american samoa - University of Hawaii at Manoa

COOPERATIVE NATIONAL PARK RESOURCES STUDIES UNIT 

UNIVERSITY OF HAWAII AT MANOA 

Department of Botany 

3190 Maile Way 

Honolulu, Hawaii 96822 

(808) 956-8218 

Technical Report 87 

BOTANICAL INVENTORY OF THE PROPOSED 

TUTUILA AND OFU UNITS OF THE NATIONAL PARK 

OF AMERICAN SAMOA 

Dr. W. Arthur Whistler 

Department of Botany 

University of Hawai'i at Manoa, 

Honolulu, HI 96822 

Cooperative Agreement 

CA 8034-2-0001 

April 1994

. . 

REPUBLIC 

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AUSTRALIA 

INWEALTH OF 

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. 

Tropic of 

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Samoa 

NIUE ' 1;. 

motional I ' Line I 

ISLANDS OF SAMOA 

0 50 100 

L 

MILES 

I 

'UPOLU I OLOSEGA 

SOCIETY 

,. ISLANiS 

ROSE 0 

American Samoa ATOLL 

ISLANDS 

Figure 1 

Location Map 

THE PACIFIC OCEAN

TABLE OF CONTENTS 

I . INTRODUCTION .................................................................................................................................. 1 

The Geography .................................................................................................................................. 1 

.. 

I hc Climate ........................... . ..................................................................................................... I 

The Geology ...................................................................................................................................... 1 

Floristic Studies on Tutuila ............................................................................................................... 4 

Vegetation Studies on Tutuila ........................................................................................................... 8 

11 . METHODOLOGY ................... . ................................................................................................. 9 

I11 . VEGETATION OF THE PARK ........................................................................................................... 10 

Primary Vegetation ....................................................................................................................... 10 

A . Littoral Vegetation ................................................................................................................. 10 

1 . Herbaceous Strand ................................... . ..................................................................... 11 

2 . Littoral Shrubland ......................................................................................................... 16 

3 . Littoral Forest .................................................................................................................. 16 

B . Wetlands ................................................................................................................................ 18 

4 . Coastal Marsh ................................................................................................................... 20 

C . Rainforest ......................... . ................................................................................................. 20 

Variation within the Rainforest ............................................................................................. 20 

Characteristics of Rainforest ................................................................................................. 22 

5 . Lowland Forest ................................................................................................................. 23 

a . Valley Lowland Forest ....................... . .............................................................. 23 

b . Coastal Lowland Forest .............................................................................................. 26 

c . Ridge Lowland Forest ................................................................................................. 27 

6 . Montane Forest ............................................................................................................... 30 

D . Summit Vegetation ................................................................................................................ 34 

7 . Montane Scrub ................................................................................................................ 34 

Secondary Vegetation ...................................................................................................................... 36 

8 . Managed Land Vegetation .............................................................................................. 36 

9 . Secondary Scrub ................... . .................................................................................... 38 

10 . Secondary Forest ........................................................................................................... 39 

IV . DISCUSSION AND RECOMMENDATIONS .................................................................................... 40 

V . ACKNOWLEDGEMENTS .................... . ..................................................................................... 42 

V1 . LITERATURE CITED ........................... . ........................................................................................ 43 

APPENDIX A: ANNOTATED CHECKLIST OF THE FLORA OF TUTUILA .............................................. 45 

APPENDIX B: PLOT DATA ........................................................................................................................... 108 

APPENDIX C: SEASONALITLY OF IMPORTANT FRUIT-BEARING TREES AND SHRUBS IN THE 

PARK AREA ...................................................................................................................... 120 

APPENDIX D: THE OFU UNIT OF THE NATIONAL PARK OF AMERICAN SAMOA ............................ 125 

APPENDIX E: PLANTS AS INDICATORS OF PAST LAND USE .............................................................. 129 

APPENDIX F: SUMMARY OF THE VASCULAR FLORA OF THE TUTUILA UNIT OF THE 

NATIONAL PARK OF AMERICAN SAMOA ................................................................... 132

LIST OF FIGURES 

1 . Map of the Pacific showing the location of Samoa ........................................................... Frontispiece 

2 . Map of Samoa ...................................................................................................................................... 2 

3 . Vegetation map of the Tutuila park ...................................................................................................... 3 

4 . Pago Pago Bay, with Mt . Matafao in the distance .................................................................................. 5 

5 . Mt . Pioa, a trachyte plug ......................................................................................................................... 5 

6 . I'olauta Ridge and Pola Islet, Vatia ......................................................................................................... 6 

7 . Amalau Valley .................................................................................................................................. 6 

8 . Amalau beach ...................................................................................................................................... 12 

................................................................. 12 

9 . Coral rubble beach north of Vatia; Pola Islet in the distance 

10 . Tuafanua Beach west of Vatia ............................................................................................................... 13 

. 

1 1 Tafeu Cove west of Vatia ........................... . .................................................................................... 13 

12 . I, &oral shrubland vegetation on windswept Sauma Point .................................................................... 15 

13 . Littoral shrubland vegetation on Polauta Ridge ................................................................................... 15 

14 . Littoral shrubland vegetation on Pola Islet .......................................................................................... 17 

............................................................................... 17 

15 . Aiga Bay, west of and outside of the Tutuila park 

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

16 . Littoral forest north of Vatia 19 

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

17 . Vatia coastal marsh 19 

18 . Streamside vegetation at Amalau Valley ........................................................................................... 24 

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

. ..................................................................... 26 

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

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

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

19 . Coastal lowland forest near Tafeu Cove 

20 . Disturbed lowland forest on ridges north of Mt Alava 

26 

21 . Ridge lowland forest west of Amalau Valley 29 

22 . Montane forest on Alava Ridge 29 

23 . Montane forest on Toa Ridge 31 

24 . Lomagramma cordipinna dominating floor of montane forest ........................................................ 33 

25 . Mauga-0-Tula Ridge behind Vatia .................................................................................................... 35 

26 . Montane scrub vegetation on Mauga-o-Tula, Vatia ............................................................................ 35 

27 . Vatia Village showing extent of managed land vegetation .................................................................... 37

LEGEND B 

\ PROPOSED PARK AREA 

OFU ISLAND f-3 

OLOSEGA 

ISLAND 

AMERICAN SAMOA 

Manu'a Islands 

0 1 

MILES 

AUNU'U 

ISLAND 


Tutuila and Aunu'u Islands 

0 2 3 4 

LEGEND 

PROPOSED PARK AREA 

TA'U ISLAND 

figure 2. Major islands of American Samoa and proposed park units

I. INTRODUCTION 

The Geography 

Samoa is a volcanic archipelago situated in the 

south I'acitic Ocean at a latitude of 13 - 15" south 

and a longitude of 168 - 173" west, and runs in a 

west-northwest direction east of Fiji, north of Tonga, 

south of Tokelau, and west of Niue and the Cook 

Islands (Fig. 1). It comprises nine inhabited 

volcanic islands and two distant atolls, and has a 

total area of ca. 3 100 km2. The archipelago is 

divided politically into Western Samoa, which is an 

independent country, and American Samoa, which is 

an unincorporated territory of the United States. 

American Samoa, which comprises the east end 

of the archipelago, consists of five main volcanic 

islands (Tutuila, 'Aunu'u, Ofu, Olosega, and Ta'u) 

and two atolls (Rose and Swains). Tutuila, on which 

this unit of the new proposed park is located (other 

units are on Ofu and Ta'u), is the westernmost and 

by far the largest of the islands of American Samoa. 

It has an area of approximately 142 km2 and a 

maximum elevation of 653 m on top of Matafao 

(Amerson et a/. 1982). 

The proposed Tutuila unit of the National Park of 

American Samoa is located on the north central part 

of the island, between the villages of Fagasa on the 

west and Afono on the east (Fig. 2 & 3). The village 

of Vatia is situated in the center of the park, and the 

boundaries around it begin at about the 200 ft 

elevational line. The northern boundary is the 

I'acific Ocean, and the southern boundary is the 

slope leading from Pago Pago Harbor up to Alava 

Ridge. The highest elevation in the Tutuila unit of 

the park is Mt. Alava at 490 m (1610 ft), where a 

TV tower is located, and the total area of the park is 

2,770 acres (National Park Service 1988). The Ofu 

unit is discussed in Appendix D. 

The CIimate 

Since American Samoa comprises small islands 

situated between the Tropic of Capricorn and the 

Equator, its climate is maritime and tropical. It 

belongs to Koeppen's "tropical climate zone" 

characterized by average monthly temperatures of 

more than 1 8"C, no winter season, and abundant 

annual rainfall that exceeds annual evaporation. The 

mean daily temperature for Tutuila is 26.7"C (80°F), 

with an average maximum daily of 29.7"C (85.4"F) 

and an average minimum daily of 23.6"C (74.6"F). 

There is not much seasonal variation, with the 

warmest month being March (27.2"C, 80.9OF) and 

the coolest being August (25.9"C, 78.7"F) 

(Nakamura 1984). 

The rainfall at the airport at Tafuna on the 

south-central coast of Tutuila averages about 3200 

mm (124 in) per year (Nakamura 1984). The 

monthly average there from October to May is 300 

mm (12 in), while in the dry season from June to 

September it is 180 mm (7 in). This variation is 

strongly affected by the tradewinds, which blow 

regularly from the southeast from about April to 

December. The months when they are absent are 

characterized by storms coming in from other 

directions. 

Topographic features strongly effect these totals 

in some places. For example, Pago Pago, which lies 

just to the west of "Rainmaker" (Matafao) mountain, 

averages nearly 5000 mm (200 in) a year. There is 

probably not too much of a rainshadow effect, since 

the island is not very high in elevation, but more 

rain would be expected to fall on the south and east 

coasts than on the north and west coasts, as in 

Western Samoa (Wright 1963). 

The relative humidity is constantly high, ranging 

between 82 and 86% (at 5 p.m.) throughout the year 

(Amerson et al. 1982). Droughts of varying duration 

occasionally occur, but these do not have much 

permanent effect on the vegetation. Hurricanes, the 

most recent of which were in 1987 (known as Tusi), 

1990 (Ofa), and 1991 (Val), periodically hit the 

islands, but the occurrence of three strong hurricanes 

in such close proximity is unusual. 

The Geology 

Samoa is a volcanic archipelago that is "oceanic" 

in origin, i.e., it was formed from basalt rising from 

a "hot spot" on the ocean floor of the Pacific plate 

beyond the continental islands to the west. The 

archipelago was born in isolation, and has never had 

a connection to any other land area. The islands 

originated in the Pliocene Epoch (over two million 

years ago), and were formed generally in a westerly 

direction, with the youngest on the west end of the 

chain (Savai'i in Western Samoa) and the oldest on 

the east end (Rose Atoll). The most recent volcanic 

activity in American Samoa was a submarine 

eruption between Ta'u and Olosega in 1866. The 

only signs of geologically recent volcanism on 

Tutuila are the Tafuna Plain, and several adjacent

Figure 3 

VEGETATION TYPE MAP 

Tutuila Unit 

National Park of American Samoa 

m 

LEGEND VEGETATION NPES 

MATAPAI POINT 

= 

PROPOSED PARK MU LOWLAND FORE- 

LHSTURBED VEGETATION 

nEW WORK CONDUCTED tM DR MT 

WHISTLER. 1993 

LITTOR4 VEGETATION 

MONTANE FOREST 

MONTANE SCRUB

tuff and cinder cones, including the island of 

'Aunu'u, but these eruptions are thousands of years 

old. There are no signs of recent volcanism within 

the boundaries of the proposed Tutuila park unit. 

Tutuila was created in a series of volcanic 

eruptions forming an island elongated in a east- west 

direction. 'This was followed by stream erosion and 

submergence, which has resulted in a rugged terrain 

and deeply embayed coastline. The most salient 

features of the island are the well-sheltered Pago 

Pago Bay (Fig. 4), the mountain peaks, such as Pioa 

(Fig. 5), otherwise known as "Rainmaker 

Mountain," and Matafao (see Fig. 4), the steep 

ridges ascending into the interior of the island, and 

the cliff-bound Tafuna plain and its associated 

craters. 

Pago Pago Harbor, which nearly bisects the island 

and lies just beyond the southern edge of the park, 

was formed in an ancient caldera that was bounded 

on the east by Pioa, on the north by Alava Ridge, on 

the west by Matafao, and on the south by a now 

non-existent wall outside the mouth of the present 

day harbor. After the volcanic series ended, erosion 

of the center of the caldera and drowning of the 

valley produced the spectacular present day harbor 

that parallels the southern slope of Alava Ridge 

(maximum elevation 490 m) which marks the 

southern boundary of the proposed park. 

Between the east and west ends of the park, 

which are marked by Afono and Fagasa villages, 

rcspectivcly, are a series of steep-sided ridges and 

stream valleys that radiate out from Alava Ridge and 

cxtend down to the north coast of the island. The 

most spectacular of these is the one at Vatia that 

extends out to a steep-sided peninsula (Polauta) and 

Pola Islet (Fig. 6). With the exception of Vatia 

village, the north coast of the park is rugged and 

uninhabited. Most of these stream valleys are 

narrow and steep-sided, but Amalau Valley, with its 

scenic cliffs and waterfalls, is a short, amphitheater 

valley (Fig. 7). The whole coastline of the park is 

steep and rugged, and the only sandy beaches are 

found in Vatia village, Amalau Bay, and on the west 

side of Polauta. The park, consequently, is formed 

mostly by the caldera north wall and the eroded 

northern slopes of the ancient volcano. 

South of both ends and beyond the boundaries of 

the park are the two previously noted peaks, Pioa 

and Matafao, which are classified as trachyte 

plugs-cores of ancient volcanoes that remain after 

the crater rims have eroded away. Eight of these 

trachyte plugs are found on Tutuila (Steams 1944), 

the largest two of which are Pioa and Matafao. 

These plugs are comprised of a fine-grained, highly 

eroded, cream-colored trachyte rock that is chara- 

cterized by a predominance of potassium feldspar. 

The light color of the soil is evident in the cliffs of 

Pioa (see Fig. 5). This soil is generally low in 

important minerals, which results in a stunted, 

scrubby vegetation called montane scrub (see the 

discussion on plant communties below) on the 

slopes and summits of these trachyte plugs. 

Although these two peaks are outside the park 

boundaries, a trachyte plug ridge of equal size but 

lower elevation, forms the spectacular Pola Islet and 

Polauta Ridge just north of Vatia village. 

Floristic Studies on Tutuila 

The first collections of plant specimens from 

Tutuila were made during the visit of the United 

States Exploring Expedition (USEE) to Samoa in 

1839. The specimens, collected by W. Bracken- 

ridge, W. Rich, and C. Pickering, are now stored at 

the Smithsonian Institution and the Gray Herbarium, 

with some duplicates at other institutions such as 

Kew. Unfortunately, the specimens were poorly 

curated, and mistakes in locality are not uncommon. 

In fact, some specimens of endemic Samoan plants 

were incorrectly labeled as having been collected in 

Tahiti. Even the correctly labeled USEE specimens 

cite only "Samoa" as the locality, so it is not certain 

which ones were collected from Tutuila -although 

there is some indication from the published list of 

USEE specimens (Pickering 1876). 

The next plant collector to visit Tutuila was 

apparently the Rev. T. Powell, an amateur English 

botanist employed as a missionary by the London 

Missionary Society in Samoa (ca. 1862-1885). 

Powell collected about 630 numbered specimens in 

Samoa, and many others that are unnumbered. 

Unfortunately, most of the specimens lack localities, 

so it is impossible to determine which ones were 

collected on Tutuila and which ones on the other 

islands (he is known to have collected on Savai'i,

Figure 4. Pago Pago Bay, with Mt. Matafao in the distance. 

Figure 5. Mt. Pioa, a trachyte plug.

Figure 6. Polauta Ridge and Pola Islet, Vatia. 

Figure 7. Amalau Valley. 

6

'IJpolu, and 'Ta'u as well). The only relevant 

publication by Powell was a list of Samoan plant 

names (Powell 1868), but there is hardly any 

mention of Tutuila in this publication. Pickering 

(1 876) often noted island localities in his descrip- 

tions of the USEE collections, but many of the 

listed specimens are unidentified and it is usually 

impossible to correlate the specimens with an island. 

Powell's specimens are stored at Kew in London, 

where they were sought and listed during an 

extensive herbarium search by the author (October 

1992). 

Another collector is known to have gathered 

specimens on Tutuila at about the time-Dr. E. 

Graeffe, a Swiss physician who traveled extensively 

in the region in the 1860s and 1870s. Unfortunately, 

many of his specimens, which are stored principally 

at Hamburg, with duplicates at many herbaria, lack 

specific localities. Several other collectors visited 

Tutuila during the last half of the 19th century, but 

their contributions to the flora of Tutuila are minor. 

The last collector of the century to visit and 

botanize in Samoa was F. Reinecke, who published 

the first flora of Samoa (1 896, 1898). Most of his 

collection is from Western Samoa, and many of his 

Tutuila specimens are unnumbered and lack critical 

data. His collection, stored in Berlin, was destroyed 

during World War 11, but duplicates are found at 

Kew and some other herbaria. Another botanist, K. 

Rechinger, visited the island a decade later (in 1905) 

and collected a few specimens, but the bulk of his 

collcctions, which are stored at Vienna, were made 

in Western Samoa (Rechinger 1907-191 5). 

The next significant collector in Samoa was W. 

A. Setchell, who visited Tutuila in 1920 and 

published a flora of the island (Setchell 1924). His 

collection, with about 580 numbers, is deposited at 

the University of California, Berkeley. He was 

followed by D. W. Garber, who collected about 190 

specimens (nos. 783-973) from the island in 1924. 

Other minor collections from that decade are those 

by Eames (1921) with about 30 specimens (nos. 

TI-T30), Bryan (1924) with about 68 specimens 

(nos. 956-1 0 18, 1375-1379), and Diefenderfer 

(1929 and 1930) with about 48 specimens (nos. 

1-29, 1-1 9). Their collections are principally stored 

at the Bishop Museum. 

The next major collection from Tutuila was made 

by E. Christophersen in 1929 and 193 1, and were 

included in his two publications on the flora of 

Samoa (Christophersen 1935, 1938). Christophersen 

collected about 407 specimens on Tutuila (nos. 

980-1231, 1252-1266, 1807-1830, 3478-3593). 

His publications still form the most complete 

account of the flora of Samoa. Christophersen was 

followed by T. G. Yuncker who worked on Tutuila 

in 1939. Although Yuncker published his results 

from Manu'a (Yuncker 1945), his Tutuila collection, 

numbering about 144 specimens (nos. 9300-9443) 

has not been published. The Christophersen and 

Yuncker collections are stored at the Bishop 

Museum. 

The next collections were made during the 1950s 

and 1960s, but this information has never been 

published. In 1959 A. Wisner collected on the 

island, and his unstudied specimens are stored at the 

Bishop Museum. In 1965, C. H. Lamoureux and C. 

R. Long collected on Tutuila, and their specimens 

are stored at the University of Hawai'i. Lamoureux's 

collection has about 80 specimens (nos. 3000-3029, 

305 1-3099) and Long's about 200 (nos. 1892-1997, 

2595-2609,3045-3 120). 

The original botanist on the study of American 

Samoa by Amerson et al. (1 982), J. Kuruc, collected 

a number of specimens on Tutuila in 1975, but 

except for a few, these are without any data and 

most are not even mounted. The largest collection 

of Tutuila specimens was made by the present 

author from 1972 to 1993. The collection numbers 

about 131 1 specimens (nos. 300-33 1, 1286-1289, 

1419-1486, 2689-2986, 3109-3124, 3343-3353, 

3531-3551, 3601-3673, 3745-3763, 3775-3778, 

3836-3857, 8031-8036, 8044-8074, 8393-8524, 

8526-8748, 8895-920 1, 9329-938 1). These 

specimens are in the author's personal collection, 

with many duplicates at the Bishop Museum, 

National Tropical Botanical Garden, Kew Garden, 

British Museum, and Berlin. Also, a duplicate set of 

the collections made during the present study has 

been turned over to the National Park Service in 

Honolulu. Little of this work has heretofore been 

published, other than revisions of two genera 

(Whistler 1986, 1988a). 

A checklist of the flora of American Samoa, 

island by island, was published as an appendix in a

vegetation study (Whistler 1980), based on the work 

up lo tI1:11 tinw, principally during the Amersori cl ul. 

study. 

Co~nparcd to the "continental" islands of 

Mclanc4ia lo the west, the Samoan archipelago has a 

highly iittcnuated flora--perhaps only a third as large 

ils that of Fiji, which lies just 700 miles to the west. 

However, this is larger than that of any other 

tropical Polynesian archipelago except Hawai'i, 

which has fewer genera but more species. The 

Samoan flora is estimated to consist of about 540 

native species of flowering plants (two thirds of 

them in the Dicotyledonae). These are included in 

about 282 genera in 95 plant families. There are 

also about 230 species of ferns and fern allies, most 

of which are listed in Christensen (1 943). The level 

of endemism of the angiosperms is estimated to be 

about 30% (Whistler 1992a) at the species level, but 

only one genus, Sarcopygme of the Rubiaceae 

family, is endemic to Samoa. Another 250 or so 

plants, most of them introduced species, are 

classilicd as weeds or adventive species (Whistler 

1988b). 

Based on the recent field work carried out by the 

author up to May 1993, and on a review of the 

previous literature and collections from the island, 

the native flora of Tutuila is estimated to consist of 

approximately 417 native species of vascular plants 

(see Appendix A for a complete listing of species). 

This total comprises about 301 flowering plants 

(angiosperms), 109 ferns, and 7 fern allies. This 

compares with Ta'u, which has about 329 native 

species, comprising 222 angiosperms, 104 ferns, and 

7 fern allies. An additional 179 species of vascular 

plants (all angiosperms) have been introduced and 

naturalized (most of them as "weeds") on Tutuila; 

some of these were brought in by Polynesians 

("Polynesian introductions") prior to the European 

Era, but most were introduced during recent times 

("modern introductions") after about 1800. 

The largest plant families represented in the 

Tutuila flora are Orchidaceae (49 native species), 

Rubiaceae (1 7), Cyperaceae (l5), Fabaceae (14), 

Poaceae (1 3), Euphorbiaceae (12), Urticaceae (1 I), 

and Myrtaceae (9). As noted earlier, the rate of 

cndernism in Samoa is about 30%, but the local 

island endemism for Tutuila is only about 1%; the 

only species reported to be endemic to the island are 

Cj~rlandru sp. nova (Gesneriaceae), Cyrtandra 

gcminafu, and Elalostemu tutuilensc (Urticaceae). 

The unidentified Ischaemum (Poaceae) may 

represent an additional endemic species. 

Most of the plants found on the island are not rare 

(although they may be hard to find), and of the 424 

native species of vascular plants, all but about 44 of 

them have been collected in the last 20 years. Most 

of the species present on the island are present in the 

park; those that have not been reported from the 

park are marked by asterisks in the checklist of the 

flora (Appendix A). 

Vegetation Studies on Tutuila 

The study of the vegetation of Samoa in general, 

and Tutuila specifically, has lagged far behind the 

study of the flora, and prior to 1976, very little had 

been published on it. The first known description of 

the island was made during the USEE visit in 1839. 

Pickering (1 876) made several general observations 

of the whole archipelago, but under the heading 

"Mountain- crests," he noted the following 

description for what is now called montane scrub in 

Samoa: 

"The only mountain-ridges I had the opportunity 

of visiting, were on Tutuila; the highest peak there 

being only 'twenty-three hundred and twenty-seven 

feet' above the sea. On one of my excursions, I 

approached within a fourth of a mile, and three 

hundred feet vertically, of this highest point; and 

much to my surprise, found the surface occupied in 

great part by a Pandanus, perhaps a peculiar species 

[Pandanus reineckei]; a decumbent Frecinetia also 

abounded; two plants greatly impeding progress, 

and as evening was approaching, I decided to 

proceed no farther. All the way to the summit, the 

vegetable growth continuing in sight was clearly the 

same with that around me: the matted bed of 

Pandanus and Freycinetia covering the exposed 

crest; while on the descending slopes, shrubs and 

low trees were intermingled, many of them species 

not growing in the country below: as, a scarlet- 

flowered Metrosideros [Metrosideros collina], a 

spreading tree, thirty feet high; a Myrtus? [Syzygium 

brevijolium?] with sessile, cherry-like fruit; an 

Ilex-like Acronychia [Acronychia richii]?; a 

Melastomac. [Astronidium pickeringii] with white 

and singularly wax-like petals; an Afstonia [Alsto-

nu pucijicuJ; an Eurya? [Eurya pickeringii], a shrub 

with inconspicuous flowers; a Weinmunnia 

[Wcinmunniu affinis]; a peculiar species of Genio- 

sloma [Geniostoma rupestre]; a Myrica-like Myrsine 

[Rapunea myricifolia]; the Grossostylis 

[Crosso.stylis hrJ'ora], having much of the aspect of 

I'hiladelphus; and a large flowered Cyrtandra 

[Cyrtandra geminata]: also peculiar epidendric 

Orchidaceae; the resemblance to the Tahitian 

mountain-ridges being further kept up by a few beds 

of Glcichenia [Dicranopteris linearis], the only ones 

I met with on the Samoan Islands." 

The first known ecological research in Samoa 

was done on wetlands of American Samoa (Whistler 

1976), but the only relevant wetland noted is the one 

at Vatia, which lies outside, but adjacent to, the 

park. Another study was carried out on the wetlands 

of 'l'utuila (Uiosystems Analysis lnc. 1992), which 

also nicntioned the Vatia marsh. 

The most comprehensive study to date of the 

vegetation of the island was carried out during a 

survey of the wildlife and vegetation of American 

Samoa (Amerson et al. 1982). The botanical portion 

of that study (Whistler 1980) included a description 

of the vegetation of Tutuila and the other islands of 

American Samoa. Although a number of study plots 

were established at that time, only one is within the 

park boundaries. Because the terminology in that 

study is now somewhat out of date, the vegetation 

units recognized in the present report are based on 

more recent and more comprehensive studies 

(Whistler 1992a). 

A more recent study of the vegetation was carried 

out by the U. S. Forest Service (Cole et al. 1988) in 

1986, but the work was oriented towards making 

vegetation maps from aerial photographs and 

ground truthing the results. One further ecological 

study was carried out by the author on the coastal 

vegetation, which included areas within the park, 

but this report has not been published yet by the Sea 

Grant program of the University of Hawai'i. 

11. METHODOLOGY 

A detailed checklist of the flora of Tutuila was 

prepared before the field work began; it was 

extracted from an unpublished checklist of the flora 

of Samoa prepared by the author. This checklist is 

based on the work of the U.S. Exploring Expedition 

(Pickering 1876), the flora of Reinecke (1896, 

l898), the publications of Rechinger (1 907-1 91 5), 

the flora of Setchell (1924), and the floristic work of 

Christophersen (1935, 1938). The only recent and 

up-to-date published checklist of the flora of the 

island is found in The vegetation of Eastern Samoa 

(Whistler 1980). In addition to these published 

works, specimens from the Kew Garden and Bishop 

Museums were extensively searched for specimens 

collected from the island. The collections of the 

author made on the island from 1972 to the present 

were added to this; these are in his personal 

collection, with duplicates scattered at various 

herbaria. The flora comprises Appendix A at the 

end of this report. 

The actual field work for this project was carried 

out on the island from 27 May to 16 June 1992, with 

a follow-up period from 17 December 1992 to 5 

January 1993. The field team consisted of the 

author with several other people, principally Natasha 

Bartley whose help was graciously provided by the 

Department of Marine and Wildlife Resources (for 

the May to June field work). During the field work 

the most interesting places were selected from aerial 

photographs and from 1 :200 topographic maps 

supplied by the Pago Pago office of the National 

Park Service. Further sites were visited based on 

field observations and information supplied by 

people familiar with the park area, particularly Rory 

West of Vatia village, who served as guide on 

several of the trips. 

Most of the field study was concentrated in the 

eastern half of the park, because the vegetation there 

is in much better condition than in the western half. 

This is probably due partly to the spate of recent 

hurricanes that seem to have taken a greater toll in 

the western half, but also partly to disturbance 

caused by recent agriculture adjacent and north of 

the Alava Ridge road in the western half and by the 

road itself. 

Field trips were made to most of the places 

named on Fig. 2. The coastline of the park from 

Afono to Tafeu Cove was viewed by boat during an 

aborted trip to Tafeu Cove (rough seas precluded a 

landing). 

Eleven sites (plus one on Ofu) were selected 

during the field work for vegetation sampling; at 

these sites approximately 100 trees picked at random

were measured for dbh to determine relative 

dominance of the species. It was deemed too 

time-consuming to actually set up measured 

boundaries for these plots, but the only data not 

obtained by using the quicker method is total dbh 

pcr unit of land area. On all the walks qualitative 

and quantitative notes were taken on the vegetation 

and flora. The plot data is shown in the tables in 

Appendix B. A single plot sampled in 1976 is 

within the park boundaries, and this is included in 

the appendix (as is the one from Ofu); however, the 

1976 plot was not relocated during the current field 

study, and was probably destroyed in the constru- 

ction of the Alava Ridge road. 

The parameter used to determine "dominance" is 

basal stem area (cross-section). The dominant tree 

species of a plot was the one that had the greatest 

total stem area in the sample of trees. The term 

"relative dominance," expressed as a percentage, is 

determined by dividing the total basal stem area of a 

species by the total for all the species in the sample. 

The tables also show the number of trees above 6 

inches dbh, in order to give some indication of the 

typical sizes of the trees. 

A trip was also made to Ofu from 13-16 June and 

similar field methods were employed. Because most 

of the park area on Ofu is either steep cliff or 

disturbed coastal vegetation, little undisturbed 

vcgctation was available for sampling. Conse- 

quently, only a single plot was sampled there 

(Appendix B), although another one from the 1976 

was included after modification of the data to 

correspond to the present field methods. 

During the field work voucher specimens were 

collected, and were prepared using a plant dryer set 

up at the Department of Marine and Wildlife 

Resources (DMWR) at 'Utulei. As noted earlier, a 

set of these voucher specimens was given to the 

National Park Service, and duplicates were 

distributed to various herbaria. The vegetation map 

was made based upon the recent field observations 

and upon the vegetation map of Cole et al. (1988). 

This map is very detailed (588 "polygons" are 

recognized - on Tutuila), and most of the native 

vegetation units recognized on the map are difficult 

to distinguish in the field. A further complication 

was the spate of hurricanes (Tusi, Ofa, and Val) that 

hit the island since the aerial photographs, upon 

which the maps were based, were taken (1984). 

Some of the vegetation, particularly in the western 

half of the park, was severely damaged. However, 

the maps are useful for determining the area of 

native and non-native vegetation, and were used to 

prepare the vegetation map shown in Fig. 3. 

111. VEGETATION OF THE PARK 

The vegetation of the park is divided into two 

categories, primary vegetation and secondary 

vegetation, and these are further divided into several 

subcategories and again into "plant communities" 

that share a common flora and structure. 

Primary Vegetation 

Primary vegetation includes all those plant 

communities that are in a relatively stable condi- 

tion-i.e., the structure and flora of the community 

change little over time. It may also be called native 

vegetation, since nearly all the species found here 

are native (indigenous or endemic). However, the 

secondary forests in Samoa are also comprised 

mostly of native species, unlike some other Pacific 

islands (e.g., Hawai'i), so this term is a little 

ambiguous. Being labeled as primary does not 

mean that the vegetation has never been disturbed 

(naturally or by man), only that it is composed of a 

combination of tree species that remains fairly 

constant until the next disturbance (i.e., "climax 

vegetation"). 

The primary vegetation within the park on Tutuila 

is divided here into three categories-littoral 

vegetation, rainforest, and summit vegetation. 

Another category, wetlands, comprises swamps 

(freshwater and mangrove), which are dominated by 

woody species, and marshes, which are dominated 

by herbaceous species. There are no wetlands 

within the park boundaries, but since the best 

remaining marsh on Tutuila is located within Vatia 

village, the community is included in the discussion 

here and the marsh is described. 

A. Littoral Vegetation 

The term "littoral" refers to something occurring 

on or near the seashore (litor- is Latin for seashore). 

Littoral vegetation comprises the native plant cover 

occurring on undisturbed seashores, and is the same

category recognized in the report on the Ta'u unit of 

the National Park of Samoa (Whistler 1992b). Even 

when the area is disturbed, the plants found on the 

shore are often still littoral species, because non- 

littoral species are not adapted to the harsh con- 

ditions present. 

Littoral vegetation typically has a zone of herba- 

ceous species that covers the foreshore. A zone of 

shrubby vegetation is often found inland from this, 

and farther inland a zone of littoral forest is usually 

present. These three plant zones or communities 

recognized here-herbaceous strand, littoral shrub- 

land, and littoral forest-are the same ones recogni- 

zed in the Ta'u report (Whistler 1992b). 

Littoral communities differ from other plant 

communities in their narrow extent-they are 

restricted seaward by the hightide mark of the ocean 

and inland by lowland forest. Because of this, some 

authors may regard the three types of vegetation 

recognized here as zones of a littoral community 

rather than communities themselves. This is 

supported by the tendency of some of these vege- 

tation types to be indistinct and to blend into each 

other with no sign of a clear boundary. Also, littoral 

vegetation sometimes lacks one or more of the zones 

all together. However, because of distinct 

differences in structure and flora, the three are 

rccognited here as communities. 

Littoral vegetation may occur on sandy, rocky, 

talus, or coral rubble shores, but there are no sandy 

shores within the park boundaries on Tutuila. Most 

of the park's shoreline has a solid rock substrate in 

the form of sea cliffs and windswept headlands. The 

only coral rubble beaches are found in Amalau 

Valley (Fig. 8), Vatia north of the village where the 

park boundary begins (Fig. 9), and Tuafanua west of 

Polauta Ridge (Fig. 10). There is also a very small 

coral rubble beach at Tafeu Cove (Fig. ll), but 

because of high waves, landing there was not 

possible during the field study. 

Environmental conditions found in areas of 

littoral vegetation are harsher than in any other plant 

community on Tutuila. Although the annual rainfall 

in all areas of the island probably exceeds 2500 mm 

(I00 in), the rocky surface retains little water for 

growth, and often the water that is present is 

brackish. This results in species on the exposed 

coasts (particularly the herbaceous ones) being 

xerophytic, i.e., adapted to dry conditions. The sub- 

strate itself is also a limiting factor; sometimes the 

plants present can survive only by clinging tena- 

ciously to cracks in the rocks. Littoral species are 

also adapted to bright light conditions, i.e., they are 

heliophytes (light-loving species). Because of this, 

littoral herbs and shrubs are generally absent from 

the littoral forest floor. This may also explain why 

littoral trees typically do not extend very far 

inland-they may be shaded out by the taller inland 

forest species that are, however, precluded from 

dominating the shore itself by their intolerance to 

the harsh environmental conditions. Temperature is 

also an important factor for littoral species, parti- 

cularly the herbaceous plants. While the mean 

daytime temperature is 26.7"C (80°F) on most of the 

island, the exposed rock heats up to higher 

temperatures from the relentless sun. 

The most critical environmental factor, however, 

is the sea. Salty sea air, brackish ground water, and 

occasional high waves make the habitat inhospita- 

ble to most other species. Plants present must be 

somewhat halophytic (salt tolerant) to survive. 

Occasional high waves, some with destructive force, 

may sweep the shore, ripping out poorly rooted 

plants and leaving behind puddles of sea water. 

Another characteristic shared by most littoral 

plants is their method of dispersal-most have 

seawater-dispersed seeds that can survive long 

periods of seawater immersion before washing up 

on a distant beach and germinating. Because of this 

adaptation for long-distance dispersal, very few 

littoral species are restricted in distribution; none of 

the Samoan littoral species is endemic to the 

archipelago. A few littoral species have seeds 

dispersed by adhering to seabird feathers (e.g., 

Pisonia grandis), and a few have colorful fruits that 

are eaten and transported internally by birds that 

frequent the shore. The three littoral communities 

are discussed below. 

1. Herbaceous Strand 

This community comprises the herbaceous 

vegetation occurring on the immediate seashore, and 

is the same community recognized on Ta'u (Whistler 

1992b). It is sometimes further divided into "rock 

strand" and "sand strand", but since the two habitats 

are not always distinct (there are intermediate 

types-coral rubble and talus) and the species that 

dominate the two types often cross over to the other 

habitat, this distinction is not retained here.

Figure 8. Amalau beach. 

Figure 9. Coral rubble beach north of Vatia; Pola Islet in the distance. 

12

Figure 10. Tuafanua Beach west of Vatia. 

Figure 1 1. Tafeu Cove west of Vatia. 

13

It is a moot point, anyway, since there are no sandy 

beaches within the park boundaries (although there 

are small deposits of coral sand on some of the coral 

rubble beaches). 

The herbaceous strand is limited seaward by the 

hightide mark, and inland by the edge of the littoral 

shrubland or littoral forest. In the park it is most 

frequently encountered in a narrow zone on wind- 

swept headlands (Fig. 12). The most characteristic 

species in these places are grasses such as Lepturus 

repens and lschaemum murinum, sedges such as 

Fimhrislylis cymosa and Mariscus javanicus (sele- 

sele), the vine lpomoea pes-caprae (beach morning- 

glory, fue moa), and the fern Acrostichum aureum 

(swamp fern, sa'ato). In protected areas on steep 

cliffs, ferns such as Phymatosorus scolopendria (lau 

magamaga) and Asplenium nidus (bird's-nest fern, 

laugapapa) and the vine Hoya australis (wax 

flower, fue selela) may dominate. Another herba- 

ceous species occurring here, Hedyotis biflora, 

belongs to the same genus as Hedyotis foetida, a 

species that is common in similar habitats on Ta'u 

but absent from Tutuila. 

Another type of rocky substrate vegetation that 

might be classified as herbaceous strand is the rocky 

point (Polauta Ridge and Pola Islet) north of Vatia, 

where a number of herbaceous species cling to the 

scantily clad cliffs. One of them, Boerhavia 

,qluhrc~lu, is a new record for American Samoa (it 

was collected twice in Western Samoa, but is 

irregularly distributed elsewhere in the Pacific). It 

has small sticky fruits that adhere to bird feathers, 

which accounts for its presence around this area 

whcre numerous seabirds nest. Other species may 

occur on Pola Islet, but its sheer sides and the 

surrounding waves make exploration unfeasible at 

this level of study. The vegetation there will be 

discussed in more detail under littoral shrubland. 

Coral rubble beaches are found in several places 

within the park. In bays where coral rubble typi- 

cally accumulates, the waves pile up the rubble at a 

steep angle. At the top of the slope there is a plateau 

where the flat ground surface extends into the forest. 

The best examples of this are Amalau Bay (see Fig. 

8) and the southern end of Tuafanua beach (see Fig. 

10). However, both of these are disturbed. Amalau 

is near habitations, and the presence of coconuts and 

disturbed vegetation inland of the beach attests to 

human activities. Tuafanua was apparently hard hit 

by hurricanes, as evidenced by the number of fallen 

trees found there. 

In the flat zone at the top of the slope where 

patches of sand may also be present, prostrate 

creeping vines typically comprise the dominant 

lifeform. The most common of these species are 

Ipomoea pes-caprae and Vigna marina (beach pea, 

fue sina). Less common are Canavalia rosea (fue 

fai va'a) Ipomoea macrantha, Operculina ventri- 

cosa, and Stictocardia tiliifolia. The latter two 

species are not typically littoral; they are more often 

found inland as weeds. All of these vines are native 

species, except Stictocardia and Operculina. Two 

other native species are sometimes found here, 

Zehneria mucronata and Zehneria samoensis, but 

they are more typically found a short distance inland 

under the shade of the littoral forest. 

Lifeforms other than littoral vines occur on coral 

rubble beaches. The most typical non-vine species 

are the grasses Thuarea involuta, Paspalum 

vaginatum, and Lepturus repens, and the sedge 

Fimbristylis cymosa. In more hospitable areas, 

inland weeds such as Mikania micrantha (mile-a 

-minute vine, fue saina) may also occur. In some 

places the Barringtonia trees that form the littoral 

forest may grow over the edge of the ocean creating 

a shade that precludes herbaceous strand from 

becoming established. 

Talus slopes are found in two places in the park 

-on either side of Pola Point. The eastern portion of 

this is Vatia beach, which is accessible. The other, 

Tuafanua, is reached from Vatia by a trek over a low 

saddle (70 m elevation) and a steep ascent through 

lowland forest. Tuafanua beach, about 1 km in 

length, is limited southward by a rocky headland 

and disappears northward at the base of the steep 

cliffs that form Pola Point. Talus dominates the 

northern end of this beach, below the cliffs. The 

talus itself is mostly barren, and the littoral 

vegetation begins mostly inland where there is an 

accumulation of coral rubble. This may be due to 

the inability of plants to find a suitable foothold 

between the irregular rocks and boulders, or possi- 

bly the recent hurricanes scraped clean this lower 

portion of the beach.

Figure 12. Littoral shrubland vegetation on windswept Sauma Point. 

Figure 13. Littoral shrubland vegetation on Polauta Ridge. 

15

2. Littoral Shrubland 

'I'his is the native shrubby vegetation typically 

occurring inland of the herbaceous strand (when 

prcscnt) and seaward of the littoral forest. It is same 

community recognized on Ta'u (Whistler 1992b). 

Like herbaceous strand, it may occur on either 

rocky, sandy, or coral rubble shores. However, it 

most commonly occurs on rocky coasts, and often 

blends into the herbaceous strand on this sub-strate, 

especially when the shrubs are dwarfed by 

prevailing sea winds. Littoral shrubs are typically 

intolerant of shade, and do not extend into the forest, 

where different shrubs, saplings, and ferns 

dominate. Within the park boundaries areas of 

littoral shrubland occur primarily where rocky 

points slope into the sea rather than forming 

cliff-bound headlands. 

Only a few examples of this community exist 

within the park boundaries, at least in areas that are 

accessible; the best examples on Tutuila are on the 

lhfuna coast on the north-central part of the island 

bctwcen Fogagogo and Vaitogi, where the lava- 

bound coast forms a suitable habitat for the shrubs. 

The best examples in American Samoa are on the 

southern tip of Olosega, and on the south corners of 

Ta'u. 

The most typical littoral shrubland species in 

Samoa are Scaevola taccada (to'ito'i) and 

Wollastonia biflora (beach sunflower, ateate). 

Scaevola is a shrub that is widespread in the Pacific. 

Wollastonia, a species with a similarly wide distri- 

bution, is more of a subshrub in most areas, app- 

earing even vinelike when it hangs down from cliffs 

of headlands. Other typical species in this habitat 

arc Clcrodendrum inerme (aloalo tai), Colubrina 

u.viutica (fisoa), Dendrolobium umbellatum (lala), 

and C'yrtandra samoensis. 

The best accessible area of littoral shrubland 

within the park is found on the precipitous slopes of 

Polauta Ridge at the north end of Vatia beach (Fig. 

13). An equally good example is Pola Islet (Fig. 

14), but this in inaccessible. The vegetation on the 

islet and on the cliffs of the ridge is actually a 

mixture of littoral herbs, shrubs, and trees that find a 

tenacious existence on the cliffs and in rock cracks. 

Because the littoral plants present are not trees, the 

vegetation cannot be classed as littoral forest. But 

since most of the species on Pola Islet are woody, it 

is probably most accurate to consider the whole 

plant cover as littoral shrubland. 

The dominant species present here is Scaevola 

taccada. Other common woody species present are 

Clerodendrum inerme, Premna serratifolia, Hibi- 

scus tiliaceus, Morinda citrifolia, and Pisonia 

grandis. The most abundant herbs are the grasses 

Lepturus repens and Stenotaphrum micranthum, and 

the fern Acrostichum aureum. Higher up the slopes 

the fern Davallia solida forms dense patches, 

probably in cracks and on ledges. A different kind 

of littoral shrubland is present at the north end of 

Vatia on the flat coral rubble bench above the 

sloping beach. This is dominated by Hibiscus 

tiliaceus, but also common here are Clerodendrum 

inerme, Barringtonia asiatica, Morinda citrifolia, 

and Ficus scabra. This may actually be the remains, 

and beginnings, of a littoral forest that was cut down 

or destroyed by hurricanes. A similar vegetation 

covers the talus slopes behind this area, and may be 

a disclimax vegetation kept from reverting to littoral 

forest by the talus that slides or falls down from the 

adjacent cliffs. 

Another variation of littoral shrubland sometimes 

mentioned in the literature is one dominated by 

Pandanus tectorius (screwpine, fasa). While there 

are areas on Tutuila where this tree-like monocot 

forms monodominant stands, such as on the Tafuna 

coast between Fogagogo and Vaitogi, the plant is 

uncommon within the park boundaries. It fact, 

Pandanus was not even noted during the survey as 

occurring on any of the beaches visited (it was, 

however, seen at higher elevations on Mauga-o- 

Tula). Consequently, the possible distinctiveness of 

Pandanus scrub vegetation is a moot point within 

the park boundaries. 

3. Littoral Forest 

This is the forest that occurs directly on the shore 

(or just inland of the other zones of littoral vegetation) 

and is dominated by trees that owe their 

distribution either directly or indirectly to the effects 

of the sea. It is the same community recognized on 

Ta'u (Whistler 1992b). It ofien occupies a narrow 

zone sometimes only 50 or 100 m in width, 

especially on level coasts where it is replaced inland 

by lowland forest. This was the case in Amalau 

before it was disturbed; here the disturbed littoral 

forest zone is replaced inland by secondary forest

Figure 14. Littoral shrubland vegetation on Pola Islet. 

Figure 15. Aiga Bay, west of and outside of the Tutuila park. 

17

itnd li~t-hx inlilr~d by lowland forcsl donlinalccl by within the park (il was only seen to be common at 

I!y.vo\ylunl tnuolu. Rclbre Amalau was disturbed, it the north end of Vatia village). On uninhabited 

was probably very similar to a site studied at Aiga Rose Atoll, however, which is the site for an 

Bay on the western end of the island (Whistler 1980) extensive seabird colony, the only patch of forest 

and shown in Fig. 15. On steep coasts, the zone of 

littoral forest may extend much farther inland. In a 

plot (Appendix B: plot 2) on Sauma Ridge at 185 m 

elevation and nearly a half mile away from the coast 

(down the ridge), a littoral tree, Barringtonia 

usiutica (fish-poison tree, futu) was the second 

dominant species with 17% relative dominance. 

present is entirely dominated by huge - Pisonia trees. 

Hibiscus is an adaptable species that may 

dominate in freshwater swamps, on the margins of 

mangrove forests, along estuaries, on beaches, on 

talus slopes, and in disturbed lowland forest up to 

the higher elevations of the island. It is a secondary 

Burringtonia is, in fact, the dominant littoral 

forest tree on all the coasts within the park, as it is 

on nearly all of the coasts of the whole island. This 

huge tree has thick, spreading branches that may 

even overhang the shore in protected areas. On the 

thrce areas of coral rubble beach within the park, 

Bwringhnia probably has a relative dominance of 

over 80%. A similar figure may apply to the littoral 

forest on steep slopes. 

The second most abundant species is either 

Terminalia catappa (tropical almond, talie) or 

Calophyllum inophyllum (Alexandrian laurel, 

fetau), which are similar in size and shape to 

Burringlonia, although perhaps somewhat taller. 

Also present in the littoral forests within the park are 

Hernandiu nymphaeifolia (Chinese-lantern tree, 

pula), Pisonia grandis (pisonia, pu'avai), Hibiscus 

~iliuceus (beach hibiscus, fau), Thespesia populnea 

(Pacific rosewood, milo), Cerbera manghas (leva), 

Cocos nucifera (coconut, niu), and Guettarda 

speciosu (puapua). 

On sandy beaches Hernandia, which is a huge 

trce like the three most abundant species noted 

above, apparently replaces or competes with 

Ilurringtoniu in dominance. Since there are no 

sandy beaches within the park boundaries, this is a 

moot point. However, Hernandia shares dominance 

with Barringtonia and Pisonia in the small area of 

littoral forest north of the village of Vatia (Fig. 16). 

species on beaches (as it is in most of the places 

where it is found), and its presence usually indicates 

some kind of disturbance. It is a smaller tree than 

the littoral trees discussed above, and often forms 

dense thickets that are penetrated only with 

difficulty. The coconut is another species whose 

presence usually indicates disturbance. Although it 

may naturally dominate on sandy beaches of atolls, 

on the coasts of Samoa it is probably mostly or 

entirely a secondary species planted for its nuts. 

The other three trees, Cerbera, Guettarda, and 

Thespesia, are smaller and less common than the 

foregoing species, and have not been reported in 

Samoa to form a significant part of any littoral 

forest. A few other species, such as Planchonella 

grayana, Diospyros samoensis ('au'auli), and 

Diospyros elliptica ('anume), are common in some 

littoral forests. However, these trees are coastal 

rather than littoral, i.e., they occur in coastal regions, 

but not usually on the immediate shore. They are 

more frequent on coastal ridges, and will be 

discussed in more detail under lowland forest. 

B. Wetlands 

As noted above, this category comprises swamps 

and marshes. Although no swamps or marshes are 

found within the Tutuila park boundaries, the marsh 

adjacent to Vatia village is discussed here because it 

is so close to the park, because it is relatively 

undisturbed, and because it is a vegetation type 

under considerable pressure in Samoa. The report 

on the Ta'u unit of the National Park (1992~) 

Pisonia is typical of undisturbed beaches where 

seabird colonies are found (its sticky fruits are discussed a wetland from that island under 

transported attached to seabird feathers). The "disturbed coastal marsh." The largest marsh on that 

probable demise of seabird colonies in the past 

because of overharvesting may account for the 

limited extent of Pisonja on Tutuila as well as 

island is located at Ta'u village; it is similar to the 

one at Vatia, but was not discussed in the 1992 

report because it is not within the park boundaries.

Figure 16. Littoral forest north of Vatia. 

Figure 17. Vatia coastal marsh. 

19

4. Coastal Marsh 

This is the herbaceous vegetation growing in 

water-logged freshwater depressions. It is typically 

found in coastal areas, usually in bays that have 

been filled in and subsequently protected from the 

Tea by a barrier of sand. In these areas water from 

incoming streams accumulates and forms wetland 

soil. If the barrier of sand is complete, a freshwater 

marsh forms; if the barrier is breached by an exiting 

stream, a mangrove forest usually forms. Not all 

freshwater marshes are coastal nor are all formed in 

bays; in Western Samoa some are found in montane 

craters, and one on 'Aunu'u Island is located in a 

coastal volcanic crater. 

The marsh at Vatia is located behind (west of) 

Vatia village (Fig. 17). It was apparently once used 

for growing taro, but there is no indication that this 

was recent. The dominant plant in the center of the 

marsh is Eleocharis dulcis (water chestnut, 

'utu'utu), an indigenous sedge that typically 

dominates Samoan marshes. Not present in the 

marsh, nor apparently on the island, is the marsh 

fern Cyclosorus interruptus, which is often abundant 

in marshes on other islands in Samoa. Also 

common in the Vatia marsh are Vigna marina (the 

littoral beach pea vine), and Rhynchospora 

corymhosa (selesele). In a couple of places 

Acrosrichum aureum (swamp fern, sa'ato) forms 

clumps. Other species present in smaller quantities 

are Commclina diffusa (mau'utoga), Mikania 

micruntha (mile-a-minute vine, fue saina), 

Ludwigia octovalvis, Ludwigia hyssopifolia, and 

Allernanthera sessilis-all typical introduced wetland 

species. Along most of the edges there are thickets 

of' Hihiscus tiliaceus (beach hibiscus, fau). 

Also present in the marsh is the small fragrant 

herb Limnophilafragrans. It is rare in Samoa, and 

over most of its former Pacific range, and should be 

a candidate for endangered species status in 

American Samoa. The only other place it is noted 

from on Tutuila is Faga'alu Stream, and that record 

is 73 years old. It is, however, found in the marsh at 

Ta'u, the one at the Ofu airport, and the one on 

Olosega. The Vatia marsh is the best preserved one 

on Tutuila, and possibly the second best in the 

territory, since the Ofu and 'Aunu'u marshes were 

devastated by recent hurricanes. 

C. Rainforest 

The tropical rainforest is the high forest 

composed mostly of native trees adapted to moist 

forest conditions. Before the arrival of the first 

Polynesians in Samoa, nearly all of the land surface 

of the archipelago was clothed in a mantle of 

tropical rainforest. The only areas not covered with 

it were the coastal zones, where the harsh conditions 

prevent rainforest from becoming established; 

wetlands (marshes, mangrove forest, freshwater 

forest), where water-logged soil inhibits rainforest 

development, small summit areas where soil factors 

allow only scrubby vegetation to grow; and cliff 

faces. These types of vegetation, including 

rainforest, are all classified as climax vegetation 

-they are relatively stable in structure and compo- 

sition over long periods of time. 

