BULLETIN OF THE UNITED STATES NATIONAL MUSEUM

CONTRIBUTIONS FROM THE UNITED STATES NATIONAL HERBARIUM

VOLUME 37

SYSTEMATIC PLANT STUDIES

ALBERT C. SMITH, BENJAMIN C. STONE, Epwin B. Bartram,
Grorce W. GIL_ett, Peter H. RAVEN, and Marie L. Farr

SMITHSONIAN INSTITUTION PRESS e WASHINGTON, D.C. e 1969

Publications of the United States National Herbarium

The United States National Herbarium, which was founded by
the Smithsonian Institution, was transferred in the year 1868 to the
Department of Agriculture and continued to be maintained by that
department until July 1, 1896, when it was returned to the official
custody of the Smithsonian Institution. The Department of Agri-
culture, however, continued to publish the series of botanical reports
entitled “Contributions from the United States National Herbarium,”
which it had begun in the year 1890, until, on July 1, 1902, the Na-
tional Museum, in pursuance of an act of Congress, assumed respon-
sibility for the publication. The first seven volumes of the series were
issued by the Department of Agriculture.

Frank A. Taynor,
Director, United States National Musewm.
Il

CONTENTS

Part 1. Studies of Pacific Island Plants, XVII: The genus
Geniostoma (Loganiaceae) in the New Hebrides,
Fiji, Samoa, and Tonga. By Albert C. Smith and
Benjamin C. Stone. (Published January 5, 1962) .
Part 2. Mosses of the Eastern Highlands, New Guinea,
from the 6th Archbold Expedition, 1959. By
Edwin B. Bartram. (Published September 21,

1965) 2...

Part 3. Studies of Pacific Island Plants, XVIII: New and
noteworthy flowering plants from Fiji. By Albert
C. Smith. (Published May 4, 1967) ......

Parr 4. The genus Cyrtandra in Fiji. By George W. Gillett.
(Published February 6,1968)......... 107-159

Part 5. A revision of the genus Camissonia (Onagraceae).

By Peter H. Raven. (Published February 5, 1969). 161-396

Parr 6. Bredin-Archbold-Smithsonian Biological Survey of
Dominica: Myxomycetes from Dominica. By

Marie L. Farr.* (Published January 31,1969). . 397-440

*Errata: Line 22, page 425, should read “Stemonitis hyperopta .. .

III

43-67

69-106

”

UNITED STATES NATIONAL MUSEUM

CONTRIBUTIONS FROM THE UNITED States NaTIoNAL HERBARIUM

VoLumE 37, Part 1

STUDIES OF PACIFIC ISLAND PLANTS, XVII
THE GENUS GENIOSTOMA (LOGANIACEAE)
IN THE NEW HEBRIDES, FIJI], SAMOA, AND TONGA

By Apert C. Smirn and
BENJAMIN C. STONE

BULLETIN OF THE UNITED STATES NaTIONAL Museum

SMITHSONIAN INSTITUTION ¢ WASHINGTON, D.C, © 1962

STUDIES OF PACIFIC ISLAND PLANTS, XVII!
THE GENUS GENIOSTOMA (LOGANIACEAE)
IN THE NEW HEBRIDES, FIJI, SAMOA, AND TONGA

ALBERT C. SMITH AND BENJAMIN C. STONE

Introduction

Specimens of Geniostoma, an essentially tropical Old World genus of
woody flowering plants of the family Loganiaceae, have been accumu-
lating in world herbaria, including that of the U.S. National Museum,
over a period of several decades, during which no serious attempt to
understand relationships within the genus has been made. Specific
identifications have usually proved difficult, and the genus as a whole
is in need of a complete systematic revision. Primarily in order to
facilitate an understanding of the Fijian specimens, which are now
reasonably abundant in herbaria, we have undertaken a regional re-
vision within a small portion of the generic range, extending from the
New Hebrides eastward to Samoa. It has been customary to throw
together a large part of the Pacific Geniostomata under the name G.
rupestre J. R. & G. Forst., in spite of the fact that Gilg & Benedict
(in Bot. Jahrb. 56: 542. 1921) considered it a local endemic of the
New Hebrides. Only in New Caledonia, Samoa, and to a lesser extent
in Micronesia have studies of Geniostoma progressed to a point where
identifications can be made with any degree of assurance. For the
Malaysian region in the broad sense there exist no dependable treat-
ments, @ situation that one may hope to see corrected as work on the
Flora Malesiana progresses.

As is so often the case in “difficult”’ genera, the older descriptions
are mostly inadequate for an understanding of the identity of species.
Fortunately we have been able to see types or isotypes of most taxa
thought to occur in our region. Especially valuable for study has
been type material of G. rupestre, the type species of the genus.
Geniostoma has in general been correctly understood and delimited,

1 This paper, based on research partially supported by a grant from the National Science Foundation,
continues the series under the same title by the first author. The place of publication of nos. X-XIV in the

series was noted in connection with no. XV, in Contr. U.8. Nat. Herb. 30: 523-575. 1953. No. XVI was
published in Journ. Arnold Arb. 34: 97-124. 1953,

1

2 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

but we believe that concepts of some of its species have been unreason-
ably broad. Sixteen species are now believed to occur between the
New Hebrides and Samoa, of which six are here described as new.
The other species of our region are redescribed on the basis of avail-
able material and older descriptions, and we append notes on distri-
butions, local names, and relationships.

In the course of this study the pertinent herbarium material of sev-
eral institutions has been examined and is here cited, with the indi-
cated abbreviations: Arnold Arboretum of Harvard University (A);
Bernice P. Bishop Museum (Bish); British Museum (BM); Gray
Herbarium of Harvard University (GH); Royal Botanic Gardens,
Kew (K); New York Botanical Garden (NY); University of Cali-
fornia at Berkeley (UC); and U.S. National Herbarium (US). We
are indebted to the administrators of these herbaria for the opportu-
nity of studying specimens under their care.

HISTORICAL SKETCH

The genus was described by J. R. & G. Forster in their “Charac-
teres Generum,” in 1776, as was also their type species, G. rupestre,
and was based upon material obtained by them on the island of Tanna,
in the New Hebrides, during Cook’s second voyage. The generic name
is formed from the Greek words genias (beard) and stoma (mouth) and
of course should be treated as neuter (cf. Int. Code Bot. Nomencel.
Rec. 75A. 1956); when necessary we have corrected epithets given a
feminine form. Additional species of Geniostoma were described in
the early part of the nineteenth century from the Mascarene Islands,
Java, Sumatra, the Philippines, New Zealand, Australia, and the
Marianas Islands. For the most part, however, specimens obtained
in Polynesia and Melanesia were identified with G. rupestre. In 1880
H. Baillon (in Bull. Soc. Linn. Paris 1: 247-248) published a list of
New Caledonian species of the genus. Various other species described
in Geniostoma, including those from the New World, were later shown
to belong to other genera; the Hawaiian species, for instance, have
been removed to the closely related genus Labordia Gaud. The first
treatment of the genus that was relatively broad in regard to geography
and in addition was acute and analytical was that of T. Valeton (in
Bull. Inst. Bot. Buitenz. 12: 1-28. 1902), who studied the species of
the vicinity of Buitenzorg and also some of those from more distant
regions. Although he did not attempt a monographic revision, he
employed a number of new criteria in his specific distinctions and
was apparently the first to recognize the sexual dimorphism of sev-
eral of the species. During the present century the number of
putative species has substantially increased and now approaches 100,
but the arrangement of the species within the genus has been neg-

SMITH & STONE—PACIFIC ISLAND PLANTS, XVII 3

lected. There almost certainly remain undescribed species in New
Guinea and the other large islands of Malaysia and Melanesia. In
Micronesia, and in the Society Islands and Polynesia generally, most
or all of the species may have been discovered. According to A.
Guillaumin (Fl. Anal. Syn. Nouv.-Caléd. 285. 1948), none of the New
Caledonian species occur outside that area except for G. rupestre. As
in other genera, the islands between Fiji and New Guinea have
disclosed fewer Geniostomata than might be anticipated, indicating
the need of further field work there.

CRITERIA OF USE IN SPECIFIC DELIMITATION

Among vegetative features of Geniostoma, the unusual nature of the
stipules of two Fijian species, G. macrophyllum and G. stipulare,
deserves mention. In these species the stipules are large, adnate to
the margins of the petioles and forming foliaceous sheaths that are
soon disrupted, leaving traces of tissue or obvious linear scars on the
petiole. Both of these species are further distinguished by several
other significant characters; they have cauline or ramuline inflores-
cences and leaves large for the genus. Their relationship may possibly
be with G. caulocarpum K. Schum., of New Guinea. With the excep-
tion of G. macrophyllum and G. stipulare, the species treated by us
have intrapetiolar stipules of a type more characteristic of the genus,
forming a short sheath scarcely exceeding 2 mm. in length.

Certain species of our region are characterized by having markedly
quadrangular or even 4-winged branchlets; the Samoan G. fleisch-
mannit has the branchlets narrowly alate, as do some species of New
Caledonia. Geniostoma insulare has its branchlets at least in the
ultimate internodes sharply quadrangular. However, most of our
species have essentially terete branchlets, which in the juvenile inter-
nodes may be compressed or subquadrangular.

Other vegetative characters of potential taxonomic usefulness
include the kind and degree of pubescence and the size and shape of
leaf-blades. Pubescence of the vegetative parts in general is limited
to the innovations, although in certain species it is persistent; in the
Fijian G. calcicola, for instance, a short, dense indument covers both
surfaces of the leaf-blades, a character noted otherwise among species
of the vicinity only in G. crassifolium Benth., of the Isle of Pines.
Leaf-size and shape are characters at best of secondary importance.
The leaf-blades may be fairly consistently large (in G. macrophyllum
and @. stipulare) or small (in @. stenocarpum), but in most species-
complexes the range of size is too great to be of taxonomic value. In
leaf-shape there is a somewhat wider array of types and perhaps a
slightly greater taxonomic value, but, as Valeton cautioned in 1902,

4 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

leaf-shape alone is not dependable in Geniostoma and must be used in
conjunction with other characters.

One species of our area, the Samoan @. biseriale, has essentially
sessile leaves that at once distinguish it from others of the region.
Other species with similarly subsessile leaves are G. sessile Kanehira
(Palau), G. rapense F. Br. (Rapa), and G. clavatum J. W. Moore
(Raiatea), but they are diverse in other respects and obviously do not
form a natural species group.

The basic structural unit of the inflorescence is a dichasium, gener-
ally of the cymose type; this is much modified, and finds its extreme
diversity in the multiflowered, glomerate, cauline inflorescences of
G. macrophyllum, which may bear up to 200 flowers, these basically
organized in congested triflorous dichasia. At the other extreme may
be mentioned the highly reduced, simple cyme of G. calcicola, with its
5 flowers on moderately long pedicels. In some New Caledonian
species the inflorescence is reduced to a single flower. The most
common type of inflorescence in the genus is a moderately open, com-
pound dichasium with perhaps 5 to 30 flowers, those of the distal axes
generally in triflorous dichasia, but those lower on any inflorescence-
branchlet sometimes irregular, single, or paired. The inflorescences in
leaf-axils (characteristic of most species) may consist of a single cyme
but frequently are fasciculate (usually at the first node of the peduncle
but sometimes in the axil itself) in pairs or sometimes in threes. In
discussing the inflorescence of Geniostoma, Valeton discerns a racemose
and a paniculate type, which he terms bostryx and cyme. According
to Rickett (in Bull. Torrey Bot. Club 82: 430. 1955), these are but
modifications of the dichasium. The verticillaster described by Vale-
ton seems also a modification of the dichasium, with an obsolete or
absent peduncle and fasciculate rachises. Corymbose panicles,
according to Valeton present in G. avene Valeton and G. australianum
F. v. Muell., are also basically dichasial. The main rachis is branched
into opposite laterals, which are subtended by bracts, each node thus
being bibracteate; on the irregular (not triflorous) branchlets there
may be single unpaired bracts, or a pair of bracts without branchlets
in their axils. In most inflorescences the calyces are subtended by
bracteoles. In general, the nature of the inflorescence is diagnostic
for only a few unusual species (e.g. in our area @. gracile, G. macro-
phyllum, G. insulare, and perhaps G. calcicola), but still it provides a
valuable supplementary character in delimiting taxa.

The pioneering work of Valeton in Genitostoma has elucidated the
incipient and actual sexual dimorphism in various species and has
explained the diverse types of gynoecia. Dimorphism is expressed
in some species by a considerable difference in stylar length, and in
addition by the form and pubescence of the stamens. In general,

SMITH & STONE—PACIFIC ISLAND PLANTS, XVII 5

it would appear that flowers with long styles are those with fertile
anthers, while those having short styles have sterile anthers. In the
species of our area, however, marked dimorphism is not obvious, and
short-styled flowers frequently appear to have fertile anthers. The
close approach to dioecism that Valeton discussed in relation to
certain Javanese species is lacking in the Fijian-Samoan area, at least
as far as can be ascertained from gross microscopic observations.
However, in certain cases, e.g. in G. vitiense, there is a fairly marked
heterostyly, accompanied by anther differences. Stylar length is
thus seen to be a character related to sexual dimorphism, and as such
it is hardly appropriate for use in specific distinctions. In this regard,
one may wonder whether the species from Saipan described as G.
longistylum by Gilg is merely a long-styled form of G. micranthum
DC. or G. hoeferi Gilg & Benedict; students of Micronesian species
should review Geniostoma with this in mind.

The presence or absence of trichomes on the anthers, found to be
of some use in the species discussed by Valeton, is apparently of little
or no consequence in our species. The shriveled thecae of the sterile
anthers mentioned by Valeton are not obvious in specimens from our
region; at least we have noted no anthers that appear to be sterile,
although of course it is possible that non-fertile pollen or reduced
fertility exist in some cases. In respect to floral dioecism in the
species of our area, there seems to be little or no sterilization of the
gynoccia. Incipient tendencies seem to be toward hermaphrodite vs.
pistillate individuals, but these tendencies are of no present taxonomic
use.

Significant specific differences are found in the venation of the
corolla, although these differences are difficult to express verbally
without resorting to lengthy and complex phrases. We have included
an illustration for each species here treated (Plate 1) showing the
corolla split between two lobes and flattened out (two lobes are omitted
from each sketch for brevity). In one basic type of corolla-venation,
illustrated by G. confertiflorum, G. clavigerum, and G. uninervium,
each corolla-lobe is supplied by a single main nerve, which may pro-
duce a few lateral branches, but these are so few or so inconspicuous
that the impression of a one-nerved lobe is given. A second type of
venation is more complex and could be described as somewhat cande-
labriform. In this type the main central nerve supplying each lobe
has one or more laterally produced pairs of branches, these either
subparallel or arcuately ascending and in turn usually laterally
branched. Among our species the nerves anastomose and form retic-
ulations only in G. dictyoneurum, in other species being free in the
lobes. Whether the main vein supplying each lobe sends out its
first branches high or low in the corolla provides a character of some

6 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

use. A division into three veins at or near the base of the corolla-tube
frequently occurs in such species as G. rupestre and G. macgregorii;
elsewhere, as in G. stenocarpum and G. insulare, the main vein usually
branches first near the summit of the corolla-tube or in its throat.
However, there is no real consistency within species as to this venation-
pattern, and it should be used with caution as a differentiating char-
acter, even though incipient trends are apparent. For the most part
the venation is readily observed in corollas of relatively thin texture;
in species with a more carnose corolla that organ must be boiled and
the outer tissue separated with dissecting tools, the venation then
being apparent. The venation of the calyx, which usually has three
nerves ascending into each lobe, provides a less useful character in
species differentiation.

In most species of Genzostoma in our area the corolla is glabrous
without, but all species observed by us have an obvious indument in
the throat. The trichomes of the corolla-throat are white, thin,
and retrorse, but as they sometimes extend to the inner surface of
the lobes they tend to become spreading or even ascending. The
length of the corolla-throat trichomes provides a very useful and
consistent character. Among our species these trichomes are either
“short” (0.1-0.4 mm. long) or “long” (0.4-0.9 mm. long); while
such a dimensional difference is not striking, there appear to be no
species in which a decision is difficult. Of course detection and
measurement of so minute a character are facilitated by adequate
microscopic and micrometric equipment, but even with a hand lens
one can usually decide whether the trichomes are ‘‘short’’ or “long.”

Pubescence similar to that within the corolla-throat extends to
the filaments; rarely the anthers also bear trichomes of the same
type. The gynoecium in Geniostoma is characteristically glabrous,
but one of our species, G. macrophyllum, differs strikingly in having
its ovary copiously hirtellous on the distal surface. The style only
rarely is weakly hirtellous. The stigmas form a globose or clavate
to obovoid capitulum that is minutely papillate, with papillae essen-
tially conical but sometimes so slender as to appear setulose. It
may be added here that the radial nature of the stigma, with the
papillae in marked radial groups as shown in the Forsters’ plate
(t. 12), has been noted in G. vitiense.

A marked and diagnostic character found in certain Malaysian
species, that of mucronulate or apiculate anthers, is of less importance
in our area. However, in some instances (e. g. in G. samoense var.
parviflorum) there does appear to be a very minute mucro on the
anther. There is also some indication that apiculation of the anthers
may be correlated with incipient sexual dimorphism, as mentioned
and illustrated under G. vitiense and G. macgregorii. However,

SMITH & STONE—PACIFIC ISLAND PLANTS, XVII 7

all of our species fundamentally have obtuse or merely acute anthers,
unlike such species as G. arboreum (Bl.) Kuntze (=G@. haemospermum
Steudel) and G. miquelianum Koorders & Valeton as described by
Valeton.

The capsules in our species are usually ellipsoid to ovoid or obovoid,
briefly apiculate by the persistent base of the style. However, a
tendency toward essentially globose capsules is seen in G. insulare
f. sphaerococeum and G. dictyoneurum. The capsules are subtended
by persistent calyces, and all of them dehisce into two recurved valves
to expose the seminal mass. The degree of recurvature of the
valves, mentioned by Valeton, seems to us of little or no diagnostic
use.

In summarizing this discussion of specific criteria, we agree with
earlier students of Geniostoma in finding none of them entirely satis-
factory. In approximate order of usefulness in differentiating the
species of our region may be mentioned: type of stipule, presence
or absence of ovary-indument, length of trichomes of the corolla-
throat, type and abundance of vegetative pubescence, position and
type of inflorescence and number of flowers, corolla-venation, stigma-
shape, capsule-shape, and terete vs. angled or winged branchets.
Various combinations of these characters and others of lesser value
permit the recognition of taxa that seem to possess stability. That
our delimitation of taxa is somewhat subjective is obvious, but this
is inevitable in a study where experimental methods are impracticable.

Systematic Treatment

Geniostoma J. R. & G. Forst. Char. Gen. 24. ¢. 12. 1776.

As presently known, @eniostoma is composed of about 95 species of
trees and shrubs. It is exclusively Old World in distribution, occur-
ring from the Mascarene Islands through Malaysia north to Kyushu
(Japan) and the Marianas and East Caroline Islands, south to Austral-
ia and New Zealand, and east to the Society Islands. The species at
the extreme north and south limits of the genus are @. ligustrifolium
A. Cunn. (New Zealand), @. petiolosum C. Moore & F. v. Muell.
(Lord Howe Island), and G@. glabrum Matsumura (Japan). Many
species are ecologically correlated with the coastal lowland vegetation
of tropical islands, but increasing numbers are becoming known from
the cool, wet forests of the high interiors of such large islands as New
Guinea.

Characteristic features of the genus are the presence of stipules, the
simple opposite leaves, the axillary or cauline dichasial inflorescences,
the tubular 5-lobed calyx and corolla, the insertion of the 5 stamens
on the corolla-tube near its throat, the bilocular ovary, the numerous
anatropous or nearly amphitropous ovules, and the bivalved, capsular,

8 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

dehiscent fruits with seeds embedded in an eventually exposed pulpy
mass. The persistence of the seed-mass between the valves of the
dehisced capsule is noteworthy; the seeds remain on persistent funicles
and are enclosed by the pulp, which dries to a thin pellicle; the placen-
tation persists until at last the mass drops or is broken from the exocarp
and then splits into two halves, corresponding to the contents of each
locule of the ovary. The seeds are ovate-angulate, usually concave
at the hilum and minutely papillate; the embryo, embedded in a small
amount of smooth endosperm, is short-oblong, usually more than half
the length of the seed, and very shortly divided at apex into two
cotyledons.
Key to the Species

Stipules large, forming foliaceous sheaths 1-3.5 cm. long, adnate to margins of
petioles and caducous, leaving obvious linear scars; leaves large for the genus,
the blades often exceeding 30X15 em. (but sometimes smaller) ; inflorescences
borne on trunks or branches or defoliate branchlets; Fijian species.

Ovary copiously hirtellous on distal surface; corolla-lobes glabrous (except at
base within) ; inflorescences cauline, glomerate to elongate (to 25 cm. long)
and with as many as 200 flowers; leaves glabrous . . 1. G. macrophyllum

Ovary glabrous; corolla-lobes densely hirtellous within; inflorescences ramuline,
dichasial, 15—20-flowered; leaf-blades (as well as other young parts) copious-
ly pubescent beneath at least on principal nerves, the trichomes predomi-
nantly 3-celled, about 0.56 mm. long .. . .... . .2 G. stipulare

Stipules intrapetiolar, small, forming inconspicuous ‘sheaths no more than 2 mm.
long, the petioles without obvious stipular scars; leaves comparatively small,
the blades not exceeding 24 13 cm. and usually much smaller; inflorescences
axillary or rarely on defoliate branchlets.

Leaves obviously petiolate, the petioles normally at least 4 mm. long, the
blades various at base (acute or attenuate to rounded) but not cordate.

Corolla-venation comparatively simple, each lobe supplied by a single nerve
with a few short faint laterals, or essentially unbranched; trichomes of
the corolla-throat obvious, 0.6—0.7 mm. long; Fijian species.

Stigma oblong-clavate, obovoid, or ellipsoid; each corolla-lobe supplied by
a faint nerve with 1 or 2 usually unbranched laterals on each side.
Leaves comparatively large, the petioles 9-20 mm. long, the blades
usually 10-23%5-12 ¢em.; inflorescences 5-—11-flowered; corolla-
lobes with a single pair of faint lateral nerves arising from the
midnerve . . . . . 3. G. confertiflorum
Leaves smaller, the petioles 5-91 mm, ‘long, the blades 3-8 X 1-3.6 cm.;
inflorescences 3-5 (rarely to 9)-flowered; corolla-lobes with 2 pairs
of lateral nerves arising from the midnerve . . 4. G. clavigerum
Stigma globose; each corolla-lobe supplied by a single dark relatively
conspicuous nerve with brief and obscure (or usually without) laterals;
petioles 5-9 mm. long, the leaf-blades ovate, usually about 10-12X
3-5em.. . .. .. . 5. G. uninervium

Corolla-venation comparatively! complex, each" lobe supplied by a nerve that
produces in the corolla-tube (or at its base) one or several pairs of obvious
lateral nerves, these ascending together into the lobe and usually re-
branched.

SMITH & STONE—PACIFIC ISLAND PLANTS, XVII 9

Trichomes of the corolla-throat short, 0.1-0.4 mm. long.

Indument of young parts of plant copious, although minute, composed
of setulose trichomes 0.05-0.2 mm. long, these obvious on the
young branchlets, petioles, and often on ventral surfaces of leaf-
blades, especially on costae; inflorescence-branches similarly pilose;
Fiji to Tonga and Niue... . .... . 6G. vitiense

Indument lacking throughout (except w ‘ithin corolla).

Inflorescences elongate, with branchlets and pedicels 7-17 mm. long,
3-7-flowered; corolla-lobes glabrous and smooth, each supplied
by a nerve that branches high in the tube or in the throat; stigma
clavate-obovoid; capsules ellipsoid, 5-9 4-7 mm.; Samoa.

7. G. gracile

Inflorescences comparatively compact, with branchlets and pedicels
2-6 mm. long, 5—15(—20)-flowered; corolla-lobes pilose proximally
or pustulose within, each supplied by nerves diverging in lower
part of tube or at its base; stigma globose; capsules narrower,
7-13X3-4.5 mm.; Fiji and Tonga. . . . . 8. G. macgregorii

Trichomes of the corolla-throat comparatively long, 0.4-0.9 mm. long.

Leaf-blades puberulent on both surfaces (hairs obvious, about 0.3 mm,
long), elliptic, 3.5-5X 1.5-2.5 cm., rounded at apex; Fiji.

9. G. calcicola

Leaf-blades glabrous, or pilose only on the ventral costae, variously
shaped but most often acute to acuminate at apex.

Corolla-venation complex, each lobe supplied by a nerve branched in
lower portion of tube, the branches rising into the lobe and there
forming a reticulum of short veinlets; leaf-blades thin in
texture, ovate, 7-123.5-7 cm., usually rounded at base and
apex; Fiji... . .... . 10. G. dictyoneurum

Corolla-venation not reticulate, the veinlets in the lobes free.

Flowers comparatively large, the corolla 4.5-5 mm. long, each lobe
supplied by a dichotomous main nerve with several laterals;
leaf-blades oblanceolate or narrowly elliptic, 4-12 1.5-3.5
cm.; branchlets quadrangular and narrowly winged distally;
Samoa . foe .... . LLG. fleischmannii

Flowers comparatively small, "the corolla 2.5-3.5 mm. long, each
lobe supplied by a single main nerve with 1 or 2 pairs of laterals;
branchlets not winged.

Leaf-blades lanceolate or narrowly ovate, 4-7.5X1.2-2.7 cm.;
each corolla-lobe ‘Supplied by a nerve divided in the throat;
Fiji... . ..... 12. G. stenocarpum

Leaf-blades various in shape, usually exceeding 8X3 cm.

Inflorescence-axes minutely puberulent with trichomes no
more than 0.03 mm. long; vegetative parts glabrous;
leaf-blades ovate to broadly ovate, 8-18X4-9 cm. (often
smaller in var. parviflorum); Samoa. . 13. G. samoense
Inflorescence-axes glabrous (or, if puberulent in some forms of
no. 15, then the young vegetative parts also puberulent).
Leaf-blades acuminate or obviously cuspidate at apex; each
corolla-lobe supplied by a nerve usually branching at or

near base of tube; New Hebrides and Santa Cruz
Islands, and perhaps westward . . . . 14. G. rupestre

10 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Leaf-blades rounded to obtuse (or sometimes broadly acute
or bluntly cuspidate) at apex; each corolla-lobe supplied
by a nerve usually branching in upper part of tube; Fiji
to Samoa, Tonga, and Niue. .... 15. G. insulare

Leaves subsessile, the petioles 2 mm. long or less, the blades subauriculate,
cordate, or subcordate at base; plant glabrous throughout except for the
usual corolla-throat trichomes, these 0.3-0.4 mm. long; Samoa,

16. G. biseriale

1, Geniostoma macrophyllum Gillespie in Bishop Mus. Bull. 91: 25, fig. 28. 1932.
PLATE 1, FIGURE 1; PLATE 2, FIGURES 1-3

Small tree to 10 m. high, glabrous throughout except for some
floral parts, the trunk to 25 cm. in diameter; branchlets stout, distally
4-8 mm. in diameter and compressed-quadrangular, becoming terete
and up to 2 cm. in diameter; stipules forming a foliaceous sheath up
to 15 mm. long, distally acute, laterally united with the alate petiole-
margin; leaves large for the genus, the petioles broadly alate, not
distinct from the blade, the blades chartaceous, ovate to elliptic or
sometimes obovate, 22-55 cm. long, 9-25 cm. broad, gradually decur-
rent at base and produced laterally along the petiole into a broad
decurrent wing, obtuse to bluntly cuspidate at apex, entire at margin,
the costa stout, conspicuous, sulcate above, prominently elevated
beneath, the secondary nerves 11-20 pairs, arcuate-ascending, irreg-
ularly anastomosing near margin, slightly sulcate above, raised
beneath, the veinlets prominulent on both surfaces; inflorescences
cauline on old wood, often near base of trunk, variable in size and
shape, when young somewhat glomerate, at length more or less
pendent on elongated branching woody axes bearing numerous con-
gested triflorous dichasia, sometimes up to 25 cm. long and bearing
as many as 200 flowers, freely branching, with deltoid, acute, incon-
spicuous bracts and small ultimate bracteoles less than 1 mm. long;
flowers subsessile in congested groups of 3; calyx cupuliform, about
2 mm. in diameter, lobed to the middle, the lobes deltoid-ovate,
acute, 1-3 (—5)-nerved, the nerves free; corolla 3-3.3 mm. long,
divided to middle, the tube 1.5-1.7 mm. long, pilose at throat, the
trichomes about 0.6 mm. long, dispersed in a uniform band around
throat down to middle of tube, the lobes deltoid-ovate, about 1.5
1.5 mm., glabrous without and within except at extreme base at
throat, the venation comparatively simple, each corolla-lobe supplied
by a single usually 3-branched vein, the branches arising midway
in the tube; stamens with short glabrous or sparsely ventrally bar-
bate filaments about 0.3 mm. long, the anthers broadly ovoid, about
0.5 mm. long, sparsely hirtellous at base, acute at apex; ovary de-
pressed-globose, conspicuously flattened distally, about 0.6 mm. high
and 1.6 mm. in diameter, laterally glabrous but copiously hirtellous
on distal surface with erect hairs about 0.2 mm. long, the style gla-

SMITH & STONE—PACIFIC ISLAND PLANTS, XVII 11

brous, 0.4-0.5 mm. long, the stigma depressed-globose, papillate,
about 0.5 mm. broad; capsules glabrous, ovoid-ellipsoid, apiculate
by the stylar remnant, 8-11 mm. long.

Tyrer Locauiry: Near Nasinu, Naitasiri Province, Viti Levu, Fiji;
the type is Gillespie 3638, cited below.

DistripuTIon: Endemic to Fiji, and known thus far only from
Viti Levu, Vanua Levu, and Ovalau, as a slender tree 3-10 m. high,
with a trunk 4-10 (-25) cm. in diameter, occurring in wet forests at
altitudes from near sea level to 850 m. The inflorescences, which
consist of dense many-flowered glomerate or elongate irregular di-
chasia, are borne on the trunk, often less than a meter from the
ground. The corolla is noted as greenish white, within hirtellous
with whitish hairs; other floral parts are whitish. The capsule at
maturity is black, with orange seeds. No local names or uses are
indicated.

FIJI: Vir1 Levu: Mba: Hills between Nandala and Nukunuku Creeks, along
trail from Nandarivatu toward Lewa, Smith 6168 (A, Bish, K, US). Serua: Hills
along coastal road, Greenwood 976A (A). Namosi: Hills east of Wainikoroiluva
River near Namuamua, Smith 8899 (US); hills bordering Wainavindrau Creek,
near Wainimakutu, Smith 8552 (US); hills east of Navua River, Greenwood
976 (A). Naitasiri: Near Viria, Parks 20461 (Bish, UC); along Tamavua-Sawani
road, Setchell & Parks 15071 (Bish); near Nasinu, Gillespie 3638 (Bish type,
GH, UC); near Suva Pumping Station, Degener & Ordonez 13746 (A, Bish, K,
NY). Naitasiri or Rewa (near border): Mt. Kombalevu, Parks 20317 (Bish,
UC, US); near Tamavua, Jeoward 65 (K); woods near Tamavua, Gillespie 2109
(Bish). Rewa: Slopes of Mt. Korombamba, Gillespie 2297 (Bish); Suva Ditch
Trail, Bryan 370 (A, Bish); Princes Road, Meebold 16791 (Bish); Central Road,
Suva, Tothill 613 (Bish, K), 617 (K). Vit Levu, without locality, Parks 20870
(Bish, UC). Vanua Levu: Thakaundrove: Southern slope of Korotini Range,
below Navitho Pass, Smith 573 (Bish, GH, K, NY, UC, US). Ovatau: Near
summit of main range west of Levuka, Gillespie 4449 (Bish, UC); Mt. Tana
Lailai, near Levuka, Graeffe in 1864 (K).

Geniostoma macrophyllum, together with the new species G. stipulare,
is isolated in relation to the other species of our area in gross and
obvious features as well as in technical characters. These two species
exhibit certain vegetative characteristics in common: unusually large
leaves; enlarged, petiolar stipules; and cauliflory or ramuliflory.
In particular, the nature of the stipules, which in both species are
adnate to the margins of the petiole and in @. macrophyllum united to
the basal part of the blade also, marks these species as amply distinct
from their Fijiian congeners, and indeed from most of the other
Pacific species which have been the subject of our attention.

Geniostoma macrophyllum is abruptly distinct from G. stipulare,
and displays very little intraspecific variation of consequence. Only
one specimen (Jeoward 65) is unusual, in that its leaves are markedly
obovate and more or less cuspidate at apex, and in addition, the
branches of the inflorescence are excessively elongated; yet the

12 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

flowers are indistinguishable from those ofthe “other cited specimens
of G. macrophyllum.

The phenomenon of cauliflory is more marked in G. macrophyllum
than in other species of the genus, with the exception of G. caulocarpum
K. Schum., of New Guinea, the inflorescences often appearing close
to the ground on the main trunk. In G. stvpulare the inflorescences are
mostly ramuline, and in addition are fewer-flowered and more strictly
cymose. In all other Fijian Geniostomata, the inflorescences are either
strictly axillary among the leaves, or both axillary and ramuline on
juvenile branches.

While we believe that the two species here discussed form a distinct
section within Geniostoma, we refrain from proposing a formal name,
since this should follow a monographic study of the entire genus.

2. Geniostoma stipulare Smith & Stone, sp. nov.
PLATE 1, FIGURE 2; PLATE 2, FIGURES 4-9

Arbor parva ad 5 m. alta, innovationibus et ramulis atque stipulis
foliorum petiolis et laminis subtus rufo-pilosis, pilis 3-cellularibus
circiter 0.5 mm. longis; ramulis subteretibus vel subcompressis apicem
versus 2-5 mm. diametro; stipulis gemmam circumcludentibus per
margines petiolorum fere ad basim laminae adnatis anguste oblongis,
1-3.5 cm. longis, caducis; foliis magnitudine disparibus, petiolis
1.5-4.5 cm. longis supra sulcatis subtus rotundatis, laminis ovatis
ellipticis vel obovatis, 6.5-34 cm. longis, 3-18 cm. latis, basi cuneato-
decurrentibus, apice acuminato-cuspidatis, margine integris, nervis
majoribus subtus copiose et areolis sparsius pilosis, costa supra leviter
sulcata subtus valde elevata, nervis secundariis utrinsecus 8-15
utrinque paullo elevatis, venulis subtus prominulis; inflorescentiis
secus ramos vel ramulos defoliatos enatis conferto-glomeratis sub-
sessilibus dichasialibus 15-20-floris, pedunculo ad 1 mm. longo,
pedicellis 1-6 mm. longis, bracteolis forma et textura calycis lobis
similibus; floribus (videtur pistillatis) in dichasiis congestis trifloris
dispositis; calyce cupuliformi, lobis orbiculari-ovatis 1-1.5 mm. longis
paulo latioribus 3—5-nervatis; corolla 4.5-5.2 mm. longa ad medium
lobata, tubo lobisque extus glabris, tubo intus basi glabro faucibus
pilis albis circiter 0.8 mm. longis hirtello, lobis ovatis intus praeter
apicem dense hirtellis, nervatio comparate multiplice, corollae quoque
loba nervis 3 e basi tubi in lobam adscendentibus praedita, quoque
nervo laterali faucibus semel diviso; staminibus in speciminibus
nostris forsan abortivis, filamentis deltoideo-ligulatis circiter 0.8 mm.
longis pilosis, antheris lanceolato-oblongis apice acutis circiter 1.3
mm. longis et 0.5 mm. latis basi breviter hirtellis, thecis fortasse
sterilibus; ovario glabro depresso-globoso 1-1.6 mm. alto 2.5-3 mm.
lato, stylo filiformi 2.5-3.5 mm. longo, stigmate globoso diametro

SMITH & STONE—PACIFIC ISLAND PLANTS, XVII 13

circiter 1.1 mm. papilloso; capsula ovoidea 9-10 mm. longa circiter
4 mm. lata apice apiculata.

Type in the U.S. National Herbarium, Nos. 2192146 and 2336118, collected
among hills east of the Navua River, near Nukusere, Province of Serua, Viti
Levu, Fiji, alt. 100-200 m., November 2, 1953, by A. C. Smith (No. 9148).
Duplicates to be distributed.

ADDITIONAL SPECIMENS EXAMINED:

FIJI: Vim1 Levu: Naitasiri or Namosi: Mt. Naitarandamu, near summit,
Gillespie 3364 (Bish, UC). Vanua Levu: Thakaundrove: Mt. Ndikeva, in
forest on eastern buttress, Smith 1860 (Bish, GH, K, NY, UC, US).

Disrrisution: The few specimens known of this species indicate
that it is a Fijian endemic restricted to Viti Levu and Vanua Levu.
It is a shrub or small, slender tree up to 5 m. high, with relatively
large leaves, for the genus, which are sparsely but rather uniformly
pubescent on the lower surfaces; it has"ramuline;inflorescences con-
sisting of congested, subsessile dichasia. The corolla and other floral
organs are white, except for the stigma, which is pale yellow.

Loca NAME: On the label of Smith 1860 the common name is given
as muskarimba, but this is questionable, since the name often refers
to members of the genus Cyrtandra (Gesneriaceae), some species of
which in fact resemble G. stipulare in foliage and superficially in
fruit.

This new species, while varying considerably in the size of its
leaves, sometimes attains a leaf-size approaching or equalling that of
G. macrophyllum—two or three times larger than the leaves of most
other Fijian Geniostomata. However, the pubescence of the leaves
and of the inner surfaces of the corolla-lobes, the glabrous ovary,
the pilose filaments, the relatively large flowers, and the unusual
stipules effectively characterize this species. The inflorescences are
noteworthy in that they are borne on older branches, being much
contracted dichasia bearing relatively few flowers.

That the relationship of G. stipulare is to G. macrophyllum is indi-
cated by the large, petiolar stipules and the position of the inflores-
cences. ‘Together they form a somewhat isolated group; nonetheless,
they are nearly as distinct from each other as they both are from their
other Fijian congeners. Even the type of trichome that G. stipulare
produces is distinct; it is usually a three-celled hair, the cells being
elongate, and on drying it becomes flattened, the cells then seeming
oriented in different planes. ‘These peculiar trichomes, however, are
limited to the leaves and innovations; those of the corolla-throat are
of the more ordinary type, cylindric and apparently nonseptate.

At mature branchlet-nodes the stipules are lacking, but their oc-
currence is indicated by the long scars that are left, one on each margin
of the petiole, from the base to a point definitely below the blade,
but about three-fourths as long as the petiole. Apparently the stipules

14 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

are split by the expansive growth of the young leaves, and the rem-
nants are caducous.

There is possibly a third undescribed species of Geniostoma of this
relationship present in Fiji as yet represented only by a few fragments
which are mounted, together with a leaf of G. macrophyllum, on a
single sheet (Degener 14027, from Vanua Levu, in the herbarium of
the Arnold Arboretum). If the remaining leaf, which manifestly
does not represent G. macrophyllum, and the inflorescences do in fact
belong together as representing one plant, there seems little doubt
that an undescribed species still awaits adequate collecting. The
leaf, which is hirtellous beneath with the same three-celled type of
trichome found in G. stipulare, is as large as those of the type of that
species, with a long slender petiole displaying the two linear stipular
scars also characteristic of G. stipulare (terminating before the base
of the blade), but the blade is basally auriculate. The inflorescences,
which are greatly elongated and appear to be cauline, bear numerous
small congested flowers like those of G. macrophyllum. However,
they exhibit glabrous ovaries and decidedly mucronulate anthers, in
the former character differing from G. macrophyllum, and in the latter
from G. stipulare.

3. Geniostoma confertiflorum Smith & Stone, sp. nov.
PLATE 1, FIGURE 3; PLATE 2, FIGURES 15, 16
Arbor parva 3-10 m. alta praeter corollae partem interiorem ubique
glabra, ramulis apicem versus compressis et 2-5 mm. diametro demum
teretibus; stipulis intrapetiolaribus in vagina rotundata circiter 1.5
mm. longa apice minute mucronulata conjunctis; foliis magnitudine
moderatis, petiolis anguste alatis 9-20 mm. longis, laminis ellipticis
vel ovatis, 7-24 cm. longis, 3-13 cm. latis, vulgo 10-23 & 5-12 cm.,
basi cuneatis vel rotundatis et decurrentibus, apice acuminatis,
margine integris, costa supra sulcata subtus elevata, nervis secundariis
utrinsecus 8-12 subtus prominulis, venulis obscuris; inflorescentiis
axillaribus dichasialibus 5—11-floris, pedunculo communo subnullo
haud 1 mm. longo, rhachidibus primariis 1-3 mm. longis bibracteolatis,
bracteolis circiter 0.6 mm. longis, pedicellis 2-3 mm. longis in calyeem
dilatatis; floribus visis videtur hermaphroditis; calyce cupuliformi
circiter 2.5 mm. longo, lobis ovatis circiter 1.5 mm. longis 3—5-nervatis;
corolla 44.7 mm. longa extus glabra, tubo 2-2.4 mm. longo intus
basim versus glabro faucibus pilis albis circiter 0.6 mm. longis hirtello,
lobis ovatis 22.3 mm. longis erectis vel subpatentibus intus praeter
basim glabris, nervatio simplice, corollae quoque loba nervo unico
subobscuro per tubum ad lobam adscendente praedita, quoque nervo
venulum unicum subsimplicem inconspicuum faucibus ferente; stami-
nium filamentis deltoideo-ligulatis 0.3-0.4 mm. longis, antheris anguste

SMITH & STONE—PACIFIC ISLAND PLANTS, XVII 15

ovoideis circiter 1 mm. longis ubique glabris apice minute apiculatis
vel subretusis; ovario depresso-globoso glabro diametro circiter 1
mm., stylo 1-1.3 mm. longo, stigmate oblongo-clavato 1-1.3 mm.
longo minute papilloso; capsula late ellipsoidea 7-9 mm. longa apice
apiculata, seminibus circiter 1 mm. longis cumulo pulposo flavo-
aurantiaco immersis.

Type in the U.S. National Herbarium, No. 2191173, collected in hills north of
Wainavindrau Creek, between the Korombasambasanga Range and Mt. Naitara-
ndamu, Province of Namosi, Viti Levu, Fiji, alt. 250-450 m., September 14, 1953,
by A. C. Smith (No. 8488). Duplicates to be distributed.

ADDITIONAL SPECIMENS EXAMINED:

FIJI: Virr Levu: Serua: Mbuyombuyo, near Namboutini, Tambualewa 15593
(A, Bish, K, NY, UC, US); hills west of Waivunu Creek, between Ngaloa and
Korovou, Smith 9240 (US). Naitasiri: Near Tamavua, Gillespie 2059 (Bish,
NY, UC, US); near Nasinu, Gillespie 3637 (Bish, UC, US). Rewa: Near Lami,
Gillespie 2317 (Bish, UC); Central Road, Suva, Tothill 611 (K). Ovatnau:
Summit of Mt. Ndelaiovalau and adjacent ridge, Smith 7381 (US), 7621 (US).
Naavu: Slopes of Mt. Ndelaitho, on northern spur toward Navukailangi, Smith
7878 (US); hills east of Herald Bay, inland from Sawaieke, Smith 7744 (US).

DistrisuTion: Endemic to Fiji, thus far known from Viti Levu,
Ovalau, and Ngau. The species is noted as arborescent, becoming 3
to 10 m. high, with a trunk up to 5 cm. in diameter; the flowers are
white or pale greenish, with the anthers and stigmas sometimes pale
yellow. The seeds are noted as yellow. The habitat is in woods or
forest, often on ridges or in rocky areas, at elevations from near sea
level to 626 m. No local names have been recorded.

This new species, as well as the next one, G. clavigerum, is char-
acterized by its clavate stigma; in other Fijian species the stigma is
globose or subglobose. In addition, both these species and a third,
G. uninervium, are similar in that the nervation of their corolla-lobes
is relatively simple, consisting of a sparsely or almost unbranched
single central nerve.

Of the specimens cited, those from Ngau (Smith 7744, 7878) and
one from Viti Levu (Gillespie 2317) exhibit rather large leaf-blades,
which are thin in the former but coriaceous in the latter. Other
specimens display considerably smaller leaves, which in respect to
size differ little from those of the other Fijian species. Also, an
unusual condition is present in Gillespie 2059, of which the styles
are minutely and sparsely hirtellous rather than glabrous. However,
with this one exception, the floral characters of all the specimens
agree in size, pilosity, shape, and other characters.

Geniostoma confertiflorum is a species not closely related to those
often confused with G@. rupestre, and it is nearly as well marked a species
as G. macrophyllum or G. stipulare. In superficial characters, such as
the average size and shape of the leaves, however, it resembles such

597008—61——2

16 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

species as G. rupestre, G. macgregorii, and perhaps glabrate forms of
G. vitiense—all species of a different group, with globose stigmas and
complex corolla-venation. More significantly, G. confertiflorum is
similar in general appearance to G. clavigerum and to G. uninervium.
The following new species, G. clavigerum, differs from G@. confertiflorum
chiefly in its small chartaceous leaves, but also in corolla-venation.
From G. uninervium the present new species is more easily distin-
guished by the fainter corolla-venation.

4. Geniostoma clavigerum Smith & Stone, sp. nov.
PLATE 1, FIGURE 4

Arbor parva ad 10 m. alta praeter corollae partem interiorem
omnino glabra, ramulis subteretibus apicem versus 1-2 mm. diametro;
stipulis intrapetiolaribus in vagina rotundata 1-1.5 mm. alta con-
junctis; foliis comparative parvis, petiolis 5-9 mm. longis non alatis,
laminis chartaceis ovatis vel elliptico-ovatis, 3-8 cm. longis, 1-3.6 cm.
latis, basi obtuso-cuneatis, apice acutis vel subacuminatis, margine
integris, costa supra paulo sulcata subtus elevata, nervis secundariis
utrinsecus 5-9 utrinque prominulis, venulis obscuris; inflorescentiis
axillaribus dichasialibus 3-5 (—9)-floris, pedunculo communo ad 1 mm.
longo apice bibracteato, axibus secundariis 1—5 mm. longis ad nodos
bibracteatis, pedicellis 1-5 mm. longis infra calycem dilatatis et
bracteolatis, bracteis bracteolisque anguste ovatis 0.4-1 mm. longis;
floribus (visis videtur hermaphroditis) in dichasiis trifloris dispositis;
calyce cupuliformi, lobis ovatis 1-1.5 mm. longis acutis minute
ciliolatis 3-5-nervatis; corolla breviter tubulosa extus glabra 4-5 mm.
longa, tubo 2—2.5 mm. longo intus basim versus glabro faucibus pilis
albis reflexis circiter 0.6 mm. longis hirtello, lobis erectis vel sub-
patentibus glabris vel basi parce pilosis, nervatio subsimplice, corollae
quoque loba nervo unico inconspicuo per tubum adscendente et fere
faucibus cum nervulis utrinque 2 lateralibus praedita, nervulis in lobas
adscendentibus haud vel obscure ramulosis; staminium filamentis
ligulatis, antheris anguste ovoideis circiter 1 mm. longis; ovario
depresso-globoso diametro circiter 1 mm., stylo filiformi 1-1.2 mm.
longo, stigmate clavato obovoideo vel ellipsoideo circiter 1 mm. longo
papilloso; capsula ellipsoidea vel ovoidea 9-10 mm. longa apiculata.

Type in the herbarium of the Bernice P. Bishop Museum, collected on slopes
near the base of Mt. Nanggaranambuluta [Lomalangi], near Nandarivatu,
Province of Mba, Viti Levu, Fiji, alt. 1,050 meters, in November 1927, by J. W.
Gillespie (No. 3916). Duplicate at UC.

ADDITIONAL SPECIMENS EXAMINED!

FIJI: Virr Levu: Mba: Tholo-i-Nandarivatu, Gibbs 724 (BM, K); Mt. Na-
nggaranambuluta [Lomalangi], Gillespie 3782 (Bish, UC), 4334 (Bish, NY, UC).

DistrinuTion: Endemic to Fiji, known only from northern Viti
Levu, in the elevated area near Nandarivatu at about 1,000 m. or

SMITH & STONE—PACIFIC ISLAND PLANTS, XVII 17

somewhat higher. Records indicate the species as a small tree 4-10 m.
high, with white odorless flowers, occurring in dense wet forests.

Geniostoma clavigerum is very close to the preceding, G. conferti-
florum, and appears to be a smaller-leaved, smaller-cymed plant.
The more complex corolla-venation distinguishes the species, but the
smaller leaves and fewer-flowered cymes also appear to be constant.
The relatively large flowers and clavate stigmas (whence the name),
as well as the corolla-venation, are features which distinguish this
species from its other Fijian congeners.

5. Geniostoma uninervium Smith & Stone, sp. nov.
Puate 1, FIGURE 5; PLATE 2, FiauREsS 10-14

Arbor parva ad 10 m. alta praeter corollae partem interiorem ubique
glabra vel innovationibus ramulis junioribus stipulis petiolis laminae
costisque dorso minute rufo-puberula mox glabrata, ramulis subtere-
tibus vel leviter compressis apicem versus 1-2 mm. diametro; stipulis
intrapetiolaribus in vagina rotundata 1-1.5 mm. alta conjunctis;
foliis magnitudine modecratis, petiolis 5-9 mm. longis non alatis,
laminis chartaceis ovatis, 4.5-15.5 cm. longis, 2-7 cm. latis, plerumque
10-12 X3-5 cm., basi rotundatis vel obtuse cuneatis et paulo decur-
rentibus, apice acuminatis, costa supra leviter sulcata subtus elevata,
nervis secundariis utrinsecus 5-9 utrinque prominulis, venulis sub-
obscuris; inflorescentiis axillaribus coarctatis dichasialibus 3-9-floris,
pedunculo obsoleto haud 1 mm. longo apice bibracteato axes 1-3
principales gerente, axibus 1-3 mm. longis apice bibracteatis 1-3-floris,
pedicellis 2-4 mm. longis plerumque ebracteolatis infra calycem haud
dilatatis; calyce cupuliformi, lobis deltoideis circiter 1.51.5 mm.
minute ciliolatis 3-nervatis, nervo medio fusco lateralibus pallidis;
corolla circiter 4 mm. longa urceolata fere ad medium lobata, tubo
intus basim versus glabro faucibus pilis albis 0.6-0.7 mm. longis
hirtello, lobis demum reflexis, nervatio simplice, corollae quoque loba
nervo unico fusco conspicuo lateraliter haud ramuloso praedita;
staminium filamentis deltoideo-ligulatis circiter 0.4 mm. longis
pilosis, antheris glabris anguste ovatis circiter 0.7 mm. longis minute
apiculatis; ovario subgloboso circiter 1.2 mm. diametro apice depresso,
stylo filiformi circiter 1 mm. longo, stigmate globoso papilloso circiter
0.7 mm. diametro; capsula ellipsoidea vel obovoidea circiter 10
mm. longa.

Type in the U.S. National Herbarium, No. 2191135, collected in hills north
of Wainavindrau Creek, between the Korombasambasanga Range and Mt.
Naitarandamu, Province of Namosi, Viti Levu, Fiji, alt. 250-450 m., September
11, 1953, by A. C. Smith (No. 8487). Duplicates to be distributed.

ADDITIONAL SPECIMENS EXAMINED:

FIJI: Virt Levu: Namosi: Trail between Nanggarawai and Saliandrau, Gillespie
3217 (Bish, US). Tailevu: Hills east of Wainimbuka River in vicinity of Ndakui-

18 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

vuna, Smith 7099 (US). Rewa: Southeastern slope of Mt. Korombamba, Gillespie
2295 (A, Bish, UC, US). Vanvua Levu: Thakaundrove: Mt. Mariko, Smith 461
(Bish, GH, K, NY, UC, US). Ovatau: Hills west of Lovoni valley, on ridge
south of Mt. Korolevu, Smith 7527 (US). Taveuni: Summit of Uluingalau,
Smith 896 (Bish, GH, K, NY, UC, US).

DistrispuTion: Endemic to Fiji; known thus far from Viti Levu,
Vanua Levu, Ovalau, and Taveuni. The plant is arborescent, be-
coming 5-10 m. high, and occurs at elevations of 100 to 1,200 m. in
dense forest or thickets. The corolla and filaments are white or
greenish white and the anthers white or yellow.

Locat NAMES: ‘‘Mboimboinda” or ‘“‘kavikaloa”’ (Smith 7099). The
former name is a rather general one for species of Geniostoma and
refers to the characteristic foetid odor.

The relationships of this species are apparently with the two
preceding ones, all three sharing a relatively simple pattern of nerva-
tion of the corolla-lobes. Geniostoma uninervium differs from both
G. confertiflorum and G. clavigerum in its globose stigmas and also in
the darker corolla-venation. In the simplicity of this venation it
differs from all its Fijian congeners.

6. Geniostoma vitiense Gilg & Benedict in Bot. Jahrb. 56: 542.1921; A. C.Sm.
in Sargentia 1: 97. 1942; Yuncker in Bishop Mus. Bull. 220: 216. 1959.
PuaTE 1, FIGURE 6; PLATE 2, FIGURES 21-23; PLATE 3, riauREs 12-15

Geniostoma crassifolium var. glaberrimum Benth. in Journ. Linn. Soc. Bot. 1:
97. 1857.

Geniostoma rupestre var. puberulum A. Gray in Proc. Amer. Acad. 4: 33,
nomen. 1859, in Bonplandia 10: 37, nomen. 1862.

Geniostoma rupestre sensu Seem. FI. Vit. 164, 1866; non J. R. & G. Forst.

Shrub or tree to 12 m. high, the innovations and all young vegetative
parts densely pubescent with stiffly erect rufous or cinereous setulose
trichomes 0.05-0.2 mm. long, these forming a close and fairly per-
sistent indument on the branchlets, petioles, and ventral side of the
costa, and often also on the ventral surface of the blade, the leaves
and branches finally glabrate; branchlets somewhat compressed-
quadrangular and 1-4 mm. in diameter distally, becoming at length
terete; stipules intrapetiolar, forming a short subtruncate sheath
1-1.5 mm. long, soon caducous; leaves petiolate, the petioles 3-10
mm. long, exalate, the blades chartaceous to subcoriaceous, ovate
to narrowly ovate or lanceolate-ovate, 2.5-15 cm. long, 1-6.5 cm.
broad, commonly about 104 cm., subtruncate to nearly subcordate
or rounded or broadly acute at base, broadly or narrowly acute to
somewhat attenuate or slightly acuminate at apex, entire, glabrous
above, pubescent beneath at least on the midrib when young, usually
also on veinlets and surfaces, at last glabrate, the costa shallowly
suleate above, raised beneath, the lateral nerves 5-12 per side, com-
monly 6-8, slightly raised on both surfaces; inflorescences axillary

SMITH & STONE—PACIFIC ISLAND PLANTS, XVII 19

among the leaves, dichasial, commonly 5-15-flowered, all axes and
bracts pubescent or in fruit somewhat glabrate, the peduncle 1-3 mm.
long, bibracteate at summit with lanceolate bracts 1.5-2 mm. long,
the major axes 2-6 mm. long, bibracteate at nodes, the pedicels 3-6
mm. long, bracteolate or bibracteolate and slightly flared into calyx;
flowers borne in triflorous (or irregular) dichasia, slightly sexually
dimorphic and apparently gynodioecious, heterostyled; calyx cupuli-
form, glabrous, the lobes deltoid-ovate, acute or slightly acuminate,
3-5-nerved, marginally ciliolate, about 0.8 mm. long and broad;
corolla short-tubular, with spreading or reflexed lobes, glabrous
externally, 2.8-3.3 mm. long, the tube half as long, glabrous within
at base but above closely pilose around the throat with pale retrorse
trichomes 0.2-0.4 mm. long, the lobes glabrous within except at the
extreme base with a few trichomes along the veins, ovate, the nervation
relatively complex, cach lobe supplied by a branching system of nerves
consisting of a central main nerve entering the base of the tube and
producing 1 or commonly 2 pairs of lateral veins in the upper part of
the tube or throat or near the base of the lobe, each lateral vein
usually branched once or twice (or rarely thrice), producing brief
lateral veinlets, these occasionally obscurely forked near their apices,
all ascending in the lobe; stamens inserted at the throat, the filaments
deltoid-ligulate, 0.3-0.4 mm. long, retrorse-pilose with trichomes like
those of the corolla-throat, the anthers in short-styled flowers narrowly
deltoid, about 0.8 mm. long, devoid of pollen, minutely apiculate at
apex, the connective pilose at base, the thecae pilose with shorter
trichomes at base, the anthers in long-styled flowers fertile, oblong-
ovate, 0.8-0.9 mm. long, minutely retuse at apex, the connective
glabrous, the thecae basally sparsely hirtellous or glabrous; ovary
glabrous, in short-styled flowers large, subglobose, about 1.2 mm. in
diameter, the style obsolete orgto 0.3 mm. long; the ovary in long-
styled flowers smaller, subglobose, about 0.9 mm. in diameter, the
style 0.8-0.9 mm. long; stigma papillate, globose, about 0.8 mm. in
diameter; capsule ellipsoid, apiculate, 5-9.5 mm. long, 3.5-4.5 mm.
broad.

Type Loca.ity: Fiji, without precise locality; the type is Seemann
301, presumably the specimen deposited in the Berlin herbarium
and subsequently destroyed. However, isotypes are available at Kew
and the British Museum and are cited below.

DistriBpuTion: Fiji, Tonga, and Niue, of general occurrence from
near sea level to 600 m. altitude in dry forests, secondary forests,
open hilly country, or near streams. The species has been recorded
as shrubby but also as a tree up to 12 m. high, with a trunk 8 cm.
in diameter and curved, pendent branches. The greenish white

20 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

flowers produce a markedly foetid odor, which accounts for the
Fijian name.

LocaL NAMES AND USE: Commonly known in Fiji as mboimboinda,
but also with the following recorded names: mbatimara (Smith 8966),
mbatimboni (Smith 8405). In Tonga, recorded names are facfailunga
and jifilunga, and on Lifuka (Yuncker), te’e pilo a Maui. On Niue,
the names tete and teatea have been recorded by Yuncker. One record
of a medicinal use is given, from the Yasawa Group, Fiji, by St. John,
where some part of the plant is used as a remedy for stomach-ache.

FIJI: Yasawa Group: Waya: Olo Creek, north of Yalombi, in woods, S¢.
John 18018 (Bish, US). Virr Levu: Mba: North of Lomolomo, Degener &
Ordonez 13650 (A, Bish, K, NY, UC, US); north of Natalau, Degener 15003 (A,
Bish, K, NY, UC, US); mountains near Lautoka, Greenwood 1237 (A, UC, US);
Nandarivatu, Gibbs 555 in part, as to pubescent specimen only (BM). Nandronga
& Navosa: Near Kalaro, H. B. R. Parham 208A (BM); hillsides above Tholo
Levu, Nokonoko District, H. B. R. Parham 246 (BM). Serua: Hills west of
Waivunu Creek between Ngaloa and Korovou, Smith 9215 (US), 9274 (US);
hills between Navua River and Wainiyavu Creek, near Namuamua, Smith 8966
(US); hills east of Navua River near Nukusere, Smith 9100 (US). Ra: Vicinity
of Rewasa, near Waileka, Degener 15461 (A, Bish, K, NY, US). Namosi: Valley
of Wainambua Creek, south of Mt. Naitarandamu, Smith 8767 (US); hills north
of Wainavindrau Creek, between Korombasambasanga Range and Mt. Naita-
randamu, Smith 8405 (US); near Namosi, in valley of Waindina, Gillespie 2619
(Bish, UC). Naitasiri: Near Korosuli, on Wainimala River, Horne 978 (K);
near Viria, Meebold 16499 (JX); along Navutu-Nanduna track, B. E. Parham
3011 (A); Tholo-i-Suva, Parks 20890 (Bish, UC, US); woods near Nasinu, Gil-
lespie 3493 (Bish, UC, NY). Naitasiri or Rewa, on border: Mt. Kombalevu,
Parks 20293A (Bish, US), 20330 (Bish, UC, US). Rewa: Near Suva, Jeoward 100
(K). Viti Levu, without precise locality: Graeffe 1371 (K). Vanua Levu:
Mbua: Rukuruku Bay, H. B. R. Parham 5 (GH). Mathuata: Seanggangga
Plateau, in drainage of Korovuli River, near Natua, Smith 6642 (A, Bish, K,
NY, US); Lambasa, Greenwood 490 (KX); summit ridge of Mt. Numbuiloa, east
of Lambasa, Smith 6507 (A, Bish, K, NY, US); Navukuru, south of Lambasa,
Gressitt 2483 (US). Thakaundrove: Eastern drainage of Yanawai River, Dege-
ner & Ordonez 14104 (A, Bish, K, NY, UC, US); hills south of Nakula valley,
Smith 326 (Bish, GH, Kk, NY, UC, US); Maravu, near Salt Lake, Degener &
Ordonez 14233 (A, Bish, K, NY, UC, US). Vanua Levu, without precise locality:
U.S. Expl. Exped. (source of the name G. rupestre var. puberulum, GH, K, NY,
US); Horne 614 (K); Guppy in 1899 (KX); Tothill F456 (K). Taveunr: Near
Waiyevo, Gillespie 4634 (Bish); near Wairiki, Gillespie 4400.2 (Bish, NY, UC,
US). Ovatau: Milne 270 (K, type of G. crassifolium var. glaberrimum); Port
Kinnaird, Seemann 300 (BM, K). Naau: Shore of Herald Bay near Sawaieke,
Smith 8000 (US). Fiji, without definite locality: Seemann 301 (isotypes, BM,
Ix, fragment at GH).

TONGA: Vava’u: Home in 1846 (BM); Barclay 338A (BM); Crosby 229 (K);
Talau Hill, MacDaniels 1092 (Bish) ; near upper rim of seaside cliff above Ha’alaufuli
on northeastern'part’of island, Yuncker 16101 (US). ‘‘Vavau and Lifuka”: Harvey
in 1855 (GH, K). Liruxa: Yuncker 15787 (US). Tonga, without definite
locality, “Capt. Cook” (BM).

NIUE: East of Alofi village, Yuncker 9625 (A, Bish, US), 10092 (Bish, US);
near Mutalau village, Yuncker 9696 (A, Bish, UC, US).

SMITH & STONE—PACIFIC ISLAND PLANTS, XVII 21

Geniostoma vitiense was correctly segregated from the inclusive
concept of G. rupestre by Gilg and Benedict in 1921, but it appears to
have been cited since only in the two publications indicated by us
above. It is a well marked species, immediately distinguishable from
most of its congeners by its close, harsh indument, and more techni-
cally by the very short rather dense trichomes of the corolla-throat.
Some specimens exhibit a tendency toward glabrescence, while others
have harsh persistent indument even on mature parts. The intra-
petiolar stipules, the leaves of variable size, and the candelabriform
venation of the corolla-lobes are commonplace characters in the genus,
but the short trichomes of the corolla-throat and the vegetative
pubescence are good distinguishing characters. The leaf-blades, while
quite variable, are generally more or less ovate or ovate-lanceolate,
often rather broadly rounded or subtruncate at base, and usually
acute rather than acuminate at apex. The puberulent dichasial axes
are another distinguishing feature of G@. vitiense, hardly unique, but
among the other Fijian species present only in G. calcicola, perhaps in
G. insulare, and sometimes in G. stenocarpum.

The sexual dimorphism of the flowers of this species, as yet not
very strongly differentiated morphologically, is quite possibly com-
plete in respect to pollen abortion within the anther. As noted in the
description, slight divergences in pubescence and in the form of anther
exist between the “‘pistillate’ and “perfect” flowers, as well as in the
gynoecia. As far as can be made out from the specimens examined,
the “perfect”’ flowers are constant on any one individual, while other
individuals produce ‘‘pistillate’’ flowers; thus an incipient, if not an
actual, gynodioecism is present. However, sterilization of the stami-
nate organs has not, apparently, been accompanied by sterilization
of the pistillate organs in other individuals; and fruits are to be found
on both ‘‘pistillate” and ‘‘perfect” individuals. There may, of course,
be some difference in the degree of fertility between these sexual
forms, but it is not apparent. The same phenomenon was reported
by Valeton for several species of Malaysia. In our area, it is probably
common in other species, and definite indications of it have been seen
in G. macgregorii, while in some species the flowers appear to be all
strictly hermaphrodite (as in G. macrophyllum). A more intensive
study of this problem, carried out in the field, would probably yield
some useful information. Perhaps associated with this phenomenon
is that of odor; the malodorous flowers of G. vitiense and of certain
other species are possibly the hermaphrodite rather than the perfect
flowers, or vice versa. The odor is such that it probably attracts
flies as pollinating vectors. Scentless, or at least not foetid, flowers
are known in such species as G. clavigerum and perhaps G. macgregorit.
Such a difference, if it is associated with a difference in the mode of
pollination, could be taxonomically significant.

22 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

7. Geniostoma gracile Rechinger in Rep. Sp. Nov. 6: 325. 1909, in Denkschr.
Akad. Wiss. Wien 85: 328. ¢. 11. 1910; Gilg & Benedict in Bot. Jahrb. 56:
544. 1921: Christophersen in Bishop Mus. Bull. 128: 75, pro parte, as to
Mac Daniels 1121 only. 1935.

PLATE 1, FIGURE 7; PLATE 3, FIGURES 9-11
Shrub or small tree with drooping branches, glabrous throughout
except for some floral parts, with slender subterete branchlets distally

subcompressed and 1-2 mm. thick; stipules intrapetiolar, forming a

gently rounded sheath about 1 mm. high; leaves petiolate, the petioles

slender, 4-7 mm. long, the blades thin-chartaceous, lanceolate-ovate
to elliptic-ovate, relatively narrow, (1.5-) 4-13 cm. long, 1-4.5 cm.
broad, rounded to subcuneate and briefly decurrent at base, long-
attenuate and sharply acuminate at apex, entire, the costa sulcate
above, raised beneath, the lateral nerves 4~10 per side, slightly raised
on both surfaces, the veinlets smooth, mostly plane and subevident or

obscure; inflorescences axillary among the leaves, dichasial, 3-7

(usually 5-)-flowered, the axes slender and notably elongate, the

peduncle 1-7 mm. long, the axes and pedicels 7-17 mm. long, with

short bracts about 1 mm. long, the pedicels slightly flared into the
calyx; flowers externally glabrous; calyx cupuliform, the lobes deltoid-
ovate, acute, 0.7-1 mm. long, 1-nerved to obscurely 3-nerved; corolla
about 3.5 mm. long, tubular, divided to middle into suberect or patent
deltoid-ovate lobes, the tube closely pubescent within, at throat with
pale reflexed trichomes about 0.3 mm. long, those nearer the base of
the tube shorter, about 0.1 mm. long, the tube pilose within on the
proximal third or half, glabrous distally, the venation relatively com-
plex, that of each corolla-lobe derived from a single central vein
ascending from below the tube up to the throat, there producing one or
usually two pairs of ascending laterals, these in turn branched laterally
once, twice, or even thrice, in the lobe, all the branches ascending and
not anastomosing; stamens inserted at throat on short retrorsely pilose
deltoid-ligulate filaments about 0.2-0.3 mm. long, the anthers narrowly
deltoid-oblong, bluntly tipped, about 0.80.35 mm., glabrous; ovary
subglobose, about 1 mm. in diameter, narrowed into the filiform
glabrous style, this about 1-1.4 mm. long, the stigma clavate-obovoid
and subbilobate, papillate, about 0.8-0.9 mm. broad and 1 mm. long;
capsule ellipsoid, apiculate, 5-9 mm. long, 4-7 mm. broad.

Type Locauity: Upolu, Samoa; cited syntypes are Rechinger 948
and 1475, of the former of which duplicates are cited below.

Disrripution: Endemic to Samoa; known only from Savaii and

Upolu. The plant is shrubby or becoming a small tree, occurring from

near sea level to about 500 m. altitude, in forest or shrubby vegetation.

The flowers are white.

Locat NAME: Taipovpo, according to Gilg & Benedict, who probably
derived their information from labels on one of Rechinger’s specimens.

SMITH & STONE—PACIFIC ISLAND PLANTS, XVII 23

SAMOA: Savami: Coastal thickets, Reinecke 532 (K). Upotu: Above U’umapu
[Utumapu] (‘“Utmapu” of Rechinger and Gilg & Benedict), Rechinger 948
(isosyntype, BM, US); Vaea, MacDaniels 1121 (Bish).

In contrast to Christophersen, who cited a large suite of specimens
under this species, we find that only the few specimens cited above
represent the species as typified by an isosyntype. Most of the speci-
mens cited by Christophersen under G. gracile are here treated as G.
samoense var. parviflorum, a variety imperfectly known to Gilg &
Benedict and apparently to Christophersen, and sustained by us as a
distinct entity, although the type has not been seen and was pre-
sumably destroyed along with the type material of G. gracile, G.
vitiense, and many other species at Berlin. Geniostoma gracile ap-
pears to be related to G. macgregorit and G. vitiense, species which also
exhibit the very short corolla-throat trichomes characteristic of G.
gracile. In the narrow, sharply acuminate leaf-blades and the un-
usually elongate cymose dichasia, G. gracile differs from its Fijian
relatives and also from most other Samoan Geniostomata. However,
specimens of G. samoense var. parviflorum sometimes bear dichasia
almost as elongated as those of @. gracile; the former appears to be a
population commonly found at rather high elevations, and of course it
differs from G. gracile in its long corolla-throat trichomes.

8. Geniostoma macgregorii (Horne ex Baker) Smith & Stone, comb. nov.
PLATE 1, FIGURE 8; PLATE 2, FIGURES 17-19
Plectronia macgregorit Horne, A Year in Fiji, 266, nomen. 1881; Horne ex Baker
in Journ. Linn. Soc. Bot. 20: 363. 1883.
Geniostoma rupestre sensu auct. quoad plantas vitienses, pro parte.

Shrub or small tree to 10 m. high, glabrous throughout except for
some floral parts; branchlets slender, subterete or slightly compressed
and 1-3 mm. in diameter distally; stipules intrapetiolar, forming a
subtruncate or gently rounded (or in youngest state, somewhat
deltoid) sheath 1-1.5 mm. high; leaves glabrous, petiolate, the petioles
4-16 mm. long, slender, proximally exalate, the blades thick-mem-
branous, chartaceous, or subcoriaceous, when juvenile subglossy, later
dull, ovate to elliptic-ovate or oblong-ovate or sublanceolate, 3-16 cm.
long, 1-6 cm. broad, commonly about 103.5 cm., attenuate, acute,
or somewhat rounded at base, briefly decurrent on the petiole, atten-
uate and acuminate at apex, often very narrowly so or even caudate,
entire, the costa sulcate above, slightly elevated beneath, the lateral
nerves 5-10 per side, slightly raised on both surfaces, the veinlets
evident or obscure, plane; inflorescences axillary among the leaves,
dichasial, usually 5-15-flowered, rarely to 20-flowered, all the axes
glabrous, the peduncle stout, flattened, 1-4 mm. long, bibracteate at
summit with lanceolate bracts 1-1.5 mm. long, the major axes slender,
2-6 mm. long, bibracteate at nodes, the pedicels 2-6 mm. long, slender,

24 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

flared and bracteolate below the calyx; flowers borne in triflorous (or
near the peduncle uni- or biflorous) dichasia, glabrous externally;
calyx cupuliform, the lobes deltoid-ovate, 1-3 (rarely 5-)-nerved,
about 1 mm. long, as broad or nearly as broad; corolla short-tubular,
slightly urceolate, with reflexed lobes, 3-4 mm. long, divided to
middle, the tube glabrous within basally, above to throat and on base
of lobes closely pilosulous with short retrorse trichomes 0.1-0.25 mm.
long, the lobes ovate, basally pilosulous like the throat, distally merely
pustulous with pustules 0.05 mm. high, at the extreme apex glabrous,
the venation relatively complex, each corolla-lobe supplied by three
main nerves entering the tube already distinct, or diverging in the
lower part of tube, ascending to throat and into the lobe, producing
2-4 short ascending (rarely forked) lateral nerves; stamens inserted
at throat, the filaments deltoid-ligulate, about 0.4 mm. long, closely
pilosulous with short retrorse trichomes like those of the throat, the
anthers narrowly oblong-deltoid, about 0.8 mm. long, acute, glabrous;
ovary glabrous, subglobose, about 1.4 mm. high and 1.8 mm. broad,
the style glabrous, filiform, 1.2-1.5 mm. long in long-styled flowers,
0.7-0.9 mm. long in short-styled flowers, the stigma globose; capsule
narrowly ellipsoid or narrowly ovoid, 7-13 mm. long, 3-4.5 mm. broad.

Type Locatity: Ovalau, Fiji; the type is Horne 261, cited below.

DistripuTion: Known only from Fiji and Tonga. The plant is a
slender shrub or small tree 3-10 m. high, bearing flowers with a
greenish white or pale green corolla and a white stigma; the fruit is
green without, the inner pulp orange. The habitat is noted as dense
or open forest, or near the margins of forest, from near sea level to
900 m. altitude, mostly between 50 and 200 m. altitude.

LocaL NAMES: Very rarely noted, although numerous specimens
are at hand; mboimboinda on a few specimens from Vanua Levu;
Graeffe’s collection from Ovalau bears the name mbatimbona. Three
collectors note that the foetid odor common to G. vitiense is not
present, and indeed one states “‘odorless” (Greenwood 541A); another
states that there is an odor, but not a foetid one (rather, says the
Tothill label, a “horsy” scent). These data, in conjunction with the
rare usage of the name mboimboinda, may indicate that the plants are
scentless or at least not malodorous.

FIJI: Vitr Levu: Mba: Mt. Evans, Greenwood 388 (K); near Lautoka,
Greenwood 49 (K); northern slopes of Mt. Namendre, east of Mt. Koromba
(‘Pickering Peak’’), Smith 4554 (A, Bish, K, NY, US). Nandronga & Navosa:
Near Mbelo and Vatukarasa, Degener 15296 (A, K, NY). Serua: Hills west
of Waivunu Creek, between Ngaloa and Korovou, Smith 9318 (US), 9469 (US);
vicinity of Ngaloa, Degener 15071 (A, Bish, K, NY, UC, US); near Vatutavathe,
Degener 15206 (A, Bish, K, NY, UC, US); hills between Waininggere and Waisese
Creeks, between Ngaloa and Wainiyambia, Smith 9357 (US). Namosi: Coastal
hills, Greenwood 541A (A). Tailevu: Uthunivanua, McKee 2817 (US). Naitasiri:

SMITH & STONE—PACIFIC ISLAND PLANTS, XVII 25

Near Viria, on Rewa River, Meebold 16507 (Bish); woods beyond Tamavua,
Gillespie 2151 (Bish), 2170 (A, Bish, NY, UC, US); Wainimbuku River area, on
ridge near Nasinu river, Shradha Nand [Fiji Dept. Agric. No.] 7358 (US); near
Nasinu, Gillespie 3528 (Bish, NY, UC); Suva Pumping Station, Degener &
Ordonez 13757 (A, US). Rewa: Near Naikorokoro River west of Suva, Meebold
21943 (Bish); Mt. Korombamba, H. B. R. Parham 84 (BM); southeastern slopes
of Mt. Korombamba, Gillespie 2230 (Bish, UC); summit of Mt. Korombamba,
Gillespie 2395 (Bish, UC); lower slopes of Mt. Korombamba, Vaughan 3327
(BM, K); near Suva, Tothill 477 (Bish, K), 623 (K), Meebold 16436 (K), 21387
(BM, NY); Central Road, Suva, Tothill 625 (Bish); mouth of Rewa River, Guppy
in 1899 (K). Kanpavu: Hills above Namalata and Ngaloa Bays, Smith 132
(Bish, GH, K, NY, UC, US). Vanvua Levu: Mathuata: Hills near Lambasa,
Greenwood 511 (K), 541 (K); southern slope of Mt. Numbuiloa east of Lambasa,
Smith 6398 (A, Bish, K, NY, US), 6576 (A, Bish, K, US). Thakaundrove:
Southwestern slope of Mt. Mbatini, Smith 668 (Bish, GH, K, NY, UC, US).
OvaLau: Graeffe 1612 (K); vicinity of Levuka, Horne 96A (GH, K); ‘‘mountains
of Ovalau,’” Horne 261 (GH, K type). Koro: Tothill 612 (Bish in part, K).
Matuxku: Milne 128 (K), 128 bis (K). Kamara: Central wooded basin, Bryan
504 (Bish). Fiji Islands, without definite locality, U. S. Expl. Exped. (US).
TONGA: Vava’u: Near Neiafu, Setchell 15656 (UC).

This species, frequent in Fiji, was described in very general terms
by Baker (and not at all by Horne); it provides not the only illustra-
tion of confusion between Geniostoma and Plectronia (i.e. Canthium,
Rubiaceae), as evidenced by G. niinoanense Kanehira (=Canthium
barbatum (Forst. f.) Seem. var. korrorense (Valeton) Fosberg, the
variety based on Plectronia korrorensis Valeton). However, there is
no question as to the identity of Horne’s specimen, and therefore his
specific epithet, corrected in accordance with Rec. 73C of the Interna-
tional Code of Botanical Nomenclature, must be used.

Although material here cited has generally been previously identified
as G. rupestre, it is less closely related to that species than it is to
G. vitiense, even though the pubescence of the latter gives a superficial
impression of difference that is not entirely warranted. The short
trichomes of the corolla-throat are a constant feature of G. macgregoru
and serve to distinguish it at once from the New Hebridean G. rupestre.
From G. vitiense, G. macgregorii differs in being glabrous and in having
its leaf-blades relatively more narrow (in most specimens) and usually
long-acuminate; in floral characters the two are very similar. The
remarks on sexual dimorphism given under G. vitiense also apply here.

Only the single specimen cited above from Tonga carries the known
distribution of G@. macgregorii beyond Fiji; this specimen, while not
entirely typical, cannot be referred to any other species occurring in
Tonga and we believe it to be correctly placed here. Outside of our
area, G. batanense Merr., of the Philippines, is somewhat similar in
general aspect to G. macgregorii, but it has externally puberulent
flowers and a setulose style.

26 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

9. Geniostoma calcicola A. C. Sm. in Sargentia 1: 99. 1942.
PLATE 1, FIGURE 9; PLATE 2, FIGURE 24

Geniostoma sp. A. C. Sm. in Bishop Mus. Bull. 141: 125. 1936.

Scandent shrub to 50 cm. high, with slender terete fusco-cinereous
branchlets at first densely short-setose with pale, conic-tapered,
several-celled setae; stipules intrapetiolar, forming a short broadly
ovate sheath; leaves densely puberulent, petiolate, the petioles 3-5
mm. long, the blades chartaceous, elliptic or oblong-elliptic, 3.5-5 cm.
long, 1.5-2.5 cm. broad, obtuse at base, rounded or slightly emarginate
or minutely mucronulate at apex, entire, with slightly recurved mar-
gins, pilose on both surfaces, the trichomes about 0.3 mm. long, the
costa shallowly sulcate above, raised beneath, the lateral nerves 4-6
per side, shallowly sulcate above, slightly raised beneath, the veinlets
obscure; inflorescences axillary, dichasial, slender, open, 5 (rarely 9-)-
flowered, the peduncle 1 mm. long, the pedicels 2-5 mm. long, the
nodes bibracteate, the flowers bracteolate just below the calyx, all the
axes sparingly setulose; flowers borne on relatively long slender
pedicels scarcely flared at apex; calyx cupuliform, glabrous except for
the ciliolate margin, the lobes acute, ovate-deltoid, 1.2-1.5 mm. long,
1-3-nerved; corolla short-tubular, 3.5—-4 mm. long, glabrous externally,
the tube about 2.5 mm. in diameter, glabrous within at base, pilose
at throat with white trichomes about 0.8 mm. long sparsely but uni-
formly disposed around throat, the lobes about 1.5 mm. long, ovate,
reflexed, glabrous except at extreme base, the venation pinnate, each
corolla-lobe supplied by one main nerve branched just below the lobe
or sometimes toward base of tube into 2 main lateral nerves, all
ascending into the lobe and there unbranched or with a few short free
laterals; stamens inserted at throat, the filaments ligulate, about 0.4
mm. long, sparsely pilose, the anthers oblong, basally cordate, apically
obtuse, about 1 mm. long; ovary glabrous, depressed-subglobose,
about 1.3 mm. in diameter, the style filiform, about 1 mm. long,
glabrous, the stigma subglobose to somewhat obcordate, about 0.8 mm.
in diameter, papillate; capsule unknown.

TYPE LocaLity: Fulanga, Fiji; the type is Smith 1116, cited below.

DistriBuTion: Known only from the type collection, collected on
cliff face in limestone formation; the corolla is white. The other
specimens tentatively referred to G. calcicola in the original publication
are now accounted for under G. insulare.

FIJI: Futanea: Near Monothaki, near sea level, Smith 1116 (Bish, GH type,
K, NY, UG, US).

Although the occurrence of this species on other islands of Lau may
be suspected, at present we do not have a single collection other than

SMITH & STONE—PACIFIC ISLAND PLANTS, XVII 27

the type referable to it. Its closest relatives are G. crassifolium Benth.,
of the Isle of Pines, and G. insulare, described below. Relationships
among the three species may be indicated in the following key:
Leaf-blades puberulent on both surfaces.

Ovary and style hirtellous, the style 0.3-0.5 mm. long, the stigma clavate;
anthers about 1.2 mm. long; dichasia closely hirtellous; leaf-blades up to
7X4 em... we ee ee G. crassifolium

Ovary and style glabrous, the style 0.8-1 mm. long, the stigma obovoid or sub-
globose; anthers about 0.9 mm. long; dichasia sparsely hirtellous; leaf-
bladesupto5X2.5em.......... 020 0888 G. calcicola

Leaf-blades glabrous or puberulent only beneath. . ....... G. insulare

Reference of Tongan plants to G. calcicola in Yuncker’s Plants of
Tonga (in Bishop Mus. Bull. 220: 215. 1959) was based, we now believe,
on too inclusive a concept, and these plants are now cited under G.
insulare.

10. Geniostoma dictyoneurum Smith & Stone, sp. nov.
PuatsE 1, FiauRE 10

Arbor gracilis ad 7 m. alta praeter corollae partem interiorem
omnino glabra, ramulis gracilibus apicem versus 1-2 mm. diametro
et subteretibus vel obscure quadrangularibus; stipulis intrapetiolaribus
in vagina rotundata vel subtruncata 1-1.3 mm. alta conjunctis;
foliis magnitudine moderatis, petiolis 8-12 mm. longis, laminis mem-
branaceis vel tenuiter chartaceis ovatis, 7-12 cm. longis, 3.5-7 cm.
latis, basi rotundatis vel subtruncatis et in petiolum subito cuneato-
decurrentibus, apice rotundatis, margine integris, costa supra sulcata
subtus conspicue elevata, nervis lateralibus utrinsecus 4-10 utrinque
prominulis, venulis obscuris; inflorescentiis axillaribus vel ramulis
paulo infra folia enatis dichasialibus, pedunculo crasso 1-2 mm. longo
apice bibracteato axes 1-3 principales graciles nodis bibracteatos
4-6 mm. longos gerente, bracteis 0.5-1 mm. longis, axibus 3-9-floris,
floribus 1-3 in dichasiis dispositis, pedicellis gracilibus 1-3 mm.
longis infra calycem dilatatis et bracteola calycem adpressa ornatis;
calyce cupuliformi, lobis deltoideis 0.6—0.8 mm. longis minute ciliolatis
uninervatis vel interdum inconspicue 3-nervatis; corolla circiter 4 mm.
longa ad medium lobata, tubo faucibus pilis circiter 0.5 mm. longis
hirtello, lobis ovatis basim versus interdum parce hirtellis, nervatio
intricato, corollae quoque loba nervis 3 e basi tubi in lobam adscen-
dentibus praedita, quoque nervo venulis lateralibus pluribus tenuibus
in reticulum laxum conjunctis ornato; staminium filamentis ad 0.2 mm.
longis copiose hirtellis, antheris anguste deltoideo-ovoideis circiter
0.90.4 mm. apice acutis basi haud parce hirtellis; ovario depresso-
globoso circiter 0.8 mm. lato, stylo filiformi ad 0.5 mm. longo, stigmate

28 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

subgloboso papilloso circiter 0.5 mm. lato; capsula subglobosa vel
ate ovoidea, 5-6 mm. longa, 4.5-5.5 mm. lata, apice apiculata.

Type in the U.S. National Herbarium, No. 2190641, collected among the hills
east of Herald Bay, inland from Sawaieke, Ngau, Fiji, alt. 300-450 m., June 18,
1953, by A. C. Smith (No. 7814). Duplicates to be distributed.

ADDITIONAL SPECIMENS EXAMINED:

FIJI: Koro: Rocky thicket at sea level on east coast, Smith 1037 (Bish, GH,
K, NY, UC, US).

DistriruTion: Endemic to Fiji, and known only from Ngau and
Koro. The plant is recorded as a slender tree 5-7 m. high, growing
in forest from sea level to 450 m. altitude, the flowers with corolla
and filaments pale green or white, the anthers white.

Locat NAME: Mboimboinda (Koro).

Geniostoma dictyoneurum, as the name implies, exhibits anasto-
mosing venation of the corolla-lobes, a character found very rarely
in other species. The leaf-blades are rather broadly ovate, apically
blunt or rounded, noticeably thin-membranous, and relatively large
for the genus, with 4-10 lateral nerves on each side of the midrib.
The pilosity of the corolla inner surface is densest on the filaments;
on the corolla-throat it is sparser and covers a narrower band. In
this respect there exists a certain resemblance to specimens of G.
vitiense, Which occasionally shows a similar variability in corolla-
indument. The completely glabrous vegetative parts of the present
species readily distinguish it from G. vitiense. A resemblance to
G. macgregorit may also be noted, but the long corolla-throat hairs
of G. dictyoneurum, and of course the anastomosing corolla-lobe
venation, quickly differentiate it. Perhaps the closest relative of
G. dictyoneurum is G. insulare, at least the form with subglobose
capsules; in G. dictyoneurum the capsules are also regularly sphaeroidal
rather than elongated.

11. Geniostoma fleischmannii Rechinger in Rep. Sp. Nov. 7: 17, as G. fleisch-
mannt. 1909, in Denkschr. Akad. Wiss. Wien 85: 329. ¢. 10. 1910; Gilg &
Benedict in Bot. Jahrb. 56: 545. 1921; Christophersen in Bishop Mus. Bull.
128: 175, as G. fleischmanni. 1935. PLATE 1, FIGURE 11

Shrub or small tree 3-4 m. high, glabrous throughout except for
some floral parts; branchlets slender, gray-brown, markedly quad-
rangular and narrowly winged, distally 1-3 mm. thick; stipules intra-
petiolar, forming a truncate sheath about 1 mm. long; leaves glabrous,
petiolate, the petioles 2-7 mm. long, the blades oblanceolate or nar-
rowly elliptic, 4-12 em. long, 1.5-3.5 cm. broad, attenuate-decurrent
or sometimes acute and decurrent at base, acute to acuminate at apex,
the costa shallowly sulcate above, slightly raised beneath, the lateral
nerves 6-12 but usually 9 or 10 per side, slightly raised on both sur-
faces, the veinlets obscure; inflorescences axillary among the leaves,

SMITH & STONE—PACIFIC ISLAND PLANTS, XVII 29

dichasial, usually 5-10-flowered, the peduncle flattened, subobsolete
or to 1.5 mm. long, the axes and pedicels 4-9 mm. long, bracteate with
bracts to 1.5 mm. long, the pedicels bracteolate and flared just below
calyx; flowers slenderly pedicellate; calyx cupuliform, the lobes del-
toid, about 1 mm. long and 0.8 mm. broad, 1-3-nerved; corolla
4.5-5 mm. long, the tube 2.5-2.7 mm. long, glabrous without, glabrous
within at the extreme base but retrorse-pilose above to throat with
trichomes 0.7—0.9 mm. long, the lobes 1.9-2.3 mm. long, glabrous dis-
tally, deltoid-ovate, the venation complex, each lobe supplied by one
main nerve, this usually narrowly divaricate in the tube, ascending
and branching into the lobe, producing 2-4 ascending lateral branch-
nerves, these occasionally briefly branched; stamens inserted at
throat, with short pilose filaments 0.3-0.5 mm. long, the anthers del-
toid, blunt-tipped, about 0.7 mm. long; ovary glabrous, about 1.5
mm. high and 1.8 mm. broad, the style filiform, glabrous, about 1
mm. long, the stigma subglobose, subbilobate, about 0.9 mm. in
diameter; capsule ellipsoid, 7-8 mm. long, about 4 mm. broad, short-
apiculate.

TypE Locauity: Savai, Samoa; the type is Rechinger 1648, pre-
sumably deposited at Berlin and subsequently destroyed. We have
seen no isotypes, but the species is very well illustrated and easily
recognized.

DistrirnutTion: Endemic to Samoa; apparently known only from
Savai. It is said to be a shrubby species becoming a small tree 4 m.
high, with markedly 4-angled branchlets and with white flowers,
occurring at high elevations (1030 m. according to Christophersen &
Hume), or in “sunny openings near Asau” (Rechinger).

SAMOA: Sava: Forest above Matavanu, Christophersen & Hume 2124 (Bish,
NY, UC, US).

Characters that amply distinguish G. fleischmannii from other
Samoan species, and in combination from its congeners throughout the
Pacific, are its relatively large corollas, its candelabriform corolla-lobe
venation of an unusual type (consisting of a dichotomous main nerve
with several laterals rather than being 3 or 5-nerved), its long, narrow,
usually oblanceolate leaf-blades, and its 4-winged branchlets.

12. Geniostoma stenocarpum A, C. Sm. in Sargentia 1: 98. 1942.
PLATE 1, FIGURE 12; PLATE 3, FIGURES 7, 8
Shrub or small tree to 4 m. high, with at first often minutely
pubescent but soon glabrate slender subterete branchlets; stipules
intrapetiolar, forming a short rounded sheath; leaves petiolate, the
petioles 2-6 mm. long, the blades lanceolate or narrowly ovate,
4-7.5 cm. long, 1.2-2.7 cm. broad, attenuate and decurrent or merely
acute at base, or very rarely subtruncate, narrowly acute to slightly

30 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

acuminate at apex, entire, the costa slightly sulcate above, raised
beneath, the lateral nerves 4-7 per side, prominulent, the veinlets
mostly obscure; inflorescences glabrous or sometimes at first minutely
pubescent and soon glabrate, axillary among the leaves, 5-12(-18)-
flowered, dichasial, compound dichasial, or often irregular, the peduncle
1 mm. long or less, the major axes 3-6 mm. long, the nodes bibracteate
(sometimes with only 1 or with no bract), the pedicels 2-5 mm. long,
bracteolate, the bracteoles about 0.5 mm. long; calyx cupuliform, the
lobes deltoid-ovate, acute or short-acuminate, 1-3-nerved, the margins
ciliolate; corolla short-tubular, 2.5-3.5 mm. long, glabrous externally,
divided to middle, the tube glabrous within basally, sparsely hirtellous
at throat with trichomes about 0.4 mm. long, similarly hirtellous in a
narrow median horizontal band and on filaments (but glabrous between
filaments in upper portion), the lobes ovate, glabrous, the venation
relatively complex, each corolla-lobe supplied by 3 nerves with several
lateral branches in the lobe, derived from a single main nerve entering
the tube and producing two main lateral nerves in the throat; stamens
inserted at throat, the filaments narrowly deltoid-ligulate, about
0.6 mm. long, pilose, the anthers glabrous except at extreme base,
obtuse or with a minute mucronulate apex, ovate-oblong, 0.8—0.9 mm.
long; ovary glabrous, subglobose, about 1 mm. in diameter, the style
filiform, glabrous, short to nearly obsolete, 0.3-0.7 mm. long, the
stigma subglobose, about 0.5 mm. in diameter, papillate; capsule
ellipsoid, 7-10 mm. long, 2.5-3.5 mm. wide, apiculate by the stylar
remnant, subtended basally by the persistent calyx.

Type tocauiry: Nandarivatu, Viti Levu, Fiji; the type, Degener &
Ordonez 13591, is cited below.

Disrripution: Endemic to Fiji and thus far known only from Viti
Levu, where it appears quite frequent at middle elevations. It has
been recorded as a shrub or small tree up to 4 m. high, occurring in or
on borders of dense forest at elevations up to 1150 m. and only rarely
as low as 150 m.; the flowers are greenish or white.

Loca names: Mbati-mbati (Gillespie 3170); mbuimbuita (Smith
4993, 6020); mbatimara (Smith 5062).

FIJI: Virt Levu: Mba: Nandarivatu and vicinity, Gibbs 555, pro parte (BM);
Tothill 626 (KX), Gillespie 3170 (Bish), 4280 (Bish, UC), Degener & Ordonez 13591
(A type, Bish, K, NY, UC, US), 13592 (A, Bish, K, NY, UC, US), 14314 (A, Bish,
K, NY, UC, US); slopes of escarpment north of Nandarivatu, Smith 6020 (A,
Bish, K, NY, US); Mt. Ndelaiyod, on escarpment west of Nandarivatu, Smith
5062 (A, Bish, K, NY, US); ridge between Mt. Nanggaranambuluta and Mt.
Namama, east of Nandarivatu, Smith 4993 (A, Bish, K, NY, US); slopes of Mt.
Tomanivi [Mt. Victoria], Smith 5226 (A, Bish, K, NY, US); hills between Nandala
and Nukunuku Creeks, along trail from Nandarivatu toward Lewa, Smith 6152
(A, Bish, K, NY, US). Nandronga & Navosa: Northern portion of Rairaimatuku

Plateau, between Nandrau and Rewasau, Smith 5394 (A, Bish, K, NY, US).
Serua: Banks of Navua River, at Namata Rapids, Gillespie 2941 (Bish, UC).

SMITH & STONE—PACIFIC ISLAND PLANTS, XVII 31

The available specimens fall into three fairly well marked forms
that appear not significant enough to merit names: a narrow-leaved
glabrous form, a narrow-leaved puberulent form, and a glabrous form
with comparatively broad, ovate leaf-blades. The first of these
includes the type collection and most of the cited specimens. Only
three collections (Degener 14314 and Gillespie 2941 and 4280) show
a marked and fairly persistent puberulence on the branchlets; a
minute indument is frequent on the innovations but is usually soon
lost. One puzzling specimen, Smith 6020, seems to have both gla-
brous and persistently puberulent branchlets on one tree. The third
form, with leaf-blades ovate and rounded to subtruncate at base, is
represented by Gibbs 555 and Smith 5394. Association of these three
forms with particular habitats or altitudes does not at this time seem
to be noteworthy.

The closest relationship of G. stenocarpum appears to be with
G. rupestre, of the New Hebrides, and less notably with G. insulare.
The small and predominantly lanceolate leaf-blades give a very
distinct aspect to G. stenocarpum. Although corolla-lobe venation
is not an entirely dependable criterion of relationship, in this respect
G. stenocarpum is closer to G. insulare than to G. rupestre, in which
the lobe-supplying nerve usually divides at or near the base of the
corolla-tube.

The flowers of G. stenocarpum are noted as foetid only by Gibbs,
whose No. 555 is a mixture with G. vitiense; her comment may apply
to the latter species, which is reported by various collectors as having
malodorous flowers.

13. Geniostoma samoense Reinecke in Bot. Jahrb. 25: 666. 1898; Rechinger in
Denkschr. Akad. Wiss. Wien 85: 328. 1910; Gilg & Benedict in Bot.
Jahrb. 56: 543. 1921; Setchell in Carnegie Inst. Publ. 341: 59, 1924.

PuaTE 1, FIGURE 13

Small tree to 10 m. high or shrub, glabrous throughout except the
axes of the inflorescence and some floral parts, or the innovations at
first minutely pubescent, soon glabrate; branchlets distally 1-3 mm.
in diameter, subterete or somewhat compressed-quadrangular, in age
terete; stipules intrapetiolar, forming a subtruncate or gently rounded
sheath 1-2 mm. high; leaves petiolate, the petioles 5-15 mm. long,
exalate, the blades chartaceous, ovate to broadly ovate, 3-18 cm.
long, 1.5-9 cm. broad, rounded or obtuse-cuneate and decurrent at
base, obtuse to acuminate at apex, entire, the costa sulcate above,
raised beneath, the lateral nerves 5-10 per side, slightly raised on
both sides, the veinlets manifest or somewhat obscure, not raised;
inflorescences axillary among the leaves, dischasial, the dichasia
strict or often irregularly branching, 5-15-flowered, commonly 9-
flowered, the peduncle about 1 mm. long and 0.7 mm. thick, bibrac-

597008—61——3

32 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

teate at summit, the major axes 2-5 mm. long, the pedicels 1-3 mm.
long, bibracteolate at nodes or sometimes irregularly 1- or 2-
bracteolate, the bracteoles when opposite continuous across the node,
sheathing, all the axes minutely puberulent with short blunt trichomes
no more than 0.03 mm. in length; calyx 1.3-1.6 mm. long, the lobes
equal in length to the tube, broadly deltoid, about 0.7 mm. long and
1mm. wide, marginally ciliolate, 3-nerved (rarely obscurely so) ; corolla
2.5-3.3 mm. long, the tube 1.2-1.8 mm. long, glabrous without,
glabrous within in the lower half, pilose in the upper half to the throat
with white trichomes 0.45-0.7 mm. long, the lobes glabrous or pilose
only at extreme base, the venation comparatively complex, each
corolla-lobe supplied by 3 main nerves diverging near base of tube
and branching laterally near throat, all ascending into the lobe, there
with brief lateral branches or none; stamens inserted at throat, the
filaments pilose, ligulate, about 0.3 mm. long, the anthers ovate,
acute or obtuse, about 0.7 mm. long and 0.5 mm. broad; ovary gla-
brous, subglobose, about 1 mm. in diameter, the style about 0.2 mm.
long, glabrous, the stigma depressed-globose or subglobose, papillate,
0.7-1 mm. in diameter; mature capsule ellipsoid-obovoid, briefly
apiculate by the stylar remnant, the lobes dehiscent-recurved, 8-10
mm. long, 4-5 mm. broad; seeds ovoid with a concave hilum, the
testa minutely papillate, about 0.06 mm. thick, the papillae no more
than 0.015 mm. high, orange in life.

Geniostoma samoense is a species of the alliance of G. rupestre,
differing chiefly in the puberulent dichasia, the relatively broader
capsules, and the larger, broader, usually more obtuse leaf-blades.
The venation of the corolla-lobes is very similar to that of G. rupestre,
there being some variability within each species as to the height in
the corolla-tube of the point of divergence of the three main nerves.
The venation of the sepals is generally 3-nerved, although the two
lateral nerves are sometimes short and obscure. The dichasia are
distinctly but minutely puberulent with trichomes no more than
0.03 mm. long, which are usually persistent even on fruiting specimens.

Several specimens cited by Christophersen (in Bishop Mus. Bull.
128:176. 1935) as G@. rupestre are here referred to G. samoense, a
species not considered by Christophersen in identifying his material.
There is no indication that Christophersen believed Reinecke’s
species a synonym of G. rupestre, but this was possibly the case.
We are satisfied that the two entities amply merit specific recognition.

The available material of G. samoense represents two varieties,
which may be differentiated as follows:

Leaf-blades usually 8-18 X 4-9 cm., rounded at base, obtuse to obtusely acuminate

at apex; anthers blunt at apex; stigma globose; plants of low elevations.
var. samoense

SMITH & STONE—PACIFIC ISLAND PLANTS, XVII 33

Leaf-blades usually 3-11 X1.5—-4.5 cm., cuneate at base, long-acuminate at apex;
anthers minutely mucronulate; stigma depressed; plants usually of
mountain forests... . 2... ee ee ee eee ee «6Vr, parviflorum

13a. Geniostoma samoense var. samoense
PLATE 2, FIGURES 25, 27, 28; PLATE 3, FIGURES 1-4
Geniostoma rupestre var. macrophyllum A. Gray in Proc. Amer. Acad. 4: 33,
nomen. 1859.

Leaf-blades large, relatively broad, rounded at base, obtuse to
obtusely acuminate at apex, comparatively thin in texture; bracts
1-1.3 mm. long; anthers blunt at apex; stigma globose, the style
short.

TypE LocaLity: Samoa; the precise locality and the identity of
the holotype are uncertain, since Reinecke cited his own collection
(275) from Upolu and three Graeffe specimens (245 and 1393 from
Upolu, and 1613 from Savaii). Rather than designate a lectotype
at this time we prefer to consider the four collections as syntypes;
duplicates of two of them (Graeffe 1393 and 1613) have been seen
and are cited below as isosyntypes.

DistriBuTION: On the basis of available material, this typical
variety is seen to occur on several islands of Samoa and also on
Uvea, Wallis Islands. It is recorded as a shrub or small tree, with
white flowers; the seeds are brown, embedded in orange pulp. Habi-
tats reported are swampy forest near the coast, thickets on rocky
bluffs, near lava fields, or in fau (Hibiscus tiliaceus) forest, from
altitudes of near sea level to 680 m. (summit of Mt. Fao).

Loca NAMEs: Taitai-ipu (Christophersen); taipoipo (Reinecke,
on Upolu); fatimatao (Powell, on Upolu); lawmafatifati (Setchell, on
Tutuila, and also Powell).

SAMOA: Sava: Matautu, Vaupel 222 (Bish, K, NY, US); between Letui
and Aopo, Christophersen 906 (Bish); Matavanu lava field, Christophersen &
Hume 1898 (Bish) ; Safotu-Manase, Christophersen & Hume 2451 (Bish, UC, US);
near Salaeula, lava flow of 1905-1911, Christophersen 2467 (Bish, NY); without
precise locality, Graeffe 1613 (K isosyntype). Upoiu: Summit of Mt. Fao,
Christophersen 556 (Bish, NY, UC); without precise locality, Graeffe 18 (BM),
1393 (K isosyntype), 1614 (K). Tutuila or Upolu: Powell 257 (K). Turura:
Near Nuuli, Setchell 117 (Bish, UC); Breakers Point, Seale in 1929 (NY); Pango
Pango harbor, Diefenderfer 29 (Bish); Pango Pango and vicinity, Yuncker 9336
(Bish), Kuntze 23007 (NY), Meebold 12646 (Bish, K), Setchell 264 (UC), Col-
larino (Setchell no.) 578 (UC). Oru: Yuncker 9453 (Bish, US), 9477 (Bish,
UC, US). Tau: South of Siufaga, Yuncker 9005 (A, Bish, US); near Siufaga,
Yuncker 9192 (Bish). Samoa, without locality, Graeffe sn. (GH); U.S. Expl.
Exped. (source of the name G. rupestre var. macrophyllum, GH, NY, US).

UVEA (Wallis Islands): Without other locality, Graeffe 42 (BM).

34 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

13b. Geniostoma samoense var. parviflorum Reinecke in Bot. Jahrb. 25: 666,
as var. parviflora. 1898; Gilg & Benedict in Bot. Jahrb. 56: 544, as var.
parviflora. 1921.

PLATE 2, FIGURE 26; PLATE 3, FIGURES 5, 6

Leaf-blades comparatively small, usually 3-111.5-4.5 em.,
cuneate at base, long-acuminate at apex, pale beneath, with obvious
venation ; anthers minutely mucronulate with a tip about 0.1 mm. long;
stigma depressed, about 0.7 mm. in diameter, broader than high.

TypE LocaLity: Upolu, Samoa; the type is Reinecke 555, which we
have not seen. Gilg & Benedict mentioned the specimen (in 1921) as
moldy and useless, but nevertheless they opined that it might represent
a distinct species.

DistrisutTion: Samoa, known only from Savaii and Upolu; the
material available to us is all from the former island. The variety is
recorded as a small tree or shrub up to 10 m. high, with white flowers
and green fruits. It usually occurs between 1,000 and 1,400 m.
altitude, but on occasion as low as 200 m.

SAMOA: Savam: Matavanu crater, Christophersen 609 (Bish, US) ; Matavanu
lava field, Christophersen & Hume 1878 (Bish, UC, NY); above Matavanu,
Christophersen & Hume 2200 (Bish, NY); Letui, Christophersen 775 (Bish, UC,

US); above Salailua, Christophersen 2675 (A, Bish, UC, US), 3118 (Bish, UC,
NY); Siuvao-Auala, Christophersen 3375 (Bish).

Although there is a strong difference in general aspect between the
two varieties here recognized, no significant floral characters appear
to distinguish them. The flowers in both are about the same size,
and why Reinecke selected the varietal epithet “parviflora” is not
clear. His original description of the variety is not very helpful, but
it seems definitely to refer to the small-leaved form with long-acumi-
nate leaf-apices.

14. Geniostoma rupestre J. R. & G. Forst. Char. Gen. 24. t. 12. 1776, Beschr.

Gatt. Pfl. ¢. IT, f. 12. 1779; Willd. Sp. Pl. 1: 998. 1797; Spreng. Pl. Min. Cog.

1: 18. 1813, Syst. Veg. 1: 588. 1825; Endl. Gen. Pl. 576. 1838; DC. Prodr.

9: 26. 1845; Benth. in Journ, Linn. Soe. Bot. 1: 97. 1857; F. v. Muell. Contr.

Phytog. New Hebr. 13. 1873; Gilg & Benedict in Bot. Jahrb. 56: 542. 1921;

Guillaumin in Bull. Soc. Bot. Fr. 72: 701. 1927, in op. cit. 76: 299. 1929, in
Journ. Arnold Arb. 13: 22. 1932, in Journ. Linn. Soc. Bot. 51: 557. 1938.

PLate 1, FIGURE 14

Geniostoma “densiflora” sensu Guillaumin in Bull. Mus. Hist. Nat. [Paris] II.

9: 295. 1937, non Baillon (1880).

Shrub or small tree to 10 m. high, glabrous throughout except for
some floral parts, the branchlets slender, subterete or somewhat com-
pressed-quadrangular, 1-2 mm. in diameter distally; stipules intra-
petiolar, forming a subtruncate or obtusely deltoid sheath 1-1.5 mm.
high; leaves glabrous, petiolate, the petioles slender, 5-10 mm. long,
the blades thin-chartaccous to subcoriaceous, ovate to elliptic-ovate,

SMITH & STONE—PACIFIC ISLAND PLANTS, XVII 35

often rather narrowly so, 3-14 cm. long, 1-4 cm. broad, acute and
decurrent or rarely somewhat rounded at base, acuminate or obviously
cuspidate at apex, entire, the costa sulcate above, raised beneath,
the lateral nerves 5-9 per side, slightly raised on both surfaces, the
veinlets plane, mostly obscure; inflorescences axillary among the
leaves, dichasial, 5-15-flowered, glabrous, the peduncle short, flattened,
1-2 mm. long, bibracteate at apex, the bracts lanceolate, to 1.5 mm.
long, the axes slender, 2-5 mm. long, bibracteate at nodes, the pedicels
2-5 mm. long, slightly flared and bracteolate below calyx; flowers
externally glabrous, borne in triflorous (or irregular) dichasia; calyx
cupuliform, the lobes deltoid-ovate, 0.7-1 mm. long, 3-nerved; corolla
2.5-3 mm. long, short-tubular, slightly urceolate, with reflexed (?)
lobes, divided to middle, the tube glabrous within at base, sparsely
pilose above and around throat with white reflexed trichomes 0.5-0.8
mm. long, these also on filaments, the lobes glabrous distally within,
sparsely pilose with similar trichomes toward extreme base along
nerves, the venation comparatively complex, each lobe supplied by a
central nerve producing a lateral branch on each side near base of
tube or sometimes higher, the branches ascending into the lobe and
there often with brief lateral branches, occasionally the lobe supplied
by two equal (or very rarely by three equal) veins ascending distinct
from base of tube; stamens inserted at throat, the filaments deltoid-
ligulate, 0.3-0.4 mm. long, pilose, the anthers narrowly oblong-deltoid,
about 0.8 mm. long, minutely apiculate at apex, distally or entirely
glabrous; ovary glabrous, subglobose, 0.6-0.7 mm. in diameter, the
style filiform, glabrous, 0.6-0.7 mm. long, the stigma globose or sub-
globose, about 0.5 mm. in diameter, capsule 7-10 mm. long, 3-4 mm.
broad.

Typr LocaLity: Tanna, New Hebrides; type collected by J. R. &
G. Forster on Cook’s second voyage, cited below.

Distrinution: Known with certainty from the New Hebrides
(several islands) and the Santa Cruz Islands; also reported from New
Caledonia, as discussed below. The specimens available to us are
noted as shrubs or small trees to 10 m. high; the flowers are said to
emit a strong foctid odor that attracts flies. Adequate altitudinal
data are not available, but presumably G. rupestre is a lowland species.

SANTA CRUZ ISLANDS: Vanrxoro: Kajewski 650 (A, UC, US).

NEW HEBRIDES: Esprriru Santo: Mt. Tabwemanana, in open grassland,
I. & Z. Baker (Oxford Univ. Exped.) 29 (BM). Amsrym: Mt. Touo, de la Rtie
(2nd Voy.) in 1935-36 (A). Erromanga: Wuppenbu (?), Cheesman 20 (K).
Tanna: J. R. & G. Forster 103 (BM type, K); ‘Herb. Pallas’’ (coll. Forster?)

(BM); W. Anderson in 1774 (BM). Anerryum: Anelgahaut Bay, Kajewskt 728
(A, Bish, NY, US).

36 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Genvostoma rupestre has been the traditional “catch-all” for Pacific
specimens of the genus, even though Gilg & Benedict in 1921 ex-
pressed the opinion that it was a local species limited to the New
Hebrides. They apparently had access to some portion of the origi-
nal Forster material from Tanna. At this time we cannot be certain
as to the precise identity of the several New Caledonian sheets re-
ferred to G. rupestre by Guillaumin, but it is quite probable that the
range of the species does extend to that island and perhaps also to the
Solomons. However, on the basis of existing material we feel assured
that it does not occur east of the New Hebrides.

Since G@. rupestre is the type species of the genus, some importance
is attached to the Forster material, fortunately preserved in good
condition. The four specimens cited above from Tanna are so similar
in all respects that one may believe them to be parts of the same
Forster collection. As holotype we designate the British Museum
sheet that is clearly labeled as a Forster collection; previously none
of these sheets has been so designated.

Among the species with comparatively long corolla-throat trichomes,
G. rupestre is readily distinguished from its only close allies, for in-
stance G@. samoense and G. insulare, by its glabrous habit and usually
acuminate leaf-blades. A closer superficial resemblance exists be-
tween G. rupestre and the Fijian G. macgregorii, but the indument of
the corolla in these two species is uniformly and strikingly different,
even though this character is not obvious upon superficial examination.

15. Geniostoma insulare Smith & Stone, sp. nov. PuaTE 1, FIGURE 15

Frutex vel arbor gracilis ad 5 m. nonnunquam ad 15 m. alta praeter
corollae partem interiorem demum glabra, vel innovationibus ramulis
petiolis foliorum costa subtus inflorescentiae axibusque arcte puberulis
(pilis patentibus rufis circiter 0.1 mm. longis), ramulis juventute saepe
quadrangularibus 1-3 mm. crassis demum subteretibus glabrisque;
stipulis intrapetiolaribus in vagina 1-1.5 mm. alta rotundata vel
obtusa conjunctis; foliis plerumque comparative parvis, petiolis 3-9
(-11) mm. longis complanatis superne subalatis, laminis chartaceis vel
subcoriaceis, ovatis vel ellipticis interdum elliptico-obovatis raro
obovatis, 2-12 cm. longis, 1-7 cm. latis, vulgo circiter 52.5 cm.,
basi rotundatis subtruncatis obtusis vel late acutis et plus minusve
decurrentibus, apice rotundatis vel acutis interdum minute mucronatis
vel cuspidatis vel retusis, margine integris, costa supra sulcata subtus
elevata, nervis lateralibus supra saepe subdepressis subtus prominulis,
venulis subobscuris; inflorescentiis axillaribus dichasialibus 5-15-floris,
pedunculo 1-3 mm. longo, axibus pedicellisque 2-5 (-7) mm. longis
interdum puberulis, bracteis inconspicuis; calyce cupuliformi, lobis

SMITH & STONE—PACIFIC ISLAND PLANTS, XVII 37

deltoideo-ovatis circiter 11.3 mm. 1-3 (-5)-nervatis; corolla
breviter tubulosa 3-3.5 mm. longa fere ad medium lobata, tubo
faucibus pilis pallidis retrorsis 0.4-0.8 mm. longis hirtello, lobis del-
toideo-ovatis circiter 1.5 mm. longis latisque supra basim glabris,
nervatio subintricato, corollae quoque loba nervo unico cum nervis
lateralibus 2 plus minusve tubi medio ornato praedita, nervis 3 sub-
parallelibus vel curvatis in lobam adscendentibus et ibi venulas breves
laterales paucas gerentibus; staminium filamentis ad 0.2 mm. longis
pilosis, antheris deltoideis circiter 0.8 mm. longis apice acutis vel
minute mucronulatis; ovario subgloboso 0.7-0.9 mm. diametro, stylo
ad 0.6 mm. longo, stigmate subclavato-obovoideo inconspicue bilobato
papilloso 0.7-0.9 mm. longo; capsula ellipsoidea vel sphaeroidea, 5-14
mm. longa, 3-5.5 mm. lata, apiculata, seminibus ovoideis circiter 1.2
0.7 mm. minute papillosis.

The species here described as new appears to be the most abundant
Geniostoma in Tonga, and in some forms it occurs more sparingly in
both Fiji and Samoa. Specimens which we now refer here have been
variously identified in herbaria as @. rupestre, @. vitiense, and G.
calcicola. Of these species, G@. insulare is most closely allied to G.
rupestre, from which it differs in its quite dissimilar foliage, the leaf-
blades being most often rounded to obtuse at apex and only occasion-
ally broadly acute or bluntly cuspidate. In general the veins supply-
ing corolla-lobes branch high in the tube rather than near or at the
base as is frequent in G. rupestre, but this character is not entirely
dependable. Our var. tongense of the new species, in its closely but
harshly puberulent vegetative parts, suggests G’. calcicola, which we
now believe to be represented only by the type collection from Fulanga,
Fiji. However, G. insulare never has its foliar indument extending
to both surfaces of the leaf-blades, this being limited even in var.
tongense to the ventral costae and a few of the laterals. The complex
involving G. insulare, G. rupestre, and G. calciola is not satisfactorily
understood from available material and the present solution is ad-
mittedly provisional, pending further observation or experiment.
Two varieties, one of which has three quite distinct forms, are dis-
tinguished as follows:

Vegetative parts glabrous... ..... 1 1 ew ee ee ew var. insulare

Leaf-blades elliptic or ovate, rarely somewhat obovate, rounded or merely

acute at base and apex, not cuspidate.
Capsules ellipsoid, comparatively narrow; Tonga and Fiji. . .f. insulare
Capsules sphaeroid; Fiji. . 2... .......042. f. sphaerococcum

Leaf-blades mostly obovate, gradually decurrent at base, rounded or broadly

acute and cuspidate at apex;Samoa.......... f. cuspidatum

Vegetative parts (innovations, branchlets, petioles, and usually leaf-blades on
ventral costae) puberulent; Tonga and Niue. ....... var. tongense

38 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

15a. Geniostoma insulare var. insulare

Varietas typica; partibus vegetativis glabris; foliorum laminis
plerumque ellipticis vel ovatis raro subobovatis, basi obtusis vel
rotundatis interdum subdecurrentibus, apice rotundatis vel obscure
mucronatis vel late acutis; capsulis ellipsoideis vel sphaeroideis.
15a (1). Geniostoma insulare var. insulare forma insulare

Geniostoma rupestre var. ellipticum A. Gray in Proc. Amer. Acad. 4: 33, nomen.

1859.

Forma typica; foliorum laminis plerumque ellipticis vel ovatis;
capsulis ellipsoideis.

Type in the herbarium of the University of California, No. 296979, collected
at edge of mangrove swamp near Nukualofa, Tongatabu, Tonga, near sea level,
June-August, 1926, by W. A. Setchell & H. E. Parks (No. 15334). Duplicates
at Bish, GH, US.

ADDITIONAL SPECIMENS EXAMINED:

TONGA: Ha’anat Group: Kao: Yuncker 15900 (US). Nomuxa: Seaside
thicket, Yuncker 15821 (US). Nruatroruraru: Summit of range east of Vaipoa,
Mirlimann 366 (US). Taran: Path to the Piu’o Tafahi, Htirlimann 432 (US).
Ton@ataBu Group: Tona@atasu: Peninsula west of Hofoa, Hurlimann 83 (US);
Vahe Ha’ake, Setchell & Parks 15357 (UC); near Ha’atafu, Setchell & Parks
15575 (UC); below Ha’akame village, Yuncker 15284 (US); near Niutoua, Yuncker
15111 (US); without precise locality, U.S. Expl. Exped. (source of the name G.
rupestre var. ellipticum, GH, NY), Lister in 1889 (K), Setchell & Parks 15231
(UC), 15247 (UC). Tonga, without further locality: D. Nelson (BM).

FIJI: Vanua Levu: Mathuata: Seanggangga Plateau, in drainage of Korovuli
River, near Natua, Smith 6868 (A, Bish, K, US). Thakaundrove: Maravu, near
Salt Lake, Degener & Ordonez 14170 (A, Bish, K). Futanaa: Limestone forma-
tion on lagoon cliff, Smith 1203 (Bish, GH, K, NY, UC, US).

Disrrinution: Tonga and Fiji; the plant is recorded as a low shrub
or slender tree not much exceeding 3 m. in height, usually occurring
in thickets near sea level or in xerophilous or fog forest upward to
500 m., frequently on limestone; the flowers are white or greenish
white.

LocaL NAMES AND use: Te’e pilo a Maui (recorded by various
collectors in Tonga); faefaelunga (Niuatoputapu); mbitimbiti (Ful-
anga). Hiirlimann indicates that the plant is used in Tongan medi-
cine.

As the nomenclaturally typical form and variety of G. insulare we
have indicated the population with glabrous habit, prevailingly elliptic
or ovate leaf-blades, and ellipsoid capsules.
15a (2). Geniostoma insulare var. insulare forma sphaerococcum Smith «&

Stone, f. nov. PLATE 2, FIGURE 20

Forma foliorum laminis ovatis acutis saepe basi rotundatis vel sub-
truncatis; e forma typica capsulis sphaeroideis distinguitur.

Type in the U.S. National Herbarium, No. 1674613, collected in hills above

Namalata and Ngaloa Bays, Kandavu, Fiji, alt. 200-400 m., October 14, 1933,
by A. C. Smith (No. 112). Duplicates at Bish, GH, K, NY, UC.

SMITH & STONE—PACIFIC ISLAND PLANTS, XVII 39

ADDITIONAL SPECIMENS EXAMINED:

FIJI: Virt Levu: Naitasiri: Woods near Tamavua, Gillespie 3610 in part (NY);
near road beyond Nasinu, Gillespie 3610 in part (Bish, UC). Ovatau: Northwest
of Levuka, Gillespie 4566 (Bish, NY, UC). Kanpavu: Hills above Namalata
and Ngaloa Bays, Smith 71 (Bish, GH, K, NY, UC, US); without precise locality,
Tothill 614 (K). Vanua Mpatavu: Nambavatu, Tothill 615 (K). FuLanaa:
Limestone formation, Smith 1142 (Bish, GH, K, NY, UC, US). Fiji, without
further locality: U.S. Expl. Exped. (US).

DistrisuTion: Fiji, known from the several islands mentioned
above, where it occurs as a tree 5-15 m. high from sea level up to 400
m. alt., often in rocky formations or on limestone.

Locaut NAME: Mboimboinda (Kandavu, Fulanga).

The cited specimens differ from the typical form of our var. insulare
primarily in having consistently globose or sphaeroidal capsules; the
leaf-blades also tend to be slightly different in shape, being predomi-
nantly ovate and rounded or nearly truncate at base. However, the
indicated nomenclatural position seems best to show the relationship
within Geniostoma of this puzzling population, on the basis of available
material. In form the capsules suggest those of G. dictyoneurum,
described above, from which G. insulare differs in the free veinlets of
its corolla-lobes and in the smaller, more coriaceous, and usually more
acute leaf-blades.
15a (3). Geniostoma insulare var. insulare forma cuspidatum Smith & Stone,

f. nov.

Forma foliorum laminis plerumque obovatis interdum ellipticis
apice cuspidatis et capsulis late ellipsoideis e forma typica differt.

Type in the herbarium of the Bernice P. Bishop Museum, collected on the sum-
mit of Mt. Fao, Upolu, Samoa, alt. 680 m., September 6, 1929, by Erling Christo-
phersen (No. 569). Duplicate at UC.

ADDITIONAL SPECIMENS EXAMINED:

SAMOA: Savau: Near Salaeula, Christophersen 2467 (Bish). Tau: Nearsummit
of main range, Garber 755 (Bish). Samoa, without further locality: U.S. Expl.
Exped. (US), Whitmee in 1876-77 (GH).

Distribution: Known only from the few scattered cited Samoan
collections, at altitudes up to 680 m.; the plants are said to be shrubs
or trees up to 5 m. high.

Although the cited Samoan material seems clearly to belong to the
typical variety of @. insulare, it represents a local variant which, at
least for the present, seems worthy of formal recognition.
15b. Geniostoma insulare var. tongense Smith & Stone, var. nov.

Planta e varietate typica partibus vegetativis et inflorescentiae
ramulis arcte puberulis distinguitur; ramulis demum glabratis; foliorum
laminis ex ovatis ellipticis utroque subacutis interdum subrotundatis;
capsulis ellipsoideis.

40 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Type in the herbarium of the University of California, No. 297257, collected
on the plateau on Eua, Tonga, in June or July, 1926, by H.E. Parks (No. 16261).
Duplicates at Bish, GH, US.

ADDITIONAL SPECIMENS EXAMINED:

TONGA: Vava’u Group: Vava’u: Coastal cliff above Leimatua, Yuncker
16057 (US). Tonaatasu Group: Eva: Above Honuma, Yuncker 15505 (US);
above Fuai, Yuncker 15379 (US), Hirlimann 190 (US); cliffs at Likio, Parks
16097 (UC), 16395 (UC); terrace north of Vaingana Creek, Htirlimann 275 (US);
without precise locality, Lister in 1889 (K), Cartwright in 1889 (K).

NIUE: Rocks near Alofi, Yuncker 10152 (Bish).

DistripuTion: Tonga and Niue; the variety is recorded as a shrub
or as a small tree approaching 5 m. in height and with a trunk 5-18
cm. in diameter, occurring in the forests of level places or along lime-
stone cliffs, elevations of sea level up to 240 m. having been indicated;
the flowers are white or greenish, with a disagreeable odor.

Locau NAME: Te’e pilo a Maui has been recorded in Tonga, as for
var. wnsulare.

As implied in our discussion of the species as a whole, these con-
spicuously puberulent individuals from Tonga and Niue comprise a
population of G. insulare that seems worthy of varietal recognition.
16. Geniostoma biseriale Rechinger in Rep. Sp. Nov. 6: 325. 1909, in Denkschr.

Akad. Wiss. Wien 85: 329. ¢. 9, f. 2. 1910; Christophersen in Bishop Mus.
Bull. 128: 175. 1935,
PLATE 1, FIGURE 16

Shrub or small tree 2-4 m. high, glabrous throughout except for
some floral parts, with slender pale gray-brown subterete or obscurely
quadrangular branchlets, these distally about 2 mm. thick; stipules
intrapetiolar, forming a short subtruncate sheath 1-1.2 mm. long;
leaves subsessile, the petioles at most 2 mm. long, the blades oblong-
elliptic or narrower and lanceolate-elliptic (rarely slightly lanceolate-
ovate), subauriculate, cordate, or subcordate at base, acuminate at
apex, 6-15 cm. long, 2-6 cm. broad, the costa sulcate above, raised
beneath, the lateral nerves 7-12 per side, slightly raised on both sur-
faces, the veinlets obscure; inflorescences axillary among the leaves,
dichasial, 5-10-flowered, the peduncle 1-2 mm. long, the axes and
pedicels glabrous, 1-3 mm. long, bracteate, the flowers glabrous ex-
ternally; calyx cupuliform, the lobes 0.8-1 mm. long, deltoid-ovate;
corolla short-tubular, 3.5-4.2 mm. long, lobed nearly to middle, the
tube glabrous basally within, at throat sparsely pilose with trichomes
0.3-0.4 mm. long, the lobes deltoid-ovate, glabrous distally, the
venation complex, that of each lobe derived from a single central main
nerve producing 1 or 2 pairs of lateral branches in the tube, these in
turn producing 1 or 2 lateral branches, all ascending into the lobe;
stamens inserted at throat, the filaments about 0.2 mm. long, the an-
thers narrowly deltoid-oblong, 0.6-0.8 mm. long; ovary glabrous,

SMITH & STONE—PACIFIC ISLAND PLANTS, XVII 4]

depressed-globose, about 0.7 mm. high and 0.9 mm. broad, the style
slender, glabrous, 1.5-2 mm. long, the stigma (in our specimens
poorly preserved) apparently obovoid, about 0.7 mm. long; capsule
ellipsoid, 7-8 mm. long, 2-4 mm. broad, apiculate.

Typr Locauity: Upolu, Samoa, in forest near Tiavi, at 700 m. alt.;
the type is Rechinger 446, presumably deposited in the Berlin Her-
barium and subsequently destroyed. The cited illustration is excel-
lent and permits adequate understanding of the concept.

Distripution: Endemic to Samoa, and thus far known from few
collections from Savaii and Upolu, occurring in forest between 400 and
1500 m. alt.; it is said to be a shrub or small tree up to 4 m. high,
with white flowers.

SAMOA: Savam: Papa’afu, crater-rim, Christophersen 2727 (Bish, UC, NY);
above Sili, Christophersen 3152 (Bish, UC, NY). Upouiu: Ridge above Malo-
lolelei, Christophersen 11 (Bish, US).

Geniostoma biseriale is readily distinguished from other species of
our region by its essentially sessile leaves, of which the petioles do not
exceed 2 mm. in length, the blades being proportionately unusually
narrow, prevailingly cordate or even subauriculate at base. The indu-
ment of the corolla-throat is short, suggesting that the closest relatives
of G. biseriale may be G. gracile and G. macgregorii, but the relation-
ship is not close.

Excluded Species

Geniostoma microphyllum Seem. in Bonplandia 10: 37, nomen. 1862, Fl. Vit.
164. 1866.

FIJI: Virt Levu: Namosi (?): “Navua River,” Seemann 304 (BM, GH, K
type); Rewa: Summit of Central Road, Suva, Tothill 611 (K).

This taxon is not a Geniostoma, but appears to be rubiaceous. We
judge it to be a juvenile form of a Psychotria, probably of a lianoid
species, suggestive of P. serpens L. The Seemann specimen was men-
tioned by A. Gray (in Proc. Amer. Acad. 5: 320. 1862), who stated
that the specimen, first labeled as a Gaertnera, was a new Gentostoma.
The nature of the stipules, the leaf structure, and the apparent climb-
ing habit (adventitious roots appearing at the nodes of the stem)
exclude this plant from Geniostoma. We hope to be able to state
more precisely the identity of this puzzling plant, but are not pre-
pared to do so at this time.

U.S. GOVERNMENT PRINTING OFFICE: 1968

PLATES

PLATE 1

Corolla-lobe venation-patterns in Gentostoma. The types of venation of 16 species are
shown, all 5; each figure illustrates 3 lobes and the corresponding part of the corolla-
tube. Ficures 1, G. macrophyllum; 2, GC. stipulare; 3, G. confertiflorum; 4, G. clavigerum;
5, G. univervium; 6, G, vitiense; 7, G. gracile; 8, G. macgregorii; 9, G. calcicola; 10, G. dictyo-
neurum; LL, G. fletschmannti; 12, G. stenocarpum; 13, G. samoense; 14, G. rupestre; 15, G
insulare; 16, G. bisertale.

CONTR. U.S. NAT. HERBARIUM, VOL. 37 SMITH & STONE, PLATE 1

[For explanation see opposite page.]

CONTR. U.S. NAT. HERBARIUM, VCL. 37 SMITH & STONE, PLATE 2

PLATE 2

Floral features of Geniostoma. Ficures 1-3, G. macrophyllum: 1, portion of inflorescence,
S25: 2, gynoecium, * 10; 3, stamen, * 10. FicuRres 4-9, G. stipulare: 4, ultimate dichas-
ium, X 5; 5, stigma, showing lobes, * 5; 6, ey noecium, 5; 7, stamens, X 5; 8, capsule
before dehiscence, * 2.5; 9, capsule after dehiscence, ~ 2.5. Ficures 10-14, G. uniner-
vium: 10, inflorescence with flowers, * 5; 11, schematic diagram of youn inflorescence,
% 5; 12, stamen and gynoecium, * 2.5; 13, calyx, showing venation, * 2.5; 14, capsule in
outline, S<.2.5. Ficures 15, 16, G. confertiflorum: 15, portion of oe porescence with flower,
2.5: 16, eynoecium, * 2.5. Ficures 17-19, G. macgregortt: 17, part of inflorescence
with flower and gynoecia, * 2.5; 18, gynoecium, * 2.5; 19, schematic diagram of inflores-
cence, 2.5. Ficure 20, G. insulare {. sphaerococcum: capsule and part of inflorescence,
S25, Ficures 21-23, G. vitiense: 21, inflorescence, 2. 5; 22, 23, capsules, X 2.5. Fic-
urE 24, G. calcicola: schematic diagram of inflorescence, * 2.5. Ficure 25, G. samoense
var. samoense: inflorescence and flowers, X 2.5. Ficure 26, G. samoense var. parviflorum:
inflorescence and flowers, X 2.5. Ficures 27, 28, G. samoense var. samoense: 27, greatly
enlarged view of inflorescence-node with bract; 28, capsule, X 2.5

PLATE 3

Floral features of Geniostoma, Figures 1-4, G. samoense var. samoense: 1, stamen,
5; 2, flower past anthesis, with pedicel, < 5; 3, seed in side view and cross-section, * 10;
4, portion of wall of seed in cross-section, * 50. Ficures 5, 6, G. samoense var. parviflorum:
5, stamen, * 5; 6, gynoecium, “ 35. Ficures 7, 8, G. stenocarpum: 7, gynoecium, 5;
8, schematic diagram of inflorescence, * 2.5. Figures 9-11, G. gractle: 9, two inflores-
cences, one with flowers at anthesis, the other slightly later, x 2.5; 10, stamen, & 5; 11,
gynoecium, < 5. Figures 12-15, G. vitiense: 12, 13, stamen and gynoecium from “long-
styled” or hermaphrodite flower, & 5; 14, 15, stamen and gynoecium from “‘short-styled”

or pistillate flower, * 5.

CONTR. U.S. NAT. HERBARIUM, VOL. 37 SMITH & STONE. PLATE 3

[For explanation see opposite page.]

UNITED STATES NATIONAL MUSEUM

CONTRIBUTIONS FROM THE UNITED States NationaAL HERBARIUM

Vo.LuME 37, Part 1

STUDIES OF PACIFIC ISLAND PLANTS, XVII
THE GENUS GENIOSTOMA (LOGANIACEAE)
IN THE NEW HEBRIDES, FIJI, SAMOA, AND TONGA

By AvBert C. Smita and
BENJAMIN C. STONE

SMITHSONIAN INSTITUTION e WASHINGTON, D.C. e 1967

(Synonyms in italics. Page numbers of principal entries in boldface.)

Canthium barbatum korrorensis, 25

Cyrtandra, 13

Faefaelunga, 38

Faifailunga, 20

Fatimatao, 33

Fau, 33

Fifilunga, 20

Gaertnera, 41

Genfostoma, 7
arboreum, 7
australianum, 4
avene, 4
batanense, 25
biseriale, 4, 40
calcicola, 3, 4, 21, 26, 37
caulocarpum, 3, 12
clavatum, 4
clavigerum, 5, 15, 16, 18, 21
confertiflorum, 5, 14, 17, 18
crassifolium, 3, 27

glaberrimum, 18

densiflora, 34
dictyoneurum, 5, 7, 27, 39
fleischmanni, 28
fleischmannii, 3, 28
glabrum, 7
gracile, 4, 22, 41
haemospermum, 7
hoeferi, 5

insulare, 3, 4, 6, 21, 26, 27, 28, 31, 36

insulare, 38
cuspidatum, 39
insulare, 38
sphaerococcum, 38

sphaerococcum, 7

tongense, 37, 39

ligustrifolium, 7
longistylum, 5

macgregorii, 6, 16, 21, 23, 28, 36, 41
macrophyllum, 3, 4, 6, 10, 13, 14,

15, 21
42

Geniostoma—Continued

micranthum, 5
microphyllum, 41
miquelianum, 7
niinoanense, 25
petiolosum, 7
rapense, 4
rupestre, 1, 2, 6, 15, 16, 18, 21, 23,
25, 31, 32, 34, 37
macrophyllum, 33
puberulum, 18
samoense, 31, 36
parviflora, 34
parviflorum, 6, 23, 34
samoense, 33
sessile, 4
stenocarpum, 3, 6, 21, 29
stipulare, 3, 11, 12, 15
uninervium, 5, 15, 16, 17
vitiense, 5, 6, 16, 18, 23, 24, 25, 28,
31, 37

Hibiscus tiliaceus, 33
Kavikaloa, 18

Labordia, 2
Laumafatifati, 33
Mbatimara, 20, 30
Mbati-mbati, 30
Mbatimbona, 24
Mbatimboni, 20
Mbitimbiti, 38
Mboimboinda, 18, 20, 24, 28, 39
Mbuimbuita, 30
Muskarimhba, 13
Plectronia korrorensis, 25

macgregori, 23

Psychotria, 41

serpens, 41

Taipoipo, 22, 33

Taitai-ipu, 33

Teatea, 20

Te’e pilo a Maui, 20, 38, 40
Tete, 20

UNITED STATES NATIONAL MUSEUM

CONTRIBUTIONS FROM THE UNITED STATES NATIONAL HERBARIUM

VoLuME 37, ParT 2

MOSSES OF THE EASTERN HIGHLANDS, NEW GUINEA,
FROM THE 6th ARCHBOLD EXPEDITION, 1959

By Epwin B. Bartram

BULLETIN OF THE Unitrep States NationaL MusSEuM
SMITHSONIAN INSTITUTION e WASHINGTON, D.C. e 1965

MOSSES OF THE EASTERN HIGHLANDS, NEW GUINEA, FROM
THE 6th ARCHBOLD EXPEDITION, 1959

By Epwin B. Bartram’

The moss collections, represented by 375-400 numbers, made by
L. J. Brass, H. M. van Deusen, and J. D. Collins in the Eastern
Highlands of northeastern New Guinea have been acquired by the
Smithsonian Institution. These were entrusted to me for determina-
tion. They are a welcome and important supplement to the collection
made by Ross G. Robbins in the Western Highlands. The results
bring into sharp focus a number of significant aspects of the New
Guinea moss flora.

As a result of active field work by Mr. Brass and Mr. Robbins this
insular moss flora has been expanded in recent years to about 850
species. ‘This is a large number for an area the size of New Guinea
and it emphasizes the astonishing richness of this tropical flora.
Every new collection adds new records and new species to a total
which may eventually reach or exceed 1,000 species. The rugged
terrain of the interior, with some peaks reaching altitudes of 14,000
to 16,500 ft., provides a wide variety of habitats which increase the
interest and complexity of the flora. While far from complete, our
present knowledge is broad enough to form the basis for some general
conclusions. The percentage of endemism is abnormally high.
Between 350 and 400 species, or nearly half of the known flora, have
been described as new from New Guinea. Critical studies may re-
duce the figure to some extent, but the ratio of endemic species is
likely to remain disproportionally high. Of the remaining 400 to 500
species by far the greater part is broadly representative of the Indo-
Malayan areas to the west from where the bulk of the present flora
has been derived. A limited number of species are common to the
New Zealand-Australian area, but the affiliation in this direction is
rather limited. A few species from the alpine crests of the higher
peaks such as Calliergon sarmentosum, Calliergon wilhelmense (near

1Kdwin B. Bartram, 1878-1964. Shortly before his death the author trans-
ferred his personal herbarium to the Farlow Herbarium at Harvard University
where he had long been honorary curator of mosses.

43

44 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

C. turgescens), and Meesea triquetra show such disjunct geographical
distribution as to preclude any logical explanation. If the North
Pole was located in the south central Pacific ages ago, as has been
suggested, these relicts may represent waifs of circumpolar distribu-
tion that have survived in sanctuary niches, but this is of course only a
random guess. The implications are too complex and extensive for
my modest attainments. The fact remains that the New Guinea
moss flora is predominantly Malaysian in character and probably
represents the southeastern front of a broad pattern of geographical
distribution extending east to Fiji and Samoa.

In the following list, species not previously recorded from New
jJuinea are preceded by an asterisk. A complete series of specimens
is deposited in the herbarium of the United States National Museum,
Washington, D.C., and in the Farlow Herbarium at Harvard University.

Andreaeaceae

Andreaea rupestris Hedw.

Mt. Wilhelm, east slopes: on stones on alpine grassland, 13,500 ft., Van Deusen
29896; cushioned in crevices near summit, 14,500-14,700 ft., Collins 29958, 29959;
cushioned on a rock near summit, 14,700 ft., Van Deusen & Collins 30027.

Andreaea (Enervia) novo-guinensis Bartr. sp. nov.

Robusta, caespitosa, caespitibus densis, fuscescentibus, nitidiu-
sculis. Caulis 5-7 em. altus, parce ramosus, ramis obtusis. Folia
erecta, imbricata, late ovata, profunde concava, obtusa, 1.5-2.5 mm.
longa, 1.2 mm. lata, ecostata; marginibus integris, erectis, superne
inflexis; cellulis superioribus parvis, rotundatis, incrassatis, laevis-
simis, diam. 5-6 y, basilaribus internis linearibus, incrassatis, porosis,
externis parvis, rotundatis. Folia perichaetialia parva caulina simi-
lia; theca in pseudopodio crasso, ovalis folia perichaetialia vix superans.

Type in the U.S. National Herbarium, collected on moist ground
near summit, at 14,700 ft. altitude, east slopes, Mt. Wilhelm, New
Guinea, June 18, 1959, by Van Deusen and Collins (no. 30029).

This remarkable species in an unusual habitat is characterized by
the robust habit, the turgid stems, and the broad, concave, bluntly
pointed leaves.

Fissidentaceae

Fissidens filicinus D. & M.
Kassam: wet, shady bank of a small stream in Castanopsis-oak forest, 1370 m.,
32414,

Ditrichaceae

Ditrichum sericeum Bartr.
Mt. Wilhelm, east slopes, 14,700 ft.: on ground near summit, Collins 29964;
in a moist crevice near summit, Van Deusen & Collins 30032.

BARTRAM—MOSSES OF NEW GUINEA EASTERN HIGHLANDS 45

Dicranaceae

Trematodon longicollis Mx.
Mt. Otto, south slopes, 7,300 ft., 29122.

Campylopodium euphorocladum (C. M.) Besch.
Purosa, Okapa area: terrestrial on roadside in rain forest, 1950 m., 31610.

Dicranella setifera (Mitt.) Jaeg.
Arau: on a clay roadbank in forest, 1400 m., 32069.

Campylopus (Eucampylopus) crispifolius Bartr. sp. nov.

Dense caespitosus, caespitibus fuscescentibus. Caulis brevis, vix
1 cm. longus, dense foliosus; folia sicca arcte flexuosa vel crispata,
humida erecto-patentia, e basi oblonga sensim acuminata, 4 mm.
longa, 0.5 mm. lata; marginibus erectis, integris, apice solum den-
ticulatis; costa inferne 375 y lata; cellulis basilaribus hyalinis, parietibus
teneris, rectangularibus, 12-18 » latis, superioribus parvis, rhomboi-
deis; alaribus nullis. Caetera ignota.

Type in the U.S. National Herbarium, collected on a seepage slope
in alpine grassland, at 3560 m. elevation, east slopes, Mt. Wilhelm,
New Guinea, June 14, 1959, by L. J. Brass (no. 29894).

The absence of alar cells suggests some affinity with C. robbinsi
Bartr., but the flexuose-crispate leaves less slenderly acuminate are
quite distinctive.

Campylopus (Palinocraspis) wilhelmensis Bartr. sp. nov.

Caespitosus, caespitibus lutescentibus intus fuscescentibus. Caulis
erectus, 4-5 cm. altus, simplex vel sparse ramosus, rubro-tomentosus.
Folia sicca erecta, humida flexuoso-patentia, e basi breviter oblonga
sensim subulato-acuminata, 5 mm. longa, ad 0.7 mm. lata; marginibus
erectis, inferne integris, superne remote et minute denticulatis; costa
inferne 300 zu lata; cellulis basilaribus breviter oblongis, 14-18 uy latis,
parietibus firmis, marginalibus e seriebus 4-5 abrupte angustioribus,
alaribus distinctis, fuscis vel hyalinis, sat auriculatis; superioribus
parvis, subquadratis. Caetera ignota.

Type in the U.S. National Herbarium, collected on a tree stump in
moist forest, at 2740 m. elevation, east slopes, Mt. Wilhelm, New
Guinea, July 1, 1959, by L. J. Brass (no. 30237).

The costal structure is similar to that of C. morobensis Bartr., but
the stems are evenly foliate, tomentose and the leaves shorter.
Campylopus comosus Bosch & Lac. (?)

Sterile. Morobe District, Botanic Gardens, Lae: on tree-fern stem supports
of epiphytic orchids, 200 ft., 29111.

Campylopus caudatus (C. M.) Mont.
Mt. Wilhelm, east slopes: on deadwood on alpine grassland, 3560 m., 29891.

46 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Campylopus (Palinocraspis) atro-fuscescens Bartr. sp. nov.

Dense caespitosus, caespitibus inferne nigricentibus superne
lutescentibus. Caulis erectus, dense foliosus. Folia erecta, flexuosa,
humida erecto-patentia, e basi ovata sensim subulato-acuminata,
6 mm. longa, 0.6 mm. lata; marginibus erectis, integris, apice solum
parcissime denticulatis; costa breviter excurrente, inferne circa
0.3 mm. lata; cellulis superioribus parvis, rotundatis, basilaribus
linearibus, porosis margines versus angustioribus, alaribus sub-
quadratis, vix distinctis, fugacissimis. Caetera ignota.

Type in the U.S. National Herbarium, forming cushions on alpine
erassland, at 13,100 ft. elevation, east slopes, Mt. Wilhelm, New
Guinea, June 15, 1959, by J. D. Collins (no. 29941).

Suggestive of C. clemensiae Bartr. in some respects but distinct. in
the darker color, the evenly foliate stems, and the differentiated alar
cells which while present are not well defined.

Campylopus exasperatus Brid.

Mt. Wilhelm, east slopes: on wet ground in alpine grassland, 13,100 ft., Collins
29939.

Campylopus archboldii Bartr.
Mt. Wilhelm, east slopes: on exposed rock in alpine grassland, 3560 m., 29821a;
on rocks of a shady stream in forest, 2770 m., 30450.

Campylopus umbellatus (Arn.) Par.

Morobe District, Wau-Edie Creek road: 5,800 ft., 29162. Kaindi: on clayey
moist roadbank, 2060 m., 29548. Mt. Wilhelm, east slopes: on alpine grassland,
13,100 ft., 29942, 30090a. Mt. Otto: on rocks of a creek bank in forest, 2200 m.,
30839, Mt. Michael, northeast slopes: on exposed rock in forest, 2000 m., 31533.

Atractylocarpus dicranoides Dix.
Mt. Wilhelm, east slopes: on deadwood in subalpine forest, 3560 m., 30045.

*Amphidium cyathicarpum (Mont.) Broth.

Mt. Wilhelm, east slopes: on dry ground under a rock overhang on alpine
grassland, probably fertilized by owl pellets and droppings, 3650 m., Van Deusen
30151.

Distribution: Guatemala, western South America, Australia, New
Zealand, Africa.

*Symblepharis reinwardtii (D. & M.) Bosch & Lac.
Mt. Michael: on rocks of summit, 3650 m., Brass & Collins 31259.

Distribution: Sikkim, Burma, Java, Borneo, Philippine Islands.

Dicranoweisia macrocarpa Bartr.
Mt. Wilhelm, east slopes: cushioned high on a tree in forest, 2770 m., 30470.

Holomitrium stenobasis Dix.
Mt. Wilhelm, east slopes: on a tree in edge of subalpine forest, 3700 m., 30119.

Holomitrium austro-alpinum Bartr.
Mt. Otto: on alpine grassland of the summit, 3540 m., 31038.

Cryptodicranum armittii (C. M.) Bartr.
Arau: on tree trunks, 1400 m., 31960a, 31961.

BARTRAM—MOSSES OF NEW GUINEA EASTERN HIGHLANDS 47

Dicranoloma perarmatum Broth.

Morobe District, Mt. Kaindi: on decayed wood in forest, 7,200 ft., 29165; on
edge of a path in Castanopsts forest, 2000 m., 29526. Mt. Wilhelm, east slopes: on
tree trunk in subalpine forest, 3650 m., 29868.

Dicranoloma laevifolium (Broth. & Geh.) Par.

Mt. Wilhelm, east slopes: in small cushions on tree trunks in subalpine forest,
3650 m., 29869; on wet ground in long-grass community of alpine grassland,
3800 m., 30121.

Dicranoloma dicarpum Hornsch.

Mt. Michael, northeast slopes: on lower trunks of big trees in Castanopsis-
Nothofagus forest, 2000 m., 31504. Purosa, Okapa area: on leaf littered ground
in Castanopsis forest, 2050 m., 31844.

Dicranoloma braunii (C. M.) Par.

Mt. Wilhelm, east slopes: on tree trunks in forest, 2740 m., 30234. Purosa,
Okapa area: on undergrowth in rain forest, 1950 m., 31667; on tree trunks in
Castanopsis forest, 2050 m., 31843. Arau: on tree in Castanopsis forest, 1400 m.,
31976.

Dicranoloma blumii (Necs) Par.

Mt. Michael: on trees in subalpine forest, 3500 m., Brass & Collins 31318.
Braunfelsia enervis (D. & M.) Par.

Mt. Wilhelm, east slopes: on a tree in moist, mossy forest, 2600 m., 30490.

Leucobryaceae

*Leucobryum chlorophyllosum C. M.

Morobe District, Oomsis: on buttress roots of a great Anisoptera tree, 200 m.,
29268a.

Distribution: Ceylon, Java, Philippine Islands, Celebes, Sumbawa,
Timor.
Leucobryum sanctum Hpe.

Purosa, Okapa area: on tree trunks in rain forest, 1950 m., 31693. Kassam:
on logs and bases of trees in Castanopsis-oak forest, 1370 m., 32433.

Leucobryum candidum (Brid.) H. f. & W.
Mt. Otto, south slopes, 2000 m.: low on trees in Castanopsis forest, 31111.

Leucobryum pentastichum D. & M.
Morobe District, Oomsis: on buttress roots of a great Anisoptera tree, 200 m.,

29266, 29267a.
Octoblepharum albidum Hedw.

Morobe District: Botanic Gardens, Lae: on tree-fern stem supports of epiphytic
orchids, 29105.
Leucophanes glaucescens C. M.

Morobe District, Umi River, Markham Valley: low on a tree in rain forest,
480 m., 32584; Botanic Gardens, Lae: on tree-fern stem supports of orchids, 200
ft., 29107b.

Leucophanes octoblepharioides Brid. var. korthelsii (D. & M.) Fleisch.
Morobe District, Botanic Gardens, Lae: 200 ft., 29106a.

48 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Calymperaceae

Syrrhopodon ciliatus (Hook.) Schwaegr.
Morobe District, Botanic Gardens, Lae: on tree-fern stem supports of epiphytic
orchids, 200 ft., 29107.

*Syrrhopodon philippinensis Bartr.

Morobe District, Oomsis: on buttress roots of a great Antsoptera tree, 200 m.,
29268.

Distribution: Philippine Islands.

Calymperes hyophylaceum C. M.
Morobe District, Botanic Gardens, Lae: on tree-fern stem supports of epiphytic
orchids, 200 ft., 29109.

Calymperes moluccense Schwacgr.
Morobe District, Botanic Gardens, Lae: on tree-fern supports of epiphytic
orchids, 29107a, 29110.

Calymperes tahitense (Sull.) Mitt.
Morobe District, Kelly Bros. place on Bulolo Road, about 3 miles west of Lae:
ca. 20 m., between buttress roots of tree, 29104a.

Pottiaceae

Hyophila involuta (Hook.) Jaeg.

Mt. Michael, northeast slopes: on rocks in Castanopsis-Nothofagus forest, 2000
m., 31535, 31537a, 31544. Morobe District, Umi River, Markham Valley: on
moist rock face in rain forest, 480 m., 32627.

Trichostomum angustatum (Mitt.) Fleisch.
Mt. Otto, south slopes: underside of a rock overhang, 2200 m., 30923.

Trichostomum subulifolium Bartr.

Mt. Wilhelm, east slopes: on a tree trunk in subalpine forest, 3560 m., 29847.
Mt. Michael, northeast slopes: on rock in Castanopsis-Nothofagus forest, 2000
m., 31543.

*Leptodontium warnstorfii Fleisch.
Mt. Wilhelm, east slopes: alpine grassland, 13,500 ft., Van Deusen 29895.

Distribution: Java.
*Streptopogon erythrodontus (Tayl.) Wils.

Mt. Wilhelm, east slopes: tufted on exposed branches of a tree in subalpine
forest 3560 m., 30068.

Distribution: Guatemala, Colombia, Ecuador, Peru, Bolivia, Ha-
wall, Madagascar, Africa.

The occurrence of this species in New Guinea presents a problem in
geographical distribution that is hard to explain with any satisfaction.
The species is fairly well distributed along the American Cordillera
from Bolivia to Guatemala. It is also known from the crater of
Haleakala, Maui, Hawaii, and the variety rutenbergii (C. M.) Salm.
from Madagascar and the African continent. This is the only known
record from the southwest Pacific area. The species appears to be of
austral origin and has fanned up through South America and Africa.

BARTRAM—MOSSES OF NEW GUINEA EASTERN HIGHLANDS 49

Grimmiaceae

Grimmia affinis Hornsch.
Mt. Wilhelm, east slopes: cushioned on rocky slopes, 14,700 ft., Van Deusen &

Collins 30026.

This species has been recorded from the Himalayas and Ceylon
at high altitudes under the name of G. ovalis (Hedw.) Lindb., but
according to the studies by Dr. Sayre (Bryologist 55:258, 1952), Horn-
schuch’s name will take precedence. Here the setae are distinctly
curved when dry but strictly erect when moist.

Rhacomitrium crispulum (H. f. & W.) H. f. & W.

Mt. Wilhelm, east slopes: exposed rock in alpine grassland, 3650 m., 29874; in
crevices near summit, 14,500-14,700 ft., J. D. Collins 29961, 29962; on a rock in
alpine grassland, 3560 m., 30086, 30091; terrestrial in alpine grassland, 3800 m.,
30122; on rocks of a shady stream in forest, 2770 m., 30448.

Rhacomitrium lanuginosum (Hedw.) Brid. var. pruinosum H. f. & W.

Mt. Wilhelm, east slopes: alpine grassland, 13,100 ft., Collins 29940; 3700 m.,
Brass 29996.

Funariaceae
Entosthodon subulatus Bartr.

Mt. Wilhelm, east slopes: on wet ground in alpine grassland, 13,100 ft., Collins
29938.

Entosthodon contortus Bartr.

Mt. Michael: wet ground edging small pools on alpine grassland, 3560 m.,
31432 bis.

Funaria hygrometrica Hedw.
Morobe District, Mt. Kaindi: roadbank, 2200 m., 29593. Mt. Wilhelm, cast

slopes: burnt ground, 2770 m., 30673. Purosa, Okapa area: burnt ground, 31841.

Splachnaceae

Tayloria octoblepharis (flook.) Mitt.
Mt. Wilhelm, east slopes: terrestrial in an open place in long-grass community

of alpine grassland, 3850 m., 30118; on log, 2740 m., 30241a.

Bryaceae

Mielichhoferia (Eumielichhoferia) novo-guinensis Bartr. sp. nov.

Paroica: caespitosa, caespitibus densis, superne pallide viridibus,
inferne fuscescentibus, nitidiusculis. Caulis erectus, usque ad 2 cm.
altus, inferne denudatus, dein dense foliosus. Folia imbricata, ovato-
lanceolata, tenuiter acuminata, 2 mm. longa, 0.45 mm. lata; marginibus
erectis, superne minute denticulatis; costa inferne 0.75 y lata, valde
excurrente; cellulis anguste linearibus, infimis brevioribus. Seta 2 cm.
longa; theca erecta vel leniter inclinata, oblongo-cylindrica, deopercu-
lata ad 4 mm. longa; operculum obtuse conicum; endostomium hya-
linum, laevissimum, corona basilaris ad 50 u alta, processus anguste
lineari, 375 u longi, medio ¢ 17 yu lati; spori ferruginei, papillosi, diam.
20 uy.

768-102—65

9
“

50 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Type in the U.S. National Herbarium, collected in a crevice near
summit, at 14,500-14,700 feet elevation, east slopes, Mt. Wilhelm,
New Guinea, June 16, 1959, by J. D. Collins (no. 29966).

This species resembles M. javanica Broth. in some respects, but
the larger, finely acuminated leaves 2 mm. long with the costa excur-
rent in a short arista are good diagnostic characters.

Pohlia elongata Hedw.
Mt. Wilhelm, east slopes: on base of tree in edge of subalpine forest, 3650 m.,

29866.

*Pohlia flexuosa Hook.
Morobe District, Wau-Edie Creek road: on moist, clayey roadbank, 5,800 ft.,

29159.
Distribution: Apparently widely distributed in tropical regions.
Himalayas, West Indies, Central & South America.

Brachymenium nepalense Hook.
Mt. Wilhelm, east slopes: on tree trunk in forest, 2740 m., 30231.

*Anomobryum cymbifolium (Lindb.) Broth.
Mt. Wilhelm, east slopes: on rocks in an alpine grassland stream, 3560 m.,
30089.

Distribution: Himalayas, Khasia, South India, Java, Amboina,
Philippine Islands.
Bryum argenteum [edw.

Morobe District, Mt. Kaindi: open ground, 2250 m., 29680. Mt. Wilhelm,

east slopes: on a stump in alpine grassland, 3560 m., 30085; on open ground in
alpine grassland, 4080 m., 30172; on bare ground in forest 2600 m., 30718.

Bryum porphyroneuron C. M.

Morobe District, Botanic Gardens, Lae: limestone of a rockery, 200 ft., 29114;
Kaindi: on wet rock, 29551; on log, 29552; Gurakor: on log in forest, 640 m.,
29414. Mt. Wilhelm, east slopes: open flood banks of Pengagl Creek, 2770 m.,
30269.

*Bryum australe Hampe.
Morobe District, Kaindi: roadbank in forest, 2060 m., 29681.

Distribution: Australia.

Bryum nitens Hook.

Morobe District: Umi River, Markham Valley, 480 m., 32627a.
Bryum (Doliolidium) wilhelmense Bartr. sp. nov.

Gracile, dense caespitosum, caespitibus pallide fuscescentibus, intus
dense intertextis. Caulis ad 5 cm. altus, ramosus. Folia erecta,
imbricata; folia caulina ovata breviter aristata, ad 0.8 mm. longa,
comalia longiora; marginibus integris, erectis; costa breviter excur-
rente; cellulis superioribus rhomboideis, 35-40 u longis, 10 yu latis,
basilaribus infimis subquadratis. Seta gracilis ad 2.5 cm. longa;
theca horizontalis, deoperculata 2 mm. longa.

BARTRAM—MOSSES OF NEW GUINEA EASTERN HIGHLANDS 51

Type in the U.S. National Herbarium, collected on exposed rock
in alpine grassland, at 3560 m. elevation, east slopes, Mt. Wilhelm,
New Guinea, June 11, 1959, by L. J. Brass (no. 29821). Paratype
at same locality, on rocks in a stream in alpine grassland, Brass 30093.

