ShIITHSONIAN CONTRIBUTIONS TO BOTANY 0 NUMBER 39 

Studies in the Genus Boerhavia L. 

(Nyctaginaceae), 1-5 

F. Raymond Fosberg 

SMITHSONTAN INSTITUTION PRESS 

City of Washington 

1978 
ABSTRACT 

Fosberg, F. Raymond. Studies in the Genus Boerhavia L. (Nyctaginaceae), 1-5. 

Smithsonian Contiibutions to Botany, number 39, 20 pages, 1978.-The genus 

Boerhavia is discussed, its infrageneric classification is reviewed, and four subgenera are recognized. Distinctions of tw.0 Linnaean species, B. diffirsa and B. 

repens are clarified. Boerhavia mtitabilis K. Brown and B. pzibescens R. Brown 

are typified. The Boerhavia repens group is discussed and delimited, as is B. 

rubicunda and several similar species. Boerhavia repens var. maris-indici, B. 

albiflora, B. crispifolia, and B. heibstii are described as new. A curious abnormal 

form of B. crispifolia is also described and discussed. 

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded 

in the Institution’s annual reprt, Sinithsonian Year. SERIES COVER DLSIGN: Leaf clearing from 

the katsura tree Cercidiphyllurn japonicurn Siebold and Zuccarini. 

Library of Congress Cataloging in Publication Data 

Fosberg, Francis Raymond, 1908- 

Studies in the genus Boerhavia L. (Nyctaginaceae), 1-5. 

(Smithsonian contributions to botany ; 39) 

Bibliography: p. 

Includes index. 

1. Boerhavia. I. Title. 

QKl.SZ747 no. 39 [QK495.S9] 581’.08s [583’.933] i7-608165 

11. Series: Smithsonian Institution. Smithsonian contributions to botany ; 

39. 
Contents 

Page 

1 . Introduction to the Genus Boerhauia L ............................. 1 

2 . 4 

3 . Typification of Boeyhauia inutabilis R . Brown and Boerhauia pubescens 

R . Brown ...................................................... 5 

4 . Boerhauia repens Group .......................................... 8 

5 . Boerhauia rubicunda Steudel and Somewhat Similar Species ........... 13 

Literature Cited .................................................... 19 

Index ............................................................. 20 

Clarification of Boerhauia diflzisa L . and Boerhauia repens L ........... 

... 

111 

Studies in the Genus Boerhavia L. 

(Nyct aginaceae), 1-5 

F. Raymond Fosberg 

1. Introduction to the Genus Boerhavia L. 

Almost 200 binomials have been proposed in 

Boerhavia L. (often spelled “Boerhaavia”), according to the Index Kewensis. Of these, some are nomenclatural synonyms and homonyms, many are 

taxonomic synonyms, and others have been transferred to other genera. Some species are polymorphic, others very uniform. They are tropical or subtropical, and some have wide geographic ranges. 

Two major sources of taxonomic difficulty with 

Boerhavia are immediately apparent to one who attempts to identify plants belonging to it or to look 

at the genus as a whole. One is the level at which 

supra-specific categories are to be classified. The 

other is the apparent plasticity or polymorphism 

within the species. 

The first problem, as in several other groups in 

the family (e.g., Pisonia L., Mirabilis L., Abronia 

L.), is the level at which to recognize genera. Considered in a broad sense Boerhavia has a distinctive 

and recognizable habit or facies. No one at all 

familiar with the family is likely to confuse Boerhavia with any other genus. Technically it is not 

easy to circumscribe. Herbaceous or rarely slightly 

woody, usually elongate stems with prominent 

nodes and in some species with a band of sticky 

mucilaginous material around the middle of the 

internodes, slender forking cymose or rarely racemose or even spicate inflorescences with small, 

F. R. Fosberg, Department of Botany, National Museum of 

Natural History, Smithsonian Institution, Washington, D.C. 

20560. 

scale-like free bracts, small flowers with campanulate, cylindric, or funnel-shaped perianth limb sep. 

arated by a constriction from the sulcate, enlarged 

lower portion of the perigone, which develops into 

a clavate or oblong, often glandular, anthocarp or 

fruit; these form a good combination of recognition 

characters. 

Within this circumscription four groups of species are easily recognized: (1) Plants with 10 welldefined sulci on a symmetrical fruiting perigone 

have been segregated by some as the genus Anulocaulis Standley. They have the sticlty band around 

the internodes prominently developed. Four or five 

species are included here, occurring from the Death 

Valley region of California and Nevada to Coahuila, Mexico. (2) Another group has rather large, 

sometimes slightly asymmetric fruits, faintly 10-sulcate, provided with a few prominent sticky glands. 

The flowers are larger and more funnelform and 

zygomorphic than is usual in the genus Boerhavia, 

sensu lato. The species with these characteristics 

have been given the generic name Commicarpus 

Standley. Two or more of the species are very 

widely distributed. (3) Still another group, with 

only one or two Mexican species, producing similarly large flowers in racemes and very asymmetric, 

rather gibbous, fruits, has been called Cyphomeris 

Standley. (4) The remaining species, Boerhavia L. 

sensu stricto, have small, almost regular flowers, and 

3-5-sulca te, glabrous or viscid-glandular symmetrical fruits,, the glands small and sessile or shortly 

1 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

stalked. The species of this group are well distributed in open habitats in the warm parts of the 

world. 

No answer satisfactory to all taxonomists is possible to the question of whether these four groups 

should be recognized as separate genera or rather 

as sections or subgenera of one broad genus, Boerhauia L. For those who are interested in relationships, who better comprehend larger complex 

groups, or who are not specialists in the family 

Njctaginaceae, the broad single genus will be preferable. Those who find smaller, more coherent, 

simpler groups easier to remember and understand, 

to whom relationships are not as important as differences, or who are Nyctaginaceae specialists may 

prefer to maintain the four genera. My own inclination is to the first of these viewpoints. Also, since 

the generic name and specific epithet are what form 

the specific name, it seems to me that a wide knowledge of plants is made easier by maintaining the 

members of these two ranks, genus and species, in 

as broad a sense as is consistent with phylogenetic 

probability and morphological continuity. In this 

study, Boerhaviu will be used in the broader sense 

indicated above and in its synonymy. 

The other source of confusion is the apparent 

extraordinary plasticity or polymorphism of some 

of the species, especially the strand and coastal ones. 

These latter have, at least the Old World ones, 

frequently been treated as belonging to one enormous wide-ranging and complex species to which 

the name Boerhauia diffusa L. has usually been applied-mistakenly in my opinion. If B. diflusu is 

removed (pp. 4-5) much of the remaining complexity, again at least in the Old World, might be included in B. repens L. 

It would be possible to treat B. repens as a protean species to include all prostrate axillary-flowered, thick-rooted boerhavias, as has been done in 

the past, under the misapplied name “Boerhavia 

difura.” This course has not been satisfactory to 

many field observers who have had much experience 

with the boerhavias of the Pacific islands or those 

of the Caribbean. Without having any illusions that 

all the problems presented by these plants are 

solved, it seems that a partial resolution of some 

of the difficulties might be made by recognizing 

some of the clearer components of the complex as 

separate species. In this study a beginning will be 

made toward doing this and toward clarification 

of the application of several of the older names 

that have been applied to members of the group. 

ACKNOWLEDGMENTS-I wish to thank the authorities of the numerous herbaria where I have, during 

many years, been able to study plants of this genus, 

some of whom have sent specimens as loans or 

gifts. Thanks are also due to Miss Dulcie A. Powell, 

who has made numerous dissections and notes on 

floral details. Her work and some of the field work 

and herbarium visits have been supported by grants 

from the Smithsonian Research Awards Program, 

administered by the Smithsonian Research Foundation. Logistic support for field work in the Phoenix 

Islands was provided by the U. S. Air Force. 

Boerhavia L. 

Boerhavia L., Gen. P1. ed. 5, 4, 1754 [1753]; Sp. P1. 3, 1753.- 

Choisy, in de Candolle, Prodr. 13(2):449-450, 1849.-A. 

Gray, Am. Jour. Sci. 11. 15:321-324, 1853.-Heimerl, Beitr. 

Syst. Syctag. 24-29, 1897; in Engler and Prantl, Nat. 

Pflanzenf. 111, l(h):26, 1889.-Standley, Contr. U. S. Nat. 

Herb. 12:375, 1909; North Amer. Fl. 21(3):204, 1918.- 

Heimerl, in Engler and Prantl, Nat. Pflanzenf., ed. 2, 

16(c):117-119, 1934. 

Boerhaauia auct. plur. [orthographic variant]. 

Lindenia Martens & Galeotti, Bull. Acad. Bruxelles 10(1):357, 

1843 [not Bentham, P1. Hartrv. 84, 351, 18421. 

Tinantia Martens gi Galeotti, Bull. Acad. Bruxelles 11(1):240, 

1844 [not Scheidweiler, in Otto and Dietrich, Allg. Gartenz. 

7:365, 18391. 

Senkenbergia Schauer, Linnaea 19:711, 1847 [not Senckenbergia Caertner f., Meyer and Scherbius, Oekon. F1. Wetterau 2:413, 18001 [orthographic variants]. 

Cornmicarpits Standley, Contr. U. S. Nat. Herb. 12:373, 1909: 

13:428, 1911; 18:101, 1916; North Amer. F1. 21(3):215, 1918. 

Andocaulis Standley, Contr. U. S. Nat. Herb. 12:374, 1909; 

13:430, 1911; North Amer. F1. 21(3):216, 1918. 

Cyphonieris Standley, Contr. U. S. Nat. Herb. 13:428, 1911: 

North Arner. F1. 21(3): 214, 1918. 

TYPE SPEcIEs.-Boerhuviu diflusa L. (lectotype). 

DEScRIPTIoN.-Herbs, rarely vine-like, creeping, 

or scrambling, rarely somewhat woody at base: 

leaves opposite, simple, entire to sinuate or repand 

lobed, estipulate, usually petiolate; flowers borne in 

variously formed cymose, umbelloid, racemose or 

rarely spicate inflorescences, these bracteate, the involucral bracts subtending ultimate partial inflorescences not connate: perianth of a single completely united series of parts termed a perigone, this 

separated into two parts by a strong transverse constriction, distal portion, corolloid, campanulate, 
NUMBER 39 3 

tubular or funnelform, caducous, proximal portion 

longitudinally sulcate, persistent, at maturity closely 

investing the fruit, termed an anthocarp; stamens 

basally attached, filaments free, anthers with 2 orbicular cells, usually exserted; pistil 1, ovary unilocular, ovule 1, basally attached, style filiform, stigma 

capitate, usually somewhat exserted; anthocarp 3, 

4, 5, or 10 sulcate, indehiscent, containing one erect 

seed. 

says toward an infrageneric classification of Boerhavia have been written by Anton Heimerl in the 

two editions of Engler and Prantl, Die natiirlichen 

Ppanzenfamilien, in 1889 and 1934. The 1889 classification was used in modified form in Heimerl’s 

studies of the Nyctaginaceae in 1897. In the 1889 

classification, Heimerl recognized five sections: Senkenbergia A. Gray, Micranthae, Pterocarpon, Solenanthae, and Adenophorae, the last four all erected 

by Heimerl in this treatment. In 1897 he used this 

same arrangement except that he here merged sect. 

Pterocarpon into sect. Micranthae. 

Standley, in 1909 and 1911, elevated three of 

Heimerl’s sections to generic rank, giving them new 

generic names. Sect. Solenanthae became Anulocaulis, sect. Adenophorae became Commicarpus, 

and Senkenbergia became Cyphomeris. For the last 

of these, three earlier generic names exist, all later 

homonyms, as shown by the synonymy. Since Senckenbergia Gaertner f., Meyer & Schprbius and Senkenbergia Schauer are both named in honor of 

Johan Christian Senckenberg (Heimerl, 1934: 115), 

they are orthographic variants and, therefore, must 

INFRACENERIC CLASSIFICATION.-The principal eSbe regarded as homonyms for nomenclatural purposes (ICBN, Art. 64 note and Art. 75). 