To the list of non-rainforest types of vegetation 

can be added two types of successional vegetation 

-that found on lava flows, and secondary vegetation 

communities (secondary scrub, secondary forest). 

These have suffered some kind of severe 

disturbance, but if left alone, they will eventually 

revert to rainforest. Lava flows are not currently 

found on Tutuila (although the Tafuna Plain was 

formed by a geologically recent lava flow) and will 

be discussed no further. The disturbance causing 

most secondary vegetation is the activity of man. 

The early Samoans cleared large areas of the island 

for growing crops and establishing villages. In 

addition to the areas now used for villages and 

plantations, some inland and isolated places in the 

park show signs of old villages and plantations 

-archaeological sites and occasional cultivated trees 

persisting far away from any currently used land. 

Thus all of the primary vegetation within the park 

boundaries can be classified as rainforest, except for 

the coastal areas (herbaceous strand, littoral 

shrubland, and littoral forest) and the one area of 

montane scrub behind the village of Vatia. 

Variation within the Rainforest 

The rainforest within the park is not uniform: 

several different types can be recognized on the 

basis of floristic composition. Four major factors 

cause these floristic differences-soil, topography, 

elevation, and disturbance.

Soils differ in origin and age of the parent 

bedrock, and younger soils tend to be more fertile 

than older clayey soils. However, most of the park 

area Ins the same origin-the old Pago Pago volcano, 

the caldera of which forms Pago Pago harbor. The 

major differences in soil are between the hillside 

soils and the alluvial soils of some of the valleys, 

which leads to differences in the forests of the 

alluvial valleys and the forests of the ridges. The 

vast majority of the park area is classified as "Fagasa 

family-Lithic Hapludolls-Rock outcrop association" 

(Nakamura 1984), which comprises the ridges and 

upper valleys. The only other significant type 

present is "Aua very stony silty clay loam," which is 

located on the lower slopes and lowlands around 

Vatia, at Va'aogeoge Cove west of Vatia, and at 

Amalau Bay east of Vatia. 

Topography is related to soil, since the alluvial 

valleys are flat to gently sloping and the hillsides are 

steep. Steep slopes increase the rate of run-off, and 

presumably create drier soil conditions. Conversely, 

the soil of the alluvial valleys is flat, with slower 

run-off and more water available from the streams 

and rivulets from the slopes, and presumably the 

trees found there thrive because of the moister soil 

conditions. 

Elevation is important for two reasons-tempera- 

ture and rainfall. Rainfall generally increases with 

elevation. In Western Samoa, Wright (1963) 

estimated it increased at a rate of 22-30% with 

every 100 m increase in elevation, at least in the 

areas he identified as "foothills." Temperature 

decreases with elevation, at a rate of about 0.7"C 

(1.3"F) per 100 m. Elevation probably makes a 

significant contribution to the variation of rainforest 

within the park, since it varies from near sea level to 

480 m elevation-a potential difference of 106 to 

144% in rainfall, if Wright's figures can be applied 

to Tutuila. Temperature is probably not that 

significant a factor, since the difference between the 

coastal areas and the summit of the park would be 

less than 3.S°C (6.3"F). 

Disturbance is a factor that is difficult to assess. 

Vegetation is not static; it is always in a state of flux. 

But when a forest becomes relatively stable in flora 

and structure for long periods of time, it is termed 

climax forest. But since there are continually 

occurring disturbances-both man-made and 

natural-that affect small and large areas of the 

forest, the result is a mosaic of vegetation at 

different ages of development. If the disturbance is 

complete, i.e., the canopy trees are removed and the 

disturbance is of short duration (a hurricane, or 

several crops grown there before the land is 

abandoned), the area will relatively quickly undergo 

a series of predictable structural and floristic 

changes, a process known as succession. 

The longer the period between disturbance, the 

more the mosaic parts tend to become uniform in 

flora and structure. The larger the area of 

disturbance, the larger the pieces in the mosaic. 

There are certain species whose presence in forest is 

an indication of past disturbance, particularly Rhus 

taitensis, Alphitonia zizyphoides, Macaranga 

stipulosa, and Hibiscus tiliaceus, which are 

discussed in more detail in the section on secondary 

forest. 

The classification of the rainforest of the park into 

smaller units (communities or associations) is a 

difficult and often unrewarding task. While 

categories can be constructed by looking at part of 

the park, the classification becomes less and less 

maintainable when more areas are sampled. That is 

because such classifications are artificial-they are 

constructed mainly for the convenience of those 

studying the subject and may not represent any 

meaningful reality in nature. Plant species each 

distribute themselves in their own unique ways, 

based on environmental factors, and to link these 

species with variable distributions into concrete 

vegetation units can only lead to inconsistencies. 

Another problem encountered when trying to put 

vegetation into categories is boundaries. If the 

community to be defined is monodominant (i.e., it is 

entirely dominated by a single species), a clear 

boundary may be present. Mangroves, for example, 

have a relatively clear boundary marked by the 

upper limits of the influx of salt water. However, 

communities with several dominant species are 

more difficult to define with boundaries. This is 

particularly a problem on ridges that extend from the 

sea to high elevation with no abrupt environmental 

boundaries (such as change in soil type). This 

uniformity of environment and absence of abrupt 

environmental boundaries is characteristic of the 

park.

Ilowever, there are differences within the 

rainforest of the park, and it is useful to recognize 

some vegetation units. The two units recognized 

here are communities and associations. Plant 

communities differ from each other in flora and 

structure; associations comprise subdivisions of 

con~nluuitic\ llrat dif'fer little in structure, but are 

dominated by different species. The communities 

and associations recognized within the park are 

determined mostly by elevation. 

Based on differences in species composition and 

elevation, the rainforest in the park is divided here 

into two communities-lowland forest and montane 

forest. A somewhat arbitrary boundary dividing the 

two is an elevation of about 350 m (1 150 ft), where 

there seems to be a decrease in tree species typical 

of the lowlands and an increase in species typical of 

the highlands. This same elevational boundary was 

used to distinguish lowland and montane forest on 

Ta'u (Whistler 1992b). The lowland forest is further 

divided into four types based on characteristic 

species, which are discussed below. 

Characteristics of Rainforest 

Rainforests share a number of similarities in 

structure and form. When undisturbed, they often 

have a fairly open floor dominated mostly by 

shade-tolerant herbs that are adapted to shady 

conditions. The herbs are mostly ferns, a few are 

terrestrial orchids, and very few are other flowering 

plants. A sizable number of the 109 fern species 

native to Tutuila (see Appendix A) are terrestrial 

species of primary forest. Of the 49 native species 

of orchids in Samoa, only about 12 are typical of the 

shady floor of primary forest (the others are mostly 

epiphytic). Grasses and sedges are relatively rare in 

undisturbed rainforest, since most of them are 

sun-loving species; they are most commonly found 

along trails in the forest, where more sunlight is 

available. 

Shrubby species are relatively sparse on the floor 

of undisturbed rainforest, with the exception of a 

few shade-tolerant species. The shrub layer is 

instead often dominated by saplings of the canopy 

species, which, if given a chance, will eventually 

grow into the canopy. Canopy trees typically have 

large, relatively poorly dispersed seeds or fruits, 

unlike secondary forest species that have smaller, 

more readily dispersed seeds or fruits. 

Several species characteristic of the floor of the 

rainforest do not readily separate into herbs or 

shrubs. The ti plant Cordylinefiuticosa is shrublike, 

but is a non-woody monocot. Another monocot is 

the giant banana-like herb Heliconia laufao (laufao) 

that sometimes forms dense patches. Tree ferns are 

also in this difficult category. The tree fern Cyathea 

vaupelii (olioli) is common in the rainforest, but it is 

usually only a couple of meters in height. Several 

other large ground ferns are found in the rainforest, 

such as species of Angiopteris and Marattia, which 

have long thick stemmed fronds that radiate out 

from a large hemispherical base. 

Disturbance that leads to the opening of the 

canopy leads to changes on the forest floor, as 

sun-loving species (many of them "weeds") rapidly 

invade. The presence of these sunlight-requiring 

invaders may be short-lived if the canopy is quickly 

restored. Some of the invaders, however, are the 

vines that ascend into the forest canopy and 

establish themselves for the long haul. 

Other significant lifeforms are present in the 

rainforest besides the trees, shrubs, and herbs noted 

above. One of the most significant comprises 

climbers. Two strategies are employed by rainforest 

climbers-trunk-climbing and twining. Trunk 

climbers are species that produce adventitious roots 

along their stems that allow them to cling to the tree 

trunk. The most significant of these in the rainforest 

of the park are Freycinetia reineckei, Freycinetia 

storckii (both 'ie'ie), Piper graeffei (fue manogi), 

and Lomagramma cordipinna (a fern). These may 

begin as creepers on the forest floor, but when they 

encounter a suitable tree, they ascend the trunk to 

where light conditions are much more favorable for 

their growth. 

The other type of climber, if it is woody, is called 

a liana. Lianas typically climb by means of a 

twining growth motion of the stem. When they 

encounter a tree trunk, they twine around it and 

eventually up into the canopy. They may eventually 

loose their connection with the tree trunk (as when 

the tree dies and disintegrates or falls), resulting in 

cable-like vines hanging from the canopy. These 

two categories are not absolute, since some plants 

seem to be intermediate between the two. Another 

method of climbing is by means of tendrils, but this 

is not common in the rainforests of Tutuila; the best

example on Tutuila is the climbing reed-like 

Flagellaria gigantea (lafo). Some other vines 

appear to be independent of the ground and grow as 

epiphytes. 

The other major lifeform in the forest is that of 

epiphytes. An epiphyte is a plant that grows on trees, 

using them as a place to grow while deriving no 

nutrition from them (as a parasite would). The most 

common epiphytes in the rainforests of Tutuila are 

ferns and orchids. Most of the 49 orchids of Samoa 

fit into this category, particularly members of the 

genus Dendrobium and Bulbophyllum. Epiphytes 

are light-loving species that cannot survive in the 

shady conditions on the forest floor, and 

consequently live perched upon the branches and 

upper trunks of the trees where the light conditions 

are better. They have to have some form of 

attachment to the tree, and they must be somewhat 

xerophytic to survive away from the moisture-laden 

soil. The two rainforest communities, lowland 

forest and montane forest, are discussed below. 

5. Lowland Forest 

This is the high tropical rain forest that covered 

most of the area of the park prior to the arrival of the 

first Polynesian settlers in Samoa. It currently 

covers the ridges and valleys on the north coast up 

nearly to the highest elevations where it is replaced 

by montane forest. Some of these ridges, 

particularly the upper portions between Fagasa and 

the TV transmitter, have recently been disturbed by 

taro plantations, and much of the same area was 

extensively damaged by the recent hurricanes. 

However, most of the lowland areas east of Toa 

Ridge (just to the east of the TV transmitter) and 

west of Afono are covered with native lowland 

forest that is relatively undamaged by the recent 

spate of hurricanes. Three types of lowland forest 

are distinguished below: valley lowland forest, 

coastal lowland forest, and ridge lowland forest. 

a. Valley Lowland Forest - This is the forest that 

covers alluvial valleys near the coast. It is the same 

as the Dysoxylum lowland forest on coral rubble 

described on Ta'u (Whistler 1992b). The alluvium 

is deposited by streams, and the larger and broader 

the valley, the greater the area of this forest. The 

species that dominate here are probably adapted to 

moist loose soil conditions, and are generally absent 

or insignificant on the ridges, which presumably 

have drier, denser soil. This is one of the habitats 

most suitable for villages, and it is likely that 

Fagasa, Vatia, and Afono were covered with this 

type of lowland forest prior to settlement. The only 

area of this now within the park is at Amalau, but it 

is disturbed. 

A plot in valley lowland forest was sampled on 

the west side of the stream in Amalau Valley inland 

from the road (see Appendix B: plot 10). This area 

is dominated by Dysoxylum maota (maota), which 

had an estimated relative dominance of 46%. The 

tree species second in dominance is Planchonella 

samoensis (mamalava). Also common are 

Inocarpus fagifer (Tahitian chestnut, ifi) and 

Myristica fatua (Samoan nutmeg, 'atone). Other 

significant trees present are Canarium vitiense 

(ma'ali), Kleinhovia hospita (fu'afu'a), Bischofia 

javanica ('o'a), Cananga odorata (ilangilang, 

moso'oi), Elaeocarpus ulianus, Macaranga 

st@ulosa (lau fatu), and Ficus scabra (mati). 

This vegetation is very similar to one sampled in 

a similar bay called Aiga, just to the west of A'asu 

on the northwest coast of the island (Whistler 1980). 

In that plot Dysoxylum spp. (listed mostly as D. 

samoense, perhaps incorrectly) had a relative 

dominance of 74%. The most numerous trees there 

were Myristica fatua and Barringtonia samoensis 

(falaga). The latter species, which is related to the 

littoral Barringtonia asiatica, is typical along 

streams. It was also common at Amalau outside of 

the plot area along the stream. While there is no 

distinct riparian forest in Samoa, two 

species-Barringonia samoensis and Inocarpus 

fagifer are typical along streams (Fig. 18). 

There is another area of this type of forest below 

the road in Amalau Valley. It is somewhat 

disturbed, but is also dominated by Dysoxylum 

maota, with a subcanopy primarily dominated by 

Myristica fatua. Another similar forest was 

described in the Ta'u report, based on a sample made 

there in 1976 (Whistler 1980). In that sample, the 

dominant tree was determined to be Dysoxylum 

samoense with a relative dominance of 86% (D. 

maota is not found on the island), an unusually high 

degree of monodominance for lowland forest.

Figure 18. Streamside vegetation at Amalau Valley.

When undisturbed, the floor of the forest is 

relatively open, and is usually dominated by shade- 

tolcrant ferns. At Amalau, the dominant ground 

cover species was the fern Lomagramma cordi- 

pinna, with lesser amounts of Arthropteris repens 

(mostly on rocks and tree trunks) and Tectaria 

sfmrrisii, also ferns. Other terrestrial ferns present 

include Asplenium nidus (bird's-nest fern, 

laugapapa), Asplenium polyodon, Pteris tripartita, 

Angiopteris evecta (king fern, nase), Bolbitis palu- 

sfris, Phymatosorus scolopendria (lau magamaga), 

Pneumatopteris bryanii, and Diplazium proliferum. 

This is very similar to the previously mentioned 

Aiga plot sampled in 1976, where Lomagramma 

was the dominant, followed by Asplenium nidus 

(Whistler 1980). 

Trunk climbers are fairly common in this 

community, especially the aroid Epipremnum 

pinnatum and the previously mentioned ferns 

Lomagramma cordipinna and Arthropteris 

repens, which are usually restricted to the lower 

trunks. Also common in some places is Piper 

graeffei. Similar results were recorded at Aiga, 

where Epipremnum and Piper were the two 

dominant species. Lianas are not very common in 

this community, at least at Amalau. The most 

typical species are Gynochtodes epiphytica, Hoya 

pottsii, and Alyxia bracteolosa (lau maile). 

Epiphytes are not very abundant, at least when 

compared to montane forest. The most common 

species is Asplenium nidus, as it was at Aiga; also 

present are Phymafosorus scolopendria, Davallia 

.solids (laugasese), Davallia epiphylla (laugasese), 

Nephrolepis biserrata, and Antrophyum planta- 

gineum, all of which are ferns, and Dendrobium 

dacfylodes, which is an orchid. 

b. Coastal Lowland Forest - This is the lowland 

forest that occurs on ridges from inland of the 

littoral forest to about 200 m elevation (Fig. 19). It 

is virtually the same thing as the "'au'auli coastal 

forest" recognized in American Samoa (Whistler 

1980) and "Diospyros coastal forest" described from 

the offshore islet of 'Upolu, Western Samoa 

(Whistler 1983), but was not reported in the park 

area on Ta'u (Whistler l992b). It is characterized by 

species that are found on lower ridges and by the 

presence of a some of littoral species. It is some- 

what transitional between the forest of interior 

ridges and littoral forest dominated by Barringtonia 

asiatica (futu). The boundaries between this forest 

and the other two are not distinct, especially since 

Barringtonia can be found at over 200 m elevation 

(and occasional trees at much higher elevations, but 

possibly planted there). 

Two plots, nos. 1 and 2 in Appendix B, were 

sampled in this forest. Plot 2 was at Matapai Point 

(shown on most maps as "Matape") at the northeast 

comer of the park (and perhaps just a little outside 

of the currently proposed boundaries) at about 50 m 

elevation, the other on Sauma Ridge just to the west 

of Amalau Valley at 185 m. These two are some- 

what heterogeneous, because they probably 

represent the lower and upper limits of this type of 

lowland forest. The lower plot had some littoral 

forest elements, such as Terminalia catappa (talie), 

and the upper had some elements from the higher 

elevation ridge forest. 

By combining the data from the two plots, the 

following trees were shown to be the typical species 

(excluding secondary forest species): Diospyros 

samoensis ('au'auli) with an average relative 

dominance of 18%, Garugafloribunda (vivao) with 

17%, Barringtonia asiatica with 12%, Planchonella 

garberi ('ala'a) with lo%, Planchonella grayana 

with 5%, Guettarda speciosa (puapua) with 3%, 

and Syzygium clusiifolium (asi vai) with 1%. Three 

other species characteristic of this vegetation are 

Diospyros elliptica ('anume), Sterculia fanaiho 

(fana'io), and Syzygium dealatum (asi vai?). A 

dominance of the Syzygium and the Diospyros 

species is probably the best indicator of this type of 

vegetation. 

The forest floor is relatively open, since it is 

much drier than the above described lowland forest 

of alluvial valleys. The dominant species is 

probably Asplenium nidus (bird's- nest fern, 

laugapa). Also sometimes common are the ferns 

Phymatosorus scolopendria (lau maga maga), 

Davallia solida (laugasese), and Tectaria stearnsii. 

In sunny places where the canopy is broken, weeds 

such as Nephrolepis hirsutula (sword fern, vao 

tuaniu), may become common. Seedlings are often 

abundant, and sometimes may even dominate the 

forest floor. The most common of these are 

probably Diospyros elliptica and Garuga j70ri- 

bunda, but composition depends upon which trees 

dominate in the area. Littoral shrubs such as 

Colubrina asiatica (fisoa) and Clerodendrum

Figure 19. Coastal lowland forest near Tafeu Cove. 

Figure 20. Disturbed lowland forest on ridges north of Mt. Alava.

incrme (aloalo tai) may also be common, 

particularly if the site is very near the ocean. 

Another shrub, Alyxia stellata (nau), is also 

sometimes common. 

Vines are not common in this type of lowland 

forest, probably because of the dry soil and salty 

winds from the nearby sea. The most frequent 

species is probably the trunk climber Epipremnum 

pinnatum. Other species present include Hoya 

australis (fue selela), Mucuna gigantea, and 

Ipomoea macrantha. Epiphytes are also relatively 

sparse for the same reasons. The most frequent 

species are Asplenium nidus, Davallia solida, and 

Davallia epiphylla, all of which are ferns, and the 

orchid Dendrobium dactylodes. 

c. Ridge Lowland Forest - This is the forest on 

the upper portion of the ridges and slopes, ranging 

from about 200 to 350 m elevation, but sometimes 

lower. This forest has indistinct boundaries because 

it blends into coastal lowland forest below it and 

montane forest community above it. It corresponds 

to the "mixed lowland forest" described for Ta'u 

(Whistler 1992b), the "asi ridge forest" of American 

Samoa (Whistler 1980), the "mafoa ridge forest" of 

'Upolu, Western Samoa (Ollier et al. 1979), and the 

"asi forest" and "tamanu forest" of Savai'i and 

'Upolu (Chandler et a[. 1978). It probably covers 

the majority of the park, but the western half of the 

area has recently been disturbed by hurricanes and 

human activities (Fig. 20). 

Ridge lowland forest is not a monodominant 

community, and any one of several trees may 

prevail at a particular site. The seven most 

characteristic canopy trees in the forest on Tutuila 

are shown in Table 1. The four canopy species 

averaged between 12 and 16% relative dominance. 

The first species, Syzygium inophylloides (asi), is 

also the dominant of the lowland forest of Ta'u 

(Whistler 1992b). Calophyllum neo-ebudicum 

(tamanu) is common throughout the archipelago in 

lowland forest of ridges, as is Canarium vitiense 

(ma'ali). Buchanania merrillii (no Samoan name) 

finds its greatest abundance on Tutuila; it is absent 

from Manu'a, and is uncommon in Western Samoa. 

The subcanopy trees may sometimes be 

dominant, but they are not as tall nor as big (in girth 

or dbh) as the other species. Diospyros samoensis is 

a dominant tree of lowland forest of coastal ridges, 

and becomes progressively less important with 

increasing elevation. Myristica fatua (Samoan 

nutmeg, 'atone) is typical of most types of lowland 

forest, but at higher elevation on ridges it is often 

replaced by Myristica hypargyraea (also called 

'atone). Canarium harveyi (mafoa) may be a 

modern introduction to Samoa, since it was not 

recorded in the islands before 1920; consequently, it 

may actually be a canopy tree that has not been in 

Samoa long enough to grow into the huge size 

characteristic of older native trees. 

Table 1: Relative dominance (%) of characteristic trees of the lowIand forest of interior 

ridges of the Tutuila park. 

CANOPY SPECIES 

PLOTS AVERAGE 

5 6 7 8 9 

Syzygium inophy No ides 16 11 22 25 4 16 

Calophyllum neo-ebudicum 11 13 17 3 12 13 

Canarium vitiense 8 7 12 11 2 5 13 

Buchanaria merrillii 1 3 1 11 - 18 12 

SlJBCANOPY SPECIES 

Diospyros samoensis 

Canarium harveyi 

Myristica fatua

The above seven characteristic trees accounted for 

about 75% of the relative dominance of the five plots; 

most of the remainder comprised occasional 

secondary forest trees (Hibiscus tiliaceus, Rhus 

taitensis, and Alphitonia zizyphoides), montane forest 

species at the higher elevations, and coastal lowland 

forest species at the lower elevations. The presence of 

the secondary forest species indicates some kind of 

past disturbance. 

It may have just been a small gap (see "gap 

replacement" in the section on secondary forest), but 

the presence of Hibiscus may indicate some human 

activity there. 

A variation of this kind of lowland forest occurs 

when Inlsiu bijuga (ifilele) is present. This tree has an 

odd distribution on Tutuila: it is found at Nu'uuli just 

inland from the mangrove swamp, and on a number of 

ridges on the north shore of the island between Vatia 

and Fagamalo; there is also a population on 

Nu'usetoga islet at Masefau. It is common all along 

the ridge behind Vatia, including the top of 

Mata-o-Tula in the montane scrub, and on the lower 

part of Faiga Ridge, which is contiguous with it. It is 

odd that this tree is not reported on the ridges of the 

south part of the island, unless it was removed in the 

past by relentless logging (it is probably the finest 

timber tree in Samoa). It may be an ancient 

introduction to Samoa that only grows in areas where 

it was originally planted, but this is only speculative. 

In plot no. 4 on Muli'ulu Ridge east of Fagasa, 

lntsia was the dominant species with a relative 

dominance of 20%. The other important species in 

that plot, in descending order of relative dominance, 

were Canurium hurveyi, Syzygium inophylloides, 

Cunurium vitiense, Buchanania merrillii, and Calo- 

phyllum nco-ehudicum, which are the same species 

typical of the lowland forest of the interior ridges. In 

the 1976 survey of the vegetation of American Samoa, 

the only plot where Intsia turned up was on 

Nu'usetoga Islet, where it was the dominant species 

with a relative dominance of 20%; the second 

dominant species there was Syzygium inophylloides. 

On a similar ridge on 'Upolu, Intsia was the dominant 

with 49% relative dominance (Park et al. 1992); the 

next four dominant species there, in descending order, 

were Syzygium inophylloides, Canarium vitiense, 

Diospyros samoensis, and Myristica fatua. 

The floor of the forest often has a relatively dense 

ground cover (Fig. 21), partly perhaps because the 

steep sides of the ridges may allow more light 

through the canopy, but more importantly, because 

of the recent hurricanes. In shady areas on 

relatively flat places on the ridge tops, Loma- 

gramma cordipinna may dominate, but in most 

areas in this forest it is relatively uncommon. The 

most common species are the ferns Asplenium 

nidus (laugapapa), Tectaria stearnsii, Christella 

harveyi, Microsorium sylvaticum, Phymatosorus 

scolopendria (lau magamaga), and Nephrolepis 

hirsutula (sword fern, vao tuaniu). In places with 

a disturbed canopy, vines such as Gynochtodes 

epiphytica, Faradqya amicorum (mamalupe), and 

Alyxia bracteolosa (lau maile) may form dense 

thickets. 

There are also several small trees that are 

particularly common in this forest. The most 

frequent of these are Ixora samoensis (filofiloa), 

Psychotria insularum (matalafi), and Aglaia 

samoensis (laga'ali). Less common are Casearia 

sp. nova, Citronella samoensis, and Flacourtia 

rukam (filimoto). These are all native species; the 

introduced shrub Clidemia hirta (Koster's curse) is 

found here in small amounts, but is much more 

common at higher, wetter elevations. Also 

common are Cordyline fruticosa (ti), Alyxia 

stellata (nau), and Zingiber zerumbet ('avapui). 

The last species is particularly common along 

ridge tops, and may be a naturalized relict of 

former cultivation once occurring in these places. 

It apparently spreads only infrequently to other 

areas. 

Epiphytes are more common here than in 

coastal lowland forest, but less common than at 

higher elevations in montane forest. The most 

common epiphytic orchids are Dendrobium 

biflorum, Dendrobium dactylodes, Flickingeria 

comata, Bulbophyllum polypodioides, and 

Phreatia micrantha. The most common epiphytic 

ferns are Humata heterophylla, Humata 

polypodioides, Asplenium nidus, Davallia solida, 

Davallia epiphylla, and Phymatosorus 

scolopendria. 

Like epiphytes, vines are more common in this 

forest than in coastal lowland forest, but less 

common than in montane forest. The most 

frequent trunk climbers are Freycinetia reineckei, 

Freycinetia srorckii, and Piper graeffei. Epi- 

premnum pinnatum, which is common on the

coastal ridges, decreases in importance with 

increasing clevation. In moister places, the terrestrial 

fern Lomagrumma cordipinna is also common 

ascending the lower portions of tree trunks. The most 

common lianas are Gynochtodes epiphytica, Alyxia 

hrucrcdo.su, and Hoyapottsii. 

6. Montane Forcst 

This is the rainforest occurring above about 350 m 

elevation and dominated by trees that are mostly 

absent at lower elevations. The highest point within 

the park is Mt. Alava (at 490 m elevation), so the 

upper limit of montane forest within the park is 

restricted to this elevation. It occurs mostly on the 

south edge of the park along Alava Ridge (Fig. 22) 

and on some of the upper valleys north of the ridge, 

mostly east of the TV transmitter on Mt. Alava where 

there is less disturbance. 

This community is the same one described on Ta'u 

(Whistler 1992b), and corresponds to the "Maotamea 

montane forest" and the "fega-vao cloud forest" 

described in an earlier publication (Whistler 1980). 

The floristic differences that serve to distinguish it 

from lowland forest are caused mostly by the higher 

elevation, and indirectly by rainfall and temperature. 

Little reliable climatic data has been recorded in this 

community or at this elevation, but if the decrease of 

0.65"C (1.17"F) per 100 m increase in elevation 

suggested by Wright (1963) for Western Samoa 

applies to Tutuila, the mean temperature is 2.3 to 

3.2"C (4.1 to 5.7'F) cooler there than at sea level. No 

rainfall data has been recorded within the park, but the 

rainfall in its montane forest may be as much or more 

than the 5000 mm (200 in) per year in Pago Pago 

Harbor, and is certainly greater than in the lowland 

forest north of it. 

When undisturbed, montane forest is similar in 

stature to lowland forest, but it has been severely 

damaged by the recent hurricanes (Fig. 23). But even 

before that, human disturbance took its toll-a road 

running from Fagasa Pass to the TV transmitter was 

built in about 1977, which has opened the area up to 

agriculture and erosion. The road, in fact, probably 

obliterated one of the marked plots set up during the 

1975 -1 976 inventory (Whistler 1980). Consequently, 

no undisturbed examples of it were found during the 

present survey. 

As noted above, the major canopy species of the 

lowland forest on ridges are Syzygium inophylloides 

(asi), Canarium vitiense (ma'ali), Canarium 

harveyi (ma foa?), Calophyllum neo-ebudicum 

(tamanu), Intsia bijuga (ifilele), and Buchanania 

merrillii. However, at higher elevations these trees 

become less important or disappear, and are 

replaced by species not found at lower elevations. 

The most characteristic species of the Tutuila 

montane forest are Dysoxylum samoense 

(maotamea), Syzygium samoense (fena vao), and 

Crossostylis biJ70ra (saitamu). Other important 

species include Hedycarya denticulata, Syzygium 

samarangense (nonu vao), Palaquium stehlinii 

(nasu, in Western Samoa), Myristica hypargyraea 

('atone), Elaeocarpus tonganus (a'amati'e), 

Trichospermum richii (ma'osina), Astronidium 

pickeringii, Weinmannia affinis, Fagraea berte- 

roana (pualulu), and Hernandia moerenhoutiana 

(pipi). 

Because no intact examples of montane forest 

were found during the present survey, no plots 

were sampled. However, in a plot sampled in 

montane forest on Alava Ridge in 1976 (Whistler 

1980), and shown in Appendix B as plot no. 12 

(recalculated so as to standardize it with the 

current plots) the dominant species, in order of 

relative dominance, were Dysoxylum huntii (22%), 

Crossostylis bzjlora (1 8%), Syzygium samoense 

(1 3%), Palaquium stehlinii (1 0%), and Hedycatya 

denticulata (9%). None of these trees are 

significant in forest at lower elevations. 

Dysoxylum huntii is the most characteristic tree 

of this community. It is also a dominant in the 

montane forest of Ta'u, as well as in Western 

Samoa, where it is the major timber tree of higher 

elevations (above 600 m) on Savai'i and 'Upolu. It 

is rarely found below 300 m elevation, where it is 

replaced by two other species of the same genus, 

Dysoxylum maota (maota) and Dysoxylum 

samoense (mamala). 

Syzygium samoense, the other characteristic 

canopy tree of the montane forest of Tutuila, is 

rarely found below 300 m elevation. In two plots 

sampled in montane forest on Olosega island 

(Whistler 1980), it ranked first and fourth in rela- 

tive dominance. Crossostylis biJlora, which is an 

inland species of the mangrove family Rhizopho- 

raceae, is more of an understory tree that is not 

reported from below 250 m elevation, and is not 

found in Manu'a.

Figure 23. Montane forest on Toa Ridge.

Scvcral other trees are characteristic of Alava ridge, 

but arc more heliophytic (sun-loving) species that do 

not do well in the shade of undisturbed montane 

forest. 'The three most significant of these are 

Me/ro.videros collina, Spiraeanlhemum samoense, and 

Glochrdion cuspidatum, Metrosideros, which is 

probably the same species that comprises most of the 

native forests in Hawai'i, dominates the scrubby 

vegetation found on the upper part of the steep slopes 

of Alava Ridge, especially around the TV transmitter. 

Spirueanthemum is also common there, and was found 

to be the dominant species in one type of cloud forest 

at I600 m elevation on Savai'i (Whistler 1978). 

Glochidion cuspidatum is more of a secondary forest 

tree closely related to Glochidion ram@orum, a tree 

that typically occurs in disturbed forests of the 

lowlands. Also common on these slopes are Neo- 

nauclea forsteri (afa), which is a secondary forest 

canopy species, and Syzygium samoense. 

Some of the ridge top from the TV transmitter to 

the beginning point of the old trail down to Vatia is 

currently dominated by thickets of Hibiscus tiliaceus 

(fau). This tree is common in many types of habitats, 

especially disturbed ones, and its presence is probably 

the result of previous cultivation along the stretch of 

ridge. Most of the rest of the ridge in this area is open 

montane forest consisting of scattered trees growing 

out of a dense ground cover. 

The distinction between the lower limit of the 

montane forest and the upper limit of lowland forest is 

not very clear, because there is a zone of variable 

width where the characteristic species of the two 

communities share dominance. In plot 11 (Appendix 

B) at 350 m elevation, for example, the five dominant 

trees were Calophyllum neo-ebudicum, Syzygium 

inophylloides, and Buchanania merrillii, which are 

lowland forest species, and Syzygium samoense and 

I-lernandia moerenhoutiana, which are montane forest 

species. 

In undisturbed montane forest, the ground cover is 

often dense, and is comprised of shade tolerant ferns. 

The most characteristic and abundant species is 

probably Lomagramma cordipinna (Fig. 24); other 

common species include Asplenium nidus, Asplenium 

polj~odon, Trichomanes spp., and Microsorium 

.sy/vdicum. 

Other plants present in the forest floor of montane 

Sorest include Angiopteris evecta (king fern, nase), 

Murallia smithii, and Cyathea vaupelii (olioli). 

Angiopteris and Marattia are very similar to each 

other-they have long thick fronds arising from a 

thick base. Cyathea is a small tree fern. These 

ferns are present in both undisturbed and dist- 

urbed forest. Also common in these places are 

Zingiber zerumbet (wild ginger, 'avapui), Helico- 

nia laufao (laufao), and Cordyline fruticosa (ti). 

Native grasses such as Centosteca lappacea (sefa), 

Oplismenus compositus, and Cyrtococcum 

oxyphyllum are also found in montane forest, most 

often in clearings or along trails. 

In the disturbed montane forest on Alava ridge, 

the native, shade tolerant ferns are outcompeted in 

the sunny conditions by introduced, weedy 

heliophytic species, the most common of which 

are Mikania micrantha (fue saina), Clidemia hirta, 

and Paspalum conjugatum (vao lima). Mikania is 

a vine that grows over low vegetation and even up 

in the lower branches of trees. Clidemia is a 

noxious shrub that is uncommon in lowland forest, 

but very common in the summit scrub community 

of Ta'u (although it apparently did not reach that 

island until after 1976). Also common here are the 

two Freycinetia species, which often form dense 

tangles that are traversed only with difficulty. 

The amount and variety of epiphytes is another 

characteristic that serves to distinguish montane 

forest from lowland forest. The difference is not 

only in the amount or biomass of epiphytes, but 

also in the number of species present. The 

montane forest is the most floristically diverse on 

Tutuila, not only for epiphytes, but also for trees 

and ground cover species. Because of the cooler, 

conditions in the montane forest, there is a pro- 

fusion of epiphytes that are mostly absent from 

lowland forest. By far the most important epi- 

phytes are ferns and orchids. Only one epiphytic 

vascular plant species reported here was neither a 

fern (or fern ally) nor an orchid-Procris pedun- 

culata, a member of the nettle family. It was not 

very common, however. 

Many of the trees on the less disturbed parts of 

Alava Ridge are covered with epiphytes. The 

most characteristic fern is probably Trichomanes 

humile, a filmy fern that cannot exist in drier 

forests. Also common are Hymenophyllum 

imbricatum (another filmy fern), Hymenophyllum

Figure 24. Lomagramma cordipinna dominating floor of montane forest.

poIvon/l~os, Dtrvulliu solidu (which occurs all the way 

down lo sea level), Humata heterophylla, Ctenopteris 

hlcchnoides, and Ctenopteris contigua. Mosses and 

liverworts are also common. 

Many different epiphytic orchids are found in this 

forest, but probably the most abundant one is 

llcribobium sludei. Other common species include 

Dcndrobium biflorum, Flickingeria comata, Liparis 

con~ylobulbon, Phreatia micrantha, Phreatia matthewsir, 

Bulbophyllum polypodioides, Bulbophyllum 

longiscapum, Glomera montana, Diplocaulobium 

fililohum, and Coelogyne lycastoides. 

Vines are also common in the montane forest. There 

are few lianas in the forest in the park, mainly because 

of the absence of a canopy. The most common 

species present are Faradaya amicorum (mamalupe) 

and Alyxiu hracteolosa (lau maile). Less common 

and snialler species include Medinilla samoensis, 

IMscorca hulbifcra, and Hoya vitiensis (fue selela). 

The most common trunk climbers are Freycinetia 

reineckei and Freycinetia storckii (both called 'ie'ie). 

Less common species include Piper graeffei ('ava'ava 

aitu). One fern, Lomagramma cordipinna, also 

becomes a trunk climber when it encounters a tree 

while growing across the forest floor (see Fig. 24). 

D. Summit Vegetation 

'The summit vegetation category comprises a single 

plant community on Tutuila, montane scrub, first 

described in Samoa by Whistler (1980). Another type 

of summit vegetation, "summit scrub" was described 

from Ta'u (Whistler 1992b); although this has a 

somewhat similar structure, it differs in origin and 

floristic composition. 

7. Montane Scrub 

'l'his is the scrubby vegetation in which trees are 

dwarfed and secondary in importance (biomass) to the 

dcnse tangle of ground cover of species-mostly ferns 

and climbers. In Samoa this community appears to be 

unique to Tutuila, where it is restricted to ancient 

"trachyte plugs" scattered around the islands. As 

noted in the earlier section on geology, trachyte plugs 

are the remains of old volcanic cores. Their unique 

white clayey soil is characterized by a deficiency of 

calcium, magnesium, and phosphorus. The white soil 

is noticeable from a distance on the cliffs that are 

often found in areas of montane scrub (see Fig. 5). 

There are several areas of trachyte with their 

associated montane scrub vegetation on Tutuila, 

the most significant of which are Mt. Matafao and 

Mt. Pioa. These two peaks, which reach 650 m 

(2142) and 520 m (17 18 ft) respectively, can 

easily be seen from Pago Pago Harbor, but are not 

within the boundaries of the park. The trees 

present are scattered and mostly shrublike, and 

most belong to species different from those found 

in the surrounding lowland forest. In an earlier 

survey of these vegetation types (Whistler 1980), 

the dominant trees recorded from montane scrub 

on Matafao, Pioa, and another area called Mt. Tau 

(near the airport) were Pandanus reineckei, 

Rapanea myricifolia, Syzygium brevifolium, 

Astronidium pickeringii, Metrosideros collina, and 

Alstonia paczjlca. The ground cover, or perhaps 

more aptly described, the vegetation between the 

scattered trees, was dominated by Dicranopteris 

linearis (false-staghorn fern, asaua), Dipteris 

conjugata (a fern reported in Samoa only from 

Matafao and Pioa), Duvallia epiphylla, and 

Freycinetia storckii ('ie'ie). 

Only a single trachyte plug is found within the 

park boundaries-Mt. Mauga-o-Tula behind the 

village of Vatia (Fig. 25). Its steep summit (a ridge 

with high points on either end) is reached only 

with difficulty via an ill-defined trail ascending the 

east-facing slope. This trachyte ridge is only half 

the elevation of Pioa and Matafao (less than 300 

m), yet it has some of the same species that 

characterize montane scrub. In a cursory sample 

on Mauga-o-Tula during the present survey, the 

dominant trees recorded were Metrosideros 

collina, Intsia bijuga (ifilele), Alstonia pacifica, 

Eurya pickeringii, Psychotria insularum 

(matalafi), and Erythrospermum acuminatis- 

simum. This is a mixture of lowland ridge forest 

species and montane scrub species. However, the 

dominant ground cover species is Dicranopteris 

linearis, which is characteristic of, although not 

entirely restricted to, montane scrub (Fig. 26). 

Also present, but of lesser importance, are Scleria 

polycarpa, Oplismenus hirtellus, and Lindsaea 

ens folia.

Figure 25. Mauga-o-Tula Ridge behind Vatia. 

Figure 26. Montane scrub vegetation on Mauga-o-Tula, Vatia.

Although Mauga-o-Tula lacks some of the chara- 

cteristic montane scrub species of other trachyte 

plugs, such as Matafao and Pioa, it still can be be 

classified as montane scrub because of the dominance 

of Metrosideros trees and by the ground cover 

dominated by Dicranopleris. 

Secondary Vegetation 

Secondary vegetation includes all those plant 

communities that are in a state of relatively rapid 

change in structure and species composition brought 

about by some kind of disturbance. The disturbance 

can be natural, as from hurricanes and landslides, or it 

can be caused by the activities of man, such as the 

clearing of forests for plantations and villages. Until 

the arrival of the first Polynesian settlers perhaps 3000 

years ago, all disturbance was natural. Nowadays, 

however, it is most often caused by the activities of 

man. 

Three secondary vegetation communities are 

recognized below. They are successional, i. e., over 

time after a disturbance, the first one, managed land 

vegetation, evolves into the second one, secondary 

scrub, and this in turn evolves into the third one, 

secondary forest. Species dominating disturbed 

vegetation have some characteristics in common: 

(I) they are relatively fast growing; (2) they have 

mostly small, readily dispersed seeds; (3) they are 

intolerant to shade and disappear when the large 

primary forest trees, or other secondary species, shade 

them out. The species of the first community, 

managed land vegetation, are mostly aliens, i.e., 

non-native, but those of the other two are mostly 

indigenous (but not endemic) species. 

The three communities are described below. They 

are basically the same as those included in the report 

for the Ta'u unit of the Park (Whistler 1992b), altho- 

ugh a few of the species are different (Ta'u has fewer 

species). 

8. Managed Land Vegetation 

Managed land comprises those areas that are 

actively managed by man, and includes villages, 

plantations, pastures, roads, and airports. The managed 

land within the park comprises only villages, 

plantations, and roads (and roadsides). Although the 

village of Vatia, which is surrounded by the park, is 

excluded from the park itself by a 200 ft elevation 

boundary line, there are several houses in Amalau 

Valley (possibly only four) above and below the 

road, and thus the land around these structures fit 

into the village category. The TV transmitter atop 

Mt. Alava and the area around a house along the 

Alava Ridge road, belong here as well. 

The plantations in the park are found mostly 

around the village of Vatia above the 200 ft 

elevational line (Fig. 27), perhaps some on the 

slope to the west of Afono, and some patches 

below and north of the Alava Ridge road. The 

most commonly cultivated plants in plantations 

(and villages) are coconuts, taro, and bananas. 

When coconut plantations are abandoned, the 

coconuts may persist for many decades, but 

eventually disappear because they do not compete 

well reproductively with rainforest trees. Taro and 

bananas are usually grown only for one or a few 

times on a piece of land cut out of the forest, after 

which the plot is abandoned to fallow. It is then 

rapidly overgrown with herbaceous weeds, and 

eventually by shrubs and small trees, at which time 

it is classified as secondary scrub (discussed 

below). 

Paved roads obviously have no vegetation, but 

their margins can be classified as managed land. 

The only paved road within the park runs from 

Afono to Vatia, and its margins are periodically 

cut or sprayed with herbicide. The road cut along 

this and other roads probably also would be cla- 

ssified as managed land, at least until they become 

vegetated with forest or scrub. 

The only other road within the park is the dirt 

road running from Fagasa Pass to the TV trans- 

mitter-the Alava Ridge road. It is passable only 

by means of 4-wheel drive vehicles because of 

erosion and mud puddles, but a family travels it 

regularly by truck to their house located part way 

along the road. The managed land vegetation of 

this road consists of the low plants on the road 

itself, the taller weedy plants along the margins, 

and the plants on the roadcuts along its route. 

Nearly all the plants found on managed land are 

introduced species. Before the arrival of the 

Europeans, the weeds introduced by the 

Polynesians and a few native species shared 

dominance. Now, however, most of the weeds, 

including the most dominant species, are modern 

introductions-mostly from tropical America. 

Because of the competition of these more recently

Figure 27. Vatia Village showing extent of managed land vegetation.

introduced weeds, many of the Polynesian weeds have 

been severely reduced or have disappeared from 

American Samoa. Polynesian weeds reported on 

'Tutuila but not collected on the island in the last half 

century include Amaranthus viridus (Amaranthaccae), 

Cyathula prostrata (Amaranthaceae), Senna 

.sophc~rti (I:abaccae), and Urcrria lagopodoides 

(Pabaceae). 

The dominant weedy plants found on managed land 

are Mikania micrantha (mile-a-minute vine, fue 

saina), Nephrolepis hirsutula (sword fern, vao 

tuaniu), Bidens alba (beggar's-tick), and Paspalum 

conjugatum (T-grass, vao lima). Many other species 

may be common, depending upon soil, moisture, and 

type of management. About 250 species introduced 

species have become naturalized in the archipelago 

(Whistler 1988b), and about 168 of these occur on 

Tutuila (Appendix A); most of these are weeds that 

thrive in sunny disturbed areas; most are flowering 

plants, but a few are ferns. Mikania, unlike the other 

three most abundant weed species, is a vine than can 

smother other herbaceous or shrubby vegetation, and 

may inhibit timely regeneration of forest from 

disturbed land. 

The dominant plants on the margins of the Alava 

Ridge road include Brachiaria mutica (California 

grass), Mikania micrantha, Calopogonium muco- 

noides, Mimosa pudica (sensitive plant, vao fefe), and 

Ludwigia actovalvis (primrose willow). There is very 

little if any current management, since the road is 

largely abandoned, so these are often mixed with 

shrubs and trees characteristic of secondary scrub. 

The roadway itself is dominated by a different set of 

species that do not thrive in the dense shade of the 

taller roadside plants; the most common plants here 

include Ludwigia octovalvis, Eleusine indica (goose 

grass, ta'ata'a), Cuphea carthagenensis, and Paspa- 

lum conjugatum. The roadsides from Afono to Vatia 

are dominated by weeds such as Desmodium tortuo- 

.sum (especially in places that have not recently been 

sprayed or cut), Bidens alba, Paspalum conjugatum, 

and Calopogonium muconoides. 

lioadcuts are similar to cliffs in that erosion on the 

steep surfaces prevents a stable climax vegetation 

from becoming established. On the Alava Ridge road, 

the dominant species are weedy vines, such as 

Mikania micrantha and Dioscorea bulbifera (bitter 

yam, soi), secondary scrub species, such as Pipturus 

argenteus, secondary forest trees, such as Neonau- 

clea forsteri (afa), and ferns, such as Pneumato- 

pteris glandulosa, Angiopteris evecta (king fern, 

nase), Pleisoneuron attenuatum, and Macrothe- 

lyptris polypodioides. Also common is the large 

' native reed Miscanthus floridulus (u). The road- 

cuts on the eastern part of the park are dominated 

mostly by secondary forest trees, such as Maca- 

ranga harveyana (lau pata), Pipturus argenteus 

(soga), and Morinda citrijiolia (Indian mulberry, 

nonu), and herbaceous species such as Dioscorea 

bulbifera (bitter yam, soi), Mikania micrantha, 

Nephrolepis hirsutula (vao tuaniu), and Macro- 

thebpteris polypodoides. In some places Dicra- 

nopteris linearis (asaua, false staghorn fern) is 

dominant, especially on cliffs forming the edge of 

the road. 

Other than the taro, coconut, and banana 

plantations around the fringes of the villages and 

along roads, there is hardly any current agricul- 

tural activity within the park boundaries. Only 

one person, a non-Samoan residing in Vatia, is 

currently known to have any inland plantations. 

One is on the hillside west of Amalau Valley, 

another is on Toa Ridge about halfway between 

the TV transmitter and the north coast, and another 

is reportedly on the ridge just to the west of Tafeu 

Bay. The Toa Ridge plantation is an area of 

Dicranopteris linearis, which is suggestive of 

ancient village or plantation sites. Currently only a 

few orange trees and a mango are planted there, all 

of them quite young. The area west of Tafeu Bay 

has some coconuts, possibly a remnant of former 

human activity (the name of the area is Nu'utoga, 

literally "Tongan village," which may indicate its 

ancient status). The farmer using this area 

believes that some of the trees there may be old 

cultivated species that he doesn't recognize, but 

unfortunately the area was not visited because of 

weather conditions. 

9. Secondary Scrub 

This is the type of disturbed vegetation that is 

dominated by shrubs and small trees. It is usually 

a successional stage between managed land, or 

land disturbed naturally (as by a landslide), and 

secondary forest. The species that dominate here 

are mostly woody rather than herbaceous. When 

the land is managed, or weeded in the case of 

plantations, these woody plants are kept in check,

since they are perennial species that take longer to 

grow and rcproduce. The lower, smaller, and faster 

growing herbaceous plants are dominant under these 

conditions, since weeding never eliminates weeds-at 

best, it controls and prevents them from overrunning 

the crop. Ifowever, when management has ended, or 

aftcr a landslide, the disturbance has ended, the small 

shrubs and trees soon take over and eventually shade 

out the herbaceous species, other than the vines. 

Vines found in secondary vegetation can be divided 

into two types-herbaceous vines and woody vines 

(lianas). Herbaceous vines are common in secondary 

scrub, since they can cover the shrubs and low trees. 

However, when taller trees prevail, herbaceous vines 

cannot climb high enough and are generally shaded 

out below the canopy. Lianas are less common in 

secondary scrub than they are in secondary forest, 

since they lack the large trees to climb, but most of 

them probably get their start at this stage. 

The most common secondary scrub trees, more or 

less in order of abundance, are Hibiscus tiliaceus 

(beach hibiscus, fau), Macaranga harveyana (lau 

pata), Pipturus argenteus (soga), Morinda citrifolia 

(Indian mulberry, nonu), Caricapapaya (papaya, esi), 

Omalanthus nutans (fogamamala), Kleinhovia 

hospita (fu'afu'a), and Trema cannabina (magele). 

Most of these have relatively large, ovate or 

heart-shaped leaves and do not grow to be very tall. 

Ilihiscus is a very adaptable tree, since it is 

common in disturbed littoral forest, in thickets on the 

margins of mangrove swamps and streams, and in- 

land on disturbed hillsides. Its distinctive gray-green 

color and growth form make it easy to spot from a 

distnncc. It is often found in secondary forest as well, 

commonly as scattered, scrambling individuals. Its 

prcscnce there probably indicates major disturbance in 

the past. 

The largest areas of secondary scrub in the park are 

on the ridges north of Alava Ridge. These are mostly 

abandoned plantations that are now overgrown with 

scrubby vegetation. It should be kept in mind, how- 

ever, that the vegetation categories outlined here are 

not distinct entities and often blend into each other. 

The vegetation on the ridges is often a mixture of 

scrub and scattered lowland forest trees that have 

survived the recent hurricanes. 

10. Secondary Forest 

This vegetation comprises high forest that is 

structurally somewhat similar to primary forest, 

but is dominated by secondary rather than primary 

forest species. Secondary species were noted earlier 

to typically have relatively small, easily dispersed 

seeds. They may be either bird dispersed, 

with succulent hits that are attractive to birds, or 

wind dispersed with very light or winged seeds or 

fruits that are transported by the wind. Secondary 

species are also characterized by their transitory 

dominance-they do not perpetuate themselves in 

areas lacking disturbance. 

Primary species, on the other hand, have larger 

seeds that give the seedling extra stored energy to 

insure a greater survival rate on the shady forest 

floor. When the forest canopy becomes establi- 

shed, the small seeds of secondary forest species 

are at a disadvantage for survival, and their repro- 

duction is inhibited. Primary forest species repro- 

duce in their own shade and maintain their domi- 

nance until the next major disturbance. The 

seedlings themselves may also have different 

tolerances to shade, with the primary forest spe- 

cies being adapted at the seedling stage to relati- 

vely low light intensities. 

The transitory nature of the dominance of 

secondary forest canopy trees in their own forest is 

reflected in population dynamics; although they 

dominate the largest size classes, they are severely 

under-represented in the smaller size classes 

-which include the trees that will eventually 

inherit the canopy. For example, at a sampled plot 

at Faleiulu on Ta'u (Whistler 1980), the two 

dominant trees were both secondary species; all 24 

sampled individuals were over 15 cm dbh. On the 

other hand, 30 of the sampled individuals were 

primary forest canopy species, but only 11 of these 

were over 15 cm dbh. Barring further disturbance 

over the short run, the primary forest trees will 

eventually replace the secondary forest species in 

that plot. Secondary forest also has less species 

diversity, since the forest is younger and has not 

had sufficient time for all the species formerly 

there to become re-established. 

A complicating factor in the distinction between 

primary and secondary forest is called "gap 

replacement." When a tree falls in primary or

even mature secondary forest, it leaves an opening in 

thc canopy it may even take a few other canopy 

individuals with it when it falls. The canopy opening 

allows secondary forest trees to become established in 

primary forest. However, many primary forests lack 

any of the major secondary forest species, which 

possibly indicates the gaps routinely occurring in 

forests do not allow enough light to favor secondary 

forest species. 

