SMITHSONIAN CONTRIBUTIONS TO BOTANY NUMBER 45 

Systematic Studies of 

Micronesian Plants 

F. Raymond Fosberg 

and Marie-Helene Sachet 

SMITHSONIAN INSTITUTION PRESS 

City of Washington 

1980 
ABSTRACT 

Fosberg, F. Kaymond, and Slarie-Helene Sachet. Systematic Studies of Micronesian Plants. Smithsoninn Contiibutions to Botany, number 45, 40 pages, 2 figures, 1980.-Taxonomic and nomenclatural treatments of various genera in a 

number of families known from Micronesia, with new species, varieties, and 

combinations. A discussion is included of the systematic position of the genus 

Fagmea Thunberg, transferring it from the family Loganiaceae to the Gentianaceae. The genus TYollastonia de Candolle (Compositae) is resurrected from the 

synonymy of Wedelia Jacquin, and the widespread species commonly called 

Wetlelia biflo~a (L.) de Candolle is discussed in relation to the Hawaiian genus 

Lipochaeta de Candolle and is moved to Wollastorzia. 

OFFICIAL 

recorded 

from the 

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

in the Institution's annual report, Smithoninn I'enr. SERIES CO~ER DESIGX: Leaf clearing 

Katsura tree Cercidiplzyllum jnponicum Siebold and Zuccarini. 

Library of Congress Cataloging in Publication Data 

Fosberg, Francis Raymond, 1908- 

S\ atematic studies of Micronesian plants. 

(Smithsonian contributions to botany ; no. 45) 

Bibliography: p. 

Includes index. 

I. BotanL-Micronesia-Classification. 2. Botany-Classification. I. Sachet, Marie-Helene joint 

author. 11. Title. 111. Series: Smithsonian Institution. Smithsonian contributions to botany : 

no. 45. 

QK1 .S2747 no. 45 [QK473.M5] 581'.08s [582'.09'965] i9-16001 
Contents 

Page 

1 

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4 

4 

5 

5 

5 

5 

6 

6 

7 

12 

12 

14 

15 

15 

15 

15 

17 

17 

Introduction ...................................................... HYMENOPHYLLACEAE .... ............... GLEICHEKIACEAE ............. ............... POLYPODIACEAE, sensu lato ...................................... GNETACEAE ........................... ............. 

HYDROCHARITACEAE ................................................ 

ARACEAE .... ............................... 

............................. 

........................... ........................ ........ .................................... 

CUCURBITACEAE .................. ............. 

RHIZOPHORACEAE .................................... 

........................... ......................... 

SAPOTACEAE ............................. GENTIAN ACEAE .......................... ........... 

LOGANIACEAE ..... ........................... BORAGINACEAE .... ........................... 

..................................... 

23 

25 

26 

27 

30 

35 

37 

iii 

Systematic Studies of Micronesian Plants 

F. Raymond Fosberg 

and Marie-Helene Sachet 

Introduction 

In the course of work on the Micronesian flora 

we have encountered many taxonomic and nomenclatural problems. Some of our conclusions have 

been published as small papers, others have accumulated, unpublished. The names and observations 

which are ready for publication are here made 

available in advance of their use in the parts of the 

Flora, which appear at irregular intervals. l'hus the 

correct names may be used and referred to by ourselves and others working on the plants of the 

region. 

This paper includes observations on the families 

Hymenophyllaceae, genera Hymenophyllurn and 

Trichomnnes; Gleicheniaceae, Gleichenia; Polypodiaceae, Thetypteris; Gnetaceae, Gnetum; Hydrocharitaceae, Hydrilla; Gramineae, Centosteca; 

Araceae, Rhaphidophora; Urticaceae, Elatostenia; 

Leguminosae, Callinndra and Stylosanthes; Eupliorbiaceae, Acalypha; Celastraceae, Loeseneriella; 

Clusiaceae, Cnlophyllum and Gnrcinin; Flacourtiaceae, Flacourtia; Cucurbitaceae, Citrullus and Zehneria; Rhizophoracene, Brugziiera; Melastornataceae, Medinilla and Melastoma; Araliaceae, 

Osnioxylon; Xlyrsinaceae, Myrsine; Sapotaceae, 

Pouterin; Gentianaceae, Fagraea; Loganiaceae, 

Geniostomn; Boraginaceae, Heliotropium; Verbenaceae, Callicnrpa; Acanthaceae, Pseuderanthemum; Rubiaceae, Geophila, Hedyotis, Psychotria, 

and Spernzacoce; Compositae, Vernonia, Wollastonin. Selected specimens are cited to support the 

observations; in some cases they may again be cited 

later in the flora itself. 

F. Raymond Fosberg and Marie-Helene Sachet, Department of Botany, National Museum of h'atural History, 

Smithsonian Zn.rtitution, Washington, D.C. 20560. 

For details of herbarium symbols, the history, circumstances, and format of the Flora of Micronesia, 

see the first three papers in the series, Smilhsonian 

Contributions to Botany, numbers 20, 24, and 36. 

i\'e are much indebted to those Ivho have sup. 

plied specimens and field ob,ervations, especially 

Mrs, hlarjorie Falanruw of Yap Island, Aliss Joan 

Canfield in Palau, and Rlr. Philip ;\loore in Guam. 

51r. Royce Oliver has given inv:iluable help in locating specimens and pointing out pertinent details. 

Miss Dulcie Powell has examined much of the 

manuscript. Dr. Harold Robinson has made valuable suggestions concerning our treatment of 

IYollastoniu. l\re are much indebted to the authorities of the various herbaria, froin which specimens 

are cited, for their courtesy in allowing us to work 

with their collections and to borrow specimens 

when necessary. The Flora Malesiana has been of 

tremendous assistance in working out many taxonomic and nomenclatural details. Grants from the 

Smithsonian Research Awards Program to the 

Smi thsonian Research Foundal.ion have supported 

our work on island floras, including that reported 

in this paper, for a number of years. IVe express our 

thanks to all of the above. 

A map of Micronesia, showing the locations of 

the islands mentioned is included as Figure 1. 

HYMENOPHYLLACEAE 

Hymenophyllum Smith 

Hymenophyllum digitaturn (Swartz) Fosberg, 

new combination 

Trichomanes digitaturn Swartz, Syn., 370, 422, 1806. 

Microtriclzonranes digitaturn (Swart7) Copeland, Phil. Jour. 

Sci., 67:1, 36, 1938. 

1 
I300 135' I400 145' I 50° 

'30' 

9'. 

b 

RYUKYU 0 

0 loo 200 300 400 500 Statute Miles I I I I I 1 

250 

200 

15' 

PHIL 

BON IN 

3Okinawa -jimo Haha-jimd I SLA N 0s 

ISLANDS I' Da i t6 - j i ma Ki to-io-jlma. 

CI / \ 

lwo Jima, VOLCANO ISLANDS 

I 

I-- Ma rcusq 

-- -Minorni-iB- jima 

/ 

/ 

/ 

/ 

+- a 

c / 

/ 

/ 

1 . Parece Vela 

I 

I 

I 

I 

/ 

I 

PPlNE I 

I SL ANDS 

: Pmor 

t 

I 

I 

I 

I . ,... U I I t h i 

Fais YARi ' ' ' 

Mindonoo 

/oj' 

5O 

0 e 

'Urocar 

+4aug 

Asuncion 

*Pagan 

VGuguan 

T Sa r igan 

Agrigon MAR I AN AS 

Al amaganb 

I SLANOS 

Anatahan' .Med i ni I lo 

Saipon 

Agiguon 

Tin i a n3g 

'Rota 

#Guam 

Santa Rosa Reef 

Gaferut ' Namonuito ... E.Fpyu . Murilo 

'lap W. Foyu. P!kelotPula Nomwin 

olinarao. ... Lomotrek , PRUK. 

Kuop- Noma 

Lukunor 

*, 

' I s L A NDS) Ba b e Id a o b Woleal .. . Elat 

I fa 1 uk %a tawat ~uluwat Eauripik Losap / Angaur. PellliU 

P'lusuk Namoluk ..5!al 

Satawdn Sonsorol 

Pulo Anna 

Mer I r CAROLINE ISLANDS 

I 

I 

\ 

.Helen Reef 

Ha Ima hers\ 

---__ 

NEW G UlNEA 

5O 
I550 160' 165' 170' I750 

FIGURE 1 .-Micronesia. 

~Pokak 

't Kopingamarangi 

I 

I 

Abaiang: iarawa Marakei 

ISLANDS -' M~,~~~ 

ur ia.. A bemama 

0, r anu ka I 

I 

.Nauru 

-- onotoat Nikunaul 

---_ Tarnono. ,Aroroi , 

Banaba- Nonouti",, Beru 

- (Oc .an1 Tabiteuea%h ' 9 

--- -z -z -- -- , SOLOMON ELLICE 

ISLANDS 

165' 170' 1750'' 1 I I 1 

180' 

3 0' 

25' 

2 oo 

15' 

I00 

50 

O0 

50 

180' 
4 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

In spite of their united involucres, transferring 

the species 1 eferred by Copeland to Alicl-otrichomanes Copeland, from Trichonzanes to Hymenoplzyllurn, removes some of the artificiality from the 

circumscription of the traditional genera of the 

H) menophyllaceae. This weakens, somewhat, the 

argument for dismemberment of these large genera. 

I am unable to find that the appropriate combination has been made for this species. 

Trichomanes L. 

Trichomanes falsinervulosum (Nishida) 

Fosberg, new combination 

Microgoniunz /alsinervulosurn h’ishida, Jour. Jap. Bat., 

32(5):156, fig. 4, 1957. 

This species resembles TI ichomanes craspedoneunivz Copeland, but differs in many details, as 

brought out in the original description. However, 

the dimensions given for the frond do not conespond \\.ell to the shape illustrated. I have not seen 

the type, Hosokawa s.ii,, in Taihoku University 

Herbarium. 

Trichomanes tahitense Nadeaud 

Trichomanes tahiteizse Nadeaud, Enum P1. Tahiti, 18, 1873. 

Microgoniuni tahitense (Nadeaud) Tindale, Contr. N.S.W. 

Ti ichomanes omphalodes C. Christensen, Ind. Fil., 646, 

Nat. Herb, Flora Ser., 201:4, 33-34, 1979. 

1906. 

Dr. Tindale points out that the species coninionl) 

known as T. on~phalodes C. Christensen is identical 

with the older T. tahitense Nadeaud. In Micronesia 

it is known from Palau and Ponape, Tvhere it is an 

epiph) te on tree trunks in Tvet forests. 

GLEICHEKIACEAE 

Gleichenia J. E. Smith 

Glekhenia linearis var. latiloba (Holttum) 

Fosberg, new combination 

Dicranopteris linearis var, lntiloba Holttum, Reinwardtia, 

43277, 1957. 

The several characters used by Holttum (1957a: 

258-259; 1957b: 168-184) to separate Dicran0pteri.s 

Bernhardi from Gleichnia Smith do not seem adequate to justify more than sectional rank. Therefore, this variety must be transferred if it is to be 

maintained. 

POLYPODIACEAE, sensu lato 

Thelypteris Schmidel 

This genus has been drastically divided by 

Holttum (1971, 19i7), but we find it very difficult 

to follow most of this segregation and think it more 

convenient, and perhaps even more natural to follow Morton (1963) in regarding the “family” 

Thel) pteridaceae as principally comprising one 

large genus. Names are available in Thelypteris 

for most Alicronesian species, but we find none for 

the following three, for which a new combination 

and two new names are proposed. 

In addition, Cycloro~us jaculosus (Christ) Ito has 

been reported from Truk. Neither Thelypteriy 

jaculosus (Christ) Panigrahi nor Thelypteris ~ubpubescens (Blume) Iwatsuki, which have been confused, are admitted from Micronesia by Holttum. 

Disposition of the Truk record will have to wait 

until we can examine the specimen, collected by 

Kusano in 1915, on which it is based. 

Thelypteris carolinensis (Hosokawa) 

Fosberg, new combination 

Dryopteris carolinensis Hosokawa, Trans. h’at. Hist. SOC. 

Christella carolinensis (Hosokawa) Holttum, Kew Bull., 

Formosa 26:74, 1936. 

31:307, 1976. 

Thelypteris peliliuensis Fosberg, new name 

Glaphyropteris palauensis Hosokawa, Trans. h’at. Hist. Sac. 

Formosa, 323285-286, 1942. 

Dryopteyis irninersa sensu Hosokawa, Trans. Nat. Hist. SOC. 

Formosa, 28:147, 1938 [non (Blume) 0. Kuntze, Rev. Gen. 

Pl., 2:813, 18911. 

Hosokawa’s description is rather full but we are 

unable, from it, to place the plant in any of Holttuin’s genera, nor has Holttum accounted for 

Hosokawa’s species. TVe have not seen the type, 

Hosokawa’s 9221, from Peliliu, Palau. Holttum 

(1971, 1977) seeins to restrict the genus Glnphgroptrri.s (Presl ex Fee) Fee to the Kew TVorld. Therefore, even if it were maintained as a genus, it would 
NUMBER 45 5 

scarcel) be an appropriate place for the species 

under consideration. Because of the alre‘idy existing 

Thelypterzy palauen,i, (Hosokawa) Reed (1968) the 

epithet palazrenszs cannot be transferred to Thelypte?is. Hence a new name is proposed, based on 

Hosokawa’s description and t) pe-specimen. 

Thelypteris rupi-insularis Fosberg, new name 

Dryopteris rupicola Hosokawa, Trans. Nat. Hist. SOC. 

Formosa, 26:73, 1936 [non C. Christensen 1917 = 

Thelypteris rupicola (C. Christensen) Ching 19171. 

Glapliyropteris rupicola (Hosokawa) Hosokawa, Trans. Nat. 

Hist. Soc. Formosa, 323285, 1942. 

Christella rupicola (Hosokawa) Holttum, Allertonia, 1:217, 

1977. 

GNETACEAE 

Gnetum L. 

Gneturn gnctnon sensu Fosberg Pi Sachet, Smithsonian Contr. 

Bot., 20:13, 197.5 [non L., Mantissa, 1:125, 17671. 

The two sterile specimens from Palau, Hosokawcc 

906I and Stone and Sabino 4541 (both BISH), referred to Gncticm gncinon L. by LIS in 1975 (following uncritically the determination on the label of 

no. 9061), when compared with fertile specimens of 

Plzaleria ni.ridai Kanehira (Thymeleaceae), seem undoubtedly to belong to the latter species. Since 

these are the only basis of reports of Gneticm in 

Microne, .I of which we know, the family Gnetaceae, ge,.us Gnetum, and species Gnetum g~emon 

should be deleted from the flora of Micronesia. 

HYDROCHARITACEAE 

Hydrilla Richard 

Hydrilla verticillata (L.f.) Royle 

HydriZZa uerticilata (L.f.) Royle, Ill. Bot. Himal., T. 376, 

Serpicula uerticillata L.f. Suppl., 416, 1781. 

Egeria densa sensu Fosberg, Phytologia, 15:496, 1968 [non 

In 1968 I (Fosberg) reported Egeria densa 

Planchon from Guam on the basis of Stone 4305, 

from the Talofofo River near the mouth. The 

Bishop liuseum sheet of this collection was determined by Harold St. ,John as Egeria densa. In his 

Flora of Guam, Stone (1971: 101-102) refers this 

numlier to both Egerin and Hydrilla, which caused 

1839. 

Planchon, Ann. Sci. Nat. Bot. 3,11:80, 18491. 

me to check the US sheet. This turned out to be 

Hydrilla, on the basis of the prominent, rather than 

exceedingly minute, spinulose teeth on the leaf 

margin. The collection is sterile, as is so often the 

case with both these aquatics. Since this is the orlly 

report of Egel-ia from Guam, this genus should be 

deleted from the flora of Guam. 

GRAhiINEAE (= POACEAE) 

Centosteca Desvaux (Centotheca Heauvois, 

orth. mut.) 

Centosteca lappacea (L.) Desvaux 

Centosteca lappacea (L.) Desvaux, Nouv. Bull. SOC. Philom. 

Paris, 2:189, 1810; Jour. de Bot. Desvaux, 1:71, 1813.- 

Beauvois, Essai . . . Agrost., 69, 1812.-Dandy in Exell. 

Cat. T’asc. P1. S. Thome, 365, 1944. 

Cenchrus Zappaceus L., Sp. Pl., ed. 2, 1488, 1763. 

HoZcus latifolius Osbeck, Dagb. Ostincl. Resa, 247, 1757. 

Centotheca [atifolia Trinius, Fund. Agrost., 141, 1820 (nom. 

superfl. illegit.).-Monod de Froideville, Blumea, 19:58, 

60, 1971. 

For the benefit of any who may have missed 

Dandy’s 1944 discussion of the nomenclature of this 

widespread grass species, and who may have read 

the recent synopsis of the genus Centosteca (as 

Centotheca) by Nonod de Froitleville (1971:57-60), 

it seems worth while to point out that IIonotl 

apparently also missed Dandy’s discussion. Dandy 

showed that Trinius based his Centothcca lntifolia 

on Cenclirus lappaceus L., not on Holcus Iatifolia 

Osbeck; therefore, any transfer of the latter to 

Centosteca will create a later 1iomon)m. Hence the 

name Cen tosteca lappacea (L.) Desvaux must stand. 

It should be pointed out that Desvaux’s original 

spelling of the generic name w.\‘;Is Cento.rteca which 

was changed (“corrected”) by Beauvois: this change 

does not seem to be warranted according to the 

International Code, even though it wa~ later accepted by Desvaux in 1813 (Soderstrom and Decker, 

1973:433). 

ARACEAE 

Rhaphidophora Hasskarl 

Rhaphidophora carolinensis (Volkens) 

Fosberg, new combination 

Epiprenzrztcm carolinen~c \.olkens. Bot. Tahrb.. 31:459, 1901 
6 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Epipremnuni palauense Koidzumi, Bot. Mag. (Tokyo), 30:401, 

Rhaphidophora koidzumii Kanehira, Enum. Micr. PI., 286, 

1916. 

1935. 

When Rhaphidophora Hasskarl, 1842, and Epipremnum Schott, 1857, are united under the name 

Rhaphidophora, the name for this species must be 

based on the earlier E. carolinense Volkens. Kanehira, though he did not unite the two genera in 

his Enumeration, did transfer E. palauense Koidzumi to Rhaphidophora, renaming it e. koidzumii 

because of the earlier Rhaphidophora palauensis 

Koidzumi (1916). However, the still earlier epithet 

of Epipremnum carolinense Volkens must prevail. 

URTICACEAE 

Elatostema Forster 

Elatostema Forster, Char. Gen. 105, tab. 53, 1775.- 

Gaudichaud, Bot. Freyc. Voy. Uranie, 493, 1826 [1830]. 

Pellionia Gaudichaud, Bot. Freyc. Voy. Uranie, 494, 1826 

[1830], [pro parte]. 

Langveldia Gaudichaud, Bot. Freyc. Voy. Uranie, 494, 

1826 [1830]. 

This genus, typified by Elatostema sessile Forster 

and taken in the broad sense here adopted, is large 

and taxonomically difficult, and there is no adequate treatment of it. 

A number of species have been described from 

Micronesia, some represented only by their type 

collections. Superficially, they are mostly rather 

similar, but have not been studied in detail, partly 

from lack of adequate material. In the Marianas, 

the oldest name available, Pellionia divaricata, was 

provided by Gaudichaud for a plant from Guam. 

hferrill (1914:76, 77) described two further species 

from Guam, Elatostema stenophyllum and E. calcareum, but made no reference to Gaudichaud’s 

species; yet the original material of Pellionia 

divaricata seen in Paris seems very similar to ElatoAtenza calcaieunz Merrill. However, in our opinion 

the reduction should not be made until all available 

material can be assembled and compared critically 

in minute as well as gross features. So far as present 

concepts go, the genus is noted for its high degree 

of local endemism, at least in insular floras. For the 

present it seems best to accept all three Marianas 

species. 

Elatostema divaricatum (Gaudichaud) 

Fosberg, new combination 

Pellionia divaricata Gaudichaud, Bot. Freyc. Voy. Uranie, 

Procrts diva? tcata (Gaudichaud) Steudel, Som., ed. 2, 2:398, 

494, 1826 [1830]. 

1841 [“In insulis Mariannis (Guam)”]. 

For Gaudichaud’s species, it is necessary to provide this new combination. This name being oldest, 

may eventually replace one or both of the two other 

names for the Marianas species. 

LEGUMINOSAE (= FABACEAE) 

Calliandra Bentham 

Calliandra haematocephala Hasskarl 

Calliandra haematocephala Hasskarl, Retzia, 1 :216, 1855. 