In some respects this species resembles B. junghuhnianum Hpe. of
Java, but the elongated stems and shorter-pointed longer leaves are
quite distinctive.

Bryum novo-guinense Bartr.

Mt. Wilhelm, east slopes: on rocks in an alpine grassland stream, 3560 m.,
30090b.

Bryum truncorum Brid.

Mt. Wilhelm, east slopes: on a tree in subalpine forest, 3560 m., 30150; ona
tree stump in forest, 2740 m., 30233; lower trunk of tree in mossy forest, 2770 m.,
30349. Mt. Otto, south slopes: on a log in forest, 2480 m., 30902.

Bryum crassulum Bartr.
Morobe District, Kaindi: on a log in forest, 2150 m., 29703.

Rhodobryum giganteum (Hook.) Schp.
Mt. Wilhelm, east slopes: terrestrial in moist forest, 2900 m., 30388.

Leptostomaceae

Leptostomum intermedium Broth.
Mt. Wilhelm, east slopes: on tree in subalpine forest, 3560 m., 29892, Mt.
Otto, south slopes: on dead small tree in forest, 2480 m., 30903.

Mniaceae

Orthomniopsis elimbata Noguchi.

Morobe District, Wau-Edie Creek road: 5,800 ft., on moist, rocky roadbanks,
29161. Mt. Wilhelm, east slopes: on thick terminal branches of a Ternstroemia
tree in forest, 2770 m., 30352.

As no. 30352 is in good fruit, the sporophyte of this species may be
described. Dr. Noguchi was correct in assuming that this species was
an Orthomniopsis. Mnium rotundifolium Bartr., which is exactly the
same thing, will therefore be reduced to synonymy.

Perichaetial leaves spatulate, rounded, unbordered, 7-8 mm.
long. Seta solitary or two from one perichaetium, 5 mm. long, 0.5
mm. thick, slightly curved; capsule nodding, oblong-cylindric, abruptly
contracted to the seta, 4 mm. long, 2 mm. thick; peristome double,
teeth brown, papillose, linear-lanceolate, 0.8 mm. long, endostome
papillose, pale, basal membrane about half the height of the teeth,
segments about equaling the teeth. Operculum, calyptra, and spores
not seen.

Rhizogoniaceae

Hymenodon parvulus Bartr.

Mt. Wilhelm, east slopes: on trees in forest, 2770 m., 30348, 30351.
No. 30348 contains many fruiting plants. The sporophyte may be

52 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

described as follows: Perichaetium small, leaves about 1 mm. long,
broadly ovate, laxly areolate, unbordered, costa excurrent. Seta 12
mm. long, pale brown; capsule erect, relatively large, oblong-cylindrie,
urn 2.5 mm. long, operculum conic-rostrate, 1 mm. long, oblique;
outer peristome lacking, endostome pale, segments 0.3 mm. long from
a rather high basal membrane projecting about 50 u above the rim;
spores pale, smooth, diameter 10 4; calyptra cucullate, small, barely
covering the operculum.

Rhizogonium spiniforme (Hedw.) Bruch.

Morobe District, Oomsis: on buttress roots of a great Anisoptera tree, 200 m.,
29267; Gurakor: base of tree, 640 m., 29505. Arau: on tree roots, 1400 m.,
31926. Kassam: on tree trunks and rotting wood, 1370 m., 32339.

Hypnodendraceae

Hypnodendron diversifolium Broth.
Purosa, Okapa area: on logs in rain forest, 1950 m., 31697. Arau: on surface

roots in oak-Castanopsis forest, 1400 m., 31926a.

Hypnodendron auricomum Broth. & Geh.

Morobe District, Kaindi: on stem of a dead tree fern in forest, 2060 m., 29789.
Mt. Wilhelm, east slopes: terrestrial in deep shade in subalpine forest, 3560 m.,
29849. Mt. Otto, south slopes: terrestrial on edge of a stream in Castanopsis
forest, 2000 m., 31112.

Hypnodendron (Euhypnodendron) brassii Bartr. sp. nov.

Robustum, pallide viride, nitidum. Caulis usque ad 10 cm. altus,
erectus, rigidus, alte stipitatus, dense rufo-tomentosus, stipite foliis
pallidis, adpressis, acuminatis, superne dense ramosus, ramis_bi-
pinnatis, ca. 3 cm. longis, densissime congestis ; folia ramea complanata,
ovato-lanceolata, breviter acuminata; marginibus erectis, superne
grosse et irregulariter serratis; costa tenuis, infra apicem folii evanida,
dorso superne serrata; cellulis linearibus, basilaribus infimis laxioribus.
Setae numerosi, ad 10, rubri, 2.5 cm. longi; theca cum collo cylindrica,
valde plicata, 5 mm. longa.

Type in the U.S. National Herbarium, collected on ground in
Castanopsis-oak forest, at 1400 m. elevation, Arau, New Guinea,
Oct. 12, 1959, by L. J. Brass (no. 32042).

This unusual and attractive species is immediately distinguished by
the tomentose stipes and the branches densely crowded in a compact,
globular head up to 6 cm. in diameter, suggestive in a way of H.
junghuhnit but much denser and more pronounced.

Mniodendron humile Lindb.

Mt. Wilhelm, east slopes: on dead tree in forest 2740 m., 30243; on a log in
forest, 2770 m., 30451. Mt. Otto, south slopes: terrestrial in stunted forest,
3300 m., 31039, Arau: on logs in forest, 1400 m., 31978.

BARTRAM—MOSSES OF NEW GUINEA EASTERN HIGHLANDS 53

Bartramiaceae

*Conostomum australe Sw.
Mt. Wilhelm, east slopes: on moist ground near summit, 14,700 ft., Van
Deusen & Collins 30030.

Distribution: Colombia to Fuegia, Falkland Islands, Auckland
Islands, Campbell Islands, Australia, New Zealand.

Bartramia sp.

The leaves up to 4 mm. long differ from those of either B. cubica
Dix. or B. conica Bartr. where they are 7-8 mm. long. In the absence
of fruit it seems unwise to propose a new species.

Mt. Wilhelm, east slopes: cushioned in crevices near summit, 14,500-14,700
ft., Collins 29963.

Philonotis secunda (D. & M.) Bosch & Lac.

Mt. Wilhelm, east slopes: wet, boggy ground in long-grass community in
alpine grassland, 3560 m., 30070; on rocks in stream, 2770 m., 30450a. Mt.
Otto, south slopes: wet stream bank, 2200 m., 30841. Morobe District, Umi
River, Markham Valley: 480 m., 32529.

Philonotis novoguinensis Reim.?
Mt. Wilhelm, east slopes: in shallow seepage water on open flood terraces of
Pengagl Creek, 2770 m., 30407.

The broader leaf cells (to 10 » wide) suggest that these plants may
be referred to Reimer’s species, but in the absence of fruit the deter-
mination is questionable.

Breutelia romeri Fleisch.
Mt. Wilhelm, east slopes: on moist ground in long-grass community of alpine
grassland, 3560-3850 m., 29819, 30117.

Spiridentaceae
Spiridens reinwardtii Nees.
Mt. Wilhelm, east slopes: on stem of a tree fern in subalpine forest, 3560 m.,
30149.

Spiridens aristifolius Mitt.
Purosa, Okapa area: in rain forest, forming large clumps often on stems of
tree ferns, 1950 m., 31682.

Spiridens perichaetialis Bartr. & Willis.

Morobe District, Wau-Edie Creek road: on dead tree fern, 1850 m., 29777.
Mt. Wilhelm, east slopes: on trees, 2770 m., 30292. Mt. Otto, south slopes: on
small trees in forest, numerous plants radiating from their supports, 2480 m.,
30892. Mt. Michael: on trees in forest, 3100-3290 m., Brass & Collins 31302.
Purosa, Okapa area: on stems of tree ferns, 1950 m., 31682a.

Orthotrichaceae
Zygodon reinwardtii Al. Br.
Mt. Wilhelm, east slopes: on deadwood in subalpine forest, 3560 m., 29820.

Zygodon intermedius B. & 5.
Mt. Wilhelm, east slopes: under a rock overhang in alpine grassland, 3650 m.,
Van Deusen 30152.
768-102—65——3

54. CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Ulota anguste-limbata Bartr.
Mt. Wilhelm, east slopes: on upper branches of trees in subalpine forest,
3650 m., 29867; on dead branch in edge of subalpine forest, 3900 m., 30120.

Ulota splendida Bartr.
Mt. Wilhelm, east slopes: in golden yellow masses on alpine grassland, 13,100
ft., Collins 29944; alpine grassland, 13,100 ft., Collins 29945a.

Desmotheca apiculata (D. & M.) Lindb.

Morobe District, Botanic Gardens, Lae: 200 ft., on roots of epiphytic orchid,
29108; Oomsis: matted high on a dipterocarp tree in forest, 150 m., 29194; Umi
River, Markham Valley, 480 m., on tree trunk, 32617.

Macromitrium erubescens Bartr.
Morobe District, Kaindi: high epiphyte in Nothofagus forest, 2150 m., 29701,
Mt. Otto, south slopes: high epiphyte in Castanopsis forest, 2000 m., 31116.

Macromitrium orthostichum Nees.
Mt. Otto, south slopes: on tree, 2000 m., 31117a.

Macromitrium (Goniostoma) brevirameum Bartr. sp. nov.

Humile, fuscescens. Caulis prostratus, ramosus, rami breves,
4-5 mm. alti, cuspidati. Folia sicca spiraliter contorta, humida
patula, anguste oblongo-lanceolata, obtusa, apiculata, 1.5 mm.
longa, 0.3 mm. lata; marginibus integris, superne sinuatis; costa
percurrente; cellulis superioribus parvis, papillosis, sat incrassatis,
basilaribus linearibus, grosse tuberculosis. Seta gracilis, 5-7 mm.
longa; theca elliptica, microstoma, ore angulato; calyptra glabra,
basi integra.

Type in the U.S. National Herbarium, collected as a high epiphyte
in Castanopsis forest, at 2000 m. elevation, south slopes, Mt. Otto,
New Guinea, Aug. 17, 1959, by L. J. Brass (no. 31117). Paratype
on a living branch in forest undergrowth, at 2000 m. elevation, north-
east slopes, Mt. Michael, Brass 31485.

This species has much in common with M. speirophyllum Bartr.,
but the distinctions are sharply defined. Here the branches are less
than 5 mm. high, the leaves only 1.5 mm. long, the upper leaf cells
7-8 » in diameter and minutely papillose and the color brownish
green, while in M. spetrophyllum the branches are about 1 cm. long,
the leaves 2-2.5 mm. long, the upper cells 12-15 w in diameter and
highly papillose and the color deep, rich red.

Macromitrium novo-guinense Bartr.

Mt. Wilhelm, east slopes: on bark of a dead tree in forest, 2750 m., 30229;
on tree trunks in subalpine forest, 3560 m., 29823.

Macromitrium sublongicaule Bartr.

Mt. Wilhelm, east slopes: on high branches of a tree in forest, 2770 m., 30353.
Mt. Otto, south slopes: high epiphyte in Castanopsis forest, 2000 m., 31118,
31117b. Arau: on tree trunks in Castanopsis forest, 1400 m., 31943, 31977. Mo-
robe District, Umi River, Markham Valley: on tree trunk, 480 m., 32615.

BARTRAM—MOSSES OF NEW GUINEA EASTERN HIGHLANDS’ 55

Macromitrium semipellucidum D, & M.

Morobe District: Kelly Bros. place about 3 miles west of Lae, on Bulolo road,
20 m., 29102; Botanic Gardens, Lae, 200 ft., on tree-fern stem support of an
epiphytic orchid, 29112; Umi River, Markham Valley, on a dead stick, 480 m.,
32548; on a tree trunk, 32614.

Macromitrium angustifolium D. & M.

Mt. Wilhelm, east slopes: on dead branch, 2740 m., 30230; on tree trunk in
forest edge, 2600 m., 30789. Mt. Otto, south slopes: on tree trunk in Castanop-
sis forest, 2000 m., 31113. Kassam: on tree trunk, 1370 m., 32376. Purosa,
Okapa area: on a big tree in rain forest, 1950 m., 31796. Morobe District,
Kaindi: open, rocky bank of stream in forest, 2060 m., 29549; Umi River, Mark-
ham Valley, tree trunk, 480 m., 32616.

Schlotheimia emarginato-pilosa Herz.
Arau: high on trees in Castanopsis-oak forest, 1400 m., 31941.

Schlotheimia pilicalyx Broth. & Geh.

Mt. Wilhelm, east slopes: tree trunks in subalpine forest, 3650 m., 29873,
29993. Mt. Otto, south slopes: on alpine grassland of summit, 3540 m., Brass &
Collins 31038a.

Schlotheimia macgregorii Broth. & Geh.

Mt. Michael: on a tree in forest, 2700 m., 31450. Morobe District, Kaindi:

high epiphyte in Nothofagus forest, 2150 m., 29702.

Rhacopilaceae
Powellia involuta Mitt.

Mt. Michael, northeast slopes: on stem of small Pandanus in Castanopsis-
Nothofagus forest, 2000 m., 31547. Arau: trunk of Pandanus in Castanopsis-
oak forest, 1400 m., 31942; on stem of Pandanus tree, 1400 m., 32039.

Rhacopilum spectabile R. & H.

Purosa, Okapa area: on rotting logs, 1950 m., 31651. Kassam Gap: on log,
1460 m., 32297. Mt. Michacl, northeast slopes: on twigs and branches, 2000 m.,
31540, 31542, 31549. Mt. Otto, south slopes: on tree in forest, 2200 m., 30840.
Morobe District, Wau-Edie Creek road: on log, 5,800 ft., 29163; Mt. Kaindi:
on log, 2200 m., 29594 in part.

Hedwigiaceae

Rhacocarpus humboldtii (Hook.) Lindb.

Mt. Wilhelm, east slopes: cushioned on moist ground near summit, 14,700 ft.,
Van Deusen & Collins 30031.
Rhacocarpus alpinus (Wright) Par.

Mt. Wilhelm, east slopes: terrestrial on alpine grassland, 3700 m., 29995.

These plants correspond in all particulars with those from Mt.
Kinabalu, Borneo, and also with the scrap from Mt. Wilhelmina
(Bartram, E. B., Lloydia 5:275, 1942). The leaves show no trace of
a hair-point and are constantly short apiculate. By contrast the
plants in no. 30031 all show hair-pointed leaves. I have therefore
referred this latter collection to R. humboldtii and this one to R.
alpinus pending a critical revision of the genus.

56 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Cyrtopodaceae

Bescherellea cyrtopus F. v. Muell.
On lower trunks of trees in Castanopsis forest, Kassam, 1370 m., 32281.

*Bescherellea elegantissima Duby.

Mt. Wilhelm, east slopes: low on trees in moist forest, 2740 m., 30245; on tree
trunks in forest, 2900 m., 30387; low on trees in open, swampy forest, 2770 m.,
30468.

Distribution: New Caledonia, Aneityum.
Ptychomniaceae

Hampeella pallens (Lac.) Ficisch.
Morobe District, Kaindi: on dead branch in forest, 2060 m., 29663.

Prionodontaceae

Neolindbergia brassii Bartr.

Mt. Wilhelm, east slopes: masses low on tree in forest, 2770 m., 30620.

By good fortune these plants are well fruited with the capsules
very young and immature, ripe and in prime condition, or old and
empty. The sporophyte characters are as follows: perichaetium
small, inner leaves erect, 3 mm. long, ligulate from an ovate base,
serrulate near apex, costa ending below the acute apex, upper cells
small, round, basal cells linear toward costa, small and round in
5-6 rows at margins. Seta 5 mm. long, erect, papillose; capsule
globose, brown, glossy, 1 mm. in diam.; operculum oblique, conic-
rostrate, 1 mm. long; calyptra cucullate, short, pilose, 2 mm. long;
peristome teeth about 0.3 mm. long, linear, hyaline, smooth, inserted
below rim; endostome none or fragmentary and rudimentary without
segments or cilia; spores 16-20 yu in diam., minutely papillose, exothecial
cells irregularly hexagonal, incrassate, smaller and colored in several
rows around mouth, annulus none.

Pterobryaceae
Trachyloma indicum Mitt.
Mt. Wilhelm, east slopes: low on trees in subalpine forest, 3560 m., 30069.

Trachyloma tahitense Besch.

Mt. Wilhelm, east slopes: low on trees in moist, mossy forest, 2900 m., 30390.
Endotrichella apiculata Bartr. sp. nov.

Sat robusta, pallide viridis, nitida. Caulis secundarius usque ad
4 cm. longus, simplex, haud complanatus, cum foliis 7 mm. latus.
Folia late patentia, rotundato-ovata, ca. 3 mm. longa, 2.2 mm. lata,
apice rotundata, apiculata, plicata; marginibus superne minute
denticulatis; costa duplex, brevis; cellulis lineari-rhomboideis, in-
crassatis, infimis fuscis, alaribus nullis. Folia perichaetialia parva,
erecta, acuta; seta 1 mm. longa; theca erecta, exserta, oblonga deoper-
culata 2 mm. longa.

BARTRAM—MOSSES OF NEW GUINEA EASTERN HIGHLANDS 57

Type in the U.S. National Herbarium, collected on undergrowth
in forest, at 1400 m. elevation, Arau, New Guinea, Oct. 6, 1959, by
L. J. Brass (no. 31909).

Paratype: On undergrowth in Castanopsis-oak forest, 1370 m.,
Kassam, New Guinea, Brass 32439a.

The short, apiculate leaves are distinctive as compared with the
other local species.

Endotrichella compressa (Mitt.) Broth.

Arau: on undergrowth in forest, 1400 m., 31910; high on trees in Castanopsis-
oak forest, 1400 m., 31925. Kassam: tufted on underbrush, 1370 m., 32437.
Endotrichella falcifolia Bartr. sp. nov.

Robusta, superne pallide viridis inferne fuscescens. Caulis secun-
darius ad 5-6 cm. longus. Folia vix conferta, faleato-secunda, oblongo-
lanceolata, filiformiter acuminata, plures plicata, ecostata, 6-7 mm.
longa, 1.5 mm. lata; marginibus superne grosse serratis; cellulis
superioribus anguste linearibus, incrassatis, basilaribus latioribus,
infimis fuscis, alaribus paucis, irregularis. Caetera ignota.

Type in the U.S. National Herbarium, collected on undergrowth
in rain forest, at 2000 m. elevation, Purosa, Okapa area, New Guinea,
Oct. 1, 1959, by L. J. Brass (no. 31840).

The falcate-secund leaves will separate this species from E. perpli-
cata Broth. to which it seems to have some affinity.

Endotrichella elegans (D. & M.) Fleisch.

Morobe District, Gurakor: on branchlets of small tree in a moist ravine in

rain forest, 640 m., 29419.

Endotrichella perplicata Broth.
Purosa, Okapa area: on branches of woody undergrowth in rain forest, 1900
m., 31766a. Kassam: on undergrowth in Castanopsis-oak forest, 1370 m., 32436.

Endotrichella novae-hannoverae C. M.
Mt. Wilhelm, east slopes: low on tree in moist forest, 2770 m., 30284.

Garovaglia mirabilis Dix.

Kassam: on underbrush in Castanopsis-oak forest, 1370 m., 32438. Mt. Michael,
northeast slopes: on a rock in Castanopsis-Nothofagus forest, 2000 m., 31546.
Morobe District, Mt. Kaindi: on dead branchlets in forest undergrowth, 7,200
ft., 29166.

Garovaglia brevifolia Bartr. sp. nov.

Gracilis, fuscescens, nitida. Caulis secundarius usque ad 5 em.
longus, simplex, sat dense foliosus, cum foliis ca. 2 mm. latus. Folia
late patentia, apice incurva, late ovata, abrupte apiculata, 2-2.5
mm. longa, 1.3-1.6 mm. lata, leniter plicata, ecostata; marginibus
planis, superne denticulatis; cellulis anguste rhomboideis, c. 12 u
latis, 36-48 y longis, incrassatis, alaribus sat numerosis, oblongis.
Perichaetium majusculum, alte convolutum, foliis abrupte breviter
acuminatis, marginibus superne denticulatis; theca immersa, oblongo-
cylindrica, deoperculata 2 mm. longa, subsessilis.

58 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Type in the U.S. National Herbarium, collected on undergrowth
in a semiswampy place in Castanopsis-oak forest, at 1370 m. elevation,
Kassam, New Guinea, Nov. 5, 1959, by L. J. Brass (no. 32439).

The short, abruptly apiculate leaves and more slender habit are
thoroughly distinctive characters in comparison with G. densifolia
Thw. & Mitt.

Garovaglia papuana Dix.
Mt. Wilhelm, east slopes: low on trees in moist forest, 2740 m., 30246.

Meteoriaceae

Meteorium miquelianum (C. M.) Fleisch.

Mt. Wilhelm, east slopes: on twigs in subalpine forest, 3560 m., 29846; on
woody undergrowth in moist forest, 2770 m., 30368. Morobe District, Umi
River, Markham Valley: on tree in rain forest, 480 m., 32585.

Floribundaria floribunda (D. & M.) Fleisch.

Arau: on leaf of a Zingiberaceous herb, 1400 m., 32185. Kassam: on woody
undergrowth, 1370 m., 32279. Morobe District, Umi River, Markham Valley:
on trees, underbrush and leaves in rain forest, 480 m., 32550, 32551.
Floribundaria aurea (Griff.) Broth.

Mt. Wilhelm, east slopes: pendant from underbrush in moist, mossy forest,
2770 m., 30409.

Meteoriopsis reclinata (C. M.) Fleisch.

Morobe District, Gurakor: on leaves, rotten wood and branchlets, 640 m.,
29417, 29420a, 29423.

Lembophyllaceae

Camptochaete subporotrichoides (Broth. & Geh.) Broth.
Mt. Wilhelm, east slopes: low on trees in forest, 2900 m., 30391.

Phyllogoniaceae

Orthorrhynchium elegans (H. f. & W.) Reh’dt.
Kassam: on lower trunk of tree in Castanopsis-oak forest, 1370 m., 32377.

Neckeraceae

Calyptothecium crispulum (Lac.) Broth.
Mt. Otto, south slopes: on a tree on forest edge, 7,300 ft., 29124.

Calyptothecium extensum Fleisch.
Morobe District, Umi River, Markham Valley: 480 m., 32587.

Calyptothecium urvilleanum (C.M.) Broth.
Kassam: on stilt roots of Pandanus, 1370 m., 32413. Morobe District: Umi
River, Markham Valley, 480 m., on trees in rain forest, 32586.

Neckeropsis gracilenta (Lac.) Fleisch.
Morobe District, Gurakor: on leaves and trees in moist rain forest, 640 m.,
29420, in part, 29421.

Neckeropsis lepineana (Mont.) Fleisch.

Purosa, Okapa area: on tree trunk, 1900 m., 31767. Arau: on tree trunks,
1400 m., 31907. Kassam: on tree trunks, 1370 m., 32334. Morobe District:
Umi River, Markham Valley, on underbrush in rain forest, 480 m., 32549.

BARTRAM—MOSSES OF NEW GUINEA EASTERN HIGHLANDS 959

Homaliodendron fliabellatum (Sm.) Fleisch.
Kassam: on tree trunks, 1370 m., 32280.

Pinnatella mucronata (Lac.) Fleisch.
Morobe District, Umi River, Markham Valley: on dead branches, 480 m.,
32531a.
Nemataceae

Ephemeropsis tjibodensis Goeb.
Mt. Michael, northeast slopes: on twigs and leaves of an undergrowth small
tree in Castanopsis-Nothofagus forest, 2000 m., 31531 in mixture.

Hookeriaceae

Daltonia angustifolia D. & M.
Mt. Wilhelm, east slopes: on deadwood in subalpine forest, 3560 m., 30044;

on twigs in forest undergrowth, 2740-2800 m., 30248, 30472, 30740.

Daltonia angustifolia D. & M. var revoluta (Broth.) Bartr.
Mt. Wilhelm, east slopes: on a log in moist forest, 2740 m., 30242.

Daltonia contorta C. M.
Mt. Wilhelm, east slopes: on a log, 2740 m., 30242a.

Daltonia aristifolia R. &. C.
Mt. Wilhelm, east slopes: on deadwood in subalpine forest, 3650 m., 29875.

Distichophyllum brevicuspidatum Bartr.
Mt. Michael, northeast slopes: on twigs and leaves, 2000 m., 31531a.

Actinodontium rhaphidostegum (C.M.) Bosch & Lac.
Arau: on a rotting log, 1400 m., 31980a.

Callicostella papillata (Mont.) Mitt.
Purosa, Okapa area: on rotting wood in rain forest, 1950 m., 31696.

Chaetomitriopsis glaucocarpa (Reinw.) Fleisch.

Purosa, Okapa area: on branches of an undergrowth tree in rain forest, capsules
a pale, glaucous blue 1950 m., 31797. Kassam: low on woody undergrowth in
forest, 1370 m., 32278.
Chaetomitrium nano-hystrix C. M.

Morobe District, Gurakor: on living twigs in rain forest undergrowth, 640 m.,
29424.
Chaetomitrium papillifolium Bosch & Lac.

Morobe District, Umi River, Markham Valley; on dead branches and bark in
rain forest, 480 m., 32531, 32583.
Chaetomitrium papuanum Bartr.

Arau: on twigs of undergrowth trees, 1400 m., 31929. Kassam: on branches
of woody undergrowth, 1370 m., 32435.
Chaetomitrium crispifolium Bartr.

Mt. Wilhelm, east slopes: on a tree trunk, 3560 m., 29850a; on a log in moist
forest, 2740 m., 30241.

Chaetomitrium integrifolium Bartr.
Arau: on leaf of an undergrowth tree in Castanopsis-oak forest, 1400 m., 31928.

Chaetomitrium pseudopapillifolium Bartr.
Morobe District, Oomsis: on bark and branchlets in rain forest, 100 m., 29243,
29244, 29292, 29293; Umi River, Markham Valley: on a tree trunk, 480 m., 32656.

60 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Thuidiaceae

Pelekium velatum Mitt.

Morobe District, Oomsis: logs in rain forest, 100 m., 29210, 29211; Umi River,
Markham Valley, on a log, 480 m., 32613.
Thuidium furfurosum (I. f. & W.) Jacg.

Morobe District, Oomsis: on rocks on bank of stream in rain forest, 100 m.,
29230.
Thuidium investe (Mitt.) Jaeg.

Morobe District, Kaindi: on bark and rotting wood in forest, 2060 m., 29550,
29555.
Thuidium cymbifolium (D. & M.) Bosch & Lac.

Mt. Michael, northeast slopes: on trees and rocks in Castanopsis-Nothofagus
forest, 31539, 31536. Kassam: on rocks, 1370 m., 32349, 32434, in part.

Amblystegiaceae

*Calliergon sarmentosum (Wahlenb.) Kindb.

Mt. Wilhelm, east slopes: on wet ground in alpine grassland, 13,100 ft., Collins
299438a.

The occurrence of this widely distributed species in the alpine region
of New Guinea is noteworthy and significant. It is evidently a waif
from the south where it occurs sparingly in New Zealand.

Distribution: Europe, Asia, Fuegia, South Georgia, New Zealand,
Kast Africa.

Calliergon wilhelmense Bartr. sp. nov.

Dense caespitosum, caespitibus fuscescentibus, nitidis. Caules
prostrati, parce ramosi, valde tumidi. Folia laxe imbricata, oblongo-
ovata, breviter acuminata, profunde concaya, usque ad 2 mm. longa,
0.8 mm. lata; marginibus crectis, integris superne inflexis; costa duplex,
brevissima; cellulis superioribus, anguste linearibus, 4-5 u latis, 40-50
# longis, basilaribus longioribus, incrassatis alaribus sat numerosis,
irregulariter quadratis, incrassatis. Caetera ignota.

Mt. Wilhelm, east slopes: forming loose cushions on wet ground on alpine
grassland, 13,100 ft., J. D. Collins 29943. June 15, 1959.

This species in appearance and detail shows a very close affinity to
C. turgescens (Th. Jens.) Kindb., but the uniformly short, double
costa and the narrower lamina cells seem to be distinguishing
characters.

Brachytheciaceae

Pleuropus luzonensis Broth.

Mt. Wilhelm, east slopes: low on a dead tree in forest, 2740 m., 30240; on bark
of a big Podocarpus papuanus tree, 2770 m., 30469.
Brachythecium plumosum (Hedw.) B. S. & G

Mt. Wilhelm, east slopes: on rocks of a stream in forest, 2600-2800 m., 30447,
30449, 30600, 30738, 30739.

BARTRAM—MOSSES OF NEW GUINEA EASTERN HIGHLANDS 61

Brachythecium integerrimum Bartr.
Mt. Wilhelm, east slopes: on rocks in an alpine grassland stream, 3060 m.,

30090.
Eurhynchium celebicum (Bosch & Lac.) Bartr.
Arau: on a bracket fungus in Castanopsis-oak forest, 1400 m., 32194 in mixture.

Rhynchostegiella stellata Bartr.
Morobe District, Kaindi: on a tree in forest, 2060 m., 29554, with hepatics.

Entodontaceae

Erythrodontium julaceum (Hook.) Par.
Morobe District. Wau: in front of District Office on a tree, 3,500 ft., 29156;
Umi River, Markham Valley, on a tree in rain forest, 480 m., 32657.

Campylodeontium flavescens (Hook.) Bosch & Lac.
Arau: on rotting wood in Castanopsis-oak forest, 1400 m., 32062.

Sematophyllaceae

Clastobryella cuculligera (Lac.) Fleisch.
Mt. Wilhelm, east slopes: on branch in forest undergrowth, 2770 m., 30471.

Acanthocladium clarkii Dix.
Mt. Wilhelm, east slopes: pendant from twigs in subalpine forest, 3700 m.,
29990.

Acanthocladium gracile Bartr.
Mt. Wilhelm, east slopes: on a tree trunk in subalpine forest, 3560 m., 29850.

Morobe District, Kaindi: on moist, clayey roadbanks, 2060 m., 29615.

The falcate-secund leaves suggest that this species may be an
Ectropothecium. The inflated alar cells of the stem leaves are hard
to demonstrate and seem to be large hyaline cells confined to the
extreme basal angles as in other species of Ectropothecium.

Trismegistia panduriformis (C. H. Wright) Broth.
Purosa, Okapa area: on surface roots in rain forest, 1950 m., 31666.

Trismegistia rigida (II. & R.) Broth.
Arau: on a log in Castanopsis-oak forest, 1400 m., 32196.

*Meiothecium attenuatum Broth.
Morobe District, Wau: on a tree trunk in front of District Office, 3,500 ft.,

29157.
Distribution: Philippine Islands.

Meiothecium longisetum Bartr.
Mt. Wilhelm, east slopes: on deadwood in moist, mossy forest, 2900 m., 30389,

Meiothecium microcarpum (Harv.) Mitt.
Morobe District, Botanic Gardens, Lae: 200 ft., on bark of frangipani tree,
29115; Lae: 10 m., on bark of garden Hibiscus tree, 29116.

Brotherella falcata (D. & M.) Fleisch.
Arau: on a rotting log in Castanopsis-oak forest, 1400 m., 31979.

Sematophyllum subhumile (C. M.) Fleisch.
Morobe District, Oomsis: tree trunk in rain forest, 200 m., 29265.

62 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Acroporium diminutum (Brid.) Fleisch.
Kassam: on tree trunks, 1370 m., 32432.

Acroporium sigmatodontium (D. & M.) Fleisch.
Arau: on a tree trunk, 1400 m., 31960.

Acroporium secundum (R. & H.) Fleisch.
Purosa, Okapa area: on trees, 1950 m., 31734.

Warburgiella subleptorrhynchoides Fleisch.

Mt. Wilhelm, east slopes: log in forest, 2740 m., 30239, 30249, 30250; on trees
in subalpine forest, 3560-3650 m., 29848, 29871, 29893. Mt. Otto: on trees and
rotting wood, 2000 m., 31114, 31115. Mt. Michael, northeast slopes: on rotting
wood and a bracket fungus, 2000 m., 31550, 31391. Purosa, Okapa area: on
rotting logs, 1950 m., 31647, 31649. Arau: on rotting wood, 1400 m., 31940a.
Kassam: on a rotting log, 1370 m., 32336. Morobe District, Mt. Kaindi: on
deadwood, 2060-2200 m., 29642, 29764.

Trichosteleum laevi-hamatum Dix.
Arau: on rotting wood, 1400 m., 32197.

Trichosteleum novae-guineae Bartr.
Arau: on rotting wood, 1400 m., 31940.

Acanthorrhynchium papillatum (Harv.) Fleisch.
Kassam: on tree trunks, 1370 m., 32337.

Taxithelium nepalense (Schwacgr.) Broth.
Morobe District, Oomsis: on deadwood in rain forest, 150 m., 29193, 29212.

Taxithelium kerrianum (Broth.) Fleisch.

Morobe District, Umi River, Markham Valley: on dead branches, 480 m.,
32531b.
Taxithelium isocladum (Lac.) R. & C.

Arau: on leaves of a ground fern, 1400 m., 31975. Morobe District, Gurakor:
on leaves of undergrowth shrub, 640 m., 29481.

Glossadelphus zollingeri (C. M.) Fleisch.
Morobe District, Gurakor: on a rock in a stream bed in rain forest, 640 m.,
294138.

Glossadelphus prostratus (D. & M.) Fleisch.
Mt. Wilhelm, east slopes: on moist granitic rocks of a stream in forest, 2600 m.,

30601.

These plants are less slender than the plants from Mt. Dayman
with leaves up to 1 mm. long and 0.45 mm. wide. It is apparently a
robust form.

Macrohymenium mitratum (D. & M.) Fleisch.

Arau: ona tree, 1400 m., 31976a.

Macrohymenium novo-guineense Reim.

Mt. Wilhelm, east slopes: on deadwood in edge of subalpine forest, 3700 m.,
29994; on exposed trunk of tree in forest, 3150 m., 30705; exposed trunk of tree in
subalpine forest, 3600 m., 30148.

These collections differ from Af, mitratum in the more robust habit,
ruddy color, and longer setae. Here as in the material from Mt.

BARTRAM——-MOSSES OF NEW GUINEA EASTERN HIGHLANDS 63

Wilhelmina and Mt. Hagen the setae are from 12-17 mm. long and
weakly scabrous above. The leaves vary from 2-2.4 mm. long and
from 0.4-0.7 mm. wide. While the leaves are slightly larger than
originally described, the robust habit, coloring, and long setae indicate
that they should be referred here.

Hlypnaceae

Ectropothecium ichnotocladum (C. M.) Jaeg.

Mt. Michael, northeast slopes: on dead bark, 2000 m., 31548. Arau: on
deadwood, 1400 m., 31980, 32195. Morobe District, Umi River, Markham
Valley: on log, 32530, in part.

Ectropothecium percomplanatum Broth.
Mt. Michael, northeast slopes: on wet rocks in stream, 2000 m., 31534.

Ectropothecium falciforme (D. & M.) Jaeg.

Mt. Wilhelm, east slopes: among tall grasses in alpine grassland, 3560 m.,
29822; on a rotting log, 2740 m., 30235. Mt. Michael, northeast slopes: on logs,
2000 m., 31486, 31545.

Ectropothecium circinatulum Ther.
Morobe District, Kaindi: on bank of stream, 2060 m., 29557.

Ectropothecium arfakense Broth. & Geh.

Purosa, Okapa area: on a log, 1950 m., 31650. Arau: on a log, 1400 m.,
31927. Morobe District: Umi River, Markham Valley: on a log, 480 m., 32720.

Ectropothecium longipedunculatum Dix.
Purosa, Okapa area: on leaf-littered ground, 2000 m., 31839.

Ectropothecium buitenzorgii (Bel.) Jaeg.

Mt. Michael, northeast slopes: on a log, 31348. Kassam: on logs and rocks
in a stream bed, 1370 m., 32335, 32434, in part. Arau: ona log, 1400 m., 31927a.
Morobe District, Mt. Kaindi: on log, 2200 m., 29594, in part.

Ectropothecium dealbatum (H. & R.) Jaeg.
Morobe District, Botanic Gardens, Lae: on orchid roots, 200 ft., 29113.

Trachythecium verrucosum (Hpe.) Fleisch.

Purosa, Okapa area: on a branch in rain forest undergrowth, 1950 m., 31648;
on rotting wood, 1950 m., 31798. Arau: on a log, 1400 m., 31908. Morobe
District, Umi River, Markham Valley: on logs and rocks, 480 m., 32530, in part,
32682, in part.

Isopterygium albescens (Schwaegr.) Jaegr.
Morobe District, Botanic Gardens, Lae: on a tree-fern stem, 200 ft., 29106.

Ctenidium luzonense Broth.
Mt. Wilhelm, east slopes: on leaves of an epiphytic fern in subalpine forest,
3700 m., 29992; on a tree trunk, 2740 m., 30236.

Hylocomiaceae

Macrothamnium hylocomioides Fleisch.
Mt. Wilhelm, east slopes: on lower trunks of trees in moist forest, 2850 m.,

30556.

64 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Polytrichaceae

Pogonatum junghuhnianum (D. & M.) Bosch & Lac.
Mt. Otto, south slopes: on clayey roadbanks, 7,300 ft., Kotuni, 29123.

Pogonatum humile Bartr.
Mt. Wilhelm, east slopes: in a crevice near summit, 14,500-14,700 ft., Collins

29967; on ground near summit, 14,700 ft., Collins 29964a.

Pogonatum klossii Dix.
Mt. Otto, south slopes: on claybanks, 29122a. Morobe District, Kaindi: on

clayey ground, 2060 m., 29578.

Pogonatum spurio-cirratum Broth.
Purosa, Okapa area: in small colonies in rain forest, 1950 m., 31736. Mt.
Klandora, Kratke Mts.: terrestrial in forest, 2130 m., Brass & Collins 32166.

Polytrichum juniperinum Hedw.
Mt. Michael: alpine grasslands, 3650 m., Brass & Collins 31260, 31432.

Polytrichadelphus archboldii Bartr.
Mt. Wilhelm, east slopes: on open eroding banks of Pengagl Creek, 2770 m.,
30408.
Dawsoniaceae

Dawsonia grandis Schleip. & Geh.
Mt. Wilhelm, east slopes: a small colony in moist, mossy forest, 2600 m., 30487.

Purosa, Okapa area: in rain forest, 1950 m., 31719.

Dawsonia pullei Fleisch.
Mt. Michael: on a path in low, mossy, myrtaceous forest, 3290 m., 31419.
Morobe District, Kaindi: on mossy ground in Nothofagus forest, 2150 m., 29633.

Dawsonia lativaginata v. d. Wijk.
Mt. Otto, south slopes: on rocks and ground in rain forest of a ravine, leaves

somewhat bluish green, 2200 m., 30828.
Dawsonia beccarii Broth. & Geh.

Morobe District, Wau-Edie Creek road: on open, clayey ground, 5,800 ft.,
29164; Kaindi: on open, clayey banks of a stream, 2060 m., 29625.

INDEX

(Synonyms in ztalics; page numbers of principal entries in boldface)

Acanthocladium clarkii, 61
gracile, 61
Acanthorrhynchium papillatum, 62
Acroporium diminutum, 62
secundum 62
sigmatodontium 62
Actinodontiuvm rhaphidostegum, 59
Amphidium cyathicarpum, 46
Andreaea novo-guinensis, 44
rupestris, 44
Anomobryum cymbifolium, 50
Atractylocarpus dicranoides, 46
Bartramia conica, 53
cubica, 53
sp., 53
Bescherellea cyrtopus, 56
elegantissima, 56
Brachymenium nepalense, 50
Brachythecium integerrimum, 61
plumosum, 60
Braunfelsia enervis, 47
Breutelia romeri, 53
Brotherella falcata, 61
Bryum argenteum, 50
australe, 50
crassulum, 51
junghuhbnianum, 51
nitens, 50
novo-guinense, 51
porphyroneuron, 50
truncorum, 51
wilhelmense, 50
Callicostella papillata, 59
Calliergon sarmentosum, 43, 60
turgescens, 44, 60
wilhelmense, 43, 60
Calymperes hyophylaceum, 48
moluccense, 48
tahitense, 48
Calyptothecium crispulum, 58
extensum, 58
urvilleanum, 58
Camptochaete subporotrichoides, 58
Campylodontium flavescens, 61
Campylopodium euphorocladum, 45

Campylopus archboldii, 46
atro-fuscescens, 46
caudatus, 45
clemensiaec, 46
comosus, 45
crispifolius, 45
exasperatus, 46
morobensis, 45
robbinsii, 45
umbellatus, 46
wilhelmensis, 45

Chaetomitriopsis glaucocarpa, 59

Chaetomitrium crispifolium, 59
integrifolium, 59
nano-hystrix, 59
papillifolium, 59
papuanum, 59
pseudopapillifolium, 59

Clastobryella cuculligera, 61

Conostomum australe, 53

Cryptodicranum armittii, 46

Ctenidium luzonense, 63

Daltonia angustifolia, 59
angustifolia var. revoluta, 59
aristifolia, 59
contorta, 59

Dawsonia beccarii, 64
grandis, 64
lativaginata, 64
pullei, 64

Desmotheca apiculata, 54

Dicranella setifera, 45

Dicranoloma blumii, 47
braunii, 47
dicarpum 47
laevifolium, 47
perarmatum, 47

Dicranoweisia macrocarpa, 46

Distichophyllum brevicuspidatum, 59

Ditrichum sericeum, 44

Ectropothecium, 61
arfakense, 63
buitenzorgii, 63
circinatulum, 63
dealbatum, 63

65

66

Ketropothecium—Continued
falciforme, 63
ichnotocladum, 63
longipedunculatum, 63
percomplanatum, 63

Endotrichella apiculata, 56
compressa, 57
elegans, 57
falcifolia, 57
novae-hannoverae, 57
perplicata, 57

Iintosthodon contortus, 49
subulatus, 49

Ephemeropsis tjibodensis, 59

Erythrodontium julaceum, 61

Eurhynchium celebicum, 61

Fissidens filicinus, 44

Floribundaria aurea, 58
floribunda, 58

Fumaria hygrometrica, 49

Garovaglia brevifolia, 57
densifolia, 58
mirabilis, 57
papuana, 58

Glossadelphus prostratus, 62
zollingeri, 62

Grimmia affinis, 49
ovalis, 49

Hampeella pallens, 56

Holomitrium austro-alpinum, 46
stenobasis, 46

Nomaliodendron flabellatum, 59

Hymenodon parvulus, 51

Hyophila involuta, 48

Hypnodendron auricomum, 52
brassii, 52
diversifolium, 52
junghuhnii, 52

Isopterygium albescens, 63

Leptodontium warnstorfii, 48

Leptostomum intermedium, 51

Leucobryum candidum, 47
chlorophyllosum, 47
pentastichum, 47
sanctum, 47

Leucophanes glaucescens, 47
octoblepharioides var. korthelsii, 47

Macrohymenium mitratum, 62
novo-guineense, 62

Macromitrium angustifolium, 55
brevirameum, 54
erubescens, 54
novo-guinense, 54

INDEX

Macromitrium—Continued
orthostichum 54
semipellucidum, 55
speirophyllum, 54
sublongicaule, 54

Macrothamnium hylocomioides, 63

Meesea triquetra, 44

Meiothecium attenuatum, 61
longisetum, 61
microcarpum, 61

Meteoriopsis reclinata, 58

Meteorium miquelianum, 58

Miclichhoferia javanica, 50
novo-guinensis, 49

Mniodendron humile, 52

Mnium rotundifolium, 51

Neckeropsis gracilenta, 58
lepineana, 58

Neolindbergia brassii, 56

Octoblepharum albidum, 47

Orthomniopsis elimbata, 51

Orthorrhynchium elegans, 58

Pelekium velatum, 60

Philonotis novoguinensis, 53
secunda, 53

Pinnatella mucronata, 59

Pleuropus luzonensis, 60

Pogonatum humile, 64
junghuhnianum, 64
klossii, 64
spurio-cirratum, 64

Pohlia elongata, 50
flexuosa, 50

Polytrichadelphus archboldii, 64

Polytrichum juniperinum, 64

Powellia involuta, 55

Rhacocarpus alpinus, 55
humboldtii, 55

Rhacomitrium crispulum, 49
lanuginosum var. pruinosum, 49

Rhacopilum spectabile, 55

Rhizogonium spiniforme, 52

Rhodobryum giganteum, 51

Rhynchostegiella stellata, 61

Schlotheimia emarginato-pilosa, 55
macgregorii, 55
pilicalyx, 55

Sematophyllum subhumile, 61

Spiridens aristifolius, 53
perichaetialis, 53
reinwardtii, 53

Streptopogon erythrodontus, 48
erythrodontus var. rutenbergii, 48

Symblepharis reinwardtii, 46

Syrrhopodon ciliatus, 48
philippinensis, 48

Taxithelium isocladum 62
kerrianum, 62
nepalense, 62

Tayloria octoblepharis, 49

Thuidium cymbifolium, 60
furfurosum, 60
investe, 60

Trachyloma indicum, 56
tahitense, 56

Trachythecium verrucosum, 63

INDEX

67

Trematodon longicollis, 45

Trichosteleum laevi-hamatum, 62
novae-guineae, 62

Trichostomum angustatum, 48
subulifolium, 48

Trismegistia panduriformis, 61
rigida, 61

Ulota anguste-limbata, 54
splendida, 54

Warburgiella subleptorrhynchoides, 62

Zygodon intermedius, 53
reinwardtii, 53

U.S. GOVERNMENT PRINTING OFFICE :1965

BULLETIN OF THE UNITED STATES NATIONAL MUSEUM

CONTRIBUTIONS FROM THE UNITED States NATIONAL HERBARIUM

VoLuME 37, Part 3

STUDIES OF PACIFIC ISLAND PLANTS, XVIII
NEW AND NOTEWORTHY
FLOWERING PLANTS FROM FIJI

By ALBERT C. SMITH

SMITHSONIAN PRESS e WASHINGTON, D.C. e 1967

STUDIES OF PACIFIC ISLAND PLANTS, XVIII!
NEW AND NOTEWORTHY
FLOWERING PLANTS FROM FIJI

ALBERT C. SMITH

Most of the plants discussed or described in this paper were obtained
during the writer’s third collecting trip to Fiji in 1953-54.2_ The first
set of this material is deposited in the U.S. National Herbarium; a
duplicate study set is in the herbarium of the Bernice P. Bishop Mu-
seum; and eight remaining sets will shortly be distributed to world
herbaria. However, earlier collections in herbaria have also been
consulted and are here cited when pertinent. In this paper 18 species
are described as new, and range extensions and emendations are noted
for a few others. The place of deposit of specimens is indicated as
follows: Arnold Arboretum (A); Bernice P. Bishop Museum (Bish);
Gray Herbarium (GH); University of California, Berkeley (UC);
U.S. National Herbarium (US). I am indebted to the staffs of these
institutions for the privilege of studying such material, and especially
to the Director of the Bishop Museum for providing permanent
working space.

Musaceae

Heliconia paka A. C. Sm., sp. nov.
Heliconia bihai sensu A. C. Sm. Sargentia 1:7. 1942; sensu J. W. Parham,
Pl. Fiji Isl. 259. 1964; non L.

Herba valida 2-6 m. alta; foliis saepe 3 m. longis vel longioribus
glabris, petiolis crassis ad 1 m. longis, laminis lanceolatis 2-3 m. longis
medium versus 30-50 cm. latis, basi rotundatis et in petiolum decurren-
tibus, apice obtusis, costa valida, nervis primariis inter se 0.8-1.5 cm.
marginem versus abrupte curvatis; inflorescentia erecta ad 60-70 cm.
longa, pedunculo et rhachide crassis 10-15 mm. diametro glabris,

1 This paper is based on research partially supported by a grant from the Na-
tional Science Foundation. Number XVII, by A. C. Smith and B. C. Stone, was
published in Contr. U.S. Nat. Herb. 37:1-41, 3 pls. 1962.

2Smith, A. C., Botanical studies in Fiji. Ann. Rep. Smithsonian Inst. for
1954: 305-315, 12 pls. 1955. For itineraries of the two earlier trips, see Journ.
New York Bot. Gard. 35:261-280. 1934; Journ. Arnold Arb. 31:188-141. 1950.

69

70 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

pedunculo 5-10 em. longo, rhachide inconspicue flexuosa; bracteis
primariis inter se 2-6 cm. plerumque 12-15, vagina subcoriacea saepe
lenticellata marginem versus pilis brunneis 0.3-0.7 mm. longis incon-
spicue hispidula, lamina subcarnosa late ovato-lanceolata, 30 (in-
feriore)—13 (superiore) cm. longa, ad 5 cm. lata, ad apicem obtusum
gradatim angustata, crasse carinata, margine scariosa, conspicue ner-
vata, praeter margines versus proxime ut vagina pilosa ubique glabra:
floribus glabris plerumque 5-8 in axillis bractearum; bracteis sub
floribus e basi lata lanceolatis 6-7 em. longis 1.5-2 em. latis acutis mul-
tinerviis inconspicue carinatis, praeter pilos rigidos brunneos adpressos
circiter 1 mm. longos ad carinam glabris; pedicellis teretibus 2-4 mm.
diametro 15-25 mm. longis sub anthesi et fructu superne paullo in-
crassatis; tepalis liberis in vivo subcarnosis in sicco chartaceis con-
spicue nervatis 55-60 mm. longis, posteriore 7-9 mm. anterioribus
4—5 min. interioribus 5-6 mm. latis; staminodio petaloideo tepalum
posterius basim versus affixo, parte libera subcarnosa ovato-cucullata
acuta ca. 5X5 mm.; staminibus et stylo sub anthesi quam tepalis
paullo brevioribus, antheris 12-13 mm. longis obtusis, stigmate
3-lobato lobis obscure bifidis; ovario elongato-turbinato sub anthesi
10-13 mm. longo et 4-5 mm. diametro; fructu in vivo carnoso in
sicco coriaceo ellipsoideo-triquetro ad 25 mm. longo et 20 mm. lato,
basi obtuso, apice anguste truncato; seminibus 3 coriaceis triquetris
ca. 18 mm. longis, 6-8 mm. latis, utroque subapiculatis, intus basim
versus operculatis, conspicue ruguloso-tuberculatis praecipue faciebus
exterioribus et carina dorsali, tuberculis obtusis vel interdum carina
complanatis et 1-2 mm. eminentibus.

Type in the U.S. National Herbarium, Nos. 2191516-2191519 (4 sheets),
collected in dense forest in the hills east of the Wainikoroiluva River, near Na-
muamua, Namosi Province, Viti Levu, Fiji, alt. 50-200 m., October 15, 1953,
by A. C. Smith (No. 8900). Duplicates at Bish, ete.

ADDITIONAL SPECIMENS EXAMINED:

FIJI: Virt Levu: Mba: Mountains near Lautoka, Greenwood 1303 (A, US);
Mt. Evans Range, Greenwood 1156 (A); vicinity of Nandarivatu, Degener 14352
(Bish, GH, UC, US). Naitasiri: Raradawai, Wainamo-Wainisavulevu Divide,
St. John 18263 (Bish, US); Suva Pumping Station, Degener & Ordonez 13988
(Bish, GH, UC, US); Central Road 8 miles from Suva, MacDaniels 1152 (Bish).
Kanpavu: Hills above Namalata and Ngaloa Bays, Smith 193 (Bish). VANUA
Levu: Thakaundrove: Vatunivuamonde Mt., Savu Savu Bay region, Degener &
Ordonez 14011 (Bish, GH, UC, US).

The local name “paka”’ is commonly applied to this indigenous
Heliconia, and St. John also records it as “vava ni Viti.’ The plant
is a fairly common component of rain forests and wet thickets at
elevations from near sea level up to at least 750 m.; it is more abundant
than indicated by the number of collections, as evidently collectors
choose not to prepare material of what they take to be a common

SMITH—PACIFIC ISLAND PLANTS, XVIII 71

coarse herb. The leaves are suitable for thatching temporary shel-
ters and the plant is thus well known to local hunters and forest
travelers. There is no indication that this species is an introduction,
and no justification for referring it to the American H. bihai L.
Degener (No. 14352) indicates that the seeds are edible when cooked,
and St. John’s label suggests that the flowers may be eaten raw or
boiled. The collection selected as the type bears good flowers, and
several of the other specimens have flowers as well as fruits. Field
notes indicate that the outer bracts are red, the inner bracts dull
yellow, and the fruit yellow, becoming orange when ripe.

Schumann (Pflanzenr. 1 (IV. 45):36. 1900) implies that his con-
cept of H. bihaz L. includes not only American plants from the West
Indies and Mexico to southern Brazil, but also specimens from Samoa
westward to the Moluccas or perhaps to Sumatra. The Fijian
material, indeed, keys to H. bihai in his system. Subsequent stu-
dents of Heliconia have realized that H. bihai could not logically be
so inclusive. Griggs (Bull. Torrey Club 30:656. 1903) more nar-
rowly defined H. bihat and suggested that it is limited to the West
Indies or Guiana. According to Griggs it has an elongate peduncle
and bracts mostly concealing the rachis. Thus far most of the Old
World taxa lack acceptable specific epithets, but an informative
discussion by Ridley (Agric. Bull. Straits Settlem. 7:129-132. 1908)
provides descriptions and notes for several of them. As far as can
be ascertained at present, none of the names there discussed
seem to apply to the species in Fiji. Specimens are now at hand from
the island groups from Samoa westward; the available material from
the New Hebrides, Solomons, and New Guinea includes several
undescribed species, but none of it requires comparison with H. paka.

The Samoan collections cited as H. bihai by Christophersen (Bishop
Mus. Bull. 128:54. 1935) are superficially similar to my new species,
but their seeds are even more coarsely and irregularly rugulose. The
indument of at least some of the Samoan specimens is more abundant
(on the sheaths and surfaces of the primary bracts), being present
also sometimes on the pedicel and ovary. The inner perianth seg-
ments, insofar as observed, appear to have somewhat thicker and
more obviously raised nerves. In view of these points, I hesitate to
combine the Samoan material with H/. paka, although subsequent
study may indicate this as the correct disposition.

Cunoniaceae

Pullea perryana A. C. Sm. Journ. Arnold Arb, 33:148. 1952.

Ovatau: Summit of Mt. Ndelaiovalau and adjacent ridge, alt. 575-628 m.,
in dense bush and thickets of crest, Smith 7613 (Bish, US, ete.) (tree 12 m. high;
perianth white; disk lobes rich pink; filaments and styles white).

72 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

The third collection of this species, previously known only from
southeastern Viti Levu, is of interest. In inflorescence and floral
characters the Ovalau material is essentially identical with the earlier
collections, but its leaf blades are consistently smaller, being only
4-7 <2-3.5 cm., and correspondingly less coarsely crenate. These
foliage differences may be merely the result of the exposed habitat on
the eastern ridge of Ovalau.

Meliaceae

Aglaia gracilis A. C. Sm. Contr. U.S. Nat. Herb. 30:489. 1952.

Virt Levu: Serua: Hills east of Navua River, near Nukusere, in dense forest,
alt. 100-200 m., Smith 9126 (Bish, US, ete.) (simple-stemmed shrub or small tree
to 5 m. high; inflorescence borne on main stem; petals dull yellow; anthers yel-
lowish white); hills between Waininggere and Waisese Creeks, between Ngaloa
and Wainiyambia, in dry forest, alt. 50-100 m., Smith 9551 (US) (slender tree
2m. high; petals and filaments yellow; anthers nearly white).

This infrequently collected species has not otherwise been recorded
below 750 m. The Serua collections provide the only flowering
material known other than the type, with which they agree excellently
in the very distinctive floral characters. However, they permit a
slight amplification of the original description as follows:

Leaves up to 70 cm. long, the petiole up to 22 cm. long; leaflet
blades up to 287 cm.; inflorescences up to 2 em. long, the lateral
branches several but inconspicuous, up to 6 mm. long.

Dysoxylum gillespieanum A. C. Sm. Contr. U.S. Nat. Herb. 30:516. 1952.

Vitt Levu: Serua: Hills east of Navua River, near Nukusere, alt. 100-200 m.,
in dense forest, Smith 9105 (Bish, US, ete.) (tree 20 m. high; young inflorescence
brownish green; fruit green, ellipsoid, up to 53.5 em.); hills between Waini-
nggere and Waisese Creeks, between Ngaloa and Wainiyambia, alt. 50-100 m.,
in dry forest, Smith 9381 (Bish, US, ete.) (tree 15 m. high; flower buds dull green);
Namboutini, roadside near Nambukelevu, Qoro & Kuruvoli 13717 (Fiji Dept.
Agriculture, Bish) (slender tree 8 m. high; fruit brown, oval).

The cited collections, obtained at lower elevations than the two
specimens thus far known, are interesting in that they verify the
position assigned to D. gillespieanum in my original treatment. The
flowers now available indicate that the species belongs in § Dysorylum
and is indeed a fairly close relative of D. lenticellare Gillespie. No.
9381 has nearly mature inflorescences, while No. 9105 has young
inflorescences and also mature fruits, somewhat larger than those
originally described. Further differentiating characters based on the
inflorescence may now be indicated. Dysoxylum gillespieanum
differs from D. lenticellare in its compact, simply racemose (rather than
branched) inflorescence, its basically 4-merous (rather than 5-merous)
flowers, its disk sericeous-puberulent within (rather than elabrous),

SMITH—PACIFIC ISLAND PLANTS, XVIII 73

and its solitary (rather than paired) ovules. Probably the flowers
are also smaller than those described for D. lenticellare (Contr. U.S.
Nat. Herb. 30:508. 1952), but this cannot be stressed in the absence
of fully mature flowers. The new collections permit amplification of
my earlier description as to foliage dimensions as well as to inflores-
cence:

Leaves up to 40 cm. long, the petiole to 9 cm. long, the petiolules to
25 mm. long; leaflet blades occasionally oblong-oblanceolate and up
to 228 cm., the secondary nerves up to 16 per side; inflorescence
arising from branchlets below leaves, simply racemose, in advanced
bud 1.5-2 cm. long, the peduncle 2-3 mm. long; peduncle, rachis,
bracts, pedicels, and calyx copiously but minutely sericeous with
yellowish hairs scarcely 0.1 mm. long; flowers 10-15 per inflorescence,
solitary in the axils of broadly deltoid, rounded bracts about 12mm. ;
pedicels stout, 1-2 mm. long; calyx gamosepalous, cupuliform,
slightly before anthesis about 2X3 mm., the tube minute, the limb
subearnose, irregularly 4-lobed, the lobes valvate, broadly ovate, up
to 1X2 mm., rounded or obtuse at apex, the sinuses acute; petals 4,
carnose, in advanced bud about 2.5 mm. long and 1.5-2 mm. broad,
subacute, sericeous-puberulent without like the calyx; staminal tube
short-cylindric, subcarnose, about 2 mm. long, glabrous, crenulate
at apex; stamens 7 or 8, the anthers sessile, oblong, obtuse, about 0.9
mm. long; disk carnose, about 1 mm. long and 1.5 mm. in diameter,
crenulate at apex, glabrous without, sparsely sericeous-puberulent
within and with a few marginal setae up to 0.5 mm. long; ovary
copiously yellowish sericeous with hairs about 0.2 mm. long, the
locules 3, each with one ovule affixed near the middle, the style stout,
terete, about 1 mm. long, glabrous, the stigma peltate, about 0.8 mm.
in diameter, crenulate at margin.

Euphorbiaceae

Macaranga (§ Adenoceras) caesariata A. C. Sm., sp. nov.

Arbor gracilis ad 8 m. alta; ramulis teretibus crassis apicem versus
1-1.5 cm. diametro, pilis stramineis 1.5-2.5 mm. longis copiosissime et
pervicaciter villosis; stipulis subcoriaceis elliptico-oblongis ca. 1.5 cm.
longis et 1 cm. latis, apice obtusis, extus copiose villosis, intus glabris;
petiolis validis 12-23 cm. longis ut ramulis villosis; foliorum laminis
amplis late ovatis 25-45 em. longis 20-35 cm. latis, basi rotundatis et
late (5-9 em.) peltatis, apice in acuminem gracilem 1-2 cm. longum
angustatis, margine inconspicue crenulatis, supra glabris vel nervis
et interdum venulis notabiliter setulosis, subtus nervis omnibus ut
petiolo copiose setuloso-villosis ac etiam conspicue miniato-glandu-
losis, nervis primariis 6-8 supra leviter elevatis subtus prominentibus,

74 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

rete venularum intricato subtus prominulo; inflorescentiis o copiose
ramulosis ad 20 cm. longis latisque, ramulis ad nodos pilis pallidis
0.5-1 mm. longis saepe hispidulis ac etiam puberulis (pilis 0.1-0.2 mm.
longis) subglabratis, bracteis majoribus lanceolato-obovatis ad 8 mm.
longis et 1.5 mm. latis irregulariter patelliformi-glandulosis saepe dorso
parce hispidulis; floribus sessilibus 3-8 in glomerulis, calyce sub an-
thesi ca. 1.2 mm. longo et 1.5 mm. diametro eglanduloso 3-lobato,
lobis deltoideo-ovatis acutis, staminibus 7-11 (saepe 9), filamentis ca.
1 mm. longis, antheris transverse ellipsoideis 0.5-0.6 mm. latis;
inflorescentiis ? minoribus sub fructu 5-7 cm. longis latisque, ramulis
copiose et persistenter setuloso-puberulis (pilis 0.1-0.3 mm. longis),
bracteis saepe deltoideo-lanceolatis ad 42 mm. stipite brevi minute
brunneo-sericeis; pedicellis sub fructu 2-3 mm. longis puberulis,
calyce rotato ca. 3 mm. diametro inaequaliter fisso parce sericeo;
fructibus levibus copiosissime sessili-glandulosis, coccis 3.5-4 mm.
diametro, stylis 2 divaricatis 1-1.5 mm. longis.

Type in the U.S. National Herbarium, Nos. 2191715 and 2191716, collected in
dense forest in the hills west of Waivunu Creek, between Ngaloa and Korovou,
Serua Province, Viti Levu, Fiji, alt. 50-150 m., November 28, 1953, by A. C.
Smith (No. 9218). Duplicates at Bish, ete.

ADDITIONAL SPECIMENS EXAMINED:

FIJI: Vitr Levu: Serua: Hills between Waininggere and Waisese Creeks,
between Ngaloa and Wainiyambia, alt. 50-100 m., in dry forest, Smith 9657
(Bish, US, ete.). Rewa: Mt. Korombamba, Meebold 16473 (Bish).

The collection selected as the type bears staminate inflorescences;
T recorded the local names of “ndavo” or “mavo.”” No. 9657 is from a
fruiting specimen, and the Meebold collection consists of a single leaf.
My notes indicate the tree as slender and freely branched, 6-8 m.
high, with copious thin, red latex; the staminate perianth is brick red
and the anthers pale yellow.

The new species is closely related only to M. magna Turrill, from
which it is readily distinguished by the copious and persistent in-
dument of its branchlets, petioles, and lower surfaces of leaf blades.
The corresponding indument of M. magna is not only sparser and
fugacious, but the individual hairs are only 0.2-0.5 mm. long, whereas
the hairs of the vegetative parts of MM. caesariata are 1.5-2.5 mm.
long and form a striking covering. The indument on the inflores-
cence branches is also more obvious in the new species, but there
appear to be no very obvious floral differences. The stamens of the
new species are fewer (7-11 as far as observed), as compared with
12-14 in Af. magna.

Glochidion atalotrichum A. C. Sm., sp. nov.

Arbor monoica multiramosa 4-8 m. alta, ramulis cinereis eracilibus
(apices versus 0.5-1 mm. diametro subflexuosis) juventute copiose

SMITH—PACIFIC ISLAND PLANTS, XVIII 75

stramineo-hirtellis (pilis 0.5-1 mm. longis manifeste septatis 5-8-
cellularibus) ac etiam copiose albo-puberulis (pilis haud 0.1 mm.
longis), indumento longiore mox caduco, breviore subpersistente;
stipulis deltoideo-lanceolatis glabratis 1-1.5 mm. longis mox caducis;
petiolis gracilibus 2-4 mm. longis primo ut ramulis saepe pilosis
mox glabratis; foliorum laminis chartaceis glabris in sicco supra
fuscis subtus pallidioribus, ovato-lanceolatis, 4-7.5 cm. longis 1.52.5
cm. latis vel interdum minoribus, basi obtusis et in petiolum decur-
rentibus, apice in acuminem gracilem 1-1.5 cm. longum gradatim
angustatis, margine integris vel haud undulatis, costa supra acute
elevata subtus prominente, nervis secundariis utrinsecus 5-8 curvatis
anastomosantibus supra subplanis subtus prominulis, rete venularum
supra obscuro subtus subprominulo; inflorescentiis @ axillaribus
breviter racemosis saepe 2-floris, rhachide 0.5-1 mm. longa, bracteis
imbricatis ad 10 deltoideis subacutis glabris haud 0.5 mm. longis,
pedicellis gracilibus glabris sub anthesi 3-6 mm. longis; perianthio
ovoideo, segmentis 6 similibus oblongis 1.8-2 mm. longis 0.8-1 mm.
latis subacutis, extus ut ramulis juvenilibus hirtellis sed interdum
subglabratis; columna staminali ca. 1.2 mm. longa, antheris 3 loculis
connatis ca. 0.7 mm. longis, connectivis in apices oblongos subacutos
ca. 0.3 mm. longos productis; floribus 9 forsan solitariis, pedicello,
perianthii segmentis, et gynoecio ut ramulis juvenilibus hirtellis,
ovario triquetro-ovoideo, columna stylari post anthesin subulata ca.
2 mm. longa, stylis 3 apicem versus liberis; capsulis depresso-globosis
maturitate ad 12 mm. diametro, stylis longe persistentibus et mani-
feste hirtellis, valvis 3 sulcatis demum bilobatis, seminibus rubris
ovoideis 3.5-5 mm. longis 2.5-4 mm. latis.

Type in the U.S. National Herbarium, No. 2191397, collected in dense forest
on the northern slopes of the Korombasambasanga Range, in drainage of
Wainavindrau Creek, Namosi Province, Viti Levu, Fiji, alt. 450-600 m., Septem-
ber 28, 1953, by A. C. Smith (No. 8747). Duplicates at Bish, etc.

ADDITIONAL SPECIMENS EXAMINED:

FIJI: Vitr Levu: Namosi: Hills north of Wainavindrau Creek, between
Korombasambasanga Range and Mt. Naitarandamu, alt. 250-450 m., in dense
forest, Smith 8463 (Bish, US, etc.). Namosi-Naitasiri boundary: Mt.
Naitarandamu, alt. 800 m., Gillespie 3103 (A, Bish, US).

Field notes indicate the plant as a freely branched, spreading tree
4-8 m. high, with yellowish-green perianth segments. My two
collections are clearly monoecious, bearing staminate flowers as well
as pistillate, although the latter are scarce; No. 8463 also bears a
few fruits. The Gillespie specimen bears detached fruits that appear
to be fully mature.

Among the species of our area, @. atalotrichum is recognized by the
characteristic multiseptate hairs of the young vegetative parts, which

228-962—67——2

76 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

are also often present on the perianth segments and on the gynoecium,
persisting on the styles in fruit. From G. seemannii Muell. Arg.,
which it resembles superficially, the new species differs not only in its
indument, but also in its long-acuminate leaf blades and its three
styles.

Anacardiaceae
Pleiogynium hapalum A. C. Sm., sp. nov.

Arbor dioica gracilis 5-20 m. alta, ramulis subteretibus apicem
versus 5-10 mm. diametro, partibus novellis pilis fusco-stramineis
mollis patentibus simplicibus 0.2-0.5 mm. longis copiose indutis; foliis
imparipinnatis (20—) 30-50 em. longis, petiolis (3-) 4-12 em. longis et
rhachide subteretibus et copiose contumaciter pilosis; foliolis (9-) 11
vel 13, petiolulis gracilibus leviter canaliculatis 5-10 mm. longis
(terminali ad 30 mm. longo) ut rhachide pilosis; foliolorum laminis
chartaceis in sicco supra fuscis subtus viridescentibus inaequaliter
ovatis, (6-) 8-13 em. longis, (2.5-) 4.5-6 cm. latis (inferioribus saepe
minoribus), basi inaequaliter subacutis vel obtusis et in petiolulum
decurrentibus, apice acutis vel in acuminem ad 1 cm. longum cuspidatis,
margine integris vel obscure undulatis, supra praeter costam et nervorum
secundariorum bases pilosas glabris, subtus conspicue et persistenter
pilosis (pilis ad 0.5 mm. longis praesertim costa et nervis permansis),
costa supra elevata subtus prominente, nervis secundariis utrinsecus
6-10 patentibus supra planis vel prominulis subtus elevatis, rete venu-
Jarum intricato supra immerso subtus haud prominulo; inflorescentiis
o@ axillaribus late paniculatis multifloris 10-30 em. longis 4-12 em.
latis, pedunculo 2-10 em. longo, ramulis numerosis gracilibus inferiori-
bus 2-7 cm. longis, axibus omnibus ut foliolorum rhachide copiose
pilosis, bracteis ad nodos deltoideo-oblongis obtusis 0.5-1 mm. longis
subacutis extus pilosis; floribus oc subsessilibus laxe aggregatis vel
apice ramulorum ultimorum brevium solitariis, pedicello super arti-
culationem ad 1 mm. longo bracteas obscuras 1-3 oblongas 0.2-0.5 mm.
Jongas inconspicue glanduloso-ciliolatas gerente; floribus glabris, calyce
subrotato 1.7-3 mm. diametro, lobis submembranaceis semiorbiculari-
oblongis 0.5-0.8 mm. longis 0.7-1.2 mm. latis margine glanduloso-
ciliolatis, petalis plerumque 5 (raro 4 vel 6) anguste imbricatis sub-
membranaceis oblongis 1.8-2.2 mm. longis 1.3-1.5 mm. latis apice
obtusis 38-5-nervatis; disco annulari-pulvinato 1.5-2 mm. diametro
manifeste crenulato; staminibus plerumque 10 (interdum 9-12), fila-
mentis filiformi-subulatis 0.6-1.5 mm. longis, antheris ovatis versatili-
bus 0.5-0.7 mm. longis apice obtusis, thecis basi divergentibus;
gynaecio in floribus & abortivo in disco immerso, stylis rudimentariis
cono Inconspicuo 5-7-suleato adnatis vel raro minute liberis; in-
florescentiis 9 quam © simplicioribus similiter pilosis 3-6 em. longis

SMITH—PACIFIC ISLAND PLANTS, XVIII 77

paucifloris; perianthio ut & sed calyce post anthesin incrassato, petalis
ad 32.5 mm.; staminodiis saepe 14, filamentis 0.7-1 mm. longis
reflexis, antheris minutis sterilibus; disco conspicue pulvinato carnoso
incrassato; ovario oblato-sphaeroideo apice rotundato basim versus
angustato, stylis crassis 7 vel 8 ca. 0.5 mm. longis e ovarii margine
distali patentibus, loculis 7 vel 8, ovulis solitariis pendulis; inflo-
rescentia sub fructu ramulis pedunculisque persistenter pilosis; drupa
late turbinata 12-14 mm. alta paullo supra medium 18-20 mm. lata
et ibi manifeste 7- vel 8-angulata, basi et apice rotundata.

Type in the U.S. National Herbarium, No. 2191567, collected in dense forest
in the hills east of the Wainikoroiluva River, near Namuamua, Namosi Province,
Viti Levu, Fiji, alt. 50-200 m., October 15, 1958, by A. C. Smith (No, 8958).
Duplicates at Bish, ete.

ADDITIONAL SPECIMENS EXAMINED:

FIJI: Virt Levu: Mba: Vicinity of Nandarivatu, Gillespie 3189 (Bish, GH,
UC, US). Namosi: Northern slopes of Korombasambasanga Range, in drainage
of Wainavindrau Creck, Smith 8699 (Bish, US, etc.); hills bordering Wainavi-
ndrau Creek, in vicinity of Wainimakutu, Smith 8531 (Bish, US, ete.) ; vicinity of
Nanggarawai, Gillespie 3227 (Bish, GH, UC, US); vicinity of Namuamua, Gil-
lespie 2977 (Bish, GH, UC), 3058 (Bish, GH, UC). Tailevu: Hills east of Wai-
nimbuka River, in vicinity of Ndakuivuna, Smith 7164 (Bish, US, ete.). Vanua
Levu: Thakaundrove: Natewa Bay region, hills west of Korotasere, Smith
1940 (Bish, GH, UC, US).

Field notes indicate the plant as a slender tree 5-20 m. high, oc-
curring in forested areas at elevations of 50-750 m.; the petals are
yellow or greenish yellow, the filaments white or greenish yellow,
and the anthers yellow. Pistillate inflorescences are found only on
Gillespie 3189, fruits on Smith 7164, and staminate inflorescences on
the other collections.

As Leenhouts has pointed out (Blumea 7:159. 1952), the correct
name for the widespread tree that has long been passing as Pleiogy-
nium solandri (Benth.) Engl. is P. timoriense (DC.) Leenh. This
occurs from at least Timor through Queensland and into the Pacific
as far as Tonga and the Cook Islands, being frequent in Fiji. It is
characteristically glabrous, but some individuals in Fiji and Tonga
(and perhaps also in Australia to judge from Engler’s description in
DC. Monogr. Phan. 4:255. 1883) are puberulent on the young parts
of the branchlets, petiole, and leaf rachis. However, these parts
bear hairs scarcely exceeding 0.1 mm. in length and they are soon
elabrate. The species here described has no consequential inflo-
rescence or fruit differences from the glabrous individuals, but the
copious and persistent indument of its vegetative parts and inflo-
rescence axes seems too striking to permit its inclusion in the wide-
spread species.

78 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Another species of this relationship, P. papuanum C. T. White
(Proc. Roy. Soc. Queensl. 45:27. 1934), differs from P. hapalum in
its smaller, glabrous leaves with fewer leaflets, its smaller flowers,
and the furfuraceous gray scurfy indument of its branchlets and
rachis.

Gillespie inscribed some of his collections which are cited above
with a binomial apparently based on Dracontomelum pilosum Seem.
and accredited to Engler; however, no such binomial appears to have
been published. At any rate Dracontomelum pilosum is quite a
different plant, recently referred to the synonymy of Dysorylum
quercifolium (Seem.) A.C. Sm. (Brittonia 14:245, 1962).

Celastraceae

Cassine vitiensis (A. C. Sm.) A. C. Sm., comb. nov.
Elaeodendron vitiense A. C. Sm. Journ. Arnold Arb. 31:289. 1950.

Vitt Levu: Serua: Hills between Waininggere and Waisese Creeks, between
Ngaloa and Wainiyambia, alt. 50-100 m., in dry forest, Smith 9385 (Bish, US,
etc.). OvaLau: Hills southeast of valley of Mbureta River, alt. 100-300 m., in
dense forest, Smith 7450 (Bish, US, ete.).

Ding Hou (FI. Males. I. 6:284. 1963) gives cogent reasons for
combining Elaeodendron and Cassine, in which conclusions it seems
advisable to follow him. The second and third collections of the
apparently uncommon Fijian species show differences from the type,
but these do not appear very significant. No. 9385 bears flowers
that are substantially smaller than those originally described, and
have the ovary only 2(rather than 3- or 4-)-locular. Fruits are present
in both the newer collections. My original description may be
amplified as follows:

Tree up to 20 m. high, the leaf blades up to 13 em. long and 8.5
cm. broad; pedicels in flower often only 1.5-2 mm. long and the flowers
only 5-6 mm. in diameter, the petals as small as 2.52 mm., the
filaments as short as 0.5 mm., the anthers only 0.4 mm. broad, the
ovary-locules sometimes only 2; pedicels in fruit up to 7 mm. long,
the disk, stamens, and stigma subpersistent; fruits ellipsoid, up to
22 mm. long and 15 mm. broad, obtuse at base, acute or cuspidate
at apex, the epicarp thin but coriaceous, smooth, the mesocarp fibrous,
hard in dried fruits, 1.5-2 mm. thick, the endocarp hard and bony,
1-2 mm. thick, inconspicuously rugulose without, forming 2 separable
putamens, these laterally flattened, up to 15X75 mm.; seed solitary,
ellipsoid-lanceolate, about 126 mm., strongly flattened, obtuse or
subacute at both ends.

SMITH—PACIFIC ISLAND PLANTS, XVIII 79

Elaeocarpaceae

Elaeocarpus (§ Blepharoceras) milnei Seem. Fl. Vit. 28. 1865; A. C. Sm. Contr.
U.S. Nat. Herb. 30:556. 1953.

Vitt_ Levu: Namosi: Northern slopes of Korombasambasanga Range, in
drainage of Wainavindrau Creek, alt. 450-600 m., in dense forest, Smith 8731
(Bish, US, ete.).

It is a satisfaction to have available a second collection of a species
otherwise known only from the type specimen, the more so as the
new material agrees almost precisely with the type (Milne 81) and
has adequate color notes. In my observation flower color provides a
dependable character in Hlaeocarpus, but it is too frequently ignored.
No. 8731 has the sepals dull pink, and the petals rich pink with pale
yellow apical laciniae. The new material permits the following slight
amplification of my 1953 description:

Tree up to 15 m. high, the branchlets up to 17 mm. in diameter
near apex; leaf blades up to 38 cm. long and 17 cm. broad, the marginal
teeth often 1 cm. apart; sepals up to 11.5 mm. long and 3.5 mm. broad;
petals up to 14 mm. long and 8 mm. broad, the apical lobes up to
12 in number; disk about 1 mm. high, with hairs about 0.3 mm. long;
anthers with dorsal awns sometimes 0.8 mm. long; hairs of the ovary
up to 0.3 mm. long, the ovules 8 per locule.

Except for the above trivia, my specimen appears identical with
the type in such important characters as leaf shape, inflorescence
dimensions, and indument. ‘These details are recorded at this time
primarily for discussion in reference to the following new species.

Elaeocarpus (§ Blepharoceras) chionanthus A. C. Sm., sp. nov.

Arbor gracilis ad 8 m. alta, partibus novellis pilis ferrugineis 0.4—0.6
mm. longis copiosissime hispidulo-hirtellis, ramulis subteretibus
crassis apicem versus 7-10 mm. diametro demum glabratis; foliis
apices ramulorum versus congestis, petiolis crassis 3-4 mm. diametro
leviter canaliculatis 2-6 cm. longis persistenter hirtellis (pilis 0.2-0.3
mm. longis); foliorum laminis subcoriaceis in sicco fusco-viridibus
oblongo-ellipticis, magnitudine diversis sed maturitate 25-30 cm.
longis et 13-17 cm. latis, basi plerumque rotundatis (obtusis vel
subcordatis), apice rotundatis vel late obtusis vel obtuse cuspidatis,
margine anguste recurvatis et dentibus plerumque 1-1.5 cm. distanti-
bus obscure calloso-apiculatis inconspicue crenatis, supra glabris vel
costa obscure strigillosis, subtus copiose et pervicaciter pilosis (pilis
ferrugineis vel stramineis mollibus 0.3-0.6 mm. longis costa secunda-
riisque densissimis sed etiam lamina manifestis), costa supra valde
elevata subtus prominente, nervis secundariis utrinsecus 9-13 patenti-
curvatis et marginem versus anastomosantibus supra paullo elevatis
subtus prominentibus, rete venularum intricato utrinque prominulo;

80 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

inflorescentiis racemosis e ramulis infra folia enatis 25-35-floris,
pedunculo subnullo (ad 38 mm. longo), rhachide gracili 12-16 cm.
longa et pedicellis pilis 0.2-0.4 mm. longis copiose ferrugineo-
setuloso-hirtellis, bracteis sub floribus lanceolatis ca. 1.5 mm. longis
dense pilosis caducis, pedicellis sub anthesi crassis ca. 5 mm. longis;
sepalis 5 subcarnosis lanceolatis 10-11 mm. longis ca. 3 mm. latis
subacutis, extus copiose puberulis (pilis 0.1-0.2 mm. longis), intus
carinatis et parce sericeis; petalis 5 basim versus incrassatis superne
papyraceis, oblongo-obovatis, 11-13 mm. longis, 5-7 mm. latis,
utrinque glabris vel intus basim versus obscure pilosis, apice lobis

8-10 fimbriatis, laciniis subaequalibus 1.5-2 mm. longis subacutis

1-4-nervatis; disco pulvinato ca. 1 mm. alto 5-lobato, lobis sulcatis

pilis 0.2-0.4 mm. longis hispidulis; staminibus 23-25 uniseriatis

6-7 mm. longis, filamentis gracilibus 2-3 mm. longis obscure puberulis,

antheris 3.8—4.2 mm. longis copiose et minute tuberculato-hispidulis,

arista dorsali 0 2-0.3 mm. longa, apice ventrali rotundato; ovario
lanceolato-ovoideo ut disco copiose sericeo-hispidulo, stylo subulato ca.

5 mm. longo superne glabro, ovarii pariete incrassato, loculis 2, ovulis

in quoque loculo 8 biseriatis.

Type in the U.S. National Herbarium, Nos. 2191946 and 2191947, collected in
dense forest in hills west of Waivunu Creek, between Ngaloa and Korovou, Serua
Province, Viti Levu, Fiji, alt. 50-150 m., December 7, 1958, by A. C. Smith
(No. 9495). Duplicates at Bish, ete.

Field notes indicate the plant to be a slender tree to 8 m. high, with
petals and filaments pale green, becoming pure white, the anthers
yellow, the gynoecium pale green, and the largest observed leaf blades
up to 80X17 cm. Small leaf blades are often mixed with the larger
ones on the same shoot. The new species is closely related only to
FE. milnei Seem., from which it differs in several characters best
summarized as follows:

Indument comparatively sparse and short (hairs of various parts usually less
than 0.2 mm. long, the leaf blades essentially glabrous beneath); leaf blades
obovate, gradually narrowed toward base; petals rich pink, with pale yellew
laciniae; anthers with dorsal awns 0.5-0.8 mm. long... . . . E. milnei

Indument comparatively copious and long (hairs of various parts 0.2-0.6 mm.
long, the leaf blades densely and persistently pilose beneath); leaf blades

oblong-elliptic, usually rounded at base; petals pure white at anthesis;
anthers with dorsal awns 0.2-0.3 mm. long... . . . . E. chionanthus

Melastomataceae
Astronidium pallidiflorum A. C. Sm., sp. nov.

Arbor ad 15 m. alta, partibus novellis, ramulis petiolisque pilis
capitato-glandulosis graciliter stipitatis 0.1-0.3 mm. longis copiose
ferrugineo-pubescentibus; ramulis apicem versus 3-7 mm. crassis et
ibi complanato-quadrangularibus; petiolis gracilibus | 1.5-2 mm.

SMITH—PACIFIC ISLAND PLANTS, XVIII 81

diametro subteretibus 3-5 cm. longis, laminis crasso-chartaceis
oblongo-lanceolatis 13-23 em. longis 4-7.5 cm. latis, supra olivaceis
et nervis principalibus ut petiolis pubescentibus demum glabratis,
subtus praecipue nervis copiose pilosis (pilis clavatis ad 0.5 mm. longis
vel superficie interdum ad 0.8 mm. longis), basi obtusis, apice in
acuminem 1.5-2 em. longum calloso-subacutum gradatim angustatis,
margine dentibus calloso-acutis manifestis 0.5-1.5 cm. distantibus
crenulatis, 5-nerviis, nervis e basi adscendentibus, exterioribus 2
comparate inconspicuis et 1-2 mm. intra marginem, costa et nervis
2 interioribus supra subplanis subtus prominentibus, venulis transver-
sis et rete venularum laxo supra planis subtus prominulis; inflorescentia
terminali trichotome cymosa sessili e basi 3-divisa ad 13 em. longa
et 20 cm. lata, ramulis crassis conspicue complanatis et pedicellis et
calyce ut partibus novellis copiose pubescentibus, bracteis bracte-
olisque mox caducis non visis, floribus apices ramulorum ultimorum
versus 2-5 aggregatis, pedicellis crassis 1-1.5 mm. longis; calyce
submaturo cupuliformi 4—4.5 mm. longo et diametro apice lobis plerum-
que 10 subdeltoideis obtusis ca. 0.5 mm. longis inaequabiliter fisso;
petalis glabris in floribus submaturis 5 oblongis ad 42.5 mm. apice
rotundatis, staminibus 10 valde inflexis, filamentis ligulatis ca. 3 mm.
longis, antheris oblongis ca. 3 mm. longis apice conspicue recurvatis,
caleari basilari conspicuo ca. 0.8 mm. longo; stylo tereti petalis
subaequali, ovarii loculis 5, placentis clavatis, ovulis numerosissimis.

Type in the U.S. National Herbarium, No. 2191790, collected in dense forest
in the hills west of Waivunu Creek, between Ngaloa and Korovou, Serua Province,
Viti Levu, Fiji, alt. 50-150 m., November 26, 1953, by A. C. Smith (No. 9313).
Duplicates at Bish, ete.

Field notes indicate the plant as a tree 15 m. high, with pale greenish
white petals, filaments, anthers, and style. The new species is closely
related only to A. kasiense A. C. Sm., with which it has in common
a copious indument of clavate hairs, these being very different from
the much longer, subulate hairs of A. storckii Seem., the only other
Fijian species of this general affinity. The principal points of dif-
ference between the new species and A. kasiense are summarized
as follows:

Indument of young parts, petioles, inflorescence branches, ete., composed of
hairs usually 0.2-0.5 mm. long; leaf blades ovate-clliptic, 7-12 cm. broad,
rounded to obscurely subcordate at base, the fourth and fifth nerves 4-5 mm.

within the margin and conspicuously connected by transverse veinlets to an
obvious submarginal nerve; flowers (as far as known) 4-merous . A. kasiense
Indument of young parts, petioles, inflorescence branches, ete., comparatively
close, composed of hairs usually 0.1-0.3 mm. long; leaf blades oblong-
lanceolate, 4-7.5 em. broad, obtuse at base, the fourth and fifth nerves
submarginal, 1-2 mm. within the margin and lacking exterior veinlets; flowers
(as far as known) 5-merous. . . «6 ee ee ee ee eA pallidiflorum

82 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Medinilla subviridis A. C. Sm. Journ. Arnold Arb. 33:101. 1952.

Vitt Levu: Namosi: Hills north of Wainavindrau Creek, between the Koro-
mbasambasanga Range and Mt. Naitarandamu, Smith 8487 (Bish, US, ete) ; north-
ern slopes of Korombasambasanga Range, in drainage of Wainavindrau Creek,
Smith 8748 (Bish, US, etc.). Serua: Hills between Waininggere and Waisese
Creeks, between Ngaloa and Wainiyambia, Smith 9541 (Bish, US, etc.).

The three cited collections of this supposedly rare plant, formerly
known from a single locality near Mt. Tomanivi, extend the known
range southward in Viti Levu, and downward to the low elevation of
50-100 m. recorded for No. 9541. Color notes are remarkably con-
sistent with those recorded in the original description. However, it
may be noted that the petioles of the larger leaves are sometimes
inconspicuous and only 2 mm. long, the blades themselves may be as
large as 178 cm., and the inflorescence may be up to 10 em. long.

Medinilla spectabilis A. C. Sm., sp. nov.

Frutex alte scandens ubique mox glaber sed partibus novellis parce
furfuraceo-puberulis (pilis stramineo-ferrugineis 0.1-0.3 mm. longis
multicellularibus lateraliter brevi-calcaratis vel obscure plumulosis),
ramulis gracilibus subteretibus, internodiis plerumque 2-4 em. longis;
foliis similibus vel disparibus; foliis majoribus: petiolis gracilibus
leviter canaliculatis 13-25 mm. longis, laminis maturis chartaceis vel
papyraceis in sicco fusco-olivaceis ellipticis, 6-12 em. longis, 3.5-6.5
cm. latis, basi obtusis et in petiolum breviter decurrentibus, apice
obtusis vel in acuminem ad 5 mm. longum obtuse cuspidatis, plerum-
que @ basi 7-nerviis, nervis superioribus et costa e basi liberis vel ad
5 mm. conjunctis supra subplanis subtus elevatis, nervis aliis debiliori-
bus, marginalibus interdum inconspicuis, venulis supra immersis
subtus prominulis vel planis; foliis minoribus: petiolis saepe haud
3 mm. longis, laminis interdum 2.52 cm., in forma et textura
majoribus similibus; inflorescentiis axillaribus vel e ramulis infra folia
orientibus pauciramosis, racemosis vel divaricato-cymosis, 5-25 cm.
longis latisque, ramulis gracilibus in vivo molliter carnosis in sicco
saepe complanatis, internodiis 10-17 mm. longis; bracteis e nodis
2 vel 3 membranaceis vel papyraceis elliptico-oblongis, 12-20 mm.
Jongis, 5-12 mm. latis, basi et apice rotundatis, sessilibus; floribus e
nodis 1-3, pedicellis gracilibus sub anthesi 5-10 mm. longis; bracteolis
apice pedicelorum 2 florem plus minusve obtegentibus late vel
manifeste imbricatis, textura et colore bracteis similibus, elliptico-
suborbicularibus, 18-22 mm. longis, 13-15 mm. latis, basi obtusis,
apice rotundatis, 7-9-nervatis, rete venularum teneri ; calyce in vivo
subearnoso, hypanthio cupuliformi 5-7 mm. longo et diametro, basi
obtuso et minute stipitato, ut pedicello obscure disperse-glanduloso
(glandulis haud 0.05 mm. longis minute stipitatis), limbo erecto
membranaceo 2-3 mm. longo integro, dentibus 4 obsoletis submar-

SMITH—PACIFIC ISLAND PLANTS, XVIII 83

ginalibus; petalis 4 membranaceis late obovatis, 17-20 mm. longis,

14-17 mm. latis, apice rotundatis vel obscure retusis, basi late obtusis,

multinerviis; staminibus 8, filamentis gracilibus ligulatis 6-7 mm.

longis basi haud 0.5 mm. latis, antheris oblongis 4.5-5 mm. longis

basi manifeste trilobulatis (lobis anterioribus ca. 0.5 mm. longis,
posteriore minore), apice in tubulum 0.5-1 mm. longum poro introrso-

terminali dehiscentem angustatis; stylo tereti subcarnoso 10-12 mm.

longo, stigmate minuto.

Type in the U.S. National Herbarium, Nos. 2191067 and 2191068, collected in
dense forest in the hills east of Somosomo, west of the old crater occupied by a
small swamp and lake, Taveuni, Fiji, alt. 660-900 m., August 18, 1953, by
A. C. Smith (No. 8362). Duplicates at Bish, ete.

The specimens were collected from a high-climbing liana and were
noted as follows: Inflorescence arising from stem near base, or some-
times associated with leaves; inflorescence branches, bracts, and
bracteoles magenta; calyx white, suffused with pink; petals pale
pink; filaments and style white; anthers rich blue, the basal lobes
yellow.

The tangimauthia of Fijian legend (MM. waterhouser Seem.) is often
considered the most beautiful indigenous plant of the archipelago; it
appears limited to the summit ridge of Taveuni and to Mount Seatura
on Vanua Levu. (A color photograph was recently published as the
frontispiece of J. W. Parham’s Plants of the Fiji Islands, 1964.)
To find another spectacular Medinilla near the “lake” of Taveuni
was a delightful surprise, the more so as the second species is nearly,
if not quite, as exquisite as the tangimauthia. However, it is in-
stantly distinguished, although notreasonably allied to any other species.
Our new plant most strikingly differs from ©]. waterhousei in the color
of its inflorescence parts and petals, and particularly in having its
flowers clasped by two imbricate bracteoles, rather than by the
strictly opposed and often obcordate bracteoles so characteristic of
M. waterhousei. Differences between the two species may be sum-
marized as follows:

Leaves of mature plants isomorphic; leaf blades 4-8X2.5-4 cm., attenuate to
acute at base, 3- or 5-nerved; inflorescence branches, bracts, bracteoles, and
pedicels brilliant red or scarlet; pedicels 15-25 mm. long at anthesis; flower-
subtending bracteoles strictly opposed, ovate, 23-35 15-30 mm., rounded
or subcordate at base; flowers comparatively large, the petals white, 23-
26X 20-25 mm.; filaments broadly ligulate, 1.2-1.5 mm. broad, 8-10 mm.
long; anthers 7-8 mm. long, with only the posterior basal lobe obvious; style
20-22 mm.long. . . . 2... ee ee te ee M. waterhousei

Leaves of mature plants isomorphic or dimorphic; leaf blades of larger leaves
6-12 3.5-6.5 em., obtuse at base, usually 7-nerved; inflorescence branches,
bracts, bracteoles, and pedicels magenta or pink; pedicels 5-10 mm. long at
anthesis; flower-subtending bracteoles obviously imbricate, elliptic-orbicular,
18-22% 13-15 mm., obtuse at base; flowers smaller, the petals pale pink,

228-962—67——3

84 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

17-20X 14-17 mm.; filaments about 0.5 mm. broad, 6-7 mm. long;
anthers 4.5-5 mm. long, obviously 3-lobulate at base; style 10-12 mm. long.
M. spectabilis

Medinilla decora A. C. Sm., sp. nov.

Frutex scandens, partibus novellis, ramulis, petiolis inflorescentiae
ramulisque ferrugineo-furfuraceis (pilis 0.1-0.4 mm. longis multicellu-
laribus lateraliter brevi-calcaratis interdum ad nodos 0.8 mm. longis),
ramulis subteretibus gracilibus, internodiis 3-4 cm. longis; foliis sub-
similibus, petiolis gracilibus leviter canaliculatis 2-4 em. longis, laminis
chartaceis in sicco fusco-olivaceis subtus viridescentibus, elliptico-
lanceolatis, (6-) 8-14 cm. longis, (2.5-) 4-6 cm. latis, basi longe at-
tenuatis et in petiolum gradatim decurrentibus, apice obtusis vel
obtuse cuspidatis, margine anguste recurvatis subintegris vel obscure
crenulatis, 5-nerviis, nervis superioribus cum costa 1-3 cm. conjunctis
supra leviter elevatis subtus prominentibus, nervis inferioribus intra
marginem 2-3 mm. paullo debilioribus, venulis transversis subtus
prominulis, rete venularum immerso; inflorescentia axillari solitaria
laxa pauciflora 3-8 cm. longa, pedunculo 3-20 mm. longo et rhachide
gracilibus; bracteis 2-4 e nodis papyraceis oblongo-lanceolatis, 3-5 mm.
longis, 0.6-1.5 mm. latis, apice obtusis, furfuraceis vel puberulis:
floribus 2-4 e nodis, pedicellis gracilibus 3-6 mm. longis; bracteolis
apice pedicellorum binis membranaceis calycem obtegentibus ellipticis,
6-7 mm. longis, ca. 4 mm. latis, 5-7-nervatis, apice rotundatis,
utrinque parce puberulis subglabratis; calyce 4-5 mm. longo et apice
diametro, hypanthio cupuliformi, limbo suberecto submembranaceo
1-1.5 mm. longo margine integro, dentibus 4 minutis submarginalibus;
petalis 4 membranaceis glabris obovatis sub anthesi 7-8 mm. longis et
6-7 mm. latis, apice rotundatis vel subretusis et obscure mucronulatis,
5-nerviis; staminibus 8 glabris, filamentis gracilibus ligulatis 3-3.5
mm. longis, antheris oblongis 1.7-2 mm. longis, basi trilobulatis (lobis
2 anterioribus ca. 0.5 mm. diametro, posteriore paullo minore), apice
obtusis poro unico terminali dehiscentibus; stylo filiformi 6-7 mm.
longo, stigmate minuto.

Type in the U.S. National Herbarium, No. 2191868, collected in dry forest in
the hills between Waininggere and Waisese Creeks, between Ngaloa and Waini-
yambia, Serua Province, Viti Levu, Fiji, alt. 50-100 m., November 30, 1953, by
A. C, Smith (No. 9398). Duplicates at Bish, ete.

The specimens were collected from a high-climbing liana; the bracts
and pedicels are green, the bracteoles dull pink, the calyx pale pink,
the petals rich pink, the filaments and style pink-tinged, and the an-
thers purple, yellow at base. Color of inflorescence parts in Medinilla,
although perhaps not basically very significant, often provides depend-
able differentiating characters and clues to morphological differences.

The new species is closely related only to M4. kandavuensis A. C. Sm.,

SMITH—PACIFIC ISLAND PLANTS, XVIII 85

which it resembles in foliage and pubescence, although the indument
of the Serua plant is somewhat more obvious and persistent. Medt-
nilla decora differs in its smaller flower-subtending bracteoles and espe-
cially in its smaller flowers; the calyx of M. kandavuensis is 8-9 mm.
long; the petals are 12-13 10-11 mm.; the anthers are 4-5 mm. long
with the posterior basal lobe slightly larger than the anterior ones;
and the style is 12-13 mm. long.

From M. rhodochlaena A. Gray, which more nearly suggests the new
species in its floral dimensions, M. decora differs in its consistently
isomorphic leaves, the blades of which are thinner in texture and longer
attenuate at base. In inflorescence characters, M/. rhodochlaena as
compared with the new species has broader bracts, flower-subtending
bracteoles that are comparatively thick in texture and orbicular or
reniform rather than elliptic, and a slightly smaller calyx at anthesis.
Gray’s species, now known from abundant material, has richly colored
inflorescences, the inflorescence bracts and bracteoles being rich pink
to purple red (rather than green and dull pink respectively in the new
species); its calyx is also a deeper pink.

Medinilla ovalifolia (A. Gray) A. C. Sm., comb. nov.

Anplectrum ovalifolium A. Gray, Bot. U.S. Expl. Exped. 1:597. 1854.
Medinilla amoena Seem. FI. Vit. 88. 1865.

Medinilla parvifolia Seem. Fl. Vit. 89. 1865.

Allomorpha ovalifolia Triana, Trans. Linn. Soc. 28:74. 1871.

The type specimen of Anplectrum ovalifolium (US 47676) was
collected by the U.S. Exploring Expedition in the vicinity of Mbua
Bay, Province of Mbua, Vanua Levu, Fiji. The available material
was taken from a plant in bud, and apparently from a branch exposed
to sun. In foliage this specimen is very similar to Smith 1671, also
from Mbua (summit of Mt. Seatura). The form represented by these
two specimens is a fairly atypical form of the species usually referred
to M. amoena Seem., in that its leaf blades are nearly rounded at base.
But in all other respects the form must be placed with M. amoena,
and it fits my description of that species in Sargentia 1:85. 1942. As
the specific epithet is available in Medinilla, I see no way to avoid a
new combination based on Anplectrum ovalifolium to replace the
frequently collected and well known M. amoena. More complete
synonymy, a description, and citations of specimens are found in my
1942 publication.

Araliaceae

Polyscias culminicola A. C. Sm., sp. nov.
Arbor ad 8 m. alta, ramulis crassis et petiolorum basibus parce

stramineo-puberulis mox glabratis; foliis ad apices ramulorum con-
gestis imparipinnatis multijugis 30-40 cm. longis, petiolo 5-9 cm.

86 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

longo supra complanato basi incrassato sed haud alato, rhachide
subtereti; foliolis 19-23, petiolulis gracilibus canaliculatis 4-6 (termi-
nali ad 15) mm. longis, laminis in sieco chartaceis fusco-olivaceis,
ovato-ellipticis lateralibus manifeste falcatis, 5-7 cm. longis (inferiori-
bus minoribus), 2.5-3 cm. latis, basi inaequilateraliter obtusis vel
rotundatis, apice subacutis, margine integris paullo incrassatis, costa
supra paullo elevata subtus prominente, nervis secundariis utrinsecus
plerumque 6-11 patentibus margines versus curvatis et anastomo-
santibus utrinque planis vel subtus prominulis, rete venularum
immerso; inflorescentiis terminalibus racemoso-paniculatis, ad 23 cm.
longis et 18 cm. latis, inflorescentiae pedunculo brevi, rhachide,
ramulis, umbellarum pedunculis pedicellisque stramineo-puberulis
(pilis patentibus pluricellularibus glandulosis 0.1-0.2 mm. longis);
umbellarum pedunculis gracilibus 1-2.5 em. longis interdum infra
medium minute bibracteatis, bracteis sub floribus oblongis obtusis
0.5-0.7 mm. longis puberulis caducis; floribus in umbellulis 11-32
dense aggregatis, pedicellis sub anthesi gracilibus 3-4 mm. longis
infra flores paullo incrassatis et articulatis; calyce obconico sub anthesi
1.5-2 mm. longo et apice 2-2.5 mm. diametro, limbo minuto obscure
5-denticulato, disco subcupuliformi carnoso; petalis 5 oblongo-
deltoideis sub anthesi 2.5-3 mm. longis ca. 1.5 mm. latis, apice acutis
et cucullatis, intus obscure carinatis; staminibus 5, filamentis sub-
carnosis teretibus 1-1.5 mm. longis, antheris oblongis 2-2.3 mm.
longis apice rotundatis, thecis basi liberis; stylis 2 fere ad basim liberis
carnosis subacutis ca. 1.5 mm. longis.

Type in the U.S. National Herbarium, No. 1965344, collected in crest forest
on a wind-swept ridge, northern slopes of Mt. Namendre, east of Mt. Koromba
(Pickering Peak), Mba Province, Viti Levu, Fiji, alt. 750-900 m., May 27, 1947,
by A. C. Smith (No. 4514). Duplicates at A, Bish, ete.

Although the new species bears a superficial resemblance to P.
joskeit Gibbs, it clearly differs in its umbellate flowers, those of P.
josket being individually scattered on the ultimate inflorescence
branches. Poyscias josket further differs from P. culminicola in
lacking the minute glandular indument described above, in having
its leaflets only 7-15 and with blades that are not strongly falcate,
only slightly inequilateral at base, and obviously undulate-serrulate,
and in its 4-merous flowers that are borne on pedicels less than 2 mm.
long. A local name recorded for the new species is “sawira.”’

Schefflera (§ Heptapleurum) euthytricha A. C. Sm., sp. nov.

Arbor gracilis 5-6 m. alta, ramulis teretibus crassis apices versus
1-1.5 cm. diametro, partibus novellis crasse et conspicue hirsutis
(pilis saepe 5-10 mm. longis, partibus vegetativis demum caducis
sed nodis, petiolis, et foliolorum costa illic persistentibus, inflorescentia

SMITH—PACIFIC ISLAND PLANTS, XVIII 87

pervicacibus); foliis digitatis mox glabratis, petiolis crassis sub-
teretibus 21-30 em. longis, basi in vaginam coriaceam 2-3 cm.
latam incrassatis, ligula obtusa ad 1 cm. longa; foliolis 7-9, petiolulis
in sicco rugulosis 2-6.5 cm. longis, laminis subcoriaceis in sicco fusco-
olivaceis subtus pallidioribus, elliptico- vel oblongo-oblanceolatis,
18-27 cm. longis, 7-10 cm. latis, basi inaequilateraliter obtusis vel
subacutis, apice in acuminem calloso-obtusum 5-10 mm. longum
cuspidatis, margine irregulariter et inconspicue crenato-serrulatis
(dentibus saepe 2 vel 3 per cm. antrorse calloso-apiculatis), costa
valida supra elevata subtus prominente, nervis secundariis utrinsecus
12-15 erecto-patentibus marginem versus coplose anastomosantibus
supra subplanis subtus valde elevatis, rete venularum subimmerso
vel interdum subtus prominulo; inflorescentia apices ramulorum
versus laterali paniculato-racemosa ad 80 cm. longa et lata, pedunculo,
rhachide, et ramulis copiose et conspicue hirsutis (pilis brunneis e
basi crassa 0.1-0.15 mm. diametro tereti-filiformibus multicellularibus
superne obtortis, pedunculo ad 9 mm. rhachide 3-6 mm. ramulisque
2-3 mm. longis), pedunculo 4-16 cm. longo bracteas 1 vel 2 sub-
coriaceas ad 4 cm. longas saepe gerente, rhachide tereti 2-3 mm.
diametro, ramulis secundariis 12-25 gracilibus sub anthesi 17-40 cm.
longis; bracteis sub ramulis papyraceis lanceolatis 25-35 mm. longis
basi 4-6 mm. latis apice subulatis multinerviis, extus copiose hirsutulis,
intus glabris vel parce tomentellis; umbellulis numerosis 2-15 mm.
distantibus, bracteis sub umbellis 5-102-2.5 mm. eis majoribus
subsimilibus; umbellularum pedunculis eracilibus 3-10 mm. longis
parce hirtellis vel glabratis infra medium 1- vel 2-bracteolatis et
apice pluribracteolatis (bracteolis lanceolatis 0.5-3 mm. longis
acutis parce hispidulo-tomen tellis) ; floribus 5—10 per umbellulam glabris
vel interdum calycis lobis et petalis inconspicue setulosis, pedicellis
sub anthesi 1-4 mm. longis; calyce cupuliformi ca. 1 mm. longo et
9 mm. diametro 5-dentato, dentibus:deltoideis acutis 0.3-0.4 mm.
longis; petalis 5 deltoideo-ovatis 1.4-1.6 mm. longis 1-1.2 mm. latis,
apice acutis et cucullatis, intus obscure carinatis; staminibus 5,
filamentis gracilibus 0.5-0.7 mm. longis, antheris ellipsoideis 0.6-0.8
mm. longis apice rotundatis, thecis basi liberis; ovario apice subplano,
stylis 5 liberis minutis sub anthesi 0.1—-0.2 mm. longis, ovulis solitariis
pendulis.

Type in the U.S. National Herbarium, Nos. 2191526 and 2191527, collected
in dense forest in the hills east of the Wainikoroiluva River, near Namuamua,
Namosi Province, Viti Levu, Fiji, alt. 50-200 m., October 15, 1953, by A. C.
Smith (No. 8908). Duplicates at Bish, ete.

ADDITIONAL SPECIMENS EXAMINED:

FIJI: Vitt Levu: Serua: Hills between Navua River and Wainiyavu Creek,
near Namuamua, in dense forest, alt. 100-200 m., Smith 8983 (Bish, US).

88 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Field notes indicate that the plant is a slender tree to 5 or 6 m.
high; the inflorescence is lateral near the apices of branchlets; the
tomentum of the inflorescence is dull brown; the calyx, petals, and
filaments are greenish white, and the anthers white. A local name
referred to No. 8908 was “sole tangane.”’

Among Fijian species, Schefflera euthytricha is closely allied only
to the abundant S. vitiensis (A. Gray) Seem., but that species lacks
the extraordinary, large, spreading, many-celled hairs that cover
the younger vegetative parts of the new species and persist so con-
spicuously on its inflorescence. The rachis and branches of the
inflorescence of S. vitiensis, if not entirely glabrous, are merely fur-
furaceous-tomentellous with inconspicuous, irregular hairs scarely
0.1 mm. long. Furthermore, the primary bracts of its inflorescence
are subcoriaceous and comparatively small (3-15 mm. long), while
its umbel-subtending bracts are usually only 1-3 (rarely 6) mm. long.

In respect to its remarkable indument, the new species is more
suggestive of S. samoensis (A. Gray) Harms, which has similar veg-
etative hairs, these sometimes also occurring on the inflorescence;
however, such hairs are comparatively sparse and do not exceed
2-3 mm. in length. In size of inflorescence bracts, S. samoensis
resembles S. vitiensis rather than S. euthytricha. In having its
leaflet blades conspicuously and closely apiculate-serrate, and in
having 6 or 7 styles, S. samoensis differs from both the Fijian species
of this alliance.

Myrsinaceae
Tapeinosperma ampliflorum A. C. Sm., sp. nov.

Arbor gracilis 4-10 m. alta inflorescentiis exceptis glabra (sed
partibus novellis parce et obscure glanduloso-puberulis), ramulis
cinereis subteretibus apices versus 4-8 mm. diametro; petiolis crassis
semiteretibus 7-12 mm. longis fere ad basim angulatis, laminis sub-
coriaceis in sicco fuscis immerso-glandulosis obovato-ellipticis, (10-)
12-20 cm. longis, (3.5-) 5-8 em. latis, basi attenuatis et in petiolum
longe decurrentibus, apice obtusis vel rotundatis, margine integris
et anguste recurvatis, costa valida supra subplana subtus prominente,
nervis lateralibus primariis utrinsecus (12-) 15-20 patentibus margines
versus anastomosantibus utrinque inconspicue prominulis, rete
venularum subimmerso; inflorescentiis apices ramulorum versus
lateralibus compacte paniculatis sub anthesi ad 6 cm. longis et 4 cm.
latis (sub fructu ad 9X6 em.), pedunculo (1-3 cm. longo), rhachide
ramulisque lateralibus crassis angulatis ut pedicellis pilosis sed demum
glabratis; floribus 3-6 apices ramulorum brevium (2-10 mm.)
lateralium versus laxe ageregatis, bracteis oblongis obtusis 2-3 mm.
longis margine ut calycis lobis ciliolatis mox caducis, pedicellis crassis

SMITH—PACIFIC ISLAND PLANTS, XVIII 89

(ad 1.5 mm. diametro) subteretibus sub anthesi 1-2 mm. longis
puberulis (pilis fusco-brunneis glandulosis patentibus 0.05-0.1 mm.
longis); calyce cupuliformi sub anthesi 3.5-4 mm. longo 4.5-5 mm.
diametro alte 5-lobato ut pedicellis glanduloso-piloso, lobis late
imbricatis ovatis 2-3 mm. longis 2.5-4 mm. latis apice cuspidatis
margine conspicue ciliolatis (pilis multicellularibus 0.2-0.3 mm. longis
saepe obscure glandulosis); corolla carnosa immerso-nigro-glandulosa
subrotata fere ad basim 5-lobata, lobis late imbricatis ovatis 3-4 mm.
longis latisque obtusis; staminibus corollae faucibus insertis, filamentis
brevibus, antheris deltoideo-ovoideis 1.5-1.8 mm. longis obtusis,
connectivo crasso immerso-glanduloso; ovario sub anthesi conico ca.
1 mm. longo conspicue immerso-glanduloso in stylum aequilongum
attenuato, stigmate truncato, placenta ovoidea apice acuta, ovulis 4;
pedicellis sub fructu ad 5 mm. elongatis, calyce demum rotato 5-7
mm. diametro persistente, fructibus subgloboso-obovoideis 7-9 mm.
diametro apice conico-subacutis basi obtusis, pericarpio in sicco
coriaceo 1-1.5 mm. crasso, semine 1.

Type in the U.S. National Herbarium, No. 2192118, collected in dense forest in
the bills east of the Navua River, near Nukusere, Serua Province, Viti Levu, Fiji,
alt. 100-200 m., October 29, 1953, by A. C. Smith (No. 9102). Duplicates
at Bish, etc.

ADDITIONAL SPECIMENS EXAMINED:

FIJI: Vitt Levu: Namosi: Northern slopes of Korombasambasanga Range,
in drainage of Wainavindrau Creek, in dense forest, alt. 450-600 m., Smith 8743
(Bish, US, etc.). Serua: Vatutavathe, vicinity of Ngaloa, in forest, alt. 0-150 m.,
Degener 15176 (A, Bish, UC, US, etc.).

The cited specimens are from often slender trees 4-10 m. high; the
bracts and flower buds are dull pink and the fruits dull brown.

Gillespie (Bishop Mus. Bull. 74:8. 1930) has noted that the dif-
ferentiation between Tapeinosperma and Discocalyx based on the
number and arrangement of ovules is not satisfactory in the Fijian
species referred to these genera. Maintenance of the two genera
should be based on other characters, but a decision whether to retain
Discocalyx (the later name) requires monographic reconsideration.

Although it has a reduced number of ovules, 7. ampliflorum is
allied to species that have been placed in Tapeinosperma. From T.
clavatum Mez it differs in its compact inflorescence, much larger
flowers, deeply lobed calyx, and closer but basically similar inflores-
cence indument. In flower size, 7. chloranthum A. C. Sm. more
closely approximates the new species, but its inflorescence rachis,
branches, pedicels, and calyx have a comparatively thick and tangled
tomentum (of many-celled bairs 0.2-0.4 mm. long). Another species
of this general relationship, 7. greenwoodii A. C. Sm., differs from the
new species in its ample inflorescence with much smaller flowers.
The four species here mentioned have the ovules either 2, 3, or 4.

90 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Rubiaceae

Squamellaria major A. C. Sm., sp. nov.

Frutex epiphyticus partibus novellis et corolla exceptis glaber,
basi incrassato-tuberosus, tubere ellipsoideo ad 40 cm. longo intus
cuniculis a formicis inhabitato, partibus novellis et stipularum
cicatricibus pilis brunneo-stramineis debilibus ad 1 mm. longis pluri-
cellularibus parce hirtellis, ramulis rugulosis crassis superne 4-8 mm.
diametro, stipulis intrapetiolaribus mox caducis; foliis pro genere
magnis subsessilibus, petiolis crassis complanatis (ad 4 mm. lato)
subnullis vel ad 5 mm. longis, laminis crasso-chartaceis in sicco
fusco-olivaceis oblongo-oblanceolatis, (8-) 12-17 em. longis, (3-)
4-6.5 cm. latis, basi obtusis et subito in petiolum decurrentibus,
apice rotundatis vel eroso-cuspidatis, margine scariosis integris vel
undulatis, costa valida utrinque prominente, nervis secundariis
utrinsecus 5-7 valde adscendentibus margines versus anastomo-
santibus utrinque conspicue elevatis, rete venularum laxo irregulari
utrinque subprominulo; inflorescentiis axillaribus cylindrico-tuber-
culiformibus, rhachide ad 10 mm. longa et 4 mm. diametro videtur
apice 6—10-flora, bracteis sub floribus scariosis late deltoideis obtusis
ad 1X2 mm. mox caducis; floribus subsessilibus, pedicello
subnullo vel sub anthesi 1-2 mm. longo; calyce subecarnoso cupuli-
formi sub anthesi 3-4 mm. longo, basi in pedicellum obscurum an-
gustato, limbo erecto apice 4-5 mm. diametro integro vel obscure
4-denticulato; corolla tenuiter carnosa elongato-tubuloso-clavata
maturitate 35-45 mm. longa, basim versus 3.5-4 mm. diametro,
superne incrassata, extus tubo obscure puberula sed superne et lobis
pilis albidis debilibus 0.1-0.3 mm. longis manifestius hirtella, intus
glabra; corollae tubo 2-4 mm. supra basim squamulis 4 aucto, eis
membranaceis antrorse semiorbicularibus ca. 0.5 mm. longis et 1-1.5
mm. latis, margine pilis patentibus albidis 0.5-1 mm. longis copiose
barbatis; corollae lobis 4 oblongis sub anthesi 7-8 mm. longis 2-2.5
mm. latis subacutis demum reflexis; staminibus 4 corollae faucibus
affixis, filamentis subteretibus 3-5 mm. longis, antheris oblongis
versatilibus 5.5-6 mm. longis et ca. 1.5 mm. latis utrinque obtusis
filamento medium versus affixis, thecis inferne liberis; disco annulari-
pulvinato carnoso ad 0.5 mm. alto et 2 mm. diametro; stylo filiformi
sub anthesi quam corolla paullo longiore, stigmate subpeltato vel
discoideo-cupulari 1.2-1.4 mm. diametro apice depresso margine
membranaceo reflexo, ovario 4-loculari, ovulis solitariis e basi erectis:
fructibus juvenilibus ellipsoideis calycis limbo persistente suberecto
coronatis.

Type in the U.S. National Herbarium, No. 2191043, collected in dense forest

on the slopes of Mt. Manuka, east of Wairiki, Taveuni, Fiji, alt. 300-600 m.,
August 14, 1953, by A. C. Smith (No. 8323). Duplicates at Bish, etc.

SMITH—PACIFIC ISLAND PLANTS, XVIII 9]

Field notes indicate the plant as an epiphytic shrub, with an
ellipsoid tuber up to 40 cm. long and with ant-inhabited canals;
the calyx tube is white, pink-tinged, with the limb green at margin;
the corolla is white, faintly pink-tinged distally within; the filaments
are white and the anthers pinkish; and the style is pale pink.

As noted by Beccari in his remarkable study of myrmecophilous
plants (Malesia 2:1-284. pls. 1-65. 1884-86), the two Fijian species
that comprise his genus Squamellaria (op. cit., 228. 1886) are amply
distinct from Hydnophytum and Myrmecodia in the barbate scales
within the corolla tube and in the entire and very different stigma.
Although Beccari implied that the species of Squamellaria are not
associated with ants, this was due only to the inadequately noted
specimens available to him. In my observation, all three species
of the genus now known have tubers with ant-inhabited canals,
similar to those of Hydnophytum; this is borne out by J. W. Parham
(Plants of the Fiji Islands, 209. 1964).

The new species is at once distinguished from its two congeners
by its much larger leaf blades, with conspicuous ascending secondary
nerves, and by its large flowers. The largest leaf blades noted for
the earlier species do not exceed 12 by 3 cm., nor the largest corollas
3 cm. in length. Both S. imberbis (A. Gray) Bece. and S. wilsonit
(Horne ex Baker) Becc. have leaf blades that are obviously acute at
apex and attenuate at base, and in the latter species the petioles are
slender, obvious, and often more than 3 cm. long.

Ixora calcicola A. C. Sm. Bishop Mus. Bull. 220:251, fig. 13. 1959.

Futanca: In thickets on limestone formation, Smith 1145 (Bish, GH, UC,
US); on lagoon cliff, Smith 1219 (Bish, GH, UC, US). Onaera Npriti: Rocky
sea coast, Bryan 387 (A, Bish).

These Lau specimens were taken from shrubs or small trees 1-4 m.
high, growing near sea level; the corolla is white, the anthers yellow,
and the fruit brown.

Previously I had referred these three specimens to J. samoensis
A. Gray, which they resemble only superficially; that species has the
inflorescence-subtending bracts acute or obtuse to a short-stipitate
base, and the calyx obviously pilose. In comparison with the type
specimen of I. calcicola, from Vava’u, Tonga, the Lau material shows
normal variability, having the petioles sometimes up to 1 cm. long,
the leaf blades up to, 12.57 cm., and the corolla with a tube 18-25
mm. long and with lobes perhaps as small as 8X4 mm., somewhat
smaller than those described. A more recent collection from Vava’u,
Hotta 4936 (Bish), differs from the type collection (Yuncker 16091) to
about the same degree as the Fijian material.

92 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Ixora arestantha A. C. Sm., sp. nov.

Frutex vel arbor 5-12 m. alta, ramulis subteretibus apices versus
2-3 mm. diametro superne pilis pallidis haud 0.1 mm. longis minute
puberulis; stipulis subcoriaceis distinctis dorso puberulis 5-7 mm.
longis basi late deltoideis in aristam subulatam 3-5 mm. longam
angustatis; petiolis saepe inconspicuis interdum manifestis 1-10 mm.
longis canaliculatis mox glabratis; foliorum laminis chartaceis in sicco
fuscis oblongo-lanceolatis, 9-21 em. longis, 2.5-7.5 cm. latis, basi
anguste rotundatis vel late obtusis, in apicem acutum vel mucronulatum
angustatis, supra glabris, subtus (interdum obscure saepe manifeste)
pilis 0.05-0.2 mm. longis puberulis, costa supra paullo elevata subtus
prominente, nervis secundariis utrinsecus 10-15 patentibus inconspicue
anastomosantibus utrinque prominulis vel supra planis, rete venu-
larum inconspicuo laxo utrinque prominulo vel subimmerso; inflore-
scentiis terminalibus trichotome corymbosis e basi divisis sub anthesi
3-6.5 cm. longis et ad 13 em. latis, ramulis gracilibus et pedicellis et
calyce pilis 0.05-0.2 (-0.3) mm. longis copiose puberulis, bracteis
bracteolisque lanceolato-subulatis 0.8-3 mm. longis puberulis; pedi-
cellis gracilibus sub anthesi plerumque 1-3 (-6) mm. longis saepe
superne pluribracteolatis; calyce oblongo-cupuliformi sub anthesi
1.7-3 mm. longo et apice 1.2-2 mm. diametro, limbo suberecto hy-
panthium subaequante conspicue 4-lobato intus obscure strigilloso,
lobis oblongo-deltoideis subacutis 0.6-1.5 mm. longis; disco annulari-
pulvinato ca. 0.5 mm. diametro; corolla extus obscure puberula vel
glabra hypocrateriformi, tubo gracili (0.8-1 mm. diametro) sub anthesi
5-7 mm. longo, lobis 4 patentibus anguste oblongis obtusis 5.5-6 mm.
Jongis 1.5-1.7 mm. latis; staminibus exsertis, filamentis filiformibus
1.5-2 mm. longis, antheris lineari-oblongis 3.5-4 mm. longis utroque
acutis ca. 1 mm, supra basim affixis, thecis inferne liberis: stylo gracili
8-9 mm. longo pilis patentibus ca. 0.1 mm. longis setuloso-puberulo
superne incrassato, stigmatibus divaricatis 1.3-1.8 mm. longis; fructi-
bus subglobosis 6-7 mm, diametro parce puberulis vel elabris calycis
limbo puberulo coronatis.

Type in the U.S. National Herbarium, No. 2191555, collected in dense forest
in the hills east of the Wainikoroiluva River, near Namuamua, Namosi Province,
Viti Levu, Fiji, alt. 50-200 m., October 15, 1953, by A. C. Smith (No. 8941).
Duplicates at Bish, ete.

ADDITIONAL SPECIMENS EXAMINED:

FIJI: Viri Levu: Serua: Vatutavathe, vicinity of Ngaloa, Degener 15199 (A);
hills between Waininggere and Waisese Crecks, between Ngaloa and Wainiyambia,
Smith 9552 (Bish, US, ete.). Namosi: Hills east of Navua River, Greenwood
988 (A). Namosi or Serua: Along stream above waterfall near Namuamua,
Gillespie 3253 (Bish). Naitasiri: Vicinity of Nasinu, Gillespie 3508 (A, Bish,
UC, US), 3592 (A, Bish). Ovanau: Tlills southeast of Mbureta River, Smath
7461 (Bish, US, ete.).

SMITH—PACIFIC ISLAND PLANTS, XVIII 93

Field notes indicate the plant as a shrub or slender tree 5-12 m.
high, occurring at elevations of 50-400 m. in dense forest or dry
forest. Color notes of some of the specimens are contradictory, but
usually the inflorescence branches and calyx are noted as deep pink
to bright red, the corolla tube as greenish to pale pink, and the lobes
as deeper pink or dull red (although for No. 9552 I noted the corolla
limb as white); the fruit is indicated as red.

In the close indument of the lower surfaces of its leaf blades and of
its inflorescence branches and calyx, the new species (of § Pavettopsis)
suggests I. pubifolia A. C. Sm. and J. greenwoodiana A. C. Sm.,
obviously differing from both of these in the rounded or broadly
obtuse bases of its leaf blades and its very short petioles. The
corollas of I. arestantha are substantially smaller than those of J.
pubifolia, and its calyx lobes are longer than those of either of these
species.

A closer relative of the new species may be I. myrtifolia A. C. Sm.,
which has subsessile leaves with the blades rounded or subcordate
at base, but that species has the leaves completely glabrous. Jzora
arestantha further differs from I. myrtifolia in the indument (short
but obvious) of its inflorescence branches and calyx, in its long-
aristate stipules (the awns in I. myrtifolia seldom exceeding 2 mm.
in length), in the filiform bracts and bracteoles of its inflorescence,
and in the obvious and comparatively elongate calyx lobes. The
corolla tube of the new species is short and slender for species of this
alliance, but comparably mature corollas are not yet known for J.
myrtifolia.

Ixora prolixa A. C. Sm., sp. nov.

Frutex vel arbor 4-10 m. alta, partibus vegetativis glabris, ramulis
subteretibus apices versus 2-4 mm. diametro, internodiis distalibus
interdum subcomplanatis et ibi raro obscure puberulis; stipulis
subcoriaceis 2-4 mm. longis basi late deltoideis in aristam lanceolatam
1-1.5 mm. longam angustatis; petiolis crassis canaliculatis 11-27
mm. longis; foliorum laminis chartaceis in sicco fusco-olivaceis
oblongo-ellipticis, (9-) 12-20 cm. longis, (4-) 5-11 cm. latis, basi
rotundatis vel obtusis et in petiolum abrupte decurrentibus, apice
obtusis vel subacutis et minute calloso-apiculatis, costa supra leviter
depressa vel plana subtus prominente, nervis secundariis utrinsecus
9-12 patentibus marginem versus anastomosantibus utrinque promi-
nulis vel supra planis, rete venularum copioso utrinque plerumque
prominulo; inflorescentiis terminalibus trichotome corymbosis amplis
e basi divisis sub anthesi 8-23 em. longis et 11-30 cm. latis, ramulis
eracilibus et pedicellis et calyce pilis 0.05-0.2 mm. longis obscure
vel manifeste puberulis, bracteis bracteolisque inconspicuis deltoideis

94 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

vel lanceolatis acutis 0.5-1 mm. longis mox caducis; pedicellis gracili-
bus plerumque 3-12 mm. longis (interdum brevioribus) apicem versus
obscure bracteolatis; calyce cupuliformi 1.5-2 mm. longo apice 1.2-
1.8 mm. diametro, limbo quam hypanthio breviore intus inconspicue
strigilloso, lobis 4 late ovato-deltoideis 0.2—-0.4 mm. longis rotundatis
vel obtusis; disco minuto annulari; corolla glabra vel extus parcissime
puberula hypocrateriformi, tubo gracili (0.7-1.3 mm. diametro) sub
anthesi 12-20 mm. longo, lobis 4 oblongis obtusis 7-10 mm. longis
2-3 mm. latis; staminibus exsertis, filamentis filiformibus 2—4 mm.
longis, antheris lineari-oblongis 7-8 mm, longis 2-2.5 mm. supra basim
affixis utroque conspicue angustatis, basi et apice mucronulatis,
thecis inferne liberis; stylo gracili exserto breviter setuloso vel sub-
glabro superne incrassato, stigmatibus divaricatis 2-4 mm. longis;
fructibus subglobosis 7-10 mm. diametro calycis limbo inconspicuo
persistente coronatis.

Type in the herbarium of the Bernice P. Bishop Museum, collected in the central
forest of the island of Tuvutha, Fiji, alt. about 100 m., September 11, 1924, by FE.
H. Bryan, Jr. (No. 548).

ADDITIONAL SPECIMENS EXAMINED:

FIJI: Vanua Msatavu: Southern limestone section, on the islet Malatta, alt.
0-100 m., Smith 1435 (Bish, GH, UC, US). Koro: Eastern slope of main ridge,
alt. 300-500 m., Smith 969 (Bish, GH, UC, US). Naav: Slopes of Mt. Ndelaitho,

on northern spur, toward Navukailangi, alt. 350-500 m., Smith 7879 (Bish, US,
etc.).

Field notes indicate the plant as a shrub or tree 5-10 m. high,
growing in forest at elevations of near sea level to 500m. The inflo-
rescence branches and calyx are pink, the corolla white, pinkish at
base, the filaments white, and the fruit red or at length brown. The
specimens from Koro and Ngau, in fruit, are referred here with reser-
vations, as corollas are required definitely to separate this new species
from J. elegans; however, in general facies these collections from Loma-
i-Viti very closely suggest the two Lauan specimens, each of which
bears ample flowering and fruiting inflorescences.

Tn its strictly glabrous leaves, puberulent inflorescence branches and
calyx, and very short calyx limb and lobes, the new species (of
§ Pavettopsis) suggests I. elegans Gillespie and J. decora A .C. Sm.
From J. elegans, I. proliza differs in its wide-spreading, larger, more
open inflorescence, and especially in its larger flowers, of which the
corolla tube is very slender and 12-20 mm. long (rather than 5-6 mm,
as in J. elegans). In foliage the two species are very similar, but the
leaf blades of I. elegans are more gradually narrowed to a predom-
inantly acute apex. The related J. decora differs from J. proliza in its
proportionately narrower leaf blades, more compact inflorescences, and
very obscure inflorescence indument.

SMITH—PACIFIC ISLAND PLANTS, XVIII 95

In the size of its inflorescence and corolla, the new species resembles
I. tubsflora A. C. Sm. (thus far known only from Taveuni), but that
species has obvious, oblong-deltoid calyx lobes, glabrous inflorescence
branches, and stipules with a conspicuous (2-5 mm. long) awn.

Psychotria brevicalyx Fosberg, Sargentia 1:132. 1942.
Psychotria gibbsiae var. velutina Fosberg, Sargentia 1:127. 1942.

In describing this species, Fosberg cited only fruiting material;
he mistook a calycine fragment for an ‘‘exceedingly short 5-denticulate
calyx-ring” and consequently applied to his concept an inappropriate
epithet. Additional material now available indicates that the calyx
limb greatly exceeds the hypanthium at anthesis. The relationship
of the species is not with P. turbinata A. Gray as stated by Fosberg,
but rather with P. pubtflora (A. Gray) Fosberg and its allies. In this
alliance, P. pubiflora (which may include P. crassiflora Fosberg),
P. gibbsiae S. Moore, and P. magnifica Gillespie all have the lower
surface of the leaf blades glabrous. The closest ally of P. brevicalyc
is P. nandarivatensis A. C. Sm., from which it may be distinguished
as follows:

Leaf blades 12-24 4-12 cm., copiously hirtellous beneath with pale or reddish
hairs usually 0.2-0.5 mm. long, these uniformly distributed and persistent;
indument of inflorescence similarly copious and persistent, the hairs 0.5-1.2
mm. long . rn P. brevicalyx

Leaf blades usually smaller, (5-) 7-16X(2-) 3-6 cm., sporadically hirtellous
beneath with reddish hairs 0.5-1 mm. long, these denser on costa and nerves
than on surface, rarely essentially lacking; indument of inflorescence similar,
often sporadic and rarely essentially lacking . . . . . . P. nandarivatensis

While the extremes of these two concepts are quite distinct, some
specimens are confusingly intermediate. Both species are here re-
tained pending a detailed review of the genus in Fiji. Psychotria
nandarivatensis is now known from many upland specimens in northern
and northwestern Viti Levu.

The type of P. gibbsiae var. velutina in foliage and indument pre-
cisely agrees with material here referred to P. brevicalyx, including its
type. The inflorescence of the varietal type is comparatively com-
pact, but its calyx is typical for P. brevicalyz.

The calyx limb of the concept here accepted as P. brevicalyx is
quite variable, but is characterized as composed of a cylindrical lower
portion 5-10 mm. long and a more or less flaring apical portion. The
apex in some specimens is subrotate and 12-15 mm. in diameter, with
broad lobes as large as 68 mm. Other flowering specimens have
the apex only 6-10 mm. in diameter, with oblong lobes 1-51.5-
2.5mm. Since foliage and corolla characters are not correlated with
these differences, and since the calyx limb itself varies substantially
on single plants, I believe this concept of P. brevicalyx to be reasonable

96 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

for Psychotria, in which intraspecific crossing must be locally common-
place. A supplementary description of the inflorescence follows:

Inflorescence branches, pedicels, and calyx copiously hirtellous, the
indument composed of pale, many-celled spreading hairs 0.5-1.2 mm.
long; calyx 10-15 (rarely to 20) mm. long, the hypanthium cupuliform,
about 2 mm. long, the calyx limb ascending in a cylindrical tube
5-10 mm. long and 3.5-4 mm. in diameter, broadening or flaring into
an apical portion 6-15 mm. in diameter, this conspicuously or obscurely
reticulate-veined, pilose without and less obviously so or glabrous
within, the lobes 5, variable, broadly deltoid to oblong, 1-6 mm. long,
1.5-8 mm. broad, obtuse at apex, the sinuses obtuse or subacute;
corolla cylindric, gradually broadened distally, 18-20 mm. long, co-
piously hirtellous without with pale few- or several-celled hairs 0.2-1
mm. long, the tube puberulent or tomentellous within distally, the
lobes oblong, 4-5 mm. long, about 1.5 mm. broad, subacute, obscurely
puberulent within; filaments slender, 2-4 mm. long, appressed to
corolla throat and similarly pilose, the anthers oblong, 2-3 mm. long,
obtuse at both ends; style filiform, about as long as the corolla, the
stigmas recurved, about 1 mm. long.

The species is known from the two large islands of Fiji, at ele-
rations of near sea level to 600 m. (the Gillespie specimen is said to
be from Mt. Naitarandamu at about 1,150 m.). Following are the
specimens referable to it, cited without details except for the island
and province:

Vitt Levu: Nandronga & Navosa: Degener 15256, 15292 (type US; Yawe,
vicinity of Mbelo, near Vatukarasa, alt. 120-300 m.), Tabualewa 15620. Serua:
Tabualewa 15605 (US, type of P. gibbsiae var. velutina: Mbuyombuyo, near

Namboutini), Degener 15154, Smith 8997, 9206, 9656. Namosi: Gillespie 313),
Smith 8726, 8893, 8898. Vawnva Levu: Thakaundrove: Smith 1856.

Psychotria stenantha A. C. Sm., sp. nov.

Frutex vel arbor parva 4-5 m. alta ubique praeter pilos obscuros
inter stipulas et inflorescentiae bracteolis et calycis mareine
glabra; stipulis 6-12 mm. longis inferne connatis superne liberis et
late bifidis; petiolis gracilibus 7-15 mm. longis, foliorum laminis
subcoriaceis in sicco fuscis et copiose immerso-glandulosis, elliptico-
oblongis, 7-10 em. longis, 2.5-5 em. latis, basi acutis, apice obtusis
vel obtuse cuspidatis, margine anguste recurvatis, costa utrinque
prominente, nervis secundariis utrinsecus 10-12 patentibus supra
subplanis subtus elevatis, rete venularum subimmerso; inflorescentin
compacta ad 2 cm. diametro e basi 3- vel 4-divisa, ramulis 2- vel 3-plo
divisis, floribus plerumque trinis, bracteolis sub floribus minutis saepe
crispato-pilosis (pilis 0.2-0.5 mm. longis), pedicellis teretibus 1-2 mm.
longis; calyce cylindrico 3-3.5 mm. longo (hypanthio obconico ad
1 mm. longo incluso), limbo ca. 1.5 mm. diametro apice abrupte sed

SMITH—PACIFIC ISLAND PLANTS, XVIII 97

inconspicue ad 2 mm. patente et ibi pilis brunneis pluricellularibus
0.1-0.2 mm. longis inconspicue ciliolato 4- vel 5-denticulato, dentibus
deltoideis acutis haud 0.3 mm. longis, sinibus rotundatis; corolla
carnosa in alabastro ad 5 mm. longa intus faucibus pilosa, lobis 4 vel
5 obtusis; staminibus 4 vel 5, filamentis in alabastro brevibus, antheris
oblongis 1.7-2 mm. longis utroque obtusis; disco minuto glabro;
stylo gracili apice bifido.

Type in the herbarium of the Arnold Arboretum, collected in thick forest on

the Mt. Evans Range, Mba Province, Viti Levu, Fiji, alt. about 1,050 m., Sep-
tember 24, 1944, by William Greenwood (No. 1062A). Duplicate at Bish.

In its narrow, erect calyx limb obviously exceeding the hypanthium
in length, and in its glabrous habit, P. stenantha is closely related only
to P. vitiensis Fosberg (Calycosia monticola Gillespie) and P. mundula
A.C. Sm. From the former it differs obviously in its comparatively
short calyx limb (2-2.5 mm. long instead of 4.5-7 mm.), and also in
its somewhat larger leaves with more numerous, spreading secondaries.
The corolla of the type collection is too young to permit comparison.
From P. mundula the new species differs in its comparatively open
habit, larger leaves, more open inflorescence, and short calyx lobes.

Psychotria hemisphaerica Gillespie, Bishop Mus. Bull. 91:32, fig. 36. 1932.

In discussing P. carnea (Forst. f.) A. C. Sm. (Bishop Mus. Bull.
141:151. 1936), based on Petesia carnea Forst. f., I placed P. hemi-
sphaerica in its synonymy. Upon further examination of the Fijian
material now available, I believe Gillespie’s concept to merit specific
recognition. It is readily distinguished from P. carnea by having
the distal branches of its inflorescence, its pedicels, bracts, and calyx
puberulent (with pale hairs that are minute but apparent, 0.05—0.1
mm. long). The indument extends to both surfaces of the calyx
limb and is usually, but not always, persistent in fruit. The cor-
responding parts of P. carnea are glabrous.

Gillespie did not cite a type, and this becomes a matter of conse-
quence because the 17 specimens cited by him are, in my opinion,
referable to four species. The following collections represent P.
carnea: Gillespie 2975.1, 3300, Parks 20226, 20265, 20465. Gillespie
3793 represents P. incompta A. C. Sm., and Gillespie 2353 P. pit-
tosporifolia Fosberg.

An exchange of correspondence between Gillespie and Miss Marie
C. Neal (then in charge of the Bishop Museum herbarium) is pertinent
in the designation of a lectotype for P. hemisphaerica. In this,
Gillespie asked Miss Neal to select a specimen most resembling his
figure 35; she has indicated Gillespie 2208, which indeed seems to be
the basis for the habit drawing and the fruit (figs. 6, ¢). I believe
that Gillespie 4592 must have served as the basis for the flower

98 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

drawings (figs. a, d, e). As the lectotype for this concept, Gillespie
2208 in the Bishop Museum herbarium may be accepted.

Psychotria hemisphaerica is known only from the islands of Viti
Levu and Vanua Levu, with a single specimen from Ongea Levu in
Lau, at elevations from near sea level to about 600 m. Following
are the specimens referable to it, cited without details except for the
island and province:

Vitt Levu: Nandronga & Navosa: Smith 4713. Serua: Degener & Ordonez
13622, Tabualewa 15581, Smith 9236, 9363. Namosi: Greenwood 1041. Naitasiri:
Gillespie 2457, 3425, 3467, 3519, Setchell & Parks 15098, 15167, Parks 20069,
Mac Daniels 1146, B. E. Parham 974, Degener & Ordonez 13763. Rewa: Gillespie
2208 (lectotype Bish; southeastern slopes of Mt. Korombamba, alt. 400 m.),
2260, 4592, Parks 20121, 20315, Meebold 16876, 17043. Vanua Levu: Mathuata:
Smith 6420. Thakaundrove: Smith 588. Onaea Levu: Bryan 422. Fiji,
without locality: Horne 705.

Psychotria hemisphaerica is one of a group of five interrelated Fijian
species, the others being P. carnea (which also occurs in Samoa and
Tonga), P. huntert (Horne ex Baker) A. C. Sm., P. archboldiana
Fosberg, and P. ineompta. This group, which scarcely merits my pre-
vious designation as Psychotria sect. Eumachia (DC.) A. C. Sm.
(Bishop Mus. Bull. 141:151. 1936), is characterized as follows:
Calyx limb suberect or erecto-patent, 2-5 mm. long, often thin and
obviously nerved, and fairly persistent in fruit; corolla limb sharply
enlarged and 4-angled or 4-winged just prior to anthesis, the corolla
lobes broad and obviously veined, appressed at their inner margins
to form the angles or wings; stipular and pyrene characters are also
reasonably constant but are duplicated in other species groups, and a
consideration of their value must await revision of all the Fijian Psy-
chotriae. The species of this alliance are not alwayssharply demarcated,
and P. hemisphaerica may be expected to intergrade with P. carnea and
P. archboldiana. Students of Psychotria (among innumerable other
insular genera) must be reconciled to the plastic nature of their con-
cepts, which cannot be discarded merely because occasional individ-
uals demonstrate intermediate characteristics.

Psychotria scitula A. C. Sm., sp. nov.

Arbor parva ut videtur; ramulis gracilibus cinereis glabris inter-
nodiis ultimis paullo complanatis; stipulis in vaginam subcoriaceam
gracilem valde connatis, vagina cylindrica superne gradatim contracta
6-12 mm. longa pilis rubiginosis multicellularibus 0.5-1 mm. longis
hirtella demum glabrata, stipularum apicibus liberis lanceolato-
subulatis inconspicuis 1-2 mm. longis; petiolis gracilibus 10-25 mm.
longis, laminis foliorum practer costam subtus ut stipulis rubiginoso-
hirtellam glabris papyraceis in sicco fuscis, obovato-lanceolatis, 9-16
cm. longis, 3-6 em. latis, basi acutis et in petiolum longe decurrentibus,

SMITH—PACIFIC ISLAND PLANTS, XVIII 99

in acuminem conspicuum gracilem 1-2 cm. longum gradatim angusta-
tis, supra cystolithis linearibus inconspicue ornatis, costa supra
elevata subtus prominente, nervis secundariis utrinsecus 11-16 arcuato-
adscendentibus marginem versus manifeste anastomosantibus supra
subplanis subtus elevatis, rete venularum laxo supra immerso subtus
prominulo; inflorescentia laxa e basi 5-ramulosa, ramulis 3.5-6 cm.
longis gracilibus debilibus glabris superne trichotome divisis, bracteolis
sub floribus oblongo-linearibus 1-3 mm. longis 0.2-1 mm. latis extus
copiose vel sparse strigillosis (pilis ut eis calycis), floribus plerumque
trinis sessilibus; calyce anguste cupuliformi sub anthesi 3-3.5 mm.
longo et apice 2-2.5 mm. diametro extus basi excepta pilis rubiginosis
0.4-1 mm. longis plerumque 5-7-cellularibus copiose hirtello, hypan-
thio parvo obconico, limbo erecto submembranaceo inconspicue
elanduloso-ornato intus glabro 2-2.5 mm. longo subtruncato, dentibus
5 minutis; corolla cylindrica ad 5 mm. longa (immatura) utrinque
glabra vel pilis paucis apicalibus inconspicue ornata, lobis 5 oblongis
1.5-2 mm. longis obtusis; staminibus 5, filamentis brevibus glabris,
antheris oblongis ca. 2 mm. longis utroque obtusis; disco annulari-
pulvinato glabro; stylo in alabastro corollam subaequante apice bifido.

Type in the herbarium of the B. P. Bishop Museum, collected on Mt. Voma,
Namosi Province, Viti Levu, Fiji, alt. 900 m., September 6, 1927, by John W.
Gillespie (No. 2789). Duplicates at GH, UC. Another collection from the same
locality, alt. 800 m., is Gillespie 2672 (A, Bish, GH, UC).

Psychotria scitula is closely allied only to P. filipes A. Gray and P.
diffusiflora A. C. Sm., differing from both in its sessile flowers, the
striking indument of its calyx, and its comparatively elongate calyx
limb. Both of the older species have pedicellate flowers with the
calyx limb glabrous and 0.5-1 mm. long. Psychotria diffusiflora
further has the corolla copiously barbellate within, while that of
P. filipes is essentially glabrous within. Now that more adequate
material is at hand, the difference in leaf shape between P. jfilipes
and P. diffusiflora is less significant than I previously thought, but
in this respect the new species more closely resembles typical P.
diffusiflora. Another species of this relationship, P. pelagica Seem.,
is in my opinion questionably separable from P. filipes.

In connection with the new species, P. vomensis Gillespie should
be mentioned; as the name implies, this also occurs on Mt. Voma.
Both species have the flowers in sessile triads and the calyx copiously
pilose. However, P. vomensis has the inflorescence comparatively
compact, each inflorescence ray be:ng usually unbranched and obvi-
ously pilose; the calyx indument is composed of usually unicellular pale
hairs scarcely 0.1 mm. long; the calyx limb is subspreading and with
obvious lobes; the leaf blades are proportionately broader and obtuse
to obtusely short-cuspidate at apex, and lack the costal indument;

100 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

and the stipules are of quite a different type, being free and deeply
bifid in the distal half.

Psychotria consobrina A. C. Sm., sp. nov.

Arbor gracilis vel frutex 3-5 m. altus, ramulis gracilibus internodiis
distalibus copiose sed minute papilloso-puberulis (pilis unicellulari-
bus) ; stipulis 8-10 mm. longis extus copiose puberulis inferne connatis
superne in lobos 3-5 mm, longos et 0.8 -1.5 mm. latos apice acutos vel
attenuatos bifidis; petiolis gracilibus semiteretibus 1.2-5 em. longis
primo puberulis demum glabratis, laminis foliorum chartaceis in
sicco fuscis ellipticis vel lanceolato-ellipticis, (9-) 10-20 em. longis,
(2-) 4-7.5 cm. latis, basi acutis et in petiolum decurrentibus, apice
acuminatis (acumine ad 1 cm. longo) vel obtuse cuspidatis, obscure
immerso-glandulosis, utrinque glabris, costa supra subplana subtus
prominente, nervis secundariis utrinsecus 12-18 patentibus curvatis
marginem versus anastomosantibus supra subplanis subtus elevatis,
rete venularum intricato utrinque subimmerso vel subtus prominulo;
inflorescentia ampla cymosa 5-12 cm. longa 4-10 em. lata manifeste
pedunculata 3- vel 4-plo divisa, axibus pedicellisque gracilibus copiose
puberulis (pilis pallidis 1-3-cellularibus 0.05-0.1 mm. longis), pedun-
culo 2-7 cm. longo valde reflexo, bracteis lanceolato-subulatis 1.5—3.5
mm. longis utrinque puberulis plerumque trifidis, lobo centrali
longissimo, bracteolis sub floribus similibus sed 1-2 mm. longis;
floribus plerumque trinis, pedicellis sub anthesi 1-5 mm. longis,
calyce et corolla minute sed copiose puberulis (pilis plerumque uni-
cellularibus) ; hypanthio cupuliformi sub anthesi haud 1 mm. longo,
calycis limbo erecto-patente 1.5-2 mm. longo et apice 2.5-3.5 mm.
diametro copiose immerso-glanduloso intus puberulo, lobis 5 semior-
bicularibus 0.5-1 mm. longis et 1.5 mm. latis ; corolla cylindrica sub
anthesi 9-10 (-13) mm. longa superne paullo amplhiata intus medium
versus pilis pallidis multicellularibus 0.5-0.8 mm. longis copiose
tomentello-barbata, lobis 5 oblongis 2-3 mm, longis obtusis intus
cucullatis; staminibus 5, filamentis glabris 1.5-2 mm. longis, antheris
oblongis ca. 2 mm. longis utroque obtusis ; disco annulari-pulvinato
ad 0.8 mm. alto et 1.3 mm. diametro; stylo corollam subaequante
superne papilloso apice bifido; fructibus obo voideo-ellipsoideis demum
glabratis in vivo carnosis in sicco inconspicue angulatis, 8-9 mm.
longis, 5-7 mm. latis, calycis limbo manifeste puberulo coronatis,
pyrenis semi-obovoideis 6—7><4-6 X2-3 mm., ventre complanatis,
dorso obtuse et inconspicue 3- vel 5-carinatis.

Type in the herbarium of the B. P. Bishop Museum, collected in limestone

hills near quarry beyond Lami village, Rewa Province, Viti Levu, Fiji, alt. 50 m.,
February 6, 1928, by John W. Gillespie (No, 4596). Duplicates at GH, UC.

SMITH—PACIFIC ISLAND PLANTS, XVIII 101

ADDITIONAL SPECIMENS EXAMINED:

FIJI: Virt Levu: Nandronga & Navosa: Vicinity of Mbelo, near Vatukarasa,
Degener 15292a (A), Tabualewa 15623 (A, Bish, UC, US). Serua: North of
Korovou, St. John 18933 (Bish, GH); hills west of Waivunu Creek, between
Ngaloa and Korovou, Smith 9300 (Bish, US, etc.) ; hills north of Ngaloa, in drain-
age of Waininggere Creek, Smith 9157 (Bish, US, etc.) ; hills between Waininggere
and Waisese Creeks, between Ngaloa and Wainiyambia, Smith 9383 (Bish, US,
ete.). Namosi or Serua: Vicinity of Namuamua, Gillespie 2996 (A, Bish, UC,
US). Ra: Vicinity of Rewasa, near Vaileka, Degener 15335 (A, Bish, UC, US),
15380 (A), 15449 (A, US), 15525 (A, US). These specimens occur from near sea
level to about 300 m., in dense wet or dry forest; the corolla is reported as white
and the fruit as red.

The complex of forms related to P. fureans Fosberg is difficult to
understand on the basis of available material, but I believe that my
discussion in Journ. Arnold Arb. 34:105 (1953) requires amending.
The closest relative of P. furcans appears to be P. pritchardit Seem.,
with which it has in common a spreading, obviously pedunculate, and
usually reflexed inflorescence. From P. pritchardn, P. furcans and
the new species here discussed differ in having the stipules compara-
tively small, not auriculate, with the free portion bifid into acute lobes,
and the inflorescence peduncle comparatively short, 2-7 cm. long.
Psychotria pritchardii has extraordinary, auriculate stipules, intercon-
nected by a flange of tisstie proximal of each petiole, the free portion
being bifid into broad, rounded lobes, and the peduncle greatly elon-
cated, 7-35 em. long. On the basis of available specimens, P. prit-
chardii seems limited to Taveuni and Vanua Levu.

The new species, as represented by the specimens cited above,
differs from typical P. furcans in the copious but minute indument of
its young parts, stipules, inflorescence peduncle and branchlets, pedi-
cels, calyx, and corolla; further its calyx limb is longer and more
spreading, with larger, rounded lobes, its pyrenes are obtusely 3- or
5-carinate rather than with a prominent, thin, median dorsal keel,
and its leaf blades are prevailingly elliptic rather than oblong-lanceo-
late. In addition to typical P. furcans, from upland Viti Levu, and
typical P. consobrina, from lowland Viti Levu, there is available a
series of confusing specimens, mostly from upland Viti Levu, that show
relationships to both concepts. Because these specimens have the
puberulent inflorescence parts (including calyx and corolla) and the
comparatively large calyx limb of P. consobrina, I believe this to be
their better position, But they have, in general, the oblong leaf-
blades, glabrous young parts, and glabrous stipules of typical P.
furcans. The calyx tube, in some of these specimens, is as much as
3 mm. long and in shape suggestive of that of P. chrysophylla Fosberg,
a species with a more pronounced inflorescence indument of several-

102 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

celled, longer, reddish hairs. These questionable specimens, now
referred to P. consobrina with reservations, are:

Vit1t Levu: Mba: Gillespie 3695, Parks 20537, 20680, Degener 14369, 14525,
14625, 14637, 14658, 14746, Greenwood 889, 889A, 1163, 1246, 1250, Smith 4091,
4277, 4354, 4372, 5980, 6038, 6270. Nanpronaa & Navosa: Smith 5411, 5477,
5608. Moruriki: Seemann 244. Vanua Levu: Mathuata: Smith 6671, 6824.

While a complex of morphological intermediates among these three
species—P. furcans, P. consobrina, and P. chrysophylla—may be ex-
pected to occur, the extremes seem to have a degree of individuality
that makes their recognition as species desirable.

In connection with the above citation of Seemann 244, it should be
noted that this was one of the two collections originally cited by
Seemann as representing his P. broweri (Seem. Fl. Vit. 135. 18686).
These two collections are not conspecific, although both came from the
island of Moturiki. The second, Seemann 254, was inf ormally chosen
as the lectotype by Gillespie, as I indicated in Journ. Arnold Arb.
34:104. 1953. Such a choice is herewith formalized, since it has been
followed in herbaria, where P. broweri is now well represented by
material from several islands of Fiji. It is an essentially glabrous
plant, characterized by having its inflorescence epedunculate, with
3-6 branches from the base; its relationship is with P. platycocea
A. Gray and P. filipes A. Gray.

Psychotria salticola A. C. Sm., sp. nov.

Arbor vel frutex 4-8 m. altus inflorescentia excepta glaber, ramulis
subteretibus internodiis distalibus subcomplanatis; stipulis conspicuis
papyraceis ellipticis, eis inflorescentiam juvenilem obtegentibus ad
3 cm. longis et 1.5 cm. latis in vaginam latam inferne connatis, superne
liberis manifeste nervatis conspicue bifidis, lobis oblongo-deltoideis
4-8 mm. longis latisque apice obtusis; petiolis gracilibus semiteretibus
2-4 cm. longis, laminis foliorum chartaceis in sicco fusco-viridibus
coplose glanduloso-punctatis, anguste ellipticis, (12-) 15-24 em. longis,
(3.5-) 5.5-10 em. latis, basi acutis et in petiolum longe decurrentibus,
apice in acuminem brevem ad 1 cm. longum obtuse cuspidatis, costa
supra elevata subtus prominente, nervis secundariis utrinsecus 11-14
erecto-patentibus arcuatis supra subplanis subtus valde elevatis, rete
venularum subimmerso vel subtus subprominulo; inflorescentia ter-
minali compacta multiflora e basi 3—5-divisa sub anthesi ad 4 em.
longa, ramulis gracilibus 2- vel 3-plo divisis et pedicellis hirtellis
(pilis pallidis vel rubiginosis patentibus 0.1-0.2 mm. longis 2-4-
cellularibus), bracteis bracteolisque inconspicuis oblongo-lanceolatis
0.5-1 mm. longis superne fimbriatis; floribus plerumque trinis, pedi-
cellis gracilibus sub anthesi 2-4 mm. longis interdum glabris; calyce
anguste cupuliformi vel subfusiformi sub anthesi 3.5-5 mm. longo,

SMITH—PACIFIC ISLAND PLANTS, XVIII 103

hypanthio obconico 1-1.5 mm. longo parce hirtello vel glabro, limbo
erecto subcarnoso 2.5-3.5 mm. longo apice 2.5-3 mm. diametro
utrinque glabro margine pilis ut eis pedicelli conspicue fimbriato,
4—6-denticulato vel lobato, lobis minutis vel interdum semiorbiculari-
bus ad 0.5 mm. longis; corolla subcarnosa cylindrico-infundibuliformi
sub anthesi ad 12 mm. longa, extus copiose (vel tantum apice) pilis
0.1-0.3 mm. longis 2—5-cellularibus hirtella, intus medium versus pilis
brevioribus saepe unicellularibus barbellata, lobis 4-6 oblongis
2-2.5 mm. longis obtusis; staminibus 4-6, filamentis minutis, antheris
oblongis 1.6-1.8 mm. longis utroque obtusis; disco annulari-pulvinato
glabro ad 0.8 mm. alto; stylo gracili corollam subaequante apice
bifido; inflorescentiae ramulis sub fructu ad 6 cm. longis et pedicellis
ad 12 mm. longis, fructibus juvenilibus ellipsoideis calycis limbo
erecto coronatis.

Type in the U.S. National Herbarium, Nos. 2191239 and 2191240, collected
in dense forest in the hills bordering Wainavindrau Creek, in the vicinity of
Wainimakutu, Namosi Province, Viti Levu, Fiji, alt. 150-250 m., September 17,
1953, by A. C. Smith (No. 8550). Duplicates at Bish, ete.

ADDITIONAL SPECIMENS EXAMINED:

FIJI: Virt Levu: Namosi: Hills north of Wainavindrau Creek, between
Korombasambasanga Range and Mt. Naitarandamu, alt. 250-450 m., Smith
8491 (Bish, US, etc.).

Field notes indicate the plant as a tree or shrub 4-8 m. high, with
the inflorescence branches, pedicels, and calyx pink or dull pink and
the corolla white. The two collections here referred are not identical,
but they agree so closely that they must be considered conspecific;
the above description is inclusive. The type has the inflorescence
parts obviously pilose, whereas No. 8491 has the inflorescence branches
and pedicels only sparingly so and the corolla (in young bud) glabrous
except at apex; the latter collection also has the calyx limb with
obvious lobes, whereas that of the type is only minutely denticulate.

The new species is most closely allied to P. brackenridgei A. Gray,
a fairly variable species that is known from many collections and
several islands, with which it has in common large, glabrous, prevail-
ingly elliptic leaf blades, bifid stipules with broad, obtuse lobes, an
inflorescence branching from the base, and a fairly obvious inflores-
cence indument composed of several-celled, often reddish hairs.
Psychotria brackenridgei, however, has the calyx limb erecto-patent,
comparatively short (seldom exceeding 1 mm. in length), and copiously
pilose on both sides, whereas in P. saléicola the calyx limb is erect,
longer (2.5-3.5 mm. long), and glabrous except for the fimbriate
margin. In general aspect P. salticola also suggests P. browert
Seem., but that species has a strictly glabrous inflorescence and a
short calyx limb (0.5-0.8 mm. long).

