580,5

FB

V, 29:1-2

1954/55

cop, 2

REVISION OF

THE HAWAIIAN MEMBERS

OF THE GENUS
TETRAPLASANDRA A. GRAY

EARL EDWARD SHERFF

FIELDIANA: BOTANY

VOLUME 29, NUMBER 2

Published by
CHICAGO NATURAL HISTORY MUSEUM

JULY 29, 1955

NATURAL

HISTORY SURVET

LIBRA/?*

REVISION OF

THE HAWAIIAN MEMBERS

OF THE GENUS
TETRAPLASANDRA A. GRAY

EARL EDWARD SHERFF

Research Associate, Systematic Botany

FIELDIANA: BOTANY

VOLUME 29, NUMBER 2

Published by

CHICAGO NATURAL HISTORY MUSEUM Tfff UB RARV Of fHI
JULY 29, 1955

fiUG 26 1955
HHIWMIWOHUINOIS

PRINTED IN THE UNITED STATES OF AMERICA
BY CHICAGO NATURAL HISTORY MUSEUM PRESS

PS

'.5

Revision of the Hawaiian Members of the Genus
Tetraplasandra A* Gray

INTRODUCTION AND HISTORICAL NOTES

The history of the genus Tetraplasandra begins with the original
description of the genus by Asa Gray (Bot. U. S. Explor. Exped.
727, tab. 94. 1854). A single species was described at that time,
T. hawaiiensis. 1 This, the type basis of the genus, had leaves densely
tomentose beneath and a paniculate inflorescence with umbellate,
racemose, and umbellate-racemose subordinate branches. In 1873,
Wawra (Flora 56: 158) described a second species, T. waimeae,
from the Island of Kauai, a species with glabrous leaves and twice-
compound umbels. As time went on, three additional species were
described from elsewhere than the Hawaiian Islands: T. paucidens
Miq. Ann. Mus. Bot. Lugd. Bat. 1: 4. 1863. East Indies; T.
Koordersii Harms, Ann. Jard. Buitenz. 19: 12. 1904. Celebes; and
T. philippinensis Merr. Philipp. Jour. Sci. 1, suppl. 219. 1906.
Philippine Islands. As these do not come within the scope of this
treatment, they need not concern us here.

In the same volume that contained the original description of
T. hawaiiensis, Gray described interrogatively (p. 726) a Gastonia
oahuensis from the mountains behind Honolulu, alt. 2,000 feet.
Seemann (Jour. Bot. 6: 139. 1868) renamed this Triplasandra
oahuensis, a name that was altered to Tetraplasandra oahuensis in
1898, when Harms (in Engl. & Prantl, Pflanzenfam. 3: 8: 30)
merged the genus Triplasandra Seem, with Tetraplasandra A. Gray.
Prior to Harms' work, however, Hillebrand (Fl. Haw. Isls. 151-155.
1888) had given treatments for Triplasandra and Tetraplasandra.
In the latter, he had included only the two species, Tetraplasandra
hawaiiensis and T. waimeae. In the former, he had retained Tri-
plasandra oahuensis as it had been renamed by Seemann and had
appended to it a var. /3. He had added three new species of his

1 Spelled hawaiensis by Gray but given by certain subsequent authors the
more conventional spelling hawaiiensis.

49

50 FIELDIANA: BOTANY, VOLUME 29

own: T. meiandra, with six varieties, a (earlier named by Wawra
op. cit. 159 with the untenable Heptapleurum waimeae), /3, 7, 8, e,
and f, 1 T. Lydgatei, and T 1 . kaalae. These three additional species of
Hillebrand's were transferred likewise by Harms (loc. cit.) to Tetra-
plasandra, making a total at that time of six species of Tetraplasandra
for the Hawaiian Islands.

In 1913, Rock (Indig. Trees Haw. Isls. 337-349, pis. 135-139)
presented an expanded treatment of the genus for the Hawaiian
Islands, listing a total of eight species, two of which, namely,
T. waialealae and T. lanaiensis, he had recently described elsewhere
(College Haw. Publ. Bull. 1: 10, pi. 1, and 12, pi. 2. 1911). Rock
had made numerous field observations on the various islands and
assembled a vast quantity of specimens, many of them of superior
quality. His success in discovering numerous new localities unknown
to Hillebrand for many of the varieties of T. meiandra, he explained
by saying that "the forests have merely been opened up nowadays
by ditch trails, while in Hillebrand's time the rain forests were
almost inaccessible." Rock (p. 347) voiced his intention of pro-
ducing a monograph later on, but he seems never to have done so.

Almost contemporaneous with the collections by Rock were
those by Mr. Charles Noyes Forbes, whose activities were halted
by his premature death a third of a century ago. Like many of
Rock's specimens, many by Forbes have since proved new to science
and will be found cited in considerable numbers in the present text.
Forbes lived to publish only one new species in the genus, his
T. racemosa, a species which, however, is best set apart in a separate
genus (see "Excluded Species" below).

Following Forbes, Skottsberg made various collections on the
islands and published several new species and varieties. These are
listed under T. pupukeensis Skottsb., T. meiandra (Hillebr.) Harms,
and T. kohalae Skottsb. and bring the number of Hawaiian species
to ten.

The most extensive of all collections of Tetraplasandra, however,
are those of Dr. Otto Degener and his associates. Dr. Degener has
spent some three decades in residence in the islands and collected
far more numerously than any other collector of Hawaiian plants.
He described one new species, T. waianensis (Fl. Haw. fam. 281), in

1 Hillebrand did not specify varietal status for these subordinate categories.
In the immediate context (pp. 152, 153) he used the word "forms," but on an
earlier page (151) he had already referred to a as a variety. I have followed
Drake del Castillo, Rock, Skottsberg, and others in citing Hillebrand's Greek
letters (somewhat arbitrarily to be sure) as applying to varieties.

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 51

1938, but his other new taxa have since been described as varieties
or formae with myself in joint authorship.

In 1952 appeared two papers by myself on Hawaiian Tetrapla-
sandra (Bot. Lean 1 , nos. 6: 19 and 7: 7) and in 1953 still a third
paper (ibid. no. 8:2). These contained numerous notes, descriptions
of new varieties and formae, and descriptions of three new species
so similar to orthodox Tetraplasandra as currently accepted that
they need merely be mentioned here: T. lihuensis, T. Munroi, and
T. bisattenuata. In the last-cited paper, the above fourteen species
were apportioned to three sections of the genus. To Eutetraplasandra
Harms (sensu magnopere restricto) was assigned the lone species
T. hawaiiensis. To the new section Neotetraplasandra were assigned
all of the other thirteen species except T. meiandra, which was
placed by itself in the third section, Nothotetraplasandra Harms.
Besides these three sections, as will be seen later (p. 54), the genus
contains a fourth, namely, Pterotropia. This section contains the
remaining five species known for Hawaiian Tetraplasandra at the
present time, and a brief statement of its inception and subsequent
history may be of value here.

In 1867, Horace Mann, Jr., the son of the great educator,
described two new Hawaiian species that he himself and William
T. Brigham had collected between May 4, 1864, and May 18, 1865.
One, Mann & Brigham 606, had come from Kauai and was named
Heptapleurum kavaiense (the v changed by several subsequent
authors, as also from here on in the present text, to u). The other,
Mann & Brigham 349, had come from Lanai and was named H.
dipyrenum. Mann considered the two species as constituting a new
section for Heptapleurum Gaertn. He created, therefore, the section
name Pterotropia, presenting an adequate description. The following
year, Seemann (Jour. Bot. 6: 103. 1868) renamed Mann's first species
Agalma kauaiense. He created the new genus Dipanax for H.
dipyrenum and renamed the one contained species Dipanax Mannii.
In 1888 Hillebrand (Fl. Haw. Isls. 149) went back to Mann's section
name Pterotropia but raised it to generic rank a course not per-
mitted by the rules of today, since the earlier Dipanax of Seemann
had been validly published and would take precedence over Ptero-
tropia on the generic level. Hillebrand gave valuable notes upon
Pterotropia as a genus, also upon related araliads. With him the
genus Pterotropia consisted of the two species originally described
by Mann, also a new var. (called Tetraplasandra micrantha in the
present text) of P. kauaiensis and a third species, P. gymnocarpa,
so named because of its anomalous, completely superior ovaries,

52 FIELDIANA: BOTANY, VOLUME 29

the other taxa in Pterotropia having the ovaries inferior or only
partially superior.

Shortly after Hillebrand's treatment, there appeared the Illus-
trationes Florae Insularum Maris Pacifici by Drake del Castillo.
Drake accepted Dipanax Mannii Seem, and Heptapleurum kauaiense
Mann (misspelling the trivial epithet as kauiense, however) and
renamed Hillebrand's new species Heptapleurum gymnocarpum. In
1897, Heller (Minnesota Bot. Studs. 1: 870) went back to the name
Dipanax Seem, for the three species known to him but rejected
Seemann's D. Mannii in favor of the new combination Dipanax
dipyrena. He rejected Agalma kauaiense (H. Mann) Seem, in favor
of the new combination Dipanax kauaiense. For Pterotropia gym-
nocarpa Hillebr. he made the new combination Dipanax gymnocarpa.

Rock, already mentioned above, preferred Pterotropia to Dipanax
on grounds of priority (although as a generic name Dipanax preceded
Pterotropia by twenty years).

Recently (Bot. Leafl. no. 7: 10. 1952), I have stated that a mono-
graphic study of Tetraplasandra "reveals so much interlocking and
overlapping of characters between Tetraplasandra" and the species
assigned to Pterotropia "as to remove all warrant for their segrega-
tion." Accordingly, Pterotropia was restored to its original rank of
section but transferred as a section to a place within the genus
Tetraplasandra where, as already stated, it constitutes the fourth
(and last) section. To it belong T. kauaiensis (type of the section)
with several additional varieties (six recently described, one the early
Heptapleurum dipyrenum H. Mann reduced to varietal rank); the
T. micrantha and T. gymnocarpa already mentioned, the latter with
three additional varieties; and, finally, the two recently described
species, T. turbans of northwestern Oahu and T. kahanana of
northeastern Oahu.

As in my previous revisional studies, numerous photographs of
the more important herbarium specimens have been taken. As
formerly, a complete set has been retained in my own private
albums and a duplicate set deposited in the Herbarium of Chicago
Natural History Museum. To the authorities of that institution,
(and especially to Dr. Theodor Just, Chief Curator, Department of
Botany) who have afforded opportunities for conducting the research
summarized in the following pages, and also to the authorities of all
other co-operating institutions, it is a pleasure to express here my
appreciation and thanks.

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 53

Genus TETRAPLASANDRA A. Gray: Description

A. Gray, Bot. U. S. Explor. Exped. 727, tab. 94. 1854; Dipanax
Seem. Jour. Bot. 6: 130. 1868; Triplasandra Seem. Jour. Bot. 6:
139. 1868; Pterotropia (H. Mann) Hillebr. Fl. Haw. Isls. 149. 1888.

Unarmed glutinous trees or shrubs. Leaves alternate, impari-
pinnate with 5-21 entire leaflets, petioles basally clasping and estipu-
late or with but rudimentary stipules; terminal petiolule usually
articulate. Inflorescence terminal or lateral, the florets umbellate-
racemose on the umbellate-racemose branches of a usually stalked
panicle, or in simple or compound umbels or at times subracemiform
umbels; bracts minute, caducous. Pedicels not articulate with
calyx, the latter in most species adnate in fruit to ovary halfway up
or more, its limb narrow, spreading, often crenulate, often conspi-
cuous. Florets hermaphrodite. Petals 5-9, epigynous, valvate in
the bud, thick or thickish, cohering or finally all or most of them
separating and spreading, often involute and even uncinulate at
tip. Stamens as many as and alternating with petals or 2-8 times
as many, arranged in 1-4 circles around an annular disk, filaments
short and thickish, anthers introrse and ovate or oblong to lanceolate.
Ovary 2-13 (rarely -16) -celled, with a pendulous, anatropous ovule
(rarely 2 ovules) in each cell, in fruit half to almost entirely inferior
or by exception (T. gymnocarpa and vars., T. turbans} completely
superior, its top depressed or truncate or more often convex to conical
and serving at center as an often ill-defined stylopodium, this some-
times contracted into a short, discrete style; stigmas 2-13 (rarely
-16), capitate (or if numerous then slender and radiating). Fruit a
drupe, not or but slightly succulent, globose to ovoid or cylindrical,
rounded to angular or compressed ; pyrenae chartaceous, crustaceous,
or coriaceous, ovoid to compressed, with 1 or 2 ridges on either side.
Seeds often ribbed or furrowed. A genus of about twenty- two
known species, all but three (namely, T. Koordersii Harms of Celebes,
T. paucidens Miq. of the East Indies, and T. philippinensis Merr.
of the Philippine Islands) belonging to the Hawaiian Islands.

1 Name from Ttrpaw\a.<7ioff, fourfold, and avdpa, stamens, four times as many
stamens as petals having been found in the flowers or florets of the type species.
The International Code of Botanical Nomenclature adopted at Stockholm in
1950 recommends (Rec. 83A) but does not require the masculine gender for Tetra-
plasandra, the andra of which comes from a masculine Greek word. In the present
work, however, the feminine gender has been maintained throughout in accordance
with general practice for generic names so derived when they end in a, as shown,
for example, in the binomials Calliandra amazonica Benth., Comandra Richard-
soniana Fern., Cyrtandra platyphylla A. Gray, Peltandra virginica (L.) Schott &
Endl., Tetraplasandra racemosa Forbes, etc.

54 FIELDIANA: BOTANY, VOLUME 29

Type species: Tetraplasandra hawaiiensis A. Gray, Bot. U.S.
Explor. Exped. 728, tab. 94. 1854.

EXCLUDED SPECIES

T. racemosa Forbes, Occ. Paps. Bishop Mus. 6: 4: 51, figs. 9 and
10. 1917= Munroidendron racemosum (Forbes) Sherff, Bot. Leafl.
no. 7: 22. 1952.

SECTIONS OF TETRAPLASANDRA

Four sections may be recognized. These are:

Sect. I. Eutetraplasandra Harms in Engler & Prantl, Nat.
Pflanzenfam. 3:8: 29. 1898 (as to T. hawaiiensis only). Stamens
four times as many as petals or fewer; inflorescence paniculate;
leaves tomentose beneath. No. 1.

Sect. II. Neotetraplasandra Sherff, Bot. Leafl. no. 8: 12. 1953
(sect. Eutetraplasandra Harms, loc. cit., exclud. T. hawaiiensis}.
Stamens two to eight times as many as petals; inflorescence umbel-
late or compound-umbellate or sometimes varying somewhat to race-
mose-paniculate; leaves commonly glabrous beneath. Nos. 2-13.

Sect. III. Nothotetraplasandra Harms, op. cit. 30. 1898.
Stamens about as many as petals; inflorescence etc. as in sect. II.
No. 14.

Sect. IV. Pterotropia (H. Mann) Sherff, Bot. Leafl. no. 7: 10.
1952; Heptapleurum Gaertn. sect. Pterotropia H. Mann, Proc. Amer.
Acad. 7: 168. 1867; Pterotropia (H. Mann) Hillebr. Fl. Haw. Isls.
149. 1888 (for a genus). Stamens as many as petals; inflorescence
coarsely paniculate, the florets umbellate-racemose on the umbellate-
racemosely disposed branches. Nos. 15-19.

CONSPECTUS OF SPECIES, VARIETIES, AND FORMS

1. Tetraplasandra hawaiiensis 65

var. j8. awiniensis 66

var. 7. gracilis 67

var. 8. microcarpa 68

2. Tetraplasandra waimeae 69

var. )8. angustior 71

3. Tetraplasandra waialealae 72

var. 0. urceolata 74

var. 7. acrior 74

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 55

var. 8. wahiawensis 75

var. e. subsessilis 76

4. Tetraplasandra pupukeensis 77

var. /3. venosa 78

var. 7. megalopoda 79

f. trigona 79

var. <5. nitida 80

var. e. decipiens 81

5. Tetraplasandra lihuensis 81

var. j8. gracilipes 82

6. Tetraplasandra kohalae 83

7. Tetraplasandra lanaiensis 84

8. Tetraplasandra Munroi 85

9. Tetraplasandra Lydgatei 86

var. /3. brachypoda 87

var. 7. Forbesii 88

var. 5. leptorhachis 89

var. e. coriacea 89

10. Tetraplasandra oahuensis 90

var. j8. longipes 93

var. 7. eradiata 93

var. 8. pseudolongipes 94

var. e. Fauriei 95

var. f. pseudorhachis 96

var. 77. venulosior 96

var. 6. hailiensis 97

11. Tetraplasandra waianensis 97

var. /3. palehuana 99

12. Tetraplasandra kaalae 99

var. j8. multiplex 101

13. Tetraplasandra bisattenuata 102

14. Tetraplasandra meiandra 103

var. j8. bisobtusa 105

var. 7. Bryanii 105

var. d. Degeneri 106

var. e. Hillebrandii 106

var. f . hiloensis 107

var. 77. leptomera . 108

var. 6. longipedunculata 108

var. i. makalehana 111

56 FIELDIANA: BOTANY, VOLUME 29

var. K. mauiensis Ill

var. X. molokaiensis 112

var. p.. occidentalis 113

var. v. olowaluana 113

var. . ovalis 114

var. o. polyantha 115

var. TT. polystigmata 116

var. p. prolifica 116

var. s. prolificoides 117

var. T. ramosior 117

var. v. rhynchocarpa 118

var. v'. rhynchocarpoides 119

var. 4>. Rockii 119

var. x- simulans 120

var. \f/. Skottsbergii 121

var. co. tenuistylis 122

var. a/, tenuistyloides 123

15. Tetraplasandra kauaiensis 123

var. |8. dipyrena 125

var. 7. grandis 126

var. d. intercedens 126

var. e. nahikuensis 127

var. f . robustior 128

var. 77. koloana 129

var. d. occidua 129

16. Tetraplasandra turbans 130

17. Tetraplasandra micrantha 131

18. Tetraplasandra gymnocarpa 133

var. /3. leptocarpa 134

var. 7. megalocarpa 135

var. d. pupukeensis 136

19. Tetraplasandra kahanana 137

DISTRIBUTION BY ISLANDS

Island of Kauai

1. T. waimeae 2. T. waialealae

var. a. waimeae var. a. waialealae

var. jS. angustior var. /3. urceolata

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 57

var. 7. acnor
var. 5. wahiawensis
var. e. subsessilis

3. T. lihuensis

var. a. lihuensis
var. j8. gracilipes

4. T. bisattenuata

5. T. meiandra

var. 6. Degeneri

6. T. kauaiensis

var. a. kauaiensis
var. f . robustior
var. 77. koloana

Island of Oahu

1. T. pupukeensis

var. a. pupukeensis
var. /3. venosa
var. 7. megalopoda

f . trigona
var. 5. nitida
var. e. decipiens

2. T. Lydgatei

var. a. Lydgatei
var. /3. brachypoda
var. 7. Forbesii
var. 6. leptorhachis
var. e. coriacea

3. T. oahuensis

var. a. oahuensis
var. j8. longipes
var. 7. eradiata
var. 5. pseudolongipes
var. e. Fauriei
var. f. pseudorhachis
var. 77. venulosior
var. 0. hailiensis

4. T. waianensis

var. a. waianensis
var. jS. palehuana

5. T. kaalae

var. a. kaalae
var. /3. multiplex

6. T 1 . meiandra

var. a. meiandra
var. 7. Bryanii
var. 0. longipedunculata
var. i. makalehana
var. X. molokaiensis

1. T. turbans

8. T. micrantha

9. T. gymnocarpa

var. a. gymnocarpa
var. /3. leptocarpa
var. 7. megalocarpa
var. 5. pupukeensis

10. T. kahanana

Island of Molokai

1. T. hawaiiensis

var. i5. microcarpa

2. T. meiandra

var. o. polyantha
var. ^. Skottsbergii

58 FIELDIANA: BOTANY, VOLUME 29

Island of Lanai

1. T. hawaiiensis 4. T. meiandra

var. a. hawaiiensis var. e. Hillebrandii

var. 6. microcarpa - m 1 . .

5. T. kauaiensis

2. T. lanaiensis var. 0. dipyrena

3. T. Munroi

Island of Maui

1. T. hawaiiensis 3. T. kauaiensis

var. 7. gracilis var. /3. dipyrena

var. 5. microcarpa (fide Hillebr.)

~ , . , var. 6. inter cedens

2. T. meiandra ,.,

, var. e. nahikuensis
var. TJ. leptomera .,

. . var. 0. occidua
var. K. mauiensis

var. /i. occidentalis
var. v. olowaluana
var. . ovaKs
var. TT. polystigmata
var. T. ramosior

Island of Hawaii

1. T. hawaiiensis 4. T. kauaiensis

var. a. hawaiiensis var. /3. dipyrena

var. )8. awiniensis (fide Hillebr.)

var. 5. microcarpa var. 7. grandis

2. T. kohalae

3. T. meiandra

var. /3. bisobtusa
var. f . hiloensis
var. p. prolifica
var. $. prolificoides
var. i>. rhynchocarpa
var. i/. rhynchocarpoides
var. 0. Rockii
var. x- simulans
var. w. tenuistylis
var. a/, tenuistyloides

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 59

KEY

a. Stamens 2-8 times as many as petals.

6. Leaves (except for seedlings and watersprouts) normally tomen-
tose underneath; inflorescence paniculate; stamens 3 or 4
times as many as petals 1. T. hawaiiensis and vars.

6. Leaves glabrous underneath; inflorescence 1-4-umbellate.
c. Stamens 6-8 times as many as petals; drupes urceolate-
globose, 1.5-3 cm. thick, strongly ribbed when dry; natives
of Isl. Kauai 2. T. waimeae and var. /8. angustior.

c. Stamens mostly fewer; drupes smaller or at least more slender.

d. Petals 8-10 mm. long or, if shorter, then drupes ovoid-

cylindric, 2.2 cm. long and up to 1 cm. thick.
e. Inflorescence once- or imperfectly twice-umbellate, um-
bels 23-28-flowered; native of northwestern Hawaii.

6. T. kohalae.

e. Inflorescence twice-umbellate or, if with solitary pe-
duncle, not from Isl. Hawaii; natives of Isl. Oahu.

4. T. pupukeensis and vars.

e. Inflorescence thrice-umbellate; natives of Isl. Kauai.

/. Flowers large, their calyx 8-15 mm. tall; peduncles
and rays without nodes. . .3. T. waialealae and vars.

/. Flowers smaller, their calyx about 5-6 mm. tall.

gr. Leaflets mostly truncate at base, coriaceous; pe-
duncles without nodes 5. T. lihuensis.

g. Leaflets contracted to rounded at base, submem-
branaceous; peduncles with one node.

5. T. lihuensis var. /3. gracilipes.

d. Petals mostly 7 mm. or less in length.

e. Drupes 5- or 6- (rarely 4-) angled; natives of southeastern

Oahu 10. T. oahuensis and vars.

e. Drupes 3-4-angled.

/. Inflorescence drooping, thrice-umbellate; secondary
umbels of 17-21 drooping, several-nodate rays 8-10
cm. long; native of Isl. Lanai 7. T. lanaiensis.

f. Inflorescence erect, or, if drooping, then with no nodes

on rays.
g. Peduncles bearing rays that bear pedicels.

60 FIELDIANA: BOTANY, VOLUME 29

h. Leaflets fusiform in outline; native of south-
eastern Kauai 13. T. bisattenuata.

h. Leaflets broader at one or both ends.

i. Rays of inflorescence 20, about 5-6.5 cm.
long; umbellets about 6-8-flowered ; native of

Isl. Lanai 8. T. Munroi.

i. Rays of inflorescence mostly fewer and shorter;
umbellets directly at tips of rays and about
10-12-flowered or at tips of subrays and 3-5-
flowered; natives of northwestern Oahu.

12. T. kaalae and var. /3. multiplex,
g. Peduncles bearing pedicels directly, rays absent.
h. Petals 5, stamens 12; natives of Koolau Range
from southeastern Oahu northwestward as far

as Wahiawa 9. T. Lydgatei and vars.

h. Petals about 8, stamens about 28; known only
from Middle Palawai Ridge and vicinity,
Waianae Range, Isl. Oahu.

11. T. waianensis and var. /3. palehuana.

a. Stamens as many as petals.
6. Ovary superior, even in fruit.

c. Umbellets dense, 10-16-flowered, at anthesis only 7-9 mm. in
diameter with pedicels under 1 mm. long. . . 16. T. turbans,
c. Umbellets and pedicels larger ... 18. T. gymnocarpa and vars.
6. Ovary more or less inferior.

c. Inflorescences mostly either simple or compound umbels. 1
d. Stigmas 2 or at times 3.

e. Drupes ovoid to urceolate-ovoid ; natives of Kilauea-

Kulani region, southeastern Kauai.
/. Drupes more or less truncate at top, stylopodium
usually contracted below stigmas into an evident

style 14. T. meiandra var. w. tenuistylis.

f. Drupes conical above calyx-limb as a stylopodium
and without an evident style.

14. T. meiandra var. a/, tenuistyloides.

1 In addition to the varieties of T. meiandra listed under this c are the ill-
named and but scantily known var. X. molokaiensis of Wailupe (on Oahu!) and the
var. 8. Degeneri of northwestern Kauai, a variety unknown as to drupes but
with thrice-compound inflorescences, numerous rays (even 18 or 21), and small
pentamerous corollas.

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 61

e. Drupes mostly more elongate, style seldom evident.
/. Inflorescence thrice-umbellate (rays present).

g. Umbellets compactly flowered, pedicels 4-6 mm.
long; native of northwestern Hawaii.

14. T. meiandra var. $. prolificoides.
g. Umbellets more open, pedicels longer.

h. Peduncles under 7 cm. long; native of north-
western Hawaii . 14. T. meiandra var. p. prolifica.
h. Peduncles longer.

i. Leaflets large, blade 12-17 cm. long and 5-7.5
cm. wide; drupe more or less ovoid, its vertex
from slightly raised to moderately conical;
local in northwestern Oahu.

14. T. meiandra var. L. makalehana.
i. Leaflets smaller.

j. Leaflets ovate to oblong, at apex somewhat
acute, at base more or less contracted;
rays mostly under 5 cm. long, umbellets
8-12-flowered; drupe slender-elongate, its
vertex manifestly conical and exserted; of
wide distribution on Oahu.

14. T. meiandra sensu stricto.
j. Leaflets oblong, obtuse at both ends; rays
5-7.5 cm. long, umbellets 15-25-flowered;
apparently confined to northwestern Ha-
waii 14. T. meiandra var. /3. bisobtusa.

/. Inflorescence twice-umbellate (rays absent).

g. Pedicels very numerous (up to 30), slender, in
fruit 0.5-1 mm. thick; leaflets not or but weakly
reticulate- venulose above; lateral petiolules 6-16
mm. long; stigmas 2; native of north westernmost
Hawaii (rarer 2-stigmatate forms of the stouter-
pedicelled var. longipedunculata of Oahu may
likewise be looked for here).

14. T. meiandra var. x- simulans.
g. Pedicels less numerous (about 10-15 to an umbellet),
in fruit 1-1.5 mm. thick; leaflets conspicuously
reticulate-venulose above; lateral petiolules 11-27
mm. long; stigmas more often 3; native of northern
Oahu 14. T. meiandra var. 7. Bryanii.

62 FIELDIANA: BOTANY, VOLUME 29

d. Stigmas mostly 5 or 6.

e. Pedicels 6-9 mm. long; peduncles on a rhachis 1 dm.
long; native of West Maui.

14. T. meiandra var. T. ramosior.

e. Pedicels 20-24 mm. long; peduncles in a sessile umbel;

not found on West Maui.

/. Drupe ovoid to cylindric-ovoid, apically truncate, its
calyx portion about 1 cm. tall; native of north-
eastern Hawaii .... 14. T. meiandra var. f . hiloensis.

f. Drupe globose- or urceolate-ovoid, at apex protruding-
conical (this portion 2-3 mm. tall), its calyx portion
about 7-8 mm. tall; native of northern East Maui.
14. T. meiandra var. IT. polystigmata.

d. Stigmas mostly 3 or 4.

e. Pedicels mostly 3-5 mm. long at anthesis; blades of
leaflets narrow-oblong to widely oblanceolate or nar-
rowly obovate; native of East Maui.

14. T. meiandra var. 77. leptomera.

e. Pedicels mostly longer.

/. Inflorescence at most twice-umbellate, peduncles 1-3.5
dm. long, up to 30-flowered; native of Isl. Oahu.

14. T. meiandra var. 6. longipedunculata.

f. Inflorescence normally thrice-umbellate.

fir. Umbellets numerously flowered, the fuller ones with
12-20 florets.

h. Trees of East Maui; pedicels subrobust at an-
thesis.

i. Drupes cylindric, about 11 X 5 mm., stylo-
podium hemispherical; native of southern
East Maui . . 14. T. meiandra var. K. mauiensis.

i. Drupes urceolate, 7x6 mm. or larger, stylo-
podium narrowly conic; native of northern
East Maui .... 14. T. meiandra var. . ovalis.

h. Trees of West Maui ; pedicels delicate at anthesis,
drupes globose-cubical or shortly trigonous-
prismatic. . 14. T. meiandra var. v. olowaluana.

h. Trees of Molokai; pedicels delicate at anthesis.

14. T. meiandra var. o. polyantha.

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 63

In. Trees of southeastern Hawaii; peduncles 6-12 cm.
long, rays 22 (1 or 2 at times attached 1-3
cm. below tip of peduncle).

14. T. meiandra var. /. rhynchocarpoides.

g. Umbellets fewer-flowered, the fuller ones with 5-12
florets.

h. Top of drupe hemispherical; rays delicate, 1
mm. thick; native of eastern Molokai.

14. T. meiandra var. x- Skottsbergii.

h. Top of drupe at most depressed-hemispherical.
i. Stylopodium elongate-conic, 3-4 mm. long
(stigmas included); native of southeastern
Hawaii . . 14. T. meiandra var. v. rhynchocarpa.

i. Stylopodium smaller.

j. Leaf -petioles stoutish, usually 3.5-4.5 rarely
to 7.5 cm. long; leaflets 5-9, broadly
oblong-oval to obovate or subrotund; na-
tive of Isl. Lanai.

14. T. meiandra var. e. Hillebrandii.

j. Leaf-petioles longer (normally 5.5-10 cm.
long).

k. Blades of 7-13 oblong to subovate, ovate,
or subobovate leaflets obtuse or rounded
at each end or even truncate at base,
moderately to conspicuously depressed-
and reticulate-venulose above, up to 11
cm. long and 7.5 cm. wide; rays 5-16;
native of southernmost Hawaii.

14. T. meiandra var. </>. Rockii.

k. Blades of 5-7 oval-elliptic leaflets gradually
narrowed at each end, at most but
weakly depressed- or reticulate-venulose,
4-8 cm. long and 2-4 cm. wide; rays
5-8; native of West Maui.