Heimerl in 1934 accepted these genera, proposed 

by Standley, and treated Boerhavia L., sensu stricto, 

as including only what he had previously, in 1897, 

called sect. Micranthae. This genus he divided into 

four sections, Spicatae, Singulifiorae, Pyramidatae, 

and Clavatae. 

The four major subdivisions are without doubt 

natural groups, whatever rank they are assigned. 

Heimerl’s (1 934) four sections of Boerhavia sensu 

stricto are somewhat less convincing, but no better 

scheme has been proposed. For the purposes of this 

series of studies they will be adopted until the genus 

is well enough known that improvements or refinements become possible, The names of these sections 

must be brought into accord with the International 

Code of Botanical Nomenclature. 

Since a broad generic concept is adopted here the 

segregate genera are regarded as subgenera. These 

seem to be clearly distinguishable by the characters 

used in the key to subgenera below. 

In subgenus Boerhavia (Boerhavia L., sensu 

stricto, Boerhavia, sensu Heimerl, 1934) Heimerl 

distinguishes the four sections indicated above. 

These are recognizable here, but are very close together and it is by no means certain that this is the 

most natural arrangement possible, or even that 

they should be distinguished as sections (see Standley, 1909: 377). 

Within these sections are recognizable species 

groups, not yet well enough defined to be given 

formal recognition as series. They will be referred 

Key to the Subgenera of Boerhauia 

(modified from Standley, 1911:382) 

1. Anthocarp very asymmetric, inflorescences racemost, upper part of perigone funnelform 

1. Anthocarp symmetric to slightly asymmetric, inflorescences cymose or cymose paniculate, 

rarely with spicate or art of perigone campanulate, 

2. Anthocarp distinctly ate), upper part of perigone 

................................................................ ............................... cyphomeris 

cylindric or funnelform ...................................................... 2 

campanulate or cylindric ...................................................................... 

to salverform .................................................. 3 

3. Anthocarp tending to be cylindrical and somewhat asymmetric with large gummy 

3. Anthocarp without glands, strongly 10-ribbed, internodes with sticky bands Anulocaulis 
4 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

to as the Boerhauia repens group, the B. diflusa 

group, etc., without formal definition. 

Subgenus Boerhavia 

Boerhnavio auct. plur. orth. mut. 

Boerhavia sect. Aficranthne Heimerl, in Engler and Prantl, 

Nat. Pflanzenf. 111, l(b):26, 1889; Beitr. Syst. Nyctag. 24, 

1897. 

TYPE-SPECIES.-BoerhaVia diflusa L. (lectotype). 

DEscRrPTrorv.-Herbs frequently with enlarged 

roots, inflorescence variously cymose or cymose paniculate, smaller branches ending in capitate, glomerate, or umbellate or irregular clusters of small 

flowers, anthocarps 3-5 sulcate, often glandular, 

apices rounded or conic, mature or almost mature 

fruits articulate to and readily detached from pedicels or common receptacle. 

DISTRIBUTION-PantrOpiCal and subtropical. 

Section Boerhavia 

Boerhavia sect. Micranthae Heimerl, Beitr. Syst. Nyct. 84, 

1897 [pro parte]; in Engler and Prantl, Kat. Pflanzenf. 

111, I(b): 26, 1889 [pro parte]. 

Boehavia sect. Clavatae Heimerl, in Engler and Prantl, Nat. 

Pflanzenf. ed. 2, 16(c):118, 1934. 

Pantropical and subtropical. 

Includes the common widespread lowland and 

strand species with usually sticky fruits with 

rounded or conical apices. 

Section Spicatae 

Boerhauia sect. Spicatae Heimerl, in Engler and Prantl, Nat. 

Phnzenf. 16(c):118, 1934. 

Southwestern U. S. to Peru. 

A few annual species with flowers rather spicately 

to rarely capitately arranged on the inflorescence 

branches, anthocarps glabrous, 4-5-ribbed. 

Section Singuliflorae 

Boerhavia sect. Singuliflorae Heimerl, in Engler and Prantl, 

Kat. Pflanzenf., 16(c):118, 1934. 

New hIexico and Texas to Mexico (and Asia and 

Africa?). 

A few species with flowers on ultimate branches 

ot inflorescence mostly reduced to one, anthocarps 

glabrous or hairy, not sticky glandular. 

Section Pterocarpon 

Boerhavia sect. Pterocarpon Heimerl, in Engler and Prantl, 

Boerhauia sect. Pyramidatae Heimerl, in Engler and Prantl, 

Southern U. S. to Brazil and Ecuador, one species 

widely introduced in the Old World tropics and 

one or two probably indigenous African ones. A few 

species of annual plants with usually obpyramidal 

anthocarps with 3-5 keels or wings. 

Nat. Pflanzenf., ed. 1, 3(1b):26, 1889. 

Nat. Pflanzenf. 16(c):118, 1934. 

2. Clarification of Boerhavia diffusa L. and Boerhavia repens L. 

The two Linnaean names, Boerhauia diflusa and 

Boeyhavia repens, have generally been applied to 

a great diversity of Old World strand and lowland 

plants characterized principally by small flowers 

and narrowly clavate 5-ribbed, mostly viscousglandular anthocarps. The extreme view on this 

complex was that of Heimerl, who, in his later 

years, placed almost all Indo-Pacific Boerhauia in 

B. difl'usa, recognizing a multitude of varieties and 

forms. Boerhauia repens has occasionally been applied to prostrate forms, but until recently, not 

consistently to any one. 

In 1956 in Colombo, Ceylon (Sri Lanka) Prof. 

B. A. Abeywickrama pointed out to me as Boerhauin diflusa L. a plant of decumbent habit with 

fine ascending, diffusely paniculate inflorescences, 

noting that this was the type-locality of this species. 

To me, this was definitely not conspecific with the 

plants commonly thought to be B. diflusa in the 

Pacific Islands and so determined by Anton Heimerl 

in 1936, when he identified the Mangarevan Expedition collections (1937). 

Later study by me of the material in the Linnaean Herbarium suggests that, though very fragmentary, the specimen 9.3 of B. difusa could, indeed, belong to the Colombo species. This is here 

designated lectotype. The Linnaean specimen 9.8 

of B. repens, also rather fragmentary, belongs also 

in the compIex of prostrate plants generally called, 

up to then, B. difusa, but characterized by axillary 
NUMBER 39 5 

inflorescences. Linnaean specimen 9.2, marked B. 

diffusa has axillary inflorescences and belongs nearer 

to B. repens, though not identical with it. 

Study of a wide range of specimens apparently of 

this relationship suggests that a reasonable division 

can be made on the basis of strictly terminal paniculate inflorescences (B. diffusa group) versus axillary 

pendunculate cymes or umbels (B. repens group). 

Both of these are complex groups, in which several 

reasonably distinct species can be recognized. Alternatively, they may be regarded as two very broad 

collective species. The principal possibility of confusing the two is in populations of B. repens where 

repeated branching takes place distally, giving a 

paniculate appearance. Here the presence of axillary peduncles with umbelloid or glomerate inflorescences on the median parts of the plants seems 

a good distinguishing mark for B. repens and its 

close relatives. 

Boerhawia diflusa L. in a narrow sense is a decumbent plant with rather elongate prostrate main 

branches radiating from a root-crown, with ample, 

diffuse, finely branching, ascending panicles of small 

purple or pinkish flowers and narrowly clavatefusiform sticky anthocarps. It is common in disturbed weedy places in Ceylon, with related forms 

widespread in South Asia, Malesia, and to Australia. 

How B. paniculata Richard, B. coccinea Miller, B. 

caribaea Jacquin, B. wiscosa Lagasca et Rodriguez, 

and B. hirsuta Willdenow, all of the New World, 

differ has not yet been worked out to my satisfaction, but several species probably exist in this 

complex. 

3. Typification of Boehavia mutabilis R. Brown and Boerhavia pubescens R. Brown 

The original publication of B. mutabilis R. was separated out as the “Public Collection” (W. T. 

Stearn, BM(NH), pers. comm.). 

Thus Brown may have had in hand as many as 

lected in “Nova Cambria: Bustard Bay, 1770” by 

Banks and Solander (as the label is in Solander’s 

Brown (Prodr. 422, 1810) follows: 

1. B. mutabilk, procumbens glabra v. tenuissime pubescens, 

caule tereti, foliis repandis subovatis obtusis: paginis discoloribus, floribus 3-2-andris (T.M!) V.V. 

Obs. B. tetrandru Forst. forsan hujus varietas. 

10 plants of his own gathering, as well as one col1. B. mutabilis, procumbent glabrous or very thinly pubescent, stem terete, leaves repand subovate obtuse: surfaces 

not of the same color, flowers with two or three stamens 

(T.M.) seen alive. 

Observation. B. tetrandra Forst. perhaps a variety of this. 

[Translation by Fosberg.] 

This name, Boerhawia mutabilis, has been a 

source of confusion in a genus that is, even without 

it, about as confusing as a genus can be. The epithet 

mutabilis suggests that Brown himself encountered 

problems with it. An examination of his material 

and his field notes in the British Museum leaves 

one more than sympathetic with Brown, especially 

as he lived at a time when special creation and 

fixity of species were normal doctrine. 

There are 8 sheets of specimens in the herbarium 

collected by Brown, possibly 9 collections-it is not 

certain if on one sheet there are two separate gatherings. Another plant is mentioned in the field 

notes, but could not be located. Of course, the best 

set of his material was lost at sea, and another set 

hand according to Drs. Marshalband Groves of the 

British Museum botany stall). In Brown’s field 

notes, these specimens were initially indicated as 

separate species and varieties of B. mutabilis. By 

the time of publication of the Prodromus they had 

been reduced to two species, B. mutabilis for most 

of them and B. pubescens for one with notably 

pubescent stem and leaves. The latter is a plant of 

the Boerhawia repens L. relationship, soft-pubescent 

with multicellular hairs and “flores subtriandri.” 

No flowers remain on this Brown specimen from 

Carpentaria, which must be regarded as the holotype of B. pubescens R. Brown. Although the label 

of this specimen bears only the locality “Carpentaria,” in Brown’s field notes is the statement, “In 

apricis arenosis prope littora insularum lalons occidentalis Sinu Carpentariae.” The specimen has 

axillary cymes, with insertion appearing pushed out 

of the axil proper by the development of a branch. 

It represents a pubescent form of Boerhavia repens 

L. that occurs rather widely distributed in the Pa- 
6 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

cific Islands. The type is soft-pilose, hairs multicellular, with ovate, obtuse, mucronulate leaves that 

are smaller distally as the stems elongate; cymes one 

at an axil, with an axillary branch pushing the 

peduncle out of the axil as it develops. The branch 

bears another cyme, peduncle about 3 cm long, 

bearing a tiny not quite umbellate cluster at the 

top with 5-8 pedicels 2-3 mm long, no flowers left, 

but the field notes say “flores subtriandri.” In the 

B. repens complex the number of stamens per 

flower is somewhat variable, usually between 2 and 

4, and cannot be relied on as a distinguishing character. However, considerable field observation suggests that forms otherwise morphologically and 

habitally distinct, are each likely to have preponderately a particular number of stamens. 

I do not at present regard this pubescent form as 

specifically distinct from B. repens L., but as it is, 

nevertheless, identifiable over a substantial geographic range, it may be given the rank of “varietas.” 