104 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Compositae
Crassocephalum crepidioides (Benth.) S. Moore, Journ. Bot. 50:211. 1912.

Vit1 Levu: Namosi: Northern base of Korombasambasanga Range, in drain-
age of Wainavindrau Creek, Smith 8669 (Bish, US, etc.); hills east of Waini-
koroiluva River, near Namuamua, Smith 9069 (Bish, US, etc.). OvaLav:
Valley of Mbureta and Lovoni Rivers, Smith 7393 (Bish, US, etc.).

The cited specimens are from coarse or succulent herbs up to 1 m.
high, naturalized along streams and on open gravel banks and rocky
shores of rivers at altitudes of 20-400 m.; the florets and stigmas are
dull yellow, distally tinged with bright or reddish orange.

Crassocephalum crepidioides is now widely distributed from Africa
to southern Asia and the Philippines; it extends in the Pacific east-
ward at least to New Guinea and Ponape, but apparently it has not
been previously recorded from Fiji. Belcher (Ann. Mo. Bot. Gard.
43:13. 1956) has commented on the fact that in herbaria this weed
is often confused with Hrechtites hieracifolia (L.) Raf. ex DC.

Index

(Synonyms in italics. Page numbers of principal entries in boldface.)

Aglaia gracilis, 72
Allomorpha ovalifolia, 85
Anplectrum ovalifolium, 85
Astronidium kasiense, 81
pallidiflorum, 80
storckii, 81
Calycosia monticola, 97
Cassine vitiensis, 78
Crassocephalum crepidioides, 104
Dracontomelum pilosum, 78
Dysoxylum gillespieanum, 72
lenticellare, 72
quercifolium, 78
Elaeocarpus chionanthus, 79
milnei, 79, 80
Elaeodendron vitiense, 78
Erechtites hieracifolia, 104
Glochidion atalotrichum, 74
seemannii, 76
Heliconia bihat, 69
paka, 69
Ixora arestantha, 92
calcicola, 91
decora, 94
elegans, 94
greenwoodiana, 93
myrtifolia, 93
prolixa, 93
pubifolia, 93
samoensis, 91
tubiflora, 95
Macaranga caesariata, 73
magna, 74
Mavo, 74
Medinilla amoena, 85
decora, 84
kandavuensis, 84
ovalifolia, 85
parvifolia, 85
rhodochlaena, 85
spectabilis, 82
subviridis, 82
waterhousei, 83
Ndavo, 74

Paka, 70

Petesia carnea, 97

Pleiogynium hapalum, 76
papuanum, 78
solandri, 77
Timoriense, 77

Polyscias culminicola, 85
joskei, 86

Psychotria § Eumachia, 98
archboldiana, 98
brackenridgei, 103
brevicalyx, 95
broweri, 102, 103
carnea, 97, 98
chrysophylla, 101, 102
consobrina, 100
crassiflora, 95
diffusiflora, 99
filipes, 99, 102
furcans, 101, 102
gibbsiae, 95

var. velutina, 95

hemisphaerica, 97
hunteri, 98
incompta, 97, 98
magnifica, 95
mundula, 97
nandarivatensis, 95
pelagica, 99
pittosporifolia, 97
platycocca, 102
pritchardii, 101
pubiflora, 95
salticola, 102
scitula, 98
stenantha, 96
turbinata, 95
vitiensis, 97
vomensis, 99

Pullea perryana, 71

Sawira, 86

Schefflera euthytricha, 86
samoensis, 88
vitiensis, 88

106 INDEX

Sole tangane, 88 Tapeinosperma ampliflorum, 88
Squamellaria imberbis, 91 chloranthum, 89

major, 90 clavatum, 89

wilsonii, 91 greenwoodii, 89
Tangimauthia, 83 Vava ni Viti, 70

\ am U.S. GOVERNMENT FRINTING OFFICE: 1967

BULLETIN OF THE UNITED STATES NATIONAL MUSEUM

CONTRIBUTIONS FROM THE UNITED StraTEs NATIONAL HERBARIUM

VoLuME 37, Parr 4

THE GENUS CYRTANDRA IN FIJI

By Grorce W. GILLETT

SMITHSONIAN INSTITUTION PRESS e WASHINGTON, D.C, e 1967

THE GENUS CYRTANDRA IN FIJI!
GreorcE W. GILLETT

Introduction

The present study is a limited revision of the genus Cyrtandra in
Fiji and is related to the series initiated by A.C. Smith on selected
families of phanerogams from Fiji and the adjacent archipelagos.
The earlier work on Cyrtandra in Fiji (Seemann, 1861, 1866; Gray,
1862; Clarke, 1883; Gillespie, 1930; Smith, 1936, 1942, 1953) neces-
sarily emphasized exploration and the description of new species. This
preliminary phase has included well over 200 collections and the
description of a total of 45 species. The present work emphasizes the
review of previously known species, the amplification of early descrip-
tions, and a comparative study to provide for the delineation of infra-
generic groups and the identification of species.

The importance of Cyrtandra in Fiji was emphasized by Smith
(1953) who noted that it was exceeded in number of species only by
Psychotria. The wide distribution and frequent occurrence of this
Malaysian-Pacific genus make it an important element in several other
floras, including that of Polynesia. Therefore, the present study in-
cluded a review of all Polynesian species and an investigation of re-
lationships between the Cyrtandrae of Fiji and those of Polynesia.

In number of species (ca. 600) Cyrtandra is the largest genus in the
Gesneriaceae, and among the families of Engler’s Tubiflorae (in-
cluding the Polemoniales, Lamiales, and Scrophulariales) it is ex-
ceeded in number of species only by the genus Solanum. Its territory
extends from Malaya to Eastern Polynesia, with centers of diversity
in Borneo and New Guinea. B. L. Burtt (written communication)
estimates the number of species to be between 130 and 150 for each of
these centers. The great diversity and the large number of species in
the genus impose formidable demands on the taxonomist who aspires
to interpret its infrageneric taxonomy. Notable among those who have

1This paper is based on research partially supported by National Science
Foundation Grants GB 3336, George W. Gillett, principal investigator, Uni-
versity of California, Riverside, and GB 2074, A. C. Smith, principal investigator,
Department of Botany, University of Hawaii, Honolulu.

107

108 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

attempted this was C. B. Clarke who, in 1888, described 2 subgenera
and 13 sections based on a study of 167 species. In 1888, Hillebrand
described five sections, these embracing only the species indigenous
to the Hawaiian Islands. Schlechter (1923) published a treatment of
the Cyrtandra of New Guinea in which he recognized 96 species,
dividing these among two subgenera and 14 sections. Unlike his
predecessors, Schlechter provided a key to his sections. His treatment
probably has considerable merit, yet it pertains to only a portion of
the genus, and surely excludes significant evolutionary lines among
the hundreds of species that do not occur in New Guinea. It is be-
cause of the incomplete, questionable status of the present infrageneric
taxonomy that I have not followed any one interpretation but instead
have designated provisional species groups that appear as homo-
geneous, natural units. None of these are formally described and
named because certain groups include species outside of Fiji, and these
will require revisions as future work is accomplished. Through suc-
cessive regional treatments of this type, perhaps the infrageneric
relationships can gradually be brought into focus and the principal
evolutionary stocks described and named with the minimum of am-
biguity and confusion.

Smith (1955) has shown that many genera in the flora of Fiji have
their maximum speciation in territory to the west, principally in New
Guinea, with a poorer representation in Fiji. This attenuated phyto-
geographic pattern is expressed by Cyrtandra, which has approximately
250-300 species in New Guinea and Borneo but only 35 species in Fiji
by the present treatment. Further exploration will surely increase the
number of species in Fiji, but the possibility of the adjusted figure
being more than a relatively small fraction of the total for New Guinea
and Borneo seems most unlikely. The examination of extensive her-
barium material representing all of Polynesia, but of only limited
material from New Guinea, Borneo, and the Philippines, suggests that
the Fijian and Polynesian species combined portray only a subsidiary
portion of the total morphological expression for the genus.

A short summary of the phytogeographical relationships of the six
species groups that are recognized in this paper is in order. Each of
these is further discussed under the particular species. Species Group 1
(the Cyrtandra vitiensis group) extends from Fiji to Samoa and Hawail.
Species Group 2 (the C. chippendaler group) extends from Fiji to
Samoa, southern Polynesia, and Hawaii. Species Group 3 (the C.
kandavuensis-C. attenuata group) extends eastward from Fiji, occurring
throughout most of Polynesia. Species Group 4 (the C. denhamia
group) extends eastward from Fiji to Samoa and the Marquesas
Islands but is not known in Hawaii. Neither Group 5 (the C. harveyr
group) nor Group 6 ( the C. prattit group) are known to occur east of

GILLETT—CYRTANDRA IN FIJI 109

Fiji. However, both Group 5 and Group 6 have closely related species
in New Guinea, verified by the comparison of herbarium material.

In his recent monograph of Cyrtandra on Oahu, Hawaiian Islands,
St. John (1966) recognizes a total of 133 species, an amazing con-
centration for an island of only 600 square miles, of which only a
small portion is Cyrtandra territory. However, a herbarium and field
study of the Oahu species and of the other Hawaiian species reveals
that they are relatively homogencous in comparison with the Fijian
Cyrtandrae. Using the key to the six species groups in Fiji, all of the
Hawaiian species would be accommodated in Group 1, Group 2, and
Group 3. Remarkable similarities exist between Group 2 (the C.
chippendalei group) and subgenus Brachycyathus in St. John’s mono-
oraph. However, the former embraces only 8 species for all of Fiji,
while the latter includes 4 sections and 89 species for the island of
Oahu.

Habitat Relationships

Guppy (1906) indicated that the Cyrtandrae of the Pacific Islands
are most frequent in moist woods among rank vegetation, but he also
stated that they may occur in more exposed and drier habitats. Field
observations that I made in Java, Borneo, and Hawaii indicate that
Cyrtandra invariably grows as an understory element in dense rain
forest, often in ravines and gorges characterized by high humidity,
very low light intensities, and an almost constant moisture supply.
In many species the root systems are shallow, often clinging to the
surfaces of moist rocks. These habitat characteristics pertain to
elevations between 50 and 2,000 meters. It is known that some species
of Cyrtandra are capable of surviving intermittent droughts, but in
the Pacific the genus is in no instance adapted to arid or semiarid,
exposed habitats. The basic relationship is with upland rain forests,
and the genus occurs only on the islands of the Pacific where sub-
stantial areas of upland rain forest are preserved. It is not an element
of the strand flora, and it does not occur on “high” islands that have a
low rainfall. Examples of the latter include Niihau, in the Hawaiian
Islands, from which no Oyrtandra collections are known, even though
populations abound on the island of Kauai, scarcely 20 miles distant.
It is not likely that Cyrtandra could have established founder popula-
tions on Pacific islands prior to the development of a relatively dense
overstory vegetation. Considering the large number of species in the
genus and its extensive geographical range, it would appear to have an
unusually narrow habitat relationship. The habitats in Hawaii seem
remarkably uniform in character to the point of making it very dif-
ficult to cite differences that could account for the presence of a given
species in one area and its exclusion from another.

110 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM
Pollination and Breeding System

The consideration of the breeding system and its relationship to
variability and the delineation of species should precede a taxonomic
study, if possible. This information provides helpful guidelines for the
interpretation of variability and morphological differences within and
between populations. The types of variation patterns to be expected
from different breeding systems have been presented by Baker (1959),
whose interpretations are followed in this study. It has not been possi-
ble to carry out a field study of Cyrtandra in Fiji, but this has been
accomplished on populations of closely related species in Hawaii—
plants that have the same basic floral morphology, and the white,
fleshy corollas that characterize at least three-fourths of the species
known in Fiji. The observations cited below have been made by me on
several Hawaiian species occurring on Kauai, Oahu, Lanai, Molokai,
Maui, and Hawaii, and together they present a consistent pattern of
floral ontogeny and pollination,

The flowers of Cyrtandra are protandrous, and there appears to be
a conspicuous provision for outcrossing. This is promoted not only by
the earlier maturity of the anthers but also through the positioning
of the anthers and stigma to minimize self-pollination. The effective-
ness of this system is indicated by the complete absence of fruit in 116
bagged flowers of the Oahu species, Cyrtandra garnotiana Gaud., and
by the production of fruit in 39 out of 42 unbagged controls on the
same plants. Self-incompatibility is not known in the Gesneriaceae,
and the above evidence is most inadequate for postulating its occur-
rence in Cyrtandra. Therefore, a measure of inbreeding remains a
possibility inasmuch as the complete exclusion of entomophilous
self-pollination would seem unlikely.

At early anthesis, the filaments extend forward in the corolla tube
and hold the basifixed, apically coherent anthers with the connectives
against the wall of the tube and the thecae face-to-face on the mid-
plane of the tube. The anthers are then parted from below and are
spread so that the thecae lie in one plane. Simultaneously, the anthers
are elevated so that the connectives are appressed to the tube opposite
the upper sinus. The thecae now occupy a position directly over the
access to the nectiferous, cupulate annular disc at the base of the tube.
At this time the immature lobes of the stigma are closely appressed.
In later anthesis the anthers are shifted in position, being lowered to
the anterior surface of the tube, the thecae appressed to the tube
opposite the lower lobe, so that any residual pollen is isolated from
contact with the mature stigma. The stigmatic lobes now expand
and separate, the cleft of separation usually extending farther down on
the anterior side of the style, so that the lobes very often form a

GILLETT—CYRTANDRA IN FIJI 111

cordate receptive surface. The extended stigma is now held over the
access to the nectary, near the position formerly occupied by the
thecae of the anthers. It is clear that two patterns of configuration
are displayed by a given flower and that two insect visits are neces-
sary for accomplishing cross-pollination. Actual pollination has not
been observed, but it is assumed that this is accomplished by night-
flying Lepidoptera attracted to the white corollas and inflorescences.
A lepidopterous antenna has been recovered from one flower. These
observations confirm earlier reports of protandry in Cyrtandra by
St. John (1966) and Carlquist (1965, 1966).

In at least one Fijian species, Cyrtandra harveyi, the outbreeding
system is further specialized through the gynodioecious habit, and
unisexual flowers are known to occur in several additional species in
New Guinea and the Pacific.

It seems clear that in Pacific species of Cyrtandra there is a very
strong trend to outcrossing which in turn would promote variability
involving many genotypes (Baker, 1959). The outcrossing mechanism
would be expected to facilitate interspecific cross-pollinations since
flowers of different species often are remarkably similar, and two,
three, or more species very often occur in the same relatively narrow
habitat. One putative interspecific hybrid, between C. milnet and
C. vitiensis, has been distinguished in the present study.

The outcrossing breeding system provides for the retention of
temporarily disadvantageous genes, including new mutations, through
their inclusion in heterozygous genotypes. Such a breeding system
would thus tend to enlarge the genetic resources, providing for flexi-
bility. It is significant that this system is, in Cyrtandra, correlated
with the propensity of the genus to establish populations on isolated
island habitats. This breeding system, through its provision of maxi-
mum genetic resources for the immigrants and of maximum flexibility
for the founder population, is probably of considerable significance
in the successful establishment of populations on island habitats.

The recombination potential of a species is increased by its capacity
for the dispersal of diaspores (Grant, 1958). In the Pacific species of
Cyrtandra, the fruit is nearly always a conspicuous, succulent berry,
and there seems little reason to doubt the thesis of Guppy (1906) that
these are eaten and dispersed by birds. The seeds are very small,
usually less than 1 mm. in diameter, with perhaps 50 to 100, or more,
per fruit. The seed dispersal would seem to be very effective, so that
a given population could include progeny of neighboring populations
that lie beyond the range of pollen vectors. This would suggest that the
variability of a given species would be a pattern of intergradations
extending over a territory of generous dimensions. However, inter-
island dispersal apparently is not common in Cyrtandra, for a great

112 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

many species are restricted to a single island, and few species are
known to occur beyond a given archipelago (Guppy, 1906). This
restriction is indicated in the Fijian species (fig. 1, table 1).

The extensive geographical territory and impressive speciation of
Cyrtandra suggest that an effective balance is maintained between
dispersal and geographical isolation. The genus has had dispersability
sufficient to accomplish its distribution over the vast expanses of the
western Pacific. However, the high level of speciation suggests that
there have been time intervals between dispersal events, and that
these have been of duration sufficient to provide the geographical
isolation requisite for evolutionary change.

Acknowledgmen ts

The herbarium material of several institutions has been examined
in this study and is here cited, with the indicated abbreviations:
Arnold Arboretum of Harvard University (A); Bernice P. Bishop
Museum (BISH); British Museum (BM); Gray Herbarium of Har-
vard University (GH); Royal Botanic Gardens, Kew (K); New
York Botanical Garden (NY); the University of California (UC);
and U.S. National Herbarium (US), Appreciation is extended to the
administrators of these herbaria for the privilege of examining speci-
mens under their care. Special thanks are extended to Mr. B. L.,.
Burtt, Royal Botanic Garden, Edinburg, who reviewed the manu-
script and communicated many helpful suggestions. Appreciation is
extended to Dr. Harold St. John, Bernice P. Bishop Museum, who
reviewed the manuscript. The encouragement, valuable suggestions,
and numerous reference materials made available to me by Dr. A.
C. Smith, Wilder Professor of Botany, University of Hawaii, are
gratefully acknowledged.

Systematic Treatment
Cyrtandra J. R. & G. Forst. Char. Gen 5. pl. 8. 1776. 2

Herbs, vines, shrubs, or small trees, the stems round to sharply
quadrangular, the trichomes septate, uniseriate, capitate or non-
capitate hairs; leaves whorled, opposite or pseudoalternate by the
reduction of one member of the pair, petiolate to sessile to connate-
perfoliate, the blades simple; inflorescence a congested or elongate
cyme, sessile or pedunculate, axillary on young branches or ecauli-
florous on older stems, sometimes arising among adventitious roots,
usually bracteate, the outer bracts persistent or deciduous, foliaceous
to corolloid and soft-textured, minute to as long as, or longer than,

? Greek kyrtos, curved, and andros, stamens.

GILLETT—CYRTANDRA IN FIJI

113

16%

172s

Liec:

AND NAVOSA

zee ae E 180° W

MATHUATA..5 dSRAMB| +

["t---THAKAUNDROVE

TAVEUNI &
2
|
KORO —
Yroro 5
Oovarau
f ‘eet ¢ sg
Ss S-NAITASIRI,
NANDRONGA (7S as h\ neau 1
~~ SeRUAN OREN
NAMOSI
ae avy =
DISTRIBUTION OF CYRTANDRA
IN THE FlJ!l ISLANDS
a TOTAL SPECIES ON THE ISLAND
|” SPECIES ENDEMIC TO THE ISLAND
10, 0 10 20 30 40 50
CHHE— E— —
MILES
18 toe E 180° W

~/6—

7°

~}§e4

194

-20° 4
s

the inflorescence, free or connate and cyathiferous, the inner bracts
like the outer but smaller; flowers perfect, often protandrous, some-
times unisexual, the calyx foliaceous or corolloid, persistent or decidu-
ous, spathaceous to variously cleft from summit nearly to base into 2 to
5 equal or unequal, valvate lobes; corolla fleshy to chartaceous, usually
white, but also green, yellow, orange, red, pink, or lavender, 1-6 cm.
long, cleft to half its length into rounded, equal to unequal, spreading
to erect lobes, these imbricate in bud, the limb often bilabiate, the

114 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

TaBLE 1.—Distribution of Cyrtandra in Fiji, by islands

Viti Vanua
Species Kandavu Levu Ovalau Ngau Koro Levu Taveuni Rambi

acutangula
aloisiana
anthropophagorum
altenuata
cephalophora
chippendalei
chlorantha

ciliata

coleoides
cyathibracteata
denhamit x
dolichocar pa

esothriz

harveyi

hornet

tnvolucrata

jugalis

kandavuensis x
leucantha

milnet >
moniana

mulliseptata

muskarimba

occulta

pratta

pritchardit

reticulata

spathacea x
taviunensis

tempestit

trichophylla

ventricosa

vicloriae

vitiensts

zanthantha

xxx x
x

xxK KKK KX
x
*
x

x MK KKK KKK KKK XK
x
x

x xX

xxx -&X

Total species: 35, all endemic to the Fiji Islands; 22 species are restricted to a
single island; 8 species occur on 2 islands; 5 species occur on 3 islands.

lower or anterior lobe usually equal to or larger than the others;stamens
2, the filaments adnate to the tube opposite the sinuses of the anterior
corolla lobe, the lower portion of each filament variously twisted, the
anthers coherent at apices; staminodes 2 or 3, 0.5-5 mm. long, adnate
to the corolla tube opposite the posterior sinuses, sometimes bearing
small, vestigial anthers; base of the ovary clasped by a nectiferous,
annular, cupulate, or one-sided disc, this often persistent in mature
fruit, the ovary and style glabrous or variously pubescent, the stigma

GILLETT—CYRTANDRA IN FIJI 115

capitate to applanate and bilobed, the plane of the expanded lobes
perpendicular, oblique, or parallel to the axis of the style; fruit white,
green, yellow, orange, red, or pink, a succulent to coriaceous berry,
ovoid to oblong, up to 4 cm. long, or cylindrical and up to 12.5 cm.
long, often beaked by the persistent style base; seeds many, usually
less than 1 mm. long, usually embedded in the fleshy placentae.

Perhaps 600 species, distributed in rain forest habitats from the
Nicobar Islands to the Malay Peninsula of the Asian mainland;
through the Philippine Islands to Botel Tobago Island southeast of
Formosa, and the Ryukyu Islands; throughout the Malay Archipel-
ago, southeast to Queensland and the Loyalty Islands; east on the
high islands of the Pacific to the Hawaiian Islands and the Marquesas
Islands. The genus is not known on Formosa, Yap, the Marianas
Islands, New Caledonia, or the Tuamotu Archipelago.

Key to Species Groups

Inflorescence capitate to subcapitate, the pedicels concealed by the congested
flowers and bracts. .... . . ... . Group 1
Inflorescence a branching cyme, not capitate or - subcapitate; pedicels visible, not
obscured by congested flowers and bracts.
Calyx persistent, not abscissing from the maturing fruit ... . . Group 2
Calyx deciduous, abscissing at or shortly after anthesis.
Flowers not borne on an elongate, woody, branchlike inflorescence axis.
Styles and anthers restricted to the corolla tube, the filaments 1-4 mm.
long, rarely 5 mm. long.
Calyx lobes about as long as the calyx tube . . . . . . . . Group 3
Calyx lobes 2 to 5 times as long as the calyx tube . . . . . Group 4
Styles, or anthers, exserted above the corolla tube, displayed in or above
the broad corolla throat; filaments 8-12 mm. tong flowers sometimes

unisexual ..... eee ee . . . . « Group 5
Flowers borne on a woody, branchlike inflorescence axis up to 20 cm. long,
bearing numerous pedicels ca. 0.6 em. long. . . ... =. . Group 6
GROUP 1

Outer inflorescence bracts fused into a cup-shaped involucre.
Leaf blades glabrous; calyx and bracts glabrous on the abaxial surfaces.
1. C. cyathibracteata
Leaf blades, calyx, and bracts sericeous on the abaxial surfaces . .2. C. occulta
Outer inflorescence bracts free; inflorescence without a cup-shaped involucre.
Inflorescence cylindrical; peduncle 10-15 mm. long; calyx and bracts with
indument of dark brown, rigid, glistening hairs . . . . 3. C. cephalophora
Inflorescence spreading, not cylindrical; peduncle 2-10 mm. long; calyx and
bracts glabrous or with light brown, crumpled, capitate and noncapitate

hairs.
Outer surfaces of calyx and bracts glabrous to gabrate; bracts lobed; petioles
glabrate, winged to base .. . .... . 4 C. vitiensis

Outer surfaces of calyx and bracts moderately to densely pubescent with
capitate and noncapitate hairs; bracts not lobed; petioles conspicuously
pubescent, rarely winged.

Hairs of the petiole shorter than the petiole diameter . . 5. C. leucantha

116 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Hairs of the petiole longer than the petiole diameter.
Inner surface of corolla glabrous; corolla green . . . 6. C. chlorantha
Inner surface of corolla pubescent; corolla green to white . 7. C. milnei

GROUP 2

Fruit long-attenuate, narrow-cylindrical, scarcely tapered to the style, up to 5.5
em. long; corollaup to 5.5 em.long .. . . . . . & C. dolichocarpa
Fruit fusiform to ellipsoid to ovoid, conspicuously tapered to the base of the
style, up to 2.5 cm. long; corolla up to 3.5 em. long.
Calyx utriculose to ventricose, concealing the fruit, the lobes erect.
Leaf blades pubescent above, the under surface with tertiary veins concealed
by pubescence; branches rounded, not sharply quadrangular.
Hairs of petioles about equal to petiole diameter; outer surface of corolla
pubescent; fruit fusiform . . . ... . . 9 C. multiseptata
Hairs of petioles 2-8 times as long as petiole diameter; outer surface of
corolla glabrous; fruit ovoid... ... 2... 10. C. ventricosa
Leaf blades glabrous above, the under surface with conspicuous tertiary veins
perpendicular to the secondary veins; branches sharply quadrangular.
11. C. acutangula
Calyx campanulate to cylindrical with spreading lobes, the developing fruit
exposed.
Inflorescence and leaves with appressed-sericeous pubescence.
Shrub or small tree up to 6 m. high with coarse, woody branches; peduncles
up toca. 0.6 em. long ... ... . . 12. C. trichophylla
Shrub or herb 1-3 m. high with slender branches: peduncles 1-2 cm. long.
13. C. muskarimba
Inflorescence and leaves with spreading, velutinous pubescence.
Calyx lobes thick, coriaceous; calyx 2.5 to 3.5 cm. long . 14. C. victoriae
Calyx lobes foliaceous, not thick or coriaceous; calyx 1.5 to 2.5 cm. long.
15. C. chippendalei
GROUP 3
Calyx spathaceous with a single cleft nearly to base. . . . . 16. C. spathacea
Calyx not spathaceous, but variously cleft or lacerated.
Peduncle subtending a branching cyme bearing 6 to 10 flowers, or more.
Calyx lobes ovate; outer surface of calyx pubescent . .17. C. kandavuensis
Calyx lobes lanceolate-attenuate; outer surface of calyx glabrous.
18. C. attenuata
Peduncle subtending 1 to 8, rarcly 4, pedicellate flowers.
Pedicels subtended by a pair of connate-perfoliate bracts.
19. C. involucrata
Pedicels subtended by free bracts.
Corolla with dense outer indument of capitate or noneapitate hairs.
Outer surface of corolla with indument of capitate hairs.
20. C. reticulata
Outer surface of corolla with indument of noneapitate hairs.
Mature leaves glabrous above. . . . . ... . 21, C. jugalis
Mature leaves pubescent above. . . . . 22, C. anthropophagorum
Corolla glabrous on outer surface.
Calyx lacerated into unequal segments, the outer surface glabrate, the
inner surface pilose with ascending hairs; leaves pubescent below.
Calyx thick, fleshy, drying to a coriaceous texture, the apices of the
lobes triangular. . . ... 0.0... 0... . 23. C. esothrix

GILLETT—CYRTANDRA IN FIJI 117

Calyx thin, not fleshy and not coriaceous upon drying, the apices of
the lobes lanceolate-attenuate. . ..... . . . 24. C. hornei
Calyx parted into 5 equal lobes, the leaves glabrous below.
Fruit orange to red; inflorescence filiform, with 2, 3, or more pairs of
bracts; style glabrous.
Young stems drying to a flaky surface; bracts lanceolate.
25. C. taviunensis
Young stems drying to a smooth surface; bracts linear.
26. C. montana
Fruit white; inflorescence not filiform, with 1 pair of bracts; style with
indument of capitate hairs. . . . . . . . .27. C. pritchardii

GROUP 4

Calyx lobes large, 1-1.5 em. long.
Peduncles 4-9 cm. long; leaf blades lanceolate to ovate . . . 28. C. denhamii
Peduncles 0.5-1.5 em. long; leaf blades ovate to obovate. . . 29. C. tempestii
Calyx lobes small, ca. 0.5 cm. long.
Outer surface of calyx glabrous, the lobes with rounded apices; peduncles up to
5Bem.long. 2... ee ee ee ee ee ee ee 2 80, C, ciliata
Outer surface of calyx with dense, velutinous pubescence, the lobes with acute
apices; peduncles up to 1.5 cm. long.
Young petioles tomentose, the hairs about 0.5 as long as the petiole diameter.
Montane habitats 500-1100 m.. ...... . . .81. C, xanthantha
Young petioles densely villous, the hairs 1-4 times as long as the petiole
diameter. Lowland habitats below 150m... . . . . 32. C, aloisiana

GROUP 5

Corolla with dense outer indument of capitate hairs; mature leaves pubescent
beneath . toe ee ee ee ee ee es . 33. C. harveyi
Corolla glabrous on outer surface; mature leaves glabrous beneath.
34. C. coleoides

GROUP 6

A single species . 2 1 1 1 ee eee ee ee es . 35. C. prattii
1. Cyrtandra cyathibracteata Gillett, sp. nov.

Frutex ad 2 m. altus, partibus novellis indumento pruinoso indutis
mox glabratis, ramulis ad 2 em. diametro; folia opposita, petiolis
alatis 1-4 cm. longis, laminis ovatis vel obovatis ad 40 cm. longis et
17 cm. latis, basi in petiolum alatum inequilateraliter attenuatis,
apice acutis, margine hydathodis ca. 0.5 mm. longis et inter se ca.
1 mm. inconspicue serratis, maturitate utrinque glabris, venis primariis
utrinsecus 7-12 adscendentibus, secundariis subtus inconspicuis;
inflorescentiae capitatae, pedunculo crasso ca. 1 em. longo apice
bracteis caducis ornato, bracteis extimis in involucrum cyathiforme
1-2 cm. longum connatis ut bracteis interioribus ovatis extus glabris
intus pilosis (pilis patentibus ca, 254 diametro ad 2 mm. longis),
pedicellis crassis 1-2 cm. longis; calyx carnosus ca. 12 mm. longus
caducus in lobis haud 1 mm. longis fissus, sinibus sub anthesi ir-
regulariter laceratis, tubo extus glabro intus piloso,; corolla alba

118 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

carnosa ad 2 cm. longa in lobis subaequalibus rotundatis ca. 0.2
fissa extus indumento eglanduloso ornata intus glabra; staminia
tubo corollae medium versus affixa, filamentis ca. 3 mm. longis,
antheris in apice adhaerentibus ca. 2 mm. infra fauces corollae;
staminodia 2, ca. 0.5 mm. longa infra filamenta 3 mm. et sinibus
lateralibus oppositis disposita; discus annulari-cupuliformis prominens
ca. 1.5 mm. altus deciduus; gynoecium ca. 12 mm. longum, ovario
glabro, stylo indumento noncapitato ornato, stigmate bilobato pelvi-
formi; fructus albi ovoidei ca. 8 mm. longi apice styli basi 1-2 mm.
longo apiculati.

Type in the herbarium of the B. P. Bishop Museum, collected on
bank of creek in Wainimala Valley, near Matawailevu, Waimano
Creek, Province of Naitasiri, Viti Levu, Fiji, alt. about 500 m.,
Aug. 3, 1937, by H. St. John (18191). Duplicate at US.

Distripution: Known only from the type specimen.

This species is closely related to Cyrtandra occulta and to C. vitiensis,
differing from both in its glabrous foliage. It appears to be sympatric
with the latter species. Further comments on the relationships in
this group are included under C. occulta.

2. Cyrtandra occulta A. C. Smith, Journ. Arnold Arb. 34: 39. 1953.3

Erect shrub 2-3 m. high with densely sericeous pubescence of
septate, uniseriate, noncapitate hairs ca. 20u in diameter and up to 7
mm. long, the foliage and inflorescences subcoriaceous when dry;
leaves opposite, the petioles 1-7 cm. long, winged, the blades ovate to
oblanceolate, up to 40 cm. long and 18 cm. broad, tapering to the
petiole acute to rounded at apex, undulate-crenate, glabrous above and
densely sericeous beneath, the pubescence obscuring the secondary
veins; inflorescences densely capitate, on stout axillary peduncles u p
to 3.5 cm. long, terminated by matted-sericeous, connate bracts
forming a cuplike involucre up to 3 cm. long subtending a bracteolate,
branched, many-flowered cyme 5-10 mm. long, the bracteoles greenish
white, ovate-acuminate, up to 3 cm. long, with matted-sericeous
pubescence on both surfaces; calyx greenish white, up to 2.5 em. long,
cleft 0.5 to 0.66 of its length into lanceolate-acuminate lobes with
dense-sericeous pubescence on both surfaces; corolla white, cylindrical
up to 2 cm. long, cleft ca. 0.25 of its length into unequal, rounded
lobes with dense-sericeous outer pubescence and inner pubescence of
very short, capitate hairs; filaments 3-4 mm. long, bearing apically
coherent anthers 2-3 mm. below the corolla sinuses; staminodes 2,

* This and most of the other species of Cyrtandra are mentioned in J. W. Parham,
Plants of the Fiji Islands (1964); most of the older species were mentioned in
llorne, A Year in Fiji (1881), and by C. B. Clarke, in DC. Monogr. Phan., vol. 5
(1888).

GILLETT—CYRTANDRA IN FIJI 119

adnate 8-10 mm. below the upper sinuses and near the middle of the
corolla tube; cupulate annular disc conspicuous at anthesis, ca. 1 mm.
high, deciduous from the maturing fruit; ovary and style ca. 12 mm.
long, the style with indument of capitate hairs on the upper 0.66 of
its length, separating 1-2 mm. above the apex of the mature ovary,
the stigma applanate, bilobed, the lobes spreading from an oblique
terminal cleft; fruit white, ovoid, ca. 1.6 cm. long and 1 cm. broad.

Type LOcALITy: Western slopes of Mount Tomanivi, Viti Levu,
Fiji. Type collected by A. C. Smith, cited below.

Distrrisution: Dense forests of Viti Levu, at altitudes of 250-
1300 m.

Virt Levu: Mba: Mount Tomanivi, Smith 5913 (A-type, BISH, US), Parks
20841 (BISH, UC, US). Namosi: Hills north of Wainavindrau Creek, Smith 8507
(BISH, US). Naitasiri: Rairaimatuku Plateau, Smith 5792 (A. US).

The present species and the related Cyrtandra cyathibracteata are
distinguished from other members of this group by the cuplike in-
volucre formed by the fusion of the two outer bracts of the inflores-
cence. This character has been observed in C. godeffroyi Reinecke
(Samoa), C. burbidgei C. B. Clarke (Borneo), and C. cumingit C. B.
Clarke (Philippine Islands). In C. burbidgei, the white involucre is
certainly a most conspicuous feature, and it probably is of significance
in attracting pollinators.

All of the species in Group 1 are characterized by bracts, calyces,
and corollas that gradually rot and fall from the inflorescences, ex-
posing the white fruits.

3. Cyrtandra cephalophora Gillespie, Bishop Mus. Bull. 74: 21, fig. 26. 1930.

Erect shrub 2-3 m. high, the young foliage glabrate, the stems up
to 1.5 cm. in diameter, quadrangular, the pith up to 1 cm. in diameter;
leaves opposite, the petioles 6-9 cm. long, the blades lanceolate to
lanceolate-ovate to ovate, up to 24 cm. long and 12 cm. broad, equi-
lateral and cuneate at base, acute to rounded at apex, entire to faintly
serrate, at maturity glabrous above, glabrate beneath with scattered,
noncapitate hairs ca. 254 in diameter and 0.3 mm. long, the lower
surface with 5-7 strongly ascending, raised primary veins on each side;
inflorescences capitate-cylindrical, on stout peduncles 1-1.5 em. long,
the flowers partially concealed by conspicuous, persistent, ovate to
orbicular subtending bracts 5-10 mm. long, the bracts with rigid,
glistening, dark brown, noncapitate hairs ca. 30y in diameter and 0.5-1
mm. long without, with similar but much larger hairs 1-3 mm. long
within; calyx persistent, ca. 10 mm. long, cleft into equal, lanceolate-
attenuate lobes ca. 4 mm. long, glabrate without with small, non-
capitate hairs, pilose within with similar, ascending hairs to 3 mm.
long; corolla white, 12 mm. long, cleft 0.33 of its length into rounded,

120 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

unequal lobes, glabrous on both surfaces; filaments 2-3 mm. long,
bearing apically coherent anthers 1-2 mm. below the corolla limb;
staminodes 2, clavate, ca. 1.5 mm. long, adnate ca. 1 mm. below the
limb; cupulate annular disc prominent, ca. 1 mm. high, persistent;
ovary and style ca. 8 mm. long, glabrous, the style curved toward the
corolla throat and bearing an applanate, bilobed stigma, the lobes
thick, ovate; fruit orange, ovoid, ca. 7 mm. long, terminated by the
persistent style base.

Tyre uocauity: Namosi, Viti Levu, Fiji; type collected by Gil-
lespie, cited below.

Distripution: Namosi, Naitasiri, and Rewa, Viti Levu; Thakaun-
drove, Vanua Levu, in rain forest habitats between 150 and 1,200
meters.

Vitt Levu: Namosi: Naitarandamu Mountain, just below summit, Gillespie
3121 (BISH-type, NY, UC). Naitasiri: Tamavua Woods, 74 miles from Suva,

Gillespie 2416 (BISH). Rewa: Mount Kombalevu, Parks 20924, (BISH, UC).
Vanua Levu: Thakaundrove: Mount Mariko, Smith 463 (BISH).

The present species is one of the most easily recognized of F ijian
Cyrtandrae by its conspicuously bracteate, capitate-cylindrical in-
florescence and the peculiar indument of the inflorescence. The clavate,
conspicuous staminodia borne near the summit of the corolla tube are
not known to occur in other Fijian species.

4. Cyrtandra vitiensis Seem. (Bonplandia 9: 257. 1861, nom. nud.; Seem. ex A.
Gray, Proc. Amer. Acad. 6: 41. 1862, nom. nud.) Fl. Vit. 182. 1866.
Cyrtandra amicta A.C, Smith, Journ. Arnold Arb. 34:38. 1953.

Erect herbaceous to shrubby plants up to 4 m, high, with coarse
branches up to 2 em. in diameter and with extensive pith ca. 1.5 em.
in diameter, the young vegetative parts with sericeous pubescence of
sepatate, uniseriate, noncapitate hairs ca. 20 in diameter and up to
1.5 mm. long; leaves opposite, with winged petioles 4-15 em. long,
the blades lanceolate to oblanceolate to ovate, up to 70 cm. long and
22 cm. broad, with an inequilateral, decurrent base and an acute to
obtuse to rounded apex, entire to undulate to faintly serrate at margin,
glabrous above, faintly pubescent beneath with noncapitate hairs, the
primary veins raised beneath, 6-20 per side, curved upward, the
secondary veins forming a conspicuous reticulate pattern beneath;
inflorescences congested-capitate, many-flowered, on stout peduncles
5-10 mm. long, these terminated by a pair of ovate, greenish white,
lobed bracts up to 3 em. long, these glabrous without and densely
sericeous within, subtending a bracteolate, branching cyme, the
bracteoles ovate-elliptic to lanceolate, 2- or 3-lobed, up to 3 em. long,
similar to the bracts in color and indument; calyx 1.5-4 em. long,
persistent, but rotting from the mature fruit, cleft ca. 0.25 of its
length into unequal lanceolate to triangular lobes, these glabrous or

GILLETT—CYRTANDRA IN FIJI 121

with scattered capitate hairs without and densely sericeous within at
the base of the tube; corolla cylindrical, 1.7-4.5 cm. long, with 5
rounded, unequal lobes 2-10 mm. long, sericeous without when young,
becoming glabrate at maturity, glabrous within; filaments 2-5 mm.
long, arising opposite the lower sinuses and bearing apically coherent
anthers at or below the corolla limb; staminodes 2, ca. 0.5 mm. long,
adnate opposite the upper sinuses at about the same level as the
stamens; cupulate annular disc prominent, ca. 1-1.5 mm. high at
anthesis, deciduous from maturing fruits; ovary and style ca. 1-1.5
em. long, the ovary glabrous, the style with sericeous indument on
upper 0.33 of its length, but with capitate hairs below the stigma, the
stigma with 2 flattened, applanate, ovate lobes perpendicular to the
style, the style separating 1-2 mm. above the summit of the mature
ovary; fruit white, ellipsoid, up to 2.5 cm. long.

Type tocauity: Namosi, Viti Levu, Fiji. Type collected by See-
mann, cited below.

Distrisution: Upland rain forests of Viti Levu, between 125 and
1300 meters.

Vitt Levu: Mba: Mount Koroyanitu, Mount Evans Range, Smith 4244 (A,
BISH, US); Mount Tomanivi, Smith 5914 (A-type of Cyrtandra amicta, BISH,
US), Smith 5103 (A, US), Gillespie 4095 (BISH, NY, UC, US); Nandala Creek,
near Nandarivatu, Smith 6223 (A, BISH, US), Degener 14925 (A, NY). Nan-
dronga & Navosa: Rairaimatuku Plateau, Smith 5654 (A, BISH, US). Serua:
Korovou, St. John 18941 (BISH, US). Namosi: Woods near Namosi, Seemann 277
(GH, K-type); Voma Mountain, Gillespie 2480 (BISH); without further locality,
Parks 20239 (BISH, UC, US), Parks 20429 (UC), Parks 20271 (BISH, UC, US);
Korombasambasanga Range, Smiih 8649 (BISH, US). Naitasiri: Matawailevu,
St. John 18184 (BISH, US). Rewa: Near Suva, Tothill 649 (BISH), Setchell &
Parks 15156 (UC), Setchell & Parks 15135 (UC).

Useful distinguishing characters of Cyrtandra vitiensis include the
large, lanceolate to oblanceolate, glabrate to faintly pubescent leaves
with decurrent bases and reticulate venation beneath, and the con-
gested, many-flowered, bracteate inflorescences, and glabrous outer
surfaces of the bracts. The corolla has a sericeous outer pubescence
when young but becomes glabrate to glabrous upon maturity, as in
specimens originally referred to C. amicta. Both phases have calyces
with pilose inner surfaces and are so closely similar in leaf and
inflorescence characters that the possibility of their being conspecific
had to be evaluated. After considerable study, I have concluded
that their separation must rest on calyx and corolla measurements
that contrast the larger flowers of C. amicta (calyx 3.5—4 cm. long;
corolla 4—4.5 cm. long) with the smaller flowers of C. vitiensis (calyx
1.5—2.5 cm. long; corolla 1.7—2.8 cm. long). It is my conclusion
that these differences in size, based on very limited material, probably
reflect the normal variability within C. vitiensis.

269-812—672

122 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Specimens (Smith 4243, A, BISH, US) intermediate between
Cyrtandra vitiensis and C. milnet have been collected on Mount
Koroyanitu, Mount Evans Range, Mba, where both species occur.
This collection surely portrays a combination of characters of the two
species, including the decurrent petioles and the lanceolate to
oblanceolate lamina of C. vitiensis, and the more dense indument of
petioles and leaves, the shorter, triangular calyx-lobes, and the high
frequency of capitate hairs, all of which characterize C. milnei. The
collection is placed under C. milnei in the present study. There seems
little doubt that the two species are very closely related.

The facies of certain species of the Hawaiian Islands suggests a
close relationship with Cyrtandra vitiensis and other species of Group 1.
These species include C. oblanceolata St. John & Storey and C.
warianuensis Rock, which have the habit as well as the congested
inflorescence and indument of C. vitiensis. In addition, the habit
inflorescence, leaves, and flowers of C. richit A. Gray, of Samoa,
suggest that it is of the same evolutionary stock as C. vitiensis.

5. Cyrtandra leucantha A. C. Smith, Journ. Arnold Arb. 34: 41. 1953.

Shrub up to 3 m. high with brown indument of septate, uniseriate
capitate and noncapitate hairs up to ca. 100u in diameter and 1-2 mm.
long; leaves opposite, the petioles 3-10 cm. long, with pubescence
shorter than petiole diameter, the blades oblanceolate to lanceolate-
ovate to ovate, up to 32 cm. long and 15 em. broad, inequilateral and
cuneate to attenuate at base, acute to acuminate at apex, coarsely
serrate to biserrate at margin, tomentose on both surfaces; inflores-
censes many-flowered, congested, the peduncles ca. 2-5 mm. long,
terminated by glabrate, ovate to elliptic to reniform, corolloid,
persistent bracts up to 1 cm. long, these subtending branching,
bracteolate cymes 5-10 mm. long, the bracteoles similar to and closely
subtended by the outer bracts; calyx white, 1.5 cm. long, unequally
cleft into 5 triangular-acuminate lobes 2-3 mm. long, persistent,
with dense outer pubescence of capitate and noncapitate hairs, with
inner pubescence of noncapitate hairs; corolla white, up to 2.5 cm.
long, cleft into subequal, rounded lobes 2-3 mm. long, with dense,
capitate and noncapitate hairs without, glabrous within; filaments
ca. 3 mm. long, arising ca. 10 mm. below the sinuses of the lower
corolla lobe and bearing apically coherent anthers well below the
corolla throat; staminodes 2, ca. 1 mm. long, recurved, adnate opposite
the upper sinuses at approximately the same level as the filaments;
cupulate annular disc conspicuous, ca. 1 mm. high, deciduous from the
mature fruit; ovary and style 12-15 mm. long, the ovary glabrous, the
style with long, capitate hairs throughout its length, terminated by an
applanate, bilobed stigma, the broadly ovate lobes of the stigma

GILLETT—CYRTANDRA IN FIJI 123

perpendicular to the axis of the style, the style separating 1-2 mm.
above the apex of the mature ovary; fruit white, ellipsoid, ca. 2 cm.
long and 1.4 cm. wide.

Type LocaLity: Mount Nanggaranambuluta, Mba, Viti Levu,
Fiji. Type collected by A. C. Smith, cited below.

Locau NAME: ‘‘Mbeta kai’ (Smith 4767).

DistripuTion: Upland rain forests between 300 and 1,100 meters,
in Mba, Viti Levu, and on Taveuni.

Vitt Levu: Mba: Mount Nanggaranambuluta, Smith 4767 (A. BISH, US),
Smith 6312 (A-type, BISH, US), Gillespie 3689 (A, BISH, UC); Nandarivatu,
Tothill 644 (BISH), Degener & Ordonez 13523 (A. BISH, NY, UC, US); Nandala,
Degener 14836 (A, BISH, NY, UC, US). Taverunt: Hills east of Somosomo, Smith

8375 (BISH, US); Somosomo, edge of woods, Gillespie 4780 (BISH, US); valley
between Mount Manuka and Mount Koroturanga, Smith 8253 (BISH, US).

The relationship between this species, Cyrtandra chlorantha A. C.
Smith, and C. milnei A. Gray is very close. A tabular summary of
distinguishing characters is presented under C. chlorantha. It has
been pointed out in the discussion under C. occulta and C. vitiensis
that this group of species is also represented in Polynesia, Borneo,
and the Philippine Islands. An additional western relationship is
portrayed by a collection from the Solomon Islands (Brass 3147, San
Cristoval, BISH) in which the calyx, leaf, and inflorescence char-
acters suggest the C. leucantha lineage.

6. Cyrtandra chlorantha A. C. Smith, Journ. Arnold Arb. 34: 42. 1953.

Shrub 3-4 m. high with pubescence of brown, septate, uniseriate
capitate and noncapitate hairs up to 120 in diameter and up to 4 mm.
long; leaves opposite, the petioles 3-5 cm. long, the blades lanceolate
to elliptic, up to 17 cm. long and 6 cm. broad, inequilateral and acute
to cuneate at base, acute to acuminate at apex, coarsely serrate,
tomentose to glabrate on both sides; inflorescences many-flowered,
congested, the penduncles up to ca. 0.5 cm. long, terminated by
persistent, green, ovate-elliptic bracts ca. 1.5 cm. long, these with a
dense pubescence of long, capitate hairs without, glabrous to glabrate
within, subtending short, branching bracteolate cymes, the bracteoles
similar to the bracts but smaller; calyx green, broad-cylindrical,
persistent, ca. 1.5 cm. long, cleft ca. 0.25 of its length into unequal
triangular lobes, with dense, capitate hairs without, glabrous to
glabrate with few capitate hairs within; corolla green, narrowly
cylindric, ca. 1.5-2.2 cm. long, slightly exserted beyond the calyx,
cleft ca. 0.2 of its length into rounded lobes, with capitate hairs
without, glabrous within; filaments adnate near the middle of the
corolla tube, ca. 3 mm. long, bearing apically coherent anthers 1-2
mm. below the corolla limb; staminodes 3, ca. 1 mm. long, adnate

124 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

opposite the upper sinuses at same level as filaments, the median
staminode ca. 1 mm. above the laterals; cupulate annular dise
prominent, ca. 1.5 mm. high; ovary and style ca. 1 em. long, the ovary
glabrous, the style with capitate hairs over its entire length, the
stigma applanate, bilobed, the lobes perpendicular to the axis of the
style, borne below the anthers.

Tyrer Locanity: Naitasiri, Viti Levu, Fiji. Type collected by A. C.
Smith, cited below.

Disrripution: Known only from the type locality, alt. 870-970 m.

Vitt Levu: Naitasiri: Northern portion of Rairaimatuku Plateau, between
Mount Tomaniviand Nasonggo, in dense forest, Smith 5789 (A-type, BISH, US).

This species and the related Crytandra milnet and C. leucantha can
be separated from each other by the following characters:

C. chlorantha C. milnet C. leucantha
Color of calyx and corolla: green greenish- white
white to
white
Inner corolla surface: glabrous pubescent glabrous
Number of staminodes: 3 3 2
Length of petiole hairs: 3-4 mm. 3-6 mm. 1-2 mm,

7. Cyrtandra milnei (Seem. in Bonplandia 9: 257. 1861, nom. nud.) Seem. ex
A. Gray, Proc. Amer. Acad. 6: 40. 1862.
Cyrtandra desvoeuxit Horne (A year in Fiji, 260. 1881, nom. nud.) ex C. B.
Clarke in DC Monogr. Phan. 5: 265. 1883.
Cyrtandra glandulosa Gillespie, Bishop Mus. Bull. 74: 22, fig. 28, 1980.

Sparsely branched herb or shrub 1-4 m. high with indument of
brown, septate, uniseriate trichomes, the capitate hairs 50-100u in
diameter and 1-2 mm. long, the noncapitate hairs up to 120 in di-
ameter and up to 6 mm. long; leaves opposite, the petioles up to 20
em. long, densely villous, the hairs equal to or longer than the petiole
diameter, the blades lanceolate to elliptic to ovate to obovate, up to
50 cm. long and 21 em. broad, rounded, obtuse, acute, cuneate, or
attenuate at base, obtuse, acute, or acuminate at apex, simply or
doubly serrate, tomentose to glabrate above, moderately pilose be-
neath; inflorescences many-flowered, congested, the peduncles to 1
em. long, terminated by corolloid, green to white, ovate to elliptic
bracts up to 2 cm. long and 1 em. wide, with a sparse to dense pubes-
cence of capitate hairs without, glabrous within, subtending a bracteo-
late, branching cyme 0.5-3 cm. long, the bracteoles similar to but
smaller than the bracts; calyx green to white, cylindrical, persistent,
1-2.5 em. long, cleft ca. 0.25 of its length into 5 unequal lobes, the 3
smaller upper lobes triangular, the 2 larger lower lobes lanceolate-
triangular, with capitate hairs on both surfaces; corolla green to white,
2-3 em. long, cylindrical, cleft ca. 0.12 of its length into unequal,

GILLETT—CYRTANDRA IN FIJI 125

rounded, erect lobes, with capitate and noncapitate hairs on both
surfaces; filaments 3-4 mm. long, arising 8-10 cm. below the sinuses of
the lower corolla lobe, bearing apically coherent anthers 1-2 mm.
below the limb; staminodes 3, clavate, adnate opposite the upper
sinuses at approximately the same level as the filaments, the medial
staminode up to 2 mm. long, the lateral to 1 mm., borne slightly
above the medial; cupulate annular disc prominent, ca. 1 mm. high,
deciduous; ovary and style 1.2-2 cm. long, the ovary glabrous, the
style with capitate hairs, separating 1-2 mm. above the apex of the
mature ovary, the stigma applanate, bilobed, the lobes perpendicular
to the axis of the style; fruit white, ovoid, up to 2 cm. long and
1.5 cm. wide.

Typr LocaLity: Ovalau, Fiji. Type collected by the U.S. Exploring
Expedition, cited below.

Locau NAME: ‘‘Me-diri-ta-bua”’ (Gillespie 2601); “beta levu’”’ (Gil-
lespie 3852); and “‘kau-bi-bi” (Gillespie 2749).

Distrinution: Dense forests between 50 and 1,250 meters on Viti
Levu, Kandavu, and Ovalau.

Vitt Levu: Mba: Mount Koroyanitu, Mount Evans Range, Smith 4129 (A,
BISH, US), Smith 4243 (A, BISH, US); valley of the Singatoka River, near
Nandarivatu, Gillespie 3852 (BISH-type of Cyrtandra glandulosa Gillespie, UC);
Mount Tomanivi, Gillespie 4108 (BISH, UC, US), Smith 5101 (A, BISH,
US), Degener 14335-a (A, NY); near Nandarivatu, Gillespie 4253 (BISH, UC,
US); Nauwanga, near Nandarivatu, Degener 14686 (A, BISH, NY, UC, US);
hills between Nggaliwana and Nandala ereeks, south of Nauwanga, Smith 5848
(A, BISH, US). Namosi: Namosi Village, Gillespie 2601 (BISII); Voma Mountain,
Gillespie 2749 (BISH); hills east of Wainikoroiluva River, near Namuamua,
Smith 8940 (BISH, US), Smith 8963 (BISH, US). Kanpavu: Mount Mbuke
Levu, Smith 287 (BISH, NY). Ovatau: Near summit of range west of Levuka
Gillespie 4445 (BISH, NY, UC); Lovoni Valley, Horne 179 (GH, K-type of
Cyrtandra desvoeuxii Horne ex C. B. Clarke); without further locality, U.S. Expl.
Exped. s.n. (GH, US-type).

Seemann used Gray’s description, verbatim, in Flora Vitiensis.
Clarke wrote an abbreviated version of this, following it with a longer
description of Seemann 281, which is Cyrtandra chippendalei, not C.
milnet.

The designation of Ovalau as the type locality for this species rests
on indirect evidence. ‘The Wilkes Expedition is known to have collected
six species of Cyrtandra in Fiji. The type material of C. dolichocarpa
A. Gray and asterile specimen of C. harveyi bear the locality of Sandal-
wood Bay, Vanua Levu. The remaining four species almost certainly
were obtained on Ovalau, where the expedition spent a total of 42
days, nearly half of the visitation to Fiji. Pickering noted four collec-
tions of Cyrtandra on Ovalau. Ovalau is designated as the locality
for three of these: C. involucrata Seem., C. anthropophagorum Seem.

126 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM
ex A. Gray, and C. pritchardii Seem. The label of the fourth collection,
the type of C. milnet, carries only “Feejee Islands” for locality; this
is cited by Gray, but it may be assumed that the collection was ob-
tained on Ovalau.

The bracts of the inflorescence are large and conspicuous, often
exceeding the calyx. There is considerable range in flower size and
this is sometimes expressed within a single inflorescence. The overall
variability of this species is extensive and includes variation in leaf
size, leaf shape, frequency and dimensions of capitate and noncapitate
hairs. The colors of the bracts, calyces, and corollas have been vari-
ously reported as from greenish white to white. This variability extends
to Cyrtandra vitiensis through one collection, Smith 4243, which is
intermediate between C. milnei and C. mtiensis. The variability of
present collections indicates that C. milnei is a complex assemblage
of populations that make a difficult taxonomic problem. A portion of
this complex was described by Gillespie as C. glandulosa. However, I
have been unable to separate Gillespie’s material from the rest of
the complex.

8. Cyrtandra dolichocarpa A. Gray, Proc. Amer. Acad. 6:40. 1862. Cyrtandra
microstigma C. B. Clarke, in DC. Monogr. Phan. 5: 284. 1883.

Shrub or spreading tree 1-4 m. high with ferrugineous pubescence
of septate, uniseriate, capitate and noncapitate hairs up to 100 in
diameter and 6 mm. long; leaves opposite, the petioles 1-4 ecm.
long, the blades lanceolate-ovate, up to 21 cm. long and 6 em. broad,
with inequilateral, obtuse, acute, or cuneate bases, acute to acuminate
at apex, serrate, with 2 or 3 teeth per cm., moderately pubescent
above, densely pubescent beneath, the veins obscure above, conspic-
uous beneath, the primary veins raised, curved upward, 6-8 per side,
the secondary veins reticulate; inflorescences pilose, the peduncles
0.5-1 em. long, terminated by persistent, ovate bracts 3-5 mm. long,
subtending 1-5 bracteolate pedicels 2—2.5 em. long, the bracteoles
ca. 3 mim. long, pilose on both surfaces; calyx 2.5-38 cm. long, per-
sistent, cylindrical, the lanceolate-acuminate lobes connivent, parted
10-15 mm. along 1 or 2 sinuses, with dense, noncapitate hairs on
both surfaces; corolla chartaceous, distinctly bilabiate, white, up
to 5.5 em. long, cleft about 0.25 to 0.33 of its length into unequal,
rounded lobes, the lower (anterior) lobe up to 2 em. long and 1 em.
broad, the limb with conspicuous longitudinal veins, with numerous
capitate hairs on both surfaces, the tube with capitate hairs without,
glabrous within; filaments ca. 3 mm. long, arising 3-4 mm. below
sinuses of the anterior corolla lobe, bearing apically coherent anthers
at the throat; staminodes 3, 1.5 mm. long, arising ca. 5 mm. below
sinuses of the upper lip, the median slightly shorter and borne 1-2

GILLETT—CYRTANDRA IN FIJI 127

mm. above the lateral; cupulate annular disc conspicuous, ca. 1
mm. high; ovary and style ca. 2.5 cm. long, extending to ca. 5 mm.
below corolla throat, densely pubescent with capitate and noncapitate
hairs below, capitate hairs above, the ovary attenuate to the style
and stigma, the stigma applanate, bilobed, the lobes thick, ca. 1
mm. long, spreading obliquely to the axis of the style; fruit cylindrical,
elongate, up to 5.5 cm. long and 5-7 mm. broad.

Typz tocautiry: Mbua Bay (Sandalwood Bay), Vanua Levu. Type
collected by the U.S. Exploring Expedition, cited below.

DistTriBUTION: Dense forests of Vanua Levu and Rambi, 100-700 m.

Locat NAME: “Muskarimba” (Smith 612, 1594).

Vanua Levu: Mbua: Mbua Bay, U.S. Expl. Exped. (GH, US-type) ; upper Ndama
River Valley, Smith 1594 (BISH). Mathuata: Northwestern slope of Mount
Numbuiloa, east of Lambasa, Smith 6530 (A, BISIT, US). Thakaundrove: South-
western slope of Mount Mbatini, Smith 612 (BISH, GH, NY, UC, US). Rambi:
No locality, Horne 439 (K-syntype of Cyrtandra microstigma C. B. Clarke, the
other syntype, Horne 439a, has not been seen.)

The present species has the largest flowers and surely the most
peculiar, distinctive fruits of all the Fijian species. The extension of
this lineage into Polynesia is indicated by similar flowers, inflores-
cences, and fruits in the Cyrtandrae of Samoa, such as C. krugert
Rein., C. nudiflora C. B. Clarke, and C. pulchella Rich., all of which
have glabrous fruits, and C. vaupelit Lauterb., in which the fruits
have a pubescence similar to that of C. dolichocarpa.

9. Cyrtandra multiseptata Gillespie, Bishop Mus. Bull. 74:23, fig. 31. 1930.

Shrub or slender tree 2-4 m. high with brown indument of septate,
uniseriate noncapitate hairs up to 60 in diameter and 3 mm. long;
leaves opposite, the petioles 1-6 cm. long, the blades lanceolate to
ovate to obovate, up to 20 cm. long and 10 cm. wide, inequilateral,
cuneate at base, acute to acuminate at apex, serrate, with 1-3 teeth
per cm., at maturity glabrate above, tomentose beneath; inflores-
cences tomentose, the penduncles 2-5 mm. long, terminated by con-
nate, deciduous bracts ca. 6 mm. long, subtending 1-3 bracteolate
pedicels ca. 0.5 cm. long, the bracteoles connate, deciduous, equal to
or smaller than the bracts; calyx 3.5 cm. long, ventricose, cleft ca.
0.33 of its length into unequal lanceolate-acuminate lobes, with non-
capitate hairs ca. 40u in diameter without, and a dense tomentum of
smaller hairs ca. 154 in diameter within; corolla with coarse, non-
capitate hairs without, glabrous within; cupulate annular disc con-
spicuous, ca. 1 mm. high, persisting as a broken, irregular ring on
mature fruits; fruit fusiform, sheathed by the calyx, up to 2.5 cm.
long and 1 cm. wide.

128 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

TypE LocaLity: Wet canyons near Namosi Village, Viti Levu, Fiji.
Type collected by Parks, cited below.

DistriputTion: Dense forests between 800 and 1,000 meters, Mba
and Namosi, Viti Levu.

Vitt Levu: Mba: Western slopes of Mount Nanggaranambuluta, east of
Nandarivatu, Smith 6327 (A, BISH, US); Mount Tomanivi, Smith 5102 (A, US).
Namosi: Canyons near Namosi Village, 800 m., Parks 20237 (BISH-type UC, US).

This species is distinguished from Cyrtandra dolichocarpa and C.
ventricosa by the differences cited in the tabular summary given under
the latter species. The habit, inflorescences, and chartaceous corollas
of C. multiseptata, C. ventricosa, and C. dolichocarpa suggests that the
three species are of a common evolutionary line. The extension of this
lineage into Polynesia is discussed under C. dolichocarpa.

10. Cyrtandra ventricosa Gillett, sp. nov.

Herba suffruticosa 1-2 m, alta, pilis septatis 120% diametro et ad
6 mm. longis induta; folia opposita, petiolis 1-7 cm. longis, laminis
ovatis vel lanceolato-ovatis ad 22 em. longis et 12 em. latis, basi
subaequilateraliter cuneatis, apice acutis vel acuminatis, margine
hydathodis serratis, supra glabratis, subtus dense pubescentibus, venis
primariis utrinsecus 6-10 curvatis; inflorescentiae pilosae pauciflorae,
pedunculis axillaribus erectis 1-2 mm. longis, bracteis terminalibus
lanceolatis caducis foliosis ca. 5 mm. longis, pedicellis bracteatis 10-12
mm. longis, bracteis interioribus caducis deltoideis basi connatis; calyx
persistente cylindricus fauce constrictus 1.5 em. longus in lobis 4 vel
5 valvatis lanceolatis 0.25 fissus utrinque pilosus; corolla chartacea
viridi-alba vel eburnea 18 mm. longa, utrinque glabra, reticulato-
venosa, in lobis 5 inaequalibus fissa, lobis 2 posterioribus 7 mm. longis
et 6 mm. latis, lobis 3 anterioribus 10 mm. longis et 9 mm. latis;
stamina 2 mm. infra sinus corollae lobi inferioris enata, antheris
apice adhaerentibus in corollae faucium ore; staminodia 3 infra
filamenta 2 mm. et sinibus superioribus opposita disposita, staminodio
medio ca. 1 mm. longo et 1 mm. supra staminodia lateralia 2 mm.
longa; discus annulari-cupuliformis prominens 1.5 mm. altus, sub
fructu interruptus; gynoecium 1.5 cm. longum glabrum, styli parte
superiore caduca, stigmate bilobato, lobis applanatis obliquis; fructus
ovoidel 12 mm. longi, 7 mm. lati.

Type in the herbarium of the Bernice P. Bishop Museum, collected
on the southern slope of the Valanga Range, Province of Thakaun-
drove, Vanua Levu, Fiji, alt. 100 m., Noy. 11, 1933, by A. C. Smith
(868). Duplicates at A, NY, UC, US.

ADDITIONAL SPECIMENS EXAMINED:

Vanua Levu: THakauNnprove: Savu Savu Bay, Vatunivuamonde Mountain,
Degener & Ordonez 13968 (A, BISH, UC, US).

GILLETT—CYRTANDRA IN FIJI 129

Locau NAME: ‘‘Merikula” (Smith 368).

The affinities of this species are with Cyrtandra multiseptata and
C. dolichocarpa. It is distinguished from the others by its indument,
fruits, and habit, as shown in the following tabular summary:

C. ventricosa C. multiseptata C. dolichocarpa
Dimensions of leaf 120% * 6-7 mm. 604 X 3 mm. 100u X 6 mm.
trichomes:
Fruit shape: ovoid fusiform cylindrical
Habit: subligneous shrub or shrub or tree
herb 1-2 slender tree 1-4 m. high
m. high 2-4 m. high

11. Cyrtandra acutangula Seem. (Bonplandia 9: 257. 1861, nom. nud.; Seem. ex
A. Gray, Proc. Amer. Acad. 6: 41. 1862, nom. subnud.) Fl. Vit. 182. 1866.
Cyrtandra utriculosa C. B. Clarke, in DC. Monogr. Phan. 5: 230. 1883.
Shrub 2-3 m. high with dark brown indument of septate, uniseriate,
noncapitate hairs 15-25y in diameter and up to 1 mm. long, the young
branches sharply quadrangular, the bark with linear striations; leaves
opposite, the petioles 1-6 cm. long, 3-4 mm. in diameter, the blades
dark brown, lanceolate to elliptic to ovate, up to 23 cm. long and 8
cm. broad, equilateral and cuneate at base, acute to obtuse at apex,
faintly serrate, when young glabrous above and densely brown-
sericeous beneath, at maturity glabrous on both surfaces, the lower
surface with conspicuous, raised veins, the secondary veins perpendicu-
lar to the primary veins; inflorescences tomentose, the peduncles 2-8
mm. long, reflexed, terminated by a pair of linear-lanceolate, free,
caducous bracts up to 1.5 cm. long and subtending 1-3 bracteolate
pedicels, the bracteoles lanceolate, free, caducous, up to 7 mm. long;
calyx cylindrical, persistent, up to 3.5 em. long, cleft to about half its
length into lanceolate, acuminate lobes, the tube glabrous without,
pilose with brown, ascending hairs 3-4 mm. long within; cupulate
annular disc 2 mm. high, persistent, forming a continuous ring in
mature fruit; fruit ovoid, about 1 cm. in diameter and 2 cm. long,
tipped by the basal 3 mm. of the style and enclosed in the persistent
calyx.
Typr tocauity: Namosi, Viti Levu, Fiji. Type collected by See-
mann, cited below.
Distripution: Known with certainty only from the upland rain
forests of Namosi Province, where it occurs at elevations near 1,000 m.
Vitr. Levu: Namosi: ‘‘Mountains near Namosi,”’ Seemann 276 (GH, K-type);
damp, shaded forests near Vienunga, Horne 911 (GH, K-type of Cyrtandra
utriculosa C. B. Clarke); Mount Naitarandamu, Gillespie 3120 (BISH, UC).
Viti Levu, without further locality: Parks 20478 (UC), 20954 (BISH, UC, US).
I have been unable to find a morphological basis for separating
Cyrtandra utriculosa C. B. Clarke from C. acutangula. Both species are

130 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

based on material from a limited region of Namosi, so that the re-
duction of C. utriculosa seems justified. The facies of this species is
very different from that of other species in Group 2, and I have seen
no Polynesian material that bears a close resemblance to it.

The present species differs from the Cyrtandra dolichocarpa-C.
ventricosa-C, multiseptata lineage in its much smaller trichomes,
sharply quadrangular stems, dark brown foliage, and glabrous leaves.
The trichomes of C. multiseptata are over twice the diameter of those
of the present species, and those of CO. ventricosa over five times.

12. Cyrtandra trichophylla A. C. Smith, Journ. Arnold Arb. 34: 44. 1953.

Shrub to slender tree 2-6 m. high with brown, appressed-sericeous
pubescence of septate, uniseriate, noncapitate hairs 20-40y in diame-
ter and up to 2 mm. long; stems up to 8 cm. in diameter 1.5 m. above
the ground, with much xylem and little pith; leaves opposite, the
petioles 1-5 cm. long, the blades ovate to obovate to oblanceolate,
up to 32 cm. long and 12 em. broad, inequilateral and acute to cuneate
at base, acute to obtuse to rounded at apex, dentate-crenate, glabrate
to glabrous above, sericeous beneath, the primary veins conspicuously
raised on lower surface, 9-12 per side, curved upward, the secondary
veins sometimes obscured by pubescence; inflorescences with brown,
dense, sericeous pubescence, few-flowered, nearly sessile, the stout
peduncles terminated by connate, caducous bracts ca. 5 mm. long,
subtending 1-3 pedicels 5-14 mm. long; calyx persistent, eventually
rotting from mature fruit, 12-15 mm. long, cleft 0.5 to 0.66 of its
Jength into lanceolate-acuminate lobes, densely sericeous on both
surfaces; corolla ca, 2 cm. long, curved-cylindrical, cleft ca. 0.25 of its
length into rounded, unequal lobes, densely sericeous without, the
limb and throat with capitate hairs within; filaments ca. 3 mm. long,
bearing apically coherent anthers at the base of the corolla limb ;
staminodes 3, ca. 0.5 mm. long, arising 5-6 mm. below the upper
sinuses; cupulate annular disc thick, prominent, ca. 1 mm. high,
breaking into plates persisting in mature fruit; ovary and style 12
mm. long, the ovary glabrous, the style with numerous capitate hairs
in the upper half of its length, separating 1-2 mm. above the mature
ovary, the stigma borne 1-2 mm. below the anthers, bilobed, applanate,
the lobes oblique to the axis of the style; fruit white, ovoid-ellipsoid,
up to 2 cm. long and 1 em. wide.

Type nLocauity: Ridge from Mount Namama toward Mount
Tomanivi, Viti Levu, Fiji. Type collected by A. C. Smith, cited below.

Distripution: Dense forests of Viti Levu from 100 to 1,300 m.

Vitt Levu: Mba: Nandala Creek, 3 mi. south of Nandarivatu, Smith 6246
(A, BISH, US). Namosi: Ridges near Namosi Village, Gillespie 2690 (BISH,
NY, UC); Valley of the Waindina, near Namosi, Gillespie 2595 (BISH); Naitaran-
damu Mountain, Gillespie 3243 (BISH), Gillespie 3244 (BISH, UC), Gillespie 3378

GILLETT—CYRTANDRA IN FIJI 131

(BISH, UC), Gillespie 3303 (BISH, UC); Wainavindrau Creek, near Wainima-
kutu, Smith 8544 (BISH, US); Vakarongasiu Mountain, Gillespie 3289 (BISH,
UC); without further locality, Parks 20269 (BISH, UC). Ra: Ridge from Mount
Namama toward Mount Tomanivi, Smith 5698 (A-type, BISH, US). Naitasiri:
Wainisavulevu-Numbulolo Divide, St. John 18324 (BISH, US); near Tamavua,
150 m., Gillespie 2455 (A, BISH), Tamavua Village, Gillespie 2116 (BISH, UC).
Rewa: Korombamba Mountain, Gillespie 2257 (BISH, UC), Parks 20138 (UC).

Among Fijian Cyrtandrae this species has the most conspicuously
woody habit; the proportion of wood in the stem and branches is
greater than it is known to be in any of the other species. It occurs
in the territory of Cyrtandra vitiensis but is easily distinguished by its
open, few-flowered inflorescences in contrast to the capitate, many-
flowered inflorescences of C. vitiensis. The contrast in the proportion
of wood in these species is shown below:

C. vitiensis C. trichophylla
Diameter of stem (mm.) 20 10; 15; ‘16
Diameter of pith cylinder (mm.) 15 4 5 4

Neither Cyrtandra trichophylla nor any closely related species are
known to occur in Polynesia.

13. Cyrtandra muskarimba A.C. Smith, Bishop Mus. Bull. 141:182. fig. 67. 1936.

Shrub or succulent herb 1-3 m. high, the young parts with dark
brown, appressed, septate, uniseriate, noncapitate hairs ca. 60-80
in diameter and up to 3 mm. long; leaves opposite, the petioles 4-11
em. long, the blades elliptic to ovate, up to 28 em. long and 15 cm.
broad, unequal and obtuse to cuneate at base, acute to acuminate at
apex, finely serrate, above with appressed, usually parallel, sericeous
hairs, beneath densely pubescent, with 9-11 upwardly curved, raised
primary veins on each side of midvein, the secondary veins obscured
by pubescence; inflorescences elongate, branching cymes with 1-4
flowers; peduncles 1-2 em. long, terminated by lanceolate bracts ca.
1.5 em. long, subtending pedicels ca. 1.5 cm. long; calyx persistent,
ca. 1.5-1.8 cm. long, cleft 0.5 to 0.66 of its length into unequal,
triangular to lanceolate lobes, with dark brown, appressed-ascending
pubescence on both surfaces; corolla white, 2.0-2.3 cm. long, cleft
into 5 rounded, unequal lobes 2-3 mm. long, cylindrical, coriaceous,
externally glandular with capitate hairs or pilose with ascending,
noncapitate hairs, within with capitate hairs; filaments 2-3 mm. long,
bearing apically coherent anthers ca. 1 mm. below the corolla limb;
staminodes 2, ca. 0.5 mm. long, arising near middle of corolla tube;
cupulate annular disc conspicuous, ca. 1 mm. high, persisting as a
broken ring in mature fruit; ovary and style 7-8 mm. long, the style
with capitate hairs in upper 0.25 of its length; stigma borne 1-2
mm. below the anthers, weakly bilobed; fruit ovoid, ca. 1.5 em. long

132 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

and 0.8 cm. wide, tipped by 1-2 mm. of the persistent style base,
sheathed by the persistent calyx.

Typr tocauity: In dense forest on the southwestern slope of
Mount Mbatini, Vanua Levu, Fiji. Type collected by A.C. Smith,
cited below.

Locau NAME: “ Muskarimba” (Smith 1914).

Distripution: Upland rain forests of Serua, Viti Levu, and
Thakaundrove, Vanua Levu, between 300 and 700 meters.

Vitt Levu: Serua: Namboutini, Damanu 70 (K); Mbuyombuyo, near Nam-
boutini, Tabualewa 15577 (A, BISH, NY, UC, US). Vanua Levu: Thakaundrove:
Southwestern slope of Mount Mbatini, Smith 673 (BISH-type, GH, NY, UC,
US); Mount Ndikeva, Smith 1914 (BISH, GH, NY, UC, US).

This species is closely related to Cyrtandra trichophylla, differing
from it in its decidedly less woody habit, larger trichomes, shorter
peduncles, and the presence of only two staminodes.

14. Cyrtandra victoriae Gillespie, Bishop Mus. Bull. 74: 25, fig. 34. 1930.

Shrub to small tree 2-5 m. high with ferrugineous, septate, uni-
seriate, noncapitate hairs ca. 100-160u in diameter and up to 5 mm.
long; leaves opposite, the petioles 2-6 em. long, with dense, velutinous
pubescence, the blades ovate, obovate, oblanceolate, or lanceolate,
up to 85 em. long and 15 cm. broad, inequilateral and acute to cuneate
at base, acute to acuminate at apex, serrate to entire, with moderate
to dense tomentum above, the hairs lying parallel and upwardly
directed on younger leaves, spreading on mature foliage, velutinous
beneath; inflorescences with few flowers, the peduncles stout, 0.5-1
em. long, terminated by velutinous, greenish white, ovate to lanceo-
late, attenuate, free to connate, caducous bracts 3-5 cm. long, these
subtending 2-4 pedicels 5-10 mm. long; calyx white, growing with
the developing fruit and persistent on mature fruits, 1.7-3 cm. long
at anthesis, 83-4 cm. long in mature fruit, cleft 0.5 to 0.66 of its length
into unequal, lanceolate-attenuate lobes, velutinous without, within
glabrous above, below with a dense tomentum of ascending non-
capitate hairs; corolla white, ca. 3.0-3.5 em. long, cylindrical, slightly
curved, cleft into rounded, unequal lobes 4-5 mm. long, with velu-
tinous outer pubescence projecting well above the lobes, the inner
surface with capitate hairs in upper third and mixed noncapitate and
capitate hairs below; filaments 4-5 mm. long, bearing apically coherent
anthers at the corolla throat; staminodes 3, adnate near the middle
of the corolla tube, up to 1.5 mm. long, the median up to 0.5 mm.
long, borne 2 mm. below the lateral; cupulate annular dise conspicuous,
ca. 1 mm. high, persisting as a broken, thick, dark ring in mature
fruit; ovary and style ca. 1.8 em. long, the ovary glabrous, the style
with dense indument of capitate hairs in upper 0.75 of its length,

GILLETT—CYRTANDRA IN FIJI 133

separating 2-3 mm. above the apex of the fruit, the stigma weakly
bilobed; fruit ovoid, up to 2 cm. long and 1.5 cm. wide.

Typs Locautity: Mount Tomanivi, Mba, Viti Levu, Fiji. Type
collected by Gillespie, cited below.

Loca NAME: ‘‘Doo’-re le’vu” (Gillespie 4088).

DistRIBUTION: Dense upland forests of central Viti Levu, between
250 and 1,250 meters.

Vitt Levu: Mba: Mount Tomanivi, Gillespie 4088 (BISH-type, UC), Smith
5915 (A, BISH, US), 5916 (A, US), 5917 (A, BISH, US); summit of Mount
Nanggaranambuluta, Gillespie 4349 (BISH); escarpment west of Nandari-
vatu, Smith 5081 (A, BISH, US). Namosi: Summit of Vakarongasiu Mountain,
Gillespie 3263 (BISH); valley of Wainambua Creek, south of Mount Naita-
randamu, Smith 8801 (BISH, US). Ra: Mount Namama, east of Nandarivatu,
Smith 5697 (A, BISH, US), 5722 (A, BISH, US).

The significant features of Cyrtandra victoriae are the very large,
fleshy (drying to coriaceous) calyx and corolla, The flowers and fruits
of Smith 5081 and 5917 indicate that the calyx grows considerably
after anthesis, a useful distinguishing character for the species.

Length of Calyx

at anthesis in mature fruit
30 mm. 40 mm.
Smith 5081 (US) 20 mm. 30 mm.
17 mm. 38 mm.
Smith 5917 (US) 30 mm. 37 mm.
30 mm.

The closest relationship of this species is with Cyrtandra chippendaler,
and this is discussed under the latter species.

15. Cyrtandra chippendalei Horne ex C. B. Clarke, in DC. Monogr. Phan. 5: 230.
1883.
Cyrtandra tomentosa A. C. Smith, Sargentia 1: 116. 1942.

Herbaceous to shrubby plants 1-4 m. high with light brown, septate,
uniseriate, noncapitate hairs 100-160y in diameter and 2-5 mm. long;
leaves opposite, the petioles 2-10 cm. long, with villous pubescence of
hairs as long as the petiole diameter, the blades ovate to lanceolate
to oblanceolate, up to 28 cm. long and 13 em. wide, inequilateral and
rounded to cuneate to attenuate at base, acute to acuminate at apex,
finely serrate, moderately pubescent above and velutinous beneath;
inflorescences with 1-6 flowers, densely villous; peduncles 1-3 em. long,
terminated by ovate, acuminate, basally connate, caducous bracts
to 4 em. long, these subtending bracteolate pedicels 0.5-2 cm. long;
calyx 1.2-2.5 em. long, distended by the maturing fruits, cleft ca.
0.5 to 0.66 of its length into 5 unequal, triangular to attenuate-

134 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

lanceolate, foliaceous lobes, densely pilose on both surfaces; corolla
white, 2-3 cm. long, fleshy, drying to a coriaceous texture, cleft into
unequal, rounded lobes 3-4 mm. long, pilose without, glabrous within
or with capitate hairs; filaments 4-5 mm. long, bearing apically
coherent anthers slightly below the corolla throat; staminodes 2,
conspicuous, ca. 1.5 mm. long, adnate opposite the upper lateral
sinuses at the same level as the filaments; cupulate annular disc
conspicuous, ca. 1 mm. high, persisting as a fragmented brown disc in
mature fruit; ovary and style 10-12 mm. long, the ovary glabrous, the
style glabrous below, with dense indument of capitate hairs in the
upper half of its length, separating 1-5 mm. above the summit of the
mature ovary, the stigma applanate, bilobed, borne near the filament
bases; fruit ovoid, ca. 2.2 em. long and 1.8 cm. wide, the color not
known.

Type LocaLity: Mountain forests near Wai-Wai, Savu Savu Bay,
Vanua Levu, Fiji. Type collected by Horne, cited below.

Locat NAME: “Mbeta” (Degener 14889); “Muskarimba” (Smith
1986).

Disrrisution: Rain forests of Viti Levu, Vanua Levu, and Taveuni,
between 150 and 1,250 meters.

Vitt Levu: Mba: Hills east of Nandala Creek, about 3 mi. south of Nandarivatu,
Smith 6237 (A, US). Nandronga and Navosa: Nandrau, Degener 14889 (A-type of
Cyrtandra tomentosa A. C. Smith, BISH, NY). Namosi: Naitarandamu Mountain,
Gillespie 3351 (BISH, NY UC); Wainavindrau Creek, near Wainimakutu, Smith
8549 (BISH, US); Wainavindrau Creek, n. slope of Korombasambasanga Range,
Smith 8720 (BISH, US). Rewa: Korombamba, Parks 20129 (UC). Vanua Levu:
Thakaundrove: Near Wai-Wai, Savu Savu Bay, Horne 577 (GH, K-type);
Natewa Peninsula, Uluingala, Smith 1986 (BISH, GH, NY, UC, US). Taveunt:
Between Somosomo and Wairiki, Smith 885 (BISH, NY), hills east of Somosomo,
Smith 8377 (BISH, US); without further locality, Seemann 281 (GH, K).

The type has triangular sepals and the inner surface of the corolla is
glabrous. This is one of the most variable of Fijian Cyrtandrae.
However, this variablity involves differences that do not seem to be
in any case of sufficient magnitude to justify the recognition of more
than one species. There is a close relationship between Cyrtandra
chippendalei and C. vietoriae. The foliaceous, smaller calyx (1.2-2.5
em. long) of C. chippendalei does not have postanthesis erowth and
therefore is easily distinguished from the fleshy (drying to coriaceous)
calyx of C. victoriae, which has considerable postanthesis growth.

There is as yet no evidence that the Cyrtandra chippendalei lineage
extends into Tonga, Samoa, or eastern Polynesia. However, the
Cyrtandrae of Hawaii that are classified under subgenus Brachycyathus
C. B. Clarke (St. John, 1966) are of this lineage. The Hawaiian species
possess the calyx, inflorescence, and indument characters of Group 2
of the Fijian Cyrtandrae and surely would be placed near C. chip-

GILLETT—CYRTANDRA IN FIJI 135

pendalet. The discontinuous distribution of this group is comparable
to that of certain other Pacific genera. These include Cheirodendron
(Araliaceae), locally common in the Marquesas Islands and the
Hawaiian Islands but not known elsewhere; Pelea (Rutaceae),
restricted to Tahiti, the Marquesas, and Hawaii; Gouania (Rham-
naceae), in Fiji and Hawaii, but not known in Samoa or Tonga;
and Exocarpos (Santalaceae) in Fiji, Rapa, and Hawaii, but not
known elsewhere in Polynesia.

Hawaiian representatives of the Cyrtandra chippendalei line cur-
rently number 89 species on the island of Oahu (St. John, 1966).

16. Cyrtandra spathacea A. C. Smith, Bishop Mus. Bull. 141: 133. fig. 69. 1936.

Slender tree to 4 m. high, the young parts with indument of dark
brown, septate, uniseriate hairs ca. 20u in diameter and 0.2 mm. long;
leaves opposite, the petioles 1.5-6 cm. long, the blades lanceolate-
ovate, to ovate-elliptic, up to 22 em. long and 14 cm. broad, unequal,
obtuse to acute at base, acute to obtuse to rounded at apex, serrate,
with 2 or 3 obtuse teeth per cm., each dentition with a distal hyda-
thode, at maturity glabrous above, glabrate to glabrous below, the
primary veins 6-10 per side, curved upward, the secondary veins
conspicuous, reticulate; inflorescences branched cymes of 1-4 flowers,
the peduncles 1-2 mm. long, terminated by a pair of caducous bracts
ca. 1 mm. long, subtending a bracteolate cyme up to 4 cm. long;
calyx spathaceous, ca. 1.5 em. long, deciduous, cleft, and lacerated
nearly to base at one sinus, on both surfaces with scattered, non-
capitate hairs; corolla cream-white, glabrous on both surfaces, ca.
3 cm. long, cleft 0.5-1 cm. into unequal, rounded lobes, the broad
throat tapering into a tube ca. 1 cm. long; filaments ca. 5 mm. long,
adnate opposite the sinuses of the lower corolla lobe, bearing apically
coherent anthers at the sinuses; staminodes 3, ca. 1 mm. long, arising
ca. 5 mm. below upper sinuses, the median staminode 1-2 mm. below
the lateral; cupulate annular disc prominent, ca. 1 mm. high, with
inner pubescence of noncapitate hairs projecting above the rim,
the disc drying to an inconspicuous, fragmented ring in mature fruit;
ovary and style 1.5 cm. long, the ovary glabrous, the style with capi-
tate and noncapitate hairs over its entire length, separating 1-2 mm.
above apex of the mature ovary, the stigma applanate, bilobed,
the lobes thick, symmetrical, perpendicular to the axis of the style,
borne 4—5 mm. below the anthers; fruit white, ellipsoid, up to 12 mm.
long and 8 mm. wide.

TypE LocaLity: Kandavu, Fiji; the type is Smith 258, cited below.

DistriBuTION: Upland rain forests of Viti Levu and Kandavu.

Vitt Levu: Mba: Mountains near Lautoka, Greenwood 51 (K). Kanpavv:
Mount Mbuke Levu, Smith 258 (BISH-type, GH, NY, UC, US).

136 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

The spathaceous calyx is also present in Cyrtandra longiflora
J. W. Moore of Polynesia, and specimens of the latter species collected
in the Society Islands bear a striking similarity to the material from
Viti Levu. The spathaceous calyx is present in several Hawaiian
species of subgenus Cyrtandra. Additional evidence of the extension
of this group into Polynesia is presented under C. kandavuensis and
C’. attenuata.

17. Cyrtandra kandavuensis A. C. Smith, Bishop Mus. Bull. 141: 132, fig. 68.
1936,

Shrub or small, spreading tree 2-4 m. high, the foliage and in-
florescences with light brown septate, uniseriate, noncapitate hairs ca.
20 in diameter and 1 mm. long; leaves opposite, the petioles 3-9 em.
long, the blades ovate to elliptic, up to 25 cm. long and 14 cm. wide,
unequal, obtuse to oblique at base, acuminate at apex, serrate, with 1
or 2 teeth per cm., above with scattered, appressed hairs ca. 1 mm. long
and parallel to primary veins, beneath densely pubescent on the veins,
with nearly glabrate areoles, the primary veins 6-8 per side, curved
upward, converging at margins; inflorescences many-flowered,
branched, open cymes, with dense, erect, capitate and noncapitate
hairs up to 0.33 mm. long, the peduncles 4-9 cm. long, terminated by
a pair of deciduous, ovate, foliaceous bracts ca. 2-2.5 cm. long, the
pedicels 2.5-4 cm. long; calyx deciduous, ca. 1.5 cm. long, cleft 0.66
of its length into 5 equal, ovate, acuminate, valvate lobes, with non-
capitate and capitate hairs without, glabrous within; corolla white,
ca. 1.5-2 cm. long, cleft ca. 0.25 of its length into rounded lobes, the
throat funnelform, with capitate and noncapitate hairs without,
glabrous within; filaments ca. 3 mm. long, arising 3 mm. below the
sinuses of the lower corolla lobe and projecting upward scarcely more
than 1 mm., bearing apically coherent anthers below the corolla limb;
staminodes 3, ca. 1 mm. long, adnate ca. 5 mm. below the upper
sinuses, the median staminode about 1 mm. above the lateral; cupulate
annular disc conspicuous, ca. 1.5 mm. high, persisting as a divided
pelviform structure in mature fruit; ovary and style glabrous, ca. 10
mm. long, the style separating ca. 3 mm. above the mature ovary,
bearing an applanate, bilobed stigma ca. 2-3 mm. below the anthers;
fruit white, ellipsoid, ca. 12 mm. long and 8 mm. wide.

Typg Locauiry: Mount Mbuke Levu, Kandavu, Fiji. Type col-
lected by A. C. Smith, cited below.

Distripution: Dense forests on Kandavu, 200-840 m.

Kanpavu: Mount Mbuke Levu, Smith 282 (BISH-type, GH, UC, US), Smith
288 (BISH, GH, NY, UC, US), Smith 214 (BISH, GH, NY, UC, US).

Related material in Polynesia includes an undetermined Cyrtandra
from Tahiti, St. John 17154, and C. pupukeaensis St. John & Storey,

GILLETT—CYRTANDRA IN FIJI 137

of Oahu, Hawaiian Islands. Both have the inflorescence, calyx,
corolla, and indument of C. kandavuensis.

18. Cyrtandra attenuata Gillett, sp. nov.

Frutex 1-4 m. altus, pilis septatis ad 20y diametro et 0.5 mm. longis
indutus maturitate glaber; folia opposita, petiolis 1-3 cm. longis,
laminis lanceolatis vel ovatis ad 15 cm. longis et 7 cm. latis, basi
inaequilateraliter acutis vel cuneatis, apice acutis vel acuminatis,
dentatis vel undulatis; inflorescentiae cymosae subpatentes 2-8
florae, pedunculis 1-6 cm. longis indumento adscendente obtectis,
bracteis terminalibus binis linearibus 2-3 mm. longis caducis, pedicellis
1 vel 2 bracteolatis ad 6 cm. longis; calyx caducus albus vel viridi-
albus membranaceus manifeste venosus ad 2 cm. longus basi inflatus
in lobis deltoideis vel lanceolatis attenuatis aequalibus 0.33-0.5 fissus
utrinque glaber vel glabratus; corolla alba chartacea ad 4 cm. longa
in lobis paullo inaequalibus 0.25 fissa extus glabra intus capitato-
elanduloso-pilosa; stamina ca. 5 mm. infra sinus corollae lobi inferioris
enata, filamentis 4-5 cm. longis, antheris in corollae faucibus in apice
adhaerentibus; staminodia 3 infra sinus superiores 4-5 mm. disposita
1.5-2 mm. longa, staminodio medio paullo supra staminodia lateralia ;
discus annularicupuliformis prominens sub anthesi ca. 1 mm. altus;
gynoecium ca. 1.5 cm. longum, ovario glabro superne gradatim
attenuato, stylo inferne glabro superne capitato-glanduloso-piloso,
stigmate applanato bilobato 1-2 mm. infra antheras; fructus maturi
desiderati.

Type in the herbarium of the Bernice P. Bishop Museum, collected
near Uluingala, Natewa Peninsula, Province of Thakaundrove,
Vanua Levu, Fiji, alt. 600-820 m., June 15, 1934, by A. C. Smith
(2006). Duplicates at GH, NY, UC, US.

ADDITIONAL SPECIMENS EXAMINED:

Tavewntr: Hills east of Somosomo, west of old crater occupied by small swamp
and lake, dense forest, Smith 8366 (US).

The distinctive features of this species include the relatively large
flowers with inflated calyces and the chartaceous corollas with glabrous
outer surfaces. Material of the Samoan Cyrtandra divertae Christopher-
son bears a strong similarity to C. attenuata. A specimen with re-
markably similar flowers and inflorescences was collected in Tahiti
(St. John & Fosberg 14158 (BISH)). Finally, the new species is re-
markably similar to C. nukuhivensis F. Brown, of the Marquesas
Islands, the inflorescences and flowers being generally comparable
in form and indument. These comparisons seem to indicate clearly
that the lineage of the new species extends into western and eastern
Polynesia.

269-812-673

138 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

19. Cyrtandra involucrata Seem. (Bonplandia 9: 257. 1861, nom. nud.; Seem. ex

A. Gray, Proc. Amer. Acad. 6: 41, 1862, nom. subnud.) Fl. Vit. 183. 1866.
Cyrtandra monticola Gillespie, Bishop Mus. Bull. 74: 23, fig. 30. 1930.
Cyrtandra gillespieana A. C. Smith, Bishop Mus. Bull. 141: 134. 1936.

Shrub or small tree up to 5 m. high, the leaves and inflorescences
with medium to dark brown pubescence of shiny, septate, uniseriate,
noncapitate hairs up to 50u in diameter and 1 mm. long, densely
tomentose on younger parts, glabrate at maturity; leaves opposite,
the petioles up to 4.5 cm. loug, the blades lanceolate to oblanceolate
to elliptic, up to 20 em. long and 10 em. wide, unequal and rounded,
obtuse, attenuate, or cuneate at base, acute to acuminate at apex,
crenate to serrate, with about 3 teeth per cm., the teeth tipped by
hydathodes, tomentose to glabrate with appressed hairs or glabrous
above, with dense, spreading hairs beneath; inflorescences appressed-
sericeous, the peduncles 3-40 mm. long, terminated by 2 ovate,
connate-perfoliate, foliaceous, deciduous bracts 5-8 mm. long, sub-
tending 1-4 pedicels up to 2 cm. long; calyx deciduous, 6-8 mm. long,
cleft about half of its length into 5 unequal, lanceolate, acuminate
lobes, with noncapitate hairs on both sides; corolla cream to light
yellow, ca. 1.5 cm. long, eleft ca. 0.25 of its length into rounded,
subequal lobes, the tube cylindrical, with dense noncapitate and
capitate hairs without, with short, capitate hairs within; filaments
2-3 mm. long, bearing apically coherent anthers 1-2 mm. below the
corolla limb; staminodes 8, adnate opposite the upper sinuses ca. 3
min. below the limb, the median staminode 1-2 mm. above the lateral;
annular disc ca. 1 mm. high, persisting as rectangular plates in mature
fruit; ovary and style up to 1 em. long, the ovary glabrous, the style
with capitate hairs over its entire length, separating at summit of
ovary, the stigma applanate, bilobed, borne 2-3 mm. below anthers;
fruit white, ovoid to ellipsoid, ca. 1.8 em. long and 1.2 em. wide.

Typs Locaxity: “Woods of Namosi,”’ Viti Levu, Fiji. Type collected
by Seemann, cited below.

Locau NAME: “Tangitangi” (Tabwalewa 15596).

Disrripution: Interior mountains of Viti Levu, Ovalau, and
Vanua Levu, between 30 and 1,150 m. in densely shaded creeks and
drainages as an understory element in the rain forest.

Vitt Levu: Mba: Mount Evans Range, Greenwood 1228 (A, BISH, NY, UC,
US); Mount Evans Range, eastern slopes of Mount Koroyanitu, Smith 4136
(A, BISH, US); Thol-i-nandarivatu Mountain, Gillespie 3866 (BISH, UC); hills
east of Nandala Creek, ca. 3 mi. south of Nandarivatu, Smith 6221 (A, BISH,
US); western and southern slopes of Mount Tomanivi, Smith 5318 (A, BISH,
US). Nandronga & Navosa: Nausori highlands, in drainage of Namosi Creek
above Tumbenasolo, Smith 4717 (A, BISH, US). Serua: Mbuyombuyo, near
Namboutini, Tabualewa 15596 (A, BISH, UC, US); hills north of Ngaloa, in
drainage of Waininggere Creek, Smith 9426 (BISH, US). Namosi: Without
locality, Seemann 279 (GH, K-type); vicinity of Namosi Village, Gillespie 2691

GILLETT—CYRTANDRA IN FIJI 139

(BISH); Naitarandamu Mountain, Gillespie 3122 (BISU-type of Cyrtandra
monticola Gillespie, UC). Ovatau: Without further locality, U.S. Expl. Exped.
s.n. (GH, US). Vanua Levu: Mbua: H. B. R. Parham 17 (A).

The connate, foliaceous bracts subtending the pedicels provide a
distinguishing feature for this species. However, the bracts are
deciduous and if they are lacking, the species is easily confused with
Cyrtandra anthropophagorum, in which the inflorescence bracts are
also deciduous but are free and not connate. In such cases the follow-
ing characters are of value:

C. involucrata C. anthropophagorum
Color of pubescence dark brown golden brown
Pedunele length up to 4.5 cm. up to 3 em.
Trichomes of peduncle up to 1 mm. long, up to 1.5 mm. long,
appressed spreading

These two species are relatively common in Fiji, but neither they
nor related species are known to occur in Polynesia.

20. Cyrtandra reticulata Gillett, sp. nov.

Frutex 2-3 m. altus, partibus novellis pilis fuscis septatis ecapitatis
ca. 30u diametro ad 0.5 mm. longis dense sericeus, maturitate glaber;
folia opposita, petiolis ad 4.5 cm. longis, laminis lanceolatis vel
ellipticis ad 20 cm. longis et 7 cm. latis, basi inaequilateraliter rotunda-
tis vel obtusis vel cuneatis, apice acutis vel attenuatis, integris vel
dentibus 1 vel 2 per cm. crenatis vel serratis, supra puberulis vel
elabris, subtus in nervis dense pubescentibus alibi subglabratis,
venis primarlis utrinsecus 6-8 curvatis, rete venularum conspicuo;
inflorescentiae pilis patentibus dense pubescentes, pendunculis 6-12
mm. longis, bracteis terminalibus lanceolatis 4 mm. longis caducis,
pedicellis 2 vel 3 et 5-12 mm. longis; calyx caducus ca. 8 mm. longus
in lobis 5 aequalibus ovato-lanceolatis acuminatis 0.5 fissus extus
ecapitato-pilosus intus glaber; corolla pallide lutea infundibuliformis
ca. 1.8 cm. longa in lobis inaequalibus rotundatis ca. 0.25 fissa, extus
capitato-glanduloso-pilosa, intus tubo glabra limbo capitato-glandu-
loso-pilosa; filamenta ca. 3 mm. longa, antheris in apice adhaerentibus
ca. 1 mm. infra sinus corollae; staminodia 3, ca. 0.75 mm. longa prope
filamenta enata, staminodio medio ca. 1 mm. supra staminodia
lateralia; discus annulari-cupuliformis conspicuus ca. 1 mm. altus
sub fructu fragmentibus persistens; gynoecium ca. 12 mm. longum,
ovario glabro, stylo albo ubique capitato-piloso demum 1-2 mm.
supra basim caduco, stigmate applanato bilobato ca. 3 mm. infra
antheras; fructu albo ca. 1 em. longi et 0.6 cm. lati.

Type in the herbarium of the Arnold Arboretum, collected on the
southern slopes of Mount Numbuiloa, east of Lambasa, Province of
Mathuata, Vanua Levu, Fiji, alt. 100-350 m., Oct. 27, 1947, by
A. C. Smith (6343). Duplicates at BISH, US.

140 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

ADDITIONAL SPECIMENS EXAMINED:

Vanua Levu: Mathuata: Same locality as type, Smith 6529 (A, BISH, US).
Thakaundrove: Eastern drainage of Yanawai River, Degener & Ordonez 14088
(A, BISH, NY, UC, US).

The new species bears a superficial resemblance to Cyrtandra
harveyi but lacks the exserted stamens and style, the long filaments,
the pedestal or “gynophore” at the base of the ovary, and the much
larger flowers of that species. Inflorescence and floral characters
seem to place it near C. involucrata and C. attenuata. | have not seen
other Melanesian or Polynesian Cyrtandrae that resemble the new
species.

21. Cyrtandra jugalis A. C. Smith, Journ. Arnold Arb. 34: 49. 1953.

Shrub or slender tree 2-5 m. high, densely pubescent on younger
parts with septate, uniseriate, light to dark brown, noncapitate hairs
ca. 30u in diameter and 0.5 mm. long; leaves opposite, the petioles
1-2 em. long, the blades lanceolate to oblanceolate to ovate, up to
16 cm. long and 5 cm. wide, unequal and acute to cuneate at base,
acute to acuminate at apex, undulate to serrate, with 2-3 teeth per
em. at margin, glabrous above, with dense tomentum of noncapitate
hairs beneath, these more pronounced on the veins, the primary veins
raised, 5-7 on each side of midvein, the secondary veins inconspicu-
ously reticulate; inflorescence a simple dichasium, tomentose to
glabrate, the peduncles 1-5 em. long, terminated by a pair of deciduous
lanceolate, free bracts ca. 0.5 cm. long, subtending pedicels 1-2.5 em.
long; calyx fleshy, drying subcoriaceous, ca. 1 cm. long, cleft ca. 0.33
of its length into 5 equal, triangular-attenuate lobes, glabrate on both
surfaces; corolla white, ca. 2 cm. long, cleft ca. 0.33 of its length into
unequal, rounded lobes, with dense, ascending, noncapitate hairs
without, with capitate hairs within; filaments ca. 2 mm. long, adnate
to corolla tube opposite lower sinuses, bearing apically coherent anthers
at the base of the corolla limb; staminodes 3, adnate opposite upper
sinuses at same level as filaments, the median staminode 1 mm. long,
slender, the lateral 2 mm. long, clavate; cupulate annular dise promi-
nent, 1 mm. high, persisting as rectangular fragments in mature fruit;
ovary and style up to 10 mm. long, the ovary glabrous, the style with
capitate hairs, separating ca. 1 mm. above apex of mature fruit, the
stigma applanate, bilobed; fruit white, ellipsoid, ca. 1.2 em. long and
0.8 cm. wide.

Type Locauity: Ridge between Mount Nanggaranambuluta and
Mount Namama, east of Nandarivatu, Viti Levu. Type collected by
A. C. Smith, cited below.

GILLETT—CYRTANDRA IN FIJI 141

LocaL NAME: “Mindra” (Smith 4994); “Mbetambeta” (Degener
14904).

Distripution: Dense forests in the highlands of central Viti Levu,
in the vicinity of Mount Tomanivi and the mountains west of Nan-
darivatu, between 700 and 1,150 meters.

Vitt Levu: Mba: Nambuisa Village, Gillespie 4134 (BISH, UC); ridge between
Mount Nanggaranambuluta and Mount Namama, east of Nandarivatu, Smith
4994 (A-type, BISH, US); summit of Mount Nanggaranambuluta, Smith 4878
(A, BISH, US); western and southern slopes of Mount Tomanivi, Smith 5319
(A, BISH, US). Nandronga & Navosa: Vicinity of Nandrau, Degener, 14904 (A);
northern portion of Rairaimatuku Plateau, between Nandrau and Nanga, Smith
5569 (A, BISH, US). Ra: Ridge from Mount Namama toward Mount Tomanivi,
Smith 5695 (A, BISH, US), Smith 5696 (A, BISH, US), Smith 5709 (A, BISH,
US), Smith 5715 (A, BISH, US).

The present species appears to be closely related to Cyrtandra
involucrata and C. anthropophagorum; it lacks the connate-perfoliate
bracts of the former and the larger corollas (up to 20 mm. long) and
‘ndument of the latter. However, neitner of the related species have the
simple dichasium inflorescence that so well characterizes Cyrtandra
jugalis. None of the Polynesian species have a close resemblance to
this species.

22. Cyrtandra anthropophagorum Seem. (Bonplandia 9: 257. 1861, nom. nud.) ex
A. Gray, Proc. Amer. Acad. 6: 41. 1862. FI. Vit. 182. 1866.
Cyrtandra buttii Horne ex C. B. Clarke, in DC. Monogr. Phan. 5: 279. 1883.

Shrub or small tree to 4 m. high, the stem up to 6 cm. diameter
1.5 m. above the ground, freely branched, the young parts covered
with golden-brown, velutinous, septate, uniseriate, noncapitate hairs
20-604 in diameter and up to 1.5 mm. long; leaves opposite, the
petioles 1-5 cm. long, the blades ovate-lanceolate, at maturity up to
30 em. long and 12 cm. wide, unequal and acute, attenuate, or cuneate
at base, acute to acuminate at apex, crenate to serrate, with about 3
teeth per cm., tomentose above, with appressed-ascending hairs
parallel to the primary veins, densely tomentose beneath with spread-
ing hairs, the primary veins 8-12 on each side of midvein, upwardly
curved, the secondary veins concealed by pubescence; inflorescence
densely tomentose, the peduncles 1-30 mm. long, terminated by a pair
of lanceolate, free, deciduous bracts ca. 1.5 cm. long, subtending 1-4
pedicels 1-3 cm. long; calyx green to dull white, 5-7 mm. long, cleft
half its length into 5 unequal, lanceolate, acute to acuminate lobes,
both surfaces tomentose; corolla white, 12-15 mm. long, the tube
cylindric, the lobes 4-5 mm. long, unequal, rounded, tomentose with
noncapitate hairs without, and short, capitate hairs within; filaments
ca. 2 mm. long, bearing apically coherent anthers slightly below the

142 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

corolla limb; staminodes 3, adnate opposite the upper sinuses near the
middle of corolla tube, the lateral staminodes nearly 1 mm. long, the
median ca. 0.25 mm. long; cupulate annular dise conspicuous, ca. |
mm. high, drying to deciduous, rectangular segments; ovary and style
ca. 8 mm. long, the ovary glabrous, the style with capitate hairs on
the upper 0.75 of its length, separating ca. 1 mm. above summit of
mature ovary, the stigma applanate, bilobed, borne below the anthers:
fruit white, ovoid, up to 1.5 em. long and | em. wide.

Typr Locaniry: Viti Levu, Fiji. Type collected by Seemann (278).

Loca namu: “Makamakandora”’ (St. John 18926).

Disrrisution: Understory of rain forests in coastal and interior
regions of southeastern Viti Levu and Ovalau, between 30 and 1,100
meters.

Vitt_ Levu: Without locality, Seemann 278 (GH-lectotype, K). Serua: Tlilis
north of Ngaloa, in drainage of Waininggere Creck, Smith 9164 (BISH, Us);
track north of Korovou, St. John 18926 (BISH); Nakavu, Parks 20364, 20365
(BISH, UC, US). Namosi: Naitarandamu Mountain, Gillespie 3366 (BISH, UC),
3084 (A, BISH, NY, UC, US), 3139 (BISH), 3135 (BISH); without locality,
Horne 818 (GH, K-type of Cyrtandra buttii Horne ex C. B. Clarke), Horne 956
(GH, K-syntype of Cyrtandra buttii Horne ex C. B. Clarke); without locality,
Gillespie 2490 (BISH, UC), 2998 (BISH, UC, US), 2848 (BISH), Parks 20262
(BISH, UC), 20262-a, 20268, 20276 (BISH, UC, US); hills north of Wainavin-
drau Creek, between Korombasambasanga Range and Mount Naitarandamu,
Smith 8411 (BISH, US); hills bordering Wainavindrau Creek, in vicinity of
Wainimakutu, Smith 8565 (BISH, US); northern slopes of Korombasambasan ga
Range in drainage of Wainavindrau Creek, Smith 8719 (BISH, US); Namuamua,
Gillespie 2992 (A, BISH); hills east of Wainikoroiluva River, near Namuamua,
Smith 8909 (BISH, US); Navua River, Greenwood 1049 (A). Naitasiri: Nasinu,
9 mi. from Suva, Gillespie 3509 (BISH, UC); Tamavua-Sawani Road, Setchell
Parks 15143 (UC); Nasinu road, 2 mi. from Suva, Gillespie 3450 (BISH, NY, UC,
US). Tailevu: Hills east of Wainimbuka River, in vicinity of Ndakuivuna, Smith
7062, 7110 (BISH, US). Rewa: Suva, Meebold 16725 (BISH), Tothill 640 (A,
BISH), 641, 507 (BISH), MacDaniels 1138 (BISH). Ovatau: Summit of Mount
Ndelaiovalau, Sith 7380 (BISH, US); without locality, undershrub in rain forest,
E. H. Bryan 599 (BISH) without locality, U.S. Kepl. Exped., June 1840 (GH, US).
VaNnvua Levu: Mathuata: Wainikoro, Greenwood 687 (K).

Gray described the flower from the material of the US. Iixploring
Expedition, the young fruit from Seemann 278. The latter is here
designated as the lectotype. This is one of the most frequently col-

lected Cyrtandrae in Fiji. Its relationship is with C. involuerata and
is discussed under that species.

23. Cyrtandra esothrix A.C, Smith, Journ. Arnold Arb. 34:45. 1953.

Shrub or small tree up to 6 m. high, the leaves and inflorescence with
light tomentum of septate, uniseriate, erect, noncapitate hairs ca. 40
uw in diameter and 0.5 mm. long, the young stems quadrangular; leaves

GILLETT—CYRTANDRA IN FIJI 143

opposite, the petioles 0.5 to 2.5 cm. long, the blades lanceolate to ovate,
up to 22 cm. long and 8 cm. wide, unequal and obtuse to acute or
cuneate at base, acute to acuminate at apex, entire to undulate to
serrate, glabrous above, beneath with glabrous to glabrate areoles,
the veins tomentose to glabrous, the primary veins raised, 6-9 per side,
the secondary veins indistinct to conspicuous; inflorescences axillary,
the peduncle up to 1 cm. long, terminated by minute, caducous bracts
subtending 1-4 pedicels up to 1 cm. long; calyx greenish white,
coriaceous, deciduous, up to 1 cm. long, cleft ca 0.25 of its length into
equal, connivent, lobes, with 1 or 2 sinuses lacerated to base, glabrous
without, pilose within, with ascending light brown hairs; corolla
white, cylindrical, ca. 2 cm. long, cleft 0.20 of its length into unequal,
rounded, spreading lobes pubescent without, with capitate and non-
capitate hairs, glabrous within; filaments ca. 2 mm. long, adnate
opposite the sinuses of the anterior corolla lobe and bearing apically
coherent anthers at base of corolla limb; staminodes 3, ca. 0.5 mm.
long, adnate opposite the upper sinuses near middle of corolla tube,
the median staminode ca. 2 mm. below the lateral; cupulate annular
disc conspicuous, ca. 0.75 mm. high, deciduous; ovary and style ca. 1
em. long, the ovary glabrous, the style with capitate hairs, deciduous
ca. 2 mm. above apex of the mature ovary, the stigma applanate,
bilobed, borne ca. 5 mm. below the anthers; fruit white, elliposidal,
up to 1.5 em. long and 1 em. broad.

Type pocatrry: Northern portion of Rairaimatuku Plateau,
Viti Levu, Fiji. Type collected by A. C. Smith, cited below.

Disrripution: Interior mountains of eastern Viti Levu and
Ovalau, in dense forest, 100 to 1,100 m.

Locau NAME: ‘“Makamakandora” (St. John 18185).

Vitt Levu: Mba: Mount Tomanivi, Gillespie 4081 (BISH, GH, NY, UC),
Smith 5098 (A, BISH, US), Smith 5267 (A, BISH, US). Namosi: Naitarandamu
Mountain, Gillespie 3113 (BISH, UC, US). Naitasiri: Rairaimatuku Plateau,
between Mount Tomanivi and Nasonggo, Smith 6134 (A-type, BISH, US);
Wainamo Creek, near Matawailevu, St. John 18185 (BISH, US). Tailevu: Hills
cast of Wainimbuka River, near Ndakuivuna, Smih 7067 (BISH, US), Smith,
7134 (BISH, US), Smith 7179 (BISH, US). Ovatau: Mountains south of Levuka,

on overland trail to the west coast, Gillespie 4538 (BISH, UC); hills east of
Lovoni Valley, Smith 7261 (BISH, US), Smith 7679 (BISH, US).

The significant distinguishing features of this species include the
thick, coriaceous calyx, which is lacerated below the short lobes, and
the pilose interior of the calyx. There is some variability in the in-
dument of the secondary veins of the lower leaf surface. In all collec-
tions from Ovalau and the adjacent territory of Tailevu, Viti Levu,
these veins are glabrous; however, they are tomentose in the collec-
tions from central Viti Levu.

144 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

There appears to be a close morphological relationship between
Cyrtandra esothriz and C. hornet; the two species can be distinguished
by the differences cited in the following tabulation:

C. hornet C. esothriz
Surface of outer/calyx: puberulent glabrous
Surface of inner/corolla: glandular glabrous
Apices of calyx lobes: attenuate acute
Staminodes: 1.5 mm. long 0.5 mm. long

Tn the examination of the Polynesian representatives of the genus,
I have seen no material that would appear to be closely related to
these two species. The bark of Cyrtandra esothrir has been used
medicinally to cause constipation after sickness, according to St. John.

24. Cyrtandra hornei C. B. Clarke, in DC Monogr. Phan. 5: 281. 1883.
Cyrtandra greenwoodiana A. C. Smith, Journ, Arnold Arb, 34: 46. 1953.

Shrub or small tree up to 6 m. high, with rubiginose tomentum of
septate, uniseriate, noncapitate hairs ca. 15p in diameter and 0.1-0.3
mm. long, becoming glabrate to glabrous; leaves opposite, the petioles
0.5-8 cm. long, the blades elliptic to ovate, up to 15 em. long and 6 em.
wide, inequilateral and obtuse to acute or cuneate at base, obtuse to
acute or acuminate at apex, entire to undulate, dentate, or serrate,
glabrous above, tomentose to glabrate beneath, the primary veins 5-9
per side, the secondary veins reticulate, glabrous; inflorescences usually
i-flowered, tomentose to glabrate, the peduncle 0.5-2 cm. long,
terminated by a pair of minute caducous bracts subtending a pedicel
3-15 mm. long; calyx ca. 11-16 mm. cleft ca. 0.33 of its length into
free or connivent, valvate, acuminate lobes, usually lacerated below
2 sinuses, glabrate without, pilose within with ascending noncapitate
hairs; corolia white, ca. 2-2.5 em. long, cylindrical, cleft ca. 0.25 of its
length into rounded, unequal, spreading lobes, glabrous without, with
rapitate hairs within; filaments ca. 2-3 mm. long, adnate opposite
sinuses of anterior corolla lobe, bearing apically coherent anthers ca.
1 mm. below the corolla limb; staminodes 8, ca. 1.5 mm. long, adnate
opposite posterior sinuses at middle of tube; cupulate annular disc
conspicuous, 1 mm. high, deciduous; ovary and style ca. 8 mm. long,
the ovary glabrous, the style with capitate hairs, separating 1-2 mm.
above apex of mature ovary, the stigma applanate, bilobed, borne 6-7
mm. below the anthers; fruit white, ellipsoid, up to 25 mm. long and
8mm. broad.

Typy Locauiry: Fiji, without further locality. Type collected by
Horne, cited below.

Disrripution: Dense forest, interior mountains of central and
northern Viti Levu, between 700 and 1,100 m.

GILLETT—CYRTANDRA IN FIJI 145

Locau NAME: ‘‘Mimila” (Smith 4768).

Fry1: Without further locality, Horne 449 (K-lectotype), Horne 570-2 (GH,
K). Horne 713 (GH).

Vitt Levu: Mba: Between Mount Vatuyanitu and Mount Natondra, Mount
Evans Range, Smith 4304 (A, BISH, US); Mount Nairosa, eastern flank of
Mount Evans Range, Smith 4412 (A-type of Cyrtandra greenwoodiana A. C.
Smith., BISH, US); Mount Evans Range, Greenwood 1260 (A, US), 871-a (A);
western slopes of Mount Nanggaranambuluta, Smith 4768 (A, BISH, US); Mount
Nanggaranambuluta, Greenwood 871 (A, UC, US); Gillespie 3684 (BISH, UC);
near Nandarivatu, Gibbs 572 (BM); Degener 14622 (A, BISH, NY, US), Gillespie
3385 (BISH, NY, UC, US), Gillespie 4038 (BISH, UC); Mount Matomba,
Nandala, near Nandarivatu, Degener 14512 (A, BISH, NY, UC, US), 14833 (A,
BISH, NY, UC, US). Namosi: Voma Mountain, Gillespie 2890.1 (UC).

Clarke’s description is based on Horne 449, 570, and 1134 (not seen),
all at Kew, of which Horne 449 is here designated as lectotype.

The relationship of this species is surely with Cyrtandra esothriz,
and a tabular contrast of the two species is given under the latter.
There is considerable intergradation in the leaves of these two species,
and I know of no reliable means of distinguishing material in which
flowers are not present.

25. Cyrtandra tavinunensis Gillespie, Bishop Mus. Bull. 74: 24, fig. 33, 1930.

Glabrous, slender shrub or small tree 3-4 m. high, the young stems
with a flaky surface when dried; leaves opposite, the petioles 0.5-2 cm.
long, the blades lanceolate to ovate, up to 20 cm. long and 6 cm. broad,
equilateral and acute to attenuate at base, acute to acuminate at apex,
undulate, glabrous above and beneath, with 5-7 upwardly curved
primary veins per side, the secondary veins faintly to conspicuously
reticulate beneath; inflorescences elongate, filiform, one-flowered, the
peduncles up to 6 em. long, bearing a pair of ovate, foliaceous bracts
10-12 mm. long, and 1-4 pairs of bracteoles, the terminal bracteoles
subtending a pedicel ca. 3 cm. long; calyx ca. 8 mm. long, cleft ca.
0.33 of its length into 3-5 ovate, valvate lobes, glabrous on both sur-
faces; corolla greenish white, ca. 1 cm. long, cleft ca. 0.33 of its length
into unequal lobes, the exterior glabrous, the interior with dense
indument of papillate hairs ca. 100u in diameter; filaments ca. 3 mm.
long, adnate opposite the sinuses of the anterior corolla lobe ca. 5 mm.
below the limb, bearing apically coherent anthers near the base of the
limb; staminodes 2, adnate ca. 5 mm. below the limb; cupulate an-
nular disc prominent, ca. 1 mm. high at anthesis, deciduous from
maturing fruit: ovary and style glabrous, ca. 7 mm. long, the style
separating near summit of the fruit; stigma applanate, bilobed, the
lobes ca. 1.5 mm. long, ovate, borne below the anthers; fruit bright
red, ovoid, up to 1.5 cm. long and 1 cm. broad.

269-812 674

146 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Type LocaLity: Taveuni. Type collected by Gillespie, cited below.

DistrisutioNn: Dense upland forests of Taveuni, 700-850 m.

TaveEun!I: Trail from Somosomo, Gillespie 4782 (BISH-type, NY, UC, US);
summit and adjacent slopes of Mount Manuka, east of Wairiki, Smith 8234
(BISH, US).

The flaky surface of the young stems provides the most useful
feature to distinguish Cyrtandra taviunensis from the closely related
C. montana. This peculiar feature is not known in other Fijian species
but does occur elsewhere in the genus, particularly in Malaysian
species.

Further discussion of the relationship between this species and
Cyrtandra montana is presented under the latter.

26. Cyrtandra montana Gillespie Bishop Mus. Bull. 74:22, fig. 29. 1939.
Cyrtandra bracteolosa A. C. Smith, Journ. Arnold Arb. 34:43. 1953.

Erect, glabrous shrub 2-5 m. high, the young foliage with indument
of scattered, papillate hairs ca. 40 in diameter, becoming glabrous
at maturity; leaves opposite, the petioles 0.5-2.5 cm. long, the blades
lanceolate to elliptic, up to 23 cm. long and 6 cm. broad, obtuse to
acute to cuneate at base, acute to acuminate at apex, undulate to
dentate, with 1-2 teeth per cm., glabrous above and beneath, 6-9
upwardly curved primary veins per side, the secondary veins faintly
to conspicuously reticulate below; inflorescences filiform, glabrous,
branching cymes, the peduncles ca. 1-3.5 cm. long, bearing a pair
of deciduous, linear bracts 2-7 mm. long, the branches with 1-3 pairs
of bracteoles, the terminal bracteoles subtending pedicels 0.5-3 cm.
long; calyx 4-8 mm. long, glabrous on both surfaces, cleft ca. 0.5 of
its length into ovate-acuminate, valvate lobes, the base slightly
inflated; corolla white, broadly funnelform, up to 1.6 cm. long,
cleft ca. 0.5 of its length into unequal, acute lobes, glabrous without,
the limb within with indument of papillate hairs ca. 100y in diameter,
the throat within with short, capitate hairs ca. 30% in diameter;
filaments ca. 1.5 mm. long, bearing apically coherent anthers 1-2
mm. below the corolla limb; staminodes 3, ca. 0.5 mm. long, adnate
ca. 3 mm. below the limb, the median staminode 1 mm. below the
lateral; cupulate annular disc conspicuous, ca. 1 mm. high, deciduous;
ovary and style glabrous, 5 mm. long, the style separating ca. 1 mm.
above apex of mature fruit; stigma applanate, bilobed, borne ca.
2 mm. below the anthers; fruit orange to red, ovoid, up to 1.5 cm.
long and 1 cm. broad.

TypE LocaLity: Wooded ridge on Naitarandamu Mountain, 1,100
m. Type collected by Gillespie, cited below.

DistrisBuTtion: Dense forests of Mba and Namosi, central Viti
Levu, between 250 and 1,100 m.

GILLETT—CYRTANDRA IN FIJI 147

Vitt Levu: Mba: Hills east of Nandala Creek, ca. 3 mi. south of Nandarivatu,
Smith 6232 (A-type of Cyrtandra bracteolosa A. C. Smith, BISH, US). Namosi:
Mount Naitarandamu, Gillespie 3144 (BISH-type, UC); Valley of Wainambua
Creek, south of Mount Naitarandamu, Smith 8833 (BISH, US); hills north of
Wainavindrau Creek, between Korombasambasanga Range and Mount
Naitarandamu, Smith 8504 (BISH, US); northern slopes of Korombasambasanga
Range, in drainage of Wainavindrau Creek, Smith 8732 (BISH, US).

A close examination of the types of Cyrtandra montana and C.
bracteolosa has failed to reveal any differences that would justify the
recognition of the latter as a distinct species. Smith (1953) indicated
differences in the length of the calyx (4 to 5 mm. long in C. bracteolosa
and 4 to 8 mm. long in C. montana), but a quantitative difference of
this magnitude would be expected within a given species. The types
were collected from the same general territory in the uplands of
central Viti Levu.

The close relationship of Cyrtandra montana and C. taviunensis
seems clearly indicated in their striking similarities in inflorescence,
indument, flower structure, fruit size, fruit color, and other features.
This lineage apparently does not extend into Polynesia, and there
are no other currently known Fijian species that appear close to this
remarkably distinctive pair. It seems reasonable to speculate that the
two species were isolated by divergent, long-distance dispersal,
probably by frugivorous birds attracted to the brightly colored fruits.
27. Cyrtandra pritchardii Seem. in Bomplandia 9: 257 (nomen), 364 (desc.).

1861; Fl. Vit. ¢. 39. 18664
Cyrtandra coriacea C. B. Clarke, in DC. Monogr. Phan. 5: 280. 1883.
Cytandra gracilipes Gillespie, Bishop Mus. Bull. 74: 21, fig. 27. 1930.

Glabrous, freely branched shrub or small tree up to 5 m. high;
leaves opposite, the petioles up to 2.5 cm. long, the blades lanceolate
to elliptic, up to 18 cm. long and 5.5 cm. broad, equal to unequal and
acute to cuneate at base, acute to acuminate at apex, undulate or
entire, glabrous above and beneath, the primary veins obscure above,
conspicious beneath, 8-12 per side, the secondary veins indistinct;
inflorescences 1-3 flowered, the peduncles 0.5-1 em. long on older
branches, up to 2.5 cm. long on young shoots, terminated by a pair
of caducous bracts ca. 0.5 mm. long, bearing 1-3 pedicels up to 2 cm.
long; calyx coriaceous, 5-7 mm. long, cleft 0.33 of its length into 5
equal, valvate, lanceolate lobes, glabrous without, with capitate hairs
within; corolla white, 1.5-1.8 cm. long, cleft ca. 0.33 of its length into
unequal, rounded lobes, glabrous without, with capitate hairs within;
filaments ca. 2 mm. long, adnate 3 mm. below the sinuses of the
anterior corolla lobe, bearing apically coherent anthers at the base
of the limb; staminodes 3, ca. 0.5 mm. long, adnate ca. 5 mm. below

4 Also figured in Parham, Pl. Fiji Isl. f. 87. 1964.

148 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

the upper sinuses; cupulate annular disc prominent, ca. 0.5 mm.
high at anthesis, drying to a fragmented, brown disc in mature fruit;
ovary and style 9-12 mm. long, the ovary glabrous, the style with
capitate hairs on upper 0.75 of its length, separating at summit or
2-3 mm. above apex of mature ovary, the stigma applanate, bilobed;
fruit white, ellipsoid to fusiform, up to 2 cm. long and 1.2 em. broad.

Typx Locauity: Ovalau. Type collected by Seemann, cited below.

Distripution: Dense forests of southeastern Viti Levu and Ovalau,
between 50 and 500 m.

Viti Levu: Serua: Hills west of Waivunu Creek, between Ngaloa and Korovou,
Smith 9304 (BISH, US); hills between Waininggere and Waisese Creeks, between
Ngaloa and Wainiyambia, Smith 9546 (BISH, US). Rewa: Summit of Korom-
bamba Mountain, Gillespie 2393 (BISH); southeast slopes of Korombamba
Mountain, Gillespie 2306 (BISH-type of Cyrtandra gracilipes Gillespie, UC),
Gillespie 2221 (BISH, UC), Korombamba Mountain, Meebold 16726 (BISH);
near Suva, Horne 698 (K-type of Cytranda coriacea C. B. Clarke), Parks 20068
(BISH, UC). Ovatau: “Mr. Pritchard’s Estate,” Seemann 283 (GH, K-type);
without locality, U.S. Expl. Exped. (US).

The Ovalau specimens have broadly elliptic leaves, while the Viti
Levu specimens have lanceolate to elliptic leaves; the material from
the two islands seems similar in all other features. With no other
differences to separate them other than slight contrasts in leaf shape,
it seems best to refer Cyrtandra coriacea and C. gracilipes to C. prit-
chardi. This species was the first Cyrtandra to be described from
Fiji; it is well illustrated in pl. 39 of Flora Vitiensis.

None of the Polynesian material I have examined bears any re-
semblance to Cyrtandra pritchardii but very similar material has been
obtained from the Solomon Islands: San Cristoval, Brass 2690; and
Guadalcanal, Kajewski 2711.

28. Cyrtandra denhamii Seem. Fl. Vit. 182. 1866.

Shrub or tree up to 5 m. high with rubiginose, septate, uniseriate,
noncapitate hairs ca. 204 in diameter and 0.25 mm. long; leaves
opposite, glabrate at maturity, the petioles 1-3 em. long, the blades
lanceolate to elliptic, up to 19 cm. long and 6 em. broad, unequal
and obtuse to acute or cuneate at base, acute to acuminate at apex,
undulate to dentate, with 1-3 teeth per cm., the venation conspicuous
beneath, the primary veins 5-7 per side, the secondary veins reticulate;
inflorescences many-flowered, branched, glabrate cymes, the pe-
duncles 4-9 cm. long, terminated by a pair of deciduous, lanceolate
to ovate bracts up to 1.5 cm. long, subtending braceteolate pedicels
0.5-3 cm. long, the bracteoles lanceolate, up to 6 mm. long; calyx
white, deciduous, up to 1.5 em. long, cleft nearly to base into 5 un-
equal, lanceolate, acuminate, valvate lobes, at maturity glabrate
without, glabrous within; corolla white, 1 cm. long, cleft ca. 0.25 of

GILLETT—CYRTANDRA IN FIJI 149

its length into unequal, rounded lobes, with capitate hairs ca. 404
in diameter and up to 0.25 mm. long on both surfaces; filaments ca.
1 mm. long, bearing apically coherent anthers at the corolla sinuses;
staminodes 3, 0.25 mm. long, adnate opposite the upper sinuses ca.
2 mm. below corolla throat, the median staminode about 0.5 mm.
above the lateral; cupulate annular disc conspicuous, ca. 0.75 mm.
high, drying to irregular rectangular fragments, persistent in mature
fruit; ovary and style ca. 5 mm. long, the ovary glabrous, abruptly
tapered to the style, the style with dense indument of capitate hairs,
the stigma capitate, shallowly bilobed, the style separating 2-5 mm.
above apex of mature fruit; fruit white, ellipsoid, up to 2.5 cm. long
and 1.3 cm. wide.

Type Locatiry: Mountains of Ngau, Fiji. Type collected by
Milne, cited below.

DistriputTion: Known only from the upland forests of Ngau, 300
to 450 m.

Neau: “Mountains,” Milne 236 (K-type); hills east of Herald Bay, inland
from Sawaieke, Smith 7755 (BISH, US).

The type includes two specimens of which the lower specimen was
annotated by Gray as a Clerodendron. However, my examination of
the type reveals that the lower specimen is indeed Cyrtandra denhamu,
as had been indicated by Seemann.

The deeply cleft calyx and sparse indument of the mature foliage
characterize both Cyrtandra denhamii and C. tempestii. The eastward
extension of this group into Polynesia is indicated by C. feanrana
F. Brown, of the Marquesas Islands, and C. longipedunculata Rech.,
of Samoa, both of which have the inflorescence and flower characters
of OC. denhamii. The capitate, shallowly bilobed stigma of C. denhami
is unique among Fijian species of Cyrtandra, and also occurs in
C. feaniana, cited above.

29. Cyrtandra tempestii Horne ex C. B. Clarke, in DC. Monogr. Phan. 5: 273.
1883.

Compact shrub or slender tree 3-4 m. high, with rubiginose
tomentum on the younger parts, consisting of small, septate, uniseri-
ate, noncapitate hairs ca. 154 in diameter and 0.2 mm. long; leaves
opposite, the petioles 5-6 cm. long, the blades ovate, up to 20 cm.
long and 10 cm. broad, oblique and acute to obtuse at base, acute at
apex, undulate to finely serrate at margin, at maturity glabrate
above, glabrate to tomentose-sericeous beneath, the primary veins
ca. 8-10 per side, the secondary veins reticulate; inflorescences many-
flowered, branched cymes, the peduncles 0.5-1.5 cm. long, termi-
nated by a pair of minute, caducous bracts subtending 2-4 branches,
the pedicels 1-2 cm. long; calyx white, ca. 1 cm. long, cleft nearly to
base into 5 equal, linear-lanceolate, valvate lobes, puberulent with

150 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

conspicuous, dark brown, noncapitate hairs without, glabrous within;
corolla funnelform, white, ca. 2 cm. long, cleft 0.25 of its length into
5 unequal, reinform lobes to ca. 5 mm. broad, glabrous without, with
capitate hairs within; filaments ca. 2 mm. long, adnate opposite the
sinuses of the anterior corolla lobe, bearing apically coherent anthers
ca. 3 mm. below the corolla limb; staminodes 3, 1-2 mm. long, adnate
1-2 mm. below filaments and opposite the posterior sinuses; cupulate
annular dise conspicuous, ca. 1.5 mm. high, drying to a pelviform
structure, persistent in mature fruit; ovary and style ca. 8 mm. long,
glabrous, the style separating at the apex of the mature ovary, the
stigma applanate, bilobed, borne ca. 4 mm. below the anthers; mature
fruit white, ellipsoid, up to 1.6 cm. long and 1 cm. broad.

TypE Locauity: Taveuni, “in woods near the sea at Na Seli
Levie.” Type collected by Horne, cited below.

DistrisuTion: Known only from two localities on Taveuni,
where it occurs in dense forest at elevations from “near the sea”
(Horne 1136) to 900 m.

Taveuni: East of Somosomo, near old crater occupied by small swamp and
lake, Smith 8388 (BISH, US); Na Seli Levie, Horne 1136 (GH, K-type).

This species is closely related to Cyrtandra denhamii both in
morphology and dispersal potential, for each species is of a lineage
that has established populations in eastern Polynesia. The Polynesian
populations related to C. tempestit occur on Rarotonga in the Cook
Islands. These have the habit, leaves, inflorescence, calyx, and
nectiferous annular dise of C. tempestii. The Cook Islands populations
are currently recognized as a distinct species, C. rarotongensis Cheesm.

30. Cyrtandra ciliata Seem. (Bonplandia 9: 257. 1861, nom. nud.) FI. Vit. 182.
pl. 41. 1866.

Shrub to slender tree up to 4 m. high, the foliage with indument of
septate, uniseriate noncapitate hairs ca. 304 in diameter and up to 1
mm. long; leaves opposite, the petioles 2-6 cm. long, tomentose, the
blades lanceolate to lanceolate-ovate or ovate, up to 28 cm. long and
12 cm. broad, oblique to inequilateral and rounded to acute or cuneate
at base, acute to acuminate at apex, undulate to coarsely serrate, with
ca. 2 teeth per cm., glabrous above, beneath glabrate to glabrous on
the areoles, tomentose on the veins and margins, the primary veins
7 or 8 per side, the secondary veins reticulate below; inflorescences
elongate, branching, many-flowered cymes with indument of non-
capitate hairs ca. 20g in diameter and 0.2 mm. long, the peduncles
0.5-5 cm. long, shorter in the inflorescences borne on older stems,
terminated by caducous, linear bracts 1-2 mm. long, subtending
bracteolate branches, the pedicels 1-5 cm. long; calyx green to white,
deciduous, ca. 4 mm. long, cleft nearly to base into 5 equal, linear-
lanceolate lobes with rounded apices, puberulent without, glabrous

GILLETT—CYRTANDRA IN FIJI 151

within; corolla white, funnelform, ca. 2-2.5 cm. long, cleft 3-4 mm.
into unequal, rounded lobes, with numerous capitate hairs without or
glabrous, glabrous within; filaments ca. 3 mm. long, bearing apically
coherent anthers at the base of the corolla limb; staminodes 3, ca. 1
mm. long, adnate ca. 4 mm. below the upper sinuses, the median
staminode borne at same level as the lateral; cupulate annular disc
prominent, ca. 1 mm. high, deciduous from the mature fruit; ovary
and style ca. 1.5 em. long, the ovary glabrous, the style with capitate
hairs in upper half of its length, separating 1-4 mm. above the rounded
apex of the mature ovary, the stigma applanate, shallowly bilobed,
the lobes thick, spreading vertically; fruit white, ovoid, up to 12 mm.
long and 7 mm. broad.

Type LocALITy: Vuna, southwest Taveuni, Fiji. Type collected
by Seemann, cited below.

Locau NAME: “Muskarimba” (Smith 1979).

DistriBuTION: Vanua Levu, Taveuni, and Koro, in dense forest,
between 300 and 900 m.

Koro: Eastern slope of main ridge, Smith 965 (BISH, GH, NY, US). Vanua
Levu: Thakaundrove: Mount Mariko, Smith 435 (BISH, GH, NY, UC, US);
Mount Ndikeva, Smith 1863 (BISH, GH, NY, UC, US); Natewa Peninsula,
Uluingala, Smith 1979 (BISH, GH, NY, UC, US). Taveuni: Vuna, Seemann 282
(GH, K-type) ; between Somosomo and Wairiki, Smith 765 (BISH, GH, NY, US);
Mount Manuka, east of Wairiki, Smith 8207 (BISH, US) ; without further locality,
Gillespie 4724 (BISH, NY, UC).

The excellent plate in Flora Vitiensis has greatly facilitated identi-
fication, making this one of the most easily determined of the Fijian
species. The affinities of Cyrtandra ciliata are not known. None of the
other Fijian species have comparable flowers and inflorescences, and
none of the Polynesian species are similar. It therefore seems likely
that this species represents a lineage that is either peculiar to Fiji, or
attenuated from Melanesian territory to the west, from which I have
been able to examine very little material.

The variability in corolla indument is notable, for the populations
from Vanua Levu are characterized by corollas with an outer indument
of capitate hairs, while populations from Taveuni have corollas with
a glabrous outer surface; however, the plants seem similar in other
features. This difference is indicative of the propensity for evolutionary
change in isolated populations of Cyrtandra. In such cases one has
the alternative of either: circumscribing species to reasonably broad
limits, or giving formal recognition to a number of separate, isolated
populations, each distinguished by a slight morphological expression,
such as the above.

31. Cyrtandra xanthantha A. C. Smith, Journ. Arnold Arb. 34: 48. 1953.

Shrub 3-4 m. high with ferrugineous, septate, uniseriate, noncapitate
hairs ca. 304 in diameter and to 1 mm. long; leaves opposite, the

152 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

petioles 1-3 cm. long, tomentose, the blades lanceolate to elliptic, up
to 16 cm. long and 6 cm. broad, inequilateral and acute or cuneate at
base, acute to acuminate at apex, entire to serrate, the serrations 1-2
per cm., papillate-puberulent to tomentose above, beneath densely
tomentose with raised primary veins 6-8 per side and curved upwardly,
the secondary veins reticulate beneath; inflorescences ferrugineous-
tomentose, the peduncles 0.5-3 cm. long terminated by a pair of
caducous, ovate-acuminate, connate bracts up to 1.7 cm. long and
0.8 cm. broad, the bracts villous above and below, subtending pedicels
0.5-1.5 cm. long; calyx deciduous, white to pale green, ca. 1 em.
long, cleft ca. 0.66-0.75 of its length into equal, lanceolate, valvate
lobes, the tube and lobes villous without, glabrous within; corolla
white to pale yellow, curved-cylindrical, up to 2.5 cm. long, cleft ea.
0.33 of its length into unequal, rounded, spreading lobes, the tube and
lobes with capitate hairs without, glabrous within; filaments 2-3 mm.
long, bearing apically coherent anthers at the base of the corolla limb:
staminodes 2, ca. 1.5 mm. long, adnate near the middle of corolla
tube; cupulate annular disc conspicuous, ca. 1 mm. high with irregular
margin, deciduous in mature fruit; ovary and style 8-9 mm. long, the
ovary glabrous, the style with capitate hairs in the upper half of its
length, separating 1-2 mm. above the rounded apex of the mature
ovary, the stigma applanate, shallowly bilobed, the lobes thick,
spreading vertically, borne ca. 3mm. below the anthers; fruit ellipsoid,
up to 1.8 em. long and 1 em. broad.

Typr Loca.ity: Slopes of the escarpment north of Nandarivatu,
Fiji. Type collected by A. C. Smith, cited below.

Disrrisution: Interior of Viti Levu in upland rain forest, 500-925 m.

Vitt Levu: Mba: Mount Evans Greenwood 376 (K); slopes of escarpment
north of Nandarivatu, Smith 6277 (A-type, BISH, US); near Nandarivatu,
Gillespie 4150 (A, BISH); valley of Singatoka River, near Nandarivatu, Gillespie
3979 (A, BISH).

The affinity of this species is with Cyrtandra aloisiana, and these
two species do not appear to be closely related to other Fijian or
Polynesian species. A tabular contrast of C. ranthantha and C. eloisiana
is offered below:

C. xanthantha C. aloisiana
Habitat: Montane inland rain Lowland coastal rain
forests 550-925 m. forests below 150 m.
Length of calyx lobes: Lobes 2-3 times length Lobes 4-5 times length
of calyx tube. of calyx tube.
Inflorescence bracts: Only outer bracts Outer and inner bracts
present. present.
Length of corolla: Corolla up to 25 mm. Corolla up to 10 mm.
long. long.
Hairs of the petiole: 0.25-1.0 mm. long. 2-7 mm. long.

Staminodes 2, ca. 1.5 mm. long. 3, ca. 0.5 mm. long.

GILLETT—CYRTANDRA IN FIJI 153

32. Cyrtandra aloisiana A. C. Smith, Sargentia 1: 116, 1942.

Shrub about 1 m. high with villous, rubiginose, septate, uniseriate
noncapitate hairs ca. 604 in diameter and up to 7 mm. long; leaves
opposite, the petioles 1-5 cm. long, densely villous, the blades lanceo-
late to elliptic to ovate, up to 22 cm. long and 8 cm. broad, inequilateral
and acute or cuneate at base, obtuse to acute or acuminate at apex,
entire to faintly serrate, sparsely villous on the upper surface, beneath
with villous pubescence and raised primary and secondary veins, the
primary veins 6 or 7 per side; inflorescences congested, with 2-8
flowers, the peduncles 3-12 mm. long, terminated by a pair of cadu-
cous, lanceolate-ovate, acuminate, villous bracts up to 15 mm. long
and 7 mm. broad, the bracts subtending bracteolate pedicels ca. 10
mm. long, the bracteoles up to ca. 7 mm. long and 3 mm. broad,
deciduous, villous on both surfaces; calyx ca. 6 mm. long, deciduous,
cleft nearly to base into 5 equal, lanceolate, acuminate lobes, these
often connivent, villous without, glabrous within; corolla pale yellow,
cylindrical, ca. 1 cm. long, cleft 2-3 mm. into unequal, erect, rounded
lobes, the tube and lobes with capitate hairs without, glabrous within ;
filaments adnate ca. 2 mm. below the corolla throat, opposite sinuses
of the anterior lobe, bearing apically coherent anthers at the base of
the corolla limb; staminodes 3, ca. 0.5 mm. long, adnate near middle
of the corolla tube; cupulate annular disc ca. 0.5 mm. high, conspicu-
ous, with irregular margin, falling from the maturing fruit; ovary
and style ca. 5 mm. long, the ovary glabrous, the style with indument
of capitate hairs on upper 0.25 of its length, separating 2-3 mm. above
the apex of the mature ovary, the stigma applanate, shallowly bilobed,
the lobes thick, spreading vertically, borne 2-3 mm. below the anthers;
fruit white, ellipsoid, 12-16 mm. long.

Type tocauity: Near Ngaloa, Serua, Viti Levu, Fiji. Type collected
by Degener, cited below.

Disrripution: Forested slopes near the ocean, south coast of Viti
Levu, 0-150 m.

Loca name: ‘“Makamakandora” (Degener 15105); ‘“‘Soronim-
bengga”’ (St. John 18952).

Vitt Levu: Nandronga & Navosa: Mbelo, near Vatukarasa, Tabualewa
15619 (A, BISH, UC, US); track north of Komave, St. John 18952 (BISH, US).
Serua: Thulanuku, near Ngaloa, Degener 15105 (A-type, BISH, NY, UC, US).

This species and the closely related Cyrtandra zanthantha have no
relationships to the Cyrtandrae of Polynesia insofar as I have been
able to determine. The contrasts between C. zanthantha and C.
aloisiana are presented under the former species.

154 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

33. Cyrtandra harveyi Seem. Fl. Vit. 182. 1866.
Cyrtandra gorriei Horne ex C. B. Clarke, in DC. Monogr. Phan. 5: 273. 1883.

Shrub 1-4 m. high with perfect or unisexual flowers and dark brown,
septate, uniseriate, noncapitate hairs ca. 204 in diameter and up to
0.5 mm. long, the pubescence dense and closely appressed on younger
parts, sparse on mature foilage; leaves opposite, the petioles 2-9 cm.
long, the blades lanceolate to ovate, up to 25 cm. long and 9 em. wide,
unequal and rounded to acute or cuneate at base, acute to acuminate
at apex, serrate, above with appressed, upwardly directed parallel
hairs, beneath with moderate to dense pubescence of spreading hairs;
inflorescences bearing 1-3 flowers, the peduncles 3-15 mm, long,
pubescent, very short and slender on older stems, longer and thicker
on young shoots, terminated by caducous, lanceolate bracts 2—3 mm.
long subtending pedicels 5-12 mm. long; calyx caducous, greenish
white, fleshy, drying subcoriaceous, ca. 7 mm. long, cleft nearly half
its length into 5 ovate to lanceolate, acuminate, free or connivent lobes,
with 1 or more of the sinuses often lacerated, the tube and lobes with
scattered, conspicuous, dark brown hairs without, glabrous within;
corolla white, 2.5-3 cm. long, funnelform, slightly curved, and cleft
ca. 0.125 of its length into rounded, unequal lobes, the tube and lobes
with conspicuous capitate hairs ca. 30u in diameter and 0.33 mm.
long without, glabrous within; filaments of bisexual flowers ca. 8 mm.
long and adnate 3 mm. below the sinuses of the anterior corolla lobe,
bearing 2 apically coherent anthers in the broad corolla throat; stami-
nodes 3, 1 mm. long, adnate ca. 7 mm. below the posterior sinuses,
the median staminode borne ca. 1 mm. above the laterals ; cupulate
annular disc prominent, ca. 1 mm. high clasping a stalk, or gynophore
ca. 1.5 mm. long, this expanding upward to the base of the ovary;
ovary and style 1.7-2 cm. long, the ovary glabrous, the style with
short, capitate hairs over its entire length, exserted 2-3 mm. above the
corolla throat, separating 1-5 mm. above the apex of the mature
ovary, the stigma applanate, shallowly bilobed; fruit white, ellipsoid,
ca. 2 cm. long and 1 cm wide.

In female flowers (Smith 367, all cited specimens) both stamens and
staminodes are absent and the style and ovary are 2—2.3 cm. long.
The style is 1 mm. in diameter, much thicker than in bisexual flowers,
and is exserted 3-4 mm. above the throat of the corolla.

Type Locauity: Nandi, Mbua, Vanua Levu, Fiji. Type collected
by Harvey, cited below.

Loca NAME: “Merikula” (Smith 367).

DistriBution: Dense rain forests of Vanua Levu, between 100 and
400 meters.

Vanua Levu: Mbua: Nandi, W. H. Harvey, s.n. (GH, K-type; Sandalwood
Bay, U.S. Explor. Exped., s.n. (GH, US). Mathuata: Southern base of Mathuata

GILLETT—CYRTANDRA IN FIJI 155

Range, north of Natua, Smith 6761 (A, BISH, US); Wainunu-Ndreketi divide,
Smith 1847 (BISH, GH, NY, US). Thakaundrove: Hills east of Mbalanga, Savu
Savu Bay region. Degener & Ordonez 13899 (A, BISH, NY, UC, US); Vatunivu-
amonde Mountain, Savu Savu Bay region, Degener & Ordonez 13969 (A, BISH,
NY, UC, US); southern slope of Valanga Range, Smith 367 (BISH, GH, NY,
UC, US); “mountains at Koro-ni-Saca” (near Nakuku), Horne 546 (K-type of
Cyrtandra gorriei Horne ex C. B. Clarke).

The affinity of Cyrtandra harveyi clearly is with C. coleowdes. Both
species have distinctive morphological features not known in other
Fijian Cyrtandrae. These include the peculiar stalk, or gynophore, at
the base of the ovary, the exserted style, and the long (8-15 mm.),
exposed filaments. The presence of gynodioecious flowers in this
species is unique in Fiji; however, the occurrence of unisexual flowers
in Cyrtandra is known in at least 16 species from New Guinea, 2 from
Borneo, and 1 from Hawaii. Lauterbach placed the unisexual material
in a separate genus, Oyrtandropsis, which Burtt (1936) considers
untenable.

34. Cyrtandra coleoides Seem. (Bonplandia 9: 257. 1861, nom. nud.) FI. Vit. 181,
pl. 40. 1866.5
Cyrtandra alba Gilliespie, Bishop Mus. Bull. 74: 20, fig. 25. 1930.

Erect shrub or small tree 2-4 m. high with pruinose indument on
younger parts, becoming glabrous at maturity; leaves opposite, the
petioles 1-4 cm. long, the blades elliptic to ovate, up to 20 cm. long
and 9 cm. wide, unequal to oblique and rounded to acute or cuneate
at base, acute to rounded at apex, serrulate to undulate or entire, the
primary veins raised beneath, 6-8 on each side of midvein; inflores-
cences borne on older, woody stems, glabrous, the peduncles ca. 1 mm.
long, bearing deciduous bracts less than 0.5 mm. long, the bracts
subtending 1-4 bracteolate pedicels, the bracteoles 0.5 mm. long,
deciduous; calyx ca. 1 em. long, parted or lacerated into irregular
lobes, glabrous on both surfaces; corolla funnelform, ca. 2.5 cm. long,
cleft 3-5 mm. into rounded, unequal lobes, the tube and lobes glabrous
on both surfaces; filaments 10-12 mm. long, adnate 2-3 mm. below
the sinuses of the anterior corolla lobe, bearing 2 apically coherent
anthers above the throat; staminodes 2, 2.5 mm. long, adnate opposite
the lateral sinuses at about the middle of the corolla tube; cupulate
annular disc conspicuous, 1.5 mm. high, deciduous, flanged at base
and constricted at rim, clasping a short stalk or gynophore ca. 1.5
mm. long, this expanding upwardly to the base of the glabrous ovary ;
ovary and style to 2.2 cm. long, at anthesis the ovary scarcely broader
than the style, the upper half of the style with capitate hairs, separat-
ing at summit of the mature ovary, the stigma applanate, shallowly
bilobed, presented above the throat and opposite the upper sinus but

5 Illustrated in Parham, Pl. Fiji Isd. f. 86. 1964.

156 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

slightly below the anthers; fruit white, ellipsoid, up to 18 mm. long
and 9 mm. wide.

Type vocaniry: Namosi, Viti Levu, Fiji. Type collected by
Seemann, cited below.

Locau NAME: ‘“‘Makamakandora” (St. John 18227); “Lembalemba”
(Smith 5628; ‘“Mandiri tambua”’ (Gillespie 2671).

Disrripution: Dense upland rain forests of central and eastern
Viti Levu, 150-1150 m.

Vitt Levu: Mba: Mount Tomanivi, Smith 5259 (A, BISIT, US); hills between
Neggaliwana and Nandala creeks, south of Nauwanga, Smith 5813 (A, BISH,
US). Nandronga & Navosa: Northern portion of Rairaimatuku Plateau, Smith
5628 (A, BISH, US). Namosi: Voma Mountain, Gillespie 2671 (BISH-type of
Cyrtandra alba Gillespie, UC), Gillespie 2924.1 (BISH); Wainavindrau Creek,
near Wainimakutu, Smith 8879 (BISH), US; without further locality, Seemann
280 (GH, K-type). Naitasiri: Wainamo Creek, St. John 18227 (BISH, US).
Rewa: Suva ditch trail, Bryan 378 (A, BISH).

It has not been possible for me to find a morphological basis for
separating Cyrtandra alba from C. coleoides. The types of both species
were collected in the province of Namosi, Viti Levu. The type of
C. alba was collected on Voma Mountain, Namosi. It is known that
Seemann collected on the same mountain so that the locality informa-
tion as well as available morphological comparisons support the
reduction of C. alba to synonymy under C. coleoides. Gillespie’s
illustration of C. alba incorrectly portrays the calyx with 5 equal lobes.
The type material clearly has the calyx lacerated into irregular
segments as in C. coleoides, and a pencil sketch attached to the type
of C. alba, obviously the basis for the published illustration, portrays
the calyx with lacerated, irregular segments. There is obviously a
close relationship between this species and Cyrtandra harveyt. The
morphological comparisons of these two species are presented under
the latter.

35. Cyrtandra prattii Gillespic, Bishop Mus. Bull. 74: 24, fig. 32. 1930.

Erect shrub up to 3 m. high, the ligneous branches to ca. 2 cm. in
diameter, the pith scanty, up to 1.5 mm. in diameter, the young foliage
tomentose with septate, uniseriate, noncapitate hairs ca. 50u in
diameter and 0.75 mm. long, becoming glabrate to glabrous at matur-
ity; leaves opposite, the petioles 1-3.5 em. long, the blades lanceolate
to elliptic, up to 20 cm. long and 8 cm. broad, acute to cuneate at
base, acute to acuminate at apex, faintly serrate, with 1-2 serrations
per cm., at maturity glabrate to glabrous above, densely pubescent
to glabrate beneath, the primary veins raised beneath, curved upward,
6-9 per side, the secondary veins conspicuously reticulate beneath ;
inflorescences with an elongate, woody axis to 35 cm. long, arising in
the lower part of stem or at ground level among the adventitious

GILLETT—-CYRTANDRA IN FIJI 157

roots, bearing at 1-3 cm. intervals ovate, connate-perfoliate, deciduous
bracts 0.5-1 cm. long, the bracts glabrate without, pilose within,
subtending tomentose pedicels 3-7 mm. long; calyx ca. 2 cm. long,
cleft ca. half its length into unequal, lanceolate, acuminate, sometimes
connivent lobes, often lacerated nearly to base below the anterior
sinus, densely tomentose on both surfaces; corolla pale yellow,
cylindrical, fleshy, drying coriaceous, ca. 3 cm. long, cleft to 0.33 of
its length into unequal, rounded lobes, the tube and lobes tomentose
without, glabrous within; filaments 2-3 mm. long, bearing apically
coherent anthers ca. 1 mm. below the corolla limb, the anthers black,
apiculate, ca. 2 mm. long; staminodes 2, ca. 0.5 mm. long, adnate
opposite posterior sinuses ca. 5 mm. below the limb; cupulate annular
disc conspicuous, ca. 1.5 mm. high, deciduous in mature fruit; ovary
and style ca. 2 cm. long, densely tomentose with noncapitate hairs
ca. 40 in diameter and up to 1 mm. long, the stigma applanate,
bilobed, the lobes ovate, ca. 4 mm. long, with inner pubescence of
capitate hairs ca. 20u in diameter and ca. 0.1 mm. long, the style
separating ca. 3 mm. above apex of mature ovary; fruit white, ovoid,
ca. 2. cm. long and 1.2 cm. broad.

Typge Locauity: Near summit of Mount Tomanivi, Mba, Viti
Levu, Fiji. Type collected by Gillespie, cited below.

DistrisuTion: Dense upland forests of central Viti Levu, 900-1300
m.
Loca NAME: “ Andalunga” (St. John 18260).

Virt Levu: Mba: Mount Tomanivi, Gillespie 4091 (BISH-type, UC). Naitasiri:
Wainamo-Wainisavulevu Divide, Rarandawai, St. John 18260 (BISH, US).
Fiji, without further locality, Parks 20242 (BISH).

No likeness to this species has been seen in other Fijian Cyrtandrae,
nor in the Polynesian material. The unique inflorescence surely justifies
the assignment of this species to a distinct group among the Fijian
species of Cyrtandra. The closest relationship of Cyrtandra pratti
would appear to be with C. nodosula Schlecht., of New Guinea.

Bibliography

Baker, H. G.
1959. Reproductive methods as factors in speciation in flowering plants.
Cold Spr. Harb. Symp. Quant. Biol. 24: 177-191.
Borrt, B. L.
1936. Melanesian plants: II. Kew Bull. 1936: 459-466.
CaRLQUIST, 8.
1965. Island life. A natural history of the islands of the world. Natural
History Press, New York.
1966. The biota of long-distance dispersal IV. Genetic systems in the floras
of oceanic islands. Evolution, 20: 433-455.
CuiarkgB, C. B.
1883. Cyrtandreae. DC. Monogr. Phan. 5 (1): 1-303.
GILLESPIE, J. W.
1930. New plants from Fiji. Bishop Mus. Bull. 74: 20-26, figs. 25-34.
Grant, V.
1958. The regulation of recombination in plants. Cold Spr. Harb. Symp.
Quant. Biol. 23: 337-363.
Gray, A.
1862. Characters of some new or obscure species of plants, of monopetalous
orders, in the collection of the U.S. South Pacific Exploring Ex-
pedition. Proc. Amer. Acad. 6: 40-41.
Guppy, H. B.
1906. Observations of a naturalist in the Pacific, vol. II: Plant dispersal.
Maemillan, London.
HILLEBRAND, W.
1888. Flora of the Hawaiian Islands. Heidelberg.
Sr. Joun, H.
1966. Monograph of Cyrtandra (Gesneriaceae) on Oahu, Hawaiian Islands.
Bishop Mus. Bull. 229: 1-465.
ScHLEcTeErR, R.
1923. Gesneriaceae papuanae. Bot. Jahrb. Engler 58: 255-379.
SEEMANN, B.
1861. Cyrtandra Pritchardii?. Bonplandia 9: 364-365.
1865-73. Flora vitiensis, 181-183; tab. 39-41. 1866. L. Reeve & Co.,
London.
Smita, A. C.
1936. Fijian plant studies. Bishop Mus. Bull. 141: 131-134.
1942. Fijian plant studies, II. Sargentia 1: 116-117.
1953. Studies of Pacific Island Plants, XIV. Notes on the Fijian species
of Cyrtandra. Journ. Arnold Arb. 34: 37-51.
1955. Phanerogam genera with distributions terminating in Fiji. Journ.
Arnold Arb. 36: 273-292.