14. T. meiandra var. ju. occidentalis.

c. Inflorescences mostly paniculate, the divisions more or less
racemose-umbellate .

d. Leaflets thin, larger lateral ones more or less parallel-sided
and 15-20 cm. long, in age sparsely whitish-farinose

64 FIELDIANA: BOTANY, VOLUME 29

beneath, petiolules 4-5 mm. long; native of northwestern

Oahu 17. T. micrantha.

d. Leaflets thick or coriaceous, mostly shorter (or if as long
then basally truncate to cordate) and with curved edges,
petiolules mostly longer.

e. Entirely glabrous; rays suberect, arcuate; pedicels
slender, elongate (12-17 mm. long); native of north-
eastern Oahu 19. T. kahanana.

e. Inflorescence and usually lower leaf-surfaces (except in
var. grandis) pubescent; rays straightish and mostly
spreading or reflexed, pedicels mostly thickish and
short. . . .15. T. kauaiensis and vars.

ABBREVIATIONS USED FOR NAMES
OF HERBARIA CITED

Arn. Herb. Arnold Arboretum, Jamaica Plain, Massa-
chusetts.

Berl. Herb. Berlin Botanical Garden, Berlin-Dahlem.

Bish. Herb. Bernice P. Bishop Museum, Honolulu.

Brit. Herb. British Museum (Natural History), London.

Calif. Acad. Herb. California Academy of Sciences, San Francisco.

Carn. Herb. Carnegie Museum, Pittsburgh.

Chi. Herb. Chicago Natural History Museum, Chicago.

Corn. Herb. Cornell University, Ithaca, New York.

Del. Herb. Delessert, Geneva.

Goth. Herb. Botanical Garden, Gothenberg, Sweden.

Gray Herb. Gray, Harvard University, Cambridge, Massa-
chusetts.

Kew Herb. Royal Botanic Gardens, Kew.

Mo. Herb. Missouri Botanical Garden, St. Louis.

N.Y. Herb. New York Botanical Garden, New York City.

Par. Herb. Museum of Natural History, Paris.

Phila. Herb. Academy of Natural Sciences, Philadelphia.

Stockh. Herb. Natural History Royal Museum, Stockholm.

Univ. Calif. Herb. University of California, Berkeley.

U.S. Herb. United States National Museum, Washington,
B.C.

Yunck. Herb. Dr. T. George Yuncker, Greencastle, Indiana.

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 65

1. Tetraplasandra hawaiiensis A. Gray, Bot. U. S. Explor.
Exped. 728. 1854; ibid. pi. 94. 1857.

a. Blade of lateral leaflets subcuneately attenuate at base.

6. Petiolules of lateral leaflets usually 0.8-2 cm. long; umbellets
7-11-flowered; pedicels 5-7 mm. long at anthesis, thickish,
very densely and conspicuously tomentose; corolla depressed-
conical to subhemispherical just before opening, about 3-4
mm. tall, not strongly angulate when dried; native of north-
westernmost Hawaii var. /3. awiniensis.

6. Petiolules of lateral leaflets 2-10 mm. long; umbellets commonly
3-6-flowered; pedicels 8-14 mm. long at anthesis, becoming
more slender and less densely and less conspicuously tomen-
tose; corolla conical just before opening, about 5 mm. tall,
becoming strongly angulate on drying; native of West Maui.

var. 7. gracilis.

a. Blade of leaflets broader and usually rounded to truncate at base.
6. Petals usually 7 or 8, stamens usually 28-32, drupe 7-10-celled
and usually 8-10 mm. thick; native of southern Hawaii.

var. a. hawaiiensis.

b. Petals 5-7, stamens commonly only 17-20, drupe 9-13-celled
and usually 5-7 mm. thick; native of Molokai, Maui, Lanai,
and northwesternmost Hawaii var. d. microcarpa.

A tall, stout, much-branching tree 12-18 m. or more tall, bark
smooth and pale, branchlets invested with tawny, coarse, stellate
tomentum. Leaves 5-9-foliolate; petiole 1.5-2 dm. long (this with
the rhachis 3-6 dm. long), stellate-tomentulose; leaflets on petiolules
0.2-2.2 cm. long (or terminal one longer), blade oblong or varying
to ovate, ovate-oblong, or broadly lanceolate-oblong, at apex obtuse,
at base (for lateral leaflets oblique) truncate to cuneately contracted,
1-1.7 dm. long and about 2.5-6 cm. broad, pale, coriaceous, glabrous
above, densely tawny-tomentose beneath except hirsute with fuzzy
hairs at and near the minutely tomentulose or partly subglabrous
midrib (for leaflets of "watersprouts" 2 dm. long and 12 cm.
wide, glabrous on both surfaces). Inflorescence of 1-3 terminal
panicles, densely stellate-tomentose, up to 3.5 dm. long including
the peduncle (this nude or subnude, often 1.5-2 dm. long); pedun-
cular or secondary branches few, umbellately arranged at tip of
peduncle (or 1-5 placed racemosely lower down), umbellately
branched directly into usually 4 or 5 pedicels (these up to 2-2.5 cm.
long in type material) or into a few tertiary branches which in turn
support the pedicels. Calyx obconic-urceolate, about 4-6 mm. tall

66 FIELDIANA: BOTANY, VOLUME 29

and broad, tomentulose. Petals usually 7 or 8, tomentulose, co-
hering at apex, 6-8 mm. long. Stamens 4 times as many as petals
or fewer, in a single circle, recurved. Ovary 7-10-celled, its apex
crowned with a short stylopodium, this with 7-10 at times indistinct
stigmas; drupe globose or slightly depressed-globose, usually 10
mm. in diameter, many- (7-10-) ribbed when dry and truncate or
slightly depressed-truncate at top, the abruptly rising stylopodium
1-2 mm. tall; pyrenae 7-10, compressed, chartaceous, closely
contiguous.

Type: United States South Pacific Exploring Expedition under
Captain Charles Wilkes, in the District of Puna, Isl. Hawaii, 1838-
1842 (U.S.).

Distribution: Southern and eastern Hawaii.

Specimens examined (all from Isl. Hawaii) : Otto Degener 2,181,
along roadside in woods, South Kona, Aug. 24, 1926 (Chi.); Degener
21,759, tree 30 ft. tall, in woods among aa lava, Panaewa Woods mauka
of Hilo, Feb. 1, 1952 (Chi.) ; Degener 21,788, tree 35 ft. tall, "water
sprouts and seedlings glabrous," alt. 2,000 ft., where overtopping
forest, Kulani Prison Road, Feb. 5, 1952 (Berl.; Bish.; Carn.; Chi.,
2 sheets; Corn.; Del.; Gray, a large glabrous "watersprout" leaf
only; Kew; N.Y.; Par.); Degener 21,834, beautiful tree 40 ft. tall,
in forest on aa lava, along main road near Hoopuloa Lava Flow,
Feb. 9, 1952 (Am.; Chi.; Phila.; Univ. Calif.); Degener & Wiebke
2,181, along roadside in woods, South Kona, Aug. 24, 1926 (N.Y.,
4 sheets; U.S.); G. 0. Fagerlund & A. L. Mitchell 799, tree 20-25 ft.
tall, Ohia-shrub-transition forest on aa flow, alt, 1,900 ft., south of
Kalapana Trail, Aug. 17, 1943 (Bish.); Joseph F. Rock, Kapua,
South Kona, January, 1912 (Gray, 2 sheets, my photograph no.
4,260); Rock 10,026, tree 50-60 ft. tall, not uncommon in upper
forest, alt. 3,200 ft., Kapua, South Kona, Jan. 28, 1912 (Am., 2
sheets); Rock (similarly) 10,026, Kapua, South Kona, February,
1912 (Bish.; Gray; U.S., 2 sheets, my photograph no. 4,259); Carl
Skottsberg 459, lava flow west of Pahoa, Sept. 9, 1922 (Bish.; Goth.) ;
United States South Pacific Exploring Expedition under Capt. Wilkes,
District of Puna, 1838-1842 (type collection, Gray, my photograph
no. 4,175; N.Y.).

Tetraplasandra hawaiiensis var. /3. awiniensis Sherff, Bot.
Leafl. no. 6: 37. 1952.

Tree 7.5-15 m. tall. Leaflets a little narrower, usually all (not
only the terminal one) gradually or subgradually narrowed to the

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 67

petiolule. Panicles of the inflorescence solitary or 2-5 congregrated
in a sessile umbel. Umbellets numerously (more often 7-1 1-)
flowered, the thickish pedicels only 5-7 mm. long and very densely
and very conspicuously yellowish-brown-tomentose. Flowers rather
small, the petals only about 4 mm. long. Drupes a little smaller,
about 6-8 mm. thick; pedicels finally elongate and more slender,
often 1.5-2.1 cm. long.

Type: Joseph F. Rock 8,386, tree 25 ft. tall, woods of Awini,
North Kohala, Isl. Hawaii, June, 1910 (Arn.).

Distribution: Northwesternmost Hawaii.

Specimens examined (Isl. Hawaii) : Otto Degener & Amy Greenwell
21,898, beautiful tree of forested slopes, Pololu-Honokanenui region,
North Kohala, Feb. 18, 1952 (topotypes, Bish.; Calif. Acad.; Chi.,
3 sheets, my photograph no. 4,264; Del.; Gray; Kew; Mo.; N.Y.;
Phila.; Univ. Calif.; U.S.); Rock 8,386 (type, Arn., my photograph
no. 4,261: isotypes, Bish., my photograph no. 4,263; Gray, 3 sheets,
my photograph no. 4,262).

Tetraplasandra hawaiiensis var. 7. gracilis Sherff, Bot.
Leafl. no. 6: 38. 1952.

Leaves 4.5 dm. long; petiole 13 cm. long; leaflets 5-9, petiolule
for the lateral ones 2-10 mm. long, blade smaller and narrow-oblong,
at apex subacute or obtuse, at base (for the lateral ones) oblique
and subcuneately attenuate, 6-12 cm. long and 3-4 cm. wide.
Branches and branchlets of inflorescence more slender, the umbellets
few- (commonly 3-6-) flowered, the pedicels slender and 8-14 mm.
long at an thesis. Calyx small toward an thesis, about 3 mm. tall
and 3-5 mm. thick; petals 5, near anthesis scarcely 6 mm. long;
stamens 15; stigmas about 10; drupes urceolate-cylindric, about
10-costate, their calyx about 5 mm. tall and thick, the truncate top
abruptly narrowed into a slender stylopodium about 1-1.3 mm. tall.

Type: Joseph F. Rock & S. T. Hashimoto 16,033, Waihee Valley,
West Maui, September, 1918 (Bish.).

Distribution: Known only from general vicinity of type locality
in West Maui.

Specimens examined (West Maui) : Degener, Greenwell, & Hathe-
way 21,142, single tree on shrubby, forested slope, 35 ft. tall with
trunk 1 ft. thick, in narrowest, rainiest part of Makamakaole Gulch,
Nov. 28, 1950 (Bish.; Chi.; Gray; N.Y.); Rock & Hashimoto 16,033
(type, Bish., my photograph no. 4,268 : isotype, Bish., my photograph
no. 4,269).

68 FIELDIANA: BOTANY, VOLUME 29

The leaflets are similar to those of var. awiniensis but apically
somewhat more acute.

Tetraplasandra hawaiiensis var. 5. microcarpa Sherff, Bot.
Leafl. no. 6: 38. 1952.

Leaflets narrowly to widely oblong, up to 2 dm. long and to
8 cm. wide, obtuse to acute at tip, tomentose beneath or sometimes
glabrous on both surfaces. Calyx smaller, at anthesis only 3-4
mm. tall and subequally thick; petals 5-7, only about 5-6 mm.
long; stamens commonly only 17-20. Drupes smaller, the depressed-
globose body only 5-7 mm. thick, truncate to convex at top, 9-13-
celled; ribs sharper on drying.

Type: Joseph F. Rock 7,006, above Kaluaaha, Isl. Molokai,
April, 1910 (Bish.).

Distribution: Molokai, Maui (where the leaflets often reach the
maximum size), Lanai, and northwesternmost Hawaii.

Specimens examined (Isl. Molokai) : Edwin H. Bryan, Jr. 618,
tree 10 m. tall, trunk diam. 2.5 dm., heartwood pithy and cream-
colored, sapwood hard and of light color, on slopes of gulches and
flat, lower edge of rain-forest, alt. 300 m., Kainalu, July 24, 1927
(Bish., 2 sheets) ; Otto Degener, east part of Kaluaaha plateau, July
3, 1928 (topotypes, N.Y.; U.S.); Degener 18,017, forest, same place
and date (topotypes, Mo.; N.Y., 3 sheets; U.S.); Abbe Urbain Faurie
269, alt. 600 m., Pukoo, May, 1910 (Am.; Bish.; Par.); Charles N.
Forbes 508-Mo, tree 25-30 ft. tall, ridges south of valley, Halawa,
September, 1912 (Bish., 2 sheets); William Hillebrand, Molokai
(Bish.; Gray; Kew; U.S., where locality is given as Leper Settlement
[cf. "Kalauwau" of Hillebrand's text, p. 155, a locality unknown
to me]); Joseph F. Rock, Kaluaha (Kaluaaha), March, 1910 (topo-
type, N.Y.); Rock 7,006 (type, Bish., my photograph no. 4,266:
isotypes, Arn., 2 sheets; Gray, 2 sheets, my photograph no. 4,265);
G. W. Russ, Ahaino, above Forest Reserve Line, December, 1932
(Bish., 2 sheets) ; H. St. John et alii 12,846, tree 25 ft. tall, rain-forest,
alt. 2,600 ft., ridge east of Mapulehu Valley, Dec. 29, 1932 (Yunck.).

East Maui: Otto Degener, along ditch trail between Honomanu
Valley and Keanae, July 14, 1927 (Mo.); Degener, near ditch trail,
Keanae Valley, July 19, 1927 (Mo.; N.Y.); Degener 17,787, near
Keanae Valley, July 19, 1927 (Mo.; N.Y.); Degener 17,788 pro parte
(this number used by Degener also for another East Maui plant,
referable to T. meiandra var. ovalis, qu. vide}, precipitous canyon

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 69

slope, along ditch trail at Keanae Valley on Honomanu Valley side,
July 14, 1927 (Mo.; N.Y., 2 sheets; U.S.); Degener & Wiebke 2,316,
only one tall tree on exposed cliff along ditch trail, Honomanu
Valley, July 14, 1927 (N.Y., 3 sheets; Univ. Calif.; U.S.).

West Maui: Degener, "with both glabrous and tomentose leaves
on same tree," half-mile north of Keahikauo, July 21, 1927 (N.Y.,
2 sheets); Degener 2,319, tall trees in dicranopteris-tangled forest,
same place and date (Mo.; N.Y., 3 sheets); Charles N. Forbes
108-M, ridge west of lao Valley, June, 1910 (Bish.); William Hille-
brand, "Kanapali" (Kaanapali; U.S., where mounted on same sheet
with Hillebrand's Leper Settlement material from Molokai) ; Horace
Mann, Jr., & William T. Brigham 378, West Maui (Bish.; Gray;
Mo.).

Isl. Lanai: Forbes 141-L, mountains near Koele, June, 1913
(Bish.); Forbes 377-L, locality not stated, September, 1917 (Bish.);
George C. Munro 39, edge of Huola, east side of mountain, August,
1913 (Bish., 3 sheets; U.S.); Munro 140, Lanaihale, west face, Nov.
17, 1913 (fruiting; Bish.); Munro 140, Maunalei mauka, July 19,
1915 (flowering; Bish.); Rock 8,016, Haalelepakai, main ridge, July
21, 1910 (Gray; U.S.); Rock 8,016a, Mahana ridge, July, 1910 (Bish.);
Rock (similarly) 8,016a, Haalelepakai, south slopes of main ridge,
August, 1910 (Gray, 2 sheets) ; Rock & Hammond 8,050, main ridge,
July 22, 1910 (Arn., 2 sheets; Bish.).

Isl. Hawaii: Degener & Wiebke 2,185, sunny, mesophytic forest,
east of Pololu Valley, Aug. 11, 1926 (Mo.; N.Y., 6 sheets; Univ.
Calif.; U.S.); L. H. MacDaniels 327, spreading tree, 15 m. tall,
Kohala ditch trail, sides of gulch in rain-forest, alt. 1,000 m., Dec.
6, 1926 (Bish.).

2. Tetraplasandra waimeae Wawra, Flora 56: 158. 1873;
cf. Sherff, Bot. Leafl. no. 6: 25. 1952; Tetraplasandra Elstonii Hochr.
Candollea2:481. 1925.

Leaflets mostly narrow-oblong var. /3. angustior.

Leaflets broader var. a. waimeae.

Trunk erect, 9-12 m. tall and 3 dm. in diameter, sparingly
divided near top into short, ascending branches; bark smooth.
Leaves clustered at ends of branches, 3-4.5 dm. long; leaflets 5-13,
petiolules variously up to 3.5 cm. long; blades oblong to ovate-
oblong or even obovate, 6-15 cm. long and 3.5-6.5 cm. wide, at
apex obtuse to truncate or even emarginate, at base rounded to

70 FIELDIANA: BOTANY, VOLUME 29

broadly cuneate (and for lateral leaflets oblique), chartaceous to
coriaceous, glabrous. Inflorescence a terminal umbel of several
rays (with or without a short common rhachis), these 5-15 cm. long
and with a terminal umbellet of 10-20 flowers on thick, compressed
or angulate pedicels 2-5 cm. long. Calyx 8-12 mm. tall, thick-
tubular to slender-urceolate, somewhat constricted below the wavy,
denticulate border. Petals 7 or 8, triangular-lanceolate, 10-12
mm. long, pink or reddish, coriaceous, glabrous, finally expanded,
the wide-open corolla up to 3.5 cm. across (this unique in the
genus for size). Stamens normally 6, 7, or 8 times as many as
petals, 8-10 mm. long, in two rows. Ovary variously 6-10-celled
(cells sometimes 2-ovulate), the stigmas on a short, compressed
stylopod of 1 mm. Drupe urceolate-globose, 1.3-3.2 cm. in diameter,
somewhat fleshy, strongly ribbed when dry, truncate and 6-10-
radiate at top, stylopodium cylindric-compressed, the 6-10 stigmas
scarcely distinct at the oval-oblong apex; pyrenae compressed,
separated by pulp, thick-coriaceous, deeply notched at upper inner
angle, sharp at back and with 2 prominent ridges on each side.

Type: Heinrich Wawra 2,114, Halemanu, Isl. Kauai (presumably
in Mus. Vienna).

Distribution : Kaholuamanu to Kumuweia Ridge and Halemanu,
northwestern Kauai.

Specimens examined (all from Isl. Kauai) : Degener & Greenwell
21,577, a 20 foot tree, in forest, Kumuweia Ridge east of Waineke
Swamp, Jan. 1, 1952 (Bish.; Carn.; Chi., 2 sheets; N.Y.) ; F. Fagerlind
& C. Skottsberg 6,608, single trees, forest near Kalalau Lookout, Mar.
14, 1948 (Stockh.) ; Charles N. Forbes 432-K, mountains in vicinity of
Kaholuamanu, September, 1909 (Bish.); Forbes 1,064-X, Waimea
Drainage Basin, west side, July 3-Aug. 18, 1917 (Bish.); William
Hillebrand, Waimea, 1871 (Gray, my photograph no. 4,211); Hille-
brand, Halemanu (fragment, Bish.); Albert S. Hitchcock 15,550,
vicinity of Kaholuamanu, Oct. 26, 1916 (U.S.); Joseph F. Rock,
Kauai, October, 1916 (Bish., my photograph no. 4,290); Rock, 1,720,
Halemanu, Feb. 14-26, 1909 (topotype, Bish., foliage only); Rock
1,744, Kaholuamanu, Mar. 3-10, 1909 (Bish.; U.S.); Rock 5,162,
same place, September, 1909 (Gray); Rock 5,165, same place, Sept.
3, 1909 (Bish., my photograph no. 4,213); Rock 5,167, same place,
September, 1909 (Gray, my photograph no. 4,214); Rock 5,168, in
the mountains above Waimea, in forest of Kaholuamanu near
Opaiwela, alt. 3,700 ft., Sept. 3, 1909 (Am., 2 sheets) ; Rock 5,913,
Opaiwela trail, alt. 3,700 ft., Sept. 3, 1909 (Bish.); Carl Skottsberg

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 71

1,009, alt. about 1,200 m., near Kokee ranger station, Waimea, Oct.
28, 1922 (Goth.).

As previously remarked in my Botanical Leaflet no. 6, page 25,
Hillebrand may have accompanied Wawra on the type-collection
trip, for his specimen at Gray Herbarium, "collected by Dr. W. Hille-
brand, 1871," has the species named in honor of Wawra. Moreover,
the fragment, a leaflet and entire flower, at Bishop Museum, "ex
coll. Hillebrand in Herb. Berlin" bears on the label, "Halemanu,"
Wawra's type locality. Had Wawra not published the binomial
Tetraplasandra waimeae in 1873, Hillebrand's binomial honoring
Wawra would doubtless have appeared in 1888, when Hillebrand's
posthumous Flora of the Hawaiian Islands was published. 1

Hochreutiner (loc. cit.) described his Tetraplasandra Elstonii
from a specimen collected by himself, his no. 3,538, alt. about
1,300-1,400 m., Kaunupalanui, near Gay cottage on Kaholuamanu,
Waimea, Isl. Kauai, Apr. 19, 1905. With him, T. Elstonii stood
intermediate between T. waimeae, described originally from Hale-
manu, and T. waialealae, a species peculiar to the high summit
swamp of Mt. Waialeale. 2 Unfortunately, T. waimeae had been
described rather narrowly, the ovary for example being called not
more than 8-celled. For T. Elstonii the ovary was described as 8-
or 9-celled. However, among the specimens examined from both
Halemanu and Kaholuamanu, and listed above for T. waimeae,
variations from 6 to 10 cells are found. Much the same is true of
Hochreutiner's other criteria listed; therefore T. Elstonii seems
entirely synonymous with T. waimeae.

Tetraplasandra waimeae var. /3. angustior Sherff, Bot. Leafl.
no. 6: 26. 1952.

Leaflets commonly narrow-oblong, the lateral margins usually
almost parallel, petiolule 4-11 mm. long, blade apically retuse to
rounded or obtuse and basally broad-cuneate to rounded (and for
the lateral leaflets very oblique), up to 11 X 3.5 cm. or 12 X 4 cm.

1 In his Flora, however, Hillebrand mentioned Wawra and Knudsen as
collectors but omitted his own name.

2 Skottsberg (Meddel. Goteb. Bot. Tradg. 15: 415. 1944), in questioning the
status of T. Elstonii as apart from T. waimeae and T. waialealae, seems to have
overlooked the ecological consideration that T. Elstonii and T. waimeae were
both from the drier forests of Waimea (on the leeward side of Kauai), while
T. waialealae was a swamp inhabitant on Mt. Waialeale (and within the wind-
ward, rainy area on Kauai).

72 FIELDI ANA: BOTANY, VOLUME 29

or 9 X 3.2 cm. or rarely 14 X 5 cm. Mature drupes up to 1.8
cm. tall and 1.5 cm. thick.

Type: Amos Arthur Heller 2,734 pro parte, in and near a bog at
the head of Wahiawa Stream, Isl. Kauai, Aug. 21, 1895 (Phila.).
Distribution: Island of Kauai; see discussion below.

Specimens examined (all from Isl. Kauai) : Heller 2,734 pro parte
(type, Phila., my photograph no. 4,209); Heller 2,734 pro parte, in
and near a bog at the head of Wahiawa Stream, Aug. 12, 1895
(Arn.); Heller 2,734 pro parte, on Kaholuamanu, Sept. 10-16, 1895
(Bish., my photograph no. 4,210; Corn., where dated Sept. 2-9,
1895; Par., 2 sheets).

Under his no. 2,734, Heller collected two kinds of material.
The first will be treated below under T. waialealae var. wahiawensis.
The second is the present var. angustior of T. waimeae. The specimen
at Philadelphia was selected as the type, since it bore a fine in-
florescence in the mature-fruit stage. The inflorescence is essentially
sessile as in T. waimeae proper (i.e., var. waimeae}, not stalked as
in T. waialealae.

Heller, it will be noted, gave two habitats, one, Kaholuamanu,
and the other, at or near what is often called Wahiawa Swamp.
Since the variety is so extremely rare as never to have been collected
apparently by anyone else, it seems incredible that he found it in
both of the widely separated habitats mentioned. In a number of
cases I have found Heller's specimens to have been distributed with
"crossed labels." Since we know that one form under his no. 2,734,
namely T. waialealae var. wahiawensis, does grow at and near Wa-
hiawa Swamp, it seems probable that his other form, the present
T. waimeae var. angustior, came from somewhere on Kaholuamanu.

3. Tetraplasandra waialealae Rock, Coll. Haw. Publica-
tions Bull. no. 1: 10, pi. 1. 1911.

a. Leaflets 11-15, sessile to shortly petiolulate (petiolule 1-10 mm.) ;
native of swampy areas in northwestern Kauai.

var. f. subsessilis.

a. Leaflets 5-11, at least the lower petiolules of moderate (9-16
mm.) to greater (16-26 mm.) length.

b. Leaflet-blades 5.5-9 cm. long and 2.5-5 cm. wide, at apex
subacuminate to obtuse; native of swampy places in southern
Kauai . . . . var. 5. wahiawensis.

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 73

6. Leaflet-blades much larger, mostly 9-15 cm. long and 4-7 cm.

wide.

c. Leaflet-blades more often broad or broadish at each end, at
apex abruptly narrowed into a shortly acuminate tip.

var. a. waialealae.

c. Leaflet-blades cuneately contracted at base, apically sharp-
attenuate and -acuminate.
d. Calyx of immature drupe cylindric-prismatic.

var. y. acrior.
d. Calyx of immature drupe thick-urceolate.

var. |8. urceolata.

Small tree 4.5-7.5 m. tall; trunk rather short, about 1.5-2 dm.
thick, its bark white; branches long and suberect, the irregularly
pinnate leaves crowded at their ends, 3-4.5 dm. long. Leaflets
7-11; blades dark-green and glossy, mostly oblong but a few oblong-
ovate, at usually broad apex sometimes gradually but more often
abruptly narrowed into a shortly acuminate tip, at base the lateral
ones oblique and broadly rounded to cuneately contracted, thick-
coriaceous, 7-14 cm. long and 3.5-6.5 cm. wide; lateral petiolules
l-2(-4) cm. long. Inflorescence a terminal, compound umbel of
usually 4 peduncles, each about 7-13 cm. long and bearing 6
rays 6 cm. long, these each bearing an umbellet of usually
5 pedicels about 2-2.2 cm. long. Calyx more or less cylindric,
purplish-black with an undulate border, 7-9 mm. tall. Petals 5-7,
triangular, thick, glabrous, with a prominent median nerve, 1 cm.
long. Stamens in 2 circles, 4 times as many as the petals. Ovary
6-celled; stigmas 6, on a conical stylopod 5 mm. tall. Mature
drupes not seen.

Type: Joseph F. Rock 8,870, abundant on summit of Waialeale,
alt. 5,000 ft., Isl. Kauai, Oct. 20, 1911 (Bish.; but see discussion
below) .

Distribution: Central to northeasternmost Kauai.

Specimens examined : Degener & Greenwell 21,730, a single fallen
tree of 35 ft. in height (inflorescence a brilliant claret), in lower
forest, ridge mauka (i.e., inland) from Papaa, Moloaa Forest
Reserve, Jan. 16, 1952 (Bish.; Chi.; N.Y.); Rock 8,870 (isotypes,
Arn., 2 sheets; Bish., my photograph no. 4,201; Gray, 3 sheets).

This species was founded upon mixed material, Rock 4,902 and
Rock 8,870. The first cited habitat, however, was "Summit of
Waialeale, elevation 5,000 ft.," and this may be taken as limiting

74 FIELDIANA: BOTANY, VOLUME 29

the species in its narrower sense to Rock 8,870, the tree with leaflets
obtuse but mostly very short- or abrupt-acuminate tips. Indeed,
Rock's full-page plate is of that form. Excellent specimens of Rock
8,870 have been available for study, as above cited.

Rock 4,902 is seen to consist of two forms, both of them different
from var. waialealae or the species proper in having apically sharp-
attenuate and -acuminate leaflets, and one of them, from Kaluiti
Stream, with urceolate (not cylindric) fruiting-calyces. These are
treated below under vars. acrior and urceolata.

Tetraplasandra waialealae var. 0. urceolata Sherff, Bot.
Leafl. no. 6: 23. 1952.

Leaves 4.5 dm. long; petiole 1.5 dm. long, toward base
ciliate, very minutely and numerously papillate, and sparsely pu-
berulous; leaflets 9, narrowly oblong or the bottom ones oblong-
ovate, at apex sharply and somewhat gradually acuminate, at base
very widely and at times obliquely cuneate, the blade 11-15 cm.
long and 3.5-5.5 cm. wide, the petiolule 4-16 mm. long. Drupes at
least when submature urceolate, about 16-17 mm. tall including
stylopodium and stigmas and 1 cm. thick below the middle.

Type: Joseph F. Rock 4,902 pro parte, Kaluiti Stream, high
plateau, alt. 4,800 ft., central Kauai, Sept. 22, 1909 (Bish.).

Distribution: Known only from type locality in central Kauai.

Specimens examined: Rock 4,902 pro parte (type, Bish., my
photograph no. 4,203).

The leaflets are more gradually attenuate toward their tip than
in var. acrior and the submature drupes, of which four are preserved
on the type sheet, are thick-urceolate (more as in T. waimeae
Wawra), not cylindric-prismatic as in that variety. Rock (loc. cit.,
also Indig. Trees Haw. Isls. 343. 1913) did not attempt delimitation
of these varieties urceolata and acrior from T. waialealae proper,
having collected them all within a distance of about 3 km. from each
other. Study of a topographic map reveals, however, that the three
habitats involved differ sharply in conditions of altitude, drainage,
etc. (Regarding the whole subject of ecological barriers, some of
which unquestionably exist between the three type habitats, see
Riley, The American Naturalist 86: no. 826: 23. 1952.)

Tetraplasandra waialealae var. 7. acrior Sherff, Bot. Leafl.
no. 6: 22. 1952.

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 75

Leaflets 5-7, oblong or at times somewhat ovate or very slightly
obovate, toward apex narrowed and very sharply acuminate, blade
8 X 3 cm. up to 18 X 7 cm. Drupes when submature cylindric-
prismatic and 1.5-2 cm. tall including the conical stylopodium,
stigmas 5 or 6.

Type: Joseph F. Rock 4,902 pro parte, Kailiili Stream, Keaku
Cave, Isl. Kauai, September, 1909 (Gray).

Distribution: Central Kauai.

Specimens examined (all from Isl. Kauai) : Albert S. Hitchcock
15,481, alt. 3,600-5,080 ft., Waialeale, Oct. 22-24, 1916 (U.S., my
photograph no. 4,206); Rock, Isl. Kauai, October, 1916 (Bish.,
my photograph no. 4,205); Rock 4,901, near summit of Waialeale,
Sept. 22, 1909 (Arn.) ; Rock 4,902 pro parte (type, Gray, my photo-
graph no. 4,202).

For remarks, see under var. /3. urceolata above.

Tetraplasandra waialealae var. 8. wahiawensis Sherff, Bot.
Leafl. no. 6: 24. 1952.

Dwarf tree. Leaves submembranaceous, 2-3 dm. long, 7-11-
foliolate, petiole 5-8 cm. long; leaflets ovate to oblong, at apex
subacuminate or subacute to obtuse, at base oblique and very
broadly cuneate, blade 5.5-9 cm. long and 2.5-5 cm. wide; petiolules
now (for the type) 2-9 mm. now 1.2-2.6 cm. long. Branches of
inflorescence: primaries (peduncles) 3, finally robust, 9 cm. long
and 7-9 mm. thick; secondaries (rays) 9, finally robust, about
4.5-6.5 cm. long and under 4 mm. thick; pedicels about 3-5, finally
robust and arcuate, 2-2.7 cm. long and 2.5-3 mm. thick.

