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F. RAYMOND FOSBERG (1908-1993)
COMMEMORATIVE ISSUE

Issued by
NATIONAL MUSEUM OF NATURAL HISTORY

SMITHSONIAN INSTITUTION
WASHINGTON, D.C. U.S.A.
FEBRUARY 1994

ATOLL RESEARCH BULLETIN NOS. 390-396

RESEARCH
BULLETIN

ATOLL RESEARCH BULLETIN

NOS. 390-396

NO.

NO.

NO.

NO.

NO.

NO.

NO.

RAYMOND FOSBERG (1908-1993) COMMEMORATIVE ISSUE

390.

391.

392.

393.

394.

395.

396.

LISTS OF TAXA NAMED FOR F. RAYMOND FOSBERG AND
BY HIM
BY DAN H. NICOLSON

ANNOTATED LIST OF THE KNOWN PUBLICATIONS OF
F. RAYMOND FOSBERG
BY ANNA L. WEITZMAN

THE FLORA OF NAURU
BY R.R. THAMAN, F.R. FOSBERG, H.I. MANNER, AND
D.C. HASSALL

FLORA OF THE PHOENIX ISLANDS, CENTRAL PACIFIC
BY F. RAYMOND FOSBERG AND DAVID R. STODDART

THE HOA OF HULL ATOLL AND THE PROBLEM OF HOA
BY DAVID R. STODDART AND F. RAYMOND FOSBERG

STORY OF AN OCEANIC ARCHIPELAGO
BY F. RAYMOND FOSBERG

COMMENTS ON ATOLL PHOSPHATE ROCK
BY F. RAYMOND FOSBERG

ISSUED BY
NATIONAL MUSEUM OF NATURAL HISTORY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C., U.S.A.
FEBRUARY 1994

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ACKNOWLEDGMENT

The Atoll Research Bulletin is issued by the Smithsonian Institution to
provide an outlet for information on the biota of tropical islands and reefs
and on the environment that supports the biota. The Bulletin is supported by
the National Museum of Natural History and is produced by the Smithsonian
Press. This issue is partly financed and distributed with funds from readers.

The Bulletin was founded in 1951 and the first 117 numbers were issued by
the Pacific Science Board, National Academy of Sciences, with financial
support from the Office of Naval Research. Its pages were devoted largely to
reports resulting from the Pacific Science Board’s Coral Atoll Program.

All statements made in papers published in the Atoll Research Bulletin
are the sole responsibility of the authors and do not necessarily represent
the views of the Smithsonian nor of the editors of the Bulletin.

Articles submitted for publication in the Atoll Research Bulletin should
be original papers in a format similar to that found in recent issues of the
Bulletin. First drafts of manuscripts should be typewritten double spaced and
can be sent to any of the editors. After the manuscript has been reviewed and
accepted, the author will be provided with a page format with which to prepare
a single-spaced camera-ready copy of the manuscript.

COORDINATING EDITOR

Ian G. Macintyre National Museum of Natural History
MRC-125
Smithsonian Institution
Washington, D.C. 20560

EDITORIAL BOARD

Stephen D. Cairns (MRC-163) National Museum of Natural History
Brian F. Kensley (MRC-163) (Insert appropriate MRC code)

Mark M. Littler (MRC-166) Smithsonian Institution

Wayne N. Mathis (MRC-169) Washington, D.C 20560

Victor G. Springer (MRC-159)

Joshua I. Tracey, Jr. (MRC-137)

Warren L. Wagner (MRC-166)

Roger B. Clapp (MRC-111) National Museum of Natural History

National Biological Survey
Smithsonian Institution
Washington, D.C. 20560

David R. Stoddart Department of Geography
501 Earth Sciences Building
University of California
Berkeley, CA 94720

Bernard M. Salvat Laboratoire de Biologie & Ecologie
Tropicale et Méditerranéenne
Ecole Pratique des Hautes Etudes
Labo. Biologie Marine et Malacologie
Université de Perpignan
66025 Perpignan Cedex, France

F. Raymond Fosberg Ohio 1968
Photograph by H. Gray Multer

F. RAYMOND FOSBERG, 1908-1993

It is with deep sadness and regret that we record the passing of our
Founding Editor Dr F. Raymond Fosberg, at his home in Falls Church,
Virginia, on 25 September 1993. He was 85 years old, and had been seriously
ill with cancer for several months.

Ray occupied a very special place in the development of coral reef and
island studies in the twentieth century. Through the Afoll Research Bulletin he
helped create an autonomous branch of science, known now for its national
and international meetings, its public importance, and its diverse and
dynamic research programs. The Bulletin was and remains central in these
endeavors.

His career in the Pacific effectively began when he took an assistantship
in the Department of Botany at the University of Hawaii with Professor
Harold St John in 1932. He had gained his B.A. at Pomona College in 1930,
and during 1930-1932 served as Botanist at the Los Angeles County Museum,
working on the floras of the southwestern United States and northern
Mexico. In 1934 he joined St John on the Bishop Museum’s Mangarevan
Expedition, led by the malacologist C. Montague Cooke, Jr. This took him
from Hawaii to Fanning in the Lines, Tahiti and Mehetia in the Societies, to
the atolls of Anaa, Tepoto, Napuka, Fangatau, Hao and Vahitahi in the
Tuamotus, Mangareva in the Gambiers, Pitcairn and nearby Henderson and
Oeno in southeastern Polynesia, to the islands of the Australs from Marotiri
to Maria, and back to Hawaii by way of Raiatea, Borabora, Vostok, Flint,
Christmas and Fanning. In total the expedition visited 25 high islands and 31
coral islands; St John and Fosberg brought back 15,000 plant specimens. It was
an extraordinary introduction to the island world of the tropical Pacific, and it
determined Ray’s interests for the rest of his life.

Following the expedition Ray gained his M.S. in 1935 at the University
of Hawaii with a monograph on the Hawaiian genus Gouldia (Rubiaceae), and
then moved to the University of Pennsylvania, taking his Ph.D. in 1939 with
a thesis on the Polynesian species of Hedyotis in the same family. Then for
several years during and after World War II he worked in South America on
the medically important genus Cinchona, also in the Rubiaceae. But he
returned to Pacific island studies in 1946 when he spent six months working
on an Economic Survey of Micronesia for the U.S. Commercial Company. By

il

the end of the 1940s he was deeply immersed in the study of the vegetation
and floristics of Pacific islands.

This coincided with the development of the Coral Atoll Program of the
Pacific Science Board, beginning with an expedition to Arno Atoll in the
Marshalls in 1950 and 1951. Ray was responsible for founding the Atoll
Research Bulletin in 1951 as an outlet for the results of the Pacific Science Board
expeditions. It became the major repository for information on coral reefs
and islands, especially their terrestrial aspects, and Ray’s first paper in the
original issue was reprinted in the Bulletin in 1992. Ray was always deeply
respectful of the director of the Pacific Science Board, Dr Harold Jefferson
Coolidge—at least to me he always called him ‘Mr Coolidge’—who in
addition to the Coral Atoll Program was also responsible for the programs
Coordinated Investigations in Micronesian Anthropology and Scientific
Investigations in Micronesia, as well as being a leading figure in the Pacific
Science Association and the International Union for the Conservation of
Nature and Natural Resources. Ray wrote an appreciation of him in the
Bulletin in 1985.

Virtually from the start of the Coral Atoll Program Ray was assisted by
Dr Marie-Héléne Sachet. Together they built up a formidable data base on
island biogeography, as well as an unprecedented collection of publications,
photographs and reports, now in the Smithsonian Institution. Resulting
from this work they published the massive Island Bibliographies and its
Supplement in 1955 and 1971. Marie-Héléne was largely responsible for the
practical editing and production of the Bulletin, especially in the early days
when it was a much more time-intensive operation than it is now. Her own
thesis had been on the natural history of Clipperton Atoll, but in 1963 she
worked in the Societies and on Rangiroa, in 1963 and 1974 in the Marquesas,
and in 1982 and 1983 especially in the high Society Islands, on Tetiaroa, and in
the Tuamotu. Many of her papers on this work appeared in the Bulletin (nos.
274, 275, 276, 277, 294). Her sudden death in 1986 was a severe blow to Ray,
since there were still so many major projects to be completed. Ray wrote a
detailed appreciation of their collaboration in a Commemorative Issue of the
Bulletin in 1987, and there is also a very perceptive notice of her by R. Koenig
in the Bulletin of the Société des Etudes Océaniennes [Tahiti] in the same year.

Their work, together with that of Royce Oliver in the Department of
Botany at the National Museum of Natural History, led to the codification of
an enormous body of knowledge, especially in their geographical checklists of
the plants of Micronesia published in 1976, 1982 and 1987, and in a continuing
project on the Flora of Micronesia (1975-1993). In collaboration with Steve
Renvoize of the Royal Botanic Gardens, Kew, Ray also published the Flora of

iii

Aldabra and nearby islands in 1980, and co-edited with M. D. Dassanayake the
Revised Handbook of the Flora of Ceylon, of which seven volumes have been
published since 1980. The Vascular flora of the northern Mariana Islands
appeared in 1975. Ray also put in order the Partial flora of the Society Islands
(1974) based on the large collections and voluminous notes assembled by his
friend Martin Grant in 1930-31: Ray had been deeply affected by Grant’s death
in 1968 before any of his wealth of knowledge had been published, and I think
this spurred him on over the last quarter of a century to ensure that it would
not happen to himself.

The complete bibliography of Ray’s publications in this issue includes
many floristic papers on reef islands over an immense biogeographic range.
In the Bulletin these include plant lists for the Marshall Islands (1955, 1990),
the Gilbert Islands (1987), Henderson Island (1983), Wake Island (1959, 1969),
and Nauru and the Phoenix Islands (this issue), as well as many other Pacific
localities; of the northern Great Barrier Reef (1991); of the Maldive Islands
(1957, 1966), Agalega (1983), Cousin (1983), and many other western Indian
Ocean islands (1970, 1983); and of Alacran (1962), the Dry Tortugas (1981), the
Belize cays (1982), and the Jamaican cays (1991) in the western Atlantic. As his
bibliography shows, this list is very far from complete. Over his lifetime Ray
collected more than 66,000 plant numbers in multiple sets, and probably
carried out and published more scientific work on more tropical islands, both
high and low, around the world than anyone else. Sadly he did not live to
see the publication of a final synthesis with Dieter Mueller-Dombois,
Vegetation of the tropical Pacific islands, though a precursor summary on the
Society Islands did appear in 1992.

While the major thrust of his work was floristic and taxonomic, Ray
had an enormously wide range of interests in natural history broadly
conceived—in geomorphology, diagenesis, soils, birds, biogeography, ecology,
conservation, geography. He wrote originally and trenchantly on all of these
topics, and it is clear that these more general writings derived from deep
conviction as well as scientific expertise. Though he spent his life on the
tropical oceans and their islands, Ray came from the land: his youth was
spent on the family melon-growing enterprise in California. I think he
learned the lessons then of environmental uncertainty, accountability, and
practical conservation.

He was not deeply interested in theory, and in fact distrustful of it. He
was a pragmatic and realistic recorder of empirical nature in all its
unpredictable variety. He tried to bring order into this complexity by
classifying and naming and by studying distributions. He belonged as an

iv

interpreter of the island world with the founders of the discipline—Darwin,
Hooker, Wallace—and used much the same methods.

In addition to the taxonomic and floristic papers—the building blocks
of his life work—Ray wrote a number of influential interpretative papers.
One of the earliest and still most widely cited was his ‘Derivation of the flora
of the Hawaiian Islands’ in volume 1 of Zimmerman’s Insects of Hawaii (1948),
which took an uncompromisingly dispersalist position. Based on an analysis
of nearly two thousand species of seed plants and ferns in over 250 genera, he
concluded that “the flora has always been insular. It is exactly the type that
might be expected to be descended from a random aggregation of chance waifs
carried overseas by a combination of factors such as storms, currents and
birds. Of seed plants, an average of but one successful arrival and
establishment every 20,000 to 30,000 years would account for the flora” (1948,
p- 119). This was robust material, especially in an age and a subject dominated
by Skottsberg and Corner, Darlington and Simpson. Another landmark paper
was his ‘Qualitative description of the coral atoll ecosystem’ (1961), which
sought to provide a framework for the diversity of data accumulating at a
rapidly accelerating rate. A third (which appealed to me) was his address on
‘Geography, ecology, and biogeography’ to the Association of American
Geographers in 1976, in which he forcefully informed geographers what they
really ought to be about .

There was one major project that he did not get completed in his
lifetime, and in many respects it was one of the more intriguing ones. From
the outset in his Pacific studies Ray had been concerned by the problem of the
botany of Cook’s voyages. He had toiled to make Merrill’s somewhat
eccentric volume on that subject publishable. He wrote on Parkinson and
Banks long before the Florilegium was finally published. But from the time of
the Mangarevan Expedition he became especially concerned with the
collections made by Johann Reinhold and Georg Forster on Cook’s second
voyage. For decades our home in Cambridge was a way station for Ray en
route to herbaria in Leningrad, Paris, Berlin, and up to two dozen other
places, where he indefatigably tracked down the widely dispersed Forster
collections from the Pacific; his intention was to lectotypify species which had
been inadequately established. The work is virtually complete, and indeed
Ray saw the proof of a precursor paper to Allertonia only a day before he died.
He was also interested in Robert Brown’s collections on the Flinders
Expedition, and re-collected at Brown’s sites in the Gulf of Carpentaria, but I
do not think he published on Brown’s material.

Ray was a large, powerfully built, imposing man—tall and handsome,
as Clifford Gessler described him at the time of the Mangarevan Expedition,

hough sadly increasingly stooped in later years. He had fairly craggy features,

a rather shy and disarmingly boyish smile, often a twinkle in clear blue eyes
which never needed glasses; indeed the only aid he needed was the
trademark hand lens he always wore around his neck. His speech was slow
and deliberate, his convictions clear, his distaste for some more recent ideas
not disguised. Both he and Marie-Héléne could be severe critics of work that
did not reach their own exacting standards, as I have reason to recall. He was
a man of some stubbornness and considerable physical determination (he
took up SCUBA, for example, when well into his sixties).

I first met him more than thirty years ago when I was just starting
work on the sand cays of Belize. E. J. H. Corner of Cambridge, one of tropical
botany’s memorable figures, told me to seek Ray out. I went to Washington,
and Ray and Marie-Héléne came to see me in the lobby of the old Raleigh
Hotel. Somehow I had the premonition that it was to be a very important
meeting for me. Something worked that day, in spite of the fact that I had
very little idea of what I was about. Indeed I recall with embarrassment the
state of the earliest collections I sent him. But he persevered. I have seen that
pattern of support repeated so many times since: Ray never grudging his
time or his advice to those he thought he could help. He did this to educate
people, especially students; to advance science in the tropical seas; and to
build the Smithsonian’s plant collections. He helped unknown and
beginning students and members of the general public as well as his scientific
peers and friends. Indeed he sometimes exasperated me by the way he
allowed himself to be deflected from the task in hand during long-planned
visits to overseas herbaria when people learned he was in town and
descended on him.

This is not the place to record Ray’s unremitting work for
conservation, but some instances of his powerful initiatives in the atoll realm
must be mentioned. From the middle 1950s he urged the preservation of
Aldabra, the western Indian Ocean elevated atoll renowned for its huge
population of giant tortoises, and he played a central rdle in the United States
in orchestrating the opposition to plans to turn it into a military base in 1965-
1967. Less well known were his successful efforts to limit the impact of U.S.
missile testing programs in the Phoenix Islands in the early 1970s. And he
was central—with his unique knowledge—in the campaign to preserve
Henderson Island in southeastern Polynesia from bizarre plans in the late
1970s and early 1980s to turn it into a private refuge from nuclear attack. The
issue of the Bulletin (321-329, 1989) recording the first results of the new wave
of Henderson studies carries an appreciation by Ray of the work of the former
Secretary of the Smithsonian, S. Dillon Ripley—again, always ‘Mr Ripley’ to
Ray—who had appointed him to the Institution and who had a particular

interest in the Henderson and Aldabra Rails. Ray pursued these causes to the
highest political levels with tenacity and immense authority, and in each case
the cause was won. Ray also advised Mr Marlon Brando over many years on
his management of Tetiaroa Atoll in the Societies.

Ray was both facilitator and communicator in international science.
He attended the Pacific Science Congresses between 1949 and 1991, and was
active in Pacific Science Association committees and in organizing symposia
and fieldtrips. The book he edited following the symposium he organized at
the Honolulu Congress in 1961 on Man’s Place in the Island Ecosystem became a
classic. The theme was re-examined at the Vancouver Congress in 1975 but
the results were unfortunately never published. At his last Congress in
Honolulu he gave a plenary lecture, published in the Bulletin in 1992, as well
as a contributed paper, and met many friends and colleagues for what was to
be the last time. He was one of that small group which founded the
International Coral Reef Symposia at Mandapam in India a quarter of a
century ago. He was at the Second Symposium for the memorable cruise
along the Great Barrier Reef in 1973. Both he and Marie-Héléne were on
home ground for the spectacular 5th meeting in Tahiti in 1985. Ray went off
on the slightly alarming Makatea fieldtrip with Josh Tracey and others
afterwards, and Marie-Héléne went to Takapoto. Both were in their element
at that meeting, and no-one could have foreseen that we should lose both of
them so soon. Ray naturally became an honorary member of the
International Society for Reef Studies when that was founded. He was an
active member of the International Botanical Congresses from Stockholm
(1950) to Berlin (1988). He was a major figure in the UNESCO Humid Tropics
Program, in the International Society for Tropical Ecology, the U.S. Nature
Conservancy, and the International Biological Program. He attended all of
Nicholas Polunin’s International Conferences on Environmental Futures.

Many honors came to him: he was truly touched by such recognition
but did not need it. He received the George Davidson Medal and the Edward
W. Browning Achievement Award. He particularly appreciated the Robert
Allerton Medal from the Pacific Tropical Botanical Garden on Kauai and the
Herbert E. Gregory Medal from the Pacific Science Association. He was glad
that his wider messages were recognised by the award of the Daly Medal by
the American Geographical Society. Pomona College, where he did his
undergraduate work and to which he left his personal plant collection,
awarded him an honorary degree in 1980, as did the University of Guam in
1986, the University of Peradeniya, Sri Lanka, in 1986, and the University of
the South Pacific in Fiji in 1989. He much appreciated this academic
recognition.

Toward the end of his life there were three public celebrations in which
he delighted. The first was a meeting at Bishop Museum in Honolulu on
March 23, 1984, for the 50th anniversary of the Mangarevan Expedition. It
was attended by Harold St John, Kenneth P. Emory, Yoshio Kondo and
Donald Anderson. C. Montague Cooke, Jr., Peter H. Buck and J. Frank
Stimson had already passed away, and Ray was in fact the last survivor of
those who met again in 1984. Only E. C. Zimmerman can now tell us about
that historic expedition. Ray was also honored by a special symposium at the
INTECOL meeting in Yokohama in 1990. This was published as a special
issue of Pacific Science, to which Ray himself contributed, in 1992.

Finally the editors of the Bulletin assembled a special 40th anniversary
issue, without Ray’s knowledge, and this appeared in 1992. In November that
year the Director of the National Museum of Natural History, Frank Talbot,
himself a coral reef specialist who had also been at Mandapam, hosted a
reception in his office at which the issue was presented to Ray. There is a
note with photographs in the News and Comment section of issue 379-389 in
1993. Among those present were long-time associates Josh Tracey and Frank
Whitmore; his colleague Royce Oliver; and Myron M. Weinstein, the Hebraic
scholar from the Library of Congress, a friend over very many years to both
Ray and Marie-Héléne. Ray was already very frail, but he gave a powerful
and moving address. It was his last public appearance, though he continued
to go to his office in the Department of Botany until August of 1993.

It is sad indeed that so soon after the 40th Anniversary Issue we
collectively recall the life, work, personality and achievements of F. Raymond
Fosberg. We do so through a Commemorative Issue of the journal he
founded and nurtured. This issue records his remarkable contribution to
knowledge of the natural world, through his publications and through the
taxa which he named and which are named for him. We publish also a
number of papers which Ray left in final form, including the Flora of Nauru
and his Sauer Memorial Lecture, the first papers from his Phoenix Islands
investigations, and a final commentary on his phosphate rock hypothesis.
His lifetime’s collection of plants will form an enduring monument in
herbaria around the world, and especially in the U.S. National Herbarium in
Washington, D.C.

These will not be the last of Ray’s publications in the Bulletin. There are
other substantial works near completion, as well as the monograph on the
Forsters. The encyclopaedic dictionary of reef terminology which he
completed years ago with Rhodes Fairbridge and many others is now again
accessible. There is a checklist of Indian Ocean reef island plants similar to
those he prepared on Micronesia.

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The Atoll Research Bulletin will stand as one of Ray’s chief memorials,
not only for the information it records but for the way it has served to codify
and institutionalize the emergent discipline of coral reef and island studies
during Ray’s professional life. Mary McCutcheon’s exemplary and
comprehensive Index to the Bulletin in 1991 gives a measure of that
achievement. In thinking of Ray and the difference he has made to all of us,
it is worth reflecting not only on the status and scope of reef and island
science when the Bulletin was founded in 1951, but also on what it was like
when Ray embarked on the Mangarevan Expedition exactly sixty years ago.

This introductory note is very far from being a comprehensive
treatment of Ray’s life and work. Others will speak with more authority than
I can on many interests that he had. It has not been an easy notice to write,
largely because I knew him for too long and too well for that to be possible.
Both Ray and Marie-Héléne entered into the folklore of our family, as we
doubtless did into theirs. Both were true friends.

There is a fine appreciation of Ray by Dieter Mueller-Dombois in the
INTECOL Symposium issue of Pacific Science in 1992. There is also a
substantial notice of him by Steve Renvoize, his co-author on the Aldabra
flora, in the London Independent for 14 October 1993.

David R. Stoddart

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The F. Raymond Fosberg Fund

Friends and colleagues of Ray Fosberg (1908-1993) are establishing The
F. Raymond Fosberg Fund in memory of the man who was the foremost
student of Pacific botany and vegetation of his time and a leading
conservationist. Extensive travel provided him with the broadest concepts of
world patterns of geography, vegetation, coral reefs, and islands and their life,
a view surpassing that of most scientists. He wrote works (more than 800) on
the plants of Hawaii, Micronesia, Aldabra, and many other places, that are the
basis of our present knowledge. He advised and helped innumerable younger
colleagues in developing their careers and pioneered the ethic to conserve the
plants and animals of the world. This fund is established in his memory, and
with the hope that through it his efforts for Pacific island botany and
conservation can be furthered. The fund will be administered by The Nature
Conservancy of Hawaii, a non-profit conservation organization, to support
the protection of native Hawaiian forests and the plants and animals which
depend on them for their survival.

Of all the places on Earth, Hawaii has the most alarming concentration
of species teetering on the brink of extinction. Hawaii amounts to only 0.2%
of the land area of the United States. Yet nearly 75% of the nation’s
historically documented plant and bird extinctions are from Hawaii. And of
all the bird species currently endangered in the 50 United States, 40% are from
Hawaii. Hawaiian plant species account for 34% of the list of endangered
plants in the nation. The Nature Conservancy of Hawaii is working to
protect the forests that these remaining native plants and animals need to
survive.

If you wish to give to The F. Raymond Fosberg Fund, write your check
to The Nature Conservancy of Hawaii and send it to:

The Nature Conservancy of Hawaii
Att: The F. Raymond Fosberg Fund
1116 Smith Street, #201

Honolulu, Hawaii 96817

ATOLL RESEARCH BULLETIN

NO. 390

LISTS OF TAXA NAMED FOR F. RAYMOND FOSBERG AND
BY HIM

BY

DAN H. NICOLSON

ISSUED BY
NATIONAL MUSEUM OF NATURAL HISTORY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C., U.S.A.
FEBRUARY 1994

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LISTS OF TAXA NAMED FOR F. RAYMOND FOSBERG AND
BY HIM

BY

DAN H. NICOLSON

SUMMARY

Fifty-one plant names, using epithets such as fosbergii or fosbergiana, honor Dr.
Fosberg. Almost 1000 taxa were named by him or with him as co-author, over half being
infraspecific taxa (subspecies, varieties, formae) in Gouldia and Hedyotis of the
Rubiaceae.

The basic tool used for these lists was Index Kewensis (mostly includes only seed
plants and only to species level) on a compact disk, read-only-memory (CD-ROM) in the
beta-test version. There are some difficulties with data misread by the optical character
reader, such as Piper "foshergii" and Diplostephium "fcsbergii". Thus, there may be some
overlooked species names masked by a misspelling. Also, Index Kewensis did not
incorporate trinomials (infraspecific names) or names of subdivisions of families and
genera until relatively recently. This may result in some overlooked names at these ranks.

More names were found in databases maintained at Missouri Botanical Garden
(TROPICOS is excellent for mosses). Marshall Crosby and Robert Magill, bryologists,
found two more moss names and four more vascular plant names, all older trinomials. A
trinomial and an unexpected binomial (Gouldia fosbergii, apparently overlooked by Index
Kewensis) was found by my colleague, Warren L. Wagner, in a database of Hawiian plant
names. The "Gray Cards" database of Harvard University (names of New World vascular
plants after 1890), now on-line, was checked.

I don’t know of tools comparable to /ndex Kewensis and these databases for other
biological groups. Dr. Paul C. Silva (University of California at Berkeley), who maintains
a card index to algae, was asked if he could locate phycological names. Dr. F. Christian
Thompson, member of the Zoological Commission, was asked if he could locate
zoological names. Dr. David Lellinger, pteridologist here at the Smithsonian, was asked
about finding fern names. All reported that they knew of no taxa named for Dr. Fosberg
but that the tools do not exist to check, let alone to be certain.

Author names are abbreviated in accordance with APN (R. K. Brummitt & C. E.
Powell’s 1992 Authors of plant names). Periodical titles are abbreviated in
accordance with B-P-H (G. H. M. Lawrence et al.’s 1968 Botanico-Periodicum-
Huntianum) and its Supplement (Bridson & Smith, 1991).

Unless some problem or inconsistency was noted, the original publication was not
checked. If errors of commission or omission are noted, the author would be grateul for
the information.

Botany Department, MRC-166, National Museum of Natural History, Smithsonian
Institution, Washington, D.C. 20560, U.S.A.

TAXA NAMED FOR F. RAYMOND FOSBERG

Shortly before Dr. Fosberg died, I reported to him that his moss, Poftia fosbergii (see
below), had been demoted to a variety, Pottia sterkeana var. fosbergii. He commented
that it was the first species named for him, based on material that he collected shortly
before his twentieth birthday in 1928 as a student of Philip Munz at Pomona College.

On a more personal note, some of Dr. Fosberg’s colleagues have told me that he was
pleased that Emilia fosbergii was named for him. Dr. Fosberg showed me how to tell the
three confused neotropical species apart, almost at arm’s length. This species, going under
a misapplied name, was new so I named it for him. In any case, wherever he went in the
tropics, he could point out this widespread weed and tell listeners that it was named for
him.

The Manual of Flowering Plants of Hawai'i says that Fosberg’s lovegrass (Eragrostis
fosbergii) is extinct. Throughout his career Dr. Fosberg supported conservation and
warned of the dangers of man’s destruction of habitats. Are others extinct?

Fifty-one names in his honor have been located, although some are transfers of the
same taxon, such as Bernardi’s transferring Cussonia fosbergiana to Schefflera. One
invalidly published name, a nomen nudum, has been omitted: Serianthes minhassae subsp.
fosbergii Kanis in Brunonia 2: 302. 1980 (= Albizia melanesica Fosberg).

There is a distinction between adjectival and substantive epithets. Adjectival epithets,
agreeing in gender, number and case with the generic name (fosbergianus, -a, -um), are
appropriate to honor someone who has not had a direct part in the taxon itself, such as
a professor, supporter, etc., and is literally translated from Latin as "the Fosbergian
Cussonia." Substantive epithets, nouns with independent gender, number and case, are in
genitive case (fosbergii) and are appropriate to honor one who has had a direct and
substantive part in the taxon, such as having collected it, recognized it as new, etc.,
literally "Fosberg’s Emilia." Recent workers may be unaware of the distinction and use
the substantive form because it is shorter. Nevertheless, of the species named in Fosberg’s
honor, he appears to have a substantive, direct part in 90% (fosbergii) and an indirect part
in 10% (fosbergianus, -a, -um), the mark of an active field worker.

Musci

Ectropothecium fosbergii E. B. Bartram in Occas. Pap. Bernice P. Bishop Mus. 15(27):
346 (1940).

Phascum fosbergii (E. B. Bartram) J. Guerra in J. Bryol. 16: 59 (1990) (Pottia
fosbergii).

Pottia fosbergii E. B. Bartram in Bryologist 33: 18 (1930).

Pottia sterkeana var. fosbergii (E. B. Bartram) R. H. Zander in Novon 3: 92 (1993)
(Pottia fosbergii).

Acanthaceae
Aphelandra fosbergii Leonard in Contrib. U. S. Natl. Herb. 31: 711 (1958).

Aquifoliaceae
Ilex fosbergiana S. Y. Hu in J. Jap. Bot. 46: 68 (1971).

Araliaceae
Cheirodendron trigynum var. fosbergii Sherff in Bot. Leafl. 5: 7 (1951).
Cussonia fosbergiana Bernardi in Ber. Schweiz. Bot. Ges. 76: 363 (1966).
Schefflera fosbergiana (Bernardi) Bernardi in Candollea 24: 117 (1969) (Cussonia
fosbergiana).

Asteraceae (Compositae)
Adenostemma fosbergii R. M. King & H. Rob. in Phytologia 29: 6 (1974).
Ageratina fosbergii R. M. King & H. Rob. in Phytologia 28: 495 (1974).
Emilia fosbergii Nicolson in Phytologia 32: 34 (1975).
Diplostephium fosbergii Cuatrec. in Bull. Torrey Bot. Club 80: 403 (1953).
Hesperomannia bushiana var. fosbergii Degener, Fl. Haw. Fam. 344 Hesperomannia
bushiana [unpaged] (1937).
Munnozia fosbergii H. Rob. in Phytologia 34: 382 (1976).
Perezia fosbergii Tovar in Publ. Mus. Hist. Nat. "Javier Prado", ser. B, Bot. 1(8):22
(1955).
Pluchea fosbergii Cooperr. & Galang in Amer. J. Bot. 52: 1025 (1965).
Senecio fosbergii Cuatrec. in Feddes Repert. Spec. Nov. Regni Veg. 55: 138 (1953).

Cactaceae
Opuntia fosbergii C. B. Wolf in Occas. Papers Rancho Santa Ana Bot. Gard. 2: 79
(1938).

Cyclanthaceae
Sphaeradenia fosbergii Harling in Acta Hort. Berg. 18: 353 (1958).

Cyperaceae
Fimbristylis fosbergiana T. Koyama in Bull. Arts & Sci. Div. Ryukyu Univ. 3 : 66
1959).

Ericaceae
Themistoclesia fosbergii A. C. Sm. in Amer. J. Bot. 40: 470 (1953).

Fabaceae (Leguminosae)
Schleinitzia fosbergii Nevling & Niezgoda in Adansonia, n. s. 18: 362 (1978).

Gesneriaceae
Cyrtandra fosbergii St. John & Storey in Occas. Pap. Bernice P. Bishop Mus. 20(16):
84 (1950).
Cyrtandra grayana var. fosbergii St. John in Phytologia 63: 479 (1987).

Lauraceae
Litsea fosbergii Kosterm. in Ceylon J. Sci., Biol. Sci. 9: 51 (1971).

Loganiaceae
Labordia kaalae var. fosbergii Sherff in Amer. J. Bot. 5: 588 (1938).

Marcgraviaceae
Gravia fosbergiana Ewan in Nat. Hist. Misc. 88: 2 (1951).

Melastomataceae
Miconia fosbergii Wurdack in Phytologia 11: 390 (1965).

Menispermaceae
Tinospora fosbergii Kundu in Ceylon J. Sci., Biol. Sci. 12: 49 (1976).

Monimiaceae
Siparuna fosbergii A. C. Sm. in Amer. J. Bot. 40: 469 (1953).

Myrsinaceae
Myrsine fosbergii Hosaka in Occas. Papers Bernice P. Bishop Mus. 16(2): 46 (1940).
Myrsine meziana var. fosbergii (Hosaka) Wilbur in Pac. Sci. 19: 522 (1965) (Myrsine
fosbergii ).
Rapanea fosbergii (Hosaka) Degener & I. Degener in Phytologia 22: 212 (1971) (Myrsine
fosbergii).

Onagraceae
Fuchsia fosbergii Munz in Aliso 7: 409 (1972).

Pandanaceae
Pandanus fosbergii St. John in Pacific Sci. 15: 331 (1961).

Piperaceae
Peperomia fosbergii Yunck. in Bernice P. Bishop Mus. Bull. 143: 46 (1937).
Piper fosbergii Trel. in Lloydia 3: 111 (1940).

Pittosporaceae
Pittosporum insigne var. fosbergii Sherff in Field Mus. Nat. Hist., Bot. Ser. 22: 413
(1941).

Poaceae (Gramineae)
Eragrostis fosbergii Whitney in Occas. Pap. Bernice P. Bishop Mus. 13(8): 75 (1937).

Polygalaceae
Monnina fosbergii Ferreyra in Phytologia 69: 356 (1990).

Portulacaceae
Portulaca fosbergii Poelln. in Occas. Pap. Bernice P. Bishop Mus. 12(9): 3 (1936).

Rubiaceae
Arachnothrix fosbergii Steyerm. in Mem. New York Bot. Gard. 17: 255 (1967).
Galium fosbergii Dempster in Madrofio 35: 3 (1988).
Gardenia fosbergii Tirveng. in Nordic J. Bot. 3: 460 (1983).
Gouldia fosbergii Degener & I. Degener in Phytologia 15: 49 (1967).
Hedyotis fosbergii W. L. Wagner & D. R. Herbst in Bishop Mus. Occas. Pap. 29: 111
(1989).
Psychotria fosbergii Steyerm. in Mem. New York Bot. Gard. 23: 630 (1972).
Psychotria fosbergii Sohmer in Bot. Notiser 129: 381 (1977) (= Psychotria sohmeri Kiehn
1986).

Solanaceae
Solanum fosbergianum D’Arcy in Ann. Missouri Bot. Gard. 60: 700 (1974).

Tiliaceae
Trichospermum fosbergii Kosterm. in Trans. Bot. Soc. Edinburgh 41: 416 (1972).

TAXA NAMED BY F. RAYMOND FOSBERG

The basic input was again taken from /ndex Kewensis on CD-ROM, this time by
Dr. Joseph Kirkbride (US Department of Agriculture). Ellen Farr (Smithsonian Botany
Department) brought Dr. Kirkbride’s data through a database (DBase IV) into text. I
am very grateful to them for their time and trouble. I then removed all entries with
Fosberg as an author in parentheses (transfer of a Fosberg taxon), such as Chamaesyce
stoddartii (Fosberg) J. Sojak, unless it was made by Fosberg, such as Casuarina
litorea var. souderi (Fosberg) Fosberg & Sachet. I also removed "Fosberg ex"
citations, such as Hedyotis angulata Fosberg ex Shinners since Dr. Fosberg was not
actually an author. At that point there were 466 entries.

A card file of taxa named by Dr. Fosberg was the basis for additions. Finally, his
recent papers were scanned for new taxa. When this was completed there were 993
entries.

The bulk of the new entries concern infraspecific names. No effort was made to
include autonyms (a trinomial combination in which the final epithet is the same as
the species epithet). Infraspecific uses of "eu-" epithets were also omitted. An example
is Gouldia terminalis var. ovata f. euovata (a classification, not a combination) which
becomes Gouldia terminalis f. "euovata" (a combination, not a classification). Names
with such epithets are invalidly published under Art. 24.3. A similar problem involves
Gouldia terminalis var. ovata f. russii (Brittonia 8: 176. 1956). This, reduced from a
classification to the combination G. terminalis f. russii, is an isonym of G. terminalis

6

f. russii Fosberg (Bernice P. Bishop Mus. Bull. 147: 49. 1937), not a new combination
based on it, there classified as a form of G. terminalis var. kaala. In short, moving a
forma from one variety to another in the same species, does not create a new name.

Fungi
Diaporthopsis metrosideri Roane & Fosberg in Mycologia 75: 165 (1983).
[Pyrenomycetes]
Pteridophyta
Gleicheniaceae

Gleichenia linearis var. latiloba (Holttum) Fosberg in Smithsonian Contr. Bot. 45: 4
(1980).

Gleichenia linearis var. tomentosa (Luerss.) Fosberg in Occas. Pap. Bernice P. Bishop
Mus. 24(2): 10 (1969).

Gleichenia weatherbyi Fosberg in Amer. Fern J. 40: 140 (1950).

Hymenophyllaceae
Hymenophyllum alternatum Fosberg in Amer. Fern J. 40: 135 (1950).
Hymenophyllum digitaria (Sw.) Fosberg in Smithsonian Contr. Bot. 45: 1 (1980).
Trichomanes javanicum var. boryanum (Kunze) Fosberg in Amer. Fern J. 40: 136
(1950).

Lycopodiaceae
Lycopodium phlegmaria var. mirabile (Willd.) Fosberg in Lloydia 3: 110. (1940).

Ophioglossaceae
Ophioglossum pendulum var. falcatum (Presl) Fosberg in Occas. Pap. Bernice P.
Bishop Mus. 23(2): 29 (1962).

Polypodiaceae

Dryopteris goggilodus (Schkuhr) Fosberg in Occas. Pap. Bernice P. Bishop Mus.
16(15): 337 (1942).

Polypodium katuii (Brownlie) Fosberg in Micronesica 23: 1 (1990).

Sphenomeris chinensis var. hawaiiensis Fosberg & Sachet in Micronesica 18(1): 131
(1983 "1982").

Sphenomeris chinensis var. tenuisecta Fosberg & Sachet in Micronesica 18(1): 132
(1983 "1982").

Thelypteris carolinensis (Hosok.) Fosberg in Smithsonian Contr. Bot. 45: 4 (1980).

Thelypteris cyatheoides (Kaulf.) Fosberg in Occas. Pap. Bernice P. Bishop Mus. 23(2):
30 (1962).

Thelypteris fadenii Fosberg & Sachet in Smithsonian Contr. Bot. 8: 9 (1972).

Thelypteris guamensis (Holttum) Fosberg & Sachet in Amer. Fern. J. 71: 82 (1981).

Thelypteris opulenta (Kaulf.) Fosberg in Smithsonian Contr. Bot. 8: 3 (1972).

Thelypteris opulenta var. hirsuta (Ching) Fosberg in Smithsonian Contr. Bot. 8: 6
(1972).

Thelypteris parasitica (L.) Fosberg in Occas. Pap. Bernice P. Bishop Mus. 23(2): 30
(1962).

Thelypteris peliliuensis Fosberg in Smithsonian Contr. Bot. 45: 4 (1980).

Thelypteris rupi-insularis Fosberg in Smithsonian Contr. Bot. 45: 5 (1980).

Thelypteris sandwicensis (Hook. & Arn.) Fosberg in Occas. Pap. Bernice P. Bishop Mus.
23(2): 30 (1962).

Thelypteris stenogrammoides (Baker) Fosberg in Occas. Pap. Bernice P. Bishop Mus.
23(2): 30 (1962).

Thelypteris wagneri Fosberg & Sachet in Smithsonian Contr. Bot. 8: 6 (1972).

Schizaeaceae
Lygodium circinnatum var. semihastatum (Cav.) Fosberg in Amer. Fern J. 40: 142
(1950).
Schizaea dichotoma var. sellingii Fosberg in Amer. Fern J. 40: 144 (1950).

Spermatophyta
Acanthaceae
Hygrophila triflora (Roxb.) Fosberg & Sachet in Baileya 21: 147 (1981).
Pseuderanthemum bibracteatum Fosberg in Lloydia 3: 119 (1940).
Pseuderanthemum bibracteatum f. ochraceum Fosberg in Lloydia 3: 121 (1940).
Pseuderanthemum carruthersii var. atropurpureum (Bull) Fosberg in Phytologia 5: 290
(1955).
Pseuderanthemum carruthersii var. reticulatum (Bull) Fosberg in Smithsonian Contr.
Bot. 45: 26 (1980).
Pseuderanthemum palauense Fosberg & Sachet in Smithsonian Contr. Bot. 45: 27
(1980).
Aceraceae
Acer nigrum var. floridanum (Chapm.) Fosberg in Castanea 19: 27 (1954).
Acer nigrum var. glaucum (Schmidt) Fosberg in Castanea 19: 27 (1954).
Acer nigrum var. grandidentatum (Torr. & A. Gray) Fosberg in Castanea 19: 27
(1954).
Acer nigrum var. leucoderme (Small) Fosberg in Castanea 19: 27 (1954).
Acer nigrum var. pseudoplatanoides (Pax) Fosberg in Amer. Midl. Naturalist 26: 695
(1941).
Acer nigrum var. schneckii (Rehder) Fosberg in Castanea 19: 27 (1954).
Acer nigrum var. sinuosum (Sarg.) Fosberg in Castanea 19: 27 (1954).

Aizoaceae
Sesuvium portulacastrum var. griseum Fosberg in Occas. Pap. Bernice P. Bishop Mus.
21(14): 47 (1952).

Amaranthaceae
Alternanthera paronychioides var. bettzickiana (Regel) Fosberg in Willdenowia 20:
259 (1991).
Lagrezia oligomeroides (C. H. Wright) Fosberg in Kew Bull. 29: 262 (1974).

Amaryllidaceae
Crinum asiaticum var. pedunculatum (R. Br.) Fosberg & Sachet in Micronesica 20:
131 (1988 "1987").

Apocynaceae

Alstonia marquisensis M. L. Grant ex Fosberg & Sachet in Micronesica 8: 46 (1972).

Alyxia stellata var. deckeri Fosberg & Sachet in Micronesica 10: 254 (1974).

Alyxia stellata var. fatuhivensis Fosberg & Sachet in Micronesica 10: 254 (1974).

Alyxia stellata var. marquesensis (F. Br.) Fosberg & Sachet in Micronesica 10: 253
(1974).

Melodinus insularis (Markgr.) Fosberg in Micronesica 23: 150 (1990).

Neisosperma acuminata (Valeton) Fosberg & Sachet in Adansonia, n. s. 17: 28 (1977)
without basionym page.

Neisosperma apoensis (Elmer) Fosberg & Sachet in Adansonia, n. s. 17: 28 (1977).

Neisosperma brownii Fosberg & Sachet in Micronesica 8: 49 (1972).

Neisosperma citrodora (Lauterb. & K. Schum.) Fosberg & Sachet in Adansonia, n. s.
17: 29 (1977).

Neisosperma ficifolia (S. Moore) Fosberg & Sachet in Adansonia, n. s. 17: 29 (1977).

Neisosperma glomerata (Blume) Fosberg & Sachet in Adansonia, n. s. 17: 29 (1977).

Neisosperma iwasakiana (Koidz.) Fosberg & Sachet in Adansonia, n. s. 17: 29 (1977).

Neisosperma kilneri (F. Muell.) Fosberg & Sachet in Adansonia, n. s. 17: 29 (1977).

Neisosperma nakaiana (Koidz.) Fosberg & Sachet in Adansonia, n. s. 17: 30 (1977).

Neisosperma oppositifolia (Lam.) Fosberg & Sachet in Micronesica 8: 48 (1972).

Neisosperma poweri (F. M. Bailey) Fosberg & Sachet in Adansonia, n. s. 17: 31
(1977):

Neisosperma rudis (Markgr.) Fosberg & Sachet in Adansonia, n. s. 17: 31 (1977).

Neisosperma sciadophylla (Markgr.) Fosberg & Sachet in Adansonia, n. s. 17: 31
i977).

Ochrosia coccinea var. peekelii (Markgr.) Fosberg & Sachet in Adansonia, n. s. 17: 25
(1977):

Ochrosia fatuhivensis Fosberg & Sachet in Micronesica 8: 48 (1972).

Ochrosia mariannensis var. crassicarpa Fosberg & Falanruw in Micronesica 11: 80
(1975).

Ochrosia minima (Markgr.) Fosberg & Boiteau in Adansonia, n. s. 17: 27 (1977).

Ochrosia nukuhivensis Fosberg & Sachet in Micronesica 8: 48 (1972).

Ochrosia solomonensis (Merr. & Perry) Fosberg & Boiteau in Adansonia, n. s. 17: 28
(1977).

Rauvolfia sachetiae Fosberg in Smithsonian Contr. Bot. 47: 21 (1981).

Araceae
Rhaphidophora carolinensis (Volkens) Fosberg in Smithsonian Contr. Bot. 45: 5
(1980).

Araliaceae

Osmoxylon mariannense (Kaneh.) Fosberg & Sachet in Smithsonian Contr. Bot. 45: 16
(1980).

Osmoxylon oliveri Fosberg & Sachet in Smithsonian Contr. Bot. 45: 16 (1980).

Osmoxylon pachyphyllum (Kaneh.) Fosberg & Sachet in Smithsonian Contr. Bot. 45:
16 (1980).

Osmoxylon truncatum (Kaneh.) Fosberg & Sachet in Smithsonian Contr. Bot. 45: 16
(1980).

Polyscias scutellaria (Burm. f.) Fosberg in Occas. Pap. Univ. Hawaii 46: 9 (1948).

Polyscias tricochleata (Miq.) Fosberg in Phytologia 5: 290 (1955).

Arecaceae (Palmae)

Clinostigma carolinense (Becc.) H. E. Moore & Fosberg in Gentes Herb. 8: 462
(1956).

Clinostigma ponapense (Becc.) H. E. Moore & Fosberg in Gentes Herb. 8: 463
(1956).

Clinostigma savoryanum (Rehder & E. H. Wilson) H. E. Moore & Fosberg in Gentes
Herb. 8: 465 (1956).

Clinostigma smithii (Burret) H. E. Moore & Fosberg in Gentes Herb. 8: 462 (1956)

Gulubia palauensis (Becc.) H. E. Moore & Fosberg in Gentes Herb. 8: 455 (1956).

Ptychosperma hosinoi (Kaneh.) H. E. Moore & Fosberg in Gentes Herb. 8: 468
(1956).

Ptychosperma ledermannianum (Becc.) H. E. Moore & Fosberg in Gentes Herb. 8:
469 (1956).

Ptychosperma palauense (Kaneh.) H. E. Moore & Fosberg in Gentes Herb. 8: 470
(1956).

Asclepiadaceae
Dischidia melanesica Fosberg in Lloydia 3: 116 (1940).
Hoya dodecatheiflora Fosberg in Lloydia 3: 118 (1940).

Asteraceae (Compositae)
Dubautia sherffiana Fosberg in Bull. Torrey Bot. Club 70: 395 (1943).
Emilia alstonii Fosberg in Ceylon J. Sci., Biol. Sci. 10: 66 (1972).
Emilia baldwinii Fosberg in Ceylon J. Sci., Biol. Sci. 10: 68 (1972).
Emilia exserta Fosberg in Ceylon J. Sci., Biol. Sci. 10: 65 (1972).
Emilia speeseae Fosberg in Ceylon J. Sci., Biol. Sci. 10: 67 (1972).
Wedelia biflora var. canescens (Gaudich.) Fosberg in Phytologia 5: 291 (1955).
Wollastonia biflora var. canescens (Gaudich.) Fosberg in Smithsonian Contr. Bot. 45:
33 (1980).
Wollastonia lifuana (Hochr.) Fosberg in Allertonia 7: 80 (1993).

Bignoniaceae
Chilopsis linearis var. arcuata Fosberg in Madrofio 3: 366 (1936).
Chilopsis linearis var. glutinosa (Engelm.) Fosberg in Madrofio 3: 365 (1936).

10

Boraginaceae
Heliotropium procumbens var. depressum (Cham.) Fosberg & Sachet in Smithsonian
Contr. Bot. 45: 24 (1980).

Brassicaceae (Cruciferae)
Lepidium bidentatum var. o-waihense (Cham. & Schtdl.) Fosberg in Occas. Pap.
Bernice P. Bishop Mus. 24(2): 17 (1969).
Lepidium bidentatum var. remyi (Drake) Fosberg in Occas. Pap. Bernice P. Bishop
Mus. 24(2): 18 (1969).

Cactaceae

Coryphantha bella (Britton & Rose) Fosberg in Bull. S. Calif. Acad. Sci. 30: 58
(1931).

Coryphantha chaffeyi (Britton & Rose) Fosberg in Bull. S. Calif. Acad. Sci. 30: 58
(1931).

Coryphantha chihuahensis (Britton & Rose) Fosberg in Bull. S. Calif. Acad. Sci. 30:
58 (1931).

Coryphantha dasycantha (Engelm.) Fosberg in Bull. S. Calif. Acad. Sci. 30: (1931).

Coryphantha lloydii (Britton & Rose) Fosberg in Bull. S. Calif. Acad. Sci. 30: 58
«i931:

Coryphantha piercei Fosberg in Bull. S. Calif. Acad. Sci. 30: 58 (1931).

Coryphantha sneedii (Britton & Rose) Fosberg in Bull. S. Calif. Acad. Sci. 30: 58
(1931).

Coryphantha tuberulosa (Engelm.) Fosberg in Bull. S. Calif. Acad. Sci. 30: 58 (1931).

Echinocactus johnstonianus (Britton & Rose) Fosberg in Amer. Midl. Naturalist 27:
257 (1942).

Echinocereus englemannii var. munzii (Parish) Fosberg in Bull. S. Calif. Acad. Sci.
322123) (1933).

Neomammillaria subg. Dolichothele (Schum.) Fosberg in Bull. S. Calif. Acad. Sci. 30:
57 (1931).

Neomammillaria subg. Galactochylus (Schum.) Fosberg in Bull. S. Calif. Acad. Sci.
30: 58 (1931), based on generic type.

Neomammillaria subg. Hydrochylus (Schum.) Fosberg in Bull. S. Calif. Acad. Sci. 30:
56 (1931).

Neomammillaria subg. Phyllospermum (Britton & Rose) Fosberg in Bull. S. Calif.
Acad. Sci. 30: 57 (1931).

Neomammillaria subg. Solisia (Britton & Rose) Fosberg in Bull. S. Calif. Acad. Sci.
30: 57 (1931).

Neomammillaria echinops (Scheidw.) Fosberg in Bull. S. Calif. Acad. Sci. 30: 56
(19311):

Neomammillaria longimamma (DC.) Fosberg in Bull. S. Calif. Acad. Sci. 30: 58
(1931).

Neomammillaria pectinata (B. Stein) Fosberg in Bull. S. Calif. Acad. Sci. 30: 58
(1931).

Neomammillaria sphaerica (Dietr.) Fosberg in Bull. S. Calif. Acad. Sci. 30: 58 (1931).

11

Neomammillaria tetrancistra (Engelm.) Fosberg in Bull. S. Calif. Acad. Sci. 30: 57
(1931).

Neomammillaria uberiformis (Zucc.) Fosberg in Bull. 8. Calif. Acad. Sci. 30: 58 (1931).

Neomammillaria zeyeriana (F. Haage) Fosberg in Bull. S. Calif. Acad. Sci. 30: 56 (1931).

Opuntia bigelovii var. hoffmannii Fosberg in Bull. S. Calif. Acad. Sci. 32: 121 (1933).

Opuntia engelmannii var. megacarpa (Griffiths) Fosberg in Bull. S. Calif. Acad. Sci. 33:
100 (1934).

Opuntia engelmannii var. wootonii (Griffiths) Fosberg in Bull. S. Calif. Acad. Sci. 30:
59 (1931).

Opuntia phaeacantha var. covillei (Britton & Rose) Fosberg in Bull. S. Calif. Acad. Sci.
33: 102 (1934).

Opuntia phaeacantha var. mojavensis (Engelm. & Bigel.) Fosberg in Bull. S. Calif. Acad.
Sci. 33: 103 (1934).

Opuntia phaeacantha var. piercei Fosberg in Bull. S. Calif. Acad. Sci. 33: 102 (1934).

Campanulaceae
Campanula divaricata f. alba Fosberg in Castanea 20: 60 (1955).
Lobelia gaudichaudii var. koolauensis Hosaka & Fosberg in Occas. Pap. Bernice P.
Bishop Mus. 14(1): 4. (1938).
Rollandia lanceolata var. kipapensis Hosaka ex Fosberg in Occas. Pap. Bernice P.
Bishop Mus. 12(15): 10 (1936).

Caprifoliaceae
Sambucus racemosa var. arborescens (Torr. & A. Gray) Fosberg in Amer. Midl.
Naturalist 37: 765 (1942).

Caryophyllaceae
Drymaria gentryi Fosberg in Proc. Biol. Soc. Wash. 62: 147 (1949).
Drymaria stipitata Fosberg in Lloydia 4: 281 (1941).

Casuarinaceae
Casuarina equisetifolia var. souderi Fosberg in Micronesica 2: 143 (1966).
Casuarina litorea var. souderi (Fosberg) Fosberg & Sachet in Smithsonian Contr. Bot.
24: 6 (1975).

Celastraceae
Loeseneriella macrantha var. palauica (Loes.) Fosberg in Smithsonian Contr. Bot. 45:
12 (1980).
Maytenus palauica (Loes.) Fosberg in Phytologia 5: 290 (1955).
Maytenus thompsonii (Merr.) Fosberg in Phytologia 5: 290 (1955).

Chenopodiaceae
Atriplex argentea var. caput-medusae (Eastw.) Fosberg in Amer. Midl. Naturalist 26:
693 (1941).

12

Atriplex phyllostegia var. draconis (Jones) Fosberg in Amer. Midl. Naturalist 26: 694

ie ee var. greenei (Nelson) Fosberg in Amer. Midl. Naturalist 27: 253

Are fremontii var. atrovirens (Rydb.) Fosberg in Amer. Mid]. Naturalist 26: 691

a oahuense var. discosperma Fosberg in Occas. Pap. Bernice P. Bishop Mus.
23(2): 35 (1962).

Clusiaceae (Guttiferae)
Calophyllum inophyllum var. takamaka Fosberg in Kew Bull. 29: 255 (1974).
Calophyllum inophyllum var. wakamatsui (Kaneh.) Fosberg & Sachet in Smithsonian
Contr. Bot. 45: 12 (1980).
Garcinia ponapensis var. trukensis (Kaneh.) Fosberg in Smithsonian Contr. Bot. 45:
13 (1980).
Garcinia rumiya var. calcicola Fosberg in Smithsonian Contr. Bot. 45: 14 (1980).
Garcinia laddii Fosberg in Pac. Sci. 31: 297 (1977). [fossil]

Combretaceae

Terminalia glabrata var. brownii Fosberg & Sachet in Smithsonian Contr. Bot. 47: 15
(1981).

Terminalia glabrata var. haroldii (Exell) Fosberg & Sachet in Smithsonian Contr. Bot.
47: 16 (1981).

Terminalia glabrata var. intonsa Fosberg in Micronesica 23: 2 (1990).

Terminalia glabrata var. koariki (Exell) Fosberg & Sachet in Smithsonian Contr. Bot.
47: 16 (1981).

Terminalia glabrata var. rarotongensis Fosberg & Sachet in Smithsonian Contr. Bot.
47: 16 (1981).

Terminalia rostrata Fosberg & Falanruw in Phytologia 28: 469 (1974).

Connaraceae
Connarus semidecandrus var. gaudichaudii (DC.) Fosberg in Micronesica 11: 82
(1975).

Convolvulaceae
Ipomoea indica var. acuminata (Vahl) Fosberg in Bot. Not. 129: 38 (1976).
Ipomoea indica var. hosakae Fosberg in Micronesica 2: 151 (1966).
Ipomoea setifera var. fimbriosepala (Choisy) Fosberg in Smithsonian Contr. Bot. 36:
24 (1977).
Ipomoea tiliacea var. merremioides Fosberg in Smithsonian Contr. Bot. 21: 14 (1975).
Ipomoea tiliacea var. smithii Fosberg in Smithsonian Contr. Bot. 21: 15 (1975).

Cornaceae
Cornus sericea f. baileyi (Coult. & Evans) Fosberg in Bull. Torrey Bot. Club 69: 588
(1942).

18.

Cornus sericea f. californica (Meyen) Fosberg in Bull. Torrey Bot. Club 69: 589
(1942).

Cornus sericea f. interior (Rydb.) Fosberg in Bull. Torrey Bot. Club 69: 588 (1942).

Cornus sericea f. occidentalis (Torr. & A. Gray) Fosberg in Bull. Torrey Bot. Club
69: 589 (1942).

Cornus sericea subsp. occidentalis (Torr. & A. Gray) Fosberg in Bull. Torrey Bot.
Club 69: 589 (1942).

Cornus sericea subsp. stolonifera (Michx.) Fosberg in Bull. Torrey Bot. Club 69: 587
(1942).

Cornus sericea subsp. stolonifera (Michx.) Fosberg in Bull. Torrey Bot. Club 69: 587
(1942).

Cucurbitaceae

Citrullus lanatus var. caffrorum (Alef.) Fosberg in Smithsonian Contr. Bot. 45: 15
(1980).

Trichosanthes hosokawae Fosberg in Occas. Pap. Bernice P. Bishop Mus. 22(1): 67
(1958).

Zehneria grayana (Cogn.) Fosberg & Sachet in Smithsonian Contr. Bot. 47: 12 (1981).

Zehneria grayana var. vitiensis (A. Gray) Fosberg & Sachet in Smithsonian Contr.
Bot. 47: 13 (1981).

Zehneria guamensis (Merr.) Fosberg in Smithsonian Contr. Bot. 45: 15 (1980).

Zehneria samoensis (A. Gray) Fosberg & Sachet in Smithsonian Contr. Bot. 47: 12
(1981).

Cunoniaceae
Cunonia parviflora var. glabrata (F. Br.) Fosberg in Micronesica 8: 45 (1972).
Cunonia parviflora var. marquesana (F. Br.) Fosberg in Micronesica 8: 44 (1972).
Weinmannia parviflora var. glabrata (F. Br.) Fosberg in Micronesica 8: 45 (1972).
Weinmannia parviflora var. marquesana (F. Br.) Fosberg in Micronesica 8: 44 (1972).
Weinmannia parviflora var. myrsinites Fosberg & Sachet in Micronesica 8: 45 (1972).

Cyperaceae
Bulbostylis basalis Fosberg in Kew Bull. 31: 829 (1977).
Cyperus bigibbosa Fosberg in Kew Bull. 31: 832 (1977).
Cyperus niveus var. leucocephalus (Kunth) Fosberg in Kew Bull. 31: 835 (1977).
Cyperus odoratus var. attenuatus Fosberg & Sachet in Micronesica 20: 167 (1988
"1987").
Cyperus odoratus var. curtispiculus Fosberg & Sachet in Micronesica 20: 167 (1988
a LOST):
Cyperus odoratus var. novae-hannoverae (Boeck.) Fosberg & Sachet in Micronesica
20: 168 (1988 "1987").
Fimbristylis juncea var. marquesana (Steud.) Fosberg in J. Jap. Bot. 63: 83 (1988).
Fimbristylis juncea var. nukahivensis (Steud.) Fosberg in J. Jap. Bot. 63: 82 (1988).
Fimbristylis juncea var. separanda (Jardin) Fosberg in J. Jap. Bot. 63: 81 (1988).
Fimbristylis juncea var. tertia (Jardin) Fosberg in J. Jap. Bot. 63: 81 (1988).

14

Fimbristylis sachetiana Fosberg in J. Jap. Bot. 63: 83 (1988).

Dioscoreaceae
Dioscorea esculenta var. tiliifolia (Kunth) Fosberg & Sachet in Micronesica 20: 135
(1988 "1987").

Ebenaceae

Diospyros sect. Cupulifera Fosberg in Bull. Torrey Bot. Club 65: 613 (1938).

Diospyros sect. Ferreola (Roxb.) Fosberg in Bull. Torrey Bot. Club 65: 609 (1938).

Diospyros christophersenii Fosberg in Bull. Torrey Bot. Club 65: 613 (1939).

Diospyros ellipticifolia var. elliptica (G. Forst.) Fosberg in Bull. Torrey Bot. Club 65:
611 (1939).

Diospyros ellipticifolia var. iridea Fosberg in Bull. Torrey Bot. Club 65: 612 (1939).

Diospyros ferrea f. degeneri Fosberg in Occas. Pap. Bernice P. Bishop Mus. 15(10):
130 (1939).

Diospyros ferrea f. kauaiensis Fosberg in Occas. Pap. Bernice P. Bishop Mus. 15(10):
131 (1939).

Diospyros ferrea f. lanaiensis Fosberg in Occas. Pap. Bernice P. Bishop Mus. 15(10):
130)(1939);

Diospyros ferrea f. obtusa Fosberg in Occas. Pap. Bernice P. Bishop Mus. 15(10): 127
(1939).

Diospyros ferrea f. ovata Fosberg in Occas. Pap. Bernice P. Bishop Mus. 15(10): 127
(1939).

Diospyros ferrea f. pubescens (Skottsb.) Fosberg in Occas. Pap. Bernice P. Bishop
Mus. 15(10): 129 (1939).

Diospyros ferrea f. sandwicensis (A. DC.) Fosberg in Occas. Pap. Bernice P. Bishop
Mus. 15(10): 125 (1939).

Diospyros ferrea f. subcoriacea Fosberg in Occas. Pap. Bernice P. Bishop Mus.
15(10): 127 (1939).

Diospyros ferrea f. waiauensis Fosberg in Occas. Pap. Bernice P. Bishop Mus. 15(10):
128 (1939).

Diospyros ferrea f. wiebkei Fosberg in Occas. Pap. Bernice P. Bishop Mus. 15(10):
131 (1939).

Diospyros ferrea subsp. sandwicensis (A. DC.) Fosberg in Occas. Pap. Bernice P.
Bishop Mus. 15(10): 122 (1939).

Diospyros ferrea var. compacta (R. Br.) Fosberg in Brittonia 40: 61 (1988).

Diospyros ferrea var. degeneri Fosberg in Occas. Pap. Bernice P. Bishop Mus. 15(10):
129 (1939).

Diospyros ferrea var. gillespiei Fosberg in Bull. Torrey Bot. Club 65: 610 (1938).

Diospyros ferrea var. kauaiensis Fosberg in Occas. Pap. Bernice P. Bishop Mus.
15(10): 130 (1939).

Diospyros ferrea var. nandarivatensis (Gillespie) Fosberg in Bull. Torrey Bot. Club
65: 610 (1938).

15)

Diospyros ferrea var. palauensis (Kaneh.) Fosberg in Bull. Torrey Bot. Club 67: 417
(1940).

Diospyros ferrea var. pubescens (Skottsb.) Fosberg in Occas. Pap. Bernice P. Bishop Mus.
15(10): 128 (1939).

Diospyros ferrea var. savaiiensis (Christ.) Fosberg in Bull. Torrey Bot. Club 65: 611
(1938).

Diospyros ferrea var. sclerophylla Fosberg in Occas. Pap. Bernice P. Bishop Mus. 15(10):
129: (1939).

Diospyros ferrea var. subimpressa Fosberg in Bull. Torrey Bot. Club 65: 611 (1938).

Diospyros ferrea var. toppingii Fosberg in Occas. Pap. Bernice P. Bishop Mus. 15(10):
129 (1939).

Diospyros globosa (A. C. Smith) Fosberg in Bull. Torrey Bot. Club 65: 612 (1939).

Diospyros hillebrandii (Seem.) Fosberg in Occas. Pap. Bernice P. Bishop Mus. 12(15):
9 (1936).

Diospyros rufa (Labill.) Fosberg in Bull. Torrey Bot. Club 65: 613 (1939).

Diospyros samoensis var. longisepala (Gillespie) Fosberg in Bull. Torrey Bot. Club 67:
418 (1940).

Diospyros sandwicensis (A. DC.) Fosberg in Occas. Pap. Bernice P. Bishop Mus. 12(15):
8 (1936).

Epacridaceae
Styphelia tameiameiae var. hexamera Fosberg & Hosaka in Occas. Pap. Bernice P.
Bishop Mus. 14(1): 4 (1938).

Ericaceae
Vaccinium calycinum f. grandifolium (Wawra) Fosberg in Occas. Pap. Bernice P.
Bishop Mus. 16(15): 342 (1942).
Vaccinium calycinum f. hamatidens (H. Lév.) Fosberg in Occas. Pap. Bernice P.
Bishop Mus. 16(15): 343 (1942).
Vaccinium calycinum var. grandifolium (Wawra) Fosberg in Occas. Pap. Bernice P.
Bishop Mus. 16(15): 343 (1942).

Euphorbiaceae

Acalypha amentacea f. circinata (Mill. Arg.) Fosberg in Smithsonian Contr. Bot. 45:
11 (1980).

Acalypha amentacea subsp. wilkesiana (Mill. Arg.) Fosberg in Smithsonian Contr.
Bot. 45: 10 (1980).

Acalypha amentacea var. grandis (Benth.) Fosberg in Smithsonian Contr. Bot. 45: 8
(1980).

Acalypha amentacea var. heterotricha Fosberg in Smithsonian Contr. Bot. 45: 9
(1980).

Acalypha amentacea var. palauensis Fosberg in Smithsonian Contr. Bot. 45: 9 (1980).

Acalypha amentacea var. trukensis (Pax & Hoffm.) Fosberg in Smithsonian Contr.
Bot. 45: 10 (1980).

16

Acalypha amentacea var. velutina (Mill. Arg.) Fosberg in Smithsonian Contr. Bot. 45:
10 (1980).

Acalypha cardiophylla var. ponapensis (Kaneh. & Hatus.) Fosberg in Smithsonian Contr.
Bot. 45: 11 (1980).

Acalypha crockeri Fosberg in Lloydia 3: 114 (1940).

Acalypha swallowensis Fosberg in Lloydia 3: 114 (1940).

Antidesma platyphylla var. hamakuaense Fosberg in Occas. Pap. Bernice P. Bishop Mus.
12(15): 8 (1936).

Claoxylon ooumuense Fosberg & Sachet in Smithsonian Contr. Bot. 47: 6 (1981).

Euphorbia mertonii Fosberg in Kew Bull. 33: 181 (1978).

Euphorbia prostrata var. candirhiza Fosberg in Micronesica 25: 189 (1992).

Euphorbia stoddartii Fosberg in Kew Bull. 33: 182 (1978).

Glochidion cleistanthoides Fosberg in Willdenowia 20: 263 (1991).

Glochidion excorticans Fosberg in Willdenowia 20: 260 (1991).

Glochidion excorticans var. calvum Fosberg in Willdenowia 20: 261 (1991).

Glochidion hosokawae Fosberg in Willdenowia 20: 261 (1991).

Glochidion websteri Fosberg in Willdenowia 20: 262 (1991).

Jatropha integerrima var. hastata (Jacq.) Fosberg in Rhodora 78: 102 (1976).

Margaritaria anomala (Baillon) Fosberg in Kew Bull. 33: 185 (1978).

Margaritaria anomala var. cheloniphorbe (Hutch.) Fosberg in Kew Bull. 33: 185 (1978).

Melanolepis multiglandulosa var. glabrata (Mill. Arg.) Fosberg in Phytologia 5: 289
(1955).

Phyllanthus maderaspatensis var. frazieri Fosberg in Kew Bull. 33: 188 (1978).

Phyllanthus mckenziei Fosberg in Kew Bull. 33: 189 (1978).

Phyllanthus sandwicensis f. rufidus Fosberg in Occas. Pap. Bernice P. Bishop Mus.
12(15): 6 (1936).

Fabaceae (Leguminosae)

Albizia sect. Pachyspermae (Benth.) Fosberg in Reinwardtia 7: 74 (1965).

Albizia eymae Fosberg in Reinwardtia 7: 87 (1965).

Albizia falcataria (L.) Fosberg in Reinwardtia 7: 88 (1965).

Albizia melanesica Fosberg in Reinwardtia 7: 85 (1965).

Albizia minhassae var. ledermannii (Harms) Fosberg in Reinwartdia 7: 85 (1965).

Albizia minhassae var. proliferata Fosberg in Reinwartdia 7: 83 (1965).

Albizia minhassae var. umbellata Fosberg in Reinwartdia 7: 84 (1965).

Albizia montana var. kostermansii Fosberg in Reinwartdia 7: 79 (1965).

Canavalia galatea var. hawaiiensis (Degener, I. Degener & Sauer) Fosberg in Occas.
Pap. Bernice P. Bishop Mus. 23(8): 132 (1966).

Canavalia galatea var. kauaiensis (Sauer) Fosberg in Occas. Pap. Bernice P. Bishop
Mus. 23(8): 132 (1966).

Canavalia galatea var. molokaiensis (Degener, I. Degener & Sauer) Fosberg in Occas.
Pap. Bernice P. Bishop Mus. 23(8): 132 (1966).

Canavalia megalantha var. falanruwiae Fosberg in Micronesica 11: 78 (1975).

Desmodium heterocarpon f. substrigosum Fosberg in Micronesica 2: 146 (1966).

Desmodium intortum var. pilosiusculum (DC.) Fosberg in Micronesica 4: 257 (1968).

Ue,

Inocarpus fagifer (Parkinson) Fosberg in J. Wash. Acad. Sci. 31: 95 (1941).

Leucaena insularum var. guamensis Fosberg & B. C. Stone in Micronesica 2: 67 (1965).

Prosopis pallida f. armata Fosberg in Occas. Pap. Bernice P. Bishop Mus. 23(8): 134
(1966).

Serianthes dilmyi Fosberg in Taxon 8: 65 (1959).

Serianthes ebudarum Fosberg in Reinwardtia 5: 307 (1960).

Serianthes kanehirae Fosberg in Reinwardtia 5: 302 (1960).

Serianthes kanehirae var. hooglandii Fosberg in Reinwardtia 5: 303 (1960).

Serianthes kanehirae var. yapensis Fosberg in Reinwardtia 5: 303 (1960).

Serianthes melanesica Fosberg in Reinwardtia 5: 312 (1960).

Serianthes melanesica var. lifouensis Fosberg in Reinwardtia 5: 314 (1960).

Serianthes melanesica var. macdanielsii Fosberg in Reinwardtia 5: 313 (1960).

Serianthes melanesica var. meeboldii Fosberg in Reinwardtia 5: 314 (1960).

Serianthes melanesica var. yunkeri Fosberg in Reinwardtia 5: 313 (1960).

Serianthes robinsonii Fosberg in Reinwardtia 5: 301 (1960).

Serianthes sachetiae Fosberg in Reinwardtia 5: 310 (1960).

Tephrosia purpurea var. piscatoria (Aiton) Fosberg in Micronesica 25: 187 (1992).

Flacourtiaceae
Flacourtia ramontchii var. renvoizei Fosberg in Kew Bull. 29: 254 (1974).
Flacourtia rukam var. micronesica Fosberg & Sachet in Smithsonian Contr. Bot. 45:
14 (1980).
Xylosma smithiana Fosberg in Allertonia 7: 66 (1993).

Gentianaceae

Fagraea berterana vat. galilai (Gilg & Benedict) Fosberg in Smithsonian Contr. Bot.
45: 20 (1980).

Fagraea berterana var. kusaiana (Hosok.) Fosberg in Smithsonian Contr. Bot. 45: 20
(1980).

Fagraea berterana var. ladronica Fosberg in Smithsonian Contr. Bot. 45: 20 (1980).

Fagraea berterana var. pogas (Hosok.) Fosberg in Smithsonian Contr. Bot. 45: 21
(1980).

Fagraea berterana var. sair (Gilg & Benedict) Fosberg in Smithsonian Contr. Bot. 45:
21 (1980) without exact basionym page.

Geraniaceae

Geranium carolinianum var. australe (Benth.) Fosberg in Occas. Pap. Univ. Hawaii
3226, (1937):

Geranium cuneatum f. rockii (Skottsb.) Fosberg in Occas. Pap. Bernice P. Bishop
Mus. 12(16): 11 (1936).

Geranium cuneatum var. menziesii (A. Gray) Fosberg in Occas. Pap. Bernice P.
Bishop Mus. 12(16): 11 (1936).

Geranium cuneatum var. tridens (Hillebr.) Fosberg in Occas. Pap. Bernice P. Bishop
Mus. 12(16): 14 (1936).

18

Geranium humile var. mauiense Fosberg in Occas. Pap. Bernice P. Bishop Mus. 12(16):
18 (1936).

Geranium multiflorum var. ovatifolium (A. Gray) Fosberg in Occas. Pap. Bernice P.
Bishop Mus. 16(15): 339 (1942).

Gesneriaceae

Cyrtandra ootensis var. fatuhivensis Fosberg & Sachet in Smithsonian Contr. Bot. 47:
30 (1981).

Cyrtandra ootensis var. fatuhivensis Fosberg & Sachet in Smithsonian Contr. Bot. 47:
30 (1981).

Cyrtandra ootensis var. mollissima Fosberg & Sachet in Smithsonian Contr. Bot. 47:
30 (1981).

Cyrtandra ootensis var. mollissima Fosberg & Sachet in Smithsonian Contr. Bot. 47:
30 (1981).

Cyrtandra ootensis var. quaylei Fosberg & Sachet in Smithsonian Contr. Bot. 47: 31
(1981).

Cyrtandra ootensis var. quaylei Fosberg & Sachet in Smithsonian Contr. Bot. 47: 31
(1981).

Cyrtandra revoluta Fosberg & Sachet in Smithsonian Contr. Bot. 47: 31 (1981).

Cyrtandra tahuatensis Fosberg & Sachet in Smithsonian Contr. Bot. 47: 31 (1981).

Cyrtandra thibaultii Fosberg & Sachet in Smithsonian Contr. Bot. 47: 28 (1981).

Goodeniaceae
Scaevola paulayi Fosberg in Micronesica 23: 3 (1990).
Scaevola sericea var tuamotuensis (St. John) Fosberg in Taxon 10: 226 (1961).

Hernandiaceae
Hernandia sonora var. nymphaeifolia (Presl) Fosberg in Micronesica 23: 142 (1990).

Hypericaceae
Hypericum degeneri Fosberg in Occas. Pap. Bernice P. Bishop Mus. 24(2): 21 (1969).
Hypericum hypericoides var. multicaule (Michx.) Fosberg in Castanea 30: 202 (1965).

Juncaceae
Luzula campestris var. hawaiiensis (Buch.) Degener & Fosberg in Degener, Fl. Haw.
Fam. 66 Juncac.. [unpaged] (1937).
Luzula campestris var. oahuensis Degener & Fosberg in Degener, Fl. Haw. Fam. 66
Juncac.. [unpaged] (1937).

Lamiaceae (Labiatae)
Monarda fistulosa var. brevis Fosberg & Artz in Castanea 18: 130 (1953).
Phyllostegia mollis var. resinosa Fosberg in Occas. Pap. Bernice P. Bishop Mus.
16(15): 345 (1942).

19

Lauraceae
Cryptocarya oahuensis (Degener) Fosberg in Occas. Pap. Bernice P. Bishop Mus.
2 GS) 3.311936):

Liliaceae

Dianella saffordiana Fosberg & Sachet in Micronesica 20: 132 (1988 "1987").

Dianella sandwicensis var. lavarum (Degener) Fosberg in Occas. Pap. Bernice P.
Bishop Mus. 24(2): 12 (1969).

Dianella sandwicensis var. multipedicellata (Degener) Fosberg in Occas. Pap. Bernice
P. Bishop Mus. 24(2): 12 (1969).

Dracaena hawaiiensis (Degener) Fosberg in Occas. Pap. Bernice P. Bishop Mus.
23(2): 32 (1962).

Loganiaceae
Fagraea berterana var. marquisensis Fosberg & Sachet in Phytologia 28: 471 (1974).
Geniostoma gagnae Fosberg & Sachet in Smithsonian Contr. Bot. 47: 18 (1981).
Geniostoma hallei Fosberg & Sachet in Smithsonian Contr. Bot. 21: 13 (1975).
Geniostoma hallei var. fatuivense Fosberg & Sachet in Smithsonian Contr. Bot. 47: 19
(1981).
Geniostoma hallei var. hivaoense Fosberg & Sachet in Smithsonian Contr. Bot. 47: 19
(1981).
Geniostoma micranthum var. hoeferi (Gilg & Benedict) Fosberg in Smithsonian Contr.
Bot. 45: 23 (1980).
Geniostoma micranthum var. paganense Fosberg in Smithsonian Contr. Bot. 45: 23
(1980).
Geniostoma quadrangulare Fosberg in Smithsonian Contr. Bot. 21: 13 (1975).
Geniostoma rarotongensis Fosberg & Sachet in Smithsonian Contr. Bot. 47: 20 (1981).
Geniostoma sykesii Fosberg & Sachet in Smithsonian Contr. Bot. 47: 19 (1981).

Loranthaceae
Phoradendron bolleanum var. capitellatum (Trel.) Fosberg in Lloydia 4: 276 (1941).
Phoradendron bolleanum var. densum (Trel.) Fosberg in Lloydia 4: 276 (1941).
Phoradendron bolleanum var. pauciflorum (Torr.) Fosberg in Lloydia 4: 276 (1941).
Phoradendron flavescens (Nutt.) Fosberg in Lloydia 4: 279 (1941).
Phoradendron juniperinum var. ligatum (Trel.) Fosberg in Lloydia 4: 274 (1941).

Malvaceae
Abutilon asiaticum var. albescens (Miq.) Fosberg in Micronesica 2: 149 (1966).
Abutilon asiaticum var. australiense (Britten) Fosberg in Micronesica 2: 150 (1966).
Abutilon asiaticum var. subasperum Fosberg in Micronesica 2: 149 (1966).
Abutilon asiaticum var. supraviride Fosberg in Micronesica 2: 149 (1966).
Abutilon mangarevicum Fosberg in Smithsonian Contr. Bot. 47: 9 (1981).
Abutilon pitcairnense Fosberg in Smithsonian Contr. Bot. 47: 10 (1981).
Abutilon sachetianum Fosberg in Smithsonian Contr. Bot. 47: 7 (1981).
Abutilon whistleri Fosberg in Smithsonian Contr. Bot. 47: 10 (1981).

20

Hibiscus australensis Fosberg in Micronesica 2: 156 (1966).

Lebronnecia Fosberg in Adansonia, n. s. 6: 509 (1966).

Lebronnecia kokioides Fosberg in Adansonia, n. s. 6: 510 (1966).

Pavonia fryxelliana Fosberg & Sachet in Ceylon J. Sci., Biol. Sci. 15: 38 (1985 "1982").

Pavonia lourteigiae Fosberg & Sachet in Bull. Mus. Natl. Hist. Nat., B, Adansonia 1: 18
(1981).

Melastomataceae
Medinilla medinilliana (Gaudich.) Fosberg & Sachet in Smithsonian Contr. Bot. 45: 15
(1980).
Medinilla nodosa Fosberg in Lloydia 3: 115 (1940).
Melastoma malabathricum var. mariannum (Naudin) Fosberg & Sachet in Smithsonian
Contr. Bot. 45: 15 (1980).

Meliaceae
Malleastrum leroyi Fosberg in Kew Bull. 29: 255 (1974).

Moraceae
Artocarpus altilis (Parkinson) Fosberg in J. Wash. Acad. Sci. 31: 95 (1941).
Batocarpus amazonicus (Ducke) Fosberg in Proc. Biol. Soc. Wash. 55: 101 (1942).
Ficus prolixa var. carolinensis (Warb.) Fosberg in Phytologia 5: 289 (1955).
Ficus prolixa var. saffordii (Merr.) Fosberg in Phytologia 5: 289 (1955).
Ficus tinctoria var. neoebudarum (Summerh.) Fosberg in Phytologia 5: 289 (1955).
Maillardia pendula Fosberg in Kew Bull. 29: 266 (1974).

Myrsinaceae

Maesa canfieldiae Fosberg & Sachet in Phytologia 44: 364 (1979).

Maesa carolinensis var. kusaiensis Fosberg & Sachet in Phytologia 44: 366 "365".
(1979).

Maesa walkeri Fosberg & Sachet in Phytologia 44: 368 (1979).

Myrsine adamsonii Fosberg & Sachet in Smithsonian Contr. Bot. 21: 4 (1975).

Myrsine andersonii Fosberg & Sachet in Smithsonian Contr. Bot. 21: 4 (1975).

Myrsine brownii Fosberg & Sachet in Smithsonian Contr. Bot. 21: 5 (1975).

Myrsine carolinensis (Mez) Fosberg & Sachet in Smithsonian Contr. Bot. 45: 17
(1980).

Myrsine cheesemanii (Mez) Fosberg & Sachet in Smithsonian Contr. Bot. 21: 5
(1975):

Myrsine fasciculata (J. W. Moore) Fosberg & Sachet in Smithsonian Contr. Bot. 21: 6
(1975).

Myrsine fusca (J. W. Moore) Fosberg & Sachet in Smithsonian Contr. Bot. 21: 6
(1975).

Myrsine gracilissima Fosberg & Sachet in Smithsonian Contr. Bot. 21: 6 (1975).

Myrsine grantii Fosberg & Sachet in Smithsonian Contr. Bot. 21: 7 (1975).

Myrsine grantii var. toviiensis Fosberg & Sachet in Smithsonian Contr. Bot. 21: 8
(1975).

21

Myrsine hartii (M. L. Grant) Fosberg & Sachet in Smithsonian Contr. Bot. 21: 8 (1975).

Myrsine ledermannii (Mez) Fosberg & Sachet in Smithsonian Contr. Bot. 45: 17 (1980).

Myrsine naiuensis Fosberg & Sachet in Smithsonian Contr. Bot. 21: 8 (1975).

Myrsine nukuhivensis Fosberg & Sachet in Smithsonian Contr. Bot. 21: 9 (1975).

Myrsine obovata (J. W. Moore) Fosberg & Sachet in Smithsonian Contr. Bot. 21: 9
(ISS)

Myrsine orohenensis (J. W. Moore) Fosberg & Sachet in Smithsonian Contr. Bot. 21: 9
(1975).

Myrsine ovalis var. wilderi Fosberg & Sachet in Smithsonian Contr. Bot. 21: 10 (1975).

Myrsine palauensis (Mez) Fosberg & Sachet in Smithsonian Contr. Bot. 45: 17 (1980).

Myrsine raiateensis (J. W. Moore) Fosberg & Sachet in Smithsonian Contr. Bot. 21: 10
(1975).

Myrsine rapensis (F. Br.) Fosberg & Sachet in Smithsonian Contr. Bot. 21: 10 (1975).

Myrsine ronuiensis (M. L. Grant) Fosberg & Sachet in Smithsonian Contr. Bot. 21: 10
(1975).

Myrsine st.-johnii (M. L. Grant) Fosberg & Sachet in Smithsonian Contr. Bot. 21: 10
COWS).

Myrsine tahuatensis Fosberg & Sachet in Smithsonian Contr. Bot. 21: 10 (1975).

Myrtaceae

Eugenia elliptica var. levinervis Fosberg in Kew Bull. 33: 134 (1978).

Pimenta racemosa vat. grisea (Kiaersk.) Fosberg in Amer. Midl. Naturalist 27: 762
(1942).

Psidium cattleianum var. littorale (Raddi) Fosberg in Occas. Pap. Bernice P. Bishop
Mus. 23(2): 37 (1962).

Psidium littorale var. longipes (Berg) Fosberg in Proc. Biol. Soc. Washington 54: 180
(1941).

Psidium littorale var. lucidum (Degener) Fosberg in Proc. Biol. Soc. Washington 54:
180 (1941).

Nesogenaceae or Verbenaceae
Nesogenes euphrasioides var. lineata Fosberg in Micronesica 23: 2 (1990).
Nesogenes rotensis Fosberg & D. Herbst in Micronesica 19: 12 (1984 "1983").

Nyctaginaceae
Boerhavia albiflora Fosberg in Smithsonian Contr. Bot. 39: 11 (1978).
Boerhavia albiflora var. heronensis Fosberg in Brittonia 40: 59 (1988).
Boerhavia albiflora var. powelliae Fosberg in Smithsonian Contr. Bot. 39: 12 (1978).
Boerhavia crispifolia Fosberg in Smithsonian Contr. Bot. 39: 15 (1978).
Boerhavia fistulosa Fosberg in Brittonia 40: 57 (1988).
Boerhavia fistulosa var. puberuliflora Fosberg in Brittonia 40: 59 (1988).
Boerhavia herbstii Fosberg in Smithsonian Contr. Bot. 39: 17 (1978).
Boerhavia repens var. maris-indici Fosberg in Kew Bull. 33: 398 (1979) with type; et
in Smithsonian Contr. Bot. 39: 9 (1978) without type.

22

Boerhavia repens var. pubescens (R. Br.) Fosberg in Smithsonian Contr. Bot. 39: 6
(1978).

Boerhavia rubicunda var. stenophylla (Boissier) Fosberg in Smithsonian Contr. Bot. 39:
14 (1978).

Pisonia wagneriana Fosberg in Phytologia 62: 177 (1987).

Oleaceae
Chionanthus sessiliflorus (Hemsl.) Fosberg in Micronesica 25: 197 (1992).

Onagraceae
Oenothera heterophylla var. curtissii (Rose) Fosberg in Amer. Midl. Naturalist 27: 763
(1942).
Oenothera heterophylla var. rhombipetala (Nutt.) Fosberg in Amer. Midl. Naturalist
27: 763 (1942).

Orchidaceae
Anoectochilus apiculatus L. O. Williams & Fosberg in Bull. Torrey Bot. Club 70: 387
(1943).
Habenaria setifera (Tuyama) Fosberg & Sachet in Micronesica 20: 142 (1988 "1987").
Liparis yamadae (Tuyama) Fosberg & Sachet in Micronesica 20: 141 (1988 "1987").
Malaxis calcarea (Schltr.) Fosberg & Sachet in Micronesica 20: 142 (1988 "1987").
Malaxis kerstingiana (Schltr.) Fosberg & Sachet in Micronesica 20: 143 (1988
PSST):
Malaxis trukensis (Fukuy.) Fosberg & Sachet in Micronesica 20: 143 (1988 "1987").
Malaxis volkensii (Schltr.) Fosberg & Sachet in Micronesica 20: 143 (1988 "1987").
Nervilia jacksoniae Rinehart & Fosberg in Micronesica 24: 81 (1991)
Rhynchophreatia carolinensis (Schltr.) Fosberg & Sachet in Micronesica 20: 152 (1988
"1987").
Rhynchophreatia pacifica (Fukuy.) Fosberg & Sachet in Micronesica 20: 152 (1988
POS7"):

Oxalidaceae
Oxalis gagneorum Fosberg & Sachet in Smithsonian Contr. Bot. 47: 3 (1981).

Passifloraceae
Passiflora suberosa var. perhastata Fosberg in Micronesica 25: 193 (1992).

Piperaceae
Peperomia mariannensis f. saipana (C. DC.) Fosberg in Phytologia 13: 240 (1966).

Peperomia palauensis var occidentalis Fosberg in Smithsonian Contr. Bot. 24: 13
(1975).

Peperomia pallida var anderssonii (Yunck.) Fosberg & Sachet in Micronesica 8: 43
(1972).

23

Peperomia ponapensis var. trukensis (Yunck.) Fosberg in Phytologia 13: 241 (1966).
Piper betle f. densum (Blume) Fosberg in Phytologia 13: 235 (1966).

Piper betle f. marianum (Opiz) Fosberg in Phytologia 13: 236 (1966).

Piper guahamense f. glabrum (Yunck.) Fosberg in Phytologia 13: 235 (1966).

Piper hosokawae Fosberg in Phytologia 13: 237 (1966).

Poaceae (Gramineae)
Cynodon dactylon var. parviglumis (Ohwi) Fosberg & Sachet in Micronesica 18(2): 45
(1984 "1982").
Digitaria pruriens var. microbachne (Presl) Fosberg in Phytologia 5: 289 (1955).
Eragrostis whitneyi Fosberg in Occas. Pap. Bernice P. Bishop Mus. 15(3): 39 (1939).
Eragrostis whitneyi var caumii Fosberg in Occas. Pap. Bernice P. Bishop Mus. 15(3):
41 (1939).
Isachne confusa var. purpurascens (Glassman) Fosberg & Sachet in Micronesica 18(2):
53 (1984 "1982").
Isachne miliacea var. minutula (Gaudich.) Fosberg & Sachet in Micronesica 18(2): 55
(1984 "1982").
Ischaemum longisetum var. raulersoniae Fosberg & Sachet in Micronesica 18(2): 62
(1984 "1982").
Ischaemum muticum vat. aristuliferum Fosberg & Sachet in Micronesica 18(2): 64
(1984 "1982").
Ischaemum polystachyum var. chordatum (Trin.) Fosberg & Sachet in Micronesica
18(2): 66 (1984 "1982").
Ischaemum polystachyum var. hillii Fosberg & Sachet in Micronesica 18(2): 67 (1984
1982):
Ischaemum polystachyum var. intermedium (Brongn.) Fosberg & Sachet in Micronesica
18(2): 67 (1984 "1982").
Lepturopetium marshallense Fosberg & Sachet in Micronesica 18(2): 72 (1984
"1982").
Lepturus gasparricensis Fosberg in Occas. Pap. Bernice P. Bishop Mus. 21(14): 293
@i955):
Lepturus repens var. cinereus (Burcham) Fosberg in Occas. Pap. Bernice P. Bishop
Mus. 21(14): 292 (1955).
Lepturus repens var. occidentalis Fosberg in Occas. Pap. Bernice P. Bishop Mus.
21(14): 291 (1955).
Lepturus repens var. septentrionalis Fosberg in Occas. Pap. Bernice P. Bishop Mus.
21(14): 291 (1955).
Lepturus repens var. subulatus Fosberg in Occas. Pap. Bernice P. Bishop Mus. 21(14):
290 (1955).
Lepturus stoddartii Fosberg in Brittonia 40: 53 (1988).
Oplismenus hirtellus var. imbecillis (R. Br.) Fosberg & Sachet in Micronesica 18(2):
78 (1984 "1982").
Oplismenus hirtellus var. microphyllus (Honda) Fosberg & Sachet in Micronesica
18(2): 79 (1984 "1982").
Panicum reptans var. burmanicum (Hook. f.) Fosberg in Micronesica 2: 154 (1966).

24

Panicum reptans var. marquisense (F. Br.) Fosberg in Micronesica 2: 154 (1966).

Paspalum cartilagineum var. biglumaceum Fosberg & Sachet in Micronesica 18(2): 81
(1984 "1982").

Paspalum orbiculare var. otobedii Fosberg & Sachet in Micronesica 18(2): 83 (1984
S820):

Pennisetum polystachion f. viviparum Fosberg & Sachet in Micronesica 18(2): 86
(1984 "1982").

Saccharum spontaneum var. insulare (Brongn.) Fosberg & Sachet in Micronesica
18(2): 89 (1984 "1982").

Sorghum bicolor var. cafer (Koern.) Fosberg & Sachet in Micronesica 18(2): 93 (1984
"1982").

Sorghum bicolor var. obovatum (Hack.) Fosberg & Sachet in Micronesica 18(2): 94
(1984 "1982").

Sorghum bicolor var. rotundulum (Snowden) Fosberg & Sachet in Micronesica 18(2):
94 (1984 "1982").

Sorghum bicolor var. subglabrescens (Steud.) Fosberg & Sachet in Micronesica 18(2):
95 (1984 "1982").