158

Index

(Synonyms in italics. Page numbers of principal entries in boldface.)

Cheirodendron, 135
Clerodendron, 149

Cyrtandropsis, 155
Cyrtranda,

Subgenus Brachycyathus,
109, 134

Cyrtandra acutangula, 114, 129

alba, 155

aloisiana, 111, 114, 152, 153

amicta, 120

anthropophagorum, 114, 125, 139,
141

attenuata, 108, 114, 136, 137, 140

bracteolosa, 146

burbidgei, 119

bultii, 141

cephalophora, 114, 119

chippendalei, 108, 109, 114, 125,
133, 134, 135

chlorantha, 114, 123

ciliata, 114, 150

coleoides, 114, 155

coriacea, 147

cumingii, 119

cyathibracteata, 114, 117, 119

denhamii, 108, 114, 148, 150

desvoeuxit, 124

divertae, 137

dolichocarpa, 114, 125, 126, 127,
128, 129, 130

esothrix, 114, 142, 145

feaniana, 149

garnotiana, 110

gillespieana, 138

glandulosa, 124

godeffroyi, 119

gorriet, 154

gracilipes, 147

greenwoodiana, 144

harveyi, 108, 111, 114, 140, 154, 156

hornei, 114, 125, 144

involucrata, 114, 125, 138, 140, 141,
142

jugalis, 114, 140

Cyrtandra— Continued
kandavuensis, 108, 114, 136
krugeri, 127
leucantha, 114, 122, 123, 124
longiflora, 136
longipedunculata, 149
microstigma, 126

milnei, 111, 114, 122, 123, 124, 125,

126
montana, 114, 146
monticola, 138

multiseptata, 114, 127, 128, 129,

130
muskarimba, 114, 131
nodosula, 157
nudiflora, 127
nukuhivensis, 137
oblanceolata, 122
occulta, 114, 118, 123
prattii, 108, 114, 156
pritchardii, 114, 126, 147
pulchella, 127
pupukeansis, 136
rarotongensis, 150
reticulata, 114, 139
richii, 122
spathacea, 114, 135
taviunensis, 114, 145, 147
tempestii, 114, 149
tomentosa, 133
trichophylla, 114, 130, 131, 132
utriculosa, 129
vaupelii, 127
ventricosa, 114, 128, 129, 130
victoriae, 114, 132, 134

vitiensis, 108, 111, 114, 118, 120,

121, 122, 123, 126, 131
waianuensis, 122
xanthantha, 114, 151, 153

Exocarpos, 135
Gouania, 135
Pelea, 135
Psychotria, 107
Solanum, 107

159

U.S, GOVERNMENT PRINTING OFFICE: 1967

Camtssonia brevipes (A. Gray) Raven subsp. brevipes, from a watercolor by Henry R. Mockel,
made near Palm Springs, Riverside County, California; original in the Hunt Botanical
Library, Pittsburgh. Reproduced by permission of Mr. Mockel and the Hunt Botanical
Library.

BULLETIN OF THE UNITED STATES NATIONAL MUSEUM

CONTRIBUTIONS FROM THE UNITED STATES NATIONAL HERBARIUM

VouuME 37, Part 5

A REVISION OF THE GENUS CAMISSONIA
(ONAGRACEAE)

By Peter H. Raven

SMITHSONIAN INSTITUTION PRESS e CITY OF WASHINGTON e 1969

For sale by the Superintendent of Documents, U.S. Government Printing Office
Washington, D.C. 20402 - Price $1.75

1

A REVISION OF THE GENUS CAMISSONIA (ONAGRACEAE)

By Peter H. Raven

Introduction

Camissonia is one of the better characterized genera of Onagraceae’
tribe Onagreae. This tribe consists of 11 genera, more than half of the
total for the family. Included in it are some 215 species, comprising
about a third of the Onagraceae. My most recent synopsis of the
genera and tribes of the family was presented in 1964 (Brittonia 16:
276) and listed 12 genera of Onagreae. Subsequently, a detailed study
of the then monotypic genera Gongylocarpus and Burragea (Carlquist
and Raven, Amer. Journ. Bot. 53: 378-390. 1966) has led to the
conclusion that they should be merged. I would now list the genera
of the tribe Onagreae as follows: Gongylocarpus (including Burragea),
Camissonia, Gayophytum, Xylonagra, Oenothera, Stenosiphon, Hauya,
Calylophus, Gaura, Clarkia, Heterogaura.

I cannot agree with Munz (N. Amer. FI. IT. 5: 53. 1965) as to the
desirability of separating Hauya as a tribe distinct from the rest of the
Onagreae solely because these plants have stipules. This appears
to be an unwarranted emphasis of a single characteristic which would
lead to a less realistic portrayal of affinities within the family. Indeed,
the evidence seems to indicate clearly that Hauya is more closely
related to the other genera of tribe Onagreae with divided sporogenous
tissue—Calylophus, Gaura, Clarkia, and Heterogaura—than these
genera are to the remainder of the tribe. It scarcely seems credible
that the division of the sporogenous tissue of the anthers into packets,
separated by sterile tissue, is a characteristic that arose more than
once in the evolution of this group, and the combination of evening
opening of the flowers (in Hauya, Calylophus hartwegii sens. lat., and
most species of Gaura) with a broad, open hypanthium can hardly be
dismissed as mere coincidence.

My own work on the genus Camissonia was initiated in August 1957
and has occupied much of my time since that date. It is based on the
rich collections of this genus in the herbarium of the United States
National Museum and other herbaria (listed below), as well as on
extensive field studies throughout the area of the group. The first

161

162 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

part of this work was published in 1962—a revision of the group then
known as Oenothera subg. Chylismia (Univ. Calif. Publ. Bot. 34:
1-122). Subsequently, I became convinced that the species recently
included in Oenothera could better be grouped as three genera: Camas-
sonia, Calylophus, and Oenothera. I explained the reasons for this
decision to Dr. Philip A. Munz, and he requested that I publish this
material in synoptical form so that he could adopt the new generic
delimitations for his forthcoming treatment of the family (N. Amer.
Fl. II. 5: 79. 1965). Setting aside my work on the present revision,
I prepared a synopsis of the species and the sectional classification of
Camissonia, making new combinations when names were available
and in general presenting a bare outline of my views on the species
in this group, but without detailed justification. I did not, however,
publish any new species for which names at some level were not
already available. I made this manuscript available to Dr. Munz and
proceeded to get the material into print as soon as possible (Brittonia
16: 276-288. 1964).

Soon after the article appeared, Dr. Munz informed me that he
had changed his mind and would not accept my concepts of the genera
involved. He did, however, accept every proposition I made about
species, subspecies, and sections, making some dozens of transfers
of these taxa to Oenothera without comment on or justification for
their new status. I would naturally have preferred to make these
changes in rank in the present paper, where they could be fully
discussed and justified in detail. I have attempted to do this herein,
even when the changes in status have been made earlier.

Delimitation of Camissonia

In order to evaluate the position of Camissonia, it is necessary to
review some of the material I presented in 1964. Clarkia, recognized
as a distinct genus by all authors for more than 50 years, is distin-
guished from Oenothera (in the broad sense of Munz) by two charac-
teristics only: its possession of divided sporogenous tissue in the
anthers and by the basal attachment of these anthers. But similarly
divided sporogenous tissue is found in Hauya, Gaura, Heterogaura,
and in that portion of Oenothera treated by Munz as the subgenera
Calylophus and Salpingia and by me as the genus Calylophus. All of
these genera are likewise similar in stigma morphology (Raven,
Brittonia 16: 278. 1964) and in their broad, relatively long hypanthia.
The evidence that they are a coherent group, with clear interrelation-
ships, appears unequivocal.

There are two other distinctive groups of genera in the tribe
Onagreae. The first consists of Oenothera (in the relatively narrow
sense) and Stenosiphon, separated most obviously by the indehiscent,

RAVEN—GENUS CAMISSONIA 163

1-seeded fruits of the latter. In these two genera, the stigma is of a
type unknown elsewhere in the family: 4-lobed, without any disc
at the base and the lobes receptive all around. The second group
consists of the genera Gongylocarpus, Xylonagra, Gayophytum, and
Camissonia, all consisting of plants with a capitate or hemispherical
stigma of a sort widespread in the rest of the family, as well as rela-
tively short hypanthia and flowers that open primarily in the morning.
Looking at the pattern of variation in the tribe from this vantage
point, it is possible to recognize three distinct groups, the relation-
ships between which are not close. In spite of its highly specialized
fruits, it seems clear that Gongylocarpus includes the most generalized
species found in the tribe. It is the only genus of the tribe that retains
the original basic chromosome number for the family, n=11. It is
further clear that the sort of capitate or hemispherical stigma found
in this genus and in Xylonagra, Gayophytum, and Camissonia is
primitive in the tribe and probably in the family as a whole. One
other genus of the tribe, Hauya, has a similar hemispherical stigma;
but in Hauya, as we have seen, the sporogenous tissue in the anthers
is divided into packets by sterile tissue. Hauya is generalized in a
number of respects: chromosome number (2=10), habit (shrubs or
trees to more than 30 m. tall), and possession of stipules. Thus Hauya
must represent a relatively early offshoot of the phyletic line that
also gave rise to Calylophus, to Gaura, to Clarkia, and to Heterogaura.
One could scarcely imagine that these groups gained their identical
anther morphology independently, and we have mentioned their
similar floral morphology and biology. The relationships of the group
comprising Oenothera and Stenosiphon are obscure, although is it
clear that they belong in the tribe Onagreae. At some point the
ancestors of these plants made the transition to an evening-opening
habit, with the deep, narrow hypanthia and expanded stigmatic
surface characteristic of moth-pollinated plants in many families.
What then is the rationale for retaining Oenothera, Camissonia, and
Calylophus in a single genus, when these three groups differ absolutely
in anther morphology and stigma morphology, and modally in floral
biology? At best, this can be justified only on historical grounds;
there are no transitions between these three groups, no species are
intermediate, and no confusion of delimitation exists. The characters
used to separate the universally recognized genera Clarkia, Gayo-
phytum, Stenosiphon, Gaura, Heterogaura, and Xylonagra are of the
same sort as those which separate Oenothera, Camissonia, and Caly-
lophus, and the groups delimited are clearly as meaningful and
coherent phenetically. Of even more crucial importance, the evidence
that these three genera are each more closely related to other genera
than they are to one another appears overwhelming; not a single fact

164 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

has been adduced in opposition to this view. If our taxonomy is to
show anything whatsoever about the relationships of plants, there
appear to be only two possible courses of action. One would result
in the grouping of all genera of the tribe Onagreae, except Hauya
and Gongylocarpus, into a single genus; the other is the generic
treatment adopted here.

Historical and Taxonomic Considerations

A historical review of generic concepts in the tribe Onagreae was
presented in 1964 (Raven, Brittonia 16: 276-288) and will not be
repeated here. Many aspects of the taxonomic history of particular
species groups are given in the general remarks on the sections con-
cerned. A few general points are, however, germane.

The genus Camissonia, as delimited in this treatment, was first
outlined in my 1964 paper. Indeed, Camissonia is a somewhat unfa-
miliar name, Sphaerostigma, Tararia, Eulobus, and Chylismia being
more frequently employed at the generic level for some of its constitu-
ent groups. Very few combinations under Camissonia were made
prior to 1964; nevertheless, this does clearly seem to be the oldest
generic name applied to the complex in question, antedating Sphaero-
stigma by 17 years. Although the name Sphaerostigma is more familiar,
it has never been used in the inclusive sense of Camissonia as applied
in the present monograph, and nothing seems to be gained by con-
serving it or any other junior synonym over Camissonia.

Most species of Camissonia are endemic to the Western United
States, with three reaching adjacent Western Canada, a number in
the bordering states of Mexico, and one in Western South America,
whence the first known species, C. dentata, was described by Cavanilles
in 1795. The increasing tempo of exploration of California and
adjacent regions throughout the 19th century led to the discovery of
most of the species knowin today. The first more or less synoptical
treatment was that of Sereno Watson in his “Revision of the extra-
tropical North American species of the genus Oenothera” (Proc. Amer.
Acad. 8: 573-618. 1873); Watson’s work was reasonable and con-
servative as to species delimitation. In 1896, John Kunkel Small
published a review of the species under the title, “Oenothera and its
segregates,’”’ Bull. Torrey Cl. 23: 167-194. 1896. This was an exceed-
ingly uncritical treatment at both specific and generi¢ levels, replete
with new combinations and hasty, superficial judgments. Starting
six years later, Hector Léveillé confused the taxonomy of Oenothera
and related groups to an unprecedented degree by publishing his
“Monographie du Genre Onothera (1902-1913). Léveillé was surely
one of the most erratic taxonomic workers ever to publish a large
amount of material, and it is unfortunate that he concentrated so

RAVEN

GENUS CAMISSONIA 165

much of his effort on Oenothera; his illustration of an annual member
of the Polygonaceae, facing p. 167 of his Monographie, labeled
“Onothera gracilis (Philippi) Lévl.” is merely one example of his
notoriously superficial work. Aven Nelson remarked about Léveillé’s
efforts in 1905 (Bot. Gaz. 40: 54. 1905), “It is extremely difficult to
believe that his grouping of the specimens can stand, especially when
one finds that the annotations do not harmonize with the final pub-
lished list, and that the nomenclature of the illustrations in some
instances does not coincide with that of the text.”

Nelson did not help matters much in his review of Sphaerostigma,
presented as “Contributions from the Rocky Mountain Herbarium.
VI.” (Bot. Gaz. 40: 54-63. 1905), for this was again a relatively
uncritical listing of species which did not advance the understanding
of the group to any marked degree. On the other hand, this paper
did, for the first time, have the advantage of some knowledge of the
plants in the field, and it probably represents an improvement over
the efforts of Small and Léveillé, if not those of Watson.

In 1928, when P. A. Munz instituted his invaluable monographic
series on Oenothera, there were probably few groups of plants in the
United States for which the taxonomy was more confused. The con-
servative approach used in Munz’ revisions of Qenothera subg.
Chylismia (Amer. Journ. Bot. 15: 223-240. 1928), Sphaerostigma (Bot.
Gaz. 85: 233-270. 1928), and Tararia and Eulobus (Amer. Journ.
Bot. 16: 246-257. 1929) brought a remarkable degree of order out of
a tremendously confused situation. These three papers provided a
sound basis for the accumulation of knowledge about the plants here
treated as Camissonia which has served very well for some 40 years.

It is instructive to consider how and why the present revision differs
from them. Munz recognized 32 species; the present paper includes 61.
Both, coincidentally, recognize 29 major infraspecific taxa. Thus the
number of species has nearly doubled in the 40 years. One obvious
reason for this change has been the accumulation of additional
herbarium material during the intervening years. In 1929 Munz had
available 13 collections of the three species of sect. Hulobus endemic
to Baja California; I have been able to study well over a hundred.
That this has not been the primary reasons for the recognition of
more species, however, is suggested by the fact that in this section,
for all the accumulation of additional information, the taxonomy has
remained unchanged.

A much more important reason for the recognition of more species
now than in 1928-9 has been the accumulation of additional informa-
tion. First has been chromosomal information so vital to an under-
standing of the polyploid complexes which sect. Holostigma and sect.
Camissonia have proved to be. It is ndt surprising that some of the

166 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

most dramatic inflations in species number have taken place in these
groups, 4 species and 7 additional varieties in the former being recog-
nized here as 14 species with 3 additional subspecies, and 2 species
with 6 additional varieties in the latter being recognized here as 12
species with 4 additional subspecies. These changes have been made
largely to accommodate the distinctive population systems that have
been revealed by chromosomal information, although a few of the
taxa recognized were unknown or uncollected in 1929. A second sort
of information concerns the breeding systems of the plants concerned ;
thus series of populations consisting of relatively large-flowered, often
self-incompatible individuals are routinely separated at present from
their relatively small-flowered, autogamous relatives at the specific
level, whereas in 1929 the biological significance of such distinctions
was largely unknown to or not appreciated by taxonomists.

The most important changes in taxonomy in the past 40 years,
however, have been philosophical. In 1929, degree of difference or
similarity was the aspect of variation patterns stressed most in making
taxonomic decisions. In 1968, the pattern itself was stressed, with
discontinuities in the pattern of variation being emphasized as much
as or In some instances even more than the degree of similarity be-
tween the two populations being compared. In 1929 relatively similar
populations were often grouped as varieties of a single species regard-
less of whether they intergraded or remained completely distinct in
nature. Very different plants were separated at the species level even if
they intergraded completely and numerous intermediates were repre-
sented in the material being studied. The taxon here called Camissonia
pubens was formerly considered a variety of Oenothera contorta (Munz,
Bot. Gaz. 85: 254. 1928), even though no intermediates were found
or expected; on the other hand, the taxa here called Camissonia
boothia subsp. desertorum and subsp. alyssoides, being quite different,
were considered to belong to distinet species, even though it was noted
that the former “grades . . . insensibly . . .”’ (Alunz, Bot. Gaz. 85:
246. 1928) into the latter. Neither treatment reflects more information
about the systems of populations being classified, or is “better’’:
rather, each is in accordance with the taxonomic spirit of the times in
which it was constructed. As usual, one must turn to the presentation
of data on which the taxonomy is based to learn anything about the
pattern of variation that underlies the taxonomy; the mere recognition
of taxa conveys only a strictly limited amount of information about
the organisms being classified. Either sort of taxonomic philosophy
might be used to construct a useful classification if the facts on which
it was based were correct and well ordered; neither could be so used
if the facts were incorrect or poorly aligned with one another. In the
final analysis, all one can ask of a taxonomic system is that the popu-

RAVEN—GENUS CAMISSONIA 167

lations classified as lower level taxa be more similar to one another
(in some respect) than are the populations classified as higher level
taxa. Thus one would logically expect two subspecies to be more
similar to one another than two species in the same group; whether
this is true or not will depend on the perceptiveness of the taxonomist
making the classification and on the information available to him.

There is more unanimity about another philosophical difference be-
tween the taxonomy of 40 years ago and the taxonomy of the present.
In 1929, it was considered quite acceptable to give names to infra-
specific taxa which were based on genetically different members of
Mendelian populations, providing that one wished to emphasize the
differences between them. The denial of John Ray’s 17th-century
dictum about the construction of taxa implicit in the taxonomic recog-
nition of such segregating entities went largely undetected until the
1930s, when populations and not individuals were clearly established
as the units of biological classification. In general, taxonomists in-
tentionally have not given different formal names to the progeny of a
single individual or pair of individuals, no matter how different these
might be, and this has been the prevalent tradition for some 300
years. In part, of course, the recognition of such entities as varieties
sometimes happens when insufficient information is available; but
in the 1920s such a taxon as Oenothera cheiranthifolia var. nitida
could be recognized as distinct, basically because ‘‘the aspect of a
wholly glabrous plant is quite different from that of a very hairy
one...” (Munz, Bot. Gaz. 85: 270. 1928), even though such plants
were well known to occur as segregating elements in normal popu-
lations. Most taxonomists would not at present accord formal recog-
nition to such an entity.

The desire to give formal taxonomic names to distinct populations
only and not to the genetically distinctive elements in such populations
underlies the modern preoccupation with the geographical race as the
only infraspecific entity worthy of recognition, a tendency that was
apparent more than a century ago. Many taxonomists of the present
day formally recognize only geographic races below the species level,
although this may sometimes pose difficulties when the organisms
being classified are largely or wholly autogamous.

Subspecies are used herein as the only infraspecific taxon for the
following reasons. The Code states explicitly that subspecies are the
primary unit into which species are divided, and it is therefore in-
correct to use any other infraspecific taxon, such as variety, for the
primary division of a species. From a formal standpoint, there is
much to recommend Pennell’s (Amer. Journ. Bot. 36: 1922. 1949)
suggestion to treat subspecies and varieties as synonymous, to use
the former word in scientific nomenclature and to relegate the latter

168 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

to informal usage, where the term “variety” has already gained a
multiplicity of meanings quite inconsistent with its employment as
a scientific term. Should such a suggestion be adopted, the nomen-
clatural problem would be simplified and the number of new com-
binations greatly reduced. A tendency toward the recognition of
only one level of taxa below the level of species is already apparent
in Kurope. For me, the recognition of more than one such level
implies a degree of precision in the understanding of relationships
which often becomes ludicrous. As H. L. Mason has often empha-
sized, two of any three taxa will always be more closely related to
one another than they are to the third; but is it practical to recognize
all such shades of relationship in our formal taxonomy, even if it
were possible to ascertain the facts of the case unequivocally? I
personally feel that the adoption of the trinomial system in zoology
at an early date was a sound solution of this particular problem of
nomenclature. I hope that botanists may ultimately adopt such a
system and consider it desirable meanwhile for names at both varietal
and subspecific levels to be considered for purposes of priority at
either one, even though it is not technically necessary to do so.

In the present revision, I have tried to indicate in the synonymy
how the concepts of Munz can be integrated with my concept. I have
not given references to other taxonomic papers, since this has been
done by Munz in his revisions, which have been the only important
basis of classification of the groups for some 40 years. I have, how-
ever, given full synonymy and citation of types in all cases. Specimens
are not cited here for many of the taxa treated in the same sense in
which they were understood by Munz; the ranges of such taxa are
already sufficiently documented in the literature. For sects. Chylismia,
Lignothera, and Chylismiella 1 have not cited specimens or given
distribution maps, since this information has been presented in my
earlier revision (Raven, Univ. Calif. Publ. Bot. 34: 1-122. 1962).

Geographical Distribution

Camissonia is primarily a group of the Western United States and
adjacent Mexico, although a few autogamous species extend beyond
this area. Three species —Camissonia andina, C. breviflora, and C.
contorta—are found in the southern part of Western Canada; all of
them are autogamous. On the other hand, 79 of the 90 taxa recog-
nized in this revision are found in California, and 21 are endemic to
the State (12 other endemics just reach southern Oregon or north-
western Baja California). Three of the four species of Camissonia
sect. Hulobus are endemic to Baja California, us are C. cardiophylla
subsp. cedrosensis (technically also found in Sonora because of its

RAVEN—GENUS CAMISSONIA 169

occurrence on Isla Tiburén), C. claviformis subsp. wigginsu, C.
proavita, and CO. guadalupensis subsp. guadalupensis, the last two in
the California floristic region. No taxon is endemic to Sonora, although
eight occur there, and C. californica and C. chamaenertoides range
widely in the State. No species of Camissonia is known from Chihuahua,
although C. chamaenerioides, the only species of the genus that occurs
in Texas, may well occur there. Only two species, C. chamaeneriordes
and C. scapoidea subsp. seapoidea, are definitely known to occur in
New Mexico, although C. refracta may also occur in that State.

In Arizona, 27 taxa occur, and C. palmeri and C. strigulosa might also
be represented. Of these, C. claviformis subsp. peeblesii, C. speculicola
subsp. hesperia and subsp. speculicola, C. confertiflora, C. scapoidea
subsp. macrocarpa, and C. exilis, all belonging to sect. Chylismia, ure
endemic. In Colorado, only C. walkeri subsp. walkeri, C. eastwoodiae,
and C. seapoidea subsp. scapoidea of sect. Chylismia, C. parvula of sect.
Camissonia, and C. breviflora and C. subacaulis of sect. Tetrapteron
are known, and only C. seapoidea occurs east of the Rocky Moun-
tains. Utah has 22 taxa, none endemic, and Nevada has 33, with C.
nevadensis and C. megalantha well-marked endemics. In Wyoming,
Camissonia scapoidea subsp. scapoidea, C. subacaulis, C. breviflora, C.
parvula, C. minor, and C. andina, all autogamous, are found, with
only C. subacaulis, C. breviflora, and C. andina reaching Montana.
Idaho has 13 taxa, Oregon 18, and Washington 10; Camissonia
hilgardii is virtually endemic to Washington and C. pygmaea endemic
to the 3-State area.

It is evident, therefore, that Camissonia centers in the California
floristic area, where the plants chiefly occur in plant associations
derived from the Madro-Tertiary Geoflora. Sect. Holostigma, with 14
species and 3 additional subspecies, is virtually endemic to this region,
as is the close-knit group of 11 taxa comprising species 44—51 of sect.
Camissonia (this latter group also including the only South American
species, the autogamous C. dentata). Two other sections (Chylismia
and Eremothera) and the four taxa comprising species 40-43 of sect.
Camissonia center in the Great Basin and adjacent deserts to the
south, as does the monotypic sect. Chylismiella. Sect. Nematocaulis,
comprising two closely related species, is associated with the high,
cold deserts north of the Great Basin, and sect. Tetrapteron, while
probably radiating from floristic regions similar to those represented
in cismontane California at the present day, has also radiated in the
mountains and deserts of these northern regions. Sect. Lignothera has
its maximum diversity in the deserts and mountains about the head
of the Golfo de California. Finally, sect. Hulobus, small but phylo-
genetically crucial, is primarily associated with the peninsula of Baja

170 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

California, well known as one of the most important refugia in the
southern part of North America, where it is associated with Gongylo-
carpus fruticulosus (Benth.) T. 5. Brandegee, perhaps the most
generalized species of the tribe Onagreae surviving at the present
day.

Phylogenetic Relationships

In the light of information presented up to this point, what can be
said about the relationships of the sections of Camissonia? First of all,
the nine sections I have recognized are sharply distinct from one
another, and there are no transitions between them. Their association
into phylogenetic lines or circles of affinity is extremely problematical,
and there seems to be absolutely no advantage to be gained by group-
ing them into subgenera as has been done by Munz (N. Amer. FI. IT.
5: 142-177. 1965); indeed, I would hold that such an association of
the sections into artificial groupings is misleading. Camissonia is
clearly the most heterogeneous of the 11 genera of the tribe Onagreae,
and its sections are amply differentiated. Keeping these facts in mind,
it is possible to offer a few deductions about the phylogeny of the
group and the relationships of its constituent parts.

We have already seen that the genus Gongylocarpus comprises the
most generalized species in the tribe, and it appears reasonable to use
it as a standard for examining phylogenetic relationships in Camis-
sonia. The other two genera closely related to Camissonia—Xylonagra
and Gayophytum—do not appear to offer much guidance in this respect;
although, as we shall see, the restriction of the monotypic Xylonagra
to central Baja California may well be significant. Within Gongylo-
carpus, the low shrub G. fruticulosus is clearly more generalized, in
view of its self-incompatibility, woody habit, and large flowers, than
the wide-spread autogamous G. rubricaulis Schlecht. & Cham. (Carl-
quist & Raven, Amer. Journ. Bot. 53: 378-390. 1966). Gongylocarpus
fruticulosus is restricted to two adjacent islands in Bahia Magdalena,
southern Baja California, where it is locally abundant.

Geographically, it is clear that, Camissonia sect. Eulobus, the only
section of the genus centering in Baja California, where three of its
four species are endemic, deserves special attention. One of its species,
Camissonia crassifolia, has the woodiest habit of any Camissonia; also
the section shares with Gongylocarpus the fleshy dise within the hy-
panthium which is, among the members of tribe Onagreae, known
only in these two genera. Thus the woody habit, the fleshy disc in the
hypanthium, and a common habitat in the Baja California littoral all
link Gongylocarpus fruticulosus with Camissonia crassifolia, and |
believe the latter to be the most generalized extant species of its genus.

RAVEN—GENUS CAMISSONIA 171

Within Camissonia, pinnately lobed or divided leaves are found only
in sect. Hulobus, sect. Chylismia, and sect. Tetrapteron, and the mem-
bers of the latter group are the only species of Camissonia which share
with sect. Hulobus a fleshy disc closing the hypanthium. The relation-
ship between Hulobus and Chylismia is obviously close; indeed, the
basal rosette of Camissonia californica (which normally withers by
the time the plants flower) is often virtually indistinguishable from
that of C. multijuga, one of the more generalized species of sect.
Chylismia. One of the most interesting lines of evidence for a close
relationship between these three groups, however, has been developed
recently; in some species of each most of each petal is strongly ultra-
violet reflective, as usual in the genus, but the basal portion is occupied
with a conspicuous, nonultraviolet-reflecting spot which is invisible to
the human eye but can be photographed with special techniques
(D. P. Gregory and P. H. Raven, MS.). No such nonultraviolet-
reflecting areas have been found on the petals of outcrossing species
of sect. Holostigma, sect. Camissonia, or sect. Hremothera, and in the
first two examples, the place of this nonultraviolet-reflecting spot is
obviously taken by the one or two large, bright brownish-red dots at
the base of each petal, which doubtless serve the same function as
nectar guides. The species of sects. Chylismia and Tetrapteron that
have the nonultraviolet-reflecting spots have petals that appear clear,
bright yellow and unspotted to the human eye, whereas those of sect.
Eulobus have a distinctive, fine red flecking throughout the non-
ultraviolet-reflective area.

In summary, I hold that sect. Hudobus is the most primitive group
within the genus Camissonia, and that from plants similar to this
croup were derived the ancestors of sect. Chylismia, which radiated
into the deserts and later gave rise to the small section Lignothera,
specialized for pollination by hawk moths. Also from plants similar
to sect. Hulobus, the species of sect. Tetrapteron, which became asso-
ciated with more mesic habitats, were derived. Presumably among
the most primitive species of sect. Holostigma is Camissonia chei-
ranthifolia, the only woody or even perennial species found among
the last five sections of the genus as I have arranged it here. These
sections were probably also derived from plants similar to those of
sect. Hulobus. All species of these five relatively advanced sections
have simple leaves that are never lobed (at the most coarsely serrate) ;
none has a dise in the hypanthium; and none has a nonultraviolet-
reflective area on the petal, so far as known.

Evident shifts from the perennial to the annual habit have charac-
terized the first five sections listed here (the last four are strictly
annual); these shifts have usually been accompanied by a change

172 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

from self-incompatibility to self-compatibility. Self-incompatible
species are distributed among the nine sections of Camissonia as

follows:
Self-incompatible Self-compatible

species species Total

I. Eulobus 2 2 4
II. Chylismia a7 6 13
III. Lignothera — 2 2
IV. Tetrapteron b2 c4 6
V. Holostigma 42 12 14
VI. Camissonia 2 10 12
VII. Eremothera 2 e5 7
VIII. Chylismiella - 1 1
IX. Nematocaulis — 2 2
17 44 61

9 C. confertiflora, C. eastwoodiae, and C. parryi not tested but listed here provisionally.

b C. tanacetifolia seems to include self-compatible populations.

¢ Some populations of C. swbacaulis might be self-incompatible.

4 C. cheiranthifolia subsp, suffruticosa in part only; subsp. cheiranthifolia wholly self-compatible.
¢ C. nevadensis might be self-incompatible, but this seems unlikely.

Of the nine sections recognized in this revision, three small ones
contain no self-incompatible species; one (Chylismia) consists mainly
of self-incompatible species; and the other five contain one or two
self-incompatible species each, with a majority of self-compatible
ones. It is interesting that of the two groups associated with the most
severe desert, Chylismia and Eremothera, the former is the only section
in which as many as half of the species, including nearly all of the
common ones, are self-incompatible, and the most widespread species
of the latter, Camissonia boothii, is likewise self-incompatible. One
normally associates exploitation of a desert habitat with self-compati-
bility or autogamy, but this has evidently not been the case here;
perhaps the ability to adjust to the rapidly shifting desert habitats
calls for a maximum of recombination, and this is precisely what
seems to be favored in both groups, which conform to the definition
of homogamic complexes (Raven, Univ. Calif. Publ. Bot. 34: 66-67.
1962).

If pinnately lobed or divided leaves are primitive for the genus
Camassonia, as the evidence appears to indicate, there have been evolu-
tionary trends toward simple leaves in sect. Hulobus (where some popu-
lations of Camissonia crassifolia, such as those on Isla de Cedros, have
lobed leaves, while most others have entire leaves) and sect. Chylismia
(where lobed or divided leaves predominate, although some species
have lost them). Sect. Lignothera appears to be a derivative of sect.
Chylismia, specialized for hawk-moth pollination in its long hypanthia,
the longest in the genus. In the two species comprising this group, the
pollen is shed in tetrads, a characteristic otherwise unknown in the
tribe Onagreae. In sect. Tetrapteron, Camissonia tanacetifolia and

RAVEN—GENUS CAMISSONIA 173

C. breviflora are characterized by deeply lobed leaves, as are some
populations of C. subacaulis; whereas the remaining perennial species,
C. ovata, has entire leaves, as do the two annuals. The superficial
similarity between these two annual members of sect. Tetrapteron
and some species of sect. Holostigma provides a clue as to the relation-
ships of the latter group, but probably tells us nothing about the
exact route of its derivation.

Within the last five groups listed here, relationships are almost
entirely obscure. Sect. Holostigma includes the only perennial species
and perhaps is morphologically the most generalized of the five sec-
tions. There does not, however, appear to be any evidence for close
relationship between this group and any other. Sect. Camissonia
consists entirely of yellow-flowered species, like sect. Holostigma;
but no species are transitional between the two groups and there is no
evidence for close relationship between them. Polyploidy is important
in both of these sections, which have diversified largely in the areas
derived from the Madro-Tertiary Geoflora and now center in Cali-
fornia, with one group of sect. Camissonia (species 40-43) centering
in the northern Great Basin. Sect. Eremothera, consisting of white-
flowered species, does not exhibit any evident relationship to any
other group, and it seems certain that white-flowered species evolved
at least three times in the history of the genus, once here, once in
C. claviformis (sect. Chylismia), and once in C. pterosperma (the only
species of sect. Chylismiella). In the first two instances, the acquisi-
tion of white flowers was undoubtedly associated with a shift from
bee-pollination to pollination by small moths.

Almost nothing can be said about the relationships of the last two
small sections, Chylismiella (with one species) and Nematocaulis (with
two very closely related ones). The white petals of the former, yellow
at the base, are found elsewhere in the tribe only in the genus Gayo-
phytum; the seeds of Camissonia pterosperma, the only species of this
section, are unique in the family. Likewise, the flattened capsules of
both species of sect. Nematocaulis are reminiscent of those of Gayo-
phytum humile A. L. de Jussieu, probably the most generalized species
of its small genus (Lewis & Szweykowski, Brittonia 16: 343-391.
1964). It appears possible that these two small, distinctive sections
of Camissonia have been derived from the same evolutionary line that
gave rise to Gayophytum, but nothing more can be said about their
relationships at this time.

Pollination Systems

Most outcrossing species of Camissonia are pollinated by bees, and
a number of these bees are oligolectic, gathering pollen from no other
plants (Linsley, MacSwain, & Raven, Univ. Calif. Publ. Entom.

174 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

33: 1-98. 1964). One group of two species, sect. Lignothera, has
shifted to opening in the evening, when the plants are pollinated
by hawk moths; these species have yellow flowers. The outcrossing
species of the entirely white-flowered sect. Hremothera and the white-
flowered and afternoon opening yellow-flowered subspecies of Camis-
sonia claviformis (sect. Chylismia) are predominantly pollinated by
small moths after their opening in the late afternoon or early evening,
but in both groups secondary associations with oligolectic bees in the
late afternoon have also evolved (Linsley et al., op. cit.). Two species
of sect. Chylismia, Camissonia megalantha and C. heterochroma, are
unique in the genus in having purple flowers. In both, the flowers
open near sunrise; the former species is pollinated by bees, and the
latter is autogamous. Finally, C. pterosperma has white petals that
are yellow at the base and flowers that open in the morning. This
pattern of petal coloration is found elsewhere in the tribe only in
Gayophytum, where pollination by flower-flies and small bees predom-
inates (MacSwain, Raven, & Thorp, MS.) ; but Camissonia pterosperma
is largely if not entirely autogamous.

One of the most interesting aspects of the pollination system in
Camissonia, and indeed in the Onagraceae as a whole, is the common
occurrence of pairs of species—one member large-flowered and out-
crossing (and sometimes self-incompatible), the other small-flowered
and self-pollinating (and often a polyploid or complex structural
heterozygote, depending on the group). In Camissonia, many examples
of such pairs are evident: C'. multijuga-C. walkeri subsp. tortilis (sect.
Chylismia) ; C. tanacetifolia-C. breviflora (sect. Tetrapteron) ; C. bistorta-
C’. lewistt (sect. Holostigma); C. kernensis subsp. kernensis-C. pubens
(sec. Camissonia); C. refracta-C. chamaenerioides (sect. Eremothera) ;
and C. hilgardii-C. andina (sect. Nematocaulis). Autogamy doubtless
arises with a certain frequency throughout the range of normally
outcrossing species, but if it is to spread and become characteristic
of entire populations, it must do so because it provides for the duplica-
tion of a genotype that is highly successful in some area. This has
obviously occurred time after time in the evolution of the genus
Camissonia.

Chromosome Numbers and Behavior

The earlier reports of chromosome number listed by Lewis, Raven,
Venkatesh, and Wedberg (Aliso 4: 73-76. 1958) and those of Gregory
and Klein (Aliso 4: 505-521. 1960) are repeated in this paper. On the
other hand, the numerous chromosome counts I reported earlier for
taxa of sect. Chylismia, Lignothera, and Chylismiella (Raven, Univ.
Calif. Publ. Bot. 34: 1-122. 1962) are not included in this revision,
although a few more recent determinations of chromosome numbers
for these groups are reported. The only other familiar reports of

RAVEN—-GENUS CAMISSONIA 175

chromosome number in Camissonia are six undocumented counts
listed by Johansen (Proc. Nat. Acad. Sci. U.S, 15: 882-885. 1929;
Amer. Journ. Bot. 16: 595-597. 1929), without citation of locality or
voucher, and the single, probably erroneous count reported by Bell
and discussed below. Johansen’s counts are mentioned in the accounts
of the taxa to which they refer.

Aneuploidy is almost unknown in Camissonia and rare in the tribe
Onagreae. Only in Clarkia (with basic chromosome numbers of «=5
6, 7, 8, and 9) does aneuploidy seem to be a regular feature of the
evolutionary pattern which often accompanies speciation. In Camis-
sonia, at least two populations of the South American tetraploid C.
dentata have n =13 instead of the usual n =14; this appears to be the
only documented instance of aneuploidy in the genus. In these popu-
lations, aneuploidy does not appear to have had any important
bearing on speciation. Bell (Sida 2: 169. 1965) reported n=6 in
Camissonia claviformis subsp. peeblesii (“Ocenothera clavaeformis var.
aurantiaca’”’) from Maricopa County, Arizona (Bell 17558, NCU). In
the drawings made from the original preparations, however, which
Dr. Bell has kindly allowed me to examine, it appears that 2 =7 is
as likely a count as n—6. All other individuals of this species that
have been examined cytologically (a total of 411) have had n =7.

Although detailed information about chromosome number and
behavior is given throughout the text of this paper and in the appen-
dix, and summaries are given for each section, it seems worthwhile
to summarize the chromosomal information available for Camissonia
at this point, as follows:

Additional Polyploid tara __
Individuals Populations Species subspecies wholly partly
I. Eulobus 108 78 4
II. Chylismia 683 239 13 14 2
III. Lignothera 36 23 2 2
IV. Tetrapteron 49 46 6 1 1 1
V. Holostigma 389 3380 14 3 D
VI. Camissonia 192 181 12 4 aQ
VII. Eremothera 67 57 7 5 1
VIII. Chylismiella 1 1 L
IX. Nematocaulis 11 11 2 1
1536 971 61 29 16 a

« Including two Chilean populations of C. dentata subsp. dentata with n=13.

As can be seen from the table, the chromosomal sampling of
Camissonia has been extensive. Counts are available for at least one
population representing each of the nine sections, and for every
species and subspecies except the following: Camissonia confertiflora,
C. speculicola subsp. hesperia, and C. scapoidea subsp. macrocarpa,
and subsp. utahensis of sect. Chylismia, and C. gouldii of sect. Ere-

295-655 O—6S8

2

176 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

mothera. Thus 59 of the 61 species recognized here and 85 of the 90
taxa have been examined cytologically. It seems unlikely that the
pattern will be modified in any substantial way by further chromo-
some counts.

The polyploid plants examined in sect. Eremothera and_ sect.
Tetrapteron, as well as the polyploids of Camissonia walkeri (sect.
Chylismia), are clearly cytologically autopolyploid, with rings of
chromosomes forming regularly at meiotic metaphase I. The hexa-
ploid in sect. Tetrapteron has almost certainly been derived directly
from the tetraploid; these two races are regarded as subspecies of
C. tanacetifolia. The single diploid plant of this species that has been
examined may well have been derived directly from a tetraploid
individual as discussed in detail following the taxonomic treatment
of the species.

With one possible exception (see appendix) no rings of chromosomes
have been observed in the polyploids of sect. Holostigma and sect.
Camissonia, and polyploidy has played the most important role in
these two groups in the course of evolution. The same is true of the
polyploids of sect. Nematocaulis. Morphological and geographical rela-
tionships make it virtually certain that the tetraploid populations in
C. californica (sect. Hulobus) and in C. seapoidea subsp. scapoidea
(sect. Chylismia) have had an autotetraploid origin, even though
they do not form rings of 4 chromosomes at meiotic metaphase I.
In all of these groups the chromosomal homology of the diploid
entities (see below) makes it clear that the formation of multivalents
in polyploids is genetically controlled, and prevented even if the
diploid genomes present in « polyploid are largely homologous. In
the further interpretation of these data, it is instructive to consider
the occurrence of translocation heterozygotes in naturally oecurring
diploid populations of these sections. It is important to stress the
occurrence of such heterozygotes in outcrossing taxa, as rings of
chromosomes tend to be eliminated relatively rapidly in the course
of autogamous reproduction, except in the special case of balanced
structural heterozygosity.

In sect. Chylismia, translocation heterozygotes comprised about 20
percent of the individuals examined (Raven, Univ. Calif. Publ. Bot.
34: 40. 1962), and the more limited amount of information available
for sect. Lremothera suggests that a comparable level of heterozygosity
may obtain here also. In one plant of Camissonia refracta, a member
of this section, a ring of 10 chromosomes was found at diakinesis ;
this was the largest ring found in the genus. The situation in other
well-sampled groups is, however, very different. In sect. Camisson id,
only about 2 percent of the outcrossing individuals examined had a
ring of chromosomes at meiotic metaphase; and in sect. Holostigma

RAVEN—GENUS CAMISSONIA 177

and sect. Hulobus, no multivalents have been observed, even though
approximately 500 plants of the two groups have been examined.
Even if reciprocal translocations occur frequently in these sections,
they must be eliminated rapidly from the populations.

One naturally associates the frequent occurrence of reciprocal
translocations within the populations of a species with the role of
such translocations in the differentiation of species in the same group.
It might therefore be expected that patterns of chromosomal differ-
entiation between species would be different in sect. Chylismia from
those observed in sects. Holostigma and Camissonia. This does not,
however, appear to be the case; interspecific hybrids in all three
groups show chromosomal configurations which indicate that their
parents differ by, usually, 2-4 reciprocal translocations. Thus the
appearance of reciprocal translocation configurations at a high fre-
quency within the populations of a species may have little to do with
the patterns of differentiation between populations of the group. In
all three sections, there is a high degree of chromosome association
in diploid interspecific hybrids; hence, the control of multivalent
formation in polyploids must have a genetic basis and not be related,
for example, to the frequent formation of univalents in interspecific
hybrids in some of the groups. It may be significant, however, that
pair-forming polyploids are frequent and evolutionarily important in
those sections in which reciprocal translocations are infrequent or
rare in natural populations, and virtually absent in those groups in
which reciprocal translocations are abundant in natural populations.

Finally, sect. Chylismia is the only group of the genus in which
supernumerary chromosomes have been noted. Here, one or more
supernumeraries were found in some 3.7 percent of the nearly 700
individuals examined (Raven, Univ. Calif. Publ. Bot. 34: 41. 1962),
this observation according well with an earlier hypothesis of a causal
relationship between translocation heterozygosity and supernumerary
formation (Lewis, Evolution 5: 142-157. 1951, Lewis & Raven,
Rec. Adv. Bot. 1466-1469. 1961). Supernumeraries might be expected
to occur in sect. Eremothera if more extensive sampling was carried out.

Acknowledgments

I am most grateful to Harlan Lewis for suggesting this study
initially and supporting it in its early stages, as well as for many useful
discussions subsequently; to Lee W. Lenz, Director, Rancho Santa
Ana Botanic Garden, for extensive support of the work during the
time I was employed at the Garden (1961-62); and to the National
Science Foundation for GB-141 (1962-65), which allowed the comple-
tion of this project. Philip A. Munz has generously shared his wide
experience of the family with me throughout the course of the work.

178 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

The Plant Biology Station of the Carnegie Institution of Washington,
Stanford, has most kindly allowed the use of their facilities in cultivat-
ing selected strains of Camissonia during the past five years. Most of
the illustrations and all of the maps were prepared by Marilyn Wright,
while Joan Burger provided the drawings of sect. Holostigma. I am
grateful to Mr. Henry R. Mockel, Twentynine Palms, Calif., and to
George H. M. Lawrence, Director of the Hunt Botanical Library,
Pittsburgh, for permission to reproduce Mr. Mockel’s attractive water-
color (frontispiece) of Camissonia brevipes subsp. brevipes. Figures 57
and 63 are drawn on a base map kindly supplied by the California
Insect Survey. Dennis E. Breedlove supplied expert technical assist-
ance in growing the plants, hybridizing them, and determining
chromosome numbers, especially during the years 1963-64.

Many individuals have contributed special materials for this study,
comprising either herbarium specimens, seeds, or fixed buds for
cytological study. Extensive collections were presented by Reid V.
Moran and Dennis E. Breedlove, and other material collected espe-
cially for this purpose was generously donated by E. R. Blakley, D. P.
Gregory, W. Klein, H. Lewis, Mildred E. Mathias, P. A. Munz,
R. Ornduff, C. L. Porter, C. F. Smith, H. J. Thompson, E. C. Twissel-
mann, Frank Vasek, and H. L. Wedberg. Clare Hardham has provided
a large number of interesting collections and done much to expedite
my field studies in northern San Luis Obispo and southern Monterey
Counties, California.

T am also most grateful to the curators of the following herbaria,
from which material has been supplied for this study: University of
New Mexico, Albuquerque (UNM); Sul Ross State College, Alpine,
Texas (SRSC); Iowa State College, Ames (ISC); University of
Michigan, Ann Arbor (MICH); University of Texas Herbarium,
Austin (TEX); University of California, Berkeley (UC); Jepson
Herbarium, Berkeley (JEPS); University of Colorado, Boulder
(COLO); Montana State College, Bozeman (MONT);. Gray Her-
barium, Harvard University (GH); Field Museum of Natural
History, Chicago (F); Pomona College (POM); Rancho Santa Ana
Botanic Garden, Claremont, Calif. (RSA); Tracy Herbarium, Texas
A & M University, College Station (TAES): Instituto de Biologia,
Concepcion, Chile (CONC); Botanical Museum and Herbarium,
Copenhagen (C); Oregon State University, Corvallis (OSC); Her-
barium of Southern Methodist University, Dallas, Texas (SMU):
University of California, Davis (DAV); Royal Botanic Garden,
Edinburgh (E); University of Oregon, Eugene (ORE); Colorado
State University, Fort Collins (CS); Fresno State College, Fresno,
California; Grand Canyon National Park, Arizona; Royal Botanic
Gardens, Kew (K); Rocky Mountain Herbarium, University of

RAVEN—GENUS CAMISSONIA 179

Wyoming (RM); Herbarium of the Komarov Botanical Institute
of the Academy of Sciences of the U.S.S.R., Leningrad (LE); Inter-
mountain Herbarium, Utah State Agricultural College (UTC);
British Museum (Natural History), London (BM); University of
California, Los Angeles (LA); Los Angeles County Museum (LAM) ;
University of Wisconsin, Madison (WIS); Tulane University Her-
barium, New Orleans (NO); The New York Botanical Garden (NY);
University of Notre Dame (ND): National Museum of Canada,
Ottawa (CAN); Plant Research Institute, Canada Department of
Agriculture, Ottawa (DAO); University of Oxford (OXF); Muséum
National d’Histoire Naturelle, Paris (P); Academy of Natural
Sciences, Philadelphia (PH); Idaho State College Herbarium, Poca-
tello (IDS); Brigham Young University, Provo, Utah (BRY);
Washington State University, Pullman (WS); University of Nevada,
Reno (RENO); Nevada Agricultural Experimental Station, Reno;
Missouri Botanical Garden, St. Louis (MO); Peck Herbarium of
Willamette University, Salem, Oregon; University of Utah, Salt Lake
City (UT); San Diego Museum of Natural History (SD); San Diego
State College (here given as SDSC); California Academy of Sciences,
San Francisco (CAS); California State Polytechnic College, San Luis
Obispo (OBI); Santa Barbara Botanic Garden, Santa Barbara,
California (SBSG); Santa Barbara Museum Of Natural History
(SBM); University of California, Santa Barbara (UCSB); Museo
Nacional de Historia Natural, Santiago, Chile (SGO); University of
Washington, Seattle (WTU); Dudley Herbarium, Stanford Universi-
ty, Stanford, California (DS); New Mexico College of Agricultural and
Mechanical Arts, State College (NMC) ; Naturhistoriska Riksmuseum,
Stockholm (S); Desert Botanical Garden, Tempe, Arizona (DES);
University of British Columbia, Vancouver, British Columbia (UBC) ;
U.S. National Museum, Washington (US); U.S. National Arboretum,
Washington (NA). In addition, I have examined the material of
Camissonia in the private herbarium of Ernest C. Twisselmann,
Cholame, California.

Systematic Treatment

Camissonia Link

Camissonia Link, Jahrb. Gewaechsk. 1: 186. 1818.

Sphaerostigma (Seringe) Fisch. & Mey., Ind. Sem. Hort. Petrop. 2: 49. 1835.
Based on Oenothera sect. Sphaerostigma Seringe, in DC., Prod. 3: 46. 1828.
Lectotype: O. dentata Cav.= Camissonia dentata (Cav.) Reiche.

Agassizia Spach, Hist. Veg. Phan. 4: 347. 1835; not Chavannes 1880.
Type: A. cheiranthifolia (Hornem. ex Spreng.) Spach=Camissonia
cheiranthifolia (Hornem. ex Spreng.) Raimann.

Holostigma Spach, Nouv. Ann. Mus. Paris III. 4: 332. 1835.

180 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Eulobus Nutt. ex Torr. & Gray, Fl. N. Amer. 1: 514. 1840. Type: E.
californicus Nutt. ex Torr. & Gray =Camissonia californica (Nutt. ex Torr.
& Gray) Raven.

Chylismia (Nutt. ex Torr. & Gray) Raimann, in Engl. & Prantl, Natiirl.
Pflanzenfam. III. 7: 217. 1893. Based on Oenothera subg. Chylismia Nutt.
ex Torr. & Gray.!. Type: O. scapoidea Torr. & Gray = Camissonia scapoidea
(Torr. & Gray) Raven.

Taraxia (Nutt. ex Torr. & Gray) Raimann, in Engl. & Prantl, Natiirl.
Pflanzenfam. III. 7: 216. 1893. Based on Oenothera subg. Tararia Nutt.
ex Torr. & Gray, Fl. N. Amer. 1: 506. 1840. Lectotype: O. breviflora
Torr. & Gray=Camissonia breviflora (Torr. & Gray) Raven.

Annual or perennial herbs, rarely subshrubs, from a taproot which
in some species branches and gives rise to new plants. Leaves basal
or cauline and spirally arranged, largest near the base, reduced up-
ward. Flowers opening near sunrise, or in a few taxa near sunset or
in the late afternoon, the stigma receptive and anthers shedding pollen
simultaneously and immediately and the flowers withering in less
than a day, 4-merous or very rarely 3-merous. Petals yellow, rose-
purple, or white, reflexed in anthesis in the outcrossing species.
Stamens and style yellowish, the stigma greenish yellow. Stamens 8,
the episepalous ones longer, or very rarely 4, the epipetalous ones
lacking; anthers versatile or basifixed (in sect. Tetrapteron), the sporog-
enous tissue in each locule undivided. Pollen yellow, shed singly or (in
sect. Lignothera) in tetrads. Stigma capitate or hemispherical. Capsule
loculicidal, usually promptly so, straight or contorted, the seeds in 1
or 2 rows in each of the 4 locules, usually adhering to the central
partition following dehiscence. Basic chromosome number, 2=7.
Self-incompatibility known in approximately 17 of the 61 species.

TYPE SPECIES: Camissonia flava Link=C. dentata (Cav.) Reiche.

In the following treatment, an effort has been made to group related
species. Within each section, generalized species are listed first, more
specialized ones later. The keys have been written for ease of identifi-
cation and are not intended to display relationships. Sympatric
entities have been listed within each section following the taxonomic
treatment of each taxon. In the same place a summary of the available
chromosomal information and that on. self-incompatibility is pre-
sented. Only a digest of my earlier treatment of sects. Chylismia,
Lignothera, and Chylismiella (Raven, Univ. Calif. Publ. Bot. 34:
1-122. 1962) is presented here, but I have tried to include enough
information to make the present revision complete and usable taxo-
nomically without reference to the earlier paper. In the citation of
collections, R has consistently been used for Raven, except in the
case of type collections. Voucher specimens are understood to be

' As to the status accorded this and comparable g BroUups, see Brizicky, Journ.
Arn. Arb. 44: fn. 1963, and Gray, Man. Bot. N. U.S., ed. 2, p. xi., 1856.

RAVEN—GENUS CAMISSONIA 181

deposited in the Dudley Herbarium, Stanford University (DS), with
duplicates in the U.S. National Museum (US) and Rancho Santa
Ana Botanic Garden, Claremont, California (RSA), in most cases,
unless some other indication of their deposition is made.

Key to Sections

Ovary with a long slender sterile projection below the hypanthium, the plants
acaulescent or nearlyso... . ... . . Section IV. Tetrapteron
Ovary lacking a sterile projection, the plants only occasionally appearing acaules-
cent, and then when immature.

Seeds with a thick papillate wing surrounding the concave face (fig. 78); petals
white, yellow near the base. . . . . . . . . Section VIII. Chylismiella

Seeds not winged; petals yellow, white, or purple.
Capsules on well-defined pedicels, not coiled or contorted; seeds in 2 rows in

each locule.

Hypanthium 0.4-8 mm. long; leaves basal or cauline, often pinnately

divided or lobed; pollen shed singly . . . . . Section II, Chylismia
Hypanthium 4.5-40 mm. long; leaves all cauline, simple, usually cordate-
orbicular; pollen shed in tetrads. . . . . . Section III. Lignothera

Capsules sessile (see also 40. C. kernensis), often coiled or contorted; seeds in

1 row in each locule.

Petals white, the flowers vespertine . . . Section VII. Eremothera
Petals yellow, the flowers matinal.

Leaves pinnately lobed, more rarely subentire; petals usually with a
pattern of minute maroon flecks near the base; seeds often with
purple dots or blotches; capsules often sharply reflexed at maturity
(Baja California, Sonora, southern California, Arizona).

Section I. Eulobus

Leaves subentire; petals unspotted or with 1 or 2 bright red spots near
the base; seeds unspotted; capsules not reflexed (widespread).

Plants with naked, capillary stems, each bearing a crowded, leafy
inflorescence at its apex; capsule strongly flattened, 0.5-1 cm.
long... .. . . Seetion IX, Nematocaulis

Plants usually leafy from the base, the inflorescence not crowded;
capsule not flattened, usually over 1 cm. long.

Capsules quadrangular, at least when dry; plants usually with
flowers at the basal nodes; leaves mostly lanceolate to ovate;
seeds dull, flattened, usually over 1 mm. long.

Section V. Holostigma

Capsules subterete; plants lacking flowers at the basal nodes;
leaves mostly linear or very narrowly elliptic; seeds shining,
triangular in transection, often under 1 mm. long.

Section VI. Camissonia
SECTION I. EULOBUS

Stigma cylindrical, 1.4-4.5 mm. long; branches prostrate, radiating from a centra

axis (central and southern Baja California)... .. . 3.C. sceptrostigma
Stigma globose; branches erect.

Woody perennials, often flowering the first year; stigma held well above the

anthers at anthesis; leaves usually entire (west coast of Baja California).

1. C. crassifolia

182 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Annuals; stigma equaled by at least the longer stamens, their anthers surround-
ing it at anthesis; leaves usually deeply pinnatifid.
Capsules usually more or less contorted; hypanthium 3-4.2 mm. long; petals

10-19 mm. long (central Baja California) . . . . . . 2. C. angelorum
Capsules straight or nearly so, deflexed at maturity; hypanthium 0.6-1.5
mm. long; petals 6-14 mm. long (widespread). . . . 4. C. californica

SECTION II. CHYLISMIA

Stamens 4 (local in Mohave and Coconino Cos., Arizona). . . . . . 15. C. exilis
Stamens 8.
Corollas lavender in anthesis.
Stigma held above anthers at maturity; hypanthium 4,5-8.5 mm. long;

petals 9-13.8mm.long. . . . . . . . . . 16. C. megalantha
Stigma surrounded by anthers at maturity: hypanthium 2-5 mm. long; petals
2-6 mm. long... . . rane Wan OF heterochroma

Corollas yellow or white at anthesis.
Capsules distinctly clavate, more than 2 mm. thick (see also 9. C. walkeri).
Stigma surrounded by anthers at maturity, the petals less than 5.5 mm.
long.
Oil cells lining veins on underside of leaves pale yellowish brown, rarely
darker; leaves often simple and entire, in compact basal rosette.

Capsules to 30 mm. long; leaf blades to 5.5 em. long; base of plants
often nearly glabrous (Wyoming, Colorado, eastern Utah, New
Mexico)... . . . 13a. C. scapoidea subsp. scapoidea

Capsules 25-40 mm. long: leaf blades to 3.5 em. long; base of plants
densely strigose (Arizona) . 13b. C. scapoidea subsp. macrocarpa

Oil cells lining veins on underside of leaves dark brown, conspicuous;
leaves remotely dentate and often pinnately compound, in loose
basal rosette.

Leaves with several palrs of lateral leaflets; capsules 16-38 mm.
long; petals 4-8 mm. long (western Utah and adjacent Neveda),

13d. C. scapoidea subsp. utahensis

Leaves with lateral leaflets few or absent: capsules rarely over 20 mm.
long; petals 1.8-4 mm. long (Oregon, Idaho, western Utah, and
Nevada). . . . . . . . 18. C. scapoidea subsp. brachycarpa

Stigma held well above anthers at maturity, the petals usually more than
6 mm. long.
Mature pedicels and capsules sharply deflexed; corollas bright yellow.
11. C. munzii
Mature pedicels and capsules ascending or spreading; corollas yellow
or white.

Branches of the inflorescence numerous and _ filiform; leaves not
strictly basal; capsules less than 10 mm. long, the pedicels slender
(southwestern Utah and northwestern Arizona) . . 14. C. parryi

Branches of inflorescence not filiform; leaves mostly in a basal rosette;
capsules often more than 10 mim. long.

Stamens differentiated into 2 sets: inflorescence loose; leaves
entire or nearly so, rounded, often simple and glabrous (Colorado
and Utah)... 2... . 2. 12. C. eastwoodiae

Stamens subequal; infloresceuce dense; leaves generally dentate,
often acutely angled, pinnate and pubescent (Oregon and
Nevada southward).

RAVEN—GENUS CAMISSONIA 183

Lower parts of the plants with villous, spreading pubescence;
sepals with free caudate projections arising below apices.
Petals fading reddish (Arizona, Sonora, Baja California).

Petals 3-5 mm. long; style 7-12 mm. long (Arizona, and
Sonora). . . . . 10k. C. claviformis subsp. rubescens

Petals 1.5-2 mm. long; style 5-7 mm. long (Baja California)

10j. C. claviformis subsp. wigginsii

Petals not fading reddish (San Diego and Imperial Cos., Cali-

fornia, and adjacent Baja California)

10h. C. claviformis subsp. peirsonii

Lower parts of plants short-appressed pubescent or glabrous

but not villous with spreading trichomes; sepals entire or

with caudate projections.

Petals white, the hypanthium orange brown.

Plants variously pubescent on stems and in inflorescence;
lateral leaflets often reduced.

Lateral leaflets reduced in number, the leaves often nearly
simple; basal rosette compact (Nevada, southern
Oregon, local in Mono and Inyo Cos., California).

10c. C. claviformis subsp. integrior

Lateral leaflets generally well developed and numerous,
the basal rosette not compact.

Sepals often with projections arising below the apices;
terminal leaflets often large and nearly cordate;
buds and inflorescences often silky-strigose (Death
Valley region, California).

10d. C. claviformis subsp. funerea

Sepals usually entire; terminal leaflets usually incon-

spicuous; buds and leaflets not silvery.
Pubescence on leaves, stems, and often inflorescences
glandular (Arizona).
10g. C. claviformis subsp. peeblesii
Pubescence on leaves and stems rarely glandular, that
of inflorescences never so (southern Nevada,
California, rare in Arizona).
10f. C. claviformis subsp. aurantiaca

Buds and inflorescences usually glabrous or nearly so;
lateral leaflets usually well developed (western deserts
of California) . . 10e. C. claviformis subsp. claviformis

Petals yellow, the hypanthium yellow or orange brown.

Plants with strigose pubescence above and in the inflorescence,
the sepals sometimes with free projections (southern
California, Arizona, and northern Mexico).

10i. C. claviformis subsp. yumae

Plants usually glabrous above, the sepals entire; northern
part of species range.

Leaves lanceolate, narrow, and evenly dentate, often
nearly entire by loss of lateral leaflets; hypanthium
dark (Inyo and Mono Cos., California).

10b. C. claviformis subsp. lancifolia

Leaves pinnate, the terminal leaflets ovate, often blunt,
the lateral leaflets often well developed (Washoc
Co., Nevada, northward).

10a. C. claviformis subsp. cruciformis

184 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Capsules elongate, often linear, usually less than 2 mm. thick; pedicels often
inconspicuous.
Stigma surrounded by anthers at maturity; petals less than 6 mm. long;
style less than 6 mm. long; inflorescence erect in bud.

Clumped perennials with somewhat woody base; leaves glabrous or
sparsely pubescent; anthers glabrous (Colorado R. in and below
Grand Canyon, Arizona).

Leaves and style glabrous. . . 8a. C. speculicola subsp. speculicola

Leaves and style pubescent . . . 8b. C. speculicola subsp. hesperia

Annuals or short-lived perennials, the base usually not at all woody;
leaves pubescent, usually densely so; anthers pubescent or glabrous
(widespread). . . . . . .. . . . 9 C. walkeri

Stigma elevated above anthers at maturity: “petals usually more than 6

mm. long; style more than 6 mm. long.

Inflorescence erect or drooping, elongating before flowers open, the mature
buds mostly individually pendulous; plants virgate; hypanthium less
than 3 mm. long (Utah, southern Nevada, and northwestern Arizona).

7. C. multijuga

Inflorescence drooping, mostly elongating after flowers open; stems
stout, not virgate; hypanthium more than 3 mm. long.

Sepals with free caudate tips arising just below apices, glandular-
pubescent; inflorescence often branched; stamens differentiated
into 2 sets (Toroweap area, Mohave Co., Arizona).

5. C. confertiflora

Sepals lacking free caudate tips, or if they are present, the sepals
variously pubescent, but not glandular; inflorescence branched
or unbranched; stamens subequal (widespread) . 6. C. brevipes

SECTION III. LIGNOTHERA

Hypanthium 18-40 mm. long; style 30-58 mm. long. . . . 19. C. arenaria
Hypanthium 4.5-14 mm. long; style 8-23 mm. long.
Leaves ovate-acuminate, longer than broad; petals less than 4.5 mm. long;
pubescence predominantly glandular.
18. C. cardiophylla subsp. cedrosensis
Leaves cordate to cordate-orbicular, about as long as broad; petals mostly more
than 4.5 mm. long.
Plants coarse; pubescence mostly glandular (Inyo Co., California).
18b. C. cardiophylla subsp. robusta
Plants more slender; pubescence mostly villous (San Bernardino Co.,
California, southward and eastward).
18a. C. cardiophylla subsp. cardiophylla

SECTION IV. TETRAPTERON

Plants perennial, strictly acaulescent; capsule not winged.

Leaves pinnatifid, the plants more or less densely pilose or appressed pubescent.
Style 9.5-20 (-25) mm. long, the stigma held above the anthers at anthesis;

petals (8-) 10-23 mm. long.
Less than 5 percent of pollen 4-pored; pubescence variable (widespread).
22a. C. tanacetifolia subsp. tanacetifolia
More than 10 percent of pollen 4-pored; plants densely gray-pubescent
(Sierra Valley area in Plumas and immediately adjacent Lassen and
Sierra Cos., California) . 22b. C. tanacetifolia subsp. quadriperforata
Style 3-6.5 mm. long, the stigma surrounded by anthers of longer stamens
at anthesis; petals 5.5-7 (-8) mm. long. . . . . . . 23. C. breviflora

RAVEN—-GENUS CAMISSONIA 185

Leaves not pinnatifid or, if so, plants subglabrous.
Leaves more or less densely white-ciliate; capsule subterete, the thin walls
much distended by the seeds . . . 2... . . +. . + 20. C. ovata
Leaves subglabrous, rarely minutely and sparsely strigulose; capsule more or
less 4-angled, the nearly flat walls not distended by the seeds.
21. C. subacaulis
Plants annual, sometimes branching; capsule broadly winged in upper half.
Petals 5-18 mm. long; style 3-5.5 mm. long; pubescence pilose.
24. C. graciliflora
Petals 2-3.5 mm. long; style 1-2.2 mm. long; pubescence strigose . 25. C. palmeri

SECTION V. HOLOSTIGMA

Plants perennial, often somewhat woody near the base.

Petals (10-)12-20 mm. long; styles 13-23 mm. long (near Goleta, Santa Barbara
Co., California, south to near San Quintin, Baja California; San Nicolas
and San Martin Isls.) ... 26a. C. cheiranthifolia subsp. suffruticosa

Petals 6-11 mm. long; styles 6-9 mm. long (Point Conception, Santa Barbara
Co., California, north to Coos Bay, Curry Co., Oregon; several islands).

26b. C. cheiranthifolia subsp. cheiranthifolia
Plants annual (except for a few populations of C. bistorta, q.v.).

Stigma evidently held above the anthers at anthesis, the flowers large (southern
California, north to Ventura Co., and adjacent Baja California).

27. C. bistorta

Stigma surrounded by at least the anthers of the longer set of stamens at
anthesis, the flowers small to large (widespread).

Capsule very heavy, 2.8-3.5 mm. thick near the base, straight or nearly so,
deeply grooved along the lines of dehiscence (Guadalupe and San Cle-
mente Isls.).

Pubescence strigose; petals not spotted (Guadalupe Isl.).
28a. C. guadalupensis subsp. guadalupensis
Pubescence spreading; petals with a prominent red dot near the base
(San Clemente Isl.) . . . 28b. C. guadalupensis subsp. clementina

Capsule 0.75-2.2 mm. thick near the base, straight or curved into 1 or more
spirals, not deeply grooved (widespread).

Pollen 25-100 percent 4- or 5-pored; hexaploid species.

Inflorescence lacking glandular trichomes; 4-pored pollen comprising
25-60 percent of the total (Santa Lucia Mts., Monterey Co., and
inner Coast Ranges south to Santa Barbara Co., California).

38. C. luciae

Inflorescence with an understory of short glandular trichomes; 4- and
5-pored pollen comprising 70-100 percent of the total (not in area
of C. luctae).

Capsule 1.3-1.6 mm. thick, subterete in transection (southernmost
Monterey to central San Luis Obispo Co., California).

37. C. hardhamiae

Capsule 1,5-2 mm. thick, more or less quadrangular in transection

(San Diego Co., California, and adjacent Baja California; off-

shore islands) ....... . 2... 4... . 39. C. robusta

Pollen up to 5 percent 4-pored (rarely more in C. intermedia); diploid

or tetraploid species.

Capsule markedly quadrangular in transection, 1.8-2.2 mm. thick
(Los Angeles Co., California, south throughout the northwestern
quarter of Baja California) . . 2... 2... . . 29 ¢. lewisii

186 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Capsule not markedly quadrangular in transection, at least in living
material, 0.75—1.2 (—1.8) mm. thick (widespread).

Plants subglabrous, usually with 50 or more flowers in the basal

rosette, subcaespitose or with very thick prostrate branches from

the base (coastal Baja California)... . . . 30. C. proavita
Plants subglabrous to densely villous, with few flowers in basal rosette
(widespread).

Upper leaves evidently petiolate, attenuate at the base; capsule
usually more contorted; plants subglabrous to strigulose, often
reddish (Yolo Co., California, to 30° N. lat. in Baja California;
Madera Co., California; Santa Cruz Isl.) . . . 33. C. ignota

Upper leaves usually not evidently petiolate, rounded or truncate
at the base; capsule straight to coiled once or twice; plants strig-
ulose to villous, always evidently so (widespread).

Conspicuously gray-strigose plants of the deserts, the branches
usually decumbent (deserts of southern California to Arizona
and northern Baja California).

Petals 6.5-13 mm, long; style 6.5-10.5 mm. long (western
Riverside and San Bernardino Cos., California).

34a. C. pallida subsp. hallii

Petals (2-) 3.5-6 (-8) mm. long; style (2.1-) 3-6.5 mm. long

(widespread on deserts) . . 34b. C. pallida subsp. pallida

Plants not conspicuously gray-strigose, barely reaching the mar-

gins of the deserts (except for C. confusa in central Arizona).
Capsule 0.75-0.9 mm. thick; upper leaves narrowly ovate or
ovate; plants erect; n=7... . . . . 82. C. hirtella
Capsule 0.9-1.2 (-1.8) mm. thick; upper leaves narrowly
lanceolate to narrowly ovate; plants erect or decumbent;
n=7 or 14.

Plants with decumbent branches from the basal rosette;
upper leaves usually narrowly lanceolate; short glandu-
lar trichomes usually absent in inflorescence; n=7
(Coast Ranges of California, Sonoma Co. to San Diego
Co). 2... 2... 81. CG. omicrantha

Plants erect; upper leaves lanceolate or narrowly ovate; short
glandular trichomes almost always present in inflores-
cence; n= 14 (widespread).

Plants gray-villous; petals (2.5-) 5-10.5 mm. long; style

2.5-) 4.5-7.5 mm. long (San Luis Obispo Co. to San
Diego Co., California; central Arizona).

36. C. confusa

Plants usually not gray-villous; petals 1.5-3.5 (-4.5) mm.

long; style 2-3.5 mm. long (Lake and Yolo Cos.,

California, to 30° N. lat. in Baja California; Santa

Cruz and Santa Catalina Isls.) . . 35. C. intermedia

SECTION VI. CAMISSONIA
Plants of North America.
Sepals reflexed singly; plants often with conspicuous spreading pubescence
(Great Basin and neighboring areas).
Stigma elevated well above the anthers at anthesis; petals 8-15 (—18) mm.
ong.

RAVEN—GENUS CAMISSONIA 187

Plants compact, the leaves clustered at the base; pubescence dense,
spreading; fruiting pedicel 3-15 mm. long (northeastern Kern Co.,
California) . . . .. . . . 40a. C. kernensis subsp. kernensis

Plants not compact, the leay es not clustered at the base; pubescence vari-
able; fruiting pedicel 0-5 (-15) mm. long (Kern, Inyo, Mono, and San
Bernardino Cos., California, and adjacent Neveda).

40b. C. kernensis subsp. gilmanii
Stigma surrounded by the anthers at anthesis; petals 1.8-4 mm. long.
Basal rosette well developed, the plants usually with long, spreading tri-

chomes.
Leaves 0.4-1.8 mm. wide; hypanthium 0.8-1.6 mm. long; style 1.6-38.2
mm. long;n=7. . ... . . 41. C. pusilla
Leaves 2-5 (-6.2) mm. wide: hypanthium 1. 3- 3 mm. long; style 3.2-4.1
mm.long;n=14..... ... . . 42. C. pubens
Basal rosette not well developed, the stems bare near the base; plants
usually with appressed pubescence or subglabrous . . 43. C. parvula

Sepals usually reflexed in pairs, remaining attached (widespread).
Stigma held above the anthers at anthesis; petals (3.5-) 4-15.5 mm. long.
Leaves lanceolate or narrowly ovate, entire or nearly so (Sierra Nevada,
California, in Madera and Mariposa Cos.)
45a. C. sierrae subsp. sierrae
Leaves linear to narrowly elliptic or narrowly oblanceolate, serrulate to
coarsely serrate (widespread in California).

Leaves linear to narrowly elliptic or narrowly oblanceolate, serrulate or
serrate; stems usually erect (widespread in central and southern
California)... ........ 44a. C. campestris subsp. campestris

Leaves narrowly elliptic, coarsely serrate; plants usually decumbent
(southern Monterey to northern Santa Barbara Co., California).

44b. C. campestris subsp. obispoensis
Stigma surrounded by the anthers at anthesis.
Leaves entire, very rarely with 1 or 2 small teeth; pubescence dense,
strigulose (upper Kern R. drainage, California) . . 50. C. integrifolia
Leaves serrulate or serrate; pubescence variable (widespread).

Leaves narrowly elliptic, rounded or obtuse at the base; n=7 (central
Sierra Nevada, California).

Petals 4-7 mm. long; style 4.5-7 mm. (Madera and Mariposa Cos.,
California) . . . .. . . . 45a. C. sierrae subsp. sierrae

Petals 2.2-4 mm. long; style 2.8-5 mm. long (Fresno Co., California).

45b. C. sierrae subsp. alticola

Leaves linear or very narrowly elliptic, narrowly cuneate or attenuate at
the base; n= 14, 21 (widespread).

Often more than 30 percent of the pollen grains 4-pored; plants us-
ually with spreading pubescence on the stems; n= 21 (widespread
in California south to Kern Co.) ... . . . 51. C. contorta

Very rarely with more than 10 percent of the pollen grains 4-pored;
pubescence variable; n= 14 (California from Lake Co. southward,
and adjacent Baja California).

Petals (4-)4.5-7 mm. long; style 4-7 mm. long; pubescence of stems
long, spreading (Lake Co. and Sierra Nevada foothills from
Eldorado Co. to Fresno Co., California). . . 46. C. lacustris

Petals 2.1-4.2(-4.5) mm. long; style 2.3-4.8 mm. long; pubescence
variable (Coast Ranges from Sonoma Co., California, south to
northern Baja California, and Sierra Nevada of Kern Co.,
California).

188 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Plants villous, the inflorescence glandular (serpentine soils, San
Benito Co., California) . . .. . . . . 47. C. benitensis
Plants strigulose or with villous trichomes on the lower stem, rarely
villous and glandular only (widespread in range given above).

48. C. strigulosa

Plants of South America.
Leaves linear or very narrowly lanceolate; plants erect, decumbent, or prostrate;
capsules 0.6-0.9 mm. thick (widespread). . 49a. C. dentata subsp. dentata
Leaves broadly elliptical, thick; plants prostrate; capsules 1.1-1.3 mm. thick,
heavy (coastal sands in Prov. Concepcién, Aruco, and Cautin, Chile).
49b. C. dentata subsp. littoralis

SECTION VII. EREMOTHERA

Capsules not thickened near the base; plants lacking flowers at the lower nodes.
Petals 3.5-7 mm. long; stigma held well above the anthers at anthesis.
52. C. refracta

Petals 1.8-3 mm. long; stigma surrounded by the anthers at anthesis.
53. C. chamaenerioides
Capsules thickened near the base; plants with or without flowers at the lower
nodes.
Inflorescence erect to the tip at anthesis; plants flowering from the base.
Petals 3-5 mm. long; style 6-7 mm. long, glabrous (west-central Nevada).
57. C. nevadensis
Petals 0.8-1.3 mm. long; style 1.2-3.2 mm. long, pubescent near the base
(widespread). . . 2... 2... ee 58. C. minor
Inflorescence nodding at anthesis; plants flowering from near the base or not.
Petals (38-) 4-7.5 mm. long; stigma held well above the anthers at anthesis.
Cluster of large basal leaves prominent at the time of flowering, the in-
florescence relatively leafless; plants blooming in the spring, never
with prominent spreading pubescence.

Capsules 2-3.8 mm. thick near the base, woody, the inflorescence very
condensed (Sonoran and eastern Mojave Desert; San Juan Co.,
Utah)... ..... =... 54e. C. boothii subsp. condensata

Capsules 1-2.3 mm. thick near the base, not woody, the inflorescence
not highly condensed.

Capsules 1.7—2.3 mm. thick near the base, curved outward (cismontane
central and southern California, south to Los Angeles Co.).

54a. C. boothii subsp. decorticans

Capsules 1-1.6 mm. thick near the base, often curved downward
(western Mojave Desert of California; upper Kern R. drainage).

54b. C. boothii subsp. desertorum

Cluster of basal leaves rarely prominent at time of flowering, the inflo-

rescence leafy; plants blooming in summer or late spring, often with
prominent spreading pubescence.

Plants villous and glandular pubescent, blooming in the summer.

Plants usually 1.5-4 dm. tall, robust; cauline leaves coarsely serrulate,
lanceolate or narrowly ovate (Mono Co., California, north to
castern Washington and southwestern Idaho; Mohave Co.,
Arizona) 2... . «6546. C. boothii subsp. boothii

Plants usually 0.5-2 dm. tall; cauline leaves serrulate, narrowly
lanceolate or lanceolate (southern Lander and eastern Churchill
Cos., Nevada, south to northeastern Inyo Co., California, and
southern Nye Co., Nevada; Kingston Range, San Bernardino
Co., California). . . . . . . 54e. C. boothii subsp. intermedia

RAVEN—GENUS CAMISSONIA 189

Plants strigose, sometimes densely so, rarely villous or with an admixture
of glandular trichomes; plants blooming in late spring and early
summer (eastern Oregon and southern Idaho south to Lassen Co.,
California, through eastern Nevada to northern Clark Co., and in
western Utah) ... .. ... . 54d. C. boothii subsp. alyssoides

Petals 1.5-2.5 mm. long; stigma surrounded by the anthers at anthesis.
Pubescence villous and glandular; seeds dimorphic, the basal ones coarsely
papillose (eastern Washington, eastern Oregon, southern Idaho).
55. C. pygmaea
Pubescence glandular; seeds monomorphic, all smooth or nearly so (south-
western Utah and Coconino Co., Arizona). . .. . . 56. C. gouldii

SECTION VIII. CHYLISMIELLA

One species. . . 2... ee ee ee ee ew. 659. C. pterosperma

SECTION IX. NEMATOCAULIS

Petals 2.5-5 mm. long; style 4.5-6 mm. long (eastern Washington and along the
Columbia R.) 2... 2. ee ee. 660. C. hil gardii
Petals 0.8-2.3 mm. long; style 1.7-3 mm. long (widespread) . . . 61. C. andina

Section I. Eulobus

Camissonia sect. Eulobus (Nutt. ex Torr. & Gray) Raven, Brittonia 16: 283.
1964.
Eulobus Nutt. ex Torr. & Gray, Fl. N. Amer. 1: 514. 1840.
Oenothera sect. Eulobus (Nutt. ex Torr. & Gray) Baill., Hist. Pl. 6: 461.
1877.
Oenothera subg. Eulobus (Nutt. ex Torr. & Gray) Munz, Amer. Journ.
Bot. 16: 254. 1929. .

Woody subshrubs or annuals, the plants caulescent, flowering from
near the base or not. Inflorescence erect. Flowers opening near sunrise.
Ovary lacking a sterile projection. Hypanthium lined with a lobed,
red brown, fleshy dise within or with a smaller, circular dise at the
base of the style. Sepals reflexed separately or united. Petals yellow,
usually finely flecked with red near the base, this area of the petals
not ultraviolet reflective, the remainder of the petals strongly so;
petals fading orange red after pollination. Stamens 8, subequal or
with the epipetalous ones shorter; anthers versatile, villous or gla-
brous. Pollen shed singly. Capsule sessile, regularly and promptly
loculicidal, the midrib of each valve yellowish brown, prominent,
straight or contorted, somewhat torulose, subterete in living material,
the seeds in 1 row in each locule. Seeds monomorphic, narrowly
obovoid, more or less triangular in transection, pointing up or down
in capsule, finely papillose.

TYPE sPEciES: Camissonia californica (Nutt. ex Torr. & Gray)
Raven.

DisTRIBUTION: Sandy slopes and flats, from southern San Benito,
central Kern, and eastern San Bernardino Counties, California, and
western and southern Arizona (from the vicinity of Kingman south),

190 OONTRIBUTIONS FROM THE NATIONAL HERBARIUM

south-central Sonora at about 28°20’ N. lat., and in Baja California to
the vicinity of Bahia de la Magdalena at about 24°30’ N.; Santa Cruz
and Santa Catalina Islands, California; Isla de Cedros, Baja Cali-
fornia; and Isla San Lorenzo, Golfo de California.

Camissonia sect. Mulobus is here taken in exactly the same sense
in which it was constituted by Munz in 1929 (Amer. Journ. Bot.
16: 253-257). At the time he prepared this revision, Dr. Munz re-
marked that herbarium material of three of the four species of this
eroup was meager. In spite of this, the examination of the abundant
material that has been collected in the succeeding 40 years has not
given me any reason to disagree with the delimitation of species
presented in 1929. I have, however, cited specimens to document the
ranges of the three species endemic to Baja California, as this has
not been possible until recently.

The woody Camissonia crassifolia appears to be the most generalized
species within the group. It is one of the two self-incompatible species
in the section and the only one known to be regularly associated with
a group of oligolectic bees, one of which is Andrena (Onagrandrena)
eulobt Linsley & MacSwain. Camissonia angelorum is closely related to
and doubtless best thought of as an annual, self-pollinated derivative
of C. crassifolia. More distantly related, but evidently derivable from
the same ancestral stock, are the self-incompatible C. sceptrostigma,
with its unique cylindrical stigma, of a form unknown elsewhere in
Onagreae and the regularly self-pollinating C. californica, the most
widespread species of the section and the only one to contain tetraploid
populations. Camissonia californica, probably the most specialized
species in the section, is rarely visited by insects.

In sect. Hulobus, 108 individuals have been examined chromo-
somally, representing 78 populations. Not one of these has had a
multivalent association of chromosomes in meiosis, all being strictly
pair-forming. In this respect, sect. Hudobus resembles sect. Holostigma;
reciprocal translocations appear to play no adaptive role in natural
populations, and multivalents have not been observed either in
diploid or in polyploid populations, even though it is highly probable
that some of the polyploids combine relatively homologous diploid
genomes (see Appendix A, p. 381).

1. Camissonia crassifolia (Greene) Raven, Brittonia 16: 283. 1964.
Oenothera crassifolia Greene, Bull. Calif. Acad. 1: 188. 1885.

Erect well-branched shrub up to 1.4 m, tall, or sometimes prostrate ;
plants flowering the first vear, and then as little as 1.5 em. tall, with
no distinct basal rosette, sometimes flowering at the basal nodes;
plants subglabrous or with a few strigulose trichomes in the inflores-
cence, especially around the hypanthium and on the upper portions
of the ovary. Leaves somewhat fleshy and glaucescent, narrowly

RAVEN—GENUS CAMISSONIA 191

elliptic to narrowly lanceolate, rarely very narrowly elliptic, 1-6 em.
long, 2.5-12 mm. wide, entire to rather coarsely serrate or crenate,
the apex obtuse, the base cuneate; leaves subsessile or with a petiole
1-2 mm. long. Inflorescence long and virgate. Hypanthium 1.5-2.5
mm. long, 2-3 mm. across at the summit, densely villous within.
Sepals 5-9 mm. long, 1.2-2 mm. wide, reflexed in pairs or groups of
3 or 4. Petals (5.5-) 7.5-15 mm. long, (4—) 6.5-13 mm. wide, finely
flecked with red in the basal third. Filaments of the episepalous
stamens 3-7 mm. long, those of the epipetalous ones 2-5 mm. long;
anthers 2.5-4 mm. long, sparsely long-ciliate or glabrous. Style (7—)
§.5-12 mm. long, densely villous in basal half; stigma globose, 0.8—1.1
mm. thick, held well above the stamens at anthesis. Capsules ir-
regularly contorted, ca. 1.5-3.5 cm. long, 1.2-2 mm. thick. Seeds 1.7—2
mm. long, 0.7-0.8 mm. thick, brownish with small purple dots, often
distorted or curved by the tough capsule walls. Gametic chromosome
number, n=7. Self-incompatible.

Type: Cabo San Quintin, Baja California, 10 May 1885, FE. L.
Greene (ND; isotypes, BM, CAS, DS, F, GH, K, NY, US).

DistTriBUTION (Figure 4): Sandy soil, especially on beaches, or
rocky headlands, along the west coast of Baja California from just
north of San Isidro, at about 31°20’ N. lat., to the region of Bahia
de la Magdalena, at about 24°40’ N. lat.; ranging away from the
immediate coast only in the region of the Desierto Vizcaffio at about
27° N. lat. Also on Isla de Cedros and Isla Santa Magdalena. From
near sea level to a few hundred feet in elevation.

Vouchers for chromosome number (5 individuals, 5 populations),
n=T:

BAJA CALIFORNIA: 3 mi. N. of San Quintin, Breedlove 4173 (progeny) ;
20.6 mi. 8. of road to San Quintin, R12391 (K, LA, UC); Pabellon Beach, ca.
10 mi. S. of San Quintin, Klein et al. 13 (RSA, UC); Punta Baja, Klein & Gregory
1236; Isla Cedros, Moran 10763.

REPRESENTATIVE SPECIMENS EXAMINED:

BAJA CALIFORNIA: About 5 mi. N. of San Isidro, Tillett 799 (RSA); San
Antonio del Mar (Johnson Ranch), Chambers 704 (DS, UC); Santo Domingo
(Hamilton Ranch), Nelson & Goldman 7174 (US); W. of Bahia de San Quintin,
Wiggins 11890 (DAO, DS, OSC, SD, UC); 14 mi. 8. of San Quintin, Raven 17024
(RM, RSA, UC); Punta Baja, Harbison in 1950 (SD); Santa Catarina Landing,
Wiggins 4438 (DS, GH, POM, US); ca. 14 mi. 8. of Puerto San José, Wiggins
& Thomas 207 (DS): Playa Maria, Street in 1875 (GH); 28 mi. SW. of Punta
Prieta, Haines & Stewart in 19385 (ARIZ, DS, GH, K, LA, NY); Lagoon Head,
1889, Palmer 769 (GH, K, US); Viscaffio Depression 8. and W. of Laguna Scam-
mon, Gentry 7353 (ARIZ, DS, RSA, UC); Picachos de Santa Clara, Geniry 7740
(ARIZ, DS, UC); stabilized dunes 13 mi. W. of Rancho San Angel, Wiggins
16267 (DS); 1.6 km. E. of Laguna de San Ignacio, 59 km. SW. of San Ignacio,
Carter et al. 2515 (DS); Punta Santo Domingo, Nelson & Goldman 7174 (POM);
San Juanico, Street in 1876 (GH). Cepros IsLanp: Slope above low beach cliffs,
southwest coast, NNW. of Cerro Redondo, Moran 10772 (DS, SD); behind

295-655 O—68

192 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

beach NNW. of Cerro Redondo, Moran 10763 (DS, 8D); W. coast 0.5-1 mi.
NE. of Red Rock, Haines & Hale in 1939 (ARIZ, CAS, COLO, DS, POM, SD,
UC). Isna Santa Magpauena: Barclay 3126 (BM, RSA), Orcutt 28 (GH, Us).
Mangrove I., Bahia de la Magdalena, Greene in 1988 (SD).

Colonies of Camissonia crassifolia may be variable in the degree of
lobing of the leaves; thus Gregory 352 (RSA), from Pabellon Beach,
has coarsely serrate leaves and Gregory 350 (RSA), which grew with
it, has entire leaves. Collections with deeply lobed leaves are rather
common, those of Palmer and Street cited above being extreme in
this respect. Plants growing on the beach at certain localities are
prostrate and distinctive in appearance. Camissonia crassifolia grows
sympatrically with tetraploid populations of C. californica.

Plants from Isla de Cedros, where the species is local, flower regularly
the first year. The flowers are borne in the rather compact rosette of
deeply lobed leaves, and the plants only occasionally persist to flower a
second year. A progeny of Moran 10763, from Isla de Cedros, and one
from Breedlove 4173, from near San Quintin, on the mainland, were
grown at Stanford, and both were found to consist entirely of self-in-
compatible plants. Reciprocal hybrids were made between these two
strains and grown to maturity. In the F, individuals, the stems elon-
gated rapidly and no flowers were formed at the basal nodes; in both of
these respects they resembled the mainland parent. The F, individuals
were self-incompatible like their parents and had 86 percent stainable
pollen (based on a sample of 200 grains in cotton blue in lactophenol),
compared with about 90 percent in most nonhybrid individuals. In
meiosis, they formed 7 pairs of chromosomes, including 6 ring bivalents,
indicating nearly complete homology between the chromosomes of
the parental plants.

2. Camissonia angelorum (S. Wats.) Raven, Brittonia 16: 283. 1964.
Ficur_Es l, 2
Oenothera angelorum S. Wats., Proc. Amer. Acad. 24: 49. 31 January 1889.
Oenothera crassiuscula Greene, Pittonia 1: 290. 16 April 1889. Type: Low
plains around the southern shore of Bahfa de San Bertélome, Baja Cali-
fornia, March 1889, Lieutenant Pond (location of specimen not known).
Sphaerostigma angelorum (S. Wats.) A. Nels., Bot. Gaz. 40: 63. 1905.

Erect, sparingly branched, virgate annual 8-90 em. tall, with no
distinct basal rosette, not flowering at the basal nodes, glabrous
or sparsely strigulose, the inflorescence strigulose. Leaves deeply
pinnatifid, very narrowly elliptic, 1-7.5 em. long, 1-15 mm. wide, the
apex acuminate, the base attenuate; leaves subsessile or with a petiole
up to 3 mm. long. Inflorescence long and virgate. Hypanthium 3-4.2
mm. long, 3-4.5 mm. across at the summit, sparsely pubescent or
subglabrous in lower half within. Sepals 6.5-11 mm. long, 1.2-2.2 mm.
wide, reflexed in pairs or groups of 3 or 4. Petals 10-19 mm. long, 8-21
mm. wide, finely flecked with red in basal third. Filaments of the

RAVEN—GENUS CAMISSONIA 193

(MW)

Ficures 1-3.—Species of Camissonia sect. Eulobus, * 34: 1-2, C. angelorum: 1, Portion of
a fruiting branch (Wiggins 18189, DS); 2, a flowering plant (Wiggins @ Thomas 299,
DS). 3, Flowering branch of C. sceptrostigma (Wiggins 16824, DS).

194 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

episepalous stamens 3-5 mm. long, those of the epipetalous ones
2-3.5 mm. long; anthers 2-3.2 mm. long, long-ciliate. Style 5-9 mm.
long, subglabrous or with scattered erect trichomes near its base;
stigma globose, 1.3-2 mm. thick, held at the level of the anthers of the
longer stamens at anthesis, or perhaps a little above them in certain
plants. Capsules contorted, but often not as much so as in C. erasst-
folia, 1.5~3.5 em. long, 1-1.4 mm. thick. Seeds as in (. crassifolia.
Gametic chromosome number, n=7. Self-compatible and capable of
self-pollination, but probably often outerossed in nature.

Type: Bahja de los Angeles, Baja California, 22 November to 20
December 1887, #. Palmer 519 (GH; isotypes, BM, C, K, NY, UC,
US).

Distripution (Figure 4): Sandy flats and washes in Baja Cali-
fornia, ranging across the peninsula, from the vicinity of Punta Final
on the gulf coast at about 29°45’ N. lat. south to the northern edges
of the Desierto Vizcafiio at about 27°40’ N. lat.; also on Isla San
Lorenzo in the Golfo de California. From sea level to about 1,500
ft. alt.

Vouchers for chromosome number (9 individuals, 9 populations),
n=7 (all with 7 pairs):

BAJA CALIFORNIA: 2 mi. inland from Ensenada de San Francisquito,
Moran 12391 (DS, SD); 17 mi. S. of Laguna Seca Chapala, Thorne & Hendrickson
32640 (DS, RSA); Salorio, Moran 12314 (DS, SD); ca. 10 mi. E. of Bahia de los
Angeles, ca. 1,500 ft., Weggins 14808 (DS, progeny); Bahfa de los Angeles, Moran
12298 (DS, SD); just N. of San Borja, Moran 12461 (DS, SD); Bahia de San
Francisquito, Moran 12618 (DS, SD); Arroyo Salinito, Moran 12860 (DS, SD);
El Datillar, 8 mi. N. of Rancho Miramar, Moran 12847 (DS, SD).

REPRESENTATIVE SPECIMENS EXAMINED:

BAJA CALIFORNIA: 25 mi. N. of Rancho Laguna Chapala on road to San
Felipe, 1,180 ft., Wiggins & Ernst 662 (DS, UC); just W. of Bahia de San Luis
Gonzaga, J. L. & D. B. Wiggins 15906 (DS); 25 mi. N. of Punta Prieta, Wiggins
9361 (CAS, DS, GH, NY, POM, RM, UC, US, WTU); 20 mi. E. of Punta
Prieta, Wiggins 7662 (DS, GH, UC, US); Bahia de Los Angeles, Wiggins 7692
(DS, GH, LA, UC, US); near San Andreas, Wiggins & Thomas 215 (DS); 18
mi. 8. of Punta Prieta, J. L. & D. B. Wiggins 18189 (CAS, DS); 12 mi. NE. of
Rancho Rosarito on road to San Borja, Porter 559 (CAS, DS, GH, RSA); Bahfa
de San Francisquito, Johnston 3572 (CAS, GH, K, US); Rancho Barril, Wiggins
7822 (Ds, F, GH, US); Laguna de Guerrero Negro, Thomas 8508 (DS); 21 mi.
E. of Laguna de Guerrero Negro on road to El Arco, Wiggins 16828 (DS); 40 mi.
inland from Lagoon Head, 1889, Palmer 771 (GH, K, POM, US); & mi. W. of
Calmalli, Haines & Stewart in 1935 (ARIZ, Ds, GH, k, LA, NY, POM, RM,
RSA, UC, WTU); El Canton Rancho near Calmalli, Haines & Stewart in 1935
(ARIZ, Ds, F, GH, K, LA, POM, RM, RSA, UC); 7 mi. SE. of Rancho La
Cantina, Wiggins 16209C (DS): Bahia de San Bertélome, Rose 16208 (GH, US);
5.5 mi. N. of El Huizache, Vizeatiio Desert, Porter 541 (CAS, DS, RSA). Sourn
San Lorenzo Isuanp: Moran 4130 (DS, UC)

RAVEN—GENUS CAMISSONIA 195

Ficure 4.—Baja California, showing ranges of Camissonia sect. Eulobus: @=C. crassifolia;
O=C. angelorum; A=C. sceptrostigma.

196 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

The collection cited from Bahia de San Bertédlome has deeply
cut leaves and very probably is similar to the unlocated type of
Oenothera crassiuscula, from the same region. Camissonia angelorum
is rather obviously an annual derivative of (. crassifolia and certain
populations, notably some to those from the Desierto Vizeaitio region
at the southern limits of @. angelorum, can be assigned to one species
or the other only with the greatest difficulty. The two species might
be regarded as geographical subspecies, but the shift in breeding
system and ensemble of morphological features distinctive of each
are sufficient to make their continued recognition as species desirable,
in my opinion. Progeny of Wiggins 14808, grown at Stanford, set
seed abundantly in the absence of insects and when isolated from one
another; nevertheless, the showy flowers of Camissonia angelorum
suggest strongly that this species must at least occasionally be cross-
pollinated by insects in nature. Camissonia angelorum has been
observed growing sympatrically with tetraploid populations of (.
californica.

3. Camissonia sceptrostigma (T. 8. Brandeg.) Raven, Brittonin 16: 283. 1964.
FIGURE 3
Oenothera sceptrostigma T. 8. Brandeg., Proce. Calif. Acad. IL. 2: 156. 1889.

Rosette-flowering annual, with prostrate branches tardily elongat-
ing from the base, these to at least 35 em. long, with the leaves,
flowers, and capsules standing erect from them; entire plant finely
strigulose, often grayish-pubescent. Leaves thin, deeply and irregu-
larly pinnatifid with narrow lobes, narrowly elliptic, 1.5-16 em. long,
4-27 mm. thick, the apex acuminate, the base attenuate. Hypanthium
3.2-6 mm. long, 2.5-5 mm. across at the summit, glabrous within.
Sepals 3-8.5 mm. long, 1-2.4 mm. wide, spreading (not reflexed) ,
separate from one another. Petals 6.5-17.5 mm. long, 8-19 mm. wide,
with diffuse red flecking near the base. Filaments of the episepalous
stamens 1.7-4.5 mm. long, those of the epipetalous ones 0.8—1.8 mm.
long; anthers 1.2-2.5 mm. long, glabrous or finely pubescent, held
erect. Style 3.5-8.3 mm. long, glabrous; stigma cylindrical, 1.4—4.5
mm. long, 0.4-1.2 mm. thick, held + above the anthers at anthesis.
Capsules in the rosette straight or nearly so, those of the branches
somewhat flextious, 1.7-4 em. long, 1.5-1.8 mm. thick. Seeds 0.8—1.2
mm. long, 0.4-0.6 mim. thick, pale brownish, not spotted. Gametic
chromosome number, n=7. Self-incompatible.

Tyre: San Gregorio, Baja California, 1 February 1889, 7S.
Brandegee (CAS; isotypes, DS, PH, Us).

Disrrinution (Figure 4): Sandy fields and washes in central and
southern Baja California, from 11 miles south of Millers Landing,
at about 28°20’ N, lat., to the south end of Bahia de la Magdalena, at
about 24°30’ N. From near sea level to ca, 1,200 ft. alt.

RAVEN—GENUS CAMISSONIA 197

Voucher for chromosome count (2 individuals, 2 populations; n=7) :

BAJA CALIFORNIA: 314 mi. W. of Rancho Miramar, 28°08’ N. lat., 1138°40’
W. long., ca. 150 m, Moran 12857 (DS, SD); Verity in 1967 (R68-302; DS).

ADDITIONAL SPECIMENS EXAMINED:

BAJA CALIFORNIA: 11 mi. 8. of Millers Landing, Wiggins 15099 (DS); 3
mi. W. of main road and ea. 4-5 mi. N. of Rancho Mesquital, Wiggins 16768
(DS) ; 24 mi. E. of Laguna de Guerrero Negro, Wiggins 16824 (DS); Lagoon Head,
1889, Palmer 772 (C, CAS, GH, K, NY, UC, US); 23.2 mi. NW. of El Arco,
Porter 207 (DS, GH); 11 mi. W. of El Arco, Wiggins 1683A (DS); El Arco, E'p-
ling & Robinson in 1935 (DS); 10 mi. E. of El Soltero, Wiggins 15148 (DS); 15 mi.
S. of El Arco, Wiggins 15156 (DS, UC); 22 mi. 8. of Pozo Aleman, Wiggins 7849
(DS, F, GH, UC, US); Arroyo de Tecolote, Desierto Vizcaino, Gentry 7835
(ARIZ, DS, UC); Picachos de Santa Clara, Desierto Vizcaino, Gentry 7759
(ARIZ, DS, UC); 5-7 mi. E. of Los Picachos de Santa Clara, Gentry 7593 (RSA,
UC); 15 mi. E. of El Soliyo, Wiggins 15148 (UC); 7 mi. S. of Los Angeles, San
Ignacio to El Arco road, Wiggins & Ernst 626 (DS, UC); E. edge of Desierto
Vizcaino, 56 km. 8. of El Arco, Carter et al. 1940 (SD); 40 mi. W. of San Ignacio,
Hake in 1921 (DS); 50 km. NW. of San Ignacio, Carter et al. 1960 (DS, F, GH, K
SD, US); 25 mi. N. of San Ignacio, Wiggins & Ernst 615 (DS, UC); 14 mi. N. of
San Ignacio, Reed 6203 (POM); 24 mi. N. of La Purisima, Wiggins & Ernst 589
(DS, UC); Punta San Juanico, Moran 3524 (DS); Santo Domingo, Orcutt (UC);
San Jorge, Gentry 4176 (DS); 51 mi. NW. of El Crucero, Porter 488 (CAS, DS,
GH); 10 km. N. of Medano Amarillo, Bahia de la Magdalena, Johansen 644
(CAS, DS); S. of El Refugio, Wiggins 5526 (DS); 14 mi. 5S. of El Refugio, Shreve
7179 (ARIZ, DS, F); Estero Salinas, Porter 462 (CAS, DS, GH, UC), Wiggins
11477 (DS, GH UC, US); Bahia de la Magdalena, Lung in 1894 (UC).

Camissonia sceptrostigma is uniform morphologically and, while an
undoubted member of sect. Hwlobus, is entirely distinct within the
group. It has not been observed growing sympatrically with any of
the other three species of the section, but occurs in the same vicinity
as each one of them in the region of the northern Desierto Vizcaino.
Four plants of the Verity collection cited above were grown in the
experimental garden and found to be self-incompatible.

4. Camissonia californica (Nutt. ex Torr.& Gray) Raven, Brittonia 16: 283.

1964.

Eulobus californicus Nutt. ex Torr. & Gray, Fl. N. Amer. 1:515. 1840. Not
Oenothera californica 3. Wats. 1876.

Oenothera leptocarpa Greene, Pittonia 1: 302. 1889.

Oenothera eulobus H. Lév., Monogr. Onoth. 231. 1905.

Oenothera crassifolia var. leptocarpa (Greene) H. Lév., Monogr. Onoth. 209
1905.

Virgate annual 2-180 cm. tall, with a well-defined basal rosett©
which is usually withered by the time of flowering; no flowers formed
in the lower nodes; stems thick, fleshy, hollow, bluish green, sometimes
glaucescent, or bright green. Plants subglabrous, the leaves occasion-
ally finely strigulose or with a few glandular trichomes on the margins
and midvein. Leaves irregularly and sharply pinnatifid-lobed, very
much reduced upward and in the inflorescence, mostly in the basal

198 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

rosette, narrowly elliptic, up to 30 cm. long and 6.5 cm. wide, the apex
acute, the base narrowly cuneate. Inflorescence with a few glandular
trichomes, very rarely strigulose. Hypanthium 0.6-1.5 mm. long,
1-1.8 mm. across at the summit, glabrous within, closed by a conspic-
uous rounded fleshy disc. Sepals 3.9-8 mm. long, 0.7-1.5 mm. wide.
Petals 6-14 mm. long, 3.5-8.5 mm. wide, with or without fine red
flecking at the base. Filaments of the episepalous stamens 3-9 mm.
long, those of the epipetalous ones 2-5 mm. long; anthers of longer
stamens 1-2.5 mm. long, those of the shorter ones greatly reduced
0.5-1.2 mm. long. Style 4-10 mm. long, glabrous; stigma 0.8-2 mm.
thick, held at the same level as the anthers of the longer stamens,
which more or less surround it at anthesis. Capsules straight or slightly
curved, (4.5—) 6-11 cm. long, 1-1.2 mm. in diameter, subterete when
fresh, becoming quadrangular in transection when dry, sharply re-
flexed at maturity. Seeds 1.3-1.6 mm. long, 0.5-0.7 mm. thick, olive,
often flecked with purple dots. Gametic chromosome numbers, n=7,
14. Mostly self-pollinating, and apparently always self-compatible;
rarely visited by insects.

Tyee: Near San Diego, San Diego County, California, March-May
1836, 7’. Nuttall (NY; isotypes, BM, GH, K, PH).

Distrisution (Figure 5): Open or brushy hillsides and washes, or
desert flats, often in loose soil: upper Austin Creek, Sonoma County,
California (Kruckeberg 5933, DS); from the southern half of San Benito
County and westernmost Fresno County, California, south through
the Coast Ranges and mountains of southern California; the low foot-
hills for about 20 miles east of Bakersfield, Kern County, California;
and across the deserts (common along their western margins; absent
in the Colorado Desert east of the Salton Sink; rare in the Providence
and Old Woman Mountains of the eastern Mojave Desert) to western
and southern Arizona (vicinity of Kingman, Mohave County; south-
ernmost Yavapai, Gila, and Graham Counties southward); whence
south to central Sonora (to ca. 7 miles west of Mina San José on road
to Misa, Wiggins 6317, DS, POM, US; north of Bocoachic, 3,730 ft.,
Wiggins 11704, DS) at about 28°20’ N. lat. and throughout Baja
California south to the Sierra Vizeaffio at about 27°15’ N. lat. (e.g.,
Gentry 7416, DS, POM; Gentry 7515, DS); also on Santa Cruz and
Santa Catalina Islands, California. From sea level to 4,000 ft. alt.

Vouchers for chromosome number (39 individuals, 24 populations),
n=7:

U.S.: CaLIroRNIA: LOS ANGELES co.: Santa Monica Mts.: Decker Canyon,
Mathias 3159 (COLO, LA, RSA, UC); Latigo Canyon, Venkatesh 6 (3 plants;
LA), 7 (2 plants; ARIZ, LA, RSA, UC); South Sepulveda Way, Lewis 1629 (LA);
Mulholland Drive, 6.3 mi. from intersection with Beverly Glen Blvd., Venkatesh 4

(LA), 2.5 mi. from intersection, Venkatesh 2 (11 plants; LA). ORANGE co.: 2.3 mi.
above gate on Silverado Canyon Truck Trail, R17749. RIVERSIDE co.: 3.5 mi. W.

RAVEN—GENUS CAMISSONIA 199

s Dy CD09 (e)
BB O2O fe) ot
\ ) OSS 0 .e) &
ae s re) {e)
ag mee @) Qo
DO-* 1?)
© ° 00
()
e)
ie) oO oo}
C) ie) Ni O65 0 (@)
) % CO
D e) ie)
,@ [@)
OO [@)
© ° 10)
L)
00 oO
& 3
“.
Q
O.\y °
4 e
#)\ Om fe)
b bO
() o
O i)
=

\ n

Ficure 5.—Portion of southwestern United States and adjacent Mexico, showing range of
Camissonia californica (O): A=diploid chromosome counts (n=7) and @=tetraploid
(n=14).

of Cabazon, Lewis 1679 (LA); 2 mi. N. of Aguanga, Klein 1299 (RSA). SAN
BERNARDINO co.: N. of Muscovy, Gregory 362 (RSA). SAN DIEGO co.: 2.3 mi. 5.
of Rincon Springs, R17399; Lake Wohlford, R20170; 2 mi. E. of Escondido,
700 ft., R17405; 2 mi. E. of San Pasqual, R17410; Rincon Springs, R20172; 1.8
mi. from Ballena on road to Ramona, R9581 (LA); 0.7 mi. W. of Banner, 3,000
ft., R17421; 0.9 mi. W. of Barrett Lake Road on State Hwy. 94, Breedlove 1852.
SAN LUIS OBISPO cO.: Parkhill Road, Calf Canyon, La Panza Campground, ca.
2,000 ft., R18325; La Panza Campground, 2,100 ft., R18324; 4.5 mi. NE. of
Santa Margarita, R20157. veNruRA co.: 11.4 mi. N. of Fillmore on Cow Springs
Road, Topatopa Mts., 2,500 ft., Breedlove 2653; 9 mi. 8. of Oxnard, State Hwy.
101A, Copp in 1962.
BAJA CALIFORNIA: Ensenada, Bates 2340.

200 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Vouchers for chromosome number (53 individuals, 38 populations),
n= 14:

U.S.: ARIZONA: MARICOPA co,: 12 mi. SW. of Roosevelt, Gregory 61 (RSA).
PIMA CO.: Santa Catalina Mts.: 2.8 mi. below Molina Basin Pienic Area, Lewis
1075 (LA); Molina Basin Picnic Area, Lewis 1077 (LA, RSA). YAVAPAI CO.: 2 mi.
E. of Congress Junction, Munz et al. 22972 (RSA). CaLirorNIA: LOS ANGELES CO.:
Soledad Canyon, Lewis & Thompson 1615 (LA); Mint Canyon at road to Acton,
R13906. sanra MONICA MTs.: Latigo Canyon, R20162; 2.5 mi. W. of Seminole
Hot Springs, R20163; 4.1 mi. NW. of Tapia Park, R14046; Topanga Canyon,
Venkatesh 5 (8 plants; LA); Mulholland Drive 1.3 mi. W. of Laurel Canyon Blvd.,
R14081 (COLO, LA, RSA); Mulholland Drive 6.3 mi. from junction with Beverly
Glen Blvd., Venkatesh 3 (6 plants; LA). RIVERSIDE co.: Aqueduct Road 5.9 mi.
S. of Garnet, Lewis 1620 (LA); Pinyon Crest. Road 2.5 mi. from Palms-to-Pines
Hwy., Theobald 61; Pushawalla Canyon, H. & M. Lewis 1610 (LA). SAN BENITO
co.: Call Mts., E. of a point 18 mi. N. of New Idria, R10875 (LA, RSA). san
BERNARDINO CO.: 1.5 mi. W. of Mill Creek Ranger Station, R17468; Morongo
Valley, Kyhos 65-125. san piEGo co.: 0.4 mi. from U.S. Hwy. 395 on road to
Fallbrook, Wedberg 205 (herb. San Diego State College); Earthquake Valley,
R11537; Borrego Valley, H. & M. Lewis 1544 (LA); 0.7. mi. W. of summit of
Yaqui Pass, R11520. saNTA BARBARA CO.: Head of Buckhorn Canyon, summit of
San Rafael Mts., on Tepusquet Canyon Road, Breedlove 2780; Santa Barbara,
Breedlove 2227; 1 mi. 8. of Prisoners Harbor, Santa Cruz I., Breedlove 2816.
VENTURA Co.: 0.5 mi. N. of turn to Matilija Hot Springs, 1,400 ft., R17495; 0.3
mi. SW. of Camp Comfort, 8. of Ojai, 600 ft., R17494; Little Syeamore Canyon,
Santa Monica Mts., Lewis 1646 (LA).

BAJA CALIFORNIA: 7.5 mi. N. of Santa Tomas, R17049; 5.5 mi. 8. of San
Vicente, R17023; 5 km. N. of Colonia Guerrero, R12152 (LA, UC); 2 km. N. of
Colonia Guerrero, R12142 (LA, UC); 14 mi. S. of San Quintin, R17031; 7.5 mi.
S. of Socorro, Klein 1222 (RSA); 2 mi. inland from Ensenada de San Francisquito,
Moran 12392; Bahia de los Angeles, Moran 12308; Las Trincheras, Moran 12608;
Cuesta la Ley, Barril road, Moran 12642.

Camissonia californica, with its virgate habit, nearly leafless
branches, and Jong, sharply reflexed capsules, superficially resembles
some species of Cruciferae more than members of the family Ona-
graceae. Its capsules have been said to be imperfectly 4-celled, or 2-
celled as in the related genus Gayophytum (Munz, Amer. Journ.
Bot. 16: 253-254. 1929; N. Amer FI. II. 5: 80. 1965), but I have
been unable to confirm this in the wide range of material examined.
This species, while very distinct, is obviously closely related to the
other three grouped with it by Munz (loc. cit.) and recognized in the
present revision as comprising Camissonia sect. Eulobus. It has been
observed growing sympatrically with C. crassifolia and C. angelorum,
but no hybrids have been found.

The only tetraploids in sect. Hulobus are found in the present species.
T have found no character or combination of characters by which
they can be separated from the diploids, and their pollen appears to be
identical in size and number of pores. As pointed out by Munz (N.
Amer. FI. IT. 5: 146. 1965), coastal populations of this species tend to
consist of relatively robust, large-flowered plants; yet it is by no

RAVEN—GENUS CAMISSONIA 201

means true that all such populations are diploid. On the basis of the
92 individuals and 62 populations reported here (including earlier
reports by Lewis et al., Aliso 4: 73-86. 1958), the diploids are much
more narrowly distributed than the tetraploids (fig. 5). Diploids are
known from coastal San Luis Obispo County, and occur sporadically
near the coast in Ventura and Los Angeles Counties. From the Los
Angeles basin to San Diego County, they range much farther inland,
occurring to the margin of the desert near San Gorgonio Pass and near
Aguanga; but they are intermixed with tetraploid populations in
some pattern not yet properly understood, occurring as far west as
the vicinity of Fallbrook in northwestern San Diego County. Un-
fortunately, no counts are available for extreme western Riverside
County or for Orange County. In San Diego County, aside from the
Fallbrook record just mentioned, most of the populations appear to
be diploid, the tetraploid being confined to the margins of the desert.
In Baja California, on the other hand, only a single diploid count is
known, that from Ensenada, and all of the interior populations and
those near the coast farther south (which consist of robust, large-
flowered individuals similar to the coastal ones found farther north)
are tetraploid. There does not appear to be any point in attempting
to separate the diploid and tetraploid populations of this species
taxonomically, and indeed there does not seem to be any practical
possibility of doing it even if it were thought to be highly desirable.

No multivalent configurations have been observed in the tetraploid,
but the lack of chromosome associations larger than bivalents is
undoubtedly genetically controlled, as it is in sect. Holostigma and
sect. Camissonia, and certainly does not indicate a complete lack of
homology between the two sets of chromosomes present in these
tetraploid populations. Indeed, an autotetraploid origin appears most
likely for these tetraploids and would be consistent with their mor-
phological relationships. If this is true, however, it is somewhat diffi-
cult to explain the occurrence of the tetraploids over a much wider
range than the diploids, unless one could postulate that they recom-
bine the genetic material present in diploid populations in some novel
way (or ways) highly adaptive in more interior sites. This favorable
genetic combination could be maintained by the tetraploidy of the
interior plants, which are probably also more highly self-pollinating
than those found nearer the coast, judging from their smaller flowers.
Two diploid populations (R20157, cited above, from San Luis Obispo
County, California, 7 plants; and R20166, San Pasqual Grade, San
Diego County, California, 8 plants) and three tetraploid populations
(Morongo Valley, San Bernardino County, California, Ayhos 65-125,
2 plants; Santa Monica Mountains, Los Angeles County, California,
R20162, 5 plants; R20163, 6 plants the last two cited above)

202 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

were tested by pollen-tube growth and found to be self-compatible.
Kruckeberg 5933, cited above, was grown in a greenhouse at the
University of Washington and found also to be self-compatible (A. R.
Kruckeberg, pers. comm.). This collection was undoubtedly also
tetraploid. In the smaller flowered interior populations, the anthers
deposit pollen directly on the stigma and there is no doubt that
autogamy predominates.

The disjunct occurrence of this species on serpentine near Austin
Creek in western Sonoma County, California, is of considerable
interest. I have no record of its occurrence on serpentine elsewhere,
but it is characteristic for species occurring in highly marginal situ-
ations to occur in edaphic conditions that are unusual or unknown in
their main area of distribution (Raven, Evolution 18: 336-338, 1964).

Section II. Chylismia

Camissonia sect. Chylismia (Nutt. ex Torr. & Gray) Raven, Brittonia 16: 281
1964.
Oenothera subg. Chylismia Nutt. ex Torr. & Gray, Fl. N. Amer. 1: 506. 1840.
Chylismia (Nutt. ex Torr. & Gray) Raimann, in Engl. & Prantl, Natiirl.
Pflanzenfam. III. 7: 217. 1893.
Oenothera sect. Euchylismia (Nutt. ex Torr. & Gray) Munz, Amer. Journ.
Bot. 15: 224. 1928.
Oenothera sect. Chylismia Raven, Univ. Calif. Publ. Bot. 34: 80. 1962.
Oenothera sect. Tetranthera Raven, Univ. Calif. Publ. Bot. 34: 114. 1962.
Type species: Oenothera exilis Raven=Camissonia exilis (Raven) Raven.
Camissonia sect. Tetranthera (Raven) Raven, Brittonia 16: 282. 1964.
Perennial or annual herbs, the plants caulescent, not flowering
at the basal nodes, with or without a well-developed basal rosette.
Leaves pinnately divided or entire. Inflorescence erect or drooping
in bud. Flowers opening near sunrise or in the late afternoon (in
C. claviformis only). Ovary lacking a sterile projection. Hypanthium
lacking a fleshy disc within. Sepals reflexed separately. Petals yellow,
white, or lavender, if yellow strongly ultraviolet reflective, with or
without one or more red dots near the base, and some yellow-flowered
species? with the basal area of the petals with a large, nonultraviolet
reflective area; petals, if white or yellow, often fading orange red
after pollination. Stamens 8 or 4; anthers versatile, long-ciliate or
glabrous. Pollen shed singly. Capsule long-pedicellate, regularly and
promptly loculicidal, each valve with a brown, prominent midrib,
terete, the seeds in 2 rows in each locule. Seeds monomorphic, lentic-
war, with a more or less pronounced membranous margin when
immature, lance-ovoid when mature, finely lacunose.
TYPE SPECIES: Camissonia scapoidea (Torr. & Gray) Raven.

* Camissonia brevipes, C. confertiflora, C. eastwoodiae, and C. multijuga. Other
species did not show a contrasting pattern of this sort.

RAVEN—GENUS CAMISSONIA 203

Disrripution: Sandy desert slopes, flats, and washes, often with
Artemisia tridentata in the northern part of its area, or on rock slides
or cliffs, from southeastern Oregon, east-central and southern Idaho,
and central and western Wyoming, south throughout the Great
Basin in eastern California, Nevada, and western Colorado, to north-
westernmost New Mexico, throughout Arizona, and widespread in
the Mojave and Colorado Deserts, south to northwesternmost Sonora
above the head of the Golfo de California, and in eastern Baja Cali-
fornia south to the vicinity of Bahia de los Angeles at about 29° N. lat.
Also, Camissonia scapoidea has been collected twice on the upper
drainage of the Arkansas River in eastern Colorado and is the only
species of its section and one of the two in the genus—the other is
(’. chamaenerioides—to occur east of the crest of the southern Rocky
Mountains, Plants of Camissonia sect. Chylismia do not occur west
of the Cascade-Sierra Nevada axis and constitute the only major
group of the genus strictly confined to the deserts; however, the
distribution of sect. Hremothera is generally similar.

The phylogeny and relationships of this group were discussed
by me in detail in 1962 (Univ. Calif. Publ. Bot. 34: 1-122), and
the section is included here only to bring it into line with my treat-
ment of the rest of the genus and to bring my account of it up to
date. The only specimens cited in the present treatment are types
and recently collected vouchers for chromosome number determina-
tions and range extensions. Distribution maps were presented in
my 1962 revision and are not reprinted here.

Only one major change is made in the present treatment. Impressed
with its loss of the epipetalous stamens, I set up a new monotypic
section, sec. Tetranthera, for Camissonia exilis in my 1962 revision.
Subsequently, however, I have found that some individuals of C.
andina (sect. Nematocaulis) likewise have lost the epipetalous stamens,
and some individuals of C. minor (sect. Eremothera) approach this
condition. In all of these cases, the loss of this whorl of stamens
appears to be associated with autogamy. In the light of this informa-
tion, and considering the close overall resemblance between C. exilis
and ('. walkeri, C. scapoidea, and C. parryi, I now feel that my earlier
segregation of C. erilis as a monotypic section reflected an unwarranted
overemphasis of a single characteristic. The relationships of this
species appear to be reflected more accurately when it is included in
sect. Chylismia with its close relatives.

5. Camissonia confertiflora (Raven) Raven, Brittonia 16: 281. 1964.
Oenothera confertiflora Raven, Univ. Calif. Publ. Bot. 34: 80. 1962.
Stout annual herb 15-50 em. tall, branched at the base and above,
with a well-developed basal rosette; stems densely villous at the base,
mixed with shorter strigose pubescence, and villous above with short

204 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

glandular pubescence. Basal leaves oblanceolate, 7-20 cm. long,
pinnate, the lateral leaflets to 2 em. long, alternating with shorter
pinnules, or reduced; terminal leaflet narrowly ovate, to 5 cm. long
and 2.5 cm. wide, sparsely villous. Inflorescence dense and corymbiform
in bud, later elongating, glandular-puberulent. Hypanthium 3-5 mm.
long, 3-5 mm. across at the summit, short-villous within below. Sepals
9-12 mm. long, 2.5-4 mm. wide, glandular-pubescent, or with a few
scattered long trichomes; free caudate portions conspicuous, arising
just below the apices, these with a median row of light brown oil
cells. Petals 12-18 mm. long, 12-17 mm. wide, bright yellow, of same
color as stamens, style, and inside of hypanthium, red dotted at the
base, fading purplish. Filaments of the episepalous stamens 6-8 mm.
long, those of the epipetalous ones 4—5 mm. long; anthers 4-6 mm. long,
ciliate. Style 11-18 mm. long; stigma ca. 1.5 mm. thick, held well
above the anthers at anthesis. Immature capsule to 35 mm. long, on a
pedicel 5-15 mm. long, glandular-pubescent. Seeds not known.
Presumably self-incompatible.

Type: In cinder soil, on east side, base of Vulcan’s Throne, Toro-
weap Valley, ca. 4,200 ft., Grand Canyon National Monument,
Mohave County, Arizona, 30 April 1952, FE. McClintock 52-294
(CAS 373481; isotypes, ARIZ, NY, WS).

DistriputTion: Known only from the type locality.

Camissonia confertiflora is most closely related to C. brevipes and to
C. multijyuga. It is easily distinguished from the latter by its larger
flowers, drooping inflorescence, and large buds; and from the former
by its glandular sepals, unequal stamens, and uniformly branched
habit. In addition, the single locality for C. confertiflora is outside the
range of C. brevipes. Camissonia multijuga grows within less than a
mile of this locality, but in this area is very distinct; some collections,
however, such as one from Searchlight Ferry, Mohave County, Arizona
(Shreve 7870), are more similar to C. confertiflora in their crowded
inflorescences and larger flowers. No more is known about this species
at present than when I described it in 1962.

6. Camissonia brevipes (A. Gray) Raven, Brittonia 16: 281. 1964.

Annual, 3-75 cm. tall, the stems often densely clothed below with
spreading white trichomes. Leaves mostly in a basal rosette, spatulate,
the terminal leaflet to 6.5 cm. long and 5 cm. wide, sometimes cordate
at the base, the lateral leaflets reduced or absent. Inflorescence
nodding, elongating in flower and fruit. Hypanthium 3-8 mm. long,
densely short-villous in lower portions within, 1.5-7 mm. across at the
summit. Sepals 5-9 mm. long, the apices covered with brown oil cells,
the caudate appendages present or absent. Petals 3-18 mm. long,
4-25 mm. wide, bright yellow, rarely red dotted, the same color as the
stamens and style. Stamens subequal; filaments 3-6 mm. long;

RAVEN—GENUS CAMISSONIA 205

anthers 2.5-6 mm. long, ciliate. Style 10-18 mm. long; stigma 1-2 mm.
thick held above the anthers at anthesis. Capsule 18-92 mm. long,
glabrous or sparsely villous, ascending or spreading, straight or curved;
pedicel 2-20 mm. long. Seeds 1-1.5 mm. long, 0.6-1 mm. wide.
Gametic chromosome number, n=7. Self-incompatible.

Distripution: Alluvial slopes, flats, and canyon sides, Washington
County, Utah, Lincoln and Esmeralda Counties, Nevada, and Inyo
County, California, south to Yuma County, Arizona, and Imperial
County, California. From ca. 230 ft. below sea level to 6,000 ft. alt.
6a. Camissonia brevipes subsp. brevipes FRONTISPIECE

Oenothera brevipes A. Gray, Pac. R. R. Rept. 4: 87. 1857.

Oenothera divaricata Greene, Bull. Torrey Cl. 10: 125. 1883. Type: CAS
866, labeled: ‘‘The specimen found among the ‘Cruciferae’ of State Geo.
Survey”; isotype, GH. As indicated in a note penciled by Katherine
Brandegee, probably the same as a specimen from gravelly banks, Fort
Mojave, Mojave County, Arizona, 15 February 1861, J. G. Cooper (GH),
bearing the characteristic printed label, ‘Geological Survey of California
1860-61.”

Chylisma brevipes (A. Gray) Small, Bull. Torrey Cl. 23: 194. 1896.

Oenothera brevipes var. typica Munz, Amer. Journ. Bot. 15: 228. 1928.

Oenothera brevipes subsp. brevipes; Raven, Univ. Calif. Publ. Bot. 34: 81. 1962.

Stout, generally unbranched above, the stems villous. Terminal
leaflet ovate. Inflorescence dense, nodding. Hypanthium 4-8 mm.
long, villous without, yellow within. Sepals usually villous, with
a caudate protection arising below the apex. Petals 6-18 mm. long,
7-25 mm. wide. Anthers 4-6 mm. long. Capsule 20-92 mm. long;
pedicel 5-20 mm. long. Gametic chromosome number, n=7. Self-
incompatible.

Type: On gravelly hills, Colorado River, California or Arizona,
February 1854, J. M. Bigelow (GH; isotypes, NY, PH, US).

DistrisputTion: Hillsides and alluvial fans, sometimes in great
abundance on sandy slopes, southern Esmeralda County, Nevada,
and southwestern Washington County, Utah, south to northern
Yuma County, Arizona, and northern Imperial County, California.
From ca. 230 ft. below sea level to 6,000 ft. alt.

Camissonia brevipes subsp. brevipes grows in more mesic habitats
than those occupied by subsp. arizonica or subsp. pallidula. In the
mountains east of Death Valley, for example, C. claviformis subsp.
aurantiaca and C. brevipes subsp. pallidula are found growing on dry
flats, C. munzii above in more mesic sites, and C. brevipes subsp.
brevipes in the best watered situations near the tops of the passes.
It grows sympatrically with C. claviformis subsp. aurantiaca, C.
claviformis subsp. claviformis, C. claviformis subsp. funerea, C. clavi-
formis subsp. integrior, C. claviformis subsp. peeblesii, C. multijguga, C.
munzi, and C. walkeri subsp. tortilis, and forms hybrids with C. clavi-
formis, C. multijuga, and C. munzit. It intergrades with subsp. pal-

206 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

iidula over a wide area in Utah, Nevada, California, and Arizona,
but not with subsp. arizonica, with which it does not come in con-
tact. The most densely villous and succulent individuals within subsp.
brevipes are found in Mohave County, Arizona, and Clark County,
Nevada, in the region about Hoover Dam. Progeny of Everett &
Balls 23034, grown at Claremont, California, were self-incompatible,
as were 12 plants from Frenchman Flat, Nye County, Nevada,
R18853; 3 from State Hwy. 95, 33.4 miles north of junction with
U.S. Hwy. 60-70, San Bernardino County, California, Kyhos 65-237;
and 3 from 2.9 miles south of Twentynine Palms, San Bernardino
County, California, Ayhos 64-31. Individuals from the last three
populations were tested by pollen-tube growth in selfed and out-
crossed individuals. It is surprising that insect visitors to this self-
incompatible species appear to be relatively uncommon; neverthe-
less, full seed set appears to be the rule, except in isolated individuals.
Probably a wide spectrum of visitors, active throughout the hours
of daylight, is effective in cross-pollinating the plants.
6b. Camissonia brevipes subsp. pallidula (Munz) Raven, Brittonia 16: 281. 1964.

Oenothera brevipes var. pallidula Munz, Amer. Journ. Bot. 15: 229. 1928.

Oenothera pallidula (Munz) Munz, Leafl. West. Bot. 2: 88. 1938.

Oenothera brevipes subsp. pallidula (Munz) Raven, Univ. Calif. Publ. Bot. 34:

83. 1962.

Slender, usually branched above, the stems generally strigose.
Terminal leaflet narrowly ovate to ovate. Inflorescence dense, nodding.
Hypanthium 4-5 mm. long, yellow within. Sepals strigose, lacking
projections, or these poorly developed. Petals 7-12 mm. long, 8-14
mm. wide, often red-dotted near the base. Anthers 4-6 mm. long.
Capsule 2-4.2 em. long; pedicel 2-10 mm. long. Gametic chromo-
some number, n=7. Self-incompatible.

Type: Las Vegas, Clark County, Nevada, 25 April 1905, M. E.
Jones (POM 38034).

DistrisuTion: Dry flats, often on desert pavement, often with
Larrea and Ambrosia dumosa, from Washington County, Utah, to
southeastern Inyo County, California, and south to Riverside Coun-
ty, California, and Mohave County, Arizona, but only along the
Colorado River south of Clark County, Nevada. From 250-3,500
ft. alt.

Fourteen plants of this entity from Frenchman Flat, Clark County,
Nevada (R18905), were tested for self-compatibility by pollen-tube
growth; all were found to be self-incompatible. Camissonia brevipes
subsp. pallidula intergrades with C. brevipes subsp. brevipes over a
wide area, and no evidence for reduction of fertility or for cytological
abnormality has been found in the intermediate plants, in spite of the
very different phenotypes of these two subspecies (Raven, Univ.
Calif. Publ. Bot. 34: 84. 1962). Camissonia brevipes subsp. pallidula

RAVEN—GENUS CAMISSONIA 207

srows sympatrically with C. claviformis subsp. aurantiaca, C. clavi-
formis subsp. peeblesii, C. munzii, C. walkert subsp. tortilis, and
C. multijuga, forming natural hybrids with the first two.
6c. Camissonia brevipes subsp. arizonica (Raven) Raven, Brittonia 16: 281.
1964.
Oenothera brevipes subsp. arizonica Raven, Univ. Calif. Publ. Bot. 34: 84.
1962.

Slender, often branched above, the stems villous. Terminal leaflet
narrowly ovate. Inflorescence nodding, the buds individually deflexed
before opening. Hypanthium 3-5 mm. long, orange brown within.
Sepals villous to strigose, with free caudate portions nearly terminal
or lacking. Petals 3-8 mm. long, 4-7 mm. wide, often red-dotted at
base, and often fading reddish. Anthers 2.5-5 mm. long. Capsule
1.8-6 cm. long; pedicel 2.5-5 mm. long. Gametic chromosome number,
n=7.

Type: Mohawk Mountains, Yuma County, Arizona, 29 March
1935, A. & R.A. Nelson 1343 (RM 172249; isotypes, GH, MO, NY,
UC, US).

Disrrisution: Rocky slopes and flats, Yuma County, Arizona,
and eastern Imperial County, California. From 250-900 ft. alt.

Plants of Camissonia brevipes subsp. arizonica from western Arizona
are similar to C. claviformis subsp. rubescens, and the two taxa occur
sympatrically in that area. Hybrids between them, however, seem to
be rare. The present taxon also grows with C. claviformis subsp.
peeblesti and with C. claviformis subsp. yumae, forming occasional
natural hybrids with each of them.

7. Camissonia multijuga (S. Wats.) Raven, Brittonia 16: 281. 1964.

Oenothera multijuga S. Wats., Amer. Nat. 7: 300. 1873.

Oenothera brevipes var. parviflora 8. Wats., Amer. Nat. 9: 271. 1875. Type:
Valley of the Virgin R., near St. George, Washington Co., Utah, 1874,
C. C. Parry 74 (GH; isotypes, MO, F); not Oenothera parviflora L. 1758.

Chylisma multijuga (S. Wats.) Small, Bull. Torrey Cl. 23: 193. 1896.

Chylisma venosa A. Nels. & Kennedy, Muhlenbergia 3: 140. 1908.

Chylisma hirta A. Nels., Bot. Gaz. 47: 428. 1909. Type: in stony washes,
Tuly’s Ranch, 13 mi. from Las Vegas, Clark Co., Nevada, 1 May 1905,
L. N. Goodding 2348 (RM 65187; isotypes, UC, GH, MO).

Chylisma parviflora (8S. Wats.) Rydberg, Fl. Rocky Mts. 1064. 1917.

Oenothera brevipes var. multijuga (S. Wats.) Jeps., Man. Fl. Pl. Calif. 687.
1925.

Oenothera multijuga var. typica Munz, Amer. Journ. Bot. 15: 230. 1928.

Oenothera multijuga var. parviflora (S. Wats.) Munz, Amer. Journ. Bot. 15:
231. 1928.

Oenothera watsoni Tidestr., Proc. Biol. Soc. Wash. 48: 41. 1935.

Oenothera phlebophylla Tidestr. Proc. Biol. Soc. Wash. 48: 41. 1935.

Virgate annual or biennial 20-150 cm. tall, with a well-developed
basal rosette, nearly leafless above, with numerous divergent branches;

295-655. O—68——4

208 OONTRIBUTIONS FROM THE NATIONAL HERBARIUM

stems villous, at least below, glabrous or even glaucous above, rarely
glandular-pubescent. Basal leaves oblanceolate, villous to nearly gla-
brous, sometimes purple-dotted, 6-30 cm. long, pinnate or bipinnate,
with well-developed narrowly ovate, irregularly serrate lateral leaflets
to 2.5 cm. long and 2 em. wide, alternating with smaller pinnules,
the leaflets often divided further; terminal leaflet irregularly serrate,
to 6 cm. long and 4 cm. wide, the veins below prominently lined
with dark brown oil cells. Inflorescence erect to drooping, sparse,
elongating in bud, the buds individually pendulous when mature.
Hypanthium 1-3 mm. long, 1-1.5 mm. wide at the summit, glandular-
puberulent, villous, or glabrous without, glabrous or villous in lower
portions within. Sepals 3-8 mm. long, 1.5-2 mm. wide, glandular-
puberulent, strigose, villous, or glabrous, short-caudate at or slightly
below the apices. Petals spreading in anthesis, 4-9 mm. long, 4-8 mm.
wide, bright yellow, rarely cream, sometimes fading purplish, and
of the same color as the stamens, style, and usually the inside of
the hypanthium, which may rarely be greenish. Filaments of the
episepalous stamens 2.5-4 mm. long, those of the epipetalous ones
1.3-3 mm. long. Anthers 2-4 mm. long, ciliate. Style 7-11 mm. long;
stigma 0.7-1 mm. thick, held well above the anthers at maturity.
Capsule 1—5.2 cm. long, 1.3-2 mm. thick, slender, straight to slightly
curved, glabrous or sparsely villous; pedicel 7-20 mm. long, spreading
or rarely slightly deflexed. Seeds 1-1.3 mm. long, 0.8-1 mm. thick.
Gametic chromosome number, n=7. Self-incompatible.

Type: Near Kanab, Washington County, Utah, 1872, Mrs. Ellen
P. Thompson (GH).

DistriBution: Forming colonies on rocky slopes and banks of eroded
sedimentaries, sometimes on gypsum or limestone, often on con-
glomerates, often with Pinus edulis and Juniperus or with Larrea
and Encelia farinosa, Washington County, Utah, and southern Lin-
coln County, Nevada, to northern Mohave County, Arizona. From
1,000-3,500 ft. alt.

Although occupying a relatively limited range, Camissonia multi-
juga is one of the most variable species in its section. Nevertheless,
it has not been possible to identify well-marked infraspecific taxa.
Plants of Gullion 259, 8 miles southwest of Davis Dam, Clark County,
Nevada, grown in the experimental garden, were self-incompatible,
as were 12 individuals from a population in Eldorado Canyon, Clark
County, Nevada, R18970, judging from the comparative growth of
pollen tubes in selfed and outcrossed individuals. The autogamous
C. walkeri subsp. tortilis was very probably derived from this entity,
and grows sympatrically with it at several localities in southern
Nevada, with no direct evidence for hybridization. Camissonia multi-

RAVEN—GENUS CAMISSONIA 209

juga also grows sympatrically with C. brevipes subsp. brevipes, C.
brevipes subsp. pallidula, C. claviformis subsp. aurantiaca, and C.
parryi, rarely forming hybrids with both subspecies of C. brevipes.
8. Camissonia speculicola (Raven) Raven, Brittonia 16: 282. 1964.

Tufted perennial 10-50 cm. tall, with numerous divergent branches
from the base; stems and leaves glabrous or sparsely villous below.
Leaves broadly elliptical, 3-20 cm. long, pinnate or often bipinnate,
mostly in a basal rosette; lateral leaflets well developed, to 15 mm.
long and 7 mm. wide, often further divided and alternating with
smaller pinnules; oil cells beneath prominent, dark brown. Inflores-
cence erect, elongating in flower and fruit, the mature buds individually
drooping. Hypanthium 1.5-2 mm. long, 1 mm. across at the summit,
glandular-pubescent without, glabrous within. Sepals 2-5 mm. long,
0.5-0.8 mm. wide, glandular-pubescent, caudate with short append-
ages at the apices, these free in bud. Petals 2-6 mm. long, 1.5-4
mm. wide, bright yellow, fading light purplish, red-dotted near the
base, of the same color as inside of hypanthium, stamens, and style;
filaments of the episepalous stamens 1.5-3 mm. long, those of the
epipetalous ones 1-2 mm. long; anthers 1.2-2 mm, long, glabrous.
Style 4-7 mm. long; stigma ca. 0.8 mm. thick, at about the same level
as the anthers at anthesis. Capsule straight to slightly curved, spread-
ing to ascending; 0.8-2 cm. long, 1-1.5 mm. thick; glandular-pubes-
cent or glabrous; pedicel 6-10 mm. long; seeds 0.6-1 mm. long, to
0.6 mm. thick. Gametic chromosome number, n=7.

DisrripuTion: Washes and debris slides, along the Colorado River
in Coconino and Mohave Counties, Arizona, in several disjunct
localities. From 2,300—5,600 ft. alt.
8a. Camissonia speculicola subsp. speculicola

Oenothera speculicola subsp. speculicola Raven, Univ. Calif. Publ. Bot. 34:
87. 1962.

Densely tufted and somewhat woody at base, occasionally with
secondary flowering axes arising above from the decumbent stems.
Leaves glabrous, glutinous, shining. Style 4-5 mm. long, glabrous.
Gametic chromosome number, n=7.

Type: Debris slides and crevices in Redwall limestone, Kaibab
Trail at 5,400 ft. alt., south rim of Grand Canyon, Coconino County,
Arizona, 8 June 1958, P. H. Raven 13119 (RSA).

Distripution: Crevices of broken limestone, south rim of the
Grand Canyon, Coconino County, Arizona. From 4,500—5,600 ft. alt.

At its type locality, Camissonia speculicola subsp. speculicola occurs
in narrow crevices in the solid limestone, (. walkeri subsp. walkeri
nearby on loose rockslides; no evidence of hybridization between
these two entities was noted.

210 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

8b. Camissonia speculicola subsp. hesperia (Raven) Raven, Brittonia 16: 281.
1964.

Oenothera speculicola subsp. hesperia Raven, Univ. Calif. Publ. Bot. 16: 84.
1962.

Less densely tufted and not woody. Leaves sparsely pubescent.
Style 5-7 mm. long, pubescent at base.

Type: Near Mooney Falls, Havasu Canyon, Coconino County,
Arizona, 23 May 1950, J. 7. Howell 26486 (CAS 359774; isotype,
ARIZ).

Disrripution: Washes and dry stream beds, often on limestone,
in two disjunct areas along the Colorado River, Arizona: Havasu and
Hualapi Canyons, Coconino County; from Separation Canyon to
Spencer Canyon, Mohave County. From 2,300-3,500 ft. alt.

Camissonia speculicola subsp. hesperia resembles the allopatric
subsp. speculicola in its glabrous anthers and distinctive habit. Both
of these taxa appear to be outcrossers, but they are probably not
self-incompatible, judging from their relatively small flowers (com-
pared with those of the self-incompatible C. multijuga). Camissonia
speculicola subsp. hesperia grows sympatrically with C. walkeri subsp.
walkeri.

9. Camissonia walkeri (A. Nels.) Raven, Brittonia 16: 281. 1964.

Slender herb 10-60 em. tall, vilious below and with a basal! rosette,
the cauline leaves well developed or absent. Basal leaves oblong, with
brown oil cells prominently lining veins below, often purple-dotted,
doubly serrate, 3-22 em. long, villous, often densely so below; ter-
minal leaflet 1-5 em. long, 0.5-3 em. wide. Inflorescence branching,
erect, elongating in flower and fruit, the buds individually drooping.
Hypanthium 0.5-1.5 mm. long, glandular-pubescent or hispid without,
glabrous to sparsely villous within, Sepals 1.2-4 mm. long, 0.5-1 mm.
wide, often purple-dotted, short-caudate at the apices. Petals 1.2-6
mm, long, 0.6—6 mm. wide, bright yellow and of same color as stamens,
style, and inside of hypanthium. Filaments of the episepalous stamens
1-3 mm. long, those of the epipetalous ones 0.3-2 mm. long; anthers
0.6—-2 mm. long, glabrous or ciliate. Style 1.5-6 mm. long, glabrous or
villous at base; stigma ca. 0.6 mm, thick, surrounded by the anthers
at maturity. Capsule 1.2-4.5 em. long, 1.2-1.8 mm. thick, spreading or
ascending, the valves often twisted at maturity, straight or slightly
curved. Seeds 0.6-1.2 mm. long, 0.4-0.6 mm, thick. Gametic chromo-
some numbers, n=7, 14. Autogamous.

Distriputton: Washes and alluvial slopes, from Mesa County,
Colorado, to southern Elko and Mineral Counties, Nevada, and south
to Inyo and northeastern San Bernardino Counties, California;
Clark County, Nevada; and Coconino County, Arizona. From ca.
2,000-6,000 ft. alt.

RAVEN—GENUS CAMISSONIA 211

9a. Camissonia walkeri subsp. tortilis (Jeps.) Raven, Brittonia 16: 281. 1964.
Oenothera scapoidea var. tortilis Jeps., Man. Fl. Pl. Calif. 687. 1925.
Oenothera multijuga var. parviflora sensu Munz, Amer. Journ. Bot. 15: 2381.

1928, for the most part, excluding the type.
Oenothera walkeri subsp. tortilis (Jeps.) Raven, Univ. Calif. Publ. Bot. 34:
90. 1962.

Annual or short-lived perennial, nearly leafless above and with
well-developed basal rosette. Leaves pinnate or more rarely bipinnate,
the lateral leaflets usually well developed, to 3 em. long and 1.5 cm.
wide, often alternating with smaller pinnules. Hypanthium 1—1.5 mm.
long. Sepals 1.5-4 mm. long, often purple-dotted. Petals 2.8-6 mm.
long, 2-6 mm. wide. Anthers 1-2 mm. long, glabrous to sparsely
ciliate. Style 4-6 mm. long. Capsule on a pedicel 5-30 mm. long.
Gametic chromosome number, n=7. Autogamous.

Typr: Wild Rose Canyon, Panamint Mountains, Inyo County,
California, 24 May 1917, W. L. Jepson 7131 (JEPS 2669).

Distripution: Colonial in rocky debris near cliffs and along
ephemeral streams, often on limestone, Logan (?) and Millard Coun-
ties Utah, southern Elko and Mineral Counties, Nevada, and Inyo
and northeastern San Bernardino Counties, California, to Clark
County, Nevada, and Washington County, Utah. Common only in
Inyo County, California. From ca. 2,000-6,000 ft. alt.

Camissonia walkeri subsp. tortilis grows sympatrically with C.
brevipes subsp. brevipes, C. brevipes subsp. pallidula, C. heterochroma,
and C. multijuga without any evidence of hybridization. Although
modally very distinct from C. walkeri subsp. walkeri, subsp. tortilzs
replaces it geographically and their variation patterns overlap to such
an extent that I consider it desirable to retain both within the same
species.
9b. Camissonia walkeri subsp. walkeri

Chylisma walkeri A. Nels., Bot. Gaz. 56: 66, 1913.

Oenothera multijuga var. orientalis Munz, Amer. Journ. Bot. 15: 232. 1928.
Type: Moab, Grand Co., Utah, 7 June 1913, M. E. Jones (POM 38608).

Oenothera walkeri (A. Nels.) Raven, Univ. Calif. Publ. Bot. 34: 88. 1962.

Annual, usually leafy above, the leaves cordate to ovate and simple
by reduction of the lateral leaflets, or more rarely these developed ;
basal rosette not conspicuous. Hypanthium 0.5-1.3 mm. long. Sepals
1.5-2 mm. long. Petals 1.2-3 mm. long, 0.6-1.8 mm. wide. Anthers
0.6-0.8 mm. long, glabrous. Style 1.5-4 mm. long. Capsule on a pedicel
5-15 mm. long. Gametic chromosome numbers, n=7, 14. Autogamous.

Type: Dry gypsum hills, Paradox Valley, Montrose County,
Colorado, 1 July 1912, E. P. Walker 200 (RM 75864; isotypes, DS,
GH, POM, US).

Distrusution: Loose slides of limestone and other sedimentaries,
or in sandy washes, Mesa County, Colorado, and Emery County,

212 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Utah, south to Montezuma County, Colorado, and central Coconino
County, Arizona. From 3,200-5,500 ft. alt.

Under conditions of cultivation, plants of this subspecies develop a
rather full rosette of leaves similar to that characteristic of subsp.
tortilis; in the field, the cauline leaves are very often the only ones
present at the time of flowering. Plants of this taxon probably oc-
casionally hybridize with C. scapoidea subsp. scapoidea; one remark-
ab'e intermediate population is represented by Eastwood 5077 (CAS)
from Grand Junction, Mesa County, Colorado, with the pubescence
and cauline leaves of C. walkeri subsp. walkeri but the habit of C.
scapoidea subsp. scapoidea. Camissonia walkeri subsp. walkert grows
sympatrically with both subspecies of C. speculicola but in less stable
soil and without any evidence of hybridization. Camissonia scapoidea
subsp. scapoidea, on the other hand, is mostly restricted to alkaline clay
flats and is thus usually ecologically separated from ©. walkeri
subsp. walkeri.

An apparent autotetraploid population of C. walkeri subsp.
walkert has been found growing on the walls of the Marble Gorge,
Coconino County, Arizona. These plants had up to 4 rings of 4 chromo-
somes at meiotic metaphase I (Raven, Univ. Calif. Publ. Bot. 34: 40.
1962). Seven other individuals from four other populations were
diploid.

10. Camissonia claviformis (Torr. & Frém.) Raven, Brittonia 16: 282, 1964,

Annual 3-70 em. tall, branching mostly from the base. Leaves in
a prominent basal rosette, reduced upward, pinnately divided or
entire by reduction, 1.5-20 em. long, broadly oblanceolate, the terminal
segment 0.8-9 em. long, 0.2-4 cm. broad, lanceolate to cordate, the
lateral segments lacking or to 2.5 cm. long; veins beneath promi-
nently lined with brown oil cells. Inflorescence nodding, not elongating
much in flower. Flowers opening in late afternoon in all except subsp.
cruciformis. Hypanthium 1.5-6.5 mm. long, 1-5 mm. across at the
summit, short-villous in lower portions within. Sepals 2-8 mm. long,
1-2.5 mm. wide, with brown oil cells on midribs above and on free tips,
if present. Petals 1.5-8 mm. long, 1.5-10 mm. wide, yellow or white,
reflexed in anthesis, and of same color as stamens and style. Stamens
subequal, the filaments 1.5-5.5 mm. long, dilated at base; anthers 1.5—
6 mm. long, long-ciliate. Style 5-16 mm. long, held well above the
stamens at anthesis; stigma greenish, 0.6-1.3 mm. thick. Capsule
0.9-3.8 em.long, 1.5-2.3 mm. thick, straight or curved, clavate, on a
spreading or ascending pedicel 4-40 mm. long. Seeds pale brown,
narrowly obovoid, lenticular, 0.6-1.5 mm. long. Gametic chromosome
number, n=7. Self-incompatible.

DistripuTion: Sandy slopes and flats, often in washes, from
southeastern Oregon and adjacent Idaho south to eastern California,

RAVEN—GENUS CAMISSONIA 213

Nevada, and Arizona, northwestern Sonora, and to near latitude 29°
N. in eastern Baja California. From ca. 250 ft. below sea level to
6,500 ft. alt.

Camissonia claviformis is the most complex and one of the most
widely distributed species of the genus. The central part of its range
is occupied by five closely related white-flowered subspecies, clavi-
formis, aurantiaca, funerea, integrior, and peeblesit, so similar morpho-
logically that they could be regarded as a single taxon. South of this
area are found four additional subspecies, all yellow-flowered, namely,
peirsonii, rubescens, wigginsii, and yumae. These entities have sepals
and flower color similar to those of C. brevipes, and it is likely that
they were derived following hybridization between that species and
white-flowered populations of C. claviformis. North of the range of
the white-flowered subspecies are found two additional yellow-
flowered subspecies, cruciformis and lancifolia, the latter apparently
derived following hybridization between the former and some of the
white-flowered subspecies. Most populations of subsp. eruciformis con-
sist of plants in which the flowers open in the early morning; all other
subspecies consist of plants that have flowers opening in the late
afternoon. In this and other morphological respects, C. claviformis
subsp. cruciformis (including subsp. citrina) resembles C. eastwoodiae
and C. munzii and is undoubtedly close to the ancestral stock from
which C. claviformis was derived.
10a. Camissonia claviformis subsp. cruciformis (Kell.) Raven, Brittonia 16:

282. 1964.

Oenothera cruciformis Kell., Proc. Calif. Acad. Sci. 2: 227, fig. 71. 1873.

Chylisma clavaeformis var. cruciformis (Kell.) Small, Bull. Torrey Cl. 23:
193. 1896.

Chylisma cruciformis (Kell.) Howell, Fl. N. W. Amer. 233. 1898.

Oenothera clavaeformis var. cruciformis (Kell.) Munz, Amer. Journ. Bot.
15: 285. 1928.

Oenothera clavaeformis var. purpurascens sensu Munz, Leafl. West. Bot.
3: 538. 1941; not S. Wats. 1873:

Oenothera clavaeformis subsp. cruciformis (Kell.) Raven, Univ. Calif. Publ.
Bot. 34: 108. 1962.

Oenothera clavaeformis subsp. citrina Raven, Univ. Calif. Publ. Bot. 34: 108.
1962. Type: Roadbanks of decomposed conglomerate, in clay soil, 1.4
mi. NE. of Oregon State line on U.S. Hwy. 95, Jump Creek Canyon,
Owyhee Co., Idaho, 23 June 1958, P. H. Raven & O. T. Solbrig 13385A
(RSA).

Plants 3-55 cm. tall, strigose or glandular below and often glandular
above, or glabrous. Lateral leaflets few to numerous; terminal leaflet
to 8 cm. long and to 4 cm. wide, narrowly ovate to nearly cordate,
serrate with scattered teeth. Flowers usually opening near sunrise.
Hypanthium 2-6.5 mm. long, sparsely strigose or glandular without,
yellow or orange brown within. Sepals sparsely strigose or glandular-
pubescent. Petals 2.5-8 mm. long, 2.5-10 mm. wide, bright yellow,

214 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

often red-dotted at the base and often fading purple. Filaments 2-5
mm. long; anthers 2-6 mm. long. Style 7-16 mm. long. Capsule
0.8-2.8 cm. long, glandular-pubescent to glabrous, pedicel 4-30 mm.
long. Gametic chromosome number, n—7. Self-incompatible.

NerortypPE: Steamboat Springs, Washoe County, Nevada, 28 May
1959, P. H. Raven 14269 (RSA). The type specimen, if one was
prepared, has not been preserved; the proposed neotype is from the
type locality and a close match for Proc. Calif. Acad. Sci. 2, fig. 71
(p. 228). See Raven, Univ. Calif. Publ. Bot. 34: 108. 1962.

Distripution: Sandy or clayey flats and slopes, with Artemisia
tridentata, Purshia tridentata, Grayia spinosa, or Chrysothamnus,
Lake, Harney, and Malheur Counties, Oregon, western Canyon and
Owyhee Counties, Idaho, central and southern Washoe County,
Nevada, and Lassen County, California. From 2,000—4,600 ft. alt.

Camissonia claviformis subsp. eruciformis is unusually variable,
probably owing to hybridization between its southern populations
(which occupy a disjunct area) and subsp. integrior, which they
contact locally. The populations of this taxon found in the vicinity
of Sparks, Washoe County, Nevada, consist of plants that have
flowers opening in the evening; all other populations of the sub-
species appear to have flowers opening in the morning, near sunrise.
This taxon intergrades with C. claviformis subsp. integrior and occurs
sympatrically with C. scapoidea subsp. brachycarpa.

Four populations of this subspecies were tested for pollen-tube
growth following self- and cross-pollination and found to be self-
incompatible: 4.5 miles north of Sparks, Washoe County, Nevada
(8 plants) ; Steamboat Springs, Washoe County, Nevada (15 plants) ;
4.2 miles west of Rome, Malheur County, Oregon (3 plants, Kyhos
65-255); Jump Creek Canyon, Owyhee County, Idaho (13 plants).
10b. Camissonia claviformis subsp. lancifolia (Heller) Raven, Brittonia 16: 282.

1964.

Chylismia lancifolia Heller, Muhlenbergia 2: 226. 1906.

Oenothera clavaeformis var. cruciformis sensu Munz, Amer. Journ. Bot. 15:
235. 1928.

Oenothera clavaeformis subsp. lancifolia (Heller) Raven, Univ. Calif. Publ.
Bot. 34: 109. 1962.

Plants 5-60 cm. tall, strigose below, glabrous and often glaucous
above; rosette not strictly basal, the leaves erect. Lateral leaflets few
or wanting; terminal leaflet to 5 cm. long and 2.8 em. wide, lanceolate,
the apex acuminate, unevenly serrate, the serrations tipped with light
brown clusters of oil cells. Hypanthium 3.5-6 mm. long, glabrous
without and orange brown within. Sepals glabrous or sparsely strigose
near the apices. Petals 3.5-7 mm. long, 4-9 mm. wide, bright yellow,

usually red-dotted near the base. Filaments 3-5 mm. long; anthers

RAVEN—GENUS CAMISSONIA 215

3.5-5 mm. long. Style 9-16 mm. long. Capsule 1.1-2.8 cm. long,
glabrous or nearly so; pedicel 8-18 mm. long, Gametic chromosome
number, n =7.

Type: Base of the White Mountains east of Laws, Inyo County,
California, 11 May 1906, A. A. Heller 8231 (US 611229; isotypes,
CAS, COLO, DS, MO, UC).

DisrRIBUTION: Sandy slopes and flats, often with Artemisia tri-
dentata or Chrysothamnus, east of the Sierra Nevada in southern Mono
and Inyo Counties, California. From 4,000—5,500 ft. alt.

The southernmost station for C. claviformis subsp. laneifolra, in the
Owens Valley, Inyo County, California, 3 miles north of Big Pine, is
approximately 24 miles north of the northernmost for subsp. clav-
formis, 7.7 miles north of Independence. The yellow-flowered sub-
species reaches its peak of bloom several weeks later than the white-
flowered one. Camissonia claviformis subsp. lancifolia grows sym-
patrically with C. heterochroma.
10c. Camissonia claviformis subsp. integrior (Raven) Raven, Brittonia 16: 282.

1964.

Oenothera scapoidea var. purpurascens 8S. Wats., Proc. Amer. Acad. 8: 595.
1873. Lectotype: Not abundant, islands in Mono L., Mono Co., California,
10-11 July 1863, W. H. Brewer 1845 (GH; isotypes, UC, US).

Oenothera clavaeformis var. typica sensu Munz, Amer. Journ. Bot. 15: 236.
1928.

Oenothera clavaeformis subsp. integrior Raven, Univ. Calif. Publ. Bot. 34: 106.
1962.

Plants 6-70 cm. tall, strigose below and usually glandular-pubes-
cent above, the leaves sometimes purple-dotted. Lateral leaflets few
or absent; terminal leaflet to 7 cm. long and 3 cm. wide, narrowly
ovate to ovate, often subcordate at the base, serrate, the serrations
often tipped with clusters of oil cells. Hypanthium 3-6 mm. long,
strigose, glandular, or nearly glabrous without, orange brown within.
Sepals rarely purple-dotted, strigose, glandular, or nearly glabrous,
rarely with short free caudate portions. Petals 4.5-8 mm. long, 4-10
mm. wide, occasionally purple-dotted at base, fading purple. Fila-
ments 3-5.5 mm. long. Anthers 3-5.5 mm. long. Style 10-16 mm.
long. Capsule 13-25 mm. long, glabrous, strigose, or glandular; pedicel
15-40 mm. long. Gametic chromosome number, n=7. Self-
incompatible.

Type: Palisade, Eureka County, Nevada, 14 June 1882, M. E.
Jones 3869 (POM 38505; isotypes, CAS, DS, MO, NY, UC, US,
UTC).