If further field study shows that local populations 

in different regions are distinguishable, its rank 

might have to be elevated to subspecies within 

which varieties could then be described. For the 

present, the following disposition will serve: Boerhavia repens var. pubescens (R. Brown) Fosberg, 

new combination and new status (Boerhavia pubescens R. Brown, Prodr. 422, 1810). 

The following eight (or nine) specimens in BXI 

(plus one mentioned in Brown’s field notes but not 

located in BM) form the basis of B. rnutabilis R. 

Brown and from among them a lectotype must be 

designated, to firmly pin down this name, which 

has been variously applied by different writers and 

collectors. They are a diverse lot and come from 

widely scattered localities around the Australian 

coast. Brief descriptions of these sheets and designation of one sheet as lectotype of B. mutabilis, 

with reasons, follow. The Brown specimens have 

been arbitrarily given numbers 1 to 8 to facilitate 

discussion, even though two of them also bear 

numbers given by Brown. 

1. This specimen from Bay XII, 10 March 1802, 

is from the South Coast of Australia according to 

Brown’s system of indications in his field notes. It 

lacks fully developed leaves and has only one umbel 

with two remaining fruits on an elongate, probably 

originally paniculate branch. It is borne on a strong 

root crown. The leaves are described as oblong 

lanceolate, 5-6 lines long, which suggests that he 

was describing the reduced distal leaves that still 

remain on the specimen. The flowers are described 

as purple, with 2 stamens, the style curved above. 

The specimen is so incomplete that I cannot at this 

time assign it to a species. 

2. A specimen (Brown 10) indicated in the field 

note? as from “island a Nov. 24, 1802,” but labeled 

“islands a, b, e, f, g, h, 1, main opp. Groote Island,” 

is typical B. difusa L., with ample leaves, a coarse 

subglabrous stem, and a paniculate inflorescence 

about 30 cni long; stamens 2. On the label and in 

the notes this is indicated as “p undulata.” Since 

this epithet was not published by Brown, it has no 

status in the nomenclature of B. mutabilis. 

Another plant is described in the notes as from 

“Island a,” collected on the same day, differing in 

having less undulate leaves with less prominent 

veins, and 3 stamens. This specimen was not located. 

3. Mounted on the same sheet as No. 1 is a plant 

labeled “A desc.” and “Boerhavia diandra? Capentaria Island e Decbr. 6.” (It is not my intention here 

to publish a new binomial but merely to quote 

Brown’s label.) This is a much-branched subglabrous plant with small oblong-elliptic leaves, a 

somewhat paniculate rather stout inflorescence with 

flowers ultimately in few-flowered glomerules rather 

than umbels, fruit not very glandular (though the 

description in the notes contradicts this), and the 

stamens are described as “3 interdum 2.” Though 

rather too coarse, the paniculate inflorescence places 

this plant close to B. difusa L., but the leaves are 

more like those of B. tetrandra Forster. 

The sheet bearing this and No. 1 is marked “type 

specimen” but with no indication by whom or that 

such a lectotypification has ever been published or 

to which of the two plants it applies. Hence it is 

here disregarded. 

4. A specimen labeled “Boerhavia mutabilis & 

parvifolia/Boerhavia pumilio/B spec. [or B. desc.]/ 

Carpentaria Island e Dec. 6, 1802,” which seems 

certainly to be two branches of a witches-broom 

caused by Albugo infection (a common phenomenon in Pacific strand Boerhavia). It is rather condensed in habit, but not as much as some others; 

leaves small 6 x 4 mm; but flowers abnormally 

large, 5-10 mm long, possibly because the perianth 

limb persistent on the perigone, continued to grow. 

5. On the same sheet with No. 4 are 2 sterile 

branches, designated ‘Iy axillaris,” of a large-leafed 
NUMBER 99 7 

plant, that look like sterile branches of B. diflusa 

L. No locality is given unless the same as that of 

var. e [4, above], “Carpentaria Island e,” which is 

on a label that could apply to both specimens. On 

the y axiElaris label is “Ramuli petioli” and “venis 

. . . folia pulvero-pubescentes” and in the field notes 

“axillaris Foliis planiusculis majoribus umbellis 

rubra ramis solitariis caulo angulato? An distincta 

species? an B. diffusa Lin.?” “Ora orientalis” [= East 

Coast?]. 

6. A specimen (Brown 44) from Keppel Bay, 10- 

14 September 1802, is a plant with subglabrous 

stems branched at base from a heavy root-crown, 

with ovate-oblong obtuse leaves and axillary, filiform, almost hair-like cymes with a few umbellate 

flowers at their tips. In some leaf axils, in addition 

to the cymes, there are axillary branchlets bearing 

more cymes. The stamens are described in the notes 

as “2.” The capillary cymes of this specimen bring 

to mind plants of the high islands of the Pacific, 

such as Hawaii and the Marianas, which have 

frequently been called B. mutabilis R. Brown, but 

which have conspicuously acute leaves, rather than 

obtuse as in this specimen. A Robert Brown specimen without data in Leningrad corresponds to this 

one. 

7 and 8. On a sheet labeled “Boerhavia mutabilis 

a glandulosa” “Northumberland Islands Sept. 30, 

1802” are two series of fragments. A diagnosis in the 

field notes says “foliis undulatis glanduloso.” 

Number 7 (the upper row) labeled as above, is of 

6 pieces, very stunted, witches-broom-like, congested, 

with leaves ovate, about 1 cm long, short pubescent. 

Number 8 (the lower row) is of 5 larger fragments, 

not so congested, and are merely labeled “Carpentaria.” The leaves are larger, more broadly ovate, 

conspicuously undulate, notably short-pubescent, 

acute to obtuse, mucronate, with axillary cymes 

shorter than the leaves. 

The “glands,” abundant on both series, but more 

so on the upper pieces (No. 7), are sessile black dots 

that seem to be fruiting pustules of a fungus. The 

description in the field notes (1 October 1802) is 

rather full, but does not mention the “glands.” It 

describes the flowers as on subaxillary peduncles, 

“1-2 floris,” limb campanulate, stamens 2, style filiform, equaling the stamens. 

This plant does not exactly resemble anything 

I know, but I am sure it is strongly affected by a 

fungus. 

Through the kindness of Dr. J. B. Marshall of 

the British Museum Herbarium, material from these 

two series was submitted to the Mycological Institute at Kew. Their report (3 August 1973) is as 

follows: 

Dr. Sutton confirms that fungal mycelium is present in 

the leaf tissues of the two collections of Boerhavia mutabilis 

var. glandzilosa (IMI 177418 and 177419). However, it does 

not appear to be consistently associated with the occluded 

cells and it appears unlikely that the latter condition is the 

direct result of fungal infection. It is not possible to identify 

the mycelium without other vegetative structures or reproductive organs. 

The mention of “occluded cells” does not refer to 

anything observed by me, nor is it clear why the 

black dots or pustules were not mentioned. 

In addition to the specimens collected by himself, 

Brown apparently had available a sheet presumably 

collected by Banks and Solander at “Nova Cambria: 

Bustard Bay, 1770” (Solander’s hand, according to 

J. B. Marshall and Groves). This seems to be a very 

wilted and crumpled specimen of B. diflusa L., with 

a paniculate inflorescence, In addition to filling out, 

somewhat, Brown’s concept of his B. mutabilis, this 

sheet indicates that the weedy B. diflusa had 

reached Australia in pre-European times. 

This account of the protologue of Boerhavia 

mutabilis shows that Brown’s concept of his species 

was most inclusive, as well as indicating how appropriate was the epithet “mutabilis.” To clarify the 

application of the name, according to modern 

standards, a lectotype must be chosen from among 

the diversity of specimens. 

Brown gave no indication of what he thought 

was his species, sensu stricto, unless the “A desc.” 

on sheet no. 3 is an indication, along with his 

indications of “B. desc.” on sheet no. 4 and and a 

on sheets 5 and 7-8. Whoever noted sheet no. 

1-3 as type presumably thought so. Brown’s brief 

description in the Prodromus (1810:422) is of little 

help. 

Most of these specimens are either too fragmentary or too abnormal (because of fungal attack) 

to be easily associated with known taxa. Sheet no. 

2, Brown 10, is clearly B. diffusa L., with a diffuse 

panicle and large undulate leaves. 

Since the application of the name is not at all 

clear from Brown’s published description and since 

his specimen, Brown 10, is the one most clearly 

referable to a known species, it seems entirely 
8 ShlITHSONIAN CONTRIBUTIONS TO BOTANY 

proper to dispose of this troublesome name, Boer- nomic synonym of Boerhavia diffusa L. (1753), and 

hnuia mutabilis R. Brown, by choosing Brown 10 need no longer be used. 

as its lectotype, which is hereby done. It is not yet The only tenable alternative would be to choose 

possible to dispose of all the remaining specimens sheet no. 6, Brown 44. This seems less advisable as 

in Brown’s protologue except as indicated in the its leaf characters are fairly different from those of 

discussion of these specimens above. Boerhnvia the plants that at times have been referred to 

mutabilis R. Brown (1810) now becomes a taxo- B. mutabilis. 

4. Boerhavia repens Group 

Perhaps the most taxonomically refractory group 

in the genus Bee?-hauia is composed of an indeterminate number of taxa of seacoast and arid-land 

plants with basically axillary inflorescences, and 

5-ribbed clavate fruits. They are mostly prostrate, 

with elongate stems, often forming loose or dense 

mats. In some forms the leaves become gradually 

reduced in the most distal parts and the branching 

appears paniculate, lateral branches developing on 

alternate sides, and what appears to be a main axis 

is strongly zig-zag. The peduncles, however, are still 

axillary. 

This is, as presently understood, an Old ‘CVorld 

group, extending from Africa to throughout Polynesia. It may be closest to the New TVorld B. coccinea 

and its relatives, as these usually bear some axillary 

peduncles and have clavate, usually glandular anthocarps. 

The taxonomy of the B. repens group is not yet 

understood. Several taxa seem reasonably clear. 

Boerhauia yepens, sensu strictissimo, is African, 

originally from h’ubia, but I have seen too little 

African material to define it to my satisfaction. TWO 

Pacific species, B. tetl-andra and B. albiflora, new 

species, seem fairly clear, but probably hybridize 

locally with related forms. They are treated below, 

along with a preliminary discussion of B. yepens 

and two of its varieties. One additional variety 

(B. i.epens var. pubesccns) is assigned above in the 

discussion of B. mutabilis Brown. More species and 

subordinate taxa remain to be clarified. 

The members of this group have usually been 

treated as B. difliisa L., which however, is a plant 

with a terminal paniculate inflorescence (pp. 4-5). 

Dr. L. E. W. Codd of the Botanical Survey of South 

Africa, (1966: 114, 121), working on African material, 

restricted B. repens to plants with axillary inflorescences. 

Field studies in the Central Pacific, Micronesia, 

and on the Great Barrier Reef islands have turned 

up a widespread rather distinctive white-flowered 

plant of the Boerhauia repens L. relationship. In 

some islands where it occurs, it seems to be the only 

Boerhauia encountered. In other islands others 

occur, but little intergradation has been noticed. 

It is described below as B. albiflora Fosberg, new 

species. 

Boerhavia repens L. 

Boerhavia repens L., Sp. P1. 3, 1753. 

Herbaceous plant with thickened root and several 

prostrate stems radiating from the root crown, 

sparingly to much branched; leaves mostly ovate to 

oblong, obtuse to acute at apex, rounded to truncate 

to subcordate at base, shortly petiolate, blade 

usually pale or even white beneath, green above; 

flowers borne in axillary pedunculate cymes or 

pseudo-umbels, usually from alternate axils, the 

peduncles appearing to be crowded aside from the 

axil proper by the growth of an axillary branch, 

which very soon bears axillary cymes, leaves often 

reduced distally, creating a paniculate appearance, 

but with a clear central axis; peduncle branched 

only near tip, branches umbellately or subumbellately branched, each branch ending in a glomerule 

of flowers, or the branch system reduced to a single 

umbellate to glomerate cluster of flowers, or the 

peduncle bearing one pair of leaves and a glomerate 

cluster of flowers; perigone strongly constricted 

below the middle, lower part glandular, 5-costate, 

upper part corolloid, campanulate, shortly 5-lobed; 

stamens 2-4, subequal with perigone; style also subequal; anthocarp clavate to ellipsoid, rounded at 

apex, 5-ribbed, very glandular. 