Calliandra inaequilatera Rusby, Mem. Torrey Bot. Club, 

6:28, 1896. 

This attractive ornamental is planted in many 

tropical countries, but has apparently not been reported before from Micronesia. Calliandra haematocephala and C. inaequilatera have usually been 

treated as separate species: R.S. Cowan (1963: 94-98) 

has given characters purporting to separate them. 

However, the cultivated Micronesian specimens 

seem to fall between the two by his criteria. 

Nevling and Elias (1 97 1 : 69-85) convincingly 

demonstrated that two species are not represented 

and that the material on which the earlier name 

was based was from cultivated plants in Java that 

originated from plants brought from the same region in Bolivia from which Rusby brought the type 

of Calliandra inaequilatera. 

This species should not be confused with Calliandra haematomma [sic] (Bertero ex de Candolle) 

Bentham, an entirely different species, though the 

name is sometimes misapplied. 

SPEcibiEX’s Stt~.-Caroline Islands: Palau; Koror, Entomology Laboratory grounds, Salsedo IIP(US). Ponape: 

Colonia, 100-150 ft [3MO m], Salomon and George B(US). 

Stylosanthes Swartz 

Two species of Stylosanthes have been reported 

from Micronesia, both by Alan Burges (ca. 1935) 

from Nauru. \\’e have not seen specimens of either. 

They were reported as Stylosanthes sundaicus 
NUMBER 45 7 

Taubert and Stylosanthes guyannensis (Aublet) 

Swartz. 

Mohlenbrock (1957:345-346) equates Stylosanthes 

sundaicus with Stylosanthes humilis Humboldt, 

Bonpland & Kunth. Since we have seen no specimen, we can only accept this disposition, and use 

Mohlenbrock’s key characters to separate the two 

Micronesian recorded species in the key given 

below. 

Mohlenbrock used the orthography guyanensis for 

the other species. Neither spelling seemed likely 

considering that Aublet was French-speaking. Reference to the original texts showed that Aublet (1775: 

776) used the binomial Trifolium guianense and 

that Swartz (1789:301) transferred it in that form to 

his new genus Stylosanthes. On Aublet’s plate, 

T.309, however, the name is “Trifolium, guyannense”. Doubtless this form originated with a nonFrench-speaking engraver. This form could only be 

correct if the plate had been published prior to the 

text. There seems to be no evidence that this happened, nor any information about the order in 

which the 72 fascicles appeared (Stafleu & Cowan, 

1976:79) if they did not come out simultaneously. 

Key to the Micronesian Species of Stylosanthes 

Calyx lobes 3-5 mm long, loment minutely beaked, beak not over 0.5 mm long ...... S. guianemk 

Calyx lobes 1.5 mm long, loment with curved or hooked beak 1.5-3.5 mm long .......... S. humilis 

Stylosanthes guianensis (Aublet) Swartz 

Stylosanthes gziianensis (Aublet) Stvartz, Svenska Vetensk. 

Trifoliurn guianense Aublet, Hist. P1. Guian. Fr., 776, 

Acad. Handl. 10:301, 1789. 

T.309, 1775. 

In Micronesia reported only from Nauru, originally from tropical America. 

Stylosanthes htamilis Humboldt, Bonpland 8c Kunth 

Stylosanthes huniilis Humboldt, Bonpland & Kunth, Nov. 

Stylosanthes sundaica Taubert, Verh. Bot. Brandenb., 32:21, 

Gen. Sp., G:506, 1823. 

1890. 

In Micronesia reported only from Nauru, originally from tropical America. 

EUPHORBIACEAE 

Acalypha L. 

This vast genus is inherently difficult, but its 

taxonomic intricacy is enhanced by the myriad of 

slightly differing forms that have been recognized 

and described as species. Certain of the complexes 

would be more manageable if considered as broad 

aggregate species, with the more recognizable pop 

ulations regarded as varieties and forms. 

One such complex, for which the oldest name 

seems to be Acalypha amentacea Roxburgh, extends 

from the Malesian area well into the Pacific. 

Acalypha amentacea Roxburgh 

Acalypha amentacea Roxburgh, F1. Ind. ed. Carey, 686, 1832. 

This was originally described as follows: 

Dioecious, shrubby. Leaves long-petioled, ovate-cordate, 

taper-pointed, three-nerved. Stipules ensiform; spikes axillary, 

as long as the petioles, the male amentaceous; involucres 

small, hairy, dentate, one-flowered; style multifid. 

A native of the Moluccas. It differs from Konig’s A. 

Oetulina (in Retz. Obs. or N. 85) in having but one female 

flower in the involucre. 

This species was reduced to A. fruticosa Forskal 

by Muller-Argau (1866: 822), this reduction being 

followed by Pax and Hoffmann (1924:169), and by 

later botanists generally, except Merrill. This reduction seems manifestly incorrect, because A. fruticosa (an African-Arabian western Indian species) 

does not reach the Moluccas. The description fits 

the large shrubby or arborescent species that is common in a myriad of variations and local races from 

India to Fiji and Tonga at least, and north to the 

Philippines and Carolines. 

IVhile it may seem drastic to propose reduction 

of such long recognized species as A. grandis, A. 

stipulaceu, A. angatemis and A. wilkesiana, no one 

has yet offered any convincing attempt to distinguish them by other than superficial key characters, 

and these are frequently unreliable. 

The differences in this assemblage are in leaf 

size, shape, length of petiole, pubescence of stem 

and leaf, size and shape of stipules, thickness, den- 
8 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

sity, length and hairiness of staminate spike, length 

and density of pistillate spike, shape and dentation 

of bracts of pistillate spike, and in the case of A. 

wilkesiana in its forms, color, and modification of 

leaves. 

It is proposed to treat this assemblage as one polymorphic species, within which many lesser units 

will eventually have to be distinguished. A few of 

these, most of which already have names, can be 

recognized at once, but to classify the variations in 

this group as a whole, and even to be certain which 

described species are to be included, and what their 

precise limits are, is a major taxonomic task and 

cannot be attempted here. 

Typification of A. a7nentncea presents problems. 

No specimen was located at Kew, either in the general herbarium or in the \\:allich Herbarium. Dr. 

J. R. Sealy, formerly of the Kew Herbarium says 

(pers. comni.) that the Xlolucca plants described by 

Roxburgh ha1.e not been located. Nothing is known 

about this collection. Most of the plants were not 

illustrated and cannot be typified at present, except 

by the descriptions. Sealy says it would be worth 

while to look in the Brussels and Calcutta herbaria 

to see if these Alolucca specimens are there, but we 

have not pursued this suggestion. In the British 

Aluseum is a specimen, “Amboyna Dec. 1801” 

[Christian Smith pencilled on label]. It represents 

a plant with large ovate-cordate or subcordate 

leaves, blades 18-22 cm long, 11-15 cm wide, 

serrate, teeth acute to someTvhat acuminate or 

mucronate, apex prominently acuminate, nerves 

densely hirsute beneath, reticulation slightly hirsute, petioles densely hirsute, 7-1 7 cm long; stipules 

ovate-lanceolate, long-acuniinate, 12-14 nini long, 

sericeous except margins. Staminate spikes slender, 

about 2 mm thick, 10-13 nim long, the rachis densely 

hirsute, the basal 1-2 cm almost bare of flowers; 

bracts scale-like, 1 niin or so long, acuminate, hirsute; a single stout pistillate spike, 1.5 mm thick, 8 

cm long, densely pilose, sparsely flowered, bracts reduced, 2 mm long, hirsute Ivithout, with 8-12(?) 

lanceolate teeth, each bract subtending one flower, 

ovary hirsute, style many-branched, 1-2 mm long. 

This specimen bears 110 name on the original 

label, but E. D. Xlerrill has annotated it in 1951 as 

Acnlypha amentacea Roxburgh with a note: “This 

117ight be the collection on which Roxburgh’s species 

was based as Charles Smith’s material went to him.” 

It is a reasonably good match for Robinson 353 

(BM), perhaps a bit more hirsute, pistillate spike 

less densely flowered, bracts less developed than on 

most spikes of no. 353 but about like those on the 

youngest spike, which seems comparable in age. It 

is undoubtedly the same species. 

It seems expedient to recognize two subspecies, 

based mainly on the dentation of the bracts of the 

pistillate spikes, the aspect of the plants, and leaf 

outline. 

Acalypha amentacea Roxburgh ssp. amentacea 

Terminal tooth of bract not conspicuously longer 

than lateral ones. 

Pending further study, all of the subordinate 

taxa excepting wilkesiana are placed in this subspecies. 

Acalypha amentacea Roxburgh var. amentacea 

Acalypha aniboynensis Bentham, Lond. Jour. Bot., 2:233, 

Acalyplia stipulacea Klotzsch in Meyen, Nov. Act. Acad. 

Acalypha gradis var. arnboinensis (Bentham) Muller-Argau, 

1843. 

Cur, 19 (suppl.):416, 1843. 

Linnaea, 34:10, 1865. 

There seems to be no significant difference between material collected in Amboina (Robinson 

353, 354) and some of the variable series of material from the Philippines distributed by Dr. E. D. 

Alerrill and well represented in the U.S. National 

Herbarium. 

Leaves broadly ovate-subcordate, strongl! acuIninate, softly pubescent beneath; young parts 

sericeous-tonientose; stipules lanceolate-acuminate; 

staminate spikes congested, becoming elongate but 

remaining densely flowered, pistillate more loosely 

flowered, equally long, with ovate incised bracts, 

these sparsel) hairy. 

Characterization based largely on Robin yon 35?, 

353, from Amboina. 

Acalypha amentacea var. grandis (Bentham) 

Fosberg, new status 

A. grandis Bentham in Hooker, Lond. Jour. Bot., 232-233, 

1843. 

Leaves cordate orbicular, somewhat acuminate. 

softly and sparsely pubescent or somewhat strigose 
NUMBER 45 9 

beneath, stipules linear acuminate to almost filiform; young growth toinentulose; staminate spikes 

moderately congested, pistillate with bracts reniform 

with about 7 acute teeth; ovary and fruit densely 

hirtellous. 

As interpreted here this variety is known definitely only from Fiji. The Barclay specimen cited 

below is here designated lectotype. The name 

Acdyplza. gmndlis Bentham has been applied to a 

diversity of plants from Melanesia, Micronesia, antl 

Malesia, especially the Philippines, and even 

Hawaii. These specimens, so far as we have seen 

them, belong to several other varieties, not all 

worked out yet. The 1Iicronesian ones are mostly 

var. palauensis and ssp. wilkesiana. 

SPECIME ss SEES.-Fiji: without locality, Hinds in 1841 (K, 

synt>x); Suhalau, Zlni.c/crs S.PZ. (K, syntype. photo in US): 

Kambara, H. F. ,\loore 48 (US). 

.licalyphu amentacea var. palauensis Fosberg, 

new variety 

.4cci/)f1/1n jrnticoco seiiFu Kanehira [noti Forskal, F1. Aegypt., 

.lco/~p/~n gi.ciiit/is seiisu auct. blicr. [non Bentham in Hooker, 

161, 17751. 

Lond. Jour. Bot., 2:232, 18431. 

Frutes partibus junioribus tomentulosis vel 

strigulosis, periantho masculo leviter puberulo, 

1 obi s br ac tear u in f oenii near 11 in (5 -) 5 -9 ova t is vel 

lanceolatis glabris vel strigosis. Type: Palau Is. 

Peliliu, Fosbcrg 476 ii (US, holotype). 

Shrub, young stems sparsely to densely strigulose 

or appressed puberulent; leaves broadly ovate, 

rounded to subcordate or cordate at base, acuminate, margins rather coarsely crenate-serrate, both 

surfaces very sparsely appressed hirsute mostly on 

ner\-es to almost glabrous, petioles thinly strigose 

to almost glalirous; stipules narrowly ovate-acuminate to linear-subulate, glabrous to lightly sericeous, 

allout 1 cni long: staminate spikes slender, longer 

or shorter than lea.i.es, rachis tomentulose or puberulent with appressed or incurved hairs, bracts shorter 

than flowers, :icuminate, peiianth sparsely pilosulous; pistillate spikes from much shorter than 

leaves antl slendei-, to as long as or exceeding leaves 

and quite stiff, rachis sparsely to densely tomentulose, flowers sparse to crowded, bracts sparsely to 

densely strigose, lobes (5) 5-9, ovate to lanceolate, 

glabrous to somewhat strigose especially on nerves 

without, middle one longest; ovary and fruit densel) 

to lightly strigose; ovate; the spikes vary enormously 

in length and density of flowering; the staminate in 

prominence of the bracts and presence or absence 

of one or two pistillate flowers at base; the pistillate 

in stiffness of rachis and from almost glabrous to 

densely strigose, the bracts from 5- to 9-toothed, the 

teeth from glabrous to hairy and from lanceolate to 

ovate. About the only completely constant featui~s 

are the appressed pubescence, the ( renate-serrate 

leaf margins, the acuminate apices, and the relatively few teeth on the ovate (vs. reniform) bracts. 

Yet there seem no reason to believe that this is 

other than a coherent, freely interbreeding probably 

monophyletic population. It is found, so far as 

known, only 011 the limestone areas of Palau. It 

differs notably in pubescence from the other native 

Palauan variety heterotricha, which has only been 

found on the volcanic part of Babeltlaob Island. 

SPECIMESS SEEN.-Caroline Islands: Palau: without locality, 

Ledermann 14072 (B); Cheatham I63 (US, BISH, UC). 

Kayangl: Ngajangl I., Gressitt 15 (US), 35 (US). Gatulel-to, 

Ailai-son, Hosokaiva 730i (Fo). Auluptagel: (Oropusyakaru), 

Hwcoknua 9096 (A); Orokusva~;aru-ii~iFoku, Hosokawa 9783 

(.4); Ngerengchol, Lee Marvin Beach, Cape Pkula-heap, 1 rn, 

Canfield 452 (US). Aulong I., Salsedo I2 (US). Aulupse’el: 

N\V end, Stone 4538 (BISH, US); Sgerebe’ed beach, E end, 

2 m, E‘osberg $if77 (US, BISH); Du’ebachel beach. 2 m, 

Fosberg 4y451 (US, BISH). Peliliu: Emnzons 58 (US, BISH); 

S end of E side, “Purple Beach,” 0-2 rn, Fosberg 26007 

(US, BISH); S part near NE end of airstrip, Fosberg 47637 

(US. holotype); Sgdlkol \illage, 2 in. Forbrrg 4i633 (US. 

BISH). Angaur: S/c/.iedo 356 (US); roadside midway between 

Lake A and Lake D, 10 m, Ca?i/ie/d 202 (US). Sonsorol: 

Hni-dy I25 (CS, BISH); Beri-y 28 (US, BISH). 

Acalypha amentacea var. heterotricha Fosberg, 

new variety 

Frutex pubescens, partibus junioribus hirsutis 

pilibus brevibus vel longis mistis, stipulis ellipticis 

vel oblongis valde acuininatis, periantho inasculo 

glabro, rhachitlibiis spicorum dense liirsutulis, lobis 

hr ac t ear um foe m i ne ai- uiii (7-) 9 1 anceol a ti s 11 i rs u ti s 

adpressis. Type: Palau Is.: Babeldaob, Fosberg 

TO576 (US, holotype, isotypes; BISH, isotype). 

Sparsely branched pubescent shrub to 3 m tall, 

young stems densely covered with a mixture of 

appressed and erect sharply pointed hairs of varying 

lengths; leaves o\.al or broadly elliptic, sharply and 

prominently acuminate, base subcordate to cordate, 

margins finely serrate, ciliate, upper surface rvith 
10 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

sparse scattered large hairs, those on veins smaller, 

under surface more densely hairy, especially on 

nerves, petioles 5-8 cm long, densely pubescent, 

nerves 5-7 from base, pinnate above; stipules oblong 

to elliptic, 1-2 cm long, 3-4 mm wide, thin, strongly 

acuminate, falcate, appressed to erect pilose within 

and without, erect hairs near margins and on lower 

part of midrib, staminate spikes densely flowered, 

4-15 cm long, slender, rachis densely hirsute, bracts 

densely hirsute without and on margin, triangular, 

scarcely exceeding flowers, perianth glabrous; pistillate spikes on separate branches or distal on same 

branch with staminate, 5-22 cm long, rachis densely 

hirsute, flowers and subtending bracts separated to 

almost touching, bracts appressed hirsute without, 

especially on nerves, glabrous within (7-)9-lobed, 

lobes lanceolate, middle one longest, the outermost 

very short, ovary and fruit shortly hirsute-pilose. 

This is the only collection of Acalypha amentacea 

from the volcanic part of Palau, and it is strikingly 

distinct in its indument from the common var. 

palauensis from the limestone areas. 

SPECIMEN Smh..-Caroline Islands: Palau: Babeldaob, west 

side, Ngerumongui, on savanna slopes on bauxitic soil, 

23 Nov 1968, Fosberg 50576 (US, holotype, isotypes, BISH, 

isotype). 

Acalypha amentacea var. trukensis 

(Pax & Hoffmann) Fosberg, new status 

Acalypha trukensis Pax & Hoffmann, Pflr., IV, 147(XVI): 151, 

1924. 

Leaves orbicular, subcordate to cordate, upper as 

well as lower surfaces of leaves with some hair, 

pubescence of petioles and young parts of long, stiff, 

erect hairs; stipules linear subulate, lightly sericeous; staminate spikes slender, loosely flowered; pistillate with longer peduncles than usual, floriferous 

portion equalling or shorter than peduncle, bracts 

broadly triangular cordate, sparsely pilose, teeth 7, 

triangular subulate. 

Differs from var. grandis principally in the stiff, 

yellowish pubescence on young growth and petioles, 

and in the less pubescent lower leaf surfaces. 

SPECIMENS SEm.-Caroline Islands: Truk: Tol, 150-200 m, 

Kanehira 1284 (US, NY); To1 Island, Uriribot, Hosokawa 

8254 (A); Moen Island, Tunuk District, Spence 434 (BISH); 

Natusima [Dublon Island], Hosokawa 6526 (A, BISH); 

Takamatsri i0 (BISH); Zogeyasi-hayasi, Hosokawa 655’4 (Fo); 

central part of Ulalu Island, 100 ft [30 m], Wong 237A 

(US, BISH, K), 237 (US, BISH, K); Melot, Hosokawa 8344 

(BISH). Lukunor Atoll Anderson 2099 (US, BISH, Fo) (sterile 

young plant, atypical in broad stipules possibly because 

young). 

Acalypha amentacea var. velutina (Muller-Argau) 

Fosberg, new combination 

Acalypha grandis var. uelutina Muller-Argau, Flora, 47:441, 

Acalypha angatensis Blancho, F1. Filip., 750, 1837. 

Acalypha tomentosa Blanco, F1. Filip., 750, 1837. 

1864. 

Leaves, ovate, rounded to subcordate at base, 

strongly but gradually acuminate, plant strongly 

tomentose, velutinous, and rarely with long stiff 

hairs, under surface of leaves softly pubescent, especially on nerves, upper surface pubescent on nerves; 

stipules ovate-lanceolate, acuminate, tomentose. 

Merrill (1918:227) equates this with Acalypha 

angatensis Blanco and Acalypha tomentosa Blanco. 

It was referred to Acalypha grandis as a variety by 

Muller-Argau. Actually there is no Acalypha grandis 

(Acalypha amentacea var. grandis) in the Philippines. Acalypha amentacea is very variable there, 

and includes the specimen, M.R. 333, that Merrill 

uses to illustrate Blanco’s Acalypha angatensis. 

Plants with narrow, gradually acuminate leaves, 

velvety young growth and under sides of leaves, 

almost all from Luzon, seem to be separable and 

to correspond with Muller-Argau’s A. grandis var. 

velutina, basionym of A. amentacea var. velutina 

(Muller-Argau) Fosberg. 

SPECIMENS SEEN.-Luzon: central, Loher 4675 (US); Benguet 

Prov.: without locality, Basani 15882 (US); Baguio, Topping 

246 (US); Elmer 5924 (US); Mearns 2525 (US). Union Prov.: 

Bauang, Elmer 5727 (US): Abra Prov.:Mt. Posuey, Rantos 

26974 (US). Cagayan Prov.: without locality, Castillo 22727 

(US). (Province not indicated) Nueva Viscaya, near Quiaugau, 

Merrill 156 (US); Batanes Is.: Sabtan, McGregor 10161 (US). 

Acalyphu amentacea ssp. wilkesianu 

(Muller-Argau) Fosberg, 

new combination, new status 

Acalypha wilkesiana Miiller-Argau in de Candolle, Prodr., 

15(2):817, 1866. 