Sorghum bicolor var. transiens (Hack.) Fosberg & Sachet in Micronesica 18(2): 96
(1984 "1982").

Tridens flavus f. cupreus (Jacq.) Fosberg in Castanea 20: 58 (1955).

Triodia flava f. cuprea (Jacq.) Fosberg in Castanea 11: 67 (1946).

Polemoniaceae
Gilia aggregata var. arizonica (Greene) Fosberg in Amer. Midl. Naturalist 37: 764
(1942).

Polygonaceae

Eriogonum abertianum var. bracteatum Fosberg in Madrono 4: 192 (1938).
Eriogonum abertianum var. cyclosepalum (Greene) Fosberg in Madrofio 4: 192 (1938).
Eriogonum abertianum var. lappulaceum Fosberg in Madrofio 4: 193 (1938).
Eriogonum abertianum var. villosum Fosberg in Madronio 4: 191 (1938).
Eriogonum abertianum var.gillespiei Fosberg in Madrofio 4: 193 (1938).
Rumex giganteus f. coccineus Fosberg in Occas. Pap. Bernice P. Bishop Mus. 24(2):

16 (1969).

Portulacaceae
Lewisia pygmaea var. nevadensis (A. Gray) Fosberg in Amer. Midl. Naturalist 27: 256
(1942).
Portulaca mauritiensis var. aldabrensis Fosberg in Kew Bull. 32: 256 (1977).
Portulaca mauritiensis var. grubbii Fosberg in Kew Bull. 32: 258 (1977).
Portulaca oleracea var. delicatula Fosberg in Kew Bull. 32: 254 (1977).

73)

Potamogetonaceae
Ruppia maritima f. curvirostris St. John & Fosberg in Occas. Pap. Bernice P. Bishop
Mus. 15(16): 178 (1939).
Ruppia maritima f. pacifica St. John & Fosberg in Occas. Pap. Bernice P. Bishop Mus.
15(16): 177 (1939).
Ruppia maritima var. pacifica St. John & Fosberg in Occas. Pap. Bernice P. Bishop Mus.
15(16): 176 (1939).

Ranunculaceae
Anemone quinquefolia var. lancifolia (Pursh) Fosberg in Virginia J. Sci. 2: 108 (1941).

Rhamnaceae
Ceanothus dentatus var. dickeyi Fosberg in Amer. Midl. Naturalist 27: 256 (1942).
Gouania mangarevica Fosberg in Bull. Torrey Bot. Club 65: 607 (1939).
Rhamnus smithii var. mulleri Fosberg in Lloydia 4: 286 (1941).

Rhizophoraceae
Bruguiera gymnorhiza f. alba (B. C. Stone) Fosberg in Smithsonian Contr. Bot. 45: 15
(1980).
Rosaceae
Spiraea alba var. septentrionalis (Fern.) Fosberg in Castanea 30: 197 (1965).
Spiraea betulifolia f. campii Fosberg in Virginia J. Sci. 2: 109 (1941).

Rubiaceae

Subtr. Coprosminae Fosberg in Acta Phytotax. Geobot. 33: 75 (1982).

Bikkia mariannensis var. longicarpa (Valeton) Fosberg in Occas. Pap. Bernice P.
Bishop Mus. 15(20): 213 (1990).

Bobea platypes Fosberg in Phytologia 62: 179 (1987).

Calycosia macrocyatha Fosberg in Sargentia 1: 139 (1942).

Calycosia macrocyatha var. kandavuensis Fosberg in Sargentia 1: 140 (1942).

Canthium barbatum f. calcicola Fosberg in Occas. Pap. Bernice P. Bishop Mus.
13(19): 261 (1937).

Canthium barbatum f. pitcairnense Fosberg in Occas. Pap. Bernice P. Bishop Mus.
13(19): 260 (1937).

Canthium barbatum f. rurutu Fosberg in Occas. Pap. Bernice P. Bishop Mus. 13(19):
DOOIGIOS TI):

Canthium barbatum f. tubuai Fosberg in Occas. Pap. Bernice P. Bishop Mus. 13(19):
259 (1937).

Canthium barbatum var. australense Fosberg in Occas. Pap. Bernice P. Bishop Mus.
13(19): 258 (1937).

Canthium barbatum var. christianii Fosberg in Occas. Pap. Bernice P. Bishop Mus.
13(19): 260 (1937).

Canthium barbatum var. gambierense Fosberg in Occas. Pap. Bernice P. Bishop Mus.
13(19): 261 (1937).

26

Canthium barbatum var. huahinense Fosberg in Occas. Pap. Bernice P. Bishop Mus.
1S@9) 725701937):

Canthium barbatum var. korrorense (Valeton) Fosberg in Occas. Pap. Bernice P.
Bishop Mus. 15(20): 218 (1940).

Canthium barbatum var. raiateense (J. W. Moore) Fosberg in Occas. Pap. Bernice P.
Bishop Mus. 13(19): 258 (1937).

Canthium barbatum var. raivavaense Fosberg in Occas. Pap. Bernice P. Bishop Mus.
1319): 25951937):

Canthium barbatum var. rapae (Riley) Fosberg in Occas. Pap. Bernice P. Bishop Mus.
13(19): 260 (1937).

Canthium barbatum var. rupestre (Hosok.) Fosberg in Smithsonian Contr. Bot. 81: 52
(1993).

Canthium barbatum var. societense Fosberg in Occas. Pap. Bernice P. Bishop Mus.
13(19): 256 (1937).

Canthium barbatum var. temehaniense (J. W. Moore) Fosberg in Occas. Pap. Bernice
P. Bishop Mus. 13(19): 258 (1937).

Canthium odoratum var. tinianense (Kaneh.) Fosberg in Phytologia 5: 291 (1955).

Coprosma cookei Fosberg in Occas. Pap. Bernice P. Bishop Mus. 13(19): 289 (1937).

Coprosma esulcata (F. Br.) Fosberg in Brittonia 8: 178 (1956).

Coprosma longifolia var. oppositifolia Fosberg in Bull. Torrey Bot. Club 70: 394
(1943).

Coprosma oliveri Fosberg in Brittonia 20: 292 (1968).

Coprosma rapensis var. benefica (Oliver) Fosberg in Occas. Pap. Bernice P. Bishop
Mus. 13(19): 288 (1937).

Coprosma rapensis var. mangarevica Fosberg in Occas. Pap. Bernice P. Bishop Mus.
13(19): 288 (1937).

Coprosma taitensis var. glabrata (J. W. Moore) Fosberg in Occas. Pap. Bernice P.
Bishop Mus. 13(19): 284 (1937).

Coprosma taitensis var. oliverii Fosberg in Occas. Pap. Bernice P. Bishop Mus.
13(19): 284 (1937).

Coprosma taitensis var. raiateensis (J. W. Moore) Fosberg in Occas. Pap. Bernice P.
Bishop Mus. 13(19): 284 (1937).

Coprosma velutina Fosberg in Occas. Pap. Bernice P. Bishop Mus. 13(19): 285
(1937).

Coprosma velutina var. andersonii Fosberg in Occas. Pap. Bernice P. Bishop Mus.
13(19): 286 (1937).

Dolicholobium oblongifolium var. degeneri Fosberg in Sargentia 1: 119 (1942).

Dolicholobium oblongifolium var. longissimum (Seem.) Fosberg in Sargentia 1: 119
(1942).

Galium hypocarpium (L.) Fosberg in Sida 2: 386 (1966).

Geophila repens var. asiatica (Cham. & Schtdl.) Fosberg in Smithsonian Contr. Bot.
45: 27 (1980).

Gonzalagunia longithyrsa Fosberg in Sida 2: 387 (1966).

Gouldia hillebrandii Fosberg in Bernice P. Bishop Mus. Bull. 147: 59 (1937).

27

Gouldia hillebrandii f. lancifolia Fosberg in Bernice P. Bishop Mus. Bull. 147: 61
(1937).
Gouldia hillebrandii var. hawaiiensis Fosberg in Bernice P. Bishop Mus. Bull. 147: 61
(19377):
Gouldia hillebrandii var. nodosa Fosberg in Bernice P. Bishop Mus. Bull. 147: 60
(1937).
Gouldia st.-johnii Fosberg in Bernice P. Bishop Mus. Bull. 147: 62 (1937).
Gouldia st.-johnii var. munroi Fosberg in Bernice P. Bishop Mus. Bull. 147: 63
(1937).
Gouldia terminalis f. acuminata Fosberg in Bernice P. Bishop Mus. Bull. 147: 45
(11937):
Gouldia terminalis f. acuta Fosberg in Bernice P. Bishop Mus. Bull. 147: 55 (1937).
Gouldia terminalis f. albicaulis Fosberg in Bernice P. Bishop Mus. Bull. 147: 32
(1937).
Gouldia terminalis f. cuneata Fosberg in Bernice P. Bishop Mus. Bull. 147: 34 (1937).
Gouldia terminalis f. heterophylla Fosberg in Bernice P. Bishop Mus. Bull. 147: 52
@937):
Gouldia terminalis f. hirtellicostata Fosberg in Bernice P. Bishop Mus. Bull. 147: 33
(1937).
Gouldia terminalis f. hirtellifolia Fosberg in Bernice P. Bishop Mus. Bull. 147: 55
@1937):
Gouldia terminalis f. impressa Fosberg in Bull. Torrey Bot. Club 70: 391 (1943).
Gouldia terminalis f. kahili Fosberg in Bernice P. Bishop Mus. Bull. 147: 33 (1937).
Gouldia terminalis f. kalaupapa Fosberg in Bernice P. Bishop Mus. Bull. 147: 51
(1937).
Gouldia terminalis f. kauensis Fosberg in Bernice P. Bishop Mus. Bull. 147: 55
(1937).
Gouldia terminalis f. kehena Fosberg in Bernice P. Bishop Mus. Bull. 147: 54 (1937).
Gouldia terminalis f. latifolia Fosberg in Bernice P. Bishop Mus. Bull. 147: 38 (1937).
Gouldia terminalis f. lydgatei Fosberg in Bernice P. Bishop Mus. Bull. 147: 53
(1937).
Gouldia terminalis f. macrophylla Fosberg in Bernice P. Bishop Mus. Bull. 147: 32
(1937).
Gouldia terminalis f. makawaoensis Fosberg in Bernice P. Bishop Mus. Bull. 147: 50
(1937).
Gouldia terminalis f. maunahui Fosberg in Bernice P. Bishop Mus. Bull. 147: 51
(937):
Gouldia terminalis f. membranacea Fosberg in Bernice P. Bishop Mus. Bull. 147: 53
(1937).
Gouldia terminalis f. molokaiensis Fosberg in Bernice P. Bishop Mus. Bull. 147: 45
(1937).
Gouldia terminalis f. nealiae Fosberg in Bernice P. Bishop Mus. Bull. 147: 45 (1937).
Gouldia terminalis f. oahuensis Fosberg in Brittonia 8: 176 (1956).
Gouldia terminalis f. oblonga Fosberg in Brittonia 8: 176 (1956).

28

Gouldia terminalis f. obovata Fosberg in Bull. Torrey Bot. Club 70: 391 (1943).

Gouldia terminalis f. parvithyrsa Fosberg in Bernice P. Bishop Mus. Bull. 147: 37
(1937).

Gouldia terminalis f. petiolata Fosberg in Bernice P. Bishop Mus. Bull. 147: 53
(11937):

Gouldia terminalis f. pittosporoides Fosberg in Bernice P. Bishop Mus. Bull. 147: 39
(937):

Gouldia terminalis f. punaula Fosberg in Bernice P. Bishop Mus. Bull. 147: 53
(ISEB).

Gouldia terminalis f. rigidifolia Fosberg in Bernice P. Bishop Mus. Bull. 147: 39
(1937).

Gouldia terminalis f. rigidifolioides Fosberg in Brittonia 8: 174 (1956).

Gouldia terminalis f. robusta Fosberg in Bernice P. Bishop Mus. Bull. 147: 29 (1937).

Gouldia terminalis f. rockii Fosberg in Bernice P. Bishop Mus. Bull. 147: 43 (1937).

Gouldia terminalis f. russii Fosberg in Bernice P. Bishop Mus. Bull. 147: 49 (1937).

Gouldia_ terminalis f. sambucina (Heller) Fosberg in Bernice P. Bishop Mus. Bull.
147: 36 (1937).

Gouldia terminalis f. santalifolia Fosberg in Bernice P. Bishop Mus. Bull. 147: 51
(1937).

Gouldia terminalis f. sclerophylla Fosberg in Bernice P. Bishop Mus. Bull. 147: 35
(1937).

Gouldia terminalis f. storeyi Fosberg in Bernice P. Bishop Mus. Bull. 147: 52 (1937).

Gouldia terminalis f. subpilosa Fosberg in Bernice P. Bishop Mus. Bull. 147: 56
(1937).

Gouldia terminalis f. suehiroae Fosberg in Bernice P. Bishop Mus. Bull. 147: 52
(1937).

Gouldia terminalis f. teres Fosberg in Bernice P. Bishop Mus. Bull. 147: 36 (1937).

Gouldia terminalis f. violetae Fosberg in Bull. Torrey Bot. Club 70: 392 (1943).

Gouldia terminalis f. wailauensis Fosberg in Bernice P. Bishop Mus. Bull. 147: 52
C1939):

Gouldia terminalis f. waipioensis Fosberg in Bernice P. Bishop Mus. Bull. 147: 37
(1937).

Gouldia terminalis var. angustifolia Fosberg in Bernice P. Bishop Mus. Bull. 147: 43
(i937):

Gouldia terminalis var. antiqua Fosberg in Bernice P. Bishop Mus. Bull. 147: 54
(1937).

Gouldia terminalis var. arborescens (Wawra) Fosberg in Bernice P. Bishop Mus. Bull.
147: 31 (1937).

Gouldia terminalis var. aspera Fosberg in Brittonia 8: 175 (1956).

Gouldia terminalis var. bodeoides Fosberg in Bernice P. Bishop Mus. Bull. 147: 37
(1937).

Gouldia terminalis var. congesta Fosberg in Bernice P. Bishop Mus. Bull. 147: 55
(1937).

Gouldia terminalis var. cordata (Wawra) Fosberg in Bernice P. Bishop Mus. Bull.
147: 44 (1937).

Gouldia terminalis var.
147: 44 (1937).
Gouldia terminalis var.

Bull. 147: 47 (1937).

Gouldia terminalis var.
(1937).

Gouldia terminalis var.
(1937).

Gouldia terminalis var.
(1937).

29

cordata (Wawra) Fosberg in Bernice P. Bishop Mus. Bull.
coriacea (Hook. & Arn.) Fosberg in Bernice P. Bishop Mus.
crassicaulis Fosberg in Bernice P. Bishop Mus. Bull. 147: 54
degeneri Fosberg in Bernice P. Bishop Mus. Bull. 147: 39

elongata Fosberg in Bernice P. Bishop Mus. Bull. 147: 32

Gouldia terminalis var. forbesii Fosberg in Bernice P. Bishop Mus. Bull. 147: 57

(1937).

Gouldia terminalis var.
(1937).

Gouldia terminalis var.

Gouldia terminalis var.
(1937).

Gouldia terminalis var.

Gouldia terminalis var.
(1937).

Gouldia terminalis var.
(1937).
Gouldia terminalis var.
Gouldia terminalis var.
147: 34 (1937).
Gouldia terminalis var.
(1937).

Gouldia terminalis var.

Gouldia terminalis var.
(1937).

Gouldia terminalis var.
50 (1937).

Gouldia terminalis var.
147: 56 (1937).
Gouldia terminalis var.

(1937).
Gouldia terminalis var.
147: 58 (1937).
Gouldia terminalis var.
(1937).

Gouldia terminalis var.

Gouldia terminalis var.
(1937).

Gouldia terminalis var.
58 (1937).

glabra Fosberg in Bernice P. Bishop Mus. Bull. 147: 36

hathawayi Fosberg in Brittonia 8: 174 (1956).
hosakai Fosberg in Bernice P. Bishop Mus. Bull. 147: 38

kaala Fosberg in Bernice P. Bishop Mus. Bull. 147: 49 (1937).
kapuensis Fosberg in Bernice P. Bishop Mus. Bull. 147: 39

konaensis Fosberg in Bernice P. Bishop Mus. Bull. 147: 37

lanai Fosberg in Bernice P. Bishop Mus. Bull. 147: 59 (1937).
macrocarpa (Hillebr.) Fosberg in Bernice P. Bishop Mus. Bull.

macrothyrsa Fosberg in Bernice P. Bishop Mus. Bull. 147: 40

myrsinoidea Fosberg in Brittonia 8: 176 (1956).
osteocarpa Fosberg in Bernice P. Bishop Mus. Bull. 147: 33

ovata (Wawra) Fosberg in Bernice P. Bishop Mus. Bull. 147:
parvifolia (Wawra) Fosberg in Bernice P. Bishop Mus. Bull.
pedunculata Fosberg in Bernice P. Bishop Mus. Bull. 147: 46
pseudodichotoma Fosberg in Bernice P. Bishop Mus. Bull.
pubescens Fosberg in Bernice P. Bishop Mus. Bull. 147: 57

pubistipula Fosberg in Brittonia 20: 291 (1968).
purpurea Fosberg in Bernice P. Bishop Mus. Bull. 147: 46

quadrangularis Fosberg in Bernice P. Bishop Mus. Bull. 147:

30

Gouldia terminalis var.

(1937).

Gouldia terminalis var.
Gouldia terminalis var.

(1937).

Gouldia terminalis var.

147: 42 (1937).

Gouldia terminalis var.

(1937).

Gouldia terminalis var.

(19317):

Gouldia terminalis var.

(1937).

rotundifolia Fosberg in Bernice P. Bishop Mus. Bull. 147: 41

sclerotica Fosberg in Brittonia 8: 175 (1956).
skottsbergii Fosberg in Bernice P. Bishop Mus. Bull. 147: 42

stipulacea (Wawra) Fosberg in Bernice P. Bishop Mus. Bull.
subcordata Fosberg in Bernice P. Bishop Mus. Bull. 147: 44
tenuicaulis Fosberg in Bernice P. Bishop Mus. Bull. 147: 57

wawrana Fosberg in Bernice P. Bishop Mus. Bull. 147: 30

Guettarda speciosa f. tahitensis (Nadeaud) Fosberg in Occas. Pap. Bernice P. Bishop

Mus. 13(19): 262 (

1937).

Gynochthodes ovalifolia var. smithii Fosberg in Bull. Torrey Bot. Club 67: 421 (1940).

Hedyotis sect
Hedyotis sect
Hedyotis sect
Hedyotis sect
Hedyotis sect
Hedyotis sect
(1943).

. Austrogouldia Fosberg in Bernice P. Bishop Mus. Bull. 174: 28 (1943).
. Bikkiocarpa Fosberg in Bernice P. Bishop Mus. Bull. 174: 25 (1943).

. Gouldiopsis Fosberg in Bernice P. Bishop Mus. Bull. 174: 56 (1943).

. Lignea Fosberg in Allertonia 6: 202 (1991).

. Protokadua Fosberg in Bernice P. Bishop Mus. Bull. 174: 23 (1943).

. Wiegmannia (Meyen) Fosberg in Bernice P. Bishop Mus. Bull. 174: 29

Hedyotis subg. Diplophragma (Wight & Arn.) Fosberg in Bernice P. Bishop Mus.
Bull. 174: 21 (1943).

Hedyotis subg. Kadua
174: 69 (1943).

(Cham. & Schltdl.) Fosberg in Bernice P. Bishop Mus. Bull.

Hedyotis subg. Oceanica Fosberg in Bernice P. Bishop Mus. Bull. 174: 67 (1943).
Hedyotis subg. Oldenlandia (L.) Fosberg in Bernice P. Bishop Mus. Bull. 174: 19

(1943).

Hedyotis subg. Polynesiotis Fosberg in Bernice P. Bishop Mus. Bull. 174: 22 (1943).

Hedyotis acuminata f.
(1943).

Hedyotis acuminata f.

Hedyotis acuminata f.

Hedyotis acuminata f.

Hedyotis acuminata f.
76 (1943).

Hedyotis acuminata f.
(1943).

Hedyotis acuminata f.

Hedyotis acuminata f.

Hedyotis acuminata f.
(1943).

alicarpa Fosberg in Bernice P. Bishop Mus. Bull. 174: 82

alicarpoides Fosberg in Brittonia 8: 171 (1956).

allanii Fosberg in Brittonia 8: 170 (1956).

basipuba Fosberg in Brittonia 20: 290 (1968).
brevimucronata Fosberg in Bernice P. Bishop Mus. Bull. 174:

brownii Fosberg in Bernice P. Bishop Mus. Bull. 174: 79
calycosa Fosberg in Brittonia 20: 289 (1968).

compacta Fosberg in Brittonia 8: 169 (1956).
cylindrica Fosberg in Bernice P. Bishop Mus. Bull. 174: 90

Sul

Hedyotis acuminata f. duripes Fosberg in Bernice P. Bishop Mus. Bull. 174: 84
(1943).

Hedyotis acuminata f. eamesii Fosberg in Bernice P. Bishop Mus. Bull. 174: 86
(1943).

Hedyotis acuminata f. elliptica Fosberg in Bernice P. Bishop Mus. Bull. 174: 79
(1943).

Hedyotis acuminata f. forbesii Fosberg in Bernice P. Bishop Mus. Bull. 174: 88
(1943).

Hedyotis acuminata f. grandis (A. Gray) Fosberg in Bernice P. Bishop Mus. Bull. 174:
90 (1943).

Hedyotis acuminata f. grayana Fosberg in Bernice P. Bishop Mus. Bull. 174: 79
(1943).

Hedyotis acuminata f. greenwelliae Fosberg in Brittonia 8: 170 (1956).

Hedyotis acuminata f. halawana Fosberg in Bernice P. Bishop Mus. Bull. 174: 76

(1943).

Hedyotis acuminata f. haupuensis Fosberg in Bernice P. Bishop Mus. Bull. 174: 76
(1943).

Hedyotis acuminata f. heteripes Fosberg in Bernice P. Bishop Mus. Bull. 174: 85
(1943).

Hedyotis acuminata f. hirtella Fosberg in Bernice P. Bishop Mus. Bull. 174: 80
(1943).

Hedyotis acuminata f. hirticalyx Fosberg in Bernice P. Bishop Mus. Bull. 174: 89
(1943).

Hedyotis acuminata f. honopui Fosberg in Brittonia 8: 171 (1956).

Hedyotis acuminata f. iaoensis Fosberg in Bernice P. Bishop Mus. Bull. 174: 87
(1943).

Hedyotis acuminata f. isae Fosberg in Brittonia 20: 289 (1968).

Hedyotis acuminata f. kaalae (Wawra) Fosberg in Bernice P. Bishop Mus. Bull. 174:
78 (1943).

Hedyotis acuminata f. koolauensis Fosberg in Bernice P. Bishop Mus. Bull. 174: 78
(1943).

Hedyotis acuminata f. lanaiensis Fosberg in Bernice P. Bishop Mus. Bull. 174: 85
(1943).

Hedyotis acuminata f. liberipes Fosberg in Bernice P. Bishop Mus. Bull. 174: 75
(1943).

Hedyotis acuminata f. microliberipes Fosberg in Brittonia 8: 169 (1956).

Hedyotis acuminata f. obovata Fosberg in Bernice P. Bishop Mus. Bull. 174: 88
(1943).

Hedyotis acuminata f. ovalicalyx Fosberg in Bernice P. Bishop Mus. Bull. 174: 88
(1943).

Hedyotis acuminata f. ovalifolia (A. Gray) Fosberg in Bernice P. Bishop Mus. Bull.
174: 87 (1943).

Hedyotis acuminata f. pilicalyx Fosberg in Bernice P. Bishop Mus. Bull. 174: 82
(1943).

32

Hedyotis acuminata f. quadricosta Fosberg in Bernice P. Bishop Mus. Bull. 174: 84
(1943). ;

Hedyotis acuminata f. sherffiana Fosberg in Bernice P. Bishop Mus. Bull. 174: 77
(1943).

Hedyotis acuminata f. skottsbergiana Fosberg in Bernice P. Bishop Mus. Bull. 174: 89
(1943).

Hedyotis acuminata f. sphaerocarpa Fosberg in Bernice P. Bishop Mus. Bull. 174: 76
(1943).

Hedyotis acuminata f. st.johnii Fosberg in Bernice P. Bishop Mus. Bull. 174: 85
(1943).

Hedyotis acuminata f. storeyi Fosberg in Bernice P. Bishop Mus. Bull. 174: 84 (1943).

Hedyotis acuminata f. subsessilis Fosberg in Bernice P. Bishop Mus. Bull. 174: 87
(1943).

Hedyotis acuminata f. toppingii Fosberg in Bernice P. Bishop Mus. Bull. 174: 77
(1943).

Hedyotis acuminata f. tuberculata Fosberg in Bernice P. Bishop Mus. Bull. 174: 86
(1943).

Hedyotis acuminata f. turbinata Fosberg in Bernice P. Bishop Mus. Bull. 174: 89

(1943).

Hedyotis acuminata f. variabilis Fosberg in Bernice P. Bishop Mus. Bull. 174: 82
(1943).

Hedyotis acuminata f. violetana Fosberg in Bernice P. Bishop Mus. Bull. 174: 81
(1943).

Hedyotis acuminata f. wailauensis Fosberg in Bernice P. Bishop Mus. Bull. 174: 85
(1943).

Hedyotis albido-punctata (Merr.) Fosberg in Lloydia 3: 123 (1940).

Hedyotis albidopunctata (Merr.) Fosberg in Lloydia 32: 123 (1940).

Hedyotis angulata Fosberg in Field & Lab. 17: 166 (1949).

Hedyotis angusta Fosberg in Bernice P. Bishop Mus. Bull. 174: 48 (1943).

Hedyotis angusta var. koolauensis Fosberg in Brittonia 8: 167 (1956).

Hedyotis angusta var. umbrosa Fosberg in Bernice P. Bishop Mus. Bull. 174: 49
(1943).

Hedyotis auricularia var. brevipila Fosberg & Sachet in Allertonia 6: 208 (1991).

Hedyotis auricularia var. dispersa Fosberg & Sachet in Allertonia 6: 208 (1991).

Hedyotis auricularia var. longipila Fosberg & Sachet in Allertonia 6: 209 (1991).

Hedyotis auricularia var. melanesica Fosberg in Bull. Torrey Bot. Club 67: 419
(1940).

Hedyotis auricularia var. tomentella Fosberg & Sachet in Allertonia 6: 210 (1991).

Hedyotis caerulea f. albiflora (Millsp.) Fosberg in Castanea 19: 30 (1954).

Hedyotis caerulea f. benkei Fosberg in Castanea 19: 31 (1954).

Hedyotis caerulea var. faxonorum (Pease & A. H. Moore) Fosberg in Castanea 19: 30
(1954).

Hedyotis caerulea var. minima (Beck) Fosberg in Castanea 19: 31 (1954).

Hedyotis canadensis (Roem. & Schult.) Fosberg in Virginia J. Sci. 2: 110 (1941).

33

Hedyotis centranthoides f. accrescens Fosberg in Bernice P. Bishop Mus. Bull. 174:
65 (1943).

Hedyotis centranthoides f. diffusa Fosberg in Bernice P. Bishop Mus. Bull. 174: 62
(1943).

Hedyotis centranthoides f. glomerata (Hook. & Arn.) Fosberg in Bernice P. Bishop
Mus. Bull. 174: 64 (1943).

Hedyotis centranthoides f. hirta Fosberg in Bernice P. Bishop Mus. Bull. 174: 63
(1943).

Hedyotis centranthoides f. kamokuensis Fosberg in Bernice P. Bishop Mus. Bull. 174:
63 (1943).

Hedyotis centranthoides f. kohala Fosberg in Bernice P. Bishop Mus. Bull. 174: 61
(1943).

Hedyotis centranthoides f. laevis (Wawra) Fosberg in Brittonia 8: 168. (1956).

Hedyotis centranthoides f. mauiensis Fosberg in Bernice P. Bishop Mus. Bull. 174: 65
(1943).

Hedyotis centranthoides f. meeboldii Fosberg in Bernice P. Bishop Mus. Bull. 174: 65
(1943).

Hedyotis centranthoides f. vestita Fosberg in Bernice P. Bishop Mus. Bull. 174: 62
(1943).

Hedyotis centranthoides f. yunckeri Fosberg in Bernice P. Bishop Mus. Bull. 174: 63
(1943).

Hedyotis centranthoides var. laevis (Wawra) Fosberg in Bernice P. Bishop Mus. Bull.
174: 63 (1943).

Hedyotis cookiana f. brighamii Fosberg in Bernice P. Bishop Mus. Bull. 174: 48
(1943).

Hedyotis cookiana f. elatior (Mann) Fosberg in Bernice P. Bishop Mus. Bull. 174: 47
(1943).

Hedyotis cookiana f. oahuensis Fosberg in Bernice P. Bishop Mus. Bull. 174: 48
(1943).

Hedyotis cookiana var. ensiformis Fosberg in Bernice P. Bishop Mus. Bull. 174: 46
(1943).

Hedyotis cookiana var. herbacea (H. Lév.) Fosberg in Bernice P. Bishop Mus. Bull.
174: 46 (1943).

Hedyotis corallicola Fosberg in Kew Bull. 33: 136 (1978).

Hedyotis coriacea f. conostyla (Gaudich.) Fosberg in Bernice P. Bishop Mus. Bull.
174: 25 (1943).

Hedyotis corymbosa var. ampla Fosberg in Allertonia 6: 220 (1991).

Hedyotis costata var. lutescens (Kaneh.) Fosberg in Smithsonian Contr. Bot. 45: 28
(1980).

Hedyotis cushingiae Fosberg in Allertonia 6: 221 (1991).

Hedyotis degeneri Fosberg in Bernice P. Bishop Mus. Bull. 174: 55 (1943).

Hedyotis degeneri var. coprosmifolia Fosberg in Bernice P. Bishop Mus. Bull. 174: 56
(1943).

Hedyotis divaricata var. capillothyrsa Fosberg in Allertonia 6: 224 (1991).

Hedyotis elatior (H. Mann) Fosberg in Brittonia 8: 167 (1956).

34

Hedyotis elatior f. brighamii Fosberg in Brittonia 8: 167 (1956).

Hedyotis elatior f. ensiformis Fosberg in Brittonia 8: 167 (1956).

Hedyotis elatior f. herbacea (H. Lév.) Fosberg in Brittonia 8: 167 (1956).

Hedyotis elatior f. oahuensis Fosberg in Brittonia 8: 167 (1956).

Hedyotis fluviatilis (C. N. Forbes) Fosberg in Bernice P. Bishop Mus. Bull. 174: 90
(1943).

Hedyotis fluviatilis f. breviflora Degener & Fosberg in Bernice P. Bishop Mus. Bull.
174: 91 (1943).

Hedyotis fluviatilis f. hathewayi Fosberg in Brittonia 8: 171 (1956).

Hedyotis fluviatilis f. kamapuaana (Degener) Fosberg in Bernice P. Bishop Mus. Bull.
174: 91 (1943).

Hedyotis fluviatilis var. kamapuaana (Degener) Fosberg in Bernice P. Bishop Mus.
Bull. 174: 91 (1943).

Hedyotis fluviatilis var. kauaiensis Fosberg in Bernice P. Bishop Mus. Bull. 174: 93
(1943).

Hedyotis foetida var. mariannensis (Merr.) Fosberg in Phytologia 5: 291 (1955).

Hedyotis foggiana Fosberg in Bernice P. Bishop Mus. Bull. 174: 65 (1943).

Hedyotis foliosa (Hillebr.) Fosberg in Bernice P. Bishop Mus. Bull. 174: 44 (1943).

Hedyotis formosa (Hillebr.) Fosberg in Bernice P. Bishop Mus. Bull. 174: 26 (1943).

Hedyotis fruticulosa var. atroglomerata Fosberg in Allertonia 6: 228 (1991).

Hedyotis fruticulosa var. yapensis (Kaneh.) Fosberg in Allertonia 6: 228 (1991).

Hedyotis glaucifolia (A. Gray) Fosberg in Bernice P. Bishop Mus. Bull. 174: 41
(1943).

Hedyotis glaucifolia f. nealiae Fosberg in Bernice P. Bishop Mus. Bull. 174: 43
(1943).

Hedyotis glaucifolia var. helleri Fosberg in Bernice P. Bishop Mus. Bull. 174: 44
(1943).

Hedyotis glaucifolia var. subimpressa Fosberg in Brittonia 8: 166 (1956).

Hedyotis glaucifolia var. waimeae (Wawra) Fosberg in Bernice P. Bishop Mus. Bull.
174: 43 (1943).

Hedyotis grantii Fosberg in Bernice P. Bishop Mus. Bull. 174: 29 (1943).

Hedyotis greenmanii Fosberg in Field & Lab. 17: 167 (1949).

Hedyotis hookeri (K. Schum.) Fosberg in Allertonia 6: 199 (1991).

Hedyotis intricata Fosberg in Lloydia 4: 290 (1941).

Hedyotis kanehirae (Hatus.) Fosberg in Smithsonian Contr. Bot. 45: 28 (1980).

Hedyotis knudsenii (Hillebr.) Fosberg in Bernice P. Bishop Mus. Bull. 174: 57 (1943).

Hedyotis korrorensis var. hosokawana Fosberg in Allertonia 6: 230 (1991).

Hedyotis korrorensis var. subglomerata Fosberg in Allertonia 6: 231 (1991).

Hedyotis laciniata var. takamatsui Fosberg in Occas. Pap. Bernice P. Bishop Mus.
15(20): 215 (1940).

Hedyotis lancifolia var. brevipes (Bremek.) Fosberg in Kew Bull. 33: 137 (1978).

Hedyotis littoralis (Hillebr.) Fosberg in Bernice P. Bishop Mus. Bull. 174: 53 (1943).

Hedyotis mannii Fosberg in Bernice P. Bishop Mus. Bull. 174: 49 (1943).

Hedyotis mannii var. cuspidata Fosberg in Brittonia 8: 168 (1956).

35

Hedyotis mannii var. laxiflora (Mann) Fosberg in Bernice P. Bishop Mus. Bull. 174:
50 (1943).

Hedyotis mannii var. munroi Fosberg in Bernice P. Bishop Mus. Bull. 174: 50 (1943).

Hedyotis mannii var. scoposa Fosberg in Bernice P. Bishop Mus. Bull. 174: 51 (1943).

Hedyotis michauxii f. alba (Alex.) Fosberg in Castanea 19: 32 (1954).

Hedyotis michauxii Fosberg in Amer. Midl. Naturalist 29: 786 (1943).

Hedyotis molokaiensis Fosberg in Bernice P. Bishop Mus. Bull. 174: 51 (1943).

Hedyotis mullerae Fosberg in Lloydia 4: 288 (1941).

Hedyotis nigricans (Lam.) Fosberg in Lloydia 4: 287 (1941).

Hedyotis nigricans var. pulvinata (Small) Fosberg in Castanea 19: 37 (1954).

Hedyotis nigricans var. scabra (Wats.) Fosberg in Lloydia 4: 288 (1941).

Hedyotis nuttalliana Fosberg in Virginia J. Sci. 2: 111 (1941).

Hedyotis parvula (A. Gray) Fosberg in Bernice P. Bishop Mus. Bull. 174: 54 (1943).

Hedyotis parvula f. sessilis Fosberg in Bernice P. Bishop Mus. Bull. 174: 55 (1943).

Hedyotis procumbens (J. F. Gmel.) Fosberg in Castanea 19: 32 (1954).

Hedyotis prolifera Fosberg in Kew Bull. 33: 138 (1978).

Hedyotis purpurea f. albiflora Fosberg in Castanea 19: 34 (1954).

Hedyotis purpurea f. leucantha (Standl.) Fosberg in Castanea 19: 36 (1954).

Hedyotis purpurea f. pubescens (Britton) Fosberg in Castanea 19: 33 (1954).

Hedyotis purpurea var. calycosa (A. Gray) Fosberg in Castanea 19: 33 (1954).

Hedyotis purpurea var. ciliolata (Torr.) Fosberg in Castanea 19: 34 (1954).

Hedyotis purpurea var. floridana (Standl.) Fosberg in Castanea 19: 36 (1954).

Hedyotis purpurea var. longifolia (Gaertn.) Fosberg in Castanea 19: 34 (1954).

Hedyotis purpurea var. setiscaphia (Carr) Fosberg in Castanea 19: 35 (1954).

Hedyotis purpurea var. tenuifolia (Nutt.) Fosberg in Castanea 19: 35 (1954).

Hedyotis raiateensis (J. W. Moore) Fosberg in Bernice P. Bishop Mus. Bull. 174: 28
(1943).

Hedyotis rapensis (F. Br.) Fosberg in Occas. Pap. Bernice P. Bishop Mus. 13(19): 250
(1937).

Hedyotis rapensis var. taverana Fosberg in Occas. Pap. Bernice P. Bishop Mus.
13@19)2 2508 C1937):

Hedyotis remyi (Hillebr.) Fosberg in Brittonia 8: 165 (1956).

Hedyotis remyi var. nuttallii (Fosberg) Fosberg in Brittonia 8: 166 (1956).

Hedyotis remyi var. plana (Fosberg) Fosberg in Brittonia 8: 166 (1956).

Hedyotis remyi var. silvicola Fosberg in Brittonia 8: 166 (1956).

Hedyotis rennellensis Fosberg in Lloydia 3: 122 (1940).

Hedyotis romanzoffiensis (Cham. & Schltdl.) Fosberg in Occas. Pap. Bernice P. Bishop
Mus. 13(19): 248 (1937).

Hedyotis sachetiana Fosberg in Allertonia 6: 235 (1991).

Hedyotis scabridifolia var. stonei Fosberg in Allertonia 6: 237 (1991).

Hedyotis schlechtendahliana f. dichotoma Fosberg in Bernice P. Bishop Mus. Bull.
174: 34 (1943).

Hedyotis schlechtendahliana f. glabrescens Fosberg in Bernice P. Bishop Mus. Bull.
174: 34 (1943).

36

Hedyotis schlechtendahliana f. glauca (Meyen) Fosberg in Bernice P. Bishop Mus.
Bull. 174: 36 (1943).

Hedyotis schlechtendahliana f. hirtella Fosberg in Bernice P. Bishop Mus. Bull. 174:
35 (1943).

Hedyotis schlechtendahliana f. kaalensis Fosberg in Bernice P. Bishop Mus. Bull. 174:
37 (1943).

Hedyotis schlechtendahliana f. secundiflora (Hillebr.) Fosberg in Bernice P. Bishop
Mus. Bull. 174: 34 (1943).

Hedyotis schlechtendahliana subsp. cordata (Cham. & Schltdl.) Fosberg in Bernice P.
Bishop Mus. Bull. 174: 32 (1943).

Hedyotis schlechtendahliana subsp. remyi (Hillebr.) Fosberg in Bernice P. Bishop
Mus. Bull. 174: 40 (1943).

Hedyotis schlechtendahliana subsp. rockii Fosberg in Bernice P. Bishop Mus. Bull.
174: 37 (1943).

Hedyotis schlechtendahliana subsp. tenuifolia (Fosberg) Fosberg in Bernice P. Bishop
Mus. Bull. 174: 39 (1943).

Hedyotis schlechtendahliana var. cordata (Cham. & Schltdl.) Fosberg in Bernice P.
Bishop Mus. Bull. 174: 32 (1943).

Hedyotis schlechtendahliana var. glauca (Meyen) Fosberg in Bernice P. Bishop Mus.
Bull. 174: 36 (1943).

Hedyotis schlechtendahliana var. nitens (Wawra) Fosberg in Bernice P. Bishop Mus.
Bull. 174: 37 (1943).

Hedyotis schlechtendahliana var. nuttallii Fosberg in Bernice P. Bishop Mus. Bull.
174: 41 (1943).

Hedyotis schlechtendahliana var. opaca (Wawra) Fosberg in Bernice P. Bishop Mus.
Bull. 174: 38 (1943).

Hedyotis schlechtendahliana var. plana Fosberg in Bernice P. Bishop Mus. Bull. 174:
41 (1943).

Hedyotis schlechtendahliana var. remyi (Hillebr.) Fosberg in Bernice P. Bishop Mus.
Bull. 174: 40 (1943).

Hedyotis schlechtendahliana var. reticulata Fosberg in Bernice P. Bishop Mus. Bull.
174: 40 (1943).

Hedyotis schlechtendahliana var. rigida Fosberg in Bernice P. Bishop Mus. Bull. 174:
38 (1943).

Hedyotis schlechtendahliana var. secundiflora (Hillebr.) Fosberg in Bernice P. Bishop
Mus. Bull. 174: 33 (1943).

Hedyotis schlechtendahliana var. tenuifolia Fosberg in Bernice P. Bishop Mus. Bull.
174: 39 (1943).

Hedyotis sieberi (Baker) Fosberg in Kew Bull. 33: 140 (1978).

Hedyotis strigulosa (DC.) Fosberg in Smithsonian Contr. Bot. 45: 28 (1980).

Hedyotis tayloriae Fosberg in Field & Lab. 17: 169 (1949).

Hedyotis thyrsoidea Fosberg in Bernice P. Bishop Mus. Bull. 174: 52 (1943).

Hedyotis thyrsoidea var. hillebrandii Fosberg in Bernice P. Bishop Mus. Bull. 174: 53
(1943).

Sy)

Hedyotis verticillata var. bartlettii Fosberg & Sachet in Allertonia 6: 211 (1991).

Hedyotis vestita var. lutescens (Kaneh.) Fosberg in Allertonia 6: 244 (1991).

Hedyotis wrightii (A. Gray) Fosberg in Lloydia 4: 290 (1941).

Hillia hathewayi Fosberg in Sida 2: 387 (1966).

Hoffmannia steinvorthii Fosberg in Sida 2: 388 (1966).

Hydnophytum stewartii Fosberg in Lloydia 3: 123 (1940).

Ixora sect. Vitixora Fosberg in Sargentia 1: 124 (1942).

Ixora biflora Fosberg in J. Bot. 76: 235 (1938).

Ixora biflora var. fleckeri Fosberg in J. Bot. 76: 277 (1938).

Ixora brevipedunculata Fosberg in Occas. Pap. Bernice P. Bishop Mus. 13(19): 271
@o377)):

Ixora carolinensis var. chartacea Fosberg in Occas. Pap. Bernice P. Bishop Mus.
15(20): 222 (1940).

Ixora carolinensis var. lanceolata (Kaneh.) Fosberg in Occas. Pap. Bernice P. Bishop
Mus. 15(20): 223 (1940).

Ixora carolinensis var. ponapensis Fosberg in Occas. Pap. Bernice P. Bishop Mus.
15(20): 223 (1940).

Ixora carolinensis var. volkensii (Hosok.) Fosberg in Occas. Pap. Bernice P. Bishop
Mus. 15(20): 222 (1940).

Ixora casei var. medialoba Fosberg & Sachet in Baileya 23: 85 (1989).

Ixora coccinea f. lutea (Hutch.) Fosberg & Sachet in Baileya 23: 79 (1989).

Ixora coccinea var. hermannii Fosberg & Sachet in Baileya 23: 79 (1989).

Ixora coccinea var. intermedia Fosberg & Sachet in Baileya 23: 80 (1989).

Ixora foetida (L.f.) Fosberg in Allertonia 7: 78 (1993).

Ixora moorensis (Nadeaud) Fosberg in Occas. Pap. Bernice P. Bishop Mus. 13(19):
269 (1937).

Ixora queenslandica Fosberg in J. Bot. 76: 234 (1938).

Ixora raivavaensis Fosberg in Occas. Pap. Bernice P. Bishop Mus. 13(19): 273 (1937).

Ixora setchellii Fosberg in Occas. Pap. Bernice P. Bishop Mus. 13(19): 266 (1937).

Ixora st.-johnii Fosberg in Occas. Pap. Bernice P. Bishop Mus. 13(19): 269 (1937).

Leptostigma longiflorum (Standl.) Fosberg in Acta Phytotax. Geobot. 33: 80 (1982).

Leptostigma pilosum (Benth.) Fosberg in Acta Phytotax. Geobot. 33: 79 (1982).

Leptostigma reptans (F. Muell.) Fosberg in Acta Phytotax. Geobot. 33: 82 (1982).

Leptostigma setulosum (Hook. f.) Fosberg in Acta Phytotax. Geobot. 33: 81 (1982).

Leptostigma weberbaueri Fosberg in Acta Phytotax. Geobot. 33: 80 (1982).

Morinda umbellata var. archboldiana Fosberg in Sargentia 1: 123 (1942).

Morinda umbellata var. bucidifolia (A. Gray) Fosberg in Sargentia 1: 123 (1942).

Morinda umbellata var. forsteri (Seem.) Fosberg in Sargentia 1: 123 (1942).

Morinda umbellata var. glandulosa (Merr.) Fosberg in Occas. Pap. Bernice P. Bishop
Mus. 15(20): 220 (1940).

Ophiorrhiza leptantha var. yasawana Fosberg in Bull. Torrey Bot. Club 67: 420
(1940).

Ophiorrhiza palauensis var. biseta Fosberg in Occas. Pap. Bernice P. Bishop Mus.
15(20): 214 (1940).

38

Psychotria \sect. Pelagomapouria Fosberg in Brittonia 16: 257 (1964).

Psychotria \sect. Straussia (A. Gray) Fosberg in Brittonia 16: 264 (1964).

Psychotria adamsonii Fosberg in Notul. Syst. (Paris) 8: 172 (1939).

Psychotria andersonii Fosberg in Allertonia 6: 248 (1991).

Psychotria archboldiana Fosberg in Sargentia 1: 131 (1942).

Psychotria atricaulis Fosberg in Candollea 38: 456 (1983).

Psychotria aurantiocarpa Fosberg in Sargentia 1: 130 (1942).

Psychotria bowermaniae Fosberg in Notul. Syst. (Paris) 8: 170 (1939).

Psychotria brevicalyx Fosberg in Sargentia 1: 132 (1942).

Psychotria carolinensis (Valeton) Fosberg in Occas. Pap. Bernice P. Bishop Mus.
15(20): 224 (1940).

Psychotria cheathamiana Fosberg in Allertonia 6: 249 (1991).

Psychotria chrysophylla Fosberg in Bull. Torrey Bot. Club 67: 423 (1940).

Psychotria cordata var. podantha Fosberg in Sargentia 1: 130 (1942).

Psychotria crassiflora Fosberg in Sargentia 1: 132 (1942).

Psychotria degeneri Fosberg in Sargentia 1: 134 (1942).

Psychotria eumorphanthus Fosberg in Sargentia 1: 127 (1942).

Psychotria fauriei (H. Lév.) Fosberg in Occas. Pap. Bernice P. Bishop Mus. 23(2): 43
(1962).

Psychotria fragrans (Gillespie) Fosberg in Sargentia 1: 126 (1942).

Psychotria furcans Fosberg in Sargentia 1: 133 (1942).

Psychotria gibbsiae var. velutina Fosberg in Sargentia 1: 127 (1942).

Psychotria glabra (Turrill) Fosberg in Sargentia 1: 126 (1942).

Psychotria grantii Fosberg in Occas. Pap. Bernice P. Bishop Mus. 13(19): 277 (1937).

Psychotria greenwelliae Fosberg in Brittonia 16: 269 (1964).

Psychotria griseifolia var. unicarinata Fosberg in Sargentia 1: 129 (1942).

Psychotria hathewayi Fosberg in Brittonia 16: 267 (1964).

Psychotria hathewayi var. brevipetiolata Fosberg in Brittonia 16: 267 (1964).

Psychotria hawaiiensis (A. Gray) Fosberg in Occas. Pap. Bernice P. Bishop Mus.
23(2): 43 (1962).

Psychotria hawaiiensis var. glabrithyrsa Fosberg in Brittonia 16: 271 (1964).

Psychotria hawaiiensis var. glomerata (Rock) Fosberg in Brittonia 16: 271 (1964).

Psychotria hawaiiensis var. hillebrandii (Rock) Fosberg in Occas. Pap. Bernice P.
Bishop Mus. 23(2): 43 (1962).

Psychotria hawaiiensis var. molokaiensis (Rock) Fosberg in Brittonia 16: 271 (1964).

Psychotria hawaiiensis var. rotundifolia (Skottsb.) Fosberg in Brittonia 16: 271 (1964).

Psychotria hawaiiensis var. scoriacea (Rock) Fosberg in Brittonia 16: 270 (1964).

Psychotria hexandra f. forbesii Fosberg in Brittonia 16: 263 (1964).

Psychotria hexandra f. hosakana Fosberg in Brittonia 16: 262 (1964).

Psychotria hexandra f. waialuana Fosberg in Brittonia 16: 260 (1964).

Psychotria hexandra subsp. oahuensis Degener & Fosberg in Brittonia 16: 261 (1964).

Psychotria hexandra var. hosakana Fosberg in Brittonia 16: 262 (1964).

Psychotria hexandra var. kealiae Fosberg in Brittonia 16: 260 (1964).

Psychotria hexandra var. oahuensis (Degener & Fosberg) Degener & Fosberg in
Brittonia 16: 261 (1964).

39

Psychotria hexandra var. rockii Fosberg in Brittonia 16: 262 (1964).

Psychotria hexandra vat. st.-johnii Fosberg in Brittonia 16: 262 (1964).

Psychotria hivaoana Fosberg in Notul. Syst. (Paris) 8: 166 (1939).

Psychotria hombroniana (Baill.) Fosberg in Phytologia 5: 291 (1955).

Psychotria hombroniana var. canfieldiae Fosberg in Allertonia 6: 253 (1991).

Psychotria hombroniana var. hirtella (Valeton) Fosberg in Smithsonian Contr. Bot. 45:
29 (1980).

Psychotria hombroniana var. kusaiensis (Kaneh.) Fosberg in Smithsonian Contr. Bot.
45: 29 (1980).

Psychotria hombroniana var. ladronica (Hosok.) Fosberg in Allertonia 6: 255 (1991).

Psychotria hombroniana var. malaspinae (Merr.) Fosberg in Smithsonian Contr. Bot.
45: 29 (1980).

Psychotria hombroniana var. mariannensis (Kaneh.) Fosberg in Allertonia 6: 255
(1991).

Psychotria hombroniana var. peliliuensis Fosberg in Allertonia 6: 255 (1991).

Psychotria hombroniana var. squarrosa (Valeton) Fosberg in Smithsonian Contr. Bot.
45: 29 (1980).

Psychotria hosokawae Fosberg in Allertonia 6: 257 (1991).

Psychotria insularum var. paradisii Fosberg in Brittonia 16: 256 (1964).

Psychotria kaduana (Cham. & Schltdl.) Fosberg in Occas. Pap. Bernice P. Bishop
Mus. 23(2): 43 (1962).

Psychotria kaduana var. longissima (Rock) Fosberg in Brittonia 16: 266 (1964).

Psychotria kaduana var. pubiflora (Heller) Fosberg in Brittonia 16: 265 (1964).

Psychotria le-bronnecii Fosberg in Notul. Syst. (Paris) 8: 168 (1939).

Psychotria le-bronnecii var. tahuatensis Fosberg in Notul. Syst. (Paris) 8: 169 (1939).

Psychotria leptocarpa (Hillebr.) Fosberg in Brittonia 16: 266 (1964).

Psychotria leptothyrsa var. leptothyrsoides (Kaneh.) Fosberg in Allertonia 6: 259
(1991).

Psychotria leptothyrsa var. yapensis Fosberg in Allertonia 6: 260 (1991).

Psychotria macroserpens Fosberg in Sargentia 1: 131 (1942).

Psychotria magnifica (Gillespie) Fosberg in Sargentia 1: 126 (1942).

Psychotria marauensis Fosberg & Florence in Candollea 38: 456 (1983).

Psychotria mariniana (Cham. & Schltdl.) Fosberg in Occas. Pap. Bernice P. Bishop
Mus. 23(2): 43 (1962).

Psychotria mauiensis Fosberg in Brittonia 16: 266 (1964).

Psychotria merrillii var. polyneura (Valeton) Fosberg & Sachet in Allertonia 6: 264
(i991):

Psychotria monocarpa Fosberg in Sargentia 1: 130 (1942).

Psychotria pickeringii var. solanoides (Turrill) Fosberg in Sargentia 1: 128 (1942).

Psychotria pittosporifolia Fosberg in Sargentia 1: 134 (1942).

Psychotria ponapensis Fosberg in Occas. Pap. Bernice P. Bishop Mus. 15(20): 224
(1940).

Psychotria psychotrioides (Heller) Fosberg in Brittonia 16: 269 (1964)>

Psychotria pubiflora (A. Gray) Fosberg in Sargentia 1: 126 (1942).

40

Psychotria raivavaensis Fosberg in Occas. Pap. Bernice P. Bishop Mus. 13(19): 279
(1937).

Psychotria rotensis var. palauensis (Hosok.) Fosberg in Allertonia 6: 267 (1991).

Psychotria rufocalyx Fosberg in Sargentia 1: 127 (1942).

Psychotria serpens var. parvula (A. Gray) Fosberg in Sargentia 1: 131 (1942).

Psychotria st.-johnii Fosberg in Bull. Torrey Bot. Club 67: 423 (1940).

Psychotria tetragonoides Fosberg in Sargentia 1: 135 (1942).

Psychotria timonioides Fosberg in Sargentia 1: 136 (1942).

Psychotria tubuaiensis Fosberg in Occas. Pap. Bernice P. Bishop Mus. 13(19): 277
(1937):

Psychotria vitiensis Fosberg in Sargentia 1: 127 (1942).

Psychotria waianensis Fosberg in Brittonia 16: 268 (1964).

Psychotria whistleri Fosberg in Micronesica 23: 3 (1940).

Randia vitiensis (Seem.) Fosberg in Sargentia 1: 120 (1942).

Readea prismoclavata Fosberg in Sargentia 1: 137 (1942).

Readea roseata Fosberg in Sargentia 1: 136 (1942).

Spermacoce buckleyi Fosberg in Brittonia 40: 64 (1988).

Spermacoce dispersa (Hook. f.) Fosberg in Phytologia 62: 182 (1987).

Spermacoce ericifolia (Hook. f.) Fosberg in Phytologia 62: 182 (1987).

Spermacoce ernstii Fosberg & D. Powell in Smithsonian Contr. Bot. 45: 29 (1980).

Spermacoce everistiana Fosberg in Brittonia 40: 61 (1988).

Spermacoce linearifolia (Hook. f.) Fosberg in Phytologia 62: 182 (1987).

Spermacoce perpusilla (Hook. f.) Fosberg in Phytologia 62: 182 (1987).

Spermacoce repens (DC.) Fosberg & D. Powell in Smithsonian Contr. Bot. 45: 30
(1980).

Spermacoce rotundifolia (Anderss.) Fosberg in Phytologia 62: 183 (1987).

Spermacoce suberecta (Hook. f.) Fosberg in Phytologia 62: 183 (1987).

Tarenna sambucina var. buruensis (Miq.) Fosberg & Sachet in Allertonia 6: 272
(1991).

Tarenna sambucina var. glabra (Merr.) Fosberg & Sachet in Allertonia 6: 272 (1991).

Tarenna sambucina var. oweniana Fosberg in Allertonia 6: 272 (1991).

Tarenna sambucina var. papuana Fosberg in Allertonia 6: 273 (1991).

Tarenna sambucina var. tahitensis (Valeton) Fosberg & Sachet in Allertonia 6: 273
(1991).

Tarenna verdcourtiana Fosberg in Phytologia 41: 357 (1979).

Timonius affinis var. sapotifolius (A. Gray) Fosberg in Sargentia 1: 121 (1942).

Timonius corymbosus var. takamatsui Fosberg & Sachet in Micronesica 20: 159 (1988
"1987").

Timonius kajewskii (Guillaumin) Fosberg in Bull. Torrey Bot. Club 70: 393 (1943).

Timonius mollis var. submollis Fosberg & Sachet in Micronesica 20: 161 (1988
W987"):

Timonius mollis var. villosissimus (Kaneh.) Fosberg & Sachet in Micronesica 20: 161
(1988 "1987").

Timonius salsedoi Fosberg & Sachet in Micronesica 20: 162 (1988 "1987").

Timonius smithii Fosberg in Sargentia 1: 121 (1942).

41

Timonius subauritus var. strigosus Fosberg & Sachet in Micronesica 20: 163 (1988
"1987").

Trukia dryadum (S. Moore) Fosberg in Phytologia 62: 175 (1987).

Trukia fitzalanii (F. Muell.) Fosberg in Phytologia 62: 175 (1987).

Trukia macarthurii (F. Muell.) Fosberg in Phytologia 62: 176 (1987).

Trukia tahitensis (Nadeaud) Fosberg in Phytologia 62: 174 (1987).

Rutaceae
Zanthoxylum semiarticulatum f. laiense (Degener) Fosberg in Occas. Pap. Bernice P.
Bishop Mus. 24(2): 20 (1969).
Zanthoxylum semiarticulatum var. sessile (Degener) Fosberg in Occas. Pap. Bernice P.
Bishop Mus. 24(2): 19 (1969).

Santalaceae
Santalum ellipticum var. latifolium (A. Gray) Fosberg in Occas. Pap. Bernice P.
Bishop Mus. 23(2): 35 (1962).
Santalum insulare var. alticola Fosberg & Sachet in Candollea 40: 463 (1985).
Santalum insulare var. deckeri Fosberg & Sachet in Candollea 40: 467 (1985).
Santalum insulare var. hendersonense (F. Br.) Fosberg & Sachet in Candollea 40: 470
(1985).
Santalum insulare var. raiateense (J. W. Moore) Fosberg & Sachet in Candollea 40:
465 (1985).

Sapindaceae
Allophylus paniger Fosberg in Allertonia 7: 61 (1993).

Sapotaceae
Manilkara achras (Mill.) Fosberg in Taxon 13: 255 (1964).

Pouteria auahiensis (J. F. Rock) Fosberg in Occas. Pap. Bernice P. Bishop Mus. 23(2):
40 (1962).

Pouteria aurantia (J. F. Rock) Fosberg in Occas. Pap. Bernice P. Bishop Mus. 23(2): 40
(1962).

Pouteria calcarea (Hosok.) Fosberg in Smithsonian Contr. Bot. 45: 17 (1980).

Pouteria ceresolii (J. F. Rock) Fosberg in Occas. Pap. Bernice P. Bishop Mus. 23(2): 39
(1962).

Pouteria grayana (St. John) Fosberg in Micronesica 25: 196 (1992).

Pouteria grayana var. florencei Fosberg in Micronesica 25: 196 (1992).

Pouteria micronesica (Kaneh.) Fosberg in Smithsonian Contr. Bot. 45: 17 (1980).

Pouteria rhynchosperma (J. F. Rock) Fosberg in Occas. Pap. Bernice P. Bishop Mus.
23(2): 39 (1962).

Pouteria spathulata (Hillebr.) Fosberg in Occas. Pap. Bernice P. Bishop Mus. 23(2): 40
(1962).

42

Saxifragaceae

Broussaisia arguta f. glabra Fosberg in Occas. Pap. Bernice P. Bishop Mus. 15(4): 58
(939):

Broussaisia arguta f. oppositifolia Fosberg in Occas. Pap. Bernice P. Bishop Mus.
15(4): 60 (1939).

Broussaisia arguta var. pellucida Fosberg in Occas. Pap. Bernice P. Bishop Mus.
15(4): 59 (1939).

Scrophulariaceae
Veronica virginica var. purpurea (Pursh) Fosberg in Castanea 20: 59 (1955).

Solanaceae
Solanum cheesmaniae (Riley) Fosberg in Phytologia 62: 181 (1987).
Solanum indicum var. aldabrense (C. H. Wright) Fosberg in Kew Bull. 33: 141
(1978).
Solanum lycopersicum var. cerasiforme (Dunal) Fosberg in Phytologia 5: 290 (1955).

Sterculiaceae
Melochia villosissima var. compacta (Hochr.) Fosberg in Micronesica 23: 149 (1990).
Sterculia ellipticifolia Fosberg in Willdenowia 20: 263 (1991).

Thymelaeaceae
Phaleria ixorioides Fosberg in Bull. Torrey Bot. Club 67: 418 (1940).

Tiliaceae
Corchorus torresianus var. yunckeri Fosberg in Micronesica 2: 147 (1966).
Tilia americana var. neglecta (Spach) Fosberg in Castanea 20: 58 (1955).

Triuridaceae
Sciaphila stemmermanniae Fosberg & Sachet in Pac. Sci. 34: 25 (1981).

Typhaceae
Typha domingensis var. sachetiae Fosberg in Bot. Bull. Acad. Sin. 30: 220 (1989).

Urticaceae
Elatostema divaricatum (Gaudich.) Fosberg in Smithsonian Contr. Bot. 45: 6 (1980).

Verbenaceae
Callicarpa candicans f. glabriuscula (H. J. Lam) Fosberg in Smithsonian Contr. Bot.
45: 25 (1980).
Callicarpa candicans var. integrifolia (H. J. Lam) Fosberg in Smithsonian Contr. Bot.
45: 25 (1980).
Callicarpa candicans var. paucinervia (Merr.) Fosberg in Smithsonian Contr. Bot. 45:
25 (1980).

Callicarpa candicans var. ponapensis Fosberg in Smithsonian Contr. Bot. 45: 26
(1980).

Clerodendrum glabrum var. minutiflorum (Baker) Fosberg in Kew Bull. 33: 143
(1978).

Lantana involucrata f. candida Fosberg in Rhodora 78: 113 (1976).

Lippia strigulosa f. parvifolia (Moldenke) Fosberg in Rhodora 78: 113 (1976).

43

i 5» iDemeday ¢

ATOLL RESEARCH BULLETIN

NO. 391

ANNOTATED LIST OF THE KNOWN PUBLICATIONS OF
F. RAYMOND FOSBERG

BY

ANNA L. WEITZMAN

ISSUED BY
NATIONAL MUSEUM OF NATURAL HISTORY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C., U.S.A.
FEBRUARY 1994

AAG HORA

i 14 jj-+ i

(MOTRIN J) 2rTaAn 2O Uae JAMOITAW
= : VOT A1Ter VADIORTIMA « :
‘ ABU Ml OTDM

bel YRAURMBY © &

ANNOTATED LIST OF THE KNOWN PUBLICATIONS OF F. RAYMOND
FOSBERG

BY

ANNA L. WEITZMAN

INTRODUCTION

F, Raymond Fosberg kept a card file and reprints of scientific publications, reviews, popular articles,
and less formally distributed writings. None of those sources is complete and I have filled in some gaps by
searching through periodicals that he used frequently. There will undoubtedly be corrections and additions to
this list. Scientific publications are numbered 1-625, with Dr. Fosberg's own numbering in parentheses
following the sequential numbers I assigned (his numbering had many publications numbered out of order or
unnumbered). I did not number republications of articles. Because this is an attempt to give a relatively
complete summation of the published views of Dr. Fosberg, I have included reviews, published letters,
published abstracts, anonymous committee reports that he attributed to himself, and informally published (often
duplicated and distributed by Dr. Fosberg at meetings) documents in this list. These publications were not
carefully documented and I am sure that this list is far from complete. Those categories are numbered
separately as R1-89 (reviews), L1-20 (letters), and D1-86 (abstracts and informal documents) respectively.

In many instances, especially later in his career, large and small publications included information
about named taxa. Many of those, such as the regional checklists, do not need further explanation, but some
include other information about a number of taxa. I have included a list of genera discussed following the
citation of the article. Those names are also indexed by citation and page numbers following the publication
list. In nearly all cases I have omitted the names of taxa when the information given is merely distributional. I
have made a judgment about what genera are included, necessarily subjective, but where there is information
that may help the user understand Dr. Fosberg's views on the taxon or taxa, or help the user identify and apply
the correct name to plants, the generic name is listed and indexed. I have sometimes included synonyms in the
list because they are lectotypified, or because I thought it useful to include them.

I have also included a list of publications that are known (by the Botany Department at the National
Museum of Natural History) to be in press or submitted for publication. I expect an additional paper of Pacific
plant distribution notes based on manuscripts found in his office. There are other publications which are
expected to be completed and published by co-authors including: The Vegetation of the Pacific Islands with
Dieter Mueller-Dombois to be published by Fischer Verlag, a list of vascular plants of Kosrae with Jerome
Ward, and various articles written with David Stoddart. Other publications which Dr. Fosberg was working on
are unlikely to appear. Those include a checklist of vascular plants of the Society Islands; a supplement to the
Micronesian checklists; a Marquesas checklist; Flora of Micronesia, part 6; the bulk of the planned Forster list;
a Spermacoce (Rubiaceae) monograph; and an A/yxia (Apocynaceae) monograph. The treatments of
Malvaceae and Rubiaceae for the Flora of Ceylon have been forwarded to others for completion. The Pacific
plant checklists will be used in preparation of a planned checklist for all of Oceania. Readers who would like to
use materials that Dr. Fosberg did not publish should contact the Botany Department of the National Museum
of Natural History.

Periodical titles are abbreviated following B-P-H (G.H.M. Lawrence et al.’s 1968 Botanico-
Periodicum-Huntianum) and its Supplement (Bridson and Smith, 1991). Unless marked with *, all citations

Department of Botany, MRC-166, National Museum of Natural History, Smithsonian Institution, Washington,
D.C. 20560.

were checked against original publications or reprints with original pagination. Where actual date of
publication or distribution differs from that on the title page. the date on the title page is placed in brackets
following the actual date of distribution according to Dr. Fosberg's records or other sources. A few citations
verified by Lu Eldredge (Pacific Science Association) are indicated with **, one verified by Scott Miller
(Bishop Museum) is indicated with ***. The assistance of Dan Nicolson throughout my work on this list is
gratefully acknowledged.

A brief list of some published photographs and biographical information of F.R. Fosberg follows the
list of publications. This is not intended to be a complete list, merely a compilation of what I found in
compiling this list of publications.

KNOWN PUBLICATIONS OF F.R. FOSBERG

1929
1 (1). Fosberg, F.R. 1929. Preliminary notes on the fauna of the giant kelp, Macrocystis pyrifera.
Pomona Coll. J. Ent. and Zool. 21: [3 unnumbered pages in reprint. ]
1931
2 (2). Fosberg, F.R. 1931. Remarks on the taxonomy of the Cactaceae and some new combinations and
names in that family. Bull. S. Calif. Acad. Sci. 30: 50-59. [in reprint pp. 1-10; Coryphantha,
Neomammillaria, Opuntia. |
. 1931. The cacti of the Pyramid Peak Region, Dona Ana County, New Mexico. Bull. S. Calif.
Acad. Sci. 30: 67-73. [*Republished in Desert Mag. 5: 147-149, 1942?]

316):

1932

4 (4). Fosberg, F.R. 1932. The study of the Cactaceae. Cact. Succ. J. (Los Angeles) 4: 270-272.

[Opuntia.]
1933

5 (5). Pierce, W.M. and F.R. Fosberg. 1933. Notes on southwestern cacti. Bull. S. Calif. Acad. Sci. 32:

121-126. [Opuntia, Echinocereus, Cereus, Echinocactus, Neomammillaria.}
1934

6 (6). Fosberg, F.R. 1934. A key to the families of Monocotyledons in the Hawaiian Islands. Occas. Pap.
Univ. Hawaii 18: 1-8.

7 (7). ——. 1934. The southern California Prickly-Pears. Bull. S. Calif. Acad. Sci. 33: 93-104. [(110a)
republished in Desert Plant Life 21: 75-80, 1949.] [Opuntia.|

1935

8 (8). Fosberg, F.R. and J. Ewan. 1935. Notes on southwestern plants. Bull. S. Calif. Acad. Sci. 34: 177-

183.
1936

9 (9). Fosberg, F.R. 1936. Plant collecting on Lanai, 1935. Mid-Pacific Mag. 49: 119-123.

10 (10). ——. 1936. Miscellaneous Hawaiian plant notes—I. Occas. Pap. Bernice P. Bishop Mus. 12(15):
3-11. [Cryptocarya, Cassytha, Schiedea, Eurya, Phyllanthus, Antidesma, Diospyros, Phyllostegia,
Rollandia.|

11 (11). ——. 1936. The Hawaiian geraniums. Occas. Pap. Bernice P. Bishop Mus. 12(16): 1-19.

12 (12). ——. 1936. The varieties of the desert willow, Chilopsis linearis. Madrono 3: 362-366.

13 (14). ——. 1936. Plant remains in Shelter Cave, New Mexico. Bull. S. Calif. Acad. Sci. 35: 154-155.

*D1 (17a). ——. 1936. Vegetation of Vostok Island, central Pacific, [abstract.] Special Publ. Bernice
Pauahi Bishop Mus. (Proc. Hawaiian Acad. Sci. 11) 30(24): 19.

*D2 (17b). ——. 1936. A study of the Hawaiian genus, Gouldia, [abstract.] Special Publ. Bernice Pauahi
Bishop Mus. (Proc. Hawaiian Acad. Sci. 11) 30(24): 20.

*D3 (17c). St. John, H. and F.R. Fosberg. 1936. Vegetation of Flint Island, Central Pacific, [abstract.]

Special Publ. Bernice Pauahi Bishop Mus. (Proc. Hawaiian Acad. Sci. 11) 30(24): 20.
1937

14 (15). Fosberg, F.R. 1937. Immigrant plants in the Hawaiian Islands. I. Occas. Pap. Univ. Hawaii 32:
3-11. [Myrica, Atriplex, Spergularia, Geranium, Sida, Myriophyllum, Elephantopus, Eupatorium,
Pluchea.|

15 (18). . 1937. The genus Gouldia (Rubiaceae). Bernice P. Bishop Mus. Bull. 147: 1-82, 3 plates.

16 (19). ——. 1937. Some Rubiaceae of southeastern Polynesia. Occas. Pap. Bernice P. Bishop Mus.
13(19): 245-293. [Hedyotis, Ophiorrhiza, Tarenna, Canthium, Guettarda, Timonius, Ixora.
Psychotria, Nertera, Coprosma, Morinda, Borreria.|

*17 (16). Degener, O. and F.R. Fosberg. 1937. Luzula campestris (L.) DC., in Degener, O., Flora
Hawaiiensis, 2 pp.

18 (13). St. John, H. and F.R. Fosberg. 1937. Vegetation of Flint Island, Central Pacific. Occas. Pap.
Bernice P. Bishop Mus. 12(24): 1-4.

1938
19 (20). Fosberg, F.R. 1938. The Lower Sonoran in Utah. Science 87: 39-40.
20 (22). ——. 1938. Eriogonum abertianum and its varieties. Madrofio 4: 189-194.
21 (24a). ——. 1938. Two Queensland /xoras. J. Bot. 76: 233-237.
22 (24b). ——. 1938. Additional Note on Queensland /xoras. J. Bot. 76: 276-277.
23 (25). ——. 1938. A central repository for type-specimens. J. Bot. 76: 327-330.
24 (26). . 1938. Notes on plants of the Pacific Islands. |. Bull. Torrey Bot. Club 65: 607-614.

[Gouania, Diospyros, Randia.]

25 (21). Fosberg, F.R. and E.Y. Hosaka. 1938. An open bog on Oahu. Occas. Pap. Bernice P. Bishop
Mus. 14(1): 1-6. [Styphelia, Lobelia.|

26 (23). St. John, H. and F.R. Fosberg. 1938. Identification of Hawaiian plants: a key to the families of
dicotyledons of the Hawaiian Islands, descriptions of the families, and list of the genera. Occas. Pap.
Univ. Hawaii 36: 1-53.

D4 (19a). Fosberg, F.R. 1938 [1937.] An aggressive Lantana mutation. Special Publ. Bernice Pauahi
Bishop Mus. (Proc. Hawaiian Acad. Sci. 12) 31: 18.

1939

27 (27). Fosberg, F.R. 1939. Safeguarding type specimens. Science 89: 245.

28 (28). ——. 1939. Plant collecting manual for field anthropologists. Botanical Supply Dept., American
Fiber-Velope Mfg. Co., 1-22, Philadelphia.

29 (29). ——. 1939. Nomenclature proposals for the 1940 Botanical Congress. Amer. J. Bot. 26: 229-
231.

30 (30). . 1939. Notes on Polynesian grasses. Occas. Pap. Bernice P. Bishop Mus. 15(3): 37-48.
[Eragrostis, Digitaria, Paspalum.

31 (31). . 1939. Taxonomy of the Hawaiian genus Broussaisia (Saxifragaceae). Occas. Pap. Bernice
P. Bishop Mus. 15(4): 49-60.

32 (32). ——. 1939. Diospyros ferrea (Ebenaceae) in Hawaii. Occas. Pap. Bernice P. Bishop Mus.
15(10): 119-131.

33 (33). ——. 1939. Taxonomy and hybridism. Chron. Bot. 5: 397-398.

34 (38). ——. 1939. Psychotria (Rubiaceae) in the Marquesas Islands. Notul. Syst. (Paris) 8: 161-173.

35 (104). ——. 1939. Reports on nature protection and related items for: Fanning Island, Christmas

Island, Vostok Island, Flint Island, Henderson Island, Oeno Island. Pitcairn Island, Society Islands,
Austral Islands, Rapa, Tuamotu Archipelago, Gambia Islands. Proc. Sixth Pacific. Sci. Congress 4:
501-504, 517-524.

36 (33b). St. John, H. and F.R. Fosberg. 1939. A new variety of Ruppia maritima (Ruppiaceae) from the
tropical Pacific. Occas. Pap. Bernice P. Bishop Mus. 15(16): 175-178.

R1 (33a). Fosberg, F.R: 1939. Artistic tropical blooms, review of McLean, O.G. and M.D. Frear, Flowers
of Hawaii, 1938. J. New York Bot. Gard. 40: 73.

1940

37 (34). Fosberg, F.R. 1940. Notes on Micronesian Rubiaceae. Occas. Pap. Bernice P. Bishop Mus.
15(20): 213-226. [Bikkia, Uncaria, Ophiorrhiza, Hedyotis, Mussaenda, Randia, Timonius, Canthium,
Gynochthodes, Morinda, Ixora, Psychotria.|

38 (37). . 1940. Notes on plants of the Pacific Islands—II. Bull. Torrey Bot. Club 67: 417-425.
[Diospyros, Phaleria, Neonauclea, Hedyotis, Ophiorrhiza, Gynochthodes, Timonius, Morinda, Ixora,
Psychotria, Digitaria.|

39 (39). . 1940. Melanesian Vascular Plants. Lloydia 3: 109-124. [Campium, Lycopodium, Ficus,
Acalypha, Medinilla, Dischidia, Hoya, Acanthus, Pseuderanthemum, Hedyotis, Hydnophytum.]|

40 (40). . 1940. The aestival flora of the Mesilla Valley Region. New Mexico. Amer. Midl.
Naturalist 23: 573-593.

41 (43). ——. 1940. Financing a taxonomic journal. Chron. Bot. 6: 55-56.

42 (44). ——. 1940. A taxonomic journal. Chron. Bot. 6: 127-128.

43 (54a). ——. 1940. Santalaceae of Nevada. Contributions toward a Flora of Nevada (U.S.D.A.) 5: 1-2.
[Comandra.]}

44 (54b). ——. 1940. Ulmaceae of Nevada. Contributions toward a Flora of Nevada (U.S.D.A.) 6: 1-3.
[Celtis.]

45 (54c). ——. 1940. Chenopodiaceae of Nevada (in part). Contributions toward a Flora of Nevada
(U.S.D.A.) 8: 1-18. [Nitrophila, Allenrolfea, Salsola, Sarcobatus, Bassia, Halogeton, Grayia,
Eurotia, Cycloloma, Monolepis.|

46 (35). St. John, H. and F.R. Fosberg. 1940. Identification of Hawaiian plants: Part 2. A key to the
families and genera of the gymnosperms and of the monocotyledons of the Hawaiian Islands, with
descriptions of the families. Occas. Pap. Univ. Hawaii 41: 1-47.

R2 (33b). Fosberg. F.R. 1940. By airplane to a Stone-Age Island. review of Archbold, R., and A. L. Rand,
New Guinea Expedition, 1940. J. New York Bot. Gard. 41: 19.

3 (41). ——. 1940. Review of Gates, F.C.. Flora of Kansas. Amer. Midl. Naturalist 24: 503-504.

R4 (42). ——. 1940. Review of Deam, C.C., Flora of Indiana, 1940. Chron. Bot. 6: 95.

1941

47. Fosberg, F.R. 1941. Names in Amaranthus, Artocarpus, and Inocarpus. J. Wash. Acad. Sci. 31: 93-
96.

48 (54h). ——. 1941. Loganiaceae of Nevada. Contributions toward a Flora of Nevada (U.S.D.A.) 23: 1-
2. [Buddleja.|

49 (54d). ——. 1941. Aizoaceae of Nevada. Contributions toward a Flora of Nevada (U.S.D.A.) 24: 1-3.
[Sesuvium, Mollugo.]}

50 (54e). . 1941. Haloragaceae (Haloragidaceae) of Nevada. Contributions toward a Flora of Nevada
(U.S.D.A.) 25: 1-3. [Hippuris, Myriophyllum.]

51 (54f). . 1941. Elaeagnaceae of Nevada. Contributions toward a Flora of Nevada (U.S.D.A.) 26: 1-
4. [Elaeagnus.|

52 (46). . 1941. For an open-minded taxonomy. Chron. Bot. 6: 368-370.

53 (47). . 1941, Observations on Virginia Plants, Part I. Virginia J. Sci. 2: 106-111. [Abies,
Anemone, Saxifraga, Spiraea, Veronica, Hedyotis.|

54 (85). . 1941. Rubiaceae, pp. 94-95, in Hermann, F.J., Check-list of plants in the Washington—
Baltimore area. Conference on the District Flora, Washington.

55 (48). . 1941. Varieties of the strawberry guava. Proc. Biol. Soc. Wash. 54: 179-180. [Psidium.]

56 (50). . 1941. Notes on Mexican plants. Lloydia 4: 274-290. [Phoradendron, Mirabilis, Drymaria,
Paronychia, Fendlera, Rhamnus, Hedyotis.|

SWS): . 1941. Notes on North American plants—1. Amer. Midl. Naturalist 26: 690-695.
[Chenopodium, Atriplex, Acer.]

58 (52). . 1941. Observations on Virginia plants, Part II. Virginia. J. Sci. 2: 284-288.

59 (49). Fosberg, F.R. and E.H. Walker. 1941. A preliminary check list of plants in the Shenandoah
National Park, Virginia. Castanea 6: 89-136.

60 (54g). McVaugh, R. and F.R. Fosberg. 1941. Index to the geographical names of Nevada.
Contributions toward a Flora of Nevada (U.S.D.A.) 29: 1-216.

D5 (55). Fosberg, F.R. 1941. Care and housing of botanical type specimens. 7 pp.. duplicated and
distributed with Taxonomic Index.

D6 (56). ——. 1941. Local floras in relation to conservation. 4 pp., duplicated and distributed with
Taxonomic Index.

1942

61 (57). Fosberg, F.R. 1942. Notes on North American plants—II. Amer. Midl. Naturalist 27: 253-258.
[Arriplex, Arenaria, Lewisia, Ceanothus, Echinocactus, Elaeagnus.|

62 (58). . 1942. Uses of Hawaiian Ferns. Amer. Fern J. 32: 15-23.

63 (59). ——. 1942. Subspecies and variety. Rhodora 44: 153-157.

64 (60). . 1942. Coniferae (Coniferales) of the Washington-Baltimore area—preliminary keys. 5 pp.,
Conference on District Flora. [Pinus, Tsuga, Taxodium, Chamaecyparis, Juniperus. |

65 (61). ——. 1942. Rubiaceae, in Smith, A.C., Fijian Plant Studies II. Sargentia 1: 118-140.

66 (62). ——. 1942. The genus Batocarpus Karst (Moraceae). Proc. Biol. Soc. Wash. 55: 99-102.

67 (63). . 1942. Notes on North American plants—III. Amer. Midl. Naturalist 27: 761-765.
[Pimenta, Oenothera, Gilia, Richardia, Sambucus.]|

68 (64). ——. 1942. Scholarship and economic usefulness. Chron. Bot. 7: 154-155.

69 (65). ——. 1942. Cornus sericea L. (C. stolonifera Michx.) Bull. Torrey Bot. Club 69: 583-589.

70 (66). ——. 1942. Segregation of type specimens. Science 96: 515-516.

71 (66c). ——. 1942. Miscellaneous notes on Hawaiian plants—2. Occas. Pap. Bernice P. Bishop Mus.

16(15): 337-347. [Dryopteris, Korthalsella, Geranium, Vaccinium, Lepechinia, Phyllostegia,
Coprosma. |

R5 (66a). ——. 1942. Reviews of Bailey, L.H. and E.Z. Bailey, Hortus Second, 1941 and Kelsey, H.P. and
W.A. Dayton, Standardized Plant Names, ed. 2, 1942. Chron. Bot. 7: 283-285.

D7 (66b). [list by editors of Chron. Bot., assistance from others including F.R. Fosberg.] 1942. A list of
plant scientists in Central and South America. Chron. Bot. 7: 97-133.

1943

72 (67). Fosberg, F.R. 1943. The Polynesian species of Hedvotis (Rubiaceae). Bernice P. Bishop Mus.
Bull. 174: 1-102.

73 (68). . 1943. Notes on North American plants. IV. Amer. Midl. Naturalist 29: 785-786.
[Portulaca, Hedyotis.|

74 (69). . 1943. Notes on plants of the Pacific Islands—III. Bull. Torrey Bot. Club 70: 386-397.
[Nephrolepis, Anectochilus, Spergularia, Malachra, Maurandva, Ruellia, Gouldia, Timonius,
Coprosma, Elephantopus, Pseudelephantopus, Ageratum, Dubautia.|

75 (70). . 1943. Colombian Cinchona Manual. Foreign Economic Administration, Bogota,
Colombia. 27 pp.

76 (72). ——. 1943. Proposal to conserve Schizaea. Chron. Bot. 7: 395.

77 (71). Fosberg, F.R. and E.H. Walker. 1943. First supplement to a preliminary check list of plants in the
Shenandoah National Park. Castanea 8: 109-115.

1944
78 (74). Fosberg, F.R. 1944. Colombian Cinchona Manual, 2nd edition. Foreign Economic
Administration, Bogota, Colombia, 33 pp.

IDG3)s . 1944. Manual de Quinas Colombianos. [Spanish translation by G. Saenz Fety.] Foreign
Economic Administration, Bogota, Colombia, 40 pp. [Cinchona.]
80 (75). ——. 1944. El Paramo de Sumapaz, Colombia. J. New York Bot. Gard. 45: 226-234.
*D8 (73a). ——. 1944? Gifts of the Americas. Quinine. | p., place and date of publication unknown.
1945

81 (76). Fosberg, F.R. 1945. Principal economic plants of tropical America, pp. 18-35, in F. Verdoorn,
ed., Plants and Plant Science in Latin America. Chronica Botanica Co. Waltham, Mass.

82 (77). ——. 1945. The Air Force's collection of aerial photographs. Science 102: 538-539.

83 (79). ——. 1945. The work of botanists of the Cinchona Mission in Colombia. Chron. Bot. 9: 119-
vile

84 (86). ——. 1945. Drymaria and Mollugophytum, pp. 95-96, in Morton, C.V., Mexican Phanerogams
described by M-E. Jones. Contr. U.S. Natl. Herb. 29: 87-116.

85 (87). ——. 1945. The Sleepy Hollow area [collecting projects in the local region.] Bull. Wash.-Balt.
Flora 9: 4-5.

1946

86 (80). Fosberg, F.R. 1946. A biologist reflects upon old age and death. Sci. Monthly 63: 158-159.

87 (81). ———. 1946. Observations on Virginia Plants. III. Castanea 11: 66-69. [Triodia.|

88 (82). ——. 1946. Application of the name Euphorbia maculata L. Rhodora 48: 197-200.

89 (83). ——. 1946. The herbarium. Sci. Monthly 63: 429-434.

90 (84). . 1946. Rubiaceae, pp. 106-108, in Hermann, F.J., A checklist of plants in the Washington-

Baltimore Area. Ed. 2., Conference on the District Flora, Washington.

91 (88a). . 1946. Gifts of the Americas: Quinine. Agriculture of the Americas, inside back cover.
[Cinchona.|
R6 (78). ——. 1946. Vegetation of the Far East described for men in the Service, review of Merrill, E.D.,
Plant Life of the Pacific World, 1945. J. New York Bot. Garden 47: 44-45.
L1 (88). ——. 1946. Present status of foreign Herbaria and Museums, [letter to the editor.] Science 103:
282-283.
1947

92 (89). Fosberg. F.R. 1947. Observations on Virginia Plants. IV. Castanea 12: 59-62. [/mpatiens,
Chelone, Taraxacum.|

93 (90). . 1947. Formaldehyde in plant collecting. Science 106: 250.

94 (91). ——. 1947. Cinchona plantation in the New World. Econ. Bot. 1: 330-333.

95 (92). ——. 1947. Euphorbia maculata again. Bull. Torrey Bot. Club 74: 332-333.

96 (93). ——. 1947. Micronesian mangroves: a journey by motorized sampan to a peak development of
land-building swamps in the Carolines. J. New York Bot. Gard. 48: 128-138.

97 (101). ——. 1947. Nyctaginac-7e. Contributions toward a Flora of Idaho 36: 1-3. [Mirabilis,
Abronia.|

98 (102). _ 1947. Cornaceae of Idaho. Contributions toward a Flora of Idaho 77: 1-3. [Cornus.]

R7 (94). ——. 1947. General biology: philosophy and education, review of Notiy, L. du, Human Destiny,

1947. Quart. Rev. Biol. 22: 315-317.
1948

99 (95). Fosberg, F.R. 1948. Island Floras, pp. 18-21, in Coolidge, H.J., ed., Conservation in Micronesia.
National Research Council, Washington, D.C.

100 (96). ——. 1948. Areas to be set aside, pp. 16-19, im Recommendations and selected papers from the
Honolulu Conference on Conservation in Micronesia.

101 (96a). ———. 1948. Island Floras, pp. 16-19, i7 Recommendations and selected papers from the
Honolulu Conference on Conservation in Micronesia.

102 (97). ———. 1948 [1946.] Preservation of natural areas for scientific reserves and watersheds.
Botanical Report on Micronesia, U.S. Commercial Company, Economic Survey of Micronesia Vol. 13-
I, pp. 17-19.

103 (98). ———. 1948. Artificial key to the Rubiaceae of the Washington-Baltimore Area. Bull. Wash.-
Balt. Flora 12: 10-12.

104 (100). ———. 1948. Derivation of the flora of the Hawaiian Islands, pp. 107-118, in Zimmerman, E.C.,
Insects of Hawaii. Vol. 1. Univ. Press Hawaii, Honolulu.

105 (103). ——. 1948. Economic Botany—a modern concept of its scope. Econ. Bot. 2: 3-14.

106 (105). ——. 1948. Salinity and Atol! vegetation. Proc. Hawaiian Acad. Sci. 23: 8.

107 (106). . 1948. Immigrant plants in the Hawaiian Islands. II. Occas. Pap. Univ. Hawaii 46: 1-17.
[Musa, Spergularia, Kalanchoé, Bruguiera, Polyscias, Kigelia, Ixora, Hippobroma, Emilia,
Eupatorium.]}

108 (99). Fosberg, F.R. and E.H. Walker. 1948. Second supplement to a preliminary check list of plants
in the Shenandoah National Park. Castanea 13: 83-92.

D9 (109). Fosberg, F.R. 1948. Popular or profusely illustrated works on Hawaiian plants. 2 pp.,
duplicated and distributed by Workshop Committee, Summer Program Committee, Nature Study
Classes, Honolulu.

1949

109 (107). Fosberg, F.R. 1949. Atoll vegetation and salinity. Pacific Sci. 3: 89-92.

110 (111). ——. 1949. A new Drymaria (Caryophyllaceae) from Mexico. Proc. Biol. Soc. Wash. 62:
147-148.

111 (113). ——. 1949. Flora of Johnston Island, central Pacific. Pacific Sci. 3: 338-339.

112 (114). ——. 1949. Early history of Cinchona. J. New York Bot. Gard. 50: 254-257.

113 (114a). . 1949. Hedyotis angulata Fosberg and Hedyotis greenmanii Fosberg, in Shinners. L.H.,
Transfer of Texas species of Houstonia to Hedyotis (Rubiaceae). Field & Lab. 17: 166-169.

114 (117). ——. 1949. Waipoua. New Zealand Sci. Rev. 7: 200-201.

115 (126). ——. 1949. Plant exploration—South America and the Pacific Islands. Bull. Hort. Soc. New
York 20(4): 9-11.

R8 (110). . 1949. The Rules reprinted, review of Camp, W.H., H.W. Rickett, and C.A. Weatherby,
International Rules of Botanical Nomenclature, 1947. J. New York Bot. Gard. 50: 92-93.

D10 (108). . 1949. Further nomenclatural considerations. 12 pp., distributed privately before 7th Int.
Bot. Congress.

D11 (112). ——. 1949. The problem of rare and vanishing plant species. Duplicated for and distributed

at IUPN Conference, UNESCO/IUPN/Conf. 2/IIf/8, Lake Success, 2 pp. [(199) republished 1950?.
Proc. & Papers Internat. Tech. Conf. Protection Nature 502-504. ]

*D12 (125). ——. 1949. A conservation program for Micronesia, [abstract.] Selected Abstracts and Final
Resolutions of the Seventh Pacific Science Congress, Auckland, p. 12.

1950
116 (116). Fosberg, F.R. 1950. Official plant names? Rhodora 52: 1-7.
117 (118). ——. 1950. Notes on Micronesian Pteriodophytes. Amer. Fern J. 40: 132-147. [Angiopteris,
Marattia, Hymenophyllum, Trichomanes, Gleichenia, Lygodium, Schizaea, Lycopodium.|
118 (120). . 1950. Translations of Japanese botanical papers. Pacific Sci. 4: 375.
119 (121). ——. 1950. Ecological notes on the upper Amazon. Ecology 31: 650-653.
120 (122). ——. 1950. Remarks on the factual basis of ecology. Ecology 31: 653-654.
121 (123). . 1950. Nature appreciation in the Netherlands. Nature 43: 482-484.

122 (118a). Diehl, W.W. and F.R. Fosberg. 1950. Taxus in Piedmont Virginia. Castanea 15: 129-131.

R9 (115). Fosberg, F.R. 1950. A much needed text for botany students, review of Newbigin, M.I., Plant
and Animal Geography, 1949. J. New York Bot. Gard. 51: 16.

**D 13 (124). [——.] 1950. Report on the Seventh International Botanical Congress. Pacific Sci. Assoc.
Inform. Bull. 50(6): 3.

1951
123 (127). Fosberg, F.R. 1951. The American element in the Hawaiian Flora. Pacific Sci. 5: 204-206.
124 (128). ——. 1951. Museums in Pacific research. Proc. Seventh Pacific Sci. Congress 1: 91-93.
125 (131). ——. 1951. Regulations for the selection of generic names for conservation. II. Suggestions

by F.R. Fosberg. Taxon 1: 15.