A widespread Indo-Pacific and African species, 
NUMBER 39 9 

type from “Nubia inter Mocho & Tangos.” This 

is a protean species most of whose varieties have 

not been clearly distinguished. 

Boerhuvia repens var. maris-indici Fosberg, 

new variety 

Herba prostrata internodiis 1-2 (-5) cm longis, 

radice gracile fusiformis striatis, foliis parvis 2 X 1.5 

cm basibus truncatis vel subcordatis, apicibus obtusis 

pedunculis axillaribus subumbellatis gracilibus, 

floribus glomeratis, anthocarpiis ellipsoidiis vel 

clavatis. 

Root narrowly fusiform, stems prostrate, puberulent, internodes mostly 1-2 cm long except toward 

base where they may be up to 5 cm, striate; leaves 

small, mostly about 1-1.5 X 0.7-1.0, rarely up to 

2 X 1.5 cm, becoming smaller distally, base truncate 

to subcordate, apex obtuse or acutish, margin often 

somewhat crispate, upper surface green, minutely 

glandular puberulent, lower pale, with about 3(-4) 

veins on a side, intricately and deeply rugose when 

dry, petiole slender, up to 1 cm long; peduncles 

very slender, 2-3, rarely up to 4 cm long, subumbelloid to rather irregularly branched above, puberulent, flowers glomerate to sub-openly arranged; 

bracts lanceolate; perigone with ribs smooth or 

somewhat glandular, intervals glandular, limb 

puberulent without, 1.5-2 times as long as lower 

part, whitish to purple, campanulate; stamens 4(-5), 

subexserted, anthers with cells orbicular; style subexserted, stigma discoid, peltate; anthocarp ellipsoid 

to somewhat clavate, 2.5-3 mm, shortly pedicellate 

to subsessile, more or less glandular with shortly 

stalked glands. 

This variety is similar in habit to Boerhauia 

repens var. minor Delile, (1824:119-120, pl. 3), described from Egypt, but which differs in that each 

axillary peduncle seems to be composed of 2 internodes, with a pair of leaflike bracts at the node 

between them. 

A number of the cited specimens, e.g. Renvoize 

1315, Gwynne 6. Wood 1258,1185,1211, and VeeversCarter 21, have all or most of the peduncles on 

small lateral branches rather than on the main trailing stems. In this they suggest var. minor Delile. 

Also a number of collections, Fosberg 52146 from 

the Seychelles, Gwynne 6. Wood 1330 from Assumption, Stoddart d. Poore 1240 and Fosberg49806 from 

Cosmoledo, Stoddart 6. Poore 1300, Fosberg 49732, 

Fosberg 49677 from Astove, and Fosberg 49551 from 

Aldabra have the leaf margins crisped somewhat as 

in B. crispifolia. This could indicate some gene 

exchange between the two. 

The anthocarps are rather variable both in shape 

and degree of glandularity, occasionally approaching those of B. crispifolia. The glands are mostly 

stipitate, however. 

SPECIMENS EXAMINED 

ALDABRA.-S.~. Abbott in 1892 (US). Mentor I.: Frazier 4 

(US, K); Renvoize 1315 (US); small islet in lagoon W of 

mouth of Gionnet Channel, Fosberg 49551 (US). 

ASSUMPTION.-Stoddart 1061 (K); Gwynne 6. Wood 1330 

(EA), close to B. elegans in leaf form; near settlement, Stoddart 1062 (K) (atypical but specimen scrappy); center of 

island, Frazier 759 (US). 

COSMOLEDO AToLL.-Menai Island: S of settlement, Stoddart Q Poore 1240 (K, US); Gwynne Q Wood 1258 (EA). Wizard I.: Thomasset in 1902 (K); central part of Wizard I., 

Fosberg 49806 (US, K), 49792 (US, K); Renvoize 1226 (US, K). 

ASTOVE I.-Ridgway 21 (US); Veevers-Carter 21 (EA); 

Fryer 9 (K); Gwynne & Wood I314 (K, EA); N of settlement, 

Stoddart k Poore I294 (K, US); Renvoize 1187 (US, K); Grand 

Anse, Fosberg 49677,49732,49702 (US, K). 

SEYCHELLEs.-cousin Island: NE corner, Fosberg 52145 

(US), 52146 (US); Roche Canon, Fosberg 52077 (US). 

FARQUHAR ATOLL.-North I.: Gwynne k Wood 1185 (EA) 

(as B. coccinea). Farquhar (South) I.: Gwynne 6 Wood 1211 

AMrRANTEs.-Remire I.: Gwynne t Wood 877 (EA), 886 

(EA) (very young). Iles Africains: Southern Is., Stoddart 6. 

Poore 1431 (US, K). Des Noeufs I. Vesey-Fitzgerald 5807 (K, 

2 sheets) (“tuberous roots fed to pigs”). 

(EA). 

Heimerl, in Schinz (18971, refers Voeltzkow 34 

(not seen) to a Boerhauia diflusa var. eudiBusa 

Heimerl f. psammophila Heimerl, possibly proposed in the place cited. I have not seen this specimen and hesitate to refer it definitely here, but from 

the brief remarks about it, that it has a leafy 

prostrate stem and axillary inflorescences, it likely 

belongs here. The uncertainty leads me to give 

my plants a new epithet rather than changing the 

rank of Heimerl’s epithet. 

Boerhaviu repens var. minor Delile 

Boerhavia repens var. minor Delile, Descr. Egypte, vol. 19 

(Histoire naturelle): 119-120, Atlas pl. 30 (by Redoute), 1824. 

Two or 3 glandular clavate-ellipsoid anthocarps 

on a bibracteate axillary peduncle. The 2 opposite 
10 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

bracts are like reduced leaves about half-way up on 

the peduncle. One cyme at a node, alternating sides, 

rarely the peduncle branched, with 2 cymelets. 

Opposite axil has a semi-abortive inflorescence or 

branchlet. Leaves unequal at a node. (Description 

drawn from Redoute plate, cited above.) 

Codd (1966: 121) regards this as synonymous with 

var. repens. The bracteate peduncles seem to justify 

maintaining var. minor for the time being, until 

more ample African material can be studied. 

Boerhavia repens var. pubescens 

(R. Brown) Fosberg 

Boerhavia pubescens R. Brown, Prodr. 422, 1810. 

For transfer and discussion see pages 5 and 6. 

Boerhmia tetrandra Forster f. 

Boerhavia tetrandra Forster f., Prodr. 2, 1786; PI. Esc. ins. 

Oc. .-lusrr. 41, li86.-Vahl, Enum. 284, 1894.-de Candolle, 

Prodr. 13(2):456, 1849.-Guillemin, Ann. Sci. Nat. Bot. 

series 2, 7:190-191, 1837.-Heimerl, Ann. Cons. Bot. Geneve 

5:188, IS‘Jl.-Setchell, Cam. Inst. Wash. Pub. 341:249-250, 

pl. 33B, 34A, E, 1924.-Taylor, Plants of Bikini, 176-177, 

1950. 

Boerhavia diflusa var, tetrandra (Forster f.) Heimerl, Beitr. 

Syst. Nyct., 27, 1897. 

Forster’s plant was from the Society Islands and 

his short description from the Prodromzts (1786:2) 

follows: “5 B. tetrandra, caule repente, floribus 

tetrandris. F. Ibidem.” 

I understand this to be a common, widespread, 

fairly characteristic plant recognizable by its rather 

long slightly arcuate internodes, firm, usually oblong or elliptic leaves not much tending to reduction distally, and axillary strong erect peduncles 

usually umbellate or (in bud) glomerate, at summit, 

the rays either ending in single pedicellate flowers 

or more usually in secondary umbels with raylets 

ending in single or more rarely 2 or 3 flowers, not 

ordinarily glomerate, and with limb of perigone 

clear pink, campanula te, margin spreading, scarcely 

lobed, almost like a diminutive morningglory 

flower in appearance. 

The Forster specimen at the British Museum, a 

photo of which was kindly sent by the then Director, 

Sir George Taylor, fits this concept quite well, as 

do the manuscript description by Forster quoted 

by Guillemin (1837: 190-191), and the descriptions 

by Setchell (1924:249-250) and by Taylor (1950: 176). 

Herbaceous prostrate plant with several to many 

elongate stems from a thickened root or cluster of 

such roots, stems branching, tending to form very 

loose mats, internodes up to 8 (-10) cm long, tending to be slightly arcuate; stems and leaves glabrous 

or minutely appressed puberulent or even rarely 

densely short-pilosulose, especially the young growth, 

tending also to be reddish; leaves rather thick and 

firm, oblong or elliptic, to almost orbicular, more 

rarely somewhat ovate or obovate, entire, base 

rounded to subcordate, apex rounded to emarginate, 

lower surface very pale, veins 4 or 5 on a side tending to be rather obscure, petiole 5-7 mm, rarely 

10-11 mm long; peduncles strong, erect, up to 

10 cm tall, either umbelloid or a single small cluster 

of flowers at summit, branches with one or several 

flowers, these in open clusters rather than close 

glomerules, flower about 4 mm long; perigone 

pubescent, with stalked glands especially on ridges 

of lower part, limb campanulate with margin 

spreading, only shallowly lobed, pink; stamens (2-) 

4 or 5, filaments straight to somewhat curved, anthers with 2 orbicular cells apparently opening 

while still in bud; style straight to somewhat curved, 

stigma peltately capitate; anthocarps clavate to 

oblong, 5-ribbed, glands especially on ribs, apex 

rounded to subtruncate. 

This species is known to me principally from low 

coral islands-atolls and barrier reef islands around 

high islands. I have seen it from the Marshalls, 

Gilberts, Phoenix Islands (Hull Atoll only), Line 

Islands, Rose Atoll, Swain’s Isand, Flint Island, 

Society Islands, Austral Islands, and the Tuamotus. 

Some Hawaiian specimens may belong here, but 

their final disposition must await study of the 

Hawaiian boerhavias as a whole. 

SPECIMENS EXAMINED 

LINE ISLANDS.-\vaShingtOn I.: N side, Long 1845 (US); SW 

road about 1.5 mi [2.4 km] E of village, Long 2824 (HAW, 

US): S road, 1 mi [1.6 km] E of village, Long 2826 (US). 

Palmyra Atoll: sl., Long 1814 (US), 1788 (US): Rock 10273 

(NY, US); Judd & Mitchell 12 (NY); islet directly N of W 

channel, Long 2832 (US); S side of runway 3/4 mi [1.2 km] 

from the W end of runway, Long 2861 (US). 

Christmas Island: Bergman 9 (NY), I1 (US). 

CENTRAL PACIFIC ISLAhDS.-HOWland I.: Bryan 1307 (NY) 

(unusually small, broadly ovate, obtuse to acutish leaves 

firm): Judd and Mitchell 155 (BISH) (some glands in furrows 

as well as on ridges), 
NUMBER 39 11 

Jarvis I.: Bergman 56 (NY) (Rowers purple, 2(-3) stamen). 

Tongareva Atoll: Cocos grove, Long 3125 (US), Long 3127 

(US), Long 3138 US; island N of the NW passage, Sibley 

(Long’s) 3173 (US). 

Vostok Island: Long 3191 (US), 3202 (US), 3208 (US). 