Shrub, much branched, twigs with short internodes, condensed toward apices, closely puberulent, 

leaf-scars prominent, reniform, narrow horizontal 

stipule scars at sides, circular or broad horizontally 
NUMBER 45 11 

elliptic inflorescence or branchlet scars above; leaves 

with blades very broadly ovate to suborbicular, 

cordate to more usually subcordate, up to 15 cm X 

13 cm, apex prominently acuminate but acumen 

with a rounded or etnarginate tip, five main veins 

from base, midrib with about 4 pairs of subopposite pinnate nerves, main nerves connected with 

ladder-like tertiary veins, intervals filled with network of decreasing prominence, margins serratecrenate with low, very obtusely pointed crenations, 

points and sinuses both terminating branch veinlets 

from the shallowly scalloped marginal vein formed 

by anastomosing nerves, each point somewhat 

glandular thickened and with a small straight hair 

or seta, easily caducous; stipules ovate, very strongly 

acuminate; spikes dark red, slender, 4-10 cm long, 

strongly ascending, the staminate much fewer and 

below the pistillate on the branchlet, staminate 

buds in dense sessile clusters, rather close together 

on rachis, pistillate spikes with flowers single, alternate and opposite, each subtended by broadly 

ovate-cordate somewhat glandular bract, this with 

a large ovate terminal tooth and 3 or 4 much 

smaller ovate teeth on each side, these becoming 

slightly smaller toward the base, ovary densely 

white-hirsute slightly shorter than bract, styles 3, 

deeply filiform-fimbriate, 5-8 fimbrils on each 

branch, these up to about 5 mm long; no fruit 

seen setting. 

The type, ITS. Expl. Exp. from “Feejee” (US), 

does not differ much from this description except 

that it shows a peculiar contraction of the basal part 

of nerves often seen in cultivated specimens. 

This plant, f. z~iilkesiana, is known throughout 

hficronesia and the rest of the tropics in cultivation 

but it is not known in an indigenous wild state anywhere. It apparently persists after cultivation and 

establiqhes small seedling populations which tend 

to “revert,” losing the characters of colored and 

deformed leaves which distinguish it from A. 

amentacea ssp. amentacea. Such plants have been 

collected in Tl’ailau Valley, LIolokai, Hawaiian Is. 

(Degcner 8381(US)) and Rota, Marianas (Necker 

RB(US)), both determined by A.C. Smith as A. 

gi~nrli,~. Smith (1952:391) distinguishes A. wilke- 

.c.iana from A. g~~~ndi.c on the basis of much shorter 

petioles often flattened at apex, and colored leaves. 

The dentation of the bracts on the pistillate spikes 

is also rather distinctive, the teeth being fewer and 

broader than in most other varieties, the apical one 

much longer than the lateral ones, and the bracts 

being less hairy. The leaves are variable and tend 

to be distorted, with the veins drawn down to the 

petiole. hfany garden forms are known, such as the 

following: 

Acalypha amentacea ssp. wilkesiana f. 

circinata ( Muller-Argau) Fosberg, 

new combination, new status 

Acalypha wilkesiana f. circinata Muller-Argau in de 

Candolle, Prodr., 15(2):819, 1866.-Pax and Hoffman, PBr. 

I\‘, 14i(XV1):154, 1924. 

Leaves much distorted, broadly cordate but 

drawn down to petiole, arcuate, petiole very short, 

blades usuallj green with white borders. 

Occasionally planted in Micronesia, as in Yap 

and Nauru. Said to have been introduced into Yap 

by the Germans. 

Acalypha arnentacea varieties 

Tl‘hat seem to be other varieties of this species are 

known from Ponape (G/a\5maj7 2916 (BISH, US)) 

and from the Gilbert Islands ant1 Nauru. They are 

as yet not well characterized and should be further 

collected and studied. 

Acalypha cardiophylla Merrill 

Acalypha cardiophylla Merrill, Phil. Jour. Sci., 1 (suppl.): 

Acalypha catzrrus sensu auct. [non Blume, Bijdr., 629, 18261. 

This species was described from the Philippines. 

Acalypha cardiophylla hlerrill seems to be the 

oldest unequivocal name for it. It has generally 

been called Acalypha catuiur Blume, but this is an 

illegitimate supei fluous name, as Acalypha rpicigera 

L. was cited as a synonym. The var. cardiophylla is 

knolvn fi om the Philippines and posSibly elsewhere 

in hfalesia. It is not at all closel) related to the 

Acalypha an7entatea complex. 

80, 1906. 

Acalypha cardiophylla var. ponapensis 

(Kanehira & Hatusima) Fosberg, 

new combination, new status 

Acalypha poizapensis Kanehira and Hatusima, Bot. Mag. 

Tokyo, 54:434, 1940. 
12 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

This variety differs from var. cardiophylla principally in being essentially glabrous. The bracts subtending the pistillate flowers are minute, triangular; 

the pedicels of the staminate flowers are short, 1 mm 

or less long. 

It is endemic in Ponape. 

CELASTRACEAE 

(including HIPPOCRATEACEAE) 

Loeseneriella A.C. Smith 

Loeseneriella macrantha (Korthals) A.C. Smith 

Loeseneriella macrantha (Korthals) A. C. Smith, Am. Jour. 

Bot., 28:439, 1941.-Ding Hou, Blumea, 12:31-32, 1963; 

F1. Males., I, 6:398, 1964 

Hippocratea macrantha Korthals, Kruidk. Verh. Nat. Gesch. 

Nederl. Ind. 187, pl. 39, 1839-1842. 

Loeseneriella macrantha (Korthals) A.C. Smith var. 

macrantha 

Extends from Ceylon through Malesia to the New 

Hebrides, not known from Micronesia. 

Loeseneriella niacrantha var. palauica (Loesener) 

Fosberg, new combination 

Hippocratea ntacrantha var. palauica Loesener, Bot. Jahrb., 

63:274, 1930. 

Found in Palau, Yap, and Truk. 

A. C. Smith (1941:439), when he established the 

genus Loeseneriella A. C. Smith with L. macuzntha 

(Korthals) A. C. Smith as type, mentioned that 

Loesener had published a variety of Hippocratea 

wzacmntha from Palau, but made no comment on 

it. Neither in Blumea (12:31-32, 1963) nor in Flora 

Malerianu (I, 6:399, 1964), where he treated LoeJeneriella macrantha, did Hou make any mention or 

disposition of the variety palauica. Hence we are 

transferring it to the correct genus. 

GUTTIFERAE (= CLUSIACEAE) 

Calophyllum L. 

Three species of Calophyllum L. have been reported from Micronesia, C. inophyllum L., C. wakamatsui Kanehira, and C. cholobtaches Lauterbach. 

Of these we are treating C. wakamatsui as a variety 

of C. inophyllum and C. cholobtaches as a synonym 

of the widespread C. soulattri Burman f. 

Dr. P. F. Stevens is permitting us to publish a 

new species, C. pelewensis Stevens, resulting from 

his as yet unpublished monographic studies of the 

genus Calophyllum. 

Calophyllum inophyllum L. 

Calophyllum inophyllum L., Sp. Pl., 513, 1753. 

This is a widespread and familiar Indo-Pacific 

strand and lowland tree, often planted as a shade 

tree or ornamental. Its var. inophyllum has large 

globose fruits and is fairly uniform. In Palau an 

endemic upland form seems varietally distinct. 

Calophyllum inophyllum var. wakamatsui 

(Kanehira) Fosberg & Sachet, 

new combination, new status 

Calophyllzciri wakamatsui Kanehira, Bot. Mag. Tokyo, 

48:401, 1934. 

This upland population in Babeldaob Island, 

Palau, has smaller, more leathery leaves and somewhat smaller slightly elongate or ovoid fruits. Dr. 

P. F. Stevens (pers. comm.) does not separate this 

from var. inophyllum, but we found it to have a 

different appearance in the field and the Palauans 

consider it distinct, calling it esmolech while var. 

inophyllum is called btaches. 

SPECIMENS SEEN.-Caroline Islands: Palau: Babeldaob: 

Aimeliik, Kanehira 2326 (FU), 200 m, 2343 (FU, holotype); 

Airai area beyond airfield, Cheatham 181 (US); between 

Nekken and Ngatpang, Cheatham 188 (US); Airai Dam Site, 

Fisher I35 (US); 2 miles [3.2 m] on road to Nekken from 

Airai, Salsedo 317 (US); Ogiwaru, Takamatsu 1430 (US, 

BISH); Marukiyoku (Melekiok), Takamatsu 1704 (US, BISH); 

Lake Ngardok, 25 m, Fosberg 32526 (US, BISH); Takamatsu 

13Z7 (US, BISH). 

Calophyllum pelewense P.F. Stevens, new species 

Calophyllunt cholobtaches auct. [non Lauterbach, Bot. Jahrb. 

59:20, 19241.-Kanehira, Bot. Mag. Tokyo, 45:329, 1931; F1. 

Micronesica, 233, fig. 105, 1933; Jour. Dept. Agric. Kyushu 

Imp. Univ., 4:370, 1935. 

A speciebus aliis Calophylii in gemma terminali 

parva 1-2.5 mm longa, innovationibus axillaribus 
NUMBER 45 13 

cicatricibus basalibus ornatis, lamina mediocra 

elliptica vel ovata costa infra elevata cum 6-10 

nervis per 5 mm, inflorescentiis cum internodiis 

infimis 0-2 mm longis, et floribus cum 8-11 tepalis, 

differt. 

Type: Palau, ,July-Aug. 1929, Kanehira 386 (4, 

holotype; BISH, NY, isotypes). 

Arbor; truncus et cortex haud cogniti. Ramulus 

compianatus, 1.52 mm in transverso, valde 4- 

angulatus vel subalatus, in siccitate brunneus, 

glaber; par superius gemmarum axillarium rotundatum, ca. 0.5 mm longum, effusum; gemma terminalis corpulenta 1-2.5 mm longa indumento 

breve appresso griseo praedita. Petiolus 0.9-1.5 cm 

longus, supra vadose et anguste vel late concavus, 

infra convexus, glaber; lamina elliptica vel ovata, 

raro obovata, 3.5-10.2 cm longa et 2.1-4.9 cm lata, 

apice acuininata raro plus minusve rotundata, basi 

acuta vel attenuata, margine unclulata leviter recurvata, coriacea, glaber, costa supra e basi sensim 

angustata, haud vel leviter depressa, 0.35-0.6 mm 

lata in medio laminae, infra elevata, plus minusve 

striata, nervis lateralibus utrinque manifestis, elevatis, 5-10 nervis per 5 mm sub angulo 55-70' e 

costa divergentibus. Inflorescentiae e axillis foliatis 

pope apices ramulorum ortae, raro terminales, 

cum 9-17 floribus, haud ramosae; axes 1.3-8 cm 

longi, basi indumento puberulo praediti, internodio infimo 0-2 mm longo; bracteae subovatae, 

circa 3.5 mm longae, mox deciduae; pedicelli 1-2.5 

cin longi, glabri. Flos hermaphroditus (?); tepala 

8-1 1, duobus exterioribus late ovatis circa 4 mm 

longis et latis, pilis brevibus fimbriatis, tepalis interioribus obovatis 5.5-6 mm longis et 3.5-4.5 mm 

latis, glabris; stamina 125-230, filamentis circa 3 mm 

longis, antheris suboblongis, 0.8-1 mm longis apicibus retusis; ovarium circa 2 mm longum, glabrum; 

stylus circa 2.2 mm longus; stigma peltatum, 0.7-1.2 

mm in transverso, t 3-radiatum. Fructus probabiliter ellipsoideus (valde oppressus), circa 1.5 cm 

longus et 1 cm latus, apice rotundatus, in siccitate 

rugosus; stratum exterius circa 0.5 mm crassum; 

putamen parietibus circa 0.2 mm crassis, laevibus. 

Endemic to Palau, Calophglliini pelewense has 

been collected on river banks at the landward edges 

of in a ngrove s . 

Calophgllzrm pelewense may be recognized by its 

very short terminal bud, axillary innovations with 

basal scars and axillary inflorescences with a short 

basal internode, usually elliptic to ovate leaf blades 

with rather prominent, distant venation, flowers 

with 8-11 tepals, and fruits of moderate size (ca. 

1.5 cm long). 

Specimens of Calophpllum pelewense have been 

labeled C. cholobtaclies (equals C. soulattri), but the 

two species are not at all close. Calophyllum soulattrz has a longer terminal bud, a greenish drying 

lamina, a branched inflorescence, flowers with four 

tepals, and a spherical fruit with a thick, compact, 

outer layer. 

SPEcImNs SEm.-Caroline Islands: Palau: without locality, 

Ledernmn I4489 (B, K); Kanehira 386 (A, holotype; BISH, 

NY, isotypes). Babeldaob: Garasumao, Takamatsu 1556 

(BISH, US); Aimiriik, Kanehira 1984 (US, NY, K, P); 

Hosokawa 7216 (BISH): Armatin, 8 m, Takamatsu 1525 

(BISH); Airai, Kanehii-a 2380 (SY); Sgerikill River, Fisher 

25 (IYS), 55 (US). Koror: Raymundus 334 (WRSL). 

Calophyllum .sozdattri Burman f. 

Caloplzylluni soulattri, Burman f., F1. Ind., 121, 1768. 

Calophyllunz cholobtaches Lauterbach, Bot. Jahrb., 59: 20, 

1924. 

Dr. P. F. Stevens (below) informs us that Calophylluna cholobtnches, described as a Palau endem i c, i s in cl is t i ngu i s h a bl e from the widespread 

C. soulattri known from southeast Asia, the Philippines, Malesia, New Guinea, and 1Ielanesia. This 

is not an unusual distribution pattern for a Palau 

plant. Dr. Stevens will designate 1,crlei ~naiziz 14251 

(IZ'RSL, fragment) as lectotype of C. cholobtache.f 

1.auterbach. This specimen is the only one of those 

originally cited that is fertile and it clearly agrees 

with the description; the other specimens were collected by Rayniundus. 

SP~CIXIENS Srrs.--C;iroline Islantl~: Palau: Koror: Leder. 

//innn If25/ (IVRSL); Uruhthapel, Dutlofi 80 (US). 

Garcinia L. 

Garcinia ponapensis Lauterbach 

Garcinia ponapensi5 Lauterbach, Bot. Jahrb. 59:21, 1924. 

Garcinia ponapensis var. trulzensis (Kanehira) 

Fosberg, new combination, new status 

Garcinia trukensis Kanehira, Bot. Mag. Tokyo 46:488, 1932. 

This plant is at most a variety of G. ponnpensi.5 

differing in slightly narrower, more oblong leaves. 
14 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

fruit less strongly ribbed and not umbonate. 

Kanehira cited two of his own collections, the first 

of which Kanehiru 1269 (FU, US) is here designated 

as lectotype. 

Garcinia rumiyo Kanehira 

Garcinia rumiyo Kanehira, Bot. Mag. Tokyo, 45:329, 1931. 

This species has been known from Yap and Palau, 

where it is not common but is frequently found. 

Examination shows that the plants from Yap (type 

locality) and Palau are varietally separable. 

Garcinia rumiyo Kanehira var. rumiyo 

Endemic to Yap. 

Garcinia rumiyo var. calcicola Fosberg, 

new variety 

Arbor foliis ad 11 X 6 cm, petiolis crassis, pedicellis 3 mm longis, fructibus 15 mm longis, calicis 

10 mm latis. Type: Palau: Kgelobel, H. Owen 5 

(US, holotype). 

Small tree, to 10 m tall, leaves somewhat larger 

than in Yap plants, blades to 11 x 6 cm; pedicels 

about 3 mni long. fruit 1.5 cm long, fruiting calyx 

10 mm across. 

Apparently confined to rough limestone substrata, 

endemic to the southern half of Palau. 

SmcIvENs Srr:r.-Caroline Islands: Palau: without locality, 

Richardson 36 (US). Limestone Islands, without further locality, Dutton 97 (US, BISH, L); Ngelobel, H. Owen 5 (US, 

liolotype); Trukodokorokke, coral island, Takamnt.su I150 

IBISH, US). Koror, Kanehira 1890 (FU, US), 350 (FU); Hosoknrcia 9791 (US). Aulupse’el, Dii’ebachel Beach, 2 m, Fosberg 

4if61 US). Peliliu, Knnrhira 24-19 IFU). 

FLACOURTIACEAE 

Flacourtia Commerson ex L’Heritier 

Flacourtia rukam Zollinger & Moritzi 

Flacourtia rukam Zollinger & Moritzi, Svst. Verh., 33, 1854. 

This species has been reported from Palau, Truk, 

;md Ponape. We have studied a series of collections 

from Micronesia and find that they differ, at least 

varietally, from the typical 1 ariety, which is known 

from nlalesia and Samoa. 

Flacourtia rukam var. micronesica 

Fosberg & Sachet, new variety 

Frutex vel arbor parva, folii costa utrinque glabra 

vel subglabra, stigmatibus 6-7 subsessilibus. Type: 

Ngarakabesang, Fosberg 25668 (holotype, US; isotypes, BISH, L). 

Small tree or large shrub; leaves broadly ovate 

or elliptic to oblong, usually about 10 X 6 cm, on 

East growing shoots or branches to 20 X 10 cm, 

bluntly acuminate, rounded at base, margins obscurely or markedly crenate to serrate-crenate, 

blades subcoriaceous when mature, midrib glabrous 

or subglabrous on both sides, petioles 7-1 1 mm long; 

racemes about 1 cm long; sepals glabrous without; 

fruit depressed globose, about 8 mm high, 10 mm 

wide, subtruncate at both ends, 6-7 lobed, at least 

when dry, with a horizontal constriction at about 

middle, the 6-7 stigmas subsessile, in a ring or 

ellipse when mature or almost so. 

The Palau and Truk specimens of F. rukam differ 

significantly from those from Indonesia, Melanesia, 

and Samoa in the glabrous or subglabrous midribs 

of the leaves and in the subsessile stigmas on the 

summit of the fruit. The non-Micronesian specimens examined have definite styles up to 1 mm 

long. We have not seen the specimen (Kanehira 

1542) reported from Ponape; but we have seen two 

sterile sheets from there, collected by Tukamatsu 

(822, 822-A) that are probably var. micronesica. 

This variety is found in secondary forests and 

thickets, at low elevations in Palau, rather higher 

in Truk. 

SPECIMENSEEN.-Caroline Islands: ’ Palau (Pelew): without locality, Kanehira 352 (US) (fruit). Babeldaob: 

Ngarekalong, Takainatsu 1674 (BISH); Ngatpng, Takamatsu 

1311 (BISH); Arukodokorokke, Takamatsu 1145 (BISH); 

Ngersuul, Ngchesar, Emnions 81 (US); Arumizu, Hosokawa 

YO82 (US); Arumonogui, near Arumatan, Hosokawa 6758 

(A, BISH). Koror: Kanehira 343 (BISH, 2 sheets); Ngermid, 

Snlsedo 307 (US); Mt. Kororu, Hosokawa 9800 (US). 

Ngarakabesang (Arakabesan): Takamatsu 125.2~ (BISH); west 

side near old Japanese seaplane base, 1-20 m, Fosberg 

25668 (US, holotype; BISH, L, isotypes) (fruit); west peninsula of Ngarakabesang, 10-20 m, Fosberg 32471 (US). 

Aulupse’el (Olopshakal): Takamatsu 1489 (BISH); Du’ebechel 

beach, 2 m, Fosberg 47443 (US, BISH) (seedling). Urukthapel: 

\Vest peninsula near old cultured pearl establishment, 2-10 

m, Fosberg 32147 (US) (pistillate flowers); limestone islet, 

Dutton 101 (US, BISH, K) (sterile, Palauan name “emeong”). 

Peliliu: Kn?iehira 2632 (US) (fruit). Angaur: East side, 3-5 

m, Fosberg 31994 (US, BISH, MO). Kasioru, Takamatsu 1516 

(BISH). Truk Is.: without locality, Takaniatsu 6 (BISH); 
NUMBER 45 15 

Kanehzra 129f (US) (staminate flowers). Moen: Takamatsu 

2f5 (BISH, 3 sheets); Mt. Trokken (Wara), Hosokawa 8407 

(BISH); track to highest part of island, 100400 m, Evans 

1268 (US, BISH, NY). Dublon (Natsushima): Takarnatsu 

162 (BISH); upper ridges and top of Mt. Tolomen (Tolowan) 200-360 m. Fosberg 24544~ (US, BISH, L, MO, Fo) 

(joung fruit), (Trowasi) Mt. Troman (Tolomen), Hosekawa 8510 (US, .A) (staminate flowers). Fefan: Mt. Tuktyap, 

Hosokawa 8385 (.4, BISH). TO]: Suiyo-to, Hosokawa 8319 

(BISH, ,A). Ponape: Toleailuka, Takarnatsu 822 (BISH), 822 

.4 (BISH). 