Type: Charles N. Forbes 274-K, Wahiawa Swamp, Aug. 11,
1909 (Bish.).

Distribution: Southern Kauai.

Specimens examined (all from southern Kauai): Forbes 2QQ-K,
several trees, low, 4 ft. tall, flat-topped, petals 5, stamens 20, ovary
5-celled, on edge of Wahiawa Swamp, August, 1909 (topotype,
Bish.); Forbes 274-K (type, Bish., my photograph no. 4,204); Amos
A. Heller 2,734 pro parte, along the Hanapepe River, near the Falls,
Aug. 14, 1895 (Gray, my photograph no. 4,208); Heller 2,734 pro
parte, in and near a bog at the head of Wahiawa Stream, Aug. 21,
1895 (N.Y.; Univ. Calif.); Heller 2,734 pro parte, between the Hana-
pepe and Wahiawa rivers, Aug. 24, 1895 (Mo.) ; Heller 2,734 pro parte,
in and near a bog at the head of Wahiawa Stream, Oct. 19, 1895

76 FIELDIANA: BOTANY, VOLUME 29

(U.S., my photograph no. 4,207); Rev. John M. Lydgate, Wahiawa
Mts. (Bish., my photograph no. 4,19s). 1

Tetraplasandra waialealae var. e. subsessilis Sherff, Bot.
Leafl. no. 6: 23. 1952.

Leaves 3-4.5 dm. long, 11- or 13-(rarely 15-)foliolate; petiole
stout, 6-9 cm. long; leaflets narrowly to moderately oblong, at
apex (where often bent in the dry state) subacuminate or rarely
obtuse, at base oblique, coriaceous, sessile to shortly petiolulate,
petiolules 1-10 mm. long, blade 8-13 cm. long and 3.5-5 cm. wide.
Peduncles 9-13 cm. long, stout; secondary branches now 3 or 4 (as
on the type) now 8, straight or curved, 4-7 cm. long; pedicels
3-6, now all straightish (as on the type) now some of them curved,
0.8-2 cm. long. Mature drupes sometimes 5- but commonly 6-ribbed
otherwise ovoid to subfusiform, 19-21 mm. tall to calyx-limb (this
spreading, brown in dried state, about 1 mm. wide); stylopodium
conical, reaching about 5 mm. above calyx, bearing usually 6 much
less often 5 or very rarely 7 stigmas.

Type: Lucy May Cranwell, Olof H. Selling, & Carl Skottsberg
2,925, forests in the vicinity of Lehua makanoi, Alakai, Isl. Kauai,
Aug. 13, 1938 (Bish).

Distribution: In the swampy areas of northwestern Kauai.

Specimens examined (all from northwestern Kauai) : Cranwell et
alii 2,925 (type, Bish., my photograph no. 4,197: isotype, Goth.,
my photograph no. 4,200); Otto Degener 21,473, in open forest at
3,000 ft., east rim of Kalalau Valley, Dec. 27, 1951 (Bish.; Chi.);
Degener & Greenwell 21,725, a 10-foot tree, among shrubs and trees,
Alakai Swamp, Jan. 19, 1952 (Chi.); Otto Degener & Henry Wiebke
2,183, sunny, open swamp, Waineke Swamp, Kokee, June 28, 1926
(N.Y., 2 sheets, my photograph no. 4,199); Fagerlind & Skottsberg
6,587, near Kawaikinana Stream, Kokee-Mohihi Road, Mar. 13,
1948 (Stockh.) ; L. H. MacDaniels 811, slender tree 4 m. tall, swampy
ground in mixed forest, Alakai Swamp, alt. 1,300 m., Feb. 17,
1927 (Bish.).

1 Forbes 205-K, vicinity of Wahiawa Swamp, August, 1909 (Bish.), is a spray
from a seedling. One leaf is about 2.5 dm. long, its 5 leaflets narrowly ovate to
lanceolate-oblong and acute to sharply attenuate-mucronate. A second leaf is
simple, its ovate-oblong blade 14 cm. long and about 7.7 cm. wide. The third
is simple, but the blade, about 11 cm. long, has on each side a large basal lobe
2.5-3 cm. long, giving an over-all width of more than 1.1 dm. The plant is doubt-
less to be referred to the present var. wahiawensis.

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 77

Skottsberg (Meddel. Goteb. Bot. Tradg. 15: 415. 1944) gave a
special description of Cranwell et al. 2,925, but referred it to the
species proper. He apparently overlooked the subsessile and coria-
ceous nature of the leaflets, which he did note, however, as being
"11-13 instead of 5-9."

4. Tetraplasandra pupukeensis Skottsb. Meddel. Goteb.
Bot. Tradg. 10: 149, fig. 17. 1936.

a. Leaflets strongly reticulate- and depressed- veiny on upper surface,

pedicels numerous (13-30), 1.4-4 cm. long.

6. Leaflets ovate to oblong-ovate, large, the terminal one up to
1.5 dm. long and 9 cm. wide, lateral leaflets 5-9 cm. wide.

var. /3. venosa.
6. Leaflets usually narrower and smaller, lateral leaflets commonly

2-4 (rarely -6) cm. wide.

c. Lateral leaflets oblong or oblong-oval, apically subacuminate
to truncate or rounded or even emarginate, usually above
3.5 cm. wide; stylopodium 3 mm. tall.

var. 7. megalopoda and f. trigona.

c. Lateral leaflets oblong-lanceolate or oblong-rhomboid to nar-
rowly ovate, apically acute to acuminate, commonly
2-3.5 cm. wide; stylopodium 1.5-2 mm. tall.

var. e. decipiens.
a. Leaflets smoothish to weakly veiny on upper surface.

6. Leaflets or some of them acutish to acuminate at tip, commonly
glossy above at least when dry; umbellets 10-25-flowered.

var. d. nitida.

b. Leaflets obtuse to rounded or truncate at tip, dull or but
weakly glossy above, umbellets mostly 10-12-flowered.

var. a. pupukeensis.

Small tree. Leaves 2-3.5 dm. long, 5-7-foliolate; petiole 4-9.5
cm. long; leaflets at points 3-5 cm. apart on rhachis, pale and at
times glossy, on upper surface obsoletely to moderately reticulate-
venulose, blade of terminal one about 8-12 X 5-8 cm., ovate to
obovate, at apex broadly rounded to subtruncate and more or less
emarginate, on a stipe 1-3 crri. long, this subarticulate near its
upper end; blade of lateral ones ovate-elliptic or rotund-elliptic,
very obtuse at each end and at base often oblique, for the inter-
mediate ones 8-12 X 5.5-7 cm., for the lowermost ones 5-8 X 4-5.5
cm.; petiolules 2-10 mm. long. Inflorescence a twice compound

78 FIELDIANA: BOTANY, VOLUME 29

umbel at end of a leafy branch, peduncles 5, with scattered nodes,
5-11 cm. long and 2-4 mm. thick, straight or barely curved; pedicels
about 10-12, angulate, becoming more or less robust and 1.5-2.2
cm. long. Calyx at anthesis obconic to urceolate, 7-9 mm. tall and
dz6 mm. broad at top. Petals 6 or 7, ovate-triangular, at apex
cucullate, 8-9 mm. long, at base 3-3.5 mm. wide, fleshy, rosaceous,
presently deciduous. Stamens about 20, the outer about 5.5, the
inner about 4.5 mm. long, fleshy; anthers 3.5-4 mm. long, curved
at top. Stylopodium conic, 2 mm. tall; stigmas commonly 6, sub-
sessile. Drupes ovoid-urceolate, obtusely 5-7-angled and deeply
5-7-sulcate; calyx 11-12.5 mm. tall and 8-10 mm. thick, with con-
spicuous, spreading border 1 mm. wide for its limb; stylopodium
2-2.5 mm. tall and thick, sharply 5-7-rayed, the 5-7 stigmas rather
sharply distinct.

Type: Carl Skottsberg 1,816, alt. about 300 m., Pupukea-Malaeha-
hana trail, "Pupukea Forest Reserve," Sept. 15, 1926 (Goth.).

Distribution: Extreme northern Oahu.

Specimens examined (all from Oahu): Edward Y. Hosaka 124,
alt. 1,800 ft., Pupukea-Paumalu Forest Reserve, Jan. 12, 1930
(Bish.); Kazuto Nitta 76, moderately dry slope, Pupukea, Jan. 12,
1930 (N.Y., my photograph no. 4,291); Skottsberg 1,816 (type,
Goth., my photograph no. 4,177: isotype, Bish., my photograph
no. 4,218).

Tetraplasandra pupukeensis var. ft. venosa Deg. & Sherff
ex Sherff, Bot. Leafl. no. 6: 40. 1952.

Blades of leaflets thicker, strongly reticulate- and depressed -
veiny on upper surface, ovate to oblong-ovate, apically obtuse to
rounded, basally roundish or more often very broadly cuneate,
terminal one up to 1.5 dm. long and to 9 cm. wide, lateral ones
mostly a third smaller in length and width. Inflorescence twice
umbellate, peduncles 3, about 7-13 cm. long, robust (5-7 mm.
thick), more or less scurfy; pedicels 13-28 in a terminal umbellet
or a single one placed 1-2 cm. lower down, 1.4-2.8 cm. long, sub-
robust. Calyx obconic at anthesis, about 7 mm. tall and at its limb
about 5 mm. wide; petals about 8; stamens 16; drupes (only
somewhat immature ones seen) cylindric-prismatic, 5- or commonly
6-ribbed, calyx about 12 mm. tall and 7-8 mm. thick, stylopodium
conic, its height about 2 mm. and diameter at base 3 mm., stigmas
5 or more often 6.

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 79

Type: Otto Degener, in gully southeast of Kahuku entrance of
Pupukea-Kahuku Trail, Isl. Oahu, Nov. 22, 1931 (N.Y., 2 sheets).

Distribution: Northern Oahu.

Specimens examined (all from Oahu) : Degener, Isl. Oahu (N.Y.,
2 sheets) ; Degener, Pupukea-Kahuku, July 19, 1931 (N.Y.) ; Degener,
in gully southeast of Kahuku entrance of Pupukea-Kahuku Trail,
Nov. 22, 1931 (type, N.Y., 2 sheets, my photographs no. 4,224 and
no. 4,225).

Tetraplasandra pupukeensis var. 7. megalopoda Sherff,
Bot. Leafl. no. 6: 40. 1952.

Leaflets rather small; their blade variously oblong, oblong-oval,
or (especially for terminal leaflet) obovate to subrhomboid-obovate,
apically subacuminate to truncate or rounded, even emarginate,
basally cuneate-contracted, under 11.5 cm. long and 6 cm. wide,
conspicuously reticulate- (and on upper surface depressed-) veiny,
more or less glossy above. Peduncles large, 1.7-2.6 dm. long and
4-9 mm. thick, at tip 30-pedicellate, perhaps solitary; pedicels
somewhat slender, straight to curved or rarely a few deflexed-
ascending, 2.5-4 cm. long. Calyx at anthesis obconic, about 8 mm.
tall and at limb about 6 mm. broad; petals 8, stamens 16;
drupes (only immature ones known) with a cylindric-prismatic calyx,
this ribbed, about 11 mm. tall and at limb 7 mm. thick; stylopodium
conic, 3 mm. tall, stigmas commonly 5 rarely 6.

Type: David LeRoy Topping 2,921, Pupukea-Kahuku Trail,
Nov. 23, 1924 (Univ. Calif.).

Distribution: Northern Oahu.

Specimens examined (all from Oahu) : Otto Degener 2,921, Pupu-
kea-Kahuku Trail, Nov. 23, 1924 (topotype, N.Y.); Degener 17,792
pro parte, along summit-divide toward east of Pupukea-Kahuku
Trail, shrubby rain-forest, Dec. 6, 1931 (topotype, N.Y., my photo-
graph no. 4,272) ; Degener 18,250 p. p., forest, directly mauka of Kawela
Bay, Apr. 2, 1933 (N.Y.) ; Degener & Emilio Ordonez 12,066, forest,
C.C.C. Trail, Kawailoa, July 3, 1938 (Mo.; N.Y.; U.S.); Topping
2,921 (type, Univ. Calif., my photograph no. 4,222).

Tetraplasandra pupukeensis var. megalopoda f. trigona
Deg. & Sherff ex Sherff, Bot. Leafl. no. 7: 7. 1952. 1

1 Through typographical error, the joint authorship Degener & Sherff was
erroneously applied (loc. cit.) to the varietal epithet, megalopoda, whereas it was
intended for the formal epithet, trigona.

80 FIELDIANA: BOTANY, VOLUME 29

Pedicels barely shorter, more or less deflexed-ascending; drupes
commonly tri- less often tetragonous, stigmas commonly 3 less
often 4.

Type: Degener 17,792 pro parte, along summit-divide toward
east of Pupukea-Kahuku Trail, shrubby rain-forest, Isl. Oahu, Dec.
6, 1931 (Mo.).

Distribution: Known only from type locality in northeastern
Oahu.

Specimens examined (Isl. Oahu) : Degener 17,792 pro parte (type,
Mo., my photograph no. 4,298); Degener 18,248, rain-forest at
summit-divide toward east of trail, Pupukea-Kahuku Dec. 6, 1931

(N.Y.).

Degener 17,792 was previously cited (Bot. Leafl. no. 6: 41. 1952)
among the specimens of var. megalopoda. The stigmas on the few
immature fruits present numbered in some cases 4 in others 5.
Recently the fine sheet from the Missouri Botanical Garden came
to hand, with an abundance (45) of mature drupes. These were
almost uniformly 3-stigmatate and 3-angulate, this being exceptional
in T. pupukeensis, the varieties of which normally show 4, 5, or 6
stigmas and drupe-angles.

Tetraplasandra pupukeensis var. <5. nitida Deg. & Sherff
ex Sherff, Bot. Leafl. no. 6: 39. 1952.

Petiole up to 12.5 cm. long, petiolules of lateral leaflets 4-12
mm. long; blades of leaflets variously oblong, lanceolate-oblong,
ovate, even subovate, at apex variously obtuse to acuminate, at
base contracted or rarely roundish, commonly glossy above when
dry, lateral nerves more often conspicuous. Inflorescence sessile,
twice umbellate; peduncles 4, curved or straight, 8-16 cm. long,
subrobust (about 5-6 mm. thick), 10-25-flowered ; pedicels 1.5-2
cm. long, robust, about 2 mm. thick. Calyx at an thesis obconic,
7 mm. tall and near top 5 mm. broad; petals about 8-10; stamens
16-28; stigmas 5; drupes unknown.

Type: Degener, Murashige, & Ken 21,017, tree 20 ft. tall, in
rainy forest, alt. 3,500 ft., Mt. Kaala, Isl. Oahu, July 10, 1949 (Chi.).

Distribution: Known only from type locality on Mt. Kaala,
northwestern Oahu.

Specimens examined: Degener, Murashige, & Ken 21,017 (type,
Chi., my photograph no. 4,271: isotypes, Arn.; Bish.; Gray; N.Y.;
Phila.;Univ. Calif.; U.S.).

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 81

Tetraplasandra pupukeensis var. e. decipiens Deg. & Sherff
ex Sherff, Bot. Leafl. no. 8: 8. 1953.

Similar to var. megalopoda in its peduncles, florets, and drupes,
but differing in its 5-9 leaflets, these smaller, oblong-lanceolate or
oblong-rhomboid to narrowly ovate or (the terminal one) obovate,
at apex acute to acuminate, at base more or less attenuate or cuneate
and oblique, blade 4.5-10 cm. long and 2-3.5 cm. wide, lateral petio-
lules 3-11 mm. long. Umbellets about 18-22-flowered, pedicels
16-20 mm. long. Stamens 10-12. Drupes 4-5-angled, prismatic,
the calyx about 1-1.1 cm. tall and 5-6 mm. thick, the stylopodium
small-conic and 1.5-2 mm. tall including the 4-6 stigmas.

Type: Otto Degener 17,974, windy rain-forest near top, Koolau
Mountains, northern Oahu, Aug. 25, 1935 (N.Y.).

Distribution: Northern Oahu.

Specimens examined (northern Oahu): Degener 17,791 p. p., in
shrubby rain-forest, growing with but rarer than no. 17,792 (under his
no. 17,792 Degener collected two forms: T. pupukeensis var. megalo-
poda and its f. trigona), along summit-divide toward east of Pupukea-
Kahuku Trail, Dec. 6, 1931 (N.Y.); Degener 17,974 (type, N.Y.,
my photograph no. 4,354) ; Degener 18,249 p. p., rain-forest at summit-
divide toward east of trail, Pupukea-Kahuku, Dec. 6, 1931 (N.Y.,
my photograph no. 4,355) .

The slender leaflets, with their more attenuate tips and bases
and their veiny, coriaceous upper surfaces, offer a deceptive re-
semblance to those of T. oahuensis var. venulosior (whence the
varietal name). The various characters of flowers and fruits, how-
ever, are more in agreement with those of T. pupukeensis, particularly
the var. megalopoda.

5. Tetraplasandra lihuensis Sherff, Bot. Leafl. no. 8: 8. 1953.

Leaflets coriaceous, at base more often truncate or barely sub-
cordate, peduncles nodeless, pedicels robust var. a. lihuensis.

Leaflets submembranaceous, at base contracted or rounded but not
truncate, peduncles mostly with node near middle, pedicels slender
at least in fruit var. 0. gracilipes.

Leaves glabrous, 5 dm. long including petiole (this 11 cm.
long and toward base 1 cm. thick), 13-foliolate; leaflets coria-
ceous, more or less oblong (the smaller terminal one ovate), sub-
attenuate above middle, at apex subacute to obtuse, at base some-
times oblique and more often truncate or barely subcordate, blade

82 FIELDIANA: BOTANY, VOLUME 29

10-19 cm. long and about 6.5-9 cm. wide; lateral petiolules sub-
robust, 5-15 mm. long. Inflorescence thrice umbellate; peduncles
3, umbellately grouped, 1-1.5 dm. long and 5 mm. thick,
irregularly scaly, without nodes, at apex umbellately 6-radiate;
rays very scaly, sharply angled, without nodes, 4-6.5 cm. long,
somewhat spreading to suberect, at apex umbellulately about
7-9-pedicellate; pedicels robust, angulate, at anthesis about 8-13
mm. long. Calyx cylindric-obconic near anthesis, about 5 mm. tall
and at middle about 2-3 mm. thick, its limb membranaceous, un-
dulate, and broadly (about 2 mm.) extended. Corolla not yet ex-
panded, about 6-7 mm. tall and thick, petals 7 or 8. Stamens
numerous, in several series. Stigmas about 8 (in young flowers).
Drupes not seen.

Type: Charles N. Forbes 164-K, Power-Line Trail from Lihue,
southeasternmost Kauai, Aug. 6-9, 1909 (Bish.).

Distribution: Southeastern Kauai.

Specimens examined: Forbes 164-lf, (type, Bish., my photograph
no. 4,196); Forbes 286-K, tall tree, leaves glaucous, Wahiawa Moun-
tains, August, 1909 (Bish.).

Both of the specimens examined had their leaflets numerously
but irregularly black-punctulate, but this character seemed due to
some fungus growth.

Tetraplasandra lihuensis var. /3. gracilipes Sherff, Bot.
Leafl. no. 8: 9. 1953.

Leaves ll-foliolate, leaflets submembranaceous, basally con-
tracted or rounded but not truly truncate; lateral petiolules slender,
8-18 mm. long. Peduncles 4, slender, about 7-9 cm. long and
2-3 mm. thick, more often with a single node near middle, at tip
about 8-14-rayed; rays slender, without nodes, 2-4 cm. long, at tip
few- or several-pedicellate; pedicels finally slender, 8-13 mm. long
(only 4 seen). Flowers not seen. Drupes ovoid-globose, calyx 8
mm. tall, glabrous, limb obsolete; stylopodium convex-hemispherical
or mitriform, 2-3 mm. tall (on type specimen densely and minutely
erect-hispidulous), abruptly contracted into a style about 1 mm.
long; stigmas 4 or imperfectly 5.

Type: L. H. MacDaniels 847, tree 10 m. tall, in rain-forest, alt.
850 m., upper Lihue ditch, southeasternmost Kauai, Feb. 21, 1927

(Bish.).

Distribution: Southeasternmost Kauai.

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 83

Specimens examined: MacDaniels 847 (type, Bish., my photo-
graph no. 4,356).

Unfortunately, T. lihuensis proper is as yet unknown in its
fruiting state, nor is var. gracilipes known so far in its flowering
state; hence, association of the two as members of the same species
is somewhat arbitrary. It seems doubtful, however, that var.
gracilipes will be found more different from T. lihuensis proper than
are the varieties of the large-flowered T. waialealae from that species.

The stylopodium of the mature drupes of var. gracilipes is
hispidulous. Small hairs can be found even at the limb of the
fruiting calyx. Most of the hairs appear to belong there naturally
(if so, affording a most anomalous character) rather than to have
resulted from some fungus growth.

6. Tetraplasandra kohalae Skottsb. Meddel. Gb'teb. Bot.
Tradg. 15:414. 1944.

Small tree. Leaves 2.5-4 dm. long, 5-7-foliolate; petiole 7.5-11.5
cm. long; blade of leaflets elliptic-oblong, at apex shortly to even
subobtuse or obtuse, at base rounded and for the lateral leaflets
somewhat oblique, on both surfaces clearly reticulate-nerved with
nerves (except for the midnerve) depressed, about 15 secondaries
at each side of midnerve, 6.5-12 cm. long and 3.5-5.5 cm. wide (for
random leaflets selected, the blade measuring 12 X 5, 11.6 X 5.5,
11 X 5.2, 9 X 4.4, 8.5 X 4, 8 X 4.4, 7 X 3.8, and 6.5 X 3.3 cm.);
petiolule 8-14 mm. long. Inflorescence of one or perhaps more um-
bellets each on a peduncle 1 dm. long; umbellets 23-28-fruited
(flowers unknown as yet), pedicels finally 1.5-1.8 cm. long. Drupe
large, ovoid-cylindric, 22 mm. tall and about 8-10 mm. thick
(submature ones of type collection having calyx 14-16 mm. tall
and stylopodium adding 2 mm. more, thickness of about 7-8 mm.),
the calyx-limb narrow, thickened-undulate; stylopodium 3 mm. tall,
depressed-conic, thus barely sloping to the very short style, this
with 2 or 3 stigmas.

Type: Lucy May Cranwell, Olof H. Selling, & Carl Skottsberg
3,151, close to second ditchman's house, Upper Hamakua ditch
trail, at head of Alakahi Valley, Kohala Mountains, Isl. Hawaii,
Sept. 7, 1938 (Goth.). 1

Distribution: Known only from type locality in northern Hawaii.

1 The type number given on the original labels was 3,151, not 3,156 as er-
roneously printed in Skottsberg's original text.

84 FIELDIANA: BOTANY, VOLUME 29

Specimens examined: Cranwell et alii 3,151 (type, Goth., my
photograph no. 4,176: isotype, Bish., my photograph no. 4,217).

7. Tetraplasandra lanaiensis Rock, Coll. Haw. Publications
Bull. 1:12, pi. 2. 1911.

Small tree about 5 m. tall, glabrous throughout, freely but
irregularly branching, with smooth, whitish bark. Leaves 3-4 dm.
long; leaflets 5-7, dark-green above and lighter beneath, subcoria-
ceous, the lateral with blade oblong or less often narrow-obovate,
at apex obtuse to abruptly subacuminate, at base subrotund or
more often broadly cuneate, 8-10 cm. long and 4-6 cm. wide,
petiolule 6-15 mm. long; terminal with blade more or less obovate,
at times a little larger, on a petiolule 2-4 cm. long and more or less
plainly articulate toward blade. Inflorescence thrice umbellate on
a rhachis 2.5 cm. long, not erect but drooping; peduncles 3-5,
usually 1.5-2.3 dm. long, bearing umbels of about 17-21 drooping
2- or 3-nodate rays, these mostly 7-9 cm. long and bearing 7-13
flowers normally in a terminal umbellet (rarely 1 or 2 placed lower
down) ; pedicels (at anthesis) slender, 1-1.3 cm. long. Calyx tubular-
obconic, about 6 mm. tall, with a denticulate border; petals 5 or 6,
lanceolate, greenish-yellow, about 7 mm. long; stamens 10 or 12;
ovary 3-celled; stigmas sessile. Drupe unknown.

Type: Joseph F. Rock 8,088, not common, only two trees ob-
served, on dry forehills in Kaiholena Valley, 1 associated with Osman-
thus sandwicensis, Suttonia lanaiensis, Sideroxylon spathulatum, Maba
sandwicensis, etc., alt. 2,000 ft., Isl. Lanai, Aug. 2, 1910 (Herb.
College Hawaii).

Distribution: Known only from central Lanai.

Specimens examined (all from Isl. Lanai): Charles N. Forbes
307-L, Lanai, September, 1917 (Bish.); Forbes 386-L, tree 9-10 ft.
tall, leaves glossy above, smooth and pale below, petals yellow, 6,
stamens 17 or 18 in 2 series, ovary 4-celled, Kaiholena, September,
1917 (Bish.); Forbes 388-L, Lanai, September, 1917 (Bish.); George
C. Munro, Lanai, Aug. 31, 1915 (U.S.); Munro, Kaiholena, Mar.
18, 1922 (U.S.); Munro 335, Hookio, Aug. 21, 1915 (Bish.); Munro
461, Isl. Lanai, same date (Bish.); Rock 8,088 (isotypes, Arn., 2

1 In his original text and on his labels, Rock gave the broader term "Mahana
Valley," but later (Indig. Trees Haw. Isls. 343. 1913) wrote: "The tree . . . was
seen only in Kaiholena Valley, crowded by other trees at an elevation of 2,000
feet. Kaiholena Valley, belonging to the drier regions of Lanai, is extremely
interesting and harbors a very multiformous tree flora."

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 85

sheets; Bish.; Gray, 4 sheets, my photographs no. 4,173 and no.

4,174).

Unfortunately, T. lanaiensis is not as yet known in its fruiting
state, while the apparently close T. Munroi is not as yet known in a
flowering state. In T. Munroi, however, the rays of the inflorescence
are without nodes, while in T. lanaiensis they have 2 or 3 nodes
each, this character alone making distinction easy. From T. mei-
andra var. Hillebrandii, apparently frequent on Lanai, T. lanaiensis
can be distinguished, in the absence of flowers and fruits, by its
less coriaceous, less venulose, apically more tapering or often sub-
acuminate leaflets.

8. Tetraplasandra Munroi Sherff, Bot. Leafl. no. 7: 7. 1952.

Tree; leaves (a single one seen so far) 4 dm. long (petiole
12 cm. long included); leaflets 9, petiolule 7-12 mm. long; blade
coriaceous, oblong-ovate to oblong or (for terminal leaflet) even
oblong-obovate, 7-12 cm. long and 3-6 cm. wide, at apex obtuse or
even emarginate, at base rounded to broadly cuneate, scarcely
oblique. A single umbel seen, this twice compound, finally about
1.6 dm. wide, its stem bent, slender (3-4 mm. thick) and 19 cm.
long; rays 20, slender (about 0.8-1.5 mm. thick), sharply angulate,
entirely without nodes, straight or arcuate, 5-6.5 cm. long; pedicels
commonly 6-8, slender or even filiform, straight or more often ar-
cuate, very sharply angulate, finally 11-16 mm. long. Flowers not
seen. Mature drupes drying to a grayish-brown, now 3- now 4-angu-
late, deeply and broadly grooved between the angles, 7-10 mm.
long (including the small, conic stylopodium and 3 or 4 stigmas)
and 4.5-5.5 mm. thick.

Type: George C. Munro 694, Kaiholena Valley, Isl. Lanai, Mar.
18, 1922 (Bish.).

Distribution: Known only from type locality in central Lanai.

Specimens examined: Munro 694 (type, Bish., my photograph
no. 4,172).

The type had been referred interrogatively to T. lanaiensis
Rock, a species known as yet only from the same general locality
in central Lanai. That species, known so far only in the flowering
state, differs in having smaller leaflets, these less blunt at the tip
and more gradually narrowed to a more oblique base, their petiolules
shorter and broader; the rays definitely 2- or 3-nodate and a third
or so longer. T. lanaiensis has, moreover, a mammoth, thrice-

86 FIELDIANA: BOTANY, VOLUME 29

compound umbel 4 dm. broad. T. Munroi should be restudied in
the field as to flowers and as to the nature of a complete inflorescence.
The detached, twice-compound umbel described from the type
sheet may, with its elongate stalk, prove to have been just one main
part of a thrice-compound umbel that had seemed too large for a
herbarium specimen. It may be remarked here that collectors of
Tetraplasandra specimens should be especially careful to preserve
enough of the complete inflorescence down to and including the
nearest leaf-petioles, to permit correct conclusions as to how com-
pound or decompound it really is.

9. Tetraplasandra Lydgatei (Hillebr.) Harms in Engl. &
Prantl, Pflanzenfam. Ill, 8: 20. 1898; Triplasandra Lydgatei Hillebr.
Fl. Haw. Isls. 153. 1888.

a. Leaflets glossy and leathery on upper surface .... var. e. coriacea.
a. Leaflets otherwise.

6. Inflorescence sessile or subsessile, its thick, scarcely visible
footstalk (rhachis) 1 cm. tall.

c. Peduncles 13 (in herbarium specimens some may have
been cut away), about 9.5-12 cm. long and 1.5-3 mm.
thick, 20-30-flowered, pedicels 15-22 mm. long.

var. 7. Forbesii.

c. Peduncles about 4-6, only 3.5-6.5 cm. long, more slender,

10-20-flowered.

d. Peduncles up to about 18-20-flowered, pedicels finally
8-10 mm. long; drupes 5-6 mm. tall and thick, mostly
3-angled var. /3. brachypoda.

d. Peduncles about 12-flowered, pedicels finally 13-18 mm.
long; drupes ovoid-globose, 8-10 mm. thick and some-
what taller, obtusely 4-angled var. a. Lydgatei.

b. Inflorescence on a definite, slenderish footstalk 1.5-2.5 cm. tall.

var. 5. leptorhachis.

Small tree, quite glabrous. Leaves 2-3 dm. long, 5-11-foliolate;
petiole 8-9 cm. long; blades of leaflets elliptic-oblong or at times
narrowly obovate, at tip obtuse to truncate and slightly emarginate,
below middle gradually narrowed to a frequently oblique base,
thin-chartaceous, 6-10 cm. long and 2-4.5 cm. wide; petiolules
delicate, 8-23 mm. long. Inflorescence twice-umbellate from a short,
common rhachis of about 13 mm.; peduncles 4 or 5, slender, 6.5

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 87

cm. long, bearing terminal umbellets of about 12 pedicels, these
slender and about 13-18 mm. long. Calyx broad-obconical, 5 mm.
tall, with undulating border. Petals 5, cohering at their tips, 6
mm. long. Stamens 12, about 4 mm. long, anthers straight or
recurved. Ovary 4-celled, its disk slightly raised with 4 sessile
stigmas. Drupe ovoid-globose, 8-10 mm. in diameter, obtusely
4-angled, the short conical apex finally elongated into a short
stylopod.

