Subterranean as well as surface 
noise is present at most fumaroles. The gases produced may be saturated, 
ejecting with them large quantities of water, or they may be dry and 
either hot or cold. The dry, hot gases seem to condense atmospheric mois- 


1962 | HOWARD, VOLCANISM AND VEGETATION 283 


ture some distance from the orifice, producing steam or clouds of vapor, 
thereby giving fumarolic areas an eerie appearance. With one exception, 
the gases and liquids produced by fumaroles in the Lesser Antilles are 
acid in reaction. This acidity is responsible for the chemical alteration, 
the coloration, and the physical decomposition of the soil and rocks char- 
acteristic of the areas around fumaroles. 

A cluster of fumaroles is called a “soufriére,” although in the Lesser 
Antilles this term is loosely applied to solitary as well as clustered vents, 
to mountains, and even to towns near sulphur-producing vents. A fuma- 
role producing an odor of sulphur compounds is occasionally called a 
“solfatara” in this area. Technically, fumaroles are also classified accord- 
ing to their location as crateral fumaroles, when located in a volcanic 
crater, and noncrateral, when the point is lateral to a volcanic mass. 
Noncrateral fumaroles may be primary or secondary, depending on their 
association with the magmatic heat source. Representatives of all these 
types are to be found in the area. When the fumarole is depressed, sur- 
face waters may collect, and the dissolution of rock forms a mud pond 
or lake. Such bodies are also produced by the damming of valleys below 
fumarole areas by land slides, as in the case of the Boiling Lake in Domi- 
nica, Clear-water hot springs commonly found in alkaline areas elsewhere 
in the world have not been found in the Lesser Antilles. 


VOLCANIC ERUPTIONS AND THE VEGETATION 


The two active volcanoes on Martinique and St. Vincent are unique 
in producing only ash accumulations and a characteristic eruptive form 
termed nuées ardentes during the eruptions observed in modern times. 
The few lava flows or blocks recorded from the Lesser Antilles are of 
great age. 

The recent and observed volcanic activity in the Lesser Antilles began 
with earthquakes and noise, the emission of gas and steam, and the 
ejection of ash And boulders. Some of the steam was derived from the 
vaporization of a crater lake from the caldera, at or near the summit. 
In the eruptions of the Soufriére in St. Vincent in both 1812 and 1902, 
either an eruptive force or a break in the crater wall produced a cascade 
of boiling water which descended the slopes to the sea. While this boiling 
torrent followed the established valley patterns on the mountain slope, its 
heat, volume, and speed killed and removed the vegetation. During its 
descent, it scoured the river valleys clear of accumulations of ash which 
thickened the consistency of the water and led to the reports of a lava 
flow. Anderson (2), who studied the 1812 deposits, concluded that it was 
a mud flow of accumulated ash and not a true lava flow. 

The most destructive of the eruptive features of West Indian volcanoes 
have been the nuées ardentes. (See Figs. 1, 2.) The French seismologist 
Lacroix (36) proposed this term for the eruptions of Mt. Pelée and the 
Soufriére, and it has now become a standard term in all languages, al- 
though it is occasionally translated into English as “incandescent ash.” 


284 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


Fic. 1. Nuée ardente descending slopes of Mt. Pelée, April 13, 1931. Photo 
ei from Perret (44). Fic. 2. ee of nuée ardente on Mt. Pelee, 
1930. Photo reproduced from Perret (46). Fic. 3. Ash accumulation on Wallibu 
fields, St. Vincent, 1902. Photo reproduced ae Anderson and Flett (4). 


Smith (63) has described the phenomenon as a “strange black cloud, 
which, laden with hot dust, swept with terrific velocity down the moun- 
tainside, burying the country in hot sand, suffocating and burning all 
living creatures in its path, and devouring the rich vegetation of the hill 
with one burning blast.” Perret (46) explains its formation as follows: 
“Tn the acidic type [of volcano| its conduit normally closed because of the 
relative infusibility of the lava... the lava will have become sur- 
charged with gases rising slowly fron below, with possible assimilation 
from meteoric precipitation from above . . . the accumulated gas charge 
will, with eventual rupture of whatever restraint may have been imposed, 
go into paroxysmal explosive manifestation in the early stages of an erup- 
tion. A good instance of this is seen in the nuée ardente. . . . The viscous 
liquid has, through its own high gas content now coming out of solution 


1962 | HOWARD, VOLCANISM AND VEGETATION 285 


in distributed gas vesicles, become autoexplosive as a whole and thus 
capable, on release, of lifting itself . . . clear out of its pocket, in a 
stupendous en masse expansion. It will be evident that this will convert 
what had been a liquid into an infinitely subdivided mass; in a word, 
largely into a cloud of vapor and solidifying particles. Its great weight 
will precipitate this upon the mountain flanks in a down rushing ava- 
lanche of very heavy materials at high temperature, emitting gas from 
every pore amid rising clouds of ash, and constituting what has been 
termed nuée ardente. This phenomenon was not described prior to 1902, 
the date of the destruction of St. Pierre, Martinique. . . . The velocity 
of the onrushing avalanche is phenomenal, and this fact is not unrelated 
to the continuing evolution of gas. Vapor films between all sclid particles 
—from which particle the gas continues to be emitted during the ‘life’ 
of the nuée — <ffectively prevent all solid contacts, leaving the moving 
mass quite frictionless and capable of flowing upon the slightest inclines. 

. The original temperature (certainly on the order of more than 1,000 
degrees C.) tends to be maintained by this insulating atmosphere.”’ Per- 
ret also indicates that in a nuée ardente there is little if any free oxygen. 
As a result, no combustion occurs during its passage and materials are 
therefore carbonized and not burned. 

The effects of a nuée ardente are localized but lethal to the life of the 
narrow belt over which it progresses. Nuées ardentes have been photo- 
graphed from close range and have even been entered by volcanologists. 
Lava flows in Hawaii may leave little islands of vegetation (kipukas) un- 
disturbed but. surrounded with incandescent molten rock. Thus, too, in 
the case of a nude ardente, areas adjacent to the descending cloud may 
be unaffected by the passage of the ‘incandescent avalanche.” A study 
of the slopes of the Soufriére in St. Vincent today will show forests of 
massive trees cleft cleanly by the passage of a muée ardente. On one 
particular slope there are 50-80-foot trees with trunks 2 to 3 feet in 
diameter and within a few yards the ash and scoria of the nuée ardente 
still barely colonized after 60 years. The uée ardente carbonizes the 
vegetation in its path, while the residue of scoria left behind (in some 
cases 80 feet thick) retains its heat and kills the Te eae parts .of 
the plants and the seeds and fruits on the ground. (See F 

Quite in contrast. is the effect of the free fall of ash ic may precede 
or follow the nuge ardente. The volume of free ash is tremendous and the 
distance it may be carried by the wind difficult to believe. Anderson and 
Fleet (3, 4) cite reports of a fall of ash on ships 830 miles east-southeast 
of Barbados, following the eruption of the St. Vincent Soufriére. Ash fell 
on Barbados, 100 miles away, at the rate of 3.94 tons per acre. Nearly 
all of St. Vincent was dusted or submerged in free ash from the eruption. 

The term “volcanic ash” carries the popular connotation of a light- 
weight material. Perret states that this is not so, particularly in the ash 
derived from a nuége ardente. The crushing effect of the masses of ash is 
clearly documented, and the flattening effect on the vegetation implied. 
In the case of free-fall ash, the weight of individual particles may be less, 


286 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLIII 


but the cumulative weight equally great on any given area. Fields of 
arrowroot and other crops in St. Vincent buried under inches or feet 
of the cold free-fall ash did not recover. The accumulation of ash on 
leaves and branches caused defoliation and breakage of plants throughout 
the island. Although the size of the ash particles may have been small, 
the abrasive effect was noted in the puncture of leaves of crop plants and 
the removal of bark. A more modern report following the eruption of 
Paricutin in Mexico appropriately termed this “plutonic sandpaper” (47). 
The effects of the free ash on the vegetation is limited to abrasion, crush- 
ing, and smothering. The ash is generally chemically inert and without 
heat. The plants affected may recover or sprout from the base. Under- 
ground portions of the plants are not killed, and seeds germinate and 
may push up through the covering of ash. 

The days which followed the eruptions of Pelée and the Soufriére have 
been reported as stormy. Heavy rain showers occurred. Perhaps the ac- 
tivity of the volcanoes caused the formation of the heavy rain clouds; but 
in any case, violent rain storms swept the area, and exceptionally heavy 
rain descended on the ash-covered slopes. Russell (52) noted that “in- 
stead of being a protection to the surface on which it rests, the fresh 
debris is in many instances of assistance in its more rapid erosion. On 
steep slopes, and even when the surface is nearly level, the rills formed 
during the numerous tropical showers quickly cut through the loose sur- 
face material, and aided by the angular particles in suspension, corrode 
the soil or rocks beneath. The rains, as it seems, are heavier than usual, 
owing to two causes: first, the great amount of water contributed to the 
atmosphere as steam, and second, the vast amount of dust blown into 
the air, each particle of which serves as a center for condensation.’ River- 
beds filled with ash and scoria were swept clean, scoured to new depths 
and often altered from their previous courses. As the ash deposits were 
removed, the lower soil areas, too, were eroded in sheets of mud which 
were deposited along the coast in fans recorded as 70 to 80 feet wide and 
seven miles in length. Subsequent earthquakes caused landslides in shaken 
and loosened terrain. North of Chateaubelair in St. Vincent, Russell re- 
ported “strips of nearly flat alluvial land, adjacent to the sea, have dis- 
appeared leaving fresh bluffs of loose debris some 30 or 40 feet high.” 

The revegetation of areas affected by volcanic activity has been the 
basis for classic studies in the case of Krakatoa. For the West Indies, 
Perret (44, 46) and Anderson (2) made initial studies and Stehlé (66) 
and Beard (6, 7) some recent summaries. A more detailed study is still 
required, with consideration given to the specific nature of the disturbance 
in each area. Perret noted the occurrence of ‘several mosses and fungi” 
within two years of the 1930 eruption of Mt. Pelée in an area affected by 
a nuée ardente. Judging from his photographs, the “fungi” are lichens 
of the genus Stereocaulon. Perret also noted the invasion of the “golden 
back fern” and a plant with “tiny blue blossoms, said to be invariably 
yellow in other localities.” As both gold and silver forms of Pityro- 
gramma calomelanos and P. chrysophyila occur in the Lesser Antilles, 


1962 | HOWARD, VOLCANISM AND VEGETATION 287 


the scientific determination of plants cited in the geologists’ reports are 
not to be trusted. Perret also commented that while the golden form 
of the fern occurred at lower elevations, only the silver form was found 
at higher level on the disturbed areas. Although the plant with blue blos- 
soms can not be identified, Pitcairnia spicata is represented by the yellow- 
flowered var. sulphurea near craters and by the red-flowered typical form 
where volcanic fumes are not present. 

Both Anderson (2) and Sands (53) visited the St. Vincent Soufriere 
in 1907, five years after the last eruption, and noted the species which 
invaded the area. Although Beard cites Anderson’s paper, he quotes only 
from Sands’ less critical report in which 154 species of plants were col- 
lected from low-altitude areas badly devastated by the eruption. Ander- 
son, who was present during the eruptions, differentiates between areas 
affected by the nuée ardente and those affected by an ash fall. He states, 
“The incandescent avalanche swept down the Wallibu valley and spread 
out over the old fan or plateau at its mouth. it then turned south round 
the lower end of the Richmond Ridge and destroyed the Richmond Works 
and all the vegetation near them. The ash still remains to a depth of two 
to six feet in different parts, and the old roots are completely buried and 
thoroughly destroyed, but the avalanche was confined to the bottom of 
the valley and none of its effects are visible on either side. . . . The sur- 
face of the ash near the Works has not consolidated, but is rapidly break- 
ing up under the influence of plant roots, and humus is being formed. . . . 

e chief new plants are Castor Oil (Ricinus communis), which grows 
in luxuriant masses along and around the ruins of the Works, and a plant, 
Cattle Tongue (Pluchea odorata) which has already formed flourishing 
bushes taller than a man. Besides these, Indigo (Jndigofera anil), Sensi- 
tive Plant (Mimosa pudica), Guinea Grass (Panicum maximum), Eupa- 
torium odoratum, and two grasses were also noted.” Where the surface 
of the ash was consolidated, no vegetation can spring up; where it was 
broken by water or the hooves of animals the first invaders proved to be 
silver fern, grasses, and Pluchea. For areas covered with free fall ash, 
Anderson also related, ‘‘On the south slope leading up to the Morne Garu 
range the return has been considerable. The surface is still studded with 
the charred and bleached skeletons of trees, which appear to have been 
killed universally with the single exception of a small lateral valley north 
of the lower part of the Richmond Ridge, where a few palm trees in a 
sheltered position have recovered. The shrubs and herbaceous vegetation, 
which were all burnt level with the ground, are gradually returning, in 
many cases from the old roots, since the removal of the thin covering of 
ashes by the rain. 

“The top of the Wallibu plateau was entirely devastated. The trees 
remain only as bleached trunks except a few which have recovered in 
sheltered positions at the ends and south edge of the plateau. The dead 
trunks show that the ash was never more than a few feet thick at the 
most, and the whole is now covered with a luxuriant growth chiefly from 
the old roots. 


jan 


288 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLIII 


“At about 800 feet Tree Ferns (Cyathea arborea) become very abundant 
and large sheets of them are common, and generally it may be said that 
vegetation is luxuriant up to a height of about 1,000 feet, abundant up 
to 1,500, and very sparse above that height, with only a few grasses and 
silver ferns; higher up nothing but mosses and lichens are found. 

“At the lower lip of the crater and just inside it mosses and lichens 
only are found. The mosses have been identified at Kew as: Pogonatum 
tortile, P. Beauv. and Philonotis tenella, Jacq. and the lichen as Stereo- 
ee sp.’ 

Beard (6) presented a study of the altitude zonation of vegetation 
types in undisturbed areas and, in comparison, the types he found on 
the slope of the Soufriere in 1940, approximately 40 years after the erup- 
tion. Beard found at 2,400 feet altitude that the “plants are knee high 
only, still composed of silver fern with now very stunted Charianthus and 
Cyathea, Pitcairnia, Lobelia, ferns and grasses. 

“At 820 m. Charianthus and silver fern begin to thin out, and Cyathea 
is only 60 cm. high. Towards 900 m. the vegetation is simulating ‘paramo’ 
and is distinctly alpine in appearance. The ground is entirely and thickly 
covered with lichens, thallose, foliose and crustose. Higher plants are 
scattered and tufted and include the Lobelia and Pitcairnia, several grasses, 
a Lycopodium and a few ferns. Cyathea is now at most 30 cm. high, and 
the general vegetal covering is ankle deep only. 

“At the highest levels one finds a tundra only. The stony ground is 
thickly covered with lichens and, in between, a few clumps of Lodelia and 
Pitcairnia, The lichens vary greatly in colour, being predominantly grey 
or orange, and having a striking appearance at a little distance.” 

It is interesting that the inside walls of the crater are covered with larger 
plants, right down to the water line nearly 1,000 feet below. Here Bac- 
charis cotinifolia, Gleichenia, Lycopodium, ferns, grasses, and a few 
Cyathea occur in greater stature than at comparable levels on the outer 
slope of the crater. 

Stehleé (66) described the revegetation of the slopes of Mt. Pelée and 
also determined that the altitudinal zonation existing before the eruption 
was re-established afterwards. On both Mt. Pelée and the Soufriére it was 
apparent that the nuée ardente completely removed all plants and their 
propagulss, The ash fall, however, may have killed the es of plants 

ove the ground but allowed subterranean root and stem ystems, as 
well as seeds, to survive and grow. In both cases, the iene was 
more rapid at lower elevations where a weedy flora was the first type to 
be established. At all but the highest elevations, lateral invasion from the 
areas of vegetation unaffected by the nuée ardente was apparent. At the 
highest elevations where no unaffected areas were present and the soil 
was isolated both vertically and horizontally from seed sources which 
would tolerate those environmental conditions, the reinvasion and _ re- 
establishment was slow, and often an unexpected composition was cre- 
ated. Lichens and mosses were the first and most successful invaders. 
Wind-blown spores of ferns were introduced, and the plants created pure 


1962] HOWARD, VOLCANISM AND VEGETATION 289 


stands of many individuals. Although spores of Lycopodium were eventu- 
ally introduced, only scattered plants were found. The seeds of Pitcazrnia, 
Cladium, and various grasses could have been distributed by wind to 
these new areas. Stehlé suggests that the fruits of Charianthus, Lisianthus, 
Lobelia, Tibouchina, Ilex, Weinmannia, and Clusia are distributed by 
birds or bats, and these plants are slow to become established in the 
devastated areas of high elevation. At lower altitudes birds, cattle, and 
man have played a role in starting new plants. Stehlé, for example, no- 
ticed the invasion of sugar cane on the slopes of Martinique and suggested 
that a wind-blown seed allowed this plant to be established on the scoria. 
Perhaps a node dropped by a wandering cane-chewing sightseer might be 
a more logical interpretation. The seeds of Acacia and Mimosa observed 
by Sands, Perret, and others were spread by cattle wandering over the 
newer ash accumulations 

It is of some interest. to note that in all cases the growth of invading 
species appeared to be more rapid inside the crater of the volcano than 
at comparable levels on the outside. This growth can probably be asso- 
ciated with the greater protection afforded within the crater walls, as 
well as the more abundant moisture available on the less porous soil 
surface. 


FUMAROLES AND THEIR EFFECTS 


The occurrence of fumaroles, either singly or clustered as soufriéres, can 
be reported for all the major islands of the Lesser Antilles from Grenada, 
in the south, to St. Kitts, in the north. The fumaroles are found at many 
elevations — submarine near Nevis, coastal on Montserrat, at low eleva- 
tion on Grenada, and at the summit of the Soufriére (4800 feet) on 
Guadeloupe. The fumaroles are present in many vegetational types, in- 
cluding the thorn scrub of Grenada, old field vegetation on Dominica, 
mesophytic forests on Montserrat, and the volcanic facies on Guadeloupe. 
Active fumaroles can be determined from some distance by the production 
of heat, steam, and fumes laden with sulphur compounds. Near by, they 
show a characteristic destruction or absence of adjacent vegetation, if 
the fumes contain sulphur compounds, or an equally characteristic altera- 
tion of the rock texture and color. Extinct or dormant fumaroles may be 
recognized in open areas where mosses, algae, or perhaps a few higher 
plants cover the surface or where the softened rock in a restricted area 
reveals the former fumarole and its activity. 

The activity of the fumarole may vary considerably in intensity over a 
long period of time or even in relation to the annual rainy or dry seasons. 
Some recurrence of fumarolic activity has been noted and associated with 
active volcanism on adjacent islands or with earthquakes of considerable 
geographic latitude. Within the Lesser Antilles are dormant fumaroles, 
those with long records of continuous activity, new ones, and those sporadic 
in their action. 

Nearly all fumaroles are characterized by the emission of heat. Stehlé 


290 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


(67) gives temperatures of the crateral fumaroles of Mt. Pelée as high as 
700° C. Robson and Willmore (50) have published the most recent 
studies of heat emission from fumaroles in the Lesser Antilles. They have 
suggested as a base point the temperature of 24° C. for meteoric water 
or ordinary cold springs and for the air temperature. Steam temperatures 
from Ie Qualibou soufriére in St. Lucia have been measured as high 
as 185° C., while water outilow temperatures from ponds in the same 
soufriére varied from 63° t On Montserrat, Robson and Will- 
more recorded the flow of water from Gages Upper Soufriére at tempera- 
tures of 34-86° C. from three sources and at Galway’s Soufriére from 

© to 97° C. from two sources. Crater lakes, which may not show vio- 
lent actions typical of fumaroles may show the effects of their proximity 
to sites of volcanism. The mean lake temperatures for the crater lake in 
the Soufriére of St. Vincent was 27.1° C., indicating at least a 4° rise in 
temperature due to the heating effect of submerged fumaroles. There is 
only one record of plant life in the boiling lakes of the Lesser Antilles, 
but this has never been verified in more recent observations. No animal 
life has been reported. Crater lakes, by contrast, have an algal or higher 
plant flora, and the crater lake on St. Vincent’s Soufriére has recently 
been stocked with fish. The heat of water flowing from fumaroles or boil- 
ing lakes is soon dissipated. Blue-green and green algae, diatoms, and 
even sedges have been collected from outflows with temperatures of 56° 
to 85° C. within five feet of vents on several islands. Unfortunately, no 
identifications have ever been received from specialists who might name 
these collections of plants properly. 

The direct emission of steam could have a noticeable destructive effect 
on the surrounding vegetation. Few plants could stand blasts of tempera- 
tures of 100° to 185° C. However, the direct blast of steam heat prob- 
ably is of minor aoe or none at all in comparison with soil heat, 
toxic gases, and cumulative acidity. Although MacGregor (39) gives in 
tabular form records of heat from soufriéres on Montserrat, he also points 
out that the maximum temperature recorded depends on how far the 
thermometer can be pushed down the orifice against the gas pressure. 
Temperatures are higher below than at the surface, and such heat as is 
emitted is dissipated rapidly in the atmosphere. 

For many of the West Indian fumaroles there appear to be few if any 
toxic substances included in the air being forced out at the present time. 
Around the fumaroles of Wotten Waven, in Dominica, the vegetation, in- 
stead of being adversely affected or killed, appears to grow more luxuri- 
antly in the local area of higher heat and humidity. However, in many 
fumaroles there must be toxic substances, for the fumes have a definite 
effect on the vegetation. MacGregor and other authors have stated that 
the fumes kill the local vegetation. Even the names of the fumaroles and 
their areas suggest the toxicity. The largest soufriére on St. Lucia is known 
as ‘“Qualibou,” a name meaning “place of death,” and some soufriéres in 
Dominica are located in the “Valley of Desolation,” It is clear that the 
fumes are often noxious and toxic to men and to plants. The death of 


1962 | HOWARD, VOLCANISM AND VEGETATION 291 


two visitors to the Boiling Lake of Dominica has been substantiated. The 
geologist Perret was overcome by fumes while residing in his field station 
within Gages Lower Soufriére, and workmen on an earlier project in the 
same area were overcome. Although several geologists have attempted to 
collect and analyze the gases, the results are inconclusive. Romer (51) 
has reported the fumes from fumaroles on Mt. Pelée to contain only car- 
bon dioxide, sulphur dioxide, and traces of hydrogen chloride; however, 
some fumaroles on Montserrat failed to give positive tests for carbon 

xide. Obviously the composition of the gases may vary with the nature 
a the activity of the fumaroles. Most fumaroles of the Lesser Antilles 
produce hydrogen sulphide, but all characteristically reveal an absence of 
fluorine, relatively few chlorides, and negligible amounts of boron. Perret 
described the gas of Gages Lower Soufriére and concluded it was ‘“‘a gas 
having a traitorously pleasant odor, but extremely irritating to the eyes 
and respiratory passages. . It was highly unstable and decomposed 
into hydrogen sulphide and siphur. It may have been one of the per- 
sulfides of hydrogen.” Willmore (71) stated, “These vents emit a mix- 
ture of steam and sulphur gases which, during quiet periods, usually have 
a temperature a little below that of boiling water. During periods of 
abnormal activity the temperature of the exhalations often rises, and the 
predominant sulphur constituent may change from hydrogen sulfide to 
sulphur dioxide. — The tendency for the temperature to become stabilized 
near boiling point is readily explained on the assumption that the sou- 
friére is supplied by hot juvenile gases which bubble through meteoric 
water on their way to the surface. The change in gaseous constitution 
during active periods is presumed to occur because the original magmatic 
gases undergo reactions on their way to the surface, so that the enc 
products depend on the temperature and velocity of the flow.” The a 
sence of fluorides and the relatively small amounts of chlorides and boron 
support the conclusion that meteoric and not juvenile water is involved 
in most soufriéres of the Lesser Antilles, at least most of the time. 

The destruction of the vegetation around fumaroles, therefore, is ap- 
parently not caused by the chemical elements of the fumes or the water 
but by their nature as compounds. The various sulphur compounds in 
the presence of water can produce a series of acids. Litmus papers and 
“pHydrion” papers respond quickly, giving acid reactions. [Perret re- 
ported that during the seismic crisis on Montserrat egg shells dropped 
in the fumarolic waters give rapid effervescence. Another mildly acid con- 
stituent of the fumarole fumes is hydrogen sulphide. The presence of 
this compound is evident, not only in the odor, but in the tarnishing of 
all silver brought into the vicinity. The town of Soufriére in St. Lucia 
is unfortunately in the lee of the Qualibou soufriére, and one is continu- 
ously aware of the derivation of the name of the town. Silverware tar- 
nishes in the town of Plymouth in Montserrat which is downwind from 
the Gages soufriéres. Perret depended on the reaction of lead-acetate 
paper to sulphur to record the intensity of such emission from the Gages 


— 


292 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


soufriéres, and during the same seismic crises even the lead paint black- 
ened on ships in the roadstead. 

The acids derived from the various sulphur compounds also affect the 
soil and rocks in the vicinity of the fumaroles. MacGregor (38, 39) con- 
cluded that ‘mineral changes brought about in the country rock render it 
locally pale to pure white in color.” The pale color of soufriére areas 
renders them conspicuous and easily recognizable from a considerable dis- 
tance. Accompanying the color change in the rock is a loss of structural 
hardness. Rocks in fumarole areas may retain their original shape and 
texture but become soft and nearly putty-like in consistency. The novice 
learns quickly of the treacherous footing in areas of fumaroles, where 
even the largest stones are not to be trusted. 

When both hydrogen sulphide and sulphur dioxide are present, a chemi- 
cal reaction may take place oxidizing the former and reducing the latter 
to produce water and crystalline sulphur. The deposits of sulphur crystals 
around the vents of Lesser Antillean fumaroles are colorful. Local needs 
for sulphur are easily met and at times small quantities of sulphur have 
been exported from the area. The commercial extraction of sulphur as 
a by-product of the use of steam for thermal power has been proposed 


(8). 


EFFECTS OF FUMAROLES ON VEGETATION 


During a visit to many of the fumaroles in the Lesser Antilles, it be- 
came apparent that there were areas around most fumaroles where no 
vegetation survived; that leeward from the vents certain plants were 
affected or killed by the emissions of the vent, while to the windward 
the same species approached or persisted much more closely; that certain 
species tolerate the effects of the fumaroles longer than others; that cer- 
tain species invaded an affected area more readily than others; that some 
physiological change took place in the species which did grow success- 
fully near the fumaroles in comparison with the same species elsewhere; 
and that the effects of the fumaroles on the vegetation were greater at 
certain periods than others. A review of the literature reveals that the 
earliest descriptions of the Boiling Lake on Dominica do not vary from 
observations made recently, and that Sapper’s sketch (57) of the limit of 
devastation of vegetation at Galway’s Soufriére on Montserrat differs little 
from a sketch map made last year. As long as the fumarole remains ac- 
tive and reasonably constant the effects on the vegetation are noticeable 
to a distance limited primarily by the terrain. Furthermore, the chemi- 
cally altered rock and soil near the fumarole remain an inhospitable area 
for plants for decades, and revegetation is an extremely slow process. 

owever, new vents do open and I have had the opportunity of record- 
ing a sequence in the change of the vegetation near a new fumarole over 
a ten-year period on the island of Montserrat. In 1950, I visited both 
Gages Lower Soufriére, studied by Perret in 1934-37, and Gages Upper 
Soufriére, which was reactivated by an earthquake in 1937 when Gages 


1962] HOWARD, VOLCANISM AND VEGETATION 293 


Lower Soufriére was dormant. Gages Upper Soufriére was active, but the 
vegetation around it appeared to have been stabilized in relation to the 
vents. Photographs were taken at both soufriéres, notes were made, and 
specimens were collected approaching and adjacent to the devastated area. 
I returned to the area in 1961 expecting to determine the identity and the 
role of the plants which had successfully invaded during the past decade. 
At Gages Lower Soufriére I was not disappointed. Several four-foot plants 
of Clusia alba, as well as other weeds, were growing in the center of the 
ruins of Perret’s field station. Young plants of Andropogon, Philodendron 
giganteum, and Pitcairnia were seen, and a coating of mosses colored 
many of the rocks. 

The situation at Gages Upper Soufriére was another matter. New 
vents had opened in a thicket downstream from the previously affected 
area, affording an opportunity of determining the initial effects of a 
fumarole on previously undisturbed vegetation. The exact date of the 
inception of this vent is not known. Martin-Kaye (40) observed the 
area affected by Gages Upper Soufriére in 1954. A marker tree, a large 
buttressed rooted specimen of Sloanea massonii from which I had col- 
lected a specimen in 1950 was now dead, and only the stump remained 
some 50 feet inside the devastated area (Fig. 13). A vent had opened 
in the last decade directly under that tree, the base of which was now 
discolored white and yellow. The remaining stump was hot from the 
steam and wet and rotted, so that an 18-inch machete penetrated easily 
to its hilt, while “pHydrion” paper pressed against the sodden wood 
showed a pH of 1.5. The activity of the Gages Upper Soufriére had not 
only been extended downstream but up the hillside nearly to the ridge 
(Fig. 12). It apparently will be only a short time before the fumarole 
activity breaks through the crest of the ridge and effects a new drainage 
system. 

In the newly affected area readings of temperature and the pH were 
taken on occasions between 8 A.M. and 5 P.M. over a period of two weeks 
with no significant variation in the figures. The vent of greatest interest 
(Fig. 18) was in a thicket composed of species of Blechnum, Charianthus, 
Clusia, Cyathea, Ficus, Ilex, Miconia, Palicourea, Philodendron, and 
Symplocos. No noticeable hole or pore had developed, and the steam 
appeared to be issuing from an area of dry soil about four inches in 
diameter. Outside of this small focal point some organic material was 
present but this was extremely dry. The diameter of the area of dry 
duff was approximately four feet. Outside of this area the debris on the 
soil surface was noticeably warm and moist, and dead and dying plants 
were obvious and readily identifiable. The effects of this new vent ex- 
tended for a distance of only 20 feet. The temperature reading in the 
vent area was 94-96° C., a measurement obtained by inserting to a depth 
of three inches a laboratory thermometer calibrated to 250° C. In the 
area of dry and barren soil around the vent the temperature was 84° C. 
at a depth of three inches. The most tolerant plants in the area of wet, 
warm soil nearest the vent were Blechnum, Clusia, and Philodendron. 


294 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


Beneath these plants the soil temperature was 54° C. Progressing out- 
ward on several radii, it was determined that species of Ficus and Lyco- 
Hey grew in soils at 48° C., while Miconia and Palicourea withstood 
sevond this point the soil temperature dropped to 30° C., and 
no ice was evident. The dead or nearly dead plants in soil temperatures 
of 40° C. and above were Charianthus, Cyathea, Ilex, Miconia, Palicourea, 
and Symplocos. About 100 yards away from this vent in the undisturbed 
forest, soil and air temperatures varied between 24° and 26° C. at dif- 
ferent times of the day. It is apparent that one previously unrecognized 
factor in the death of plants around new fumaroles is the heating effect 
of the soil. Although the fumes from this small vent did discolor lead 
acetate paper they gave no reading on “pHydrion” paper. Litmus paper 
gave an acid reaction very slowly. The same papers used on the moist 
soil zone where plants had been killed gave readings of pH 1-1.5. The 
vegetable material present in the dry zone crumbled easily in my fingers, 
That of the moist zone, by contrast, was slippery or almost mucilaginous. 
The structure of the duff had been altered, and I judged from the ease 
with which my feet slipped that a heavy shower of rain would cause 
severe erosion. The areas unaffected by heat or low pH were more fibrous 
to the touch and more resistant under foot. Readings taken in the vege- 
table material in the unaffected forest showed pH 4—4.2. Hardy, Rod- 
rigues, and Nanton (19) report that the majority of agricultural soils on 
Montserrat range from a pH of 6.1 to 7.4. Their two sets of figures for 
forest sites indicate pH readings of 4.3—4.8 (for lower montane rain forest 
soil on Frenchman's Hill) to a pH of 5.2—6.6 (for soils in a palm brake 
on Chance’s Mountain). The initial killing effect of fumaroles may well 
be, first, soil temperatures beyond the range of tolerance of many species 
and, secondly, high acidity. 

General observations were made on the vegetation within a few yards 
of the older area of devastation around Gages Upper Soutfriére. Although 
the heat of the steam emissions could be felt in this area, it was not 
oppressive, and temperature readings were only a degree or two above 
those recorded deeper in the forest. The odor of hydrogen sulfide was 
noticeable, as was a higher humidity. Wet litmus paper did not indicate 
an acid reaction during a 30-minute exposure. Nevertheless, plants were 
dying and a slowly cumulative toxic effect of the fumes is the only ex- 
planation. The genera most susceptible appeared to be Cyathea, Fr eziera, 
Hirtella, Licania, Palicourea, Peperomia, Philodendron (except P. gigan- 
teum), Psychotria, Symplocos and Weinmannia. Species of Gonzalagunia, 
Hex, and Sloanea appeared more tolerant and those of Clusia and Tern- 
stroemia, as well as Philodendron giganteum, most resistant. 

The oldest areas of Gages Upper Soufriére showed scattered plants of 
Clusia alba, Cyperus ligularis, Philodendron giganteum, and Pitcairnia 
spicata, It is not clear whether these plants were residual from the former 
vegetation or more recent invaders. They grew both as individuals and 
in groups. In numbers they were more abundant on the fumarole affected 
soil than in comparable areas of the undisturbed forests. Vents of two 


1962 | HOWARD, VOLCANISM AND VEGETATION 295 


types were present in the areas where these plants grew. One type issued 
only fumes while the other had steam so laden with moisture that it con- 
densed around the orifice and ran off, forming a small stream. The air 
issuing from the dry vents was recorded as high as 93° C., but the soil 
temperature immediately around the vent was 37° C. Incrustations of 
crystalline sulphur a such orifices. The moist steam vents, however, 
gave readings of 37° to 70° C. taken in a comparable manner, and the 
“pHydrion” paper gave a reading of pH 1.5. Barren soil temperatures 
around these vents were 30° C. Temperatures recorded from around 
the root systems of Philodendron and Pitcairnia were 32° C. and the 

was 3. The highest water temperature recorded in any of the small pools 
within Gages Upper Soufriere was 94 

The Qualibou soufriere in St. Lucia alowed an example of vegetation 
stabilized to the long-term effects of fumes from eee This sou- 
friére is located in a valley with a high hill on the windward side so that 
fumes from the fumaroles tend to rise along the hill face. The vegeta- 
tion of this hillside consisted of stunted low plants of Blechnum, Clusia, 
Cyperus, Gleichenia, Lycopodium, and Pityrogramma. Just over the crest, 
in an area where the fumes were dissipated by the prevailing winds, the 
native forest flourished in an unaffected manner. The species which tol- 
erated the fumes on the hillside were those with heavy cuticles, for the 
wax-rigid nature of the foliage was conspicuous in contrast to that of 
the same species growing in an area where they were unaffected by the 
fumes. The waxy coating on the leaves apparently made this vegetation 
more Susceptible to burning, for a large fire scar dominated the face of 
th n a few places a clear line of demarcation was evident between 
the plants which burned and those which only wilted down. This de- 
marcation appeared to be associated with the plants exposed to the fumes 
and those protected from it. 

An example of the periodicity of activity of fumaroles and the killing 
effect on the vegetation can be found on the slopes of the Soufriére on 
Guadeloupe. Stehlé (65) has described the units of vegetation which 

ave been established at successive altitudes to the summit. His “Clusi- 
etum,” ‘“‘Lobelietum,” ‘“Pitcairnietum” represent, in part, tolerance of the 
component species to temperature, wind, and cloud cover but also to the 
pH of the ash terrain and to the effects of the fumaroles at the summit. 
During periods of little fumarole activity, the shrubs and trees may de- 
velop normally. When the fumaroles become active, the acid fumes will 
kill many of the branches and occasionally entire plants. Generally these 
plants recover by sprouting from the lower branches or the base, but 
the upper branches remain dead and leafless. 

A unique fumarole area, perhaps more properly termed a hot springs, 
is found in Chambourg section of St. David’s parish in northern Grenada. 
The twelve small bubbling hot springs apparently are influenced by the 
old volcanic mass below the crater lake, Lake Antoine. The soils appear 
to be coral limestone and a sandstone which includes volcanic ash. Dur- 
ing certain seasons the waters of the springs overflow and several sizeable 


296 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


cones have been created. An alkaline litmus reaction was obtained from 
the waters collected in overflow pools. This, to my knowledge, is the only 
alkaline fumarole in the Lesser Antilles. No plants from the surrounding 
scrub of Acacia, Acrostichum, Avicennia, Chrysobalanus, and Laguncularia 
have encroached on the rock areas built up by the hot spring. 

The Grenada hot-spring area is the only one in the Lesser Antilles ven- 
erated by the local people. One vent had an animal sacrifice placed there 
by religious cultists. Several springs had fresh or wilted bunches of flowers 
in the orifices. Candles and coatings of wax were present in the largest 
vent, along with a painting of the Sacred Heart. The late Father Bryan 
Proudman, who was my host in this area attributes the tokens to “‘shandoo”’ 
ceremonies. 


CATALOGUE OF THE FUMAROLE AREAS OF THE 
LESSER ANTILLES 


Grenada 


THe GraNnp Etanc. This crater lake is in the caldera of an old vol- 
cano. Beard (7) has described and illustrated this area and its vegeta- 
tion. The lake temperature indicates the presence of some heat source. 
Sulphur odors are noticeable at times, and particles of sulphur occasion- 
ally appear on the surface of the lake. No effect of the heat or acidity can 
be noticed on the surrounding vegetation. 

LAveRA Hot Sprincs. (Figs. 4, 5.) About twelve small vents or pot 
holes near Lake Antoine produce an occasional weak odor of hydrogen 
sulfide. These are the only alkaline reacting springs that I have encoun- 
tered. The vegetation has not encroached on the sheets of hard rock 
formed by the overflow from these springs. 


St. Vincent 


THE SOUFRIERE. Interest in the eruptive volcano on St. Vincent centers 
on the accuracy of the reports of its eruption in 1902. Prior to that time, 
eruptions have been recorded for 1718 and 1812. Reports of the former 
eruption have been considered by Anderson and Flett (4). They point 
out that while Huggins (32) discredits such reports, Defoe (37) only in 
part exaggerates in his story, which may indeed have been based on fact. 
The letters of Rev. Smith (64) may be considered corroborative evidence 
that the Soufriére did erupt in 1718. 

Mr. James Anderson apparently was the first white man to ascend the 
volcano. In an account published in 1785 (1) he described the summit. 
“We found very little obstruction in our way up until we got to the place 
where I returned, and there, for about a quarter of a mile, we had con- 
siderable difficulty to clear our way through grass and ferns. After we 
came within a quarter of a mile from the top, we found ourselves in an- 
other climate all at once, the air very cold, and the vegetable productions 
changed; here was nothing but barrenness over the whole summit of the 


1962] HOWARD, VOLCANISM AND VEGETATION 297 


mountain. On the confines of the grassy region and the barren I found 
some beautiful plants. Moss grows here in such plenty that I frequently 
sunk up to my knees in it. This is the only place in the West Indies that 
produced any moss that I have seen. About noon we gained the top of 
the peak . . . when in an instant, we were surprised with one of the 
grandest and most awful scenes I had ever beheld.” For the crater An- 
derson reported, “In the centre of the bottom is a burning mountain, of 
about a mile in circumference, of a conic form, but quite level. . From 
the external appearance of this mountain, I imagine it has only Perce to 
burn lately, as on several parts of it I saw small shrubs and grass, which 
looked as if they had been scorched and burnt. ... On ae opposite 
sides of the burning mountain, . . . are two lakes of water.’ 

An anonymous account published in the Evening News of July 30th, 
1812, is reprinted by Anderson and Flett (4). This account described an 
ascent of the Soufriére on April 26th, only a few hours before its eruption 
at noon on the 27th. On viewing the crater, the anonymous report stated, 
“Exactly in the centre of this capacious bowl rose a conical hill, about 
250 or 300 feet in height, and about 200 in diameter, richly covered and 
variegated with shrubs, brushwood, and vines about half-way up, and for 
the remainder powdered over with virgin sulphur to the top. . . . The 
precipitous sides of this magnificent ampitheatre were fringed with various 
evergreens and aromatic shrubs, flowers, and many alpine plants. On 
the north and south sides of the base of the cone were two pieces of water, 
one pure and tasteless, the other strongly impregnated with sulphur and 
alum.”’ A comparison of the two reports indicates a secession of the ac- 
tivity of the cone and a re-establishment of the vegetation. The eruption 
changed all this, for as Shephard (62) stated, ‘all the former beauty of 
the Soufriére was, of course destroyed; the conical mount disappeared, 
and an extensive lake of yellow-coloured water, whose agitated waves 
perpetually threw up vast quantities of black sand, supplied its place.” 
Hooper’s account in 1886 (24) suggests that a single lake remained in 
the crater. Anderson and Flett (4) reported that the crater “contained 
three small lakes of water, greenish and turbid, that in the southeast cor- 
ner was throwing up jets of mud and steam with a hissing noise.” In the 
years which followed, an accumulation of rain water apparently raised 
the surface of the three lakes and made them confluent. Beard (6) found 
only one lake in the crater during his visit in 1940, and such is the situa- 
tion today. The southeast corner of the lake, however, is often discol- 
ored with suspended material and flakes of sulphur, indicating that all 
is not quiet in the fumarole area. 

Beard has described the progress of plant succession on the Soufriére in 
a paper published in 1945. A new study, now that an additional two 
decades have passed, appears to be in order. I visited the summit of the 
Soufriére in 1950 and found sufficient differences from Beard’s description 
to suggest that the initial invasion of plants had been successful, that a 
favorable environment had been created, and that rapid changes were 
then taking place. At that time, the floral composition of the summit was 


298 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLIII 


closer to that described by Hooper prior to the 1902 eruption, with the 
sole exception of an abundance of fleshy liverworts unmentioned by either 
Hooper or Beard. 


St. Lucia 

THE QUALIBOU SOUFRIERE. (Figs. 20, 21.) This appears to be the 
only active soufriere on the island of St. Lucia and is situated about a 
mile and a half southeast of the town of Soufriére. Hooper visited the 
area in 1SS6 and reported briefly that “the exhalation . . . has reduced 
the vegetation on both sides within reach to a carpet of brake fern.” In 
1903, Sapper (56) and, in 1904, Hovey (26) described the area as between 
two and three acres in extent, but neither made reference to the vege- 
tation. In 1949, Beard (7) noted that a specialized vegetation was found 
around the Sulphur Springs (the Qualibou vents). “Clumps of Pitcairnia 
are as usual found nearest to the vents with some Cyperus ligularis. Far- 
ther out, these are joined by various species of ferns, principally Blech- 
num serrulatum dominated by large clumped bushes of Clusia plukenetii 
and C. alba. The whole hillside above the Springs is carpeted with Blech- 
num ferns with an occasional Clusia.” 

The vegetation around the Qualibou soufriére appears to be com- 
pletely stabilized. Algae are found in dense mats in the drainage of the 
hot springs. Cyperus and Eleocharis plants are common in shallow warm 
pools of seepage areas. None of the larger plants seen in the area showed 
any damage from the constant emissions of the fumaroles. 

Recent interest has centered in the possible utilization of the thermal 
heat for generating electricity. The reports of Bodvarsson (8) and of 
Robson and Willmore (49) are pertinent to this problem. 


Martinique 

Mr. PeLveée. The destructive eruption of Mt. Pelée in 1902 eliminated 
the town of Saint Pierre, killing all but one person. The latter stages of 
this eruption, as well as that which occurred in 1930, are well described 
by numerous authors (2, 3, 29, 36, 39, 44, 52). Bibliographies of signifi- 
cant publications are given by Hovey ( (28) and Russell (52). Few specific 
data are available on the regrowth of the vegetation, and a modern survey 
seems desirable. Stehlé visited the area and published in 1938 a general 
consideration of the regrowth (66). He recognized conditions comparable 
to those of Guadeloupe, excepting the sulphur fumes, and suggested that 
the vegetational zones were a ‘Clusietum martinicense,” a ‘*Lobelietum 
martinicense,’ and a “Pitcairnietum martinicense.” Stehlé attributed to 
larger animals the introduction of Mimosa pigra and Acacia tortuosa to 
the areas of recent lavas. Birds, he felt, accounted for the introduction of 
Ficus, Melia, Trema, Tetrazvgia discolor, and Miconia striata. At the 
lowest elevations on new lavas, Stehlé found common weedy species of 
Crotalaria, Cassia, Emilia, Fuphorbia, Bidens, Ageratum, Indigofera, 
Pluchea, etc. Slightly higher and probably introduced by wind were plants 


1962 | HOWARD, VOLCANISM AND VEGETATION 299 


of Ochroma pyramidale, Ceiba antillana, and Tecoma stans. Near the 
top of Mt. Pelée, Stehlé reports the occurrence of two species of Des- 
modium and single species each of Stenotaphrum, Erigeron, Sauvagesia, 
Mikania, Arthrostemma, Hypoxis, Cuphea, Rubus, Phenax, and Cyperus. 
The majority of these are not found at higher altitudes in undisturbed 
vegetation on Martinique or nearby islands. No information is available 
on the length of their persistence. Stehlé indicated that the most power- 
ful colonizers were Pitcairnia bracteata, Guzmania plumieri, Tibouchina 
chamaecistus, Baccharis cotinifolia, Pityrogramma calomelanos, and two 
species of Lpidendrum. 

Thermal springs are recorded from the vicinities of the towns of Frégate 
and La Redoute, along the coast near Marigot, and on the siepes of Mt. 
Pelée above Macouba. I have not located descriptive literature regarding 
their activity as fumaroles or the vegetation around them. 


Dominica 


The earliest record of fumaroles on Dominica appears in Atwood’s ac- 
count of the island pend in 1791 (5). His vivid, but questionably 
accurate description stated, ‘“‘These sulphurous mountains are certainly 
among the most wonderful phaenomena of nature, and command our 
astonishment and admiration. To see vast tracts of land on fire, whose 
smoke, like clouds, stretches far around: brimstone in flames, like streams 
of water issuing from the sides of precipices; in the vallies large holes 
full of bituminous matter, boiling and bubbling like a caldron; the earth 
trembling under the tread, and bursting out with loud explosions, are 
objects truly terrefic to the beholder; who, on the spot, are struck with 
awe and admirations, on viewing such dreadful works of the Almighty, 
who causes them to exist, for purposes only known by him.” Edwards 
(15) also reported about Dominica that ‘several of the mountains 
contain ee volcanoes, which frequently discharge vast quan- 
tities of burning sulpht ‘rom these mountains also issue springs of 
ot water. ...° There are no reports of eruptions on Dominica in his- 
toric Himes! and these two early descriptions can not be definitely located 
geographically. It is possible that these both refer to the imposing Val- 
ley of Desolation and its famous Boiling Lake, and represent its earliest 
record. The discovery of the Boiling Lake is generally credited to Dr. 
H. A. Nicholls around 1880. Nicholls’ route was soon followed by Ober 
(43), Johow (33), and others, all of whom comment in one way or an- 
other on the nature and composition of the vegetation in the Valley of 
Desolation and around the Boiling Lake. 

Hovey (30) and Earle (14) were the first to consider the geology of 
the area. Later in the twentieth century Domin (13a) and Hodge (21, 
22) studied the vegetation of Dominica, and both wrote extensively on 
this subject. Hodge has said of the Valley of Desolation, ‘““The combined 
sulphurous fumes of all the vents and fumaroles have had a blighting 
effect far up the slopes, wherever they have come against the vegetation. 


= 


300 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


The end result has been the formation of what is known locally as the 
Valley of Desolation.’ [See Fig. 6.| Along the ridges above lies Elfin 
woodland but in the valley only patches of it remain and in these Clusia 
venosa and an /lex seem best able to tolerate the noxious gases. In the 
proximity of the fumaroles vegetation is absent or sparse and except for 
the colorful algae to be seen in the warm water of the streams, is appar- 
ently unable either to tolerate the noxious fumes or to root in the mineral- 
impregnated soil. The only pioneer species observed close to the vents 
were the bromeliad Pitcairnia spicata var. sulphurea, a grass, Ischaemum 
latifolium, and occasional plants of Dicranopteris bifida.” Hovey (30), 
Sapper (56), and Robson and Willmore (51) give data on the water 
temperatures of the Boiling Lake. 

Earle (14) reported additional soufriéres in Dominica at Ravine D’Or 
(Layou Valley), Morne au Diable (Portsmouth), Laudat and Soufriére 
villages, and Wotten Waven. He states, “All of these are insignificant 
besides the Grande Soufriére, though the Wotten Waven springs are quite 
numerous and besides a few violent fumaroles also display small mud 
volcanoes.’ 

At Wotten Waven I was able to visit the fumarole area which is located 
in an old field now devoted largely to pasture. The fumaroles at the time 
of my visit were noisy and emitting steam, but almost without sulphurous 
odors. The high humidity and heat appeared to encourage plant growth, 
and the shrubs of Clusia alba were more luxuriant than specimens we had 
passed a few yards away (Fig. 7.) 

The fumarole area behind Soufriére village was represented by an enor- 
mous scar on the hillside where landslides had recently exposed fresh sur- 
faces and obscured some of the vents. The fumaroles were not noticeably 
active, and although the odors of sulphur dominated the air, there was 
no evident destruction of the vegetation. The area was well stabilized, but 
few plants other than species of Blechnum, Dicranopteris, and Cyperus 
had invaded the areas. 


Guadeloupe 


THERMAL SPRINGS. The island of Guadeloupe is reported to have ther- 
mal springs along the coast at the towns of Boiuillante, Monchy, and 
Pigeon; somewhat inland near La Lise and Dolé; on the slopes of the 
Soufriére near Baines Jaunes; and at the summit of the Soufriére. All 
of these are on Basse Terre and probably are associated with the volcanic 
massif of the Soufriére. I have not visited the thermal springs at low 
altitude and have been unable to find any descriptive literature regarding 
them or the vegetation surrounding them. 

The oldest reference to hot springs and fumaroles is found in Pére 
Labat’s writings. His description of the thermal springs of “Ance de 
Goyave”’ presumably refer to the area currently called Boiillante and not 
to Goyave on the west coast. Labat also climbed the Soufriére and de- 
scribed the stunted vegetation near the summit. A trip from St. Claude 


1962] HOWARD, VOLCANISM AND VEGETATION 301 


to Baines Jaunes and along the old carriage trails past the warm and 
cold baths to the summit of the Soufriére er that few changes have 
occurred in the intervening 260 years (Figs. 8, 9). 

Stehlé (65, 67, 68) has published several ‘admirable treatments of the 
vegetation of the slope of the Soufriére. He indicates that the effects of 
high altitude and of the sulphur vapors cause a limitation in height of 
plants but seem to encourage the development of many lateral branches. 
He also states, “Les gaz sulfureux émis par la Soufriére sont néfastes a 
la végétation environnante, qu’ils détruisent peu a peu.” Stehlé concludes 
that the species which exist in this environment are particularly interest- 
ing and rare. Many are thought to be endemics, but as more is known of 
the vegetation of the high mountain peaks from St. Kitts to St. Vincent 
it becomes increasingly difficult to separate those species which are re- 
stricted to high elevations on the ash of volcanic ejecta and those which 
are there because of their tolerance to fumarole emissions. However, the 
red-flowered form of the bromeliad Pitcairnia spicata occurs at high alti- 
tudes on St. Kitts, Guadeloupe, Dominica, Martinique, and St. Vincent, 
while the yellow-flowered form, P. spicata var. sulphurea, has not been 
found on St. Kitts and occurs on the other islands in areas influenced by 
the crateral fumaroles. The endemic species of these high mountains 
are not the most distinct of species, and recent collections from other 
islands have indicated that their distribution is wider in the Lesser An- 
tilles than has been previously reported. 


Montserrat 


GALWAY’s SOUFRIERE. (Fig. 15.) One of the earliest accounts of the sou- 
friéres on Montserrat was made by Dr. Nicholas Nugent, an honorary mem- 
ber of the Geological Society of London who visited Montserrat in 1810. 
His point of view in visiting Galway’s Soufriére was entirely geological, but 
his description applies to the present as well. “A rugged horse-path was 
traced along the brink of the ravine, which we followed amidst the most 
beautiful and romantic scenery. At the head of this ravine is a small 
amphitheatre formed by lofty surrounding mountains, and here is situated 
what is termed ‘The Sulphur.’ Though the scene was extremely grand 
and well worthy of observation, yet I confess I could not help feeling a 
good deal disappointed, as there was nothing like a crater to be seen, or 
anything else that could lead me to suppose the place had any connexion 
with a volcano. On the north, east and west sides were lofty mountains 
wooded to the tops, composed apparently of the same kind of porphyry 
we had noticed all along the way. On the south, the same kind of rock 
of no great height, quite bare of vegetation, and in a very peculiar state 
of decomposition. And on the south-eastern side, our path, and the outlet 
into the ravine. The whole area thus included, might be three or four 
hundred yards in length and half that distance in breadth.” 

Karl Sapper visited Montserrat in 1903 (57) and visited both Gages 
and Galway’s soufriéres. He published a sketch map of Galway’s Sou- 


302 JOURNAL OF THE ARNOLD ARBORETUM [| VOL, XLIII 


friere indicating the principal fumaroles where he recorded temperatures 
of 34.2-93.2° C. and commented on the production of hydrogen sulfide y 
each. Sapper estimated the area of destruction of vegetation as between 
three and four hectares. MacGregor (39) reproduced Sapper’s sketch- 
map and concluded that the site of fumarole activity had changed by the 
time of his visit in 1936. Perret also visited Galway’s Soufriére during the 
volcanic crisis in Montserrat in 1934. His description of the area is 
meager, for he concluded that, ‘“Galways Soufriére was a more ordinary 
solfatara having less chemical activity and water of less acidity — alto- 
gether, a vent less capable of serving as an indicator of increasing vol- 
canic activity than the more accessible Gages Soufriére.” He did note 
that the water from Gages Soufriére had an acid content ten times as 
great as that from Galway’s. My tests in 1961 with “pHydrion” paper 
gave a reading of pH 5 for the waters in Galway’s fumaroles and 1.5 for 
those in Gages. 

Sapper commented only on the dead and dying vegetation in the 
vicinity of Galway’s Soufriére, and his outline map of the area of activity 
and destruction has recently been questioned by Martin-Kaye (40), who 
felt from seeing MacGregor’s reproduction of Sapper’s map that the 
main stream was in reality the southeast stream as drawn by Martin- 
Kaye (40). I find it difficult to agree with this conclusion and suggest 
that Sapper and MacGregor are correct. Certainly the locations of the 
principal vent are similar on the two maps in relation to the area of 
destroyed vegetation. 

Martin-Kaye has reproduced temperature records at Galway’s mae 
friere taken by the Montserrat Department of Agriculture from 1936 t 
1952. My own reading taken in January, 1961, showed a vent pe 
ture of 96° C. (200° F.) and pool temperatures 83° and 77° C. (180° 
and 170° F.), respectively, for the pools where, I was told, the tempera- 
tures were generally recorded. 

The vegetation at Galway’s appears to be completely stabilized. The 
prevailing winds blowing down the mountain gorges across the soufriére 
and towards the sea apparently have carried the toxic fumes for years at 
definite elevations. Plants have developed in the lee of fans, piles of scree, 
and even behind boulders. To the windward, the vegetation is unaffected, 
and not at all stunted. The principal plants nearest the soufriere were 
Clusia, Freziera and Ternstroemia. Within the soufriére proper the ex- 
isting and often flourishing plants belong to the genera Andropogon, Clusia, 
Cyperus, Philodendron, Pitcairnia, Pityrogramma, and Ternstroemia. 

GacES LOWER SOUFRIERE. (Fig. 14.) Nugent does not mention Gages 
soufrieres and Shafer’s (61) description of his visit to Montserrat speaks 
so generally of the three soufriéres that one can not place his actual visit. 
Gages Lower Soufriére is said to have originated in 1896 following a 
cloudburst and to have been moderately active during an earthquake series 

98. Perret’s visit to and study of Gages Lower Soufriére during its 
period of renewed activity beginning in 1933 is well recorded. Gages 
Lower Soufriere consists of two parallel valleys, one old and inactive, 


— 


1962 | HOWARD, VOLCANISM AND VEGETATION 303 


the other a “new rift” which opened around 1930. The one valley re- 
mained active through 1935, but by the 6th of May, 1937, the gas phase 
had subsided and the water from the gorge was nearly chemically inert. 
I visited this area in 1950 and again in 1961. Little change was observed 
in the first visit from the condition shown in Perret’s photographs. In 
1961, only minor fume emissions were noticeable with faint odors of 
hydrogen sulfide with sufficient heat and steam to cloud one’s glasses. 
Fume temperatures at the most active vents were 94-96° C. The area of 
devastation, which Perret judged to be 600 feet long and 350 feet wide, 
had not increased, but by 1961 an occasional patch of green from pro- 
thallia or young gametophy tes of mosses was evident, with an occasional 
plant of Andropogon, Capraria biflora, Clusia, or Cyperus present in the 
valley. On the limonitized zone at the head of the valley grasses and 
sedges were becoming more abundant, apparently developing from seed 
sources above the devastated area. 

Gaces Upper SourrikreE. (Figs, 10-13, 18, 19.) Accurate dating is 
not available for the age of Gages Upper Soufriére. Sapper (57) visited 
it in 1903, and Perret indicates that it was dormant prior to the earth- 
quake of November 11, 1935, when its activity increased enormously. 
Perret does not go into detail on his observations on Gages Upper Sou- 
friere, although he delayed his departure from Montserrat to study this 
new outbreak. He does indicate that the temperatures were 99° C, i 
the steam vent and 96° and 98° C. in the pools. His illustrations, and par- 
ticularly Fig. 31 reproduced here, are noteworthy for the remnants of the 
vegetation shown. Clearly, the area of Gages Upper Soufriére had been 
relatively small prior to the 1935 activity, for the density of the vegetation 
indicates a recent kill. Perret noted that there was a threat for Gages 
Upper Soufriére to “extend its domain by reason of gas emission and rain 
erosion.” Martin-Kaye’s observations and map for his visit in October, 
1954, led to the conclusion that ‘“‘much of this area has comparatively re- 
cently opened up as many rotted tree stumps are still to be seen more 
or less in the position of growth.” My own comparative observations al- 
ready mentioned were made in 1950 and again in 1961 and lead only to 
the conclusion that Gages Upper Soufriére is not only as active as it has 
been in the past but is continuing to increase to the southwest and may well 
cross the ridge into the next valley. The area devastated at present is 
about eight acres at an altitude between 1300 and 1400 ft. along the 
stream bed, but extending up the hill to around 1700 ft. 

Tar River SourrizRe. MacGregor (39) has illustrated in his Figure 
37 the appearance of the “New Cow Hill soufriére” (Tar River) shortly 
after its inception in 1936. The fumaroles apparently appeared in the 
center of a banana plantation. When Martin-Kaye and Willmore visited 
the area the fumarole was relatively quiet, and only about an acre of 
vegetation was affected by the fumes. In 1961 steam and fumes were not 
apparent from a convenient observation point on the slopes leading to 
English’s Mountain 

LANG’ s Soursrtre. (Figs. 16, 17.) With the expert guidance of Kings- 


304 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


ley Howes we visited the recently discovered Lang’s Soufriére on the 
north flank of the Soufriére Hills above Paradise Estate. The soufriére 
itself is essentially dormant, giving only occasional indication of hydrogen 
sulfide and some degree of heat. It is located at the head of a deep ravine, 
and landslides not of recent time have covered the actual fumaroles. The 
decomposition of the rock in the area, plus the occurrence of one large 
dead specimen of Sloanea and the abundance of Philodendron giganteum 
attested to the past activity of this fumarole. 