CUCURBITACEAE 

Citrullus Schrader 

Citrullus lanatus var. caffrorum (Alefeld) 

Fosberg, new combination 

Cifrulliis vulgaris var. caflrorum Alefeld, Landwirth. FI., 

Cucirrbita raflra Ecklori & Leyher, Coll. Sem. 1833. 

Cilrullits caffer Schrader, Linnaea 10, Lit. Ber., 109, 1834. 

Citritllits lanatit.t var. cafler (Schrader) Mansfeld, Die 

210, 1866. 

Kul~urpllanre. kill., 2:422, 1962. 

Var. cagrorum Alefeld here lectotypified by 

Cucurbita cagra Ecklon & Zeyher seems to be the 

earliest varietal epithet for the common red-fleshed 

cultivated watermelon. It has not apparently been 

transferred to Citrullus lanatus the oldest specific 

name for what has usually been called Citrullus 

vulgaris Schrader. 

Zehneria Endlicher 

Zehneria guamensis (Merrill) Fosberg, 

new combination 

Melothria guamensis Merrill, Phil. Jour. Sci. Bot., 9: 151-152, 

Studies by Charles Jeffrey (1962:342-344) indicate 

that the genus Melothria L. occurs only in America. 

The Pacific island species belong to Zehneria. 

1914. 

RHIZOPHORACEAE 

Bruguiera Lamarck 

Bruguiera gymnorhiza f. alba (Stone) 

Fosberg, new combination 

Rrugiriera conjugata f. alba Stone, Pac. Sci. 13: 102, 1959. 

Hou (1958:453, 463) has shown that the correct 

name for what has been called Bruguiera conjugata (L.) Lamarck is B. gymnorhiza Lamarck (as 

gymnorrhiza). 

MELASTOMATACEAE 

Medinilla Gaudichaud 

Medinilla medinilliana (Gaudichaud) 

Fosberg & Sachet, new combination 

Melastoma medinilliana Guadichaud, Bot. Voy. Uranie 69, 

Medinilla rosea Gaudichaud, Bot. Voy. Uranie, Tab. 106, 

1826. 

1829, 484, 1830. 

Gaudichaud (1 826:69), used the name Melastoma 

medinilliana accompanied by a few words of description, effecting valid publication of the binomial. Later he (1 830:484) described Medinilla 

rosea, placing Melastoma medinilliana in synonymy. 

The earlier ,epithet must be restored (ICBN Art. 11). 

Melastoma L. 

Melastoma malabathricuna var. mariannum 

(Naudin) Fosberg & Sachet, 

new combination, new status 

Melastoma mariannum Naudin, ,41111. Sci. Nat. 111 Bot., 

13:276, 1849. 

The differences between M. malabathricum and 

its immediate relatives, including M. mariannum, 

seem trivial, and several of these entities merit, at 

most, varietal rank. M. mariannum has white 

flowers and a small few-flowered inflorescence. 

ARALIACEAE 

Osmoxylon Miquel 

Osmoxylon Miquel, Ann. Mus. Bot. Lugd.-Batav. 1:5, 1863. 

Boerlagiodendron Harms in Engler and Prantl, Nat. 

Pflanzenfam. 111, 81:31-32, 1894. 

Philipson (1976) demonstrated that in all probability Boerlagiodendron Harms is congeneric with 

the previously poorly known Osmoxylon Miquel. 

He does not treat the Micronesian species. Accepting his conclusion, we here provide the new combinations for the Micronesian species in Osmoxy- 
16 SMITHSOSIAN COSTRIBUTIONS TO BOTANY 

lon, and describe one new species to replace a 

misidentification current in Micronesian literature. 

Osmoxylon mariannense (Kanehira) 

Fosberg & Sachet, new combination 

Roe, lagzodendron niai iannense Kanehira, F1. Micr., 288, 

1933; Bot. Mag. Tok\o, 47:676, 1933. 

Endemic to Rota, Marianas, first collected by 

hfarche (264 P, US), but the species remained undescribed until 1933, when it was based on Kanehira 

1773,1774 (FU, syntypes). 

Osmoxylo~i truucatum (Kanehira) 

Fosberg & Sachet, new combination 

Iloerlagiodendron trrincatiim Kanehira, Bot. Mag. Tokyo, 

48:403, fig. 2, 1934. 

Known only from Aimeliik in the southwest part 

of Babeldaob, Palau, Kanehim 2364 (FU, holotype) 

and 2303 (FU). 

Osmoxyloti pachyphyllum (Kanehira) 

Fosberg & Sachet, new combination 

h’ori Ingiodendrou $acli~phslltim Kanehira, Bot. Mag. Tokyo, 

48:401, 1934. 

Known from Babeldaob Island, Palau, described 

from 3 syntype collections, Aimeliik, Kanehira 

2301,2311, and 2452 (all FU). 

Osmoxylon oliveri Fosberg & Sachet, new species 

Iloerlngioderidron piilrherriiniim sensu Kanehira, F1. Micr., 

185, fig. 143, 1933 [non OsnioxyIon pulcherrimum Vidal 

ex F.-Villar, Sovis. App., 102. 18801. 

Arbor parva, folk grandis reniformis 11-13 

lobatis glabris, alis basalibus petioli integris non 

pectinatis, umbella multiradiata bracteis linearibus 

vel o\.atis acuminatis inarginibus non-fimbriatis, 

floribus bacciformibus sterilibus pedicellatis. 

Type: Palau: Babeldaob, Cheatham 54 (US, holotype; BISH, NY, L, Fo, isotypes). 

Tree to 5 m tall, unbranched (always?), leaves 

with blades reniform to somewhat orbicular, 

glabrous, deeply 11-13 lobed, 50 cm or more in 

diameter, lobes elliptic to lanceolate, apices of lobes 

obtuse to acute or acuminate, basal sinus deep, 

rounded to acute, petiole angular in cross section at 

least when dried, 80-100 cm long; base swollen 

with 2-5 thin, entire, nonpectinate, transverse wings 

or flanges (petiolar crests); stipules clasping, bifid, 

lobes ovate, acute or acuminate, 1.5-2.5 cm long; 

inflorescence a sessile or very shortly pedunculate 

hemisplierical compound umbel of many trifid rays, 

these to 15 cm long, lowest internode lenticellate, up 

to 5 cm long, subtended by a linear hard bract with 

a thin white margin, with 2 such bracts at summit, 

subtending the 2 outer rays, outer rays 6-8 cm long 

of 2 internodes, the lower 1.5-2 cm long with 2 

ovate-lanceolate bracts at summit, upper 4-5 cm 

long, ending in a tight subcapitate umbel or head 

of hermaphrodite flowers on extremely short pedicels, attached to an ellipsoid-capitate receptacle or 

enlargement of the summit of the ray, central ray 

very short, 0.5 to at most 1 cm long, ending in an 

umbel of about 10-12 “sterile bacciform flowers” 

these on slender pedicels up to 5 mm long; perfect 

flowers with calyx reduced to a persistent fleshy 

ring, corolla cylindric to cylindric-campanulate, 

gamopetalous, 3-4 mm long, leathery, lobes 4-5, 

1 mm long, ovate, incurved, tips slightly hooked, 

stamens 4-5 strongly exserted, filaments rather 

thick, basally attached, hooked at the slender tip, 

anthers oblong, attached near base; disk elevated, 

star-like with wide blunt points, in the center a ring 

of 5-6 oval stigmas depressed in center; fruit turbinate to broadly obovoid, deeply 5 lobed, 6-8 mm 

long; sterile flowers pea-like, 5-8 mm diameter, with 

3 radiating abortive locules. 

This species was indicated as amply distinct from 

Osmoxylon pitlchenimzim Vidal ex F.-Villar (with 

which it has been equated up to now) by our assistant, Nr. Royce Oliver, to whom we have dedicated it. TVith an ample series of collections it has 

been possible to draw up a rather full description. 

The most obvious distinctions from 0. pukherriniitnz are in the pedicellate baccifom sterile flowers, 

in the entire, nonpectinate petiolar flanges and the 

nonfimbriate bracts. From Osmoxylon pachyphylIum, it is distinguished by thinner leaves with many 

more lobes. It is, so far as known, endemic to Palau, 

mostly on Koror and Babeldaob islands. 

SPI c 1111 \s Sr.rx.-Caroline Islands: Palau: withorit locali[i, Clipntiicii~i ii (CS), /Oi (t.S); Richardson 55 (US). 

Dabrldaoh: DiilioJi 7f ((3, BISH), Cizentham 8 (US); Marikyoku, Kanehira 2057 (FU) (sterile, could conceivably be 0. 

lvioicotiiin); Ogiwal. Knriehirn 2065 (FU), 2066 (FC), Dudiii’s 
NUMBER 45 17 

homestead near boundary between Ngeremlengui and 

Sgardmau, below 300 ft [90 m] Cheatham 54 (CS, holotype; 

BISH, NY, L, Fo, isotypes); Ngerremlengui, Cheathain I07 

(US, BISH, Fo), 108, 109, I10 (all US), Ill (US, BISH, Fo), 

Ngliaklolubed, old Japanese botanical garden site, Fosberg 

32332 (US, BISH, Fo), west coast, 1 m, Fosberg 32435 (US, 

BISH); Sgaraard Slun., on tributary of Ngereakl River, 

0.8 mi [l km] SSTT of Kulotauk, 2 m, Canfield 397 (CS); 

Sgerelong, 0.2 mi C0.3 km] W of Pkulrengerelong, 10 m, 

Canfield ?04 (US). Koror: Kanehira 129 (FU), 1853 (FU); 

(cult.) “Kesiamel” Blackburn 280 (US, BISH); A‘gerbe’edesa~c, 

5-10 m, “siama” k‘osberg 12379 (US, BISH). Aulupse’el: 

Risong, Matuker Bay. south side of island, 10-15 m, 

“kesiamel,” k‘otbel-g 47560 (US, BISH), 

MYRSINACEAE 

Myrsine L. 

Ji)rsirie L . Len P1 eti. 7. 90, 1754 [15i3]. 

IZapanea 4ublet, Hist. P1. Guiane Franc., 1:121, t. 46, 1775. 

Following the study by Hosaka (1940) and our 

own further consideration (1975:3-11) of the Polynesian species, we regard Rapanea Aublet as insufficiently distinct from Myrszne L. Hence we here 

transfer the three Micronesian species of Rapanea 

to Myrsine. 

Myrsine carozinensis (Mez) Fosberg & Sachet, 

new combination 

Rapnnea rarolinensis Slez, Bot. Jahrb., 56:538, 1921. 

T)pe: Ponape, Lerleimann 13695 (B, 2 isotypes). 

Myrsine ledermannii (Mez) Fosberg & Sachet, 

new combination 

Rnfmnea ledermnnnzi Sler, Bot. Jahrb., 56:538-539, 1921. 

Type: Ponape, Ledermann 13786 (B, 2 isotypes). 

Myrsine palnuensis (Mez) Fosberg & Sachet, 

new combination 

Rapanea fialaiiensis Slez, Bot. Jahrb. 56:539, 1921. 

Type: Palau, Babeldaob, Ngatkip, Ledermann 

14529 (B). 

SAPOTACEAE 

Pouteria Aublet 

We have consistently followed Baehni (1942) in 

including in the large pantropical genus Pouteria 

Aublet the Pacific species described in Sideroxylon 

L., and usually comprising the genus Planchonella 

Pierre. This in spite of the opinion of van Royen 

(1957:239), who separates Planchonella from 

Pouteria on the basis of the relative thickness of 

the cotyledons and the albumen. All other characters, according to van Royen, overlap. He attaches 

considerable importance to the predominantly 

American distribution of Pouteria and the predominantly Pacific distribution of Planchonella. 

Our present impression is that the single clear difference between the two is insufficient for generic 

separation. We are not adamant in this opinion 

and will reconsider the matter critically when preparing the treatment of the family for the Flora. 

The classification of the Sapotaceae is in a most unsatisfactory state and no disposition can be regarded 

as very stable at present. We are making, however, 

two new combinations that are needed for consistent use of Pouteria for plants of this affinity in 

Micronesia. 

Pouteria calcarea (Hosokawa) Fosberg, 

new combination 

5rderoxylon calcareicin Hosokawa, Trans. Nat. Hist. Soc. 

Planchonella? calcarea (Hosokawa) 1 an Royen, Blumea, 

Formosa 32:li, 1942. 

8:422. 1957. 

This species seems closest to Pouteria micronesica 

but differs in its smaller leaves and 1-2 seeded 

fruits. 

Caroline Islands: “Hab. Palau, Islet Oropsyacal 

[~-Iulupse’el] rare in primar) forest on a coral rocky 

place (Hosok. no. 9790!-Type in Herb. Taihoku 

Imp. Univ.-Aug. 30, 1941).” I\’e have not seen this 

specimen. 

Pouterin micronesica (Kanehira) 

Fosberg, new combination 

Sideroxplon micronesicurn Kanehira, Bot. Mag. Tokyo, 46:671, 

1932; F1. SIicr., 308, pl. 158, 193.3; Enum. P1. SIicr., 388, 

1935. 

Planchonella micronesicn (Kanehira) Kanehira ex Lam, 

Blumea, 5:12, 1942; Proc. 6th Pac. Science Congr., 4:681, 

1910.-\an RoLen, Blumea, 8:402-404, fig. 45, 1%;. 

~,‘~i~~.\~~~~~~~~~fcri~ sp? Baelini. Candollea, 0:128, 1‘342. 

This species is amply distinct from the widespread 
18 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Pouteria obovata by the large, elliptic-oblong leaves 

with strongly acuminate apices and more numerous 

lateral veins. 

SPECIMEN SEEx.-Caroline Islands: Kusaie: without local. 

ity, Kanehira I322 (DISH, isotype). 

GENTIANACEAE 

Systematic Position of the Genus Fagraea 

Dr. P. W. Leenhouts (pers. comm., 1976) has questioned our removal of the genus Fagraea Thunberg 

from the Loganiaceae to the Gentianaceae (Fosberg 

and Sachet, 1974:471-472; Sachet 1975: 18-19>, and 

we grant this requires some explanation. These 

families are usually placed close together, but seem 

to be separated on the basis of presence or absence 

of stipules, primarily woody vs. herbaceous habit, 

and by the Gentianaceae seldom having a baccate 

fruit. 

However, the Gentianaceae do have woody members: Lisianthus P. Browne, Macrocarpaea Gilg, 

Symbolanthus G. Don, all of tropical America, and 

Gentianothamnus Humbert of Madagascar. Rusbynnthur Gilg has “capsule berry-like, with a thin 

cartilaginous wall, rupturing irregularly” according 

to Ewan (1948:212). Also, Gentianothamnus Humbert has at the bases of the leaf or of the petiole 

two small lobes resembling the “stipules” of 

Fagreae. 

Tachia Aublet, another woody member of the 

Gentianaceae in South America, has a low stipular 

collar (Figure 2) and traces of lobes similar to those 

of Gentinnothamnus. That no attention has been 

paid to these may be due to the abundant gum that 

a b 

FIGURE 2.-Nodes of Tachia gztianensis Aublet: a, glandular 

hairs in collar surrounding young leaves; b, older node with 

collar separate from leaf bases and surrounding flower stalks 

in axils. 

covers the nodes in this genus, probably secreted by 

the glandular hairs lining the inside of the collar 

(Figure 2). 

When, in 1942-1944, I (Fosberg) saw living in the 

Colombian Andes the three genera of woody 

Gentianaceae subtribe Tachiineae, Macrocarpaea, 

Symbolanthus, and Lisianthus, I was strongly impressed by their resemblance to the Indo-Pacific 

loganiaceous genus Fagraea Thunberg, though I did 

not doubt that these three genera belonged to the 

Gentianaceae. The genus Lisianthus serves to connect the first two with more typical members of the 

Gentianaceae. I had not then seen Anthocleista 

Afzel, the African representative, and Potalia Aublet, the American representative of Fagraea, especially a form of Potalia amara Aublet from the Rio 

Negro, Vaupes, Colombia (Schultes et al. 18127 

(US), I8304 (US), and Schultes Q Lopez 9339 (US)), 

which has inflorescences similar to those of Mncmcarpaea. These specimens incidentally, probably 

represent a species distinct from P. amara. 

After re-examining herbarium material of the 

three genera of the Tachiineae listed above and of 

the three genera of Potalieae, our conclusion remains that they are closely related. If the transfer 

of the Potalieae to the Gentianaceae, which is our 

choice, is unacceptable, we could consider transferring the Tachiineae to the Loganiaceae, increasing 

the artificiality of the latter family. 

Long ago, Asa Gray (1859:323) pointed out that 

the Loganiaceae is an artificial family, and that 

Fagraea seemed to show similarities to Lisianthus 

and related Gentianaceie, and to connect the latter 

with Rubiaceae, Orobanchaceae, and Loganiaceae. 

‘CVe are merely agreeing with him and making the 

necessary transfer. We would suggest that the 

Potalieae and the Tachiineae be united to form 

the tribe Potalieae, which name, being the oldest 

with the rank of tribe, must take precedence over 

Tachiineae, and be transferred to the Gentianaceae. 

In this tribe would be included Potalia Aublet, 

Ant h ocleista Afzel, Fagraea Thunberg, Lisiant hus 

L., Tachia Aublet, Tachiadenus Grisebach, Rusbyanthus Gilg, Chorisepalum Gleason and Wodehouse, Macrocarpaea Gilg, and Symbalanthus Don. 

Eustoma Salisbury does not seem to belong here. 

Several other genera, placed in the Tachiineae by 

Gilg (1895:90-94), we do not exclude, but neither 

are we familiar enough with them to formally include them. 
NUMBER 45 19 

We are not impressed with the “naturalness” of 

the subdivision of either the Gentianaceae by Gilg 

(1895) or of that of the Loganiaceae by Solereder 

(1892). Leenhouts (1962b:293-297) has discussed 

critically the subdivision of Loganiaceae. His classification takes into account a number of lines of 

evidence, and his conclusions are eminently conservative. His circumscription of the family is traditional. We differ from it in the placement of the 

Potalieae. 

Fagraea Thunberg 

Four species and one variety of Fagraea Thunberg have been described from western Micronesia, 

all considered endemic. Additionally, two names in 

this genus, F. berteriana and F. morindifolia, had 

been used earlier. 

To the five endemic taxa we axe adding a new 

one, and these six we consider to represent two 

species: Fagiaca ksid, which is endemic on Palau, 

and F. berteriana, which is a widespread Pacific 

species with five Micronesian varieties. 

References to Fagraea (mostly as Fragraea 

sphalm.), published earlier than the paper by Gilg 

and Benedict (1921) and given as Fagraea sp., to 

F. berteriana A. Gray ex Bentham, or to F. morindifolia Blume, record the genus from Nauru and the 

Gilbert Islands. To the best of our knowledge, they 

were not based on specimens. Burges (ca. 1935:4, 5) 

recorded Fagraea sp. in a manuscript list, citing a 

1910 record by Hambruch. Burges gave no further 

indication as to the source of the record. About all 

one can conclude from this is that Fagraea, probably 

F. berteriana, or a plant so identified, may have 

existed on Nauru. Fagraea sp. (written Fragraea) 

reported by Grimble (1924:103) and Fagraea (as 

Fragraea) berteriana, quoted by Loumala (1953: 

117-118) from Kramer (1906:268, 320, 351, 456) as 

from the Gilbert Islands, specifically Onotoa, both 

with the vernacular name, “te uri” or “to ori”, 

commonly used in the Gilberts for Guettarda, we 

regard as misidentifications of Guettarda speciosa. 

The occurrence of Fagraea on dry Onotoa Atoll 

seems very unlikely. Kramer (1905: 140), referring 

to Schumann and Lauterbach (1901), reports 

Fagraea morindifolia Blume, from the Marshall 

Islands, with the vernacular names, “wut” and 

“libiruk.” Schumann and Lauterbach (1901:499) 

record this species, quite distinct in its racemiform 

inflorescence, from the Solomon Islands, with a 

Malesian distribution, but make no mention of 

Micronesia or the Marshalls. Judging by the Marshallese name “wut” we believe this to be a misidentification of Barringtonia asiatica (L.) Kurz, 

which has somewhat similar leaves. TVe have no 

specimens or reliable records of Fagraea from 

Micronesia east of Kusaie. 

The taxonomy of Fagraea is ‘in an unsatisfactory 

state, despite the monographic efforts of Leenhouts 

(1962a, 196213.) The six Micronesian taxa are obviously close, but specimens can be sorted on even 

superficial characters into six piles, each with a 

definite and coherent geographic range. Workers 

publishing prior to 1962 unhesitatingly accepted 

five species as distinct, one of them with a variety. 