Type: Rev. John M. Lydgate, Wailupe, Isl. Oahu (formerly in
Berl.; see remarks below).

Distribution : Known only from type locality in southeasternmost
Oahu.

Specimens examined (Isl. Oahu): Charles N. Forbes 2,511-0,
Wailupe Valley, May 4, 1917 (topotype, Bish., my photograph no.
4,293); Hillebrand & Lydgate, Wailupe (Bish., a leaf and peduncle
with 7 drupes, ex herb. Lydgatei, type collection, my photograph
no. 4,186).

This species was founded by Hillebrand upon a plant attributed
to Lydgate and which had been collected at Wailupe, southeastern-
most Oahu. The type was in the Berlin Herbarium and perished
with the destruction of that herbarium in World War II. I have
before me, however, a sheet from the Bishop Museum Herbarium,
bearing a 7-foliolate leaf and a single peduncle about 6.5 cm. long,
with seven pedicellate fruits still attached. These were from Lyd-
gate's own herbarium and are authentic for the species. The original
label gives "Niu or Wailupe" but presumably Hillebrand, who
collected jointly with Lydgate many of the specimens that went
into Lydgate's herbarium, 1 had definite knowledge that Wailupe
was the real locality. The pedicels are slender and 13-16 mm.
long. Hillebrand (loc. cit.} gave the length as 16-18 mm. ("8-9
lines"). The species in its narrower sense seems never to have been
collected again in recent years except for Forbes 2,511-0.

Tetraplasandra Lydgatei var. /3. brachypoda Sherff, Bot.
Leafl. no. 6: 19. 1952.

Peduncles normally 4-6, about 3.5-5.5 cm. long and 1-1.5 mm.
thick, up to about 18-20-flowered, pedicels slender and finally 8-10

1 Indeed, on the label itself, Hillebrand and Lydgate are both named as
the collectors. (For a special note regarding a "collection of Hawaiian plants
collected or named by Dr. Hillebrand" and possessed by Mr. Lydgate, see Occas.
Papers, Bishop Mus. 6: 5: 8. 1918).

88 FIELDI ANA: BOTANY, VOLUME 29

mm. long; drupes when dry 5-6 mm. thick and tall, a few 4-angled
the rest 3-angled.

Type: D. LeRoy Topping & Will Bush 3,701, tree 30-40 ft. tall,
same one as for Degener et al. 6,099, Dec. 3, 1933 (N.Y.).

Distribution: Southeasternmost Oahu.

Specimens examined (all from Isl. Oahu) : Otto Degener 18,244,
moderately dry, open forest, western slope of central ridge of Niu
Valley, June 4, 1932 (topotypes, Mo.; N.Y.); Degener, Swezey, Park,
& Nitta 6,099, in moderately dry, open woods on side of trail on
western slope of central ridge of Niu Valley, June 4, 1932 (topotypes,
Gray; N.Y.; U.S.); Charles N. Forbes with J.C. Bridwell 2,461-0,
ridge between Niu and Wailupe, Apr. 11, 1917 (Bish.); Topping &
Bush 3,701 (type, N.Y., my photograph no. 4,185: isotypes, Mo.;
N.Y., 2 sheets; U.S., my photograph no. 4,189).

The foliage of herbarium specimens examined is notably pallid
and many of the leaves are 11- or 13-foliolate. Degener (Fl. Haw.
fam. 281. June 29, 1934) has presented a beautiful plate of this
variety, drawn from one of the Topping & Bush specimens. With
him, however, it was assumed to represent T. Lydgatei proper.

Tetraplasandra Lydgatei var. 7. Forbesii Sherff, Bot. Lean 1 ,
no. 6: 20. 1952.

Tree 6 m. tall, bark dark or blackish, leaves 7-foliolate. Pe-
duncles 13 to an umbel, 9.5-12 cm. long and 1.5-3 mm. thick,
drying reddish-brown, 20-30-flowered ; pedicels very slender, com-
monly 15-22 mm. long; drupes ovoid-globose, sharply 3- or most
often 4-angled, 4-6 mm. thick and 8 mm. tall including stylopodium
and stigmas.

Type: Charles N. Forbes 1,942-0, tree about 20 ft. tall, darkish
bark, leaves with 7 leaflets . . . turning yellow with age . . . ridge
west of Waialae Valley, near the top, rather dry locality, Isl. Oahu,
Oct. 15, 1914 (U.S.).

Distribution: Known only from type locality, southeastern
Oahu.

Specimens examined (Isl. Oahu) : Forbes 1,942-0 (type, U.S., my
photograph no. 4,187: isotype, Bish., a mere fragment, but with
fuller data) ; L. H. MacDaniels 466, Palolo- Waialae Ridge, Jan. 27,
1927 (topotype, Bish.).

The elongate, numerous peduncles and pedicels at once dis-
tinguish this variety.

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 89

Tetraplasandra Lydgatei var. 5. leptorhachis Deg. & Sherff
ex Sherff, Bot. Leafl. no. 6: 20. 1952.

Leaflets 5 or 7. Inflorescence a twice compound umbel on a
rhachis about 3-4 mm. thick and 1.5-2.5 cm. long; peduncles 4-8,
drying a greenish-brown, 4-7.5 cm. long and about 2 mm. thick,
18-32-flowered ; pedicels slender, 12-16 mm. long, greenish-brown
when dry; drupes ovoid, sharply 3- or 4-angulate, when submature
4-5 mm. thick and 6-7 mm. tall.

Type: Otto Degener 17,796, open, windy rain-forest, slope north-
east of Nuuanu Valley, Isl. Oahu, Nov. 20, 1926 (U.S.).

Distribution: Southeastern and central Oahu.

Specimens examined (all from Isl. Oahu) : Otto Degener, slope
northeast of Nuuanu Valley, Nov. 20, 1926 (topotype or perhaps
isotype, N.Y.) ; Degener, Wahiawa, Nov. 25, 1926 (N.Y., my photo-
graph no. 4,191); Degener 17,796 (type, U.S., my photograph no.
4,190: isotypes, Mo.; N.Y.; Yunck.).

In T. Lydgatei proper (i.e., var. Lydgatei), also in the vars.
brachypoda and Forbesii, the flowering axis or rhachis is reduced to
a short, thick stump 1 cm. or so long, a mere anchorage for the
peduncles of the compound umbel. In var. leptorhachis the rhachis
stands out distinctly as a slender stalk.

The geographic range is remarkable, extending as it does north-
westward from Nuuanu Valley in far southeastern Oahu to Wahiawa,
well past the center of Oahu. Recourse to my private copy of
Degener's "Rosetta Stone" (a key prepared by him some years ago
to summarize his collection data and deposited in my keeping)
shows that on the date given for Wahiawa, he collected "in the
mountains east of Wahiawa."

Tetraplasandra Lydgatei var. e. coriacea Sherff, Bot. Leafl.
no. 6: 21. 1952.

Leaflets 5-9, their blades elliptic-oblong, 8-12 cm. long and
3.5-5.7 cm. wide, glossy and leathery on upper surface. Peduncles
of the inflorescence (an immature one seen) 10, about 3-3.5 cm.
long, provided at the middle with a bract (this widely ovate, sharply
acuminate, 3 mm. long, finally deciduous), umbellately clustered
upon a thick rhachis about 1 cm. long, 35-flowered; pedicels
slender, about 6 mm. long, calyx about 1.5-2 mm. tall; immature
corolla as yet unopened, sharply hexagonal, about 3.2 mm. tall and
about 2.5 mm. thick.

90 FIELDIANA: BOTANY, VOLUME 29

Type: Richard S. Cowan 563, alt. 1,800 ft., Halawa Valley
Ridge Trail, Koolau Range, southeastern Oahu, Mar. 2, 1947 (Bish.).

Distribution: Known only from type locality in southeastern
Oahu.

Specimens examined (Isl. Oahu) : Cowan 563 (type, Bish., my
photograph no. 4,192).

The glossy, leathery upper surface of the leaves is very distinctive.
The packet on the type sheet contains three more or less mature
drupes, but there is nothing to indicate that they actually came from
the type tree. Their pedicels are 8 mm. long, two are 3- and one is
4-gonous, and all are about 1 cm. tall, stylopodium and stigmas
included.

10. Tetraplasandra oahuensis (A. Gray) Harms in Engler
& Prantl, Pflanzenfam. Ill: 8: 30. 1898; Gastonia? oahuensis A. Gray,
Bot. United States Explor. Exped. 726. 1854; Triplasandra oahuen-
sis (A. Gray) Seem. Jour. Bot. 6: 139. 1868; Gastonia oahuensis
A. Gray ex Hillebr. Fl. Haw. Isls. 153. 1888.

a. Inflorescence having rays between peduncle and pedicels.

6. Leaves 4.5 dm. long including petiole, or if smaller, with
rays usually 4-7 cm. long.

c. Lateral petiolules 1-4 mm. long; leaflets 5 or 7; peduncle
robust (5 mm. thick) and short (4.5 cm. long), rough
and transversely or circumferentially pocked; rays 15-
pedicellate; native of subcentral Oahu.

var. f . pseudorhachis.

c. Lateral petiolules 5-20 mm. long; leaflets 9; peduncle more

slender and elongate (8- 16 cm. long), smoothish unless

for a few bracteal scars; rays 5-12-pedicellate.

d. Leaflets 11; peduncle slender and elongate (1.6 dm.

long); rays 3-3.7 cm. long; pedicels about 5-7; native

of southeastern Oahu var. d. pseudolongipes.

d. Leaflets 7-9; peduncle about 8-10 cm. long; rays more
often 4-7 cm. long; pedicels 8-12; native of north-
westernmost Oahu var. d. hailiensis.

b. Leaves about a third shorter including 6 cm. long petiole,

leaflets 5-9.

c. Median and lower leaflets oblong to oval or obovate; rays
under 3 cm. long var. a. oahuensis.

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 91

c. Median and lower leaflets elliptic to elliptic-oblong or nar-
rowly oval; rays 3.5-4 cm. long var. e. Fauriei.

a. Inflorescence lacking rays, the pedicels attached directly to the

peduncle.

6. Leaflets for most part very coriaceous and on usually glossy
upper surface depressed-venulose; umbellets 30-flowered;
native of northeastern Oahu (Punaluu and Kaaawa).

var. rj. venulosior.
b. Leaflets otherwise; umbellets up to 20-flowered; natives

farther west or south.

c. Peduncle 5-8.5 cm. long, leaflets ovate to broad-oblong,
4-6.5 cm. broad, obtuse to subacute at apex, abruptly
broad-cuneate to subtruncate at base; petiolules 3-10 mm.

long var. 7. eradiata.

c. Peduncle 16-18 cm. long, lateral leaflets oval-oblong to
narrowly oblong or oblong-oblanceolate, mostly under 4
cm. broad, obtuse to rounded at apex, narrowly to mod-
erately cuneate at base; petiolules 7-20 mm. long.

var. j8. longipes.

Evidently arborescent, branchlets stout, marked with the leaf-
scars. Leaves 5-9-foliolate, glabrous (as is the whole plant), 22
cm. long including the slender petiole (this 6 cm. long); leaflets
moderately fleshy and thick, dull, closely but inconspicuously
feather- veined; blades 4.5-9 cm. long and 2-3.8 cm. wide, the median
ones oblong or oval, terminal and basal ones inclining toward obovate
or terminal pair (on type, U.S.) oblanceolate, all obtuse (on type)
to barely subacute (isotype, Gray) at tip and broadly to narrowly
cuneate at base; petiolules 5-8 mm. long. Umbels probably thrice
compound, but only a single, detached compressed-angled peduncle
about 9.5 cm. long present on type sheet, this having borne several
short (2.5-2.8 cm.), slender or subrobust, compressed-angled rays
(three left on type and one on isotype [Gray] at and just below its
tip) ; bracts if any caducous; umbellet at tip of ray about 6-pedicellate
(in fruit); pedicels 5-10 mm. long. Petals 5, triangular-lanceolate,
valvate in aestivation, expanding in anthesis but caducous. Stamens
10; filaments short, subulate; anthers 2-celled, oblong, opening
longitudinally. Drupes immature; their calyx- tube clavate, 4-5-
angled, 4.5 mm. tall and about 3 mm. thick at the very short,
truncate, repand limb; their ovary inferior, 4- or 5-celled, crowned
with a tiny, conical, now 4- now 5-rayed stylopodium about 1 mm.
tall, bearing an (as yet) undeveloped and entire stigma.

92 FIELDIANA: BOTANY, VOLUME 29

Type: United States Exploring Expedition under Captain Charles
Wilkes, at altitude of 2,000 feet, on the mountains behind Honolulu,
Island of Oahu (U.S.).

Distribution: Known only from vicinity of Konahuanui and
Olympus, southeastern Oahu.

Specimens examined (all from Isl. Oahu): Charles N. Forbes,
Konahuanui, Nov. 4, 1908 (topotype, Bish., my photograph no.
4,349) ; Forbes (with Mrs. G. E. Kelly] 2,390-0, Castle Trail between
Konahuanui and Olympus, Aug. 1, 1916 (Bish.); D. Wesley Garber
245, Konahuanui-Olympus Trail, Feb. 15, 1920 (Bish.); U.S. Ex-
plor. Exped. under Capt. Wilkes, alt. 2,000 ft., mountains behind
Honolulu (type, U.S., my photograph no. 4,295: isotype, Gray, my
photograph no. 4,294).

The above description has been made to follow the original one
by Gray very closely, with a few additions made directly from the
type and isotype, both now before me. Hillebrand (Fl. Haw. Isls.
153. 1888) gave a treatment for this species (under the name Tripla-
sandra oahuensis) and cited "the slopes of Waiolani and Konahuanui,"
Isl. Oahu, for its habitat. A study of his specimens available to
me, however, shows that his own specimens were of a tree varietally
different, treated below as var. eradiata.

Gray's immature material had the stigmas as yet entire, and he
regarded the flowers as sterile because of the diminutive ovary.
However, the ovary was 4- or 5-locular and, from our knowledge
today of the genus we know that the "entire" stigma doubtless
would have divided presently into several distinct stigmas as the
flowers developed. 1 Gray's type is all too scanty but is sufficient
to show that a peduncle about 9.5 cm. long bears rays and that
these bear in turn pedicels. (How many peduncles were present in
an umbel or whether the umbel was sessile or mounted on a rhachis
is not evident from the type collection.)

The material confused by Hillebrand with this species differed
at once in having pedicels placed directly on the peduncles, not on
the rays. This and other differences are noted in the description
for var. eradiata.

The specimen collected by Forbes at Konahuanui has leaflets
somewhat more coriaceous and depressed-venulose. The two stout

1 Commonly the number of stigmas on a mature drupe in Tetraplasandra
equals the number of pyrenae or cells, but at least in var. eradiata of this species
the stigmas may be fewer.

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 93

peduncles are umbellately grouped, 7.5-8.5 cm. long, and one has
two of its fifteen 1.5-2.4 cm. long rays scattered to 3.5 cm. lower
than the terminal umbellet; its pedicels are but 3-6 mm. long.

Tetraplasandra oahuensis var. /3. longipes Sherff, Bot. Leafl.
no. 8: 4. 1953; Triplasandra oahuensis var. /3. Hillebr. Fl. Haw.
Isls. 152. 1888.

Terminal and lowermost leaflets rhomboid-obovate, blade
5.5-6 cm. long and 4 cm. wide; petiolules 7-20 mm. long.
Peduncles umbellately 2-4-congregated, 16-18 cm. long, moder-
ately scurfy, 22-flowered (rays lacking); pedicels very slender,
sparsely scurfy, 1.2-2 cm. long; corollas not yet expanded, under
5 mm. tall; calyx glabrate to more or less tomentulose-scurfy or
-mealy, obconic, about 6 mm. tall; stamens at least 10-15; im-
mature stigmas 4 or 5. Drupes unknown in mature state; immature
ones cylindric, black (at least when dry), calyx about 12 mm. tall
and about 5 mm. thick, limb brown-margined, spreading and
conspicuous, the small and conical stylopodium under 1.5 mm. tall,
stigmas commonly 2 at times 3.

Type: Joseph F. Rock 4,831, Niu Valley, Island of Oahu,
Aug. 22, 1909 (Bish.).

Distribution: Known only from type locality in southeastern-
most Oahu and from upper part of Kipapa Gulch far to the north-
west.

Specimens examined : Edward Yataro Hosaka 956 p. p., a 20 ft. tree,
wooded slope, alt. 1,600 ft., south ridge, Kipapa Gulch, Apr. 2,
1933 (Bish.); Joseph F. Rock, Niu Valley, 1911 (Am.); Rock 4,831
(type, Bish., my photograph no. 4,273).

Hillebrand's var. /3. was collected by him at Niu and was this
variety.

Tetraplasandra oahuensis var. 7. eradiata Sherff, Bot. Leafl.
no. 8: 5. 1953.

Small tree, leaves about 3 dm. long, leaflets 7-13, coriaceous,
glabrous; blade ovate to broad-oblong, 6-8.5 cm. long and 4-6.5
cm. wide, at apex obtuse to subacute or more rarely subacuminate,
at base oblique and abruptly wide-cuneate to subtruncate; petiolules
3-10 mm. long. Peduncles 3-5, at times arising from a rhachis
under 1.3 cm. long, themselves 5-8.5 cm. long, at their tip umbel-
lulately about 16-20-pedicellate, pedicels finally 13-15 mm. long.
Calyx cylindric-prismatic, 4-6 mm. tall. Petals 5 or 6, about 6 mm.

94 FIELDIANA: BOTANY, VOLUME 29

long. Stamens 10-15, a half shorter that the petals, anthers re-
curved. 1 Ovary 5- or 6-loculate. Drupes prismatic-ovoid to shortly
prismatic-cylindric, 5- or 6-angled, the calyx 7.5-8 mm. tall and
about 5-5.5 mm. thick; stylopodium 5- or 6-rayed, depressed-conic,
about 2 mm. tall (including in the same umbellet the now 2 or 3
now 5 or 6 stigmas).

Type: William HiUebrand, Nuuanu, Island of Oahu, 1867

(Gray).

Distribution: Southeastern Oahu.

Specimens examined (Isl. Oahu) : HiUebrand, Konahuanui (Bish.,
a liberal fruiting fragment ex herb. Hillebrandii olim in Herbaria
Berolinensi} ; HiUebrand, Nuuanu, 1867 (type, Gray, my photo-
graph no. 4,2236: isotype, Kew, my photograph no. 4,274&).

The two HiUebrand sheets each bear two specimens and the
label in each case gives the name Triplasandra oahuensis Seem.,
also both Nuuanu and Mt. Kaala for the habitat. One specimen
on each sheet is in the flowering state and is seen to belong to the
type collection of Hillebrand's T. kaalae, an inhabitant of Mt. Kaala.
The other specimen on each sheet has mature fruit and is seen to
match very closely Hillebrand's description (Fl. Haw. Isls. 153.
1888) of Triplasandra oahuensis Seem., our Tetraplasandra oahuensis. 2
It differs, however, from Gray's type material in such important
respects that it may well be separated varietally. These important
respects are seen quickly on reference to the varietal key given above.

Tetraplasandra oahuensis var. d. pseudolongipes Deg. &
Sherff ex Sherff, Bot. Leafl. no. 8: 4. 1953.

Leaves (at least in dry state) pallid, 4.5 dm. long (including
petiole, this 13 cm. long), ll-foliolate; blade of leaflets under
1 dm. long and under 4.5 cm. wide, more often oval-oblong to
oblong or subobovate, at apex more often weakly acuminate, at base
cuneately contracted; petiolules slender, 7-10 mm. long. Peduncle

1 In the absence of flowers on the type and isotype specimens examined, I
have relied upon Hillebrand's text for the petals and stamens.

2 Since HiUebrand knew his T. kaalae only from Mt. Kaala, it is of course
obvious that the specimens in fruit were the ones that came from Nuuanu. In
his text, HiUebrand cited the "slopes of Waiolani and Konahuanui." As Kona-
huanui slopes down on its northwest side into Nuuanu, we may well suppose
that by naming Konahuanui specifically, HiUebrand was merely revealing more
explicitly the exact habitat of the specimens previously attributed in a loose way
to "Nuuanu" (in reality a long valley extending upwards of 8 km. to the south-
west).

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 95

(a single one seen) 1.6 dm. long, smooth (but one scar present),
at its tip 10-radiate; rays curved, suberect, smooth, about 3-3.7
cm. long, at tip umbellulately about 5-7-pedicellate; pedicels sharply
angled, smooth to scurfy, 6-9 mm. long. Flowers not seen. Drupes
in submature state about 11 mm. tall including stylopodium (this
about 1.5 mm. tall); stigmas now 2 now 3.

Type: Otto Degener 2,968, Konahuanui, Island of Oahu, Dec. 28,
1924 (N.Y.).

Distribution: Southeastern Oahu.

Specimens examined (Isl. Oahu): Degener 2,968 (type, N.Y.,
my photograph no. 4,347); C. N. Forbes (with Dean Lake) 1,973-0,
Waimano Ridge, Oct. 27-30, 1914 (Bish., leaflets 7 or 9, more
coriaceous and more venulose).

In the absence of flowers, the fruits, which are essentially
typical for T. oahuensis var. oahuensis except with fewer stigmas,
have been relied upon for placing this variety in this species. Closely
related to var. pseudolongipes apparently is Degener 17,976, a 20
ft. tree, Middle Halawa Ridge, Oahu, May 26, 1935 (N.Y.). The
leaves are smaller; leaflets 5 or 7, mostly broader, also thicker and
more venulose; peduncles 5, rays more slender at least at an thesis;
stamens 15. Mature drupes are much to be desired.

Tetraplasandra oahuensis var. e. Fauriei Sherff, Bot. Leafl.
no. 8: 3. 1953.

Leaflets now dull now more or less coriaceous and glossy and
on upper surface also depressed-venulose, median ones narrowly
oblong, their blade more often 8-9.5 cm. long and 2.5-3.3 cm.
wide; lowermost elliptic-oblong to scarcely oval, blade 5-8 cm. long
and under 3.3 cm. wide; three terminal ones narrowly elliptic-oblong
to oblanceolate, apically at times truncate-rounded or even retuse.
Peduncle (a single one seen) 12 cm. long, more or less scurfy
toward base, elsewhere glabrous, at tip about 8-rayed; rays slender,
arcuate, suberect, 3.5-4.5 cm. long, at apex umbellulately 5-8-pedi-
cellate; pedicels slender, glabrous, 6-8 mm. long. Drupes immature,
their calyx cylindric to cylindric-obconic, about 9 mm. tall; stylo-
podium under 2 mm. tall including stigmas (these 2 or 3 even 4 or 5
but as yet not discrete).

Type: Abbe Urbain Faurie 270 pro parte, Konahuanui, Island of
Oahu, October, 1909 (Par.).

Distribution: Known only from type locality in southeastern
Oahu.

96 FIELDIANA: BOTANY, VOLUME 29

Specimens examined: Faurie 270 p. p. (type, Par., my photograph
no. 4,346: isotype, Arn., my photograph no. 4,353).

Tetraplasandra oahuensis var. f. pseudorhachis Sherff,
Bot. Leafl. no. 8: 3. 1953.

Leaflets 5 or 7, lateral ones subsessile with petiolules only 1-4
mm. long; petiole slender, up to 1 dm. long. Peduncle solitary,
robust, 4.5 cm. long and 5 mm. thick, more or less marked with
transverse pocks, at tip 12-rayed; some rays conspicuously elon-
gate (up to 8 cm. long), all more or less pocked-scurfy, 15-
flowered; pedicels slender, 8-11 mm. long; stamens 10-16; drupes
ovoid-prismatic, commonly 3- or 4-angled, calyx about 11 mm. tall,
stylopodium hardly 1 mm. tall, stigmas 3 or 4 or more rarely 5.

Type: G. W. Russ, Waimano Gulch, Island of Oahu, August,
1929 (Bish.).

Distribution: Known only from type locality in subcentral Oahu.

Specimens examined: Russ, Waimano Gulch, August, 1929
(type, Bish., my photograph no. 4,350).

The type had been determined as T. meiandra, but the florets
have 10-16 stamens as in T, oahuensis and the ripe drupes too are more
typical for the latter species. The lone stout, short peduncle might
be mistaken for the rhachis found in some Tetraplasandra taxa,
whence the epithet pseudorhachis. However, it bears a multiplicity
of secondary branches ( a rhachis so-called bearing seldom more than
5 or 6) and these tend to disjoint from the apex as rays often do,
leaving conspicuous circular pocks 5 mm. across.

Tetraplasandra oahuensis var. 77. venulosior Sherff, Bot.
Leafl. no. 8: 2. 1953.

Leaflets 5-7, lowermost ones ovate or oval-oblong, median ones
ovate-oblong or oblong, terminal ones obovate or widely oblan-
ceolate, all rounded to obtuse at apex or rarely abruptly subacute,
at base oblique and cuneate to rounded, coriaceous, on upper
surface more often subglossy and conspicuously depressed-venulose,
lateral petiolules 3-8 mm. long; petiole 6.5-8 cm. long. Peduncle
(a single one seen) slender, about 9 cm. long, smoothish above, at
tip umbellulately and densely 30-pedicellate; pedicels compressed,
smooth, 9-14 mm. long. Florets immature, stamens 10-16. Drupes
unknown.

Type: Edward P. Hume 165, alt. 450 m., steep hillside, north
ridge, Kaaawa, Island of Oahu, Apr. 12, 1931 (Bish.).

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 97

Distribution: Northeastern Oahu.

Specimens examined : Otto Degener, Pig-God Trail, rainy summit,
Punaluu, Isl. Oahu, May 31, 1931 (N.Y., my photograph no. 4,352);
Degener 18,015 pro parte, Pig-God Trail, windy, forested summit,
Punaluu, Jan. 17, 1932 (Mo.; N. Y.) ; Degener 18,242, same place and
date (Mo.; N.Y., my photograph no. 4,292); Hume 165 (type, Bish.,
my photograph no. 4,351).

Tetraplasandra oahuensis var. 6. hailiensis Deg. & Sherff
ex Sherff, Bot. Leafl. no. 8: 6. 1953; Tetraplasandra meiandra var.
hailiensis Deg. & Sherff ex Sherff, Bot. Leafl. no. 7: 10. 1952.

Leaves up to 4 dm. long including petiole (this 5.5-10 cm.
long), leaflets 7-9, petiolules 5-15 mm. long; blades of leaflets
oblong or more often oval-oblong or ovate, apically obtuse to
subacute, basally oblique and commonly truncate to broadly rounded,
coriaceous, reticulately depressed-venulose above, up to 11 cm.
long and to 7.5 cm. broad. Inflorescence sessile or subsessile, thrice
umbellate; peduncles 4, about 8-10 cm. long and 3-5 mm. thick;
rays terminal or a single one placed lower down on peduncle, about
12-16, subrobust, more often 4-7 cm. long; pedicels 8-12, sub-
robust, 9-16 mm. long. Petals 7, stamens 11. Drupes globose-
prismatic, strongly 4- or at times 5-angled when dry, calyx 7-8 mm.
long, the depressed-conic stylopodium under 2 mm. tall, stigmas
4 or sometimes 5.

Type: Otto Degener 18,016, decadent forest, top of Haili Gulch,
Isl. Oahu, Nov. 28, 1931 (first sheet, N.Y.; second sheet, U.S.).

Distribution: Known only from type locality in northwestern-
most Oahu.

Specimens examined: Degener 18,016 (first type sheet, N.Y.,
my photograph no. 4,283; second type sheet, U.S., my photograph
no. 4,284: isotypes, Mo., 2 sheets; N.Y.); Degener 18,247, about
4 dying trees in dryish, decadent forest, top of Haili Gulch (Makua
vicinity), Nov. 28, 1931 (Mo., my photograph no. 4,285; N.Y.).

11. Tetraplasandra waianensis Deg. Fl. Haw. fam. 281, with
plate. 1938; Tetraplasandra waianeensis Deg. ex Skottsb. Meddel.
Goteb. Bot. Tradg. 15: 414. 1944 (sphalm).

Leaflets oblong, 3-4 cm. wide var. a. waianensis.

Leaflets broadly oval-oblong or sometimes subcordate-subrotund,
5-8.5 cm. wide var. /3. palehuana.

98 FIELDIANA: BOTANY, VOLUME 29

Glabrous tree commonly 3-4 m. tall and with few, thick branches,
or at times a much-branched shrub. Leaves 1.5-3 dm. long,
5-9-foliolate; petiole 4-8 cm. long; leaflets coriaceous, oblong, at
apex bluntly acute to shallowly cuspidate, the lateral obliquely
rounded at base with blades 4-9 cm. long and 3-4 cm. wide, on
petiolules 1-6 mm. long, the terminal one evenly cuneate at base
and at times more or less obovate, its petiolule jointed near blade
and commonly 2-3 cm. long. Inflorescence sessile or upon a thick
rhachis up to 6 cm. long, twice umbellate; peduncles 4-8, spreading,
6-12 cm. long, each bearing about 3 scars of caducous, triangular-
ovate, 4 mm. long bracts, 20-30-flowered ; pedicels slender, 2 cm.
long. Calyx cylindric-obconic, 6 mm. tall, with undulate border;
petals about 8, subequal, lanceolate, acute, 7 mm. long, about 2
mm. wide; stamens about 28, unequal in length, filaments 1-2 mm.
long, anthers 2-3 mm. long; ovary 4-celled, style 1 mm. long, stigmas
4. Drupe (known as yet only in submature state) narrowly urce-
olate-cylindric, truncate at apex, 10-12 mm. tall as to its calyx and
3-4 mm. thick, stylopodium subslenderly conic, 1 mm. tall in-
cluding the 4 or rarely 3 stigmas.

Type: Otto Degener 11,318, in forest, Middle Palawai Ridge,
Waianae Range, Isl. Oahu, May 12, 1936 (herb, not cited).

Distribution: At and near Palawai Gulch, Waianae Mountains,
Isl. Oahu.

Specimens examined (all from Isl. Oahu): Degener 11,318 (type
collection, Gray, my photograph no. 4,220; Mo.; N.Y., my photo-
graph no. 4,219) ; Degener 17,973, northeast slope of Puu Kumakalii,
Waianae Range, Apr. 1, 1936 (with neither flowers nor fruits;
N.Y.); Grady L. Webster 1,443, much-branched shrub 3.5 m. tall,
alt. 2,600 ft., South Palawai Gulch, Mar. 27, 1948 (Bish.).

The "T. waianeensis" of Skottsberg's text seems to have been
an unintentional misspelling. 1

1 Curiously enough, Skottsberg elsewhere (ibid. 508) intentionally altered
waianensis to waianaeensis for Bidens waianensis Sherff, remarking: "Named for
the Waianae Mts.; the specific epithet has to be waianaeensis; waianensis is
inadmissible." The International Rules, Art. 70 (Brittonia 6: 26. 1947), de-
manded, however, the retention of the original spelling of a name or epithet
"except in the case of a typographical error, or of a clearly unintentional ortho-
graphic error." My Bidens waianensis was published in 1920 (Bot. Gaz. 70:
104) with an intended omission of the final ae of Waianae in the interest of brevity.
No typographical error was involved in the resulting epithet waianensis, nor,
since the resulting epithet was intentionally created with the ae missing, can there
be said to have been a clearly unintentional orthographic error. Thus it seems
that the spelling waianensis must stand. I suspect, moreover, that Degener 's
later use of this spelling for a new Tetraplasandra came from his naturally wishing

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 99

In general habit of foliage and shape of mature drupes, T.
waianensis is all too close to T. pupukeensis of the northern part of
the Koolau Range on Oahu. In the shape of the submature drupes,
however, both T. waianensis and its var. palehuana appear specifi-
cally different from T. pupukeensis and the several varieties and
forms of that species. The submature (about half-matured) drupes
in T. waianensis and its variety are slenderly cylindric or cylindric-
subprismatic, the calyx 2.5-3 times as tall as thick. In T. pupuke-
ensis and its subordinate entities, they are urceolate to obconic, the
calyx less than 2 times as tall as thick.