DistrispuTion: Dry flats in Great Basin desert, with Artemisia
tridentata, Chrysothamnus, or Juniperus, often forming large colonies
on banks and flats, from southern Harney County, Oregon, southward
nearly throughout Nevada to Lincoln and Esmeralda Counties and to

216 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Mono and northeastern Inyo Counties, California. From 4,000-6,500
ft. alt.

This is the most widely distributed subspecies of C. claviformis,
and is markedly variable in kind and amount of pubescence. It inter-
grades with subsp. aurantiaca and with subsp. cruciformis and grows
sympatrically with C. brevipes subsp. brevipes (forming occasional
hybrids), (. heterochroma and C. scapoidea subsp. brachycarpa.
Three populations from Nevada were sampled for self-compatibility,
and 15 plants from 20 miles northeast of Lovelock, Pershing County
(R18542), 9 from Palisade, Eureka County, and 20 from 28.9 miles
southwest of Lovelock, Churchill County (Kyhos 65-254) were found
to be self-incompatible by pollen-tube growth.
10d. Camissonia claviformis subsp. funerea (Raven) Raven, Brittonia 16: 282.

1964.

Oenothera clavaeformis var. aurantiaca sensu Munz, Amer. Journ. Bot. 15:
237. 1928; pro parte.

Oenothera clavaeformis subsp. funerea Raven, Univ. Calif. Publ. Bot. 34: 106.
1962,

Plants 6-60 cm. tall, strigose below, often densely so. Lateral leaflets
well developed or more often wanting; terminal leaflet to 8 cm. long
and to 4.5 cm. wide, ovate, often cordate at the base, very prominent,
dentate. Hypanthium 3-5.5 mm. long, strigose without, usually
densely so, orange brown within. Sepals usually densely strigose,
with conspicuous free caudate segments arising on midribs just below
apices. Petals 3.5-7.5 mm. long, 3.5-7.5 mm. wide, often fading purple.
Filaments 3-5.5 mm. long; anthers 3-4.5 mm. long. Style 8-12 mm.
long, Capsule 1.2-3.2 cm. long, strigose; pedicel 8-22 mm. long.
Gametic chromosome number, n=7.

Type: Dry wash, 2 miles east of Stovepipe Wells, Death Valley,
Inyo County, California, 7 April 1928, P. A. Munz & C. L. Hitchcock
11030 (POM 159488; isotypes, GH, UC, US).

Distripution: Dry slopes and flats, with Larrea and Ambrosia
dumosa, Eureka Valley, Saline Valley, and the region of Death Valley,
Inyo County, and northernmost San Bernardino County, California.
From ca. 250 ft. below sea level to 3,000 ft. alt.

This taxon is geographically coherent and distinct in its more
extreme forms, but intergrades with subsp. aurantiaca and subsp.
claviformis over a broad front, producing an array of intermediate
forms. It occurs sympatrically with C. munzii and with C. brevipes
subsp. brevipes, forming occasional hybrids with each of these species.
10e. Camissonia claviformis subsp. claviformis

Oenothera clavaeformis Torr. & Frém., Rep. Expl. Exped. Rocky Mts. 314.
1845.

Oenothera scapoidea var. clavaeformis (Torr. & Frém.) 8. Wats., Bot. King
Expl. 109. 1871.

RAVEN—GENUS CAMISSONIA 217

Chylisma scapoidea var. clavaeformis (Torr. & Frém.) Small, Bull. Torrey Cl.
23: 194. 1896.

Chylisma clavaeformis (Torr. & Frém.) Heller, Muhlenbergia 2: 105. 1906.

Oenothera clavaeformis var. typica Munz, Amer. Journ. Bot. 15: 236. 1928.

Chylismia clavaeformis var. typica (Munz) Johansen, Amer. Journ. Bot. 16:
597. 1929.

Oenothera clavaeformis subsp. clavaeformis Raven, Univ. Calif. Publ. Bot. 34:
99. 1962.

Plants 6-55 cm. tall, glabrous or strigose below and occasionally
glandular, the leaves sometimes purple-dotted. Lateral leaflets usually
well developed; terminal leaflet to 6 cm. long and 3.5 cm. wide,
narrowly ovate, irregularly sinuate-dentate. Hypanthium 3-5.5 mm.
long, glabrous, sparsely strigose, or rarely glandular without, orange
brown within. Sepals glabrous, sparsely strigose, or rarely glandular,
with short-caudate terminal portions. Petals 3.5-8 mm. long, 4-8 mm.
wide, white, very rarely pale yellow, sometimes purple-dotted at
base, often fading purple. Filaments 3-5 mm. long; anthers 3-4 mm.
long. Style 8-12 mm. long. Capsule 1.2—3 cm. long, glabrous or sparse-
ly pubescent; pedicel 10-25 mm. long. Gametic chromosome number,
n =7. Self-incompatible.

Type: Frémont’s 2d Expedition (NY). Collected in 1844 by J. C.
Frémont. The herbarium specimen selected as the lectotype by Munz
(Amer. Journ. Bot. 15: 236. 1928) and cited by him as ‘‘Fremont’s
Pac. R. R. Exp., 1843-’44, ‘probably collected in California,’ ” has
not been located, but this may be an erroneous citation of a specimen
from the Torrey Herbarium (NY) labeled ‘Fremont’s Pacific R.
Road Expedition, 1853-54 (Probably collected in California).’’ This
collection might be an isotype, but if so it is incorrectly labeled.

Distrrpution: Alluvial slopes and flats, with Larrea and Ambrosia
dumosa, western Inyo County, eastern Kern County, northern Los
Angeles County, western San Bernardino County, and northern River-
side County, California, almost entirely on the Mojave Desert.
From 2,800—5,500 ft. alt.

This subspecies intergrades gradually and throughout a broad area
with subsp. aurantiaca and with subsp. funerea. It occurs sym-
patrically with C. brevipes subsp. brevipes, forming occasional hybrids
with this entity. Isolated plants of this subspecies fail to set fruit,
and more direct evidence of its self-incompatibility was provided by
a study of pollen-tube growth following self- and cross-pollination in
three plants from each of the following localities in California: near
Pearblossom, Los Angeles County, Kyhos 64-14; Joshua Tree, San
Bernardino County, AKyhos 64-33.
10f. Camissonia claviformis subsp. aurantiaca (S. Wats.) Raven, Brittonia 16:

282. 1964.
Oenothera scapoidea var. aurantiaca 8S. Wats., Proc. Amer. Acad. 8: 595. 1873.

Chylismia scapoidea var. aurantiaca (S. Wats.) A. Davids. & Moxley, FI. 8.
Calif. 254. 1923.

218 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Oenothera clavaeformis var. aurantiaca (S. Wats.) Munz, Amer. Journ. Bot.
15: 237. 1928.

Chylismia aurantiaca (S. Wats.) Johansen, Desert 3: 78. 1931.

Oenothera clavaeformis subsp. aurantiaca (S. Wats.) Raven, Univ. Calif.
Publ. Bot. 34: 100. 1962.

Plants 5-50 cm. tall, strigose, especially below, and rarely glandular-
pubescent above, the leaves sometimes purple-dotted. Lateral leaflets
up to 25 on each side of rachis, irregular, and well developed; terminal
leaflet to 3 cm. long and 1.5 em. wide, narrowly ovate, irregularly
sinuate-dentate. Hypanthium 3-5 mm. long, strigose without, orange
brown within. Sepals strigose, rarely with short free caudate tips in
bud. Petals 2.5-8 mm. long, 3.5-8 mm. wide, white, rarely purple-
dotted at base, often fading purple, rarely orange. Filaments 2-5 mm.
long; anthers 2.5-4.5 mm. long. Style 7.5-14 mm. long. Capsule
1.3-3 em. long, sparsely strigose or glabrous; pedicel 8-25 mm. long.
Gametic chromosome number, n=7. Self-incompatible.

Lectotype: Fort Mohave, Mohave County, Arizona, 1861, J. G.
Cooper (GH; isotypes, UC, US); Munz, Amer. Journ. Bot. 15: 237.
1928.

Disrrisution: Common on sandy flats and washes, with Larrea,
Ambrosia dumosa, and Fouquieria splendens, from Lincoln County,
Nevada, south to northeasternmost Baja California, and in Arizona
only in westernmost Mohave and Yuma Counties. From 230 ft. below
sea level to 3,000 ft. alt.

Camissonia claviformis subsp. aurantiaca is variable in leaf outline
and color and in amount of pubescence. It intergrades completely
with the other four white-flowered subspecies, claviformis, funerea,
integrior, and pebblesii. It likewise intergrades with subsp. yumae and
subsp. peirsonii where it comes in contact with them. Occasional
plants have large amounts of glandular pubescence in the inflorescence
and elsewhere, as subsp. peeblesii, but nowhere west of the Colorado
River do such individuals exceed 50 percent of any population known
to me. Camissonia claviformis subsp. aurantiaca grows sympatrically
with C. brevipes subsp. brevipes, C. brevipes subsp. pallidula, C. multi-
juga, and C. munzii, forming occasional hybrids with the first two and
last entities. Three individuals each from two populations of this
subspecies from California were shown to be self-incompatible by
pollen-tube growth in selfed and outcrossed flowers: 0.7 mile from
U.S. Hwy. 99 on road to Snow Creek, Riverside County, Kyhos 64-3;
4.3 miles south of Mortmar, Imperial County, Kyhos 64-13.
10g. Camissonia claviformis subsp. peeblesii (Munz) Raven, Brittonia 16: 282.

1964,

Oenothera clavaeformis var. aurantiaca sensu Munz, Amer. Journ. Bot. 15:
237. 1928.
Oenothera clavaeformis var. peeblesii Munz, Leafl. West. Bot. 2: 158. 1939.

RAVEN—GENUS CAMISSONIA 219

Camissonia clavaeformis subsp. peeblesii (Munz) Raven, Univ. Calif. Publ.
Bot. 34: 101. 1962.

Plants 5-60 cm. tall, glandular-pubescent, especially above, and
often also strigose below. Leaves glandular-pubescent, the lateral
leaflets numerous, irregular, and well developed, the terminal leaflet
narrowly ovate, irregularly sinuate-dentate, to 7 cm. long and 3 cm.
wide. Hypanthium 3-5.5 mm. long, glandular-pubescent or strigose
without, orange brown within. Sepals glandular-pubescent, strigose,
or both, usually lacking caudate appendage. Petals 3-7.5 mm. long,
3-5 mm. wide, often fading purple. Filaments 2-3 mm. long; anthers
2.5-4 mm. long. Style 7.5-14 mm. long. Capsule 1.5-3 cm. long,
glandular-pubescent or strigose; pedicel 4-32 mm. long. Gametic
chromosome number, n=7.

TypEe: Casa Grande, Pinal County, Arizona, 25 February 1927,
Peebles & Harrison 3537 (US 1367424; isotype, ARIZ).

DistriBuTION: Flat sandy plains and washes, with Prosopis,
Carnegiea gigantea, Larrea and Ambrosia dumosa, throughout almost
all the southwestern half of Arizona and locally in northwesternmost
Sonora. From 400-2,000 ft. alt.

I have included in Camissonia claviformis subsp. peeblesvi all popula-
tions in which a majority of the plants have some glandular pubescence,
and the resulting taxon has a coherent geographical range. It inter-
grades completely with subsp. aurantiaca and with subsp. rubescens,
growing in part in the same area as the latter. It also occurs sympat-
rically with all three subspecies of C. brevipes, forming occasional
hybrids with them.
10h. Camissonia claviformis subsp. peirsonii (Munz) Raven, Brittonia 16: 282.

1964.
Oenothera clavaeformis var. peirsonit Munz, Amer. Journ. Bot. 15: 238. 1928.
Chylismia peirsonit (Munz) Johansen (as “piersonii’’), Desert 3: 78. 1931.
Oenothera clavaeformis subsp. peirsonii (Munz) Raven, Univ. Calif. Publ.
Bot. 34: 105. 1962.

Plants 5-60 cm. tall, spreading-villous below and often above,
more rarely strigose or with glandular pubescence. Lateral leaflets
usually well developed; terminal leaflets to 9 cm. long and to 3.5 cm.
wide, narrowly ovate, irregularly sinuate-dentate. Hypanthium
2.5-4.5 mm. long, villous or strigose without, orange brown within.
Sepals villous or strigose, with conspicuous free caudate segments
arising just below apices of midribs. Petals 4.5-7 mm. long, 4-9 mm.
wide, yellow, more rarely white, not changing color in fading. Fila-
ments 3.5-5 mm. long; anthers 3.5-5 mm. long. Style 9-14 mm. long.
Capsule 1.2-3.8 cm. long, spreading pubescent (rarely glabrous) ;
pedicel 10-20 mm. long. Gametic chromosome number, n=7.

Typrr: Twenty-eight miles south of Coachella, Imperial County,
California, 12 April 1922, F. W. Peirson 4512 (POM 138409; isotype,
RSA).

220 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Distrrpution: Sandy fields and washes, with Larrea, Ambrosia
dumosa, and Fouquieria splendens, east of the Salton Sea and Imperial
Valley in San Diego and Imperial Counties, California, and in north-
eastern Baja California to the Gulf of California. From 240 ft. below
sea level to 1,000 ft. alt.

This subspecies intergrades with subsp. aurantiaca in northeastern
San Diego County, California, where it is extremely variable in
pubescence, leaf color, and flower color. Since publishing my revision
of the group, I have seen a specimen of C. claviformis subsp. peirsonii
from 41 miles south of San Felipe, Baja California, Wiggins 18162
(DS), this record establishing a new southern limit for the subspecies.
One additional determination of chromosome number has been made
in this entity, n=7, in Klein 1175 (RSA), from 2 miles south of San
Felipe, Baja California.
10i. Camissonia claviformis subsp. yumae (Raven) Raven, Brittonia 16: 282. 1964.

Oenothera clavaeformis subsp. yumae Raven, Univ. Calif. Publ. Bot. 34: 104.
1962.

Plants 5-40 em. tall, strigose pubescent below, often densely, and
sometimes glandular-pubescent above. Lateral leaflets well developed
or reduced; terminal leaflet to 6.5 em. long and to 2 cm. wide, lance-
olate, irregularly sinuate-dentate. Hypanthium 2.5-4 mm. long,
strigose without, orange brown within. Sepals strigose with short free
caudate portions arising just below the apices, or these absent. Petals
3-5 mm. long, 4-5.5 mm. wide, pale yellow, fading reddish or not
changing color after pollination. Filaments 2-4 mm. long, anthers
2-4 mm. long. Style 7.5-10 mm. long. Capsule 1.2-3.2 em. long,
strigose; pedicel 5-30 mm. long. Gametic chromosome number,
n=7. Self-incompatible.

Type: Covering dunes, Yuma Desert 4.8 miles west of summit of
Telegraph Pass, Yuma County, Arizona, 28 February 1958, P. H.
Raven 11724 (RSA).

Distripution: Very dry dunes and sandy flats with Larrea and
Ambrosia dumosa, southeastern Imperial County, California, Yuma
Desert, Arizona, and from El Gran Desierto to Puerto Penasco in
northwestern Sonora, and in northeastern Baja California (8 miles
northwest of La Ventana, Moran 12354, DS, SD). From sea level to
300 ft. alt.

This subspecies, which is unusually variable, was almost. cer-
tainly derived following hybridization between the yellow-flowered
subsp. peirsonii and the white-flowered subsp. aurantiaca. It inter-
grades with subsp. aurantiaca in California and Baja California
and with subsp. peeblesii in Arizona and Sonora. It erows sympatri-
cally with C. brevipes subsp. arizonica, but no hybrids have been
found. Three individuals from a population along U.S. Hwy. 80,

RAVEN—GENUS CAMISSONIA 221

0.4 mile west of the junction with U.S. Hwy. 98, Imperial County,
California, Kyhos 64-11, were found to be self-incompatible, judging
from pollen-tube growth in self-pollinated flowers.
10j. Camissonia claviformis subsp. wigginsii (Raven) Raven, Brittonia 16:
282. 1964.
Oenothera clavaeformis subsp. wigginsii Raven, Univ. Calif. Publ. Bot. 34: 103.
1962.

Plants 5-20 cm. tall, covered with spreading white trichomes
below and glandular pubescence above. Lateral leaflets reduced or
absent; terminal leaflet to 3 cm. long and to 1.5 cm. wide, narrowly
ovate, irregularly sinuate-dentate. Hypanthium 1.5-2 mm. long,
villous and often purple-dotted without, orange brown within. Se-
pals villous, often purple-dotted, with short free caudate portions
arising on midribs below the apices. Petals 1.5-2 mm. long, 1.5-2
mm. wide, yellow, fading brick red after pollination. Filaments 1.5-2
mm long; anthers 1.5-2 mm. long. Style 5-7 mm. long. Capsule (1.5-)
2.5-3.5 cm. long, often purple-dotted, villous; pedicel 8-20 mm. long.
Gametic chromosome number, n=7.

Type: Wash 3 miles from Los Angeles Bay, Baja California, Mex-
ico, 20 February 1935, J. L. Wiggins 7684 (DS 265606; isotypes F,
GH, UC, US).

DistripuTion: Central Baja California, in washes. From ca. 500-
ca. 2,000 ft. alt.

This is apparently a rare and rather restricted subspecies most
similar to subsp. peirsonii and subsp. rubescens, from which it differs
principally in flower size. It is, however, separated from the former
subspecies, the nearest geographically within the species, by a gap
of about 75 miles. In spite of its relatively small flowers, the stigma
is held well above the anthers and this subspecies is presumably self-
incompatible like the rest of the species. The chromosome number
was determined in a collection from the Arroyo San Francisquito
5 miles northwest of Las Arrastras, 29°37’ N. lat., 114°27’ W. long.,
Moran 12402 (DS, SD).
10k. Camissonia claviformis subsp. rubescens (Raven) Raven, Brittonia 16:

282. 1964.
Oenothera clavaeformis subsp. rubescens Raven, Univ. Calif. Publ. Bot. 34:
103. 1962.

Plants 5-40 cm. tall, covered, especially below, with spreading
white trichomes, and also with short glandular trichomes. Lateral
leaflets few or absent; terminal leaflet to 4 cm. long and 2 cm. wide,
lanceolate, irregularly dentate. Hypanthium 3.5-5 mm. long, glandu-
lar-pubescent or villous or both, orange brown within. Sepals villous
or glandular-pubescent, with conspicuous free caudate segments
arising from midribs below the apices. Petals 3-5 mm. long, 3-5
mm. wide, yellow, fading brick red after pollination. Filaments 2.5-5

222 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

mm. long; anthers 2.5-4 mm. long. Style 7-12 mm. long. Gametic
chromosome number, n=7.

Type: On sandy flats, 24 miles south of Sonoyta on road to Rocky
Point (Punta Pefiasco), Distrito de Altar, Sonora, 14 March 1936,
D. D. Keck 4178 (DS 287997; isotypes, GH, POM, UC, US).

DistriBuTION: Sandy flats, with Larrea, Ambrosia dumosa, and
Carnegiea gigantea, western Maricopa, eastern Pima, and southeastern
Yuma Counties, Arizona, and northwestern Sonora. From 200—1,000
ft. alt.

This entity grows sympatrically with C. brevipes subsp. arizonica
and occasionally forms natural hybrids with it. It intergrades with
C. claviformis subsp. peeblesii.

11. Camissonia munzii (Raven) Raven, Brittonia 16: 281. 1964.
Oenothera munzii Raven, Univ. Calif. Publ. Bot. 34: 91. 1962.

Annual herb 8-50 cm. tall, with numerous branches at the base
and above, the leaves mostly in a subbasal rosette. Stems and leaves
strigose, often densely so above and in the inflorescence. Leaves to
20 cm. long, pinnate, with well-developed lateral leaflets; terminal
leaflet ovate, to 6 cm. long and 3 cm. wide, the veins lined below
with brownish oil cells. Inflorescence nodding, not congested, elongat-
ing in mature bud. Hypanthium 2-3 mm. long, 2-3 mm. across at
the summit, strigose without, villous and orange brown within. Sepals
4—7 mm. long, 1.5-2 mm. wide, strigose. Petals 3-10 mm. long, 3-10
mm. wide, bright yellow, red-dotted near the base. Stamens subequal,
the filaments 4-8 mm. long; anthers 3-6 mm. long, ciliate. Style 8-18
mm. long; stigma 0.6-0.8 mm. thick, held well above the anthers at
anthesis. Capsule 0.8-2.4 cm. long, 1.5-2 mm. thick, clavate; pedicel
8-28 mm. long, widely divergent and becoming sharply deflexed at
maturity. Seeds pale brown, 0.8-1.6 mm. long, 0.5-0.8 mm, thick.
Gametic chromosome number, n—7. Self-incompatible.

TypE: Sandy desert wash, Salsberry Pass, south end of Death
Valley, Inyo County, California, 9 April 1940, P. A. Afunz 16474
(POM 255191; isotypes, CAS, GH, NA, POM, RSA, UC, US, WS,
WTU).

Disrripution: Relatively mesic slopes and washes at middle ele-
vations in the mountains at the north end, eastward, and south of
Death Valley, from Saline Valley and the Grapevine Mountains, Inyo
County, California, and Yucca Flat, Nye County, Nevada, south-
ward to the Kingston Range, San Bernardino County, California.
From 2,000-5,000 ft. alt.

Camissonia munzii grows sympatrically with C. brevipes subsp.
brevipes, C. brevipes subsp. pallidula, C. claviformis subsp. funerea, C.
claviformis subsp. aurantiaca, and C. heterochroma, forming occasional
hybrids with the first and fourth mentioned taxa. It is readily dis-

RAVEN—GENUS CAMISSONIA 223

tinguished by its long, deflexed pedicels and clavate capsules. A popu-
lation from 7.2 miles south of Mercury, Nye County, Nevada, R18865,
was tested for self-compatibility, and 14 individuals were found to
be self-incompatible by pollen-tube growth.
12. Camissonia eastwoodiae (Munz) Raven, Brittonia 16: 282. 1964.
Oenothera scapoidea var. eastwoodae Munz, Amer. Journ. Bot. 15: 234. 1928.
Oenothera eastwoodiae (Munz) Raven, Univ. Calif. Publ. Bot. 34: 92. 1962.

Succulent annual 3-30 cm. tall, the stems glabrous, glandular-
pubescent, or villous below. Leaves mostly basal, entire or sparsely
denticulate, simple, the blades oblanceolate to cordate, darker green
above, 0.8-7.5 cm. long, 0.4-3 cm. wide; veins below lined with pale
brown oil cells; petioles 0.5-8 cm. long. Inflorescence drooping,
elongating in flower and fruit. Hypanthium 2-4.5 mm. long, 1.5-3.4
mm. across at the summit, glandular-pubescent or strigose without,
villous in lower portions within. Sepals 3-8 mm. long, 1-2.5 mm.
wide, glandular-pubescent, strigose, or glabrous, without caudate
appendage. Petals 5.5-9 mm. long, 4-10 mm. wide, bright yellow and
of same color as stamens, style, and inside of hypanthium, red-
dotted near the base. Filaments of the episepalous stamens 3-8
mm. long, those of the epipetalous ones 2.8-5.5 mm. long; anthers
2-4 mm. long, ciliate. Style 10-17 mm. long; stigma 0.6-0.8 mm. thick,
held well above the stamens at anthesis, villous near the base. Capsule
1.8-4 cm. long, curved, erect, on a spreading or slightly deflexed
pedicel 4-28 mm. long. Seeds tan, 1.2-1.7 mm. long. Gametic chromo-
some number, n=7.

Type: Grand Junction, Mesa County, Colorado, May 1892,
A. Eastwood (GH; isotypes, UC, US).

Distripution: Clay flats, on gray alkaline marine-deposited
gumbo, and in sandy draws, Mesa County, Colorado, and Emery
County, Utah, south to San Juan County, Utah. From 4,000-5,500
ft. alt.

Camissonia eastwoodiae, the only outcrossing species in the eastern
part of the range of sect. Chylismia, is undoubtedly self-incompatible.
It grows sympatrically with the autogamous C. scapoidea subsp.
scapoidea and in the same vicinity as the ecologically distinct C.
walkeri subsp. walkeri.

13. Camissonia scapoidea (Torr. & Gray) Raven, Brittonia 16: 95. 1964.

Annual 3-45 cm. tall; stems strigose, villous, or glandular-pubescent
below. Leaves mostly in a basal rosette, greatly reduced upward,
spatulate, 1-18 cm. long, the lateral leaflets well developed or absent.
Inflorescence drooping, glandular-pubescent, strigose, or nearly
glabrous. Hypanthium 1-4 mm. long, 1-3.5 mm. across at the summit,
glandular-pubescent, strigose, or glabrous’ without, sparsely villous
to glabrous within. Sepals 1.2-5 mm. long, 1-2 mm. wide, strigose,

295-655 O—68 3

224 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

glandular-pubescent, or glabrous, entire or short-caudate at the
apices. Petals 1.5-5.5 mm. long, 1-4 mm. wide, bright yellow, of
same color as stamens, style, and inside of hypanthium. Filaments
of the episepalous stamens 1.2-6 mm. long, those of the epipetalous
ones 0.5-4 mm.; anthers 1-2.5 mm. long, ciliate or glabrous. Style
glabrous or short-villous at the base, 3-11 mm. long; stigma 0.3-0.6
mm. thick, surrounded by anthers, at least the longer ones, at anthesis.
Capsule ascending, curved to nearly straight, 0.8-4 cm. long, 1.8-2.6
mm. thick, sparsely pubescent to glabrous, the valves sometimes
twisted at maturity, on an ascending or spreading pedicel 4-20 mm.
long. Seeds pale brown, 1-2 mm. long. Gametic chromosome numbers,
n=7, 14. Autogamous.

DisTRIBUTION: Sandy or clayey slopes and flats, from south-
eastern Oregon, southwestern and central Idaho, and western and
central Wyoming, western Colorado, northeastern Arizona, and
adjacent New Mexico, throughout Utah, and in northeastern Nevada.
From 2,500-6,000 ft. alt.

13a. Camissonia scapoidea subsp. scapoidea

Oenothera scapoidea Torr. & Gray, Fl. N. Amer. 1: 506. 1840.

Chylisma scapoidea (Torr. & Gray) Small, Bull. Torrey Cl. 23: 193. 1896.

Oenothera brevipes var. scapoidea (Torr. & Gray) H. Lév., Monogr. Onoth.
146. 1905.

Chylismia scapoidea var. seorsa A. Nels., Bot. Gaz. 54: 140. 1912. Lectotype:
Evanston, Uinta Co., Wyoming, 27 July 1897, A. Nelson 4125 (RM 10458).

Oenothera scapoidea var. typica Munz, Amer. Journ. Bot. 15: 233. 1928.

Oenothera scapoidea var. seorsa (A. Nels.) Munz, Amer. Journ. Bot. 15: 233.
1928.

Oenothera scapoidea subsp. scapoidea Raven, Univ. Calif. Publ. Bot. 34: 94.
1962.

Leaves usually simple, more rarely with a few reduced lateral
leaflets, the blades narrowly ovate to ovate, often cordate at the
base, to 5.5 em. long and 3 cm. wide, subglabrous, very rarely villous
on both surfaces, the oil cells below pale yellowish brown, rarely
darker. Petals acute, 1.7-5 mm. long, 1.5-3.5 mm. wide, often red-
dotted near the base. Filaments of the episepalous stamens 1-4 mm.
long, those of the epipetalous ones 0.5-2.5 mm. long; anthers 0.5-1.9
mm. long, glabrous or ciliate. Style 2.5-9 mm. long. Capsule (1-)1.5-3
cm. long; pedicel 5-18 mm. long. Gametic chromosome numbers
n=7, 14. Autogamous.

Type: Clay hills, probably near the rendezvous of 1834, on Ham’s
Fork of the Green River, Sweetwater County, Wyoming, 22 June to
2 July 1834, T. Nuttall (NY; isotypes BM, GH). See Raven, Univ.
Calif. Publ. Bot. 34: 93. 1962.

Disrripution: Sandy or clay flats, throughout western Wyoming,
west to Lake County, Oregon, and Elko and White Pine Counties,

RAVEN—GENUS CAMISSONIA 225

Nevada, south to Beaver County, Utah, northern Coconino, Navajo,
and Apache Counties, Arizona, San Juan County, New Mexico, and
western Colorado; also on the headwaters of the Arkansas River in
eastern Colorado. The only member of sect. Chylismia that occurs
east of the Continental Divide. From 4,000-8,000 ft. alt.

Camissonia scapoidea subsp. scapoidea grows sympatrically with
C. eastwoodiae and in the same vicinity as the ecologically distinct
C. walkeri subsp. walkeri. It is highly variable, as would be expected
in such widely ranging, principally autogamous entity. Both diploid
and tetraploid populations occur in this taxon. The first series of
tetraploids I studied (Univ. Calif. Publ. Bot. 34: 38-39. 1962), from
Emery County, Utah, was growing with C. eastwoodiae and diploid
plants of C. scapoidea subsp. scapoidea. At the time, it seemed possible
that these tetraploids might have had an allopolyploid origin between
the two diploids. It seemed even more likely, however, that they
were autotetraploids derived from diploid C. scapordea subsp.
scapoidea, from which they were morphologically almost indistinguish-
able. This possibility is now considerably strengthened by the dis-
covery of a tetraploid individual in Sweetwater County, Wyoming,
along Interstate Hwy. 80, 4.8 miles southwest of junction with US.
Hwy. 30N, R19544. This locality is some 150 miles north of the nearest
station for Camissonia eastwoodiae. The plant examined formed 14
pairs of chromosomes, like the individuals from Utah studied earlier.
A diploid individual was likewise found in southwestern Wyoming,
1.7 miles west of Fort Bridger, Uinta County, Mosquin & Mulligan
5128. These two chromosome counts constitute the first from the
northern portion of the range of the subspecies and suggest the neces-
sity for further chromosome counts of this taxon before its cy tological
pattern of variation can properly be assessed.
13b. Camissonia scapoidea subsp. macrocarpa (Raven) Raven, Brittonia 16: 282.

1964.
Ocenothera scapoidea subsp. macrocarpa Raven, Univ. Calif. Publ. Bot. 34: 95.
1962.

Basal rosette compact, the leaves simple, rarely with reduced
lateral leaflets. Leaf blades ovate, often cordate at the base, to 3.5 cm.
long and 1.5 cm. wide, nearly glabrous, the oil cells below pale yellowish
brown. Petals 1.5-2 mm. long, ca. 1.5 mm. wide. Filaments of the
episepalous stamens ca. 2 mm. long, those of the epipetalous ones ca.
1 mm. long; anthers ca. 1 mm. long, ciliate. Style ca. 3 mm. long. Cap-
sule 2.5-4 cm. long, stout, very large for size of plant, ascending;
pedicel 4-6 mm. long. Autogamous.

Type: 9 miles northeast of Black Point, 5,700 ft., Apache County,
Arizona, 11 June 1937, R. H. Peebles & E. G. Smith 13529 (US 1739264;
isotypes, ARIZ, US).

226 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Distripution: On detrital clay knobs and gravelly flats, northern
Apache and Navajo Counties and northeastern Coconino County,
Arizona. From 5,000-6,500 ft. alt.
13c. Camissonia scapoidea subsp. brachycarpa (Raven) Raven, Brittonia 16: 282.

1964.
Oenothera scapoidea subsp. brachycarpa Raven, Univ. Calif. Publ. Bot. 34: 95.
1962.

Slender, the leaves simple or often pinnate with well-developed
lateral leaflets. Terminal leaflets narrowly ovate, often cordate at the
base, to 10 cm. long and 5 cm. wide, glabrous or weakly pubescent,
the oil cells below dark brown, prominent. Petals 1.8-4 mm. long,
acute, 1-4 mm. wide, often red-dotted near the base. Filaments of the
episepalous stamens 1.2-3.2 mm. long, those of the epipetalous ones
0.5-1.5 mm. long; anthers 0.9-1.8 mm. long, ciliate. Style 4-7.5 mm.
long. Capsule 0.8-2 (-2.8) em. long; pedicel 10-16 mm. long. Gametic
chromosome number, n=7. Autogamous.

Type: Loose dry open slopes with sagebrush and shrubs, 7 miles
north of Brogan, T. 148., R. 42 E., sec. 27, 3,000 ft., Malheur County,
Oregon, 10 June 1955, A. Cronquist 7808 (NY; isotypes, DS, GH, ID,
MO, RM, RSA, UC, US, UTC, WS, WTU).

Distripution: Sandy slopes and flats, with Grayia, Artemisia, or
Juniperus, southeastern Oregon, southwestern Idaho, northwestern
Utah, and northeastern Nevada. From 2,500-6,000 ft. alt.

Camissonia scapoidea subsp. brachycarpa grows sympatrically with C.
claviformis subsp. cruciformis and subsp. integrior, and overlaps morpho-
logically with C. seapoidea subsp. seapoidea, from which it is separated
geographically. The possible derivation of this species from populations
similar to C. claviformis subsp. cruciformis and C. eastwoodiae has been
discussed earlier (Raven, Univ. Calif. Publ. Bot. 34: 71. 1962).
13d. C. scapoidea subsp. utahensis (Raven) Raven, Brittonia 16; 282. 1964.

Oenothera scapoidea subsp. utahensis Raven, Univ. Calif. Publ. Bot. 34: 96.
1962.

Leaves pinnate, sparsely pubescent, the lateral leaflets well devel-
oped, up to 1 cm. long; terminal segment ovate, cordate at the base,
up to 4.5 em. long and 3 cm. wide, the oil cells below dark brown.
Petals 4-5.5 (-8) mm. long, 3-4 mm. wide, blunt, often red-dotted
near the base. Filaments of the episepalous stamens 5-6 mm. long,
those of the epipetalous ones 3.5-4 mm. long; anthers 2-2.5 mm.
long, ciliate. Style 8-11 mm. long. Capsule 1.6-3.8 em. long; pedicel
8-20 mm. long. Autogamous.

Type: Black Rock, 4,300 ft., Salt Lake County, Utah, June 1869,
S. Watson (US 70527; isotype, GH).

Disrrisution: On dry rocky slopes and flats, with Atriplex and
Chrysothamnus, western Utah and extreme northeastern Nevada (Elko
County). From 4,100-5,500 ft. alt.

RAVEN—GENUS CAMISSONIA 227

No new information is available about this enigmatic subspecies,
which stands in an intermediate position, both morphologically and
geographically, between C. scapoidea and C. walkeri subsp. tortilis.
14. Camissonia parryi (S. Wats.) Raven, Brittonia 16: 282. 1964.

Oenothera parryi 8. Wats., Amer. Nat. 9: 270. 1875.

Oenothera scapoidea var. parryi (S. Wats.) M. E. Jones, Proc. Calif. Acad. IT.
5: 682. 1895.

Oenothera tenuissima M. E. Jones, Proc. Calif. Acad. II. 5: 683. 1895. Type:
In clay washes, Rockville, 4,000 ft., Washington Co., Utah, 26 September
1894, M. E. Jones 6083 (POM 38644; isotypes, MO, NY, UC, US).

Chylisma parryi (S. Wats.) Small, Bull. Torrey Cl. 23: 193. 1896.

Oenothera brevipes race parryi (S. Wats.) H. Lév., Monogr. Onoth. 146. 1905.

Chylismia tenuissima (M. E. Jones) Rydb., Bull. Torrey Cl. 40: 66. 1913.

Oenothera parryi f. tenwissima (M. E. Jones) Raven, Univ. Calif. Publ. Bot.
34: 111. 1962.

Erect and often intricately branched annual herb 5-80 cm. tall, with
poorly defined basal rosette, densely covered below with white
trichomes ca. 2 mm. long, forming a villous pubescence on the stems
and leaves. Leaves simple or very rarely with a few small lateral
leaflets, ovate, often cordate at the base, sparsely dentate, reduced
upward, the veins below lined with pale or dark brown oil cells.
Inflorescences nodding, mostly glabrous, with intricate, filiform
branches, corymbiform. Hypanthium 0.5-2 mm. long, 0.3-0.5 mm.
across at the summit, glabrous or villous without and within. Sepals
1.5-4 mm. long, 0.5-1.3 mm. wide, villous to glabrous, with clusters
of light-brown oil cells at tip, the caudate appendages absent. Petals
2-7 mm. long, 2-6 mm. wide, bright yellow, of same color as stamens,
style, and inside of hypanthium, often red-dotted near the base,
sometimes fading rose. Filaments of the episepalous stamens 1.7—3.5
mm. long, those of the epipetalous ones 1.2-2.5 mm. long; anthers
0.9-1.2 mm. long, glabrous. Style 4-9 mm. long, glabrous; stigma
0.3-0.6 mm. thick, held well above the anthers at anthesis. Capsule
0.4-1 em. long, 1.2-1.5 mm. thick, glabrous or glandular-pubescent,
on a filiform pedicel 4-20 mm. long, the pedicel widely spreading or
reflexed but the capsule erect. Seeds light brown, finely pitted, lenticu-
lar, with narrow cellular rim, few and crowded in 4-septate capsule
so as to appear 1-rowed, 0.7—-1.2 mm. long, 0.5-0.6 mm. thick. Gametic
chromosome number, n=7. Outcrossing.

Type: Abundant on bare gypseous clay hills near St. George,
Washington County, Utah, 1874, C. C. Parry 72 (GH; isotypes, DS,
F, GH, MO, ND, NY).

Distripution: Red clay and sand slopes weathered from red
(fresh-water-deposited) sandstone cliffs, with Larrea or Juniperus,
southwestern Utah and northwestern Arizona. From 2,700-4,000 ft.
alt.

228 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Ficure 6.—Camissonia arenaria, X 3% (Raven 11743, DS).

Camissonia parry? is highly restricted geographically and edaphically
and distinctive morphologically. It no longer appears desirable to
accord formal taxonomic recognition to the late season plants Jones
described as Oenthera tenuissima. Camissonia parryi is very likely
self-incompatible. It grows sympatrically with C. multijuga, but the

RAVEN—GENUS CAMISSONIA 229

latter is confined to firm rock walls, whereas C. parryi grows on red-

clay flats, which it often carpets.

15. Camissonia exilis (Raven) Raven, Brittonia 16: 383. 1964. FIGURE 7
Oenothera exilis Raven, Univ. Calif. Publ. Bot. 34: 114. 1962.

Slender and copiously branched, with purplish-brown stems,
covered with glandular puberulence, and with occasional scattered
white-villous pubescence. Leaves to 3.5 cm. long, reduced upward,
with narrowly ovate blades to 2 cm. long and 0.7 em. wide, the margins
feebly denticulate, the veins below lined with brownish oil cells.
Inflorescence erect, sparse, elongating in fruit, each flower subtended
by a bract to 3.5 mm. long, with a petiole to 1 mm. long. Hypanthium
0.4-0.5 mm. long, 0.4 mm. across at the summit, glabrous within.
Sepals 1-1.2 mm. long, ca. 0.5 mm. wide, with small clusters of oil
cells at apices. Petals 1-1.5 mm. long, ca. 1 mm. wide, yellow fading
purplish, the same color as stamens, style, and inside of hypanthium.
Stamens 4, episepalous; filaments ca. 0.5 mm. long; anthers ca. 0.5
mm. long, glabrous. Style ca. 1.5 mm. long, glabrous; stigma ca. 0.2
mm. thick, surrounded by the anthers at anthesis. Capsule 0.45-0.7
em. long, ca. 1.5 mm. thick, clavate; pedicel 4-9 mm. long; pedicels
spreading, the capsules spreading or ascending. Seeds light brown,
ca. 0.8 mm. long, ca. 0.5 mm. wide. Gametic chromosome number,
n=7. Autogamous.

Type: “Gypsum-covered flat,” near Cottonwood Spring, east
base of Virgin Mountains, 3,500 ft., Mohave County, Arizona, 6 June
1941, P. A. Munz 16779 (POM 264197; isotype, DS).

DistriBution: Travertine-covered flats, east base of Virgin Moun-
tains, ca. 3,500 ft., Mohave County, Arizona; Buckskin Mountains,
between Willow Springs and Lees Ferry, ca. 5,000 ft., Coconino
County, Arizona.

This exceedingly rare and local species is the only member of the
genus characterized by the absence of the epipetalous stamens. On
the other hand, certain plants of C. andina have also lost this whorl
of stamens, and the epipetalous stamens are sometimes greatly re-
duced in C. minor. In all other respects, C. ezilis is closely similar to
C. parryi and to other species of sect. Chylismia, and I no longer
recognize it as constituting a distinct, monotypic section, which
procedure would confer undue emphasis to a single distinctive
characteristic.

16. Camissonia megalantha (Munz) Raven, Brittonia 16: 282. 1964.
Oenothera heterochroma var. megalantha Munz, Leafl. West. Bot. 3: 52. 1941.
Oenothera megalantha (Munz) Raven, Univ. Calif. Publ. Bot. 34: 111. 1962.

Robust annual with glandular pubescence, 0.1-2 m. tall, with an
ill-defined basal rosette, the cauline leaves larger below; secondary
branches numerous. Leaves simple, broadly ovate, cordate at the

CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

230

(SC “9601 2¢npo3190gq) vsoytpr9043
‘OD ‘8 ‘(Sq ‘edAjos: “Ez/9] zunpy) sipxa ry ‘LM X ‘wiuosstwpy yo saisadg—g-/ saunorgy

RAVEN—GENUS CAMISSONIA 231

base, sinuate-dentate, the veins below prominently lined with yellowish
oil cells, the blades up to 8 cm. long and 7 cm. wide, the petioles to
5.5 cm. long. Hypanthium 4.5-8.5 mm. long, 1.5-3 mm. across at the
summit, glandular-pubescent without, cream (greenish yellow near
summit) with matted villous pubescence within, except for the glabrous
lower 2 mm. Sepals 4.5-9 mm. long, 1.5-2.4 mm. wide, with clusters
of oil cells at the apices, often reflexed in pairs; caudate appendages
absent. Petals 9-13.8 mm. long, 6-15.5 mm. wide, pale to dark
lavender, diffusely flecked with purplish near the base, and white at
the very base. Filaments pale greenish yellow, those of the episepalous
stamens 6-12 mm. long, those of the epipetalous ones 3.5-8 mm. long;
anthers ca. 2 mm. long, glabrous. Style 14-22.5 mm. long, white,
densely villous above the lowest 3-4 mm., up to a point above the
summit of the hypanthium (ca. 5-5.5 mm. below the stigma) ; stigma
depressed-globose, ca. 1 mm. across, ca. 0.5 mm. high, held well
above the anthers at anthesis. Capsule 0.8-1.4 cm. long, 1.3-2 mm.
thick, clavate; pedicel 2-3.5 mm. long. Seeds brown, 1-1.3 mm. long.
Gametic chromosome number, n=7. Outcrossing but self-compatible.

Type: Volcanic alkali soil, Cane Springs, Skull Mountains, Nye
County, Nevada, 24 August 1938, P. Train 2358 (POM 253923; iso-
types, ARIZ, CAS, DS).

DistripuTion: Known only from rubble derived from volcanic
tuff, partly on moist soil about springs, at the type locality, at ca.
4,100 ft. alt., in southern Nye County, Nevada. Flowering mostly from
June to October; germinating in the late spring.

Camissonia megalantha is outcrossed and visited by such large,
polylectic bees as Anthophora urbana Cress. and Centris rhodopus
Ckll. soon after its flowers open near sunrise (Raven, MS.). It is,
however, self-compatible, as shown by comparative studies of pollen-
tube growth in 46 selfed and outcrossed individuals from the type
locality. This species and its obvious derivative, the autogamous
C. heterochroma, are the only lavender-flowered species in the genus
Camissonia, and two of the very few that regularly bloom in the
autumn. They are highly specialized vegetatively within this group.
In the tribe Onagreae, species with lavender or purplish petals are
found in the following genera: Gongylocarpus, Clarkia, Heterogaura,
and Oenothera. In species of most genera, however, the petals may fade
purplish after fertilization, indicating that the capability of producing
such pigments exists throughout the tribe. Camissonia megalantha
occurs sympatrically with C. claviformis subsp. claviformis.

17. Camissonia heterochroma (S. Wats.) Raven, Brittonia 16: 282. 1964.
Oenothera heterochroma S. Wats., Proc. Am. Acad. 17: 373. 1882.

Chylisma heterochroma (S. Wats.) Small, Bull. Torrey Cl. 23: 193. 1896.
Oenothera heterochroma var. typica Munz, Aliso 2: 83. 1949.

232 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Oenothera heterochroma var. monoensis Munz, Aliso 2: 84.1949. Type: Sherwin
Grade, 5,500 ft., Mono Co., California, 16 August 1947, J. 7’. Howell 24172
(CAS 342092; isotypes, DS, RSA).

Oenothera heterochroma subsp. heterochroma Raven, Univ. Calif. Publ. Bot.
34: 112. 1962.

Oenothera heterochroma subsp. monoensis (Munz) Raven, Univ. Calif. Publ.
Bot. 34: 113. 1962.

Annual herb 0.1-1 m. tall, with an ill-defined basal rosette; second-
ary branches numerous; entire plant covered with glandular pubes-
cence, or nearly glabrous and glaucous above. Leaves simple, ovate,
often cordate at the base, sinuate-dentate, glandular-pubescent, to
7 cm. long and 5 cm. wide, the veins below prominently lined with
brown oil cells, the petioles to 6 em. long. Inflorescences erect, elon-
gating in flower, mostly axillary. Hypanthium 2-5 mm. long, 0.5-1.7
mim. wide at the summit, glandular-pubescent without, villous within.
Sepals 1.5-3.5 mm. long, 0.6-1.5 mm. wide, with clusters of oil cells
at tips; caudate appendages absent. Petals 2-6 mm. long, 1-4 mm.
wide, lavender, paler below, often yellow at very base and often with
lavender flecks below. Filaments creamy, those of the episepalous
ones 1.8-3 mm. long, those of the epipetalous ones 1-2.5 mm. long;
anthers 0.6-1 mm. long, glabrous or sparsely ciliate. Style creamy,
villous near the base, 4-7 mm. long, stigma ca. 0.8-1 mm. thick,
surrounded by the anthers at anthesis. Capsule 0.7-1.3 em. long, 1.3-2
mm. thick, erect, clavate, straight, glandular-pubescent; pedicel 2-5
mm. long. Seeds brown, 1-1.2 mm. long. Gametic chromosome number,
n=7. Autogamous.

Type: Candelaria, Mineral County, Nevada, 1881, W. H. Shockley
19 (GH).

Disrripution: Alluvial slopes and rocky slides, often on rocky
soil, Churchill and Lander Counties, Nevada, south to Lincoln and
southern Nye Counties, Nevada, Mono Lake, Mono County, and
central Inyo County, California. From 2,200-7,000 ft. alt.

Camissonia heterochroma is highly colonial, rare, and autogamous.
Although it grows adjacent to C. brevipes subsp. brevipes, C. claviformis
subsp. funerea, C. claviformis subsp. integrior, C. claviformis subsp.
lancifolia, and C. munzii, its time of flowering only rarely coincides
with that of any of these entities. I can no longer see any justification
for separating the relatively glabrous and glaucous populations
within this species (=subsp. monoensis) as a separate entity, particu-
larly in view of their poorly separated geographical ranges; plants
from Lincoln County, Nevada, are more or less intermediate in
pubescence characters.

The following two collections, seen since I completed my revision
of the group in 1962, extend the then known range: Cerro Gordo,
3.8 miles east of Keeler, Inyo Mountains, Inyo County, ea. 5,300 ft.,

RAVEN—GENUS CAMISSONIA 233

California, Thompson 3383 (DS); east side road south of Rainier Mesa,
5.4 miles north of junction of Tippipah Road, East Forty-Mile
Drainage, 5,000 ft., Nye County, Nevada, Beatley 391 (DS).

Section III. Lignothera

Oenothera subg. Chylismia sensu Munz, Amer. Journ. Bot. 16: 224. 1929; N. Amer.
F]. IT. 5: 163. 1965; pro parte.
Oenothera sect. Euchylismia sensu Munz, Amer. Journ. Bot. 16: 224. 1929; pro
arte.
Oenothera sect. Lignothera Raven, Univ. Calif. Publ. Bot. 34: 76. 1962.
Camissonia sect. Lignothera (Raven) Raven, Brittonia 16: 281. 1964.
Suffrutescent perennials, flowering the first year, the plants caules-
cent, not flowering at the basal nodes. Inflorescence nodding. Flowers
opening in the late afternoon. Ovary lacking a sterile projection.
Hypanthium lacking a fleshy disc within. Sepals reflexed separately.
Petals dull yellow or rarely cream, unmarked, fading brick red or
yellowish, with no contrasting ultraviolet-reflective pattern. Stamens
and style yellow, the stigma depressed globose, greenish yellow.
Stamens 8, subequal; anthers versatile, ciliate. Pollen shed in tetrads.
Capsule subsessile or with a prominent, stout pedicel, straight or
slightly curved, subterete, the seeds in 2 rows in each locule. Seeds
monomorphic, narrowly obovoid, finely lacunose.
TypE spEcIES: Camissonia cardiophylla (Torr.) Raven.
Distripution: Alluvial slopes and washes from Inyo County,
California, south through San Diego County, California, and Pinal
County, Arizona, to northwestern Sonora and about 27°50’ N. lat. in
Baja California; Isla de Cedros and islands in the Golfo de California.
In 1962, I regarded this group as an early evolutionary offshoot
within Camissonia (Univ. Calif. Publ. Bot. 34: 67). In my present
view, however, it is more logical to regard the late afternoon opening
of the flowers, pollen shed in tetrads, and perhaps woody habit as
specializations within the tribe Onagreae, and consequently to regard
sect. Lignothera as a specialized offshoot of some species similar to the
relatively generalized ones in sect. Chylismia. The hypanthium in
plants of sect. Lignothera is not approached in length by any other
species of Camissonia, and it now appears likely that the two closely
related species comprising this group are best regarded as primarily
moth-pollinated derivatives in a mainly bee-pollinated (and morning-
opening) group. Hawk-moth pollination is probably important here,
particularly with respect to C. arenaria, in which the hypanthium
may be up to 4 em. long. The relationship of sect. Lignothera to the
remainder of Camissonia might therefore be analogous to the relation-
ship between the hawk-moth-pollinated Gaura mutabilis Cav. and the
rest of the genus Gaura. There seems to be no reason to regard the
shedding of pollen in tetrads as primitive in the tribe Onagreae or in

234 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

the family, and relatively generalized species of sect. Hulobus and
sect. Chylismia are somewhat woody.

Of the 36 plants of sect. Lignothera for which chromosome number
determinations have been made, pairing has been observed in 30.
Two of these had a ring of 4, and one had 2 rings of 4. Thus 10 percent
of the plants examined were chromosomally heterozygous, which is a
relatively high proportion but much lower than that found in sect.
Chylismia.

18. Camissonia cardiophylla (Torr.) Raven, Brittonia 16: 281. 1964.

Plants to 1 m. tall, covered with villous or glandular pubescence-
Leaves reduced upward, ovate to cordate-orbicular, blades to 5.5 cm.
long and 5.5 cm. wide, pubescent, erose-dentate, the teeth tipped with
clusters of brown oil cells, the petioles to 7.5 cm. long. Hypanthium
4.5-14 mm. long, 3-7 mm. across at the summit, villous within,
pubescent without; sepals 3-9 mm. long, 2-3.5 mm. wide. Petals 3-12
mm. long, 2-13 mm. wide, yellow or cream. Filaments 1-3 mm. long;
anthers 2-4 mm. long. Style 8-23 mm. long, stigma 2-3.5 mm. thick
surrounded by or held just above the anthers at anthesis. Capsule
2-5.5 cm. long, 2.5-3.5 mm. thick, straight or slightly curved, as-
cending; pedicel 1-18 mm. long. Seeds 0.5-0.7 mm. long. Gametic
chromosome number, n=7. Self-compatible but often outcrossing.

Distripution: Alluvial slopes and washes from Inyo County,

valifornia, south through San Diego County, California, and Pinal

County, Arizona, to 27°50’N. lat. in Baja California. Also on Isla de
Cedros and on several islands in the Golfo de California, both in Baja
California and in Sonora. From sea level to 4,500 ft. alt. See Univ.
Calif. Publ. Bot. 34: fig. 1. 1962.

18a. Camissonia cardiophylla subsp. cardiophylla

Oenothera cardiophylla Torr., Pac. R. R. Rep. 5: 360. 1858.

Oenothera cardiophylla var. petiolaris M. E. Jones, Proc. Calif. Acad. II, 5:
682. 1895. Lectotype: Rosario Mission, Baja California, 30 April 1886, C.
R. Orcutt 1333 (US 47096; isotypes, F, GH, MO, NY, PH).

Chylismia cardiophylla (Torr.) Small, Bull. Torrey Cl. 23: 193. 1896.

Oenothera cardiophylla var. typica Munz, Amer. Journ. Bot. 15: 226. 1928.

Oenothera cardiophylla subsp. cardiophylla; Raven, Univ. Calif. Publ. Bot. 34:
78. 1962.

Pubescence villous, sometimes glandular, often mixed. Leaves
cordate. Hypanthium 4.5-12 mm. long, villous of more rarely glan-
dular-pubescent without. Petals 3-12 mm. long, 2-12 mm. wide.
Style 8-23 mm. long, often held above the anthers at anthesis. Gametic
chromosome number, n=7. Self-compatible but often outcrossing.

Tyre: Fort Yuma, Yuma County, Arizona, Major G. H. Thomas
(NY).

RAVEN—GENUS CAMISSONIA 235

DistRIBUTION: Rocky walls and sandy alluvial flats with Hyptis
emoryi, Larrea and Ambrosia dumosa, southern San Bernardino
County, California, south to eastern San Diego County, California,
and. Yuma County (and possibly western Pinal County), Arizona,
and south in northeastern and central Baja California, Mexico, to
approximately 27°50’ N. lat.; also on Isla Angel de la Guarda, Isla
San Marcos, Isla San Luis, Isla San Pedro Martir, and Isla San
Esteban in the Golfo de California. Isla San Esteban is the only
locality for this subspecies in Sonora. From sea level to 2,000 ft. alt.
See Univ. Calif. Publ. Bot. 34: fig. 1. 1962.

Progeny of Balls & Everett 22918, grown at Claremont, California,
were self-compatible, as were two individuals from Yaqui Pass, San
Diego County, California, Kyhos 64-6, judging by pollen-tube growth
in selfed individuals, and two from Box Canyon, Riverside County,
California, Kyhos 65-109 (DS). Gametic chromosome counts of n=7
have recently been made in two collections from Mexico: Isla San
Esteban, Golfo de California, Sonora, Moran 13047; Las Trincheras,
28°29’ N. lat., 113°07’ W. long., ca. 430 m., Baja California, Moran
12600. Wiggins 18159 (DS), from 41 miles south of San Felipe, Baja
California, represents a previously unreported station for this taxon.
Camissonia cardiophylla subsp. cardiophylla has been found growing
sympatrically with C. arenaria, with no evidence of hybridization.
18b. Camissonia cardiophylla subsp. robusta (Raven) Raven, Brittonia 16: 281.

1964.
Oenothera cardiophylla subsp. robusta Raven, Univ. Calif. Publ. Bot. 34: 79.
1962.

Entire plant glandular-pubescent and with scattered long, white,
eglandular trichomes. Leaves broadly ovate, often cordate at the
base, the petioles stout. Hypanthium 9-14 mm. long, olandular-
pubescent without. Petals 7-11 mm. long, 9-13 mm. wide. Style
14-20 mm. long, the stigma usually held above the anthers at anthesis.
Gametic chromosome number, n=7. Self-compatible but often
outcrossed.

Type: Rocky canyon slopes, Jail Canyon, west slope of Panamint
Range, 2,500 ft., Inyo County, California, 14 April 1937, P. Train
(US 1737210; isotypes, ARIZ, COLO, DS, NA, OSC, UC).

Disrrisution: Rocky borders of washes and_ hillsides, with
Larrea and Ambrosia dumosa, western and southern margins of Death
Valley, Inyo County, California. From 2,000-4,500 ft. alt. See Univ.
Calif. Publ. Bot. 34: fig. 1. 1962.

Camissonia cardiophylla subsp. robusta is presumably self-com-
patible, as isolated individuals have on two occasions been observed
setting fruit.

236 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

18c. Camissonia cardiophylla subsp. cedrosensis (Greene) Raven, Brittonia 16:
281. 1964.
Oenothera cedrosensis Greene, Bull. Calif. Acad. 1: 187. 1885.
Oenothera cardiophylla subsp. cedrosensis (Greene) Raven, Univ. Calif. Publ.
Bot. 34: 79. 1962.

Mostly annual, the plants entirely glandular-pubescent, with
scattered long, eglandular trichomes. Leaves ovate, acuminate at
the apex, much longer than broad, broadly cuneate or very rarely
slightly cordate at the base. Hypanthium 8-12 mm. long, glandular-
pubescent without. Petals cream or even (?) white, 3-4.5 mm. long,
3-4 mm. wide. Style 10-15 mm. long; stigma surrounded by the
anthers at anthesis. Gametic chromosome number, n=7. Autogamous.

Type: Isla de Cedros, Baja California, 1859, J. A. Veatch (CAS
862).

Distripution: Arroyos and borders of washes, with Pachycormus,
Larrea, Pachycereus, and Bursera, Isla de Cedros west to Bahia de los
Angeles and El Desierto Vizcaffio, Baja California; Isla Tiburén,
Sonora. From near sea level to perhaps 1,000 ft. alt. See Univ. Calif.
Publ. Bot. 34: fig. 1. 1962.

Camissonia cardiophylla subsp. cedrosensis is local and apparently
not common. This entity is undoubtedly more highly self-pollinating
than the other races of this self-compatible species, judging from its
smaller flowers. A plant from Bahia de los Angeles, Baja California,
Moran 12419, had 7 pairs of chromosomes at meiotic metaphase I.

19. Camissonia arenaria (A. Nels.) Raven, Brittonia 16: 281. 1964. FIGuRE 6

Chylisma arenaria A. Nels., Amer. Journ. Bot. 21: 575. 1934.

Oenothera cardiophylla var. splendens Munz & Johnst., Bull. Torrey Cl. 49:
354. 1923. Type: UC 196590, wrth same data as type of Oenothera
cardiophylla var. longituba Jeps.; not located.

Oenothera cardiophylla var. longituba Jeps., Man. Fl. Pl. Calif. 686. 1925.
Type: High among rocks at the foot of the Needles, Colorado R., Mohave
Co., Arizona, 7 March 1910, J. Grinnell (JEPS 2668); see above.

Oenothera arenaria (A. Nels.) Raven, Univ. Calif. Publ. Bot. 34: 76. 1962.

Plants to 1.8 m. tall, villous, occasionally with scattered glandular
trichomes in the inflorescence. Leaves reduced upward, the petioles
to 6 cm. long, the blades to 6 em. long and 6 cm. wide, villous above
and densely so below, ovate, cordate at the base, the apex sometimes
acuminate, coarsely dentate or doubly dentate. Inflorescence less
compact than that of C. cardiophylla. Hypanthium 18-40 mm. long,
5-8 mm. across at the summit, villous without, finely pubescent
within. Sepals 8-15 mm. long, 3-5.5 mm. wide. Petals 8-20 mm. long,
7-28 mm. wide. Filaments 5-9 mm. long; anthers 5-8 mm. long.
Style 30-58 mm. long; stigma 2.5-3 mm. thick, held above the anthers
at anthesis. Capsule 3—-4.4 cm. long, 2.5-3.5 mm. thick; pedicel 2-5 mm.
long. Seeds 0.5-0.7 mm. long. Gametic chromosome number, n=7.
Outcrossed.

RAVEN—GENUS CAMISSONIA 237

Type: Sandy washes in the Fortuna Range (western Gila Moun-
tains), ca. 20 miles east of Yuma, Yuma County, Arizona, 26 February
1930, A. Nelson 10140 (RM 138602).

DistriBuTION: Sandy washes, rocky conglomerate cuts, and granitic
slopes and draws, with Larrea, Prosopis, Carnegiea, and Ambrosia
dumosa, foot of the Needles, Mohave County, Arizona; from the
north end of the Salton Sea, Riverside County, California, southeast-
ward to the Tinajas Atlas Range, Arizona and Sonora. From below
sea level to 1,400 ft. alt. See Univ. Calif. Publ. Bot. 34: fig. 1. 1962.

This species has twice been found growing sympatrically with C.
cardiophylla subsp. cardiophylla, but no intermediates have been
observed.

Section IV. Tetrapteron

Oenothera subg. Heterostemon Nutt., Journ. Acad. Sci. Philad. 7: 22. 1834. Type:

Oenothera heterantha Nutt. = Camissonia subacaulis (Pursh) Raven.

Oenothera subg. Primulopsis Torr. & Gray, Fl. N. Amer. 1: 507. 1840; nom. subs.
Oenothera subg. Tarazia Nutt. ex. Torr. & Gray, Fl. N. Amer. 1: 506. 1840.

Lectotype: Oenothera breviflora Torr. & Grya = Camissonia breviflora (Torr. &

Gray) Raven.

Tarazia (Nutt. ex. Torr. & Gray) Raimann, in Engl. & Prantl, Natiirl. Pflanzen-

fam. III. 7: 216. 1893.

Oenothera sect. Tetrapteron Munz, Amer. Journ. Bot. 16: 247. 1929.
Oenothera sect. Eutarazia Munz, Amer. Journ. Bot. 16: 247. 1929.
Oenothera sect. Heterostemon (Nutt.) Munz, N. Amer. FI. IT. 5: 143. 1965.

Terennials or annuals, the plants acaulescent or subacaulescent with a
crown of leaves and flowers at summit of the taproot. Flowers opening
near sunrise. Ovary with a long, slender, sterile projection below the
hypanthium on which the flower is placed. Hypanthium closed by an
evident fleshy disc. Sepals reflexed separately or in pairs. Petals
yellow, unspotted, strongly ultraviolet reflective, with a large non-
reflective area near the base in C. ovata. Stamens, style, and inside of
the hypanthium yellow, the stigma greenish yellow. Anthers attached
near the base, erect in anthesis. Pollen shed singly. Capsule subsessile,
irregularly loculicidal, the seeds in two rows in each locule, these
rows often much crowded. Seeds monomorphic, with a linear raphe
on one side, variable.

LEcroTyPE SPECIES: Camissonia graciliflora (Hook. & Arn.) Raven
(cf Raven, Brittonia 16: 283. 1964).

DistriputTion: South-central British Columbia and Alberta at
about 51° N. lat. and southwestern Saskatchewan, and from the
region of the Continental Divide westward south to southern Utah,
central Nevada, and throughout California to northwestern Baja
California.

This section of six species consists of four groups very distinct from
one another: C. ovata, C. subacaulis, C. tanacetifolia-C. breviflora,
and ©. graciliflora-C. palmeri. The last two species are annual, the

238 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

others perennial. In view of their habit and clearly specialized capsules,
it is believed that C. graciliflora and C. palmeri were derived from a
perennial stock; but they are not especially similar to any of the
extant perennials. Camissonia ovata and C. tanacetifolia consist largely
or entirely of self-incompatible individuals, whereas C. subacaulis and
C. graciliflora are facultatively outcrossed although self-compatible.
Camissonia breviflora and C. palmeri are rather obviously specialized
autogamous derivatives of species similar to C. tanacetifolia and C.
graciliflora, respectively. Five of the six species in this section are
diploid (n=7) but C. tanacetifolia is primarily tetraploid (n=14),
with a local hexaploid (n=21) race and at least one diploid (n=7)
population. A further discussion of chromosomal evolution in. this
species is included following its taxonomic description (p. 247).

Chromosomal heterozygosity is unknown in natural populations of
this section, for which 173 diploid counts are available. Multivalent
associations in the 25 tetraploid and 4 hexaploid plants of C. tanaceti-
folia examined clearly reflect: autopolyploid pairing, not structural
rearrangement.

20. Camissonia ovata (Nutt. ex Torr. & Gray) Raven, Brittonia 16: 283. 1964.
Fiacures 9, 15
Ocenothera ovata Nutt. ex Torr. & Gray, Fl. N. Amer. 1: 507. 1840. Munz,
Amer. Journ. Bot. 16: 251. 1929; N. Amer. Fl. II. 5: 144. 1965.
Tarazia ovata (Nutt. ex Torr. & Gray) Small, Bull. Torrey Cl. 23: 185. 1896.
Oenothera primuloidea H. Lév., Monogr. Onoth. 65. 1902.

Subsucculent acaulescent perennial from a thick taproot which
often branches at the crown in age. Leaves ovate to very narrowly
elliptic, 3-15 cm. long, 1.6-5 em. wide, more or less densely white-ciliate
with erect trichomes ca. 0.5 mm. long, subentire to shallowly sinuate
or crisped, rarely deeply sinuate, the apex acute to acuminate, the
base attenuate to a narrowly winged petiole 8-150 mm. long, slightly
dilated at the base. Buds erect just before anthesis. Sterile prolonga-
tion of ovary 2.5-18 em. long. Hypanthium 2-3 mm. long, 3.5-5 mm.
across at the summit, sparsely pubescent outside, with short matted
trichomes inside near the base. Sepals 11-19 mm. long, 1.2-4 mm.
wide, sparsely pubescent, reflexed separately. Petals often apiculate,
8-23 mm. long, 5-19 mm. wide. Filaments of the episepalous stamens
3.5-8 mm. long, those of the epipetalous ones 2-6 mm. long; anthers
3-4.8 mm. long. Style 4.5-11 mm. long, short-pubescent near the base;
stigma 0.8-2 mm. in diameter, held slightly above the anthers at
anthesis. Capsule 11-30 mm. long, 3-5 mm. thick, subterete, the walls
thin, much distended by the seeds, rarely with a pedicel up to 0.4
mm. long. Seeds elongate-ovoid, uniform brown, densely and coarsely
papillose, 1.8-2.2 mm. long, 1.2-1.4 mm. thick. Gametic chromosome
number, n=7. Self-incompatible.

RAVEN—GENUS CAMISSONIA 239

CMW)

Ficures 9-20.—Seeds and capsules of Camissonta sect. Tetrapteron: 9-14, Seeds, X 20: 9,
C. ovata (K. Brandegee 127, DS); 10, C. subaculis (Raven 19601, DS); 11, C. tanacetifolia
subsp. tanacetifolia (Thompson 11962, DS); 12, C. breviflora (Cottam et al. 15285, DS);
13, C. graciliflora (Dudley in 1899, DS); 14, C. palmeri (Parish 4861, DS). 15-20, Cap-
sules, from same collections as seeds unless otherwise noted, X 2: 15, C. ovata; 16, C.
subaculis; 17, C. tanacetifolia subsp. tanacetifolia (Applegate 7635, DS); 18, C. breviflora
(Raven 19551, DS); 19, C. graciliflora, entire fruiting plant (Sharsmith 3395, DS);
20, C. palmert.

295-655 O—68——6

240 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Ficure 21.—Western United States, showing ranges of Camissonia sect. Tetrapteron:
A=C. ovata; @=C. subacaulis.

Type: Moist plains in the immediate vicinity of Monterey, Mon-
terey County, Calif., March 1836, T. Nuttall (NY; isotypes, BM, GH).

DistriBuTion (Figure 21): Grassy fields, often in clay soil, not far
from the coast; Umpqua Valley, Douglas County and Josephine
County, Oregon; Humboldt, Mendocino, and Lake Counties south to
the vicinity of Monterey Bay, Monterey County and again south of
the Santa Lucia Mountains in northern San Luis Obispo County,
from about 2 miles north of the mouth of Arroyo de la Cruz south to
Los Osos Valley near Morro Bay, California. From sea level to about
1,500 ft. alt.

RAVEN—GENUS

CAMISSONIA

\

241

Ficures 22-23.—Species of Camissonia sect. Tetrapteron, X 34: 22, Plant of C.

breviflora (Cottam et al. 15825, DS); 23, flower of C. tanacetifolia subsp. tanacetifolia

(Thompson 11953, DS).

242 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Vouchers for chromosome number, n=7 (no structural heterozy-
gosity observed; 10 individuals, 7 populations) :

U.S.: CALIFORNIA: LAKE CO.: 0.8 mi. W. of Lakeport, Breedlove 4567. MONTEREY
co.: Carmel Valley, R18148; 7 mi. SW. of San Juan Bautista on Old San Juan
Grade, R18194. sAN BENITO co.: 1 mi. W. of San Juan Bautista, R18193. SAN

LUIS OBISPO cO.: 1 mi. N. of Cambria, R18140; 1 mi. S. of Cambria, R18747 (4
plants). SAN MATEO CO.: Pigeon Point, Ornduff 4746.

Individuals of Camissonia ovata are self-incompatible (contrary to
the assertion of Katherine Brandegee, Univ. Calif. Publ. Bot. 6: 46.
1914), as judged by the lack of pollen-tube growth in 71 selfed indi-
viduals from a population near the intersection of Sand Hill Road and
Whiskey Hill Road, near Woodside, San Mateo County, California;
7 individuals from the Lake County, California, population cited above,
and 12 plants from the population from south of Cambria, San Luis
Obispo County, California, cited above. I am grateful to Gretchen M.
Mills for these observations on the San Mateo County population.
Camissonia ovata has been found growing sympatrically with C.
gracdiflora. The oligolectic bee Andrena (Diandrena) chalybea (Cress.)
is consistently associated with it throughout its range and usually
effects pollination in the early morning (MacSwain, Raven & Thorp,
MS.). Other species of bees gather pollen from Camissonia ovata and
visit it locally, sometimes in competition with Andrena chalybea.

This species might possibly occur or have occurred in the Sierra
Nevada foothills of central California, judging from a specimen
labeled ‘Roseville, May 1884, M.K.C.” (DS); it is uncertain whether
the record refers to the community of this name in Sacramento
County, California. At any rate, the occurrence of this species east of
the Sacramento Valley has not been confirmed by modern collections.

The chromosome number of this species was earlier reported by
Johansen (Amer. Journ. Bot. 16: 597. 1929; Ann. Bot. 45: 111-124,
pl. 6. 1931), but without any definite indication of locality or voucher
specimen.

Camissonia ovata is a distinctive species without any close relatives.
It might be confused superficially with C. subacaulis, but the struc-
ture of the capsules and seeds in these two species is very different
(figs. 9, 10, 15, 16), as is the marking of the petals when viewed in
ultraviolet light. White-flowered plants are found in a very low
frequency.

21. Camissonia subacaulis (Pursh) Raven, Brittonia 16: 283. 1964.
FiaureEs 10, 16
Jussiaea subacaulis Pursh, Fl. Amer. Sept. 1: 304. 1814.
Oenothera heterantha Nutt., Journ. Acad. Philad. 7: 22.1834. Type: Near
Camas Creek, in or near Jefferson Co., Idaho, 23 June 1833, N. J. Wyeth
(PH; isotypes, BM, NY).

RAVEN—GENUS CAMISSONIA 243

Oenothera heterantha var. tarazacifolia S. Wats., Proc. Amer. Acad. 8: 589.
1873. Type: Near Austin, Lander Co., Nevada, July 1868, S. Watson
(GH).

Tarazia heterantha (Nutt.) Small, Bull. Torrey Cl. 23: 186. 1896.

Tararia heterantha var. tarazxacifolia (S. Wats.) Small, Bull. Torrey Cl.
23: 186. 1896.

Tarazia tarazacifolia (S. Wats.) Heller, Muhlenbergia 1: 1. 1900.

Tarazia subacaulis (Pursh) Rydb., Mem. N.Y. Bot. Gard. 1: 281. 1900.

Oenothera primuloidea H. Lév., Monogr. Onoth. 65. 1902, pro parte; nom.
subs.

Oenothera subacaulis (Pursh) Garrett, Spr. Fl. Wasatch 64. 1911.

Oenothera subacaulis var. tarazacifolia (S. Wats.) Jeps., Man. FI. Pl. Calif.
683. 1925.

Subsucculent acaulescent perennial from a deep, thick taproot
which may branch in age to produce more than a single rosette.
Leaves lanceolate to narrowly elliptic, 2-22 cm. long, 0.7-4.2 cm. wide,
rarely minutely and sparsely strigulose along veins and margins, sub-
entire to sinuate or less commonly deeply and irregularly pinnatifid,
the apex acuminate, the base attenuate to a narrowly winged petiole
1-12 cm. long. Buds erect just before anthesis. Sterile prolongation of
ovary 1.5-8 cm. long. Hypanthium 1.5-3 mm. long, 3-4 mm. across
at summit, subglabrous outside, with short matted trichomes inside
near the base. Sepals 4.1-13 mm. long, 0.8-3 mm. wide, sometimes
very minutely strigulose, reflexed separately. Petals often apiculate,
5-16 mm. long, 5-16 mm. wide. Filaments of the episepalous stamens
1.8-6.5 mm. long, those of the epipetalous ones 0.5-2.5 mm. long;
anthers 0.9-2 mm. long. Style 4-8.5 (-11) mm. long, glabrous or
sparsely pubescent near the base; stigma 1-2 mm. in diameter, usually
surrounded by the longer anthers at anthesis, but very rarely held
above them. Capsule 11-28 mm. long, 5-8 mm. in diameter, 4-angled
with flat walls, these thick and smooth and scarcely distended by the
seeds, rarely with a pedicel up to 1 cm. long, becoming blackened
and persistent on the plants for one or more years after shedding
seeds. Seeds oblong, uniform tan to light brown, 1.3-1.9 mm. long,
0.6-1 mm. thick, coarsely pitted, the pits in ca. 25 parallel rows.
Gametic chromosome number, n=7. Self-compatible but often out-
crossed.

Type: Moist ground, Quamash Flats (Squamash Flats), on Weippe
(Oyipe) Prairie, Shoshone County, Idaho, 14 June 1806, M. Lewis
(PH). See Coues, Proc. Acad. Nat. Sci. Philad. 1898: 302. 1898.
Locality incorrectly given by Pursh as “the banks of the Missouri,” to
which he credited many of the species from the Lewis and Clark Ex-
pedition even when the labels indicated other, more specific data
(letter from F. W. Pennell to P. A. Munz, Feb. 17, 1928, filed at
POM). See also Torrey & Gray, Fl. N. Amer. 1: 507. 1840; Raven,

244 OONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Brittonia 16: 283. 1964, as to the application of this name to this
species.

Distrisution (Figure 21): Wet, often clayey meadows, sometimes
growing in deep grass, from Yakima, Kittitas, Chelan, Lincoln, and
Spokane Counties, Washington, and Philips County, Montana (Lake
Bowdoin, Blankinship in 1903, MONT), south through Sheridan and
Fremont Counties, Wyoming, to Routt County, Colorado; Summit
Duchesne, and Sanpete Counties, Utah; White Pine, northern Nye,
and Douglas Counties, Nevada; and in California east of the Cascade
axis to the vicinity of Bodie, Mono County, and again on the Kern
Plateau, Tulare County; 1,500-8,600 ft. alt.

Vouchers for chromosome number, n=7 (no structural hetero-
zygosity observed; 4 individuals, 4 populations):

U.S.: CALIFORNIA: MONO co.: 4.4 mi. NW. of Bridgeport, R18548. Ipano:
ADAMS CO.: 3 mi. W. of Big Payette Lake, R18508. vALLEY co.: 2 mi. S. of McCall,
R18510. OREGON: LAKE co.: Hart Mountain Antelope Refuge Headquarters,
R18430.

Individuals of the two populations of Camissonia subacaulis tested
by pollen-tube growth have been found to be self-compatible and
facultatively self-pollinating. In the first Idaho population mentioned
above 16 individuals were tested, and 17 were tested in the California
population. Pollinators were extremely rare at both localities, and the
flowers were more or less hidden from view at the base of the rosette.
In a few populations (such as Hitchcock & Muhlick 14048, Ds,
POM, Valley County, Idaho) the stigmas seem to be held above the
anthers at anthesis, and some of the populations of. this species
might prove to consist of self-incompatible individuals. Camissonia
subacaulis has been found growing sympatrically with C. tanacetifolia
subsp. tanacetifolia and very probably occurs sympatrically with C.
breviflora as well. This species appears to have no near relatives.
An earlier approximate report of the chromosome number of. this
species was made by Johansen (Amer. Journ. Bot. 16: 597: 1929),
but without any citation of voucher or locality.

22. Camissonia tanacetifolia (Torr. & Gray) Raven, Brittonia 16: 283. 1964.

Acaulescent perennial from a woody, deep taproot from which
numerous slender branches arise in age to produce additional rosettes.
Leaves very narrowly elliptic, 6.5-32 em. long, 0.7-3.3 em. wide,
more or less densely short-pilose, or the trichomes appressed; deeply
and irregularly pinnatifid, the apex acute to long acuminate, the base
attenuate to narrowly cuneate to a petiole 1-8 em. long. Buds erect
just before anthesis. Sterile prolongation of ovary 1.4-5.5 em. long.
Hypanthium 4—6.5(-8.5) mm. long, d-7(-11) mm. across at summit,
short-pilose outside, soft-pilose in lower half within. Sepals 5.5-13
mm. long, 1.8—4.2 mm. wide, short-pilose, reflexed separately. Petals

RAVEN—GENUS CAMISSONIA 245

(8—) 10-23 mm. long, 7-15(-18) mm. wide. Filaments of the episepalous
stamens 5.5-12 mm. long, those of the epipetalous ones 2.5-8 mm.
long; anthers (2.3—-) 2.8-3.5 mm. long. Style 9.5-20(-25) mm. long,
pilose near the base; stigma depressed-globose, more or less 4-notched,
1.3-3.5 mm. in diameter, held well above the anthers at anthesis.
Capsule 7-25 mm. long, 3-5 mm. thick, gradually attenuate to the
sterile prolongation, subterete, the walls thick but evidently distended
by the seeds, sessile, disintegrating irregularly the season after their
formation. Seeds oblong, uniform tan to brown, 1.5—-2 mm. long, 0.6-—
0.8 mm. thick, pitted in rows, with ca. 20 rows of pits. Gametic
chromosome numbers, n=7, 14, 21. Self-incompatible, probably
with some self-compatible populations.

Distrisution (Figure 24): Open clayey fields and moist slopes,
sometimes by streams or lakes, Klickitat, Douglas, and Spokane
Counties, Washington, and Owyhee, Ada, Elmore, Camas, and
Blaine Counties, Idaho, south on the east side of the Sierra-Cascade
axis to Elko, northernmost Nye, and Douglas Counties, Nevada, and
the vicinity of Bodie and Conway Summit, Mono County, California.
From about 1,300 ft. alt. in the north to 8,200 ft. in the south. Gre-
garious and often coloring large fields yellow when in bloom.
22a. Camissonia tanacetifolia subsp. tanacetifolia. Ficures 11, 17, 23

Oenothera nuttallii Torr. & Gray, Fl. N. Amer. 1: 506. 1840; not Sweet
1830. Type: Said to be from “Plains in the Rocky Mountains, near Black-
foot River,’ which would put it in or near Caribou County, Idaho, 10-11
July 1834, T. Nuttall (isotype PH), at the type locality for the following
species and far from any known locality for the present one; therefore, the
locality as given is probably in error.

Oenothera tanacetifolia Torr. & Gray, Pac. R. R. Rept. 2: 121, pl. 4. 1854.

Tarazia longiflora Nutt. ex Howell, Fl. N.W. Amer. 231. 1891-1896.

Tararia tanacetifolia (Torr. & Gray) Piper, Contr. U.S. Nat. Herb. 11: 405.
1906.

Taraxia tikurana A. Nels., Bot. Gaz. 54: 140. 1912. Lectotype: Tikura,
Blaine Co., Idaho, 4,500 ft. elev., 22 July 1911, A. Nelson & J. F. Macbride
1302 (RM; isolectotypes, DS, GM, NMC, POM, US.)

Plants more or less densely pilose or the trichomes appressed;
rarely more than 5 percent of the pollen with 4 pores. Gametic chro-
mosome numbers, n=7, 14. Self-incompatible, probably with self-
compatible populations.

Type: Vicinity of Pah-Rum Peak, Lake Range, Washoe County,
Nevada, 18 June 1854, E. G. Beckwith 43 (GH, the specimen from
which the illustration was prepared; isotype, NY). Type locality
given as “On the higher parts of the Sierra Nevada” but the actual
locality was reconstructed from Beckwith’s account of the expedition
(Pac. R.R. Rept. 2(1b): 1-70. 1854).

Disrrisution (Figure 24): That of the species, but not found in
and near Sierra Valley in Plumas and immediately adjacent portions
of Lassen and Sierra Counties, California.

246 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Figure 24.—Western United States and a portion of adjacent Canada, showing ranges of
species of Camissonia sect. Tetrapteron: O=C. tanacetifolia subsp. tanacetifolia, with
@= tetraploid chromosome counts (n=14) and @=the single diploid count (n=7);
A=C. tanacetifolia subsp. quadriperforata (all hexaploid, n==21); X = C. breviflora.

Voucher for chromosome number, n=7 (1 structurally homozygous
individual) :

U.5.: OREGON: HARNEY co.: 0.6 mi. N. of Wagontire, R18432.

Vouchers for chromosome number, n= 14 (with variable numbers
of rings of 4 chromosomes in meiotic metaphase I, the maximum
number of such rings observed in each case indicated in parentheses

RAVEN—GENUS CAMISSONIA 247

following the collection number; other chromosomes present as _bi-
valents; 25 individuals, 25 populations) :

U.S.: CALIFORNIA: LASSEN co.: U.S. Hwy. 395, 11.1 mi. N. of Litchfield,
R18423 (3); 0.9 mi. N. of Ravendale, R18426; 11.6 mi. N. of Litchfield, R17875
(3); 0.56 mi. N. of Ravendale, R17876 (5). Mopoc co.: 13 mi. NW. of Canby,
R17893; State Hwy. 139 at S. road to Lava Beds National Monument, R17896
(4). pLuMas co.: Middle Fork of Feather R. just N. of Mohawk, Mosquin &
Gillett 5302 (3). stiskryou co.: Grass Lake, Bates 2436; 14 mi. 8S. of Macdoel,
R17900; 4.4 mi. 8. of Dorris, R17898. IpaHo: camMas co.: Near Camas Creek
and State Hwy. 46, R18526 (5). ELMORE co.: 2.7 mi. N. of Mountain Home,
R18523. OWYHEE co.: U.S. Hwy. 395, 9.3 mi. NE. of the Oregon line R19576.
Nevapa: pouatas co.: U.S. Hwy. 395, 2.9 mi. N. of junction with State Hwy.
3, R18546; S. end of Double Spring Flat, R14267. wasHor co.: 20 mi. E. of
Cedarville (Calif.) on State Hwy. 8A, R17885 (3). OREGON: crooK co.: N. road
from Prineville to Big Summit Prairie, Chambers & Lewis 1934. HARNEY CO.:
U.S. Hwy. 20, 7.4 mi. E. of junction with U.S. Hwy. 395, R18457; 8.4 mi. N. of
Narrows, R18456; 23.2 mi. S. of Narrows, R18454 (4); U.S. Hwy. 20, 2.1 mi.
W. of junction to Ft. Harney, T. & L. Mosquin 4345 (DAO). LAKE co.: 0.9 mi.
E. of Hart Mountain Antelope Refuge Headquarters, R18341 (5); 4.1 mi. E. of
Drew’s Reservoir, W. of Lakeview, R18428. MALHEUR co.: 6 mi. N. of Jordan
Valley, R19583. WASHINGTON: ADAMS CoO.: 4 mi. NE. of Ritzville, R18496.

Twelve individuals from the Washington population just cited
appeared to be self-compatible, judging from pollen-tube growth in
selfed, caged individuals. On the other hand, 25 plants of the first-
mentioned Lake County, Oregon, population proved to be self
incompatible, as did 17 individuals from the Douglas County, Nevada,
population. Andrena (Onagrandrena) raveni Linsley & MacSwain is
oligolectic on plants of the tribe Onagreae, and females of this species
have been observed abundantly gathering pollen on Camissonia tan-
acetifolia subsp. tanacetifolva in a number of populations. Outcrossing
is certainly the mode in Camissonia tanacetifolia, and the existence
and distribution of self-compatible plants needs to be studied in
greater detail.

From a chromosomal point of view, 25 of the 26 populations of
this subspecies examined have been effectively autotetraploid, usually
with 3 rings of 4 chromosomes in each diakinesis or metaphase I
cell, but occasionally with up to 5 rings of 4. It is not known whether
the diploid set of chromosomes duplicated twice in these plants is
structurally homologous with that in the related diploid C. breviflora,
but that species is autogamous and small-flowered, and cannot be
identical with the ancestor of the self-incompatible, large-flowered,
polyploid C. tanacetifolia. No other related diploid is known.

The single diploid strain of C. tanacetifolia subsp. tancetifolia was
found during the course of an extensive survey of chromosome num-
bers throughout the range of this taxon (fig. 24). It is morphologi-
cally identical to the tetraploid plants that occur in the same area,
and it could therefore be their diploid ancestor. In my opinion, how-

248 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

ever, it is at least equally probable that this single diploid population
(perhaps no more than a single plant) is a polyhaploid derivative of
the predominant tetraploid. The chromosomes of the tetraploid are
like those of other tetraploids that have given rise occasionally to
fertile polyhaploid (=diploid) derivatives (Raven & Thompson,
Amer. Nat. 98: 251-252. 1964). Meiotic accidents that could lead
to the formation of such a diploid derivative are known to occur
sporadically in plants of this sort. I can think of no way to distin-
guish these two competing hypotheses concerning the origin of this
diploid plant or population at present.

Camissonia tanacetifolia subsp. tanacetifolia has been found grow-
ing sympatrically with C. subacaulis. It is closely related to the au-
togamous, diploid C. breviflora but sharply distinct in flower size.

22b. Camissonia tanacetifolia subsp. quadriperforata Raven, subsp. nov.

A subsp. tanacetifolia differt: foliis villose incanis; plus quam 10
percentum pollinis grana plerumque quadripora; chromosomatum
numero gametico, n=21.

Tyre: Common on flats with Artemisia, 6.2 miles north of Sier-
raville on road to Loyalton, Sierra County, California, 28 May 1963,
P. H. Raven 18416 (DS 502795; isotypes RSA, Us).

DistriBuTION (Figure 24): Common on more or less open clay flats,
often with Artemisia, in Sierra Valley, Plumas County, and imme-
diately adjacent portions of Lassen and Sierra Counties, California;
4,300—5,100 ft. alt.

Vouchers for chromosome number, n=21 (with many rings of 6
at diakinesis and meiotic metaphase I; 4 individuals, 4 populations):

U.S.: CALIFORNIA: PLUMAS Co.: The type collection, cited above; 3.1 mi. W.
of Vinton, R18419; 6.9 mi. S. of Vinton, R18417: ca. 2.5 mi. SE. of Beckwourth,
ca. 5,000 ft., Ornduff 4347 (LA). stprRA co.: State Hwy. 49, 2.4 mi. WNW. of
Sierraville, R18415.

ADDITIONAL SPECIMENS EXAMINED:

U.S.: CALIFORNIA: LASSEN CO.: Long Valley, sec. 34, T. 45 N., R. 17 E., 4,300
ft., Sawyer 219 (RSA, UC): Chat, 5,000 ft., Hillman in 1897 (POM), Jones in
1897 (POM): 3 mi. N. of Omira, Howell 11857 (CAS). pLUMAs co.: Sierra Valley,
Bolander & Keller in 1872 (NY), Lemmon 87 (GH), in 1873 (GH), Hillman in
1898 (POM); Portola, Eastwood 7051 (CAS); 1.7 mi. NE. Vinton, 5,000 ft.,
Cantelow in 1941 (CAS, RSA); 8 mi. N. of Chileoot, 5,000 ft., Munz 11823 (COLO,
NY, RSA, RM, SD, UT, WTU); Chilcoot, 4,995 ft., Rose 34411 (CAS, RM);
434 mi. SE. of Sugarloaf, 4,900 ft., Sawyer 90 (POM, RSA); along road to Beck-
wourth, from Calpine, Sierra Valley, ca. 4,800 ft., Bacigalupi 4282 (JEPS, RSA):
Beckwith Pass, 4,800 ft., Jepson 7767 (DS, JEPS): near Beckwourth, 4,900 ft.,
Howell 30753 (CAS), 37769 (CAS). sterra co.: Loyalton, Clemens in 1920 (CAS).
WITHOUT DEFINITE LOCALITY: Lemmon in 1875 (POM), 104 (NY).

Camissonia tanacetifolia subsp. quadriperforata is narrowly dis-
tributed and can be recognized most easily by a characteristic directly

RAVEN—GENUS CAMISSONIA 249

related to its hexaploid chromosome number: namely, the relatively
high proportion of 4-pored pollen grains. Plants of this taxon are
essentially identical in all other respects to those of subsp. tanacetifolra
found in neighboring areas, although they do differ morphologically
from a majority of the populations of subsp. tanacetifolia found in
other areas. There is no known diploid population that could have
combined with the tetraploid subsp. tanacetifolia to give rise to this
hexaploid. In view of this and the local distribution of subsp. guadriper-
forata, its lack of morphological distinctness from adjacent tetraploid
populations of the species, and the behavior of its chromosomes in
meiosis, it seems likely that this hexaploid arose directly from the
adjacent tetraploid populations by the functioning of an unreduced
gamete. Probably the hexaploid originated recently and only once
and has since spread to occupy its present area, which is less than 25
miles across at its broadest point. The habitat occupied is uniform
and continuous. In order to understand how this postulated series of
events could have occurred, however, it is necessary to assume that
the hexaploid is physiologically superior to the tetraploid in its area,
for it has apparently replaced the tetraploid where it does occur.

The nearest stations of hexaploid and tetraploid plants appear to be
approximately 9 miles apart at present, with subsp. tanacetifolia at
Graeagle (Mason 1037, DS, POM, UC) and subsp. quadriperforata
to the east at Portola. Both of these stations are on the Middle Fork
of the Feather River in Plumas County, California.

23. Camissonia breviflora (Torr. & Gray) Raven, Brittonia 16: 283. 1964.
Ficures 12, 18, 22
Oenothera breviflora Torr. & Gray, Fl. N. Amer. 1: 506. 1840.
Tarazia breviflora (Torr. & Gray) Nutt. ex Small, Bull. Torrey Cl. 23: 185.
1896.

Similar to C. tanacetifolia. Plants densely clothed with appressed
long trichomes. Leaves 1.7-10 cm. long, 0.35-3.3 cm. wide, the petiole
1.5-3 cm. long. Sterile prolongation of ovary 0.4-1.5 em. long. Hypan-
thium 1.8-2 (-2.5) mm. long, 2.3-4 mm. across at the summit. Sepals
3.8-5 (-7.5) mm. long, 0.9-2.2 mm. wide. Petals 5.5-7 (—8) mm. long.
2.8-5 (-6) mm. wide. Filaments of the episepalous stamens 2.8-4 (—5)
mm. long, those of the epipetalous ones 1.5-1.8 (—2) mm. long;
anthers 0.9-1.3 mm. long. Style 3-6.5 mm. long; stigma globose, 1.1-
1.6 mm. in diameter, surrounded by and receiving pollen directly
from the anthers of the longer stamens at anthesis. Capsule 8-17
mm. long, 2-5 mm. thick. Seeds 1.5-1.8 mm. long, 0.7-0.9 mm.
thick. Gametic chromosome number, n=7. Autogamous.

Tyre: Near the Blackfoot River, in or near Caribou County,
Idaho, 10-11 July 1834, 7. Nuttall (GH; isotypes, BM, K, NY, PH).

Distripution (Figure 24): Moist, often muddy or sandy flats,
sometimes on lakeshores or in the dried beds of ponds in the vicinity

250 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

of Chilko Lake (Nemiah Trail, 4,500 ft., Copley 440, DAO), at ca.
51° N., 124° W. in central British Columbia; from the region of
Drumheller, Alberta (Handhills Lake, 51°30’ N., 112°07’ W., Macoun
144, BM, GH, NY), and Loomis, southwestern Saskatchewan
(Hudson 1966, DAO), south to Montana (Glacier, Cascade-Meagher,
Park, and Madison Counties), Wyoming (Yellowstone National
Park, Teton, Sublette, Sheridan, Sweetwater, and Lincoln Counties),
Colorado (Larimer County: North Park, Osterhout 1036, RM),
Idaho (Caribou, Bear Lake, Franklin, Custer, and Cassia Counties),
and Utah (Summit, Wasatch, Sevier, Piute, and Garfield Counties,
reaching its southern limit on the Aquarius Plateau); from 3,500 ft.
(at its northern limits) to 10,000 ft. elevation, apparently always above
6,500 ft. in the United States, In south-central Idaho, populations
of C. breviflora are found within 65 miles of populations of the related
C. tanacetifolia, but the former is found in mountain meadows at
approximately 8,000 ft. elevation, the latter on more or less alkaline
flats from 3,000-4,500 ft. elevation. Records of C. breviflora from
south and west of Cassia County, Idaho, are based on misidentifi-
cations of sometimes stunted specimens of C. tanacetifolia. One such
erroneous record is that of Munz (Calif. Fl. 952. 1959), based on
Wheeler 3744 (POM, NY, US) from the Warner Mountains of Modoc
County, California; Camissonia breviflora is not known from any
locality in or near California.

Voucher for chromosome number, n=7 (1 structurally homozygous
individual):

U.S.: WYOMING: SWEETWATER CO.: Eden Valley Reservoir, ca. 6,600 ft., R19551.

Plants of the collection just cited were grown at Stanford for
several years and found to be autogamous, shedding pollen directly
on the stigma and setting full capsules even when protected from
insects. It is presumed that Camissonia breviflora, which is relatively
uniform morphologically, is an autogamous derivative of the diploid
ancestor of C. tanacetifolia. These two species have been confused
in the past, but are sharply distinct both morphologically and eco-
geographically. Camissonia breviflora almost certainly occurs sym-
patrically with C. subacaulis.

24. Camissonia graciliflora (Hook. & Arn.) Raven, Brittonia 16: 283. 1964.
Oenothera graciliflora Hook. & Arn., Bot. Beechey Voy. 341. 1838.
Tarazia graciliflora (Hook. & Arn.) Raimann, in Engl. & Prantl, Natiirl.
Pflanzenfam. III. 7: 216. 1893.

Annual, acaulescent or less commonly with short ascending lateral
branches up to 2.5 cm. long. Leaves linear to very narrowly lanceolate,
1-9.8 cm. long, 0.1-0.9 cm. wide, more or less densely pilose, entire or
very sparsely serrulate, sessile, dilated at the base, the margins of the
basal portion membranous. Buds nodding before anthesis but flowers

RAVEN—GENUS CAMISSONIA 251

erect. Sterile prolongation of ovary 0.6-4.5 cm. long. Hypanthium
1.6-3.2 mm. long, 1.8-5 mm. across at the summit, pilose outside, more
sparsely so inside, especially near the base. Sepals 4.5-8 mm. long,
1.2-2.9 mm. wide, pilose, reflexed in pairs. Petals 5-18 mm. long, 4.8-13
mm. wide. Filaments of the episepalous stamens 1.8-3.2 mm. long,
those of the epipetalous ones 0.8-1.6 mm. long. Style 3-5.5 mm. long,
short-pubescent near the base; stigma 1—1.6 mm. in diameter, sur-
rounded by the anthers of the longer stamens at anthesis. Capsule 4-8
mm. long, 2.6—4.8 mm thick, sharply 4-angled and with a pointed wing
near the center top of each valve, thick-walled (almost woody) and
very tardily dehiscent, splitting for only about a third of its height.
Seeds obovoid, tan mottled with brown, finely papillose, 1.2-2 mm.
long, 1-1.2 mm. thick. Gametic chromosome number, n=7.

Type: California, 1833, ). Douglas (K; isotypes, BM, GH, NY).

DistriBuTION (Figure 25): Colonial on open or brushy slopes, often
in clayey soil, from near sea level to about 2,500 ft. elevation; from
Josephine County, Oregon, south in the Coast Ranges and foothills of
the Sierra Nevada to Ventura and Los Angeles Counties, California;
San Ysabel, San Diego County, Henshaw in 1893 (NY); red clay soil
in once-cultivated field, Rancho Aguajito, 300 m., 29°59’N., 115°15’W.,
Baja California, Raven et al. 12675 (UC; the only known occurrence of
the section in Mexico). Usually away from the coast, but collected
both in Marin County (Mt. Tamalpais (Wood in 1915, CAS) and in
San Francisco (Cannon in 1896, CAS), California. Once collected as
an introduced plant in a dry, weedy field, St. George, Rockland
County, New York, 20 July 1933, Moldenke 8008 (NY).

Voucher for chromosome number, n=7 (1 structurally homozygous
individual) :

U.S.: Catirornia: Monterey Co.: Jolon, Breedlove 4279.

Camissonia graciliflora has been collected growing sympatrically
with the very closely related C. palmeri (1 mile west of Pearblossom,
Los Angeles County, California, Mosquin 3264, DAO; 3260, C.
palmeri) and with C. ovata. It is self-compatible and facultatively self-
pollinated, as shown by pollen-tube growth in three plants of R18153,
Monterey County, California, that were caged in the field, and eight
plants of R18177, San Luis Obispo County, California, that were
grown in the experimental garden at Stanford. This species is probably
not regularly outecrossed, judging from observations made at Jolon,
Monterey County, California, on 31 March 1963 (Raven, MS.). With
C. palmeri it forms a very distinctive group in the section, marked by
annual habit, distinctive capsule morphology, obovoid seeds, nodding
buds, and sepals reflexing in pairs; nonetheless, these two species so
closely resemble the perennial members of the section in details of
floral morphology (such as the sterile projection on the ovary, and the

252 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Ficure 25.—Portion of the western United States and adjacent Mexico, showing the
ranges of species of Camissonia sect. Tetrapteron: A=C. graciliflora; @=C. palmeri.

fleshy ring closing the bottom of the hypanthium), that I believe they
are best kept together in a single section.
25. Camissonia palmeri (S. Wats.) Raven, Brittonia 16: 283. 1964. FicureEs 14, 20

Oenothera palmeri 8. Wats., Proc. Amer. Acad. 12: 251. 1877.
Tarazia palmeri (S. Wats.) Small, Bull. Torrey Cl. 23: 184. 1896.

Similar to C. graciliflora. Branches swollen, to 5 mm. thick, with
conspicuously exfoliating white rhytidome. Leaves narrowly oblanceo-
late, 1.5-5.5 cm. long, 0.18-0.6 cm. wide, strigose, rarely with a few
spreading trichomes near the base, sparsely but evenly serrulate.

RAVEN—GENUS CAMISSONIA 253

Sterile projection of ovary 0.55-1.2 cm. long. Hypanthium 0.8-1.3 mm.
long, 0.8-1.2 mm. across at the summit, strigose outside, glabrous
within. Sepals 1.6-2.3 mm. long, ca. 1 mm. wide, strigose. Petals
2-3.5 mm. long, 1.8-4 mm. wide. Filaments of the episepalous stamens
0.8-1 mm. long, those of the epipetalous ones ca. 0.2 mm. long.
Style 1-2.2 mm. long, glabrous; stigma 0.3-0.6 mm. in diameter,
surrounded by the anthers of the long and short stamens, both of which
shed pollen directly onto it at anthesis. Capsule 5-7 mm. long, 4.5-7
mm. thick, with much more prominent points than in C. gracilzflora.
Seeds narrowly obovoid. Gametic chromosome number, n=7.
Autogamous.

Type: Arizona, 1876, EL. Palmer 597 (GH), perhaps from the
Colorado River valley. The only known collection from this State,
where the occurrence of the species needs to be confirmed; cf. McVaugh
& Kearney, Amer. Midl. Nat. 29: 775-778. 1943.

DistrisuTion (Figure 25): Deserty flats, often under sagebrush,
2,000-4,650 ft. elevation, along the Malheur River near Vale, Malheur
County, Oregon; 20 miles north of Winnemucca, Humboldt County.
(Ripley & Barneby 4568, NY) and Empire City, Ormsby County.
(M. E. Jones in 1882, GH, POM, UC), Nevada; in California fairly
common from southern Inyo County, to the southwestern borders
of the Mojave Desert, west to the vicinity of Tejon Pass and south-
eastern San Luis Obispo County in the inner South Coast Ranges,
and also east of Jacumba on road to Mountain Springs, San Diego
County (Eastwood 9553, CAS). May also occur in Arizona (cf. the
type collection).

Voucher for chromosome count, n=7 (1 structurally homozygous
individual) :

U.S.: CALIFORNIA: SAN LUIS OBISPO co.: Temblor Grade, Elkhorn Valley
Road, south of Crocker Grade, Lewis & Thompson 1635 (LA, RSA).

As noted under that species, Camissonia palmeri has been collected
growing sympatrically with C. graciliflora. Its floral structure provides
unequivocal evidence of its autogamy. Indeed, some plants of both
C. palmeri and C. graciliflora become cleistogamous under certain
circumstances, these conditions being poorly understood. Such cleis-
togamous individuals, or other plants lacking flowers, can be deter-
mined only with difficulty. Plants of both species remain rooted in the
eround long after they have died, the capsules shedding the seeds
slowly after the first autumn rains.

Section V. Holostigma

Agassizia Spach, Hist. Vég. Phan. 4: 347. 1835; not Chavannes 1830.
Holostigma Spach, Nouv. Ann. Mus. Paris IIT. 4: 332. 1835.

Oenothera subg. Sphaerostigma sensu Munz, Bot. Gaz. 85: 234. 1928; pro parte.
Oenothera sect. Holostigma (Spach) Munz, N. Amer. Fl. II. 5: 159. 1965.

254 OONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Perennials or annuals, the plants caulescent but with a well-devel-
oped basal rosette, flowering from the base. Inflorescence nodding at
anthesis, becoming erect in fruit. Flowers opening near sunrise. Ovary
lacking a sterile projection. Sepals reflexed separately or in pairs.
Petals yellow, often with one to several red dots near the base, fading
reddish, with no contrasting pattern visible in ultraviolet light.
Stamens, style, and inside of hypanthium yellow, the stigma yellow
or greenish yellow. Capsule regularly loculicidal, often contorted, not
torulose, with a prominent brown rib down the center of each valve,
the seeds in one row in each locule. Seeds with a scar at the micropylar
end and a poorly developed pellucid chalazal crest, dull, brownish
black, minutely lacunose in rows, narrowly obovoid.

TYPE SPECIES: Camissonia cheiranthifolia (Hornem. ex Spreng.)
Raimann.

DistrisuTion: Brushy or open slopes and flats throughout. cis-
montane California and the northern half of Baja California, with
two species ranging across the deserts from southern Inyo County,
California, and Esmeralda County, Nevada, across southern Arizona
to the vicinity of Tucson, and one species ranging north along the
coast to Coos Bay, Curry County, Oregon. Not known from, but
possibly to be expected in, Sonora.

In all species of sect. Holostigma, the leaves of the basal rosette
and the first formed capsules are more densely pubescent than those
formed later. The leaves of the basal rosette tend to be broadest near
the middle, and those of the stems and inflorescence tend to be
broadest near the base and often subsessile or with shorter petioles.
The first formed flowers are small, the largest ones are formed later
and decrease markedly in size as the flowering season comes to an
end. For all of these reasons it is especially necessary to deal with
comparable stages when evaluating material of some of the critical
species in this section.

The group of species comprising sect. Holostigma has always been
a particularly difficult one taxonomically, both for the reasons just
mentioned and because of their reticulate, polyploid pattern of evolu-
tion (see Appendix). In the treatment presented here, I have recognized
14 species, 3 of which are divided into 2 subspecies each. Of these 14
species, 9, including all of the polytypic ones, are diploid; 2 are tetra-
ploid; and 3 are hexaploid.

All 9 diploids are relatively distinct from one another, although
there is some intergradation between them. Both self-incompatible
entities in the section are diploid: the ditypic, perennial C. cheiranthi-
folia, a species of the coastal strand from Coos Bay, Oregon, south to
northern Baja California (partly consisting of self-compatible indi-
viduals), and the mostly annual C. bistorta that intergrades broadly

RAVEN—GENUS CAMISSONIA 255

with C. cheiranthifolia along the coast of southern California. Of the
remaining seven diploids, C. guadalupensis of San Clemente and
tuadalupe Islands, C. ignota of southern California and scattered
localities northward, and C. proavita of coastal northern Baja Cali-
fornia are extremely distinct and do not appear to have contributed
to the origin of any of the polyploid species.

The difficulty lies with the remaining four diploid species and
their polyploid derivatives. Camissonia lewisti, found along the coast
of southern California and the northern half of Baja California, may
have been derived more or less directly from coastal populations of
(. bistorta. In northern and central California and also extending into
the main area of the group in southern California, C. micrantha is
largely a species of sandy beaches, washes, and flats at relatively low
elevations, whereas (. hirtella is a species of cleared brushy slopes and
chaparral margins away from the coast. Although these two diploids
are modally extremely distinct, they have apparently combined to
produce a tetraploid, C. intermedia, which bridges the morphological
and ecological gap between them and contributes materially to the
taxonomic difficulties of the group. Camissonia pallida, a diploid with
a very wide range in the deserts of southern California and adjacent
Nevada, western and southern Arizona, and northeastern Baja
California, is very closely related to C. hirtella and at times difficult
to distinguish from it, although the species come into contact only at
the margins of their ranges. Given this pattern of variation, it is not
at all surprising that C. confusa, the probable allotetraploid between
Camissonia hirtella and C. pallida, should compound the taxonomic
problems of separating its two diploid ancestors and be difficult to
recognize when compared with certain populations of the diploids.

Although Camissonia confusa, the rarer of the two tetraploids, does
not appear to have contributed to the origin of any hexaploid species,
©. intermedia has rather obviously done so. In Monterey and San
Luis Obispo Counties, California, north of the main area of the
section, C. intermedia appears to have combined with each of its
parents, C. hirtella and C. micrantha, to have given rise to the local
hexaploids C. luciae and C. hardhamiae, respectively. In Baja Cali-
fornia, C. intermedia has evidently combined with C. lewisii to give
rise to the hexaploid C. robusta.

With a reticulate pattern of variation of this sort, it is no wonder
that taxonomic treatments accorded the group have varied widely,
particularly in the absence of knowledge about chromosome number.
Although there has been some variation in the number of species and
varieties recognized, the perennial C. cheiranthifolia and the annual,
self-incompatible C. bistorta generally have been recognized as dis-
tinct from the remainder of the group by virtue of their large flowers

295+-650 O—68——7

256 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

and long styles. The same is largely true of C. guadalupensis, described
in 1876 but excessively rare in collections until the past five years.
Most of the difficulty then has concerned the remaining species of
the section, usually treated collectively as Oenothera micrantha.

How have different authors treated the Oenothera micrantha com-
plex? E. L. Greene in 1891 subdivided this group into the more or
less prostrate or trailing coastal Oenothera micrantha, which he allied
to Oenothera bistorta of southern California, and the erect plant of the
interior with broader leaves that he called Oenothera hirtella. Greene
was the first author who had seen many populations in nature to
deal with this group. In 1905, LeRoy Abrams realized the distinctive-
ness of the densely strigose desert populations, which he described
as Oenothera pallida. Later the same year Aven Nelson, working
largely with herbarium material, treated some material of both species
that had been described as varieties of his Sphaerostigma micranthum,
but also recognized Sphaerostigma hirtellum and described another
species based on material of the same diploid taxon as S. arenicola.
Nelson was evidently puzzled by the striking changes in plants of
this group that occur with increasing maturity and had no acquaint-
ance with them in the field; his work cannot be regarded as a forward
step.

In 1907 appeared what were clearly the most perceptive remarks

on the plants of this section written without knowledge of chromosome
numbers. This paper was entitled ‘Notes on Sphaerostigma,” by
Anstruther Davidson, an enthusiastic amateur botanist of southern
California. From his excellent vantage point Davidson was able to
present a remarkably accurate picture of the entities represented
in the group. He recognized as species S. pallidum, S. hirtellum (“a
good and quite distinct species, and not a merely hirsute variety of
micranthum as some have supposed”), and S. micranthum, and also
described S. hallii, here regarded as a subspecies of Camissonia pallida.
Davidson’s remarks about the pattern of variation in his S. micran-
thum are remarkable and deserve to be quoted at some length (Muhlen-
bergia 3: 107. 1907):
The typical plant of this species is, I presume, the ordinary moderately hirsute,
semi-prostrate form found from San Francisco southward. In the coast region
round Los Angeles there are two well marked forms. One, the congesta form, has
very congested foliage, with the leaves larger and more hirsute, and with very
small flowers; the other, the macrocarpa form, with the same foliage, has the short
stout capsule of bistorta. The latter is the rarer form, examples of which are found
from Los Angeles, San Diego, and Lower Califormia. A typical interior form is
found in Parish’s collections from San Bernardino county, and is frequently met
with in other interior valleys. It is to the eye quite smooth in all its parts.

In retrospect, it is obvious that Davidson was quite correct about
the typical form of the species, which is here recognized as Camissonia

RAVEN—GENUS CAMISSONIA 257

micrantha; this also includes his ‘“‘congesta” form. Davidson’s ‘‘macro-
carpa” form is obviously the species described 60 years later in the
present revision as C. lewisii; and his ‘typical interior form” is the
plant described by Jepson in 1925 as a variety of Oenothera micrantha
and here regarded as a distinct species Camissonia ignota. In other
words, Davidson recognized and quite accurately described all seven
diploids with which he was familiar; it is probable that he saw no
material of C. proavita or C. gudalupensis. The relatively few polyploid
populations that he encountered were presumbly referred to one of the
diploids, depending on their morphological relationships.

No subsequent author gained quite as clear a picture of the major
patterns of variation in the group, possibly because of an increasing
awareness of the way in which the polyploids recombined the charac-
teristics of what otherwise would have been distinctive diploid races.

In 1928, P. A. Munz published his comprehensive review of the
section. His concept of this complex was based on a comprehensive
species, Qenothera hirta (= Camissonia micrantha), with four varieties
corresponding broadly to the diploids Camissonia micrantha, C.
hirtella, C. ignota, and C. pallida. Dr. Munz did not realize the distine-
tiveness of C. lewisii and C. proavita and had very little material of the
latter upon which to base any judgment. Again the polyploids were
redistributed among the diploid taxa they most closely resembled.

In 1964, I outlined the limits of sect. Holostigma accepted here and
made new combinations at the specific level for the available names in
the group (Brittonia 16: 276-288). This led Dr. Munz to accept his
earlier varieties as species (FI. N. Amer. IT. 5: 159-163. 1965). He was
unable to list C. lewisii, C. proavita, and the five polyploid species, as
no earlier names were available for these entities.

The present study of Camissonia sect. Holostigma is clearly not
a definitive one, and much work remains to be done. On the other hand,
it represents 10 years of intensive fieldwork, cultivation of the plants
in the experimental garden, and the determination of the chromosome
numbers of 389 individuals from 335 populations (see Appendix). I
believe that the taxonomy presented here leads to a clearer picture of
the pattern of variation in the section than anything published pre-
viously, and I have had much more material to draw on than any
earlier worker concerned with. the group. Lines within and between
the nine diploid species appear clearly drawn, but the nature and
derivation of both tetraploids need to be investigated in much greater
detail, preferably by the techniques of experimental hybridization.
The delimitation of the three hexaploids seems clearer, but the same
sort of work could profitably be done here also. At any rate, it is
hoped that the present taxonomic synopsis offers a sound basis for
such studies. As pointed out in the Appendix, the group appears to

258 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

have several cytogenetic peculiarities which should make a further

investigation of the evolutionary patterns in it of more than usual

interest.

26. Camissonia cheiranthifolia (Hornem. ex Spreng.) Raimann, in Engl. & Prantl,
Natiirl. Pflanzenfam. III. 7: 217. 1893.

Venothera spiralis f. arcuata H. Lév., Monogr. Onoth. 222. 1905. Léveillé says
simply ‘Fruit simplement arque—falque” and this cannot be placed with
either of the two subspecies in the absence of authentic material.

Oenothera spiralis f. clypeata H. Lév., Monogr. Onoth. 222. 1905. Léveillé
says ‘‘Feuilles scutiformes, sessiles, comme mucronees au sommet”’; to assign
this name to either subspecies in the absence of authentic material would be
a guess.

Sphaerostigma spirale var. clypeatum (II. Lévy.) A. Nels., Bot. Gaz. 40: 60. 1905.

Short-lived perennial, usually flowering the first year, with a
circle of prostrate, decumbent, or more or less ascending stems
radiating from a central rosette which withers in age, the stems to
0.6 (-1.3) m. long, ultimately somewhat woody and developing a
conspicuously exfoliating reddish brown (less commonly whitish)
rhytidome. Plants more or less densely strigose pubescent on all parts,
rarely glabrous. Leaves narrowly ovate, 0.5-5 cm. long, 0.25-2.2 cm.
wide, sparsely and weakly serrulate, the apex acute, the base cuneate
to cordate or (in the basal rosette) attenuate; petioles to 2.5 cm. long
in the basal rosette, 0-10 mm. long on the branches. Inflorescence
with two types of trichomes, short-erect and long-villous. Hypan-
thium 2.1-8.5 mm. long, 1.5-5 mm. across at the summit, glabrous
within. Sepals 4-11.5 mm. long, 1.3-3.4 mm. wide, reflexed in pairs.
Petals 6-20 mm. long, 5-22 mm. wide, often red-dotted near the base.
Filaments of the episepalous stamens 2.8-8 mm. long, those of the
epipetalous ones 1.5-6 mm. long; anthers 1.3-3 mm. long. Style 6-23
mm. long, glabrous; stigma 0.4-1.5 mm. in diameter, surrounded by
or held above the anthers at anthesis. Capsule 1-2.5 em. long,
ca. 2-2.5 mm. thick, quadrangular in transection, often curled in one
or two spirals. Seeds 1.2-1.3 mm. long, 0.5-0.7 mm. thick. Gametic
chromosome number, n=7. Self-incompatible, self-compatible but
outcrossing, and autogamous races known.

Disrripution (Figure 30): Sandy slopes and flats along the im-
mediate coast and on islands from Coos Bay, Curry County, Oregon,
to the vicinity of San Quintin, Baja California; also known from the
east shore of San Francisco Bay and locally on sand dunes along the
lower Sacramento River, California. Near sea level to several hundred
feet elevation.

Although some 35 miles separate the southern limit of subsp.
cheiranthifolia, at Point Conception, from the northern limit of subsp.
suffruticosa, just above Goleta, both of the subspecies occur on San
Nicolas Island. Here, as on the other Channel Islands where the

RAVEN—GENUS CAMISSONIA 259

species occurs, subsp. cheiranthifolia is common and has been collected
a number of times. On 24 April 1966, I found a small colony of about
40 plants of subsp. suffruticosa on the open dunes ca. 1 mile east-
southeast of Seal Beach, at the northwest end of San Nicolas Island
(my no. 20761, collected with H. J. Thompson). The colony was sur-
rounded by about 15 intermediates and located near a road in a much
larger population of subsp. cheiranthifolia. It is possible that subsp.
suffruticosa was introduced on the island, but it was collected as early
as 1901 (Trask 45, GH, NY), so it more likely reached the island by
natural means. The population I studied presumably consists of self-
compatible plants, although the stigma was held well above the anthers
as usual in this subspecies.

Experimental hybrids were made between plants of subsp. suffruti-
cosa from Point Dume, Los Angeles County, California, and subsp.
cheiranthifolia from Jalama Beach, Santa’ Barbara County, Cali-
fornia. Both parental strains had 7 pairs of chromosomes at meiotic
metaphase I, and both consisted of self-compatible plants. The F;
hybrids (Raven 17746, 17747, 17934) were intermediate between the
parents morphologically, self-compatible, and fully fertile, although
the stigma was held above the anthers as in the subsp. suffruticosa
parent. Reciprocal hybrids were indistinguishable, and one hybrid
made in each direction was examined cytologically and found to form
7 pairs at meiosis, indicating a lack of structural differences. In terms
of pollen stainability, three plants with subsp. suffruticosa as the
pistillate parent had, respectively, 93, 80, and 88 percent stainable
pollen (based on samples of 100 grains in cotton blue in lactophenol),
and five with subsp. cheiranthifolia as the pistillate parent had,
respectively, 66, 72, 58, 53, and 77 percent stainable pollen. Both
parents had consistently more than 95 percent stainable pollen, thus
indicating a definite, but limited, reduction in fertility in the F,
hybrids apparently not associated with chromosomal differentiation
between the parental strains.
26a. Camissonia cheiranthifolia subsp. suffruticosa (S. Wats.) Raven, Brittonia 16:

284. 1964. FiGurEs 26, 27
Oenothera viridescens Hook., Fl. Bor. Amer. 1: 214. 1833. Type: From Santa
Barbara, Santa Barbara Co., Calif., or San Diego, San Diego Co., Calif,,
November or December 1793, 4. Menzies (KX).
Sphacrostigma viridescens (Hook.) Walp., Rep. 2: 77. 1845.
Ocenothera cheiranthifolia var. sufiiruticosa S. Wats., Proc. Amer. Acad. 8: 592
1873.
“Oenothera virescens Hook.,”’ Greene, Fl. Francise. 214. 1891.
Sphaerostigma spirale var. viridescens (Hook.) A. Nels., Bot. Gaz. 40: 60. 1905.
Oenothera spiralis f. viridescens (Hook.) H. Lév., Monogr. Onoth. 222. 1905.
Oenothera spiralis var. linearis Jeps., Man. Fl. Pl. Calif. 684. 1925. Type:
Sunnyside, San Diego Co., Calif., April 1908, H. \. Ha 3908 (UC).
Oenothera spiralis var. viridescens (Hook.) Jeps., Man. FI. Pl. Calif. 684. 1925.

260 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Ficures 26-29,—Camissonia cheiranthifolia, x 1: 26-27, C. cheiranthifolia subsp. suffruticosa
(Gregory 468, DS): 26, Flowering branch; 27, portion of fruiting branch. 28-29, C.
chetranthifolia subsp. cheiranthifolia, flowering branches: 28, Raven 20825, DS; 29,
Randall 220, DS.

RAVEN—GENUS CAMISSONIA 261

Often bushy, the branches ascending and the plants quite woody;
pubescence often dense and silvery, particularly in the summer and
fall. Hypanthium 5-8.5 mm. long, 3-5 mm. across at the summit.
Sepals 6-11.5 mm. long, 2.4-3.4 mm. wide. Petals (10-) 12-20 mm.
long, 14-22 mm. wide, usually with one or two red dots at the base.
Filaments of the episepalous stamens 5-8 mm. long, those of the epi-
petalous ones 3-6 mm. long. Anthers 2.2-3 mm. long. Style 18-238
mm. long; stigma held well above the anthers at anthesis. Gametic
chromosome number, n=7. Outcrossing, with self-incompatible and
self-compatible populations known.

LecrotyPe: Santa Barbara, Santa Barbara County, California,
Brewer 307 (GH). This appears to be the only specimen in the Gray
Herbarium annotated with this name by Watson. Considering this,
and the fact that it is from a definite locality, it appears to be a better
choice of lectotype than the first-listed specimen assumed by Munz
(Bot. Gaz. 85: 269. 1928) to be the type.

DistriBuTIoN (Figure 30): Sand dunes and beaches along the im-
mediate coast, from Deveraux Dunes just west of Goleta, Santa
Barbara County, California, to 6 miles south of San Quintin, Baja
California (Ballou & Canby in 1925, C, F, POM, UC); San Nicolas
Island, Ventura County, California (see below); San Martin Island,
Baja California. From near sea level to several hundred feet elevation.

The chromosome number has been determined from 8 individuals
from 7 different populations scattered throughout the range of this
entity (see Appendix). Populations from 5.5 miles southeast of Point
Mugu, Ventura County, California (R20161—6 plants tested for
growth of pollen tubes), Point Dume (see below), and Playa del Rey
(Kyhos 65-47, 1 plant grown at Stanford), Los Angeles County,
California, consisted of self-compatible plants, which were however
outcrossing, with the stigma elevated well above the stamens at
anthesis and not contacting them even when the flowers closed. On
the other hand, populations from farther south, in San Diego County,
California, consisted of self-incompatible individuals: Del Mar Beach,
R20167 (11 plants tested for pollen-tube growth); DeAnza Cove.
Mission Bay, Wedberg (15 plants grown at Stanford as R66-224).
Andrena (Onagrandrena) oenotherae Timb. was an abundant visitor to
the flowers of this subspecies at all localities investigated, extending
from the northern limit to south of Ensenada, Baja California; this
large, black, oligolectic bee doubtless is a very important agent
promoting outcrossing in this subpecies. Andrena (Diandrena) para-
chalybea Viereck, 2 somewhat smaller oligolectic bee, is locally fre-
quent as an early morning visitor to the present subspecies also.

It appears likely that populations very similar to the self-incom-
patible ones presently referred to Camissonia cheiranthifolia subsp.

262 OONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Ficure 30.—Portion of the western United States and adjacent Mexico, showing the
ranges of species of Camissonia sect. Holostigma: @=C. cheiranthifolia subsp. suffrutt -
cosa; (Q=C. cheiranthifolia subsp. cheiranthifolia; A=C. bistorta.

suffruticosa gave rise on the one hand to the self-incompatible, annual
C. bistorta and on the other to the autogamous, perennial C. cheiran-
thifolia subsp. cheiranthifolia. An account of the anatomy of subsp.
suffruticosa has been given by Purer, Ecol. Monogr. 6: 1-88. 1936.
At the present time, it occurs sympatrically with both of its pre-
sumed derivatives, and its relationship with subsp. eheiranthifolia has

RAVEN—GENUS CAMISSONIA 263

been considered above. It also occurs sympatrically with C. lewisi at
many localities.

Often Camissonia bistorta and C. cheiranthifolia subsp. suffruticosa
occupy adjacent but sharply distinct habitats. At Point Dume, Los
Angeles County, California, the former occurs on the upper reaches
of the sandy beach and the latter forms colonies in the sandy grassland
on the tops of the bluffs. Here the two do not grow immediately
adjacent to one another, and intermediates have not been observed.
On the other hand, where the habitats of the two species do intergrade
more subtly, hybrids occur rather commonly. Populations of perennial
plants otherwise resembling (. bistorta are characteristic of some areas
and occur frequently along the entire coast of San Diego County,
California, and in northern Baja California. Extensive series of inter-
mediate populations occupy the sandy riverbeds in the vicinity of
San Diego, a habitat that is clearly intermediate between the coastal
dunes and flats inhabited by (. cheiranthifolia and the firmer sandy
soil farther inland where (. bistorta occurs. Where intermediate plants
do occur, they generally appear as relatively uniform local populations.
The fertility of such plants is usually not reduced; for the collections
indicated with an asterisk in the following list, pollen stainability was
at least 96 percent for the average of two samples each of 200 pollen
grains in cotton blue in lactophenol. In the Jones collection from
Pacific Beach, pollen stainability was reduced to 79 percent; this
collection might represent an F,, or might be sterile for some other
reason.

Specimens of apparent hybrids between Camissonia cheiranthifolia
subsp. suffruticosa and C. bistorta examined :

U.S.: CALIFORNIA! SAN DIEGO cOo.: Encinitas, Lewis 1202* (LA), Epling &
Robison in 1932 (LA); Cardiff, Benson 10352* (POM); Oneonta, Chandler 5094
p.p. (NY); U.S. Hwy. 101 N. of La Jolla, Benson 4286 (POM); Valley Center,
Woodcock in 1928 (SD); Crown Point near Pacific Beach, Purer in 1935 (POM)r
Sunnyside, Hall 3908 (F); Foster, 400 ft., Smith 5257 (F); San Diego River nea;
the Mission, Abrams 3412 (CAS, GH, LE, NMC, NY, PH, POM, UC); Mission
Valley, Purer 6124 (DS, POM, SD), Gander 4929 (SD); Cuyamaca Road 0.5
mi. N. of Mission Gorge Road, San Diego R., Santee, Wedberg 533* (DS) El;
Monte Park Grove, Alderson in 1934 (SD); San Luis Rey, Gander 3309 (;SD)
foot of San Miguel Mt., Youngberg 44 (POM), Gander 154.5 (SD); 5. San Diego,
Chandler 5097 (DS, NY); wash between National City and Chula Vista, Wolf
2086 (DS, RSA, UC), Gander 4920 (SD); Otay Creek, 2 mi. below dam, Peirson
3410 (RSA); Otay, Higgins in 1954 (UC); Nestor, near Tia Juana, Wiggins 3229
(DS LA, UC).

BAJA CALIFORNIA: Dry Tijuana R. bed, 80 ft., Smith 5206 (F, PH); 2
mi. N. of Rosarito Beach, Wiggins & Gillespie 3882 (CAS, DS, F, GH, NY,
POM, RSA); Descanso Dunes, R17059* (DS) Higgins in 1949 (RSA), in 1950
(RSA, SD), Creighton 525 (COLO), Purer in 1932 (SD); sea bluffs 32 mi. N. of
Ensenada, Munz 22702 (RSA); Pinita, Harbison in 1950 (SD); Hot Springs,
Jones in 1882 (POM, RM, UC); Cariso Creek, T. S. Brandegee in 1893 (UC).

264 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

These two entities have been hybridized experimentally, using a
strain of C. bistorta from 6.5 miles east of San Juan Capistrano, Orange
County, California (R18782), and one of C. cheiranthifolia subsp.
suffruticosa from De Anza Cove, Mission Bay, San Diego County,
California, collected by H. L. Wedberg. Both parental strains were
self-incompatible, and both regularly formed 7 pairs of chromosomes
at meiotic metaphase I. Seeds were readily obtained in both directions,
but only those derived from the cross using C. cheiranthifolia as the
female parent germinated. Four individuals from a large F, progeny
were examined cytologically. The maximum association of chromo-
somes observed was a ring of 8 and 3 ring bivalents, with the large
ring most often present as a chain and one of the bivalents being a
rod (1.e., forming only one chiasma) in most cells. Samples of 200
pollen grains from each of three individuals, stained in cotton blue
in lactophenol, gave the following percentages of apparently viable
pollen: 24.5, 24.5, and 34.7%. Although the hybrids were self-incom-
patible like both of their parents, cross-pollination gave essentially
full seed-set, and doubtless these plants would easily backcross with
either parent, thus producing individuals similar to many of the
putative hybrids observed in the field.

Two of the apparent hybrids (possibly backcrosses with C. cheir-
anthifolia) found in the field were examined cytologically and showed
no evidence of structural heterozygosity:

BAJA CALIFORNIA: Meadows E. of Rosarito, Bates 2331; ca. 29 mi. S. of
Tijuana, Klein & Gregory 1274* (RSA).

One of the early binomials applied to plants of this section was
Holostigma bottae Spach, Nouv. Ann. Mus. Paris 4: 336. 1835. Its
description is such that Munz (N. Amer. Fl. IT. 5: 163. 1965) appro-
priately stated, “Possibly an earlier name for Oenothera bistorta’’
[= Camissonia bistorta]. Thus it is of considerable interest and impor-
tance to establish the identity of Spach’s plant, with its type locality
“In California australi.”” Through the kindness of Dr. Alicia Lourteig,
I have been able to ascertain that there is apparently no specimen in
the Paris herbarium annotated “/olostigma bottae.’’ There is, how-
ever, a specimen collected by Paolo Emilio Botta at San Diego,
California, in 1827, labeled (probably in Spach’s hand) “Sphaero-
stagma.”’ Spach used the names Sphaerostigma and Holostigma essen-
tially interchangeably, and this appears to be the only specimen of
the group present at Paris that was collected by Botta. Furthermore,
it agrees perfectly and in detail with Spach’s rather extensive descrip-
tion of Holostigma bottae. Consequently, I designate it the lectotype
of this species.

Unquestionably, the specimen is one of the series of populations
intermediate between Camissonia bistorta and C. cheiranthifolia subsp.

RAVEN—GENUS CAMISSONIA 265

suffruticosa so common in the vicinity of San Diego and perfectly
matched by many individuals growing today along the sandy bed
of the San Diego River (e.g., Wedberg 533 or Abrams 3412, cited above).
Its percentage of stainable pollen based on a sample of 200 grains,
is 93%. In view of this, the name Holostigma bottae cannot be regarded
as belonging to either of its parental species, and there seems no need
to transfer it to Camissonia to serve as a binomial designation for
the series of interspecific hybrids from which its type collection was
derived more than 140 years ago.

26b. Camissonia cheiranthifolia subsp. cheiranthifolia FiGurEs 28, 29

Oenothera cheiranthifolia Hornem. ex Spreng., Syst. 2: 228. 1825.

Ocenothera spiralis Hook., Fl. Bor. Amer, 1; 213. 1833. Type: Coastal Cali-
fornia (either at Trinidad Head, San Francisco, Santa Cruz, or Monterey),
1792-1794, A. Menzies (K).

Holostigma cheiranthifolium (Hornem. ex Spreng.) Spach, Nouv. Ann. Mus.
Paris III. 4: 335. 1835.

Sphaerostigma cheiranthifolium (Hornem. ex Spreng.) Fisch. & Mey., Ind.
Sem. Hort. Petrop. 2: 50. 1835.

Sphaerostigma spirale (Hook.) Fisch. & Mey., Ind. Sem. Hort. Petrop. 2:
50. 1835.

Agassizia cheiranthifolia (Hornem.) Spach, Hist. Nat. Vég. 4: 348. 1835.

Holostigma spirale (Hook.) Spach, Nouv. Ann. Mus. Paris III. 4: 336. 1835.

Oenothera nitida Greene, Pittonia 1: 70. 1887. Type: San Miguel I., Santa
Barbara Co., Calif., September 1886, EF. L. Greene (ND; isotypes, NY,
PH, UC, US).

Sphaerostigma nitidum (Greene) Small, Bull. Torrey Cl. 23: 190. 1896.

Oenothera spiralis var. nitida (Greene) Jeps., Man. Fl. Pl. Calif. 684. 1925.

Ocenothera cheiranthifolia var. typica Munz, Bot. Gaz. 85: 268. 1928.

Ocnothera cheiranthifolia var. nitida (Greene) Munz, Bot. Gaz. 85: 269.1928.

Branches usually prostrate or decumbent and largely herbaceous;
pubescence rarely dense and silvery. Hypanthium 2.1-4.2 (—4.8) mm.
long, 1.5-3 mm. across at the summit. Sepals 4-5.6 (—6.7) mm. long,
1.3-1.9 (-2.1) mm. wide. Petals 6-11 mm. long, 5-12 mm. wide, rarely
with one or two red dots near the base. Filaments of the episepalous
stamens 2.8-4.5mm. long, those of the epipetalous ones 1.5-3 mm. long.
Anthers 1-1.5 mm. long. Style 6-9 mm. long; stigma surrounded by
both sets of anthers which shed pollen directly onto it at anthesis, or
very rarely held above the anthers. Gametic chromosome number,
n=7. Facultatively autogamous, but sometimes outcrossing; self-
compatible.

Lecrotyre: “Ex hort. semina e California mis. Wormskiold” (C),
doubtless from seed collected in the vicinity of Fort Ross, Sonoma
County, or San Francisco, California. During most of the 19th
century, the species was erroneously attributed to Chile, based on
Sprengel’s statement in the protologue; no evidence for this has ever
been found, and it is assumed that Sprengel made a simple mistake
(cf. Munz, Bot. Gaz. 85: 268. 1928).

266 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Disrrispution (Figure 30): Sandy slopes and flats along the coast,
from Coos Bay, Curry County, Oregon, south to Point Conception,
Santa Barbara County, California; also on the east shore of San
Francisco Bay in Alameda and Contra Costa Counties, California;
sand dunes and flats along the lower Sacramento River, California,
often with Lupinus arboreus, another beach species (4 miles west. of
Antioch, Contra Costa County, Heller in 1942, WTU; between
Antioch and Rio Vista bridges, Sacramento County, Eastwood &
Howell 8636, CAS; Brannan Island State Park, Sacramento County,
R20178, DS); San Miguel, Santa Rosa, Santa Cruz, San Nicolas, and
San Clemente Islands.

The gametic chromosome number of this subspecies has been
determined as n=7, with no evidence of structural heterozygosity, in
single individuals from eight populations throughout its range (see
Appendix). Although self-pollination generally occurs in this taxon
upon the closing of the flowers, some populations, especially in the
San Francisco Bay area, are much visited by oligolectic bees of the
genus Andrena (Onagrandrena) and are doubtless outcrossed to a
large extent. Other populations, such as those around Monterey Bay
and farther south, appear not to be visited often by insects of any
kind and are probably predominantly self-pollinated. The flowers of
subsp. chetranthifolia generally open more tardily than those of subsp.
suffruticosa, which also occupies areas where the mornings are not
often foggy.

This subspecies is quite variable in pubescence, and there appears
to be a cline extending from the San Francisco area, where the plants
are usually decumbent and greenish, to the region of Morro Bay, San
Luis Obispo County, where many of the plants are more erect and
silvery pubescent, resembling indiviuals of subsp. suffruticosa. Such
plants may also have larger flowers than usual for subsp. eheiranthifolia
but in general are not exceptional in flower size.

Other variants in pubescence may occur as individual plants,
mixed in populations of more usual individuals. For example, at the
mouth of Hazard Canyon, San Luis Obispo County, California,
Robert Hoover collected an interesting local race with large bright
green but still pubescent leaves, up to 4.5 em. long and 2 cm. wide,
growing with the usual more heavily pubescent and smaller leaved
plants. Occasional completely glabrous or subglabrous plants occur in
populations; these individuals have been named Qenothera nitida
Greene. Such plants agree with the rest of the species in chromosome
number, They obviously comprise only one of the more conspicuous
segregating elements in their interbreeding populations, and do not
merit formal taxonomic recognition. Such glabrous plants have been
collected around Monterey Bay, Monterey County, California, where

RAVEN—GENUS CAMISSONIA 267

they are well known; in Santa Barbara County, California, at Surf
(Pierson 8308, POM, RSA) and at Jalama Beach (Smith 2848, RSA);
and on Santa Rosa (East Point, Munz & Hoffmann 11744, POM) and
San Miguel Islands.

On San Martin Island, off the coast of Baja California, individuals
of subsp. suffruticosa occur which are more prostrate than usual for
that subspecies and thus resemble subsp. chetranthifolia: Moran 10571
(ARIZ, COLO, BS, RSA, SD, UC). As these plants agree with other
Baja California populations of subsp. suffruticosa in flower size and
pubescence, I have regarded them as representing a maritime ecotype
of that taxon. The possibility that they may in fact belong with
subsp. cheiranthifolia and represent a relictual station for that more
northern subspecies cannot, however, be completely ruled out at
present. It does seem more likely that they have converged with it
in habit on the basis of their similar ecology.

Camissonia cheiranthifolia subsp. cheiranthifolia occurs sympat-
rically with C. micrantha, and with C. guadalupensis subsp. clementina
and locally intergrades with C. cheiranthifolia subsp. suffruticosa as
discussed above.

27. Camissonia bistorta (Nutt. ex Torr. & Gray) Raven, Brittonia 16: 284.
1964. FiIGuRE 31
Oenothera heterophylla Nutt. ex Hook. & Arn., Bot. Beechey Voy. 341.
1938; non Spach 1835. Type: San Diego, San Diego Co., California,
April-May 1836, 7. Nuttall (BM).

Oenothera bistorta Nutt. ex Torr. & Gray, Fl. N. Amer. 1: 508. 1840.

Sphaerostigma bistortum (Nutt.) Walp., Rep. 2: 77. 18438.

Oenothera bistorta var. veitchiana Hook., Bot. Mag. 84: pl. 5078. 1858.
Lectotype: Plains about San Gabriel Mission, Los Angeles Co., California
W. Lobb 416, grown at Hort. Veitch in 1857 (K).

Sphaerostigma veitchianum (Hook.) Small, Bull. Torrey Cl. 23: 191. 1896.

Sphaerostigma bistortum var. veitchianum (Hook.) A. Nels., Bot. Gaz. 40: 59.
1905.

Oenothera cheiranthifolia f. delicatula H. Lév., Monogr. Onoth. 216, fig. on
p. 214. 1905.

Oenothera bistorta var. typica Munz, Bot. Gaz. 85: 266. 1928.

Annual, or rarely a short-lived perennial, with one to several
ascending or several spreading, decumbent stems radiating from a
central rosette, these to 0.8 mm. long, with a conspicuously exfoliating
pale rhytidome. Plants strigose or more commonly villous. Leaves
very narrowly elliptic in the basal rosette to narrowly lanceolate or
lanceolate on the stems and in the inflorescence, rarely all linear,
1.2-12 em. long, 0.15-1.5 em. wide, denticulate, but often only sparsely
and inconspicuously so, the apex acute, the base narrowly cuneate
(in the basal rosette) to cuneate or subcordate; petioles to 4 cm. long
in the basal rosette, the upper leaves usually subsessile. Inflorescence
with two types of trichomes, short-erect and long-villous. Hypanthium

268 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Ficures 31-32.—Species of Camissonia sect. Holostigma, * 1:31, C. bistorta (Raven 16935
DS); 32, C. lewistt (Raven 17021, DS).

RAVEN—GENUS CAMISSONIA 969

2-5 (-7.5) mm. long, 1.5-4 mm. across at the summit, glabrous to
pubescent in lower portions within, often with prominent red dots
near the summit within. Sepals (2.3—-) 5-8 (-11) mm. long, (0.8—) 1-2
mm. wide, reflexed in pairs. Petals (4.2—-) 7-15 mm. long, (2.5—-) 6.5-12
(-16) mm. wide, each petal almost always with a bright red spot
(rarely two) near the base. Filaments of the episepalous stamens (1—)
1.5-3.5 mm. long, those of the epipetalous ones (0.5—) 1-2.5 mm. long;
anthers (0.5—) 1.3-2 (-2.5) mm. long. Style (5.5-) 7-12 mm. long, us-
ually glabrous but occasionally pubescent near the base; stigma (0.7—)
1-1.8 mm. in diameter, held well above the anthers at anthesis. Cap-
sule 1.2-4 em. long, 1.5-2.5 mm. thick, more or less quadrangular in
transection, straight or somewhat contorted. Seeds 0.9-1 mm. long,
0.3-0.55 mm. thick. Gametic chromosome number, n=7. Self-
incompatible.

TypE: San Diego, San Diego County, California, April-May 1836,
T. Nuttall (NY; isotypes, BM, Ik, GH, PH).

Distrisution (Figure 30): Sandy fields near the coast or clayey
grassland, often in openings of coastal sage, from Piru Creek and the
Ojai Valley in Ventura County south and east through southern Los
Angeles, southwestern San Bernardino, Orange, western Riverside,
and the western two-thirds of San Diego Counties, California, and
south in cismontane Baja California to Ojos Negros and San Vicente,
never far from the coast; reaching the margins of the desert in San
Bernardino (near Hesperia) and San Diego Counties. Usually at low
elevations, sea level to 2,000 ft., but exceptionally as high as 8,000
ft. alt. in the mountains, where it occurs about the borders of meadows.
Collected as a waif beyond its main area in the Goleta salt marsh,
Santa Barbara County, California, ‘Introduced apparently with
Stream gravel... ,’? 20 June 1959, Pollard (CAS) and on ballast
heaps at Nanaimo, Vancouver Island, British Columbia, 13 July
1893, Macoun (NMC). Camissonia bistorta has occasionally been
cultivated in Europe, at least in the 19th century.

The chromosome number has been determined from 29 individuals
derived from 22 populations scattered throughout the range of the
species (see Appendix). Populations from 6.5 miles northeast of San
Juan Capistrano, Orange County, California, R18782 (8 individuals) ;
lower edge of Cleveland National Forest, just above San Juan Camp-
ground, Santa Ana Mountains, Orange County, California, R18783
(6 individuals) ; Point Dume, Los Angeles County (2 individuals) ; and
Claremont, Los Angeles County, California (10 individuals), were
grown in the experimental garden and found to be self-incompatible.
In addition, the Claremont strain was grown by Hagen (Indiana
Univ. Publ. Sci. Ser. 16: 309. 1950) who also found it to be self-
incompatible. Several species of large bees of the genus Andrena sub-

270 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

genera Diandrena and Onagrandrena are frequent oligolectic visitors to
the flowers of Camissonia bistorta soon after they open in the early
morning and doubtless are the most important agents in effecting
cross-pollination in this species.

Camissonia bistorta may well be an annual derivative of popula-
tions similar to the self-incompatible ones referred to C. cheiranthifolia
subsp. eheiranthifolia, although the reverse hypothesis cannot. be
excluded at present. Depauperate individuals of this species, found
in dry years or unfavorable sites, often have relatively small flowers,
as do plants at the end of their flowering period. The stigma is, how-
ever, consistently held above the anthers, and such plants can thereby
easily be separated from the numerous related autogamous species
in sect. Holostigma. This species occurs sympatrically with C. lewisii,
and a single obvious F; hybrid between the two species has been col-
lected at Playa del Rey, Los Angeles County, California, Wedberg 272
(herb. San Diego State College). This plant had about 23 percent
stainable pollen and apparently full seed-set; at meiosis it formed
variable configurations of chromosomes, the maximum observed in
14 cells being 1 pair, 1 ring of 4, 1 chain of 8, and 5 univalents (H. L.
Wedberg, pers. comm.). Camissonia bistorta also occurs adjacent to
and hybridizes extensively with C. cheiranthifolia subsp. suffruticosa,

Ficure 33,—California and Baja California, showing ranges of species of Camissonia sect.
Holostigma: A=C. lewisii, with A=diploid chromosome counts (n=7); O=C. pallida
subsp. hallti with @=diploid counts (n=7).

RAVEN—GENUS CAMISSONIA 271
as explained in the section dealing with that taxon. It has likewise
been observed growing sympatrically with C. micrantha, C. hartella,
C. ignota, C. intermedia, C. confusa, and C. robusta, but hybrids between
these predominantly autogamous species and C. bistorta have not been
found. Camissonia bistorta is probably relatively similar to the popula-
tions ancestral to at least some of the autogamous taxa in this section,
the most obvious relationship being that with C. lewrs??.

In his original treatment of this species (Bot. Gaz. 85: 266-267.
1928), Munz included two varieties in addition to the typical one.
The first of these, Oenothera bistorta var. veitchiana Hook., was applied
to inland populations with relatively slender, long, and straight cap-
sules which differed in these respects from the series of coastal popu-
lations, including the type of the species, which have thicker, shorter,
usually coiled capsules. These two entities intergrade so broadly that
it does not appear useful to segregate them taxonomically. The other
variety recognized by Munz, 0. bistorta var. hallii (Davids.) Jeps.,
intergrades completely with the entity designated by Munz 0. hirta
var. eafoliata (A. Nels.) Munz and is here treated, together with
that taxon, as the species Camissonia pallida (Abrams) Raven. Cam-
issonia pallida and C. bistorta do not occur together, as far as I know,
and, although both entities are diploid (n—7), there appears to be
no evidence for morphological intergradation or even of overlapping
patterns of variation.

The report of Camissonia bistorta from Santa Catalina Island,
California (Millspaugh & Nuttall, Field Mus. Publ. Bot. 5: 183.
1923: as Sphaerostigma bistortum) is based on misidentified specimens
of C. robusta and C. californica, which I have seen.

28. Camissonia guadalupensis (S. Wats.) Raven, Brittonia 16: 284. 1964.

Erect annual with short branches usually arising above the base;
plants thick-stemmed, heavy-set, with conspicuously exfoliating epi-
dermis, subsucculent, villous or strigose all over, 2-18 (-35) em.
tall. Leaves narrowly elliptic in the basal rosette to narrowly ovate
on the stems and in the inflorescence, 1.2-3.8 (-9.5) cm. long, 0.5—
1.2 (-1.8) em. wide, sparsely and weakly denticulate, the apex acute
(lower leaves) to obtuse (upper leaves), the base attenuate (lower
leaves) to rounded or truncate (upper leaves); leaves subsessile.
Inflorescence with short glandular trichomes in addition to the pre-
dominant sort found on other parts of the plant. Hypanthium 1.6—
2.4 mm. long, 1.2-1.6 mm. across at the summit, sparsely pubescent
with short erect trichomes in lower portions within. Sepals 1.9-3.2
mm. long, 0.8-1.6 mm. wide. Petals 2.8-4.2 mm.long,2-3.6 mm. wide,
sometimes with a prominent red dot near the base of each one. Fila-
ments of the episepalous stamens 1.3-2.3 mm.long, those of the epi-
petalous ones 0.4-1.6 mm. long; anthers 0.4-0.8 mm. long. Style

295-655 O—68 -S

27? CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

3.2-4.5 mm. long, short-pubescent near the base; stigma 0.5—0.8
mm. in diameter, surrounded by the anthers at anthesis. Capsule
10-18 mm. long, 2.8-3.5 mm. thick near the base, very heavy in appear-
ance, quadrangular in transection and deeply grooved along the
lines of dehiscence, straight or slightly curved outward. Seeds 0.75—
0.9 mm. long, 0.4-0.55 mm. thick. Gametic chromosome number,
n=7. Autogamous.

DistRiBUTION (Figure 36): Sandy flats, dunes, and arroyo bottoms,
San Clemente Island, Los Angeles County, California (subsp. elem-
entina), and Isla Guadalupe, Baja California (subsp. guadalupensis).
Sea level to perhaps 1,200 ft. alt.

Reciprocal F, hybrids were made between progenies of R17628
(subsp. clementina) and Moran & Ernst 6737 (subsp. guadalupensis).
These plants had a mixture of all three sorts of pubescence found in the
two subspecies, the strigose pubescence of subsp. guadalupensis,
however, being relatively sparse. The hybrids had a red dot near the
base of each petal, as in subsp. clementina, but this was reduced in
size. They appeared to have full seed-set, comparable to that of the
parents. Two of the F; individuals in which subsp. guadalupensis
was the pistillate parent (=R18771) and one in which subsp. clem-
entina was the pistillate parent had 7 bivalents at meiotic metaphase I
and diakinesis, and thus no evidence of chromosomal structural
heterozygosity. The pollen stainability in cotton blue in lactophenol,
calculated from a sample of 200 grains in each case, was measured in
three plants in which subsp. guadalupensis was the pistillate parent
and found to be 41.5, 49.5, and 41.5 percent; in two F, plants in which
subsp. clementina was the pistillate parent, the percentages of stainable
pollen were, respectively, 58 and 38 percent. As compared with the
98-100 percent stainable pollen found in samples taken from both
parents grown under the same conditions, these figures indicate a
substantial reduction in fertility in the hybrids, although seed-set
does not appear to be impaired. Nonetheless. the very close mor-
phological relationship between these two entities and their evident
distinctiveness when compared with other species can seemingly best
be indicated by retaining them within a single taxonomic species.
28a. Camissonia guadalupensis subsp. guadalupensis FIGURE 34

Oenothera guadalupensis S. Wats., Proc. Amer, Acad. 11: 115. 1876.
Oenothera quadalupensis subsp. guadalupensis Munz, N. Amer. Fl. II. 5: 160.
1965.

Pubescence strigose with some longer trichomes in inflorescence,
but no glandular ones. Petals without red dot at base. Gametic
chromosome number, n=7. Autogamous.

Type: Isla Guadalupe, Baja California, February—May 1875, F.
Palmer (GH).

RAVEN—GENUS CAMISSONIA 273

Ficures 34-35,—Camissonia guadalupensis, X 1:34, C. guadalupensis subsp. guadalupensis

(Raven 18175, DS); 35, C. guadalupensis subsp. clementina (Raven 17284, DS).

274 OONTRIBUTIONS FROM THE NATIONAL HERBARIUM

DisrripuTion (Figure 36): Isla Guadalupe, Baja California, known
with certainty only from the sandy bed of Arroyo Melpomone,
900-1,300 ft. alt.

ADDITIONAL SPECIMENS EXAMINED:

BAJA CALIFORNIA: Ista Guapatupe: Bed of Arroyo Melpomone, Moran
12070 (SD), Moran & Ernst 6737 (SD); progeny of Moran & Ernst 6737, R17530
(DS), 18175 (Ds).

Ficure 36.—Portion of the southwestern United States and northwestern Mexico, showing
the ranges of species of Camissonia sect. Holostigma; +-=C. guadalupensis subsp.
guadalupensis; %=C. guadalupensis subsp. clementina; A=C. proavita, with A=dip-
loid chromosome counts (n=7); O=C. micrantha, with @=diploid counts (n=7).

RAVEN—GENUS CAMISSONIA 275

The chromosome number has been determined from two plants of
this population, grown in the experimental garden (see Appendix).
Plants of this subspecies were more highly autogamous than those of
subsp. clementina when the two were grown in the experimental garden.
28B. Camissonia guadalupensis subsp. clementina (Raven) Raven, Brittonia

16: 284, 1964. FIGURE 35
Oenothera guadalupensis subsp. clementina Raven, Aliso 5: 332. 1963.

Pubescence dense, villous, with short glandular trichomes also
present in the inflorescence. Petals each with a red dot near the
base. Gametic chromosome number, n=7. Autogamous.

Type: Common on pebbly dune above ocean south of Eel Point,
San Clemente Island, elevation 40 ft., Los Angeles County, California,
9 April 1962, P. H. Raven 17125 (RSA).

Distripution (Figure 36): Restricted to San Clemente Island,
Los Angeles County, California, where it is common on dunes around
the north end and down the west shore, perhaps to the south end;
up to 100 ft. elevation.

ADDITIONAL SPECIMENS EXAMINED:

U.S.: CALIFORNIA! SAN CLEMENTE I., LOS ANGELES Co.: Trask 1684 (E, POM,
US), Murbarger 154 (UC), Piehl 631101 (SBBG), Blakley 5224, 5254 (both SBBG),
R17260, 17284, 17301, 17628, 17954, 18023 (all DS).

The chromosome number has been determined from single indivi-
duals of three populations, including the type collection (see Appendix).
Plants of the subspecies were facultatively autogamous when grown
in the experimental garden at Stanford, the flowers not selfing as
early in the day on which they open as in subsp. guadalupensis. In the
latter, the anthers scarcely left contact with the stigma at any time
after opening. Camissonia guadalupensis subsp. clementina has been
observed growing sympatrically with C. cheiranthifolia subsp.
cheiranthifolia.

29. Camissonia lewisii Raven, sp. nov. FIGuRE 32
Ocnothera hirta var. typica Munz, Bot. Gaz. 85: 262. 1928; pro parte.
Oenothera micrantha sensu Munz, N. Amer. FI. II. 5: 106. 1965; pro parte.

Herba annua villosa, ad C. micrantha similis, ramis plurimis decum-
bentibus e rosula centrali radiantibus, rariore caule singulari erecto;
rami ad 0.6 m. longi. Folia 1-8 cm. longa. 0.2-1.1 cm. lata, denticulata.
Inflorescentia pilis brevibus gladulosis longioribusque erectis patenti-
bus pilosa. Hypanthium 1.5-4 mm. longum, ostio 1-2.5 mm. diameter.
Sepala 1.7-3.5 mm. longa, 1-1.5 mm. lata. Petala (2.5 —) 3-5.5 mm.
longa, (2-)2.5-5 mm. lata, plerumque prope basin puncto rubro uno
vel duo. Filamenta staminum episepalorum 2-2.8 mm. longa, illa
epipetalorum 1-1.7 mm. longa; antherae 0.7-1.2 mm. longae. Sylus
2.8-4.5(-5.5) mm. longus; stigma 0.7-1.8 mm. diametro, sub antheris
circumnexum. Capsula 13-20 mm. longa, 1.8-2.2 mm. crassa, tran-

276 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

sectione valde quadrangularis in spiram laxam contorta. Semina
0.7-0.8 mm. longa, 0.5-0.6 mm. crasa, obovoidea. Chromosomatum
numerus gameticus, n=7. Autogama.

Type: Common in open clayey grassland, with EHschscholzia,
Abronia, and grasses, 1.1 mile north of Colonia Guerrero, elevation
200 ft., Baja California, 30 March 1962, P. H. Raven 17035 (DS;
isotypes, RSA, US).

Disrripution (Kigure 33): Open sandy and clayey grassland,
coastal dunes and beaches, never far from the coast, from sea level
to 1,000 ft. alt., Point Dume and the Los Angeles Basin, Los Angeles
County, California, south to Cardén Grande at the northern edge of
Territorio de la Baja California Sur.

SELECTED SPECIMENS EXAMINED:

U.S.: CALIFORNIA: LOS ANGELES co.: Hollywood, Davidson 1437 (LAM);
Inglewood, Abrams 3232 (DS, GH, LA, NMC, NY, PH, POM, UC, US); Roscoe,
Eastwood 242 (CAS); Point Dume, Mars (LA); Ballona Harbor, Abrams 1192
(DS, POM); San Pedro, Eastwood 166 (CAS). ORANGE co.: Santa Ana, King in
os (JEPS); Bee Canyon, Rancho Santa Ana, Howell in 1928 (RSA). SAN DIEGO

: Fallbrook, Jones in 1882 (DS, GH, NY, POM, RM, UC); Agua Hedionda,
eae 3409 (DS, RM, RSA); 4.6 mi. E. of San Pasqual, R17406 (DS); El
Segundo, Purer in 1931 (SD); San Luis Rey, Gander in 1937 (SD); Crown Point,
Gander 143.8 (SD); Oneonta, Chandler 5094 p.p. (NY); Bird Rock, F. E. & EF. S.
Clements 184 (COLO, F, GH, NY, PH, UC, UCSB), 185 (COLO, F, GH, NY,
PH, UC, UCSB); Point Loma, Eastwood in 1913 (CAS, GH, NY, US); Ocean
Beach, Brandegee 1645 (CAS, F, GH, LE, NMC, NY, POM, RM, UC, US);
1.5 mi. from Jamul toward Otay Dam, Wiggins 1923 (DS).

BAJA CALIFORNIA: Tijuana valley, Orcutt 82 (F, GH, US); ranch 29 mi.
SW. of Tijuana, Jones in 1925 (POM); Descanso Dunes, 1817060 (DS); s. end
of Ensenada Bay, R17006 (DS); 9.3 mi. S. of San V icente, R17044 (DS); 2.5 mi.
W. of San Vicente, Kadish in 1956 (L A); 7.4 mi. N. of Ejido México, R17041
(DS); 4 km. 8. of Ejido Méxieo, R12230 (DS); 4 mi. W. of Colonia Guerrero,
Blakley 6302 (DS, SBBG); Santo Domingo (Hamilton’s Ranch and _ vicinity),
Wiggins 4494 (CAS, DS, GH, LA, NY, POM, UC, US); near Red Rock above
Hamilton’s Ranch, Thomas 130 (DS, OSC); San Quintin, Epling & Robison in
1935 (DS, UC, NY), San Quintin Bay, 1889, Palmer 617 (CAS, F, GH, NY,
PH, UC, US); Socorro, Cronemiller 3076 (DS, POM); 20.6 km. S. of road to San
Quintin, R12396 (DS); 20 mi. N. of El Rosario, Harbison in 1935 (SD); 3.7 mi. E.
of Punta Baja toward El Rosario, Gregory 356 (RSA); 16 km. SE. of El Rosario,
R12466 (DS); El Caiién Rancho, near Calmalli, Haines & Stewart in 1935 (DS);
El] Poso Allemin, Brandegee in 1889 (UC); Cardén Grande, Brandegee in 1889
(UC).

Camissonia lewisii, an autogamous diploid species, can easily be
recognized by its short, usually once-coiled capsules that are square
in transection in living material. It is quite similar to some coastal
populations of C. bistorta, such as those found in San Diego County,
California, and may have been derived from populations of this sort.
The chromosome number has been determined from 23 individuals
representing 19 populations, scattered throughout the range of the

RAVEN

GENUS CAMISSONIA 277

species, including the type collection (see Appendix). An apparent

hybrid with C. bistorta was once collected, as discussed under that

species, and C, lewisii likewise occurs sympatrically with C. cheiranthi-
folia subsp. suffruticosa, C. proavita, C. micrantha (at Point Dume and

Playa del Rey, Los Angeles County, California), C. ignota, C. inter-

media, and C. robusta. No hybrids with these species have been ob-

served.