*126 (132). ——. 1951. Temprana historia de la quina. Revista Acad. Colomb. Ci. Exact. 8: 273-274.
127 (133). ——. 1951. Atoll Research Bulletin—Editorial Introduction. Atoll Res. Bull. 1: i-ii.
128 (134). ——. 1951. Ecological research on coral atolls. Atoll Res. Bull. 1: 6-8. [(714). republished

1992. Atoll Res. Bull. 355: 9-11.]
129 (135). ——. 1951. Flora and vegetation of coral atolls. Atoll Res. Bull. 1: 13-14.

130 (136). ——. 1951. Literature on coral atolls. Atoll Res. Bull. 1: 23-25.

131 (137). . 1951. Land ecology on coral atolls. Atoll Res. Bull. 2: 7-11.

132 (129). Tweedie, M.W.F. and F.R. Fosberg. 1951. The research museum or herbarium. Proc. Seventh
Pacific Sci. Congress 1: 97.

R10 (137a). ——. 1951. Plants of a coral atoll, review of Taylor, W.R., Plants of Bikini, 1950. Sci.
Monthly 72: 196-197.

R11 (137b). ———. 1951. Before men came, review of Braun, E.L., Deciduous Forests of Eastern North
America, 1950. Sci. Monthly 73: 66-67.

R12 (137c). ——. 1951. Regional ecology, review of Pearsall, W.H.. Mountains and Moorlands, 1950.
Sci. Monthly 73: 135.

R13 (137d). ——. 1951. Earth's largest ocean, review of Freeman, O.W., ed., Geography of the Pacific,
1951. Sci. Monthly, 73: 203-204.

**D 14 (130). [——.] 1951. Interim report on the activities of the Standing Committee on Pacific

Conservation. Pacific Sci. Assoc. Inform. Bull. 51(8): 3-4.
1952

133 (140). Fosberg. F.R. 1952. Introduction and chapter 6: Botany, pp. 1, 43-51, 55-56, in Sachet, M.-H.,
ed., Handbook for Atoll Research, preliminary edition.

*134 (150). . 1952. Lignes biogéographiques dans l'ouest du Pacifique. Compt. Rend. Sommaire
Séances Soc. Biogéogr. ?(256): 161-166. [No volume number on reprint. ]

135 (139). Fosberg. F.R. and H. St. John. 1952. Vegetation et flore de I'atoll Maria, iles Australes. Rev.
Sci. Bourbonnais Centr. France. 1951: 1-7 [translated into French by M.-H. Sachet.]

136 (138). Degener. O. and F.R. Fosberg. 1952. A central Pacific Sesuvium. Occas. Pap. Bernice P.
Bishop Mus. 21(14): 45-47.

R14 (140a). Fosberg, F.R. 1952. Keys to local woody plants, review of Blackburn, B., Trees and shrubs in
Eastern North America, 1952. Castanea 17: 142-144.

R15 (151). ——. 1952. Review of Porsild, A-E., Botany of Southeastern Yukon adjacent to the Canol
Road, 1951. Quart. Rev. Biol. 27: 394.

D15 (140b). ——. 1952. Plants suitable for Naval station, Kwajalein, and suggestions for their
cultivation. [Prepared for Commanding Officer, U.S. Naval Station, Kwajalein. ]

1953

137 (141). Fosberg, F.R. 1953. Nomenclature of Premna obtusifolia R. Br. Taxon 2: 88-89.

138 (142). ——. 1953. Proposal to conserve the generic name Citrul/us Schrader. Taxon 2: 99-101.

139 (143). ——. 1953. Proposals to amend the Stockholm Code of Botanical Nomenclature. Taxon 2:
107-109.

*140 (145). ——. 1953. La flora de los atolones de Oceania. Rev. Geogr. Chile 9: 30-42.

141 (147). ——. 1953. Typification of Euphorbia maculata L. Rhodora 55: 241-242.

142 (149). ——. 1953. Vegetation for a southern California arboretum. Année Biol. 29: 429-442.

143 (155). . 1953. A conservation programme for Micronesia. Proc. Seventh Pacific Sci. Congress 4:
670-673.

144 (152a). ——. 1953. Directions for studying vegetation and flora, pp. 44-56, in Fosberg, F.R. and M.-
H. Sachet, eds. Handbook for Atoll Research (second preliminary edition). Atoll Res. Bull. 17: 1-129.

145 (157a). ——. 1953. Chapter 6—Botany, pp. 44-56, in Fosberg, F.R. and M.-H. Sachet, eds., Handbook
for Atoll Research, second preliminary edition.

146 (157). ——. 1953. Vegetation of central Pacific atolls, a brief summary. Atoll Res. Bull. 23: 1-26.

147 (148). Fosberg, F.R. and L. Artz. 1953. The varieties of Mfonarda fistulosa L. Castanea 18: 128-130.

148 (144). Fosberg. F.R. and M.-H. Sachet. 1953. Reinstatement of Eranthemum pulchellum Andr.
Taxon 2: 135-136.

149 (15la). Fosberg, F.R. and M.-H. Sachet, eds. 1953. Handbook for Atoll Research (second preliminary
edition). Atoll Res. Bull. 17: 1-129.

R16 (151). Fosberg. F.R. 1953. Review of Lawrence, G.H.M., Taxonomy of Vascular Plants, 1951.
Quarterly Rev. Biol. 28: 58-61.

1954

150 (146). Fosberg, F.R. 1954. Comments on recent nomenclature proposals. Taxon 3: 1-3.

151 (152). . 1954. Notes on plants of the Eastern United States. Castanea 19: 25-37. [Equisetum,
Acer, Oenothera, Hedyotis.|

152 (154). . 1954. Soils of the northern Marshall atolls, with special reference to the Jemo Series.
Soil Sci. 78: 99-107.

*153 (156). . 1954. Vanishing island floras and vegetation. Proces-verbaux et rapports de la reunion
technique, Caracas, 1952, IUPN 538-543.

154 (156a). . 1954. The synopsis of proposals for the 8th Congress. Taxon 3: 150-151.

155 (153). Dandy, J.E. and F.R. Fosberg. 1954. The type of 4maryllis bella-donna L. Taxon 3: 231-232.

R17 (156a). Fosberg, F.R. 1954. Review of Good, R., The Geography of the Flowering Plants, ed. 2,
1953. Science 120: 132.

**D16 (210). [——.] 1954. Note on Eighth International Botanical Congress. Pacific Sci. Assoc. Inform.
Bull. 6(3): 6.

**D17 (211). [——.] 1954. Conservation in the U.S. Trust Territory of the Pacific. Pacific Sci. Assoc.
Inform. Bull. 6(3): 6-7.

**D18 (209). [——.] 1954. Eighth International Botanical Congress. Pacific Sci. Assoc. Inform. Bull.

6(6): 10.
1955

156 (158). Fosberg, F.R. 1955. Ants and dermestids on a mountain top. Ecology 36: 367-368.

157 (160). ——. 1955. Northern Marshall Islands Expedition, 1951-1952. Narrative. Atoll Res. Bull.
38: 1-36.

158 (161). ——. 1955. Northern Marshall Islands Expedition, 1951-1952. Land biota: vascular plants.
Atoll Res. Bull. 39: 1-22.

159 (164). ——.. 1955. Observations on Virginia plants V. Castanea 20: 58-60. [Tridens, Tilia,
Monotropa, Veronica, Campanula. |

160 (168). . 1955. Observations on Hedyotis caerulea var. minor. Castanea 20: 104-106.

161 (173). ——. 1955. Systematic notes on Micronesian Plants. Phytologia 5: 289-292. [Digitaria, Ficus,
Melanolepis, Maytenus, Polyscias, Solanum, Pseuderanthemum, Canthium, Hedvotis, Psvchotria,
Wedelia.|

162 (172). . 1955. Pacific forms of Lepturus R. Br. (Gramineae). Occas. Pap. Bernice P. Bishop
Mus. 21(14): 285-294.

*163 (159). Fosberg, F.R., P. Dansereau, and W.R.B. Oliver. 1955. Botanical report on Ambuklao.
[perhaps published in The Ambuklao Symposium by IUPN]

164 (165). Fosberg, F.R. and E.H. Walker. 1955. Third supplement to a preliminary check list of plants in
the Shenandoah National Park. Castanea 20: 61-70.

*165 (163). Sachet, M.-H. and F.R. Fosberg. 1955. Island Bibliographies: Micronesian Botany, land
environment and ecology of coral atolls, vegetation of tropical Pacific Islands. National Academy of
Sciences—National Research Council, publication #335, 577 pp.

R18 (162). Fosberg, F.R. 1955. Review of Gressitt. J. L., Insects of Micronesia. vol. 1, Introduction,
1954. Science, 121: 560-561.

*R19 (259). ——. 1955. Review of Odum, E.P. Fundamentals of Ecology. Austral. J. Sci. 18: 37-38.

R20 (260). ——. 1955. Review of Gates Clarke, J.F., Catalogue of the type specimens of Microlepidoptera
in the British Museum (Natural History) described by Edward Meyrick, vols. 1 & 2. 1955. Science
122: 1274-1275.

1956
166 (174). Fosberg, F.R. 1956. Studies in Pacific Rubiaceae: I-IV. Brittonia 8: 165-178. [Hedvotis,

Gouldia, Coprosma.]|
167 (175). ——. 1956. Disposition of Urera gaudichaudiana Henslow. Ann. Mag. Nat. Hist., ser. 12. 9:

423-424.

*168 (179). . 1956. Le Role des réserves dans la conservation d'espéces et d'associations végétales,
pp. 28-33, in Heim et al. Derniers refuges, Atlas commenté des Réserves Naturelles dans le monde.

169 (169a). . 1956. Citrullus, Citrullus vulgaris Schrader still correct. Nomina conservanda
proposita. Taxon 5: 15.

170 (169). Fosberg, F.R. and M.-H. Sachet. 1956. The Indo-Pacific strand Scaevola. Taxon 5: 7-10.

171 (170). Moore, H.E., Jr. and F.R. Fosberg. 1956. The palms of Micronesia and the Bonin Islands.
Gentes Herb. 8: 422-478. [Livistona, Metroxylon, Heterospathe, Areca, Pinanga, Gulubia,
Clinostigma, Ptychosperma, Cocos, Nypa.]

*D19 (166). Fosberg, F.R. 1956. Coral atoll ecology. Working paper for UNESCO Humid Tropics
Committee. Preparatory Meeting, Kandy, Ceylon, UNESCO/NS/HT/47, Paris, 5 pp.

*D20 (167). ——. 1956. The ecology of Man in the tropics. UNESCO Humid Tropics Committee
Background Paper. UNESCO/NS/HT/44, Paris, 5 pp.
L2 (179a). ——. 1956. On Axis deer, [letter to the editor.] Honolulu Advertiser, 3 March.
1957

172 (176). Fosberg, F.R. 1957. The community ecologist. A.I.B.S. Bull. 7: 24-25. [(*176a.) republished
1957. Audubon Magazine 59: 197-198; (*265.) republished 1960. Bio-dynamics 56: 24-28.]

173 (177). ——. 1957. Some geological processes at work on coral atolls. Trans. New York Acad. Sci.,
ser. 2, 19: 411-421.

174 (181). ——. 1957. Slicks on ocean surface downwind from coral reefs. Atoll Res. Bull. 53: 1-4.

175 (182). ——. 1957. The Maldive Islands, Indian Ocean. Atoll Res. Bull. 58: 1-37.

176 (183). . 1957. Description and occurrence of atoll phosphate rock in Micronesia. Amer. J. Sci.
255: 584-592.

177 (184). ——. 1957. Lonely Pokak. Living Wilderness 22(62): 1-4.

178 (185). ——. 1957. The naturalized flora of Micronesia and World War II. Proc. Eighth Pacific Sci.
Congress 4: 229-234.

179 (186). ——. 1957. The vegetation provinces of the Pacific. Proc. Eighth Pacific Sci. Congress 4: 15-
2B)

180 (187). ——. 1957. Vegetation of the Oceanic province of the Pacific. Proc. Eighth Pacific Sci.
Congress 4: 48-55.

181 (207). ——. 1957. Notes on the vegetation of Iriomote-Zima, Ryukyu Islands. Sci. Bull. Agric. and
Home Economics Div., Univ. Ryukyus, 4: 123-130.

**182 (222). ——. 1957. Soils, vegetation and agriculture on coral atolls. Proc. Eighth Pacific Sci.
Congress 3A: 1037-1047.

183 (204). ——. 1957 [1956.] The protection of nature in the islands of the Pacific. Comptes Rendus des

séances, Huitieme Congress International de Botanique, Paris, 1954, Sect. 21-27: 104-116.

184 (180). Fosberg, F.R. and M.-H. Sachet. 1957. Plantes récoltées en Micronésie au XIX® Siécle. Bull.
Mus. Hist. Nat. (Paris), 2nd sér., 29: 428-438.

*185 (189). Jenkins, A.E. and F.R. Fosberg. 1957. Records of citrus canker and sweet potato stem and
foliage scab in Micronesia. Pl. Dis. Reporter 41(12): 1055-1056.

R21 (197a). Fosberg, F.R. 1957. Review of Zaneveld, J.S., Economic marine algae of tropical south and
east Asia and their utilization, 1955. Quart. Rev. Biol. 32: 396-397.

L3 (197b). . 1957. Botanists are essential for success, [letter to the editor.] Honolulu Star-Bulletin,
[copy undated, after 1 November. ]

L4 (197c). ———. 1957. Advice on a botanical garden, [letter to the editor.| Honolulu Star-Bulletin, 9
November.

D21 (178). ——. undated but distributed 1957. Preliminary list of botanists interested in Pacific Basin

problems. 41 pp., duplicated and distributed by Pacific Science Association Standing Committee on

Pacific Botany.
D22 (188). ——. 1957. Report on the implementation of the botanical resolutions passed at the various

Pacific Science Congresses. Proc. Eighth Pacific Sci. Congress 4: 2-8.

11

D23 (195). ——. 1957. Dynamics of atoll vegetation, [abstract.] Ninth Pacific Sci. Congress Abstracts,
pp. 42-43.

D24 (192). ——. 1957. Tropical Pacific grasslands and savannas, [abstract.] Ninth Pacific Sci. Congress
Abstracts, p. 43.

D25 (194). ——. 1957. Qualitative description of the coral atoll ecosystem, [abstract.] Ninth Pacific Sci.
Congress Abstracts, p. 56.

D26 (193). ——. 1957. The herbarium as a research museum, [abstract.] Ninth Pacific Sci. Congress
Abstracts, pp. 149-150.

D27 (196). ——. 1957. Introduction [to Symposium on climate, vegetation, and rational land utilization
in the humid tropics; abstract.] Ninth Pacific Sci. Congress Abstracts, p. 253.

D28 (197). ——. 1957. The preservation of man's environment, [abstract.] Ninth Pacific Sci. Congress

Abstracts, pp. 264-265.
D29 (205). . 1957. List of botanists interested in Pacific Basin problems. Duplicated and distributed
by Pacific Science Association Standing Committee on Pacific Botany, pp. 1-64.

1958
186 (198). Fosberg, F.R. 1958. Notes on Micronesian Pteridophyta, I]. Amer. Fern J. 48: 35-39.
[Ceratopteris.]

187 (200). . 1958. Vascular flora of Pagan Island, Northern Marianas. Pacific Sci. 12: 17-20.

188 (201). ——. 1958. Mapping of vegetation types. Study of Tropical Vegetation, UNESCO, 219-220.

189 (202). ——. 1958. Vegetation of the islands of Oceania. Study of Tropical Vegetation, UNESCO 54-
60.

190 (203). . 1958 [1956.] Habitat availability and the development of floras. Comptes Rendus des

séances, Huitieéme Congress International de Botanique, Paris, 1954, Sects. 3-6: 96-104.

191 (206). . 1958. Zanthoxylum L., "Xanthoxylon Mill.", and Thylax Raf. Taxon 7: 94-96.

192 (212). ——. 1958 [1957.] List of plant species reported from Pagan, p. 107, in [Corwin, G., L.D.
Bonham, M.H. Terman, and G.W. Viele,] U.S. Army Engineers, U.S. Geological Survey, Military
Geology of Pagan Mariana Islands, Intelligence Division, Office of Engineer, Headquarters, U. S. Air
Force (Far East), Tokyo. [Table is four pages long. ]

193 (213). . 1958. Nomenclature proposals for the Montreal Congress. Taxon 7: 149-151.

*194 (216). ——. 1958. Algunos campos de investigacion basica significativos en la evaluacion de los
recursos naturales, [translated by T. Arango B.] Econ. Colomb. 19(54): 43-52.

195 (217). ——. 1958. Una experiencia Colombiana, [translated by T. Arango B.]. Revista Nac. Agric.
641: 15-21.

196 (218). ——. 1958. A new Trichosanthes (Cucurbitaceae) from Palau. Occas. Pap. Bernice P. Bishop

Mus. 22(1): 67-69.

197 (220). . 1958. Vascular plants from Rennell and Bellona Islands, pp. 101-102, in Wolff, T., The
Natural History of Rennell Island, British Solomon Islands, Danske Rennelleskpedition, 1951, vol. 3.

198 (199). Fosberg, F.R. and G. Corwin. 1958. A fossil flora from Pagan, Mariana Islands. Pacific Sci.
12: 3-16. [Pteris, Freycinetia, Pandanus, Miscanthus, Cocos, Alocasia, Dianella, Spathoglottis,
Trema, Hernandia, Macaranga, Melanolepis, Aglaia, Tristiropsis, Elaeocarpus, Eurya, Premna.|

L5 (215). Fosberg, F.R. 1958 [1957.] Forestry at the Ninth Pacific Science Congress, [letter to the editor.]
Malayan Forester 21: 201-202.

D30 (214). ——. 1958. Manas a dispersal agent. 1-5, 1958. Duplicated for and distributed at Missouri
Bot. Gard. Annual Systematics Symposium.

D31 (219). ——. 1958. Natural areas as a botanical resource. 4 pp. Duplicated by Nature Conservancy
for distribution at American Society of Plant Taxonomists annual meeting.

**T)32 (208). [——.] 1958. [List of standing committee on Pacific Botany.] Pacific Sci. Assoc. Inform.
Bull. 10(6): 2-3.

1959
199 (221). Fosberg, F.R. 1959. Upper limits of vegetation on Mauna Loa, Hawaii. Ecology 40: 144-146.
200 (223). ——. 1959. Pinus contorta and its variations. Baileya 7: 7-10.

201. ——. 1959. [Proposal] 308 [amend] Article 24 [Montreal Congress.] Taxon 8:19.

202. ——. 1959. [Proposal] 312 [amend] Article 73 [Montreal Congress.] Taxon 8: 19.

203 (224). ——. 1959. Nomenclatural notes on Datura L. Taxon 8: 52-57.

204 (225). ——. 1959. Serianthes dilmyi nomen novum. Taxon 8: 65.

205 (226). ——. 1959. Typification of Zanthoxylum L. Taxon 8: 103-105.

206 (227). ——. 1959. Long-term effects of radioactive fallout on plants? Atoll Res. Bull. 61: 1-11.

207 (228). ——. 1959. Vegetation and flora of Wake Island. Atoll Res. Bull. 67: 1-20.

208 (229). ——. 1959. Additional records of phanerogams from the northern Marshall Islands. Atoll
Res. Bull. 68: 1-9.

209 (232). ——. 1959. Plants and fall-out. Nature 183: 1448.

210 (234). ——. 1959 [1958.] Manas a dispersal agent. Southw. Naturalist 3: 1-6.

211 (236). ——. 1959 [1958.] Climate, vegetation, and rational land utilization in the humid tropics.
Introduction. Proc. Ninth Pacific Sci. Congress 20: 1-3.

212 (237). ——. 1959. The preservation of man's environment. Proc. Ninth Pacific Sci. Congress 20:
159-161.

*213 (241). ——. 1959. The research frontier. Saturday Review 42(46): 72-73.

*214 (242). ——. 1959. Who will identify the plants? Pl. Sci. Bull. 5(5): 4.

215 (244). ——. 1959 [1958.] The herbarium as a research museum. Proc. Ninth Pacific Sci. Congress
14: 13-14.

216 (247). ——. 1959. Conservation situation in Oceania. Proc. Ninth Pacific Sci. Congress 7: 30-31.

217 (249). ——. 1959. Notes on the Shenandoah National Park flora. Castanea 24: 135-143.

[Thelypteris, Setaria, Monarda.|

218 (250). . 1959. Vegetation [of Guam], pp. 167-217, in Tracey, J.I. Jr.. C.H. Stensland, D.B. Doan,
H.G. May, S.O. Schlanger, and J.T. Stark, Military geology of Guam, Mariana Islands, Parts I and II.
Intelligence Div., Office of the Engineer, U.S. Army Pacific.

219 (252). ——. 1959. The importance of biological research in the Pacific region. Gard. Bull. Singapore
17: 219-224.
220 (252c). . 1959. Flora and Fauna of Peru, Flora of Melanesia, Flora of Colombia. [Articles in the

1959 edition of Collier's Encyclopedia, cited from page proof. |

221 (238). Fosberg, F.R. and J.R. Swallen. 1959. Agnes Chase. Taxon 8: 145-151.

222 (246). [Fosberg. F.R.] 1959 [1958.] The vegetation of Micronesia. Dept. of the Army, Office, Chief of
Engineers, Engineer Intelligence Study # 257, xviii + 160 pp. [also published as: Fosberg, F.R.
1959. Scientific Investigations in Micronesia 1949-; The vegetation of Micronesia—Part 1. Pacific
Science Board, National Research Council, National Academy of Sciences, xx + 158 pp.]

223 (230). [Fosberg, F.R., ed.]. 1959. Atoll news and comment. Atoll Res. Bull. 70: 1-7.

224 (171). [Fosberg, F.R.. T. Arnow, and F.S. MacNeil.] 1959 [1956.] U.S. Army Engineers, U.S.
Geological Survey. Military geography of the Northern Marshalls. Intelligence Division, Office of
Engineer, Headquarters, U. S. Air Force (Far East), Tokyo, ix + 320 pp.

R22 (245). Fosberg. F.R. 1959. Tropical ecology, review of Misra, R. and G.S. Puri, Indian manual of
plant ecology, 1958. Ecology 40: 753-754.

R23 (252a). ——. 1959. Review of Koedoe, [Journal for Scientific Research in the National Parks of the
Union of South Africa] vol. 1, 1958. National Parks Mag. 33: 14-15.
R24 (252b). ——. 1959. Review of Gates Clarke, J.F., Catalogue of the Type Specimens of

Microlepidoptera in the British Museum (Natural History) described by Edward Meyrick, vol. 3, 1958.
Science 130: 1246.

L6 (231). ——. 1959. Balance in cultivated ecosystems, [letter to the editor.] Science 129: 1499.

*D33 (222a). . 1959. Maps of the humid tropics: Report of the Bloomington Conference. Working
paper for Abidjan Meeting. UNESCO/NS/HT/82. 5 pp.

D34 (233). ——. 1959. Summation by the Rapporteur Général of the Symposium of the vegetation of the
humid tropics. Berita Madjelis Ilmu Pengetahuan Indonesia. (Council for Sciences of Indonesia) 1:
16-21. [(*243.) republished 1959. Humid Tropics Newslett. 1: 2-7.]

D35 (235). . 1959 [1958.] Report on the symposium on climate, vegetation, and rational land
utilization in the humid tropics. Proc. Ninth Pacific Sci. Congress 20: v-vi.
D36 (239). ——. 1959. Structural-functional classification of vegetation for small-scale mapping,
[abstract.] Proc. Ninth International Bot. Congress 2: 118.
D37 (240). ——. 1959. Vegetation and the geologist, [abstract.] Proc. Ninth International Bot. Congress
2: 118-119.
1960

225 (248). Fosberg, F.R. 1960. The vegetation of Micronesia. 1. General descriptions, the vegetation of
the Marianas Islands, and a detailed consideration of the vegetation of Guam. Bull. Amer. Mus. Nat.
Hist. 119: 1-76 + 40 plates.

226 (251). . 1960. Check-list of the seed plants of Guam. i-ii, 1-51. Military Geology Branch, U.S.
Geological Survey. Duplicated by U.S. Navy, Agana, Guam).

227 (253). ——. 1960. Classification of vegetation for small scale mapping purposes (Symposium on
Forest Classification). Bull. Trop. Ecol. 1(1): 17-18.

*228 (254). ——. 1960. Amateur scientific studies on natural areas. Ecol. Stud. Leafl. 1: 1.

*229 (255). ——. 1960. A plant census of a natural area. Ecol. Stud. Leafl. 2: 1.

230 (256). ——. 1960. Introgression in Artocarpus (Moraceae) in Micronesia. Brittonia 12: 101-113.

231 (257). ——. 1960. Plant collecting as an anthropological field method. El Palacio 67: 125-139.

232 (258). ——. 1960. Vegetation [of Ishigaki-Shima], pp. 51-84, in H.L. Foster, C.H. Stensland, H.G.

May, F.R. Fosberg, and R.J. Alvis, Military Geology of Ishigaki-Shima, Ryikyi-Rett6. Intelligence
Division, Office of the Engineer, U.S. Army Pacific).

233 (261). ——. 1960. Serianthes Benth. (Leguminosae—Mimosoideae-Ingeae). Reinwardtia 5: 293-317.

234 (262). ——. 1960. Vegetation [of the Miyako Archipelago], pp. 165-187, in Doan, D.B., J.E. Paseur,
and F.R. Fosberg, Military Geology of the Miyako Archipelago, Ryiikyi-Retté. Intelligence Division,
Office of the Engineer, U.S. Army Pacific.

235 (263). . 1960. Le réle de la végétation dans la conservation du sol et de l'eau. Comptes rendus de
la Réunion Technique d'Athenes de 1'U.I.C.N. 2: 31-36.

236 (264). . 1960. Random notes on West African palms. Pricipes 4: 125-131.

*237 (266). ——. 1960. Statement, in "Bor Tornetrask regleras?". Kung. Svenska Vetenskapsakad.
Skrifter Naturskyddsarenden 50: 92-93.

*238 (267). . 1960. Our country's landscape—its beauty and diversity, pp. 4-18, in Woodward, C.H..,
The Vanishing Natural Areas of the United States. (Nature Conservancy Brochure).

239 (268). ——. 1960. Remarks on superfluous names and later homonyms. Taxon 9: 243-244.

*240 (268a). ——. 1960. Saturday Review, 14 November. [quoted from by G. Ashford, Conservation?

Of Course—But What Shall We Conserve? San Antonio Express and News, p. 8-G, 10 January.

241 (269). . 1960. Notes on the vegetation of Spruce Knob, West Virginia. Castanea 25: 120-122.

L7 (269a.) ——. 1960. Our brand of smog, [letter to the editor.] The Evening Star (Washington, D.C.)
26 April.

L8 (269b.) . 1960. Of mice and men, [letter to the editor.] The Evening Star (Washington, D.C.) 3
September.

1961

242 (270). Fosberg, F.R. 1961. Humid tropics vegetation. Science 133: 904-906.

243 (273). ——. 1961. Pacific vegetation progress report (1956-1958). Proc. Symposium on Humid
Tropics Vegetation, UNESCO, Tjiawe, Indonesia, 1958, 32-33.

244 (274). ——. 1961. On the possibility of a rational general classification of humid tropical vegetation.
Proc. Symposium on Humid Tropics Vegetation, UNESCO, Tjiawe, Indonesia, 1958, 34-59.

245 (275). ——. 1961. The study of vegetation in Europe. A.I.B.S. Bull. 11(3): 17-19.

246 (277). ——. 1961. Guide to excursion III. Tenth Pacific Science Congress and University of Hawaii,
Honolulu, ix + 207 pp.

247 (282). ——. 1961. Scaevola sericea Vahl versus S. taccada (Gaertn.) Roxb. Taxon 10: 225-226.

248 (283). ——. 1961. What should we map? Colloq. Internat. Centre Nac. Recherche Sci. 97: 23-35.

14

249 (288). ——. 1961. Vegetation-free zone on dry mangrove coasts. U.S. Geol. Survey Prof. Pap. 424-
D: 216-218. =
250 (306). ——. 1961. Soils, pp. 47-50, in Blumenstock, D.I., ed. A report on typhoon effects upon Jaluit

Atoll. Atoll Res. Bull. 75: 1-105.

251 (306a). . 1961. Flora and vegetation, pp. 51-55, in Blumenstock, D.I., ed. A report on typhoon
effects upon Jaluit Atoll. Atoll Res. Bull. 75: 1-105.

252 (306b). ——. 1961. Typhoon effects on individual species of plants. Pp. 57-68. in: Blumenstock,
D.I., ed. A report on typhoon effects upon Jaluit Atoll. Atoll Res. Bull. 75: 1-105.

253 (306c). . 1961. Table of plant species by Islets, pp. 95-104, in Blumenstock, D.I., ed. A report
on typhoon effects upon Jaluit Atoll. Atoll Res. Bull. 75: 1-105.

254 (289). ——. 1961. Qualitative description of the coral atoll ecosystem. Atoll Res. Bull. 81: 1-11.
[(328.) reprinted 1963 [1962.] Proc. Ninth Pacific Sci. Congress, 1957. 4: 161-167.]

255 (290). ——. 1961. Description of Heron Island. Atoll Res. Bull. 82: 1-4.

256 (293). ——. 1961. Southern distribution of Botrychium oneidense and B. multifidum. Amer. Fern J.
51: 175-180.

257 (295). ——. 1961. A classification of vegetation for general purposes. Trop. Ecol. 2: 1-28.

258 (302). ——. 1961. Conservation, pp. 270-273, in Gray, P.. ed., The Encyclopedia of the Biological
Sciences. Reinhold Publishing Corp., New York.

259 (303). ——. 1961. Rubiales, pp. 895-897, in Gray. P., ed., The Encyclopedia of the Biological

Sciences. Reinhold Publishing Corp., New York.

260 (276). Fosberg, F.R., B.J. Garnier, and A.W. Kichler. 1961. Delimitation of the humid tropics.
Geogr. Rev. 51: 333-347. [(276a.) partially republished 1961. Ekistics 12: 448-452.]

261 (291). Fosberg, F.R. and R.F. Thorne. 1961. Vascular plants of Heron Island. Atoll Res. Bull. 82: 5-
13;

262 (292). [Fosberg. F.R., ed.]. 1961. Atoll news and comment. Atoll Res. Bull. 84: 1-14.

263 (271). Blumenstock, D.I., F.R. Fosberg. and C.G. Johnson. 1961. The re-survey of typhoon effects on
Jaluit Atoll in the Marshall Islands. Nature 189(4765): 618-620.

R25 (287). Fosberg, F.R. 1961. The salt-marsh ecosystem, review of Chapman, V.J.. Salt marshes and salt
deserts of the world, 1960. Ecology 42: 847-848.

L9 (291la). ——. 1961. Against Billboards, [letter to the editor.] The Evening Star (Washington, D.C.),
p. A-7, 27 January.

L10 (291b). ——. 1961. Our smog rating, [letter to the editor.] The Evening Star (Washington, D.C.),
10 December.

D38 (272). ——. 1961. Report of representative on the National Research Council. Bull. Ecol. Soc.
Amer. 42: 26-29.

D39 (279). ——. 1961. An inventory of the rare and vanishing plants of the Pacific Basin, [abstract. ]
Abstracts Symp. Pap. Tenth Pacific Sci. Congress, pp. 131-132.

D40 (280). ——. 1961. Plant dispersal in the Pacific. [abstract.] Abstracts Symp. Pap. Tenth Pacific Sci.
Congress, pp. 225-226.

D41 (281). ——. 1961. The ecosystem concept: Introduction, [abstract.] Abstracts Symp. Pap. Tenth
Pacific Sci. Congress, pp. 473-474.

*D42 (285). ——. 1961. Introduction to Symposium on Pacific Basin Biogeography: Tropical Pacific
Relationships. Duplicated for and distributed at Tenth Pacific Sci. Congress, Honolulu.

*D43 (286). ——. 1961. Plant dispersal in the Pacific. Duplicated for and distributed at Tenth Pacific Sci.

Congress, Honolulu.
D44 (278). Fosberg, F.R. and M.S. Doty. 1961. List of botanists of the Pacific area. Duplicated for and
distributed at the Tenth Pacific Science Congress, 126 pp.

1962

264 (294). Fosberg. F.R. 1962. Limits of effective publication. Taxon 11: 27.
265 (299). ——. 1962. Status of the name Geophila D. Don (Rubiaceae). Taxon 11: 180-181.
266 (300). ——. 1962. The Indo-Pacific strand Scaevola again. Taxon 11: 181.

15

267 (304). ——. 1962. Miscellaneous notes on Hawaiian plants—3. Occas. Pap. Bernice P. Bishop Mus.
23(2): 29-44. [Ophioglossum, Thelypteris, Dracaena, Santalum, Chenopodium, Argemone, Leucaena,
Bruguiera, Psidium, Pouteria, Ipomoea, Vitex, Asarina, Psychotria, Sambucus.]

268 (308). . 1962. A brief study of the cays of Arrecife Alacran, a Mexican atoll. Atoll Res. Bull. 93:
1-25+6 unnumbered pages.

269 (307). Fosberg, F.R. and M.-H. Sachet. 1962. Vascular plants recorded from Jaluit Atoll. Atoll Res.
Bull. 92: 1-39.

**270 (305). Oliver, W.R.B., P. Dansereau, and F.R. Fosberg. 1962 [1961]. Report on a botanical
inspection of Ambuklao Dam site and drainage basin. Proc. Eighth Pacific Sci. Congress 6: 210-212.

271 (309). [Fosberg, F.R. and M.-H. Sachet, eds.] 1962. Atoll news and comments. Atoll Res. Bull. 94:
1-19.

D45 (296). Fosberg, F.R. 1962. Report of representative on the National Research Council. Bull. Ecol.
Soc. Amer. 43: 16-17.

D46 (325). . 1962. Report of the chairman of the Standing Committee on Pacific Botany. Proc.
Ninth Pacific Sci. Congress 4: 1-3.

L11 (297). . 1962. Heredity, environment, and culture, [letter to the editor.] Science 136: 914.

*L12 (298). ——. 1962. Rails, rand, and service, [letter to the editor.] Saturday Review 45: 17-18.

L13 (309a). ——-. 1962. Wasted resource, [letter to the editor.] The Evening Star (Washington, D.C.) 21
May.

R26 (301). ——. 1962. Review of Reid, G.K., Ecology of inland waters and estuaries, 1961. Atlantic

Naturalist 17: 203-205.

. 1962. Flora of the Tremblor Range, review of Twisselmann, E.C., Flora of the Tremblor
Range and the neighboring part of the San Joaquin valley, Wasmann J. Biol. 14(2): 161-300, 1956.
Taxon 11: 205.

R27.

1963

272 (311). Fosberg, F.R. 1963. Serianthes dilmyi again. Taxon 12: 34-35.

273 (313). . 1963. The physical background of the humid tropics—substratum, pp. 35-37, in
Symposium on the impact of man on humid tropics vegetation. UNESCO and New Guinea
Government. 402 pp.

274 (314). . 1963. A theory on the origin of the coconut, pp. 73-75, in Symposium on the impact of
man on humid tropics vegetation. UNESCO and New Guinea Government. 402 pp.

275 (315). . 1963. Grazing animals and the vegetation of oceanic islands, pp. 168-169, in
Symposium on the impact of man on humid tropics vegetation. UNESCO and New Guinea
Government. 402 pp.

276 (316). . 1963. Nature and detection of plant communities resulting from activities of early man,
pp. 251-262, in Symposium on the impact of man on humid tropics vegetation. UNESCO and New
Guinea Government. 402 pp.

277 (318). . 1963. Standley and tropical American botany, pp. 97-99, in Williams, L.O., ed., Homage
to Standley, Chicago Natural History Museum, Chicago.

278 (320). ——. 1963. A buttressed elm from Ontario. Rhodora 65: 366.

279 (326). ——. 1963 [1962.] Tropical Pacific grasslands and savannas. Proc. Ninth Pacific Sci.
Congress 4: 118-123.

280 (327). ——. 1963 [1962.] Dynamics of atoll vegetation. Proc. Ninth Pacific Sci. Congress 4: 114-
117.

281 (284, 344). ——. 1963. The island ecosystem, pp. 1-6, in Fosberg, F.R., ed., Man's place in the island
ecosystem, a symposium. Tenth Pacific Sci. Congress, Honolulu.

282 (345). ——. 1963. Introduction [to section I-B, Tropical relationships], pp. 187-188, in Gressitt, J.L..
ed., Pacific Basin Biogeography, a symposium. Tenth Pacific Sci. Congress, Honolulu.

283 (346). ——. 1963. Plant dispersal in Pacific, pp. 273-281, in Gressitt, J.L., ed., Pacific Basin

Biogeography, a symposium. Tenth Pacific Sci. Congress, Honolulu.

16

284 (347). ——. 1963. Disturbance in island ecosystems, pp. 557-561, in Gressitt, J.L., ed., Pacific Basin
Biogeography, a symposium. Tenth Pacific Sci. Congress, Honolulu.

285 (343). Fosberg, F.R., ed. 1963. Man's place in the island ecosystem, a symposium. Tenth Pacific Sci.
Congress, Honolulu. Bishop Museum Press, Honolulu, viii + 264 pp.

286 (330). [Fosberg, F.R. and M.-H. Sachet, eds.] 1963. Atoll news and comment. Atoll Res. Bull. 100:
1-16.

R28 (310). Fosberg, F.R. 1963. Physical geography: effects of nuclear explosions on vegetation comment
on Shields, L.M. and P.V. Wells, Effects of nuclear testing on desert vegetation (Science 135: 38-40,
1962). Geogr. Rev. (New York) 53: 135-137.

R29 (317). ——. 1963. Pesticides and ecology. review of Carson, R., Silent Spring, 1962. Ecology 44:
624.

*D47 (312). ——. 1963. Report of the Ecological Society of America Representative on the National
Research Council. Bull. Ecol. Soc. Amer. 44: 15-16.

D48 (319). ——. 1963. UNESCO meetings (report). Newslett. Hawaiian. Bot. Soc. 2: 110.

D49 (347a). ——. 1963. Report of the standing committee on Pacific Botany, pp. 169-186, in Tuthill,
L.D., ed., Proc. Tenth Pacific Sci. Congress, Honolulu.

D50 (347b). Fosberg, F.R. [chairman] et al. 1963. Report of the American Association for the
Advancement of Science council study committee on natural areas as research facilities. 80 pp. +
appendices.

1964

287 (322). Fosberg, F.R. 1964. Conversations on ecology IX. Garden J. New York Bot. Gard. 14: 63-64.

288 (329). ——. 1964. Studies in Pacific Rubiaceae: V. Brittonia 16: 255-271. [Psychotria.]

289 (331)..——. 1964. Comments on Dr. Donk's proposals. Taxon 13: 175-177.

290 (332). ——. 1964. Do we want a code or a committee? Taxon 13: 177-178.

291 (334). ——. 1964. The correct name of the chiclé. Taxon 13: 254-255. [Manilkara.|

*292 (335). ——. 1964. The question of collecting specimens. Ecol. Stud. Leafl. 3: 4.

*293 (336). ——. 1964. How to investigate the history of a natural area. Ecol. Stud. Leafl. 4: 5-6.

*294 (338). . 1964. Phenological and autecological studies of plant species in natural areas. Ecol.
Stud. Leafl. 5: 7-9.

*295 (339). ——. 1964. Quiet observation. Ecol. Stud. Leafl. 5: 9-10.

296 (340). . 1964. Résumé of the Cycadaceae. Federation Proceedings 23(6) (Part I [Proceedings of
the third conference on the Toxicity of Cycads]): 1340-1341.

297 (340a). [Fosberg, F.R. and M.-H. Sachet, eds.] 1964. Atoll news and comment. Atoll Res. Bull. 108:
1-8.

R30 (321). Fosberg. F.R. 1964. The illness of the world ecosystem, review of Dasmann, R.F., The last
horizon, 1963. Ecology 45: 201-202.

R31 (356). ——. 1964. Review of, Lawrence, G.H.M., ed., Adanson, the Bicentennial of Adanson's
"Familles des Plantes" Part One. Hunt Monograph Series 1, 1963. Torreya 91: 508-509.

R32 (333). ——. 1964. Pefialosa—Tiburon Flora, review of Pefialosa, J., A flora of the Tiburon Peninsula,
Marin County, California (The Wasmann Journal of Botany 21(1): 1-74, 1963). Taxon 13: 219-220.

*R33 (337). ——. 1964. Studies at Pine Hills, review of McCormick, Vascular Flora of Shades State Park
and Pine Hills Natural Area, Indiana. Ecol. Stud. Leafl. 4: 6.

L14 (340b). ——. 1964. Rights and wrongs, [letter to the editor.] The Evening Star (Washington, D.C.)
27 April.

D51 (323). ——. 1964. Report of Representative on the National Research Council. Bull. Ecol. Soc.
Amer. 45: 24-25.

DS52 (324). [Ross, R. and F.R. Fosberg.] 1964. Report of the special committee on neotypes. Taxon 13:
110-114.

1965

298 (341). Fosberg, FR. 1965. Natural bird refuges in the Galapagos. Elepaio 25: 60-67.

299 (348). ——. 1965. Revision of Albizia Sect. Pachysperma (Leguminosae—Mimosoideae).
Reinwardtia 7: 71-90.

300 (349). . 1965. Nomenclature of the Horseradish (Cruciferae). Baileya 13: 1-4. [Cochlearia,
Armoracia.]|
301 (350). . 1965. Lauryl pentachlorphenate protecting herbarium specimens: Application with

brushes. Taxon 14: 165-166.
302 (351). ——. 1965. The entropy concept in ecology, pp. 157-163, in Symposium on ecological
research in humid tropics vegetation. UNESCO, Kuching, Sarawak.

303 (352). . 1965. Field excursions during and after the symposium, pp. 271-288, in Symposium on
ecological research in humid tropics vegetation. UNESCO, Kuching, Sarawak.

304 (355). ——. 1965. Vegetation and the geologist. Trop. Ecol. 6: 3-18. [(355a.) republished (date
unknown), The International Society for Tropical Ecology, Varansi, India, pp. 1-18.]

305 (358a). ——. 1965. Introduction, im Fosberg, F.R. and D. Carroll, eds. Terrestrial sediments and
soils of the northern Marshall Islands. Atoll Res. Bull. 113: 1-6.

306 (358b). ——. 1965. Geology, in Fosberg, F.R. and D. Carroll, eds. Terrestrial sediments and soils of

the northern Marshall Islands. Atoll Res. Bull. 113: 7-22 + 22 plates.

*307 (360, 382). . 1965. Nature Conservancy registry of sites of special scientific interest. Nature
Conservancy News 15: 6.

308 (358d). Fosberg, F.R. and D. Carroll. 1965. Unconsolidated sediments considered as soils, in Fosberg,
F.R. and D. Carroll, eds. Terrestrial sediments and soils of the northern Marshall Islands. Atoll Res.
Bull. 113: 43-65 + 4 plates, 6 tables.

309 (358). Fosberg, F.R. and D. Carroll, eds. 1965. Terrestrial sediments and soils of the northern
Marshall Islands. Atoll Res. Bull. 113: 1-156.

310 (381). Fosberg, F.R. and P.M. Mazzeo. 1965. Further notes on Shenandoah National Park plants.
Castanea 30: 191-205. [Spiraea, Oxalis, Euphorbia, Tilia, Viola, Galium.]

311 (354). Fosberg, F.R. and M.-H. Sachet. 1965. Manual for tropical herbaria. Regnum Veg. 39: 1-132.

*312 (357). Fosberg, F.R., M.-H. Sachet, J.M. Goldberg, and A. Reimer. 1965. World distribution of soil,
rock, and vegetation. U.S. Dept. of the Interior, Geological Survey Report TEI-865: 1-37.

313 (368). Fosberg, F.R. and B.C. Stone. 1965. Leucaena insularum in Guam. Micronesica 2: 67-70.

314 (358c). Fosberg, F.R. [with J.W. Wells, M.S. Doty, and R. Todd.] 1965. Other features, in Fosberg,
F.R. and D. Carroll, eds. Terrestrial sediments and soils of the northern Marshall Islands. Atoll Res.
Bull. 113: 23-30 + 5 plates.

315 (340b). [Fosberg, F.R. and M.-H. Sachet, eds.] 1965. Atoll news and comment. Atoll Res. Bull. 112:

1-14.

*R34 (353). Fosberg, F.R. 1965. Review of Carson, R., The sense of wonder. Nature Conservancy News
IS ISy,

L15 (360a). ——. 1965. Perils of Pesticides, [letter to the editor.] The Evening Star (Washington, D.C.)
p. A-8, 12 January.

L16 (360b). ——. 1965. De Gaulle's Gall, [letter to the editor.] The Evening Star (Washington, D.C.) 2
March.

D53 (342). ——. 1965. [Report of the] Representative on National Research Council. Bull. Ecol. Soc.
Amer. 46: 20.

*D54 (383). [Fosberg, F.R.] 1965. Inter-American specialized conference to deal with problems relating
to the conservation of renewable natural resources in the Western Hemisphere. Nature Conservancy
News 15(4): 5. [Excerpts from report to Nature Conservancy by F.R. Fosberg.]

1966
316 (359). Fosberg, F.R.. 1966. Northern Marshall Islands land biota: birds. Atoll Res. Bull. 114: 1-35.
317 (362). ——. 1966. Miscellaneous notes on Hawaiian plants—4+. Occas. Pap. Bernice P. Bishop Mus.

23(8): 129-138. [Dryopteris, Xyris, Canavalia, Prosopis, Osmanthus, Emilia.]

318 (365). ——. 1966. Vegetation as a geological agent in tropical deltas, pp. 227-233, in Scientific
problems of the Humid tropical zone deltas and their implications. Symposium. UNESCO, Dacca,
Pakistan.

319 (366). ——. 1966. Concluding remarks [of symposium], pp. 409-413, in Scientific problems of the

humid tropical zone deltas and their implications. Symposium. UNESCO, Dacca, Pakistan. [(*367.)
republished in part 1966. Nature and Resources II: 1-5.]

320 (369). ——. 1966. Systematic notes on Micronesian plants. 2. Phytologia 13: 233-241. [Piper,
Peperomia. |

321 (370). ——. 1966. Scirpus americanus Persoon (Cyperaceae) in Costa Rica. Sida 2: 347-348.

322 (371). . 1966. Studies in American Rubiaceae 1. New and noteworthy Costa Rican species. Sida
2: 386-389. [Galium, Gonzalagunia, Hillia, Hoffmannia, Psychotria.|

323 (372). ——. 1966. Restoration of lost and degraded habitats, pp. 503-515, in Darling, F.F. and J.P.
Melton, eds., Future environments of North America. Natural History Press.

324 (373). ——. 1966. The oceanic volcanic island ecosystem, pp. 55-61. in Bowman. R.I., ed. The
Galapagos. University of California Press, Berkeley.

325 (375). ——. 1966. The correct name of the horseradish (Cruciferae). Baileya 14: 60. [Cochlearia,
Armoracia.|

***326 (376). ——. 1966. Vascular plants, pp. 153-238, in Doty, M.S. and D. Mueller-Dombois, Atlas

for bioecology studies in Hawaii Volcanoes National Park. Hawaii Bot. Sci. Pap. no. 2, University of
Hawaii, Honolulu, iii + 507 pp. [republished 1972. Hawaii Agricultural Experiment Station
Miscellaneous Publication no. 89.]

*327 (384). ——. 1966. Some ecological effects of wild and semi-wild exotic species of vascular plants.
Pacific Vegetation Project, National Research Council, Washington, D.C. I[UCN/R.T.10/III/1.2.
[(394.) republished 1967. IUCN pub. n.s., 9: 98-109.]

328 (387). ——. 1966. Critical notes on Pacific Island plants. 1. Micronesica 2: 143-152. [Casuarina,
Alternanthera, Desmodium, Tristellateia, Corchorus, Abutilon, Ambroma, Bruguiera, Ipomoea,
Hemigraphis. |

329 (363). Fosberg, F.R.. E.W. Groves, and D.C. Sigee. 1966. List of Addu vascular plants, pp. 75-92, in
Stoddart, D.R., ed., Reef Studies at Addu Atoll, Maldive Islands: preliminary results of an expedition
to Addu Atoll in 1964. Atoll Res. Bull. 116: 1-122.

330 (374). Fosberg, F.R. and W.L. Klawe. 1966. Preliminary list of plants from Cocos Island, pp. 187-
189, in R.I. Bowman, ed. The Galapagos. University of California Press, Berkeley.

331 (380). Fosberg, F.R. and M.-H. Sachet. 1966. Lebronnecia, gen. nov. (Malvaceae) des Iles Marquises.
Adansonia, n.s., 6: 507-510.

332 (386). —— and ——. 1966. Plants of southeastern Polynesia. Micronesica 2: 153-159. [Panicum,
Crotalaria, Hibiscus, Pavonia, Terminalia, Cordia, Blumea, Vernonia.|

333 (364). [Fosberg, F.R. and M.-H. Sachet, eds.] 1966. Atoll news and comments. Atoll Res. Bull. 117:
1-8.

R35 (377). Fosberg, F.R. 1966. Review of Steele. A.R., Flowers for the King: the expedition of Ruiz and
Pavon and the Flora of Peru, 1964. Bull. Torrey Bot. Club 93: 283.

R36 (378). . 1966. Review of Harris, D.R., Plants, animals and man in the Outer Leeward Islands,
West Indies, 1965. Bull. Torrey Bot. Club 93: 284.

DS55 (361). . 1966. [Report of the] Representative on the National Research Council. Bull. Ecol. Soc.
Amer. 47: 26-27.

D56 (379). ——. 1966. Observations on vegetation patterns and dynamics on Hawaiian and Galapageian
volcanoes, [abstract.] Proc. Eleventh Pacific Sci. Congress 5(3): 3.

D57 (379a). ——. 1966. Report to the Nature Conservancy on the [Ninth] I.U.C.N. General Assembly. 21
pp. [Duplicated for the Nature Conservancy. ]

19

1967
334 (385). Fosberg, F.R. 1967. The Smithsonian Tropical Biology Program [presidential address. |
Symposium on Recent advances in Tropical Ecology, Varanasi, India: 7-12. [(410.) also republished

separately, pp. 1-10.]

335 (389). ——. 1967. Unique Aldabra. Atlantic Naturalist 22: 160-165.
336 (391). ——. 1967. The cult of the expert and numerical taxonomy. Taxon 16: 369-370.
337 (395). . 1967. Appendix 1, a classification of vegetation for general purposes, pp. 73-120, in

Peterken, G.F., Guide to the Check Sheet of IBP Areas, IBP Handbook No. 4.

338 (398). . 1967. Opening remarks: Island ecosystem symposium. Micronesica 3: 3-4.

339 (400). . 1967. Deistveniya podchod k kartirovaniyn Rastetelnocti v praktisheckich tselich (A
pragmatic approach to the practical vegetation mapping). Geobot. Kartogr. 1967: 9-17. [in Russian].

340 (403). ——. 1967. Observations on vegetation patterns and dynamics on Hawaiian and Galapageian
volcanoes. Micronesica 3, 129-134.

341 (407). ——. 1967. Succession and condition of ecosystems. J. Indian Bot. Soc. 46(4): 351-355.

342 (416). ——. 1967. Some relations between ecosystem size and cultural evolution. Proc. Symp.

Recent Adv. Trop. Ecol., Varanasi, India, pp. 702-704.

343 (404). [Fosberg, F.R. and M.-H. Sachet, eds.] 1967. Atoll news and comment. Atoll Res. Bull. 119:
1-6.

344 (388). Buechner, H.K. and F.R. Fosberg. 1967. A contribution toward a world program in tropical
biology. BioScience 17: 532-538.

345 (465). Ripley. S.D. 1967. Perspectives in tropical biology. BioScience 17: 538-540. [Text of address
delivered by S.D. Ripley, slightly paraphrased from text written by F.R. Fosberg at Panama
Conference on Tropical Biology, 9 November 1966. ]

R37 (392). Fosberg, F.R. 1967. Botanical lingua franca, review of Stearn, W.T., Botanical Latin, 1966.
Ecology 48: 175-176.

R38 (393). ——. 1967. Observing bamboos, review of McClure, F.A., The Bamboos-a fresh perspective,
1966. Ecology 48: 176.

R39 (409). ——. 1967. Review of Wenkam, R., Kauai and the park country of Hawaii, 1967. Atlantic
Naturalist 22: 233-234.

D58 (390). ——. 1967. [Report of the] Representative on the National Research Council. Bull. Ecol. Soc.
Amer. 48: 27-28.

D59 (405). Fosberg, F.R., C. Cottam, A.M. Stebler, and J. Major. 1967. An ecological context for the use
and conservation of natural resources. 6 pp. duplicated and distributed by Applied Ecol. Sect., Ecol.
Soc. Amer.

1968

346 (397). Fosberg, F.R. 1968. Systematic notes on Micronesian plants, 3. Phytologia 15: 496-502.
[Sagittaria, Egeria, Lepturus, Crinum, Spathoglottis, Lepidium, Mimosa, Cassia, Clitoria, Cordia.|

347 (406). . 1968. Investigation of Aldabra Atoll. Bull. Regional Centre for Science and Technology
for Africa, Nairobi, Kenya 3(2): 39-41.

348 (408). . 1968. Polypodium vulgare on Long Island. Amer. Fern J. 58: 153-154.

349 (411). ——. 1968. Studies in Pacific Rubiaceae: VI-VII. Brittonia 20: 287-294. [Hedyotis, Gouldia,
Coprosma.|

350 (429). ——. 1968. Critical notes on Pacific Island plants. 2. Micronesica 4: 255-259. [Thelypteris,

Crassula, Desmodium, Alyxia, Scaevola.|

351 (434). . 1968. Scientific need for parks and reserves, pp. 380-381, in Conservation in tropical
South East Asia, IUCN Publ., new series vol. 10.

R40 (396). ——. 1968. Gigantic grasses, review of McClure, F.A., The Bamboos, a fresh perspective,
1966. BioScience 18: 55.

R41 (399). ——. 1968. An oceanographic encyclopedia, review of Fairbridge, R.W., ed., The
encyclopedia of oceanography, 1966. Ecology 49: 187-188.

20

R42 (401). ——. 1968. A comprehensive text, review of Kiichler, A.W., Vegetation Mapping, 1967.
BioScience 18: 143-144. ;

R43 (413). ——. 1968. Review of Twisselmann, E.C., A Flora of Kern County, California, 1967. Taxon
17: 565-566.

R44 (402). . 1968. A major synthesis of biology, review of Platt, R.B. and G.K. Reid, Bioscience,

1967. BioScience 18: 248-249.
1969

352 (417). Fosberg, F.R. 1969. Aura of pollution. Biol. Conservation 1: 231-232.

353 (421). ——. 1969. Sabah (North Borneo), Malaysia: Sepilok Forest Endangered. Biol. Conservation
1: 339-340.

354 (422). ——.. 1969. The problem of isolation in the lowland tropical rain forest. Eleventh Int. Bot.
Congress Abst. 62.

355 (423). ——-. 1969. Miscellaneous notes on Hawaiian plants—5. Occas. Pap. Bernice P. Bishop Mus.
24(2): 9-24. [Gleichenia, Adiantum, Polypodium, Thelypteris, Setaria, Dianella, Arundina,
Odontochilus, Santalum, Polygonum, Rumex, Lepidium, Fragaria, Cassia, Medicago, Zanthoxylum,
Abutilon, Hypericum.|

356. . 1969. Correction on Serianthes dilmyi (Leguminosae). Taxon 18: 351-352.

357 (424). ——. 1969. The nomenclature of Sphenomeris chinensis [Filic.]. Taxon 18: 596-598.

358 (425). ——. 1969. Plants of Satawal Island, Caroline Islands. Atoll Res. Bull. 132: 1-13.

359 (428). ——. 1969. Observations on the Green Turtle in the Marshall Islands, in Island news and
comment. Atoll Res. Bull. 135: 9-12.

360 (431). ——. 1969. Commentary—A Balanced Ecology. Ecology, 50: 463-464.

361 (433). ——. 1969. Paleobotany of the oceanic Pacific Islands, in Leopold, E.B., Miocene pollen and
spore flora of Eniwetok, Marshall Islands. U.S.G.S. Prof. Pap. 260-II: 1140.

*362 (437). ——. 1969. Diversity and cooperation among conservation organizations. II Simposio y

Foro de Biologia Tropical Amazonica. Leticia, Colombia: 164-167.

363 (426). Fosberg, F.R. and M. Evans. 1969. A collection of plants from Fais, Caroline Islands. Atoll
Res. Bull. 133: 1-15.

364 (414). Fosberg, F.R. and M.-H. Sachet. 1969. Wake Island vegetation and flora, 1961-1963. Atoll
Res. Bull. 123: 1-15.

365 (415). [Fosberg, F.R. and M.-H. Sachet, eds.] 1969. Island news and comment. Atoll Res. Bull. 126:
1-19.

366 (427). [Fosberg, F.R. and M.-H. Sachet, eds.] 1969. Island news and comment. Atoll Res. Bull. 135:
1-17.

R45 (412). Fosberg, F.R. 1969. The California Islands, review of Philbrick, R.N., ed., Proceedings of the
symposium on the biology of the California islands, 1967. Ecology 50: 167.

*R46 (418). ——. 1969. Review of Brower, K., ed., Galapagos, the Flow of Wildness, 2 vols.,
photographs by E. Porter, Sierra Club, 1968. [reprint does not show place of publication. }

R47 (418a). ——. 1969. Review of Nelson, B., Galapagos, Islands of Birds, 1968. Atlantic Naturalist 24:
224-225.

R48 (430). ——. 1969. Vegetation map bibliography, review of Kiichler, A.W., ed., International

bibliography of vegetation maps. Vol. 3—U.S.S.R., Asia & Australia, 1968. Ecology 50: 749-750.
D60 (419). . 1969. Remarks on the International Code and proposals to change it. I.-X. Duplicated
for and distributed at the Eleventh International Botanical Congress, Seattle, pp. 1-9.
*D61 (420). ——. 1969. Geomorphic cycle on Aldabra—hypothesis, [abstract.] Mandapam symposium on
corals and coral reefs, abstracts, pp. 26-27.

1970
367 (432). Fosberg, F.R. 1970. Desert Wilderness. The Living Wilderness, 34(109): 17-24.
368 (436). ——. 1970. The tropical agriculture panacea. BioScience 20: 793.
369 (445). ——. 1970. The problem of isolation in the lowland tropical rain-forest. Trop. Ecol. 11: 162-

168.

21

370 (448). ——. 1970. Island faunas and floras, pp. 309-311, in McGraw-Hill Encyclopedia of Science
and Technology, ed. 3.

371 (440). Fosberg, F.R. and T. Blunt. 1970 [1969.] Vernon Black Gum Swamp. Rhodora 72: 280-282.

372 (441a). Fosberg, F.R. and S.A. Renvoize. 1970. Plants of Farquhar Atoll, pp. 27-34, in Stoddart,
D.R., ed., Coral Islands of the Western Indian Ocean. Atoll Res. Bull. 136: 1-224.

373 (441b). —— and ——. 1970. Plants of Cosmoledo Atoll, pp. 57-66, in Stoddart, D.R., ed., Coral
Islands of the Western Indian Ocean. Atoll Res. Bull. 136: 1-224.
374 (44 1c). and ——. 1970. Plants of Astove, pp. 101-112, in Stoddart, D.R., ed., Coral Islands of

the Western Indian Ocean. Atoll Res. Bull. 136: 1-224.

375 (441d). and . 1970. Plants of Assumption Island, pp. 147-154, in Stoddart, D.R., ed., Coral
Islands of the Western Indian Ocean. Atoll Res. Bull. 136: 1-224.

376 (441e). —— and ——. 1970. Plants of Desroches, pp. 167-170, in Stoddart, D.R., ed., Coral Islands
of the Western Indian Ocean. Atoll Res. Bull. 136: 1-224.

377 (441f). —— and . 1970. Plants of Remire (Eagle) Island Amirantes, pp. 183-186, in Stoddart,
D.R., ed., Coral Islands of the Western Indian Ocean. Atoll Res. Bull. 136: 1-224.

378 (441g). and . 1970. Plants of African Banks (Iles Africaines), pp. 193-194, in Stoddart.
D.R., ed., Coral Islands of the Western Indian Ocean. Atoll Res. Bull. 136: 1-224.

379 (441). Doty, M.S., F.R. Fosberg, and D. Mueller-Dombois. 1970. Initial site studies for the
international program in the tropical and far western Pacific. Micronesica 5: 283-293.

R49 (438). Fosberg, F.R. 1970. S.E. Asiatic Floras, review of Reed, C.F., Bibliography to Floras of
Southeast Asia, 1969. Taxon 19: 432.

R50 (439). ——. 1970. A local flora from San Francisco, review of McClintock, E., W. Knight, and N.
Fahy, A Flora of the San Bruno Mountains, San Mateo County, California, 1968. Taxon 19: 432.

L17 (435). ——. 1970. Floating gobs of oil, [letter to the editor.] Science 168: 917.

L18 (445a). ——. 1970. Insect overkill, [letter to the editor.] The Evening Star (Washington, D.C.), 19
June.

1971

380 (442). Fosberg, F.R. 1971. Preliminary survey of Aldabra vegetation. Philos. Trans., Ser. B, 260:
215-225.

381 (446). ——. 1971. Endangered island plants. Bull. Pacific Trop. Bot. Gard. 1(3): 1-7.

382 (451). ——. 1971. The Fern Vegetation of Aldabra Atoll. Amer. Fern J. 61: 97-101. [Acrostichum.]

383 (452). ——. 1971. Psidium L. (Myrtaceae) in Ceylon. Ceylon J. Sci., Biol. Sci. 9: 58-60.

384 (456). ——. 1971. Mangroves v. Tidal Waves. Biol. Conservation 4: 38-39.

385 (447). Fosberg, F.R. and A.A. Bullock. 1971. List of Diego Garcia vascular plants, in Stoddart, D.R.
& J.D. Taylor, eds. Geography and ecology of Diego Garcia Atoll, Chagos Archipelago. Atoll Res.
Bull. 149: 143-160.

386 (444). [Fosberg, F.R., M.-H. Sachet, and D.R. Stoddart, eds.] 1971. Island news and comment. Atoll
Res. Bull. 148: 1-38.

387 (461). Sachet, M.-H. and Fosberg, F.R. 1971. Island bibliographies supplement—Micronesian botany;
land, environment. and ecology of coral atolls; vegetation of tropical Pacific islands. National
Academy of Sciences, Washington D.C., 427 pp.

388 (443). Stoddart, D.R., J.D. Taylor, F.R. Fosberg, and G.E. Farrow. 1971. Geomorphology of Aldabra
Atoll. Philos. Trans., Ser. B, 260: 31-65.

R51 (449). Fosberg, F.R. 1971. Review of Wiggins, I.L. and D.M. Porter, Flora of the Galapagos Islands,
1971. Atlantic Naturalist 26: 139-141.

R52 (450). ——. 1971. Review of Howell, J.T., Marin Flora, second ed., 1970. Taxon 20: 369.

R53 (457). ——. 1971. Amazonia, review of Meggers, B.J., Amazonia Man and Culture in a Counterfeit
Paradise, 1971. Ecology 52: 952.

1972

389 (453). Fosberg, F.R. 1972. The axis deer problem, pp. 201-202, in Mueller-Dombois, D., Island

ecosystems stability and evolution subprogram, U. S. International Biological Program Techn. Rept. 2.

22

390 (454). ——. 1972. Synopsis of views of F.R. Fosberg relative to the harvesting of koa and hapuu on
the Big Island, p. 205, in D. Mueller-Dombois, Island ecosystems stability and evolution subprogram,
U. S. International Biological Program Techn. Rept. 2.

391 (455). ——. 1972. Man's effects on island ecosystems, pp. 869-880, in Farvar, M.T. and J. P. Milton,
The Careless Technology, Natural History Press, Garden City, New York.
392 (464). ——. 1972. Guide to excursion III, Tenth Pacific Science Congress, revised edition. University

of Hawaii, Honolulu, x + 249 pp.
393 (466). _ 1972. Our native plants—a look to the future. National Parks and Conservation
Magazine 46(11): 17-21.

394 (467). . 1972. On the importance of preserving diversity in ecosystems. Comptes Rendus Conf.
Int. Cons. Nat. Res. Madagascar, I.U.C.N. N.S. Suppl. 36: 81-84.

395 (469). ——. 1972. Emilia (Compositae) in Ceylon. Ceylon J. Sci., Biol. Sci. 10: 61-69.

396 (470). ——. 1972. The value of systematics in the environmental crisis. Taxon 21: 631-634.

397 (479). ——. 1972. Geomorphic cycle on Aldabra—hypothesis. Proc. Symp. Corals and Coral Reefs
(Mar. Biol. Assoc. India): 469-475.

398 (495). ——. 1972. List of vascular flora, pp. 9-13, in Stoddart, D.R., Reef islands of Rarotonga. Atoll

Res. Bull. 160: 1-14 + 7 figs.

399 (497a). _ 1972. Morotiri (Bass Rocks) Austral Islands, in Island news and comment. Atoll Res.
Bull. 162: 9-10.

400 (458). Fosberg, F.R. and M.-H. Sachet. 1972. Thespesia populnea (L.) Solander ex Correa and
Thespesia populneoides (Roxburgh) Kosteletsky (Malvaceae). Smithsonian Contr. Bot. 7: 1-13.

401 (460). and . 1972. Three Indo-Pacific 7he/lypteris species reinterpreted and a new African
species described. Smithsonian Contr. Bot. 8: 1-10.

402 (468). —— and . 1972. Status of floras of western Indian Ocean islands. Comptes Rendus Conf.
Int. Cons. Nat. Res. Madagascar, I.U.C.N. N.S. Suppl. 36: 152-155

403 (471). ——— and ——. 1972. Plants of southeastern Polynesia. 2. Micronesica 8: 43-49.
[Peperomia, Ascarina, Weinmannia, Alstonia, Ochrosia, Neiosperma.|

404 (497). [Fosberg, F.R.. M.-H. Sachet, and D.R. Stoddart, eds.] 1972. Island news and comment. Atoll
Res. Bull. 162: 1-26.

405 (496). Stoddart. D.R. and F.R. Fosberg. 1972. South Indian sand cays. Atoll Res. Bull. 161: 1-23.

R54 (459). Fosberg, F.R. 1972. Island biology, review of Stern, W.L., ed., Adaptive aspects of insular
evolution. 1971. BioScience 22: 261.

R55 (463). ——. 1972. Small island ecosystem, review of van Zinderen Bakker, Sr., JM. Winterbottom,
and R.A. Dyer, eds., Marion and Prince Edward Islands. Report on the South African Biological and
Geological Expedition, 1965-1966, 1971. Science 176: 662-663.

D62 (462). _ 1972. The Ceylon flora project status report, 1972. Duplicated. 17 pp.

1973

406 (472). Fosberg, F.R. 1973. Temperate zone influence on tropical forest land use: a plea for sanity.,
pp. 345-350, in Meggers, B.J., E.S. Ayensu, and W.D. Duckworth. Tropical Forest Ecosystems in
Africa and South America. A comparative review. Smithsonian Institution Press, Washington, D.C.,
vili + 350 pp.

407 (473). ——. 1973. Caesalpinia major a legitimate name. Taxon 22: 162-163.

408 (474). ——. 1973. Reflections after the symposium on systematic biological collections, July 6-8,
1972. Pl. Sci. Bull. 19(1): 4-6.

409 (475). ——. 1973. Vascular plants—widespread island species, pp. 167-169, in Costin, A.B., and R.H.
Groves, Nature Conservation in the Pacific, Australian National University Press, Canberra.

410 (476). ——. 1973. Past, present and future conservation problems of oceanic islands, pp. 209-215, in

Costin, A.B., and R.H. Groves, Nature Conservation in the Pacific, Australian National University
Press, Canberra.

23

411 (480). ——. 1973. On present condition and conservation of forests in Micronesia. Pacific Sci.
Assoc. Standing Comm. on Pacific Bot. Symposium: Planned Utilization of the Lowland Tropical
Forest, Bogor, Indonesia, 1971: 165-171.

412 (481). ——. 1973. The name of the Octopus Tree. Baileya 19: 45-46. [Brassaia.]

413 (484). ——. 1973. Type specimens of Buxus sempervirens Linnaeus. Boxwood Bull. 13(2): 18-21.

414 (483). Sachet, M.-H. and F.R. Fosberg. 1973. Remarks on Halophila (Hydrocharitaceae). Taxon 22:
439-443.

R56 (477). Fosberg, F.R. 1973. Review of Lewis, D.. We, the navigators, 1972. Geogr. Rev. (New York)
63: 428-430.

R57 (485). . 1973. Review of Zwinger, A.H. and B.E. Willard, Land Above the Trees, 1972. Biol.

Conservation 5: 317-318.

*L19 (482). ——. 1973. Letter on the proposed road through Abisko from Kiruna to Narvik, p. 21, in
Andersson et al, Project K-N, Royal Academy of Sciences, Stockholm.

D63 (478). Sachet, M.-H. and F.R. Fosberg. 1973. The Marquesas in Pacific Biogeography, [abstract. |
Island Ecosystem, I.R.P. Techn. Rept. 26: 37.

1974

415 (486). Fosberg, F.R. 1974. Sketch of the St. Croix flora, pp. 239-244, in Multer, H.G. and L.C.
Gerhard, eds., Guidebook to the Geology and ecology of some marine and terrestrial environments of
St. Croix. Special publication No. 5, West Indies Laboratory, Fairleigh Dickinson University,
Christiansted, St. Croix, 304 pp.

416 (487). . 1974. Notes on Micronesian Pteridophyta, III. Ophioglossum nudicaule in the Caroline
Islands. Amer. Fern J. 64: 1-2.

417 (489). ——. 1974. Studies in American Rubiaceae 2. Ayuque, Balmea stormae, an endangered
Mexican species. Sida 5: 268-270.

418 (494). ——. 1974. Miscellaneous notes on the flora of Aldabra and neighboring islands: III. Kew
Bull. 29: 253-266. [Flacourtia, Calophyllum, Malleastrum, Poupartia, Lactuca, Plumbago, Ehretia,
Ipomoea, Lagrezia, Acalypha, Euphorbia, Maillardia.|

419 (490). Fosberg, F.R. and M.V.C. Falanruw. 1974. A new Micronesian Terminalia (Combretaceae).
Phytologia 28: 469-470.

420 (491). Fosberg, F.R. and M.-H. Sachet. 1974. A new variety of Kagraea berteriana (Gentianaceae).
Phytologia 28: 470-472.

421 (493). —— and ——. 1974. Plants of southeastern Polynesia. 3. Micronesica 10: 251-256.
[Waltheria, Alyxia, Ochrosia, Neiosperma.]|

422 (492). Grant, M.L., F.R. Fosberg, and H.M. Smith. 1974. Partial flora of the Society Islands:
Ericaceae to Apocynaceae. Smithsonian Contr. Bot. 17: 1-85. [Vaccinium, Styphelia, Ardisia,
Rapanea, Plumbago, Palaquium, Manilkara, Planchonella, Lucuma, Chrysophyllum, Styrax, Olea,
Jasminum, Geniostoma, Fagraea, Allamanda, Lepinia, Plumeria, Alstonia, Catharanthus, Ervatamia,
Alyxia, Ochrosia, Cerbera, Thevetia, Nerium.]

423 (498). Mueller-Dombois, D. and F.R. Fosberg. 1974. Vegetation map of Hawaii Volcanoes National
Park (at 1: 52,000). Cooperative Nat. Park Resources Unit: Tech. Rept. 4: 1-44, map sheets 1-25.

R58 (488). Fosberg, F.R. 1974. Review of Polunin, N., ed., The Environmental Future, 1972. Geogr. Rev.
(New York) 64: 445-447.

1975
424 (500). Fosberg, F.R. 1975. The deflowering of Hawaii. National Parks and Conservation Magazine
49: 4-10.
425 (501). ——. 1975. Dr. Raven's proposals. Taxon 24: 192-193.
426 (502). ——. 1975 [1974.] Phytogeography of atolls and other coral islands. Proceedings of the
Second International Coral Reef Symposium. 1: 389-396.
427 (503). ———. 1975. Revised check-list of vascular plants of Hawaii Volcanoes National Park.

Cooperative National Park Resources Studies Unit Hawaii, Technical Report no. 5: 1-19.

24

428 (511). ——. 1975. Identification of vascular flora of Namoluk Atoll, eastern Caroline Islands, in
Marshall, M. the natural history of Namoluk Atoll. eastern Caroline Islands. Atoll Res. Bull. 189:
23-48.

429 (512). . 1975. Typification and author citation of \/erremia tridentata ssp. hastata Van

Ooststroom. Taxon 24: 541.

430 (513). . 1975. Vascular plants of Aitutaki, pp. 73-84, in Stoddart, D.R. and P.E. Gibbs, eds.,
Almost-Atoll of Aitutaki: Reef Studies in the Cook Islands, South Pacific. Atoll Res. Bull. 190: 1-
158.

431 (558). ——. 1975. Phytogeography of Micronesian mangroves, pp. 23-42, in Walsh, G., S. Snedaker,

and H. Teas, Proceedings of International Symposium on Biological Management of Mangroves, The
University of Florida Press, Gainesville.

432 (507). Fosberg, F.R., M.V.C. Falanruw, and M.-H. Sachet. 1975. Vascular flora of the northern
Marianas islands. Smithsonian Contr. Bot. 22: 1-45.

433 (508). Fosberg, F.R. and M.V.C. Falanruw. 1975. Noteworthy Micronesian Plants. 1. Micronesica
11: 77-80. [Eustachys, Boehmeria, Canavalia, Euphorbia, Ochrosia, Eupatorium.|

434 (499). Fosberg, F.R. and D. Herbst. 1975. Rare and endangered species of Hawaiian vascular plants.
Allertonia 1: 1-72.

435 (505). Fosberg, F.R. and M.-H. Sachet. 1975. Flora of Micronesia, 1. Gymnospermae. Smithsonian
Contr. Bot. 20: 1-15. [Cycas, Araucaria, Podocarpus, Pinus, Cryptomeria, Cupressus, Juniperus,

Gnetum.|

436 (506). —— and ——. 1975. Flora of Micronesia. 2. Casuarinaceae, Piperaceae, and Myricaceae.
Smithsonian Contr. Bot. 24: 1-28. [Casuarina, Peperomia, Piper, Myrica.]

437 (509). —— and ——. 1975. Noteworthy Micronesian Plants. 2. Micronesica 11: 81-84. [Portulaca,

Connarus, Salomonia, Bacopa, Limnophila, Striga.|

438 (510). and . 1975. Polynesian plant studies. 1-5. Smithsonian Contr. Bot. 21: 1-25.
1. Vaccinium cereum (L.f.) Forster f. in Tahiti and the Marquesas, pp. 1-3.
2. Myrsine L. in eastern Polynesia, pp. 3-11.
3. Three species of Geniostoma J. R. & G. Forster, pp. 11-14.
4. Ipomoea tiliacea (Willdenow) Choisy in the Pacific, pp. 14-15.
5. Annotations to the Hawaiian flora, pp. 15-23. [Adiantum, Salicornia, Crotalaria, Melochia,

Waltheria, Plantago, Hedyotis.|

439 (510a). [Fosberg, F.R. and M.-H. Sachet, eds.] 1975. Island news and comments. Atoll Res. Bull.
185: 1-39.

R59 (515). Fosberg, F.R. 1975. Review of Faulkner, D. This Living Reef, 1974. AAAS Science Books
and Films 11: 74.

D64 (504). ——. 1975. Outline of the contents of Biogeography. Two page duplicated excerpt from talk,
distributed to audience at talks Geography, Ecology, and Biogeography at AAG annual meeting,
Milwaukee, Wisconsin, 21 April.

D65 (515c). ———. 1975. Biogeoclimatic patterns in Micronesia, [abstract.] 13th Pacific Science
Congress, Vancouver, Proc. 1, Abstracts of papers, pp. 97-98.
D66 (515d). ——. 1975. An overview of natural area preservation in the world, [abstract.] 13th Pacific

Science Congress, Vancouver, Proc. 1, Abstracts of papers, p. 139.

D67 (515e). . 1975. Development of ecosystem concept, [abstract.] 13th Pacific Science Congress,
Vancouver, Proc. 1, Abstracts of papers, pp. 248-249.

D68 (515b). Fosberg, F.R. and M.-H. Sachet. 1975. Micronesia: status of floristic knowledge, [abstract.]
13th Pacific Science Congress, Vancouver, Proc. 1, Abstracts of papers, p. 98.

D69 (515a). Sachet, M.-H. and F.R. Fosberg. 1975. The Marquesas in Pacific plant geography, [abstract. ]
13th Pacific Science Congress, Vancouver, Proc. 1, Abstracts of papers, p. 110.

1976

440 (514). Fosberg, F.R. 1976. Coral island vegetation. Chapter 8, pp. 255-277, in Jones, O.A. and R.
Endean, Biology and Geology of Coral Reefs. Volume III Biology 2, Academic Press, New York.

25

441 (516). ——. 1976. Status of the name Chloris barbata (L.) Swartz. Taxon 25: 176-178.

442 (517). ——. 1976. Typification of Sadleria hillebrandii Robinson. Taxon 25: 187-188.

443 (518). ——. 1976. Bobea elatior again. Taxon 25: 188.

444 (519). ——. 1976. Revisions in the Flora of St. Croix, U.S. Virgin Islands. Rhodora 78: 79-119.

[Jatropha, Lantana, Lippia.]

445 (520). . 1976. Geography, ecology, and biogeography. Ann. Assoc. Amer. Geogr. 66: 117-128.

446 (521). ——. 1976. Ipomoea indica taxonomy: a tangle of morning glories. Bot. Not. 129: 35-38.

447 (526). ——. 1976. Distributional extensions of marine Spermatophytes. Micronesica 12: 317-318.

R60 (522). ——. 1976. Review of Whitmore, T.C., Tropical Rain Forests of the Far East, 1975. Taxon
25: 510.

R61 (523). ——. 1976. Review of Udvardy, M.D.F., A Classification of the Biogeographical Provinces of
the World, 1975. Environm. Conservation 3: 313-314.

R62 (528). ——. 1976. Review of Carlquist, S., Island Biology, 1974. Economic Botany 30: 6-7.

R63 (529). ——. 1976. Review of Everist, S.L., Poisonous Plants of Australia, 1974. Economic Botany
30: 6.

R64. ——. 1976. Review of Sterling, T., The Amazon, 1975. Taxon 25: 658.

1977

448 (533). Fosberg, F.R. 1977. Paspalum distichum again. Taxon 26: 201-202.

449 (534). . 1977. Miscellaneous notes on the flora of Aldabra and neighbouring islands: IV. A new
Bulbostylis and observations on Cyperus (Cyperaceae). Kew Bull. 31: 829-835.

450 (535). . 1977. Miscellaneous notes on the flora of Aldabra and neighbouring islands: V.
Pandanus tectorius Parkinson, sensu latissimo (Pandanaceae). Kew Bull. 31: 837-840.

451 (539). . 1977. Miscellaneous notes on the flora of Aldabra and neighbouring islands: VI.
Portulaca (Portulacaceae) in the Aldabra Group. Kew Bull. 32: 253-258.

452 (551). . 1977. Documentation of identifications in ecological work. Trop. Ecol. 18: 97-98.

453 (541). Fosberg, F.R., P. Boiteau, and M.-H. Sachet. 1977. Nomenclature of the Ochrosiinae
(Apocynaceae): 2. Synonymy of Ochrosia Juss. and Neiosperma Raf. Adansonia, n.s., 17: 23-33.

454 (532). Fosberg, F.R., M.V.C. Falanruw, and M.-H. Sachet. 1977. Additional records of vascular
plants from the northern Mariana Islands. Micronesica 13: 27-31.

455 (524). Fosberg, F.R. and M.-H. Sachet. 1977. Flora of Micronesia. 3. Convolvulaceae. Smithsonian
Contr. Bot. 36: 1-34. [Aniseia, Argyreia, Cuscuta, Evolvulus, Ipomoea, Merremia, Operculina,
Stictocardia.]|

456 (540). and . 1977. Nomenclature of the Ochrosiinae (Apocynaceae): 1. Application of the
Names Neiosperma Raf. and Calpicarpum G. Don. Adansonia, n.s., 17: 19-22.

457 (536). [Fosberg, F.R., D.R. Stoddart, and M.-H. Sachet, eds.] 1977. Island news and comment. Atoll
Res. Bull. 219: 1-30.

458 (542). Tsuda, R. T., F.R. Fosberg, and M.-H. Sachet. 1977. Distribution of seagrasses in Micronesia.
Micronesica 13: 191-198.

R65 (527). Fosberg, F.R. 1977. Review of Wace, N.M. and M.W. Holdgate, Man and Nature in the
Tristan da Cunha Islands. Environm. Conservation 4: 73.

R66 (530). ——. 1977. Review of Dasmann, R.R., Environmental Conservation, 4th ed. 1976.
Environm. Conservation 4: 155-156.

R67 (531). ——. 1977. Review of Hoare, M.E., The Tactless Philosopher: Johann Reinhold Forster
1729-1798, 1975. Taxon 26: 120-121.

R68 (537). ——. 1977. Review of Chapman, V.J., Mangrove Vegetation, 1976. Taxon 26: 113.

R69 (538). ——. 1977. Review of Herklots, G., Flowering Tropical Climbers, 1976. Taxon 26: 119.

L18 (525). . 1977. Expert on Hawaiian plants responds [reader comment.] National Parks and

Conservation Magazine 51(2): 28-29.

26

1978
459 (543). Fosberg, F.R. 1978. Studies in the genus Boerhavia L. (Nyctaginaceae), 1-5. Smithsonian

Contr. Bot. 39: 1-20.
1. Introduction to the genus Boerhavia L., pp. 1-4.
2. Clarification of Boerhavia diffusa L. and Boerhavia repens L., pp. 4-5.
3. Typification of Boerhavia mutabilis R. Brown and Boerhavia pubescens R. Brown, pp. 5-8.
4. Boerhavia repens group, pp. 8-13.
5. Boerhavia rubicunda Steudel and somewhat similar species, pp. 13-18.

460 (545). _ 1978 [1977.] A fossil Garcinia fruit from the New Hebrides, Melanesia. Pacific Sci. 31:
293-297.

461 (546). ——. 1978. Miscellaneous notes on the flora of Aldabra and neighbouring islands: VII.
Eugenia elliptica var. levinervis (Myrtaceae). Kew Bull. 33: 133-134.

462 (547). ——. 1978. Miscellaneous notes on the flora of Aldabra and neighbouring islands: VIII.

Hedvotis subgenus Oldenlandia (Rubiaceae). Kew Bull. 33: 135-140.
463 (548). . 1978. Miscellaneous notes on the flora of Aldabra and neighbouring islands: IX. The
native Aldabra Solanum. Kew Bull. 33: 141-142.

464 (549). . 1978. Miscellaneous notes on the flora of Aldabra and neighbouring islands: X. The
Aldabra Clerodendrum (Verbenaceae). Kew Bull. 33: 143-144.

465 (550). ——. 1978. Miscellaneous notes on the flora of Aldabra and neighbouring islands: XI.
Critical notes on Euphorbiaceae. Kew Bull. 33: 181-190. [Euphorbia, Margaritaria, Phyllanthus.|

R70 (544). ——. 1978. Review of Bryson, R.A. and T.J. Murray, Climates of Hunger: Mankind and the
World's Changing Weather. Environm. Conservation 5: 75.

R71 (544a). ——. 1978. Review of Heintze, C., Summit Lake, Four Seasons in the Sierras, 1976.

Environm. Conservation 5: 160.
1979
466 (552). Fosberg, F.R. 1979. The woody Rubiaceae of Aldabra Island (Indian Ocean). Phytologia 41:
347-362. [Canthium, Guettarda, Polysphaeria, Psychotria, Tarenna, Triainolepis, Tricalysia.|

467 (554). ——. 1979. Miscellaneous notes on the flora of Aldabra and neighbouring islands: XII.
Boerhavia repens L. var. maris-indici Fosberg (Nyctaginaceae) validated. Kew Bull. 33: 398.

468 (561). ——. 1979. Tropical floristic botany—concepts and status—with special attention to tropical
islands, pp. 89-105, in Larsen, K. and B. Holm-Nielsen, Tropical Botany. Academic Press, London.
xi + 453.

469 (562b). . 1979. Plants of D'arros Island, pp. 19-25, in Stoddart, D.R., M.J. Coe, and F.R.
Fosberg. D'arros and St. Joseph, Amirante Islands. Atoll Res. Bull. 223: 1-48.

470 (562c). ——. 1979. Plants of St. Joseph Atoll, pp. 43-48, in Stoddart. D.R., M.J. Coe, and F.R.
Fosberg. D'arros and St. Joseph, Amirante Islands. Atoll Res. Bull. 223: 1-48.

471 (574). ——. 1979. Conservation of diversity. Environm. Aware. 2: 109-130.

472 (553). Fosberg, F.R. and M.-H. Sachet. 1979. Callicarpa erioclona Schauer (sensu stricto).
Phytologia 41: 363-365.

473 (555). ——- and ——. 1979. Maesa (Myrsinaceae) in Micronesia. Phytologia 44: 362-369.

474 (555a). —— and ——. 1979. Correction to Af/aesa (Myrsinaceae) in Micronesia. Phytologia 44: 502.

475 (557). Fosberg, F.R., M.-H. Sachet, and R.L. Oliver. 1979. A geographical checklist of the
Micronesian Dicotyledonae. Micronesica 15: 41-295.

476 (562a). Stoddart, D.R..M.J. Coe, and F.R. Fosberg. 1979. D'arros and St. Joseph. Amirante Islands.
Atoll Res. Bull. 223: 1-48.

1980

477 (562). Fosberg, F.R. 1980. Whither terrestrial ecosystems? Preservation of the habitat of man, pp.
64-93, in Polunin, N., ed. Growth without Ecodisasters?. MacMillan Press, London and Basingstoke,
Xxvi + 675 pp.

478 (567). ——. 1980. The plant eco-system for Moenjodaro. UNESCO, Restricted Technical Report
RP/1977-78/4.121.6:1-15.

27

479 (556). Fosberg, F.R., D. Otobed, M.-H. Sachet, R.L. Oliver, D.A. Powell, and J.E. Canfield. 1980.
Vascular plants of Palau with vernacular names. Duplicated, Smithsonian Institution, 43 pp.

480 (560). Fosberg, F.R. and S.A. Renvoize. 1980. The flora of Aldabra and neighbouring islands. Kew
Bull., Additional Series, 7: 1-358. [4crostichum, Nephrolepis, Annona, Cissampelos, Brassica,
Capparis, Cleome, Gynandropsis, Maerua, Flacourtia, Ludia, Portulaca, Calophyllum, Abutilon,
Gossypium, Hibiscus, Sida, Thespesia, Corchorus, Grewia, Triumfetta, Erythroxylum, Tribulus,
Oxalis, Citrus, Suriana, Ochna, Malleastrum, Xylocarpus, Apodytes, Maytenus, Mystroxylon,
Colubrina, Gouania, Scutia, Allophylus, Dodonaea, Macphersonia, Operculicarya, Moringa,
Caesalpinia, Cassia, Delonix, Tamarindus, Calliandra, Dichrostachys, Abrus, Canavalia, Clitoria,
Crotalaria, Erythrina, Indigofera, Sophora, Tephrosia, Teramnus, Vigna, Brexia, Bruguiera,
Cassipourea, Ceriops, Rhizophora, Lumnitzera, Terminalia, Eugenia, Pemphis, Sonneratia, Turnera,
Passiflora, Carica, Cucumis, Cucurbita, Lagenaria, Momordica, Peponium, Trichosanthes, Sesuvium
Trianthema, Mollugo, Canthium, Guettarda, Hedyotis, Polysphaeria, Psychotria, Tarenna,
Triainolepis, Tricalysia, Bidens, Launaea, Melanthera, Synedrella, Tridax, Vernonia, Scaevola.
Plumbago, Sideroxylon, Jasminum, Azima, Salvadora, Carissa, Catharanthus, Pandaca, Pentopetia,
Pleurostelma, Sarcostemma, Pleurostelma, Tylophora, Cordia, Ehretia, Tournefortia, Evolvulus,
Ipomoea, Capsicum, Datura, Nicotiana, Solanum, Bacopa, Bryodes, Striga, Tabebuia, Asystasia,
Barleria, Hypoestes, Justicia, Ruellia, Avicennia, Clerodendrum, Conega, Lantana, Premna,
Stachytarpheta, Nesogenes, Leonotis, Ocimum, Boerhavia, Mirabilis, Pisonia, Achyranthes,
Alternanthera, Amaranthus, Deeringia, Lagrezia, Arthrocnemum, Cassytha, Hernandia, Bakerella,
Viscum, Acalypha, Euphorbia, Margaritaria, Pedilanthus, Phyllanthus, Ricinus, Laportea, Obetia,
Ficus, Maillardia, Casuarina, Acampe, Angraecum, Hederorkis, Dioscorea, Agave, Asparagus,
Dracaena, Lomatophyllum, Commelina, Cocos, Phoenix, Pandanus, Najas, Cymodocea, Halodule,
Ruppia, Syringodium, Thalassodendron, Halophila, Thalassia, Bulbostylis, Cyperus, Fimbristylis,
Bambusa, Cenchrus, Cymbopogon, Dactyloctenium, Daknopholis, Digitaria, Eleusine, Enteropogon,
Eragrostis, Eriochloa, Ischaemum, Lepturus, Panicum, Paspalum, Pennisetum, Sclerodactylon,
Sporobolus, Stenotaphrum, Zea.|

481 (559). Fosberg, F.R. and M.-H. Sachet. 1980. Systematic Studies of Micronesian Plants. Smithsonian
Contr. Bot. 45: 1-40. [Hymenophyllum, Trichomanes, Gleichenia. Thelypteris, Centosteca,
Rhaphidophora, Elatostema, Calliandra, Stylosanthes, Acalypha, Loeseneriella, Calophyllum,
Garcinia, Flacourtia, Citrullus, Zehneria, Bruguiera, Medinilla, Melastoma, Osmoxylon, Myrsine.
Pouteria, Fagraea, Geniostoma, Heliotropium, Callicarpa, Pseuderanthemum. Geophila, Hedvotis,
Psychotria, Spermacoce, Vernonia, Wollastonia.|

482 (565). and ——. 1980. Flora of Micronesia. 4. Caprifoliaceae-Compositae. Smithsonian
Contr. Bot. 46: 1-71. [Sambucus, Hippobroma, Scaevola, Adenostemma, Ageratum, Artemisia, Aster,
Bidens, Blumea, Chromolaena, Chrysanthemum, Cichorium, Conyza, Coreopsis, Cosmos,
Crassocephalum, Cynara, Eclipta, Elephantopus, Emilia, Epaltes, Erigeron, Gaillardia, Glossogyne,
Lactuca, Mikania, Pluchea, Pseudo-Elephantopus, Pseudogynoxys, Sonchus, Spilanthes, Synedrella,
Tagetes, Tithonia, Tridax, Vernonia, Wedelia, Wollastonia, Youngia, Zinnia.|

483 (566). Dassanayake, M.D. and F.R. Fosberg, eds. 1980. A Revised Handbook to the Flora of Ceylon.
Vol. I. Amerind Publishing Co. Pvt. Ltd., New Delhi, 508 pp.