Tetraplasandra waianensis var. 0. palehuana Sherff, Bot.
Leafl. no. 8: 11. 1952.

Leaf -petiole 1 dm. long; leaflets 5-11, broadly oval-oblong or
sometimes subcordate-subrotund, at apex obtuse or rounded or
rarely subacuminate, mostly 7-12.5 cm. long and 5-8.5 cm. wide.
Flowers unknown. Mature drupes ovoid-prismatic, coarsely 4- or
sometimes 5-angled, deeply and broadly grooved between the
angles, truncate at top, calyx portion 10-11 mm. tall and narrowly
undulate-bordered at top, the slenderly conic stylopodium with
its ill-defined style and usually 4 but sometimes 5 stigmas rising
2.5 mm. above calyx-limb.

Type: Charles N. Forbes 1,674-0, Palehua, Waianae Mountains,
Isl. Oahu, Apr. 1-4, 1911 (Bish.).

Distribution: At and near Palehua, Isl. Oahu.

Specimens examined (Isl. of Oahu) : Otto Degener 17,971, Middle
Palawai Ridge, Waianae Mts., May 12, 1936 (topotype, N.Y., my
photograph no. 4,297) ; Forbes 1,674-0 (type, Bish., my photograph
no. 4,296).

12. Tetraplasandra kaalae (Hillebr.) Harms in Engl. &
Prantl, Pflanzenfam. Ill, 8: 30. 1898; Triplasandra kaalae Hillebr.
Fl. Haw. Isls. 154. 1888.

to use a Latin adjective already in use for Waianae. I have therefore followed
his spelling for the epithet in the genus Tetraplasandra. There seems an abundance
of precedents for this course. To take but one example: Pyrus ioensis (Wood)
Bailey is the official name for a North American crab-apple and is spelled as here
given in all standard works (e.g., Gray's Man. Bot. edit. 8: 759. 1950). The
epithet ioensis refers to Iowa, but might equally well have referred to many such
geographic names as loka, lola, and lona, to be found in a standard atlas. By
contrast, the epithet waianensis could well apply to no other place in the Ha-
waiian Islands than Waianae except Waianu and Waianui, both localities so
inconsequential that they are omitted from the official United States topographic
maps (Maui, ed. of 1933, and Oahu, eds. of 1917 and 1930).

100 FIELDIANA: BOTANY, VOLUME 29

Umbels thrice compound, umbellets 10-15-flowered, drupes globose-
ovoid, their calyx as broad as tall; leaves 2-3 dm. long, leaflets
7-11, petiolules 12-25 mm. long var. a. kaalae.

Umbels now 3 times now 4 times compound, umbellets 3-5-flowered,
drupes ovoid, their calyx about 8 mm. tall and 6 mm. thick;
leaves 2 dm. long, leaflets 5, lateral petiolules only 2-7 mm.
long var. j3. multiplex.

A small tree up to 9 m. tall, glabrous, ultimate branchlets
slightly scurfy. Leaves about 2-3 dm. long; petiole 4-12 cm. long,
clasping at base. Leaflets 7-11; blade ovate or ovate-oblong, at
apex subacute or obtusely acuminate to rounded or even subretuse,
at base broadly cuneate to rounded and (except for terminal leaflet)
oblique, 7-12 cm. long and 3.5-8 cm. wide, coriaceous, glabrous
on both surfaces, moderately or inconspicuously reticulate- venulose,
drying a dark- or light-green and dull or in places glossy; petiolule
12-25 mm. long. Inflorescence thrice compound; 3-5 peduncles
from a short (under 1 cm.), common rhachis, each 4-7 cm. long, com-
pressed-angular and with 3 bract-scars, bearing at top a radiate
umbel; rays 10-15, angular, curved or straightish, 2.5-4.2 cm. or
in some topotype or near-topotype material (e.g., Degener et al.
21,018 and Selling 3,338) 4-8 cm. long, with usually 2-4 bract-scars,
10-15-pedicellate; pedicels 5-13 mm. long, in a terminal umbellet.
Calyx narrowly to broadly obconic, glabrous, 2-3 mm. tall; petals
6, finally expanded, 6-8 mm. long, stamens 12-18; ovary 4- or
rarely 3-celled. Drupe globose-ovoid, when dry strongly 4- or
rarely 3- or 5-angled, broadly and deeply grooved between the
angles, its calyx 7-8 mm. tall and narrowly undulate-bordered, its
top convex and abruptly constricted to a very short, slender stylo-
podium, this holding the 3-5 (commonly 4) stigmas 2.5-3 mm.
above calyx-limb.

Type: William Hillebrand, alt. 4,000 ft., on top of Mt. Kaala,
northwestern Oahu (formerly at Berl.).

Distribution: On and near Mt. Kaala, northwestern Oahu.

Specimens examined (Isl. Oahu) : Otto Degener 18,014, forest, up
ridge on right side of head of Makua Valley, June 26, 1932 (N. Y.) ;
Degener 18,245, ridge at head of Makua Valley on south side, June
26, 1932 (N.Y.); Degener 18,246, only one dying tree above forest,
southeast side of Makua Valley near its head, Jan. 3, 1932 (N. Y.) ;
Degener 21,016, second ridge east of Dupont Trail, north of Kaala,
July 24, 1949 (Bish.; Chi.; Del.; Gray; Kew; N.Y.); Degener &
Miss Amy Greenwell 21,015, forested slope at 1,800 ft. alt., Dupont

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 101

Trail, north slope of Kaala, Oct. 23, 1949 (Berl.; Bish.; Chi.; Del.;
Gray; Kew; Mo.; N.Y.; Par.; Phila.; U.S.); Degener, Greenwell, &
Hatheway 20,682, single tree on forested western slope, alt. 1,400
ft., east branch of East Makaleha Valley, July 4, 1950 (Bish.; Chi.;
N.Y.; U.S.); Degener, Murashige, & Kerr 21,018, tree 25 ft. tall,
dryish, forested slope, alt. 2,500 ft., Mt. Kaala, Oahu, July 10,
1949 (Arn.; Bish.; Calif. Acad.; Chi., 2 sheets; Del.; Gray; Kew;
Mo.; N.Y.; Par.; Phila.; Univ. Calif.; U.S.); Degener, Ordonez, &
Selling 12,236, single tree in decadent forest, southeast of Kolekole
Pass, Sept. 3, 1938 (N.Y.); William Hillebrand, Kaala (Bish.,
fragments ex type coll. in herb. Hillebr. olim in Herb. Berol., my
photograph no. 4,194); Hillebrand, Mt. Kaala, 1867 (topotypes,
perhaps even isotypes, Gray, my photograph no. 4,223a; Kew, my
photograph no. 4,274a) ; Olof H. Selling 3,338, near Kolekole Pass,
July 3, 1938 (Bish.).

Hillebrand apparently had not seen mature drupes. Degener,
Ordonez, & Selling no. 12,236, from near Kolekole Pass, matches type
material very closely but excels in having submature drupes. Sell-
ing 3,338 from the same locality has mature drupes and longer rays,
four of one umbel measuring 7-8 cm. long. Skottsberg, who de-
termined Setting 3,338 as T. kaalae (Meddel. Goteb. Bot. Tradg.
15: 413. 1944) expressed uncertainty as to his determination, since
the Selling plant possessed no flowers and for T. kaalae no drupes
had been described. However, until such time as more complete
suites can be obtained with both flowers and fruits, we may derive
some assurance from what is known about T. kaalae var. multiplex.
In that variety, the type sheet of which bears both flowers and
mature fruits, the flowers are essentially as in Hillebrand's flowering
material of T. kaalae and the fruits are very similar to those of
Selling's fruiting material of T. kaalae, though a trifle smaller and
more nearly ovoid (i.e., less globose-ovoid) .

Hillebrand (op. cit. 151) set his T. kaalae in that portion of his
key having the "drupe cylindrical, truncate," although his failure
to describe a drupe in his text description makes it seem fairly
certain that he had not seen one for T. kaalae and was surmising
its shape from a glance at the floral calyx. For T. kaalae none but
globose-ovoid, convexly topped drupes are known.

Tetraplasandra kaalae var. /3. multiplex Sherff, Bot. Lean 1 ,
no. 8: 11. 1953.

Leaves 2 dm. long, 5-foliolate; leaflets moderately smaller,
lateral petiolules only 2-7 mm. long; inflorescence now 3 times now

102 FIELDIANA: BOTANY, VOLUME 29

4 times umbellate, peduncles 3-7 cm. long; rays now few (8)
now more numerous (17), only 2-4.5 cm. long, at apex sometimes
branched umbellately into 5-7 subordinate rays (subrays), these
sharply angulate, more or less arcuate, 1.5 cm. long; pedicels
3-5 in a terminal umbellet, slender, about 3-4 mm. long. Drupes
ovoid, the calyx portion about 8 mm. tall and 6 mm. thick, the
top more pyramidal.

Type: G. W. Russ, Makua, northwestern Oahu, January, 1930
(Bish.).

Distribution: Known only from type locality in northwestern
Oahu.

Specimens examined: Russ, Makua, Oahu, January, 1930 (type,
Bish., my photograph no. 4,193).

The type had been determined at the Bishop Museum as T.
kaalae, but the differences are so numerous as to justify at least
a distinct varietal status. The type sheet bears a flowering spray
having two 5-foliolate leaves and a sessile, thrice compound umbel
with but two peduncles. Separately mounted are two more in-
florescences. These have a long stalk, one 4.5 cm. and one 7 cm.
long. By comparisons within the genus, these stalks appear better
designated as peduncles than as rhachises. It is assumed that two
or more are umbellately clustered; this being the case, the whole
inflorescences (that is, the two separately mounted ones) are 4 times
umbellate, with a set of subrays between the rays proper and the
pedicels.

13. Tetraplasandra bisattenuata Sherff, Bot. Leafl. no. 6:
26. 1952.

Leaves 5- or 7-foliolate, 2-2.5 dm. long; petiole 6-7 cm. long;
leaflets fusiform in outline, their blade attenuate at both ends other-
wise oblong, only 6-9 cm. long and 2.5-4.5 cm. wide; petiolules
rather long (up to 3.5 cm.). Inflorescence similar to that of Tet-
raplasandra waialealae; peduncles (a single one seen) 8.5 cm. long;
secondary branches 3.5-4 cm. (not "mm." as erroneously printed
in original description) long, about 4- or 5-flowered; pedicels about
1.5-1.8 cm. long; calyx (more or less cylindrically) obconic at
anthesis, about 7-8 mm. tall and 4-5 mm. thick, presently 4- or
5-angled; corolla hemispheric-conic when shed, about 5.5 mm. tall
and slightly broader. Petals about 6, stamens about 12-19.

Type: Rev. John M. Lydgate, 4-5 ft. tall, Hii Mountains, south-
eastern Kauai (Bish.).

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 103

Distribution: Known only from type locality in southeastern
Kauai.

Specimens examined: Lydgate, 4-5 ft. tall, Hii Mts., Isl. Kauai
(type, Bish., my photograph no. 4,195).

The type had been labeled Tetraplasandra waialealae Rock, but
from that species it differs in its small, long-petioluled leaflets, which
are gently attenuate or acuminate at both ends, also in the small
calyces and doubtless small fruits.

14. Tetraplasandra meiandra (Hillebr.) Harms in Engler &
Prantl, Pflanzenfam. Ill, 8: 30. 1898; Heptapleurum (?) waimeae
Wawra, Flora 56: 159. 1873; Triplasandra meiandra and var. (?)
Hillebr. Fl. Haw. Isls. 152. 1888; Triplasandra waimeae (Wawra)
Heller, Minn. Bot. Studs. 1: 871. 1897.

For key to varieties, see general key, p. 59.

A small, glabrous tree up to 7 m. tall, branching from near
base. Leaves 2-4.5 dm. long, petiole 4-9 cm. long and dilated-
clasping at base; leaflets 7-13, blades 6-11 cm. long and 2-5 cm.
wide, variously oblong or ovate or (especially for terminal ones)
oblanceolate to obovate, at apex obtuse to subacute or acuminate, at
base cuneately contracted or sometimes rounded, coriaceous and often
glossy when dried, more or less reticulate-venulose with venation
depressed above and salient below, lateral petiolules now (for type
material) 3 mm. long now (for most material) 5-15 mm. long,
terminal petiolule 1-3 cm. long and jointed near blade. Inflo-
rescence thrice umbellate, sessile or on a short, stout rhachis; ped-
uncles 3-5, often 1-2.5 dm. long, nodulose with 3-5 bract-scars,
bearing at top an umbel of 12-21 rays (or 1-3 of these irregularly
placed lower down on peduncles) ; rays angulate, straight or curved,
2-5 cm. long, mostly enodulate, with an umbellet of 8-12 florets
at top, their pedicels slender to somewhat thickish, 6-14 mm. long.
Bracts broadly ovate, 4-8 mm. long, caducous long before anthesis.
Calyx obconic-cylindrical, with a short, repand-denticulate border.
Petals 5-8, triangular-lanceolate. Stamens as many as petals and
shorter. Ovary 2- rarely 3- celled. Drupe urceolate-cylindrical,
compressed, calyx 10-11 mm. tall and 5-6 mm. thick, stigmas 2 or
rarely 3, sessile on the broad, conical, exserted vertex, this about
3 mm. tall.

Type: Heinrich Wawra 1,638, western ridge of Nuuanu, south-
eastern Oahu (Mus. V.). 1

1 Wawra (loc. cit.) cited Waiolani, Isl. Oahu, as the type locality. For the
type locality I have followed, however, Hillebrand, who says: "Wawra's specific

104 FIELDIANA: BOTANY, VOLUME 29

Distribution: Island of Oahu, especially in the Koolau Range.

Specimens examined (all from Isl. Oahu) : Richard S. Cowan 557,
fruits red-tipped, alt. 2,000 ft., Halawa Valley ridge trail, Koolau
Range, Mar. 2, 1947 (Bish.); Otto Degener 17,972, Waimano, June
9, 1935 (Mo.; N.Y.) ; Degener 17,978, forest, C.C.C.Trail, Aiea, Feb.
16, 1936 (N.Y.) ; Degener, Salucop, & Arlantico 11,820, open summit
rain-forest, Aiea, Dec. 6, 1937 (Del.; Gray; Mo.; N.Y.; U.S.);
Faurie 270 pro parte, Konahuanui, October, 1909 (Arn.; Bish.);
Faurie 273, alt. 800 m., Waianai, May, 1910 (Arn.); Charles N.
Forbes, Konahuanui, Nov. 4, 1908 (Bish.); Forbes, Waiolani Ridge,
Sept. 17, 1908 (Bish.) ; Forbes 1,034-0, Konahuanui, Jan. 16, 1909
(Bish.); William Hillebrand, Nuuanu (fragment of practically type
material see footnote at Bish., my photograph no. 4,253);
Vaughan MacCaughey, gulch slope, "table-land," Kaena, Mar. 27,
1915 (Bish.) ; L. H. MacDaniels 62, small tree 3 m. tall, alt. 700 m.,
Waipio-Waiawa Ridge, Oct. 6, 1926 (Bish.); Mann & Brigham,
Konahuanui (Bish.); Joseph F. Rock, Punaluu Mountains, Aug. 23,
1908 (Bish.); Rock 8,722, Konahuanui, Jan. 8, 1910 (Bish., my
photograph no. 4,254); Rock & Shaw, Olympus, October, 1912
(Bish.) ; H. St. John 20,215, tree 7 m. tall, trunk diam. 1 dm., moist
woods, alt. 1,400 ft., Waimano Trail, Waimano, Mar. 23, 1941
(Bish.); D. L. Topping 2,968, Konahuanui, December, 1926 (Univ.
Calif.).

This species is remarkable within the genus, indeed within the
Hawaiian Araliaceae and perhaps as much so throughout the Hawai-
ian phanerogams, for the extreme amount of varietal diversification
displayed. Hillebrand regarded it as a "collective species," and
his view has been accepted here. Unfortunately, however, the
materials for many of the recognizable varieties are so scanty as to
make futile any attempt at phylogenetic alignments of the numerous
varieties with reference to each other. Accordingly they are listed
here in purely an alphabetical sequence.

In addition to the above cited specimens is one (R. S. Cowan
980, tree 7 m. tall, leaves thick, rigid, dark green above, Halawa
Ridge Trail, Isl. Oahu, Apr. 25, 1948 Bish.) with slender subrays
between rays and pedicels, as in T. kaalae var. mulitiplex. Florets
are lacking, however, and the several drupes are too immature to
place the specimens certainly with the T. meiandra complex.

name is wrong; his specimens were collected during a joint excursion with the
author upon the western ridge of Nuuanu, Oahu." Just why Wawra should have
used the specific epithet waimeae, pertaining to Waimea of far-northern Oahu,
for a species that he himself attributed to Waiolani of southeastern Oahu, I do
not know.

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 105

Tetraplasandra meiandra var. /3. bisobtusa Sherff, Bot.
Leafl. no. 6: 36. 1952; Triplasandra meiandra var. 7. Hillebr. Fl.
Haw. Isls. 152. 1888.

Tree 3.5-9 m. tall. Leaflets 11, their blades 7.5-9 cm. long
and 3-3.7 (-4.5) cm. wide, oblong, obtuse at both ends or narrowed
at base, thick-chartaceous, the lateral ones on petiolules 3 mm.
long. Inflorescence compound-umbellate; peduncles 5-11 cm. long,
15-25-flowered, pedicels 10-14 mm. long. Petals and stamens 5-8;
stigmas 2 or 3, sessile.

Type: William Hillebrand, Kawaihae iuka (Kawaihae-Uka),
North Kohala, Isl. Hawaii (formerly in herb. Hillebr., Berl.).

Distribution: Known only from type locality in North Kohala,
Isl. Hawaii.

Specimens examined: Hillebrand 264, tree 30 ft. tall, Kohala
Range, Isl. Hawaii (Bish.; Kew; topotypes or perhaps isotypes, my
photograph no. 4,314).

Tetraplasandra meiandra var. 7. Bryanii Sherff, Bot. Leafl.
no. 8: 10. 1953.

Leaves 7- or 9-foliolate, petiole slender, about 8-9 cm. long;
leaflets coriaceous on upper surface (where also somewhat glossy and
conspicuously venulose), ovate or oblong (terminal one subrhom-
boid to obovate), at apex more or less acuminate, at base scarcely
or not at all oblique but widely or moderately cuneate, the blade
8.5-11 cm. long and 4-6.5 cm. wide; lateral petiolules 11-27 mm.
long. Peduncles (only single, unattached ones seen) perhaps sol-
itary, smoothish, few-nodate, 11-23 cm. long, at apex umbellulately
about 10-15-pedicelled; pedicels finally 2.2-3.2 cm. long, straight-
ish or curved, erect or spreading-ascending, moderately slender
(1-1.5 mm. thick). Florets not seen. Drupe ovoid-cylindric, com-
pressed or three-angled; its calyx slenderly urceolate, 11-14 mm.
tall and 6-8 mm. thick, the expanded limb conspicuous; stylopodium
conic, 4-5 mm. tall including the 2 or more often 3 stigmas.

Type: Edwin H. Bryan, Jr., 878, small tree 5-6 m. tall, in very
wet rain-forest near the summit-ridge, alt. 2,150 ft., Kawailoa, Isl.
Oahu, Nov. 20, 1934 (Bish.).

Distribution: Northern Oahu.

Specimens examined: Edwin H. Bryan, Jr., 878 (type, Bish.,
my photograph no. 4,357: isotype, U.S., my photograph no. 4,358).

106 FIELDIANA: BOTANY, VOLUME 29

Tetraplasandra meiandra var. 5. Degeneri Sherff, Bot. Leafl.
no. 6: 27. 1952.

Tree, leaves 4 dm. long (including petiole, this 1 dm. long);
leaflets up to 9 or 11, their slender petiolules mostly 8-10 mm. long;
blades diversely oblong oval-oblong or oblong-obovate, coriaceous,
at apex retuse or rounded, at base oblique and very broadly
cuneate to rounded, now 7 X 4.7 cm. now 9 X 6.8 cm. now 12 X
5.5 cm. Inflorescence thrice umbellate, more or less verruculose;
primary branches or peduncles 4 (apparently supported on a thick
rhachis 2 cm. long), spreading, 9-14.5 cm. long and 3-6 mm.
thick; secondary branches or rays numerous, even 18 or 21, slender,
sharply angulate, 3-5.5 cm. long and 1-2 mm. thick; tertiary branches
or pedicels more often 8-12, sharply angulate, 10-12 mm. long
shortly before anthesis. Fruits and fully developed flowers not
seen. Flowers close to anthesis small; calyx cylindric-prismatic,
6 mm. tall and about 3 mm. thick, at top very narrowly brownish-
margined; corolla as yet unopened, conic, 4 mm. tall, petals 5,
stamens 5.

Type: Otto Degener 18,021, in rain-forest, Kokee, northwestern
Kauai, June 24, 1926 (N.Y.).

Distribution: Known only from type locality in northwestern
Kauai.

Specimens examined: Degener 18,021 (type, N.Y., my photo-
graph no. 4,215: isotype, Mo., my photograph no. 4,216).

The large inflorescence on the type spray measures 2.7 dm. in
diameter. The abundance of rays or secondary branches and the
diminutive size of the florets distinguish this handsome tree at once
from the other Kauai members of the genus.

Tetraplasandra meiandra var. c. Hillebrandii Sherff, Bot.
Leafl. no. 6: 31. 1952.

Leaves 1.8-2.5 dm. long including the stoutish petiole (this
3.5-7.5 cm. long), leaflets 5-9; blades thick and coriaceous, con-
spicuously and depressedly reticulate-venulose above, broadly
oblong-oval to subrotund, at apex rounded (sometimes even emargi-
nate) to obtuse or rarely almost subacuminate, at base more or
less oblique and abruptly contracted or at times widely cuneate,
4.5-11.5 cm. long and 3.5-8.5 cm. wide; petiolules stout, lateral ones
5-16 mm. long. Inflorescence thrice or at times twice umbellate,
peduncles 1, or more often 3 or 4, frequently robust, smoothish,

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 107

angulate, 4-13 cm. long; rays 8-10(-19), sharply angulate, 6-12-
flowered, 2.5-7 cm. long, devoid of nodal scars; pedicels finally
slender, 6-12 mm. long; petals 5-7, about 4 mm. long; stamens
5-7. Drupes cylindrical, becoming prismatic on drying, 1.5 cm.
long (including the broadly conic, almost 2 mm. tall stylopodium
and 3 or 4 stigmas) and 5-7 mm. thick.

Type: William Hillebrand, Isl. Lanai, 1870 (Gray).
Distribution: Island of Lanai.

Specimens examined (all from Isl. Lanai): Charles N. Forbes
104-L, mountains near Koele, June, 1913 (Bish.); Forbes 199-L,
same place and date (Bish.; U.S.); Hillebrand, Lanai, 1870 (type,
Gray, my photograph no. 4,228 : isotype fragment ex herb. Hillebr. in
Herb. Berol., Bish.) ; George C. Munro, Lanai (Bish.) ; Munro, Poomai,
Apr. 18, 1914 (U.S.); Munro 199, Hookio Ridge, Sept. 28, 1913
(Bish.) ; Emilio Ordonez (Degener distrib. no.} 12,988, Waialala Gulch,
July 7, 1940 (Mo.; N.Y.; U.S.); Joseph F. Rock 8,015, Mahana
ridge, Aug. 3, 1910 (Bish., my photograph no. 4,230); Rock &
Hammond 8,015, alt. 2,600 ft., top of ridge leading to Haalelepakai,
July 21, 1910 (Am.; Gray, my photograph no. 4,229).

Hillebrand had determined his collection originally as represent-
ing a new species of Triplasandra, but later (Fl. Haw. Isls. 153.
1888) he treated it, along with Oahu and Molokai material, as
constituting his Triplasandra meiandra var. 8.

Tetraplasandra meiandra var. f . hiloensis Sherff, Bot. Leafl.
no. 6: 34. 1952; Triplasandra meiandra var. f. Hillebr. Fl. Haw.
Isls. 153. 1888.

Somewhat similar to var. longipedunculata, but leaflets larger,
10-15 cm. long. Inflorescence either twice or thrice umbellate,
peduncles 10-17.5 cm. long, bearing at their apices an umbel of
15-20 pedicels or of rays each of which in turn bears an umbellet of
pedicels; drupes ovoid- truncate, 1 cm. tall and 6-7 mm. thick, the
5, 4, or 6 stigmas on a slender stylopodium and reaching about 1
mm. above calyx-limb; pedicels finally 2-2.4 cm. long, slender,
arcuate.

Type: Rev. John M. Lydgate, woods of Hilo, Isl. Hawaii (formerly
in herb. Hillebr., Berl.).

Distribution: Hilo, northeastern Hawaii.

Specimens examined: Lydgate, woods of Hilo (Bish., one leaflet
and one mature drupe from Hillebrand's type, formerly at Berlin).

108 FIELDIANA: BOTANY, VOLUME 29

C. N. Forbes 719-H (Bish.) lacks flowers and fruits but may
belong to this variety. It was collected along a trail between
Halelouolu and Wailuku River, South Hilo, Isl. Hawaii, June 5,
1915. The single leaf examined has a petiole 1.8 dm. long, a rhachis
2.7 dm. long (measured to joint of terminal petiolule), leaflets
mostly obtuse at each end or broadly contracted below, blade
12-13 cm. long and 6-7 cm. wide.

Tetraplasandra meiandra var. 77. leptomera Sherff, Bot.
Leafl. no. 6: 30. 1952.

Tree 9 m. tall, buds resinous; leaves delicate, 2.5 dm. long;
leaflets 7-9 (-11), blades narrow-oblong to broadly oblanceolate
(for terminal leaflet narrowly obovate), at apex rounded to obtuse
or rarely subacute, more or less gradually narrowed below to a
cuneate base, 8-10 cm. long and only 2-3 (for terminal leaflet -4)
cm. wide, median nerve doubtless scarlet in fresh state; petiolules
slender, 5-10 mm. long. Inflorescence thrice umbellate; peduncles
3 or 4, slender, angulate, irregularly scurfy, 8.5-10.5 cm. long; rays
15-23, glabrate, angulate, at anthesis 4.5 cm. long and 1-1.5 mm.
thick, about 10-flowered; pedicels slender, at anthesis only 3-5 mm.
long; petals narrowly lanceolate, 3-4 mm. long, outwardly reddish
and inwardly greenish-white (according to Forbes), alternating with
the 6 stamens. Drupes unknown.

Type: Charles N. Forbes 1,930-M, tree 30 ft. tall, buds gummy,
petals reddish outside, greenish-white inside, stamens 6 and alter-
nate with petals, 3 or 4 main peduncles . . . Lauman Forest, south
slope of Haleakala, East Maui, Mar. 11, 1920 (U.S.).

Distribution: Known only from type locality in East Maui.

Specimens examined : Forbes 1,930-M (type, U.S., my photograph
no. 4,239: isotype, Bish., my photograph no. 4,240).

Tetraplasandra meiandra var. 9. longipedunculata Skottsb.
Meddel. Goteb. Bot. Tradg. 15: 412. 1944; Triplasandra meiandra
var. e. Hillebr. Fl. Haw. Isls. 153. 1888; Tetraplasandra meiandra
var. e. Rock, Indig. Trees Haw. Isls. 347. 1913.

Petiole 3.5-7 cm. long; leaflets (5-)7-9(-13); blades oblong to
oblong-lanceolate or moderately oblong-obovate, apically rounded
to subacuminate or even acuminate, basally oblique and mostly
broad-cuneate, coriaceous, 6-9(-12) cm. long and 2.5-4.8(-6.5) cm.
wide; petiolules 5-15 mm. long. Inflorescence once or twice um-
bellate; rhachis or scape rudimentary or lacking; peduncles solitary

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 109

or 2-6, elongate, 1-2.5 or even to 3.5 dm. long; pedicels 15-30,
terminal, straightish to deflexed-ascending, 11-18 mm. long. Flower
5-8-merous, calyx about 5.5 mm. tall, limb manifest, its broadly
subtriangular lobes apiculate and 1 mm. long; petals triangular,
4 mm. long, 2-3 mm. wide; stamens 5-8, filament 1.7 mm. long,
anthers 1.5 mm. long and 0.85 mm. wide; stylopodium conic,
1.5 mm. tall, stigmas 3-5 but mostly 4, or very rarely all or a few 2
(see discussion below), sessile. Drupe subovoid-prismatic to elon-
gate-urceolate-prismatic, 3-5-angled, round-grooved between angles,
calyx 9-10 mm. (in type) to 14 or even 16 mm. tall, its border
often crenulate and conspicuously expanded, stylopodium broadly
pyramidal, 2-3 mm. tall including stigmas.

Type: William Hillebrand & Rev. John M. Lydgate, Niu, south-
easternmost Oahu (formerly in herb. Hillebr. in Berl.).

Distribution: Throughout the Koolau Range, Isl. Oahu.