OTHER SOUFRIERES. Through the Soufriére Hills are other indications 
of dead or dormant fumarole areas. Several were seen in Gages Gut en 
route to Gages Upper Soufriére. Neither fumes nor heat is emitted from 
these areas at the present. The older ones were green with mosses and 
lichens, and the younger ones still showed the color of the altered rocks. 
These fumaroles were apparently never of major importance, for the sur- 
rounding vegetation was neither affected nor characterized by the genera 
usually found in active areas. Mulcair Soufriére was known by reputa- 
tion to the local people, but, as Martin-Kaye discovered, it seems to be 
inaccessible, for no local individual was capable of guiding me to it. 

Hot WATER Ponp. This coastal pond is thought to be heated by sub- 
terranean springs, possibly the drainage from the Gages soufriéres. It did 
not support any unusual plant life nor was the surrounding vegetation 
affected in any way. 


St. Kitts 


Martin-Kaye (40) supplies the only references I have encountered to 
fumarole activity on the island of St. Kitts. My visits have been at times 
when the cloud cover made any ascent of Mt. Misery of doubtful value. 
Martin-Kaye reports however, that the crater of Mt. Misery is between 
one-half and three-fourths of a mile in diameter at the rim, 500 to 60 
feet deep, and possesses a small shallow lake in the southwest part of the 
floor, which “except in the vicinity of two small areas of fumarole activity, 
is otherwise covered with vegetation.” Martin-Kaye also reports that 
“now defunct fumarolic activity has caused rock alteration at Rainbow 
Cliff, Frigate Bay where variously colored ochres are associated with 
yellowish, brownish and grey clays, kaolin, some gypsum and probably 
opal powders and alunite as are associated with the active soufriéres in 
Montserrat, Nevis, Mt. Misery itself and elsewhere in the Lesser Antilles.’’ 
Regrettably no data on the surrounding vegetation are available. 


Nevis 


The Rev. Mr. Smith (64) in a series of letters presents some of the 
earliest records of the natural history of Nevis. In his second letter he 
refers to a hot river and to baths “supposed to flow from the sulphur 
ground which is not above three quarters of a mile higher in the country.” 
He also notes that “In my parish of St. John in the Island of Nevis, there is 


1962] HOWARD, VOLCANISM AND VEGETATION 305 


a considerable spot of sulphurous ground on the south side, at the upper end 
of a deep rupture in the earth vulgarly called Sulphur Gut, which is so 
excessive hot . . . as to make us immediately feel it through our Shoe 
Rev. Smith refers to another at a “place two miles and _ half 
south-ward of Charles Town” and reports that ‘a new hot spring was in 
1718, discovered in Windward Parish, upon clearing of a wood . . . just 
above Camp-ground.” 

Bryan Edwards (15) said for Nevis, “That the island was produced 
by some volcanic explosion, in ages long past, there can be no doubt; 
for there is a hollow, or crater, near the summit, still visible, which con- 
tains a hot spring strongly impregnated with sulphur; and sulphur is 
frequently found in substances, in the neighboring gullies and cavities of 
the earth.” In his excellent treatment of the geology of the Leeward 
Islands, Martin-Kaye has reported two active soufriéres on Nevis at 
Farm’s Estate and Cades Bay, the latter first noticed in 1953. In addi- 
tion there is some gas seepage noticeable in the Belmont area and thermal 
pools at Bath Springs near Charlestown and in the Camp Springs area. 
The thermal pools apparently are those described by the Rev. Smith, in- 
dicating an activity of nearly 250 years. 

The Farm’s Estate Soufriére is both old and relatively quiet. Hydrogen 
sulphide odors and wisps of steam are noticeable. Willmore recorded tem- 
peratures of 42-44° C. in small fissures and temperatures of 96° C. in 
a vent. Apparently the vegetation has been affected by the activity of 
the fumaroles but has now become stabilized. 

The Cades Bay Soufriére, according to Martin-Kaye, was first noted 
in 1953 when the “vegetation began to die and hydrogen sulfide was 
evident.” In six months an area twenty by thirty feet containing sunken 
boiling steam holes had developed and odors of sulphur dioxide were evi- 
dent. Shortly thereafter several areas became confluent by the loss of 
vegetation and a barren area of about 1.5 acres was produced containing 
small pools with water registering a temperature of 118° F. Martin-Kaye 
reports that in 1958 the vents “seem to have mainly concentrated in one 
small bubbling pool a few feet across’ and that the ‘‘vegetation had re- 
established itself over the western part.’’ Regrettably no specific informa- 
tion on the nature of the vegetation of the area is available. 


SUMMATION AND COMPARISONS 


Volcanism, either eruptive or fumarolic, is destructive to vegetation, as 
ample examples show in the Lesser Antilles. Nuées ardentes, character- 
istic of recent eruptions, are totally destructive of plant life, and ash falls 
less so. The revegetation of ash-covered areas is related to the type of 
destruction but also to the altitude of the area considered and its prox- 
imity to sources of new plants. At lower elevations or adjacent to un- 
disturbed areas invasion by weedy species is rapid. Plants covered 
only with free-fall ash appear to sprout rapidly from the basal portions 
and quickly supply a covering vegetation. At highest elevations where a 


306 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLII 


specialized vegetation occurs the re-establishment of vegetation is much 
slower. Even when the proper seeds are introduced from adjacent moun- 
tain tops the environment is inhospitable, and decades pass before the 
original introductions have formed an even moderately complete cover. 

Yew areas of fumarolic activity appear to act first through a heating 
effect of the soil to which the native species are selectively tolerant. The 
emission of sulphur compounds creates an acid environment in the soil 
and air-borne acidic fumes. Natives species appear to be killed off in 
direct relationship to their ability to stand increasing acidity. The air- 
borne acid particles appear to be cumulative in their action on the sur- 
rounding vegetation. Ferns with leaves of delicate textures and broad- 
leaved plants without heavy cuticles are most sensitive to these fumes. 
Plants with coriaceous leaves or heavy cuticles are most resistant. At the 
highest elevations on soils composed of volcanic ash it is difficult to de- 
termine the role of fumarole emissions in the establishment and _per- 
sistence of the species present. 

In the Lesser Antilles the species which appear to be most tolerant of 
fumarolic gas emissions and soil acidity are the following: Andropogon 
pertusus, Blechnum indicum, Cladium restioides, Clusia alba, Clusia 
mangle, Clusia plukenettu, Cyperus ligularis, Cyperus planifolius, Dicra- 
nopteris bifida, Guzmania plumieri, Lycopodium cernuum, Lycopodium 
meridionale, Lycopodium tortum, Philodendron giganteum, Pitcairnia an- 
gustifolia, Pitcairnia albucifolia, Pitcairnia spicata var. sulphurea, Pityro- 
ramma calomelanos, Pityrogramma chrysophylla, and Ternstroemia pe- 
duncularis. 

A comparison of the plant species from other geographic areas which 
are tolerant of fumarolic environments or which are invaders of volcani- 
cally disturbed areas is interesting yet difficult due to the differences in 
composition of the floras. Much has been written of the thermal springs 
of the United States, particularly the area of Yellowstone National Park 
where emphasis has been given to heat tolerant algae and to the animal 
life of the hot springs. Similar studies are available for the hot-spring 
areas of Iceland. In both cases, the families, genera, and species of the 
plants can not be compared with those of the Lesser Antilles. The ex- 
tensive fumarole areas of Indonesia and of Japan have received some 
study. Mizushima (41) studied the plant community of the fumarole 
areas of the Idzu Islands, midway between Tokyo and the Bonin Islands. 
He found that the plants tolerant of fumarole conditions were: Chamae- 
syce hirta, Chrysanthemum pacificum, Cyperus polystachyos, Fimbri- 
stylis dichotoma {. floribunda, Imperata cylindrica var. koenigii, Linder- 
nia crustacea, Lycopodium cernum, Lygodium japonicum, Ophioglossum 
petiolatum, Paspalum orbiculare, and Pouzolzia zeylanica (Urticaceae). 
He noted that plants of Lindernia crustacea are of interest as the indi- 
viduals gain in stature as they approach the fumaroles. Hedyotis, Pas- 
palum and Pouzolzia were limited in their distribution on the island to 
the areas of the fumaroles. Cyperus, Fimbristylis, Imperata, Lycopodium, 
and Lygodium were indicated as abundant near the fumaroles. 


1962] HOWARD, VOLCANISM AND VEGETATION 307 


The locations of the numerous areas of volcanism in Indonesia are 
effectively shown in a map published by Ter Braake (9). Braun-Blanquet 
(10) indicates that Pteris incisa thrives on the soft, sulphurous soils ad- 
jacent to springs that are rich in alum. Polypodium vulcanicum has been 
reported to survive on rocks which are occasionally flooded by waters 
with temperatures as high as 75° C, According to Holtermann (23), the 
dominant shrubs in such areas are Agapetes vulgaris and Rhododendron 
retusum., 

Braun-Blanquet suggests that the plants of such fumarole areas are 
mesophytic in nature. Schimper (59) however, indicates that the flora 
of the solfatara areas is composed of xerophilous species. Zollinger (72) 
and later Junghuhn- (34) were the first to observe in Java that the vegeta- 
tion close to fumaroles was composed of alpine species, even when the 
fumarole was 1000 to 1500 meters below the natural alpine region. Schim- 
per described the fumarole area as follows: ‘Where these pools are col- 
lected, usually in large numbers and of different sizes, the soil is a wet 
white clay, which is said by Junghuhn to arise by the action of sulphuric 
acid on trachyte; it is usually covered by a yellowish efflorescence of 
sulphur. The ground is frequently so hot that to remain standing on it is 
impossible. From all the crevices and pools there escape hot vapors of 
suffocating odour, sometimes of sulphuretted hydrogen, at other times of 
sulphurous acid. The water has an acid taste and sets the teeth on edge.” 
He noted further that “the bushes of the solfataras are much lower in 
stature than the surrounding forest and quite sharply marked off from it. 
None of the small trees and shrubs that form the underwood in the 
high-forest appear among them; of forest herbs only a few species repre- 
sented by detached individuals occur; plants that pecan elsewhere in 
the open situations of the same region are entirely abse 

In the vicinity of solfataras Schimper indicates hee are es: numer- 
ous plants of Ficus diversifolia, Rhododendron javanicum, and Vaccinum 
varingiacfolium. In solfataras of lower regions some other species, notably 
Gaultheria leucocarpa, Pree aes javanensis, Myrsine avenis, Rhododen- 
dron retusum, and Rhododendron tubiflorum, occur, and, in addition, 

“isolated ferns with leathery leaves and lycopods are always present in 
such spots.” Plants of the Melastomataceae and Myrsinaceae, although 
abundant in the Lesser Antilles, do not tolerate fumarolic environments. 
The Ericaceae are poorly represented in the Lesser Antilles, and the genera 
do not occur in areas where fumaroles are active. 

Junghuhn noted that in solfatara areas the shrubs are swept clear of 
all epiphytic vegetation, including mosses and lichens, This is also true 
throughout the Lesser Antilles. A new fumarole seems to be most toxic 
to the epiphytic moss, liverwort, and lichen flora of trunks and branches 
of the adjacent vegetation. In dormant fumarole areas, however, mosses 
are among the first invaders. 

The steam vents on Mauna Loa, Hawaii, contain minimal amounts of 
sulphur and do not seem to be influential in determining the vegetational 
components around them. Ferns, particularly Nephrolepis, are abundant, 


308 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


while Dodonea, Styphelia, Vaccinium, and many rubiaceous genera grow 
more vigorously in the humid environment of the vent than at a distance. 

The revegetation of surfaces disturbed by volcanic eruptions has been 
the subject of a classic study in the case of Krakatoa (69). Fosberg (16) 
has outlined the nature of the invasion and revegetation of lava flows and 
ash-cinder areas of Hawaii. Only preliminary studies are available for 
the Paricutin eruptions in Mexico. Calvert and Calvert (12) discuss the 
effects of volcanic activity and subsequent fumarolic activity for Volcan 
Irazu, Costa Rica. However, the altitude and the species involved in all 
these areas do not permit comparisons with similar areas of the Lesser 
Antilles. 


BIBLIOGRAPHY 

1. ANpeRson, J. An account of Morne Garou, a mountain in the island of 
ot, Vincent, with a description of the volcano on its summit. Phil. 
Trans. Roy. Soc. London 75: 16. 

2. ANDERSON, T. Report on the eruptions of the Soufriére in St. Vincent in 
1902, and on a visit to Montagne Pelée in Martinique. Part II. Phil. 
Trans. Roy. Soc. London A. 208: 275-304. 1908. 

3. ANDERSON, T. & FLeETT, J. S. Preliminary report on the recent eruption 
of the Soufriére in St. Vincent, and of a visit to es Pelée, in Mar- 
tinique. Proc. Roy. Soc. London 70: 423-445. 

4, : Report on the eruptions of the Soufriére, in - Vincent in 1902, 


and on a visit to Montagne Pelée, in Martinique. Part I. Phil. Trans. 
Roy. Soc. London A. 200: 353-553. 1903. 
5. Atwoop, T. The history of the island of Dominica. London 


6. BEARD, 7 S. The progress of plant succession on the i ae hes St. Vin- 
cent. Jour. Ecol. 33: 1-9. 5, 

i . The natural vegetation of the Windward and Leeward Islands. 
Oxford, 1949. 

8. Bopvarsson, G. Production of thermal energy on the Island of St. Lucia. 
United Nations, Technical Assistance Administration. New York, 1951. 

9. Braake, A. L. TER. Volcanology in the Netherlands Indies. /n: Honic 


& VERDOORN, Science and scientists in the Netherlands Indies. New 
York, 1945. 

10. BrauN-BLANQUET, J. Plant Sociology. McGraw Hill, 1932. 

11. Burtrertin, J. La constitution geologique et la structure des Antilles. 
Gap, France, 1956. 

12. CALVERT, A. S., & CaAtvert, P. P. A year of Costa Rican natural history. 
Macmillan, 1917. 

13. Davis, W. M. The Lesser Antilles. Am. Geogr. Soc. Publ. 2. 1926. 

13a. Domin, K. The Pteridophyta of the island of Dominica. Mem. Roy. 
Czech. Soc. Sci. Div. Nat. Hist. Math. ser. 2, 2: 1-259. 

14. Earte, K. W. Geological notes on the island of Dominica, BW. Geol. 
Mag. 65: 169-187. 1928. 

15. Epwarps, B. History, ee and commercial, of the British Colonies in the 
West Indies. Vol. 1. 

16. FosBerc, F. R. ae excursion III, Tenth Pacific Science Congress. 
Honolulu, 1961. 


1962 | HOWARD, VOLCANISM AND VEGETATION 309 


17. 


Frotow, S. Les ee de la Montagne Pelée. Revue Martinique 1: 
39-54. 1944. [Not se 

GovAULT, J. orn ee sur la Soufriére de la Guadeloupe pour servir a 
l’étude des appareils éruptifs Peléens. Pointe-a-Pitre, Guadeloupe. 1942. 
[ Not seen. | 

Harpy, F. & Ropricues, G., with Nanton, W. R. E. The agricultural soils 
of Montserrat. Trinidad, 1949. 

Hirt, R. T. Pelé and the evolution of the Windward Archipelago. Bull. 
Geol. Soc. Am. 16: 243-288. 1905 


. Hopce, W. H. The vegetation of Dominica. Geogr. Review 33: 349-375. 


1943. 
. Flora of Dominica, B.W.I. Lloydia 17: 1-238. 1954 


. HOLTERMANN, C. Der Einfluss des Klimas auf den Bau der Pflanzengewege. 


Leipzig. 1907: 
Hooper, E. D. M. Report upon the forests of St. Vincent. London, 1886. 
. Report upon the forests of St. Lucia. Madras, 1887. 


Hovey, E. O. Some erosion phenomena observed on the islands of Saint 


Vincent and Martinique in 1902 and 1903. Bull. Geol. Soc. Am. 15: 
560-1. 1904 
_ Grinde Soufriére of Guadeloupe. /bid. 15: 561-2. 1904. 
. Bibliography of literature of the West Indian eruptions published 
in the United States. /bid. 15: 562-566. 1904. 
. The present condition of Mount Pelé. /bid. 16: 566-9. 1905. 
. Boiling lake of Dominica. /bid. 16: 570-571. 1905. 
—. Soufriére of Saint Lucia. /bid. 18: 569-570. 1905. 
Huccins, P. F. Report on the Soufriére, Saint Vincent, B.W.I. St. Vincent, 
1902. 


. JoHow, F. Vegetationsbilder aus West-Indien und Venezuela. Kosmos 


Stuttgart 2: 270-285. 1884. 


. JUNGHUHN, F. Java, deszelfs gedaante, bekleeding en invendige structuur. 


German ed. (ed. Hasskarl) 1: 453. 


. LaspaTt, PERE. Nouveau voyage aux Isles ‘de Amerique. Vol. 2. 1772. 
. Lacrorx, M. A. L’Eruption de la Martinique. 44 pp. Privately printed, 


Paris, 1903. 


. Lee, W. Daniel Defoe: his life and recently discovered miscellaneous 


writing. Vol. 2. 1869. 

MacGrecor, A. G. Royal Society Expedition to Montserrat, B.W.I.: Pre- 
liminary report on the geology of Montserrat. Proc. Roy. Soc. London B. 
121: 232-252. 1936. 

. The volcanic history and petrology of Montserrat, with observa- 
tions on Mt. Pelé in Martinique. Phil. Trans. Roy. Soc. London B. 229: 
1-90. 1938. 

Martin-Kave, P. H. A. Reports on the geology of the Leeward and 
British Virgin Islands. 1-117. St. Lucia, 1959. 


. Mizusui1ma, M. A preliminary report on the florula of Isl. Aogashima, 


Idzu Islands. Misc. Rep. Res. Inst. Nat. Resources 38: 102-126. 1955. 


. Nucent, N. An account of “The Sulphur” or “Soufriere’” on the island 


of Montserrat. Trans. Geol. Soc. London 1: 185-190. 


: See F. A. Camps in the Caribbees. Boston, 1880. 
. PERR F. A. The eruption of Mt. Pelée, 1929-1932. Carnegie Inst. 


ae Publ. 458. 1935. 


on 
Ww 


nn 


JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


. The volcano-seismic crisis at Montserrat, 1933-1937. Carn. Inst. 
Wash. Publ. 512. 1 
Volcanological observations. Carnegie Inst. Wash. Publ. 549. 


1950. 

Poucn, F. H. Paricutin comes.of age. Nat. History 53: 342-348. 1944. 

REED, F. R.C. The geology of the British Empire. Ed. 2. London, 1949. 

Rogpson, G. R. & Wit_more, P. L. Geological and eps eee 
on the Qualibou Soufrieére of St. Lucia. 1-6. Trinidad, 

. Some heat measurements in West Indian ee “Bull. Vol- 
can. ser. 2. 17: 13-39. 

Romer, M. [Title unknown.| Bull. Serv. Meterol. et de ?Observ. Geophys. 
1953. | Not seen. | 

Russet, I. C. Volcanic En uOns on ec and St. Vincent. Ann. 
Rep. Smithson. Inst. 331-34 903. 

Sanps, W. M.. An account e the return Fe vegetation and the revival of 
agriculture, in the area devastated by the Soufriere of St. Vincent in 
1902-03. West Indian Bull. 12: 22-33. 1912. 

Sapper, K. Besuch den Insel Grenada. Centralbl. Min. Geol. Palaeont. 1903: 
182-186. 1903. 

Bericht iiber einen Besuch von St. Vincent. /bid. 248-251. 1903. 

Zur Kenntniss der Insel S. Lucia in Westindien. Jbid, 273-278. 


— 


1903. 

. Ein Besuch der Insel Montserrat. /bid. 279-283. 1903. 

Scuwabe, G. H. Beitrage zur kenntnis Islandischer Thermalbiotope. Archiv 
Hydrobiol, puDpL, Ser, 6: 161-352... TO3D. 

Scuimper, A. F. W. Plant-geography, upon a physiological basis. English 

Pacing by W. R. Fisuer. Revised and edited by P. Groom & I. B. 

our. Oxford. 1903. 

ees C. Historical geology of the Antillean-Caribbean Region. New 
York, 1935 

SHAFER, Wi Report on a visit to the Island of Montserrat. Jour. N.Y. Bot. 
Gard. 8: 81-88. 1907. 

SHEPHARD, Mr. Historical account of the Island of St. Vincent. London, 
1831 

SmitH, LONGFIELD. Volcanic eruptions in the West Indies. West Indian 
Bull. 3: 271-293. 1902 

SmirH, Rev. Mr. W. A natural history of Nevis. Cambridge, 1745. 

STEHLE, H. Flore de la Guadeloupe et Dépendances. Essai d’écologie et 
de eae botanique. Vol. 1. Basse Terre, Guadeloupe, 1935. 

ae associations végétales de la Martinique. Bull. Agr. 

19 


Martniga 6: etre 

; rest” aie of she) Carib bean Islands. Carib. For. 6(Suppl.): 
273- — ecrc 
s types forestiers des Iles Caraibes. /bid. 7: 337-709. 1946. 


VAN pone M. N. Catalogues of the active volcanoes of Indonesia. 


(Part I in Catalogue of the active volcanoes of the world.) Naples, 1951 
— I. Herbaceous angiosperms of the Lesser Antilles. Puerto Rico, 


195 

Wittmore, P. L. The earthquake series in St. Kitts-Nevis, 1950-51. Nature 
69: 770-772. 1952. 

ZOLLINGER, H. Ueber Pflanzenphysiognomie im Allgemeinen und diejenige 
der Insel Java insbesondere. Zurich, 1 


1962] HOWARD, VOLCANISM AND VEGETATION Sl 


EXPLANATION OF PLATES 
PLATE I 


Fic. 4. Cone of fumarole at Lavera rae Springs, Grenada. Fic. 5. Path of 
mineral deposit from cone in Fig. 4, nada. Fic. 6. Valley of Desolation, 
Dominica. Fic. 7. Steam vent at Tee en Dominica. Fic. 8. eClaseran 
guadelupense” on windward slopes of La Soufriére, Guadeloupe. Dicranopteris 
bifida and Cladium restioides are seen in foreground, Clusia mangle in middle, 
and /lex species on distant slope. Fic. 9. The same on leeward slopes, showing 
killing by periodic emissions of sulfur fumes. Dead plants are Clusia, Ilex, and 
Freziera; living plants are Philodendron giganteum. 


PLATE II 


Fic. 10. Gages Upper Soufriére, Montserrat, after its reactivation on No- 
vember 11, 1935. Photo reproduced from Perret (45). Fic. 11. Gages Upper 
Soufriére, ‘Montserrat, as it appeared in June. 1950. Fic. 12. Gages Upper 
Soufriére, Montserrat, as it appeared in January, 1961. Fic. 13. Dead and 
acid-rotted stump of Sloanea dentata at Gages Upper Soufriére in January, 
1961. A fumarole had opened under this tree within the last decade, for the tree 
was alive in 1950. Fic. 14. Gages Lower Soufriére, Montserrat, in January, 
1961. Vegetation has descended on the limonitized area from above. Fic. 15. 
Main steam vent at Galway’s Soufriere, Montserrat, in January, ge Effect 
of the prevailing winds from the mountain is shown by the path of the steam. 
Effects of the fumes are visible on the hillsides by the reduction of vegetation. 


PLATE III 


Fic. 16. Lang’s Soufriére, Montserrat, as seen from Paradise Ridge. Point 
of arrow locates the dead Sloanea trunk shown in Fic. 17. Rear of the arrow 
marks dense growth of Philodendron giganteum. Fic. 17. Point of emission 
of fumes from Lang’s Soufriére, Montserrat. A dead trunk of Sloanea is on 
the left. Recent rock fall shows the depth of the effects of the acidity on the 
rock face. Fic. 18. A new area of fumarole activity at Gages Upper Soufriére, 
Montserrat, January, 1961. Plants of Clusia alba are in the foreground. Cyathea 
and other components of adjacent woodland are shown in the background, as 
yet, not affected by fumes of the new fumarole. Destructive effects to the pres- 
ent are due to heat and soil acidity. Fic. 19. Gages Upper Soufriére, Montserrat, 
January, 1961. The upper valley of this soufriére showing the proximity sat 
the undisturbed woodland on the windward side of the fume area. 

Qualibou Soufriére, St. Lucia. Boiling lakes and steam vents with Saas 
rock are in the foreground. The profile of the hill in the background ee 
the absence of shrubs or trees. Fic. 21. Qualibou Soufriere, St. Lucia. A fac 
view of the hill shown in Fig. 20. The hillside vegetation is of Blechnum a 
Lycopodium where affected by the fumes of the soufriére. The edge of the 
affected area is shown on the right, 


Jour. Arnotp Ars. VoL. XLII PLATE I 


Howarp, VOLCANISM IN THE LESSER ANTILLES 


Jour. ARNOLD Ars, VoL. XLIII PriatTE II 


eh 


Howarpb, VOLCANISM IN THE LESSER ANTILLES 


Jour. ARNOLD Ars, VoL, XLIII PiatTE III] 


Howarb, VOLCANISM IN THE LESSER ANTILLES 


1962 | ERNST, PAPAVERACEAE & FUMARIACEAE 315 


THE GENERA OF PAPAVERACEAE AND FUMARIACEAE 
IN THE SOUTHEASTERN UNITED STATES ! 


WaLLace R, ERNST 


PAPAVERACEAE A. L. de Jussieu, Gen. Pl. 235. 1789, nom. cons. 
(Poppy FAMILY) 

Annual or perennial, sometimes rhizomatous herbs |shrubs, or small 
trees|, often glaucous and pubescent with multicellular [or unicellular | 
hairs or glabrous, usually with acrid, reddish-orange to yellow |or white | 
sap. Leaves rosulate and/or cauline, usually alternate, reduced upward, 
exstipulate, petiolate or sessile, pinnately (seld om palmately) [or tern- 
ately | veined, lobed, or dissected |or entire]. Inflorescences terminal 
and determinate, 1-many -flowered, usually cymose [or rarely paniculi- 
form|; buds erect [or nodding], subtended by a leaf or bract. Flowers 
erect, bisexual, open, essentially regular and hypogynous [seldom perig- 
ynous ]. Perianth cyclic, imbricate, usually 3-seriate, 2- or 3-merous. 
Sepals 2 or 3 [seldom synsepa lous}, usually with dissimilar left-right- 
oriented margins [seldom reversed], tightly inclosing the buds until 
anthesis, then caducous. Petals usually 2-seriate and twice as many as 
sepals (sometimes polypetalous) |or seldom apetalous], opening diurnally, 


"Prepared for a generic flora of the southeastern United States, a joint project of 
the Gray Herbarium and the Arnold Arboretum made possible through the support 
of George R. Cooley and the National Science Foundation and under the direction of 
Reed C. Rollins an Carroll E. Wood, Jr. This et follows the style established 
in the first paper the series, Jour. Arnold Arb. 39: 296-346. 1958 (and continued 
through volume a The area covered in this, as in eeyaiae treatments, is bounded by 
and includes North Carolina, Tennessee, Arkansas, and Louisiana. The deseriptions 
apply primarily to the plants of this area with any supplementary information in 
brackets. References which have not been verified by the eee a marked by an 
asterisk. The il Ee were prepared by Mrs. Dorothy H. Marsh, the manuscript 
by Mrs. Gordon W. 

This ee ie is ine in part, on a survey of the floral morphology of Fuma- 
riaceae and a doctoral dissertation on the floral morphology of Papaveraceae. The 
subfamilies described here, discussed in detail in the above dissertation, represent a 
re-evaluation of the generic affinities. Altogether, these four subfamilies constitute the 
subfamily Papaveroideae, as used by Fedde, their names and included genera more 
or less corresponding to of his five tribes. In the terminology of Fedde, the 
Romneyeae Benth. & Hook. (including Rone eva Harv. and Arctomecon Torr. 
Frém.) are placed with the Papavereae, and Glauciunt Mill. has been removed from 
the ee ereae and placed with the Chel ee Reichen 

e Russian literature citations were translated by Dr. ie K. Brizicky; the elo- 
Bene cae descriptions were prepared by Dr. R. C. Foster; Dr. G. W. Gillett spe 
Mr. R. C. Bean provided material for cytological study; rs. H. Francia Hom 
mersand verified some distribution records in the herbarium of the University of 
North Carolina; and Dr. C, E. Wood, Jr., has given much expert advice. 


worn 


316 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


sometimes wrinkled, usually fugacious; the outer series larger and alter- 
nate with the sepals. Stamens usually many, indefinite, varying in num- 
ber with the size of the flower, usually shorter toward the petals; the 
filaments sometimes expanded lor toothed| and often constricted below 
the anther: anthers basifixed, 2-locular at anthesis, usually extrorse, 
dehiscent by longitudinal slits; pollen often 3- colpate, sometimes poly- 
porate |polycolpate or rugate]. Gynoecium syncarpous, of 2 to many 
carpels; stigmas laterally confluent |or discrete|, as many as and alternate 
with the placentae, the margins of adjacent stigmas often conspicuous 
over the placentae [see Eschscholzia|; style definite to absent; ovary 
1-locular [locule rarely occluded; or rarely multilocular; or rarely the 
carpels conduplicate about a central cavity]; ovules usually numerous 
[seldom 1], obovoid, amphitropous, 2- integumented, the micropyle be- 
neath the funiculus; placentation parietal, sometimes intruded into the 
locule. Fruit dehiscent by 2 to many valves alternate with the persistent 
placentae [or fruit nonvalvate and dehiscent through the placentae|; de- 
hiscence usually incomplete, acropetal or basipetal. Seeds [1 to] many, 
obovoid to -_ reniform, free [or rarely fruit fragmenting into 1-seeded 
joints|, variously sculptured or smooth, sometimes arillate and attractive 
to ants; endosperm present, often oily; embryo small, straight or some- 
what curved. Seedling cotyledons linear | very rarely bifid] or expanded 
ace petiolate. (Excluding Fumariaceae DC. and Hypecoum L.) TYPE 
Nus: Papaver L. 


About 25 genera and at least 200 species, widely distributed in the 
Northern Hemisphere from sea level to ca. 5800 m. (19,000 t.)* 
Papaver, local in both South Africa and Australia, extends northward 
into the Arctic; Bocconia and Argemone occur in the New World tropics. 
Of the 13 genera indigenous to the United States, only three occur in our 
area. 

he genera may be divided conveniently into four subfamilies. The 
Chelidonioideae Ernst, the most heterogeneous of the subfamilies, tend- 
ing to have ~aulecelluiay uniseriate hairs, 2-merous perianths, 3-colpate 
or polyporate pollen, 2-valved gynoecia (3-S-valved in Stylophorum 
diphyllum) with well-developed median carpellary traces, and arillate 
seeds, are represented in our area by Sanguinaria and Stylophorum, and 
by Chelidonium majus, a naturalized weed. Glaucium flavum Crantz (2n 

* Chelidonioideae, subfam. nov. Papaveraceae pilis mollibus, bog multicellu- 
laribus, terminaliter uniseriatis, septis perpendicularibus (in onia Y in 
Maes ramosis, fastigatis et imbricatis) ; perianthio dimero Gn Eomecone synse- 
palo; in Saucers polypetalo; in Macleaya et a apetalo) ; gynoecio biplacen- 
tato et bivalvato (valvis 3-5 in Stylophoro diphyllo); vena carpellare mediana bene 
explicata; valvis acropetaliter vel basipetaliter as lait sed dehiscentia plerumque 
laterale et subcompleta; seminibus saepe arillatis (arillus deest in Glaucio, Dicranostig- 
mate, et Macleaya microcarpa) ; polline tricolpato vel polyporato. Herbae perennes, 
saepe rhizomatosae vel in Bocconia ogee in hemisphaerio septentrionale. Typus 
GENERICUS SUBFAMILIAE: ae ie ee L. Genera AltA: Bocconia L., Coreanomecon 
Nakai, Dicranostigma Hook. & Thom age Hance, Glauctum Mill, Hylomecon 
Maxim., Macleaya R. Br., Sanguinaria L, Stylophorum Nutt 


Ww 
—. 
a 


1962 | ERNST, PAPAVERACEAE & FUMARIACEAE 317 


= 12, 24) and Macleaya cordata (Willd.) R. Br. (2n = 20) are reported 
as garden escapes or weeds to the north of our area. The dehiscence of 
the valves is acropetal in Sanguinaria and Chelidonium but basipetal in 
Stylophorum, Glaucium Mill., and Macleaya R. Br. 

The Papaveroideae,* see with us by Argemone and at leas 
three introduced species of Papaver, tend to have multicellular-multi- 
seriate hairs, 2- or 3-merous perianths, 3-colpate |or several-aperturate | 
pollen, and usually more than 3-valved gynoecia; the median carpellary 
traces tend to be absent and are replaced, in Papaver and some other 
genera, by pse eee ae traces originating from the placental bundles; 
dehiscence of the valves is basipetal, and the seeds usually lack arils. 

e Seeder Ernst,? occasionally represented in our area by 
Eschscholzia californica, are glabrous or have unicelluiar hairs, usually 
2-merous perianths, polycolpate pollen, and 2-valved, conspicuously ribbed 
gynoecia with well-developed median carpellary traces: the dehiscence of 
the valves is acropetal and violent; the seeds usually lack arils. 

latystemonoideae Ernst,’ vernal herbs of the western United 
States, have multiseriate hairs, 3-merous perianths, 3-colpate pollen, 3- 
or several-carpellate gynoecia with median carpellary traces, and discrete 
stigmas; the fruits open by splitting through the placentae without the 
formation of valves, and the seeds lack arils. 
he family is rich in alkaloids, some of them important narcotics, with 
both medicinal and poisonous properties claimed. The alkaloid protopine, 
originally thought common and unique to both Papaveraceae and Fu- 
mariaceae, has been identified in the Berberidaceae; berberine, having 


e same empirical formula, is reported for Papaveraceae and six other 
families (see Willaman & Schubert). The acrid sap, usually colored and 
4 averoideae, subfam. nov. Papaveraceae pilis multicellulari-multiseriatis ; 
perianthio di- vel trimero (aliquando polypetalo) ; gynoecio triplacentato vel placentis 
valvisque numerosioribus; vena carpellare mediana plerumque deest, saepe vena pseudo- 
mediana de fasce placentale substituta; fructu basipetaliter oe nte; in Arctomecone 


humile seminibus arillatis; polline tricolpato vel pluvi-aperturato. Herbae annua vel 
perennes (raro sublignoste), praecipue in hemispherio Cee TYPUS GENERI- 
CUS SUBFAMILIAE: Papav . GENERA ALIA: Arctomecon Torr. & Frém., Argemone 


L., Canbya Parry, Meconepss Vig. (Cathcartia Hook. f.), Roemeria Medic., Romneya 
Harvey, Sivlomecon G. Tayl. 

‘Eschscholzioideae, subfam. nov. VPsapaveraceae plerumque glabrae, vel pilis 
Trea et papillatis; perianthio trimero vel plerumque dimero (in Eschscholzia 
synsepalo); gynoecio biplacentato et bivalvato, longitudinaliter conspicue nervato; 
vena carpellare mediana bene explicata; valvis acropetaliter ees dehiscenti- 
us; in Dendromecone seminibus arillatis; polline polycolpato. erbae annuae vel 
perennes breve victi, vel lignosae, Civitatium Consociatorum bee et Mexiconis 
septentrionalis orientalisque. YPUS GENERICUS SUBFAMILIAE: Fschscholzia Cham. 
GENERA ALIA: D 


Platyste ape 
foliis angustis et integris, saepe verticillatis; perianthio trimero; gynoecio tri- 
pluri-placentato, vena carpellare mediana rikawern ndo tarde explicata; fructu semi- a 
carpico, carpellis partim vel in toto divisis per placentis sine !ormatione valvaru 
seminibus exarillatis; polline tricolpato. Herbae vernales Californiae, etiam erin 
et Washingtonis. TyPUS GENERICUS SUBFAMILIAE: Platystemon Benth. GENERA ALIA: 
Hesperomecon Greene et Meconella Nutt. ex Torr. & Gray 


318 JOURNAL OF THE ARNOLD ARBORETUM [voE Stir 


darkening after exposure, apparently contained in articulated tubes or 
nonarticulated sacs, is an important characteristic of the family. 

The interpretation of the gynoecial structure has been hotly contested 
by morphologists, some concluding, for example, that the 2-valved fruits 
are the equivalent of four carpels arranged in two series (see Bancroft), 
while Arber favored the classical 2-carpel interpretation. The latter view, 
both simple and appealing, is applicable with equal success to all mem- 
bers of the Papaveraceae and the Fumariaceae. The 2-valved fruits in 
Papaveraceae seem less specialized in some respects than the multivalved 
ruits, 

It seems generally agreed that the Papaveraceae are to be placed near 
the Ranales, on the one hand, and near Fumariaceae, Cruciferae, and 
Capparaceae, on the other. Papaveraceae and Fumariaceae have 2-seriate 
corollas of twice as many petals as sepals, while the corollas of Cruciferae 
and Capparaceae are + uniseriate and the petals as many as, and alternate 
with, the sepals. The insertion of the perianth segments is fundamentally 
similar in Fumariaceae and Papaveraceae, but the patterns of the traces to 
the perianths are + different. The sepals of Papaveraceae inclose the buds 
until anthesis, while those of Fumariaceae being mere scales, sometimes 
without traces, do not 

Chromosome aamibers are reported for many species (2n = 12, 14, 16, 
18; 20, 22). 29, 20, Cay, 32,84, 20, 38, 40,42, 44, 50;-Caz 58,. 70,00, 34, 

1. 112). Gametic numbers of 6, 7, or multiples thereof are + common, 
while 8-11, 16, and 19 are uncommon and usually from rather exceptional 
taxa in the family. Species small in habit and usually annual tend to 
have low chromosome numbers, while those large in habit, among them 
the woody members of the family, tend to have higher numbers. 

Economically the family is important as the source of showy garden 
plants; edible poppy seed, poppy-seed oil, and opium and its derived 
alkaloids are obtained from Papaver somniferum. 


= 


REFERENCES: 

ALEXANDER, I. Entwicklungsstudien an Bliiten von Cruciferen und Papaveraceen. 
Planta 41: 125-144. 1952. 

ARBER, A. Studies in flower ae IV. On the gynaeceum of Papaver and 
related genera. Ann. Bot. II. 2: 649-663. 1938. | Basic structure of gyno- 
ecilum, with detailed Keren of the inverted portions of the placental 


bundles. | 

Bancrorr, H, A review of researches concerning floral morphology. Bot. Rev. 
bE 77-99, 1935, [Synopsis of controversy over carpel polymorphism, 80- 
84. | 


BerNuaArpr, J. Uber den Charakter ae die eg der Papaveraceen 
und Fumariaceen. Linnaea 8: 401-477. 1833. [See also ibid. 12: 651-668. 
1838. | 

Bie. J.-D., & G. Bocguer. se campylotrope : les Rhoeadales. Actes Soc. 
Helvét. Nat. Argovie 1960: 

BeRSILLON, G. Recherches sur ae ee iéracées; contribution a l’étude du 
développement des dicotylédones herbacées. han Sci. Nat. Bot. XI. 16: 
225-447. 1955. [Primarily Papaveraceae; a general coverage of phyllo- 


> 


1962] ERNST, PAPAVERACEAE & FUMARIACEAE 319 


taxy, histology, differentiation of conductive tissue, and floral morphology ; 
extensive bibliography. 

Buxatscu, F. Zur Analyse der Bakterienhemmstoffe aus der Wurzel vom 
Schoéllkraut und ahnlichen Pflanzen. Arch. Mikrobiol. 24: 281-296. 1956. 
[Includes Chelidonium majus and Sanguinaria canadensis. | 

Cuist1aKkov, I. D. Organogensis in the family Papaveraceae Hook. Moscow. 
1871.* [Fide A. L. TAKHTAJAN, pp. 21, 22. Problems in evolutionary plant 
morphology. AIBS English translation by O. H. GANKIN, ed. G. L. STEB- 
bins. 139 pp. Washington. 1959. | 

Crete, P, Papavéracées. Le développement de l’embroyn chez le Dicranostigma 
franchetianum (Prain) Fedde. Compt. Rend. Acad. Sop. Paris 245: 720= 
fide. 1997; 

Dickson, J. Studies in floral anatomy. II. The floral anatomy of Glaucium 
flavum with references to other members of the Papaveraceae. Jour. Linn. 
Soc. 175-224. 1935. [A polycarpellate interpretation of the 
gynoecium. | 

Eames, A. J. The vascular anatomy of the flower with refutation of the theory 
of carpel polymorphism. Am. Jour. Bot. 18: 147-188. 1931. | Directed 
at the works of E. R. Saunders. | 

Morphology of the angiosperms. xili + 513 pp. McGraw-Hill, New 

York. 1961. [Reaffirms polycarpellate interpretation of the gynoecia, 198, 

243: reports embryos undifferentiated when seeds shed in several genera 


___— & C. L. Wutson. Crucifer carpels. Am. Jour. Bot. 18: 638-656. 1930. 
[Spirited reassertion of polycarpellate explanation of the cruciferous gyno- 
ecium, extended to closely allied families; gynoecium of Corydalis aurea 
shown as though 3-locular, 654; see earlier paper (1928) for application to 
Fumariaceae and Capparaceae, therefore indirectly to Papaveraceae. | 

Ernst, W. R. Chromosome numbers of some western Papaveraceae. Contr. 
Dudley Herb. 5: 109-115. 1958. Chromosome numbers of some Papaver- 
aceae. Ibid. 137-139. 1959. 

A comparative morphology of the Papaveraceae. vii + 213 pp. 202 
figs. Ph.D. Dissertation, Stanford Univ., Calif. 1962. | Excludes Fumaria- 
ceae; deals primarily with floral morphology at the generic level, recognizes 
4 subfamilial groups. | 

Feppe, F. Die geographische Verbreitung der Papaveraceae. Bot. Jahrb. 
36(Beibl. 81): 28-43. 1905. [Includes Fumariaceae. | 

———. Papaveraceae-Hypecoideae et Papaveraceae-Papaveroideae. Pflanzen- 
reich IV. 104(Heft 40): 1-430, 199° [The basic work; family treated as 
composed of subfamilies Hypecoideae, Papaveroideae, and Fumarioideae, 
the last omitted. | 

Papaveraceae. Nat. Pflanzenfam. ed. 2. 17b: 5-145. 1936. {Important 
synopsis covering for the first time subfam. Fumarioideae. | 

Gates, B. N. Carunculate seed dissemination by ants. Rhodora 45: 438-445. 
1943. [Includes Chelidonium majus and Dicentra spectabilis. | 

Gray, A. Ordo Papaveraceae. Gen. Pl. U. S. 1: 108-116. pls. 47-49, 1848. 
[Argemone, Stylophorum, Sanguinaria. | 

Harms, H. Reihe Rhoeadales. Nat. Pflanzenfam. ed. 2. 17b: 1-4. 1936. 

Harvey-Grpson, R. J., & M. Brapitey. Contributions towards a knowledge of 
the anatomy of the lower dicotyledons. I. The anatomy of the stem of the 
Papaveraceae. Trans. Roy. Soc. Edinb. 51: 589-608. pls. 1-3. 191 


320 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLIII 


HeGNAUER, R. Die Gliederung der Rhoeadales sensu Wettstein im Lichte der 
Inhaltstoffe. Planta Med. 9(1): 37-46. 1961.* 

HoneyMan, J. M. On the occurrence of cyanogenetic glycosides in the order 
Rhoeadales. Taxon 5: 33, 34. 1956. 

Hurcuinson, J. Bocconia and Macleaya. Kew Bull. 1920; 275-282. 1920. 

—. Contributions towards a phylogenetic classification of flowering plants: 
V. The genera of Papaveraceae. /did. 1925: 161-168. 1925. [A radical 
classification with keys to tribes and genera; suggests a phylogeny orig- 
inating from forms with few stamens. 

Lecer, I.. Recherches sur l’appareil végétatif des Papavéracées Juss. Mém 
Soc. Linn. Normandie 18: 195-623. 1894-1895. [| Anatomical study, meladiag 
Fumariaceae, of vegetative organs and laticiferous system; an unlikely 
phylogeny presumes Fumaria basic to Fumariaceae, these giving rise to 2 
ultimately confluent phylads of Papaveraceae, 5 7] 

LIGNIER, O. Notes anatomiques sur l’ovaire de quelques Papavéracées. Bull. 
Soc. Bot. Fr. 58: 279-284, 337-344, 429-435. 1911 

ManesHwakr!, P., & K. K Intraovarian selina in Eschscholzia cali- 
fornica Chan, Argemone mexicana L., and A. ochroleuca Sweet. Nature 
191; 304. 1961.* 

MarkKcrafF, F. ci ee phylogenetische Studien an Bluten von Cruciferen 
und Papaveraceen. Proc. 7th Int. Bot. Congr. 1950: 352-354. 1953. 
eae phylogenetische Studien an Bliiten der Rhoeadales. J/bid. 
857, 

ee g, Untersuchungen Uber den Blutenbau der Papaveraceen. Vet-akad. 

Handl. 50: 1-168. pls. 1-28. 1912. |General morphological approach to 
most genera of Papaveraceae and Fumariaceae showing many floral anom- 
alies; separates Pteridophylloideae from Hypecoideae and Fumarioideae. | 

NorDHAGEN, R. Remarks on some new or little known myrmecochorous plants 
from North America and East Asia. Bull. Res. Council Israel D 
201. 1959. fees ee Chelidonium, etc.; discusses termi- 
nology of seed-ant relationship; thinks stem anatomy degenerate. 

Norris, T. Torus aan and nectary ee as phylogenetic criteria 
in Seni Am. Jour. Bot. 28: 101-113. 1941. [Includes Fumariaceae. | 


Payer, J. Traité d’organogénie comparée de la fleur. 2 vols. Masson, Paris, 
Lot Excellent illustrations of developmental stages, including Fumaria- 
ceae. | 


Popov, M. G. On the relationships and the history of Papaver and Roemeria. 
(In Russian.) Bot. Zhur. 42: 1389-1397. 1957. | Published posthumously ; 
deals little with Papaver and Roemeria at the generic level; a general 
phylogenetic consideration of the Papaveraceae and Fumariaceae, strongly 
influenced by weg of color and hue of the petals and modern dis- 
tribution of the s 

Roper, I. Ae ctont atte ond fluoreszenzoptische Untersuchungen an Same 
Papaveraceen. Osterr. Bot. Zeitschr. 104: 370-381. 1958. | Shows ane 

of seed coats, including some Fumariaceae. | 

SAGDULLAJEVA, A. L, Pollen morphology of Papaveraceae. (In Russian; very 
brief paeee sy hae Probl. Bot. 4: 11-50. 1959. [Reports on 84 spp. 
in 28 genera, including Fumariaceae; 10 types of pollen are described; 
well oe a: i of Dendromecon shown is atypical. 

SAUNDERS, E. R. Floral morphology. Vol. 1. vii + 132 pp. W. Heffer & 
Sons, Ltd., Cambridge. 1937. | Papaveraceae and Fumariaceae, 45-56; her 


1962 | ERNST, PAPAVERACEAE & FUMARIACEAE 321 


complete literature references, 23-25; see particularly New Phytol. 29: 
44-55. 1930; advocates the polycarpellate interpretation of the gynoecium. | 

SHaw, C. H. Note on the sexual generation and the development of the seed- 
coats in certain of the Papaveraceae. Bull. Torrey Bot. Club 31: 429-433. 
pl. 15. 1904. [Includes Sanguinaria, Chelidonium, and Eschscholzia; re- 
ports division of microsporocytes of Sanguinaria in early spring. | 

SIMS, . Bocconia cordata. Bot. Mag. 44: pl. 1905. 1817. [= Macleaya cor- 
dat 


SLAVIK, "y, & L. StavixovaA. Alkaloids of the Papaveraceae. V. Chem. Listy 
48: 1557-1559. 1954; VI. Ibid. 49: 106-110. 1955; VII. Lbid. 1546-1549; 
IX. Ibid. 51: 1923-1926. 1957. (In Czech); VIII. Collect. Czech. Chem. 
Commun. 22: 279-285. 1957; XIV. Ibid. 25: 756-760. 1960; XVI-AVII, 
XV. bid. 1663— 1675, 1698, 1699. (In German.) [None seen; order of 


Sources, R. Embryogénie des Papavéracées. Développement de l’embryon 
chez le Roemeria violacea Medic. (R. hybrida DC.). Compt. Rend. Acad. 
Sci. Paris 226: 979-981. 1948. 

WittaMaN, J. J., & B. G. Scuusert. Alkaloid-bearing plants and their con- 
tained alkaloids. U. S. Dep. Agr. Tech. Bull. 1234: 1-287. 1961. [Gives 
botanical names and reported alkaloids, cross-referenced with empirical 
formulas and plant index; literature citations through 1959, from Chem. 
Abstr. through 1957. | 


Key TO THE GENERA OF PAPAVERACEAE 


General characters: plants herbaceous, sometimes rhizomatous, with acrid, 
usually colored sap; leaves rosulate or usually alternate (rarely opposite) ; in- 
florescence terminal and cymose (rarely paniculiform|; flowers solitary or 
clustered, subtended by a leaf or bract, bisexual, hypogynous (or rarely perig- 
ynous), essentially regular; perianth cyclic, 2- or 3-merous; sepals inclosing 
the buds until anthesis, then caducous; petals twice as many as sepals and 2- 
seriate (occasionally polypetalous) frarely apetalous|; stamens many; gynoe- 
cium 1-locular with 2 or more parietal placentae; fruit dehiscent by 2 or more 
valves 
A. Inflorescence ees or cymose, flowers solitary or several and clustered; 

petals 4-6 or m 

B. Gynoecium ae 2 placentae and valves 

C. Leaf blades palmately veined and lobed, glabrous; flowers solitary on 
scapes; dehiscence - valves acropetal; seeds arillate; low perennial 
fromthick: rhizOme:. 45 co.ee ee ee ee tet eae 1. Sanguinaria. 

C. Leaf blades Reet veined, lobed or dissect 
D. Flowers hypogynous, aposepalous: leaves esac broadly pin- 

nately dissected. 
E. Inflorescence subumbellate, flowers subtended by small, nar- 
row bracts; dehiscence of valves acropetal; seeds arillate. 
Pe ET ee en tates cy Ce gt eee ae 3. Chelidonium. 
E. Inflorescence an expanding cyme, flowers subtended by 
bracts; locule occluded, dehiscence of valves basipetal; seeds 
nonarillate; fruit linear, becoming 10 cm. or more long; ne liage 
BEVig: tt 2 Rests tees aon A Mee mc Ree neni een Ash St a | Glaucium., | 
D. Flowers perigynous, synsepalous; leaves pinnately finely eee 


oz2 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLIII 


glabrous (or hairs pagan, fruits longitudinally ribbed, de- 
NISGERCE CIO DOIAls. @ 4 arcade kw ee Ro egw Ad Verma 4. Eschscholzia. 

B. Gynoecium with 3 or more placentae and valves; dehiscence basipetal. 
F. Flowers on + weak pedicels subtended by small bracts and 2 or 3 
+ opposite leaves on a naked shoot; hairs soft, multicellular-uniseri- 
ate; fruits nodding, dehiscence + complete: seeds dae Ae sae 
OVENS coined a Sine 4d eee we Wee ore deb ee We ye . Stylophorum. 
“, Flowers on ‘rigid pedicels subtended by leaves or hone bracts on 
leafy shoots; hairs multicellular- oO fruits erect, dehiscence 

incomplete; seeds nonarillate; ann 

G. Flowers on short pedicels’ eee undulate, style short; leaves 
Tes A OI cess cd Sem, had Gon Bini SusseG Sana wa cee : Arsemone, 
G. a on long pedicels; stigmas appearing radiate and embedded 
a cartilaginous disc, style absent; leaves not prickly. 6. Papaver. 
A. ipa paniculiform, flowers numerous, apetalous; fruits 2-valved, 
flattened, nodding, dehiscence basipetal; tall herbaceous perennials wit th 
broad, long: DeLwIte LEAVES, 6 oc unica wekey ad ee ekwwe ee xu eas | Macleaya. | 


— 
4 


Subfam. CHELIDONIOIDEAE Ernst 
1, Sanguinaria Linnaeus, Sp. Pl. 1: 505. 1753; Gen. Pl. ed. 5. 223. 1754. 


Low, glabrous, perennial herb with red-orange sap, from a thick rhizome. 
Leaves few, with broad, basically reniform-sagittate, palmately veined 
and variously lobed Bladas, long-petiolate, subtended by 2-6 ovate to 
spatulate bracts. Flowers solitary on naked scapes (or very rarely 2 
or 3, each subtended by a bract); scape at first enfolded by a leaf. Sepals 
2, thin, + notched apically. Corolla often appearing quadrate, 2 (or 
more )-seriate, with 8 (rarely 6-12), narrow, white or pale-pink, sometimes 
irregularly inserted petals. Stamens many; anthers oblong; pollen poly- 
porate with 9-16 apertures. Stigma 2-lobed; style + definite; placentae 
2, Fruit broadly fusiform, 2-valved; dehiscence + acropetal, usually in- 
complete apically. Seeds few to many, black or reddish-orange with a 
conspicuous aril along the upper surface. Seedling cotyledons hypogeal, 
oblong, short-petiolate. Typr species: S. canadensis L. (Name from 
Latin, sanguinarius, bleeding, from the color of the sap.) — BLooproor, 
PUCCOON 


A monotypic genus of eastern North America. Sanguinaria canadensis, 

= 18,° with autumnal meiosis of the pollen mother cells, is among the 
earliest vernal herbs of deciduous woodlands, often occurring on limestone 
or calcareous soils, from Texas to northern Florida north to Nova Scotia 
and Manitoba. The leaf shape is variable; the seeds are reported to be 
attractive to ants. 

The closest affinity is with the monotypic Eomecon Hance, of eastern 
Asia, which shares with Sanguinaria a preference for the forest habitat, 

* Original determination by the author from squash preparations of microsporocytes 
from plants cultivated in the Beal-Garfield Botanic Garden, Michigan State University, 
East Lansing, Michigan (Gillett 1264, collected 1 November 1961); nine pairs of 
chromosomes were observed in several cells 


1962] ERNST, PAPAVERACEAE & FUMARIACEAE o23 


palmately veined, long-petiolate leaves with broad blades, white petals, 
2-valved, fusiform fruits, arillate seeds, and polyporate pollen; but the 
leaves of Eomecon are cordate and sparsely pubescent, the calyx is syn- 
sepalous, and the inflorescence is a branched scape with each of the 
several flowers pedicellate and subtended by a small bract. 


REFERENCES: 
Under family references see BUKATSCH, GRAY, NoRDHAGEN, and SLAVIK & 
SLAVIKovA (XVII 
BAKER, R. O. The en development, gross structure, ecology and ee 
ical distribution of Sanguinaria canadensis. Nat.-Study Rev. 15: 62-73 
1919,* 


Bastin, E. S. Some further observations on the structure of Samguinaria 
canadensis. Am. Jour. Pharm. 67: 4-9. 1895. 

Curtis, W. Sanguinaria canadensis. Bot. Mag. 5: pl. 162. 1791. 

Gates, B. N. The dissemination by ants of the seeds of bloodroot, Sanguinaria 
canadensis. Rhodora 44: 13-15. 1942. [Seeds red-orange. | 

erg S. Some notes on the pollination of flowers. Bull, Wis. Nat. Hist. 

_ II. 4: 12-21. 1906. [Flowers proterogynous. | 

eo G. A. Alkaloids from Sanguinaria canadensis and their influence 
on growth of Phymatotricum omnivorum, Pl. Physiol. 14: 377-380. 1939. 

GREENE, E. L. ee regarding Sanguinaria. ee 5: 306-308. 1905. 
[Treats 6 spp., 4.n 

Harris, J. A. A mire ane study of the morphology of ue fruit of the 
bloodroot, Sanguinaria canadensis. Biometrika 7: 305-351. 

ion in the inflorescence of Sanguinaria. Biol. eared 30: 
629-633. 1910. 

ae J. W. Juvenile and adult forms of bloodroot. Pl. World 6: 106- 
108. 19 iE lustrates variation in leaves 

Hom, T. Sa to the knowledge ae the germination of some North 
American plants. Mem. Torrey Bot. Club 2: 57-108. 1891. [Cotyledons 
of Sanguinaria hypogea al. | 

Medicinal plants of North America. 18. Samguinaria canadensis L. 
Merck’s Rep. 17: 209-212. 1908. 

Jounson, R. H. Aberrant societies of Sanguinaria and Trillium, Torreya 9: 
5, 6. 1909. [Petals number 6-12, primarily 8.] 

MEEHAN, Sanguinaria canadensis. American blood-root. Meehans’ Monthly 
J oes 900. 


eae J. A. Notes on the seedlings of bloodroot. Am. Midl. Nat. 1: 
199-203. 1910. | Extensive observations on morphology and history. | 
Spencer, W. P. Variation in petal number in the blood-root, See 
canadensis. Am. Nat. 78: 85- a 1944. 

SURFACE, F. M. Contribution to the life history of Sanguinaria canadensis. Ohio 
Nat. 6: 379-385. 1905. |Eight-celled embryo sac; autumnal division of 
microsporocytes. | 


2. Stylophorum Nuttall, Gen. N. Am. Pl. 2: 7. 1818. 


Low, pubescent, perennial herbs with yellow to orange sap, developing 
a stout rhizome; hairs multicellular, terminally uniseriate. Leaves ros- 
ulate, petiolate, deeply 5-7-pinnatifid, pale beneath. Inflorescence um- 


324 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


belliform, few flowered, bracteate, and subtended by 2 or 3 + opposite 
leaves terminating an elongated naked shoot. Sepals 2, attenuate, api- 
cally involute and reflexed, the upper left margin lobed. Petals 4, 2- 
seriate, yellow. Stamens many; filaments narrow, often constricted below 
the oblong anthers; pollen usually with several + obscure apertures. 
Stigma truncate with 3 or 4 shallow lobes; style conspicuous; placentae 
3 or 4 [or placentae 2, stigma deeply 2-lobed, and style short]. Fruits 
ellipsoid, usually 4-valved |or + linear and 2-valved], pubescent and 
nodding; dehiscence basipetal and + complete. Seeds few to many, with 
a conspicuous aril along the upper surface. Lectotype sprcies: S. 
diphyllum (Michx.) Nutt.; see Britton & Brown, Illus. Fl. No. U. S. ed. 
2. 2: 140. 1913. (Name from Greek, stylos, style, and phoros, bearing, 
“from the distinct and conspicuous style which distinguishes this plant 
from all the rest of the Papaveraceae,”’ a distinction no longer true. 


carpellate ovaries, in cross section, diagramma tic, x 4 of, mature fruit 7 
g, four-valved fruit after dehiscence, one valve removed, x 4%; h, arillate seed, 
the micropyle below, < 7 


A genus of two or three species of eastern North America and eastern 
Asia. Stylophorum diphyllum, celandine poppy, wood poppy, mock poppy, 
or yellow poppy, 2” = 20,‘ occurs in rich woods and on bluffs, sometimes 
on calcareous soil, from northern Arkansas and Tennessee to Pennsylvania 
and Wisconsin. The degree of dissection of the leaves is variable. Our 
species, described by Michaux as a species of Chelidonium and reunited 
with Chelidonium by Prain (1895), is remarkably similar in the vegetative 


* Original determinations by the author from squash preparations of microsporocytes 
from a plant cultivated at Stanford University, California (grown from seeds from the 
Royal Botanical Garden, Uppsala, Sweden), and from a plant cultivated at Waverley, 
Massachusetts (£rust 800 [GH]); twenty chromosomes were observed in the former 
and ten pairs were observed in several cells of the latter 


1962] ERNST, PAPAVERACEAE & FUMARIACEAE 325 


stage to the Old World C. majus. Stylophorum, with characteristic com- 
pact inflorescences usually subtended by two + opposite leaves on a naked 
shoot, pubescent fruits with basipetal dehiscence, and pollen with several 
+ obscure apertures, contrasts markedly with Chelidonium majus, in 
which the subumbellate inflorescences arise from branched, leafy stems, 
the dehiscence of the glabrous fruits is + acropetal, and the pollen is 
3-colpate. The closest affinities of S. diphyllum, characterized by ellip- 
soidal, usually 4-valved fruits and + rounded divisions of the leaves, is 
with S. lasiocarpum (Oliver) Fedde (described by Oliver as a species of 
Chelidonium) which has elongated, almost linear, 2-valved fruits, and 
+ pointed divisions of the leaves. 