Camissonia lewisii is named in honor of Harlan Lewis, who intro-
duced me to the study of Onagraceae when I was a graduate student
at the University of California, Los Angeles. Judging from its mor-
phology and distribution, this species is certainly one of the parents
of the hexaploid C. robusta, which can, however, easily be distin-
guished by its more robust, erect habit, less sharply angled capsules,
and high proportion of 4- and 5-pored pollen grains. Camissonia
lewisii is probably not directly involved in the ancestry of any other
polyploid species.

30. Camissonia proavita Raven, sp. nov. FIGURE 38
Oenothera hirta var. typica sensu Munz, Bot. Gaz, 85: 262. 1928; pro parte.
Oenothera micrantha sensu Munz, N. Amer. FI. II. 5: 160. 1965; pro parte.

Herba annua sparsissime strigulosa, omnino eglandulosa, ad C.
micrantha similis, rosula densissime florens, ramis prostratis crassius-
culis ad 45 cm. longis tarde e rosula crescentibus. Folia conferta
lineares vel angustissime elliptica, 2-9 cm. longa, 0.2-0.55 cm. lata,
ad petiolum gradatissime attenuate, sparsissime denticulata; petiolus
ad 2 cm. longus. Inflorescentia densiore strigulosa, conferta, interdum
pilis paucis longioribus patentibus praedita. Hypanthium 1.2-1.8 mm.
longum, ostio 1-1.2 mm. diametro. Sepala 1.8-2.8 mm. longa, 0.8—1.2
mm. lata. Petala 2.6-3.5 mm. longa, 1.8-2.2 mm. lata, immaculata.
Filamenta staminum episepalorum 1.2-1.6 mm. longa, illa epipet-
alorum 0.5-1.3 mm. longa; antherae 0.6-0.8 mm. longae. Stylus 2-3.2
mm. longus; stigma 0.8-1 mm. diametro, sub anthesi antheris cir-
cumnexum. Capsula 15-20 mm. longa, 1.3-1.5 mm. crassa, tran-
sectione in sicco quadrangularis sed in vivo subquadrangularis,
angulis subrotundatis; capsulae rosulae rectae, illae ramorum in
spiram unam vel in spiras duas contortae. Semina 0.8-0.9 mm. longa,
0.5-0.6 mm. crassa. Chromosomatum numerus gameticus, n=7.
Autogama.

Tyre: Disturbed soil by roadside, with Orthocarpus purpurascens,
Eschseholzia, Brassica, near kilometer 201, 5.5 miles south of San
Vicente, 950 ft., Baja California, 29 March 1962, P. H. Raven 17020
(DS; isotypes, RSA, US).

Distrispution (Figure 36): Sandy flats and dunes along the beaches
or in the valleys up to 2,200 ft. alt., northern Baja California from
the vicinity of San Vicente to near Misién San Borjas.

278 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Figures 37-38.—Species of Camissonia sect. Holostigma X 1: 37, C. micrantha (Raven
18245, topotype, DS); C. proavita (Raven 17020, holotype, DS), showing only a portion
of the dense central tuft and one of the branches.

RAVEN—GENUS CAMISSONIA 279

ADDITIONAL SPECIMENS EXAMINED:

BAJA CALIFORNIA: 21 mi. 5. of Santo Tomas, Wiggins 4273 (DS, NY,
POM, US); San Telmo Plains, 7 mi. E. of San Telmo on road to Rancho San José
(Meling’s Ranch), Wiggins 9735A (DS); San Telmo, Brandegee in 1893 (UC);
Los Alisos, ca. 5 mi. 8. of Meling’s Ranch, ca. 750 m, Moran 11341 (SD); San
Quintin Bay, 1889, Palmer 656 (C, F, GH, NY, PH, US); 20.6 km. 5. of road to
San Quintin, R12397 (DS); near Rosario, Orcutt in 1886 (UC); 16 km. SE. of El
Rosario, R12493; sandy wash at junction of El Marmol and San Fernando roads,
25 mi. W. of El Mdérmol, Wiggins 4343 (DS); Rancho Ramona, 29°53’N. lat.,
115°06’ W. long., Huey in 1958 (SD); San Borjas, Brandegee in 1889 (UC).

Camissonia proavita is an autogamous species obviously closely
related to C. micrantha but easily distinguished by its very crowded
habit and subglabrous vestiture. Most collected plants have no side
branches and present a very dense cluster of 50 or more capsules
in the basal rosette when mature. The branches are thicker than in
any other member of the section, and the plant presents a very dis-
tinctive aspect at all stages of its life cycle. More than 200 miles sepa-
rate the northernmost station of this species from the southernmost
mainland locality of its close relative, C. micrantha.

The name ‘‘proavita’”’ (=ancestor) was assigned to this species
because of the hypothesis that it was one of the diploid ancestors of
tetraploid populations of Baja California assigned in the present revi-
sion to C. intermedia, the other being C. ignota. At present, it appears
more likely that all populations of C. intermedia have had a common
allotetraploid origin involving C. micrantha and C. hirtella, but
concrete evidence is not available to disprove the earlier hypothesis
completely.

Camissonia proavita occurs sympatrically with C. cheiranthifolia
subsp. suffruticosa, C. lewisii, C. hirtella, C. ignota, C. intermedia,
and C. robusta, but no natural hybrids have been observed. The
chromosome number of these species has been determined from 4
individuals, each from a different population, including the type col-
lection (see Appendix).

31. Camissonia micrantha (Hornem. ex Spreng.) Raven, Brittonia 16: 284. 1964.
FIGURE 37
Oenothera hirta Link, Enum. Hort. Berol. 1: 378. 1821; non Oenothera hirta
L. 1760. Type: Seeds from California, grown in the Berlin Botanical
Garden (authentic material, NY).
Oenothera micrantha Hornem. ex Spreng., Syst. 2: 228. 1825.
Holostigma micranthum (Hornem. ex Spreng.) * Spach, Nouv. Ann. Mus.
Paris 4: 335. 1835.
Sphaerostigma hirtum (Link) Fisch. & Mey., Ind. Sem. Hort. Petrop. 2: 22.
1835.
Sphaerostigma micranthum (Hornem. ex Spreng.) Walp., Rep. 2:77. 1843.
Oenothera hirta var. typica Munz, Bot. Gaz. 85: 261. 1928.

Annual with sprawling branches up to 0.6 m. long from a basal

rosette, or nfore rarely with a single erect stem. Plants more or less

280 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

densely villous all over. Leaves very narrowly elliptic in the basal
rosette, the upper ones proportionately broader near the base but
rarely broad enough to be called lanceolate, 1-12 cm. long, 0.2-1.7
mm. wide, denticulate, the apex acute, the base narrowly cuneate (in
the basal rosette) to rounded; petioles of lower leaves to 2 cm. long,
the upper leaves subsessile. Inflorescence more densely villous than
the rest of the plant, usually grayish, rarely with an understory of
short, glandular trichomes. Hypanthium 1.2—2 mm. long, 0.8-1.5 mm.
across at the summit, a few trichomes within near the base. Sepals
1-2.2 (—2.5) mm. long, 0.4-1 mm. wide. Petals 1.5-3.5 (—4.5) mm.
long, 0.8-2.2 (—2.5) mm. wide, occasionally with 1 or 2 red dots near
the base. Filaments of the episepalous stamens 0.8-1.5 mm. long, those
of the epipetalous ones 0.5-0.8 (—1) mm. long; anthers 0.4-0.6 mm.
long. Style 2-3.5 mm. long, with a few trichomes near the base;
stigma 0.5-0.6 mm. in diameter, surrounded by the anthers at anthesis.
Capsule 1.3-2 (-2.5) em. long, 1.1-1.2 (-1.8) mm. thick, subterete in
living material but drying more or less quadrangular in transection,
straight or curved in up to slightly more than one complete spiral.
Seeds 0.7-1.1 mm. long, 0.3-0.6 mm. thick. Gametic chromosome
number, n=7. Autogamous.

LecroryPeE: Grown in the Copenhagen Botanical Garden, probably
in 1821, from seeds doubtless collected in the vicinity of Fort Ross,
Sonoma County, California (probably at Bodega Bay) (C).

Disrripution (Figure 36): Endemic to California, where it grows
on coastal dunes and beaches or away from the ocean in sandy fields
and in washes, from the vicinity of Bodega Bay, Sonoma County,
and near Rio Vista, Sacramento County, south in the Coast Ranges
to the Los Angeles Basin and the northern edge of San Diego County;
also on San Miguel, Santa Rosa, Santa Cruz, and Santa Catalina
Islands. From sea level to about 1,000 ft. alt. very rarely up to about
2,500 ft. Also collected once, probably on ballast heaps, in the vicinity
of Nanaimo, Vancouver Island, British Columbia, 13 July 1898,
Macoun (BM). Rather widely grown in European botanical gardens
from the 1820s to the present, as a curiosity.

REPRESENTATIVE SPECIMENS EXAMINED:

U.S.: CALIPORNIA: CONTRA CosTA Co.: San Leandro, Edwards in 1877 (NY);
Strawberry Canyon, Berkeley Hills, ca. 1,300 ft., R8982 (CAS); sand hills E. of
Antioch, Heller 8892 (DS, F, GH, NY, PH, US, WT'U); near Byron, Heller 14496
(DS). rrEsNo co.: Los Gatos Canyon 4 mi, N. of Coalinga, Howell 5804 (CAS),
KERN CO.; Alee Cook Rocks, Cedar Canyon, 2,200 ft., Twisselmann 1194 (CAS).
LOS ANGELES CO.: Big Tujunga wash, Abrams 1390 (DS, POM); Pomona, Munz
et al. 2300 (DS, POM); Playa del Rey, Ewan 7536 (DS, LA, RM, UC); mouth of
Hamilton Canyon, Santa Catalina I., Fosberg $4315 (F, LAM, NY, PH, POM,
SBM, UC, US); Avalon, Santa Catalina I., Trask in 1898 (US). MARIN co.:
Near Radio Station, Pt. Reyes Peninsula, Howell 20956 (CAS, RSA). MoN-

RAVEN—GENUS CAMISSONIA Preal

TEREY co.: Near Del Monte, Heller 6659 (DS, F, GH, LE, NY, PH, POM, UC,
US); San Ardo, Hardham 10004 (DS); Pajaro Hills, Chandler 386 (UC); 20 mi.
from Carmel up Carmel R., road to Tassajara Springs, Wolf 3773 (UC); Pacific
Valley, Santa Lucia Mts., Plaskett in 1898 (CAS); Vineyard Canyon Road 12
mi. W. of Parkfield, Wiggins 8020 (DS, GH, UC); Los Laurelos Road, 8 mi. N. of
Carmel Valley, ca. 600 ft., Rose 57052 (COLO, RSA, US); Sycamore Canyon,
1 mi. S. of Big Sur State Park, Bristol 27 (Fresno State College herb.). ORANGE
co.: Bryant Ranch, Wolf 3778 (DS, RSA). RIVERSIDE co.: Elsinore, McC. in 1892
(DS); 1.4 mi. W. of junction of Hwy. 60 with Van Buren Ave., Wedberg 227 (San
Diego State College herb.). sAcRAMENTO co.: Brannan I. State Park, near Rio
Vista, R20177 (DS). sAN BENITO co.: About 4 mi. N. of Pinnacles Lodge, Baker
9282 (RSA). SAN BERNARDINO cCo.: Red Hill near Upland, Johnston 1196 (DS,
POM, UC, US). san pigGo co.: U.S. Hwy. 395, 2 mi. S. of turn to Lilac, R16950,
R17457 (both DS). san FRANcIsco co.: Fleishacker Pool, Abrams 12192 (DS;
basis for fig. 3434 in Abrams, Il. Fl. Pac. States 3: 208. 1951). SAN JOAQUIN CO.:
Corral Hollow, Hoover 3034 (UC). SAN LUIS oBIsPo co.: Baywood Park, Morro
Bay, Hardham 9072 (DS, RSA); 3 mi. E. of Santa Margarita, Linsley & Mac-
Swain 59-84 (RSA, UC); 10 mi. E. of Creston on La Panza Road, Hoover 7099
(RSA); Arroyo Grande, Brewer 446 (GH, UC, US). san Margo co.: San Pedro,
Elmer 4813 (ARIZ, CAS, DS, NY, OSC, POM, UC, US); Ano Nuevo Point,
Thomas 10178 (DS). SANTA BARBARA CO.: 5 mi. S. of Surf, Ferris 7543 (DS, POM,
UC); 5 mi. from Lompoc, Munz 9269 (C, POM); Fox Canyon, Santa Barbara,
Smith 1698 (RSA, WTU); 3 mi. 8. of Carpinteria, Lew7s 1298 (LA); San Miguel I.,
Blakley 5083 (DS, SBBG); Santa Rosa I., Water Canyon, Munz & Crow 11694
(GH, LA, OSC, POM, RSA); Santa Cruz L, vicinity of Pelican Bay, Abrams «&
Wiggins 41 (CAS, DS, GH, UC). santa cruz co.: Ben Lomond sand hills, Rose
33241 (WTU); ca. 3.5 mi. W. of Watsonville, Thomas 3104 (DS, RSA). sonoMA
co.: Bodega Point, Eastwood 4831 (CAS, GH, PH). venrura co.: Beach near
mouth of Ventura R., Pollard in 1947 (COLO, RSA).

Camissonia micrantha is a diploid, autogamous species which has
in the past been interpreted much more broadly (Munz, Bot. Gaz.
85: 260-265. 1928), including all species of this section except C.
cheiranthifolia, C. bistorta, and C. guadalupensis. As interpreted here,
it includes most of the material referred by Munz (loc. cit.) to Oeno-
thera hirta var. hirta, excluding mostly collections of the tetraploid
C. intermedia, which in part blurs the morphological distinctions
between its two diploid ancestors, C. micrantha and C. hairtella. The
chromosome number of (. micrantha has been determined in 47
individuals representing 39 populations from throughout the range of
the species (see Appendix).

Camissonia micrantha is most closely related to the allopatric
C. proavita, as discussed under that species. It is one of the diploid
ancestors not only of the tetraploid C. intermedia, as just mentioned,
but has probably combined with this tetraploid to give rise to the hexa-
ploid C. hardhamiae. Camissonia micrantha occurs sympatrically with
C. cheiranthifolia subsp. cheiranthifolia, C. bistorta, C. lewisii, C. hirtella,
C’. intermedia, C. confusa, C. hardhamiae, C. luciae, and C. robusta, but
no hybrids with any of these species have been observed.

282 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

The populations of C. micrantha growing on the coastal dunes and
beaches of northern Marin and southern Sonoma Counties consist of
plants that are more robust and larger flowered than those found farther
south. Such plants match the type material very closely and also are
identical with plants grown from seed sent from the Copenhagen
Botanical Garden and grown at Stanford (R18242). These seeds may
have been derived from direct descendants of the original introductions,
the species apparently having been in cultivation in Europe continu-
ously since the 1820s.

32. Camissonia hirtella (Greene) Raven, Brittonia 16: 284. 1964.
Fiaures 39, 40

Oenothera hirtella Greene, Fl. Francise. 215. 1891.

Sphaerostigma hirtellum (Greene) Small, Bull. Torrey Cl. 23: 190. 1896.

Sphaerostigma arenicola A. Nels. (‘‘urenicolum’’), Bot. Gaz. 40: 58. 1905.
Type: Tassajara Hot Springs, Santa Lucia Mts., Monterey Co., Cailif.,
June 1901, A. D. EF. Elmer 3192 (DS: isotype, US).

Oenothera hirta var. jonesii Il. Lév., Monogr. Onoth. 213. 1905. Lectotype:
Drytown, Amador Co., Calif., 500 ft., G. Hansen 543 (DS; isotypes, NY,
POM, UC. Cf. A. Nelson, Bot. Gaz. 40: 59. 1905).

Sphaerostigma micranthum var. jonesii (H. Lév.) A. Nels., Bot. Gaz. 40: 59.
1905.

Sphaerostigma bistortum var. reedii Parish, Muhlenbergia 3: 60. 1907. Lectotype:
Waterman Road, 3,000 or 3,500 ft., San Bernardino Mts., San Bernardino
Co., Calif., S. B. Parish 5794 (isotypes, GH, NY, UC).

Sphaerostigma hirtellum var. montanum Davidson, Muhlenbergia 3: 108.
1907.

Oenothera micrantha var. hirtella (Greene) Jeps., Man. Fl. Pl. Calif. 684. 1925.

Oenothera micrantha var. reedii (Parish) Jeps., Man. Fl. Pl. Calif. 684. 1925.

Ocenothera hirta var. jonesii f. reedii (Parish) Munz, Bot. Gaz. 85: 263. 1928.

Oenothera micrantha var. jonesii (H. Lév.) Munz, Amer. Journ. Bot. 19: 778.
1932.

Erect annual, generally similar to C. micrantha, with one or more
ascending branches from the basal rosette. Upper leaves narrowly
ovate or ovate (rarely lanceolate), the apex acute, the base cordate
to truncate. Inflorescence almost always with an understory of short
glandular trichomes. Hypanthium 1-3 mm. long, 1-3 mm. across at
the summit. Sepals 2.5-6 mm. long, 0.6-2 mm. wide. Petals 2-9 mm.
long, 1.3-7 mm. wide, sometimes red-dotted near the base, occa-
sionally with a salient tooth arising from the emarginate apex. Fila-
ments of the episepalous stamens 1.2-6 mm. long, those of the
epipetalous ones 0.5-3 mm. long; anthers 0.4-1 mm. long. Style 2-8
mm. long; stigma 0.5-1 mm. in diameter, surrounded by the anthers
at anthesis. Capsule 0.75-0.9 mm. thick, terete, once or twice con-
torted. Gametic chromosome number, n=7. Autogamous, rarely
cleistogamous.

LecrotyrE: Mt. Hamilton, Santa Clara County, California, July
1891, Lf. L. Greene (ND).

Disrripution (Figure 42): On brushy hills and slopes, often locally
abundant on burns, from T rinity and Amador Counties southward

RAVEN—GENUS CAMISSONIA 283

Ficures 39-41,—Species of Camissonia sect. Holostigma, X 1: 39-40, C. hirtella: 39, small
flowered plant (Raven 17473, DS); 40, large-flowered plant (Raven 17416, DS). 41 C.
robusta (Raven 17043, DS).

284 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

O LT
° fe)
° fe)
om. O
&
fe)
O50 6
BQ &
&
It ob
e
08 8 0
<C)
C)
C) ° % A
ee
oO Os “
e Og oho
beet rN
° og De
20 os &N\)o
EP °S'8 0 a ° ,>
on Coban 6 Aad
“<at> $s 4 a,
@ o 06
fe)
« yD ¢ o8Se
OgsO
é\
\ oot
O Cras
Oo fm
&
NI
of
fe)
fe
O

NN
Ficure 42,—Portion of the southwestern United States and northwestern Mexico, showing
the ranges of species of Camissonia sect. Holostigma: Q=C. hirtella, with @=diploid

chromosome counts (n=7); A=C. pallida subsp. pallida, with A=diploid counts
(n=7).

in the Coast Ranges and Sierra Nevada of California to the Sierra
Judrez and Sierra San Pedro Martir (south to ca. 30°45’ N. lat.) of
Baja California, as well as the hills to their west; usually away from
the immediate coast and barely reaching the margins of the desert.
From near sea level to 7,500 ft. alt.

REPRESENTATIVE SPECIMENS EXAMINED:

U.S.: CALIFORNIA: AMADOR Co.: 6 mi. N. of Jones Butte, 450 ft., Roseberry
121 (RSA, UC); Tone, Hoover 2416 (RSA, UC). caLtaveras co.: Cosumnes R.,
Rattan 6420; Mokelumne Hill, Blaisdell (CAS, GH, NY, US). eLpoRapo co.:
Coloma, Eastwood 14168 (CAS). FresNo co.: Tehipite Valley, Hall & Chandler 508

RAVEN—GENUS CAMISSONIA 285

(UC). GLENN co.: Grade ca. 10 mi. E. of Alder Springs, Heller 11443 (CAS, DS,
F, GH, NY, OSC, UC, WTU). Kern co.: W. end of Teyuca Ridge, 6,800 ft.
Twisselmann 9576 (DS); Breckenridge Mt., 7,400 ft., Twisselmann 4704 (CAS,
LA); 1 mi. E. of Havilah, 3,600 ft., R17569 (DS); Erskine Creek, Purpus 5099
(GH, UC, US); 8-10 mi. E. of Caliente, Thorne 31714 (RSA); 5 mi. E. of Alta
Sierra, ca. 5,000 ft., Greenhorn Mts., Thorne 31772 (RSA). LAKE co.: W. base of
Snow Mt., Ackley in 1965 (CAS); foot of Mt. Sanhedrin, Reynolds (CAS); Kelsey
Creek, Kelseyville, Benson 2178 (POM); 2 mi. W. of Iloughs Springs, Abrams
12483 (DS); Cobb Mt. to Adams Springs on the Binkley Ranch, Jussel 154
(CAS, POM, UC). Los ANGELES co.: Mandeville Canyon, Santa Monica Mts.,
Anderson & Epling in 1930 (F, LA, US); San Dimas Canyon, 1,500 ft. Munz &
Harwood 3686 (POM, UC, WTUS); Soledad Canyon between Acton and Ravenna,
Craig 480 (POM); Bear Canyon Trail to Mt. San Antonio, Crazg 415 (POM, UC).
MADERA Co.: State Hwy. 41, 8. of junction with Bass Lake road, Wedberg 441
(San Diego State College herb.). Marin co.: Mill Valley, Bioletti in 1892 (RM).
MARIPOSA co.: Mt. Bullion, 2,300 ft., Jepson 10722 (JEPS). MENDOCINO Co.:
Anderson Valley, Purdy in 1882 (GH); summit on road between Ukiah and
Boonville, 2,300 ft., Swensen 39 (UC). MONTEREY Co.: 3 mi. up Salinas R. from
Monterey, Peirson 3793 (DS, RSA); Pajaro Hills, Chandler 436 (UC); Hanging
Valley, Arroyo Seco Road, 3,000 ft., Hardham 10871 (DS, RSA); between Junipero
Serra and Pinyon Peaks, 4,700 ft., Howell 30177 (CAS); Big Sur, Baker 7863
(CAS). napa co.: Ridge W. of Oakville, ca. 500 ft., Reed 325 (UC, WTU); near
Knoxville, Jepson in 1892 (UC); La Jota, Howell Mt., Jepson 17569 (JEPS).
ORANGE Co.: Glen Ivy Trail to Santiago Peak, 4,000 ft., Munz 7064 (POM, UC).
RIVERSIDE Co.: Junction Main Divide Truck Trail with Ortega Hwy., 2,300 ft.,
Santa Ana Mts., R17762 (DS); Chalk Hill, 5,000 ft., San Jacinto Mts., Hall 2089
(UC); W. of Lake Elsinore, 1,600 ft., Munz 5086 (POM). sAn BENITO co.: Near
Hernandez, Dudley in 1899 (DS); Chalone Creek, Pinnacles National Monument,
Burgess 238 (UC); brushy slopes of San Benito Peak above New Idria Mine,
4,000 ft., Lyon 919 (UC). SAN BERNARDINO Co.: Devore, mouth of Cajon Canyon,
600 m, Munz 11054 (C, GH, POM, UC, US); near Big Bear L., 6,000 ft., Goodman
& Hitchcock 1743 (NY); Fredalba, Abrams 2791 (DS, GH, LE, NMC, NY, PH,
US). san pieGo co.: Above Henshaw Dam, Palomar Mts., 3,500 Munz 8334
(GH, NY, POM); Cuyamaca L., 4,700 ft., Munz & Harwood 7276 (POM);
Laguna Junction, Epling ct al. in 1932 (DS, LA, UC); 10 mi. W. of Campo,
3,000 ft., Munz 12633 (POM). SAN LuIs oBIsPo Co.: 4.5 mi. NE. of Santa Margarita,
R20158 (DS); Bryson, Santa Lucia Mts., Hardham 10728 (DS); Arroyo Grande,
Brewer 443 (US); McChesney Peak, 3,500-4,000 ft., Hardham 11040 (DS). sanra
BARBARA co.: Junction of Zaca Ridge and Catway roads, Figueroa Mt., San
Rafael Mts., 4,200 ft., Blakley 48300 (CAS, RSA, SBEG); Carpinteria, Jones in
1929 (POM); S. slope of Santa Ynez Peak, Santa Ynez Mts., 3,700 ft., Breedlove
2696 (CAS); Cuyamaca Valley 55 mi. E. of Santa Maria, Munz 11417 (POM);
Pelican Bay, Santa Cruz I., Clokey 5010 (DS, GH, NY, POM, SD, US). santa
CLARA Co.: Sierra Azul Ridge near Loma Prieta, 3,600 ft., Hichborn in 1920 (DS) ;
Rattan’s Ranch, near San Jose, Rattan in 1885 (DS, RSA); Colorado Creek,
Red Mts., 2,500 ft., Mt. Hamilton Range, Sharsmith 3800 (UC). SANTA CRUZ
co.: 1 mi. up Boulder Creck from town of Boulder Creek, Thomas 1780 (DS,
OSC, RSA); Ben Lomond Mt., 2,500 ft., Thomas 3471 (DS, OSC, RSA), soLaNno
co.: Summit Vaca Mts. via Collins Camp, Jepson 14482 (JEPS). sonoMA Co.:
Petrified Forest, Eastwood 4618 (CAS). surrer co.: North Butte, Marysville
Buttes, Bacigalupi 2241 (DS). reHama co.: W. of Paskenta along logging road,
Wagnon 12659 (RSA). TRINITY co.: The Knob, near Deer Lick Springs, Azildale
in 1931 (DS). TULARE co.: Road to Mineral King, ca. 5,000 ft., Baker 4195a

286 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

(DS). TUOLUMNE co.: South Fork Bridge, Tuolumne R., Grant 843 (JEPS).
VENTURA Co.: Upper Sespe Creek, N. of Wheelers Hot Springs, 3,500 ft., Munz
13189 (POM, UC); Goodenough Meadow, Mt. Pinos region, Dudley & Lamb
4698 (DS); between Nordhoff and Big Chief peaks, ca. 4,000 ft., Pollard in 1945
(CAS); burn near Kennedy Canyon, Pollard in 1946 (ARIZ, CAS, SBM).

BAJA CALIFORNIA. 18 mi. SE. of Tecate, Munz 9516 (POM); 50 mi. SE.
of Tecate, Munz 9558 (POM, US); 9 mi. W. of La Rumorosa, 4,020 ft., Hevly
& Pitman 2053 (ARIZ); 3.5 mi. FE. of Rancho San José (Meling’s Ranch), Wig-
gins 9787 (DS, UC, US); 19.5 km. SE. of San Vicente, 400 m, R12285 (DS);
10 mi. N. of La Huerta, pine-oak belt, 4,000 ft., Weggins 11845 (DS, OSC, SD,
UC); 1 mi. NE. of Santa Catarina, 64 mi. SE. of Ensenada, ca. 3,875 ft., Broder
683a (DS, UCSB); Los Emes, 8. Sierra San Pedro Martir, 3,500 ft., Wiggins
9903 (DS, UC, US).

As I conceive it, this taxon includes most of the populations gen-
erally referred to Oenothera micrantha var. jonesii (H. Lév.) Munz.
As discussed under Camissonia micrantha, the tetraploid C. intermedia
in part bridges the morphological hiatus between its two putative
diploid parents, (. micrantha and (. hirtella, and thus renders the
separation of all three taxa difficult. The chromosome number of
Camissonia hirtella was determined in 84 individuals derived from 76
populations scattered throughout the range of this species (see
Appendix).

This widely distributed species is extremely variable in flower size,
some individuals having petals up to 9 mm. long. Even in a collection
with such large flowers, like R19076 (grown from R17402) from San
Diego County, California, the longer set of anthers surrounds the
stigma and deposits pollen directly on it, and the plants are fully
self-compatible, setting a full complement of seed even when pro-
tected from insects. Insects are in general very rare at the flowers of
this species.

In addition to being one of the diploid parents of the tetraploid
(’. intermedia, Camissonia hirtella has very probably combined with
the diploid C. pallida subsp. pallida to give rise to the tetraploid C,
confusa, and with one of these two tetraploids to give rise to the
hexaploid C. luciae. Camissonia hirtella occurs sympatrically with
C. bistorta, C. micrantha, C. ignota, C. pallida subsp. pallida, (.
intermedia, C. confusa, C. hardhamiae, C. luciae, and C. robusta.

At times it is difficult to distinguish relatively narrow-leaved strains
of this species from (. intermedia. For example, some populations
from the central Sierra Nevada foothills of Calaveras County, Calli-
fornia (e.g., Quick 52-01, CAS, 0.5 mile from Parrots Ferry bridge
on road to Vallecito), temporarily referred here, might actually prove
to be C. intermedia if their chromosome number could be determined.
Similarly, Heller 7337 (DS, F, GH, LE, NY, OSC, PH, RM, UC,
US), a widely distributed gathering from the foothills west of Los
Gatos, Santa Clara County, California, might belong to either species.

RAVEN—GENUS CAMISSONIA 287
Indeed, the existence of the tetraploid populations here referred to
(. intermedia was the primary reason that plants as different as C.
micrantha and C. hirtella could ever have been referred to a single
species, Serious difficulty likewise exists in separating some of the
large-flowered, southern California populations of (. hirtella from the
geographically much more restricted tetraploid C. confusa, Camissonia
confusa bridges the morphological hiatus between C. hirtella and C.
pallida subsp. pallida in the same way that the other tetraploid
known in the section, (. intermedia, bridges the gap between (.
hirtella and C. micrantha. As C. hirtella is sometimes more similar to
(. pallida subsp. pallida than it ever is to C. micrantha, the taxo-
nomic problem can be even more serious, although the vast majority
of populations can be assigned to one species or the other without
difficulty. The three hexaploid species in the section can readily be
separated from C. hirtella on the basis of the proportion of 4- and
5-pored pollen grains they possess.

A single plant that was apparently a natural F, hybrid between
(. hirtella and C. ignota was found with the two parents in Lyons
Valley, 8 miles northeast of Jamul, San Diego County, California.
This plant had 6 pairs of chromosomes and 2 univalents in 2 cells
and 5 pairs plus 4 univalents in 3 cells at diakinesis. It appeared to
have set a full complement of seeds.

33. Camissonia ignota (Jeps.) Raven, Brittonia 16: 284. 1964. FIGuRE 45
Oenothera micrantha var. ignota Jeps., Man. Fl, Pl. Calif. 684. 1925.
Oenothera hirta var. ignota (Jeps.) Munz, Bot. Gaz. 85: 263. 1928.

Ocenothera ignota (Jeps.) Munz, N. Amer. FI. IT. 5: 161. 1965.

Erect, often subsucculent and reddish-tinged annual, similar to
C. mierantha, with one or more ascending branches from the basal
rosette; plants to 55 cm. tall, very finely strigulose with a sparse
admixture of longer trichomes. Upper leaves very narrowly to nar-
rowly elliptic, to 6 cm. long and 1.4 em. wide, finely serrulate, the
apex acute, the base attenuate; almost all leaves evidently petiolate,
the petiole up to 2.5 cm. long. Bracts truncate at base, narrowly
lanceolate or lanceolate. Inflorescence subglabrous or glandular-
pubescent with a few longer erect trichomes, rarely entirely strigulose.
Hypanthium (1.1-) 1.8-3 mm. long, 1.1-2 mm. across at the summit,
elabrous within. Sepals 2.6-5.5 mm. long, 1-1.7 mm. wide. Petals
(3-) 4-8 mm. long, (2—-) 3-6 mm. wide, sometimes red-dotted near the
base. Filaments of the episepalous stamens (1.2-) 2.5-3.6 mm. long,
those of the epipetalous ones (1-) 1.3-2 mm. long; anthers (0.6)
0.8-1.6 mm. long. Style (3-) 4.5-7 mm. long; stigma 0.8-1.2 mm. in
diameter, surrounded by the anthers at anthesis. Capsule mostly 2-3
em. long, 0.8-1 mm. thick, terete in living material, very slender and
usually much contorted, coiled irregularly up to five times, or more

a)

ISS CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Ficures 43-45.—Species of Camissonia sect. Holostigma, X 1:43, C. pallida subsp. pallida
(Raven 13995, DS); 44, C. pallida subsp. hallit (Alexander & Kellogg 2072, DS); 45, C.
ignota (Raven et al. 12524, DS).

RAVEN—GENUS CAMISSONIA 289
rarely simply flexuous. Seeds 1.2-1.3 mm. long, 0.5-0.6 mm. thick.
Gametic chromosome number, n=7. Autogamous.

Tyrer: Jurupa Hills, west of Riverside, Riverside County, Cali-
fornia, April 1905, Mrs. C. M. Wilder 90 (UC 69875).

DistrisuTion (Figure 46): Most common in clay fields and slopes
at low elevations, but occasional on sandy soil and higher in the
mountains in the Coast Ranges ‘and bordering valleys from Yolo
County, California, south to the southern end of the Sierra San Miguel,
at about 30° N. lat., in Baja California, usually away from the im-
mediate coast and barely reaching the margins of the desert; Santa
Cruz Island, Santa Barbara County, California; collected once in the
lowest foothills of the Sierra Nevada in Madera County, California.

a
@0 re
Kom .
's 1)
yy e
i) en
\ ree
Oe
1e)
e
e
©)
oO

Nan

Ficure 46.—Portion of the southwestern United States and northwestern Mexico, showing
the ranges of species of Camissonia: CQ=C. ignota, with @=diploid chromosome
counts (n=7); A=C. kernensis subsp. gilmanti; A=C. kernensts subsp. kernensis.

290 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

From about 400 to 3,500 ft. alt. Most abundant in western Riverside
and San Diego Counties, California, and generally local and sporadic
elsewhere.

REPRESENTATIVE SPECIMENS EXAMINED:

U.S.: CALIFORNIA: FRESNO co.: Alcalde, Eastwood 13549 (CAS). Los ANGELES
co.: 1 mi. W. of Acton, Soledad Canyon, Lewis in 1954 (LA); 3 mi. 8. of Boquet
Canyon Guard Station, Tavares & Brinson 263 (DAO, RM, SD, UC); San
Fernando wash, Eastwood in 1913 (CAS); Tujunga Canyon, 1,500 ft., Pecrson 414
(JEPS, RSA); Griffith Park, Davidson 1451 (LAM); San Dimas Canyon, 1,500
ft., Harwood 3692 (POM); Johnson’s Pasture, 3 mi. N. of Claremont, Jones in
1928 (CAS). MADERA co.: San Joaquin R. ca. 5 mi. above Pollasky, Heller 8166
(CAS, DS, F, GH, NY, PH, UC, US). MErcep co.: Ortigalita Peak, San Carlos
Range, Lyon 1529 (RM, UC, WTU); Pacheco Pass, Eastwood 14092 (CAS).
MONTEREY CO.: Lowes Canyon, Hardham 3317 (CAS, LA). ORANGE co.: Trail to
Santiago Peak from Trabuco Canyon, in Holy Jim Canyon, 3,500 ft., Peirson
3505 (RSA); 2.3 mi. above gate on Silverado Canyon Truck Trail, 2,500 ft.,
R17750 (DS); Anaheim plains, King in 1908 (JEPS). riveRsIDE co.: Temescal
Canyon, ca. 12 mi. SE. of Corona, Hitchcock & Muhlick 22151 (DS, RM, UC,
UT, WTU); Elsinore, McClatchie in 1892 (DS, NY); Temecula, Cleveland in 1882
(SD); Coldwater Trail, Santa Ana Mts., Pequegnat 2 (LA); Rattlesnake Mts.,
Wilder in 1905 (DS); Saunders Meadow, Meyer 161 (JEPS). sAN BENITO CO.:
Pinnacles, Sutliffe in 1920 (CAS). SAN BERNARDINO CO.: Vicinity of San Bernar-
dino, Parish 4756 (NY, US); near Colton, Parish 78 (DS, UC). san pIEGo co.:
Moro Hills near Fallbrook, Abrams 3326 (CAS, DS, GH, LA, LE, NY, PH, POM,
UC, US); 2.3 mi. S. of Rincon Springs on road to Escondido, 1,100 ft., R17402
(DS); 1 mi. E. of Oak Grove, 2,700 ft., R17370 (DS); 2 mi. W. of Banner, Gander
173.3 (SD); Barrett, R16936 (DS); 1.1 mi. E. of Campo, R17445 (DS); Jacumba,
Gander 181.6 (SD). SAN LUIS OBISPO co.: Upper Navajo Creek, La Panza Range,
Hoover 6904 (CAS, RSA); Cuyama Valley 44 mi. W. of Maricopa, 2,000 ft.
Benson 5787 (POM). SANTA BARBARA CO.: Santa Ynez Mts., Eastwood in 1904
(CAS); Valley Anchorage, Santa Cruz I., Hoffmann in 1929 (POM). STANISLAUS
co.: Near head of Del Puerto Canyon, Hoover 4883 (UC). ventura co.: Burn
near Kennedy Canyon, Ojai Valley, Pollard in 1946 (RSA). yoLo co.: Grade on
Rumsey to Arbuckle road, Hoover 3199 (POM, RSA, UC, US).

Camissonia ignota as here constituted is, in general, a very distinc-
tive entity that includes most of the populations referred by Munz
(Bot. Gaz. 85: 263. 1928) to Oenothera hirta var. ignota and later to
Oenothera micrantha var. ignota. Many of the populations here con-
sidered to belong to the tetraploid C. confusa were considered by
Munz to belong to this taxon also. These differ in their more robust
habit, broader leaves, abundant gray pubescence, less contorted
capsules, and tetraploid chromosome number. At times, however, the
separation of C. confusa from C. ignota can be very difficult, and it is
not impossible that C. ignota has contributed to the origin of some of
the tetraploid populations IT have included with C. confusa. In general,
however, it appears more probable that the diploid parents of the
latter species are C. hirtella and C. pallida subsp. pallida. It does not
appear likely that C. ignota has contributed to the formation of any

RAVEN—GENUS CAMISSONIA 291

other polyploid species, although at one time I thought that the
tetraploid Baja California populations here referred to C. intermedia
might have been derived from the diploids C. proavita and C. ignota
(see discussion under C. intermedia). The chromosome number of
Camissonia ignota has been determined in 22 individuals taken from
18 populations from throughout the range (see Appendix).

Camissonia ignota has been observed growing sympatrically with
C. bistorta, C. lewisii, C. proavita, C. hirtella, C. intermedia, and C.
confusa. A single F; hybrid with C. hirtella was found in San Diego
County, California, and is discussed under that species.

34. Camissonia pallida (Abrams) Raven, Brittonia 16: 284. 1964.

Similar to C. micrantha, but covered with dense, strigose pubescence.
Leaves entire or very sparsely denticulate, the upper ones undulate,
mostly 1-3 em. long, 0.35-0.7 cm. wide, with a short but distinct
petiole up to 2 mm. long. Inflorescence occasionally with an admixture
of glandular trichomes, usually conspicuously gray-pubescent. Hy-
panthium (1-) 1.8-4 mm. long, (0.7-) 1.2-3.5 mm. across at the
summit. Sepals (1.5-) 2.5-8 mm. long, 0.6-3 mm. wide. Petals (2-)
3.5-13 mm. long, (1.3-) 3-14.5 mm. wide, occasionally with 1-3 red
dots near the base. Filaments of the episepalous stamens (0.5-)
1.5-6.5 mm. long, those of the epipetalous ones (0.2—) 0.5-3.8 mm.
long; anthers (0.4—) 0.8-2.2 mm. long. Style (2-) 3-10.5 mm. long;
stigma 0.6-1.5 mm. in diameter, surrounded by at least the anthers
of the longer stamens and often by both sets of anthers at anthesis.
Capsule 1-1.2 mm. thick, more or less quadrangular in transection,
straight or once or more coiled, rarely up to three times. Seeds 1—1.45
mm. long, 0.6-0.7 mm. thick. Gametic chromosome number, n=7.
Autogamous.

DistTRIBUTION (Figures 33, 42): Desert slopes and flats, often along
washes, from the head of the San Joaquin Valley in Ventura and
Kern Counties, California, across the Mojave and Colorado Deserts,
north to the vicinity of Independence, Inyo County, California, and
Esmeralda County, Nevada (the only member of the section in
Nevada); east to Mohave and Yavapai Counties, Arizona, and also
in the vicinity of Tucson, Pima County; and south along the eastern
side of Baja California to the Sierra de San Borjas at about 29° N.
lat.; from about 100-6,000 ft. alt.

Camissonia pallida consists of two subspecies: subsp. alli, which
is relatively large-flowered, and subsp. pallida. The former occupies
a much more restricted range than the latter, and often grows with
it. Numerous intermediate individuals are found in such localities,
and the plants appear to differ only in flower size. No meiotic disturb-
ances or evidences of sterility have been found in individuals with in-
termediate-sized flowers. Presumably the distinctness of these two

292 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

species is maintained by their predominant autogamy. Camissonia
pallida subsp. hallii was considered a variety of Oenothera bistorta
(=Camissonia bistorta) by Munz (Bot. Gaz. 85: 267. 1928). Despite
their similarity in flower size, however, C. bistorta is self-incompatible
and the stigma is elevated above the anthers at anthesis, whereas C.
pallida subsp. hallii is autogamous, the anthers shedding pollen di-
rectly on the stigma at anthesis. Vegetatively, these two entities are
sharply distinct.
34a. Camissonia pallida subsp. hallii (Davids.) Raven, Brittonia 16: 284. 1964.
Figure 44
Sphaerostigma hallit Davids., Muhlenbergia 3: 107. 1907.
Oenothera bistorta var. hallii (Davids.) Jeps., Man. Fl. Pl. Calif. 685. 1925.
Oenothera hallit (Davids.) Munz, N. Amer. FI. II. 5: 160. 1965.

Hypanthium 3.8-4.2 mm. long, 2.2-3.5 mm. across at the summit.
Sepals 4.8-8 mm. long, 1.2-3 mm. wide. Petals 6.5-13 mm. long,
5-14.5 wide, with 1-3 red dots near the base of each one. Filaments
of the episepalous stamens 3-6.5 mm. long, those of the epipetalous
ones 1.83.8 mm. long; anthers 1.5-2.2 mm. long. Style 6.5-10.5 mm.
long, the longer stamens just equal to and their anthers shedding
pollen directly on the stigma at anthesis. Gametic chromosome
number, n=7. Autogamous, but clearly facultatively outcrossing.

Type: Banning, Riverside County, California, 17 April 1897,
H. M. Hall 446 (UC).

Distripution (Figure 33): Sandy washes in Riverside and San
Bernardino Counties, California, ca. 100-4,300 ft. alt., from Banning
east throughout the Little San Bernardino Mountains and_ their
northern slopes and southeast to the vicinity of Mecca and Box
Canyon.

REPRESENTATIVE SPECIMENS EXAMINED:

U.S: CALIFORNIA: RIVERSIDE Co.: Big Oaks Canyon, Banning, 4,000 ft., Cooper
1645 (RSA); Whitewater Canyon 2 mi. N. of Banning-Indio boulevard, Munz
15660 (CAS, DS, GH, POM, UC, US, WS, WTU); near Mecea, Winblad in 1937
(CAS); Box Canyon, Winblad in 1937 (CAS). SAN BERNARDINO Co.: Just S. of
Morongo Valley, 2,600 ft., R17532 (DS); E. of Quail Spring, Moran 844 (DS,
POM, UC).

Camissonia pallida subsp. hallii has not been observed growing
sympatrically with any other member of the section, except C. pallida
subsp. pallida, with which it intergrades. Its chromosome number has
been determined in 4 individuals from 3 different populations (see
Appendix). The combination Camissonia hallii, attributed to me by
Munz, N. Amer. Fl. II. 5: 160. 1965, has not been published elsewhere.

Relatively large-flowered plants similar to subsp. hallii are occasion-
ally found elsewhere in the area of subsp. pallida. One such collection
was made in the pinyon-juniper association of the Cerbat Mountains,

RAVEN—GENUS CAMISSONIA 293

Mohave County, Arizona, 5,000 ft. elevation, Jaeger in 1941 (POM);.
the petals of one plant are 11.5 mm. long and 8 mm. wide, and have red
dots near the base similar to those usually found in subsp. pallida.
One of the plants mounted on this sheet is typical of subsp. pallida,
with petals ca. 3.5 mm. long. It would be highly desirable to have more
material for study before extending the range of subsp. hallii to this
disjunct station.

34b. Camissonia pallida subsp. pallida FIGuRE 43

Sphaerostigma pallidum Abrams, Bull. Torrey Cl. 32: 539. 1905.

Sphaerostigma micranthum var. exfoliatum A. Nels., Bot. Gaz. 40: 59. 1905.
Lectotype: Colorado Desert, probably in San Diego Co., Calif., April 1889,
C. R. Orcutt.

Oenothera abr ymsi J. F. Macbride, Contr. Gray Herb. 65: 41. 1922.

Oenothera micrantha var. abramsii (J. F. Macbride) Jeps., Man. Fl. Pl. Calif.
684. 1925.

Oenothera hirta var. exfoliata (A. Nels.) Munz, Bot. Gaz. 85: 264. 1928.

Oenothera micrantha var. exfoliata (A. Nels.) Munz, Amer. Journ. Bot. 19:
778. 1932.

Hypanthium (1-) 1.8-3 mm. long, (0.7—) 1.2-2.5 mm. across at the
summit. Sepals (1.5-) 2.5-5.5 mm. long, 0.6-2 mm. wide. Petals (2-)
3.5-6 (-8) mm. long, (1.3-) 3-5 (-6.5) mm. wide, very rarely with red
dots at the base. Filaments of the episepalous stamens (0.5—-) 1.5-4 mm.
long, those of the epipetalous ones (0.2-) 0.5-2.2 mm. long; anthers
(0.4-) 0.8-1.2 mm. long. Style (2.1-) 3-6.5 mm. long, the stigma sur-
rounded at anthesis by both sets of anthers, or sometimes by only the
longer set. Gametic chromosome number, n=7. Autogamous.

Typr: Near the station, Cabazon, Riverside County, California,
6 April 1903, L. R. Abrams 3228 (NY; isotypes, CAS, DS, GH, LA,
LE, NY, RM, RSA, SBM, UC, US, WS, WTU).

DisTRIBUTION (Figure 42): That of the species.

REPRESENTATIVE SPECIMENS EXAMINED:

U.S.: ARIZONA: MOHAVE co.: 9 mi. NW. of Kingman, 3,300 ft., Munz et al.
92982 (DAO, RSA, UC). pima co.: Bear Canyon, Santa Catalina Mts., Mason
et al. in 1960. YAVAPAI co.: 5.4 mi. NE. of Congress, 4,000 ft., R17367 (DS);
Skull Valley, 4,300 ft., Jones in 1903 (POM). CaLiroRNIA: IMPERIAL CO.: 0.5
mi. E. of Mountain Springs, Ferris 7078 (DS, NY). 1Nyo co.: North Fork of
Oak Creek, Independence, 3,000 ft., Train in 1937 (ARIZ, DS, OSC); 2.5 mi. E.
of Coso Hot Springs, Coso Range, 3,550 ft., Alezander & Kellogg 2768 (UC).
KERN co.: 8. end of Soledad Mt., 2,775 ft., Twisselmann 4308 (CAS, LA); 9.7
mi. S. of Weldon, 3,100 ft., R17565 (DS); Poso Creek, slope on S. side, Greenhorn
Mts., ca. 2,600 ft., Smith 299 (JEPS). Los ANGELES CO.: N. of Pearblossom,
Wedberg 250 (DS); 2 mi. N. of Saugus, Munz 10013 (POM, UC); Rock Creek to
Big Pines, 4,500 ft., Pierson 7951 (RSA). RIVERSIDE CO.: Dos Palmos Spring,
N. end of Santa Rosa Mts., 3,600 ft., Munz 15107 (CAS, GH, POM, UC); San
Gorgonio Pass, 1 mi. W. of Whitewater, 1,200 ft., Wolf 8416 (ARIZ, GH, NY,
RSA, WTU); Palm Canyon near Palm Springs, Johnston 1110 (DS, POM, US);
Indio, McGregor in 1917 (DS); Fried Liver Wash, Pinto Basin, Joshua Tree

294. CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

National Monument, R11863 (DS); 5 mi. S. of Gold Coin Mine, Little San
Bernardino Mts., Epling & Robinson in 1933 (DS, LA, RSA). SAN BERNARDINO
co.: Cedar Canyon, Mid Hills, 5,000 ft., R17353 (DS); 3.5 mi. N. of Lake Arrow-
head, 4,550 ft., Axelrod 273 (UC); Cajon Pass, 3,800 ft., Jones in 1903 (POM);
Cactus Flat, San Bernardino Mts., 6,000 ft., Munz 10518 (POM); Granite Mts.,
N. of Amboy, Jaeger in 1989 (DS). san piEGO co.: Hell Hole Canyon, Borrego
Valley, Epling & Robinson in 1932 (DS, F, LA, NY, RSA, UC); extreme SE.
corner of the county, Gander in 1940 (SD); 11 mi. E. of Banner, Gould 2268
(RSA, UC); 5 mi. E. of Laguna Station, 1,090 m, Bracelin 5 (POM, UC). ventura
co.: Flood plain of Piru Creek, Hoffmann in 1930 (CAS, SBM); sandy field near
Camulos, Hoffmann in 1929 (SBM). NEVADA: ESMERALDA co.:2 mi. E, of Bell-
ville, Purpus in 1898 (UC).

BAJA CALIFORNIA. Vicinity of Japs, Orcutt 1191 (UC); 3 mi. W. of Santa
Catarina, 64 mi. SE. of Ensenada, 3,755 ft., Broder 659 (DS, UCSB); Rancho
San Matias, Sierra San Pedro Martir, ca. 1,050 m, Moran 10844 (DS, SD);
San Juan Mine, Sierra de San Borjas, ca. 1,200 m, Moran 8127 (RSA, SD, UC).

This extraordinarily variable taxon is at times difficult to separate
from C. hirtella where the ranges of these two species approach one
another along the margins of the desert. There is, however, no evi-
dence of genuine intergradation, and most plants are easily referred
to one species or the other. An even more difficult problem is the
separation of C. pallida subsp. pallida from C. confusa, which is
presumably an allotetraploid derived from the diploids C. pallida
subsp. pallida and C. hirtella. In California, the problem is rarely
a difficult one, but in Arizona further chromosome counts will clearly
be necessary to elucidate the situation completely. All collections
from the area of the Tonto National Forest in Gila and Maricopa
Counties appear to be C. confusa, and this is documented by chromo-
some counts from the area. Collections from Paradise Valley and
Avondale, farther west in Maricopa County, and the vicinity of
Sacaton and Casa Grande Ruins, to the south in Pinal County,
also are morphologically similar to the tetraploids of California.
Populations in Yavapai and Mohave Counties, on the other hand,
are known to be diploid. A series of populations from the vicinity
of Tucson and the Santa Catalina Mountains in Pima County,
Arizona, appears morphologically similar to the diploid and are
tentatively referred here, even though they would, if this disposition
is correct, represent a disjunct occurrence for C. pallida subsp. pallida.
Unfortunately none of these populations has been examined cyto-
logically.

The chromosome number of this subspecies has been determined
in 26 individuals from 24 populations scattered throughout the range
(see Appendix). Camissonia pallida subsp. pallida intergrades with
C. pallida subsp. hallii and occurs sympatrically with C. hirtella
and probably C. ignota (the latter in the vicinity of Santa Catarina,
Baja California).

RAVEN—GENUS CAMISSONIA 295

35. Camissonia intermedia Raven, sp. nov. Ficure 48
Oenothera hirta var. typica sensu Munz, Bot. Gaz. 85: 261, 1928; pro parte.
Oenothera hirta var. jgonesit sensu Munz, Bot. Gaz. 85: 262. 1928; pro parte.
Oenothera micrantha sensu Munz, N. Amer. FI. II. 5: 160. 1965; pro parte.

Herba annua dense villosa erecta, ad C. micrantha similis, usque ad
6 dm alta, ramis plurimis erectis e rosula centrali radiantibus, rariore
caule singulari erecto. Folia his C. micranthae basi parum latiora.
Inflorescentia plus minusve cano-villosa, persaepe pilis brevibus
glandulosis pubescentes. Usque ad partem vicensimam pollinis gran-
orum quadriporam, rariore ad partem dimidiam. Chromosomatum
numerus gameticus, n=14. Autogama.

Type: Common along disturbed roadsides in chaparral, with Adeno-
stoma fasciculatum and Arctostaphylos glauca, Mulholland Highway
2.6 miles east of Seminole Hot Springs, 1,800 ft. alt., Santa Monica
Mountains, Los Angeles County, California, 1 May 1962, P. H. Raven
17517 (DS 514938; isotypes, RSA, US).

DistriBuTION (Figure 50): Mostly on disturbed brushy slopes,
often common on burns, from Lake and Yolo Counties (where rare)
south in the Coast Ranges of California to the western San Gabriel
Mountains, western Riverside County, and San Diego County, and
south in Baja California to the south end of the Sierra San Miguel at
about 30° N. lat.; also on Santa Cruz and Santa Catalina Islands.
From (500-) 1,000-2,500 ft. alt. Most abundant in Monterey County
and from coastal Santa Barbara County to the Santa Monica Moun-
tains of Los Angeles and Ventura Counties, California, and again in
coastal Baja California.

REPRESENTATIVE SPECIMENS EXAMINED:

U.S.: CALIFORNIA: CONTRA cosTA co.: W. slope of Mt. Diablo, ca. 900 ft.
Rose 41229 (GH, NY, UC). uake co.: Blue Lakes, Baker 264a (DS). Los ANGELES
co.: Wilson’s Trail, Abrams 1505 (DS); foothills near Sherman, Braunton 65
(US); Puddingstone Canyon, Munz et al. 2444 (POM); Liveoak Canyon Dam,
San Gabriel Mts., 1,500 ft., Wheeler 2543 (LA); Mandeville Canyon, 1,200 ft.,
Santa Monica Mts., Templeton & Clokey 4458 (DAO, NY, POM, UC, US); Los
Alisos Canyon, Santa Monica Mts., Epling in 1931 (CAS, F, LA, UC, US);
Avalon Hill, Santa Catalina I., Knopf 428 (F). MonrEeREY co.: Priest Valley,
2,250 ft., Howell 39114 (CAS); limestone ridge above Posts, ca. 2,000 ft., Big Sur,
Hoffman 1180 (UC). Napa co.: Oakville to Glen Ellen, ca. 900 ft., R2860 (CAS);
Capell Valley, Kamb & Chisaki 1760 (UCSB). oRANGE co.: Santiago Peak Trail,
Abrams 1792 (NY); 5 mi. 8. of Laguna, Mason 2926 (UC). RIVERSIDE co.: 5 mi.
NE. of Murietta, 1,500 ft., Munz & Johnston 5354 (POM). san BENITO co.: 10.5
mi. from junction near Bitterwater on road to Hernandez, R9152 (CAS, LA,
RSA, UC); Bear Valley, Jepson 12246 (JEPS). san DIEGO co.: Near County line
on U.S. Hwy. 395, Gander in 1937 (SD); Jamul, 1,500 ft., Stokes in 1895 (DS).
SAN LUIS OBISPO co.: Chimney Ranch, North Road to Adelaida, Hardham 3347
(LA, SBBG); east School Canyon, San Luis Obispo, Condit in 1908 (UC). SANTA
BARBARA co.: N. of Lompoc, Chandler 2570 (DS); Los Feliz School at end of
Mission Canyon Road, 1,000 ft., Smith 920 (RSA); Gibraltar Road to El Camino

296 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Ficures 47-49.—Species of Camissonia sect. Holostigma, X 1: 47, C. hardhamiae (Raven
18315, holotype, DS); 48, C. intermedia (Breedlove 2305, DS); 49, C. luciae (Raven
18267, holotype, DS).

RAVEN—GENUS CAMISSONIA 297

Ficure 50.—Portion of the southwestern United States and northwestern Mexico, showing
the ranges of species of Camissonia: A=C. intermedia, with A= tetraploid chromosome
counts (n= 14); @=C. chamaenerioides; the dotted-line outline indicates the range of
the related, larger flowered, self-incompatible C. refracta.

Cielo, 2,000 ft., Hardham 10854 (DS); Sierra Blanca, Santa Cruz I., Wolf 2842
(POM, RSA); Coches Prietos Canyon to Alberts Anchorage, 400 ft., top of ridge,
Blakley 3314 (SBBG). SANTA CLARA co.: Evergreen, Burtt Davy 71 (UC); Loma
Prieta, Burtt Davy 665 (UC). SANTA cRUz co.: Near summit of Santa Cruz to
Los Gatos highway, Covel 323 (CAS). sTanisLAus co.: Near head of Del Puerto
Canyon, Hoover 4884 (RSA, UG). ventura co.: Santa Susanna Pass, Epling in
1926 (LA); Foster Park burn, Ojai Valley, Pollard in 1945 (CAS, RSA, SBM).
yoLo co.: Grade on Rumsey-Arbuckle road, Hoover 3195 (POM, RSA, UC).

BAJA CALIFORNIA: 10 mi. S. of Ensenada, Purer 7140 (SD); near San
Jacinto, Gander in 1939 (SD); near Refugio, Cronemiller 3083 (POM); 13.1 mi. 8.
of Santo Tomas, Turner et al. 2074 (DS); 9.3 mi. 8. of San Vicente, R17045 (DS);
2.5 mi. S. of Ejido México, R17039 (DS).

Camissonia intermedia is almost certainly an allotetraploid, its
two diploid ancestors being C. micrantha and C. hirtella. The diffi-
culties of distinguishing it from these two diploids have been dis-
cussed in the sections dealing with each. At one time, I thought
that the tetraploid populations of Baja California here referred to
C. intermedia might have been derived from the diploids C’. ignota
and C. proavita, as mentioned under the latter species. Evidence for

298 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

this view is the fact that C. micrantha, one of the parents of the
northern populations of C. intermedia, ranges no farther south than
northern San Diego County, California, where the main area of the
tetraploid stops also. No tetraploids have been found in central San
Diego County, despite intensive sampling, and the Baja California
populations range north only to Jamul in the southernmost part of
the county. In general, the plants found in Baja California seem to
be somewhat more slender and glabrous than the ones from farther
north; but this is far from an absolute difference, and I have found it
difficult to quantify. It might be possible to elucidate this problem
with a program of experimental hybridization, but at present it
appears best to retain all of these obviously closely related tetraploid
populations in a single species.

The chromosome number of this species has been determined in
60 individuals from 53 populations from throughout the range of the
species including the type (see Appendix). Camissonia intermedia
has been found growing sympatrically with C. bistorta, C. lewisii,
C. proavita, C. micrantha, C. hirtella, C. ignota, C. confusa, and C.
robusta. Individuals of this species were referred by Munz (Bot.
Gaz. 85: 260-265. 1928) to Oenothera hirta var. typica and O. hirta
var. jonesii, entities corresponding for the most part to the diploid
parents of C. intermedia.

36. Camissonia confusa Raven, sp. nov. FIGuRE 51
Oenothera hirta var. ignota sensu Munz, Bot. Gaz. 85: 263. 1928; pro parte.
Oenothera ignota sensu Munz, N. Amer. FI. II. 5: 161. 1965; pro parte.

Herba annua robusta dense cinerero-villosa rare strigosa erecta,
ad C. micrantha similis, usque ad 6 dm. alta, ramis plurimis erectis
e rosula centrali radiantibus, rariore caule singulari erecto. Folia
superiora lanceolata vel anguste ovata, 1-5 cm. longa, 0.4-2 cm. lata,
basi rotundata vel truncata, sparse et exigue denticulata, undulata,
apice longe acuminata. Inflorescentia glandulosa, rariore pilis longi-
oribus admixta vel strigosa, rarissime subglabra. Hypanthium (1.8—)
2-3.8 mm. longum, ostio 1.2-2 mm. diametro. Sepala (1.5-) 3.2-8.5
mm. longa, 1.2-1.5 mm. lata. Petala (2.5-) 5-10.5 mm. longa, (1.5-)
2.8-6 mm. lata, plerumque prope basin puncto rubro uno vel duo.
Filamenta staminum episepalorum (1.2-) 2.5-4.5 mm. longa, illa
epipetalorum (0.8—) 1.5-2.5 mm. longa; antherae (0.4-) 0.8-1.5 mm.
longae. Usque ad partem vicensimam pollinis granorum quadriporum.
Stylus (2.5-) 4.5-7.5 mm. longus; stigma 1-1.2 mm. diametro, sub
anthesi antheris saltem longioribus cireumnexum. Capsula 0.9-1.2
mm. crassa surrecta vel in spiras 1 vel 2 contorta. Semina 1.2—-1.35
mm. longa, 0.5-0.7 mm. crassa. Chromosomatum numerus gameticus,
n=14. Autogama.

299

CAMISSONIA

RAVEN—GENUS

Ficure 51.—Camissonia confusa (Raven 17388, DS), X 1.

300 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Typé: Common in coastal sage by edge of citrus orchard, along dirt
road 2.3 miles west of Mill Creek Ranger Station, San Bernardino
Mountains, 2,300 ft. alt., San Bernardino County, California, 28 April
1962, P. H. Raven 17466 (DS 515446; isotypes, RSA, US).

Distripution (Figure 52): Dry slopes away from the coast, from
the La Panza Range of central San Luis Obispo County south through
the Coast Ranges to the San Bernardino Mountains and southern
San Diego County, California; also in central Arizona (westernmost
Gila, northern Pinal, and Maricopa Counties); from about 1,100-
6,600 ft. alt.

REPRESENTATIVE SPECIMENS EXAMINED:

U.S.: ARIZONA: GILA co.: Tonto Hill, 690 m., Collom 265 (GH, NY); 12 mi. N.
of Roosevelt, 2,300 ft., Peebles & Smith 11503 (ARIZ); Jana Gordo Wash, 1,800

Se
XK
‘ke
jb ©)
rs
22
“be & De
oa \F Bop O |
0 eo
bh a 0 Oo 8
A @)
oe

Figure 52,—Portion of the southwestern United States and northwestern Mexico, showing
the ranges of species of Camtssonia sect. Holostigma: QO=C. confusa, with @=tetra-
ploid chromosome counts (n=14); X =C. hardhamiae; O=C. luciae, with @=hexa-
ploid chromosome counts (n=21); A=C. robusta, with A= hexaploid counts (n=21).

RAVEN—GENUS CAMISSONIA 301

ft., Wittbank 143 (US); Collom Camp, Mazatzal Mts., 1,200 m, Collom 268
(ARIZ, US). monave co.: Tempe, Gamong & Blaschke in 1892 (US); 12.1 mi.
SW. of Roosevelt on State Hwy. 88, 2,300 ft., Gregory 60B (DS, RSA, UC);
Oak Flats, mountains between Miami and Superior, A. & R. A. Nelson 1858 (K,
RM); 6 mi. SW. of Superior, Russell 11271 (OSC); Paradise Valley, 25 mi. N. of
Phoenix, Peebles ct al. 5102 (ARIZ, US); 25 mi. E. Scottsdale on Beeline Hwy.,
1,160 ft., Kennedy 98 (DS); on canyon road 12 mi. from Apache Lodge, Eastwood
17161 (CAS). pinaL co.: Superstition Mt., Beckett 10784 (ARIZ); Sacaton,
Peebles & Harrison 1759 (US); sandy washes of the Gila R. near Casa Grande
Ruins, A. & R. A. Nelson 1677 (RM). CALIFORNIA: ORANGE CO.: Lower San Juan
Campground, Bates 2318 (DS). RIVERSIDE co.: 3.6 mi. NE. of Aguanga, Vestal
63113.8 (DS); Main Divide Truck Trail at Ortega Hwy., Santa Ana Mts., 2,300
ft., R17763 (DS); ca. 4 mi. E. of Redlands, Klein et al. 265 (DS); 10 mi. 8. of
Hemet, Munz 10819 (POM, SBM); 18 mi. from Banning on road to Idyllwild,
5,000 ft., Munz 8142 (GH, POM). SAN BERNARDINO CO.: Oak Glen, Munz &
Hitchcock 10916 (GH, POM); San Antonio Canyon, San Gabriel Mts., 5,500 ft.,
Peirson 2706 (POM, RSA); N. of Upland, Breitung 15060 (DAO); foothills, San
Bernardino Mts., Parish in 1891 (NY). San pieGo co.: 3.1 mi. from U.S. Hwy. 80
on road to Morena L., 3,400 ft., R17439 (DS); Banner, Keck et al. 6089 (NY);
1 mi. E. of Warners Ranch, 3,400 ft., McCully 19 (POM); 6 mi. N. of Santa
Ysabel, Munz 9808 (POM).

Camissonia confusa is a rather rare allotetraploid species which can
readily be confused with certain populations of either of its presumed
diploid parents, C. hirtella and C. pallida subsp. pallida. The problems
of separating it from its two putative ancestors have been discussed
under the treatments of the diploids, and it should be reemphasized
at this point that further chromosome counts are badly needed from
Arizona. As mentioned in the treatment of C. ignota, many of the col-
lections of C. confusa have been confused with that diploid entity,
and it is possible that C. ignota may have participated in the origin of
some of the tetraploid populations here referred to C. confusa.

The chromosome number of C. confusa has been determined in 29
individuals from 22 populations, including the type collection (see
Appendix). It has been observed growing sympatrically with C.
bistorta, C. micrantha, C. hirtella, C. ignota and C. intermedia. North of
San Bernardino County, California, populations are very scattered,
but the following collection from Santa Barbara County may belong
here: canyon of Mono Creek, 1,600 ft., upper Santa Ynez River water-
shed, Chandler 1460 (SBBG). The plants represented in this collection
are immature, and the species is otherwise not known from Santa
Barbara County.

37. Camissonia hardhamiae Raven, sp. nov. FIGURE 47

Herba annua robusta villosa pilis glandulosis brevioribus admixta,
ad C. intermedia similis. Folia superiora lanceolata vel anguste ovata.
Hypanthium 1.7—2 mm. longum, ostio 1.3-1.5 mm. diametro. Sepala
1.8-3.2 mm. longa, 0.8-1.2 mm. lata. Petala 2-4 mm. longa, 1.8—3.5
mm. lata, immaculata. Filamenta staminum episepalorum 1.5—2 mm.

302 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

longa, illa epipetalorum 1-1.5 mm. longa; antherae ca. 0.7 mm. longae.
Pollinis granorum 70-80 per centum, 4— vel 5-porum. Stylus 3-4 mm.
longus; stigma 0.7-1 mm. diametro, sub anthesi antheris cireumnexum.,
Capsula 1.3-2.5 cm. longa, 1.3-1.6 mm. crassa surrecta vel in spiram
unam contorta. Semina ca. 1.2 mm. longa, 0.6-0.7 mm. crassa. Chromo-
somatum numerus gameticus, n=21. Autogama.

Type: Calf Canyon, 3.7 miles northeast of Santa Margarita, San
Luis Obispo County, California, 5 May 1963, P. H. Raven 18315 (DS
516765).

Distripution (Figure 52): Very local, known only from a few
localities in sandy soil in disturbed oak woodland, southernmost
Monterey to central San Luis Obispo County, California; ca. 800-ca.
2,000 ft. alt.

SPECIMENS, EXAMINED:

U.S.: Catirornia: Monterey co.: Sandy Valley, Big Sandy ca. 1.5 mi. NE. of
intersection with Indian Valley Road, Hardham 10035A (D8). SAN LUIS OBISPO CO.:
4.5 mi. NE. of Santa Margarita, Linsley & MacSwain 59-87 (RSA), 59-88 (RSA),
Thorp 62-33 (DS); 1.1 mi. N. of intersection with Parkhill Road on Calf Canyon
Road, Hardham 2293 (LA); 1 mi. E. of Red Kagle Mine, 12 mi. S. of Shandon,
McMillan 105 (CAS).

This local hexaploid species has almost certainly been derived
from the diploid C. micrantha and the tetraploid C. intermedia, both
of which grow in the area of C. hardhamiae. The hexaploid is quite
similar to C. micrantha, but it can easily be distinguished by an
examination of the pollen, C. micrantha having only an occasional
4-pored grain. Camissonia hardhamiae has broader leaves and thicker
capsules than C. micrantha and is generally more robust. Its chromo-
some number has been determined in single plants from 3 popula-
tions, including the type collection (see Appendix). It has been found
growing sympatrically with C. micrantha and C. hirtella.

Cammissonia hardhamiae is named in honor of Clare B. Hardham of
Paso Robles, California, enthusiastic student of the flora of the area
where it grows. Mrs. Hardham has contributed much to our knowledge
of the flora of her area through her numerous collections.

38. Camissonia luciae Raven, sp. nov. FIGURE 49
Oenothera hirta var. jonesii sensu Munz, Bot. Gaz. 85: 262. 1928; pro parte.
Oenothera micrantha sensu Munz, N. Amer. FI. II. 5: 160. 1965; pro parte.

Herba annua erecta robusta villosa eglandulosa, ad C. micrantha
similis. Folia superiora lanceolata, 1.3-5.5 em. longa, 1.2-2.5 em.
lata, basi rotundata vel truncata, apice acuminata, sessilia, sparse
denticulata. Inflorescentia plerumque cinereo-pubescens. Hypanthium
2-3 mm. longum, ostio 1.5-1.8 mm. diametro. Sepala 2.5-4.5 mm.
longa, 1-1.8 mm. lata. Petala 4-7 mm. longa, 2.2-5 mm. lata, saepe

RAVEN—GENUS CAMISSONIA 303

mucronata, plerumque prope basin puncto rubro uno. Filamenta
staminum episepalorum 1.2-6 mm. longa, illa epipetalorum 0.81.6
mm. lata; antherae 0.4-1 mm. longae. Pollinis granorum 25-60 per-
centum quadriporum. Stylus 3-6 mm. longus; stigma 1-1.2 mm. diame-
tro, sub anthesi antheris staminorum longiorum vel rarissime et longi-
orum et breviorum circumnexum. Capsula ca. 1.5-2 cm. longa, 1.3-2
mm, crassa, transectione quadrangularis exsiccatis, subteretis in statu
vivo non dessicata, in spiras 1.5-2+ contorta. Semina 1.3-1.5 mm.
longa, 0.7-0.8 mm. crassa. Chromosomatum numerus gameticus,
n=21. Autogama.

Type: Openings in Adenostoma chaparral, The Indians, 2,200 ft.,
Santa Lucia Mountains, Monterey County, California, 3 May 1963,
P. H. Raven 18267 (DS 545885).

Distripution (Figure 52): Locally common in openings in the
chaparral, Santa Lucia Mountains, Monterey County, California,
and scattered southward to San Benito, San Luis Obispo, and Santa
Barbara Counties; 1,000-4,500 ft. alt.

SPECIMENS EXAMINED:

U.S.: CALIFORNIA! MONTEREY CO., SANTA LUCIA MTs.: Hastings Reservation,
near Jamesburg, Hoover 2963 (POM, RSA, UC), 2964 (POM, RSA, UC), Lins-
dale 43 (CAS), 123 (COLO), in 1943 (CAS), Cook & Cook 842 (CAS, mix with
C. intermedia), 680 (CAS); Conejo Creek 2 mi. below Jamesburg, Bacigalupi
1143 (DS, POM); road to Jamesburg 0.8 mi. from main road up Carmel Valley,
R18226 (DS); The Indians, 2,200 ft., Hardham 5443a (CAS, RSA); trail to
Ventana Double Cone between Pat Springs and Puerto Suelo, 4,000 ft., Hardham
10151 (DS, with C. hirtella); Tassajara Springs, Kelley in 1917 (CAS, with C.
intermedia); Cachagua Creek, Santa Lucia Mts., Howell 30253 (CAS). SAN LUIS
opispo co.: Bill Hill Ranch at the foot of Caliente Mt., Caliente Range, 2,300
ft., Twisselmann 3489 (CAS); Crocker Grade, Hardham 12548 (DS). SANTA
BARBARA CO.: “Santa Barbara,” Elmer 3943 (DS, F, POM, RSA; probably from
above Santa Barbara, in the mountains); upper Cuyama Valley, Hoffmann in
1929 (POM); Pine Corral Potrero near Salisbury Potrero, 4,500 ft., Sierra Madre
Mts., Muller & Blakley 1014 (DS, SBBG); trail to Zaca Peak, Eastwood 616
(CAS); Miranda Pine Mt., 3,250 ft., Graham 545 (UC).

Camissonia luciae is a hexaploid species that parallels the wide-
spread diploid C. hirtella in the variable notching of its petals. Pre-
sumably it has been derived from the diploid C. hirtella and the
tetraploid C. intermedia, but it is rather easily separated from both
by its relatively large flowers and pollen characteristics. The chro-
mosome number of ©. luciae has been determined in 7 individuals
derived from as many populations, and including the type collection
(see Appendix). It has been found growing sympatrically with C.
micrantha, C. hirtella, and C. intermedia, especially often with the
last-mentioned species.

295-655, O—68——10

304 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

39. Camissonia robusta Raven, sp. nov. Figure 41
Oenothera micrantha sensu Munz, N. Amer. Fl. II. 5: 160. 1965; pro parte.

Herba annua erecta robusta villosa, ad C. micrantha similis. Folia
superiora latiora quam in (. micrantha, usque ad 12 cm. longa et 2 cm.
lata. Inflorescentia semper pilis brevioribus glandulosis admixta.
Hypanthium 1.8-3.7 mm. longa, ostio 1-2 mm. diametro. Sepala
2.6-4.2 mm. longa, 1.1-1.8 mm. lata. Petala 3.2-7 mm. longa, 2.5-6
mm. lata, plerumque prope basin puncto rubro uno vel punctis
duabus. Filamenta staminum episepalorum 1.8-3 mm. longa, illa
epipetalorum 1-1.5 mm. longa; antherae 0.8-1. 6 mm. longae. Pollinis
granorum maximam partem 4-porum, rariore 5- vel 3-porum. Stylus
3-6.2 mm. longus; stigma 1-1.5 mm. diametro, sub anthesi antheris
staminorum longiorum circumnexum. Capsula 1.4-2.5 cm. longa, 1.5—
2 mm. crassa, transectione subquadrangularis in statu vivo, surrecta
vel in spiram unam contorta, Semina 0.9-1.2 mm. longa, 0.6-6.7 mm.
crassa, Chromosomatum numerus gameticus, n=21. Autogama.

Type: Disturbed soil along roadside, with Orthocarpus purpurascens,
Eschscholzia, Brassica, near kilometer 201, 5.5 miles south of San
Vicente, 950 ft., Baja California, 29 March 1962, P. H. Raven 17022
(DS 516799).

Distrinution (Figure 52): Coastal sage or chaparral, often in
disturbed or open places, coastal San Diego County, California, and
coastal northwestern Baja California, south to the vicinity of El
Rosario at about 30° N. lat.; also on Santa Cruz, Santa Catalina,
San Clemente, and Guadalupe Islands. From sea level to about 1,000
ft. alt. (to 3,500 ft. on Guadalupe Island ).

REPRESENTATIVE SPECIMENS EXAMINED:

U.S.: CALIFORNIA: LOS ANGELES CO. SANTA CATALINA I: Avalon, Trask in
1896 (NY, UC, US). san CLEMENTE L.: Vicinity of Mosquito Harbor, Abrams &
Wiggins 345 (DS, UC); N. coast, Trask 194 (NY, US); canyon 8. of Lemon Tank,
W. coast, Munz 6743 (POM). san pigco co.: U.S. IIwy. 395, 2 mi. 8. of turn to
Lilac, R17458 (DS); 2 mi. E. of Pala, Munz 10364 (POM); 2 mi. E. of San Pasqual,
R17408 (DS); Howard Canyon, La Jolla, F. E. & E. S. Clements 183 (COLO, F,
GH, NY, PH, UC, UCSB); N. of Pt. Loma, Stover 149 (SD); bank of Dulzura
Creek 3 mi. below Dulzura, Wiggins 2176 (CAS, DS, UC, WTU); Sweetwater R.,
base of San Miguel Mt., Gander 154.4 (POM, SD). SANTA BARBARA CO., SANTA
CRUZ 1.: Vicinity of Pelican Bay, Abrams & Wiggins 41 (F).

BAJA CALIFORNIA: Nachoguero Valley, Schoenfeldt 3409; Ensenada,
Bates 2341 (DS); Todos Santos Bay, Fish in 1882 (F, UC); 6 mi. E. of El Rosario,
Wiggins 17889 (DS); 6 mi. N. of Rancho Mesquital, Wiggins 16767 (DS).
GUADALUPE I.; Near mouth of canyon at Northeast Anchorage, Moran 2891 (CAS,
DS, GH, RSA); Northeast Anchorage, Wiggins & Ernst 222 (DS, UC), Moran
5661 (CAS, DS, RSA, SD, UC); near spring at top of island, N. end, ca. 1,200 m,
Lindsay in 1948 (SD); Barracks Cove Cafion, Copp 151 (Ds),

Camissonia robusta is one of the commoner species of its group
throughout its mainland area, and the commonest species of the sec-

RAVEN—GENUS CAMISSONIA 305

tion on Santa Catalina Island. On Santa Cruz Island this species is
very rare. On San Clemente and Guadalupe Islands, the only other
species of the section is the very different C. quadalupensis, but the
two species are ecologically different and not known to grow together.
The pollen of C. robusta is absolutely diagnostic, and its stout capsules,
which are evidently quadrangular but not nearly so sharply so in
living material as are those of C. Jewisii, and erect, robust habit make
this species distinctive in gross morphology also. It has been collected
erowing sympatrically with Camissonia bistorta, C. lewisit, C. micrantha,
C. hirtella, and C. intermedia.

The chromosome number of Camissonia robusta has been determined
in 29 individuals derived from 24 populations from localities scattered
throughout the range of the species, including the type collection
(see Appendix). This hexaploid is almost certainly derived from two
species with which it occurs nearly throughout its rather limited range,
the diploid C. lewisti and the tetraploid C. intermedia. It is morpho-
logically intermediate between these two entities in all respects,
although on the average somewhat more robust.

Moran (Madrofio 11: 160. 1951; as Oenothera micrantha) has sug-
gested that this species might be introduced on Guadalupe Island,
where it was first collected in 1948. He bases his argument largely
on the fact that this species was not discovered by any of a succession
of collectors who visited the island and inevitably landed in the same
region where C. robusta is now fairly common. This may be true, but
judging from the phytogeographic relationships of Guadalupe Island
and the scarcity of this species on San Clemente Island, I believe that
it is equally likely that the species may simply have gone undetected
and become more common recently. It will probably not be possible
to resolve this question definitely.

Section VI. Camissonia

Camissonia sect. Camissonia

Oenothera sect. Sphaerostigma Seringe, in DC., Prod. 3: 46. 1828.

Oenothera subg. Sphaerostigma (Seringe) Torr. & Gray, Fl. N. Amer. 1: 508.

1840.
Camissonia sect. Sphaerostigma (Seringe) Raven, Brittonia 16: 284. 1964.
Annuals, the plants caulescent, with no well-defined basal rosette,

but the leaves sometimes clustered near the base, plants not flowering
from the base. Leaves narrow, subsessile. Inflorescence nodding at
anthesis, becoming erect in fruit. Flowers opening near sunrise. Ovary
lacking a sterile projection. Sepals reflexed singly or in pairs. Petals
yellow, often with one to several red dots near the base, fading red-
dish, with no contrasting pattern visible in ultraviolet light. Stamens,
style, and inside of hypanthium yellow, the stigma yellow or greenish
yellow. Capsule regularly loculicidal, straight or somewhat flexuous,

306 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

subterete, more or less torulose, the seeds in one row in each locule,
the friable central column much distorted by the seeds at maturity.
Seeds with a scar at the micropylar end, blunt at the chalazal end,
shining, dark brown, minutely lacunose, narrowly obovoid, more or
less triangular in transection.

TYPE SPECIES: Camissonia dentata (Cav.) Reiche.

Distriputton: Brushy or open slopes and flats, southern Van-
couver Island, British Columbia, southern Washington, southern
Idaho, and western Wyoming south to western Colorado, central
and northern Nevada, and throughout California to northernmost
western Baja California (south to about 30°43’ N. lat.); said to have
been collected once at Fort Huachuca, southern Arizona; also in
western South America, mostly west of the Andes, from Arequipo
(ca. 16° S. lat.), southern Peru to the vicinity of Lago Argentino
and Santa Cruz (ca. 50° S. lat.), in Patagonia.

Camissonia sect. Camissonia is a close-knit group divided herein
into 12 species with 4 additional subspecies. Two of these, C. ker-
nensis and C. campestris, are self-incompatible diploids; two others,
C. pusilla and C. sierrae, are self-compatible diploids; one is an auto-
gamous hexaploid; and the remaining seven are autogamous tetra-
ploids. One obviously closely interrelated group of species comprise
the self-incompatible diploid C. kernensis, the autogamous diploid
C. pusilla, and the two autogamous tetraploids C. pubens and C.
parvula. These species differ from the others in their Great Basin
habitat, and in having the sepals separating from one another when
the flowers open. In the remaining species the sepals remain united
in pairs. Relationships within this group are close, and the popula-
tions included here have generally been regarded as comprising only
two species, the self-incompatible C. campestris (usually known, in-
correctly, as Oenothera dentata) and the autogamous (C. contorta
(usually divided into Oenothera contorta var. typica and Q. contorta
var. strigulosa). The species that I have named C. sierrae, C. beni-
tensis, and C. integrifolia have hitherto been too poorly represented
in collections to attract attention; C. lacustris has been confused with
C. campestris and C. contorta (sens. lat.); and the extremely closely
related trio consisting of the tetraploid C. strigulosa, the hexaploid
C. contorta, and the South American tetraploid C. dentata (some
populations with n=13, the only instance of aneuploidy known in
the genus) has in North America been regarded as comprising Veno-
thera contorta var. strigulosa, var. epilobioides, and var. contorta, the
lines between these taxa not corresponding to those between the
species recognized here. These might all conservatively be treated
as C’. dentata sens. lat., but I prefer to keep them separate for reasons
that will be discussed in some detail below.

RAVEN—GENUS CAMISSONIA 307

Camissonia kernensis subsp. kernensis was initially attributed to
Oenothera subg. Chylismia, but I excluded it from that group in 1962
(Univ. Calif. Publ. Bot. 34: 115). On the other hand, C. kernensis
subsp. gilmanii was first described as a variety of C. campestris (“‘Oeno-
thera dentata’”’); I pointed out the conspecificity of the two taxa here
erouped as C. kernensis to Dr. Munz in 1963, and he brought them
together in 1965 (N. Amer. Fl. II. 5: 158-159). Part of what is here
regarded as C. kernensis subsp. gilmanti was included by Munz
(Bot. Gaz. 85: 259. 1928) in his concept of Oenothera dentata var.
johnstonii, the type of which is Camissonia campestris. In the same
1928 treatment, Munz included all of the autogamous entities in this
group in Oenothera contorta, with his var. pubens corresponding to
C. pubens, his var. fleruosa to C. pusilla and C. parvula, his var.
typica to much of C. contorta, and his var. epilobioides and var. strigu-
losa including the other autogamous taxa known to him.

Determinations of chromosome number, and in many cases pairing,
have been made of 177 individuals from 166 populations of this
section, including a few counts reported earlier by Lewis, Raven,
Venkatesh, & Wedberg (Aliso 4: 73-86. 1958) and two reported by
Gregory & Klein (Aliso 4: 505-521. 1960). The chromosome number
for Camissonia campestris (as Sphaerostigma dentatum and var. cam-
pestre) was reported by Johansen (Proc. Nat. Acad. Sci. U.S. 15:
884. 1929; Amer. Journ. Bot. 16: 597. 1929), but without any indica-
tion of voucher specimens. As far as I know, these are all of the avail-
able reports for the section.

The degree of chromosomal structural heterozygosity in this section
is low: of the 35 individuals of Camissonia campestris subsp. campestris
examined, 3 had a ring of 4 chromosomes and 5 pairs and 1 had a
chain of 3 plus a univalent and 5 pairs. Considering the two self-
incompatible species in the section, C. campestris and C. kernensis,
49 individuals have been examined chromosomally, with the 4 just
indicated being the only ones that were heterozygous—approximately
2 percent of the total. This is much lower than the more than 20 percent
of 676 individuals of sect. Chylismia that were heterozygous for at
least one reciprocal translocation, but significantly higher than in
sect. Holostigma, where reciprocal translocations appear to play no
role whatever in natural populations.

Several interploid natural hybrids, discussed in the following pages,
shed light on the pattern of chromosomal evolution in sect. Camissonaa.
For example, in hybrids between the diploid (n=7) C. campestris
subsp. campestris and the hexaploid (n =21) C. contorta, the maximum
association of chromosomes observed was a chain of 8 chromosomes,
a ring of 4, 2 chains of 4, 2 ring bivalents, and 2 rod bivalents. It is
therefore obvious that the complete absence of multivalent configura-

308 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

tions in the hexaploid is genetically controlled, and does not reflect
any great degree of differentiation between the three genomes present
in this species. No multivalents have been found in any polyploid
in this section, and it would appear that even if the two diploid genomes
present in a tetraploid would, on the basis of their homology, pair
completely, they are prevented from doing this genetically. Natural
hybrids between the tetraploids C. integrifolia and C. strigulosa, with
up to 10 bivalents, and artificial hybrids between the North American
tetraploid C. strigulosa and the South American tetraploid C. dentata,
which show few multivalent associations despite their high degree of
sterility, tend to support this hypothesis (Raven & Moore, MS.).

In sect. Camissonia, therefore, it appears that diploid genomes may
be brought together in any way that will produce highly fit combina-
tions for particular ecological situations, and that these polyploids
will soon (perhaps initially) be pair-forming and highly fertile regard-
less of the degree of chromosomal divergence between their diploid
antecedents. There appears to be a wide gap between the group of
species consisting of C. kernensis, C. pusilla, C. pubens, and C. parvula,
and the other species of the section; none of the polyploids appears
to bridge the gap between these two groups.

The synonymous name Onothera torulosa H. Lév., Monogr. Onoth.
176. 1905, requires special mention here; this name was an illegitimate
substitute for most of the species in this section published earlier,
and it is not practical to cite it in the synonymy of all of these names
or to attempt to typify it by one of these concepts; as far as I know,
the concept has never been taken up by any author other than Léveillé.
40. Camissonia kernensis (Munz) Raven, Brittonia 16: 284. 1964.

Robust, erect annual, often well branched, from a prominent or
poorly developed basal rosette, the plants 5-30 em. tall, densely
villous or glandular pubescent to subglabrous. Leaves very narrowly
to narrowly elliptic, more rarely lanceolate, 1-3.8 (—5.5) em. long,
0.2-0.5 cm. wide, sparsely serrate, the apex acuminate, the base
narrowly cuneate, much reduced and bract-like in the inflorescence.
Inflorescence usually densely glandular-pubescent, sometimes with an
admixture of longer trichomes. Hypanthium 2.2-3.8 (—5) mm. long,
1.8-2.8 (-4) mm. across at the summit, villous within, with a red
dot within at the base of each anther. Sepals 5-9 (-11) mm. long,
1.3-2 mm. wide, reflexed separately. Petals 8-15 (—18) mm. long,
5.5-12 (-16) mm. wide, with two large red dots at the base of each
one. Filaments of the episepalous stamens 3.5—5.5 (—7) mm. long,
those of the epipetalous ones 1.3-2 (4.5) mm. long; anthers 1.8-2
(-3) mm. long. Style 7-10 (-14) mm. long, villous near the base;
stigma 1-2 (-2.5) mm. in diameter, held well ubove the anthers at
anthesis. Capsule 2.2-3.7 em. long, 1.5-1.7 mm. thick, subsessile or

RAVEN—GENUS CAMISSONIA 309

on a pedicel up to 15 mm. long. Seeds 1.1—1.2 mm. long, 0.5-0.6 mm.
thick. Gametic chromosome number, n =7. Self-incompatible.

DistRiBuTION (Figure 46): Desert washes and in canyons, north-
eastern Kern County to near Victorville and the Providence Moun-
tains in San Bernardino County, southern Inyo County, and near
Benton Station, southern Mono County, California; also in western
Clark and Nye Counties, Nevada; 2,500—6,000 ft. alt.

Although most populations of the two subspecies comprising this
species are easily assignable to one or the other, a number of others
are intermediate and assignable only with great difficulty. Collec-
tively, the species is sharply distinct from all others in its flower size,
pubescence, and sepal characteristics.
40a. Camissonia kernensis subsp. kernensis

Oenothera kernensis Munz, Amer. Journ. Bot. 18: 737. 1931.
Oenothera kernensis subsp. kernensis; Munz, N. Amer. FI. II, 5: 159. 1965.

Plants compact, 5-15 (-22) em. tall, covered everywhere with long,
white trichomes, the leaves clustered at the base. Capsule on a promi-
nent pedicel 3-15 mm. long. Gametic chromosome number, n=7.
Self-incompatible.

Type: Sandy slopes and openings, east slope of Walker Pass,
Freeman Canyon, 4,500 ft., Kern County, California, 13 May 1930,
F. W. Peirson 8822 (POM;; isotype, K).

DistripuTion (Figure 46): Sandy slopes and flats, often with
Artemisia tridentata and Yucca brevifolia, northeastern Kern County,
California, from El Paso Peaks to the vicinity of Claraville and near
Lake Isabella, north to Grapevine Canyon; 2,800—5,700 ft. alt. Often
locally abundant.

Vouchers for chromosome number (4 individuals, 3 populations),
n=T7:

U.S.: CALIFORNIA: KERN Co.: Walker Pass, W. slope, 5,000 ft., Munz 22581
(DS, RSA): 2.8 mi. W. of summit of Walker Pass, R12811; Freeman Canyon 3.4
mi. E. of summit of Walker Pass, R12763 (2 plants: ARIZ, DS, GH, SD, UC, US,
WTU).

REPRESENTATIVE SPECIMENS EXAMINED:

U.S.: CALIFORNIA: KERN co.: Grapevine Canyon, 2,800 ft., Twisselmann
7117 (CAS, mixture with C. campestris) ; upper end of Short Canyon, Linsley &
MacSwain 60-18 (RSA, UC); head of Indian Wells Canyon, 5,700 ft., Twisselmann
4525 (CAS, DS); NE. flank of Morris Peak, 4,175 ft., Twisselmann 7140 (CAS,
DS, SBBG); Horse Canyon, Scodie (Kiavah) Mt., 5,300 ft., Twisselmann 8178
(CAS, DS, RSA); Weldon, 2,400 ft, R17559 (DS); 9.7 mi. S. of Weldon, 3,100
ft., R17563 (DS); 5 mi. NW. of Dove Well, Rude 408 (DS) ; divide between Kelso
Canyon and Kelso Valley, 4,650 ft., Twisselmann 6912 (CAS, DS); Piute Mts.,
Harris Grade above Kelso Canyon, 3.1 mi. W. of Shorty’s Place, 5,420 ft., Twissel-
mann 2799 (CAS); Erskine Creek, 4-5,000 ft., Purpus 5083 (UC, mixture with
C. hirtella); Iron Canyon, El Paso Mts., ca. 3,300 ft., Budgett 30 (CAS); Dove
Spring Canyon 25 mi. SW. of Inyokern, Budgett 54 (CAS).

310 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

First described as a member of Oenothera subg. Chylismia because
of its stalked capsule, Camissonia kernensis subsp. kernensis is in
the overwhelming majority of its characteristics obviously a member
of sect. Camissonia. At the margins of its rather restricted range,
it intergrades completely with C. kernensis subsp. gilmanii (Oenothera
dentata var. gilmanti), an undoubted member of this group. Ten plants
from the locality of R17559, cited above, were tested for self-in-
compatibility by caging the plants in the field and selfing and out-
crossing them, with subsequent examination of pollen-tube growth.
All were found to be self-incompatible. Camissonia kernensis subsp.
kernensis is, at the head of Short Canyon, the principal pollen source
of the large oligolectic bee Andrena (Onagrandrena) mojavensis Linsley
& MacSwain, not known from any other locality (Linsley, MacSwain,
& Raven, Univ. Calif. Publ. Entom. 33: 59-98. 1964). Other oligolectic
species of Onagrandrena likewise visit the plant here and on Walker
Pass and are undoubtedly important in the outcrossing of these self-
incompatible individuals. The flowers of Camissonia kernensis subsp.
kernensis open before sunrise, and considerably before those of C.
campestris at most of its localities (Linsley et al., op. cit.).

Camissonia kernensis subsp. kernensis grows sympatrically with C.
campestris, which like it is self-incompatible at a number of localities:
no hybrids have been observed, even when, as at Short Canyon,
Kern County, California, the plants grow intermixed and are known
to be visited by the same species of bees. Camissonia kernensis subsp.
kernensis likewise grows sympatrically with C. pubens, C. strigulosa,
C. integrifolia, and C. contorta.
40b. Camissonia kernensis subsp. gilmanii (Munz) Raven, comb. nov.

Oenothera dentata var. johnstonii sensu Munz, Bot. Gaz. 28: 259. 1928; pro
parte.

Oenothera dentata var. gilmanii Munz, Leafl. West. Bot. 2: 87. 1938. Oen-
othera kernensis subsp. gilmanii (Munz) Munz, N. Amer. FI. II. 5: 159.
1965.

Ocnothera kernensis subsp. mojavensis Munz, N. Amer. Fl. II. 5: 159. 1965.
Type: 3 mi. W. of Pilot Knob, 900 m, western San Bernardino Co., Calif.,

20 March 1947, P. A. Munz 11666 (RSA 38224; isotypes, ARIZ, CAS,
COLO, SD, WTU).

Plants less compact, to 30 cm. tall, covered everywhere with short
glandular pubescence, usually with a few scattered longer trichomes,
or subglabrous, with few glandular trichomes, the leaves not markedly
clustered at the base. Capsule subsessile or on a pedicel to 5 mm.
(rarely to 15 mm.) long. Gametic chromosome number, n=7. Self-
incompatible.

Type: Bradbury Wash, 3,000 ft., Inyo County, California, 6 June
1937, M. F. Gilman 2587 (POM).

DisrriputTion (Figure 46): Desert washes and slopes, from the
vicinity of Red Rock Canyon, Kern County and Victorville and the

RAVEN—GENUS CAMISSONIA 311

Providence Mountains, San Bernardino County, California, north to
westernmost Clark and Nye Counties, Nevada, and the southern
half of Inyo County, California; also collected once near Benton
Station, southern Mono County, California. From 2,500—6,000 ft. alt.

Vouchers for chromosome number (5 individuals, 5 populations),
n=T7:

U.S.: CALIFORNIA: INYO Co.: Trona to Death Valley Road, N. base of Panamint
Mts., edge of Panamint Valley, Lewis & Mosquin 1119; 5.4 mi. from Emigrant
Junction on road to Wildrose Station, Lewis & Mosquin 1123. sAN BERNARDINO
co.: Dawes Siding, between Cima and Kelso, R11926; road to Camp Irwin 26.6 mi.
N. of U.S. Hwy. 466 at Barstow, R11944. NEVADA: NYE co.: Wash below Cane
Springs, R18947.

REPRESENTATIVE SPECIMENS EXAMINED:

U.S.: CALIFORNIA! INYO CO.: 10 mi. N. of Shoshone on road to Bradbury Well,
ca. 3,000 ft., Hitchcock 6106 (DS NY, POM, RM, RSA, US, WS, WTU); Great
Falls Canyon, Argus Mts., Wheeler & Richardson in 1930 (CAS, F, LA, LAM,
UC, US); Mill Creek Canyon, Panamint Mts., 1,300-1,700 m, Coville & Funston
759 (C, DS, GH, PH, US); Epsom Salt Works, Hidden Spring region S. of Wingate
Wash, 2,000 ft., Train 568 (ARIZ, COLO, DS, OSC, US); Cave Springs, Jaeger in
1927 (POM); Lee District (N. of Darwin), at Grape Vine Spring, Austin 539
(UC); 6 mi. NE. of Pilot Knob, Meerchaum Spring, Miller in 1920 (CAS). KERN
co.: U.S. Hwy. 395, 5 mi. N. of Johannesburg, 3,150 ft., Campbell 13406 (RSA) ;
Red Rock Canyon, 2,800 ft., Benson 3411 (POM); Haiwee, Davidson 2820 (LAM) ;
E. fork of Last Chance Canyon, El Paso Mts., 3,000 ft., Pefialosa 473A (CAS).
MONO co.: Benton Station, 5,500 ft., Robinson & Linder in 1935 (Fresno State
College herb.). SAN BERNARDINO Co.: County road from Wheaton Springs to Nip-
ton road S., ca. 11 mi. N. of Cima, Ferris & Bacigalupi 13293 (ARIZ, DS, RM,
US, WS, WTU); 10 mi. SW. of Garlic Spring, Munz & Keck 7869 (GH, POM); 2
mi. N. of third summit out of Barstow, 8 mi. SW. of Bicycle Lake, 2,800 ft., Wolf
6532 (ARIZ, DS, LA, NY, RM, RSA, UT, WS); 5 mi. E. of Cima, Munz 13947
(DS, POM, UC); 5 mi. W. of Columbia Mine, Providence Mts., 1 mi. E. of Union.
Pacific R.R., 3,300 ft., Wolf 10649 (DS, RSA, WTU); 8 mi. E. of Victorville,
Jaeger in 1932 (POM). Nevapa: cLARK Co.: Nipton, 3,000ft., Jones in 1907 (POM) ;
5 mi. below Deer Creek, 6,000 ft., Charleston Mts., Raedar 181 (COLO); French-
man Flat 11 mi. N. of Mercury, R18854 (DS). NyE co.: W. of Cane Springs, ca.
4,000 ft., Rickard & Beatley in 1959 (RSA, UC); W. Yucca Flat Drainage Basin,
4,800 ft., Beatley 2924 (DS, RSA).

Plants of Camissonia kernensis subsp. giumanii in the vicinity of
Cima, San Bernardino County, California, approach those of subsp.
kernensis in their relatively low stature and unusually abundant
eglandular pubescence. Closer to the restricted range of subsp.
kernensis, the two subspecies intergrade completely. In general,
plants from the southern part of the range of subsp. gilmanii are rela-
tively lower and less glandular, and often with more obviously crisped
leaves, than those from farther north; it was on this basis that Munz
described a third subspecies, Oenothera kernensis subsp. mojavensis.
Study of more abundant herbarium material has convinced me that
these trends do not need to be reflected in the formal taxonomy of the

312 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

group, especially since the geographical separation of the two sorts
of plants is not complete.

Camissonia kernensis subsp. gilmanii has been found growing
sympatrically with C. campestris at several localities without any
evidence of hybridization, even though both entities consist entirely
of self-incompatible individuals and are generally visited by the same
bee species. Thus, 3 miles west of Pilot Knob, San Bernardino County,
California, at an elevation of ca. 3,400 ft., P. A. Munz collected these
two entities growing together; here C. kernensis subsp. gilmanii was
restricted to the higher ground, and often grew in coarse gravelly
soil, whereas C. campestris grew in the loose sandy soil of a wash. A
mixed collection from Red Rock Canyon, Kern County, California
(Brown in 1946, LA) suggests that the two species grow sympatrically
at this locality also. Similar ecological relationships were observed
near the head of Short Canyon, Kern County, California, where
C. campestris grows in the sandy soil of the wash bottom, while
C. kernensis subsp. kernensis is largely restricted to more stable slopes
at the head of the wash.

The self-incompatibility of Camissonia kernensis subsp. gilmanii
was determined by pollen-tube growth following self- and cross-
pollination in six individuals of R18947 from Nye County, Nevada,
cited above.

41. Camissonia pusilla Raven, sp. nov. FIGuRE 53
Oenothera contorta var. flecuosa sensu Munz, Bot. Gaz. 85: 253. 1928; and
N. Amer. FI. IT. 5: 156. 1965; pro parte.

Herba annua tenuis 2-22 cm. alta, plerumque e basi ramis plurimis
rosulaque bene evoluta. Plantae glanduloso-pubescentes, praecipue
in inflorescentibus, et plerumque pilis longioribus eglandulosis ad-
mixtae, prope basin plantarum densiore. Folia linearia, 1-3 cm.
longa, 0.4-1.8 mm. lata, plerumque purpureo-maculata, serrulata.
Hypanthium 0.8-1.6 mm. longum, ostio 0.7-1.1 mm. diametro, intus
glabrum. Sepala 1.2-2 mm. longa, 0.3-0.8 mm. lata, unumquidque
sub anthesi discedens, alabastro apices sepalorum discreti. Petala
1.8-3.1 mm. longa, 0.8-1.5 mm. lata, rare prope basin punctibus
duabus rubris maculata. Filamenta staminorum episepalorum 0.8-2
mm. longa, illa epipetalorum 0.4-0.9 mm. long; antherae 0.3-0.4 mm.
longae. Stylus 1.6-3.2 mm. longus, glaber; stigma 0.5—-1 mm. diametro
sub anthesi antheris circumnexum. Capsula 1.8-3.2 cm. longa,
0.6—0.9 mm. crassa, subsessilis vel pedicelo usque ad 2 mm. longo,
non rostrata. Semina 0.7—-0.8 mm. longa, ca. 0.4 mm. crassa. Chromo-
somatum numerus gameticus, n=7. Autogama.

Type: On sagebrush (Artemisia tridentata) covered slopes 2.5 miles
west of U.S. Hwy. 395 on U.S. Hwy. 50, 5,100 ft., Washoe County,
Nevada, 28 May 1962, P. H. Raven 17867 (DS).

RAVEN—GENUS CAMISSONIA 313

fp (MW)

Ficures 53-55.—Species of Camtssonia sect. Camissonia, X 34:53, C. pusilla (Raven 17463,
DS); 54, C. pubens (Twisselmann 7048, DS); 55, C. parvula (Raven 17864, DS).

314 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Disrripution (Figure 56): Scattered in sandy soil on open or
brushy slopes, Franklin and Grant Counties, southeastern Washing-
ton, and Elmore and Payette Counties, west-central Idaho, south
through the Great Basin to southern Inyo and the Providence Moun-
tains of northeastern San Bernardino County, California; southern
Nye County, Nevada; and Washington County, southwestern Utah;

Ficure 56.—Western United States, showing ranges of mainly autogamous species of
Camissonia sect. Camissonia, the range of the related larger flowered, self-incompatible
C. kernensis outlined by dotted line: A=C. pusilla, with A=diploid chromosome
counts (n=7); O=C. pubens, with @=tetraploid counts (n=14); O=C. parvula,
with @=tetraploid counts (n= 14).

RAVEN—GENUS CAMISSONIA 315

local in the eastern San Bernardino Mountains, San Bernardino
County, California. From ca. 400—-9,500 ft. alt.

Vouchers for chromosome number (5 individuals, 5 populations),
n=T7:

U.S.: CALIFORNIA: Inyo co.: Carroll Creek ca. 1 mi. above Mt. Whitney
Pack Station, 5,900 ft., E. slope of the Sierra Nevada, R17541; ca. 4 mi. 8. of
Oasis, Klein 354 (RSA); near Black Ace Mine, just W. of summit of Westgard
Pass, White Mts., Lew7s 1091 (LA, RSA). SAN BERNARDINO Co.: Cactus Flat,
San Bernardino Mts., 5,900 ft., R17463. Nevapa: WASHOE co.: 2.5 mi. W. of
U.S. Hwy. 395 on U.S. Hwy. 50, 5,100 ft., R17867 (type collection).

REPRESENTATIVE SPECIMENS EXAMINED:

U.S.: CALIFORNIA: INYO Co.: 4 mi. W. of Lone Pine, 4,700 ft., Alexander &
Kellogg 2786 (mixture with C. parvula); Lake Sabrina road, 8,500 ft., Benson
6020 (POM); Bishop Creek, 5,000 ft., Hall & Chandler 7244 (POM, UC); 0.5
mi. W. of Willow Spring, Last Chance Mts., 5,800 ft., Roos 6363 (CAS, DS,
RSA, UC); Jackass Spring, Hunter Mt., Inyo Mts., 6,600 ft., DeDecker 1400
(RSA); Alabama Hills, 4,800 ft., Howell 33226; Lone Pine, 7,000 ft., Jones in
1897 (GH, NY, POM, US); Argus Mts., 5,800 ft., Hall & Chandler 7089 (UC);
Wonoga Peak, 9,000-9,500 ft., Howell 25419 (CAS, RSA; with C. pubens); Mill
Creek Canyon, Panamint Mts., 1,300-1,700 ft., Coville & Funston 762 (DS,
NY, US); near Crystal Spring, Coso Mts., 1,850 m., Coville & Funston 938 (DS,
NY, US). Lassen co.: 2 mi. N. of Susanville, Gillespie 9346 (DS). MoNo co.:
Rock Creek, 6,700 ft., Munz 11785 (NY, RSA); near Benton, Robinson & Linder
c26 (COLO, RSA, WTU), Eastwood & Howell 9547 (CAS, mixture with C.
parvula; RSA); 5 mi. N. of Sherwin Grade, 1,750 m., Munz 11075 (C, GH, POM,
UC, US); S. of Mono Lake, 4 mi. N. of Mono Mills, Keck 2882 (DS). SAN BER-
NARDINO Co.: 1 mi. SE. of Tecopa Pass, Kingston Range, Alexander & Kellogg
2365 (POM, UC); Cactus Flat, San Bernardino Mts., 6,000 ft., Munz 10519
(POM), Jones in 1926 (CAS, POM). Ipano: ELMoRE co.: Glen’s Ferry, Davis
85-36 (IDS). payeTTE co.: Willow Creek E. of Payette, Christ 9447 (NY)
NEVADA: ELKO Co.: Cobre, 6,000 ft., Jones in 1906 (POM); Elko to Wendover
road, Hutchings in 1932 (UT); Toana Range, 5 mi. E. of Shafter, Maguire 16892
(NY, mixture with C. parvula). ESMERALDA co.: Candelaria, Shockley 247 (NY,
mixture with C. parvula). EUREKA co.: U.S. Hwy. 40, 2.9 mi. W. of Carlin, R18534
(DS). LANDER co.: 3 mi. N. of Birch Creek, J. M. & M. A. R. Linsdale 780A
(CAS). nrE co.: Near Potts Ranch, near Potts, Goodner & Henning 330 (POM) ;
4 mi. 8. of Millett, J. M. & M.A. R. Linsdale 694 (CAS); 8. end of Rainier Mesa,
7,500 ft., Beatley & Carl 2616 (herb. Nevada Test Site, Mercury, Nev.). WASHOE
co.: 2 mi. SW. of Steamboat Springs, Canby 181 (POM); Truckee Pass, 4,500
ft., Kennedy 1306 (DS, RM, UC, US). waite PINE co.: 2 mi. SW. of Baker,
Maguire 20849 (UT). OrEGoN: Blue Mts., Cusick in 1897 (GH); eastern Oregon,
Howell in 1880 (US). HARNEY co.: Ca. 26 mi. S. of Folly Farm and near Manns
Lake, Hitchcock & Muhlick 21131 (DS, RM, WS, WTU); 8 mi. W. of Riley,
Peck 13840 (DS, F); Squaw Butte, Gilkey (OSC); 4.7 mi 8S. of Narrows, R18444
(DS). mMALHEUR co.: 4 mi. W. of Vale, Peck 20641 (UC); near Harper Ranch,
900 m., Leiberg 2221 (DS, F, GH, NY, POM, UC, US); ca. 15 mi. SW. of Vale,
Hitchcock 20636 (WTU); Rome, Peck 21789 (UC). UTau: 1ron co.: NE. of Enter-
prise, 5,500 ft., Hall EN16 (F, UT). wasuinaton co.: Anderson’s Ranch, Maguire
& Blood 1459 (POM, RM, UC); mountain meadows 50 mi. NW. of St. George,
1877, Palmer 166 (GH, NY). WASHINGTON: FRANKLIN Co.: Pasco, Piper 2965
(GH). grant co.: Junction of Crab and Douglas Creeks, 2,000—3,000 ft., Sandberg
& Leiberg 263 (C, CAS, DS, F, GH, NMC, NY, RM, UC, US, WS, WTU).

316 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

This diploid species is closely similar to the tetraploids C. parvula
and C. pubens, differing from the former in its long, spreading tri-
chomes and well-developed basal rosette and from the latter in its
more slender habit, narrower leaves, and smaller flowers and seeds.
At its type locality and elsewhere, C. pusilla grows sympatrically
with C. pubens, and, as indicated above, it often grows sympatrically
with C. parvula; at all such localities the entities are sharply distinct.
On the other hand, northern populations of C. pusila are occasionally
somewhat difficult to distinguish from those of C. parvula, commoner
northward, when the plants are relatively glabrous and depauperate,
and chromosome counts from the northern localities of C. pusilla
would be most helpful.

A collection said to be from Kramer, San Bernardino County,
California, A. Brandegee in 1913 (UC), consists of a mixture of this
species with C. parvula; the locality is very probably in error (cf.
Raven, Univ. Calif. Publ. Bot. 34: 110. 1962).

Camissonia pusilla includes much of the material referred by Munz
(Bot. Gaz. 85: 253-254. 1928; and subsequently) to Oenothera con-
torta var. flexuosa, that entity consisting of a mixture of this species,
the tetraploid C. parvula, and misidentified specimens of the very
different hexaploid C. contorta. In 1964 (Brittonia 16: 284. 1964),
I considered C. pusilla, C. parvula, and C. pubens to be best regarded
as comprising a single taxonomic species, C. parvula; I also annotated
a number of specimens during the period when I held this view.
On the other hand, Munz (N. Amer. FI. IT. 5: 156. 1965) recognized
C. pubens as a distinct species, but unaccountably retained C. parvula
and C. pusilla as Oenothera contorta var. fleruosa as he had done in
1928. As mentioned above, C. contorta is a very distinct hexaploid,
in which the sepals are reflexed in pairs; however, depauperate
individuals can sometimes be confused with those of C. parvula.

42. Camissonia pubens (8S. Wats.) Raven, comb. nov. Ficure 54

Chamissonia contorta var. pubens (S. Wats.) Kearney, Trans. N.Y. Acad.
Sci. 14: 37. 1894.

Sphaerostigma contortum var. pubens (S. Wats.) Small, Bull. Torrey Cl. 23:
189. 1896.

Sphaerostigma pubens (S. Wats.) Rydb., Bull. Torrey Cl. 33: 146. 7 April 1906.

Sphaerostigma orthocarpum A. Nels. & Kennedy, Proc. Biol. Soc. Wash. 19:
155. 12 November 1906. Type: Pyramid Lake, Washoe Co., Nevada,
19 May 1905, P. B. Kennedy 1015a (RM).

Oenothera pubens (S. Wats.) Munz, N. Amer. FI. II. 5; 156. 1965.

Similar to C. pusilla, but differing in the more robust habit, the
stouter stems to 38 cm. long. Leaves narrowly lanceolate, 1.5—4.5 em.
long, 0.2-0.6 cm. wide, undulate-serrate, acuminate at the apex,
narrowly cuneate at the base. Hypanthium 1.3-3 mm. long, 0.8-2 mm.
wide at the summit, usually pubescent in lower half within, often
with a small red dot at the base of each filament. Sepals 2.2-3.8 mm.

RAVEN—GENUS CAMISSONIA 317

long, 0.8-1.3 mm. wide. Petals (2.2—) 3-4 mm. long, (1.3-) 1.8-3 mm.
wide, often with one to several red dots near the base. Style 3.2-4.1 mm.
long; stigma 1.2-1.4 mm. in diameter. Capsule (1.8—) 2.6—-5 cm. long,
0.8-1.2 mm. thick. Seeds 1-1.1 mm. long, ca. 0.4 mm. thick. Gametic
chromosome number, n= 14.

Type: Carson City, Ormsby County, Nevada, May 1868, S. Watson
(GH; isotype, US).

DistriBuTION (Figure 56): Sandy, often sagebrush-covered slopes
and flats from Lassen County, California, and northern Washoe
County, Nevada, south to southwestern Lander and central Mineral
Counties, Nevada, and southern Inyo and northeastern Kern Coun-
ties, California; 3,000-9,500 ft. alt.

Vouchers for chromosome number (9 individuals, 9 populations),
n=14:

U.S.: CALIFORNIA: KERN co.: Weldon, R17537; 3.4 mi. E. of summit of Walker
Pass, R12764. LASSEN co.: 4.8 mi. N. of Milford, R17873. Mono co.: 2.2 mi. 8.
of N. turn to Sherwin Grade, R14294. Nevapa: ormsBy co.: U.S. Hwy. 50,
21 mi. W. of junction with U.S. Hwy. 395, R17862; 2 mi. 8. of Carson City
R18406. wasHoE co.: 7.5 mi. S. of Vya, ca. 6,500 ft. R17883; 4.7 mi. W. of
U.S. Hwy. 395 on road to Mt. Rose, 5,200 ft., R18410; 2.5 mi. W. of U.S. Hwy.
395 on U.S. Hwy. 50, 5,100 ft., R17866.

REPRESENTATIVE SPECIMENS EXAMINED:

U.S.: CALIFORNIA: INYO co.: Wonoga Peak, 9,000-9,500 ft., Howell 25420
(CAS, RSA, US); near end of Mt. Whitney road, Kerr in 1937 (CAS); Carroll
Creek, 5,500 ft., Howell 33378 (CAS); Olancha, Hoffmann in 1929 (CAS); Bishop
Creek, 6,700 ft., Howell 33520 (CAS); 5 mi. W. of summit of Mountain Spring
‘anyon, Hoffmann in 1929 (POM); Onion Valley road, Howell 33430 (CAS);
Sage Flat to Olancha Pass, ca. 7,000 ft., Howell 26701 (CAS); Taboose Creek,
6,500 ft., R9702 (CAS); Nine-mile Canyon, 5,000 ft., Ripley & Barneby 5867a
(CAS); Snake Hill, 5 mi. W. of Independence, 5,000 ft., Kerr in 1937 (CAS).
KERN co.: 19 mi. above Weldon on road to Walker Pass, 4,500 ft., Munz 13357
(RSA, mixture with C. strigulosa; WTU); 1.9 mi. E. of summit of Walker Pass,
R12814 (DS); W. slope of Walker Pass, near summit, T'wisselmann 7048 (CAS,
DS); Piute Mt. to Walker Basin road, 5,950 ft., Twisselmann 3650 (CAS); Weldon,
R18790 (DS); Horn Brothers Sawmill 3.6 mi. W. of Claraville, Piute Mts., 6,200
ft., Breedlove 4008 (DS); Erskin Creek, 4,000-5,000 ft., Purpus 5365 (UC).
LASSEN cCo.: Honey Lake, Brandegee in 1892 (UC); between Susanville and Leavitt
Lake, Gillespie 935la (DS); vicinity of Doyle Station, 1,290 m, Eggleston 6741
(GH, NY, US). mono co.: Sonora Junction, 7,200 ft., Ripley & Barneby 5804
(CAS, NY). Nevapa: pouaias co.: Clear Creek, 10 mi. SW. of Carson City,
Archer 6055 (POM). LANDER co.: Along roadside to Buzanes, ca. 25 mi. N. of
Ione, Goodner & Henning 690 (DS, NY, POM). MINERAL co.: E. slopes of Wassuk
Range, near Big Indian Mt., 9,500 ft., Archer 7153 (ARIZ, POM); Hawthorne,
Jones in 1882 (POM); 6-7 mi. from mouth, up Cory Creek, Wassuk Range,
7,400 ft., Archer 6951 (POM). ormsBy co.: Empire City, Jones 3870 (CAS, F,
PH, UC); Carson City, 1,446 m, Baker 971 (CAS, F, GH, NMC, NY, POM,
RM, UC, US). srorey co.: Virginia City, Jones in 1882 (POM); 1.8 mi. SE. of
Five Mile House, 6,800 ft., Adams 31 (UC). wasHor co.: 5 mi. N. of Poeville,
NW. of Reno, 5,000 ft., Tillotson 99 (RSA, UC); Reno, Jones in 1903 (POM);

318 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Verdi, Sonne in 1889 (F); Lakeview, between Carson City and Franktown,
K. Brandegee in 1913 (UC).

This distinctive tetraploid species occurs sympatrically with C.
pusilla, It has also been found growing together with C. parvula,
C. kernensis subsp. kernensis, C. strigulosa, C. integrifolia, and C. con-
torta, with no evidence of interspecific hybridization. The tetraploid
chromosome voucher from northern Washoe County, Nevada, is
rather doubtfully referred here, and more information is needed about
the populations in the northern portion of the range of this species.
This entity has generally been regarded as distinct and has rarely
been confused with any other species since it was first described in
1873, as can readily be seen from the synonymy.

43. Camissonia parvula (Nutt. ex Torr. & Gray) Raven, Brittonia 16; 284. 1964.
FiagureE 55
Oenothera parvula Nutt. ex Torr. & Gray, Fl. N. Amer. 1: 511. 1840.
Sphaerostigma parvulum (Nutt. ex Torr. & Gray) Walp., Rep. 2: 78. 1843.
Sphaerostigma contortum var. flecuosum A. Nels., Bot. Gaz. 40: 58. 1905.
Type: Point of Rocks, Sweetwater Co., Wyo., 16 June 1898, A. Nelson
4760 (RM; isotypes, GH, US).
Sphaerostigma filiforme A. Nels., Bot. Gaz. 40: 57. 1905. Type: New River
(Reese’s River), Utah, 28 May 1889, collector not known (MO).
Sphaerostigma flecuosum (A. Nels.) Rydb., Fl. Rocky Mts. 601. 1917.
Oenothera contorta var. fleruosa (A. Nels.) Munz, Bot. Gaz. 85: 253. 1928.

Slender annuals closely similar to C. pusilla, but differing as follows:
basal rosette absent, with one or more wiry, naked stems arising from
the base; pubescence strigulose, dense or wanting, or very rarely
spreading, the plants usually with a sparse admixture of glandular
trichomes, especially in the inflorescence, but often subglabrous;
flowers sometimes larger, the hypanthium up to 2 mm. long; sepals to
2.2 mm. long and 1.1 mm. wide; petals to 3.6 mm. long.

TypE: Probably from southern Idaho (“toward Lewis’ River’’),
July or August 1834, T. Nuttall (NY; isotype, BM).

Disrrisution (Figure 56): Sandy slopes, often under sagebrush,
from Okanagan and Yakima Counties, Washington; Elmore and
Fremont Counties, Idaho; and Teton, Washakie, and Carbon Coun-
ties, Wyoming, south through westernmost Colorado (Moffat and Mesa
Counties), throughout Utah, to westernmost Nye and southern
Mineral Counties, Nevada and central Inyo County, California;
from ca. 400 (in southern Washington) to 6,600 ft. alt.

Vouchers for chromosome number (5 individuals, 5 populations),
n=14:

ULS.: CALIFORNIA: INYO Co.: 2.5 mi. N. of Big Pine, R17860. LasseN co.: U.S.
Hwy. 395, 32 mi. 8. of Milford, R17870. Ipano: FREMONT co.: 4.8 mi. E. and 2.5
mi. N. of St. Anthony, R19561. Nevapa: ormsBy co.: Just E. of New Empire,
R17864. OREGON: HARNEY Co.: 12.9 mi. 8. of Narrows, R18449.

RAVEN—GENUS CAMISSONIA 319

REPRESENTATIVE SPECIMENS EXAMINED:

U.S. CaLiForRNIA: INYO Co.: 2.8 mi. N. of Big Pine, R13825 (DS); 4 mi. W. of
Lone Pine, 4,700 ft., Alerander & Kellogg 2786 (UC, mixture with C. pusilla) ;
head of Division Creek road, 6,500 ft., Kerr in 1938 (CAS) ; foothills W. of Bishop,
Heller 8288 (CAS, DS, F, GH, NY, PH, UC, US, WS). Lassen co.: Secret Valley,
Applegate 8930 (DS, RSA); 10 mi. 8. of Amadee, 4,000 ft., Jones in 1897 (POM).
MONO Co.: Long Valley, Noldecke in 1938 (CAS) ; near Benton, Eastwood & Howell
9547 (CAS, mixture with C. pusilla). CoLorapo: mMEsA co.: Grand Junction,
Eastwood in 1892 (OSC). Morrat co.: 5 mi. W. of Maybell along Cedar Springs
draw, Weber 5450 (COLO, DS). Ipano: county UNKNowN: Lannans Spur,
Henderson 5339 (DS). cassta co.: Raft R., Davis 8-37 (IDS). Goop1NeG co.: 3 mi.
S. of Hagerman, Davis 1950 (IDS, POM, UC). sERome co.: Blue lakes 1893,
Palmer 74 (US). owYHEE co.: N. of Bruneau, Ripley & Barneby 6503 (CAS) ;
Indian Cove near Hammett, Christ 9533 (NY); Bruneau, Christ 4483 (NY).
TWIN FALLS co.: W. of Buhl, Piemeisel 40-211 (WTU). NEVADA: ELKO Co.: 2 mi.
S. of Contact, Holmgren 949 (WS); Toana Range, 5 mi. E. of Shafter, Maguire
16892 (NY, mixture with C. pusilla). ESMERALDA co.: Candelaria, Shockley 247
(DS; F, mixture with C. pusilla; JEPS). HuMBOLDT co.: 10 mi. N. of Winnemucca,
4,500 ft., Breene 185 (POM). nyE co.: Tonopah, Shockley 147 (DS, POM), Twin
R., 6,000 ft., J. M. & M.A. R. Linsdale 858 (CAS); 10 mi. S. of Calloway, 4,900
ft., Ripley & Barneby 3633 (CAS). ormssy co.: Carson City, 5,000 ft., Jones in
1897 (POM); Steamboat Springs, Stokes in 1903 (UC). wasHOE co.: 2.1 mi. NE.
of Pyramid, 4,100 ft., Stmontacchi 553 (RSA, UC). OREGON: HARNEY CO,: 21 mi.
N. of Frenchglen, Hitchcock & Muhlick 21204 (WTU); Narrows, Peck 6963 (GH) ;
Squaw Butte, Steward in 1956 (OSC); 3 mi. S. of Frenchglen, Peck 25205 (UC).
LAKE co.: Alkali L., Peck 19440 (CAS) ; near Summer L., 1,360 m, Eggleston 6861
(NY, US). maLHeur co.: Near Harper Ranch, 900 m, Leiberg 2108 (UD). Urau:
BEAVER Co.: Beaver Mts., near Milford, 4,900 ft., Stokes in 1903 (US). GRAND Co.:
Thompsons Spring, Jones in 1891 (POM). satr Lake co.: Antelope I., 4,300 ft.,
Watson in 1869 (GH, NY, US). saNnpEreE co.: Sevier Bridge, Jones in 1880 (POM,
US). TOOELE co.: Gold Hill, Jones in 1891 (F, NY, POM, RM); Deep Creek,
Jones in 1891 (F, GH, POM, UC, US). uinTAH co.: 2 mi. 8. of Jensen, 4,700 ft.,
Graham 9019 (POM); between Dinosaur National Monument and Green River,
4,800 ft., Graham 7739 (POM). wasHincTon Co.: Silver Reef, 2,500 ft., Jones 5149n
(POM). WasHINGTON: FRANKLIN Co.: Pasco, Piper 2965 (WS, mixture with C.
pusilla; GH specimen is all C. pusilla), OKANAGAN Co.: Nespelem Road at Hopkins
Canyon, Fiker 1874 (WS). yaKima co.: Near North Yakima, Henderson in 1892
(WTU); Donald Pass, Rattlesnake Hills, Hoover 5789 (UC). WYOMING: CARBON
co.: Sierra Madre, Nelson in 1907 (RM); Ft. Steele, 6,500 ft., Tweedy 4432 (NY,
US). SWEETWATER co.: Granger, Nelson 4760 (NY), 4698 (RM, US); Point of
Rocks, 6,550 ft., Ripley & Barneby 7922 (CAS). TETON co.: Pacific Creek, Merrill
& Wilcox 602 (GH, NY, RM, US). wasHaxie co.: W. of Worland, 4,100—4,300
ft., Nichols 380 (RM).

Camissonia parvula is closely related to C. pusilla and C. pubens
and grows sympatrically with both species. It is the only species of
sect. Camissonia in Wyoming and Colorado and the most widespread
one in Idaho and Utah. It can usually be distinguished readily by its
linear leaves, lack of a definite basal rosette, and strigose pubescence,
but the plants are occasionally subglabrous or very rarely have a few
spreading longer trichomes, in which cases they are difficult to dis-

295-655 O—68——-11

320 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

tinguish from the diploid C. pusilla that occurs in the same general
areas in northern Nevada and eastern Washington and Oregon.
Camissoma parvula has also been found growing sympatrically with
C’. contorta.

44. Camissonia campestris (Greene) Raven, Brittonia 16: 284. 1964.

Slender, erect or decumbent annual, usually well branched, with
wiry stems and no definitely defined basal rosette, the plants 5-25
(-50) cm. tall, glabrous, hirsutulous, strigulose, or glandular-pubescent;
stems with a conspicuously exfoliating white rhytidome. Leaves linear
to narrowly elliptic or narrowly oblanceolate, 0.5-2.5 (-3) em. long,
0.1-0.15(-0.5) cm. wide, sparsely serrulate to coarsely serrate, the apex
acuminate, the base attenuate. Inflorescence glandular-pubescent,
sometimes very sparsely so or glabrous, or with an admixture of
longer eglandular trichomes, or strigulose, sometimes densely so, or
sparsely short-villous. Hypanthium 1.5-5.5 mm. long, 2.5-6 mm. across
at the summit, more or less densely villous in lower half within, often
with a red dot at the summit within at the base of each filament.
Sepals 3-8 (—12) mm. long, 1.3-3 mm. wide, reflexed in pairs. Petals
(3.5-) 5-15.5 mm. long, (3—) 5-13 mm. wide, usually with two (or one)
red dots at the base of each one. Filaments of the episepalous sta-
mens (1.4—) 2.1-5.5 mm. long, those of the epipetalous ones (0.7-) 1.2—
3.2 mm. long; anthers 1—-2.4 mm. long. Style (3.2—) 4-12 (-15) mm. long,
glabrous or sparsely hirsutulous near the base; stigma 0.7-3 mm. in
diameter, held well above the anthers at anthesis. Capsule 2—4.3 cm.
long, 0.7-1.5 (-2) mm. thick, subsessile. Seeds 0.8-1.6 mm. long, 0.4—
0.6 mm. thick. Gametic chromosome number, n=7. Self-incompatible.

DistRiBuTIoN (Figure 57): Open sandy flats, endemic to California;
often abundant and covering large areas, from the vicinity of Antioch,
Contra Costa County, San Joaquin County and northern Stanislaus
County, California, south in the San Joaquin Valley, inner Coast
Ranges, and Sierra Nevada foothills to Santa Barbara, Ventura, the
northern half of Los Angeles, and western Riverside County; near
Campo, San Diego County (Eastwood 9447, CAS); and western
borders of the Mojave Desert from the vicinity of Twentynine Palms
and the Little San Bernardino Mountains in northern Riverside and
southern San Bernardino Counties northwest to the vicinity of Daggett
and Calico, east of Barstow, San Bernardino County, the area about
Red Mountain, in westernmost San Bernardino and easternmost
Kern Counties, and the southwesternmost corner of Inyo County,
where it extends north almost to Olancha. Not found near the coast,
and occurring from near sea level on the lower San Joaquin River to
about 6,400 ft. alt., in the Sierra Nevada of Kern County. I have also
seen a collection from Painted Canyon, in the northwestern corner
of the Colorado Desert, Riverside County (Randall 219 SD); the

RAVEN—GENUS CAMISSONIA 321

occurrence of Camissonia campestris at this locality should be con-
firmed by additional collections, as should the following: Santa Cruz
Mountains, 2,300 ft., Pendleton 945 (UC); Panamint Mountains,
Inyo County, Winblad in 1937 (CAS). Collections of this species
cited from the Sacramento Valley and bordering mountains are all
polyploid and here referred to C. lacustris and C. contorta; otherwise
all material cited by Munz (Bot. Gaz. 85:258-260. 1928) under
Oenothera dentata var. campestris, var. johnstonii (except for collec-
tions cited above as Camissonia kernensis subsp. gilmanii), and var.
parishii belongs here, and specifically to Camissonia campestris subsp.
campestris,

This species has usually been referred to the South American
Camissonia dentata, and consequently known as Oenothera dentata
Cav.; all South American populations, however, are tetraploid (n=14
or 13) and self-compatible, the stigma being surrounded by at least
the anthers of the longer stamens at anthesis.

44a, Camissonia campestris subsp. campestris

Oenothera dentata sensu auct. Amer. bor. mult.; non Cav. 1798.

Oenothera campestris Greene, Fl. Francisc. 216. 1891.

Sphaerostigma campestre (Greene) Small, Bull. Torrey Cl. 23: 189. 1896.

Sphaerostigma campestre var. parishit Abrams, Fl. Los Angeles 272. 1904.
Type: Plains, San Bernardino, San Bernardino Co., Calif. May 1900,
S. B. Parish (DS).

Oenothera torulosa race helianthemiflora H. Lév., Monogr. Onoth. 178, pl. 1905.
Lectotype: Mojave Desert, Calif., May 1882, S. B.& W. F. Parish 1304
(MO).

Sphaerostigma campestre var. helianthemiflorum (H. Lév.) A. Nels., Bot. Gaz.
40: 57. 1905.

Oenothera torulosa f. permizta H. Lév., Monogr. Onoth. 180. 1905. Type:
Fresno, Fresno Co., Calif., 21 March 1889, collector unknown (presumably
MO).

Oenothera dentata var. campestris (Greene) Jeps., Man. FI. Pl. Calif. 685, 1925.

Oenothera dentata var. parishii (Abrams) Munz, Bot. Gaz. 85: 259. 1928.

Oenothera dentata var. johnstonii Munz, Bot. Gaz. 85: 259. 1928. Type: Near
Mojave, Kern Co., Calif., 5 May 1920, J. M. Johnston (POM 8844).

Sphaerostigma dentata subsp. campestris (Greene) Johansen, Amer. Journ. Bot.
16: 597. 1929.

Oenothera campestris subsp. campestris Munz, N. Amer. FI. II. 5: 158. 1965.

Oenothera campestris subsp. parishii (Abrams) Munz, Fl. N. Amer. II. 5:
158. 1965.

Oenothera cruciata sensu Munz, N. Amer. FI. II. 5: 157, 1965; pro parte.

Plants usually erect; leaves linear to narrowly elliptic or narrowly
oblanceolate, serrulate. Gametic chromosome number, n=7. Self-
incompatible.

Lecroryre: California, 1831-2, D. Douglas (GH); cf. Munz, Bot.
Gaz. 85: 258. 1928.

DIsTRIBUTION (Figure 57): That of the species.

322 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Ficure 57.—California, showing ranges of species of Camissonia sect. Camissonia: @ =C
campestris subsp. campestris; X=C. campestris subsp. obispoensis; %—=C. sterrae
subsp. sterrae; C. sierrae subsp. alticola, area outlined by heavy black line.

Vouchers for chromosome number (35 individuals, 32 populations),
n=7 (all with 7 pairs, except where noted):

U.S.: CALIFORNIA: FRESNO CO.: 3.3 mi. N. of Selma, R17064; 7 mi. SW. of
Tollhouse, Breedlove 5199. KERN co.: Jawbone Canyon, Munz & Gregory 23310
(RSA); 1.2 mi. SE. of Blackwells Corner, 650 ft., R16982; 0.7 mi. S. of Taft, 800
ft., Breedlove 1947; Wheeler Ridge, E. of Maricopa, Lewis 1684 (LA); 7.3 mi. N.
of Bakersfield, R17078; S. of Arvin, 450 ft., R17089; near mouth of Kern R.
canyon, Lewis et al. 1675 (LA; 5 pairs + ring of 4); W. of mouth of Kern R.
canyon, Lewis 1683 (LA; 5 pairs + ring of 4); Kern R. canyon, 8.2. mi above road
to Redwood Meadow, Lewis & Wedberg 1109 (LA, RSA; two plants, 7 pairs; one,
5 pairs + ring of 4);8.9 mi. S. of Weldon, R12802; Miracle Hot Springs, R18804;
ca. 5 mi. S. of Walker Basin on road to Caliente, Lewis 1253; 10 mi. N. of Red
Rock Canyon, Munz & Gregory 23313 (RSA); Black Mt. burn, Greenhorn Mts.,
3,900 ft., R18765; head of Short Canyon, R13985. Los ANGELES co.: 0.5 mi. E. of
Pearblossom, Lew?s 1691 (5 pairs + chain of 38 + univalent); 3.7 mi. W. of Little-
rock, Theobald 88; 1 mi. W. of Littlerock, R13994. MADERA co.: 7.6 mi. 8. of Coarse-
gold, 1,300 ft., R18330; San Joaquin Experimental Range, between Coarsegold and
O’Neals, Lewis 1296 (2 plants). MONTEREY co.: 6 mi. N. of Mission San Antonio,
Breedlove 5579; 1.5 mi. W. of Bradley, R18205. sAN BERNARDINO CO,: 0.3 mi. N. of
junction of Hwy. 138 with Hwy. 395, Wedberg in 1957 (LA); Barstow, R11894.
SAN LUIS OBISPO co.: 0.4 mi. S. of San Juan Creek 8. of Shandon, 1,150 ft.,R1697 3;
5.5 mi. 8. of Shandon on San Juan Creek, 1,200 ft., R16967; 8.6 mi. W. of Simm-

RAVEN—GENUS CAMISSONIA 323

ler, 1,800 ft., Breedlove 2061; W. edge of Carrizo Plain, Breedlove 2085. SANTA
BARBARA CO.: 11.8 mi. W. of New Cuyama, 2,100 ft., Breedlove 1939. TULARE CO.:
2 mi. 8. of Tipton, 500 ft., R17063.

Two early reports of chromosome number in this species were made
by Johansen (Proc. Nat. Acad. Sci. U.S. 15: 884. 1929) but no localities
or vouchers were cited.

Plants of this taxon are abundantly visited by oligolectic bees,
especially Andrena spp. The following populations were found to
consist of self-incompatible individuals: 7.6 miles south of Coarse-
gold, 1,300 ft., Madera County, R18330 (nine plants, tested in field) ;
Black Mountain burn, Greenhorn Mountains, 3,500 ft., Kern County,
R18765 (one plant, in greenhouse); 1.5 miles west of Bradley,
Monterey County, R18205 (12 plants, tested in field); 3 miles south-
west of Livingston, Merced County, R18809 (12 plants, tested in
field).

Populations of Camissonia campestris from the foothills of the
Sierra Nevada tend to have relatively small flowers; but these plants
are probably all self-incompatible, judging from the results with the
Madera County population mentioned above. Some populations from
relatively high elevations in the Greenhorn and Tehachapi Mountains
of Kern County consist of plants with relatively broad leaves, but the
leaves are attentuate at the base and the plants do not closely resemble
C. sierrae from farther north. Such collections are (all from Kern
County, California): Black Mountain burn, Greenhorn Mountains,
3,500 ft., R18765 (self-incompatible; see above); Wagy Creek Fire
Road, east side of Greenhorn Mountains, Hardham 3682a (LA);
east slope of Bear Mountain, Tehachapi Mountains, 6,200 ft., Twvssel-
mann 9633 (DS). The status of these plants should be further
investigated.

Oenothera cruciata, as recognized by Munz (N. Amer. FI. II. 5:
157. 1965), consists of a mixture of late-season plants of Camissonia
campestris subsp. campestris, whence the chromosome count, n=7;
relatively large-flowered plants of the hexaploid C. contorta; and the
Clear Lake, Lake County, California, populations of the tetraploid
C. lacustris.

Hybridization between Camissonia campestris and the hexaploid
C. contorta was studied 1.5 miles due west of Bradley, Monterey
County, California, on April 20, 1963. Here both parents were common
on a sunny, west-facing hillside in open oak woodland, with C. cam-
pestris predominant in the more sandy localities and C. contorta every-
where. Widely scattered intermediate individuals made up a very small
proportion of the population, with some 15 collected among many
thousands of the parents. Ten of these were studied cytologically,
with observations of meiosis possible in eight. Of these, five had

324 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

2n—=28 chromosomes and were presumably F, individuals. The maxi-
mum configuration observed was a chain of 8 chromosomes, a ring of
4, two chains of 4, and 4 pairs, two of which were ring bivalents and
two heteromorphic rod bivalents. Less extensive chromosome con-
figurations were observed in most cells of most plants, with trivalents
rather frequent and univalents occasional. These observations clearly
show that the complete lack of multivalent formation in the hexaploid
is under genetic control, which breaks down in the hybrids. Pollen
stainability in these presumed F, individuals was as follows (samples
of 200 grains): 0 percent (one plant), 15.5 percent, 20.5 percent, 22.5
percent, 42 percent.

The remaining three plants on which chromosomal observations
were made had 2n=21 and were presumably backcross individuals
with C. campestris as the recurrent parent. One of these had a ring of 4
chromosomes, a chain of 4, 6 pairs (including 3 ring pairs), and a
univalent, with 7 percent stainable pollen; the second had a chain of 4,
a chain of 3, 6 pairs, and 2 univalents, with 0.5 percent stainable pollen;
and the third had 2 chains of 3, 7 pairs, and a univalent, with 4 percent
stainable pollen. The petals in these plants were 5-5.5 mm. long, in-
stead of 4-4.5 mm. as in the presumed F, individuals. Despite their
low fertility, these individuals strongly suggest the possibility of
interspecific hybridization enhancing the variability of C. campestris.
It is not surprising that no backcrosses with C. contorta were obtained
in this small sample, for C. contorta is largely autogamous, C. cam-
pestris self-incompatible and outcrossing. Thus backcrossing to
C’. campestris is far more likely than to C. contorta.

Another apparent F, hybrid plant was studied from 2.9 miles east of
Paso Robles, San Luis Obispo County, California, Keck 2110 (DS).
This completely pollen-sterile individual was mounted on a herbarium
sheet with plants of the putative parents. Another similar mixed
collection with one apparent F, hybrid was collected about 1 mile
southeast of Havilah, Kern County, California, 975 m., Coville &
Funston 1082 (US).

An unusual individual that was presumably not of hybrid origin
is S. B. & W. F. Parish 1304A (GH), collected with normal individuals
of C. campestris (1304). In this plant, the petals are narrowed greatly
in their upper two-thirds, thus resembling an exaggerated form of the
sort of petals found occasionally in C. hirtella (sect. Holostigma).

Camissonia campestris subsp. campestris grows sympatrically with
C. kernensis subsp. kernensis, C. kernensis subsp. gilmanii, C. strigulosa,
C’. integrifolia, and C. contorta. It is one of the only two diploid species
in that part of sect. Camissonia in which the sepals are reflexed in
pairs, and thus may have contributed to the origin of the tetraploids
C’. lacustris, C. benitensis, C. strigulosa, C. dentata, and C. integrifolia;

RAVEN—GENUS CAMISSONIA 325

these species have very likely had a common origin at the tetraploid
level. Judging from the fact that the hexaploid C. contorta occupies the
hot, dry San Joaquin Valley of California, almost to the exclusion of
the tetraploid C. strigulosa, which is restricted to the foothills and
margins of the valley, it appears likely that C. contorta (at least in
this area) is an allohexaploid combining genomes from the diploid
C. campsteris (abundant with it in the valley) and the tetraploid C.
strigulosa. Whether the hexaploid populations found north and east
of the area of C. campestris originated in this way or by the functioning
of an unreduced gamete in a tetraploid plant cannot at present be
determined. It is likely, however, considering the sort of pairing ob-
served in natural hybrids between the diploid C. campestris and the
hexaploid C. contorta, that a hexaploid formed by the functioning
of such an unreduced gamete would be pair-forming, and therefore
probably fertile, from the time of its origin.

44b. Camissonia campestris subsp. obispoensis Raven, subsp. nov.

A subsp. campestri differt: foliis anguste ellipticis, grosse serratis ;
ramis persaepe decumbentibus. Chromosomatum numerus gameticus,
n=7.

TypE: Opening in chaparral on sandy soil, 1.8 miles west of Bee
Rock, San Luis Obispo County, California, 5 May 1963, P. H. Raven
18301 (DS 520310).

DistRIBUTION (Figure 57): Marine sand deposits in openings in
chaparral and oak woodland, endemic to central coastal California
from southern Monterey County through northern and western San
Luis Obispo County to northwesternmost Santa Barbara County;
from ca. 300-1,600 ft. alt.

Vouchers for chromosome number (5 individuals, 5 populations),
n=7:

U.S.: CALIFORNIA: MONTEREY Co.: E. edge of San Antonio Valley, Santa Lucia Mts.,
1,350 ft., Breedlove 2268; Bradley, R18306; Big Sandy Valley 2 mi. N. of Indian

Valley Road, R18290. sAN LUIS OBISPO co.: 1.8 mi. W. of Bee Rock, R18301;
Atascadero, R18278.

REPRESENTATIVE SPECIMENS EXAMINED:

U.S.: CALIFORNIA: MONTEREY co.: Big Sandy, N. of San Miguel, Hardham
10823 (DS); Big Sandy, 3 mi. NE. of Indian Valley, N. of San Miguel, Hardham
in 1962 (DS); Lowes Canyon, 15 mi. N. of San Miguel, Hardham 3332 (CAS, LA),
R18296 (DS); ca. 3 mi. N. of summit of Hog Canyon Road 8. of Parkfield, R18300
(DS); Cholame Creek (upper end), Jepson 15903 (JEPS); 28.1 mi. NE. of San
Miguel, Big Sandy to Parkfield road, Hardham 10773 (DS, RSA). SAN LUIS
Osispo co.: Nacimiento Dam to Bradley-Pleyto road, Hardham 4582 (SBBG);
Cantinas Creek, near N. end of Nacimiento L., Hardham 4567 (LA, SBBG); sand
dunes on top of Tierra Redonda, Hardham 1859 (LA, SBBG); 1.6 mi. W. of Bee
Rock, Hardham 2259 (LA); 5 mi. ESE. of Bryson, 1,500 ft., Graham 379 (UC);
Oak Flat Road intersection with road to Adelaida, Hardham 2058 (LA); upper

326 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Navajo Creck, Hoover 6898 (CAS); Yaro Creek district N. of Pozo, Hoover 6993
(CAS); Chimney Rock Ranch, South Road to Adelaida, Hardham 829 (LA); Santa
Margarita, Summers in 1886 (ARIZ, POM), Wall in 1933 (CAS, POM); upper
Arroyo Grande, Hoover 6860 (CAS, RSA); road from Arroyo Grande to Huasna,
Eastwood 14988 (CAS). SANTA BARBARA Co.: 3 mi. NE. of Mt. Solomon, 800 ft.,
Lee 241 (UC); Cuyama Valley, Pollard in 1953 (CAS, RM).

This distinctive local endemic intergrades completely with subsp:
campestris wherever the ranges of these two taxa approach one
another. A series of hybrids between C. campestris subsp. obispoensis
and C. strigulosa is represented by Hardham 3240 (LA, RSA, SBBG),
from a damp grain field at Bee Rock, northern San Luis Obispo
County, California. The fertility of four representative putative F,
hybrids, based on the stainability of a sample of 200 pollen grains
from each in cotton blue in lactophenol was, respectively, 11, 1.5,
6.5, and 9 percent. A similar population is represented by Hardham
10823 (DS), from the Big Sandy, north of San Miguel, 21.2 miles
north of the Salinas River bridge, Monterey County, California,
where it grew on the borders of the road. This collection includes two
plants of C. campestris subsp. obispoensis, one of C. strigulosa, and
two hybrids, one of which had a pollen stainability of 0.5 percent,
based on a sample of 200 pollen grains.

An apparent backcross to this subspecies from a hybrid between
it and the hexaploid C. contorta was found with the presumed parents
at Atascadero, San Luis Obispo County, California (R18281). This
individual had 7 pairs of chromosomes, 3 univalents, and a chain of
4;it was not setting any seed.

45. Camissonia sierrae Raven, sp. nov.

Herba annua hirsutulosa humilis, plerumque ramis multis e basi:
caules tenuissimi, ad 15 cm. longi. Folia lanceolata vel anguste ovata,
aliquando elliptica, 0.5-1.8 em. longa, 0.2-0.5 cm. lata, subintegria,
apice acuta, basi rotundata vel obtusa. Inflorescentia pilis glandulosis
brevioribus admixta. Hypanthium 1.3-2.2 mm. longum, ostio 1.2-2.2
mm. diametro, intus in dimidio inferiore hirsutulosum, summum
puncto rubro prope basin omnium filamentorum. Sepala 3—4.2 mm.
longa, 1-1.3 mm. lata, binatim reflexa. Petala 4-7 mm. longa, 3.5-5
mm. lata, unumquidque prope basin punctis duabus rubris maculatum.
Filamenta staminorum episepalorum 2.4-3.2 mm. longa, illa epi-
petalorum 1.2-2 mm. longa; antherae 0.6-1.2 mm. longae. Stylus
4.5-7 mm. longus; sub anthesi insuper antheribus minime portatum
vel antheris staminorum longiorum stigma 0.6-0.8 mm. diametro,
circumnexum. Capsula tenuissima, 2-3 em. longa, ca. 0.5-0.7 mm.
crassa, sessilis. Semina his C. campestris similia. Chromosomatum
numerus gameticus, n=7. Plus minusve autogama.

Tyrr: Meadow in forest dominated by Pinus ponderosa, 0.7 miles
west of junction in Oakhurst on road to Mariposa, 2,400 ft., Madera

RAVEN—GENUS CAMISSONIA 327

County, California, 7 May 1963, P. H. Raven 18343 (DS; isotypes,
BM, GH, NY, RSA, US).

DistTRIBUTION (Figure 57): Carpeting the ground locally in open
forest of Pinus ponderosa and at the upper limits of the Pinus sabini-
ana-Quercus douglasii forest, from 1,800—4,000 ft. alt., Sierra Nevada
foothills of central California, in central Mariposa and Madera
Counties.

Vouchers for chromosome number (2 individuals, 2 populations)
n=7.

U.S.: CALIFORNIA: MADERA Co.: 0.7 mi. W. of Oakhurst, R18343 (type col-
lection); 4.3 mi. W. of Oakhurst, R18345.

ADDITIONAL SPECIMENS EXAMINED:

U.S.: CALIFORNIA: MADERA Co.: 3.7 mi. W. of Oakhurst, Thorp 63-10 (DS);
S. end of Oakhurst, Thorp 63-12 (DS); the Falls, Bass Lake, 3,450 ft., H. & M.
Dearing 5955 (DS, SBBG); Bass Lake, H. & M. Dearing 6018 (SBBG, UCSB).
MARIPOSA co.: Yosemite Valley, Corring in 1884 (GH), Bioletts in 1900 (NY),
Head in 1906 (CAS), 4,060 ft., Wright & Patchett in 1917 (JEPS); Royal Arches,
Yosemite Valley, Jepson 10478 (DS, JEPS); Little Yosemite, Flinn in 1925 (F);
2 mi. off state highway, 7 mi. below Mariposa on road to Ben Hur, 1,800 ft.,
Wolf 4836 (COLO, POM, RSA, SD, WS, WTU).

Camissonia sierrae is a delightful and distinctive endemic found
above the upper limits of C. campestris, its closest relative, in the
central Sierra Nevada of California. The latter species occurs locally
in much drier habitats. Eighteen plants of Thorp 63-10, cited above,
were tested for pollen-tube growth and found to be self-compatible,
in contrast to all known populations of C. campestris. Despite this,
C. sterrae is probably outcrossed to a large extent, and is abundantly
visited by bees, including a large species of Andrena (Onagrandrena).
Its stigma is often held above the anthers and this, together with
the red markings at the summit of the hypanthium and on the petals,
is highly correlated with outcrossing in Camissonia, as is the abundant
pubescence in the lower half of the hypanthium within and near the
base of the style.

This species is very probably a derivative of C. campestris, but is
usually easily distinguished by its habit (ef. fig. 58) and much broader,
relatively shorter leaves. The populations from Mariposa County
have narrower leaves and a laxer habit than those from Madera
County, but form a distinctive morphological and ecological unit
with them. Camissonia sierrae has been observed growing sympatri-
cally with C. contorta at the type locality of the former.

45a. Camissonia sierrae subsp. sierrae Fiaure 58
All of the preceding remarks apply to this subspecies. It has been

necessary in this instance to depart from the usual format of this

paper, as both subspecies are being described as new at this time.

328 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

D >

Ficures 58-60.—Species of Camissonia sect. Camissonia, X 34: 58, C. sierrae subsp. sierrae
(Raven 18343, holotype, DS); 59, C. sierrae subsp. alticola (Russell 75, DS); 60, C.
benitensis (Raven 15084, holotype, DS).

RAVEN—GENUS CAMISSONIA 329

45b. Camissonia sierrae subsp. alticola Raven, subsp. nov. FIGureE 59

A subsp. sierrae differt: floribus parvioribus; hypanthio 1-2.2 mm.
longo, ostio 0.9-2.2 mm. diametro, intus subglabro; sepalis 1.2-3 mm.
longis, 0.8-1.5 mm. latis; petalis 2.2-4 mm. longis, 1.8-2.2 mm. latis,
immaculatis; filamentis staminorum episepalorum 1.8-2.8 mm. longis,
illis epipetalorum 0.8-1.8 mm. latis; antheris 0.6-0.9 mm. longis;
stylo 2.8-5 mm. longo, glabro; stigmate 0.6—-0.8 mm. diametro, sub
anthesi antheris cireumnexo. Chromosomatum numerus gameticus,
n=7. Autogama.

Typsr: Grassy pocket of soil in seepage over granite ledge 100 yards
north of Mono Hot Springs Campground, in yellow pine forest, 6,500
ft., Fresno County, California, 8 June 1963, D. EF. Breedlove 5212 (DS;
isotypes, BM, GH, NY, RSA, US).

DistTrRIBUTION (Figure 57): Shallow soil on granite outcrops, yellow
pine forest, in the Sierra Nevada of northeastern Fresno County,
California, from 6,500—7,700 ft. alt.

Voucher for chromosome number (1 individual, 1 population), n=7:

U.S.: CALIFORNIA: FRESNO co.: Mono Hot Springs, Breedlove 5212 (type
collection).

ADDITIONAL SPECIMENS EXAMINED:

U.S.: CALIFORNIA: FRESNO co.: Near Bear Dam, 7,300 ft., R7157 (CAS);
Mono Hot Springs, 6,500 ft., R6217 (CAS), S. & C. H. Quibell 2326 (DS), Ferguson
430 (DS, JEPS); between Florence L. and Blaney Meadows, 7,500 ft., R4284
(CAS); Florence L., Russell 75 (DS); 3/8 mi. above E. end of Vermillion Valley,
ca. 7,700 ft., C. H. & E. M. Quibell 2670 (herb. Fresno State College).

A distinctive local race, Camissonia sierrae subsp. alticola is obviously
very closely related to C. sierrae subsp. sierrae but is highly autogamous
and occurs at much higher elevations. I have seen a collection from
Lake Merced on the Merced River of Mariposa County, 7,200 ft.,
Jepson 3181 (JEPS), which is very similar to Fresno County plants
of subsp. alticola; additional collections from the Merced River
drainage basin would make it possible to evaluate the relationship
between these two series of populations.

46. Camissonia lacustris Raven, sp. nov. FicureE 61

Oenothera dentata var. campestris sensu Munz, Bot. Gaz. 85: 258. 1928; pro
parte.

Oenothera contorta var. epilobioides sensu Munz, Bot. Gaz. 85: 256. 1928; pro
parte.

Oenothera dentata sensu Munz, N. Amer, FI. II. 5: 157. 1965; pro parte.
Oenothera cruciata sensu Munz, Amer. FI. II. 5: 157. 1965; pro parte.
Oenothera campestris subsp. campestris sensu Munz, N. Amer. FI. II. 5: 158.
1965; pro parte.
Herba annua C. strigulosae persimilis. Plantae pilis longis erectis
dense vestitae; inflorescentia glanduloso-pubescens. Hypanthium
intus in partem dimidiam pubescens. Sepala (3-) 3.8-5.5 mm. longa.

330 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Ficures 61-62.—Species of Camissonia sect. Camissonia, X %: 61, C. lacustris (Raven
18164, holotype, DS); 62, C. integrifolia (Raven 12806, holotype, DS).

RAVEN—GENUS CAMISSONIA 331

Petala (4-) 4.5-7 mm. longa, (2.8-) 3-6.5 mm. lata, plerumque
unumquidque prope basin punctis duabus rubris maculatum. Fila-
menta staminorum episepalorum 2.5-3.5 mm. longa, illa epipetalorum
1.7-2.5 mm. longa; antherae 0.8-1.3 mm. longae. Stylus (3.5-) 4-7
mm. longus, prope basin pubescens. Chromosomatum numerus
gameticus, n=14. Autogama.

Typx: In open grassland of serpentine hill 0.2 mile west of State
Highway 29 near Lakeport, Lake County, California, 13 April 1963,
P. H. Raven 18164 (DS).

DistrisuTion (Figure 63): Open grassland of central California,
in two disjunct areas: serpentine soil along the northwest side of
Clear Lake, Lake County, 1,400-1,800 ft. alt.; Sierra Nevada foot-
hills from Eldorado to Fresno County, 600-5,000 ft., scattered.

Vouchers for chromosome number (5 individuals, 4 populations),
n=14:

U.S.: CALIFORNIA! LAKE CO.: 0.2 mi. W. of Lakeport, R18164 (type collection),

Breedlove 4546. MARIPOSA cO.: Bootjack Road at Chowchilla School, 2,900 ft.,
R18346. TUOLUMNE co.: 1 mi. NE. of Tuolumne City, Breedlove 4817; Carnegie

Ficure 63.—California, showing ranges of some tetraploid (n=14) species of Camissonia
sect. Camtssonia: northern limits of the widespread C. strigulosa indicated by a dotted
line; O=C. lacustris, with @=tetraploid chromosome counts; X= C. benitensis; A=C.
integrifolia, with A= tetraploid chromosome count.

332 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

REPRESENTATIVE SPECIMENS EXAMINED:

U.S.: CALIFORNIA: CALAVERAS co.: Mokelumne Hill, Blaisdell (GH, US).
ELDORADO co.: N. fork of the Cosumnes R. NW. of Youngs, R9107 (CAS, RM).
FRESNO Co.: Pine Flat, 600 ft.. C. H. & E. Quibell 1712 (NY, RSA); 4 mi. W.
of Pine Ridge, Breedlove 5275 (DS). Lake co.: Cold Creek Canyon, 1,800 ft.,
Schulthess in 1928 (POM); near Lakeport, Baker 3067 (CAS, NMC, POM, UC);
Kelseyville, Blankinship in 1923 (RSA). Mariposa co.: Near Leconte Memorial,
Yosemite Valley, 4,000-4,500 ft., Abrams 4426 (DS, GH, NY); Camp Curry,
Stone in 1915 (PH); Little Yosemite, Smith in 1908 (ARIZ, PH). TuoLUMNE
co.: Hog Ranch, Hetch-Hetchy Valley, 4,700 ft., Grant 974 (POM); Phoenix L.,
Grant 946 (JEPS).

Camissonia lacustris is very closely related to and probably derived
from the widespread tetraploid C. strigulosa. The latter occurs in the
Sierra Nevada north only to Kern County and in the North Coast
Ranges only on the immediate coast to the vicinity of Bodega Bay.
Thus C. lacustris occupies a series of widely scattered enclaves in
the area now within the range of the hexaploid C. contorta. It is of
interest that the northernmost population of tetraploids in this
section in the Coast Ranges, that near Clear Lake, Lake County,
occurs on serpentine: marginal populations of plants often occur on
substrates unusual for the species as a whole, and such a situation is
ideal for the derivation of edaphically restricted endemics (cf. Raven,
Evolution 18: 336-338. 1964). In Camissonia contorta, the hexaploid
which occurs in the same areas, often a third or more of the pollen
grains are 4-pored, whereas in C. lacustris and other tetraploids of
sect. Camissonia it is rare for as many as 5 percent of the pollen grains
to be 4-pored. The long, spreading trichomes on the stems of C.
lacustris and its relatively large flowers are likewise useful in its
recognition.