The most characteristic secondary forest species in 

Samoa is Rhus taitensis (tavai), which grows to be a 

huge tree. Large areas of forest dominated by this tree 

occur on Tutuila. It can be recognized from a distance 

by its layered branches tinted red with young leaves, 

and in aerial photographs by the even texture of the 

canopy. It may dominate alone, or it may share 

dominance with another species, Alphitonia 

zizyphoides (toi). Other typical secondary species 

include Canunga odorata (ilangilang, moso'oi), 

Bischqfia juvanica ('o'a), Elattostachys falcata 

(taputo'i), Neo-nauclea forsteri (afa), and 

Macarangm stipulosa (lau fatu). However, these 

rarely become dominant, and the latter species is 

probably typical of gap replacement. 

Several other trees are somewhat intermediate 

between primary and secondary forest species. Dyso- 

xylum maola (maota) and Dysoxylum samoense 

(mamala) are species that appear in young secondary 

forest. In alluvial valleys they may completely domi- 

nate, but elsewhere they may disappear or comprise 

only a minor part of the relative dominance. Pometia 

pinnalri (tava) is a dominant primary forest species on 

area of rocky soil from recent lava flows (i.e., as it 

was on thc 'fafuna plain before the native vegetation 

was dccimatcd), but on hillsides it appears to be a 

pioneer species. This is a moot point, however, 

because I'ometia has not been recorded from the north 

side of the island and consequently not within the park 

boundaries. Another species should also be 

mentioned, Paraserianthes falcataria. This intro- 

duced tree is common along the beginning part of the 

Alava Ridge road, but since it has small, poorly 

dispersed seeds, it does not readily spread. 

The most significant secondary forest in the park is 

found on the slopes below Alava Ridge road and 

above Fagasa Bay. The dominant species here is 

probably Neonauclea forsteri (afa), but nearly as 

common are several other species, including Dyso- 

xylum samoense, Rhus taitensis, Alphitonia zizy- 

phoides, Myristica fatua (Samoan nutmeg, 

'atone), Bischofia javanica ('o'a), and Cananga 

odorata. It is not apparent why these ridges are 

covered with secondary forest. There is no sign of 

current agriculture, although there is on the ridges 

farther east along the road (see Fig. 20), and the 

presence of large trees suggests that agriculture is 

not the cause. Although the area seems far enough 

above Fagasa to be unlikely as a plantation area, it 

may have been utilized for this long ago-long 

enough ago that no signs of any plantation exist 

now. Perhaps the most likely explanation is that 

the steep slopes are unstable, resulting in a 

"disclimax" vegetation of secondary forest. This is 

somewhat similar to the "Dysoxylum lowland 

forest on talus slopes and cliffs" described from 

Ta'u (Whistler 1992b). The disturbed condition 

may have been abetted by erosion from the ridge 

road. 

Another area of secondary forest is found on the 

slope below the road just to the west of Amalau 

Valley. The dominant species here is probably 

Dysoxylum maota (maota), but also common are 

Cananga odorata, Kleinhovia hospita, Garuga 

jloribunda (vivao), Planchonella samoensis (ma- 

malava), and Ficus scabra (mati). This forest is a 

combination of ridge lowland forest, valley 

lowland forest, and secondary forest species, and 

is possibly on land that was formerly cultivated. 

DISCUSSION AND 

RECOMMENDATIONS 

Two or three decades ago, Samoa had the 

highest percentage of intact native vegetation of 

any Polynesian archipelago. This may still be 

true, but since then nearly all of the forests of the 

lowlands (up to about ca. 400 m elevation) have 

been felled for their timber or cut and burned for 

agriculture to keep up with the population growth 

of the islands. 

The park is important because of the native 

vegetation it contains. Some of the best lowland 

forest remaining in the archipelago is found on 

Tutuila. It occurs in two main areas, one within 

the park boundaries between Fagasa and Afono, 

the other on the northwest Tutuila coast between 

the villages of Fagamalo and Fagasa. These two 

areas are similar in being mostly uninhabited (both

have one village in the middle) and consist of rugged 

coastlines and ridges covered with native forest. Thus 

the park represents a significant portion of the 

remaining lowland forest in American Samoa. It also 

represents a significant portion of the quality lowland 

forest of the whole archipelago as well, since much of 

the remaining lowland forest of Western Samoa was 

devastated by the recent hurricanes, while most of that 

within the Tutuila park is in relatively good condition. 

In addition to the lowland forest within the park, there 

is one area of another unique plant community, 

montane scrub, a vegetation type that is restricted in 

the archipelago to Tutuila. 

The park is also important for floristic reasons. The 

native flora of the whole archipelago consists of about 

540 species of flowering plants and 230 species of 

ferns and fern allies. Tutuila is home to 57% of the 

native flowering plants of the archipelago and 50% of 

the native fern and fern allies. The park itself contains 

39% (209 species) and 35% (81 species), respectively. 

There are 49 species of native orchids on the island, a 

number similar to that found in the rest of Polynesia 

(outside of Samoa) combined, and 30 of these have 

been recorded in the park. Consequently, this 

relatively small park on a small island comprising less 

than 5% of the total land area of the archipelago, is 

home to a significant percent of the total native flora 

of Samoa. 

The need to protect this area, and other sensitive 

areas is heightened because of the recent rapid 

population growth. While the population of Western 

Samoa is rising only slowly because of out-migration, 

and the population in Manu'a is decreasing, the population 

of Tutuila is rapidly increasing. Over the last 

10 years, the population has increased 44% (Craig 

1992) to about 46,600 (as of 1990) and at this rate will 

top 100,000 by the year 2010. Much of this 

population has settled in the Tafuna area, where the 

unique Pometia lowland forest that once covered the 

whole area has now been almost entirely wiped out. 

There will be increasing pressure for land, and for the 

development of roads into hitherto inaccessible areas. 

Thus the need to protect the few remaining areas of 

native vegetation cannot be over-emphasized. 

By merely designating an area as a National Park is 

not enough to protect the plants and animals in the 

area. The concept of a park is unheard of in the 

traditional "fa'asamoa," and the conflict that arises is 

complicated by the Samoan land tenure system and 

ancient concepts of land use. Unlike the situation 

with bats and some bird species, there are no 

individual plant species that are threatened by 

human activity at this time. But the plants are 

threatened by the loss of their pollinators and 

dispersers-typically the birds and bats. Their 

habitat is threatened. Thus the following 

recommendations are made. 

1. Strict enforcement of hunting laws. 

There are laws on the books for American 

Samoa that prohibit shooting birds and bats, but 

such laws do not conform to Samoan tradition and 

for the most part are ignored. This lack of local 

support leads to weak enforcement. Even though 

there is a law against hunting birds and bats, there 

is no prohibition against importing shotgun shells, 

which are hardly used for anything but birds and 

bats. As far as enforcement in the park is concerned, 

stiffer penalties, including federal prison 

terms (if possible) may be effective, since nothing 

else has been found to work. Honest and diligent 

enforcement officers are needed-that have the 

integrity and courage to enforce laws and apprehend 

violators, even if the offenders are members 

of their own extended families. Perhaps some kind 

of village enforcement could be feasible, if the 

villagers can be made to understand how the loss 

of wildlife will eventually effect them. 

2. Control of feral pigs within the park. 

There is currently a problem of feral pigs in 

parts of the park. Well-traveled pig trails were 

seen on some of the ridges. In the past the feral 

population was kept in check by local hunters, but 

with increasing urbanization and rising material 

culture, pig hunting has decreased to practically 

nothing. Perhaps a bounty on feral pigs, or hired 

hunters who, are allowed to keep the slain pigs, 

would be effective. The feasibility of traps and 

exclosures should also be studied. 

3. Limiting vehicular access to the park. 

Most of the disturbance within the park is along 

the Alava Ridge road. As the pressure on land 

increases in the future, individuals may claim and 

clear even more land along the road, destroying 

the native vegetation. While traditional farming is 

allowed by the park agreement, there is no need to 

make it easier by improving access. The road 

should be blocked so that few if any vehicles can 

use the road (this would have to be worked out 

with families currently farming the area or living

there). No increased access should be promoted at 

this time, until it is certain that it will not lead to 

additional development of the interior of the park. 

Access roads to new projects in Samoa have always 

opened up the roadside areas to destruction. 

4. Establishing a list of endangered plant and 

animal species. 

There is currently no Federal list of endangered 

species in American Samoa. Federal regulations 

apply to Guam, but so far have not been used in 

American Samoa. This has probably been due to lack 

of local interest in such a listing. The Federal 

government and the American Samoa government 

should be prodded into becoming involved; the least 

they could do is make a list of candidate species of 

animals and plants that are rare or endangered; this 

could be culled from Appendix A of this report. 

5. Other critical areas that may need protection. 

It may be feasible to extend the boundaries of the 

park to include other critical habitats or areas of native 

vegetation. Two areas adjacent to the park, Mt. 

Matafao and Mt. Pioa, have a unique native montane 

scrub vegetation that is home to a number of rare 

Samoan species. On the other hand, because these 

two areas are on poor soil, have steep slopes, and are 

relatively inaccessible, they may not need further 

protection. A much larger project, left to a later time 

(to see if the current park works out), is to study the 

feasibility of expanding the park to include the area 

from Fagasa to Fagamalo on the northwest Tutuila 

coast, which is largely native lowland forest. 

6. Control of harmful weeds. 

There are 96 alien plant species recorded from the 

park, most of which are modern introductions 

(introduced during the European Era beginning about 

1830) rather than Polynesian introductions. A few of 

these are trees that are naturalized in the forest, but the 

vast majority are weeds adapted to living in disturbed 

places. Two of the weeds are particularly common in 

the park. Mikania micrantha (mile-a-minute vine, fue 

saina) has been present in Samoa for nearly a century, 

but many foresters believe that it retards forest 

regeneration because of its smothering growth. It is 

one of the few introduced weeds that occurs in 

undisturbed native forest. It is not much of a problem 

within this kind of forest, since it does not thrive in 

the shade, but in cut-over areas it is. Farmers don't 

mind it, in taro anyway, since it is easy to pull up and 

clear. The other plant is Clidemia hirta (Koster's 

curse). Although it is found in lowland forests, it is 

not very common there. However, in montane 

forest, where it is wetter, it is a dominant species 

in sunny places. It is a very serious problem on 

the top of a Ta'u, despite the fact that it has 

apparently been on that island for less than 20 

years (Whistler 1992b). These two weeds may be 

good candidates for biological control. 

There are a number of other potentially harmful 

species present in Western Samoa, such as Fun- 

tumia elastica (pulu vao) and Solanum torvum, 

that are not yet found in American Samoa. These 

should not be allowed into the country under any 

circumstances. Another potentially harmful 

species, Castilla elastica (pulu mamoe) is an 

invasive species in Western Samoa, and was 

recently recorded from Tutuila (near Fagamalo) 

and Ta'u (personal observation, 1993). It should be 

eliminated before it becomes further established. 

7. Control of other pests. 

In reaction to the establishment of the African 

snail on Tutuila, the predatory snail Euglandina 

was introduced as a biological control. Although 

it may help with the African snails, it unfortu- 

nately is leading to the demise of some of the 

native land snails on the island. A similar 

sequence of events has recently taken place on 

Ta'u as well (Whistler 1992b). Some native 

species are currently known from only a few 

isolated localities within the park. A comprehen- 

sive study of the snails within the park, as well as 

on the whole island, is needed, to see what can be 

done to save the threatened species. 

ACKNOWLEDGMENTS 

The author would like to give special thanks to 

Natasha Bartley for her invaluable help with the 

field work during the first part of the project. He 

would also like to thank Dr. Pepper Trail, Dr. 

Peter Craig, Doug Cuillard, Rory West, and Mike 

Harrington for their assistance during parts of the 

field work, and Henry Sesepasara for making 

available the facilities and staff of the Department 

of Marine and Wildlife Resources. 

This research was supported under a co- 

operative agreement (CA 8034-2-0001) between 

the University of Hawai'i at Manoa and the 

National Park Service.

LITERATURE CITED 

Amerson, A. B., Jr., W. A. Whistler, & T. D. Schwaner. 1982. Wildlife and wildlife habitat of American 

Samoa. I. Environment and ecology; 11. Accounts of flora and fauna. U. S. Department of the Interior, 

Washington D. C. 

Biosystems Analysis, Inc. 1992. A comprehensive wetlands management plan for the islands of Tutuila 

and Aunu'u, American Samoa. Econ. Develop. Planning Office, Government of American Samoa. 

Brownlie, G. 1977. The pteridophyte flora of Fiji. J. Cramer, Vaduz, Germany. 397 pp. 

Chandler, K. C., A. T. Larsen, & F. P. Wallis. 1978. The forest resources of Western Samoa. P. F. Olsen & 

Co. Ltd., Rotorua, new Zealand. 2 Vols. 

Christensen, C. 1943. A revision of the Pteridophyta of Samoa. Bernice P. Bishop Mus. Bull. 177: 1-138. 

Christophersen, E. 1935, 1938. Flowering plants of Samoa. Bernice P. Bishop Mus. Bull. 128: 1-221; 11. 

154: 1-77. 

Cole, T. G., C. D. Whitesell, W. A. Whistler, N. McKay, & A. H. Ambacher. 1988. Vegetation survey and 

forest inventory, American Samoa. Pac. Southwest Forest and Range Experiment Station, Berkeley. 

14 PP. 

Craig, P. 1992. How many people can American Samoa support? Editorial, Samoa News, 4 April. 

Nakamura, S. 1984. Soil survey of American Samoa. US. Dept. Agriculture, Soil Conservation Service. 

95 PP. 

National Park Service. 1988. National park feasibility study: American Samoa. ,U. S. Dept. of the Interior. 

Ollier, C., W. A. Whistler, & A. B. Amerson, Jr. 1979. '0 le Pupu-Pu'e National Park. U. N. Develop. 

Adv. Team for the So. Pacific; Suva, Fiji. Vol. I-Main Report, 79 pp. Vol.2-Interpretive Material, 83 

PP. 

Park, G., R. Hay, A. Whistler, and T. Lovegrove. 1992. The national ecological survey of Western Samoa. 

Department of Conservation, Wellington, N. Z. 205 pp. 

Pickering, C. 1876. The geographical distribution of animals and plants in their wild state. (From U.S.E.E. 

Vol. 19, pt. 2: 276-3 1 1). Naturalists' Agency, Salem, Mass. 

Powell, T. 1868. On various Samoan plants and their vernacular names. J. Bot. 6: 278-285, 342-347, 

355-370. 

Rechinger, K. 1907-1 9 15. Botanische und Zoologische Ergebnisseeiner wissenschaflichen Forschungsreise 

nach den Samoa-Inseln, ... Denkschr. Akad. Wiss. Wien 81: 197-317. 1907; 84: 385-562. 1908; 85: 

175-432. 1910; 88: 1-65. 1911; 89: 443-708. 1913; 91: 139-213. 1915. 

Reinecke, F. 1896, 1898. Die Flora der Samoa-Inseln. Bot. Jahrb. 23: 237-368; 25: 578-708. 

Setchell, W. A. 1924. American Samoa. Part I. Vegetation of Tutuila Island: Part 11. Ethnobotany of the 

Samoans; Part 111. Vegetation of Rose Atoll. Publ. Carnegie Inst. Wash. 341: (Dept. Marine Biol. 20): 

1-175. 

Smith, A. C. 1979-1991. Flora vitiensis nova: a new flora of Fiji. National Trop. Bot. Garden, Lawai, 

Kaua'i, Hawai'i. 5 Vols. 

Stearns, H. T. 1944. Geology of the Samoan Islands. Bull. Geolog. Soc. Amer. 55: 1279-1332.

Whistler, W. A. 1976. Wetlands of American Samoa. US. Army Corps of Engineers, Honolulu. 74 pp. 

(Mimeograph) 

Whistler, W. A. 1980. The vegetation of eastern Samoa. Allertonia 2(2): 45-190. 

Whistler, W.A. 1983. Vegetation and flora of the Aleipata Islands, Western Samoa. Pac. Sci. 37(3): 227- 

249. 

Whistler, W. A. 1984. Annotated list of Samoan plant names. Econ. Bot. 

Whistler, W. A. 1986. A revision of Psychotria (Rubiaceae) in Samoa. Jour. Am. Arb. 67: 341-370. 

Whistler, W. A. 1988a. A revision of Syzygium (Myrtaceae) in Samoa. Jour. Am. Arb. 69: 167-192. 

Whistler, W. A. 1988b. Checklist of the weed flora of Western Polynesia. So. Pacific Commission Techn. 

Paper 194. Noumea, New Caledonia. 69 pp. 

Whistler, W. A. 1992a. The vegetation of Samoa and Tonga. Pacific Sci. 46(2): 159-178, 

Whistler, W. A. 1992b. Botanical inventory of the proposed Ta'u unit of the National Park of American 

Samoa. Techn. Rept. 83. Cooper. Nat. Park Resources Studies Unit, Univ. Hawaii. 85 pp. 

Wright, A. C. S. 1963. Soils and land use of Western Samoa. New Zealand Dept. Indust. & Sci. Res., Soil 

Bur. Bull. 22: 1-191. 

Yuncker, T. G. 1945. Plants of the Manua Islands. Bernice P. Bishop Mus. Bull. 184: 1-73.

APPENDIX A 

ANNOTATED CHECKLIST OF THE FLORA OF TUTUILA 

'Thc following checklist of the flora of Tutuila is 

based on field work carried out on the island from 

27 May to 12 June 1992 and 17 December 1992 to 5 

January 1993, as well as on collections of all bota- 

nists since 1839. Not all of the species listed have 

been recorded within the park; some, especially 

those restricted to habitats not existing in the park, 

are highly unlikely to occur there. Species not yet 

recorded from the park are marked with an asterisk; 

specimens collected within the park are followed by 

an exclamation point. 

The species on the checklist are divided into four 

groups, Pteridophyta (ferns), Fern Allies, Dicoty- 

ledonae, and Monocotyledonae. Each of these 

taxonomic categories is further divided into plant 

families arranged in alphabetical order; the species 

within the families are also arranged alphabetically. 

Each entry includes the scientific name, common 

names (if any; English names, when present, are in 

brackets), a brief description, habitat, reported 

elevation, geographic range, and a list of all 

specimens noted from the island. Because the 

average number of specimens collected for each 

species is low, the elevations listed are those 

recorded for the species in the whole archipelago. 

The cited specimens, all of which are from 

Tutuila, can be found in the collections of the 

Bishop Museum, the Smithsonian Institution, the 

National Tropical Botanical Garden, and the Univer- 

sity of Hawai'i (the author's personal collection). A 

voucher set was also prepared for the National Park 

Service. Specimens noted in previous publications 

but not verified by the author are noted as "n.s." (not 

seen). Specimens lacking a collection number are 

noted as "s.n." (without number). 

The taxonomy employed in the list follows The 

Pteridophyte jlora of Fiji (Brownlie 1977) for the 

ferns and fern allies, and Flora vitiensis nova (Smith 

1979-1991) for the flowering plants, with only a 

few exceptions. The Samoan names were taken 

from Whistler (1 984).

Acrostichum aureum L. 

[swamp fern] sn'ato 

PTERIDOPHYTA 

AD1 ANTIACEAE 

I,argc, ercct, leathery fern up to 2 m or more in 

height with pinnate, leathery fronds and sori densely 

covering the lower surface of the upper, fertile 

pinnae. Common on coastal rocks and in coastal 

marshes, recorded only near the coast. Indigenous, 

pantropical in distribution. McMullin 43; Setchell 

93, 238; Eames T23; Garber 967; Bryan 967, 980!; 

Yuncker 9366, 9423; Lamoureux 3028; Whistler 

2823,2853,3292, 8906!. 

Adiantum philippense L. 

[maidenhair fern] 

Delicate terrestrial fern with shiny black to dark 

brown stipes, pinnate fronds with a lamina 15-20 cm 

long, and sori extending all along the outer margins 

of the fertile lobes. Uncommon in shady places, 

reported from 50 to 200 m elevation. Indigenous, 

widely distributed in the Old World tropics. 

Whistler 8412, 9355!. 

Bolbitis palustris (Bracken.) Hennipman 

Medium-sized terrestrial fern with a short, creeping 

rhizome, dimorphic pinnate fronds, margins cut 

about 113 of the way to the midrib into rounded 

lobes, and sori covering the entire lower surface of 

the pinnae of fertile fronds. Occasional to locally 

common in shady forest, reported from 30 to 600 m 

elevation. Indigenous, reported from Australia to 

Rapa. Reinecke 5c; Whistler 3534, 3616, 9078, 

0121. 

Pityrogramma brackenridgei (Carruthers) Maxon 

Small terrestrial fern with a short rhizome covered 

with linear brown scales, fronds tufted, lamina 

pinnatifid, and sori covering the silvery 

undersurface of the lobes. Occasional in sunny 

places such as roadcuts, reported from near sea level 

to 1550 m elevation. Endemic. A Kew specimen 

was determined by Tyron in 1965 as P, chrysophylla 

(Sw.) Link; if this is to be followed, the Samoan 

plant would be indigenous rather than endemic. 

OSEE s.n.; Setchell 118, 237; McMullin 42; 

Lamoureux 3029; Whistler 3753, 8064, 8408, 

8615!, 8988!. 

Pteris comans Forst. f. 

Medium-sized terrestrial fern with a short, erect 

rhizome, deltoid-ovate, deeply bipinnatifid fronds, 

reticulate venation, lanceolate lobes, and sori 

arranged along the entire margin. Occasional to 

common in littoral to lowland forest, reported from 

near sea level to 200 m elevation. Indigenous, 

ranging from Australia to Polynesia. Setchell 124; 

Steams s.n.; Garber 973; McMullin 26; Long 3098; 

Lamoureux 3099; Whistler 2909, 8422,9033!. 

Pteris ensiforrnis Bum. f. 

Small terrestrial fern with a creeping rhizome, 

clumped pinnate fronds, 3 or 4 pairs of long, 

narrowly lanceolate, finely toothed lobes (fertile 

ones often 3-branched), and sori continuous along 

the margins. Occasional in coastal and lowland 

forest, reported from near sea level to 200 m eleva- 

tion. Indigenous, ranging from India to Polynesia. 

Reinecke 15a; Setchell 108; Garber 877; Yuncker 

9339; Whistler 8452, 8540!, 8693!. 

Pteris pacifica Hier. 

Medium-sized terrestrial fern with an erect rhizome, 

clumped, deeply pinnatifid, glossy fronds with up to 

8 pairs of lateral pinnae cut to near their midrib into 

narrowly lanceolate, rounded lobes, and sori nearly 

continuous along the pinnae margins. Occasional in 

coastal to lowland forest, reported from 20 to 200 m 

elevation. Indigenous, ranging from the Solomon 

Islands to Polynesia. Reinecke 14a; McMullin 9; 

Garber 872; Wilder s.n.; Dumas 8; Kuruc 143; 

Whistler 8446, 8574!, 903 1 !, 9128. 

Pteris tripartita Sw. 

Large terrestrial fern up to 2 m or more in height, 

with a short, erect rhizome, tripartite fronds, open 

venation, narrowly lanceolate pinnae mostly 2-3 cm 

wide and cut to near the midrib, and long sori 

extending to near the sinus on the lower edge of the 

lobes. Occasional in disturbed places and forest, 

reported from near sea level to 800 m. elevation. 

Indigenous, ranging from Africa to Tahiti. Veitch 

s.n.; McMullin 31; Whistler 3777, 8435, 8595!, 

9031!, 9117. 

*Pteris vaupelii Hier. 

Large terrestrial fern with an erect rhizome, deeply 

pinnatifid, glossy fronds with lateral pinnae cut to 

near the midrib into narrowly lanceolate, rounded

lobes, and sori nearly continuous along the pinnae 

margins. Uncommon in lowland to cloud forest, 

reported from near sea level to 1700 m elevation. 

Endemic. Whistler 9092. 

ASPIDIACEAE 

Arachniodes aristata (Forst. f.) Tindale 

Medium-sized terrestrial fern with a scaly, creeping 

rhizome, deltoid, tripinnate and quadripinnate 

fronds, aristate segment tips, and indusiate sori in a 

single row on each side of the midvein of the ulti- 

mate segments. Uncommon in lowland to montane 

forest, reported from 50 to ca. 600 m elevation. 

Indigenous, ranging from southern Africa to Tahiti. 

Whistler 8554!, 8645!. 

Ctenitis samoensis (Christen.) Holttum 

Medium-sized terrestrial fern with a creeping 

rhizome, deltoid, quadripinnate fronds, tomentose 

ribs, medial sori on veins, and small, reniform, caducous 

indusia. Uncommon in lowland to montane 

forest, reported from 240 to ca. 600 m elevation. 

Indigenous, also found on Rarotonga. Reinecke 

92? (n.~.); Sledge 1829; Whistler 8591!, 8555!. 

Pleocnemia leuzeana (Gaud.) Presl 

Large terrestrial fern with an erect rhizome, fronds 

up to 4 m in height, lamina bipinnate to tripinnatifid, 

yellow glandular hairs present on the undersurface 

of the minor veins, and large exindusiate sori. 

Occasional in lowland to montane forest, reported 

from near sea level to 1000 m elevation. Indigenous 

to Samoa, ranging eastward to Indonesia. Graeffe 

s.n. (n.~.); Yuncker 9432!?; Whistler 8063, 8498, 

8741!, 9071. 

Tectaria setchellii Maxon 

Large terrestrial fern with a decumbent rhizome, 

unwinged stipes, deltoid pinnate-bipinnate lamina 

25-40 cm long, 2-5 pairs of lobed pinnae, and small, 

irregularly scattered sori lacking indusia. Occa- 

sional in coastal and lowland forest, reported from 

near sea level to 250 m elevation. Endemic. Veitch 

s.n.; Setchell 119, 360; Wilder s.n.; Yuncker 9364, 

94 1 1 ; Long 309 1, 3 107; Whistler 8423, 8425, 

8550!. 

Tectaria stearnsii Maxon 


tufted fronds, obovate, mostly bipinnatifid lamina 

30-50 cm long, small sori at the ends of short 

vcinlets, and usually with no indusia. Common in 

coastal and lowland forest, reported from near sea 

level to 450 m elevation. Endemic. Veitch sun.; 

Reinecke s.n.; Setchell 40, 208, 227; Garber 927; 

Meebold 8261, 16863; Wilder s.n.; Yuncker 9322, 

9433, 9440; Wisner 16, 62, 63, 64; Whistler 3844, 

8424, 8465, 8700!, 9124. The difference between 

this and T. dissecta Forst. f. found on adjacent 

islands needs to be explored. 

ASPLENIACEAE 

Asplenium australasicum Hooker 

Large epiphyte or terrestrial fern similar to A. nidus, 

but with the midrib on lower surface angled rather 

than rounded. Occasional in lowland to montane 

forest, reported from 250 to 390 m elevation. Indigenous, 

widespread in the South Pacific. Whistler 

9337!, 9344!. 

Asplenium cuneatum Lam. 

Terrestrial fern with a short-creeping rhizome, 

bipinnate lamina up to 25 cm long, pinnae up to 12 

on each side and merging into a lobed apex, 1-4 

elongate sori on each pinnule, and entire indusia. 

Occasional in coastal to cloud forest, reported from 

50 to 1550 m elevation. Indigenous, widespread in 

the tropics. Reinecke 3f; Setchell 395; Meebold 

16619; Dumas 17; Garber 844, 854; Long 1956, 

1975, 1997b; Kuruc 147; Whistler 8556!, 9074, 

9130. 

*Aspleniurn feejeense Bracken. 

Terrestrial fern with simple, narrowly lanceolate 

fronds up to 50 x 7 cm, tip attenuate and ending in a 

terrestrial, proliferous bud, and sori in parallel rows 

between the margin and midrib. Occasional in 

coastal to montane forest, reported from 50 to 900 m 

elevation. Endemic. Yuncker 9408; Kuruc 144. 

Asplenium horridum Kaulf. 

Terrestrial fern with pinnatifid fronds up to 2 m 

long, hairy stipes, numerous, narrowly lanceolate 

pinnae 10- 15 cm long and lobed to near the midrib, 

and linear sori arranged several to a lobe.

Uncommon in montane to cloud forest, reported 

from 400 to 1500 m elevation. Indigenous, ranging 

from Malaysia to Hawai'i. Reinecke 76b; Whistler 

9049!, 9085. 

Asplenium lascrpitiifoliurn Lam. 

I,arge terrestrial f'crn with a stout, long-creeping 

rhizome, tufted, dark brown or black stipes, a 

deltoid, tripinnate to quadripinnatifid lamina up to 

60 x 20 cm, rhomboid-trapezoid ultimate segments, 

and 1 or 2 linear sori on the ultimate lobes. Occa- 

sional in lowland to montane forest, reported from 


ranging from the Mascarenes to Polynesia. 

Setchell 21 3; Garber 926; Yuncker 9319; Whistler 

3843, 8062, 8577!, 91 18. 

Aspleniurn nidus L. 

[bird's-nest fern] laugapapa 

L,arge epiphytic or terrestrial fern with simple, 

sessile, glossy fronds up to 2 m long arranged in a 

rosette, basal portion of the midrib rounded on the 

lower surface, and numerous, closely parallel rows 

of sori on the lower surface of the frond from the 

midrib to halfway to the margin. Common in littoral 

to cloud forest, reported from near sea level to 1100 

m elevation. Indigenous, found throughout the Old 

World tropics. Reinecke 103b; Setchell 13; Whistler 

2838,8704!, 9 126. 

Asplenium polyodon Forst. f. 

Small terrestrial fern with a short-creeping rhizome, 

tufted fronds, deltoid, pinnate lamina 10-45 cm 

long, up to 15 pairs of falcate to lanceolate, acumi- 

nate pinnae, and linear sori almost covering the full 

length of the vein. Common in lowland to montane 

forest, reported from near sea level to 700 (1300) m 

elevation. Indigenous, ranging from Madagascar to 

Polynesia. Reinecke 37a?, 58; Setchell 47; Garber 

855, 864, 865; Eames TI8 (n.s.); Yuncker 9321, 

9377; Wisner 65; Whistler 8528!, 9120. 

Aspleniurn tenerum Forst. f. 

Small to medium-sized epiphytic or terrestrial fern 

with a short, erect rhizome, tufted fronds, pinnate 

lamina cut into 15-30 pairs of toothed, round-tipped 

oblong pinnate, and linear sori along the veins from 

near the midrib to near the margins. Uncommon in 

lowland to montane forest, reported from 30 to 700 

m elevation. Indigenous, ranging from Ceylon to 

Polynesia. Yuncker 9327; Whistler 3533, 9129, 

9352!. 

*Asplenium unilaterale Lam. 

Small terrestrial fern with a creeping rhizome, shiny 

black stipes, linear-lanceolate, pinnate fronds up to 

20 x 4 cm having a pinnatifid tip, 20 or more pairs 

of pinnae, and numerous linear sori. Uncommon in 

coastal to montane forest, reported from 180 to 700 

m elevation. Indigenous, ranging from East Asia to 

Polynesia. Reinecke 35b, 35f; Whistler 9073. 

ATHYRIACEAE 

Diplazium bulbiferum Bracken. 

Medium-sized terrestrial fern with a short, erect 

rhizome, pinnatifid lamina, broadly lanceolate 

pinnae up to 15 cm long divided to near the midrib 

into oblong, toothed lobes, and linear sori on most 

of the veins of the lower surface of the lamina. 

Uncommon in lowland forest, reported from 50 to 

ca. 400 m elevation. Indigenous, ranging from 

Vanuatu to Samoa. Wilder s.n.; Whistler 8492, 

8575!. 

Diplazium dilatatum BI. 


broadly deltoid, pinnate or bipinnate lamina up to 50 

x 20 cm, largest pinnules distinctly lobed, forked 

veins bearing linear sori, and indusia almost ciliate 

in appearance. Uncommon in lowland to cloud 

forest, reported from 300 to 1400 m elevation. 

Indigenous, ranging from India to Polynesia. 

Whistler 3601, 8526!, 8652!, 9157. 

Diplazium harpeodes Moore 

Large terrestrial fern with an erect rhizome, bipin- 

natifid to tripinnatifid lamina up to 1 m in length, 

lanceolate pinnules cut about halfway to the midrib 

into blunt, often toothed lobes, and linear sori on 

most of the veins of the lower surface. Uncommon 

in lowland to cloud forest, reported from near sea 

level to 1300 m elevation. This species is quite 

variable and may need further taxonomic work. 

Indigenous, ranging from Fiji to Pitcairn Island. 

USEE s.n.; Veitch s.n.; Setchell 204; Long 1982; 

Whistler 3625, 3846?, 3847, 8635!, 9075, 9076, 

913 1.

Diplaziurn proliferurn (Lam.) Thouars 

Large terrestrial fern with a short, stout, erect 

rhizome, spiny stipe and rachis, bulbils often 

present, pinnate lamina up to 1 m in length, lanceo- 

late pinnae with shallowly lobed margins, and sori 

along all the veins of the lower surface. Uncommon 

in lowland to montane forest, reported from near sea 

level to 600 m elevation. Indigenous, ranging from 

tropical Africa to Samoa. Graeffe 444; McMullin 

15, 16; Setchell 15; Garber 836; Bryan 1377; Kuruc 

148; Whistler 8447, 8527!, 8596!, 9139. 

*Lunathyriurn japonicurn (Thunb.) Kurata 

Medium-sized fern with scaly stipes and rhizomes, 

deeply bipinnatifid fronds, short sori along the veins 

midway between the costule and margins, and thin, 

entire indusia. Uncommon in lowland to cloud 


Indigenous, ranging from Japan to Polynesia. 

Veitch sen.; Whistler 9077. 

BLECHNACEAE 

* Blechnurn doodioides (Bracken.) Brownlie 

Medium-sized terrestrial fern with erect an rhizome, 

tufted stipes, dimorphic pinnatifid fronds, basal 

segments of sterile fronds reduced to rounded lobes, 

pinnae of fertile fronds narrowly lanceolate, and sori 

linear and parallel to the costa. Locally common in 

sunny areas in montane to cloud forest, reported 

from 750 (but lower if actually occurring on 

Tutuila) to 1650 m elevation. Indigenous, ranging 

from Southeast Asia to Samoa. Reinecke 145a. 

This record should be checked. 

Blechnurn orientale L. 

Large terrestrial fern with a stout, erect rhizome, 

simply pinnate fronds up to 2 m in height, and sori 

in a continuous line on either side of the midrib. 

Common in lowland to montane forest, reported 

from ca. 50 to 700 m elevation. Indigenous, ranging 

from tropical Asia to the Austral Islands. Setchell 

28; Eames TI 6; Christophersen 103 1, 35 16; Kuruc 

149; Whistler 861 1 !, 8907!. 

Blechnurn vulcanicurn (Bl.) Kuhn 

Large terrestrial fern with an erect rhizome, dimor- 

phic pinnate fronds, sterile fronds cut nearly to the 

midrib into narrowly lanceolate, finely parallel- 

veined lobes, and sori covering the entire lower 

surface of the linear lobes of the fertile fronds. 

Occasional in montane scrub to cloud forest, 


ranging from Malaysia and New Zealand to the 

Marquesas. Christophersen 1129!, 3561. 

CYATHEACEAE 

*Culcita straminea (Labill.) Maxon 

Large terrestrial fern with a large prostrate to sub- 

erect stem, tufted fronds up to 4 m long, deltoid, 3- 

to 4-pinnate lamina, quaternary segments cut into 

acute lobes bearing 1 or sometimes 2 sori, and an 

indusium of 2 valves. Locally common in montane 

forest and scrub, reported from 300 to 650 m eleva- 

tion. Indigenous, ranging from Vanuatu to central 

Polynesia. Setchell 389; Christophersen 1027, 

354 1; Sledge 1825; Whistler 9060. 

Cyathea decurrens (Hooker) CopeI. 

Small tree fern with large fronds, smooth stipes, 

upper surface of costa and costules densely hairy, 

bipinnate lamina, ultimate segments deeply lobed, 

and globose sori lacking indusia. Uncommon in the 

montane forest, reported from 450 to 910 m eleva- 

tion. Indigenous, ranging from Australia and New 

Caledonia to the Cook Islands. Whistler 9046!. 

Cyathea lunulata (Forst. f.) Copel. 

[tree fern] olioli 

Tree fern with large fronds having warty, somewhat 

glacous stipes, bipinnate lamina with pinnules lobed 

to the costules, linear-lanceolate lobes 2-3 mm wide, 

and inconspicuous indusia. Common in disturbed 

forest, reported from 20 to 900 m elevation. Indige- 

nous, ranging from the Marianas to Samoa. Setchell 

2, 73, 225; McMullin 32; Yuncker 9417, 9424; 

Whistler 8982a!. 

*Cyathea truncata (Bracken.) Copeland 


Tree fern with densely tomentose and warty stipes, 

tripinnate lamina, tertiary pinnules small and 

petiolulate, small sori, and rudimentary indusia of 

thin scales hidden beneath the sori. Occasional to 

common in montane scrub, reported from 500 to 

1300 m elevation. Indigenous, ranging from the 

Solomon Islands to Samoa. Powell 1 19; Reinecke

171 ; Setchell 393; Christophersen 1029, 353 1; 

Whistler2717, 3551, 8481,9061,9194. 

Cyathea vaupelii Copeland 


'I'rue fcrn with large fronds, smooth rachises, 

bipinnale lamina with pinnules lobed to the costules, 

oblong lobes mostly 4-5 mm wide, and conspicuous 

indusia. Occasional in lowland to montane forest, 

reported from 50 to 820 m elevation. Endemic. 

Reinecke 89d; Setchell 2, 225; McMullin 14; Eames 

'r2 1 ; Christophersen 3546; Yuncker 9430; Whistler 

3550, 8517!, 8674!, 8901!. 

DAVALLIACEAE 

Arthropteris repens (Bracken.) Christen. 

Epiphytic or terrestrial fern (typically on lower tree 

trunks) with a long-creeping, thin rhizome, pinnate 

fronds up to 30 x 5 cm, and sori between the costa 

and the margin. Common in lowland to cloud 

forest, reported from near sea level to 1500 m 

elevation. Indigenous, ranging from Borneo to 

Samoa. Reinecke s.n.; Whistler 3532, 8578!, 9123. 

Davallia epiphylla (Forst. f.) Spreng. 

Terrestrial or epiphytic fern with a widely creeping 

rhizome covered with hairlike scales, lamina broadly 

deltoid, 4-5-pinnate, fertile segments bifid at the 

apex, sori immersed, and an indusium with a free 

acuminate tip. Common in lowland to montane for- 

est, reported from near sea level to 650 m elevation. 

Indigenous, ranging from Fiji to Tahiti. McMullin 

12; Setchell 36; Wilder s.n.; Yuncker 9303; Long 

1978, 3 106; Wisner 58; Kuruc 150; Whistler 3545, 

8703,8715!, 8983!, 9154. 

*Davallia plumosa Baker 

Medium-sized epiphytic fern with a densely scaly, 

creeping rhizome, non-ciliate rhizome scales, usual- 

ly tripinnate fronds, immersed sori, and cylindrical 

indusia with the rounded outer edge reaching the 

edge of the segment or a little longer. Uncommon 

in the montane to cloud forest, reported from 900 to 

1650 m elevation. If the record is correct, this 

elevation would have to be down to at least 650 m. 

Endemic to Samoa. Reinecke s.n. (ns.). A que- 

stionable record. 

Davallia solida (Forst. f.) Sw. 

Epiphytic fern with a stout, creeping rhizome, 

tripinnate to tripinnatifid, deltoid fronds, sori on 

terminal subdivisions of lobes, and tubular indusia. 

Common in lowland to cloud forest, reported from 


ranging from Malaysia to eastern Polynesia. 

Reinecke s.n.; McMullin 2 1; Setchell 6 1; Garber 

824; Diefenderfer 7; Whistler 2825, 8702!. 

Humata heterophylla (Smith) Desv. 

Epiphytic fern with a long-creeping rhizome, 

dimorphic fronds, simple, lanceolate, finely veined 

sterile fronds, lobed fertile fronds having marginal 

sori, and thin indusia. Occasional to locally 

abundant in lowland to montane forest, reported 

from near sea level to 700 m elevation. Indigenous, 

ranging from Sumatra to Polynesia. Safford 940; 

Setchell 280; Garber 827, 881; Lamoureux 3016; 

Long 1966; Whistler 28 19,3016, 8538!. 

Humata polypodioides Bracken. 

Epiphytic fern with a long-creeping rhizome cov- 

ered with appressed, brown scales, uniform pinnate 

fronds with lamina and stipe of similar length, and 

terminal, submarginal sori covered with an indusium 

attached only at the base. Uncommon in lowland to 

montane forest, reported from near sea level to 700 

m elevation. Indigenous, also found in Fiji and 

Vanuatu, and possibly New Guinea. Safford 16; 

McMullin 18; Setchell 2 19; Garber 826; Yuncker 

9347; Long 1976b; Whistler 2820, 8681 !, 8701 !. 

"Humata serrata Bracken. 

Epiphytic fern with a long-creeping rhizome cov- 

ered with narrow scales, dimorphic, 2-3-pinnate 

deltoid fronds, terminal, submarginal sori covered 

with an indusium and attached only at the base, and 

an acute "horn" extending beyond the sori. Uncom- 

mon in montane to cloud forest and montane scrub, 


also found in New Caledonia, or perhaps endemic to 

Samoa. USEE s.n. (n.s.); Powell 136; Whistler 

9087. 

Nephrolepis biserrata (Sw.) Schott 

Medium-sized epiphytic or terrestrial fern with a 

short rhizome, erect, pinnate fronds up to 2 m in 

length, scales on young stipes, pinnae with crenate 

margins lacking an auricle, sori between the margins

and the costules, and reniform indusia. Common in 

lowland to montane forest, reported from near sea 

level to 1000 m elevation. Indigenous, pantropical 

in distribution. Reinecke 172; Setchell 18, 209; 

Christophersen 1032; Yuncker 9330; Long 1992, 

3047; Whistler 3544, 8979!. 

Nephrolepis hirsutula (Forst, f.) Presl 

Medium-sized terrestrial fern with a short, thick 

rhizome, erect pinnate fronds up to 1.2 m long, 

rachis and surfaces usually with paIe scales, pinnae 

with an auricle and crenate margins, sori near the 

pinnae margins, and reniform indusia. Common in 

disturbed places, reported from near sea level to 

1550 m elevation. Indigenous, ranging from 

tropical Asia to Polynesia. Reinecke lb; Setchell 

3 17; Garber 873; Christophersen 3499; Yuncker 

9367; Whistler 9099!, 9140. 

Oleandra neriiformis Cav. 

Epiphytic fern with a long brittle rhizome, simple 

linear-lanceolate fronds with close, conspicuous 

veins, sori in a single row close to the midrib, and 

reniform indusia. Occasional to common in mon- 

tane forest to cloud forest, reported from 500 to 

1500 m elevation. Indigenous, ranging from tropi- 

cal Asia to Samoa. Veitch s.n.; Reinecke 108b; 

Setchell 398; Whistler 3549, 8739!, 9086. 

DENNSTAEDTIACEAE 

Orthiopteris tenuis (Bracken.) Brownlie 


tufted deltoid, compound fronds, grooved rachis and 

costa, and terminal marginal sori enclosed within a 

cup-like indusium d~awn downwards into a point 

with the margins fused to the segment. Uncommon 

in montane forest, reported from 400 to 900 m elevation. 

Indigenous, ranging from Samoa to New 

Guinea. This genus is in need of revision. Brownlie 

(1977) noted that this name may apply only to an 

endemic Fijian species. Whistler 9050! 

Dicranopteris linearis (Bum. f.) Underwood 

[false-staghorn fern] saua 

Large terrestrial fern with a creeping, reddish-hairy 

rhizomes, fronds forked 1-3 times, lamina bipin- 

natifid, segments narrowly oblong, and sori with up 

to 15 sporangia. Occasional to common in disturbed 

places, fernlands, and montane scrub, reported from 


widely distributed in the Old World tropics and 

subtropics. Setchell 110, 367? (n.~.); Garber 868; 

Wilder s.n.; Christophersen 1026, 3493; Mitchell 

5 16; Y uncker 9344; Wisner 145; Lamoureux 3026; 

Whistler 2929, 86 1 O!, 8686!. 

GRAMMITIDACEAE 

Ctenopteris blechnoides (Grev.) Wagner & Grether 

Small epiphytic fern with a short-creeping rhizome, 

a black rachis, lanceolate, pinnate lamina less than 

1.8 cm wide, triangular sterile lower lobes, linear- 

oblong upper fertile lobes, and oblong to round sori 

sometimes nearly covering the segment. Occasional 

in mangrove to montane forest, reported from near 

sea level to 900 m elevation. Indigenous, ranging 

from Ceylon to Tahiti. Reinecke 176; Setchell 391 

p.p.; Garber 787; Christophersen 1054, 12 17, 3553; 

Sledge 1827; Whistler 2788, 8944, 8997!, 9045!, 

9340!. 

Ctenopteris contigua (Forst, f.) Holttum 

Small epiphytic fern with a short rhizome, clustered, 

winged stipes, lanceolate, subpinnate lamina lobed 

to near the midrib, and sori embedded at the tip of 

the upper lobes. Occasional in lowland to cloud 

forest, reported from 300 to 1500 m elevation. Indi- 

genous, ranging from Ceylon to Tahiti. Reinecke 

124a; Christophersen 1 127!, 12 16, l825!; Whistler 

8547!. 8943!. 

*Ctenopteris lepidum (Brause) ..... 

Small epiphytic fern with rhizome short-creeping, 

stipes 2-6 cm long, lamina lanceolate, subpinnate, 

lower segments triangular and sterile, upper ones 

linear and fertile, and exindusiate oblong or round 

sori almost covering undersurface of segment. 

Occasional in montane forest, reported from 450 to 

725 m elevation. This species has apparently not yet 

been transferred to the genus Ctenopteris. Endemic. 

Reinecke 166b; Setchell 39 1 p.p.; Christophersen 

1055, 1072; Whistler 9084. 

*Grammitis hookeri (Bracken.) Copeland 

Small epiphytic fern with a short-creeping rhizome, 

reddish hairy stipes 2-4 cm long, simple, lanceolate 

lamina, and round sori in a single row on either side

of the costa. Uncommon in montane to cloud forest, 


also found in I-lawai'i and Fiji. Sine coll, s.n. (at 

Kew); Christophersen 3543. 

HY MENOPHYLLACEAE 

Ilyn~cnopliyllu~n irnbricatum BI. 

Medium-sized filmy fern with a widely creeping 

rhizome, winged rachis, spaced fronds up to 25 cm 

long, 2- to 3-pinnatifid lamina, lobes narrowed 

below the tip, and a globose sorus enclosed within 

the suborbicular lips of the indusium. Locally 

common in montane to cloud forest, reported from 

500 to 1600 m elevation. Indigenous, ranging from 

Sumatra to Tahiti. Setchell 392; Long 3059; 

Whistler 8636!, 8735!, 9043!. 

Hymenophyllum polyanthos Sw. 

Small filmy fern with a widely creeping, wiry 

rhizome, winged rachis, spaced fronds 5-25 cm 

long, 2- 3-pinnatifid lamina, lobes wider than the 

sorus, and the obovate sorus enclosed within the 

rounded lips of the indusium. Occasional in 


level to 1000 m elevation. Indigenous, widely 

distributed in the wet tropics. Reinecke 175; 

Christophersen 1056, 113 1 !, 1195, 1196, 1214, 

3529; Whistler 8543!, 8713!, 8998!, 9042!, 9089!. 

*Hyrnenophyllum praetervisum Christ 

Small epiphytic filmy fern with a long-creeping 

rhizome, unwinged stipes, thin fronds, lamina 1-2 

cm wide with toothed margins, and a sorus enclosed 

within a turbinate indusium having dentate margins. 

Uncommon in montane to cloud forest, reported 

from 500 to 1300 m elevation. Endemic. Reinecke 

88b. 

"Trichomanes apiifolium Presl 


stipes covered with long reddish hairs, 3-4-pinnati- 

lid fronds up to 50 cm high, and a sorus enclosed 

within a tubular indusium. Uncommon in montane 


genous, ranging from the Philippines and Malaysia 

to Samoa. Veitch s.n.; Reinecke 115, 174. 

*Trichomanes bimarginatum van den Bosch 

Tiny epiphytic fern with a creeping rhizome covered 

with dark, simple hairs, short, dark hairy stipes, a 

simple, oblong to obovate lamina less than 2 cm 

long, 1-3 sori on the lamina apex, and an immersed 

tubular indusium. llncommon in montane to cloud 

forest, reported from 400 to 1100 n~ elevation. 

Indigenous, ranging from Ceylon to Samoa. 

Reinecke sen. 

Trichomanes bipunctatum Poiret 

Small epiphytic fern with a slender, long-creeping 

rhizome, deltoid-ovate, tripinnatifid fronds 4-8 cm 

long with a submarginal, false vein in the segments, 

and a sorus enclosed within a tubular indusium. 

Uncommon in lowland to cloud forest, reported 

from near sea level to 1200 m elevation. Indige- 

nous, ranging from Madagascar to Tahiti. Reinecke 

135; Christophersen 1 145!; Whistler 8699!, 8991 !, 

9334!. 

Trichomanes boryanum Kuntze 

Small to medium-sized terrestrial fern with an erect 

rhizome, tufted, pinnate fronds with narrowly 

oblong pinnae, and a sorus enclosed within a tubular 

indusium located on the upper margins of the upper 

pinnae. Occasional to common in lowland to mon- 

tane forest and in streambeds, reported from near 


from the Vanuatu to Samoa. Garber 884 (n.s.); 

Christophersen 112 1 !, 1144!; Meebold 16865; 

Yuncker 93 15; Long 3 112; Whistler 8644!, 8731 !. 

*Trichomanes caudatum Bracken. 

Small epiphytic filmy fern with a creeping rhizome, 

a rachis winged in the upper part only, lanceolate 

tripinnatifid fronds 5-30 cm long, sori conspicuous 

on short axillary segments or terminal, and a cylin- 

drical indusium. Rare in montane to cloud forest, 


ranging from Australia to Tahiti. Veitch s.n. 

Trichomanes dentatum van den Bosch 

Small terrestrial fern with an erect rhizome, 

clustered fronds with a tripinnatifid, broadly deltoid 

lamina up to 20 cm long, few hairs on the rachis,


Uncommon in lowland to montane forest reported 


from New Caledonia to Polynesia. Yuncker 9323; 

Whistler 8710!, 8743!, 9006!. 

*'llriclwmuncs cndlichcrianum Prcsl 

Small epiphytic fern with a slender creeping 

rhizome, winged rachis, irregularly bipinnatifid 

fronds 1.5-6 cm long, 2-6 pairs of narrowly oblong 

to lanceolate pinnae, margins with 2 rows of 

specialized cells, and a sorus enclosed within a 

tubular indusium. Apparently rare in montane forest, 


ranging from Norfolk Island and New Zealand to 

Tahiti. Reinecke 135. 

Trichomanes humile Forst. f. 

Small epiphytic fern with a slender, long-creeping 

rhizome, winged rachis, bipinnatifid fronds 2-5 cm 

long, lobed margins with 2 rows of elongate cells, 


Common in littoral to cloud forest, reported from 


ranging from Malaysia to Tahiti. Reinecke s.n.; 

Garber 940; Christophersen 1146!; Yuncker 9333; 

Long 3 1 19; Whistler 8529!, 8944!, 9095!, 9336!, 

9350!. 

Trichomanes intermedium van den Bosch 

Medium-sized fer~ with an erect rhizome, stipe 

having caducous wings, fronds tufted, lamina 

tripinnate to quadripinnatifid, 8-30 cm long and 

bearing some appressed hairs, ultimate segments 

narrow, and the sorus enclosed within a tubular 

indusium. Occasional in lowland to montane forest, 


Indigenous, ranging from New Guinea to Samoa. 

Setchell 218; Garber 843, 885; Wilder s.n.; Christophersen 

18 15 !; Mitchell 5 18; Yuncker 9406; 

Whistler 8638!. 

*Trichomanes maximum B1. 

Medium-sized fern with a short-creeping rhizome, 

stipes having caducous wings, lamina quadripinnate, 

ultimate segments less than 0.5 mm wide, and the 

sorus enclosed within a tubular indusium. Uncom- 

mon in lowland to cloud forest, reported from 300 to 


Malaysia to Tahiti. Setchell218; Garber 733?. 

Trichomanes saxifragoides Presl 

Small fern with a wiry, creeping rhizome, flabeli- 

form, reniform fronds mostly less than 1.5 cm long 

and wide, and a sorus enclosed within a tubular 

indusium. Locally common in lowland to montane 


elevation. Indigenous, ranging from Africa and 

Japan to Hawai'i. Reinecke 88b; Whistler 8648!. 

*Trichomanes taeniatum Copeland 

Small epiphytic fern with a creeping rhizome, fronds 

digitate or dichotomous, margins ciliate but without 

a specialized margin or false vein, and sori enclosed 

within a tubular indusium. Rare in montane scrub 

on and probably montane forest elsewhere in 

Samoa, reported from 450 m elevation. Indigenous, 

also found in the Society Islands. Sledge 1824. 