REFERENCES: 
Under family references see Gray and NORDHAGEN , under Chelidonium see 
PRAIN. 
Boynton, K. R. Stylophorum diphyllum. Addisonia 3: 31, 32. pl. 96. 1918. 
Hooker, W. J. Stylophorum diphyllum. Bot. Mag. 81: pl. 4867. 1855. 
SCHLOTTERBECK, J. O., & C. R. Ecker. The development and structure of the 
seed of Stylophorum diphyllum. Proc. Am. Pharm. Assoc. 50: 401-404. 


3. Chelidonium Linnaeus, Sp. Pl. 1: 505. 1753; Gen. Pl. ed. 5. 224. 


Low, branched, pubescent, biennial or perennial herb with yellow sap, 
developing a stout rhizome or taproot; hairs multicellular, terminally 
uniseriate. Leaves rosulate and cauline, + petiolate below, deeply 5-7 
(or more)-pinnatifid, with rounded |or laciniate| divisions. Inflorescences 
subumbellate, with several flowers, bracteate, terminal and axillary, aris- 
ing from branched, leafy shoots. Sepals 2, somewhat hooded, the upper 


Fic. 2. Chelidonium. a—h, C. majus: a, flowering and fruiting shoot, X %; 
b, opening flower bud, showing emerging stigma, X 3; c, flower, X 3; d, stamen, 

10: e, gynoecium from flower bud just before anthesis, X 10; f, two-valved 
fruit after dehiscence, X 2; g, portion of persistent placenta showing attachment 
of seeds, X 4; h, arillate seed, X 20. 


326 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


left margin lobed. Corolla appearing cruciform; petals 4, 2-seriate, yeilow. 
Stamens several to many; anthers elliptical; filaments narrow, usually 
constricted below the anther; pollen 3-colpate. Stigma 2-lobed; stvle 
short or indefinite; placentae 2. Fruit narrowly oblong or linear, some- 
times curved, 2-valved; dehiscence + acropetal, usually incomplete api- 
cally. Seeds few to many, with a conspicuous aril on the upper surface. 
Seedling cotyledons petiolate. Lectotype species: C. majus L: see 
Britton & Brown, Illus. Fl. No. U. S. ed. 2. 2: 141. 1913; typified by 
removal of other Linnaean species to other genera. (Name from Greek, 
chelidon, swallow, because, according to Aristotle and other scholars, the 
mother-swallows bathed the eyes and thereby strengthened the sight of 
their young with the saffron-colored sap.) — CELANDINE, SWALLOWWORT, 
ROCK POPPY. 


An Old World genus, probably monotypic. Chelidonium majus, 2n = ee 
usually of damp, shady areas, roadsides, rock walls, and waste places, 
widely naturalized in eastern North America, occurs in northwestern North 
Carolina and probably elsewhere in our area. Forms with large flowers or 
with variously dissected or laciniate leaves are occasionally seen in collec- 
tions from both Europe and Asia. 

Linnaeus (1753) included two species of Glaucium and one of Roemeria 
Medic. in his Chelidonium. Later, Prain united Stylophorum, Hylomecon 
Maxim., and Dicranostigma Hook. & Thoms. with Chelidonium: but 
Glaucium, its generic distinction from Dicranostigma debatable, was not 
included. The vegetative stages of Chelidonium majus and Stylophorum 
diphyllum (q. v.) are similar. 

REFERENCES: 

Under a references see BuKATSCH, NoRDHAGEN, and SLAvIK & SLAVI- 
KOVA (XVII 
BANDELIN, : J.. & W. Matesn. Alkaloids of ipa majus, L., leaves 

and stems. I. Jour. Am. Pharm. Assoc. Sci . 45: 1956.* 

BoRATYNSKA, W. Die Variabilitat des rn aes im Schéllleraut (Cheli- 
donium maius L.) unter dem Einfluss des radioaktiven Isotopes 60Co. (In 
Polish; German summary.) Poznan. Inst. Przem, Zielarskiego. Bull. 6(34): 
255- 262. 60. 

Crete, P. A propos de la strophiole du Chelidonium majus L. Bull. Soc. Bot. 
Fr. 91: 88. 1944, [Complains that his work (bid. 84: 196-199. 1937) re- 
garding the epidermal origin of the strophiole was overlooked by Szemes. | 

KaczMarEK, F., & B. MALex. Zur Papier- chromatographie der Alkaloide von 
sided majus L. (English summary.) Planta Med. 7(2): 171-173. 

59. 


ea A. La chélidoine double et la chélidoine laciniée en Belgique. Bull. 
Jard. Bot. Bruxelles 25: 409, 410. 1955. 

Prain, D. A revision of the genus Chelidonium. Bull. Herb. Boiss, 3: 570-587. 
1895. [Combines Chelidonium, Stylophorum, a and Dicrano- 


stigma: oo intimately related to this complex, is not considered. | 
SZEMES, G. ur Entwicklung des Elaisoms von Oaioaaa. majus. Wiener Bot. 
ae 92: 215-219, 1943 


1962] ERNST, PAPAVERACEAE & FUMARIACEAE 327 


Ternpau, M. Die aus Chelidonium majus L. isolierten Alkaloid. (Abstr.). 
Pharm. Zentralh. Deutsch. 97(3): 147. 1958.* 


Subfam. ESCHSCHOLZIOIDEAE Ernst 
4. Eschscholzia Chamisso in Nees, Horae Phys. Berol. 73. pl. 15. 1820. 


Low, annual or perennial herbs with thin yellow to orange sap, sometimes 
developing a stout taproot; usually glabrous (or hairs 1-celled). Leaves 
rosulate and/or cauline, alternate or subopposite, petiolate, + ternately 
finely dissected [or rarely entire]. Flowers and fruits erect on long pedi- 
cels; early flowers usually terminating short shoots, later flowers often 
terminal and axillary on long shoots; receptacles perigynous. Calyx syn- 
sepalous, usually attenuate. Petals usually 4, 2-seriate, yellow or orange 
[sometimes pinkish or white|. Stamens usually many, often adhering to 
base of petals; filaments thick, sometimes with a dark spot; anthers usually 
narrowly oblong; pollen polycolpate with 5-7 [10] slitlike |sometimes 
confluent] apertures. Stigmas 2 to several (the secondary stigmas over 
the placentae); style absent; placentae 2. Fruit columnar, constricted be- 
low the apex, conspicuously ribbed, 2-valved; dehiscence acropetal, in- 
complete, usually violent. Seeds many, reticulate [papillate or variously 
pitted]. Seedling cotyledons linear, bifurcate [but usually entire]. TYPE 
species: Eschscholzia californica Chamisso. (Named for J. F. Eschscholtz,® 
1793-1831, zoologist of Dorpat, friend and companion of Chamisso on the 
Kotzebue expedition which visited California in 1816 on the Rurick.) — 
CALIFORNIA POPPY. 


At least 14 species of northwestern Mexico and the western United 
States, concentrated in California. Perhaps half of the 123 species listed 
by Fedde are referable to Eschscholzia californica, 2n = 12, cultivated 
for its attractive flowers, adventive in many parts of the world, and col- 
lected in our area at least in North and South Carolina; it is distinguished 
by the apparently unique bifurcated cotyledons and the fleshy receptacle 
rim. In nature, as in cultivation, the species is exceedingly variable. 

The 2-valved, deeply ribbed fruits of Eschscholzia, sometimes described 
as polycarpellate, are most easily interpreted as the equivalent of two 
carpels. The longest stigmas are centered over the median carpellary 
traces; secondary stigmas, perhaps a peculiarity of reduction in the vas- 
culature, often develop over the placentae. The combination of synsepaly, 
perigyny, and stigmatic structure suggest that Eschscholzia is relatively 


specialized. 
The genus is allied most closely with the monotypic Hunnemannia 
Sweet, 22 = 56, of eastern Mexico, in which the flowers are somewhat 


perigynous but aposepalous. Synsepaly is also characteristic of Homecon, 


°-The name of the man is usually given thus, but the generic name was rendered 
Eschscholzia in the original description (misspelt Escholzia on the plate) and in the 
later reprint (Linnaea 1: 553-555. 1826), seemingly confirming the intended orthogra- 
phy. 


328 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIIT 


of eastern China, but the affinities of the latter are with Sanguinaria and 
other Chelidonioideae. 

Chromosome numbers of 27 = 12, 14, 22, 24, 34, and 36 have been 
reported 


REFERENCES: 
Under family references see MAHESHWARI & KANTA. 

Hooker, W. J. Eschscholzia californica Cham. Bot. Mag. 56: pl. 2887. 1829. 
[Includes history; see also /bid. 63: pl. 3495. 1863 (as E. crocea).] 

Jepson, W. L. Eschscholtzia. Fl. Calif. 1: 564-575. 1922. [The most complete 
account of variation in E. californica, with observations on transplants. | 

Lewis, H., & F . Went. Plant growth under controlled conditions. IV. 
Response of finns annuals to photoperiod and temperature. Am. Jour. 
Bot. 32: 1-12. 1945. | Brief account of modification of vegetative characters 
in E. californica and E. lobbii. | 

Licnier, O. Eschscholtziées. Explication anatomique de la fleur. Bull. Soc, Bot. 
Fr. 62: 298-315. 1915. 

SACHAR, R. C., & H. Y. M. Ram. The hea of Eschscholzia californica 
Cham, Phy tomorphology 8: 114-124. 1958. | Polygonum-type embryo sac. | 

Saito, K. On a haploid plant eae from the induced autotetraploid plant 
of common California poppy. (In Japanese; English summary.) Jap. Jour. 
Breed. 7(3): 152-156. 1958.* 

Sources, R. Embryogénie des Papavéracées. Développement de Vembryon chez 
PEschscholzia californica Cham. Compt. Rend. Acad. Sci. Paris 229: 485— 
487. 1949 


Subfam. PAPAVEROIDEAE 
5. Argemone Linnaeus, Sp. Pl. 1: 508. 1753; Gen. Pl. ed. 5. 225. 1754. 


Stout, prickly, annual [or biennial] herbs [rarely shrubs] with yellow 
[or white to reddish-orange| sap; prickles and hairs multicellular-multiseri- 
ate, harsh, smooth and polished [or rough and/or branched]. Leaves 
rosulate and cauline, pinnately veined and lobed, sessile, clasping, or some- 
what decurrent, sometimes narrowed into a petiole- like base. Inflorescence 
an expanding cyme, few to several flowered and bracteate. Sepals 2 or 3, 
+ hooded, usually attenuated into a sharp prickle. Corolla 2-seriate, oath 
twice as many petals as sepals (occasionally polypetalous), yellow or litte 
[bronze or lavender]. Stamens numerous; filaments narrow; anthers nar- 
rowly oblong, apically revolute after anthesis; pollen 3-colpate. Stigmatic 
region deeply undulate, the lobes 3 — several (margins of adjacent stigmas 
forming conspicuous lobes over the placentae); style short or obscure; 
placentae as many as stigmatic lobes. Fruit usually more than 3- valved; 
dehiscence basipetal and incomplete. Seeds numerous, reticulate. ee 
ling cotyledons linear-lanceolate. LECTOTYPE SPECIES: A. mexicana L.: 
Britton & Brown, Illus. Fl. No. U. S. ed. 2. 2: 138. 1913. (Name ne an 
herb mentioned by Pliny; perhaps “from the Greek, argema, the cataract 
of the eye for which the juice of a poppy-like plant of the same name was a 
supposed remedy” [Ownbey].) — PrickLy poppy. 


1962] ERNST, PAPAVERACEAE & FUMARIACEAE 329 


About 28 species (Ownbey) lacking clear subgeneric groupings, often of 
disturbed habitats in the New World; one presumed endemic in the 
Hawaiian Islands. Argemone mexicana, devil’s fig, 2n = 28, with yellow 
petals and leaves usually blotched with white along the main veins, 
occasional (mostly in the Coastal Plain) in our area, is a cosmopolitan 
weed of warmer climates, and may be native to the West Indies, Central 
America, and Florida (Ownbey). Forma /eiocarpa (Greene) G. B. Ownbey, 
of southwestern peninsular Florida, where perhaps introduced from Key 
West (Ownbey), is prickly only on the margins of the leaves. It is in- 
frequently collected, and the chromosome number is unreported. <Arge- 
mone albiflora Hornem. (A. alba of authors), with white petals, occasion- 
ally polypetalous, is reported from Florida to Texas and discontinuously 
northward to Illinois and Connecticut. Subspecies albiflora, 2n = 28, with 
oblong to oblong-elliptic capsules and uniform, evenly spaced spines, pri- 
marily of the Coastal Plain, is perhaps native only to northern Florida and 
introduced northward; nian texana G. B. Ownbey, 2” = 28, with nar- 
rowly elliptical capsules and unequal spines and prickles, is native only to 
eastern Texas and is introduced in Arkansas. The two subspecies are com- 
pletely interfertile or nearly so (Ownbey). 

The degree of prickliness, often inconsistent and difficult to describe, is 
apparently of diagnostic importance. Interspecific hybrids, possible be- 
tween many species, can, if the original parents were of the same polyploid 
level, be carried beyond the first generation, but show a decline in vigor and 
fertility. Sterility barriers probably have not played the primary role in 
speciation (Ownbey). Chromosome numbers of 2n = 28, 56, 84, and ca. 
112 have been reported. 

The characteristic hooded sepals terminated by a harsh prickle are 
rarely found outside of Argemone. As in Arctomecon Torr. & Frém., 
Papaver, Roemeria, Stylomecon G. Tayl., some species of Mecano pss 
Vig., and usually in Romneya Harv., the median carpellary traces are lack- 
ing, the fruits are at least three-valved, and the continuous margins of 
adjacent stigmas become conspicuous over the piacentae. 


REFERENCES: 
U family references see GRAY, MAHESHWARI & Kanta, and SLAvIK & 

SLAVIKOVA (VII, XIV). 

Bort, H.-G., & H. FLentje. Alkaloide aus Argemone platyceras. Naturwissen- 
schaften 47: 323. 1960 

Bose, P. K. Cytological studies of Argemone mexicana Linn. Jour. Indian Bot. 
Soc. 16: 197-208. pls. 12, 13. 1937. [Reports normal 8-nucleate embryo 
sac, ovules anatropous with 2 integuments, mature pollen uninucleate; 
suggests polyembryony in Glaucium. | 

CuHopra, R. N., & K. S. Rar. Response of ovules of Argemone mexicana L. to 
colchicine treatment in vivo. Phytomorphology 8: 107-113. 1958. [Some 
seeds with 2 embryos, the smaller possibly haploid. | 

Crete, P. A propos de l’embryologie de l’Argemone mexicana L. Phytomor- 
phology 6: 144-148. : 

Curtis, W. Argemone mexicana. Bot. Mag. 7: pl. 243. 1793. 


330 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


GiraAL, F., & A. SoreLo. Alcaloides de Argemone ochroleuca. Ciencia 19: 67, 
68. 1959 

Hakim, S. A. E. Extraction and detection of poppy alkaloids. Jour. Physiol. 
138: 8P. 1957. | Argemone. | 

———. Poppy alkaloids and glaucoma. J/bid. 40P, 41P. | Argemone. | 

eee G. S., & W. G. Rose. Mexican prickly poppy seed oil. Oil Soap 
20(3): 33- 35. 1943.* 

con. S. kK. Natural hybrid between Argemone mexicana and A. ochroleuca. 
Curr. Sci. Bangalore 29: 282. 1960.* 

Ownbsry, G. B. Monograph of the genus Argemone for North America and the 
West Indies. Mem. Torrey Bot. Club 21: 1-59. 1958. | Illustrations, maps, 
and new chromsome numbers. | 

. The genus Argemone in South America and Hawaii. Brittonia 13: 91- 
109, 1961. 

Pratn, D. An account of the genus Argemone, Jour. Bot. London 33: 129-135, 
176-178, 207-209, 307-312, 325-333, 363-371. 1895. [Monograph; the 
first section a history. Footnote, p. 134: “. . . organically the placentas 
[sic! = stigmas] are in Papaver, as in every other Papaveraceous genus, 
alternate with the placentas.” 

SACHAR, R. C. The embryogeny of Argmone mexicana L.—a criticism. Phyto- 
morphology 6: 148-151. 1956. |The important paper and reference guide 

a controversy with Crété. | 
Sims, J. Argemone albiflora Hornem. Bot. Mag. 49: pl. 2342. 1822. 


6. Papaver Linnaeus, Sp. Pl. 1: 506. 1753; Gen. Pl. ed. 5. 224. 1754. 


Annual [biennial or perennial] herbs, usually with wae sap and pu- 
bescent with multicellular-multiseriate [rarely prickly] hairs. Leaves 
rosulate and/or cauline, pinnately veined and lobed or dissected, usually 
without a distinct petiole. Buds nodding |or erect]; flowers on long 
peduncles subtended by a leaf or bract [or rarely sessile between a pair 
of foliaceous bracts on a scape, or in a cymose spike]. Sepals 2 or 3, 
somewhat hooded. Corolla 2-seriate, with twice as many petals as sepals 
(occasionally polypetalous) white to purple or scarlet [or yellow], 
often with a dark spot toward the base. Stamens many; filaments filiform, 
often constricted below the elliptical or oblong anthers; pollen often 3- 
colpate |or seldom with several apertures]. Stigmatic furrows, usually 
more than 3, appearing radiate, embedded in a flattened or + conical, 
cartilaginous, disclike structure [or disc rarely absent]; placentae as 
many as stigmas. Fruit globose to clavate, truncate, usually more than 
3-valved; dehiscence basipetal and incomplete. Seeds numerous, some- 
what reniform or obovoid-falcate, usually reticulate. Seedling cotyledons 
+ linear. Lectotype species: P. somniferum L.: see Britton & Brown, 
Illus. FI. No. U.S. ed. 2. 2: 136. 1913. (Name oe ae poppy, ap- 
parently derived from papa, pap, thick milk.) — Po 


At least 40 species of the Northern Hemisphere, including in the New 
World the Arctic, the Rocky Mountains, and western California; also 
reported locally in South Africa and Australia. Representatives of three 


1962] ERNST, PAPAVERACEAE & FUMARIACEAE 331 


of nine sections recognized by Fedde, adventive from Europe or garden 
escapes, are to be expected in our area. These include Papaver Rhoeas L. 
(§ OrtHorHorapEes Fedde), 2” = 14, or its cultivated derivative, 
“Shirley Poppy,” with deeply dissected, pubescent leaves and smooth 
fruits about as wide as long; P. dubium L., 2n = 42, resembling P. Rhoeas, 
with fruits about twice as long as wide; and er comm ierumn (S$ PAPAVER, 
§ Mecones Bernh.), opium poppy, 2” = 22, with smooth, globose fruits 
and usually glabrous, broad, clasping, shallowly dissected or incised Sate 
Papaver Argemone L. (§ ARGEMONORHOEADES Fedde), 2n = 42, 
clavate, sparsely pubescent fruits, has been collected immediately to oe 
north of our area. 

Several weedy species of Papaver often associated with the cultivation 
of cereal crops may inhabit the same or nearby habitats in Britain and 
have common insect visitors, but interspecific hybrids are rarely found. 
The relatively self- incompatible diploid, P. RAoezs, and the relatively self- 
compatible hexaploid, P. dubium, can be hy ‘pridized experimentally in 
either direction, the hybrids showing anomalous variation and reduced 
fertility (McNaughton & Harper); bees sometimes discriminate between 
these species. 

The absence of a style and the more than three stigmatic furrows (i.e. 
the margins of adjacent deltoid stigmas) usually embedded in a disc 
characterize the genus. In most species of Papaver, as in those of Roemeria, 
in Meconopsis nirics (L.) Vig., and in the monotypic Séylomecon, the 
median carpellary traces are absent and replaced by pseudomedian traces 
originating from the placental bundles; the vascular bundles of the 
pedicels of some species are + scattered. 

Chromosome numbers of 2” = 12, 14, 15, 21, 22, 28, 40, 42, 44. 56, 70, 
and 84 have been reported. 


REFERENCES: 

Under family references see ARBER and Popov. Many references from the 
large pee have been omitted here. 

AcHESON, R. M., J. L. Harper, & I. H. McNaveHTon. Distribution of antho- 
cyanin oe in poppies. Nature 178: 1783, 1284. 

Anprews, A. C. The opium poppy as a food and spice in the necical period. 
Act ish 2674152—15 5. 1952, 

BocpasHevskata, O. V. Biosynthesis of morphine in poppy (Papaver somnif- 
erum) and its movement in the plant. (In Russian.) Fiziol. Rast. 5(2): 
185-187. 1958.* 

Bort, H.-G., & H. Eumxe. Alkaloide von Papaver paeoniflorum. Naturwis- 
senschaften 45: 315. 1958. 

Cie1j, G. Beet vellows in poppy. Euphytica 10: 225-228. 1961. [Beet virus, 
transmitted by aphids, also infecting P. sommniferum, controllable by s 
temic eta 

CoMMISSION INTERNATIONALE DES INDUSTRIES AGRICOLES. Bibliographie sur la 
morphologie et l’histologie de certaines variétés de pavots (Papaver). 
Commiss. Int. Ind. Agr. Bibliogr. B/2570. 6 pp. 1954.* 

Danert, S. Zur systematik von Papaver somniferum L,. Kulturpflanze 6: 61-88. 
1958. [Describes 52 Varietéten divided into 4 Convarietaten. 


332 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLUI 


Faperce, A. C. Genetics of the Scapiflora section of Papaver. III. Interspecific 
hybrids and genetic homology. Jour. Genet. 46: 125-149. pl. 3. 

FAIRBAIRN, J. W., & L. D. Kapoor. 5 7 ase vessels of Papaver somnif- 

Sri. Planta Med. 8(1): 49-61. 

Harve, J. L., & I. H. McNaucuron. cs inheritance of dormancy in inter- 
and intraspecific hybrids of Papaver. Heredity 15: 315-320. 1960. 

HasirscHKA, G. Bildung von Chromosomenbindeln nach Art der Speichel- 
driisenchromosomen, i igs es Ruhekernchromosomen und andere Struk- 
ear aipeneanchieiian n den endopolyploiden Riesenkernen der Antipoden 
von Papaver Rhoeas. nee 8: 87-113. 1956. 

Hiuis, kK. L., & C. N. RopwEL_. The recombination of some varietal characters 
in the opium poppy. Austral. Jour. Agr. Res. 1: 118-131. 1950. | Quality 
and quantity of opium—morphine content: some plants self-compatible. | 

Hrisui, N. J. Cytogenetical studies on Papaver somniferum L. and Papaver 
setigerum DC. and their hybrid. Genetica 31: 1-130. 

Iv’ina, G. M. Embryological investigations of Papaver somuniterum L. (In 
Rusern English summary.) Bull. Mosk. Obshch. Isp. Pri. Biol. II. 66(4): 
13-25. 

Kant, K. Take -ovarian pollination in Papaver Rhoeas L. Nature 188: 683, 
634. 1960. 

Karp, M. L. On the occasional absence of opium in the opium poppy. Compt. 
Rend. Acad. Sci. URSS II. 52: 537-540. 1946. 

KawatTanlI, T., & H. ASAHINA. External characters and alkaloids of the arti- 
ficial interspecific F, hybrid between Papaver orientale L. (2) and P., 
somniferum L. (6). Jap. Jour. Genet. 34: 353-362. : 

KLEINSCHMIDT, G., & K. Motues. Zur Physiologie und Biosynthese der Alka- 
loide von Papaver somniferum, Zeitschr. Naturf. 14b(1): 52-56. 1959.* 

KNABEN, G. On the evolution of the Radicatum-group of the Scapiflora Papavers 
as studied 7 70 and 56 chromosome species. B. Experimental studies. Op. 
Bot. 3(3): 1-98. pls. 1-6. 1959. 

Lisovsku, G. r Anomalies in the structure of the perianth in Papaver somnif- 
erum L. (In Russian.) Bot. Zhur. 42: 1282-1285. 1957. 

Love, A. Cytotaxonomical remarks on the Icelandic Papaver. Nytt. Mag. Bot. 

-18. 

Tee I. H. Internal breeding barriers in Papaver. Scott. Pl. Breed. 
Sta. Rep. 1960: 76-84. 196?. [Chromosome numbers, some interspecific 
hybrids, and pairing patterns of chromosomes in some hybrids 

. L. Harper. The comparative biology of closely related species 
living in the same area. I. External breeding-barriers between Papaver 
species. New Phytol. 59: 15-26. 1960; II. Aberrant aaa and virus- 
like syndrome in hybrids between Papaver Rhoeas L. and P. dubium L. 
Ibid. 27-41; III. The nature of ee Page sympatric populations of 
Papaver dubium and P. Lecoqit. Ibid. ts 

MauesHwarl, N., & M. Lav. In vitro sa of excised ovules of Papaver 
somiferim L. Phytomorphology 11: 307-314. 1961. 

MIKA, Studies on the growth and development and morphine content of 
opium ee Bot. Gaz. 116: 323-339. 1955. 

Nanpa, K. K. Some observations on the emergence, growth and flowering of 
branches i in Papaver Rhoeas L. Phyton Buenos Aires 16(1): 27-43. 1961.* 

SARKANY, S., & E. Percs. Histogenetical observations in the stem tip of 
ned somniferum L. Acta Biol. Acad. Sci. Hungar. 7(2/3): 183-201. 


1962] ERNST, PAPAVERACEAE & FUMARIACEAE 333 


ScHIJFSMA, L., M. Hoespercen, & F. E. Nrjpam. A study of the colour and 
other characters of the seed of some varieties of oil seed poppy. Euphytica 
9: 127-140. ; 
Sosa-BourpouiL, C. Sur le chimisme des étamines de Pavot. Différences entre 
lignées fertiles et lignées stériles. Compt. Rend. Acad. Sci. Paris 223: 43- 
45, 1946 


STERMITZ, F. Re & H. Rapoport. The biosynthesis of opium alkaloids. Alka- 
loid interconversions in Papaver somniferum and P. orientale. Jour. Am. 
Chem. Soc. 83: 4045-4050. 1961 

True, R. H., & W. W. SrockBerceER. Physiological observations on alkaloids, 
latex and oxidases in Papaver somniferum, Am. Jour. Bot. 3: 1-11. 1916. 

Vototov, E. N. The effect of polyploidy upon the tapetum in poppy. Compt. 
Rend. Acad. Sci. URSS II. 53: 561-564. 1946. 

WiLpeMAN, E. be. La variabilité du nombre de stigmates chez Papaver Rhoeas 
L. Bull. Acad. Sci. Bruxelles V. 31: 459-470. 1946. 

ZaiTsEvA, A. A. On the latex of the opium poppy. (In Russian.) Agrobiologiia 
3: 122-126. 1951.* 


FUMARIACEAE A. P. de Candolle, 
Reg. Veg. Syst. Nat. 2: 105. 1821, nom. cons. 


(FuMIToRY FAMILY) 


Annual, biennial, or perennial, sometimes scandent herbs from taproots, 
bulblets, or rhizomes [or tubers]; plants usually glaucous, rarely pu- 
bescent with unicellular [or multicellular] hairs. Leaves rosulate and/or 
cauline, usually alternate, reduced upward, exstipulate, petiolate, or 
sessile, pinnately or ternately veined, lobed, or dissected. Inflorescences 
terminal and/or axillary, determinate or indeterminate, cymose or race- 
mose, scapose or branched, few [rarely 1| to many flowered; buds erect 
or nodding, bracteate. Flowers erect to nodding, bisexual, hypogynous, 
usually + closed by the apical connation of the inner petals (sometimes 
cleistogamous), bilaterally symmetrical or irregular; perianth cyclic, 3- 
seriate, 2-merous. Sepals 2, bractlike, often + peltate, with + similar 
margins, not inclosing the buds at anthesis [seldom persistent]. Petals 
4, in 2 series, imbricate (very rarely sympetalous) ; outer petals alternate 
with the sepals, 1 or both [rarely neither] + saccate basally; inner 
petals opposite the sepals, characteristically connate over the stigmas |or 
seldom free]. Stamens 6, diadelphous, the phalanges alternate with the 
sepals, sometimes spurred basally and/or nectariferous; anthers di- 
morphic, 2- and 1-locular at anthesis, applied to the stigmatic region [or 
stamens 4, free, anthers 2-locular|]; pollen 3-colpate or several-rugate 
[rarely 2-colpate or polyporate]. Gynoecium syncarpous, of 2 carpels; 
style narrow, persistent |or articulated and deciduous], usually flattened 
apically in the nonplacental plane into a 2- to several-lobed structure 
bearing the stigmatic surface [or seldom attenuated into 2 simple stig- 
mas]; ovary 1-locular [rarely occluded]; placentae parietal, ovules few 
to many, 2-integumented, usually campylotropous, lenticular and + 
reniform [rarely anatropous and obovoid|. Fruit 2-valved [seldom in- 


334 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


flated|, the inner wall of valves often transversely ribbed; dehiscence 
usually incomplete, acropetal or basipetal, the placentae persistent [or 
fruit indehiscent, or very rarely dimorphic]. Seeds 1—many, free [or 
fruit rarely fragmenting into 1-seeded joints], + flattened, reniform or 
orbicular [rarely obovoid], often lustrous, arillate, and attractive to ants; 
endosperm present; embryo small, straight or somewhat curved. Seedling 
cotyledons 2 [1], often petiolate (seldom + linear-lanceolate). (Papa- 
veraceae subfam. Fumarioideae Endl. Gen. Pl. 858. 1839, ““Fumariaceae”’ ; 
including Hypecoum L. and Pteridophyllum Sieb, & Zucc.) TYPE GENUS: 
Fumaria L. 


About 19 genera and perhaps 425 species, primarily of the Northern 
Hemisphere, concentrated in Asia where the species are poorly known, oc- 
curring also in Africa. The family has not been monographed in detail. 

Of the three subfamilies recognized on the basis of floral structure, the 
monotypic Pteridophylloideae Murbeck (restricted to Japan), and the 
monogeneric Hypecoideae Prantl & Kiindig (Eurasian), are distinguished 
by free petals and four free stamens. The genera of subfam. Fumarioideae, 
characterized by diadelphous stamens applied to the stigmatic mechanism 
and the apical connation of the inner petals, fall into two tribes. The 
Corydaleae Reichenb., of worldwide distribution in the Northern Hemis- 
phere, represented in aie area by indigenous species of Dicentra, Adlumia, 
and Corydalis have persistent, nonarticulated styles; the more specialized 
Fumarieae, of limited Old World distribution, represented in our area 
by at least one species of Fumaria, have articulated, deciduous styles. 
The character of the styles seems more important than either the dehis- 
cence or indehiscence of the fruits or the number of seeds in the fruits, the 
criteria by which the two tribes are usually separated. Some of the Old 
World species now referred to Corydalis have deciduous styles, suggesting 
that their taxonomic status should be reconsidered. 

The Fumariaceae, with some medicinal and poisonous properties claimed, 
appear to be as rich in alkaloids as the Papaveraceae, sharing about ten 
per cent of their approximately 140 known alkaloids (Willaman & Schu- 
bert). Of their common alkaloids, eight are reported only for these two 
families, two (with identical empirical formulas) are found in six other 
families, and three (with identical empirical formulas) are found in four 
other families. The sap of the Fumariaceae, somewhat acrid, watery, and 
apparently colorless, is contained in nonarticulated sacs; some authors 
consider the family devoid of laticifers. 

The Fumariaceae and Papaveraceae share bracteate inflorescences, two- 
seriate corollas of twice as many petals as sepals, and similar two-valved 
fruits, the latter most easily understood as the equivalent of two carpels; 
however, the patterns of the vascular traces to the perianths and the 
nature of the sepals, stamens, and seeds usually are different. Hutchinson 
has suggested that the Fumariaceae, nearly as closely allied with some 
Berberidaceae as Papaveraceae, probably have not arisen directly from the 


1962] ERNST, PAPAVERACEAE & FUMARIACEAE SE) 


ancestors of the present Papaveraceae. The least specialized Fumariaceae 
do not closely resemble Berberidaceae, however. 

Chromosome numbers reported for Fumariaceae include 2n = 12, 14, 
16, 18, 20, 24, 28, 32, 48, 56, 64, and ca. 150; somatic numbers of 18 and 
ca. 150 are exceptional. Gametic numbers of 8 and multiples thereof are 
common, while 6 and 7, or multiples thereof, are rare or have not been 
verified in recent years (cf. Papaveraceae); only one report of 2m = 28 
is verified (Ryberg). 


REFERENCES: 


Under Papaveraceae see BANCROFT, BERNHARDI, BERSIER & BoQueET, BERSIL- 
LON, Eames (1961), Eames & WiLson, FEeppe (1905, 1909, 1936), LEGER, 
MurBECK, PAYER, RODER, SAGDULLAJEVA, SAUNDERS, and WILLAMAN & ScHU- 
BERT. 


ARBER, A. Studies in floral morphology. III. On the Fumarioideae, with special 

reference to the androecium. New Phytol. 30: 317-354. 1931. [Analysis 
of fundamental structure of the fumariaceous flower; includes Corydalis, 
Dicentra, and Fumaria.| IV. On the Hypecoideae, with special reference to 
the androecium. /bid. 31: 145-173. 1932. [Establishes morphological rela- 
tionship of Hypecoum to the Fumarioideae. | 

Ernst, W. R. On the family status of the Fumariaceae. (Abstr.) Am. Jour. 
Bot. 48: 546. 1961. 

Gray, A. Ordo Fumariaceae. Gen. Pl. U. S. 1: 117-124. pls. 50-52. 1848. 
[Dicentra, Adlumia, Corydalis. | 

HvuTcHINson, J. e genera of the Fumariaceae and their distribution. Kew 
Bulle 1921 Obs. Loo: 

KLOIMWIEDER, R. Beitrage zur Kenntnis der Schlauchzellen der Fumariaceen, 
speziell der aes Dicentra s.). Akad. Wiss. Wien Sitz-ber. 138: 517- 
55.0) 

LIGNIER, O. a fleur des Crucifeéres comparée a celle des Fumariées. Assoc. 
Fr. Avanc. Sci. Compt. Rend. 25(1): 403-406. 1897. 

MartTEL, E, ane sur le diagramme floral des Cruciféres et des Fumariacées. 
Jour. Bot. Morot 12: 29-31. 1898 

RysBerc, M. A morphological study of the Fumariaceae and the taxonomic sig- 
nificance of the characters examined. Acta Horti Berg. 19: 121-248. pls. 
I-12. 1960. [Indispensable survey of the family; includes chromosome 
numbers and extensive bibliography. | 

SLAVIK, J., & L. Sztavixova. Alkaloide der Mohngewdchse (Papaveraceae). 
XVIII. Alkaloide aus Hypecoum procumbens a und H. eee 
Hook. f. et Thoms. Collect. Czech. Chem. Commun. 26: 1472-147 = 

SouEcEs, R. L’embryon chez le Corydalis cheilanthifoli Hemsl. Les Fumaria 
cées et la classification embryogénique. Ann. Sci. Nat. Bot. 17 
1946. [Comparisons with Medicago, Fumaria, and Bip couny 


KEY TO THE GENERA OF FUMARIACEAE 


General characters: plants herbaceous, with watery sap, sometimes rhizomatous 
or with bulblets (rarely scandent) ; leaves rosulate or alternate, glabrous. In- 
florescence terminal or axillary, bracteate, racemose or cymose; flowers bi- 
sexual, hypogynous, bilaterally symmetrical or irregular; perianth cyclic, 2- 
merous; sepals not inclosing the buds at anthesis; petals 4, two-seriate (very 


336 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLIII 


rarely sympetalous), 1 or both outer petals + saccate basally, the inner petals 

connate apically; stamens 6, diadelphous, 1 or both median stamens basally 

spurred and/or nectariferous; anthers applied to the lobed and flattened stig- 
matic mechanism; gynoecium 1-locular with 2 parietal placentae ; fruit 2-valved 

[rarely ndehiscent|, 

A. Flowers bilaterally symmetrical, nodding, usually bracteolate and bracteate; 
petals usually + free (or sympetalous); outer petals identical; inner petals 
symmetrical. 

B. Plants acaulescent or rhizomatous; petals + free basally; stamen fila- 
ments usually + partially free basally; valves of fruit em basi- 
petally, only the median vein persistent apically. . 

Plants caulescent and scandent; petals united through half ae length, 

spongy and persistent; stamen filaments united basally and adnate to 

corolla; valves of fruit dehiscent acropetally, free at the base. 2. Adlumia. 

A. Flowers irregular, variously ascending or descending, bracteate; petals + 
free at base (or adnate to the stamens); outer petals dissimilar, the upper 
petal conspicuously saccate, the lower sometimes gibbous; inner petals 
ages but asymmetrical. 

. Style persistent; fruit elongate, several seeded, dehiscent + bi are 


<3 


BnTAScOY DIGMMIALS:. d.2 6.6 cod Kaede andwit yaaa aay ee weeaee 3. Corydalis. 
C. sa articulated and dehiscent; fruit globose, hard, 1-seeded, indehiscent ; 
AP vente MRNAS on Sica ca we Rede pe hee eet a eames 4, Fumaria. 


Tribe CoryDALEAE Reichenb. 

1. Dicentra Bernhardi, Linnaea 8: 457, 468. 1833. nom. cons. 

Perennial [sometimes scandent| herbs from rhizomes, bulblets [tubers 
or taproots], glabrous [or very rarely with a few multicellular-multiseriate 
hairs] and occasionally glaucous. Leaves few to many, rosulate [or cauline 
and alternate], pinnately or ternately dissected |petioles very rarely sub- 
terranean and absorptive]. Inflorescence terminal, simple or branched 
{sometimes axillary or leaf-opposed], determinate [or rarely indeter- 
minate]|, cymose or racemose [very rarely with solitary, 1-flowered scapes |. 
Flowers usually nodding [or seldom erect], bilaterally symmetrical, 
usually bracteolate as well as bracteate. Sepals ovate to lanceolate, non- 
peltate. Outer petals identical, + saccate basally and reflexed apically; 
inner petals symmetrical and identical, + spatulate, externally crested 
and apically connate. Stamens in 2 similar phalanges with filaments fused 
or usually partly free basally; median stamens spurred basally and/or nec- 
tariferous; pollen 3-colpate or 12-rugate. Style narrow, persistent, flattened 
apically into a lobed stigmatic structure. Fruit nodding [or seldom erect], 
2-valved, elongate, dehiscence basipetal, incomplete basally, the median 
caepellaey vein persistent apically [or dehiscence acropetal, free at the 
base and incomplete apically|. Seeds few to many, reniform-suborbicular, 
flattened, usually lustrous and + arillate. (Bikukulla Adans. |= Bicu- 
culla of authors], nom. rejic.) Type species: D, Cucullaria (L.) Bernh. 
(Name from Greek, dis, twice, and kentron, a point.) — BLEEDING HEART. 


1962] ERNST, PAPAVERACEAE & FUMARIACEAE 337 


About 19 species and three subgenera distributed in North America and 
eastern Asia. Subgenus DactyLicapNnos (Wall.) K. R. Stern, is scandent 
and confined to northern India and adjacent China; CHrysocaPpNos 
Engelm. occurs in California and eastern Asia; DicENTRA, with one species 
in eastern Asia and several in North America, is represented in our area 
by three species. Dicentra Cucullaria, Dutchman’s breeches, 2n = 32, 
occurring from South Carolina to Arkansas, north to Minnesota and Nova 
Scotia, and discontinuously in the Pacific Northwest, is distinguished by 
white to pink, asymmetrical bulblets and basally fused stamen filaments 
with the nectariferous spur directed diagonally; D. canadensis (Goldie) 
Walp., squirrel-corn or turkey-corn, 2” = ca. 64, flowering somewhat 
earlier and sympatric almost throughout the eastern distribution of the 
above, has yellow, pea-shaped bulblets and filaments partially free basally 
with the short nectariferous spur oriented vertically. These two, seem- 
ingly most closely related to each other, have similar scapose, racemose 
inflorescences terminating blindly in a linear lobe, an aborted bud, or a 
flower. A strictly racemose inflorescence, also occurs in D. spectabilis 
(L.) Lem., 2n = 16, of Asia. Dicentra eximia (Ker) Torr., bleeding 
heart, 27 = 16, with cymose, usually branched inflorescences, strongly 
allied with D. formosa (Haw.) Walp., 2n = ?24, of the Pacific States, is 
distributed from eastern Tennessee and western North Carolina and 
northward to New Jersey and Pennsylvania. 

The flowers of Dicentra, as in most Fumariaceae, are somewhat tubular 
and closed by the apical connation of the inner petals over the anthers 
and stigmas. Although abundant pollen is shed directly on the stigmas 
and adheres to them, the germinating pollen tubes barely penetrate the 
stigmatic tissue in a number of species (Stern, 1961). In some, ovaries 
abort if protected from the visits of insects; bees sometimes gain direct 
access to the nectaries by biting through the base of the corolla. 


REFERENCES: 


Under family references see GRAY; under Papaveraceae see GATES. 


Camp, W. H. Dicentra Cucullaria. Addisonia 20: 21, 22. pl. 651. 1937. 
Hooker, W. J. Dielytra canadensis. Bot. Mag. 57: pl. 3031. 1830. [Dicentra 


MEEHAN, T. Dicentra Cucullaria. Meehans’ Monthly 6: 41, 42. 1896. [Brief 


history. | 

Roginson, C. Flowers and insects. I. Bot. Gaz. 14: 120-126. 1889. [Insect 
visitors of D, Cucullaria and biting of holes through corolla. ] 

Rypperc, P. A. Bicuculla eximia. Addisonia 14: 53, 54. pl. 474. 1930. [D. 
eximia. | 

Stms, J. Fumaria Cucullaria. Bot. Mag. 28: pl. 1127. 1808. [D. Cucullaria.] 

Seay K. R. Revision of Dicentra Ree Brittonia 13: 1-57, 1961. 
[Includes illustrations, distribution maps, and review of alkaloidal prop- 
erties with literature citations, 10-12; treats meee as a subgenus 
of Dicentra. See pollen, Am. Jour. Ree 49: 362-368. 1962.] 


338 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLIII 
2. Adlumia Rafinesque ex A. P. de Candolle, Reg. Veg. Syst. Nat. 2: 


¢ 


111. 1821, nom. cons.* 


Delicate, taprooted biennial vine usually with a single, indeterminate 
leader. eave: pinnately divided, at first rosulate, then cauline and al- 
ternate, the petiolules + opposite, distally tendriliform. Inflorescences 
many, | or 2 ina leaf axil, cymose, often branched, usually with many nod- 
ding, bracteate flowers. Sepals somewhat peltate. Corolla sympetalous, 
white to purplish, persistent and spongy, bilaterally symmetrical, basally 
+ saccate alternate with the sepals, apically with 2 reflexed outer lobes, 
and 2 connate inner lobes. Stamens 6, diadelphous, but basally + mon- 
adelphous and united with the corolla; median stamens nectariferous 
basally but not spurred; pollen 3-colpate or 6-rugate [9—12-rugate]. Style 
narrow and persistent, terminating in a flattened, + 2-lobed stigmatic re- 
gion. Fruits + linear, 2-valved, dehiscence acropetal, the valves free 
basally. Seeds few, flattened. suborbicular, lustrous, nonanilate: Seedling 
cotyledons 2, petiolate. TyPE SPECIES: A. cirhosa Raf. ex A. 
fungosa (Ait.) Greene ex BSP. (Named for John Adlum, 1759-1836, sol- 
dier of the Revolution, major under the elder Adams, later brigadier gen- 
eral of the Pennsylvania Militia; enthusiastic advocate of American viti- 
culture. ) 


A genus, probably monotypic, of eastern North America and Korea. 
Adlumia fungosa, Allegheny vine, mountain fringe, or climbing fumitory, 


(1904, p. 343), who stated that his specimens, insofar as he could de- 
termine, were identical to the American species, a conclusion substantiated 
by his description. Ohwi (1931, p. 387), described A. asiatica, but the 
description and plate are indistinguishable from the American plant. The 
name A, Komarovii was used by Popov (1937, posthumous 
The unique, sympetalous corolla of Adlumia may culminate a trend 
toward the tubular condition evident in the scandent species of Dicentra 
and in the analogous, long-spurred petals of some species of Corydalis. The 
affinities of Adlumia are with Dicentra subg. DactyLicapnos, which also 
has tendrilliform leaflets, scandent growth, and acropetal dehiscence of the 
fruits, the valves becoming free basally. Current growth of other species of 
Dicentra is not scandent, lacks tendrilliform leaflets, and is terminated by 
an inflorescence; and dehiscence of the fruits is basipetal, the valves re- 
maining attached apically only by the filamentous median carpellary vein 
and basally where no abscission zone forms. Stern favored the retention 
both of Adlumia (indeterminate-monopodial growth with axillary, cymose 
inflorescences, sympetalous corolla and nectariferous but nonspurred 
median stamen filaments) as a genus and of DactyLicapNnos (indetermi- 
mia was unnecessarily conserved over Bicuculla Borckh., a later homonym of 
Bikukulla Adans. which is equal to Fumaria Cucullaria L. and now equivalent to the 
type species of the conserved Dicentra Bernh. (See Rickett & Stafleu, Taxon 8: 
274, 1959.) 


1962 | ERNST, PAPAVERACEAE & FUMARIACEAE 339 


nate-sympodial growth with leaf-opposed, racemose inflorescences, apopet- 

alous corollas, elaborately spurred and nectariferous median stamen fila- 

ments) as a Subgenus of Dicentra. 

REFERENCES: 

Under family references see GRAY. 

Britton, E. G. Adlumia fungosa, Addisonia 5: 21, 22. pl. 171. 1920. [Re- 
see garden plants annual, attaining a length a 10 feet. 

Komaroy, V. L. Baas Zn: Fl. Manshuriae. vol. 2. (In Russian.) 
cere Horti Petrop. 22: 337-352. 1904. [Reports discovery in 1897 in 
Korea of Adlumia, a and, as far as could be determined, identical to 
the American 

Onwt., J. Adlumia gee sp. nov. Zz: Symbolae ad Floram Asiae Orientalis 
Il n Latin.) Bot. Mag. Tokyo 45: 387. 1931. ne the sp. for 
Korea, (2) biennial; does not seem to differ from A. fungos 

Popov, M. G. Papaveraceae. In: Fl. URSS 7: 573-717. 1937. (In Russian.) [A. 
asiatica Ohwi given as perennial, 647; pl. 45, fig. 6 


3. Corydalis Ventenat, Choix de Plantes. pi. 19. 1803, nom. cons.!° 


Annual, biennial [or perennial, very rarely scandent] herbs, from tap- 
roots [tubers or rhizomes], often glaucous, usually glabrous (or very 
rarely with 1-celled hairs). Leaves variously pinnately or ternately dis- 
sected, rosulate or cauline, alternate, petiolate toward the base. Inflores- 
cence terminal, simple or branched, few to many flowered [or scapose] 
usually racemose (or very rarely cymose); flowers irregular, often 
horizontally oriented, bracteate. Sepals ovate, + peltate. Outer petals 
dissimilar, the upper conspicuously saccate, the lower sometimes + gib- 
bous; inner petals similar but asymmetrical, connate apically. Stamen 
filaments fused in 2 dissimilar phalanges, the upper median stamen spurred 
and nectariferous; pollen 3-colpate or 6—12-rugate. Gynoecium elongate, 
usually upwardly curved; style persistent [or seldom articulated and 
deciduous], apically flattened into a lobed stigmatic structure. Fruit erect 
or nodding [sometimes geniculate at the receptacle], usually elongate, 
2-valved, sometimes falcate, often torulose [or very rarely inflated], 
dehiscence + acropetal, the valves free at the base and often transversely 
ribbed between the seeds. Seeds few to many, flattened, reniform or + 
orbicular, lustrous, and usually arillate. Seedling cotyledons 2 [or 1], 
usually long-petiolate. Lecrorype species: C. sempervirens (L.) Pers. 
(Name from Greek, korydos, or korydaillis, the crested lark of Greece, and 
presumably the name of a plant with a corolla spur resembling the spur 
of the lark.) 

* When first conserved, Corydalis was attributed to Medicus who applied the name 
to Fumaria vesicaria L., now Cysticapnos vesicarius (L.) Fedde. To preserve the 
modern concept of Corydalis, the authority was changed to Ventenat who stated in a 


ournefort and Gaertner. Tournefort cited Fumaria siliquosa sempervirens of Corn uti, 
presumably now equivalent to Corydalis sempervirens oe Pers., the lectotype species. 
Gaertner presumably illustrated Corydalis lutea (L.) D 


340 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLUI 


Fic. 3. Corydalis. a-k, C. sempervirens: a, flowering and fruiting shoot and 
basal leaf, X Y%: b, flower, lateral view — note peltate sepal, X 4; c tip of 


stamens, the lower two-locular, the two upper one-locular, * 10; h, gynoecium 

and pedicel, lateral view, X 45 i i, lobed stigmatic region, lateral view, Oe 4 

fruit after dehiscence, show z persistent style, placentae, and attachment. of 
seeds, X 1%; k, arillate seeds the micropyle at left edge below aril, * 20 


A genus of the Northern Hemisphere, probably of fewer than 300 species, 
concentrated in south-central Asia (where poorly known), with one species 
in East Africa, variously segregated without agreement into several sections 
and subsections. 


1962] ERNST, PAPAVERACEAE & FUMARIACEAE 341 


In our area, five taxa of sect. Coryparis (§ encom aa Prantl) are 
widely distributed (Ownbey). Corydalis sempervirens, 2n 16, pink or 
pale corydalis, rock fumeroot, or roke harlequin, a biennial ec anpated from 
northern Georgia to Newfoundland and central Alaska, tentatively re- 
ported from Siberia (Fl. USSR.), is distinguished by its branched, leafy 
shoots, pinkish flowers, and apparently unique cymose inflorescences. Four 
winter annuals bearing a general resemblance to C. aurea Willd., 2n = 
16, the latter widespread to the west and north of our area, were defined by 
Ownbey (1947): C. crystallina Engelm., with conspicuously pubescent 
fruits, occurring in Arkansas and the states immediately west and north; 
C. micrantha (Engelm.) Gray subsp. micrantha, 2n = 16, slender fume- 
wort, with erect fruits, occurring in northwestern Arkansas northward 
and westward; C. micrantha subsp. australis (Chapm.) G. B. Ownbey, 
intergrading with subsp. micrantha, usually with + longer racemes and 
flowers, and longer, narrower fruits, occurring from Arkansas southeast- 
ward along the Gulf Coast, across Blond and north along the Atlantic 
Coast to North Carolina; and C. flavula (Raf.) DC., yellow fumewort or 
yellow harlequin, similar to C. micrantha subsp. australis, with nodding 
fruits, occurring from Louisiana to Missouri eastward to North Carolina 
and Connecticut. 

The taxon Corydalis micrantha subsp. australis, first described as C. 
aurea var. australis Chapm., was considered conspecific with C. micrantha 
(Engelm.) Gray (originally C. aurea var. micrantha Engelm.), and re- 
described as Capnoides Halei Small (with part of the original material 
referable to Corydalis crystallina), later Corydalis Halei (Small) Fernald 
& Schubert. 

Both Corydalis micrantha (including subspp. micrantha and australis) 
and C, flavula are complicated biologically by the occasional occurrence of 
individual, or whole inflorescences of, cleistogamous flowers, apparently 
sometimes correlated with certain vegetative peculiarities. Normal flowers 
in this instance are closed by the permanent apical connation of the inner 
petals, and the stamens shed pollen on the stigmatic mechanism to which 
they are appressed. In some species the stigmatic region can be exposed 
suddenly by probing along the path leading to the nectariferous spur. 

Chromosome numbers of 2m = 12, 14, 22, 24, 34, and 36 have been re- 
ported. 


REFERENCES: 

Under family references see Gray; under Papaveraceae see FEDDE (1936). 
BERSILLON, G. Le renouvellement annuel du tubercule de Corydalis solida 
Sw.; mécanismes histogénétiques. Revue Gén. Bot. 66: 469-488. 1959. 
Bort, H.-G., & H. Eumxe. Alkaloide aus Corydalis angustifolia und C. de- 

cipiens. ere 46: 427. 1959 
Curtis, W. Fumaria glauca. Bot. Mag. 5: fl. 179. 1792. [C. sempervirens. ] 
FERNALD, M. L. TaonaGane and reidentifications of North American plants. 
Rhodora 48: 207-216. 1946. [History of Corydalis Halei, 207. 
Gray, A. Corydalis aurea and its allies. Bot. Gaz. 11: 188, 189. 1886. 
MansrFELp, R. Bemerkungen zur Liste der Nomina Generica Conservanda. 


342 . JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLIII 


a German.) Taxon 2: 187-190. 1953. [Ventenat, rather than Medicus, 
ust be the authority for proper conservation of the name Corydalis. | 
isan M.R. The toxicity of Corydalis Caseana, Jour. Agr. Res. 42: 239-243. 
1931. 


Ownsey, G. B. Monograph of the North American species of Corydalis. Ann. 
Missouri Bot. Gard. 34: 187-259. 1947. [Includes citation of specimens 
and distribution maps. | 

n the cytotaxonomy of the genus Corydalis, section Eucorydalis. Am. 
Midl. Nat. 45: 184-186. 1951. | Reports from root tips 2” = 16 for C 
aurea, C. micrantha, and C, sempervirens; identical chromosomal mor- 


RYBERG, M. Studies in the taxonomy and fertility of some Scandinavian Cory- 
dalis species of the sect. Pes-Gallinaceus Irmisch. Acta Horti Berg. 15: 
207-284. 1950. [Analysis of variation and breeding behavior in northern 
European spp. of Corydalis; fall meiosis. | 

A taxonomical survey of the genus Corydalis Ventenat with reference 
to cultivated species. bid. 17: 115-175. pls. eS 

A morphological study of Corydalis nobilis, C. cava, C. solida, and 

some allied species with special reference to their underground organs. 
Ibid. 19: 15-119. pls. 1-16. 1959. 

SovEcEs, R. Embryogénie des Fumariacées. Les premiers ee du deével- 
oppement de l’embryon chez le Corydalis lutea DC. Compt. Rend. Acad. 
Sci. Paris 222: 161-163. 1946. La différenciation des Ras foudamient: ales 
du corps chez le Corydalis lutea DC. Ibid. 253-255. 

SPRAGUE, T. A. The correct spelling of certain generic names — III. Kew Bull. 

928: 337-365. 1928. [Review of use and spelling of the name Corydalis. | 

TraBerT, C. H., & U. SCHNEIDEWIND. Uber Inhaltsstoffe von Corydalis cava 
Schweigger und Koerte, 2. Die Untersuchung der Samen. Pharm. Zentralh, 
Deutsch. 99(1): 2-7. 1960.* 


Tribe FUMARIEAE 
4, Fumaria Linnaeus, Sp. Pl. 1: 699. 1753; Gen. Pl. ed. 5. 314. 1754. 


Low, glabrous, compact or diffuse herbs from taproots. Leaves rosulate 
and/or cauline, alternate, variously pinnately or ternately dissected |some- 
times + tendriliform]. Inflorescences many flowered, terminal or axillary, 
indeterminate, and racemose; flowers irregular, bracteate. Sepals small, 
peltate-ovate, variously dissected. Outer petals dissimilar, the upper sac- 
cate at the base; inner petals similar but asymmetrical, connate apically. 
Stamen filaments fused in 2 dissimilar phalanges, somewhat adnate later- 
ally with the base of the petals; upper median stamen spurred and _nec- 
tariferous; pollen polyporate with 6-12 apertures. Style narrow, + 
ascending, articulated and dehiscent, apically flattened into + 2 lobes; 
ovary short, + ovate; ovules 4, + reniform, Fruit subhemispheric, often 
with 2 shallow depressions apically, hard, indehiscent, filled with the single 
nonarillate seed. Seedling cotyledons linear-lanceolate, nonpetiolate. 

YPE SPECIES: F. officinalis L. (Name from Latin, fwmus, smoke, perhaps 
alluding to the odor of the plant.) — Fumirory, EARTH-SMOKE, FUME- 
ROOT, FUMEWEED, FUMEWORT. 


1962] ERNST, PAPAVERACEAE & FUMARIACEAE 343 


A genus of cosmopolitan weeds, concentrated in the Mediterranean 
region, with many species and infraspecific taxa; represented in our area 
by Fumaria officinalis, 2n = 32 (also ?14 and Oy occurring from western 
Florida along the Coastal Plain into North Carolina. Other species, the 
differences appearing trivial in view of the apparent edaphic plasticity of 
the genus, may occur in our area. The environment readily modifies the 
character of the vegetative organs and, under some conditions, the fre- 
quency of cleistogamous flowers which are paler or whitish in color, re- 
duced in size and lacking nectaries but producing fruit. Hybrids in the 
Old World are uncommon, isolated, and sterile (Pugsley). Chromosome 
numbers of 2” = 14, 28, 32, ca. 48, 56, and 64 have been reported. 


REFERENCES: 

GreceER, J. Uber Kalkoxalatkristalle in der Samenschale von Fumaria officinalis 
L. Planta 12: 49-52. 1930 [1931]. 

Lerort, L. Quelques groupes infra-spécifiques luxembourgeois de Fumaria 
Di aerttes Ib, Soc. Nat. Luxembourg. Bull. II. 44: 180, 181. 1950 [1951].* 

Pucstey, H. W. A revision of the genera Pumaria and Rupicapnos. Jour. Linn. 
Soc. Bot. 44: 233-355. pls. 9-16. 1919. 

SouEGES, R. Embryogénie des Fumariacées. L’origine du corps de l’embryon 
chez le Fumaria officinalis L. Compt. Rend. Acad. Sci. Paris 213: 528-530. 
194] 


344 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLIII 


NOTE ON DAPHNOPSIS CRASSIFOLIA (THYMELAEACEAE) 


Lorin I. NEVLING, JR. 


AMONG A NUMBER OF SPECIES which were poorly or incompletely known 
at the time of my revision of the genus Daphnopsis (1959) was D. crasst- 
folia (Poiret) Meissner. The description of this species necessarily was 
sketchy and incomplete because of the availability of relatively few and 
fragmentary specimens. I was not able to see fruiting material, for ex- 
ample, although it had been described previously. Since my publication, 
a relatively large number of specimens referable to this species have been 
rediscovered and examined. On the basis of new data drawn from re- 
cently examined herbarium material, the description has been recast and 
completed. In addition, these collections give the first precise knowledge of 
the range of the species (see Map). 