R72 (562)a. Fosberg, F.R. 1980. Reviews of Heikoff, J.M., Coastal resources management: institutions
and programs, 1977, and Barnes, R.S.K., ed., The Coastline: a contribution to our understanding of
its ecology and physiography in relation to land-use and management and the pressures to which it is
subject, 1977. Environm. Conservation 7: 81-82.

R73 (563). ——. 1980. Island botany, review of Bramwell, D., ed., Plants and Islands, 1979. Nature 285:
680-681.
R74 (564). ——. 1980. Review of Bramwell, D., ed., Plants and Islands, 1979. Environm. Conservation

Ie Do,

28

D70 (568). [Fosberg, F.R.] 1980. Island ecosystems, a review. United Nations Environmental Program
Report No. 3, 1-34, Nairobi. [retyped, duplicated, and published from a manuscript review furnished
to UNEP by Fosberg in 1979.]

D71 (569). [Fosberg, F.R.] 1980. Coastal ecosystems, a review, United Nations Environmental Program
Report No. 4, 1-65, Nairobi. [retyped, duplicated, and published from a manuscript review furnished
to UNEP by Fosberg in 1979.]

1981

484 (573). Fosberg, F.R. 1981. The names published in Stickman's Herbarium Amboinense or Casuarina

litorea L., a valid Rumphian name. Taxon 30: 218-227.

485 (575). . 1981. Introduction, p. 1, in Raulerson, L. (ed.). Plant Biogeography of Guam. Univ. of
Guam, Marine Lab. Tech. Report. 69.

486 (586). ———. 1981. An overview of natural area preservation in the world, pp. 1-6. in Foster, J.B., ed.
Ecological Reserves. [British Columbia Provincial Museum] Heritage Record No. 12.

487 (590, 622). ——-. 1981 [1980.] A revision of the Flora of Ceylon (Sri Lanka). Spolia Zeylanica 35(1-
2): 9-14, 19.

488 (578). Fosberg, F.R. and J. Canfield. 1981 [1980.] Noteworthy Micronesian Plants 3. Micronesica
16: 189-200. [Tricomanes, Humata, Nephrolepis, Panicum, Eriocaulon, Aneilema, Achyranthes,
Piriqueta, Ammannia, Eugenia, Melaleuca, Leucas, Utricularia, Andrographis, Hedyotis,
Spermacoce, Timonius.|

489 (579). Fosberg, F.R. and M.V.C. Falanruw. 1981 [1980.] Noteworthy Micronesian Plants. 4.
Micronesica 16: 201-210. [Digitaria, Lemna, Achyranthes, Galphimia, Kolreuteria, Lepisanthes,
Eugenia, Allamanda.|

490 (580). Fosberg, F.R., M.V.C. Falanruw, and M.-H. Sachet. 1981 [1980.] Additional records of
vascular plants from the northern Marianas. 2. Micronesica. 16: 211-214. [Randia, Aidia, Setaria,
Hedyotis, Eupatorium, Chromolaena, Bidens, Cosmos, Wedelia, Wollastonia.|

491 (570). Fosberg, F.R. and M.-H. Sachet. 1981 [1980.] A New Sciaphila (Triuridaceae) from Palau
Islands. Pacific Sci. 34: 25-27.

492 (571). and ——. 1981 [1980.] Typification of the genus Ochrosia Jussieu, pp. 211-215, in
Gunawardana, T.T.P. et al, eds., P-E.P. Deraniyagala Commemoration Volume, Lake House
Investments, Ltd, Colombo, Sri Lanka.

493 (572). ——— and ——. 1981. Polynesian plant studies 6-18. Smithsonian Contr. Bot. 47: i-iv, 1-38.
6. Tricholaena Schrader (Poaceae) in the Pacific Islands, pp. 1-3.

7. Anew woody Oxalis L. (Oxalidaceae) from the Marquesas, pp. 3-5.

8. A new Claoxylon A. Jussieu (Euphorbiaceae) from the Marquesas, pp. 5-6.

9. Abutilon Miller (Malvaceae) in southeastern Polynesia and Samoa: The Abutilon sachetianum
Group, pp. 6-11.

10. New combinations in Zehneria Endlicher (Cucurbitaceae), pp. 11-13.

11. Terminalia L. (Combretaceae) in eastern Polynesia, pp. 13-17.

12. Further notes on Afyrsine L. (Myrsinaceae) in eastern Polynesia, p. 18.

13. New Geniostoma J. R. & G. Forster (Loganiaceae) taxa, pp. 18-21.

14. A new Rauvolfia L. (Apocynaceae) from the Marquesas, pp. 21-24.

15. Leucas R. Brown (Lamiaceae) in the Pacific Islands, pp. 24-26.

16. Observations on Cyrtandra J. R. & G. Forster (Gesneriaceae) in the Marquesas Islands, pp. 26-33.

17. Nomenclature of Dichrocephala integrifolia (L.f.) O. Kuntze (Asteraceae), pp. 33-35.

18. Miscellaneous taxonomic and distributional notes, pp. 35-37. [Vigna, Melicoccus, Triumfetta,
Miconia, Cordia.]

494 (583). and _ 1981. Clarification of Hygrophila triflora (Acanthaceae). Baileya 21: 145-
148.
495 (584). —— and ——. 1981. Pavonia (Malvaceae) in Society Islands. Bull. Mus. Natl. Nat. Hist.. B,

Adansonia 1: 15-18.

29

496 (585). and ——. 1981. Nomenclatural notes on Micronesian Ferns. Amer. Fern J. 71: 82-84.
[Thelypteris, Trichomanes.]

497 (576). Dassanayake, M.D. and F.R. Fosberg, eds. 1981. A Revised Handbook to the Flora of Ceylon.
Vol. II. Amerind Publishing Co. Pvt. Ltd., New Delhi, 511 pp.

498 (587). —— and ——., eds. 1981. A Revised Handbook to the Flora of Ceylon. Vol. III. Amerind
Publishing Co. Pvt. Ltd., New Delhi, 499 pp.

499 (577). Macnae, W. and F.R. Fosberg. 1981. Rhizophoraceae, pp. 487-500, in Dassanayake, M.D. and
F.R. Fosberg, eds., A Revised Handbook to the Flora of Ceylon. Vol. Il. Amerind Publishing Co.
Pvt. Ltd., New Delhi, 511 pp. [Anisophyllea, Bruguiera, Ceriops, Kandelia, Rhizophora.|

500 (588). and . 1981. Lecythidaceae, pp. 195-201, in Dassanayake, M.D. and F.R. Fosberg,
eds., A Revised Handbook to the Flora of Ceylon. Vol. III. Amerind Publishing Co. Pvt. Ltd., New
Delhi, 499 pp. [Barringtonia, Careya, Couroupita.|

501 (589). —— and . 1981. Sonneratiaceae, pp. 450-453, in Dassanayake, M.D. and F.R. Fosberg,
eds., A Revised Handbook to the Flora of Ceylon. Vol. III. Amerind Publishing Co. Pvt. Ltd., New
Delhi, 499 pp. [Sonneratia.]

502 (595). Stoddart, D.R. and F.R. Fosberg. 1981. Bird and Denis Island, Seychelles. Atoll Res. Bull.
252: 1-55.

503 (596). —— and ——. 1981. Topographic and floristic changes, Dry Tortugas Florida, 1904-1977.
Atoll Res. Bull. 253: 1-76.

R75 (581). Fosberg, F.R. 1981 [1980]. Review of van Steenis, C.G.G.J, ed., Flora Malesiana, series 1, vol.
8. Micronesica 16: 365-366.

R76 (582). ——. 1981 [1980.] Review of Vogel, E.F. de, Seedlings of Dicotyledons; structure,
development, types, descriptions of 150 woody Malesian taxa. Micronesica 16: 366-367.

1982

504 (594). Fosberg, F.R. 1982. A preliminary conspectus of the genus Leptostigma (Rubiaceae). Acta
Phytotax. Geobot. 33: 73-83.

505 (599). Fosberg, F.R. and M.-H. Sachet. 1982. Notes on Sphenomeris chinensis (L.) Maxon ex
Kramer. Micronesica 18(1): 129-133.

506 (598). Fosberg, F.R., M.-H. Sachet, and R.L. Oliver. 1982. Geographical checklist of Micronesian
Pteridophyta and Gymnospermae. Micronesica 18(1): 23-82.

507 (593). Fosberg, F.R., D.R. Stoddart, M.-H. Sachet, and D.L. Spellman. 1982. Plants of the Belize
cays. Atoll Res. Bull. 258: 1-77.

508 (597). Stoddart, D.R. and F.R. Fosberg. 1982. Species-area relationships on small islands: floristic
data from Belizean sand cays, pp. 527-539, in Riitzler, K. and I.G. MacIntyre, eds., The Atlantic
Barrier Reef Ecosystem at Carrie Bow Cay, Belize, 1: Structure and Communities. Smithsonian
Contr. Mar. Sci. 12: 1-539.

509 (592). Stoddart, D.R., F.R. Fosberg, and M.-H. Sachet. 1982. Ten years of change on the Glover's
Reef cays. Atoll Res. Bull. 257: 1-17 + 24 pp. figs. and plates.

510 (591). Stoddart, D.R., F.R. Fosberg, and D.L. Spellman. 1982. Cays of the Belize barrier reef and
lagoon. Atoll Res. Bull. 256: 1-76.

1983

511 (602). Fosberg, F.R. 1983. The human factor in the biogeography of oceanic islands. Comt. Rend.

Séances Soc. Biogéogr. 59: 147-190.

512 (606). ——. 1983. Two new species of Psychotria from the Society Islands. Candollea 38: 455-458.

513 (607). ——. 1983. A possible new pathogen affecting Metrosideros in Hawaii. Newslett. Hawaiian
Bot. Soc. 22: 13-16.

514 (614). ——. 1983. Natural history of Cousin Island. Atoll Res. Bull. 273: 7-38.

515 (605). Fosberg, F.R. and M.-H. Sachet. 1983. Henderson Island threatened. Environm. Conservation
10: 171-173.

516 (61 1a). and ——. 1983. Plants of the Society Islands, pp. 76-107, in Carr, D.J., ed., Sydney

Parkinson, artist of Cook's Endeavour voyage. University of Hawaii Press, Honolulu, xvi + 300 pp.

30

517 (608). Fosberg, F.R., M.-H. Sachet, and D.R. Stoddart. 1983. Henderson Island (southeastern
Polynesia): summary of current knowledge. Atoll Res. Bull. 272: 1-47, pl. 1-12.

518 (615). ——, ——, and ——. 1983. List of the recorded vascular flora of Agalega. in Sachet, M.-H.,
D.R. Stoddart. and F.R. Fosberg, eds., Floristics and ecology of western Indian Ocean islands. Atoll
Res. Bull. 273: 109-142.

519 (601). Dassanayake, M.D. and F.R. Fosberg. eds. 1983. A Revised Handbook to the Flora of Ceylon.
Vol. [V. Amerind Publishing Co. Pvt. Ltd., New Delhi, 532 pp.

520 (600). Roane, M.K. and F.R. Fosberg. 1983. A new pyrenomycete associated with \/etrosideros
collina subspecies polymorpha (Myrtaceae). Mycologia 75: 163-166.

521 (619). Robertson, I.A.D., S.A. Robertson, and F.R. Fosberg. 1983. List of plants collected on
Alphonse Island, Amirantes. Atoll Res. Bull. 273: 177-184.

522 (616). Robertson, S.A. and F.R. Fosberg. 1983. List of plants collected on Coetivy Island, Seychelles.
Atoll Res. Bull. 273: 143-156.

523 (617). and . 1983. List of plants collected on Platte Island, Seychelles . Atoll Res. Bull.
273: 157-164.

524 (618). —— and ——. 1983. List of plants of Poivre Island. Amirantes. Atoll Res. Bull. 273: 165-
176.

525 (611). Sachet, M.-H. and F.R. Fosberg. 1983. An ecological reconnaissance of Tetiaroa Atoll, Society
Islands. Atoll Res. Bull. 275, 1-67, pl. 1-42.

526 (610a). Sachet, M.-H., D.R. Stoddart, and F.R. Fosberg, eds. 1983. Floristics and ecology of western
Indian Ocean islands. Atoll Res. Bull. 273, 1-253.

R77 (603). Fosberg, F.R. 1983. Review of Mueller-Dombois, D.. K.W. Bridges. and H.L. Carson, eds.,
Island Ecosystems: Biological Organization in Selected Hawaiian Communities, 1981. Environm.
Conservation 10: 82-33.

R78 (604). ——. 1983. Review of Hayward, S.J.. V-H. Gomez, and W. Sterrer, eds., Bermuda's Delicate
Balance--People and the Environment, 1981. Environm. Conservation 10: 179.

R79 (609a). ——. 1983. Review of Long, S.P. and C.F. Mason, Saltmarsh Ecology, 1983. Environm.
Conservation 10: 369.

R80. ——. 1983. Review of Smith, A.C., Flora Vitiensis Nova, vol. 2, 1981. Taxon 32: 156.

D72 (608a). ——. 1983. A possible new pathogen affecting \/etrosideros in Hawaii, [abstract.] Pacific
Science Association, 15th Congress, Abstracts. p. 75.

D73 (608b). Stoddart, D.R., C.D. Woodroffe, and F.R. Fosberg. 1983. Substrate-specific species
distributions on Suwarrow Atoll, Central Pacific: implications for island biogeography theory,
{abstract.] Pacific Science Association, 15th Congress, Abstracts, pp. 227-228.

1984

527 (621). Fosberg, F.R. 1984. Henderson Island saved. Environm. Conservation 11: 183-184.

528 (624). ——. 1984. Phytogeographic comparison of Polynesia and Micronesia, pp. 33-44, in
Radovsky, F.J., P-H. Raven, and S.H. Sohmer, eds., Biogeography of the Tropical Pacific, Proceedings
of a Symposium. Special Publ. Bernice Pauahi Bishop Mus., 72. x + 221 pp.

529 (629). Fosberg. F.R. and D. Herbst. 1984 [1983.] A Nesogenes (Chloanthaceae) from Micronesia.
Micronesica 19: 11-15.

530 (609, 610). Fosberg, F.R. and M.-H. Sachet. 1984. Identity and typification of Ruellia triflora Roxb.
and R. difformis L.f. (Acanthaceae). Baileya: 22: 138-140.

531 (612). and . 1984 [1982.] Micronesian Poaceae: critical and distributional notes.
Micronesica 18(2): 45-102. [Cynodon, Digitaria, Echinochloa, Imperata, Isachne, Ischaemum,
Lepturopetium, Oplismenus, Paspalum, Pennisetum, Pogonatherum, Polytrias, Saccharum, Sorghum.]

532 (613). and ——. 1984. [1982 | The identification of Desmodium purpureum (Roxburgh) Hooker
and Arnott. Micronesica 18(2): 195-196.

533 (620). Fosberg. F.R. and D.R. Stoddart. 1984. Vegetation and floristics of western Indian Ocean coral
islands, pp. 221-238, in Stoddart, D. R., ed., Biogeography and Ecology of the Seychelles Islands. Dr
W. Junk Publishers, The Hague.

en

31

R81 (636). Fosberg, F.R. 1984. Review of: Synge, H. The IUCN/WWF Plants Conservation Programme

1984-1985. Environm. Conservation 11: 285-286.
1985

534 (625). Fosberg, F.R. 1985. Cordyline fruticosa (L.) Chevalier (Agavaceae). Baileya 22: 180-181.

535 (628). . 1985. Origin and early history of Flora Neotropica, pp. 25-27, in Cuatrecasas, J., Flora
Neotropica Monograph number 2 Supplement, Brunelliaceae.

536 (632). . 1985. Range extension of Diaporthiopsis meterosideri to Bird Park, Hawaii Volcanoes
National Park. Newslett. Hawaiian Bot. Soc. 24: 18.

537 (637). . 1985. Coral reef destruction in Ryukyu Islands. Environm. Conservation 12: 270.

538 (638). ——. 1985 [1986?]. Present state of knowledge of the flora and vegetation of emergent reef
surfaces. Proceedings of the Fifth International Coral Reef Congress, Tahiti, 5: 107-112.

539 (678). ——. 1985. Classification of emergent reef surfaces. Atoll Res. Bull. 292: 29-36, .

540 (679). ——. 1985. Botanical visits to Krakatau in 1958 and 1963. Atoll Res. Bull. 292: 39-48.

541 (625a). Fosberg, F.R. and M.-H. Sachet. 1985. History of the generic name Pseuderanthemum
Radlkofer (Acanthaceae). Baileya 22: 178-179.

542 (633). and . 1985. Rare, endangered, and extinct Society Island plants. Natl. Geogr. Soc.
Res. Report 21: 161-165. 2

543 (634). —— and ——. 1985. Santalum in eastern Polynesia. Candollea 40: 459-470.

544 (635). —— and ——. 1985 [1982.] A new Pavonia from Ceylon. Ceylon J. Sci., Biol. Sci. 15: 38-
40.

545 (623). Fosberg, F.R. and E.E. Terrell. 1985. A recently established exotic in west Florida and
Alabama (Hedyotis salzmannii or Oldenlandia salzmannii Rubiaceae). Castanea 50: 49-51.

546 (638a). Dassanayake, M.D. and F.R. Fosberg, eds. 1985. A Revised Handbook to the Flora of Ceylon.
Vol. V. Amerind Publishing Co. Pvt. Ltd., New Delhi, 476 pp.

547 (627). Lourteig, A. and F.R. Fosberg. 1985. Application of the name Calophyllum calaba L.,
Guttiferae (Clusiaceae). Phytologia 57: 153-55.

R82 (630). Fosberg, F.R. 1985 [1983.] Review of Hezel, F.X., The First Taint of Civilization: A History of
the Caroline and Marshall Islands in Pre-Colonial Days, 1983. Micronesica 19: 180-181.

*R83 (631). . 1985. Review of Cuzent, G., revised and expanded by Florence, J., M. Guerin, F.
Margueron, D. Margueron, D. Koenig, and R. Koenig, Archipel de Tahiti: Recherches sur les
principales productions vegetales, 1983. Pacific Stud. 8: 125-129.

1986
548 (639). Fosberg, F.R. 1986. Biodiversity. Environm. Aware. 9: 125-129.
549 (639a). ——. 1986. (264) Proposal to abolish conservation of specific names. Taxon 35: 847.
550. ——. 1986. (265) Proposal regarding autonyms. Taxon 35: 847-849.
551. ——. 1986. (266)-(268) Proposals relating to the rejection of names under Article 69. Taxon 35:
849-850.
552. ——. 1986. (269)-(274) Proposals to rationalize and make consistent and understandable Art. 73.8
and Rec. 73g.1 on connecting forms in epithets made up of two elements. Taxon 35: 850-851.
553. ——. 1986. Marie-Hélene Sachet (1922-1986). Taxon 35: 936.
D74 (707). ——. 1986. Vegetation of Bikini Atoll, 1985. Report prepared for Bikini Atoll Rehabilitation
Committee, 39 pages, duplicated.
1987
554 (640). Fosberg, F.R. 1987. The genus 7rukia Kanehira (Rubiaceae). Phytologia 62: 171-176.
555 (641). ——. 1987. A new Pisonia (Nyctaginaceae) from the Hawaiian Islands. Phytologia 62: 177-
178.
556 (642). ——. 1987. Bobea platypes Fosberg, new species (Rubiaceae) from Maui, Hawaiian Islands.
Phytologia 62: 179-180.
557 (643). ——. 1987. New nomenclatural combinations for Galapagos plant species. Phytologia 62:

181-183. [Solanum, Spermacoce.|
558 (646). ——. 1987. Artificial Diversity. Environm. Conservation 14: 74.

32

559 (647). ——. 1987. A living museum of Ceylon trees. Environm. Conservation 14: 79.

560 (648). ——. 1987. Papala Képau, Pisonia umbellifera (Forster) Seeman. Newslett. Hawaiian Bot.
Soc. 26(2): 23-25.

561 (648b). ——. 1987. Geophila (Rubiaceae), pp. 320-321, in Dassanayake, M.D. and F.R. Fosberg, eds.
A Revised Handbook to the Flora of Ceylon. Vol. VI. Amerind Publishing Co. Pvt. Ltd., New Delhi,
424 pp.

562 (650). ——. 1987. Marie Helene Sachet: islands, atolls and reefs, in Introduction to Marie-Héléne

Sachet Commemorative Issue. Atoll Res. Bull. 293: 1-7.

563 (651). Fosberg, F.R. and M.-H. Sachet. 1987. Flora of Maupiti, Society Islands. Atoll Res. Bull. 294:
1-70.

564 (652). —— and ——. 1987. Flora of Gilbert Islands, Kiribati, checklist. Atoll Res. Bull. 295: 1-33.

565 (648a). Dassanayake, M.D. and F.R. Fosberg, eds. 1987. A Revised Handbook to the Flora of Ceylon.
Vol. VI. Amerind Publishing Co. Pvt. Ltd., New Delhi, 424 pp.

*D75 (644). Fosberg, F.R. 1987. Undisturbed vegetation on Fernandina. | p. abstract distributed at
workshop on vegetation research and management, Charles Darwin Research Station, 12 April, Santa
Cruz Island, Galapagos.

*D76 (645). ——. 1987. Revegetation of Bikini Atoll, 1987. 1 p. abstract distributed at workshop on
vegetation research and management, Charles Darwin Research Station, 12 April, Santa Cruz Island,
Galapagos.

D77 (653). ——. 1987. Workshop on botanical research and management in Galapagos, held at the

Charles Darwin Research Station, Santa Cruz Island, Galapagos, Ecuador. during 13-19 April 1987.
Environm. Conservation 14: 277-278.

1988

566 (654). Fosberg, F.R. 1988. New and noteworthy plants from Great Barner Reef Sand Cays, Australia.
Brittonia 40: 52-65. [Lepturus, Boerhavia, Euphorbia, Spermacoce, Diospyros, Abutilon.]|

567 (649, 655). ——. 1988. Native Hawaiian flora surveyed. Naturalist Review, Winter,. p. 3.
568 (656). ——. 1988. Fimbristylis juncea (Forst. f.) R. and S. and its relatives. J. Jap. Bot. 63: 78-85.
569 (656a). ——. 1988. The vegetation of Bikini Atoll, 1985. Atoll Res. Bull. 315: 1-28+ 5S figs.

570 (657). Fosberg, F.R. and M.-H. Sachet. 1988 [1987.] A geographic checklist of the Micronesian
Monocotyledonae. Micronesica 20: 19-129.

571 (658). —— and . 1988 [1987.] Noteworthy Micronesian plants 5. Micronesica 20: 131-156.
[Crinum, Dianella, Dioscorea, Curcuma, Didymoplexis, Liparis, Habenaria, Peristylus, Microstylis,
Malaxis, Nervilia, Spathoglottis, Rhynchophreatia, Taeniophyllum, Trachoma.|

572 (659). and ——. 1988 [1987.] The genus 7imonius (Rubiaceae) in the Palau Islands.
Micronesica 20: 157-164.
573 (660). and ——. 1988 [1987.] Notes on Micronesian Cyperaceae. Micronesica 20: 165-170.

[Cyperus, Fimbristylis, Machaerina.|

574 (656b). [Fosberg, F.R., I.G. MacIntyre, K. Riitzler, and L.L. Cole, eds.] 1988. Island news and
comments. Atoll Res. Bull. 320: 1-12.

*L20 (656c). Fosberg, F.R. 1988. Priorities in Science, [letter to the editor.] The Washington Post, 14

1989

May.

*D78 (656d). ——. 1988. A review of the natural history of the Marshall Islands. Prepared for East-West
Center, MacArthur Foundation Project.

575 (662). Fosberg. F.R. 1989. Interest and beauty in the world. Environm. Conservation 16: 4.

576 (663). ——. 1989. Society Island plants 1. Typha L. (Typhaceae). Bot. Bull. Acad. Sin. 30: 219-221.

577 (666). ——. 1989. Garbage dump in the Marshall Islands? Environm. Conservation 16: 78-79.

578 (668). ——. 1989. Nuclear garbage proposal for the Marshall Islands. Environm. Conservation 16:
DB.

579 (669a). ——. 1989. A skeptic sees a miracle. Environm. Conservation 16: 353-354.

580 (669b). ——. 1989. Success against shifting sands in Uzbekistan. Environm. Conservation 16: 371.

33

*581 (697). ——. 1989. Foreword, pp. [X-X, in Campbell, D.G. and H.D. Hammond, eds., Floristic
Inventory of Tropical Countries, New York Botanical Garden.

582 (669c). ——. 1989. Henderson Island: dedicated to S. Dillon Ripley [dedication of issue.] Atoll Res.
Bull. 321: 3 unnumbered pages.

583 (669e). ——. 1989. [Tribute to] Truman G. Yuncker. Brittonia 41: 248-249.

584 (661). [Fosberg, F.R.] 1989. In the beginning .... Univ. S. Pacific Bull. 22(15): 1-3. [Condensation of
a lecture. ]

585 (664). Fosberg, F.R. and M.-H. Sachet. 1989. Three cultivated /xoras (Rubiaceae). Baileya 23: 74-
85.

586 (665). —— and ——. 1989. Lectotypification of /xora coccinea L. (Rubiaceae). Taxon 38: 486-489.

587 (669d). Fosberg, F.R., G. Paulay, T. Spencer, and R. Oliver. 1989. New collections and notes on the
plants of Henderson, Pitcairn, Oeno, and Ducie Islands. Atoll Res. Bull. 329: 1-18.

1990

588 (669). Fosberg, F.R. 1990. Endangered species? Why worry? Environm. Aware. 13: 69-71.

589 (671). . 1990. New and noteworthy Cook Island plants. Micronesica 23: 1-4. [Polypodium,
Terminalia, Nesogenes, Psychotria, Scaevola.|

590 (676). . 1990. Nyctaginaceae, pp. 976-988, in Wagner, W.L. et al. Manual of the Flowering
Plants of Hawaii. [Boerhavia, Mirabilis, Pisonia.|

591 (690). ——. 1990. Undisturbed vegetation on Fernandina. 1964. in Lawesson, J.E., O. Hamann, G.
Rogers, G. Reck, and H. Ochoa, eds., Botanical Research and Management in Galapagos, Mongr.
Syst. Bot. Missouri Bot. Gard. 32: 59-65.

592 (691). . 1990. Revegetation on Bikini Atoll, 1985, in Lawesson, J.E.,O. Hamann, G. Rogers, G.
Reck, and H. Ochoa, eds., Botanical Research and Management in Galapagos, Mongr. Syst. Bot.
Missouri Bot. Gard. 32: 265-269.

*593 (69la). ——. 1990. Do we want a monotonous world? Environm. Aware. 13: 55-57.
594 (698a). ——. 1990. A review of the natural history of the Marshall Islands. Atoll Res. Bull. 330: 1-
100.

595 (677). Fosberg, F.R. and L. Raulerson. 1990. Noteworthy Micronesian plants. 6. Micronesica 23:
139-152. [Eragrostis, Cyperus, Alpinia, Hernandia, Oxalis, Bombax, Melochia, Coccinia, Melodinus,
Cerbera, Paederia, Psychotria.|

*R84 (668a). Fosberg, F.R. 1990. Review of Wallace, A.R. The Malay Archipelago: the land of the
orang-utan, and the bird of paradise, new edition, 1990. Science Books and Films 26: 118-119.

D79 (667). . 1990. The vegetation of the Society Islands, [abstract.] Fifth International Congress of
Ecology, Yokohama, Japan, Aug. 27-30, 1990, p. 141.

*D80 (673). ——. 1990. Commentary on Westing's keynote address. Duplicated (and corrected) handout
distributed at 4th International Conference on Environmental Future, Budapest, April 26.

1991

596 (683). Fosberg, F.R. 1991. Casuarina in Miscanthus savanna on Guam. Micronesica 24: 163.

597 (685). ——. 1991. After zero-growth? Environm. Conservation 18: 3-4.

598 (689). ——. 1991. A Code or a bureaucracy? pp. 145-149, in Hawkesworth, D.L., ed., Improving the
Stability of Names: Needs and Options. Regnum Veg. 123.

599 (694). ——. 1991. Hawaii conservation biology initiative. Environm. Conservation 18: 263-264.

*600 (696). ——. 1991. Pacific oceanic island biodiversity in a geohistoric perspective [Appendix D,
Plenary address.] Proc. XVII Pacific Sci. Congress 71-73. [(715.) republished 1992. Atoll Res. Bull.
355: 13-17.]

601 (702). ——. 1991. Opening address, p. 1, in Chase, R. and R.R. Thaman, eds., Proceedings of the
International Conference on Developing Agricultural Research Programs for Atolls.

602 (705). ——. 1991. The atoll environment [condensation of a lecture,] p. 16, in Chase, R. and R.R.

Thaman, eds., Proceedings of the International Conference on Developing Agricultural Research
Programs for Atolls.

603 (681). Fosberg, F.R. and R.L. Oliver. 1991. C.L. Ledermann's collection of flowering plants from the
Caroline Islands. Willdenowia 20: 257-314. :
1. Fosberg, F.R. Introduction, pp. 257-259.
Dh . Miscellaneous notes, pp. 259-267. [Alternanthera, Bridelia, Celtis, Cerbera, Clitandropsis,
Glochidion, Gmelina, Homalanthus, Sterculia.|
3. Fosberg, F.R. and R.L. Oliver. Numerical list of Ledermann Micronesian collections, pp. 267-299.
[lectotypes designated in taxa of: Amaracarpus, Andruris, Badusa, Bikkia, Borreria,
Buchanania, Bulbophyllum, Canthium, Cerbera, Clitandropsis, Coffea, Decaspermum,
Dendrobium, Diplocaulobium, Discocalyx, Elaeocarpus, Euodia, Fagraea, Fimbristylis,
Garcinia, Geniostoma, Gmelina, Habenaria, Hedyotis, Hetaeria, Hippocratea, Ilex,
Loeseneriella, Maesa, Malaxis, Melicope, Microstylis, Moerenhoutia, Morinda, Myrsine,
Oldenlandia, Phreatia, Plectronia, Premna, Pseuderia, Psychotria, Rapanea, Rhynchophreatia,
Salacia, Salacicrates, Spermacoce, Stemonurus, Svmplocos, Syzygium, Taeniophyllum,
Timonius, Trichoglottis, Urandra, Vandopsis.|
4. Fosberg, F.R. and R.L. Oliver. Missing Ledermann types, pp. 299-301.
5. Ledermann species listed by family, pp. 301-314.
604 (698). Fosberg, F.R. and M.-H. Sachet. 1991. Studies in Indo-Pacific Rubiaceae. Allertonia 6: 191-
278.
1. Fosberg, F. R. Introduction. 191-193.

2. ——. Introductory remarks on Hedyotis L. sensu lato. 193-195.

3. Fosberg, F. R. and M.-H. Sachet. Disposition of Leptopetalum Hook. & Arnott in Hedyotis L.
195-205.

4. Fosberg, F. R. and M.-H. Sachet. Variation in Hedyotis auriculata L. and H. verticillata Lam.
205-212.

5. Fosberg, F. R. Hedyotis L. in Micronesia. 212-244.

6. ——. The Micronesian species of Psychotria L. 244-268.

7. Fosberg, F. R. and M.-H. Sachet. Variations in Zarenna sambucina. 268-274.

605 (686). Fosberg, F.R. and D.R. Stoddart. 1991. Plants of the reef islands of the northern Great Barrier
Reef. Atoll Res. Bull. 348: 1-82.

606 (700a). Dassanayake, M.D. and F.R. Fosberg, eds. 1991. A Revised Handbook to the Flora of Ceylon.
Vol. VII. Amerind Publishing Co. Pvt. Ltd., New Delhi, 439 pp.

607 (682). Rinehart, A.F. and F.R. Fosberg. 1991. Nervilia jacksoniae nov. sp., a new species of
Orchidaceae from Guam and Rota. Micronesica 24: 81-85.

608 (687). Stoddart. D.R. and F.R. Fosberg. 1991. Phytogeography and vegetation of the reef islands of
the northern Great Barrier Reef. Atoll Res. Bull. 349: 1-19.

609 (688). —— and . 1991. Plants of the Jamaican Cays. Atoll Res. Bull. 352: 1-24.

R85 (684). Fosberg. F.R. 1991. Ravages of Man, review of Cuddihy, L.W. and C.P. Stone, Alteration of
Native Hawaiian Vegetation. Effects of Humans, Their Activities and Introductions, 1990.
Micronesica 24: 167-168.

R86 (692). ——. 1991. Review of Wagner, D.R. Herbst, and $.H. Sohmer, Manual of the Flowering
Plants of Hawaii, 1990. Environm. Conservation 18: 191.
R87 (695). ——. 1991. Review of Mitchell, A., The Fragile South Pacific, an Ecological Odessey, 1991.

Environm. Conservation 18: 285.

R88 (708). . 1991. Review of Schultes, R-E. and R.F. Raffauf, The healing forest: Medicinal and
toxic plants of the northwest Amazonia, 1990. Taxon 40: 157-158.

*D81 (674). ——. 1991. A Code or a bureaucracy? Duplicated and distributed at Kew meeting on
Improving the Stability of Names, Needs, and Options, February 20.

*D82 (680). . 1991. Captain Cook's Voyages and Pacific Science, [abstract.] Toward the Pacific
Century Symposium, Abstracts, p. 39.

35

D83 (675). Fosberg, F.R. and S. Pearsall. 1991. Classification of ecosystems. Background paper
duplicated and distributed by Nature Conservancy and SPRED at Ecosystem Classification Workshop,
East-West Center, Honolulu, March 11-15, 1991, 41 pp.

*D84 (700). [Fosberg, F.R. with thirty other workshop members.] 1991. An ecosystem classification and
criterion for conservation in the tropical insular Pacific. Honolulu, duplicated [by the Nature
Conservancy?], 81 pp.

1992
610 (699). Fosberg, F.R. 1992. Vegetation of the Society Islands. Pacific Science 46: 232-250.

*611 (701). . 1992. Sustainability. Environm. Aware. 15: 15-16.

612 (703). ——. 1992. An essay on lectotypification. Taxon 41: 321-323.

613. ——. 1992. (82-87) Six proposals to amend the Code. Taxon 41: 354-355.

614 (706). ——. 1992. The Coral island environment and its relevance to Atoll agriculture, pp. 45-54, in

Chase, R.G., Review of Agricultural Development in the Atolls. University of the South Pacific, Apia,
Western Samoa, 1992.

615 (713). . 1992. Pacific Island plants—taxonomic and distributional notes. Micronesica 25: 175-
199. [Polypodium, Lemaphyllum, Nephrolepis, Spathoglottis, Vanilla, Peperomia, Lepidium,
Erythrina, Tephrosia, Euphorbia, Phyllanthus, Passiflora, Luffa, Pouteria, Chionanthus, Ochrosia.]

R89 (670, 709). . 1992. Review of Beehler, B., A Naturalist in New Guinea, 1991. Environm.
Conservation 19: 190, 1992.

*D85 (704). ——. 1992. Plant evolution in the Pacific region, [abstract.] AIBS meeting, Honolulu,
August, 1992.
D86 (710). ——. 1992. Phytogeographic observations in Okinawa and the southern Ryukyu Islands,

{abstract.] International symposium on Okinawan Studies Abstract, Natural Science Session, pp. 7-8.
[Also published p. 105 of symposium volume. }
1993

616 (716). Fosberg, F.R. 1993. The Forster Pacific Islands collections from Captain Cook's Resolution
Voyage. Allertonia 7: 41-86. [precursor account of a project to locate and interpret with examples
collections by the Forsters; 7mesipteris, Lycopodium, Polypodium, Arachniodes, Agrostis, Paspalum,
Miscanthus, Dianella, Peperomia, Korthalsella, Balanophora, Ximenia, Acacia, Sesbania, Euphorbia,
Allophylus, Grewia, Waltheria, Xylosma, Wikstroemia, Rhizophora, Terminalia, Alyxia, Melodinus,
Cyrtandra, Dicliptera, Guettarda, Ixora, Wollastonia.]

617 (711). Fosberg, F.R. and S.H. Pearsall III. 1993. Classification of non-marine ecosystems. Atoll Res.
Bull. 389: 1-38.

618 (712). Fosberg, F.R., M.-H. Sachet and R.L. Oliver. 1993. Flora of Micronesia, 5: Bignoniaceae-
Rubiaceae. Smithsonian Contr. Bot. 81: i-iv, 1-135. [Catalpa, Clytostoma, Crescentia, Cydista,
Dolichandrone, Jacaranda, Kigelia, Macfadyena, Mansoa, Parmentiera, Pyrostegia, Saritaea,
Spathodea, Tabebuia, Tecoma, Tecomaria, Sesamum, Aeschynanthus, Cyrtandra, Episcia,
Saintpaulia, Utricularia, Acanthus, Andrographis, Aphelandra, Asystasia, Barleria, Blechum,
Crossandra, Eranthemum, Fittonia, Graptophyllum, Hemigraphis, Hygrophila, Justicia, Odontonema,
Pseuderanthemum, Ruellia, Thunbergia, Myoporum, Plantago, Aidia, Badusa, Bikkia, Canthium,
Cephaelis, Cinchona, Coffea, Dentella, Gardenia, Geophila, Guettarda, Gynochthodes, Hedyotis,
Hydnophytum, Ixora, Mitracarpus, Morinda, Mussaenda, Mycetia, Ophiorrhiza, Paederia, Pentas,
Psychotria, Scyphiphora, Spermacoce, Tarenna, Timonius, Trukia, Uncaria.]

Publications in press:

619. Thaman, R.R., F.R. Fosberg, H.I. Manner, and D.C. Hassall. In press. Flora of Nauru. Atoll Res.
Bull. 392. [in this issue. ]

620. Fosberg, F.R. and D.R. Stoddart. In press. Flora of Phoenix Islands, Central Pacific. Atoll Res. Bull.
393. [in this issue. ]

621. Stoddart, D.R. and F.R. Fosberg. In press. The Hoa of Hull Atoll and the problem of Hoa. Atoll Res.
Bull. 394. [in this issue. ]

622. Fosberg, F.R. In press. Story of an oceanic archipelago. Atoll Res. Bull. 395. [in this issue. ]

36
623. ——. Inpress. Comments on Atoll Phosphate rock. Atoll Res. Bull. 396. [in this issue. ]
624. ——. Inpress. /xora casei vs. Ixora longifolia (Rubiaceae). Baileya 23(4). [expected 1994.]
625. ——. Inpress. A new species of Scleria (Cyperaceae) from New Caledonia. Adansonia. [expected

1994]
Submitted publication:
Fosberg, F.R. Biodiversity in India, 1993. Herbaria, plant diversity, and conservation—new challenges
ahead. Botanical Survey of India.

SOME PUBLISHED PHOTOGRAPHS, DEDICATIONS, AND BIOGRAPHICAL
INFORMATION

Gregory, H.E. et al. 1935. Mangarevan Expedition. Bernice P. Bishop Mus. Bull. 133: 33-71.
[Photograph and information about F.R. Fosberg. |

Fosberg, F.R. 1961. Pacific vegetation progress report (1956-1958). Proc. Symposium on Humid Tropics
Vegetation, UNESCO, Tjiawe, Indonesia, 1958, 32-33. [Includes biographical sketch of F.R. Fosberg.]

1971. Bull. Pacific Tropical Bot. Gard. 1(3): 8. [Photograph of F.R. Fosberg with other famous tropical
botanists. |

Harney, T. 1978. Spreading the News on Atolls. The Smithsonian Torch 78-10: 6. [Description of Atoll
Research Bulletin, comments about and photograph of F.R. Fosberg. |

Mason, C.N. 1980. Browning Award to Dr. Fosberg. Environm. Conservation 7: 124. [Appreciation of
F.R. Fosberg’s work and as recipient of 1979 Edward W. Browning medal for distinguished
international services in the cause of environmental conservation. |

[Fosberg, F.R.] 1989. In the beginning .... Univ. S. Pacific Bull. 22(15): 1-3. [condensation of a lecture,
accompanied by a photograph of Dr. Fosberg receiving an Honorary degree from the Vice-Chancellor. ]

Meurer-Grimes, B. 1989. Itinerary of Truman G. Yuncker’s expeditions. Fig. 1, Brittonia 41: 226.
[Photograph of Erling Christopersen, E. Y. Hosaka, F. Raymond Fosberg, Harold St. John, and
Truman G. Yuncker in Oahu, 1932.]

Stoddart, D.R. 1992. F. Raymond Fosberg and the Atoll Research Bulletin 1951-1991. Atoll Res. Bull.
355: 1-6. [Photographs of F.R. Fosberg and others on pages 8 & 12.]

Anonymous. 1993. News and Comments, 40th Anniversary Reception. Atoll Res. Bull. following 389, 2
unnumbered pages (30-40). [Photographs with F.R. Fosberg and appreciation of his work. }

Mueller-Dombois, D. 1992. F. Raymond Fosberg: An Appreciation. Pacific Science 46: 107-110.

van Balgooy, M.M.J., ed. 1993. Pacific plant areas, volume 5. Rijksherbarium/Hortus Botanicus, Leiden.
[Final volume dedicated to F.R. Fosberg. ]

Nicolson, D.H. In press. Lists of taxa named for F. Raymond Fosberg and by him. Atoll Res. Bull. 390.
[in this issue. ]

In addition, several newspapers carried obituaries of F. Raymond Fosberg, including the Washington
Post and London Independent. Obituaries will also appear in scientific journals including Zaxon. The
National Tropical Botanical Garden intends to dedicate an upcoming issue of its Bulletin to F.R. Fosberg.
Undoubtedly other scientific journals will include obituaries and other dedications will appear in the coming
months.

A

Abies, 53-4

Abronia, 97-6

Abrus, 480-27

Abutilon, 328-18, 355-20, 480-27,
493-28, 566-32

Acacia, 616-35

Acalypha, 39-4, 418-23, 480-27,
481-27

Acampe, 480-27

Acanthus, 39-4, 618-35

Acer, 57-4, 151-9

Achyranthes, 480-27, 488-28,
489-28

Acrostichum, 382-21, 480-27

Adenostemma, 482-27

Adiantum, 355-20, 438-24

Aeschynanthus, 618-35

Agave, 480-27

Ageratum, 74-5, 482-27

Aglaia, 198-11

Agrostis, 616-35

Aidia, 490-28, 618-35

Albizia, 299-17

Allamanda, 422-23, 489-28

Allenrolfea, 454

Allophylus, 480-27, 616-35

Alocasia, 198-11

Alpinia, 595-33

Alstonia, 403-22, 422-23

Alternanthera, 328-18, 480-27,
603-34

Alyxia, 350-19, 421-23, 422-23,
616-35

Amaracarpus, 603-34

Amaranthus, 47-4, 480-27

Amaryllis, 155-9

Ambroma, 328-18

Ammannia, 488-28

Andrographis, 488-28, 618-35

Andruris, 603-34

Anectochilus, 74-5

Aneilema, 488-28

Anemone, 53-4

Angiopteris, 117-7

Angraecum, 480-27

Aniseia, 455-25

Anisophyllea, 499-29

INDEX TO GENERIC NAMES

Annona, 480-27
Antidesma, 10-2
Aphelandra, 618-35
Apodytes, 480-27
Arachniodes, 616-35
Araucaria, 435-24
Ardisia, 422-23

Areca, 171-10

Arenaria, 61-5
Argemone, 267-15
Argyreia, 455-25
Armoracia, 300-17, 325-18
Artemisia, 482-27
Arthrocnemum, 480-27
Artocarpus, 47-4, 230-13
Arundina, 355-20
Asarina, 267-15
Ascarina, 403-22
Asparagus, 480-27
Aster, 482-27

Asystasia, 480-27, 618-35
Atriplex, 14-3, 57-4, 61-5
Avicennia, 480-27
Azima, 480-27

B

Bacopa, 437-24, 480-27

Badusa, 603-34, 618-35

Bakerella, 480-27

Balanophora, 616-35

Balmea, 417-23

Bambusa, 480-27

Barleria, 480-27, 618-35

Barringtonia, 500-29

Bassia, 454

Batocarpus, 66-5

Bidens, 480-27, 482-27, 490-28

Bikkia, 37-4, 603-34, 618-35

Blechum, 618-35

Blumea, 332-18, 482-27

Bobea, 443-25, 556-31

Boehmeria, 433-24

Boerhavia, 459-26, 467-26, 480-
27, 566-32, 590-33

Bombax, 595-33

Borreria, 16-3, 603-34

Botrychium, 256-14

Brassaia, 412-23

Brassica, 480-27

37

Brexia, 480-27
Bridelia, 603-34
Broussaisia, 31-3
Bruguiera, 107-7, 267-15, 328-18,
480-27, 481-27, 499-29
Bryodes, 480-27
Buchanania, 603-34
Buddleja, 48-4
Bulbophyllum, 603-34
Bulbostylis, 449-25, 480-27
Buxus, 413-23

Cc

Caesalpinia, 407-22, 480-27

Calliandra, 480-27, 481-27

Callicarpa, 472-26, 481-27

Calophyllum, 418-23, 480-27,
481-27, 547-31

Calpicarpum, 456-25

Campanula, 159-9

Campium, 39-4

Canavalia, 317-17, 433-24, 480-
Di:

Canthium, 16-3, 37-4, 161-9, 466-
26, 480-27, 603-34, 618-35

Capparis, 480-27

Capsicum, 480-27

Careya, 500-29

Carica, 480-27

Carissa, 480-27

Cassia, 346-19, 355-20, 480-27

Cassipourea, 480-27

Cassytha, 10-2, 480-27

Casuarina, 328-18, 436-24, 480-
27, 484-28, 596-33

Catalpa, 618-35

Catharanthus, 422-23, 480-27

Ceanothus, 61-5

Celtis, 44-4, 603-34

Cenchrus, 480-27

Centosteca, 481-27

Cephaelis, 618-35

Ceratopteris, 186-11

Cerbera, 422-23, 595-33, 603-34

Cereus, 5-2

Ceriops, 480-27, 499-29

Chamaecyparis, 64-5

Chelone, 92-6

Chenopodium, 57-4, 267-15

38

Chilopsis, 12-2

Chionanthus, 615-35

Chloris, 441-25

Chromolaena, 482-27, 490-28

Chrysanthemum, 482-27

Chrysophyllum, 422-23

Cichorium, 482-27

Cinchona, 75-5, 78-5, 79-5, 83-6,
91-6, 94-6, 112-7, 618-35

Cissampelos, 480-27

Citrullus, 138-8, 169-10, 481-27

Citrus, 480-27

Claoxylon, 493-28

Cleome, 480-27

Clerodendrum, 464-26, 480-27

Clinostigma, 171-10

Clitandropsis, 603-34

Clitoria, 346-19, 480-27

Clytostoma, 618-35

Coccinia, 595-33

Cochlearia, 300-17, 325-18

Cocos, 171-10, 198-11, 480-27

Coffea, 603-34, 618-35

Colubrina, 480-27

Comandra, 434

Commelina, 480-27

Conega, 480-27

Connarus, 437-24

Conyza, 482-27

Coprosma, 16-3, 71-5, 74-5, 166-
9, 349-19

Corchorus, 328-18, 480-27

Cordia, 332-18, 346-19, 480-27,
493-28

Cordyline, 534-31

Coreopsis, 482-27

Cornus, 69-5, 98-6

Coryphantha, 2-2

Cosmos, 482-27, 490-28

Couroupita, 500-29

Crassocephalum, 482-27

Crassula, 350-19

Crescentia, 618-35

Crinum, 346-19, 571-32

Crossandra, 618-35

Crotalaria, 332-18, 438-24, 480-
27

Cryptocarya, 10-2

Cryptomeria, 435-24

Cucumis, 480-27

Cucurbita, 480-27

Cupressus, 435-24

Curcuma, 571-32

Cuscuta, 455-25

Cycas, 435-24

Cycloloma, 45-4

Cydista, 618-35

Cymbopogon, 480-27

Cymodocea, 480-27

Cynara, 482-27

Cynodon, 531-30

Cyperus, 449-25, 480-27, 573-32,
595-33

Cyrtandra, 493-28, 616-35, 618-
35

D

Dactyloctenium, 480-27

Daknopholis, 480-27

Datura, 203-12, 480-27

Decaspermum, 603-34

Deeringia, 480-27

Delonix, 480-27

Dendrobium, 603-34

Dentella, 618-35

Desmodium, 328-18, 350-19, 532-
30

Dianella, 198-11, 355-20, 571-32,
616-35

Diaporthiopsis, 536-31

Dichrocephala, 493-28

Dichrostachys, 480-27

Dicliptera, 616-35

Didymoplexis, 571-32

Digitaria, 30-3, 38-4, 161-9, 480-
27, 489-28, 531-30

Dioscorea, 480-27, 571-32

Diospyros, 10-2, 24-3, 32-3, 384,
566-32

Diplocaulobium, 603-34

Dischidia, 39-4

Discocalyx, 603-34

Dodonaea, 480-27

Dolichandrone, 618-35

Dracaena, 267-15, 480-27

Drymaria, 564, 84-6, 110-7

Dryopteris, 71-5, 317-17

Dubautia, 74-5

E

Echinocactus, 5-2, 61-5
Echinocereus, 5-2
Echinochloa, 531-30
Eclipta, 482-27

Egeria, 346-19

Ehretia, 418-23, 480-27
Elaeagnus, 51-4, 61-5

Elaeocarpus, 198-11, 603-34

Elatostema, 481-27

Elephantopus, 14-3, 74-5, 482-27

Eleusine, 480-27

Emilia, 107-7, 317-17, 395-22,
482-27

Enteropogon, 480-27

Epaltes, 482-27

Episcia, 618-35

Equisetum, 151-9

Eragrostis, 30-3, 480-27, 595-33

Eranthemum, 148-8, 618-35

Erigeron, 482-27

Eriocaulon, 488-28

Eriochloa, 480-27

Eriogonum, 20-3

Ervatamia, 422-23

Erythrina, 480-27, 615-35

Erythroxylum, 480-27

Eugenia, 461-26, 480-27, 488-28,
489-28

Euodia, 603-34

Eupatorium, 14-3, 107-7, 433-24,
490-28

Euphorbia, 88-6, 95-6, 141-8,
310-17, 418-23, 433-24, 465-
26, 480-27, 566-32, 615-35,
616-35

Eurotia, 454

Eurya, 10-2, 198-11

Eustachys, 433-24

Evolvulus, 455-25, 480-27

F

Fagraea, 420-23, 422-23, 481-27,
603-34

Fendlera, 564

Ficus, 39-4, 161-9, 480-27

Fimbristylis, 480-27, 568-32, 573-
32, 603-34

Fittonia, 618-35

Flacourtia, 418-23, 480-27, 481-
27

Fragaria, 355-20

Freycinetia, 198-11

G

Gaillardia, 482-27

Galium, 310-17, 322-18
Galphimia, 489-28

Garcinia, 460-26, 481-27, 603-34

Gardenia, 618-35

Geniostoma, 422-23, 438-24, 481-
27, 493-28, 603-34

Geophila, 265-14, 481-27, 561-
32, 618-35

Geranium, 11-2, 14-3, 71-5

Gilia, 67-5

Gleichenia, 117-7, 355-20, 481-27

Glochidion, 603-34

Glossogyne, 482-27

Gmelina, 603-34

Gnetum, 435-24

Gonzalagunia, 322-18

Gossypium, 480-27

Gouania, 24-3, 480-27

Gouldia, 15-3, 74-5, 166-9, 349-
19

Graptophyllum, 618-35

Grayia, 454

Grewia, 480-27, 616-35

Guettarda, 16-3, 466-26, 480-27,
616-35, 618-35

Gulubia, 171-10

Gynandropsis, 480-27

Gynochthodes, 37-4, 38-4, 618-35

H

Habenaria, 571-32, 603-34

Halodule, 480-27

Halogeton, 454

Halophila, 414-23, 480-27

Hederorkis, 480-27

Hedyotis, 16-3, 374, 38-4, 394,
DB -AND6-4) 72-51 3-5 Se
151-9, 160-9, 161-9, 166-9,
349-19, 438-24, 480-27, 481-
27, 488-28, 490-28, 545-31,
603-34, 604-34, 618-35

Heliotropium, 481-27

Hemigraphis, 328-18, 618-35

Hernandia, 198-11, 480-27, 595-
33

Hetaeria, 603-34

Heterospathe, 171-10

Hibiscus, 332-18, 480-27

Hillia, 322-18

Hippobroma, 107-7, 482-27

Hippocratea, 603-34

Hippuris, 50-4

Hoffmannia, 322-18

Homalanthus, 603-34

Houstonia, 113-7

Hoya, 394

Humata, 488-28

Hydnophytum, 39-4, 618-35
Hygrophila, 494-28, 618-35
Hymenophyllum, 117-7, 481-27
Hypericum, 355-20

Hypoestes, 480-27

I

Ilex, 603-34

Impatiens, 92-6

Imperata, 531-30

Indigofera, 480-27

Inocarpus, 474

Ipomoea, 267-15, 328-18, 418-23,
438-24, 446-25, 455-25, 480-27

Isachne, 531-30

Ischaemum, 480-27, 531-30

Ixora, 16-3, 21-3, 22-3, 374, 38-
4, 107-7, 585-33, 586-33, 616-
35, 618-35, 624-36

J

Jacaranda, 618-35
Jasminum, 422-23, 480-27
Jatropha, 444-25
Juniperus, 64-5, 435-24
Justicia, 480-27, 618-35

K

Kalanchoé, 107-7
Kandelia, 499-29

Kigelia, 107-7, 618-35
Kolreuteria, 489-28
Korthalsella, 71-5, 616-35

L

Lactuca, 418-23, 482-27
Lagenaria, 480-27
Lagrezia, 418-23, 480-27
Lantana, 444-25, 480-27
Laportea, 480-27
Launaea, 480-27
Lebronnecia, 331-18
Lemaphyllum, 615-35
Lemna, 489-28

Leonotis, 480-27
Lepechinia, 71-5
Lepidium, 346-19, 355-20, 615-35

39

Lepinia, 422-23

Lepisanthes, 489-28

Leptopetalum, 604-34

Leptostigma, 504-29

Lepturopetium, 531-30

Lepturus, 162-9, 346-19, 480-27,
566-32

Leucaena, 267-15, 313-17

Leucas, 488-28, 493-28

Lewisia, 61-5

Limnophila, 437-24

Liparis, 571-32

Lippia, 444-25

Livistona, 171-10

Lobelia, 25-3

Loeseneriella, 481-27, 603-34

Lomatophyllum, 480-27

Lucuma, 422-23

Ludia, 480-27

Luffa, 615-35

Lumnitzera, 480-27

Luzula, 17-3

Lycopodium, 39-4, 117-7, 616-35

Lygodium, 117-7

M

Macaranga, 198-11
Macfadyena, 618-35
Machaerina, 573-32
Macphersonia, 480-27
Macrocystis, 1-2

Maerua, 480-27

Maesa, 473-26, 474-26, 603-34
Maillardia, 418-23, 480-27
Malachra, 74-5

Malaxis, 571-32, 603-34
Malleastrum, 418-23, 480-27
Manilkara, 291-16, 422-23
Mansoa, 618-35

Marattia, 117-7
Margaritaria, 465-26, 480-27
Maurandya, 74-5

Maytenus, 161-9, 480-27
Medicago, 355-20
Medinilla, 39-4, 481-27
Melaleuca, 488-28
Melanolepis, 161-9, 198-11
Melanthera, 480-27
Melastoma, 481-27
Melicoccus, 493-28
Melicope, 603-34

Melochia, 438-24, 595-33
Melodinus, 595-33, 616-35

40

Merremia, 429-24, 455-25

Metrosideros, 513-29, 520-30

Metroxylon, 171-10

Miconia, 493-28

Microstylis, 571-32, 603-34

Mikania, 482-27

Mimosa, 346-19

Mirabilis, 56-4, 97-6, 480-27,
590-33

Miscanthus, 198-11, 596-33, 616-
35

Mitracarpus, 618-35

Moerenhoutia, 603-34

Mollugo, 49-4, 480-27

Mollugophytum, 84-6

Momordica, 480-27

Monarda, \47-8, 217-12

Monolepis, 454

Monotropa, 159-9

Morinda, 16-3, 374, 384, 603-
34, 618-35

Moringa, 480-27

Musa, 107-7

Mussaenda, 37-4, 618-35

Mycetia, 618-35

Myoporum, 618-35

Myrica, 14-3, 436-24

Myriophyllum, 14-3, 504

Myrsine, 438-24, 481-27, 493-28,
603-34

Mystroxylon, 480-27

N

Najas, 480-27

Neiosperma, 403-22, 421-23, 453-
25, 456-25

Neomammillaria, 2-2, 5-2

Neonauclea, 38-4

Nephrolepis, 74-5, 480-27, 488-
28, 615-35

Nerium, 422-23

Nertera, 16-3

Nervilia, 571-32, 607-34

Nesogenes, 480-27, 529-30, 589-
33

Nicotiana, 480-27

Nitrophila, 45-4

Nypa, 171-10

O

Obetia, 480-27

Ochna, 480-27

Ochrosia, 403-22, 421-23, 422-
23, 433-24, 453-25, 492-28,
615-35

Ocimum, 480-27

Odontochilus, 355-20

Odontonema, 618-35

Oenothera, 67-5, 151-9

Oldenlandia, 462-26, 545-31,
603-34

Olea, 422-23

Operculicarya, 480-27

Operculina, 455-25

Ophioglossum, 267-15, 416-23

Ophiorrhiza, 16-3, 374, 384,
618-35

Oplismenus, 531-30

Opuntia, 2-2, 4-2, 5-2, 7-2

Osmanthus, 317-17

Osmoxylon, 481-27

Oxalis, 310-17, 480-27, 493-28,
595-33

P

Paederia, 595-33, 618-35

Palaquium, 422-23

Pandaca, 480-27

Pandanus, 198-11, 450-25, 480-
2,

Panicum, 332-18, 480-27, 488-28

Parmentiera, 618-35

Paronychia, 564

Paspalum, 30-3, 448-25, 480-27,
531-30, 616-35

Passiflora, 480-27, 615-35

Pavonia, 332-18, 495-28, 544-31

Pedilanthus, 480-27

Pemphis, 480-27

Pennisetum, 480-27, 531-30

Pentas, 618-35

Pentopetia, 480-27

Peperomia, 320-18, 403-22, 436-
24, 615-35, 616-35

Peponium, 480-27

Peristylus, 571-32

Phaleria, 38-4

Phoenix, 480-27

Phoradendron, 56-4

Phreatia, 603-34

Phyllanthus, 10-2, 465-26, 480-
27, 615-35

Phyllostegia, 10-2, 71-5

Pimenta, 67-5

Pinanga, 171-10

Pinus, 64-5, 200-11, 435-24
Piper, 320-18, 436-24
Piriqueta, 488-28
Pisonia, 480-27, 555-31, 560-32
590-33
Planchonella, 422-23
Plantago, 438-24, 618-35
Plectronia, 603-34
Pleurostelma, 480-27
Pluchea, 14-3, 482-27
Plumbago, 418-23, 422-23, 480-
27
Plumeria, 422-23
Podocarpus, 435-24
Pogonatherum, 531-30
Polygonum, 355-20
Polypodium, 348-19, 355-20, 589-
33, 615-35, 616-35
Polyscias, 107-7, 161-9
Polysphaeria, 466-26, 480-27
Polytrias, 531-30
Portulaca, 73-5, 437-24, 451-25,
480-27
Poupartia, 418-23
Pouteria, 267-15, 481-27, 615-35
Premna, 137-8, 198-11, 480-27,
603-34
Prosopis, 317-17
Pseudelephantopus, 74-5
Pseuderanthemum, 39-4, 161-9,
481-27, 541-31, 618-35
Pseuderia, 603-34
Pseudo-Elephantopus, 482-27
Pseudogynoxys, 482-27
Psidium, 55-4, 267-15, 383-21
Psychotria, 16-3, 34-3, 37-4, 38-
4, 161-9, 267-15, 288-16, 322-
18, 466-26, 480-27, 481-27, |
512-29, 589-33, 595-33, 603-
34, 604-34, 618-35
Pteris, 198-11
Ptychosperma, 171-10
Pyrostegia, 618-35

—eeeeeeeeeeee

R

Randia, 24-3, 37-4, 490-28

Rapanea, 422-23, 603-34

Rauvolfia, 493-28

Rhamnus, 56-4

Rhaphidophora, 481-27

Rhizophora, 480-27, 499-29, 616-
35

Rhynchophreatia, 571-32, 603-34

Richardia, 67-5

Ricinus, 480-27

Rollandia, \0-2

Ruellia, 74-5, 480-27, 530-30,
618-35

Rumex, 355-20

Ruppia, 36-3, 480-27

S

Saccharum, 531-30

Sadleria, 442-25

Sagittaria, 346-19

Saintpaulia, 618-35

Salacia, 603-34

Salacicrates, 603-34

Salicornia, 438-24

Salomonia, 437-24

Salsola, 454

Salvadora, 480-27

Sambucus, 67-5, 267-15, 482-27

Santalum, 267-15, 355-20, 543-31

Sarcobatus, 454

Sarcostemma, 480-27

Saritaea, 618-35

Saxifraga, 53-4

Scaevola, 170-10, 247-13, 266-14,
350-19, 480-27, 482-27, 589-33

Schiedea, 10-2

Schizaea, 76-5, 117-7

Sciaphila, 491-28

Scirpus, 321-18

Scleria, 625-36

Sclerodactylon, 480-27

Scutia, 480-27

Scyphiphora, 618-35

Serianthes, 204-12, 233-13, 272-
15, 356-20

Sesamum, 618-35

Sesbania, 616-35

Sesuvium, 49-4, 136-8, 480-27

Setaria, 217-12, 355-20, 490-28

Sida, 14-3, 480-27

Sideroxylon, 480-27

Solanum, 161-9, 463-26, 480-27,
557-31

Sonchus, 482-27

Sonneratia, 480-27, 501-29

Sophora, 480-27

Sorghum, 531-30

Spathodea, 618-35

Spathoglottis, 198-11, 346-19,
571-32, 615-35

Spergularia, 14-3, 74-5, 107-7

Spermacoce, 481-27, 488-28, 557-
31, 566-32, 603-34, 618-35

Sphenomeris, 357-20, 505-29
Spilanthes, 482-27
Spiraea, 53-4, 310-17
Sporobolus, 480-27
Stachytarpheta, 480-27
Stemonurus, 603-34
Stenotaphrum, 480-27
Sterculia, 603-34
Stictocardia, 455-25
Striga, 437-24, 480-27
Stylosanthes, 481-27
Styphelia, 25-3, 422-23
Styrax, 422-23

Suriana, 480-27
Symplocos, 603-34
Synedrella, 480-27, 482-27
Syringodium, 480-27
Syzygium, 603-34

T

Tabebuia, 480-27, 618-35

Taeniophyllum, 571-32, 603-34

Tagetes, 482-27

Tamarindus, 480-27

Taraxacum, 92-6

Tarenna, 16-3, 466-26, 480-27,
604-34, 618-35

Taxodium, 64-5

Taxus, 122-7

Tecoma, 618-35

Tecomaria, 618-35

Tephrosia, 480-27, 615-35

Teramnus, 480-27

Terminalia, 332-18, 419-23, 480-
27, 493-28, 589-33, 616-35

Thalassia, 480-27

Thalassodendron, 480-27

Thelypteris, 217-12, 267-15, 350-
19, 355-20, 401-22, 481-27,
496-29

Thespesia, 400-22, 480-27

Thevetia, 422-23

Thunbergia, 618-35

Thylax, 191-11

Tilia, 159-9, 310-17

Timonius, 16-3, 37-4, 38-4, 74-5,
488-28, 572-32, 603-34, 618-35

Tithonia, 482-27

Tmesipteris, 616-35

Tournefortia, 480-27

Trachoma, 571-32

Trema, 198-11

Triainolepis, 466-26, 480-27

4]

Trianthema, 480-27

Tribulus, 480-27

Tricalysia, 466-26, 480-27

Trichoglottis, 603-34

Tricholaena, 493-28

Trichomanes, 117-7, 481-27, 496-
29

Trichosanthes, 196-11, 480-27

Tricomanes, 488-28

Tridax, 480-27, 482-27

Tridens, 159-9

Triodia, 87-6

Tristellateia, 328-18

Tristiropsis, 198-11

Triumfetta, 480-27, 493-28

Trukia, 554-31, 618-35

Tsuga, 64-5

Turnera, 480-27

Tylophora, 480-27

Typha, 576-32

U

Uncaria, 374, 618-35
Urandra, 603-34

Urera, 167-9

Utricularia, 488-28, 618-35

V

Vaccinium, 71-5, 422-23, 438-24

Vandopsis, 603-34

Vanilla, 615-35

Vernonia, 332-18, 480-27, 481-
27, 482-27

Veronica, 53-4, 159-9

Vigna, 480-27, 493-28

Viola, 310-17

Viscum, 480-27

Vitex, 267-15

W

Waltheria, 421-23, 438-24, 616-
35

Wedelia, 161-9, 482-27, 490-28

Weinmannia, 403-22

Wikstroemia, 616-35

Wollastonia, 481-27, 482-27, 490-
28, 616-35

42

X

Xanthoxylon, 191-11
Ximenia, 616-35
Xylocarpus, 480-27
Xylosma, 616-35
Xyris, 317-17

Y

Youngia, 482-27

Z

Zanthoxylum, 191-11, 205-12,
355-20

Zea, 480-27

Zehneria, 481-27, 493-28

Zinnia, 482-27

ATOLL RESEARCH BULLETIN

NO. 392

THE FLORA OF NAURU

BY

R.R. THAMAN, F.R. FOSBERG, H.I MANNER
AND D.C. HASSALL

ISSUED BY
NATIONAL MUSEUM OF NATURAL HISTORY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C., U.S.A.
FEBRUARY 1994

DEDICATION

We dedicate this Flora of Nauru to Joseph Detsimea
Audoa, his family and the people of the Republic of Nauru
who have had their precious island and its flora destroyed
and degraded as a result of wars and exploitation beyond
their control.

ACKNOWLEDGEMENTS

The authors would like to acknowledge, in particular, the late Honorable Joseph
Detsimea Audoa, the Minister of Health and Education at the time of the commencement
of the study and later Minister of Justice in the Government of Nauru, who, because of
his vision and commitment to the culture and environment of Nauru, initiated and
provided the financial support for the study of the flora of Nauru. He was particularly
concerned that the plants of Nauru and their cultural uses be recorded before such
knowledge was lost.

We also acknowledge Mr. Lisle Newby, the then Director of Education, who,
along with Joe Audoa, were the main supporters of the project, and who provided
valuable logistical support throughout. Special thanks are also given to our main local
informants and assistants, the Reverend James Aingimea and the late Henry Michael
Heine; and to Daphne Fotu, Jacob Gabwinare, Katarina Satto, Kenia Raidinen, Reynold
Capelle, Eda Adam and Montiba Star, our main informants in relation to the cultural uses
and Nauruan names of plants.

Our thanks also go to the Honorable Lawrence Stephen, Minister of Education
during part of the project; Obera Menke, Robert Kaierua, Leo Keke, Delilah Capelle,
Eddie Borak, John Healy, Gary Bailey, Dennis and Ria Berdinner, Julie Olsson, Dennis
Ketner, Sio Fotu, Pine Harrison, John Brechtefeld, Rene Harris, Porthos Bop, Jacob
Aroi, Leon Thompson, Benjamin Morgan, Iosefa Elisala and Teaora Tabanou, all of
whom contributed in some way to the success of the study. To others who helped in any
way during our study, we also give thanks.

Thanks are also due to those people, who over the past 100 years, have collected
and identified plants on Nauru; and to Saula Vodonaivalu of the South Pacific Regional
Herbarium of The University of the South Pacific, Suva, Fiji who identified, preserved
and is the curator of the herbarium specimens collected by Thaman, Manner and Hassall.
Without their efforts, this flora, would have been impossible. Similarly, we would like to
thank The University of the South Pacific for the support it has provided throughout the
duration of the study, both as the institution where the study was initiated and completed
and as Nauru’s own university of which it is one of twelve regional member countries.

In addition to the above acknowledgements, Fosberg wants to mention the
substantial assistance given him during his week-long visit to Nauru in 1983 This visit
was at the invitation of Mr. Richard Wood, accountant for the Nauru government, who
made the arrangements for the visit and who served as guide during the first day,
reaching areas not otherwise accessible. The success of this visit was also made
possible by transport and guidance by Mrs. Joan Nichols of the Australian High Commis-
sion’s office. In her company much of the interior of the island and some of the local
gardens were visited and many plant specimens were gathered. The success of this visit

was largely due to the two people mentioned above, and their company in the field was
much enjoyed and appreciated. Many thanks!

Finally, we wish to express our heartfelt thanks to the people of Nauru whose
warmth and hospitality made our work on their beautiful but damaged island so enjoyable
and worthwhile.

To all of you, TUBWA KOR,
R.R Thaman, F.R. Fosberg, H.I. Manner and D.C. Hassall

Suva, Fiji
June 1993

CONTENTS

PART I: INTRODUCTION AND ANALYSIS OF THE
VEGETATION AND FLORA

INTRODUCTION
PREVIOUS STUDIES
CURRENT STUDY
BACKGROUND ON NAURU

Physical Environment
The People
Development History
Contemporary Economy
Impact on the Flora

VEGETATION TYPES

Coastal Strand Vegetation

Mangroves and Coastal Marsh Vegetation

Relict Stands of Inland Forest

Limestone Escarpment or Pinnacle Vegetation

Coconut-Palm- and Pandanus-Dominated
Agricultural Lands

Houseyard Gardens and Urban Vegetation

Ruderal Vegetation

Phosphate-Mined Lands

THE FLORA
Nature of Indigenous Species
Comparison with Other Island Floras
Nature of Exotic Species

ECOLOGICAL AND CULTURAL UTILITY OF EXISTING FLORAS

CONCLUSION

PAGE

ii

PART II: A COMPILATION OF THE VASCULAR FLORA OF NAURU

KEY

Latin/Scientific Names

Family

Common Names

Vernacular Names

Antiquity Status

Geographical Origin

Abundance or Frequency Occurrence
Species Description

Uses and Cultural Utility

Collectors and Herbarium Specimens

VASCULAR PLANTS OF NAURU

PTERIDOPHYTA (Ferns and Fern Allies)

ASPLENIACEAE
DAVALLIACEAE
OPHIOGLOSSACEAE
POLYPODIACEAE
PSILOTACEAE
PTERIDACEAE

GYMNOSPERMAE (Gymnosperms)

ANGIOSPERMAE (Angiosperms or Flowering Plants)

ARAUCARIACEAE
CYCADACEAE

MONOCOTYLEDONAE

ARACEAE
ARECACEAE/PALMAE
BROMELIACEAE
CANNACEAE
COMMELINACEAE
CYPERACEAE
DIOSCOREACEAE
IRIDACEAE

LILIACEAE (Including Agavaceae and Amaryllidaceae)

MARANTACEAE
MUSACEAE
ORCHIDACEAE
PANDANACEAE

POACEAE OR GRAMINEAE
PONTEDERIACEAE
STRELITZIACEAE
TACCACEAE
ZINGIBERACEAE

DICOTYLEDONAE

ACANTHACEAE
AMARANTHACEAE
ANACARDIACEAE
ANNONNACEAE

APIACEAE OR UMBELLIFERAE
APOCYNACEAE
AQUIFOLIACEAE
ARALIACEAE
ASCLEPIADACEAE
ASTERACEAE OR COMPOSITAE
BALSAMINACEAE
BASELLACEAE

BEGONIACEAE
BIGNONIACEAE

BOMBACEAE

BORAGINACEAE
BRASSICACEAE OR CRUCIFERAE
CACTACEAE
CAPPARIDACEAE OR CAPPARACEAE
CARICACEAE
CASUARINACEAE
CHENOPODIACEAE
CLUSIACEAE OR GUTTIFERAE
COMBRETACEAE
CONVOLVULACEAE
CRASSULACEAE
CUCURBITACEAE

ERICACEAE

EUPHORBIACEAE

FABACEAE OR LEGUMINOSAE
GENTIANACEAE
GERANIACEAE

iv

GESNERIACEAE
GOODENIACEAE
HERNANDIACEAE

LAMIACEAE OR LABIATAE

LAURACEAE
LECYTHIDACEAE
LYTHRACEAE
MALPIGHIACEAE
MALVACEAE
MELIACEAE
MORACEAE
MORINGACEAE
MYRTACEAE
NYMPHAEACEAE
NYCTAGINACEAE
OLEACEAE
ONAGRACEAE
OXALIDACEAE
PASSIFLORACEAE
PIPERACEAE
POLYGALACEAE
POLYGONACEAE
PORTULACACEAE
RHAMNACEAE
RHIZOPHORACEAE
ROSACEAE
RUBIACEAE
RUTACEAE
SAPINDACEAE
SAPOTACEAE
SAXIFRAGACEAE
SCROPHULARIACEAE
SOLANACEAE
STERCULIACEAE
SURIANACEAE
TILIACEAE
URTICACEAE
VERBENACEAE
VITACEAE

BIBLIOGRAPHY

153
155
156
156
159
160
161
161
162
169
170
173
174
175
176
177
178
178
179
181
181
182
183
183
184
185
185
191
193
194
194
194
196
200
201
201
202
203
208

209

Table 1.

Table 2.

Table 3.

Table 4.

Appendix I.

LIST OF TABLES
Antiquity status of the flora of Nauru

Species indigenous or possibly indigenous
to Nauru

Frequency of use for specified purposes of plant
species present in Nauru

Species of particular cultural utility on Nauru

LIST OF APPENDICES

Class, subclass, family and antiquity status of the
vascular flora of Nauru

Appendix II. Nature and ecological and cultural (ethnobotanical)

importance of coastal plant species of the
tropical Pacific Ocean

15

16

23

26

216

219

NAURU

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{hn

THE FLORA OF NAURU
A COMPILATION AND ANALYSIS OF THE VEGETATION AND FLORA OF
THE EQUATORIAL PACIFIC OCEAN ISLAND OF NAURU
BY

R.R. THAMAN|, F.R. FOSBERG?, H.I. MANNER? AND D.C. HASSALL*

PART I.

INTRODUCTION AND ANALYSIS OF THE VEGETATION AND
FLORA

INTRODUCTION

The recorded flora of the raised phosphatic limestone island of Nauru in the
equatorial Pacific Ocean consists of approximately 493 species. Only 59 are possibly
indigenous, none of which are endemic. The balance is composed of ornamentals, weedy
exotics, food plants, and a limited number of other useful cultigens. Twelve of the
recorded species are either extinct or were never successfully established on Nauru. Long
human settlement, expansion of coconut monoculture during the colonial period,
widespread destruction during World War II, and almost a century of open-cast phos-
phate mining, have led to serious vegetation degradation, disturbance, and displacement

'. Professor of Pacific Islands Biogeography, Department of Geography, School of Social
and Economic Development, The University of the South Pacific, Suva, Fiji.

*. Emeritus Professor of Botany, Botany Department, National Museum of Natural
History, Smithsonian Institution, Washington D.C.

3. Professor of Geography, College of Arts and Sciences, Department of Social Science
and Anthropology, University of Guam, Mangilao, Guam.

*. Former Senior Lecturer in Biology, University of the South Pacific, Suva, Fiji.

Manuscript received 12 July 1993; revised 6 August 1993.

2

of the indigenous flora. Although greatly outnumbered by exotics, indigenous species still
dominate some of the most disturbed habitats, as well as constituting the most culturally-
utilitarian and ecologically-important species.

The flora is in two parts. PART I: INTRODUCTION AND ANALYSIS OF THE
VEGETATION AND FLORA, includes a brief discussion of: 1) previous studies and the
current study; 2) the physical environment, people, development history and contem-
porary economy of Nauru; 3) the nature of the existing flora and vegetation associations
of Nauru; and, 4) an analysis of the ecological and cultural importance of the flora in the
context of modern small-island development.

PART II: A COMPILATION OF THE VASCULAR FLORA OF NAURU,
consists of a listing of all vascular plants reported to have been present on Nauru, along
with relevant information on each species. Although a number of persons have collected
plants on Nauru over the past century, this constitutes the first extensive flora the of
vascular plants of the island. The plants are listed in alphabetical order by family and
species within families, starting with the ferns and gymnosperms and then angiosperms,
with monocotyledons listed first. Information on each species, variety or subspecies
includes: 1) scientific (Latin) name and synonyms; 2) common (mainly English) name
(s); 3) Nauruan, Kiribati, Tuvaluan, Chinese (Cantonese), Filipino and Solomon Island
names, when available; 4) origin; 5) antiquity status (i.e., whether a species is indigenous
to Nauru, an aboriginal introduction, an early post-European-contact introduction or a
recent post-World War II introduction; 6) its status in terms of abundance (i.e., whether
it is abundant, common, rare, extinct, etc); 7) a detailed botanical description; 8) its
habitat or distribution on Nauru; and 9) its cultural or ethnobotanical importance.

PREVIOUS STUDIES

Although collections and observations of the flora have been made on Nauru by
Finch prior to 1900; Burges in 1933; Fosberg in 1980; Scully in 1980; Thaman, Hassall
and Manner in 1980 and 1981; Thaman and Manner in 1987; Swarbrick in 1988; and
Raulerson in the early 1980s, little has been published. The only substantial publications
on Nauru’s vegetation include studies by Manner, Thaman and Hassall (1984 and 1985)
on vegetation changes induced by phosphate mining; a list of Nauruan plant names by
Thaman, Manner and Hassall (1985); and the "Vegetation of Nauru and the Gilbert
Islands" by Thaman (1992). Fosberg, Sachet and Oliver’s "A geographical checklist of
Micronesian Dicotyledonae" (1979); "Geographical checklist of Micronesian Pteridophyta
and gymnosperms" (1982); and "A geographical checklist of the Micronesian Mono-
cotyledonae" (1987), are also important and list most of the species cited and specimens
examined prior to 1980.

————

CURRENT STUDY

The current study is based on an in-depth analysis of the above sources; five-
weeks fieldwork, including two visits by R.R. Thaman, H.I. Manner and D.C Hassall of
The University of the South Pacific, Suva, Fiji in Nauru in 1980, 1981; one week each
by Thaman and Hassall in 1987; and a three-day visit by F.R. Fosberg of the Botany
Department of the Smithsonian Institution, Washington, D.C. in 1987. During these visits
herbarium specimens were collected and identified and the Latin and vernacular names of
all plant species recorded and listed alphabetically. These lists were then cross-checked,
consolidated, amended and augmented, both during and after the field studies, using lists
and names from previous studies. Information was also collected on the cultural uses
(ethnobotany) of the flora.

BACKGROUND ON NAURU
Physical Environment

The Republic of Nauru is an isolated, uplifted limestone island located 41 km
south of the equator at 166 deg 56 min E longitude, some 2000 km east-northeast of
Papua New Guinea, 4450 km south-southeast of the Philippines and an equal distance to
the southwest of Hawaii. The nearest island is Banaba (Ocean Island), 300 km due east,
which is part of the Republic of Kiribati. The Gilbert Islands, the main islands of
Kiribati, lie a further 400 km to the east.

The island, with an area of only 22 km’, consists of a narrow coastal plain,
ranging from 50 to 300 m wide, encircling a limestone escarpment rising some 30 m to
the central plateau (See map of Nauru). The escarpment ranges in gradient from vertical
cliffs to gradually-sloping areas of colluvial soil interspersed with limestone outcrops and
pinnacles. The plateau, with a maximum elevation of 70 m, consists of a matrix of coral-
limestone pinnacles and limestone outcrops, between which lie extensive deposits of soil
and high-grade tricalcic phosphate rock (Viviani 1970, Tyrer 1963). Buada Lagoon, a
landlocked brackish lake, and its associated fertile depression (about 12 ha in size), is
located in the low-lying southwest-central portion of the island.

Apart from Buada Lagoon, there are no surface freshwater resources on Nauru,
although there are a few brackish ponds on the northeast of the island and an under-
ground lake in Moqua Cave in the southeast (Viviani 1970:4). The only significant
permanent freshwater resource is groundwater in the form of a "lens" of often slightly
brackish freshwater, hydrostatically "floating" on higher density saltwater beneath it. The
height of the freshwater lens above sea level and the level of salinity vary in relation to
the elevation, geology, texture and shape of the island, and with the amount of water use
and rainfall. Replenishment or recharge of the lens is dependent on rainfall.

Climatically, Nauru is located in the dry belt of the equatorial oceanic zone, with
mean daily temperatures ranging from 26 to 32°C. Annual rainfall is extremely variable,
averaging 1500 mm per year with a range of 300 to 4572 mm. Severe prolonged
droughts are common and place severe stress on even the most hardy coastal strand
species, lead to the death of non-coastal exotics (such as breadfruit), and severely restrict
the production of even coconut palms (Catala 1957). For example, in 1917 and 1918,
during an unprecedented drought, when only 465 and 483 mm of rain fell, "thousands of
coconuts and other fruit trees died" (Griffiths 1923).

The coastal soils of Nauru are among the poorest in the world. They are shallow,
alkaline, coarse-textured and have carbonatic mineralogy. They are composed of a
variable layer of organic matter and coral sand and fragments, which overlay a limestone
platform. The coastal soils are only about 25 cm deep, and contain more coral gravel
than sand in the lower horizons. Potassium levels are often extremely low, and pH values
of up to 8.2 to 8.9 and high CaCO, levels make scarce trace elements, particularly iron
(Fe), manganese (Mn), copper (Cu) and zinc (Zn), unavailable to plants. Fertility is,
therefore, highly dependent on organic matter for the concentration and recycling of plant
nutrients, lowering soil pH, and for soil water retention in the excessively well-drained
soils. Although levels of organic matter can be relatively high in undisturbed soils under
natural vegetation, it can decrease dramatically as a result of clearance by fire or
replacement by coconuts and other introduced plants (Morrison 1987ab).

The plateau soils of Nauru vary from shallow soils, on the tops of limestone
pinnacles, composed primarily of organic material and sand or dolomite, with very little
phosphate, to deep phosphatic soils and sandy phosphatic rock, up to over 2 m deep
between the pinnacles. Topsoils range from 10 to 25 or 30 cm in depth, overlaying a
deeper material which is frequently reddish yellow and between 25 and 75 cm deep,
changing to pinkish grey at greater depth. Undisturbed plateau soils have a high level of
organic material and are generally fertile. Calcium dominates the exchange complex and
exchangeable magnesium is also high. Exchangeable potassium is low, while extractable
phosphate values are generally high and sulphate moderate. The trace elements man-
ganese, copper, cobalt and molybdenum levels are very low, and these, plus iron and
zinc, are rendered unavailable to plants under pH values >6.5 (Morrison 1987b).

Around Buada Lagoon and in some poorly drained swampy areas near the base of
the escarpment on Nauru, there are poorly developed, but relatively fertile, wet soils.

The People

The indigenous people of Nauru are Micronesians, who have probably inhabited
the island for up to 3000 years or more. There is some evidence of Melanesian, and
possibly Polynesian, influence. The Nauruan language is quite distinct from all other
Pacific languages, reportedly a fusion of elements from the Gilbert, Caroline, Marshall
and Solomon Islands. Early this century there was evidence of distinct racial types or

5

groups of mixed origin. The people were divided into twelve distinct, originally totemic,
matrilineal clans, most of which spoke different dialects, some of which were still in
evidence when trying to obtain vernacular plant names in the early 1980s. However, most
dialects have become obsolete, having been replaced by the principle dialect, which was
used for Bible translation by European missionaries early this century (Viviani 1970:4-7).

The traditional subsistence economy of Nauru was based on coconut and pandanus
as the main staples, a limited range of wild terrestrial food products, sea birds, such as
the black noddy tern (Anous tenuirostris) and a very wide range of fish and other marine
foods. Milkfish or ibija (Chanos chanos) fry, collected from the reef at low tide, were
farmed in family-owned divisions of Buada Lagoon to provide fish for special occasions
and when other supplies failed. Their housing, tools, clothing, medicines, fuel, fishing
equipment, canoes, dyes, ornamentation, perfumes, toys and other material and many of
their non-material needs were satisfied from their environment, especially from plants.

The Nauruan population suffered from introduced diseases against which they had
no natural resistance, and from incessant tribal warfare, with Nauruans numbering only
1250 in 1910, a decline of 150 from the total of 1400 in 1840 and 300 less than recorded
in the German census in 1905 (Viviani 1970:37).

The estimated population of Nauru in the most recent census in 1983 was 8042, of
whom 4964 were Nauruan, with the balance comprised mainly of I-Kiribati (people of
Kiribati, formerly the Gilbert Islands of the British Gilbert and Ellice Islands Colony),
Tuvaluan, Chinese, Filipino or Solomon Island contract workers in the phosphate
industry. There are also European, Indian and Pacific island expatriates working mainly
for the Nauruan Government. The Nauruans live on the coastal strip and around Buada
Lagoon, the phosphate workers in the Nauru Phosphate Company dormitory accom-
modation at Location near the phosphate loading cantilevers, and the expatriate civil
servants in residential areas on the escarpment. The town center is located between the
airport and Location near the cantilevers (Figure 1), with most government offices near
the airport.

Development History

After the first recorded European sightings of Nauru by John Fearn of the British
Ship Hunter in 1798 who named it "Pleasant Island", there was little regular contact with
the island until the 1830s when British and American whalers made regular stops for
water and food, and beachcombers arrived. The beachcombers, with the introduction of
new weaponry, intensified a period of almost incessant clan warfare. At the time of the
beginning of formal colonial influence when Nauru was incorporated into Germany’s
Marshall Islands Protectorate in 1888, the island "had the appearance of a battlefield"
(Viviani 1970:22). In 1919, after World War I, Nauru became a League of Nations
mandate of Great Britain, Australia and New Zealand, administered by Australia. Nauru
became independent in 1968.

Apart from the provisioning of whaling vessels, mainly with fish, pigs and
coconuts, the first regular non-traditional economic activity was a sporadic copra trade
established in the 1880s. By the late 1880s Nauru produced about one million pounds of
copra annually, one of the main reasons that Germany, on the recommendation of
resident German traders, annexed the island in 1888. In the 1890s, due to drought and
infrequency of ships, the copra trade failed to reach its potential with Nauruans refusing
to make more copra than needed to pay their taxes (Viviani 1970; Carter 1984).

The strategic and economic importance of Nauru increased dramatically with the
discovery, in 1900, of high grade phosphate rock, containing up to 78 per cent tricalcic
phosphate. The mining of phosphate, without approval of the indigenous inhabitants,
began in 1907. Caroline Islanders and Chinese contract-laborers were recruited to mine
the deposits. Since then, I-Kiribati, Tuvaluan, Filipino and Solomon Islands contract
workers have been recruited to work in the phosphate industry. The deposits have been
mined continuously since 1907, except for disruptions during World War I and again
during World War II. Copra production continued to fluctuate, with over 300 tons having
been exported the year before the phosphate trade began in 1906, 277 tons in 1916, and
falling to only 10 tons in 1918 due to a prolonged drought (Viviani, 1970: 22-38).

With the beginning of the phosphate trade, the whole pattern of Nauruan life
began to change. Although barter continued, money became the medium of exchange,
and trade stores with their array of goods, further encouraged the trend. As Viviani
(1970:38) argues:

Old crafts such as mat making began to be forgotten as woven
materials become available. Rites and customs were debased.
Faced on all sides by the white man’s attempt to dominate their
environment and the disintegration of their culture, the Nauruans
sought a new orientation for their lives. They could no longer
follow the old ways completely and so settled for a combination
of the basic elements of the old culture, clinging strongly to
their family life, and some of the advantages of Western civili-
sation. They were able to achieve this because, although royal-
ties were low - only about 230 Pounds Sterling per annum at
this time for the whole population - this money, together with
land rents and some return from copra, freed them from the
necessity of working on the phosphate fields to pay their taxes.

The most disruptive period for Nauru was World War II, during which the island
was continuously bombed by Japanese and Americans planes. Beginning in 1940, five
phosphate vessels were sunk off Nauru and the island shelled by German warships.
Nauru was again bombed by Japanese planes in 1941 and 1942 prior to Japanese
occupation of the island. After a Japanese military airstrip was completed in 1943, the
island was bombed almost continuously by Allied planes. By the end of 1943, due to the

7

importation of at least 3000 Japanese marines, some 1500 Japanese and Korean laborers,
and the relocation of 700 Banabans to Nauru, the food situation became so serious that
1201 Nauruans, seven Chinese and two priests were deported to Truk. Malnutrition and
dysentery were widespread. Allied bombings increased considerably until the Japanese
surrender to an Australian occupation force in 1945. At this time, of nearly 5200 people,
only 591 were Nauruan, and the destruction of the phosphate works and buildings on
Nauru was almost total. The 737 Nauruans who had not died under the harsh conditions
imposed by the Japanese in Truk returned home in 1946 (Viviani 1970; Carter 1984). As
argued by Viviani (1970:85):

The Japanese had destroyed the Nauruan’s homes, schools, and
churches, placed them on a semi-starvation level and destroyed
much of what was left of their old way of life. The deportation
of two-thirds of the Nauruans and the death of nearly 500,
mostly the old and the young, left the society after the war with
a gap in generations and a disruption of family life. Again the
Nauruan population had fallen well below the 1,500 level which
the Nauruans themselves regarded as a minimum for survival.

Contemporary Economy

Nauru’s sole export continues to be phosphate, with the sporadic export of copra
having ceased in the 1950s. Phosphate earnings have made Nauru among the wealthiest
nations in the world in terms of per capita income, although the distribution of wealth is
uneven due to unequal land rights to phosphate deposits. Nauru is considered totally
urbanized, with Nauruans having almost completely abandoned subsistence production,
except for the harvest of coconuts and pandanus fruit for consumption; pandanus leaves
for plaited ware; the acquisition of fish and other seafood; and the hunting of noddy
terns, which are considered a delicacy of chiefly status, and the capture and caring for of
frigate birds as pets, both traditional pastimes. Most of the limited subsistence agricul-
tural production is in the hands of immigrant communities.

The establishment of its own heavily subsidized international airline, Air Nauru,
in 1970, which flies to Asia, Australia, New Zealand and other Pacific islands, and the
extension of the runway over the reef have accelerated the processes of urbanization and
an increasing dependence on imported products. Of concern is the impact that the
destruction of the traditional subsistence food system, rapid urbanisation and the almost
total dependence on nutritionally-poor imported food and drink, including extremely high
rates of alcohol consumption, have had on the health of Nauruans, which have the among
highest or most rapidly increasing rates in the world of obesity, dental disease, al-
coholism and nutrition-related non-communicable diseases such as diabetes, cardiovas-
cular disease, gout and hyperuricemia and liver cancer. All are causes of premature

8

mortality (Zimmet ef al. 1977, 1978; Speake et al. 1979; Taylor 1983; Coyne 1984;
Thaman 1982, 1983, 1985, 1988a). ;

In terms of Nauru’s economic future, the phosphate deposits on Nauru are
projected to be depleted shortly after the turn of the century. Fortunately, a significant
proportion of recent phosphate returns have been wisely invested in overseas properties,
businesses and investment funds to provide income in post-phosphate-mining Nauru. A
further source of income is derived from Nauru’s status as a "financial center" or "tax
haven" for overseas companies wishing to register there. Finally, negotiations and
litigation arising out of a recent Commission of Inquiry into the Rehabilitation of the
Worked-out Phosphate Lands in Nauru to determine culpability and the extent of damages
to the Nauruan culture and environment are currently in process and could lead to
substantial payments to Nauru and the development of plans for the rehabilitation of the
island.