PHOENIX ISLANDS.-HU~~ Atoll: S leg of E Islet near W end, 

Gardner Atoll: S side of Main Channel, Fosberg 6. Smith 

SOCIETY ISLANDS.-S.~. Forster (BM, type). Tahiti: Tautira 

District: beach, Nadeaud 322 (G). Scilly Islands: Motu Roa, 

Quayle 232 (US). Tetiaroa Atoll: Motu-Aie, Fosberg 6. Sachet 

54624 (US, NY). 

COOK Isums.-Aitutaki: Akitua, Stoddart 2222 (US); Motukitiu, Stoddart 2342 (US); Tavaema, Motu, Stoddart 2345 

AUSTRAL ISLANDS.-Raivavae, Whitney Expedition 298 (Us). 

Tubuai I.: Tapapatauai I., 1 m, St. John 16404 (NY, 2 

Maria Atoll: Middle I., Fosberg 12121 (NY); S.E. Islet, 

TUAMOTU ARcHIPELACo.-Tatakoto Atoll: Chapin 809 (NY) 

Hao Atoll: Chapin 819 (NY); Barclay 3260 (US) (as Bow I.). 

Rangiroa I.: Moore 196 (US). 

Raroia Atoll: Doty 6. Newhouse 11111 (US, NSW), 11172 

Oeno Island: St. John 6 Fosberg 15191 (US). 

MARSHALL IsLANDs.-Eniwetok Atoll: Rujoru I., St. John 

23806 (US): Rigili I., St. John 23747 US; Rainey s.n. (US); 

Bogon I., St. John 23761 (US); Igurin I., Fosberg 24306 (US, 

BISH, NY); Bogombago I., Taylor 46-1307 (US, G). 

Bikini Atoll: Eniirikku I., Taylor 46-1030 (NY), 46-1034 

(US, BM); Airukiraru I., Taylor 46-1152 (NY, BM, G); Ailinginae, Sifo I., Fosberg 36685 (US, BISH, Fo). 

Rongelap Atoll: Rongelap I., Fosberg 36653 (US, BISH, 

Fo): Kabelle I., Fosberg 36668 (US, BISH, Fo); Gegen I., 

Fosberg 36729 (US, BISH, Fo); Rigonman I., Taylor 46-1376 

Utirik I.: Fosberg 33644 (US, BISH, NY) (somewhat paniculate). 

Taka Atoll: s.L, Amermon (Long’s) 2921, 2938, 2941 (all 

HAW); Taka I., Fosberg 33746 (US, NY); Watwerok I., Fosberg 33761 (US, BISH, NY). 

Fosberg 6 Stoddart 54820 (US). 

55777 (US). 

(US). 

sheets). 

Fosberg 12071 (US). 

(unusually short and bushy): 808 (NY) (normally elongate). 

(US), 11007 ,(US). 

(US, IVY). 

Jemo Island: Fosberg 33881 (US, NY). 

Likiep Atoll: Aikini I., Fosberg 27058 (US, BISH, NY) (fls. 

pink); Fosberg 27057 (US, BISH, IVY) (Rs. white): Fosberg 

27006 (US, BISH, NY) (fls. pink). 

Lae Atoll: Bikenaj I., Fosberg 34074 (US, BISH, NY); Fosberg 34076 (US, BISH). 

Ujelang Atoll: Bokan I., Fosberg 34163 (US, BISH, NY). 

Wotho Atoll: Wotho I., Fosberg 34277 (US, BISH). 

Ujae Atoll: Bock I., Fosberg 34320 (US, BISH, NY). 

Kwajalein Atoll: Enewetok I., Eosberg 3.1136 (US, BISH, 

Arno Atoll: Arno I., Stone 1103 (US); Hatheway 802 (Us, 

BISH, Fo); Enirik I., Anderson 3742 (US, NY), 3731 (US, NY). 

Namorik Island: Betche in 1881 (NSW). 

NY). 

Jaluit Atoll: Schnee in 1902 (NSW); Mejatoo I., Fosberg 

41418 (NY, US). 

GILBERT ISLANDS.-S.~., Woodford in 1884 (BM); Jensen 

in 1878 (BM). 

Onotoa Atoll: Moul 8202 (US) North I., Moul 8151 (NY, 

US); Tabuarorae I., Moul 8223 (NY, US). 

Butaritari Atoll: Butaritari I., W end of island near Ukiangang Village, Herbst 6. Allerton 2747 (US). 

Tarawa: Bikenibeau, Catala 27 (P); near Teachers Training 

College, Sachet 1443 (US); Ranapa, between Bairiki and Bikeniku, Sachet 1424 (US). 

Marakei Atoll, Adair 189 (US). 

SAMOA.-ROSe Atoll: center of island few yards north of 

monument, Harris 303 (BISH), 304 (NY) (leaves small broad, 

slightly emarginate, cymes axillary, unusually short, slender); 

Swerdlofl C Needham in 1970 (US); Schultr 24 (US). 

SWAIN’S ISLAND.-WhiSth W3382 (us). 

Boerhavia albiflora Fosberg, new species 

Planta herbacea, caulibus pluribus prostratis pallide-viridibus, foliis formis variis infra pallidis, 

pedunculis axillaribus apice umbelliformis, floribus 

albis glomeratis limbo perigonio campanulato, staminibus plerumque 2, filamentis styloque valde 

curvatis, antheris stigmateque leviter exsertis, anthocarpio ellipsoidale vel plus minusve clavato 5-sulcato 

sulcis valde glandulo-pubescentibus. 

Herbs with several to many long, pale green, 

creeping stems radiating out from a very thick (up 

to 8 cm) root-stock; leaves broadly oval, obtuse, dull 

green above, white beneath, tending to become 

smaller distally and more ovate, 4 or 5 pairs of 

veins, each leaf tending to produce a leafy branchlet 

in its axil, the branchlets at successive nodes on 

alternate sides developing, with a very reduced 

vestige on the other side; on the same side as the 

branchlet but off to one side a long stout peduncle, 

or a scar of one; peduncle to 12 cm, with an umbelloid summit, branches usually 5, of different lengths, 

each branch subtended by a subulate bract, and 

with several much reduced branches in a tiny umbel 

at its summit, each of these with a small glomerule 

of 4-10 or more subsessile flowers or fruits; perigone 

with lower segment 5-sulcate with glandular pubescence principally in the furrows, ridges tending to 

be mucilaginous, upper segment white, campanulate, with 5 emarginate lobes, externally puberulent 

especially in tiny tufts of hairs at summits of lobes; 

stamens 1-3, mostly 2, with strongly curved filaments and orbicular anther cells, anthers slightly 

exserted; style curved, making the flower appear 
12 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

somewhat zygomorphic, stigma slightly exserted; 

anthocarp ellipsoidal to slightly obovoid, narrowed 

toward apex, then rounded-substruncate, pentagonal 

in cross-section, with 5 costae, strongly glandular 

pubescent, especially in furrows, sticky. 

This is the principal species found in the Phoenix 

Islands and is very abundant there. It extends northward and westward in the Marshall Islands and to 

Wake Island, southward and eastward through the 

central Pacific islands, and also occurs on the southern islands of the Great Barrier Reef. In all this 

range, except for the degree of reduction of the 

leaves distally and the strength of the tendency 

toward paniculate inflorescences distally, there is 

surprisingly little variation. One specimen, Long 

2612 (US) from Phoenix Island, with flowers pink, 

leaves scarcely exceeding 12 mm in length and stems 

rather densely short puberulent, probably belongs 

here, as it has flowers with tufts of puberulence at 

top of upper perianth, 2 stamens, and glands mainly 

in grooves of anthocqrp. The plants from Tl’ake 

Island and the northern Marshalls have a characteristic leaf shape and may be treated as a geographical variety. It is newly described below as 

var. powelliae. The collections, cited below, from 

Malden and Starbuck islands tend to have 3 or 

even 4 stamens and in this respect are intermediate 

writ11 B. telrandra. The glands tend to be on the 

sides of the furrows in the Malden plants. 

Boerhauk albiflora Fosberg var. albiflora 

Described above. 

HoLo-rwE.--Fosberg 6- Stoddart 54748 (US), 

Enderbury Island. 

SPECIMENS EXAMINED 

GILBERT IsLAws.-Onotoa Atoll: South Island, Movl 8198 

(GH): North Island, Mod 8340 (GH), N end Korth Island, 

Moul 8084 (US). 

PHOENIX ISLAWS.- Canton I.: s.L, Shultz 21 (US); s.l., Degener 6- Hatheway 21313 (US); Browne in 1939 (BISH); around 

Air Base, FosOerg L- Walker 30207 (US); around Pan-American 

Hotel settlement, Fosberg 30883 (US): area back of docks, 

Fosberg 6 Stoddart 54892 (US). 

McKean I.: on E rocky slopes, Long 2030 (US); on the NW 

beach, Long 2046 (US): on the SW end on the inner slope, 

Long 2041 (US): on the W side of the lagoon, Long 2034 

(US); SW side, Long 2428 (US); near SW end, Long 2439 

(US); on the inner NE side of lagoon, Long 2443 (US): S 

side of island, Fosberg 55795 (US): W side of island, Fosberg 

55788 (US). 

Birnie I.: s.I., Fosberg G Stoddart 54724 (US). 

Phoenix I.: SIV side, Long 2622 (US): N side, Long 2077 

(HAIV); E side, Long 2081 (US): midwest rim above lagoon, 

Long 2083 (US); S end, Long 2087 (US): W of N end of 

lagoon, Fosberg 6 Stoddart 54762 (US): SW shore of island, 

Fosberg L- Stoddart 54767a (US), Fosberg & Stoddart 54768 

(US); S margin of lagoon, Fosberg b Stoddart 54770 (US). 

Enderbury I.: inside C. S. Air Force Station enclosure, 

Fosberg Stoddart 54747 (US): in center of flats near N end 

of island, Fosberg &- Stoddart 54759 (US); nearby U. S. Air 

Force Station enclosure, Fosberg G Stoddart 54748 (US, holotype, BISH, A, K, BM, L, G, NY, UC, TI, BO, BRI, P, B, 

F, S, isotypes). 

Gardner Atoll: ‘Gardner Isld.” U. S. Expl. Exped. (GH). 

(Leaves small, ovate or oblongish, cymes rather slender, axillary, ending in a single glomerule. No color given): SE end 

of lagoon, Fosberg 55761 (US): W coast somewhat inland, 

Fosberg 55772 (US). 

Hull Atoll: s.L, E. H. Bryan Jr. 53 (US): small islets on N 

reef, Fosberg & Stoddart 54832 (US); near middle of S leg of 

East I., FosbeTg 5 Stoddart 54823 (US). 

Sydney Atoll: S side of atoll, lagoon shore, Fosberg 6. Stoddart 51864 (US); back of abandoned Gilbertese village site 

on \tT side of island, Fosberg G. Stoddart 54859 (US): in abandoned Gilbertese village, W side of island, Fosberg & Stoddart 54854 (US): SE shore of lagoon, Long 2588 (US): SW side 

of island, Long 2580 (US). 

Vostok Island: Long 3204 (US), 3203 (US): Anderson in 

1935 (US). 

Malden Island: SW side, Long 3382 (US); S side, Long 3385 

Jarvis Island: G m, Bergman 52 (US); Bryan 1388 (BISH). 

Starbuck Island: SJ,, Long 3302 (US); N peninsula, Long 

3319 (US), 3300 ‘(US): E end, Long 3307 (US); W beach, Long 

3351 (US). 

AUSTRALIA.-QueenSland: Capricorn Is., Heron Island, Fosberg 55063 (US). 

(US). 