Leenhouts (1962b:335) placed all of these taxa in 

the synonymy of the widespread F. berteriana, not 

recognizing any of them, not even F. ksid, at varietal 

or other infraspecific rank. 

Fagraea ksid, with its slightly zygomorphic flowers 

and its oblongobovoid, somewhat dorsiventrally 

compressed fruit, held horizontally, would satisfy 

even a moderately broad species concept. The other 

five, recognizably though only slightly distinct, seem 

to fit very well as varieties within the broad and 

polymorphic F. berteiiana Gray ex Bentham (see 

Fosberg and Sachet 1974:471-472, for remarks on 

this species and its taxonomy). JVe are still not 

ready to publish a treatment of this species over its 

entire range, but the Micronesian varieties seem 

reasonably clear, though forming a closely related 

group. Mre are publishing our disposition of them 

here so that their names may be used. 

Key to the Micronesian Species of Fagraea 

Calvx lobes 8-10 mm long, acute, corolla tube o\er 6 cm long, fruit oboxoid, slightlv dorsiventrally corn pressed F. ksid 

Calyx lobes 3-7 mm, rounded or obtuse. Corolla tube 2-4 cm long, fruit ovoid to globose, subglobose or ellipsoid, not at all compresed F. berterianu 
20 SXIITHSOSIAN CONTRIBUTIOSS TO BOTAZIY 

Fagraea berteriana A. Gray ex Bentham 

Fcigi-nra berteriann A. Gray ex Bentham, Jour. Linn. Soc. 

Carirsa grnndis Bertero ex Bentham, Jour. Linn. SOC. Bot., 

Fngraea grnndis Pancher in Sebert, Sot. Bois Souv. Caled., 

Frcgrnen Jciclechteri Gilg & Benedict, Bot. Jahrb., 56:551, 

Bot., 1:98, 1855 [1836]. 

I:;?. 118, 1857 [1836]. 

184, 187.1. 

1921. 

T) pe: “Hab. In ins. Societatis (Bertero, Bidwill, 

Hin(ls, hrcL~\), in\ Suhdlii\ a e Marquesas (Barcla)), in Archipelago Louisiade dicto (Xfacgilli\ra))” 

Bentham. The Bertero specimen is indicated as t) pe 

[lectot) ~)e?] b! Leenliouts (1962a:-l20). 

Glabrous tree, branchlets rather flesh), tending to 

be quadrangular, leaves elliptic to obovate, thincoriaceous to coriaceous, petiolate, petioles auriculate at base; inflorescence terminal, basically dichasial, 3-4 times branched, a pair of scale-like bracts 

at each node; flowers pedicellate, very fragrant, 

pedicels to 15 mm long; calyx united below, with 

5 imbricate, thin-margined rounded lobes, erect, 

then spreading; corolla salverform, fleshy-cartilaginous, cream-white aging yellow, tube 2-7 cm long, 

somewhat to notably dilated in the upper part, 

lobes obovate, rounded distally; stamens inserted on 

a slightly thickened ring inside the corolla tube, 

anthers linear; ovary ovoid, style somewhat thickened near base, filiform, almost equalling tube to 

somewhat exserted, stigma of two flattened lobes 

erect and pressed together, then spreading; fruit 

an ovoid to subglobose or globose, usually umbonate, not strongly beaked by style base, orange to 

red many-seeded berry, the skin firm and glossy, the 

m;in! seeds enibetlded in fleshy pulp. 

The type locality of this species is Tahiti, and var. 

beq.teyiniin is not in Micronesia. In Micronesia it is 

represented by five very closely related varieties. 

They may be keyed out as follows: 

Key to the Micronesian Varieties of Fagraea berteriana 

1. Petiole mod\ under 20 mni long. slightl\ \\inget1 at least abole, b\ decurrent blade 

2. Cal\x lobes 3-4 (-5) mm 

2. Cal\x 3-6 mm 

1. Petiole inostlx o\er 20 inm, not winged. 

3. Corolla tube 20-27 nim long 

3 Corolla tube 28-35 mtn long 

4. Cal\u lobes 34 (-3) mm long 

4 Gal\\ lobes 4 5-7 nim long 

Fagraea Derteriana var. galilai (Gilg & Benedict) 

Fosberg, new combination, new status 

1.q)o‘eo go‘iiini Lilg k Uenctlict. Bot. Jahrb.. .56:5X, 1921. 

Leaves oval to obovate-oval, rounded to very 

shortly umbonate at apex, petioles 15-30 mm long, 

calyx lobes (3-) 3.54-5) nim long, corolla tube 

about 28 mm long, style equalling it or slightly 

longer, fruit globose to ovoid-globose, 16-18 X 

20-22 mm. 

Known only from Palau (Truk records all seem 

to belong to var, pops). 

Type: Palau Islands, Babeldaob: “bei Ngalkip, 

100 m, Mittelwdd,” Letlernianii 14476 (B, lectotype, 

in bud only, here designated). 

Fagraea berteriaiia var. kusaiaiia (Hosokawa) 

Fosberg, new combination, new status 

lag, a?ri k ucain,ia Hosokawa, Trans. Sat. Hist. SOC. Formosa, 

24.904, 1934. 

Far, sair 

\ ar. kusaiana 

var. fiogas 

kar. galilai 

\ar. ladronica 

Leaves broadly obovate, slightly acuminate, base 

cuneate, T eins somewhat prominent; petiole (12-) 

16-23 mm; calyx lobes 5(-6) mm long; corolla tube 

25-39 mm long, stigma included or exserted (possibly heterost) lous?); fruit broadly ellipsoid to subglobose or globose, 21-24 mm long, bluntly beaked. 

Close to vai. stti], but petioles longer scarcely 

winged except at summit. Knou-n onlj from Kusaie. 

T) pe: Caroline Islands: Kusaie: 5It. Buache, 

H o r o I< (I ic N 62SR (.I, i so t ) pe) . 

Fagraea berteriana var. ladronica Fosberg, 

new variety 

Planta foliis obovatis, petiolis usiter 17-30 mm 

longis, lobis caljcis 5-7 mm longis, tub0 corollae 

31-35 mm longo, fructu globoso vel ellipsoideo, 

25-30 mm longo. 

TJpe: Rota, Fosberg 25142 (US, holotype; BISH, 

Fo, isotjpes). 
NUMBER 45 21 

Leaves obovate, very slightly to not pointed, base 

somewhat cuneate, veins quite visible, petiole (12-) 

17-30 (-31) mm long, calyx lobes (4-) 5-7 mm long, 

corolla tube 31-35 mm, style subequal with corolla 

tube or slightly exserted; fruit globose to ovoid or 

ellipsoid, 25-30 mm long. 

This is close to var. galilai, to which it has been 

referred, but differs especially in its much longer 

calyx lobes, longer corolla tube and larger fruit. 

Known from Guam and Rota in the Marianas. 

SITCINFTS Sixs.-Marianas Islands: Rota: mi. [1.2 km] 

E of Sabana 420-450 m. Fosberg 25142 (US; type; BISH, Fo, 

isot!pes); Sabana, Secker 1iS-l (US), A’ecker RSlj (US); road 

to Sabana, 200-400 in, Sachet 1807 (US); 150-250 m. Evans 

2096 (US). Guam: without locality, Anderson 136 (US); N of 

Almagosa, 330 m. Brjan 1-726 (SY); Mt. Lamlam, Glassman 

2ii (.I): dforan 4713 (US, Fo); 900 ft [275 m], Moore 260 

(US, H.4IV): llt. .Umagosa, Stone 4340 (US); 300 m, Stone 

4109 (GUAM): Sa\al Magazine area, near Almagosa Springs, 

Stone 4S99 (GL.4JI) (corolla tube not or scarcely exserted, 

throat 22-23 mm long, lobes 12-14 mm long, fruit oval, 

slightly limbonate, 19 X 25 mm). 

Fagraea berteriana var. pogas (Hosokawa) 

Fosberg, new combination 

Fagiaea sair iar. pogas Hosokawa, Bull. Biogeugr. SOC. Japan, 

7:198, 1937. 

Leaves obovate to oblong-obovate, petioles (16-) 

20-30 (-35) mm long, not or scarcely winged, calyx 

lobes 34 mm, corolla tube 20-25 mm long, style 

strongly exserted, 23-30 (-38) mm long, fruit oval, 

15-18 mm long, with a low umbo 2-4 mm high. 

Found on Truk and in the hlortlock Atolls. 

Type: Caroline Islands: Truk Group, Toll Island, 

Hosokawa 8333 (A, isotype). 

Fagraea berteriana var. sair (Gilg & Benedict) 

Fosberg, new combination, new status 

Fagraea $air Gilg 85 Benedict, Bot. Jahrb., 56:555-557, 1921. 

Leaves oblong to obovate, slightly pointed, base 

cuneate to slightly attenuate, petiole 7-15 (-30) mm 

long, narrowly winged by decurrent base of blade, 

venation somewhat obscure but visible, calyx lobes 

(3-) 4 mm long, corolla tube 23-36 mm long, style 

included to usually exserted; fruit globose or rarely 

subglobose, 25-30 mni in diameter. 

Knolvn onl) from the central eastern Carolines, 

native in Ponape, probably introduced in Kamoluk 

and Nukuoro. 

Type: Caroline Islands: Ponape, “bei Palapolap 

im Garten,” Ledermann 13932 (B, isotype). 

Fagraea ksid Gilg & Benedict 

Fagraea ksid Gilg & Benedict, Bot. Jahrb., 56:552-553, 1921. 

Small tree, 4-12 m tall, glabrous; leaves broadly 

obovate to oval, thick, glossy, to 15 X 9 cm, rounded 

at apex, cuneate to slightly attenuate at base, veins 

subopposite, 6-8 pairs, visible but not prominent, 

petioles 2-2.5 cm long, first pair on a branchlet 

tending to be reduced to obovate cataphylls; cjme 

terminal several times trichotomously branched, 

with reduced leaves at first ramification and at first 

ramification of central branch; flowers slightly 

zygomorphic, pedicellate, pedicels erect, calyx lobes 

8-10 mm long, obtuse, closel) erect around base of 

corolla; corolla tube 6-10 cm long only very slightly 

dilated upward, lobes obovate 2.5 cm long, 1-1.6 

cm wide rounded at apex; anthers included, style 

exceeding tube, stigma bilobed, exserted; pedicels in 

fruit nodding and fruit either horizontal or somewhat drooping, caljx spreading in fruit, berry 

ellipsoidal to narrowly pyriform, dorsiventrally 

somewhat compressed to about 4.5 X 2 X 1 cm, 

apex rounded but abruptly strongly beaked by the 

stiff, narrow stjle-base, this 4-6 mm long, deep 

orange when ripe, 

Endemic to Palau, rather common on Babeldaob 

on weathered volcanic rock, on Koror reported by 

collector as on limestone. The syntypes from Koror 

Island (Ledermann 14,189, Gibbon (Kersting’s) 

1205, and Raymundus 209) were probably all destroyed in the ’1Vorld War 11 bombing of the Berlin 

herbarium. 

LOGANIACEAE 

Geiziostoma J. R. & G. Forster 

The genus Geniostoma In the Marianas Islands 

presents much difficult) in interpietation. Plants of 

this genus have been collected in the Marianas on 

Guam, Rota, Saipan, Alamagan, Pagan, and Agrigan. In all, four species have been described: G. 

micianthum by A. de Candolle, G. hoefeiz by Gilg 

and Benedict, its var. glabium and Geniostoma 
22 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

longistylum by Gilg and Geniostoma glaberrima by 

Hosokawa. In addition, Kanehira (1935:392) proposed the name G. saipanense “in sched.” but apparently never published his description. These 

four all represent distinguishable forms, differing 

mainly in hairiness or lack of it, and length of style. 

In leaf-shape and size and fruit size, there is also 

much variation but this is not well correlated with 

the variations in pubescence and style-length. 

If only the Guam plants were involved it might 

be possible to uphold two species, differing in several characters, one, glabrous growing on the volcanic soil, the other, generally hairy, on limestone. 

The glabrous plant has the summit of the ovary 

glabrous and the style 0.5-1 mm long. The hairy 

plant has the style 0.5 mm or less long and a slight 

hairiness at the summit of the ovary. One plant 

(Marche 282) from Guam but without locality is 

glabrous but with very short style and hairs at the 

summit of the ovary. 

Plants similar to the common glabrous one from 

Guam occur on Rota (on volcanic soil), on Agrigan 

and on Alamagan (one specimen from Alamagan 

with a few hairs at top of ovary). Agrigan and 

Alamagan are strictly volcanic islands. The Alamagan specimens have short styles. Essentially glabrous 

plants, similar to the Marche 282 collection are also 

known from Saipan, where they occur on volcanic 

soil. 

On Pagan is a plant with leaves very slightly 

tomentulose beneath, along midrib and acuminate, 

with very small fruit, 2.5-4 mm long, style 0.5 mm 

long, ovary slightly hairy at summit. 

Taking into consideration only the separation 

based on pubescence, the correlation between glabrous plants and volcanic substratum is striking, 

and that between hairy plants and limestone is 

complete except for the Pagan Island specimen. In 

each of the two, however, both short-styled and 

long-styled plants occur. Even without desirable 

field studies on the populations, the probability 

seems strong that G. micranthum is a heterostylous 

species. Accepting this, it seems reasonable to regard the glabrous and the pubescent populations, 

with the exception of the Pagan plant, as varieties, 

each inhabiting a distinct habitat. The Pagan specimen, which does not fit this picture, also differs in 

having a much smaller fruit. The best disposition 

of it, at present, seems to be to make it a separate 

variety of G. micmnthum. 

Geniostoma micranthum A. de Candolle 

Geniostoma rnicranthuni A. de Candolle, Prodr., 9:27, 1845.- 

Gilg. and Benedict, Bat. Jahrb., 56:541, 1921.-Merrill, Phil. 

Jour. Sci. Bot., 9: 128, 1914.-Fosberg, Falanruw and Sachet, 

Smithsonian Contr. Bot., 22336, 1975.-Stone, Micronesica, 

63473-474, 1971 [as G. rupestre Forster]. 

Geniostoma hoeferi Gilg & Benedict, Bot. Jahrb., 56:541, 1921. 

Geniostonm Iongist~li~nt Gilg, Sotizbl., 12:221, 1934. 

Geniostoma hoeferi var. glabrum Gilg, Notizbl., 12:221, 1934. 

Geniostonia gluberriina Hosokawa, Tr. Sat. Hist. Soc. Formosa, 25334, 1935. 

Geniostoma rupestre sensu Stone, Leenhouts [nec J. R. and 

G. Forster, Char. Gen., 24, 1775, nec A. de Candolle, 

Prodr., 9:26-27, 18453. 

This species has been generally recognized as distinct since the time of de Candolle. However, in 

1962, Leenhouts 1962b:369-373) took a much 

broader view of Geniostoma rupestre J. R. & G. 

Forster, the type-species of the genus. He submerged 

a vast number of Pacific and Malesian species in 

G. rupestre, not even giving them varietal status. 

Included was G. micranthum A. de Candolle. He 

was followed in this course by Stone (1971:473-474). 

Conn, of Lae, New Guinea, who is studying the 

genus, has also annotated material of G. mzcranthum as G. rupestre. On the other hand, in a critical 

and detailed study of the Western Polynesian, 

Fijian, and New Hebridean species, Smith and 

Stone (1962:32-36), have restricted G. rupestre to 

the New Hebrides, Santa Cruz Islands, and possibly 

New Caledonia. 

Genzostoma mzcranthum is obviously very close 

to G. rupestre, but differs in having much less 

acuminate, usually smaller leaves with closer venation, usually shorter, thicker petioles; cymes with 

flowers more congested, usually more numerous, and 

with peduncles longer. 

It seems more consistent with our treatment of 

the eastern Polynesian species of Geniostoma (Fosberg and Sachet, 1975:ll-13) to keep the two sep 

arate, at least until Conn’s monograph appears. 

Leenhouts (1962b3372) has stated the situation in 

this genus briefly but rather well; however, coming 

to opposite conclusions from ours. 

Endemic in the Marianas Islands. 
NUMBER 45 23 

Key to the Micronesian Varieties of Geniostoma micranthum 

Plants essentially glabrous var. micranthum 

Plants tomentulose at least on the underside of the leaf-midrlb, young stems and inflores1. 1. 

cences pilosulous. 

2. Fruit 5-10 mm long 

2. Fruit 2.54 mm long 

Geniostoma micranthum A. de Candolle var. 

micranthum 

Plants essentially glabrous except flower parts; 

fruit 5-10 mm long. Known from volcanic soils on 

Agrigan, Alamagan, Saipan, Rota, and Guam. 

Type: Marianas Islands: Guam: Gaudichaud 138 

(G, holotype; P, isotype). 

Geniostoma micranthum var. hoeferi 

(Gilg 8c Benedict) Fosberg, 

new combination, new status 

Geniostoma hoeferi Gilg & Benedict, Bot. Jahrb., 56: 541, 

1921.-Kanehira, Enum. Micr. Pl., 391, 1935.-Stone, Micronesia, 6:474, 1971. 

Geniostoma longistylum Cilg, Sotizbl., 12:221, 1934.-Kanehira, Enum. Micr. Pl., 392, 1935.-Stone, Micronesica, 6:474, 

1971. 

Youngest growth and lower side of midribs tomentulose, cymes pilosulous, apex of ovary pilosulous, fruit 5-10 mm long. 

Found on limestone on Saipan, Rota, and Guam. 

Types: Marianas Islands: Saipan: Schnee XI; 

Hoefer 64, 77, 84, syntypes, not seen, probably destroyed. Kanehira 2232 (NY, isotype of Geniostoma 

longistylum). 

Geniostoma micranthum var. paganense 

Fosberg, new variety 

Folium lamina elliptico-oblonga usque ad 8-10 

cm longa leviter acuminata costa infra tomentulosa; 

fructus late ellipsoidalis 2.5-4 mm longus summi 

pilosulus. Type: Pagan, Kanehira 2209 (NY, holoLeaves elliptic oblong, up to 8-10 cm long, somewhat acuminate, midrib very lightly tomentulose 

beneath; style 0.5 mm long; fruit broadly ellipsoidal, 2.5-4 mm long, slightly hairy at summit. 

Apparently endemic on Pagan. The principal 

distinctive character is the small fruit. 

tYP+ 

var. hoeferi 

var. paganense 

SPECIMEN Smrv.-Marianas Islands: Pagan: without locality, Kanehira 220Y (XY, holotype). 

BORAGINACEAE 

Heliotropium L. 

Heliotropium procumbazs Miller 

Helzotropium procumbens Miller, Gard. Dict. ed. 8, Heliotropium 10, 1768 [unpaged].-Johnston, Contr. Gray Herb., 

Heliotroptum inundatum Swartz, Prodr., 40, 1788; F1. Ind. 

81:52-54, 1928. 

Occ., 1:343, 1797. 

Similarity between specimens from Guam, commonly referred to Heliotropium ovalifolium var. 

depressum, and Mexican specimens labeled Heliotropium procumbens Miller or Heliotropium inundatum Swartz suggested that they might belong to 

the same taxon. If so, this species may also have 

been another of the many Mexican and other tropical American plants that were carried to the Philippines and Guam very early by the Spanish “Manila 

Galleons.” This led to a preliminary comparison of 

Heliotropium ovalifolium Forskal with its New 

World counterpart, Heliotropium procumbens 

Miller, of which Heliotropium inundatum Swartz 

is a synonym. This rather cursory study showed 

almost no differences between the two. I. M. 

Johnston had no<ted this similarity but continued 

to maintain the two species. The fact emerged from 

our studies that the Guam plant is virtually identical with the narrow-leafed form of Heliotropium 

procumbens found on the west coast of Mexico, the 

area from which the Manila Galleons sailed. Our 

conclusion is that, whatever the disposition of 

Heliotropium ovalifolium Forskal, its Guam variety 

should be transferred to Helioti opium procumbens 

and its circumscription made to include the narrowleafed west Mexican plants, as well as those of the 

western Pacific area. Before combining the two 

species ovalifolium and procumbens, it seems advisable to carry on a more detailed study, but it is 
24 ShIITHSON'IAN COSTRIBUTIONS TO BOTANY 

probable that more difference may exist between 

populations within each of the species than between 

the two species. Such a study will have to be postponed for the present, however. Here we will discuss 

only the western Pacific Island variety. 

Heliotropium procumbens var. depressum 

(Chamisso) Fosberg & Sachet, new combination 

Heliotropiunz graci/e R. Brown p depessum Chamisso, 

Linnaea, 4:45i, 1829. 

Heliotropiuirz roromandelianum var. depressum (Chamisso) 

A. de Candolle, Prodr., 9:542, 1845. 

Heliotropium coromandelianum var. oblongifolium A. de 

Candolle, Prodr., 9:541-542, 1845. 