Impact on the Flora

Due to environmental factors described above - extreme isolation from major plant
source regions, small island size, extremely poor soils, and climatic and physiological
drought - the indigenous flora of Nauru is among the poorest and most restricted on
earth. Moreover, the long settlement history, widespread destruction during World War
II, monocultural expansion of coconut palms as the sole cash crop, increasing ur-
banization and contact with, and importation of plants from the outside world, and over
75 years of open-cast phosphate mining (in Nauru and Banaba), have all played a role in
the serious degradation, disturbance, and displacement of the indigenous flora and
vegetation.

VEGETATION TYPES

The terrestrial primary vegetation types of Nauru are limited to:

Ie coastal strand vegetation;

2 limited areas of mangroves and coastal marsh vegetation;
3 relict stands of inland forest; and,

4. limestone escarpment or pinnacle vegetation.

Secondary and cultural vegetation types include:

lt coconut-palm-dominated lands under various degrees of maintenance;

2: houseyard gardens and urban vegetation;
3: extensive and variable areas of ruderal vegetation; and,
4. three-quarters of the entire island under severely-modified disclimax

vegetation in various stages of succession after some 80 years of open-cast
phosphate mining.

These vegetation types will each be described in turn.

Coastal Strand Vegetation

The coastal strand vegetation of Nauru has been severely modified as a result of:
1) thousands of years of human habitation and selective removal of indigenous species for
construction, boatbuilding, firewood and other purposes; 2) the expansion of monocul-
tural coconut groves for export production of copra; 3) the expansion of coastal set-
tlements which occupy most of the coastal plain; and, 4) the widespread practice of
allowing pigs to forage freely along beach flats (Viviani 1970).

The dominant species in the outer coastal zone in Nauru include the herbaceous
species Lepturus repens, Cyperus javanicus, Ipomoea pes-caprae and Vigna marina; the
woody species, Scaevola taccada, Tournefortia argentea and Morinda citrifolia, plus the
aboriginal introduction, Cocos nucifera. Species common on rocky limestone outcrops
along the coast include the same species plus Polypodium scolopendria, Capparis
cordifolia, Clerodendrum inerme, Terminalia catappa and Calophyllum inophyllum.
Species present on non-rocky, somewhat disturbed inland coastal sites between the strand
and the base of the escarpment include Hibiscus tiliaceus, Cocos nucifera, Premna
serratifolia, Calophyllum inophyllum, Pandanus tectorius, Morinda citrifolia, Terminalia
catappa, Ochrosia elliptica, and isolated specimens of Barringtonia asiatica, Thespesia
populnea and Hernandia nymphaeifolia. Shrubby species include Scaevola taccada,
Colubrina asiatica, Abutilon indicum and Phyllanthus societatis; herbaceous species
include Cyperus javanicus, Digitaria setigera, Vigna marina, Ipomoea macrantha and the
ferns, Polypodium scolopendria and Nephrolepis biserrata. Caesalpinia bonduc, Euphor-
bia chamissonis, Sida fallax and Triumfetta procumbens are scarce, but were probably
more abundant in the past.

Mangroves and Coastal Marsh Vegetation

Shallow-water habitats with muddy bottoms and protected from strong wave action
are extremely limited on Nauru. Although reportedly present in the past around Buada
Lagoon, Nauru’s single mangrove species, Bruguiera gymnorhiza, is now restricted to a
system of landlocked brackish ponds or small lagoons near the base of the escarpment in
Menen, Anabar and Anetan Districts. The largest concentration is found around Araro

10

Lake in Anetan. Fosberg (c.1972) also reports the occurrence of B. gymnorhiza in similar
landlocked ponds, sink-holes and small inland swamps in Palau.

Other species commonly associated with mangroves and present in Nauru include
Derris trifolia, encountered on limestone outcrops, and Vitex negundo, which is present
in depressions near the base of the escarpment in Menen District.

The swampy areas surrounding Buada lagoon and near the base of the escarpment
on Nauru are dominated by Cyperus javanicus and C. compressus, with one specimen of
Ludwigia octovalvis collected from a coastal depression.

Relict Stands of Inland Forest

In terms of relict stands of primary inland forest on Nauru, there seem to be two
distinct types: 1) plateau forest, which probably covered up to 90 per cent of the island
before the onset of phosphate mining, and 2) escarpment forest, including forest on
unmined limestone outcrops or pinnacles on the plateau.

The former, four-fifths of which has been removed during phosphate mining, is
dominated almost entirely by 16 m-tall Calophyllum inophyllum. Infrequent canopy trees
include Guettarda speciosa, Premna_ serratifolia and Terminalia catappa, with the
understorey dominated by Scaevola taccada, Morinda citrifolia and Dodonea viscosa, the
parasite Cassytha filiformis, Psilotum nudum, and the ferns, Polypodium scolopendria and
Nephrolepis biserrata. Also occasional in open sites is Phyllanthus societatis. Exotic
species dominant in disturbed sites include Psidium guajava, Lantana camara and two
herbaceous species, Euphorbia hirta and Desmodium triflorum (Manner et al. 1984,
1985).

Limestone Escarpment or Pinnacle Vegetation

The dominant species on the limestone cliffs of the escarpment and on emergent
pinnacles on Nauru is Ficus prolixa, with Terminalia catappa, Ochrosia elliptica and
Guettarda speciosa constituting important second stratum species (Manner et al. 1985).
Isolated relict stands of Barringtonia asiatica and Pisonia grandis are also found along
the crest of the escarpment above Anibare Bay. The ferns, Nephrolepis biserrata and
Polypodium scolopendria, and the liana, Ipomoea macrantha, are locally abundant, and
the herb, Laportea ruderalis, is found in moist shady habitats at the base of the es-
carpment. On the more gradually-sloping colluvial portions of the escarpment almost
impenetrable thickets of Hibiscus tiliaceus are found. Understorey species include
Colubrina asiatica and Tacca leontopetaloides. In Anetan District, in the north (Figure
1), Clerodendrum inerme festoons limestone outcrops and cliffs. In some areas the exotic

11

fruit trees, soursop and sweetsop (Annona muricata and A. squamosa) have become
naturalised and constitute the dominant understorey species.

Coconut-Palm- and Pandanus-Dominated Agricultural Lands

As stressed above, coconuts and pandanus were the two most important staple
crops in pre-European-contact Nauru. Coconut groves, although far less important today
than in the past when copra was the main export, are still found in a number of sites on
Nauru’s coastal strip. In most cases, the plantations are comprised of randomly scattered
trees of varying heights and ages. In poorly-maintained groves, coconut seedlings and
fallen leaves and husks dominate the understorey.

Planted groves of edible Pandanus tectorius cultivars in forest clearings, both on
the plateau and on the more gradually sloping areas of the escarpment, were also once a
prominent feature of Nauru’s traditional agricultural vegetation. Today, they are
restricted to a few relict groves, some of which were found to be present in the unmined
areas of Anibar District in 1980.

Houseyard Gardens and Urban Vegetation

Although indigenous and aboriginally introduced species are important com-
ponents, most houseyard gardens and urban vegetation are dominated by recently
introduced exotic species. The high diversity of recently introduced exotics in Nauru is
due mainly to the almost complete destruction of the subsistence economy, urbanization
and increasing air transport contact with overseas sources of non-quarantined planting
materials.

In indigenous Nauruan houseyard gardens the dominance of recently introduced
ornamental species is very pronounced, with some 118 of 140 species being classified as
recent introductions. Indigenous species include Abutilon indicum, Barringtonia asiatica,
Calophyllum inophyllum, Cerbera manghas, Clerodendrum inerme, Cordia subcordata,
Ficus prolixa, Guettarda speciosa, Morinda citrifolia, Ochrosia elliptica, Premna
serratifolia and Terminalia catappa, which in some cases, such as with Cerbera manghas
and Cordia subcordata, are only present today in Nauru in houseyard gardens. Common
food plants include, in order of importance, coconut, breadfruit, bananas, pandanus,
papaya, Citrus spp. and guava. Common ornamentals, found in at least four of 16 sample
gardens, include Acalypha amentacea vars., Bougainvillea spp., Caesalpinia pulcherrima,
Caladium bicolor, Casuarina equisetifolia, Catharanthus roseus, Codiaeum variegatum,
Cordyline fruticosa, Crinum spp., Delonix regia, Dieffenbachia spp., Hibiscus rosa-
sinensis, Hosta plantaginea, Hymenocallis littoralis, Ixora spp., Jasminum sambac,
Jatropha integerrima, Nerium oleander, Pentas spp., Plumeria spp., Polyscias spp.,
Pseuderanthemum carruthersii, Sansevieria trifasciata, Tabernaemontana divaricata,
Tecoma stans and Thunbergia erecta.

12

The houseyard gardens of I-Kiribati, Tuvaluan, Chinese and Filipino contract
workers, and in the European and Indian expatriate communities of Nauru, are very
different. Each reflect distinctive preferences in food and ornamental plants, and are
commonly dominated by food plants. I-Kiribati and Tuvaluan gardens at Location, where
there is very little space for planting, usually consist of a single banana, coconut, papaya
or breadfruit tree, or a few cassava, sweet potato, taro, tannia (Xanthosoma sagit-
tifolium), pineapple, sugarcane, hibiscus spinach (Hibiscus manihot), or chilli (Capsicum
frutescens) plants. All are often grown in boxed or fenced areas filled with imported soil
or mulch. Chinese gardens at Location focus more on short-term vegetable plants, such
as Chinese cabbages (Brassica spp.), onions and garlic (Allium spp.), amaranth spinach
(Amaranthus spp.), coriander (Coriandrum sativum), long beans (Vigna sesquipedalis)
and a range of cucurbits. Filipino workers plant sweet potato, hyacinth bean (Dolichos
lablab) and horseradish or drumstick tree (Moringa oleifera). European expatriates plant
tomatoes, lettuce and parsley, whereas the expanding Indian expatriate community has
planted eggplant (Solanum melongena), okra (Hibiscus esculentus), horseradish tree
(Moringa oleifera) and bilimbi (Averrhoa belimbi). A similar range of food species are
cultivated behind the workshops on Topside, although the areas under crops are greater,
with some gardeners growing taro, tannia and giant swamp taro, employing the tradi-
tional intensive mulching systems of Kiribati and Tuvalu (Thaman 1987a, 1988b).

The balance of the “urban vegetation" is composed of many the same species
which are occasionally planted as roadside trees or around government and Nauru
Phosphate Corporation buildings and parking lots. The remaining area of extensive urban
vegetation is the golf course in Aiwo District which is lined with trees, including
Hibiscus tiliaceus, Thespesia populnea, but dominated by banyan trees (Ficus spp.).

Ruderal Vegetation

Extensive areas of highly disturbed ruderal vegetation in settlements, waste places,
along roadsides and airstrips, and in areas associated with pre-mining vegetation
clearance are found in Nauru. The dominant species in most areas are pioneering grasses,
annuals and shrubby weedy species.

Common species include; 1) the grasses, Cenchrus echinatus, Chloris inflata,
Cynodon dactylon, Dactyloctenium aegyptium, Digitaria spp., Eleusine indica, Eragrostis
amabilis, Lepturus repens and Tricholaena rosea; 2) the sedges, Cyperus javanicus, C.
rotundus and Fimbristylis cymosa (which, along with Digitaria setigera and Lepturus
repens, are probably indigenous); and, 3) the herbaceous species, Ageratum conyzoides,
Alysicarpus vaginalis, Amaranthus dubius, A. viridis, Bidens pilosa, Cassia occidentalis,
Cleome rutidosperma, C. viscosa, Crotalaria goreensis, C. spectabilis, Desmodium
tortuosum, Euphorbia spp., Hedyotis corymbosa, Indigofera hirsuta, Malvastrum
coromandelianum, Passiflora foetida, Phyllanthus amarus, Physalis spp., Portulaca
oleracea, Sida rhombifolia, Spermacoce assurgens, Stachytarpheta urticifolia, Synedrella

13

nodiflora, Tridax procumbens and Vernonia cinerea, plus the indigenous species,
Phyllanthus societatis and the parasitic Cassytha filiformis.

A few tree or tree-like species have become naturalized in dense stands in Nauru.
Mangifera indica is dominant in dense forests behind residences in the Buada Lagoon
depression; Annona muricata and A. squamosa, as mentioned above, form dense stands
on gradually sloping areas of the escarpment; Adenanthera pavonina, Lantana camara
var. aculeata, Leucaena leucocephala and Psidium guajava form dense stands or thickets
between Buada Lagoon and the decalcination plant on the coastal strip in Aiwo District;
and Casuarina equisetifolia and Muntingia calabura have colonized roadside areas and
portions of the phosphate-mined area. Of particular interest, is the almost monospecific
colonization of the large topsoil stockpile on Topside by the cucurbit, Luffa cylindrica
var. insularum.

Phosphate-Mined Lands

As a result of almost 80 years of open-cast phosphate mining, some three-quarters
of Nauru is under severely-modified disclimax vegetation in various stages of succession.
Prior to mining the vegetation is removed by bulldozer and the topsoil removed to expose
the phosphate deposits which lie between coral-limestone pinnacles. The extraction of
phosphate then causes dramatic changes in local relief, which varies between 4 and 8 m
from the top of the pinnacles to the pit bottoms, with about three to four pinnacles
occurring within each 100 m?. Because mining is only about 20 per cent efficient,
unconsolidated phosphate deposits remain in the pit bottoms and on the saddles and scree
slopes between the pinnacles. These deposits (which might be mined at a later date) and
the pinnacle surfaces, constitute the main sites for recolonization (Manner et al. 1984,
1985).

Although there is widespread evidence that exotics commonly replace indigenous
species in highly disturbed habitats, the Nauru study by Manner, Thaman and Hassall
(1984, 1985) supports the conclusion of Mueller-Dombois (1975) that indigenous
(pioneer) species are often better adapted to edaphically harsh environments, given the
cessation of human disturbance. Their study shows a very rapid colonization of mined
areas by indigenous ferns and exotic herbs, followed by a fairly rapid replacement by
native, primarily coastal strand, species.

Early pioneer species include the non-woody exotics, Alysicarpus vaginalis,
Cleome rutidosperma, Crotalaria goreensis, Emilia sonchifolia, Eragrostis amabilis,
Euphorbia cyathophora, E. hirta, E. prostrata, Hedyotis corymbosa, Synedrella nodi-
flora, Tricholaena rosea, Tridax procumbens and Vernonia cinerea, plus the indigenous
ferns, Nephrolepis biserrata and Polypodium scolopendria. Of these, only the two ferns
species, and Euphorbia hirta, E. prostrata, Tricholaena rosea, Tridax procumbens and
Vernonia cinerea remain significant components of the flora in the later stages of
succession, usually in open disturbed sites.

14

Species entering the succession early and remaining dominants in the 40- to 80-
year-old sites include: the trees, Calophyllum inophyllum, Dodonea viscosa, Ficus
prolixa, Guettarda speciosa, Morinda citrifolia and Premna_ serratifolia; the shrubs,
Phyllanthus societatis and Scaevola taccada; the grasses and sedges, Lepturus repens,
Fimbristylis cymosa and Cyperus javanicus; the parasite, Cassytha filiformis; and the
diminutive fern, Ophioglossum petiolatum, All are indigenous. Larger exotics found in
the later stages of succession in more open habitats include Lantana camara, Psidium
guajava and Stachytarpheta urticifolia.

The study suggests that the potential natural disclimax vegetation of the open-cast
mined plateau will probably be dominated by Calophyllum inophyllum and Guettarda
speciosa, with the epiphytic Ficus prolixa dominating the more ecologically severe
pinnacle habitats. Morinda citrifolia, Premna_ serratifolia and the exotics, Lantana
camara and Psidium guajava, could become important components of the subcanopy. The
exotics, Casuarina equisetifolia (which is native to limestone habitats on other Pacific
islands) and Muntingia calabura, both now locally abundant in some mined areas, could
enter into the succession as well.

As argued by Manner e¢ al. (1985), given no deliberate human intervention, the
succession to a disclimax vegetation association capable of sustaining human life will
probably take "many thousands of years". It is stressed that it is ironic that Nauru’s
central plateau, from which Nauruans formerly obtained some of the necessities of life,
will be a "topographic jungle" stripped of its natural vegetation, before the next century,
in order to provide the phosphate needed to revive phosphate-poor soils to fuel the
development of Australia and New Zealand.

THE FLORA

Like the extremely limited, degraded and displaced vegetation types, the indige-
nous terrestrial flora of Nauru exhibits extreme poverty and current numerical domination
by exotics. Of a total of 493 species or hybrid cultivars reported to have been present on
Nauru, only 59 (12%) are possibly indigenous (Tables 1 and 2). There are no reported
endemics, reflecting the lack of habitat diversity and the predominance of ubiquitous,
easily-dispersed pantropical or paleotropical coastal species. Two species (Achyranthes
canescens and Tarenna sambucina are presumably now extinct, Aidia cochinchinensis
possibly now extinct, and half (28) of the remaining 56 species are severely restricted in
distribution, endangered or possibly extinct, due to removal and severe habitat modifica-
tion or limitation (Table 2). The high number of recent introductions in Nauru reflects its
increasing contact with the outside world via its national airline, Air Nauru, increasing
urbanization and the total absence of quarantine regulations.

15

Table 1. Antiquity status of the flora of Nauru in terms of whether species are presumed
to be indigenous to Nauru; aboriginal or recent post-European-contact introductions; or
now extinct.

Class/Genera Indigenous Aboriginal Recent Extinct Total

Pteridophytes 8 - 3 - 11
Gymnosperms - S 2 - 2

Monocotyledons 6 2 137, - 145
Dicotyledons 45 3 278 9 335
Total 59 5 420 9 493

In terms of diversity at the family level, only 97 vascular plant families are
represented on Nauru (Appendix I). Of these, only 33 are represented by indigenous
species.

The pteridophytes and gymnosperms are represented by six families, all of which
are represented by an indigenous species. The Gymnosperms are represented by only two
families, none of which are indigenous.

The 145 monocotyledons fall into 18 families, only three of which, Cyperaceae,
Pandanaceae and Poaceae, are indigenous. Araceae, Liliaceae and Poaceae are the only
families with over ten separate species (26, 34 and 26 respectively). Other monocotyl-
edon families represented by five of more species include Arecaceae, Commelinaceae,
Cyperaceae, Marantaceae, Orchidaceae and Zingiberaceae. The majority of species from
these families are cultivated ornamentals, or in the case of Poaceae, weedy grasses.

The dicotyledons are represented by 71 families, 25 of which are possibly indige-
nous. Families represented by two or more indigenous species each are Apocynaceae,
Boraginaceae, Capparidaceae, Convolvulaceae, Euphorbiaceae, Fabaceae, Malvaceae,
Rubiaceae and Verbenaceae (see Part II and Appendix I).

16

Table 2. Species indigenous or possibly indigenous to Nauru (? = status uncertain,
possibly an aboriginal or recent introduction; E = possibly extinct; e = endangered or

rare).

Type/Species

PTERIDOPHYTES

Asplenium nidus
Nephrolepis biserrata
Nephrolepis hirsutula
Ophioglossum petiolatum
Polypodium scolopendria
Psilotum nudum

Pteris tripartita

Pyrrosia adnascens

Subtotal

HERBS

Achyranthes canescens
Heliotropium procumbens
Laportea ruderalis
Triumfetta procumbens

Subtotal

GRASSES AND SEDGES

Cyperus javanicus
Digitaria setigera
Fimbristylis cymosa
Lepturus repens
Stenotaphrum micranthrum

Subtotal

Status

bce Ptr a is ae
~~

ca?)

oo

VINES AND LIANAS

Canavalia cathartica
Canavalia rosea
Capparis quiniflora
Cassytha filiformis
Derris trifoliata
Ipomoea littoralis
Ipomoea macrantha
Ipomoea pes-caprae
Vigna marina

Subtotal

SHRUBS

Abutilon asiaticum var. albescens

Caesalpinia bonduc
Capparis cordifolia
Clerodendrum inerme
Colubrina asiatica
Dodonaea viscosa
Euphorbia chamissonis
Phyllanthus societatis
Scaevola taccada

Sida fallax

Suriana maritima

Subtotal

TREES

Aidia cochinchinensis
Barringtonia asiatica
Bruguiera gymnorhiza
Calophyllum inophyllum
Cerbera manghas
Cordia subcordata
Erythrina variegata
Fagraea berteriana
Ficus prolixa

Guettarda speciosa

17

18

Hernandia nymphaeifolia sae
Hibiscus tiliaceus =F
Morinda citrifolia +
Ochrosia elliptica =e
Pandanus tectorius oF
Pisonia grandis ar
Premna serratifolia =F
Tarenna sambucina E?
Terminalia catappa all
Thespesia populnea SHE
Tournefortia argentea ata
Vitex negundo ic
Subtotal 20 (2E)
TOTAL SPECIES 56 (3E)

Sources: An extensive review of the available literature and personal records and
observations by the authors; see in particular Thaman 1992; Manner, Thaman and
Hassall 1984, 1985.

Nature of Indigenous Species

Of the indigenous species, eight are widespread pantropical or paleotropical
pteridophytes, including Psilotum nudum, Polypodium scolopendria, Ophioglossum
petiolatum, Pteris tripartita and Nephrolepis spp., with Pyrrosia adnascens, a common
epiphyte, also present on Nauru (Table 2). Asplenium nidus, although almost certainly
indigenous, was found only as an ornamental in houseyard gardens during the current
study.

There are no indigenous gymnosperms, although the widespread Cycas circinalis is
found in cultivation.

Indigenous monocotyledons are restricted to Pandanus tectorius, some cultivars of
which are undoubtedly aboriginal introductions, and a small range of sedges and grasses
(Cyperaceae and Poaceae), some of which might be aboriginal or recent introductions.
The coconut palm (Cocos nucifera) is classified as an aboriginal introduction. The grass
Stenotaphrum micranthrum (reported present by Fosberg ef al. 1987) is considered to be
endangered or now absent.

19

The dicotyledons are comprised almost exclusively of salt-tolerant, widely-
dispersed, pantropical coastal species. Of the 42 herbaceous and woody dicotyledons, half
(21) are endangered or rare (Table 1). Species such as Heliotropium procumbens,
Laportia ruderalis, Triumfetta procumbens, Abutilon asiaticum, Caesalpinia bonduc,
Euphorbia chamissonis, Sida fallax, Suriana maritima, Aidia cochinchinensis, Bar-
ringtonia asiatica, Cerbera manghas, Erythrina variegata, Hernandia nymphaeifolia,
Pisonia grandis, Thespesia populnea and Vitex negundo are represented by only a few
remaining individuals, often in houseyard gardens, or by localized relict communities.
Prior to widespread disturbance, Nauru would have undoubtedly had more species than it
has at present.

Comparison with Other Island Floras

The extreme poverty of the indigenous flora of Nauru becomes more obvious when
compared with estimates of the indigenous floras of island groups which are larger or
closer to the Asian plant source region. The estimated number of indigenous species for
the following island groups are: Malaya (c. 20,000), Philippines (10,000), Bismarck
Archipelago (700), Vanuatu (750), Fiji (more than 1,100), Tonga (257), Samoa (548) and
French Polynesia (600). Only extremely isolated small islands such as Easter Island and
the three small atolls of Tokelau, with indigenous floras of 32 and 33 species respec-
tively, have poorer floras than Nauru (Good 1947 in Manner 1987, Parham 1971). When
taken individually, the floras of Pacific atolls range from as few as three to perhaps 150
indigenous species, compared to some Indian Ocean atolls, nearer to continental areas,
which have close to 300 indigenous species (Fosberg 1952).

The floristic poverty of Nauru becomes even more pronounced based on a
comparison of the frequency occurrence of 142 widespread coastal species in eleven
Pacific island groups (see Appendix II and Thaman 1992). All of the species considered
have the ability to cope successfully in environments characterized by loose shifting
sands, wave action, soil-less limestone and volcanic terraces and rock outcrops, high
salinity, strong sunlight, strong winds, seaspray and associated physiological drought
(Fosberg 1952, 1960) and, in some cases, periodic inundation and waterlogging, all
conditions common in Nauru.

The island groups analyzed ranged from large, geographically-older high island
groups such as Fiji, composed of over 300 islands with a total area of 18,376 km’, to the
three small isolated atolls of Tokelau with a total land area of only 12.2 km’. Also
included in the comparison were high island groups with diverse habitats, such as Guam,
a volcanic island with extensive areas of limestone and an area of 549 km?; Samoa and
Hawaii, recent basaltic volcanic island groups, with little or no limestone; Palau, a group
of some 340 volcanic and uplifted limestone islands, including an atoll, Kayangel, and
the raised phosphate island of Angaur, located only 850 km to the east of the Philippines;
Tonga, a group of about 150 uplifted limestone and some volcanic islands, with a total
area of 697 km’; Niue, an isolated uplifted limestone island, like Nauru, but with an area

20

of 258 km’, and few beaches; and Makatea, an uplifted phosphatic island about the same
area as Nauru, and like Nauru, with no protective barrier reefs or coastal lagoons
(Thaman 1992).

Although these comparisons are strongly biased by the size and geologic age of the
islands, their distance from plant source areas and the unavailability of information on
other analogous islands such as Banaba (Ocean Island), the poverty of the coastal floras
of the two small phosphate islands of Nauru and Makatea (in the Tuamotu Archipelago of
French Polynesia) is clearly apparent, both with only 55 of the 142 widespread coastal
species each. The Gilberts and Tokelau, both groups of low-lying atolls, have only 74
and 34 species each. All other island groups, including Hawaii and Niue, have at least
two-thirds of the 142 species present as indigenous or long-established introductions, an
indication of greater habitat diversity and/or less habitat degradation (Thaman 1992). Of
interest is that 55 of Nauru’s 59 indigenous species are among the 142 widespread coastal
or mangrove species listed in Appendix I.

As atoll groups, the Gilbert and the Tokelau Islands, have the fewest ferns,
although Nauru has the fewest widespread coastal herbs. It could be assumed that the
fern, Davallia solida, and widespread herbaceous species, such as Boerhavia, Hedyotis
and Portulaca spp. and Sesuvium portalucastrum, were all originally present on Nauru,
but eliminated due to widespread destruction of coastal habitats. In terms of coastal
grasses and sedges, Nauru, the small atolls of Tokelau, and Makatea have the fewest
species, possibly due to the combination of widespread habitat destruction and a relative
absence of marsh or wetland environments. The widespread destruction of the areas
around Buada Lagoon for cultivation by the Japanese during World War II may have
destroyed many natural wetland environments. Species which might have been present
include the grasses Paspalum distichum and Thuarea involuta. Whether a given sedge
species arrived naturally or was introduced deliberately due to its cultural utility is
uncertain.

Coastal vines and lianas are noticeably fewer on Makatea and Nauru, with only
three and eight out of the 14 widespread species. Noticeably absent on Nauru are Abrus
precatorius, Entada phaseoloides and Mucuna gigantea.

With the exception of the Tokelau Islands (with only three species), Nauru and
Makatea have the poorest shrub flora, with only 10 and 11 species respectively, out of a
possible 27 species. Similarly, the Tokelaus have only 16 of 62 common coastal tree
species, with Nauru and Makatea having only 23 and 21 species respectively. Conspicu-
ously absent on Nauru are the shrubs, Allophylus timoriensis, Pemphis acidula, Sophora
tomentosa and Wollastonia biflora; and the trees, Ficus tinctoria, Neisosperma op-
positifolia, Pipturus argenteus and Terminalia samoensis, most of which are present on
the smaller elevated phosphate-rich island of Fais and the coral-limestone island of
Satawal in the western Caroline islands (Fosberg and Evans 1969).

21

There are other widespread non-coastal species which might have been present in
the past on Nauru, which were present in 1932 (before mining, which ceased in 1966,
almost completely destroyed the island’s inland vegetation) on the analogous phosphate
island of Makatea, which is located much further from the centers of plant diversity than
Nauru. These include: the fern, Ophioglossum pendulum; orchids, such as Oberonia and
Taeniophyllum spp.; the herb, Procris pedunculata; the vines, Abrus precatorius
(mentioned above) and Dioscorea bulbifera (the most widespread of all yam species and
present, in many cases probably as an aboriginal introduction, from East Africa to
Micronesia and eastern Polynesia)(Stone 1970); and the shrubs and trees, Alyxia sp.,
Canthium barbatum, Celtis paniculata, Glochidion ramiflorum, Ixora sp., Melochia
odorata, Planchonella (Pouteria) sp., and Timonius sp. (Wilder 1934). Tarenna sam-
bucina, also present on Makatea, was reported present on Nauru by Burges in 1933, but
is now considered to be extinct.

Makatea has one presumably endemic species, Euprichardia vuylstekeana, and it
might be expected, given the diversity of pre-mining microhabitats and the isolation of
Nauru, that there could have been endemic species there also. Interestingly, the one plant
that was thought to be possibly endemic on Nauru turned out to be Phyllanthus societatis,
"a common plant among coral rocks" on Makatea in 1933 (Wilder 1934).

A similar comparison with the relatively undisturbed flora of Henderson Island, a
remote raised limestone island with a similar limestone plateau and pinnacle topography,
provides further insight into some of the plant species that might have existed in the past
on Nauru. Because it was unsuitable for permanent habitation and had no economic
phosphate deposits, Henderson has survived successive Polynesian and European impacts,
with only five known introduced plant species. Species of widespread genera found on
Henderson, but not on Nauru include: the fern, Davallia solida; the herbs, Boerhavia
tetrandra, Euphorbia sparrmannii, Lepidium bidentatum, Peperomia hendersonensis,
Portulaca lutea, Procris pedunculata and Sesuvium portalucastrum; the shrubs, AI-
lophylus sp., Alyxia sp., Canthium barbatum and C. odoratum, Glochidion pitcairnense,
Ixora fragrans, Jasminum didymum, Eugenia reinwardtiana (Eugenia rariflora), Morinda
umbellata var. forsteri, Pemphis acidula, Timonius polygamus and Xylosma suaveolens
var. haroldii; and the trees, Celtis paniculata var. viridis, Geniostoma hendersonense,
Meryta brachypoda, Pittosporum arborescens, Sesbania coccinea and Santalum hender-
sonense. Nine species or varieties are presently recognized as endemic, all of which are
found in the interior. These are the very areas on Nauru that have been so devastated by
phosphate mining, long settlement and devastation and scavenging for food and firewood
during the Second World War. Not present on Henderson, but present on Nauru are
Calophyllum inophyllum and Barringtonia asiatica (Paulay and Spencer 1989, Fosberg et
al. 1983, Fosberg et al. 1989).

22

Nature of Exotic Species

Exotic species, which constitute 88 per cent (434 out of a total of 493 reported
species) of the flora of Nauru, dominate ruderal, houseyard and urban vegetation. Exotic
species include a wide range of ornamentals, weedy species, food plants and a number of
other useful species.

Ornamentals, which are normally confined to houseyard and village gardens,
comprise some 59 per cent (257) of the 434 exotic species. On Nauru, introductions by
travellers from Australia, Fiji and other areas with highly developed ornamental gar-
dening traditions; the absence of quarantine restrictions; and the almost total breakdown
in the subsistence economy, seem to be the main reasons for the disproportionate
importance of ornamental plants. Some of these ornamental, of course, have other uses
such as living fencing or for the preparation of medicines or garlands.

The proportions of the exotic flora composed of weedy species is 18 per cent (80
of 434 species), an indication of both the poverty of the competitive indigenous flora and
the highly disturbed nature of the vegetation.

Although food plants represent 16 per cent of the exotic flora, due to the harsh
environment, limited land area and limited focus on food production in Nauru, many of
these species are restricted in numbers or utility and are often represented by experimen-
tal attempts to diversify food production or by individual, often immature specimens of a
given species. Exotic food plants of particular importance on Nauru include numerous
edible pandanus cultivars (Pandanus tectorius), some of which are undoubtedly aboriginal
introductions, and the coconut (Cocos nucifera), also an aboriginal introduction. Recent
introductions of more localized importance, or of particular importance to contract
worker communities on Nauru include: the vegetables, hibiscus spinach (Hibiscus
manihot), Chinese cabbage cultivars (Brassica spp.), long beans (Vigna sesquipedalis),
amaranthus spinach (Amaranthus spp.) and pumpkin (Cucurbita pepo); the staple root
crops, taro (Colocasia esculenta), tannia (Xanthosoma_ Ssagittifolium), sweet potato
(Ipomoea batatas) and cassava (Manihot esculenta); a range of banana and plantain
cultivars (Musa cultivars); and the tree crops, lime (Citrus aurantifolia), guava (Psidium
guajava), mango (Mangifera indica), soursop (Annona muricata) and the horseradish or
drumstick tree (Moringa oleifera), all of which seem to do well in Nauru’s harsh
environment. Important emergency or pig foods include Polynesian arrowroot (Tacca
leontopetaloides) and purslane (Portulaca spp.), both of which are found as naturalised
plants in the coastal vegetation of in ruderal sites.

Other useful exotic species include kapok (Ceiba pentandra), cotton (Gossypium
barbadense), tobacco (Nicotiana tabacum), and bamboo (Bambusa vulgaris), which were
all reportedly more abundant in the past. As suggested above, some larger weedy exotics,
such as Adenanthera pavonina, Annona spp., Casuarina equisetifolia, Lantana camara,
Leucaena leucocephala, Mangifera indica, Muntingia calabura and Psidium guajava,
some which are classified as ornamentals, food plants or other useful plants, have

23

become naturalized and competitive with the indigenous species in some disturbed and
relatively undisturbed sites.

ECOLOGICAL AND CULTURAL UTILITY OF EXISTING FLORAS

Although highly disturbed, outnumbered and, in some ways, "enriched" by
introduced exotics, the vegetation and flora of Nauru still constitute a critical ecological
and cultural resource to the people of Nauru. This is particularly true for the indigenous
species, virtually all of which had wide cultural utility within the traditional subsistence
economy.

In terms of the more specific ecological attributes of Nauru’s plant resources, the
most important functions include the provision of shade and animal and plant habitats,
protection from wind, erosion, flood and saltwater incursion, land stabilization, protec-
tion from the desiccating effects of salt spray, soil improvement and mulching.

In terms of more strictly cultural utility, preliminary analyses indicate 169 purposes
or use categories for 39 indigenous species, an average of 4.3 uses per species. There are
434 uses for 354 exotic species, an average of 1.2 uses per species (Table 3). This gives
a combined total of 603 use/purpose categories for 393 species (1.5 uses per species).
Twenty (20) indigenous and 80 exotic species had no reported uses. The relative
importance of the indigenous flora would undoubtedly be much more pronounced if: 1) a
more systematic in-depth survey of the cultural utility of each indigenous species had
been conducted; 2) Nauru had not experienced such widespread devastation of its
population, traditional economy, traditional education system and its indigenous flora
(and associated ethnobotanical knowledge) over the past 100 years; and 3) planted
ornamentals, by far the most widespread use of exotic species, were excluded from the
analysis of indigenous species.

Table 3. Frequency of use for specified purposes of plant species present in Nauru (Note:
Introduced includes both aboriginal introductions such as coconut and recent post-
European-contact introductions).

Purpose/Use Indigenous Introduced Total

x/59 x/434 x/493

Cultivated Ornamentals 9 Doi 266
Food Plants 2 64 66
Body Ornamentation 16 23 39
Medicinal/Health 17 13 30
Staple Foods l 13 14

24

—
—
—=
Ww

General Construction
Scenting Oil/Perfumery
Firewood/Fuel
Emergency/Famine Foods
Tools/Utensils
Boat/Canoe Building
Handicrafts

Games/Toys

Food Parcelization
Living Fences/Hedges
Cordage/Fibre

Hair Conditioner
Woodcarving
Adhesive/Glue/Caulking
Earth Oven Cover
Magic/Sorcery
Drinks/Beverage

Fishing Equipment
Clothing

Animal Feed

Plaited Ware
Legends/Mythology
Furniture

Animal Cages/Roosts
Fish Poisons

Fire by Friction
Strainers/Filters
Thatching/Roofing
Dyes/Pigments
Nets/Traps

Fans 1
Chewing Gum/Masticants -
Oils/Lubricants
Corks/Stoppers l
Other Uses* 12

—
Ww

ho WNN RR IE RK ON RANNY

NNR RK KK NRK WNWNWN! PNK NAW WK ND ~) CO WAN OO
a OO NO ee

DONYNNYWNNNNNNWWWARAAHRHRANANADIIIWMDWMWOS

aArren

TOTAL 169 434 603

NO USES 20 80 100

* Other uses include aphrodisiacs, appetite stimulants, brushes, toilet paper, il-
lumination, soap/shampoo, containers, deodorants/air fresheners, fishnet floats, green

25

manure, groundcover, meat tenderizer, insect repellents/fumigants, love potions, wild
animal food, fishing bait, cigarette wrappers and tobacco.

Moreover, if distinct uses within use/purpose categories (e.g., tools with distinct
functions, different types of fishing equipment, foods or ornamentation for different
occasions or purposes, medicines for different ailments, or plants used for specific parts
of boats or houses) are counted (see individual uses for each species as detailed in Part
II), the economic and cultural utility of plants becomes even more pronounced. The
coconut (Cocos nucifera) palm, for example, has 33 reported uses in Nauru (Table 4),
almost undoubtedly a gross underestimate, in light of at least 128 reported uses (many of
which are almost ubiquitous) for the coconut palm throughout the Pacific Islands
(Thaman 1992). Next in order of importance, are 19 species, all with 5 or more reported
uses. These include, in order of importance, Hibiscus tiliaceus, Pandanus tectorius,
Scaevola taccada, Morinda citrifolia, Guettarda speciosa, Calophyllum inophyllum,
Cordia subcordata, Terminalia catappa, Artocarpus altilis, Thespesia populnea, Tour-
nefortia argentea, Premna serratifolia, Triumfetta procumbens, Vitex negundo, Ochrosia
elliptica, Cassytha filiformis, Musa ABB Group, Bambusa vulgaris and Carica papaya.
Of these 20 species, only Cocos nucifera, Artocarpus altilis, Musa ABB Group, Bambusa
vulgaris and Carica papaya, are aboriginal or recent introductions. The rest are probably
indigenous or very early aboriginal introductions (e.g., some authorities suggest that
species such as Hibiscus tiliaceus, Morinda citrifolia, Cordia subcordata and Terminalia
catappa might have been aboriginal introductions into some Pacific islands because of
their cultural utility.

Another 13 species, 7 of which are indigenous (Dodonea viscosa, Hernandia
nymphaeifolia, Plumeria rubra, Psidium guajava, Erythrina variegata, Bruguiera
gymnorhiza, Barringtonia asiatica, Vigna marina, Clerodendrum inerme, Gardenia
taitensis, Jasminum sambac, Crinum asiaticum and Hibiscus rosa-sinensis), have at least
3 uses each. Another 22 species have at least two reported uses each (Table 4). There is
some usage overlap between categories, such as supplementary and emergency foods,
medicinal, magical, ceremonial and body ornamentation plants, or plants used for
handicrafts, woodcarving, cordage and clothing. Conversely, the categories could be
further broken down to yield an even greater list of uses. Moreover, the list does not
include the more strictly ecological functions of coastal plants, such as shade, protection
from wind, sand and salt spray, erosion and flood control, coastal reclamation, animal
and plant habitats, and soil improvement, all of importance, particularly on an ecological-
ly devastated post-mining Nauru.

26

Table 4. Species of particular cultural utility on Nauru based on an analysis of different
individual uses listed under each species in Part II (Notes: 1) uses do not include a wide
range of ecological functions or uses; A = probably an aboriginal introduction; R =
recent post-European-contact introduction; all undesignated species are possibly indige-

nous).

Latin Name

Cocos nucifera (A)
Hibiscus tiliaceus
Pandanus tectorius
Scaevola taccada
Morinda citrifolia
Guettarda speciosa
Calophyllum inophyllum
Cordia subcordata
Terminalia catappa
Artocarpus altilis
Thespesia populnea
Tournefortia argentea
Premna serratifolia
Triumfetta procumbens
Vitex spp.

Ochrosia elliptica
Cassytha filiformis
Musa ABB Group (R)
Bambusa vulgaris (R)
Carica papaya (R)
Dodonea viscosa
Hernandia nymphaeifolia
Plumeria rubra (R)
Psidium guajava (R)
Erythrina variegata
Bruguiera gymnorhiza
Barringtonia asiatica
Vigna marina
Clerodendrum inerme
Gardenia taitensis (R)
Jasminum sambac (R)
Crinum asiaticum (R)

Hibiscus rosa-sinensis (R)

Uses

PHWOWKHWWHKWWARARARAUNNUNUNUNANNNNADARWOSLHONHO

27

Note: Species with two (2) recorded uses each: Polypodium scolopendria, Cyperus
Javanicus, Abutilon asiaticum, Ficus prolixa, Pisonia grandis, Sida fallax, Ipomoea pes-
caprae, Spondias dulcis (A), Ageratum conyzoides (R), Asclepias curassavica (R),
Bougainvillea spp. (R), Cassia occidentalis (R), Catharanthus roseus (R), Citrus
aurantifolia (R), Delonix regia (R), Ixora casei (R), Lantana camara (R), Mangifera
indica (R), Mirabilis jalapa (R), Ocimum basilicum and O. sanctum (R).

In terms of specific uses, cultivated ornamentals and food plants are by far the
most common. If, however, only indigenous plants are considered, the most widely
reported uses are for medicine, body ornamentation, general construction, cultivated
ornamentation, boat or canoe building, tools or utensils, handicrafts, scenting coconut oil
or perfumery, firewood or fuel, games and toys, hair conditioners, food parcelization,
woodcarving, covering or insulating the earthen oven, magic or sorcery, emergency or
famine foods, cordage or fibre, clothing, plaited ware and furniture.

It must be stressed that the analyses are based on traditional uses, many of which
have lapsed or are only employed in emergency, because modern technology has pre-
empted them. Modern medicine, clothing, fishing lines, matches, crockery, plastic bags,
soap, and emergency food rations (food aid) have, for example, replaced traditional
plant-derived products in Nauru. Moreover, many of the current generation, schooled in
the modern educational system and living in the cash economy, often know few of the
traditional uses of plants, let alone their vernacular names . . . a state which could be
referred to as "devegetation of the mind" . . . and which has undoubtedly contributed to
the degradation of the indigenous and long-established aboriginal vegetation in Nauru.

Of particular note is the importance of traditional food and beverage crops, the
abandonment of which, for highly imported foods such as sugar, white rice and flour,
cabin biscuits, noodles canned fish, soft drinks, alcohol and tea, has led, as stressed
above, to dangerous levels of food dependency and some of the highest, or most rapidly
increasing, incidences in the world of vitamin and mineral deficiency and nutrition related
diseases. Diseases such as iron-deficiency anemia, vitamin-A-deficiency-induced night
blindness, diabetes, cardiovascular disease, hypertension and stroke, gout and hyper-
uricemia, some forms of cancer and dental disease, which were rarely encountered in the
past are now serious causes of morbidity and mortality in Nauru, and among other
Pacific island populations (Speake et al. 1979; Coyne 1984; Taylor 1983; Zimmet et al.
1977, 1978; Thaman 1982, 1983, 1988a).

CONCLUSION

The vegetation and flora of Nauru are among the most impoverished, degraded,
disturbed and displaced in the Pacific islands. Long habitation, almost a century of open-
cast phosphate mining, continuous bombing, destruction and displacement of the people
during World War II, rapid urbanization and the abandonment of agriculture and

28

subsistence activities have arguably produced one of the most severely modified natural
and cultural floras on earth. }

Although both the vegetation and limited indigenous flora of Nauru has been
severely degraded and outnumbered by exotic species, many of the native species are still
present, unfortunately often in an endangered state. The indigenous flora also still
dominates most habitats, including the later stages of the phosphate-mined pit and
pinnacle topography of Nauru. Even in ruderal habitats and in houseyard gardens and
villages, where they are outnumbered by exotics, indigenous species constitute important
components.

It is argued that, while floristic degradation in Nauru appears to be among the
most severe in the Pacific, the current flora still constitutes an important ecological and
cultural resource that must be protected as part of the development process, and NOT as
an afterthought. Even in the case of Nauru, it may not be too late.

29

PART I
A COMPILATION OF THE VASCULAR FLORA OF NAURU

Part II, "A Compilation of the Vascular Flora of Nauru", consists of a listing of,
and relevant information on, the 493 vascular plant species or distinct cultivars and six
widely recognised varieties of common species reported to have been present at some
time on Nauru.

It begins with Pteridophyta (ferns and fern allies), followed by Gymnosperms and
then Angiosperms. Within Angiosperms, Monocotyledons precede Dicotyledons. Under
these headings individual families are listed in alphabetical order (e.g. Acanthaceae,
Amaranthaceae, Anacardiaceae . . .), with individual species being listed in alphabetical
order by genus within each family (e.g. Asystasia gangetica, Barleria cristata, Barleria
prionitis, Blechum brownei . . .).

KEY

Under each species/entry the types of information and order of presentation is as
follows: 1) Latin or scientific name; 2) common name (s); 3) local vernacular names used
by the main ethnic groups in Nauru; 4) synonyms for the Latin or scientific name; 5)
antiquity status of the species, i.e., whether it is indigenous to Nauru, an aboriginal
introduction or a recent introduction to Nauru; 5) geographical origin of the species; 6)
abundance or frequency of occurrence; 7) description of the species; 8) habitat or
distribution in Nauru; 9) uses or cultural utility; and 10) an indication of the persons who
have recorded or collected a given species, including numbers corresponding to her-
barium specimens. This information, its organization and the symbols used under each
category are explained below.

Latin/Scientific Names

ie The first name listed in bold print is what the authors consider to be the currently
most widely accepted published Latin binomial for a given species (usually the
earliest published name or basionym). All names follow the International Code of
Botanical Nomenclature.

D: The Latin names provided in italics after the common and the vernacular names
include Latin binomial synonyms (syns.) or older names no longer in use for the
species, and, in some cases, incorrect names commonly applied to the species,
which are indicated by sensu auct. non.

30

4. The name (s) or the abbreviation of the name (s) of the authority or authorities
(persons responsible for describing and publishing a given species name) are
provided after each species name, e.g., (L.) Anders.

Family

ile Family names (e.g., POLYPODIACEAE, ACANTHACEAE or RUBIACEAE)
are centered in bold capitals immediately before the first species entry in each
family.

2D: Where a family is known by two different names or a species placed in either of

two families, both are listed (e.g., FABACEAE OR LEGUMINOSAE, POA-
CEAE OR GRAMINAE or CLUSIACEAE OR GUTTIFERAE)

Common Names

i English or common names for a species, and other widely-used names, are
included in quotation marks, e.g., "coconut", immediately to the far right of the
Latin name.

Vernacular Names

ll. The vernacular names include the Nauruan names and other names used by the
dominant ethnic communities of Nauru. These are found on the second line, after
the common English names.

2. The letter (B) after a Nauruan name indicates names listed by Burges (1933). All
other names were collected by Thaman, Manner and Hassall as part of the current
study.

Se N, K, T, C, SI, Philippines and Hindi, are used in parentheses to indicate the
Nauruan, Kiribati, Tuvaluan, Chinese (Cantonese), Solomon Island, Filipino and
Hindi names, respectively.

4, The question mark (?) designates unverified or doubtful names.

3. In terms of pronunciation, the Nauruan phonetics are difficult to match with the
accepted sounds and orthography of the Latin alphabet. The closet approximations
of the correct Nauruan pronunciation of a given name are provided instead of
resorting to the use of strange combinations of letters or special phonetic symbols.

6. In the cases of other vernacular languages used in Nauru, the ‘g’ in Tuvaluan is
pronounced as if it were like ‘ng’ as in the word ‘sing’; in Kiribati the t, when

31

followed by an i, is pronounced like an s, and when ti falls at the end of a word,
the i remains silent, e.g., ‘roti’ is pronounced as if it were ‘rose’.

Antiquity Status

Antiquity status indicates whether a given species is presumed to be indigenous to
Nauru; an aboriginal introduction by Nauruans or other indigenous Pacific Islanders
before European contact; or a post-European-contact introduction. In some cases it is
suggested that a species may have been successfully introduced prior to European
contact, but either not successfully established or brought to extinction before botanical
collections or observations of the flora were made. In the case of recent introductions,
some species are categorized as to whether they are assumed to be pre- or post-World
War II introductions. This is based on information received from informants and/or
whether a species was reported present before World War II by Burges in 1933. The ?
indicates that the true status of a species is in doubt (e.g., whether it is really indigenous
or an aboriginal introduction).

Geographical Origin

Geographical origin refers to what seems to be the original natural distribution of
a given species before humans began to acts as dispersal agents for plants. In many cases
it is difficult to be sure what the original pre-human or pre-European-contact range of a
given species was because species introduced either deliberately or accidentally by the
Pacific Island colonizers of the islands have often become naturalized and integral
components of what now seems to be indigenous vegetation.

With respect to terminology, Malesia (sometimes spelled Malayasia) is a biogeo-
graphical term referring to an area encompassing insular southeast Asia, the Indonesia,
Philippines and the island of New Guinea; Indomalaysia refers to an area encompassing
the Indian Ocean and Malesia; Indopacific refers to an area extending from the Indian
Ocean to the Pacific Islands; Paleotropics refers to the Old World tropics including
tropical Africa, Asia and the tropical Pacific Islands; pantropical indicates that a species
is found throughout the Old and New World tropics; and cosmopolitan indicates that a
species is found almost worldwide. In some cases (usually in the cases of easily dispersed
weedy pioneer species) information is provided on both the assumed original distribution
and whether a species is now more widespread (e.g., pantropical).

Abundance or Frequency Occurrence
The estimates of abundance or frequency occurrence or whether a given species is

now endangered or extinct are based on in-the field observations by Thaman, Manner and
Hassall from 1979 through 1987, plus information included in herbarium vouchers of

82

other collectors. In-the field data included in-depth analysis of all vegetation associations,
transects at ten locations around the coastal plain, transects across the entire island and
extensive sampling using 100 m? quadrats in areas where open-cast phosphate mining had
occurred.

Species Description

Descriptions of individual species (and selection of the most widely accepted
scientific names and authorities) were based on the analysis of living plants in Nauru,
herbarium specimens and a synthesis of the best existing descriptions in the works listed
in the Bibliography. Works of particular value included Fosberg, Sachet and Oliver 1979,
1982, 1987; Haselwood and Motter 1976; Hay, McQuown, Beckett and Beckett 1974;
Henderson and Hancock 1989; Henty and Pritchard 1973; Herklots 1972; Macoboy 1969,
1979; Neal 1965; B.E.V. Parham 1972; J.W. Parham 1972; Purseglove 1972, 1974;
Rotar 1968; St. John 1973; Smith 1979, 1981, 1985, 1988, 1991; Stemmermann 1981;
Stone 1970; Sykes 1970; Whistler 1980, 1983, 1992; Wilder 1934; Wright, Minter and
Carter 1984; and Yuncker 1959.

Uses and Cultural Utility

Ethnobotanical information (including vernacular names) on uses or cultural utility
of the individual plant species was obtained through in-depth interviews with elderly
person known for their knowledge of the traditional uses of Nauru’s plants; other
respondents, whenever possible; and from information in available documents and
publications. Main informants included Joseph D. Audoa, James Aingimea, Henry
Michael Heine, Daphne Fotu, Jacob Gabwinare, Katarina Satto, Kenia Raidinen, Reynold
Capelle, Eda Adam and Montiba Star.

Collectors and Herbarium Specimens

The numbers listed at the end of the information on each species indicate which
collectors or observers collected or recorded that species as being present on Nauru; the
numbers in parentheses identify the numbers of the herbarium vouchers or specimens
collected by each collector (s), e.g., 2, 3(58802), 4(168N), 5(92), 6, 7(27812).

The numbers and the collectors/observers are as follows:

1 refers to citations by persons such as Finch prior to 1900, about which there is
very little information;

2 refers to citations by Alan Burges of Sydney University who collected in 1933
and most of whose specimens are lodged at Kew Botanical Gardens, London;

33

3 refers to citations by F.R. Fosberg of the National Museum of Natural History
of the Smithsonian Institution, Washington D.C. in 1980, whose specimens are
lodged with the Smithsonian Institution;

4 refers to Brian Scully of the University of California at Riverside who also
collected in 1980 and most of whose specimens are also lodged with the Smith-
sonian Institution;

5 and 6 refer R. R. Thaman, H.I. Manner and D.C Hassall of The University of
the South Pacific, Suva, Fiji who collected over two 2-week periods in November
1980 and July 1981, respectively, and whose specimens are lodged with the South
Pacific Regional Herbarium at The University of the South Pacific, Suva;

7 refers to Thaman and Manner (then with the University of Guam) who collected
again in July-August 1987 and whose specimens are also lodged at the South
Pacific Regional Herbarium; and,

8 refers to John Swarbrick, of the University of Queensland, Gatton College who
collected weedy species on Nauru in June 1988, and whose specimens are lodged
in Queensland with duplicates at the South Pacific Regional Herbarium..

When possible, F.R. Fosberg of the Smithsonian Institution examined and verified
the identifications of all herbarium specimens. These include some specimens collected in
the early 1980s by Dr. Lynn Raulerson of the University of Guam.

34

VASCULAR PLANTS OF NAURU

PTERIDOPHYTA (Ferns and Fern Allies)
ASPLENIACEAE

Asplenium nidus L. "bird’s-nest fern"
laulu, laukatafa, laukatapa (T)
syns. Neottopteris nidus (L.) Sm.; N. curtisorus (Christ) Hosok.; Asplenium
curtisorum Christ; A. phyllitidis D. Don; Neottopteris phyllitidis (D. Don)
J. Sm.

Indigenous? Paleotropical. Rare. Large epiphytic fern with a short creeping
rhizome; fronds, up to 1 m or more long and 15 cm wide, simple, bright green,
elongated, swordlike, with prominent midribs, forming a rosette-like cluster; sori
numerous, linear, borne on distal parts of fronds, reaching nearly to the margin.
Reported by Burgess as an epiphyte on Calophyllum inophyllum in 1935, but seen only as
a ornamental by Thaman, Hassall, and Manner in 1980 and 1981. 2, 5, 6, 7(27813).

DAVALLIACEAE

Nephrolepis biserrata (Sw.) Schott "sword fern"
dakeang, dageang (N); te keang, te keang ni Makin (K); sulufe (T)
syns. Aspidium biserratum (Sw.); Nephrodium biserratum (Sw.) Gaud.; N.
splendens (Willd.) Gaud.; Aspidium splendens Willd.

Indigenous. Pantropical. Abundant. Terrestrial tufted fern with short scaly
creeping to erect rhizome; stipes, up to 30 cm or longer, clustered; fronds, up to 2 m or
longer and 30 cm wide, oblong-elliptic in outline, pinnate; pinnae up to 15 cm or longer
and 2 cm wide, margins usually serrate, apex acuminate; sori, large, in a row at a
distance from the margin (submarginal). Found in colonies and dense populations in
unmined areas and in pits between pinnacles in mined areas; one of first plants to
colonize mined areas; occasional as an ornamental. Leaves used occasionally in garlands.
3(58600), 4(136N), 5(44), 6, 7.

35

Nephrolepis exaltata (L.) Schott "Boston fern"
syn. Polypodium exaltatum L.

Recent introduction. Pantropical. Occasional. Fern with graceful spreading fronds.
Ornamental pot plant. 3, 5(86), 6(181).

Nephrolepis hirsutula (Forst.f.) Presl "sword fern", "fishtail fern"
dakeang, dageang (N); te keang (K); sulufe (T)
syns. Polypodium hirsutulum Forst.f.; Nephrodium gibbosum (Willd.) Gaud.;
Aspidium gibbosum Willd.

Indigenous. Indopacific. Occasional. Terrestrial fern with a creeping rhizome;
stipes, clustered, scaly; fronds, up to 50 cm or more long, pinnate; pinnae, up to 10 cm
or more long and 1 cm wide, sessile, oblong-lanceolate, serrullate, obtuse or acutish,
pubescent; sori, large, round, submarginal. "Abundant amongst moist rocks." 8 (9586).

OPHIOGLOSSACEAE (Adder’s Tongue Fern Family)

Ophioglossum petiolatum Hook. "adder’s tongue fern"

Indigenous. Pantropical. Occasional. Diminutive erect terrestrial fern, about 6 cm
high, with creeping rhizomes; fronds, ovate to rhombic, 2.5 to 6 cm long. Found as
scattered individuals in sandy open and partly shaded areas, primarily in older strip-
mined areas on floors of pits between pinnacles. No reported use. 5(151), 6.

POLYPODIACEAE

Polypodium scolopendria Burm.f. "lawai fern" (Hawaii)
dakeang, dageang (N); te keang (K); maile (T)
syns. Phymatodes scolopendria (Burm.) Ching; Polypodium phymatodes L.;
Phymatodes phymatodes (L.) Maxon; Tectaria phymatodes (L.) Cav.;
Polypodium grossum Langsd. & Fisch.; P. hemionitis Cav.; Microsorum
scolopendria (Burm.f.) Copel.

Indigenous. Paleotropical. Very abundant. Terrestrial and epiphytic fern with a
stout creeping rhizome; stipes, up to 30 cm or longer; fronds, up to 40 cm or more long,
broadly triangular-ovate in outline, simple but deeply and broadly divided into lanceolate
acute lobes; sori, large, commonly in two rows on lower surface of lobes. Found in

36

colonies and dense populations in unmined areas, in pits between pinnacles in mined
areas, and on escarpment and cliffs below the plateau. Fragrant fronds used for making
garlands, leis, and other ornamentation and boiled in coconut oil to scent it. 3(58594),
4(121N), 5(43), 6, 7(27811), 8(9578A).

Pyrrosia lanceolata (L.) Farw.
syns. Arostichum lanceolatum L.; Pyrrosia adnascens (Sw.) Ching; Cyclophorus
lanceolatus (L.) Alst.; C. adnascens (Sw.) Desv.; Polypodium adnascens
Sw.; Niphobolus varius Kaulf.; Cyclophorus varius (Kaulf.) Gaud.;
Pyrrosia varia (Kaulf.) Farw.

Indigenous. Trop. Asia to Polynesia. Rare. Small inconspicuous epiphytic
dimorphic fern with a long creeping slender rhizome; fronds at intervals along slender
rhizomes; sterile fronds, 4 to 10 cm long and 1 to 1.5 cm wide, simple, oblong-elliptic
or oblanceolate; fertile fronds, up to 15 cm or longer and about 7 mm to 1 cm wide;
sori, small, densely packed together on lower surface. Growing on tree. 6(173).

PSILOTACEAE

Psilotum nudum (L.) Beauv. "psilotum", "reed fern"
ibiribir (N); te kimarawa (K)

Indigenous. Tropics and subtropics. Occasional. Small erect terrestrial perennial
herb, up to about 30 cm or higher, arising from a stout rhizome, with many successively
two-forked green 3-angled, longitudinally-ribbed branches, about 2 mm in diameter;
leaves, minute, scale-like; sporangia, about 1 mm in diameter. axillary, subglobose,
three-lobed. Found as scattered individuals and small clusters in shady areas under
unmined vegetation on the central plateau and uncommon under trees and shrubs on
escarpment. No reported use. 2(53.5), 3(58764, 58596), 5(102), 6, 7(22314).

PTERIDACEAE

Adiantum sp. "maiden-hair fern"

Recent introduction. Occasional. Fern with finely-dissected fronds and wiry black
stalks. Small ornamental fern grown as a pot plant. 5, 6.

37

Pteris ensiformis Burm.f. "sword brake"

Recent introduction. Tropical and subtropical Asia to Polynesia. Small dimorphic
fern, 15 to 50 cm high, with a brown scaly rhizome; stipes, up to 30 cm or longer straw-
colored; sterile fronds, 6 to 10 cm long, blades lanceolate, smooth, bipinnate, pinnules
variable in size and shape, mostly oblong to linear-lanceolate, sharply serrate; fertile
fronds, 15 to 25 cm long, pinnate or bipinnate, pinnules, up to 15 cm long and 7 to 10
mm wide, widely-spaced, linear; sori, marginal, near serrate. Rare. Ornamental pot
plant. 6(172).

Pteris tripartita Sw. "sword brake"
dakeang, dageang (N)
syn. P. marginata Bory

Indigenous. Paleotropical. Occasional. Large terrestrial fern with a short rhizome;
stipes, up to 60 cm or longer, smooth, brown; fronds, up to 1 m or longer, tripartite,
deltoid, each part divided into lanceolate pinnae; sori forming a continuous row along
margins of pinnae. Found as individuals or isolated clusters at base of limestone cliffs of
escarpment and in waste places near cliff base; occasionally a planted ornamental.
4(135N), 5(55), 6, 7(27815).

GYMNOSPERMAE (Gymnosperms)

ARAUCARIACEAE (Araucaria Family)

Araucaria heterophylla (Salisb.) Franco "Norfolk Island pine"
syn. A. excelsa (Lamb.) R. Br.

Recent introduction. Norfolk Island. Infrequent. Stately symmetrical tree with
about 5 horizontal or drooping branches, each with many branchlets, radiating from each
tier; leaves, up to about 1.2 cm or longer, evergreen, stiff awl- or scale-shaped,
narrowly-triangular, overlapping, borne on branchlets; cones, 7.5 to 10 cm in diameter,
ovoid, woody; pollen borne in catkins, about 5 cm long, which develop singly at the ends
of branchlets. Planted immature ornamental trees in home gardens near airport. 5, 6, 7.

38
CYCADACEAE (Cycad Family)

Cycas circinalis L. "cycad", "sago palm"
te bam (K); laupama, laukimoa (T)
syn. C. rumphii Miq.; C. seemannii (A. Br.) Schuster; C. undulata Desf.

Recent introduction. Trop. Asia, Australia, and the Pacific islands. Occasional.
Small dioecious palm-like rarely-branching plant, up to 2 m or taller, with a sturdy
brown-ringed trunk; leaves, up to 1 m or longer and 30 cm or more wide, clustered in a
rosette at the crown, frondlike, smooth, pinnate; pinnae, up to 30 cm long and | to 2 cm
wide, narrowly lanceolate, with prominant midribs; male inflorescence, up to about 50
cm by 10 cm or more, borne in center of leaves, brown, cone-shaped, consisting of
pollen-bearing scales; female inflorescence and fruit borne on edges of modified brown,
woolly leaves, up to 30 cm long; fruit, up to 5 cm long, ovoid, somewhat compressed,
with a thin fleshy orange-brown covering and nutlike seeds, with poisonous kernels.
Planted ornamental. No reported use in Nauru; seed kernels processed into flour as a
famine or ceremonial food in areas of Melanesia, Polynesia and Micronesia. 3, 5(46), 6,
ids

ANGIOSPERMAE (Angiosperms or Flowering Plants)

MONOCOTYLEDONAE

ARACEAE (Arum or Taro Family)

Aglaonema commutatum Schott "aglaonema"

Recent introduction. Indonesia to Pacific Is. Rare. Perennial herb, up to 1.5 m;
leaf blades, 8 to 20 cm long and 3 to 10 cm wide, elliptic- to oblong-lanceolate, dark
green with silver markings and 4 to 5 pairs of primary side veins; petioles, 5 to 15 cm
long. Ornamental pot plant. 5, 6(274), 7.

Aglaonema costatum N.E. Br. "aglaonema"
Recent introduction. S$. E. Asia. Rare. Perennial herb, up to 1 m; leaf blade,

about 12.5 by 7.5 cm, stiff, ovate to heart-shaped, dark green with white markings and 5
to 10 pairs of primary side veins. Ornamental pot plant. 6(273), 7.

39

Aglaonema marantifolium BI. "aglaonema"

Recent introduction. S. E. Asia. Occasional. Perennial herb, up to 1 m; leaf
blade, 15 to 35 cm long and 7.5 to 12.5 cm wide, oblong, pointed, dark green with light
green markings along main and side veins. Ornamental pot plant. 5, 6(271), 7.

Aglaonema cv. "Pseudobracteata" "Chinese evergreen", "aglaonema"

Recent introduction. S. E. Asia? Occasional. Perennial herb, up to 80 cm; leaf
blades, 15 to 35 cm long and 7.5 to 12.5 cm wide, dark green with large white and light
and dark green markings; petioles white. Ornamental pot plant cultivar. 5, 6(272).

Alocasia cucullata (Lour.) G. Don "Chinese taro"
syn. Arum cucullatum Lour.

Recent introduction. India. Occasional. Perennial herb, up to 1.5 m; leaves, up to
30 cm or more in diameter, dark green, long-stalked, rounded heart-shaped; petiole,
long. Ornamental pot plant. 5, 6.

Alocasia lowii Hook.f.

Recent introduction. Malaysia. Rare. Perennial herb, up to 2 m; leaves, 30 cm or
longer, metallic green, heart- or arrow-shaped, with silver-white veins and edges, edges
entire or somewhat wavy; petioles rose colored, up to 30 cm or longer. Ornamental pot
plant. 6.

Alocasia macrorrhiza (L.) Schott "giant taro", "elephant ears"
te kabe (K); taamu (Tuvalu)
syns. Arum macrorrhizon L.; Alocasia indica (Roxb.) Spach; Colocasia gigantea
Hook.f.; Colocasia macrorrhiza (L.) Schott

Recent introduction. Trop. Asia. Occasional. Tall thick-stemmed (up to 15 cm in
diameter) herbaceous plant, up to 2 m tall; leaves, up to 1 m long and 60 cm wide,
broadly arrow-shaped, leathery, entire or wavy margins; petiole, up to 1 m_ long.
Possibly an aboriginal introduction into Nauru, which was either never adopted as a food
plant or was not used at the time of European contact. No reported Nauruan name.
Planted ornamental and rare Tuvaluan food plant at| Location and in food gardens near
Topside Workshops, where a large specimen had been planted and mulched in a plaited
pandanus-leaf basket set in the ground. Important staple food plant in Samoa and Tonga
and an important supplementary staple food plant in Fiji, eastern Polynesia and on some

40

atolls in Tuvalu and the Tuamotu and Caroline Islands. Swollen tuberous stem cooked as
a staple vegetable. Wild or naturalized, ornamental (often variegated), and edible
varieties or cultivars exist in many countries, some of which are used only as a famine or
emergency food. 5, 6, 7.

Alocasia sanderiana Bull. "alocasia", "kris plant"

Recent introduction. Philippines. Occasional. Perennial herb, up to 1.5 m; leaves,
30 cm or longer, heart- or arrow-shaped, dark green with silver-white veins and edges,
reddish below, deeply-lobed edges; petioles up, to 25 to 30 cm long. Ornamental pot
plant. 5, 6, 7.

Alocasia cv. "Amazonica"

Recent introduction. Rare. Potplant. Hybrid cross with A. sanderiana. 6(185).

Anthurium andraeanum Lind. "anthurium"

Recent introduction. Trop. America. Rare. Erect perennial, up to | m tall; leaves,
15 to 20 cm long and 7.5 to 12 cm wide, oblong, heart-shaped; petiole, longer than
blade; flowers subtended by a plastic-like, bright red or orange-pink, heart-shaped spathe,
10 to 12 cm long, flowers borne on a yellow or white spadix, up to 15 cm long.
Ornamental pot plant. 6.

Caladium bicolor (Ait.) Vent "artist’s pallet", "caladium"
syn. Arum bicolor Ait.

Recent introduction. Brazil. Occasional. Perennial herb, up to 35 cm tall; leaf
blade, 10 to 30 cm long, attractive, heart-shaped, variegated, patterned with green pink,
red or white spots; leaf stalks, 3 to 6 times longer than blade. Ornamental pot plant.
SO Sil7,.90195),.5, 0,7.

Colocasia esculenta (L.) Schott "taro", "dasheen"
detaro (N); te taroro (K); talo (T)
syns. C. antiquorum Schott; Caladium esculentum Vent.

Pre-World War I introduction; possibly originally an unsuccessful aboriginal
introduction. Trop. Asia. Occasional. Large perennial herb rising on petioles, up to 1 m,
from an underground tuber; leaves, 25 to 50 cm long and 15 to 30 cm wide, clustered,
green, heart-shaped or ovate-cordate, peltate, tips pointing down; petioles, up to 1 m;

41

inflorescence shorter than leaves, spathe yellow, spadix shorter, cylindrical; tuber or
corm, up to 30 cm long and 15 cm in diameter, often with up to 2 to 15 side tubers or
cormels. Possibly an aboriginal introduction into Nauru, which was either never adopted
as a food plant or was not used at the time of European contact. Food plant in Tuvaluan,
I-Kiribati, and Chinese gardens at Location and Topside workshops; occasionally planted
and mulched in plaited pandanus-leaf baskets. Very important, often dominant staple in
other Pacific countries, although recently becoming less important in western Melanesia
because of widespread infestations of taro leaf blight (Phytophthora colocasiae) and
Alomae and Bobone viruses. Corms cooked as a staple food and the tender green leaves,
and sometime the petioles, as a spinach or green vegetable. 5, 6, 7(27825).

Cyrtosperma chamissonis (Schott) Merr. "giant swamp taro"
dababai (N); te babai (K); pulaka (T)
syns. Arisacontis chamissonis Schott; Cyrtosperma edule Schott; C. merkusii
(Hassk.) Schott; Caladium cordifolium Hartzer

Pre-World War I introduction from other areas of Micronesia; possibly originally
an unsuccessful aboriginal introduction. New Guinea and western Pacific Is. Uncommon.
Large massive perennial tuber-forming herb, up to 3 m high, rising from a large corm;
leaf blades, up to 2 m long, but usually less, dark olive-green, erect, ovate-sagittate
(arrow-shaped), only very slightly peltate, tip pointing upward, basal lobes rather
pointed, sinus deep; petiole, up to 2.5 m or more, but usually less, erect, spiny on the
lower part. Immature food plant at Location; small patch in poorly-drained area surroun-
ding Buada Lagoon and six plants cultivated in moist area in mulched plaited pandanus-
leaf baskets in Topside Workshop food gardens in 1987. Very important staple root crop
and ceremonial food in Tuvalu and Kiribati and other low-lying atoll countries of
Micronesia, but evidently not traditionally important in Nauru. Corms cooked as a staple
vegetable. 5, 6, 7(27826).

Dieffenbachia leonii Hort. "dumb cane"

Recent introduction. Colombia. Rare. Perennial herb, up to about 70 cm high;
leaves, variable in shape, green with more or less regular white areas; probably a
horticultural hybrid between D. maculata and D. seguine. Ornamental pot plant. 6.

Dieffenbachia maculata (Lodd.) Bunt. "dumb cane"
syn. D. picta Schott

Recent introduction. Brazil. Common. Thick-stemmed erect perennial herb, less
than 1 m tall, with a ringed, cane-like stem; leaves, up to 30 cm long and 15 cm wide,
narrowly ovate-oblong but variable, green with many irregular ivory-white flecked

42

markings, midrib strong with 15 to 20 pairs of curved-ascending lateral veins; petiole,
about 12 cm long, broadly grooved. Ornamental pot plant; occasionally planted in
gardens. Sap causes dermatitis when applied externally and causes mouth paralysis and
severe pain when taken in mouth. 3(58677, 58774), 5, 6, 7.

Dieffenbachia seguine (Jacq.) Schott "dumb cane"

Recent introduction. N. S. America and Caribbean. Occasional. Thick-stemmed
erect perennial herb, up to | m, with a ringed, cane-like stem; leaves variable, narrowly
oblong or ovate-oblong but variable, dark-green with irregular white spots and 9 to 15
side veins; petiole, not, or only narrowly grooved. Ornamental pot plant. Exhibits same
properties as described for D. maculata. 3(58677), 6, 7.

Epipremnum aureum (Lind. ex Andre) Bunt. "taro vine", "pothos aureus"
syns. Rhaphidophora aurea (Lind. ex Andre) Birds.; Scindapsus aureus (Lind.
ex Andre) Engl.; Pothos aureus Lind. ex Andre; Epipremnum pinnatum

cv. "Aureum".

Recent introduction. Solomon Is. Occasional. Branched high climbing vine with
thick rope-like stems with adventitious rootlets; leaf blades, up to 50 cm long and 35 cm
wide, ovate-subcordate, deeply-lobed, petiolate, variegated green and cream or pale-
yellow with irregularly-spaced bands on each side of the midrib, lateral veins slightly
ascending. Planted ornamental. 3(58724), 5, 6, 7.

Monstera deliciosa Liebm. "monstera", "fruit salad plant", "taro vine", "ceriman"

Pre-World War II introduction. C. America and Mexico. Rare. Perennial
epiphytic stout-stemmed herbaceous vine, nodes often with long hanging aerial roots; leaf
blades, up to 80 m long, thick, dark-green, heart-shaped, pinnately-lobed or parted,
grooved, often conspicuously perforated; petioles, up to 90 cm long; inflorescence with
stalk, up to 15 cm long, spathe rather thick, up to 24 cm long, pale-yellowish, spadix up
to 20 cm long and 5 cm wide, cone-shaped; fruit, edible berries, up to 1 cm long,
numerous, yellowish-green to purplish, edible, borne on spadix. Pot plant and planted
ornamental. 3(58727), 5, 6, 7.

Philodendron hastatum C. Koch & Sellow "philodendron"
Recent introduction. Brazil. Rare. Perennial herbaceous climbing vine; leaf

blades, up to 25 cm long, oblong heart-shaped to sagittate or hastate, medium-green,
shiny; petiole, 15 cm or longer. Ornamental pot plant. 6, 7.

43

Philodendron scandens C. Koch & Sellow ssp. oxicardium (Schott) Bunt."philodendron"
syn. P. oxycardium Schott

Recent introduction. Trop. America. Rare. Perennial herbaceous climbing,
twining vine; leaf blades, up to 30 long and 20 cm wide, shiny green, heart-shaped,
tapering to a fine point; petioles 10 to 20 cm long. Ornamental pot plant. 6(238), 7.

Philodendron sp. "philodendron"

Recent introduction. Trop. America. Rare. Ornamental pot plant. 5 (238), 7.

Scindapsus pictus Hassk. var. argyraeus (Engl.) Engl. "silver vine"

Recent introduction. Indomalaya. Rare. Tall fleshy perennial herbaceous vine;
leaves, up to 30 cm or longer, thick, leathery, oval to heart-shaped, pointed tips curved
to one side, dark green with silver dots or marbling. Ornamental pot plant. 6(225).

Spathiphyllum cv. "Clevelandii" "spathiphyllum", "white sails"

Recent introduction. Rare. Hybrid perennial herb, up to 80 cm; leaf blade, up to
25 cm by 10 cm, dark green, narrow ovate or oblong; flowers, arising on long slender
flower stems from the base to above the leaf tops, bearing attractive white flower bracts
or spathes and white spikes or spadices, 5 to 10 cm long. Ornamental pot plant. 6, 7.

Syngonium angustatum Schott "syngonium"
syn. S. podophyllum sensu Souder non Schott

Recent introduction. Mexico. Occasional to common. Climbing or creeping
perennial herb; leaf blades, up to 20 cm long, shining green, palmate, deeply three- to
nine-parted, held erect on rigid stems. Pot plant and planted ornamental. 3(58722), 5, 6,
Us

Xanthosoma lindenii (Andre) Engl.

Recent introduction. Colombia. Rare. Perennial erect herb; leaf blades, up to 30
cm long and 7.5 cm wide, spear-shaped, green, principal midrib and side veins white,
under surface green; petioles green, up to 30 cm long. Ornamental pot plant. 6.

44

" " " "

Xanthosoma sagittifolium (L.) Schott "“tannia", "yautia", "cocoyam", "American taro"
detaro (N); te taororo (K); talo Palagi (T) :
syn. Arum sagittifolium L.

Pre-World War II introduction. W. Indies. Occasional. Large taro-like tuberous
perennial herb, up to 2 m or higher; leaf blades, up to 50 cm or longer and 40 cm wide,
thick, sagittate (arrow-shaped), glaucous, green to purple-green; petioles, up to 1 m long,
arising from a central tuber or corm with up to 10 or more lateral tubers or cormels,
each 15 to 25 cm long. Food plant in home gardens at Location and near Nauruan home
at Buada; occasionally planted and mulched in pandanus-leaf baskets at Topside work-
shops; found primarily in Tuvaluan gardens. Important staple food crop throughout
Melanesia and Polynesia. Possibly introduced into Nauru in the late 19th or early 20th
century, but never becoming as important as in other areas of the Pacific. Side cormels
cooked as a staple and tender young leaves cooked as a green vegetable or spinach. 5, 6,
he

ARECACEAE/PALMAE (Palm Family)

Caryota urens L. "fishtail palm", "Wine palm", "toddy palm"

Recent introduction. Trop. Asia. Rare. Erect single-stemmed palm, up to 10 m;
fronds, up to 5 to 6 m long, drooping, bipinnate, with wedge-shaped fishtail-like leaflets,
irregularly and jaggedly toothed; flowers, numerous, grouped in threes, one female
between two male flowers, hanging in clustered panicles, each produced successively
lower on the trunk from the leaf (frond) nodes, until the lowest node flowers and
produces seed, after which the plant dies; fruit, ovoid, red, juicy, surrounding a kidney
shaped seed. Planted ornamental. 6.

" "

Chrysalidocarpus lutescens H. Wendl. "golden cane palm", "golden-fruited palm"

Recent introduction. Madagascar. Rare. Small erect many-trunked clump-forming
palm, up to 5 m or more high, with smooth bamboo-like ringed trunks; fronds, arching,
light green, turning yellow-orange with age, deeply divided into long narrow forked
segments, each about 2.5 cm wide; flowers, borne in clusters among the leaves, male and
female flowers separate in the same cluster; fruit, yellow, date-like. Planted ornamental.
3.0:

45

Cocos nucifera L. "coconut palm"
ini (N); te ni (K); niu (T)

Aboriginal introduction. S. Asia and Indian Ocean Islands. Abundant. Tall erect
single-stemmed palm with a slender, more or less curved or inclined, trunk, up to 30 m
tall; leaves, up to 4 m or longer, clustered at top of trunk, frond-like, pinnate, with
numerous narrow oblong-lanceolate leaflets (pinnae), 25 to 90 cm long; flowers, in
clusters on a simple-branched spadix, up to 1.2 m or longer, which originates in the axils
between the leaves, with female flowers at the base of the branchlets and small male
flowers near the tips and over the remainder of the branchlets; each spadix subtended by
a stout woody, boat-shaped, beaked bract or spathe; fruit, 15 to 30 cm long, subglobose
to ovoid or ellipsoid, one-seeded, with a thick fibrous husk surrounding a hard nut filled
with hard white oily edible pulp and, when young, with sweet water. Planted extensively
on coastal strip, around Buada Lagoon, near roads in strip-mined areas; occasional on
plateau and escarpment; common along strand; common in Nauruan home gardens and
occasionally planted around contract workers quarters at Location and Topside work-
shops. Cultivars include inur, ito, ita, inamaro and ini. Formerly important for copra
production for export; trunks used in house construction and for animal pens; midrib of
frond used for flooring and walls of houses; young fronds used for weaving baskets, food
containers and parcels, mats, housing thatch, fans, hats, dividers for communal fish
farming in Buada Lagoon, and other plaited ware and for making skirts (ridi); old and
young fronds used for roofing; coir and dry leaves important as tinder in making fire by
friction and carrying fire; midrib of leaflets or pinnules used in brooms and in weaving;
soft endosperm (meat) and water (milk) of young, green nuts (ini) consumed; meat of
mature nuts (eanikiwi) grated and eaten in a variety of ways and squeezed and boiled,
usually with flowers or leaves, to make perfumed coconut oil; coconut endosperm
("meat") has been an important staple throughout the small-island Pacific and in coastal
areas of larger islands, with some people receiving up to 70% of their dietary calories
from this source; it was also undoubtedly the main staple plant food of Nauruans in the
past and a major food for chickens and pigs; coir or husk of both green and mature nuts
used to make strong fibre and cordage (sennit) for strainers, affixing tool handles, boat
and house lashings, fishnets and lines, belts, canoe caulking, corks or stoppers, slings;
dried fronds, husks, and shells used as fuel for cooking; shells used for making charcoal,
one of the main uses being to fuel hand irons in the past; shells used to make drinking
cups in the past; sap from flower spathe tapped to make toddy (karawai) and then often
boiled down to make a molasses-like syrup (kKamaimai); toddy often allowed to ferment to
become alcoholic "sour" toddy; loose burlap-like tissue (inini) at the base of the fronds
used to strain coconut milk; dust or pollen collected on lower ends of fronds used as a
blood coagulant and disinfectant; coconut oil use to treat tinea; inside of very small
immature nuts used in post-natal medicine; very young leaves, without the midrib,
chewed and used as a medicine for fever and infantile beriberi. 2, 3, 5, 6, 7.

46

Livistonia chinensis (Jacq.) R. Br. "Chinese fan palm", "fountain palm"
syn. Latania chinensis Jacq.

Recent introduction. China, Ryukyu and Bonin Islands. Rare. Attractive single-
stemmed erect palm, up to 10 m tall, but normally less than 5 m, with a ringed gray
trunk; fronds, rounded or palmate, fan-like, about 1.5 m or more in diameter, pleated in
middle and divided radially into 70 to 90 leaflets, about 4 cm wide, each with yellow-
green midribs and split (bifid) drooping tips; petioles, up to 2 m long and 15 cm wide,
with stout tooth-like spines, up to about 1.5 cm long, on the lower part; flowers, borne
in large numbers in clusters on branchlets of long branched inflorescences; fruit, up to 2
cm long, dull bluish-green, olive-like. Ornamental pot plant and planted ornamental. 6,
Ts

Phoenix sp. "date palm"

Recent introduction. Rare. Erect single-stemmed palm with feather-like pinnate
fronds. Planted ornamental on Command Ridge. 5, 6(61).

Pritchardia pacifica Seem. & Wendl. "Pacific fan palm", "Fiji fan palm"
dabam (N); te bam (K)
syn. Eupritchardia pacifica (Seem. & Wendl.) O. Ktze.

Recent introduction. Fiji, Tonga, and Samoa. Rare to occasional. Erect single-
stemmed spineless palm up to 10 m tall; fronds, 1 m long and 1.2 m wide, spreading
from the crown, wedge- or fan-shaped, pleated or folded in middle, when young, and
divided radially into numerous leaflets with forked, scarcely drooping tips; petioles, about
1.5 m long, stout; flowers, numerous brownish, borne in clusters on branched axillary
panicles; fruit, about 1.2 cm in diameter, ovoid, green to blackish, 1 seeded. Planted
ornamental at the Meneng Hotel. Small edible fruit consumed in Fiji, Tonga and
elsewhere in the Pacific but reportedly not eaten in Nauru. 3, 5, 6, 7.

Roystonea elata (Bartr.) Harper "royal palm"
dabam (N)
syns. Palma elata Bartr.; Roystonea regia (HBK.) O.F. Cook; Oreodoxa regia
HBK.

Recent introduction. Cuba and Florida. Rare. Erect single-stemmed symmetrical
palm, up to 15 m or taller, with a smooth greenish-grey to whitish trunk; fronds, up to 3
m long, pinnate, arching from crown and radiating from a long green cylinder of leaf
sheaths which looks like an extension of the trunk; leaflets, nearly 1 m long, diverging at
various angles; inflorescence, up to | m long, borne below the leaves, many-branched,
with a long, slender boat-shaped spathe; flowers, usually in groups of 3, of which 2 are

47

staminate and 1 pistillate, cream-colored with violet stamens; fruit, less than 2 cm long,
ovoid, purplish, one-seeded. Planted ornamental. 5(137), 6, 7.

BROMELIACEAE (Pineapple Family)

Ananas comosus (L.) Merrill "pineapple"
te bainaboro (K); fara, painapolo (T)
syns. Bromelia comosa L.; B. ananas L.; Ananas sativus Schult.f.; A. ananas
(L.) Karst.

Pre-World War I post-European contact introduction. Brazil. Rare. Green to
bluish-grey short-stemmed perennial herb, up to 1 m _ high; leaves, sword-shaped,
succulent, pointed and sometimes armed (spiny), arranged in a bushy rosette; inflores-
cence, terminal, many-flowered head with small blue flowers; fruit, a pinecone-shaped or
globose syncarp, 15 to 30 cm long, yellow-brown to green, surmounted by a crown of
small leaves, which is composed of many 6-sided berries arranged spirally and embedded
in yellow pulp. Food plant in home gardens at Location and Topside Workshops and
occasionally in expatriate home gardens. Fruit edible. 3, 5, 6.

Nidularium innocenti Lem.

Recent introduction. Brazil. Rare. Perennial rosette-forming herb, up to 50 cm of
higher; leaves, strap-shaped, finely-toothed, radiating from a rosette, overlapping bases
forming water-holding reservoirs; flowers, central, small, white with showy red-orange
bracts. Ornamental pot plant. 6.

Tillandsia usneoides L. "Spanish moss", "Florida moss", "long moss"
Recent introduction. Trop. America. Rare. Pendulous, rootless epiphyte with

slender, wiry stems, up to 4 m or longer; leaves, 2.5 to 7.5 cm long, grey, narrow,

curved, threadlike, scattered at regular intervals along stem; flowers, nearly 12 mm long,

axillary, inconspicuous, single, yellowish. Planted ornamental. 6.

Vriesia sp. "vriesia"

Recent introduction. Trop. America. Rare. Perennial herb with stiff smooth-edged
variegated leaves arranged in a rosette. Ornamental pot plant. 6.

48
CANNACEAE (Canna Family)

Canna indica L. "Indian shot", "canna"

Recent introduction. W. Indies. Rare. Perennial herb, to over 1 m high; leaves,
15 to 50 cm long and 10 to 25 cm wide, oval to narrow, rather fleshy, borne on long
slender erect green stems, up to | to 1.5 m long; flowers, orchid-like, red, yellow or
speckled (variegated), in loose erect clusters; fruit, warty, black, capsular, nearly
globose, containing a variable number of round black seeds. Planted ornamental. Most
ornamental specimens are showy hybrids or selections. 6(189), 7.

COMMELINACEAE (Dayflower Family)

Dichorisandra thyrisiflora Mikan "blue ginger"

Recent introduction. Brazil. Rare. Perennial herb up to 1 m tall; leaves, shiny-
green, spirally arranged; flowers, terminal, blue to violet-blue and white, borne on a
stalk in a cone-shaped flower panicle. Planted ornamental. 6.

Rhoeo spathacea (Sw.) Stearn "tradescantia", “oyster plant", "Moses in a boat"
syns. Tradescantia spathacea Sw.; T. discolor L.’Her.; R. discolor (L’Her.)
Hance

Recent introduction. Mexico and W. Indies. Occasional. Perennial short-stemmed
herb, up to 40 cm high; leaves, up to 20 to 30 cm long and 4 to 6 cm wide, lance- or
sword-shaped, stiff, succulent, shining green above, purplish beneath, borne in crowded
rosettes; flowers, small, white, in axillary clusters within boat-shaped purple bracts; fruit,
capsular, 3-valved. Pot plant and planted ornamental; planted along borders in ornamen-
tal gardens. 3(58703), 5, 6, 7.

Setcreasia purpurea B. K. Boom "purple tradescantia", "purple heart"

Recent introduction. Mexico. Rare. Perennial smooth-stemmed trailing herb, up to
30 cm high; leaves, up to 15 cm long and 3.5 cm wide, ovate-acuminate, black-purple
above and purple beneath, with woolly hairs along margins; long purple flowering stems,
up to 30 cm, bearing purple bracts and rose-pink flowers in dense clusters. Ornamental
pot plant. 3(56779), 6, 7.

49

Tradescantia fluminensis Vell. "wandering Jew"

Recent introduction. S. America. Rare. A trailing perennial herb with green or
purplish wiry stems; leaves, 2.5 to 7.5 cm long, oblong, bright green or white-striped,
sometimes purplish beneath; flowers, small rose-purple to whitish, each with two leafy
bracts. Ornamental pot plant. 6.

Zebrina pendula Schnizl. "purple wandering Jew"

Recent introduction. Mexico. Occasional. A fleshy trailing perennial herb; leaves,
up to 6 cm long, ovate, reddish-purple beneath and silver-green with purple bands down
the middle and around the edges above; flowers, 3-petaled, pink to rose-purple, borne in
bract-like leaves near stem tips. Ornamental pot plant. 3(56780), 5, 6, 7.

CYPERACEAE (Sedge Family)

Cyperus alternifolius L. "umbrella plant", "umbrella sedge"

Recent introduction. Madagascar. Rare. Tufted sedge with many stout dark green
flowering stems, up to 120 cm or higher; basal leaves reduced to lanceolate, acuminate
sheaths, 10 to 20 cm long; flower heads, terminal, umbel-like, composed of arching leaf-
like bracts, up to 25 cm long and 1 cm wide, borne at the end of stems, and umbel-like
inflorescences, 7 cm across, on rays, up to 10 cm long, with numerous spikelets, up to 3
cm long, crowded at the tips; fruit, 3-angled achene, less than | mm long. Planted
ornamental. 6, 7.

Cyperus compressus L. "sedge"

Recent introduction? Pantropical. Occasional. Tufted sedge with erect to
spreading thin triangular stems, up to 40 cm long; leaves, 1.5 to 3 mm wide, threadlike,
shorter than sheathing stems, which are up to 3 cm long, and often reddish; spikelets, 1
to 2.5 cm long and 3 to 5 mm wide, green, 12 or more borne in umbellate clusters
subtended by 3 or more leaf-like bracts, each about 3 mm long; fruit, a 3-angled,
obovoid achene, sides slightly concave, brown to almost black. Weed growing in rather
dense populations in low ground near Buada Lagoon and swampy area near bottom of
escarpment. 3(58644), 6.

50
Cyperus iria L. "sedge"

Recent introduction. Sedge up to 40 cm high; leaves, few, shorter than culms;
inflorescence, branched spikes, about 2.5 cm long, 12- to 20-flowered, spikelets about 6
mm long, bracts longer than inflorescences; fruit, a 3-angled, narrowly ovoid, yellow-
brown achene. Rare weed. 5(90a).

Cyperus javanicus Houtt "sedge", "marsh cypress"
reyenbangabanga (N); te ritanin (K); mouku (T)
syns. C. canescens Vahl; C. pennatus Lam.; C. stuppeus Forst.f.; Mariscus
Javanicus (Houtt.) Merr.; M. albescens Gaud.; M. pennatus (Lam.)
Domin; M. stuppeus (Forst.f.) Merr.

Indigenous? Paleotropical. Abundant. Large perennial tufted sedge, up to 1 m
high, with triangular stems; leaves, 7 mm or more wide, are often longer than flowering
stems, firm, rough, serrulate; inflorescence, compound umbrella-like flower clusters, to
about 15 cm across, with branched rays, up to 10 cm long, bearing spikes, to nearly 2.5
cm to 3.5 cm long, and crowded green to brownish spikelets, about 4 to 8 mm long;
fruit, 1.5 mm long, 3-angled, ellipsoid, dark-brown or black achene. Found growing wild
in isolated clusters and tufts and in colonies or dense populations in moist habitats on the
coastal strip, surrounding Buada Lagoon, on the inner border of the coastal strand, and
occasionally in mined areas. Stems used as stringers for garlands and for stringing fish;
swollen bottoms eaten occasionally in the past. The name "reyenbangabanga" means
literally the "surrounding border", referring to the way in which C. javanicus surrounds
parts of Buada Lagoon. 3(53634), 4(160N), 5(90), 6(218), 7(27824), 8(9576).