Trichomanes tahitense Nadeaud 

Small epiphytic, prostrate fern with a creeping rhi- 

zome, sessile, peltate fronds adhering to the tree 

surface, veins radiating from the point of attach- 

ment, and 1-3 sori enclosed within a tubular indu- 

sium. Occasional in lowland to montane forest, 

reported from ca. 20 to 500 m elevation. Indige- 

nous, ranging from Java to eastern Polynesia. 

Whistler 3646, 8490, 8591!, 9335!. 

HYPOLEPIDACEAE 

Histiopteris incisa (Thunb.) J. Sm. 

Large, often scrambling terrestrial fern with a long- 

creeping rhizome, shiny black stipes, tripinnatifid 

fronds, pinnules cut into triangular lobes with the 

lowest ones like stipules, and sori continuous along 

the margins except at the tips. Common in montane 

to cloud forest and montane scrub, reported from 


widely distributed in the Southern Hemisphere 

tropics and temperate zone. Setchell 394; Whistler 

9O37!. 

*Hypolepis tenuifolia (Forst. f.) Bemhardi 

Large terrestrial fern with a densely hairy, long- 

creeping rhizome, fronds broadly deltoid or ovate, 

quadripinnate, and sori covered by a pseudoindu- 

sium formed by the reflexed margin. Uncommon in 

sunny places in lowland to cloud forest, reported 

from 50 to 1700 m elevation. Indigenous, ranging

liom China to I'itcairn Island. USEE s.n.?; Setchell 

10 I?, 300. 'l'he identity of these specimens has not 

been verified. 

LINDSAEACEAE 

Lindsaca ensifolia Sw. 

ssp. agatii (Bracken.) Kramer 

Small terrestrial fern with a creeping rhizome 

clothed in reddish brown scales, pinnate to bipinnate 

lamina having 8-1 5 pinnae per side, anastomosing 

veins, and sori covered with an indusium and nearly 

continuous on the pinnae margins. Occasional to 

locally common in littoral to montane forest and 

fernlands, reported from near sea level to 600 m 

elevation. Indigenous, ranging from Australia and 

the Marianas to Samoa. Sledge 1826; Whistler 

2975, 3752, 3756, 8662!, 8730!, 9192. 

Lindsaca harveyi Carruthers ex Seem. 

Medium-sized terrestrial fern with a short-creeping 

rhizome, bipinnate or rarely simply pinnate fronds 

10-50 cm long, 1-5 pinnae to a side with a similar 

terminal one, an oblong sorus one to a lobe on 2-4 

nerve endings, and an indusium not quite reaching 

the margin. Uncommon in lowland to montane 

forest, reported from ca. 100 to 1000 m elevation. 

Indigenous, ranging from New Caledonia to Samoa. 

Setchell 272; Garber 883; Christophersen 1828!, 

35 14; Whistler 3655, 8942, 8995! Another species 

reported from Tutuila, L. pacifica Kramer, is 

virtually indistinguishable from this and its 

existence there as a separate entity is doubthl. 

* Lindsaca pulchra (Bracken.) Carruthers ex Seem. 

Small terrcstrial or epiphytic fern with a long- 

creeping rhizome covered with reddish brown 

acuniinatc scales up to 3 mm long, some veins 

anastomosing, lamina simple, 15-40 pinnules per 

side, sorus 1 to each lobe on 2-4 vein endings, and a 

delicate indusium not reaching the margin. Uncom- 

mon in lowland to cloudforest, reported from 320 to 


Bismarck Archipelago to Samoa. USEE sen.; 

Reinecke 72a. 

Lindsaea tetragona Kramer 


rhizome clothed in yellowish brown triangular 

Scales, fronds bipinnate with 3-9 pairs of lateral 

pinnae and a similar terminal one, veins free, and 

marginal sori covered by an indusium. Uncommon 

in lowland to montane forest, reported from 300 to 


Celebes to Tahiti. USEE s.n.; Tetens s.n.; Graeffe 

s.n.; Whistler 9340!. 

Microlepia speluncae (L.) Moore 

Medium-sized terrestrial fern with a creeping rhi- 

zome, tripinnate lamina, narrowly deltoid pinnules, 

secondary pinnules with crenate or lobed margins, 

and sori situated at the sinuses and covered with an 

indusium. Uncommon in sunny places in lowland to 


m elevation. Indigenous, pantropical in distribution. 

Setchell 90, 359; Bonaparte 4393; Whistler 8592!, 

9122. 

LOMARIOPSIDACEAE 

Elaphoglossum feejeense Bracken. 

Medium-sized epiphytic fern with a thick, short- 

creeping rhizome, leathery, simple, oblanceolate 

fronds attenuate at the base, and sori covering the 

entire undersurface of the fertile fronds. Uncommon 

in lowland to montane forest, reported from ca. 500 

m elevation. Indigenous, also found in Fiji. Chri- 

stophersen 1204, 1210, 3489, 3520; Whistler 8727!, 

8941, 9052!. The difference between this and E. 

reineckei Hier. et Lauterb. of the other Samoan 

islands is not always apparent. 

Lomagramma cordipinna Holttum 

Medium-sized terrestrial or epiphytic fern with a 

long-creeping or climbing rhizome, pinnate imma- 

ture fronds, pinnate to bipinnate mature fronds, and 

sori covering the lower surface of fertile pinnae. 

Common to locally abundant in lowland to montane 


tion. Indigenous, also found in Fiji. Safford (ac- 

cording to Holttum); Reinecke 55b; Wilder sun.; 

Long 1977; Sledge 1831; Whistler 2967, 8489, 

8594!, 91 74. 

MARATTIACEAE 

Angiopteris evecta (Forst. f.) Hoffman 

nase 

Large terrestrial fern with fronds up to 3 m in length 

that radiate from a short, massive, fleshy stock,

ipinnate lamina, lanceolate pinnules 10-20 cm long 

having an enlarged base, and sori of 7-12 sporangia 

free from each other, in a dense row 1-2 mm from 

the margins. Common in lowland to cloud forest, 


Indigenous, ranging from Malaysia to Tahiti. Saf- 

Sortl 404 I; McMullin 17; Comstock s.n.; Sctchell 

38, 207; Eames 727; Garber 838; Long 1950; Whi- 

stler 8975!, 9125. 

Angiopteris opaca Copeland 

nase 

Large terrestrial fern similar to the preceding spe- 

cies, but with rachis and stipe densely scaly rather 

than bearing scattered hairs. Occasional in lowland 

to montane forest, reported from 50 to 600 m eleva- 

tion. Indigenous, also found in Fiji. Christophersen 

99 1 ; Whistler 9047!, 9198. 

Marattia smithii Mett. 

nase 

Large terrestrial fern similar to the preceding two 

species, but with sporangia fused laterally to form a 

single structure. Uncommon in montane forest, 


ranging from Vanuatu to Samoa. Wilder s.n.; 

Sledge 1823; Whistler 8734!, 9090,9197. 

OPHIOGLOSSACEAE 

Ophioglossum pendulum L. 

Epiphytic fern with a small rhizome, pendulous, 

strap-shaped fronds 40-120 cm long, and sori 

embedded in a spike attached to the center of the 

frond. Uncommon in coastal to montane forest, 


Indigenous, ranging from Madagascar to Hawai'i. 

Lamoureux 3001; Sledge 1822; Whistler 2818, 

2926, 300 1, 8746!. 

*Ophioglossum reticulatum L. 

Tiny terrestrial fern with a single ovate lamina and 

sori embedded in a single erect spike. Uncommon 

in lowland to cloud forest, reported from 300 to 

1700 m elevation. Indigenous, ranging from tropi- 

cal Asia to Polynesia. Powell 127. 

POLYPODIACEAE 

*Dipteris conjugata Reinwardt 

Large terrestrial fern with a creeping rhizome, re- 

mote stipes,veins dichotomously branched, lamina 

divided to the base into two spreading halves, these 

divided into four lobes which arc further divided, 

and numerous sori scattered on the lower surface. 

Locally abundant in montane scrub, reported from 


Thailand to Samoa. Reinecke 173; Setchell 380; 

Garber 784; Christophersen 1034, 3564; Sledge 

1832; Whistler 2714,9162. 

*Lemmaphyllum accedens (Bl.) Donk 

Small epiphytic fern with a long-creeping rhizome, 

short stipes, dimorphic fronds, the sterile ones 

elliptic to ovate, the fertile ones similar but with an 

attenuate tip, and sori in a single row on both sides 

of the midrib of the upper attenuate part. Occasional 

in lowland to montane forest, reported from 300 to 

900 m elevation. Indigenous, ranging from Malaysia 

to Samoa. Graeffe 459 (n.s.); Powell 93? ("Manua 

and Tutuila"); Whistler 9093, 9186. 

Microsorium sylvaticum (Bracken.) Copeland 

Large terrestrial fern with a creeping rhizome, sim- 

ple, deeply lobed, simply pinnate fronds up to 2 m 

in height, and numerous tiny sori scattered 

irregularly over the lower surface of the lobes. 


from 25 to 850 m elevation. Indigenous, apparently 

ranging eastward to Tahiti. Kuruc 151; Whistler 

3842, 8501,8557!. 

Phymatosorus nigrescens (Bl.) Pichi Serm. 

Medium-sized terrestrial fern with a stout, creeping 

rhizome, frond deeply cut into 1-10 pairs of lobes 

(or sometimes simple and lanceolate), and deeply 

embedded sori forming conspicuous tubercles on the 

upper surface. Occasional in lowland to montane 


genous, ranging from Fiji to Tahiti. Christophersen 

997; Long 1996d; Whistler 8643!, 9132. 

Phymatosorus scolopendria (Burm, f.) Pichi Serm. 

lau auta; lau magamaga 

Large terrestrial or epiphytic fern with a long- 

creeping, scaly rhizome, simple, glossy-green

lamina deeply cut into 1-10 lobes, and large round 

sori in I or 2 rows on either side of the midrib of 

each lobe. Common to abundant in littoral to 


m elevation. Indigenous, ranging from tropical 

Africa to the Marquesas. Setchell 62; Garber 804; 

Wildcr s.11.; C'on~slock 4; Yuncker 9385; Whistler 

2824, 8630!. 

Polypodium subauriculatum Bl. 

1,arge epiphytic fern with a creeping rhizome clo- 

thed in dark brown scales, pinnate pendant fronds up 

1 m long, lanceolate pinnae, and immersed sori in a 

single row on both sides of the costa. Uncommon in 


ni elevation. Indigenous, ranging from tropical Asia 

to Samoa. Reinecke 165; Meebold 16862; Whistler 

3637!, 9048!. 

I'yrrosia lanccolata (L.) Farwell 

Small epiphytic fern with a long-creeping rhizome, 

simple, thick, narrowly lanceolate to oblanceolate 

fronds, and closely arranged sori covering the apical 

half of the lower lamina surface. Common in low- 

land to montane forest and in plantations and 

villages, reported from near sea level to 540 m ele- 

vation. Indigenous, ranging from India to Samoa. 

Reinecke 15 1; Rechinger 1265; Setchell 4, 538; 

Garber 803; Wilder s.n.; Meebold 165 15, 26515; 

Yuncker 9306; Wisner 2; Lamoureux 3000; 

Whistler 1458,2821, 8535!. 

SCHIZAEACEAE 

Scliizaca dichotoma (L.) Smith 

'I'errestrial fern with a creeping, underground rhi- 

zome, clumped fronds dichotomously branched 2-8 

times ultin~atcly into narrow, spreading lobes, and 

terminal sori with sporangia in two rows. Uncom- 

mon in lowland to montane forest reported from 100 

to 580 m elevation. Indigenous, ranging from 

Madagascar to the Marquesas. Reinecke 57b; 

Setchell 271; Garber 939; Christophersen 3536; 

Whistler 8464, 8583!. 

THELYPTERIDACEAE 

Christella dentata (Forssk.) Brownsey & Jenny 


rhizome, pubescent stipes, lamina pinnate, pinnae 

lobed about halfway to the costa, the lowest 2 or 3 

pairs gradually reduced and generally reflexed, the 

basal pair of veins united to form an excurrent vein 

to the sinus, pubescent surfaces, yellow glands on 

the lower surface, and a cordate-renifom indusium. 

Uncommon in sunny disturbed places, reported from 


widespread throughout the Old World tropics and 

subtropics. Whistler 3 845, 8636!. 

Christella harveyi (Mett.) Holttum 

Large terrestrial fern with a long-creeping rhizome, 

pubescent stipes, pinnate lamina, pinnae lobed to 

near the costa, up to 5 pairs of reduced pinnae, basal 

veins free, pubescent surfaces, yellow glands present 

on the lower surface, and a reniform, glabrous 

indusium. Common in disturbed places and lowland 


tion. Indigenous, widespread from the Bismarck 

Archipelago to Samoa. Setchell 25; Stearns sn; 

McMullin 2; Meebold 2658 1; Whistler 8442, 853 1 !. 

*Christella pacifica Holttum 

Medium-sized terrestrial fern with a long-creeping 

rhizome, pinnate fronds, pinnae lobed almost 

halfway to the costa, lowest pairs gradually reduced, 

basal pair of veins united, lower surface pubescent, 

sori medial, and indusium short-hairy. Uncommon 

in disturbed areas, reported at ca. 200 m. Indige- 

nous, also found in Fiji and New Caledonia. 

Yuncker 9346 (n.s.). 

*Christella parasitica (L.) Leveille 

Medium-sized terrestrial fern with a creeping rhi- 

zome, pinnate fronds, pinnae cut to about 213 of the 

way to the costa, lowest pairs hardly reduced, 

surfaces pubescent, orange glands present, the 

lowest veins united, sori in a row along the costa, 

and indusium hairy. Occasional in disturbed places, 


Indigenous, ranging from east Africa to Hawai'i. 

Whistler 8396. 

*Christella subjuncta (Baker) Holttum 


rhizome, fronds pinnate, cut to near the costa into 30 

pairs of pinnae, the lowest 2 or 3 pairs reduced, 

lower pinnae surface pubescent, veins free or one 

pair united, sori submarginal, and indusia small and

pubescent. [Jncommon in disturbed places, reported 

from ccr. 50 m elevation. Endemic. Powell 75; 

McMullin 2a; Setchell lob. 

Macrothelypteris polypodioides (Hooker) Holttum 

Large terrestrial fern with a short rhizome, bipinnate 

totripinnatifid fronds, the largest pinnules 7 x 2 cm, 

pale scales with marginal hairs on the rachis and 

lower surfaces of pinnules, small sori, and small 

indusia bearing a few capitate hairs. Occasional in 

open sunny places such as roadcuts, reported from 


ranging from Thailand to Tahiti. Whistler 1456, 

8426,8448, 8614,8976!. 

Macrothelypteris torresiana (Gaud.) Ching 

Large terrestrial fern with a stout, short-creeping 

rhizome, large bipinnate-tripinnatifid fronds, largest 

pinnae up to 5 cm long, hairs but no scales on 

rachises and lower surface of pinnules, small sori, 

and a small indusium bearing a few capitate hairs. 

Occasional in sunny disturbed places, reported from 


ranging from the Madagascar to Hawai'i. Reinecke 

177?; Wilder s.n.; Meebold 16864, 26587; Yuncker 

9394; Whistler 8432, 8732!, 9083. 

Plesioneuron attenuatum (Bracken.) Holttum 

Large terrestrial fern with a short erect rhizome, pin- 

nate lamina, 25-30 pairs of pinnae, the basal basio- 

scopic lobe of pinna much reduced, apex attenuate, 

sori near the costules, and the indusium dark and 

pubescent. Occasional in sunny, disturbed places, 


ranging from the Bismarck Archipelago to Tahiti. 

Wilder s.n.? (n.s.); Whistler 8523!, 8546!, 8695!, 

8900!. 

Pneurnatopteris bryanii (Christen.) Holttum 

Large terrestrial fern with an erect rhizome, pinnate 

laminalobed about 2/3 of the way to the costa, 

lowest 2-4 pairs abruptly reduced, 1-1.5 pairs of 

veins united, sori near the costules, and indusia 

small and glabrous. Uncommon in disturbed places, 


Endemic. Whistler 3626, 8593!. 

Pneurnatopteris glandulifera (Bracken.) Holttum 

Large terrestrial fern with a decumbent rhizome, 

many reduced pinnae each consisting of an crophore 

and a green rim at its base, costules with small 

acrophores at their bases, lower 1 or more veins 

united, and no indusia present. Occasional in sunny 

places, reported from near sea level to 300 m 

elevation. Indigenous, ranging from the Solomon 

Islands to Rarotonga. USEE s.n.; Setchell 283; 


*Pneurnatopteris rodigasiana (T. Moore) Holttum 


lamina, pinnae lobed about halfway to costa, the 

lowest 9 pairs reduced, the basal pair of veins uni- 

ted, sori inframedial, and the indusium pubescent. 

Uncommon in disturbed places, reported from 50 to 


Solomon Islands to Niue. Setchell 5 19 (n.~.). 

'Pneumatopteris transversaria (Bracken.) 

Holttum 


lamina, pinna lobed about halfway to the costa, 

lowest six or so pairs reduced, the basal pair of veins 

united, sori near the costules, and the indusium 

small and glabrous. Uncommon in disturbed places, 


also found in Fiji. USEE s.n.; McMullin 10. 

"Sphaerostephanos reineckei (Christen.) Holttum 


up to 15 pairs of reduced pinnae consisting of an 

acrophore and a tiny lamina, the basal pair of veins 

united, lower lamina surface pubescent and bearing 

glands, and no indusia. Uncommon in montane 


Endemic. Sledge 1820. 

Sphaerostephanos unitus (L.) Holttum 


rhizome, pinnate, coriaceous lamina, pinna divided 

about halfway to the costa, the lowest 6 or more 

pairs reduced, small yellow glands present on the 

lower surface, 1.5 pairs of veins united, sori supramedial, 

and indusium pubescent. Uncommon to 

locally common in disturbed places, reported at 120 

m elevation. Indigenous, ranging from east Africa 

to Samoa. Setchell 10, 226; Kuruc 142; Whistler 

803 1,8978!.

Antrophyum alatum Bracken. 

VITTARIACEAE FERN ALLIES 

Epiphytic fern with a short-creeping rhizome, tufted 

simple spathulate fronds, and sori arranged in 

longitudinal rows on veins of the lower surface of 

the lamina. Occasional in lowland to montane 


nous, ranging from New Caledonia to Tahiti. Gar- 

ber 839; Mitchell 508; Meebold 16636; Yuncker 

9400; Kuruc 146; Whistler 8576!, 9127. 

Antrophyum plantagineum (Cav.) Kaulfuss 

Small epiphytic fern with a short rhizome, stipes 3- 

10 cm long, lamina strap-shaped, and linear sori on 

the lower surface following along the veins. Occa- 

sional to common in lowland forest, reported from 


ranging from Malaysia to the Marquesas. Veitch 

s.n.; Setchell 46, 151; McMullin 34; Garber 840, 

903; Yuncker 9316; Wisner 6; Long 1958a, 1997a, 

3054; Whistler 8549!, 91 19,9353!. 

Vaginularia angustissima (Bracken.) Mett. 

Epiphytic, clump-forming fern with a creeping 

rhizome, simple, linear fronds ca. 2 mm wide, and 

sori in a groove opening towards the costa on the 

lower surface of the lamina. Occasional in lowland 

to montane forest, reported from 30 to 550 m ele- 

vation. Indigenous, also found in Fiji and probably 

Vanuatu. Setchell 43a, 251; Garber 904; Whistler 

8582!. 

Vittaria elongata Sw. 

Epiphytic fern with a creeping rhizome, simple, 

linear fronds 10-40 cm long and 3-7 mm wide, and 

sori in a marginal groove. Occasional in lowland to 

cloud forest, reported from near sea level to 1400 m 

elevation. Indigenous, ranging from tropical Asia to 

the Marquesas. Reinecke 40, s.n.; McMullin 47; 

Long 1957, 3095; Whistler 2822,8542!. 

Vittaria scolopendrina (Bory) Thwaites 

Epiphytic fern with a creeping rhizome, simple, 

sessile, pendant, linear fronds 40-90 cm long and up 

to ca. 2.5 cm, and sori in a marginal groove. Un- 

common in lowland to montane forest, reported 


from Madagascar to Samoa. Sledge 182 1 !; Whistler 

9200. 

LYCOPODIACEAE 

Lycopodium carinatum Desv. 

Epiphytic herb with dimorphic leaves densely 

arranged around the stems, with the sterile leaves 

gradually passing into the smaller ovate fertile 

leaves. Occasional in lowland to cloud forest, 


Indigenous, ranging from India to Samoa. Garber 

924; Christophersen 1824!; Lamoureux 30 17; Long 

1995, 1996e; Whistler 3628,8532!. 

Lycopodium cernuum L. 

Terrestrial herb with much-branched stems, linear- 

subulate leaves 2-3 mm long, and numerous strobili 

solitary on the tips of the branchlets. Uncommon in 

sunny places, reported from 70 to 1650 m elevation. 

Indigenous, found throughout the temperate and tro- 

pical regions. Reinecke 1 lb; Setchell 109,273,403; 

Bryan 976!; Christophersen 1033, 3548; Mitchell 

5 12; Long 2609; Whistler 2934, 8406, 87O9!. 

Lycopodium phlegmaria L. 

Epiphytic herb with dimorphic leaves abruptly 

differing from each other, the sterile leaves 2-4 mm 

wide, the tiny fertile leaves arranged in long, 

slender, forked spikes. Uncommon in lowland to 


elevation. Indigenous, found throughout the Old 

World tropics. Veitch s.n.; Reinecke 10a; Setchell 

60, 182, 269, 396; Bryan 89; Christophersen 1125!, 

1823!, 1830!; Yuncker 9318; Wisner 144; Long 

1958, 1995b, 3075; Whistler 3629, 8500, 8545!, 

89O4!. 

Lycopodium squarrosum Forst. f 

Epiphytic herb with stems branched 2-4 times, 

sterile leaves narrowly lanceolate, 1-2 cm long, and 

passing imperceptibly into the upper, fertile leaves. 

Uncommon in lowland to montane forest, reported 


nous, ranging from tropical Asia to Tahiti. Setchell 

270; Christophersen 12 18, 3525; Lamoureux 3002; 

Whistler 8499, 8544!, 8729!.

*Psiloturn complanatum Sw 

PSILOTACEAE DICOTYLEDONAE 

Leafless pendulous epiphytic herb with a short- 

creeping rhizome, flattened branches 2-3.5 mm 

wide, and conspicuous yellow sporangia. Uncom- 

111011 ~n lowlond Ibrcst, rcportcd from near sea level 

to 360 m elevation. Indigenous, widespread in the 

tropics. Lamoureux 301 5; Yuncker 9415; Whistler 

2789. 

Selaginella laxa Spring 

SELAGINACEAE 

Tiny terrestrial herb with prostrate stems, tiny 

dimorphic leaves, and dimorphic sporophylls. 

Uncomn~on on rocks in sunny places, reported from 


ranging from Fiji to Tahiti. USEE s.n.; Veitch s.n.; 

Reinecke s.n.; Meebold 21358; Setchell 107, 152; 

Y uncker 9334; Whistler 8402,845 1, 8696!, 8905!. 

Selaginella whitmeei Baker 

Erect terrestrial herb with ovate acuminate leaves on 

the main stem, ovate to lanceolate lateral leaves, and 

sporcs borne in tetragonous strobili. Occasional in 

lowland to cloud forest and in rocky stream beds, 

reported from 150 to 1650 m elevation. A number 

of virtually indistinguishable "species" have been 

named from Samoa, and further taxonomic work is 

needed to show if they are actually distinct species. 

Endemic. Reinecke 62, 62a, 62b, 62f, 62v?; Steams 

s.n.; Setchell 49, 71, 183, 223, 277; Garber 845, 

923; Bryan 77; Meebold 17123; Yuncker 9325; 

Christophersen 1215!, 1812!, 1813!, 1814!, 1816!, 

l8l7!, 3502, 3524, 3544; Wisner 143; Long 1996f, 

3053; Lamoureux 3014; Sledge 1830; Whistler 

8407, 8522!, 8548!, 8940!. 

NOTE: Several other species have been attributed to 

Tutuila, but these claims are not supported by 

voucher specimens or are otherwise likely to be 

mistakes. This includes the following species: (1) 

Lusfreopsis davalloides (Bracken.) Tindale, which 

was attributed to the island by Christ; (2) Aspleniurn 

lobulaturn Mett., based on Reinecke 37a, which may 

be just a form of another species, and is, in any case, 

otherwise only reported from 1400 to 1700 m 

elevation; and (3) Dennstaedtiaflaccida (Forst. f.) 

Bernhardi, attributed to the island by Christensen. 

ACANTHACEAE 

"Asystasia gangetica (L.) T. Anders. 

[Chinese violet] 

Weak-stemmed herb with opposite leaves, white to 

purple campanulate flowers in terminal racemes, and 

a clavate capsule. Uncommon to rare as an adven- 

tive or an escape in disturbed areas, reported only 

from the lowlands. A modern introduction, native to 

the Old World tropics. Yuncker 9393; Whistler 307. 

Blechum brownei Juss. 

Herb with opposite leaves and small white, campa- 

nulate flowers borne among ovate bracts in terminal 

spikes. Uncommon in disturbed areas, reported 

from near sea level to 610 m elevation. A modern 

introduction, native to tropical America. Whistler 

9058. 

*Hemigraphis alternata (Bum. f.) T. Anders. 

Prostrate herb with opposite purple leaves, purple 

bracts, and white campanulate flowers. Locally 

common as a weed in lawns, reported only from the 

lowlands. A modern introduction, native to tropical 

Asia. Whistler 8933. 

Justicia procumbens L. 

Small herb with opposite leaves and tiny lavender, 

bilabiate flowers in dense, ovoid, bract-bearing, 

terminal spikes. Common in disturbed areas, report- 

ed only from the lowlands. A modem introduction, 

native to tropical Asia. Whistler 320, 2757, 8958!, 

9176. 

Ruellia prostrata Poir. 

vaouli 

Herb with opposite leaves, solitary, axillary, laven- 

der, campanulate flowers, and a club- shaped cap- 

sule. Common in disturbed areas, reported only 


the Old World tropics. Whistler 9082, 9lO5!. 

*Ruellia tuberosa L. 

Tough-stemmed herb with opposite leaves, large, 

lavender, campanulate flowers solitary in the leaf 

axils, and club-shaped capsules. Uncommon as a

weed in villages and along roadsides, reported only 


introduction, native to the Old World tropics. 

Whistler 2760, 8440. 

AIZOACEAE 

"Scsuvium portulacastrum (L.) L. 

Prostrate succulent herb with opposite leaves and 

white to lavender flowers bearing numerous sta- 

mens. Uncommon on coastal rocks, reported only 

near sea level. Indigenous, widespread in the 

Pacific. Whistler 8428. 

"Achyranthes aspera L. 

tamatama 

AMARANTHACEAE 

Subshrub with opposite leaves, tiny purple flowers, 

and sharp, grass-like fruits in terminal spikes. Occa- 

sional in coastal areas and sometimes weedy, report- 

ed from near sea level to 125 m elevation. Indige- 

nous or a Polynesian introduction, widespread in the 

tropics. Setchell 146; Wilder 9; Garber 962; Wisner 

62: Whistler 8427. 

Altcrnanthera sessilis (L.) R. Br. ex DC. 

Creeping herb with opposite leaves and inconspi- 

cuous flowers in white, sessile, axillary, globose 

clusters. Occasional to common in disturbed places, 

particularly in wetland crops such as taro, reported 


introduction, possibly native to southern China. 

Setchell 303; Wilder 26; Garber 862; Lamoureux 

3085; Whistler 3954, 8601!. 

"Amaranthus viridis L. 

Red-stemmed herb with alternate leaves notched at 

the tip, and terminal and axillary spikelike panicles 

of tiny inconspicuous, green flowers. Uncommon in 

croplands and waste places, reported only from the 

lowlands. A Polynesian introduction, now pantropic 

in distribution. Setchell 319; Garber 968. 

"Cyathula prostrata (L.) BI. 

Low, weak-stemmed herb with opposite leaves, tiny 

grecn, inconspicuous flowers in narrow, long- 

stalked terminal spikes, and bur-like fruits. Uncom- 

mon in disturbed places including secondary forest, 

reported from near sea level to 1100 m elevation. A 

Polynesian introduction, native to the Old World 

tropics. Setchell215, 326. 

ANACARDIACEAE 

Buchanania merrillii Christoph. 

Large tree with simple, alternate, long-petioled 

oblanceolate leaves, white flowers in axillary 

panicles, and hard, purple, lens-shaped fruits. 


from 250 to 450 m elevation. Endemic. Setchell 

430, 437; Garber 921; Christophersen 1010; tong 

1971 ; Spence 474!; Whistler 2691 !, 2962, 85 16!. 

Rhus taitensis Guillemin 

tavai 

Large tree with milky sap, alternate pinnately 

compound leaves, tiny white flowers, and small, 

purple, fleshy fruits. Common to abundant in 

lowland and especially secondary forest, reported 


from Futuna to Tahiti. Setchell 65; Bayliss s.n.; 

Yuncker 9332; Wisner 7; Spence 476!; Whistler 

2734,3642. 

ANNONACEAE 

Cananga odorata (Lam.) Hook. f. & Thoms. 

[ilangilang] moso'oi 

Straight, medium-sized tree with alternate leaves all 

in one plane, large, fragrant, yellow apocarpus 

flowers, and black subglobose fruits. Occasional to 

common in disturbed forest, reported from 15 to 600 

m elevation. A Polynesian introduction, or possibly 

indigenous, widespread westward to South Asia. 

Setchell 179; Diefenderfer 3; Whistler 3 1 19, 9108!. 

APIACEAE 

Centella asiatica (L.) Urb. 

[Asiatic pennywort] tono 

Low herb with creeping stems rooting at the nodes, 

alternate, kidney-shaped leaves, and tiny green, 

inconspicuous, axillary flowers. Occasional to 

common in sunny disturbed places, reported from 

near sea level to 700 m elevation. Probably a 

Polynesian introduction, native to tropical Asia. 

Reinecke 524; Garber 965; Whistler 1455,908 1 !.

APOCYNACEAE 

Alstonia pacifica (Seem.) A. C. Smith 

Small tree with glossy opposite leaves, milky sap, 

small white flowers, and long, curved, cylindrical 

capsules. Common in open forest on ridges in the 

tnoul~ti~ins, rcported from 150 to 1300 rn clcvation. 

Indigenous to Samoa, ranging westward to the 

Solomon Islands. Setchell 38 1, 387; Christophersen 

1042, 1061, 1 IB!, 1257, 1265, 3540; Long 1970, 

1973; Whistler 2707, 2715, 3301, 8519!, 8676!, 

9066. 

Alyxia bracteolosa Rich 

lau maile 

Vine with milky sap, shiny opposite leaves, tiny 

white flowers, and bluish-black globose fruits. 


10 to 1300 m elevation. Indigenous, also found in 

Fiji, Home Island, Wallis, and Tonga. Setchell 21 1, 

279; Swezey & Wilder 35; Garber 856; Christopher- 

sen 3539; Diefenderfer 9; Mitchell 521; Yuncker 

9349; Wisner 149, 150; Long 3109; Spence 467!; 

Whistler 2771 !, 2907A, 2982,3109, 8671 !, 9097!. 

Alyxia stellata (Forst.) Roemer & Schultes 

nau 

Scandent shrub with small, shiny opposite leaves, 

milky sap, tiny white flowers, and small subglobose 

to ellipsoid fruits. Common in open forest and on 

ridges, reported from 20 to 1700 m elevation. Indi- 

genous, ranging from New Caledonia to the Mar- 

quesas. Setchell294, 542; Bryan 84; Christophersen 

3551; Mitchell 521; Wisner 126; Lamoureux 3020; 

Whistler 1277,2889, 8672!. 

Cerbera manghas L. 

leva 

Medium-sized tree with glossy whorled leaves, 

milky sap, showy white flowers with a red throat, 

and a large, red, ellipsoid fruit. Occasional in littoral 

forest and sometimes cultivated and persisting in 

secondary forest, reported from near sea level to 650 

nl elevation. Indigenous, ranging from Malaysia to 

the Marquesas. Setchell 92; Diefenderfer 20; 

Yuncker 9427; Lamoureux 3011; Whistler 1420, 

2835, 8663!. 

"Ervatamia obtusiuscula Markgraf 

pulu 

Small tree with milky sap, opposite leaves, white 

flowers with contorted corolla lobes, and orange, 

2-lobed fruits. Uncommon in lowland forest, 


ranging from Vanuatu to Makatea and Tahiti. 

Reinecke 539, 553, 569. 

ARALIACEAE 

Meryta macrophylla (Rich) Seem. 

ma'ulu'ulu; fagufagu 

Small dioecious tree with large, simple, alternate 

leaves, thick, compact male inflorescences, greenish 

female flowers, and large, green, fused fruits. Occa- 

sional in coastal to montane forest, reported from 15 

to 800 m elevation. Indigenous to Samoa, also 

found in Tonga. Garber 929; Christophersen 1006, 

1 l3O!, 1253, 1254; Long 3094; Spence 468!; 

Whistler 27 18. 

Polyscias samoensis (A. Gray) Harms 

tagitagi; afia 

Small tree with alternate, pinnately compound 

leaves, flowers in compound umbels, and flattened, 

round, striate fruits. Common in coastal to montane 

forest, reported from near sea level to 400 m elevation. 

Indigenous, also found on Niuafo'ou. Graeffe 

251; Setchell 255, 374; Wilder 44, 87; Long 3101; 

Rainey 2393; Whistler 1436, 2722, 2799, 8989!, 

9146. 

ARISTOLOCHIACEAE 

Aristolochia cortinata Reinecke 

Woody climber with alternate, ovate leaves, curvedtubular, 

purple flowers, and an ovoid capsule. Uncommon 

in lowland to montane forest, reported 

from 20 to 600 m elevation. Endemic. Garber 937; 

Mitchell 519; Whistler 2869, 2899, 8580!, 8902!. 

ASCLEPIADACEAE 

*Axlepias curassavica L. 

[milkweed] 

Erect, scarcely branching shrub with milky sap, 

opposite leaves, orange and red flowers in umbels, 

and a follicle bearing comose seeds. Uncommon as 

a weed, particularly in pastures, reported from 70 to

650 111 clevi~tion. A modern introduction, native to 

tropical America. Setchell 184; Garber 890. 

Hoya australis R. Br. in Traill 

Cue selela; suni; olive vao 

Vinc with milky sap, fleshy opposite leaves, white 

flowers usually marked with red at the corolla base, 

and a follicle bearing comose seeds. Occasional 

from littoral to montane forest, reported from near 


from Australia to Samoa. Lister s.n.; Setchell 515; 

Wisner 1 18, 124; McKee 2956; Long 3090; 

Whistlcr 2833. 

Hoya pottsii Traill 

fue selela 

Viric with opposite leaves 3-5 palmately veined 

from the ba~e, white waxy flowers in umbels, and a 

cylindrical follicle bearing comose seeds. Occasional 

in the coastal to montane forest, reported 


ranging from Hainan Island to Samoa. Christophersen 

1007; Meebold 16769; Yuncker 9379; Long 

1985; Whistler 1460, 8505, 9098!. 

Hoya vitiensis Turrill 

fue selela 

Vine with milky sap, opposite, thick, mostly lanceolate 

leaves, white or maroon flowers 15-28 mm 

across, and a cylindrical follicle bearing comose 

seeds. Occasional in lowland to montane forest, 


also found in Fiji. Christophersen 101 1, 3562; Long 

3045; Whistler 2765!, 2793, 31 10,3111, 8717. 

"1-loya sp. nova 

file selela 

Vine with milky sap, opposite, thick, mostly lanceolate 

leaves, white, waxy flowers, and a cylindrical 

follicle bearing comose seeds. Occasional in 


level to 600 m elevation. Apparently endemic to 

American Samoa (Tutuila, Ofu, and Ta'u), but 

further work needs to be done on this genus in 

Samoa. Whistler 2705. 

ASTERACEAE 

Acmella uliginosa (Sw.) Cass. 

Erect herb with toothed, opposite leaves and yellow 

disc and ray florets in small, ovoid heads. Uncommon 

in disturbed places, especially in wet areas, 


modern introduction, native to tropical America. 

Whistler 3 18, 8049, 85 13!. 

Ageratum conyzoides L. 

Erect herb with coarse, opposite, fragrant leaves and 

lavender disc florets in heads arranged in terminal 

panicles. Common as a weed in sunny, disturbed 

places and croplands, reported from near sea level to 

1650 m elevation. A modem introduction, native to 

tropical America. Eames T5; Lamoureux 3058; 


Bidens alba (L.) DC. 

[beggar's-tick] 

Herb with pinnately lobed or compound, opposite 

leaves, yellow disc florets, white, spreading ray 

florets, and small, cylindrical, barbed achenes. 

Abundant in disturbed places, apparently displacing 

the following species, reported only from the lowlands. 

A modern introduction, native to tropical 

America. Whistler 2970, 8570!. 

Bidens pilosa L. 

[beggar's-tick] 

Herb with pinnately lobed or compound, opposite 

leaves, yellow disc florets in heads, and small, 

cylindrical, barbed achenes. Now uncommon (in 

American Samoa) in dry, sunny disturbed places, 



USEE sen.? (n.s.); Eames T29, Whistler 8926!. 

"Conyza bonariensis (L.) Cronquist 

[hairy fleabane] 

Erect, scarcely branching herb with hairy stems, 

alternate, gray-green leaves, branching panicles of 

small heads, and small, plumed achenes. Rare as a 

weed in disturbed places, perhaps now extirpated 

from Samoa, reported at ca. 10 m elevation. A 

modern introduction, native to South America. 

Whistler 3605.

Crassoccpllalum crepidioides (Benth.) S. Moore 

Tall herb with toothed or lobed, alternate leaves, 

drooping heads of disc florets red-brown at the tips, 

and plumed achenes. Common to abundant in 

disturbed places, especially in taro fields, reported 


introduction, native to tropical Africa. Lamoureux 

3060; Whistler 8619!. 

"Eclipta prostrata (L.) L. 

Small herb with opposite leaves and small, aster- 

like heads of white ray and disc florets. Uncommon 

in disturbed places, mostly in wet soil, reported only 

from the lowlands. A modern introduction, pan- 

tropic in distribution. Lamoureux 3072; Whistler 

328. 

Eleutheranthera ruderalis (Sw.) Schultz-Bip. 

Coarse herb with opposite leaves and drooping 

stalked heads or several disc florets. Uncommon in 

disturbed places. A modern introduction, native to 

tropical America. Trail 280. 

Emilia sonchifolia (L.) DC. 

fua lele 

Erect herb with alternate, sessile, variously shaped 

leaves with clasping auricles, a panicle of several 

heads of lavender disc florets, and plumed achenes. 

Common in sunny disturbed places, reported from 

near sea level to 450 m elevation. A modern intro- 

duction, native to the Old World tropics. Setchell 

1 2; Eames T9; Garber 8 1 1 ; Lamoureux 306 1, 3062? 

(n.~.), 3063; Sachet 37; Whistler 304, 322, 2935, 

8950!. 

Ercchtites valerianifolia (Wolf) DC. 

fua lele 

Tall, erect herb with alternate, deeply pinnately 

lobed leaves, pink disc florets in heads arranged in 

terminal panicles, and plumed achenes. Occasional 

to common in open, disturbed places, reported from 

near sea level to 1750 m elevation (but uncommon 

at low elevations). A modern introduction, native to 

tropical America. Setchell l I; Christophersen 957; 


Mikania micrantha H. B. K. 

[mile-a-minute vine] fue saina 

Herbaceous vine with opposite leaves, white disc 

florets in heads arranged in panicles, and plumed 

achenes. Abundant in disturbed places, reported 


introduction, native to tropical America. Long 

2606a; Whistler 8632!, 9133. 

*Pseudelephantopus spicatus (B. Juss. ex Aubl.) 

Baker 

vao malini 

Wiry, deep-rooted herb with alternate leaves and 

purple to white disk florets in heads arranged in a 

narrow, spike-like inflorescence. Occasional in 

lawns and disturbed places, reported only from the 

lowlands. A modem introduction, native to tropical 

America. Whistler 9072. 

"Struchium sparganophorum (L.) Kuntze 

Erect herb with alternate leaves and inconspicuous, 

white disc florets in sessile, axillary heads. Occa- 

sional on streamsides and other wet places, reported 



3229. 

Synedrella nodiflora (L.) Gaertn. 

tae'oti 

Coarse herb with opposite leaves and yellow disc 

florets arranged in sessile, axillary heads. Common 

in disturbed places, reported from near sea level to 

550 m elevation. A modem introduction, native to 

tropical America. Setchell 19, 540; Eames Tll; 

Wilder 29; Garber 814; Lamoureux 3067, 3074; 

Whistler 8626!, 8957!. 

"Tridax procumbens L. 

[coat buttons] 

Ascending herb with opposite, toothed leaves and 

solitary, long-stalked heads of white ray and yellow 

disc florets. Occasional in disturbed places, reported 

only from the lowlands. A modern introduction, 

native to tropical America. Whistler 9346.

Vernonia cinerea (L.) Less. 

[ironweed] 

Small herb with alternate leaves, purple disc florets 

in heads arranged in loose terminal cymes, and 

plumed achenes. Occasional to common in disturhcd 

placcj, rcported from near sea level to 360 m 

clcvation. A modcrn introduction, native to tropical 

America. Eames T3; Lamoureux 3057; Whistler 

2758. 9012!. 

*Wedelia trilobata (L.) Hitchc. 

Low-growing herb with opposite leaves having 

toothed margins, showy yellow composite flowers 

on long peduncles, and yellow disk and ray florets. 

Uncommon as an escape from cultivation and 

sometimes weedy, reported only from the lowlands. 

A modern introduction, native to the New World 

Tropics. Whistler 9059. 

Wollastonia biflora (L.) DC. 

[beach sunflower] ateate 

Subshrub with opposite leaves, panicles of composite 

flowers, and yellow disc and ray florets. Common 

to abundant in sunny coastal areas, sometimes 

extending inland in coconut plantations, reported 


ranging from tropical Asia to the Austral Islands. 

Setchell 17, 125; Garber 812; Bryan 975; Yuncker 

9368; Mitchell 529; Wisner s.n.; Lamoureux 3091; 

Whistler 1434, 29 15, 8562!. 

Youngia japonica (L.) DC. 

[Oriental hawksbeard] 

Rosette-forming herb with lobed, basal leaves, 

yellow ray disc florets in heads in long, thin, 

branching panicles, and plumed achenes. Uncommon 

in disturbed places, reported from near sea 

level to ca. 300 m elevation. A modern introduction, 

native from India to Japan. Whistler 5733, 

8419, 8647!. 

BARRINGTONIACEAE 

Barringtonia asiatica (L.) Kurz 

[fish-poison tree] futu 

Large tree with large, alternate, obovate leaves, 

large flowers with numerous long, white and pink 

stamens, and large top-shaped fruits. Common to 

abundant in littoral, coastal, and ridge forests, 

reported from near sea level to ca. 200 m elevation. 

Indigenous, ranging from Madagascar to the 

Marquesas. Setchell 105, 144, 160, 344; Long 

2597; Whistler 2913, 8916!. 

Rarringtonia samoensis A. Gray 

thlaga 

Medium-sized tree with large alternate leaves, 

flowers in hanging racemes, numerous showy red 

stamens, and top-shaped fruits. Occasional in 

lowland to montane forest, reported from 5 to 600 m 

elevation. Indigenous, ranging westward to Indonesia. 

USEE s.n.; Home 3, 4; Setchell45, 159, 202, 

250; Bryan 87; Garber 851, 857, 860, 883, 944; 

Christophersen 1036; Yuncker 9362; Wisner 122, 

125; Spence 482!; Whistler 1276,2863, 3287, 3654, 

91 10. 

BORAGINACEAE 

Corda aspera Forst. f. 

tou 

Small tree with alternate leaves, small white sympetalous 

flowers, and white fleshy fruits. Rare in 

lowland forest, reported from 30 to 275 m elevation. 

Indigenous or a Polynesian introduction, ranging 

from the Philippines to Samoa. Whistler 3 112, 

3113,9347!. 

*Cordia subcordata Lam. 

tauanave 

Medium-sized tree with alternate leaves, showy 

orange flowers and green, subglobose fruit. Uncommon 

in littoral forest, reported only near sea level. 

Indigenous, ranging from tropical Asia to eastern 

Polynesia. Setchell 369; Christophersen 1175; 

Yuncker 9381; Whistler 2808. 

*Heliotropium procumbens Mill. 

Low herb with narrow, pubescent, alternate leaves 

and tiny white sympetalous flowers in scorpeoid 

cymes. Uncommon in disturbed places, reported 


native to tropical America. Whistler 2968. 

"Tournefortia argentea L. f. 

[beach heliotrope] tausuni 

Small tree with silvery-pubescent, alternate leaves, 

small white flowers in compound, scorpeoid cymes,

and small, round, green fruits. Occasional on the 

seaward edge of littoral forest, reported only near 

sea level. Indigenous, ranging from Mauritius to 

southeastern Polynesia. Setchell 128, 23 1, 557; 

Cornstock 11; Wisner 89; Long 1939; Whistler 

1430. 

BRASSICACEAE 

*Rorippa sarmentosa (DC.) Macbr. 

[Polynesian cress] a'atasi 

Small herb with pinnately lobed leaves, tiny white 

flowers in terminal racemes, and small linear pods. 

Occasional as a weed, sometimes occurring in sunny 

undisturbed habitats, reported from near sea level to 

1700 m elevation. Indigenous or perhaps a Polynesian 

introduction, ranging from New Caledonia to 

Hawai'i. Setchell 325; Whistler 1466, 8433. 

BURSERACEAE 

Canarium harveyi Seem. 

mafoa? 

Medium-sized tree with alternate, pinnately compound 

leaves, 5-9 ovate to oblong leaflets, and a 

large purple fruit. Occasional in coastal to montane 


A modern introduction or possibly indigenous, 

also found in Fiji, Tonga, and Niue. Swezey 

& Wilder 256; Whistler 2693,2873, 8508!, 91 13. 

Canarium vitiense A. Gray 

ma'ali 

L,arge tree with fragrant resin, alternate, pinnately 

compound leaves bearing 7-13 elliptic leaflets, and 

ellipsoid to ovoid fruits. Occasional in lowland to 

montane forest, reported from 10 to 700 m elevation. 

Indigenous, also found in Fiji and Tonga. 

Setchell420, 574; Whistler 1281,2939, 3123. 

Caruga floribunda Decne. 

vivao 

Large tree with pubescent, alternate, pinnately 

compound leaves with leaflets that turn red individually 

when aging, small white flowers, and blueblack 

globose fruits. Occasional in coastal to 

lowland forest, reported from 10 to 250 m elevation. 

Indigenous, ranging westward to the Philippines and 

Indonesia. Setche11432; Whistler 31 16. 

CARICACEAE 

Carica papaya L. 

[papaya] esi 

Scarcely branching, dioecious tree with large, 

alternate, palmately lobed leaves, cream- colored, 

salverform male flowers in pendulous racemes, 

solitary, axillary, cream- colored flowers, and a 

large edible orange fruit. Common in disturbed 

places and cultivated in villages, reported only from 

the lowlands. A modern introduction, native to 

tropical America. USEE s.n.? (n.s.); Setchell 70; 

Garber 972; Whistler 8928!, 9134. 

CASSYTHACEAE 

Cassytha filiforrnis L. 

fetai 

Leafless, green to orange, string-like parasitic vine 

with small white flowers and globose fruits. 

Uncommon on Tutuila in littoral areas, reported 


pantropic in distribution. Whistler 8683 ! . 

CELASTRACEAE 

Maytenus vitiensis (A. Gray) Ding Hou 

Shrub or small tree with alternate leaves, toothed 

leaf margins, small white flowers, and 3-angled capsules. 

Occasional in open forest on ridges and lava 

flows, reported from 25 to 600 m elevation. Indigenous, 

ranging from Fiji to the Gambier Islands. 

Wilder 36; Garber 899 (n.~.); Christophersen 1015; 

Wisner 158; Whistler 2706, 8455, 8685!. 

Atuna racemosa Raf. 

ifiifi 

CHRYSOBALANACEAE 

Medium-sized tree with alternate simple leaves, 

white flowers, and a large, flattened-ovoid, hard- 

shelled fruit. Now rare in lowland to montane for- 

est, sometimes forming small groves, reported from 

near sea level to 450 m elevation. A Polynesian 

introduction, ranging from tropical Asia to Samoa. 

Setchell325; Garber 892; Spence 480!. 

*Parinari insularum A. Gray 

sea 

Medium-sized tree with alternate leaves, small white 

flowers, and fragrant, mottled brown, ovoid fruits.

Uncomnion in secondary forest, probably a relict of 

cultivation, reported only from the low-lands, per- 

haps now extirpated from Samoa. A Polynesian 

it~troduction from Fiji, also found in Tonga and 

Wallis. Setchell 560. 

Calophyllum inophyllum L. 

fetau 

Large tree with fissured bark, glossy, finely veined, 

opposite leaves, showy yellow and white flowers, 

and a subglobose green fruit. Common and some- 

times dominant in littoral forest, reported from near 

sea level to ca. 100 m elevation. Indigenous, 

ranging from tropical Africa to eastern Polynesia. 

Setchell 99, 99a; Wisner 11 1; Whistler 2832,2921. 

Calophyllum neo-ebudicum Guillaumin 

tamanu 

Large tree with fissured bark, finely veined, 

opposite leaves, showy white and yellow flowers, 

and a subglobose purplish drupe. Occasional to 


rrom 25 to 650 m elevation. Indigenous, ranging 

from New Britain to Niue. Setchell 221, 334; 

Spence 496!; Whistler 2937, 8677!. 

Garcinia myrtifolia A. C. Smith 

Medium-sized tree with opposite, coriaceous leaves, 

white flowers bearing numerous yellow stamens, 

and an ellipsoid fruit. Uncommon to occasional in 


tion. Indigenous, also found in Tonga and Fiji. 


Mammea glauca (Merr.) Kost. 

manapau 

Medium-sized tree with opposite, coriaceous leaves, 

showy white flowers, numerous yellow stamens, and 

an ellipsoid fruit. Uncommon in coastal to montane 

forest, reported from 20 to 900 m elevation. Endem- 

ic. Whistler 2764!, 3836, 8534!, 8698!. 

Terminalia catappa L. 

talie 

COMBRETACEAE 

Large tree with alternate, subsessile, obovate leaves, 

spikes of small white flowers, and large, conspicu- 

ously winged fruits. Occasional in littoral to lowland 

forest, reported mostly from near sea level, but 

occasionally to 260 m elevation. Indigenous or 

possibly a European introduction, widespread in the 

Old World tropics. Setchell 351, 356; Comstock 

17; Whistler 375 1, 8917!. 

Terminalia richii A. Gray 

malili 

Large tree with alternate, lanceolate leaves, small, 

spikes of white flowers, and purple, compressed- 

globose fruits. Uncommon in lowland to montane 


nous, also found in Fiji and Niue. Whistler 2955, 

9055!. 

*Terminalia samoensis Rechinger 

talie 

Medium-sized tree with alternate, petiolate, obovate 

leaves, spikes of white flowers, and unwinged, 

compressed-globose fruits. Uncommon in undis- 

turbed littoral forest, reported only near sea level. 

Indigenous, ranging from the Celebes to Tahiti. 

Christophersen 1 176; Whistler 2954, 5730. 

CONNARACEAE 

Rourea minor (Gaertn.) Alston 

Woody climbing vine with simple, alternate, ovate 

leaves, white flowers, and a fruit that opens to 

expose the single red seed. Occasional in lowland to 


tion. Indigenous to Samoa, ranging from India to 

Niue. Whistler 2883, 9201. 

*Ipomoea alba L. 

[ung-choi] 

CONVOLVULACEAE 

Creeping or low-climbing vine with adventitious 

roots, alternate, cordate leaves, and a large, showy 

white, salverform corolla having a long, narrow 

tube. Uncommon in disturbed places, reported from 

ca. 100 to ca. 600 m elevation. A modern intro- 

duction, pantropic in distribution. Whistler 2936. 

"Ipomoea aquatica Forssk. 