Heliotropium ovalifoliitm var. depressum (Chamisso) Merrill, Philippine Jour. Sci. Bot., 9:134, 1914; 10:340, 1915; 

Enum. Phil. F1. Pl., 3:378, 1923. 

Generally, both Heliotiopium ovalifolium Forska1 and Heliotiopiurn procumbens Miller have 

obovate to elliptic petiolate leaves that vary in 

width and shape of apex and in degree of canescence. In New IYorld specimens (H. proczimbens) 

the spikes tend to be more slender and more often 

trichotomous. These differences are by no means 

sharp or constant. However, some specimens, mostly 

from the west and north parts of Mexico and a few 

scattered ones from farther south and east, as well 

as the Pacific Island population (which we are assigning to Heliotiopium procumbens), stand out 

because their leaves are narrow, linear-lanceolate to 

oblanceolate (or elliptic-lanceolate), acutely or acuminately pointed and frequently notably canescent. 

The cymes are mostly dichotomous, occasionally 

simple and axillary, rarely trichotomous. The habit 

is occasionally depressed, decumbent, as indicated 

by the epithet depiessum, but it also may be ascending or erect. In habit this variety occasionally 

resembles Heliotropium cuiassauicum L., but the 

latter is never pubescent, while Heliotropium procumbens is always more or less strigose. The cymes, 

in age, become conspicuously elongate. 

As indicated above, we suggest that var. depressum came to the Philippines and then to Guam 

by the Manila Galleons several hundred years ago. 

Judging by the facts that there is only one doubtful 

Philippine specimen in the U.S. National Herbarium and that Merrill only cites one Philippine 

collection, this introduction has not prospered in 

the Philippines, but became common in Guam, 

where Chamisso found it in 1817 (Chamisso s.n. 

(K,G,LE)), and Guadichaud in 1819 (Gaudichaud 

15 (P) s.n. (G)). At least by 1889 it had spread to 

Rota (Marche 271 (P, US)). In 1939 it was found on 

Tinian (Fujikawa). In 1965 it was found in Saipan 

(Fosbeig 47720 (US)) and in 1950 in Pagan (Fosberg 

?1640 (US)). By 1965 it had reached Ulithi in 'the 

TYestern Carolines (Fosberg 46488 (US)). In the 

Marshalls it was established in Kwajalein in 1952 

(Fosberg 34124 (US)) and in Majuro at least by 1978 

(Fosbeig 58810 (US)). On Wake Island it was found 

in 1961 (Sachet 904 (US)) and in 1963 (Fosberg 

43537 (US, RISH)). By 1971 it had reached Canton 

in the Phoenix group (Clapp photo), (Fosbeig and 

Stoddart 54897 (US)) and in 1973 spread to Hull 

Atoll, in the same group (Fosberg Q Stoddard 54842 

(US, HAIY)) where it still persisted in 1975 (Fosberg 

6 Stoddaid 55741 (US, BISH)). Herbst found it in 

the Hawaiian Islands at Sand Island, Oahu, in 1975 

(Heibst 5466 (US)), and Fosberg found it at 

Kaneohe Marine Corps Air Station in 1978 (Fosberg 

and Euans 588?4A (US)). It is, of course, not to be 

assumed that the date on which the species was first 

collected on an island corresponds to the date of 

introduction, The paucity of collections before 

I\'orld TVar I1 of this and other exotics, and their 

abundance since that period, illustrate a greatly increased dispersion of these plants, presumably due 

to greatly stepped-up inter-island travel. 

The narrow-leafed plant, introduced into the 

Pacific, corresponds well with plants from Mexico, 

especially western and northern Mexico, but represented slightly in the more central and southern 

part. It is there not the exclusive form, even in the 

northwest part, as broad-leafed plants are fairly well 

distributed in Mexico. No narrow-leafed specimens 

were seen in our brief study from either the West 

Indies or South America where the broad-leafed 

ones seem to be common. In our experience this 

variety is a plant of sparsely vegetated weedy places. 

Representative specimens are cited below. A full 

range will be cited with treatment of the Boraginaceae in a later fascicle of the Flora of Micronesia. 

We have seen only one specimen from Ithe Philippines. 

SPECIMES SE~S .--3larianas Islands: without locality, 

Gaitdiclsaud s.n. (G, type of Heliotropium coromandelianum 

var. oblongifolium); Guam: without locality, Chamisso s.n. 

(K,G,LE) (isorypes of Heliotropium gracile var. depressum). 
NUMBER 45 25 

C;omtnarianas (Fonte), hills N of hlt. .\lutom, 175 m, Fosberg 

25fi5 (US, BISH, XI’, L). Rota: Alarche 271 (P, US); Songsong, Fosberg 25111 (US BISH, NY); Sasan Ha)a ba), Sachet 

ii9l (US, BISH). Saipan: Marpi Point, 15 m, Fosberg 47720 

(VS, BISH). Pagan: near landing, Fosberg 31640 (US). 

C.aroline Is.: Clithi Atoll, Falalap Islet, Fosberg 46488 (US). 

Slarshall Islands: Kwajalein Atoll, Kwajalein Islet, Fosberg if124 (VS), fS032 (CS, BISH). Slajuro Atoll, Long Islet, 

Fosbrtg 58810 (CS, BISH). 

IVake Island: IVakc Islet. Fosberg 43537 (US, BISH). 

l’liwnix Islands: Canton Island, Fosberg Q Stoddart 54897 

(US). Hull .Atoll: Forbr.7-g i- Stoddard 54842 (US, HAW), 

557fl (LS, BISH). 

Hawaiian Islands: Oahu, Sand Island, Herbst 5466 (US); 

Kaneohe, 1Iokapii l’eniiisiila, Fosberg 2 Ez~ans 5SS3fA (US). 

AIesico: Baja California: Arroyo San Juan, 30 m, Moran 

L Rcival 2012‘1 (US), Agua \’erde Bay, I. M. Johnston 

?8S3 (US); 17.G km E of San Ignacio, Curter et al. 1970 (US); 

San Jose del (:ah, II’iggins 56SfA (US); Rose 16482 (US). 

Sonora: without locality, Palmer in 1865 (US); San Bernardo, 

Kio lIa\.o, C,e,itq lib’? (US); Sail Pedro Bay, I. M. John- 

\ton $327 (YS). Sinaloa: liazatlan, TV. G. Wrigl~t 1238 (US); 

Ortega 5437 (L.S); Culiacan, Palmer 1506 (US). San Luis 

I’otosi: Xiinas de San Rafael, Piirpus 5ifi (US). Tamaulipas: San \‘icerite, Jaurnaol, von Rozpnski 376 (1;s). Suevo 

Leon: hfonterrey, Pringle 1879 (US); Cerro del Obispado, 

near Slonterrcy, fic?~rnnde: 3 Balkley If 515 (US). Jalisco: 

(;uadalajaro. Pn(itrer 2i8 (US). State not given: Yaqui River, 

Poltrier i?i 186? (LiS); Lodiego, Palmer 1563 (Us). 

VERBEWACEAE 

Callicarpa L. 

The Micronesian plants of this genus are a part 

of a taxonomically almost insoluble complex extending from southern Asia to the eastern Carolines, 

for which the correct name seems to be Callicarpa 

candicans (Burman f.) Hochreutiner, based on 

Urtica candicnns Burman f. Two species have been 

segregated from this in LIicronesia. Callicarpa 

elegans Hayek and Cullicarpa lamii Hosokawa (as 

larnmii). A third, Callicarpa paucinervia Merrill 

seems no more than a variety of Callicarpa candicans. 

T~vo other varieties of this species are separable in 

the Caroline Islands. These dispositions are tentative 

and may be changed when the Philippine and 

hlalesian plants of this affinity are critically studied. 

Tl1e n;inics for tlie JIicronesian plants are given 

here with necessary new combinations. A more intensive study of these plants is in progress but the 

names are needed now for the taxa that seem reasonably definite. 

Callicarpa candicans (Burman f .) Hochreutiner 

Caliicarpa candicans (Burman f.) Hochreutiner, Candollea, 

5:190, 1934. 

L~tica f(o?du(o!) Burmait t, FI Indlcd 297 [I971 1iG8 

Calltcarpa cana L., Mantissa Pl., 2:198, 1771. 

Callicarpa candicans (Burman f.) 

Hochreutiner var. candicans 

Not known from Micronesia, originally described 

from Java. 

Callicarpa candicans var. integrifolk (H. J. Lam) 

Fosberg, new combination 

Collicaipa cana tar. integrifolia H. J. Lam, Verb. Malay 

itch., 74, 1919. 

Caroline Islands. 

Callicarpa candicans var. integrifolia (Lam) 

Fosberg, f. integrifolia 

Common almost throughout the Caroline Archipelago. Some plants are intermediate with the following. 

Callicarpa candicans var. integrifolia f. 

glabriuscula (H. J. Lam) Fosberg, new 

combination 

Callicarpa cana var. integrifolia f. glabriuscula H. J. Lam, 

I‘erb. Xiala! .Arch., 74, 1919 [pro parte, Caroline Islands 

specirncns]. 

Callicarpa e?-ioclotin f. glnbrescew Sioltlenke, 1’111 tologia. 

8:385, 1‘362. 

Tirestern Caroline Islands, occasional. Some plants 

are intermediate with f. integrifolia. 

Cnllicarpa candicans var. pauciizervia (Merrill) 

Fosberg, new combination 

Cailirn?pa purici~iervia Sferrill, Philippine Jour. Sci. Bot., 

Callicnrpa erioclona var. paucinervilz (Merrill) Xioldenke, 

9:134, l:Il+. 

Phytologia, 8:57, 1961. 

XI ar i ;I 11 21 s . \ r ch i pe la go, probably e nde IT] i c. Plants 

approaching this have been recorded as Callicai-pa 

pazicinewia from Palau, but should probably be 

included in var. integrifolia. 
26 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Callicarpa candicans var. ponapensis 

Fosberg, new variety 

Calltra~pu elroclonu sensu Glassrnan 1952 [non Schauer in de 

Candolle, I’rodr., 11:643, 18471. 

Arbuscula foliis valde acuminatis marginibus 

valde dentatis, pagina infra dense arcte albotomentosa. Type: Ponape, Takamatsu 796 (US, 

holotype). 

A shrub with slender, densely light brownish 

stellate-tomentose branches; leaves elliptic to ovateelliptic, apices strongly acuminate, bases decurrent, 

margins strongly dentate with triangular teeth, 

upper surface nearl) glabrous except midrib and 

principal veins, lower surface closely and densely 

white stellate-tomentose. 

Caroline Islands: Ponape, Jokaj, in forest near 

coast, 8 1Iar 1!)36, ~akllmC1tS11 796 (US, holotype). 

Callicarpa lamii Hosokawa 

Callicat-pa elfguns Hayek, Fedde Repert. Nov. Sp., 2:88, 1906. 

Caroline Islands: Palau and Yap, type from the 

Philippines (Cuming 1460). 

Callicarpa lamii Hosokawa 

Callicarpa lamii Hosokawa, Jour. Soc. Trop. Agr., 6:206, 

Callicarpa glabra H. J. Lam, Verb. Malay Arch., 82, 1919 

1934 (as lammii). 

[non Koidzurni, 19181. 

Marianas, endemic, only rather weakly distinct 

from C. candicans, but maintained until further 

field observations can be made. 

ACANTHACEAE 

Pseicderanthemum Kadlkofer 

The species of this genus in the western Pacific 

badly need revision. Two or possibly three are 

natil e in Micronesia, plus two cultivated species 

originating elsewhere. The Ponape plant Pseuderanthemum laxipoiurn may be native or an accidental introduction. The data in the label are not 

very helpful in this respect. The plants native to 

Palau have, up to now, been placed in Pseutieranthemum inclusum Hosokawa. On careful study 

they prove to belong to two distinct species, the 

other of which (P. palauense) is herein described as 

new. The relationship of these to other western 

Pacific species is not clear, but at least superficially 

they seem to be distinct from those of other islands. 

Pseuderaizthemum acuminatissimum (Miquel) 

Benoist 

Pseud eran th em um mum ina t issirnun (Miquel) Benoist in 

Lecomte, FI. (;en. I’lndochine, 4:i21, 1935.-Radlkofer, 

Sitz. Bayer Akad. Wiss. Miinchen. Sat.-Phys., 13:286, 1884. 

Eranthemrrm acuminatissimum Miquel, F1. Ind. Bat., 2:835, 

1858. 

Preridern,illre;,rri~,f \p.. of Glassman, Risllop Sfus. Bull., 209: 

102, 1952. 

Glassman compared his Ponape specimen with P. 

sumatrense (Ridley) Merrill, but the inflorescence 

is far too short and dense for that species. It is a 

much better match for Pseuderanthemum acuminatissimum of Thailand and Indo-China, and may 

tentatively be referred to that species. Radlkofer 

may be said to have published this combination, 

but so much ambiguity surrounds his attempted 

transfer that it seems best to ascribe it to Benoist 

who published it very clearly. 

Pseuderanthemum carruthersii var. 

atropurpureum (Bull) Fosberg 

Pseudeianlhe~niint carruthersii var. atropurpureunt (Bull) 

I’,\eriderant/refriirfri jolriileuse Lindau in Fedde. Repert. Sp. 

Fosberg, Ph, tologia, 5:290, 1955. 

Sov., 13:553, 1915. 

Pseudeianthemum jaluitense seems, from the description, especially that of the red color of leaves 

and flowers and the form of the corolla, to belong 

to the common Pseudeianthemum carruthersii var. 

atropurpureum (Bull) Fosberg. The type, Gibbons 

1068, was most likely destroyed in the bombing of 

Berlin, so this is probably the best disposition of 

this name that will be possible. Also Pseuderanthemum carrutliersii is the only species of the genus 

known from the Marshall Islands, or likely to occur 

there. 

Pseuderanthemum carruthersii var. reticulatum 

(Bull) Fosberg, new combination, new status 

Eranthernum reticulaturn Bull, Gard. Chron., ns., 3:619, 1875. 

This differs from var. carruthersii mainly in the 

pronouncedly ovate leaves with a yellow zone along 
KUMBER 45 27 

the midrib and veins. It is occasionally planted as 

an ornamental. 

Pseuderanthemum laxiflorum (Gray) Hubbard 

Pseiirlera,itlreniui,1 laxi/?orlon (Gray) Hubbard in Bailey, 

Era?tlhemum laxiflorzifn Gray, Proc. Amer. Acad. Sci., 5:349, 

Erantlie,rzu,ti pulchellzrin Hort., Gartenmag., 176, t. 17, 

Pseitderaiitli~,nu,ri ptrlchellum (Hort.) Merrill, Phil. Jour. 

Psettderanthemum bicolor sensu auct. Micr. [non (Schrank) 

Rhodora, 183159, 1916. 

1 86 1-1 862. 

1810 [non Andrews, Bot. Repos., t. 88, 18001. 

Sci. Bat., 7:248, 1912. 

Radlkofer]. 

The widely planted ornamental species, commonly called Pseuderanthemum bicolor, seems to be 

this, Pseuderanthemum bicolor is a wild Philippine species. The taxon Pseuderanthemum pulchellum has been used for this species, but ilts basionym 

is a later honionym and hence illegitimate. 

Pseuderanthemum palauense Fosberg & Sachet, 

new species 

Planta herbacea, foliis ovatis acuminatis, racemis 

terminalibus, bracteis ovatis, sepalis lanceolatis 

ceribus, corolla curvata 2 cm longa, semine diametro 

3-4 mm discoideo faciebus verrucatis margine 

sinuato dentato. 

Herb a few decimeters tall, young parts and 

grooves on 2 sides of stem pilosulous, nodes rather 

prominent; leaves thin, ovate, strongly acuminate, 

obtuse or acute at base, up to 7 X 3 cm, petiole 

slender, about 1 cm long, blade with up to 7 veins 

on a side, these appressed puberulent above, much 

less so on under sides; racemes slender, slightly 

pilosulous, terminal and on small side branchlets, rachis 4-6 cm long, occasionally branched, 

bracts small, ovate, about 1 mm long, subtending 

pedicels 2-3 mm long, dilated upward, occurring 

opposite or subopposite moderately spaced along 

rachis, sepals narrowly lanceolate, 3-4 mm long, free 

almost to base, almost glabrous, sharply pointed; 

corolla about 2 cm long, curved, glabrous without, 

limb in bud 3-4 mm long (buds only seen), style and 

stamens about 3 cm long (only detached ones seen 

intact); fruit glabrous, about 2 cm, beak short; seed 

disk-like, margin scalloped, lobe at one end of scar 

more prominent, faces conspicuously warty, graybrown, 3-4 mm across. 

This is quite distinct from P. inclusum Hosokawa 

in its leaf shape, more slender and lax inflorescences, 

much longer corollas, styles and stamens, and in 

scalloped warty seeds. It seems closest to Philippine 

plants commonly referred to P. bicolor (Schrank) 

Radlkofer, which have predominantly axillary 

cymes, linear subulate bracts and sepals, and entiremargined seeds. The Philippine plants do not seem 

to be the same as the cultivated species known as 

P. bicolor (Schrank) Radlkofer (see above). 

SPECIMESS SEEN.-Caroline Islands: Palau: Peleliu Island, 

in forest, Takanrntclc li6R (BISH, holotvpe); Babeldaob 

Island, Aiekalong, in field, Takamatsu 1681 (BISH) 

RUBIACEAE 

Geophila D. Don 

Geophila repens (L.) Johnston 

Geophila repens (L.) Johnston, Sargentia, 8:281, 1949. 

Koiidrletia repenc L., S,st. Nat., ed. 10:928, 1759. 

Geophila repem var. asiatica (Chamisso & 

Schlechtendal) Fosberg, new combination 

Gmphila ie,iifor,nis iar, asiatica [Rheede ex] Chamisso & 

Schlechtendal, Linnaea, 4:137, 1829 

The Old World populations of Geophila repens 

have pyrenes with flat inner faces, while the New 

World ones have them consistently curved or 

warped. Chamisso and Schlechtendal based their 

variety on a specimen collected by Chamisso on 

Guam, but also referred to Rheede, Hortus (1690:41, 

pl. 21). This seems to be the oldest varietal epithet 

applicable to the Old World variety. 

Hedyotis L. 

Hedyotis costata (Roxburgh) Kurz 

tfedyutis tostata (Roxburgh) Kurz, Jour. Asiatic SOC. Bengal, 

Vprirnacocr costata Roxburgh, Hort. Beng., 10, 1814 [nom. 

43 (2):135, 1876. 

nud.]: F1. Intl., 1:376, 1820. 

The typical form, var. costata, is not known from 

Micronesia (but see discussion below). 
28 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Hedyotis costata var. lutescens (Kanehira) 

Fosberg, new combination, new status 

Hedyolis Iule~cens Kanehira, Trans Nat. Hist. SOC. Formosa, 

2534, 1935. 

This has been reduced outright to Heclyotu 

costata (Roxburgh) Kurz by Hatusima (1936:222), 

and by Kanehira, himself (1936:607). However, the 

much denser conspicuously yellow pubescence seems 

to justify retaining it as a variety. The type locality 

is Aimiriik, Babeldaob, Palau. We have not seen 

this specimen (Kanehira 1980), but Takamatsu 

I512 (BISH) is apparently the same. Some specimens from Hainan and Sumatra may belong here, 

though their habit is more open and the pubescence 

more sparse. 

Hedyotis katiehirae (Hatusima) 

Fosberg, new combination 

Leptopetalum kanehirae Hatusima, Trans. Nat. Hist. SOC. 

Hedyotis kunehiroe Hatusha, Trans. Nat. Hist. SOC. ForFormosa, 26:218, 1936. 

mosa, 26:218, 1936 [in synonymy]. 

This species was placed in Leptopetalum Hooker 

and Arnott by Hatusima. That genus differs from 

Hedyotis L. only in its stamens being inserted at 

the base of the corolla tube, rather than part way 

or all the way up. This seems an inadequate basis 

for maintaining it as a separate genus, hence I 

(Fosberg) am transferring L. knnehiiae to Hedyotis 

L. It is close to Hetlyoti.~ foetidn var. mnriannensis 

(.Vei~~ill) Fo$bcrg. 

The coiiilin;~ tion Hdyo/is kmaelrirne (Hatusima) 

Fosberg was inadvertently pblished by us (Fosberg, 

Falanruw, and Sachet, 1975:40) without its basionym. As a valid name it dates from the present 

pub 1 i ca t i on. 

Hedyotis strigulosa (Bar ding ex de Candolle) 

Fosberg, new combination 

Oldenlandia strigulosa Bartling ex de Candolle, Prodr., 

Hedyolis coreona Leveille in Fedde, Repert. Sp. h’ov., 

Oldenlandio nlbido-punctntn Merrill, Phil. Jour. Sci. Bot. 