Cyperus papyrus L. "papyrus", "Egyptian paper"

Recent introduction. Africa and Mediterranean. Rare. Tall perennial clump-
forming green-stemmed sedge, 1 to 4 m high; leaves, basal ones reduced to bladeless
sheaths, the involucral leaves much shorter than inflorescence; inflorescence, umbel-like
flower cluster of tufted yellow-green flower heads on drooping threadlike rays, 10 to 40
cm long, with linear spikelets, 6 to 10 mm long and 1 mm across, bearing 6 to 20 scales;
fruit, an oblong, 3-angled achene. Planted ornamental. 5, 6.

Cyperus rotundus L. "nut sedge", "nut grass"
ibugibugi (N); te mutemute (K); mouku (T)
syn. C. hexastachyos Rottb.

Pre-World War II introduction? Cosmopolitan. Common. Perennial tufted erect
sedge, 10 to 50 cm high, with hard, scaly brown-black tubers borne on underground
runners or rhizomes; leaves, 5 to 15 cm long and 2 to 5 mm wide, grass-like, folded

Sil

along the midrib; inflorescence, reddish-brown flower spikelets, 1 to 2 cm long, and 2 to
4 leaf-like bracts, 1 to 12 cm long, borne on unequal rays, to 6 cm long, in loose
terminal umbels borne at the top of three-angled flowering stems (culms), 10 to 40 cm
tall; fruit, about 1.5 mm long, brown, oblong, 3-sided achene. Weed in gardens; growing
in extensive stands as lawns; in swamps in Meneng. 3(58686), 4(149N), 5(89), 6, 7, 8.

Eleocharis ochrostachys Steud. "sedge"
syn. E. laxiflora (Thw.) H. Pfeiff.

Recent introduction. Asia. Rare. Erect perennial sedge, 35 to 70 cm tall, with
pithy stems; leaves, reduced to basal sheaths; inflorescence, a cylindrical spikelet, 10 to
20 cm long and 3 to 4 mm across, with numerous green to yellowish-brown scales, 4 to
5 mm long, and 5 to 7 bristles; fruit, 1.5 to 2 mm long, 2-sided, obovate, shiny brown to
gray achene. Planted ornamental. 6(210).

Fimbrystylis cymosa R. Br. "sedge", "beach sedge"
ibugibugi, ibiugbiugi; te uteute ni mane (K); mouku (T)
syns. F. spathacea Roth; F. pyncnocephala Hillebr.; F. glomerata (Retz.) Nees
ex K. Schum. non (Schrad.) Nees; F. atollensis St. John; F. wightiana
Nees

Indigenous. Pantropical. Abundant. Perennial tufted sedge, 10 to 50 cm high, with
erect 3-angled stems; leaves, 5 to 30 cm long and 1.5 to 4 mm wide, numerous, stiff,
linear, densely-clumped; inflorescence, a head or simple umbel with 3 to 8 primary rays,
1 to 4 cm long, and single or clustered brown spikelets, 1 to 4 mm long, in globose
heads, 5 to 10 mm in diameter, borne on 3-angled flowering stems (culms), 10 to 60 cm
high; fruit, less than 1 mm long, 2- to 3-sided, obovate, brown-black achene. Found
growing in clusters or tufts in open and semi-open places on the coastal strip and in
mined areas on the plateau. No reported use on Nauru. 2, 3(58613, 58670), 5, 6(210), 7.

DIOSCOREACEAE (Yam Family)

Dioscorea alata L. "yam", "greater yam", "winged yam"
te iam (K); ‘ufi (T)

Pre-World War II introduction. S. E. Asia. Rare. Herbaceous or shrubby,
twining, high-climbing tuberous vine with square, more or less winged, unarmed, four-
angled stems; leaves, up to 12 to 25 cm or longer and 8 to 15 cm wide, opposite, ovate
or heart-shaped (cordate), palmately S- to 11-nerved; petiole, nearly -half, to nearly as
long as blade; flowers, rarely-seen, small, greenish, in narrow terminal axillary panicles,

a2

up to 30 cm or more long; tubers, large, up to 10 kg or more in weight, variable-sized,
globose, cylindrical or lobed, black- to brown-skinned, with white to purplish flesh. Food
plant in home gardens at Location and Denigomodu. Important staple food crop in many
parts of Melanesia, Polynesia and Pohnpei (Ponape), Yap and other high islands in
Micronesia, where numerous named cultivars are recognized, but insignificant in Nauru.
Tuber cooked as a staple vegetable. 5, 6.

Dioscorea esculenta (Lour.) Burkill "lesser yam", "sweet yam", "Goa yam"
syns. Oncus esculentus Lour.; Dioscorea fasciculata Roxb.

Pre-World War II introduction. S. E. Asia. Rare. Herbaceous or woody, twining,
high-climbing tuberous vine with cylindrical thorny or armed stems; leaves, 10 to 15 cm
long and wide, alternate, heart-shaped or orbicular, woolly-pubescent; petioles as long as
the blades; flowers, 4 to 5 mm wide, rarely-seen, small, green, borne in slender axillary
panicles, up to 40 cm long; tubers, 15 to 20 cm long, clustered, ovoid, potato-like, with
thin brownish skin and white, slightly sweet flesh. Food plant in home gardens at
Location and Meneng. Important staple in parts of Papua New Guinea and Solomon
Islands and a supplementary staple crop in many areas of Melanesia, Polynesia and
Micronesia. Tuber cooked as a staple vegetable. 5, 6.

IRIDACEAE (Iris Family)

Gladiolus sp. "gladiolus"

Recent introduction. S. Africa. Rare. Erect herbaceous bulbed annual, up to 1 m
or higher, of which many hybrid cultivars exist; leaves, up to over 60 cm long and 2 to 3
cm wide, erect, strap- or sword-shaped; flowers, 5 cm or more in diameter, few to
many, showy, white, yellow, red or purplish, often variegated, with curved funnel-
shaped tubes and oblong segments, born in 1-sided spikes on flowering stems which are
longer than the leaves. Planted ornamental. 5, 6, 7.

Iris sp. TintSy

Recent introduction. Origin? Rare. Erect herbaceous rhizomatous perennial, up to
60 cm or higher, of which many hybrid cultivars exist; leaves, several, flat, narrow,
forming a fan-shaped cluster at the base of an erect flower stem; flowers, blue to
purplish-blue. Planted ornamental. 5, 6(100).

353

Tigrida pavonina (L.f.) Ker-Gawl. "tiger flower"

Recent introduction. Mexico and Guatemala. Rare. Erect herbaceous bulbed
perennial, up to 75 cm tall; leaves, about 30 cm long, stiff, narrow, almost pleated;
inflorescence, a branched or unbranched flower stem, up to 60 cm or more, bearing
three-lobed, red, orange, yellow or white, often spotted, flowers, 7 to 15 cm in diameter,
which appear one after another from bracts, 7 to 12 cm long. Planted ornamental. 6.

LILIACEAE (Including Agavaceae and Amaryllidaceae) (Lily Family)

Agapanthus africanus (L.) Hoffmannsegg "African lily", "lily of the Nile"

Recent introduction. S. Africa. Rare. Erect perennial herb, up to 60 cm tall;
leaves, strap-shaped, evergreen; inflorescence, tall flowering stems bearing showy flower
heads with many blue, violet-blue or white funnel-shaped flowers, each about 5 cm long.
Planted ornamental. 6.

Ww "

Agave americana L. "century plant", "malina"

Recent introduction. Mexico. Rare. Large stemless erect perennial succulent herb,
up to 2 m or higher; leaves, 80 to 160 cm long and 15 to 20 cm wide, thick, fleshy,
pointed and spined, variegated with yellow to white margins, borne in a massive rosette
of 30 to 60 leaves; flowers, 7.5 cm long, yellow-green, borne in clusters on a tall pole-
like branching stem, up to 6 to 10 m high, arising from the center of the rosette. Planted
ornamental. 5(68), 6.

Agave rigida Mill. "sisal", "sisal hemp", "malina", "agave"
te robu (K)
syn. A. sisalana Perr.

Pre-World War II introduction? Mexico. Occasional. Large stemless perennial
succulent herb, up to 2 m or higher; leaves, up to 150 cm long and 10 to 15 cm wide,
thick, stiff, fleshy, spine-tipped, bluish, arranged in a large rosette; flowers, up to 5 cm
long, shortly-tubular, yellow-green, borne in panicles on a tall branching stem, up to 8 m
high, arising from the center of the rosette. Established locally, especially along edges of
old strip-mined area. Grown for export in some tropical areas for the fibre from its
leaves which is made into rope and other products. 3(58739), 5, 6(215), 7.

54

Allium ascalonicum L. "shallot"
te anian (K); ts’ung (C) ;

Pre-World War II introduction. Palestine. Occasional. Small erect green herb, up
to 35 cm; leaves, 20 to 35 cm long and 5 to 15 mm wide, narrow, strap-shaped, growing
from many clustered individual white to reddish bulbs, surrounding a single planted bulb,
which separate into segments or "cloves". Cultivated in Chinese food gardens in beds and
containers at Location and Topside workshops. Pungent bulbs and tender leaves eaten
raw or cooked as a spice or vegetable. 5, 6, 7.

Allium cepa L. "bulb onion", "common onion"
te anian (K)

Recent introduction. Persia. Rare. Herb with hollow flattened tubular leaves and a
well-developed white or light-green, generally single, bulb with white to yellow-brown
skin. Single immature plant in Chinese food garden at Location. 6.

Allium fistulosum L. "green onion", "spring onion", "Welsh onion",
"Japanese bunching onion" te anian (K); ts’ung (C)

Pre-World War II introduction. E. Asia. Occasional. Erect green herb, up to 50
cm tall; leaves, 25 to 50 cm long and | to 2 cm wide, green, round, tubular, clustered,
with white, slightly swollen stems which produce no bulbs. Cultivated food plant in
Chinese gardens at Location. Pungent tender leaves and stems eaten raw or cooked as a
spice or vegetable. 5, 6.

Allium porrum L. "leek"

Recent introduction. Eurasia. Rare. Herb with long flattened and folded leaves, up
to 70 cm long, a long white neck below the leaves, and a small white bulb. Single plant
growing in food garden at Location. 6.

Allium sativum L. "garlic"
suen, suen t’au (C)

Pre-World War II introduction. S. Asia. Occasional. Erect green herb, up to 30
cm high; leaves, 10 to 30 cm long and 1 to 2 cm wide, flat, strap-shaped, with a well-
developed segmented bulb with a strong persistent odor, which separates into several
segments or "cloves". Plant in Chinese gardens at Location. Grown from bulbs, mainly
for its pungent edible green leaves which are used as a spice or green vegetable in
Chinese cooking. 5, 6.

55

Allium schoenoprasum L. "chives"

Recent introduction. N. hemisphere. Rare. Erect herb, up to 25 cm high, forming
grass-like clumps; leaves, 10 to 25 cm long and 1.5 to 3 mm wide, fine, tubular, with a
poorly developed bulb. Pot herb cultivated in container at Cliff Lodge. Tender green
leaves used as a spice by European residents. 5, 6.

Allium tuberosum Rottler ex Sprengle "Chinese chives"
sai ts’ung (C)

Pre-World War II introduction. E. Asia. Common. Erect herb, up to 30 cm high;
leaves, 20 to 30 cm long and 3 to 4 mm wide, flat, ribbon-like, with a poorly developed
bulb. Cultivated in Chinese food gardens at Location and Topside workshops for its
edible green leaves. 5, 6, 7.

Asparagus aethiopicus L. “asparagus fern", "Sprenger asparagus"
syns. A. densiflorus (Kunth) Jessup; A. sprengeri Reg.

Recent introduction. S. Africa. Occasional. A much-branched arching wiry-
stemmed perennial, up to 1 m, bearing open whorled fern-like leaf clusters; "leaves", up
to 2 to 2.5 cm long, shiny, flat, narrow, pale-green, lanceolate; flowers, small, fragrant,
pinkish; fruit, small, red, 1- to 3-seeded berries, about 8 mm in diameter. Ornamental
pot plant. 3(58711), 5, 6(167, 180).

Asparagus setaceus (Kunth) Jessup "asparagus fern"
syns. Asparagopsis setacea Kunth; Asparagus plumosus Baker

Recent introduction. S. Africa. Rare. Slightly woody and spiny climbing, many-
branched perennial, up to 1 m, forming flat horizontal fern-like sprays; "leaves", up to 8
mm long, fine, fern-like or needle-like; flowers, minute, white; fruit, small, blackish, 1-
to 3-seeded berries. Ornamental pot plant. 6(169), 7.

Chlorophytum capense (L.) Voss "spider plant", "ribbon plant", "bracket plant"
syns. C. comosum (Thunb.) Jacq.; C. elatum R. Br.

Recent introduction. Africa. Rare. Evergreen perennial herb, up to 30 cm or
higher, with tuberous rhizomes; leaves, 15 to 30 cm long and 1.5 to 2.5 cm wide,
rosetted or tufted, grass-like, arching, variegated with mid-green and white or yellow

56

longitudinal bands; flowers, small, starry-white, 6-parted, borne in loose clusters on
flower stems, up to 1.5 m long, and tipped with leafy offshoots or plantlets; fruit, a
leathery 3-angled capsule with flat seeds. Ornamental pot plant. 3(58690), 6.

Cordyline fruticosa (L.) A. Chev. "cordyline", "ti-plant" (Hawaii)
te rauti (K); ti (T)
syns. Convallaria fruticosa L.; Cordyline terminalis (L.) Kunth; Taetsia fruticosa
(L.) Merr.; 7. terminalis (L.) W. F. Wight

Recent introduction. Trop. Asia or Australasia? Occasional. Woody erect shrub
or tree-like branched or unbranched perennial, up to 2 m or taller; leaves, up to 60 cm
long and 15 cm wide, lanceolate or oblong, smooth, tough, shiny dark-green to rust or
red, borne in rosettes clustered in spirals near the tips of the branches; petioles, 5 to 15
cm long; flowers, numerous, white to lilac-tinted, borne on large branching terminal
panicles, about 30 cm long; fruit, globose, about 5 mm in diameter, thinly fleshy, red,
purplish, or yellowish; seeds,, obovoid, black, glossy; some cultivars with a large edible
tuberous root. Planted ornamental and pot plant. Very important ceremonial and magico-
religious plant, a traditionally important supplementary food plant and famine food, and
important decorative plant, with numerous other cultural uses in Melanesia and Polyn-
esia, where numerous named cultivars and hybrids exist. The large sweet white tubers of
some cultivars are baked for days in earthen ovens to be consumed as food, sweets or
confectionery, or, in Hawaii, made into an alcoholic beverage known as okolehao; leaves
important for parcelling food and dancing skirts and body ornamentation. Apparently a
recent introduction into Nauru with no reported non-ornamental local uses. 3(58676), 5,
6(186), 7.

Crinum asiaticum L. "spider lily", "crinum lily", "grand crinum"
dagiebu, dagibu (N); te kiebu (K); tapua, talotalo (T)
syns. C. pedunculatum R. Br.; C. procerum Bak.

Recent introduction. Trop. Asia. Occasional. Perennial erect bulbous herb, up to
1.5 m or taller; leaves, up to 1.5 m long and 20 cm wide, many, arching, fleshy, strap-
shaped, green to yellow, rising from a whitish stalk; inflorescence, an umbel-like
flowerhead bearing 10 or more spider-like, 6-parted, fragrant, white flowers, with
flowering tubes, up to 10 cm long, white filaments and purplish styles, borne on a
flattened fleshy flowering stem (scape); fruit, subglobose, beaked, 1- to 2-seeded; seeds,
large, fleshy. Planted ornamental. Flowers used in garlands; roots crushed for the
treatment of filariasis. 3, 5(122), 6(216), 7.

Si,

Crinum augustum Roxb. "crinum lily", "Queen Emma lily"
dagiebu, dagibu (N); te kiebu (K)

Pre-world War II Introduction? Mauritius and Seychelles. Large perennial erect
bulbous herb, up to over 1 m tall; leaves, up to 1.5 m long and 12 cm wide, stemless,
strap-shaped, tapering; flowers, white with dark reddish-purple markings, segments, 15
to 16 cm long by 2 to 2.5 cm wide, filaments and style purple, borne on a flower stalk
which divides into numerous short cluster-bearing stalks. Occasional. Planted ornamental;
flowers used in garlands. 5, 6, 7(22319).

Crinum macrantherum Eng.
dagiebu, dagibu (N)?
syn. C rumphii Merr.

Pre-World War II introduction. Listed as C. macrantherum on Burgesses’(1935)
list. Rare? Planted ornamental. 2.

Crinum moorei Hook.f. "veld lily"

Recent introduction. S. Africa. Rare. Perennial bulbous herb, up to 1 m high;
leaves, 60 to 90 cm long and 7.5 to 10 cm wide, lanceolate, smooth; flowers, fragrant,
rose, pink or white, funnel-shaped with curved tubes, up to 10 cm long and 2.5 cm wide,
and segments, up to 10 cm long. Planted ornamental. 5(63).

Dracaena deremensis Engler "dracaena"
syn. Pleomele deremensis (Engler) N.E. Br.

Recent introduction. Trop. Africa. Rare. Woody palm-like erect shrub, up to 1.5
m or taller; leaves, up to 45 cm by 5 cm, sword-shaped, longitudinally green and white-
striped, apetiolate. Ornamental pot plant. 6, 7.

" "

Dracaena fragrans (L.) Ker-Gawl. "dracaena", "dragon flower", "pleomele"
syns. Alectris fragrans L.; Pleomele fragrans (L.) Salisb.

Recent introduction. Trop. Africa. Rare. Woody palm-like erect shrub, up to 3 m
or higher; leaves, up to 90 cm long by 10 cm wide, arching, green or longitudinally
green and yellow-striped, apetiolate; flowers, fragrant, greenish-yellow, borne in
panicles; fruit, orange-colored, unpleasant-smelling berry. Ornamental pot plant. 5, 6,
AGA):

58

Dracaena sanderiana Sander "dracaena", "ribbon plant"
syn. Pleomele sanderiana (Sander) N.E. Br.

Recent introduction. Trop. Africa. Rare. Erect woody shrub, up | m or taller;
leaves, to about 17 cm long and 3 cm wide, petiolate, arching, dark green, with white
edges, set at intervals along the stem; petioles, 7.5 to 10 cm long. Ornamental pot plant.
Tk

Furcraea foetida (L.) Haw. "Mauritius hemp"
syns. F. gigantea Vent.; Agave foetida L.

Recent introduction. Trop. S. America. Rare. Stemless perennial herb, up to 1.5
m or higher; leaves, 1 to 1.8 m long and 10 to 15 cm wide, stiff, fleshy, sword-shaped,
spine-tipped, arranged in a rosette; flowers, strong-scented, greenish-white, borne in
terminal panicles on a tall branching stem, up to 7 m high. Found in ruderal sites. May
have been confused with Agave rigida by Swarbrick. 8.

Gloriosa superba L. "climbing lily", "gloriosa", "glory lily"

Recent introduction. Trop. Africa. Occasional. Climbing perennial lily, reaching
up to 1.5 m in length; leaves, glossy, twisted, narrow, tapering into tendrils; flowers, up
to 7.5 cm long, striking, 6-parted, bright red and yellow, with curled petals which turn
inside out and point upwards when fully open. Planted ornamental and pot plant. 5,
6(177).

" "

Hippaestrum puniceum (Lam.) Urban "Barbados lily", "amaryllis"
syns. Amaryllyis punicea Lam.; Hippaestrum equestre (Ait.) Herb.; Amaryllis
equestris Ait.

Recent introduction. Trop. America. Bulbous herb, up to 60 cm or higher; leaves,
30 to 48 cm by 4 cm, 6 to 8 in number, strap-shaped, dark green, developing after the
flowers have died; flowers, 10 to 12 cm across, funnel-shaped, red or salmon-colored
with green centers, borne in groups of 2 to 4 on a stout flowering stem, 30 to 60 cm
high. Planted ornamental. 3(58718), 6, 7.

Hosta plantaginea (Lam.) Asch. "plantain lily", "funkia"

Recent introduction. China and Japan. Rare. Clump-forming herb, up to 50 cm
high; leaves, ovate with many oblique grooves or ribs spreading from the midrib;
flowers, funnel- or bell-shaped, small, white or blue-lilac, borne in narrow spikes on a
flower stalk. Planted ornamental. 5(62), 6.

59

Hymenocallis littoralis (Jacq.) Salisb. "spider lily"
lili (N); te ruru ni mane (K); lili (T)
syn. Pancratium littorale Jacq.

Recent introduction. Trop. America. Occasional. Erect bulbous perennial herb, up
to 75 cm or higher; leaves, 45 to 75 cm long and 2 to 7 cm wide, strap-shaped, fleshy;
inflorescence, an umbel of 3 to 8 striking, white, funnel-shaped, spider-like, fragrant
flowers with narrow tubes, 10 to 17 cm long, and long slender spreading petals,
connected at the base by a thin web, borne at the top of a thick, flattened flowering stem,
30 to 75 cm long; fruit, green, smooth, 3-celled. Planted ornamental. 3(58782), 5(129),
OH Ws

Littonia modesta Hook "climbing lily"

Recent introduction. S. Africa. Rare. Unusual climbing herb, reaching 60 to 90
cm in length; leaves, narrow, shining, green, spaced along an unbranched stem, each
ending in a small tendril; flowers, rich-orange, wide, spreading, bell-shaped, borne
singly in the axils of the leaves. Planted ornamental. 6.

Narcissus sp. "daffodil", "narcissus"

Recent introduction. Europe. Rare. Bulbous herb, up to about 25 cm; leaves, flat;
flowers, fragrant, white to yellow, on individual stalks borne at the top of a flattened
stem. Ornamental pot plant. 6.

Sandersonia aurantiaca Hook.f. "golden lily of the valley", "Chinese lanterns"

Recent introduction. Natal. Rare. Tuberous perennial herb, up to 45 cm; leaves,
stalkless, spear-shaped, tapering to a fine point, borne up the slender stem in decreasing
size; flowers, showy, orange, lantern-shaped, borne on long pendant stalks from the axils
of the leaves on the upper half of the main stem. Ornamental pot plant. 6.

Sansevieria trifasciata Prain "bowstring hemp", "mother-in-law’s tongue"

Recent introduction. Trop. W. Africa. Occasional. Erect evergreen perennial, up
to 1 m high, with orange turmeric-like rhizomes; leaves, 30 to 90 cm long and 2.5 to 7.5
cm wide, fleshy, erect, sword-shaped, tapering to a sharp subulate point, decorative and
cross-banded on both sides with shades of dark green to yellowish to greyish-green, some
varieties with creamy-yellow vertical margins; flowers, whitish to yellowish or greenish,
tubular, borne in clusters on a flowering stem (scape) which is slightly shorter or as long

60

as the leaves; fruit, globose, orange with fleshy seeds. Planted ornamental and pot plant.
3(98637) 95, 6;. 7.

Sprekelia formosissima (L.) Herb. "Jacobean lily", "Aztec lily"

Recent introduction. C. America. Rare. Bulbous plant, about 30 cm high; leaves,
strap-shaped, developing after the flowers; flowers, 7.5 to 10 cm across, striking
crimson-red, single, irregularly funnel-shaped, 6-segmented, 3 together forming a lip, 2
curled back at the sides and the sixth held upright at the top of the bloom, borne on a
long hollow stem. Ornamental pot plant. 6.

Yucca gloriosa L. "yucca", "Spanish bayonet”

Recent introduction. Trop. America and S.E. United States. Rare. Tree-like
perennial shrub, up to 2 m or taller; leaves, about 60 cm long and 5 cm wide, sharp-
pointed, green to grey-blue, borne in rosettes on a branched or unbranched stem;
flowers, up to 7.5 cm across, many, white to creamy, short-stalked, cup- or bell-shaped,
borne in clusters on a much-branched flower stem extending above the leaf rosette.
Planted ornamental. 3(58683), 5, 6, 7.

Zephyranthes candida (Lindl.) Herb. "zephyr flower", "white star of Bethlehem",
"westwind flower", “storm lily"

Recent introduction. Argentina and Uruguay. Rare. Bulbous herb, up to 25 cm
high; leaves, seldom over 25 cm, green, fleshy; flowers, 3.5 to 5 cm long, solitary, 6-
segmented, white inside, rose-tinged or not outside, borne on erect or bent flowering
stems, 10 to 20 cm long; fruit, a 3-valved capsule with flat black seeds. Planted
ornamental. 6.

Zephyranthes rosea Lindl. "pink lady", "pink star of Bethlehem", "pink zephyr flower"
susana (T)
syn. Atamosco rosea (Lindl.) Green

Recent introduction. Guatemala and W. Indies. Rare. Bulbous herb, rarely up to
10 cm; leaves, up to 25 cm long, narrow, strap-shaped, downward-curved; flowers,
about 2.5 to 3 cm long, solitary, pink to rosy-red, borne on erect or bent flower stems,
up to 15 cm long. Planted ornamental. 3(58692), 5, 6, 7.

61
MARANTACEAE (Arrowroot Family)

Calathea ornata (Lem.) Koern. "calathea"
syn. Maranta ornata Lem.

Recent introduction. Northern S. America. Occasional. Perennial herb, up to 45
cm or higher; leaves, up to 65 cm long and 22 cm wide, 1 to 7 in number, narrowly
oval, green with varied striped markings above, which range from rose to white when
young, but to entirely green above and purplish-red below when mature. Ornamental pot
plant. 3(58688), 5, 6, 7.

Calathea wiotiana Makoy "calathea", "rattlesnake plant"
syns. C. insignis Bull; C. lanceolata Boom

Recent introduction. Brazil. Rare. Perennial, low, bushy herb, up to 30 cm or
taller; leaves, about 30 cm long, wavy-edged, parallel-sided, velvety green, marked with
oblique, dark olive-green blotches above and reddish-purple below. Ornamental pot plant.
6.

Calathea zebrina (Sims) Lindl. "calathea"
syn. Maranta zebrina Sims

Recent introduction. Brazil. Rare. Perennial unbranched short-stemmed herb, up
to more than 45 cm high; leaves, 30 to 60 cm long and 15 cm wide, oblong, with
somewhat wavy edges, as many as 20 in number, spirally-arranged, tufted, velvety, dark-

green and light- or yellow-green bands above and purple beneath. Ornamental pot plant.
6.

Ctenanthe lubbersiana (Morr.) Eichl. "ctenanthe", "bamburanta"

Recent introduction. Brazil. Rare. Tufted perennial, up to 50 cm high, with
slender forking stems; leaves, 15 to 20 cm long, narrowly oblong, long-stalked, varie-
gated, deep green above, lined and mottled with yellow, and paler beneath. Ornamental
pot plant. 6.

Ctenanthe oppenheimiana (Morr.) K. Schum. "ctenanthe", "never-never plant"
Recent introduction. Brazil. Rare. Clump-forming perennial, up to 75 cm high;

leaves, up to 25 cm or longer, lance-shaped, green with silver-grey bands on the upper
surface and purple-red beneath. Ornamental pot plant. 6.

62

Maranta leuconeura Morr. "maranta", "prayer plant", "rabbit tracks"

Recent introduction. Brazil. Rare. Perennial herb, up to 30 cm high; leaves, up to
15 cm long, oblong, stalked, light emerald green with brown-purple patches both sides of
the mid-vein, lying almost horizontal by day becoming more vertical at night like hands
in prayer; flowers, about 2.7 cm long, white to violet, accompanied by two narrow
bracts. Ornamental pot plant. 6, 7.

MUSACEAE (Banana Family)*

*The nomenclature for the genus Musa is confused, with most of the common
seedless cultivars or clones being triploid crosses of the fertile species Musa acuminata
Colla and M. balbisiana Colla. The Latin binomials M. nana Loureiro, M. sapientum L.,
and M. paradisiaca L. are commonly used as follows: M. nana for the "dwarf Caven-
dish", and M. sapientum for the taller bananas, which are generally eaten ripe, but which
are also cooked throughout the Pacific as starchy staples, and M. paradisiaca for the
starchier bananas or plantains, which are usually eaten cooked as a staple starch, but
occasionally eaten ripe as fruit. The nomenclature most widely used by agronomists is
that developed by Simmonds, which classifies all cultivars or clones on the basis of their
assumed genetic background, eg. Musa ABB Group would be a triploid cross of one M.
acuminata group and two M. balbisiana groups. Both nomenclature systems are presented
here to more precisely identify the clones that are currently present in Nauru.

Musa (AAA Group) Simmonds “panana", "Robusta", "poyo", "Mons Marie"
dabanana (N); te banana (K); fuamaoluga (T)
syns. M. sapientum L.; M. paradisiaca L. var. sapientum (L.) Kuntze; M.
paradisiaca L. ssp. sapientum (L.) Kuntze; M. acuminata Colla cvs

Pre-World War II introduction. S$. E. Asia. Occasional. Clump- or stand-forming
giant perennial herb, up to 6 m tall, with green pseudostems (trunks) composed of leaf
sheaths; leaves, up to 2.5 m long and 75 cm wide, broad-bladed, broadly feather-shaped,
bright green, smooth, at first entire, but soon splitting like a feather along parallel side
veins, spirally arranged in a terminal crown through which the inflorescence emerges,
leaf stems and midribs thick; inflorescence, thick, stalk-like, terminal, and bearing male
flowers, in a large budlike tip with dark purplish-red scales, and female flowers along the
stalk which turn into large hanging fruit bunches; fruit, seedless, blunt-tipped, medium-
thick-skinned, greenish-yellow, turning bright yellow on ripening; new pseudostems
sprout from base of old pseudostems which die or are cut after bearing fruit. Food plant
in Tuvaluan and I-Kiribati food gardens at Location and Topside workshops; occasional
in Nauruan and expatriate home gardens. Important food and export crop in many areas

63

of the Pacific, especially in Tonga and Western Samoa, where bananas are a major
export crop, and in Tuvalu, where bananas are a major staple food. Not a traditional food
plant, and of only limited importance on Nauru. 2, 3, 5, 6, 7.

Musa (AAB Group) Simmonds "lady’s finger banana", "pisang rajah" (Indonesia)
dabanana (N); te banana, te oraora (K); tamatamailima, inisi (T)
syns. Musa X paradisiaca L. var. hort. "Pisang raja" (M. acuminata Colla x
M. balbisiana Colla)

Pre-World War II introduction. S. India. Occasional. Clump- or stand-forming
giant perennial herb, up to 7 m tall, with bronze-green pseudostems (trunks) composed of
leaf sheaths; leaves, up to 3 m long and 80 cm wide, broad-bladed, broadly feather-
shaped, bright green, smooth, at first entire, but soon splitting like a feather along
parallel side veins, spirally arranged in a terminal crown through which the inflorescence
emerges, leaf stems and midribs thick; inflorescence, thick, stalk-like, terminal, and
bearing male flowers, in a large budlike tip with dark purplish-red scales, and female
flowers along the stalk which turn into large hanging fruit bunches; fruit, tightly-packed,
light-yellow, short, slightly rounded, plump, very thin-skinned, seedless; new pseudos-
tems sprout from base of old pseudostems which die or are cut after bearing fruit. Food
plant in Tuvaluan and I-Kiribati gardens at Location and Topside workshops. Eaten ripe
as a fruit throughout much of the Pacific and a very important staple in Samoa, where it
is cooked green. Not usually found in Nauruan gardens. 3, 5, 6, 7.

Musa (ABB Group) Simmonds "cooking banana", "plantain", "bluggoe"
dabanana (N); te banana, te umuumu (K); pata (T)
syns. Musa X paradisiaca L. var. hort. "Bluggoe" (M. acuminata Colla x M.
balbisiana Colla )

Pre-World War II introduction? §S. E. Asia and Pacific. Common. Clump- or
stand-forming giant perennial herb, up to 6 m tall, with pale green pseudostem (trunks)
composed of leaf sheaths; leaves, up to 2.5 m long and 75 cm wide, broad-bladed,
broadly feather-shaped, bright green, smooth, at first entire, but soon splitting like a
feather along parallel side veins, spirally arranged in a terminal crown through which the
inflorescence emerges, leaf stems and midribs thick; inflorescence, thick, stalk-like,
terminal, and bearing male flowers, in a large budlike tip with dark purplish-red scales,
and female flowers along the stalk which turn into large hanging fruit bunches; fruit,
light-green, waxy, thick-skinned, angular, with a tapering blunt-tip; new pseudostems
sprout from base of old pseudostems which die or are cut after bearing fruit. Food plant
in well mulched areas and planting boxes at Location and near Topside workshops in
Tuvaluan and I-Kiribati gardens. Important traditional supplementary staple in many areas
of the Pacific, where it seems to be an aboriginally introduced cultivar and known as bata
or pata in Fiji and Polynesia respectively. The most common banana cultivar in Kiribati,
from where it may have been introduced into Nauru. Leaves and pseudostem used

64

medicinally to wrap sick persons to lower fevers; leaves used to parcel food and to cover
earthen oven; green fruit cooked as a staple food and ripe fruit eaten raw. Evidently not a
successful aboriginal introduction to Nauru, where Musa cultivars do not seem to have
been a traditional food crop. 5, 6, 7.

ORCHIDACEAE (Orchid Family)

Cattleya sp. “cattleya orchid"

Recent introduction. Trop. America. Rare. Bulbous or slender-stemmed orchid
with thick, slightly-folded leaves and large showy flowers. Ornamental pot plant. 6.

Dendrobium undulatum R. Br. "dendrobium orchid"

Recent introduction. Australia. Rare. Orchid with short thick leaves and many
showy flowers borne along a flowering stem. Ornamental pot plant. 6.

Dendrobium sp. "dendrobium orchid"

Recent introduction. Rare. Orchid with short thick leaves and many showy
flowers borne along a flowering stem. Planted ornamental at H.M. De Robert’s home. 6.

Spathoglottis plicata Bl. "Malayan ground orchid"

Recent introduction. Indomalaysia. Rare. Terrestrial erect perennial herb, up to 60
cm or taller; leaves, 50 cm to 120 cm long and 5 to 20 cm wide, oblong-lanceolate or
elliptic-lanceolate, plicate, with numerous parallel veins; flowers, 2 to 3 cm wide, pink to
rose-lavender, with some magenta and bright yellow in the center, borne in clusters on a
long flowers stalk (scape), up to 150 cm or higher; fruit, a narrow oblong seed-bearing
capsule, about 3.7 cm long. Planted ornamental. 5, 6.

Vanda teres Lindl. "vanda orchid"

Recent introduction. Burma. Rare. Evergreen orchid with tall climbing stems, up
to 2 m; leaves, about 15 cm long, dark green, cylindrical; flowers, 2 to 5 flowers, each
10 cm across, with rounded creamy white to rose-tinted sepals, rounded rose-colored
petals, orange or yellow throats with red markings, and the middle lobe purple or rose,
borne in clusters on the climbing stems. Planted ornamental. 6.

65

Vanda sp. "vanda orchid"

Recent introduction. S. E. Asia. Rare. Evergreen orchid with showy flowers.
Planted ornamental. 6.

PANDANACEAE (Pandanus Family)*

*The nomenclature for the genus Pandanus is, like Musa, confused, with some
taxonomists classifying many of the common cultivars and wild clones or species, both
edible and non-edible, as forms or varieties of P. tectorius. Other taxonomists consider
them distinct species, often listing numerous species or varieties for a given area. For
example, P. odoratissimus L.f. has long been thought to be synonymous with P.
tectorius, but is not considered, by many authorities, to occur east of Malaysia. Similar-
ly, P. odoratissimus L.f. var. pyriformis Mart. has been used as a synonym for a wild
and doubtful variety of P. tectorius, whereas Stone (1970) considers P. fragrans Gaud. to
be the common wild species on Guam, and does not consider P. fectorius to be present.
Thus, because the fruit of many of the named cultivars or varieties found on Nauru were
not collected and identified, the identifications here must be considered provisional, with
most named cultivars being grouped under P. tectorius. Other widespread forms, such as
P. dubius Spreng., a widespread edible species; and P. spurius Miq. cv. "PUTAT"
(syns. P. tectorius Warb. var. laevis Warb.; P. odoratissimus L.f. var. laevis (Warb.)
Mart., which are widely cultivated for their leaves for use in plated ware, are also
possibly present, but not listed here. P. dubius, if present, however, is a very different
plant, not likely to be confused with P. tectorius. It has no edible fleshy tissue, but a
large globose fruit, 30 to 40 cm in diameter, composed of many drupes, 8 to 18 cm long,
each containing a single seed with edible white endosperm.

Pandanus tectorius Warb. var. pulposus Warb. "pandanus", “screw pine"
epo, epuh (N); te kaina (K); fala (T)
syn. P. pyriformis Gaud.

Indigenous and probably an aboriginal introduction in the case of some cultivars.
Pacific Is. Common. Stout, branching tree, up to 5 m or more tall, with numerous aerial
roots and thick forking stems; leaves, seldom over 10 cm wide and 1 to 3 m long,
spirally-arranged, pointed, with armed or spiny margins and midribs; male inflorescence,
fragrant, pendant, with cream-yellow bracts and white spikes; female inflorescence,
similar but smaller, on separate trees; fruit, pineapple-like, ovoid, 20 to 30 cm long and
10 to 20 cm wide, with 50 or more wedge-shaped, yellow to red-orange, 1 to 12-celled
woody drupes, each about 4 to 8 cm long. Commonly planted on the coastal strip in
home gardens and in open areas and in stands on the unmined portions of the plateau

66

(Topside); women were formerly responsible for the care and cultivation of pandanus,
although men helped in the initial clearing of land. An important staple to the Nauruans
and to the I-Kiribati and Tuvaluans on their home islands. Very important fresh fruit and
staple in Nauru and other atoll and Micronesian countries. Named cultivars which still
exist on Nauru include enaben (enabun), erabaite (erabwaite), eragadibyaw, eragomogom
(eragumugum), inaparabei (inaporabei), inimenoiya, irireiab, and iriribe; other named
cultivars, which are reportedly now extinct due to mining, bombing during World War
II, and failure to replant, include eraburabur (eraburbur), eramwimwi, erarapaiwa
(erarapeiwa, erarapwiewa), erkibwir, erokwoi, eronubwe, erwuro, inibiterin, and itoidi
(etoidi). Some of these cultivars may be P. dubius (see above). Ripe fruit of all cultivars
eaten in Nauru as a vitamin-A-rich snack food and also cooked or fermented and put on
pandanus mats or leaves in the sun to dry to make a "rich man’s" food known as edongo,
which is also a traditional food for the sick; leaves, which are commonly soaked in fresh
water or boiled, are plaited into mats (itubare), baskets, and other plaited ware, and
make the best traditional thatching and roofing; main trunk and stilt roots used in house
construction; wood and dried fruit sometimes burned as fuel; thin outside bark of the stilt
roots scraped and mixed with coconut juice to cure constipation and poor appetite; I-
Kiribati use leaves as cigarette wrappers. According to Wedgewood (1936), during the
yearly pandanus harvest (ineded), which usually occurred around August or September,
people used to leave their homes on the coast to stay in temporary bush huts on the
pandanus lands in the interior. 2, 3(58760), 5(64), 6, 7(27814?).

Pandanus sanderi Hort. ex Masters "variegated pandanus"

Recent introduction. Indomalaysia to the Pacific Is. Rare. Perennial shrub with
long pointed variegated green leaves with yellow-white margins. Planted ornamental. 6.

POACEAE OR GRAMINAE (Grass Family)

Andropogon sp.

Recent Introduction. Rare. Small perennial grass. Local on strip-mined land.
3(58736).

Arundo donax L. "giant reed"

Recent introduction. Old World. Occasional. Large perennial grass or reed with
strong, bamboo-like, erect hollow stems, 2 to 4 cm in diameter, rising in clumps, up to
4 m high; leaves, 50 to 70 cm long and 2 to 6 cm wide, flat, pale bluish-green, smooth;
inflorescence, a large feathery flowering panicle, up to 70 cm long, with whitish to

67

purplish spikelets, 8 to 12 mm long. Planted ornamental and spontaneous in some areas.
3(58742), 3(171N), 5(95), 6, 7(22318).

Bambusa vulgaris Schrad. ex Wendl. “common bamboo", "feathery bamboo"
ebarabaratu, embarabaraba (B)(N); te kaibaba (K)
syns. Arundo bambos L.; Bambos arundinacea Retz.; Bambusa arundinaria
Willd. ex Merr. (Sphalm.); B. arundinacea (Retz.) Willd.

Pre-World War II post-European contact introduction. Trop. Asia. Rare. Giant
perennial clump-forming woody grass with segmented green to yellowish stems (culms),
up to 15 m tall and 8 cm wide; leaves, 9 to 30 cm long and 1 to 4 cm wide, feathery,
lanceolate, rough below and on the margins; flowers, spikelets, 2 to 3.5 cm long, borne
in large leafy clusters at the nodes. Planted on coastal strip. Woody stems used in the
construction of perches for pet frigate birds, for fishing rods, net handles for noddy-bird
nets and reef and flying-fish nets; splinters used in the past to mend fishing nets; parts
used medicinally. 2, 5, 6(217), 7.

Cenchrus brownii R. & S. "Brown’s burgrass", "Brown’s sandbur"
eakung, iyakong (N)
syn. C. viridis Spreng.

Recent introduction. Trop. America. Occasional. Annual grass, up to 50 cm or
higher; leaves, 5 to 12 cm long, narrow, slightly hairy; inflorescence, a cylindrical
flowering spike bearing many crowded somewhat delicate globose spiny burrs. Weed in
open places and ruderal habitats on the coastal strip. 5(35), 6, 7.

Cenchrus echinatus L. "purgrass", "sand bur"
eakung, iyakong (N); te kateketeke (K); mouku talatala (T)

Pre-World War I introduction. Trop. America. Common. Annual grass, 10 to 60
cm tall, lower parts often prostrate, rooting at the nodes; leaves, 3 to 10 cm long and 3
to 10 mm wide, narrow, slightly hairy on the upper surface near base, smooth on lower
surface; inflorescence, a dense cylindrical spike-like raceme, 3 to 8 cm long, bearing 5 to
15 well-spaced (not crowded) spiny burs, usually bearing 2 to 4 spikelets, 5 to 7 mm
long; fruit, a globose bur, 3 to 6 mm in diameter, purplish or straw-colored with age,
with numerous irregularly arranged spines, up to 5 mm long. Weed occurring in clusters
or tufts in open and ruderal habitats on the coastal strip. 2, 3(58607), 4(146N), 5(34), 6,
7k 8?

68

Chloris inflata Link "finger grass"
syn. C. barbata sensu auct. non (L.) Sw.

Recent introduction. Trop. America. Occasional. Tufted perennial grass, up to 90
cm tall, erect or somewhat bent at the base, rooting at the lower nodes, which are often
purplish; leaves, up to 3 to 20 cm long and 2 to 6 mm wide, flat, usually bluish-green,
rough on the edges, often hairy near the base on the upper side; inflorescence, flower
heads of 2 to 11 conspicuous, purplish, finger-like, feathery terminal flower spikes, 2.5
to 7.5 cm long, with spikelets, about 3 mm long, with 3 slender bristles, borne at the tips
of flowering stems (culms), up to 60 to 90 cm high. Weed occurring locally in isolated
clusters or tufts in open and ruderal places on the coastal strip. 3(58519), 4(151N),
5(40), 6, 7, 8(9549).

Chrysopogon aciculatus (Retz.) Trin. "needle grass", "seed grass", "golden beard grass"
syns. Andropogon aciculatus Retz.; Rhaphis aciculatus (Retz.) Desv.

Recent introduction. S. E. Asia and Pacific Is. Occasional. Perennial grass, up to
60 cm high, usually with prostrate or creeping stems; leaves, 2 to 5 cm long and 3 to 6
mm wide, flat or folded, linear-lanceolate, rough on the edges; inflorescence, a finely-
branched flower stem, up to 25 cm long, bent at the base, bearing slender erect clustered
purplish panicles, 2.5 to 6 cm long, with awned or barbed, spikelets, 3 to 6 mm long.
Weed occurring locally on bare soil, roadsides, waste places, and occasionally in lawns.
3(58625, 58709), 5(82), 6, 7.

Cynodon dactylon (L.) Pers. “Bermuda grass"
ibugibugi (N); te uteute (K); mouku (T)
syns. Panicum dactylon L.; Capriola dactylon (L.) O. Ktze.

Recent introduction. Old World. Common. Small, low creeping perennial sod-
forming grass, rooting at the lower nodes; leaves, 2.5 to 10 cm long and 3 to 6 mm
wide, rough on the edges; inflorescence, a somewhat ascending or erect flowering stem
(culm), up to 10 to 15 cm, bearing 2 to 6, often 4 to 5, slender, radiating purplish finger-
like terminal flower spikes, 2.5 to 7 cm long, and spikelets, up to 3 mm long. Common
in open places forming mat or sod and in lawns. 3(58625), 5, 6(270), 7.

Dactyloctenium aegyptium (L.) Beauv. "four-finger grass", "beach wire grass",
“crowfoot grass"
syns. Cynosurus aegyptius L.; Dactyloctenium aegyptiacum (L.) Willd. (orth.
mut.); Eleusine aegyptiaca (L.) Desf.

Recent introduction. Paleotropics. Occasional. More or less prostrate annual
grass, 15 to 60 cm high, rooting from the lower nodes; leaves, up to 15 cm to 30 cm

69

long, but mostly shorter, and up to 6.2 mm wide, flat, the edges lined with hairs;
inflorescence, an erect flowering stem (culm), up to 50 cm tall, bearing 2 to 6 radiating
thick, somewhat purplish, dark-colored terminal flower spikes, each up to nearly 3 to 6
cm long, the rachis pointed and extending beyond the spikelets, the spikelets awned, up
to 3 mm long. Weed in clusters or tufts in open and ruderal habitats on the coastal strip.
3(58603, 58606), 4, 5, 6(209), 7, 8(9550).

Dactyloctenium cteniodes (Steud.) Besser?

Recent introduction. Africa. Annual grass. Occasional locally in disturbed places.
4(153N).

Dichanthium sp. "blue grass"

Recent introduction. Occasional. Perennial grass with bluish stems (stolons).
Occurs locally in open weedy places. 3(58518), 5(85).

Digitaria bicornis (Lam.) R. & S. "crab grass", "large crab grass"
syns. Panicum bicorne Lam.

Recent introduction. Trop. Asia. Occasional? Perennial creeping grass. Weed in
open and ruderal sites on the coastal strip. 4(152N).

Digitaria ciliaris (Retz.) Koel. "crab grass", "large crab grass"
syns. Panicum ciliare Retz.; Digitaria adscendens (HBK) Henr.; Syntherisma
ciliaris (Retz.) Schrader

Recent introduction. Trop. Asia. Occasional? Perennial creeping and ascending
grass, rooting by runners at the lower nodes and forming dense mats, with smooth,
usually flattened, ascending stems; leaves, 2 to 13 cm long and 3 to 6 mm wide, flat,
green to blue-green, sometimes purplish, slightly hairy on upper surface; inflorescence, a
branched raceme, 4 to 9 in number, up to about 17 cm long, with 2 to 4 slender finger-
like branches, 2.5 to 8 cm long, remaining close together, forming compact delicate
heads; rachis, winged; spikelets, about 3 mm long, narrow, ellipsoid. Weed in garden.
3(59604).

Digitaria radicosa (Presl) Miq. "crab grass"
syns. Panicum radicosa Presl.; Digitaria borbonica Desv.; D._ timorensis
(Kunth) Balansa

70

Recent introduction. Old World tropics. Occasional? Annual erect or ascending
grass, up to 60 cm high; leaves, 2 to 10 cm long and 3 to 7 mm wide, sheaths puberu-
lent; inflorescence, a branched raceme, 4 to 10 cm long, usually 2 to 3, rarely 6 to 8, in
number, smooth, but puberulent at the axils, with fingerlike, very narrow spikelets, 3 by
0.6 to 0.75 mm, linear-lanceolate, on pedicels, 2.5 to 3.5 mm long. Roadside weed.
3(58608)

Digitaria setigera Roth "crab grass", "slender crab grass"
ibugibugi, ibiugibiugi (N); te uteute (K)
syns. Panicum pruriens Fisch. ex Trin.; Digitaria pruriens (Fisher ex Trin.)
Buse (in Burgess’ list 1935); D. microbachne (J. S. Presl) Henr.

Indigenous. S.E. Asia to Polynesia. Occasional. Densely-tufted creeping or
ascending grass, up to 60 cm high, with smooth slender stems; leaves, 6 to 20 cm long
and 3 to 8 mm wide, smooth hairy, with leaf sheaths at the base; inflorescence, an erect
slender raceme, 5 to 15 cm long, 3 or 4, rarely more, somewhat pubescent, with
numerous spikelets, about 2.5 to 3 mm long. Weed occurring in clusters or tufts in open
and ruderal habitats on the coastal strip. 2, 3, 5(83), 6(156), 7, 8(9551?).

Digitaria violascens Link. "smooth crab grass", "violet crab grass"
syns. D. argyrostachya (Steud.) Fern.

Recent introduction. Trop. Asia. Rare. Erect to spreading annual, sometimes
perennial, smooth-stemmed grass, up to 60 cm tall, forming dense clumps; leaves, 2 to 9
cm long and 3 to 6 mm wide, flat, slightly reddish, usually smooth, but often slightly
rough on the edges and sometimes with a few hairs near the base on the upper side;
inflorescence, composed of 2 to 9 slender finger-like branches, 2.5 to 10 cm long, at the
tip or along the upper 2.5 to 5 cm of the flowering stem, and very small dark-brown
spikelets, up to 2 mm. Weed occurring locally near Buada Lagoon. 3(58786).

Echinochloa colona (L.) Link. "jungle rice"
syns. Panicum colonum L.; Oplismenus colonus (L.) HBK.

Recent introduction. India. Occasional. Tufted erect to ascendant annual grass, 20
to 80 cm high, often decumbent at the base and rooting at the lower nodes, with flattened
often reddish-purple stems; leaves, 2.5 to 15 cm long and 3 to 10 mm wide, flat,
tapering, margins smooth or sometimes scabrous; inflorescence, a flowering stem (culm),
20 to 60 cm high, bearing a panicle, 4 to 12 cm long, with very short green or purple-
tinged alternating racemes, 1 to 3 cm long, with ovate spikelets, 1 to 3 mm long,
crowded in 4 rows on short pedicels. Weed in gardens around buildings. 4(150N), 5
(155).

71

Eleusine indica (L.) Gaertn. "wiregrass", "goosegrass"
ibugibugi (N); te uteute (K); mouku (T)
syn. Cynosurus indicus L.

Pre-World War II introduction. India; long naturalized in Old and New Worlds.
Abundant. Ascending tufted annual or perennial grass, usually growing in tufts, with
flattened pale green stems, with hairy nodes; leaves, 6 to 30 cm long and 3 to 8 mm
wide, mostly glabrous, scabrous on the midvein and upper margins; inflorescence, 2 to 7,
flattened, fingerlike terminal branches or spikes, 4 to 12 cm long and 3 to 7 mm wide,
radiating from the tip of a flowering stem (culm), up to 30 cm tall, with one or two
spikes slightly below the tip; spikelets 3 to 6 mm long; seeds, about 1.5 mm long, dark
green to dark-reddish-brown, ridged, crowded in two rows on one side of the spike.
Growing in rather dense concentrations or colonies in gardens and waste places. 2,
3(58660), 4(147N, 154N), 5(65), 6, 7, 8(9544).

Eragrostis amabilis (L.) Wight & Arn. ex Hook. & ArHove grass", "Japanese lovegrass"
ibugibugi, ibiugibiug, bahibahi (N); te uteute n’ aine (K); mouku (T)
syn. Poa amabilis L.; Eragrostis tenella (L.) Beauv. ex R. & S.

Recent introduction? Old World. Common. Small, rather delicate, annual grass,
usually branched or spreading at the base but sometimes erect, up to 30 cm tall, with
delicate wiry stems, 15 to 45 cm long; leaves, 3.5 to 10 cm long and 1.5 to 4 mm wide,
slightly rough on the upper surface, smooth on the lower, with tufts of long hairs at the
summits of the sheaths; inflorescence, a flowering stem (culm), 5 to 15 cm long, bearing
short, rather delicate, branches and numerous, often reddish-purple, 4- to 6-flowered
spikelets, about 1 to 1.5 mm long, in open panicles. Weed occurring in scattered clusters
around buildings, paths, and other ruderal places. 2(28.5), 3(58626), 4(148N), 5(66), 6,
7, 89539).

Lepturus repens (Forst.f.) R. Br. var. subulatus Fosb"bunchgrass", "beach bunchgrass"
ibugibugi, ibiugibiugi (N); te uteute (K); mouku (T)
syns. Rottboellia repens Forst.f.; Monoerma repens (Forst.f.) Beauv.

Indigenous. Pacific Is. Occasional. Freely-branching, creeping grass with stems,
up to 40 cm long, rooting at the nodes; leaves, 7 to 20 cm long and 3 to 7 mm wide,
narrowly lanceolate to linear, often with inrolled margins; inflorescence, protruding,
solitary, long-awned spikelets embedded in a narrow, jointed, cylindrical flowering spike,
6 to 15 cm long and | to 2 mm in diameter, which disarticulates at maturity. Occurring
in clusters among strand vegetation. 3(58605, 58750), 5(84), 5, 7.

72

Oplismenus hirtellus (L.) Beauv. var. "basket grass"
syns. Panicum hirtellum L.; Orthopogon imbecillus R. Br.; Oplismenus imbecil-
lus (R. Br.) R & S.; O. undulatifolius (Ard.) Beauv.

Recent introduction. Pantropical. Rare. Slender, much-branched creeping or
ascending perennial grass, up to 30 cm or higher; leaves, 5 to 10 cm long and 8 to 12
mm wide, lanceolate, with an acute tip and an asymmetrical base, pubescent, especially
below, blade wavy; inflorescence, small flower clusters, with purplish bristles, borne on
the upper 7 to 15 cm of a erect unbranching flower stem. In open grea in Topside forest.
7(22322).

Panicum subquadriparum Trin.
syn. Brachiaria subquadripara (Trin.) Hitch.

Recent introduction. Trop. Asia. Rare. Perennial creeping grass with stems
(culms) 20 to 60 cm long, rooting at the lower nodes; leaves, 2.5 to 12 cm long and up
to 4 to 10 mm wide, acuminate, nerves pubescent, with a hairy leaf sheath; inflores-
cence, an open, slender, elongated panicle, 5 to 15 cm long, with 2 to 10 spike-like,
solitary, horizontally spreading racemes, 2 to 10 cm long, bearing lanceolate, sharp-
tipped spikelets, 2 to 5 mm long, in 2 rows on one side. Rare around Buada Lagoon.
5(83), 6(155).

Saccharum officinarum L. "sugar cane"
tugage (N)("sugarcane"); te kai tioka, te kai soka ("the sugar tree")(K); kaleve
gau, tolo (T)
syn. S. chinense Roxb. ex Nees in H & A.

Pre-World War I introduction? New Guinea and Trop. Asia. Occasional. Large
clump-forming erect perennial grass with strong thick unbranched stems, 2 to 4 m tall
and 2.5 cm or more in diameter, with short conspicuous internodes filled with solid juicy
pulp; leaves, 45 cm or more long and 3 cm or more wide, smooth, overlapping, finely-
saw-toothed, loosely clustered near the top of stems, with hairy overlapping leaf sheaths;
inflorescence, straight wandlike terminal stalk, panicle about 20 to 50 cm or more long,
borne at the top of each stem, with numerous densely-branched, feathery, pinkish to
silvery-white, many-jointed racemes or flower tassels, with spikelets, 4 to 5 mm long,
although most chewing cultivars rarely flower. Food plant in I-Kiribati and Tuvaluan
gardens at Location and Topside workshops. An important supplementary food plant
throughout most of the Pacific, with the sweet juicy pulp a source of sugar and an
important snack food, the leaves widely used for high quality house thatching, and the
chewing of the fibrous stems reportedly a main factor in good dental hygiene, which is
ironic, given the role of processed sugar in tooth decay. Interspecific hybrids of S.
officinarum and more fibrous wild canes, such as S$. spontaneum and S. robustum, form

73

the basis for the export sugar industries of Fiji and Hawaii. Apparently not traditionally
important on Nauru. 5, 6, 7.

Sporobolus diander (Retz.) Beauv. "Indian dropseed"
syn. Agrostis diander Retz.

Recent introduction. Southern Asia. Rare. Tufted, smooth, slender, erect
perennial grass, 30 to 70 cm high, arising from an abruptly bent base; leaves, 7.5 to 30
cm long and 2 to 6 mm wide; inflorescence, a smooth and sometimes drooping, spike-
like, flowering stem or panicle, 10 to 30 cm long and 5 cm wide, bearing numerous open
feathery branches, 1.5 to 4 cm long, with dull grey spikelets, 1 to 2 mm long, and brown
seeds, less tha 1 mm long. Weed in waste place on coastal strip. 6(154).

Stenotaphrum micranthum (Desv.) Hubb.
syns. Ophiurinella micrantha Desv.; Stenotaphrum subulatum Trin.

Indigenous. Mascarene Is. in the Indian Ocean through Malesia to eastern
Polynesia and the Marshall Islands in Micronesia. Erect to ascending, tufted, freely
branching grass, up to 45 cm high, with prominent nodes and slightly compressed
sheaths, rooting at the lower nodes; leaves, 2.5 to 12 cm long and 5 to 12 cm wide,
lanceolate, finely pointed, glabrous; inflorescence, 3 to 15 cm long, a slender to rather
stout, corky cylindrical raceme; spikelets, 2 to 4 on short branchlets, 1.5 to 3 mm long,
oblong to oblong-lanceolate, obtuse, embedded in the rachis. Rare grass, cited by
Fosberg et al. 1987, but not seen or collected in 1979 or 1980.

Tricholaena rosea Nees "Natal grass", "Natal red top"
syns. Rhynchelytrum roseum (Nees) Stapf. & Hubb.; Rhynchelytrum repens
sensu auct. non (Willd.) Hubb.; Tricholaena repens (Willd.) Hitchc.

Recent introduction. S. Africa. Common. Erect perennial grass, 40 to 90 cm high,
rooting at the lower nodes, usually much-branched at the base, with green to bluish-green
stems, often purplish at the joints; leaves, 5 to 15 cm long and 2 to 6 mm wide, flat
bluish-green; inflorescence, a feathery panicle, 8 to 15 cm-long, bearing dark red or
purplish flowers, which fade to silver-pink when old, borne on fine ascending branchlets;
spikelets, about 5 mm long, clothed with fine silky hairs, about 3 to 5 mm long. Found
in clusters in waste places on coastal strip and plateau and occasionally in mined areas.
3(58655), 4(170N), 5(49), 6, 7, 8(9567).

74
PONTEDERIACEAE (Pickerel Weed Family)

Eichhornia crassipes (Mart. & Zucc.) Solms-Laub. “water hyacinth"
syns. Pontederia crassipes Mart. & Zucc.; Eichhornia speciosa Kunth

Recent introduction. Trop. and Subtrop. America. Rare. Usually floating,
perennial herb, up to 60 cm tall, with masses of fleshy, more or less horizontal, black
roots, rooted only at flowering time by long slender roots; leaves, up to 7 to 10 cm wide,
rounded or oblong, on bulbous air-filled petioles which act as floats; inflorescence, a
flower spike, about 40 cm long, bearing showy 6-petaled pale violet flowers, about 5 cm
in diameter, with a blue patch with a spot of bright yellow on the larger upper lobe,
about eight to a stem, rising above the leaves; some forms have pink and yellow flowers.
Water weed in Buada Lagoon and planted in tubs at Location. Commonly planted as an
ornamental; has escaped to become a serious pest in many areas of the world where it
clogs rivers and causes flooding and obstructs navigation. All parts are reportedly edible
6(255).

STRELITZIACEAE (Bird of Paradise Family)

Heliconia collinsiana R.F. Griggs “hanging heliconia", "fish-pole heliconia"
syn. H. pendula Wawra

Recent introduction. Guatemala. Rare. Erect herb, up to 2 m tall; inflorescence,
up to 30 cm or more long, hanging, almost plastic-looking, parrot-beak-, or claw-like,
with bright red flower bracts, each 5 cm or longer, with yellow and green lower
margins, and white waxy powder. Planted ornamental. 5, 6.

Heliconia humilis (Aubl.) Jacq. "heliconia", "lobster claw"
syn. Musa humilis Aubl.

Recent introduction. Trop. S. America. Rare. Erect herb, less than 2 m tall;
leaves, broad, paddle-shaped, 3 to 6 in number; inflorescence, an erect flowering stalk,
up to 120 cm tall, among the leaves, bearing pointed lobster-claw-shaped bright red
flower bracts with dark green upper margins, each up to 12 cm long. Planted ornamental.
5, 6(236).

IS

Heliconia psittacorum L. "heliconia"

Recent introduction. Trop. S. America. Occasional. Erect herb, less than 1.5 m
tall; leaves, few, 3 to 5 in number; inflorescence, an erect flowering stalk bearing small
bright orange flowers with green tips. Planted ornamental. 5, 6, 7.

Heliconia sp. "heliconia"

Recent introduction. Trop. America. Rare. Erect herb. Planted ornamental. 5
(174).

TACCACEAE (Polynesian Arrowroot Family)

Tacca leontopetaloides (L.) O. Kuntze "Polynesian arrowroot"
damagmag, damogmog (N); te makemake (K); masoa, vatia (T)
syns. Leontice leontopetaloides L.; Tacca pinnatifida Forst.

Aboriginal introduction. Paleotropics. Occasional. Large stemless herb, up to 1 m
high; leaves, large, palmately 3-parted, deeply-lobed, on long petioles, 60 cm or more in
length, emerging directly from starchy underground tubers resembling potatoes, 20 to 25
cm in diameter; inflorescence, a hollow flower stalk (scape), up to over 1 m high,
bearing a terminal umbel or cluster of 10 to 40 hanging green flowers, about 15 mm
long, surrounded by 6 leafy bracts, 4 to 5 cm long and 2 cm wide, and numerous
hanging green and purplish filaments, up to 25 cm long; fruit, 2 to 2.5 cm in diameter,
fleshy, ribbed, globose, which turns yellow when mature, and is crowned with floral
parts. Occurring spontaneously in old gardens and in escarpment forest. Tubers grated
and washed to eliminate poisonous substances, and made into edible starch in the past,
but apparently not used by Nauruans to the extent that it was used in other parts of
Micronesia and Polynesia. Paste from tuber used as an adhesive for barkcloth and other
handicrafts in Polynesia and Melanesia and the fibers from the flower stem for weaving
in parts of Polynesia. 5, 6(119).

ZINGIBERACEAE (Ginger Family)

Alpinia purpurata (Vieill.) K. Schum. "red ginger"
syns. Guillainia purpurata Vieill.; Languas purpurata (Vieill.) Kaneh.

76

Recent introduction. Indonesia to Pacific Is. Occasional. Erect herb, 1.2 to 4 m
tall, with leafy stems; leaves, up to 30 cm long, lance-like; inflorescence, a flower spike,
about 25 to 30 cm long, erect or drooping, with large, open, dark red bracts, each
accompanied by a small inconspicuous white ephemeral flower, about 2.5 cm long, with
new plantlets, which sprout among the bracts, taking root as the dying flower spike
collapses to the ground. Planted ornamental. 3(59710), 5, 6, 7.

Alpinia zerumbet (Pers.) Burtt & R.M. Smith "shell ginger"
syns. Costus zerumbet Pers.; Alpinia nutans (Andr.) Roscoe; A. speciosa
(Wendl.) K. Schum.; Catimbium speciosum (Wendl.) Holttum.

Recent introduction. S.E. and E. Asia. Rare. Erect herb, up to 2.5 m tall; leaves,
50 to 70 cm long and 7 to 15 cm wide, oblong-lanceolate; inflorescence, an arching
flower stem, 1.5 to 3 m long, bearing a somewhat lax or pendant open flower cluster.
about 25 cm long, bearing irregularly bell-shaped flowers with waxy white bracts and
red-tipped corollas, up to 5 cm long, with yellow lips with reddish lines or veins; fruit,
about 2 cm in diameter, globose, ridged, red. Planted ornamental. 6.

Hedychium coronarium Koen. "white ginger"

Recent introduction. India. Rare. Erect herb with stems, up to 1 m tall; leaves,
about 50 to 60 cm long and 10 cm wide, narrowly oblong or lance-shaped, with short
petioles; inflorescence, about 20 cm long, dense, spike-like, terminal, with overlapping
bracts, each bearing 2 to 3 fragrant white flowers, 6 to 8 cm long, with slender tubes and
a pale green spot on the lip, which turn yellow with age. Planted ornamental and pot
plant. 3(59671), 5, 6, 7.

Nicolaia elatior (Jack) Horan. "torch ginger"
syns. Alpinia elatior Jack; Phaeomeria speciosa (Bl.) Koord; P. magnifica
(Roscoe) K. Schum

Recent introduction. Mauritius. Rare. Large clump-forming perennial herb, up to
2 to 5 m high, with arching leaf stems; leaves, 25 to 60 cm long and 10 to 15 cm wide,
pointed, numerous, alternating in two rows up the stem; inflorescence, highly ornamen-
tal, consisting of numerous red, white-margined bracts, the basal bracts being large,
waxy, flowerless and forming a nest for a cone-like flowerhead, about 12 cm long,
composed of numerous overlapping bracts, which are spirally arranged and accompanied
by small flowers. Planted ornamental. 6.

T

Zingiber officinale Roscoe "ginger"
keung (C)
syn. Z. zingiber Karst.

Pre-World War II introduction. India and China. Rare. Erect smooth herb, up to
90 cm high, arising from edible, thick, hard, often palmately-branched, pale-yellow
(within) rootstalks or rhizomes, about 1.5 to 2.5 cm in diameter; leaves, 5 to 30 cm long
and 2 to 3 cm wide, annual, lance-shaped, sheathed at the bases; inflorescence, a
flowering stem, about 6 to 12 cm high, bearing flower spikes, 4 to 7 cm long and 1.5 to
2.5 cm wide, with bracts, 2 to 3 cm long, and greenish-yellow three-lobed flowers, about
1.5 to 2.5 cm long. Planted in Chinese food garden at Location. Rhizome used as a
spice. An increasingly important commercial crop for export and local processing in Fiji.
Dp, (05

Zingiber zerumbet (L.) Sm. "wild ginger"
syn. Amomum zerumbet L.

Recent introduction; reintroduced recently by Fijian expatriate community Trop.
Asia. Rare. Erect perennial herb, up to 1.5 m tall, with leafy stems rising from tuberous
aromatic rootstalks or rhizomes; leaves, mostly 15 to 30 cm long, usually shorter, oblong
lance-shaped, sheathing at the bases; inflorescence, a spike-like flower stalk (scape), up
to 30 cm long, arising from the rhizomes, bearing ovoid to cylindrical reddish or green
flower heads, 5 to 20 cm long, with many bracts, 2 to 3 cm long, and inconspicuous
three-lobed white to cream flowers, about 5 cm long. Planted ornamental or medicinal
plant. An important aboriginal introduction throughout much of Melanesia and Polynesia,
where it is an important medicinal plant. 3, 6.

78

DICOTYLEDONAE

ACANTHACEAE (Acanthus Family)

Asystasia gangetica (L.) Anders. "asystasia", "Chinese violet"
syns. Justicia gangetica L.; Asystasia coromandeliana Nees

Recent introduction. Paleotropics. Occasional. Perennial trailing herb, or sub-
shrubby plant, with ascending stems, up to 30 to 50 cm or longer, especially if climbing
among taller vegetation; leaves, opposite, ovate to heart-shaped, up to 4 to 6 cm long;
flowers, 6 to 10, tubular or bell-shaped, somewhat narrower or curving upwards at the
base, and borne in racemose clusters, up to 15 cm or more long, with a 5-parted calyx,
and a light-violet and white, or yellow, in some varieties, corolla, about 2 to 3 cm long,
with five broad spreading lobes; fruiting capsule club-shaped, four-seeded. Planted
ornamental and naturalized in some ruderal places. 3, 4(127N), 5, 6(231), 7.

Asystasia sp.

Recent introduction. Pot plant. 3(58702).

Barleria cristata L. "Philippine violet", "bluebell barleria"

Recent introduction. India. Rare. Small erect shrub, 60 cm to 1.2 m or higher,
with downy branches; leaves, 2.5 to 10 cm long rough, hairy, short-stemmed, oval and
pointed at both ends; flowers, 1 to 3, attractive stemless and funnel-shaped, with a four-
parted calyx, about 2 cm long, and a five-lobed violet or white and violet corolla, about
5 to 7 cm long, developing at the leaf axils, each accompanied by 2 spiny-edged green to
white narrow lanceolate bracts, about 2 cm long. Planted ornamental. 3(58797), 6.

Barleria prionitis L. "porcupine flower"

Recent introduction. Paleotropics. Occasional. Erect shrub, up to 1 m high;
leaves, 5 to 6 cm long by about 2 cm wide, elliptic to ovate or obovate, obtuse, acute or
even somewhat petiolate, narrowed to base; flowers, up to 3.5 cm long, with yellow
corollas and bracts converted into three-forked spines, 1 to 2.5 cm long. Planted

79

ornamental and naturalized in ruderal sites and on disturbed slope below plateau.
3(68772);:6, 7.

Blechum brownei Juss.
syns. Barleria pyramidatum Lam.; Blechum pyramidatum (Lam.) Urb.

Recent introduction. Peru. Rare. Erect, prostrate or ascending branched shrub, 10
to 50 cm tall, with pubescent stems; leaves, 2 to 7 cm long, simple, ovate to lanceolate,
pubescent, pointed at both ends, and on short petioles; flowers borne on terminal spikes,
2 to 5 cm long, with greenish or whitish overlapping ovate bracts, 1 to 1.5 cm long,
concealing a white or pale violet funnel-shaped, five-lobed corolla, scarcely longer than
the bracts; seed capsule oblong, 5 to 6 mm long. Weed in lawns. 5(60), 6.

Crossandra infundibuliformis (L.) Nees "crossandra"
syns. Justicia infundibuliformis L.; Crossandra undulaefolia Salisb.

Recent introduction. India. Rare. Shrub, 30 to 90 cm high; leaves, 7.5 to 12.5 cm
long, shiny green, narrow, ovate, wavy-margined, and pointed at both ends; flowers
borne on long narrow downy spikes, about 10 cm long, with overlapping bracts bearing
showy tubular salmon-orange flowers with a five-lobed calyx and a corolla with a narrow
tube, about 2 cm long, and one lip which is 2.5 to 5 cm across and has 3 to 5 shallow
lobes; fruiting capsule, oblong, containing 4 scaly seeds. Planted ornamental. 6.

Eranthemum pulchellum Andr. "blue eranthemum"
syns. Justicia nervosa Vahl; Eranthemum nervosum (Vahl) R. Br.

Recent introduction. India. Rare. A smooth shrub, 60 cm to 1.5 m high; leaves,
10 to 20 cm long, oval, pointed at both ends, with prominent veins and shallow-toothed
margins; flowers borne on narrow spikes, 2.5 to 7.5 cm or longer, with overlapping
whitish, green-veined oval bracts bearing a five-lobed calyx and a narrow-tubular bright-
blue corolla, up to 2.5 cm long, the five lobes overlapping spirally in bud, but spreading
to a diameter of under 2 cm; fruiting capsule ovoid or oblong, 4-seeded. Planted
ornamental. 6.

Fittonia argyroneura Coem. "snail plant", "nerve plant", "silver-net leaf"
syn. F. verschaffeltii var. argyroneura Nichols.

Recent introduction. Peru. Rare. Perennial creeping herb, up to 30 cm high,
rooting at the joints; leaves, 5 to 10 cm long and 2.5 to 6 cm wide, broad, ovate,
ornamental, with a network of fine white veins; petioles 0.5 to 3 cm long; inconspicuous
flowers borne on spikes, up to 6 cm long, with obovate bracts, less than 1 cm long, with

80

a calyx about 0.5 cm long and a two-lipped tubular yellow corolla, 1.5 to 2 cm long.
Ornamental pot plant. 6. ;

Fittonia verschaffeltii (Hort. ex Lemaire) Coem. "snail plant", "nerve plant",
"painted net-leaf"

Recent introduction. Peru. Rare. Perennial creeping herb, up to 30 cm high,
rooting at the joints; leaves, 6 to 12 cm long and 3 to 8 cm wide, broad, smooth, ovate,
ornamental, with a network of fine pink to bright carmine veins; petioles 0.5 to 3 cm
long; inconspicuous flowers borne on erect narrow spikes, up to 6 cm long, with
overlapping obovate greenish bracts, less than 1 cm long, a calyx about 0.5 cm long, and
a tubular yellow two-lipped corolla, 1.5 to 2 cm long. Ornamental pot plant. 6.

Graptophyllum pictum (L.) Griff. “caricature plant", "morado"
syns. Justicia picta L.; Graptophyllum hortense Nees

Recent introduction. New Guinea. Rare. Erect shrub, up to 2 m high; leaves, 8 to
20 cm long and 3 to 13 cm wide, smooth, leathery, variegated, elliptic-oblong, opposite,
with slightly wavy margins, and pointed at both ends, some green, with irregular
yellowish blotches or markings in the center and other varieties with red-purple leaves
with pinkish to cream-colored blotches along the center; petioles 1 cm long or less;
flower spikes with small bracts bearing clusters of flowers with a short 5-small-lobed
calyx and a 2-lipped, funnel-shaped crimson-purple corolla, about 4 cm long, the upper
lip with 2 short lobes, the lower with 3 long narrow lobes; fruiting capsule 2-seeded.
Planted ornamental. 6, 7.

Hemigraphis alternata (Burm.f.) T. Anders. "cemetery plant"
syns. Ruellia alternata Burm.f.; Hemigraphis colorata (Bl.) Hall.f.; Ruellia
colorata Bl.

Recent introduction. Java. Rare. Creeping perennial herb, rooting at the joints;
leaves, 2 to 8 cm long, ovate to heart-shaped, metallic- to greenish-purple above and
reddish-purple beneath, and with sunken veins and scalloped margins; petioles nearly as
long as blades; erect flowering spikes, 2 to 3 cm long, with crowded overlapping narrow
purple bracts, about | cm long, bearing a calyx with 5 narrow lobes, each less than 1 cm
long, and a white, purple-lined corolla, about 2 cm long; fruiting capsule slender, 4- to
20-seeded. Planted ornamental ground cover. 6.

Justicia fulvicoma Schlecht. & Chamisso "shrimp plant", "red shrimp plant"

syns. Beloperone guttata Brand. non Wallich; Justicia brandegeana Wassh. & L.
B. Smith; Drejerella guttata (Brandeg.) Bremek.

81

Recent introduction. Mexico. Rare. A weak-stemmed much-branched shrub, 60
cm to 2 m or higher; leaves, 2.5 to 6 cm long, opposite, entire, pointed, ovate to heart-
shaped, somewhat shiny-green and hairy; petioles, 1 to 2 cm long; flower spikes, up to
10 cm long, consisting of conspicuous brick-red, heart-shaped overlapping downy bracts,
1 to 1.5 cm long, those near the tip yellow-green with red veins, from which protrude
small white flowers with a calyx with five narrow lobes, and a white corolla, 2.5 to 4 cm
long, two-lipped to near the middle, with two rows of maroon spots on the lower, shortly
three-lobed lip; fruiting capsule, club-shaped and four-seeded. Planted ornamental and pot
plant. 3(58720), 5, 6.

Nicoteba betonica (L.) Lindau “white shrimp plant", "squirrel’s tail"
syn. Justicia betonica L.

Recent introduction. Trop. Africa to Malaya. Rare. Weak-stemmed shrub, up to
1.5 m or higher; leaves, 7.5 by 3.5 cm or much larger, ovate or narrower, acuminate,
smooth, dark green; flower spikes, about 10 cm long, borne at branch tips in erect
spikes, with white and lilac flowers, each accompanied by three conspicuous overlapping
heart-shaped and ovate pointed bracts which are white with green veins. Planted
ornamental. 6, 7.

Odontonema strictum (Nees) O. Ktze. "odonotema", "red justicia"
syns. O. tubiforme (Bertol.) O. Ktze.; Justicia tubaeformis Bertol.; Thrysacan-
thus strictus Nees; O. nitidum (Jacq.) O. Ktze.; Justicia coccinea Aubl.(?)

Recent introduction. C. America. Occasional. Erect shrub, up to 1.5 m tall;
leaves, mostly 5 to 20 cm long and 4 to 9 cm wide, ovate-oblong or lanceolate-oblong,
wavy margins, and smooth above and slightly hairy beneath; petioles, 0.5 to 2 cm long;
branching terminal flower spike, with very small bracts and straight slender bright-red
corolla tubes, 2.5 cm long and only about 5 mm across, with 5 lobes, each about 3 mm
long, crowded at branch tips. Planted ornamental. 5, 6, 7.

Pachystachys lutea Nees "yellow shrimp plant"

Recent introduction. Brazil. Rare. Erect branched shrub, up to 45 cm tall; leaves
rich, dark-green, elliptical, and pointed; stems, erect, each terminating in a striking 2.5
to 10 cm-long cone-like spike of conspicuous rich yellow overlapping bracts, from which
protrude white tubular flowers, about 4 cm long. Pot plant. 3(58720).

82

Pseuderanthemum bicolor (Schrank) Radlk.
syn. Eranthemum bicolor Schrank

Recent introduction. E. Malaysia. Rare. Erect shrub, up to 1 m tall; leaves,
mostly 5 to 15 cm long and up to 6 cm wide, purple or purple-brown, ovate-oblong or
lanceolate-oblong, acuminate, glabrous or finely puberulent; flowers borne in axillary
cymes, the uppermost sometimes in racemes, with puberulent calyxes, up to 1 cm long,
and puberulent 5-lobed white tubular corollas, 3.5 to 4 cm long with red throats, one
lobe with a red blotch, and other lobes white; fruiting capsule 2.5 to 3 cm long. Planted
ornamental. 6, 7(27810).

Pseuderanthemum carruthersii (Seem.) Guill. var. carruthersii "false eranthemum"
te iaro (K)
syns. Eranthemum eldorado Hort.; Pseuderanthemum eldorado (Williams)
Radlk.

Recent introduction. Melanesia? Occasional. Erect shrub, up to 1 m or higher;
leaves, 5 to 15 cm long, smooth, elliptic-ovate leaves, mottled yellow-green, with
conspicuous netlike veins; petioles 1 to 3 cm long; erect spike-like racemes bearing
flowers with inconspicuous bracts, green or reddish 5-lobed calyxes, up to 8 mm long,
and white and purple or rosy-purple tubular corollas, the tube, about 1.3 cm long, and
the 5 lobes spreading to about 2.5 cm or more in diameter; fruiting capsule, four-seeded,
club-shaped. Planted ornamental shrub. 3, 5, 6(201), 7.

Pseuderanthemum carruthersii ( Seem.) Guill. var. atropurpureum (Bull) Fosb.
"purple false eranthemum", "false face" te iaro (K); lakauuli (T)
syns. P. atropurpureum (Bull) Radlk.; Eranthemum atropurpureum Bull.; P.
versicolor (Hort.) Radlk.; Eranthemum versicolor Hort.

Recent introduction. Melanesia? Occasional. Erect shrub, up to 1.5 m or more
high; leaves, 5 to 15 cm long, dark purplish-red, elliptic-ovate; petioles, 1 to 3 cm long;
erect spike-like racemes bearing flowers with inconspicuous bracts, green or reddish 5-
lobed calyxes, up to 8 mm long, and 4-lobed white and purple or rosy-purple tubular
corollas, the tube and lobes about 1.3 cm long; fruiting capsule, four-seeded, club-
shaped. Planted ornamental shrub. 3(58777, 58792), 5, 6, 7.

Sanchezia speciosa Leonard "sanchezia"
syn. S. nobilis sensu auct. non Hook.f.

Recent introduction. Ecuador. Rare. Erect shrub, up to 1 m or higher, with
smooth four-angled branches; leaves, 6 to 45 cm long, oval, pointed at both ends, and
green, with yellow mid- and side veins; narrow flowering spikes bearing wide red bracts,

83

about 2.5 cm long, each pair with clusters of 8 to 10 narrow bright-yellow tubular
flowers with corollas about 5 cm long and 5-lobed calyxes; fruiting capsule, narrow,
cylindrical, 6- to 8-seeded. Planted ornamental shrub 6(161).

Thunbergia alata Bojer ex Sims "black-eyed Susan"

Recent introduction. Trop. Africa. Rare. Herbaceous or slightly woody pubescent
climbing or trailing vine with angular stems, up to 2 m or longer; leaves, 4 to 8 cm long,
mid-green, opposite, deltoid ovate or ovate-lanceolate, pointed, palmately-veined,
irregularly toothed; winged petioles, about as long as the blade; flowers solitary on
axillary stems (pedicels), up to 5 cm long, 2 large bracts, about 2 cm long, calyx ring-
like, corolla, yellow-orange to cream, 3 to 4 cm across, with a purple tube, 2 cm long;
fruiting capsule depressed-globose, beaked, about 1 cm long; seeds warty and ribbed.
Planted ornamental vine. 5.

Thunbergia erecta (Benth.) T. Anders. "bush thunbergia"
syn. Meyenia erecta Benth.

Recent introduction. Trop. W. Africa. Occasional. Erect, somewhat sprawling
shrub, up to 2 m tall, with 4-angled stems; leaves, 3 to 8 cm or longer, opposite, oval-
ovate, palmately 5- to 7-nerved; petioles, slender, up to 4 cm or longer, winged nearly to
base; flowers solitary on axillary stems (pedicels), 1 to 2 cm long, with 2 large bracts, up
to 2 cm long, calyx, 3 to 6 mm long, corolla curved, funnel-form, 3.5 to 6 cm in
diameter and tube 2 or 3 cm long, irregularly 5-lobed, deep blue-violet or white with a
yellow throat; fruiting capsule 2 to 2.5 cm long. Planted ornamental erect shrub.
3(58700), 5(110), 6, 7.

Thunbergia grandiflora (Roxb. ex Rottler) Roxb."Bengal clock vine", "Bengal trumpet",
"large-flowered thunbergia"
syn. Flemingia grandiflora Roxb. ex Rottler

Recent introduction. India. Rare. Climbing vine; leaves, 7.5 to 20 cm long, dark
green, nearly as broad, cordate, palmately lobed, 5 to 7- nerved, pubescent; petioles, 4 to
12 cm long; flowers in pendent racemes, calyx pubescent and reduced to a narrow ring,
corolla violet or whitish, 6 to 8 cm in diameter, the tube pale yellow, 3 to 3.5 cm long;
capsule up to 1.5 cm, the beak to 3 cm long. Planted ornamental. 6(187).

84
AMARANTHACEAE (Amaranth Family)

Achyranthes canescens R. Br.
syn. A. velutina H. & A.

Indigenous? Extinct? Somewhat shrubby herb, up to | m tall; leaves, up to 12 cm
or longer, opposite, ovate or elliptic, acuminate, slightly blunt; erect unbranched flower
spikes bearing spiny bracts, often pinkish or purplish, and greenish calyx and corolla
green, about 6 mm long; fruit, 5-seeded, adherent to fur or clothing. Reported by
Schumann (1888) as collected by Finsch; reported present by Burges, 1933; not seen
since. Extinct? 2.

Alternanthera ficoidea L. var. bettzickiana (Reg.) Backer "joyweed", "alternanthera",
"telanthera", "calico plant"

syns. Telanthera bettzickiana Reg.; A. versicolor Reg.; Alternanthera bettzick-
iana (Reg.) Nichols.

Recent introduction. Brazil. Rare. Small erect variable herb with red branching
stems, 15 to 30 cm high; nodes usually with tufts of white hair; leaves, about 2.5 cm
long, opposite, lanceolate-ovate or ovate, variegated green, pink, red, yellow, and
sometimes cream; petiole equal to or longer than the blade; flowers, small white in
sessile heads in axils of the upper leaves. Planted ornamental border. 6.

Alternanthera sessilis (L.) R. Br. ex R. & S. "joyweed"
syns. Gomphrena sessilis L.; Alternanthera denticulata R. Br.; A. nodiflora R.
Br.; A. amoena (Lem.) Voss

Recent introduction. Pantropical. Rare. Spreading or semi-prostrate branching
herb, up to about 50 cm long, rooting at the lower nodes; internodes with 2 pubescent
lines; leaves, 1 to 6 cm long and 0.4 to 1 cm wide, opposite, green, narrowly elliptic-
obovate or oblanceolate, obtuse or acutish, minutely toothed; petiole up to about 1 cm
long; flowers, white, small, stemless (sessile) in compact headlike clusters in the leaf
axils; fruit enclosed in a small bladder which does not split open. Garden weed. 6(188).

Amaranthus dubius Mart. ex Thell. "spleen amaranth"

Recent introduction. Trop. America. Occasional. Small erect herb, up to about 70
cm high; stem, smooth, except upper portions which are slightly hairy; leaves, 3.5 to 8
cm long and 2 to 5 cm wide, ovate, smooth on both surfaces, margin smooth; petiole, 2
to 7 cm long; flowers, greenish, in dense axillary and terminal clusters, male and female

85

flowers separate, each with 5 sepals; seeds, small, oval, black. Weed in home gardens
and at Location. 3, 6, 7(22309, 22312).

Amaranthus hypochondriacus L. "prince’s feather"
syn. A. hybridus L. var. hypochondriacus (L.) Robins.

Recent introduction. Trop. America. Rare. Smooth erect herb, up to 1 m or
higher; leaves, bright red-purple to purple-green; flowers, bright red-purple in thick
showy feathery panicles. 6.

Amaranthus spinosus L. "spiny amaranth", "thorny amaranth"
ma si han (C)

Pre-World War II introduction? Pantropical. Occasional. Smooth erect, branched
herb, 25 to 70 cm high, armed with a pair of sharp needle-like spines, about 1 to 2 cm
long, at the base of each petiole; leaf blades, 2.5 to 10 cm long and 1.5 to 4.5 cm wide,
green, alternate, broadly lanceolate, pointed at the tip; petioles, about as long as leaf
blades, clasping the stem; female flowers, pale green, clustered at leaf axils; male
flowers, pale green, borne in terminal panicles; fruit a circumscissile utricle; seeds, very
small, lens-shaped, shiny dark-brown. Weed in waste places and in gardens; occasionally
cultivated in Chinese contract workers’ gardens at Location. Used as a medicinal plant
and leaves reportedly occasionally cooked for spinach by Chinese. 5, 6(140).

Amaranthus tricolor L. "Joseph’s coat", "Chinese spinach". "amaranth", "pigweed"
te mota, te moota (K); in ts’oi (C)
syns. A. gangeticus L; A. melancholicus L.

Pre-World War II introduction? Trop. Asia. Occasional. Erect unarmed,
branching herb, up to 1.5 m tall; leaves, 3 to 7 cm long, oval, pointed to blunt, long-
petiolate, green or blotched with red or purple; flowers in rounded stemless clusters in
leaf axils and in terminal panicles, about 5 cm long; fruit, a 1-seed utricle; seed, small,
disc-shaped, dark brown, shiny. Food plant in Chinese gardens at Location and Topside
workshops. 5(27), 6.

Amaranthus viridis L. "slender amaranth", "green amaranth", "pigweed"
syn. A. gracilis Desf.

Recent introduction. Pantropical. Occasional. Erect or decumbent, unarmed herb,
up to 20 to 60 cm high; stems, reddish, longitudinally-grooved; leaf blades, 3 to 7.5 cm
long and 2.5 to 5 cm wide, alternate, green above, lighter below, broadly lanceolate or
ovate, obtuse or slightly notched at tips, smooth on both surfaces; petioles, 2.5 to 5 cm

86

long; flowers, greenish to purplish, in small dense axillary clusters and terminal panicles;
fruit, a l-seeded utricle; seed, small, disc-shaped, dark brown. Weed in waste places,
primarily at Location. 4(143N), 5(26), 6.

Celosia argentea L. var. cristata (L.) Ktze. "cock’s comb"
syn. C. cristata L.

Recent introduction. Trop. Americas. Erect annual herb, 30 cm to 75 cm high,
stems usually ribbed; leaves, 2 to 16 cm long and up to 5 cm wide, although usually
smaller, alternate, narrow lanceolate to ovate, often reddish-green; inflorescence, a
dense, flatly-crested, elongated, fan-shaped flower head, 6 to 15 cm across, bearing
numerous tiny flowers, pinkish or yellow, but usually magenta or bright red; each fruit
with two small, glossy, black seeds. Rare. Planted ornamental. 7.

Gomphrena globosa L. "globe amaranth", "pearly everlasting"

Recent introduction. Trop. America. Rare. Erect branched annual herb, up to 45
cm high; leaves, 10 by 5 cm or smaller, opposite, obovate-oblong, pale-green, hairy,
petioles clasping; flowers, numerous, crowded, in globose heads, about 1.5 to 2.5 cm in
diameter, white, yellow, pink, red, purple or variegated, borne at the ends of branches,
each head having two leafy bracts. Planted ornamental. Flowers used by Nauruans in
head garlands and other ornamentation. 5(93)

Iresine herbstii Hook.f. "iresine", "bloodleaf", "achyranthes"

Recent introduction. Brazil. Rare. Fast-growing, erect shrubby perennial herb, up
to 30 to 70 cm high, with bright red stems and branches; leaves, up to 6 cm long,
opposite, fleshy, round to heart-shaped, the tip blunt or notched, brilliant red-purple to
bronze-red, with wide, arching prominent veins, the blade continuing down the petiole;
flowers, which may not develop, greenish-yellow, minute, numerous, in fine, much-
branched panicles, up to 50 cm long. Planted ornamental. 6.

ANACARDIACEAE (Cashew or Rhus Family)
Mangifera indica L. "mango"
damanko (N); te mangko (K); mago (T)

Pre-World War I introduction. Indo-Burma. Common. Large dense, broad-
crowned tree, with dark roughened bark, up to 20 m high; leaves, 10 to 35 cm long and

87

2 to 8 cm wide, green to light-green, alternate, spirally arranged, simple, oblong-
lanceolate, acute to acuminate, leathery (coriaceous) and somewhat shiny, pinnately-
nerved, midrib prominent, young leaves often reddish; petioles up to 5 cm or more long,
somewhat flattened on the upper surface; flowers, small, about 4 to 8 mm in diameter,
numerous, yellowish-white to pinkish-white, 4- to 5-parted corolla, densely covered with
short yellowish hairs, in large branched terminal panicles, 10 to 50 cm long; fruit, 8 to
20 cm long, fleshy, fibrous, ovoid, pointed or rounded edible drupe, with thick, smooth,
green to yellow-orange or reddish skin, sweet, juicy, yellow-orange to dark orange pulp,
and a large, flattened, fibrous, ribbed seed case containing one seed. Large fruit and
shade tree planted in home gardens and found growing in mature spontaneous stands near
Buada Lagoon. Fruit eaten ripe and green, with ripe fruit occasionally made into jam on
Nauru; used for firewood. The leaves, and sap from leaves and fruit, can cause an
allergic rash. Common introduced fruit tree, found in houseyard gardens, agricultural
areas and naturalized throughout Melanesia, Polynesia and Micronesia, possibly an
aboriginal introduction in some areas, but not on Nauru; an important cash crop for local
sale and export in Polynesia, Melanesia and the larger islands of Melanesia. 2, 3(58643),
5(147), 6, 7.