In the vicinity of Clear Lake, Lake County, California, Camissonia
lacustris occurs on serpentine soils and the hexaploid C. contorta
nearby on the nonserpentine soils now under moderately heavy
cultivation. The records of C. campestris (Oenothera dentata auct.)
from Lake County apply to C. lacustris, as do some of the populations
included in Munz’ (N. Amer. FI. II. 5: 157. 1965) concept of Oenothera
cruciata (S. Wats.) Munz, a diverse assemblage of unrelated elements
with several different chromosome numbers. Twelve individuals from
the type locality were selfed in the field and found to be self-compatible
by pollen-tube growth; the stigma is surrounded by at least the anthers
of the four longer stamens even in the largest flowered individuals
of this species.

47. Camissonia benitensis Raven, sp. nov. Figure 60

Herba annua C. strigulosae persimilis. Plantae 3-18 mm. altae,
plus minusve purpurascentes, ramis secondariis tenuibus numerosis
ad angulum 90° patentibus, pilis longis 0.2-0.3 mm. longis erectis

RAVEN—GENUS CAMISSONIA 333

dense vestitae; inflorescentia glanduloso-pubescens et villosa. Folia
angustissime elliptica, sparsa, 0.7-2 cm. longa, 0.15-3 em. lata, apice
acuta. Hypanthium ca. 1.2 mm. longum ostioque diametro, intus in
partem dimidiam dense brevipubescens. Sepala 3.2-3.5 mm. longa,
ca. 1.5 mm. lata. Petala 3.5-4 mm. longa, ca. 3 mm. lata, unumquidque
prope basin punctis duabus rubris maculatum. Filamenta staminorum
episepalorum ca. 2 mm. longa, illa epipetalorum ca. 1.2 mm. longa,
unumquidque basi puncto rubro. Stylus 2.1-2.5 mm. longus, glaber;
stigma sub anthesi antheribus circumnexum. Chromosomatum
numerus gameticus, n=14. Autogama.

Type: On serpentine by small stream 5.4 miles from Hernandez
on road to New Idria, along Clear Creek, San Benito County, Califor-
nia, 19 April 1960, P. H. Raven 15084 (DS; isotype, US).

DistriBuTION (Figure 63): Clay soils derived from serpentine,
lower Clear Creek Drainage, San Benito County, California, at an
elevation of ca. 2,000 ft.

ADDITIONAL SPECIMEN EXAMINED:

U.S.: CALIFORNIA: SAN BENITO co.: Along wash of Clear Creek 1.8 mi. from
Hernandez on road to New Idria, R15094 (DS).

This distinctive species is apparently an endemic of serpentine
soils in the vicinity of the type locality. Nonserpentine soils in the
vicinity are occupied by populations of the hexaploid Camissonia
contorta, which species tends to occur in light sandy soil, and expecially
along washes. Both of the collections of C. benitensis cited above were
tetraploid, with 14 pairs of chromosomes at meiotic metaphase I.
Like C. lacustris, it occupies an enclave in an area that is now domin-
ated by the hexaploid, and, also like C. lacustris, it was probably
derived from C. strigulosa.

48. Camissonia strigulosa (Fisch. & Mey.) Raven, comb. nov.

Sphaerostigma strigulosum Fisch. & Mey., Ind. Sem. Hort. Petrop. 2: 50, 1835-

Oenothera strigulosa (Fisch. & Mey.) Torr. & Gray, Fl. N. Amer. 1: 512. 1840.

Oenothera strigulosa var. epilobioides Greene, Fl. Francisc. 216. 1891. Lecto-
type: Near Los Angeles, Los Angeles Co., Calif., 1890, A. Davidson (ND).

Sphaerostigma contortum var. greenei Small, Bull. Torrey Cl. 23: 189. 1896;
nom. subs.

Oenothera contorta var. strigulosa (Fisch. & Mey.) Munz, Bot. Gaz. 85: 255.
1928.

Oenothera contorta var. epilobioides (Greene) Munz, Bot. Gaz. 85: 256, 1928.

Oenothera dentata sensu Munz, Fl. N. Amer. II. 5: 157. 1965; pro parte.

Erect or sometimes decumbent annual, to 5 dm. tall, usually well
branched, with wiry stems and no well-defined basal rosette, the stems
white or later brown, with exfoliating rhytidome; plants strigulose
and often also glandular-pubescent, especially in the inflorescence,
or occasionally glandular-pubescent only and then subglabrous;

334 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

hirsute pubescence sometimes present near the base, but this almost
always accompanied by strigulose pubescence higher on the stems.
Leaves linear to very narrowly elliptic, 0.8-3.5 cm. long. 0.1-0.3
cm. wide, sparsely serrulate. Hypanthium 1.6-2.7 mm. long, 1.1-2.4
mm. across at the summit, moderately to very sparsely pubescent
within in lower half, or more rarely glabrous. Sepals 1.6—4 mm. long,
1-1.65 mm. wide, reflexed in pairs, the short tips of the sepals sub-
terminal and distinct in bud. Petals 2.1-4.2 (-4.5) mm. long, 1.4-3.1
mm. wide, sometimes with two red dots at the base of each one. Fila-
ments of the episepalous stamens 0.9-2 (-2.2) mm. long, those of the
epipetalous ones 0.5-1.3 mm. long; anthers 0.3-0.6 mm. long. Usually
less than 10 percent of the pollen with four pores. Style 2.3-4.8 mm.
long, pubescent or rarely glabrous near the base; stigma 0.8-1.4 mm.
in diameter, surrounded by the anthers of both sets of stamens at
anthesis, and these shedding pollen directly on the stigma. Capsule
straight or somewhat flexuous, 1.5-4.5 cm. long, 0.8-1.3 mm. thick,
subsessile, with a sterile beak usually 1.5-3 mm. long. Seeds 0.6-0.8
mm, long, 0.3-0.4 mm. thick. Gametic chromosome number, n=14.
Autogamous.

Lectotype: Sphaerostigma strigulosum F. M. California. Cult.—
C. A. Meyer” (ex herb. Ledebour), LE. This specimen closely matches
a modern collection from the dunes between the town of Bodega
Bay and Bodega Head, Sonoma County, California, Raven 18246,
which is known to be tetraploid. It was probably grown from seeds
gathered in the vicinity of Fort Ross, Sonoma County, California,
at the northern limit of the species.

Disrripution (Figure 64): Widespread in open sandy grassland,
vicinity of Bodega Bay, Sonoma County, California, south through
the Coast Ranges to the vicinity of Santa Eulalia, at about 30°43’
N. lat. in Baja California; common in cismontane southern Cali-
fornia, and in the Tehachapi Mountains, through which it reaches
the southern Sierra Nevada of Kern County, California; Santa Rosa
Island. A single collection supposed to be from Fort Huachuca,
Arizona, has been seen, but the presence of this species in Arizona
should be confirmed; this collection might represent a short-term
introduction, for example. From sea level to 6,700 ft. alt.

Vouchers for chromosome number (54 individuals, 50 populations),
n=14:

U.S.: CALIFORNIA: KERN co.: Near Havilah, R18802; 1 mi. 8. of Havilah,
Lewis 1345, 3.4 mi. 8., R20251; 0.5 mi. W. of Weldon, R18799 (sympatric with
C. contorta); Weldon, R18777 (sympatric with C. integrifolia), 18796 (sympatric
with C. contorta); 5 mi. 8. of Weldon, R18793 (sympatric with C. integrifolia) ;
9.7 mi. S. of Weldon, 3,100 ft., R17561; NE. of Lake Isabella, 2,600 ft., Howell
37164 (CAS; progeny= R18844, 19750); Black Mt. burn, Greenhorn Mts., 3,500
ft., Howell 38215 (CAS; progeny= R18784; sympatric with C. campestris and

335

RAVEN—GENUS CAMISSONIA

fa ea —_
Rane
. 0.
yt [a
sae
Ze
ry
i]
tes
aN
ra
ume 3
ta
\
‘, A
\ f .
‘ ( IN —<$<$< —<—
+ ¢ .
‘ \ fi 1
\ \ : /
1, X ! i
i \ f (
‘, a -_
\ Sa, ” ~
‘ —_ \
‘ -
fo “a, oe. 7
! , Ne ee a
fo}
iS
: ‘.,
~ ~~
‘ ss
~
f ~.
1 ‘. ——
oe
' N\ 1
.y .
\ ‘, \ fl
. “N,
\ . \ |!
i ! \ \
\ ' .
“A \. 1
. x. Se
. ..
\ ‘s,
. \
e 2 \.
0 \
\ O \
\ ,
oe) \
\ SD \
» OO ‘ ———<——_
, ©) ~
me's .
\
1. ‘
‘ \ \
NOR 4
\ e. \
$ \ \ .
ee PPO
a ; .
G me ta \o 4
~ fo)

Ficure 64.—Western United States and a portion of northwestern Mexico: O=range of
Camissonia strigulosa, with @=tetraploid chromosome counts (n=14); area of the

closely related hexaploid C. contorta is outlined with dotted line.

295-655 O—68——12

336 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

C. contorta). MARIN cO.: Dillons Beach, R18244. MONTEREY co.: Paloma Creek,
road from Arroyo Seco to Jamesburg, R18217; Big Sandy Valley, R18282.
ORANGE CoO.: 5.6 mi. NE. of Mission San Juan Capistrano, Wedberg 179. RIVERSIDE
co.: Reche Canyon, Wedberg 228; 4.1 mi. W. of Dripping Springs, R14022;
Cajaleo Road 0.1 mi. W. of Mockingbird Canyon Road, Wedberg 197; Machado
Road near Elsinore, Wedberg 192; 7.7 mi. E. of Hemet, Wedberg 199 (herb. San
Diego State College); 1.8 mi. S. of Hemet, Wedberg 201 (herb. San Diego State
College); San Jacinto River wash 2.1 mi. SW. of Cranston Ranger Station, San
Jacinto Mts., Lewis 1250; Hemet Meadows, San Jacinto Mts., R14245; Lake
Fulmor, San Jacinto Mts., R14234; Vanderbilt Flat, San Jacinto Mts., Wedberg
in 1957 (LA, RSA). san piEGo co.: Warner Hot Springs, Kuijt in 1958 (progeny);
Dodge Valley, Wedberg 225 (herb. San Diego State College) ; 2.5 mi. W. of Julian,
H. & M. Lewis 1672 (LA); 2.4 mi. N. of Pala, R16919; 2.2 mi. W. of Campo
R16925. sAN FRANCISCO Co.: Sunset District, Breedlove 4907. SAN LUIS OBISPO CO.:
4.5 mi. KE, of Santa Margarita, R20159; 8.6 mi. W. of Simmler, La Panza Range,
1,800 ft., Breedlove 2059; Morro Bay, R17935; Nipomo Mesa, Breedlove 2099,
4267, R17956; Camatti Creek, R18322; 2.9 mi. E. of San Miguel, R18311;
Simmler to Creston Road, 3.6 mi. W. of junction with Pozo to Simmler Road
H. & M. Lewis 1609 (LA); summit of road between Arroyo Grande and Edna,
Lewis et al. 1664 (LA). SANTA BARBARA Co.: 4 mi. N. of Lompoc, R15509; Juneal
Campground, upper Santa Ynez Valley, 1,800 ft., Breedlove 2234; E. of Torrey
Pines, Santa Rosa I., R15000. SANTA CLARA CO.: Upper Arroyo del Puerto, Mt.
Hamilton Range, R18231, Breedlove 4890. santa cRUz co.: Summit of Quail
Hollow Road near Ben Lomond, R18184; 1 mi. S. of Ben Lomond, Breedlove
4636. SONOMA co.: Between town of Bodega Bay and Bodega Head, R18246.
VENTURA CO.: Cow Spring Campground, 17.6 mi. N. of Fillmore, Topatopa Mts.
3,500 ft., Breedlove 2645.
BAJA CALIFORNIA: 7.5 mi. N. of Santo Tomas, R17048.

REPRESENTATIVE SPECIMENS EXAMINED:

U.S.: CALIFORNIA: ALAMEDA co.: Alameda, Jepson in 1891 (JEPS, NY, UC
US); Oakland, Brewer 2735 (US). conTRA costa co.: West Berkeley, Tidestrom
in 1893 (POM). FRESNO co.: Priest Valley to Parkfield Junction, Thomas 497
(DS; all pollen 3-pored). KERN co.: Miracle Hot Springs, Buckalew 23 (RSA);
Cuddy Valley, near Mt. Pinos, 5,500 ft., Hall & Chandler 6356 (UC); W. end ot
Tecuya Ridge, 6,800 ft., Twisselmann 9577 (CAS, DS); Black Bob Canyon, 4,200
ft., Twisselmann 8339 (CAS, DS); Kernville, 2,650 ft., Howell 37114 (CAS) ;
Temblor Range W. of McKittrick, Eastwood & Howell 4099 (CAS). LoS ANGELES
co.: Browns Flat, San Gabriel Mts., 4,300 ft., Wheeler 915 (CAS, LA, UC); near
Inglewood, Abrams 3237 (DS, GH, LE, NMC, NY, PH, POM, UC, US); vicinity
of Elizabeth Lake, 2,750 ft., Hall 3051 (DS, NY, POM, RM, UC, US; possibly
C. contorta, and one individual with 40 percent 4-pored pollen); summit Mt.
Wilson, 6,000 ft., Grant in 1906 (DS); Tuna Canyon, Verdugo Hills, MacFadden
in 1931 (GH, NY, RM, UC); Santa Monica Mts., 800 ft., Abrams 1261 (DS,
NY, POM); Tujunga Creek at Barley Flats Trail crossing, 4,046 ft., Hwan 7271
(POM). MARIN co.: Dunes near the radio station, Pt. Reyes Peninsula, Howell
21755 (CAS). MONTEREY co.: Tassajara Hot Springs, 1,550 ft., Ferris 8314 (DS,
GH, UC); Pacifie Grove, Patterson & Wiltz in 1907 (DS); near Soledad, Howitt
1510 (CAS); San Simeon, 7. S. Brandegee in 1888 (UC). oRANGE co.: Rancho
Santa Ana, 500 ft., Howell 900 (RSA). RIVERSIDE Co.: Banning, 2,300 ft., Jones
in 1903 (DS, POM); Temeseal Canyon, 8 mi. SE. of Corona, Hitchcock & Muhlick
22136 (DS, UC, WS, WTU); Idyllwild, San Jacinto Mts., 5,400 ft., Spencer 2201
(POM). SAN BENITO co.: San Juan Bautista, Brewer 715 (UC). SAN BERNARDINO

RAVEN—GENUS CAMISSONIA 337

co.: San Bernardino Valley, S. B. & W. F. Parish 81 (DS, F, GH, NY, UC, US);
N. of Cajon Pass, 3,600 ft., Peirson 4514 (RSA); Mojave R. at Deep Creek, 3,000
ft., Lewis 1428 (LA); Redlands, Lewis in 1937 (LA); Baldwin Lake, 6,700 ft.,
Munz 10745 (POM, UC). san piEGo co.: 2 mi. EK. of Pala, Munz 10371 (POM);
Warner Springs, 3,132 ft., Rose 49097 (COLO, RSA); Palomar Mt., 6,000 ft.,
Cooper 1494 (NY); pasture between Middle and South Peaks, Cuyamaca Mts.,
5,000 ft., Munz 9792 (DS, POM); Descanso, Munz & Harwood 7137 (GH, POM,
RM, UC); Jacumba, Munz 8091 (POM); Campo, Eastwood 9383 (CAS); base of
San Miguel Mt., Gander in 1935 (POM). SAN FRANCISCO CO.: Presidio, Lake
Merced, Rose 41282 (ARIZ, C, CAS, COLO, DS, GH, LA, NMC, NY, OSC,
PH, POM, RM, UC, US, WS); Sutter St., San Francisco, Rattan in 1878 (D8).
SAN LUIS OBISPO CO.: Salinas R. 5 mi. N. of Paso Robles, Wiggins 3421 (DS);
Queen Bee Public Camp, La Panza Range, 2,300 ft., Munz 16325 (NY, RSA,
WTU); Price Canyon, Hoover 9171 (CAS); Pismo Beach, Abrams 6513 (D8);
foothills of Caliente Mt., Carissa Plains, Hardham 1786 (LA); Black Lake, 5. of
Arroyo Grande, Hoover 7069 (CAS, RSA). SAN MATEO co.: South San Francisco,
Eastwood in 1894 (GH); Colma, K. Brandegee in 1905 (UC). SANTA BARBARA CO.:
Santa Barbara, Elmer 4001 (CAS, COLO, DS, F, GH, NY, POM, US); 10 mi.
W. of Buellton, Munz 10297 (C, POM); base of Figueroa Mt., Breedlove 209
(UCSB); Big Pine, San Rafael Mts., Hoffmann in 1929 (CAS); E. fork of Lion
Canyon, below Pine Corral Potrero, Sierra Madre Mts., 4,500 ft., Blakley &
Muller 3425 (CAS, DS, JEPS, SBBG, UCSB; might be C. contorta) ; near edge of
Pine Corral Potrero, near Salisbury Potrero, 4,500 ft., Muller & Blakley 1015
(DS, SBBG; might be C. contorta—22 percent 4-pored pollen); Santa Rosa I,
Brandegee in 1888 (UC). SANTA CLARA co.: Mt. Hamilton, Elmer 4625 (ARIZ,
CAS, DS, NY, OSC, POM, UC, US, WS). santa cruz co.: Swanton, Rick in 1912
(DS); Sunset Beach State Park, Thomas 3073 (DS); near Boulder Creek, ca. 650
ft., Thomas 1781 (DS). sonoma co.: Bodega Bay, Heller & Brown 5613 (COLO,
DS, F, GH, NY, PH, POM, RM, US); “Colon Ross,” Wrangell (LE). STANISLAUS
co.: Head of Adobe Creek, Adobe Valley, 1,600 ft., Mt. Hamilton Range, Shar-
smith 3564 (RSA, UC). ventura co.: North Fork of Ventura R. 1.7 mi. above
Wheelers Hot Springs, 1,900 ft., Wolf 6898 (CAS, DS, GH, NY, RSA, UC, WS,
WTU); Red Reef Canyon, Topatopa Mts., 2,800-3,500 ft., Abrams & McGregor
155 (DS, GH, NY, US); Matilija Canyon area above dam, Pollard in 1947 (CAS) ;
Dry Lakes Ridge, Matilija-Sespe Divide, ca. 4,700 ft., Pollard in 1962 (CAS).

BAJA CALIFORNIA: 1.5-2.5 mi. upstream from Rincén, 4.5 mi. NE. of
Santa Catarina, 64 mi. SE. of Ensenada, ca. 4,100 ft., Broder 716 (DS, UCSB);
9 mi. SE. of Tecate, Munz 9477 (POM, UC); 15 mi. NNW. of Rosarito, 3,780 ft.,
Wiggins 10009 (DS, US); Laguna Hansen, 5,450 ft., Sierra Judrez, Gander 3294
(SD); Rancho San José (Meling’s Ranch), 2,300 ft., Wiggins 9771 (DS, UC, US);
Rancho San Matias, 31°17’N. lat., 115°33’W. long., ca. 1,050 ft., Moran 10843
(DS, SD); Santa Eulalia, 30°43’N. lat., 115°20’W. long., ca. 1,850 m, Moran
11135 (DS, SD).

Camissonia strigulosa is a tetraploid species very closely related to
the South American tetraploid C. dentata and at times distinguishable
from it only with difficulty. Hybrids between these two species are,
however, highly sterile (Raven & Moore, MS.), and most individuals
can be distinguished by their habit and flower size, which is often
larger in the South American populations. Camissonia strigulosa is
likewise very closely related to the North American hexaploid C.
contorta, although the latter can usually be distinguished by its posses-

338 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

sion of more than 10 percent 4-pored pollen grains, a character which
is of course directly correlated with its higher chromosome number.
In Kern County, California, in the southern Sierra Nevada, where
the ranges of the two overlap, tetraploid individuals (=C. strigulosa)
have appressed pubescence on the stems, hexaploid individuals
(=C. contorta), spreading pubescence. This distinction does not
always hold true in the Coast Ranges, but is sometimes useful.

Despite the undoubtedly close relationship between these three
species, I prefer to maintain them as distinct, particularly in view of
the other closely related but more easily distinguishable entities that
are accorded taxonomic recognition in this revision. It is likely that
differentiation at the tetraploid level has given rise to the local
endemics C. benitensis and C. integrifolia, and the relatively local
C. lacustris in North America, and to C. dentata following the intro-
duction of a seed of this tetraploid group to South America (cf. Raven,
Quart. Rev. Biol. 38: 151-177. 1963). The three relatively uncommon
North American tetraploids are now at the margins of the range of
the common tetraploid, C. strigulosa, and in two cases completely
surrounded by populations of the hexaploid. As mentioned in the
discussion of the diploid C. campestris, it is likely that some populations
of the hexaploid C. contorta (those in the San Joaquin Valley of
California, especially) have originated following hybridization between
the diploid C. campestris and the tetraploid C. strigulosa; others have
very probably originated by the occasional functioning of unreduced
gametes in tetraploid populations.

Hybridization between C. campestris subsp. obispoensis and C.
strigulosa has been mentioned in the account of the former taxon,
above. From a bud collection made at Miracle Hot Springs, Kern
County, California (R17567), I obtained a meiotic chromosome
configuration of ca. 5 weakly joined pairs, a chain of 3 chromosomes,
and 22 univalents (2n=35), suggesting hybridization between the
hexaploid C. contorta and the tetraploid C. strigulosa at this locality ;
unfortunately, the individual from which these buds were taken was
not identified in the collection. Another individual from the same
collection had 5 pairs, a chain of 3 chromosomes, and 8 univalents,
suggesting hybridization between C. campestris subsp. campestris, also
present at this locality, and the tetraploid C. strigulosa. In addition
to the three taxa just mentioned, (C. strigulosa also has been observed
growing sympatrically with the tetraploids C. pubens (Munz 13357,
RSA) and C. integrifolia; hybridization with the latter is discussed
under the treatment for that species, and with the diploid C. kernensis
subsp. kernensis.

A single collection of what appears to be this species is supposed
to have been made at Fort Huachuca in southern Arizona (May

RAVEN—GENUS CAMISSONIA 339

1892, Wileor, NY; Trans. N.Y. Acad. Sci. 14: 37. 1894). If the
collection is properly labeled, the plant may have been a temporary
introduction at this locality, perhaps with feed for animals; it is
not otherwise known from Arizona.

49, Camissonia dentata (Cav.) Reiche, Fl. Chile 2: 264. 1898.3

South American plants very similar to the North American C.
strigulosa, but differing in most cases by a combination of the following
characteristics. Plants ascending, decumbent, or prostrate. Leaves
linear to broadly elliptic, 0.3-2.8 cm. long, 0.07—0.45 cm. wide, usually
denticulate but more rarely subentire. Hypanthium 0.8-3 mm. long.
Sepals 1-4(-5) mm. long, 0.7-2 mm. wide. Petals 1.5-6(-8) mm. long.
Filaments of the episepalous stamens 0.5-1.8 mm. long, those of the
epipetalous ones 0.5-1.3 mm. long; anthers 0.5—-1.3 mm. long. Style
0.8-5 mm. long; stigma 0.5-0.8 mm. thick, usually surrounded by the
anthers in anthesis, but elevated above them in the plants with the
largest flowers. Capsule 0.8-3.5 cm. long, 0.6-1.3 mm. thick. Seeds
0.8-1.1 mm. long, 0.3-0.5 mm. thick. Gametic chromosome numbers,
n=14, 13. Self-compatible and usually autogamous.

Distrrpution: In South America, from Arequipa, Peru, southward
west of the Andes to Concepcién Province, Chile, and southeastward in
Argentina to’ Lago Argentino and Deseado in Gobernacién de Santa
Cruz. From sea level to ca. 3,500 ft. alt. throughout most of its area,
but in northernmost Chile and southern Peru mostly between ca.
9,500 and 12,500 ft. alt. A map of the range of this species will be
given by Raven and Moore (MS.).

The extreme variability of this complex species, which occupies
a north-south range of some 2,500 miles in South America, has been
the basis of a great deal of taxonomic confusion. The two recent
treatments of the South American members of Camissonia (Munz,
Physis 11: 266-292. 1933; Farm. Chil. 3-12. 1934) divide the popu-
ulations here assigned to this species into five taxa: Oenothera contorta
var. divaricata (Gay) Munz, O. contorta var. epilobioides (Greene)
Munz, 0. dentata Cav. (var. dentata), O. dentata var. campestris
(Greene) Jeps., and O. micrantha var. acuminata (Phil.) Munz. Cam-
issonia cheiranthifolia, erroneously reported from Chile in its proto-
logue by Gay (FI. Chile 2: 330. 1846) and by Reiche (FI. Chile 2:
263. 1898), was correctly excluded from the flora of South America by
Munz (Farm. Chil. 10. 1934; Rev. Univ. Cat. Santiago 22: 275. 1937).
Munz distinguished Oenothera dentata from 0. contorta on the basis of
the larger flowers of the former, with petals 4-7 mm. long, those of

3 The treatment of this species is by P. H. Raven and D. M. Moore, Department
of Botany, The University, Leicester, England. Moore’s fieldwork in Chile and
Argentina was supported by grants from the National Science Foundation to
Harlan Lewis and from the Royal Society of London to Moore.

340 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

the latter given as 2.5-3.5 mm. long. In our study of a wide range
of Chilean material, however, we have found a complete range of
flower size, the petals ranging from 1.5-6 or even 8 mm. long. With
one exception, this variation does not appear to be geographically
correlated. Collections from the central part of Concepcién Province
tend to have larger flowers than those from elsewhere, although plants
with petals longer than 3 mm. are scattered from Coquimbo to
Concepcién. Sample measurements of petal length are given in the
list of cited specimens.

Munz (1933-34, 1937) separated Oenothera contorta var. divaricata
from var. epilobioides on the basis of the larger size and longer capsules
of the latter, which often had a sterile tip at the apex. We have been
unable, however, to confirm these relationships in our studies of South
American material. Neither have we found the nature of the pubes-
cence to be a reliable or geographically correlated characteristic.
Many of the plants in this group have glandular pubescence in the
inflorescence, but a large number have strigose pubescence, and these
two types are sometimes found mixed in a single gathering. Other
plants are subglabrous.

Oenothera dentata var. campestris was separated from 0. dentata
in Munz’ treatment of the Chilean species (1934) by its narrower
leaves. Such plants are sporadic throughout central Chile and ought
not to be confused with the California diploid Camissonia campestris
for reasons which will be made clear below.

Finally, as regards Oenothera micrantha var. acuminata, we have seen
only a photograph made of the type in Santiago (GH). The plant
shown is exceptional in its long-acuminate upper leaves, but in other
respects there seems to be no reason why it could not be an unusual
form of the complex so widespread throughout South America. It
certainly seems unreasonable, on the basis of this single specimen, to
credit the North American sect. Holostigma to South America. It is,
however, highly desirable that similar plants be re-collected, and they
may exist in Biobfo or neighboring provinces.

Despite the fact that all of the South American populations of this
genus seem to belong to a single species, there are two distinctive
geographical races within this species that deserve formal taxonomic
recognition. The more local and uniform series of populations consists
of a group of heavyset, broad-leaved plants with relatively large
flowers and thick, heavy capsules found along the coast of Concepcién
and the neighboring provinces. This series is described as C. dentata
subsp. littoralis.

We have grown 15 populations of this species, including 2 popu-
lations of subsp. littoralis, in the experimental garden. Of these, 13
had n=14, and 2, one belonging to the series of relatively large-

RAVEN—GENUS CAMISSONIA 341

flowered populations from Concepciédn mentioned above and the
other to subsp. littoralis, had n=13. The two populations with n=13
represent the only naturally occurring aneuploid populations in the
genus, but they are not strongly separated by sterility barriers from
other South American populations with n=14. A summary of our
cytogenetic and experimental studies with this group is given else-
where (Raven & Moore, MS).

It appears likely that Camissonia may have reached South America
only once, and relatively recently, by long-distance dispersal across
the tropics (Raven, Quart. Rev. Biol. 38: 151-177. 1963), a phenom-
enon which has probably occurred twice in Gayophytum, once in
Clarkia, and twice in Boisduvalia, three other genera of annual plants
in the Onagraceae. Camissonia dentata is obviously most closely re-
lated to the North American tetraploid C. strigulosa, from which it
may well have been derived, but the South American entity is far
more variable, and most populations can easily be distinguished from
those of North America. In these respects, the situation is closely
similar to that in Clarkia. In view of this, the high degree of sterility
between North and South American plants (Raven & Moore, MS)
and the degree of morphological distinction between the species
recognized in North America in this assemblage, it appears best to
consider the North and South American plants as specifically distinct.
If this is not done, the oldest name for the collective group would be
Camissonia dentata, a name which (as Oenothera dentata) has usually
been incorrectly applied to the diploid species here called Camissonia
campestris in North America. Should Camissonia dentata be under-
stood in this collective sense, it would probably be best to include
the hexaploid C. contorta in the same species, but it would probably
still be possible to maintain the more local tetraploids C. benitensis
and C. lacustris as species distinct from this complex. Camissonia
integrifolia, a third local tetraploid species, is so distinct morphologi-
cally that it does not really enter into this problem.

49a. Camissonia dentata subsp. dentata
Oenothera dentata Cav., Icon. 4: 67, t. 398. 1797, excl. verbis falsis ‘stigma .
quadrifidum, laciniis ovatis minimis.”’
Camissonia flava Link, Jahrb. Gewachsk. 186. 1818.
Oenothera chamissonis Link, Enum. Pl. Hort. Berol. 1: 378. 1821. Type:
Chile, presumably cultivated at Berlin, and probably destroyed (not seen).
Holostigma argutum Spach, Ann. Nat. Sci. Bot. Paris II. 4: 270. 1835; nom.

subs.

Holostigma heterophyllum Spach, Ann. Sci. Nat. Bot. Paris II. 4: 271. 1835;
nom. subs.

Sphaerostigma chamissonis (Link) Fisch. & Mey., Ind. Sem. Hort. Petrop. 2:
49. 1835.

Holostigma tenuifolium Spach, Ann. Sci. Nat. Bot. Paris IT. 4: 270. 1835; non
Oenothera tenuifolia Cav. 1797. Type: Chile, Bertero (P), not seen. The

342 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Unio Itineraria distribution of Bertero’s 466 and 1190 (BM, NY, 8), from
“Quintero, Quillota, in petrosis aridis planitiei fluvii Cachapual,’’ is a
mixture of large- and small-flowered plants, probably from two different
localities; the small-flowered plants may be isotypes of this species, or the
type collection may be Bertero 467 (BM), from Rancagua, Prov. O'Higgins.

Oenothera chilensis D. Dietr., Syn. Pl. 2: 1285. 1840; non Fisch. & Mey. 1835.

Oenothera heterophylla (Spach) Steud., Nom., ed. 2, 2: 206. 1841; non Spach
1835, nee Nutt. ex Hook. & Arn. 1838. ;

Oenothera minutiflora D. Dietr., Syn. Pl. 2: 1285. 1840. Type: Cultivated,
from Chile, C. A. Meyer (isotype, LE).

Sphaerostigma dentatum (Cav.) Gay, FI. Chil. 2: 326. 1846.

Sphaerostigma heterophyllum (Spach) Gay, Fl. Chil. 2: 327. 1846.

Sphaerostigma tenuifolium (Spach) Gay, Fl. Chil. 2: 327. 1846.

Sphaerostigma divaricatum Gay, Fl. Chil. 2: 238, t. 22. 1846. Type: Sandy
places in the low mountains around Santiago, Chile (not seen).

Sphaerostigma ramosissimum Philippi, Linnaea 33: 68. 1864. Type: Near
Concumen, Aconcagua, Chile, Landbeck (SGO, not seen; photographs,
GH, POM).

Sphaerostigma acuminatum Philippi, Anal. Univ. Chile 84: 629. 1893. Type:
Sandy places near Nacimiento, Bfobfo, Chile, January 1877, Philippi
(SGO; isotype, NY).

Chamissonia acuminata (Phil.) Reiche, Anal. Univ. Chile 98: 481. 1897.

Chamissonia tenuifolia (Spach) Reiche, Fl. Chile 2: 263. 1898.

Chamissonia paradoza var. divari-ata (Gay) Reiche, Fl. Chile 2: 264. 1898.

Oenothera torulosa f. chilensis (D. Dietr.) H. Léy., Monogr. Onoth. 180. 1905

Oenothera contorta var. divaricata (Gay) Munz, Physis 11: 288. 1933.

Oenothera micrantha var. acuminata (Phil.) Munz, Farm. Chil. 11. 1934.

Oenothera dentata var. campestris sensu Munz, Farm. Chil. 10. 1934.

Oenothera contorta var. epilobioides sensu Munz, Farm. Chil. 10. 1934.

Plants erect, decumbent, or prostrate. Leaves linear or very nar-
rowly lanceolate. Capsules 0.6-0.9 mm. thick. Gametic chromosome
numbers, n= 14, 13. Self-compatible and mostly autogamous.

Type: Near Coquimbo and Taleaguano, Coquimbo, Chile, Nov.,
Ruiz & Pavon (MA; photographs, CONC, F). Judging from the
photographs I have seen, the flowers are probably shown a little too
large in the illustration that accompanies the type description; the
stigma was certainly not 4-lobed in this plant.

Disrrinution: That of the species, except coastal Arauco and
Cautin Provinces, Chile.

REPRESENTATIVE SPECIMENS EXAMINED:

ARGENTINA: BurEnos ArrEs: Carmen de Patagones, Miccio-Peralta 67 (DS).
Cuusut: “Rio Carren”’ (?), Skottsberg in 1908 (UPS); ‘“Futalenfa,” Skottsberg in
1908 (8S, UPS); Lago Colue Huapi, Riggs 67 (F, GH). Comovoro Rrvapavia: 50
km. W. of town of Lago Buenos Aires, Eyerdam et al. 24491 (GH, K, UC). Nrv-
QuEN: Neuquén, 240 m, Comber 37 (K); Rio Caliefu, Cabrera 5109 (F, POM);
Piedra de Aquila, 40°02’S. lat., 70°01’ W. long., Senn 4244 (RSA); Cerro Lotena,
900 m, Ammann 83 (F). Rio NeGro: San Carlos de Bariloche, 800 m, Buchtein
in 1905 (US); General Roca, 250-360 m, Fischer 85 (BM, F, GH, K, NY, US);
shore of Lago Nahuel Huapi, Cordini 155 (POM, US). Santa Cruz: Deseado,
Darwin 129 (K); Tehuelches, 300 m, Donat 87 (BM, F, GH, K, 8, UC); 160 km.

RAVEN—GENUS CAMISSONIA 343

S. of Deseado on road to San Julian, Eyerdam et al. 23912, (GH, UC); San Julian,
50 m, Blake 318 (K); 78 km. S. of San Julian on road to Santa Cruz, Rfo Chico,
Eyerdam et al. 23994 (GH, UC); S. shore of Lago Argentino 3 km. E. of Calefate,
270 m, Eyerdam et al. 24279 (GH, K, UC).

CHILE: Aconcaaua: Papudo, FE. W.D. & M M. Holway 65 (GH, US); Los
Andes, Mancilla in 1952 (CONC). Arauco: Isla Santa Maria, Beck 75 (K).
ATACAMA: Sandy washes around El Morro, ca. 14 km. ENE. of Caldera, Worth &
Morrison 16165 (GH, UC); La Varilla, ca. 25 km. N. of Vallenar on road to Capi-
ap6, Worth & Morrison 16262 (GH, K, UC); Cachiyuyo, Ricardi & Marticorena
4885/1270 (CONC). Bfosfo: Candelaria, Arenales de Los Setenta, Junge in 1935
(CONC; petals 5 mm long). Cotcnacua: La Rufina, Rfo Claro, Ricard: in 1951
(CONC); Talearihue, San Fernando, Ricardi in 1950 (CONC). CoNcEPcI6n:
Puente Itata, road from Concepcién to Bulnes, Pfister 810 (CONC; petals 6.5 mm
long); Quilacoya, Pfister 298 (CONC; petals 6 mm long); between Yumbel and
Estacién Yumbel, Marticorena et al in 1959 (CONC; petals 4.5 mm long) ; Pangal
del Laja, W. of Yungay, Marticorena et al. in 1959 (CONC; petals 3.5 mm long),
Ricardi & Marticorena 4991/1375 (CONC; petals 5 mm long). CoqurmBo: Punta
Colorado, Rose & Rose 19340 (NY, US); Paihuano, Valle del Cochiguaz, 1,120 m.
Behn in 1948 (CONC); Parque Lambert, ca. 6 km, NW. of La Serena on road to
Islon, 30 m, Worth & Morrison 16337 (GH, K, UC); Coquimbo, 30 m, Elliot 548
(K; petals 4.5 mm long); Fray Jorge, Mufioz B112 (GH); Los Vilos, Behn (CONC;
petals 3-4 mm long); ca. 5 km. above Los Molles, Moore 215 (DS). Curtco: Los
Quefies, Milner 45 (CONC). Matueco: Fundo Tambillo near Nacimiento, Hempel
7422 (CONC; petals 6-8 mm long), Pfister in 1934 (CONC; petals 2 mm long).
Mavte: Caquenes, Reid (K, NY; some plants with petals 2.5 mm long, others
5.5-6 mm). Nusie: Fundo Santa Ana, Behn in 1937 (CONC); Rfo Cholguan,
road from Yungay to Huépil. Pfister in 1946 (CONC). O’Hiaarns: Callanas, Rfo
Pangal valley, Moore 263 (DS). Santraco: Batuco, over 500 m, Looser 3652
(GH); San Antonio, coast, Claude-Joseph 1449 (US); Esmeraldar, Departmento
Melpilla, Reed 1867/10 (GH). Tauca: Talea, Claude-Joseph 1692 (US). Tara-
pack: Road from Azapa to Chapiquifia, Episcacha, Km. 94, 3,500 m, Ricardi &
Marticorena 4760/1145 (CONC). Vauparafso: Limache, Cuesta La Dormida,
1,200 m, Behn in 1940 (CONC); Valparafso, 60 m, Sandeman 202 (BM, K); Vina
del Mar, Buchtein in 1895 (S). WITHOUT DEFINITE LocALITY: Gay (GH, K,
NY, RSA; ‘‘Sphaerostigma tenuifolium Spach’’).

PERU: Arequipa: Chachani, 3,350 m, Stafford 607 (BM, K); near Chiguata,
3,100 m., Vargas 8093 (RSA); Pampa de Arrieros, 3,750 m., Pennell 13327 (F,
GH, NY, PH); lower slopes of Misti, 3,050 m, Sandeman 4002 (K); Tiabaya,
2,100-2,200 m., Pennell 13096 (PH). WirHovur DEFINITE Locality: Dombey 726
(F).
49b. Camissonia dentata subsp. littoralis Raven & Moore, subsp. nov.

A subsp. dentata differt: foliis latioribus, 0.5—-1.8 cm. longis, 0.2-0.45
cm. latis, crassioribus; plantis prostratis; capsulis crassioribus, 1.2-2.5
cm. longis, 1.1-1.3 mm. latis. Numerus chromosomatum gameticus,
n=14, 13. Autogama.

TypE: Near Concepcién, Concepcién, Chile, 1855, P. H. Germain
(GH; isotypes, BM, F, K). An isotype is illustrated in H. Lév.,
Monogr. Onoth., facing p. 175. 1905.

DisrriBuTION: Sandy soil and dunes on the coast of Concepcién,
Arauco, and Cautin Provinces, Chile; from sea level to a few hundred
feet alt.

344 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

SPECIMENS EXAMINED:

CHILE: Arauco: Laraquete, Moore 290 (DS). Cautin: Bajo Imperial, Moreton
Middleton in 1906 (BM, 8). Concerct6n: San Vicente, Scott Elliott 197 (BM), Junge
in 1934 (CONC): Concepcion, Jaffuel 2906 (GH), Cuming 116 (BM): mouth of
the Bfobfo, Ricard 8520 (CONC); N. side, mouth of the Bfobfo, Junge in 1935
(CONC), Pfister 297 (CONC), in 1941 (CONC), in 1948 (CONC), Ramirez in 1959
(CONC); Tomé, Junge in 1935 (CONC); Coronel, Hill 167 (K); Coronel, Mina
Schwager, Sparre 10111 (CONC); Lengua, in mouth of Estero Lengua in Bahia
San Vicente, Moore 276 (DS). Province UNKNOWN: Taliahua, Philippi (US); I.
Quiriquina, Gunckel 2805 (GH).

This distinctive and fairly localized entity occupies the southern
part of the coastal range of this species in Chile. It has relatively large
flowers, the petals ranging from 3-5.5 mm. long.

A specimen from Concepcién (Elliot 549, GH, K) is spreading in
habit but does not have the broad leaves of subsp. littoralis; one from
San Pedro, Concepcién (Sparre 14229, CONC) is closer to littoralis
but intermediate. Likewise, a single collection that has been examined
from farther north (Santo Domingo, south of Llolleo, Santiago,
Looser 1455 GH) approaches subsp. littoralis, as does one from
Linares, Playa de Pelluhue, Aravena 61 (UC, US).

For some unknown reason, Hector Léveillé (Monogr. Onoth. 175.
1905) considered this taxon a species distinct from his Qnothera
torulosa, in which he included all other members of sect. Camissonia.
He called this species Onothera hyssopifolia Molina, Saggio Chile, ed.
2, 134. 1810, but Molina says of his species “fiore . . . terminale
violetto, di pit. d’un pollice di diametro . . .”’; it seems likely that, as
suggested by Munz (Farm. Chil. 7. 1934), this name refers to some
form of Clarkia tenella (Cav.) H. & M. Lewis (Godetia tenurfolia (Cav.)
Spach). Onothera hyssopifolia, however, was not mentioned by Moore
and Lewis in the most recent revision of the South American species of
Clarkia (Bol. Soc. Arg. Bot. 10: 332-340. 1965).

50. Camissonia integrifolia Raven, sp. nov. FIGuRE 62

Herba annua C. strigulosae persimilis. Inflorescentia dense cinerero-
strigulosa; plantae sparse strigulosa vel subglabrae. Folia linearia,
integria, rare dentibus parvulis obscuris 1-2. Hypanthium intus in
partem dimidiam pubescens. Stylus prope basin pubescens vel glaber.
Capsula stricta, erecta, 4.5-6 cm. longa. Semina 1-1.2 mm. longa,
0.4-0.5 mm. crassa. Chromosomatum numerus gameticus, n=14.
Autogama.

Type: On bank covered with Artemisia tridentata, Weldon, Kern
County, California, 3 May 1958, P. H. Raven 12806 (DS).

DistriBution (Figure 63): Dry, often deserty slopes along the Kern
River from Miracle Hot Springs to the vicinity of Weldon, Kern
County, California; 2,450-3,000 ft. alt.

RAVEN—GENUS CAMISSONIA 345

Vouchers for chromosome number (1 population, 1 individual),
n=14:
U.S.: CALIFORNIA: KERN CO.: Weldon, R18776 (grown in experimental garden).

ADDITIONAL SPECIMENS EXAMINED:

U.S.: CALIFORNIA! KERN CO.: Weldon, 2,450 ft., R17536 (DS), 18791 (DS);
mouth of Kelso Canyon at State Hwy. 178, 2,800 ft., T’wisselmann 7041 (CAS,
DS, SBBG); 5 mi. S. of Weldon, R18792 (DS); 6 mi. from Weldon on road to
Kelso Valley, 3,000 ft., Weatherby 1158 (RSA); 8.3 mi. 8. of Weldon on road to
Kelso Valley, 2,900 ft., R17560 (DS); Miracle Hot Springs, ca. 2,500 ft., Buckalew
23 (RSA).

Camissonia integrifolia is a narrow endemic of a portion of the upper
Kern River drainage. It is distinctive in its entire leaves, long, erect
capsules, and distribution of its pubescence. Raven 17536, cited above,
is predominantly this species but includes two plants of C. strigulosa
and one probable hybrid. A chromosome determination was made of
one plant of this collection, which had an extremely irregular polyploid
meiosis, suggesting hybridity. Progeny of this collection was grown
at Stanford, yielding R18776 (C. integrifolia, n=14), R18777
(C. strigulosa, n=14), and R18778, a single obvious hybrid between
these two species. The hybrid had variable meiotic configurations
ranging from 8 univalents and 10 pairs of chromosomes to 1 chain of
3, 13 univalents, and 6 pairs. It had 11 percent stainable pollen, based
on a sample of 200 grains in lactophenol. Thus C. integrvfolia appears
amply distinct from C. strigulosa, from which it may have been derived,
and grows sympatrically with it, with the occasional formation of
sterile hybrids. These two species were also observed growing sym-
patrically at the locality of R18792, cited above. Camissonia integri-
folia has also been observed growing sympatrically with C. kernensis
subsp. kernensis, C. pubens, C. campestris, and C. contorta.

51. Camissonia contorta (Dougl.) Kearney, Trans. N.Y. Acad. Sci. 14: 37. 1894.
Oenothera contorta Dougl., in Hook., Fl. Bor. Amer. 1: 214. 1834.
Sphaerostigma contortum (Dougl.) Walp., Rep. 2: 78. 1843.

Oenothera dentata var. cruciata S. Wats., Proc. Amer. Acad. 8: 594. 1873.
Lectotype: Sandy places in the Sacramento Valley, Calif., Hartweg 1733
(GH, NY).

Oenothera campestris var. cruciata (S. Wats.) Greene, Fl. Francisc. 216. 1891.

Sphaerostigma campestre (Greene) Small var. minus (“minor”) Small, Bull.
Torrey Cl. 23: 189. 1896.

Oenothera cheiranthifolia var. contorta (Dougl.) H. Lév., Monogr. Onoth. 216.
1905.

Oenothera torulosa f. cruciata (S. Wats.) H. Lév., Monogr. Onoth. 180. 1905.

Oenothera torulosa f. mixta H. Lév., Monogr. Onoth. 180. 1905. Type: Possibly
at MO; cf. A. Nelson, Bot. Gaz. 40: 57. 1905.

Sphaerostigma campestre var. miztum (H. Lév.) A. Nels., Bot. Gaz. 40: 57.

1905.
Oenothera contorta var. typica Munz, Bot. Gaz. 85: 253. 1928.

346 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Oenothera contorta var. strigulosa sensu Munz, Bot. Gaz. 85: 255. 1928; pro
parte.

Oenothera contorta var. epilobioides sensu Munz, Bot. Gaz. 85: 256. 1928;
pro parte.

Oenothera contorta var. contorta; Munz, N. Amer. FI. IJ. 5: 156. 1965.

Oenothera cruciata (S. Wats.) Munz, N. Amer. Fl. IT. 5: 157. 1965.

Oenothera dentata sensu Munz, N. Amer. FI. II. 5: 157. 1965; pro parte.

Very similar to C. strigulosa, but almost always with hirsute
pubescence near the base and often with only hirsute pubescence or
a limited admixture of glandular trichomes in the inflorescence,
very rarely entirely strigulose and glandular-pubescent. Leaves
often broader, to narrowly elliptic, the foliage often more bluish
green than that of C. strigulosa. Flowers often larger, the petals to
5 mm. long and 3.3 mm. wide. Longer filaments to 2.6 mm. long, the
shorter to 1.5 mm. long; anthers to 0.8 mm. long. Often more than
30 percent of the pollen with 4 or 5 pores. Style to 5.1 mm. long.
Seeds 0.7-0.9 mm. long. Gametic chromosome number, n=21.
Autogamous.

Type: On barren sandy soils of the interior parts of the Columbia
River, Washington or Oregon, 1825-6, D. Douglas (K; isotype, BM).

DistripuTion (Figure 65): Open or grassy slopes and flats, usually
on sandy soil: southern Vancouver Island, British Columbia, and
San Juan and Whidbey Islands, northwesternmost Washington;
Klickitat and Walla Walla Counties, southern Washington, and
Adams and Ada Counties, Idaho, south through east-central Oregon
to the vicinity of Reno, western Nevada, and from southwestern
Oregon (Douglas, Jackson, and Josephine Counties) south nearly
throughout California to the southern San Joaquin Valley and border-
ing foothills in Kern County; generally away from the coast from
Sonoma County, California, southward, where replaced by (. stri-
gulosa. From sea level to 7,500 ft. alt.

Vouchers for chromosome number (42 individuals, 40 populations),
n=21:

U.S.: CALIFORNIA: CONTRA COSTA CO.! Antioch, R18142; ca. 1 mi. 8. of Oakley,
R18144. rREsNO co.: 1.6 mi. 8. of Easton, R18328° Fresno, R17067; 3.3 mi. N.
of Selma, R17065. HUMBoLDT co.: Near Arcata, R18373; 1 mi. E. of Dinsmores,
R18367. KERN co.: West of mouth of Kern River canyon, Wedberg & Lewis 1671
(LA); Miracle Hot Springs, R17567 (progeny = R18773), R18805; Isabella,
R18800; 0.5 mi. W. of Weldon, R18798 (sympatric with C. strigulosa) ; Weldon,
R18795 (sympatric with C. strigulosa); Havilah, R17568; Black Mt. burn, Green-
horn Mts., 3,500 ft., Howell 38215 (CAS; progeny = R18785; sympatric with
C. campestris subsp. campestris and C. strigulosa). LAKE Co.: 3 mi. 8. of Kelse y-
ville, Breedlove 5143; 2 mi. NE. of Lower Lake, Breedlove 4540; 9.9 mi. N. of
Lower Lake, R18240; Lakeport R18239, LassEN co.: 4.8 mi. N. of Milford,
R17872. MADERA co.: 7.6 mi. 8. of Coarsegold, R18340; 0.7 mi. W. of Oakhurst,
R18344. MERCED co.: Ca. 20 mi. W. of Merced, Klein 444 (RSA; progeny, DS).
MONTEREY CO.: 1.5 mi. W. of Bradley, R18206: 4.1 mi. N. of Mission San Antonio,

RAVEN—GENUS CAMISSONIA 347

Ficure 65.—Western United States: O=range of Camissonia contorta, with @= hexaploid
chromosome counts (n=21). Northern limits of the closely related tetraploid C. strigu-
losa indicated by dotted line.

348 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Breedlove 2278; 1.6 mi. N. of junction to Jamesburg, Carmel Valley, R18149;
Jolon, R18154; 10 mi. N. of Hunter Liggett Military Reservation Headquarters
on road to The Indians, R18272. prumas co.: 3.1 mi. W. of Vinton, R18418;:
4.8 mi. W. of Vinton, R18420. san BENITO co.: Panoche Creek Campground,
Pinnacles, Breedlove 2182; 1.1 mi. 8. of Cienaga School, near Hollister, R18199.
SAN LUIS OBISPO CoO.: 1.8 mi. W. of Bee Rock, R18302, 18303; Atascadero, R18280.
SONOMA co.: 3 mi. 8. of Sebastopol, Breedlove 3706. TULARE Co.: Posey to White
River, R20239; Kern River Canyon 6 mi. 8. of Roads End P.O., 3,100 ft., Howell
33134 (CAS; progeny = R18770). NEVADA: STOREY co.: 4.7 mi. N. of Carson
City, R18544. wasHoE co.: Ca. 5 mi. N. of Reno, Mosquin & Gillett 5310 (DAO;
progeny = R18780), OREGON: HARNEY co.: 5.3 mi. N. of Narrows, R18455.
JACKSON Co,: Just N. of Central Point, R18381. wasco co.: 2.2 mi. W. of Sherman
Co. line along Columbia R., R18385.

REPRESENTATIVE SPECIMENS EXAMINED:

CANADA: Saanich Spit, Hastham in 1942 (NMC, UBC); Cedar Hill, near
Victoria, Macoun in 1887 (NMC).

U.S.: CALIFORNIA: AMADOR co.: 2 mi. NNE. of Buena Vista, ca. 300 ft., Howell
29808 (CAS). BUTTE co.: Chico, Copeland 3033 (POM); hills 8 mi. N. of Oroville,
Heller 11330 (CAS, DS, GH, NY, OSC, PH, UC, US, WTU). caLaveras co.: 2
mi. W. of Camanche, 400 ft., Belshaw 1978 (UC); Mokelumne Hill, Blaisdell
(CAS). conrra costa co.: Alhambra Valley, Rattan in 1887 (DS); Nortonville,
Bacigalupit 4977 (JEPS). ELpoRADO co.: Coloma, Reed in 1941 (UC). FRESNO Co.:
Hwy. 180 just below 4,000 ft. marker, Quibell 1963 (RSA); Shaver, 5,200 ft., Hall
& Babcock 292 (UC); Millwood Meadow, near Sequoia Lake, 5,100 ft., Simonian
482 (herb. Fresno State College); Clovis, Thompson in 1902 (DS, POM). num-
BOLDT co.: Klamath R. at mouth of Slate Creek, 400 ft., Tracy 16244 (UC).
KERN CO.: Cedar Canyon just below mouth of Fulton Creek, Greenhorn Range,
3,250 ft., T’wisselmann 8495 (DS); Walker Basin, 3,400 ft., Twisselman, 9518
(CAS, DS, RSA); 16 mi. 8. of Bodfish, Thorne 31736 (DS, RSA); Poso Creek,
Hall & Babcock 5012 (DS, GH, PH, UC). taxes co.: Dashiels, Mt. Sanhedrin,
Eastwood in 1925 (CAS); W. base of Snow Mt., Ackley in 1965 (CAS); Jordan
Park, Jussel in 1932 (POM). LAssEN co.: Milford, Baker in 1890 (UC); Chat,
5,000 ft., Jones in 1897 (POM). MaprERA co.: N. side of Millerton L., 550 ft.,
Bacigalupt 7672 (JEPS); Bass Lake, 3,300 ft., Quibell 2452 (POM); 5 mi. NE.
of Madera, Howell 41073 (CAS). Mariposa co.: Just 8. of Mormon Bar, 1,800 ft.,
Howell 40571 (CAS). mENpocINo co.: Sand in the Russian R., Ukiah, Bolander
3857 (F, NY); Navarro, Byxbee in 1895 (JEPS); near Longvale, 1,500 ft., Tracy
9924 (UC, WTU). MERcED co.: Delhi, Howell 35293 (CAS); 2 mi. SW. of Living-
ston, Stage 59-59 (RSA, UC); Merced R. bottom between Snelling and Hopeton,
Howell 1049 (CAS). mopoc co.: Parker Creek SE. of Alturas, 5,200 ft., Payne
637 (JEPS); sink along highway between Malin and Canby, several miles N. of
quarantine station, Baker 8309 (POM). MonTEREY co.: Trail to Ventana Double
Cone, 4,000 ft., Hardham 10127 (DS); 3 mi. NE. of Pleyto Well, SW. of Bradley,
550 ft., Graham 266 (RSA, UC). napa co.: Howell Mt., Jepson 14478 (JEPS) ;
Capell Valley, 700 ft., Kamb & Chisaki 1761 (UCSB). ptumMas co.: Near Beck-
wourth, 4,880 ft., Howell 37800 (CAS). sAN BENITO co.: 5.4 mi. 8S. of Willow
Creek School on road to the Pinnacles, Ferris 8347 (CAS, DS, GH, POM, UC);
Panoche Pass, Abrams & Borthwick 7984 (DS, NY, POM); 3.7 mi. from Hernandez
on road to New Idria, R15088 (DS). san soaQquin co.: 8.8 mi. S. of Stockton,
R9132 (CAS); Mokelumne, 200 ft., Roseberry 160 (UC); Tracy, Baker 2775
(CAS, F, GH, NY, POM, RM, UC, US). san Luts optspo co.: Palo Prieto

RAVEN—GENUS CAMISSONIA 349

Canyon, Ferris 12816 (DS); Atascadero, Hardham 3054 (LA, SBBG). santa
CLARA co.: Stanford University, Elmer 2368 (NY, US). santa cruz co.: Bielawski
Look Out Station, 3,270 ft., Thomas 1514 (DS, OSC); Deer Ridge Farm, summit
of Santa Cruz Mts., 2,800 ft., Pendleton 371 (POM, UC). sHasta co.: Near
McArthur, Eastwood & Howell 7955 (CAS, POM); Hat Creek ca, 20 mi. W. of
Fall River Mills, Hitchcock 6554 (POM, WTU); near Redding, Heller 7888
(GH, NY, PH, US). stskryou co.: French Creek Logging Road W. of Calla-
han, 3,500-4,000 ft., Ferris & Lorraine 11757 (DS, RSA, UC, WTU); Scott
Valley, 2,900 ft., Parker 406 (NMC, UC); Yreka Creek, Siskiyou Mts., 2,800 ft.,
Wheeler 3527 (POM, RSA, UC); near Yreka, Greene in 1876 (F). SONOMA CO.:
Pitkin Marsh near Forestville, Rubtzoff 1027 (UC). sTANISLAUS co.: 8 mi. E. of
Oakdale, Hoover 3956 (UC); Turlock, Fosberg in 1927 (POM); Modesto, Hoover
889 (UC). suTTER co.: South Peak, Marysville Buttes, Jepson 14478 (JEPS,
UC). TEHAMA co.: 6 mi. W. of Paskenta, Bailey in 1938 (JEPS). TRINITY Co.:
Three Forks of Mad River, 2,900 ft., Tracy 10175 (DS, UC). TuLareE co.: Sand
along Tule R., Dudley in 1897 (DS); Johnsondale, 4,670 ft., Howell 37343 (CAS).
TUOLUMNE co.: Tuolumne R. just below junction of Middle and South Forks,
R8234 (CAS). Ipano: apa co.: Boise, 2,880 ft., Clark 38 (DS, F, GH, NY, POM,
RM, UC, US, WS, WTU). canyon co.: New Plymouth, 2,200 ft., Nelson &
Macbride 777 (RM); Big Willow, 3,000 ft., Macbride 158 (GH, NY, RM, US).
OWYHEE Co.: Jump Creek Canyon, R13391 (DS). wAsHINGTON co.: Weiser, 2,200
ft., Jones in 1900 (POM). NEvapa: sToREY co.: Carson City, 5,000 ft., S. Watson
in 1868 (NY). WASHOE co.: Reno Hot Springs to Steamboat Springs, 5,200 ft.,
Train 3557 (NY, UC). OreGon: crook co.: Hay Creek, 730 mi., Leiberg 204
(F, GH, NY, ORE, UC, US, WS). povaias co.: Bluffs of the Umpqua R.
opposite Roseburg, Constance & Rollins 2959 (DS, GH, NMC, NY, RM, RSA,
UC, US, WS, WTU). crutram co.: 6 mi. E. of Arlington, R18390 (UC, WS,
WTU). HARNEY co.: 28 mi. from Burns on road to Lakeview, Howell 28697
(CAS); 12 mi. N. of Frenchglen, Peck 25262 (OSC, UC). Hoop RIvER co.: Hood
River, Suksdorf in 1895 (WS). sackson co,: 2 mi. N. of Central Point, Peck 14968
(US). JOSEPHINE co.: Grants Pass, Henderson in 1886 (DS). Morrow co.: 0.5
mi. E. of Boardman, Cronquist 6239 (NY, OSC, RSA, US, WS, WTU). sHErR-
MAN cO.: Mouth of John Day R., Hitchcock 20435 (WT U). UMATILLA co,: Uma-
tilla, Jones in 1902 (POM). wasco co.: The Dalles, Sheldon 10263 (D5, F, GH,
NY, POM, US, WS); Tygh Valley, Flinn in 1913 (ORE). WHEELER Co.: 2 mi.
SW. of the Painted Hills, ca. 10 mi. NW. of Mitchell, 2,300 ft., Cronquist 7269
(NY, OSC, RSA, US, WS, WTU). WasHINGTON: BENTON co.: Collidge, ZFR
in 1910 (WS). FRANKLIN co.: Pasco, Piper 2965b (WS). IsLAND co.: Whidbey
I., Gardner (POM, UC). KuicxiTat co.: Carley to Roosevelt, Pickett et al. 1433
(WS); near Rockland, Suksdorf 5098-5101 (DS, F, US, WS). san suan co.: Cattle
Point, San Juan I., Peck 12711 (WS, WTU). waLLa wa.ua co.: Wallula, Booth
in 1944 (WS).

As mentioned in the discussions of the preceding species of this
section, some populations of Camissonia contorta, the only hexaploid
species of the group, are very difficult to distinguish from those of
the tetraploid C. strigulosa. Camissonia contorta probably arose, at
least in part, following hybridization between the diploid C. campestris
subsp. campestris and the tetraploid C. strigulosa, but some popula-
tions referred to this species may also have originated following
the functioning of an unreduced gamete in a tetraploid plant. Camis-

300 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

sonia contorta has been found growing sympatrically with.C. kernensis
subsp. kernensis, C. pubens, C. parvula, C. campestris subsp. cam-
pestris, C. campestris subsp. obispoensis, C. sierrae, C. lacustris (popu-
lations in near vicinity), C. strigulosa, and C. integrifolia. Hybrids
have been observed with C. campestris subsp. campestris, C. campestris
subsp. obispoensis, and C. strigulosa, but these were highly sterile,
and there is no evidence of intergradation between C. contorta and
any other species. Whether the northern and southern populations
of this hexaploid differ from one another enough to merit formal
taxonomic recognition (particularly in view of the large gaps in the
range) remains to be seen; I cannot identify any consistent differences
between them at present.

Hexaploid populations were referred by Munz (Bot. Gaz. 85: 251-
257. 1928) to Oenothera contorta var. typica (virtually all of this taxon),
var. epilobioides, and var. strigulosa (a few northern populations only;
mostly=C. strigulosa). In his more recent treatment of the group
(N. Amer. Fl. Il. 5: 156-157. 1965) he presented a very confusing
treatment, recording Oenothera contorta var. contorta as diploid, which,
so far as I am aware, it never is; placing some hexaploid populations
with his Oenothera dentata, which name I now restrict to South Amer-
ican plants; and recognizing as a species Oenothera cruciata, said to
have n=7, and consisting of a mixture of late-season plants of Camis-
sonia campestris subsp. campestris (n=7), the Clear: Lake, Lake
County, California, populations of C. lacustris (n=14), and some
relatively large-flowered plants of C. contorta (n=21).

Section VII. Eremothera

Camissonia sect. Eremothera Raven, Brittonia 16: 285. 1964.
Oenothera subg. Sphaerostigma sensu Munz, Bot. Gaz. 85: 234. 1928; pro
parte.
Oenothera sect. Eremothera (Raven) Munz, N. Am. FI. II. 5: 148. 1965.
Annuals, the plants caulescent, with no well-defined basal rosette,
but the leaves sometimes clustered near the base; plants usually with
conspicuous exfoliating white rhytidome, often reddish; plants flower-
ing from the base or not. Leaves subsessile or petiolate, sometimes
purple-dotted. Inflorescence erect or nodding at anthesis and becoming
erect in fruit. Flowers opening near sunset. Ovary lacking a sterile
projection. Petals white, only rarely reddish initially but always fading
reddish after fertilization, without any dots or markings. Anthers
yellow; stigma and inside of hypanthium greenish; filaments and
style white. Capsule sessile regularly but sometimes tardily loculicidal,
straight or much contorted, often somewhat torulose, the seeds in
one row in each locule, the friable central column much distorted by
the seeds at maturity. Seeds either of two kinds, those of the body of

RAVEN—GENUS CAMISSONIA 351

the capsule sharply angular, truncate-ellipsoid, dark brown, coarsely
papillose, those of the beak narrowly cbovoid or narrowly oblanceolate,
smooth or finely lacunose, or all of the second type (figs. 71, 72).

TYPE SPECIES: Camissonia refracta (S. Wats.) Raven.

DistriBution: Mainly in the interior deserts and bordering areas,
from east-central Washington, central and southern Idaho, and west-
ern Wyoming south through western Colorado to trans-Pecos Texas,
northern Sonora, northeastern and central Baja California, and the
deserts of California, whence north in the inner Coast Ranges to
Alameda and San Joaquin Counties; loose, sandy, or clay soil, on
brushy or open slopes and flats.

The clearly related species of Camissonia sect. Eremothera can easily
be divided into four distinctive groups: The first consists of the self-
incompatible C. refracta and its more widely distributed autogamous
derivative, C. chamaenerioides; second, the polymorphic, self-incom-
patible C. boothiz, here divided into six subspecies, and its two auto-
gamous, smaller flowered derivatives, C. pygmaea and C. gouldii;
third, C. nevadensis, a localized endemic of clay soil in west-central
Nevada which is very likely an outcrosser and may or may not be
self-incompatible; and fourth, the highly autogamous, widespread
C. minor, which does not appear to be closely related to any of the
other groups. In contrast to sect. Holostigma and sect. Camissonia,
which contain majorities of polyploid species, sect. Eremothera re-
sembles sect. Chylismia and sect. Eulobus in being almost exclusively
diploid, the one exception being one of the two strains of the auto-
gamous C.. pygmaea that was examined cytologically; this plant was
tetraploid (n=14). I cannot make any useful suggestions as to
whether C. refracta or C. boothii is more primitive within this group,
but it is clear that the autogamous species have all been derived from
self-incompatible ancestors. Camissonia nevadensis is comparable with
Boisduvalia cleistogama Curran, an annual species of another tribe of
the family (Epilobieae), in inhabiting heavy clay soil and having
decumbent branches with tardily dehiscent capsules which shed their
seed only following rains and many months after the flowers were
open and the plants were green. As pointed out earlier in this paper,
Camissonia graciliflora and C. palmeri also share this unusual habit;
I do not know of any comparable species of Onagraceae, although a
number of genera in other families that occur in the same places have
produced one or more species that are somewhat comparable.

In general, the outcrossing species of Kremothera are visited and
pollinated by small moths in the evening after they open and some-
times also by bees early the following morning. At a few, widely
scattered localities, Camissonia boothii subsp. decorticans opens its
flowers about an hour before sunset and is visited and pollinated by

295-655 O—68——13

352 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

the large, oligolectic bee Andrena (Onagrandrena) vespertina Linsley

& MacSwain. Most populations of this subspecies, and most (if not all)

other populations of the section open their flowers at or shortly after

sunset, and there is generally not enough time for visitation by bees
before darkness.

In his revision of this group, Munz (Bot. Gaz. 85: 233-270. 1928)
recognized C. refracta, C. chamaenerioides, C. nevadensis, and C. minor
with the same limits accepted in the present paper. No problems have
arisen subsequently concerning the delimitation of these distinctive
species. Camissonia pygmaea was considered a variety of C. boothit,
but I raised it to specific rank in 1964 (Raven, Brittonia 16: 285)
because of its nonoverlapping pattern of variation. This entity is
autogamous, in contrast to the self-incompatible and much larger
flowered C. boothii; the separation of these two taxa at the specific
rank was subsequently accepted by Munz (N. Amer. Fl. I. 5: 154.
1965). Camissonia gouldvi is described in the present revision from
material that was not available to Munz at the time of his 1928 paper;
like C. pygmaea, it is evidently an autogamous derivative of C. boothti.

The populations here grouped as C. boothit were separated by
Munz (Bot. Gaz. 85: 233-270. 1928) into three species: Oenothera
decorticans, with four intergrading varieties; O. boothii; and O. alys-
soides. Three of the four varieties of O. decorticans are here regarded
as subspecies of Camissonia boothii; the fourth, var. rutila, is regarded
as a series of unrelated populations with relatively small flowers
in which the petals are red from the time the flowers first open. In-
termediate populations are found joining the three species recog-
nized by Munz in 1928, and this is the reason that I considered them
best treated as subspecies in 1964. In his 1965 monograph, Munz
followed this treatment, describing an additional subspecies, subsp.
intermedia, which occurs in central Nevada and adjacent eastern-
most California and links subsp. boothit with subsp. alyssoides. Dr.
Munz and I thought earlier that an additional subspecies, which he
described in 1965 as subsp. inyoensis, should be recognized; but I
have since come to believe that this entity is best treated as part
of subsp. desertorum as delineated here. At any rate, we both agree
that this complex series of populations is best regarded as a single,
polytypic species.

52. Camissonia refracta (5S. Wats.) Raven, Brittonia 16: 285. 1964. FiaurE 67
Sphaerostigma refractum (S. Wats.) Small, Bull. Torrey Cl. 23: 192. 1896.
Oenothera deserti M. EK. Jones, Contr. West. Bot: 12: 15. 1908. Type: Needles,

San Bernardino Co., Calif., 5 May 1884, M. EF. Jones 3828 (POM: isotypes,
DS, GH, US).
Sphaerostigma deserti (Jones) Heller, Muhlenbergia 9: 68. 1913.

Erect, 6-45 cm. tall, often well branched at the base and above,
not flowering near the base, sparsely strigulose or with an admixture

RAVEN—GENUS CAMISSONIA 353

66. 60

(MW)

Ficures 66-67.—Species of Camissonia sect. Eremothera, X 34: 66, C. chamaenerioides
(Gould 1473, DS); 67, C. refracta (Hitchcock &@ Muhlick 22246, DS).

of glandular trichomes in the inflorescence or all over. Leaves very
narrowly elliptic to very narrowly lanceolate, the basal ones largest,
to 6 cm. long and 0.8 cm. wide, often withered by the time of flow-
ering; leaves sparsely and weakly denticulate; apex acuminate, the
base attenuate; basal leaves with a petiole up to 2 ecm. long, the

354 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

upper ones subsessile. Inflorescence nodding before anthesis. Hy-
panthium 4-6 mm. long, 1.6-2 mm. across at the summit, villous
in lower half within. Sepals 4-6 mm. long, 1.2-2 mm. wide. Petals
3.5-7 mm. long, 2-6 mm. wide. Filaments of the episepalous stamens
2-4.5 mm. long, those of the epipetalous ones slightly shorter; an-
thers 1.5-2.5 mm. long. Style 9-13 mm. long, the lower portions
villous; stigma 1-1.5 mm. in diameter, held well above the anthers
at anthesis. Capsule 2-5 cm. long, 0.7-1 mm. thick, straight to con-
torted, erect, spreading, or reflexed, terete. Seeds all of one type,
0.9-1.5 mm. long, 0.45-0.5 mm. thick, gray, finely lacunose, the lacu-
nae in lines. Gametic chromosome number, n=7. Self-incompatible.

Lectotype: Gravelly hills near the Colorado River, Arizona or
California, 17 February 1854, J. Bigelow (GH); cf. Munz, Bot. Gaz.
85: 237. 1928.

DISTRIBUTION (Figure 68) : Sandy desert slopes and flats from south-
ernmost Esmeralda, southern Nye, and Clark Counties, Nevada, and
Washington County, southwestern Utah, south throughout the
Mojave and Colorado Deserts of Inyo, San Bernardino, Imperial,
central and eastern Riverside, and eastern Kern and San Diego
Counties, California; Mohave, Yuma, and western Pima Counties,
Arizona. A single collection has been seen from Hidalgo County, New
Mexico (east of Lordsburg, Jones in 1930, POM), but this is from
some 250 miles out of the usual range. I have not mapped it and feel
that the locality should be reconfirmed by further collections. From
about 100 ft. below sea level to 4,500 ft. alt.

Vouchers for chromosome number (15 individuals, 12 populations),
n=7:

U.S.: ARIZONA: YUMA Co.: Ca. 20 mi. E. of Yuma, H. & M. Lewis 1640 (LA; 2
pairs + ring of 10); 20 mi. N. of Yuma, R14837 (n=7); 4.3 mi. E. of Ehrenburg,
R11597 (2 plants, both with 5 pairs + ring of 4). CALIFORNIA: IMPERIAL CO.:
23 mi. N. of Ogilby, Alein 119 (7 pairs). INyo co.: Sheppard Canyon, Panamint
Valley, T. & L. Mosquin 4312 (4 pairs + ring of 6); Harrisburg Flats, Panamint
Mts., Gregory 399 (RSA; 5 pairs + ring of 4). RIVERSIDE co.: Box Canyon, Lewis
1623 (LA; 5 pairs + ring of 4), 1633 (LA; 3 pairs + 2 rings of 4); Corn Springs,
Chuckwalla Mts., R11577 (2 plants, both with 5 pairs + ring of 4); Fargo Canyon,
R114389 (2 plants, both with 7 pairs). SAN BERNARDINO Co.: Near Whipple Mts.,
Klein 1394 (7 pairs). NEVADA: CLARK Co.: Ranger Mts., R18916 (n=7).

Camissonia refracta is distinctive and has rarely been confused
with any other entity. It often grows sympatrically with C. chama-
enerioides, C. boothvi subsp. condensata, and C. boothia subsp. deser-
torum, but no hybrids have been observed in nature. The high degree
of chromosomal heterozygosity in this species is striking and could
well be investigated further; of the 13 individuals in which chromo-
some association has been observed, 4 had 7 pairs; 6 had 5 pairs and
a ring of 4; 1 had 3 pairs and 2 rings of 4; 1 had 4 pairs and a ring of 6;

RAVEN—GENUS CAMISSONIA

359

2
o oY?
o
o
* So
© o
AK K
* Se PS
"he °3°/°
e e
ONY
e
.¢ .
= ee
eo § ee
mrt e*®
ee? e
. % - e
\ e* ° e
ee
e

Ficure 68.—Western United States, showing the ranges of species of Camissonia sect.
Eremothera: @=C. refracta; X=C. nevadensis; O=C. pygmaea, with @=diploid
chromosome count (n=7) and &=tetraploid count (n=14); A=C. gouldii.

306 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

and 1 had 2 pairs and a ring of 10. There is no evidence that any of
these configurations is associated with balanced lethals as in certain
species of Oenothera, Gayophytum, Calylophus, and Gaura; but it
would seem on the basis of our present limited amount of evidence
that well over half of the individuals of this species are heterozygous
for at least one reciprocal translocation, a situation unknown else-
where in Camissonia. Six individuals from the Nevada population
cited above were found to be self-incompatible by pollen-tube growth.

53. Camissonia chamaenerioides (A. Gray) Raven, Brittonia 16: 285. 1964.
FIGURE 66
Sphaerostigma chamaenertoides (A. Gray) Small, Bull. Torrey Cl. 23: 189. 1896.
Sphaerostigma erythrum A. Davids., Bull. 8. Calif. Acad. Sei. 1: 118, pl. 9.
1902. Type: Dry hill slopes, south of Clifton, Greenlee Co., Ariz., 10 April

1899, A. Davidson 244 (LAM; isotypes, DS, UC).

Oenothera erythra (A. Davids.) Macbride, Contr. Gray Herb. 65: 41. 1922.

Erect, 8-50 cm. tall, usually branched near the base, not flowering
near the base, glandular-pubescent with an admixture of strigulose
pubescence in and near the inflorescence. Leaves very narrowly
elliptic, more rarely narrowly elliptic, the basal ones largest, to
8 cm. long and 2.5 em. wide; leaves entire to very sparsely dentic-
ulate; apex acute or acuminate, the base attenuate; petioles of basal
leaves 1-3.5 cm. long, the upper leaves subsessile. Inflorescence
nodding before anthesis. Hypanthium 1.6—-2.3 mm. long, 0.9-1.4 mm.
across at the summit, pubescent in lower half within. Sepals 1.5-2.5
mm. long, 0.7-1.38 mm. wide. Petals 1.8-3 mm. long, 1-1.8 mm. wide.
Filaments of the episepalous stamens 0.7-1.5 mm. long, those of the
epipetalous ones very slightly shorter; anthers 0.5-1.1 mm. long.
Style 2.3-4.5 mm. long, the lower portions pubescent; stigma 0.7—1
mm. in diameter, surrounded by the anthers at anthesis. Capsule
3.5-5.5 cm. long, 0.8-0.9 mm. thick, spreading, terete. Seeds similar
to those of C. refracta, 0.9-1 mm. long, ca. 0.3 mm. thick. Gametic
chromosome number, n =7. Autogamous.

Type: El Paso, El Paso County, Texas, 1852, C. Wright 1377 (GH;
isotypes, BM, K, PH, US).

Disrripution (Figure 50): Sandy desert slopes and flats from
northern Inyo County, California, Nye and Lincoln Counties, Nevada,
and Washington County, Utah (also known from Dugway, Tooele
County, Jones 1891, POM), south to the vicinity of Bahia de los
Angeles in northeastern Baja California, in Sonora to the vicinity of
Luja midway between Hermosillo and Guaymas, throughout Arizona
except for the northeastern corner, rare in New Mexico (Dona Ana
and Luna Counties), and in trans-Pecos Texas east to Brewster
County; also known from Isla Angel de la Guarda in the Gulf of
Cahfornia. Not known from, but to be expected in, Chihuahua. From
about 150 ft. below sea level to 4,600 ft. alt.

RAVEN—GENUS CAMISSONIA 357

Vouchers for chromosome number (10 individuals, 5 populations),
n=7:

U.S.: ARIZONA: PIMA cO.: 3 mi. N. of headquarters, Organ Pipe National
Monument, H. & M. Lewis 1639 (3 plants; LA). yuma co.: 4.3 mi. E. of Ehren-
burg, R11596 (DS); 1 mi. 8. of Ehrenburg, R11593 (DS). CALIFORNIA: RIVERSIDE
co.: Corn Springs, Chuckawalla Mts., Mosquin 3209 (4 plants; DAO).

BAJA CALIFORNIA: Bahia de los Angeles, Moran 12422.

Camissonia chamaenerioides is distinctive and has rarely been con-
fused with any other species. It is similar to C. refracta and probably
derived from populations relatively similar to that species, but differs
not only in flower size but also in leaf shape, pubescence, and seed
size. Camissonia chamaenerioides is the only species of the genus that
occurs in Texas. This species occurs sympatrically with C. refracta, C.
boothii subsp. condensata, and C. boothii subsp. desertorum; no natural
hybrids have been observed.

54. Camissonia boothii (Dougl.) Raven, Brittonia 16: 285. 1964.

Erect, 12-65 cm. tall, often well branched at the base and above:
usually not flowering near the base, subglabrous, strigulose, villous, or
elandular-pubescent. Leaves very narrowly elliptic to narrowly ovate,
the basal ones largest, 2-10 cm. long, 0.5-3.8 cm. wide, often withered
by time of flowering, sparsely denticulate, the apex acuminate or
acute, the base narrowly cuneate to attenuate; basal leaves with a
petiole 0.4-6 cm. long, the upper ones subsessile. Inflorescence nodding
before anthesis. Hypanthium 4-7 (-8) mm. long, 1.7-2.5 mm. across at
the summit, villous in lower half within. Sepals (2.7—) 4-8 mm. long,
1.4-2.3 mm. wide. Petals (3-) 4-7.5 mm. long, (1.75-) 2.6-7 mm. wide.
Filaments of episepalous stamens (1.5—-) 2-5.8 mm. long, those of the
epipetalous ones slightly shorter; anthers (1—) 1.8-2.3 mm. long. Style
(6.5-) 8.2-13.5 (-15) mm. long, villous near the base; stigma 1.2-2 mm.
across, depressed-globose, held well above the anthers at anthesis.
Capsule slightly curved outward to contorted, 0.8-3.5 cm. long, 1-3.8
mm. thick at the base, terete, held on the dried plant and shedding seeds
tardily. Seeds all of one type or of two types, the relatively smooth
kind (which may be the only kind) 1.4-2.1 mm. long, 0.5-0.7 mm. thick,
more or less triangular in transection, light brown, those of the body
0.5-0.9 mm. thick, dark brown, coarsely papillose. Gametic chromo-
some number, n =7. Self-incompatible.

Distripution (Figure 69): Deserty, often brushy or open slopes
and flats, from Klickitat, Adams, and Whitman Counties, eastern
Washington, and southwestern Idaho, south to eastern California
(whence north in the inner Coast Ranges to Alameda and San Joaquin
counties), northeastern Baja California, northwestern Sonora, and
through western Utah to western Arizona, south to Pima and Pinal
Counties. From about 200 ft. below sea level to 6,600 ft. alt.

358 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

As explained in the introductory remarks for the section, Camis-
sonia boothi is here treated as a polytypic species with six subspecies -
that replace one another geographically. The seeds are apparently
always monomorphic in subsp. alyssoides but very rarely so in popu-
lations of the other subspecies.

Of the 21 individuals of this species for which meiotic chromosome
association was observed, 2 had a ring of 4 chromosomes and 5 pairs,
1 had a ring of 6 and 4 pairs, and 18 had 7 pairs. This suggests the
presence of translocation heterozygosity of about the same order of
magnitude (about 20 percent) as found in sect. Chylismia, but much
less than characteristic of C. refracta.
54a. Camissonia boothii subsp. decorticans (Hook. & Arn.) Raven, Brittonia 16:

285. 1964.

Gaura decorticans Hook. & Arn., Bot. Beechey Voy. 3438. 1838.

Oenothera gauraeflora Torr. & Gray, Fl. N. Amer. 1: 510. 1840.

Sphaerostigma gauraeflorum (Torr. & Gray) Walp., Rep. 2: 78. 1843.

Oenothera decorticans (Hook. & Arn.) Greene, Fl. Francise. 217. 1891.

Oenothera rutila A. Davids., Erythea 2: 62. 1894. Type: Big Rock Creek,
San Gabriel Mts., Los Angeles Co., Calif., 8 July 1894, A. Davidson
(LAM).

Sphaerostigma decorticans (Hook. & Arn.) Small, Bull. Torrey Cl. 23: 191.
1896.

Sphaerostigma rutilum (A. Davids.) Parish, Erythea 6: 89. 1898.

Oenothera alyssoides var. decorticans (Hook. & Arn.) Jeps., Man. Fl. Pl.
Calif. 686. 1925.

Oenothera decorticans var. typica Munz, Bot. Gaz. 85: 245. 1928.

Oenothera decorticans var. rutila (A. Davids.) Munz, Bot. Gaz. 85: 245. 1928.

Oenothera boothii subsp. decorticans (Hook. & Arn.) Munz, N. Amer. FI. IT.
5: 152, 1965

Oenothera boothit subsp. rutila (A. Davids.) Munz, Fl. N. Amer. II. 5: 153.
1965.

Stems 12-65 mm. long, hollow, with conspicuous, exfoliating white
or somewhat pinkish or brownish rhytidome. Plants subglabrous,
with strigulose or glandular trichomes in the inflorescence. Capsule
nearly straight, curved outward, swollen at base, 1.7-2.3 mm. thick
near the base, rapidly tapering upward. Seeds dimorphic, 1.4-2.1
mm. long, 0.5-0.7 mm. (smooth type) or 0.8 mm. (papillose type)
thick. Gametic chromosome number, n=7. Self-incompatible.

Type: California, 1833, D. Douglas (K; isotypes, BM, GH, NY).

Distrisution (Figure 69): Open, usually steep slopes, often on
shale or other loose rocky substrates, endemic to California: inner
south Coast Ranges from Alameda and southwestern San Joaquin
Counties south to Santa Barbara and Ventura Counties and the San
Gabriel Mountains and easternmost Santa Monica Mountains of
Los Angeles County, whence north through the Tehachapi Mountains
to the western slopes of the southern Sierra Nevada in Kern and
southernmost Tulare Counties; from near sea level to 6,000 ft. alt.

RAVEN—GENUS CAMISSONIA 359

—

Figure 69.—Western United States and a portion of northwestern Mexico, showing the
range of Camissonia boothii: A=C. boothit subsp. decorticans; X =subsp. desertorum:
@=subsp. condensata; A=subsp. intermedia; C=subsp. alyssoides; = subsp.
boothii.

360 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Vouchers for chromosome number (19 populations, 10 individuals),
n=7 (7 pairs in plants marked with an asterisk):

U.S.: CALIFORNIA: KERN Co.: Grocer Grade 5 mi. W. of Maricopa, 2,000 ft.;
Breedlove 2395*; near Havilah, Lewis 1106 (2 plants, 1*; LA, RSA, UC). mon-
TEREY CO.: 4.9 mi. E. of King City, Breedlove 2175*. sAN LUIS OBISPO Co.: Carrizo
Plain at 8S. end of Soda Lake, Bates 2407; 8.6 mi. W. of Simmler, Breedlove 2062;
N. road to Adelaida near junction to Nacimiento Dam, R18326; Paso Robles,
Lewis & Thompson 1101* (LA). SANTA BARBARA Co.: 4.3 mi. W. of Pendola Flats
Guard Station, upper Santa Ynez R., 1,600 ft., Breedlove 2241. STANISLAUS Co.:
Arroyo del Puerto, Breedlove 5579.

Camissonia boothii subsp. decorticans is the only member of sect.
Eremothera to occur off the deserts, but it still occurs in related,
highly xeric communities. Twenty plants of R18326, cited above,
were tested for pollen-tube growth following self- and cross-pollina-
tion and found to be self-incompatible, as have four from east San
Luis Obispo County, California, Ayhos 65-173. This subspecies has
not been found growing sympatrically with any other member of the
section, but it intergrades with subsp. desertorum at the margins of
its range. It is usually found away from the coast, but has been found
on the bluffs at Taylor Ranch, 1.5 miles east of the mouth of the
Ventura River, Pollard in 1963 (CAS, SBBG).

Occasional populations of this subspecies at relatively high eleva-
tions in the southern part of its range have somewhat smaller flowers
and reddish petals; they have been distinguished as Oenothera rutila
or, more recently, Oenothera boothia subsp: rutila. In my opinion,
there is no evidence that the various populations with this combina-
tion of characteristics had a common origin, and, even if they did,
they do not appear to constitute a major geographical race comparable
with the other subspecies recognized here.
54b. Camissonia boothii subsp. desertorum (Munz) Raven, Brittonia 16: 285.

1964.

Oenothera decorticans var. desertorum Munz, Bot. Gaz. 85: 246. 1928.

Oenothera boothii subsp. desertorum (Munz) Munz, Fl. N. Amer. II. 5: 153.
1965.

Oenothera boothii subsp. inyoensis Munz, Fl. N. Amer. IT. 5: 153. 1965. Type:
Willow Creek, Saline Valley, 800 m, Inyo Co., Calif., 25 April 1942, A. M.
Alexander & L. Kellogg 2702 (RSA, 41213).

Similar to subsp. decorticans but shorter, usually less than 35 cm.
tall. Capsule flexuous-contorted, the beak often directed downward,
1-1.6 mm. thick near the base. Gametic chromosome number, n=7.

Type: 10 miles southwest of Garlic Springs, San Bernardino
County, California, 8 April 1924, P. A. Munz & D. D. Keck 7881
(POM 48926).

Disrrisution (Figure 69): Desert slopes and washes, endemic
to California: from vicinity of Benton Station, Mono County, south in
the deserts and east slope of the Sierra Nevada to eastern Kern,

RAVEN—GENUS CAMISSONIA 361

northern Los Angeles, and western San Bernardino Counties; also in
the upper Kern River drainage, west to the vicinity of Kernville,
and in southeasternmost Tulare County. From 1,500-6,500 ft. alt.

Vouchers for chromosome number (6 individuals, 5 populations),
n=7:

U.S.: CALIFORNIA: INYO Co.: 3 mi. N. of Big Pine, R14258; road to Westgard
Pass, Lewis 1085 (LA, RSA, UC; 7 pairs); Marble Canyon, Inyo Mts., R17551
(7 pairs). KERN co.: Road from Randsburg to Red Rock Canyon, Wedberg in 1957
(LA; 5 pairs+ring of 4). Mono co.: Sherwin Grade, 5,000 ft., Lewzs 1673 (LA).

As here conceived, Camissonia boothit subsp. desertorum occupies
a position intermediate between subsp. decorticans and subsp. con-
densata, both geographically and morphologically. Populations of
subsp. desertorum from the northern portion of its range are in general
more pubescent than those from farther south and were referred by
Munz (Bot. Gaz. 85: 241-247. 1929; N. Amer. Fl. II. 5: 151-154.
1965) to subsp. alyssoides. They may represent intermediates with
that taxon, but do not agree with it in habit, being much more lax,
and have much more appressed pubescence. In my view, the only
populations of subsp. alyssoides in California are those in eastern
Lassen County, and the nearest populations to those of subsp. de-
sertorum are in the Sheep Range of northern Clark County, Nevada,
and in Churchill County, Nevada. I have also included in subsp.
desertorum those populations from the Panamint Mountains Inyo,
County, California, listed by Munz (Bot. Gaz. 85: 245. 1929) as
approaching Oenothera decorticans var. rutila; these were apparently
also included in subsp. desertorum by Munz in his more recent work
on the group (N. Amer. Fl. II. 5: 153-154. 1965).

Populations from the eastern slope of the Inyo Mountains and
elsewhere in Inyo County, California, where they grow on limestone,
have relatively small flowers and lax inflorescences, which make them
appear quite distinct in appearance. They were named Oenothera
boothit subsp. inyoensis by Munz in 1965, and at the time, I too thought
that they deserved formal taxonomic recognition. Intergradation with
other populations more typical of subsp. desertorwm is so broad and
complete that it is often impossible to assign particular plants to one
or the other with confidence. Therefore I now consider that they are
best assigned to subsp. desertorum, despite the fact that their inclusion
makes subsp. desertorum quite heterogeneous.

A few collections made within the range of subsp. condensata on
the western Colorado Desert of California appear to be this subspecies ;
for example, Parish in 1882 (JEPS) from Whitewater, Riverside
County. These may be extreme variants of subsp. condensata or
may represent relict populations of subsp. desertorum in an area now
largely occupied by subsp. condensata, which may be of relatively
recent derivation.

362 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

54c. Camissonia boothii subsp. condensata (Munz) Raven, Brittonia 16:
285, 1965.
Oenothera decorticans var. condensata Munz, Bot. Gaz, 85: 247. 1928.
Oenothera boothii subsp. condensata (Munz) Munz, N. Amer. Fl. II. 5:
154. 1965.

Similar to subsp. desertorum, but shorter, 5-20 (—30) em. tall, the
stems very thick, the inflorescence more crowded. Capsules 2-3.8 mm.
thick near the base, more or less quadrangular, tapering abruptly,
the midribs of the valves very prominent, yellowish. Gametic chromo-
some number, n=7. Self-incompatible.

TypEe: Dos Palmos Spring, Riverside County, California, 31 Jan-
uary 1926, P. A. Munz 9960 (POM 98708).

DistriBuTION (Figure 69): Desert slopes and washes, in sandy soil,
from the vicinity of Death Valley, Inyo County, California, south
to the region around Barstow, San Bernardino County, and throughout
the Colorado Desert of Riverside, Imperial, and eastern San Diego
Counties, to the region about the head of the Gulf of California in Baja
California and Sonora (south to about 31°30’ N. lat.); eastward to
Esmeralda, southern Nye and Lincoln, and Clark Counties, Nevada,
and Washington County, Utah, and western Mohave, Yuma, Mari-
copa, southwesternmost Pinal, and westernmost Pima Counties,
Arizona; also in Glen Canyon, San Juan County, Utah (Gaines 927,
1011, WS). From about 250 ft. below sea level to 4,000 ft. alt.

Vouchers for chromosome number (8 individuals, 7 populations),
n=7:

U.S.: ARIZONA: MARICOPA cCo.: Near Sentinel, R14826. MoHAVE co.: 48 mi.
NW. of Kingman, Munz et al, 22985 (5 pairs -+ ring of 4). CALIFORNIA: IMPERIAL
co.: Coyote Wells, Breedlove 1847; 1 mi. N. of U.S. Hwy. 80 on road to Ogilby,
Klein 94 (2 plants, 7 pairs). RIVERSIDE co.; Box Canyon, Lewis 1663 (LA; 7
pairs). SAN DIEGO co.: Borrego Valley, Lewis 1624 (LA; 7 pairs), Wedberg in 1957
(LA).

As noted by Munz in the protologue of this taxon (Bot. Gaz.
85: 247. 1928), it is very distinctive in fruit, the plants persisting in a
dried condition for a long time and shedding their seeds tardily.
Camissonia boothti subsp. condensata intergrades broadly with subsp.
desertorum where their ranges approach one another. Twelve plants
from Frenchman Flat, Nye County, Nevada (R18853), were found
to be self-incompatible by pollen-tube growth following cross- and
self-pollination.
54d. Camissonia boothii subsp. alyssoides (Hook. & Arn.) Raven, Brittonia 16:

285. 1964.
Oenothera alyssoides Hook. & Arn., Bot. Beechey Voy. 340. 1838.
Holostigma alyssoides (Hook. & Arn.) Hook., Icon. 4: 339. 1840.
Sphaerostigma alyssoides (Hook. & Arn.) Walp., Rep. 2: 78. 1843.

Oenothera alyssoides var. villosa 8. Wats., Proc. Amer. Acad. 8: 591. 1873.
Type: Near Salt Lake, Utah, Capt. H. Stansbury (GH; isotype, NY).

RAVEN—GENUS CAMISSONIA 363

Sphaerostigma utahense Small, Bull. Torrey Cl. 23: 191. 1896. Lectotype:
Milford, Beaver Co., Utah, June 1880, M. E. Jones (NY).

Sphaerostigma alyssoides var. macrophyllum Small, Bull. Torrey Cl. 23: 192.
1896; nom. subs.

Oenothera gauraeflora var. hitchcockit H. Lév., Monogr. Onoth. 226. 1905.
Type: Simpson’s Park, Utah, 6 July 1859, collector not known (MO).

Sphaerostigma hitchcockii (H. Lév.) A. Nels., Bot. Gaz. 40: 226. 1905.

Oenothera utahensis (Small) Garrett, Spring Fl. Wasatch Reg. 64. 1911.

Sphaerostigma implerum A. Nels., Bot. Gaz. 52: 267. 1911. Type: Falk’s
Store, 2,200 ft., Canyon Co., Idaho, 17 May 1910, J. F. Macbride 27 (RM;
isotypes, GH, UC, US, WS, WTU).

Sphaerostigma macrophyllum Rydb., Bull. Torrey Cl. 40: 66. 1913.

Oenothera alyssoides var. typica Munz, Bot. Gaz. 85: 242. 1928.

Oenothera boothii subsp. alyssoides (Hook. & Arn.) Munz, N. Amer. FI. II.
5: 154. 1965.

Similar to subsp. desertorum, the plants 3-35 cm. tall, sometimes
flowering near the base, with leafier stems and a less prominent basal
rosette, the leaves relatively small. Plants strigulose, often densely so,
especially in the inflorescence, or more rarely villous or with an
admixture of glandular trichomes. Capsule 1-1.4 mm. thick, usually
very contorted and crowded. Seeds monomorphic, smooth. Gametic
chromosome number, n=7. Self-incompatible.

Typre: Pine Creek, Snake River plains, southern Idaho, probably
in July 1830, D. Douglas (K; isotype, GH).

Distripution (Figure 69): Sandy slopes and flats, often with
Artemisia tridentata, from Grant and northern Malheur Counties,
eastern Oregon and Payette, Ada, and Elmore Counties, southwestern
Idaho, south to eastern Lassen County, California, and Churchill,
northernmost Nye, Lincoln, and northern Clark Counties, Nevada,
and throughout western Utah, south to Piute and Washington Coun-
ties. From 2,000 to 5,500 ft. alt.

Vouchers for chromosome number (8 individuals, 8 populations),
n=7 (7 pairs in those collections marked with an asterisk) :

U.S.: CALIFORNIA: LASSEN co.: 19 mi. S. of Ravendale, R13273. Nevapa:
CHURCHILL co.: 28.9 mi. SW. of Lovelock, Kyhos 65-253.* EUREKA Co.: 2.9 mi.
W. of Carlin, R18535*. HUMBOLDT co.: 3.5 mi. W. of Golconda Summit, R18537*.
LINCOLN co.: E. of Panaca, R11255 (CAS, LA, RSA). PERSHING Co.: 20 mi. NE.
of Lovelock, R18540. OREGON: LAKE Co.: 3.4 mi. E. of Headquarters, Hart Mt.
Antelope Refuge, R13347*. UTAH: MILLARD Co.: 62 mi. SW. of Delta, Mathias
3025* (COLO, LA, RSA, UC).

The number of synonyms listed with this taxon attests to its wide
range rather than to any extraordinary degree of variability. Self-
incompatibility was demonstrated by pollen-tube growth in 21 plants
from the locality of R18540, cited above, and three from 28.9 miles
southwest of Lovelock, Churchill County, Nevada, AKyhos 65-253.
Camissonia boothii subsp. alyssoides intergrades broadly with subsp.
intermedia and is very similar to subsp. desertorum. Certain populations

364 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

cited by Munz (Bot. Gaz. 85: 241-244. 1928) are here regarded as
belonging to C. boothii subsp. desertorum, as noted in the treatment of
that subspecies.

In general, the last three subspecies of this species consist of plants
that persist and flower over a much longer season than the first three,
which flower chiefly following the winter rains.
54e. Camissonia boothii subsp. intermedia (Munz) Raven, comb. nov.

Oenothera alyssoides var. villosa sensu Munz, Bot. Gaz. 85: 243. 1928: pro
arte.

Oenothera boothii var. typica sensu Munz, Bot. Gaz. 85: 248. 1928: pro parte.

Oenothera boothti subsp. intermedia Munz, Fl. N. Amer. II. 5: 152. 1965.

Similar to subsp. alyssoides, but densely villous with a dense ad-
mixture of glandular trichomes in the inflorescence, the plants usually
5-20 cm. tall, the leaves mostly less than 2.5 em. long. Seeds dimorphic.
Gametic chromosome number, n=7.

Type: Dry sandy gravel wash, 4 miles south of Cloverdale Ranch
on Reese River to Tonopah road, Nye County, Nevada, 16 September
1939, P. Train 3455 (RSA 56046; isotypes, UC, WTU).

DistripuTion (Figure 69): Sandy washes and flats, often with
Artemisia tridentata, in Nevada from southeastern Churchill and
southern Lander Counties south throughout Nye and Esmeralda
Counties and in southern Mineral County, and in northeastern Inyo
County, California; also in the Kingston Range, northeastern San
Bernardino County, California. From 5,000-7,000 ft. alt.

Vouchers for chromosome number (2 individuals, 2 populations),
n=7 (7 pairs observed in 1 plant of population marked with asterisk) :

U.S.: CALIFORNIA: INYO co.: Ca. 4 mi. S. of Oasis, Fish Lake Valley, Klein

353 (RSA). NEvapDaA: NYE co.: 10 mi. W. of Lockes, Thompson & Mathias 1692*
(ARIZ, LA, RSA, UC, WTU).

REPRESENTATIVE SPECIMENS EXAMINED:

U.S.: CaLIForNrA: Inyo co.: Deep Springs Valley, Ferris 1357 (DS): 2 mi.
NE. of Willow Spring, Last Chance Mts., 6,000 ft., Roos 6396 (DS, RM, RSA,
UC). SAN BERNARDINO co.: Kingston Mts., 0.1 mi. E. of summit of grade between
Horse (Thief) and Beck Springs, ca. 5,000 ft., Wolf 10515 (RSA). Nevapa:
CHURCHILL co.: 2 mi. KE. of Frenchman, Mills & Beach C-20 (UC): vicinity of
East Gate, 5,100 ft., Allen 349 (DS, NY, POM). esMERALDA co.: 3 mi. W. of
Lida, Silver Peak Range, Maguire & Holmgren 25625 (GH, NY, UC, US, WTU);
Montezuma Mt. W. of Goldfield, 2,000 m, Tidestrom 9764 (GH, US): Icehouse
Canyon, W. slope of Silver Peak Range, 5,300 ft., Alexandria & Kellogg 5682 (DS,
RM, UC, US, WTU); wash leading to summit between Fish Lake Valley and
Basalt, Ferris 6681 (DS, POM). LANDER co.: Austin to Big Creek, 6,000 ft.,
Kennedy 4507 (CAS, DS, GH, PH, RM); Austin, 6,000 ft., Hitchcock & Martin
5626 (DS, NY, POM, UC, WS, WTU); 18.5 mi. E. of Austin, Goodner & Henning
836 (DAO, F, POM, UC). nye co.: Warm Springs Valley, Maguire & Holmgren
25458 (GH, NY, UC, WTU); Smoky Valley, Maguire & Holmgren 25365 (ARIZ,
DS, GH, NMC, NY, OSC, UC, US, WS, WTU); 2 mi. NW. of Penelas Mill, 15
mi. NW. of Ione, Beach 865 (ARIZ, DS, IDS, NMC, NY, POM, RM, UC, US, WS,

RAVEN—GENUS CAMISSONIA 365

WTU); pass over Pancake Mts., 84 mi. SW. of Ely, 6,000 ft., Rollins &
Chambers 2497 (DS, GH, RM, UC, US); foothills of Pahute Mesa, N. Forty Mile
Drainage, 6,000 ft., Beatley 1694 (DS). MINERAL Co.: 10 mi. N. of Basalt, T7zde-
strom 10040 (GH); Candelaria, 6,000 ft., Shockley 324 (DS, UC).

This entity bridges the morphological and geographical gap between
subsp. boothii and subsp. alyssoides in Nevada completely. Neverthe-
less, it consists of a large number of morphologically relatively uniform
populations that occupy a sizable range in central Nevada.

54f. Camissonia boothii subsp. boothii

Oenothera boothii Dougl., in Hook., Fl. Bor. Amer. 1: 213, 1834.

Sphaerostigma boothii (Dougl.) Walp., Rep. 2: 77. 1843.

Oenothera gauraeflora race boothii (Dougl.) H. Lév., Monogr. Onoth. 226.
1905.

Sphaerostigma senex A. Nels., Proc. Biol. Soc. Wash. 18: 173.29 June 1905.
Type: Pyramid Lake, Washoe Co., Nev., 9 June 1903, G. H. True 750
(RM).

Sphacrostigma lemmoni A. Nels., Bot. Gaz. 40: 61. 1905. Type: “East flank of
the Sierra Nevada,” presumably in Nevada, 1875, J. G. Lemmon 103 (MO;
probable isotype, US).

Oenothera boothii var. typica Munz, Bot. Gaz. 85: 248. 1928.

Oenothera boothii subsp. boothii Munz, N. Amer. Fl. IT. 5: 151. 1965.

Similar to subsp. intermedia, but often more robust, and the leaves
broader, lanceolate or narrowly ovate, the cauline ones especially
proportionately broader, 0.8-6 cm. long, 0.6-2.2 mm. broad, more
coarsely serrulate, the plants mostly 15-60 cm. tall. Gametic chromo-
some number, n=7. Self-incompatible.

Type: On high sandy and gravelly hills near the junction of the
Snake and Clearwater Rivers, vicinity of Lewiston, Idaho (either in
Washington or in Idaho), approximately 24 July 1826, D. Douglas
(K; isotype, BM).

DistriputTion (Figure 69): Sandy flats or steep, loose slopes, in
Adams, Whitman, and Walla Walla Counties, southeastern Wash-
ington, west along the Columbia River to Klickitat County and
Hamilton Island, Skamania County, Washington, Washington,
Canyon, Owyhee, and Jerome Counties, southwestern Idaho; south
through eastern Oregon (Grant, Harney, and Malheur Counties) ;
and again in west-central Nevada (southern Washoe, western Church-
ill, easternmost Lyon, and Mineral Counties) and adjacent Mono
County, California, in the vicinity of Mono Lake and Benton. Indis-
tinguishable populations occur in the vicinity of Toroweap Valley,
Mohave County, Arizona (e.g., Cottam 13936, CAS, UT). From
2,000-7,500 ft. alt.

Vouchers for chromosome number (3 individuals, 3 populations),
n=7:

U.S.: NEVARA!: MINERAL co.: W. side of Walker L., R11246 (7 pairs). WASHOE
co.: Pyramid L., 18.8 mi. N. Sutcliffe, R13197 (7 pairs); ca. 15 mi. N. of Nixon,
ca. 3,500 ft., Ornduff 4203 (LA; 4 pairs + ring of 6).

366 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Camissonia booth subsp. boothii occurs in three disjunct areas,
intergrading completely with subsp. intermedia and subsp. alyssoides
in the Nevada portion of its range. A few collections from two areas
on the Mojave Desert of California, within the range of subsp.
desertorum, resemble subsp. boothii closely in their glandular pubes-
cence, habit, and summer blooming:

U.S.: Cauirornia: Inyo co.: 10 mi. N. of Little Lake, 1931, Hoffmann 576
(POM). SAN BERNARDINO co.: Brown’s Crossing of the Mohave R., 1882,
S. B. & W. F. Parish 1504 (DS, F, GH); Hesperia, 1901, Abrams 2166; near
Victorville, 1916, Peirson 792 (RSA).

Plants of this sort have not been collected in these areas for nearly
40 years. The relationship of these populations to the other subspecies
should be investigated when additional material becomes available.
It is possible, but seems unlikely, that the plants were introduced
in these areas. A similar specimen has been seen from Fort Mojave,
Mohave County, Arizona, 1860-1861, Cooper (GH).

Self-incompatibility was demonstrated in 40 plants from the lo-
cality of R11246, cited above, by examining pollen-tube growth in
selfed and outcrossed stigmas.

55. Camissonia pygmaea (Dougl.) Raven, Brittonia 16: 285. 1964.
Figures 71-73
Oenothera pygmaea Dougl., in Hook., Fl. Bor. Amer. 1: 213. 1834.
Oenothera boothii var. pygmaea (Dougl.) Torr. & Gray, Fl. N. Amer. 1:
510. 1840.
Sphaerostigma boothit var. pygmaeum (Dougl.) Walp., Rep. 2: 78. 1843.

Very similar to C. boothii subsp. boothii, the plants 1.5-35 cm.
tall. Flowers smaller, the hypanthium 1.7-2.2 (-4) mm. long, 0.8-1
(-1.5) mm. across at the summit; sepals 1.8-2 mm. long, 0.9-1 mm.
wide; petals 1.5-2.5 mm. long, 0.8-1.3 (-2) mm. wide; filaments of
episepalous stamens 1-2.2 mm. long, those of the epipetalous ones
slightly shorter, the anthers 0.45-0.5 (0.7) mm. long; style 3.2-4
mm. long, the stigma 0.5-0.8 mm. thick, surrounded by the anthers
at anthesis. Gametic chromosome numbers, n=7, 14. Autogamous.

Type: On the barren sands of the interior near the Umatilla (““Utal-
la”) River, east of Pendleton, Umatilla County, Oregon, June 1826,
D. Douglas (K; isotype, BM).

Distrripution (Figure 68): Rare and local on steep, loose slopes,
often in scree, sometimes on gravelly flats, at scattered localities
in eastern Washington (Douglas, Grant, and Kittitas Counties),
eastern Oregon, and at one locality in adjacent southern Idaho
(Jerome County). From 1,200-4,000 ft. alt.

Voucher for chromosome number (1 individual), n=7:

U.S.: OREGON: WHEELER Co.: 10 mi. NW. of Mitchell, R18474.

Voucher for chromosome number (1 individual), n=14:

RAVEN—GENUS CAMISSONIA 367

CMW)

2.

Ficures 70-73.—Species of Camissonia sect. Eremothera: 70, C. gouldii, X % (Gould 1423,
holotype, POM). 71-73, C. pygmaea (Henderson 5411, DS): 71-72, Seeds, X 36: 71
dark papillose seed from lower part of capsule; 72, lighter, smoother seed from upper
part of capsule. 73, Flowering branch, X %.

U.S.: WASHINGTON: GRANT co.: Dry Falls, R18489 (12 pairs + ring of 4).

ADDITIONAL SPECIMENS EXAMINED:

U.S.: IDAHO: JEROME co.: Blue Lakes, Snake Plains, 1893, Palmer 71 (CAS,
UC), 523 (CAS, UC; these two collections with relatively large flowers and
doubtfully referred here). OREGON: GILLIAM Co.: Near camp on Pine Creek, 1470
m, Leiberg 199 (F, GH, NY, ORE, UC, US). rant co.: Muddy Station, John
Day Valley, T. Howell in 1885 (F, GH, NY, ORE, PH, US, WS, WTU); John
Day Valley NW. of Dayville, Ripley & Barneby 6628 (CAS, NY); Squaw Creek,
Humphreys Ranch, Henderson 5411 (CAS, DS, GH, ORE); 5 mi. W. of Mt.

295-655 O—68—_14

368 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Vernon, Peck 19880 (CAS). HARNEY co.: Base of Steens Mt., 7. Howell in 1885
(NY). WHEELER co.: Dry gravelly washes between Sutton Mt. and Bridge Creek,
10 mi. NW. of Mitchell, 2,200 ft., Cronquist 7239 (CAS, DAO, DS, GH, NY,
RM, UBC, UC, WS, WTU); 17 mi. E. of Mitchell, Peck 21056 (UC) ; 22 mi. E. of
Mitchell, R18480 (DS). WasHINGTON: poUGLAS co.: Rock Island, Sandberg &
Leiberg 441 (CAS, GH, LE, NMC, ORE, PH, POM, UC, US, WS, WTU). arant
co.: Dry Falls, Thompson 9105 (DS, GH, NY, POM, RSA, UC, US, WTU);
Grand Coulee near Park Lake, Thompson 11657 (WTU); Soap Lake, Eyerdam
642 (UC). kirtiTas co.: Above Blue Lake, near Cle Elum, McCalla 4438 (UBC).

This rarely collected species is obviously an autogamous de-
rivative of populations similar to those of the much larger flowered,
self-incompatible C. boothit subsp. boothit. As mentioned in the list
of cited specimens, the occurrence of C. pygmaea in Idaho is some-
what doubtful and needs to be confirmed. I have been unable to
separate diploid and tetraploid populations of this species morpho-
logically or by any character of the pollen. It is likely that the tetra-
ploid individual examined was of autopolyploid origin, and the
observation of a ring of four chromosomes in it tends to confirm this
view. This plant was the only tetraploid invidudual found in the
section.

56. Camissonia gouldii Raven, sp. nov. FiGcurE 70

A C. pygmaea persimilis, differens pilis omnibus glandulosis; foliis
ellipticis; capsulis 8-12 mm. longis, 1.7-1.8 mm. latis; seminis omnibus
sublaevibus, nullis papillosis. Autogama.

Type: Associated with Phacelia palmeri, on steep slope of volcanic
cone among loose cinders, Diamond Valley, 12 miles north of St.
George, 3,500 ft., Washington County, Utah, 15 October 1941,
Frank W. Gould 1423 (POM; isotypes, ARIZ, CAS, F, GH, NY,
UC, US).

DistriBuTion (Figure 68): Volcanic scree or cinder flats, rare and
local; south-central Washington County, Utah, and east-central
Coconino County, Arizona; 3,500-5,400 ft. alt.

ADDITIONAL SPECIMENS EXAMINED:

U.S.: ARIZONA: COCONINO Co.: Cinder flats E. of San Francisco Peaks, 1,700
m., Leiberg 5808 (US); San Francisco Mt., Knowlton 200 (US). WuPATKI NATL.
MONUMENT (COCONINO Co.): Deadmans Wash below Wupatki Ruin, occasional on
volcanic ash, ca. 4,800 ft., Whiting & Jones 1089/5213 (US); Wupatki Rim,
Jones 337 (ARIZ); E. slope of Doney Mt., 2 mi. W. of Wupatki, deep cinder soil,
5,200 ft., Jones in 1939 (ARIZ); deep cinder near Hall Canyon, 5,400 ft., Whiting
1089/5274 (ARIZ, POM).

Camissonia gouldii is obviously autogamous and, like C. pygmaea,
probably a derivative of populations similar to those of the self-
incompatible C. boothii subsp. boothit. The monomorphic seeds and
entirely glandular pubescence of C. gouldii (fig. 70) set it off from C.
pygmaea, the nearest stations of which are some 500 miles to the

RAVEN—GENUS CAMISSONIA 369

northwest. In addition, its reddish color and relatively broad leaves

give C. gouldii a very distinctive aspect. The chromosome number of

C. gouldia is unknown.

It is a pleasure to name this distinctive local endemic in honor of
Prof. Frank W. Gould of Texas A. & M. University, student of the
grasses, who has collected widely and critically in the western United
States, in Mexico, and elsewhere in Latin America. It adds another
to the long list of edaphically sharply restricted endemics of its region,
one of which is Camissonia parry?.

57. Camissonia nevadensis (Kell.) Raven, Brittonia 16: 285. 1964. Figure 74
Oenothera nevadensis Kell., Proc. Calif. Acad. 2: 224, fig. 70. 1863.
Sphaerostigma tortuosa A. Nels., Proc. Biol. Soc. Wash. 17: 95. 1904. Type:

Truckee Pass, Virginia Mts., Washoe Co., Nev., 16 June 1902, P. B.
Kennedy 766 (RM; isotypes, DS, UC).

Oenothera gauraeflora var. caput-medusae H. Lév., Monogr. Onoth. 226. 1905.
Lectotype: Foothills, Reno, 5,500 ft., Washoe Co., Nev., 10 June 1897,
collector unknown (MO; isotypes, POM, UC, US).

Oenothera gauraeflora var. vermiculata M. E. Jones, Contr. West. Bot. 12:

16. 1908.
Sphaerostigma nevadense (Kell.) Heller, Muhlenbergia 6: 51, with cover

illustration. 1910.

Erect, densely flowering from the base, the central stem 1-5 cm. tall,
the larger individuals with slender decumbent branches to 18 cm. long
radiating from-near the base of the plant, these naked below and with
a dense tuft of leaves and flowers at the ascending end; plants sub-
glabrous, the inflorescence strigulose. Leaves oblanceolate or narrowly
oblanceolate, the blades 1-3.5 cm. long, 2-7 mm. wide, entire, acumi-
nate at the apex, attenuate at the base; petioles 1-3 cm. long. Inflores-
cence erect. Hypanthium 2.2-3.2 mm. long, 1.2-1.3 mm. across at the
summit, glabrous within. Sepals 3.2-3.5 em. long, 1-1.1 mm. wide.
Petals 3-5 mm. long, 2.2-4.2 mm. wide. Filaments of the episepalous
stamens 4.5-4.8 mm. long, those of the epipetalous ones 3-4 mm. long;
anthers 0.4-1.5 mm. long. Style 6-7 mm. long, glabrous; stigma 0.5—-0.8
mm. thick, held somewhat above the anthers at anthesis. Capsule
highly contorted, 0.8-1.4 cm. long, 1-2 mm. thick near the base,
quadrangular in transection, tapering strongly from the swollen base,
with a prominent pale brown midrib running down the center of each
valve, held on the dead plant and shedding seeds very tardily. Seeds
monomorphic, 1.2-1.5 mm. long, 0.3-0.4 mm. thick, gray, finely
lacunose, much distorted by the walls of the capsule. Gametic chromo-
some number, n=7. Probably outcrossing, and possibly self-
incompatible.

Typr: From an unknown locality (doubtless the vicinity of Reno,
Washoe County, Nevada), CAS 838 in part (fragment).

Distripution (Figure 68): Local and colonial on vernally moist
clay flats in west-central Nevada, southernmost Washoe, Ormsby,

370 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Ficures 74-75.—Species of Camissonia, X 34: 74, C. nevadensis (Raven 17869, DS); 75, C.
hilgardii (Raven 18485, DS).

Storey, northernmost Lyon, and western Churchill Counties; 3,900-
4,600 ft. alt.
Voucher for chromosome number (1 individual), »=7 (7 pairs):

U.S.: NEVADA: WASHOE Co.: 2 mi. N. of Sparks, R17869 (Ds).
REPRESENTATIVE SPECIMENS EXAMINED:

U.S.: NEVADA: CHURCHILL co.: Fallon, Ross in 1914 (JEPS); ca. 8 mi. NW.
of Fallon, toward Soda L., Mills & Beach C-108 (RSA, UC). tyon co.: 5 mi.
5. of Fernley on U.S. Hwy. 95A, Mason 15189 (UC). ormsBy co.: Empire City,
Jones 3862 (CAS, GH, NY, POM, RM, UC, US); hills N. of Carson City, Stretch
in 1865 (NY). storey co.: 13 mi. E. of Sparks, Goodner & Henning 4 (POM).
WASHOE Co.: University Heights, Reno, 4,550 ft., Heller 9697 (DS, GH, PH, UC);
12 mi. N. of Reno on road to Pyramid Lake, 4,000 ft., Ripley & Barneby 4492
(CAS, NY); Truckee Pass E. of Reno, 4,500 ft., Heller 8647 (DS, GH, NY, PH,
US), Kennedy 1592 (NY, UC, US); near Wadsworth, Kennedy 2053 (DS); 9 mi.
N. of Poeville, 5,200 ft., Tillotson 120 (RSA, UC); 2.6 mi. NW. of Spanish Spring,
4,500 ft., Hendrix 823 (RSA, UC); 6 mi. SW. of Reno, Canby 99 (POM); 2 mi. N.

RAVEN—GENUS CAMISSONIA 371

of Sparks on Pyramid L. road, 4,700 ft., Archer 5077 (POM); near Pyramid L.,
Lemmon 239 (F; locality may be only an approximation).

Mature plants of this species are unmistakable; occasionally younger
ones are identified as Camissonia boothii subsp. alyssoides. The habitats
of these two taxa are distinct, and C. nevadensis is not known to occur
sympatrically with any other member of its section. Floral morphology
suggests that C. nevadensis is outcrossed, but it is not known whether
this species is self-incompatible. It probably is not, judging from what
is known of the other local entities within the genus and the tribe.
Camissonia nevadensis, one of the two species of its genus endemic to
Nevada (the other C. megalantha), is relatively rare, and a number of
its localities are being destroyed by urban expansion in the vicinity of
Reno. A few collections made by J. G. Lemmon suggest the occurrence
of this species in adjacent California, but Lemmon’s localities are
notoriously inaccurate, and it would seem to be much more probable
that he obtained this material in the vicinity of Reno or Pyramid
Lake, where he is known to have collected the species.

58. Camissonia minor (A. Nels.) Raven, Brittonia 16: 285. 1964. FIGuRE 76

Oenothera alyssoides var. minutiflora S. Wats., Bot. King Rep. 111. 1871.
Lectotype: Stansbury I., 2,300 ft., Tooele Co., Utah, June 1869, S. Watson
(US: isotype, NY); non Fisch. & Mey. 1835.

Sphaerostigma minus (‘minor’) A. Nels., Bull. Torrey Cl. 26: 130. 1899.

Sphaerostigma nelsoni Heller, Muhlenbergia 1: 1. 1900, nom. subs.

Oenothera chamaenerioides var. torta H. Lév., Monogr. Onoth. 230. 1905.
Lectotype: Granger, Sweetwater Co., Wyo., 3 June 1898, A. Nelson 4691
(MO; isolectotypes, F, GH, NY, RM, UC, US).

Sphaerostigma tortum (H. Lév.) A. Nels., Bot. Gaz. 40: 60. 1905.

Sphaerostigma tortum var. eastwoodae A. Nels., Bot. Gaz. 40: 61. 1905. Type:
Grand Junction, Mesa Co., Colo., May 1892, A. Eastwood (MO; isotypes,
GH, NMC, NY, OSC, UBC, US).

Sphaerostigma alyssoides var. minutiflorum (S. Wats.) Small, Bull. Torrey
Cl. 33: 146. 1906.

Oenothera minor (A. Nels.) Munz, Bot. Gaz. 85: 238. 1928.

Oenothera minor var. typica Munz, Bot. Gaz. 85: 239. 1928. .

Qenothera minor var. cusickit Munz, Bot. Gaz. 85: 240. 1928. Type: Malheur
R. and adjacent hills, Malheur Co., Oreg., 6 June 1901, W. C. Cusick 2545
(UC 35348; isotypes, F, GH, K, NY, POM, RM, UC, US).

Erect, flowering from the base, and often well branched from
below, the stems 3-30 cm. long; plants densely strigulose, often
grayish, the inflorescence often with a more or less evident admixture
of glandular trichomes. Leaves oblanceolate or narrowly oblanceolate,
the blades of the basal leaves 0.5-2.5 cm. long, 0.3-1.5 cm. wide, the
petioles 5-20 mm. long; leaves acuminate at the apex, attenuate at
the base. Inflorescence erect. Hypanthium 0.5-1.9 mm. long, 0.5—-1.2
mm. across at the summit, pubescent in lower half within. Sepals
0.8-1.8 mm. long, 0.35-0.7 mm. wide. Petals 0.8-1.3 mm. long,
0.4-1.3 mm. wide. Filaments 0.3-1.3 mm. long, the epipetalous ones

372 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Ficures 76-79.—Species of Camissonia: 76, C. minor, X 34 (Train 191, DS). 77-79, C.
pterosperma. 77, Habit, < 34 ( Leiberg 2075, DS); 78, ventral view of winged seed, X 65
(Heller 8376, DS); 79, habit of fruiting plant, X 34 (Heller 8376, DS).

RAVEN—GENUS CAMISSONIA 373

shorter than the episepalous ones and sometimes apparently abortive;
anthers 0.5-0.8 mm. long. Style 1.2-3.2 mm. long, pubescent near the
base; stigma 0.5-0.6 mm. thick, surrounded by the anthers at anthesis.
Capsule contorted, 1-2.5 em. long, 0.8-1.2 mm. thick, subterete, not
tapering sharply. Seeds monomorphic, 1.1-1.2 mm. long, 0.4-0.45 mm.
thick, gray, finely lacunose. Gametic chromosome number, n=7.
Autogamous or rarely cleistogamous.

Type: Green River, Sweetwater County, Wyoming, 31 May 1897,
A. Nelson 3047 (RM; isotypes, GH, NY, US).

DistriputTion (Figure 80): Occasional on sandy slopes and flats,
often with Artemisia tridentata, from Douglas and Lincoln Counties,
eastern Washington; Payette, Lehmi, Butte, and Bannock Counties,
southern Idaho; and Wyoming (Big Horn, 12 miles north of Graybull,
4,050 ft., Hamner 43, RM; Washakie, west of Worland, 4,100—4,300
ft., Nichols 374, RM; Sweetwater; and Carbon Counties) south
through southeastern Oregon (Lake, Harney, and Malheur Counties),
Utah (south to Beaver, Sevier, San Pete, Duchesne, and Uintah
Counties), and westernmost Colorado (Moffat and Mesa Counties)
to northern and central Nevada (Humboldt, White Pine, Eureka,
Elko, Pershing, Lander, Nye, and Washoe—near Verdi, Stokes in
1903, US—Counties), and northeastern California (valley east of
Cedarville, 4,000 ft., Modoc County, Ripley & Barneby 6006, CAS).

Vouchers for chromosome number (3 individuals, 3 populations),
n=7 (all 7 pairs):

U.S.: NEVADA! EUREKA co.: Near Lone Mt., cultivated at Los Angeles,
R15418. OREGON: HARNEY co.: 4.7 mi. 8. of Narrows, R18443; 12.9 mi. 5. of
Narrows, R18448.

This species is here taken in the same sense in which it was con-
stituted by Munz (Bot. Gaz. 85: 238-240) in 1928. Some of the popula-
tions from the western portions of the range of this species consist of
relatively small-flowered individuals, but these do not constitute the
sort of well-marked geographical race that I consider to merit formal
taxonomic recognition. Thus I do not recognize the taxon described as
Oenothera minor var. cusicki.

In some of the plants of Hitchcock & Muhlick 21200 (DS), from 21
miles north of French Glen, Harney County, Oregon, the epipetalous
stamens are extremely reduced, the apparently abortive anthers be-
ing subsessile. It is conceivable that some plants of this species might
be found which lack this set of anthers completely, as is the case in
some populations of C. andina and in both known populations of C.
exilis. In these cases, it would appear that the drastic reduction or loss
of the smaller set of anthers is associated with autogamy; in the other
genus of the tribe Onagreae containing species in which there has been
a loss of the epipetalous stamens, namely Clarkia, this is certainly not
the case.

374 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

é e
2° e °
e
ef ° .
e e ° ‘3 e °
e e
° e
e bd ®eee ® °
. z
e a e e e
ele * e %
e
e e
oe . A .e *
\ fe e
ENN
7
an

Ficure 80.—Western United States, showing range of Camissonia minor.

Section VIII. Chylismiella

Camissonia sect. Chylismiella (Munz) Raven, Brittonia 16: 282. 1964.
Oenothera sect. Chylismiella Munz, Amer. Journ. Bot. 15: 224. 1928.
Oenothera subg. Chylismia sensu Munz, Amer. Journ. Bot. 15; 224. 1928.

Annuals, the plants caulescent, not flowering at the basal nodes, the
leaves cauline; plants with exfoliating white rhytidome. Leaves sub-
sessile, sometimes purple-dotted. Inflorescence nodding at anthesis,
becoming erect in fruit. Flowers opening near sunrise. Ovary lacking a
sterile projection. Petals white, yellow at the base, fading purplish
after fertilization, without any dots or other markings. Stamens,
style, and inside of hypanthium yellow. Pollen shed singly. Capsule
stalked, regularly and promptly loculicidal, straight, or slightly curved,
somewhat torulose, terete, the seeds in two rows in each locule, appear-
ing as one by crowding. Seeds monomorphic, brown, with a convex
side and a concave side, the concave side surrounded by a thick wing,
this and the back densely covered with glassy, clavate trichomes.

TYPE SPECIES: Camissonia pterosperma (S. Wats.) Raven.

DisrrisuTion: Southeastern Oregon, south through Nevada and
western Utah to Inyo County, California, and northern Coconino and
Mohave Counties, Arizona.

The single species of this section is extremely distinctive and cannot
be allied with any other group of the genus with confidence. There is no

RAVEN—GENUS CAMISSONIA 375

reason to consider it related to sect. Chylismia, the only common feature
being the prominently stalked capsule. The color of the petals, white
with a yellow band near the base, is unique in Camissonia but wide-
spread in the related genus Gayophytum and found nowhere else in the
tribe Onagreae.
59. Camissonia pterosperma (S. Wats.) Raven, Brittonia 16: 282. 1964.
Figures 77-79

Oenothera pterosperma S. Wats., Bot. King Rep. 112, pl. 14. 1871.

Chylismia pterosperma (S. Wats.) Small, Bull. Torrey Cl. 23: 193. 1896.

Sphaerostigma pterospermum (S. Wats.) A. Nels., Bot. Gaz. 40: 63. 1905.

Slender and branching freely, 2-14 cm. tall; stems and leaves
entirely covered with hispid pubescence. Leaves narrowly lanceolate
to oblanceolate, 0.3-3 em. long, 0.1-0.6 cm. wide, entire, with a single
prominent vein, acute or acuminate at the apex, the base narrowly
cuneate or attenuate. Inflorescence glandular pubescent. Hypan-
thium 1-2 mm. long, 1-1.2 mm. across at the summit, glabrous with-
in. Sepals 1.5-2.5 mm. long, 0.8-1.2 mm. wide. Petals 1.5-2.5 mm.
long, 1-1.9 mm. wide. Filaments of the episepalous stamens 1-1.7 mm.
long, those of the epipetalous ones 0.5-1 mm. long; anthers 0.3—0.4
mm. long. Style 2.2-4 mm. long, glabrous; stigma 1-1.5 mm. thick,
surrounded by the anthers at anthesis. Capsule 1.2-1.8 cm. long, 1.2-
1.6 mm. thick, ascending or spreading; pedicel 4-8 mm. long, spread-
ing. Seeds 1-1.5 mm. long, 0.6-0.8 mm. thick, broader at one end, and
truncate at each end (where contacting next seed in row), the tri-
chomes longer at one end than at the other and shortest in the middle,
the two areas of trichomes (back and wings) separated by a narrow
glabrous band running around the seed. Gametic chromosome number,
n=7. Autogamous.

Type: Growing under sagebrush, Trinity Mountains, 5,000 ft.
(probably Churchill County), Nevada, May 1868, S. Watson (US
47918; isotypes, GH, NY).

Distripution: Well-drained slopes, often of volcanic orgin, with
Pinus edulis, Juniperus, and Artemisia tridentata, from south-eastern
Oregon (Lake and Malheur Counties) south through Nevada and
western Utah to Inyo County, California, and northern Mohave and
Coconino (upper end of House Rock Valley, Goodding 112-49, RM)
Counties, Arizona; widely distributed but rare. From 2,500-8,000 ft.
alt. See Univ. Calif. Publ. Bot. 34: fig. 2. 1962.

Section IX. Nematocaulis

Camissonia sect. Nematocaulis Raven, Brittonia 16: 285. 1964.
Oenothera subg. Sphaerostigma sensu Munz, Bot. Gaz. 85: 234. 1928; pro
parte.
Oenothera sect. Nematocaulis (Raven) Munz, N. Amer. Fl. II. 5: 155. 1965.
Annuals, the plants caulescent, flowering from near the base, but
the lower stem usually naked, the leaves cauline and densely tufted,

376 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

nearly every one bearing a flower in its axil; many slender branches
often arising from near the base, these naked below, the leaves and
flowers tufted near the end of each as on the main stem. Leaves sub-
sessile. Inflorescence erect. Flowers opening near sunrise. Ovary lack-
ing a sterile projection. Petals yellow, not fading purplish after fertili-
zation, without any dots or other markings. Stamens, style, and inside
of hypanthium yellow. Stamens eight, the epipetalous ones sometimes
greatly reduced, or four. Pollen shed singly. Capsule sessile, regularly
loculicidal, straight or nearly so, somewhat torulose, strongly flattened,
the central axis falling free at maturity, the seeds in one row in each
locule, crowded. Seeds monomorphic, narrowly obovoid, light to dark
brown, smooth and shining.

TYPE SPECIES: Camissonia hilgardit (Greene) Raven.

DistrRiBuTion: Southern British Columbia and Alberta, south
through westernmost Montana, western Wyoming, eastern Washing-
ton, and eastern Oregon to northern and central Utah and Nevada
and northeastern California, east of the Cascade axis.

The two species assigned to this section are very closely related
but entirely distinct within the genus; they are not similar to any
other group with which IT am familiar. The smaller flowered tetraploid
and hexaploid populations, comprising Camissonia andina, have an
extremely wide distribution, whereas the larger flowered diploids,
C. halgardiz, have a relatively narrow range almost entirely in eastern
Washington. On the basis of present knowledge of the group, it would
appear likely that the tetraploids have had an autopolyploid origin,
and that the hexaploids may have been derived directly from them
by the functioning of an unreduced gamete. Such a hypothesis would
if true postulate the existence of strict genetic control of pairing in
this group, as no multivalents have been observed at diakinesis and
metaphase I in the polyploids; but this must be true of the members of
sect. Holostigma and sect. Camissonia, so it may also apply here. As
there is only one series of diploid populations known in this extremely
distinct group, no alternative seems to suggest itself.

Occasional plants of the highly autogamous, sometimes even
cleistogamous, member of sect. Nematocaulis, Camissonia andina,
have 3-merous flowers; these have been named Oenothera andina f.
tripetala by Léveillé and O. andina var. anomala by M. E. Peck.
Such flowers are unknown elsewhere in the genus but appear again
in another genus of the tribe, Gaura, where they characterize one
autogamous entity, G. hexandra Gémez Ortega subsp. hexandra, and
are found very frequently in another autogamous species, G. angust?-
folia Michx. Elsewhere in the family 3-merous flowers occur regularly
only in the highly autogamous (sometimes cleistogamous) African
Ludwigia sect. Prieurea, consisting of a single variable species. Thus

RAVEN—GENUS CAMISSONIA 377

there appears to be a strong correlation between autogamy and the
reduction in number of flower parts in the family Onagraceae. The
same may be said of the loss of epipetalous stamens, which occurs
in occasional plants (perhaps entire populations?) of Camissonia andina
(e.g., two from Blaine County, Idaho: Macbride & Payson 3056,
POM; Macbride 806, DS). As noted above, there is a strong tendency
toward this condition in some individuals of the highly autogamous
Camissonia minor, and it is a consistent feature in the two known
populations of C. ezilis. On the other hand, there is certainly no con-
nection between autogamy and the loss of the epipetalous stamens in
Clarkia or in Ludwigia, with a single exception in the latter: relatively
small-flowered (and presumably more highly autogamous) individuals
and populations of L. inelinata (L. f.) Raven tend to have only four
stamens and have been named L. verticillata and placed in another
section of the genus by Munz.
60. Camissonia hilgardii (Greene) Raven, Brittonia 16: 285. 1964. Figure 75

Oenothera hilgardi Greene, Bull. Torrey Cl. 10: 41. 1883.

Sphaerostigma hilgardi (Greene) Small, Bull. Torrey Cl. 23: 188. 1896.

Sphaerostigma andinum var. hilgardii (Greene) A. Nels., Bot. Gaz. 40: 56.

1905.
Oenothera andina var. hilgardii (Greene) Munz, Bot. Gaz. 85: 251. 1928.

Similar to C. andina, but the flowers larger. Hypanthium 1.5-2 mm.
long, ca. 1.2 mm. across at the summit, sparingly pubescent in lower
portions within. Sepals 2-3 mm. long, 0.9-1.1 mm. wide. Petals 2.5-5
mm. long, 2.2-3.5 mm. wide. Filaments of the episepalous stamens
2-4 mm. long, those of the epipetalous ones 1.5-2.8 mm. long; anthers
of the longer stamens 0.7-0.8 mm. long, those of the shorter 0.5-0.6
mm. long. Style 4.5-6 mm. long, sparsely pubescent near the base;
stigma 0.7-0.8 mm. thick, held at or slightly above the anthers of the
longer stamens at anthesis. Gametic chromosome number, n=7.
Probably self-compatible but outcrossing.

Type: Moist alkaline soil of the Klickitat Swale, Klickitat or
Yakima County, Washington, July 1882, £. Hilgard (GH).

Disrripution (Figure 81): Sandy or clay slopes under Artemisia
tridentata, scattered but locally common, occasionally growing with
C. andina, in Okanagan, Chelan, Douglas, Grant, Kittitas, and
Yakima Counties, Washington; collected twice along the lower
Columbia River, at Bingen, Klickitat County, Washington, and
Hayden Island, Multnomah County, Oregon. From near sea level to
ca. 1,600 ft. alt.

Vouchers for chromosome number (2 individuals, 2 populations),
n=7 (7 pairs in each):

U.S.: WASHINGTON: KITTITAS co.: 16.7 mi. W. of Ginkgo Petrified Forest,
R18485. yAKIMA co.: 2.3 mi. 8S. of L. Wenas, R18483.

378 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

61. Camissonia andina (Nutt.) Raven, Brittonia 16: 285. 1964.

Oenothera andina Nutt., in Torr. & Gray, Fl. N. Amer. 1: 512. 1840.

Sphaerostigma andinum (Nutt.) Walp., Rep. 2: 79. 1843.

Oenothera andina f. tripetala H. Lév., Monogr. Onoth. 182. 1905. No specimens
cited.

Oenothera andina var. typica Munz, Bot. Gaz. 85: 250. 1928.

Oenothera andina var. anomala M. E. Peck, Torreya 32: 151. 1932. Type:
Dry slope 6 mi. NW. of Paisley, Lake Co., Oregon, 15 July 1927, M. E.
Peck 15176 (WILLU).

Plants finely strigulose throughout, more densely so in the inflores-
cence, especially on the ovaries. Branches 1-15 cm. long. Leaves
very narrowly oblanceolate, 1-3 cm. long, 0.1—-0.25 cm. wide, entire,
acuminate at the apex, attenuate at the base. Hypanthium 0.8-2 mm.
long, 0.75-1.1 mm. across at the summit, glabrous or more rarely
sparsely pubescent in lower half within. Sepals 0.8-2 mm. long,
0.5-1.1 mm. wide. Petals 0.8-2.3 mm. long, 0.6-1.2 mm. wide. Fila-
ments of the episepalous stamens 0.45-2.2 mm. long, those of the
epipetalous ones 0.1-0.45 mm. long, the epipetalous stamens rarely
lacking; anthers of the longer stamens 0.2-0.45 mm. long, those of
the shorter ones 0.08-0.2 mm. long. Style 1.7-3 mm. long, glabrous
or more rarely sparsely pubescent near the base; stigma 0.4—-0.6 mm.
thick, surrounded by the anthers at anthesis. Capsule (0.5-)0.8-1 cm.
long, 1-1.3 mm. thick along the wider axis, ascending. Seeds 0.7-1.3
mm. long, 0.3-0.35 mm. thick. Gametic chromosome numbers, n= 14,
21. Autogamous, more rarely cleistogamous.

Typr: Dry plains near the Blackfoot River, Bingham or Caribou
County, Idaho, 10-12 July 1834, T. Nuttall (BM; isotypes, GH, K,
NY, PH).

Disrripution (Figure 81): Vernally moist flats, often in clay soil
under Artemisia tridentata or in pinyon-juniper association, inconspic-
uous and often overlooked, from the vicinity of Lake Osoyoos, British
Columbia, and Medicine Hat, Alberta, south to Missoula County,
Montana (Missoula, Hitchcock 2326, CAS, POM, RSA, WS); the
western half of Wyoming; Cache, Wasatch, and Beaver Counties,
Utah; Elko, Eureka, Humboldt, northern Nye, and Ormsby Counties,
Nevada; and from Sierra County, California, north along the east
side of the Cascades throughout eastern Oregon and Washington, and
west along the Columbia River to Skamania County, Washington;
from ca. 1,600-ca. 6,500 ft. alt. I have seen a specimen labeled ‘Golden
City, etc., Colorado,” Greene in 1873 (US), but the occurrence of
Camissonia andina in that state needs confirmation.

Vouchers for chromosome number (4 individuals, 4 populations),
n=14:

U.S.: CaLirorNIA: LASSEN co.: Near Eagle L., Ornduff 4205 (LA). Ipauo:
FREMONT Co.: Near St. Anthony, R19560. OREGON: LAKE co.: W. of Lakeview,

RAVEN—GENUS CAMISSONIA 379

:
xe" C re)
ra }
Zi 4a
! s
ao
4 40 9,40 oo
fa200%o 9 PO
4 02
4,44 0 O
oO
St O ay ie)
° fo)
a_ 0-04 00 2
fe)
°
O eo) fe) —
Oo O
fe) e
fe) o fe)
fo)
0 oO o fe) tc)
Coe 0% o 5 ° 0
ro SC ogo 8 ° fe) °
fe) oe Oo ®@ fe)
fe) Q Oo fe)
fe) Qo oO fo) fe)
oo ° °
o fe) (e) ° fe}
fe)
Oo, ra) °
fe) Oo fo)
O° Oo O° °
o°
e a
0°? 5
fo) ° o ro)

: cv

Ficure 81.—Western United States, showing range of species of Camissonia sect. Nema-
tocaulis: M=C. hilgardii; O=C. andina, with (=tetraploid chromosome counts
(n= 14) and @=hexaploid chromosome counts (n=21).

R18429. WASHINGTON: DOUGLAS co.: State Hwy. 10, 4.6 mi. N. of junction with
U.S. Hwy. 2, R18495.

Vouchers for chromosome number (5 individuals, 5 populations),
n=21:

U.S.: IDAHO: ELMORE co.: 19 mi. E. of Dixie, R18524 (54 percent of pollen
4-pored). NEvapA: ELKO co.: East Humboldt Mts., R13456. OREGON: HARNEY
co.: 3.4 mi. E. of Buchanan, R18458 (46 percent of pollen 4-pored) ; 14 mi. NE.
of Wagontire, R13356. MALHEUR co.: Idaho State line on U.S. Hwy. 95, R19581.

Camissonia andina grows sympatrically with its diploid relative,
C. hilgardvi, at a number of localities within the range of the latter.
With the accumulation of additional chromosome number determi-
nations from C. andina, it should become possible to develop a method
for distinguishing tetraploids and hexaploids. Whether it will then

380 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

become desirable to accord formal taxonomic recognition to tetrat-
ploid and hexaploid populations remains to be seen. In general, it now
appears that hexaploids often have 40-90 percent 4-pored pollen grains,
tetraploids mostly 10 percent or fewer (often none). Several of the
collections examined (without chromosome number determinations)
had 10-40 percent 4-pored pollen, and more chromosome counts will
be necessary to draw a clear line between plants at the two different
polyploid levels. At any rate, it is virtually certain that the type
collections of this species is hexaploid (n=21), as an isotype (N Y) had
27 of the 50 pollen grains examined 4-pored.

The limited amount of evidence now available suggests that tetra-
ploid races occupy the margins of the range of this species and are
infrequent in the main portion of its range, where hexaploids appear to
predominate. I have thus far been unable to discover any morpho-
logical distinctions between the two races other than the pollen
difference just discussed.

Excluded Species

Sphaerostigma andinum var. minutum A. Nels., Bot. Gaz. 40: 56. 1905=Gayo-
phytum sp. (Type: without data, MO).

Oenothera micrantha Presl, Rel. Haenk. 2: 31. 1827; non Hornem. ex Spreng’
1825. Type: Chile, Haenke (PR). Holostigma paradoxum Spach, Nouv. Ann:
Mus. Paris III. 4: 334. 1835. Sphaerostigma paradoxum (Spach) Gay, Fl. Chil:
2: 329. 1846. Chamissonia paradoxa (Spach) Reiche, Anal. Univ. Chile 98: 482°
1897=Gayophytum micranthum Hook. & Arn. (cf. Lewis & Szweykowski’
Brittonia 16: 389. 1964).

Appendix

Chromosome numbers of Camissonia sect. Holostigma *

In the following lists are grouped the vouchers for chromosome
counts in Camissonia sect. Holostigma. Collections made by Raven
are preceded by Ff, those by Breedlove by B, and those by Wedberg
by W. As usual throughout this paper, vouchers are presumed to be
deposited at DS unless some other indication of place of deposit is
made. In this section, “SDSC” is used for the herbarium of San Diego
State College, where some of the vouchers are deposited; abbreviations
for other herbaria are standard. In general, Wedberg counted his own
collections and Raven counted the others, often with the assistance
of Breedlove.

In addition to the original counts reported here, we have also
included 34 counts reported for plants of this group by Lewis, Raven,
Venkatesh, & Wedberg (Aliso 4: 73-86. 1958). The only earlier
chromosome number determinations for the group appear to be those
by Johansen (Amer. Journ. Bot. 16: 595-597. 1929; Proc. Nat. Acad.
Sei. U.S. 15: 882-885. 1929) of n=7 in “Sphaerostigma veitchianum”’
and “S. spirale” (=Camissonia bistorta and C. cheiranthifolia, respec-
tively), but without any indication of the source of the materials
examined.

Including the 34 earlier counts for which voucher specimens are
available, a total of 389 individuals from 335 populations have been
examined cytologically. These populations represent all 17 taxa recog-
nized for the section, including 14 species and 3 additional subspecies.
Cytogenetically, the most striking characteristic of these counts is the
almost complete lack of rings of chromosomes such as are associated
with the presence of reciprocal translocations. Lewis et al. (1958, p. 80)
reported 12 pairs and a ring of 4 chromosomes in one plant of Camis-
sonia confusa (Lewis & Lewis 1669, LA; as Oenothera micrantha var.
ignota). This ring of 4 chromosomes may have resulted from the pres-
ence of a reciprocal translocation, or it may have been because of ho-
mology between the pairs of chromosomes involved in it. At any rate,
it was the only individual, out of 389 examined, in which such a ring
of chromosomes was found. In contrast, reciprocal translocations oc-
curred in over 20 percent of the 676 individuals of Camissonia sect.

‘This work was done in cooperation with Dennis E. Breedlove and Hale L.
Wedberg (San Diego State College).
381

382 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Chylismia that were examined cytologically (Raven, Univ. Calif. Publ.
Bot. 34: 20-43, 1962), and similar percentages probably occur in sect.
Eremothera and are characteristic of other groups of the tribe Onagreae,
such as the genus Clarkia. In view of the complete absence of reciprocal
translocations in natural populations of sect. Holostigma, it is also of
considerable interest that not one of the 389 individuals of this section
(with the possible exception noted above) examined cytologically had
a supernumerary chromosome, as compared with the 26 of the 676
individuals of sect. Chylismia. This would appear to lend considerable
weight to Lewis’ (Evolution 5: 142-157, 1951) suggestion of a causal
relationship between reciprocal translocation (presumably by occa-
sional misdivision of rings of chromosomes) and supernumerary
chromosomes in Onagreae.

It is also noteworthy that little evidence of autopolyploid pairing has
been found in the two tetraploids and three hexaploids of this section,
even though the diploid species contributing to their formation are
obviously extremely closely related. This subject is discussed further
on p. 176.

Camissonia bistorta (Nutt. ex Torr. & Gray) Raven, (29 individuals, 22 popu-
lations), n=7:

U.S.: CALIFORNIA: LOS ANGELES Co.: Point Dume, W13, in 1958, 169 (SDSC),
in 1957 (LA), H (LA), Lewis 1634 (LA); Playa del Rey, WI (LA), 562 (SDSC);
Mulholland Drive, Santa Monica Mts., W260 (SDSC). orANGE co.: Corona del
Mar, W172; 6.5 mi. E. of San Juan Capistrano, R17491 (progeny = R18782); 5.6
mi. NE. of San Juan Capistrano, W178; Dana Point, W176; lower edge of Cleve-
land National Forest just above San Juan Guard Station, 850 ft., R18783. RIVER-
SIDE Co.: 6.5 mi. SE. of Hemet, B1896; 5 mi. N. of Alberhill, W3; Upper San Juan
Campground, W188 (SDSC); Lake Mathew Road, W196; 5.1 mi. W. of Dripping
Springs, R14023; 0.5 mi. NW. of Alberhill, WB (LA); Temeseal Canyon, Lewis
1630 (2 plants; LA); ca. 2 mi. N. of Aguanga, Klein 1300 (RSA). SAN BERNARDINO
co.: U.S. Hwy. 70-99, 0.15 mi. W. of Etiwanda Ave., W26 (LA). SAN DIEGO Co.;
La Mesa, W922 (SDSC); 2.2 mi. W. of Campo, R16926; Torrey Pines Park, W211
(SDSC); Balboa Park, San Diego, W213 (SDSC).

BAJA CALIFORNIA. 3 mi. 8. of Ensenada, Klein & Gregory 1272.

Camissonia cheiranthifolia (Hornem. ex Spreng.) Raimann subsp. cheiranthifolia
(8 individuals, 8 populations), n=7:

U.S.: CALIFORNIA: HUMBOLDT Co.: 8.8 mi. N. of Big Lagoon, T. & L. Mosquin
4489. LOS ANGELES Co.: San Clemente I., R17627 (progeny). SACRAMENTO CO.:
Brannan I, State Park, near Rio Vista, R20178. sANTA BARBARA co.: Jalama
Beach, R14040; San Miguel I., Blakley 5099 (the form called Oenothera nitida
Greene, at its type locality); Santa Cruz I., Fraser Point, R15312; Santa Rosa I.,
R14995. sonoma co.: Bodega Bay to Bodega Head, R18247.

Camissonia cheiranthifolia subsp. suffruticosa (S. Wats.) Raven (7 individuals,
7 populations), n=7:

U.S.: CALIFORNIA: LOS ANGELES Co.: Point Dume, R13855 (LA, RSA).
SAN DIEGO Co.: 8. of Carlsbad, R14027; Torrey Pines State Park, W210 (SDSC).
SANTA BARBARA Co.; Devereaux Point, B1766 (UCSB); Goleta Point, R14044.
VENTURA CO.: Ventura, R13999,

BAJA CALIFORNIA: 3 mi. S. of Ensenada, R17009 (2 plants).

RAVEN—GENUS CAMISSONIA 383

Camissonia confusa Raven (29 individuals, 22 populations), n=14:

U.S.: ARIZONA: MARICOPA Co.: 12.6 mi. E. of Tortilla Flat on State Hwy. 88,
2,700 ft., R173738, 12.9 mi. E., R17375. CatiroRNIA: LOS ANGELES CO.: Native to
Rancho Santa Ana Botanic Garden, Claremont, R15535, 17748, 18139. ORANGE
co.: Lower San Juan Campground, Santa Ana Mts., Lewis 1642 (LA), W183.
RIVERSIDE Co.: Upper San Juan Campground, Santa Ana Mts., W187 (SDSC) ;
Ortega Hwy. 3.9 mi. above its E. base, Santa Ana Mts., 2,400 ft., R17488; Reche
Canyon, Loma Linda, 1,900 ft., R17471, 18766; 7.7 mi. E. of Hemet, WO (LA),
198; 10.3 mi. S. of Hemet, W204 (SDSC); 6.6 mi. from Banning on Idyllwild road,
Lewis 1667 (LA); 7.4 mi. from Banning on Idyllwild road, W303 (SDSC); Hwy.
74 ca. 11 mi. W. of Elsinore, Santa Ana Mts., W191 (SDSC). SAN BERNARDINO
co.: Oak Glen Road 2.5 mi. E. of junction with road from Yucaipa to Mill Creek,
San Bernardino Mts., 3,300 ft., R17469; 2.3 mi. W. of Mill Creek Ranger Station,
2,300 ft., R17466 (type collection). SAN DiEGO co.: 2.7 mi. SE. of Aguanga, 2,400
ft., R17388; 0.6 mi. W. of Banner Queen Store, 3,000 ft., R17422; State Hwy. 79,
2.2 mi. S. of junction with State Hwy. 76, N. of Santa Ysabel, 3,300 ft., R17395;
road to Mt. Palomar 3.9 mi. NW. of turn to L. Henshaw, 4,000 ft., R17397; L.
Henshaw, 2,800 ft., R17396; 2.5 mi. W. of Julian, H. & M. Lewis 1669 (2 plants;
LA). SAN LUIS oBISPo co.: La Panza Campground, 2,000 ft., R16968, 18323.
VENTURA Co.: Cow Spring Campground, 17.6 mi. N. of Fillmore, Topatopa Mts.,
3,500 ft., B2643.

Camissonia guadalupensis (S. Wats.) Raven subsp. clementina (Raven) Raven
(3 individuals, 3 populations), n=7:
U.S.: CALIFORNIA! SAN CLEMENTE I., LOS ANGELES Co.: Eel Point, R17125 (type
collection); beach due W. of Wall 2, R17260; W. side of island opposite Wilson
Cove, R17628 (progeny = R18176).

Camissonia guadalupensis subsp. guadalupensis (2 individuals, 1 population),
n=T7:
BAJA CALIFORNIA: Arroyo Melpomone, Guadalupe I., Moran & Ernst
6737 (2 plants in progeny =R17530, 18175).

Camissonia hardhamiae Raven (3 individuals, 3 populations), n=21:

U.S.: CALIFORNIA! SAN LUIS OBISPO Co.: 3 mi. E. of Santa Margarita, R18313;
Calf Canyon, 3.7 mi. NE. of Santa Margarita, R18315 (type collection) ; 4.2 mi.
NE. of Santa Margarita, at junction to Parkhill District, R18318.

Camissonia hirtella (Greene) Raven (84 individuals, 76 populations), n=7:
U.S.: CALIFORNIA: AMADOR Co.: 1.5 mi. N. of Buena Vista, R18351. FRESNO Co.:
Copper Creek Trail, Kings Canyon, 5,500 ft., Howell 34332 (progeny = R19082,
19748); Simpson Meadow, Middle Fork of Kings R., Howell 33793 (progeny =
R18989, 19755); 8 mi. N. of Pine Ridge, B15277. KERN Co.: Black Mt. burn,
Greenhorn Mts., Howell 18779 (progeny =R18779, 19746) ; Breckinridge Mt.,
7,400 ft., T’wisselmann, 4704 (progeny = R19080) ; 3.5 mi. E. of Alta Sierra, Green-
horn Mts., R20249; ca. 1 mi. S. of Havilah, Lewis 1344. LAKE co.: 4 mi. E. of Clear
Lake, B5150; 4 mi. NW. of Lower Lake, B4542; 2 mi. NE. of Lower Lake, B4539;
Lower Lake, R18241. Los ANGELES co.: Santa Monica Mts.: Latigo Canyon Road
0.4 mi. from Mulholland Hwy., W287 (SDSC) ; Corral Canyon, R17512A; Topanga
Canyon, W308; Piuma Road 5.1 mi. NW. of Rambla Pacifica, 700 ft., R15369;
Mulholland Hwy. just E. of junction with Decker Road, 1,600 ft., R17516; Saddle
Peak Road 0.9 mi. above Stuntz Canyon Road, 2,200 ft., R17510; 0.6 mi. above
junction of La Tuna Canyon on road to Saddle Peak, 2,050 ft., R17506; Soledad
Canyon, Lewis 1677 (LA). MARIPOSA co.: 3.7 mi. NW. of Coulterville, 2,000 ft.,
R18347. MONTEREY co.: Hanging Valley, NW. side of Junipero Serra Peak, B5599.
RIVERSIDE Co.: 2 mi. S. of Banning, WM (LA); west of Mountain Center, San

295-655 O—sb8——15

384 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Jacinto Mts., W304, 305 (SDSC) ; Upper San Juan.Campground, Santa Ana Mts.,
Lewis 1655 (LA); Upper San Juan Campground to Hwy. 74, W190 (SDSC);
10.3 mi. S. of Hemet on Hwy. 79, W202 (SDSC); Dripping Springs Camp-
ground, 1,700 ft., R17384; Ortega Hwy. 1 mi. above E. base, 1,600 ft., R17487,
3.9 mi. above, R17489; Main Divide Truck Trail at Ortega Hwy., 2,300 ft.,
R17762 (progeny = R18845); Main Divide Truck Trail 8.1 mi. S. of Santiago
Peak, Santa Ana Mts., 4,050 ft., R17760; Vandeventer Flat, San Jacinto Mts.,
WN-1(LA). SAN BERNARDINO Co.: 1.5 mi. W. of Mill Creek Ranger Station,
2,300 ft., R17467; State Hwy. 30, 2 mi. N. of National Forest Boundary, 2,000 ft.,
R17465, 4.5 mi. N. of boundary, 2,800 ft., R17464; road to Barton Flats 6.5 mi. N.
of Mill Creek, R17926 (cleistogamous). SAN DIEGO CO.: 1 mi. from Santa Ysabel
road to Julian, 3,000 ft., W928 (SDSC); Lake Wohlford, R20169; 0.9 mi. from
U.S. Hwy. 395 on Deer Springs Road to San Marcos, 700 ft., R17473; State Hwy.
94, 1.5 mi. W. of turn to Tecate, 1,600 ft., R17449; 0.5 mi. E. of Santa Ysabel,
3,300 ft., R17339, 17392, 17398; 1.1 mi. E. of Campo, 2,900 ft., R17444; 1.1 mi. E.
of Campo, 2,900 ft., R17442, 17443; 3.7 mi. N. of Cameron Corners on road to Pine
Valley, 3,500 ft., R17441; 3.1 mi. from U.S. Hwy. 80 on road to Morena Lake,
3,400 ft., R17440; 1.0 mi. E. of Oak Grove, 2,700 ft., R17389; County Road S6
just 8. of turn to Felicita County Park, R16948; 5.6 mi. W. of Campo, 2,600 ft.,
R17448; 3.8 mi. from turn to Escondido on road to Mt. Palomar, 4,100 ft., R17398;
2.3 mi. 8. of Rincon Springs on road to Escondido, cultivated at Stanford, R19076;
3.5 mi. NE. of Lake Wohlford, 1,700 ft., R17404; 8.6 mi. E. of Ramona, 2,800 ft.,
R17413; 2.5 mi. W. of Santa Ysabel, 3,100 ft., R17416; 2.1 mi. E. of Santa Ysabel
3,700 ft., R17418; 0.6 mi. above Banner Queen Store, R16922, 1.3 mi. W., 3,000
ft., R17419, 0.8 mi. B., R16924; Jamul-Lyons Valley road 0.7 mi. NE. of turn to
Camp Barrett, 2,200 ft., R17432; U.S. Hwy. 80, 0.7 mi. W. of Guatay Summit,
4,000 ft., R17436. san Lu1s OBISPO co.: S. road to Adelaida 6.4 mi. from its june-
tion with Oak Flat Road, Zewis & Thompson 1099 (LA, RSA, UC). sanTa BAR-
BARA CO,: 1.5 mi. W. of Juncal Campground, upper Santa Ynez Valley, 1,750 ft.,
B2236; 4.4 mi. E. of Santa Ynez Peak along El Camino Cielo, 3,300 ft., B2689;
Tunnel Road 2.5 mi. N. of U.S. Hwy. 150, Mission Canyon, 800 ft., B1902; above
Mono Debris Dam, upper Santa Ynez Velley, 1,500 ft., B2244. SANTA CLARA CO.:
1.5 mi. E. of Red Mt. near summit of Del Puerto Road, Mt. Hamilton Range,
B4889; upper Arroyo del Puerto 2.4 mi. E. of junction in San Antonio Valley,
R18230; summit of Loma Prieta, Santa Cruz Mts., B4675, 5011. VENTURA Co.:
Cow Spring Campground, 17.6 mi. N. of Fillmore, Topatopa Mts., 3,500 ft.,
B2644; U.S. Hwy. 399 at turn to Rose Valley, 3,600 ft., R17499; just N. of
Wheeler Gorge Guard Station, 1,950 ft., R17498; 1.2 mi. S. of Wheeler Springs,
1,400 ft., R17496A; Camp Ozena, Ozena Valley, 3,800 ft., B2436; Sespe Creek 7.6
mi. 8. of Sandstone Camp, 3,700 ft., R17500.