Daphnopsis crassifolia (Poir.) Meissn. in DC. Prodr. 14: 524. 1857. 


Daphne crassifolia Poir, Encycl. Méthod. Bot. Suppl. 3: 316. 1813 (Tyee: 
Nectoux sn. [é]!). 

Daphnopsis oe var. eggersii Krug & Urb. Bot. Jahrb. 15: 350. 1892 
Type: Eggers 2317 [9 ]). 

Hyptiodaphne crassifolia (Poir.) Urb. Symb. Ant. 2: 454. 1901. 

Hyptiodaphne crassifolia var. 8. eggersii (Krug & Urb.) Urb. ibid. 455. 


Dioecious shrubs or trees, to 4 m. tall, the young branches terete, densely 
ochraceous-hirsute and glabrescent, rugose. Leaves simple, approximately 
whorled by irregular condensation of internodes or rarely alternate on 
very vigorous shoots, usually 3 or 4 per whorl; leaf blades rarely narrowly 
elliptic to elliptic to oblanceolate to obovate or very reduced and bractlike, 
the expanded blades 3—6(—13.5) cm. long, 1-3.5 cm. broad, acute to 
acuminate or rotund and mucronulate at the apex, cuneate to subauriculate 
at the base, coriaceous, (sparsely) to densely ochraceous-sericeous and 


neath, arcuate-ascending and forming a prominent submarginal vein; 
petiole 2-4 mm. long; exstipulate. Inflorescences borne terminally (but 
appearing axillary or extra-axillary) from young bracteate stems, umbel- 
liform to subracemiform, hirsute, the primary peduncle 10-40 mm. long, 
the rachis 1-3 mm. long, the secondary peduncles 3-7 mm. long. Flowers 
unisexual, 4-merous, perigynous. Staminate flowers: 2—5 per inflorescence, 
white; pedicel obsolete to 0.5 mm. long; calyx tube tubular to broadly 
obconic, 7-10 mm. long, 2—4.5 mm. broad at the orifice, hirsute without, 
glabrous within; calyx lobes subequal, indefinitely papillate within, 2.5—4 
mm. broad; petals connate into an obscure faucal annulus; antisepalous 


1962] NEVLING, DAPHNOPSIS CRASSIFOLIA 345 


stamens inserted at the orifice, subexserted, the alternisepalous inserted 
2-3 anthers’ lengths below the orifice, included, the anthers oblong, about 
1 mm. long, 0.5 mm. broad, sessile; disc of irregular lobes, free, 1—3.5 mm. 
tall, glabrous; pistillode tenpin-shaped, 0.75-1.5 mm. tall, glabrous. Pistil- 
late flowers: 2 or 3 per inflorescence, white; pedicel 1-1.5 mm. long; 
calyx tube more or less tubular, 4-8.5 mm. long, 2 mm. broad at the orifice, 
hirsute without, glabrous within; calyx lobes unequal, indefinitely papillate 
within, the outer 3—4.5 mm. long, 2.5—3.5 mm. broad, the inner 2.5-3 mm. 
long, 2 mm. broad; petals connate into an obscure faucal annulus; stami- 
nodia 8, papilliform, minute; disc of 4 lobes, the lobes irregularly shaped, 
free, 1-2 mm. tall, glabrous; gynoecium single, tenpin-shaped, 4.5—-5.5 
mm. long, the ovary 1-locular, glabrous, the single ovule anatropous, the 
style ca. 1.5 mm. long, thick, the stigma capitate, obscurely bilobed, in- 
cluded or very rarely exserted. Drupe ellipsoid, ca. 15 mm. long, 10 mm. 
in diameter, glabrous. 


Endemic to the island of Hispaniola where it is abundant on lateritic 
soils, according to Ekman. It has been collected along river banks, in 
thickets and woods from 600 to 2000 m. altitude. Flowering is known 
from January through May and October; fruiting in April and May. The 
tough fibrous bark is used, according to Holdridge, for making rope. It 
is known locally as mahaut (Holdridge) and jayao (Jaquez). 


Map of Hispaniola showing distribution of Daphnopsis crassifolia. 


Dominican Republic. Azua: Sierra de Ocoa, San José de Ocoa, Loma del 
Rancho, Ekman H11641 (us, 6). BaraHona: Barahona, Fuertes 891 (a, 
sterile); Firme Noche Buena, Fuertes 1433 (a, NY, 6), 1498 (a, Ny, US, 2) 
BENEFACTOR: Piedra del Aguacate, n. of Rio Arriba, Howard & Howard 9436 
(A, ?). La VecA: prope Constanza, Von Tiirckheim 3103 (cu, ?; Ny, 6, 9; 
us, ¢, @). Without precise locality: Pico de Igua, Jaguez 2028 (us, @). 
Haiti. ARTIBONITE: Massif du Nord, Hinche, Ekman H11641 (Ny, 6); Massif 
des Bahos, Pte- Rio de l’Artibonite, Perodin, bridge above Ingrand, Ekman 
H3454 (us, 6). OUEST: vicinity of Mission, Fonds Varettes, Leonard 3905 


346 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


(GH, Ny, US, 6); Massif de la Selle, Petionville, top of Mt. Tranchant, Ekman 
H1159 (vs, 9): Massif des Cahas group, Las Caobas [Cahobas], Belladire 
[ Belladére|, Morne Lagoume-Hire (?), Ekman H35627 (ny, 6); Massif de la 
Selle. group Créte-a-Piquants. Port-au-Prince, Morne, Bercy, erupt. Ekman 
H5910 (xy, 6); Gros Cheval, Mornes aes Comnmissaires, Holdridge 951 (GH, 


MO, NY, US, : ). Without precise locality: La Revellue (?), collector unknown 
sm. (C, 2); “Saint-Domingue,’ Nese sn. (P- ae pe of Daphne crassifolia, 
3 ) 

The precise locality at which the holotype of this species was col- 


lected by Nectoux is unknown: consequently, it has not been mapped. 
Nectoux was “Jardinier en chef du Jardin du Roi” (Nectoux, 1791) in 
what is now Haiti. I have been unable to ascertain to what extent he 
travelled and collected. 

Ekman’s number 11641 is a split collection, part collected in the 
Dominican Republic and part in Haiti. Both localities have been mapped. 

The included stigma is an unusual feature of the pistillate flower of this 
species. The only exception I have seen, in which the stigma is exserted, 
is on Von Tiirckheim 3103 (us). In this specimen the gynoecium is of 
average size, but the calyx tube is relatively short for the species, and 
this may account for the exsertion of the stigma. 

Krug and Urban’s Daphnopsis crassifolia var. eggersi is based on an 
Eggers specimen (2317) which is larger in respect to petiole length and 
leaf size than var. crassifolia. They give the following measurements for 
var. crassifolia: petiole 2 mm. long; leaf blade 3-3.5 cm. long, 1.3-1.7 
cm. broad. In contrast, for var. eggersii they give: petiole 3-4 mm. long: 
leaf blade 5—8 cm. long, 2—3 cm. broad. These are the only contrasts, for 
var. crassifolia was known from staminate material and var. eggerst only 
from fruiting material. The measurements of leaf and petiole may be 
applied successfully to separate Nectoux s.n. (type of var. crassifolia) 
from a few specimens (the holotype of var. eggersi (Eggers 2317), was 
destroyed at Berlin and T have not located an isotype); but many speci- 
mens display enough variability on a single branchlet to overlap sig- 
nificantly the measurement ranges given for the two varieties. I believe 
that the leaf variation exemplified by this species is of a phenotypic, 
rather than of a genotypic nature. 

In 1901, when Urban erected the genus Hyptiodaphne to accommodate 
Daphne crassifolia Poir. (= Daphnopsis crassifolia (Poir.) Meissn.), he 
indicated as one of the main features of the new genus the orthotropous 
ovule attached basally in the ovary. Unfortunately, there was (and is) 
little or no information available concerning ovule position and attach- 
ment within the genus Daphnopsis except that the ovules are anatropous. 
Domke has figured (1934, figs. 39a,6,c) several ovules from plants refer- 
able to Daphnopsis crassifolia which show, quite strikingly, considerable 
variation in ovule attachment. The ovules, however, are always of the 
anatropous type. A pistillate flower (Holdridge 951, GH) was cleared and 
stained to recheck this point. The ovary wall is heavily vascularized with 
veinlets composed of short and sinuous tracheary elements. The ovular 


pee 


1962 | NEVLING, DAPHNOPSIS CRASSIFOLIA 347 


trace is developed poorly, but the trace and ovule position is nearly 
identical with that figured by Domke in Fig. 396. If Urban based his ob- 
servation on sectioned material he may have sectioned the ovule at right 
angles to the attachment, thus leading to the false impression of an ortho- 
tropous ovule (demonstrated in Domke’s fig. 39c). Regardless of these 
difficulties, a slight shift in ovule position alone is not enough to warrant 
recognition of Hyptiodaphne, 


LITERATURE CITED 


DomkeE, W. Untersuchungen uber die systematische und geographische Glhede- 
rung der Thymelaeaceen. Bibliot. Bot. 27(111): 1-151. 

Nectoux, M. Observations. Mém. d’agric. d’econ. rurale et domestique 110-123. 
1791. | Reference incomplete; seen only as a separa 

Neviinc, L. I., Jr. A revision of the genus Dies Ann. Missouri Bot. 
Gard. 46: 257 —358. 1959. 


348 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


TWO NEW ASIATIC PANDANACEAE 


BENJAMIN C. STONE 


DuRING A SHORT visit to the Harvard University Herbarium I was 
enabled, through the courtesy of the directors, to study the collections of 
Pandanaceae. Among the specimens studied were two which represent 
undescribed species, one in the genus Freycinetia, the other in Pandanus. 
Descriptions of these follow. 


Freycinetia kostermansii B. C. Stone, sp. nov. (Sect. FREYCINETIA), 
(Fie.1); 

Planta scandens, caulibus subrobustis subteretibus, plerumque 8-9 mm. 
crassis, internodiis 5-15 mm. longis; foliis obovato-oblanceolatis, basi 
breviter contractis, apice abrupte acuminatis breviter caudatis, vulgo 10- 
15 cm. longis et 12-28 mm. latis, margine basim versus serrato-dentatis, 
dentibus ad 2 mm. longis, et apicem versus serrato-denticulatis (dentibus 
ad 0.5—1 mm. longis), costa media ad apicem et in acumine denticulata; 
syncarpiis terminalibus cylindricis usque ad 8 cm. longis et 3 cm. latis, 
pedicellis in angulis et distaliter scabridulis; baccis rubris, 14-16 mm. 
longis, circiter 4 mm. latis, lageniformibus, apice rostrato-contractis, 
truncatis (areola stigmatica minima 1-2 mm. lata), parte inferiore succu- 
lentibus, stigmatibus (5—)6—-9, seminibus circiter 1.6—-1.9 mm. longis line- 
aribus rhaphide et strophiolo instructis. 

Hototype. Moluccas: Morotai, 1000 m. alt., May 27, 1949, A.J. G. H. Koster- 
mans 1193 (A). 

This new species is apparently related to Freycinetia oblanceolata Mar- 
telli, of New Guinea, and perhaps also to F. Aemsleyi Warb., of Borneo. 


Pandanus gressittii B. C. Stone, sp. nov. (Sect. ACROSTIGMA). 


P. Gressittii Merr. ex G. F. Sauer, List Pl. Lingnan Univ. Campus 94, 1947, 
nomen nudum. 


Frutex foliis circiter 30-101 cm. longis et (7—) 11-14 mm. latis, margine 
robuste armatis dentibus antrorsis acicularibus, costa media dorsaliter 
armatis dentibus robustis antrorsis, apicem versus adpressibus; syn- 
carpium solitarium, ellipsoideum, circiter 5-7 cm. longum et 3.5-5 cm. 
diametro; pedunculo erecto bracteato; drupis anguste obclavatis circiter 
14 mm. longis et 2-3 mm. latis, apice rotundato-conicis, pileo 2-3 mm. 
longo; stigmate breviter erecto-curvato, aceroso, circiter 2 mm. longo et 
0.75 mm. lato, endocarpio supramediale, semen unicum. 


DISTRIBUTION: endemic to Hainan Island. 


1962] STONE, NEW ASIATIC PANDANACEAE 349 


Fic. 1. FREYCINETIA KOSTERMANSIT: a, leaf. X 0.8; b, ¢, berry, lateral and top 
views, X 8; d, seed, X 8 and & 35. (Drawn from the hatety pe.) 


Ho.Lotype. Hainan Island: small jungle stream at Liamui, 400 m. alt., June- 
July 1935, J. Linsley Gressitt 1185 (A). ADDITIONAL SPECIMENS EXAMINED. 
Lokwui, near stream, 15 May 1935, F. C. How 72350 (a); Bo-Ting, in thickets, 
17 Oct. 1936, S. K. Law 28026 (A). 

The existence of this species was first called to my attention by Dr. 
J. Linsley Gressitt, who collected the type specimen. The name conferred 
by Merrill and long carried in manuscript appears never to have been 
formally published. I take pleasure in bringing into use the name honoring 
Dr. Gressitt, now Entomologist and Head of the Entomology Depart- 


350 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLII 


ment of the Bernice P. Bishop Museum, Honolulu, whose labors in the 
two fields of insect taxonomy and zoogeography of the Pacific are of 
more than casual interest to the botanist. 

The new species may be recognized by its shrubby habit; solitary 
syncarp set on a long or slightly nodding peduncle furnished with bracts; 
and the unilocular drupes with conoid pilei about 2-3 mm. long produced 
into a short acerose stylose stigma slightly curved or hooked in form, with 
the stigmatic groove uppermost. Gressitt’s notes indicate that the type 
plant was about 1 m. tall, with a trunk 1.5 cm. in diameter; the fruits were 
green. The leaves, which are quite small and narrow for the genus, are 
evenly tapered to a sharp, elongated apex. 

COLLEGE oF GUAM 

AGANA, GUAM 


oe 
anaadeume 


a 


ERNEST JESSE PALMER 


JOURNAL 


OF THE 


ARNOLD ARBORETUM 


VoL. XLII OctTOBER 1962 NUMBER 4 


ERNEST JESSE PALMER, 1875-1962 
CLARENCE E. Kosusxk1! 


With portrait 


ERNEST JESSE PALMER died at Webb City, Missouri, on February 25, 
1962, in his 87th year. This marked the end of a long and in many 
ways remarkable career in the fields of botany and allied sciences. 

He was born in Leicester, England, April 8, 1875, and, in 1878, when 
he was three years old, he and his sister, Louise, were brought to this 
country by their parents, Amos and Annie Palmer, who settled temporarily 
near Warrensburg, in Johnson County, Missouri. When he was 14, his 
father, responding to the stories of quick wealth offered by the mining 
opportunities in the southwestern part of Missouri, moved the family 
to Webb City and built a home for them at 321 South Main Street. 
The wealth never materialized! Throughout the rest of his life, although 
he spent many of his years away from Webb City, the Palmer home 
was maintained, and after retirement, Ernest Palmer moved with his 
family and lived out the rest of his life there. 

In this mining area, young Palmer did find a wealth of his own liking. 
It was here that he began his studies of natural phenomena. In the 
piles of waste material from the mines, fossils of plants and animals 
caught his fancy, and as in all boys of his age the collecting instinct 
came to the fore. He developed a great keenness for unearthing most 
interesting artifacts. In later years at the Arnold Arboretum he was 
always “picking up” Indian arrowheads and other Indian deposits. 
This was easy for him, but not necessarily for others. I can recall his 
telling me of a spot in the Arboretum where arrowheads were in abundance. 
I hurried to the exact location but found nothing. The next day we went 
out together— and sure enough, he picked up several more artifacts. 
Eventually two attractive trays of Palmer’s finds in the Arboretum were 
mounted and have been on display there ever since. 

His formal education was broken off suddenly while he was attending 
high school. A physical collapse of his father made it necessary for him 
to leave school and seek employment. This was undoubtedly a severe 

This article on Mr. Palmer will be ene also in the current volume of 
Rhodora, but without the bibliography. C.E.K 


352 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


blow to his pride because he wanted an education above all things. He 
loved study and research — and he intended to continue it at all costs. 
Just recently [I learned from an article by Marcia W. Kershaw, a Special 
Correspondent of the St. Louis Post-Dispatch, that Palmer managed to 
study Latin and Greek at old Webb City Baptist College. I had often 
wondered where he obtained his profound knowledge of these languages, 
especially Latin which he used so ably in his original technical descrip- 
tions of new species of plants. Mrs. Kershaw’s article brought to light 
many other facets of Mr. Palmer’s career which were previously unknown 
to me and [ am very thankful to her for her assistance. 

All his life he had managed to study not only the natural sciences but 
also English literature, mathematics, economics, and poetry until he 
was considered very proficient in each of them. 

The portrait which accompanies this article was taken in 1958 and 
shows Mr. Palmer at his desk with a copy of his published book of poems 
entitled, “Gathered Leaves, Green, Gold and Sere.” These poems, I 
understand, are truly gathered leaves, ‘leaves’ which had been lying 
around in various places during the years and were gathered together 
by his wife and published through her efforts when Palmer was eighty- 
three years old. 

His scientific papers were noted not only for his Latin descriptions but 
also for the fine phrasing which he always employed. He wrote so well 
and so easily that often he found it necessary to delete whole paragraphs 
from his papers because he considered the contents “unscientific.” It 
was a pity because everyone would have enjoyed reading them. 

Although he was dedicated to the life of a naturalist it was not until 
his meeting with Benjamin Franklin Bush, also from Missouri, that he 
tried his wings and ventured away from his home setting. When Bush 
died in 1937, Palmer published a resumé of his life in The American 
Midland Naturalist. On re-reading this biography one cannot help but 
be struck by the similarity in the early backgrounds, the interests, and, 
in fact, the entire careers of the two men. 

In 1901, Bush, then a collector for the Arnold Arboretum, visited the 
Palmer home and spent a week there using it as a base for his botanical 
collections. Bush was especially interested in obtaining specimens of 
Crataegus for C. S. Sargent, then director of the Arnold Arboretum, and 
when he departed from Webb City he left with Palmer a supply of driers 
for further collections. Thus at the age of 25 Palmer began his lifelong 
interest in Crataegus and his association with Sargent and the Arnold 
Arboretum. For the next twelve years he collected extensively in his 
home area, concentrating on the flora of Missouri, and it was not until 
1913, when he was 38, that he became associated with the Missouri 
Botanical Garden. Two years later at the age of 40 he began his many 
collecting trips to the Southwest under the aegides of both the Missouri 
Botanical Garden and the Arnold Arboretum. 

Palmer began publishing as early as 1910, at which time his first 
paper, “Flora of the Grand Falls Chert Barrens” appeared in the Trans- 


1962 | ERNEST JESSE PALMER, 1875-1962 Siok) 


actions of the Academy of Science, St. Louis. During the next ten years 
he published seven papers, mostly on ferns. In 1920, while still a collector- 
at-large he sent in from Webb City to the newly organized Journal of the 
Arnold Arboretum his first of many papers to be published by this 
Harvard University publication. Two more papers were published in this 
journal before he came to Jamaica Plain in 1921 as a regular staff member. 
He retained this status at the Arnold Arboretum for twenty-seven years, 
until he retired in 1948. 

My personal association with Palmer began in 1927. By that time he 
was firmly established at the Arboretum as a collector-botanist. Since 
he was away so much of the time on his many collecting trips and since 
he was still a bachelor he maintained personal quarters in a rooming house. 

He was a very busy man during this period. He managed to make a 
collecting trip nearly every year, usually alone, occasionally with a single 
companion. Upon his return from these trips his time was spent identifying 
his specimens, making up sets for distribution, and writing occasional 
papers — and getting ready for his next trip. He once mentioned to me 
that he had hopes of collecting in all the states of the Union. At the 
time he had nearly accomplished his intention. If I recall correctly he 
said that there were only two states in which he had not made “official” 
collections — and, as one might suspect, one of them was surprisingly 
close by! 

The greater portion of his collections was made in the southwestern 
United States. However, he always maintained his interest in the Missouri 
flora, and, in 1935, there appeared in the Annals of the Missouri Botanical 
Garden the 385-page article by Palmer and Steyermark, “An Annotated 
Catalogue of the Flowering Plants of Missouri.” This catalogue is more 
complete than is usual in such a publication. The first seventy-five pages 
are devoted to the botanical history, terrain, climate, plant regions, etc., 
of the state. Under the actual listing of the taxa may be found the 
habitat and a record of the counties wherein specimens of each entry had 
been collected. A footnote states that an earlier publication on the ferns 
and fern allies of Missouri published by the same authors three years 
earlier in the American Fern Journal should be considered supplementary 
to the 1935 Catalogue. 

A recent letter to me from Julian Steyermark, the junior author of 
the above two publications, mentions that his latest publication, Flora 
of Missouri, which has been in press since 1959 is being dedicated to 
Bey), Palmer. Palmer had hoped to see the book and knew of the proposed 
dedication. 

In 1930, when he reached the age of 55 he surprised us all by getting 
married. We had thought of him as a confirmed bachelor. He married 
Elizabeth McDougal, a bacteriologist at the Massachusetts State Lab- 
oratory which is located alongside the Arnold Arboretum. Naturally, 
a great change took place in Palmer’s life. He eventually set up a fine 
home in one of the houses belonging to the Arnold Arboretuin in Jamaica 
Plain. He and his wife had three children: Ernest Macdougal Palmer, 


354 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XLII 


born in 1931; Grace Elizabeth Palmer, in 1932; and Theodore Windle 
Palmer, in 1935 

Palmer possessed a physique that defied age. He was the slender, wiry 
type and at the time of his retirement and departure from Jamaica Plain 
for Webb City could outlast any of us on a collecting trip. Although 
I did not see him again, I understand from his long letters and from 
various conversations with others that he continued his same brisk manner 
in everything which he undertook. His letters show that when he retired 
he merely transferred his activities from Massachusetts to Missouri. In 
a letter to me in February 1955 he says: “Of course we miss many 
friends there [Jamaica Plain], and I miss many of the associations at 
the Arboretum. But I have never been sorry that I retired when I did, 
for my life on the whole has been quite happy here. I have never been 
busier at any time or interested in more things. In April I will pass 
another anniversary — and you can figure it out as the records show that 
I was born in 1875. It is hard for me to believe it as I still feel as strong 
in body and mind — so far as I can tell —as I did at fifty. I can easily 
walk ten miles any day; and I think I could climb a good sized tree, if 
there was anything at the top of it that I wanted badly enough. . 
Steyermark and I are still working actively on the Flora of Missouri .. . 
Specimens of Crataegus, Quercus and other genera keep coming to me 
for determination or revision. . . Other collections in my museum also 
occupy much time. . . I have agreed to talk to a joint meeting of all 
the Garden Clubs of Carthage at the public library. While I do not 
solicit or particularly enjoy speaking engagements, I fill them occasion- 

t is so fine and warm today that I think I will go fossil hunting 
this afternoon.” For a man retired and eighty years old he certainly was 
enjoying life to the fullest, doing exactly what he wanted to do — and he 
seemed so well equipped to do it! 

During his life time Ernest Palmer joined many scientific organiza- 
tions, among them: the American Fern Society, the American Associa- 
tion for the Advancement of Science, the American Society of Plant 
Taxonomists, the Boston Mineral Club, the Botanical Society of America, 
the Missouri Archaeological Society, the New England Botanical Club 
(President, 1944-1945), and the St. Louis Academy of Science. 


BIBLIOGRAPHY * 

— 1910 — 

Flora of the Grand Falls chert barrens. Trans. Acad. St. Louis 19: 97-112. 
— 1914 — 

Ophioglossum Engelmanni in Missouri. Am. Fern Jour. 4: 66-68. 
— 1915 — 

Trailing the first settlers. Archeol. Bull. 6: 37-41. 

* Compiled by Lazella Schwarten and Elizabeth M. Palmer. 


1962] 


ERNEST JESSE PALMER, 1875-1962 


— 1916 — 
Catalogue of the plants of Jasper County, Missouri 
plants.) Ann. Missouri Bot. Gard. 3: 3 


(Fernworts and flowering 
Texas Pteridophyta—I. Am. Fern Jour. 9: 17-22 


— 1919 — 
Texas Pteridophyta — II. /bid. 50-56 
Texas Pteridophyta — III. /bzd. 81-85 


— 1920 — 
mes canyon flora of the Edwards Plateau of Texas. 
—239. 


Jour. Arnold Arb 
ea ligneous flora of the Staked Plains of Texas. /bid. 2: 90-105. 


ae 
— 1921 — 
Botanical reconnaissance of southern a 
The forest flora of the Ozark region. [bid. 21 


Jour. Arnold Arb. 2: 129-153. 
32, 

— 1923 — 

The flowers that bloom in the ing. Cornell Rural School Leafl 


The Red River Forest at Fulton, Arkansas Jour. Arnold Arb. 4: 8-3 
— 1924 — 
Arb. 5: 


16: 1-58. 
Two interesting ferns from Arkansas. 
108-134. 


Am. Fern Jour. 14: 39-4 
The ligneous flora of Rich Mountain, Arkansas and eee “Jour. Arnold 
— 1925 — 
Is Quercus arkansana a hybrid? Jour. Arnold Arb. 6: 195-200 
Synopsis of North American Crataegi. Ibid. 5-12 
— 1926 — 


Leaves from a collector’s note book. Jour. Arnold Arb. 7: 136-145. 


The ligneous flora of Hot Springs National Park and vicinity. Ibid. 
Supplement to the catalogue of Arkansas Plants. 
Trans. Acad. St. Louis 25: 91 


104-135. 
(With Buchholtz, J. T.) 
— 1927 — 
Ferns of the Davis Mountains, western Texas 
On Nuttall’s trail through Arkansas 


Am. Fern Jour. 17 
Jour. Arnold Arb. 8: 24-55 
— 1928 — 
A botanical trip through the Chisos Mountains of Texas. Jour. Arnold Arb 
9: 153-173. 
Leaves from a collector’s note book. Jbid. 173-187 
— 1929 — 

The ligneous flora of the Davis Mountains, Texas. Jour. Arnold Arb. 10: 8 45 

— 1930 — 
Th 


Additional notes on Texas ferns. Am. Fern Jour. 20: 


e spontaneous flora of the Arnold Arboretum. a fe Arb. 11: 63-119 


356 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLIII 


— 1931 — 
A conspectus of the genus Amorpha. Jour. Arnold Arb. 12: 157-197. illust. 
— 1932 — 
Crataegus arnoldiana. Addisonia 17: 
The Crataegus problem. Jour. oe ie: om 342-362. 
The ferns and fern allies of Missouri. (With Steyermark, J. A.) Am. Fern 
Jour. 22: 105-122, 
Leaves from a collector’s note book. Jour. Arnold Arb. 13: 417-437. 
Notes on Ophioglossum Engelmanni. Am. Fern Jour. 22: 43-47. 


— 1933 — 
The family Isoétaceae in Missouri. (With Steyermark, J. A.) Am. Fern Jour. 
23: 65, 66 


— 1934 — 
Adventures in fern collecting. Am. Fern Jour. 24: 104-109. 
The beach plum in Michigan. Jour. Arnold Arb. 15: 88. 
Indian relics of the Arnold Arboretum. Bull. Pop. Inf. IV. 2: 61-68. illust. 
Notes on some plants of Oklahoma. i Arnold Arb. 15: 127-134 
Quercus ellipsoidalis in Missouri. Ibid. 
Trees of the Southeastern States. ae Ibid. 266. 


— 1935 — 
Additional fern records for Missouri. (With Steyermark, J. A.) Am. Fern 
Jour. 25: 1-6. 


An annotated catalogue of the flowering Nata of Missouri. (With Steyermark, 
Ann. Missouri Bot. Gard. 22: -758 

uae ae i the spontaneous flora of . Arnold Arboretum. Jour. Arnold 
Arb. 16: 81-97. 

Two new species of Crataegus from Missouri. Ibid, 353-357. 

Uvidaria perfoliata in Louisiana. (With Anderson, E.) Rhodora 37: 58, 59. 

— 1936 — 

Adventures in fern collecting, I]. Am. Fern Jour. 26: 136-139. 

Brauneria atrorubens and B. paradoxa. Rhodora 38: 197-199. 

Deciduous shrubs and trees; Evergreen shrubs and conifers: Broad leaved 
evergreens. Landscape Gardening and Landscape Art, Garden Clubs of Ark. 

. 22-39, 

The hawthorns (Crataegus: over 50 species in Arkansas). Native plant material 
(famous in garden lore). Arkansas state program for garden clubs. (Mimeo- 
graphed.) 

— 1937 — 

Benjamin Franklin Bush. Am. Midl. Nat. 18(3): i-vi. portra 

The identification of plant material at the Arnold oe. Bull. Pop. Inf. 
IV. 5: 13-16. 

Notes on North American trees and shrubs. Jour. Arnold Arb. 18: 133-140. 

— 1938 — 

Additions, corrections, and revisions of the annotated catalogue of the flowering 
plants of Missouri. (With Steyermark, J. A.) Ann. Missouri Bot. Gard. 25: 
775-794 


1962 | ERNEST JESSE PALMER, 1875-1962 Sou. 
New varieties and forms from Missouri. (With Steyermark, J. A.) /bid. 
769-773 
— 1939 — 


Mines and minerals of the Tri-State District. Rocks & Minerals 14: 35-49, 
Some woody plants of Rhode Island. Rhodora 41: 314-316 


— 1940 — 
Adventures in fern collecting, III. Am. Fern Jour. 30: 1-9. 
Crataegus of Indiana. In: C. C. Deam, Flora of Indiana. 533-557. 


Geodes. Rocks and Minerals 15: 120-123. 
A new form of Crataegus viridis Linn. (With Pickens, A. L.) Neighborhood 
Research 4: 61. 
— 1941 — 
Answering isolationists. Changing World 13: 10-12. 
— 1942 — 


Panicum recognitum in Rhode Island. Rhodora 44: 227. 
The red oak complex in the United States. Am. Midl. Nat. 27: 732-740. 


— 1943 — 
Quercus prinus Linnaeus. Am. Mid]. Nat. 29: 783-784. 
The species concept in Crataegus. Chron. Bot. 7: 373- Hise 


Hed (eye 
Food plants in the Arnold Arboretum. Arnoldia 4: 1-7. bs. Ao? 


— 1945 — 
Quercis Durandii and its allies. Am. Midl. Nat. 33: 514-519. 


— 1946 — 
Crataegus in the northeastern and central United States and adjacent Canada. 
Brittonia 5: 471 


eso) ee 

Ferns and fern allies of Cumberland, Rhode Island. Am. Fern Jour. 37: 33-38. 

Second supplement to the s eo ieee flora of the Arnold Arboretum. Jour. 
Arnold Arb. 28: 410-418 


— 1948 — 
Hybrid oaks of North America. Jour. Arnold Arb. 29: 1-48. 


— 1950 — 
Crataegus. In: M. L. Fernald. Gray’s Manual of Botany (8th ed.) pp. 767-801. 
Notes on eae EDIT Mackenzie. (With Steyermark, J. A.) Bull. 
Torrey Club 77: Pai 


— 1951 — 


Trees and plants of the Southwest Missouri Region. 21 articles appearing 
weekly in the Webb City Daily Sentinel Oct. 1951-Mar. 1952. 


358 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


— 1952 — 

Crataegus. In: Henry A. Gleason, The new Britton and Brown illustrated 
flora of the northeastern United States and adjacent Canada 2: 338-375 
New pteridophyte records from Missouri. (With Steyermark, J. A.) Am. Fern 

Jour. 42: 61-66. 


— 1953 — 
A hybrid Amorpha and new forms and records from Missouri. Rhodora 55: 
157-160 
— 1955 — 
Plants new to Missouri. (With Steyermark, J. A.) Rhodora 57: 310-319. 


— 1956 — 
Crataegus in Ohio with description of one new species. Ohio Jour. Sci. 56: 
205-216. 


— 1958 — 
Cephalopods aes the Burgner Formation in oe (With Unklesbay, 
A. G.) Univ. Missouri Jour. Paleont. 32: 1070-10 
Plants new to “Missouri. (With Steyermark, J. A.) Balt 10: 109-120. 


— 1962 — 
The Ozark Forest. Missouri Log (pub. annually by School of Forestry, Univ. 
Missouri). 


1962] BRIZICKY, GENERA OF ANACARDIACEAE 399 


THE GENERA OF ANACARDIACEAE IN THE 
SOUTHEASTERN UNITED STATES * 


GEORGE K. BrizIckKy 


ANACARDIACEAE Lindley, Intr. Nat. Syst. 127. 1830. nom. cons. 
CASHEW FAMILY) 


Trees or shrubs, rarely subshrubs or vines, with resin ducts in bark and 
often in leaves, flowers, and fruits. Leaves alternate {very rarely oppo- 
site or verticillate], simple, 3-foliolate, or pinnate, deciduous or persistent, 
exstipulate. Flowers small, usually regular, hvpogynous [rarely perigy- 
nous or epigynous], usually unisexual by abortion or both uni- and bi- 
sexual [rarely bisexual throughout], in axillary and/or terminal thyrses 
or panicles, more rarely in solitary or panicled spikelike racemes; bracts 
deciduous or persistent, bractlets 2, deciduous or persistent, or wanting: 
Perianth double [rarely simple or absent], usually 5-merous. Sepals 
usually connate at least at base, imbricate [or valvate]. Petals usually 
distinct, imbricate [or valvate]. Stamens 5-10 [or very rarely more], 
sometimes only 1 or 2 fertile in ¢ flowers, distinct [or basally connate], 
reduced and sterile [or wanting] in @ flowers: anthers versatile, 2-locular 
at anthesis, introrse, longitudinally dehiscent. Intrastaminal (or rarely 
extrastaminal) nectariferous disc usually 5- or 10-lobed [sometimes pro- 
duced into a gynophore, rarely absent]. Gynoecium syncarpous [or apo- 
carpous], 3[-5]-carpellate or 1-carpellate by reduction, rudimentary 
[or absent] in 4 flowers; stigmas 1-3[—5]; styles 1-3[-5]; ovary 1[3— 
5]-locular; ovules anatropous, apotropous, solitary in a locule, the funicle 
usually elongate with a basal, parietal, or apical insertion. Fruit usually 
drupaceous, with + resinous and sometimes waxy or oily mesocarp and 
crustaceous or bony endocarp (stone). Seeds with scanty endosperm or 
endospermless; embryo + curved [or straight]. (Terebinthaceae Juss., 
1789: Spondiaceae Kunth, 1824.) Type GeNus: Anacardium 


1 Prepared for a generic flora of the southeastern United States, a joint project of 
under the direction of Reed C. Rollins and Carroll E. Wood, Jr. This treatment are 
the pattern established in the first paper in the series (Jour. Arnold Arb. 39: 296- 
346. 1958) and continued through those in volumes 40-43 (1959— 1962). It should be 
plants of ae eee with any supplementary material in brackets. References which 

Since is aie to Dr. Carroll E. Wood is for his criticism and valuable 


en and to Mrs. Gordon W. Dillon, for her careful help in the preparation of 
the manuscript. 


360 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLUI 


A family of about 70 genera with nearly 600 species, primarily pan- 
tropical, extending with several genera into the temperate zones of both 
hemispheres. Of the five tribes, only Rhoéae (Rhoideae) and Anacardieae 
(Mangifereae Marchand emend. Engler) a1 are represented in our area, the 
former by three indigenous genera and a naturalized species of chile 
the latter by a species of Mangifera naturalized in southernmost Flori 
Since generic lines in the oéae are often weak and differently ane 
preted, various authors have recognized one to five indigenous genera in 
our area. 

The family, with a few exceptions (e.g., Pistacia L.), is considered to be 
entomophilous. Since the species are mostly dioecious or polygamo-dioeci- 
ous, cross-pollination should be the rule. Apomixis (adventitious poly- 
embryony) has been found in some species of Mangifera, parthenocarpy 
in several genera. Cytological data presently known for about 25 species 
in ten genera (2m = 24, 28, 30, 32, 40, 42, 48, 60) suggest polyploidy. 
Resin of many species is poisonous on contact. 

The family is believed to be closely allied with Sapindaceae and Juliana- 
ceae. Wood-anatomical and palynological evidence seems to support the 
assumption of a close relationship with Burseraceae, from which Anacar- 
diaceae differ essentially in the uniovulate ovary locules and in apotropy of 
he ovules. 

Anacardiaceae are economically important for edible fruits (e.g., Mangi- 
fera indica, mango: Spondias spp., mombin) or nuts (Anacardium occi- 
dentale L., cashew; Pistacia vera L., pistachio), for resins (e.g., Pistacia 
Lentiscus L., mastic) and lacquers (Rhus verniciflua), for tannins (Cotinus 
Coggvgria, Rhus spp., Schinopsis spp.), and for timbers (Schinopsis spp., 
red quebracho; Astronium spp., zebrawood or kingwood). Some species are 
ornamentals. 

REFERENCES: 
Bark Ley, I’. A. A monographic study of Rhus a its ee allies in North 
and bernie America, including the West In . Missouri Bot. Gard. 
24: 265-498. pls. 10-26. 1937. | Includes Eo ieee Rhus, Toxi- 
codendron, Malosma, and Actinocheita. 
. A key to the genera of the Anacardiaceae, Am. Midl. Nat. 28: 465- 
474. 1942. [Bibliographical footnotes; Searsia Barkley and Duckera Bark- 
ley segregated from Rhus. | 
Generic key to the sumac family (Anacardiaceae). Lloydia 20: 255- 
265. 1957. [Includes introductory notes on economic uses. | 
BLANK, H. Dermatitis from Anacardiaceae. Proc. Fla. Mango Forum 17: 24-26. 
oro Beas 
Encier, A. Uber die morphologischen Verhaltnisse und die geographische Ver- 
breitung der Gattung Rhus, wie der mit ihr verwandten, lebenden und aus- 
gestrobenen Anacardiaceae, Bot. Jahrb. 1: 365-420. 
. Burseraceae Anacardiaceae. Ja: CANDOLLE, A. & C. pe. Monogr. 
Phaner. 4: 1-500, 536-573. pls. 1-15. 1883. | Anacardiaceae, 171-500, 537, 


wal 
wa 
CO 
>> 
=~ 
a 


2] 
Anacaedincéas Nat. Pflanzenfam. III. 5: 138-178. 1892. 
Heimscu, C., Jr. Wood anatomy and pollen morphology of Rus and allied 


— 


1962 | BRIZICKY, GENERA OF ANACARDIACEAE 361 


genera. Jour. Arnold Arb. 21; 279-291. pls. 1-3. 1940. {Includes Actino- 
cheita, Malosma, Metopium, Cotinus, Rhus, and Toxicodendron 
mparative anatomy of the Seconda, xylem in the “Gruinales” and 

“Terebinthales,” of Wettstein with reference to taxonomic grouping. Lilloa 
8: 83-198. pls. 1-17. 1942. [Anacardiaceae, 136-144, pls. 9-12.| 

Kryn, J. M. The anatomy of the wood of the Anacardiaceae and its bearing on 
the Beas and relationships of the family. Diss. Abs. 13: 290. 1953.* 

Martin, A. C. The comparative internal morphology of seeds. Am. Midl. Nat. 
36: 513-660. 1946. [Anacardiaceae 628, 629 

Merritt, E. D. Dermatitis caused by various representatives of the Ana- 
cardiaceae. Jour. Am. Med. Assoc. 124: 222-224. 

Recorp, S. J. American moods of the family Annee Trop. Woods 60: 
11-45. 1939. 

SARGENT, C. S. Manual of the trees of North America (exclusive of Mexico). 
ed. 2. 910 pp. Boston & New York. 1922. [Anacardiaceae, 655-665. | 

Sweet, H. R., & F. A. Barkrey. A most useful plant family, the Anacardiaceae. 
Missouri Bot. Gard. Bull. 24: 216-229. 1936 

VENNING, F. D. The ontogeny of the laticiferous canals in the Anacardiaceae 
Am. Jour. Bot. 35: 637-644. 1948. 

West, E., & L. E. Arnotp. The native trees of Florida. 212 pp. Gainesville. 
1946. [ Anacardiaceae, 112-114. ] 


KEY TO THE GENERA OF ANACARDIACEAE 


General characters: woody plants with resin ducts in bark, leaves, flowers, and 
fruit; flowers small, usually hypogynous, unisexual or uni- and bisexual, in 
axillary and/or terminal thyrses or panicles; perianth double, usually 5-merous ; 
stamens 5-10; nectariferous disc intra- or rarely extrastaminal; gynoecium 1-3 
| -5 |-carpellate, ses usually 1-locular by abortion; fruit drupaceous 
Flowers é and bisexual; petals with longitudinal ridges; disc oe eae 
stamens 5, only 1 or 2 fertile; gynoecium 1-carpellate; drupes large to ve 
large, with copious juicy flesh; leaves simple; naturalized tree . a cacal 
FSO TULA aye tose cts, Cn ses eke ole te ee ce 1d cola 1. Manigifera. 
. Flowers usually unisexual; petals not ridged; disc ee stamens 

5 or 10, all fertile in ¢, all reduced and sterile in 2 flowers; gynoecium 3- 

carpellate; drupes cum not exceeding 15 mm. in length, with meager dry- 

He flesh; leaves variou 

. Stamens 10; ae spicy-aromatic, containing oil cavities adherent to 
stone ; eaves pinnate, with winges: rachises; naturalized ee or small 
treessor southern «blOmGa. 2 4a ote ee eee en tts: . Schinus. 

B. Stamens 5; drupes not aromatic, without oil cavities; leaves ae if 

pinnate, rachises not winged, except in Rhus copallina. 

C. Stigma 1, 3-lobed; style 1, very short, stout; drupes 10-15 mm. long, 
ellipsoid to obovoid, scarlet to orange, glabrous; endocarp thin, 
crustaceous; leaves odd-pinnate, leaflets long petioluled; A a 
trees or shrubs of subtropical: Floridasgnna ss .25 2.50. . Bh topium. 

3-8 mm. long, subglobular or oe 
form (covered with glandular club-shaped hairs if red or orange); 
endocarp thick, bony; leaves various (if pinnate, leaflets sessile or 
short petioluled). 

D. Styles appearing sublateral in flowers; drupes obliquely subreni- 


> 


> 


& 
WM 
Hag 

q 
soo) 
wm 
pe?) 
jo] 
jor) 
wm 
os 
al 
SB 
oO 
wm 
ww 
fol} 
en 
o 


362 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


form, usually much compressed laterally, prominently reticulate, 
elabrous, the remnants of styles lateral; mesocarp very scanty; 
pedicels of aborted flowers becoming plumose-villous; seal 
fee innocuous trees or shrubs. ................ 4. Cotinus, 
. Styles terminal; drupes subglobular, usually slightly pains 
laterally, not reticulate, sometimes with simple and/or glandular 
hairs; mesocarp thickish, resinous and sometimes also waxy; 
pedicels never plumose; leaves 3-foliolate or odd-pinnate; trees, 
shrubs, or vines, some poisonous. .................... 5. Rhus. 


=) 


Tribe ANACARDIEAE 
1. Mangifera Linnaeus, Sp. Pl. 1: 200. 1753; Gen. Pl. ed. 5. 93. 1754. 


Mostly large evergreen trees, the leaves alternate [rarely verticillate | , 
simple, entire, membranaceous to coriaceous, petiolate, persistent. Plants 
polygamous, usually andromonoecious. Flowers small, ¢ and bisexual, in 
terminal [and/or axillary] thyrses; pedicels articulated; bracts and bract- 
lets deciduous. Sepals 5 (4 or 6), deciduous. Petals 5 (4 or 6), distinct [or 
adnate to the disc at base], the veins thickened into [1—|]3—5 prominent 
ridges, deciduous. Disc extrastaminal, tumid [or minute, sometimes stipe- 
like, or wanting], S-lobed. Stamens usually 5 [rarely 10-12], inserted 
just within [or on] the disc, distinct [or connate at base], usually 1 or 2 
[rarely 5 or 6] fertile, the rest remaining sterile with imperfect small an- 
thers [or reduced to toothlike projections]; pollen 3-colpate, small to me- 
dium sized, ellipsoidal, finely reticulate. Gynoecium 1-carpellate (excep- 
tionally 2-carpellate, apo- or syncarpous), rudimentary or wanting in ¢ 
flowers; stigma simple; style sublateral; ovary obliquely subglobular, 1- 
locular, sessile; ovule solitary, funicle subbasal. Fruit a large [or relatively 
small] drupe of various shapes and colors; exocarp + leathery; mesocarp 
thick, fleshy, sweet [to acid], sometimes resinous; stone compressed, usual- 
ly hard, woody, coated with fibers. Seed large, compressed; testa papery; 
endosperm lacking (or scanty?); embryo subreniform with plano-convex, 
often unequal and lobed cotyledons and small ascendent radicle; germina- 
tion hypogeous. Type species: M. indica L. (From Portuguese, mango, 
the common name for this fruit, and Latin -fera, bearing, i.e., “bearing 
mangoes.”’) 


An Indo-Malayan genus of about 40 species. Mangifera indica (includ- 
ing M. laurina Bl.), common mango, 2n = 40, presumably native to India 
(and perhaps to Indochina and the Sunda Islands), widely cultivated 
throughout the tropics for its edible fruits, has become naturalized in ham- 
mocks of southern Florida and the Florida Keys. A tree with relatively 
large, narrow leaves and large to very large, ovate to subreniform or sub- 
cylindrical, yellowish or greenish fruits sometimes flushed with scarlet or 
crimson, the species has been in cultivation in the tropics of the Old World 
for over four thousand years and shows a wide range of variation, about a 
thousand wild forms and cultivars occurring in India. 


1962 | BRIZICKY, GENERA OF ANACARDIACEAE 363 


Mangifera indica is the only economically important species. Its fruits, 
mangoes, are said to be eaten by at least one-fifth of the world’s popula- 
tion. In India the astringent flowers, seeds, and bark find medicinal appli- 
cation, especially in diarrhoea. Bark and leaves yield a yellow dye used 
locally in southern Asia. The wood, said to be of excellent quality, is used 
extensively in India for carpentry, cabinet-work, boat-building, e 

Short-tongued insects, especially Diptera, Hymenoptera, and Coleoptera, 
seem to be the principal pollinators of Mangifera indica. Cross-pollination 
seems to be the rule. Since self-pollination has not been recorded, one may 
assume that the morphologically bisexual flowers are (at least in most 
cases) functionally (physiologically) carpellate. The somatic chromosome 
number 2” = 40 has been recorded for 23 grafted varieties and one wild 
race of Mangifera indica, as well as for M. sylvatica Roxb., M. caloneura 
Kurz, M. caesia Jack, and M. foetida Lour. Although no polyploid series 
has been found in the genus, Mukherjee (1950) suggested on the basis of 
his cytological analysis that M. indica and its allies are allopolyploids. No 
records of natural or artificial interspecific hybrids have been available, 
but Mukherjee (1957) remarked that “similarity in the chromosome num- 
ber and morphology and in the pollen size and morphology . . . indicates 
that there may be close compatibility among the species during hybridiza- 
tion and in stock-scion relationship in grafting . . .’ 

Apomixis (adventitious polyembryony) has been recorded in Mangifera 
indica and in M. odorata Griff., the embryos originating either from the 
nucellus or by budding from the cotyledons and hypocotyl. While Indian 
“seedling races” and “horticultural [grafted] varieties” are almost exclu- 
sively monoembryonic, the ‘‘seedling races” common in culture in some 
other countries (‘‘Philippine” or ‘‘Manila” races) mostly are polyembry- 
onic. There are, however, records indicating that some “Indian races” 
considered monoembryonic in India yielded a considerable percentage of 
polyembryonic seeds when grown in the Philippines. 

Although Mangifera indica is usually innocuous and dermatitis caused by 
its resin is infrequent, several wild species (e.g., M. caesia, M. foetida, M. 
odorata, and M. lagenifera Griff.) sometimes cultivated in the Malaysian 
region for their edible fruits are regarded as positively poisonous. 

The genus seems to be related to the Malayan Bouea Meisn. and to the 
tropical American Anacardium L. 


REFERENCES: 


The vast number of references has been reduced here primarily to those of 
general interest. Under family references see ENGLER (1883, pp. 195-215 & 
1892, pp. 146, 147), Hermscu (1942, pp. 136, 137), and MeErRILt. 

ArNpT, C. H. Notes on polyembryony and multiple aie from the seed in 
Mangifera indica. Am. Jour. Bot. 22: 26-28. pls. 1, 935. 

BaRNES, H. V. The mango; a list of references. U. S. ae Agr. Libr. List 29. 
62 pp. 1946.* 

CHoupHury, K. R., & S. K. MuKkuHerjee. Floral biology in mango. Proc. 
Indian Sal Congr: Assoc. 42: 370, 371. 1955.* 


364 JOURNAL OF THE ARNOLD ARBORETUM yOu, xn 


HAMMERSTEIN, C. P. Mango dermatitis experiences. Proc. Fla. Mango Forum 
19: 14-16. 1959.* [Control of allergy to mangoes. 

JULIANO, J B. Origin of embryos in the strawberry mango. Philip. Jour. Sci. 
54: 553-559. 1934. [See also Philip. Agr. 25: 749-760. 1937.] 

. E. Cuevas. Floral morphology of the mango ee indica 
Li) with eer eos to the Pico variety from the Philippines. Philip. 
Agr, 21: 449-472 * 

KENNARD, W. C. Bite of the fruit, seed and embryo of the Paneri 
mango. Bot. Gaz. 117: 28-32. 1955 

Leroy, J. F. Sur un ‘complexe aeamicue? des manguiers et sur l’origine et 
la phylogénie des variétés cultivées. Revue Int. Bot. Appl. Agr. Trop. 
27: 304-309. 1947. 

Lurz, B. Estudos sobre a biologia floral de pee es ba es ees sum- 
mary.) Mus. Nac. Rio de Janeiro 26: 125- pls. 1-4. 

Lyncu, S. J.. & R. O. Netson. Current methods of ees See of 
avocado, mango, lychee and guava in Florida. Ceiba 4: 315-337. 1956. 
| Mangifera indica, 320-328. | 

Matix, P. C. Morphology and biology of the mango flower. Indian Jour. 
Hort. 14: 1-23. 1957.* 

MUKHERJEE, S. Kk. anaes oe of the species of Mangifera Linn. 
Bull. Bot. Soc. Bengal 2: 15 

The varieties of mango ca indica L.) and their classification. 
ibid. 2: 101-133. 1948. 
. The iaononite value of the anatomical Sane of inflorescence 
axes of citi tale L. Jour. Indian Bot. Soc. 28: 162-171. hi 
graph on the genus Mangifera is ant 12: 73-136. 1949. 
: unas investigation of the mango (Mangifera indi L. ) a the 
allied eu species. Proc. Natl. Inst. Sci. India 16: 287-30 
The mango: its allopolyploid nature. Nature 166: 196, 197, 1950. 
; Pollen analysis in Mangifera in relation to fruit-set and taxonomy. 
Jour. Indian Bot. Soc. 30: 49-55. 1951 
. Origin, distribution and phylogenetic Sos of the species of Mangi- 
fera L. Tou Linn. Soc. Bot. 55 =83: 

The mango: its botany, a uses and future improvement, 

panne as observed in India. Econ. Bot. 7: 130-162. . 

i, 2G y of some Malayan species of Mangifera. Cytologia 22: 239- 

241. 1957. “TM. foetida and M. caesia, 2n = 40. |] 

Mustarp, M. J., & S. J. Lyncu. Flower-bud formation and development in 
Manegifera indica. Bot. Gaz. 108: 136-140. 1946. 

Rao, P. L. N. The essential oil of the mango flower. Sci. Cult. 11: 70. 1946.* 

Rao, U.N., & S. seers AMY. A note on ses formation from the axillary 
buds of the mango. S. Indian Hort. 3: 30, 955. 

Ramee. Po. he FOR, - W. C. Cooper. ae studies of floral induction 
in the Haden mango Series indica L.), Am. Jour. Bot. 36: 734-740. 
1949. [See also ibid. 33: , 210. 1946.] 

RvuEHLE, G. D., & R. B. .. Man growing in Florida. Fla. Univ. Agr. Ext. 
Bull. 174. 88 a map. 1960. 

SacHar, R. C., & R. N. CHopra. A study of ae — and embryo in 
Mangifera L. Sone Jour: Agr. Scl..272-21 

SHARMA, M. R. Studies in the family etme eres ¥ weal anatomy of 
the flower of Mazgifera indica L. Phytomorphology 4: 201-208. 1954. 


~ 


1962 | BRIZICKY, GENERA OF ANACARDIACEAE 365 


SincH, L. B. The mango: botany, cultivation, and utilization. 438 pp. London 

& New York. 1960.* [ Bibliography, 395-420. ] 

& R. N. Sincu. A monograph on the mangoes of Uttar Pradesh. 16 

1. 144 pp., 80 pls.; vol. 2. 138 pp., 75 pls. Lucknow. 1956. [Descriptions 
of vars. accompanied by colored pls. | 

Sincu, R. N. Sex ratio and fruit setting in mango (Mangifera indica L.). 
Science 119: 389, 390. 1954. 

SPENCER, J. L., & W. C. Kennarp. Limited stigmatic receptivity may con- 
tribute to low fruit set in the mango (Mangifera indica L.). Proc. Am. Soc. 
Hort. Sci. 67: 287-289. 1956. 

VENKATARATNAM, L. Hormone induced set and Perey in mango 
(Mangifera indica L.). Curr. Sci. Bangalore 18: 409. 1949.* 

Wesser, H. J. The economic importance of apogamy in Citrus and Manegifera. 
Proc. Am. Soc. Hort. Sci. 28: 57-61. 1931. 

Younc, T. W. Investigations of the unfruitfulness of the Haden mango in 
Florida. Proc. Fla. State Hort. Soc. 55: 106-110. 1943.* 

Influence of temperature on growth of mango pollen. /bid. 68: 308- 

SiSe 195525 


Tribe RHOEAE Marchand, ‘“Rhoideae” 

2. Schinus Linnaeus, Sp. Pl. 1: 388. 1753; Gen. Pl. ed. 5. 184. 1754. 

Trees or shrubs [rarely subshrubs, sometimes thorny]. Leaves odd-pin- 
nate or rarely even-pinnate [or simple]; the leaflets membranaceous to 
subcoriaceous [or coriaceous |, (3)5-13[-41], usually opposite [or al- 
ternate], relatively small to medium sized, entire or toothed, sessile or 
subsessile, the rachis often winged. Plants usually dioecious. Flowers 
usually unisexual, small, pediceled, in axillary [and terminal] thyrses |or 
in contracted raceme-like inflorescences]; bracts small, bractlets 2, minute, 
deciduous. Sepals 5 [4], persistent. Petals 5 [4], white [or yellow], con- 
siderably longer than the sepals. Stamens 10 [8], in 2 series, the anti- 
petalous shorter than the antisepalous, inserted below and between the 
lobes of a saucer-shaped 10[8]-lobed intrastaminal nectariferous disc, re- 
duced and sterile in 2 flowers; pollen 3-colpate, medium sized, ellipsoidal, 
finely reticulate. Gynoecium 3-carpellate, rudimentary in 4 flowers; 
stigmas 3 [1], capitellate; styles 3 [1], connate at least at base; ovary 
sessile, 1-locular by abortion; ovule solitary, 2-integumented, with a thick 
nucellus, suspended from near the top of the locule. Drupe small, pea-like, 
bright red [pink or lavender]; exocarp thin, chartaceous, shining; meso- 
carp resinous, the innermost layers (adherent to endocarp) with large cavi- 
ties containing an aromatic oil; stone laterally compressed, cartilaginous 
to bony. Seed sublenticular; testa thin, membranaceous; endosperm fleshy, 
scanty; cotyledons flat; radicle elongated, incurved upwards. (Including 
Duvaua Kunth). Lecrotypr species: S. Molle L., 2n = 28, 30; see A. S. 
Hitchcock, im Int. Bot. Congr. 1930. Nomencl. Propos. Brit. Bot. 153. 
1929. (An ancient Greek name for the mastic tree, Pistacia Lentiscus L., 
applied by Linnaeus to this genus.) — PEPPER-TREE. 


A genus of about 28 species of warm-temperate and tropical South 


366 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


America, from Chile, Argentina, and Uruguay, north to Venezuela and 
Colombia. Schinus terebinthifolius Raddi, Brazilian pepper-tree or Christ- 
mas-berry, with 5-13 oblong-ovate leaflets, and S. Molle, Peruvian pepper- 
tree, with 15-41 very narrow leaflets, are frequently cultivated in the 
warmer parts of the Eastern and Western hemispheres as ornamentals. 
Both are grown in Florida, and the former, a shrub, has become naturalized 
in southern Florida and the Keys where it seems to be spreading rapidly; 
the exact extent of naturalization needs to be determined. Schinus Molle 
has become naturalized in California and Mexico. The infructescences are 
often used as Christmas decorations. In Mexico and South America the 
fruits are used in various beverages, the seeds are sometimes used to 
adulterate pepper, and the resin, bark, and fruits are used locally in 
medicine. 

The genus seems to be most closely related to the South American 
Lithraea Miers and the East Australian Rhodosphaera Engl. The group is 
entomophilous, and honeybees have been reported as frequent visitors of 
flowers of Schinus Molle. No hybrids have been recorded. 


REFERENCES: 
Under family references see ENGLER (1883, pp. 331-345 & 1892, pp. 162-164), 
HeErIMScH (1942, pp. 139-141), and Recorp (pp. 38-40). 
BARKLEY, F. A. Schinus L. Brittonia 5: 160-198. 1944. [Monograph. | 
. A study of Schinus L. Lilloa 28: 5-110. 1957. [ Monograph. | 
CopeLAND, H. F. The reproductive structures of Schinus Molle (Anacardiaceae). 
Madrono 15: 14-25. 1959. [Floral morphology and embryology; 2n = 


30.] 

Dickson, J. D., HII, R. O. Woopsury, & T. R. ALEXANDER. Check list of flora 
of Big Pine Key, Florida and surrounding keys. Quart. Jour. Fla. Acad. 
Sci. 16: 181-197, 1953. [S. terebinthifolius, an escape on Big Pine Key, 


92.] 

RAINERI, L. Lo sviluppo dei fiori femminili di Schinus Molle L. (English sum- 
mary.) Nuovo Gior. Bot. Ital. II. 59: 49-63. 1952. [Development of the @ 
flowers. | 

——. Lo sviluppo dei fiori maschili di Schinus Molle L. (English summary.) 
Ibid, 61: 409-413. 1954. [Development of the 4 flowers. | 


3. Cotinus Miller, Gard. Dict. Abr. ed. 4. 1754. 


Small trees or shrubs with orange-yellow wood and _ strong-smelling, 
resinous sap. Leaves simple, entire, medium sized [or small], membrana- 
ceous, deciduous, the petioles short to long. Plants usually dioecious 
[occasionally polygamo-dioecious or -monoecious|. Flowers small, usually 
unisexual [rarely also bisexual], many abortive, in loose terminal thyrses, 
the slender pedicels much elongated after flowering, those of abortive 
flowers becoming plumose-villous; bracts narrow, + scarious, in part 
persistent. Sepals 5, persistent. Petals 5, yellowish- or greenish-white 
[rarely crimson], twice as long as the sepals. Stamens 5, distinct, inserted 
below an annular, shallowly 5-lobed intrastaminal disc, alternate with 
and shorter than the petals, reduced and sterile in @ flowers; filaments 


1962 | BRIZICKY, GENERA OF ANACARDIACEAE 367 


subulate; anthers broadly ovate in outline, somewhat shorter than the 
filaments, 2-locular at anthesis; pollen 3-colpate, medium sized, sub- 
spherical, striate-reticulate, with round to slightly oblong, smooth germ 
pores. Gynoecium 3- carpellate, rudimentary in ¢ flowers; stigmas 3, 
small, subcapitate; styles 3, appearing sublateral, unequal, that of the 
fertile carpel conspicuously longer than the 2 sterile ones; ovary 1-locular 
by abortion, obliquely obovoid, sessile; ovule raised on a basal funicle. 
Drupes small (4-5 mm. long), obliquely obovate to subreniform, usually 
much compressed, conspicuously reticulate veined, the style-remnants 
lateral; exocarp membranaceous; mesocarp very scanty; stone (endocarp) 
bony, subreniform, 1-seeded. Seed reniform; testa thin, membranaceous; 
endosperm wanting (?); embryo with flat, elliptic cotyledons and an 
elongate radicle incurved upwards (toward the hilum). Typr sPrciEs: 
Rhus Cotinus L. = Cotinus Coggygria Scop. (Derived from Greek, 
kotinos, an ancient name for wild olive, used by Pliny for an unidentified 
shrub of the Apennines, but applied by some pre-Linnaean botanists, e.g., 
Tournefort, to C. Coggygria.) — SMOKE-TREE. 