Hedyotis olbido-punchto (Merrill) Fosberg, Lloydia, 3:123, 

Bartling’s name apparently applies to this common species of coastal limestone rocks in the western 

4:427, 1830. 

11:64, 1912. 

9:147, 1914. 

1940. 

Pacific. The type is from “Isl. Marianne, Haenke” 

(G, holotype). Hedyotis coreana LkveillC seems most 

probably the same, but is not described in detail. 

Psychotria L. 

The genus Amaracarpus Blume was set up to 

accommodate species (e.g., Amaracarpus muscosus) 

with strikingly unique habit, but little technical 

difference from Psychotria. The inclusion in it of 

numerous species that do not share its distinctive 

character but have smali or slender axillary inflorescences reduces to insignificance its difference from 

Psychotiia. .\ series of these species were described 

from Micronesia by Valeton and by Japanese botanists who followed him. In 1940 I (Fosberg) failed 

to find reason to maintain Amaracarpus for the 

Micronesian plants, and failed, also, to see much 

justification for the plethora of Micronesian Amaracarpus species. I reduced them, summarily, to two 

species of Psychotiin: P. carolinensis and P. ponapensis. Since then, I have been unable to see even 

two good species in the complex and have also 

found that two older binominals, Psyclrotria malaspinae Merrill (1914) and Uragoga hombroniann 

Baillon (1879) have been proposed for members of 

this complex. Field studies and comparison of many 

herbarium specimens indicated that, although many 

of the entities are discernible, they intergrade and 

are not very different at best. The solution, admittedly tentative, but closer to reality than the 

earlier alternatives, seems to be to recognize a number of varieties of the one species, Psychotria hombiwiiann (Baillon) Fosberg. Names are provided for 

these so that they can be referred to, pending the 

intensiye study that will be necessary to elucidate 

a more final taxonomy of the group in Xlicronesia 

and to find its relationships elsewhere. 

Pspchotria hombroniana (Baillon) Fosberg 

Uragogo hombroniuno Baillon, Adansonia, 12:333, 1879. 

Psychotria hornbroniana (Baillon) Fosberg, Phytologia, 5:291, 

1955. 

Found in several varieties throughout the Caroline and Marianas high islands. Coral atolls do not, 

apparently, provide suitable habitats for these 

plants. Synonyms will be listed under the appro- 
NUMBER 45 29 

priate varietal names. Descriptions and citation of 

the abundant specimens will be deferred until the 

treatment of the Rubiaceae in the Flora of Micronesia. 

Psychotria hombroniana (Baillon) 

Fosberg var. hombroniana 

Uragoga iioinbroiiiana Baillon, Adansonia, 12:333, 1879. 

Marianas Islands: known from Rota and Guam. 

Type from Guam, Hombron in 1841 (P, 2 sheets). 

Psychotria hombroniana var. hirtella (Valeton) 

Fosberg, new combination, new status 

.4iiiarata~piis liirtellus Valeton, Bot. Jahrb., 63:320, 1930 

Psi c iiotrin poiiapeiists Fosberg, Bishop Mus. Occ. Pap., 

Ps\r/iotttn trinriop/i>/la Valeton, Bot. Jahrb., 63:318, 1930. 

[non Psyc/cotttn /izttella Oli\er, 18871. 

35.224, 1940. 

Caroline Islands: Ponape and Truk. 

Psychotria hombroniana var. kusaiensis (Kanehira) 

Fosberg, new combination, new status 

..limiarnipiis kusnierisis Kanehira, Bot. Mag. Tokyo, 49:276, 

fig. 26, 20 Ma) 1935. 

.4via,-acarpus kanehirae Hosokawa, Trans, Nat. Hist. Soc. 

Iormcisa, ?5:35. April 1935 [non Psychotria kanehirae 

kleri-ill, 19161. 

The two names in Amaracarpus are obviously 

synonymous and A. kaneliirae is earlier. However, 

selection of the other epithet for transfer to varietal 

status seems less likely to be confusing. 

Caroline Islands: Kusaie. 

Psychotria hombroniana var. malaspinae 

(Merrill) Fosberg, new combination, new status 

F.~),c/iotria iiialaspiiiae Merrill, Phil. Jour. Sci. Bot., 9: 148- 

Atnnrnrnrpirs irinriaii~ie~isis Kanehira, Bot. Mag. Tokyo, 

.-I ii~~i~~i~~~i~~irs Indt-oriicii,! Hosokawa, Trans. Nat. Hist. SOC. 

.i!riniarni.pii! iiia/n\pinae (hIerril1) Kanehira & Hatusima, 

‘-1 o~nrmrf;ir\ rote?isis Hosokawa, Trans. Nat. Hist. Soc. ForI-l!], 1914. 

IR:Wi, 1934. 

Formosa, Y7: 269, 193.5. 

Bot. Mag. TokIo. 50:607. 1936. 

n1oza. 25::(5. 1935. 

Marianas Islands: Alamagan; Rota: Guam. 

Psychotria hombroniana var. squarrosa 

(Valeton) Fosberg, new combination 

Aniaracarpus carolitiensis \ ar. squarrosa Valeton, Bot. Jahrb., 

Ainaracarpus carolinensis Valetosn, Bot. Jahrb., 63:319, 1930. 

Ariiaracc~rpus kmeinei i Valeton, Bot. Jahrb., 63:320, 1930. 

Arrini-atarpus lzetrropoides I’aleton, Bot. Jahrb., 63:320, 1930. 

Psychotria carolinensis (Valeton) Fosberg, Bishop hlus. OCC. 

63:319, 1930. 

Pap. 353224, 1940. 

Caroline Islands: Palau, Truk, Ponape, Kusaie. 

Spermacoce L. 

Spermacoce ernstii Fosberg & Powell, new species 

Planta herbacea diffusa vel erecta ramosa glabra 

vel sparse scabrida, foliis ellipticis acutis vel acuminatis, lobis calycis linearis 1-2 mm longis, 

corolla brevi lobis triangulo-ovatis, pauce barbata, 

antheris stigmateque valde inclusis, capsula hispidula breve late cylindrica, semine late elliptic0 

nititlo castilneo reticulato sulco lentreali lato. 

Type: Columbia: Huila, above Galilea, Fosberg 

I9632 (US, holotype). 

This taxon is published here to validate the name 

so that it can be used. A full treatment with other 

related species is to appear in the nex tutiire Tills 

species has been introduced into Micronesia and 

has spread to a number of islands, in some of which 

it has become common. 

.-llarianas Islands: Saipan: 1.: of Ogso 

Tapotchau. just S of Kannat Tadung Laulau, 170-190 m, 

Fosberg 50545 (US). Rota: Road to Sabana area, 200-400 in, 

Snc/irt /,YUK (1;s); 150-250 m, E7~iii.s 212,’ (US). Guam: 

Dedetlo, 100 in, Fosberg iiili‘) (US); Campana)a Ba), 20 m, 

P’osberg f 3f 1 i (US); Trust Territory Compound, NAS, 

Agana, 70 ni, I.osberg f6215 (US), f6216 (US); Andersen Air 

Forte Base, .Ifornri ff6i (US). Caroline Islands: Palau: 

Kayangel. along trail in middle of main islet, Cheatham 86 

(1’s). Koror: SF, coriicr of island along load tn .Ail.ai Ferry 

dock. Fo\brig 50620 (US). Peliliu: around the S landing, 

1-2 ni. fio,sbrrg ;/9iS ([‘S); N end, ikalokul at boat landing. 2 111. Fojbej-g fi616 (VS). .Angaur. west of Lake D, 0.3 mi 

[O.,; kin] SE of tillage. ‘1 in. Cnii/ie/tl 150 (CS). 

Spermacoce hispida L. 

Spermacoce hispida L., Sp. Pl., 102, 1753. 

Yf~~i~nr~re nrticulu,-is L.f., Suppl. Pl., 119-120, 1781. 

/!or7-?iio liiJpitla (L.) Schumann in Engler & Prantl, Die 

/!o? ierin roltiiidifolia Valeton, not. Jahrb., 63:323, 1930. 

Naturlichen 1’fianzenfamilien, 4(4): 144. 1891. 
30 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

The identity of Borreria rotundifolia has been a 

problem ever since the study of Micronesian botany 

became active after LVorld War 11. There seemed 

libtle chance of solving it, however, since the type 

was probably destroyed in Berlin. The discovery of 

a partial set of duplicates of Ledeimann’s collection, 

the Micronesian portion of which was kindly loaned 

by the authorities of the Berlin Herbarium, has 

enabled us to resolve this and other similar difficulties. The Ledermann collection on which the 

species is based is clearly Spel-macoce hispida L., an 

Asiatic species previously recorded from Micronesia 

from Yap Island on the basis of a Volkens specimen 

(no. 215). This specimen was also destroyed in the 

bombing of Berlin and was probably Spermacoce 

repens. 

SPECIMFX Swx.-Caroline Islands: Ponape. Pailapalap, 

I,ederiiiniiir 13800 (B, lectotype of Borreria rotztndifolia). 

Spermacoce repem (de Candolle) Fosberg & Powell, 

new combination 

/{or, erin reperrs de Candolle, Prodr., 4:542, 1830. 

,S/w nto~occ ocyiuoides sensii auct. [non Burman f., F1. Ind., 

34, 17681. 

Type from Mauritius. 

SITCIVF\S Sr~.\.-Caroline Islands: Palau: Babeldaob 

Island. Me’ebe’nbul billage, Ngatpang, Fosberg 32403 (US). 

Garamiscan Colon\, Foshrrg 25il1 (US). Yap Island: Volkens 

215 (not seen): Leuj (Leuis) village, 10-15 m, Fosberg 

i660f ((IS, BISH), Ponape Island, Classman 2583 (US, BISH): 

Retail-nepiichi, Hosokaztla i5i2 (BISH). 

COMPOSITAE (= ASTERACEAE) 

V‘erizoizia Schreber 

Vernonk patula (Dryander) Merrill 

I’utiotiin p~tulo (Dryander) hferrill, Phil. Jour. Sci. Bot., 

3:439, 11)08: 9: 154. l!)l4.-Koster. Blumea, 1 :433, 1935.- 

Stone, Xlicronesica, 1: 129, 1964; 6:594, 1971. 

C(J?I?X pntrrla Dr)andei in Aiton, Hort. Kew., 3:184, 1789. 

T’eriioiiio cliiriemis sensu Endlicher, Ann. Wien. Mus. 

Saturgesch. I: 168. 183i.-Schumann and Lauterbach, F1. 

Siids., 595, 1901. [Non (L.) Lessing, Linnaea, 6:105, 18311. 

<‘)oufJpiy pi~O~,~~cio tle Candolle. Prodr.. 3:69, 1836.-Safford. 

Contr. I..S. Xat. Hed)., 9:252, 1905 [as Cyanoptis]. 

r’ei?iniirn ?ji/loccr (Blume) \Vight in Safford, Contr. US. Nat. 

Heih.. 9:396, 19U.;.--Stoiie. Rlicronesica, 1:129, 1964. 

I’eriiorria patirln var, pubescens (Blume) Koster, Blumea, 

I :430-43.i, 1935. 

Vernonia patula is an eastern Asiatic-New 

Guinea species which extends to Guam. There it is 

probably introduced, but was found by Haenke in 

1792. It has been reported from Guam by a number 

of avthors under several names, as shown by the 

above synonymy. The correct name is apparently 

Vernonia patula, though this has not been established with certainty until now. Its nomenclature has 

been somewhat confused, as it has been considered 

as sjnonymous with the earlier Vcmonia chinensis 

(L.) Lessing. Thanks to information supplied by 

Dr. Kare Bremer (in litt., 1979), it can be reasonably 

established that the type of Conyza chinensis L., 

basionym of V. chinensis, is the specimen in the 

Linnean Herbarium, London, numbered 993: 22. 

This is marked “Suratt,” which is apparently 

equivalent to Surate, where Toren, the collector of 

the specimen cited by Linnaeus, spent some time 

collecting. From a microfiche photo of this specimen, we can determine positively that it is not the 

plant under discussion, though we are not certain 

what it is. This effectively removes Vernonia chiHCIISIS from consideration in the nomenclature of 

Vernonia patula, which thus remains the correct 

name for the plant found in Guam. 

Wollastoizia de Candolle ex Decaisne 

Il’ollastmin de Candolle ex Decaisne, Nouv. Ann. Mus. 

Ll’ede/ia sensu auct. plur. pro parte [non Jacquin, Enum. 

.Sleinmodontia sensu auct. Micr. [non Cassini, Bull. SOC. 

Paris, 3:414, 1834: Prodr., 5:546-549, 1836. 

PI. Carib., 8, 28, 17601. 

Philom. Paris, 11, 18171. 

Weak shrubs, suffrutescent herbs, and herbs with 

a resinous odor, leaves opposite, tending to be 

trinerved, exstipulate; inflorescences very open, terminal with 3-5 or more long-pedunculate heads or 

heads solitary; involucre of one principal whorl 

with a few smaller outer phyllaries; receptacular 

bracts strongly nerved, obtuse, sometimes apiculate, 

pubescent, enfolding achenes; ray florets mostly 

6-10, yellow, limbs usually slightly 2- or 3-toothed 

at apex, disk corollas yellow or greenish yellow, 

5-lobed; achenes thick, cuneiform, those of ray 

flowers 3-angled, of disk flowers 4-angled, ithose of 

ray flowers convex on outer face, all or mostly well 

developed, underdeveloped disk achenes scattered 

among fully developed ones, pappus of 1 short awn 

(or this absent) and summit of achenes blunt, 

strongly hirtellous, lower part glabrous. 
NUMBER 45 31 

The common Indo-Pacific strand plant usually 

called Wedelia biflora (L.) de Candolle ex Wight 

at first sign usually impresses botanists familiar 

with the Hawaiian flora as being a member of the 

Hawaiian genus Lipochaeta DC (e.g., Degener, in 

litt., 1942). 

Since first seeing W. biflora in the field in Micronesia in 1946, I (Fosberg) have been convinced that 

Lipochaeta and Wedelia biflora were congeneric, 

but refrained from transferring the Hawaiian 

species to Wedelia because I was familiar wi’th too 

few of the American and African species of Wedelia. 

It seemed possible that Wedelia biflora belonged in 

Lipochaeta but other Wedelia species might remain 

as a separate genus Wedelia. 

In 1965, H. Wild (1965:4-5) transferred Wedelia 

biflora and most other African species of Wedelia 

to the genus Melanthera Rohr. This genus is typified by the American Melanthera niuea (L.) Small, 

pro parte, including Rohr’s specimen (Strother, 

1970:336-338), a plant with white ray flowers, and 

not too close to the Pacific species under discussion 

except as all species in the Wedelia relationship are 

similar. The basis for uniting them with Melanthem was the angular achenes characteristic of most 

African species. The exception, Wedelia trilobata, 

was left by Wild as the only African representative 

of Wedelia sensu stricto. He does not characterize 

Wedelia proper, but in correspondence he mentions 

flat achenes. Actually the disk achenes of W. trilobata when fertile and developed are somewhat compressed-clavate, rather than flat. WedeZia biflora 

seems far closer to Lipochaeta than to either Melanthera niuea or Wedelia trilobata, or to what material I have seen of Wedelia fruticosa Jacquin, type 

of Wedelia. 

Many of the genera related to or segregated from 

Verbesina L. seem so close to each other that perhaps the logical course would be to lump them all 

into a single large genus. However, the limits of 

such a genus would likely be as hard to establish 

as are those of the segregates, and only a major revision of the large number of species in this assemblage might give any satisfactory delimitation. 

A number of people are working on parts of this 

complex, and when their results are in and the 

parts of the puzzle are compared on a tropics-wide 

basis, logical and recognizable generic lines may 

emerge. 

Regardless of this, the disposition of Wedelia 

biflora vis-a-vis Lipochaeta remains a problem in 

the Pacific Islands. It seems desirable to fit the IndoPacific species under discussion into the present 

generic arrangement more convincingly than where 

it is now, as a Wedelia. 

Our examination of mature capitula of ?Yedelta 

biflora shows that while it has plump, angular 

achenes, it does not fit Melanthera in that its raj 

flowers do produce mature achenes. Flowers that do 

not mature achenes occur randomly in the heads, 

but the rays are characteristically fertile unlike those 

of Melanthera in Wild’s key (1965: 1). 

Wedelia as typified by Wedelia fruticosa Jacquin 

has plump, but slightly compressed, non-angular disk 

achenes. Wedelia trilobata has only part of the disk 

achenes fertile. Wedelia biflora has thick, angular 

achenes, those of the ray florets with pubescence on 

and near the summits. In this character, as well as 

in its fertile rays and its general appearance, it 

resembled Lipochaeta. However, in the latter, the 

sterile florets are in the center of the disk, rather 

than scattered through it. 

Lipochacta has been recently revised by R. C. 

Gardner (1976b) who admits 25 species showing 

great diversity, especially in vegetative characters, 

and falling into two groups on the basis of four vs. 

five disk corolla lobes, correlated with a chromosome difference. In a published paper, Gardner 

(1976a:384) indicates that the closest relatives of 

Lipochaeta are in the genus Wedelza, especially 

Wedelia biflora, and that Lipochueta might have 

arisen from “an ancestor of W. biflora.” 

Dissection of heads of TVedelia biflora var. 

canescens (Gaudichaud) Fosberg shows a strong 

similarity to Lipochaeta in the involucre of one 

principal series of phyllaries, not sharply distinct 

from receptacular bracts (chaff), with a few gradually shorter ones forming an outer whorl (Melanthera has a 2-3 serate involucre according to IVild, 

1965:48). The ray florets produce fully developed 

achenes that are thick and 3- (or 4-)angled and 

rather truncate and hirtellous at the top, as in 

Lipochaeta. Most of the disk florets produce welldeveloped angular achenes, but some are sterile, 

sporadically distributed in the head and tending to 

be rather peripheral rather than centrally concentrated as in Lipochaeta. The pappus consists of a 

single short but prominent awn (sometimes absent), 

as in some species of Lipochaeta. 

Thus, Wedelia biflora seems very close to Lipo- 
32 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

chaeta and, if the difference between fertile and 

sterile ray florets is important, less close to Melanthem, and if thick angular achenes versus plump 

but somewhat compressed ones is significant, less 

close also to Wedelia. Lumping the three genera 

into a single large genus would seem a not unreasonable possibility. However, at least until the several other Pacific island plants generally referred to 

TYedelia can be carefully studied, a more conservative course might be to transfer Wedelia biflora to 

Lipochaetu, which it strong11 resembles, thus vastly 

extending the range of this genus usually considered 

endemic to Hawaii. 

De Candolle, however, placed Verbesina biflora 

L. in his genus Wollastonia, an earlier genus of a 

small number of Old TYorld species. This, though 

usually combined with Wedelia, seems altogether 

too close to Lipociliaeta de Candolle. We do not find 

that a type has been designated for Wollastonia de 

Candolle in Decaisne. The genus was published by 

Decaisne in his account of Timor plants, ascribed 

to de Candolle as in the forthcoming volume of the 

Prodromus and briefly distinguished from Wedelia 

with just enough description to effect valid publication. Three species were listed, two of them briefly 

characterized, the other, Wollastonia scabriuscula 

de Candolle ex Decaisne, described in detail, and 

the “Observation” u.here the name Wollastonia is 

validated accompanies this species. It seems clear 

enough that this was the species that Decaisne principally had in mind and it is here designated lectot!~ie of tlie genus. It must be noted, however, that 

Decaisne cited Verbrsi77a bifiora Blume (by indirect 

reference Verbesina bifloia L.) in the synonymy of 

Wollastonia scabsiuscuZn, making the latter nomenclatui-ally, as well as taxonomically, a synonym of 

Wollastonia biflora (L.) de Candolle. 

TVollastonia scabriu.xula is maintained by de 

Candolle in his Prodromus treatment, but reported 

from the Marianas, as well as from Timor, India, 

and several places in southeast Asia. The Marianas 

record certainly must have been based on the plant 

treated below as Wollastonia bipom var. canescens. 

This variety differs from var. biflorn in its strongly 

canescent to almost sericeous foliage and shorter 

nchnees. It was originally described by Gaudichaud 

as TYollastonin ci2nescen.r and again as Wollastonin 

uipn tea . In the de Candolle Prodromus Herbarium at 

Geneva are four sheets from Asia and two from 

Timor labeled LVollastoniu scubiiuscula (Gaudichaud) de Candolle. All are exactly the plant generally known as Wedelia bipora (L.) de Candolle. 