Boerhavia albiflora var. powelliae Fosberg, 

new variety 

Foliis anguste ovatis vel lance-ovatis valde acutis 

Leaves narrowly ovate to lance-ovate, apices very 

Named for Miss Dulcie Powell, who has made a 

vel acuminatis. 

acute to acuminate. 

great many dissections of these difficult flowers. 

SPECIMENS EXAMINED 

WAKE ISLAND.-U. S. Expl. Exped. (GH); Wake Islet, Fosberg 34947 (US), 34454 (US). 

MARSHALL ISLANDS.-Bikini Atoll: Bikini Is., Taylor 46-1052 

(US, A): Taylor 46-1097 (US, A): Namu I., Taylor 46-1117 

(US); Rongerik Atoll: Enyvertok I., Taylor 46-1432 (US, A); 
NUMBER 39 13 

Eniwetok Atoll: Araanbiru I., Taylor 46-1337 (US); Japan 

I., Taylor 46-1295 (US); Eniwetok I., Fosberg 36700 (us); 

Rongelap Atoll: Kabelle I., Fosberg 36670 (US); Gegen I., 

Fosberg 36728 (US, holotype). 

(HAW), Young 147 (US), and Oahu, Fosberg 27127 

(US)-approach this species in their white or 

whitish flowers and few stamens, also in having 

the glands mostly in the furrows of the lower part 

of the perigone. In habit they are rather diverse 

and are, for the present, excluded until the Hawaiian Boerhauia population can be studied as a 

whole. 

Boerhavia aff. albiflora 

A group of specimens from the Hawaiian Islands 

-Laysan Island, Long 2360 (US), 2390 (US), 2388 

5. Bornhavia rubicunda Steudel and Somewhat Similar Species 

One of the most distinctive species of Boerhavia 

subgenus Boerhauia is an Asian-African species that 

has usually been called Boerhauia elegans Choisy, 

indeed an appropriate, even if incorrect, name for 

it. It does not seem particularly related to any of the 

other species known to me, though one species, 

described here as Boerhauia crispifolia Fosberg has 

been confused with it. The fruits of the two show 

considerable similarity, also with those of Boerhauia 

herbstii Fosberg, also described below. These are 

not similar enough to be called a “group” in the 

sense used above for the Boerhauia repens group, as 

their similarity principally lies in the form and 

indument of their anthocarps. It is convenient to 

treat them here together for comparative purposes. 

Technically, B. rubicunda Steudel (B. elegans 

Choisy) falls into ’ Boerhauia sect. Singuliflorae 

Heimerl, although I am by no means certain that 

it is closely related to the American members of 

that section. 

The nomenclature of Boerhauia ru bicunda illustrates the difficulty of writing a code of rules that is 

universally satisfactory. The name Boerhauia eleguns Choisy is accompanied by a reasonably wellprepared and adequate description. However, 

Choisy (1849:453) cited the somewhat earlier name 

Boerhauia ru bicunda Steudel #( 1840: 2 13) as a synonym. Examination of the original publication of 

this latter name shows that the only descriptive 

information accompanying it is the symbol Y, explained by him as meaning “perennial.” A specimen 

“Arab. fel. Boerhavia nr. 744 Herb. arab. un it.” 

is cited that is certainly the same Schimper collection cited by Choisy for B. elegans. This is borne 

out by two sheets of this collection, one in P, the 

other in US with “Herbarium Steudel” written on 

the upper margin of the label and named Boerhauia 

rubicunda Steudel. 

This symbol is certainly not an adequate description. However, the International Code of Botanical 

Nomenclature, Art. 32, dealing with conditions for 

valid publication requires only a “description,” 

not necessarily an adequate one. I see no way to 

avoid regarding B. rubicunda as validly published, 

and no way to avoid using it as the correct name for 

the species under discussion, since its type is known 

and identifiable. There is also no way to avoid 

considering B. elegans Choisy as superfluous and 

illegitimate as Choisy did not take up the earliest 

available epithet. 

Boerhuvia rubicunda Steudel 

Boerhauia rubicunda Steudel, Nom., ed. 2, 1:213, 1840.- 

Heimerl, Beitr. Syst. Nyctag., 28, 1897 [type: Arabia Felix, 

Montis Sedder, Schimper 744 (P)]. 

Boerhavia elegans Choisy, in A. de Candolle, Prodr. 13(2):453, 

1849 [nom. superfl. illegit.]. 

Erect herb, or suffrutescent below, to at least 

90 cm tall, branching at base from a woody root 

crown, stems appearing more or less glaucous, this 

appearance reinforced by short incurved puberulence, especially in lower parts, basal part of stem 

gnarled wih stubs of branches; leaves mostly basal 

and at several nodes, cauline somewhat smaller, 

sometimes with small ones singly or fascicled in 

axils, blades ovate to oblong, subfleshy or subcoriaceous, 1-1.5 cm X 0.6-1.0 cm, apex acutish to 

obtuse and apiculate, base obtuse to subcordate, 

both surfaces more or less gray-scurfy and puberulent, nerves obscure, about 3-5 on a side; petiole 

short, slender; upper ’/* or s/4 of plant paniculate, 

leafless, branches gracefully spreading, curvingdivergent, repeatedly, up to 8 or even 12 times, 

dichotomous but tending in each fork to have a 

long capillary pedicel bearing a flower, outer 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

branches of a dichotomy tending to become more 

divergent than inner; pedicels about 1 cm long, 

perigone 2-3 mm long, constricted about y3 the 

way up, appearing less strongly constricted than 

usual in the genus, but actually strongly but extremely abruptly constricted, limb narrowly campanulate, pink or striped with red especially on 

lower part, more or less pubescent especially at or 

above tips of stripes, rarely (one sheet of Schzmper 

744) glabrous, lower segment of perigone obovoid, 

with 5 glabrous ridges, grooves between them 

slightly to densely puberulent, not glandular or 

with sessile glands only; stamens 3, sometimes 4, 

rarely 5, filaments straight to somewhat curved, 

anthers exserted, suborbicular, when dehisced appearing didymous; style with a strong curve in the 

middle or rarely straight, longer than stamens, stigma 

capitate, exserted; anthocarp about 4 mm long, 

clavate to ellipsoid, round at apex, with 5 heavy, 

rounded, glabrous ridges, grooves puberulent or 

glandular with subsessile glands, in age glabrate, 

man) conspicuous white raphide bundles in grooves. 

The superficial resemblance of this species to a 

number of species of the western American genus 

Eriogonzim (Polygonaceae) is remarkable. 

Ranges from India and Pakistan to Iran, Arabia 

and eastern Africa. The plants are rather poorly 

separable into two varieties, var. rubiczinda from 

mostly rather low elevations mostly near the coast, 

from Iran to Africa, and var. stenophylla, almost 

entirely at higher elevations from India to Iran, 

and very rarely from Arabia. Many of the labels 

give localities that are hard to locate, and some may 

be variant spellings. 

Boerhavia rubicunda Steudel var. rubicunda 

This is as described above. Its leaves are ovate to 

oblong or even obovate, gray, margins tending to be 

somewhat wavy, pedicels tending to be about 1 cm 

long. 

SPECIMENS EXAMINED 

IRAN.-M. Gazy Manzal [Gazeh Manzil or Minzil] on Rude-Kaju, branch of Slop [or Selup] RiLer, about 55 km N of 

Chahbakar, Gadde 64 (LE, 2 sheets); Baluchistan, Zahedan 

Province, 18.5 mi 130 km] NE of Geh (=Nikahaht), road to 

Qasrqad, c. 1600 ft [490 m], Gray-Wilson & Hewer 302 (K). 

ARABIA.-Arabia Felix: “In rupibus montis Sedder, W. 

Schinper 744 (type collection of both B. rubicunda and B. 

elegans, P, holotype, 3 isotypes, US, 2 sheets, L, 4 sheets, LE, 

GH, NY, BM, all isotypes); Shugra, Schweinfurth I12 (US, in 

part). Oman: sl., Jayakar s. n. (BM); Muscat (Maskat), Mutrak in Wadi, 50-100 ft [15-30 m], Waring in 1958 (K); Muscat, Bent 14 (K): from Hameira (near Shinas) to Ajib, Fernandez 838 (K), 2314 (K), 2320 (K). Aden: s.1. Baudouin in 

1867 (P); Balfour in 1880 (K); Perry in 1878 (K); Hooker 92 

(K); Thornson in 1837 (K); Yerbury 34 (BM); Germain 2 (G); 

de Marchesetti s.n. (K); Khusaf Valley, 50-100 ft [15-30 m], 

Waring in 1958 (K); Goldmore Valley, Schweinfurth 61 

(BM, G, P 2 sheets): Steamboat Point, Lunt in 18934 (BM); 

Hadramaut, Jut, Kerfoot 3058 (K); Hadramaut Wadi, north 

of Saiyun, 2000 ft [610 m], Popov, Tillin 6 Gilliland 4158 (K). 

Hejaz: Jidda (Jedda, Djeddah) Kruijt 199 (L, 2 sheets): 25 

miles [40 km] NNE, Trott 1259 (K). Not located on map: 

Djair et Khara, Ehrenberg 137 (LE); Djara et X’iara Ehrenburg in 1820-1826 (K) (perhaps same as foregoing); km 32 

Mudhailif Bani Sharfa, Mosnier 2938 (P); Ad regionem inferiorum Kefr et Sidi, Fischer 159 (LE, 3 sheets, L, 2 sheets). 

A~FRICA.-sOmaliland: 5 mi [8 km] west of Berbera, Hemming 2337 (K); Guban, Berbera, Courbon 388 (P); Busteh, 10 

mi [I6 km] inland from Berbera, Bally in 1957 (K), 11856 

(G); Mt. Busteh, near Berbera, Bally 7218 (G); Wagga Mountain, Phillips in 1897 (BM); Diddin Tug Valley, Glover 6. 

Gilliland 721 (BM, K); Boundary road at Hormo (or Homo?), 

Glover & Gilliland 936 (BM, K); “Chessala”, Courbon in 

1859-1 860 (P), “Abyssinie: Golfe d’Adoulis” [ = Eritrea: 

Dulla], Courbon in 1859-GO (P). Cote Francaise des Somalis: 

Aoussa (bord des lacs), ChPdeville 667 (P). “Junction of tugs, 

44” 18’ x 9” 57”” Gillet 4346 (K) (leaves narrowly oblong, 

2.5 x 0.6 cm, distributed on young sprouts, slightly smaller 

ones fascicled in axils; much like var. stenophylla but leaves 

somewhat broader and mostly rounded at apex, petioles to 5 

mm, slender). 

EGYPT (?)-Wady Rabdeit, Jebel Elba, Shabstei F. 1728 (K) 

(very young plant, inflorescences not much developed, mostly 

in bud, 1-12 stems from branching woody caudex). 

Nrcm.-“Air-Greboun” “2005 N 0838 E,” 37@+ ft [1130 

m], Davies 71 (BM) (sterile, a strong woody caudex much 

branched at top but plant grazed down to a few cm, stubs of 

branches covered with leaves that could be of this species, 

but this would be hard to prove). 

If the latter two specimens are correctly identified, 

it suggests that B. rubicunda may extend much 

beyond its established range and that further collecting should be done in northern East Africa. 

Boerhauia rubicunda var. stenqhylla (Boissier) 

Fosberg, new combination 

Boerhavia elegans var. stenophylla Boissier, Flora Orientalis 

4:1046, 1879. 

Boerhauia elegans sensu Stewart, Annot. Cat. Vasc. PI. W. 

Pakistan 233, 1972 [not Choisy in A. de Candolle, Prodr. 

13(2):453, 18491. 

Erect, leaves elliptic to lanceolate, prominent on 

cauline nodes, with fascicles of smaller ones in 
NUMBER 99 15 

axils; inflorescence very ample, to 30 cm high, 20 cm 

wide, open, branches with tendency to curve outward, up to 10 times ramified; pedicels 1-2 cm; 

anthocarp ellipsoid, ribs smooth, grooves whitish 

papillate, raphide bundles not noted. 