Creeping vine with alternate, hastate leaves and a 

white to light purple, rotate corolla. Uncommon in 

disturbed wet places, reported only from the low-

lands. A modern introduction, native to the Old 

World tropics. Whistler 8033a. 

lpomoea littoralis B1. 

palulu 

Scrambling herbaceous vine with alternate, cordate 

leaves and ashowy purple, rotate corolla. Occa- 

sional in sunny coastal and lowland areas, and 

sometimes weedy in plantations, reported from 10 to 

300 m elevation. Indigenous, widespread throughout 

the Pacific. Setchell 106; Eames T13; Wilder 88; 

Garber 91 I; Diefenderfer 16; Meebold 8233; 

Yuncker 9384; Wisner 85; Whistler 8637!. 

lpomoea macrantha Roemer & Schultes 

Sprawling or scrambling vine with alternate cordate 

leaves and showy white flowers with a long corolla 

tube and spreading limb. Occasional to common in 

sunny coastal areas, reported from near sea level to 

75 m elevation. Indigenous, widespread throughout 

the tropics. Long 3097; Whistler 2910, 8567!. 

Ipomoea pes-caprae (L.) R. Br. 

[beach morning-glory] fue moa 

Prostrate vine with purple stems, alternate leaves 

bilobed at the tip, and a showy purple, rotate corolla. 

Common to abundant on rocky and sandy shores, 

reported only near sea level. Indigenous, pantropic 

in distribution. Setchell 89; Wisner 21, 110; 

Whistler 1425, 8919!. 

*Merremia peltata (L.) Men. 

fue vao; fue lautetele 

Sprawling or high climbing vine with alternate 

peltate leaves, a showy white, rotate corolla, and a 

4-seeded capsule. Common in forest clearings and 

open forest of lowland to montane forest, reported 


from Africa to Tahiti. Wisner 103; Whistler 2813, 

3118a. 

Merremia umbellata (L.) Hall. f. 

Vine with alternate, hastate leaves and yellow, 

campanulate flowers arranged in umbels, locally 

common around central Tutuila in disturbed places, 


modern introduction, pantropic in distribution. 

Whistler 324, 2813, 8512!. 

*Operculina turpethum (L.) S. Manso 

Prostrate or climbing vine with winged stems, 

alternate cordate leaves with a mucronate tip, and a 

white campanulate corolla 2.5-4.5 cm long. Un- 

common in disturbed places in plantations and open 


tion. Indigenous, widespread in the Old World 

tropics. Wisner 1 13. 

Operculina ventricosa (Bertero) Peter 

Prostrate or climbing vine with unwinged stems, 

large, alternate, cordate leaves with an acuminate 

tip, and a white campanulate corolla 5-9 cm long. 

Occasional to locally common in disturbed places 

including beaches, and climbing into native forest 

canopy, reported from near sea level to 300 m 

elevation. A modern introduction indigenous to the 

Caribbean, frst reported from Polynesia in ca. 1939, 

now found in the Cook Islands, Niue, Tonga, 

Samoa, and the Marianas. It may hybridize with the 

preceding species. Whistler 2842, 8476. 

Stictocardia tiliifolia (Desr.) Hall. f. 

palulu 

Climbing vine with large, alternate, cordate leaves 

finely black-dotted on the lower surface, and a large, 

showy, lavender, rotate corolla. Occasional climb- 

ing in low vegetation in disturbed places, reported 

from near sea level to 100 m elevation. A modem 


Setchell248; Whistler 271 9. 

"Cucumis melo L. 

[Polynesian melon] 'atiu 

CUCURBITACEAE 

Prostrate vine with rough, alternate leaves, yellow, 

unisexual flowers solitary in the axils, and a small 

ovoid melon fruit. Rare in heavily disturbed places, 

reported only from the lowlands. A Polynesian 


Wilder 42; Wisner 92. 

*Luffa cylindrica (L.) Roemer var. insularum 

(A. Gray) Cogn. 

Prostrate or low-climbing vine with alternate leaves, 

large yellow flowers, and an ellipsoid fruit filled 

with a spongy material. Uncommon in sunny coastal 

areas and sometimes weedy in plantations, reported

from near sea level to 400 m elevation. Indigenous 

or a Polynesian introduction, widespread in the 

Pacific. Setchell 285; Meebold 16695; Yuncker 

9300. 

Momordica charantia L. 

[balsam pear] 

Herbaceous vine with alternate, deeply-palmately 

lobed leaves, yellow unisexual flowers on long, thin 

pedicels, fleshy, lumpy, orange, fusiform fruits, and 

large red seeds. Common in disturbed places, re- 

ported only from the lowlands. A modem intro- 

duction, native to the Old World tropics. Christo- 

phersen 989; Yuncker 9348; Whistler 1438, 2840, 

8952!, 9158. 

Zehneria mucronata (Bl.) Miq. 

Herbaceous vine with alternate, thin, mostly deltoid 

leaves, tiny white axillary flowers, and orange fruits 

rounded at the ends. Occasional in clearings in the 


tion. Indigenous, ranging from Taiwan to Tahiti. 

Whistler 2958, 8483, 8486, 8923!. 

Zehneria samoensis (A. Gray) Fosb. & Sachet 

Prostrate herbaceous vine with alternate, scabrous, 

deltoid leaves, tiny white axillary flowers, and 

orange, cylindrical fruits acute at the ends. Occa- 

sional in the littoral to coastal forest and sometimes 

inland as a weed, reported from near sea level to ca. 

200 m elevation. Indigenous, also found in Tonga 

and Niue. Lister s.n.; Setchell 290; Whistler 2908, 

8431.8931!. 

CUNONIACEAE 

Spiraeanthemum samoense A. Gray 

Medium-sized tree with simple, opposite leaves and 

racemes of tiny white flowers. Common to abun- 

dant in lowland (uncommonly) or montane to cloud 

forest, reported from 180 to 1850 m elevation. En- 

demic. Setchell 404, 549; Christophersen 1048, 

1067, 3506, 3576; Long 3063; Whistler 3838, 8518, 

8650. 

Weinmannia affinis A. Gray 

Small tree with simple (rarely 3-7-foliate), opposite 

leaves, conspicuous stipules, and racemes of tiny 

white flowers. Common in montane forest and 

scrub, reported from 300 to 1700 m elevation. Indi- 

genous, also found in Fiji and possibly Rarotonga. 

USEE s.n.?; Reinecke 487?; Christophersen 1201, 

3565; Whistler 2905, 3641 !, 9064. Weinmannia 

samoensis A. Gray is also reported from Tutuila, 

based on two specimens. The main difference bet- 

ween this and W. aflnis is compound versus simple 

leaves; however, individuals with both type of 

leaves have been found. Further field research is 

needed to sort this out. 

EBENACEAE 

"Diospyros major (Forst. f.) Bakh. 

Small tree with alternate ovate leaves bright green 

below and dark above, white, axillary 3-lobed urce- 

olate flowers, and solitary ovoid fruits. Uncommon 

in native forest, perhaps a relict of cultivation. 

Similar if not identical to D. christophersenii Fosb. 

of Western Samoa. Probably a Polynesian intro- 

duction, indigenous to Fiji and Tonga. Whistler 

8948. 

Diospyros elliptica (Forst.) P.S. Green 

'anume 

Small to medium-sized tree with alternate leaves, 

3-merous flowers, and red to yellow, ellipsoid fruits. 

Common to abundant in coastal and lowland forest, 


ranging from the Lau Islands (Fiji) to Niue. USEE 

s.n.; Rechinger 3729; Wilder 46; Diefenderfer 11; 

Wisner 123; Whistler 288 1,29 17, 3620. 

Diospyros samoensis A. Gray 

'au'auli 

Medium-sized tree with alternate leaves, white, 4- 

merous flowers, and a globose fruit with a persistent 

calyx of four reflexed lobes. Common to abundant 

in coastal and lowland forest, reported from near sea 

level to 350 m elevation. Indigenous, also found 

from the Lau Islands to Niue. Setchell 63, 333; 

Diefenderfer 16; Christophersen 1000; Yuncker 

9378; Wisner 121; Spence 481 !; Whistler 2882, 

3619. 

ELAEOCARPACEAE 

Elaeocarpus tonganus Burkill 

a'amati'e 

Medium-sized tree with alternate, ovate leaves 

turning red with age, many-petaled white flowers,

and blue spherical fruits. Occasional in lowland to 


tion. Indigenous, ranging from Tonga to the Austral 

Islands. Whistler 3632, 3849, 9006!. 

Elaeocarpus ulianus Christoph. 

Mctl~um-sizcd tree with alternate, ovate leaves, 

many-petaled white flowers, and large ovoid fruits 

containing a single irregularly edged seed. Occa- 


100 to 900 m elevation. Endemic. Whistler 2692!, 

2794,2801,3227,3603,8471,9170. 

EUPHORBIACEAE 

*Acalypha lanceolata Willd. 

Erect herb with pubescent, alternate leaves and 

inconspicuous, green, unisexual, axillary inflore- 

scences lacking a stalked, terminal flower. Uncom- 

mon in disturbed places, reported only from the 

lowlands. A Polynesian introduction, native to the 

Old World tropics. Whistler 2754,2949. 

Alcurites moluccana (L.) Willd. 

[candlenut] lama 

Medium-sized to large tree with alternate, pale 

green, palmately lobed leaves, small white flowers, 

and large green spherical fruits. Occasional in 

coastal and lowland forest, but most often found in 

cultivation, reported from 20 to 150 m elevation. A 

Polynesian introduction, ranging from Asia to 

Hawai'i. Setchell 199; Yuncker 9396; Whistler 

1445. 

Antidesma sphaerocarpum Muell. Arg. 

Shrub or small tree with alternate leaves, tiny 

inconspicuous flowers in racemes, and small, 

spherical, purple berries. Occasional in lowland 


elevation. Indigenous, also found in Futuna, 'Uvea, 

and Rotuma. Meebold 265 11, 26512; Yuncker 

9320, 9335; Long 1961, 1968; Whistler 1483, 2800, 

2941,2942,8034,8477,8494, 8538!, 8587!, 9147. 

Baccaurea taitensis Muell. Arg. 

Small tree with alternate leaves, tiny white, uni- 

sexual flowers in racemes, and brown, spherical 

fruits. Occasional in lowland to montane, reported 


385; Christophersen 11 19!, 1135!, 3522; Rainey 

2392; Whistler 27Ol,2747,2956a, 3541,8744!. 

Bischofia javanica BI. 

'o'a 

Medium-sized to large tree with alternate trifoliate 

leaves, tiny greenish flowers, and small, brown 

spherical fruits. Occasional in disturbed places and 

secondary to undisturbed forest, reported from near 

sea level to 850 m elevation. Indigenous or possibly 

a Polynesian introduction, ranging from India to 

Rarotonga. Setchell66; Bryan 984!; Diefenderfer 6; 

Mitchell 571; Yuncker 9338; Long 3085, 3114; 

Spence 5 16!; Whistler 1443. 

*Chamaesyce atoto (Forst. f.) Croizat in Degener 

pulu tai? 

Prostrate herb or low shrub with milky sap, opposite 

leaves, and greenish flowers and fruits in cyathia. 

Uncommon in sunny littoral areas, reported only 

near sea level. Indigenous, ranging from Ceylon to 

the Tuamotu Islands. Setchell 235, 304; Garber 

871 ; Whistler 8404. 

Chamaesyce hirta (L.) Millsp. 

[garden spurge] la'au fai moti 

Small succulent herb with milky sap, opposite, 

pubescent leaves, and tiny green flowers in dense 

axillary cyathia. Common in disturbed places, 

reported only from the lowlands. A modem intro- 

duction, pantropic in distribution. USEE s.n.? (n.s.); 

Setchell48, 56; Garber 800, 825; Lamoureux 3073; 

Whistler 89 15!. 

Chamaesyce hypericifolia (L.) Millsp. 

Erect herb with milky sap, glabrous, opposite leaves, 

and tiny white flowers in axillary cyathia. Occa- 

sional on roadsides and other disturbed places, 


duction, native to tropical America. Whistler 844 1, 

86OO!. 

Chamaesyce hyssopifolia (L.) Small 

Erect delicate herb with milky sap, small, opposite, 

oblong to linear leaves, and inconspicuous white 

flowers in leafy cyathia. Common in disturbed 

places, reported only from the lowlands. A modem 

introduction, native to tropical and subtropical 

America. Whistler 8045, 8914!.

*Charnaesyce prostrata (Ait.) Small 

[proslratc spurge] 

Small prostrate herb with milky sap, tiny opposite 

leave$, purple, nearly glabrous stems, and incon- 

c,picuous axillary flowers in cyathia. Occasional 

itround houses and in other sunny disturbed places, 


duction, native to tropical America. Setchell 504; 

Garbcr 9.50. 

Charnaesyce thyrnifolia (L.) Millsp. 

Prostrate herb with milky sap, tiny opposite leaves, 

pink, pubescent stems, milky sap, and inconspi- 

cuous flowers in axillary cyathia. Occasional 

around houses in in disturbed areas, reported only 


tropical America. Lamoureux 3066; Whistler 5 185, 

8953!. 

Drypetes vitiensis Croizat 

Small to medium-sized dioecious tree with leathery, 

alternate leaves, green flowers bearing numerous 

stamens, and female trees producing ellipsoid fruits. 

Uncommon to locally common in coastal to lowland 


tion. Indigenous, also found in Fiji, Tonga, and 

Niue. Whistler 3670a, 8069!, 8071!, 8413. 

*Euphorbia cyathophora Murray 

[wild poinsettia] 

Erect herb with milky sap, alternate to opposite 

leaves notched on the sides, and red-based floral 

bracts below the green, and apetalous flowers ar- 

ranged in cyathia. Occasional in disturbed places 

and pcrhaps casually cultivated, reported only from 


tropical America. Whistler 3535. 

"Euphorbia reineckei Pax 

Small, erect, scarcely branching shrub with milky 

sap, opposite leaves, greenish flowers in axillary 

cyathia, and green, 3-lobed fruits. Uncommon in 

lowland to cloud forest, reported from 280 to 1700 

m elevation. Endemic. Whistler 3673. 

Glochidion cuspidaturn Pax 

masame 

Small to medium-sized tree with alternate leaves 

pubescent below, tiny yellow, 3-merous flowers, 

and wheel-like fruits. Occasional in forest clearings, 


ranging from Melanesia to Samoa. Setchell 388; 

Christophersen 1028; Long 305 1; Whistler 2743, 

8514!. 

Glochidion ramiflorum Forst. 

masame 

Small to medium-sized tree with glabrous alternate 

leaves, tiny yellow, 3-merous flowers, and wheel- 

like fruits. Common in open and secondary forest, 


Indigenous, ranging from Melanesia to eastern 

Polynesia. Setchell 220, 537; Bryan 986; Garber 

790; Yuncker 9343; Long 3108, 3120a; Spence 

475!; Whistler 1472,2809, 3762. 

Macaranga harveyana (Muell. Arg.) Muell. Arg. 

lau pata; pata 

Small to medium-sized tree with peltate alternate 

leaves, tiny green flowers in spreading axillary 

panicles, and soft-spiny fruits. Common in dis- 

turbed places and early secondary forest, reported 


from Fiji to Tahiti. Setchell 253, 316; Swezey 57; 

Meebold 21361, 26591; Yuncker 9388; McKee 

2940; Comstock 18; Wisner 81; Spence 473; 

Whistler 1419,9020!, 91 11. 

Macaranga stipulosa Muell. Arg. 

lau fatu; patafatu 

Medium-sized tree with huge peltate, alternate 

leaves, tiny green flowers in spreading axillary 

panicles, and small, 3-lobed fruits. Occasional in 

lowland to montane forest, often in clearings, 


Setchell206; Diefenderfer 7; Whistler 2733. 

Omalanthus nutans (Forst. f.) Guillemin 

fogmamala; fanua mamala 

Small tree with milky sap, glacous, deltoid, alter- 

nate leaves, racemes of tiny cream-colored male 

flowers, solitary female flowers, and bilobed fruits. 

Occasional in open forest and disturbed places, 


ranging from New Caledonia and the Caroline 

Islands to Tahiti. Setchell 77, 153, 580; Wilder 31, 

92; Bryan 81, 966, 1013; Garber 847; Mitchell 503; 

Meebold 26566; Yuncker 9434 (n.s.); Wisner 82; 

Spence 487; Whistler 2862, 8767!, 9165.

Phyllanthus amarus Sch. & Thon. FABACEAE 

Erect herb with spirally arranged branches bearing 

small, alternate, elliptic, round-tipped leaves ar- 

ranged in one plane, and tiny green flowers and 

schizocarps on the lower side of the branches at the 

i~xils. Common in disturbed places, reported only 

liom the lowlands. A modern introduction, native to 

tropical America. Setchell 14, 534; Wilder 19; 

Garber 95 1, 955; Diefenderfer 25; Whistler 8954!. 

Phyllanthus debilis Klein ex Willd. 

Erect herb with longitudinally ridged stems, alternate, 

acute-tipped, . . oblanceolate leaves arranged in 

one plane, and tiny green flowers on the lower side 

of the branches at the axils. Occasional in disturbed 

places, reported from near sea level to 250 m elevation. 

A modern introduction, native to the Old 

World tropics. Whistler 8521 !. 

Phyllanthus urinaria L. 

Low herb with spirally arranged branches bearing 

small, alternate leaves arranged in one plane, reddish 

stems, tiny green, solitary, axillary flowers on the 

lower side of the branches, and a tiny globose 

schizocarp. Occasional in lawns and other disturbed 

places, rcported only from the lowlands. A modern 

introduction, native to tropical America. Lamo- 

ureux 3083; Whistler 8955!. 

Phyllanthus virgatus Forst. f. 

Erect herb with spirally arranged branches bearing 

small alternate leaves, tiny green flowers, and tiny 

green, spherical fruits. Uncommon in sunny or 

disturbed places in lowland forest, reported from 40 

to 260 m elevation. Indigenous or a Polynesian 

introduction, ranging from Asia to eastern Poly- 

nesia. Setchell 330; Whistler 8694!. 

*Richus communis L. 

Shrub with large, alternate, purplish, palmately 

lobed leaves, unisexual flowers with the apetalous 

male flowers in dense, yellow heads, and soft-spiny 

fruits containing six mottled seeds that are poison- 

ous. Uncommon in disturbed places, reported only 


Africa. Whistler 8897. 

Abrus precatorius L. 

[rosary pea] matamoso 

Climbing vine with alternate, pinnately compound 

leaves, lavender papilionaceous flowers, and red and 

black sceds in pods. Uncommon in coastal vege- 

tation, reported from near sea level to 100 m eleva- 

tion. Indigenous or possibly a Polynesian intro- 

duction, widespread in the tropics. Setchell 312; 

Wilder 18; Whistler 1484,2952,9030!. 

Adenanthera pavonina L. 

lo~a 

Medium-sized tree with pinnately compound leaves, 

small yellow flowers in spikes, and red seeds in 

long, twisted (when open) pods. Occasional in dis- 

turbed forest and casually cultivated, reported from 

near sea level to 200 m elevation. A modem intro- 

duction, widespread in cultivation in the tropics. 

Diefenderfer 17. 

Alysicarpus vaginalis (L.) DC. 

Prostrate herb with simple, alternate, oblong leaves, 

red papilionaceous flowers in racemes, and narrow, 

cylindrical pods. Common in lawns and sunny dis- 

turbed places, reported only from the lowlands. A 

modern introduction, native to the Old World 

tropics. Lamoureux 3076; Whistler 8972!. 

Caesalpinia major (Medic.) Dandy & Exell 

[yellow nickers] 'anaoso 

Scandent to climbing shrub with thorny stems, 

bipinnately compound leaves, distinct stipules, 

yellow flowers, and yellow-gray seeds in spiny 

pods. Occasional in littoral habitats to montane 


elevation. Indigenous, pantropic in distribution. 

Setchell 363; Whistler 2814?, 8932!. Caesalpinia 

bonduc Roxb., which is present on 'Aunu'u, is hard 

to distinguish from this species, and may also be 

present on Tutuila. It has conspicuous leafy 

stipules, which are lacking in C. major. 

Calopogonium muconoides Desv. 

Vine with stiffly hairy foliage and stems, alternate 

trifoliate leaves, and stalked, short racemes of 

lavender flowers. Locally common in disturbed 

places, reported from near sea level to ca. 600 m

elevation. A modern introduction, native to tropical 

America. Whistler 8569!. 

Canavalia cathartica Thou. 

Creeping or climbing vine with altemate, trifoliate, 

acute-tipped leaves, and mauve, papilionaceous 

llowers. Common climbing in trees of coastal and 

littoral forest, sometimes weedy farther inland, 


Indigenous, widespread in the tropics. Wisner 107, 

108; Whistler 8938. 

Canavalia rosea (Sw.)DC. 

fue fai va'a 

Creeping vine with alternate, trifoliate, notched or 

round-tipped leaves, and mauve, papilionaceous 

flowers. Occasional on sandy beaches, reported 

only near sea level. Indigenous, widespread in the 

tropics. Setchell 194; Whistler 892O!. 

"Crotalaria pallida Ait. 

[rattlepod] 

Shrub with alternate, trifoliate leaves, dense ter- 

minal racemes of yellow papilionaceous flowers 

marked with red lines, and glabrous inflated pods. 

Common in disturbed places, reported only from the 

lowlands. Setchell 57. 

*Crotalaria retusa L. 

Subshrub with simple, alternate, oblanceolate 

leaves, yellow and reddish papilionaceous flowers, 

and inflated pods. Uncommon in disturbed places, 


duction, native to Asia. (Whistler). 

Dendrolobium umbellatum (L.) Benth. 

lala 

Shrub with alternate trifoliate leaves, small white 

papilionaceous flowers, and small, jointed, several- 

seeded pods. Common in the littoral strand, repor- 

ted from near sea level to 260 m elevation. Indige- 

nous, ranging from Mauritius to Niue. Setchell287; 

Garber 820; Bryan 979 (n.~.), 1375; Wilder 224; 

Meebold 8242, 26584; Comstock s.n.; Wisner 17; 

Long 3092; Spence 500; Whistler 2855. 

Derris trifoliata Lour. 

fue o'ona 

Climbing woody vine with alternate trifoliate leaves, 

small mauve, papilionaceous flowers, and a flat- 

tened, suborbicular pod. Uncommon in littoral and 

coastal forest, reported from near sea level to 130 m 

elevatjon. Indigenous, widespread in the Pacific as 

far east as Rarotonga. Garber 853; Whistler 3539, 

8748!, 9172. 

*Desmanthus virgatus (L.) Willd. 

Erect subshrub with alternate, bipinnately compound 

leaves, globose heads of flowers having showy 

white stamens, and a long narrow pod. Uncommon 

in disturbed places, reported only from the lowlands. 

A modem introduction, native to tropical America. 

Whistler 5 186, 8445. 

*Desmodium heterocarpon (L.) DC. 

Small shrub with altemate trifoliate leaves, lavender 

papilionaceous flowers, and jointed fuzzy pods. 

Occasional in sunny disturbed areas and femlands, 


Polynesian introduction or perhaps indigenous, 

ranging from Southeast Asia to Hawai'i. USEE s.n. 

(n.s.); Wilder 13; Whistler 1486,2933. 

Desmodium incanum DC. 

Herb with alternate, trifoliate leaves, variegated, 

oblong, leaflets, mauve papilionaceous flowers, and 

fuzzy, jointed pods. Occasional to common in lawns 

and sunny disturbed areas, reported only from the 


America. Whistler 329, 1452, 8956!. 

Desmodium tortuosum (Sw.) DC. 

Tall, erect, somewhat woody herb with alternate 

trifoliate leaves, mauve papilionaceous flowers, and 

twisted, jointed pods. Occasional in disturbed 



303, 1454, 891 l!. 

Desmodium triflorum (L.) DC. 

Prostrate herb with altemate trifoliate leaves, short 

racemes of mauve, papilionaceous flowers, and a

papery, jointed pod. Occasional in disturbed places, 


duction, native to the Old World tropics. Setchell 

233; Eames T2; Garber 799, 945; Lamoureux 3056; 


*Entada phaseoloides (L.) Merr. 

[St. Thomas bean] fue inu 

High-climbing woody vine with alternate, pinnately 

compound leaves, small, dense racemes of flowers 

bearing showy white stamens, and large woody pods 

containing large, flat round seeds. Occasional in 

coastal to lowland forest, reported from near sea 


east Africa to the Austral Islands. Whistler 3624, 

3746,8485,9138. 

* Erythrina fusca Lour. 

lalapa 

Thorny, medium-sized tree with alternate trifoliate 

leaves, oval leaflets, and large showy red papilio- 

naceous flowers. Occasional in swampy coastal 

places, reported only near sea level. A modern 

introduction or possibly indigenous, pantropic in 

distribution. Christophersen 1035; Whistler 8061. 

* Erythrina subumbrans (Hassk.) Merr. 

[dadap] gatae palagi 

Medium-sized tree with alternate, trifoliate leaves, 

racemes of reddish orange flowers, and long pods. 

Occasional in disturbed places, planted as a 

nitribing plant for cacao and escaping, reported 

from near sea level to ca. 600 m elevation. A 

modcrn introduction, native from Southeast Asia to 

Malaysia. Whistler 3082. 

Erythrina variegata L. 

[coral tree] gatae 

Large thorny tree with alternate trifoliate leaves, 

broadly ovate leaflets, large showy red papilionaceous 

flowers in racemes, and large, black, 

cylindrical pods. Occasional in littoral to lowland 


Indigenous, ranging from tropical Asia to the 

Marquesas. Setchell 197, 302; Mitchell 587; 

Comstock 20, 21, 27; Yuncker 9352; Wisner 112; 

Dutton 1 12; Whistler 2834. 

"Indigofera hirsuta L. 

Small subshrub with alternate, pinnately compound 

leaves, small, racemes of salmon-colored papilionaceous 

flowers, and cylindrical pods. Rare in 

lawns and other disturbed places, reported only from 

the lowlands. A modern introduction, native to the 

Old World tropics. Whistler 2969. 

*Indigofera suffruticosa Miller 

[indigo] 

Widely branching shrub with alternate, pinnately 

compound leaves, racemes of small, salmoncolored 

papilionaceous flowers, and short, cylindrical 

pods. Occasional to common in disturbed 

places, reported from near sea level to ca. 200 m 


America. USEE s.n.? (n.~.); Wilder 49. 

Inocarpus fagifer (Parkinson) Fosb. 

[Tahitian chestnut] ifi 

Large tree with a fluted trunk, large simple, alternate 

leaves, tiny white flowers, and a large, thick-shelled 

fruit containing a large edible seed. Occasional to 

common in coastal to lowland forest, reported from 

near sea level to 350 m elevation. A Polynesian 

introduction, ranging from tropical Asia to the 

Marquesas. Graeffe 1386; Setchell 84, 355; 

Christophersen 893; Diefenderfer 12; Yuncker 

933 1; Whistler 3286. 

Intsia bijuga (Colebr.) Kuntze 

ifilele 

Large tree with alternate, pinnately compound leaves, 

showy red and white flowers, and a large 

woody pod. Uncommon to occasional in coastal to 

lowland forest, reported from near sea level to 200 

m elevation. Indigenous, ranging from East Africa 

to Samoa. Setchell 129a; Whistler 8046, 8664!. 

Leucaena leucocephala (Lam.) de Wit 

[wild tamarind] fuapepe 

Shrub or small tree with alternate, bipinnately compound 

leaves, globose heads of flowers having 

showy white stamens, and flattened pods. Occasional 

to locally abundant in dry, disturbed places, 


A modern introduction, native to tropical America. 

Setchell 180; Yuncker 9395; Whistler 8604!.

Macroptilium lathyroides (L.) Urb. 

Erect to ascending, scarcely branching herb with 

alternate, trifoliate leaves, maroon papilionaceous 

flowers, and long, narrow, cylindrical pods. Occa- 

sional in disturbed places, reported only from the 


Amcrica. Lamoureux 305 1; Whistler 361 1,9018!. 

Mimosa invisa Mart. ex Colla 

vao fefc I'alagi 

Scrambling, thorny shrub with alternate, bipin- 

nately compound leaves with the pinnae arranged 

pinnately, and pink flowers in globose heads. 

Common in disturbed places such as roadsides, 




Mimosa pudica L. 

[sensitive plant] vao fefe 

Somewhat woody herb with thorny stems, alternate, 

bipinnately compound leaves with palmately arran- 

ged pinnae, and pink flowers in dense globose 

heads. Common in lawns, croplands, and waste 

places, reported only from the lowlands. A modern 

introduction, native to tropical America. Setchell 

33; Whistler 8974!. 

Mucuaa gigantea (Willd.) DC. 

Sue inu 

High-climbing woody vine with alternate trifoliate 

leaves, hanging racemes of showy white papilio- 

naceous flowers, and smooth pods. Occasional to 

common in littoral to lowland forest, reported from 

near sea level to 200 m elevation, rarely up into 

cloud forest at 1200 rn. Indigenous, ranging fiom 

India to Hawai'i. Setchell 322; Yuncker 9389; 

Wisner 129; Whistler 2884. 

Paraserianthes falcataria (L.) I. Nielsen 

tamaligi 

Large tree with alternate, bipinnately compound 

leaves with small leaflets (ca. 5 mm wide), short 

spikes of flowers with showy white stamens, and a 

flattened pod. Occasional in disturbed areas, 

reported from 200 to 350 rn elevation. A modern 

introduction, native to Melanesia. Whistler 8605!. 

*Pongamia pinnata (L.) Men. 

Large tree with alternate, pinnately compound 

leaves, racemes or panicles of white to pink, papi- 

lionaceous flowers, and woody pods. Rare in littoral 

to coastal forest, reported only near sea level. Indi- 

genous, ranging from the Seychelle Islands to 

Samoa. Setchell3 1 1. 

Pueraria lobata (Willd.) Ohwi 

[kudzu] a'a 

Hairy vine with large, alternate, lobed, trifoliate 

leaves, violet papilionaceous flowers with a blotch 

of yellow, and a long, hairy pod. Occasional in 

disturbed places, reported only from near sea level 

to 350 m elevation. A Polynesian introduction, 

native to Asia. Garber 849; Whistler 1288, 9010. 

*Senna alata (L.) Roxb. 

[candelabra plant] la'au fai lafa 

Shrub with alternate, pinnately compound leaves 

bearing oval leaflets, showy orange and yellow 

flowers, and a black, winged pod. Uncommon in 

disturbed places and sometimes in wetlands, 



Whistler 2837, 3288. 

*Senna sophera (L.) Roxb. 

Subshrub with alternate, pinnately compound 

leaves, ovate leaflets, yellow, 5-merous flowers, and 

narrow, flattened pods. Rare in disturbed places, 

perhaps no longer found in Samoa, reported only 

from the lowlands. A Polynesian introduction, 

native to the Old World tropics. Setchell293. 

.*Senna tora (L.) Roxb. 

[peanut weed] vao pinati 

Subshrub with alternate, pinnately compound 

leaves, obovate leaflets, yellow, 5-merous flowers, 

and long, linear pods. Uncommon in pastures and 

other disturbed places, reported only from the low- 

lands. A modern introduction, native to the Old 

. World tropics. Setchell 529. 

*Sophora tomentosa L. 

Shrub with silvery, alternate, pinnately compound 

leaves, showy yellow papilionaceous flowers, and 

narrow pods constricted between the seeds. Uncom-

mon on sandy beaches and spits, reported only near 

sea level. Indigenous, pantropic in distribution. 

Setchell 127; Whistler 353 1. 

"Uraria lagopodoides (L.) Desv. ex. DC. 

Low subshrub with alternate, trifoliate leaves and 

purple, papilionaceous flowers in a dense, hairy, 

terminal raceme. Uncommon in lawns and other 

disturbed places, reported only in the lowlands. A 

Polynesian introduction, native to South Asia. 

Setchell 350; Wilder l I. 

*Vigna adenantha (G. F. W. Meyer) Markchal, 

Mascherpa, & Stainier 

Prostrate vine with alternate, trifoliate leaves, lave- 

nder and white, contorted papilionaceous flowers, 

and cylindrical pods. Uncommon in coastal areas 

and plantations, reported only from the lowlands. 

Probably indigenous, pantropic in distribution. 

Mitchell 538; Wisner 91; Whistler 308, 8733. 

Vigna marina (Bum.) Merr. 

[beach pea] fue sina 

Prostrate vine with alternate trifoliate leaves, yellow, 

papilionaceous flowers, and black cylindrical pods. 

Common to gbundant on the littoral strand, occa- 

sionally inland on roadsides, reported from near sea 

level to ca. 300 m elevation. Indigenous, pantropic 

in distribution. Setchell 101, 559; Wilder 28; 

Garber 954; Comstock s.n.; Yuncker 9360; Wisner 

133; Whistler 1428,3293,9022!. 

Casearia sp. nova 

FLACOURTIACEAE 

Small tree with alternate leaves, white flowers, and 

an ovoid capsule. Occasional in lowland forest, 


Setchell 282; McKee 2957; Kuruc 122, 153; 

Whistler 1463,2723,2868, 8584!, 9166. 

Erythrospermum acuminatissimum (A. Gray) 

A. C. Smith 

Small tree with alternate leaves, racemes of white or 

pink flowers, and a 3-valved capsule. Occasional in 


m elevation. Indigenous, also found in Fiji. 

Setchell 267, 278; Christophersen 999, 3487, 3542; 

Whistler 2892, 5726, 8050, 8462, 8669!. 

Flacourtia rukam Zoll. & Mor. ex Mor. 

filimoto 

Medium-sized tree with alternate leaves having 

toothed margins, small, white, apetalous flowers 

bearing many stamens, and a red spherical edible 

fruit. Occasional in lowland to montane forest, 

reported from ca. 50 to 500 m elevation. Indi- 

genous, also found in Tonga and westward 

(excluding Fiji). Setchell 262, 573; Garber 920; 

Bryan 982 (n.s.); Mitchell 570; Comstock 1; 

Yuncker 940 1 a; Wisner s.n.; Spence 466! ; Whistler 

I474,2781,3114,3633,86l7!. 

Homalium whitmeeanum St. John 

Small tree with alternate, toothed leaves and white, 

many-petaled flowers in dense, hanging racemes. 

Uncommon in open forest on lava flows and in 

scrub forest, reported from near sea level to 550 m 

elevation. Indigenous, also found in Tonga. 

Christophersen 3491; Whistler 2976, 8454, 8694!. 

GESNERIACEAE 

*Cyrtandra geminata Reinecke 

Shrub with opposite leaves, large, showy white 

flowers, and a pair of terminal bracts on the 

peduncle. Occasional in montane scrub, reported 

from 450 to 600 m elevation. Endemic to Tutuila. 

Reinecke 477; Setchell 548; Christophersen 104 1, 

3563; Long 3069; Whistler 3669,9068. 

*Cyrtandra longipedunculata Rechinger 

Shrub with opposite ovate to elliptic, cymes of 

showy white flowers on long peduncles, and ovoid 

fleshy fruits. Uncommon in lowland to montane 

forest, reported from 250 to 910 m elevation. Ende- 

mic. Bryan 83. 

Cyrtandra pulchella A. Gray 

Shrub with opposite, elliptic to falcate, glabrous 

leaves, large, 1-7-flowered cymes with very small 

bracteoles, and large, showy white flowers. Occa- 


200 to 800 m elevation. Endemic. USEE s.n.; 

Setchell340, 553; Bryan 78; Yuncker 9413; Wisner 

153; Christophersen 983, 1266, 1818!, 1829!; 

Spence 5 12!; Whistler 2709.

Cyrtandra samoensis A. Gray 

momolea? 

Shrub with large elliptic to ovate, opposite leaves, 

small white sympetalous flowers in axillary cymes, 

and fleshy orange berries. Occasional in littoral to 


m clcvation. Indigenous, also found in Tonga and 

Niue. Graeffe 493, 496; Setchell 9; Bryan 75, 957; 

Christophersen 98 1 ; Mitchell 577; Meebold 8258, 

16724, 265 13; Bayliss s.n.; Wisner 155; Lamoureux 

301 9; Long 3088, 3099, 3 113; Spence 478!; Clay 

s.n.; Fosberg 54920; Whistler 2859, 2878, 3 1 18, 

86O9!. 

*Cyrtandra sp. nova 

Shrub with large opposite leaves, cymes of small 

white flowers subtended by sheathing bracts, and 

orange berries. Occasional in lowland forest, 

reported from 250 to 450 m elevation. Endemic to 

Tutuila. Setchell 368, 567; Christophersen 3583?; 

Long 1962, 3071, 3078, 3 110; Whistler 2731. 

Hernandia nymphaeifolia (Presl) Kub. 

[Chinese lantern tree] pula 

Large, spreading tree with alternate, peltate leaves, 

small white flowers, and a green to white or pink, 

lantern-like fruit enclosing a single black marble- 

like seed. Common in littoral forest, particularly on 

sandy shores, reported only near sea level. Indige- 

nous, ranging from East Africa to the Marquesas. 

Setchell 143; Garber 961 ; Mitchell 53 1 ; Wisner 96, 

105; Whistler 2946, 902 1 !. 

ICACINACEAE 

Citronella samoensis (A. Gray) Howard 

Small tree with glossy, alternate leaves, racemes of 

- - 

small white flowers, and large, ellipsoid fruits. 

Occasional in lowland to cloud forest, reported from 

50 to 1600 m elevation. Indigenous, also found in 

Tonga. Setchell 2 17, 261, 443; Wilder 36; Garber 

899; Mitchell 522; Yuncker 9409; Whistler 1462, 

2702,2865, 8706!. 

GOODENIACEAE 

*Medusanthera samoensis (Reinecke) Howard 

matamo 

Scaevola taccada (Gaertn.) Vahl 

to'ito'i 

Small to medium-sized tree with oblong, alternate 

leaves, white flowers with hairy, protruding 

Shrub with waxy, opposite, spathulate leaves, white stamens, and a flattened-ellipsoid fruit with a pink, 

one-sided flowers, and white fleshy fruits. Abundant fleshy protuberance on one side. Occasional in 

on the seaward margin of littoral forest and some- lowland to cloud forest, reported from 60 to 1400 m 

times inland up to 300 m elevation. Indigenous, elevation. Endemic. Garber 919; Whistler 2961. 

widespread throughout the tropical Pacific. Setchell 

I 16, 229, 346; Garber 947; Diefenderfer 6; Wisner 

LAMIACEAE 

20; Long 2600; Lamoureux 3022; Whistler 1426, 

2854. 

*Pogostemon cablin (Blanco) Bentha 

[patchouli] patiale 

HERNANDIACEAE 

Hernandia moerenhoutiana Guillemin 

pipi 

Large, soft-wooded tree with alternate leaves, small 

white flowers, and a reddish, lantern-like fruit en- 

closing the single large black seed. Occasional in 

lowland to cloud forest, reported from 100 to 1300 

m elevation. Indigenous, ranging from the Solomon 

Islands to the Tahiti. Setchell 365, 445; Christo- 

phersen 1058, 3523; Spence 469!(n.s.); Whistler 

2803,2964,3 124,866 1. 

Subshrub with opposite, toothed, ovate, fragrant 

leaves and verticils of bilabiate flowers in spicate 

racemes or panicles (but uncommonly flowering). 

Uncommon to locally abundant in disturbed places 

and along trails, reported from near sea level to 200 

m elevation. A modem introduction, native to South 


LAURACEAE 

Litsea samoensis (Christoph.) A. C. Smith 

papaono 

Medium-sized to large tree with alternate, reddish- 

tomentose young leaves, orange flowers, a persi- 

stent, cup-like calyx, and an ovoid fruit. Occasional

in lowland to cloud forest, reported from 100 to 

1200 m elevation. Indigenous, also found on Niue. 

Graeffe s.n.; Setchell 286, 448, 462, 571; Long 

3052; Whistler 2782, 3647, 8642!. 

LOGANIACEAE 

Fagraea berteroana A. Gray ex Benth. 

pualulu 

Large tree with glossy opposite leaves, large, showy, 

tubular, white to pale orange flowers, and a large 

orange fruit containing many black seeds. Common 

in lowland to montane forest, reported from ca. 50 

to 930 m elevation. Indigenous, ranging from Fiji to 

the Marquesas. Setchell 367; Garber 882; Chri- 

stophersen 1208; Meebold 21415; Yuncker 9341, 

9392; Lamoureux 3009; Long 3 105; Whistler 2804, 

8987!. 

Geniostoma rupestre Forst. 

taipoipo; lau mafatifati 

Small tree with simple, opposite leaves, tiny white 

flowers, and a small greenish capsule containing 

many red seeds. Common in coastal to cloud forest, 

reportcd from near sea level to 1600 m elevation. 

Indigenous, ranging from Taiwan and the Celebes to 

the Cook Islands. Setchell 117, 264, 578; Seale s.n.; 

Diefenderfer 29; Yuncker 9336; Kuntze 23007; 

Meebold 12646; Whistler 1272, 1480, 2766!, 2850, 

2940,3645, 8670!, 9067. 

LORANTHACEAE 

Decaisnina forsteriana (J. A. & J. H. Schultes) 

Barlow 

tapuna 

Hemiparasitic shrub with opposite leaves, long, 

narrow red, erect flowers with a flared calyx, and a 

small, ellipsoid, fleshy red fruit. Occasional in 

littoral to montane forest, reported from near sea 


Fiji to the Cook Islands. Reinecke 159a; Setchell 

372, 575; Wilder 227; Mitchell 569; Spence 491!; 

Whistler 2783, 2965. 

LYTHRACEAE 

Cuphea carthagenensis (Jacq.) Macbr. 

Sticky herb with opposite leaves, purple, 6-parted 

axillary flowers, and small ovoid capsules surroun- 

ded by the longitudinally ribbed calyx. Occasional 

in pastures and wet, sunny places, reported only 

from near sea level to 650 m elevation. A modem 


542a; Lamoureux 3069; Whistler 3234, 8621 ! . 

*Gossypium hirsutum L. 

[native cotton] vavae 

MALVACEAE 

Shrub with alternate, palmately Iobed leaves, white 

monadelphous flowers, and a splitting capsule filled 

with cotton. Occasional on open coastal slopes of 

tuff-cone islets, reported only from the lowlands. 

Indigenous, widespread from Samoa to South 


*Hibiscus abelmoschus L. 

fau Tagaloa 

Small hairy subshrub with alternate, palmately lobed 

leaves, large showy yellow, monadelphous flowers 

with purple at the base, and a hairy ovoid capsule. 

Uncommon to occasional in disturbed (often wet) 

places, reported from near sea level to 350 m ele- 

vation. A Polynesian introduction, probably native 

to tropical Asia. Setchell 59, 241a, 525. 

Hibiscus tiliaceus L. 

[beach hibiscus] fau 

Medium-sized, scrambling tree with alternate cor- 

date leaves, yellow flowers purple at the base, and a 

5-celled capsule. Common to abundant in littoral 

forests, mangrove swamp margins, and disturbed 

and secondary forest, reported from near sea level to 

650 m elevation. Indigenous, pantropic in distri- 

bution. Setchell 53, 242; Eames T30; Diefenderfer 

15; Cornstock 30; Mitchell 534; Yuncker 9399; 

Wisner 15; Dutton 6; Long 2596; Herbst 6564; 

Whistler 2812,3285, 8612!. 

Sida rhombifolia L. 

mautofu 

Erect subshrub with ovate, alternate leaves, pale 

orange, monadelphous flowers on long axillary 

pedicels, and a beaked, wheel-shaped, 9-12-merous 

schizocarp. Common in disturbed places, reported 

from near sea level to 600 m elevation. A Polyne- 

sian introduction, native to the Old World tropics 

but now pantropical. Setchell 8; Wilder 23; Garber 

8 15; Diefenderfer 8; Whistler 86 18!.

'Thespesia populnea (L.) Sol. ex Corr. 

[Pacific rosewood] milo 

Medium-sized tree with glossy, alternate, cordate 

leaves, yellow flowers purple at the base, and a 

wbglobose, non-splitting fruit. Occasional to com- 

mon in littoral forest, reported only near sea level. 

Indigenous, ranging from tropical Africa to eastern 

Polynesia. Setchell 95; Christophersen 1206; 

Yuncker 9372; Whistler 1449,3283,3763, 8918!. 

"Urena lobata L. 


Erect subshrub with alternate leaves, pink monadel- 

phous flowers, and a subglobose, bur-like fruit. 

lJncommon in disturbed places, reported only from 

the lowlands. A Polynesian introduction, probably 

nntivc to tropical Asia. Garber 897; Diefenderfer 


MELASTOMACEAE 

*Astronidium navigatorum Christoph. 

Small tree with opposite leaves 3-5-veined from the 

base, petioles of mature leaves orange and less than 

2 cm long, small white, 4-merous flowers with geni- 

culate stamens, and tiny capsules. Occasional in 


m elevation. Endemic. Reinecke 473; Christopher- 

sen 1059, 1070, 3509, 3534; Whistler 2710,3649, 

3839, 8937. 

Astronidium pickeringii (A. Gray) Christoph. 

Small tree with opposite leaves 3-5-veined from the 

base, pctioles of mature leaves red and more than 2 

cm long, calyx entire to shallowly lobed, and 

5-merous while flowers with geniculate stamens. 

Occasional to common in lowland to montane forest, 

reported from 130 to 900 m elevation. Plants 

found on trachyte soil have a distinctly lobed calyx 

and more compact inflorescence, and may represent 

a separate endemic species. Endemic. USEE s.n. 

(n.~.); Powell s.n.; Graeffe s.n.; Reinecke 572?; 

Setchell 34 1, 40 1; Bryan 92; Christophersen 1047, 

1064, 3507; Mitchell 567; Long 2607, 3005, 3072; 

Spence 471 !; Whistler 2712, 2748, 2977, 3758, 

3840, 872 1 !. 

Clidemia hirta (L.) D. Don 

[Koster's curse] 

Shrub with opposite, hispid leaves palmately veined 

from the base, small white flowers in axillary clust- 

ers, geniculate stamens, and purple fruits. Common 

to abundant in sunny disturbed places and native 

forests and scrub, reported from near sea level to 

900 m elevation. A modern introduction, native to 

tropical America. Wisner 5; Long 1966, 2604, 3048; 

Whistler 3 1 1. 

Dissotis rotundifolia (Sm.) Triana 

Low herb with opposite, ovate leaves, 4-merous 

flowers with showy purple-pink petals, and yellow, 

geniculate stamens. Uncommon but sometimes 

locally common in coconut plantations and other 

disturbed places, reported only from the lowlands. 

A modern introduction, native to tropical Africa. 

Whistler 8480, 8968!. 

Medinilla samoensis (Hochreut.) Christoph. 

Woody climber with opposite, 3-5-veined leaves, 

pink 4-merous flowers with geniculate stamens, and 

a subglobose capsule. Occasional in montane forest, 


probably also found on Futuna. Graeffe s.n.; Chri- 

stophersen 1 14 1 !, 357 1; Yuncker 9404; Lamoureux 

3006; Long 1952; Whistler 2779. 

Melastoma denticulatum Labill. 

fua lole 

Shrub with opposite pubescent leaves 3-5-veined 

from the base, white flowers with geniculate sta- 

mens, and a scaly fruit opening to expose the red 

pulp. Common in forest clearings and disturbed 


vation. Indigenous, ranging from New Caledonia to 

Tahiti. Reinecke 529; Setchell 30, 378, 524; Eames 

T4; Bryan 82; Wilder 1, 14, 81; Christophersen 

1060, 3492, 3568, 3578; Mitchell 507; Wisner 76, 

137; McKee 2942; Long 2602, 31 17; Whistler 2927, 

8613!. 

MELIACEAE 

Aglaia samoensis A. Gray 

laga'ali 


leaves, tiny fragrant flowers in long panicles, and an

ellipsoid drupe. Occasional in lowland to cloud 

forest, reported from 20 to 1030 m elevation. En- 

demic. USEE s.n.; Setchell 156; Garber 879, 914; 

Diefenderfer 8; Mitchell 502; Meebold 25571, 

26590; Yuncker 9403; Wisner 15 1 ; Lamoureux 

3004; Whistlcr 2870, 8470,9168. 

I)ysoxylurn huntii Merr. 

maota mea 


leaves, 4-6 pairs of leaflets, white tubular flowers in 

short panicles, and subglobose fruits containing 4 

red seeds. Common in montane forest, rarely in 

lowland forest, reported from 150 to 1750 m ele- 

vation. Endemic. Setchell 366, 435, 438; Christo- 

phersen 3537; Spence 507!; Whistler 2763!, 2776!. 

Dysoxylurn maota Reinecke 

maota 


leaves, more than 6 pairs of leaflets, white tubular 

flowers with a whorl of bracts below the calyx, and 

wrinkled, yellow-brown, subglobose fruits contain- 

ing 4 red seeds. Common in coastal to montane 


tion. Indigenous, also found in 'Uvea and Futuna. 

Setchell 434; Bryan 1006; Wisner 50, 51; Kuruc 

105; Whistler 3536, 8397, 8568!, 9135. 

Dysoxylurn samoense A.Gray 

mamala; maota mamala 


leaves, more than 6 pairs of leaflets, white tubular 

flowers lacking bracts, and brown, subglobose fruits 

containing four red seeds. Common to abundant in 


level to 750 m elevation. Endemic. Setchell 150, 

358, 424, 433; Garber 916; Meebold 16720, 26595; 

Whistler 2843, 5732. 

*Xylocarpus moluccensis (Lam.) Roemer 

[puzzlenut] le'ile'i 

Small to medium-sized tree with alternate, pin- 

nately compound leaves, tiny white flowers in long 

panicles, and large green globose fruits containing 

several large tan seeds packed tightly together. 

Uncommon in littoral forest and mangrove swamp 

margins, reported only near sea level. Indigenous, 

ranging from Madagascar to Samoa. Setchell 439; 

Garber 1 1 18; Christophersen 1 177; Yuncker 9380; 

Whistler 28 10,28 1 1. 

MONIMIACEAE 

Hedycarya denticulata (A. Gray) Perk. & Gilg 

Small tree with alternate, dark green leaves, head- 

like inflorescences of tiny sessile yellow flowers, 

and glossy red fruits. Common in the lowland to 

cloud forest, reported from 150 to 1550 m elevation. 

Endemic. Setchell384, 566; Wilder & Swezey 229; 

Garber 846, 910; Christophersen 1140!; Mitchell 

568; Yuncker 9351; Long 3065, 3090; Spence 506; 


*Castilla elastica Sess6 

pulu mamoe 

MORACEAE 

Large tree with milky sap, large, oblong, alternate 

leaves, and inconspicuous flowers forming a sessile, 

cauliflorus aggregate fruit. Uncommon in disturbed 

forest, noted so far only on the western portion of 

the island (where it is locally common at Maloata) 

and 'lli'ili (where it is rare), reported from ca. 100 to 

700 m elevation (in Western Samoa). A modem 


8449. 

Ficus godeffroyi Warb. 

mati 

Small to medium-sized tree with milky sap, glabrous 

alternate leaves having a rounded base, and 

fig-like, often clustered fruits. Occasional in lowland 

to montane forest, reported from 20 to 900 m 

elevation. Endemic. Reinecke 76b; Setchell 68; 

Garber 936; Christophersen 982, 998, 1143!; 

Meebold 26517; Yuncker 9337; Spence 498, 499; 

Whistler 1282,2726, 3602, 8589!, 8992!. 

Ficus obliqua Forst. f. 

[banyan] aoa 

Large banyan tree with relatively few hanging roots, 

milky sap, alternate lanceolate leaves, and subglobose 

red fruits subtended by caduc~us bracts. 

Occasional to common in lowland to montane forest, 


ranging from Australia and the Celebes to 

Samoa. Setchell 198, 315, 422; Swezey s.n. (n.~.); 

Whistler 1467, 2923, 3671.

Ficus prolixa Forst. f. 

[banyan] aoa 

Large banyan tree with milky sap, alternate lanceo- 

late leaves, and yellow subglobose fruits subtended 

by persistent bracts. Occasional to common in 

lowlnnd forest, reported from 20 to 150 m elevation. 

Indigenous, ranging from New Caledonia to the 

Marquesas. Swezey sn.; Whistler 3671, 8467. 

Ficus scabra Forst. f. 

Small to medium-sized tree with milky sap, alter- 

nate scabrous leaves, and red to yellow fig-like 

fruits. Common to abundant in littoral forest, less so 

in lowland forest, reported from near sea level to 

300 m elevation. Indigenous, ranging, from New 

Caledonia to Niue. Setchell 148, 3 14, 421, 427, 

Meebold 8250, 16732; Comstock 31; Yuncker 9437; 

Wisner 11; McKee 2954; Whistler 1286, 2914, 

8598!, 91 14. 

Ficus tinctoria Forst. f. 

[dyer's fig] mati 

Small tree with milky sap, glossy alternate leaves, 

bracts at the base of the peduncle, and red to or- 

ange, subglobose fruits. Common in lowland to 


tion. Indigenous, ranging from India to the 

Marquesas. Graeffe 13 18; Bryan 76, 1010; Diefen- 

derfer 13, 18; Swezey s.n. (2); Mitchell 510, 574; 

Yuncker 9382; McKee 2955; Wisner 54, 120; 

Whistler 1441, 2856,2857, 8551!, 9143. 