The Timor material in the de Candolle herbarium 

must be regarded as syntypes of Wollastonia 

scabriusczila, since the first publication, by Decaisne, 

was in his account of Timor plants, except that by 

automatic typification (ICBN Art. 7, 63) the type 

must be that of Verbesina biflora L. 4s a result, 

Wollastonia de Candolle (1834), typified by the 

plant now usually called Wedelia biflora (L.) de 

Candolle, has two years priority over Lipochaeta 

de Candolle (1836). Wedelia biflora (L.) de Candolle 

(= Wollastonia biflora (L.) de Candolle) cannot be 

transferred to Lipochaeta de Candolle. Any transfer 

must be in the other direction if the two genera 

are combined. 

Our object here is to establish the name of the 

widespread Indo-Pacific strand plant, so common 

in Micronesia. It seems better not to transfer the 

species of Lipochaeta to Wollastonia until Gardner’s 

monograph is published. Then only the species that 

seem good in the light of his work need to be transferred. Ilre are satisfied that, as the genera of the 

Helianthae are now delimited, at least part, and 

probably all, of Lipochaeta is congeneric with 

Wollastoniu. 

Wollastonia biflora (L.) de Candolle 

Ii‘olhtonin hifiorn (L.) de Candolle, Prodr., 5:546, 1836. 

T’erbesina biflora L., Sp. Pl., ed. 2, 1272, 1763. 

IVidely distributed on strand and in lowlands in 

the Indo-Pacific region from East Africa to the 

Marshall Islands, Samoa, Tahiti, and the Cook 

Islands; also found in Rurutu and Rapa, where pssible it is introduced. 

Xlicronesian synonymy and references under var. 

b iflora. 

The species is quite variable but mostly in such 

superficial characters as density of pubescence, size 

and shape of leaves, toothing of margins, number 

of heads in an inflorescence, and toothing of ray 

apices. The achenes vary in size and shape; the size 

difference between the Marianas populations and 

those of the rest of Micronesia seems significant, 

especially when supported by the striking difference 

in pubescence. Hence we are recognizing two varieties of Micronesia, probably others occur in other 

parts of the yast range of the species. 
NUMBER 45 33 

Wollastonia biflora (L.) de Candolle var. biflora 

M’edelia bi/io,n (L.) de Candolle ex TVight, Contr. Bot. Ind., 

Wedelia aristata Lessing, Linnaea, 6: 160, 1831. 

Il’ollastottia uabriitscitla de Candolle ex Decaisne, Nouv. 

Inn. >fits. Paris, 3:414, 1834; Prodr. 5:947, 1836. 

I’erbesina striguIosa Gaudichaud, Bot. Voy. Uranie, 463, 

1826 [1829]. 

Ti’ollntfonin st?igitloso (Gaudichaud) de Candolle ex Decaisiie, Sou\, .Inn. 3111s. Hist. Sat., 3:414, 1834. 

Tl’etielio .ctrigulo.ra (de Candolle) Schumann, Bot. Jahrb., 

!l:223, 1888 (preptint 1887;) [de Candolle cited in error]. 

,Stf>!?i?izodoii/in bifiorcz (L.) TV, F. TVight in Safford, Contr. 

I‘.$. Sat. Herb, 9:3ii, 1905 [non sensu TVight]. 

18, 1834. 

Large sprawling, tangled or even loosely matforming herbs or plants slightly woody below, stems 

glabrous to liispidulous or shortly sparsely hirsute, 

irregularly sulcate when dry, strongly resinous; 

leaves opposite, ovate to broadly ovate, bases acute 

to obtuse or subcordate, slightly decurrent, apices 

acuminate, margins serrate, teeth tending to be uneven, main veins few, 3-5 plinerved appearing 

palmately 3-nerved from just above the base of 

blade, branch nerves anastomosing, network finer 

and finer, both surfaces finely strigose especially on 

veins, upper less so, general appearance green, 

petioles long, rather slender, leaves of sterile shoots 

often much larger and subcordate to cordate; heads 

in loose irregularly corymbiform panicles of several 

to 20 or more, about 2 cm across, phyllai-ies in 2 

series, ovate or ovate-lanceolate, outer subfoliaceous; 

rays yellow, 6-10 or more, subentire or slightly 

toothed at apex, disk corollas with short tube, limb 

funnelform-campanulate to prismatic-campanulate, 

lobes deltoid, ray flowers with deeply bifid styles, 

disk becoming globose at maturity, achenes 2.5-4 

mm long, those of the ray-flowers 3-angled, depressed in the center of the top, those of the disk 

cuneoid, 4angled. 

Known in Xlicronesia. From tlie Carolines: Palau, 

Sonsoral, Yap, Ulithi, Fais, Eauripik, TVoleai, 

Faraulap, Ifaluk, Lamotrek, Satawal, Puluwat, 

Namonuito, Murilo, Nomwin, Truk, Nama, Losap, 

Namoluk, Etal, Lukunor, Satawan, Nukuoro, 

Kapingamarangi, Ant, Ponape, Mokil, Pingelap, 

Kusaie. 1;rom the ~larshalls: Enitvetok, Bikini, 

Ailinginae, Rongelap, Ujelang, Ujae, TYotho, Lae, 

Kwajalein, Ailuk, Likiep, Aur, Ailinglapalap, 

Xlajuro, hno, Namorik, Jaluit. From the Gilberts: 

.4 b a i a n g . Detailed citation of Micronesian specimens as 

well as ethnobotanical information will appear in 

the Flora of Micronesia treatment of the Compositae (Fosberg and Sachet, in press). 

Wollastonia biflora var. canewens (Gaudichaud) 

Fosberg, new combination 

Yerbesinn ca?ie.trrit\ Gaudichaud, Bot. Voy. Uranie, 463, 

18% [1829].--Safford, (:ontr. I’.S. Sat. Herb.. 9:3‘33, 1905. 

f.evbesina nrge7itP.n Gaudichaud, Rot. Voy. Uranie, 463, 1826 

TVollmto~iia canescens (Caudichautl) de Candolle, Prodr., 

.5:947, 1836. 

S/eiitr?iodoritin cnnescens (Gaudichaud) TV. F. ITight in Safford, Contr. U.S. Sat. Herb., 9:3i;’, 190.5. 

Il’etlelia cane.rcens (Gaudichaud) Merrill, Phil. Jour. Sci. 

llot.. 9:lij, 1914. 

Wedelin nrgentea (Gauclichaud) Metrill, Phil. Jour. Sci. Bot., 

9:l5,5, 1914. 

Il’<~tir~lin biflo~n xai’. cnnrscens (Gaudichaud) Fosberg, Phy. 

tologia, 3:291, I9~5i.-Fosberg. Falanriiw, and Sachet. Smithsonian Contr. Rot., 22:43, 1975. 

Sle?ii~iiodoiitin bilioro sensu Safford, Contr. C.S. Sat. Herb., 

9:377, 1W3 [noii (L.) TI‘. F. TI‘ight in Safford, Contr. U.S. 

Sat. Herb., 9:371, 190.51, 

f’erbe,\inn bi//orn semu Safford, Contr. C.S. Nat. Herb., 93399, 

1905 [non L., Sp. 1’1. etl. 2, 1272, 17631. 

Il’edeiin bi/7orri sensu Saffoi-(1, Contr. U.S. Sat. Herb., 9:398, 

1903 [non (L.) de Candolle in TVight, Contr. Bot. Ind., 18 

183.11. 

Il’oiicrctofiin bi(lo?o senrii Safforti, Contr. U.S. Nat. Herb., 

. I905 [non (L.) tle Candolle ex \\light, Contr. Bot. 

[ 18291. 

ln:l., 18. 18341. 

Differs from var. biflora in being vegetatively 

much more densely strigose and in its shorter, 1.5-2 

(-3) mm long achenes. The density of the pubescence varies considerably. .\ctually, specimens can 

be selected exhibiting all degrees of hairiness from 

the silvery strigose plant described as Veibesina 

argcntea Gaudichaud to the greenest extreme of var. 

bifiora, perhaps a specimen (St. John cb Mail-eau 

15464 (US)) from Rapa, at the southern extreme of 

the range of the species. However, in general, the 

Marianas populations are more densely strigose 

and all that we have studied in detail have small 

achenes, so we are continuing to maintain this 

variety. Examination of three collections from the 

Bonin Islands, kindly sent by Prof. Momiyama, 

Hahajima (Solinza et nl. 715173 (MAK); Ono 6 

Kobaynslii in 19il (MAK); ilfomiyamn et al. in 

1972 (hl.%K)), shows that, though they are not conspicuously more hairy, the one mature achene present is 2.5 mm long. These plants, should probabl), 
34 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

be associated with var. canescens. Far to the south, 

on the island of Rarotonga, Cook Islands, a specimen (Stoddart 2019 (US)) has likewise very small 

achenes, about 2 mm long, though vegetatively the 

plant is not unusually hairy. Further mature fruiting 

material is needed to make a firm disposition of this 

population. Flowering specimens seem to have unusually short ray florets. 

In addition, two collections from Palau (Babeldaob, Hosokawa 7026 (BISH) and Angaur, Fosberg 

25917 (US, BISH)) seem to belong here, as they are 

markedly canescent and have rather small achenes. 

A further discussion of var. canescens may be found 

in Fosberg (19553291). 

Known in Micronesia. From the Marianas: Maug, 

Agrigan, Pagan, Alamagan, Guguan, Sarigan, 

Anatahan, Medinilla, Saipan, Tinian, Rota, Guam. 

From the Carolines: Palau: Babeldaob and Angaur. 

Detailed citation of specimens and ethnobotanical 

information will appear in the Flora of Micronesia 

treatment of the Compositae (Fosberg and Sachet, 

in press). 
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.4calypha, 1, 7 

amboynensis, 8 

amentacea, 7, 8, 10 

amentacea ssp. amentacea, 8, 11 

amentacea var. amentacea, 8 

amentacea var, grandis, 8, 10 

amentacea var, heterotricha, 9 

amentacea var. palauensis, 9, 10 

amentacea var. trukensis, 10 

amentacea varieties, 11 

amentacea var, velutina, 10 

amentacea f. wilkesiana, 11 

amentacea ssp. wilkesiana, 9, 10 

amentacea ssp. wilkesiana f. circinata, 11 

angatensis, 7, 10 

betulina, 7 

cardioph:lla, 11 

cardioptillla var. cardiophylla, 11, 12 

cardiopliLlla Lar. ponapensis, 11 

caturus, 11 

fruticma, 7, 9 

grandis, 7, 8, 9, 10, 11 

grandis \ ar. amboynensis, 8 

grandis xar. velutina, 10 

ponapensis, 11 

spicigera, 11 

stipulacea, 7, 8 

tomentosa, 10 

trukensis, 10 

wilkesiana, 7, 8, 10, 11 

Acanthaceae, 26 

Amaracarpus, 28 

carolinianus, 29 

heteropoides, 29 

hirtellus, 29 

kanehirae. 29 

kiisaiensis, 29 

kraemeri. 29 

ladronicus, 29 

inalaspinae, 29 

mariannensis, 29 

miiscosiis, 28 

rotensis, 29 

,411 t tiocleist a, 1 R 

Araceae, .5 

.Araliaceae, 15 

Bar1 ingtonia a3iatica. 19 

Roe1 lagiodendron, 15 

mariannense, 16 

pachrphjllum, 16 

piilclierrimum, 16 

truncatum. 16 

37 

Boraginaceae, 23 

Borreria hispida, 29 

repens, 30 

rotundifolia, 29, 30 

Brtiguiera, 1, 15 

Calliandra, 1, 6 

haematocephala, 6 

haematomma, 6 

inaequilatera, 6 

cana, 25 

cana xar. integrifolia, 25 

cana var. integrifolia f. glabriuscula, 25 

candicans, 25, 26 

candicans \ar. candicans, 25 

candicans tar, integrifolia, 25 

cancticans var. integrifolia f. glabriuscula, 25 

candicans 1 ar. integrifolia f. integrifolia, 25 

candicans ~ar. paucinervia, 25 

candicans tar. ponapensis, 26 

elegans, 26 

erioclona, 26 

erioclona f. glabrescens, 25 

erioclona var. paucinerbia, 25 

glabra, 26 

lamii, 26 

lammii, 26 

paucinervia, 25 

Calophrllum, 1, 12 

cholobtaches, 12, 13 

inophyllum, 12 

inophyllum var. wakamatsui, 12 

pelewensis, 12, 13 

soulattri, 12, 13 

wakamatsui, 12 

Carissa grandis, 20 

Celastraceae, 12 

Cenchriis lappaceus, 5 

Centosteca, 1, 5 

lappacea, 5 

Centotheca, 5 

latifolia, .i 

Chorisepalum, 18 

Christella carolinensis, 4 

rupicola, 5 

Chr!soph)llum sp., 17 

Citriillus, 1, 15 

caffer, 15 

lanatus, 15 

lanatus var, caffer, 15 

lanatus Iar. caffrorum, 15 

vulgaris var. caffrorum, 15 

Callicarpa 1, 25 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

macrantha Yar. palauica, 12 

Hippocrateaceae, 12 

Holcus latifolius, 5 

Hvdrilla, 1, 5 

verticillata, 5 

Hydrocharitaceae, 5 

Hymenophyllaceae, 1 

Clusiaceae, 12 

Compositae, 30 

Conyza chinensis, 30 

patula, 30 

Cucurbita caffra, 15 

Curbitaceae, 15 

CIanopis pubescens, 30 

Cyclosorus jaculosus, 4 

Dicranopteris, 4 

linearis lar. latiloba, 4 

Drpopteris carolinensis, 4 

immersa, 4 

rupicola, 5 

Fgeria, 5 

densa, 5 

Elatostema, 1, 6 

calcareum, 6 

diIaricatum, 6 

sessile, 6 

stenoph\llum, 6 

Epipremnum, 6 

carolinense, 5, 6 

palauense, 6 

laxiflorum, 27 

pulchellum, 27 

reticulatum, 26 

Euphorbiaceae, 7 

Eustoma, 18 

Eranthemum acuminatissum, 26 

Fabaceae, 6 

Fagraea, I, 18 

berteriana, 19 

berteriana lar. berteriana, 20 

berteriana var. galilai, 20,21 

berteriana var. kusaiana, 20 

berteriana var. ladronica, 20 

berteriana iar. pogas, 20,21 

berteriana var. sair, 20, 21 

galilai, 20 

grandis, 20 

ksid, 19, 21 

kusaiana, 20 

morindifolia, 19 

sair, 21 

sair Tar pogas, 21 

schlechteri, 20 

Flacourtia, 1, 14 

rukam, 14 

riikam lar. micronesica, 14 

Flacourtiaceae. 14 

Garcinia, 1, 13 

ponapensis, 13 

ponapensis var. trukensis, 13 

ritmijo, 14 

rumi\o \ar calcicola, 14 

rumiyo var. rumiyo, 14 

trukensis, 13 

Geniostoma, 1, 21,22 

glaberrima, 22 

hoeferi, 21, 22, 23 

hoeferi var. glabrum, 21, 23 

longistylum, 22, 23 

micranthum, 21, 22 

michanthum bar, hoeferi, 23 

micranthum var. micranthum, 23 

micranthum var. paganense, 23 

rupestre 

saipanense, 22 

Gentianaceae, 18 

Gentianothamnus, 18 

Ceophila, 1,27 

reniformis var. asiatica, 2i 

repens, 27 

repens var. asiatica, 27 

Glaphpropteris palauensis, 4 

rupicola, 5 

Gleicheniaceae, 1 

Gleichenia, 1, 4 

Gnetaceae, 4 

Gnetum, 1,5 

gnemon, 5 

Gramineae, 5 

Guettarda, 19 

speciosa, 19 

Guttiferae, 12 

linearis var. latiloba, 4 
SUhlBER 45 39 

Hvmenophyllum, 1, 4 

digitatum, 1 

Langeveldia, 6 

Leguminosae, 6 

Leptopetalum, 28 

kanehirae, 28 

Lipocliaeta, 31, 32 

I> i gi an t 21 11 s, 18 

Loesineriella, 1, 12 

macrantha, 12 

macrantha \ar. macrantha, 12 

macrantha var. palauica, 12 

Lmga n iaceae, 2 1 

Macrocarpaea, 18 

Medinilla, 1, I5 

niedinilliana, 15 

rosea, 1.5 

\Ielantliera, 31, 32 

nivea. 31 

\felo~hria. 15 

giiarnensis. 15 

Xfelastoma, 1, 15 

ma laba t h ricum , 15 

malal~athricurn t’ar. marianniim 

rnariannuni, 1.5 

niedinilliana. I5 

\felastoma taceae, 1.5 

Microgoniurn falsinervulosum, 4 

tahitense, 4 

Xlicrotrichomanes, 4 

digitatrim, 1 

Xfvrsinaceae, 17 

Mynine, 1, 17 

carolinensis, 17 

ledermannii, 1 i 

palauensis. 17 

Olden la ntlia a1 h ido- punctata , 28 

strigulosa, 28 

Osmoxylon, 1, 15 

rn a r ia n neuse , 1 6 

oli\eri, 16 

p~~lcher~-irn~~rn, 16 

truncatum. Ifi 

pachvpl~vllllrn, 16 

Pellionia. 6 

cli\aricata, 6 

Phaleria nisidai, 5 

I’lanclionella, 1 7 

calcarea. 17 

rnicronesica. 17 

Poaceae, 5 

Poljpodiaceae. sensu lato, 4 

Potalia, 18 

amara. 18 

Potaliae, 18 

I’outeria, 1, 17 

calcarea, 17 

micronesica, 17 

obovata, 17 

Procris divaricata, 6 

Pseuderanthemum, 1, 26 

acuminatissimum, 26 

bicolor, 27 

carruthersii var, atropurpureum, 26 

carruthersii var, carruthersii, 26 

carruthersii var, reticulatum, 26 

inclusum 26, 27 

jaluitense, 26 

laxiflorum, 26, 27 

palauense, 26, 27 

pulchellum, 27 

sumatrense, 26 

l’s?.chotria, 1, 28 

carolinensis, 28, 29 

hirtella, 29 

hombroniana, 28 

hombroniana var. hirtella, 29 

hombroniana var. hombroniana, 20 

hombi oniana ar. kirsaiensis, 21) 

hombroniana \ ar. malaspinae, 29 

hornbroriiana tar. squarrosa, 29 

niacroph\lla, 29 

malaspinae, 28, 29 

ponapensis, 28, 29 

Rapanea, 17 

carolinensis, 17 

ledermannii, 17 

palauensis, 17 

Rhaphidophora, 1, 5 

carolinensis, 5 

koidzumii, 6 

palauensis, 6 

Rhizophoraceae, 15 

Rondeletia repens, 27 

Rubiaceae, 27 

Rusbyanthus, 18 

Sapotaceae, 17 

Serpicula t.erticillata, 5 

Sideroxjlon, 17 

calcareum, 1 7 

micronesicurn, 17 

Spermacoce, 1, 29 

articularis, 29 

costata, 27, 28 

ernstii, 29 

hispida. 29, 30 

ocrmoides, 30 

repens, 30 

Stemmodontia, 30 

biflora, 33 

canescens, 33 

Symbolanthus, 18 
40 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Stxlosanthes, 1, 6 

guianensis, i 

guJannensis, i 

humilis, 7 

sundaica, i 

Fundaicus, 6, i 

rachia, 18 

guianensis, 18 

Tachiadenus, 18 

Tachiineae, 18 

l'helypteris, 1, 4 

carolinensis, 4 

jaculosus, 4 

peliliuensis, 4 

rupicola, 4 

rupi-insularis, 4 

subpubescens, 4 

I.richomanes, 1, 4 

craspedoneurum, 4 

digitaturn, 1 

falsinen ulosum, 4 

omphalodes, 1 

tahitense, 4 

guyannense, 7 

Trifolium guianense, 7 

L'ragoga hombroniana, 28, 29 

Crticaceae, 6 

t'rtica candicans, 25 

L'erbenaceae, 25 

Verbesina, 31 

argentea, 33 

biflora, 32, 33 

canescens, 33 

strigulosa, 33 

Vernonia, 1, 30 

chinensis, 30 

patula, 30 

patula var. pubescens, 30 

1 illosa, 30 

IVedelia, 30, 31, 32 

aigentea, 33 

aristata, 33 

biflora, 31, 32, 33 

biflora \ar. canescens, 31, 33, 34 

canescens, 33 

fruticosa, 31 

strigulosa, 33 

trilobata, 31 

argentea, 32 

hiflora, 32, 33 

hiflora lar. biflora, 32, 33 

hiflora lar. canescans, 32 

canescens. 32, 33 

scabriuscula, 32, 33 

strigulosa. 33 

Lehneria, 1, 15 

il'ollastonia, 1, 30, 32 

guamensis, 15