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 UBRARV 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 T1. 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. Lean1, 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. T1. 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. Lean1,

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. Lean1,

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.

Lean1, 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. Lean1,

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