Specimens examined (all from Isl. Oahu) : Richard S. Cowan 30,
Waikane-Schofield Trail, Koolau Range, Sept. 22, 1946 (Bish.);
Cowan 504, lower side of trail, moist slope, Kaunala-Waimea,
Koolau Range, Feb. 9, 1947 (Bish.); Cowan 711, tree 5 m. tall,
alt. 1,500 ft., near trail, Kipapa Gulch, Oct. 5, 1947 (Bish., forma
with longer drupes and 2 stigmas, my photograph no. 4,226) ; Cowan
745, tree 6 m. tall, fruits reddish-brown, Anahulu Trail, Kawailoa,
Koolau Range, Nov. 30, 1946 (Bish.); Cowan 963, leaves thick-
coriaceous, dark-green, glossy, Halawa Ridge Trail, Koolau Range,
Apr. 25, 1948 (Bish., the dried leaflets dull, pale-green, and sub-
chartaceous) ; Otto Degener, rainy summit, Pig-God Trail, May 31,
1931 (N.Y.); Degener 17,791 p. p., shrubby rain-forest, along sum-
mit-divide toward east of Pupukea-Kahuku Trail, Dec. 6, 1931
(N.Y.); Degener 17,977, Pig-God Trail, Punaluu, July 4, 1935
(N.Y.); Degener 18,243, windy, rainy top of ridge north of
South Halawa Gulch, Apr. 17, 1932 (N.Y.); Degener 18,249 p.
p., rain-forest at summit-divide toward east of trail, Pupukea-
Kahuku, Dec. 6, 1931 (N.Y.); Degener 18,250, directly mauka of
Kawela Bay, forest, Apr. 2, 1933 (Mo.) ; Degener & Clarence Nihei
20,222, single large tree in rain-forest at 2,200 ft., Waialae Nui
Ridge (U.S.); C. N. Forbes, mountains between Punaluu and Kaipa-
pau, Nov. 14-21, 1908 (Bish.); Forbes 2,210-0, Headgate trail,
Wahiawa, Aug. 17-20, 1915 (Bish.); Forbes 2,215-0, Wahiawa-
Kahana trail, Aug. 17-20, 1915 (Bish.) ; Hillebrand & Lydgate, Niu
(Bish., leaf and fruiting peduncle from type collection in Berl., my

110 FIELDIANA: BOTANY, VOLUME 29

photograph no. 4,257); Edward Y. Hosaka 613, tree 20 ft. tall, on
denuded ridge, alt. 2,000 ft., south ridge, Kipapa Gulch, July 4,

1932 (Bish.); Hosaka 956 p. p., 20 ft. tree, wooded slope, alt. 1,600
ft., Kipapa Gulch, Apr. 2, 1933 (Bish.) ; Joseph F. Rock, main ridge,
Punaluu Mts., Nov. 14-21, 1908 (Gray); Rock 188, Oahu (Arn.);
Rock 191, Punaluu Mts., August, 1908 (Bish.); Rock 1,307, same
place, Feb. 5, 1909 (Bish., forma approaching var. 7. BryanO);
Rock 8,826, shrub 6 ft. tall, on the ridge leading to Hauula, Punaluu-
Hauula, Koolau Mts., August, 1911 (Arn.; Bish.; Gray, 2 sheets,
my photograph no. 4,256); Rock 8,832, small shrub, Punaluu,
Koolau Mts., trail to Punaluu Camp, August, 1911 (Bish.; Gray);
Rock & C. N. Forbes, top of main ridge, Punaluu Mts., Nov. 14-21,
1908 (Arn.; Gray); Harold Saint-John 10,579, shrubby tree, "stig-
mas 2, stamens 5," alt. 2,300 ft., narrow divide, Punaluu-Kaluanui,
Sept. 28, 1930 (Bish., where stigmas are 3; Par.; Univ. Calif., where
stigmas are 2-5; U.S.); Olof H. Selling 3,631, Punaluu, Koolau Mts.,
Sept. 27, 1938 (Bish. ; Goth., my photograph no. 4,180; cited by Skotts-
berg for var. longipedunculata) ; Carl Skottsberg 1,862, alt. 700-750 m.,
ridges above Kahana Bay, Sept. 17, 1926 (Bish.) ; Amy Suehiro, alt.
2,400 ft., near main divide, Waikane-Schofield Trail, Oct. 16, 1932
(Bish.); Suehiro, 10 ft. off trail, alt. 3,000 ft., Kaipapau, Oct. 15,

1933 (Bish., stigmas mostly 3, less often 2).

Skottsberg seems to have seen only the collection by Selling.
His designation of longipedunculata as a "nov. nom.," followed
immediately by "Syn. Var. e. Hillebr. p. 153," places the type,
however, as the Niu specimen originally cited by Hillebrand. The
Niu specimen was in Hillebrand's private herbarium, which later
went into the Berlin Herbarium, which was destroyed recently by
war. It appears, however, that the type collection was obtained
jointly by Hillebrand and Lydgate. Fortunately, Lydgate's own
specimen (now before me) was preserved at Bishop Museum and
remains today to typify the variety.

The type locality for this variety, Niu, and that for var. meian-
dra, Nuuanu, are both in southeasternmost Oahu and only about
10 km. apart. Typically, var. meiandra differs in its thrice com-
pound umbels, these with rays between the peduncles and pedicels,
also in its normally 2, not 3-5, stigmas. However, occasionally
material of var. longipedunculata occurs with but 2 stigmas for some
or all of the flowers and fruits (e.g., Cowan 711; St. John 10,579;
Suehiro, Kaipapau), thus displaying a partial approach toward var.
meiandra.

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 111

Tetraplasandra meiandra var. i. makalehana Sherff, Bot.
Leafl. no. 6: 36. 1952; Triplasandra meiandra var. 0. Hillebr. Fl.
Haw. Isls. 152. 1888.

Tree 4.5 m. tall, trunk 2 dm. thick; leaflets 7-9; blades
ovate to oblong, at apex obtuse to barely acuminate, at base rounded
or cuneately contracted, coriaceous to chartaceous, 10-17.5 cm.
long and 5-7.5 cm. wide; petiole 6.5-8.5 cm. long; petiolules of
lateral leaflets 4-10 mm. long. Inflorescence thrice (more rarely
unipedunculate and then twice) umbellate, peduncles 2 (at times
5 and standing on an elongate rhachis according to Hillebrand),
slender, 6-8 cm. long; rays 8-14, smooth, about 2.5-4 cm. long,
8-14-flowered; pedicels slender, 7-11 mm. long; bracts 4 mm. long.
Petals and stamens 5, stigmas now 2 now 3, stylopodium depressed-
conic and short; drupe ovoid and compressed or trigonous, about
8 mm. tall.

Type: William H. Hatheway 209, tree 15 ft. tall, trunk 8 inches
in diameter, base of fruit green, its tip scarlet, pollination by bees,
on dry, sunny slope . . . alt. 1,200 ft., east side, Makaleha Stream,
Isl. Oahu, Aug. 21, 1950 (Chi.).

Distribution: Known only from type locality in northwestern
Oahu.

Specimens examined: Hatheway 209 (type, Chi., my photograph
no. 4,275: isotype, Gray, my photograph no. 4,276); Joseph F. Rock
17,053, Makaleha Ridge, Isl. Oahu, April, 1918 (topotype, Bish.);
Rock 17,155, right ridge of Makaleha, May 4, 1918 (topotype, Bish.).

The large leaflets are especially noteworthy for this variety.

Tetraplasandra meiandra var. K. mauiensis Sherff, Bot.
Leafl. no. 6: 29. 1952.

Leaves large, 3-4 dm. long including the 5-8 cm. long petiole;
leaflets 5-9; blades variously elliptic-oblong to ovate or subcuneate-
obovate, at apex obtuse to rounded or slightly emarginate, at base
oblique and cuneately contracted, up to 13.5 cm. long and to 7.2 cm.
wide, median vein scarlet beneath; petiolules rather elongate, 1-2. 5 cm.
long. Inflorescence sessile or subsessile, thrice umbellate, more or
less scurfy; peduncles 3-6, often robust, 8-15 cm. long, subcom-
pressed; rays 10-25 (at times 1-3 arising lower down on peduncle),
sharply angulate, 5-10 cm. long, commonly 10-20-flowered ; pedicels
5-9 mm. long; petals 3.5-4 mm. long. Drupes cylindrical, about 11
mm. long including the hemispherical stylopodium (this about 2.5
mm. tall) and about 5 mm. thick; stigmas 3 or 4, small.

112 . FIELDIANA: BOTANY, VOLUME 29

Type: Joseph F. Rock 8,666, lava fields, Auahi (Auwahi), East
Maui, November, 1910 (Gray).

Distribution: Known only from East Maui.

Specimens examined (all from East Maui): Charles N. Forbes
1,713-M, large tree 35 ft. tall, trunk 1+ ft. in diameter, peduncles
5 or 6, rays 15-23 (occasionally 1-3 below), pedicels 13, leaflets
11 or 10, their midrib crimson beneath, ridge, left-hand side of Kipa-
hulu, above the waterfall, Nov. 26, 1919 (Bish.); Forbes 1,726-M,
large tree, 25-30 ft. tall, trunk diameter 3 ft., above Mokulau Land-
ing, Dec. 7, 1919 (Bish.) ; Forbes 2,051-M, south slope of Haleakala,
Mar. 20, 1920 (topotypes, Bish.; U.S.); George C. Munro 401, Auhi
(Auwahi) or Ulupalakua, Feb. 12, 1915 (Bish.); Rock 8,666 (type,
Gray, my photograph no. 4,233: isotypes, Arn., 2 sheets; Bish., my
photograph no. 4,234; Gray, my photograph no. 4,236; U.S.,
my photograph no. 4,235) ; Rock 12,506, lava fields, Auwahi, March,
1912 (Bish.).

This variety is the one illustrated by Rock (Indig. Trees Haw.
Isls. pi. 138. 1913) for an "Auahi" (Auwahi) tree of T. meiandra
var. 8., growing at an elevation of 3,000 ft., and with trunk diameter
of 2 ft. An isotype at Gray Herbarium has a glaucous hue to portions
of the leaflets' upper surface. Definitely distinctive of the variety,
however, seem to be the large leaves and leaflets (the latter with
basally more cuneately contracted blade and with their median
nerve crimson beneath when fresh and at times even when dry),
the elongate petiolules, and the more elongate rays.

Tetraplasandra meiandra var. A. molokaiensis Skottsb.
Meddel. Goteb. Bot. Tradg. 15: 412. 1944; Tetraplasandra meiandra
var. d. Hillebr. Fl. Haw. Isls. 152. 1888.

Skottsberg's varietal epithet molokaiensis technically rests upon
T. meiandra var. 6. Hillebr. (see discussion below under var. Skotts-
bergii). Hillebrand's description of the var. 8. is a generalized or
composite one drawn evidently from varietally unlike taxa native
severally to the islands of Oahu, Molokai, and Lanai. His herbarium
perished with the destruction of the great Berlin Herbarium in
World War II, hence his cited material is no longer extant. His
first locality cited was Wailupe of far southeastern Oahu, which
must be taken as the type locality. To date, however, I have found
no specimens from Wailupe or vicinity that would seem to represent
the var. 5. and so permit a fresh and exclusive description to be
drawn (as apart from his Molokai material my var. polyantha

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 113

and his Lanai material my var. Hillebrandii} . In the absence of
such a description, it is of course impossible for the present to
assign its equivalent, i.e., var. molokaiensis, to its proper place in
the analytical key to the numerous varieties of T. meiandra.

Tetraplasandra meiandra var. /z. occidentalis Skottsb.
Meddel. Goteb. Bot. Tradg. 15: 413. 1944.

Small tree. Petiole 6-7.5 cm. long. Leaflets 3-5 or more com-
monly 7; blades oval-elliptic to oval-oblong, at apex rounded to
acute, at base mostly oblique and more or less broad-cuneately
contracted, 4-8 cm. long and 2-4 cm. wide, smoothish; petiolules
5-13 mm. long. Peduncles (solitary so far as known) 3.5-9.5 cm.
long; rays 5-8, about 2.5-5 cm. long, 6-8 (-10) -flowered; pedicels
8-12 mm. long. Flower 5- or 6-merous, 12-13 mm. long. Calyx
cylindric-campanulate, 7 mm. tall, about 3 mm. thick, its limb
subentire; petals narrowly triangular, reddish-violet, 6 mm. long,
2.5-3 mm. wide; stamens 5 or 6, filament 4 mm. long, the strongly
curved anthers about 2.2 mm. long and 1.3 mm. wide; stylopodium
broadly conic, 1.5-1.7 mm. wide; stigmas 3 or 4, sessile. Drupe
unknown.

Type: Cranwell, Selling & Skottsberg 2,675 (misprinted 2,695
originally but corrected in Skottsb. loc. cit. p. 518), alt. about 1,400
m., headwaters of Honokahua Stream, near Nakalalua, West Maui,
July 26, 1938 (Goth.).

Distribution: Summit of Puu Kukui and northwestward along
headwaters of Honokahua Stream, West Maui.

Specimens examined (all from West Maui): Albert S. Hitchcock
14,824, upper forest, alt. 3,000-5,000 ft., Puu Kukui, Sept. 24-26,
1916 (U.S.); Skottsberg 781 (misprinted 785 in Skottsberg's text),
Puu Kukui, Oct. 9, 1922 (Bish.; Goth., my photograph no. 4,181);
Cranwell, Selling, & Skottsberg 2,675 (type, Goth., my photograph
no. 4,178: isotype, Bish.); Selling 3,879, between Nakalalua and
summit of Puu Kukui, July 28, 1938 (Bish.; Goth., my photograph
no. 4,179).

Tetraplasandra meiandra var. v. olowaluana Sherff, Bot.
Leafl. no. 6: 37. 1952.

As to foliage more or less similar to var. ramosior, but with
leaflets at times more acuminate or more rounded at apex. In-
florescence thrice umbellate (rhachis absent); peduncles 3, elon-
gate, more often 1-1.5 dm. long, 8-15-radiate; rays angulate, 3-6

114 FIELDIANA: BOTANY, VOLUME 29

cm. long, 8-15-flowered; pedicels slender, 5-8 mm. long. Drupes
globose-cubical or shortly (breviter) trigonous-prismatic, calyx about
6 mm. tall and about as thick, stylopodium conic and 1.6 mm.
tall, stigmas now 3 now 4.

Type: Charles N. Forbes 2,276-M, tree 30 ft. tall, one seen in
gully far up, Olowalu Valley, West Maui, May 9, 1920 (Bish.).

Distribution: Known only from type locality in West Maui.

Specimens examined (all from West Maui) : Otto Degener 2,315,
rain-forest, near summit of Mt. Eke, Aug. 29, 1927 (Bish.; Mo., 2
sheets; N.Y., 2 sheets); Charles N. Forbes 422-M, Honokahau
Drainage Basin, Sept. 25-Oct. 17, 1917 (Bish.); Forbes 2,276-M,
Olowalu Valley, May 9, 1920 (type, Bish., my photograph no.
4,278) ; Forbes 2,313-M, ridge, left-hand side, Olowalu Valley, May
10, 1920 (topotype, U.S., my photograph no. 4,279) ; Joseph F. Rock
8,207, dense forest on the Honokawai (Honokowai) ridge leading to
summit of Puu Kukui, Aug. 23, 1910 (Arn.; Gray); Rock 16,009, Mt.
Eke, September, 1918 (Bish.) ; Harold St. John 10,231, tree 25 ft. tall,
wooded ridge, alt. 4,000 ft., Haelaau, above Honokowai, Feb. 6,
1930 (Bish.; N.Y.); R. L. Wilbur & G. L. Webster 835, 1.5 m. shrub,
rain-forest at edge of bog, alt. 4,500 ft., Puu Kukui, June 23 & 26,
1948 (Bish.) ; Wilbur & Webster 863, rain-forest, alt. 3,000 ft., Puu
Kukui, June 28, 1948 (Bish.); Wilbur & Webster 890, 5 m. tree,
"stamen 5-8," rain-forest, alt. 3,000 ft., Puu Kukui, June 29, 1948
(U.S.).

Tetraplasandra meiandra var. . ovalis Sherff, Bot. Leafl.
no. 6: 31. 1952.

Leaves 3.3 dm. long, petiole 4.5-9 cm. long; leaflets 9-11,
blade commonly oblong- or elliptic-oval, at apex obtuse or rounded,
at base oblique and more or less wide-cuneately narrowed, smooth
and obscurely or moderately venulose, 6-11 cm. long and 3.2-6.2
cm. wide; petiolule slenderish, 5-8 or even -17 mm. long. In-
florescence thrice umbellate; peduncles 3, robust to subrobust,
subangulate, scurfy, 6.5-14 cm. long; rays 20-22, sharply angu-
late, finally 3-5.2 cm. long, about 9-14-flowered; pedicels slender,
finally 7-11 mm. long; petals moderately to broadly or even del-
toidly lanceolate, about 3.5-4 mm. long. Drupes (only immature
ones known) urceolate, about 7 mm. tall including the narrowly conic
stylopodium and the usually 4 stigmas, about 6 mm. thick.

Type: Joseph F. Rock 8,582, Waikamoi Trail, Olinda, September,
1910 (Gray).

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 115

Distribution: East Maui.

Specimens examined (all from East Maui): Otto Degener, Pipe-
Line Trail, Olinda, north-central East Maui, June 16, 1927 (topo-
type, N.Y., my photograph no. 4,238); Degener, same place, July
16, 1927 (N.Y.); Degener (& Wiebke) 2,317, rain-forest, ditch-trail,
Olinda, June 16, 1927 (topotypes, N.Y., 2 sheets, my photograph
no. 4,237) ; Degener 17,788 pro parte (this number used by Degener
also for another plant, referable to T. hawaiiensis var. microcarpa,
qu. vide), Pipe-Line Trail, Olinda, June 16, 1927 (Mo.; N.Y.); Rock
8,582 (type, Gray, my photograph no. 4,246: isotype, Bish., my
photograph no. 4,247); Rock 10,070, Kailua, trail to Honomanu,
October, 1910 (Bish.).

Tetraplasandra meiandra var. o. polyantha Sherff, Bot.
Leafl. no. 6: 28. 1952.

Petiole 6-8 cm. long; leaflets 5-9, blade oblanceolate or narrowly
to moderately obovate to elliptic-rectangular, at apex obtuse to
rounded or more rarely truncate or very rarely emarginate, at base
oblique and cuneate-contracted to rounded, 5.5-10.5 cm. long and
3-5.6 cm. wide, petiolule 4-10 mm. long. Inflorescence more or less
scurfy, thrice umbellate; peduncles 3, about 8-12 cm. long,
robust, compressed; rays now 12 now (for the same specimen)
24, sharply angulate and more or less compressed, 3-4.7 cm. long,
8-18-flowered, at times one or two placed lower down; pedicels
slender, at anthesis 4-7 mm. long; petals 4-5 mm. long. Drupes
unknown.

Type: Otto Degener 17,795, rain-forest, near Pepeopae Bog, Apr.
23, 1928 (N.Y.).

Distribution: Eastern Molokai.

Specimens examined (all from eastern Molokai): Otto Degener,
west of Pepeopae, Apr. 12, 1928 (topotype, N.Y.); Degener, south-
east of Pepeopae Bog, May 9, 1928 (topotype, N.Y.); Degener
17,793, rain-forest, west of Pepeopae, Apr. 12, 1928 (topotypes,
Mo.; N.Y.) Degener 17,795, (type, N.Y., my photograph no.
4,231: isotype, N.Y., my photograph no. 4,232); Degener 18,018,
rain-forest, west of Pepeopae, Apr. 12, 1928 (topotype, N.Y.);
Degener 18,019, rain-forest, vicinity of Pepeopae Bog, Apr. 23, 1928
(topotypes, Mo.; N.Y.); Abbe Urbain Faurie 271, alt. 600 m., Pukoo,
May, 1910 (Arn.; Par.); Joseph F. Rock 6,175, edge of gulch above
Kamoku, Mar. 21, 1910 (Gray; U.S.).

116 FIELDIANA: BOTANY, VOLUME 29

Hillebrand's material from Maunahui, within 5 km. of the
Pepeopae Bog, was included by him, as stated below under var.
Skottsbergii, in his Triplasandra meiandra var. 8., called by later
writers Tetraplasandra meiandra var. <5. His Maunahui specimen
in the Berlin Herbarium was destroyed in World War II, but doubt-
less was the present variety.

Tetraplasandra meiandra var. TT. polystigmata Sherff, Bot.
Leafl. no. 7: 9. 1952.

Leaves up to 3.5 dm. long, petiole 7-11.5 cm. long; petiolules
(5-) 10-20 mm. long; blades of leaflets oval-oblong to oblong or
ovate, at apex truncate or truncate-emarginate to rounded or
subacute or even subacuminate, at base oblique and truncate to
rounded, coriaceous, on upper surface reticulately depressed-venu-
lose, 6-12.5 cm. long and 4-7 cm. wide. Inflorescence sessile, thrice
umbellate, peduncles 5, 11 cm. long, slender; rays 10-20, all
terminal or 1-3 placed lower down on peduncle, 3.5 cm. long;
pedicels about 6-8, slender, finally 1-1.5 cm. long. Drupes globose-
or urceolate-ovoid, their calyx about 7-8 mm. long, their apex
exserted-conic and 2-3 mm. long including the 5 or at times 6
stigmas.

Type: Joseph F. Rock 12,508, Honomanu trail, Haleakala, East
Maui, May, 1911 (Gray).

Distribution: East Maui.

Specimens examined (East Maui) : Otto Degener 17,785, Opuola
Stream in vicinity of ditch trail, July 7, 1927 (topotype, N.Y.);
Degener 17,786, forest, Pipe-Line Trail, Olinda, July 16, 1927 (N.Y.,
my photograph no. 4,288); Joseph F. Rock 12,508 (type Gray, my
photograph no. 4,286: isotype, Bish.); Carl Skottsberg 878, Kula
Pipe Line, Haleakala, alt. about 1,200 m., Oct. 18, 1922 (Bish., my
photograph no. 4,287).

Closely allied to the var. ovalis of the same region but having
larger and mostly more veiny leaflets, commonly 5 or 6 not 4 stigmas,
6-8 rather than 9-14 florets to an umbellet, etc.

Tetraplasandra meiandra var. p. prolifica Sherff, Bot. Leafl.
no. 6: 32. 1952.

Leaves 3.5 dm. long, more often 9-13-foliolate; leaflets on a
petiolule 4-5 or more often -10 mm. long; blade oblong or oblong-
oval or at times oblanceolate-obovate, at apex obtuse or rounded
and sometime emarginate, at base oblique and rounded or wide-

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 117

cimeately narrowed, up to 12 cm. long and to 5.5 cm. wide, veinlets
scarcely reticulate or conspicuous on upper surface but without
doubt often crimson in the living state; inflorescence sessile, thrice
umbellate; peduncles 3, subrobust, smoothish, angulate, 3. 5-7 cm.
long; rays 8-15, umbellately or at times subumbellately clustered,
angulate, smooth, 3-5 cm. long, about 12-18-flowered ; pedicels
finally 8-12 mm. long; petals not seen; drupes ovoid, their calyx
about 9 mm. tall and thick, stylopodium hemispherical (about 6
mm. broad and 3-3.5 mm. tall, its apex contracted into a short and
slender style), stigmas 2.

Type: Joseph F. Rock 12,509, Puuwaawaa, Isl. Hawaii, Feb-
ruary, 1912 (Bish.).

Distribution: Known only from type locality in northwestern
Hawaii.

Specimens examined (all from Isl. Hawaii): Rock, Puuwaawaa,
August, 1917 (topotype, Bish.); Rock 3,857, same place, June 17,
1909 (topotype, Bish.); Rock 12,509 (type, Bish., my photograph
no. 4,241: isotype, Gray, my photograph no. 4,242).

The shape and dimensions of the mature fruit are described from
a Forbes specimen, a forma with the leaflets slightly narrower and
apically less rounded, also somewhat more clearly reticulate-venu-
lose and lacking the reddish hue of the type's principal veins.

Tetraplasandra meiandra var. s. prolificoides Sherff, Bot.
Leafl. no. 6: 33. 1952.

Tree 9 m. tall, very similar to the var. prolifica but with
leaflets a little smaller and proportionately wider, the veinlets
somewhat more conspicuous, petiolule a little shorter, pedicels only
4-6 mm. long, the umbellets hence more compact.

Type: Joseph F. Rock 3,855, woods back of Puuwaawaa, Isl.
Hawaii, June 17, 1909 (Gray).

Distribution: Known only from Puuwaawaa and vicinity,
northwestern Hawaii.

Specimens examined (all from Isl. Hawaii): G. 0. Fagerlund &
A. L. Mitchell 945, tree 30 ft. tall, Waiho, Puuwaawaa, Nov. 21,
1944 (Bish.); Rock 3,855 (type, Gray, my photograph no. 4,243);
Rock 3,858, Puuwaawaa, June 16, 1909 (topotype, Gray.)

Tetraplasandra meiandra var. r. ramosior Sherff, Bot. Leafl.
no. 6: 36. 1952.

118 FIELDIANA: BOTANY, VOLUME 29

Tree 6 m. tall, leaflets 9-11; blades narrowly oblong to oval
or (of terminal leaflet) obovate, coriaceous, at apex subacuminate,
at base cuneately contracted, 7-12 cm. long and 2.5-4.8 cm. wide,
on upper surface more or less depressed-reticulate-venulose; petio-
lules elongate, 1-3 cm. long, petiole 7-9 cm. long. Inflorescence
on a rhachis, this 1 dm. long; peduncles about 6-9, about 4.5-6
cm. long, 8-radiate; rays compressed or angulate, 1.5-3.5 cm.
long, 7-10-flowered; pedicels slender, 6-9 mm. long. Petals and
stamens not seen; immature drupes urceolate-cylindric, their calyx
about 6 mm. tall and 4 mm. thick, stylopodium conic and about
2 mm. tall including the commonly 5 rarely 4 stigmas.

Type: G. R. Ewart, III, & G. C. Munro 129, growing 6 m. tall, alt.
3,900 ft., Haelaau-Puu Kukui trail, West Maui, Dec. 18, 1928

(Bish.).

Distribution: West Maui.

Specimens examined (West Maui) : G. R. Ewart, III, no. 100, alt.
3,900 ft., in rain-forest, Haelaau, Dec. 19, 1928 (topotype, Bish.,
my photograph no. 4,282); Ewart & Munro 129 (type, Bish., my
photograph no. 4,281).

The presence of a rhachis supporting the inflorescence is ex-
ceptional in T. meiandra (but cf. quotation from Hillebrand under
var. makalehana).

Tetraplasandra meiandra var. u. rhynchocarpa Sherff, Bot.
Leafl. no. 6: 33. 1952.

Large tree, up to 18 m. tall. Leaves 3.5 dm. long; leaflets
9, their blades oblong or more rarely ovate or subobovate, at both
ends obtuse or at the oblique base rounded or more rarely sub-
truncate, 6-12(-15) cm. long and 3.5-5.5(-6.8) cm. wide; petiolules
slender, 3-8 mm. long. Inflorescence twice or possibly thrice com-
pound; peduncles (a single one seen) 10.5 cm. long, somewhat
scurfy, compressed, about 5 mm. thick; rays umbellate, 9+, sub-
glabrate, angulate, 5.5-6.7 cm. long and 2-3 mm. thick, 10-flowered ;
pedicels finally 8-15 mm. long, scarcely slender; petals not seen;
drupes elongately cylindric-prismatic, their calyx finally 12-14 mm.
long and 5-6 mm. thick, topped by a conic, elongate, snout-like
stylopodium, this 3-4 mm. tall including the 3 or 4 stigmas.

Type: Joseph F. Rock 12,993, Kilauea Volcano Forest Reserve,
Isl. Hawaii, August, 1917 (Bish.).

Distribution: Southeastern Hawaii.

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 119

Specimens examined (Isl. Hawaii) : Otto Degener 17,790, in jungle,
between Glenwood and 29 Miles, June 24, 1929 (N.Y.); Degener
18,252, same place and date (Mo.; N.Y.); Degener & Wiebke 2,184,
wet jungle, between Glenwood and Kilauea, July 27, 1926 (topotypes,
Mo., 2 sheets; N.Y., 3 sheets; U.S.); Rock, Kilauea Volcano, De-
cember, 1914 (topotype, Bish.); Rock, a large tree 50-60 ft. tall,
near 29 Miles Forest, August, 1917 (Bish.); Rock 12,993 (type,
Bish., my photograph no. 4,251).

Tetraplasandra meiandra var. i/. rhynchocarpoides Deg.

& Sherff ex Sherff, Bot. Leafl. no. 7: 8. 1952.

Tree 9 m. tall, very similar in its leaves to var. v. rhynchocarpa.
Inflorescence sessile, thrice umbellate; peduncles 5, more or less
robust, 6-12 cm. long; rays 22, all placed at or one or two placed
1-3 cm. below tip of peduncle, 1-1.5 mm. thick, 3-7 cm. long, at
tip 12-20-flowered; pedicels slender, finally 9-13 mm. long, straight
or at times somewhat arcuate. Submature drupes deltoid-ovate in
side view, basally truncate; calyx about 5.5 mm. tall and toward
base about 0.5 mm. thick; stylopodium conic, elongate and snout-
like, about 3.5 mm. long including the 3 or 4 stigmas.

Type: Otto Degener 21,787, tree 30 ft. tall, overtopping forest
at alt. 2,000 ft., Kulani Prison Road, Isl. Hawaii, Feb. 5, 1952
(Chi.).

Distribution: Southeastern Hawaii.

Specimens examined: Degener 21,787 (type, Chi., my photograph
no. 4,303: isotypes, Berl.; Bish.; Del.; Gray; Kew; Mo.; N.Y.;
Par.; Phila.; U.S.); Fagerlind & Skottsberg 6,298, alt. about 6,000
ft., woods near clearing (watering place), Kulani forest, Isl. Hawaii,
Feb. 8, 1948 (Stockh.).

A sister variety to var. rhynchocarpa (likewise of southeastern
Hawaii). It differs, however, in having more numerous (22, not
9), slender (1-1.5 mm., not 2-3 mm. thick) rays, slender not
thickish pedicels, these 12-20 not 10 in an umbellet, and drupes
much smaller, their calycular portion 5.5 mm. not 12-14 mm. tall.

Tetraplasandra meiandra var. <. Rockii Sherff, Bot. Leafl.
no. 6: 33. 1952.

Leaves 3.5 dm. long; leaflets 7-13, petiole 5-10 cm. long;
blades thicker and more coriaceous, oblong or at times subovate
or subobovate, at both ends obtuse or rounded, at base oblique,
commonly depressed- and reticulate-venulose above, 7-11 cm. long

120 FIELDIANA: BOTANY, VOLUME 29

and 4-6.5 cm. wide; petiolules 4-12 mm. long, now narrow now
broader. Inflorescence thrice umbellate; peduncles about 5, robust,
compressed, rugose, 5-9.5 cm. long; rays 5-16, angulate, somewhat
scurfy, more often 3-6 cm. long, 5-12-flowered; pedicels thick, 5-8
mm. long; petals not seen; drupes cylindric-ovoid, 11-12 mm. tall
(including the short, depressed-hemispherical stylopodium and the
3 or 4 stigmas) and about 7 mm. thick.

Type: Joseph F. Rock 10,006, alt. 3,000 ft., woods of Naalehu,
Kau, Jan. 9, 1912 (Gray).

Distribution: Known only from type locality in southernmost
Hawaii.

Specimens examined: Rock 10,006 (type, Gray, my photographs
no. 4,248 and 4,249: isotypes, Arn.; Bish., my photograph no.
4,250; Gray, 2 sheets).

Rock (Indig. Trees Haw. Isls. 349, pi. 139. 1913) gave a short
description and an excellent illustration of this tree, which he
referred to Tetraplasandra meiandra var. f . The type of T. meiandra
var. f . (var. f . hiloensis of this text) came, however, from the woods
of Hilo, northeastern Hawaii, and had peduncles 10-17.8 cm. long,
fruiting pedicels very slender and elongate (Hillebrand gave the
length as 10-12 lines, and the fragment of his type from Berlin at
Bishop Museum has a pedicel 22 mm. long!), and 4-6 stigmas on
a short, slender stylopodium resting upon the truncate or slightly
depressed apex of the drupe, this surrounded by the conspicuous
brownish border of the calyx-limb.

Tetraplasandra meiandra var. x- simulans Sherff, Bot. Leafl.
no. 6: 35. 1952.

Leaves 3.5 dm. long, petiole elongate (11-13.5 cm. long), leaf-
lets few (7); their blades somewhat narrowly oblong to oblong-
obovate, at apex more often subabruptly contracted and shortly
acuminate or rarely rounded or even emarginate, at base rounded or
wide-cuneately narrowed, not conspicuously reticulate-venulose,
sometimes apparently glaucous above, 10-13.5 cm. long and 4-6.4
cm. wide; petioles 6-16 mm. long. Inflorescence normally sessile and
only twice umbellate; peduncles 3 or 4, more often elongate and
even 12-18 cm. long, compressed and subangulate; rays lacking;
pedicels very numerous (up to 30), more elongate, after anthesis
22-25 mm. long and only 0.5-1 mm. thick; petals 6 or 7, 6.5
mm. long; mature drupes unknown; very immature drupes obconic-
cylindrical, calyx about 6 mm. tall and 3 mm. thick, stylopodium
conic and much compressed, 2 mm. tall including the 2 stigmas.