Rajasthan to eastern Iran, and rarely eastern 

Arabia, tending to occur inland and at higher elevations than is usual for var. rubicunda. 

“Folia anguste lineari-lanceolata” “Hab. in siccis 

Arabiae Mascatensis l(Auch. 5250!) [corrected in 

pencil in British Museum copy to 52511. Beluschiae 

inferioris (Stocks!)” (Boissier, 1879: 1046). 

SPECIMENS EXAMINED 

INDIA.-Rajasthan: Jais a1 Mer Distr.: Mohanjar, Wadhwa 

5060 (LE, L). 

Inferra” E. Stocks [in], 1851 (G-BOISS, syntype, lectotype; 

Baluchistan: Kargoshk Kah, coast, Pierce in 1880 (K); Kalat, 

Jhal Jhao to Awaran “mountain areas,” Lamont 338 (RAW, 

G). Northwest Frontier Province: Bannu, Salt Range, Rakman 

25871 (RAW). Hyderabad: Tandojam District: 1.5 mi [2.4 km] 

from Telephone Saebony, Rotai, Jalis in 1959 (RAW). 

Province not indicated on labels: Paniala Forest, Dera Ismail 

Khan [Northwest Province] Monsi 9 (RAW); 6 mi [9.7 km] 

from T. B. Khan on way to Hyderabad, Qaisir et al. 583 

(KAR); 12 mi [19.3 km] from T. B. Khan, Northern Hills, 

Qaisir et al. 640 (KAR); 3 mi [4.8 km] Teejahban on way to 

Turbat, Aredin & Hussain 6052 (KAR), 6053 (KAR); 10 mi 

[16.1 km] before Mund, Turbat-Mund Rd., Abedin 6 Hussain 

6241 (KAR); 2 mi [3.2 km] from Pasni on way to Turbat, 

Abedin Q Hussain 6474 (KAR), 6488 (KAR); 10 mi [16.1 km] 

from Turbat on way to Awaran, Ali et al. 1540 (KAR, 2 

sheets); 30 mi [48.3 km] from Turbat on way to Hoshab, 

Abedin & Hussain 6550 (KAR); 34 mi [54.7 km] from Awaran 

on way to Hoshab, Qaisir 797 (KAR); 1.5 mi E2.4 km] from 

Teejaban on way to Hoshab, Abedin & Hussain 6049 (KAR); 

46 mi [74 km] from Thaljhus on way to Bela, Abedin 6 

Qaisir 7493 (KAR), 7494 (KAR); Lau Pass, 8 mi [12.9 km] 

from Bela on way to Awaran, Ghafoor & Qaisir I (KAR); 17 

mi [8.4 km] from Bela on way to Awaran, Qaisir et a1 747 

(KAR); 30 mi [48.3 km] from Bela on way to Awaran, Abadin 

6. Qaisir 7465 (KAR); 11 mi 117.7 km] before Awaran, Bela, 

Awaran Hills, Ah’ et al. 1450 (KAR); 8 mi [12.9 km] from 

Bela on way to Awaran, Abedin 2133 (KAR); 10 mi [16.1 km] 

from Sehwan on way to Jamshoro, Farooqi Q Qaisir 2192 

(KAR); 10 mi [16.1 km] from Hoshab on way to Panjgur, 

Qaisir et al. 1066 (KAR) 10 mi [16.1 km] from Nasurabad on 

way to Tump, Turbat-Mund Road, Abedin & Hussain 6176 

(KAR), 6177 (KAR); 50 mi [80.5 km] from Mund on way to 

Suntsar, Ghafoor Q Qaisir 243 (KAR); 21 mi [33.8 km] from 

D. G. Khan on way to Fort Munro, Abedin 6. Hussain 9643 

(KAR), 9636 (KAR); 28 mi r45.1 km] from D. G. Kahn on 

way to Fort Munro, Abedin Q Hussain 9661 (KAR); 16 km 

W of Bela, E side of Gan Pass, Lamond 29f (G); Bikaner, 

Popov G.P. 631412 (BMJ. 

PAKISTAN.-sCind [=Sind]: Stocks S.n. (G, H, L, E). “Bel. 

ImN.-Prov. Balucistan: inter Khash (Vasht, Kwash) et 

Iranshahr (Bampur), Montes Kawandar, 150&1600 m, Rechinger et al. 3986 (G, BM, K). 

ARABIA.-Arain, “Haidwan”, Philby 23 (BM); Mascate, 

Aucher-Eloy 5251 (G-BOISS, syntype, US, BM, K, isosyntypes). 

The following specimens have leaves somewhat 

broader than is typical for the variety stenophy Zla, 

but narrower than usual for var. rubicunda, otherwise as in var. stenophylla. 

PAKISTAN.-S. of Prohdab, Mile 76, Nushki Road, 4000 ft 

[I220 m], Crookshank 141 (K); 3 mi [4.8 km] from Teejahban 

on way to Turbat, Abedin 6 Hussain 6054 (KAR), 6055 (KAR); 

15 mi [24.1 km] from Teejahban on way to Turbat, Abedin Q 

Hussain 6057 (KAR); 25 mi [40.2 km] from Teejahban to Turbat, Abedin Q Hussain 6081 (KAR); 5.5 mi [8.8 km] from 

Mund on way to Turbat, Ali et al. 920 (KAR), 1068 (KAR); 33 

mi [53.1 km] from Mund on way to Teeran Fezath, Ali et al. 

1776 (KAR); 4 mi [6.4 km] before Hushab, Awaran-Hushab 

Road, Ali et al. 1461 (KAR), I465 (KAR); 12 mi [19.3 km] 

from Hushab on way to Awaran, Ghafoor & Qaisir 74 (KAR); 

34 mi [54.7 km] from Awaran on way to Hushab, Qaisir et al. 

797 (KAR); 11 mi [17.7 km] before Awaran, Bela-Awaran 

Road, Ali et al. 1449 (NY, KAR), 1450 (KAR); 6 mi [9.7 km] 

from Thaljao on way to Awaran, Abedin 6 Qaisir 7474 

(KAR). 

Boerhavia crispifoliu Fosberg, new species 

Boerhavia elegans auct. Aldabrae [not Choisy, in A. de Candolle, Prodr. 13(2):453, 18491. 

Herba gracilis puberula, radicem crassiuscula, 

caulibus pluribus, ascendentibus infra ramosis, distaliter sensim tenuiter paniculatis, foliis parvis 

tenuiter petiolatis margine crispatis, cymis axillaribus desunt, floribus parvis 3-6 glomeratis bracteolatis perigonio campanulato lobis emarginatis, 

staminibus 2-3(-4$, curvatis, anthocarpio anguste 

elliptic0 5-costato costis glabris apice rotundato 

intervalis glandulosis vel glabris albo-lineolatis. 

Tap-root to 8 (-12) mm thick, branched to, rarely, 

nearly simple; several stems from slightly thickened root crown, prostrate to somewhat ascending, 

branched, especially near base, puberulent, grayish to white below, purplish distally, internodes 

from very short to rarely as much as 6 cm long in 

leafy portions, longer in lower part of inflorescence, 

stem leafy in proximal 5-30 cm, changing gradually 

or somewhat abruptly from leafy stems to rather 

bare panicle; leaves ovate to oblong, obtuse to acute, 

mostly under 10 X 5 mm, rarely to 20 X 10 mm, 

upper surface usually purplish, to brownish green, 

beneath pale, margins usually strongly crispate, 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

blades containing abundant cystoliths or crystals 

and a few raphide bundles, petiole 2-6 mm long, 

rarely longer; axillary cymes none, panicle slender, 

capillary or almost so distally, 10-30 cm long, puberulent, main axis with branches on alternate sides at 

alternate nodes in lower part, 2 leaf-like bracts at 

each lower node, distal nodes tending more and 

more to develop 2 branches, with axis tending to be 

reduced, bracts on distal nodes reduced to ovate 

ciliate scales; flowers sessile in glomerules of 3-6, 

subtended by ovate-lanceolate, glandular-ciliate, 

scale-like bractlets, central flower opening first; entire flower about 2.5 mm long, very strongly constricted, part above constriction campanulate to 

cylindric, caducous, 1.5 mm long, puberulent, 

especially in distal half, which has longitudinally 

arranged raphide bundles, slightly lobed, tending 

to be erose or ciliolate at tips in bud, sinuses between the (4 or) 5 lobes glandular ciliolate, lobes 

emarginate; stamens 2-3(-4), slightly exserted, or included, filaments strongly curved, anthers broader 

than high, each half almost orbicular, both ends 

emarginate; style subequal with stamens, curved 

in upper part, stigma strongly depressed globose, 

(peltate); anthocarp or fruiting perigone narrowly 

ellipsoidal to subclavate, about 2 mm long, ends 

rounded to subtruncate, 5-ribbed, ribs broad, smooth, 

intervals glandular to almost or quite glabrous. 

Most parts of the plant are marked by white shortlinear or elliptic raphide bundles, especially conspicuous in the flowering perianth and in the intervals between the ribs of the anthocarp. 

This species is notably variable in habit, especially in the degree to which the inflorescence is 

developed, and in leaf size and shape. In flower 

and fruit morphology it is rather uniform, varying 

only in distribution and amount of puberulence on 

the flowering perigone, slightly in fruit shape, and 

considerably, from glabrous to densely but shortly 

glandular puberulent in the intervals between the 

ribs of the anthocarp. The crispate character of the 

leaf-margins is much more evident in living than 

in pressed and dried specimens. 

SPECIMENS EXAMINED 

ALDABRA.-w’est I.: near settlement, Stoddart 965 (US); 

Fosberg 49471 (US, K); inland from settlement near edge of 

mangrove, Fosberg 48713 (US, K); Bassin Cabri, Hnatiuk 

731744, 731745, 731746, 731747, 731748, 731750, 731751, 731752, 

731753, 731754 (all US). 

Middle I.: Camp at Passe Gionnet, Renuoize 1161 (US, K); 

East Channel, Renuoize 1282 (US, K). 

South I.: 0.7 km WNW Cinq Cases Camp, Fosbmg 49223 

(US, K); near Cinq Cases Camp, Fosberg 48873 (US, holotype, 

isotypes, K, MO, L); Fosberg 48892 (US); Fosberg 49188 (US), 

Fosberg 48976 (US, K); Renoize 781 (US, K); Hnatiuk 731936 

(US); Fosberg 49152 (US, K); 1-2 km south of Cinq Cases 

Dune, Fosburg 49170 (US, K); Fosberg 49171 (stems and leaves 

green, flowers white) (US, K); Takamaka, Hnatiuk 731931, 

731940, 731941, 731942, 731943, 731944 (all US); Renvoize 868 

(K, ZIS): south of Takamaka, Renuoize 11231 (K, US); Hadoul 

Point, Renvoize 891 (K, US). 

Common fairly generally on pitted limestone and 

coral gravel near the coast on South Island; on 

platin inland on West Island but rare. 

ABNORMAL FORM 

One plant of a strange Boerhauia (Fosberg 48929) 

was found that I do not understand. It seems almost 

like a caricature of a Boerhavia. In appearance it 

resembles a small prostrate Euphorbia of subgenus 

Chamaesyce, with fine, intricately branched stems, 

internodes 1 cm long or less, the entire plant about 

20 cm across, more densely puberulent than B. 

crispifoh, purplish, the leaves like those of B. crispifolia in size and shape, but thinner and flatmargined, not noticeably crispate, to about 5 x 3 

mm, bracteal ones reduced but still foliaceous, to 

about 3 X 1.5 mm; inflorescences not strongly distinct from vegetative part of plant, to at most 3-4 

cm long, showing some tendency for one branch at 

a node to be suppressed, cymes 2-4 times branched; 

flowers in glomerules of 2-3(-4), perigone about 2-3 

mm long, strongly puberulent, especially distally 

on buds and perigone lobes, noticeably marked with 

raphide bundles, perigone only weakly constricted, 

the lower part only obscurely 5-ribbed, distal part 

funnelform-campanulate, or urceolate above, lobes 

ovate, not at all corolloid, lobes ovate, short, densely 

puberulent, ovary lanceoloid, about ’/s the length 

of the flower, stamens 2, subequal with ovary, anthers not well developed; no tendency toward 

development of anthocarps, but perhaps the plant 

was too young. 

