JOURNAL

OF THE

ARNOLD ARBORETUM

HARVARD UNIVERSITY

ALFRED REHDER JOSEPH H. FAULL and CLARENCE E. KOBUSKI

VOLUME XVI

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

New York

0E(

1968

DATES OF ISSUE

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

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

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

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

Printed in U.S.A.

TABLE OF CONTENTS

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

South America. By Ivan M. Johnston 1

and one text figure. By Alfred Rehder 65

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

tum. By Ernest J. Palmer 81

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

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

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

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

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

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

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

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

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

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

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

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

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

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

iv TABLE OF CONTENTS

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

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

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

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

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

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

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

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

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

Staff of the Arnold Arboretum 1934-1935 469

JOURNAL

OF THE

ARNOLD ARBORETUM

^I JANUARY, 1935

^UDIES IN BORAGINACEAE, X

THE BORAGINACEAE OF

NORTHEASTERN SOUTH AMERICA

Ivan M. Johnston

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

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

2 JOURNAL OF THE ARNOLD ARBORETUM Lvol. xvi

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

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

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

1935] JOHNSTON, STUDIES IN BORAGINACEAE 3

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

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

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

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

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

inconspicuous 2. Lepidocordia, p. 45.

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

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

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

4 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi

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

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

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

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

1935] JOHNSTON, STUDIES IN BORAGINACEAE 5

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

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

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

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

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

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

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

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

mm. long ; ovary and fruit glabrous ; inflorescences

terminating leafy branchlets.

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

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

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

3. C. fallax.

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

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

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

JOURNAL OF THE ARNOLD ARBORETUM Ivol. x

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

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

••••; 5. C. nodes.

Stems without subnodal swellings, not myrmeco-

philous ; calyx not with fibrous structure; plant

not bristly.

Fruit glabrous ; calyx with a fine minute strigose-

puberulence; lower leaf-surfaces glabrous

or practically so 6. C. lacvifron

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

crowded branches 8. C. ncrvosc

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

ovoid 9. C. fnlrc

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

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

•■■•■ 11. C. scahnjolic

Lower surface of leaves pallid, with a feltv

JOHNSTON, STUDIES IN BORAGINACEAE 7

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

12. C. bicolor.

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

of abundant slender appressed hairs 12. C. bicolor.

Lower surface of leaves not felty with a pallid

indument.

Stone globose or depressed globose, quite rugose ;

calyx with abundant long slender hairs on the

inner surface which project beyond the edge of

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

ation on their margins.

Calyx outside covered with abundant slender

silky hairs ; leaves rather thin, more or less

dimorphic at the stem-forks, lower surface

much paler than the upper 13. C. sericicalyx.

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

than upper 14. C. panicularis.

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

Hairs on lower leaf-surface erect.

Lower leaf-surface somewhat scabrid with

minute stout hairs; leaves large, 15-28

cm. long, with evidently falcate midrib ;

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

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

long, midrib weakly falcate; branchlets

Hairs on lower leaf-surface appressed.

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

leaves 8-20 cm. long; inflorescence large

and stiffish ; Lower Amazon and the

Guianas.

Leaf-blades broadest at or above middle,

". . . 17. C. exaltata.

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

17a. var. melanonenra.

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

6-11 cm. long; inflorescence slender and

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

la-lobes distinctly broader than long ; calyx cupulate or

ylindrical at maturity ; inflorescence dense, globose or

picate, or exceptionally a small loose cyme ; shrubs

JOURNAL OF THE ARNOLD ARBORETUM

witli erect or ascending I)ranche>^

;, frequently

sub-

scandent.

Corolla very large, 35-50 mm. long,

, the slender

tube

abruptly and much expanded into i

I coarse cylindrical

throat; inflorescence capitate; tip i

of calyx-lobe.

; con-

spicuously long-attenuate

19.

C. granc

iiflora

Corolla small, about 5 mm. long or le;

is, tube and t

weakly differentiated.

Inflorescence glomerate or cymose

20.

C. polycephala

Inflorescence distinctly spicate.

Leaves not hairy above, merely more or less v

erru-

cose or muriculate, elongate

; spikes tern

linal ;

petioles not decurrent on the

peduncles

21. C,

, macrost

achya

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

lustrous 22. C. Schomburgkii.

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

long acuminate 2i. C. tomentosa.

Calyx with tube nearly glabrous, lobes broadly

triangular, short acuminate 24. C. muUispicaia.

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

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

1935] JOHNSTON, STUDIES IN BORAGINACEAE 9

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

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

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

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

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

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

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

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

10 JOURNAL OF THE ARNOLD ARBORETLM [vol. xvi

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

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

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

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

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

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

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

Endemic to British Guiana.

1935] JOHNSTON, STUDIES IN BORAGINACEAE 11

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

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

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

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

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

12 JOURNAL OF THE ARNOLD ARBORETUM Tvol. xvi

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

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

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

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

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

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

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

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

19351 JOHNSTON, STUDIES IN BORAGINACEAE 13

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

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

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

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

14 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi

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

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

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

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

1935] JOHNSTON, STUDIES IN BORAGINACEAE 15

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

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

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

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

16 JOURNAL OF THE ARNOLD ARBORETUM [vol. xvi

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

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

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

The species is very