BAJA CALIFORNIA: 10.9 mi. N. of Santo Tomds, 600 ft., R17014; Santa
Catarina, Broder 436 (progeny = R18992, 19754).

Camissonia ignota (Jeps.) Raven (22 individuals, 18 populations), n=7:

U.S.: CALIFORNIA: LOS ANGELES Co.: Soledad Canyon, Lewis 1676 (3 plants;
LA, RSA). orance co,: Lower San Juan Campground, Santa Ana Mts., W180
(SDSC). r1veRsipE co.: 5 mi. NW, of Alberhill, W in 1967 (2 plants; LA); State
Hwy. 74, 1.6 mi. NE. of junction with State Hwy. 71, near Elsinore, R16914;
5.7 mi, SE. of U.S. Hwy. 395 on State Hwy. 74, road to Hemet, 1,600 ft., B1898;
Reche Canyon, R17472; Dripping Springs Campground, 1,700 ft., R17385; Lake
Mathews road 14.7 mi. W. of U.S. Hwy. 395, R16909. san pIEGo co.: NW. of
Fallbrook, W206 (SDSC); 2.3 mi. 8S. of Rincon Springs on road to Escondido,
1,100 ft., R17402 (progeny =R18787, 19751); 2.7 mi. SE. of Aguanga, 2,400 ft.,

RAVEN—GENUS CAMISSONIA 385

R17387; Lyons Valley along road 8 mi. NE. of Jamul, 1,800 ft., R17434. san LUIS
OBISPO Co.: 1.2 mi. S. of Santa Margarita to Simmler road on road to Navajo
Camp, La Panza Range, 2,000 ft., R16970. sANTA BARBARA CoO.: Juncal Camp-
ground, upper Santa Ynez Valley, 1,800 ft., B2229; 8. bank of Cuyama R. on
State Hwy. 166, 1.6 mi. E. of Tepusquet Canyon Road, 1,800 ft., B1930. VENTURA
co.: Boney Ridge, Santa Monica Mts., grown at Stanford, R18173.

Camissonia intermedia Raven (60 individuals, 53 populations), n= 14:

U.S.: CALIFORNIA: CONTRA Costa co.: Mt. Diablo, B4698. LAKE co.: 4 mi. W.
of Reiff, R20213. Los ANGELES co.: Soledad Canyon, Lewis & Thompson 1616
(LA); near Bird Park, Memorial Canyon, Avalon, Santa Catalina I., R17857,
18171. SANTA MONICA MTS., LOS ANGELES Co.: just S. of Agoura, 850 ft., R17520;
Corral Canyon, 1,900 ft., R17512 (progeny =18756); 0.9 mi. W. of Seminole
Hot Springs, 800 ft., R17518; 2.6 mi. Ek. of Seminole Hot Springs, R17517 (type
collection); Mulholland Hwy. just E. of junction with Decker Road, 1,600 ft.,
R17515; Stuntz Canyon Road 3.1 mi. below junction to Saddle Peak, 1,200 ft.,
R17511; Saddle Peak Road 0.9 mi. above Stuntz Canyon Road, 2,200 ft., R17509;
Saddle Peak Road 2.4 mi. above junction of La Tuna Canyon Road, 2,400 ft.,
R17507; La Tuna Canyon 3.6 mi. N. of Coast Highway, 1,800 ft., R17504.
MONTEREY Co.: Big Sandy Valley 2 mi. N. of Indian Valley Road, R18289;
Hastings Reservation, R18220, 18218; just N. of Hastings Reservation, R18225,;
road to Jamesburg 0.8 mi. from main road up Carmel Valley, R18227; W. edge
of San Antonio Valley, Santa Lucia Mts., 1,350 ft., B2267; 2.1 mi. SW. of Arroyo
Seco Guard Station, 1,250 ft., B2305; 6 mi. W. of Priest Valley on State Hwy. 198,
2,500 ft., B2376; 0.6 mi. SW. of San Benito Co. line on road from Bitterwater to
King City, R18201; The Indians, Santa Lucia Mts., R18213, 18264, 18268;
Lowes Canyon, R18293, 18298, RIVERSIDE co.: Machado Road 1 mi. SW. of
U.S. Hwy. 71, W193, 194 (SDSC); 3.3 mi. W. of Dripping Springs Campground,
1,500 ft., R17382. san BENITO Co.: 3.7 mi. from Hernandez on road to New Idria,
R15087. SAN DIEGO co.: 2.7 mi. W. of De Luz on road to Fallbrook, 600 ft.,
R17483; 2.4 mi. from Fallbrook on road to De Luz, R17477 (progeny = R18758) ;
road to Fallbrook at junction with U.S. Hwy. 395, 950 ft., R17484. san LUIS
OBISPO CO.: 1.2 mi. W. of Bee Rock, R18305; near junction 1.8 mi. W. of Paso
Robles, R18327. SANTA BARBARA Co.: Head of Buckhorn Canyon, summit of San
Rafael Mts., 2,600 ft., B2779; Alamo Pintado Creek, 7-8 mi. N. of Los Olivos,
Birbent Canyon, base of Figueroa Mt., ca. 1,400 ft., B2498; Little Agua Caliente
Hot Springs, 1.5 mi. NE. of Mono Debris Dam, 1,700 ft., B2230; 0.9 mi. N. of
State Hwy. 150 on San Roque Road, Santa Barbara, 800 ft., B2228; Portazuela
Grade, Santa Cruz I., R15197; Prisoners Harbor, Santa Cruz I., R18172. stan-
ISLAUS Co.: 12 mi. W. of Patterson on road to Mt. Hamilton, B4856. VENTURA Co.:
Cow Springs Campground, 17.6 mi. N. of Fillmore on Cow Springs Road, Topatopa
Mts., 3,500 ft., B2617; 0.3 mi. S. of Camp Comfort, 8S. of Ojai, R17493 (progeny =
18755); 1.2 mi. S. of Wheeler Springs, R17496 (progeny = 18768) ; Little Sycamore
Canyon, Santa Monica Mts., Lewis 1666 (LA, RSA).

BAJA CALIFORNIA: 10 mi. S. of Ensenada, R17053; 15.2 km. N. of San
Vicente, R12308; 5.5 km. S. of San Vicente, R17019; 19.5 km. SE. of San Vicente,
R12263; 2.5 mi. 8. of Ejido México, R17040; 4km. S. of Ejido México, R12234;
11.3 mi. N. of Colonia Guerrero, R17037; 8.5 km. E. of El Rosario, R12429; Sum-
mit of Aguajito Grade, 2 km. E. of Rancho Aguajito, R12523.

Camissonia lewisii Raven (23 individuals, 19 populations), n=7:

U.S.: CALIFORNIA! LOS ANGELES Co.: Point Dume, W13, 170; Playa del Rey,
W273. oRANGE co.: Lower edge Cleveland National Forest, just above San Juan
Guard Station, R17490 (progeny =R18764, 19079). SAN DIEGO co.; 2.7 mi. from

386 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Fallbrook on road to De Luz, R17478, W207; Carlsbad, R14032; Switzers Canyon,
San Diego, W212; Balboa Park, San Diego, W859 (SDSC); Otay Lakes Road 1.8
mi. E. of Bonita Road, R16971.

BAJA CALIFORNIA: 2.1 mi. 8. of El Descanso, Klein & Gregory 1186; ca.
3 mi. 8. of Ensenada, Klein & Gregory 1273, R17011; 7.5 mi. N. of Santo Tomds,
R17017, 17050; 5.5 mi. S. of San Vicente, 950 ft., R17021; 1.1 mi. N. of Colonia
Guerrero, R17035 (type collection); 2 km. N. of Colonia Guerrero, R12135; San
Quintin, R12362; 3 mi. S. of San Quintin, Gregory 335 (RSA, US); 14 mi. S. of
San Quintin, R17030; 5 mi. NE. of El Rosario, Klein & Gregory 1266; ca. 13 mi.
SE. of El Rosario, Klein & Gregory 1249.

Camissonia luciae Raven (7 individuals, 7 populations), n=21:

U.S.: CALIFORNIA: MONTEREY CO., SANTA LUCIA MTS.: Road to Jamesburg 0.8
mi. from main road up Carmel Valley, R18150; shale roadbank 7.8 mi. SW. of
Arroyo Seco Guard Station on road to Jolon, B2299; Hastings Reservation, near
Jamesburg, R18221; The Indians, R18267 (type collection). SAN BENITO co.: 3.1
mi. W, of Fresno Co. line on road from Coalinga to Hernandez, Lewis et al. 1342-1.
SANTA BARBARA CO., SAN RAFAEL MTs.: 1.3 mi. N. of Cachuma Guard Station,
Lewis 1678 (LA); Davy Brown Campground, Lewis 1647 (LA).

Camissonia micrantha (Hornem. ex Spreng.) Raven (47 individuals, 39 popula-
tions), n=7:

U.S.: CaLirorNIA: CONTRA costa co.: Antioch, R18141; 1 mi. S. of Oakley,
R18143. Los ANGELES co.: Claremont, W271 (SDSC), R17120, 17121; Point
Dume, W13 (SDSC), 171. MonrEREY co.: Hastings Reservation, R18224; Big
Sandy Valley, R18283; 3 min. N. of summit of Hog Canyon Road, S. of Parkfield,
R18299; Bradley, R18307. RIveRSIDE co.: 11 mi. W. of Riverside, Snow & Mos-
quin in 1957 (LA). SAN BENITO Co.: Pinnacles National Monument, B2183, Lewis
1340; 1.1 mi. S. of Cienaga School, R18198; 5.1 mi. N. of Cienaga School, R18196.
SAN BERNARDINO Co.: Alta Loma, R16987. san piEGo.: U.S. Hwy. 395, 2 mi.
S. of turn to Lilac, R16950, 17457. saN FRANCISCO Co.: Lake Merced, R18160.
SAN LUIS OBISPO Co.: Near mouth of Arroyo de la Cruz, Hardham in 1963; 2 mi
W. of Nipomo, B4269; 1.2 mi. N. of Simmler-Creston road on road to Shandon,
R16972, 18321; near Atascadero, R18277; 1.8 mi. W. of Bee Rock, R18034; 2.9
mi. E. of San Miguel, R18310; 1.2 mi. E. of Santa Margarita, R18312, 3.8 mi. E.,
B2026, 3 mi. E., R18314, 3.7 mi. NE., R18316, 4.2 mi. NE., R18317, 18.6 mi.
NE., R18319, 18320; Nipomo Valley 1.5 mi. N. of Santa Barbara Co. line, R16955;
S. road to Adelaida 6.4 mi. W. of junction with Oak Flat Road, near Paso Robles,
Lewis & Thompson 1100 (LA). SANTA BARBARA CoO.: 1 mi. W. of Goleta, B1967;
Jalama Beach, R14039; 4 mi. N. of Lompoc, R15510; 1.7 mi. N. of Santa Ynez R.
on State Hwy. 1, B2556; San Miguel I., R18178; Cherry Canyon, Santa Rosa I.,
R14962. santa cruz co.: Sunset Beach, R18187; near Ben Lomond, R18183,
B4637. sonoma co.: Bodega Bay, R18245.

CULTIVATED: Seeds from Copenhagen Botanic Garden as Ocenothera hirta,
R18242.

Camissonia pallida (Abrams) Raven subsp. hallii (Davids.) Raven (4 individuals,
3 populations), n=7:

U.S.: CALIFORNIA: RIVERSIDE co.: Jumbo Rocks Campground, Joshua Tree
National Monument, R17116; Whitewater Canyon 2.7 mi. from U.S. Hwy. 70-99,
R17109; Morongo Canyon, Lewis 1682 (LA, RSA); Morongo Canyon, ca. 8 mi.
from U.S. Hwy. 60-70-999, W231 (SDSC).

RAVEN—GENUS CAMISSONIA 387

Camissonia pallida subsp. pallida (26 individuals, 24 populations), n=7:

U.S.: ARIZONA: MOHAVE Co.: 9 mi. N. of Kingman, 2,950 ft., R17359. YAVAPAI
co.: 5.4 mi. NE. of Congress, R18367 (progeny = R18767, 19752).

U.S.: CALIFORNIA: KERN co.: Kern R. 6 mi. S. of Roads End P.O., 3,100 ft.,
Howell 33144 (progeny = R19081, 19747); W. of the mouth of Kern R. Canyon,
Lewis 1670 (LA, RSA). Los ANGELES co.: Big Rock Creek, San Gabriel Mts.
R11961; 0.5 mi. N. of Pearblossom, 2,800 ft., R17104; 0.2 mi. E. of Pearblossom,
Lewis & Wedberg 1692 (LA); 1 mi. W. of Littlerock, R13995. RIVERSIDE Co.:
Whitewater Canyon, W in 1957 (LA); near San Bernardino Co. line in Morongo
Canyon, W232; Pinyon Crest Road 2.5 mi. from Palms-to-Pines Hwy., Theobald
62. SAN BERNARDINO co.: Cushenbury Grade 0.5 mi. below Cactus Flat, San
Bernardino Mts., 5,900 ft., R17461; Bonanza King Mine, Providence Mts.,
R17358 (progeny = R18769, 19753). SAN DIEGO Co.: 8.7 mi. S. of Scissors Crossing
on road to Vallecito Valley, R16902, 9.1 mi. SE., R11536, 6.8 mi. 8., R16900;
San Felipe Creek wash 2 mi. E. of road to Yaqui Pass, R16895, 2.2 mi. W. of road
W219 (SDSC), in 1957 (LA); Vallecito Valley just outside Anza-Borrego State
Park boundary, R16904; N. end of Vallecito Valley, W222; 0.8 mi. E. of Banner
Queen Store, Banner Grade, R16923; 2.5 mi. inside Anza-Borrego State Park on
road from Scissors Crossing, W221; 3 mi. from Banner on road to Borrego Valley,
W216 (SDSC); Ensign Ranch, Borrego Valley, H.& M. Lewis 1637A (LA),
1637B (LA).

Camissonia proavita Raven (4 individuals, 4 populations), n=7:

BAJA CALIFORNIA: 5.5 mi. S. of San Vicente, R17020 (type collection) ;
19.5 km. SE. of San Vicente, R12264; 1.2 mi. N. of Colonia Guerrero, R17034;
San Quintin, R12363.

Camissonia robusta Raven (29 individuals, 24 populations), n=21:

U.S.: CALIFORNIA: LOS ANGELES CO.: SANTA CATALINA I.: Haypress, R17807
(progeny = 18170). SAN CLEMENTE 1.: Piehl 62387 (progeny = R18174). SAN DIEGO
co.: U.S. Hwy. 395, 2 mi. S. of turn to Lilac, R17458 (progeny = 18846) ; 4.8 mi. W.
of Pala, R17061; 1.6 mi. E. of Pala, R16921, 17062; 4.6 mi. E. of San Pasqual,
R17407; 2.3 mi. E. of Ramona, R17411; 0.38 mi. from U.S. Hwy. 395 on Deer
Springs Road to San Marcos, R17474; 0.7 mi. N. of Felecita County Park, R17458 ;
Balboa Park, San Diego, W214 (SDSC); Sweetwater Road, San Diego, W860
(SDSC); Descanso to Lyons Valley road 3.2 mi. NE. of turn to Camp Barrett,
R17431; 2 mi. E. of Chula Vista on road to Bonita, B1809; Barrett, R16937
(3 plants). SANTA BARBARA CO., SANTA CRUZ 1.: Top of Ridge between Coches
Prietos Canyon and Alberts Anchorage, 400 ft., Blakley 3314 (progeny = R18756,
19077).

BAJA CALIFORNIA: Above La Misién, between Tijuana and Ensenada,
R17056; 10 mi. 8. of Ensenada, R17054; ca. 16 mi. 8. of Ensenada, Klein & Gregory
1195; 10.9 mi. N. of Santo Tomds, R17012; 7.5 mi. N. of Santo Tomas, R17016,
17051A; 5.5 mi. 8. of San Vicente, R17022 (2 plants) ; 7.4 mi. N. of Ejido México,
500 ft., R17043. GuapALUPE 1.: NE. side of Barracks Cove Canyon, Copp 151
(progeny = R18759); Northeast Anchorage, Wiggins & Ernst 222 (progeny =
R18754, 19078).

Index

(Synonyms and page numbers of figures in 2talzcs. Page numbers of principal
entries in boldface.)

Ayassizia, 179, 253
Agassizia cheiranthifolia, 179, 265
Andrena, 323
subg. Diandrena, 270
subg. Onagrandrena, 266, 270, 310,
327
chalybea, 242
eulobi, 190
mojavensis, 310
oenotherae, 261
parachalybea, 261
raveni, 247
vespertina, 352
Anthophora urbana, 231
Boisduvalia, 341
Boisduvalia cleistogama, 351
Burragea, 161
Calylophus, 161, 162, 163, 356
Calylophus hartwegii, 161
Camissonia
Sect. Camissonia, 165, 169, 171,
172, 173, 174, 176, 176, 177, 181,
186, 201, 305, 313, 314, 319, 322,
324, 328, 330, 331, 332, 339, 344,
351, 376
Sect. Chylismia, 168, 169, 171, 172,
173, 174, 175, 176, 177, 180, 181,
182, 202, 233, 234, 307, 310, 351,
358, 375, 382
Sect. Chylismiella, 168, 169, 172,
173, 174, 175, 180, 181, 189, 374
Sect. Eremothera, 169, 171, 172,
173, 174, 175, 176, 177, 181, 188,
203, 350, 353, 355, 367, 382
Sect. Eulobus, 168, 169, 170, 171,
172, 175, 176, 177, 180, 181, 188,
189, 193, 195, 233, 234, 351
Sect. Holostigma, 165, 169, 171,
172, 173, 174, 176, 176, 177, 179,
181, 185, 189, 201, 253, 262, 270,
274, 278, 283, 284, 288, 296, 300,
307, 324, 340, 351, 376, 381, 382

Camissonia—Continued

Sect. Lignothera, 168, 169, 171, 172,
174, 175, 180, 181, 184, 233

Sect. Nematocaulis, 169, 172, 173,
174, 175, 176, 181, 189, 203, 375,
379

Sect. Sphaerostigma, 305

Sect. Tetranthera, 202, 203

Sect. Tetrapteron, 169, 171, 172,
173, 174, 175, 176, 180, 181, 184,
237, 239, 240, 241, 246, 252

acuminata, 342

andina, 168, 169, 174, 189, 203, 229,
373, 376, 377, 378

angelorum, 182, 190, 192, 193, 196,
196, 200

arenaria, 184, 228, 233, 236

benitensis, 188, 306, 324, 328, 331,
332, 338, 341

bistorta, 174, 185, 254, 255, 262,
263, 264, 267, 268, 276, 277, 281,
286, 291, 292, 298, 301, 305, 381,
382

boothii, 172, 351, 352, 357, 359

boothii subsp. alyssoides, 166, 189,
352, 358, 359, 361, 362, 364, 365,
366, 371

boothii subsp. boothii,
359, 365, 366, 368

boothii subsp. condensata, 188, 354,
357, 369, 361, 362

boothii subsp. decorticans, 188, 351,
358, 359, 360, 361

boothii subsp. desertorum, 166, 188,
354, 357, 359, 360, 361, 362, 363,
364, 366

boothii subsp. intermedia, 188, 359,
363, 364, 365, 366

breviflora, 168, 169, 173, 174, 180,
184, 237, 238, 239, 241, 244, 246,
247, 248, 249

188, 352,

389

390 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Camissonia—Continued Camissonia—Continued

brevipes, 184, 202, 204, 209, 213,
219

brevipes subsp. arizonica, 205, 206,
207, 220, 222

brevipes subsp. brevipes, 178, 205,
206, 209, 211, 216, 217, 218, 222,
232

brevipes subsp. pallidula, 205, 206,
209, 211, 218, 219, 222

californica, 169, 171, 176, 180, 182,
189, 190, 196, 197, 199, 200, 271

campestris, 306, 307, 309, 310, 312,
320, 321, 323, 324, 326, 327, 334,
338, 340, 341, 345

campestris subsp. campestris, 187,
307, 321, 322, 325, 326, 338, 346,
347, 350

campestris subsp. obispoensis, 187,
322, 325, 338, 350

cardiophylla, 233, 234, 236

cardiophylla subsp. cardiophylla,
184, 234, 237
cardiophylla subsp. cedrosensis,

168, 184, 236

cardiophylla subsp. robusta, 184,
235

chamanerioides, 169, 174, 188, 203,
297, 351, 352, 353, 354, 356

cheiranthifolia, 171, 179, 254, 255,
258, 260, 281, 339, 381

cheiranthifolia subsp. cheiranthi-
folia, 172, 185, 259, 260, 262,
265, 270, 274, 281, 382

cheiranthifolia subsp. suffruticosa,
172, 185, 258, 259, 260, 262, 266,
267, 270, 277, 279, 382

claviformis, 173, 174, 202, 205, 212,
213

claviformis subsp. aurantiaca, 183,
205, 207, 209, 213, 216, 217, 218,
219, 220, 222

claviformis subsp. citrina, 213

claviformis subsp. claviformis, 183,
205, 213, 215, 216, 218, 226, 231

claviformis subsp. cruciformis, 183,
212, 213, 226

claviformis subsp. funerea, 183, 205,
213, 216, 217, 218, 222, 232

claviformis subsp. integrior, 183,
205, 2138, 214, 215, 218, 226, 232

claviformis subsp. lancifolia, 183,
213, 214, 232

claviformis subsp. peeblesii, 169,
175, 183, 205, 207, 213, 218, 220,
222

claviformis subsp. peirsonii, 183,
213, 218, 219, 220, 221

claviformis subsp. rubescens, 183,
207, 213, 219, 221

claviformis subsp. wigginsii, 169,
183, 213, 221

claviformis subsp. yumae, 183, 207,
213, 218, 220

confertiflora, 169, 172, 175, 184,
202, 203, 204

confusa, 186, 255, 271, 281, 286,
287, 290, 291, 294, 298, 299, 300,
381, 383

contorta, 168, 187, 306, 307, 310,
316, 318, 320, 321, 323, 324, 325,
326, 327, 332, 333, 334, 335, 336,
337, 338, 341, 345, 347, 350

contorta var. pubens, 316

crassifolia, 170, 171, 181, 190, 194,
195, 196, 200

dentata, 164, 169, 175, 179, 180,
306, 308, 321, 324, 337, 338, 339

dentata subsp. dentata, 175, 188,
341, 343

dentata subsp. littoralis, 188, 340,
341, 343, 344

eastwoodiae, 169, 172, 182, 202,
213, 223, 225, 226

exilis, 169, 182, 202, 203, 229, 230,
373, 377

flava, 180, 341

gouldii, 175, 189, 351, 352, 355,
367, 368

graciliflora, 185, 230, 237, 238, 239,
242, 250, 242, 253, 351

guadalupensis, 256, 257, 271, 273,
281, 305

guadalupensis subsp. clementina,
174, 185, 255, 267, 272, 273, 274,
275, 283

guadalupensis subsp. guadalupen-
sis, 169, 185, 272, 273, 274, 275,
383

hallii, 292

hardhamiae, 185, 255, 281, 286,
296, 300, 301, 383

heterochroma, 174, 182, 211, 215,
216, 231

INDEX 391

Camissonia—Continued

hilgardii, 169, 174, 189, 370, 376,
377, 379

hirtella, 186, 255, 257, 271, 279,
281, 282, 283, 284, 286, 290, 291,
294, 297, 298, 301, 302, 303, 305,
309, 324, 383

ignota, 186, 255, 257, 271, 277,
279, 286, 287, 288, 289, 294, 297,
298, 300, 301, 384

integrifolia, 187, 306, 308, 310, 318,
324, 330, 331, 334, 338, 341, 344,
350

intermedia, 185, 186, 255, 271, 279,
281, 286, 287, 291, 295, 296, 297,
301, 302, 303, 305, 385

kernensis, 306, 307, 308, 314

kernensis subsp. gilmanii, 187, 287,
289, 307, 310, 321, 324

kernensis subsp. kernensis, 174,
187, 287, 289, 307, 309, 310, 311,
312, 318, 324, 338, 345, 350

lacustris, 187, 306, 321, 323, 324,
329, 330, 331, 333, 338, 341, 350

lewisii, 174, 185, 255, 257, 263, 268,
270, 271, 275, 279, 281, 291, 298,
303, 305, 385

luciae, 185, 255, 281, 286, 296, 300,
302, 286

megalantha, 169, 174, 182, 229, 371

micrantha, 186, 255, 256, 257, 267,
271, 274, 275, 277, 278, 279, 282,
286, 287, 291, 295, 297, 298, 301,
302, 303, 304, 305, 386

minor, 169, 188, 203, 229, 351, 352,
371, 372, 374, 377

multijuga, 171, 174, 184, 202, 204,
205, 207, 210, 211, 218, 228

munzil, 182, 205, 207, 213, 216, 218,
222, 232

nevadensis, 169, 172, 188, 351, 352,
355, 369, 370

ovata, 173, 185, 237, 238, 239, 240,
242, 251

pallida, 255, 256, 257, 271, 291

pallida subsp. hallii, 186, 270, 288,
292, 294, 386

pallida subsp. pallida, 186, 284, 286,
287, 288, 290, 291, 292, 293, 301,
387

palmeri, 169, 185, 237, 238, 239,
251, 252, 253, 351

paradoza, 380

Camissonia—Continued

paradoza var. divaricata, 342

parryi, 169, 172, 182, 203, 209, 227,
228, 369

parvula, 169, 187, 307, 308, 313,
314, 315, 316, 318, 350

proavita, 169, 186, 255, 257, 274,
277, 278, 281, 291, 297, 298, 387

pterosperma, 173, 174, 189, 372,
374, 375

pubens, 166, 174, 187, 307, 308, 310,
313, 314, 316, 319, 338, 345, 350

pusilla, 187, 307, 308, 312, 313, 514,
316, 318, 319, 320

pygmaea, 169, 189, 351, 352, 346,
366, 367, 368

refracta, 169, 174, 176, 188, 297,
351, 352, 355, 3465, 356, 357, 358

robusta, 185, 255, 271, 277, 279,
281, 283, 286, 298, 300, 304, 387

scapoidea, 169, 180, 202, 203, 223

scapoidea subsp. brachycarpa, 182,
214, 216, 226

scapoidea subsp. macrocarpa, 169,
175, 182, 225

scapoidea subsp. scapoidea, 169,
176, 182, 212, 223, 224, 226

scapoidea subsp. utahensis, 175,
182, 226

sceptrostigma, 181, 190, 193, 196,
196

sierrae, 306, 323, 326, 350

sierrae subsp. alticola, 187, 322,
328, 329

sierrae subsp. sierrae, 187, 322, 327,
328, 329

speculicola, 209, 212

speculicola subsp. hesperia, 169,
175, 184, 210

speculicola subsp. speculicola, 169,
184, 209, 210

strigulosa, 169, 188, 306, 308, 310,
317, 318, 324, 325, 326, 329, 331,
332, 333, 535, 338, 339, 341,
344, 345, 346, 347, 349, 350

subacaulis, 169, 172, 173, 185, 237,
238, 239, 240, 242, 248, 250

tanacetifolia, 172, 174, 176, 237,
238, 244, 249, 250

tanacetifolia subsp. quadriperfor-
ata, 184, 246, 248

tanacetifolia subsp. tanacetifolia,
184, 239, 241, 244, 245, 246, 248

392

Camissonia—Continued
tenutfolia, 342
walkeri, 176, 182, 184, 203, 205,
210, 211
walkeri subsp. tortilis, 174, 205,
207, 208, 211, 212
walkeri subsp. walkeri,
210, 211, 212, 222, 225
Centris rhodopus, 231
Chamissonia—see Camissonia
Chylisma—see Chylismia
Chylismia, 164, 180, 202
arenaria, 236
aurantiaca, 218
brevipes, 205
cardiophylla, 234
clavaeformis, 217
clavaeformis var. cruciformis, 213
clavaeformis var. typica, 217
cruciformis, 213
heterochroma, 231
hirta, 207
lancifolia, 214
multtjuga, 207
parryt, 227
parviflora, 207
peirsonit, 219
pterosperma, 375
scapotdea, 224
scapoidea var. aurantiaca, 217
scapoidea var. seorsa, 224
tenuissima, 227
venosa, 207
walkeri, 211
Clarkia, 161, 162, 163, 175, 231, 341,
344, 373, 377, 382
tenella, 344
Cruciferae, 200, 205
Epilobieae, 351
Eulobus, 164, 165, 189
californicus, 180, 197
Gaura, 161, 162, 163, 233, 356, 376
angustifolia, 376
decorticans, 358
hexandra subsp. hexandra, 376
mutabilis, 233
Gayophytum, 161, 163, 170, 173, 174,
200, 341, 356, 375, 380
humile, 173
micranthum, 380
Godetia tenuifolia, 344

169, 209,

CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Gongylocarpus, 161, 163, 164, 170, 231
fruticulosus, 170
rubricaulis, 170
Hauya, 161, 162, 163, 164
Heterogaura, 161, 162, 163, 231
Holostigma
alyssoides, 362
argutum, 341
bottae, 264
cheiranthi folium, 265
heterophyllum, 341
micranthum, 279
paradozum, 380
spirale, 265
tenuifolium, 341
Jussiaea subacaulis, 242
Ludwigia, 376, 377
Sect. Prieurea, 376
inclinata, 377
verticillata, 377
Oenothera, 161, 162, 163, 164, 165, 231,
356
subg. Chylismia, 162, 165, 180, 202,
233
subg. Heterostemon, 237
subg. Primulopsis, 237
subg. Sphaerostigma, 235, 305, 350,
375
subg.
Sect.
Sect.
Sect.
Sect.
Sect.
Sect.
Sect.
Sect.
Sect.
Sect.

Taroxia, 180, 237
Chylismia, 202, 374
Chylismiella, 374
Eremothera, 350
Euchylismia, 202, 233
Eutaraxia, 237
Heterostemon, 237
Holostigma, 253
Lignothera, 233
Nematocaulis, 375
Sphaerostigma, 179, 305
Sect. Tetranthera, 202

Sect. T'etrapteron, 237
abramsi, 293

alyssoides, 352, 362

alyssoides var. decorticans, 358
alyssoides var. minutiflora, 371
alyssoides var. typica, 363
alyssoides var. villosa, 362, 364
andina, 378

andina var. anomala, 376, 378
andina var. hilgardit, 377
andina f. tripetala, 376, 378
andina var. typica, 378
angelorum, 192

INDEX 393

Oenothera—Continued

arenaria, 236

bistorta, 256, 264, 267

bistorta var. hallii, 271, 292

bistorta var. typica, 267

bistorta var. veitchiana, 267, 271

boothti, 352, 365

boothit subsp. alyssotdes, 363

boothit subsp. boothit, 365

boothii subsp. condensata, 362

boothit subsp. decorticans, 358

boothiz subsp. desertorum, 360

boothii subsp. intermedia, 364

boothit subsp. inyoensis, 360, 361

boothit var. pygmaea, 366

boothit subsp. rutila, 358, 360

boothit var. typica, 364, 365

breviflora, 180, 237, 249

brevipes, 205

brevipes subsp. arizonica, 207

brevipes subsp. brevipes, 205

brevipes var. multijuga, 207

brevipes subsp. pallidula, 206

brevipes var. pallidula, 206

brevipes race parryi, 227

brevipes var. parviflora, 207

brevipes var. scapoidea, 224

brevipes var. typica, 205

californica, 197

campestris, 321

campestris subsp. campestris, 321,
329

campestris var. cruciata, 345

campestris subsp. parishii, 321

cardiophylla, 234

cardiophylla subsp. cardiophylla, 234

cardiophylla subsp. cedrosensis, 236

cardiophylla var. longituba, 236

cardiophylla var. petiolaris, 234

cardiophylla subsp. robusta, 235

cardiophylla var. splendens, 236

cardiophylla var. typica, 234

cedrosensi:, 236

chamaenerioides var. torta, 371

chamissonis, 341

chetranthifolia, 265

cheiranthifolia var. contorta, 345

cheiranthifolia f. delicatula, 267

cheiranthifolia var. nitida, 167, 265

chetranthifolia var. suffruticosa, 259

chetranthifolia var. typica, 265

chilensis, 342

clavaeformis, 216

Oenothera—C ontinued

clavaeformis subsp. aurantiaca, 218

clavaeformis var. aurantiaca, 175,
216, 218

clavaeformis subsp. citrina, 213

clavaeformis subsp. clavaeformis, 217

clavaeformis subsp. cruciformis, 213

clavaeformis var. cruciformis, 213,
214

clavaeformis subsp. funerea, 216

clavaeformis subsp. tntegrior, 215

clavaeformis subsp. lancifolia, 214

clavaeformis var. peeblesiz, 218

clavaeformis subsp. peirsoni1, 219

clavaeformis var. peirsonit, 219

clavaeformis var. purpurascens, 213

clavaeformis subsp. rubescens, 221

clavaeformis var. typica, 215, 217

clavaeformis subsp. wigginsit, 221

clavaeformis subsp. yumae, 220

confertiflora, 203

contorta, 166, 307, 339, 345

contorta var. contorta, 306, 346, 350

contorta var. divaricata, 339, 340,
342

contorta var. epilobioides, 306, 307,
329, 333, 339, 340, 342, 346, 350

contorta var. flexuosa, 307, 312, 318

contorta var. pubens, 307

contorta var. strigulosa, 306, 307,
333, 346, 350

contorta var. typica, 306, 307, 345,
350

crassifolia, 190

crassifolia var. leptocarpa, 197

crassiuscula, 192, 196

cruciata, 321, 323, 329, 332, 346, 350

cructiformis, 213

decorticans, 352, 358

decorticans var. condensata, 362

decorticans var. desertorum, 360

decorticans var. intermedia, 352

decorticans var. inyoensis, 352

decorticans var. rutila, 352, 358, 361

decorticans var. typica, 358

dentata, 179, 306, 307, 321, 332, 333,
339, 340, 341, 346, 350

dentata var. campestris, 321, 329,
339, 340, 342

dentata var. crucitata, 345

dentata var. dentata, 339

dentata var. gilmanit, 310

dentata var. johnstonii, 307, 310, 321

394

Oenothera— Continued

dentata var. parishii, 321

deserti, 352

divaricata, 205

eastwoodiae, 222

erythra, 356

eulobus, 197

exilis, 202, 229

gauraeflora, 358

gauraeflora race boothit, 365

gauraeflora var. caput-medusae, 369

gauraeflora var. hitchcockti, 363

gauraeflora var. vermiculata, 369

graciliflora, 250

gracilis, 165

guadalupensis, 272

guadalupensis subsp. clementina,
275

guadalupensis subsp. guadalupensis,
272

hallit, 292

heterantha, 237, 242

heterantha var. tarazacifolia, 243

heterochroma, 231

heterochroma subsp. heterochroma,
232

heterochroma var. megalantha, 229

heterochroma subsp. monoensis, 232

heterochroma var. monoensis, 232

heterochroma var. typica, 231

heterophylla, 267, 342

hilgardi, 377

hirta, 257, 279

hirta var. exfoliata, 271, 293

hirta var. hirta, 281

hirta var. ignota, 287, 290, 298

hirta var. jonesiz, 282, 295, 298, 302

hirta var. jonesii f. reediz, 282

hirta var. typica, 275, 277, 279, 295,
298

hirtella, 256, 282

hyssoptfolia, 344

tgnota, 287, 298

kernensis, 309

kernensis subsp. gilmanit, 310

kernensis subsp. kernensis, 309

kernensits subsp. mojavensis, 310,
311

leptocarpa, 197

megalantha, 229

micrantha, 256, 257, 275, 277, 279,
295, 302, 304, 305, 380

micrantha var. abramsit, 293

CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Oenothera—Continued

micrantha var. acuminata, 339, 340,
342

micrantha var. exfoliata, 293

micrantha var. hirtella, 282

micrantha var. ignota, 287, 290, 381

micrantha var. jonesii, 282, 286

micrantha var. reedit, 282

minor, 371

minor var. cusickit, 371, 373

minor var. typica, 371

minutiflora, 342

multyjuga, 207

multijuga var. orientalis, 211

multijuga var. parviflora, 207, 211

multijuga var. typica, 207

munzi, 222

nevadensis, 369

nitida, 265, 266, 382

nuttallat, 245

ovata, 238

pallida, 256

pallidula, 206

palmeri, 252

parryi, 227

parryt f. tenuissima, 227

parviflora, 207

parvula, 318

phlebophylla, 207

primuloidea, 238, 243

pterosperma, 375

pubens, 316

pygmaea, 366

rutila, 358, 360

scapoidea, 180, 224

scapoidea var. aurantiaca, 217

scapoidea subsp. brachycarpa, 226

scapoidea var. clavaeformis, 216

scapoidea var. eastwoodae, 222

scapoidea subsp. macrocarpa, 225

scapoidea var. parryi, 227

scapoidea var. purpurascens, 215

scapoidea subsp. scapoidea, 224

scapoidea var. seorsa, 224

scapoidea var. tortilis, 211

scapoidea var. typica, 224

scapotdea subsp. utahensts, 226

sceptrostigma, 196

speculicola subsp. hesperia, 210

speculicola subsp. speculicola, 209

sptralis, 265

sptralis f. arcuata, 257

spiralis f. clypeata, 257

INDEX

Oenothera—Continued
spiralis var. linearis, 259
spiralis var. nitida, 265
spiralis f. viridescens, 259
spiralis var. viridescens, 259
strigulosa, 333
strigulosa var. epilobioides, 333
subacaulis, 243
subacaulis var. taraxacifolia, 243
tanacetifolia, 245
tenuifolia, 341
tenuissima, 227, 228
torulosa, 308, 344
torulosa f. chilensis, 342
torulosa f. cruciata, 345
torulosa race helianthemiflora, 321
torulosa f. mixta, 345
torulosa f. permizta, 321
utahensis, 363
virescens, 259
viridescens, 259
walkeri, 211
walkeri subsp. tortilis, 211
watsonit, 207
Onagraceae, 161, 277, 341, 351, 377, 382
Onagreae, 161, 162, 163, 164, 170, 175,
231, 233, 247, 373, 375, 382
Onothera—See Oencthera
Polygonaceae, 165
Salpingia, 162
Sphaerostigma, 164, 179, 264
acuminatum, 342
alyssoides, 362
alyssoides var. macrophyllum, 363
alyssoides var. minutiflorum, 371
andinum, 378
andinum var. hilgardii, 377
andinum var. minutum, 380
angelorum, 192
arenicola, 256, 282
bistortum, 267, 271
bistortum var. reedi1, 282
bistortum var. veitchianum, 267
boothii, 365
boothii var. pygmaeum, 366
campestre, 321
campestre var. helianthemiflorum, 321
campestre var. minus (‘minor’), 345
campestre var. miztum, 345
campestre var. parishii, 321
chamaenerioides, 356
chamissonis, 341
cheiranthifolium, 265

395

Polygonaceae—Continued

contortum, 345

contortum var. fleruosum, 318
contortum var. greenei, 333
contortum var. pubens, 316
decorticans, 358

dentata subsp. campestris, 321
dentatum, 307, 342

deserti, 352

divaricatum, 342

erythrum, 356

filuforme, 318

fleruosum, 318
gauraeflorum, 358

hallti, 256, 292
heterophyllum, 342

hilgardi, 377

hirtellum, 256, 282
hirtellum var. montanum, 282
hirtum, 279

hitchcockii, 363

umplexum, 363

lemmont, 365
macrophyllum, 363
micranthum, 256, 279
micranthum var. exfoliatum, 293
micranthum var. jonesti, 282
minus (‘‘minor’’), 371
nelsoni, 371

nevadense, 369

nitidum, 265

orthocarpum, 316

pallidum, 256, 293
paradozum, 380

parvulum, 318
pterospermum, 375

pubens, 316

ramosissimum, 242
refractum, 352

rutilum, 358

senex, 365

spirale, 265, 381

spirale var. clypeatum, 258
spirale var. viridescens, 259
strigulosum, 333, 334
tenutfolium, 342, 343
tortum, 371

tortum var. eastwoodae, 371
tortuosa, 369

utahense, 363

veitchianum, 267, 381
viridescens, 259

396 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Stenosiphon, 161, 162, 163 Tarazia— Continued

Tarazia, 164, 180, 237 palmeri, 252
breviflora, 249 subacaulis, 243
graciliflora, 250 tanacetifolia, 245
heterantha, 243 taraxacifolia, 243
heterantha var. taraxactfolia, 243 tikurana, 245
longiflora, 245 Xylonagra, 161, 163, 170
ovata, 238

U.S. GOVERNMENT PRINTING OFFICE : 1968—-O-295-655

BULLETIN OF THE UNITED STATES NATIONAL MUSEUM

CONTRIBUTIONS FROM THE Unitep States NATIONAL HERBARIUM

VotumeE 37, Part 6 (Enp oF Votume)

BREDIN-ARCHBOLD-SMITHSONIAN
BIOLOGICAL SURVEY OF DOMINICA

Myxomycetes from Dominica

By Marie L. Farr

SMITHSONIAN INSTITUTION PRESS ¢ CITY OF WASHINGTON e 1969

BREDIN-ARCHBOLD-SMITHSONIAN
BIOLOGICAL SURVEY OF DOMINICA

Myxomycetes from Dominica !

By Marie L. Farr?

Introduction

This paper is based mostly on the collections made during my
participation in the Bredin-Archbold-Smithsonian Biological Survey
of Dominica (British West Indies). Species of myxomycetes pre-
viously reported from this island but not found on this expedition
are also included to bring the record up to date. The present report
although based on a very short collecting period and, therefore, far
from complete, evidently represents the first extensive investigation
of the slime molds of Dominica.

From January through March 1966, over 500 myxomycete speci-
mens were collected in the field and 63 were obtained from 187 moist
chamber cultures. These efforts yielded 96 taxa (1 doubtful) of which
2, 1 of Diachea and 1 of Physarum, are described here as new species.

Dominica, the largest of the Windward Islands and third largest
of the Lesser Antilles, is located at 15°10’-15°40’ N. latitude and
61°44’-61°30’ W. longitude, about halfway between Guadeloupe
and Martinique, and has an area of slightly more than 300 square
miles; its maximum length is ca. 29 miles and its greatest width, ca.
15 miles. The highest peaks are Morne Diablotins (ca. 4,700 ft.),
Morne Trois Pitons (ca. 4,500 ft.), Watt Mountain (ca. 4,000 ft.),
and Morne Micotrin (ca. 3,900 ft.). Because of its relatively recent
volcanic origin, the island is characterized by a much craggier topog-
raphy and a (consequently) wetter climate than other major Caribbean
Islands. The difficulties in road building, timber harvesting, etc.,
caused by the rugged terrain; the heavy downpours; and the strong

1 This is the first in a series of floristic studies. Other floral papers in this series
will appear in ‘Smithsonian Contributions to Botany.’’ A companion series on
the fauna appears in the ‘Proceedings of the United States National Museum’’
and ‘Smithsonian Contributions to Zoology.”

2 Mycologist, Crops Research Division, Agricultural Research Service, U.S.
Department of Agriculture, Beltsville, Maryland 20705.

397

398 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

winds have retarded the advancement of lumbering, agriculture
(less than half of Dominica was under cultivation in 1963), and
tourism, and have resulted in the preservation of relatively large
tracts of the typical Caribbean forests that have largely disappeared
from the more highly developed islands. The last decade or so, how-
ever, has witnessed an acceleration in road construction and the
erection of an airport, which, among other factors, are greatly hasten-
ing the conversion of virgin lands to crop cultivation and the develop-
ment of tourist trade and facilities.

Unpublished climate maps * show that only a very narrow coastal
fringe (mostly less than 2 miles wide) receives less than 100 inches
of yearly rainfall and experiences a true dry season; most of the
mountainous interior lies in the zone characterized by 200-300 inches,
or even more. The relative humidity is always high (50-80% in
Roseau, 85% or more in the interior) and a considerable amount of
precipitation occurs even during the drier parts of the year. The
temperature range is ca. 65-90° F., averaging in the upper seventies.
According to Hodge (16), from whose extensive treatise much of the
present introductory information was taken, a dry season extends
from mid-January to mid-June, April being the driest and calmest
month. The wet season consists of a summer portion (mid-June to
mid-September), a “break” of “complete sunny days with occasional
heavy showers only at night” (mid-September to mid-November),
and a winter portion (mid-November to mid-January). Fentem (12)
mentions an “undependable and indefinite drought between mid-
January and mid-June on the leeward coast’ which generally lasts
about 6 weeks” and a tendency toward a second period of minimum
rainfall in September and October.

In view of the generally high precipitation and humidity one would
imagine the dry season to be the best time on most parts of the
island for gathering myxomycetes. Enough moisture is present, even
then, for their development, yet fructifications can ripen completely
without being washed away immediately by intense downpours or
rapidly disfigured by filamentous molds. The validity of this assump-
tion remains to be tested by collecting in Dominica during the wet
season. Certain areas subjected to pronounced droughts, however,
may be expected to produce myxomycetes only after the onset of
rainy periods. In Jamaica, for example, certain pastured and wooded
tracts exhibiting distinct seasonal changes in verdure were myxomyco-
logically barren during the dry season, but very fruitful during or
immediately following the rainy season (9).

Most of my time on Dominica supposedly coincided with the “dry”
season, but the weather was somewhat different from that anticipated.

3 Kindly supplied by the Lands & Surveys Department of Dominica.

FARR—MYXOMYCETES FROM DOMINICA 399

An unusually dry spell in December and January, which sometimes
necessitated a partial curtailment of electric power, but provided
very pleasant weather, was followed by an extremely wet February
and an only slightly drier March. February, especially, was marked
by nightly violent downpours changing into milder, more or less
intermittent rain of varying intensity nearly every day (at least,
inland). Although the nearly continuous rain impeded slime-mold
collecting considerably, my dampened spirits were revived by the
discovery of a number of myxomycetes fruiting in good condition,
even during the rain, in certain rather sheltered niches; the undersides
of very large logs, and substrates protected by “umbrellas” of over-
lapping banana leaves or buried within piles of leaf litter, yielded
collections of several species.

Topographic and climatic features of the island have given rise to a
diversity of soil types and microenvironments, resulting in a corre-
sponding distribution of vegetation types. Hodge (16) lists the fol-
lowing five primary plant formations, of which all but the first are
climatic: 1. Swamp forest, an edaphic climax occurring only in a few
small places on the island and represented in my collections by
locality no. 14. 2. Dry evergreen formation, including the littoral wood-
land vegetation of the windward (E.) coast. 3. Seasonal formations,
such as the thorny scrub vegetation along the dry leeward (W.) coast
and more or less evergreen forests in other areas exposed to periodic
drought; this formation has been largely replaced by crop cultivation.
4. Rain forest, the dominant, luxurious vegetation type of the wet
interior regions, much of which has become secondary or suboptimal
as a result of encroachment by shifting agriculture. Partially cleared
areas, in which considerable decay of stumps and felled trees has
already taken place, usually offer great promise for the collector.
The accumulation of rotten logs and other plant debris, the presence
of living crop plants (the most common substrate for some slime
molds), the more open, sunny situations created by removal of the
forest, and the surrounding border of undisturbed, shaded woodland
provide a variety of habitats and microenvironments and a corre-
sponding diversity of myxomycete species. This type of situation is
represented in my collections by localities 7-12, of which no. 8 yielded
the greatest number of species. The undisturbed rain forests of the
mountain slopes, on the other hand (except for locality no. 13), gen-
erally were found to be relatively unproductive despite the seemingly
ideal combination of high moisture, deep shade with sporadic, moderate
sunlight, windstillness, and abundance of rotten wood. (Rather
surprisingly, however, several myxomycetes developed in bark cul-
tures of the strongly resinous codominant tree, ‘““gommier’’ [Dacryodes
excelsa Vahl.].) The paucity of myxomycetes, which seems to prevail
in this type of habitat in other Caribbean islands as well, has puzzled

400 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

collectors for some time. Possibly the substrates are too acid to permit
spore germination or subsequent development of many species. The
density of the vegetation and the relative lack of air movement could
be expected to retard spore dispersal within the forest. Most wood-
inhabiting animals that might feed on myxomycetes were compara-
tively uncommon in the deep forest and can probably be ruled out as
cause of the sparse myxomycota. 5. Montane formations, found on
mountain summits and similar habitats exposed to strong, desiccating
winds, and consequently characterized by stunted vegetation types
such as “elfin woodland.” These are unproductive for field collecting
of slime molds. Two rather poor but interesting moist chamber devel-
opments (Physarum tleucopus and P. ?bubalinum) were obtained from
plant material collected at the top of Morne Diablotins. One would
surmise that great numbers of spores would be deposited in such an
open habitat, and that the cooler temperatures of the summits might
permit the survival of some species not normally thriving in the
tropics. Extensive collecting of substrate samples from this fo1ma-
tion, followed by moist chamber culture, would be a worthwhile
undertaking for energetic collectors possessing the endurance to make
repeated climbs to the summits and the stamina to collect after
reaching them.

Politically, Dominica is divided into 10 parishes of greatly differing
size; the largest ones are St. Joseph on the west side, St. David on the
east side, and St. Andrew to the north and northeast. Most myxomy-
cete collecting was carried out in these, as well as in St. Paul and
St. George, mainly because of accessibility. Although these parishes
proved highly fertile, some of the others may, upon thorough explora-
tion, turn out to be at least equally rich in myxomycetes.

Slime molds (including field collections and moist chamber fruit-
ings) were acquired from about three dozen localities, of which (to
save space) the 16 represented by the greatest number of field speci-
mens are coded by number and pinpointed on map 1. The first 10
constituted more or less ‘‘favorite” collecting spots because of their
combined accessibility and high yield (mostly more than 20 speci-
mens). Nos. 11-16, lower in yield, appeared very promising but, owing
to distance or inaccessibility, were less frequently and thoroughly
searched for myxomycetes. Nos. 1-3 are in St. Joseph; 11-12 in
St. Peter; 4 in St. John; 5, 13, and 14 in St. Andrew; 6, 8, 15, and 16
in St. David; 9 in St. George; 7 and 10 in St. Paul. They are briefly
described in the following paragraphs. The absence of road numbers
or names, and of accurately defined or permanent landmarks, accounts
for the sometimes awkward citations of locality.

4 On certain other islands, well-established footpaths to the mountain summits
mike this habitat more easily accessible than it is, at present, in Dominica.

FARR—-MYXOMYCETES FROM DOMINICA 401

\ 14

Cobrits ‘
Portsmouth 1
4 P
ST, JOHN
* $\ Melville Hall airport
\
ees ae ‘ st, ANDREW
‘= ‘
ier ‘ Pagua Boy
O. t ,
‘. 4
ww} .
ST, PETER AM, Dioblotins ‘sy 13
oe Naess 7
12 f yt y,
oO’ *s od
VW ee" \. ect!
a” \ /
©
ST. JOSEPH ‘
' NS
; °
ae f ST. DAVID
— 3
Le Ge __. ae
~e- -" le 7 4

> < et —— Cg

— x Pont iCassé
'

‘ *f Rosalie
7. _—- > M. Trois Pitons
AN

ST. PAUL ?

ery see
~

ee AM Micotrin we---
e 4 .

10 pom Yee woe
-- io ——
“re ST, GEORGE Jawan Mtn. 9
Roseav® vo
Y

S$ PATRICK
™. ‘
sek ‘

ST, LUKE *

Scotts Head 4

Map 1.—Dominica, B.W.I., showing the 16 most productive collecting localities for
myxomycetes.

1. Sr. Joseru: Clarke Hall Estate and vicinity (ca. half-mile
radius), along the Layou River (the largest river on the island),
extending E. approximately to the swinging footbridge and SE. to the
“Café” area, elev. 100-200 ft. This region, traversed by one of the
major roads, is almost entirely cultivated and most of the gatherings

402 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

were made in old, well-established cacao, banana, or coconut planta-
tions between highway and river, or along the roadside.

2. Sr. JoserH: From Clarke Hall W. to the highway bridge near
the mouth of the Layou River, elev. ca. 150 ft. to nearly sea level, an
area essentially similar to no. 1.

3. Sr. JosepH: Mannetts Gutter, a small creek (marked but not
labeled on current maps) ca. %-mile NE. of Clarke Hall, continuing
N. from the road passing Brookhill Estate, elev. ca. 200 ft. Most myxo-
mycetes were collected on large, more or less shaded, wet, rotten logs
and surrounding plants and debris near an ascending footpath along
the W. bank of the creek.

4. Sr. Jonn: S. of Portsmouth, near the mouth of the Indian River,
elevation near sea level, a dry area (at least when visited) on the lee-
ward (W.) coast. The species found, predominantly those tolerant of
some exposure (Lycogala epidendrum, the common Hemitrichia spp.,
Stemonitis webberi, etc.), were collected mostly on relatively dry, more
or less shaded logs in open areas.

5. St. ANDREW: Governor Estate, 3-4 miles W. of the airport, and
also beyond the end of the jeep road on a footpath ascending into the
disturbed rain forest, elev. ca. 400-600 ft. Myxomycetes were picked
up on the estate itself in low-lying plantations next to the Clyde River,
as well as along the forest path.

6. St. Davin: St. David Bay road, ca. 1.5 miles N. of its inter-
section with Rosalie road, near L’Arouma Popotte, elev. ca. 1,000 ft.
Slime molds were collected mostly in clearings and banana plantations
adjoining the highway, on rotting logs and in leaf litter.

7. St. Pau: 1-2 miles N. of Pont Cassé, in banana plantations
on the E. side of the highway leading to the airport, elev. 1,350-1,450
ft., habitat and substrates essentially similar to those of locality no. 6.
Collecting efforts on the W. side of the road were unrewarding, prob-
ably because those plantations were experimental plots for poisonous
chemicals.

8. St. Davip: About 100 yards E. of the intersection between the
roads to St. David Bay and Rosalie, along the latter, elev. ca. 1,400 ft.
This locality, reached by a short, narrow, steep footpath descending
from a roadside parking area, was characterized by an ideal diversity
of habitats, consisting of a dense, secondary rain forest encircling a
newly cut, sunny clearing next to a slightly older one planted to
banana trees and root crops; the entire cut area was lined with nu-
merous large logs (many of which were covered with moss, filmy ferns,
and various other epiphytes) and stumps from the original forest in
various stages of decay. Of all stations visited, this spot proved the
richest in number of species, but its productivity declined rather
sharply during the last month or so of my stay.

FARR—MYXOMYCETES FROM DOMINICA 403

9. St. Georce: Along the road between Laudat and Fresh Water
Lake, and on the basal S. slope of Morne Micotrin, elev. ca. 2,400 ft.
Usable by passenger cars nearly all the way, this is the highest
paved road on the island and cuts through the rich montane rain
forest of Morne Micotrin; thus, botanists find many species typical
of this plant formation by merely hiking along the road or making
short forays into the forest. The average annual rainfall measured at
a weather station near the lake is about 250 inches. Most of the
myxomycetes were collected near the road (at the edge of the forest)
and in adjacent citrus orchards.

10. St. Paut: Along the basal part of the trail ascending Morne
Trois Pitons on its NW. slope, near Pont Cassé, elev. 2,000-2,200
ft., a region of disturbed montane rain forest grading into newly cut
clearings and banana plantations.

11. St. Prerer: Rochette, ca. 2-3 miles inland from Colihaut,
elev. 2,200 ft., also an area of disturbed rain forest and cultivation.

12. St. Persr (border of St. Joun): S. end of Syndicate Estate,
near the road junction and on the basal slope of Morne Diablotins,
elev. 2,000-2,100 ft., an area of more or less disturbed rain forest
merging with banana and citrus plantations. Myxomycetes were
collected mostly in the forest.

13. St. ANDREW: Central Forest Reserve, Borne area, elev. ca.
1,400 ft., in rain forest.

14. St. Anprew: Near the road junction ca. 1.5 miles W. of Eden
Estate, in the swamp-woodland formation by the Eden River (prob-
ably the spot cited by Hodge as 2 miles W. of Calibishie). Myxomy-
cetes were found on a group of huge Pterocarpus officinalis Jacq. logs
lying in the swamp-woodland, on decaying logs on solid ground
across the road, or were cultured from bark of nearby Yerminalia
catappa L. and Carica papaya L. trees.

15. St. Davin: Near the road bridge by the mouth of the Castle
Bruce River, elevation near sea level; a dry (at least when visited),
flat coastal area partly planted to coconut and banana.

16. St. Davip: Taberi Estate, near the mouth of the Bibiay
River, elev. 150-200 ft., essentially similar to locality no. 15.

Certain places, such as Felicite (St. Parrick) and South Chiltern
House (St. Luxe) were infertile with respect to field collections but
brought forth a good number of specimens from bark samples in
moist chamber culture, denoting a potential productivity.

In addition to the field collections, nearly 200 moist chamber cul-
tures were maintained during the 3 months. They contained mostly
pieces of bark from living trees, but occasionally also deadwood from
the beach, various types of plant detritus, pieces of burlap, sclerotia,
and living plasmodia with pieces of their substrates. The cultures

404 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

were prepared usually the same day as collected by soaking the samples
(except active plasmodia) overnight in new, stacked finger bowls that
had been washed in tap water, covered on the bottom with filter
paper, and filled with tap water to cover the sample. After the excess
water was poured off, the dishes were observed every few days for
myxomycete developments and substrate pieces bearing ripe fructifi-
cations were removed. Since the only purpose of this work was to
supply further records of species occurring in Dominica, no attempt
at sterilization of the cultures was made. The use of new materials
and the performance of the entire work on the island rendered foreign
contamination unlikely. The cultures were kept at least one month
and those not disintegrated or overrun by filamentous molds were
again soaked overnight and maintained approximately another
month; no effort was made, however, to provide a uniform timetable
for all cultures. The rewetting procedure added an unexpectedly large
number of specimens, bringing the proportion of moist chambers
producing myxomycetes to ca. 35%.

Geographical Considerations

The present known total for Dominica of 99 or 100 species of
myxomycetes compares very favorably with approximate figures
published for other Caribbean islands (Antigua, 50; Cuba, 24, His-
paniola, 30; Jamaica, 110; Puerto Rico and Virgin Islands, 70; Trini-
dad, 50) and indicates that this verdant island abounds in slime molds.
In addition to the 2 newly described taxa, 9 known species appear to
constitute new records for the Antilles: Cribraria minutissima, Licea
operculata, Perichaena minor, Comatricha subcaespitosa, Echinostelium
cribrarioides, Diderma chondrioderma, Didymium comatum, Physarum
psittacinum, and P. pulcherripes. Especially interesting among these is
Echinostelium cribrarvoides, previously noted only from the Greek
type collection. Badhamia panicea, an early record (17), is still known,
in the West Indies, only from Dominica.

Table 1 shows several other species to be noted from only one
additional Caribbean island (Ceratiomyza sphaerosperma, Hemitrichia
paragoga, Comatricha aequalis, Macbrideola martinii, Stemonitis
flavogenita, Badhamia affinis, Didymium intermedium, Physarum
galbeum, P. roseum, and P. rubiginosum from Jamaica; Perichaena
vermicularis and Didymium minus from Antigua; Didymium erustaceum
from Cuba; Badhamia nitens and Didymium difforme from Hispaniola;
and Physarum auriscalpium from St. John, U.S. Virgin Islands).
These listings, however, probably do not reflect the actual Antillean
distributions for the species concerned, because the myxomycota of
most of the islands has not yet been intensively surveyed. Floristic

FARR—MYXOMYCETES FROM DOMINICA 405

studies of the myxomycetes inhabiting the Caribbean islands are
still in the beginning stages.

A number of species more or less common in the West Indies were
not encountered in Dominica, but are undoubtedly present and per-
haps prevalent during some other time of the year. Badhamia nitens,
B. panicea, Stemonitis herbatica, and Diderma rugosum, reported by
Lister (17), were not recovered; Arcyria nutans and Comatricha nigra
were conspicuous by their absence; Arcyria insignis and Diderma
testaceum likewise will probably be added to the record by future
collectors. As might be expected, Ceratiomyzxa fruticulosa, Arcyria
cinerea, A. denudata, Hemitrichia stipitata, H. vesparium, H. serpula,
Stemonitis fusca, Fuligo septica, and Physarum melleum were practically
ubiquitous throughout the collecting period. H. stipitata, as in Jamaica
and elsewhere, fruited under widely differing conditions of moisture
and exposure, its habitat ranging from dry, burnt, exposed wood to
very wet, rotten, deeply shaded logs. Cribraria violacea (in moist
chamber), Diderma effusum, Didymium nigripes, D. squamulosum,
Physarum compressum, P. nucleatum, and, oddly, Trichia favoginea
(a temperate-zone species not very often collected in the tropics)
likewise showed up frequently. Lycogala epidendrum was abundant,
but L. exiguum was almost equally common. Tubifera microsperma,
Comatricha typhoides, Clastoderma debaryanum, and Physarum nutans
were not as prevalent as in Jamaica, but Didymium squamulosum Was
by far more plentiful in Dominica.

Lister (18) remarked on the scarcity of the genus Trichia in Antigua
and speculatively attributed its [then] absence in Dominica to the
rapid destruction, by termites, of rotten wood, which is the substrate
for plasmodia of Trichia spp. This reasoning seems unfeasible, since
other wood-inhabiting slime molds (Hemitrichia spp., Stemonitis spp.,
Lycogala spp., etc.) were then known from Antigua and are widely
distributed over the Caribbean region. In Jamaica 8 species of Trichia
are known, but there the genus is generally confined to the higher al-
titudes (4,000 ft. or more) having a more temperate climate. The more
tropical environment and absence of such high elevations on Antigua
and (except on a few windswept peaks) Dominica may provide a more
plausible explanation. Trichia favoginea, however, is well established
on the latter island, and additional species of Trichia may well de-
velop at other times of the year.

One disappointing surprise was the lack of myxomycetes on or
within the abundant epiphytic bromeliads, a habitat frequently
colonized by certain species in Jamaica.

Obviously a minimum of one year’s study would be required to
gain even a preliminary knowledge regarding any seasonal distribu-
tion of the various slime-mold species on Dominica. Since the short

TABLE 1.—Hitherto known Antillean distribution of myzomycetes collected in

Dominica
Ls]
q
© od
oO o
Species a 8 8 a 1 eo & 2 a3 References
® « £ 8 8 $8 gud 3
3 2 g 3 q BSa s
q 3 © 5 4 &€ 3 Sp4 &
“<- 0 ADH S SF g a
Ceratiomyxales
Ceratiomyza fruticulosa x X X X X x X 2,4, 7,8, 9, 11, 14,17
C. sphaerosperma x 2,9
Liceales
Cribraria intricata x xX xX 4,9,14
C. languescens x xXx x X 4,9, 14,17
C. microcarpa x x 2,9, 14
C. minutissima New Antillean record
C. tenella xX xX x 2, 9,17
C. violacea x x x X 2,4, 9, 14,17
Dictydium cancellatum x xX xX x X 2,4, 8,9, 14,17
Licea operculata New Antillean record
Lycogala epidendrum xX x xX X X x 4,5, 6,7, 8,9, 14,17
LL. exiguum xX xX X 8,9
Reticularia lycoperdon x x 9, 14
Tubifera ferruginosa x X X x 7,9, 11, 14
T’. microsperma x x x xXx X 4, 5,8, 9,17
Trichiales
Arcyria cinerea xX x x X X x X 2,4, 6,6, 7,8, 9, ll,
13, 14, 17, 21
A, denudata X BPI X X X X x xX X 2,4, 7,9, 11, 14, 17
A. incarnata xX x xX x xX 4,9, 11, 14,17
Hemitrichia paragoga x 9
Hi. serpula x xX xX xX X x X 2,4, 5, 7,8, 9, 14,17
Hi. stipitata xX xX xX XxX X x X 2,4, 5a, 6, 89, 9, 11,
13, 148, 178
FY. vesparium xX xX x xX X 2,4, 5, 9, 14, 17
Perichaena chrysosperma x x x X x X = 2,4, 5,7, 9, 11, 14, 17
P, depressa xX xX xX X 2,4,9, 14,17
P. minor New Antillean record
P. vermicularis x 17
Trichia favoginea BPI°¢ xXx X xX X = 2,4b, 7b, 9, Ide
Echinosteliales
Echinostelium cribrarioides New Antillean record
E. minutum xX X 24,9
Stemonitales
Clastoderma debaryanum X X 2,4,9
Comatricha aequalis x 9
C. elegans BPI Xx XX X 2,4,9,11
C. irregularis x x 9, 14
C. longa xX xX X 4,9,14,17
C. subcaespitosa New Antillean record
C. typhoides >. x x xX xX X 2,4,5,7,9, 14,17
Diachea bulbillosa xX xX 9, 14
D. leucopodia x x xX X 2,4,5,9, 11,17
D, silvaepluvialis New Antillean record
(new species)
Lamproderma arcyrionema xX xX XX x 2, 7,9, 11, 14, 17
L. scintillans x xX xX 9,11, 17
Macbrideola martinii X 2,3
Stemonitis axifera x x Cc XxX X 5, 6, 7,8, 9, 11,13
S. smithii x 4,9
S. flavogenita xX 9
S. fusca xX x X xX X X xX 4,5, 7,8, 9, 14,17
S. herbatica x xX X X x 6, 7,8, 9, 14, 17, 21
S. hyperopta xX x 9, 14
S. nigrescens xX xX 2,9, 14
8S. splendens x xX xX X 4,9, 14,17
S. webberi xX x 9, 20

See footnotes at end of table.

TABLE 1.—Hitherto known Antillean distribution of myxomycetes collected in

Dominica—Continued
Ls}
z a3
oO i]
Species s 8 8 2 3 & a = 3% References
Ep e . a 8 a & 859 8
3S 2 of B goa 4
q =) o.6U68lU g S sP4 <2
< o AO HR § 2 «A mo
Physarales
Badhamia affinis xXx 9
B. nitens x xX 7,17
B. panicea x 17
Craterium aureum xX x 6,7, 9, 14
C. leucocephalum x x xX xX X 4,5,9,14,17
Diderma chondrioderma New Antillean record
D. effusum x x x X 4,9,14,18
D. hemisphaericum x xX xX x X 4,9, 11, 14,17
D. rugosum x xX X 4,9,17
D. spumarioides x x xX 9, 14,17
Didymium clavus x xX xX X 2,4,9,11,17
D. comatum New Antillean record
D. crustaceum x 5
D. difforme x 9
D. inter medium x 2,9
D. iridis x x xX X 4,6,9, 11, 17, 21
D, minus x 18
D. nigripes xX xX X x X = 2,4,7,9,11, 14, 18
D, sguamulosum xX xX X x X = 4,5,7,9, 11, 14,17
Fuligo cinerea x xX 9, 17
F. septica xX xX X X X xX X 4,5,7,8,9, 14,17
Physarella oblonga x x x X 2,4,9, 14,17
Physarum aeneum x xX 14,17
P. auriscalpium x 9
P. bogoriense xX x x X 2,4,9,14,17
P. bubalinum New Antillean record
(new species)
P. cinereum x x x xX X = 4,5,7,9, 11, 14,17
P. compressum x x xX 6, 7, 9, 11, 14, 17, 21
P. didermoides xX x 6, 7, 9, 14, 21
P. echinosporum x x 9, 20
P. galbeum x 9
P. lateritium x x 9, 18
P. leucophaeum x x xX 2, 6, 7, 8, 9, 13, 17
P. leucopus x 9
P. melleum x x x X 4, 7,9, 11, 14, 17
P. notabile x x 8, 9
P. nucleatum x x 2,9, 18
P. nutans x xX x X 4,7, 9, 13, 14
P. oblatum x 9
P. penetrale x 9
P. polycephalum x xX x xX 4, 5,9, 14
P. psittacinum New <Antillean record
P. pulcherripes New Antillean record
P. pusillum x xX xX xX 2, 7, 9, 11, 14, 17
P. roseum x 9
P. rubiginosum x 9
P. stellatum x x x x X 2, 4,9, 14,17
P, tenerum x x x X 4,9, 14,17
P, viride x x x x X 2, 4, 8, 9, 14, 17

« As H. clavata; H. stipitata was formerly a synonym of H. clavata and, as indicated by Martin (20, p.

68), early tropical collections so cited were H. stipitata,

b As T. affinis.

eAs T. persimilis; T. favoginea, T. affinis, and T. persimilis were united recently (10).

408 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

collecting period encompassed only +1 season and a limited number
of localities (many of which could not be visited regularly), no mean-
ingful statements on this subject can be made at this time. However,
the number and monthly distribution of collections of each species,
listed in table 2, may be of interest and serve future collectors as a
basis for comparison. It should be pointed out that the progressive
decrease revealed by Ceratiomyza fruticulosa, Arcyria cinerea, A.
denudata, Hemitrichia serpula, H. stipitata, H. vesparium, Didymium
nigripes, and Physarum compressum may be due largely to change in
collecting habits rather than season. After the first month or so of
collecting, these ubiquitous species were frequently passed over, except
as locality vouchers or otherwise especially desirable specimens.
Similarly, the more or less pronounced increase in frequency of the
Lamproderma spp., Diderma hemisphaericum, Didymium squamulosum
Physarum aeneum, and P. melleum can perhaps be ascribed, at least
partly, to the fact that their preferred habitat (buried in leaf litter)
was less thoroughly investigated during the first few weeks.

The number of myxomycetes acquired from moist chambers is high
compared with results obtained previously in Jamaica by Alexopoulos
and Beneke (2) and by me (9). More than one-third of the cultures
produced myxomycetes and many yielded more than 1 species,
especially after being resoaked. The often heavy infestation of the
cultures with mites, springtails, etc., may have suppressed the appear-
ance of additional species.

The development time of each of the 26 species obtained from moist
chamber culture (not including those collected as plasmodia, which
are considered field collections) is included in table 2; 14 of these
species were also found in the field, including nearly all cited members
of the Physarales. Cribraria violacea was the species most frequently
harvested, appearing in more than a dozen moist chambers; it was,
surprisingly, never observed in the field. No previously published
notices of moist chamber fruitings were found for the following species:
Diderma hemisphaericum, Fuligo cinerea, Physarum leucophaeum, P.
leucopus, P. melleum, P. notabile, and P. viride.

The relatively long time needed by all myxomycetes to fruit in the
moist chambers is noteworthy and perhaps due, at least partly, to
the use of tap water (distilled water being unavailable). Even members
of such genera as Cribraria, Licea, and Echinostelium, which often
fruit within a few days (1), required at least 1 to 2 weeks. In the
cases of Arcyria cinerea, Hemitrichia vesparium, Lamproderma arey-
rionema, and Physarum pusillum, the disparity of development periods
shown by different cultures within the species is perplexing, but the
later fruitings (as well as some of the other species appearing after
very long incubation) may have resulted from contamination during
handling and rewetting.

FARR—MYXOMYCETES FROM DOMINICA 409

Acknowledgments

I wish to thank G. W. Martin and C. J. Alexopoulos for their help
in the identification of specimens, J. A. Stevenson and P. L. Lentz
for determining fungal substrates of certain collections, and Arlene
Frank for technical assistance.

New Species

Diachea silvaepluvialis, sp. nov. Fiaurss 1, 2

Sporangia gregaria, stipitata, globosa vel late ovoidea, 1-1.25 mm
alta, 350-450» diam., hypothallo circulari, membranaceo, brunneo
plerumque subtenta; stipes 500-670 (—750)u altus, membrana brun-
nea translucenti tectus, crystallis calcareis, albis 5-154 diam. dense
farctus, basaliter dilatatus, apicaliter pro columella plus minusque
cylindracea apicem sporangii attingens. Peridium tenue, plus min-
usque argenteo-iridescens, ad basim pro disco vel cupula vadosa
leniter rugulosa saepe persistens; capillitium ab columella facile
separatum, reticulum laxum filorum eracilium (1-2 diam.) atro-
brunneorum levium, ramificantium et anastomosantium, cum ramu-
lis numerosis, formans. Sporae atrobrunneae, globosae, dense sed
inaequaliter spinulosae, 10-14 (-16)y diam.

Sporangia gregarious, stalked, globose to broadly ovoid, 1-1.25 mm
high, 350-450» in diameter, many subtended by a brownish, mem-
branous circular hypothallus; stalk %-% (-%) of total height, ex-
ternally dark orange or dark brown, glossy, faintly striate, consisting

Ficures 1-2.—Diachea silvaepluvialis sp. nov.: 1, Habit, 25; 2, portion of capillitium and
4 spores, X 900.

410 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

TABLE 2.—Monthly distribution of myxomycete species collected and development

periods for moist chamber cultures

Number of specimens collected

Development periods

Species (days) for moist chamber
January February March specimens
Ceratiomyxales
Ceratiomyra fruticulosa 5 6 3
C. sphaerosperma 1
Liceales
Cribraria intricata 1 1 1
C. languescens 1 1
C. microcarpa 2a 3b 1 10, 14, 23, 25, 25
C. minutissima Ja+1b 12, 24
C. tenella 3
C. violacea 33+2b 34+3b 7, 10, 11, 12, 14, 14, 14,
15, 16, 23, 25
Dictydium cancellatum 3 6
Licea operculata 2a 3b 11, 14, 20, 23, 28
Lycogala epidendrum 1 4 3
L, exiguum 3 1 1
Reticularia lycoperdon 3 1 1
Tubifera ferruginosa 1
T. microsperma 2
Trichiales
Arcyria cinerea 8+2b+106 5+1b 1 12, 15, 24, 54
A, denudata 10 6 2
A. incarnata 3 4
Hemitrichia paragoga 1
HI. serpula 8 3 3
H. stipitata 10 6 2
FT, vesparium 4+1b 2+1> 1 28, 51
Perichaena chrysosperma 1 1
P. depressa 8 2 1
P. minor 1
P. vermicularis 1b 2b 23, 25, 34
Trichia favoginea 2 6 2
Echinosteliales
Ecchinostelium cribrarioides 1s 24
E. minutum 1a 2b 17, 20, 27
Stemonitales
Clastoderma debaryanum 1 2
Comatricha aequalis 2 2
C. elegans 1b 24
C. irregularis 1 3
C. longa 7 1
C. subcaespitosa 1 3
C. typhoides 5+16 1 15
Diachea bulbillosa 1 1
D, leucopodia 2
D, silvaepluvtalis 1 1
Lamproderma arcyrionema 1+2b 1+2b 2 14, 43, 66, 77
L. scintillans 1 1 4
Machrideola martinii 1b 20
Stemonitis axifera 3 1 2
S. smithii 2 1
S. flavogenita 2or3
S, fusca 6 3 2
S. hyperoptera 2
S. nigrescens 1
S. splendens 3 3
S. webberi 1 2 3 or 4

See footnotes at end of table.

FARR—-MYXOMYCETES FROM DOMINICA 411

TaBLE 2.—Monthly distribution of myxomycete species collected and development
periods for moist chamber cultures—Continued

Number of specimens collected Development periods
Species (days) for moist chamber
January February March specimens
Physarales
Badhamia affinis lor2
Craterium aureum 1
C. leucocephalum 3
Diderma chondrioderma 1
D, effusum 2 8+1b 44
D, hemisphaericum 1b 1 4 17
D, spumarioides 1
Didymium clavus 1 2
D. comatum 1
D. crustaceum 1
D. difforme 1
D, intermedium 1
D. tridis 2 1+15 28
D. minus 1
D, nigripes 3 7 3
D., squamulosum 7 12
Fuligo cinerea 1 1 26
F, septica 1 9 3
Physarella oblonga 2 4 1
Physarum aeneum 1 4
P, auriscalpium 1b 1+1> 15, 31
P. bogoriense 2 4 3
P. bubalinum 1+?1¢ ca, 29
P. cinereum 3
P. compressum 7 8 2
P. didermoides 4
P. echinosporum 2
P. galbeum 1
P, lateritium 1
P. leucocephalum ja+2b 15, 20, 23
P. ?leucopus le ca. 29
P. meliewm le 2 12 67
P. notabile 148 23
P. nucleatum 6 8 4
P. nutans 2 2+18 1 7
P, oblatum 1
P. penetrale 1
P. polycephalum 1 1 3
P. psittacinum 2 2
P. pulcherripes 1 1 1
P. pusillum 1s 1+18+41> 12, 43, 50
P. roseum 1
P. ?rubiginosum 1
P. stellatum 3 2
P, tenerum 3 1
P. viride 2+1¢ 44+1b 23, 36

a =m.ch., collected and harvested the same month.
b =m.ch., harvested the following month.

¢ =m.ch., collected in January, harvested in March.
Unmarked =field collections.

313-143—68———-2

412 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

of a translucent, brown membrane densely packed with white, single
or aggregated, more or less angular lime crystals 5-15y in diameter;
basally enlarged, apically continued through the entire sporangium
as a more or less cylindrical columella. Peridium delicate, moderately
silvery-iridescent, usually persisting at base of sporangium as a broad,
slightly rugose disc or shallow cup, pale brown and smooth by trans-
mitted light. Capillitium arising all along columella, consisting of
an open, loose, easily detached network of delicate, dark brown,
smooth, slightly wavy, branching and anastomosing threads 1-2y
in diameter, with many concolorous or slightly paler free ends. Spores
dark brown in mass, dark purplish brown by transmitted light,
globose, closely but unevenly spinulose, sometimes with a densely
warted area 3.5-5.0u in diameter, 10-124 (2197) or 10-14(-l6u
(2998) in diameter.

Holotype: B.W.I., DOMINICA: Sr. Grorae parish, road from Laudat to Fresh
Water Lake, elev. 2,200-2,500 ft. (9), on moist, decaying leaves on shaded ground,
Farr 2197 (BPI). Paratype: St. ANprEew parish, Central Forest Reserve,
Borne area, elev. 1,400 ft. (13), on moist, dead leaf on shaded ground in rain
forest, Farr 2998 (BPI).

Two species of Diachea with nonwhite stalks have been described.
In D. thomasii Rex the lime itself is bright orange; the hypothallus is
venulose and likewise orange; the sporangia are sessile or short-stipi-
tate and larger, and the columellae much smaller than in D. silvae-
pluvialis. Diachea megalospora Thind & Manocha evidently is very
similar to D. thomasii, differing from the latter only in having slightly
larger and more coarsely marked spores.

Physarum bubalinum, sp. nov. Figures 3-5

Sporangia fasciculos vel rosulas gregarias 1-3 mm diam. formantia,
sessilia vel pseudostipitata, bubalina, plusminusve convoluta, leviter
compressa, ca. 250u lata, unitunicata; hypothallus nigellus, corneus,
inconspicuus; peridium membranaceum, albidum, granulis calcareis
bubalinis incrustatum; columella nulla; capillitium ex nodulis albis
maximam partem grandibus, angularibus vel elongatis et filis paucis
subhyalinis constans, aliquando pseudocolumellam formans; sporae
atrobrunneae, globosae vel ovoideae vel ellipsoideae, minute et dense
spinulosae, 10-12 vel 1012u diam.

Sporangia in gregarious clusters or rosettes 1-3 mm in diameter,
sessile or borne on black, weak, stalklike extensions of the hypothallus,
convoluted to nearly pseudoaethalioid, buff, slightly compressed
laterally, ca. 250 thick. Hypothallus blackish, horny, inconspicuous.
Peridium single, membranous, whitish, densely encrusted with tawny
or pale brown, sometimes fading, clusters of lime granules. Columella
none. Capillitium consisting of white lime nodes greatly varying in
size and shape but predominantly large and angular or elongated (occa-

FARR—MYXOMYCETES FROM DOMINICA 413

Figures 3-5.—Physarum bubalinum sp. nov.: 3, Habit, X25; 4, two spores, 900; 5,
capillitial lime knots, % 900.

sionally aggregated into a small pseudocolumella), and few, subhyaline
connecting threads. Spores dark brown in mass, purplish brown by
transmitted light, globose to ovoid or ellipsoid, minutely and densely
spinulose, 10-12 or 10X12, in diameter.

Holotype: B.W.I., DOMINICA: Sv. Josers parish , along Layou River near
Clarke Hall Estate, elev. ca. 200 ft. (1), on outside of rind of cacao fruit (Theo-
broma cacao L.) rotting in deep shade, Farr 2101 (BPI).

A scanty moist chamber development with broken but apparently
similarly shaped, iridescent sporangial clusters and similar spores
may represent a completely limeless phase of this species (collected

414 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

in Sr. PevEr parish, summit of Morne Diablotins, elfin woodland, elev.
4,700 ft., on stem of Lobelia stricta Sw. in moist chamber, Farr 3083b).

This myxomycete has similarities to several known species of Phy-
sarum, but cannot be accommodated in any of them. From P. nic-
araguense and convoluted forms of P. compressum it differs mainly in
its sessile habit and tawny color; from P. gyrosum in color, capillitial
structure, and spore size. Two closely related, temperate-zone species,
P. virescens and P. confertum, sometimes produce clumped, off-color
sporangia resembling those of the new species in shape, but consider-
ably smaller and with more delicate capillitium; P. virescens, further-
more, has smaller, smoother spores.

Annotated List of Species

The species of myxomycetes are listed in alphabetical order within
the genera; these, in turn, are arranged alphabetically under the
nowadays generally accepted orders Ceratiomyxales, Liceales, Tri-
chiales, Echinosteliales, Stemonitales, and Physarales. The localities
are cited according to parishes, starting with Sr. J OSEPH, and arranged
in a clockwise direction. Within each parish, numbered localities,
listed in ascending order (locality numbers printed in boldface),
precede those without a number. For the unnumbered locations, only
those elevations above 100 ft. are mentioned.

Order CERATIOMYXALES

Ceratiomyxa fruticulosa (Miill.) Macbr. No. Amer. Slime-Moulds, 18. 1899.

DisrriBution: Cosmopolitan.

Sr. Josepu: 1, near underside of wet, decaying log, 2122; 2 (Cocoa
Centre), on underside of wet, rotting, decorticated log, 2552, 2568, ca.
% mile S. of Mero, at mouth of Mero River, on rotting log, 2596. Sr.
ANDREW: 5, on underside of wet, rotten log, 2590. Sr. Davin: 8, on wet,
rotting log, 21517, 2289, 2915, 2916 (the latter poor) ; 15, on rotten logs by
river and in banana plantation, 2737 (scanty); 1.5 miles S. of Castle
Bruce, on underside of very rotten, wet, shaded log in forest, 21/71
(yellowish). Sr. GEorau: 9, on wet, rotting, shaded log, 2199 (scanty).
Sr. Pau: 7, in hollow of decaying, wet stump, 2708; on side of wet,
rotting log, 3009.

Ceratiomyxa sphaerosperma Boed. Misc. Zool. Sumatr. 24: 1. 1927.

Disrripurion: Central America, Antilles, Indonesia.

St. Davip: Rosalie Agric. Exp. Sta., ca. 1 mile NE. of town, near
intersection of road to St. Sauveur, on fibers of decaying coconut
husks, 2139; the colonies appear globose but are seen, under a dis-
secting microscope, to consist of branches borne on gelatinous, more
or less dichotomously branched stalks.

FARR—MYXOMYCETES FROM DOMINICA 415

Order LICEALES

Cribraria intricata Schrad. Nov. Gen. Pl., 7. 1797.

DIsTRIBUTION : Cosmopolitan.

Sr. Perer: 11, on underside of very wet, rotten, shaded log, 2078a.
St. ANDREW: 5, on underside of wet, very rotten log, 2592 (moldy).
Sr. Pauu: 7, on rotting twig buried in leaf litter, 2821; sporangia
yellow, with fairly abundant free ends in the net.

Cribraria languescens Rex, Proc. Acad. Phila. 1891: 394. 1891.

Distrisution: Probably cosmopolitan.

Sr. JosePH: 2 (Cocoa Centre), on extremely rotten, wet, decorti-
cated log, 2560. St. ANDREW: Ca. 1 mile W. of Porte-La-Fin, on
decaying polypore on rotting log, 2352 (moldy).

Cribraria microcarpa (Schrad.) Pers. Syn. Fung., 190. 1801.

DistriBuTION: Probably cosmopolitan.

Sr. JosePH: 1, on bark of large tree (? Inga) behind “cabana” at
Clarke Hall swimming area, in moist chamber, 2846 (a reddish,
slightly atypical fruiting with spores 8-10, in diameter and many “‘giant
spores’); also from bark of ?leguminous tree in cacao plantation, in
moist chamber, 2989a, associated with Perichaena vermicularis (29896) ;
also on bark of Mangifera indica L. in moist chamber, 2990; 2, on
bark of Mangifera indica in moist chamber, 2367; on bark of rotten
log, 2783 (sporangia dull orange); Macoucheri Est., ca. } mile inland
from W. coast, on bark of indeterminate tree in moist chamber,
2366.

Cribraria minutissima Schw. Trans. Amer. Philos. Soc. II, 4: 260. 1832.

Distrisution: USA, South America, Europe, Asia, Dominica.

Sr. Jospru: 1, on bark of Cocos nucifera L. in moist chamber,
2368 (sporangia golden yellow). St. Parricx: Felicite, from exposed
log on beach in moist chamber, 2476a; associated with Arcyria cinerea
(2476b).

Cribraria tenella Schrad. Nov. Gen. Pl., 6. 1797.

DisTRIBUTION: Cosmopolitan.

Sr. Davip: Ca. 1-1.5 miles E. of north branch of Ravine Deux
Dleaus, S. of Bois Diable Ridge, elev. ca. 1,100-1,200 ft., on wet,
decaying, decorticate log at edge of open, partly cleared rain forest,
8030. Sr. Pauu: 7, on rotting log in banana plantation, 2831, 2833,
the latter associated with Arcyria cinerea; free ends in net not numerous.
Lister (17) describes Dominica material as having ‘“‘scarcely the rudi-
ments of a cup,” mostly round nodes, and no “free rays.”’

Cribraria violacea Rex, Proc. Acad. Phila. 1891: 393. 1891.

DIsTRIBUTION: Cosmopolitan.
Str. Josreru: 1, on bark of indeterminate tree (? Inga) in moist
chamber, 2332a, associated with Licea operculata (28326); on bark of

416 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

“Salamanda” (botanical name as yet undetermined) in moist chamber,
2698a (abundant); on bark of Artocarpus communis Forst. in moist
chamber, 2864, teste G. W. Martin; 2, on bark of Mangifera indica
in moist chamber, 2368 (4 sporangia); Mero, on bark of Lagenaria
siceraria (Mol.) Standl. in moist chamber, abundant, 2595, teste
G. W. Martin; associated with Physarum nutans (2596). St. ANDREW:
14, on bark of Carica papaya in moist chamber, 2478, teste G. W.
Martin (sporangia of last 2 harvests larger); on bark of Terminalia
catappa in moist chamber, 2369. St. Parrick: Felicite, from exposed
log on beach in moist chamber, 2475. St. Luxe: Pointe Guignard,
from wood of exposed log near beach, in moist chamber, 2860; South
Chiltern House, elev. ca. 1,200 ft., on bark of indeterminate large
tree with light gray, splintery, easily detachable bark, in moist
chamber, 2722b, associated with Physarum notabile (2722a). Sr.
Pauu: N. of Mahaut, near Rodney’s Rock, on bark of Terminalia
catappa in moist chamber, 2855.

Dictydium cancellatum (Batsch) Macbr. No. Amer. Slime-Moulds, 172. 1899.

DisrriBuTION: Cosmopolitan.

St. Josepn: 1, on underside of rotting log, 2362, 2480 (both very
moldy), 2674 (good); 2 (Cocoa Centre), on underside of wet, rotting
log, 2661 (mostly moldy), teste G. W. Martin; 3, on top of wet, rotting
log, 2616, 2902 (a beautiful specimen) ; Macoucheri Est., mile inland
from W. coast, on bark of decaying, shaded log, 2061. Sr. Davin: 8,
near underside of wet, rotting log, 2661. St. Grorae: Along trail to
Trafalgar Falls, elev. ca. 1,000 ft., on wet log in shade, collected by
Mrs. Clifford Evans, 2326 (3 sporangia).

Licea operculata (Wing.) Martin, Mycologia 34: 702. 1942.

Disrrinution: USA (New England and Midwest), Panama,
Europe, Japan, Dominica.

Sr. JosePH: I, on bark of indeterminate tree in moist chamber, 2766,
determined by G. W. Martin; on bark of Znga sp. in moist chamber,
2859b, associated with Physarum leucophaeum (2859a); on bark of
indeterminate tree (?/nga) in moist chamber, 23326, associated with
Cribraria violacea (2332a) (placement of this specimen into moist
chamber for ca. 1 week, in May 1966, in Beltsville, Md., yielded 2
additional sporangia). Sr. Anprew: 14, on bark of Pterocarpus
officinalis in moist chamber, 2370. St. Pauu: Beauclair Est., ca.
0.9 mile W. of Springfield Est., elev. ca. 1,000 ft., on bark of tall,
acacia-like leguminous tree in moist chamber, 2854. My identifica-
tions of this species are based on Dr. Martin’s determination of
2766.

Lycogala epidendrum (L.) Fr. Syst. Myce. 3:80. 1829.

DistrrRIBUTION: Cosmopolitan.

Sr. JosepH: 1, on top of shaded, moist, decorticate log, 2599;
2, on wood of rather dry, rotten log, 2226 (peridium black); 3, on

FARR—MYXOMYCETES FROM DOMINICA 417

top of shaded, wet, decorticate, rotten log, 2613, 2669, 2890, 2891;
2890 resembles L. exiguum, but the peridial warts are not chambered
(perhaps a young stage of the latter species). St. Joun: 4, on top of
rotting log, 2977. St. Pau: Beauclair Est., ca. 0.9 mile W. of Spring-
field Est., elev. ca. 1,000 ft., on top of smooth, decorticate, rotting
log, 2562.

Lycogala exiguum Morg. Journ. Cincinnati Soc. Nat. Hist. 15: 134. 1893.

DisrriputTion: Cosmopolitan.

Sr. JoserH: 1, on rotting log, 2029; 2, on bark at top of rotting
log, 2784 (blackish). Sr. ANDREW: Ca. 1 mile W. of Porte-La-Fin,
on log, 2343. Sr. Parricx: La Pleine (Plaisance Bay, ca. 1 mile N.
of La Ronde River), on side of decorticate decaying log, 2307.
Sr. Paut: 7, on side of moist, rotting log, 2697. This species seems
to be more common in Dominica than in Jamaica and some other
islands, where L. epidendrum predominates.

Reticularia lycoperdon Bull. Hist. Champ. France, 95. 1791.

DistrIBUTION: Cosmopolitan.

Sr. JosErH: 1, on exposed side near bottom of decaying log,
2033; on bark of decaying logs, 2843. Sr. ANDREW: 14, near underside
of fairly exposed, rotten member of log pile in clearing, 2246; and on
same log, 2341. St. Grorau: Roseau Botanic Gardens, on side of
rotting hardwood log, 2636 (small aethalium).

Tubifera ferruginosa (Batsch) Gmel. Syst. Nat. 2: 1472. 1791.

DistRIBUTION: Cosmopolitan.

Sr. Joun: Northern outskirts of Portsmouth, along road to Cabrits,
on decaying coconut log, 2958.

Tubifera microsperma (Berk. & Curt.) Martin, Mycologia 39: 461. 1947.

Distrisution: Probably cosmopolitan.

Sr. JosErH: 3, fairly abundant on rotten log, 2911. St. Pauu: 7,
on rotting logs in banana plantation, 2825 (somewhat moldy).

Order TRICHIALES

Arcyria cinerea (Bull.) Pers. Syn. Fung., 184. 1801.

DisrrIBuTION: Cosmopolitan.

Sr. Josep: 1, on end of decaying, wet log, 2031 (very poor),
2308. Sr. Perer: 11, on underside of wet, rotting, shaded log, 2083
(digitate form). Sr. ANnprEw: 5, on underside of rotting log, 2589
(digitate form) and 2591 (associated with Trichia favoginea); on
burned wood inside of hollow stump, 2625 (teste G. W. Martin); 13,
on rotting debris in rain forest, 2996. This specimen, although micro-
scopically typical of the species, manifests an unusual color variation.
The sporangia are black on the outside and olivaceous yellow or drab
internally, with black or iridescent, olivaceous-yellow calyculi and

418 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

long, black stalks. The globose, pale yellow spores, 7-Sy in diameter,
exhibit, under the 90X oil-immersion objective, a faint but regular
reticulation of 12-14 meshes per hemisphere. This characteristic, not
mentioned for the species in any of the standard monographs, was
also found in several other, authentic collections of A. cinerea (in-
cluding the digitate form) both from the tropics and from temperate
regions. St. Davin: 8, on underside of wet, decaying log, 2286 (digi-
tate form); 16, on underside of moist, shaded, rotting log, 2313
(digitate form). Sr. Parricx: Felicite, in moist chamber from exposed
log on beach, 24766; in moist chamber from another log, 2984. Sr.
Marx: Scotts Head, on bark of indeterminate tree along beach, in
moist chamber, 2988; slightly aberrant in the pale flesh color of the
sporangia, and in the larger spores (8-10u). Sr. GrorGeE: 9, in rain
forest in shade, from a moist chamber in which an orange plasmodium
was placed, 23666; along trail to Trafalgar Falls, elev. ca. 1,000 ft.,
on underside of wet, decaying, mossy log in shade, 2829 (very poor
and moldy). Sr. Paun: 6, on underside of wet, rotting log, 2203; 7,
2702 (very poor); 10, 2273 (poor), 2427.

Arcyria denudata (L.) Wettst. Verh. Zool.-Bot. Ges. Wien 35: Abh. 535. 1886.

Distripution: Cosmopolitan.

Sr. JosePH: 1, on underside of rotting log, 2359, 2361 (the latter
weathered and moldy); on underside of very damp, rotting branch
on ground, 2773; 3, on underside of wet, decorticate, very rotten log,
2610. St. Peter (bordering Sr. Joun): 12, on underside of rotting
logs, 2374; a weathered, chocolate-colored specimen having capillitium
sculptured with nearly complete rings and some reticulate fragments.
St. ANDREW: 5, on underside of rotting log in banana plantation,
2619; 13, on wet, rotting stump in rain forest, 3000, collected as im-
mature, brown sporangia; 14, on underside of rotting log covered by
leaves, etc., 2246, 2247. Sv. Davin: 6, on rotting logs in banana planta-
tion, 2726; 8, on underside of rotting log, 2158 (sparse) and near
underside of wet, burned, decorticate, rotten log, 2656 (associated
with Hemitrichia stipitata); on bark near underside of rotting log,
2473 (weathered); 16, on log, 2311. Sv. Grorae: 9, on underside of
wet, rotting log, 2/94 (sparse). Sr. Pauu: 7, on underside of wet,
rotting log, 2695; 2 miles NE. of Springfield Est., in pasture, elev.
1,800 ft., on underside of moist, decaying log, 2337 (sparse).

Arcyria incarnata (Pers.) Pers. Obs. Mycol. 1:58. 1796.

DisrriBuTion: Cosmopolitan.

Sr. JosEPu: 1, on underside of moist, decaying log, 2301, 2842, a
pale-violet gathering with persistent patches of peridium, but with
microscopic structure normal for the species; same data, 2844, a
“typical” specimen; 3, on decaying herbaceous stem on rotten log,
3018. St. Joun: Along border between W. and E. Cabrits, elev. ca.

FARR—MYXOMYCETES FROM DOMINICA 419

100 ft., on logs, 2965 (a weathered, brown, scanty, collection). Sr. Paut:
10, near underside of decaying log, 2270 (an anomalous development
characterized by capillitium 6-8, in diameter and covered only with
closely set rings, and by many “giant spores”); same data, 2275 (a
bright-pink fruiting consisting of extremely long and elastic sporangia).
Hemitrichia paragoga Farr, Bull. Inst. Jamaica, Sci. Ser. No. 7:34. 1957.

Distrispution: Antilles (Jamaica and Dominica).

Sr. Pau: 7, on underside of wet, rotting log, 2879. Although in
battered condition, this specimen can be recognized as almost cer-
tainly conspecific with the Jamaican type of the species, collected
in the rain forest of the John Crow Mountains; the Dominica fruit-
ing has larger spores (9-11 instead of 8-9), less spiny capillitium,
and shorter tapered free ends.

Hemitrichia serpula (Scop.) Rost. ex List. Monog. Mycetozoa, 179. 1894.

DisTRIBUTION: Cosmopolitan.

Sr. Josepu: 1, on bark of decaying log, 2120, 2125; 2, on underside
of greatly decayed, decorticate log, 2229; on decaying coconut log
near road, 2489, 2490; Macoucheri Est., 4 mile inland from W. coast,
on decaying, shaded branch, 2060, (1 annular plasmodiocarp). Sr.
Joun: 4, on side of mossy, shaded, rotting log, 2972, associated with
H. stipitata (2971). St. Davin: 8, near underside of decaying log,
2166; on roots of filmy fern on wet, decaying log, 2293; on underside
of rotten log, 2461; 16, on bark on underside of rotting log, 2309,
2312. St. Pau: 7, on dead leaves buried in litter pile, 2826; on rotting
twigs, roots, and banana debris, 2878.

Hemitrichia stipitata (Mass.) Macbr. No. Amer. Slime-Moulds, 207. 1899.

DistriBution: Cosmopolitan.

St. Josepu: 1, on wet, decaying log, 2082 (scanty), 2121, 2123
(the latter teste G. W. Martin); on very wet, rotten wood, 2725
(sporangia collected in red droplet stage, mature after 2 days); 3, on
decaying coconut petiole on ground, 2609. St. Perrr: 11, on wet,
shaded, decaying log, 2077 (poor). Sr. Joun: 4, on exposed under-
side of rotting log, 2971, associated with H. serpula (2972). Sr.
ANDREW: 5, on underside of moist, rotting log, 2623. Sr. ANDREW/
Sr. Davin border: Pagua Bay, on burned, fairly dry, shaded, rotting
log on black sand beach, 2241 (sparse). St. Davin: 6, on underside
of wet, rotten log, 22/9; 8, on bark near underside of decaying log,
2287; on wet, decaying stump, 3006 (approaching H. clavata by its
vaselike sporangia and slightly roughened capillitium); on underside
of decaying log in rain forest, 2419; Rosalie Agric. Exp. Sta., ca. 1
mile NE. of Rosalie, at junction of road to St. Sauveur, on underside
of wet, rotting log, 2137. Sr. Parricx: Felicite, near mouth of Sarisari
River, on protected side of burned, dry, decorticate log ca. 10 ft.
from beach, 2305. St. Grorce: Along trail to Trafalgar Falls, elev.

420 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

ca. 1,000 ft., on underside of very moist, shaded, decaying log, 2328.
St. Pauw: 7, on underside of moist, very rotten log, 270/; 10, on very
rotten, crumbling log in rain forest, 2433, teste G. W. Martin.
Hemitrichia vesparium (Batsch) Macbr. No. Amer. Slime-Moulds, 203. 1899.

Distripution: Cosmopolitan.

Sr. Josepu: 2 (Cocoa Centre), on decaying ?coconut log, 2549; 3,
on underside of rotting log, 2614. Sr. Perrr: 11, on bark on under-
side of rotting log, 2081, 2082. Sr. Joun: 4, on wood, mossy bark,
and protruding from loose bark on side of rotting log, 2976. Sv.
AnprREw: Pointe Baptiste, elev. ca. 100 ft., on wood, 2258. Sr.
Parrick: Felicite, near mouth of Sarisari River, on rotting, moist,
decorticate log in shade of Coccoloba grove ca. 15 ft. from beach,
2306, teste G. W. Martin; from exposed, rotting log near road, in
moist chamber, 2633. Sr. Luxe: South Chiltern House, elev. ca.
1,200 ft., on bark of Mangifera indica in moist chamber, 2982 (good
material).

Perichaena chrysosperma (Curr.) List. Mycetozoa, 196. 1894.

DistTRIBUTION: Cosmopolitan.

Sr. Perer: 11, on bark of rotten log, 2079; with capillitium bearing
long spines, and with spores 10-12y in diameter, subglobose to ovoid,
some thinner-walled on one side. Sr. Pau: Beauclair Est., ca. 0.9
mile W. of Springfield Est., elev. ca. 1,000 ft., on underside of wet,
very rotten, decorticate log, 2567.

Perichaena depressa Lib. Pl. Crypt., 378. 1837.

DisTRiIBuTION: Cosmopolitan.

Sr. Josep: 3, on side of moist, rotting log, 2611; between bridge
near mouth of Layou River and W. coast, on decaying coconut
petioles on ground, 2401; Macoucheri Est., ca. 4 mile inland from
W. coast, on bark of shaded, decaying branch, 2059; sporangia hoary,
pale greenish gray, with deep-orange capillitium. Sr. Perer: 11, on
bark of rotten log, 2078. Sr. Joun: 4, on underside of log, 2385
(moldy), 2974. Sr. ANDREW: 14, on inner bark and protruding from
cracks in outer bark of rotting log, 2248, 2339, 2340. Sr. Davin:
8, on decaying rootlets of filmy fern on bark of decaying log, 2285;
16, on decorticate, decaying log, 2308.

Perichaena minor (G. List.) Hagelst. Mycologia 35:130. 1943.
Distripution: North America, Great Britain, Japan, Dominica.
Sr. Davin: 8, on rotting log, 2800, sporangia stipitate to sessile;

capillitium yellow.

Perichaena vermicularis (Schw.) Rost. Monog. Sluzowce (Mycetozoa), Append.

34. 1876.

DisrrisuTiIon: Cosmopolitan.

St. JosepH: 1, on bark of ?leguminous tree in moist chamber,
29896, associated with Cribraria microcarpa (2989a); 2, on bark of

FARR—MYXOMYCETES FROM DOMINICA 4?1

indeterminate tree in moist chamber, 2486, partly ruined in transit;
on bark of ?Tabebuta sp. in moist chamber, 2853 (capillitium showing
long spines).

Trichia favoginea (Batsch) Pers. Neues Mag. Bot. 1:90. 1794.

Distripution: Probably cosmopolitan.

Sr, JoserH: 3, on rotting debris on decaying log, 2892. Sr.
Davip: 8, on decaying twig on ground, 2280; on dead leaf on decaying
log in rain forest, 2422 (battered) ; in hollow, rotten stump in freshly
cut forest, 2474; on dead banana leaf, 2654; on decaying, mossy log,
2659a; on decaying, small root, 2914; 16, on bark of shaded, rotting
log, 2310. Sr. Pauu: 7, on underside of rotting log, 2700; 10, on
decaying log, 2431. All specimens reveal elaters 5-64 or 6-7y in
diameter, and spores covered with the high, sharp, complete reticu-
lation peculiar to T. favoginea sensu strictu. The elaters of 2654
bear short spines, and those of 2422 have many bulbous free ends;
those of the other collections are smooth, and have short-tapered
apices.

Order ECHINOSTELIALES

Echinostelium cribrarioides Alexop. Amer. Mid]. Nat. 66:391. 1961.

Disrripution: Greece, Dominica.

Sp. Josepu: 1, on bark of “Salamanda” (botanical name as yet
undetermined) in moist chamber, 2364, teste C. J. Alexopoulos. The
sporangia differ from those of the Greek type collection in being
chocolate brown rather than cream-colored, in lacking a conspicuous
basal collar, and in exhibiting a denser capillitium.

Echinostelium minutum deBy. in Rost. Monog. Sluzowce (Mycetozoa), 215. 1874.

Disrrisution: Probably cosmopolitan.

Sr. JoserH: 1, on bark of indeterminate tree (?Jnga) behind
“cabana” at Clarke Hall swimming area, in moist chamber, 2857;
Macoucheri Est., ca. 4 mile inland from W. coast, on bark of Mangifera
indica in moist chamber, 2331 (very sparse, pinkish). Sr. ANDREW:
5, on bark of Dacryodes excelsa in moist chamber, harvested in abun-
dance, 2927 (nearly white). Both pink and white forms are known
from Jamaica; no color is indicated for collections from Trinidad.

Order STEMONITALES

Clastoderma debaryanum Blytt, Bot. Zeit. 38: 343. 1880.

Disrripution: Probably cosmopolitan.

Sr. Josepu: 1, on underside of very damp, rotting log in shade,
9127b (mostly moldy); 3, on side of wet, rotten log, 2668. St. Davin:
8, on white mold on underside of very rotten, wet, decorticate log,

2421.

422 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Comatricha aequalis Pk. Ann. Rep. N.Y. State Mus. 31:42. 1879.

Disrrisution: North America, Europe, Antilles, Africa.

St. Anprew: “Hill,” ca. 2 miles W. of Pagua Bay, in banana
plantation, on wet twig in leaf litter, 2951. Sr. Paut: 7, on underside
of moist, very rotten log and on polypore, 2698 (sporangia cocoa-
colored, somewhat clavulate); on underside of wet, rotting log, 3007
(moldy); Harris Soltoun Est., ca. 1.5 miles W. of Pont Cassé, elev.
ca. 1,400 ft., on wet, shaded, rotting stump near footpath in disturbed
rain forest, 2657 (spores 8-10 in diameter).

Comatricha elegans (Racib.) List. Guide Brit. Mycetozoa ed. 3, 31. 1909.

Distrisution: USA, Europe, Southern Asia, Japan, Antilles.

St. Perer (near border of Sr. Joun): 12, on bark of Daeryodes
excelsa in moist chamber, 2632.

Comatricha irregularis Rex, Proc. Acad. Phila. 1891: 393. 1891.

Disrrisution: North and South America, Antilles, Malaya, Japan,
Australia.

St. JosErPH: 2, on top of rotting coconut log, 2785. Sr. Jown: 4,
on side of rotting log and on old polypores, 2975; near mouth of
Picard River, Prince Rupert Bay, near exposed underside of rotting
log, 2956. Sr. Davin: 16, on fairly dry, decaying log, 2314 (slightly
aberrant).

Comatricha longa Pk. Ann. Rep. N.Y. State Mus. 43: 70. 1890.

Distrisution: Probably cosmopolitan.

Sr. JosepH: 2, on bark on underside of dry, decaying log, 2230,
teste G. W. Martin. Sr. ANDREW: 14, in swamp-woodland, pendent
from old fruiting body of Trametes corrugatus (Pers.) Bres. (2256, det.
J. A. Stevenson) growing on underside of large, very rotten, decorticate
Pterocarpus officinalis log, 2244; on same log, 2347, det. G. W. Martin.
These specimens correspond well with Lister’s (17) description (quoted
from the collector’s letter) of early Dominica material as “on a root
[,] hanging down like a goat’s beard.’ Sr. Davin: 16, on bark of
decaying log and on adjacent green leaves, 2316, 2317. Sr. Pauu:
10, on log, 2278 (a large fruiting on a silvery hypothallus, ca. 2 inches
in diameter, similar in color to Stemonitis flavogenita); 1.5 miles E. of

Pont Cassé, in banana plantation, elev. ca. 1,900 ft., on rotting log,
29380,

Comatricha subcaespitosa Pk. Ann. Rep. N.Y. State Mus. 43:71. 1890.

Disrrisution: North and South America, Europe, Dominica.

St. Davin: 8, on bark of rotten logs and on twigs and debris
covering bark, 2290; practically completely covering a 20.5 ft.
area on underside of wet, rotting log, 2918; ca. 0.5-1 mile E. of north
branch of Ravine Deux Dleaux, S. of Bois Diable Ridge, elev. ca.

FARR—MYXOMYCETES FROM DOMINICA 423

1,100-1,200 ft., on wet rotting log in open, partly cleared rain forest,
3028 (capillitium with brown thickenings; spores globose to ovoid,
8-10u in diameter, finely warted). St. Paun: 7, over entire underside
of rotting log and on nearby fern roots, leaves, moss, etc., 3010
(mostly moldy).

Comatricha typhoides (Bull.) Rost. ex List. Monog. Mycetozoa, 120. 1894.

Distripution: Cosmopolitan.

St. Josepu: 1, on end of decaying log and on adjacent dead leaf,
2302; 3, on very rotten, wet log, 3020, long-stalked. St. ANDREW:
14, on underside of very rotten, decorticate, wet Pterocarpus o ffici-
nalis log, 2242, 2243, associated with Arcyria denudata. St. Davin:
6, on underside of wet, rotting log, 2207 (abundant but moldy);
1.5 miles S. of Castle Bruce, elev. ca. 400 ft., on underside of shaded,
wet, decaying log in forest, 21/72. St. Parrick: Felicite, near mouth
of Sarisari River, on bark of Calophyllum (probably C. antillanum
Britton) along road, in moist chamber, 2484.

Diachea bulbillosa (Berk. & Br.) List. Journ. Bot. 36: 165. 1898.
DistripuTion: North America, Panama, Antilles, Asia, Africa.
St. Davip: Ca. 1 mile N. of St. Sauveur, on underside of living

Pandanus leaves near roadside, 2/68; sporangia globose, silvery-

iridescent; spores 10-14u in diameter. Sr. Paut: 10, on underside

of dead leaves in forest litter, 2432.

Diachea leucopodia (Bull.) Rost. Monog. Sluzowee (Mycetozoa), 190. 1874.

DistrisutTion: Cosmopolitan.

St. JosEPH: 1, on wet, dead leaves and twigs in roadside litter
pile, 2881, a fine, abundant collection; 3, on dead leaves, 2899, with
spores atypically small (6—-8y in diameter).

Diachea silvaepluvialis Farr (see p. 409)
Lamproderma arcyrionema Rost. Monog. Sluzowce (Mycetozoa), 208. 1874.

DistRIBUTION: Cosmopolitan.

St. JosePH: 1, on bark of Juniperus sp. in moist chamber, 2849, a
beautiful collection consisting of small sporangia (ca. 0.2-0.4 mm. in
diameter); on bark of Artocarpus communis in moist chamber, 2863;
3, near underside of very rotten, wet, decorticate log, 2663; Macoucheri
Est., ca. }; mile inland from W. coast, on bark of Mangifera indica in
moist chamber, 2986 (scanty but good). Sr. Davin: 8, on underside
of wet, rotting log, 2502 (moldy); ca. 1 mile N. of St. Sauveur, elev.
ca. 400 ft., on decaying leaf debris by rotting, roadside log, 2/69
(sporangia were collected in white droplet stage and turned purple
within 3-4 hrs). Sr. Grorace#: 9, on bark of Clusia plukenetii Urb.
in moist chamber, 2983a, associated with 2 sporangia of Physarum
pusillum (2983b). Sr. Pau: 7, on dead leaf in litter pile, 2941.

424 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Lamproderma scintillans (Berk. & Br.) Morg. Journ. Cincinnati Soc. Nat. Hist.
16: 1381. 1894.

Distrisution: Probably cosmopolitan.

Sr. Josepu: 1, in wet leaf litter along road, 2884; 2, in wet leaf litter
along road, 2778, associated with Didymium crustaceum (2777). Sr.
Pau: 7, on wet, rotting plant debris on ground, 2713, associated
with Didymium squamulosum (2712); on dead leaves in litter pile,
2822 and 2829, the latter with gold-colored peridia; Brantridge
Est., 4 mile EK. of Pont Cassé, elev. ca. 2,000 ft., on bark of decaying
twig among roadside herbs, 21/29 (poorly developed). In my col-
lections as in those cited by Lister (17), the capillitium is not con-
spicuously paler near the columella.

Macbrideola martinii (Alexop. & Beneke) Alexop. Mycologia 59: 114. 1967.

Disrrisution: Antilles (Jamaica, Dominica).

Sr. Josepu: 1, on bark of large, indeterminate tree (?Jnga) behind
“cabana” at Clarke Hall swimming area, in moist chamber, 2558,
teste C. J. Alexopoulos; the second find of this species, which was
discovered in Jamaica.

Stemonitis axifera (Bull.) Macbr. No. Amer. Slime-Moulds, 120. 1899.

DisrriBution: Cosmopolitan.

Sr. Josepnu: 1, near underside of shaded, decorticate, moist, rotten
log, 2948, with sporangia 5 mm. tall, and spores 4-7 in diameter; 3,
near underside of rotting log, 2895a, associated with 1 sporangium of
Physarum psittacinum (28956) (sporangia pale-ferrugineous, 6 mm.
tall, containing spores 5—6yu in diameter); ca. 1 mile N. of Clarke Hall,
elev. ca. 250 ft., on fairly dry, exposed, rotten log, 2100. St. Davin:
8, on wood of rotting log, 2/55; 15, on rotten logs by river and in
banana plantation, 2735. St. Pauu: 2 miles NE. of Springfield Est.,
elev. ca. 1,800 ft., on underside of very rotten, moist log in pasture,
2838. The sporangium and spore measurements of 2948 and 25954
place these two collections on the border of S. smithii.

Stemonitis smithii Macbr. Bull. Nat. Hist. Univ. Iowa 2: 381. 1893.

Distrisution: Probably cosmopolitan.

Sr. Josep: 3, on rotting log, 2893. Sr. Pauu: 10, on wood of
rotting log, 2271, 2272; sporangia pale reddish brown, subclavulate,
3-4 mm. tall; spores 5-6y diameter. The numerous forms intergrading
with S. acifera suggest that S. smithit may well represent a depauperate
condition or diminutive form of S. axifera rather than a distinct
species, although the possibility of two extensively hybridizing species
is not ruled out.

Stemonitis flavogenita Jahn, Verh. Bot. Ver. Brandenburg 45: 165. 1904.

DistriBution: Probably cosmopolitan.

Sr. Josepn: 1, near underside of rotting log, 2360. (This identifi-
cation is somewhat doubtful. The surface net is rather stout and lacks

FARR—MYXOMYCETES FROM DOMINICA 425

free ends, but the meshes are too small for S. splendens; the capilitium
is strongly webbed). St. Pauu: 4, on underside of decaying log, 2269;
sporangia very slender, clustered; 7, on dead leaves in litter pile, 2944.
Stemonitis fusca Roth, Mag. Bot. Romer & Usteri 1°: 26. 1787.

DIsTRIBUTION: Cosmopolitan.

Sr. JosePH: 2, on bark and splinter on decaying log, 2227; 3,
near underside of rotting log, 2672; on rotting plant debris and on
wood, 2898; on dead leaf, 2900, partly associated with Physarum
pulcherripes (2901) (spore markings faint); on cut end of wet, de-
corticate, rotting log, 3019. Sr. ANprRew: Pointe Baptiste, elev. ca.
100 ft., on bark near underside of fairly exposed, damp, rotten log,
2257 (spores slightly smaller than usual); ca. 1 mile W. of Porte-La-
Fin, abundant laterally on bark of rotting log, 2350. St. Davin: 6, near
underside of rather wet, decaying log, 2/83; 15, on rotten logs by
river and in banana plantation, 2734; 16, on bark of decaying shaded
log, 2314 (surface net weakly developed); on decorticate, rotting log,
2318. The spore markings of nos. 2734 and 2898 approach those of
S. virginiensis Rex.

Stemonitis herbatica Peck, Ann. Rep. N.Y. State Mus. 26:75. 1874.

Reported by Lister (17).

Distrisution: Probably cosmopolitan.

Stemonitis hyperoptera Meyl. Bull. Soc. Vaud. Sci. Nat. 52:97. 1918.

Distripution: North and South America, Europe, Antilles,
Japan.

Sr. Davin: 8, on wet, rotten, decorticate log, 2152. Sr. Paut:
10, on bark near underside of rotting log, 2277 (spores 4—-5y).
Stemonitis nigrescens Rex, Proc. Acad. Phila. 1891: 392. 1891.

Distripution: USA, South America, Wales, Antilles.

St. JoserH: 3, on side of rotten log, 2667 (surface net large-
meshed).

Stemonitis splendens Rost. Monog. Sluzowce (Mycetozoa), 195. 1874.

DistTRIBUTION: Cosmopolitan.

Sr. Josrpy: 1, on underside of rotting log, 2482, with meshes of
surface net 8-40, in diameter (average 20u), but mostly less than 20x.
Sr. Perer (border of Sr. Jonn): 12, on underside of rotten logs,
2378. St. Davip: Rosalie Agric. Exp. Sta., 1 mile NE. of Rosalie,
near junction with road to St. Sauveur, on bark and wood of decaying
branch, 2140; on fairly exposed rotting log in shade, 2321. No. 2140
consists of fuscous, stunted sporangia borne in groups of dense
fascicles; however, the columellae are tortuous above, dissipating
below the apex, and other microscopic characteristics likewise are
typical for the species. No. 2321 is a poorly developed but interesting
collection approaching S. confluens Cke. & Ell. St. Grorce: 9, on

426 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

underside of rotting log in citrus orchard, 2448; Roseau Botanic
Gardens, on top of decaying hardwood log, 2635.

Stemonitis webberi Rex, Proc. Acad. Phila. 1891: 390. 1891.

DistrriBution: Probably cosmopolitan.

St. Joun: 4, laterally on bark of decaying log, 2970; along border
between W. and E. Cabrits, elev. ca. 100 ft., on side of rotting log,
2964. St. ANpREw: In banana plantation ca. 100 yards inland (W.)
from Pagua Bay, on burned log, 2953. Sv. Davin: 15, on rotten logs,
2732, 2739; Baut Sable Bay, S. of Ravine Soudé, on shaded end of
fairly exposed rotting log at S. end of black sand strip, 2319. St. Pat:
7, on side of exposed log at edge of banana plantation, 2942.

Order PHYSARALES

Badhamia affinis Rost. Monog. Sluzowce (Mycetozoa), 143. 1874.

DistriBution: Cosmopolitan.

Sr. Joun: Along border between W. and E. Cabrits, elev. ca. 100 ft.,
on bark of decaying log, 2962; the specimen fits the species in all
characters except for the unusual length of the stalks (up to nearly
1.5 mm.).

Badhamia sp. (B. ?affinis)

St. Joun: 4, on moss on rotting log, 2973. The fruiting bodies of
both Badhamia collections are more or less completely dehisced, so
that their original shapes are not clearly evident. The capillitium of
no. 2962 is typical, reticulate, and the spores are blackish in mass,
smoky brown with many showing a pale dehiscence line by trans-
mitted light, globose to ovoid or ellipsoid, densely spinulose, and mostly
14-16u in diameter. No. 2973 fits the description of var. orbiculata
(Rex) G. List. (19; 15), reported by Lister from Antigua and con-
sidered by Martin (20) a synonym of B. affnis, although it does not
compare well with some authentic specimens of var. orbiculata. The
fruiting bodies (usually only their basal halves remaining) are sessile,
probably sporangiate and plasmodiocarpous, and apparently somewhat
flattened, as indicated by the single layer of parallel white spikes
constituting the capillitium of each; the spores are distinctly violaceous
brown in mass, light brown by transmitted light, globose, spinulose,
and 10-12, in diameter, which is below the size range generally known
for B. affinis. Both collections are white, without any red or brown
tinges even near the sporangial bases; they differ conspicuously from
each other in habit, capillitium, and spore characteristics and are
probably not conspecific.

Badhamia nitens Berk. Trans. Linn. Soe. 21: 153. 1853.

Reported by Lister (17).
Distrisution: USA, Europe, Antilles, Asia, S. Africa.

FARR—MYXOMYCETES FROM DOMINICA 427

Badhamia panicea (Fr.) Rost. in Fekl. Jahrb. Nassau Ver. Nat. 27-28: 71. 1873.

Reported by Lister (17).

Disrrisution: North and South America, Europe, Dominica.
Craterium aureum (Schum.) Rost. Monog. Sluzowce (Mycetozoa), 124. 1874.

DistRIBUTION: Cosmopolitan.

Sr. Davin: 8, on dead leaf in wet leaf litter at edge of freshly cut
rain forest, 2913; sporangia yellow, subglobose to turbinate, with
small but distinct, thickened basal cup.

Craterium leucocephalum (Pers.) Ditm. in Sturm, Deuts. Fl. Pilze 1:21. 1813.

DistrisuTION: Cosmopolitan.

Sr. JosEern: 3, on dead leaf in litter pile, 3017. St. Joun: Northern
outskirts of Portsmouth, along road to Cabrits, on dead coconut
leaflet in litter pile, 2957. St. Pauw: 7, on dead banana leaf in litter
pile, 2835, teste G. W. Martin.

Diderma chondrioderma (dBy. & Rost.) G. List. in List. Monog. Mycetozoa ed.
3: 258. 1925.

Distripution: USA (Western and Midwestern), Europe, Asia,
Dominica.

Sr. Paut: 10, on decaying log, 2428; fructification subsessile, con-
sisting of several sporangia and 1 elongated plasmodiocarp; capillitium
mostly slender.

Diderma effusum (Schw.) Morg. Journ. Cincinnati Soc. Nat. Hist. 16: 155. 1894.

DistriBuTION: Cosmopolitan.

Sr. Josep: 1, on dead coconut leaf sheath still attached to young
plant, 2758; 2, in moss on deeply shaded, decaying log, 2228. Sr. LUKE:
South Chiltern House, elev. ca. 1,200 ft., on bark of Calophyllum sp.
in moist chamber, 2981. Sr. Davin: 8, on leaf, twig, and rotten log,
2160; on bark of decaying twig, 2417; on dead leaves, 2424; on decay-
ing banana sheath, 2466; on living fern leaf and dead phanerogamic
leaf, 2658; and on decaying, mossy log, 26596, associated with Trichia
favoginea (2652a) ; on living moss and small leaves on rotten log, 2747
(poor); no. 2160 consists of extensive, sheetlike, perforated plasmodio-
carps. St. Pauw: 7, around base of living banana plant and on nearby
rotting debris, 2696.

Diderma hemisphaericum (Bull.) Hornem. Fl. Dan. 33:13. 1829.

DistriButTion: Cosmopolitan.

Sr. JosepH: 1, on decaying bamboo cane and on dead leaves in wet
litter pile along road, 2880; on dead leaf, 2950; 2, on dead leaf in wet
litter pile near road, 2780. St. GroraE: 9, from moist chamber
containing a -torange plasmodium in leaf litter, 2366a, associated
with Arcyria cinerea (2866b). The identity of the plasmodium was not
ascertainable; Martin (1949) reports white or nearly white plasmodia

313-143—68——3

498 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

for both the Diderma and the Arcyria. Sr. Pauw: 7, on rotting plant

debris on ground, 2711; on dead leaves in litter pile, 2827.

Diderma rugosum (Rex) Macbr. N. Amer. Slime-Moulds, 105. 1899.
Reported by Lister (17).

Distripution: USA (Eastern and Midwestern), Europe, Antilles,
Asia.

Diderma spumarioides (Fr.) Fr. Syst. Mycol. 3: 104. 1829.
DisTRIBUTION: Cosmopolitan.

Sr. Davin: 8, on living herbaceous leaves among debris on ground,
2284; peridium areolate; spores unevenly warted, 9-11 in diameter.
These traits agree essentially with Lister’s (17) description of Dominica
material.

Didymium clavus (Alb. & Schw.) Rab. Deuts. Krypt. Fl. 1: 280. 1844.
DisrriBution: Cosmopolitan.

St. AnpreEw: “Hill,” SW. of Pagua Bay, elev. ca. 700 {t., on under-
side of rotting log in banana plantation, 3004; sporangia single and
discoid, or fused in groups of 2-4, with stalks remaining distinct.
St. Davin: 8, on underside of wet, rotten log, 2546 (sporangia widely
scattered). St. Pauu: 7, on bark of rotting log, 2832,

Didymium comatum (List.) Nann.-Bremek. Proc. Nederl. Akad. Wetens. Ser.

C, 69: 361. 1966.

Distrisution: North America (Pa., Que.), Great Britain, Japan,
Dominica.

St. Pauu: 7, on dead leaf in litter pile, 2830; a well-developed collec-
tion consisting of sporangia and plasmodiocarps and exhibiting the
characteristic elastic capillitium and spore markings described by
Nannenga-Bremekamp.

Didymium crustaceum Fr. Syst. Mycol. 3:124. 1829,

Disrrisution: North and South America, Europe, Antilles, Hawaii,
Japan.

St. JosepH: 2, in wet roadside litter pile, 2777, associated with
Lamproderma scintillans (2778). Spores dark brown, densely and
prominently spinulose, 10-14 in diameter; outer peridial crust almost
completely disappeared, perhaps because of the wet environment;
inner peridium loosely covered with yellowish lime crystals; columellae
large, whitish or yellow, apparently formed by the strongly invaginated
sporangial bases.

Didymium difforme (Pers.) Gray, Nat. Arr. British Pl. 1:571. 1821.
Disrrisution: North and South America, Europe, Antilles, Japan.
St. Joseru: 3, on leaf litter and rotting banana debris on ground,

26656, associated with D. squamulosum (2665a).

FARR—-MYXOMYCETES FROM DOMINICA 429

Didymium intermedium Schroet. in P. Henn. Hedwigia 35:209. 1896.

Distripution: USA (Mo.), Central and South America, Antilles,
Japan.

Sr, Anprew: 13, on very wet, rotting log in rain forest, 2999;
sporangia white, fascicled; stalks straw-colored.

Didymium iridis (Ditm.) Fr. Syst. Mycol. 3:120. 1829.

DistrisutTion: Cosmopolitan.

Sr, JosepH: 1, on living and dead leaves and stems lying among
decaying cacao fruit on ground, 2763 (see comments under D. nigripes) ;
slightly E. of Coulibistri, on outer surface of cacao fruit and on large
pebble inside, 2097 (a poorly developed fruiting with prominent, globose,
whitish columellae). St. Perer (border of St. Joun): 12, on dead leaf
buried in litter, 2373 (poor). St. Pau: 10, on bark of Cecropia
peltata L. in moist chamber, 2769; fruited on bark, filter paper, and
glass.

Didymium minus (List.) Morg. Journ. Cincinnati Soc. Nat. Hist. 16:145. 1894.

Distripution: Cosmopolitan.

Sr. Davin: 8, on living banana plant, 250/a (sporangia stipitate
to sessile).

Didymium nigripes (Lk.) Fr. Syst. Mycol. 3: 119. 1829.

DistrisuTION: Cosmopolitan.

Sr. Josprx: 1, on living and dead leaves and stems among decaying
cacao fruits on ground, 2761. Sr. ANprew: 18, on dead leaf in litter
pile at edge of rain forest, 2997; sporangia nearly limeless; spores very
unevenly warted, as in Mig. 102g of Lister (19). St. Davip: 8, on
dead leaf sheath, 2/54; on living and decaying banana sheaths on
eround, 22916, abundant; on dead moss and leaves, 2425 (poor); on
dead leaf, 2471; on underside of basidiocarp of Hymenochaete (2532;
probably H. berkeleyana [Mont.] Cke., fide P. L. Lentz), 2541 (very
poor); on living leaves of Carludovica plumiert Kth., 2653; on living
petiole of Colocasia sp., 2744; on living banana plant, 28016; on rotting
leaf litter, 2928 (spores 10u in diameter). St. GeorGE: 9, on living
leaves of small, shaded herb, 2198. St. Pauu: 7, on dead leaf on
ground, 2704, on dead grass leaf in debris pile, 2707. Nos. 2763 (D.
iridis) and 2761 were growing together; they are similar in micro-
scopic structure and macroscopically as well, except that the former
specimen has pale stalks and white columellae; the latter collection
exhibits dark columellae and stalks darker than those of 2763, but
sometimes becoming translucent in the upper portion. Dr. Martin,
who examined and discussed (personal correspondence) 2763, 2761,
and 2801b, believes that these stalk variations may be governed by
different maturation rates. He also found the spores of 2 adjacent
sporangia of no. 28016 to differ in color and size, those of 1 sporangium

430 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

being 8-9y and dark, and those of the other, 10-11 and much paler.

The relationship between D. iridis and D. nigripes is still contro-
versial. Laboratory culture work tends to support the existence of 2
distinct species (C. J. Alexopoulos, G. W. Martin, personal com-
munications), but field collections provide numerous connecting
forms. These may represent another example of 2 strongly hybridizing
species.

Didymium squamulosum (Alb. & Schw.) Fr. Symb. Gast. 19. 1818.

DistrrBputTion: Cosmopolitan,

St. Josepu: 1, on wet, dead leaves piled by roadside, 2883, 2885,
2887, 2889; 2 (Cocoa Centre), on wet, decaying coconut leaflet on
ground, 2550 (sparse) ; on dead banana leaf, 2774; on wet, dead leaves
by roadside, 2775, associated with Physarum melleum (2776), and
£779; 3, on leaf litter and rotting banana debris on ground, 2665a,
associated with Didymium difforme (26656); on dead leaf, 2594.
St. Anprew: 5, on rotting banana leaf on ground, 2617 (malformed).
St. Davin: 8, on decaying twig and dead banana leaf, 2651; on
rotting banana debris, 2796; in rain forest leaf litter, 2932. St. Paut:
7, on rotting banana leaves on ground, 2705, 2706, 2712, associated
with Lamproderma scintillans (2718); on leaf litter, 2828; in piles of
decaying banana leaves, 2874. This species occurs in two distinct
phases. Nos. 2828, 2883, 2887-2889, 2550, 2775, and 2932 are char-
acterized by globose sporangia covered by a powdery lime coating,
and globose or somewhat depressed columellae; the remaining speci-
mens exhibit discoid sporangia with strongly rugose peridia, and flat
white columellae. No. 2594 has globose sporangia, but with strongly
rugulose peridial lime; the columellae are formed by the deeply
umbilicate sporangium bases. This exceedingly variable species
appears to be much more common on this island than in Jamaica.
Fuligo cinerea (Schw.) Morg. Journ. Cincinnati Soc. Nat. Hist. 19:33. 1896,

DistriBuTION: Cosmopolitan.

Sr. Joswpu: Slightly E. of Coulibistri, on rind and inside of decay-
ing cacao fruit, 2096. Sr. Grorcy: Chapara Est. (near road junction),
S. corner of parish, elev. ca. 1,550 ft., on rotting fruit of Artocarpus
sp. (A. ?communis) on ground, in moist chamber, 2767; the plasmodium
turned from tan through pinkish to white before fruiting.

Fuligo septica (L.) G. H. Weber in Wiggers, Prim. Fl. Holsat., 112. 1780.

DisrrisuTiIon: Cosmopolitan.

St. JoserH: 2, on decaying coconut logs, 2493-2496; 2493 with
more or less cream-colored cortex, others with rust-colored cortex and
yellow capillitial lime; 2 (Cocoa Centre), on side of rotting log, 2548,
brown and largely moldy. Sr. Joun: 4, on top of rotting log, 2969;
aethalium salmon-colored with darker or yellow margin. St. ANDREW:
5, around thin herbaceous stems and roots hanging from underside

FARR—MYXOMYCETES FROM DOMINICA 431

of rotten log, 2588 (reported from rootlets of higher plants also by
Duss, 8); aethalia pale tan or orange. St. Davin: 8, on roots and on
very rotten wood and debris in open, hollow stump, 2655, with bright
yellow, thick, rugose but fragile cortex and abundant, somewhat
elastic, yellow capillitium; on wet, rotting log in disturbed rain forest,
in somewhat open spot near trail, 2804, resembling 2655 and associated
with Physarum pulcherripes (2803); on top of rotting log, 2933; a
large fructification (ca. 3.52.5 inches) with cortex salmon-colored
above, yellow below. St. GrorcE: Roseau Botanic Gardens, on
decaying hardwood log, 2634; Chapara Est., S. corner of parish, near
road junction, elev. ca. 1,550 ft., in banana plantation by creek,
girdling decaying stem of Bambusa vulgaris Schrad. 3-5 inches above
root crown, and on stem of woody vine, 2444; a poorly developed,
whitish specimen covered with a nectriaceous fungus. Sr. Pavu:
1.5 miles E. of Pont Cassé, elev. ca. 1,800 ft., on top of rotten log in
rain forest, 2/11; cortex hard, light brown. The brilliant yellow color
and, especially, the more or less strongly elastic capillitium, of nos.
2655 and 2804, bring to mind Erionema aureum Penz. Lister (17)
reports 1 yellow aethalium of F. septica.

Physarella oblonga (Berk. & Curt.) Morg. Journ. Cincinnati Soc. Nat. Hist.

19: 7. 1896.

DistriBuTION: Cosmopolitan.

Sr. Josepu: 1, near underside of decaying coconut log among
moss and living leaves, 2759 (possibly polyploid, since the sporangia
are almost twice the normal size with correspondingly thick stipes;
microscopic features are typical in size and appearance); 2 (Cocoa
Centre), on underside of rotten logs, collected as yellow plasmodium,
fruited 16 days later, 2765; 3, on underside of rotting log, 2616.
Sp, Perer: 11, on underside of wet, rotten, shaded log, 2080. St.
Davin: 8, on underside of wet, rotting log, 2146, teste G. W. Martin,
2512 (a freshly formed fruiting in excellent condition). Sr. Pauw: 7,
on underside of wet, rotting logs, collected as yellow plasmodium,
fruited in moist chamber, 2865 (a beautiful, abundant collection).

Physarum aeneum (List.) R. E. Fr. Ark. Bot. 1: 62. 1903.

Distrisution: USA (Eastern and Midwestern), South America,
Antilles, Hawaii, Asia.

Sr. Davin: 8, near base of living banana plant, 24/6; plasmodio-
carps brown, with iridescent inner peridium; capillitium whitish.
Sr. Pau: 7, on dead leaves in litter pile, 2836, 2877, 2945, 2946.

Physarum auriscalpium Cke. Ann. Lyc. N.Y. 11: 384. 1877.

DistrisuTion: USA, South America, Antilles.

Sr. JosepH: Macoucheri Est., ca. 4 mile inland from W. coast,
on bark of Mangifera indica in moist chamber, 2483; spores 11—-l4y
in diameter, densely spinulose. Sr. Davin: 8, on decaying banana

432 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

sheath, 2469. Sr. Luxe: Pointe Guignard, on wood from exposed
log near beach, in moist chamber, 2861 (fruiting abundantly on wood,
filter paper, and glass).

Physarum bogoriense Racib. Hedwigia 37:52. Feb. 18, 1898.

DisrrisuTion: Probably cosmopolitan; pantropical.

Sr. Davin: 8, on decaying twigs and leaf sheaths, 2283; on decaying
leaf, 2291a, associated with Didymium nigripes (2291b); 2426 (with
Didymium nigripes sporangia growing on some plasmodiocarps) ; on
base of living banana plant, 2660; on dead banana leaf, 2743; on dead
leaf in rain forest, 2794. Sv. Pau: 7, on dead banana leaf, 2709; on dead
leaf, 2820; in pile of wet, dead leaves, 3011. All collections appear
typical for the species, but Lister (17) reports sporangial color varia-
tions from reddish brown to buff, bright yellow, or nearly white.

Physarum bubalinum Farr (see p. 412).

Physarum cinereum (Batsch) Pers. Neues Mag. Bot. 1:89. 1794.

DistriBuTIoN: Cosmopolitan.

Sr. JosEPH: 2, on living and dead leaves and stems buried in debris
pile in cacao-banana plantation, 2786; N. side of Brook Hill Est.,
behind Clarke Hall, on living Paspalum conjugatum Berg. leaves,
collected by J. L. Cunningham 66-393-D, det. M. L. Farr. Sr. Davi:
8, on living banana plant, 2795. Sr. Paut: 7, in pile of decaying leaves,
2824.

Physarum compressum Alb. & Schw. Consp. Fungi, 97. 1805.

DistRIBUTION: Cosmopolitan.

Sr. Joseru: 1, inside decaying cacao fruit, 2080; on and in decaying
cacao fruits and on dead leaf, 2102; 2126 (sporangia liquid when
collected, ripened in collecting basket); 2760; the last 3 collections
represent the convolute form. 2, on dead leaf in debris pile, 2787;
3, on decaying leaf on rotten log, 2666; on moss on moist, rotting log,
“671 (sporangia fused in fascicles of 2 or 3); slightly E. of Coulibistri,
in and on moist, shaded, decayed cacao fruits and on dead leaf,
collected C. and B. Evans and M. L. Farr, 2098, 2099; spores of the
latter specimen, cultured in the laboratory by C. J. Alexopoulos,
yielded clustered, sessile, contorted and lobed (sometimes partly
immature) fructifications which, upon subculture by Charles Mims,
produced a crop of individual sporangia typical for P. compressum.
Sr. Anprew: 13, on dead leaf in rain forest, 2993 (a depauperate
voucher specimen). St. Davin: 8, on decaying plant fibers, 2418; on
wet, dead leaf and twig, 2543 (convoluted form similar to 21 26); on
dead banana leaf, 27/8; 6, on stalk of decaying banana, inflorescence,
2727; 15, on dead leaf and rotting twig, 2733 (both 2727 and 2733
contain more or less typical sporangia mixed with fascicled and

FARR—-MYXOMYCETES FROM DOMINICA 433

convoluted fruit bodies); ca. 1 mile N. of St. Sauveur, on decaying
leaf debris by rotting roadside log, 2167. St. Pauw: 10, on decaying
plant debris, 2279 (considered by Dr. Martin as an “excellent example
of the species’). An early collection of this species reported by Lister
(17) consisted of ‘compressed simple or lobed sporangia on gray
stalks.”
Physarum didermoides (Pers.) Rost. Monog. Sluzowce (Mycetozoa), 97. 1874.
DistRIBUTION: Cosmopolitan.
Sr, Davin: 8, on decaying filmy fern debris on rotting log, 2420;
on living filmy fern and associated leaf litter on rotting log, 2467,
2468; on tree fern roots and debris on wet, rotting log, 2662.

Physarum echinosporum List. Journ. Bot. 37: 147. 1899.

Distrisution: Antilles, Indonesia.

Sr, Josep: 1, on dead leaves in wet litter pile along road, 2582,
2949.

Physarum galbeum Wing. ex Macbr. No. Amer. Slime-Moulds, 53. 1899.

Distrisution: North and South America, Europe, Antilles.

Sr, Davip: 8, near underside of moist, very rotten log, 2652; a
good, fairly typical specimen with bright yellow, globose sporangia,
floriform dehiscence, and capillitium containing numerous elongated
and branching yellow lime knots. Two additional fruitings observed—
a bright yellow one from St. Davin and one with orange stalks from
Sr. Perer—were too dilapidated to retain.

Physarum lateritium (Berk. & Rav.) Morg. Journ. Cincinnati Soc. Nat. Hist.
19: 23. 1896.

Distripution: North and South America, Europe, Antilles, Asia,
Hawaii.

Sr. Pau: 7, on dead leaf in litter pile, 2947; a rather battered
fructification containing mostly sessile sporangia with limeless, iri-
descent peridia enclosing capillitium fading from bright orange to
pale yellowish, and spores 9-10 in diameter, finely warted, with
clusters of more prominent warts.

Physarum leucophaeum Fr. Symb. Gast., 24. 1818.

Distrisution: North and South America, Europe, Antilles, New
Zealand.

Sr, Joszpu: 1, on bark of Inga sp. in moist chamber, 2859a, associated
with Licea operculata (28596) (stalks dingy-yellowish). St. Marx:
Summit of Scotts Head, elev. ca. 100 ft., on bark of Tabebuia pallida
(Lindl.) Miers in moist chamber, 2562 (somewhat abnormal). St.
Luxe: South Chiltern House, elev. ca. 1,200 ft., on bark of Mangifera
indica in moist chamber, 2723 (spores slightly atypical, 10-14p,
spinulose).

434 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Physarum ‘leucopus Lk. Ges. Nat. Freunde Berlin Mag. 3:27. 1809.

DistRIBUTION: Cosmopolitan.

Sr. Perer: Summit of Morne Diablotins (end of 2d ridge), elev. ca.
4,700 ft., on stem of Lobelia stricta in moist chamber, 3033a; fruited
somewhat abnormally on glass, destroyed during transit. This species
is known from the Blue Mountains and Port Royal Mountains in
Jamaica and apparently prefers the more temperate environments.
The Dominica record must be regarded as uncertain for the present.
Physarum melleum (Berk. & Br.) Mass. Monog. Myxogastres, 278. 1892.

DistrriIBuTION: Cosmopolitan.

St. Josepn: 1, on decaying twigs among rotting cacao fruits, 2762;
on dead leaves, 2886 (a depauperate fruiting partly associated with
Didymium iridis); 2, on dead leaves, 2776, associated with Didymium
squamulosum (2775) ; 3, on dead leaf, 2896, teste G. W. Martin (stalks
tawny to dark yellow, peridia dark yellow); 2897, (stalks white ). Sr.
Anprew: 13, on dead leaf on rain forest floor, 2995. Sr. Davin: 8,
on decaying, wet banana leaf after 4 days of continuous rain, 2609;
on decaying log, 2798 (resembling 3026); among moss on side of
rotting log, 2929; near north branch of Ravine Deux Dleaus, S. of
Bois Diable Ridge, elev. ca. 1,100-1,200 ft., on dead leaf in litter pile
in banana plantation, 3026, 8027; both with whitish, slightly turbinate
sporangia and strongly developed, white columellae. Sr. Parrick:
Laronde, on bark of indeterminate tree in moist chamber, 3034
(fruited on glass). Sr. Pauu: 7, on dead leaves in litter pile, 2837,
teste G. W. Martin (a specimen with well-developed columellae and,
often, additional lime aggregations in the center of the sporangia); in
piles of decaying banana litter, 2875, 2876. This species displayed
considerable variation in sporangial color and shape and in develop-
ment of columella, as also noted by Lister (17) for Antiguan material.
Physarum notabile Macbr. No. Amer. Slime-Moulds ed. 2, 80. 1922.

Distrisution: North and South America, Europe, Antilles.

Sr. Luxe: South Chiltern House, elev. ca. 1,200 ft., on bark of
indeterminate tree in moist chamber, 2722a.

Physarum nucleatum Rex, Proc. Acad. Phila. 1891:389. 1891.

Distrisution: Pantropical; probably cosmopolitan.

St. JosePH: 3, near underside of wet, decorticate, rotten log, 2612;
“670; on underside of wet rotting log and on adjacent living weeds,
3021, very abundant. St. Prrer (bordering Sr. Jonn): 12, on under-
side of rotting logs, 2375, 2377; on decaying small roots, 2376. Sr.
AnpDREWw: 5, 2587, (a very moldy voucher specimen). Sr. Davin: 8,
on upper side of wet, decaying log, 2149; on decaying twig, 2157;
on inner bark near top of decaying log, 2472; on loose inner bark of
decaying twig, 2444; on rotting plant debris on decayed log, 2746;
on wet, rotting log, 29/7, 2931. St. Pauu: 7, on living leaf and adja-

FARR—MYXOMYCETES FROM DOMINICA 435

cent debris and on part of rotting log, 2943; 10, on wood of decaying
twig, 2274; on decorticate, rotten log, 2429, 2430. This species oc-
curred on the island in 2 forms: (a) typical delicate sporangia with
globose, white pseudocolumellae which, however, readily fell out of
the dehisced sporangia; (b) a robust (?polyploid) variant, represented
by specimens 2149, 2274, 2376, 2429, 2480, and 2746, Although of
the same structure and color as typical material, the robust fruitings
were much larger, lacked pseudocolumellae, but exhibited an abundant
capillitium with numerous, large, rounded, white lime nodes.

Physarum nutans Pers. Ann. Bot. Usteri 15:6. 1795.

DistTRriBuTION: Cosmopolitan.

Sr. JosepH: Mero, on bark of Lagenaria siceraria in moist chamber,
2596, associated with Cribraria violacea (2595); spores 9-12u in
diameter. Sr. Davin: 8, on underside of decaying log, 2423, abundant
but more or less moldy; after 4 days of continuous rain, 2511 (limeless) ;
on bark of rotting log, 2799; 6, near underside of moist, decaying log,
2134. Sr. Pauw: 10, on rotting twig, 2276 (battered).

Physarum oblatum Macbr. Bull. Nat. Hist. Univ. Iowa 2:384. 1893.

Disrrisution: Cosmopolitan.

Sr. JosepH: 2, on decaying coconut log, 2491 (faded).

Physarum penetrale Rex, Proc. Acad. Phila. 1891:389. 1891.

Disrripution: North America, Europe, Antilles, Asia, Africa.

Sr. Davin: 6, on rotting log, 2205; sporangia subovoid, yellow,
with very delicate, orange stalks.

Physarum polycephalum Schw. Schr. Natur. Ges. Leipzig 1:63. 1822.

DisTRIBUTION: Cosmopolitan.

St. ANDREW: 5, on underside of rotting log in roadside banana
plantation, 2618; Porte-La-Fin, on bark near underside of rotting
log, 2349, an abundant fruiting. St. Pauw: 7, on dead leaf on ground
2884; on dead leaves, stems, etc. in litter pile, 2838, 2839 (abundant).
The capillitium of 2834 strongly suggests P. rigidum (G. List.) G. List.
Nos. 2349, 2834, and 2839 consist of unfused sporangia and represent
“var, obrusseum (Berk. & Curt.) List.”; no. 2838, as noted by Dr.
Martin, contains fused as well as simple sporangia. Dr. Alexopoulos
obtained typical P. polycephalum fruitings from laboratory cultures
of 2349 (private communication). This evidence obviously refutes
previous recognition by several investigators (including me) of the
variety obrusseum.

Physarum psittacinum Ditm. in Sturm, Deuts. Fl. Pilze 1:125. 1817.

Disrrisution: USA, Europe, Japan, Dominica.

St. Josppu: 3, near underside of rotting log, 28956 (1 sporangium,
associated with Stemonitis azxifera [2896a]). St. Davin: 8, on under-
side of decaying, moist log, 2147, 2148; on end of wet, rotting log in

436 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

somewhat open spot in rain forest, 2803, associated with Fuligo
septica (2804). The sporangia of 2147 are globose, rusty orange, some-
times limeless and iridescent near the base; the stalks are limeless,
red or orange brown; the peridial lime scales are orange and the
capillitial lime, orange to whitish; 2/48 is a moldy specimen having
yellowish inner capillitial lime nodes, dark brown stalks, and brownish-
iridescent peridia with red lime scales. No. 2803 is a collection inter-
mediate between P. psittacinum and P. pulcherripes. Dr. Martin
favors the latter identification, but, after restudying the BPI speci-
men (which is more plentiful and in better condition than the IA
duplicate), I am inclined to place it tentatively in P. psittacinum
because of the red stipes which are not calcareous throughout, but
often partly covered with a thin layer of lime, and because of the rather
large, angular lime nodes of the capillitium. Columellae, furthermore,
appear to be lacking, but this is true also of Dominica collections
considered to be P. pulcherripes. The yellow lime of this specimen
imparts an atypically pale aspect to the sporangia, but this may be
due to fading.

Physarum pulcherripes Pk. Bull. Buffalo Soc. Nat. Sci. 1:64. July 1873.

Distrisution: North America, Panama, Ireland, Dominica.

St. Josepu: 3, on decaying, decorticate log and dead leaf, 2901,
partly associated with Stemonitis fusca (2900); an abundant, well
developed fruiting with yellow, calcareous stalks and deep golden-
or rust-colored peridia. Sr. Davin: 6, on rotting log, 2204. Sv. Paut:
7, on underside of wet, rotting log, collected as big, yellow, “knotty”
plasmodium, 2772. This species appears to be closely allied to P.
psittacinum, differing from the latter (according to the standard mono-
graphs) by having calcareous stalks slightly prolonged inside the
sporangial bases into short columellae, and small, rounded (rather
than large, angular) capillitial nodes. Both species appear to be capable
of considerable color variation, as judged from material examined.
Since the spores of the 2 species are similar and since no columellae
were found in any Dominica collections cited, stalk and capillitium
traits were the diagnostic features most strongly relied on. A com-
parison with part of the type of P. pulcherripes revealed that the
capillitia of the latter collection and of all Dominica gatherings are
identical, but also that all contain numerous angular nodes of various
sizes. The stalks of the type are strongly rugulose, bright orange, and
calcareous in the upper half, but smoother, brown, and limeless in
the lower; those of nos. 2772 and 2901 have strongly rugulose, yellow
or whitish, calcareous portions, and slightly slimmer, smoother, brown,
limeless sections; the latter may be apical, basal, or both. No. 2204,
a rather battered collection, has nearly limeless stalks. In all speci-
mens the calcareous and noncalcareous sections of most stalks are
sharply delimited.

FARR—MYXOMYCETES FROM DOMINICA 437

Physarum pusillum (B. & C.) G. List. in Lister, Monog. Mycetozoa ed. 2, 64.
1911.

DisTRIBUTION: Cosmopolitan.

St. Josepu: 1, on bark of Deloniz regia (Boj.) Raf. in front of
Clarke Hall annex, in moist chamber, 2235; on bark of Juniperus
sp. next to Clarke Hall (main bldg.), in moist chamber, 2721 (capil-
litium scanty); 2771, an improperly matured field collection. Sr.
GeorGE: 9, on bark of Clusia plukenetit in moist chamber, 2983),
associated with Lamproderma arcyrionema (2983a).

Physarum roseum Berk. & Br. Journ. Linn. Soc. 14: 84. 1873.
Distrispution: Florida, Brazil, Antilles, Africa, Asia, Oceanica.
Sr. Davin: 1-1.5 miles E. of north branch of Ravine Deux Dleaus,

S. of Bois Diable Ridge, elev. 1,100—1,200 ft., among moss on side of

rotting stump at edge of partly cleared rain forest, 3029 (a very

poor, moldy, but distinctly recognizable specimen).

Physarum rubiginosum Fr. Symb. Gast., 21. 1817.

Distrispution: USA, Europe, Antilles.

Sr. Perer: 11, on bark of fallen branch, 2076; a very small fructi-
fication consisting of short-stipitate sporangia and a small plasmodio-
carp with deep orange lime scales on the peridium and white capillitial
lime; tentatively identified by C. J. Alexopoulos and also by T.
Brooks.

Physarum stellatum (Mass.) Martin, Mycologia 39: 461. 1947.

Distrisution: North and South America, Antilles, Africa, Asia;
probably pantropical.

Sr. Perer: 12, on rotting log in rain forest, collected by G. Brooks,
2383 (moldy). St. Davin: 6, on underside of decaying, wet log, 2206
(good, abundant material). St. Paun: 7, on underside of mossy,
rotten log, 2699 (with large pseudocolumellae); on decaying roots
and other plant debris on ground, 2710; 10, on underside of moist,
decaying log, 2268a (a large fruiting).

Physarum tenerum Rex, Proc. Acad. Phila. 1890: 192. 1890.

DisTRIBUTION: Cosmopolitan.

Sr. AnprEw: 14, on rotting log in swamp-woodland, 2249, collected
as yellow plasmodium. St. Davin: 8, on very rotten log, 2445; collected
as yellow “knotty” plasmodium, ripened overnight in moist chamber.
St. Paut: 7, on side of wet, rotting log, 2703; stalks bright orange in
2445 and 2703. Nos. 2249, 2708, (and 2170, a depauperate collection
from St. Davip, which was later discarded) are of doubtful affinity.
Although these collections fit P. tenerum in every other respect, their
stalks are limeless; 2703, furthermore, has globose pseudocolumellae
similar to those of P. stellatum, but yellow; 2249, while lacking pseudo-
columellae, shows enlarged or massive lime nodes in the center of the
sporangia. The presence or absence of lime in the stalk has generally

438 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

been considered a reliable diagnostic character and is often used to
separate species in this genus; a number of authentic collections
examined (including the type) of P. tenerum all had calcareous stipes.
Since, in general appearance, dehiscence, capillitium, and spores, the
Dominica specimens are indistinguishable from typical members of
this species, they are filed here until more is known about the signifi-
cance and stability of lime content in the stalks.

Physarum viride (Bull.) Pers. Ann. Bot. Usteri 15:6. 1795.

Distrisution: Cosmopolitan.

St. JosepH: 2 (Cocoa Centre), near underside of rotting log, 2598,
teste G. W. Martin (sporangia deep yellow). St. Perer (bordering
St. Jonny): 12, on bark of Dacryodes excelsa in moist chamber, 2846.
St. AnpREw: 5, on bark of Dacryodes excelsa in moist chamber, 2856
(largely moldy). Sr. Davin: 8, on bark near underside of wet, rotting
log, 2159 (a beautiful, deep orange fructification with concolorous
stalks); 2161, collected as yellow plasmodium, which produced, by
the next day, ripe, globose, nodding, yellow-orange sporangia with
small, limeless basal discs and orange stalks; 2542 and 2545, with
same coloration as 2161, collected in slightly immature state. Sr.
GEoRGE: 9, in moist chamber from plasmodium in rotten wood, 2764a.

12.

13.

14,

16.

17.

Literature Cited

. ALEXopOULOS, C. J.

1964. The rapid sporulation of some myxomycetes in moist chamber culture.
Southwest. Nat. 9: 155-159.

. ALExoPouLos, C. J., and Benrekg, E. 58.

1954. Myxomycetes from Jamaica. Trans. Brit. Mycol. Soc. 37: 306-313.

ALEXOPOULOS, C. J., and Brenexg, E. 5S.

1954. A new species of Comatricha from Jamaica. Mycologia 46: 245-247,
figs. 1-2.

. Barngs, R. F.

1963. Myxomycetes from Trinidad. Trans. Brit. Mycol. Soc. 46: 453-458.

. BERKELEY, M. J.

1869. On a collection of fungi from Cuba. Part. 11... Journ. Linn. Soc.
Bot. 10: 341-392 (Myxogastres on pp. 346-350).

. Cirerri, R.

1929. Micoflora domingensis. Est. Agron. Moca Ser. B, Bot., No. 14, 260 pp.
(Myxomycetes on pp. 49-53).

. CirerRI, R.

1961. Mycoflora domingensis integrata. Quaderno Ist. Bot. Univ. Lab.
Critt. Pavia No. 19, 5389 pp. (Myxomycetes on pp. 9-14).

. Duss, A.

1903. Enumération méthodique des champignons recueillis a la Guadeloupe
& ala Martinique. 94 pp. (also publ. 1904 as Fl. Crypt. Antilles
frang., Champ., pp. 210-304).

. Farr, M. L.

1957. A checklist of Jamaican slime-moulds (myxomycetes). Bull. Inst.
Jamaica, Sci. Ser. No. 7, 67 pp., 4 figs., 1 table, 1 map.

. Farr, M. L.

1958. Taxonomic studies in the myxomycetes. I. The Trichia favoginea
complex. Mycologia 50: 357-369.

. Farr, M. L.

1967. Notes on myxomycetes. Mycopath. Mycol. Appl. 31: 305-313, figs. 1-9.

FEentTEM, ARLIN D.

1960. Commercial geography of Dominica. Office Naval Res. Tech. Rept.
No. 5, 18 pp., offset, illus.

GonzaLes Fracoso, R., and Cirerri, R.

1927. Hongos Parasitos y saprofitos de la Republica Dominicana. Ser. 6-10.
Est. Agron. Moca Ser. B, Bot., No. 8, 99 pp. (Myxomycetes on pp.
5-6).

1928. Ser. 16, ibid. No. 13, 17 pp. (Myxomycetes onp.3).

HAGELsTEIN, R.
1932. Revision of the Myxomycetes. Sci. Surv. Porto Rico and Virgin
Islands 8: 241-248.

. HAGELSTEIN, R.

1944. The Mycetozoa of North America. Published by the author. Mineola,
N.Y., 306 pp., 16 pl.

Hover, W. H.

1954. Flora of Dominica. Lloydia 17: 1-238.

Lister, A.

1898. Mycetozoa of Antigua and Dominica. Journ. Bot. 36 (424): 113-122,
pl. 385.

439

440 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

18. ListEerR, A.
1898. Mycetozoa of Antigua. Journ. Bot. 36(430) : 378-379.
19. Listpr, A.
1925. A Monograph of the Mycetozoa, ed. 3 (revised by G. Lister, xxxii+-296
pp., 60 figs., 222 pls., many col.).
20. Martin, G. W.
1949. Fungi, Myxomycetes. No, Amer. Flora. 1(1): 1-151. Bibliog. by H. W.
Rickett, pp. 153-157.
21. Toro, R. A.
1926. Mixomicetos de Santo Domingo. Est. Agron. Haina Ser. B, Bot., No.
6, 7 pp.

U.S. GOVERNMENT PRINTING OFFICE: 1968