Spondias dulcis Park. "Polynesian vi apple", Polynesian plum", "Otaheiti apple"
dagimadere, "Egigu’s tree" (N)
syn. S. cytherea Sonn.

Aboriginal introduction? Pacific Islands. Rare. Medium to large stiff-branched,
smooth, grey-barked, deciduous tree, up to 15 m or taller; leaves, 20 to 40 cm long,
alternate, odd-pinnate, clustered at branch ends; leaflets, up to 8 cm or longer and 2.5
cm wide, commonly 4- to 12-paired, with a single unpaired terminal leaflet (8 to 25 in
total), shiny, bright green, oval-lanceolate or oblong, acuminate, base acute, entire to
finely toothed (crenulate); flowers, tiny, 5-parted, male, female and perfect, develop in
large, lax crowded panicles; fruit, up to 8 cm or longer, oval-obovate edible drupe with
green to yellow-orange skin and light green to dark yellow pulp, surrounding a single, 5-
ridged, fibrous seed case with 1 to 5 seeds. Reported present by Burgess in 1935. The
tree, which formerly stood near Buada Lagoon, was reportedly damaged during World
War II, and although the Nauruans tried to save it by shoring it up with cement, it died
shortly thereafter. Four seeds sent by R. Thaman to J. Audoa in 1981 in an attempt to
reintroduce S. dulcis, but the result of plantings is unknown. One tree, 3 m tall, seen re-
established in fenced food garden surrounding Buada Lagoon in July 1987. Ripe fruit
eaten. Tree features in the well-known Nauruan legend concerning a young woman,
Egigu, who became the Nauruans’ "woman in the moon", after climbing the tree
(dagimadere), restoring the sight of a blind women Enibarara who lived at the top, and
marrying her third son, Maramen (the moon). 2, 7.

88

Spondias mombin L. ‘ "hog plum"
syn. S. lutea L.

Extinct? Trop. Asia. Reported present by Burges. Possibly a doubtful iden-
tification. 2.

ANNONNACEAE (Custard Apple Family)

Annona muricata L. “soursop"
dawatsip (N)

Pre-World War II introduction. Trop. America. Common. Small tree, rarely over
8 m tall; leaves, about 10 to 16 cm long and 4 to 7 cm wide, bright green, alternate,
entire, oblong or oblong-elliptic, acute or acuminate, glabrous, somewhat glossy above,
pungent; petioles short; flowers, 2 to 3 cm or more long, yellowish-green, solitary or in
pairs, with 3 thick, broadly deltoid or heart-shaped light yellow sepals and 6 cordate
outer petals, borne on the branches and trunk (cauliflorous); fruit, up to 25 cm or longer
and weighing 0.5 to 2 kg, fleshy, oblong or ovoid, irregularly heart- or kidney-shaped,
with green to yellowish-green skin, covered regularly spaces, short, slightly curved fleshy
spines, and white, juicy, somewhat acid, aromatic, cotton-like, edible pulp; seeds, about
2 cm long, numerous, black, embedded in pulp. Planted in home gardens by Nauruans
and others and spontaneous on coastal strip, in some areas of escarpment forest, and in
the Buada Lagoon area. Ripe fruit eaten raw, often with ice cream. Common recently
introduced fruit tree throughout the Pacific. 2, 3(58586), 5, 6, 7.

Annona reticulata L. “custard apple", "bullock’s heart"

Pre-World War II introduction. Trop. America. Rare. Small to medium tree,
rarely over 9 m tall; leaves, up to 20 cm long and 6 cm wide or more, light green,
smooth, alternate, elliptic- or oblong-lanceolate, acute or acuminate, pinnately nerved,
brittle; petioles about 12 mm long; flowers, about 2.5 cm long, greenish, axillary, in
groups of 2 or 3 on lateral peduncles; fruit, 7 to 13 cm in diameter, yellowish- to
greenish-red, ovoid or heart-shaped, with whitish pulp, surface divided by impressed
lines into rhomboidal or hexagonal sections; seeds, numerous, glossy brown, embedded
in pulp. Planted fruit tree; found growing, possibly spontaneously, behind settlement on
Military Ridge. 2, 5(146), 6, 7.

89

Annona squamosa L. "sweetsop", "Sugar apple"
dawatsip (N); nameana (T)

Pre-World War II introduction. Trop. America. Occasional. Small tree, rarely
over 8 m tall; leaves, 7 to 14 cm long and 4 to 5 cm wide, pale bluish-green (glaucous),
two-ranked, thin, oblong or elliptic-lanceolate, acute, sparsely puberulent on both
surfaces when young; petioles, about 1 cm long; flowers, about 2.5 cm long, greenish-
yellow, axillary, solitary, pendent, petals narrowly lanceolate, obtusish, concave at base;
fruit, about, 8 cm in diameter, light grey-blue-green, subglobose, somewhat heart-shaped,
surface divided into prominent protruding knobs or tubercles, which separate, sweet,
edible cream-colored soft pulp; seeds, numerous, blackish, embedded in pulp. Fruit tree
planted in home gardens and growing spontaneously in inland coastal and lower es-
carpment forest. Ripe fruit eaten raw. 3(58589), 5(37), 6, 7.

" "

Cananga odorata (Lam.) Hook.f. & Thoms. "ylang-ylang", "perfume tree"
derangerang, derangirang (N)
syns. Canangium odoratum (Lam. ) Baill. ex King; Uvaria odorata Lam.

Recent introduction. Indomalaysia. Uncommon. Tree, up to 15 m or taller, with a
crooked trunk, smooth grey bark and drooping, brittle branches; leaves, 7 to 20 cm long
and 4 to 9 cm wide, dark green, alternate, simple, entire, elliptic-oblong, acute, base
rounded, obtuse, slightly pubescent, pinnately nerved, midrib prominent; petiole, mostly
less than 2 cm; flowers, yellowish-green turning yellow, drooping, very fragrant, sepals
3, petals 6, up to 6 or 8 cm long, wavy, linear-lanceolate, on long puberulent pedicels, in
axillary hanging (pedunculate) umbellate clusters of 4 to 12 flowers; fruit, 1.5 to 2.5 cm
long, greenish-black, oblong, olive-like, fleshy, borne in bunches from a stalk which
lengthens as the fruit develops; seeds, 6 to 12. Planted ornamental tree in home gardens
in Buada Lagoon area. Flowers used in garlands and for scenting coconut oil. An
important aboriginal introduction of considerable cultural importance throughout
Melanesia and Polynesia, where the flowers are used in garlands and to scent coconut oil.
Used in the commercial production of essential oil in the Philippines and Indonesia.
d(L7)31 6378

APIACEAE OR UMBELLIFERAE (Parsley Family)

Apium petroselinum L. "parsley"
syns. Petroselinum petroselinum (L.) Karst.; P. crispum (Mill.) Mansf.

Recent introduction. S. Europe and W. temperate Asia. Rare. Erect biennial or
short-lived perennial herb, up to 25 to 40 cm tall, with a stout tap root; leaves, aromatic,
shiny, dark green, in dense, tufted rosettes, 2 to 3-pinnate, with leaflets, 1 to 2 cm long,

90

deltoid-ovate, but deeply toothed or lobed, often much crisped; flowers, 2 mm in
diameter, greenish-yellow, umbellets, 10 to 20 flowered, borne in flat-topped compound
umbels, 2 to 5 cm in diameter; fruit (carpel), 2 to 3 mm long, grey-brown, ovoid,
laterally compressed, with 5 slender ridges. Pot herb in European home gardens; planted
in a halved 50-gallon drum on Command Ridge. Leaves and stems used by European
inhabitants as a spice in cooking. 5.

Coriandrum sativum L. "coriander", "cilantro", "Chinese parsley"
dhania (Hindi), uen sai (C)

Recent introduction. S. Europe and the Mediterranean region. Occasional. Erect
annual herb, 15 to 30 cm high; lower leaves, bright green, broad with crenately-lobed
margins, upper leaves finely cut with narrow linear lobes; flowers, white to pinkish, in
terminal umbels; fruit, about 3 mm in diameter, globose, yellow-brown, ribbed, 2-
seeded, with an unpleasant smell when unripe, but later becoming pleasantly aromatic.
Pot herb grown in Chinese home food gardens at Location and at Topside workshops.
Aromatic leaves used as a spice by Chinese and Indians. Dried, imported seeds an
important spice in curries. 5, 6, 7.

APOCYNACEAE (Dog-bane Family)

Adenium coetanum Stapf "desert rose", " mock azalea"
syn. A. obesum Balf.

Recent introduction. E. Africa. Rare. A smooth succulent shrub, 40 to 150 cm
high, with a thick trunk, short branches and milky sap which is said to be poisonous;
leaves, 2.5 to 10 cm long, dark green, shiny, ovate, arranged spirally in tufts at branch
tips; flowers, about 5 cm long and 3.8 cm across, red-pink, short-stemmed, funnel-
shaped, developing in clusters, 2 to 10 together. Pot plant in home gardens. 3(58716), 5,
6(176), 7.

Allamanda hendersonii Bull "allamanda", "cup of gold"
syn. A. cathartica L. var. hendersonii (Bull) Bailey & Raff.

Recent introduction. Brazil. Rare. Robust woody vinelike climbing shrub, with
milky poisonous sap; leaves, 8 to 15 cm long and mostly 4 to 5 cm wide, bright green,
shiny, opposite in pairs or in whorls of 3 or 4, elliptical to sub-obovate, but broadest
above the middle, abruptly acuminate, entire, thick and somewhat leathery (coriaceous),
midrib prominent; petiole very short; flowers, corolla tube 4 to 5 cm long, showy bright
yellow, slightly fragrant, trumpet-shaped, 5-parted with broad, spreading rounded lobes,

Of

totalling 6 to 7 cm or wider, borne in axillary and terminal racemose clusters of about 10
flowers; fruit, a prickly capsule. Planted ornamental. 3(58697), 5, 6.

Allamanda violacea Gardn. & Field "purple allamanda"

Recent introduction. Brazil. Rare. Woody vinelike climbing shrub; leaves, similar
to A. hendersonii, but bearing short, stiff hairs; flowers, similar to A. cathartica, but red-
purple and 5 to 6.5 cm in diameter. Planted ornamental. 3, 5, 6.

Catharanthus roseus ( L.) G. Don "periwinkle", "Madagascar periwinkle"
denea (N); te buraroti (K); losa (T)
syns. Vinca rosea L.; Lochnera rosea (L.) Reichenb.

Pre-World War II introduction. Madagascar. Common. An everblooming erect
perennial herb, up to 50 cm or taller, with white milky sap; leaves, 2.5 to 8 cm long and
2 to 3 cm wide, opposite, oblong or oblong-obovate, apex rounded, tipped with a sharp
point, base acute, finely puberulent on both surfaces; midrib pale; flowers, showy white
or pink, with or without a red throat, 5-parted, corolla tube, 2.5 to 3 cm long, lobes, 1.5
to 2 cm long, calyx, about 6 mm long, with awl-shaped lobes; fruit, up to 3 cm long,
with paired, 2.5 cm long puberulent podlike cylinders containing several cylindrical
seeds. Planted ornamental. Flowers used in garlands and reportedly boiled by some
people and drunk as a cure for diabetes. 3(58758), 5(127), 6, 7.

Cerbera manghas L. "cerbera", "sea mango"
dereiongo, dereiyongo (N); te reiango (K?)
syns. C. odollam sensu auct. non Gaertn.; C. lactaria (G. Don) Ham.; Tan-
ghinia lactaria G. Don

Indigenous. Trop. Asia to the Pacific Is. Rare. Medium-sized tree, up to 7 m or
taller, with white sap; leaves, 15 to 30 cm long and 5 to 8 cm wide, dark green, shiny,
simple, entire, lanceolate or obovate-lanceolate, acuminate, base acute-decurrent, spirally
and closely arranged; petiole 2 to 3 cm long, narrowly winged by decurrent blade
margins; flower, 2.5 to 6 cm wide, tube about 2.5 cm long, 5-petaled, white with red
throat, fragrant, calyx with 5 narrow lobes, 1.3 cm long, borne in branching terminal
clusters about as long as the leaves; fruit, globose or ovoid, 5 to 7 cm long, smooth,
deep purple to black when ripe, single or paired; seed, a woody stone, poisonous. Found
near homes on the coastal strip and near church at Buada. Seems to be planted, or at
least protected in built-up areas. No reported use by Nauruans, but its poisonous fruit is
reportedly used medicinally and to poison fish in Samoa, Tonga and elsewhere in the
Pacific. 5(16), 6, 7.

o2

Nerium oleander L. var. oleander "oleander"
te orian (K)

Pre-World War II introduction. S. Europe to Iran. Occasional. Large erect shrub,
up to 4 m or higher, with arching stems rising from the ground, young growth finely
pubescent, young branches slightly 3-angled and all parts extremely poisonous; leaves, 10
to 20 cm long and 2 to 3.5 cm wide, dull grey-green, narrowly oblong-lanceolate, acute
at both ends, entire, flat, leathery (coriaceous), midrib prominent, lateral nerves
numerous, fine; flowers, 4 to 5 cm wide, showy, white, pink to deep red, slightly
scented, with five toothed appendages in the corolla throat, borne in terminal cymose
clusters; fruit, 10 to 20 cm long, seldom seen, sometimes developing from single
flowers. Planted ornamental. 2, 3(58783), 4, 6, 7.

Nerium oleander L. var. indicum (Mill.) Deg. & Deg. "oleander", "sweet scented
oleander" te orian (K)
syns. N. indicum Mill.; N. odorum Soland.

Post World War II introduction? Iran to Japan. Occasional. Large erect shrub, up
to 4 m or higher, with arching stems rising from the ground, young growth finely
pubescent, young branches slightly 3-angled and all parts extremely poisonous; leaves, 10
to 20 cm long and 1.5 to 3 cm wide, dull grey-green, narrowly oblong-lanceolate, acute
at both ends, entire, edges rolled back, leathery (coriaceous), midrib prominent, lateral
nerves numerous, fine; flowers, 4 to 5 cm wide, corolla usually double, showy, white,
pink to deep red, fragrant, funnel-shaped, with fringed appendages in the corolla throat,
borne in terminal cymose clusters; fruit, 10 to 20 cm long, paired, pod-like, seldom seen,
sometimes developing from single flowers. Planted ornamental shrub. 3, 5, 6, 7.

Ochrosia elliptica Labill.
eorara, eoerara (N)
syn. Bleekeria elliptica (Labill.) Koidz.

Indigenous. Australia to the Pacific Is. Occasional to common. Small to medium-
sized tree, up to 8 m or higher, with white milky sap; leaves up to 15 cm long, oval or
inverted ovate, blunt or short-pointed, leathery (coriaceous), arranged in pairs, threes or
fours, with blades tapering into petioles about 1.2 cm long; flowers, small, cream-
colored, fragrant, stemless, developing in short cymose clusters at or near the branch
tips; fruit, 2.5 to 4.5 cm long, bright red, ovoid drupe, slightly keeled or flattened on the
margins, pointed, twinned, with mealy violet, scented pulp surrounding one or two seeds.
Tree in forest remnants on rocky outcrops on the central plateau, in escarpment forests
on southern half of the island, and occasionally in home gardens on coastal strip. Wood
used by Nauruans for rafters and small timber; leaves used medicinally, being crushed
with coconut cream to treat rashes, especially for children; fruit used in children’s
games, and fruit and flowers used in garlands. 2, 3(58802), 4(168N), 5(92), 6, 7(27812).

93

Plumeria obtusa L. "white frangipani", "plumeria"
demeria (N); te meria (K); melia, pua Solomona, melia Solomona (T)

Recent introduction. Trop. America. Common. Small to medium-sized, soft-
wooded, broad-crowned, knobby, branching evergreen tree, up to 5 m or taller, with
thick fragile branch tips and white milky sap; easily planted from cuttings; leaves, 20 to
35 cm long and 6 to 10 cm wide, dark green, glossy, alternate, obovate rounded, blunt,
smooth (glabrous), tapered at the base, pinnately nerved, midrib prominent, lateral nerves
loop-connected near margin, clustered near the ends of branches; petioles, 4 to 9 cm,
stout; flowers, 4 to 6 cm in diameter, all white with a central yellow eye, fragrant, but
less so than P. rubra, 5-parted, narrowly tubular, petal lobes obovate, longer than tube,
borne in terminal cymose clusters; fruit, 15 to 24 cm long, dark green later black, single
or twinned, cylindrical, fusiform, with many flat winged overlapping seeds. Flowers used
in garlands and dried in sun and used to scent coconut oil (eir). 3(58775), 5(111), 6, 7.

Plumeria rubra L. "frangipani", "plumeria", "temple tree", "graveyard tree”
demeria, arabaneit (N); te meria (K); melia, pua Fiti, melia Nauru (T)
syns. P. acuminata Ait.f. and P. acutifolia Poir.

Pre-World War II introduction. Trop. America. Common. Small to medium-sized,
soft-wooded, broad-crowned branching deciduous tree, up to 5 m or taller, with thick
fragile branch tips and white milky sap; easily planted from cuttings; leaves, 20 to 35 cm
long and 6 to 10 cm wide, pale green to grey-green, alternate, elliptic-oblong, acute or
acuminate, smooth (glabrous), tapered at the base, pinnately nerved, midrib prominent,
lateral nerves loop-connected near margin, clustered near the ends of branches; petioles,
4 to 9 cm, stout; flowers, 3 to 5 cm in diameter, waxy-white to yellow, pink, deep red
or multicolored variations, very fragrant, 5-parted, narrowly tubular, petal lobes obovate,
longer than tube, borne in terminal cymose clusters; fruit, 15 to 20 cm long, dark green
later black, single or twinned, cylindrical, fusiform, with many flat winged overlapping
seeds. Planted ornamental. The name demeria seems to be applied to all color forms and
to both species of Plumeria, with the older name arabaneit being reserved for the bright
yellow to yellow-white cultivars, which are longer established in Nauru. Leaves used
medicinally and mixed with coconut oil for curing fever; flowers used in garlands and
dried in the sun and used to scent coconut oil (eir). 2, 3(58654), 5(50), 6, 7.

Tabernaemontana divaricata (L.) R. Br. "false gardenia", "paper gardenia",
"crepe jasmine", "scentless gardenia"
syns. WNerium divaricatum L.; N. coronarium Jacq.; Tabernaemontana coronaria
(Jacq.) Willd.; Ervatamia divaricata (L.) Burkill; E. coronaria (Jacq.)
Stapf; Tabernaemontana alternifolia L.; Nyctanthes acuminata Burm.f.

94

Recent introduction. India. Occasional. Smooth erect shrub, up to 2 or 3 m tall,
resembling a gardenia, with scanty, white milky sap; leaves, 6 to 15 cm long and 2 to
4.5 cm wide, dark green, lighter below, glossy, opposite, elliptic oblong, acute or
obtuse, acute at base, glabrous, thin, with 6 to 10 pairs of curving veins; petiole short,
mostly about 1 cm long, clasping the stem; flowers, 2.5 to 5 cm in diameter, usually
several, up to about 8, together or rarely solitary, in flat-topped corymbose clusters near
branch tips or leaf axils, white, nearly odorless, calyx about 5 mm long, segments rather
broad, deltoid and obtuse, corolla, 5-parted, tube 1.5 to 2.5 cm long, cylindrical, lobes,
about 2 cm long, often doubled and ruffled, crinkled or crepe-like; fruit, rarely seen in
cultivation, 3 to 7 cm long, yellow-orange, paired, oblong, podlike, with 1 to 3 ridges,
pubescent, red within; seeds, many, embedded in pulp. Planted ornamental. 3(58706), 5,
6222) a7:

Thevetia peruviana (Pers.) K. Schum. "be-still tree", "yellow oleander"
syns. Cerbera peruviana Pers.; C. thevetia L.; Thevetia neriifolia Juss. ex
Steud.; 7. thevetia (L.) Millsp.

Recent introduction. Peru. Occasional. Everblooming shrub or small tree, up to 8
m or taller, propagated from seeds and cuttings; all parts reportedly poisonous; leaves, 5
to 15 cm long and 0.5 to less than 1 cm wide, numerous, dark green, somewhat shiny
above, paler beneath, leathery (coriaceous), alternate, linear-lanceolate, acute or
subobtuse, acute at base; petiole, very short, about 3 mm long; flowers, about 4 to 7 cm
long, 5-parted to more than halfway down, showy yellow, mildly fragrant, funnel- or
trumpet-shaped, borne in terminal cymose clusters; calyx, 1 to 1.3 cm long, 5-lobed,
acute-acuminate; fruit, 4 to 5.5 cm broad, a juicy red drupe, ripening to black, con-
taining a thin layer of pulp and 2 oily seeds. Planted ornamental. 3(58704), 5(78), 6, 7.

AQUIFOLIACEAE (Holly Family)

Ilex sp. "holly"

Recent introduction. Eurasia. Rare. Evergreen shrub with short-stemmed leaves
bearing several strong spines. Planted ornamental. 6(164).

95
ARALIACEAE (Panax Family)

Polyscias balfouriana (Andre) Bailey "panax"

Recent introduction. Melanesia. Rare. Large erect shrub, up to 3 m or taller, with
few, rapidly ascending branches; leaves, 20 to 50 cm long, compound, 1-pinnate;
petioles, 8 to 22 cm long, petiolules, 2 to 4 cm long; leaflets, 5 to 20 cm long and 5 to
18 cm wide, ovoid to orbicular, cordate, shiny, dark-green, opposite, mostly 3 or 5 in
number, terminal leaf largest, crenate. Planted ornamental, commonly in hedges. 5, 6,
7(22324).

Polyscias cumingiana (Presl) Fern.-Vil. "panax"
lautagitagi (T)
syns. Paratropia cumingiana Presl; Nothopanax cumingii Seem.; Aralia filicifolia
C. Moore ex Fourn.; Polyscias filicifolia (Moore) Bailey

Recent introduction. Malesia, possible to Melanesia. Rare. Large erect shrub, up
to 3 m or taller, with few, rapidly ascending weak branches; leaves, mostly 15 to 70 cm
long, alternate, 1-pinnate, uppermost leaves turning a showy light-yellow, giving the
plant a feathery appearance; petioles, 5 to 20 cm long, petiolules, 1 to 2.5 cm long;
leaflets, 6 to 15 cm long and about 5 cm wide, opposite, entire to sharp-toothed or
narrowly lobed, 6 to 7 pairs, plus terminal leaflet (13 to 15 leaflets), anise-scented;
petioles 1 to 2 cm long. Planted ornamental, commonly in hedges. 6, 7.

Polyscias fruticosa (L.) Harms
te mamara (K?); lautagitagi (T)
syns. Panax fruitcosum L.; Nothopanax fruticosus (L.) Miq.

panax

Pre-World War II introduction. India to W. Polynesia. Occasional. Erect, rather
few-branched shrub, | to 3 m tall; leaves, up to 30 cm or longer, alternate, pinnate,
usually bi- or tri-pinnate; leaflets, 5 to 10 cm long, terminal leaf usually the largest,
green or variegated, commonly edged with white, glabrous, anise-scented, irregularly-
shaped, more or less lanceolate, usually toothed, lobed laciniate or pinnatifid; flowers,
small, greenish-white, borne in small umbels, forming large terminal paniculate clusters;
fruit, flattened, orbicular, ribbed 2-seeded. Planted ornamental, commonly in hedges or
as a living fence. 2, 3(58698), 5, 6(171), 7(22323).

Polyscias guilfoylei (Cogn. & March.) Bailey "panax", "hedge panax"
te toara (K); lautagitagi (T)
syns. Aralia guilfoylei Cogn. & March.; Nothopanax guilfoylei (Cogn. &
March.) Merr.

96

Pre-World war II introduction. Melanesia to S. Polynesia. Common. Large shrub,
2 to 6 m tall, with weak, rapidly ascending vertical branches and light grey bark; leaves,
mostly 15 to 50 cm long, alternate, variable, but commonly broadly ovate or elliptic and
coarsely sharply-toothed or laciniate, rarely subentire, commonly variegated with white
or pale yellow margins, or sometimes all dark green, shiny; petioles, 2 to 3 cm long,
clasping the stem; flowers, rarely seen, small, 5-parted, petals, 2.5 by 1.2 mm, in
umbels, borne on a large, much-branched terminal panicle; fruit, usually 3- to 4-celled,
with 3 to 4 seeds, 4 by 5 mm. Planted ornamental, commonly as a hedge or living fence.
2, 3(58696), 5(20), 6, 7(27822).

Polyscias scutellaria (Burm.f.) Fosb. "panax
te toara (K); lautagitagi (T)
syns. Crassula scutellaria Burm.f.; Polyscias pinnata J.R. & G. Forst.; Notho-
panax scutellaria (Burm.f.) Merr.

Recent introduction. S.E. Asia. Occasional. Medium-sized erect shrub, 1 to 3 m
tall, with weak, ascending branches; leaves, simple (unifoliate) or pinnate, with 3 or
sometimes 5 leaflets; petioles, at least 12 cm, petiolules, 2 to 5 cm or longer; leaflets, 5
to 14 cm in diameter, the terminal leaflet of compound leaves largest, orbicular-cordate,
concave and saucer-like, glossy, base subcordate-cordate, sometimes obliquely so, green
or with white or yellow variegations, some forms wavy or with blunt or rounded teeth
(crenate); flowers, borne in umbels on a much-branched panicle, small, about 3 mm
long, petals, about 2.5 mm long, 6 or 7 in number; fruit, flattened, 2-seeded (rarely 3-
celled and 3-seeded). Planted ornamental, commonly as a hedge or living fence.
368693); 5, 6, 7:

Polyscias tricochleata (Miq.) Fosb. "panax"
syn. PP. pinnata Fosb. cv. tricochleata Stone; Nothopanax tricochleatus Miq.

Recent introduction. Pacific Islands. Rare. Erect shrub, 1 to 2 m tall; leaves
usually trifoliate (rarely 1 or 5 leaflets); leaflets, orbicular, white- or yellow-margined;
flowers and fruit unknown. Known to be a mutant form of P. pinnata. Planted erect
ornamental shrub with white-margined leaflets. 3(58674), 7(22325).

Schefflera actinophylla (Endl.) Harms "Queensland umbrella tree", "octopus tree"
syn. Brassaia actinophylla Endl.

Recent introduction. N. Australia. Occasional. Small to medium-sized, soft-
wooded tree, up to 10 m or taller, with thick branches marked by conspicuous leaf-scars,
sometimes epiphytic; leaves, large, up to 90 cm or wider in diameter, palmately-
compound, umbrella-shaped, with 7 to 15, usually 7 to 9, leaflets, forming rosettes at the

oF,

branch ends; main petioles 15 to 45 cm long, petiolules, 2.5 to 8 cm long; leaflets, 10 to
30 cm long, elliptic-obovate, oblong, sub-acuminate, the central ones longest, shiny,
dark-green, leathery (coriaceous); flowers, in small red heads, arranged more or less
spicate-racemosely along stout wide-spreading axes, up to 60 cm or longer, several of
which radiate, like the tentacles of an octopus, from a central hub-like axis arising from
each leaf cluster; fruit, 10- to 12-seeded, nutlets, red-purple. Planted ornamental.
3(58672), 5, 6, 7.

ASCLEPIADACEAE (Milkweed Family)

Asclepias curassavica L."milkweed", "butterfly weed", "red cotton bush", "bloodflower"
dupaimdupaim, dupaimdupwaim (N)

Pre-World War II introduction? Trop. America. Rare. Slender erect, branched,
woody perennial herb, up to 1 m or higher, with milky sap; leaves, 6 to 15 cm long and
1 to 3 cm wide, opposite, oblong-lanceolate, acute at both ends, pinnately-nerved,
pubescent on nerves beneath; flowers, many, small, about 60 mm in diameter, borne in
4- to 15-flowered terminal and axillary umbels on finely pubescent peduncles, 3.5 to 6
cm long; corolla, 7 to 9 mm long, red to reddish-purple, with 5 deeply-parted lobes, bent
backward (reflexed), and a protruding orange to yellowish-orange scaly crown, 3.5 to 4
mm long; fruit, 5 to 7.5 cm long, erect, pointed, ovoid, podlike, smooth or downy,
containing numerous round, flat seeds, 6 to 7 mm long, each bearing a tuft of long,
silky, white hairs, 2 to 2.7 mm long. Planted ornamental; flowers used for body
ornamentation and making garlands; reportedly poisonous to livestock. 5, 6.

Calotropis gigantea (L.) R. Br. "crown flower", "giant milkweed"
te bumorimori (K)
syn. Asclepias gigantea L.

Recent introduction. India to Indonesia. Rare. A large shrub, up to 4 m or higher,
with thick downy branches and copious milky sap; leaves, 7.5 to 25 cm long, nearly
stemless (shortly petiolate), broad ovate-oblong, thick, pale green, downy, woolly
beneath, indented at base; flowers, about 3 to 4 cm in diameter, pale lavender or white,
sweetish fragrance that deteriorates when crushed, with 5 curled-back petals and a
prominent symmetrical crown, sometimes only consisting of the flower crowns. Planted
ornamental at the Meneng Hotel. 3(58771), 5, 6(152), 7.

98

Hoya carnosa (L.) R. Br. “wax plant", "wax flower"
syns. Asclepias carnosa L. ;

Recent introduction. S$. China. Rare. Fleshy waxy climbing vine, climbing by
roots, up to 2 m or longer; leaves, 5 to 10 cm long, opposite, narrow to broad ovate-
oblong, mid-green, thick, fleshy, shiny, flat; flowers, 1 to 1.3 cm in diameter, 5-parted,
white with pink centers, convex, wheel- or star-shaped, waxy, fragrant. Planted ornamen-
tal. Single specimen; could have been the Australian species, H. australis R. Br. (syn. H.
bicarinata Gray). 6, 7.

ASTERACEAE OR COMPOSITAE (Aster, Sunflower or Composite Family)

Ageratum conyzoides L. "goat weed", "ageratum"
bwiyat tsige, bwiyat ziege (N)

Pre-World War I introduction. Trop. America. Occasional. Erect to somewhat
sprawling, branching, weak-stemmed, strong smelling, annual herb, up to 80 cm high;
leaves, 2 to 10 cm long and | to 5 cm wide, opposite, ovate or rhombic-ovate, acutish,
base rounded and acute at petiole, edges scalloped, pubescent on both surfaces, glandular
dorsally; petiole up to 5 cm or longer; flowerheads, small, 4 to 6 mm long, bluish,
corymbose, in loose, terminal and axillary clusters; involucral bracts, 3 mm long, acute-
acuminate, subglabrous; florets, about 75 per head, about 1 mm long, white or light
purplish-blue; fruit (achenes), 1.5 to 2 mm long, numerous, angled, nearly glabrous;
pappus, of 5 awned scales. Found on low ground near Buada Lagoon and occasionally in
other ruderal habitats. Leaves and flowers used in garlands and body decoration and for
scenting coconut oil. 3(58652), 5(11), 6, 7, 8(9574).

Bidens alba (L.) DC. “cobbler’s peg", "Spanish needle" kauen oe, kawen oe (N)
syns. Coreopsis alba L.; C. leucanthema L.; C. leucantha L.; B. pilosa L. var.
radiata sensu auct. non Sch.-Bip.; B. leucantha (L.) Willd.

Recent introduction. Trop. America. Rare. Erect branching annual herb (in big
plants, the branches sometimes straggling), up to 1.5 m tall; leaf blades, 1 to 12 cm long
and up to 6 cm wide, lower leaves simple, ovate and serrate, but upper leaves trifoliate
or imparipinnate, leaflets ovate or ovate-oblong, acute, basally decurrent, petiolulate,
serrate; petioles, up to 6 cm long; flowerheads, few, in panicles; involucral bracts, about
7 or 8, linear-spatulate; heads, 6 to 8 mm long and 6 to 8 mm wide; ray florets bright
white, 6 to 8 mm long; disc-florets yellow; fruit (achenes), linear, 7 to 13 mm long,
black or dark brown, about 1 mm wide, flat, 4-angled, short-strigose or glabrous, with 2
to 4 barbed awns, about 3 mm long, at the tip. Weed on dirt pile near Topside sports
oval; not seen in 1987. 4(142N), 5(76), 6.

99

wo

Bidens pilosa L. "cobbler’s peg"’ Spanish needle"

Recent introduction. Trop. America. Common? Slender, erect, branching annual
herb, 20 to 90 cm tall; stems, 4-angled, glabrous; leaves, mostly 1 to 5 cm long, simple
to compound or deeply 3 to 5-lobed, opposite; lobes or leaflets ovate to lanceolate, acute,
decurrent at the base, serrate, mostly glabrous; flowerheads, 5 to 10 mm long, on
terminal or upper axillary panicles or solitary on long stalks (peduncles), up to 3 cm
long; involucral bracts, 2 to 3-ranked, lanceolate, the outer ones somewhat leaflike; ray
florets usually absent; disc-florets, yellow, tubular; fruit, 6 to 12 cm long, a black,
ribbed, straight or curved, linear achene, tipped with 2 to 4 barbed awns, which adhere
to clothing. Weed in wasteplaces, ruderal habitats and gardens. 8.

Conyza bonariensis (L.) Cronq. "hairy horseweed"
syns. Erigeron bonariensis L.; E. crispus Pourr.; E. albidus (Willd. ex Spr.) A.
Gray; Conyza albida Willd. ex Spr.

Recent introduction. Pantropical. Rare. Slender, erect annual herb, 30 cm to 1 m
or higher, with pubescent stem, the lower portion with leaf scars; leaves, 2.5 to 12 cm
long, dark greyish-green, narrowly lanceolate, lower ones, oblanceolate, coarsely dentate,
upper ones, linear, entire, pubescent on both surfaces, attached to stem by a broad base;
flower heads, 1 to 1.3 cm in diameter, in racemes or racemose panicles; involucral
bracts, about 5 mm long, grey-pubescent (downy); ray-florets, usually many, in-
conspicuous, white to cream; fruit, about 1.6 mm long, narrow, oblong, pale achene
tipped with a pappus of light silky hairs, 3 to 5 mm long. Weed in waste places.
4(133N), 5.

Dahlia pinnata Cav. "dahlia"

Recent introduction. Mexico. Rare. Perennial herb, up to 1 m or higher, with
tuberous roots; leaves, opposite, simple or divided feather-fashion one to three times;
leaflets, toothed or lobed, arranged in pairs with an extra leaflet at the tip; flowerheads,
2.5 to 10 cm in diameter, long-stemmed, single or double, flat to globose; ray-florets,
yellow central florets surrounded by conspicuous flat, tubular or rolled outer ray-florets,
ranging in color from white and yellow to pink, purple and red, which range from eight
to many and often conceal the central ray-florets; fruit, tiny, narrow, flattened and tipped
with 2 teeth or untipped. Planted ornamental. 5(79), 6.

100

Emilia sonchifolia (L.) DC. "purple sow thistle", "floras paintbrush"
syn. Cacalia sonchifolia L.

Recent introduction. Pantropical. Occasional. Erect, soft-stemmed, branching
annual herb, 10 to 40 cm tall, lower stems pubescent, upper ones nearly glabrous; leaves,
3 to 15 cm long and 5 to 7 cm wide, simple, alternate; lower leaves, deeply irregularly
lobed or dentate, subdeltoid to spoonshaped, clasping stem at the base; upper leaves,
smaller, lanceolate, toothed to subentire, sometimes purplish beneath on midrib and
bluish-green (glaucous) on blade; flower heads, 0.8 to 1 cm long and 2.5 to 6.5 mm
wide, few, cylindrical, on slender lax peduncles in loose, branching terminal clusters;
involucral bracts, about 8, cylindrical, fused, which split apart and reflex at maturity,
almost covering the flowers; ray-florets absent; disc-florets, 30 to 60 in number, 5-lobed,
white below, lavender to pink at the tips, tubular, just slightly longer than the involucre;
fruit, about 2.5 to 3 mm long, brown, prism-shaped, 5-ribbed achene, with a pappus of
numerous, silky white bristles, 6 to 8 mm long. Weed in waste places and as a pioneer in
recently mined areas. 5(115), 6, 7(22313), 8.

Gerbera jamesonii Bolus "Transvaal daisy", "gerbera", "African Daisy"

Recent introduction. S. Africa. Rare. Hairy perennial herb, 30 to 35 cm high;
leaves, 15 to 30 cm long (blade about 2/3 as long as stem)), many, deeply lobed, feather-
fashion, woolly beneath, arranged in a basal rosette; flowerheads, 7.5 to 10 cm in
diameter, attractive, daisy-like, pink, red, orange, yellow or cream-colored, solitary,
borne on the crest bare flower stalks, about 25 to 40 cm long; central disc-florets, many,
surrounded by | or 2 rows of narrow orange ray-florets; many single and double hybrid
forms exist. Planted ornamental. 5, 6, 7.

Gynura aurantiaca (Bl.) DC. "purple passion flower", "velvet plant"

Recent introduction. Java. Rare. Erect or semi-climbing, up to 75 cm; ovate,
deeply lobed or toothed, leaves, dark green, covered with thick, bright purple hairs,
giving the plant a velvety appearance; flowerheads, about 2.5 cm in diameter, like a
petal-less daisy; disc-florets orange. Ornamental pot plant. 6.

Synedrella nodiflora (L.) Gaertn. "synedrella", "nodeweed"
syn. Verbesina nodiflora L.

Recent introduction. Trop. America. Occasional. Coarse, erect to ascending,
branched annual herb, 10 to 75 cm high, often rooting at lower, decumbent nodes;
leaves, 1.5 to 8 cm long and 1.5 to 5 cm wide, larger leaves usually at the top, simple,
opposite, ovate to elliptic, acute tip and cuneate base, subentire to crenate-serrate,
scabrous, 3-nerved from base,; petiole, about 1 to 1.5 cm long, winged, pubescent on

101

both surfaces; flowerheads, 0.6 to 1.2 cm long and 5 to 6 mm across, 10 to 20-flowered,
subsessile, solitary or few together on short axillary or terminal peduncles; involucral
bracts, nearly 1 cm long, only 4 or 5, lanceolate, 2-ranked (2-seriate); ray-florets, about
3.5 to 4 mm long, yellow, strap-shaped, 3-lobed; disc-florets, yellow, with tubular
corollas; fruit, 4 to 5 mm long, a black or dark-brown achene, those of ray-florets flat,
oval and spiny-edged or winged with 2 terminal awns, those of the disc-florets cylin-
drical, unwinged with 2 erect spiny terminal awns about half as long as the achene. Weed
in low ground around Buada Lagoon and in other ruderal habitats. 3(58648), 4(134N),
5(97), 6, 7, 8(9573).

Tagetes erecta L. "marigold", "Aztec marigold", "African marigold"

Recent introduction. Mexico. Rare. Erect, smooth, few-branching, strongly-
scented, annual herb, up to 60 cm high; lower leaves opposite, upper leaves alternate,
pinnately parted or pinnate, up to 12 cm long, the segments more or less oblong, acute,
distally dentate, up to 9 cm long and 1.5 cm wide; flower heads, 4 to 8 cm in diameter,
solitary, flat to globose, borne on peduncles which flare distally; involucral bracts, about
2 cm long, arranged cylindrically; ray-florets yellow to red-orange, many, often
concealing the central disc-florets; fruit, achene tipped with pappus, about 1 cm long, and
several short scales, 3 to 4 mm long. Planted ornamental. 5, 6.

Tridax procumbens L. "wild daisy", "coat buttons"

Pre-World War II introduction. Trop. America. Common. Pubescent perennial
herb with prostrate to ascending stems, up to about 80 cm long; leaves, 1 to 6 cm long
and 1 to 3.5 cm wide, opposite, broadly lanceolate, coarsely toothed, acute or subacute,
cuneate at base, hispid on both surfaces; petioles, 5 to 15 mm long; flowerheads, about 1
cm long and 2 cm wide, terminal, borne on erect pilose peduncles, 10 to 30 cm long;
involucral bracts, 2- or 3-ranked, puberulent; ray-florets, about 4 to 5 mm long, 5 or 6 in
number, with a narrow corolla tube and broad ligulate limb, about 3 by 5 mm, white to
light yellow; disc-florets, many, the corolla narrow-campanulate, about 8 mm long,
bright yellow, 5-lobed and hairy at top; fruit, about 2 mm long, dark grey-brown,
densely hairy achene, with a spreading pappus, 5 to 6 mm long, of plumose hairs. Weed
in settled areas and near airport; pioneer in recently mined areas. 2, 3(58657), 4(145N),
5(25), 6, 7, 8(9548).

Vernonia cinerea (L.) Less. "iron weed"
syns. V. parviflora Reinw. ex Bl.; V. cinerea var. parviflora (Reinw. ex Bl.);
Conyza cinerea L.

Pre-World War II introduction. Trop. Asia. Common. Erect loosely branching
annual herb, 15 to 60 cm tall, stems longitudinally ribbed; leaves, 1 to 7 cm long and 1

102

to 2.5 cm wide, simple, alternate, lanceolate to ovate, acute, or variably shaped, the
upper ones narrower, smaller and subsessile, the lower ones with surfaces minutely
puberulent, base decurrent, irregularly toothed, with winged petioles, 1 to 3 cm long;
flowerheads, 6 to 7 mm long, in loose terminal paniculate clusters on slender peduncles;
involucral bracts, 4 to 5 mm long, green with brownish tips, lanceolate, 4-ranked (4-
seriate), puberulent; ray-florets, absent; disc-florets, about 4 mm long, 20 to 25 in
number, tubular, violet or pinkish violet, exerted; fruit, 1.5 to 2 mm long, a grey-brown,
cylindrical, appressed pubescent achene, 1.5 to 2 mm long, bearing a terminal pappus of
numerous white bristles which spread apart at maturity. Weed in settled areas; pioneer in
recently mined areas. 2, 3(58611), 4(118N), 5(67), 6, 7, 8(9538).

Wedelia trilobata (L.) Hitchc. "wedelia"
syn. Silphium trilobatum L.

Recent introduction. Trop. America. Occasional. Perennial creeping herb,
reaching a thickness (height) of about 40 to 70 cm; leaves, 2.5 to 7 cm long, bright
green, simple, opposite, three-lobed to lanceolate, toothed; petioles, short, extremely-
winged, or lacking, clasping the stem; flowerheads, about 2 to 2.6 cm in diameter, daisy-
like, solitary, on slender peduncles, about 5 to 7 cm long; involucral bracts, 6 to 9 mm
long, green, lanceolate, 2-ranked, 5 short inner and 5 longer outer bracts; ray-florets,
about 1 cm or longer and 0.5 cm wide, 8 to 10 in number, bright yellow; disc-florets,
about 8 mm long, light yellow, tubular, 20 to 30 in number, fruit, not seen. Planted
ornamental groundcover. 3(58609), 5, 6(237), 7.

Zinnia elegans Jacq. "Zinnia"
£ q

Recent introduction. Mexico. Erect annual herb, up to 60 cm tall, stems branched;
leaves, 2.5 to 14 cm long and 1.5 to 6 cm wide, opposite, ovate to elliptic or oblong,
obtuse or acute, hispid, sessile; flowerheads, 3 to 8 cm in diameter, solitary, showy,
developing at stem and branch tips; involucral bracts, about 1 cm long, chaffy, in about
3-ranks; ray-florets, about 2 cm long and 1 cm wide, 14 to 20 in number, pistillate,
violet, pink, orange, yellow or white; receptacle, convex, paleaceous, bearing numerous
5-lobed central disc-florets, about 5 mm long, yellow; fruit, 3-angled, compressed
achene, with | to 3 terminal awns. Planted ornamental. 5(128), 6.

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BALSAMINACEAE (Balsam Family)

Impatiens balsamina L. "balsam", "garden balsam"

Recent introduction. India or Africa. Rare. Succulent, watery, erect, branching
annual herb, up to about 60 cm high; leaves, up to 5 cm long and 4 cm wide, simple,
alternate, lanceolate, acuminate, toothed (serrate); petiole, usually glandular at the base;
flowers, up to 2.5 cm or more in diameter, short-stemmed, borne on short axillary
pedicels on the stem below the leafy tip; sepals, 3 or 5, one usually spurred; petals, 3-
lobed, lateral ones bifid, commonly double or long-spurred, showy yellow, white, pink,
purple or red flowers; fruit, a woolly, 5-valved capsule, explosively dehiscent, the valves
incurling instantly when separated; seeds, subglobose, usually brown, finely pitted.
Planted ornamental. 5(117), 6.

Impatiens walleriana Hook.f. "snapweed", "patience plant", "Zanzibar balsam"
syn. I. sultanii_ Hook.f.

Recent introduction. Zanzibar. Rare. Erect, branching, sub-shrubby perennial
succulent herb, up to 50 cm high; leaves, alternate along the stem and arranged in a
rosette at branch tips, narrow-ovate, acuminate at tip and base, toothed or scalloped, with
soft point at each indentation; flowers, 2.5 to 3.5 cm in diameter, petals in one plane,
borne, singly or 2 to 3 together, on long axillary pedicels; sepals, 3 or 5, 1 usually
spurred; petals, showy, bright scarlet or red to pink, accompanied by a long spur; fruit.
which may or may not develop in cultivation, a 5-valved capsule. Planted ornamental. 6.

BASELLACEAE (Basella Family)

Basella rubra L. "Indian spinach", "Ceylon spinach", "Malabar nightshade"
shaan ts’oi (C)
syn. B. alba L.

Recent introduction. Trop. Asia. Occasional. Succulent dark-green to reddish-
purple-stemmed, glabrous, branching, herbaceous, perennial twining vine, up to 4 m or
longer; leaves, 5 to 15 cm long and 4 to 14 cm wide, alternate, entire, broad-ovate,
almost heart-shaped, tender, fleshy; petioles, 1 to 2.5 cm long, green or purplish;
flowers, 4 to 5 mm in diameter, sessile, closed, white to light reddish-purple, clustered
on axillary spikes, 2 to 20 cm long; fruit, about 8 mm in diameter, ovoid, black, berry-
like. Food plant in Chinese gardens and containers at Location. Leaves and tender stems
cooked as a spinach. 5(138), 6.

104

BEGONIACEAE (Begonia Family)

Begonia coccinea Hook.f. “angel-wing begonia"

Recent introduction. Brazil. Occasional. Smooth shrub-like herb, up to 1 m or
higher, with bamboo-like stems; leaves, 10 cm or longer, oblong, pointed, glossy green
above, reddish below, finely edged with red, with a deep indentation at the base and
wavy edges; flowers, about 2.5 cm in diameter, coral-red, borne in branched red-
stemmed drooping clusters; male flowers, about 2.5 cm in diameter, 4- petaled, with two
petals larger than the other two; female flowers, showy, with red, 3-winged ovary, about
2.5 cm long. Ornamental pot plant. 5, 6.

Begonia rex Putz x B. spp. "hybrid begonia"

Recent introduction. Trop. America. Occasional. Erect perennial herb, up to 30
cm or higher, with short, thick, underground stems; leaves, variable, more or less ovate,
pointing downward, angled or lobed, sometimes waxy, commonly with a silvery, pink,
red, copper or bronze patterning on a background of all shades of green zone, and
reddish beneath; flowers, variable small, pale pink, in small clusters rising from the
leaves. Ornamental pot plant. 7.

Begonia spp. "begonia" cultivars

Recent introduction. Trop. America. Occasional. Pot plants and planted ornamen-
tals. 5(262), 6, 7.

BIGNONIACEAE (Bignonia Family)

Jacaranda mimosifolia D. Don "jacaranda"
syns. J. acutifolia Humb. & Bonpl.; J. ovalifolia R. Br.

Recent introduction. Brazil. Rare. Tree, up to 15 m or higher; leaves, up to 50
cm or longer, opposite, compound, fernlike, the primary axis with 20 to 40 branchlets
(10 to 20 pairs of divisions), each which bears 14 to 24 pairs of oblong, hairy, pointed
leaflets, about 6.5 mm long; flowers, terminal and axillary, 40 to 90 in number, borne in
erect or drooping panicles, about 20 to 30 cm long; corolla, about 5 cm long, showy
light blue-violet, perfumeless, bell-shaped, 2-lipped, S-lobed; fruit, about 5 cm in

105

diameter, a round, flattened, wavy-edged capsule, which looks like a brown bivalve
mollusc. One immature seedling planted as an ornamental. 6, 7.

Spathodea campanulata Beauv. "African tulip tree", "flame of the forest",
"fountain tree"

Recent introduction. Trop. Africa. Occasional. Medium-sized tree, up to 20 m
tall, with soft, weak wood and a ridged or buttressed trunk; leaves, 30 to 50 cm long,
dark-green, shiny, opposite, odd-pinnate, divided feather-fashion; leaflets, 4 to 12 cm
long and 2.5 to 5.5 cm wide, 3- to 9-paired with a terminal leaflet, elliptic or ovate,
acute-acuminate, entire, deep-veined, pubescent dorsally on nerves; petioles, 1 to 3 mm
long; flowers, up to about 10 to 13 cm long, in showy terminal racemose clusters,
opening a few at a time; calyx, hairy, closed in bud, opening along one side, boat-
shaped, curving upward, nearly as long as corolla, containing water when closed;
corolla, about 10 cm long and 5 cm wide, with 5 oval lobes, vivid red-orange to
vermilion, the margins dark yellow, obliquely campanulate, gaping, segments crisped-
undulate, somewhat tulip-shaped; stamens with yellow filaments; fruit, 15 to 20 cm long
and 4 to 5 cm wide, black flattened canoe-shaped capsules; seeds, woody, with a wide
parchment-like circular wing. Planted ornamental tree. Flowers used in garlands and for
ornamentation. 3(58682), 5(123), 6, 7.

Tecoma stans (L.) Juss. ex HBK. "yellow elder", "yellow bells", "tecoma",
"ginger Thomas" yellow flower (N); nei karairai (Miss or Mrs. Karairai)(K);
neikarairai (T)
syns. Bignonia stans L.; Stenolobium stans (L.) D. Don

Recent introduction. Trop. America. Common. Erect shrub, up to 4 m tall;
leaves, 10 to 30 cm long, opposite, pinnate; leaflets, 3 to 10 cm long and 2 to 3 cm
wide, 5 to 13 in number, lanceolate to elliptic-oblong, serrate, acute-acuminate, base
cuneate, sessile or short-petiolate, pinnately nerved; flowers, in large terminal racemose
clusters; pedicels to 1 cm long; calyx, 3 to 5 mm long, tubular, campanulate, 5-toothed;
corolla, 3 to 5 mm long, bright yellow, slightly fragrant, funnel-form-campanulate or
bell-shaped, slightly bilabiate, with 5 wavy lobes; fruit, 10 to 20 cm long and 5 to 6 mm
wide, narrow, flattened, rostrate capsule; seeds, many, winged. Planted ornamental.
Showy flowers used in garlands and for ornamentation, especially by I-Kiribati. 3-
(58656), 5(193), 6, 7, 8(9570).

106
BOMBACACEAE (Bombax Family)

Ceiba pentandra (L.) Gaertn. "kapok tree", "silk-cotton tree" duwoduwo (N)
syns. Bombax pentandrum L.; Eriodendron anfractuosum DC.; Ceiba casearia
Medic.; Bombax orientale Spreng.

Pre-World War II introduction. India or Africa. Occasional. Tall deciduous, soft-
wooded, light-grey-barked tree, up to 25 m or higher, with a buttressed trunk, commonly
spiny below and cylindrical and smooth above, and horizontal tiers (whorls) of widely-
spaced spreading branches; leaves, palmately compound, with 5 to 9 leaflets; leaflets, 7
to 18 cm long and 1 to 3.5 cm wide, elliptic to oblanceolate, acuminate, blue-green
(glaucous) beneath, entire or obscurely toothed, distinctly petiolate; petioles, 7 to 20 cm
long; flowers, which appear just before the leaves, clustered on branches; corolla, 5-
petaled, cream-white or pale pink, pubescent; petals about 2.5 cm long; fruit, 7 to 15 cm
long, oblong-ellipsoid, capsular, smooth, pendulous, 5-celled, eventually dehiscent, filled
with numerous long, soft silky or cotton-like fibers, each 0.8 to 3 cm long; seeds, many,
brown. Planted; some possibly spontaneous near Buada Lagoon. Fiber used in the past
for stuffing pillows and mattresses. 3(58642), 5, 6, 7.

BORAGINACEAE (Heliotrope Family)

Cordia subcordata Lam. "sea trumpet", "kou" (Hawaii)
eongo, eoongo, eowongo (N); te kanawa (K); kanava (T)

Indigenous. Indian Ocean to Hawaii. Rare to occasional. Medium-sized tree, up to
10 m or taller, with spreading branches and pale greyish slightly fissured bark; leaves, 5
to 20 cm long and 4.5 to 15 cm wide, alternate, pale green, thinly coriaceous, ovate,
acute, base rounded, obtuse or truncate, lateral nerves 4- to 6-paired, margins entire or
slightly wavy; petiole, 2 to 8 cm long; flowers, few to several, in short-stalked axillary
and terminal cymose clusters; calyx, about 15 mm long, 3- to 6-lobed, lobes short-
deltoid, pubescent ventrally; corolla, 2.5 to 4 cm long and 3 to 5.5 cm across the mouth
(limb, which is plicate in bud)), pale to bright orange, crepe-like, trumpet-shaped,
scentless, 5 to 7-lobed, lobes 15 to 25 mm long, rounded; fruit, 2 to 3 cm long, ellipsoid
or nearly round, apiculate-acute, enclosed in a fibrous, somewhat corky calyx, green
when mature but aging to brown or black; seeds, 1 to 2 (rarely 3 or 4), coarsely
muricate. Found on coastal strip near settlement areas, either planted or protected. Soft,
durable wood considered by Nauruans to be excellent timber for woodcarving, boatbuil-
ding, construction and furniture; leaves crushed and mixed with coconut milk to prevent
baldness; flowers used in garlands. Trunk highly prized for woodcarving and canoe hulls
throughout Micronesia, Polynesia and Melanesia. 3(58756), 5(77), 6, 7.

107

Heliotropium procumbens Mill. var. depressum (Cham.) Fosb. & Sachet "heliotrope"
syns. H. gracile var. depressum Cham.; H. coromandelianum var. depressum
(Cham.) A. DC.; H. ovalifolium Forsk. var. depressum (Cham.) Merr.

Indigenous. Trop. America. Rare. Prostrate branched perennial herb with a stout
taproot; leaves, up to 3 cm long and mostly 3 to 6 mm wide, greenish-gray, pubescent,
oblanceolate or linear-oblong, subsessile; flowers, only moderately crowded, rather
sparsely pubescent, borne on one-sided coiled spikes on slender forked stalks (cincinni),
5 to 6 cm long; calyx lobes, unequal, longest to nearly 3 mm, lanceolate; corolla, 1.5 to
2 mm long, 5-lobed, white; fruit, globose, breaking into 2 to 4 1-seeded nutlets, about 1
mm high. Found on coastal strip near limestone outcrops? 4(141N), 7(22316).

Tournefortia argentea L.f. "beach heliotrope"
irin (N); te ren (K); tausunu, tauhunu (T)

syns. Messerschmidia argentea (L.f.) I.M. Johnst.; Argusia argentea (L.f.)
Heine; Tournefortia sericea Cham.

Indigenous. Indian Ocean to S.E. Polynesia. Common. Small to medium-sized,
wide-spreading, short-trunked tree, 2 to 12 m tall, with rather stout twigs and deeply
grooved bark; leaves, 10 to 30 cm long and 3 to 12 cm wide, alternate, spiralled,
crowded near branch ends, obovate-oblanceolate, decurrent at base, rounded-obtuse to
acute at apex, densely appressed, silvery-grey-pubescent on both sides, softly coriaceous,
somewhat fleshy; petiole, stout, winged; flowers, small, about 6 mm across, tube 2 mm
long, numerous, white, sessile, borne terminally on crowded, many-branched clusters of
tightly-coiled, densely-pubescent, scorpion-tail-like spikes or cymes; fruit, 5 to 8 mm in
diameter, round, greenish-white to brown, 4-parted, minutely apiculate; seeds (nutlets), 2
to 4, embedded in corky tissue. Found on flats behind beaches. Leaves eaten by pigs;
tender leaves and meristem pounded to prepare medicines for curing children’s rashes,
diarrhea, and fish poisoning; fruit blown through hollow papaya petioles by children.
3(58669), 5(32), 6, 7.

BRASSICACEAE OR CRUCIFERAE (Cabbage or Mustard Family)

Brassica alboglabra Bailey "Chinese kale"
kai laan ts’o1 (C)
syns. B. oleracea var. albiflora O. Kunze; B oleracea var. alboglabra (Bailey)
Musil

Pre-World War II introduction. Asia. Occasional. Erect, sometimes branching
perennial herb, up to 30 cm or higher, grown as an annual; leaves, 6 to 20 cm long and
4 to 15 cm wide, alternate, ovate, pale bluish-green (glaucous), glabrous, dull to shiny,

108

petiolate or not clasping; petioles, 4 to 10 cm long; flowers, up to 2.5 cm in diameter,
white, perfect, 4-sepaled, 4-petaled, without bracts, borne on slender pedicels along an
elongated fleshy terminal flower stem (raceme); fruit, a long, narrow pod, with a conical
beak; seeds, small, globose. Planted in Chinese food gardens at Location. Leaves, stems
and flowers cooked as a vegetable. 5, 6, 7.

Brassica chinensis L. var. chinensis "Chinese cabbage", "Chinese white cabbage"
te kabiti n Tiaina (K); kapisi Saina (T)
syn. B. chinensis Juslenius

Pre-World War II introduction. Asia. Common. Erect, loose-heading, biennial
herb, grown as an annual, up to 40 cm or higher, with a tap-root; leaves, 10 to 40 cm
long and 5 to 20 cm wide, alternate, shiny, dark-green, prominently white-veined, ladle-
shaped, terminally rounded, upstanding, radical (coming from the roots) leaves not lobed,
stem leaves usually clasping, not forming a compact head; petiole, 3 to 12 cm long,
thickened, ivory-white, somewhat fleshy; flowers, about 1 to 1.5 cm in diameter, light to
bright yellow, perfect, 4-sepaled, 4-petaled, without bracts, held above unopened buds,
petals orbicular, borne on slender pedicels, 1 to 4 cm long, along an elongated terminal
flower stem (raceme), up 50 cm long; fruit, 3 to 6 cm long, slender pods, with a conical
beak, usually 2-grooved, opening lengthwise; seeds, about 1 to 1.5 mm in diameter,
globose, brown to blackish-grey. Commonly cultivated in Chinese food gardens at
Location and Topside workshops. Leaves and stems cooked as a green vegetable. 5, 6, 7.

Brassica chinensis L. var. parachinensis (Bailey) Tsen & Lee
"flowering white cabbage" paak ts’oi sum (C); te kabiti n Tiaina (K)
syns. B. parachinensis Bailey; B. chinensis var. oleifera Makino

Pre-World War II introduction. Asia. Common. Erect, branched, biennial, non-
heading herb, grown as an annual, up to SO cm or higher; leaves, 10 to 50 cm long and 4
to 10 cm wide, darker-green above, lighter below, stem leaves not clasping, central stem
leaves, long and narrow, ovate to lanceolate to oblong, upstanding, not heading, radical
leaves more similar to those of var. chinensis; petiole, 3 to 12 cm long, slender, light
green; flowers, about 1 cm in diameter, light yellow, perfect, 4-sepaled, 4-petaled,
without bracts, borne on slender pedicels, 1 to 4 cm long, along an elongated terminal
and sometimes axillary flower stem (raceme), up 40 cm long; fruit, 3 to 6 cm long,
slender pod, with a conical beak, opening lengthwise; seeds, about 1 to 1.5 mm in
diameter, globose, brown to blackish-grey. Commonly cultivated in Chinese food gardens
at Location. Leaves, stems, and flowers cooked as a green vegetable. 5, 6.

109

Brassica juncea (L.) Czern. "mustard cabbage", "Indian mustard", "Chinese mustard"
kai ts’oi (C); te kabiti n Tiaina (K)
syn. Sinapis juncea L.

Pre-World War II introduction. Asia. Occasional. Erect, much-branched, non-
heading, annual herb, up to 80 cm or higher, with a deep tap root; stem leaves, not
clasping, basal (radical) leaves, up to 20 cm long, stalked, usually lyrate-lobed with very
large ovate terminal segment; flowers, about 1 cm in diameter, bright yellow, perfect, 4-
sepaled, 4-petaled, without bracts, not held above the central unopened buds, petals oval
to obovate with a long claw, borne on slender pedicels along an elongated terminal and
sometimes axillary flower stem (raceme), up 40 cm long; fruit, about 1.5 to 4 cm long,
slender pod, rounded in cross section, with a 4-angled short beak, opening lengthwise;
seeds, about | mm in diameter, globose, blackish-grey. Food plant in Chinese food
gardens at Location. Leaves and stems cooked as a green vegetable. 5, 6, 7.

Brassica oleracea L. var. capitata L. "English cabbage"
te kabiti ni Imatang (K); kapisi Palagi (T)

Pre-World War II introduction. Europe. Rare. Biennial herb, usually grown as an
annual, up to 30 cm or higher, or 30 cm in diameter, with a short stem and an arrested,
compact, much-swollen terminal bud surmounted by a mass of thick overlapping whitish-
green to light blue-green leaves forming a loose or compact, round or somewhat pointed
head; flowers, virtually never seen, not developing during the first year. Planted in
gardens near Buada Lagoon. Leaves cooked as a vegetable. 3, 6, 7.

Brassica pekinensis (Lour.) Ruprecht "celery cabbage", "Shantung cabbage",
"Peking cabbage", "pe-tsai" wong pa’ak, wong bok (C); te kabiti n Tiaina (K)
syns. B. chinensis var. pekinensis (Rupr.) Sun; B. petsai Bailey

Recent introduction. N. China. Rare. Erect, loose- to compact-heading, biennial
herb, grown as an annual, up to 50 cm or higher, somewhat resembling a large yellow-
green romaine or cos lettuce; leaves, 15 to 50 cm long and 5 to 25 cm wide, alternate,
light yellow-green, thin, veiny, crinkled undulate, dentate, without distinct petiole, the
wide, flat, long whitish base with a dentate wing, radical leaves often forming a compact
head, with midrib on the lower surface sometimes having sparse bristle-like hairs;
flowers, not seen; seeds, plump, without grooves. Planted in Chinese food garden at
Location. Leaves and stems cooked as a vegetable. 6.

110

Rhaphanus sativus L. var. longipinnatus Bailey “white radish", "daikon", "Chinese,
Japanese or Oriental radish" loh paak (C)

Pre-World War II introduction. E. Asia. Occasional. Erect annual bristly herb, up
to 50 cm or higher, with a large white cylindrical tap root, up to 30 cm long and 10 cm
in diameter, or larger, and normally weighing up to 500 g to | kg or more; radical
leaves, up to SO cm and 12 cm wide, dark green, bristly (setulose), lyrate-pinnatifid, with
8 to 11 pairs of pinnae; flowers, rarely seen, white to lilac, small, 4-petaled; fruit, 3 to 7
cm long, up to 1.5 cm in diameter, inflated, indehiscent, with 6 to 12 seeds; seeds, about
2.5 to 3 mm in diameter, dark brown, globose. Cultivated in Chinese food gardens at
Location. Leaves and fleshy white tap root cooked as vegetables or eaten raw. 5, 6.

CACTACEAE (Cactus Family)

Cephalocereus sp. "Cephalocereus"

Recent introduction. Tropical America. Rare. Jointed cactus with short, funnel-
shaped, night-blooming flowers. Planted ornamental. 6(204).

Hylocereus undatus (Haw.) Britt. & Rose "night-blooming cereus"

Recent introduction. Mexico. Rare. Large fleshy, green, epiphytic, rambling or
climbing, branching, night-blooming cactus, with 3-angled (3-winged) stems, about 4 to 5
cm in diameter, with scalloped margins, which become more or less horny and bear
cushions 2.5 cm or more apart, each with | to 3 short spines, and with aerial roots
growing from the underside of the stems; flowers, about 25 cm long, night-blooming,
lasting only until about noon of the following day, beautiful and showy, fragrant with a
mildly spicy odor, waxy, cup- or funnel-shaped, with many narrow, curved, yellowish-
green outer segments surrounding many white, erect petals, which are longer than the
mass of long, yellow-tipped stamens and a long style tipped with a radiating stigma
having about 24 lobes; fruit, rare, oblong, about 10 cm long, red, containing numerous
black seeds in white pulp, which is refreshing to eat. Immature ornamental pot plant. 6.

Opuntia sp. “prickly pear cactus"

Recent introduction. Tropical America. Rare. Succulent, erect, shrub-like cactus,
up to 2 m or higher, with flattened light-green, thick (up to 1 cm wide), oboval, paddle-
blade-like joints or cushions, about 12 to 30 cm long and 5 to 10 cm wide, smooth or
with spines, up to 1 cm long; leaves, small, nearly cylindrical, short-lived; flowers and
fruit, not seen. Planted ornamental. 5, 6.

111

w Ww

Schlumbergia truncata (Haw.) Moran "Christmas cactus", "Easter cactus", "crab cactus"
syn. Zygocactus truncatus (Haw.) Schum.

Recent introduction. Brazil. Rare. Small, smooth, day-blooming cactus, up to 30
cm or higher, with numerous, flat, dark glossy-green, jointed branches, about 4 to 5 cm
long and 1.5 to 2 cm wide, coarsely toothed, blunt-ended, that fork repeatedly in pairs,
resembling crab’s claws; flowers, many, 5 to 6 cm long and 1 to 1.5 cm wide, rose-red,
rather narrow tubular, curved, with many narrow oblong petals and sepals bent back
from long, clustered, protruding white, yellow-tipped stamens and a purple style; fruit.
Ornamental pot plant and planted ornamental. 6.

Unknown cactus "cactus"

Recent introduction. Rare. Small cactus pot plant. 6.

CAPPARIDACEAE (CAPPARACEAE) (Caper Family)

Capparis cordifolia Lam. “oceanic caper"
ekabobwiya, ekabobwija (N)
syns. C. mariana Jacq.; C. spinosa var. mariana (Jacq.) K. Schum.

Indigenous. Micronesia and Polynesia. Common. Small, sometimes sprawling,
spineless, woody shrub, up to about 1 m high, with rather soft and light wood; leaf
blades, about 3 to 5 cm long (rarely to 7 cm) and 2.5 to 5 cm wide, alternate, grayish-
green, slightly fleshy, broadly ovate or orbicular, rounded at both ends or truncate or
nearly subcordate at base, sometimes notched at the apex, lateral nerves 5- to 8-paired;
petiole, slender, about 8 to 14 mm long; flowers, large, attractive, fragrant, pea-flower-
like, asymmetric, axillary, solitary, borne on pedicels, 3 to 5 cm long, the bud opening
late in the evening and withering the next morning; sepals, up to 2 cm long, 2-seriate,
reflexed at anthesis, geleate; petals, 4, up to 2 cm long, attractive, white, glabrous,
obovate-suborbicular, glabrous; stamens, numerous (80 to 108), 3 to 4 cm long, showy,
white, but fading to pink or even purple; anthers, 3 mm long, purple; fruit, up to 4 to 5
cm long, an oblong or ellipsoid berry, with reddish ribs and elsewhere green, ripening to
dull yellow; seeds, about 3 mm long, dark brown, reniform. Found on limestone cliffs
and limestone rock outcrops on coastal strip. Crushed leaves used by Nauruans as a fish
poison. 2, 3(58636), 4(132N), 5(101), 6, 7.

2

Capparis quiniflora DC.
syn. C. richii A. Gray

Indigenous. E. Indonesia (Celebes and Lombok) to Melanesia and Nauru.
Occasional. Thorny, woody, high-climbing vine, with paired recurved spines, 1 to 3 mm
long; leaves, up to 10 cm long and 6 cm wide or larger, variable, narrowly lanceolate to
ovate; petioles, 5 to 20 mm long; flowers, small, 2 to 10 in number, serially arranged in
axillary or subterminal rows on pedicels, 6 to 20 mm long, which thicken and reach 5 cm
long when in fruit; sepals, up to 5 by 3 mm; petals up to 7 by 4 mm; fruit, about 40 by
35 mm, subglobose-ellipsoid. Found in plateau forest and on limestone pinnacles and
cliffs of the escarpment. 3(58591, 58799, 58804a), 5, 6.

Cleome rutidosperma DC.

Recent introduction. Origin? Common. Slender ascending to sprawling or
creeping, elongate, sparingly pilose, annual or short-lived perennial, odorless or nearly
odorless herb; leaves trifoliate on very slender petioles, up to 2.5 cm long; leaflets, 2 to
2.5 cm by about 1 cm, thin, the middle leaflet longest; flowers, axillary, small, petals
blue-violet or pink, fading to white, on filiform pedicels, up to 3 cm long; fruit, 3 to 4
cm long, a linear-fusiform capsule, on stipes 5 to 10 mm long; seed depressed-globose,
conspicuously rugose with long curving ridges or wrinkles, dark reddish-brown. Weed in
waste places, around buildings, and in areas recently cleared for phosphate mining.
3(58601, 58616, 58751), 4(108N), 5, 6, 7(22303), 8(9540).

Cleome viscosa L.
syns. C. icosandra L.; Polonisia icosandra (L.) W. & A.; P. viscosa (L.) DC.

Recent introduction. Trop. Asia or Old World Tropics. Common. Erect, sticky,
scarcely branching annual herb, up to 20 cm high, with longitudinally-grooved and
densely glandular hairy stems and a offensive odor; leaves, up to 7 cm by 7 cm,
alternate, compound, palmately 3- to 5-foliate, long petiolate, glandular pubescent;
leaflets, 1.5 to 4 cm long, the terminal leaflet longest, obovate to elliptic, tip acute to
rounded, base cuneate to oblique, margins entire; petiole, about as long a terminal
leaflet; flowers, solitary in upper leaf axils, on pedicels, 8 to 25 mm long; calyx,
caducous, 4-lobes, about 5 mm long, falling soon; petals, 4, about 7 to 12 mm long,
yellow, oblanceolate; fruit, 3 to 7 cm long, podlike, curved, longitudinally grooved,
densely pubescent, 2-valved, splitting longitudinally (dehiscent); seeds, about 1.3 mm in
diameter, roughly circular, numerous, small, reddish-brown, ridged (reniform). Weed
found primarily in lowland waste places, roadsides, and areas recently cleared for
phosphate mining. 3(58653), 4(125N), 5(45), 6, 7(22310), 8(9542, 9572).

113
CARICACEAE (Papaya Family)

Carica papaya L. “papaya”, "pawpaw"
dababaia, dababaiya (N); te mwemweara, te babaia (K); olesi (T)

Pre-World War I introduction. Trop. America. Occasional. Soft-wooded, un- or
few-branched, rather palm-like, small, quick-growing tree, up to 4 m or higher, with
thick, hollow, tapering, nearly smooth trunks or stems with light bark and numerous
almost heart-shaped leaf-scars, copious, thick, sticky, irritating milky sap, and leaves that
drop as the tree grows; trees usually male or female, although some are bisexual
(hermaphroditic) with both male and female flowers on the same tree; leaves, up to 60
cm or more across, alternate, clustered near top, large, round in outline, deeply pal-
mately 7- to 11l-lobed, the lobes irregularly acuminately toothed and lobed; petioles,
longer than blade, often over 1 m long, hollow; male flowers, about 2 cm wide,
numerous, white or cream-colored, fragrant, borne on loose clusters on long axillary
peduncles, 30 to 90 cm long; female flowers, about 4 to 6 cm wide, white, fragrant,
subsessile, 5-petaled, solitary or few together in leaf axils; fruit, 10 to 40 cm long,
variably shaped from subglobose or pear-shaped to cylindrical, green turning yellow or
orange, with orange to red-orange flesh; seeds, 3 to 5 mm long, globose, many, around
the edges of a central cavity, dark gray-green to black, wrinkled, caper-like, enclosed in
a firm gelatinous membrane. Fruit tree in home gardens and in contract worker gardens
at Location and the Topside Workshops. Ripe fruit eaten, and made into jam, primarily
by resident European community; fruit known to be a laxative; juice and flesh from
green fruit (which contains the enzyme papain) used to tenderize pork and beef; white sap
from small immature fruit used as a cure for ringworm; fragrant flowers used in
garlands; and hollow leaf petioles used by children as "pea shooters" for the fruit of
Tournefortia argentea (irin). Common, often naturalized, in houseyard gardens and
agricultural areas and an important local cash crop throughout the Pacific, and a
commercial export crop in areas such as Hawaii, Fiji, Tonga and the Cook Islands. 2,
3(58694), 5, 6, 7, 8.

CASUARINACEAE (Casuarina Family)

Casuarina equisetifolia L. "casuarina", "she oak", "ironwood", "beefwood"
tanenbaum (German for Christmas tree)(N); te katurina, te burukam (K); toa (T)
syn. C. litorea L.

Pre-World War I introduction. Indian Ocean to Polynesia and Micronesia.
Occasional. Medium to large, hard-wooded, fast-growing, pine-like tree, up to 15 m or
higher, with numerous, short-lived, long, thin, drooping, needle-like, gray-green,
cylindrical, jointed, striate, photosynthetic branchlets; leaves, whorled, reduced to

114

lanceolate or subulate, awl-shaped scales at each branchlet node; flowers, unisexual, in
catkins; male flowers, brown, terminal, with whorled bracts, in spike-like inflorescences
at the ends of branchlets; female flower heads, reddish-brown, condensed, lateral, each
flower in the axil of a bract, protected by two lateral bracteoles, the female catkin
enlarging, the bracts becoming woody in fruit; fruit, 1 to 2 cm long, globose, cone-like,
dull-green ripening to brown, many-seeded. Planted, occasionally as a street tree or
windbreak, and spontaneous on the coastal strip, and as a naturalized pioneer on some
mined areas to where they have spread from trees planted near the Topside workshops.
Although a common indigenous tree on sandy and rocky shores, and sometimes inland,
throughout most of the western Pacific, and possibly an aboriginal introduction to some
groups, such as Samoa, the casuarina seems to have been a recent post-European
introduction to Nauru. 2, 3(58776), 5(81), 6, 7.

CHENOPODIACEAE (Goosefoot or Saltbush Family)

Atriplex nummularia Lindl. "Australian saltbush"

Said to have been introduced in 1916, but not seen in 1978 or later.

Beta vulgaris L. var. cycla L. "Swiss chard", "silverbeet", "leaf beet"

Recent introduction. Europe. Rare. Biennial glabrous herb, up to 40 cm or taller;
leaves, up to 40 by 20 cm or more, dark green, shiny, alternate, simple, feather-shaped,
fleshy with thickened white midribs; flowers, rare, small, 3- to 4-flowered cymes
arranged on a spike, each subtended by a small narrow leaf; fruit, mostly an aggregate
formed by the cohesion of 2 or more fruits forming an irregular nutlike cluster held
together by swollen perianth bases. Food plant in Chinese garden at Location. Leaves
cooked as a spinach. 5, 6.

Spinacea oleracea L. "spinach"
poh ts’oi (C)

Recent introduction. S. W. Asia. Rare. Small biennial herb, up to 25 cm high;
leaves, 5 to 15 cm long and 3 to 8 cm wide, ovate to cordate, acuminate or somewhat
Sagittate; petioles, 2 to 10 cm long; fruit, an aggregate formed by the cohesion of 3 to 6
fruits forming an irregular nutlike cluster. Food plant in garden at Location. Leaves
cooked as a spinach. 5, 6, 7.

115
CLUSIACEAE OR GUTTIFERAE (Mangosteen Family)

Calophyllum inophyllum L. "Portia tree", "Alexandrian laurel", "beach mahogany",
"tomano" (Hawaii) iyo, ijo (N); te itai (K); fetau, itai (T); fetau (T)

Indigenous. Trop. Africa to E. Polynesia and Micronesia. Very abundant.
Medium to large, hard-wooded, often crooked, slow-growing tree, 10 to 20 m tall, with a
broad, low-branching, spreading crown, rough gray bark, and sticky yellowish sap;
leaves, 10 to 20 cm long and 6 to 10 cm wide, simple, opposite, dark green, glossy,
glabrous above, densely covered with minute silvery-white scales beneath, stiff, leathery,
elliptic-oblong, rounded or emarginate, base acute, pinnately nerved, with a stout pale
yellow-green midrib, lateral nerves very fine, numerous, parallel; petioles, 1 to 2.5 cm
long, stout, caniculate; flowers, about 2 cm in diameter, showy, waxy white, very
fragrant when fresh, 4- to -5 petaled (rarely 6 to 8), with numerous yellow stamens,
borne on pedicels, up to 3 cm long, in solitary axillary, 4 to 15-flowered racemose
clusters, up to 10 cm long; sepals, 4, concave-orbicular, the outer pair smaller, about 8
mm long, the inner pair, 10 mm long; fruit, about 2.5 to 5 cm long, green to purple-
black, globose, hard, heavy, thin-fleshed, with a bony shell which encloses a somewhat
poisonous single kernel or seed surrounded with cork, hanging from a long stalk.
Dominant large tree in original pre-mining plateau (Topside) forest vegetation, common
on escarpment slopes and on coastal strip. Timber provided the best wood for house posts
(iyor, yor), furniture, woodcarving and for canoe hulls in the past; sticky sap (erebeniyo)
used for caulking canoes; kernel of green and mature fruit (i kuan iyo ) crushed to yield
oil which is applied to hair to make it long and black; old decayed fruit skewered on
coconut midribs (engow) in past and burned as traditional Nauruan light; and mature fruit
burned as a mosquito repellent. Trees now being indiscriminately felled and burned as
refuse at a Topside dump to prepare land for phosphate mining. A tree highly valued for
its timber and other purposes throughout the Pacific, the seed kernel (known commercial-
ly as "punnai nut") yielding a dark green oil formerly exported from Fiji. 2, 3(58740),
4(164N), 5(120), 6, 7.

COMBRETACEAE (Terminalia Family)

Quisqualis indica L. Rangoon creeper"

Recent introduction. Trop. Asia. Occasional. An erect, vigorous shrub, growing
into a large woody vine, from 2 to 7 m long, with slender stems and rusty-brown young
growth; leaves 7 to 15 cm long, light green, soft, opposite, oval or oblong, acute, base
rounded; flowers, borne in short showy drooping clusters in the leaf axils, changing color
from pink to white as they open, and then darkening to red, pleasantly fragrant, calyx
tube, 0.5 to 7.5 cm long, tipped with 5 short points, 5 oblong petals and 10 short

116

stamens; fruit, about 2.5 cm long, dry, narrow ovoid, 5-angled, 1-seeded. Planted
ornamental. 3(58790), 5(113), 6, 7. :

Terminalia catappa L. “beach almond", "Indian almond", “Malabar almond", "tropical
almond", "coastal almond" __ etetah, eteto (N); te kunikun (K); talie, te ipe (T)

Indigenous. Trop. Asia and Australia to W. Polynesia and Micronesia. Common.
Medium to large, deciduous or semi-deciduous tree, up to 30 m tall, with whorled,
horizontal, wide-spreading branches arranged in tiers, and reddish timber; leaves, 15 to
30 cm long and 10 to 22 cm wide, clustered in rosettes at the ends of the branches entire,
leathery, shiny, dark-green turning red and yellow before dropping, new leaves appearing
almost immediately, obovate, broadly rounded, obtuse or acute, base narrowing,
narrowly and abruptly obtuse at petiole, pinnately-nerved, midrib slightly pubescent;
petioles, 0.5 to 1 cm long, stout; flowers, small, numerous (35 to 80 ), white, petal-less,
with a 5-lobed, bell-shaped calyx, which soon falls, and 10 stamens, borne in axillary,
spikelike racemes, about 10 cm to 24 long, near ends of branches; fruit, 5 to 7 cm long,
green turning yellow, then red, ovoid, subelliptic or subobovoid, hard, flattened, two-
keeled (narrowly-winged), drupe, with thin fleshy pulp surrounding a single edible
almond-like kernel. Tree in original plateau forest, on escarpment and on the coastal
strip; occasionally planted or protected in home gardens. Timber used in light construc-
tion and for woodcarving; roots used by some people to prepare a cure for dysentery;
outside rind of fruit eaten when yellow and kernel eaten after the mature fruit has fallen;
fruit strung as necklaces and used in black magic or sorcery. 3(58662), 5(116), 6, 7.

CONVOLVULACEAE (Morning-Glory Family)

Ipomoea aquatica Forsk. “water spinach", "swamp cabbage", "water convolvulus"
Lorenzo (N); ung ts’oi (C); cangcong (F)
syns. J. reptans Poir.; Convolvulus reptans L.; I. repens Roth

Pre-World War II introduction. Trop. Asia, Africa and Australia. Occasional.
Creeping, trailing or sometimes floating, semi-aquatic, hollow-stemmed, perennial
herbaceous vine, producing long shoots, rooting freely at the nodes; leaves, 5 to 15 cm
long and 2 to 10 cm wide, light green, soft and limp (flaccid), variable, commonly
oblong-lanceolate or heart-shaped, the base hastate or truncate, long-petiolate; petioles, 3
to 15 cm long, hollow; flowers, 1 to 7 in cymes, peduncles, 2 to nearly 18 cm long,
pedicels, 2 to 6 cm long; sepals, obtuse, ovate-oblong, 7 to 9 mm long; corolla, 3.5 to
5.5 cm long, pinkish-violet, often darker in the throat, rarely white; fruit, to 1 cm long,
ovoid; seeds, finely pubescent. Food plant in Chinese and Filipino gardens at Location
and Topside Workshop gardens; naturalized in muddy areas of Buada Lagoon, where it

LAG,

was formerly planted by the Japanese during World War II. Tender leaves and shoots
cooked as a green vegetable; usually propagated by cuttings. 5(24), 6, 7.

Ipomoea batatas (L.) Lam. "sweet potato", "kumara"
te kumara (K); kumala (T); fan shue (C)
syn. Convolvulus batatas L.

Pre-World War I introduction. Trop. America. Occasional. Smooth or pubescent
creeping, herbaceous, perennial, vine with milky sap, forming large, edible tuberous
roots; leaves, 4 to 15 cm long and 3 to 11 cm wide, green, limp, variable, deltoid-ovate
to ovate orbicular, entire or palmately 3- to 5-lobed or -parted, cordate or subcordate,
commonly with 2 prominent basal lobes, acute; petioles, slender, 3 to 15 cm long, often
longer than blade; flowers, several to many in long-peduncled cymes; corolla, usually 3
to 5 cm long, tubular and spreading, pale rose-violet, darker purplish in the throat, rarely
white; sepals (calyx-lobes), about 10 to 15 mm long, briefly mucronate; fruit, a capsule;
tubers, about 5 to 10 per plant, normally 10 to 30 cm long and 100 g to 1 kg in weight,
fusiform to globular, usually smooth but sometimes ridged, skin white, yellow, orange,
red, purple or brown, flesh waxy-white, yellowish, orange, reddish or purple, developing
in the top 25 cm of the soil by secondary thickening of adventitious roots. Cultivated in
contract laborers’ food gardens at Location and Topside workshops; spontaneous along
roadsides in some areas of Topside. Tuberous roots cooked as a staple and young leaves
of some varieties occasionally cooked as a green vegetable. Dominant staple food plant
and livestock feed in most of Papua New Guinea, parts of Solomon Islands and an
important supplementary staple in many areas of Polynesia and Micronesia; leaves are
also cooked as a vegetable green and an important livestock feed in some areas of the
Pacifies516; 7:

Ipomoea fistulosa Mart. ex Choisy "bush morning-glory"
syn. I. crassicaulis (Benth.) Rob.

Recent introduction. Brazil. Occasional. Perennial shrub, up to 3 m tall, with stiff
but rather rambling branches; leaves, 6 to 25 cm long and 4 to 17 cm wide, ovate to
heart-shaped, acuminate, midrib dorsally with 2 basal glands; petiole, 3 to 15 cm long;
flowers, mostly 5 to 9 cm long, tubular, rather pale pink, but darker within the tube,
day-blooming, borne in axillary, few-flowered cymes; sepals, 5, about 5 mm long, ovate
orbicular; fruit, 1.5 to 2 cm long, ovoid; seeds, brownish pubescent. Planted ornamental,
often along borders of home allotments. Has become naturalized in other areas of the
Pacific and in S. E. Asia. 3(58617), 5(198), 6, 7.

118

Ipomoea hederifolia L.
syn. I. angulata Lam.

Recent introduction. Trop. America. Rare to occasional. Slender, branching,
herbaceous, nearly glabrous, right-twining vine, up to 4 m high; leaves, up to 5 o 6 cm
or longer and about 4 cm wide, variable, entire or irregularly lobed, ovate-cordate,
acute, mucronate, basal sinus wide, marginally more or less coarsely dentate or sub-
lobate, 7-nerved from base, midrib branched upward, long-petiolate; petiole, slender, as
long as blade; flowers, few to several in branching, long-pedunculate, axillary clusters;
corolla, 3 to 4 cm long, bright red, attractive, tubular, limb flaring to about 2.5 cm in
diameter, calyx lobes long-pointed; fruit, 7 mm in diameter, a globose capsule, opening
by 4 valves; seeds, 4, about 4 mm long, black, densely short hairy. Found growing
spontaneously in ruderal sites on the coastal strip, climbing in a Leucaena leucocephala
thicket and over roadside shrubbery in Ijuw near Anibare boundary in 1987. 5(149), 6,
7(22307).

Ipomoea littoralis Bl.
syns. J. denticulata (Desv.) Choisy; Convolvulus denticulatus Desv.; I. gracilis
sensu auct. non R. Br.; /. choisiana Wight ex Safford

Indigenous. Malaysia and the Pacific. Rare. Slender, creeping or twining
herbaceous maritime vine, with glabrous stems; leaves, 4 to 10 cm long and 3 to 8 cm
wide, alternate, light-green, subglabrous, ovate-cordate (heart-shaped) to reniform, entire
or 3-lobed, acute or acuminate to obtuse, sometimes emarginate, base cordate with broad
sinus, 7-nerved from base, midrib branched upward; flowers, solitary or few on
peduncles, 1 to 4 cm long, pedicels 2 to 5 cm; sepals (calyx-lobes), all approaching 1
cm, but unequal, apex slightly mucronate, inner lobes wide as long; corolla, about 3 to 4
cm long, funnelform with spreading limb, dark pink to lavender, darker in the throat;
fruit, approaching 1 cm, a glabrous, 2-celled capsule; seeds, 4, glabrous. Roadside and
waste places. 8.

Ipomoea macrantha R. & S. "wild moon flower"
erekogo (N); te ruku (K); fue (T)
syns. Convovulus tuba Schlect.; 7. tuba (Schlecht.) G. Don; J. grandiflora sensu
Koidz. non Lam.; /. glaberrima Boj. ex Bouton; J. alba sensu Taylor non
L.; Calonyction tuba (Schlect.) Colla; C. comosperma Boj.

Indigenous. Pantropical. Occasional. Coarse, somewhat woody (sub-herbaceous),
creeping or twining, night-blooming (climbing) vine; leaves, 5 to 16 cm long and about
equally wide, alternate, ovate-suborbicular to heart-shaped, acuminate, mucronate, base
rounded, deeply cordate with open sinus, entire to slightly undulate, 7- to 9-nerved from
base, midrib branched upward; petioles, up to 8 cm or longer; flowers, solitary or few,
commonly in pairs, on elongate axillary peduncles, 8 to 14 cm long and 10 to 14 cm

119

across the limb, pedicels, about 2 cm long, bracteate; sepals (calyx-lobes) subequal, 1.5
to 2.5 cm long, broadly suborbicular, rounded, persistent, containing slimy sap; corolla,
7 to 12 cm long, funnelform, attractive, white with pale greenish bands, opening about
midnight, fading and wilting by morning; fruit, 2 to 3 cm long, globose, dry, borne in
the persistent calyx, splitting into 4 1-seeded capsules; seeds, about 1 cm long, dark,
glabrous or with two rows of minute hairs. Found climbing on trees in coastal and
plateau forests and creeping on open ground in some ruderal places. Leaves possibly used
medicinally by Nauruans? 3(58666, 58735), 4(131N), 5(118), 6, 7.

Ipomoea pes-caprae (L.) Sweet ssp. brasiliense (L.) v. Ooststr. "beach morning-glory"
erekogo, irekogo (Burges)(N); te ruku (K); fue (T)
syns. Convolvulus brasiliensis L.; C. maritimus Desr.; I. brasiliense (L.) Sweet

Indigenous. Pantropical. Abundant. Vigorous, prostrate, creeping, somewhat
fleshy, herbaceous, vine, with a long main root and reddish-purple stems, rooting at
nodes and extending up to 5 to 10 m; leaves, 5 to 12 cm long and wide, sometimes wider
than long, alternate, glabrous, somewhat goat-foot-shaped and folded along the midvein,
somewhat succulent, ovate-orbicular, base cordate to rounded, apex notched or 2-lobed,
areolate veiny; petioles, 2 to 10 cm long, sometimes longer than blade, with 2 glands at
apex; flowers, borne in solitary or several-flowered cymes, on long axillary peduncles to
about 15 cm long; sepals (calyx-lobes) to 1 cm long, ovate, mucronate or notched at
apex; corolla, 4 to 5 cm long, funnelform, dark rosy-pink to violet, purplish in the
throat: fruit, about 1.3. to 2.5 cm across, subglobose capsule, splitting at the top into 4
sections; seeds, 4, densely pubescent. Found on beach sand, not far from the sea, and on
coastal vegetation. Leaves crushed to yield juice which prevents hair from falling out. 2,
3(58729), 4(129N), 5(29), 6, 7.

Merremia quinquefolia (L.) Hall.f.
Convolvulus quinquefolia L.

Recent introduction. W. Indies. Herbaceous prostrate or climbing vine; leaves,
palmately compound with 5 leaflets, leaflets oblong to lanceolate, 2.5 to 6 cm long;
petioles, 2 to 5 (rarely up to 9) cm long; flowers, borne in axillary, solitary or several-
flowered cymes, the peduncles glandular in the upper part and sometimes mixed with
spreading bristly hairs; sepals, 4 to 8 mm long, narrowly ovate to oblong, obtuse,
subequal or the outer ones shorter; corolla, 1.8 to 2.5 cm long, pale yellow or whitish;
fruit, about 9 mm long, capsular, globose, stramineous, 4-valved; seeds, about 4.5 mm
long, shortly curled-pilose, blackish. Weed seen in one place in semi-open area on
escarpment. 3(58765).

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CRASSULACEAE (Orpine Family)

Kalanchoe pinnata (Lam.) Pers. “air plant", "miracle plant", "life plant"
te ang (K)
syns. Cotyledon pinnatum Lam.; Bryophyllum pinnatum (Lam.) Kurz; B. cal-
ycinum Salisb.