Ficus uniauriculata Warb. 

Small to medium-sized tree with milky sap, alter- 

nate leaves having a one-sided auriculate base, 

scabrous surfaces, and fig-like fruits. Occasional in 

coastal to montane forest, reported from near sea 

level to 800 (1600?) m elevation. Endemic. Set- 

chell 69; Swezey 7; Christophersen 998; Meebold 

16732; Mitchell 509; Whistler 2879, 8981!, 9079, 

91 12. 

Myristica fatua Houtt. 

[Samoan nutmeg] 'atone 

MY RISTICACEAE 

Medium-sized tree with alternate leaves pale 

beneath, tiny yellow flowers, and a tan, ellipsoid 

fruit containing a large seed surrounded by a red 

aril. Common to abundant in lowland to montane 


tion. Indigenous, ranging from Vanuatu to Samoa. 

Setchell 1, 67; Wilder 91; Bryan 73; Garber 908; 

Christophersen 984, 985; Meebold 21351; Yuncker 

9326; Wisner 59, 66; Long 1976, 3084; Spence 

470!; Whistler 2721,2872, 8585!. 

Myristica hypargyraea A. Gray 

'atone 

Medium-sized tree with alternate leaves gray-green 

beneath, tiny yellow flowers, and a reddish-brown 

subglobose fruit containing a large seed surrounded 

by a yellow aril. Common to abundant in lowland 

to montane forest, reported from near sea level to 

930 m elevation. Indigenous, also found on Futuna, 

Alofi, and possibly Rotuma. Reinecke 445; Setchell 

342; Garber 917, 918, 928; Swezey & Wilder 58; 

Christophersen 996; Yuncker 94 19; Wisner 58; 

Whistler 2744, 8659!. 

MYRSINACEAE 

Embelia vaupelii Mez 

Woody climber with alternate leaves, white-dotted 

stems, and small white flowers. Uncommon in lowland 

to cloud, reported from near sea level to 1750 

m elevation. Indigenous, also found in Tonga. 

Setchell569; Wisner 56; Whistler 9149. 

Maesa tabacifolia Mez 

Shrub with alternate leaves, white-dotted stems, tiny 

white flowers in racemes or panicles, and small, 

flesh-colored hits. Occasional in lowland to montane 


elevation. Indigenous, also found in Tonga and Fiji. 

Lister s.n.; Setchell 581; Diefenderfer 3; Seale s.n.; 

Christophersen 3522; Yuncker 9359; Wisner 154; 

Lamoureux 3003; Spence 505!; Whistler 2762!, 

2887, 8815!. 

"Rapanea myricifolia (A. Gray) Mez 

Small tree with alternate coriaceous leaves, tiny 

white cauliflorus flowers, and small subglobose 

hits. Occasional in coastal to cloud forest, reported 

from 5 to 1500 m elevation. Indigenous, also 

found in Fiji. Setchell 382; Christophersen 1024, 

1053, 1065, 1202, 3547; Whistler 3851.

MY RTACEAE 

Decaspermum fruticosum Forst. 

nu'anu'a 

Shrub or small tree with opposite leaves, white 

5-merous flowers bearing numerous stamens, and a 

capsule fruit. Uncommon in fernlands, disturbed 

places, and montane scrub, reported from 100 to 500 

m elevation. Indigenous, ranging from Futuna to 

the Society Islands. Christophersen 1183, 35 11, 

3575; Whistler 3660, 8689!. 

"Eugenia reinwardtiana (BI.) DC. 

unuoi 


flowers bearing many stamens, and red succulent 

fruits. Uncommon in littoral and coastal forest, 

reported only from the lowlands. Indigenous, 

ranging from Micronesia and Fiji to Hawai'i. Powell 

s.n.; Whistler 3353, 8401. 

Mctrosideros collina A. Gray 

Medium-sized tree with opposite leaves, clusters of 

flowers with showy red stamens, and capsules 

containing numerous tiny seeds. Occasional to 

common in montane scrub, reported from 100 to 


Vanuatu to Hawai'i. Reinecke 643; Setchell 266; 

Christophersen 988, 1184, 1 188, 1194, 1197, 1198, 

1205, 3494, 3500, 3569, 3577, 3585!; Spence 497!; 

Whistler 3662, 8687!, 8723!. 

Psidium guajava L. 

[guava] ku'ava 

Small tree with flaky bark, opposite leaves, white 

flowers bearing many stamens, and a yellow, pink- 

fleshed, many-seeded fruit. Occasional in pastures 

and other disturbed places, reported from near sea 

level to 600 m elevation. A modem introduction, 

native to tropical America. Setchell 55; Whistler 

1485, 8960!. 

Syzygium brevifolium (A. Gray) C. Muell. 

Medium-sized tree with subsessile opposite leaves, 

terminal clusters of flowers bearing numerous white 

stamens, and small ovoid fruits. Occasional in 

montane scrub to cloud forest, reported from 235 to 

1300 m elevation. Endemic. USEE s.n.; Christo- 

phersen 1025, 1051, 1071; Whistler 3850, 8620!. 

Syzygium carolinense (Koidz.) Hosokawa 

Small to medium-sized tree with opposite leaves, 

loosely branching inflorescences, and small white 

flowers bearing numerous stamens. Occasional in 


m elevation. Indigenous, ranging from Micronesia 

to Samoa. Yuncker 9428!?; Spence 472!; Whistler 

2773!, 2780,2796a, 3543,3651,8484. 

Syzygium clusiifolium (A. Gray) C. Muell. 

asi vai 

Medium-sized tree with finely veined opposite 

leaves, widely branching, cauliflorus inflorescences 

of white flowers, and purple ellipsoid fruits. Com- 

mon to abundant in coastal to lowland forest, 


Indigenous, also found in Tonga, Niue, Wallis, and 

the Horne Islands. Lamoureux 3024a; Whistler 

1280,3755,8469, 8954!, 9107!. 

Syzygium dealatum (Burkill) A. C. Smith 

asi vai 

Medium-sized tree with opposite leaves, terminal 

inflorescences of white flowers bearing many 

stamens, and purple ovoid fruits. Occasional in 

coastal to lowland forest, reported from near sea 

level to 310 m elevation. Indigenous, also found in 

Tonga, Niue, and Uvea. Whistler 2886, 3750, 

8588!. 

Syzygium inophylloides (A. Gray) C. Muell. 

asi 

Large tree with finely veined leaves having a twisted 

attenuate tip, white flowers bearing numerous 

stamens, and yellowish ovoid fruits. Common in 

lowland to cloud forest, reported from near sea level 

to 750 m elevation. Indigenous, also found in Niue 

and Futuna. USEE sn.; Setchell 337, 570; Bayliss 

s.n.; Wilder 93; Diefenderfer 14; Christophersen 

1255; Whistler 2802, 8955!. 

*Syzygium neurocalyx (A. Gray) Christoph. 

'oli; fena 

Small tree with long, sessile, opposite leaves, large 

terminal sessile flowers bearing numerous white 

stamens, and large edible fruits. Rare in montane 

forest (in Western Samoa at least) and formerly 

cultivated, perhaps no longer found on Tutuila, 


Polynesian introduction or perhaps indigenous, also

found in Tonga, Futuna, and Fiji. Setchell 295, 

298b, 5 14; Mitchell 562. 

Syzygium samarangense (BI.) Merr. & Perry 

nonu vao? 

Small to medium-sized tree with opposite, sub- 

sc.;sile leaves, showy white flowers with numerous 

stamens, and large red fruits. Common in coastal to 


tion. Probably a modern introduction, native to 

Indo-Malaysia. Kuruc 107; Whistler 1461, 2871, 

9173. 

Syzygium samoense (Burkill) Whistler 

fena vao? 

Medium-sized tree with opposite leaves, inflore- 

scences with small bracts, white flowers bearing 

numerous stamens, and ovoid fruits. Occasional to 

common in lowland to cloud forest, reported from 

180 to 1200 m elevation. Endemic. Christophersen 

1008, 1 199, 3482, 3498; Rainey 2394; Whistler 

2688,2749,2957, 8658!, 9096!, 9341!. 

Boerliavia albiflora Fosb. 

Prostrate herb with alternate leaves, stalked clusters 

of tiny white flowers, and tiny sticky cylindrical 

seeds. Uncommon on rocky sea cliffs, reported only 

in coastal areas. Indigenous, widespread in the 

tropical Pacific. Whistler 8909!, 9026! 

Boerhavia repens L. 

ufi 'atuli 

Prostrate herb with alternate leaves, reddish stems, 

tiny pink flowers in stalked clusters, and tiny sticky 

fruits. Occasional as a weed in coastal habitats and 

villages, reported only from the lowlands. Indige- 

nous or possibly a Polynesian introduction, wide- 

spread in the Pacific. Garber 861; Wisner 45; 


I'isonia grandis R. Br. 

pu'a vai 

Large tree with a white trunk, alternate to whorled 

leaves, small white flowers in terminal clusters, and 

sticky cylindrical fruits. Common on undisturbed 

sandy shores, reported only near sea level. Indige- 

nous, ranging from Madagascar to Hawai'i. Setchell 

3 13; Whistler 2953, 9019!. 

Pisonia urnbellifera (Forst.) Seem. 

Small tree with leaves somewhat whorled, pink or 

white flowers, and sticky, long-cylindrical fruits. 

Occasional to common in lowland to montane 


tion. Indigenous, ranging from Madagascar to 

Hawai'i. Wisner 52; Whistler 2966, 8597!, 9 159. 

OLACACEAE 

*Anacolosa insularis Christoph. 

Small tree with alternate leaves, tiny white, axillary 

subsessile flowers, and small ovoid fruits. Rare in 


level to 450 m elevation. Endemic. Whistler 3348. 

*Ximenia americana L. 

moli tai 

Thorny shrub with small alternate leaves, white 

flowers hairy inside, and orange, globose, large- 

seeded fruits. Rare at the seaward margin of littoral 

forest, reported only near sea level. Indigenous, 

ranging from India to the Tuamotu Islands. 

Whistler 466 1, 8074. 

OLEACEAE 

Chionanthus vitiensis (Seem.) A. C. Smith 

Medium-sized tree with opposite leaves, small yel- 

low flowers bearing two stamens, and large, single- 

seeded, spindle-shaped fruits. Rare in coastal and 

lowland forest, reported only from the lowlands. 

Indigenous, also found in Fiji, Tonga, and Niue. 


*Jasminum betchei F. Muell. 

Thin, climbing vine with opposite leaves, showy 

white sympetalous flowers containing two stamens, 

and dark subglobose fruits. Uncommon in coastal to 


m elevation. Indigenous, also found in Fiji, Tonga, 

and Niue. Wisner 57; Whistler 3609,9175. 

Jasminum didymum Forst. f. 

Thin climbing vine with opposite, trifoliate leaves, 

white flowers with two stamens, and black, subglo- 

bose fruits. Occasional in coastal to cloud forest, 


Indigenous, ranging from Australia to the Society

Islands. Setchell 289, 348; Christophersen 149, 

1262; Whistler 9029!, 91 16. 

ONAGRACEAE 

Ludwigia hyssopifolia (G. Don) Exell 

Erect, red-stemmed herb with alternate leaves, 

small, yellow, 4-merous flowers, and a linear, longi- 

tudinally ribbed capsule. Occasional in disturbed 

wet places, reported from near sea level to 240 m 

elevation. A modern introduction, now pantropic in 

distribution. Lamoureux 3052; Whistler 3233, 

8961!. 

Ludwigia octovalvis (Sacq.) Raven 

[primrose willow] 

Tall, erect herb with alternate leaves, 4-parted 

flowers with yellow, notched petals, and a cylin- 

drical, many-seeded, longitudinally ribbed capsule. 

Common in wet, disturbed places, reported from 

near sea level to 900 m elevation. A modem intro- 

duction, now pantropic in distribution. Setchell 6, 

24, 361, 521 ; Eames T29; Garber 989; Lamoureux 

3053: Whistler 8929!. 

"Oxalis barrelieri L. 

vine 

OXALIDACEAE 

Erect herb with opposite or alternate, trifoliate 

leaves, long-stalked axillary inflorescences of pink, 

5-merous flowers, and a ribbed, cylindrical fruit 

with an acute tip. Occasional in disturbed places, 


duction, native to tropical America. (Whistler). 

Oxalis corniculata L. 

[wood sorrel] 'ii 

Low, often prostrate herb with alternate, longpetiolate, 

trifoliate leaves having obcordate leaflets, 

yellow, 5-merous flowers, and a capsule that bursts 

open to expel the seeds. Occasional in lawns, pastures, 

and other sunny, disturbed places, reported 

from near sea level to 550 m elevation. A Polynesian 

introduction, cosmopolitan in distribution. 

Setchell 323, 520; Garber 819, 946; Whistler 9036!. 

PASSIFLORACEAE 

Passiflora foetida L. 

[love-in-a-mist] pasio vao 

Hairy, herbaceous vine with alternate, palmately 

3-lobed leaves, coiled tendrils, showy white flowers, 

and a small, globose, edible fruit surrounded by 

branching calyx lobes. Common in disturbed 



1464, 895 1 !. 

*Passiflora maliformis L. 

Climbing vine with large, alternate, ovate to elliptic 

leaves, coiled tendrils, showy white and purple 

flowers, and a globose, hard-shelled fruit. Uncommon 

in secondary forest, perhaps an escape from 

cultivation, reported from near sea level to 300 m 

elevation. A modem introduction, native to tropical 

America. Whistler 3606,9152. 

PIPERACEAE 

"Macropiper puberulum Benth. 

'ava'ava aitu 

Shrub with alternate, cordate leaves, white, solitary 

male spikes, and solitary female spikes bearing 

numerous tiny red fruits. Occasional in lowland to 


elevation. Indigenous, ranging from Fiji and Futuna 

to Rapa. USEE s.n.; Whistler 2792. 

*Peperomia leptostachya Hooker & Am. 

Small terrestrial herb with puberulent opposite 

leaves and long terminal spikes of minute green 

flowers. Occasional in coastal forest, rarely in 


m or more elevation. Indigenous, widespread from 

Australia to Hawai'i. Setchell288; Whistler 2950. 

"Peperomia lonchophylla C. DC. 

Epiphytic herb with relatively large, alternate leaves 

over 2.7 times longer than wide and spikes (2 or 

more per axil) of minute green flowers. Occasional 

in lowland to cloud forest, reported from 40 to 1500 

m elevation. Endemic. Graeffe s.n.; Long 3049; 

Whistler 2985, 3615.

Peperomia pellucida (L,.) Kunth POLYGALACEAE 

Weak-stemmed herb with alternate, cordate to ovate 

leaves and terminal spikes of minute green flowers. 

Occasional in croplands and disturbed places, 


duction, native to tropical America. Yuncker 9397; 

I mnoureux 3075; Whistler 2756. 

Peperomia tutuilana Yuncker 

Small erect herb with alternate leaves and solitary 

axillary spikes (usually solitary in the upper axils) of 

minute green flowers. Occasional on rocks or trees 

in the coastal to lowland forest, reported from near 

sea level to 400 m elevation. Indigenous, also found 

in Tonga ('Eua). Reinecke 40a?; Garber 863; 

Yuncker 9386; Whistler 1274,2797,8590!, 9333!. 

Piper graeffei Warb. 

fue manogi; 'ava'avaaitu sosolo 

Climbing vine with alternate, palmately veined 

leaves, long, hanging unisexual spikes of minute 

green flowers, and tiny red fruits. Common to 

abundant in coastal to montane forest, reported from 

20 to 600 m elevation. Indigenous, also found on 

Futuna. Setchell 96, 463; Wilder 84; Garber 912; 

Christophersen 985, 1260, 3566; Diefenderfer 15; 

Meebold 16730; Mitchell 582; Yuncker 93 17, 9357; 

Spence 489!; Whistler 1422, 1470, 3122. 

PLANTAGINACEAE 

*Plantago lanceolata L. 

[narrow-leaved plantain] 

Rosette-forming herb with parallel-veined lanceolate 

leaves, narrow scapose spikes, and inconspicuous 

flowers with exserted stamens. Rare in disturbed 

places and perhaps not persisting in Samoa, 

reported only from the lowlands. A modern introduction, 

native to the Old World but now cosmopolitan. 

Rechinger 3702. 

"Plantago major L. 

[plantain] 

Rosette-forming herb with broad, parallel-veined 

leaves, a narrow scapose spike, and inconspicuous 

flowers with exserted stamens. Uncommon on 

roadsides and in other disturbed places, reported 


native to the Old World, but now cosmopolitan. 

Setchell 126: Whistler 3 19. 

Polygala paniculata L. 

pulunamulole 

Small herb with fragrant roots, alternate leaves, and 

tiny white flowers in terminal and axillary racemes. 

Occasional in disturbed places, reported from sea 

level to 700 m elevation. A modern introduction, 

native to tropical America. Setchell 16, 563; Eames 

T10; Wilder 10; Garber 795; Lamoureux 3070; 

Long 2599; Whistler 1439, 8910!. 

PORTULACACEAE 

*Portulaca lutea Sol. ex Forst. f. 

tamole 

Prostrate succulent herb with spirally arranged, 

glabrous leaves, large, showy yellow solitary 

flowers, and a capsule opening by the top splitting 

off. Occasional on coastal rocks, reported only from 

near sea level. Indigenous, ranging from New Caledonia 

to Pitcairn Island. Setchell52,414. 

*Portulaca oleracea L. 

[purslane] tamole 

Succulent prostrate to ascending herb with glabrous, 

spirally arranged leaves, yellow flowers in terminal 

clusters, and a capsule opening by the top splitting 

off. Common on rocky and sandy shores and often 

weedy, reported only from the lowlands. A Polyne- 

sian or early European introduction, now cosmopo- 

litan in distribution. Setchell 113; Garber 821, 970; 


*Portulaca quadrifida L. 

Tiny prostrate herb with opposite leaves, small 

yellow, 4-merous flowers, and a capsule opening by 

the top splitting off. Uncommon in sunny littoral 

habitats and occasionally weedy in villages, reported 

only from the lowlands. Probably a Polynesian 

introduction, pantropic in distribution. Garber 816, 


Portulaca samoensis Poelln. 

Prostrate, hairy herb with small, spirally arranged 

leaves, small yellow, 5-merous flowers and a cap- 

sule opening by the top splitting off. Occasional on 

coastal rocks, reported only near sea level. Indige- 

nous, ranging from New Guinea to Niue. Setchell 

232, 347, 357; Long 2598; Whistler 2912.

RHAMNACEAE 

Alphitonia zizyphoides (Spreng.) A. Gray 

toi 

Tall tree with alternate lanceolate leaves gray 

beneath, clusters of small white flowers, and purple 

globose fruits. Common in secondary forest and 



Southeast Asia to eastern Polynesia. Setchell 214, 

456; Wilder 94; Wisner 10, 10a, 104; Spence 465!; 

Whistler 2778,2938, 8603!, 9142. 

Colubrina asiatica (L.) Brongn. 

fisoa 

Spreading shrub with glossy alternate leaves, small 

white flowers, and green globose fruits. Occasional 

to common on the seaward margins of littoral forest, 

reported only near sea level. Indigenous, ranging 

from tropical East Africa to the Marquesas. Setchell 

3 10, 32 1; Wilder 22, 71, 75, 89; Mitchell 584; 

Yuncker 9303; Wisner 83; Whistler 1469, 8559!. 

RHIZOPHORACEAE 

*Uruguiera gymnorrhiza (L.) Lam. 

[Oriental mangrove] togo 

Large tree with glossy opposite leaves, a calyx of 

many linear lobes, orange petals, and a cigar-shaped 

fruit. Locally abundant in mangrove swamps, 

reported only near sea level. Indigenous, ranging 

from tropical Asia to Samoa. Setchell 3 1 ; Garber 

953; Yuncker 937 1; Bryan 963; Whistler 28 16. 

Crossostylis biflora Forst. 

saitamu 

Medium-sized tree with stilt roots, opposite leaves, 

and greenish flowers bearing many stamens. Occa- 

sional in lowland to cloud forest, reported from 200 


Tahiti to the Marquesas. USEE s.n.; Setchell 554; 

Bryan 93; Christophersen 1038, 1045, 3512; Mit- 

chell 575; Yuncker 9407; Wisner 152; Long 1963, 

1974; Spence 477!; Whistler 2700, 8520!, 9342!. 

*Rhizophora mangle L. 

[red mangrove] togo 

Small to medium-sized tree with stilt roots, glossy 

opposite leaves, small cream-colored flowers, and a 

long pendulous fruit. Common in estuaries and 

mangrove swamps, reported only near sea level. 

Indigenous, native to the New World tropics, New 

Caledonia, Fiji, Tonga, and Samoa. Setchell 98, 

309, 532; Christophersen 3479; Yuncker 9370; 

Whistler 2817, 3284. 

RUBIACEAE 

*Aidia cochinchinensis Lour. 

ola mea 


flowers in dense clusters, and small red fruits. 

Occasional in coastal to lowland forest, reported 


ranging from tropical Asia to western Polynesia. 

(Rechinger); Whistler 8459. 

Calycosia sessilis A. Gray 

Unbranched shrub with terminal, subsessile leaves, 

white flowers in a dense, flattened, sessile terminal 

head, and small red fruits. Occasional in montane to 


Endemic. Setchell 386; Christophersen 1 123 !; 


Canthium merrillii (Setchell) Christoph. 

ola sina 

Medium-sized tree with opposite leaves, white 

axillary flowers, and bluish, laterally compressed, 

subglobose fruits in axillary clusters. Occasional in 

coastal to montane forest, reported from 20 to 700 m 

elevation. Indigenous, also found in Tonga (the 

Niuas). Setchell 423, 442, 5 18; Diefenderfer 13; 

Long 1989; Whistler 3672. This should be 

transferred to the genus Psydrmc. 

Cyclophyllum barbatum (Forst, f.) Hall6 & 

Florence 

ola? 

Small tree with small opposite leaves, small white 

flowers, and a red, irregularly subglobose pyrene. 

Occasional to common in montane scrub and coastal 


Indigenous, ranging from Fiji to the Marquesas. 

Christophersen 1004, 3528; Whistler 2891, 

3665, 8050, 8458, 8656!, 8678!.

*Gardenia taitensis DC. "Hedyotis corymbosa (L.) Lam. 

[Tahitian gardenia] pua Samoa 

Small herb with opposite, narrowly elliptic leaves 

and thin, axillary peduncles bearing 2 or 3 tiny white 

flowers. Uncommon in disturbed places, reported 


native to Africa. Sykes 17; Whistler 5817. 

Shrub with glossy opposite leaves, large showy 

white flowers, and a green, ribbed capsule. Uncom- 

mon in coaslal forest and on coastal rocks, reported 

only ncar sca level but lormerly commonly culti- 

vated. I'robably a Polynesian introduction, ranging 

from Vanuatu to Hawai'i. USEE s.n.; Graeffe 227; 

Setchell 13 1, 236; Garber 95, 11 17; Amerson s.n.; 

Whistler 3855, 842 1 . 

Ceophila repens (L.) I. M. Johnston 

tono 

Prostrate herb with kidney-shaped leaves, small 

white flowers, and a red globose fruit. Occasional in 


level to 700 m elevation. Indigenous, pantropic in 

distribution. Setchell 41, 81; Garber 894; Wisner 1; 

Long 1957; Whistler 2943, 8639!. 

Cuettarda speciosa L. 

puapua 

Medium-si~ed tree with opposite obovate leaves, 

showy white, long-tubed, fragrant flowers, and a 

hrown, globose fruit. Common in littoral to lowland 


elevation. Indigenous, ranging from east Africa to 

the Marquesas. Whistler 9Ol 5!. 

Ixora samoensis A. Gray 

filofiloa 

Shrub with opposite leaves, long, tubular, white to 

pink flowers in 3s, and a small globose fruit. Occa- 

sional in coastal to cloud forest, reported from near 

sea level to 1200 m elevation. Endemic. Wilder & 

Swezey 58; Garber 835, 925; Christophersen 1005, 

1132!, 3481, 3535, 3549, 3554; Spence 509!; 

Whistler 2885, 3297,ii 3621, 3663, 8053, 8460, 

8633!, 8682!. 

Morinda citrifolia L. 

[Indian mulberry] nonu 

Shrub or small tree with large, glossy dark green, 

opposite leaves, small white flowers, and a large 

yellowish, irregularly shaped, fleshy, compound 

fruit. Common in coastal forest and disturbed 


elevation. Probably a Polynesian introduction, 

ranging from India to Hawai'i. Setchell 34, 572; 

Whistler 1440,3 120,9024!, 9144. 

Gynochtodes epiphytica (Rechinger) A. C. Smith Morinda myrtifolia A. Gray 

& S. Darwin 

Woody climbing vine with opposite, round-tipped 

Woody climber with opposite leaves, small axillary leaves, small white flowers, and a subglobose, 

flowers, and greenish, marble-like fruits. ccasional compound fruit. Occasional in coastal to lowland 

to common in coastal to lowland forest, reported forest, reported from near sea level to 200 m elevafrom 

20 to 400 m elevation. Indigenous, also found tion. Indigenous, ranging from New Caledonia to 

in Fiji, Niue, and Tonga. Whistler 2874, 3618, the Marquesas. Whistler 3670,8541!. 

8482.9 150. 

Mussaenda raiateensis J. W. Moore 

Hedyotis biflora (L.) Lam. 

aloalo vao 

Small herb with opposite leaves, tiny white flowers, Terrestrial or epiphytic shrub with opposite leaves, 

and a papery, subglobose capsule. Occasional on yellow flowers subtended by a white, leaf-like bract, 

coastal rocks, rarely inland and weedy, reported and a brown, ovoid hit. Occasional in lowland to 

from near sea level to 350 m elevation. Indigenous, montane forest, reported from near sea level to 700 

ranging from Mauritius to the Cook Islands. Lister m elevation. Indigenous, ranging from Vanuatu to 

s.n.; Setchell 120, 320; Garber 969; Diefenderfer 4; the Society Islands. Veitch s.n.; Reinecke 475; 

Whistler 2922, 8399, 8692!. Wilder s.n.; Garber 791; Bryan 981!; Yuncker 9355; 

Long 2601; Whistler 8536!.

Neonauclea forsteri (Seem.ex Havil.) Merr. 

afa 

Large tree with opposite leaves, conspicuous oval 

stipules on the terminal bud, and white flowers in a 

terminal head. Common in8 lowland to cloud forest, 


ranging from the Solomon Islands to Tahiti. Lister 

sen.; Setchell 565; Christophersen 181 1; Wisner 86; 

Spence 501 !?; Whistler 1284,2956,3841,9016!. 

Psychotria forsteriana A. Gray 

Small tree with opposite leaves, many tiny white 

flowers, and red subglobose fruits 2-lobed when dry. 

Common in lowland to montane forest, reported 


nous, also found in Fiji. Graeffe 37b, s.n.; Garber 

841; Meebold 8253; Long 3061; Whistler 2730, 

8495, 8990!, 9338!. 

Psychotria insularum A. Gray 

matalafi 

Small tree with opposite leaves, small white flowers, 

and glossy, globose, red fruits. Common in 

coastal to cloud forest, reported from near sea level 

to 1400 m elevation. Indigenous, also found in 

Tonga, Futuna, 'Uvea, Rotuma, and Niue. Graeffe 

s.n.; Setchell 82, 256, 572; Wilder 17, 47, 78; Garber 

792, 932; Bryan 85; Diefenderfer 11; Christophersen 

1009, 1043, 1820, 1826, 3485, 3521, 3550, 

3557, 3558; Mitchell 506, 517; Yuncker 9350; 

Wisner 61 (n.~.), 67, 130; Lamoureux 3007, 3012; 

Long 1966, 1983, 1990, 1994, 3065a, 3068, 3086, 

3102; Spence 484!; Whistler 1437, 2737, 2767!, 

3542, 3638!, 3666,3667, 8634!. 

"Psychotria pacifica K. Schum. 

Small tree with opposite leaves, panicles termi- 

nating in umbels, white sessile flowers, and red 

subglobose fruits. Common in montane to cloud 


Endemic. Graeffe 1383. A questionable record. 

Sarcopygme pacifica (Reinecke) Setchell & 

Christoph. 

u'unu 

Medium-sized, scarcely branched tree with large 

opposite leaves, long showy white flowers in heads, 

and a large pendulous compound fruit. Common in 

lowland to cloud forest, reported from 50 to 1400 m 

elevation. Endemic. Setchell 362, 419, 450; Bryan 

88; Christophersen 1 l36!, 1264, 1807!, 1808!, 

1809!, 1810!, 3586!, 3587!, 3588!; Mitchell 580; 

Long 3064; Whistler 2732, 8742!. 

Spermacoce assurgens Ruiz and Pavon 

Erect herb with opposite, lanceolate to narrowly 

elliptic leaves, unwinged stems, and tiny white 

flowers in sessile, axillary clusters. Common in 

croplands and disturbed places, reported only from 


tropical America. Lamoureux 307 1 ; Fosberg 54926; 

Whistler 2755, 9009!. 

Spermacoce mauritiana Gideon 

Small herb with smooth stems, opposite, mostly 

glabrous leaves, and tiny white flowers in sessile, 

axillary clusters. Uncommon in lawns and other 

sunny, disturbed places, reported only from the 


America. Whistler 3 17,2867, 573 1, 8524!. 

Tarenna sambucina (Forst. f.) Durand in Drake 

ma'anunu 

Small tree with opposite leaves, dense inflore- 

scences of white flowers bearing exserted stamens, 

and small globose fruits. Occasional in open forest 

and disturbed places, reported from near sea level to 


Vanuatu to the Marquesas. Rechinger 3791; Set- 

chell 83, 528; Bryan 983! (ns.); Meebold 16871; 

Whistler 1479,2925, 3775. 

RUTACEAE 

Acronychia heterophylla A. Gray 

Shrub or small tree with opposite, simple or trifo- 

liate leaves, small white flowers, and a 4-lobed fruit 

containing 4 shiny black seeds. Occasional in 


m elevation. This species should be transferred to 

the genus Melicope. Endemic. USEE s.n.; Setchell 

329, 429; Wilder 34; Garber 909; Christophersen 

1137!; Yuncker 9426!?. 

'Acronychia richii A. Gray 

Shrub or small tree with simple leaves, tiny white 

flowers, and a 4-lobed splitting fruit containing 4 

shiny black seeds. Uncommon in montane scrub 

and cloud forest, reported from 500 to 1700 m

elevation. This species should be transferred to the 

gcnus Melicope. Endemic. Christophersen 1052, 

1066, 3532, 3556; Whistler 3664. 

*Citrus macroptera Montr 

moli u'u 

Small to medium-sized thorny trees, alternate 

leaves, white 5-merous flowers, and small sour 

oranges. Uncommon in coastal to lowland forest, 

formerly cultivated but now apparently disappear- 

ing. A Polynesian introduction, widespread in the 

Pacific. Setchell 136: Yuncker 9429. 

Euodia hortensis Forst. 

LIS~ 

Shrub with opposite, simple or trifoliate, aromatic 

leaves, small white flowers in panicles, and 4-lobed 

fruits splitting to release the four shiny black seeds. 

Uncommon in disturbed places, mostly as a relict of 

cultivation, and perhaps not truly naturalized, 


Polynesian introduction, native to Melanesia. USEE 

s.n. (n.s.); Veitch s.n.; Garber 931, 960; Wilder 83; 

Christophersen 96 1 ; Whistler 1475,2896, 8654!. 

Euodia samoensis Christoph. 

so'opine 

Small tree with opposite, trifoliate leaves, small 

white flowers, and a 4-lobed h it splitting to release 

four shiny black seeds. Occasional in lowland to 


tion. Endemic. Whistler 8660!. 

Micromelum minutum (Forst. f.) Seem. 

talafalu 


leaves, small white flowers, and red fruits. Occa- 

sional in coastal to foothill forest, reported from 


ranging from the Philippines and Australia to Niue. 

Setchell 257; Christophersen 1012; Meebold 26569; 

Wisner 94; Whistler 2944, 3607, 8461, 8553!, 

8560!. 9 167. 

SAPINDACEAE 

Allophylus timoriensis (DC.) B1. 

Small to medium-sized tree with trifoliate leaves, 

small white flowers in racemes, and red, globose 

fruits. Occasional in littoral to montane forest, 


Indigenous, ranging from Malaysia to the 

Marquesas. Rechinger 3730; Yuncker 9365; 

Whistler 1273, 8070!, 8571 !. 

Arytera brackenridgei (A. Gray) Radlk. 

taputo'i 

Medium-sized tree with alternate, pinnately 

compound leaves, lanceolate leaflets, and tiny white 

flowers. Occasional to common in lowland forest, 


Indigenous, also found in Fiji. Setchell 517; Chri- 

stophersen 1 142!; Spence 5 l7!; Whistler 8073, 

8533!. 

Elattostachys falcata (A.Gray)Radlk. 

taputo'i 

Medium-sized to large tree with pinnately com- 

pound leaves, inconspicuous flowers with red 

stamens, and small 3-lobed capsules. Occasional in 

lowland to cloud forest, reported from 30 to 1450 m 

elevation. Indigenous, also found in Vanuatu, Fiji, 

Tonga, and Niue. Whistler 3650, 8068. 

*Guioa rhoifolia (A. Gray) Radlk. 

Medium-sized tree with pinnately compound leaves 

having small domatia at the base of the lower sur- 

face of the subfalcateti leaflets, and tiny apetalous 

flowers. Rare in lowland to montane forest, repor- 

ted from 250 to 750 m elevation. Indigenous, also 

found in Fiji. Christophersen 3483; Whistler 9065. 

*Pometia pinnata Forst. 

tava 

Large buttressed tree with alternate, pinnately 

compound leaves, dense panicles of apetalous 

flowers, and edible, large, subglobose fruits. Com- 

mon to abundant in lowland to lower montane 


genous, ranging from Ceylon to Samoa. Whistler 

9177. 

Sapindus vitiensis A. Gray 


leaves, winged petioles (at least in saplings), 

apetalous flowers in a large panicle, and 2-lobed or 

subglobose, red fruits. Occasional in coastal forest, 

reported only in the lowlands. Indigenous, also 

found in Fiji and Tonga. Whistler 8066!, 9056!.

SAPOTACEAE 

Manilkara dissecta (L. f.) Dubard 

pan i 

Medium-sized tree with glossy, alternate, obovate 

leaves and solitary axillary 6-merous flowers. Rare 

in coastal forest, reported from near sea level to 200 

m elevation. Indigenous, ranging from Vanuatu to 

Samoa. Guest sen.; Whistler 8466, 8679!. 

Palaquium stehlinii Christoph. 

nasu (Western Samoa) 

Large tree with ovate leaves, milky sap, small 

flowers with at least twice as many stamens as 

petals, and a large, single-seeded, curved-oblong 

fruit. Occasional in lowland to montane forest, 


Whistler 8655!, 8747!, 9339!. 

Ylanchonella garberi Christoph. 

'ala'a 

Medium-sized to large tree with milky sap, lanceo- 

late leaves, small axillary flowers, and a subglobose 

fruit red to black at maturity and containing one or 

more shiny black seeds. Occasional in coastal to 

foothill forest, reported from near sea level to 500 m 

elevation. Indigenous, ranging from Fiji to Niue. 

Setchell 291; Wisner 46, 47, 48; Whistler 2690!, 

2864,3298,3302,3349,3537,8504,8673!, 8718!. 

Planchonella grayana St. John 

'ala'a 

Large tree with milky sap, elliptic leaves, small 

axillary flowers, and a flesh-colored, globose fruit 

containing several shiny black seeds. Uncommon in 

littoral and coastal forest, reported from near sea 


Fiji to the Tuamotu Islands (Makatea). Whistler 

2945. 8430. 

Planchonella samoensis H. J. Lam ex Christoph. 

mamalava 

Large tree with milky sap, large alternate leaves, 

small axillary flowers, and an ellipsoid, red to 

yellow fruit containing a single shiny black, com- 

pressed-fusiform seed. Common to abundant in 


m elevation. Indigenous, also found in Tonga (the 

Niuas) and Niue. Setchell 259; Wisner 80; Whistler 

1442,2875,2963,3627. 

SCROPHULARIACEAE 

"Limnophila fragrans (Forst, f.) Seem. 

tamole vai 

Prostrate herb with fragrant, opposite leaves and 

small sympetalous, white to pale-blue flowers. Rare 

in wet places such as taro patches, streambeds, and 

crater lake margins, reported from near sea level to 

650 m elevation. Indigenous, ranging from Austra- 

lia and Melanesia to the Society Islands. USEE s.n. 

(n.s.); Setchell370; Whistler 8044, 9034. 

"Limnophila rugosa (Roth) Merr. 

Erect herb with opposite, oblong leaves, axillary, 

sessile flowers, and a lavender and yellow campa- 

nulate corolla. Uncommon in wet disturbed areas, 


duction, native to tropical Asia. Whistler 3228. 

*Lindernia antipoda (L.) Alston in Trimen 

Small prostrate herb with opposite sessile leaves, 

small lavender flowers oppositely arranged in 

terminal racemes, and linear capsules. Uncommon 

or rare in disturbed places, reported only from the 



Lindernia crustacea (L.) F. Muell. 

Small herb with opposite leaves, purple-blue 

flowers, and a tiny, many-seeded capsule. Occa- 

sional in wet places such as streambeds, reported 


pantropic in distribution. Setchell 76, 327; Whistler 

277O!, 8640!. 

SOLANACEAE 

*Capsicum frutescens L. 

[chili pepper] polo feu 

Subshrub with alternate leaves, cream-colored, 

rotate flowers reflexed at anthesis, and a bright red, 

cylindrical berry. Occasional in disturbed places, 

reported only from the lowlands. A modem introduction, 

native to tropical America. Setchell 5; 

Garber 802; Diefenderfer 19; Christophersen 10 14; 

Whistler 9 162.

"Physalis angulata L. 

1 wild cape gooseberry] vivao 

Erect herb with alternate leaves, pale yellow, rotate 

flowers, and a bladder-like calyx enclosing a small 

edible berry. Common in croplands and other dis- 

turbed places. Apparently a Polynesian introdu- 

ction, native to tropical America. USEE s.n. (n.s.); 

Lamoureux 3059; Whistler 32 1, 1468, 8437. 

*Solanurn americanum Mill. 

[black nightshade] mgalo 

lirect herb with alternate leaves, small white flowers 

bearing yellow stamens and reflexed petals, and a 

shiny black, globose berry. Occasional in disturbed 


vation. Apparently a Polynesian introduction, or 

perhaps indigenous, native to tropical or subtropical 

America. USEE sen.; Wilder 45; Whistler 8436. 

*Solanurn viride Forst. f. ex Spreng. 

polo 

Shrub with alternate, mostly pubescent leaves, 

many-flowered inflorescences of white flowers, and 

small red, tomato-like fruits. Rare in coastal forest, 

reported only from the lowlands. Indigenous or 

possibly a naturalized Polynesian introduction, but 

now rare. Wilder 20?; Meebold 8236?; Whistler 

8067. 

Kleinhovia hospita L. 

f'u'afu'a 

STERCULIACEAE 

Medium-sized tree with alternate cordate leaves, 

pink tlowers, and an inflated capsule. Common in 

disturbed places and secondary forest, reported from 


ranging from tropical Asia to western Polynesia. 

Setchell 373; Mitchell 537; Yuncker 9358; Wisner 

94; Whistler 901 7!. 

"Melochia aristata A. Gray 

ma'o 

Shrub with alternate cordate leaves, pink flowers, 

and a small ovoid capsule. Occasional in disturbed 

places and secondary forest, reported from near sea 

level to 700 m elevation. Indigenous, also found in 

Tonga and the Tuamotu Islands. Setchell 5 12; 


Sterculia fanaiho Setchell 

fana'io 

Medium-sized tree with alternate leaves having the 

blade angled to the petiole, small reddish flowers, 

and a large reddish-brown capsule containing a few 

large seeds. Occasional in coastal to montane forest, 


Indigenous, also found in Tonga, Niue, Rotuma, and 

Futuna. Setchell 103, 147, 426, 436, 577; 

Christophersen 1 134!. 

THEACEAE 

Eurya pickeringii A. Gray 

Shrub with toothed, alternate leaves, small cream- 

colored flowers, and a subglobose fruit. Occasional 

in montane scrub to cloud forest, reported from 100 

to 1700 m elevation. This identification needs veri- 

fication, since several questionable species names 

have been used in Samoa. Indigenous, also found 

on Rapa. USEE s.n. (n.s.); Christophersen 1049, 

1069, 3510, 35 13, 3530; Whistler 271 1, 2769!, 

3347,3350,8688,9004!. 

THYMELAEACEAE 

*Phaleria disperma (Forst. f.) Baill. 

suni 

Shrub or small tree with opposite leaves, fragrant, 

long, white, tubular flowers, and red subglobose 

fruits. Occasional in littoral and coastal forest, 


Indigenous, also found in Fiji, Tonga, and the Home 

Islands. Whistler 2848, 3856, 8434. 

Phaleria glabra (Turrill) Domke 

suni vao 

Shrub with opposite leaves, long tubular white flowers, 

and red hits. Occasional in coastal to montane 


Indigenous, also found in Fiji. Reinecke 171; 

Christophersen 1001; Wisner 1 19; Kuruc 1 18; 

Whistler 2866, 3608, 3644, 8478, 8653!. 

Wikstroemia foetida (L. f.) A. Gray 

Shrub with opposite leaves, small yellow flowers in 

few-flowered clusters, and a fleshy orange, subglobose 

fruit. Occasional in sunny disturbed forest 

and scrub, reported from 75 to 1530 m elevation. 

Indigenous, ranging from Fiji to Tahiti. Setchell

331; Garber 786, 878; Christophersen 3555; 

Yuncker 9345, 94 18; Sledge s.n.; Whistler 2907, 

2928,871 11, 8999!. 

TILIACEAE 

Grewia crenata (Forst.) Schinz & Guillaumin 

fauui 

Small tree with alternate leaves, white 5-merous 

flowers with many stamens, and a 2- to 4-lobed 

fruit. Uncommon in coastal forest, reported only 

from the lowlands. Indigenous, ranging from New 

Caledonia to the Society Islands. Setchell 149; 

Bryan 969, 1015; Meebold 8212, 21350, 26579; 

Spence 494; Whistler 1451, 1471,2839. 

Trichospermum richii (A. Gray) Seem. 

ma'o sina 

Medium-sized tree with alternate leaves having a 

pair of basal glands, stellate pubescent, white 

flowers with many stamens, and a flattened, winged, 

obovate fruit. Occasional in lowland to montane 


Indigenous, also found in Fiji. Christophersen 3570; 

Meebold 2 1350; Whistler 2777,2959. 

*Triumfetta procumbens Forst. f. 

mautofu tai 

Prostrate shrub with alternate leaves, showy yellow 

flowers having numerous stamens, and a bur-like 

fruit. Uncommon on sandy beaches, reported only 

from near sea level. Indigenous, ranging from 

Malaysia to the Tuamotu Islands. Setchell 409; 


"Triumfetta rhomboidea Jacq. 


Shrub with alternate, palmately lobed leaves, small, 

sessile, axillary, yellow flowers bearing many sta- 

mens, and a small, globose, bur-like fruit. Occa- 

sional to common in disturbed places, reported only 

in the lowlands. A modem introduction, native to 

the Old World tropics. Setchell33 8; Wilder 37. 

Trema cannabina Lour. 

magele 

ULMACEAE 

Small tree with coarse lanceolate leaves, tiny green- 

ish white flowers, and dense clusters of drupes. 

Occasional in disturbed forest and clearings, 


Indigenous, ranging from India to Niue and perhaps 

farther eastward. Setchell 263; Wilder 25; Garber 

902; Wisner 18; Whistler 3 1 15, 8982!. 

URTICACEAE 

*Boehmeria virgata (Forst. f.) Guillemin 

Small tree with coarse, ovate to elliptic, mostly 

glabrous, serrate to crenate leaves, and tiny incon- 

spicuous flowers in long spikes. Uncommon in 

disturbed forests and clearings, reported from 200 to 


Vanuatu and the Caroline Islands to the Marquesas. 

Christophersen 987; Whistler 9080. 

Cypholophus macrocephalus Wedd. 

faupata 

Shrub with ovate, rugose leaves, tiny greenish 

flowers in dense, sessile axillary clusters, and tiny 

orange drupes. Occasional in montane forest, 


ranging from Fiji to the Society Islands. Setchell 

44, 79; Bryan 80; Mitchell 576; Yuncker 9435!; 


Dendrocnide harveyi (Seem.) Chew 

[stinging-nettle tree] salato 

Medium-sized to large, soft-wooded tree with ovate 

leaves 3-5-veined from the base, stinging hairs, tiny 

green flowers, and tiny green achenes. Rare in low- 

land to montane forest on Tutuila, reported from 


ranging from Fiji to Niue. Christophersen 3583!; 

Whistler 8065, 9169. 

*Elatostema grandifolium Reinecke 

Large herb with large alternate leaves having crenate 

margins, and tiny green to white flowers. Occa- 

sional to common in montane forest, reported from 

400 to 890 m elevation. Endemic. Graeffe 1334. 

*Elatostema samoense Reinecke 

Small herb with alternate leaves, serrate leaf margins, 

variable appearance, and tiny greenish flowers. 

Rare in montane forest, reported from 550 to 700 m 

elevation. Endemic. Reinecke 503.

*Etatosterna scabriusculum Setchell 

Small herb with alternate leaves, long stipules, and 

tiny green flowers. Uncommon in montane forest, 


Setchell375; Yuncker 941 0; Long 1961, 1963. 

*Elatostema tutuilense Whistler 

Small herb with alternate leaves, deeply pinnately 

toothed leaf margins, and tiny greenish flowers. 

Rare in montane forest, no elevation reported (but 

over 200 m). Endemic to Tutuila (and collected 

only once). Reinecke 590. 

*Laportea interrupta (L.) Chew 

ogoogo 

Erect herb with toothed, ovate, alternate leaves, 

mildly stinging hairs, and a long, loose, axillary 

inflorescence of greenish, apetalous flowers. Un- 

common in disturbed places, reported only from the 

lowlands. A Polynesian introduction, possibly 

native to Southeast Asia. Setchell 145, 533; 


Leucosyke corymbulosa (Wedd.) Wedd. 

'ala'alatoa 

Small tree with alternate, palmately veined leaves 

gray on the lower surface, tiny greenish flowers, and 

a white fleshy compound fruit. Uncommon in low- 

land forest, reported from 100 to 250 m elevation. 

Indigenous, ranging from Vanuatu to the Cook 

Islands. Setchell 258; Wilder 86; Garber 809, 922; 

Yuncker 9340, 94 12; Long 1981, 3 104; Spence 

5 lo!; Whistler 2689!, 2890, 8608!, 8980!. 

*Maoutia australis Wedd. 

Small tree with alternate serrate leaves white 

beneath, and tiny green flowers in axillary inflore- 

scences. Uncommon in lowland to montane forest, 


ranging from Fiji to the Society Islands. Reinecke 

522 ("Tutuila and the Manu'a Islands!"). This 

record is questionable. 

Pilea microphylla (L.) Liebm 

Small prostrate, somewhat succulent herb with tiny 

leaves and greenish, inconspicuous flowers. Occa- 

sional in sunny disturbed areas and especially on 

rock walls, reported only from the lowlands. A 


Diefenderfer 24; Yuncker 9373; Long 3096; 

Whistler 2759, 9035!. 

Pipturus argenteus (Forst. f.) Wedd. 

soga; fau soga 

Small tree with alternate leaves gray beneath, tiny 

flowers in branching inflorescences, and white, 

fleshy compound fruits. Common in disturbed 

places and early secondary forest, reported from 


ranging from Vanuatu to the Marquesas. Reinecke 

506; Setchell 85, 122, 544; Bryan 974; Garber 813; 

Wilder 90 (n.s.); Christophersen 1073; Mitchell 565; 

Meebold 26589; Yuncker 9375; McKee 2941; 

Comstock 2; Wisner 59; Long 3095, 3096, 3103; 

Spence 479!; Whistler 2861, 8572!, 9145. 

Procris pedunculata (Forst.) Wedd. 

fua lole 

Terrestrial or epiphytic herb with tiny white flowers 

and red, strawberry-like aggregate fruits. Occasional 

in littoral to montane forest, reported from 


ranging from Malaysia to the Marquesas. Graeffe 

s.n.; Setchell 7, 212, 513; Garber 859, 869, 934; 

Bryan 932; Christophersen 986; Mitchell 573; 

Yuncker 9405; Whistler 2704, 2725, 2768, 8473, 

8996!, 91 15. 

VERBENACEAE 

Clerodendrum chinense (Osbeck) Mabberly 

losa Honolulu 

Large shrub with opposite, cordate leaves and 

showy white, many-petaled flowers in dense 

clusters. Occasional in plantations and along 

roadsides, reported only from the lowlands. A 

modern introduction, native to tropical Asia. 

Whistler 8898, 8967!. 

Clerodendrum inerme (L.) Gaertn. 

aloalo tai 

Sprawling shrub with opposite leaves and showy 

white flowers having exserted stamens. Common in 

the littoral strand and sometimes as a scandent shrub 

in littoral forest, reported only in coastal areas. Indigenous, 

ranging from tropical Asia to Niue. Setchell 

97, 292; Wilder 74; Garber 963; Diefenderfer

14; Yuncker 9369; Long 2595; Whistler 2836, 

9O27!. 

Faradaya amicorum Seem. 

mamalupe 

I-ligh-climbing vine with opposite leaves, white 

tubular flowers having exserted stamens, and a large 

red, curved-ellipsoid fruit. Common in lowland to 


Indigenous, also found in Tonga. Lister s.n.; 

Kuntze 23001, s.n.; Seal s.n.; Setchell 64, 539; 

Garber 913; Diefenderfer 1 (n.s.), 12, 28 (n.s.); 

Christophersen 990, 3478; Bryan 956; Mitchell 504; 

Meebold 8230 (n.s.), 16485, 21356 (n.s.), 26582; 

Yuncker 9301, 9387; Wisner 53, 128; McKee 2945; 

Wisner 69; Long 31 16; Spence 490!; Whistler 1482, 

2951, 8641!, 9163. 

Premna serratifolia L. 

aloalo 

Shrub or small tree with broad leaves marked with 

leaf miners, tiny greenish flowers, and a small black, 

globose fruit having a persistent calyx. Common in 

littoral strand and scrub vegetation, rarely inland, 


Indigenous, ranging from tropical Asia to the 

Marquesas. Setchell 155, 564; Garber 789, 901; 

Comstock s.n., 5, 3 la; Meebold 8239; Long s.n.; 

Whistler 2774, 9100!. 

*Stachytarpheta jamaicensis Vahl 

[Jamaica vervain] 

1,ow shrub with glabrous, opposite leaves and 

lavendcr sympetalous flowers embedded in long, 

thickened spikes. Uncommon on roadsides and 

other disturbed places, reported only from the low- 

lands. A modern introduction, native to tropical 

America. Wisner 1 16; Lamoureux 3054; Whistler 

3 15, 8439. 

Stachytarpheta urticaefolia (Salisb.) Sims 

mautofu [blue rat's-tail] 

Coarse shrub with opposite, rugose, toothed leaves 

and long, narrow, terminal spikes of purple, sympetalous 

flowers. Common in disturbed places, reported 



23, 558; Lamoureux 3055; Whistler 8973!. 

*Vitex trifolia L. 

namulega 

Small tree with opposite, palmately compound 

leaves gray beneath, tiny purple flowers, and black, 

globose fruits. Occasional in the littoral strand, and 

sometimes cultivated, reported only from the lowlands. 

Indigenous, ranging from tropical East Africa 

to the Marquesas. Setchell53 1 ; Wilder 48; Whistler 

1450,8403. 

MONOCOTYLEDONAE 

AGAVACEAE 

Cordyline fruticosa (L.) Chev. 

[ti plant] ti; lau ti 

Shrub with parallel-veined leaves, pink flowers in 

branching panicles, and bright red berries. Common 

in cultivation and naturalized in lowland to cloud 

forest, reported from near sea level to over 1100 m 

elevation. A Polynesian introduction, widespread 

from tropical Asia to Hawai'i. Setchell 75, 12 1 137, 

138, 139, 140, 141, 142, 332; Garber 808, 870; 

Bryan 74, 1017, 1018; Wisner 55, 60; Long 1965, 

3074; Whistler 1433,2724,2849, 8552. 

AMARYLLIDACEAE 

*Zephryanthes rosea (Spreng.) Lindl. 

[zephyr lily] 

Stemless herb with strap-shaped leaves arising from 

an underground bulb, and showy, pink, scapose 

flowers. Occasional in lawns, reported only from 


tropical America. (Setchell). 

ARACEAE 

Amorphophallus paeoniifolius (Dennst.) 