A genus of three or more species, primarily of warm-temperate Eurasia 
(southern France to the eastern subtropical Himalaya and central China) 
and eastern North America. Cotinus obovatus Raf. (C. americanus Nutt., 
Rhus americanus (Nutt.) Sudworth), American smoke-tree,” a tree, some- 
times a low shrub, with obovate-cuneate leaves turning orange to crimson 
in autumn, occurs sparingly on wooded, rocky cliffs and river bluffs, 
usually on limestone, in disjunct populations in the mountains of northern 
Alabama and adjacent Tennessee (Franklin County); in Daviess County, 
Kentucky (introduced?); in southwestern Missouri, northwestern Ar- 
kansas, and eastern Oklahoma; and on the Edwards Plateau, Texas. “Its 
rarity, discontinuous distribution, lack of related species in the New 
World, and occurrence as a pioneer on rocky cliffs all suggest that [it] 
is an old species formerly of general distribution but now approaching 
extinction” (Little, p. 23). Trees to about 10 m. tall have been recorded, 
but Sargent (1892) wrote that “during the War of Secession nearly all 
the large specimens were cut down for the dye which the wood yields . . .” 


and monoecism, as well as flowers with 3-locular ovaries, have been ob- 
served occasionally. Short-tongued Hymenoptera and Diptera apparently 
are the principal pollinators. Parthenocarpy seems to be of common oc- 
currence, since many fruits fail to develop seeds. Although birds have 


? Since the above was written, chromosome numbers of three taxa of Cotinus have 
been determined by Dr. Otto T. Solbrig, of the Gray Herbarium, from staminate 
material collected by Dr. C. E. Wood from the living ae of the Arnold Arbo- 
retum: C. obovatus (Arnold Arb. No. 1814-B, from seed, Charles Mohr, Mobile, 
Alabama, ae 5, 1882; Wood 9427 [aan]), 2n = 15%; C. Coggygria (Arnold Arb. 
No. 276, from Harvard Bot. Gard., 1876; Wood 9428 [aaH]), 2n = 15; and C. 
Coggygria ‘Atropurpureus’ (Arnold Arb. a 708-49-A, from seed, New York Bot. 
Gard., 1948; Wood 9429 [AaH]), 2n = 1 


368 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLIII 


been mentioned, wind is assumed to be the principal means of seed dis- 
persal by transport of the whole infructescence. Asexual reproduction 
takes place by root- and stump-sprouts. 

In spite of many similarities with Rhus, from which Cotimus differs 
mainly in the asymmetrical ovaries and fruits, unequal styles appearing 
sublateral in flower and lateral in fruit, very scanty mesocarp, and pe- 
culiar infructescences, the two genera do not seem to be very closely 
related. According to Engler, Cotinus represents a branch of a line of 
evolution different from but with a common origin with that of RAws. 
Some relationship to the South African genera Laurophyllus Thunb., 
Smodingium E. Mey., and Loxostylis Spreng. f. ex Reichenb, (Botryceras 
Willd.) seems possible. 

The leaves and bark of Cotinus Coggygria are important sources of 
tannin and have been employed extensively in the tanning industry in 
southern Europe. Leaves and flowers yield an aromatic oil. The orange- 
yellow dye (fustic, ‘young fustic’’) extracted from the heartwood of 
C. Cogevgria and C. obovatus was formerly used extensively for dyeing 
silk, wool, etc. Both species are ornamental. 


REFERENCES: 

See also under family references BARKLEY (1937, pp. 300-303), ENGLER 
(1881, p. 378; 1883, pp. 349-352; 1892, pp. 164, 165), HermscH (1940; 1942, 
pp. 139-141), Recorp (p. 23), and SARGENT (pp. 657, 658). 

Buck Ley, S. B. Rhus cotinoides, Nutt. Proc. Acad. Nat. Sci. Phila. 1881: 
125. 1881. gece in Alabama. | 

HorHAmMMer, L., H. WacNner, J. IzQuterpo, & H. Enpres. Isolierung eines 
aie aus Cotinus Coggygria Scop. (Rhus Cotinus L.) 
durch Chromatographie an Polyamid. Arch. Pharm. 291: 269-273. 1958. 

Kinc, H. G. C., & T. WHire. colouring matter of Rhus Cotinus wood 
(young fustic). Chem. Soc. Jour. 1961: 3538, 3539. 1962.* 

Litter, E. R. American ae (Cotinus obovatus Raf.), one of 
Oliahoma rarest tree species. Proc. Okla. Acad. Sci. 23: 21-23. 1943. 
[C. obovatus Raf. correct name for the sp.; distribution and a new locality 
in Oklahoma. | 

MEEHAN, T. Sex in Rhus Cotinus. Proc. Acad. Nat. Sci. Phila. 1873: 300. 1873. 

On hermaphroditism in Rhus Cotinus (the Mist Tree) and in Rhus 
ee (Common Sumac). Proc. AAAS 22(B): 73-75. 1874. 

Osmota, N. Ku. The course of accumulation of tannides in the leaves of 
Cotinus Coggygria Scop. (In Russian.) Bot. ry. 43: 581-583. 1958. 
| Includes notes on economic importance. | 

Penzes, A. Data to the ecology and taxonomy of the Cotimus genus. Acta Bot. 
Sinica 7: 167-169. 1958.* 

SARGENT, C.S. Cotinus. Silva N. Am. 3: 1-6. pls. 98, 99. 1892. 

ScuHuiz, A. Beitrage zur oe und Biologie der Bluthen. IJ. Ber. 
Deutsch. Bot. Ges. 395-409. 1892. [C. Coggygria, 395-401. | 


ic 


4. Metopium P. Browne, Civ. Nat. Hist. Jamaica 177. 1756. 


Trees or shrubs with resin ducts in bark and wood. Leaves odd- 
pinnate, (1)3-—7-foliolate, stoutly petioled, usually clustered near the tips 


1962 | BRIZICKY, GENERA OF ANACARDIACEAE 369 


of branches, persistent; leaflets long-petioluled, + leathery, entire, 
lustrous, veins and veinlets prominent. Plants usually dioecious. Flowers 
small, usually unisexual; pedicels short, stout, and articulate near base, 
in loose, ascending, determinate axillary panicles; bracts and bractlets 
minute, in part persistent. Sepals 5, connate into a + cupuliform 5-lobed 
calyx; lobes broadly ovate to almost semicircular [or truncate], + un- 
equal, thin-leathery with scarious margins, persistent. Petals 5, twice 
as long as sepals, yellow-green, with brownish to blackish veins. Stamens 
5, distinct, alternate with and shorter than the petals, inserted at base 
and between the glandlike lobes of an intrastaminal nectariferous disc, 
reduced and sterile in ? flowers; anthers oblong in outline, nearly as long 
as jor longer than] the subulate filaments, 2-locular at anthesis; pollen 
3-colpate, medium sized, ellipsoidal, reticulate- striate, with round, smooth 
germ pores. Gynoecium 3-carpellate, rudimentary in ¢ flowers. Stigma 
3-lobed; style very short, stout; ovary 1-locular by abortion, obovate to 
subglobular; ovule raised on a basal funicle. Drupe ellipsoidal to some- 
what obovoid, about 1-1.5 cm. long, glabrous, shining, orange to scarlet 
when ripe, tipped with the remnant of the style; exocarp membranaceous, 
mesocarp resinous, endocarp thin, crustaceous, all permanently united 
(coherent). Seed compressed, almost quadrangular in outline, the broad 
funicle covering one of its margins; testa thin, smooth, dark brown; 
endosperm scanty or wanting; embryo oriented eerccally, the cotyledons 
accumbent, flat, the radicle long, incurved upwards toward the hilum. 
TYPE SPECIES: Terebinthus Brownei Jacq. = M. Brownei (Jacq.) Urban. 
(An ancient name for some oriental plant [perhaps some species of 
Ferula|, yielding the gum resin galbanum, applied by Browne to his genus; 
etymology obscure.) 


A genus of three species, of the West Indies, southern Florida, British 
Honduras, Guatemala, and southern Mexico. The West Indian Metopium 
toxiferum (L.) Krug & Urban, poisonwood or coral sumac (also hog gum, 
doctor gum), occurs in hammocks, pinelands, and coastal sand dunes in 
southern Florida (about as far north as Martin County) and on the 
Florida Keys. A large tree in hammocks, a shrub in pinelands, in its 
appearance the species resembles Bursera Simaruba, with which it is often 
confused. 

The genus is very closely related to RAus, but the combination of the 
single style and stigma, the thin, crustaceous endocarp, the vertical 
embryo, and anatomical characters (diffuse-porous wood; abundant, often 
banded, vasicentric parenchyma; septate wood fibers) seems to support 
the maintenance of Metopium as distinct. 

All species are poisonous, and all parts of the plants act as a contact 
skin-poison. Leaves and resin were formerly used in local medicine in the 
West Indies. The floral biology probably does not differ much from that 
of Rhus. Birds apparently are responsible for seed dispersal. 


370 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


REFERENCES: 

See also under family references BARKLEY (1937, pp. 303-309), ENGLER 
(1881, p. 378; 1883, pp. 367-369; 1892, p. 167), Hermscu (1940; 1942, pp. 
139- Recorp (pp. 28, 29), SARGENT (pp. 658, 659), and West & ARNOLD 
(p. 112). 

Harrar, E. S., & J. G. Harrar. Guide to southern trees. 712 pp. New York. 
1946. [M. toxiferum, 435-437. ] 

SARGENT, C. S. Rhus Metopium. Silva N. Am. 3: 13, 14. pls. 100, 101, 1892. 
[M. toxiferum; not Rhus Metopium = M. Brownei el 


5. Rhus Linnaeus, Sp. Pl. 1: 265. 1753; Gen. Pl. ed. 5. 129. 1754. 


Trees, shrubs [sometimes thorny], subshrubs, or vines climbing by 
aérial, clinging, adventitious roots, often pubescent with glandular and/or 
nonglandular hairs, some poisonous by contact. Leaves 3-foliolate | rarely 
palmately 5-foliolate] or odd-pinnate (occasionally even-pinnate) [or 
simple], petioled, deciduous [or persistent]; leaflets usually opposite, 
entire or toothed, sometimes lobed, sessile or + distinctly petioluled, 
rachis sometimes winged. Plants usually dioecious. Flowers small [to 
minute], usually unisexual by abortion, pediceled [or sessile], in termi- 
nal [and]/or axillary thyrses [or panicles], or in solitary or panicled 
racemes [or spikes]; bracts deciduous or persistent, bractlets 2, at base 
of pedicel, deciduous or persistent, or wanting. Sepals 5, connate at base 
[or higher], usually persistent. Petals 5, longer than the sepals, greenish- 
white to yellow [or rarely pink], inserted at base of disc, imbricate. 
Stamens 5, alternipetalous, inserted below a 5—10-lobed, annular, patelli- 
or cupuliform intrastaminal nectariferous disc, reduced and _ sterile in 
@ flowers; anthers ovate to oblong in outline, usually shorter than the 
+ subulate filaments; pollen 3-colpate, medium sized [or small], ellip- 
soidal to spherical, reticulate, reticulate-striate or striate, more rarely 
smooth, with oblong, mostly irregular to ragged, germ pores. Gynoecium 
3-carpellate, rudimentary in ¢ flowers; stigmas 3, capitellate, sometimes 
slightly depressed; styles 3, terminal, distinct or partly [rarely + com- 
pletely] connate; ovary 1-locular; ovule raised on an elongate ascendent 
basal funicle. Drupe subglobular to globular, 3-8 mm. [rarely over 1 
cm.| in diameter, often somewhat laterally compressed, white, tan to 
brown, or red [or black], smooth or sometimes striate [rarely verruculose], 
glabrous or pubescent with glandular and/or nonglandular hairs; exocarp 
membranaceous, sometimes fragile and bursting irregularly; mesocarp 
thin or thick, + resinous, sometimes also waxy; stone + laterally com- 
pressed, smooth or ridged, sometimes bumpy, with thick, bony endocarp. 
Seed + laterally compressed, ovate to subreniform in outline; testa 
membranaceous; endosperm scanty or wanting; embryo large, generally 
transverse, with flat transverse cotyledons and a + elongate radicle in- 
curved upwards toward the hilum. (Including Lobadium Raf. |Schmaltzia 
Desv. ex Small emend. Greene], Toxicodendron Mill.). Lectotypr 
spEcIES: R. Cortaria L.; see E. L. Greene, Leafl. Bot. Obs. Crit. 1: 114. 


1962 | BRIZICKY, GENERA OF ANACARDIACEAE 371 


1905. (Name Latin, from Greek, rkous or rhoys, an ancient name for the 
Sicilian sumac, R. Coriaria; etymology obscure.) 


A genus of over 150 species, primarily of the warm-temperate areas of 
both hemispheres, but extending into tropical and cold-temperate regions. 
The genus seems to be somewhat heterogeneous, and its taxonomy is 
difficult and confused. Both the four sections of Engler, based mainly on 
fruit morphology, and some fractions of the sections have been treated 
by Barkley as distinct genera. Although thorough investigations may 
perhaps justify at least some of these segregations, a complete study of 
the entire complex is necessary. The relationships appear to be reticu- 
late, with speciés showing transitions in respect to most characteristics 
used for delimiting genera, and many supposed differences are based on 
incomplete observations or on insufficient material. At present it appears 
preferable to regard Lobadium Raf. and Toxicodendron Mill. as sub- 
genera of Rhus. 

Subgenus Ruus (subg. Sumac Torr. & Gray, 1838; § Trichocarpae 
Engler, 1881, in part), includes about ten species (primarily of warm- 
temperate North America and Eurasia) with flowers in terminal thyrses 
expanding after the leaves; thin, narrow, caducous bracts and bractlets; 
reddish drupes covered with + club-shaped hairs and sometimes also with 
slender red to colorless nonglandular hairs; exocarp adherent to the 
resinous mesocarp and both easily detachable from the smooth, bony 
stone; and odd-pinnate leaves. All (at least in our area) are innocuous 
trees or shrubs. The northeastern American Rhus typhina L. (R. hirta 
(L.) Sudw.); the wide-ranging R. glabra L.; the very local R. Michauxii 
Sarg., of the Piedmont of North Carolina and Georgia; and the widely 
distributed R. copallina L. (including R. leucantha Jacq. and R. obtusi- 
folia Small) represent the subgenus with us. Presumed natural hybrids of 
R. glabra & typhina have been known as R. glabra var. borealis Britt. 
(R. borealis (Britt.) Greene, R. pulvinata Greene), while R. Ashei 


copallina (including vars. copallina, latifolia Engler, leucantha (Jacq.) 
DC., obtusifolia (Small) Fern. & Grisc., and lanceolata Gray [the last 
supposedly beyond our range] is much needed. 

Subgenus Lopapium (Raf.) Torr. & Gray, 1838 (Lobadium Raf., 1819; 
Schmalizia Desv. ex Small emend. Greene, 1905; Rhus § Trichocarpae 
Engler, 1881, in part; including Styphonia Nutt. and Rkoeidium Greene) 
includes species with flowers in short, dense, spikelike racemes usually 
forming terminal panicles expanding before the leaves (more rarely in 
axillary and/or terminal panicles expanding after the leaves); leathery, 
broad, persistent (rarely caducous) bracts and bractlets; drupes essentially 
as in subg. Ruvus (but in some with the inner layers of mesocarp, striate 
by numerous resin ducts, remaining attached to the stone); and leaves 
3-foliolate (in ours). The nearly 35 species, all innocuous shrubs [or 
trees], are centered in Mexico, extending north to Alberta and Quebec 


oie JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


and south to Costa Rica. The northeastern RAus aromatica Ait. var. 
aromatica (Schmaltzia crenata (Mill.) Greene), fragrant sumac, with 
precocious flowers, is of frequent occurrence on calcareous soils through 
our area southward to northwestern Florida and westward and northward 
beyond our limits. The var. serotina (Greene) Rehd. (R. ¢trilobata Nutt. 
var. serotina (Greene) Barkley), with coetaneous flowers, occurring north 
and west of our region has been recorded from Arkansas. Hybridization 
between R. aromatica and the largely western American R. trilobata, as 
well as between some other species of the subgenus in areas of overlap, is 
believed possible (Barkley, 1937). 

Subgenus ToxIcoDENDRON (Mill.) K. Koch, 1853, emend. Gray, 1856 
($ Venenatae Engler; § Trichocarpae Engler, as to R. trichocarpa Miq.; 
Toxicodendron . Mill., 1754), comprises species with flowers in axillary 
thyrses expanding after the leaves; caducous bracts; no bractlets; vel- 
low-white to -brown, usually smooth drupes, glabrous or pubescent with 
nonglandular hairs; usually thin, fragile exocarp which finally breaks, 
exposing the whitish, waxy mesocarp striate with black resin ducts; stones 
bony with a few riblike longitudinal ridges (or more rarely smooth); and 
leaves 3-foliolate or odd-pinnate [rarely simple]. The nearly 15 species, 
including trees, shrubs, or woody vines climbing by aérial roots and all 
producing a contact dermatitis, are largely of temperate North American— 
eastern Asiatic distribution, but with at least R. striata Ruiz & Pavon in 
tropical South America (Colombia, Venezuela, Peru). Our representatives 
are Rk. Vernix L. (Toxicodendron Vernix (L.) Kuntze), poison sumac, 
a shrub or small tree with odd-pinnate leaves, chiefly of the Coastal Plain: 
R. radicans L. (T. radicans (L.) Kuntze), poison ivy, 2n = 30; and R. 
Toxicodendron L. (T. Toxicodendron (L.) Britt., 7. quercifolium (Michx.) 
Greene), poison oak. RAus radicans is both widely distributed and vari- 
able, and a number of varieties and forms have been recognized. According 
to Gillis (1962, p. 21), the eastern Asiatic R. orientalis (Greene) Schneider 
is conspecific with R. radicans. The external and internal morphology of 
R. diversiloba Torr. & Gray, western poison oak, 2n = 30, is better known 
than that of other species. 

The genus seems to be rather uniform in regard to floral morphology 
and biology. Flowers are usually unisexual and the species dioecious, but 
the occasional occurrence of bisexual flowers, and consequent polygamy, 
is probable. Various Hymenoptera (especially short-tongued bees) and 
Diptera have been recorded as the most frequent visitors. Chalazogamy 
has been found in all the species thus far investigated. The chromosome 
number 2m = 30 has been recorded for three or four species of subg. Toxt- 
CODENDRON, 2” = 32 for the North African Rhus oxyacantha Schousb. 
ex Cav. (Searsia Barkley; § Gerontogeae Engler). Seed dispersal by 
animals, especially by birds, apparently may be assumed for the genus as 
a whole. 

Dried leaves of Rhus glabra, R. typhina, and R. copallina, and espe- 
cially of R. Coriaria, are important sources of tannin. Leaf galls produced 
on R. javanica L. (R. chinensis Mill., R. semialata Murr.) vield tannin 


1962 | BRIZICKY, GENERA OF ANACARDIACEAE Bike. 


which is also sometimes used in making ink. The Asiatic R. verniciflua 
Stokes and sometimes R. succedanea L. are sources of natural lacquer. 
Commercial vegetable wax is obtained from the mesocarp of fruits of the 
last and some allied species. 


omacraces 
e large number of references has been reduced here primarily to those 
ae of general interest or dealing specifically with the southeastern United 
States. Under family reference see BARKLEY (1937, pp. 312-441), ENGLER 
(1881; 1883, pp. 371-452; 1892, pp. 167-172), HermscH (1940; 1942, pp. 
139-141), Recorp (pp. 3034. 43-45), SARGENT (pp. 660-665), and West & 
ARNOLD (pp. 113, 114). Under Cotinus see MzEHAN (1874). 
ASHE, W. W. Magnolia cordata and other woody plants. Bull. Torrey Bot. 
Club 54: 579-582. 1927. [“ Schmaltzia “shel hyb. nov.” (= Rhus Ashei), 


= 
Baer, J. E., & A. F. Sievers. Sumac; its collection and culture as a source of 
tannin U.S. Dep. Agr. Prod. Res. Rep. 8. 14 pp. 1957.* 
Bark ey, F. A. Five-leaved poison ivy. Biologist 17: 122-124. 1936.* 
. Studies in the Anacardiaceae. III. A note concerning the status of 
Rhus pulvinata Greene (R. glabra & typhina Koehne). Am. Midl, Nat. 
1938 


19: 598-600. 

——.. Schmaltzia. ee 24: 647-665. 1940. 

& E. BaRKLEY. A short history of Rhus to the time of Linnaeus. 

Ibid. 19: 265-333. 1938. 

Boyp, I. L. Germination tests on four species of sumac. Trans. Kan. Acad. Sci. 
46: 85, 86. 1943 

Crausen, R. T. Northeastern limits of the known range of Rhus Toxico- 
dendron. Torreya 41: 58, 59. 1941. [R. radicans.] 

CopELAND, H. F., & B. E. Dovet. Some features of the paar of Toxicoden- 

0.] 


dron diversiloba. Am. Jour. Bot. 27: 932-939. 1941. [2n 

Crooxs, D. M. L. W. Kepuart. Poison-ivy, poison-oak, i poison sumac: 
identification, precautions, eradication. U. S. Dep. Agr. Farmers’ Bull. 
1972: 1-30. 1958. 

Dawson, a R. The toxic principle of poison ivy and related plants. Rec. Chem. 
Prog. 15: 39-53. 1954.* 

: ci chemistry of poison ivy. Trans. N. Y. Acad. Sci. II. 18: 427- 
443. 1956.* 

Driers, L. Die Epharmose der Vegetationsorgane bei Rhus § Gerontogeae Engl. 
Bot. Jahrb. 24: 568-647. pl. 14. 1898 

Duncan, W. H., & T. J. Jones. Poisonous plants of Georgia. Bull. Univ. Ga. 
School Veterin. Med. 49(13): i-iv, 1-46. 1949. [R. Vernix, 17-20; R. 
radicans, 20. | 

FERNALD, M. L. Another century of additions to the flora of Virginia. Rhodora 
43: 559-630. pls. 672-692. 1941. [“Some varieties and forms of Rhus 
radicans and R. To: einen. 589-599. pls. 683-685; “The variations 
of Rhus aromatica in the Gray’s Manual range,” 599-603. pls. 686, 687. | 

—— &L. Griscom. Three days of botanizing in southeastern Virginia. /bid. 
37: 167-189. pls. 345-351. 1935. [Variations of R. copallina, 167, 168.] 

FRANKLIN, J. J. Another plant to be labeled poisonous. Pl. Gard. 8: 271. 1952. 
[R. trichocarpa. | 


374 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLIII 


Gitpert, E. F. Phenology of sumacs. Am. Midl. Nat. 66: 286-300. 1961.* 
[R. glabra and R. typhina. See also Diss. Abs. 20: 1553, 1554. 1959.] 
Giiuis, W. T. Taxonomic problems in poison ivy. Pap. Mich. Acad. Sci. I. 45: 
27-34. 1960. 
. Poison-ivy and the poison-oaks. Cranbrook Inst. Sci. News Lett. 30: 
98- 107. es 


on-ivy in northern Michigan. Mich. Bot. 1: 17-22. 1962. [R. 
asda radicans var. Rydbergii, delimitation, range in Michigan. | 

GREENE, E, Segregates of the genus Rhus. Leafl. Bot. Obs. Crit. 1: 114-144. 
1905. 

——. A study of Rhus glabra. Proc. Wash. Acad. Sci. 8: 167- 196. 1906. 
[Includes descriptions of 28 segregates, 24 proposed as new spp. 

Grimm, J. Entwicklungsgeschichtliche Untersuchungen an RAus und Coriaria, 
Flora 104: 309-334. pls. 10, 11. 1912. [R. “Toxicodendron’” (R. radicans 
?), 2n = 30, embryology, 311-323; R. glabra and R. typhina, chalazogamy, 
parthenocarpy, 323-325. 

Harapa, M. On the distribution and construction of the resin canal[s] in Rhus 
succedanes, Bot. Mag. Tokyo 51: 846-856. 1937. 

stigation on the development of resin canals and the formation of 

ne in a mesocarp of fruits of the Rhus plants found in Japan. Bull. 
Sci. Fak. Terkult. Kyushu Univ. 8: 179-191. 9.* 

Kexar, S. S. Embryology of Rhus mysurensis Heyne. Jour. Indian Bot. Soc. 
37: 114-122. 1958. 

LittLe, E. L., Jr. Miscellaneous notes on nomenclature of United States trees. 
Am. Midl. Nat. 33: 495-513. 1945. [RAus, 498-500. | 

McFappen, G. H., & R. L. McMurry. Rhus glabra. Am. Jour. Pharm. 109: 


McNair, J. B. Fats from Rhus [Malosma] laurina and Rhus [ Toxicodendron | 
diversiloba, Bot. Gaz. 64: 330-336. 1917. [Fats identical; includes fruit 
morphology of R. diversiloba. 

Secretory canals of Rhus diversiloba. Ibid. 8: 179-191. pls. 3, 4. 

ZA, 


The taxonomy of poison- ee with a note on the origin of the generic 

name. Publ. Field Mus. Bot. 4: 55-70. pls. 14-24. 1925. 

Five-leaflet poison oak. a Torrey Bot. Club 63: 473-476. 1936. 
[| R. diversiloba, pinnate-leaved form. | 

Meyer, A. Ueber die Entwickelung des Wachses der Frucht von Rhus Toxico- 
dendron Mich. Arch. Pharm, III. 15: 514-516. 1879.* 

Mosivus, M. Der japanische Lackbaum, Rhus vernicifera DC. Abh. Senckenberg. 
Naturf. Ges. 20: 203-247. pl. 1. 1899. [Morphology and anatomy of R 
verniciflua. See also Ber. Deutsch. Bot. Ges. 15: 435-441. 1897. ] 

MorceneverR, W. Untersuchungen tber die Beseitigung der Keimhemmung 
beim Samen des Hirschkolbensumachs (Rhus typhina). Arch. Forst. 5: 
203-242. 1956.* 

MuENSCcHER, W. C., & J. M. Kincssury. Poison ivy and poison sumac. Cor- 
nell Ext. Bull. 191: 1-12. 1960. 

RosBertson, C. Flowers and eS 17. Bot. Gaz. 22: 154-165. 1896. [Rhus, 
159-164. ] 


Flowers and insects. 221 pp. Carlinville, Illinois. 1928. [R. aromatica 
(“R. canadensis”) R. copallina, R. glabra, R. radicans (“R. Toxicoden- 
dron’’), list of insect visitors, 24-26. 
SARGENT, C. S. Rhus. Silva N. Am. 3: 7-11, 15-28. pls. 102-109. 1892. 


1962] BRIZICKY, GENERA OF ANACARDIACEAE 375 


. Rhus Michauxii. Garden Forest 8: 404. 1895. [Considered to be very 

poisonous, but see Warren. 

. Rhus trichocarpa. Ibid. 10: 383, 384. 

SCHNEIDER, C. K. Illustriertes Handbuch der Dine ances Vola2.. 21070" pp: 
Jena. 1907-1912 

SCHONLAND, S. The South African species of Rhus L. Bothalia 3: 1-115. 1930. 


Sizer, I. W., & C. E. Proxescu. Inactivation of the irritant toxicants of 
poison i and related compounds by tyrosinase. Science 101: 517, 518. 
1945. 


TapaTa, S. Ueber die Friichte und Keimpflanzen von Rhus succedanea, L. 
Jour. Coll. Sci. Univ. Tokyo 23(1): 1-11. 1 pl. 1907 

WARAWDEKAR, S. S. Analysis of poison ivy fruit fat. Diss. Abs. 17: 2164. 1957.* 

WarreEN, L. E. Rhus AC POE: A nonpoisonous plant. Am. Jour. Pharm. 
1910: 499-506. 1910 


376 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


COMPARATIVE ANATOMY OF THE LEAF-BEARING 
CACTACEAE, VI 
THE XYLEM OF PERESKIA SACHAROSA 
AND PERESKIA ACULEATA 


I. W. Batvey ! 


THE MOST PRIMITIVE living representatives of the Cactaceae have been 
considered to be Pereskia sacharosa Griseb. and P. aculeata Mill. Such a 
conclusion is based largely upon the claim that the flowers of these 
species have “superior” ovaries (Berger, 1926; Buxbaum, 1953). It is of 
interest to determine whether evidence from other parts of the plants 
supports such a phylogenetic generalization. Evidence from the xylem is 
considered in this paper. Additional evidence from other parts of the plants 
will be considered subsequently. 


PERESKIA SACHAROSA 


During its earlier stages of growth, this species is considered to be 
a shrub which ultimately may become a small tree eight meters high. 
As in the case of other wide-ranging putative species of Pereskia, plants 
from different localities exhibit more or less conspicuous differences in 
their leaves, general habit of growth, spination, and floral characters. 
At present, it is uncertain whether such differences are produced b 
different environmental influences or are due to genetic differences at 
racial or specific levels. 

My anatomical specimens of this species have been obtained from the 
following sources: 1) From a wild plant collected by Carenzo and Legname 
in Jujuy Province, Argentina; 2) from a wild plant collected by Cardenas 
in the Department of Cochabamba, Bolivia; 3) from a plant growing in 
the Missouri Botanical Garden; and 4) from a plant growing at the 
New York Botanical Garden. The ranges of variability in the xylem of 
the four plants are so closely similar as to provide no reliable anatomical 
criteria for differentiating one plant from the others. 

In the outer secondary xylem of larger stems the vessels occur singly 
and in small clusters (Fics. 1, 3), but tend at times in their overall 
arrangement to exhibit more or less conspicuous concentric patterns (Fic, 
1). The wood parenchyma is scanty paratracheal (Fic. 3), having lignified 
walls. The multiseriate rays are composed of cells of varying size, form, 
and orientation which likewise have comparatively thick lignified walls 

'This investigation was financed by a grant from the National Science Founda- 
tion. I am indebted to the American Philosophical Society for the loan of a Wild 
microscope. 


1962 | BAILEY, LEAF-BEARING CACTACEAE, VI 377 


(Fics. 3, 5). Where crystals of calcium oxalate are present in ray cells 
they occur singly or as several independent ones, druses being absent. 
The distribution of vessels and wood parenchyma in comparable secondary 
xylem a roots is similar (Fic. 2) but the multiseriate rays are broader 
(Fics. 2, 6). The rays, as in those of the stem, may be composed, 
ate their radial extension, of cells with lignified walls. In some 
cases, however, the first-formed mmultiseraite rays in the center of the 
root are composed at first of cells having thin, unlignified walls. These 
cells differ from subsequently formed lignified ones in containing isotropic 
granular contents, rather than starch. 

It is evident that the secondary xylem in stems and roots of Pereskia 
sacharosa is of a phylogenetically highly evolved structure such as occurs 
in trees and shrubs of normal form in a number of other dicotyledonous 
families. For a discussion of pitting, perforation plates, and other struc- 
tures visible under higher magnification, the reader is referred to the 
fourth paper of this series (Bailey & Srivastava, 1962 

There are certain ontogenetic changes in rays that should be considered 
in dealing with the secondary xylem of the leaf-bearing Cactaceae. The 
multiseriate rays of dicotyledons commonly exhibit significant changes 
in passing from the first-formed to the later-formed secondary xylem 
(Chattaway, 1933; Barghoorn, 1940, 1941a). The first-formed part of 
multiseriate rays adjacent to gaps in the primary body (particularly of 
stems having elongated internodes) frequently tend to be narrow and 
vertically extensive as seen in tangential longitudinal sections (Fic. 8). 
During subsequent radial extension of such rays they tend to become 
wider and dissected sooner or later into vertically less extensive parts which 
ultimately may assume fusiform outlines (Fic. 6). Of the various cyto- 
logical changes involved in the modification of such rays the transforma- 
tion of ray initials into fusiform cambial initials is particularly significant. 
Furthermore, the dissected parts of the original rays tend to be laterally 
displaced during increase in girth of the cambium. In addition, the cells 
of the ravs generally vary more or less in form and orientation during 
successive modifications of the multiseriate rays, the cells in the first- 
ormed part of the rays tending to have a longer vertical axis (i.e., being 
“erect’’), whereas in subsequently formed parts of the rays they become 
more or less isodiametric or even radially extensive (i.e., ‘‘procumbent”’). 
Such changes in multiseriate rays vary in degree, and may be precocious 
and relatively abrupt, gradual, or considerably delayed, varying at least 
in part with different rates of growth and the enlargement of stems. 

In roots of Pereskia sacharosa, the first-formed multiseriate rays broaden 
conspicuously during their radial extension outward (Fic. 2). In their 
first-formed part they are not only much narrower, but also more ex- 
tensive vertically. During their radial extension outward they tend to 
become dissected into lower derivatives which assume fusiform outlines 
as seen in tangential longitudinal sections (Fic. 6). In stems from some 
parts of a mature plant, the first-formed parts of the rays are narrower 
and more extensive vertically, but subsequent widening and dissection of 


378 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


the rays is much less conspicuous than in roots (compare Fics. | and 2, 
5 and 6). In other stems from the same plant, the first-formed rays of 
the secondary xylem are broader, subsequently becoming narrower (Fic. 
4) and then broadening somewhat in the later-formed xylem (Fic. 3). 

Such variations in the form of the multiseriate rays appear to be corre- 
lated with differences in the development of the primary body. The 
diameter of the primary vascular cylinder and pith commonly varies in 
different stems of the same plant from three millimeters to one centimeter. 
As seen in transverse sections, the two extreme forms of eusteles may be 
composed of the same number of fascicular and interfascicular parts. 
Although there is some variation in the breadth of fascicular parts, the 
conspicuous differences in circumference of the eusteles appears to be due 
largely to differences in the width of interfascicular parts (i.e., paren- 
chymatous gaps). In the case of the smaller young stems, cambial activity 
develops precociously in the parenchyma of the narrow gaps, and the 
multiseriate rays of the secondary xylem are relatively narrow when first 
formed. On the contrary, in the larger young stems the parenchymatous 
gaps tend to widen by division and transverse expansion of their con- 
stituent cells for a considerable time after cambial activity is initiated in 
the fascicular parts of the eustele (Fic. 4). It should be noted in this 
connection that the widening of the interfascicular parts and the delayed 
initiation of cambial activity within them leads, not only to a conspicuous 
increase in the circumference of the eustele, but also permits a marked 
increase in the diameter of the pith. Furthermore, when multiseriate rays 
of the secondary body are initiated they tend at first to be considerably 
broader than in the case of smaller stems (Fic. 4). 

In both stems and roots of Pereskia sacharosa, the parenchymatous 
cells of the first-formed part of the multiseriate rays are of “erect” form 
but are shorter than the fusiform initials of the cambium. Subsequent 
changes in the size, form, and orientation of the ray cells, as seen in 
radial longitudinal sections of the xylem, are precocious and abrupt. 
Most of the cells become more or less isodiametric, except on the sides 
of the rays and their upper and lower margins where they may retain 
an erect form. In addition, some radial tiers of slender cells of procumbent 
orientation tend to occur within the rays. 


PERESKIA ACULEATA 


This species is widely distributed in the West Indies and in eastern 
and northern South America. Because of the edibility of its fruits and 
leaves, its ornamental value in gardens, and its use in hedges, its original 
range has been considerably extended by man. It is therefore difficult 
at times to determine with certainty whether plants now growing in the 
wild (e.g., Florida or Mexico) are native or were introduced and escaped 
from cultivation. 

The species is commonly described as a shrub, frequently producing 
long clambering branches and forming vines three to ten meters long 


1962 | BAILEY, LEAF-BEARING CACTACEAE, VI 379 


(cf. Britton & Rose, 1919). The larger, older stems may have a diameter 
of two to three centimeters with a primary vascular cylinder and pith 
five to six millimeters in diameter. The ultimate terminal branches of 
mature plants are, in general, more numerous and more slender than 
those of Pereskia sacharosa, grading down in many cases to as little as 
two millimeters in diameter with correspondingly slender primary vas- 
cular cylinders and pith. Furthermore, in branches of comparable diameter, 
the eusteles of P. aculeata are composed of fewer fascicular strands and 
oe ane broader parenchymatous gaps. This conspicuous difference 

n the primary bodies of the two species appears to be correlated with 
ce tendency of P. aculeata to have two-trace unilacunar nodes, whereas 
P. sacharosa commonly has four to six vascular strands at its unilacunar 
nodes. (For illustrations see Bailey, 1960.) 

The young branches of Pereskia aculeata are commonly of two different 
forms: slender woody ones and broader, very succulent ones, referred to 
in Boke’s (1954) developmental investigations as long shoots and spur 
shoots respectively. In the former stems cambial activity and the forma- 
tion of secondary xylem are precocious, whereas in the latter stems the 
formation of secondary xylem is delayed and the thickness of the pith 
and cortex is relatively greater. At times, the most slender woody stems 
bear short, broad, highly succulent laterals. In such cases the slender 
woody stems have long internodes, whereas the succulent laterals have 
short internodes. These abrupt differences in external form and internal 
structure resemble those between long shoots (pycnoxylic”) and short 
shoots (manoxylic *) that occur in Ginkgo biloba L. (Gunckel & Wetmore, 
1946a, 1946b) in Cercidiphyllum japonicum Sieb. & Zucc. (Titman & 
Wetmore 1955), and a number of other plants. 

The structure of the secondary xylem of Pereskia aculeata varies 
markedly in different parts of a mature plant. As so frequently happens 
in scandent representatives of other dicotyledonous families, e.g.. Hippo- 
crateaceae, Icacinaceae, Schisandraceae, etc., many stems exhibit more or 
less abrupt transitions from first-formed, denser, more nearly normal 
secondary xylem to phylogenetically highly modified, softer, more porous 
forms of tissue. The denser, earlier-formed secondary xylem (illustrated 
in Fic. 7) resembles the secondary xylem of P. sacharosa (Fic. 7). The 
vessels, allowing for higher magnification, are of comparable size, form, 
and distribution. The wood parenchyma is scanty paratracheal and has 
strongly lignified walls. The multiseriate rays, although somewhat nar- 
rower than those of P. sacharosa, have thick, strongly lignified cell walls. 
On the contrary, the later-formed secondary xylem has much enlarged 
vessels, as well as radial rows of much smaller ones. In addition to scanty 
wood parenchyma about vessels in the denser parts, it has arcs or con- 
centric zones of unlignified parenchyma. The cells of the broadened rays 
(compare Fics. 2 and 3) are very thin, unlignified, and many of them 
contain druses (Fic. 16). Furthermore, the number of thick-walled libri- 
form fibers is greatly reduced. 


? Using these terms as redefined by Titman and Wetmore. 


380 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


In other stems, even from the same plant, the transition from normal 
dense to modified secondary xylem occurs at varying distances from the 
pith. The transition to xylem containing huge vessels may be abrupt or 
gradual (Fic. 11). Furthermore, the volume and distributional patterns 
of the unlignified ® parenchyma varies markedly from stem to stem. In 
the xylem illustrated in Fic. 10, the unlignified parenchyma occurs in 
broad concentricities alternating with zones of strongly lignified tissue. 
The relatively narrow multiseriate rays, unlike those of Fic. 7, have ligni- 
fied walls, except where they pass radially through the broad zones of 
unlignified parenchyma. In contrast to this, the outer secondary xylem 
in Fic. 11, although exhibiting a conspicuous enlargement of vessels has 
unlignified parenchyma that is confined to narrow arcs and concentricities. 

The roots of Pereskia aculeata superficially resemble those of P. sacha- 
rosa, having numerous rapidly broadening multiseriate rays (compare 
Fics. 2 and 14), but the ray cells of P. aculeata, many of which contain 
druses of calcium oxalate, differ in having thin, unlignified walls (Fic. 14). 
It should be noted in this connection that the first-formed ray cells in 
the center of some roots of P. sacharosa, although having thin unlignified 
walls, contain no druses and little if any starch. Furthermore, in roots, 
as in stems (Fics. 8, 9), of P. aculeata, the multiseriate rays tend to 
remain vertically extensive, and in passing radially outward do not 
exhibit precocious and conspicuous dissection into laterally displaced parts 
as illustrated for P. sacharosa in Fic. 6. The unlignified rays of both 
te and stems of P. aculeata, not only contain abundant druses (Fic. 

16) but also at times varying numbers of mucilage cells (Fic. 17). 

In many stems of Pereskia aculeata, owing to the breadth of the 
parenchymatous gaps in the eustele, the multiseriate rays tend to be 
broader when first formed on in the subsequently formed secondary 
xylem. As indicated in Fic. 11, this reduction in ray width resembles 
that which occurs in very stout young stems of P. sacharosa (Fic. 4). 
However, in the case of P. aculeata, the cells of the first-formed ray 
tissue tend to resemble fusiform cambial initials in their vertical exten- 
sion (Fic. 12). In other words, the first-formed ray initials of the 
cambium resemble fusiform initials in size and orientation. Subsequent 
transverse divisions of the ray initials lead to the formation of less ‘erect’ 
ray cells (Fic. 13) and ultimately to more or less isodiametric or some- 
what procumbent ones. It should be noted in this connection that simi- 
larities between ray initials and fusiform initials occur in the first-formed 
cambium of some other dicotyledonous families (Barghoorn, 1941). 
Tyloses are of sporadic occurrence in the vessels of Pereskia aculeata. 
n some stems, they are not only abundant but contain starch (Fic. 15). 


— 


—_ 


“The unlignified primary walls of leaf-bearing Cactaceae give a red coloration 

sections stained ae Haidenhain’s perenne and safranin. Therefore, in 
testing for lignificati it is advisable to use the phloroglucin- eae and Maule 
reactions where unlignified walls remain ieee as in Fics. 10 and 


1962] BAILEY, LEAF-BEARING CACTACEAE, VI 381 


DISCUSSION 


In the fourth paper of this series (Bailey & Srivastava, 1962) we dis- 
cussed the finer structural details in derivatives of the fusiform initials 
of the cambium. We found that the ranges of variability of potential 
diagnostic criteria in the leaf-bearing genera Pereskia, Pereskiopsis, and 
Ouiabentia are extensive, not only in different collections of the same 
taxon and in the same clone grown under different environmental influ- 
ences, but also in different parts of the same mature plant. It is evident 
that the grosser anatomical features of Pereskia aculeata (e.g., size and 
distribution of vessels, abundance, distributional patterns. and internal 
structure of wood parenchyma, and the size, form and internal structure 
of the multiseriate rays) likewise vary markedly even in different parts 
of a single plant. 

Anatomical surveys of the dicotyledons as a whole demonstrate that 
the xylem of vines and lianas commonly exhibits trends of highly ad- 
vanced phylogenetic modification, i.e., in comparisons with the structure 
of related trees and large woody shrubs of the same genus, family, or 
order. This obviously occurs in Pereskia aculeata in contrast to P. sacha- 
rosa. Thus, although P. sacharosa may have retained a form of xylem 
characteristic of ancestral leaf-bearing cacti, the internal structure of 
stems and roots of P. aculeata negates any possibility of considering this 
species to be the most primitive living representative of the Cactaceae. 
Therefore, in searching for such a primitive representative, it will be of 
interest in succeeding papers of this series to compare the internal struc- 
ture of P. sacharosa with that of other arborescent and woody, shrubby 
species of Pereskia. 

The tendencies in the xylem of Pereskia aculeata toward the formation 
of parenchymatous cells having thin, unlignified walls, with concomitant 
inclusions of druses, are extensively developed in roots of Andean pereskias 
and in stems of Pereskiopsis and Quiabentia. They seem to be correlated 
in some manner with increasing succulence and merit detailed investiga- 
tion. Furthermore, the tendency in some stems of both P. sacharosa and 
P. aculeata toward increase in circumference of the primary vascular 
cylinder and in diameter of the pith after cambial activity is initiated 
in the fascicular parts of the stele is a phenomenon which becomes greatly 
exaggerated in many representatives of the Cactaceae. It is a conspicuous 
feature in dealing with the larger basal stems of pereskias from Southern 
Mexico and Central America and in stems of certain species of Pereskiopsis 
and Ouiabentia. 


LITERATURE CITED 


Bartey. I. W. 1960. Comparative anatomy of the leaf-bearing Cactaceae, 
L. Foliar vasculature of Pereskia, Pereskiopsis and Quiahentia. Jour. Arnold 
Arb. 41: 341-356. 

L. M. Srivastava. 1962. IV. The fusiform initials of the cambium 

and the form and structure of their derivatives. /bid. 43: 187-202. 


382 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLHI 


BarcHoorn, E. S. 1940. The ontogenetic development and_ phylogenetic 
specialization of rays in the xylem of dicotyledons, I. The primitive ray 
structure. Am. Jour. Bot. 27: 918-928 

ky ce of the multiseriate and uniseriate rays. Am. Jour. 

Bot. 28: 273-282. 

. 1941b. III. The elimination of rays. Bull. Torrey Club 68: 317-325. 

Bercer, A. 1926. Die Entwicklungslinien der Kakteen. Gustav Fischer, Jena. 

Boxe, N. H. 1954. Organogenesis of the vegetative shoot in Pereskia. Am. 
Jour. Bot. 41: 619-637. 

BRITTON, N. L., & J. N. Rose. 1919. The Cactaceae. Vol. I. Carnegie Inst., 


BuxpauM, F. 1953. Morphology of Cacti. Sect. IJ. The flower. Abbey 
Garden Press, Pasadena, Calif. 

Cuatraway, M. 1933. Ray development in the Sterculiaceae. Forestry 7: 
93-108. 

GuNCKEL, J. E., & R. H. Wermore. 1946a. Studies of development in long 
shoots and short shoots of Ginkgo biloba L. 1. The origin and pattern of 
development of the cortex, pith and procambium. Am. Jour. Bot. 33: 285-295. 

. II. Phyllotaxis and the organization of the primary vascular system; 

Laie phloem and xylem. /bid. 532-543. 

AN, P. W., & R. H. Wermore. 1955. The growth of long and short 

pens in Corgi phuluti: Am. Jour. Bot. 42: 364-372 


EXPLANATION OF PLATES 


PLATE I 
Fics, 1-3. Transverse sections - the secondary gen: of Pereskia sacharosa. 
1, Outer xylem from stem 7 cm. in diameter, & 11. 2, Xylem ar root 3 cm. 


in diameter, X 11. 3, Xylem of Fic. 1 more highly acute’. x 8 


PLATE II 
Fics. 4-6. Transverse and tangential longitudinal sections of Pereskia sacha- 
rosa showing structure of multiseriate rays. 4, Transverse section of inner 
xylem from a stem 7 cm. in diameter, * 34. 5, Tangential section from outer- 
most xylem of Fic. 1, & 43. 6, Tangential section of outermost xylem of Fic. 2, 
x 43. 


PLATE III 
. 7-9, Transverse and tangential longitudinal sections of Pereskia 
aculeata from stem 2.5 cm. in diameter. 7, Transverse section of xylem, X 34. 
8, Tangential section of denser xylem, X 43. 9, Tangential section of outer, 
softer xylem, * 43. 


PLATE IV 

‘ics. 10-13. Transverse and longitudinal sections of the xylem from stems 
of eae aculeata. 10, Transverse section treated with phloroglucin-HCl, 
showing concentric zones of unlignified parenchyma (white), X 34. 11, Trans- 
verse section of inner xylem, showing ace in ray structure and narrow arcs 
of thin-walled wood parenchyma, * 43. 12, Tangential section of first-formed 
secondary xylem, * 88. 13, Radial Bere showing transitional changes in the 
height of ray cells, & 88. 


1962 | BAILEY, LEAF-BEARING CACTACEAE, VI 383 


PLATE V 
Fics. 14-17. Transverse and tangential longitudinal sections of Pereskia 
aculeata. 14, Transverse section of root, treated with phloroglucin-HCl, showing 
unlignified multiseriate rays (white), X 34. 15, Transverse section of vessel, 
showing starch in tyloses, X 107. 16, Part of Fic. 9, more highly magnified, 
showing druses in multiseriate ray, X 107. 17, Tangential section of xylem, 
showing mucilage cells (arrows) in multiseriate ray, X 107. 


Jour. ARNOLD Ars. VoL. XLIII 


+e 
* 

4, 

6¢ 


ma) ewe?’ 
‘ene a* 94 
vse! vee 
Natt o'e que M MASALA > 
Xd Paree eg & ot, s00 “ 
| Se ketones eo! “9. Oa, Fx 
ey 0 9 ‘se, That e Re 
Al “ed 


“0088 Ry 
F Pee ey yee 


Ud EA | 
a Ore @,;, ‘3 
ae Cees er 


; = é 4 

4s e ote ONS! 

Spey. Seite tes 

Ener 

en NERO Cat 

: OSI Beak: a 

be die Fy } 6. VOM Oe are Ses 
16-@ 9 bak i as shale RN aE 
Ly : 4 8.06 0 hey 
baie tae Por i RR 
oagg lt ; sine: ry soe SPEER) LS, 
ented ‘ae. ered LED 3 Tie SO A IN Diem NE 
BOTY “AO NR SR 


BAILEY, LEAF-BEARING CACTACEAE, VI 


PLATE II 


Jour. ARNOLD Ars. VoL. XLII 


‘©; —— 
ILS 


Es 


i : etienece 
SSeS: 


= 


iF AY 


sesegs ame 


Bes 


dy 


Ss 


VI 


) 


-BEARING CACTACEAE 


, LEAF 


BAILEY 


Jour. ARNOLD Ars, VoL. XLIII 


PLaTE III 


cat. 


a 
tj 


\ 
iy 
ire 


ee ee 

= Peon a, 
few te con a a ae 
wwe SES 
bee 


a a 
” 


sod 


ee we 


> 


es 


Se 
ee 
om 

r 


Jour. ARNoLtp Ars. VoL. XLII PLATE IV 


ee om 
ten 
To 
Soe eS 
eda 
< 


Raemeeel 
fran an 


i 

i 
Trout | 
in oan 


fA 


re) 


iN 
ss tel oli m 
(rary 


\ ECS 


7a 


H 


bake oe 
pat mwas 2 
os wer: "9 

bad he , 

Shed! BEA a 

4 Piety) 
\) “3 


e rt 
eatery et 
ae Oa war 
ee: 
“ratterae 


nest 
AE cae aera 


es. 


MN 
il 
i 


Ed 


BAILEY, LEAF-BEARING CACTACEAE, VI 


PLATE V 


Jour. ARNoLD Ars. VoL. XLITI 


BAILEY, LEAF-BEARING CACTACEAE, VI 


1962 | HOWARD, GUTTIFERAE 389 


SOME GUTTIFERAE OF THE LESSER ANTILLES 


Ricuarp A. Howarp 


IN THE cCouRSE of preliminary work leading towards the preparation 
of a flora of the Lesser Antilles, I have encountered several nomenclatural 
problems in the genera Clusia, Calophyllum, and Rheedia. The following 
notes are presented as discussion and clarification of these problems. 

I have received most generous assistance from Miss Alicia Lourteig, 
Mr. George Proctor, and Mr. James Dandy in checking historic specimens 
and illustrations in European herbaria. It is also with gratitude that I 
acknowledge the support of this work through a grant from the National 
Science Foundation. 


CLUSIA 


Britton and Millspaugh (Bahama Fl. 281. 1920) and Hitchcock and 
Green (Prop. Brit. Bot. 160. 1929) selected Clusia major L. as the 
type species of the genus. More recently the /ndex Nominum Genericorum 
formalized this status. Unfortunately, Clusia major, proposed by Linnaeus 
in the first edition of Species Plantarum, was abandoned by him in the 
second edition and has not been used in floras or monographs in the 
intervening 209 years. In the interest of stability of well known specific 
names, one questions the value and the necessity of resurrecting such an 
epithet. There is, however, no option in the present rules of nomenclature, 
and so Clusia major L. must replace the better known Clusia alba Jacq. 
In the process of this investigation, it became apparent that the nomen- 
clature of nearly every species of Clusia in the Lesser Antilles was in- 
volved, and several others in the Greater Antilles presented one or more 
additional problems. These will be discussed in the following paragraphs. 


Clusia major L. 

In the first edition of Species Plantarum o 509. 1753), Linnaeus 
described two species of Clusia, namely, C. major and C. minor. The 
protologue of C. major, with the modern ees ts of its supporting 
literature given in brackets, is the following: 


— 


CLUSIA foliis aveniis. major. 
Clusia flore albo, fructu coccineo. Plum. gen. 21. [C. alba Jacq. | 
Cenchramidea arbor saxis adnascens, obrotundo pingui 

folio, fructu pomiformi. Pluk. alm. 92 t. 157. f. 2. 

[C. plukenetit Urb. | 
Terebinthus folio singulari non alato rotundo succu- 

Jento, flore pallide luteo. Sloan. jam. 167. hist. 2. 

p. 97 t. 200. f. 1 [C. flava Jacq.] Raj. dendr. 51 [unknown] 


390 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLIII 


8. Clusia flore roseo major, fructu subviridi, Plum. gen. 21. 
[C. rosea Jacq. | 

y. Clusia alia minor: flore albo, fructu virescente. Plum. 
gen. 21. [possibly C. plumieri, or not known according 
to Planchon & Triana’ 


In considering this species, the specimens in the Linnaean herbarium 
are of little assistance. There are five sheets included under the name 
of Clusia. Sheet 1224.1 does not appear to be a Clusia. Sheet 1224.2 is 
annotated, “Clusia alba H. Aublet vix Linnaei J. E. S.”’ The specimen is 
probably a Clusia. Sheet 1224.3 bears only two detached leaves, One 
is obviously sessile and is probably Clusia mangle L. C. Rich. ex Planch. 
& Triana. The other leaf has a petiole and is annotated as Clusia flava 
ex Herb. Jacq. The remaining two sheets, 1224.4 and 1224.5, represent 
Mammea americana and Chrysobalanus icaco respectively. 

In 1760, in his Enumeratio Systematica Plantarum, Jacquin described 
four species of Clusia but made no reference to Species Plantarum. The 
species are briefly but validly described and each carries a reference 
to an illustration. Clusia rosea has the reference “Catesb. Car. 2. t, 99.”: 
C. alba carries “Plum, ic. 87. f. 1.°; C. flava bears “Sloane hist. Jam. 2. 
t. 200. f. 1.”; and C. venosa is supported by a reference to “Plum. ic. 87. 
f. 2.” The species published in Jacquin’s Enumeratio were normally based 
on material he collected in the West Indies. Mr. Dandy has pointed out 
in correspondence that Jacquin sometimes cited published figures from 
Browne, Sloane, Catesby, and others, but in doing this his intention was 
to provide the reader with what he supposed was an illustration of his 
own plant. In his later Selectarum Stirpium Americanarum Historia the 
Same species were usually described at length, more complete references 
were given, type localities were cited, and often a figure drawn from 
his own material was published. 

Unfortunately, Jacquin’s herbarium was badly damaged while in the 
West Indies. Although the remains were purchased by Sir Joseph Banks 
and are in the herbarium of the British Museum, there is no material of 
Clusia available. Jacquin, in his Selectarum, in 1763, did publish complete 
descriptions of the four species of Clusia, along with illustrations of two 
of them, previously briefly described in the Enumeratio of 1760. It is 
necessary to accept 1760 as the date of publication of Jacquin’s species 
but to typify them with the data and illustrations of 1763. Three of 
Jacquin’s species, C. alba, C. rosea, and C. flava represent segregates from 
Linnaeus’ C. major. The fourth species, C. venosa, appears to be identical 
with Linnaeus’ C. minor and will be discussed later. In the Selectarum 
in 1763 Jacquin gave for his C. alba the basic polynomial “Clusia foliis 
aveniis” and the reference “Linn. sp. pl. I. p. 509,” as well as the 
reference “Clusia flore albo, fructu coccineo. Plum. gen. 21. ic. 87. | ae 
although he did not use the Linnaean specific epithet. Jacquin’s illustra- 
tion clearly indicates the same plant as in Plumier’s unpublished plate 
(Fic. 1) which was copied (with alterations) for the Burmann edition. 


1962] HOWARD, GUTTIFERAE 391 


Chl five albe 
fue te | 


Fic. 1. Lectotype of Clusia major L. ce 85, oh aquarellé “Clusia flore 
albo, fructu coccineo” from Manuscript No. 6, Plumier, Botan nicum Americanum, 
I. Courtesy of the Muséum National ee ticle Paris. 


392 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


There is no doubt in my mind that Jacquin was renaming Clusia major 
by dividing this composite species and assigning new names to all three 
parts of it. In the second edition of Species Plantarum, also published in 
1763, Linnaeus accepted Jacquin’s divisions of Clusia major, used Jac- 
quin’s specific names, credited him, and cited Jacquin’s publications. By 
this act Clusia major disappeared and was never again used until Britton 
and Millspaugh cited it as the type species of the genus. 

In 1860, Planchon and Triana (Ann. Sci. Nat. IV. 13: 318-376. 1860) 
published a detailed treatment of Clusia in a larger work considering 
the Guttiferae as a whole. They concluded that Clusia major should be 
regarded as a nomen confusum and cited C. major in part under each of 
the three species mentioned “by Jacquin and accepted by Linnaeus. Other 
monographers have followed suit. 

Although Hitchcock and Green (loc. cit.) selected Clusia major as the 
type species of the genus, they qualified this by stating “senus C. rosea 
Jacq.” The Index Nominum Genericorum refers to the Hitchcock and 
Green selection. The qualification is obviously incorrect, for Jacquin’s 
application of the basic references and illustration of C. major is to 
C. alba Jacq. 

The typification of Clusia major must rest on Jacquin’s selection of the 
Linnaean reference to Plumier’s work. The plant in the Linnaean her- 
barium annotated by J. E. Smith as “Clusia alba H. Aublet vix Linnaeus” 
is neither the plant Jacquin described, nor does it correspond to the earlier 
Plumier description. Specimens collected by Plumier exist but none is 
available for the genus Clusia. Planchon and Triana selected as basic 
material an unpublished plate of Plumier (Fic. 1) but noted that the 
Burmann reproduction of this plate contained inaccuracies and excluded 
the flower and the analytical drawing. Thus the choice of a type then is 
between a previously unpublished plate and the illustration of C. alba 
published by Jacquin. I prefer the former, now published in this paper, 
to detract a bit from the long established status of C. alba. 