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 121

Type: Joseph F. Rock 8,429, Awini, Kohala, north westernmost
Hawaii, June, 1910 (Bish.).

Distribution: Northwesternmost Hawaii.

Specimens examined: Rock 8,429 (type, Bish., my photograph
no. 4,244: isotypes, Arn.; Bish., my photograph no. 4,245; Gray;

U.S.).

In its lack of rays between peduncles and pedicels, also in its
large number of pedicels, this variety simulates the otherwise quite
different T. kohalae Skottsb., the type locality of which is only a few
kilometers to the southeast of Awini. Besides these two characters,
the peduncular length, the acutish tips of most of the leaflets, the
elongate petioles, and the small number of leaflets for such large
leaves, combine to distinguish the var. simulans easily from all others
on the Island of Hawaii. Indeed, were it not for the multiplicity
of connecting intermediate or overlapping varieties known in the
great T. meiandra complex, var. simulans might seem better
segregated as a distinct species.

Tetraplasandra meiandra var. \f/. Skottsbergii Sherff, Bot.
Lean 1 , no. 6: 28. 1952; T. meiandra var. molokaiensis Skottsb. Meddel.
Goteb. Bot. Tradg. 15: 412. 1944 as to Molokai material but exclud.
syn. var. <5. Hillebr.

Petiole up to 10.5 cm. long; leaflets 7-9, their blades widely
elliptic-rectangular to subobovate, at apex broadly rounded or very
obtuse and often emarginate, at base oblique and rounded or con-
tracted, 6-10 cm. long and 3-6.5 cm. wide ("9 X 6.1, 8.5 X 5.3,
7.8 X 4.7, 6.5 X 3.7, 6 X 4.7, 6X3 cm.") ; petiolules 6-9 mm. long.
Inflorescence twice (or thrice?) umbellate; peduncles 8-19 cm. long;
rays 11-17, commonly 4-6.5 cm. long, 5-8-flowered; pedicels 7-10
mm. long. Drupes (only immature ones known) cylindrical,
9-10 mm. long including the hemispherical stylopodium, this 2.5-3
mm. tall; stigmas 3 or 4 (more rarely 5), sessile.

Type: Lucy May Cranwell 3,528, Kawela, Isl. Molokai, June
26, 1938 (Goth.).

Distribution: Eastern Molokai.

Specimens examined: Miss Cranwell 3,528 (type, Goth., my
photograph no. 4,182: isotype, Bish., my photograph no. 4,227);
C. N. Forbes 562 Mo, Wailau Valley, September, 1912 (Bish.).

Skottsberg (loc. cit.} advanced the name molokaiensis as a nomen
novum for Hillebrand's original, Triplasandra meiandra var. 5., of

122 FIELDIANA: BOTANY, VOLUME 29

the Tetraplasandra meiandra var. 8. of later authors. He then gave
a fresh description (evidently modified from his observations of the
Cranwell material in the direction of Hillebrand's really irreconcilable
description) and cited Miss Cranwell's material from Molokai as an
example. His real type basis for the name molokaiensis was, however,
the type of Hillebrand's var. 6. Reference to Hillebrand (Fl. Haw. Isls.
153. 1888) shows Wailupe, Isl. Oahu, as the first cited locality.
Two additional localities were listed, namely (1) Maunahui on
Molokai and (2) Isl. Lanai; these are referred in this text to var. o.
polyantha and var. e. Hillebrandii respectively. The identity of
the Wailupe and other Oahu specimens constituting a satisfactory
basis for Hillebrand's var. 6. in its properly restricted sense is
discussed elsewhere (vide p. 112). We may observe for the present,
however, that henceforth the var. 5., typically of Oahu, must bear
under our International Rules of Nomenclature the inapt name
molokaiensis; also, that the Cranwell specimen from Molokai,
the specimen that led Skottsberg originally to create the epithet
molokaiensis, must be separated varietally as another variety, namely
var. Skottsbergii.

Tetraplasandra meiandra var. co. tenuistylis Sherff, Bot.
Lean. no. 6: 35. 1952.

Leaves 4 dm. long, ll-foliolate; blades of leaflets commonly
oblong, apically obtuse to rounded, at base wide-cuneately contracted
and sometimes oblique, somewhat noticeably reticulate-venulose,
8-14 cm. long and 4-6.5 cm. wide; petiolules slender or fairly broad,
8-18 mm. long. Peduncles (perhaps solitary) 5.5-10 cm. long, more
or less robust (4-11 mm. thick lower down), becoming smooth; rays
10-15 angulate, smoothish, 3.5-5.5 cm. long, about 10-15 flowered;
pedicels subrobust, finally 8-12 mm. long; petals not seen; drupes
urceolate-ovoid, truncate at apex, about 12-12.5 mm. tall including
the slender stylopodium (style) and the 2 or 3 stigmas, 7 mm.
thick.

Type: Joseph F. Rock 10,362, Volcano Kilauea, Isl. Hawaii,
December, 1914 (Bish.).

Distribution: Known only from Volcano Kilauea, southeastern
Hawaii.

Specimens examined: Rock 10,362 (type, Bish., my photograph
no. 4,252: isotype, Gray).

The leaflets have a brownish venation in the dry state, but the
original color seems to have been reddish or crimson.

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 123

Tetraplasandra meiandra var. a/, tenuistyloides Deg. &
Sherff ex Sherff, Bot. Leafl. no. 8: 10. 1953.

Tree 0.5 m. tall, sparingly branched. Leaves up to 4.5 dm.
long (including petiole, this up to 13 cm. long), up to 9-foliolate;
leaflets with blade oblong or subobovate, apically obtuse to rounded
or even emarginate, basally oblique and obtuse to subtruncate, up
to 13 cm. long and to 7 cm. wide, coriaceous, more or less reticulate-
venulose above; lateral petiolules 5-20 mm. long. Inflorescence
(a single one seen) thrice-umbellate, sessile, smoothish; peduncles
4, about 6-7.5 cm. long and 5 mm. thick, scarcely nodulate; rays
8-10, terminal, weakly curved, almost entirely enodulate, slender,
2-4 cm. long, terminally about 8-10-pedicelled; pedicels umbellu-
lately grouped, finally 6-8 mm. long and 1-2 mm. thick. Florets
unknown. Drupes (sub-mature ones seen) ovoid, compressed or
trigonous; calyx about 1 cm. tall and 7-8 mm. thick, apically con-
stricted; stylopodium conic, about 3 mm. tall (including the 2 or 3 or
rarely 4 stigmas, these-scarcely definite in shape), true style lacking.

Type: Otto Degener 21,790, tree 30 ft. tall, sparingly branched
and spindly, Kulani Forest, Isl. Hawaii, Feb. 3, 1952 (Chi.).

Distribution : Known only from type locality in eastern (slightly
southeastern) Hawaii.

Specimens examined: Degener 21,790 (type, Chi., my photo-
graph no. 4,362a, not 4,362 as originally printed: isotypes, Berl.;
Bish.; Calif. Acad.; Gray; Kew; N.Y.; Par.; Phila.; Univ. Calif.).

Close to var. co. tenuistylis but differing distinctly above the
calyx-limb in the drupes. From var. v. rhynchocarpa of the same
region, its thicker and shorter drupes seem easily to separate it.

15. Tetraplasandra kauaiensis (H. Mann) Sherff, Bot. Leafl.
no. 7: 10. 1952; Heptapleurum kauaiense H. Mann, Proc. Amer.
Acad. 7: 168. 1867; Agalma kauaiense Seem. Jour. Bot. 6: 103. 1868;
Pterotropia kauaiensis (H. Mann) Hillebr. Fl. Haw. Isls. 150. 1888;
Heptapleurum kauiense Mann ex Drake del Cast. 111. Fl. Ins. Mar.
Pacif. 183. 1890 (sphalm); Dipanax kauaiensis (H. Mann) Heller,
Minnesota Bot. Studs. 1: 871. 1897.

a. Stigmas commonly 6 or 7; tree of Wahiawa Mts., southern
Kauai var. 77. koloana.

a. Stigmas fewer, trees from elsewhere.
6. Stylopodium none or obsolete,
c. Stigmas normally 3 or 4.

124 FIELDIANA: BOTANY, VOLUME 29

d. Leaflets large, their blade often 1.5-2 dm. long and 7-11 cm.
wide, lower petiolules mostly 1-3 cm. long, pedicels
finally 5-13 mm. long.
e. Leaflets more or less glabrescent beneath; native of

Kau, southern Hawaii var. 7. grandis.

e. Leaflets densely tomentose beneath; native of north-
eastern East Maui var. e. nahikuensis.

d. Leaflets smaller, their blade 4-10 cm. long and 2-5 cm.
wide, lower petiolules under 5 mm. long, pedicels in
fruit only 2-4 mm. long; native of southeasternmost

West Maui var. 9. occidua.

c. Stigmas normally 2, leaflets with definitely cordate base;

native of Lanai var. /3. dipyrena.

b. Stylopodium normally distinct (but in var. robustior very

short),
c. Stigmas normally 5 (rarely 4); native of northwestern

Kauai var. f . robustior

c. Stigmas normally 3 (rarely 2); native of southwestern East

Maui var. 5. intercedens.

c. Stigmas normally 4 (sometimes 3 or 5); native of western
Kauai var. a. kauaiensis.

Tree 9-15 m. tall, the smooth trunk erect and sparingly divided
at its top into a few ascending, stout branches, these bearing large,
terminal leaf-whorls, the young shoots and inflorescence tomentose
with fawn-colored or tawny, stellate scales. Leaves 4-9 dm. long,
leaflets 11-21; petiole 8-18 cm. long; blade narrowly oblong to
ovate-oblong, at base more or less oblique and rounded to truncate
or barely subcordate, at apex acuminate to rounded, chartaceous to
somewhat coriaceous, on lower surface densely stellate-tomentose,
on upper surface glabrescent or here and there very minutely but
at times abundantly sprinkled with stellate hairs or scales, 6-18 cm.
long and 3-8 cm. wide; petiolules 2-15 mm. long. Inflorescence a
large panicle, its usually 5 primary branches 1-3 dm. long and
mostly alternate on a rhachis about 5-7 cm. long; the rays or
secondary branches 4-7 cm. long, mostly alternate except for the
10 terminal umbellately clustered ones; flowers racemose toward
and subumbellulate (umbellets 10-20-flowered) at the tips of the rays
on pedicels 3-5 (finally sometimes up to 11) mm. long. Corolla
about 11-12 mm. across at anthesis; petals (spreading at anthesis,
5-6 mm. long, tomentose) and stamens 6 or 7 or rarely to 9. Ovary

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 125

usually 3-5-(usually 4-) celled. Drupe ovoid but 4-angled, about
10-12 mm. tall, ringed with calyx-limb below, at, or above its middle;
stylopodium distinct but under 1 mm. long, bearing 3-5 usually
4 stigmas.

Type: Horace Mann, Jr., & William T. Brigham 606, alt. 2,000-
3,000 feet, mountains above Waimea, Isl. Kauai (Gray).

Distribution: Island of Kauai, western part.

Specimens examined (all from Isl. Kauai) : Degener & Greenwell
21,578, forested gulch, edge of Kumuweia Ridge east of Waineke
Swamp, Jan. 8, 1952 (Bish.; Brit.; Carn.; Chi.; N.Y.; etc.); Degener
& Greenwell 21,579, tree 40 ft. tall, commonly locally in forest at
2,000 ft., Kopewai (Kopiwai) Trail, Kokee region, Jan. 11, 1952
(Berl.; Bish.; Brit.; Cam.; Chi.; Gray; Kew; N.Y.); C. N. Forbes
430-K, Kaholuamanu behind Waimea, September, 1909 (Bish.; Par.;
U.S.); Forbes 1,097-K, large tree 40-50 ft. tall, trunk 1.5-2 ft. in
diam., bark gray and lichen-covered, Waimea Drainage Basin, west
side, July 3-Aug. 18, 1917 (Bish., 2 sheets) ; Mann & Brigham 606
(type, Gray, my photograph no. 4,310: isotype, U.S., my photo-
graph no. 4,311); J. F. Rock, drier districts below Kaholuamanu,
Mar. 5, 1909 (Am.); Rock 2,488, Kaholuamanu, Mar. 5, 1909
(Arn.; Bish.; Gray); Rock 2,488a, Halemanu, October, 1909 (Bish.;
Gray) ; Rock & Marshall, borders of a meadow called Malua Poha,
back of Halemanu, October, 1909 (Arn.).

Tetraplasandra kauaiensis var. /3. dipyrena (H. Mann)
Sherff, Bot. Leafl. no. 7: 11. 1952; Heptapleurum dipyrenum
H. Mann, Proc. Amer. Acad. 7: 168. 1867; Dipanax Mannii Seem.
Jour. Bot. 6: 141. 1868; Pterotropia dipyrena (H. Mann) Hillebr.
Fl. Haw. Isls. 150. 1888; Dipanax dipyrena (H. Mann) Heller,
Minnesota Bot. Studs. 1: 870. 1897.

Small tree, probably little more than 6 m. tall. Leaves 3-6 dm.
long, leaflets 11-15, somewhat smaller and more coriaceous than in
var. kauaiensis, also more acuminate at tip and more definitely
cordate at base, on upper surface finally more nearly or completely
glabrous. Rhachis of inflorescence 5-8 cm. long; peduncles 5,
subumbellately or somewhat alternately disposed, at times 4 dm.
long or more; rays 2-5 cm. long; pedicels shorter, 2-5 mm. long
even in fruit. Drupe ovoid but usually somewhat 4-angled, 12 mm.
long, stylopodium none, stigmas 2 (in type) or at times 3 (or even
4, fide Hillebrandii) ; calyx-limb a ring at or above middle.

Type: Horace Mann, Jr., & William T. Brigham 349, a small
tree 15 or 20 ft. tall, Isl. Lanai (Gray).

126 FIELDIANA: BOTANY, VOLUME 29

Distribution: Known today only from Island of Lanai, but Hille-
brand (loc. cit.) listed also East Maui and Hawaii.

Specimens examined (all from Lanai) : William Hillebrand, ann.
1870 (Gray); Mann & Brigham 349 (type, Gray, my photograph
no. 4,317) ; George C. Munro, valley south of Puhielelu, Apr. 10, 1916
(Bish.); Munro, Lanai, Nov. 17, 1915 (U.S., 2 sheets); Munro 469,
ridge of Halelepakai (Haalelepaakai), Oct. 17, 1915 (Bish.; Univ.
Calif.).

Tetraplasandra kauaiensis var. 7. grandis Sherff, Bot. Leafl.
no. 7: 14. 1952.

Leaflets rather large, blade often 2 dm. long and 7-11 cm. wide,
oblong to (especially for the lower ones) ovate, at apex obtuse to
acuminate, at base oblique and truncate to cordate, often more or
less glaucous above (at least in dry state), becoming more or less
glabrous beneath; median petiolules 4-8 mm. lowermost ones 1.5-3
cm. long. Panicle very large, often almost 1 m. long, its rhachis
at times 1.5 dm. long, peduncles 5-7.5 dm. long; rays numerous
(15-20 placed racemosely along the peduncle and 7-9 umbel-
lately at peduncle's apex), up to 9 cm. long; pedicels slender,
commonly 9-16 in a terminal umbellet and 3-6 irregularly scattered
along the ray, finally 7-13 mm. long. Petals 6-8, lanceolate to
ovate-triangular, at anthesis 6-8 mm. long; stamens 6-8. Drupes
commonly trigonous otherwise slenderly ovoid, annulate above the
middle, mitriform above the annulus (calyx-limb), at apex bearing
a very tiny or obsolete stylopodium and 3 or more rarely 2 stigmas.

Type: Joseph F. Rock 10,019, alt. 2,000 feet, dense Hilea forest,
gulch from Kanolohu to Kumanua, Kau, Isl. Hawaii, Jan. 17,
1912 (Gray).

Distribution: Known only from type locality in southern Hawaii.

Specimens examined (Isl. Hawaii) : Rock 10,019 (type, Gray, my
photograph no. 4,320: isotypes, Arn., 7 sheets, my photographs nos.
4,321, 4,322, and 4,323; Bish.; Gray, 3 sheets; Par., 2 sheets; Univ.
Calif.. 3 sheets).

With this variety should be compared a tree found in eastern-
most Hawaii: Carl Skottsberg 1,125, lava field west of Pahoa, Sept. 9,
1922 (Bish.; Goth.). I have seen two leaves only; these glabrous
on both surfaces, 9-foliolate, the leaflets with faintly sinuate margins.
Flowers and fruits are much to be desired.

Tetraplasandra kauaiensis var. 5. intercedens Sherff, Bot.
Leafl. no. 7: 13. 1952.

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 127

Leaflets 13-19, blade commonly oblong, that of the laterals
apically acute or acuminate and basally truncate or barely cordate.
Peduncles 4-6, umbellately grouped, often 3-3.5 dm. long, the rays
slender and up to 8 cm. long; a few (4-10) florets grouped in an
umbellet, the others racemose along the ray; pedicels about 2-3 mm.
long at anthesis, in fruit about 4-5.5 mm. long. Drupes globose-
ovoid, about 9-10 mm. tall (including the stylopodium 1 mm.
long and the commonly 3 rarely 2 stigmas), annulate above the
middle with the calyx-limb, hemispherical or tumid-conic at top.

Type: Joseph F. Rock 8,641, a very tall tree, alt. 3,000 ft., dry
districts, Auwahi, southern slopes of Mt. Haleakala, East Maui,
Nov. 12, 1910 (Bish.).

Distribution: Southwestern East Maui.

Specimens examined (all from East Maui) : C. N. Forbes 1,973-M,
80 ft. tall, trunk 2 ft. in diam., only about 14 trees seen, 30-40 ft.
average height, Auwahi, Mar. 15, 1920 (topotypes, Bish.; Calif.
Acad.) ; Forbes 2,169-M, common in the dying forest, 30-35 ft. tall,
trunk 6-8 inches in diam., bark smooth and gray, umbellately
branched or nearly so at top, slope of Haleakala, Kula side, Apr. 8,
1920 (Bish., 2 sheets; Par.; Univ. Calif.; U.S.); William Hillebrand,
East Maui (U.S.); Rock 8,641 (type, Bish., my photograph no.
4,318: isotypes, Arn., 4 sheets; Goth.; Gray, my photograph no.
4,319; Univ. Calif., 2 sheets; U.S., 2 sheets).

In this variety are found the few (3 or sometimes 2) stigmas
characteristic of most var. dipyrena material combined with the
possession of a distinct stylopodium as found in var. kauaiensis
(hence the name "intercedes"). It is the Pterotropia dipyrena of
Rock's illustrations (Indig. Trees Haw. Isls. pis. 144 and 145. 1913),
based upon his collections on the southwestern (his "southeastern"
was an error!) "and strictly southern slopes of Mt. Haleakala."

Tetraplasandra kauaiensis var. e. nahikuensis Sherff, Bot.
Leafl. no. 7: 13. 1952.

Leaflets oblong or at times ovate-oblong or even ovate, blade
often 1.5-1.8 dm. long and 8-10 cm. wide, at apex commonly acumi-
nate, at base oblique and rounded to truncate, petiolule up to 2.5
cm. long. Inflorescence large, the drooping peduncles 4 dm. long;
rays slender, 4-10 cm. long, irregularly alternate and a few termi-
nally umbellate; pedicels 5-12 mm. long, moderately slender, 5-8
disposed in a terminal umbellet the rest racemosely. Petals 4-5
mm. long. Drupes ovoid, annulate at or slightly below the middle

128 FIELDIANA: BOTANY, VOLUME 29

with the calyx-limb, apically conic, without a distinct stylopodium,
about 8-9 mm. tall, stigmas now 3 now 4.

Type: Joseph F. Rock, Nahiku, northeastern East Maui, May 10,
1911 (Am.).

Distribution: Known only from northeastern East Maui.

Specimens examined (all from East Maui) : C. N. Forbes 239-M,
large tree, 35 ft. tall, Nahiku, July, 1910 (topotypes, Bish.; U.S.);
Joseph F. Rock, Nahiku, Jan. 14, 1909 (topotypes, Bish., 2 sheets,
my photograph no. 4,325); Rock, Nahiku, May, 1911 (topotypes,
Arn., 2 sheets) ; Rock, Nahiku, May 10, 1911 (type, Arn., my photo-
graph no. 4,324: isotypes, Gray, 2 sheets); Rock 8,641a, Nahiku,
Jan. 1-8, 1909 (topotypes, Bish., 3 sheets) ; Rock 12,510, alt. 2,000
ft., Nahiku, April, 1911 (topotypes, Bish., 3 sheets; Gray, 2 sheets;
Univ. Calif.; U.S.).

Rock (Indig. Trees Haw. Isls. 359. 1913) treated this and the
tree above described as var. intercedens as Pterotropia dipyrena.
Var. intercedens, from southern and southwestern East Maui, has
the drupes annulate with the calyx-limb above the middle, the upper
part swollen-conic or hemispherical, the 3 or rarely 2 stigmas on a
definite stylopodium; pedicels at fruiting about 4~5.5 mm. long.
Var. nahikuensis, from northeastern East Maui, has the drupes
annulate at or a little below the middle, the upper part conic, the
stigmas 3 or as often 4, a definite stylopodium lacking; pedicels at
fruiting 5-12 mm. long.

Tetraplasandra kauaiensis var. f . robustior Sherff , Bot.
Leafl. no. 7: 12. 1952.

Branches and branchlets of inflorescence twice as thick as in
var. kauaiensis, the flowering umbellets a half wider, petals of the
larger florets lanceolate and about 6 mm. long. Drupes broadly
oblong in outline, about 12 mm. tall and about 10 mm. thick,
obtusely rounded at apex, stylopodium very short, stigmas com-
monly 5 rarely 4.

Type: Amos Arthur Heller 2,884, large tree, 30-40 ft. tall, with
soft wood, on the brow of the plateau, at an elevation of about
3,500 feet, above Kaholuamanu, Isl. Kauai, Oct. 16, 1895 (Bish.).

Distribution : From above Kaholuamanu to Kokee, northwestern
Kauai.

Specimens examined (all from Isl. Kauai): Cranwell, Selling,
& Skottsberg 3,095, Kokee, Aug. 25, 1938 (Bish.; Goth.); Degener &

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 129

Wiebke 2,180, large trees in open forest with much precipitation,
Kokee, June 23, 1926 (Calif. Acad.; N.Y., 4 sheets; Univ. Calif.);
A. A. Heller 2,884 (type, Bish., my photograph no. 4,337: iso-
types, Am., my photograph no. 4,338; Chi.; Corn.; Gray; Mo.,
2 sheets; Phila.; Univ. Calif.; U.S.); L. H. MacDaniels 800, height
25 m., open forest, alt. 1,300 m., Kokee, Feb. 18, 1927 (Bish.);
Carl Skottsberg 1,010, near Kokee ranger station, Oct. 28, 1922
(Goth.).

Typical T. kauaiensis is found on Kauai at Halemanu, Kumu-
weia Ridge, drier districts below Kaholuamanu, and (type locality)
at 2,000-3,000 ft. altitude, Waimea. It is replaced at Kokee and
on the brow of the plateau above and northeast of Kaholuamanu
with the var. robustior.

Tetraplasandra kauaiensis var. 77. koloana Sherff, Bot. Lean 1 ,
no. 7: 12. 1952.

Leaflets 9, their blade narrowly oblong to widely ovate, at
times 1.8 dm. long and 1.1 dm. wide, that of the lateral ones more
or less obtuse at the apex and rounded or subtruncate at the oblique
base. Peduncles 3, spreading, 17 cm. long; rays slender, more
often 4-6 cm. long; pedicels about 8-12 in a terminal umbellet,
slender, 11-16 mm. long. Stigmas commonly 6 rarely 7; drupes
immature, the conical but laterally concave apex 3.5 mm. tall,
its stylopodium scarcely 0.5 mm. tall.

Type: Charles Noyes Forbes 276-K, tall tree, Wahiawa Mountains,
southern Kauai, August, 1909 (Bish., 2 sheets).

Distribution: Known only from type locality in southern Kauai.

Specimens examined: Forbes 276-K (2 type sheets, Bish., my
photographs nos. 4,339 and 4,340).

Tetraplasandra kauaiensis var. 0. occidua Deg. & Sherff ex
Sherff, Bot. Leafl. no. 8: 7. 1953.

Tree 8 m. tall. Leaves 1.5-3 dm. long; leaflets 9-15, lowermost
ones with ovate or even oval-subrotund blade, the others with an
oblong or the terminal one with an obovate blade, all small (blade
4-10 cm. long and 2-5 cm. wide), at tip obtuse to scarcely acute, at
base oblique and truncate to weakly cordate, coriaceous, glabrous
and often glossy above, more or less stellate-furfuraceous beneath;
lateral petiolules only 1-2 (rarely -5) mm. long. Inflorescence
smaller, its rhachis 1-3 cm. long, primary branches 3, under
1.5 dm. long; rays (secondary branches) slender, finally more often

130 FIELDIANA: BOTANY, VOLUME 29

2.5-4 cm. long, racemosely and (at apices of 5 branches) um-
bellately disposed; florets (only a few seen) rarely below commonly
at tips of rays and about 5 or 6 in an umbellet; pedicels at first
very short (under 2 mm.) and thick, finally (with fruits) 2-4 mm.
long. Petals and stamens 5. Mature drupes ovoid, calyx about
6-7 mm. tall, the swollen-conic apex about 4.5-5 mm. tall including
the 3 or more often 4 subsessile stigmas.

Type: Degener, Tarn, Tousley, & Barber 22,025, a 25-foot tree
(its seed eaten by rats), topping lowly forested slope in rainy region,
mauka of [i.e., inland from] McGregor, southeasternmost West
Maui, Mar. 9, 1952 (Chi.).

Distribution: Known only from type locality in West Maui.

Specimens examined: Degener et alii 22,025 (type, Chi., my
photograph no. 4,361: isotypes, Bish.; Brit.; Carn.; Chi., my photo-
no. 4,362; N.Y.).

16. Tetraplasandra turbans Sherff, Bot. Leafl. no. 7: 16. 1952.

Tree 12.5 m. tall with trunk 2.5 dm. thick, with whitish
and pulverulent branchlets, leaves 5 dm. long (including the very
minutely pulverulent-hispidulous petiole, this about 11-12.5 cm.
long and 6 mm. thick); leaflets about 17-19, at apex acute to
almost acuminate, at base oblique and truncate to subcordate,
coriaceous, glabrous above, more or less densely stellate-furfuraceous
beneath, blade of lowermost ones deltoid-oblong and 5.5 cm. long
by 3 cm. wide, blade of the others narrowly to moderately oblong
and 7-10.5 cm. long by 2.5-3.5 cm. wide; petiolules 1-5 mm. long,
pulverulent-furfuraceous; rhachis more or less glabrescent. Inflores-
cence furfuraceous (a lone complete one seen), terminating a rhachis
4 cm. long and 7 mm. thick, umbellately 5-branched or
-pedunculate; peduncles slender (3 mm. thick), 12-15 cm.
long, covered with scars, bearing toward apex about 10-15 very
small branchlets, these only 1 cm. long, covered with scars, at
and near their tip very densely floriferous, 10-16 florets most often
crowded into an umbellet only 7-9 mm. thick. Pedicels under 1
mm. long. Calyx basal, disk-like; corolla (scarcely mature) globose-
ovoid, about 3 mm. tall, 7-8-angulate; petals (as yet cohering but
doubtless spreading at full anthesis) and stamens 7 or 8. Drupe
(a lone, very immature one seen) on a pedicel 3.8 mm. long; its
body (the ovary perfectly free from the basal calyx and hence com-
pletely superior) tetragonal (and obovoid), 4.5 mm. tall and 3.5
mm. thick, at the depressed-truncate top crowned with a small mar-
ginal limb; stigmas 4, sessile.

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 131

Type: Charles Noyes Forbes, Makaha Valley, Kaala Range, Isl.
Oahu, Feb. 12-19, 1909 (Bish., 2 sheets).

Distribution: Northwestern Oahu.

Specimens examined: Degener & Hatheway 20,865, a 40-ft. tree
with 10-in. trunk, open sunny forest at 1,700 ft., easternmost gulch
of West Branch of East Makaleha Valley, Oct. 9, 1950 (Am.; Berl.;
Bish., Brit.; Cam.; Chi.; N.Y.; etc.); Forbes, Makaha Valley, Isl.
Oahu, Feb. 12-19, 1909 (2 type sheets, Bish., my photograph nos.
4,335 and 4,336: isotypes, Calif.; Par.).

Forbes had labeled his collection Pterotropia kauaiensis var. /3.
Hillebr., a synonym for Tetraplasandra micrantha of this text. In
T. micrantha, it is true, the calyx is similiar at anthesis to the calyx
here, but rapidly elongates upward in early fruit so as to be adnate
nearly or fully half-way up the ovary; in T. turbans, on the other
hand, the calyx remains discoid at the bottom of a completely
superior ovary in the maturing drupe, much as in T. gymnocarpa.
In both species (T. turbans and T. micrantha), the leaflets are very
furfuraceous beneath with myriads of tiny, whitish, stellately
setulose scales, but the leaflets of T. micrantha are thinner and attain
a length and breadth nearly twice as great. Moreover, in T. micran-
tha the inflorescence is much larger, its rhachis or peduncles often
3 dm. long; the florets are many fewer (only about 8-10 in a lone
terminal umbellet, with several florets placed singly 1-3 cm. lower
down on the flowering-branchlet) and on slender pedicels becoming
5-6 mm. long in early fruits (ripe fruits not seen).

Much to be desired are mature drupes. The description of an
immature drupe rests for the present on the one found in the packet
accompanying an isotype (Calif.). The thin, shallow, coroniform
limb at the apical margin is a perplexing or disturbing character
(whence the epithet, "turbans"), in that it simulates a calyx-limb
and thus falsely suggests an inferior ovary.

17. Triplasandra micrantha Sherff, Bot. Leafl. no. 7: 11.
1952; Pterotropia kauaiensis var. /3. Hillebr. Fl. Haw. Isls. 151.
1888; Pterotropia kaalae Skottsb. Meddel. Goteb. Bot. Tradg. 10:
151. 1936.