Nicholson 

teve 

Stemless herb with large deeply divided leaves 

arising from an underground rhizome, and a large, 

purple, bell-shaped spathe surrounding the thick 

cylindrical spadix. Uncommon in secondary forest 

and plantations, reported only from the lowlands. A 

Polynesian introduction, widespread from tropical 

Asia to Hawai'i. Diefenderfer 17; Whistler 8579!, 

9332!.

Epipremnum pinnaturn (L.) Engl. 

Iligh-climbing vine with fenestrate leaves and 

flowers in a fleshy cylindrical spadix surrounded by 

a spathe. Common in lowland to foothill forest, 


Indigenous, widespread in the Pacific to eastern 

Polynesia. Setchell 203; Diefenderfer 2?, 51 

Rhapidophora graeffei Engl. 

fue laufao 

High-climbing vine with entire leaves and an inflo- 

rescence of flowers in a fleshy cylindrical spadix 

surrounded by a spadix. Common in lowland to 


elevation. Endemic. Setchell 86; Meebold 8209; 

Whistler 361 514, 9148, 9354!. 

Cocos nucifera L. 

[coconut] niu 

ARECACEAE 

Large palm with pinnately divided fronds, thick 

panicles of unisexual flowers, and large fruits with a 

thick fibrous husk. Common in coastal areas, most- 

ly cultivated, reported from near sea level to 500 m 

elevation. Probably indigenous, originating some- 

where in the Old World tropics, but now pantropic 

in distribution. (Whistler). 

*Canna indica L. 

[canna] fanamanu 

CANNACEAE 

Tall erect herb with large ovate to lanceolate leaves, 

showy red flowers, and a soft-spiny fruit containing 

pea-like seeds. Occasional in disturbed places, 

especially in wet areas, reported only from the 


America. Christophersen 1003; Whistler 2786. 

*Aneilima vitiense Seem 

COMMELINACEAE 

Herb with alternate lanceolate leaves, white 3-mer- 

ous flowers, and a fuzzy fruit. Occasional in coastal 

to montane forest, reported from near sea level to 

750 m elevation. Indigenous, also found in Fiji and 

Tonga. (Whistler). 

*Commelina benghalensis L. 

Weak-stemmed herb with alternate, ovate leaves and 

blue, 3-merous flowers enclosed within a folded 

bract. Uncommon in disturbed places, reported only 

from the lowlands. A modem introduction, native to 

tropical Asia. Yuncker 9402; Whistler 8036. 

Commelina diffusa Burm. f. 

mau'utoga 

Weak-stemmed herb with alternate, lanceolate 

leaves and blue 3-merous flowers enclosed within a 

folded bract. Common in disturbed places, especia- 

lly in wet areas, reported from near sea level to 650 

tn elevation. A Polynesian introduction, native to 

the Old World tropics. Setchell2 16, 3 18. 

Cyperus compressus L. 

CYPERACEAE 

A small sedge with a terminal cluster of green, 

flattened-lanceolate spikelets subtended by several 

linear bracts. Occasional in disturbed areas, reported 


now .pantropic in distribution. Eames T1; Whistler 

8962!. 

"Cyperus rotundus L. 

[nut sedge] 

Sedge with swollen tubers, creeping rhizome, basal 

leaves, and terminal, flattened, red-brown spikelets 

on several spreading rays. Apparently uncommon in 


A modern introduction, native to the Old World 

tropics and now cosmopolitan. Graeffe 1227. 

*Cyperus stoloniferus Retz. 

Sedge with thick bulb-like tubers, basal linear 

leaves, and terminal clusters of brown, flattened- 

cylindrical spikelets. Common on rocky coasts, 

reported only on the coast. Indigenous, native to the 

Old World Tropics. Whistler 8349. 

*Eleocharis dulcis (Burm. f.) Hens. 

[water chestnut] 'utu'utu 

Leafless sedge with tall cylindrical stems bearing a 

brown, terminal, cylindrical inflorescence as wide as 

the stem. Locally abundant in coastal and montane 

marshes, reported from near sea level to 900 m 

elevation. Indigenous, ranging from West Africa to 

Samoa. Whistler 8895.

Fimbristylis autumnalis (L.) Roemer & Schultes 

Small to medium-sized sedge with three short, ter- 

minal bracts subtending clusters of small, brown, 

flntlcned-ellipsoid spikelets. Occasional in disturbed 

wet places, reported only from the lowlands. A 

~notlcrn introduction, native to tropical Asia. 

I.amourcux 3088; Whistler 323, 1285, 8035, 8963!. 

Fimbristylis cymosa R. Br. 

Small clump-forming sedge with linear leaves and 

subglobose clusters of spikelets born atop a short 

scape. Locally abundant on rocky shores, reported 

only from coastal areas. Indigenous, ranging from 

Malaysia to Hawai'i. USEE s.n.; Garber 807; 

Wisner 37; Lamoureux 3087, 3089; Whistler 1429, 

2851, 8566!. 

Fimbristylis dichotoma (L.) Vahl 

Clump-forming, thin-stemmed sedge with basal 

leaves and long stalks bearing brown ellipsoid 

spikelets in terminal corymbs. Common in disturbed 


elevation. A Polynesian, or perhaps a modern 

introduction, now pantropic in distribution. USEE 

s.n. (n.~.); Eames T7; Garber 887; Christophersen 

993: Whistler 8966!. 

Kyllinga brevifolia Rottb. 

Small sedge with creeping rhizomes, long, narrow 

stems, and small green, globose, head-like clusters 

of spikelets subtended by 3 linear, leaf-like bracts. 

Common in lawns and other sunny, disturbed 

places, reported from near sea level to 700 m eleva- 

tion. A modern introduction, now pantropic in 

distribution. Setchell 583; Whistler 2978, 8965!. 

Kyllinga nemoralis (Forst.) Dandy ex Hutch. & 

Dalziel 

Small sedge with creeping rhizomes and small 

white, globose, head-like clusters of spikelets sub- 

tended by three leaf-like bracts. Common in lawns 

and other sunny, disturbed places, reported only 


native to the Old World tropics. Setchell 11 1, 51 1; 

Eames T4; Bryan 79; Garber 866, 888; Whistler 

9106!. 

*Macharha falcata (Nees) Koyama 

Large sedge with basal, lanceolate leaves and an 

erect panicle of brown spikelets. Uncommon in 

montane scrub and along streams in montane to 

cloud forest, reported from 450 to 1 I00 m elevation. 

Indigenous, ranging from the Philippines and 

Malaysia to Samoa. Setchell 379; Christophersen 

1023,3490; Whistler 2972,3668. 

Mapania macrocephala (Gaud.) K. Schum. 

Large sedge with basal leaves and a large, brown, 

irregularly lobed subglobose inflorescence. Occa- 



ranging from the Philippines to Samoa, but not 

found in Fiji. Graeffe 1222; Setchell 397; Christo- 

phersen 1 122!, 3508, 3584!; Mitchell 520; Whistler 

3653,5727. 

"Mapania vitiensis (Uittien) T. Koyama 

Large sedge with flat, strap-like basal and cauline 

leaves and terminal corymbs of small ovoid spike- 

lets. Rare in montane scrub, reported from 500 m 

elevation. Indigenous, also found in Fiji. Christo- 

phersen 1209,3484. 

Mariscus cyperinus (Retz.) Vahl 

Erect sedge with basal leaves with green spikelets in 

a dense, head-like cluster surrounded by leaf-like 

bracts. Uncommon to occasional in disturbed 


vation. Indigenous or perhaps a Polynesian 

introduction, originally native to tropical Asia. 

Setchell 582; Eames T15; Christophersen 992; 

Meebold 8216; Yuncker 9438; Whistler 8628!. 

Mariscus javanicus (Houtt.) Men. & Metcalfe 

selesele 

Large sedge with narrow, sharp-edged basal leaves 

and a dense terminal inflorescence of numerous 

dull-brown spikelets. Occasional on rocky or sandy 

shores and in coastal taro patches, reported only 

from coastal areas. Indigenous, widespread from 

tropical Africa to Hawai'i. Graeffe 1233; Setchell 

507; Eames T20; Garber 832; Bryan 961; Wisner 

39; Whistler 1423, 9023!.

Mariscus secmannianus (Boeck.) Palla Scleria lithosperma (L.) Sw. 

Medium-sized sedge with basal leaves and terminal 

inflorescences of brown spikelets borne on several 

rays. Occasional in lowland to montane forest, 

rcportcd from near sea level to 600 m elevation. 

Indigenous, ranging from Fiji to Tahiti. Setchell 26, 

506; Bryan 989; Eames T22; Christophersen 1819 

(n.~.); Long 3060; Whistler 2806, 2860, 2900, 2916, 

3648,8414,8415?, 9032!. 

*Mariscus sumatrensis (Retz.) Raynal 

Medium-sized sedge with green spikelets in several 

cylindrical spikes borne on spreading rays and 

subtended by several, long, leaf-like bracts. Un- 

common to occasional in disturbed places, reported 

only from the lowlands. Indigenous or perhaps a 

Polynesian introduction, originally native to the Old 

World tropics. Eames T12; Wilder 40, 72; Garber 

83 1 ; Whistler 2930. 

Medium-sized sedge with thin, erect stems and 

few-flowered inflorescences bearing tiny, hard, 

white, globose fruits. Occasional in coastal to 

lowland forest, reported from near sea level to 300 

m elevation. Indigenous, pantropic in distribution. 

Garber 828, 875; Wisner 146; Whistler 1478, 2846, 

8510!. 

Scleria polycarpa Boeck. 

tafatolu; selesele 

Coarse sedge with triangular stems, sharp-edged 

leaves, and branching inflorescences bearing tiny, 

hard, white globose fruits. Occasional in lowland to 

montane forest, reported fiom near sea level to 650 

m elevation. Indigenous, ranging fiom Australia to 

Samoa. USEE s.n. (n.s.); Graeffe 1230; Reinecke 

484; Setchell 224, 527; Wilder 82; Garber 874; 

Christophersen 1030, 126 1, 3572; Meebold 17007; 

McKee 2947; Whistler 2888,2898, 3.289, 863 1 !. 

I'ycreus polystachyos (Rottb.) Beauv. 

Small, erect sedge - with basal leaves and brown 

spikelets in irregular, head-like terminal clusters. 

*Torulinium odoratum (L.) S. Hooper 

Medium-sized sedge with basal leaves and a termi- 

Common on roadsides and disturbed places, repor- nal inflorescence bearing numerous linear spikelets 

ted only from the lowlands. A modem introduction, arranged perpendicular to the spreading rays. Occanow 

pantropic in distribution. Lamoureux 3080; sional in coastal marshes, on streamsides, and in 

Whistler 8964!. montane marshes, reported from near sea level to 

950 m elevation. Probably indigenous, pantropic in 

*Rhynchospora corymbosa (L.) Britten distribution. Setchell 94, 523; Yuncker 9401; Whiselesele 

stler 2858, 3604, 8450, 8896. 

Coarse sedge with long, sharp-edged leaves and 

branching terminal corymbs bearing red-brown 

DIOSCOREACEAE 

spikelets. Common in coastal and montane marshes, Dioscorea bulbifera L. 

reported from near sea level to 950 m elevation. [bitter yam] soi 

Indigenous, pantropic in distribution. 

Wilder 253; Kuruc 44; Whistler 3232. 

USEE s.n.; 

Herbaceous climbing vine with bulbils born on the 

stem, alternate cordate leaves, tiny flowers in 

*Scirpodendron ghaeri (Gaertn.) Merr. 

hanging in racemes, and longitudinally winged, 

oblong fruits. Common in disturbed places and 

secondary forest, reported from near sea level to 400 

m elevation. A Polynesian introduction, widespread 

from Africa to Hawai'i. Setchell 100; Garber 823; 

Christophersen 101 6; Whistler 1421, 3303, 8398, 

8444. 

Large pandanus-like sedge with basal, spine-edged 

leaves and large, irregularly globose inflorescences. 

Rare on Tutuila in mangrove forest at Tafuna, else- 

where in Samoa occurring in coastal marshes, estu- 

aries, montane craters, and swamp forests, reported 

from near sea level to ca. 400 m elevation. Indige- 

nous, ranging from Ceylon to Samoa. USEE s.n. 

(n.s.); Whistler 8056, 8506. 

*Dioscorea pentaphylla L. 

[five-fingered yam] pilita 

Vine with alternate, palmately divided leaves and 

tiny flowers in hanging racemes (but flowering

arely). Uncommon in disturbed places and second- 

ary forest, reportcd from near sea level to 350 m 

clevation. A Polynesian introduction, widespread 

from tropical Asia to Hawai'i. Diefenderfer 5; 

Whistler 8052, 8057, 8472. 

FLACELLARIACEAE 

*Flagellaria gigantea Hook. f. 

lafo 

High-climbing cane with large grass-like leaves 

modified at the tips into tendrils, and tiny white 

flowers in a branching terminal inflorescence. 

Uncommon to occasional in lowland to montane 


elevation. Indigenous, also found in Fiji and Niue. 

Whistler 3837, 8947, 9070. 

HELICONIACEAE 

Heliconia laufao Kress 

laufao 

Large herb with large, banana-like leaves, orange 

flowers borne in a large bract, and orange to red, 

fleshy fruits. Occasional in lowland to montane 

forest, reported from near sea level to ca. 900 m 

elevation. Endemic. Setchell 371; Garber 943; 

Diefenderfer 4; Whistler 9 17 1. 

HYDROCHARITACEAE 

"Halophila minor (Zoll.) Den Hartog 

Creeping marine herb with small, alternate, elliptic 

to obovate bullate leaves and inconspicuous axillary 

flowers. Common in shallow sandy lagoons. Indigenous, 

ranging from East Africa to Samoa. 

Sctchell 1 14, 181. 

*Halophila ovalis (R. Br.) Hook, f. 

Creeping marine herb with alternate, elliptic to 

oblong-elliptic leaves and inconspicuous axillary 

flowers. Locally common in shallow sandy lagoons. 

Indigenous, widespread from Africa to Samoa. 

Tsuda 262, 3 10; Whistler 3857; Cox 1128. 

ORCHIDACEAE 

Appendicula bracteosa Reichenb. f. 

Small, erect, laterally compressed epiphytic orchid 

with small white pendulous flowers in a terminal 

raceme. Occasional in montane to cloud forest, 


also found in Fiji. Christophersen 1 138!; Whistler 

8740!. 

*Bulbophyllurn atrorubens Schlechter 

Small epiphytic orchid with basal, obovate leaves 

arising from a bulb-like base, and tiny white flowers 

on a long stalk. Uncommon in montane forest, 


also found in New Caledonia and the Solomon 

Islands. Christophersen 1 185; Whistler 2902. 

Bulbophyllurn longiscapurn Rolfe 

Epiphytic orchid with acute-tipped leaves arising 

from a bulb-like base, and white flowers spotted 

with red, borne atop a long stalk. Occasional in 



the Solomon Islands to Niue. Setchell 274; Christophersen 

1037; Whistler 2827, 8059, 8724!, 9003!. 

*Bulbophyllurn pachyanthurn Schlechter 

Somewhat succulent epiphytic orchid with thick 

leaves arising from a bulb-like base, and large white 

flowers mottled red, borne atop a long stalk. Uncommon 

in montane forest, reported from 300 to 

600 m elevation. Indigenous, ranging from New 

Caledonia to Tonga and Samoa. Christophersen 

1044, 1 179, 1212; Whistler 3344. 

Bulbophyllum polypodioides Schlechter 

Small creeping epiphytic orchid with leaves arising 

from a bulb-like base, and a raceme of small white 

flowers. Common in lowland to montane forest, 


Indigenous, ranging from New Caledonia to Samoa. 

Setchell 383; Christophersen 1063, 1 18 1 ; Whistler 

8503, 8539!, 8726!, 8737!, 9005!. 

Bulbophyllum rosticeps Reichenb. f. 

Small creeping, epiphytic orchid with ovate leaves 

arising from a bulb-like base, and small, shortstalked, 

red solitary or paired flowers. Occasional to 

locally common from mangrove to montane forest, 


Indigenous, ranging from Vanuatu to Samoa. 

Wisner 140; Whistler 2901, 8695!, 8946!.

*Bulbophyllurn samoanum Schlechter 

Small epiphytic orchid with leaves arising from 

bulb-like base, and solitary yellowish flowers tinged 

red, borne atop a long stalk. Occasional in montane 


genous, ranging from New Caledonia to Samoa. 

Whistler 2787. 8496. 

Bulbophyllum sp. nova 

Prostrate epiphytic orchid with leaves arising from a 

bulb-like base, and a solitary yellow flower on a thin 

scape. Uncon~mon in lowland forest, reported from 

CYI. 1 SO to 270 m elevation. Indigenous, also found 

in Niue. Whistler 3757!, 8714!, 9348!. 

"Calanthe hololeuca Reichenb. f. 

Large ground orchid with basal, plicate leaves, and a 

tall raceme of showy white, spurred flowers. Occa- 

sional in montane to cloud forest, reported from 300 


New Caledonia to Samoa. Setchell 376, 546; 

Garber 783, 785; Christophersen 1020; Whistler 

2980.9 183. 

Calanthe triplicata (Wille. f.) Ames 

Large ground orchid with plicate leaves and tall 

racemes of showy white, spurred flowers marked 

with yellow. Common in lowland to cloud forest, 


Indigenous, ranging from southern China to the 

Marquesas. Setchell 408; Christophersen 1126!, 

1827!; Mitchell 561 ; Long 3050. 

Coelogyne lycastoides F. Muell. and Kraenzl. 

Large epiphytic orchid with large plicate leaves 

arising from a bulb-like base, cream colored flowers 

marked with brown, and a large, pendulous, winged 

capsule. Common in montane to cloud forest, re- 

ported from 300 to 1550 m elevation. Indigenous, 

ranging from New Caledonia to Samoa. Garber 

788; Christophersen 1021, 1068; Whistler 2796, 

8648!. 

Dendrobium biflorum (Forst, f.) Sw. 

Epiphytic orchid with stiff stems bearing grass-like 

leaves and paired white flowers with long-attenuate 

tepals. Common to abundant in lowland to mon- 

tane forest, reported from near sea level to 600 m 

elevation. Indigenous, ranging from Vanuatu to 

Tahiti. Garber 935; Whistler 2696,2791, 905 1 !. 

Dendrobium calcaratum A. Rich. 

Epiphytic orchid with long, longitudinally striate 

stems and short-stalked, axillary clusters of orange 

flowers borne on the leafless part of the stem. 



ranging from the Solomon Islands to Samoa. 

Whistler 2830, 3623, 8708!. 

"Dendrobiurn catillare Reichenb. f. 

Small epiphytic orchid with thick, longitudinally 

striate stems, scattered leaves, and clusters of white 

or pink flowers in the axils of leafless stems. Un- 

common in montane forest, reported from 250 to 

500 m elevation. Indigenous, also found in Fiji. 

Whistler 2903, 8945. 

Dendrobium dactylodes Reichenb. f. 

Epiphytic orchid with lanceolate leaves, and paired, 

axillary, pale orange to white flowers. Common to 

abundant in lowland to montane forest, reported 

from near sea level to 800? m elevation. Indige- 

nous, ranging from Vanuatu to Samoa. Setchell 

205, 205a, 247; Diefenderfer 21; Bryan 960; 

Christophersen 1 190, 1252; Yuncker 944 1, 9442; 

Whistler 2695,2751,283 1,2919, 5734, 8058. 

Dendrobium goldfinchii F. Muell. 

Erect, laterally compressed epiphytic orchid with 

tiny white flowers in a terminal raceme. Occasional 

in coastal to lowland forest, reported from near sea 


New Guinea to Samoa. Whistler 2895, 3346, 3622, 

8060, 8668!, 9103!. 

Dendrobium samoense Cribb 

Epiphytic orchid with somewhat succulent, purple- 

edged leaves and showy cream-colored flowers in a 

large terminal raceme. Occasional in lowland to 


m elevation. Endemic. Setchell 354; Yuncker 

9304; Whistler 2795, 8667!. 

Dendrobium sladei J. J. Wood and Cribb 

Epiphytic orchid with ovate leaves, flowering bran- 

ches arising above the leaf sheath, yellow stems, and

paired white flowers marked with red. Occasional 

in montane forest, reported from ca. 30 to ca. 600 m 

elevation. Indigenous, ranging from Vanuatu to 

Samoa. Christophersen 1256?; Yuncker 9391; 

Whistler 2694, 8497,9038!, 9088. 

Didymoplexis micradenia (Reichenb. f.) Hemsley 

Leafless, purple-stemmed, saprophytic ground 

orchid with one or a few white flowers in a terminal 

raceme. Occasional in lowland to foothill forest, 


ranging from New Caledonia to Niue. Whistler 

2847,2924,3656 9330!, 9345!. 

Diplocaulobiurn fililobum (F. Muell.) Kraenzl. 

Epiphytic orchid with long stems bearing a single 

leaf on top, and with long-stalked, cream-colored 

flowers bearing long-attenuate tepals. Occasional in 

montane to cloud forest, reported from 400 to 1550 

m elevation. Endemic. Christophersen 1203, 1213; 

Kuruc 10; Whistler 8788!, 9196. 

Eria robusta (Bl.) Lindl. 

Medium-sized epiphytic or ground orchid long, 

fleshy, imbricate leaves and a dense raceme of 

numerous, cream-colored flowers borne perpen- 

dicular to the rachis. Uncommon in lowland to 


Indigenous, ranging from India to Samoa. Wisner 

145; Whistler 8722!, 8936. 

* Ery th rodes purpurascens Schlechter 

Small ground orchid with white flowers in a termi- 

nal raceme. Uncommon in montane forest, reported 

from 300 to 890 m elevation. Indigenous, also 

found in Fiji and Tonga. Whistler 274 1. 

Flickingeria comata (Bl.) A. Hawkes 

Epiphytic orchid with large thick, oval leaves, and 

white flowers mottled red and bearing a yellow 

"beard." Occasional in mangrove to montane forest, 


Indigenous, ranging from Taiwan and Malaysia to 

Samoa. Setchell 335; Garber 933; Bryan 973; 

Christophersen 1 186, 3526; Whistler 2697, 2829, 

2894,904 1 !. 

Glomera montana Reichenb. f. 

Long-stemmed epiphytic orchid with a dense 

terminal head-like cluster of white flowers pink 

inside. Occasional in lowland to cloud forest, 

reported from 50 to 1000 (1200?) m elevation. 

Endemic. Christophersen 1040, 3527; Long 2606, 

3070. 

*Habenaria monogyne Schlechter 

Large ground orchid with lanceolate leaves and 

green, spurred flowers in a terminal raceme. Un- 



547. 

*Liparis caespitosa (Thou.) Lindl. 

Small epiphytic orchid with leaves arising from a 

bulb-like base, and white flowers in short, few- 

flowered racemes. Uncommon in montane scrub, 

reported from 500 m elevation. Indigenous, ranging 

from East Africa to the Austral Islands. Christo- 

phersen 1 182, 1200. 

Liparis condylobulbon Reichenb. f. 

Medium-sized epiphytic orchid with leaves arising 

from a bulb-like stem base, and small white flowers 

in several terminal racemes. Occasional to common 

in lowland to montane forest, reported from near sea 


Southeast Asia to Samoa. Garber 848; Wilder 15; 

Christophersen 1263; Wisner 138; Whistler 2699, 

2739,9040!. 

*Liparis gibbosa Finet 

Small epiphytic orchid with leaves arising from a 

bulb-like base and a long-stalked raceme of small 

yellow to orange flowers. Uncommon in montane 

scrub, reported from 500 to 600 m elevation. Indi- 

genous, ranging from India to Samoa. Christopher- 

sen 1062,1187, 1191. 

*Malaxis reineckeana (Schlechter) Kores 

Medium-sized ground orchid with plicate leaves and 

a terminal raceme of spurred, yellow flowers. Occa- 


300 to 750 m elevation. Endemic. Reinecke 620?; 

Long 3062.

Malaxis resupinata (Forst. f.) Kuntze 

Medium-sized ground orchid with plicate ovate 

leavcs and a narrow raceme of dark maroon flowers. 



lrorn Vanuatu lo Tahiti. Setchell 387; Bryan 91; 

Christophersen 1 1281, 3582!; Yuncker 943 117; 

Wisner 141; Whistler 2727. 

Malaxis samoensis (Schlechter) Whistler 

Small ground orchid with subfalcate leaves and 

small pale yellow flowers on a few-flowered scape. 


from 300 to 800 m elevation. Endemic. Christo- 

phersen 3567; Yuncker 943 1 !? (n. s.); Whistler 

2742,5725, 8507!, 8716!, 8994!. 

"Mediocalcar paradoxum (Kraenzl.) Schlechter 

Creeping epiphytic orchid with leaves arising from 

bulb-like bases and short racemes of yellow flowers. 



ranging from the Santa Cruz Islands to Samoa. 

Setchell 402; Christophersen 3488; Whistler 3853, 

91 87. 

Moerenhoutia heteromorpha (Reichenb. f.) Benth. 

& Hook. f. 

Medium-sized ground orchid with a terminal raceme 

of pubescent white flowers. Occasional in lowland 

to montane forest, reported from 300 to 750 m 

elevation. Endemic. Christophersen 1822!; Whistler 

2729,2775!. 

"Nervilia aragoana Gaud. 

Small ground orchid with cordate leaves arising 

from the ground and yellowish green flowers on a 

flowering scape that appears after the leaves die 

back. Uncommon in lowland to montane forest, 


ranging from India to eastern Polynesia. Whistler 

3776. 

Oberonia equitans (Forst. f.) Mute1 

Small, laterally compressed epiphytic orchid with 

tiny white flowers in a dense terminal raceme. 



ranging from New Caledonia to the Tuamotu 

Islands. Rechinger 16, 1281; Setchell 210, 336; 

Garber 880; Christophersen 1039; Long 3087?; 

Whistler 2906, 8666!. 

Oberonia heliophila Reichenb, f. 

Small to medium-sized, laterally compressed epi- 

phytic orchid with long strap-shaped leaves, and tiny 

white flowers in long, narrow terminal racemes. 



ranging from the Solomon Islands to Samoa. 

Reinecke 6 15; Diefenderfer 22; Yuncker 9307; 

Long 3 100; Whistler 2790, 9039!. 

"Peristylus tradescantifolius (Reichenb. f.) Kores 

Small erect ground orchid with lanceolate leaves and 

small greenish flowers in a terminal raceme. Occa- 

sional in lowland to cloud forest, reported from 200 

to 1000 m elevation. Indigenous, also found in Fiji 

and Tonga. Long 3 120; Whistler 2686. 

"Phaius amboinensis B1. 

Large ground orchid with basal, plicate leaves and 

tall racemes of large, showy white flowers yellow 

within. Occasional in lowland to montane forest, 


also found in Fiji. Whistler 2687. 

Phreatia matthewsii Reichenb. f. 

Small, laterally compressed orchid with linear leaves 

and several basal, pedunculate racemes mostly 

shorter than the leaves and bearing tiny white flow- 

ers. Common to abundant in mangrove to montane 


tion. Indigenous, ranging from New Caledonia to 

Tahiti. Setchell 238; Garber 938; Christophersen 

1057, 1193, 1211, 1259, 1821!, 3573; Yuncker 

9305; Wisner 140; Long 1951a; Whistler 2698, 

9OO4!. 

Phreatia micrantha (A. Rich.) Schlechter 

Medium-sized, laterally compressed epiphytic 

orchid with several basal racemes of tiny white 

flowers. Common in mangrove to cloud forest, 


Indigenous, also found in Fiji and Tonga. Setchell 

400; Christophersen 1 l33!, 1189, 3503, 3538; 

Mitchell 581; Long 3018; Whistler 2740, 9OO2!.

Phrcatia obtusa Schlechter 

Medium-sized epiphytic orchid with a swollen base 

and small white flowers borne in a long raceme. 

Ilncommon in montane forest, reported from 400 to 



*Pseuderia ramosa L. 0. Williams 

Scrambling, epiphytic or terrestrial orchid with a 

raceme bearing several greenish flowers. Occa- 

sional in montane scrub and montane forest, 


also found on Futuna. Christophersen 1190a, 3533; 


Spathoglottis plicata B1. 

Large ground orchid with basal plicate leaves and 

showy pink flowers on a long scape. Common in 

disturbed sunny places and fernlands, reported from 


Vanuatu to Samoa. Setchell 193; Mitchell 514; 

Yuncker 9416; Whistler 8558!. 

Taeniophyllum fasciola (Forst.) Reichenb. f. 

Tiny leafless and stemless epiphytic orchid, consi- 

sting of flattened photosynthetic roots, short race- 

mes of tiny white flowers, and a cylindrical fruit. 

Common in lowland forest, reported from near sea 


Guam and New Caledonia to Tahiti. Whistler 2918, 

2984,3 1 l7,3 1 l7a, 3343,8657!. 

Tacniophyllum gracile (Rolfe) Garay 

Tiny leafless and stemless epiphytic orchid consi- 

sting of flattened photosynthetic roots, tiny white 

flowers on a long scape, and an ellipsoid fruit. 

Uncommon in lowland forest, reported from 150 to 


Whistler 3 1 17,9102!. 

Thrixspermum graeffei Reichenb. f. 

Small epiphytic orchid with small white flowers 

borne on several long, spreading scapes. Occa- 

sional in montane forest, reported from 400 to 725 

m elevation. Indigenous, also found in Fiji. Chri- 

stophersen 1 180, 1192; Whistler 3749!, 3852. 

*Vrydagzenea vitiensis Reichenb. f. 

Small ground orchid with ovate leaves bearing a 

silvery median strip, and small white flowers in a 

terminal raceme. Occasional in lowland to montane 


genous, also found in Vanuatu, Tonga, and Fiji. 

USEE s.n.; Lister s.n.? (no locality listed). 

"Zeuxine stenophylla (Reichenb. f.) Benth. & 

Hook. f. ex Drake 

Medium-sized ground orchid with a terminal raceme 

of white flowers with rounded sepals. Occasional in 


m elevation. Indigenous, also found in Tonga and 

Fiji. USEE s.n.; Whistler 2747?. 

*Species indet. 

An unidentified, leafless orchid with a raceme of 

white (?) flowers was collected by W. Sykes (no. 36, 

27 Sept. 1974), at Malaeimi on Tutuila, and P. 

Kores was unable to identify it even to genus (pers. 

comm.). This may be Stereosandra javanica B1. 

PANDANACEAE 

Freycinetia reineckei Warb. 

'ie'ie 

Coarse climber with pandanus-like leaves 2.5-5 cm 

wide, auricles up to 13 cm long, unisexual inflorescences, 

ovaries with 2 stigmas, large oblong 

compound fruits 5.5-1 1 cm long, numerous tiny 

fruitlets up to 13 mm long, and the fruit stalk 

smooth. Common in lowland to cloud forest, 


Garber 906, 907; Long 3013, 3076?, 3077; Whistler 

2772!, 8935!, 8993!. 

Freycinetia storckii Seem. 

'ie'ie 

Coarse climber with pandanus-like leaves 1.8-2.8 

cm wide, unisexual inflorescences, ovaries with 4-6 

stigmas, and a large cylindrical compound fruit 4-7 

cm long by 1.5-2.3 cm wide on a scabrid stalk. 

Common to abundant in lowland to cloud forest, 


Indigenous, also found in Fiji. Reinecke 355c; 

Garber 930; Christophersen 3495; Long 3017; 

Whistler 2750, 2983, 3548, 3640!, 3661, 9136, 

9137. Another species was reported from Tutuila,

F. homhronii Mart.; it differs from F. storckii in 

having narrower leaves (1-2 cm wide) and shorter 

compound fruits (2.5-5 cm long). However, the 

specimen identified as F. hombronii, Garber 930, is 

sterile, and is likely to be just a juvenile form of F. 

slorckii. The two species may, in fact, be one in the 

same. 

"Pandanus reineckei Warb. 

fasa 

Dioccious tree-like plant with prop roots, long tough 

leaves spiny on the margins and lower midrib, and 

conlpound fruits bearing I-3-lobed keys. Common 

in montane scrub, reported from 450 to 1000 m 

clevation. Endemic. Reinecke 459; Setchell 72a, 

407; Christophersen 1022, 1219; Whistler 2713, 

3659. 

I'andanus tectorius Parkinson 

fasa 

Dioecious tree-like plant with prop roots, long tough 

leaves spiny on margins and lower midrib, and 

compound fruits bearing 4-9-lobed keys. Common 

on exposed shores, reported from near sea level to 

100 m elevation. Indigenous, widespread 

throughout the tropical Pacific. Reinecke 459; 

Setchell 32, 412; Garber 984; Whistler 1427, 3617, 

3754. 

*Pandanus whitmeeanus Mart. 

pogo 

Dioecious tree-like plant with prop roots, long tough 

leaves spiny on the margins and midribs, and 

compound fruits bearing long keys with stigmas 

lateral on the tip. Commonly cultivated and sometimes 

found in montane scrub, reported from near 

sea level to 550 m elevation. Probably a Polynesian 

introduction, ranging from Vanuatu to the Cook 

Islands. Setchell 300; Christophersen 3580; 


POACEAE 

Axonopus compressus (Sw.) Beauv. 

[carpet grass] 

Mostly prostrate, mat-forming grass with narrow 

inflorescences bearing 3 spikes (a terminal pair and 

a lower one). Common in lawns and sunny, disturbed 



America. Whistler 323 1, 8627!, 8899!. 

Bothriochloa bladhii (Retz.) S.T. Blake 

Medium-sized to tall grass with a branched panicle 

bearing awned, purple spikelets. Uncommon in dry, 


lowlands. A modem introduction, native to Asia. 

Whistler 2971, 9014!, 9160. 

Brachiaria mutica (Forssk.) Stapf 

[California grass] 

Large, soft-hairy grass with a branching inflorescence 

bearing awnless spikelets with conspicuous 

black stigmas. Common to abundant in disturbed 

areas, particularly in wet places, reported from near 

sea level to 300 m elevation. A modern introduction, 

probably native to north Africa, but now 

pantropic in distribution. Whistler 8969!, 9346!. 

Brachiaria paspaloides (Presl) C. E. Hubb. 

Erect, medium-sized grass with branching panicles 

of awnless, ovoid, acute-tipped spikelets. Occasional 

in disturbed places, reported from near sea 

level to 550 m elevation. Probably a Polynesian 


Setchell 509; Eames T24; Wilder 50; Whistler 327, 

1283,8561!. 

Brachiaria subquadripara (Trin.) Hitchc. 

Low grass with awnless, obovoid spikelets blunt at 

the tip, borne on several spreading branches of the 

rachis. Occasional in lawns and other sunny, 


A modern introduction, native to the Old World 

tropics. Whistler 8624!. 

*Cenchrus calyculatus Cav. 

Medium-sized to tall grass with softly spiny subglobose 

spikelets in a terminal spike. Rare in coastal 

areas and perhaps now extinct in Samoa, reported 

only from the lowlands. Indigenous, ranging from 

New Caledonia to the Society Islands. Powell s.n. 

*Cenchrus echinatus L. 

[bur grass] vao tuitui 

Medium-sized grass with unbranched spikes of 

bur-like fruits bearing hard spines. Common in 

disturbed places, reported only from coastal areas.


Setchell 502; Garber 830. 

Cet~tosteca lappacea (L.) Desv. 

sefa 

Erect grass with broad leaves and tiny awnless 

spikclets in a large panicle. Occasional to common 

in shady disturbed places and along forest trails, 


Possibly indigenous, or a Polynesian introduction, 

ranging from tropical Africa to the Marquesas. 

Graeffe s.n.; Setchell 78, 516; Garber 867, 889; 

Yuncker 9439; Wisner 161, 162; Long 3046; 

Whistler 2974, 3230, 8474, 8599!, 91 8 1. 

Chloris barbata (L.) Sw; 

[fingergrass] 

Erect grass with digitately arranged panicle branches 

of awned, purple spikelets. Common in dry 

disturbed places, reported only from coastal areas. 

A modern introduction, native to tropical America. 

Setchell 508; Eames T17; Wilder 21; Garber 801; 

Christophersen 980; Whistler 901 1 !. 

*Chloris radiata (L.) Sw. 

Erect grass with digitately arranged spikes of awned, 

silvery spikelets. Rare in pastures, reported only 

from the lowlands. Perhaps no longer found in 

American Samoa. A modern introduction, native to 

to tropical America. Whistler 305. 

*Chrysopogon aciculatus (Retz.) Trin. 

Low, mat-forming grass with awned, purplish spikelets 

borne in erect, scapose panicles. Common in 

lawns, streambeds, and fernlands, reported only 


native to the Old World tropics. Setchell 500; 

Eames T17; Garber 798; Lamoureux 3077. 

Coix lacryma-jobi L. 

[Job's tears] sagasaga 

Tall, robust grass with large leaves and spikelets that 

form white to gray, bead-like spikelets. Occasional 

to common in wet, disturbed places, reported only 


native to Asia. Setchell 296; Garber 896; Diefenderfer 

23; Christophersen 1207; Whistler 8616!. 

"Cynodon dactylon (L.) Pers. 

[Bermuda grass] 

Prostrate, mat-forming grass with erect inflore- 

scences usually bearing four digitately arranged, 

spicate branches. Common in lawns and in sunny, 


A modern introduction, possibly native to tropical 

Africa. Whistler 3300. 

Cyrtococcum oxyphyllum Stapf 

Medium-sized grass with lanceolate leaves and 

awnless spikelets borne on fine stalks in narrow 

panicles. Locally common in shady forest areas and 

disturbed places, reported from near sea level to 830 

m elevation. Indigenous or a Polynesian introdu- 

ction, ranging from Malaysia to Niue. Christo- 

phersen 3559; Whistler 2845, 851 l!, 8939?. 

*Cyrtococcum trigonum (Retz.) A. Camus 

Small, somewhat prostrate grass with pubescent 

awnless spikelets in a short, narrow panicle. 

Common in lawns and other sunny, disturbed 


elevation. A modern introduction, native to 

Southeast Asia. Lamoureux 3068; Long 2603; 


Dactyloctenium aegyptium (L.) Willd. 

Low grass with a short inflorescence stalk bearing 

1-5 thick, digitately arranged branches of purplish, 

bristly spikelets. Occasional in disturbed dry places, 


modern introduction, native to the Old World 

tropics. Wisner 26; Whistler 8623!. 

Digitaria ciliaris (Retz.) Koel. 

Medium-sized to tall grass with multiple whorls of 

digitately arranged panicle branches bearing awnless 

spikelets. Common in plantations and sunny, disturbed 


elevation. A modem introduction, native to Asia. 

Whistler 306, 8925!, 9013!. 

"Digitaria henryi Rendle 

Small grass with spikelets in a short panicle whose 

branches do not spread or extend much out of the 

sheath at maturity. Uncommon in disturbed places, 

reported only in coastal areas. Some authors include

this in D. ciliaris, but the two have entirely different Eleusine indica (L.) Gaertn. 

growth forms. A modern introduction, native to [goosegrass] ta'ata'a 

tropical Asia. Whistler 3760, 8416. 

Digitaria horizontalis Willd. 

Medium-sized grass with shortly awned spikelets in 

thick panicle branches (usually 2 or 3 of them terminal 

with one lower). Common in sunny distur- 

Ikect grass with awnless spikelets in digitately bed places, especially along dirt roads, ;eported 

arranged panicle branches bearing scattered hairs. from near sea level to 300 m elevation. A Polyne- 

Occasional in disturbed places, reported from near sian introduction, native to the Old World tropics. 

sea level to 350 m elevation. A modern introdu- Reinecke 268; Setchell 510; Garber 794, 810; 

ction, native to tropical America. Whistler 8913!. Whistler 8620!. 

*Digitaria radicosa (Presl) Miq. *Eragrostis pilosa L. 

Small grass with awnless spikelets in 2-4 digitately Delicate grass with fine, spreading panicle branches 

arranged panicle branches. Occasional in disturbed and tiny, flattened spikelets. Uncommon in disturplaces 

in coastal areas, reported from near sea level bed places, reported only from the lowlands. A 

to cu. 300 m elevation. Probably a Polynesian modern introduction, native to Southern Europe. 


Whistler 302. 

Lamoureux 3065. 

"Eragrostis tenella (L.) Beauv. ex Roemer & 

Digitaria setigera Roth ex Roemer & Schultes Schultes 

Medium-sized grass with paired spikelets on long 

branches arranged subdigitately and spreading little 

at maturity. Common along the coast and in sunny 

disturbed places, reported from near sea level to 950 

m elevation, Indigenous or a Polynesian intro- 

duction, ranging from India to Hawai'i. Garber 

797; Wilder 30; Wisner 27; Lamoureux 3079; 

Whistler 8924!, 8927!. 

Digitaria violascens Link 

[lovegrass] 

Delicate grass with narrow panicle branches bearing 

tiny flattened spikelets. Common in lawns and in 


lowlands. A modem introduction, native to the Old 

World tropics. Setchell 157, 505; Eames T6; Garber 

796; Christophersen 1178. 

Eriochloa procera (Retz.) C. E. Hubb. 

Narrow grass with a panicle of small awnless 

Medium-sized grass with shiny black inner glumes spikelets having a conspicuous purple ring at the 

and awnless spikelets on digitately arranged panicle base. Occasional in disturbed places, reported only 

branches. Uncommon to occasional in disturbed from the lowlands. A modem introduction, native to 

places, reported from near sea level to 600 m eleva- the Old World tropics. Lamoureux 3081; Whistler 

tion. A modern introduction, native to tropical Asia. 326, 2920, 3290, 8970!. 

Whistler 301, 8629!. 

*Imperata conferta (Presl) Ohwi 

Echinochloa colona (L.) Link. 

[barnyard grass] 

Erect grass with awnless spikelets closely packed on 

several short panicle branches. Common in sunny, 

disturbed places, reported only from near sea level 

to 300 m elevation. A modem introduction, native 

to the Old World tropics. Setchell 503; Eames T26; 

Wilder 21a; Garber 966; Lamoureux 3086; Whistler 

8625!. 

Medium-sized grass with silky spikelets arranged in 

a narrow, plumose inflorescence. Uncommon on 

Tutuila in sunny disturbed places, elsewhere in 

Samoa also found on montane lava and ash flows 

and femlands, reported from near sea level to 1500 

m elevation. . Indigenous, ranging from Southeast 

Asia to Samoa. USEE s.n. (n.s.); Whistler 293 1. 

*Ischamemurn indicum (Houtt.) Merr. 

Small grass with a pair of terminal, spreading race- 

mose branches of long-awned spikelets. Common

as a pasture grass and sometimes escaping in distur- 

bed places and roadsides, reported only from the 


Asia. Whistler 3 10, 3 13,375. 

lschaemum murinum Forst. f. 

Small clump-forming grass with awned spikelets 

borne in paired spikes. Occasional to locally 

common on rocky shores, reported only near the 

coast. Indigenous, also found in Tonga and Niue. 

USEE s.n.; Garber 806; Whistler 291 1, 8072, 8409, 

841 1,8417,8429,8564!, 8684!. 

"Ischaemum sp. 

Tall grass with awned spikelets in a long pair of 

spikes. Uncommon in the montane scrub of Mata- 

fao and Le Pioa Mountains of Tutuila, reported from 

ca. 450-650 m elevation. Endemic to Tutuila? 

Setchell 405?; Christophersen 3545; Whistler 5704, 

91 82. This species has not been identified with 

certainty yet, but was reported by Setchell to have 

been introduced by Mrs. Robert Louis 

Stevenson-which is doubtful. 

"Lepturopetium sp. 

Small grass with a pair of terminal spikes with 

embedded spikelets. Rare on coastal rocks, reported 

only on the coast at the east end of the island. Two 

species have been named in this recently recognized 

genus, but these may be formed from an inter- 

generic hybrid (Fosberg, pers. comm.). Probably 

indigenous, the same as or similar to other indivi- 

duals found on several other tropical Pacific islands. 

Whistler 5728, 8395. 

Lcpturus repens (Forst. f.) R. Br. 

Small grass with narrow leaves and spikelets em- 

bedded in a corky rachis. Common on rocky and 

sandy sea coasts, reported only along the coast. 

Indigenous, ranging from Ceylon to Hawai'i. USEE 

s.n. (n.s.); Setchell415; Whistler 2852, 8565!. 

"Melinus minutiflora Beauv. 

Tall, hairy, strong-smelling grass with a dense 

panicle of awned spikelets. Uncommon in disturbed 

places or perhaps just escaping from pastures, 


duction, native to Africa. Whistler 5729 

"Microstegium glabratum (Brongn.) A. Camus 

Small grass with several unjointed racemes of long- 

awned spikelets. Uncommon in montane forest, 

reported from 150 to 450 m elevation. Probably a 

recent introduction, but indigenous from the 

Philippines to Tahiti. Whistler 9193. 

Miscanthus floridulus (Labill.) Warb. 

Tall grass with large plumose inflorescence bearing 

silky, windborne spikelets. Common in sunny 

places in the lowlands and on fernlands, reported 


ranging from at least Micronesia to eastern Poly- 

nesia. Setchell 252; Eames T19; Garber 793; 

Whistler 1448, 9109!. 

Oplismenus compositus (L.) Beauv. 

Medium-sized grass with awned spikelets on several 

branches from the rachis. Common in disturbed 

shady places, reported from near sea level to 1750 m 

elevation. Probably a Polynesian introduction, now 

pantropic in distribution. USEE s.n. (n.s.); Wisner 

60; Whistler 8475, 8586!, 8719!, 9141. 

Oplismenus hirtellus (L.) Beauv. 

[basket grass] 

Small grass with awned spikelets in short clusters on 

the rachis. Common in shady lowland forest and 

disturbed areas, reported from near sea level to 250 

m elevation. Probably a Polynesian introduction, 

native perhaps from Australia to Southeast Asia. 

Long 3079; Whistler 2897, 8509!, 8728!, 9199. 

Paspalum conjugatum Bergius 

[T- grass] vaolima 

Creeping grass with flat, round spikelets on a pair of 

spreading, terminal inflorescence branches. Com- 

mon to abundant in sunny disturbed places, reported 



550; Eames T25; Garber 964; Whistler 8912!. 

"Paspalurn dilatatum Poiret 

Erect grass with several spreading panicle branches 

bearing 2 rows of flattened, round, hairy spikelets. 

Uncommon in disturbed places, reported only from 


South America. Sykes 75.

Paspalurn orbiculare Forst. f. 

Mcdiuni-sizcd grass with several branches of 

flattened spikelets on the main axis. Common in 

wct places, reported from near sea level to 1650 m 

elevation. Indigenous, widespread from Southeast 

Asia to Ilawai'i. Garber 805; Eames T8; Christo- 

phersen 3560; Whistler 2716, 2932, 2973, 8908!, 

9063.9 179. 

Paspalurn paniculaturn L. 

Icrect grass with numerous long panicle branches 

bearing small, round, flattened spikelets. Common 

to abundant in sunny disturbed places and forest 

clearings, reported from near sea level to 200 m 


America. Whistler 8622!. 

*Paspalurn setaceum Michx. 

Small grass with a solitary inflorescence branch on 

the apex of the rachis, and round, flattened spike- 

lets. Uncommon to locally common in lawns and 

sunny disturbed places, reported only from the 

lowlands. A modem introduction, native to North 


*Paspalum urvillei Steud. 

Tall grass with several long panicle branches 

bearing round, flattened, hairy spikelets. Uncom- 

mon in disturbed places, reported only from the 


America. Whistler 325, 1453, 8418. 

Paspalurn vaginaturn Sw. 

Medium-sized grass spreading by means of under- 

ground rhizomes and bearing spikelets on two 

terminal inflorescence branches. Common along 

estuaries and on rocky or sandy shores, reported 

only on the coast. Probably an early European 

introduction, pantropic in distribution. Whistler 

8921!. 

"Pennisetum purpureum Schumacher 

Robust grass with bristly spikelets in large yellow, 

cylindrical panicles. Occasional in clumps or 

thickets in disturbed places such as roadsides, 


modern introduction, native to tropical Africa. 


*Sacciolepis indica (L.) Chase 

Small grass with a long scape bearing a dense, 

narrow, cylindrical spike of short-awned spikelets. 

Uncommon in disturbed places, reported from 350 

to 650 m elevation. A modem introduction, native 

to the Old World tropics. Whistler 9 180. 

*Setaria glauca (L.) Beauv. 

Small grass with a dense, yellow, cylindrical inflo- 

rescence bearing spikelets subtended by basal 

bristles. Occasional in disturbed places, reported 



Whistler 2815, 9161. 

"Sorghum sudanense (Piper) Stapf 

Tall, erect, annual grass with leaves spotted with 

purple, and large, brown, awned or awnless spike- 

lets. Occasional to common in sunny, disturbed 


introduction, native to the Mediterranean area. 


Sporobolus diander (Retz.) P. Beauv. 

Clump-forming grass with linear leaves and a 

narrow cylindrical panicle with short ascending 

branches bearing awnless spikelets from which the 

brown seed drops. Common in lawns and disturbed 


elevation. A modem introduction, native to India. 

Larnoureux 3064,3084; Whistler 8971 !. 

Stenotaphrum micranthum (Desv.) C. E. Hubb. 

Medium-sized grass with spikelets embedded in a 

thickened rachis that does not break apart at 

maturity. Uncommon on sandy shores, reported 


widespread from Southeast Asia to the Society 

Islands. USEE s.n.; Rechinger 3703; Whistler 8420, 

8691!, 9025!. 

*Stenotaphrum secundatum (Walter) Kuntze 

Low, mat-forming grass with a spicate inflorescence 

with the spikelets embedded in the thick rachis. 

Uncommon in lawns and sunny disturbed places, 

reported only near the coast. A modem introdu- 

ction, perhaps native to both Africa and tropical 

America. Setchell201, 501 ; Whistler 1289, 8438.

Thuarca involuta (Forst. f.) Roemer & Schultes 

Prostrate grass with lanceolate leaves and a fruit 

which at maturity is enclosed within a subtending 

bract to make a water-tight case. Occasional to 

locally abundant on sandy shores, reported only on 

the coast. Indigenous, ranging from Southeast Asia 

to the 'I'uamotu Islands. USEE s.n.; Whistler 2948, 

8922!. 

*Zoysia tenuifolia Willd. ex Trin. 

[Japanese temple grass] 

Small, mat-forming grass with narrow, involute 

leaves and a short spike bearing tiny awnless 

spikelets. Locally common in villages in coastal 

areas, sometimes in littoral habitats, reported only 

from the lowlands. A modem introduction, native to 

Southeast Asia? (Whistler). 

TACCACEAE 

Tacca leontopetaloides (L.) Kuntze 

[Polynesian arrowroot] masoa 

Erect stemless herb with tall, deeply lobed, longstalked 

basal leaves and green flowers and fruits 

borne atop a long, longitudinally grooved scape. 

Common in littoral to lowland forest, reported from 

near sea level to 200 m elevation. Indigenous or a 

Polynesian introduction, ranging from India to 

Hawai'i. Setchell 129, 185; Meebold 265 10; 

Whistler 2844,9029!. 

ZINGIBERACEAE 

Costus speciosa (Koenig) Sm. 

[crepe ginger] 

Coarse unbranched herb up to 2 m or more in height 

with alternate leaves, terminal inflorescences 

bearing reddish bracts, and large, showy white, 

crepe-like flowers. Uncommon in disturbed places, 

probably an escape from cultivation, reported from 

near sea level to ca. 300 m elevation. A modem 

introduction, indigenous from the Himalayas 

eastward to perhaps New Guinea. Whistler 8706!. 

Etlingera cevuga (Seem.) R.M. Smith 

'avapui vao? 

Tall herb 2 or more m in height with very short, 

leafless inflorescences of orange flowers. Occasional 

to locally abundant in lowland to montane 

forest, especially along streams, reported from 300 

to 800 m elevation. Indigenous, or perhaps a 

Polynesian introduction, ranging from Fiji to the 

Society Islands. Christophersen 1 124! . 

Zingiber zerumbet (L.) Smith 

[shampoo ginger] 'avapui 

Medium-sized ginger with white fragrant flowers 

borne among bracts of an ovoid terminal inflorescence 

on a leafless stalk. Occasional in 

secondary forest and plantations, reported from near 

sea level to 600 m elevation. A Polynesian 

introduction, widespread from tropical Asia to 

Hawai'i. Setchell 37; Garber 900; Diefenderfer 1; 

Wisner 68; Whistler 1275; Whistler 3848, 8986!.

APPENDIX B: PLOT DATA 

(Basal area is measured in square inches.) 

PLOT 1 : MATAPAI POINT LOWLAND COASTAL FOREST (50 m elev.) 

No. No. Basal Rel. 

Species trees 6 in. + Area Dom. 