Clusia major L. (syn. C. alba Jacq.) appears to be characterized by the 
elongate fruits borne on a cymose inflorescence which has a short peduncle. 
The species is represented by recent collections of material of the Lesser 
Antillean islands from St. Eustatius, St. Kitts, Montserrat, and Antigua, 
southward to St. Vincent. Urban’s C. plukenetii is similar in having a 
much elongated peduncle to the cymose inflorescence and a globose fruit. 
Urban cited a collection from Martinique (Duss 1829) which I have 
not seen. I have seen more recent material from Barbados (the type 
locality) and St. Lucia which agrees with Urban’s description. Regret- 
tably, considerable variation is found in the shape of the fruits of Clusia 
major. Although the length of the peduncle appears to be a reliable 
difference between these species, additional field study is necessary to 
determine if two taxa are truly represented. 


1962 | HOWARD, GUTTIFERAE 393 


Clusia flava Jacq. 

There is no confusion in the use of this name. Linnaeus (Sp. Pl. 509. 
1753) included a Sloane reference (Hist. Jam. 2. ¢. 200, f. 1.) in the 
literature cited with the original publication of Clusia major. Jacquin 
(Enum. 34. 1760; Select. 272. 1763) cited the same reference under the 
name C. flava. In the second edition of Species Plantarum (p. 1495. 1763) 
Linnaeus credited the epithet to Jacquin and cited as supporting literature 
references to Jacquin (1763), Sloane, and Browne. Fawcett and Rendle 
(Fl. Jam. 5: 193, 194, 1926) selected as the lectotype the Browne specimen 
in the Linnaean Herbarium, Presumably this is part of sheet 1224.3 and 
is the solitary detached leaf on the right-hand side. 

Fawcett and Rendle cited the distribution of Clusia fzva as Jamaica, 
Barbados, Grand Cayman, and the Florida Keys. The species is indeed 
well represented by recent collections from Jamaica and has been te- 
collected recently by George Proctor on Grand Cayman (Proctor 15141 
[GH]). The occurrence of this species in Barbados is not supported by 
specimens in any herbarium collections I have seen. The reference to its 
occurrence in the Florida Keys is apparently obtained from the writings 
of Nuttall (N. Amer. Sylva 2: 58. pl. 77. 1859) who stated of Clusia flava, 
“This singular and splendid tree is a native of Jamaica, and Cayenne in 
South America, where it is found among rocks on the declivities of moun- 
tains. We have now also to record it as a native of Key West in Florida, 
where it has recently been found, with so many other tropical productions, 
by Dr. Blodgett.” The illustration given by Nuttall is clearly that of 
C. flava; however, there are no supporting herbarium vouchers cited 
and one wonders if the illustration was not made from other herbarium 
material. Blodgett’s collections are preserved in the herbarium of the 
New York Botanical Garden where there are two sheets labelled “Clusia 
flava” collected by Dr. Blodgett. One sheet bears the common name 
“Bull Bay” and is from Pine Key. The other sheet without a common 
name was collected on Key West. Both specimens are sterile; however, 
both have heavier leaf blades than does ¢’. flava and both speciniens, I 
believe, should be referred to C. rosea. In further reference to Nuttall’s 
statement, I have seen no material of C. flava from Cayenne. Clusia flava 
appears to be restricted to Jamaica and Grand Cayman. 

Fawcett and Rendle and authors 4! other modern floras of the Antilles 
do not accept the two varieties of Clusia flava proposed by Planchon and 
Triana (q.v.). 


Clusia rosea Jacq. 

This species was described briefly by Jacquin (Enum. 34. 1760) with 
the supporting citation “Catesh. car. 2, p. 99. #. 99.” In the Selectarum 
(270. 1763) Jacquin supplied the additional references of “Plum. gen. 21.” 
and “Pluk. alm. 92. t. 157. f. 2.” and gave a full description. In the first 
edition of Catesby’s work (1743), the plant is described and illustrated 
with white petals. In the second edition (1754), the illustration shows rose- 


394 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


colored petals. The text refers to the petals as ‘white with rose streaks.” 
Jacquin did not illustrate the species in his Selectarum, but he reported 
it to be from Santo Domingo. In 1926, Fawcett and Rendle selected as 
the “type” the Bahama specimen collected by Catesby and now in the 
herbarium of the British Museum. 

In modern floras, Clusia rosea is reported to occur in Florida, the 
Bahamas, the Greater and Lesser Antilles, Trinidad, Central America, 
and Venezuela. I have not seen all of the specimens cited by various 
authors for this species but a study of many specimens labeled C. rosea 
led to the conclusion that it occurs only in Southern Florida, the Bahamas, 
the Greater Antilles (Cuba, Jamaica, Hispaniola, and Puerto Rico), and 
the Virgin Islands (St. Thomas, St. Martins, St. Jan, and Anguilla). All 
of the material I have seen from Trinidad is best referred to C. palmicida 
L. C. Rich., although I have some doubt about the application of that 
name. The specimens labeled C. rosea from northern South America 
and from Central America do not represent that species as typified by 
the Catesby plant from the Bahamas. 


Clusia plukenetii Urb. 

Both Jacquin (Select. 270. 1763) and Linnaeus (Sp. Pl. ed. 2. 1495. 
1763) cited the polynomial by Plukenet in the references given for 
Clusia rosea, Fawcett and Rendle (FI. Jam. 5: 192. 1926) did the same. 
Urban, in 1908 (Symb. Antill. 5: 432.), described C. plukenetii, gave 
the Plukenet reference and polynomial, and cited specimens from Martin- 
ique, St. Lucia, and Barbados, but did not designate a type. The Plukenet 
reference is to a poor illustration of a plant reported to occur in Barbados. 
It shows a branch with alternate leaves except for two very small leaves 
at the apex of the stem. Mr. George Proctor has informed me that a 
specimen of Clusia credited to Plukenet is in the Sloane Herbarium (Vol. 
95, p. 152, upper right). It consists of only three leaves but is probably 
the holotype of Plukenet’s polynomial and therefore of Urban’s species. 
The common name of ‘Balsam apple” reported by Plukenet is appropriate 
for the genus Clusia, This species has been discussed under C. major. 
It is not comparable to C. rosea, and the name should not be used in the 
synonymy of that species as it has been by many recent authors. 


Clusia minor L. 

The protologue of this species as given by Linnaeus is the following: 
CLUSIA foliis venosis, minor 
Clussa, flore roseo, minor, fructu flavescente, Plum. 

gen. 21. 
Habitat in America meridionali, 4 
Arbor foliis venosis. Racemus florum terminalis. 


bho 


Jacquin in his Enumeratio described Clusia venosa as “C. foliis venosis” 
and cited “Plum. ic. 87. f. 2.” The Plumier references used by Linnaeus 
and Jacquin are comparable, although the latter (Ic. 87. f. 2) expands on 


1962 | HOWARD, GUTTIFERAE 395 


aa Clufia fbr ree money, cei Phra Vales: 


Fic. 2. Lectotype of Clusia minor L. Fic. 88, Dessin a la plume ‘‘Clusia flore 
roseo minor, fructu e viridi rubra” from Manuscript No. 6, Plumier, Botanicum 
Americanum, t. VI. Courtesy of the Muséum National d’Histoire Naturelle, 

aris. 


396 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLII 


the former (Gen. 21) and is accompanied by an illustration. One can 
conclude, therefore, that Jacquin in his Enumeratio supplied an illegiti- 
mate substitute name for C. minor L. In the Selectarum, however, onan 
refers to ‘Clusia (venosa) foliis venosis. Linn. sp. pl. 20 0 DiC DME 
supplements this with a vague description which some subsequent authors 
felt represented a different species. It is clear from Jacquin’s use of 
Clusia venosa in 1760 that this name must be considered a synonym of 
C. minor L. In the second edition of Species Plantarum (p. 1495. 1763) 
Linnaeus accepted the specific epithet ‘“‘venosa” given by Jacquin and 
abandoned his own ‘‘minor,” but did not use Jacquin’s name nor refer 
to Jacquin’s publication as he did for the three segregate species of 
Clusia major. This action seems to indicate that Linnaeus felt C. venosa 
Jacq. was the same as his C. minor. 

Planchon and Triana typified Clusia minor by the unpublished plate 
numbered 88 in the Plumier manuscript. They recognized that this draw- 
ing was not finished in the characteristic manner of Plumier’s other draw- 
ings but stated that with alterations it was comparable to fig. 2 of plate 87 
in the Burmann edition of Plantae Americanae. Planchon and Triana, to 
clarify this species, printed the original, previously unpublished descrip- 
tion and commentary by Plumier and compared these with existing her- 
barium specimens available to them. Either the original Plumier drawing, 
reproduced here as Fic. 2, or the Burmann version with corrections will 
serve to typify the species. Clusia minor L. is clearly defined in modern 
floras. 


Clusia venosa Jacq. (1763 not 1760) 

Planchon and Triana (loc. cit. 369) were troubled by the description 
of Clusia venosa supplied by Jacquin in the Selectarum (273. 1763). In 
their manuscript, they described Clusia mangle, crediting the name to L. 
C. Richard on the basis of a manuscript notation. In a discussion of this 
species they state, “D’apres le nom de Palétuvier de montagne que porte 
a la Martinique le Clusia venosa de Jacquin (non L.) on pourrait croie 
que cette espéce est identique avec celle que nous décriverons ici. Mais la 
description de la plante de Jacquin ne justiferait en aucun point une telle 
détermination.”” Planchon and Triana do not otherwise place C. venosa 
Jacq. (1763). 

In 1893 J. Vesque (DC. Monographiae Phanerogamarum 8: 140, 57. 
1893) listed both C. venosa Jacq. (1763) and C. mangle Rich. ex Planch. 
& Triana. For C. venosa Vesque stated, ‘Il est impossible de classer avec 
certitude la plante visée par Jacquin. C’est probablement une espéce de 
la section Anandrogyne, voisine du Cl. Mangle qui, a ce qu'il parait, porte 
le méme nom vulgaire de ‘palétuvier de montagne’.” Vesque placed as 
supporting literature the polynomial and plate references which Planchon 
and Triana had used in defining C. minor. Vesque did not cite Jacquin’s 
Enumeratio of 1760, but listed only the Selectarum of 1763. There is an 
implication in the work of these men that C. venosa Jacq. as used in 1760 
and defined in 1763 represent two different plants. The description Jacquin 


1962] HOWARD, GUTTIFERAE 397 


used in 1763 does not clearly define either C. minor L. (C. venosa Jacq. 
1760) or Clusia mangle L. C. Rich. ex Planch. & Triana, except as the 
common name applies to the later species. 

Engler (Nat. Pflanzenfam. 21: 201. 1925) considered Clusia mangle 
to be a synonym of Clusia venosa Jacq. This decision can not be accepted. 
Grisebach (Flora Brit. W. Indies 107. 1859) used Clusia venosa Jacq. 
“exclus. syn. Plum.” and the Imray collection he cited is Clusia mangle. 
This treatment, too, is invalid. Clusia mangle L. C. Rich. ex Planch. & 
Triana is typified by a Richard collection from the Soufriére in Guadeloupe. 
The species is known from Martinique, Dominica, and Guadeloupe. It is 
characterized by long peduncled cymes, small globose fruits about 2 cm. 
in diameter and by subsessile or short-petioled leaves. The identity of 
Clusia venosa Jacq. 1763 remains unsolved but the epithet is a later 
homonym of Clusia venosa Jacq. 1760 which is a synonym of Clusia minor 
EP Otis oe 


Clusia grisebachiana (Planchon & Triana) Alain 

Grisebach described Tovomita clusioides for a plant from Cuba 
(T. clusioides Griseb. Mem. Amer. Acad. IT. 8: 166. 1860, not T. clusioides 
[ Choisy ] Cambessédes, 1828). Planchon and Triana recognized the earlier 
homonym and renamed the species in honor of Grisebach. They retained 
the species in the genus Tovomita, but expressed some doubt as to its 
proper assignment (Tovomita (?) grisebachiana Planch. & Triana, Ann. 
Sci. Nat. IV. 14: 284. 1860). Alain correctly transferred the species to 
the genus Clusia (FI. Cub. 3: 314. 1953). 

Urban described Clusia krugiana (Repert. Sp. Nov. 20: 340. 1924) 
from Puerto Rico and C. abbottii (Symb. Antill. 1: 367. 1899) from the 
Dominican Republic. Schmidt indicated on the herbarium labels of several 
Ekman collections that he believed C. abbottii belonged in the synonymy 
of C. Arugiana. In as much as Schmidt’s work was never published, this 
lead was reexamined on the basis of more recent collections and the type 
collections of each species. Only minor differences in leaf size, those partly 
of age, separate these three supposedly endemic species and they should 
be considered as one. 


CALOPHYLLUM CALABA 


A common tree of the Lesser Antilles, often used as a wind break, has 
a widely used common name of ‘‘galba.”’ Regrettably, the scientific name 
used in modern floras is less consistent. Grisebach (Flora Brit. W. Indies 
108. 1859), Urban (Symb. Antill. 8: 438. 1920), and Duss (Ann. Inst. 
Colon. Marseille 13: 103. 1896) use Calophyllum calaba L. Britton & 
Wilson (Sci. Surv. Porto Rico 5: 584. 1924) and Williams (Fl. Trinidad 
& Tobago 1: 62. 1929) use Calophyllum antillanum Britt. Fawcett and 
Rendle (Fl. Jam. 5: 200. 1926) use Calophyllum jacquinii Fawc. & Rend., 
while Moscoso (Cat. Fl. Dom. 378. 1943) and Leon and Alain (FI. Cub. 
3: 309. 1953) use Calophyllum brasiliense Camb. var. antillanum (Britt.) 


398 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


Standl. Recently Furtado (Gard. Bull. Straits Settlements 11: 258-260. 
1941) suggested the typification of Calophyllum calaba L. and his pro- 
posals seem acceptable. 

Linnaeus (Sp. Pl. 514. 1753) proposed the name Calophyllum calaba 
citing four earlier references and referring to the origin of the plant as in 
“Indiis,” thereby implying that the plant occurred in both the Old World 
and the New World. The first generic description is given by Linnaeus in 
1754 (Gen. Pl. ed. 5. 229.) where only the reference to Plumier’s Calaba 
is given. The Plumier reference is not given as such in the first edition of 
Species Plantarum, but is included in the reference Linnaeus gave to his 
Flora Zeylanica (90. 1747). In 1763, Jacquin elaborated on Linnaeus’ 
description (Select. Stirp. Amer. 269. ¢. 165. 1763), citing both the 
works of Plumier and Linnaeus, thereby implying a New World origin. 
In the sixth edition of Genera Plantarum (p. 266. 1764), Linnaeus ac- 
cepted Jacquin’s treatment. The monographers Planchon and Triana 
(Ann. Sci. Nat. IV. 15: 249. 1861), Vesque (DC. Monogr. Phan. 8: 588. 
1893), and Engler (Nat. Pflanzenfam. 21: 196. 1925) restricted Calo- 
phyllum calaba in application to plants of the New World but credit the 
name to Jacquin. 

Hitchcock and Greene (Prop. Brit. Bot. 161. 1929) suggested that 
the species be typified by Jacquin’s interpretation of Linnaeus’ name. 
More recently, Swartz in preparing the Index Nominum Genericorum 
cards cited as the lectotype for Calophyllum L. (Sp. Pl. 513. 1753) “C. 
calaba Linnaeus vide Gen. Pl. ed. 5. 229. 1754; etiam vide M. L. Green, 
Prop. Brit. Bot. 161. 1929).” 

The acceptable synonymy therefore is: 


Calophyllum calaba L. Sp. Pl. 514. 1753; Gen. Pl. ed. 5. 229. 1754; 
Jacquin, Sel. Stirp. Amer. 269. ¢. 165. 1763. 
Calophyllum antillanum Britt. in Britton & Wilson, Sci. Surv. Porto Rico 5: 
584. 
Calophyllum jacquinu Fawc. & Rendl., Fl. Jam. 5: 584. 
Calophyllum brasiliense Camb. var. guido (Britt.) er Trop. Woods 
30: 6. 1932 


Britton had rejected the Jacquin interpretation of Calophyllum calaba 
as a species of the New World and had proposed the name C. antillanum 
for the American elements. Fawcett and Rendle reached the same con- 
clusion, proposing C. jacquinii in apparent unawareness of Britton’s earlier 
piblicatzan. Standley felt that the West Indian plants represented only 
a geographical extension of a Brazilian species and proposed several 
varieties including the combination Calophyllum brasiliense var. antil- 
lanum., the Antillean material is different from that of Central and 
South America at the varietal level, many of Standley’s varieties must be 
transferred to the older specific name Calophyllum calaba L. typified by 
plants in the Antilles. 


1962 | HOWARD, GUTTIFERAE 399 


RHEEDIA LATERIFLORA 


Linnaeus (Sp. Pl. 1193. 1753) based this species on the work of Plumier 
(Gen. 45. 1703). Plumier did not specify the country of origin of the 
plant, but Lamarck (Encyc. 2: 245. 1786) noted the plant to be abundant 
in the Cul-de-sac aux Frégates in Martinique, an area visited by Plumier. 
Urban cites the distribution of the species as Jamaica, Hispaniola, Mont- 
serrat, Guadeloupe, Dominica, Martinique, St. Vincent, and Trinidad 
(Repert Sp. Nov. Beih. 5: 98. 1920). In spite of a study of recent 
collections from Guadeloupe, Dominica, Martinique, Marie Galante, Gre- 
nada, and Jamaica the species remains poorly understood. There have been 
no recent collections from Hispaniola and even the assignment of plants 
from Jamaica to this species is questionable. 

The synonymy of this species is the following: 

Rheedia lateriflora L. Sp. Pl. 1193. 1753. 
Mammea humilis Vahl, Eclog. Amer. 2: 40, ¢. 20. 1798. 
Mammea humilis var. macrophylla (Mart.) Duss, Ann. Inst. Col. Marseille 
13: 102. 1897. 
Mammea humilis var. vahlii Griseb. Fl. Brit. W. Indies 108. 1859. 
Mammea humilis var. plumieri Griseb. Ibid. 


Vahl’s species is based on a Ryan collection from Montserrat. Grise- 
bach’s two varieties are based respectively on the Vahl and Plumier types. 
Grisebach described var. vahlii as shrubby with the leaves pointed at 
both ends. His var. plumieri was a tree with the leaves rounded or sub- 
cordate at the base. The specimens I have seen are variable in leaf shape 
and both types of leaf-bases can be found on one branch. Usually the 
leaves of young plants and of the lateral or axillary branches of older 
plants have the acute leaf bases, 

Most modern workers consider Garcinia macrophylla Mart., the basio- 
nym of Rkeedia macrophylla (Mart.) Planchon & Triana and of Duss’ 
var. macrophylla, to be a distinct species. The specimens Duss cited 
are to be referred to Rheedia lateriflora. 


400 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


DATES OF PUBLICATION OF THE JOURNAL LINNAEA 


ROBERT C. FOSTER 


THE NINETEENTH CENTURY GERMAN periodical Linnaea is of consid- 
erable importance to American taxonomy because of the large numbers 
of New World plants described in it. For some time it has been known 
that certain title-page dates were incorrect. Suspicion was also felt that 
others might well be incorrect. When the Gray Herbarium Card Index 
began to include varietal names from Linnaea, it became imperative to 
establish the dating as accurately as possible. Between my own investi- 
gations and those of six previous workers, it has been possible to do this 
for many of the volumes. It seems desirable to combine these results in one 
publication to obviate the necessity of searching in seven different places. 
The earlier references are as follows: 

BLAKE, S. F., Jour. Wash. Acad. Sci. 46: 192. 1933. 

Kuntze, 0. Rev. Gen. 3(2): 158, 159. 1898. 

Moore, H. E., Gent. Herb. 8: 375. 1954. 

STEENIS- KRUSEMAN, M. J. vAN, Fl. Males. ser. 1. 4: cxevili-cxcix. 1954. 


VOLUME DATE COMMENTS AND SOURCES 
Volume 1. 
TITLE-PAGE DATE: 1826. 
Heft 1: 1-165 P. 65: paper dated oe 1825, 
Heft 2: 166-332 P. 279: ref. to Mart. Nov. Gen. 
fasc. 4, publ. ae Apr. 1826 
P) 
Heft 3: 333-511 After Aug. 1826 P. 468: reviewed Pohl, Pl. Bras. 
Tc. ere 6; this appeared in 
Aug. 1826. 
Heft 4: 512-677 
Volume 2. 
TITLE- oe DATE: 1827. 
Heft 1: 1-144 
Heft 2: ee 
Heft 3: 277-540 After Aug. 1827 ? P. 527: reviewed Lindl. Orch. 


Scelet.; copy by Lindl. to 
Brit. Mus., Aug. 
Heft 4: 541-738 


Volume 3. 
TITLE-PAGE DATE: 1828. 
Heft 1: 1-114 P. 69: paper dated Aug. 1827. 
Heft 2: 115-198 P, 173: paper dated Dec. 1827. 


Heft 3: 199-308 
Heft 4: 309-440 


1962 | FOSTER, THE JOURNAL LINNAEA 401 


VOLUME DATE COMMENTS AND SOURCES 


Volume 4. 
TITLE-PAGE DATE: 1829, 
Heft 1: 1-128 Jan. 1829 Hook. Bot. Misc. 3: 1. 1832. 
Heft 2: 129-288 
Heft 3: 289-450 P. 358: paper dated 16 Mar. 1829. 
Heft 4: 451 to end P. 598: note dated Oct. 1829. 


Volume 5. 

TITLE-PAGE DATE: 1830. 

Heft 1: 1-176 Jan. 1830 

Heft 2: 177-337 Apr. 1830 

Heft 3: 338-496 July, 1830 

Heft 4: 497-688 Oct. 1830 

Heft 5: 689-756 [1830] All dates from Van Steenis-Kruse- 

man. 


Volume 6. 
TITLE-PAGE DATE: 1831. 
Heft 1: 1-208 After Mar. 1831 P. 65: ref. to Mar. 1831. 
Heft 2: 209-384 
Heft 3: 385-544 
Heft 4: 545-736 
Heft 5: 737-796 


Volume 7. 

TITLE-PAGE DATE: 1832. 
Heft 1: 1-144 

Heft 2: 145-272 

Heft 3: 273-400 

Heft 4: 401-560 

Heft 5: 561-688 

Heft 6: 689-778 


Volume 8. 
TITLE-PAGE DATE: 1833. 
112 


ref. to July, 1833. 


8 
Heft 4: 385-512 After July, 1833 Sys 
9: ref. to 27 July 1833. 


P 
Heft 5: 513-624 After 27 July 1833 P. 
Heft 6: 625-684 


39 
52 


Volume 9 
TITLE-PAGE DATE: 1834; 
1835 at bottom of page. 


Heft 1: 1-144 After Jan. 1834 P. 124: letter dated 20 Jan. 1834. 

Heft 2: 145-272 

Heft 3: 273-402 P. 331: counts 216 Annonaceae 
known in 1834. 

Heft 4: 403-514 Early 1835 P. 506: manuscript dated 27 Dec. 
1834. 

Heft 5: 515-642 P. 566: notes dated 7 Feb. 1835. 


Heft 6: 643-758 


402 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


VOLUME DATE COMMENTS AND SOURCES 


Volume 10. 
TITLE-PAGE DATE: 1836; 
1835-1836 at bottom 


of page. 
Heft 1: 1-128 After July, 1835 P. 123: paper dated 1 July 1835. 
Heft 2: 129-224 
Heft 3: 225-368 P. 289: paper dated 25 Oct. 1835. 
Heft 4: 369-464 After Apr. 1836 P. 463: see date of letter. 
Heft 5: 465-608 After Apr. 24, 1836 P. 484: see date of letter. 
Heft 6: 609-758 
Volume 11. 
TITLE-PAGE DATE: 1837. 
Heft 1: 1-128 P. 108: paper dated 16 Oct 1836. 
Heft 2: 129-256 P. 164: paper dated 23 Nov. 1836. 
Heft 3: 257-432 P. 280: paper dated Feb. 1837. 
Heft 4: 433-544 P. 486: letter dated 15 Apr. 1837. 
Heft 5: 545-608 
Heft 6: 609-728 P. 650: paper ey ees 1837. 
At end of volum Pl. Ex- 
sicc. Afr. eae be ee dated 
20 Mar. 1838. 
Volume 12. 
TITLE-PAGE DATE: 1838. 
Heft 1: 1-128 P. 119: paper dated Jan. 1838. 
Heft 2: 129-256 P. 200: paper dated 16 Feb. 1838. 
Heft 3: 257-352 
Heft 4: 353-496 
Heft 5: 497-576 
Heft 6: 577-700 P. 686: ref. to plants collected in 
June, 1838. 
Volume 13. 
Tink sie DATE: 1839. 
Heft 1: 1-112 


Heft 2: os 272 
Heft 3: 273-336 
Heft 4: 337-448 P. 377: paper dated 7 Aug. 1839. 
Heft 5: 449-560 
Heft 6: 561-744 


Volume 14. 
TITLE-PAGE DATE: 1840. 
Heft 1: 1-160 
Heft 2: 161-240 F..1732 te dated 31 Dec. 1839 
(Old Style). 
Heft 3: 241-304 Po2iT a dated 12 Apr. 1840. 


Heft 4: 305-384 
Heft 5: 385-528 
Heft 6: 529-728 Early 1841 P. 706: bears date Feb. 1841. 


1962] FOSTER, THE JOURNAL LINNAEA 403 


VOLUME DATE COMMENTS AND SOURCES 
Volume 15. 
TITLE-PAGE DATE: 1841. 
Heft 1 60 P. 160: paper dated Jan. 1841. 
Heft 2: 161-288 P. 282: paper dated 15 Apr. 1841. 
Heft 3: 289-384 P. 384: paper dated May, 1841. 
Heft 4: 385-480 P. 442: paper dated 1 July, 1841. 
Heft 5: 481-704 P. 703: paper dated Oct. 1841. 
Volume 16. 
TITLE-PAGE DATE: 1842, 
Heft 1: 1-112 P. 112: paper dated + Jan. 1842. 
Heft 2: 113-224 
Heft 3: 225-336 P. 336: note, dated June, 1842. 
Heft 4: 337-400 P. 388: paper dated 9 Aug. 1842. 
Heft 5: 401-512 P. 512: paper dated Sept. 1842. 
Heft 6: 513-592 Noticed, Bot. Zeit. 1: 288. 28 
pr. 
Introduction to volume dated 
March, 1843. 
Volume 17. 
TITLE-PAGE DATE: 1843. 
Heft 1: 1-128 Vol. 16, p. iv, dated Mar. 1843, 
says a Ad o oh de be will soon 
pea 
Heft 2: 129-256 P. 244: ae dated May, 1843. 
Heft 3: 257-384 
Heft 4: 385-512 P. 446: paper cites material col- 
lected 29 Aug. 3. 
Heft 5: 513-640 P. 577: paper dated Dec. 1843. 
Heft 6: 641-764 Early 1844 P, 676: paper dated Jan. 1844. 
Volume 18. 
TITLE-PAGE DATE: 1844. 
Heft 1: 1-112 P. 50: paper dated May, 1844. 
Heft 2: 113-256 lst paper dated Sept. 1844 
Heft 3: 257-384 P. 301: paper a Oct 1844, 
Heft 4: 385-512 P. 505: review dated 1 Dec. 1844. 
Heft 5: 513-640 Mid-1845 P. 626: preface to paper dated 
May, 1845. 
Heft 6: 641-774 Py fin paper read 12 June 1845. 
Volume 19. 
TITLE-PAGE DATE: 1847. 
Heft 1: 1-128 Jan. 1846 Kuntze, Rev. Gen. 3(2): 158. 
1898. 
Heft 2: 129-256 Feb. 1846 Kuntze, loc, cit. 
Heft 3: 257-384 1846 Before 18 Dec. 1846; noticed Bot. 
Zeit. 4: 875, of that date. 
Heft 4: 385-512 Dec. 1846 Kuntze, loc. cit. 
Heft 5: 513-640 Feb. 1847 Kuntze, loc. cit. 


Heft 6: 641-765 Apr. 1847 Kuntze, loc. cit. 


404 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLIII 


VOLUME DATE COMMENTS AND SOURCES 

Volume 20. 

nena — DATE: 1847. 

Heft 1: 1-128 May, 1847 All dates from Van Steenis-Kruse- 

Heit ace June, 1847 man. 

Heft 3: 257-384 July, 1847 

Heft 4: 385-512 Aug. 1847 

Heft 5: 513-640 Oct. 1847 

Heft 6: 641-781 Dec. 1847 
Volume 21. 

TITLE-PAGE DATE: 1848. 

Heft 1; 1-128 

Heft 2: 129-256 P. 202: paper dated Jan. 1848. 


Heft 3: 257-384 
Heft 4: 385-512 


Heft 5: 513-640 Hefte 1-5 noticed, Bot. Zeit. 6: 
870. 15 Dec. 1848. 

Heft 6: 641-780 Noticed, Bot. Zeit. 7: 750. 19 Oct. 
1849. 

Volume 22. 

TITLE-PAGE DATE: 1849, 

Heft 1: 1-128 P. 113: ee ref. to Bot. Zeit. 
10 Nov. 1848. 

Heft 2: 129-256 P. 144: paper dated 20 Dec. 1848. 

Heft 3: 257-384 

Heft 4: 385-512 P. 464: paper dated 24 Mar. 1849. 


Heft 5: 513-640 
Heft 6: 641-768 
Heft 7: 769-898 


Volume 23. 
TITLE-PAGE DATE: 1850. 
Heft 1: 1-128 
Heft 2: 129-256 P. 212: paper dated 30 Mar. 1850. 
Heft 3: 257-384 
Heft 4: 385-512 
Heft 5: 513-640 
Heft 6: 641-770 P. 736: note dated 18 Sept. 1850. 


Volume 24. 
TITLE-PAGE DATE: 1851. 
Heft 1: 1-128 
Heft 2: 129-256 
Heft 3: 257-384 P. 300: paper dated July, 1851. 
Heft 4: 385-512 
Heft 5: 513-640 


Heft 6: 641-804 Early 1852 P. 649: paper dated Jan. 1852. 
Volume 25. 

TITLE-PAGE DATE: 1852. 

Heft 1: 1-128 June, 1852 All i ae Kuntze, Rev. Gen. 

Heft 2: 129-256 Dec. 1852 a2): 1898. 


1962 | FOSTER, THE JOURNAL LINNAEA 405 


VOLUME DATE COMMENTS AND SOURCES 
Heft 3: 257-384 Feb. 1853 
Heft 4: 385-512 Apr. 1853 
Heft 5: 513-640 June, 1853 
Heft 6: 641-772 Dec. 1853 
Volume 26. 
TITLE-PAGE DATE: 1853. 
Heft 1: 1-128 Feb. 1854 All dates from Kuntze, Rev. Gen. 
Heft 2: 129-256 Apr. 1854 3(2): 159. 1898. 
Heft 3: 257-384 Aug. 1854 
Heft 4: 385-512 Feb. 1855 
Heft 5: 513-640 May, 1855 
Heft 6: 641-807 Sept. 1855 
Volume 27. 
TITLE-PAGE DATE: 1854. 
Heft 1: 1-128 Nov. 1855 Engl. Bot. Jahrb. 19: 562. 1894. 
Heft 2: 129-256 Jan. 1856 Ibid. 
Heft 3: 257-384 Jan. 1856 Ibid. 
Heft 4: 385-512 Feb. 1856 Ibid. 
Heft 5: 513-640 [1856] 
Heft 6: 641-799 Aug. 1856 Gent. Herb. 8: 375. 1954. 
Volume 28. 
Tire. -PAGE DATE: 1856. 
Heft 1: 1-128 Aug. 1856 All dates from Blake in Jour. 
Heft 2: 129-256 Sept. 1856 Wash. Acad. Sci. 46: 192. 1933. 
Heft 3: 257-384 Jan. 1857 
Heft 4: 385-512 June, 1857 
Heft 5: 513-640 Aug. 1857 
Heft 6: 641-767 Feb. 1858 
Volume 29. 
TITLE-PAGE DATE: 1857-1858. 
Heft 1: 1-128 Feb. or later, See cover of Vol. 28, Heft 6. 
1858 
Heft 2: 129-256 June, 1858 Engl. Bot. Jahrb. 19: 526. 1894. 
Heft 3: 257-384 Sept. 1858 Engl. Bot. Jahrb. 19: 526. 1894. 
Heft 4: 385-512 Feb. 1859 Gent. Herb. 8: 375. eek 
Heft 5: 513-640 Apr. 1859 Gent. Herb. 8: 375. 
Heft 6: 641-764 June, 1859 Bull. Soc. Bot. Fr. 7. oe 1860. 
Volume 30. 
TITLE-PAGE DATE: 1859-1860. 
Heft 1: 1-128 
Heft 2: 129-256 Before 25 Nov. 1859; cf. Bull. 
Soc. Bot. Fr. 6: 742. 1859 


Heft 3: 257-384 
Heft 4: 385-512 Before 23 Nov. 1860; cf. Bull. 
Soc. Bot. Fr. 7: 867. 1860. 


406 JOURNAL OF THE ARNOLD ARBORETUM 


VOLUME 
Heft 5: 513-640 
Heft 6: 641-779 
Volume 31. 

Tine -PAGE DATE: 
Heft 1: 1-128 
Heft 2: 129-256 
Heft 3: 257-384 
Heft 4: 385-512 
Heft 5: 513-640 


Heft 6: 641-751 


Volume 32. 


TITLE-PAGE DATE: 


Heft 1: 1-128 

Heft 2: 129-256 
Heft 3: 257-384 
Heft 4: 385-512 
Heft 5: 513-640 


Heft 6: 641-801 


Volume 33. 


TITLE-PAGE DATE: 


Heft 1: 1-128 


Heft 2: 129-256 


Heft 3: 257-384 
Heft 4: 385-512 


Heft 5: 513-640 


Heft 6: 641-770 


DATE 


Mar. 1861 


1861-1862. 


1863. 


1864-1865. 


[ VOL. XLII1 
COMMENTS AND SOURCES 


Before 25 Jan. 1861; me Bull. 
Soc. Bot. Fr. 8: 24. 
Engl. Bot. Jahrb. 19: A “1894. 


Before 11 Apr. 1862; cf. Bull. 
Soc. Bot. Fr. 9: 194. 1862. 
Before 11 Apr. 1862; 


Soc. Bot. Fr. 9: 419. 1862. 
Before 13 Mar. 1863; cf. Bull. 
Soc. Bot. Fr. 10: 122. 1863 
Before 13 Mar. 1863; 
Soc. Bot. Fr. 10: 122. 1863. 


Before 13 Nov. 1863; 
Soc. Bot. Fr. 10: 459. 1863. 
Before 11 Dec. 1863; 
c. Bot. Fr. 10: 539, 1863. 
Before 11 Dec. 1863; 
Soc. Bot. Fr. 10: 539. 1863. 
Before 1 Apr. 1864; 
Soc. Bot. Fr. 11: 87. 1864. 
Before 13 May 1864; 
Soc. Bot. Fr. 11: 149. 1864. 
Before 13 Jan. 1865; 
Soc. Bot. Fr. 12: 4. 1865 


Before 31 Aug. 1864; reviewed 
in Flora, Repert., Halfbogen 3- 
4, 31 Aug. 1864. 

Betore 14 no 1864; reviewed 
in Flora, Repert., Halfbogen 5, 
14 Dec 


Before 24 Mar. 1865; cf. Bull. 
4 


Before 20 May 1865; reviewed 
in Flora, ae Halfbogen 10, 
20 May 1865. 

Before Sept.—Oct., 1865; cf. Bull. 
Soc. Bot. Fr. 12: 227. 1865. 


1962] FOSTER, THE JOURNAL LINNAEA 407 


VOLUME DATE 


Volume 34. 
TITLE-PAGE DATE: 1865-1866. 
Heft 1: 1-128 
Heft 2: 129-256 
Heft 3: 257-384 


Heft 4: 385-512 


Heft 5: 513-640 


Heft 6: 641-752 


Volume 35. 
TITLE-PAGE DATE: 1867-1868. 
Heft 1: 1-128 


Heft 2: 129-256 
Heft 3: 257-352 
Heft 4: 353-384 


Heft 5: 385-512 


Heft 6: 513-637 Nov. 1868 
Volume 36. 

TITLE-PAGE DATE: 1869-1870. 

Heft 1: 1-128 Late 1869 

Heft 2: 129-256 Late 1869 

Heft 3: 257-384 Jan. 1870 

Heft 4: 385-512 Apr. 1870 

Heft 5: 513-640 

Heft 6: 641-790 Dec. 1870 
Volume 37. 

TITLE-PAGE DATE: 1871-1872. 

Heft 1: 1-128 [Early 1872] 

Heft 2: 129-256 Apr. 1872 

Heft 3: 257-432 July, 1872 


Heft 4: 433-512 July, 1872 


COMMENTS AND SOURCES 


Before Mar. 1865; cited in DC. 
Prodr. 15(2): 543 1865. 
Before 22 Dec. 1865: cf. Bull. 
Soc. Bot. Fr. 12: 409. 1865. 
Before 22 Dec. 1865; cf. Bull. 
Soc. Bot. Fr. 12: 409. 1865. 
Before 23 Jan. 1866; reviewed in 
Flora, Repert. (1865), Half- 
bogen 3, 23 Jan. 1866. 
Before 11 May 1866; cf. Bull. 
Soc. Bot. Fr. 13: 253. 1866. 


Before 13 Oct. 1867; reviewed in 
Flora, Repert. 13 Oct. 1867, 
Z 


Dena: 

Before 13 Oct. 1867; reviewed in 

Flora, Repert. 13 Oct. 1867, 
7 


Dok 

Before June, 1868; reviewed in 

Flora, Repert. Mid-June, 1868, 
26 


p. 26. 

Before June, 1868; reviewed in 
Flora, Repert. Mid-June, 1868, 
p. 26. 


Date from Van Steenis-Kruseman. 


Se in Flora, Repert. 15, 

Jan. ps9: 

Me in Flora, Repert. 15. 
Jan. 1870, 

Van Steenis-Kruseman. 

Van Steenis-Kruseman. 


Van Steenis-Kruseman. 


Jour. Bot. 10: 220. 1872. 
Jour. Bot. 10: 382. 1872. 
Jour. Bot. 10: 383, 1872. 


408 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 
VOLUME DaTE COMMENTS AND SOURCES 
Heft 5: 513-544 Oct. 1872 Jour. Bot. 11: 61. 1873. 
Heft 6: 545-663 Sept. 1873 Jour. Bot, 12: 125. 1874. 
Volume 38. 
TITLE-E es DATE: 1874. 
Heft 1: 1-144 Dec. Lore Jour. Bot: 122 125. 1874: 
Fett 23 peer Jan. 1874 Jour. Bot. 12: 125. 1874. 


Heft 3: 257-384 
Heft. 42 395=512 
Heft 5: 513-640 
Heft 6: 641-753 


Volume 39. 


TITLE-PAGE DATE: 


Heft 1: 1-128 

Heft 2: 129-224 
Heft 3: 225-256 
Heft 4: 257-352 


Heft 5: 353-448 
Heft 6: 449-526 


Volume 40. 
TITLE-PAGE DATE: 
Heft 1 


Heft 2-3: 97-192 


Heft 4: 193-288 


Heft 5: 289-384 
Heft 6: 385-468 


Volume 41. 
TITLE- or DATE: 
Heft 1: 


Heft 2: 113-192 
: 193-288 
| 289-384 
3852570 
Heft 6: 577-655 


May, 1874 
July, 1874 


1875. 


Feb. 


1875 


June, 1875 
June, 1875 


1876. 


Aug. 
Dec. 


1876 
1876 


ec. 1876 


. 1877 
PLery 
. 1877 
oe eas 

. 1878 


Van Steenis-Kruseman. 

Van Steenis-Kruseman 

sea Bot. Zeit. 333 751. 13 
Nov. 1874 


Jour, Bot, 133253._1875. 

Jour. Bot. 13: 253. 1875. 

Bot. Zeit. 33: 455. 2 July 1875. 

Noticed, Bot. Zeit. 33: 769. 19 
Nov. 1875. 

Noticed, Bot. Zeit. 33: 790. 26 
Nov. 1875. 

Noticed, Bot. Zeit. 34: 64. 28 
Jan. 1876 

Noticed, Bot. Zeit. 34: 352. 2 
June 1876. 

Noticed, Bot. Zeit. 34: 480. 25 
uly 


Svenson, Rhodora 41: 313. 1939. 

Svenson, Rhodora 41: 313. 1939; 
noticed, Bot. Zeit. 35: 16. 5 
Jan. 1877. 


All dates from original covers 
bound in a volume in the Oakes 
Ames Orchid Herbarium Li- 
brary. 


1962 | FOSTER, THE JOURNAL LINNAEA 409 


VOLUME DATE COMMENTS AND SOURCES 

Volume 42. 

TITLE-PAGE DATE: 1878-1879. 

12 Feb. 1878 Jour. Bot. 16: 123. 1878. 
Heft 2: 113-192 Noticed, Bot. Zeit. 36: 318. 17 
May 1878. 

Heft 3: 193-288 1879 Van Steenis-Kruseman. 

Heft 4: 289-384 1879 Van Steenis-Kruseman. 

Heft 5: 385-480 1879 Van Steenis-Kruseman. 

Heft 6: 481-667 Nov. 1879 Van Steenis-Kruseman. 
Volume 43. 

‘TITLE-PAGE DATE: 1880-1881. 

Heft 1: 1-66 Sept. 1880 All dates from Van Steenis-Kruse- 

Heft 2: 67-138 July, 1881 


Heft 3-4: 139=252 Aug. 1881 
Heft 5-6: 253-486 June. 1882 
Heft 7: 487-554 July, 1882 


Gray HERBARIUM, 
HarVARD UNIVERSITY 


410 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLIII 


A MONOGRAPH OF THE GENUS PLATYDESMA (RUTACEAE)! 


BENJAMIN C. STONE * 


THE GENUS PLATYDESMA was described by Horace Mann, Jr. (1866), 
to accommodate an unusual rutaceous species collected by him and William 
Brigham ‘‘on the mountains behind Honolulu” on the island of Oahu 
in the Hawaiian Islands. Mann presented a rather full description of the 
genus and its single species, which he named Platydesma campanulata. 
A few years later (1869) he again wrote of the genus and species, with 
almost unchanged descriptions, but added a few sentences pertaining to 
the circumstances of the discovery, and a fine full-page half-tone plate 
illustrating its habit and flowers. The species was noted by Heinrich 
Wawra (1873) without further description or commentary. In 1888, how- 
ever, Wilhelm Hillebrand, in his great Flora of the Hawaiian IJslands, 

added three species to the genus, two of them, P. cornuta and P. rostrata, 
described as new, and a third to which Hillebrand applied the name Pp. 
auriculaefolia, based on Pelea auriculaefolia A. Gray. As has previously 
been pointed out (Rock 1913, 1918; Stone 1961, 1962), Hillebrand erred 
in considering Gray’s species to be a Platydesma, for it is a true Pelea of 
the typical section. However, the specimens which led Hillebrand to 
include a fourth species in Platydesma are, in fact, members of the genus. 
Rock (1913) named the taxon represented by these specimens Platydesma 
campanulatum var. sessilifolia (sic). This taxon is here considered to be 
of specific rank, agreeing essentially with Hillebrand’s treatment, but 
clearly less distinct from P. spathulatum (the correct name of Mann’s P. 
campanulata) than either P. cornutum or P. rostratum which Hillebrand 
first described. 

"Studies in the Hawaiian Rutaceae, VI. 

* The field work for this revision was accomplished while the writer was Research 
Assistant, Department of Botany, University of Hawaii, Honolulu. Some studies of 


National a Histoire Naturelle: Paris; and Dr. Otto Degener, Waialua, Oahu, Hawaii, 
who ee both herbarium material and fresh collections. I also wish to thank 
Dr. . Smith of the U. S. National Museum for his kind assistance. At the Bishop 
Museum, Honolulu, where a rather large number of collections of the various species 
has been accumulating, I was provided with all necessary facilities for study. I am 
much indebted to Marie C. Neal and to E. H. Bryan, Jr., of the Museum staff, for 

R L. M 


Perry provided generous assistance in my studies of the materials in the herbarium of 
the Arnold Arboretum and of the Gray Herbarium, for which I am most grateful. 


1962 | STONE, THE GENUS PLATYDESMA 411 


The name now applied to this species has a curious history. Material 
collected by Jules Remy was provisionally determined by H. Baillon in 
Paris as Claoxylon insigne sp. nov. (Claoxylon is a genus of Euphorbiaceae, 
the fruits of which [in the Hawaiian species] bear a slight resemblance to 
those of Platydesma.) Remy’s specimen bears only fruit. The name C. 
insigne was never validly published, but was listed (without description 
and only a herbarium reference) by Drake del Castillo (1890). Many 
years later, while revising the Hawaiian members of Claoxylon, E. E. 
Sherff happened to see the Remy collection and, relying all too heavily 
on the authority of Baillon, published the name Claoxvlon Remy Sherff 
sp. nov., with a full deen orion, Recently the real identity of this 
“Claoxylon” became apparent to Dr. Otto Degener, whe received con- 
firmation from the Muséum National d’Histoire Naturelle of Paris that 
the plant was rutaceous. A lifelike drawing of the specimen was prepared 
(see Fic. 4) in which the character of Platvdesma may readily be seen. 
The proper transfer of the name was made in Degener’s Flora Hawaiiensis 
(1960), where the present writer was able to show that this species is 
identical with the one intended by Hillebrand to include his specimens 
from Kohala, Hawaii, to which he had appended the name Platydesma 
auriculaefolia. The species is now called Platydesma Remyi (Sherff) Deg., 
Sherff & Stone. 

Only two other names must be considered in the genus, both published 
by H. Léveillé, whose erratic work, discussed at some length both by Rock 
and Rehder in recent years, scarcely needs further mention. The first is 
Platydesma oahuensis Lévl. (1911), which is simply a synonym of Mann’s 
original species. The second, Platydesma Fauriei Lévl. (1911) is borne 
by a plant not a member of the Rutaceae, but of the Solanaceae, Notho- 
cestrum longifolium A. Gray, as was first pointed out by Rock (1914). 

Recently it was found necessary to take up an older specific epithet for 
Mann’s original species (Stone, 1962), which must now be called Platy- 
desma spathulatum (A. Gray) Stone. The basionym is Melicope spathu- 
lata A. Gray, published a decade earlier than Mann’s species. Gray also 
described Melicope ? grandifolia in the sarne work (1854), which is the 
same species. Photographs of the type specimens were published in a 
preceding paper (Stone, 1962). 


Platydesma H. Mann,’ Proc. Boston Soc. Nat. Hist. 10: 317. 1866; Mem. 
Boston Soc. Nat. Hist. 1: 529. 1869. Hillebrand, Fl. Hawaiian Is. 71. 
1888. Rock, Indig. Trees Hawaiian Is. 241. 1913. Engler, Nat. Pflanzen- 
fam. ed. 2. 19a: 240, fig. 101, A-G. 1931. 


Shrubs or small trees with opposite or subopposite, simple, punctate, 


® Mann attributed feminine gender to the generic name, and was followed in this 
usage by most later authors. The name, however, like several others taken from 
Greek (such as Geniostoma) is of a class which, although ending in the apparently 
Latin feminine -a, takes neuter modifiers. Therefore the specific and eens epithets 
are corrected, where necessary, to neuter gender. 


412 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLIII 


elongate, nonstipulate leaves; wood when freshly broken with a pepsin- 
like odor, somewhat brittle; inflorescences axillary, cymose or subsessile 
and contracted; flowers hermaphrodite, with 4 broad, rounded sepals in 
imbricated pairs; petals 4, white, but slightly imbricated; disk plane, 
obscurely 4—8-lobate; stamens 8, connate into a staminal tube, white, 
only the anthers and a short, broad, deltoid filament (adnate to the con- 
nective) free, the anthers narrow, elongate, subhastate at the base, with 
linear thecae; ovary of 4 carpels, rounded above, adnate to the apex, each 
with 5-8 pendulous ovules on short, broad funiculi; style central, of 4 
connate elements, columnar, the linear, erect, united stigmas somewhat 
thickened at apex; fruit capsular, dry, 4-lobed, the carpels remaining 
connate, loculicidally (sometimes tardily) dehiscent, rounded or horned at 
apex, the endocarp thin, glabrous, finally separable, the seeds subovoid 
or ellipsoid with angles and faces (if any) from mutual pressures, the 
testa black and shining, the wall osseous, the cotyledons of the embryo 
thin, broad, rounded, the hypocotyl very short, the embryo enclosed by 
endosperm. 


Type species: Platydesma campanulatum H. Mann = P. spathulatum 
(A. Gray) Stone. 


DIsTRIBUTION: Endemic to the Hawaiian Islands; four species: Platy- 
desma Spathulatum, widespread in the archipelago, with insular varieties 
and forms; P. cornutum, endemic to Oahu; P. rostratum, endemic to 
Kauai; and P. Remyi, apparently endemic to the Kohala Mountains, 
Hawaii. 


RELATIONSHIPS: Platydesma is placed by Engler in the subfamily 
Rutoideae, tribe Xanthoxyleae, subtribe Choisyinae, as genus 28 between 
Choisya HBK., a Mexican and southwestern American genus, and Dutail- 
/yea Baill., a small genus of two species endemic to New Caledonia. Neither 
of these genera, however, appears to be either very close in relationship 
or similar in superficial or technical characters. Medicosma Hook. f., a 
monotypic Australian genus, is closer in appearance, and possibly in 
affinity, although placed before (as genus 25) the above-mentioned 
genera. It is perhaps more plausible to consider that Platvdesma may 
be ultimately an Old World derivative, but to exclude American affinities 
would be premature. 


EcoLocy: Platydesma spathulatum, and its subtaxa, are both more 
common and more widely distributed in the Hawaiian group than the 
other species. None, however, could be considered dominant or even 
frequent members of the vegetation. The species are to be found chiefly 
in the rainforests at moderate elevations, from roughly 2000 to 5000 feet, 
and also on windswept crests in low, stunted vegetation. Platydesma 
cornutum and P. rostratum usually occur as single plants or in small 
groups, often along streams or gullies under heavy shade, while P. spathu- 
latum may be found under open or closed canopy forests, or exposed on 
slopes, and can apparently tolerate drier situations. In a few localities 


1962 | STONE, THE GENUS PLATYDESMA 413 


(such as around Kokee, Kauai) P. spathulatum may be found so fre- 
quently as to constitute a considerable, though never dominant, component 
of the association. 


Hasit: Platydesma spathulatum is a polymorphic species, but older 
individuals in more protected areas become small trees. In forests such 
as those about Kumuwela, Kauai, trees of five to eight meters in height 
form an understory in a forest containing species of Psychotria, Pelea, 
Bobea, Pleomele, Pritchardia, and other genera. In exposed, windswept 
areas such as open crests along the Koolau Range, Oahu. a form of the 
same species may be found occurring as a small shrub. Both Platydesma 
cornutum and P. rostratum are shrubby, with erect, usually slender stems 
and erect or ascending branches. They can accurately be called ‘‘Schopf- 
baume” with their rather large leaves clustered toward the ends of the 
branches (see Fic. 1). Platydesma Remyi has the habit of P. spathulatum. 


LOCAL NAME AND USE: The Hawaiian name for all species is ““Pilo kea,” 
meaning ‘‘white pilo.” If the Hawaiians had a particular use for the plant, 
it is not presently known. Certain species have recently been the subject 
of chemical investigations by P. J. Scheuer and his associates at the Uni- 
versity of Hawaii. Evidence has been found that some species contain 
appreciable amounts of alkaloids and essential oils. In this connection, it 
is worthy of note that Rock (Indig. Trees 242. 1913) reported that the 
odor of the plant when bruised is like pepsin. The wood, when broken, 
emits a starchy odor. 


SPECIFIC CHARACTERS: Characters of taxonomic value are found chiefly 
in the structure, shape, and size and presence or absence of indument of 
the fruits: the structure of the inflorescence; the shape and size of the 
leaf, especially the petiole; the habit; and, to a lesser extent, the amount 
of pubescence both on leaves and parts of the inflorescence. The other 
indigenous genera of Rutaceae, Fagara (Zanthoxylum) and Pelea, are 
greatly different both in appearance and in technical characters, although 
seedling stages of Pelea and Platydesma could perhaps be confused with 
one another. 


KEY TO THE SPECIES 


1. Carpels in fruit rounded, the capsule subglobose; habit shrubby or usually 
arborescent, small trees with several to many irregularly arranged spreading 
or ascending branches. Section PLATYDESMA. 


2. Petioles conspicuous, 1-6 cm. long; fruits glabrous, roe , Maui, 
Hawaii (probably also Molokai). ................ Les nnn 
2. Petioles obsolete or at most 5 mm. ee oe ae with a more 
or less persistent minute pubescence; Hawai. ............ 2. P. Remyt., 


Carpels in fruit rostrate, cornute, or sharply Peed and somewhat divari- 
cate; habit shrubby, with erect, sparsely branched or simple stems closely 
foliose toward the apices. Section CORNUTIA. 

3. Carpels in fruit conic-pointed or shortly attenuate; leaves elongate, spath- 


— 


414 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


ulate, narrowed at base to a small rounded or aes lamina; Oahu. 
asa, A see eet ge nee ae ae ee P. cornutum. 
Carpels | in ‘fruit tapered into a slender, elongate beak; cee narrow 
oblong or subspathulate, the base broad and abruptly tauneate or obtuse; 
WA Gg yee oo ehh eos AGS ee Ooo and LE 4. P. vastus. 


ios) 


Section PLATYDFSMA 


1. Platydesma spathulatum (A. Gray) B. C. Stone, Madrono 16: 165. 
1962. 


Melicope spathulata A. Gray, Bot. U. _ a Exped. 15: 352. 1854. 

Melicope? grandifolia A. Gray, op. cit. 

Platydesma begga Mann, st a Soc. Nat. Hist. 10: 317. 1866; 

osto . Hist. 1: 529-531. pl. 22. 1869. Hillebrand, A 

Hawaiian Is. a, es “companulata.” Heller, Minn. Bot. Stud. 1: , 
1897. Rock, Indig. Trees Hawaiian Is. 241. 1913 (including f. lee 
Skottsberg, Acta Horti Gothob. 10: 120. 1936; ibid. 15: 388. 1944 

Platydesma campanulata y var. macrophylla Hillebrand, loc. cit. 

Platydesma oahuensis Léveillé, Rep. Sp. Nov. 10: 153. 1911. 


A shrub or small tree to about 12 m., with spreading branches foliose 
toward the ends, trunk straight, to about 20 cm. thick, glabrous through- 
out or with fulvous pubescence on the new leaves and inflorescence (and 
in some varieties the mature leaves pubescent beneath); juvenile branches 
greenish, at last clothed with pale grayish bark; all parts copiously glandu- 
lar, emitting odor of pepsin; wood, when broken, with a starchy odor; 
leaves variable in size, distinctly petiolate, the petiole 10-60 mm. long, 
the blades usually 10-50 cm. long (sometimes less), 5-20 cm. wide (some- 
times less), spathulate, lanceolate, obovate-lanceolate, or sometimes elliptic, 
generally about 2.5 to 4.5 times longer than broad, coriaceous or thickly 
so, sometimes the margins revolute, rounded to obtuse, acute, or even 
slightly acuminate at apex, rounded, obtuse, or acute at base, entire, 
moderately to very dark green above, usually paler beneath, pinnately 
veined, the lateral nerves subopposite, ascending from the dorsally promi- 
nent ventrally sulcate costa, straight for about 34 their length, thereafter 
dividing but not uniting into a distinct marginal vein; inflorescences 
axillary, initiated among the leaves, pedunculate, cymose, usually 3—5- 
flowered; peduncles about as long as the petioles, nodose, with ovate- 
subulate or narrowly deltoid acute bracts; pedicels up to 1 cm. long, 
bracteolate; flowers about 15-20 mm. long, the 4 calyx lobes imbricate 
in pairs, green, glabrous except for the ciliolulate margins, or sparsely to 
densely puberulent with appressed hairs, ovate-orbicular, about 8-10 mm. 
long, minutely carinate apically without, more or less persistent into fruit- 
ing; petals 4, clear somewhat creamy white, somewhat (never widely) 
7. at anthesis, slightly obovate- oblong to ovate-oblong, minutely 
thickened unguiculate within at apex, sometimes obscurely emarginate 
and mucronulate in the notch, about 14-21 mm. long; stamens 8, united 
into a tube, white, the tube 12-13 mm. long, the anthers connivent on 
short deltoid oblong processes adnate to the connective; ovary 4-locular, 


1962 | STONE, THE GENUS PLATYDESMA 415 


cornutum rostratum 


spathulatum 


“1G. 1. Leaves, fruits, and habit sketches of the four species of Platydesma; 
semi-diagrammatic. 


416 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


4-lobed, at first minutely puberulous; style about 9 mm. long, whitish, 
slightly clavate and greenish at apex, with linear stigmas; fruit globose, 
slightly 4-lobed, to 26 mm. in diameter, the carpels adnate to the summit, 
in age slightly separating laterally, at last dehiscent; seeds 5—8 per locule, 
with a shining black testa, attached by short funicles to the glabrous 
endocarp. 


Horotype: United States Exploring Expedition, 1838-1842, island of 
Kauai (us) (Melicope spathulata A. Gray) 


Key TO SUBTAXA OF PLATYDESMA SPATHULATUM 


1. Calyx glabrous or very sparsely puberulent only at extreme base. 
2. Leaves glabrous beneat 

Petioles slender, slonwate, usually 2-6 on long; blades ample, ob- 
long-spathulate or obovate-spathulate, 3-4 times longer than wide, 
mostly rounded at apex; Kauai, Oahu, Hawaii; perhaps also else- 
NPE. cc batdeimdotess aduenenes var. Spathulatum (typical form). 

3. Petioles shorter, somewhat flattened, about 1—2 cm. long. 
4. Leaves usually less se 15 cm. long, spathulate but tapered and 


acute or even acuminate at apex, tapered at hee pale, rigid, the 
margins often rev a the blade usually 3-4 times longer than 
Wide? OA, cece vd yam Cewie te ae ce var. Pe es form b. 


Leaves generally 20-50 cm. long or more, relatively broader (about 
2-3 times oe than wide), generally rounded or obtuse at both 
pet TAAL. 644 20 dential pt-deeo% var. eee form c, 

2. Leaves pubescent te densely villous at least on the midrib, the 
laminar surface sometimes glabrescent; blades ea dark, rather broadly 


“ll 


spathulata: Kauai, Oak. soc oscviega cca nerds ri dian es var. pubescens. 
1. Calyx densely cinereous-puberulent; leaf-blades usually somewhat pubescent 
beneath; Oahu, Maui, Hawaii. ......................005. var. pallidum. 


Platydesma spathulatum var. spathulatum. 
Platydesma campanulatum {. coriaceum Rock, Indig. Trees Hawaiian Is. 
242. 1913 


Glabrous throughout; petioles slender, elongate, usually 2—6 cm. long; 
blades ample, oblong-spathulate or obovate-spathulate, usually 3—4 times 
longer than wide, mostly rounded at apex, often somewhat rounded at 
base. 