Leaves up to 7.5 dm. long and to 3 dm. wide, rhachis glabrous
except when young; leaflets at least 7 or 9 (doubtless more), thin,
lateral ones oval-oblong to elongate-oblong and more or less parallel-
sided (terminal one obovate), at apex acuminate or occasionally
emarginate, at base truncate or rounded-truncate, their blade up to

132 FIELDIANA: BOTANY, VOLUME 29

2 dm. long and to 6.5 cm. wide (that of terminal one 8 cm. long
and 6 cm. wide), glabrous above, on lower surface at first densely,
later sparsely whitish-farinose with minute stellate hairs; lateral
petiolules slender and but 4-5 mm. long. Inflorescence apparently,
if we may rely upon the two cited sheets of Hillebrand material, a
slender peduncle 3 dm. long, with 15-20 bracteal scars throughout
its length, scantily and very minutely hispidulous, umbellately
branched at apex with 6 delicate rays; these up to 9.5 cm. long
and 1.5 mm. thick, stellate-hispidulous, in distal third or half race-
mosely 1-5-pedicellate and at tip umbellulately 4-10-pedicellate;
pedicels very delicate (about 0.5 mm. thick), 3-7 mm. long, stellate-
hispidulous. Florets small; calyx obconic, about 2 mm. tall and
broad, stellate-hispidulous; petals 8, outwardly stellate-hispidulous,
3-3.5 mm. long and at base 1.1-1.3 mm. wide, at tip cucullately
inflexed. Stamens 7 or 8, about 2.5 mm. long, the short, curved
filament 1.2 mm. long, the anthers 2-2.2 mm. long and 0.8-0.9 mm.
thick. Ovary 3-3.5 mm. thick. Stylopodium at most 1.3 mm. tall,
disappearing with growth of drupe, leaving the commonly 2 or 3
(rarely 4) stigmas sessile on the conical apex. Mature drupes (a
single, detached one seen in packet at Kew) ovoid, glabrous or
glabrate; calyx broadly hemispherical, 11 mm. thick at top and
6 mm. tall, its limb subobsolete; portion above calyx swollen and
mitriform-conic, 9 mm. tall including the sessile 2 or 3 (or even 4)
stigmas.

Type: William Hillebrand, Kaala, northwestern Oahu (formerly
in Berl., but later destroyed by war).

Distribution: Northwestern Oahu.

Specimens examined (Isl. Oahu): Hillebrand, Makaleha, Kaala
(topotype or perhaps isotype, U.S., my photograph no. 4,316);
Hillebrand, Kaala Mountains, 1871 (topotype or perhaps isotype,
Kew, my photograph no. 4,312, excluding inverted fragment of
panicle at left); Hillebrand 266, Waianae Range, June, 1861 (Kew,
my photograph no. 4,313).

Certain details of flower and fruit are taken from Skottsberg
(loc. cit.}. Apparently Hillebrand had somewhat mixed material
in mind for his description. The Kew sheet above cited has, in
addition to leaves and inflorescence matching those on the U.S.
sheet (and, with those, serving as the principal basis of my des-
cription), an inverted fragment of an immature panicle of some
other species of Tetraplasandra. This has three scattered, stoutish,
squarrose lateral branches 4-5 cm. long (the apex is missing) and

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 133

is densely stellate-hispidulous. It has been ignored in drawing my
description.

18. Tetraplasandra gymnocarpa (Hillebr.) Sherff, Bot. Leafl.
no. 7: 12. 1952; Pterotropia gymnocarpa Hillebr. Fl. Haw. Isls. 150
and 151. 1888; Heptapleurum gymnocarpum (Hillebr.) Drake del
Cast. 111.. Fl. Ins. Mar. Pacif. 183. 1890; Dipanax gymnocarpa
(Hillebr.) Heller, Minnesota Bot. Studs. 1: 870. 1897.

a. Stigmas commonly 2 rarely 3; drupes prismatic, 13-15 mm. tall,
1-4 in an umbellet; native of the Konahuanui-Olympus region,

southeastern Oahu var. 0. leptocarpa.

a. Stigmas usually 3-5; drupes globose-ovoid to ovoid, more to an

umbellet.
6. Petals usually 6; leaflets often 6-8 cm. wide; stigmas now 3

now 4, very rarely 2.

c. Petals 6-8 mm. long, pedicels finally often 1.5-2.5 cm. long;
drupes ovoid, 11-14.5 mm. tall; native of Punaluu region,

northeastern Oahu var. 7. megalocarpa.

c. Petals about 4 mm. long, pedicels under 1.4 cm. long even
in fruit; drupes globose-ovoid to ovoid, 6-9 mm. tall;
widespread almost throughout the Koolau Range of

Oahu var. a. gymnocarpa.

b. Petals usually 5; leaflets mostly 3-5 less often to 6 cm. wide;
stigmas now 3 now 4 or at times 5; native of the Pupukea-
Kahuku Trail region of far northern Oahu.

var. 8. pupukeensis.

Small tree 3.6-6 m. or sometimes to 7.5 m. tall, only the youngest
shoots mealy, otherwise glabrous. Leaves 3-5.5 dm. long, rather
dark-green; leaflets 9-17, oblong to ovate-oblong or the lowermost
ones ovate, at apex obtuse or more often obliquely acuminate, at
base oblique and rounded, chartaceous to coriaceous, glabrous on
both surfaces, shiny above, blade 7.5-15 cm. long and 3-8 cm. wide,
petiolule 4-18 mm. long. Rhachis of panicle rather short (2-5 cm.)
with 3-5 umbellately radiating primary branches or peduncles, these
1-3.7 dm. long; secondary branches or rays slender, elongate (more
often 5-11 cm. long), disposed umbellately at tip of and racemosely
along the peduncle, floriferous mostly by terminal umbellets of 10-16
florets and often by a few florets racemosely disposed somewhat
lower along the ray. Pedicels slender, 6-9 mm. or in fruit up to
14 mm. long. Calyx very short, with a loosely undulating border.
Petals 6 or rarely 7, cohering at the apex, about 4 mm. long. Stigmas

134 FIELDIANA: BOTANY, VOLUME 29

now 3 now 4, rarely 2; stylopodium none. Drupes globose-ovoid
to ovoid, 6-9 mm. tall and 4.5-6.5 mm. thick, several-angled. Ovary
practically free and naked, 3- or 4- or very rarely 2-celled; pyrenae
thin-papery, ovoid, beaked above and faintly notched below the beak.

Type: Rev. John M. Lydgate, valley of Niu, Island of Oahu
(formerly extant at Berlin) .

Distribution: Koolau Range of Oahu.

Specimens examined (all from Isl. Oahu) : R. W. Baxter, tree 20
ft. tall, alt. 1,100 ft., Pupukea-Kahuku Trail, February, 1939 (Mo.,
2 sheets) ; E. H. Bryan, Jr., tree 10 m. tall, trunk diameter 15 cm.,
alt. 900 ft., Pupukea, Jan. 23, 1929 (Bish., 2 sheets) ; Otto Degener,
Pupukea-Kahuku region, Mar. 31, 1929 (N.Y.); Degener & Martin
Martinez 11,920, in rain-forest, C.C.C. Trail, Aiea, Mar. 15, 1936
(U.S.) ; C. N. Forbes 1,638-0, tree 20-25 ft. tall, flat-topped, branches
subumbellate, umbels in threes, leaves thick, dark-green above and
pale beneath . . . midrib dark-red, stamens 6, petals 6, trail between
Konahuanui and Olympus, back of Manoa Valley, Feb. 14, 1911
(Bish.) ; Forbes 2,062-0, ridge north of Waimea Valley, Feb. 10-13,
1915 (Bish.) ; Forbes 2,182-0, slopes of Konahuanui, Mar. 11 and 12,
1915 (Bish.) ; Forbes with C. M. Cooke, Jr., mountains between Puna-
luu and Kaipapau, May 3-9, 1909 (Bish.) ; Forbes with J. C. Bridwell
2,457-0, between Niu and Wailupe, Apr. 11, 1917 (Bish.); D. Wesley
Garber 238, Konahuanui-Olympus Trail, Feb. 15, 1920 (Bish., 2
sheets); William Hillebrand, Niu (topotype, Gray, my photograph
no. 4,327); Hillebrand & Lydgate, Niu (isotype, Bish., my photo-
graph no. 4,326) ; E. Y. Hosaka 111, in woods, alt. 1,500 ft., Pupukea-
Paumalu Forest Reserve, Jan. 12, 1930 (Calif. Acad.) ; L. H. Mac-
Daniels 114, small tree 5 m. tall, exposed rain-forest, lee slope, alt.
850 m., Konahuanui, Dec. 21, 1926 (Bish.) ; MacDaniels 535, Pupu-
kea Military Trail, Jan. 29, 1927 (Bish.); A. Meebold, alt. 1,500-
1,800 ft., Pupukea, May, 1932 (Bish.); Kazuto Nitta 73, slightly
damp slope, Pupukea, Jan. 12, 1930 (N. Y.) ; Joseph F. Rock 406a, Ko-
nahuanui, Jan. 8, 1910 (Bish.; Calif. Acad.; U.S.) ; D. P. Rogers, 20 ft.
below Kipapa Trail, head of Waiawa Valley, Waipio, Sept. 29,
1946 (Bish.); Harold St. John 10,104, tree 20 ft. tall, in thicket,
alt. 1,900 ft., Kaluanui, Nov. 30, 1929 (Bish., 2 sheets; Par.; Univ.
Calif.); St. John 10,147, tree 20 ft. tall, woods along ridge, alt.
1,200 ft., Pupukea-Paumalu Forest Reserve, Jan. 12, 1930 (Am.;
Bish., 2 sheets; Par.; Univ. Calif.).

Tetraplasandra gymnocarpa var. /3. leptocarpa Sherff, Bot.
Lean. no. 7: 15. 1952.

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 135

Leaflets 11, blades more or less oblong or (the three terminal
ones) oblong-oblanceolate, at apex acute to subacuminate, at base
oblique and rounded, weakly powdery-mealy beneath, the slender
petiolules and the rhachis glabrescent. Inflorescences becoming
glabrous, rhachis 16 cm. long, its branches or peduncles 20-23
cm. long, umbellately grouped; rays 4-6 at tip of and up to 4
irregularly but remotely scattered along a peduncle, slender, 4-6 cm.
long. Florets not seen. Drupes few (1-4) in an umbellet (this
terminal on a ray), prismatic, 4-5-angled, toward apex conic or
mitriform, at apex commonly 2- rarely 3-stigmatate, 13-15 mm.
tall, under 5 mm. thick, stylopodium lacking; pedicels finally 10-15
mm. long.

Type: Charles Noyes Forbes 1,608-0, trail between Konahuanui
and Olympus, Island of Oahu, November, 1910 (Bish., 2 sheets).

Distribution: Known only from type locality in southeastern
Oahu.

Specimens examined: Forbes 1,608-0 (2 type sheets, Bish., my
photographs nos. 4,330 and 4,331).

Tetraplasandra gymnocarpa var. 7. megalocarpa Sherff,
Bot. Lean. no. 7: 15. 1952.

Leaflets often 6-7 cm. wide, very oblique at base. Peduncles
(primary branches of the inflorescence) 3.5 dm. long; rays (second-
ary branches) very slender, elongate (often 10-13 cm. long) ; pedicels
more elongate, finally often 1.5-2.5 cm. long. Petals 6, lanceolate
or lanceolate-linear, 6-8 mm. long; stamens 6. Drupes ovoid,
11-14.5 mm. tall, stigmas now 3 now 4 very rarely 2, stylopodium
absent.

Type: Joseph F. Rock & Genet P. Wilder 8,827, alt. 2,000 feet,
in one of the inner gulches of the Koolau Mountains, Punaluu,
Island of Oahu, August, 1911 (Gray).

Distribution: Known only from type locality in northeastern
Oahu.

Specimens examined (all from Isl. Oahu) : Degener, Park, Bush,
Potter, & Topping 11,921, in rain-forest, summit of Koolau Range
beyond Pig-God Trail, Punaluu, July 4, 1935 (topotypes, Bish., my
photograph no. 4,334; Corn.; Del., 2 sheets; Mo., 2 sheets; U.S.);
Rock 406, Punaluu, Dec. 24-29, 1908 (topotypes, Bish.; Gray; Univ.
Calif.); Rock 421, same place, Dec. 24-28, 1908 (topotypes, Gray,
2 sheets); Rock 449, in the woods of Punaluu, above Hauula, Dec.
24-28, 1908 (topotype, Arn., my photograph no. 4,315); Rock &

136 FIELDIANA: BOTANY, VOLUME 29

Wilder 8,827 (type, Gray, my photograph no. 4,332: isotypes, Arn.,
4 sheets, my photograph no. 4,333; Bish.; Gray, 2 sheets; U.S.);
Olof H. Selling 3,671, Koolau Mts. at Pimaluu, Sept. 27, 1938
(topotype, Bish.).

Rock (Indig. Trees Haw. Isls. 355. 1913), in his treatment of
T. gymnocarpa (Pterotropia gymnocarpa in his text), described the
drupes of his own specimens as "12 to 15 mm. long, and about 7
mm. in diameter," thus showing that his plate 143, which was made
from his own material, was none other than an illustration of var.
megalocarpa, as it has since been named.

Tetraplasandra gymnocarpa var. <5. pupukeensis (Deg.)
Sherff, Bot. Leafl. no. 8: 12. 1953; Pterotropia gymnocarpa var.
pupukeensis Deg. Fl. Haw. fam. 281. Aug. 24, 1938.

Large tree with spreading crown bearing few, thick branches.
Leaves 2-5.5 dm. long, coriaceous, on upper surface dull (in type
collection) or at least in dry state more or less glossy; leaflets 9-25,
oblong-ovate, at apex acuminate to acute or rarely rounded, at base
truncate to obliquely rounded or rarely acute, the central blades
6-13 cm. long and 3-5 or less often to 6 cm. wide but the basal ones
shorter and relatively wider and terminal one shorter and relatively
narrower, petiolule commonly 3-7 mm. long. Inflorescence some-
what furfuraceous, borne laterally when flowering and fruiting, the
rhachis 5 cm. long, bearing about 8 umbellately or subumbellately
grouped primary branches or peduncles; these spreading to some-
what drooping, 15-25 cm. long, bearing rays scattered racemosely
and at the tip grouped umbellately; rays mostly 3-6 cm. long,
floriferous now only in terminal umbellets of 8-12 florets now also
more or less racemosely in their terminal half. Petals 5 or very
rarely 6, maroon, thick, 6 mm. long, 2.5-4 mm. wide in same flower,
spreading to reflexed, ovate-lanceolate, apically acute. Stamens 5
or very rarely 6. Stigmas now 3 now 4 or at times 5; drupes broadly
globose-ovoid, 10-12 mm. tall.

Type: Degener, Salucop, & Arlantico 11,919, several trees in
dense forest, Pupukea-Kahuku Trail near Puu Moa, Isl. Oahu, Apr.
3, 1938 (herb, not specified).

Distribution: Koolau Range from northern end interruptedly
southeastward to the upper parts of Kipapa Gulch, Island of Oahu.

Specimens examined (all from Isl. Oahu) : Otto Degener 18,015
pro parte, lower forest, Pupukea-Kahuku Trail makai of Puu Moa,
May 28, 1932 (topotypes, Mo.; N.Y., my photograph no. 4,329);

SHERFF: REVISION OF THE GENUS TETRAPLASANDRA 137

Degener 18,253, "precise locality unknown," "Pupukea-Kahuku
region, on Pupukea end," Mar. 31, 1929 (Mo.; N.Y.); Degener,
Salucop, & Arlantico 11,549, tree 7 ft. tall, near summit, C.C.C.
Trail, Aiea, Dec. 6, 1937 (Bish. ; petals 5, drupes not seen) ; Degener,
Salucop, & Arlantico 11,919 (type collection, Bish., 2 sheets; Corn.,
my photograph no. 4,328; Del.; Mo.; U.S.); F. Fagerlind & C.
Skottsberg 6,085, Pupukea forest reserve, Koolau Range, Jan. 25,
1948 (Stockh.; leaflets truncate to somewhat subcordate); H.
Hapeman, Oahu, Apr. 14, 1908 (Mo.); E. Y. Hosaka 301, tree 35
ft. tall, along wooded stream-bed, alt. 1,000 ft., Waikakalaua Gulch,
Sept. 14, 1930 (Bish.) ; Hosaka 688, tree 25 ft. tall, on denuded slope,
alt. 2,500 ft., south ridge, Kipapa Gulch, Waipio, Sept. 18, 1932
(Bish., stigmas mostly 3, rarely 4); H. Morley, small tree 12-15
ft. tall, in rain-forest, alt. 2,500 ft., ridge south of Kipapa Gulch,
Waipio, Dec. 10, 1933 (U.S., petals 5, stigmas 4 or imperfectly 5) ;
Carl Skottsberg 1,814, alt. about 300 m., Pupukea Forest Reserve,
Sept. 15, 1926 (Goth.).

Seemingly this is a good variety, but its recognition is at times
more difficult than that of most varieties, certainly of the other
varieties of T. gymnocarpa.

19. Tetraplasandra kahanana Deg. & Sherff ex Sherff, Bot.
Leafl. no. 8: 6. 1953.

Entirely glabrous, without doubt a tree; leaf-scars on the branch^-
lets finally large and conspicuous. Leaves 4 dm. long including
the slender petiole (this 1 dm. long and 3-4 mm. thick), 7- or
9-foliolate; leaflets dull, weakly coriaceous and subobscurely veiny,
narrowly to moderately oblong or oblong-ovate or (especially the
terminal three) obovate; their blade 8-12.5 cm. long and 3-6 cm.
wide, at apex subobtuse to subacute, at base scarcely oblique and
broadly to rather narrowly cuneate; petiolules slender, lateral ones
10-15 mm. long. Inflorescence (a single one seen) 3 dm. long,
its 3 peduncles umbellately grouped, curved-erect, about 2-2.7 dm.
long, remotely and alternately branched or radiate from near base
to top; rays slender, curved-suberect, a few (5) placed along the
peduncles and still more (7) at the top, 6-10 cm. long, terminally
and umbellulately about 7-10-pedicellate; pedicels slender, at anthe-
sis straightish to curved and erect to spreading, commonly 12-17
mm. long (rarely one of them transformed into a subray 3 cm. long
and terminally subdivided into a few smaller pedicels). Calyx at
anthesis obconic, 3.5-4 mm. tall and almost as broad. Corolla

138 FIELDIANA: BOTANY, VOLUME 29

before opening scarcely 4.5 mm. tall and 6 mm. thick, petals 5 or
6, a lone mature one (seen in packet on type sheet) lanceolate,
reflexed, 6 mm. long. Stamens 5, filaments flat and scarcely 2.5
mm. long; anthers ovate, about 1.5 mm. long, attached below their
center. Drupes (only very immature ones seen) obovoid; ovary
inferior as to its lower half, its upper part depressed-hemispherical;
stylopodium central, barely conical but several-rayed, stigmas not
yet well-formed.

Type: Otto Degener 18,013, in forest, from Kahana church up
ridge to summit of mountain southeast of Kahana Bay, Isl. Oahu,
July 3, 1932 (N.Y.).

Distribution: Known only from type locality in northeastern
Oahu.

Specimens examined: Degener 18,013 (type, N.Y., my photo-
graph no. 4,344: isotype, N.Y., my photograph no. 4,345).

The five stamens place this species next to T. meiandra, but the
racemosely (as well as at the tip umbellately) branched peduncles
give an appearance to the inflorescence suggestive of T. hawaiiensis,
T. kauaiensis and T. gymnocarpa. (In T. meiandra var. meiandra a
single lateral branch or ray is very rarely emitted from a peduncle,
somewhere near the top.) A plate was prepared some years ago
by Kwan Kee Park from the type material of T. kahanana but
seems never to have been published.

INDEX OF EXSICCATAE

Collectors' names and page numbers in italic type

Baxter, R. W., Pupukea-Kahuku Trail,

13^
Bryan, Edwin H., Jr., Pupukea, 134

no. 618, 68

no. 878, 105

Cowan, Richard S., no. 30, 109

no. 504, 109

no. 557, 104

no. 563, 90

no. 711, 109

no. 745, 109

no. 963, 109

no. 980, 104

Cranwell, Miss Lucy May, no. 3528, 121
Cranwell, Miss Lucy May, Olof Hugo
Selling, & Carl Skottsberg, no. 2,675,
113

no. 2,695, 113

no. 2,925, 76, 77

no. 3,095, 128

no. 3,151, 83, 84

no. 3,156 (sphalm), 83

Degener, Otto, between Honomanu
Valley and Keanae, Oahu, 68, 79

gully s.e. of Kahuku entrance of
Pupukea-Kahuku Trail, 79

east part of Kaluaaha plateau, 68

H mile north of Keahikauo, 69

Keanae Valley, 68

slope n.e. of Nuuanu Valley, 89

west of Pepeopae, 115

s.e. of Pepeopae Bog, 115

Pig-God Trail, 97, 109

Pipe-Line Trail, Olinda, 115

Pupukea-Kahuku region, Mar. 31,
1929, 134

Pupukea-Kahuku, July 19, 1931, 79

Wahiawa, 89

no. 2,181, 66

no. 2,315, 114

no. 2,319, 69

no. 2,921, 79

no. 2,968, 95

no. 11,318, 98

no. 17,785, 116

no. 17,786, 116

no. 17,787, 68

no. 17,788, pro parte, 68, 115

no. 17,790, 119

no. 17,791 pro parte, 81, 109

no. 17,792 pro parte, 79, 80, 81

no. 17,793, 115

no. 17,795, 115

no. 17,796, 89

no. 17,971, 99

no. 17,972, 104

no. 17,973, 98

no. 17,974, 81

no. 17,976, 95

no. 17,977, 109

no. 17,978, 104

no. 18,013, 138

no. 18,014, 100

no. 18,015 pro parte, 97, 136

no. 18,016, 97

no. 18,017, 68

no. 18,018, 115

no. 18,019, 115

no. 18,021, 106

no. 18,242, 97

no. 18,243, 109

no. 18,244, 88

no. 18,245, 100

no. 18,246, 100

no. 18,247, 97

no. 18,248, SO

no. 18,249 pro parte, 81, 109

no. 18,250 pro parte, 79, 109

no. 18,252, 119

no. 18,253, 137

no. 21,016, 100

no. 21,473, 76

no. 21,759, 60

no. 21,787, 119

no. 21,788, 66

no. 21,790, 123

no. 21,834, 66

Degener, Otto, & Miss Amy Greenwell,
no. 21,015, 100

no. 21,577, 70

no. 21,578, 125

no. 21,579, 125

no. 21,725, 76

no. 21,730, 73

no. 21,898, 67

Degener, Greenwell, & William Hath-
eway, no. 20,682, 101

no. 21,142, 67
Degener & Hatheway, no. 20,865, 131

139

140

FIELDIANA: BOTANY, VOLUME 29

Degener & Martin Martinez, no. 11,920,

13
Degener, Toshio Murashige, & Mark

Kerr, no. 21,017, 80
no. 21,018, 101
Degener & Clarence Nihei, no, 20,222,

109
Degener & Emilio Ordonez, no. 12,066,

79
Degener, Ordonez, & Olof Hugo Selling,

no. 12,236, 101
Degener, Park, Bush, Potter, & Topping,

no. 11,921, 135
Degener, Salucop, & Arlantico, no.

11,549, 137
no. 11,820, lOlt
no. 11,919, 136, 137
Degener, Otto Swezey, Kwan K. Park,

& Kazuto Nitta, no. 6,099, 88
Degener, Tarn, Tousley, & Barber,

22,025, 130

Degener & Henry Wiebke, no. 2,180, 129
no. 2,181, 66
no. 2,183, 76
no. 2,184, 119
no. 2,185, 69
no. 2,316, 69
no. 2,317, 115

Ewart, G. R. Ill, no. 100, 118
Ewart, G. R. Ill, & George C. Munro,
no. 129, 118

Fagerlind, F., & Carl Skottsberg, no.

6085, 137

no. 6,298, 119

no. 6,587, 76

no. 6,608, 70

Fagerlund, G. 0., & A. L. Mitchell, no.

799, 66
no. 945, 117

Faurie, Abbe Urbain, no. 269, 68
no. 270 pro parte, 95, 96, 104.
no. 271, 115
no. 273, 10k
Forbes, Charles Noyes, Konahuanui, 92,

10k

Makaha Valley, 131
Mountains between Punaluu and

Kaipapau, 109
Waiolani Ridge, 10k
(with C. M. Cooke), mts. between

Punaluu and Kaipapau, 13k
no. 104-L, 107
no. 108-M, 69
no. 141-L, 69
no. 164-K, 82
no. 199-L, 107
no. 200-K, 75
no. 205-K, 76
no. 239-M, 128
no. 274-K, 75

no. 276-K, 129

no. 286-K, 82

no. 307-L, 8k

no. 377-L, 69

no. 386-L, 84.

no. 388-L, 8k

no. 422-M, Ilk

no. 430-K, 125

no. 432-K, 70

no. 508-Mo, 68

no. 562-Mo, 121

no. 719-H, 108

no. 1,034-O, 10^

no. 1,064-K, 70

no. 1,097-K, 125

no. 1,608-0, 135

no. 1,638-0, 13k

no. 1,674-0, 99

no. 1,713-M, 112

no. 1,726-M, 112

no. 1,930-M, 108

no. 1,942-O, 88

no. 1,973-M, 127

no. 2,051-M, 112

no. 2,062-O, 13k

no. 2,169-M, 127

no. 2.182-O, 13k

no. 2,210-O, 109

no. 2,215-O, 109

no. 2,276-M, 11 k

no. 2,313-M, Ilk

no. 2,511-0, 87
Forbes (with J. C. Bridwell), no. 2,457-0,

13k
Forbes (withJ. C. Bridwell), no. 2,461-O,

88
Forbes (with Mrs. G. E. Kelly), no.

2,390-O, 92

Forbes (with Dean Lake), no. 1,973-0,
95

Garber, D. Wesley, no. 238, 13k
no. 245, 92

Hapeman, H., Oahu, 137
Hatheway, William, no. 209, 111
Heller, Amos Arthur, no. 2,734 pro
parte, 72, 75

no. 2,884, 128
Hillebrand, William, anno 1870, 126

East Maui, 127

Halemanu, 70, 71

Kaala, 101, 132

Mt. Kaala, alt. 4,000 ft., 100

Mt. Kaala in 1867, 9k, 101

Kaala Mts., 132

Kanapali, 69

Kawaihae iuka, 105

Konahuanui, 9k

Lanai, 1870, 107

Makaleha, Kaala, 132

Maunahui, 116

INDEX

141

Molokai, 68

Niu, 93, 13 It

Nuuanu, 104

Nuuanu, 1867, 94

Waimea in 1871, 70

no. 264, 105

no. 266, 132

Hillebrand, William, & Rev. John
Lydgate, Niu, 109, 131+

Wailupe, 87

Hillebrand (with Wawrd), Nuuanu, 104
Hitchcock, Albert Spear, no. 14,824, 113

no. 15,481, 75

no. 15,550, 70

Hochreutiner, B.P.G., no. 3,538, 71
Hosaka, Edward Yataro, no. Ill, 134

no. 124, 78

no. 301, 137

no. 613, 110

no. 688, 137

no. 956 pro parte, 93, 110
Hume, Edward P., no. 165, 96, 97

Lydgate, Rev. John M., Hii Mountains,

102, 103

Woods of Hilo, 107
Valley of Niu, 13 4
Wahiawa Mountains, 76
Wailupe, 87

MacCaughey, Vaughan, Kaena, 104
MacDaniels, Laurence Howland, no. 62,
104

no. 114, 134

no. 327, 69

no. 466, 88

no. 535, 134

no. 800, 129

no. 811, 76

no. 847, 82, 83

Mann, Horace, Jr., & William T. Brig-
ham, Konahuanui, 104

no. 349 pro parte, 51, 125

no. 378, 69

no. 606 pro parte, 51, 125, 126
Meebold, Alfred, Pupukea, 134
Morley, Harold, ridge south of Kipapa

Gulch, 137
Munro, George C., Kaiholena, 84

Lanai, 107

Lanai, Aug. 31, 1915, 84

Lanai, Nov. 17, 1915, 126

Poomai, 107

Valley south of Puhielelu, 126

no. 39, 69

no. 140, 69

no. 199, 107

no. 335, 84

no. 401, 112

no. 461, 84

no. 469, 126

no. 694, 85

Nitta, Kazuto, no. 73, 134
no. 76, 78

Ordonez, Emilio, (Degener distrib.) no.
12,988, 107

Rock, Joseph Francis, below Kaholua-

manu, 125
Kaluaaha, 68
Kapua, 66

Kauai, October, 1916, 70, 75
Kilauea Volcano, 119
Near 29 Miles Forest, 119
Nahiku, Jan. 14, 1909, 128
Nahiku, May, 1911, 128
Nahiku, May 10, 1911, 128
Niu Valley, 1911, 93
Punaluu Mountains, Aug. 23, 1908,

104
Punaluu Mts., main ridge of, Nov.

14-21, 1908, 110
Puuwaawaa, 117
no. 188, 110
no. 191, 110
no. 406, 135
no. 406-a, 134
no. 421, 135
no. 449, 135
no. 1,307, 110
no. 1,720, 70
no. 1,744, 70
no. 2,488, 125
no. 2,488a, 125
no. 3,855, 117
no. 3,857, 117
no. 3,858, 117
no. 4,831, 93
no. 4,901, 75

no. 4,902 pro parte, 73, 74, 75
no. 5,162, 70
no. 5,165, 70
no. 5,167, 70
no. 5,168, 70
no. 5,913, 70
no. 6,175, 115
no. 7,006, 68
no. 8,015, 107
no. 8,016, 69

no. 8,016a, July, 1910, 69
no. 8,016a, August, 1910, 69
no. 8,088, 84
no. 8,207, 114
no. 8,386, 67
no. 8,429, 121
no. 8,582, 114, 115
no. 8,641, 127
no. 8,641a, 128
no. 8,666, 112
no. 8,722, 104
no. 8,826, 110
no. 8,832, 110
no. 8,870, 73, 74

142

FIELDIANA: BOTANY, VOLUME 29

no. 10,006, 120

no. 10,019, 126

no. 10,026, Jan. 28, 1912, 66

no. 10,026, February, 1912, 66

no. 10,070, 115

no. 10,362, 122

no. 12,506, 112

no. 12,508, 116

no. 12,509, 117

no. 12,510, 128

no. 12,993, 118, 119

no. 16,009, Ilk

no. 17,053, m

no. 17,155, 111
Rock & Charles Noyes Forbes, Punaluu

Mountains, 110
Rock & Hammond, no. 8,015, 107

no. 8,050, 69

Rock & Hashimoto, no. 16,033, 67
Rock & Marshall, borders of meadow

called Malua Poha, 125
Rock & Shaw, Olympus, 104.
Rock & Wilder, no. 8,827, 135
Rogers, D. P., head of Waiawa Valley,

134
Russ, G. W., Ahaino, 68

Makua, 102

Waimano Gulch, 96

Saint John, Harold, no. 10,104, 134
no. 10,147, 134
no. 10,579, 110
no. 20,215, 104
no. 10,231, 114

Saint John, Baker, Coulter, Fosberg, &

Yuncker, no. 12,846, 68
Selling, Olof Hugo, no. 3,338, 101

no. 3,631, 110

no. 3,671, 136

no. 3,879, 113
Skottsberg, Carl, no. 459, 66

no. 781, 113

no. 785, 113

no. 878, 116

no. 1,009, 70

no. 1,010, 129

no. 1,125, 126

no. 1,814, 137

no. 1,816, 78

no. 1,862, 110

Suehiro, Miss Amy, Waikane-Schofield
Trail, Oct. 16, 1932, 110

Kaipapau, Oct. 15, 1933, 110

Topping, David LeRoy, no. 2,921, 79

no. 2,968, 104.
Topping & William Bush, no. 3,701, 88

United States South Pacif. Explor. Ex-

ped., mts. behind Honolulu, 92
District of Puna, 66

Wawra, Heinrich, no. 1,638, 103

no. 2,114, 70

Webster, Grady L., no. 1,443, 98
Wilbur, R. L., & G. L. Webster, no. 835,

114

no. 863, 114
no. 890, 114

Publication 749

UNIVERSITY OF ILLINOIS-URBANA

30112041653715