1. Garuga floribunda 

2. Dysoxylum maota 

3. Planchonella garberi 

4. Terminalia catappa 

5. Planchonella grayana 

6. Barringtonia asiatica 

7. Diospyros samoensis 

8. Erythrina variegata 

9. Guettarda speciosa 

10. Myristica fatua 

1 1. Cocos nucifera 

12. Diospyros elliptica 

13. Sterculia fanaiho 

14. Syzygium clusiifolium 

15. Polyscias samoensis 

16. Meryta macrophylla 

17. Morinda citrifolia 

18. Ficus tinctoria 

19. Buchanania merrillii 

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

PLOT 2: SAUMA LOWLAND COASTAL FOREST (185 m elev.) 



1. Diospyros samoensis 16 9 2358 31% 

2. Barringtonia asiatica 4 3 1268 17% 

3. Canarium harveyi 37 8 978 13% 

4. Garuga floribunda 2 2 784 10% 

5. Myristica fatua 9 6 369 5% 

6. Planchonella garberi 2 2 354 5% 

7. Elattostachys falcata 1 1 346 5% 

8. Guettarda speciosa 2 2 304 4% 

9. Canarium vitiense 1 1 20 1 3% 

10. Macaranga stipulosa 3 3 166 2% 

1 1. Planchonella samoensis 1 1 133 2% 

12. Calophyllum neo-ebudicum 1 1 95 1% 

13. Aglaia samoensis 6 1 62 1% 

14. Syzygium clusiifolium 2 1 45 1% 

15. Casearia sp. nova 6 1 44 1% 

16. Planchonella grayana 1 1 38 + 

17. Ficus scabra 1 0 20 + 

18. Polyscias samoensis 3 0 13 + 

19. Citronella samoensis 2 0 6 + 

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

100 43 7604 100% 

PLOT 3: TUFANUA LOWLAND RIDGE FOREST (60 m elev.) 



1. Calophyllum neo-ebudicum 

2. Intsia bijuga 

3. Calophyllum inophyllum 

4. Garcinia vitiensis 

5. Syzygium inophylloides 

6. Metrosideros collina 

7. Glochidion ramiflorum 

8. Homalium whitmeeanum 


10. Manilkara dissecta 

1 1. Elattostachys falcata 

12. Erythrospermum acuminatissimum 


14. Syzygium clusiifolium 

15. Eurya pickeringii 

16. Chionanthus vitiensis 

17. Psychotria insularum 

18. Alstonia pacifica 

19. Canarium harveyi

PLOT 4: MULIULU LOWLAND RIDGE FOREST (190 m elev.) 



1. Intsia bijuga 

2. Canarium harveyi 


4. Canarium vitiense 



7. Hibiscus tiliaceus 



10. Rhus taitensis 

11. Elaeocarpus ulianus 

12. Alphitonia zizyphoides 

13. Hernandia moerenhoutiana 

14. Syzygium dealatum 

15. Fagraea berteroana 


17. Barringtonia samoensis 


19. Citronella samoensis 

20. Flacourtia rukam 

2 1. Garuga floribunda 

22. Planchonella garberi

PLOT 5: SAUMA LOWLAND RIDGE FOREST (320 m elev.) 



I. Diospyros samoensis 37 29 2486 27% 

2. Syzygium dealatum 8 7 1533 17% 

3. Syzygium inophylloides 11 10 1476 16% 

4. Calophyllum neo-ebudicum 7 5 1022 11% 

5. Canarium vitiense 2 2 769 8% 

6. Planchonella garberi 4 4 676 7% 

7. Rhus taitensis 3 2 415 5% 

8. Myristica fatua 7 3 312 3% 

9. Aglaia samoensis 5 2 119 1% 

10. Elattostachys falcata 2 1 92 1% 

11. Citronella samoensis 2 2 88 1% 

12. Sterculia fanaiho 2 1 70 1% 

13. Garcinia vitiensis 1 1 38 + 

14. Meryta macrophylla 4 0 30 + 

15. Diospyros elliptica 1 1 28 + 

16. Buchanania merrillii 2 0 20 + 

17. Ixora samoensis 2 0 20 + 

100 70 9194 100%

PLOT 6: TIATAUALA LOWLAND RIDGE FOREST (180 m elev.) 










8. Rhus taitensis 



I 1. Myristica hypargyraea 


1 3. Sy~ygium samoense 

14. Ixora samoensis 

15. Aglaia samoensis 

16. Elattostachys falcata 

17. Syzygium samarangense 


19. Cerbera manghas 

20. Cyathea vaupelii 


22. Cyclophyllum barbatum

PLOT 7: FAIGA LOWLAND RIDGE FOREST (260 m elev.) 



I. Syzgium inophylloides 






7. Terminalia catappa 



10. Ficus tinctoria 

1 1. Syzygium samoense 


13. Myristica hypargyraea 






19. Crossostylis biflora

PLOT 8: OLO LOWLAND RIDGE FOREST (250 m elev.) 



1. Syzygium inophylloidcs 

2. Khus taitensis 





7. Bischofia javanica 

8. Planchonella samoensis 



1 1. Aglaia samoensis 





16. Gucttarda speciosa 


18. Elaeocarpus ulianus 

19. Myrtistica fatua 


21. Psychotria insularum

PLOT 9: AMALAU LOWLAND RIDGE FOREST (280 m elev.) 



I . Canarium vitiense 





6. Dysoxylum samoense 

7. Syzygium dealatum 

8. Cananga odorata 




12. Macaranga stipulosa 



16. Crossostylis biflora 


18. Canthium merrillii 


20. Planchonella samoense 


22. Antidesnla sphaerocarpum 


24. Hedycarya denticulata

PLOT 10: AMALAU LOWLAND VALLEY FOREST (50 m elev.) 



1. Dysoxylum maota 

2. Planchonella samoensis 

3. Inocarpus fagifer 


5. Kleinhovia hospita 


7. Bischofia javanica 

8. Cananga odorata 

0. Elacocarpus ulianus 

10. Macaranga stipulosa 

I I. Ficus scabra 


13. Pisonia umbellifera 



16. Barringtonia asiatica 


18. Planchonella garberi

PLOT 11: VATIA PASS LOWLAND RIDGE FOREST (350 m elev.) 



Calophyllum neo-ebudicum 

Syzygium inophyhlloides 

Syzygium samoense 

Buchanania merrillii 

Iiernandia moerenhoutiana 

Myristica fatua 

Fagraea berteroana 

Rhus taitensis 

Elaeocarpus tonganus 


Elatostachys falcata 

Astronidium pickeringii 

Garcinia myrtifolia 

Syzygium samarangense 

Planchonella garberi 

Alstonia pacifica 

Baccaurea taitensis 

Crossostylis biflora 

Canthium merrillii 

Cyathea truncata? 

Meryta macrophylla 


Dysoxylum huntii 

Palaquium stehiinii 

Psychotria insularum

PLOT 12: ALAVA MONTANE FOREST* (400 m elev.) 



I. Dysoxyll~n~ huntii 

2. Crossostylus bitlora 

3. Syzygium samoense 

4. Palaquium stehlinii 

5. Hedycarya denticulata 





10. Syzgyium inophylloides 



13. Fagraea berteroana 


15. Canthium merrillii 

16. Baccaurea taitensis 

17. Garcinia myrtifolia 

1 8. Elacocarpus tonganus 


20. Psychotria insularurn 

21. Ixora samoensis 



24. Glochidion cuspidatum 


26. Cyathea vaupelii 



*Takcn from data collected during the Amerson et al. survey (1982).

PLOT 13: SUNITAO LITTORAL FOREST (ca. 30 m elev.) 


Svecies trees 6 in. + Area Dom. 

Hernandia nymphaeifolia 

Erythrina variegata 

Pisonia grandis 


Hibiscus tiliaceus 

Dysoxylum samoense 

Ficus scabra 

Guettarda speciosa 

Syzygium dealatum 

Slerculia fanaiho 

Chionanthus vitiensis 


Myrtistica fatua 

Ervatamia obtusiuscula 

Morinda citrifolia 

PLOT 14. OFU LOWLAND DISCLIMAX FOREST* (ca. 150 m elev.) 





Terminalia catappa 


Neonauclea forsteri 

Gucltarda speciosa 


Ficus scabra 

Sterculia fanaiho 


Bischofia javanica 


Geniostoma rupestre 

Barringtonia samoensis 

Psychotria insularum 

Ervatamia obtusiuscula 

17. Ficus tinctoria 1 0 7 + 

*Taken from data collected during the Amerson er al. survey (1982). 

114 44 6462 100%

APPENDIX C: SEASONALITY OF IMPORTANT FRUIT-BEARING TREES AND 

SHRUBS IN TKE PARK AREA. 

The table below was complied from herbarium specimens at the Bishop Museum and in the personal 

collection of the author, augmented by notes from Dr. Pepper Trail (pers. comm.). Some of the species 

includc only llowering or fruiting dates; in mosl of these the mature fruits are small and not always 

clishguishablc from the flowers. Many of the gaps in the data reflect inadequate sampling; most of the trees 

on Tutuila can be found flowering at anytime of the year. This list should be considered only tentative, 

because data for many of these species is currently being recorded by Dr. Pepper Trail of the Department of 

Marine and Wildlife Resources in American Samoa. 

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

SPECIES FAMILY FLOWERING DATES FRUITING DATES 

ANACARDIACEAE 



ANNONACEAE 

Cananga odorata 

APOCYNACEAE 

Alyxia bracteolosa 

Alyxia stellata 

Ccrbcra manghas 

ARALIACEAE 

Meryta macrophylla 

Polyscias samoensis 


Barringtonia asiatica 


BURSERACEAE 


Canarium vitiense 

Garuga floribunda 

CLUSIACEAE 

Calophyllum inophyllum 



Mammea glauca 

COMUKETACEAE 


Tcrminalia richii 

Terminalia samoensis 

CUNONIACEAE 

Spiraeanthemum samoense 

Weinmannia affinis 

EBENACEAE 

Diospyros elliptica 




Elaeocarpus ulianus

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


EUPHORBIACEAE 

Aleurites moluccana 

Antidesma sphaerocarpum 

Raccaurea taitensis 


Drypetes vitiensis 

Glochidion cuspidatum 

Glochidion ramiflorum 

Macaranga harveyana 

Macaranga stipulosa 

Omalanthus nutans 

FABACEAE 


Inocarpus fagifer 

Intsia bijuga 



Erythrospermum acuminatisimum 

Flacourtia rukam 

GESNERIACEAE 

C y rtandra samoensis 

GOODENIACEAE 

Scaevola taccada 

HERNANDIACEAE 

Hcrnandia moerenhoutiana 

ICACINACEAE 

Citronel la samoensis 

Medusanthera samoensis 

LAURACEAE 

Litsea samoensis 

LOGANIACEAE 



LORANATHACEAE 

Decaisnina forsteriana 

MALVACEAE 


MELASTOMACEAE 

Clidemia hirta 

Mcdinilla samoensis 

Melastoma denticulatum 

MELIACEAE 

Aglaia samoensis 


Dysoxylum maota 


MONlMlACEAE 

Hcdycarya denticulata

MORACEAE 

Ikus godcfl'royi 

1;icus obliqua 

Ficus prolixa 

Ficus scabra 

Ficus tinctoria 

Ficus uniauriculata 

MYRISTICACEAE 


Myristica hypargyraea 

MYRSINACEAE 

Embelia vaupelii 

Maesa tabacifolia 

MYRTACEAE 

Descaspermum fruticosum 

Metrosideros collina 

Psidium guajava 

Syzygium brevifolium 

Syzygium carolinense 

Syzygium clusiifolium 


Syzygium inophylloides 



OLEACEAE 

Jasminum betchei 

Jasminum didymum 

PII'ERACEAE 

Macropiper puberulum 

Piper graeffei 

RHAMNACEAE 

Alphitonia zizyphoides 

RHIZOPHORACEAE 

Crossostylis bifora 

RUBIACEAE 

Aidia cochinchinensis 

Calycosia sessilis 


Cyclophyllum barbatum 

Geophila repens 


Gynochtodes epiphytica 

lxora samoensis 


Morinda myrtifolia 

Mussaenda raiateensis 


Psychotria forsteriana 

Psychotria insularum


RUBIACEAE (cont'd.) 

Sarcopygme pacifica 

Tarenna sambucina 

RUTACEAE 

Euodia samoensis 

Micromelum minutum 

SAPINDACEAE 

Allophylus timoriensis 

Arytera brackenridgei 

Elattostachys falcata 

Sapindus vitiensis 

SAPOTACEAE 

Manilkara dissecta 

Palaquium stehlinii 


Planchonella grayana 

Planchonella samoensis 

SOLANACEAE 

Solanum viride 

STERCULIACEAE 


THEACEAE 

Eurya pickeringii 

THYMELAEACEAE 

Phaleria disperma 

Phaleria glabra 

Wikstroemia foetida 

TILIACEAE 

Grewia crenata 

Trichospermum richii 

ULMACEAE 

Trema cannabina 

URTICACEAE 

Cypholophus macrocephalus 

Leucosyke corymbulosa 

Pipturus argenteus 

Procris pedunculata 

VERBENACEAE 

Faradaya amicorum 

Premna serratifolia


MONOCOTS 

AGAVACEAE 

Cordyline fruticosa 1 2 - 4 - - -8-101112 1 - -4 -678 - -1112 

ARACEAE 

Epipremnum pinnatum - - - - - - - - - 10 -- -- 

Rhaphidophora graeffei - - - -5 -78 -10 -- -- 

FLAGELLARIACEAE 

Flagellaria gigantea - - - - - - - - - ---- -- - - - - - - - 8-10 -- -- 

IIELICONIACEAE 

Heliconia laufao - - -45 - -89 -- -- -- - - -45 - -89 -- 

PANDANACEAE 

Freycinetia reineckei - - - - -67 - - -- -- 12 1 - - - - - 7 8 - -- 

Freycinetia storckii - - - - - -78 - -- - - - - - - 7 8 9 10 -- 12 

Pandanus reineckei - - - - - - - - ------- - - - - 5 - - - - 10 -- -- 

Pandanus tectorius 1 - - - - - 7 - - -- 11 -- 123 - - -7 - - -- 

TACCACEAE 

Tacca leontopetaloides 123 -567 - -10 -12 

ZINGIBERACEAE 

Zingiber zerumbet 1 2 3 - - - - - 9 -- 11 12

APPENDIX D. THE OFU UNIT OF THE NATIONAL PARK OF 


INTRODUCTION 

Part of the island of Ofu in Manu'a is to be 

included in the National Park of American Samoa, 

principally because of the superb reefs located there. 

A botanical survey was carried out on the island on 

the 14th to the 16th of June 1992. The island, which 

is about 7.5 km2 (3 mi2) in area, is located about 95 

km (60 mi) east of the main island of Tutuila. It is 

part of the caldera rim of an ancient volcano, and is 

separated from the somewhat similar, adjacent 

island of Olosega by a narrow strait now crossed by 

a concrete bridge. The highest elevation of Ofu is 

Mt. Tumu at nearly 500 m (1621 ft) elevation. A 

single village is present on the west side of the 

island, but several houses and a small terminal 

building are located next to the airstrip, which lies in 

an east-west direction at the south-central tip of the 

island. Perhaps only two houses are found adjacent 

to the park area itself. 

The area under consideration is a narrow strip of 

land along the southeast-facing coast, extending 

from just east of the airport to Asaga Strait that 

separates Ofu from Olosega. This includes the 

pinnacle called Sunitao Peak that extends up to 230 

m (750 ft) elevation. The road from O h Village to 

Olosega represents the landward edge of the park 

boundary. Although not in the original proposed 

boundaries, the steep slopes leading to Mt. Tumu are 

important to the integrity of the park, so they, along 

with the summit of the island, were included in the 

present study. 

Ofu is a very small island, and it is not surprising 

that it has a flora much smaller than that of Tutuila; 

its flora is also much smaller than that of the nearby 

Ta'u documented in the early study for the National 

Park Service (Whistler 1992b). A checklist of the 

island is found in the vegetation study of American 

Samoa (Whistler 1980). Perhaps only three species 

found on Ofu are not found on Tutuila; Psychotria 

garberiana, which is endemic to Manu'a, Gyrocar- 

pus americanus, which in American Samoa is repor- 

ted only from Ofu (Nu'utele Islet), and Crateva 

religiosa, which in American Samoa is reported 

only from Ofu. 

THE VEGETATION 

The plant communities found on Ofu are the 

same ones described from the park area on Tutuila. 

Each of the natural communities present will be 

briefly described. For more general remarks about 

these communities and the disturbed communities 

(managed land, secondary scrub, secondary forest), 

see the comparable sections in the discussion of the 

vegetation of Tutuila. 

1. Herbaceous Strand 

The herbaceous strand is found along the whole 

length of the beach from the airstrip to the bridge 

separating the two islands. It is very narrow in 

extent, and is mostly not distinct from the littoral 

forest, largely because of its extensively damaged 

condition. The shore from the airport to Mt. Sunitao 

is a sandy beach, but at Sunitao it becomes rocky 

and almost vertical in places, before returning to 

sand just before the bridge. 

The dominant plants on the sandy beach is the 

beach morning-glory vine Ipomoea pes-caprae 

(beach morning-glory, fue moa) with lesser 

amounts of the littoral grasses Lepturus repens, 

Stenotaphrum micranthum, and Thuarea involuta. 

Other species present include Vigna marina (beach 

pea, fue sina), Triumfetta procumbens, Portulaca 

samoensis, Digitaria setigera, Tacca leonto- 

petaloides (Polynesian arrowroot, masoa), and 

Chamaesyce atoto. All of these, except for 

Chamaesyce and Triumfetta are found within the 

Tutuila park. Several other species of weeds are 

present, but they are secondary in importance to the 

native species. 

The dominant herbaceous plants on ~initao 

Peak 

are Phymatosorus scolopendria (lau magamaga) on 

rocky places and Lepturus repens on the gravel of 

talus slopes. Other native plants present include 

Portulaca samoensis, Portulaca lutea, Hedyotis 

foetida, and Mariscus javanicus. These plants are 

mixed in with shrubs, which will be noted below 

under littoral shrubland.

2. Littoral Shrubland 

Two types of littoral shrubland are present in the 

park, one on the sandy beaches where it alternates 

with littoral forest and herbaceous strand, and the 

other on the rocks of Sunitao and steep maritime 

cliff running inland along most of the length of the 

park. The most common shrub mixed in with the 

other littoral plants on the sandy beach is the beach 

sunflower Wollastonia bijlora (beach sunflower, 

skate), with lesser amounts of Scaevola taccada 

(to'ito'i), Sophora tomentosa, Colubrina asiatica 

(fisoa), and Clerodendrum inerme (aloalo tai). 

Sophora should be considered a rare plant in Samoa; 

it is known in the archipelago from only one beach 

on 'Upolu, one on Tutuila (Nu'uuli Point), the south 

part of 'Aunu'u, and this Ofu beach. 

The littoral shrubland on the steep sides of 

Sunitao and the cliffs of the main part of Ofu are 

much more complex and inaccessible. The slopes of 

Sunitao are dominated by a mixture of littoral trees 

and shrubs. Because the trees are often dwarfed, it 

is perhaps best to consider this vegetation as littoral 

shrubland, as was done with the vegetation of Pola 

Islet and the cliffs of Pola'uta Ridge on Tutuila. 

Littoral forests, however, are present on the gentler 

slopes of Sunitao and are discussed below. 

The dominant plants in the littoral shrubland of 

Sun itao are Dendrolobium umbellatum (lala), 

Clerodendrum inerme, Hibiscus tiliaceus (beach 

hibiscus, fau), and Pisonia grandis (pu'avai). To 

label the vegetation between the littoral forest and 

lowland forest on the cliffs on the main part of Ofu 

as littoral shrubland seems a bit incongruous since 

this community is supposed to be between the shore 

and the littoral forest. However, since it is compo- 

sed of shrubby species rather than trees, it is best to 

consider it littoral shrubland, regardless of its 

location. 

The dominant species on the steep slopes are 

Dendrolohium umbellatum, Phymatosorus scolo- 

pendria, Davallia solida (laugasese), Clerodendrum 

inerme, and Wollastonia bipora. Other species 

noted here include Morinda citrifolia (Indian mul- 

berry, nonu), Syzygium dealatum (asi vai), Ipomoea 

macrantha, Pipturus argenteus (soga), Pisonia 

grandis, Macaranga harveyana (lau pata), Colu- 

hrina asiafica, and GarugafIoribunda (vivao). 

Two other less common species were also noted 

here. The first, Gossypium hirsutum (vavae), is the 

native species of cotton that is very rare on Tutuila 

(one location recorded) and 'Aunu'u, and this was a 

surprising record for Ofu. The other, Plumbago 

zeylanica, was reported twice from the adjacent 

Olosega, but this is a new record for Ofu. Both 

species should be considered rare in American 

Samoa. 

3. Littoral Forest 

The division between this community and littoral 

shrubland is not very distinct on Ofu, especially 

when the slopes get steeper and trees get more 

shrubby. However, three types of littoral forest 

were recorded. The first is found on the rocky 

shores of Sunitao. Some of these slopes have been 

disturbed--the forest has been cut down and the area 

replanted with coconuts and bananas. This is parti- 

cularly the case on the east side of the mountain. 

Probably the natural littoral forest there was domi- 

nated by Barringtonia asiatica, as it is on the 

southwest side, and is similar to what is found in the 

Tutuila park unit. 

The gentle slopes leading up to the steeper slopes 

and cliffs are dominated by the second kind of lit- 

toral forest. A sample on the southwest side, inland 

of the Barringtonia littoral forest, showed it to be 

dominated by a mixture of littoral forest trees. In 

descending order of dominance, these are Hernan- 

dia nymphaeifolia (Chinese lantern-tree, pu'a), 

Erythrina variegata (coral tree, gatae), and Pisonia 

grandis (pu'avai). Of lesser importance are Diospy- 

ros samoensis ('au'auli), which was the most com- 

mon tree present, Hibiscus tiliaceus (beach hibiscus, 

fau), Dysoxylum samoense (mamala), Ficus scabra 

(ma ti), Guettarda speciosa (pua pua), Syzygium 

dealatum (asi vai), and Sterculia fanaiko (fana'io). 

Strangely, there was no Barringtonia in the sample 

on this slope. The understory was strongly domi- 

nated by seedlings and saplings of Syzygium 

dealatum and Diospyros samoensis, with lesser 

amounts of Psychotria insularum (matalafi), and 

Morinda citrifolia. Other significant plants included 

the bird's-nest fern Asplenium nidus (laugapapa) 

and Cordyline fruticosa (ti), with a few other 

species. The ground cover was relatively light, with 

perhaps only 15% cover. The only vines noted were 

Epipremnum pinnatum, Dioscorea bulbifera (bitter 

yam, soi), and Faradaya amicorum (mamalupe). A

similar forest is probably found on the same kind of 

slopes below the cliffs on the main part of Ofu, 

often mixed with lowland forest trees. This area is 

further discussed below. 

'Two other significant small trees are found here. 

Chionanthus vitiensis is a new record for American 

Samoa, and was collected on Tutuila (Vatia) and on 

Ofu. It is known from only two records in Western 

Samoa; only a single tree was found in each of these 

four records. Ervatamia obtusiuscula is uncommon 

on Ofu, and has not been collected on Tutuila since 

1894. Both of these species should be considered to 

be rare in American Samoa. 

The littoral forest on the sandy beach was differ- 

ent in species composition. In most places it is 

highly disturbed and dominated by planted and 

naturalized coconuts. At the east end this was a 

mixture of coconuts and Hernandia nymphaeifolia, 

which were shattered by one or more of the recent 

hurricanes. Also present on the beach is Terminalia 

samoensis (talie), which is not present in the Tutuila 

park unit. The west end of the beach, just to the east 

of the airstrip, is rocky and narrow (between the 

beach and the road), and is dominated by a combi- 

nation of Hernandia nymphaeifolia, Thespesia 

populnea (Pacific rosewood, milo), Barringtonia 

asiatica, and Hibiscus tiliaceus. Much less common 

there are Terminalia catappa, Morinda citrifolia, 

Guettarda speciosa, Macaranga harveyana, and 

Erythrina variegata. 

4. Coastal Marsh 

The wetland situation on Ofu is similar to that on 

'htuila: on both there is a coastal marsh just outside 

the boundaries of the park. But since this is an 

important type of vegetation, and it is so close to the 

park, it is discussed here. Although the two marshes 

are similar in their geological structure, they differ 

greatly in their vegetation. While the Vatia marsh 

has been abandoned, the one on Ofu is actively 

being cultivated. It was, that is, until storm waves of 

a recent hurricane swept across it. Today it is still 

mostly barren mud, probably unable to support a 

native marsh or cultivated taro because of the 

residual salt concentration. It is a habitat for native 

ducks (toloa). 

Originally the vegetation was dominated by 

Eleocharis dulcis, possibly in combination with 

Cyclosorus interruptus, and these species will 

probably return to dominate the uncultivated 

portions of the marsh. Also present is the small 

fragrant herb Limnophila fragrans. This plant is 

currently known in the territory only from the Ta' 

village marsh, the Ofu marsh, the Olosega marsh, 

and the Vatia marsh, and should be considered rare 

in Samoa. It is probably even rarer in Western 

Samoa. It was found to be present in small quan- 

tities in the brackish mud of Ofu marsh, and will 

probably become more common as the marsh 

returns to its normal state. 

5. Lowland Forest 

The lowland forest within the park is disturbed 

-on the gentler slopes by past and present agri- 

culture, and on the steeper slopes by landslips. A 

plot was sampled in this vegetation in the 1976 

survey (Whistler 1980) on the talus slope east of the 

airstrip (Appendix B: plot no. 14, modified from the 

original to correspond to the methods used in the 

present study). The dominant species in that plot 

were Hibiscus tiliaceus (18% relative dominance), 

Dysoxylum samoense (1 8%), Terminalia catappa 

(13%), Myristica fatua (12%), Neonauclea forsteri 

(1 I%), Guettarda speciosa (1 I%), and Diospyros 

samoensis (8%). Other species noted to be present 

on the slopes and cliffs during the June 1992 survey 

are Pisonia grandis, Sterculia fanaiho, Pometia 

pinnata, and Ficus obliqua. 

This vegetation is best classified as a disclimax 

lowland forest and probably best corresponds to the 

"Dysoxylum lowland forest on talus slopes and 

cliffs" noted in the Ta'u report (Whistler 1992b) as 

occurring in similar situations on that island. It is 

also similar to the vegetation in the Tutuila park 

between Alava Ridge road and Fagas, and probably 

on the steep slopes of Alava Ridge above Pago Pago 

(which are mostly outside the Tutuila park 

boundaries). 

MONTANE FOREST 

The vegetation on top of the ridge above the Ofu 

park area was also briefly examined. The top of the 

island has been extensively disturbed by past and 

recent agriculture and by the recent hurricanes. 

There is also a road leading from Ofu village up to 

the summit of the island (Mt. Tumu) to a microwave 

station. 

Since the summit is at 500 m elevation, the forest 

is or was probably best classified as montane forest, 

similar to that on Tutuila, but with fewer species. In

the relatively undisturbed places, the dominant 

species are Syzygium samarangense, Calophyllum 

neo-ehudicum (tamanu), Syzygium samoense (fena 

vao), and Myristicafatua ('atone); the two Syzy- 

gium species are typical montane forest trees, the 

other two are lowland forest species. Also present 

are f)ysoxylum huntii and Garcinia myrtifolia, which 

are typical montane forest species, and several other 

tree species characteristic of lowland forest. One 

tree found here, Celtis harperi, was a new record for 

Ofu-Olosega. 

In disturbed places, the dominant tree was Hibi- 

scus liliaceus, which often forms dense thickets. 

These thickets are similar to those on Alava Ridge 

east of the TV transmitter on Tutuila. Also common 

is another secondary forest tree, Trichospermum 

richii. The ground cover is dense here, and is 

dominated by Christella harveyi, Lomagramma 

cordipinna, and Davallia solida, all of them ferns. 

As the ridge extends east from the microwave 

station, it decreases in elevation and the vegetation 

probably goes from ridge lowland forest to coastal 

lowland forest before it meets the road just west of 

Sunitao. The ridge at this point is very narrow, and 

the vegetation is sparse, somewhat like that found on 

the steep slopes leading from the summit down to 

the park area. One tree noted on this ridge was 

Drypetes vitiensis, which is a new record for 

Manu'a.

APPENDIX E. PLANTS AS INDICATORS OF PAST LAND USE 

Prior to the arrival of the first Polynesian settlers, 

most of the surface area of the Samoan archipelago 

was covered with native rainforest vegetation. The 

only areas with vegetation not classified as rain- 

I'orcst were littoral communities, summit vegetation 

(montane scrub and summit scrub communities), 

recent lava flows and ash deposits, and disturbed 

vegetation. Nowadays over half of the vegetation of 

the islands is classified as disturbed or secondary (as 

opposed to primary), a figure that has been rising in 

recent years because of the expansion of agriculture 

to meet the needs of an expanding population and 

because of a forestry industry that has devastated the 

lowlands of the archipelago (particularly in Western 

Samoa). Prior to human settlement, all the disturbed 

vegetation was natural, caused mostly by hurricanes 

and landslips, and possibly by occasional forest 

fires. Nowadays, however, most of the disturbed 

vegetation is the result of human activities such as 

agriculture and the development of villages. The 

disturbed communities recognized in the present 

report are managed land vegetation, secondary 

scrub, and secondary forest. 

It is sometimes useful to know the past uses of an 

area of land. In order to set lease prices within the 

park, there is a need to determine which areas are 

currently or were historically used for agriculture. 

This was done mainly by the use of aerial photo- 

graphs and the maps of Cole et al. (1988). More 

accurate studies involve the structure and floristic 

composition of the vegetation. Native Samoan plant 

species require certain environmental conditions for 

germination and growth, and it is often possible to 

determine the recent historic uses of the area by 

analyzing the species composition, community 

structure, and population dynamics of an area of 

vegetation. This is most easily done by looking for 

indicator species, plants that are indicative of a 

certain type of human activity or climatic event. 

The most obvious sign of recent disturbance is the 

dominance of herbaceous vegetation. The only kind 

of natural vegetation in Samoa that is herbaceous is 

in littoral areas (herbaceous strand) and montane 

scrub (which is characterized by ferns and climbers 

dominating the ground between scattered trees and 

shrubs). The most common species in the recently 

129 

disturbed areas, most of which are abandoned taro 

or banana plantations, are Mikania micrantha 

(mile-a-minute vine, fue saina), Bidens alba 

(beggar's-tick), Paspalum conjugatum (T-grass, vao 

ha), and Nephrolepis hirsutula (sword fern, vao 

tuaniu), but over 250 species that fit into this 

category (weeds) have been recorded from Tutuila 

(see Appendix A). 

Several other species planted as ornamentals 

sometimes remain significant long after cultivation 

has ended. One such species, the mint Pogostemon 

cablin (patchouli, patiale), sometimes forms dense 

patches by vegetative means; it has not been 

recorded from the Tutuila park, however. Other 

weedy species are typical of forest trails. Axonopus 

fissifolius and Eleusine indica (ta'ata'a) are often 

found in lowland plantations and along trails in 

lowland forest. Two other grasses, Oplismenus 

compositus and Oplismenus hirtellus, are often 

found in secondary forest and thus can be indicative 

of past disturbance. Another two grasses, Cento- 

steca lappacea (sefa) and Cyrtococcum oxyphyllum, 

occur in primary vegetation, and are thus less 

valuable as indicator species. 

One species of fern, Dicranopteris linearis (false 

staghorn fern, asaua), is a natural dominant species 

in montane scrub (which is usually on trachyte soil), 

but its presence in other places, such as ridges, is an 

indication of past disturbance. There are only a few 

areas (outside of montane scrub) on Tutuila that are 

reported to be dominated by this fern. These are 

probably areas of former plantations or villages on 

ridges, from a period when the bulk of the Samoan 

population lived inland (for protection from 

marauders and enemies). The soil has been so 

over-utilized and the soil minerals so depleted that 

the native vegetation (mostly ridge lowland forest) 

has never been able to recover. Similar vegetation is 

found on 'Upolu, and is attributed to the same cause 

(Wright 1963). 

When managed land is left alone for long enough, 

the herbaceous vegetation is shaded out by a number 

of characteristic woody shrubs and trees. The most 

typical of these are Pipturus argenteus (soga), 

Omalanthus nutans (fogamamala), Macaranga

harvcyana (laupata), Morinda citrifolia (Indian 

mulberry, nonu), and Kleinhovia hospita (fu'afu'a). 

Less common, but still typical species, include 

7'rema cannabina (magele) and Melochia aristata 

(ma'o). Vegetation dominated by any combination 

of these species is probably on land that was in culti- 

vation in the last decade or two (with the exception 

of "disclimax" vegetation on steep slopes, discussed 

below). 

Another indicator species is Hibiscus tiliaceus 

(beach hibiscus, fau). This tree is one of the most 

adaptable species in Samoa, where it is common in 

disturbed lowland and littoral forest, on the margins 

of mangrove swamps, on talus slopes, and along 

streams. The tree is very useful, since its wood is 

widely used for firewood, artifacts (such as canoe 

outriggers and the sticks used for making fire by 

friction), and light construction. The bark is also 

useful for making cordage of a temporary or perma- 

nent nature. 

Hibiscus does not seem to have good dispersal 

ability, since the fruits and seeds are not wind- or 

bird-dispersed. It probably gets around by being 

planted (since it is so useful) or may spread slowly 

from adjacent areas where the trees are present. It is 

often found in what otherwise appears to be undis- 

turbed ridge lowland forest. It seems to be able to 

persist for long periods of time, but is probably 

eventually shaded out when the secondary forest 

canopy closes. Its presence in these situations may 

indicate that the area was disturbed long ago for 

inland villages or plantations. A dense stand of this 

along Alava Ridge east of the TV transmitter is 

probably related to the presence of an ancient trail 

li-om Vatia village to Pago Pago Bay. The presence 

of Hibiscus on steep talus slopes is probably natural, 

since this type of "disclimax vegetation" is in a 

continual state of disturbance due to landslips and 

rock falls. Another tree with somewhat similar 

distribution, ecological needs, and stature as 

Hibiscus is Kleinhovia hospita (fu'afu'a) mentioned 

above. but this is less common than Hibiscus. 

Cocos nucifera (coconut, niu) is another good 

indicator tree of past disturbance. Although it is 

rarely found very far inland in native vegetation, its 

presence in significant numbers in secondary forest 

is an almost certain indicator of a former plantation. 

It is not able to compete with inland species in these 

habitats. It is also probably unable to naturally 

compete with littoral forest trees, and its presence 

along uninhabited coast is usually an indication that 

it was planted there. Other trees, such as the inland 

species Paraserianthes falcataria (tamalini), 

Flueggeaflexuosa (poumuli), and Erythrina subum- 

brans (dadap, gatae palagi), do not readily spread, 

and are indicators of more recent disturbance, since 

they are all of modem introduction. 

A number of native canopy species are also indi- 

cators of past disturbance-from both natural and 

human causes. The most common of these is Rhus 

taitensis (tavai). This tree often forms monodomi- 

nant forests, particularly on the western half of the 

island. Apparently the tree, which has readily 

dispersed seeds (in small succulent fruits), germi- 

nates mostly in bright sunlight, because seedlings 

and saplings are common in areas where native 

forest has recently been cleared for agriculture, and 

possibly also when forests are flattened by hurri- 

canes. Seedlings in closed-canopy forest are un- 

common. Thus forests dominated by Rhus are most 

likely areas once extensively disturbed decades 

earlier (the tree probably lives for decades, and per- 

haps more than a century, since individuals of nearly 

2 m dbh have been recorded (Whistler 1980). 

A species with similar ecological needs and life 

history as Rhus is Alphitonia zizyphoides (toi), 

which is sometimes a co-dominant species with 

Rhus. Other species that likewise indicate past 

major disturbance are Pometia pinnata (tava), 

which, however, is not found within the park, 

Neonauclea forsteri (afa), Cananga odorata 

(ilangilang, moso'oi), and Bischofia javanica ('o'a). 

Two other secondary forest species, Elattostachys 

falcata (taputo'i) and Macaranga stipulosa (lau 

fatu), are probably more indicative of "gap replace- 

ment" and are not useful as indications or major 

disturbance to the vegetation. Seedlings of Elatto- 

stachys are often common in lowland forest, but 

adult trees are not, indicating that they may only 

mature if a canopy gap appears while they are a 

seedling or sapling; if no gap is forthcoming, the 

seedlings and saplings die. Of the secondary species 

noted above, all have small seeds, except Pometia 

pinnata. Pometia, however, is the dominant tree in 

one kind of lowland forest-that found on relatively 

recent lava flows.

Two species that appear to be common in both 

secondary and primary forest are Dysoxylum maota 

(maota) and Dysoxylum samoense (mamala). On 

the 'Tafuna plain, when forest was still present there, 

the two Dysoxylum species appeared to be succes- 

sional species that replaced Macaranga harveyi and 

in turn was replaced, or shared dominance with, 

Pomctiu pinnata and Planchonella sarnoensis (ma- 

malava). In alluvial valleys, the two Dysoxylum 

species seem to be the dominant species in the cli- 

max forest for this habitat, judging by the population 

dynamics in plots of this on Tutuila and Ta'u 

(Whistler 1980). 

Another category of plants often useful as indi- 

cators of past disturbance comprises cultivated 

Polynesian introductions. The coconut mentioned 

above would also fit into this category, except that it 

is probably native; it otherwise fits the general 

characteristics. Trees and shrubs that belong here 

are Aluna rucemosa (ifiifi), Citrus macroptera (mob 

u'u), Diospyros major, Euodia hortensis (usi), 

lnocarpus fagifer (Tahitian chestnut, ifi), Parinari 

insularum (sea), and Syzygium corynocarpum 

(seasea), Syzygium rnalaccensis (Malay apple, nonu 

fi'afi'a), Syzygium neurocalyx (oli, fena). Ino- 

curpus is now common and naturalized along 

streams. The others are probably not naturalized, 

and individuals found in forest are probably relicts 

of previous cultivation. Further information about 

these species can be found in Appendix A. 

Another species that partly fits here is Cerbera 

manghas, a littoral tree that is sometimes found 

inland where it presumably was planted (for its 

showy flowers). It was found on the ridge just west 

of Vatia in mature forest. Also found in this area 

was Syzygium corynocarpum. 

Several non-woody cultigens are also sometimes 

found in forest appearing to be primary. This 

includes Alocasia macrorrhiza (giant taro, ta'amu), 

Amorphophallus paeoniifolius (teve), Etlingera 

cevuga ('avapui vao), Schizostachyum glaucifolium 

(Polynesian bamboo, 'ofe), Solanum viride (polo- 

'iti), and Zingiber zerumbet ('avapui). Alocasia is 

not uncommon in forest and in streambeds, Etlin- 

gera, which may be native, was found once in the 

park, Schizostachyum is reported from the park (R. 

West, pers. comm.), and Solanum viride was col- 

lected there once. Zingiber is sometimes common 

along ridges, and may have been present in these 

places and has subsequently spread, since it does not 

appear to be easily dispersed.

APPENDIX F. SUMMARY OF THE VASCULAR FLORA OF THE TUTUILA 

UNIT OF THE NATIONAL PARK OF AMERICAN SAMOA. 

Species FAMILY Status* E I P M 

ADIANTIACEAE 

Acrostichum aureum 

Adiantum philippense 

Bolbitis palustris 

Pityrogramma brackenridgei 

Pteris comans 

Pteris ensiformis 

Pteris pacifica 

Pteris tripartita 

ASPIDIACEAE 

Arachniodcs aristata 

Ctenitis samoensis 

Pleocnemia leuzeana 

Tectaria sctchellii 

Tectaria stearnsii 

ASPLENIACEAE 

Asplenium auslralasicum 

Asplenium cuneatum 

Asplenium horridum 

Asplenium laserpitiifolium 

Asplenium nidus 

Asplenium polyodon 

Asplenium tenerum 

ATHYRIACEAE 

Diplazium bulbiferum 

Diplazium dilatatum 

Diplazium harpeodes 

Diplazium proliferum 

BLECHNACEAE 

Blechnum orientale 

Blechnum vulcanicum 

CYATHACEAE 

Cyathea decurrens 

Cyathea lunulata 

Cyathea vaupelii 

DAVALLIACEAE 

Arthropteris repens 

Davallia epiphylla 

Davallia solida 

Humata heterophylla 

Humata polypodioides 

Nephrolepis biserrata 

Nephrolepis hirsutula 

Oleandra neriiformis 

DENNSTAEDTIACEAE 

Orthiopteris tenuis 

FERNS

GLEICHENIACEAE 

Dicranoptcris linearis 

GRAMMITIDACEAE 

Ctcnoptcris blechnoides 

Ctenopteris contigua 

HYMENOPHYLLACEAE 

Hymenophyllum imbricatum 

Hymenophyllum polyanthos 

Trichomanes bipunctatum 

Trichomanes boryanum 

Trichomanes dentaturn 

Trichomanes humile 

Trichomanes intermedium 

Trichomanes saxifragoides 

Trichomanes tahitense 

HYPOLEPIDACEAE 

Histiopteris incisa 

LINDSAEACEAE 

Lindsaea ensifolia 

Lindsaea harveyi 

Lindsaea tetragona 

Microlepia speluncae 

LOMARIOPSIDACEAE 

Elaphoglossum feejeense 

Lomagramma cordipinna 

MARATTIACEAE 

Angiopteris evecta 

Angiopteris opaca 

Marattia smithii 

OPHIOGLOSSACEAE 

Ophioglossum pendulum 

POLYPODIACEAE 

Microsorium sylvaticum 

Phymatosorus nigrescens 

Phymatosorus scolopendria 

Polypodium subauriculatum 

Pyrrosia lanceolata 

SCHIZAEACEAE 

Schizaea dichotoma 

THELYPTERIDACEAE 

Christella dentata 

Christella harveyi 

Macrothelypteris polypodioides 

Macrothelypteris torresiana 

Plesioneuron attenuatum 

Pneumatopteris bryanii 

Pneumatopteris glandulifera 

Sphaerostephanos unitus

VIlTAKIACEAE 

Amrophyurn alaturn 

Antrophyurn plantagineum 

Vaginularria angustissirna 

Viltaria elongata 

Vittaria scolopendrina 

LYCOPODIACEAE 

Lycopodium carinatum 

Lycopodiurn cernuum 

Lycopodium phlegmaria 

Lycopodium squarrosum 

SELAGINACEAE 

Selaginella laxa 

Selaginella whitmeei 

ACANTHACEAE 

Blechum brownei 

Justicia procumbens 

Ruellia prostrata 

AMAKANTHACEAE 

Alternanthera sessilis 

ANACARDIACEAE 



ANNONACEAE 

Cananga odorata 

APIACEAE 

Centella asiatica 

APOCYNACEAE 

Alstonia pacifica 

Alyxia bracteolosa 

Alyxia stellata 

Cerbera manghas 

ARALIACEAE 

Meryta rnacrophylla 

Polyscias samoensis 

ARISTOLOCHIACEAE 

Aristolochia cortinata 

ASCLEPIACACEAE 

Hoya australis 

Hoya pottsii 

Hoya vitiensis 

ASTERACEAE 

Achrnella uliginosa 

Ageratum conyzoides 

Bidens alba 

Bidens pilosa 

Crassocephalum crepidioides 

FERN ALLIES 

DICOT ANGIOSPERMS

Species FAMILY 

ASTERACEAE (cont 'd) 

Emilia sonchifolia 

Erechtites valerianifolia 

Mikania micrantha 

Synedrella nodiflora 

Vernonia cinerea 

Wollastonia biflora 

Youngia japonica 


Barringtonia asiatica 


BORANGINACEAE 

Cordia aspera 

BURSERACEAE 


Canarium vitiense 

Garuga floribunda 

CARICACEAE 

Carica papaya 

CASSYTHACEAE 

Cassytha filiformis 

CELASTRACEAE 

Maytenus vitiensis 

CHRYSOBALANACEAE 

Atuna racemosa 

CLUSIACEAE 

Calophyllum inophyllum 



Mammea glauca 

COMBRETACEAE 


Terminalia richii 

CONNARACEAE 

Rourea minor 

CONVOLVULACEAE 

Ipomoea littoralis 

Ipomoea macrantha 

Ipomoea pes-caprae 

Merremia umbellata 

Operculina ventricosa 

Stictocardia tiliifolia 

CUCURBITACEAE 

Momordica charantia 

Zehncria rnucronata 

Zehneria samoensis 

CUNONIACEAE 

Spiraeanthemum sarnoense 

Weinmannia affinis


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

EBENACEAE 

Diospyros elliptica 




Elaeocarpus ulianus 

EUPHORBIACEAE 

Aleurites moluccana 

Antidesma sphaerocarpum 

Baccaurea taitensis 


Chamaesyce hirta 

Chamaesyce hypericifolia 

Chamaesyce hyssopifolia 

Chamaesyce thymifolia 

Drypetes vitiensis 

Glochidion cuspidatum 

Glochidion ramiflorum 

Macaranga harveyana 

Macaranga stipulosa 

O~nalanthus nutans 

Phyllanthus amarus 

Phyllanthus debilis 

Phyllanthus urinaria 

Phyllanthus virgatus 

FABACEAE 

Abrus precatorius 

Adenathera pavonina 

Alysicarpus vaginalis 

Caesalpinia major 

Calopogonium muconoides 

Canavalia cathartica 

Canavalia rosea 

Dendrolobium umbellaturn 

Derris trifoliata 

Desmodium incanum 

Desmodium tortuosum 

Desmodium triflorum 


lnocarpus fagifer 

lntsia bijuga 

Leucaena lcucocephala 

Macroptilium lathyroides 

Mimosa invisa 

Mimosa pudica 

Mucuna gigantea 

Paraserianthes falcataria 

Pueraria lobata 

Vigna marina



Erythrospermum acuminatis imum 

Flacourtia rukam 

Homalium whitmeeanum 

GESNERIACEAE 

Cyrtandra pulchella 

Cyrtandra samoensis 

GOODENIACEAE 

Scaevola taccada 

HERNANDIACEAE 

Hernandia moerenhoutiana 

Hernandia nymphaeifolia 

ICACINACEAE 

Citronella samoensis 

LAURACEAE 

Litsea samoensis 

LOGANIACEAE 



LORANATHACEAE 

Decaisnina forsteriana 

LYTHRACEAE 

Cuphea carthagenensis 

MALVACEAE 


Sida rhombifolia 

Thespesia populnea 

MELASTOMACEAE 

Astronidium pickeringii 

Clidemia hirta 

Dissotis rotundifolia 

Medinilla samoensis 

Melastoma denticulatum 

MELIACEAE 

Aglaia samoensis 


Dysoxylum maota 


MONIMIACEAE 

Hedycarya denticulata 

MORACEAE 

Ficus godeffroyi 

Ficus obliqua 

Ficus prolixa 

Ficus scabra 

Ficus tinctoria 

Ficus uniauriculata 

MYRISTICACEAE 


Myristica hypargyraea

MYRSINACEAE 

Embelia vaupelii 

Maesa tabacifolia 

MYKTACEAE 

Descaspcrmum fruticosum 

Metrosideros collina 

Psidium guajava 

Syzygium brevifolium 

Syzygium carolinense 

Syzygium clusiifolium 


Syzygium inophylloides 



NYCTAGINACEAE 

Boerhavia albiflora 

Boerhavia repens 

Pisonia grandis 

Pisonia umbellifera 

OIXACEAE 

Chionanthus vitiensis 

Jasminum didymum 

ONAGRACEAE 

Ludwigia hyssopifolia 

Ludwigia octovalvis 

OXALIDACEAE 

Oxalis corniculata 

PASSIFLORACEAE 

Passiflora foetida 

PIPERACEAE 

Peperomia pellucida 

Peperomia tutuilana 

Piper graeffei 

POLYGALACEAE 

Polygala paniculata 

PORTULACACEAE 

Portulaca samoensis 

RHAMNACEAE 

Alphitonia zizyphoides 

Colubrina asiatica 

KHIZOPHORACEAE 

Crossostylis bifora 

KUBIACEAE 

Calycosia sessilis 


Cyclophyllum barbatum 

Geophila repens 


Gynochtodes epiphytica 

Hedyotis biflora


RUBIACEAE (cont'd) 

Ixora samoensis 


Morinda myrtifolia 

Mussaenda raiateensis 


Psychotria forsteriana 

Psychotria insularum 

Sarcopygme pacifica 

Spermacoce assurgens 

Spermacoce mauritiana 

Tarenna sambucina 

RUTACEAE 

Acronychia heterophylla 

Euodia hortensis 

Euodia samoensis 

Micromclum minutum 

SAPINDACEAE 

Allophylus timoriensis 

Arytera brackenridgei 

Elattostachys falcata 

Sapindus vitiensis 

SAPOTACEAE 

Manilkara dissecta 

Palaquium stehlinii 


Planchonella grayana 

Planchonella samoensis 

SCROPHULARIACEAE 

Lindernia crustacea 

STERCULIACEAE 

Kleinhovia hospita 


THEACEAE 

Eurya pickeringii 

'11 IYMELAEACEAE 

Phaleria glabra 

Wikstroemia foetida 

TILIACEAE 

Grewia crenata 

Trichospermum richii 

ULMACEAE 

Trema cannabina 

URTICACEAE 

Cypholophus macrocephalus 

Dendrocnide harveyi 

Leucosyke corymbulosa 

Pilea microphylla 

Pipturus argenteus 

Procris pedunculata


VEKBENACEAE 

Clcrodendruni chinense 

(:lcrotlendrum incrme 

Faratlaya amicorum 

Premna scrratifolia 

Stachytarpheta urticaefolia 

MONOCOT ANGIOSPERMS 

AGAVACEAE 

Cordyline fruticosa 

ARACEAE 

Amorphophallus paeoniifolius 

Epipremnum pinnatum 

Rhapidophora graeffei X 

ARECACEAE 

Cocos nucifera 

COMMELINACEAE 

Commelina diffusa 

CYPERACEAE 

Cyperus compressus 

Fimbristylis autumnalis 

Fimbristylis cymosa 

Fimbristylis dichotoma 

Kyllinga brevifolia 

Kyllinga nemoralis 

Mapania macrocephala 

Mariscus cyperinus 

Mar iscus javanicus 

Mariscus seemannianus 

Pycreus polystachyos 

Scleria lithosperma 

Scleria polycarpa 

DIOSCOREACEAE 

Dioscorea bulbifera 

NELICONIACEAE 

Heliconia laufao X 

OHCHIDACEAE 

Appendicula bracteosa 

Bulbophyllum longiscapum 

Dulbophyllum polypodioides 

Bulbophyllum rosticeps 

Bulbophyllum sp. nova X 

Calanthe triplicata 

Coelogyne lycastoides 

Dendrobium biflorum 

Dendrobium calcaratum 

Dendrobium dactylodes 

Dendrobium goldfinchii 

Dendrobium samoense 

Dendrobium sladei

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


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

ORCHIDACEAE (cont 'd) 

Didymoplexis micradenia 

Diplocaulobium fililobum 

Eria robusta 

Flickingeria comata 

Glomera montana 

Liparis condylobulbon 

Malaxis resupinata 

Malaxis samoensis 

Moerenhoutiana heteromorpha 

Oberonia equitans 

Oberonia heliophila 

Phreatia matthewsii 

Phreatia micrantha 

Phreatia obtusa 

Spathoglottis plicata 

Taeniophyllum fasciola 

Taeniophyllum gracile 

Thrixspermum graeffei 

PANDANACEAE 

Freycinetia reineckei 

Freycinetia storckii 

Pandanus tectorius 

POACEAE 

Axonopus compressus 

Bolhriochloa bladhii 

Brachiaria mutica 

Brachiaria paspaloides 

Brachiaria subquadripara 

Centosteca lappacea 

Chloris barbata 

Coix lacryma-jobi 

Cyrtococcum oxyphyllum 

Dacytloctenium aegyptium 

Digitaria ciliaris 

Digitaria horizontalis 

Digitaria setigera 

Digitaria violascens 

Echinochloa colona 

Eleusine indica 

Eriochloa procera 

lschaemum murinum 

Lepturus repens 

Miscanlhus floridulus 

Oplismenus compositus 

Oplismenus hirtellus 

Paspalum conjugatum 

Paspalum orbiculare 

Paspalurn paniculatum 

Paspalurn vaginaturn

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


POACEAE (cont'd) 

Sporobolus diander 

Stenotaphrum micranthum 

Thuarca involuta 

TACCACEAE 

Tacca lcontopctaloides 

ZINCIBERACEAE 

Costus spcciosa 

Etlingera cevuga 

Zingibcr zerumbet 

E = Endemic; I = Indigenous; P = Polynesian introduction; M = Modern Introduction.

american samoa - University of Hawaii at Manoa

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