Kauai. Hanalei, Faurie 241 (A); Makaweli, Faurie 246 (4); Waimea, Faurie 
245 (A, BISH), Forbes 911.K. (ptsu); K Kokee, Rock, 1956 (prstt) : Kaholua- 
manu, Forbes 328.K. (BISH); Rock 1968, 1976, 5338 (GH), 1978, 5344, 5969 
eee) 6021 (A); Kaunuohua ridge, Kaluapuhi trail, Wichman (cB); Kokee 
mp, Degener 8640 (A); Kokee-Kilohana, aie 981 (BISH, GB); Lehua- 
bea cea Kilohana, Cranwell et al. HBS. 2938 (BIsH, GB); Halemanu, Rock 
2298 (BisH), 2314 (GH); Neal, 1929, 1930 (BIsH), Kusche (GH); Kumuwela, 
Cranwell et al. HBS. 2841 (sisH, K), 2847 (GB), Stone 3360 (BISH, BM, CU, E 
GH, ae _— aia cB); Awa’awapuhi trail, Lane 56-614 (BIsH), Stone 1609 
BISH 0 (BISH, P, US); Kohua ridge, Degeners & Hansen 23919 (BIsH); 
Tao) aed Forbes (Bist). 


— 


1962 | STONE, THE GENUS PLATYDESMA 417 


Oahu. WatIANAE RANGE: Piko trail, Makua, eee forest reserve, Davis, 
1932 (BISH). KOOLAU RANGE: Northen end: r summit 1 mi. se. of Black 
Junction, Kahuku, Degener & Carroll 20554 ae Kabul, Degener 8624 (a); 
Punalu’u and vicinity, Faurie 242 (a), Forbes. 1908, 1909 (B1sH); Rock 459, 
640, 657, 8834 (BisH); MacDaniels 382 (sisH), Hume 76, 106 (BisH); Kali- 
uwa’a, Rock 65 (sisH); Waipio-Waiawa, MacDaniels 16, 53 (BISH); Warawa. 
Anon., 1930 (ex Herb. Bd. Comm. Agric. For. Terr. Hawaii) (s1sH); Kawailoa, 
Bryan 867 (stsH); Kahana, Lyon, 1926 (pisH); Wahiawa-Kahana, Forbes 
2202.0. (BIsH); back of SALES Swezey, 1920 (s1sH); Budd & ner son 1155 
(sisH); Waikane-Schofield trail, St. John 10161 (s1sH); Kipapa gulch trail, 
Cowan 700 (s1sH); Anahulu Grail. Degener 10092 (A). sonthern end: Pauoa, 
Hillebrand (s1sH); Konahuanui, Forbes, 1908 (s1sH), Heller 2373 (cH), Mann 
& Brigham 94 (cu, type of Platydesma pe BISH, GH, K); Mt. 
Olympus, Rock 10225 (BISH, GH), Forbes 16. a. (sisH); Koolau range, with- 
out locality, Hillebrand 242 (K); Rock 134 (GR 

Hawaii. Kohala, Rock, 1957 (BisH), Rock a (cu); Hualalai (?), Rock 
3800 (BIsH); Ola’a aie Forbes 652.H. (sisH); Ten-Miles, Fullaway & Gif- 
ford, 1919 (sis); Glenwood, Faurie 244 (a, BIsH), Rock, 1918 (BisH), 1914 
(piso); Kilauea, Rock 12999 (sisH); “Sandwich Isl,” U. S. Exploring Ex- 
pedition (GH). 


Platydesma spathulatum var. spathulatum, form b. (Fic. 2.) 
Shrubby, with slender branches and small, narrowly spathulate, pale, 
rigid leaves tapered at both ends, the margins often revolute; petiole 
1-2 cm. long. 
This form is the one most commonly found on the summit crests of the 
Koolau Range, Oahu, where the vegetation is low, stunted, and windswept. 


Oahu. KooLtau Rance: Punalu’u, summit of Castle trail, Stone 3551 (BISH) 
(see Fic. 2), 1142 (Bisu), Degener, Park & Nitta 8633 (us); Laie, Malaeka- 
hana trail, Degener et al. 10093 (cu); ridge above Kahana, Skottsberg 1853 

BISH, GB); near summit of Poamoho trail, Degener 27328 (us); Punalu’u, 
Faurie 243 (a, BIsH); Rock 460 (A). 


_~ 


Platydesma spathulatum var. spathulatum, form c. 

Arborescent, with thick, often rugulose branches and ample leaves up 
to 50 cm. long, usually 7-25 cm. wide, obtuse at apex and rounded at 
base, dark green, quite glabrous. 

This form appears to be limited to Kauai, where it occurs in the high 
forested tableland around the rim of Kalalau Valley. 


Kau Honopu, Awa’awapuhi trail, Lane 56-586 (BISH); east rim of 
Kalalau Valley Degeners & Cadenheads 27150 (BISH, GB, K, US). 


Platydesma spathulatum var. pubescens (Skottsberg) B. C. Stone, 
Madrono 16: 165. 1962 


Platydesma campanulata var. pubescens Skottsberg, Acta Horti Gothob. 15: 
388 


Arborescent; petioles distinct, up to 5-6 cm. long; blades ample, rela- 
tively broad, pubescent beneath (on the lamina sometimes glabrescent, but 


418 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


3 
. Ao 
am A 
ao Me) 
ey 
x 


Fic. 2. Habit, with flower and ae of Platydesma spathulatum var. spathu- 
latum form b. (From Punalu’u, Oa 


the costa with a persistent villosity), broadly spathulate to subelliptic, 
dark above (the lower surface paler); calyx glabrous or nearly so 

Horotype: Oahu: Waianae Mountains, slope of Kaala, O. Selling 3710 
(GB). 

Kauai. Wahiawa Mountains, Forbes 231.K. (sisH); Waimea drainage basin 
west side, Forbes 782.K. (p1sH); Halemanu, Rock 2318 (BIsH) ; aes an tral, 
Forbes 1046.K. (BisH); Kaholuamanu, Rock 5339, 10226, (BIsH), 5 
Kumuwela plateau, Stone 3358 (A, BISH, BM, E, GR): Pendle: -Lihue csea 


1962 | STONE, THE GENUS PLATYDESMA 419 
trail, Forbes, 1909 (BIsH); Hanapepe, Faurie 240 (A). Oahu. Waianae Moun- 
tains, slopes of Kaala, Mokuleia, Morley, 1934 (BISH). 


Platydesma spathulatum var. pallidum (Hillebrand) B. C. Stone, 
Madrofio 16: 165. 1962. (Fic. 3.) 


Platydesma campanulata 6 var. pallida Hillebrand, Fl. Hawaiian Is. 71. 1888. 


Similar in most respects to the typical variety, but with densely ciner- 
eous-puberulent calyx lobes (sometimes also the outer surfaces and 


Fic. 3. Habit, flowers, and fruit of Platydesma spathulatum var. pallidum. 
(From Olinda, Maui, coll. Degener; courtesy of Dr. Degener.) 


420 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLIII 


margins copiously ciliolulate), the pubescence extending to the mid- 
region of the petals; leaf blades pubescent beneath; hairs pale or fulvous, 
erect to somewhat spreading, about 0.2-0.4 mm. long, those on the calyx 
(and axes of the inflorescence) appressed, those of the leaves more erect. 


SyNTyPEs: Oahu, Kaala, Hillebrand; Maui, Hamakua, Rev. John Lyd- 
gate, both formerly at Berlin, but destroyed in World War I]. Nrorype: 
pied (Degener 8615) (BIsH), cited below. 


) 


u. WAIANAE Mountains: Hillebrand, 1869 (cH); Peahinaia trail, 
oa a 12771 (a). Kootau Rance: Pupukea, Degener, Park & Nitta 8622 
(A, NY); Poamoho trail, Degener et al. re (A). Maui. Woods near Ukulele, 
hee Olinda, Forbes 205.M. (stsH); Olinda, Rock 8534 (BIsH, GH) : Tapping 
(Degener 8615) (BISH, CU, K); Kaea, Forbes 2552 (BISH); Nahiku, Forbes 
240.M. (isu); below Kula pipeline, Munro 796 (BISH) ; Kipahulu valley, 
ridge on left side, Forbes 1642.M. (Bis); ridge between west and east Wailuai- 
lai, Forbes 2664.M. (stsu); Honomanu, east bank of gulch, Forbes 2658.M. 
(BISH); west trail behind Hoaomani: Forbes 2587.M. (s1sH); Honomanu trail, 
Rock 10228 (BisH), Rock, 1911 (Brsit) : Kailua, Rock, 1908 (stsH); without 
definite locality other than “East Maui,” Rock, 1911 (BrsH) ; without locality, 
Forbes, 1920 (BisH). Hawaii. Kohala, Monekancant Rock 8367 (A, BISH); 
Papaikou near Honomu, Bryan, 1937 (Bist). 


bo 


Platydesma Remyi (Sherff) Deg., Sherff, & Stone, FI. Hawaiiensis, 
fam. 179. 1960. Stone, Adansonia IT. 1: 98, 99, 1961. (Fic, 4.) 


Claoxylon Remyi Sherff, Publ. Field Mus. Bot. 17: 557. 1939. 

ecg insigne H. Baillon ex Drake del Castillo, Tilustr. Fl. Ins. Mar. Pac. 
291. 1892 (nomen nudum). 

Ae see campanulatum var. sessilifolia Rock, Indig. Trees Hawaiian Is. 
243. 1913. 


Platydesma auriculaefolia sensu ar lebrand, Fl. Hawaiian Is. 72. 1888, not 
Pelea auriculaefolia A. Gray, 


A sparingly branched light-wooded shrub or laxly branched small tree 
up to 2 or 3 m. tall, with branches loosely foliose and leaves subdistant, 
opposite; habit similar to that of preceding species; leaves sessile or sub- 
sessile, the short obsolete petiole broad, flattened, up to 5 mm. long 
sparsely hispidulous; blades obovate or broadly spathulate, Bene 14— 
38 cm. long, 4.5--18.5 cm. broad, often about 30 « 12 cm. (2—3 times 
longer than broad), punctate- glandular, entire, rounded or iene obtuse 
to subacute at apex, narrowed and at last rather abruptly rounded, sub- 
truncate, or even subauriculate at base, glabrous above, sparingly puberu- 
lent heneath at least when young, the costa sulcate above and raised 
beneath; inflorescences axillary, 1-3 (—5?)-flowered, on short peduncles and 
axes, the pedicels up to 2 cm. long, the axes nodose with opposed ovate 
bracts 1-2 mm. long; flowers as in Platydesma spathulatum, but th 
calyx lobes puberulent on both faces; capsule about 25 mm. in Pieces 
minutely puberulent, with rounded carpels. 


Horotype: Hawaii. Location unknown, Jules Remy 604, 1851-55 (Pp). 


1962 | STONE, THE GENUS PLATYDESMA 421 


3 


Dbado! « Mure, 


=e 


Fic. 4. Holotype of Platydesma Remyt. eae by Mme. Godot de 
Mauroy, commissioned by O. Degener; courtesy Muséum National d’Histoire 
Naturelle, Paris, and Dr. Degener. Coll. J. Remy 604, Hawaii 1851-55.) 


422 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


Hawaii. Kohala Mountains, Hillebrand 244 (cH, x; this collection is the 
basis of Hillebrand’s Platydesma auriculaefolia) ; upper Hamakua ditch trail, 
above Koiawe valley, Cranwell & Skottsberg HBS. 3192 (BISH, GB); Kohala, 
Forbes 499.H. (sisH); mauka of Waipio valley (inland side), Lyon, 1919 
(Bis); Kohala, Rock 8354 (sis, GH), 4222 (GH, type of var. sessilifolia), 
Rock, 1912 (a). 


This species is very close to Platydesma spathulatum, differing mainly 
in the obsolete petioles, the pubescence of the leaves and calyx (similar to 
that of P. spathulatum var. pallidum) and the puberulence of the fruits. 

The drawing of the holotype given here (Fic. 4) was made at the re- 
quest of Dr. Otto Degener, and has been published previously in Flora 
Hawatiensis. 

A specimen from Kauai (Alakai, MacDaniels 781) may belong here, 
but it is fragmentary and somewhat dubious. The locality would not be 
implausible in relation to the type of habitat, but the wide distributional 
gap would be most unusual. At present it seems apparent that this species 
is endemic to the Kohala region of Hawaii. Because of the number and 
recentness of some collections there seems to be no basis for considering 
it extinct, but new collections would be desirable. It may be possible 
eventually to show that this taxon should better be considered as a sub- 
species of Platydesma spathulatum. 


Section CorNutTIA, sect. nov. 


Carpidii maturitatem divaricati cornuti vel rostrati. 
Type species: Platydesma cornutum Hillebrand. 


3. Platydesma cornutum Hillebrand, Fl. Hawaiian Is. 72. 1888. (Fic. 
a) 


Melicope grandifolia sensu Wawra, Flora 56: 138. 1873, non A. Gray, Bot. 
U.S. Explor. Exped. 15: 354. 1854. 


Erect unbranched or sparsely. branching shrub with soft wood up to 
7m. high, the main stem about 2 cm. diameter, the few branches erect 
or ascending, closely foliose near the ends, the youngest branches green, 
soon clothed with a pale grayish bark, the wood when freshly broken 
having a starchy odor; glabrous throughout except for the leaf buds 
and the proximal parts of the inflorescences, these sparsely hirtellous, but 
glabrate; leaves petiolate, petioles short, broad, flattened, 2-10 mm. long; 
blades glabrous, dark green when mature, linear-spathulate or narrowly 
obovate, 12-40 cm. long, 5-13 cm. broad, commonly about 30 « 9 cm., 
tapering gradually to the base, decurrent or finally obtuse, rounded or 
more commonly bluntly obtuse or acute at apex, the costa sulcate above, 
prominent beneath, the lateral nerves similar, ascending from the costa, 
merging distally with the deeply arched connecting vein. Inflorescences 
axillary, borne among the leaves but maturing on the lower stems after 
the leaves fall, fasciculately cymose, (3—)9—15-flowered, the peduncle and 


1962 | STONE, THE GENUS PLATYDESMA 423 


axes obsolete, pedicels thus subsessile, bractlets and bracteoles at first 
hirtellous, pedicels glabrous or nearly so, 5-10 mm. long, flaring into 
the base of the calyx; sepals glabrous except for the ciliolulate margins, 
broadly rounded, 3-4 mm. long and 5-6 mm. broad, imbricate in pairs, 
the inner pair slightly smaller; petals white, glabrous except for the 
minutely ciliolulate margins, 9-12 mm. long; stamens 8, connate into 
a staminal tube, the anthers on very short deltoid filaments, unequal, 4 
somewhat larger, equalling the corolla, 4 slightly shorter; tube at base 
very slightly connate with base of corolla; ovary 4-lobed. on an obscurely 
8-lobed disk, the styles at first connate, with 4 linear stigmas, later 
separating as the fruit matures; capsule with thin, chartaceous walls, 
glabrous, about 12 mm. long, the carpels united along their mutual faces 
nearly to the apex or free down to the middle, the distal portions divergent, 
each carpel terminating in a sharply acuminate horn-like tip; carpels 
usually 8-seeded; seeds with a shining black crustaceous testa. 


Key TO VARIETIES OF PLATYDESMA CORNUTUM 


Leaves abruptly truncate or obtuse at base, petiole distinct. var. cornutum. 
Leaves tapered and gradually, evenly decurrent at base. ...... var. decurrens. 


Platydesma cornutum var. cornutum. 


Leaves tapering toward the base but at last briefly but abruptly trun- 
cate or obtuse, the petiole distinct. 


Ho.otypre: Oahu. Without definite locality except ‘“Helemano, Wailupe, 
and Pauoa” (Koolau Range), Hillebrand (x; isotype, GH). 


Oahu. Kootau RANCE: Northern end: Pupukea, Skottsberg 1815 (BISH, GB) ; 
Pupukea-Malaekahana, Lyon, 1926 (pisH); south Opaeula gulch, Pa’ala’a, 
St. John 10642 (a, BISH); Pupukea-Kahuku, Degener 8623 (A, cu); Paumalu, 
Hosaka 123 (pts); Punalu’u, Rock 8833 (BisH), Rock and Forbes, 1908 (A), 
Rock, 1908 (GH), Anon., 1908 (Herb. Bd. Agric. For. Terr. Hawaii) (BISH); 
between Kaipapau and Punalu’u, Forbes, 1908 (BisH); Kaluanui, summit and 
beyond, Castle trail, Punalu’u, Webster 1610 (sisH); Wahiawa gulches, Forbes 
1712.0. (s1sH); southern end, Puu Konahuanui, Forbes 1010.0 (sIsH); head of 
Manoa valley, Rock & Shaw, 1912 (BIsH). 


mountains of Oahu. There does appear, however, to 
between the varieties, since var. decurrens, though mostly restricted to the 


of the Koolau Range nearest the Waianae mountains. It is here that 
intermediate forms, if any are to be found, could be expected to occur. 
Platydesma cornutum var. decurrens, var. nov. 

Folia longe spathulata, base attenuata laminis in petiolam decurrens. 


Hoioryvpe: Oahu. Waianae Range, Pu’u Kanehoa, southeastern side, 


424 


L 
O 


JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


eaf, flower, and fruit of Platydesma cornutum var. cornutum. (From 
ahu; courtesy of Dr. Degener.) 


1962 | STONE, THE GENUS PLATYDESMA 425 


on steep moist banks of stream in valley about 200 m. below summit at 
altitude of about 700 m., March 26, 1960, B. C. Stone & G. Pearsall 3263 
(BISH ). 


Oahu. Makaha valley, ridges of Mt. Kaala, Forbes, 1909 (sts); Mokuleia 
valley, Forbes 1822.0. (ptsH); Pahole gulch in Mokuleia, Degener & Hatheway 
20664 (pm); Makaleha valley, Russ, 1929 (sIsH); valley south-east of trail 
to Pu’u Kanehoa, type locality, Stone 3430 (pIsH, US); Mt. Kaala, Degener, 
Park, Topping & Swezey 8631 (A) 


This variety is still another of the many examples of taxa endemic to 
the Waianae mountain range of western Oahu. This range, which is 
geologically much older than the larger Koolau Range. may be about 
the age of the island of Kauai. Several examples are known of taxa re- 
stricted to Kauai and to the Waianae Mountains, as well as species with 
distinct varieties in the Waianae Mountains and in Kauai. The specimens 
cited above are mostly sterile or in flower, but Forbes, 1909, bears an 
immature fruit. There are no apparent differences in floral or fruit charac- 
ters between the two varieties, but the difference in the leaves is both 
obvious and constant. The holotype and Stone 3430 are both from the 
same location, where a small group of these tall, erect, unbranched shrubs 
was growing in a moist, rocky habitat near a stream, in company with 
species of Urera, Touchardia, Cyrtandra, Alectryon, Straussia, Morinda, 
Pelea, and Athyrium. The flowers, like those of the next species, Platy- 
desma rostratum, are borne on the slender trunks and mature at some 
distance below the leaves. The sepals are green, the petals and staminal 
tube clear white. 


4, Platydesma rostratum Hillebrand, Fl. Hawaiian Is. 72. 1888. (Fic. 
6.) 


Erect unbranched or sparsely branching shrub with soft wood, branches 
ascending, juvenile stems green, later clothed in thin, pale, grayish bark, 
trunks to one or two (or more) meters high, 1-2 cm. thick, freshly broken 
wood with a starchy odor; leaves clustered near the ends of the branches, 
opposite, subsessile or with short, flattened petioles about 3-18 mm. long; 
blades linear-oblong or long, narrowly elliptic to subspathulate, ae 
20-40 cm. long, and 4-12 cm. broad at maturity, usually about 33 x 
at base broadly and abruptly truncate or obtuse (rarely Se Sy 
scarcely or not attenuate, at apex bluntly acute to coarsely acuminate (the 
tip bluntly acute) or somewhat rounded, in bud hirtellous but soon glab- 
rate, glabrous at maturity; costa sulcate above, raised beneath, the lateral 
nerves numerous, subopposite or subalternate, almost at right angles to 
the costa, united distally 3-10 mm. from the margin by a deeply and sym- 
metrically arching connecting vein; inflorescences axillary, at first hidden 
among the leaves, maturing below the leaves on defoliate young branches, 
cymose, the cymes 3—-9-flowered (and sometimes fasciculate, two or three 
together), less than 2 cm. long overall, the peduncle stout, 1-3 mm. long 


426 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


and 1—1.3 mm. broad, the axes and pedicels up to 9 mm. long, the pedicels 
with an opposed pair of bractlets near the flaring apex at base of calyx, 
the axes with 1-3 pairs of decussate bractlets, these, with the bracteoles, 
axes, and peduncle, hirtellous when young, soon glabrescent; calyx slightly 
puberulent, the lobes imbricate in pairs, the outer pair larger, each lobe 
about 3 mm. long and 5 mm. wide, the inner pair with each lobe slightly 
smaller; petals clear white, glabrous, oblong, obtusely rounded at apex, 
about 5-nerved, valvate, about 12-13 mm. long and 5 mm. broad; stamens 
united into a staminal tube about 11 mm. long, anthers on brief deltoid 
filaments, these alternating in size, 4 about 2.5 mm. long, 4 about 2 mm. 
long, versatile, the anther affixed in the middle of the dorsal side; ovary 
4-lobed, on a disk about $ mm. broad, the carpels free distally, about 1.8 
mm. high, subconic and shortly attenuate into a brief style; capsule of 
4 carpels, each with a distal rostrum about 8 mm. long, connate nearly 
their whole length but the beaks free; capsule about 4 mm. long, exclud- 
ing beaks; carpels 8-seeded; seeds black with a shining crustaceous testa. 


O mm 5 O mm 5 


Fic. 6. Floral details of Platydesma rostratum. (Three upper figures from 
Skottsberg 3100, Kauai. Petals and stamens from Rock 6081. Original.) 


1962 | STONE, THE GENUS PLATYDESMA 427 


HouotypPe: Kauai, without locality, Anudsen 68 (Berlin, probably 
destroyed). 


Kauai. Waimea, Eee Kalalau trail, Cranwell, Selling & Skottsberg HBS. 
3100 (BISH, GB); uohua ridge, Kaluapuhi trail, Wichman (Skottsberg 
2937) (GB): eee Rock 2000, 1999 (A), 2323, 6081, 6084 (GH); Awa’awa- 
puhi trail, Stone 1599 (BISH, K, us): Lane, 1956 (sts); Pearsall, 1956 (sisH) ; 
Stone 3329 (pisH); Hanalei valley, Forbes 133.K. (BISH); upper Lihue ditch 
trail, MacDaniels 837 (Bis); without locality or date, Rock 17302 (a). 


This species is to be found in the mesic to wet forests of the high plateau 
region of Kauai, where it is well distributed but not common. The leaves 
are usually rather dark green and glossy above, the flowers clear white. 
The small, erect, sparingly branched shrubs form an understory element 
in rain-forest associations. 

The relationship of the species is clearly with Platydesma cornutum, 
but the two are easily distinguished by both vegetative and fruit characters. 
If (as seems probable) evolutionary divergence closely followed the se- 
quence of island formation, this species may antedate P. cornutum; or the 
two may be derived from a common ancestor. The relationship of these 
species to the arborescent, globose-fruited P. spathulatum and P. Remyi 
is obvious, but not particularly close, despite the small size (in number of 
species) of the genus. 


LITERATURE CITED 

DeEGENER, O. Flora Hawaiiensis. Unpaged. Privately printed. Honolulu. 1960-. 

DRAKE DEL CASTILLO, E. Illustrationes florae insularum maris Pacifici. Paris. 
1890-1892 

Gray, A. Botany: ees ae States oe eae oes 
Gade Charles Wilkes... 1: 1- (Cf. pp. 352-354) 1 

HILLEBRAND, W. Flora of the oe Islands. 1-673. eee London, 
and New York. 

LEVEILLE, cs Platydesma oahuensis and P. Fauriei. Repert. Sp. Nov. 10: 153- 
154. = 


MANN, ae Revision of Schiedea and the Hawaiian Rutaceae. Proc. Boston 

Soc. Ne Hist. 10: 317. 1866. 
Peis _Blayesmo, and Brighamia. Mem. Boston Soc. Nat. 

Hist. 1: 529-53 

Rock, J. F. The ae trees of the Hawaiian Islands. 1-518. Privately 
printed. Honolulu. 19 

. Revisio plantarum Hawaiiensium a Léveillé descriptarum. Repert. Sp. 

Nov. 13: 352-361. 1914. 

. Pelea and Platydesma. Bot. Gaz. 65: 261-267. 1918. 

Stone, B. C. Studies in the Hawaiian Rutaceae III]. On the New Caledonian 
species of Pelea and a misunderstood species of Platydesma. Adansonia 1: 
94-99, 


. Taxonomic and nomenclatural notes on feels esma (Hawaii) and a new 
name for a Melicope (Solomon Islands). Madrofio 16: 161-166. 1962. 
Wawra, H. Beitrage z. Flora d. Hawaiischen Inseln. Bae 56: 138. 18 73% 


COLLEGE OF GUAM 
AGANA, GUAM 


428 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XLIII 


THE THYMELAEACEAE 
IN THE SOUTHEASTERN UNITED STATES ! 


Lorin I. NEVLING, JR. 


THYMELAEACEAE A. L. de Jussieu, Gen. Pl. 76. 1789, 
“Thymelaeae,’ nom. cons. 


(DAPHNE FAMILY) 


Erect shrubs [sometimes dwarf or climbing; trees, lianas, rarely herba- 
ceous annuals|; stems generally with a layer of tough cortical fibers, often 
with internal phloem. Leaves alternate |approximate, opposite or irregu- 
larly pseudowhorled|, simple, entire, laminar [to needle-like or scalelike |, 


punctate]; petiolate or sessile, exstipulate. Inflorescences terminal at 
shoot apex, extra-axillary [axillary, or cauliflorous], pedunculate or ses- 
sile, racemose |spicate, umbelliform, or capituliform], simple |or com- 
pound]; bracts and bracteoles present or absent, [sometimes resembling an 
involucre|; pedicel articulated when present. Flowers bisexual [or uni- 
sexual by abortion and the plants polygamo-dioecious or dioecious], 
regular or rarely somewhat irregular. Calyx lobes free or, more generally, 
connate and adnate to the corolla and androecium to form a variously 
shaped calyx tube, open [or imbricate, valvate], 4/6, 5 or 3]-merous, 
often greenish yellow |but sometimes brightly colored, sometimes articu- 
lated at the middle]. Corolla nonpetaloid; petals adnate to the calyx, the 
lobes alternisepalous, free [or connate into a faucal corona, either simple 
or variously divided, of various shapes, or absent, generally inserted near 
the orifice of the calyx tube|. Androecium generally diplostemonous 
[rarely haplo- or very rarely poly- or hemistemonous, in @ flowers re- 
duced to staminodia or absent|. in 1 or 2 whorls, the upper whorl anti- 
sepalous and the lower alternisepalous; anthers fila mented [or sessile], 
longitudinally dehiscent, introrse |extrorse or horseshoe-like|, basifixed 


‘Prepared for a_ biologically sla generic flora of the southeastern United 
States, a joint project of the Arnold Arboretum and the Gray Herbarium made 
ciel through the support of George R. Cooley and the National Science Founda- 

tion, and under the direction of C. E. Wood, Jr., and R. C. Rollins. The scheme 
tstiows that outlined at the beginning Pe the series (Jour. “Arnold Arb. 39: 296-346. 
1958) except that the family description and bibliography are more extensive than 
usual. The area covered in this, as in former treatments, is bounded by and includes 
North Carolina, Tennessee, Arkansas, and Louisiana. Material included in descrip- 
tions which is inapplicable to the species of this area is placed in brackets. References 
which have not been verified are designated by an asterisk. The figure of Dirca was 
drawn by Dorothy H. Marsh under the supervision of R. B. Channell and C. E. 
Wood, Jr 


1962 | NEVLING, THYMELAEACEAE 429 


[or dorsifixed, the connective sometimes thickened or produced beyond 
the pollen sacs]; pollen globose, polyporate. Disc annular [cupular, tubu- 
lar, scalelike, or absent], free [or sometimes adnate to the calyx tube], 
surrounding the base of the gynoecium. Gynoecium syncarpous, | 12-, 8-, 
5-, 2- or| 1(pseudomonomeric)-carpellate, |reduced to a pistillode in ¢ 
flowers|; stigma usually capitate; style 1, [terminal or] eccentric, some- 
times obsolete, [rarely with parastyles|; ovary superior; ovules 1 in each 
locule, mostly pendent from the locule apex, anatropous and with ventral 
funicles [or rarely hemi-anatropous to nearly orthotropous], 2-integu- 
mented, [often carunculate, often with an obturator at the base of the 
style]. Fruit a berry [loculicidal capsule, nut, or drupe, often accom- 
panied by an accrescent calyx]. |Seeds sometimes arillate;| embryo 
Straight, with flat or thickened, narrow or broad cotyledons; endosperm 
present [or absent]. Type Genus: Thymelaea P. Miller, nom. cons. 


Approximately 55 genera, with some 500 species now recognized, all 
tropical or temperate, with the greatest speciation in tropical and sub- 
tropical regions. It is expected that the number of genera will be reduced 
by nearly one-fifth in the near future, and a further reduction could be 
realized by the elimination of a number of genera of “convenience.” A 
single genus composed of two species, only one in our area, is native to 
the United States. Thymelaea Passerina (L.) Coss. & Germ., an annual 
weed, has been reported in Iowa and Nebraska; and Daphne Mezereum 
L. is a more or less common escape from cultivation in the northeastern 
United States. 

The family is divided into four subfamilies, according to Domke’s sys- 
tem which is followed here. Some authors exclude the three genera of sub- 
fam. Gonystyloideae Domke and the three of subfam. Aquilarioideae Gilg, 
tribe Microsemmatideae Domke, treating them as a separate family, Gony- 
stylaceae Gilg, a segregation which seems both unnecessary and unwise, 
contradicting as it does both anatomical and palynological evidence. A 
third subfamily, Gilgiodaphnoideae Domke, is monogeneric. The Thyme- 
laeoideae, to which our genus belongs, contains the bulk of the genera 
and species of the family, although a number of genera still remain un- 
placed in the family system. American genera of uncertain position are 
Goodallia Bentham, Lasiadenia Bentham, and Linodendron Grisebach. 

On the basis of the few and scattered chromosome-number reports, a 
base number of nine is assumed. The majority of the species are diploid; 
the only known exceptions are a triploid Daphne and polyploid series in 
Edgeworthia, Pimelea, and Wikstroemia. Apomixis has been demonstrated 
in one species of Wikstroemia and is suspected in others. 

The family is of little general economic importance except, perhaps, 
in very localized areas. The genus Daphne is of considerable horticultural 
interest and is grown in many temperate regions for its sweetly scented 
flowers. In local areas, almost the world over, the tough extraxylary fibers 
of various species are employed as cordage and in the manufacture of 
paper or “cloth.” The heartwood of some Asian species is highly prized 


430 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


as incense wood. The fruits, which seem to have some poisonous principle, 
are used as a purgative or emetic in primitive areas. Unidentified alka- 
loids are reported from plants of a number of genera. 


REFERENCES: 
a Suaw, H. K. Thymelaeaceae-Gonystyloideae. Jn: C. G. G. J. VAN STEENIS, 
. Males. I. 4: 349-365. 19 

poets A. M. Daphnes. jour, Roy. Hort. Soc. 78: 5-18. 1953. [Section-by- 
section discussion of spp. useful in horticulture. | 

Aymonin, G. Essai d’une monographie écologique du Daphne Cneorum i 
Revue Gén. Bot. 66: 281-328. 1959. [Only detailed ecological study of any 
species in the hee with extensive bibliography. | 

BANDONI, A. J., _A, O’DonELL. La anatomia de la timeleacea ‘‘Ovidia pillo- 
pillo,” planta ae de los Andes patagénicos. Physis 15: 377-385. 1939. 

Burrows, C. J. Studies in Pimelea. I—The breeding system. Trans. Roy. 
Soc. New Zealand 88: 29-45. 1960. 

Cortrst, R. Anatomie de quelques plantes alpines. Chapitre I. Le Daphne 
Blagayana Freyer. Bull. Soc. Bot. Genéve I. 33: 145-157. 1942. | Includes 
interesting grate | 

oi de Vhistométrie dans l’examen de certaines drogues naturelles. 
Pharm. hee Helvet. 18: 361-368. 1943.* [ Bark of 4 spp. of Daphne studied. ] 

CruicKsHANK, R. H. Chromosome numbers in the genus ‘Pimelea.’ Pap. 
Proc. Roy. Soc. Tasmania 87: 13-16. 1953. [Demonstration of polyploid 
series. | 

Domke, W. Untersuchungen itber die systematische und geographische Glieder- 
ung der Thymelaeaceen. Bibliot. Bot. 27(111): 1-151. 1934. [Basic taxo- 
nomic treatment to the generic level. ] 

Facertinp, F. Zytologie und Gametophyten bildung in der Gattung W7k- 
stroemia, Hereditas 26: 23-50. 

Fucus, A. Beitrage zur Embryologie der Thymelaeaceae. Osterr. Bot. Zeitschr. 
87: 1-41. 1938. 

Gitc, E. Thymelaeaceae. Pflanzenfam. III. 6a: 216-245. 1894. 

Goswamt, P. C. Hand-made paper industry in Tawang area of the Kameng 
Frontier Division. Indian Forester 87: 765. 1961. [Paper-making with 
Daphne cannabina. | 

Guerin, P. Recherches sur la structure anatomique de l’ovule et de la graine 
des Thyméléacées. Ann. Jard. Bot. Buitenzorg II. 14: 3-35. 1916. [Broad 
study including representatives of many genera. 

Hamaya, T. A dendrological monograph on the Thymelaeaceae plants of 
Japan. Bull. Tokyo Univ. Forests 50: 45-96. 1955. [Discussion of taxonomic 
snug and taxonomic treatment. | 

endrological studies of the Japanese and some foreign genera of the 
ee Ibid. 55: 1-80. 1959. [Detailed morphological and ana- 


Hernic, K. H. Studies in the floral morphology of the Thymelaeaceae. Am. 
Jour. Bot. 38: 113-132. 1951. [Floral morphology of 34 spp. in 11 genera 
of subfams. Aquilarioideae and Thymelaeoideae. | 

Hiraoka, T. Somatic syndesis in Daphne odora I. The chromosome behavior 
in mitosis. Proc. Japan Acad. 34: 700-705. 1958. 

: ic syndesis in Daphne odora IJ. The chromosome behavior in 

meiosis. /bid. 706-711. [Chromosome behavior in a triploid sp.] 


1962 | NEVLING, THYMELAEACEAE 431 


Hou, D. Thymelaeaceae. Jn: C. G. G. J. VAN STEENIS, Fl. Males. I. 6: 1-48. 
1960. [Floristic treatment, ner ng the Gonystyloideae. ] 

Jones, L. R. Daphne Mezcreum in Vermont. Rhodora 2: 142. 1900. 

LEANDRI, J. Structure particuli¢ére du rhizome d’un Daphne. Bull. Soc. Bot. 
Fr. 75: 243-248. 1928 

. Recherches anatomiques sur les Thyméléacées. Ann. Sci. Nat. Bot. 
X. 12: 125-137. 1930. [Basic anatomical study. ] 

Martin, H. The genus Daphne. Rep. 3™ Int. Rock Gard. Pl. Conf. 125-130. 
1961. [Of horticultural interest. ] 

MEISSNER, C. F. Thymelaeaceae. DC. Prodr. 14: 493-605. 1857. [First im- 
portant taxonomic resumé of the family. ] 

NaKAHiIRA, K. Studies on the breeding of trees of special use. 4. Artificial 
hexaploid plants of mitumata (Edgeworthia papyrifera). (In Japanese; 
English summary.) Jap. Jour. Breed. 7: 112-118. 1957. [See also ibid. 157- 
160. 1958.*] 

Neviinc, L. I., Jr. A revision of the genus Daphnopsis. Ann. Missouri Bot. 
Gard. 46: 257— 358. 1959. [Includes key to New World genera. 

Nye, H. A. A new station for Daphne. Rhodora 25: 45, 46. 1923. [Somerset 
Co., Maine. ] 

Oxura, E., & M. Kono. Cytogenetical studies of Edgeworthia papyrifera Sieb. 
et. Zuce. I. Karyotype analysis de two varieties, Kochi and Shizuoka. Biol. 
Jour. Okayama Univ. 4: 60-66 58. 

Cytogenetical sion of Daphne odora Thunb. based on its 
ary ote. Ibid. 5: 51-56. 1959. 

PoHL, R. W. Thymelaea Passerina, new weed in the United States. Proc. Iowa 
Acad. Sci. 62: 152-154. 1955. 

Riptey, H. N. Garu and Chandan. Jour. Straits Branch Roy. Asiat. Soc. 35: 
73-82. 1901. [Important article on incense wood from Agquilaria ions 
Lam., A. dirta Ridl., and Wikstroemia Candolleana Meissn 

STOIANOV, G. A. orca features of rose daphne anaes Cneorum) bloom- 
ing. (In Russian.) Priroda Leningrad 45: 114, 115. 

Tuopay, D. On the behavior during drought of leaves 2 two Cape species 
of Barna with some notes on their anatomy. Jour. Bot. London 35: 
585-601. 1921 

. The geographical distribution and ecology of Passerina. Ann. Bot. 39: 
175-208. 1925 

VENKATESWARLU, J. Embryological studies in the Thymelaeaceae, II: Daphne 
cannabina Wall. and Wikstroemia canescens Meissn. Jour. Indian Bot. Soc. 
26: 13-39. 1947.* 

YaMaHA, G. Experimentelle zytologische Beitrage. III. Mitteilung. Uber die 
Wirkung einiger Chemikalien auf die Pollenmutterzellen von Daphne odora, 
Thunb. Bot. Mag. Tokyo 41: 181-211. pl. 4. 1927. 

ZAZHURILO, K. Uber die Anatomischen Probleme in der Karpologie der 
Gegenwart. (In Russian; German summary.) Acta Univ. Voroneg. 7: 21-42. 
1935. [Extensive bibliography. ] 


1. Dirca Linnaeus, Sp. Pl. 1: 358. 1753; Gen. Pl. ed. 5. 167. 1754. 


Slow-growing, deciduous shrubs with slender, flexible branches swollen 
at the nodes. Mature plant-parts usually containing simple or compound 
crystals of calcium oxalate. Leaves ovate, broadly elliptic, or obovate, 


432 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLIII 


sparsely sericeous and glabrescent, the petioles calyptrate over the axil- 
lary bud. Flowers appearing before or simultaneously with the leaves, 
borne in few-flowered, nodding racemes from the axils of last years’ leaves, 
the primary and secondary peduncles conspicuous [or nearly obsolete]; 
bud scales woolly. Sepals connate into an infundibuliform or narrowly 
campanulate calyx tube, straight [or strongly bent], yellowish green to 
yellow, glabrous, the lobes connate, the sinuses shallow and becoming ob- 
scure with age (in our species). Petals minute, lobes inserted on the calyx 
tube between the insertion of the filaments. Stamens 8, in 2 closely adja- 
cent whorls; filaments filiform, inserted between the lower and upper 
one-third of the calyx tube, glabrous; anthers exserted even in bud. Disc 
a minute, irregularly lobed annulus surrounding the base of the ovary. 
Gynoecium pseudomonomeric, glabrous; stigma punctate, exserted; style 
filiform, glabrous; ovary 1-locular with a single anatropous ovule. Fruit 
a bilaterally symmetrical berry, yellow or yellowish green to white, be- 
coming reddish or purplish. Seeds with a fleshy outer and hard inner 
seed coat; endosperm scanty; embryo with thick, plano-convex cotyledons. 
Type species: D. palustris L. (Name from Greek mythology, after Dirce, 
second wife of Lycus, who was bound by Amphion and Zethus, sons (by 
Zeus) of Lycus’ first wife, Antiope, to the horns of a wild bull to be killed 
and who then was transformed into the fountain of Dirce near Thebes.) 
— LEATHERWOOD. 


Two species of temperate North America, of obscure affinities, dis- 
tinguished from other American genera by the 4-merous, bisexual flower 
with eccentric style. Dirca occidentalis Gray is restricted to six counties 
in the San Francisco Bay area of California, while D. palustris ranges 
sporadically from New Brunswick, Quebec, and Ontario, southward to 
northwestern Florida and Alabama, and westward to Minnesota, Iowa, 
Missouri, and Oklahoma. The two generally are considered to be closely 
related, and in some instances their distinctness has been questioned. 

Our species is generally restricted to rich woods where considerable 
moisture is available. It is found, however, in a variety of habitats and 
seems to ‘be a facultative calciphile. Although of sporadic occurrence, 
local populations often are extensive. 

There is considerable disagreement in the literature concerning color 
of the fruits, which, being eaten by birds as soon as ripe, are rarely seen 
in the wild. The color is described as reddish or purplish in some studies 
and yellowish or yellow-green in others. The large majority of field obser- 
vations favor the yellow or yellow-green color, with the fruit often turning 
white and then purplish after falling or during the preparation of specimens 
for the herbarium. Further field observations are in order. 

The extreme flexibility of the stems of Dirca (as well as of many other 
Thymelaeaceae), well known to field botanists, is due largely to the slight 
degree of lignification, as determined by chemical tests, of the secondary 
wood. Poor lignification is found sporadically throughout the entire family. 
The wood is very light, with a specific gravity of about 0.41, as compared 


1962 | NEVLING, THYMELAEACEAE 433 


to 0.21 in Leztneria floridana Chapm., the lightest native wood of the 
United States. The low density of the wood in Leitneria is due to exten- 
sive amounts of parenchyma, but in Dirca the bulk of the wood is com- 
posed of fibers. The stems of Dirca, as well as many other Thymelaeaceae, 
have an extensive development of extraxylary fibers which accompanies 
all vascularization. Although these fibers can be considered as an added 
supporting device, they, too, are nonlignified (in Dirca). 


Fic. 1. Dirca. a-1, D. palustris: a, pee twig, X 1; b, flower, X 3; c, 
ae calyx tube opened lengthwise, to show insertion of stamens and minute 
tals ae tween filaments, X 3; d, cena in vertical section, diagrammatic, 

e, fruiting twig, - Ya; f, mature berry, X 2; g, seed, with fleshy outer 
‘coat removed to show bony inner coat, x 2; h, fruit, with single seed in 


xs 
o 
Ow B 


ection — semidiagra a 
ony pa seed coat (hatched), thin layer of endosperm (white) surrounding 
large embryo, ovular trace (broken line), X 3; i, fruit in cross section to show 
seed eee seed coats, endosperm, and cotyledons, xen 


REFERENCES: 


Under family references see Domke (p. 129), Gite (p. 239), HEINIG (1951), 
MEISSNER (p. 527 

ANDERSON, E. Leatherwood (Dirca palustris). Arnold Arb. Bull. Pop. Inf. IV. 
1: 25-27. 1933. 

CHoguETTE, L. Contribution 4 l’étude du Dirca palustris L. ou bois de plomb. 
Trav. Lab. Mat. Méd. Paris 17(4): 1-93. 1926. [Nearly complete anatomical 
ne morelolon cal study plus pharmacology and comprehensive bibliography. ] 

N, J. G. Leatherwood for early ee bloom. Jour. N. Y. Bot. Gard. 50: 
ce 59. 1949. [D. palustris as an ornamental. ] 

FERNALD, M. L. The fruit of Dirca ae Rhodora 45: 117-119. 1943. 


434 JOURNAL OF THE ARNOLD ARBORETUM | VOL, XLIII 


Hom, T. Dirca palustris L. A morphological study. Am. Jour. Sci. 2: 177-182. 
1921. [Anatomy of root, stem, and leaf. | 

Howe Lt, J. T. Plantae concer —JI. Madrofio 2: 11-15. 1930. [Descrip- 
tion of fruit of D. occidentalis, 

. Concerning fruit-color in ae occidentalis. Leafl. West. Bot. 7: 176. 
1954. 

Lecours, J. E. W. Le bois de plomb “Dirca palustris.” Bull. Sci. Pharmacol. 
31: 112-116. 1924.* 

Loew, F. A. Observations on the growth of an injured plant of Dirca palustris. 
Proc. Indiana Acad. Sci. 55: 48, 49. 1946. [Growth of injured plant more 
vigorous than uninjured. | 

Minn, H. E., & B. ForperHase. Notes on western leatherwood, Dirca oc- 
cidentalis Gray. Madrono 3: 117-120. 1935. [Only report of this sp. repro- 
ducing by underground stems. | 

McVaucuH, R. The fruit of the eastern leatherwood. Castanea 6: 83-86. 1941. 
[Best description of fruit; includes literature citations. | 

Suggested pay laeeny of Prunus serotina and other wide-ranging 
phylads in North America. Brittonia 7: 317-346. 1952. |Dzrca, 345; ques- 
tions distinctness of spp.; for follow-up see VOGELMANN. 

SHarp, A. J. Relationships between the floras of California and southeastern 
United States. Contr. Dudley Herb. 4: 59-61. 1951. [Indicates close rela- 
tionship of the 2 spp. of Dirca.| 

Skori¢, V. Mikoriza u nekih Thymelaeacea. (In Croatian; German summary. ) 
Acta Bot. Zagreb. 1: 22-24. 1925. [Discussion of mycorrhiza in Dirca pa- 
lustris and several spp. of Daphne. ] 

VoGELMANN, H. A comparison of Dirca palustris and Dirca occidentalis (Thy- 
melaeaceae). Asa Gray Bull. IT. 2: 77-82. 1953. 


1962] CHANNELL & WOOD, LEITNERIACEAE 435 


THE LEITNERIACEAE 
IN THE SOUTHEASTERN UNITED STATES ! 


R. B. CHANNELL AND C. E. Woop, Jr. 


LEITNERIACEAE Bentham in Bentham & Hooker, 
Gen. Pl. 3: vi, 396. 1880, “Leitnerieae,” nom. cons. 


(CorKwoop FAMILy) 


A monotypic family distinguished by secretory canals in the pith and 
leaves, nonaromatic foliage, erect catkins, single-styled gynoecium with 
a superior, l-locular ovary, single parietal anatropous ovule, and seed 
with a large embryo and thin, fleshy endosperm. 


1. Leitneria Chapman, Fl. So. U.S. 427. 1860. 


Dioecious deciduous shrub or small tree to 6 m. tall, up to 15 cm. in 
diameter: current stems hairy; bark brown; wood very light in weight. 
Leaves alternate, 5-ranked, simple, exstipulate, pubescent; blades 10-15 
cm. long, lanceolate to elliptic-lanceolate, acute, entire, somewhat coria- 
ceous and glossy in age, finally rugose; petioles half-cylindric, ca. 2.5 cm 
long; nodes with 3 traces from 3 gaps. Flowers appearing before the leaves 
in erect, preformed aments with weak axes. Staminate catkins curving 
outward, the axis lax, with 40-50 cymules in the axils of spirally deltoid- 
ovate scales, each cymule of (3—)10—-12(-—15) free stamens (apparently 
representing about 3 flowers); bracteoles and perianth absent; filaments 
short, slightly dilated at the base; anthers oblong, 2-locular at anthesis, 
slightly versatile, nearly extrorse, dehiscing longitudinally; pollen nearly 
globose, smooth, 3—6-colpate. Carpellate catkins stiffly erect, spikelike, 

1Prepared for a generic flora of the southeastern United States, a joint project 
of the Gray Herbarium and the Arnold Arboretum made possible through the support 


was prepared by Dorothy H. Marsh from living plants cultivated at the Arnold 


by Dr. Robert K. Godfrey, of Florida State University. We are further grateful to 
Dr. Wilbur H. Duncan, of the University of Georgia, for information on the occur- 
rence of Leitneria in Georgia, and to Dr. Delzie Demaree, of Hot Springs, Arkansas, 
for data from that state and from Missouri. 


436 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


relatively few flowered; carpellate flowers sessile, solitary in the axils of 
the spirally arranged primary bracts, each with 2 bractlets at the base and 
surrounded by a perianth (involucre) of (3) 4(-8) minute tepals (scales?) 
(2 often somewhat larger than the others); style linear-lanceolate, reddish, 
deciduous, the grooved stigmatic surface facing the bract; ovary superior, 
ovoid, green, 1-locular with a single parietal, pendulous, anatropous, 2- 
integumented ovule on the side toward the bract; embryo sac of the ‘Poly- 
gonum”’ type. Fruit an erect, smooth, oblong-ovoid, somewhat compressed, 
dry drupe (the thin flesh heavily reticulated with vascular bundles), 
chestnut-brown, green when young, with a terminal stylar scar. Seed with 
a thin layer of endosperm and a large, straight embryo. TYPE SPECIES: 
L. floridana Chapm. (Named for Dr. E. T. Leitner, —1838, a German 
naturalist who traveled in Florida and was killed during the Seminole 
War.) — CorKWOOD. 


FIG Leitneria. a-k, L. eee a — shoot, X 4; b, twig with 
staminate catkins, Us t d, s of staminate catkin oe without an 
within, showing stamens, ye €, oe eer ee catkins, ~ carpellate 
catkin, X 3; g, carpellate { flower AeA subtending bract removed, ‘tzmatic sur- 
face toward viewer — note tepals and two lateral bractlets, 4; : oe 
flower in vertical section, ee surface to the right, De ci 
at top, diagrammatic, X 4; i, twig with immature fruits, X 4; j, k, Be te 
fruits, X 1 


1962 | CHANNELL & WOOD, LEITNERIACEAE 437 


A single species, Leitneria floridana, known from muddy, brackish tidal 
shores, river swamps, Swampy prairies, and sloughs ® in scattered localities 
in three widely separated areas: southern Georgia (McIntosh and Dough- 
erty counties) and northern Florida (Clay, Franklin and Levy counties) ; 
southeastern Missouri (Butler, Dunklin, Pemiscot, and Ripley counties) 
and eastern Arkansas (Arkansas, Clay, Craighead, Jackson, Jefferson, and 
Lincoln counties); and southeastern Texas (Brazoria and Chambers 
counties). 

According to the literature, the species is variable in respect to height, 
leaf shape, length of catkins, and fruit size, but no comprehensive study 
of variation has been made, and no formal taxonomic segregates have 
been proposed. 

Individual plants spread clonally, apparently from adventitious buds on 
the shallow root system. Pollination is by wind, and fruits are not set in 
the absence of pollination. Sometimes cultivated as an ornamental oddity 
in appropriate wet situations, the Missouri form is hardy as far north 
as Boston, Massachusetts, and Rochester, New York. The soft, only 
slightly porous wood, the lightest of any North American plant (sp. gr. 
0.21) has sometimes been used locally for floats on fishing nets. 

Generally admitted to be highly specialized (reduced) in structure, 
Leitneria shows few features which assist in determining its relationships. 
It has been associated with Ranales, Rosales, Geraniales, Sapindales, 
Parietales, and Myricales, and in most modern works it is placed in the 
“Amentiferae,” usually as a separate order in a position close to Myricales, 
despite various lines of evidence to the contrary. Comparative morphologi- 
cal, anatomical, and embryological studies indicate that the closest rela- 
tionships may be with the primitive Rosales (including Hamamelidaceae) 
or Geraniales, but more diagnostic evidence from these groups is needed. 
The development of the male gametophyte and fertilization have not been 
studied, and the chromosome number is unknown. 

Vascular and other evidence suggests that the flower of Leitneria is 
derived from a bisexual ancestral form with a perianth of two cycles, at 
least one cycle of stamens, and a 2-carpellate, perhaps apocarpous, gynoe- 
cium. The staminate inflorescence apparently represents a many-flowered 
compound ament with a cymule of three florets in the axil of each primary 
bract. 

The secondary xylem is so advanced in various respects, including the 
simple perforation plates and the alternate pit arrangement on the side 
walls of the vessel elements, that there would be little difficulty in deriv- 
ing it from that of the primitive types of any of the groups with which 

Delzie Demaree has written to Channell (April 1, 1962) of an exceptional 
see of Leitneria in valley farmland on the property of Mr. Ronney G. Mattics 
about 214 miles east of Senath, Dunklin County, Missouri. “The area has been 
cleared of large Taxodium trees. Waste areas and ae rows are covered by Leitneria. 
These plants are a troublesome weed to this farm . I have watched this area 
for over ten years and before any of it was cleared tee of plants were present. 
The soil is sandy loam and fertile.” The plants in the area are three to five feet 
tall and were in abundant flower 


438 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


Leitneria has been associated. The most distinctive anatomical feature 
is the presence of secretory canals in the outer part of the pith which 
extend in association with the vascular bundles throughout the leaves. 
This and the stratified Seta led Solereder to note the similarity to 
Dipterocarpaceae, from which, however, it must be excluded on other 
grounds. (See Metcalfe & Chalk: Heim; Van Tieghem & Le Conte.) 


REFERENCES: 


Asse, E. C., & T. T. Earve. Inflorescence, floral anatomy nee 
of Leitneria floridana. Bull. Torrey Bot. Clu b 67: 173-193. 

BaILLon, H. Castanéacées. Hist. Pl. 6: 217-259. 1875. es 239-241; 
carpellate flowers incorrectly drawn. | 

BENTHAM, G., & J. D. Hooker. Leitnerieae. Gen. Pl. 3: 396, 397. 1880. 

CANDOLLE, C. de. Myricaceae. Prodr. 16(2): 147-155. 1864. [Leitneria, soles 

Dunpar, J. Leitneria floridana. Gard. Chron. II. 47: 228. 1910. [Use 
an ornamental. | 

ENGLER, A. Leitneriaceae. Nat. Pflanzenfam. III. 1: 28, 29. 1894; Nachtrage 
II- IV. 117, 363. 1897. 

Heim, F. Sur le genre: Leitneria Chapm. Assoc. Fr. Avanc. Sci. Cong 
Marseille 1891. 16 pp. 2 pls. Paris. 1892. [Allies Leztneria with ae. 
daceae; carpellate flowers incorrectly illustrated, as in Baillon. | 

echerches sur les Diptérocarpacées. 11 + 186 pp. pls. 1-11. Paris. 

Z ' Leitneria, 175, 176, pls. 10, 11; excluded from Dipterocarpaceae; 
illustrations as above 

Hyermavist, H. Studies on the floral morphology and phylogeny of the Amen- 
tiferae. Bot. Not. Suppl. 2: 1-171. 1948. [Comparative morphology; Leit- 
meria, 71-76, retained as order Leitneriales in Amentiferae nearest Myri- 
caceae. 

MetTcaLFe, C. R., & L. CHALK. Leitneriaceae. Anat. Dicot. 2: 1282, 1283. 1950. 

OLIVER, D. lai floridana, Chapm. Hooker’s Ic. Pl. 11: 33, 34. pl. 1044. 
1867-18 

PASFIELD, - H. Leitneria floridana. Gard. Chron. III. 107: 185. 1940. 

PFEIFFER, W.M. The morphology of Leitneria floridana. Bot. Gaz. 53: 189-203. 
pls. 18-20. 1912. [Includes 2 gametophyte and embryology. | 

SARGENT, C.S. Leitneria. Silva N. Am. 7: 109-112, pl. 330. 1895. 

SOLEREDER, H. Systematische peas der Dicotvledonen. xvi -+ 984 pp. 
Stuttgart. siried | Leitnerieae, 879, 880. | 

TiecHeM, P. van, & H. Le Conte. Structure et affinités du Leztneria. Bull. 
soc. Bot. Fr. 33: 181-184. 1886. [Assigned to Dipterocarpaceae. | 

TRELEASE, W. Leitneria floridana. Missouri Bot. Gard. Rep. 6: 65-90. front., 
pls. 30-44, 1895. [Comprehensive morphological study; wood anatomy. | 


DEPARTMENT OF BIOLOGY, 
VANDERBILT UNIVERSITY 


THE ARNOLD ARBORETUM, 
HARVARD UNIVERSITY 


1962] THE DIRECTOR’S REPORT 439 


THE DIRECTOR’S REPORT 


THE ARNOLD ARBORETUM DURING THE FISCAL YEAR ENDED 
JUNE 30, 1962 


IN RECORDING the activities of the staff of the Arnold Arboretum during 
the past year, the event that stands out most vividly is the completion 
of the Charles Stratton Dana Greenhouses and the associated | ‘open house” 
days for the Friends of the Arnold Arboretum and interested public. The 
planning of this new facility has occupied the staff for several years and 
the actual construction of it for much of the present one. The contractors 
finished their work in March, leaving April and early May for the tasks 
of moving the contents of the old greenhouses and laboratory to the new 
location and of preparing the surrounding grounds. The complete coopera- 
tion and the hard work of the horticultural staff of the Arnold Arboretum, 
best described as dedicated, made it possible to meet our schedule. For 
once, the fickle New England climate cooperated with our plans, produc- 
ing excellent weather for the moving operations and also one of the most 
extended flowering seasons in the recent history of the Arboretum. The 
frequent newspaper notices and radio announcements of the progress of 
the flowering season were also helpful in giving us the largest weekly 
attendance since before the last war. The general response to the pub- 
licity culminated in an historic traffic tie-up on surrounding streets during 
lilac weekend with an estimated 25,000 visitors on the grounds between 
two and three P.M. on Sunday, May 20 


Staff: 


New appointments to the staff meee har year were Dr. Bernice G. 
Schubert and Dr. Wallace R. Ernst. Schubert, formerly with the 
U. S. Department of Agriculture, a te Maryland, joined the staff 
on January Ist as Associate Curator. Dr. Er nst, a recent graduate of 
Stanford University, was eee jointly with the Gray Herbarium to 
work with Dr. Wood on the generic flora of the southeastern United 
States. Two scholars were appointed Mercer Fellows during the year. 
Mr. Don M. A. Jayaweera, Director of the Royal Botanic Gardens, 
Peradeniya, Ceylon, held a Rockefeller Foundation Fellowship during the 
last year and was appointed a Mercer Fellow to complete his work on 
the genus Mussaenda. Dr. Lalit ooo Srivastava, a graduate of the 
University of California at Davis, was appointed a Mercer Fellow to 
work with Dr. I. W. Bailey on ane studies of the cambium and 
secondary phloem of vascular plants. 


440 JOURNAL OF THE ARNOLD ARBORETUM | VOL. XLIII 


The resignations of Dr. Joab L. Thomas and Dr. Burdette L. Wagen- 
knecht were accepted at the beginning of the year. Dr. Thomas accepted 
a position at the University of Alabama and Dr. Wagenknecht one at 
Norwich University. 

It is a pleasure to record the horticultural award of the Colman Medal 
to Dr. Sax by the American Association of Nurserymen in recognition of 
the work which he accomplished at the Arnold Arboretum. Dr. Ernst was 
awarded the George R. Cooley Prize for the best paper presented at the 
annual meeting of the American Association of Plant Taxonomists. This 
paper, entitled “The Familial Status of the Fumariaceae,”’ summarized 
his morphological comparisons of this family with its relatives, the Papa- 
veraceae. Dr. Wyman completed his term as President of the American 
Horticultural Society and was elected to the Board of Directors of the 
Society at its annual meeting. 


Horticulture: 


The completion of the Charles Stratton Dana Greenhouses and the 
transfer of the propagation work to these new quarters summarize in 
large measure the activities of the staff in horticulture during the past 
year. Previous reports have indicated the nature of the physical plant 
proposed, and a full description of these greenhouses was published in 
Arnoldia, volume 22, combined issues 5 and 6. The new greenhouses 
are built on land owned by Harvard University for the Arnold Arboretum 
and adjacent to the city-owned land occupied by the main collections. 
The development consists of four units of construction, a main building 
with three attached greenhouses, a cold storage house built into an earthen 
bank, a free-standing slat house of modern design to house the Larz 
Anderson bonsai collection, and a pipe-frame construction to be covered 
with saran cloth to function as a shade house for nursery stock. In the 
surrounding area, over an acre of ground is devoted to nursery stock; 
additional space is for planned future expansion, appropriate landscaping 
for the buildings, bank plantings of suitable materials, as well as a demon- 
stration area for many varieties of such plants, a collection of genetically 
dwarf plants near the dJonsai collection, and a new location for the 
Arboretum hedge collection. The entire location is fenced and can be 
locked, thus affording for the first time excellent protection to the green- 
house and nursery areas. 