This plant is so different from B. crispifolia that 

it is only its association with populations of that 

species, the occurrence (so far as is known) of only 

a single individual, and the rudimentary nature of 

the stamens that makes me associate this plant with 

B. crispifolia. 

The change in habit of B. repens with attack by 
NUMBER 39 17 

the fungus Albugo platensis is well known, but 

although the present specimen was examined at the 

time of collection and later, no Albugo pustules 

were found. 

After the above plant had been studied and 

described, a shipment of Boerhavia from Aldabra 

arrived from S. A. Renvoize (Royal Botanic Gardens, 

Kew), including a number of B. crispifolia, several 

B. repens var. maris-indici, and three specimens 

resembling to some extent the plant described 

above. Two of these, Renuoize 769 and 981, were 

from the same general area, Cinq Cases, as Fosberg 

48929, and the other, 869, was from “Takamaka 

turn off.” No. 869 is much more condensed than 

48929, but has the same sort of scarcely constricted, 

vase-shaped flowers. No, 769 is of several unattached 

branches, all much more elongate than 48929. The 

main stems of these have the nodes 3-6 cm long, but 

all but one piece have most of the branches condensed and the flowers abnormal in the same way 

as in 48929. One branch has an almost normal appearance, but some of the flowers are normal, some 

abnormal. 

No. 981 is of several pieces, all extremely slender, 

the smaller branches filiform with the general 

appearance of 48929 but more loose and elongate, 

the leaves ovate, obtuse, not at all crispate, not more 

than 5 mm long. The distal parts are rather irregularly paniculate, bracteate, puberulent, flowers in 

clusters of 3 or 4, pedicellate, buds notably clavate, 

perigones clavate, with no constriction whatever, 

2.5-3.5(-4) mm along, tube slender below, gradually 

dilated upward, lobes short, incurving, notably 

pubescent, the tube tending to absciss from receptacle but to persist around the ovary. 

The presence, on no. 769, of both abnormal and 

normal flowers shows that these plants are indeed 

Boerhauia crispifolia, but I have no suggestion as 

to a possible cause for these abnormalities. Comparable, but not similar, abnormalities have been found 

in Aldabra populations of at least two other, unrelated, genera, Nesogenes and Euphorbia. 

Boerhavia herbstii Fosberg, new species 

Herba prostrata decumbens foliis triangulo-ovatis 

infra pallidis, pedunculis gracilibus foliis valde 

superantibus, partibus plantae laxe paniculatis ad 

sexies ramosis, floribus perigoniis parte proximo 

obovato glabro parte distale ventricoso staminibus 

styloque perigonio excedens, anthocarpio 5-costato 

subglabro ellipsoideo. 

Prostrate herb, stems apparently decumbent, 

pink, to 60 cm long or more, finely and thinly 

puberulent, internodes 4-15 cm long, very sparsely 

branched except in inflorescence, nodes with a 

transverse ridge bearing a line of pubescence; leaves 

triangular-ovate, to 4 x 2 cm, light green above, 

very pale beneath, subglabrous, margin slightly 

scabrous, apex bluntly acute, base obtuse to subtruncate, petiole 0.4-1 cm long, puberulent; nodes 

bearing long slender peduncles 6-7.5 cm long, just 

at side of leaf axil on alternate sides, umbellately 

branched at summit, branches bearing glomerules of 

flowers; distal half or more of plant leafless, glabrous or nearly so, very openly paniculate, the 

panicle branches ascending, two inflorescence 

branches at each node of main axis and of secondary 

axis, one branch always much larger than other, 

larger ones on alternate sides, ramified up to 6 

times, penultimate ramification tending to be umbelloid, each ray bearing a glomerule of 2-4 flowers, 

glomerules rarely proliferous, or tending to be racemulose, each branch of inflorescence subtended by 

a bract, these successively smaller, and more narrowly lanceolate, penultimate ones lance-subulate, 

those subtending glomerules and flowers ovate, subulate-acumina te, straw-colored-hyaline with dark 

purple acumen, glandular-ciliolate; perigone 2-2.5 

mm long, constricted about at middle, lower half 

green, obovoid, shallowly 5-ribbed, glabrous or with 

slight puberulence in grooves, limb very broadly 

cylindric-campanulate, ventricose at base, glabrous 

except for slight puberulence distally, pink but drying yellowish pink; stamens 3, anthers exserted, 

opened cells orbicular; style curved, stigma well exserted; anthocarp 3-3.5 mm long, ellipsoid, somewhat constricted at base, apex rounded-subtruncate, 

5 ribs heavy, glabrous, intervals exceedingly minutely glandular, appearing glabrous, with a few 

very narrow raphide bundles longitudinally placed. 

HAWAIIAN IsANos.-Lanai: Hulope Beach, on rock ledge 

above ocean, common in this area, 31 March 1976, D. Herbst 

5798 (US, holotype). 

This species does not much resemble any of the 

known Hawaiian forms, all relegated to varieties of 

B. diflusa L. by St. John (1973:154) following 

Heimerl (1937), unless it be B. diflusa var. gymnocarpa Heimerl. The photo of the type (Heimerl, 
18 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

1937, pl. 3) somewhat resembles it, but the description does not fit very well and is not sufficiently detailed to justify either its status as a variety of B. 

diflicra L. or inclusion in it of the present plant. 

Boerhauia herbstii in all likelihood is not really 

related to either B. rzibicunda Steudel or B. crispifolia Fosberg, but it does have an ample leafless 

panicle as do these two, and the fruits of all three 

are closely similar. 

The axillar) peduncles of B. herbstii, below the 

panicle. suggest a place in the Boerhauia repens 

group, where it fits also geographically. \$'hen that 

group has been adequately studied this species may 

find a place there. 
Literature Cited 

Brown, R. 

1810. Prodromus florae novae Hollandiae. Volume 1, 

viii + 592 pages. London. 

1849. Nyctaginaceae. In A. de Candolle, Prodromus Systema tis Nu turalis Regn i Vegeta bil is, 13 (2) :425-458. 

Choisy, J. D. 

Codd, L.E.W. 

1966. Notes on Boerhauia in Southern Africa. Bothalia, 

9(1):113-121. 

Delile, A. Raffeneau1824. Description de I'Egypte: Histoire naturelle. Volume 

Forster, G. 

19, 472 pages; Atlas, 62 plates. Paris. 

1786. FIoruEae insularum Australium Prodromus. 103 

pages. Gottingen. 

1837. (Zephyritis Taitensis.) Enumeration des plantes decouvertes par les voyageurs, dans les Iles de la Societe, principalement dans celle de Taiti. Annales 

des Sciences Naturelles, second series (Botanique), 

Guillemin, J.B.A. 

7:177-192. 

Heimerl, A. 

1889. 

1897. 

1934. 

1937. 

Nyctaginaceae. In Engler and Prantl, Die naturlichen Pflanzenfarnilien 111, l(b): 14-32. 

Beitrage zur Systematik der Nyctaginaceen. 40 pages. 

Wien. 

Nyctaginaceae. In Engler and Prantl, Die naturlichen Pflanzenfamilien, edition 2, 16(c):86-134. 

Nyctaginaceae of Southeastern Polynesia and Other 

Pacific Islands. Occasional Papers of the Bernice P. 

Bishop Museum, 13:2747, plates 1-3. 

1973. List and Summary of the Flowering Plants in the 

Hawaiian Islands. 519 pages. Hawaii. 

Schinz, H. 

1897. Zur Kenntnis der Flora der Aldabra-Inseln. Abhandlungen herausgegeben von der Senckenbergischen 

naturforschenden GeseElschaf t, 21 (1): 77-91. 

St. John, H. 

Setchell, W. A. 

1924. American Samoa, Part I: Vegetation of Tutuila 

Island; Part 11: Ethnobotany of the Samoans: Part 

111: Vegetation of Rose Atoll. Carnegie Institution 

of Washington Publications, 341:l-275. 

1972. International Code of Botanical Nomenclature. 426 

pages. Utrecht. 

1909. The Allioniaceae of the United States, with Notes on 

Mexican Species. Contributions from the United 

States National Herbarium, 12:303-389. 

1911. The Allioniaceae of Mexico and Central America. 

Contributions from the United States National Herbarium, 13:377-430. 

Stafleu F. A., et al., editors 

Standley, P. C. 

Steudel, E. G. 

1840. Nomenclator Botanicw. 2nd edition, volume 1, 852 

pages. Stuttgart and Tiibingen. 

Taylor, W. R. 

1950. Plants of Bikini and Other Northern Marshall 

Islands. xv + 221 pages. Ann Arbor, Michigan. 

19 
Index 

Abronia L., 1 

Albugo, 6 

Aiiulocaulis Standley, 1, 2, 3 

Boerhavia aff. alliiflora Fosberg, 13 

albiHora Fosberg, 8, 11 

albiflora Fosberg var. albiHora, 12 

albiflora tar. pwelliae Fosberg, 12 

caribaea Jacquin, 5 

coccinea Miller, 5 

crispifolia Fosberg, 9, 15, 16, 17, 18 

diffusa L., 1, 2, 4, 5, 6, 7, 8, 17, 18 

diffusa I ar. eudiffusa f. psammophila Heimerl, 9 

tliffusa {ar. gymnocarpa Heirnerl, 17 

diffusa Tar. tetrandra (Forster f.) Heimerl, 10 

elegans Choisy, 9, 13 

elegans yay. stetiophylla Boissier, 14, 15 

herbstii Fosberg, 17, 18 

hirsuta Willdenow, 5 

rnutabilis R. Brown, 5, 6, 7, 8 

mutabilis var. glandulosa, 7 

paniculata Richard, 5 

pubescens R. Brown, 6 

repens L., 1, 4, 3, 6, 8, 13, 16, 18 

repens var. rnaris-indici Fosberg, 9, 17 

platensis, 17 

repens var. minor Delile, 9 

repens var. pubescens (R. Brown) Fosberg, 6, 8, 10 

repens L. \ar. repens, 10 

rubicuiida Steudel, 13 

rubicuiida Steudel var. rubicunda, 14 

rubicunda var. stenophylla (Boissier) Fosberg, 14, 15 

Boerhavia sect. Adeiiophorae Heimerl, 3 

sect. Claiatae Heimerl, 3, 4 

sect. Micranthae Heimerl, 3, 4 

sect. Pterocarpon Heimerl, 3 

sect. Pyramidatae Heimerl, 3, 4 

sect. Senkenbergia (Schauer) Heimerl, 3 

sect. SinguliHorae Heimerl, 3, 4, 13 

sect. Solenanthae Heimerl, 3 

sect. Spicatae Heimerl, 3, 4 

tetrandra Forster filius, 5, 6, 8, 10 

viscosa Lagasca S: Rodriguez, 5 

Cornmicarpus Standley, 1, 2, 3 

Cyphomeris Standley, 1, 2, 3 

Liiidenia Martens S: Galeotti, 2 

Mirabilis L., 1 

Pisonia L., 1 

Senkenbergia Schauer, 2, 3 

Senkenbergia Gaertner, Xieyer S: Scherbius, 2, 3 

Tinantia Martens S: Galeotti, 2 

20