The Dana Greenhouses have a main building, the headhouse, 36 * I11 
feet with full basement and first floor and a smaller second floor with an 
apartment 22 68 feet for a resident guard. Included in the main build- 
ing, in addition to ample areas for the work of the propagation staff, are 
a small conference-lecture room, a laboratory for anatomical or cytological 
work, two walk-in cold rooms for controlled temperature experiments in 
ranges of plus 40° to minus 20° F., and abundant storage space. Three 
greenhouses, each 17 & 51 feet, are attached, and there is space for a 
fourth. Expansion of each is possible on standard modules. The heating 


1962] THE DIRECTOR’S REPORT 441 


Two views of the new lath house for the Japanese bonsai of the Larz 
Anderson Collection of the Arnold Arboretum. The lath house is opposite the 
main building of the Charles Stratton Dana Greenhouses and overlooks the 
edge collection. 


442 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


The Charles Stratton Dana Greenhouses of the Arnold Arboretum. 
“a build; 


AxsovE: Main building and green 
BeLtow: Greenhouses, shade houses, and cold storage house. 


1962 | THE DIRECTOR’S REPORT 443 


plant is designed to handle twice the present glass area, An auxiliary 
generator was installed to provide automatic take-over when the line 
voltage drops below 70 per cent of normal. This generator will operate 
both the heating and refrigeration units for an extended emergency. Ease 
of maintenance and operation were the primary considerations in the de- 
sign of the building and are exemplified by the uniform floor levels 
throughout the building, the ample aisles for trucks, an electric elevator 
for transport of supplies to the basement storage, glazed tile walls for 
cleanliness, chutes to an incinerator or bins for debris. washable floors 
which can be hosed and then dried with ‘‘squeegees,” and soil bins which 
are filled from the outside and are unloaded from the inside. 

The cold storage house is another feature of the experimental oppor- 
tunities made possible through the new construction. This building, 
15 & 100 feet, is of concrete block construction and is insulated with 
slabs of “styrofoam.” The house has heating as well as refrigeration units. 
One section will house the bonsai collection during the winter months, and 
a separate section can be used to produce an early cold season or extend a 
winter season for plants larger in size than are usually cared for in a 
greenhouse. Nursery stock normally subjected in New England to un- 
seasonable early warm periods followed by a late freeze now can be main- 
tained in a dormant condition until all frost danger is over. 

The erection of the new greenhouses, named for Charles Stratton Dana, 
was made possible through a generous bequest by his daughter, Martha 
Dana Mercer. This development has met a long-standing need of the 
Arboretum for modern greenhouses with experimental facilities. Their 
completion and occupancy make possible continued contributions by the 
staff to the study of the ornamental plants hardy in New England. 

To move from the old greenhouse area as quickly and as completely as 
possible required long planning and hard work on the part of the staff. 
During the fall, plants to be moved were planted in cans or were root 
pruned. Accumulations of many years were sorted and discarded or 
packed for the move. Soil was conditioned to receive the transplants. 
New equipment and supplies had to be anticipated and ordered. Finally, 
in March when construction was completed the move took place. During 
May and again during commencement week the greenhouse area was on 
display. New lawns were planted, newly transplanted materials had to be 
watered, mulches were spread, much pruning was required, and, finally, the 
ever-present weeds of newly developed areas required attention. At the 
time of this report operations are about back to normal, and much of the 
work which remains to be done can be fitted into a regular schedule. A 
special word of appreciation is due Dr. Wyman, horticulturist, who worked 
with the architects and contractors throughout the planning and the com- 
pletion of this construction; Mr. Williams, superintendent, for his own 
efforts combined with those of the grounds crew; and Mr. Fordham, 
propagator, and his staff. The good job expected was done. 

The weather of the past year was extremely favorable to the living 
collections, The season was marred only by the passage of hurricane 


444 JOURNAL OF THE ARNOLD ARBORETUM PyOL, seni 


“Esther” on September 21st. Moderate damage to branches resulted 
from gale-force winds, which also destroyed our only specimen of Juglans 
mandshurica in a localized gust. A replacement specimen has since been 
obtained from Finland, but the loss emphasizes the value of the efforts 
of the greenhouse staff to propagate plants now represented in our collec- 
tions by single individuals which have proven extremely difficult to repro- 
duce by the usual propagating techniques. The heavy snow coverage of 
February gave adequate protection to the plants during the month of 
most violent weather, and little or no killing of flower buds or branches 
was experienced. The beneficial result of natural winter protection was 
revealed in a most floriferous spring season of moderate temperatures 
which saw the major collections remain in flower for longer periods 
than usual. 

The labor requirements associated with the new greenhouses caused a 
reduction in the cultural efforts in the main collections during the year. 
Very little planting was done during the fall or spring season, and the 
regular distribution of plants to cooperating nurserymen was omitted 
this year. Materials for both programs are on hand and both will be 
reactivated during the transplanting season in the fall. 

The Department of Parks and Recreation of the City of Boston con- 
tinued its regular attention to the Arboretum road system. A major road 
unit from the Forest Hills gate to the pond area and a second unit from the 
forsythias past the lilac collection to the rockery were resurfaced, the 
drains relocated, and the sidewalks repaired. This is a major improve- 
ment which will facilitate snow plowing during the winter and make many 
areas more accessible for winter-time work. With the cooperation of 
the representatives of the Department of Parks and Recreation, the City 
of Boston scheduled a hearing concerning necessary repairs to a storm 
sewer which passes through the new greenhouse area. Since this land 
belongs to Harvard University and not the city of Boston, a division of 
the costs of repairs has been agreed upon, and it is expected that the 
needed repair will be completed in the next year. 

At the annual meeting of the American Association of Botanical Gardens 
and Arboretums, the Arboretum staff was asked to serve for another two- 
year period as registration authority for cultivated woody plants not 
represented by special societies. Although no additional registration lists 
of cultivars were published during the year, several lists have been com- 
pleted, and work is in progress on others. Dr. Wyman completed the regis- 
tration list for Fagus and Mr. Green that for Ulmus. 

Dr. Howard, with the assistance of Miss Carroll, Miss Herron, and 
Mrs. Walsh, completed the compilation of a directory of botanical gardens 
of the world which is to be published by the International Association for 
Plant Taxonomy as a volume of Regnum Vegetabile. A grant from the 
International Union of Biological oe will assist in the publication of 
this directory which lists the physical characteristics, staff members, and 
the research and resources of over 500 botanical gardens. 

During the past year the staff of the propagation department received 


A WNT | 


The Charles Stratton Dana Greenhouses of the Arnold Arboretum. 
LEFT, ABOVE: Office and conference room; BELOW: Research laboratory. RIGHT, ABOVE and BELow: Apartment 
for greenhouse guard. 


[Z961 


LYOdaa S AOLOAAIG AHL 


446 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLIII 


ee 
8 

| | 
2 


Two views of the work area for plant es in the Charles Stratton 
Dana Greenhouses of the Arnold Arboretu 


164 shipments of plant materials representing 579 species and varieties 
from 23 different countries. One hundred and thirty of these lots (repre- 
senting 458 taxa) came as plants or cuttings, while only 30 shipments (121 
taxa) consisted of seeds and fruits. By contrast, 214 shipments comprising 


1962 | THE DIRECTOR’S REPORT 447 


852 taxa were distributed on request to botanists in nine countries. Of 
these, 157 shipments (645 taxa) were as plants or cuttings and 57 ship- 
ments (207 taxa) as fruits or seeds. In addition to these, we were able to 
fill 52 requests for pollen, leaf samples, soil samples, fruits, or wood speci- 
mens obtained from the living collections in support of research requests 
of scientists in thirteen countries, not including the United States. 

The requirements of the projected plantings around the greenhouses and 
the development of new groups in the Arboretum collection necessitated 
the propagation of 467 taxa. Outside requests for materials from the 
Arboretum collections not available elsewhere numbered 62 items which 
have been propagated for distribution at the proper stage of development. 
The Arboretum staff requested the propagation of 74 taxa for taxonomic, 
cytological, or morphological studies. Finally, 84 ee, were handled by 
the propagation staff to obtain or to check propagation 

Experimental work in the greenhouses, although are by the 

move, continued on problems concerned with winter survival of rooted 
cuttings, methods of rooting species which defy normal procedures, tech- 
niques of handling and breaking seed dormancy, methods of obtaining 
more complete and uniform germination of seeds, and viability studies in 
the storage of scions. The results of these studies appear as notes or more 
comprehensive papers in Arnoldia and other horticultural journals. 


Case Estates: 

The Case Estates in Weston continue to serve as a nursery testing, and 
demonstration area, a quarantine zone, and a place to retain under condi- 
tions of easy maintenance plant material not desired in the main collections 
in Jamaica Plain. Plants propagated in the Arboretum greenhouses are 
held in Weston until they reach flowering size. During this period their 
growth habits, hardiness, and flowering characteristics can be determined. 
The best plants from the cultural or ornamental points of view later are 
planted in Jamaica Plain. Less desirable plants, the distribution of which 
may be restricted by law (e.g., Ribes or Berberis), or plants marginally 
hardy may be retained in permanent non-display plantings in Weston. Of 
the demonstration areas, the perennial garden, the ground cover plots, and 
the small stature trees attract the greatest attention from visitors. In- 
creased interest is seen in the test plantings of Narcissus varieties contrib- 
uted by Dr. Helen Scorgie and other members of the New England Section 
of the American Daffodil Society. Plants received from abroad, subject to 
plant quarantine restrictions, are maintained in special sections and 
screened houses on the Case Estates prior to their clearance for distribution 
by representatives of the Department of Agriculture. In addition, certain 
areas of the various plantings on the Case Estates can be used experi- 
mentally for trial of new horticultural practices. For example, the search 
continues for safe, yet effective, chemical weed killers for use in display 
nurseries, One of the most promising materials tried during the past year 
was “Simazine” which, when applied in the fall, made hand hoeing un- 
necessary in nursery rows until mid-June. 


448 JOURNAL OF THE ARNOLD ARBORETUM (VOL: REN 


f wet aoe 
REA 


: | oo oo . 


The Arnold Arboretum display of dwarf conifers at the Spring Flower Show 
of the Massachusetts Horticultural Society, Revere, Massachusetts, April 17-25, 
1962 


The grounds of the Case Estates are used for teaching activities of the 
staff. In addition to an “open house,” field classes have been held for the 
general public in the spring and the fall. Special tours are arranged for 
interested groups which can be shown certain plants, plantings, and prac- 
tices not demonstrable in Jamaica Plain, and the grounds are used for field 
work in biology classes of Harvard University and the Weston Public 
Schools. In addition, staff members of the Bussey Institution, the Depart- 
ment of Biology, the abot Foundation, and the Gray Herbarium, of 

rvard University, have been allowed to use small plots of land for 
eae studies. Currently, three high-school and_private-school 
students have “science fair” projects under way on the grounds. 


Herbarium: 


During the year, 16,467 specimens were mounted and added to the 
herbarium, bringing the total collection to 742,811 specimens on June 30, 
1962. During the same period, 10,920 specimens were received as acces- 
sions. Of these 9292 were in exchange, 1277 through subsidy, and the 
emainder as gifts or for identification. In conformity with the joint 
policy of having the Gray Herbarium maintain all exchanges with coun- 


1962 | THE DIRECTOR'S REPORT 449 


tries and institutions of the New World and the Arnold Arboretum those 
of the Old, the above accessions represent plants of the Eastern Hemi- 
sphere. All collections of cultivated plants are credited to the Arnold 
Arboretum and, whatever their source, are added to the horticultural 
herbarium in Jamaica Plain. Only 438 specimens were sent out as ex- 
change during the year, although many collections are being prepared for 
exchange in the near future. 

The staff filled 117 requests for loans of herbarium material, amounting 
to 12,056 specimens sent to 69 institutions — 48 in the United States and 
21 to other countries. For their study, the staff requested 94 loans com- 
prising 7278 specimens from 20 American herbaria and 22 foreign institu- 
tions. Outgoing loans averaged 103 specimens per loan and included 
materials from the Arnold Arboretum and the Gray Herbarium. Incoming 
loans averaged 80 specimens per loan, again emphasizing the wealth of 
material in our herbaria. It is of interest to note that of the outgoing loans, 
29 per cent, representing 38 per cent of the specimens sent, were for the 
use of advanced students, the remainder for professional taxonomists. 

Forty-three steel herbarium cases were purchased from the Art Metal 
Company for installation in the Administration Building in Jamaica Plain. 
Two additional cases were purchased for staff use in Cambridge. After the 
installation of the new cases, the entire horticultural herbarium was shifted 
to allow room for expansion throughout the collection and to provide case 
space for individual staff members and for the use of the mounters. It is 
eratifying that there is a gradual increase in the number of specimens of 
cultivated plants being sent for identification and in exchange. These 
have been received from many individuals in the United States and offer 
more exact evidence of the distribution of plants under cultivation. Our 
colleagues in foreign countries are also cooperating in response to our re- 
quest for specimens from cultivation in addition to those from the wild. 

Although the publications cited in the bibliography speak for the scien- 
tific achievements of the taxonomists, it is also desirable to record work 
in progress: Mr. Green, studies in the Oleaceae, particularly Notelaea in 
New Caledonia, Australia, and New Zealand; Dr. Howard, studies on the 
anatomy of the petiole of the dicotyledons and floristic studies of the West 
Indies, particularly in the Guttiferae and Leguminosae; Dr. Hu, studies 
of the Compositae of China, as well as the Commelinaceae and Juncaceae 
of the same region; Mr. Jayaweera, studies on Asiatic Mussaendae and 
the orchids of Ceylon; Dr. Kobuski, the Theaceae of Asia, particularly the 
genus Ternstroemia; Dr. Nevling, studies of the Thymelaeaceae; Dr. 
Perry, with the assistance of Mrs. Metzger, studies of the medicinal plants 
of Southeast Asia; Dr. Schubert, the genus Desmodium in tropical East 
Africa and in Panama, as well as studies toward a monograph of the 
American species of Dioscorea; and Drs, Wood, Brizicky, and Ernst, 
studies of families and genera of seed plants in the southeastern United 
States. During the year Mrs. Metzger visited libraries in England and 
Germany, while Dr. Perry consulted libraries in New York and Washing- 
ton to lend completeness to many of the medical references being re- 


450 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLIII 


viewed by them. Mr. Jayaweera completed revisions of the rubiaceous 
genus Mussaenda in India and Ceylon and in the Philippine Islands. 

In addition to many visitors who studied in the herbarium during short 
visits, we have had two scholars working for longer periods of time. Dr 
Shun Ching Lee, Professor of Botany, National University of Taiwan, is a 
Fulbright Fellow working on a revision of his book on the forests of China. 
Lieutenant Robert Bird, U. S. Army, undertook special studies of the 
vegetation of several countries of southeastern Asia, using the library and 
herbarium. 


Members of the botanical field trip after the 10th Pacific Science Congress 
at the summit of Haleakala, Maui, Hawaii. 


Library: 


The librarians continued their regular services during the year, since 
the shifting of books described in previous reports has been completed. 
Three hundred forty-seven volumes obtained by purchase, gift, and bind- 
ing were added to the library, making the total number of volumes 51,453 
on June 30, 1962. A total of 624 pamphlets was also catalogued and added 
to the collection, making a total of 18,926. The work of cross-indexing the 
main catalogue continues, with 2088 such cards being added, neat 
the new acquisitions. Four issues, totalling 3000 cards, were added to 
Gray Herbarium Card Index of American Plants. The Torrey index c 
American Botanical Literature was enriched with the addition of 2600 


1962] THE DIRECTOR’S REPORT 451 


cards, and issue number 15 was added to the Index Nominum Genericorum. 

In response to requests, sixty-six volumes were sent on interlibrary loan. 
This represents about one half the number of volumes lent in previous 
years. The staff is filling a larger number of requests by the use of “con- 
tura,” “xerox,” or microfilm reproduction methods to avoid lending old 
volumes, It was necessary to request only eight volumes from outside 
sources to meet the research needs of the staff, so extensive are the libraries 
of the Arboretum and the Gray Herbarium and of the neighboring depart- 
ments of Harvard University. 

The librarian, Mrs. Schwarten, along with Dr. Howard, attended the 
dedication of the Rachel McMasters Miller Hunt Botanical Library at the 
Carnegie Institute of Technology, Pittsburgh, Pennsylvania. 


Comparative Morphology: 


Irving W. Bailey, Professor of Plant Anatomy, Emeritus, has continued 
to serve as curator of the wood collection. During the year, Professor 
Bailey continued his research on the leaf-bearing cacti of the genera 
Pereskia, Pereskiopsis, and Quiabentia. Additional preserved specimens 
were received from tropical America and prepared for anatomical studies 
of xylem and phloem. As a Mercer Fellow, Dr. Lalit M. Srivastava has 
worked with Professor Bailey in studying the cambium and phloem of 
these genera. Dr. Srivastava completed his doctoral dissertation on the 
secondary phloem in the Abietineae, and the manuscript has been sub- 
mitted to the University of California Press for publication. At present, 
he is continuing his studies involving ontogenetic and histochemical inves- 
tigations of the vascular cambium and its derivatives on other genera 
within the collections of the Arboretum. 

Since the wood collection of the Arnold Arboretum is one of the best in 
existence, frequent requests, which are filled as materials are available, 
are received for study samples. During the past year, wood samples were 
sent to 26 investigators in 14 countries. Where specimens have not been 
sectioned previously for our own slide collection duplicate slides are re- 
quested in return, adding to the available slides for local study. The wood 
samples supplied on request are acknowledged in published papers, for in 
many cases supporting herbarium vouchers are preserved in the Arboretum 
herbarium. 


Education: 


No formal classes were offered by members of the staff during the past 
year. Informal classes on horticultural topics were represented by the field 
classes held in the fall and the spring at Weston and Jamaica Plain. The 
staff members also took part in two seminar series held weekly and bi- 
weekly in Cambridge and open to all students. The weekly series dealt 
with botanical problems in Latin America, while the biweekly one com- 
prised a discussion of the research projects of staff members and students 
of the Arboretum and Gray Herbarium. A series of lectures open to the 


452 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


public was given evenings during the fall at the administration building 
in Jamaica Plain. Attendance varied at these meetings depending on the 
topic and the weather, It seems unlikely that evening programs in Jamaica 
Plain will ever be completely successful due to the remote location and 
difficulty of public transportation. 

We again experienced an increase in the number of groups visiting the 
Arboretum and the Case Estates and requesting guided tours. There was 
an unexpected increase in the number of requests by mail for information 
on the Arboretum and in individual requests for information on or the 
location of specific plants in the Arboretum. These can be explained only 
in terms of the recent increased publicity given the Arboretum in news- 
papers, national magazines, and on local radio programs, The Arboretum’s 
exhibit at the Massachusetts Horticultural] Society’s Spring Flower Show 
was mentioned and complimented in the March 30th issue of Time, which 
drew attention to the dwarf plants. Life of May 4th listed the Arnold 
Arboretum first in a column on American arboreta and referred to our 
introduction of Metasequoia. The Harvard Alumni Bulletin of February 
17th had a cover illustration of the Arboretum, drawing attention to the 
conifer collection. The New Vork Times of April 22nd published an article 
by Dr. Walter Hodge on botanic gardens which contained an illustration 
of our lilac collection and comment that “Harvard University’s Arnold 
Arboretum, America’s best known arboretum. has been rated since Ernest 
Wilson’s day as ‘America’s greatest garden’.” Dr. Wyman’s timely articles 
in the Boston Herald feature the plants in flower at the Arnold Arboretum 
and the flowering calendar in the Sunday edition of the New Vork Times 
lists the plants in bloom at the Arnold Arboretum each week during the 
spring season. Such publicity is welcome for the interest it focuses on the 
contributions of the staff and the educational values of the living collections. 

The increase in requests for Arboretum staff members as speakers for 
individual garden club meetings now poses a real problem. Whenever 
possible requests are filled, but largely at the discretion of the staff member 
invited as speaker. Joint meetings of garden clubs are one way of utilizing 
speakers more efficiently. In order to compensate in some measure for the 
loss of working or research time. it is necessary to charge a standard fee 
for speaking engagements of staff members. Such receipts are used to 
further the work of horticultural education. 

Dr. Howard appeared on the science lecture series of the Royal Canadian 
Institute in Toronto and that of the American Association for the Advance- 
ment of Science at Franklin and Marshall College. He gave the evening 
address on Hawaiian botany at the 16th Congress of the American Horti- 
cultural Society and at the annual meeting of the Massachusetts Dietetic 
Association spoke on the economic uses of plants. Following the Pacific Sci- 
ence Congress, Dr. Howard was invited to address an open meeting of the 
Garden Club of Honolulu. Dr. Wyman addressed meetings of nurserymen 
in Iowa and Michigan. He took part in short courses in horticulture 
sponsored by the Oregon State University and the University of Massa- 
chusetts. Dr. Wyman also appeared on the lecture program of Longwood 


1962 | THE DIRECTOR’S REPORT 453 


Gardens. Mr. Green described the work of the Arboretum at Pine Manor 
Junior College and reported on methods of vegetation mapping for the 
New England Botanical Club. He also talked about the plant introduction 
and distribution program at the annual meeting of the American Associa- 
tion of Botanical Gardens and Arboretums. Mr. Fordham discussed dwarf 


Strollers in the Arnold Arboretum on lilac weekend, May 20, 21, 1962. 


454 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


and abnormal conifers at the nurserymen’s short course at the Waltham 
Field Station and spoke on methods of accelerating seed germination at a 
meeting of the New England Nurserymen’s Association. Mr. Heman 
Howard is aiding the development of Bartlett Park in Chelmsford as a 
local arboretum and spoke about the selection of plants to its supporters, 
Dr. Wyman and Mr. Williams conducted a day-long demonstration lecture 
on pruning methods for the New England Electric Public Utility Services 
which was attended by over forty line superintendents involved in main- 
tenance of electric lines. 

Radio programs often involving telephone interviews and audience 
questions have included several members of the staff. Dr. Wyman and 
Mr. Williams have also appeared on local television stations. The majority 
of these programs take place in the spring, when interest in gardening 
practices is highest. 


Exhibits and Displays: 


The living collections of the Arnold Arboretum are planted on 400 acres 
in Jamaica Plain and Weston. Only the professional horticulturists or the 
most determined visitors take time to see all of the plants of a given group. 
In some areas the many representatives of a group may be quite loosely 
associated, but more often the individual plants are widely distributed and 
located where they will show the best growth. A flower show, by contrast, 
offers an opportunity for displaying small plants or branches or portions of 
plants in a small area, conveniently displayed and studied. Such displays 
may be seen by more people in a few hours than may visit the living collec- 
tions in a full week. As a part of its effort in horticultural education, the 
staff of the Arboretum will prepare educational displays of plant materials. 
Regrettably, we cannot fill all requests or accept all invitations which, 
during the past year, were received from seven states, including Hawaii, 
from Canada, and from Europe. A display of ornamental fruiting shrubs 
and trees was shown at the Fall Show of the Massachusetts Horticultural 
Society attended by about 5000 people and at the comparable show of the 
Worcester County Horticultural Society which had about 9000 visitors. 
The staff cooperated with members of the Massachusetts Horticultural 
Society in preparing a Christmas Show which drew 3000 visitors to Horti- 
cultural Hall in Boston. The Arboretum received a silver medal for its 
display of cones and evergreens. A separate exhibit area featured a display 
of fruiting branches of selected hollies native to New England. 

During the spring season our largest exhibit was at the Massachusetts 
Horticultural Society’s Spring Flower Show at Revere, Massachusetts. 
This display, awarded a first prize and a gold medal, featured a collection 
of dwarf evergreen plants which will eventually be located near the Dana 
Greenhouses. Eighty-six thousand people attended this exhibition, At the 
request of the New York Horticultural Society the Larz Anderson collection 
of bonsai was taken to New York for the New York International Flower 
Show. A final exhibit, again by request, was a demonstration of methods 


1962] THE DIRECTOR’S REPORT 455 


of pruning trees and shrubs at the Jordan Marsh Spring Show sponsored 
by the Garden Club Federation of Massachusetts and seen by an estimated 
50,000 people. The various exhibits were designed and executed by Dr. 
Wyman, Mr. Williams, and Mr. H. Howard. Considerable time was spent 
this year in the preparation of permanent labels engraved in colored 
plastic for these displays, in an attempt to have a type of exhibit more 
easily assembled in the future. 


: f as ‘ie a 
pa i all e . 
a 

= as me Pe \\ 


Japanese bonsai of the Larz Anderson Collection of the Arnold Arboretum 
on exhibition at the International Flower Show, New York City, March 10-18, 
1962. 


ge 


Travel and Exploration: 


Pacific Science Congress, the National Shade Tree Conference, the Ameri- 
can Nurseryman’s Association, and the American Society of Horticultural 
Sciences. 

Dr. Howard collected some special plant materials for his research while 
in Hawaii for the Pacific Science Congress. The expedition to Burma of 
Mr. James Keenan, of the Royal Botanic Garden, Edinburgh, Scotland, 
sponsored in part by the Arnold Arboretum, was completed during the year, 


456 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XLII 


and the arrangement of materials collected is under way. Mrs. Claude 
Weber made a special trip to gardens and nurseries in Ohio and Illinois to 
study cultivars of Chaenomeles in flower. The trip was rewarding in the 
discovery of several old cultivars previously not known to be still in culti- 
vation. Mrs. Weber also participated in a class in tropical botany which 
spent a month in Costa Rica. This trip, financed by a grant from the 
Fernald Fund, allowed her to make general and special collections of plant 
materials for the herbarium and special collections for several staff mem- 
bers with research problems involving plants in that area. Dr. Wyman 
travelled extensively in the United States during the year, visiting many 
gardens and arboreta during peak seasons to continue his studies of orna- 
mental plants and to obtain new plants for trial at the Arnold Arboretum. 
Dr. Ernst left in late June for field work in Texas with Dr. Henry J. 
Thompson, of the University of California at Los Angeles, related to their 
joint research interests. 


Gifts and Grants: 


The Friends of the Arnold Arboretum who contribute regularly to the 
work of the Arboretum responded generously to an appeal during the 
spring. To these contributors we again express our continuing appreciation. 

Through the initiative of Mr. Seth Kelsey, of East Boxford. a member 
of the Committee to Visit the Arnold Arboretum, eight Massachusetts 
nurseries contributed over 1300 plants of various types and sizes to be used 
in establishing the basic landscape plantings around the Dana Greenhouses. 
Through their generosity, the greenhouses lost immediately a portion of 
the bleak appearance so characteristic of new buildings and freshly moved 
soils. 

One group of plantings proposed for the fenced-in area of the Dana 
Greenhouses was a collection of native hollies with most attractive fruits. 
A selection was offered the Arboretum in the fall of 1961 by Mr. Wilfrid 
Wheeler, of Hatchville, Massachusetts. Although Mr. Wheeler died on 
Christmas Day, 1961, his wishes were carried out by his sons, Wilfrid, Jr., 
Richard, and Charles. We regret that Wilfrid Wheeler could not see his 
collection of outstanding cultivars in their new location, and we are grateful 
to his family for this gift of plants which so well represent his long interest 
in and his contribution to horticulture in New England. 

Portions of the research of six staff members involving the services of 
six assistants continue to be supported by grants from the National Science 
Foundation, the National Institutes of Health, and the gifts of Mr. George 
R. Cooley. 


Publications: 


Four quarterly issues of the Journal of the Arnold Arboretum including 
most of the scientific publications of the staff were distributed during the 
year, as were the twelve numbers of Arnoldia which appear at irregular 


1962 | THE DIRECTOR’S REPORT 457 


intervals. These comprise the regular publications of the Arnold Arbo- 
retum. An issue of Arnoldia titled, “The Walter Street ‘Berrying’ Ground” 
was prepared by Mrs. Mary Lehmer, formerly of the Arboretum staff. This 
reviewed the history of the Walter Street Church and its adjacent cemetery 
now included within the boundaries of the Arboretum. Although remains 
are visible of the church, a memorial plaque, along with thirteen old head- 
stones and a crypt, mark the Revolutionary War dead and are decorated 
each Memorial Day by the historic commission of Boston. This article, 
which drew attention to a remote section of the Arboretum, is one of a 
projected series of articles on the history of the land we occupy. Another 
number of Arnoldia issued at the dedication of the Dana Greenhouses has 
been requested frequently by other botanical gardens and by architectural 
students. 


Bibliography of the Published Writings of the Staff and Students 
July 1, 1961 — June 30, 1962 

BarLey, Irvinc W. Comparative anatomy of the leaf-bearing Cactaceae, III. 
Form and Deve onelon of crystals in Pereskia, Pereskiopsis and Oi onic. 
ae Arnold Arb. 42: 334-346. 1961. 

(with Srivastava, Lattr M.). Comparative anatomy of the leaf-bearing 
Cactaceae, IV. The fusiform initials of the cambium and the form and 
structure of their derivatives. Jour. Arnold Arb. 43: 1874202. 1962 

BrizicKy, GEorGE K. The genera of Rutaceae in the southeastern United States. 
Jour. Arnold Arb. 43: 1-22. 1962. 

The genera of Simaroubaceae and Burseraceae in the southeastern 

Gaited States. Jour. Arnold Arb. 43: 173-186. 1962. 

. The genera of Violaceae in the southeastern United States. Jour. 

Arnold Arb. 42: 321-333. 1961. 

. A synopsis of the genus Columellia (Columelliaceae). Jour. Arnold Arb. 
42: 363-372. 1961. 
Taxonomic and nomenclatural notes on Zanthoxylum and Glycosmis 
(Ruta ceae). Jour. Arnold Arb. 43: 80-93. 1962. 

ERNST, WALLACE R. Blue grama at Goleta, California. Leafl. West. Bot. 9: 180. 

1961. 


(with Tryon, Rotta M.). Comments on the International Organization 
of Biosystematists. Taxon 11: 139. 1962. 
FORDHAM, ALFRED J. Germination of double-dormant seeds. Combined Proc. 
Plant eee Soc. 1960: 206-210. 1961. 
GREEN, Peter S. Herbaceous aliens in the Arboretum. Arnoldia 22: 49-56. 
1962 
. Studies in the genus Jasminum, II. The species from New Caledonia 
and the Loyalty Islands. Jour. Arnold Arb. 43: 109-131. 1962. 
. Watercress in the New World. Rhodora 64: 32-43. 1962. 
——- (with Lavener, L. A.). Catalogue of the names published by Hector 
Leveillé. I. Notes Bot. Gard. Edinburgh 23: 573-596. 1961. 
(with Tuomas, JoaB L.). The bulbiferous Ranunculus ficaria. Rhodora 
63: 289-291. 1961. 
Howarp, RicHarp A. Botanical and other observations on Redonda, the West 
Indies. Jour. Arnold Arb. 43: 51-66. 1962. 


458 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLII 


. The Charles Stratton Dana Greenhouses of the Arnold Arboretum. 
Arnoldia 22: 33-47, 1962. 
. The correct name for “Diospyros ebenaster.” Jour. Arnold Arb. 42: 
430-435. 1961. 

’ e Director’s report. The Arnold Arboretum during the fiscal year 
ended June 30, 1961. Jour. Arnold Arb. 42: 447-467. 1961. 
Naming and registering cultivated plants. Boxwood Bull. 1: 41-45. 


1962. 
(with JAYAWEERA, Don M. A.). The genus Duabanga in cultivation. 

Baileve 10: 8-13. 1962. 

(with oo. Don M. A.). A shrub called “Malitta”’ (Woodfordia 

fruticosa (L.) Kurz). Baileya 10: 14-18. 1962 

(With Nowburainig, TycHo). The i opincation of Diospyros ebenum and 

Diospyros ebenaster. Jour. Arnold Arb. 43: 94-107. 1962. 

JAYAWEERA, Don M. A. Some observations of abnormal flowers in the cultivated 
species of orchids. Orchid Rev. 69: 206-216. 1961. 

(with Howarp, RicHarp A.). The genus Duabanga in cultivation. 

Baileya 10: 8-13. 1962. 

with Howarp, Ricuarp A.). A shrub called “Malitta” (W oodfordia 
ae a (L,) Kurz). Baileya 10: 14-18. 

Kosusktr, CLARENCE E. Studies in the eas: XXXII. A review of the 
genus Ternstroemia in the Philippines. jour, Arnold Arb. 42: 263-275. 
1961. 


. Studies in the Theaceae, XXXIII. Variation in the fruit of Tern- 
stroemia kwangtungensis. Jour. Arnold Arb. 42: 426-429. 1961. 

NEVLING, Lortn I., JR. On the status of Psilaea (Thymelaeaceae). Jour. Arnold 
Arb. 43: 220, 221. 1962. 

A revision of the Asiatic genus Enkleia (Thymelaeaceae). Jour. Arnold 
Arb. 42: 373-396. 1961. 

. A revision of the Asiatic genus Linostoma (Thymelaeaceae). Jour. 
Arnold Arb. 42: 295-320. 1961, 

Perry, Liry M. Problems in the compilation of a native medicinal flora of 
southeastern Asia. Econ. Bot. 15: 241-244, 1961. 

SAX, Kary. Effects of X-rays on aging of seeds. Nature 194: 459, 460. 1962. 

Aspects of aging in plants. Ann. Rev. Pl. Physiol. 13: 489-506, 1962. 

(with Sax, Hatty J.). The effect of age of seed on the frequency = 

spontaneous and gamma ray induced chromosome aberrations. Rad. 

1: 80-83. 1961. 
SCHWARTEN, LazELLA (with RoGERSON, CLARK T., Rickett, H. W., & BECKER 
Index to American botanical literature. Bull. Torrey Club 88: 126- 
142, 187-213, 269-297, 370-385, 429-445. 1961; 89: 64-76, 

SRIVASTAVA, Lari M. (with Batrey, Irvine W.). Compatative anatomy of the 
leaf- bearing Cactaceae, IV. he fusiform initials of the cambium and the 
form and structure of their derivatives. Jour. Arnold Arb. 43: 187-202. 

1962. 


d 


Woop, Carroiy E., Jr. Magnolia. Encyclopaedia Britannica, 1961 ed. 14: 671, 
672 G1, 


(with BAKHUIZEN VAN DEN BRINK, R. C., BucHHEIM, E., Cowa AN, 


R. S., and Danpy, J. E.). Appendix II. Nomina familiarum SouseHvantla, 
Int. Code Bot. Nomenel. 1959. 187-201. 1961. 


1962 | THE DIRECTOR’S REPORT 459 
Wyman, Donatp. Barberries. Gard. Jour. N. Y. Bot. Gard. 11: 123-125; 138 
1. 


Barberries. Arnoldia 22: 9-16. 1962. 
The best of the dogwoods. Am. Hort. Mag. 41: 1-10. 1962. 
Best trees for 1962. Flower Grower 49(1): 46-48; 66. 1962. 
The birches. Arnoldia 22: 17-23. 1962 
. Dozen varieties include best in shrub Altheas. Am. Nurseryman 114(1): 
19: 121-127. 
. Few mountain ash species popular despite borers. Am. Nurseryman 
114(5): 12, 13; 106-112. 1961. 
———. The Forsythia story. Plants & Gardens 17(4): 17-19. 1962. 
. Green is what you make it. Better Homes & Gardens 40(4): 122B. 
62. 


. Hawthorns. Arnoldia 22: 25-32. 1962. 
_ Horse chestnuts and buckeyes now supplanted. Am. Nurseryman 
115(11): 11, 44-49. 1962. 
. The majestic beeches. Arnoldia 22: 1-7. 1962. 
Massive oaks are best long-lived ee trees for public areas. Am. 
Nurseryman 114(7): 12, 13, 104-110. 196 
es offer wide ae of choice es ee ornamental planting. Am. 
ape 114(11): 13, 87-97. 
The Saturday ene gardener. A guide to easy maintenance. New 
York, Macmillan. 236 pages. ilust. 1962. 
_ Shrubs and trees that flower in summer. Flower Grower 48(8): 22-24. 


— 
Ko) 


a eee popular despite limitations. Am. Nurseryman 114(9): 12, 13, 
106— 111. 1961. 

: Weeping willows leaders in group often misnamed. Am. Nurseryman 

114(3): 18, 19, 100-105. 1961. 

: the Arnold Arboretum is. Green Thumb 18(9): 309-313. 

1961. 19(1): 14-16. 1962. 


RicHArD A. Howarp, Director 


460 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XLIII 


Staff of the Arnold Arboretum 
1961-1962 


RicHarpD ALDEN Howarp, Ph.D., Arnold Professor of Botany, Professor 
of Dendrology, and Director. 


IrvinG WIDMER BaILey, S.D., Professor of Plant Anatomy, Emeritus. 


KARL SAX, S.D., Professor of Botany, Emeritus. 


GEORGE KONSTANTINE Brizicky, R. N. Dr., Botanist, Southeastern Flora 
Project. 

MicHaAeL, ANTHONY Canoso, M.S., Curatorial Assistant.* 

Henry Draper, Superintendent, Case Estates. 

WALLACE Roy Erwst, Ph.D., Botanist, Southeastern Flora Project.* 

ALFRED JAMES ForpHAM, Propagator. 

PETER SHAW GREEN, B.S., Horticultural Taxonomist. 

KATHERINE ANN HERRON, A.B., Business Secretary. 

HEMAN ARTHUR Howarp, Assistant Horticulturist. 

SHIU-YING Hu, Ph.D., Botanist. 

Don MartTINn ARTHUR JAYAWEERA, M.A., Mercer Fellow. 

CLARENCE EMMEREN KosuSkI, Ph.D., Curator.* 

MArGARET CATHERINE LeFAvouR, Herbarium Secretary. 

SUSAN DELANO MCKELVEY, A.B., Research Associate. 

Lorin Ives NEVLING, JR., Ph.D., Associate Curator. 

Lity May Perry, Ph.D., Botanist. 

BERNICE Gipuz SCHUBERT, Ph.D., Associate Curator. 

LAZELLA SCHWARTEN, Librarian.* 

Lait MOHAN SrIvVASTAVA, Ph.D., Mercer Fellow. 

Ropert Grerow WILtiaAMs, B.S., Superintendent. 

CarroLt Emory Woop, Jr., Ph.D., Associate Curator and Editor. 


Donatp Wyman, Ph.D., Horticulturist. 


* Appointed jointly with the Gray Herbarium 


1962] 


INDEX 


461 
INDEX 
Acarospora ee 63 BrizicKy, GEORGE K. The ie ) 
Adlumia, 338-339 nacardiaceae in the ieee 
gave ce eae 64 United States, 359 
— sisalana, 64 Brizicky, GEorGE K. Genera of 
Ageratum houstonianum, 66 Rutaceae in the Southeastern United 
Ailanthus, 179-180 States, 1 
Alvaradoa, 182-183 BRIZICKY, SORGE K. e Genera of 
Amaranthus dubius, 64 Sim aroubaceae and Burseraceae in the 
Amyris, 11-12 Southeastern United States, 173 
Anacardiaceae in the Southeastern United Brizicky, GEorRGE K axonomic and 
States, The Genera of, Nomenclatural Notes on Zaathoxylum 
Anacardiaceae, 359-360 and Glycosmis (Rutaceae), 80 
— tribe An pana reg 362 Buellia prospersa, 63 
— tribe Rhoé Bursera, 184, 185-186 
Anacardium, Be urseraceae in the Southeastern United 
Anatomy of the Leat- bearing Cactaceae, es e Genera of Simaroubaceae 


The Fusiform Initials 


of Cambium and the Form and 
Structure of Their Derivatives, 187; 
he Secondary Phloem, 234; VI. 


The Xvlem of Pereskia sacharosa and 
Pereskia aculeata, 376 

Annona squamosa, 65 

Antilles, Some Guttiferae of the Lesser, 
389 


Antilles, Volcanism and Vegetation in the 
Lesser, 279 

Argemone, 

Ash, prickly 

Asiatic ee 


oes 330 


Two New, 348 

Battey, I. W. Comparative Anatomy of 
the Leaf-bearing Cactaceae, VI. he 
Xvlem of Pereskia  sacharosa 
Pereskia Sey ‘ 

BaAILey, I. W., and Lair M. SRIVAST. AVA, 


and 


and Structure of Their Derivatives, 187; 
V. The ear Phloem 

Balsam apple, 

Bay cedar tee 

Bibliography, Ernest Jesse Palmer, 

ee eph Horace Faull, 


354 
230 


Boerhaavia coccinea, 
Botanical and ae Observ see on Re- 
donda, the West Indies 


Baneainy illea at a 


nd, 173 
Burseraceae, 183-186 


Cactaceae, Comparative Anatomy 


Pereskia aculeata, 376 
Calamus, 33 


ee ee 327 
a, 


ntillanum, 398 
— brasiliense “antillanurm, 398 
—calaba, 3 3 

— jacquinii, 3 
Cambium re fie Form and 
f ir Derivatives, The 
[ Comparative 
Leaf-bearing Cactaceae, IV, 

66 


Structure 
e203 


*, Grandiflorae, 
— ser. Spinosae, 

— arborescens, 2 
—aurantiaca, 204 
—chamlagu, 203 
— frutescens ecu 208 
— frutex, 203, 204 

— frncees 


204 


462 


Caragana age ee 209 
204 


, 40 

Cashew Family, 35 
Casuarina oe 64 
Catalpa silvestril, 217 
Catharanthus roseus, 66 


326 
Centrosema virginiana, 65 
Centrostachya indica, 64 
Cephalocereus ease 65 
CHANNELL, R d C. E. Woop, Jr. 
The 2 Pn in the Southension 
35 


A New Genus of 
ophulariaceae from, 215 
Chioris — 63 


Choisya 
Chry balan icaco, 390 
Citrus, 17-22 


jorantiitellt: 65 

oxylon ee 411, 420 
—remyi, 411, 

Cleome viscosa, 65 

Clusia, 389 

—sect. Anandrogyne, 396 


—venosa, 390, 396-397 

Cocos, 36 

Comparative Anatomy of the Leaf- 
bearing Cactaceae, IV. The Fusiform 


Croton flavens, 65 


JOURNAL OF THE ARNOLD ARBORETUM 


[ VOL. XLII 
Croton lobatus, 65 
Cynanchum parviflorum, 66 
Cyperus rung 63 
— sphac 63 
Pe ey gard of the Genus Vibur- 
um, A, 132 
Daphne ear! 344 
Daphne Family, 4 
Daphnopsis pee (Thymelaeaceae), 
Note on, 344 
Daphionss crassifolia eggersii, 344 
Dates ublication of the Journal 
Linnaea, 
Dicentra, 336 
avanne sanguinalis, 63 
Diospyros ebenum and Diospyros eben- 


eo The Typification of, 94 
Diospyros digyna, 
e 


ter, 100 
—ebenum, 1 
— glaberrima, 100 


Director’s Report, The. The Arnold Ar- 
boretum During i Fiscal Year Ended 

1962, 

9 

Ducal en. 412 
Earth-smoke, 342 

Ebony, 94 

Echinocitrus, 14 

eS OLF, DONALD A ms Study 

the Genus Viburnum, 

Elaphrium, 186 

Emilia coccinea, 66 

Enkleia, 220 

Eragrostis ciliaris, 63 

Ernest Jesse ae 1875-1962, 351 

ERNST, WALLACE R. The Genera of 

Fumariaceae in the 

315 


ae heterophylla, 65 


— hirta 

Fagara, 7, 8, 80 

FauLtt, ANNA F. Joseph Horace Faull, 
23 


1870-1961, 2 
Ficus citrifolia, 64 
— ROBERT Dates of Publication 
the Journal ae 
Frevcneti hemsleyi 
ostermansii, 34 
a-ahianceolata. 348 


1962] 


sie 334, 342-343 
ariaceae in the Southeastern United 
a The Genera of Papaveraceae 
and, 31 


cee eee e, —343 
— tribe ees 336 


Fumitory Family, 

Fusiform ate of the Cambium and the 
d Structure of Their Deriva- 
tives ao Comparative Anatomy of 
the Leaf-bearing Cactaceae, IV, 187 


Galactia, 65 


— stricta, 6 
Galba, 397 
Garcinia macrophylla, 


399 
Genera of ear dince se in oe South- 
eastern United States, The, 3 
Genera of Papaveraceae Ate ariaceae 
in the eae ae States, The, 
315 


Genera of Rutaceae in the Southeastern 
United States, The, 

Genera or Simaroubaceae and Bursera- 

he Southeastern United States, 


ceae in: 
The, 

Glaucium, 321 

— flavu 6 

Glycosmis (Rutaceae), Taxonomic 


d 
Nomenclatural Notes on Zanthoxylum 


and, 
Glycosmis, 12-13 
b 90 


XITl. 


Studies in the Genus Ses 
minum Species from 
Caledonia and the Loyalty ae 
109 


Grenada, 2 se 
Guadeloup 


Guttiferae th ae Lesser Antilles, Some, 


Haplophyllum, 3 


H 
Heterospat 
Hop-tree, ‘ 


INDEX 


463 


Howarp, RicH A. Botanical and 
Other Observations on Redonda, the 
West Indies, 51 

Howarp, Ree A. The Director’s Re- 
port, 

Howarp, Ricuarp A. Some Guttiferae 
os the Lesser ee 389 

WARD, RICH A. Volcanism and 
Vestn in aie mec Antilles, 279 

Howarp, RICHARD A., TycHo Nor- 
LINDH. The aden of Diospyros 
ebenum and Diospyros ebenaster, 94 

Hyphaene, 33 

eps ee crassifolia, 344 


gger 
Hyptis aes 66 
Tresine angustifolia, 64 


Jasminum, Studies in ue Genus, II. The 
w ledonia and the 


Jasminum sect. Sera: 112 
Unifoliolata, 115 


— brevistylis, 127 


15-12 
—linearifolium, 126-127 


— neocaledonicum, 122-123 


< 
isa 
oc 
E: 
5 
c 
B 
Yu 


Jatropha gossypifolia, 65 
Joseph Horace Faull, 1870-1961, 223 
Justicia periplocifolia, 66 


USKI, CLARENCE E. Ernest Jesse 


Dae 1875-1962, 351 


Lantana camara, 66 


464 


Lantana involucrata, 66 
Leaf Base in Palms, The. ie ee 
and Mechanical Bidlosy 
od, 432 


—438 

Leitneriaceae in the Southeastern United 
States, The, 4 

Leonotis nepetifolia, 66 

Limeberry, 14 

Limonia arborea, 90 


—sect. Psilaea, 221 


eit. iotands, The Species from New 
Caledonia and. Studies in the Genus 
Jasminum, II, 109 


Macleaya, 322 
—cordata, 317 

mmea americana, 390 
— humilis, 399 
—-— macrophylla, 399 
— — plumieri, 399 

—_ hii, 399 


Melicope gr randifolia, 414, 422 
— spathulata, 
Melocactus es 65 


ote on the Rela- 
tionships of Pinus merkusii, 10 
Mirov, NICHOLAS T. Ph enology of Tropi- 


Genus Platydesma 


OORE, Ravmonp J. On the Origin of 
errren sinica, 

Morphology and Mechanical aoe Its. 
The Leaf Base in Palms, 


Nageia minor, 76 

Nevis, 304 

NEVLING, Lorin I., Jr. Note on eral 
sis crassifolia Sag ae earae 

NEVLING, On eStats of 
Petlaen (hy melacaccse), ; 


JOURNAL OF THE ARNOLD ARBORETUM 


[ VOL. XLIII 


NEVLING, Lorin I., Jr. The Thymelae 
ceae in the Southeastern United sere 
428 


New Caledonia and the Loyalty eee 
The ee from. Studies in the Genu 
min II, 1 

Miechann: ees 


NorLINDH, TycHo, oad RICHARD A. How 
R 


e on Daphnopsis crassifolia (Thyme- 
laeaceae), 344 
Note on the Relationships of Pinus mer- 


ii, 
Nuées eanee 282 


Oldenlandia corymbosa, 66 
Oleaceae, 109 
Opuntia ee 65 


pe 
a 63 
Orange, oe 
Origin of Abie sinica, On the, 203 
Pacit, J. Shiuyinghua, 
of Scrophulariaceae from China, 215 
PALMER, ELIzABETH, and LAZELLA een 
TEN. Bibliography, Ernest Jesse Palm- 
354 
Palmer, Ernest Jesse, 1875-1962, 351 
Palms, The L eaf Base in. Its Morphology 
chan Biology, 23 
are fn o New foes 348 
Pandanus an 348 


A New Genus 


nd riacea 
cae “United : ae The Genera 
of, 
oe 315-: 
— subfan a ioe 316, 322 


-- eta Bechacholeisiden Sly 320 
—subf Papaveroideae, 317, 328 
— sub onoideae, 317 


. Platyste 
Pappophorum pappiferum, 63 


Paspalu m, 63 
Paulownia iverri 217 
Pelea, 413 


= dn icalaetott, 420 
Peperomia simplex, 64 
Pepper-tree, 365 
Pereskia eee The Xylem of aoe 
sacharosa and. Comparative Anat 
of the Leaf-bearing Cactaceae, VI, "376 
Pereskia, 189 


1962] 


Pereskia aculeata, 376, 378-380 

— sacharosa, 234, 376-378 

Pereskiopsis, 189, 381 

— chapistle, 234 

Phenology of Tropical Pines, 218 

Phloem, The Secondary. Comparative 
Anatomy a the Leaf-bearing Cacta- 
ceae, V, 23 

Phoenix, 35 

Phyllanthus amarus, 65 

Picramnia, 181-182 

Pilea microphylla, 64 

Pilo kea 

Pines, Spee of Tropical, 2 

eae merkusii, Note on the Ber tlie: 

, 108 


Pinus [group] Insignes, 108 
— [group] paar 108 


— elliottii 

— insignis, a 

— kasya, 21 

_ ener 108, 219 
— oocarpa 


8 
Pityrogramma chrysophylla, 63 
Platydesma (Rutaceae), A Monograph of 
the Genus, 410 
Platydesma, 411-413 


— sect. Cornutia, 413, 422-427 


—campanulata, 414 
—-— macrophylla, 414 
— — pallida, 4 


419-420, 422 


— — spathulatum, 416-417 
Platystemon, 317 
Plumbago scandens, 66 
Podocarpus, A Taxo 


South Pacific, 67 
Podocarpus sect. Afrocarpus, 68 
— sect. Se ere: 9 
— sect. Nage 
— sect. Stach carpus 68 


INDEX 


onomic Revision of, 
XIII, oecbon Polypodiopsis in the 


Podocarpus araucoensis, 67 


4 

Polypodiopsis in the South Pacific, Section. 
A Taxonomic Revision of Podocarpus, 
XIII, 67 

Povanie 14-17 

Poppy, 330 

Poppy, California, 327 

Poppy, prickly, 3 

Poppy, rock, 326 

Portulaca halimoides, 65 


Prickly poppy , 328 
i 2 
Psilaea (Thymelaeaceae), On the Status 
220 


Psi lea es 221 
Psilot 
Ptelea, on 
Pterocaulon virgatum, 66 
Puccoon, 322 


Quassia Family, 173 
Quiabentia, 189, 381 
—chacoensis, 234 


Ramalina subasperata, 63 
Red sandalwood, 94 
est Indies, aes and 


— subg. Sera 371 
— subg. Rhus, 

— subg. Toxtudendon 372 
Richard: 
Ricinus communis, 65 
Robinia sinica, 07 
Roccella babingtonii, 63 
Rock poppy, 326 
R 


Southeastern United 
States, Genera of The, 

Rutaceae, 1— 

—subfam. Aurantioideae, 12 


466 


Rutaceae subfam. Rutoideae, 5, 412 
ae, 9 


— tribe Xanthoxyleae, 412 


Sagra, Ramon de la, 84 


Sancalwood: ey 94 

Sanguinaria, 322-323 

Schinus, 365-366 

SCHWARTEN, LAZzELLA. Bibliography, Jo- 

3 
d ExizasetH M. 

Bibliography, Ernest Jesse 
Palmer, 354 

Scrophulariaceae from China, Shiuying- 
hua, A New Genus of, 215 

Section Polypodiopsis in the South Pacif- 
ic. axonomic Revision of Podocar- 
pus, XIII, 67 

Setaria setosa, 63 

Shiuyinghua, A New Genus of Scrophu- 
lariaceae from China, 215 

Shiuyinghua silvestrii, 217 

Shrubby trefoil, 9 

Sida cordifolia, Ne 

Sieve elements, 

Simarouba, i He a ay 

Simaroubaceae and Burseraceae in the 
Southeastern United Sales The Genera 


of, 173 
Simaroubaceae, 
b 


—subfam. Surianoideae, 176 

rie Picrasmeae, 179 

be Simaroubeae, 177 

ee tree, 367 

se Guttiferae of the Lesser Antilles, 


ee 283 

outh Pacific, Section Polypodiopsis in 
the. A Sag mic Revision of Podo- 
carpus, XI 

Southeastern she eee Pgs Genera 
of Anacardiaceae in the, 

reer ee United Ae aie Genera 
of Papaveraceae and Fumariaceae in 


the, 
Southeastern wna States, The Genera 
of Rutaceae in t 
Southeastern Deiter Brates, The Gen 
imaroubaceae and eae. in 
the, 173 


JOURNAL OF THE ARNOLD ARBORETUM 


[ VOL, XLIII 


Southeastern et States, The Leitneri- 
aceae in 

Southeastern United States, The Thyme- 
laeaceae in the, 4 

SRIVASTAVA, Latit i. and I. W. BaILey, 
Cc Leaf- 


and ledsciaads of Their 
187; 


Derivatives, 
. The Secondary Phloem, 234 
Stark viarohes jamaicensis, 66 

Staff of the Arnold ees 1961-1962, 


460 

Status of Psilaea (Thymelaeaceae), On 
the, 220 

Stonr, BenyAMIN C. A. Monograph of 
the enue eager nag 410 

STONE, BENJAMIN C o New Asiatic 
Pandanace as 

Studies in i Genus Jasminum, IJ. The 
Species from New Caledonia and the 
Loyalty Islands, 109 

Stylophorum, 323-325 

Suriana, 


Swallowwort, 326 
Talinum triangulare, 65 
182 


oa i, 
Taxonomic and Nomenclatural Notes on 


lication, oa The ae oe of Gly- 


cosmis, 86; Summary, 

Taxonomic sacs of . docarpus, A, 
XIII. Section Polypodiopsis in the 
South Pacific, 

Tephrosia cinerea, 65 

Thymelaea, 429 

Thymelaeaceae in the Southeastern United 

ates, The, 4 


Tillandste recurvata, 64 
B. The Leaf Base in 
Palms, Its Morphology and Mechani- 
cal sae : 
Torchwood, 
Torchwood amily 183 


Trefoil, shrubby, 9 


1962] 


Trianthema portulacastrum, 65 
Trichachne, 

—insularis, 63 
Tricholaena repens, 63 


Two New Asiatic Pandanaceae, 348 
Typification of Di 
Diospyros ebenaster, The, 


Veitchia, 


Viburnum, 8 Cytological Study of the 
Gen 13 


enus, 
Viburnum a Lentago, 147 
—sect. Megalotinus, 14 
—sect. Odontotinus, 149 
—sect. Opulus, 154 
—sect. Pseudotinus, 146 
— sect. oan 140 
—sect. Tin 8 
Se yaaakea new 149 
—alnifolium, 146 
—atrocyaneum, 148 
— awabuki, 
—betulifolium, 14 
—— ‘Aurantiacum,’ 149 
— bitchiuense, 142 
— x bo cL 140 
—-— ‘Dawn,’ 140 
— — ‘Deben,’ 140 
—bracteatum, 150 
— buddleifolium, 142 

—x Pea 144 

— burejaeticum, 14 


— — ‘Park Farm meer 142 
calvum, 14 
—x pee ie 142, 143 
— carlesii, 
—x are 143 
—— macrocephalum, 142, 143 


pubescens, 150 
ee 150 


Diospyros ebenum and 
94 


INDEX 


Viburnum eee hispidum, 150, 
— — pilosum, 
Le 151 
—edule, 154 
—ellipticum, 151 
erosum, 1 


-- secre lana 151 
— fragrans, 140 

—— album, 140 

—— candidissimum, 140 
—— gran ndiflorum, 140 


— icine 1 2 


—xj , 143 
— kansuensis, 154 
—lantana, 1 


44 
— — ‘Aurea Marginata, 144 
— — ‘Floribundum,’ 144 
—— ‘Lanceolatum,’ 144 
— — ‘Lees,’ 144 
—-— ‘Macrophyllum,’ 144 
4 


— lobophyl ; 
—macrocephalum sterile, 144 
—microphyllum 

—molle, 15 

— — leiophyllum, 152 
—mongolicum, 144 
—nudum, 14 


467 


151 


468 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XLIII 


Viburnum opulus Surely 154 Viburnum tinus hirtulum, 149 
‘Com mpactum,’ —— lucidum, 149 

— ; —— — ‘Variegatum,’ 149 

— — ‘Notcutt,’ 154 — — ‘Purpureum,’ 

——roseum, l —— variegatum, 1 

—— variegatum, 155 —tomentosum, 146, 147 

xanthocarpum, 155 — — mariesii, 

— orientale, 1 — — sterile, 155 

—ovatifolium, 152, 153 —trilobum, 155 

—parvifolium, 153 ‘Andrews,’ 156 

—phlebotrichum, 153 —— ‘Compactum,’ 156 

— photinioides, 141 ssee s,’ 156 

—plicatum, 14 — entworth,’ 156 

—— glabrum, 146 — urceolatum, 14 

— —‘Lanarth,’ 146 — utile, 

—— lanceolatum, 146 —veitchil, 145 

——ma ‘ —wilsonii, 15 

—— ‘Roseum,’ 146 ae 153, 154 

—— ‘Rowallane,’ 146 os andulosum, 154 
— ‘St. Keverne,’ 146 ae ssei, 154 


Volcanism and Vegetation in the Lesser 


— ‘Lanceolatum,’ 148 the Lesser Antilles, 280; Types of Vol- 
— prunifolium, 147 canic Activity, 282; Volcanic Erup- 
—rafinesquianum, 153 tions and the Vegetation, 283; Fuma- 
— —affine, 153 roles and Their Effects, 289; Effects of 
— recognitum ‘umaroles on Vegetation, 292; Cata- 
— rhytidocarpum, 144 logue of the Fumarole Areas of the 
ached helieides, 144, 145 Lesser Antilles, 296; Summation and 
—rhytidophyllum, 145 Comparisons, 305 
— — aureovariegatum, 145 
——  lantana, 144, 145 Wedelia calycina, 66 

— roset 145 West Indies, Botanical and Other Obser- 
— rigidum, 148 vations on Redonda, the, 51 
—roseum, 146 Wightia, 217 
—rufidulum, 147 Woop, C. E., Jr., and R. B. 
—sandankwa, 142 The Le jmeraceas in the a 

sargentii, 154, 155 United States, 435 

calvescens, 155 
— — flavum, 155 Xanthoxylum, 7 
— — ‘Puberulum’, 155 — americanum 
—scabrellum, 153 Xylem of Pevsken sacharosa and Peres- 
—schensianum, 145 kia aculeata, The. Comparative An- 


atomy of fhe Leaf-bearing Cactaceae, 
—setigerum, 15 ava ei) 

aurantiacum, 153 
= sieboldit, 


Zanthoxylum and Glycosmis (Rutaceae), 
Taxonomic and Nomenclatural Notes 


—stellulatum, 145 on, 80 
—suspensum, 142 eee ae = A 
—sympodiale, 146 — subg, 

— theiferum, a 


— subg. ee 7 
—coriaceum, 8 
=== 'Prendh ae 149 Zombia, 39