Pre-world War II introduction. Indian Ocean Islands. Occasional. Succulent,
glabrous, erect, un- or sparingly-branched, pale gray-green, perennial herb, with reddish
stems marked with oblong light spots, somewhat woody at the base, up to 60 cm or
higher; leaves, up to 20 cm long by 10 cm, opposite, simple or pinnately compound with
3 to 5 leaflets, broadly elliptic, margins crenate (scalloped), each crenation bearing at the
notch a vegetative bud which (when leaf is detached) may produce rootlets and stem,
eventually yielding a complete new plant; petioles, up to 10 to 12 long; petiolules, 3 to 5
mm long; flowers, 3 to 6 cm long, red or red-pink and green, cylindrical, pendent, calyx
and corolla 4-parted, dangling in terminal panicles; fruit, follicular. Planted ornamental.
Common pot plant; naturalized and spreading around water tank near Topside Workshop
food gardens. 2, 3, 5, 6, 7.

Kalanchoe tubiflora (Harvey) Hamet "chandelier plant"
syns. Bryophyllum tubiflorum Harvey; B. verticillata Scott-Elliot

Recent introduction. Madagascar. Rare. Succulent erect perennial herb, up to 50
cm or higher; leaves, 2 to 12 cm long and about 6 mm thick, generally dull green, with
purplish blotches or transversely striped, slender, nearly cylindrical or tubular (subterete),
usually whorled in threes at the stem nodes or crowded toward stem ends, 7-toothed at
the apex where vegetative buds develop; flowers, about 2 to 2.5 cm long, reddish,
pendent, corolla much longer than calyx, both 4-parted. Ornamental potplant. 3(58715), 4
(52).

CUCURBITACEAE (Melon Family)

Benicasa hispida (Thunb.) Cogn. "wax gourd", "ash pumpkin", "winter melon",
"white gourd" tung kwa (C)
syns. B. cerifera (Fisch.) Savi; Cucurbita hispida Thunb.; C. cerifera Fisch.

Pre-World War II introduction. Java. Occasional. Wide-spreading, herbaceous,
pubescent or hispid, annual climbing vine, to several meters, with short, branched
tendrils; leaves, 10 to 35 cm across, rounded, cordate, pubescent, palmately 5- to 11-
lobed; petioles, 10 to 20 cm long; flowers, 6 to 12 cm in diameter, monoecious, solitary,

121

axillary; calyx, 5-lobed; corolla, yellow, 5-petaled; male flowers on peduncles, 5 to 15
cm long; female flowers, subsessile, ovary densely hairy; fruit, 20 to 120 cm long and
15 to 80 cm wide, weighing up to 10 kg or more, melon-like, globose to oblong-
cylindrical, green, with a easy-to-remove white waxy covering, more or less hairy when
young; flesh (pulp), white, spongy; seeds, about 1 to 1.5 cm long and 5 to 7 mm wide,
numerous, central, buff-colored, smooth, flat, ovate-elliptic, with narrow base. Food
plant in Chinese gardens at Location. Flesh of fruit cooked as a vegetable. 5, 6, 7.

Citrullus lanatus (Thunb.) Matsum. & Tan. var. caffrorum (Alef.) Fosb. "watermelon"
te meren (K); meleni (T)
syns. Citrullus vulgaris var. caffrorum Alef.; C. vulgaris Schrad. ex Eckl. &
Zeyh.

Pre-World War II introduction. S. Africa. Rare. Slender, much-branched, wide-
spreading, prostrate, hairy, herbaceous, annual creeper, with rather thin, angular,
grooved stems, 1.5 to 5 m long, branched (bifid or trifid) tendrils and an extensive and
superficial root system; leaves, 5 to 20 cm long and 2 to 12 cm wide, alternate, scabrid
or harshly pubescent, deeply pinnately lobed, the lobes again pinnately lobed and toothed,
with broad apices; flowers, monoecious, solitary, axillary, on short pedicels, 3 to 5 cm
long, bell-shaped (campanulate), usually more male than female flowers; calyx, 5-lobed;
corolla, 2.5 to 3 cm in diameter, deeply 5-parted, pale yellow, petals, 1 to 1.5 cm long;
male flowers, greenish; female flowers, longer-pedunculate, ovary ovoid with woolly
hairs; fruit, large, up to over 50 cm long and 25 cm in diameter and weighing up to 4 to
20 kg, rounded (globose) or oblong; rind, mostly glabrous, dark-green, striped or
patterned, hard, but not durable; flesh (pulp), red to whitish-pink (rarely yellow), watery,
sweet; seeds, 6 to 15 mm long and 5 to 7 mm wide, many, black, sometimes white
mottled, smooth, flattened, with swellings on either side of the apex. Food plant in home
gardens and as spontaneous juveniles around residences and dump heaps. Fruit eaten raw.
DESKS 0 she

Cucumis melo L. var cantalupensis Naud. "cantaloupe", "rock melon"
te meren (K); meleni (T); heung kwa, t’im kwa (C)

Recent introduction. S. W. Asia and Africa to the Mediterranean. Occasional.
Softly hairy, trailing, herbaceous, annual vine, with ridged or striated stems, tender
unbranched tendrils, and an extensive and superficial root system; leaves, 7 to 13 cm
across, simple, rounded or kidney-shaped, usually S5-angled and sometimes shallowly 3-
to 7 lobed; petiole, 8 to 16 cm long; flowers, about 2.5 cm across, generally andro-
monoecious (with hermaphroditic and male flowers on the same plant), yellow; male
flowers, often in groups, each on a slender pedicel; hermaphroditic (and female) flowers,
solitary, with shorter and thicker pedicels; fruit, 12 to 24 cm long and 10 to 18 cm in
diameter, globose or oblong, hollow; rind, thick, scaly, rough, netted or often deeply
grooved; flesh (pulp), soft, juicy, pale-orange, slightly sweet or fragrant; seeds, 8 to 12

122

mm long, many, whitish or buff, smooth, flat, in a central cavity with fleshy placentas.
Food plant in Chinese gardens at Location. Fruit flesh eaten raw. 5,-6.

Cucumis melo L. var. conomon Makino "Oriental pickling melon"
ts’it kwa (C)

Pre-World War II introduction. China. Rare. Softly hairy, branching, trailing,
occasionally climbing, herbaceous, annual vine, with ridged stems, tender unbranched
tendrils, and an extensive and superficial root system; leaves, 7 to 13 cm across, simple,
rounded to heart- or kidney-shaped; flowers, about 2.5 cm across, generally andro-
monoecious (with hermaphroditic and male flowers on the same plant), yellow; male
flowers, often in groups, each on a slender pedicels; hermaphroditic (and female)
flowers, solitary, with shorter and thicker pedicel; fruit, 20 to 30 cm long and 6 to 8 cm
in diameter, oblong, cylindrical, cucumber-like, slightly hollow; rind, thin, pale whitish-
green to yellowish-green, longitudinally lined, smooth or slightly pubescent; pulp, soft,
white to greenish-white, almost tasteless; seeds, 5 to 10 mm long, many, whitish or buff,
smooth, flat, in a central cavity with fleshy placentas. Trailing or climbing food plant in
Chinese gardens at Location and Topside. Fruit cooked as a vegetable, often in soups. 5,
6.

Cucumis sativus L. "cucumber"
te kukamba (K); kukampa (T); tseng kwa, wong kwa (C)

Pre-World War II introduction. N. India. Rare. Scabrid, climbing or trailing,
herbaceous, annual vine, climbing from 1 to 5 m, with strongly 4-angled stems, un-
branched tendrils and an extensive and superficial root system; leaves, 8 to 20 cm long,
hispid or rough, base cordate, apex acuminate, scarcely angled, unlobed or shallowly 3-
to 5-lobed with acute sinuses, dentate, palmately S- to 7-nerved; petioles, 5 to 18 cm
long; flowers, 2 to 4 cm long, monoecious, yellow, bell-shaped, deeply 5-partite, hairy,
wrinkled; calyx, 5 to 10 mm long, with 5 narrow lobes; male flowers, predominant,
borne in axillary clusters on slender pedicels; female flowers, solitary or few, axillary,
on stout peduncles; fruit, 15 to 25 cm long, pendulous, slenderly oblong or cylindric,
often slightly curved, dark to light-green skinned, glabrous or covered with tiny bristly
tubercles or warts (echinate), particularly when young; flesh, pale whitish-green; seeds, 8
to 10 mm long and 3 to 5 cm wide, many, whitish, oblong, flat, margin defined only at
the apex. Food plant in houseyard gardens. Fruit eaten raw, and occasionally cooked. 5,
ea

123

Cucurbita maxima Duch. "pumpkin", "winter squash", "autumn squash"
dabamakin (N); te baukin, te bamakin, te bangke (K)

Recent introduction? S. America. Rare. Prostrate, long-trailing, slightly rough,
herbaceous, annual vine, with branched tendrils; stems, soft, round in cross-section;
leaves, 6 to 19 cm long and 7 to 20 cm wide, slightly kidney-shaped (subreniform),
nearly orbicular in outline, shallowly 5-lobed, cordate with a very deep sinus, margins
denticulate, green, occasionally with white blotches, coarsely pubescent, not rigid;
petioles, up to 20 cm long; flowers, solitary, monoecious, bell-shaped, acute or obtuse in
bud, corolla lobes curved outward, bright yellow; male flowers, 4 to 7 cm long on
peduncles, 10 to 17 cm long; sepal lobes, linear; calyx lobes, narrow subulate; female
flowers, on a soft corky peduncle, thickening to a fruit-stalk, up to 7 cm long, thicker
than stem, spongy, nearly cylindrical, not expanded at point of attachment to fruit; fruit,
variously shaped, hard, green to dull orange or yellow, often mottled, usually hollow;
flesh, yellow to orange, rather fibrous; seeds, 16 to 20 mm long and 8 to 12 mm wide,
not separating easily and cleanly from pulp, dull white or buff, usually smooth or glossy,
sometimes with fine wrinkles, ovate, attachment acute and asymmetrical, apex nearly
straight across or oblique, margin or rim, smooth, obtuse, single, slightly raised. Food
plant, often adventive in gardens. Fruit cooked as a vegetable. 3.

Cucurbita pepo L. "pumpkin", “field pumpkin", "summer squash", "marrow"
dabamakin (N); te baukin, te bamakin, te bangke (K); panikeni (T)

Pre-World War II introduction. Trop. America. Prostrate, long-trailing (rarely
somewhat erect), slightly rough, hard-stemmed, herbaceous, annual vine, with 4- to 5-
branched tendrils; stems, harsh to touch and spiculate, round in cross-section; leaves, 6
to 19 cm long and 7 to 20 cm wide, broadly triangular or heart-shaped, often pointed,
palmately, deeply 5-lobed, broad sinus between lobes, central lobe longest, rather finely
toothed, harsh to the touch and spiculate, more prickly than C. maxima, cordate with
narrow, acute sinuses, green, occasionally with white blotches, stiff and more or less
rigid, foliage held upright; petioles, up to 15 cm long; flowers, solitary, monoecious,
bell-shaped, acuminate in bud, corolla lobes nearly always upright, bright yellow to
yellow-orange; sepals, short, awl-shaped; calyx lobes, narrow subulate; male flowers, 5
to 6 cm long on peduncles; female flowers, on a hard, sharply 5-angular, grooved
peduncles, maturing into a short fruit-stalk, without cork development and not, or only
slightly enlarged at point of attachment to fruit; fruit, variable, usually spherical-oblate,
dull orange to orange-brown, sides radially round-ridged, hollow; flesh, orange, coarse-
grained, rather fibrous; seeds, 10 to 18 mm long and 8 to 11 mm wide, separating easily
and cleanly from pulp, dirty white or creamy white, smooth or finely granular, ovate,
attachment obtuse and symmetrical, apex usually straight across, thin or fairly plump,
margin or rim, smooth, obtuse, clearly defined, usually appearing double. Occasional.
Food plant, often adventive in waste places and gardens at Location and elsewhere on the
coastal strip. Fruit cooked as a vegetable. 3, 5, 6, 7.

124

Luffa acutangula (L.) Roxb. "angled loofah", "vegetable sponge", "ridge gourd",
"dish-cloth gourd" sze kwa (C)
syn. Cucumis acutangulus L.

Pre-World War II introduction. India, Paleotropics. Occasional. Stout, climbing,
glabrous, herbaceous, annual vine, with pentagonal stems, 3- or more-forked tendrils and
fetid when bruised; leaves, 10 to 24 cm long and wide, rounded, rather shallowly
palmately 5- to 7-lobed, coarsely and shallowly toothed, scabrous, pale beneath; flowers,
4 to 5 cm in diameter, monoecious, axillary, pale yellow, 5-petaled, petals almost
completely distinct, male and female flowers borne together in the same axil, opening in
the late afternoon or evening; male flowers, in racemes of several flowers, peduncle 10
to 25 cm long, stamens 3; female flowers, solitary, stigmas 3, ovary 10-ribbed; fruit, 15
to 40 cm long and 5 to 10 cm in diameter, green to gray-green, very narrowly obovoid,
with 10 acute longitudinal ridges or ribs, crowned with enlarged sepals and style; flesh,
white, somewhat spongy, becoming fibrous when old; seeds, 10 to 13 mm long and 7 to
9 cm wide, black, pitted, flattened. Climbing foodplant in Chinese gardens at Location
and Topside. Young green fruit cooked as a vegetable. 4(162N), 5, 6, 7, 8(9577).

Luffa cylindrica (L.) Roem. var. insularum (A. Gray) Cogn. "smooth loofah",
"wild vegetable sponge", "scrubber gourd", "dish-cloth gourd"
syns. Momordica cylindrica L.; M. luffa L.; Luffa aegyptica Mill.; L. insularum
A. Gray

Pre-World War II introduction; although variety insularum seems to be indigenous
to many Pacific islands, it was probably introduced to Nauru. Trop. Asia. Locally
common. Vigorous, climbing, somewhat pubescent, herbaceous annual vine, with
pentagonal stems and 2- to 3-, or more-forked tendrils; leaves, to 20 cm long and nearly
as wide, much longer than petiole, orbicular-ovate to almost kidney-shaped (subreni-
form), deeply 5- to 7-angled or -lobed, acuminate or acute, base deeply cordate with
open sinus; petiole, S to 10 cm long, scabrid or hispid; flowers, 5 to 9 cm in diameter,
monoecious, axillary, yellow, petals almost completely distinct, male and female flowers
in the same axil, opening in the early morning; male flowers, 4 to 20, crowded near the
end of a solitary axillary peduncle, stamens 5; female flowers, solitary, stigmas 3, ovary
not ribbed; fruit, 12 to 30 cm long, oblong-cylindrical, smooth or slightly ribbed or
striped, crowned with stout sepals and style, fleshy at first, becoming dry; flesh, white,
spongy, becoming fibrous when old; seeds, 10 to 15 mm long, black, smooth, flat,
faintly winged. Weedy found on edges of forest, waste places, and spreading over
Topside topsoil dump on plateau. 3(58587, 58731), 5(75), 6, 7.

125

Momordica charantia L. "bitter gourd", "bitter melon", "balsam apple"
fu kwa (C); ampalaya (P)

Pre-World War II introduction. Paleotropics. Occasional. Slender, high-climbing,
herbaceous, annual vine, 2 to 4 m long, with slightly pubescent, 5-angled, furrowed.
stems and unbranched tendrils; leaves, 5 to 15 cm in diameter, palmately 3- to 5- lobed,
the lobes more or less undulate or coarsely toothed, base cordate, smooth or slightly
pubescent; petiole, 10 to 12 cm long; flowers, 2.5 to 3.5 cm in diameter, monoecious,
axillary, yellow, corolla deeply-lobed; calyx, deeply 5-parted; male flowers, single or
racemose, stamens 3; corolla, deeply 5-parted, 1.5 to 2 cm long; female flowers,
solitary, on peduncles with a kidney-shaped bracteole at base, stigmas 3; fruit, 5 to 20
cm long, pendulous, ovoid to oblong-cylindric or pear-shaped, irregularly longitudinally
ridged with numerous bumpy tubercles, short-pointed at tip, light green to orange or dark
yellow when ripe, splitting at the tip into three irregular valves; pulp, bitter when green
and slightly sweetish when ripe; seeds, 12 to 16 mm long and 5 to 9 mm wide, numer-
ous, light brownish to light gray patterned, covered with a soft, fleshy red aril. Food
plant in Chinese and Filipino gardens at Location and in Chinese gardens at Topside
workshops and surrounding Buada Lagoon. Fruit and young leaves occasionally cooked
as vegetables. 4(144N), 5, 6, 7.

ERICACEAE (Heath Family)

Rhododendron sp. "azalea"

Recent introduction. Rare. Asia. Evergreen shrub. Flowers not seen. Planted
ornamental. 6.

EUPHORBIACEAE (Spurge Family)

Acalypha amentacea Roxb. var. wilkesiana (Muell.-Arg.) Fosb.
“copper leaf", Jacob’s coat" "beefsteak plant", "fire dragon plant"
Kayser bush (N); te aronga (K); kalakalaapuki, kakarapus, ogoogo, lakau kula (T)

syn. A. wilkesiana Muell.-Arg.

Pre-World War I introduction. Melanesia. Common. Perennial shrub, up to 2 m
or higher, with pubescent young branches; leaves, 5 to 20 cm long and 3 to 15 cm wide,
alternate, simple, mostly ovate, crenate-serrate, acuminate, pubescent, rather curved and
coarsely crisped, pinnately-nerved, attractive, commonly dark or bright-red, red-green or

126

green, often mottled with various shades of red, dark pink or bronzy green, sometimes
with whitish or reddish-white margins resembling a thin marbling of fat on a beefsteak;
petioles, 2 to 10 cm long; stipules, 3 to 11 mm long, oblong-lanceolate; flowers,
monoecious, tiny, about | cm across, petal-less, borne alternately on slender axillary or
terminal spikes, 4 to 15 cm or longer; bracts, about 4 to 10 mm across, red to deep
purple, S- to 13-toothed, only slightly cupular, subtending single flowers; male spikes
slender, elongated, hanging among lower leaves, accompanied by smaller bracts; female
spikes somewhat shorter, nearly erect, among leaves at branch tips, each cluster (1 to 5)
accompanied by a larger triangular, toothed bract, often with a rounded central lobe;
ovary, 1.5 to 2 mm wide, green, puberulent; styles whitish, hair- or bristle-like, curved;
fruit, a small capsule. Planted ornamental and hedge plant. 3(58743, 58757), 5(41), 6, 7.

Alcalypha amentacea Roxb. var. wilkesiana f. circinata (Muell.-Arg.) Fosb.
te aronga (K)
syn. A. wilkesiana Roxb.f. circinata Muell.-Arg.

Recent introduction. Pacific Is. Occasional. Perennial shrub, up to 2 m tall;
leaves, 3 to 14 cm long and wide, suborbicular or reniform or broadly ovate, often
rounded at the apex, somewhat flabellinerved, green, commonly variegated with a white
margin. Planted ornamental, often as hedges. 3(58769), 5, 6, 7.

Acalypha hispida Burm.f. "cats’ tail", "chenille plant", "red-hot poker"

Recent introduction. Indonesia. Rare. Perennial shrub, up to 2 m high; young
branches pubescent; leaves, 5 to 18 cm long and 3 to 10 cm wide, green, alternate,
ovate, dentate-crenate, acuminate, pinnately-nerved; petioles, 3 to 9 cm long; stipules, 5
to 10 mm long, oblong-lanceolate; flowers (female), axillary, petal-less, borne in
pendulous, bright-red, velvety, tassel-like spikes, 10 to 50 cm long and 1 to 2.5 cm in
diameter, resembling a cat’s tail; bracts, minute, entire, subtending flower clusters; fruit,
not seen. Planted ornamental. 3(58705), 5, 6,.

Breynia disticha J.R. & G. Forst var. disticha f. nivosa (W.G. Smith) Croizat
"leaf-flower of the Pacific islands", "snow bush" eomonon (N)(Burges, 1933)
syns. B. nivosa (Bull) Small; Phyllanthus nivosus Bull

Pre-World War II introduction. Pacific Is. Rare. Perennial glabrous shrub, up to 3
m or taller, with slender, densely, two-ranked branches; leaves, 2.5 to 5 cm long,
alternate, 2-ranked or spirally arranged, oval-ovate, entire, green, variegated with white,
pink ,light red or purple, obtuse or abruptly pointed; petioles, short; stipules, very small,
triangular acute; flowers, monoecious, small, green, petal-less, on short axillary pedicels;
male flowers, 3 to many, borne in close clusters (fascicles) in the lower axils; female

127

flowers, solitary, with a green, 6-lobed or -toothed, bell-shaped or cuplike calyx, borne
in the upper axils. Planted ornamental. 2, 3, 5, 6, 7.

Codiaeum variegatum (L.) A. Juss. "croton", "codiaeum"
syns. Croton variegatum L.; Phyllaurea variegata (L.) Wight; P. codiaeum
Lour.

Pre-World War II introduction. Malaysia to Melanesia. Common. Glabrous,
densely-branched, shrub or small tree, up to 4 m or taller; leaves, 7 to 32 cm long and 2
to 15 cm wide, alternate, spirally arranged, thick, leathery, extremely variable in shape,
size and color, commonly entire, sometimes interrupted, wavy, twisted or 3- or more-
lobed, linear-lanceolate, oblong or ovate to obovate, mostly acuminate, pinnately-nerved,
plain, spotted, mottled, variegated, or decoratively patterned green, red, purple, pink,
yellow and white, nerves often brightly colored, red or yellow; petioles, 1 to 9 cm long;
flowers, monoecious, small, petal-less, borne in lax axillary narrow spikes, 15 to 25 cm
long; male flowers, about 6 mm wide, with 5 or 6 sepals and petals and 15 to 35 free
stamens, chestered; female flowers, solitary, lacking petals, scattered; fruit, small, a
subglobose, thin-walled, 3-lobed capsule. Planted ornamental and hedge plant. 3(58714),
5, 6(199, 200), 7.

Euphorbia antiquorum L. "cactus spurge", "false cactus", "Malayan spurge tree"

Recent introduction. India, S.E. Asia. Rare. Erect, spiny, cactus-like shrub, up to
3 m or higher, with thick, green, succulent, ridged, 3- to 4-angled stems and branches;
spines, up to 7 mm long, brown, on ridge elevations; leaves, none or commonly falling
soon, 6 to 13 mm long, thick, oval; flowers, small, yellowish or greenish, borne in upper
leaf axils. Planted ornamental. 3(58680), 5(130), 6, 7.

Euphorbia chamissonis (K]. & Gke.) Boiss. "beach spurge"
e mai (B)(N); te tarai (K); teluna (T)
syns. £. atoto sensu auct. (Burges) Micr. non Forst.f.; Anisophyllum
chamissonis Kl. & Gke; Chamaesyce atoto sensu auct. non (Forst.f.)
Croizat

Indigenous. Trop. Pacific. Rare. Erect to semi-erect or sprawling, freely-branch-
ing, glabrous subshrub, up to 1 m tall, with decumbent or ascending stems, which are
branched, appear jointed and thickened at the nodes, a woody rootstock and white sticky
latex; leaves, 1.5 to 3 cm long and | to 2 cm wide, opposite, entire, somewhat fleshy,
ovate-oblong, oval-ovate or oblong subspatulate, base cordate and somewhat unequally 2-
sided, apex acute to obtuse, sometimes mucronate, palmately veined, glabrous, light-
green or glaucous above, whitened beneath; petioles, up to 2 mm _ long; flowers,
numerous, monoecious, white or greenish, petal-less, in cyathia borne in small axillary or

128

terminal branching clusters (cymes); involucres of cyathea usually 1.7 to 2.3 mm long,
glands with minute appendages; fruit, about 3 mm long, a glabrous, dehiscent capsule;
seeds, small, smooth, released when dry fruit opens. Spreading shrub on seashore.
2(38.5), 3, 5(106), 6, 7.

Euphorbia cyathophora Murr. “painted leaf", "Mexican fire plant", "hypocrite
plant", "wild poinsettia", "Christmas bush", "dwarf poinsettia"
deriba, deribeh (N); te kabekau (K)
syns. E. heterophylla L. var. cyathophora (Murr.) Griseb.; Poinsettia cyatho-

Phora (Murr.) Kl. & Gke.

Pre-World War II introduction. Trop. America. Occasional. Erect annual herb, up
to 80 cm or higher, with glabrous, green, longitudinally-grooved stems, sparingly
pubescent young shoots and white latex; leaves, 3 to 8 cm long and 1.5 to 3.5 cm wide,
alternate below, opposite above or in threes, somewhat fiddle-shaped (pandurate),
pinnately, sharply and shortly 2- to 4-lobed, usually with a deep, rounded notch or
incision on each margin, those of the basal leaves sometimes simply ovate, entire or
obscurely distantly serrate, mostly glabrous, the upper leaves crowded beneath the
flowers, the uppermost, bract-like leaves near the flowers with a bright red basal blotch;
petioles, 1 to 4 cm long, slender; flowers, monoecious, small, inconspicuous, greenish,
petal-less, in cyathia packed in short terminal clusters (cymes), 2 to 4 cm wide; involucre
of cyathium, 3 to 5 mm long, on a stalk of equal length, cup-shaped, 5-lobed, green,
enclosing several male flowers and 1 female flower, and bearing a funnel-shaped gland
with a large elongated opening, about 1.5 cm long; male flowers, tiny, numerous,
surrounding the female flower and reduced to a single stamen, anthers, yellow; female
flowers, one per cyathium, consisting of a stalked, 3-lobed ovary; fruit, 3 to 5 mm long,
glabrous, subglobose, 3-lobed capsule (schizocarp), splitting at maturity into three 1-
seeded segments; seeds, 2 to 2.5 mm across, dark brown, tuberculate, obovoid or
globose, basally flattened, apically subconic. Roadside and waste place weed. 1(23.R), 2,
3(58628), 4(104N), 5(31), 6, 7(27818), 8(9556).

Euphorbia geniculata Ortega "wild spurge"

Recent introduction. Texas, Mexico and the W. Indies. Occasional. Erect annual
herb, to about 60 cm or higher, with somewhat hairy, green hollow stems and white
latex; leaves, 3 to 8 cm and 1.5 to 3.5 cm wide, basal and apical leaves opposite, other
leaves alternate, entire, ovate or oblong-rhomboidal, acute-acuminate, obscurely dentate
toward base, lower leaves all green, glaucous beneath, the uppermost bract-like leaves
near the flowers with a white or pink basal blotch; petioles, 1 to 5 cm long; flowers,
monoecious, small, inconspicuous, greenish, petal-less, in cyathia packed in short
terminal clusters (cymes), 2 to 4 cm long; involucre of cyathium, cup-shaped, 5- to 7-
lobed, green, enclosing several male flowers and 1 female flower, and bearing a single
funnel-shaped gland at one side; male flowers, tiny, numerous, surrounding the female

128

flower and reduced to a single stamen, anthers, yellow; female flowers, one per
cyathium, consisting of a stalked, 3-celled ovary with 3 bifid styles; fruit, 3 to 5 mm
across, subglobose, 3-lobed capsule (schizocarp), splitting at maturity into three 1-seeded
segments; seeds, 2.5 to 3.2 mm across, grey-brown, angular, sharply keeled on one side,
slightly tuberculate. Weed of roadsides and wasteplaces. 5(72), 6, 7, 8(9580).

Euphorbia heterophylla L. "wild spurge"
syn. Poinsettia heterophylla (L.) Klotzsch & Garcke

Recent introduction. Mexico. Occasional. Erect, unbranched annual herb, up to 70
cm high, with hollow stems; leaves, alternate, the lower entire, oval or elliptic, with
irregularly-toothed margins, the upper leaves deeply lobed, the topmost leaves green or
with a purple-spotted (never red) base and clustered beneath the inflorescences; inflores-
cences, separately male and female borne in terminal cyathia; cyathium, a 5-lobed green
cup enclosing several male and one female flower and bearing a gland with a small
circular opening; fruit, a 3-celled capsule, explosively dehiscent; seeds, 1 to a cell, 2 to
2.5 mm across, subglobose, dark brown to black, rugose or tuberculate, with a keel and
a transverse groove at right angles to it. Weed of roadsides and ruderal sites. 3(58623,
58667), 4(102N).

Euphorbia hirta L. "garden spurge", "asthma plant", "hairy spurge", "old blood"te
tarai, te tarai Kutaie ("Kusaie, Kosrae")(K)
syns. E. pilulifera L.; Chamaesyce hirta (L.) Millsp.

Pre-World War II introduction. Pantropical. Abundant. Erect or decumbent,
scarcely branching, widely spreading, densely hairy annual herb, 6 to 60 cm high, with
white latex; leaves, 1 to 4 cm long and 0.5 to 1.5 cm wide, opposite, simple, elliptical,
oblong or ovate-rhomboidal, acute to obtuse tip, base obliquely acute, finely serrate,
stipulate, leaf surfaces appressed pubescent, green, often with red, brown or purplish
tinge, paler beneath; petioles, 2 to 3 mm long; flowers, monoecious, very small, greenish
white, petal-less, in cyathia arranged in 1 to 2 dense, axillary, globose clusters (cymes)
on short pedicels, 3 to 15 mm long; involucre 4- to 5-lobed, enclosing several male
flowers and | female flower, and with minute glands; male flowers, minute, surrounding
the female flower, each consisting of a single stamen; female flowers, on per cymatium,
consisting of a 3-celled ovary; fruit, 1 to 1.3 mm across, globose, hairy, brown, 3-lobed
schizocarp splitting into three 1-seeded segments at maturity; seeds, about 1 mm long,
irregular oblong with some faint transverse ridges. Weed in waste places and open areas;
pioneer plant in recently mined areas. 2, 3(58627, 58677), 4(114N), 5(21), 6, 7,
8(9554).

130

Euphorbia hypericifolia L. "spurge", "graceful spurge"
te tarai (K) :
syn. Chamaesyce hypericifolia (L.) Millsp.

Recent introduction. Trop. America. Occasional. Slender, reddish-stemmed,
glabrous, branching herb, 10 to 50 cm high, with a taproot and white latex; leaves, 1 to
2.5 cm long and 0.3 to 1 cm wide, thin, opposite, elliptic or elliptic-lanceolate, entire or
minutely toothed, base oblique, green, rarely pinkish or reddish-purplish, glaucous
below; petioles, short, about 1 to 2 mm long; flowers, monoecious, small, inconspicuous,
greenish, petal-less, in cyathia borne in small cymes, on pedicels, 1.2 to 3 cm long, in
upper axils; involucral bracts, white, aging to pink; fruit, a 3 lobed capsule, with one
seed to a lobe, and forked styles; seeds, less than 1 mm long, ribbed, reticulate, dark red
to purplish. Garden and ruderal weed. 3(58658), 5(72), 6(223), 7, 8(9555).

Euphorbia milii Ch. des Moul. var. splendens (Bojer) Ursch & Leandri
"crown of thorns"
syn. E. splendens Bojer

Recent introduction. Madagascar. Rare. Erect branching shrub, up to 1 m or
higher, with hard, succulent, nearly cylindrical, subterete or slightly ridged stems,
sometimes twining, beset with many slender spines, up to 2.5 cm long; leaves, 2 to 5 cm
long and 1.5 to 4 mm wide, obovate, thinly fleshy, bright green, crowded toward branch
tips, soon falling; flowers, monoecious, very small, petal-less, in cyathia borne on long
axillary peduncles near the branch tips, each subtended by two opposite, rounded, wing-
like, bright- or salmon-red bracts, 8 to 10 mm long. Planted ornamental and pot plant.
6(178).

Euphorbia prostrata Ait. "prostrate spurge"
te tarai (K)
syn. Chamaesyce prostrata (Ait.) Small

Pre-World War I introduction. Trop. America. Abundant. Prostrate annual herb,
much-branched at base, with white latex and slender purplish or pinkish stems, 2 to 20
cm long, which are glabrous or minutely puberulent on one side only; leaves, 2 to 10
mm long and 1.5 to 6 mm wide, opposite, simple, oblong-elliptic to oblong-obovate, tip
rounded to obtuse, base rounded to obliquely unequilateral, margins finely serrate
(serrulate), glabrous or sparsely puberulent dorsally, glaucous with a purplish tinge;
petioles, short, about 1 mm long; stipules, on upper side of stem distinct, linear, pilose,
those of lower side fused, apically toothed; flowers, monoecious, very small, purplish,
petal-less, borne in cyathia arranged in short, few-flowered axillary clusters (cymes) on
short pedicels; involucre, 4 to S- lobed, enclosing several male and 1 female flowers, and
bearing purple glands; male flowers, minute, surrounding the female flower, each
consisting of a single stamen; female flowers, one per cymatium, consisting of a 3-celled

131

ovary; fruit, about 1 mm long, subglobose, 3-lobed schizocarp, sparsely hairy on the
margins, pinkish or purplish, on a stalk as long as the fruit, splitting at maturity into
three 1-seeded segments; seeds, about 0.8 mm long, narrowly ovoid, truncate basally,
apically obtuse, tetragonal, reddish or grayish, 5- to 7-grooved transversely. Weed along
paths and open waste places; pioneer plant in recently mined areas. 1(62.R), 3(58630,
58631), 4(116N), 5(22), 6, 7, 8(9552).

Euphorbia pulcherrima Willd. ex Klotszch "poinsettia"
syn. Poinsettia pulcherrima (Willd.) R. Grah.

Recent introduction. Mexico. Rare. Erect, glabrous, branching, soft-wooded
shrub, up to 2 m or higher, with white latex; leaves, 8 to 25 cm long and 5 to 15 cm
wide, elliptic to elliptic-ovate, acute, entire or slightly sinuate-toothed or lobed, often
somewhat fiddle-shaped, bright green, pubescent; petioles, 3 to 8 cm long; uppermost
bracteal leaves bright red (rarely pink, white or yellow), lanceolate, forming a rosette,
borne at the tips of long arching stems; flowers, monoecious, inconspicuous, greenish,
petal-less, in rather large cyathia; involucre, 8 to 10 mm high, green, with reddish 5-
laciniate lobes, with a large yellowish gland about 4 mm long; styles, bifid, red; fruit,
about 1.5 cm long, a long-stipulate capsule, glabrous, nodding; seeds, about 6 mm
across, subglobose, pale, smooth. Planted ornamental. 6.

Euphorbia rubicunda BI. "thyme-leafed spurge", "dwarf spurge"
te tarai (K)
syn. E. thymifolia sensu auct. non L.

Recent introduction. India. Occasional. Small prostrate, hispid, annual herb, with
pinkish stems and white latex; leaves, up to 1 cm long, opposite, obliquely oblong
obtuse, crenulate, glabrous or puberulent, coppery; stipules, less than 1 mm long,
elongate fimbriate; flowers, monoecious, minute, petal-less, borne in cyathia arranged in
short, few-flowered axillary clusters (cymes) on short pedicels; involucre enclosing
several male and | female flowers, gland small or none; fruit, about 1 mm long, very
shortly-stalked, white-pubescent, 3-celled capsule; seeds, about 0.7 mm long, 1 per cell,
blunt, light-brown, 4-angled, with 5 or 6 shallow transverse furrows. Weed along
roadsides and around buildings. 3(58824, 58661), 6, 7, 8(9561).

Euphorbia tirucalli L. "pencil plant", "pencil tree", "naked lady", "milk hedge",
"milk bush"

Recent introduction. E. Africa and India. Occasional. Medium to large, freely
branching shrub, up to 4 m or higher, with numerous slender, cylindrical, leafless, green
articulated branches; leaves, 0.7 to 2.5 cm and up to 0.5 cm wide long or less, narrow,
reduced to linear scales on youngest branch tips, early caducous; flowers, inconspicuous,

132

borne in cyathia in sessile, axillary and terminal, clusters (cymes); involucre, with 5
glands (nectaries); fruit, a 3-valved capsule; seeds, smooth, ovoid. Planted ornamental. 5,
6(168).

Jatropha integerrima Jacq. "rose-colored jatropha", "red-flowered jatropha"
syns. J. hastata Jacq.; J. panduraefolia Andr.

Recent introduction. Cuba. Common. Slender shrub up to 1 to 3 m high, with
little or no watery latex; leaves, 6 to 16 cm long and 4 to 10 cm wide, alternate and
spirally arranged, ovate to fiddle-shaped (panduriform), acuminate, mostly entire with
obscure basal teeth or marginally incised or toothed;; petioles, 3 to 10 cm long; stipules,
minute, subulate-deltoid, entire; flowers, monoecious, 5-petaled, dissected, attractive rich
pink to bright red, petals about 10 to 13 mm long, borne in terminal or axillary umbels
with long, branched stalks, about 6 to 14 cm long; male flowers, distal, usually with a
dissected disc, stamens, 8 to 10, anthers yellow to salmon-orange; female flowers, lower,
with cupular or pulviniform disk, ovary 3-locular; fruit, about 1 cm long, shallowly
lobed, breaking into 3 segments; seeds, ellipsoid, with crustaceous testa. Planted
ornamental. 3(58796), 5, 6(276), 7.

Jatropha podagrica Hook. "gout stalk", "coral plant"

Recent introduction. Central America. Rare. Succulent, un- or little-branched
perennial shrub, up to | m or higher, with a swollen stem, much swollen at the base, and
white latex; leaves, 10 to 30 cm in diameter, peltate to broadly ovate, entire to palmately
3- or 5- angled or -lobed, whitish beneath, sinuses rounded, lobes broad and acute,
attached within the margin by long petioles; petioles, 5 to 12 cm long; stipules, divided
into small rigid segments; flowers, about 12 mm across, orange-red to vermilion, borne
in dense umbels on long peduncles; calyx, eglandular, orange-red, 5-petaled, petals 5 to
8 mm long, red to vermilion; pistil, prominent, about 4 mm long, light orange-yellow
(ovary) to yellowish-green (stigma and style); fruits, about 1.5 cm long, capsular, broad-
ellipsoid, breaking into 3 segments; seeds, ellipsoid, with crustaceous testa, poisonous.
Ornamental pot plant. 6.

Manihot esculenta Crantz "cassava", "manioc", "tapioca"
te tabioka (K); tapioka, kasava (T)
syns. M. utilissima Pohl; M. manihot (L.) Karst.; Jatropha manihot L.

Pre-World War II introduction. Brazil or Trop. S. America. Common. An erect,
glabrous, slightly glaucous, half-woody, branched or unbranched shrub, up to 3 m or
higher, with milky latex, stems bearing prominent nubby leaf scars, and large edible
tuberous roots; leaves, 10 to 30 cm across, alternate, spirally arranged, somewhat
pendulous, palmately 3- to 9-lobed (usually 5- to 7-lobed), the lobes, 4 to 22 cm long and

133

1 to 6 cm wide, very slenderly lanceolate-elliptic, or obovate-lanceolate, acuminate,
deeply separated, dark to light green and glabrous above, glaucous and pale beneath,
sometimes red-tinged, sometimes slightly hairy beneath, midrib green to deep red;
petioles, 5 to 32 cm long, green to deep red; stipules, usually with 3 to 5 lanceolate lobes
to 1 cm long, deciduous; flowers, about 1 to 2 cm across, monoecious, petal-less, often
not developing, pendulous, borne in few-flowered, lax, axillary paniculate clusters, 3 to
10 cm long, with male and female flowers in the same inflorescence, the latter near the
base; bracts, linear, deciduous; sepals 5, pale yellow or tinged with red, glabrous
without, puberulent within; male flowers, on pedicels 0.5 to 1 cm long, calyx, 3 to 8 mm
long, campanulate with 5 triangular lobes extending to middle of calyx, stamens 10 in 2
whorls, alternately short and long, filaments free, anthers small, basal disk orange, fleshy
with 5 double lobes; female flowers, on pedicels 1 to 2.5 cm long, usually larger than
males, calyx 5-partite to base, 1 cm long or more, ovary, 3 to 4 mm long, 3-carpellary,
6-ridged, glabrous, greenish-white with pink or red streaks, on a 10-lobed disc, style
connate, surmounted by 3 lobed white stigmas, each lobe much divided; fruit, 1.5 to 2
cm long, subglobose to ellipsoid capsule with narrow longitudinal wings or ridges, 3-
seeded, splitting explosively; seeds, about 12 mm long, ellipsoid, smooth, gray mottled
with dark blotches, carunculate, with a thin crustaceous testa; tubers, developing as
swellings on adventitious roots at a short distance from the stem, usually 4 to 10 per
plant, 10 to 100 cm long and 3 to 15 cm in diameter, usually weighing 1 to 10 kg or
more, cylindrical or tapering, occasionally branched, outer skin, rough or smooth, light
to darker brown to white or pinkish, inner rind or cortex usually white, but sometimes
tinged with pink or brown, core or pith consisting of rich white or yellowish starch with
few xylem bundles and latex tubes. Food plant in I Kiribati and Tuvaluan gardens at
Location and Topside workshops. This easy-to-grow plant, which thrives on the poorest
of soils, has become the dominant staple root crop, displacing taro and yam as the
dominant staple in many areas of high-island Micronesia, and in Fiji, Tonga, parts of the
Cook Islands, and in the drier areas of Vanuatu, New Caledonia, and Papua New
Guinea. 5, 6, 7.

Pedilanthus tithymaloides (L.) Poit. "slipper flower", "slipper spurge", "shoe spurge",
"zigzag plant", "redbird cactus", "ribbon cactus"
syns. P. carinatus Spreng.; Euphorbia tithymaloides L.

Recent introduction. Caribbean. Occasional. Erect or somewhat decumbent,
sparsely branching, slightly succulent subshrub, up to | m or higher, with white latex and
fleshy, cylindrical stems which zigzag from node to node; leaves, 5 to 10 cm long,
alternate, soon falling, fleshy, waxy, ovate to lanceolate, subsessile, often variegated or
streaked green, white, yellowish or pink tinted, or in some forms completely albino;
flowers, monoecious, inconspicuous, petal-less, yellowish, borne in cyathia in axillary or
terminal cymose clusters; cyathium, about 12 mm long, narrow, boat- or slipper-shaped
red or pink bracts containing 1 female and several male flowers, with 4 glands; fruit, 7.5
by 9 mm, capsular; seeds, 5 mm long, ovate. Planted ornamental. 3(58681), 5, 6, 7.

134

Phyllanthus amarus Sch. & Th. "sleeping plant"
te kaimatu (K) :
syn. P. niruri L. sensu auct. plur. non L.

Recent introduction. Trop. America (despite African type locality). Common.
Erect to semi-prostrate, diffusely branched annual herb, somewhat woody at the base, 10
to 50 cm high; leaves, 4 to 10 mm long and 2 to 4 mm wide, numerous, crowded,
simple but appearing pinnately compound, alternate, in two rows on opposite sides of the
branchlets, oblong-elliptic or squarish, tip and base rounded, entire, green, lighter
beneath, glabrous, subsessile; stipules, about 1.5 mm long, narrow-triangular; flowers,
monoecious, minute, petal-less, greenish, borne in axillary cymules, on the branchlets,
under the leaves, and containing 1 male and 1 female flower per node; calyx lobes 5,
acute; pedicels 2 mm long; male flowers, with 3 stamens; female flowers, with a 3-celled
ovary and 3 bifid styles; fruit, about 1.5 mm wide, solitary, capsular, 3-celled, depressed
globose, glabrous, greenish to yellow-brown, borne on a stalk about 1.5 mm long,
splitting at maturity into three 2-seeded segments; seeds, about 1.5 mm long, wedge-
shaped, rounded dorsally, light brown, longitudinally 5- to 7-ribbed on back. Weed of
gardens and waste places. 3(58685, 58733), 4(155N), 5(47), 6, 7, 8(9571).

Phyllanthus societatis Muell. Arg.
eoemangemang, ewemangemang, eoemangmang (N)

Indigenous. Polynesia and Micronesia. Common. Erect, or half-erect, branching
shrub, up to 50 cm or taller; leaves, 0.6 to 2.5 cm long, alternate, simple but appearing
pinnately compound, alternate, in two rows on opposite sides of the branchlets, sessile,
mucronate, cuneate; flowers, monoecious, minute, petal-less, greenish, borne in axillary
cymules; male flowers, numerous, slender stalked, sepals 6, oblong-ovate, having 6
alternate glands at base, stamens 3, filaments united; female flowers, mostly solitary,
sepals 6, waxy white, ovary 3-lobed subtended by an entire glandular disc; fruit, 6 to 7
mm in diameter, depressed, splitting at maturity into three 2-seeded sections. Found as
scattered individuals and small communities in unmined areas on plateau, on slopes
around plateau, at base of escarpment and on coastal strip, around cemeteries, and
occasionally in revegetated mined areas and as a pioneer plant on limestone pinnacles on
the coastal strip. Straight main stems used as toy spears by children. 3(58595, 58748,
58801), 5(48, 51), 6, 7(22317), 8(9590).

Ricinus communis L. "castor bean", castor oil plant"

Recent introduction. Africa. Rare. Coarse, erect, branching, glabrous, semi-
woody shrub, up to 3 m or higher, with spreading, hollow greenish or reddish stems and
branches with well-marked nodes and leaf scars, watery sap, and a well-developed
taproot; leaves, 10 cm to 75 cm in diameter, alternate, spirally arranged, round-ovate in
outline, peltate, mostly deeply palmately 5- to 11-lobed, lobes sharply acuminate or

135

acute, serrate, palmately nerved, green to somewhat purplish when young, paler beneath;
petiole, 8 to 50 cm long, with 2 glands at apex; flowers, monoecious, small, petal-less in
large terminal (appearing axillary) paniculate clusters (cymules), 10 to 35 cm long, with
male flowers at the base and female flowers on the top; male flowers, with 3 to 5 calyx
lobes, very numerous stamens variously connate in branching clusters; female flowers,
with caducous, spathaceous calyx, ovary 3-celled, styles spreading usually bifid; fruit, 1
to 2 cm long, ovoid, capsular, softly spiny, green to purplish-brown, with three 1-seeded
lobes, dehiscent; seeds, 0.5 to 1.5 cm long, oblong-ellipsoid, compressed dorsally,
smooth, mottled with brown, black and grey or white, very oily. Weed of roadsides and
waste places. 3, 4(137N), 5(145), 6, 7, 8.

Synadenium cupulare (Boiss.) Wheeler

Recent introduction. Trop. Africa. Rare. Large, succulent shrub, | to 1.6 m high,
with thick cylindrical, thornless branches; leaves, 5 to 10 cm long, alternate, inverted
ovate or obovate, thick, cuneate, taprring to a short petiole; flowers, small, red, borne in
cyathia in axillary and terminal umbels, with a greenish-yellow gland. Pot plant at Cliff
Lodge. 5, 6.

FABACEAE OR LEGUMINOSAE (Bean, Pea or Legume Family)

Acacia farnesiana (L.) Willd. "sweet acacia", "West Indian blackhorn", "cassie flower",
"klu" (Hawaii) katin, debena (B)(N); te bakoa, te kai bakoa (K)
syns. Mimosa farnesiana L.; Vachellia farnesiana (L.) W. & A.

Pre-World War II introduction. Trop America. Occasional. Erect, much-branching
shrub, up to 3 m or higher, the trunk and branches bearing numerous straight, slender
stipular spines, 4 to 30 mm long; leaves, dark green, finely divided, bipinnate, with 2 to
6 (rarely 8) pairs of compound leaflets (pinnae), 1 to 4 cm long, each with 8 to 25 pairs
of small leaflets, 3 to 6 mm long and 1 to 1.5 mm wide; flowers in pedunculate axillary
heads, 1 to 3 heads together, peduncles, 1 to 3 cm long; flower heads, about 1.3 cm
across, subglobose, yellow to yellow-orange, fragrant, stamens numerous; fruit, a pod, 5
to 8 cm long and 8 to 15 mm in diameter, plump, cylindrical or subterete, curved, dark
brown to black, indehiscent; pulp, sweetish; seeds, compressed, elliptic, brown. Found in
waste places and on roadsides. Glue made from seed pod. 2, 3(58640), 4(159N), 5, 6, 7,
8(9568).

Acacia sp. "acacia"

Recent introduction. Australia? Rare. Planted ornamental tree. 3(58699).

136

Adenanthera pavonina L. "red-bead tree", "false wiliwili" (Hawaii)
bin ("bean")(N); lopa (T)

Pre-World War II introduction? Malaysia. Common. Medium-sized, deciduous
tree, up to 7 m or higher; leaves, bipinnate, with 2 to 8 pairs of compound leaflets
(pinnae), each with 6 to 12 pairs of alternate leaflets, 2 to 5 cm long and 1 to 3 cm wide,
ovate, obtuse, glabrous, rather pale green, thin; petiolules about 2 mm long; flowers,
greenish-white to yellowish, sweet-fragrant, short-pedicellate, borne in slender, elongate
axillary or aggregated terminal pedunculate racemose clusters, up to 15 cm or more long;
calyx, campanulate, 4- to 5-toothed; stamens 8 to 10; fruit, a pod, up to 25 cm or longer
and 2 cm wide, slender, flattened, linear at first, becoming falcately curved or contorted,
brown, smooth and shiny inside, 2-valved, dehiscent, becoming spirally twisted after
dehiscence; seeds, about 8 mm across, 10 to 12 per pod, lenticular, suborbicular, hard,
very ornamental, dark or bright red. Spontaneous tree in escarpment and Buada forests
and in older strip-mined areas. Seeds used in necklaces and eaten by children. 3(58804),
5(142), 6, 7.

Alysicarpus vaginalis (L.) DC. “alysicarpus", "one-leaved clover"
syns. Hedysarum vaginale L; A. nummularifolius (L.) DC.

Pre-World War II introduction. Paleotropics. Common. More or less prostrate,
somewhat suffruticose hispid branching, spreading herb, with jointed stems up to 1 m or
more long; leaves, 0.3 to 5 cm long and 0.2 to 3 cm wide, alternate, somewhat dimor-
phic, the proximal elliptic-subobovate, the distal narrower, lanceolate, rounded at the
tips, obtuse or truncate at base; petioles, 1 to 8 mm long, slender; stipules, prominent;
flowers, about 6 mm long, pinkish to reddish purple, short-pedicellate, borne in rather
dense short terminal racemose clusters, up to 8 cm long, 6 to 12 flowers to a cluster;
fruit, pods, 1 to 2.5 cm long, crowded, 5 to 7 seeded, cylindric, rugose, jointed,
indehiscent, but breaking at the joints between seeds at maturity; seeds, about 1.5 mm
long, oblong-oval, brown to pale brown or yellowish. Weed in open or semi-open places,
along jeep tracks in unmined forest, and a pioneer plant in recently mined areas. 2,
3(58744, 58763), 4(115N), 5(88), 6, 7.

Bauhinia monandra Kurz “pink bauhinia", "orchid tree", "pink butterfly tree",
"St. Thomas tree"

Recent introduction. Burma. Occasional. Shrub or small tree, up to 5 m or higher,
with a rounded crown, smooth gray bark and brown-hairy young growth; leaves, 7 to 20
cm long and wide, alternate, ovate-oblong in outline, cordate to rounded at base,
palmately 3- to many-nerved, pubescent beneath when young, deeply divided at apex into
2 wing- or butterfly-like lobes which extend about a fifth to half way to the base, lobes
obtuse to subacuminate; calyx, spathaceous or S-lobed; flowers, orchid-like, up to 10 cm

137;

across, several together in terminal or axillary racemes; petals 5, erect or spreading (4
obovate, 1 ovate), each about 3 to 5 cm long and 2 to 3 cm across, white to pink with
red or purple blotches or dots, the uppermost one with deeper red, yellow or yellow
margined; fertile stamen 1, staminodes 5, small; pods, 15 to 18 cm long and about 3 cm
wide, oblong or linear, flattened, leathery, rather shiny brownish or black; seeds, ovate,
compressed, dull brown, free or separated by tissue or septa. Planted ornamental.
3(58785), 5, 6(224), 7.

" "

Bauhinia variegata L “white bauhinia", "orchid tree", "butterfly tree", "purple
orchid tree", "mountain ebony"

Recent introduction. India, Burma and China. Rare. Shrub or small tree, up to 8
m high; leaves, 5 to 14 cm long and wide, alternate, broadly ovate to suborbicular,
cordate to truncate at based, palmately-nerved, split at apex into 2 wing-like lobes which
extend about a third of the way or less to the base, lobes rounded; flowers, several
together in terminal or axillary racemes; petals 5, obovate, 4 to 6 cm long and 2 to 3 cm
across, the uppermost one broader, variable from pale purple to rose or white or yellow,
with one petal variegated with red and yellow; fertile stamens 5, staminodes 5, about half
as long as the stamens; pods, up to 30 cm long and 2.5 cm wide. The tree is variable
with the two main varieties being B. variegata var. variegata, with pale purple to rose
petals, with purple or crimson veins or blotches, and var. candida with white petals with
green veins, and somewhat purplish on the exterior. Planted ornamental tree. 6(194), 7.

Caesalpinia bonduc Roxb. "peach nicker", "gray nicker", "nicker bean"
dugienae, dogienae (N)

Indigenous. Pantropical. Rare. Scrambling to climbing shrub with long branches
with sharp, recurved, hooked spines, young parts brown pubescent; leaves, 30 to 80 cm
long, bipinnate with 6 to 12 pairs of pinnae; stipules pinnate, subpersistent, leafy,
composed of 2 or 3 superimposed parts, caducous; leaflets (pinnae), 1.5 to 6.5 cm long
and 0.5 to 3 cm wide, opposite or subopposite, ovate to elliptic-oblong, inequilaterally
rounded or cuneate at base, obtuse to subacute at apex, slightly pubescent; petiolules, 2
to 3 mm long; flowers, about 1 cm long, numerous, in supra-axillary racemose clusters,
calyx greenish brown, corolla yellow, pedicels at antithesis 6 to 12 mm long; pods, 4.5
to 9 cm long and 3 to 5 cm wide, oblong, inflated, prickly, dehiscent; seeds 1 or 2,
about 1 to 1.8 cm in diameter, subglobose, smooth, glossy gray or olive green, very
hard. Large shrub in coastal thicket and in overgrown garden on Command Ridge.
5(114), 6(158, 212, 269).

Caesalpinia pulcherrima (L.) Sw. "pride of Barbados", "dwarf poinciana",
"Barbados flower fence"
syn. Poinciana pulcherrima L.

138

Recent introduction. Trop. America. Occasional. Shrub, up to 2 to 3 m tall, with
branches bearing short, stiff, scattered prickles; leaves, up to 30 cm long, alternate,
bipinnate, pinnae 4 to 8 pairs, each with 5 to 13 pairs of leaflets; leaflets, about 2 to 2.5
cm long and up to 1 cm wide, light green, elliptic, obtuse or with shallow notch at apex,
obliquely inequilateral; flowers, on long pedicels in long terminal racemes, petals 5, 15
to 25 mm long, spreading, orbicular or oblong, crinkly-edged, usually long-clawed,
scarlet to orange-red or red-and-yellow, usually with yellow wavy margins, or all yellow
(var. flava); filaments, 5 to 7.5 cm long, mostly scarlet, longer than petals; pedicels, 2.5
to 10 cm long; pods, up to 10 cm long and 2 cm wide, coriaceous, oblong, smooth,
brown to black, several-seeded, dehiscent; seeds, 6 to 8 per pod, brown, compressed,
Planted ornamental. 3(58789), 5(38), 6, 7.

Cajanus cajan (Mill.) Millsp. “pigeon pea", “red gram", arhar dhal (Hindi)
syns. C. indicus Spreng.; C. flavus DC.; Cystisus cajan L.

Pre-World War II introduction. India and southeast Asia. Said to have been
introduced in 1935, but not seen in 1978 or thereafter, but possibly re-introduced since
then by expatriate Indian contract employees. Erect, short-lived, perennial pubescent
shrub, 1 to 3 m or higher, with prominently ribbed stems and a pronounced taproot,
sometimes grown as an annual; leaves, spirally arranged, pinnately trifoliate; leaflets, 2.5
to 12 cm long and 1.5 to 4 cm wide, elliptic to lanceolate, acute, dotted beneath with
resin glands, grayish-green pubescent, soon glabrate above, finely silvery-pilose beneath,
longer-stalked terminal leaflet usually larger than short-stalked laterals; petiole, grooved,
about 2 to 8 cm long; stipules, ovate, hairy, about 4 mm long; flowers, about 2.5 cm
long, in long-pedunculate terminal panicles, also axillary and subcapitate-racemose,
subequal to the leaves in length; calyx 4-lobed, two upper lobes united (connate); corolla,
12 to 17 mm in diameter, petals, usually suborbicular, reflexed auriculate at base, bright
yellow, marked with dark reddish-brown to crimson lines within and often flushed with
brown to red without, wings yellow, keel yellow-green; pods, 4 to 10 cm long and 6 to
15 mm _ wide, linear-oblong, flattened, inflated, pubescent-glandular, pointed, with
diagonal depressions between the seeds, often streaked with purplish black to reddish
brown, tardily dehiscent; seeds, variable in shape and color, 2 to 8, up to 8 mm in
diameter, globose, compressed, cream-colored to reddish or brownish or speckled with
small white hilum, edible. Cultivated food plant.

Calopogonium mucunoides Desv. "calopogonium", "calopo"

Recent introduction. Trop. America. Rare. Slender creeping or twining perennial
herbaceous vine, coarsely brown-tawny pubescent; leaves pinnately trifoliate; leaflets,
petiolate, nearly equal in size, about 3 to 7 cm long and 2 to 6 cm wide, ovate to
rhomboid, obtuse to apiculate-subacute at apex, densely pubescent; flowers, 7 to 10 mm
long, axillary, short- to long-pedunculate, sessile or nearly so, medium to pale blue or
slightly purplish, sometimes with proximal yellow blotches; pods 2 to 4 cm long and 3 to

139

5 mm wide, compressed-convex, slightly constricted between the seeds, coarsely
pubescent; seeds 4 to 8, cube-like, flattened, about 3.5 by 3 by 2 mm, pale brown.
Locally abundant weed in low ground around Buada Lagoon. Probably originally
introduced as a green manure and nitrogen-fixing plant, but now naturalized. 3(58651),
6(153).

Canavalia cathartica Thouars "Mauna Loa bean" (Hawaii)
erekogo, irekogo (N); te kitoko (K); saketa (T)
syns. C. microcarpa (DC.) Piper; C. turgida Graham (of Burges list)

Indigenous. Pantropical. Occasional. Rather coarse creeping and high climbing
herbaceous vine; leaves, about 20 cm long, alternate, trifoliate; leaflets, 8 to 15 cm long
and 7 to 10 cm wide, ovate to acuminate, base rounded, obtuse, papery, whitish
puberulent; flowers, 3 to 4 cm long, fragrant, short-pedicellate, few, borne on rather
long, axillary peduncles up to 20 cm long, pedicels about 2 mm long; calyx, 5-lobed,
about 14 mm long, white-puberulent, upper lip much shorter than calyx tube; corolla 5-
petaled, pink to lavender, the wings and keel petals somewhat paler than the others;
pods, 10 to 12 cm long and 3 to 4.5 cm wide, thick, inflated, indehiscent or tardily
dehiscent, 3-angled on one edge, pointed, tan or brown; seeds, about 6, about 1.8 cm
long, ovoid, slightly flattened, brown to dark reddish-brown, hilum 9 to 14 mm long.
Climbing on trees in escarpment forest of Anibare. 2(49.5), 3(58737, 58800), 5(135), 6,
Us

Canavalia ensiformis (L.) DC. "Jack bean", "horse bean"
syn. Dolichos ensiformis L

Pre-World War II introduction. W. Indies. Reported by Schumann (1898) as
collected by Finch, but not seen in 1978. Bushy erect annual herb, 1 to 2 m high, the tips
of branches tending to twine; leaves trifoliate; leaflets, 6 to 15 cm long and 5 to 10 cm
wide, ovate-elliptic, obtuse or shortly acute, shortly white puberulent especially on
petioles and petiolules; petioles, stout, grooved above, usually longer than leaflets,
terminal leaflet long-stalked, lateral leaflets short-stalked with unequilateral base; flowers,
2 to 2.8 cm long, short-pedicellate, borne on axillary peduncles of 10 to 5O flowers in
groups of 3 to 5 on swollen pedicelar glands, pedicels 2 mm long; calyx, 14 mm long, 5-
lobed, with 2 upper and 3 smaller lower lobes, puberulent, the upper lip only slightly
shorter than the calyx tube; corolla 5-petaled, rose to violet; pods, 20 to 35 cm long and
2 to 3.5 cm wide, stout, somewhat compressed, pale tan or straw-colored when ripe,
pendant, spirally dehiscent, ribbed near upper suture; seeds, 8 to 20, about 2 cm long
and 1.3 cm wide, oblong, somewhat compressed, ivory white with brown scar, hilum
about 9 mm long, pale brown with orange margin.

140

Canavalia rosea (Sw.) DC. "sea bean", "bay bean"
erekogo (N); te kitoko (K) ;
syns. C. maritima (Aubl.) Thouars; C. obtusifolia (Lam.) DC.

Indigenous. Pantropical. Rare. Prostrate, rather coarse trailing or weakly climbing
glabrous or slightly pubescent herbaceous vine; leaves, alternate, trifoliate; leaflets, 6 to
12 cm and nearly as wide, fleshy-coriaceous, elliptic to suborbicular, subovate or
subobovate, obtuse or slightly emarginate-notched at apex, base rounded, obtuse to acute,
with pubescent appressed hairs; flowers, 2 cm to 3 cm long, short-pedicellate, in long-
pedunculate axillary few-flowered racemes, up to 20 cm long, slightly shorter than the
leaves; pedicel about 3 mm long; calyx 5-lobed, about 12 mm long, whitish-puberulent,
upper lip much shorter than tube, lowest tooth 2 mm long; corolla, 5-petaled, about 3 cm
long (standard), pink to purple, commonly with white blotch or yellowish proximally, the
wings and keel petals pale magenta; pods, 8 to 15 cm long and 2 to 3 cm wide, broadly
linear, inflated or only slightly compressed, spirally dehiscent, pale tan; seeds, about 1.8
cm long, elliptic, slightly compressed, brown with dark marbling, hilum about 7 mm
long. Found in beach vegetation. 7(27804).

Cassia alata L. "golden candelabra bush", "Roman candle tree", "candle bush",
"ringworm bush"
syn. Senna alata (L.) Roxb.

Recent introduction. Mexico. Rare. Coarse erect shrub or small tree, 2 to 5 m
tall, with downy green branches and strong-smelling foliage and bark; leaves, 50 to 80
cm long, with 5 to 13 pairs of large leaflets; leaflets, 5 to 18 cm long and 3 to 9 cm
wide, the distal ones largest, ovate-oblong, obtuse, truncate, or even slightly notched at
apex, subsessile; stipules, obliquely deltoid, 6 to 16 by 3 to 10 mm; inflorescence, a
striking long-pedunculate erect dense cone-like racemose spike, 15 to 60 cm long,
bearing many crowded and overlapping flowers; calyx, 5-parted, sepals up to 16 mm
long, yellow or orange; petals 5, 16 to 23 mm long, bright yellow, subtended by yellow
or yellow-orange floral bracts; pods, 12 to 19 cm long and 2 to 3 cm wide, straight,
widely ascending, sharply tetragonal, papery or leathery in texture, winged lengthwise
down the middle of each valve; seeds, numerous, up to 60, flat. Planted ornamental;
leaves widely used as a cure for ringworm, but reportedly not in Nauru. 5, 6.

Cassia fistula L. “golden shower tree", "Indian laburnum", "pudding-pipe tree"
te katia? (K)

Recent introduction. Trop. Asia. Occasional. Small to medium tree, up to 10 to
15 m or higher, with a short bole and spreading crown; leaves, with 3 to 8 pairs of
leaflets; leaflets, 8 to 21 cm long and 5 to 9 cm wide, subsymmetrically ovate or elliptic,
acute to subacuminate; inflorescence, 15 to 65 cm long, a drooping many (15 to 75)-
flowered axillary raceme, resembling in size and shape a large bunch of grapes, often

141

occurring 2 or 3 together; pedicels 3 to 6 cm long; calyx 5-sepaled; corolla, 4 to 6.5 cm
across, 5-petaled, petals 1.6 to 3.2 cm long, bright golden yellow, subequal, shortly
clawed; stamens 10, strongly accrescent (crescent-shaped) toward the abaxial side of the
flower, the filaments of 3 long abaxial stamens sigmoidally arcuate, gradually and
slightly thickened in the middle, and much longer than the anthers; pods, 30 to 60 cm
long and 2 cm or less thick, cylindrical or narrowly terete, straight, smooth, hard-walled,
dark brown, with seeds embedded in sweet, sticky, blackish pulp; seeds, many, up to
100, brownish. Planted ornamental. 3(58529, 58788), 5, 6, 7.

Cassia glauca Lam. "scrambled-egg tree?"
syns. C. sulfurea DC. ex Colladon; Senna sulfurea (DC. ex Colladon)
Irwin & Barneby

Recent introduction. India. Rare. Small tree, 4 to 6 m high; leaves, up to 30 cm
long with 4 to 7 pairs of leaflets; leaflets, up to 8.5 cm long and 3.8 cm wide, usually
elliptic, paler beneath; inflorescence, a 7 to 15-flowered axillary raceme; pedicels, 2 to 4
cm long; calyx 5-sepaled, innermost sepal 8 to 11.5 mm long; corolla 5-petaled, petals,
up to 23 to 30 mm long, ovate to oblong-obovate, bright yellow to orange-yellow; pod,
10 to 17 cm long and 1.3 to 1.8 cm wide, plano-compressed, brown; seeds, transverse,
compressed parallel to valves. Planted ornamental. 6(170, 175), 7.

Cassia grandis L.f. "pink shower tree", “horse cassia"
te kasia? (K)

Recent introduction. Central America. Rare. Medium-sized tree up to 10 m or
higher; leaves with 8 to 17 pairs of leaflets; largest ones 3.5 to 6.5 cm long and 1.2 to
2.5 cm wide, oblong, obtuse at apex; inflorescence, 8 to 27 cm long, a 20 to 45-flowered
raceme, axillary or borne on branches, becoming obliquely geotropic from drooping
branches; pedicels, 8 to 20 mm long subtended by bracts; bracts, 2 to 5 mm long, ovate,
caducous as pedicels begin to elongate; corolla, 5-petaled, longest petals 8.5 to 11 mm
long; pods, 40 to 90 cm long and 3.5 to 5 cm wide, massively linear-oblong, slightly
laterally compressed, keeled dorsally by 1 and ventrally by 2 parallel blunt ribs; seeds,
14 to 16 mm long and 9 to 10 mm wide. Planted in home garden at Buada. 7.

Cassia occidentalis L. ’coffee senna", "arsenic bean"
tan braua ("sunflower")(N); te katia? (K)
syn. Senna occidentalis (L.) Link

Pre-World War II introduction. Trop. America. Abundant. Erect, branching fetid
annual subshrub, up to 2 m high, with glabrous longitudinally grooved stems and often
woody near the base; leaves, 11 to 24 cm long, alternate, pinnately compound, with 4 to
6 pairs of opposite leaflets; leaflets, 3 to 12 cm long and 2 to 4 cm wide, the terminal

142

pair largest, elliptic or ovate-lanceolate, acute to acuminate, base inequalaterally rounded,
entire, glabrous; petiole with conspicuous gland near base, petiolules 1 to 2 mm long;
inflorescence, a short 1- to 5-flowered axillary or terminal raceme, each flower subtended
by an acute, caducous bract; calyx deeply divided into 5 pinkish or brown-tinged sepals
with rounded tips; corolla, 5-petaled, each petal about 9 to 16 mm long, obovate, yellow,
drying whitish and brown-veined; pods, 6 to 15 cm long and 5 to 9 mm wide, erect or
narrowly ascending, linear, flattened, glabrous, falcate, the margins somewhat thickened,
purplish in colour with green margins turning brown; seeds, 15 to 50, about 3 mm long,
ovate, brown, mostly with broad faces turned toward septa. Weed of waste places and
roadsides. Dried seeds boiled as a tea substitute during World War II, a use learned from
the Japanese; soft leaves cooked as a spinach and used medicinally by the Chinese. 2,
3(58529), 4(128N), 5(13), 6, 7, 8(9547).

Centrosema pubescens Benth. "centro"

Pre-World War II introduction. Trop. S. America. Extinct. Trailing or climbing
herb, with slender pilose stems, often forming tangled mats; leaves trifoliate, petiolate,
the terminal leaflet larger; leaflets, 3 to 9 cm long and 1.5 to 5 cm wide, ovate to
elliptic, apex mucronate, base rounded, short-pilose but glabrate on both sides; inflores-
cence, compact, axillary, several-flowered pseudo-raceme, each flower subtended by
bracteoles about 8 mm long and equal in length or smaller than calyx; flowers blue to
purple or mauve with darker veins and a white or yellowish patch down the middle of
largest petal (standard), the standard, up to 4 cm broad and 3 cm long, broadly orbicular,
rather reflexed with a spur scarcely | mm long, pubescent without, the keel petals nearly
as long as the wings; pods, up to 17 cm long and 7 mm wide, excluding persistent style,
compressed, margins thickened, with 4 prominent ribs or wings near the sutures,
dehiscent, the style often persistent as a beak; seeds, often about 20, 5 by 3 by 2 mm,
oblong or subglobose, compressed, olive-green with red markings to red-brown with
black streaks, hilum small. Reported in 1935 to have been introduced, but not seen in
1978 or thereafter. A widely used pasture legume and nitrogenous cover or green manure
crop.

Cicer arietinum L. "chick pea", “garbanzo bean", "Indian gram", "common gram",
"chana" (Hindi)

Pre-World War II introduction. W. Asia. Extinct. Erect or spreading, much-
branched, annual herb, 25 to 50 cm tall, downy with all parts covered with clavate
glandular hairs and with tetragonous stems and a well developed tap root; leaves, about
5 cm long, imparipinnate with 9 to 15 leaflets, yellowish-green to dark bluish-green;
stipules, about 8 mm long, ovate, notched; leaflets, 0.8 to 2 cm long and 0.5 to 1.5 cm
wide, opposite or alternate, ovate, elliptic or obovate, serrate; inflorescence, an axillary
jointed peduncle, bearing solitary flowers, 2.4 to 4 cm long, borne on pedicels 5 to 10
cm long and slightly longer than peduncles; calyx, united with 5 teeth; corolla, white,

143

greenish, pink, purplish red or blue, pink flowers fading to blue with age, standard
(largest petal), 1 to 1.5 cm long, broad and clawed, wings free, keel incurved; stamens
10; pods, 1.5 to 3 cm long and 1 to 2 cm across, 1- to 2-seeded, oblong, swollen, beak
obliquely placed, sometimes sterile; seeds, 0.5 by 1 cm in diameter, angular, oblong-
obovoid, with pointed beak and small hilum, testa smooth, wrinkled or rough, ranging in
color from white, yellow, red, brown to nearly black. Common food and fodder legume
reported in 1935 to have been introduced, but not seen in 1978 or thereafter.

Clitorea ternatea L. “butterfly pea"

Recent introduction. Trop. America or Pantropical. Fast growing perennial
climbing or sprawling herb with slender stems, up to 3 m long, and a woody rootstalk,
young growth somewhat pubescent; leaves, alternate, compound, odd-pinnate, with 5 to 9
leaflets, one terminal; leaflets, 2 to 7 cm long and 1.5 to 4 cm wide, oblong to elliptic,
obtuse, base acutish, apressed-pilose but subglabrate on both sides; petiolules, about 2
mm long; stipules, short linear; flowers, mostly solitary, sometimes paired in leaf axils
on pedicels about as long as petioles; calyx, about 1.5 to 2 cm long, longish-tubular
subtended by 2 conspicuous basal bracteoles and lobes about half as long as tube; petals
large, up to 4 by 5 cm, the standard the largest, other petals very small; standard, erect,
half-tubular, wavy-margined, bright vivid blue to violet with a pale yellow or white
proximal blotch, finely puberulent dorsally, wings much shorter, also blue with pale basal
parts; pods, 8 to 12 cm long awns 0.8 to 1 cm wide, linear-oblong, compressed, 2-
valved, sharp-beaked, thick-margined and slightly broad-margined distally, apressed-
pilose but subglabrate, dehiscent; seeds, 5 to 10, up to 8 by 4 by 2.5 mm, ellipsoid or
subglobose, compressed, greenish to pale or dark brown with darker mottling, hilum
small. Occasional. Planted ornamental. 3(58778), 5(71), 6(246), 7.

Crotalaria goreensis Guitl. & Pers. "rattlepod"

Recent introduction. W. Africa. Common. Erect subshrub, 50 cm to 1 m tall,
branching below; leaves, alternate, long-petiolate, trifoliate; petioles, 2 to 4 cm long;
stipules foliaceous, trifoliate, similar to, but smaller than leaflets, stipular leaflets, 3 to 8
mm long and | to 3 mm wide, stipular petiolules 3 to 8 mm long; leaflets, 1 to 4 cm
long and 4 to 8 mm wide, the terminal leaf longest, narrowly elliptic, acute, obovate or
slightly mucronate, base acute, light gray-green, glabrous; inflorescence, a leaf opposed
peduncle, 3 to 7 cm long, bearing solitary flowers, pedicels about 1 to 2 mm long; calyx
funnelform, about 2 mm long; corolla, yellow; pods, 1 to 1.8 cm long, 2-valved, oblong,
inflated, beaked, dehiscent; seeds, small, about 3 mm long, club- or kidney-shaped, dark
brown. Widespread weed of waste places, roadsides and other ruderal sites on coastal
strip and on plateau. 3(58593, 58612, 58738), 4(106N),5(3, 58), 6, 7(22311), 8(9569).

144

Crotalaria retusa L. "rattlepod"

Recent introduction. Trop. Asia. Rare. Coarse short-lived subshrub or stiff herb,
40 to 100 cm high, with pilose ribbed stems; leaves, 4 to 8 cm long and 1.5 to 2.5 cm
wide, simple, alternate, obovate-oblong to oblanceolate, rounded, emarginate or mucro-
nate, cuneate at base, shortly-stalked, finely pubescent, the hairs minute and closely
appressed; stipules, 1 to 5 mm long, subulate; petioles, about 3 mm long; inflorescence,
a dense terminal many-flowered raceme, up to 30 cm long; flowers, 2 to 3 cm long,
borne on short downward bent pedicels; calyx, 2-lipped, the upper deeply bifid, the
lower 3-toothed; petals bright yellow, the standard, about 2 cm high and 3 cm wide,
reflexed, yellow marked with purplish veins or blotches, the wings, about 1.5 cm long,
yellow, almost hiding the keel, the keel with a short twisted beak; pods, 3 to 5 cm long
and 1.3 to 1.8 cm wide, oblong-clavate, inflated, dehiscent, held horizontally, black
when dry; seeds, about 20, about 4 mm long, yellowish or golden-brown. Weed of
ruderal habitats and roadsides. 4(158N).

Crotalaria spectabilis Roth “rattlepod"
syn. C. sericea Retz.

Recent introduction. India and Paleotropics. Occasional. Erect robust, smooth-
stemmed subshrub, up to | m or higher, leaves, 5 to 13 cm long, obovate, oblanceolate
or sometimes spoonshaped, base acuminate or cuneate, shortly petiolate, glabrous above,
with soft silky hairs (villose) below; stipules sessile and persistent; inflorescence, a loose
peduncie, 30 cm or longer, bearing 20 or more flowers; flowers, 2.5 cm or longer,
bright yellow or purple?; floral bracts, leaflike, oval or cordate; calyx, about 12 mm
long, glabrous, cleft more than halfway to the base; pods. 2.5 to 5 cm long, 2-valved,
bladder-shaped, glabrous, containing a small number of seeds, which at maturity become
loose in the pod. Weed near Buada Lagoon, in gardens, and near coast in Nibok;
occasionally cultivated in home gardens. 3(58647), 4, 6, 7, 8(9579).

Cyamopsis tetragonoloba (L.) Taub. “cluster bean", "guar" (Hindi)
syns. C. psoraloides (Lam.) DC.; Psoralea tetragonoloba L.; Dolichos psora-
loides Lam.

Pre-World War II introduction. India. Extinct. Robust bushy annual, 1 to 3 m
high, with stiff, erect, angled, grooved, white-pilose branches; leaves, alternate,
trifoliate; leaflets, ovate, sparsely serrate; inflorescence, a dense axillary raceme bearing
6 to 30 solitary flowers in axils of caducous bracts; calyx, cupuliform, oblique, the
lowermost tooth the longest; petals 5, strongly veined, the standard, about 8 mm long,
obovate, not clawed, white, the wings, oblong free from the keel, pinkish, the keel petals
erect, slightly gibbous or short-calcarate; pods, 4 to 10 cm long, linear, compressed, with
a marked beak, a double ridge on the dorsal side and a single ridge on the ventral side,
septate between the seeds, dehiscent, borne in stiff erect clusters; seeds 5 to 12, about 5

145

mm long, compressed-oval, white to gray or blackish. Cultivated food plant, the young
tender pod which are cooked as a vegetable; reported in 1935 to have been introduced,
but not seen in 1978 or thereafter.

Delonix regia (Bojer) Raf. "poinciana", "royal poinciana", "flame tree", "flamboyant",
"flame of the forest" bin ("bean"), red tree (N); te tua (K); fuatausaga (T)

Pre-World War II introduction. Madagascar. Occasional. Rapidly growing,
medium-sized, spreading, broad-crowned deciduous tree, up to 10 to 12 m high and
several times as wide, with smooth, light-colored bark; leaves, mostly 20 to 60 cm long,
alternate, bipinnate, frond-like, with 10 to 20 pairs of pinnae; pinnae, about 10 cm long,
each with 25 to 35 pairs of leaflets; leaflets, 0.5 to 1 cm long, opposite, oblong-elliptic,
obtusish, base oblique, subsessile, glabrous, medium green; stipules, inconspicuous,
forked at base, the divisions pinnate, with 3 to 6 leaflets; inflorescence, an axillary
corymbose-raceme bearing large, showy flowers, 7.5 to 10 cm across, aggregated near
ends of branchlets; bracts, small, caducous, bracteoles none; calyx, short-tubed, sepals 5,
valvate, subequal, free at antithesis; petals 5, about 4 to 7 cm long, subequal in size,
brilliant crimson to scarlet or red-orange, conspicuously clawed, imbricate (overlapping),
the uppermost with whitish or yellowish streaks or mottling, broad-bladed and stalked
with their distal part abruptly expanded, orbicular with wavy edges; pods, 13 to 60 cm
long and 3 to 5 cm wide, linear-oblong, parallel-edged, flattened, shortly-beaked at tip,
woody or coriaceous, pendent, septate, dark reddish-brown turning black, filled within
between the seeds, many-seeded, persistent, dehiscent; seeds, oblong-cylindric, trans-
verse, gray with pale margins, testa hard. Planted ornamental; flowers used in garlands.
3(58620), 5(121), 6, 7.

Derris trifoliata Lour. "peach derris root", "beach poison vine"
syns. Derris uliginosa Benth.; Robinia uliginosa Willd.

Indigenous. Trop. Africa to Polynesia. Occasional. Glabrous creeping and
climbing shrub with prostrate rooting stems and rather large, scattered lenticels, climbing
to a height of 5 to 6 m; leaves, alternate, 3 to 7-foliate; leaflets, 4 to 12 cm long and 2 to
6 cm wide, ovate to elliptic-ovate, acute-acuminate or cuspidate at apex (acumen rounded
or emarginate at tip), base obtuse or rounded-subtruncate, subcoriacious, rather glossy
medium-green, slightly paler but not glaucous dorsally, the secondary nerves prominulous
beneath, petiolate; petioles, 4 to 8 mm long; inflorescences, axillary short-pedunculate
racemes bearing flowers on short slender pedicels; calyx, 2 to 3 mm long, pale green;
corolla, 6 to 8 mm long, petals, pale pink or greenish-white and faintly pink-tinged; the
filaments and styles white; pods, 3 to 5 cm long and 2 to 3.5 wide, subreniform or
subrhomboid to broadly oblong, flat, with a wing, 1 to 2 mm broad, along the upper
suture, greenish or tan, conspicuously veiny, 1- to 2-seeded; seeds, 1.5 to 2.4 cm long,
oblong-reniform. In forest on cliffs and steep slopes of escarpment surrounding the

146

central plateau. No reported use in Nauru, although the roots, which contain rotenone,
are often used for fish poison throughout the Pacific. 3(58803), 6. -

Desmodium tortuosum (Sw.) DC. "Florida beggarweed"
syns. D. purpureum (Mill.) Fawc. & Rendle; Hedysarum purpureum Mill.; H.
tortuosum Sw.

Recent introduction. W. Indies and Central America. Common. Erect subshrub or
annual, somewhat woody (ligneous), branching herb, up to about 2 m high, with slender
densely and finely pubescent longitudinally grooved stems; leaves, alternate, trifoliate;
petioles, often reddish; stipules, asymmetrical, distinctly auriculate at base on side away
from petiole (and less obviously on side toward petiole), acuminate from a broad base,
sometimes reflexed, persistent; leaflets, 2.5 to 13 cm long and 1.5 to 7 cm wide, the
terminal leaflet largest, narrowly elliptic to ovate, acute to mucronate, base acute to
rounded, laxly pilose with long hairs and/or short, uncinate hairs; inflorescences, simple
or branching racemes, 20 to 50 cm long; calyx, deeply 5-lobed, densely pubescent, borne
on a thin pedicel, 5 to 17 mm long, with many basally thickened hairs; corolla, 4 to 5
mm long, pea-like (papilionaceous), white or greenish-yellow to pink shading to mauve;
stamens 10, diadelphous; pods, 1.5 to 3 cm long, twisted, brown, segmented, each
segment (article) 3 to 6.5 mm long and 3 to 4 mm wide, the articles suborbicular, the
isthmi comparatively narrow, about one-fourth as broad as articles, 2- to 7-seeded, each
seed in a separate compartment which separates at maturity; seeds, compressed. Weed on
roadsides, on slopes in settled areas, and in waste places. 3(58639, 58741), 4(110N),
5(87), 6, 7(22308).

Desmodium triflorum (L.) DC. "tropical trefoil", "three-flowered beggarweed"
syn. Hedysarum triflorum L.

Recent introduction. Pantropical. Occasional. Diminutive prostrate freely-
branching perennial creeping herb with slender stems with appressed or weakly spreading
hairs, up to 1 mm long, and a woody rootstalk, commonly forming a thick mat; leaves
alternate, trifoliate, clover-shaped, lower leaves sometimes unifoliate; stipules sym-
metrical at base, lanceolate, acuminate; leaflets, 4 to 10 mm long and almost as wide,
terminal leaflet largest, sometimes bearing 2 white marks, obovate to elliptic-obovate or
obocordate, rounded (obtuse) and usually distinctly (but slightly) emarginate, base
narrowed to a short petiolule; inflorescence, a lax, 2- to 3-(rarely 5)-flowered fascicle
(occasionally solitary), up to 6 cm long, borne opposite the leaves, each fascicle
subtended by a soon caducous primary ovate-acuminate, not congested bract,; pedicels or
peduncles, 3 to 10 mm long, lengthening in fruit to over 1 cm, puberulent with straight
or weakly spreading hairs, 0.7 to 1 mm long, or glabrous; calyx, appearing subequally 5-
lobed, lobes shallowly connate at base for less than half their length; petals pale or rich
pink to reddish-violet, the standard obovate, 4 to 5 mm long; pods, 6 to 20 mm long and
about 2.3 mm broad, indented on the lower suture, segmented into 2 to 5 articles, each 2

147

to 4 mm long and broad, indehiscent, with only uncinate hairs, the isthmi two-thirds to
three-quarters as broad as the articles; seeds, bean-shaped (reniform). Weed in gardens,
lawns and in shaded ruderal habitats on plateau. Possibly deliberately introduced as a
green manure and cover crop. 3(58708, 58803), 4(126N), 5(148), 6, 7.

Dolichos lablab L. "hyacinth bean", lablab bean", "dolichos", "bovanist bean", "Egyptian
bean", "Indian bean", "bataw or batani bean (Philippines)
syns. D . purpureus L.; Lablab purpureus (L.) Sweet; L. niger
Medik.; L. vulgaris Savi; L. cultratus DC.

Recent introduction. Paleotropics. Rare. Vigorous perennial, glabrous or finely
pubescent twining or scrambling herbaceous vine, from 1.5 to 6 m long or high, all parts
often flushed with purple; leaves, alternate, trifoliate; petioles, 6 to 26 cm long; leaflets,
5 to 15 cm long and nearly as wide, entire, broadly triangular ovate, lateral leaflets
oblique, acute-acuminate, base obtuse, subglabrous or softly pubescent, 3-nerved from
base, midrib branched upward, the terminal leaf longer stalked; inflorescences, erect,
terminal and axillary peduncles, 5 to 40 cm long, bearing pseudoracemose clusters of 1
to 5 flowers together at nodes along the peduncle, each about 1.5 cm long, on slender
pedicels up to 1 cm long; bracts and bracteoles, 2 to 5 mm long, caducous; calyx, about
equal in length to the pedicels, 4-lobed, campanulate, bilabiate, the 2 upper lobes joined
into an entire or emarginate lip, the lower lip 3-lobed; petals, small, violet or whitish,
the standard up to 14 mm long and 20 mm across, orbicular, reflexed, auriculate at base,
the wings obliquely obovate, longer than the keels, the keel petals incurved at a right
angle, obtuse with a small convex pocket at base; stamens 10; pod, 6 to 12 cm long by 2
to 4 cm wide, 2-valved, obliquely oblong, flattened, often curved, beaked at tip with
persistent style; seeds, 3 to 6, usually not larger than 15 by 9 by 5 mm, ovoid, slightly
compressed, dark brown, reddish or pale tan or white, often speckled, the hilum linear
with whitish rim-aril. Food plant in Filipino gardens at Location; the young pods and
tender seeds cooked as a vegetable. Said to have been introduced in 1935 and established
by 1936. 5, 6.

Erythrina variegata L. var. variegata "variegated coral tree"

Recent introduction. Pacific Is. Rare. Differs from E. variegata var. orientalis
(see description below) in having narrower, more leathery leaves, which are mottled or
variegated with light yellow along the midribs and side veins, and larger lighter-colored
flowers in longer erect clusters. Planted ornamental. 3, 5(39), 6, 7.

Erythrina variegata var. orientalis (L.) Merr. "coral tree", "dadap"
yora, yoreh (N)
syns. E. indica Lam.; E. corallodendron var. orientalis (L.) Merr.

148

Indigenous. Indopacific. Occasional. Medium to large deciduous tree, up to 25 m
high, the trunk and branches usually with coarse spiny thorns; leaves, alternate, trifoliate,
membranous, rachis not including the petiole 12 to 40 cm long; petioles, 9 to 20 cm
long; leaflets, 6 to 30 cm long and nearly as wide, broadly triangular-ovate, rhombic or
cuspidate, acute-acuminate, rounded, acutish or truncate at base, lower leaflets somewhat
oblique, with prominent glands below the base of petiolules; petiolules, 5 to 14 mm long;
inflorescence, a dense axillary or terminal pedunculate raceme, up to 40 cm long, bearing
numerous flowers, about 6 cm long, clustered near the end of the raceme; calyx tube in
bud closed at orifice, early broken by the emergent corolla; petals, bright dark red or
scarlet, very unequal, the standard 5 to 8 by 2 to 3.5 cm, broadly elliptic, narrowed
proximally to a short claw, the keel petals separate, obovate, rounded at apex, nearly as
long as the wings, these obovate, rounded, about one-third as long as the standard; pod,
10 to 45 cm long and 2 to 3.5 cm in diameter, not or slightly constricted between the
seeds, the valves coarsely reticulate-veined, glabrous, black, tardily dehiscent; seeds, 1 to
12, 1.2 by 2.5 cm, ovoid, bright red to brownish red, the hilum oblong. Planted or
spontaneous on coastal strip and in old strip-mined land; a number of large specimens
along road near Nauru Phosphate Company tennis court area. Trunks and thick branches
used for living fence posts, and in canoe construction in the past; flowers used in
garlands in the past. 3(58737a), 5, 6, 7.

Gliricidia sepium (Jacq.) Kunth ex Walp. "gliricidia", "madre de cacao (mother of
cocoa)", "Nicaragua cocoa shade"
syn. Robinia sepium Jacq.

Recent introduction. Central and northern So. America. Small semi-deciduous
fast-growing tree, up to 10 m tall, with pale bark and ascending stems; leaves, 15 to 25
cm long, imparipinnate, with 5 to 17 leaflets; leaflets, 3 to 8.5 cm long and 1.5 to 5 cm
wide, opposite, ovate, ovate-oblong to elliptic-oblong, obtusely acuminate, often with
brown blotches or bronzy and glaucous beneath, dull green above, glabrate; inflorescen-
ces, stiff short racemes, 5 to 13 cm long, on the older branches after the leaves have
fallen, bearing single flowers, each about 1.8 to 2 cm long, on the rachis; bracts, small,
bracteoles none; pedicels, 8 to 12 mm long, jointed at tip; calyx, cupuliform, puberulent,
red-tinged; petals 5, rose or paler pink, the standard, large, suborbicular, reflexed,
sometimes with small inflexed auricles, claw very short, with a pale yellow central
blotch, the wings falcate-oblong, free, shorter than the keel, the keel petals incurved,
sometimes partly yellowish; stamens, 10, the filaments of 9 connate into the sheath;
pods, 8 to 16 cm long and 1.5 to 1.9 cm wide, 2-valved, linear-oblong, flattened, non-
partitioned, dehiscent, the valves coriaceous; seeds, 2 to 9, purplish-brown. Planted
roadside ornamental on Meneng Terrace. Widely used in tropical America and elsewhere
as a shadetree for tree for cocoa, bananas and coffee and as a living fence and wind-
break. 7(22320).

149

Hardenbergia violacea (Schneev.) F.C. Stern "sarsaparilla"

Recent introduction. Australia. Rare. A climbing or scrambling evergreen herb;
leaves, simple; inflorescence, clusters of violet flowers with yellow markings. Ornamen-
tal pot plant. 6.

Indigofera hirsuta L. "hirsute indigo"

Recent introduction. Africa and Madagascar to southern Asia and Australia.
Occasional. Erect or low spreading shrub, up to 1 m tall, stiffly and copiously brown-
hirsute with long hairs and woody at the base; leaves, imparipinnate with 5 to 7 leaflets;
leaflets, up to 4.5 by 2.5 cm, opposite, elliptic-obovate, densely pilose on both sides;
inflorescence, a dense raceme, 20 to 30 cm long, the peduncle, 2.5 to 8 cm long, both
peduncle and pedicels brown pubescent; calyx about 4 mm long, divided nearly to base
with linear-setaceous lobes, brown pubescent; corolla, small, with brick-red to salmon-
red or rose-colored petals; pods, 1.2 to 2 cm long and about 2 mm in diameter, straight,
cylindrical, somewhat tetragonal, with spreading dark brown hairs; seeds, 6 to 9, nearly
square or rectangular in outline. In plateau forest along path. 3(58614, 58730), 5(136),
8(9545).

Indigofera spicata Forssk. "creeping indigo"
syns. JI. hendecaphylla Jacq.; 1. endecaphylla Jacq. ex Lam. (sphalm?)

Recent introduction. Africa and Madagascar to Yemen, southeast Asia and
Australia. Occasional. Spreading or prostrate annual herb, with stems sometimes
ascending to 60 cm; leaves imparipinnate with 5 to 11 leaflets (usually 7); leaflets, 3 to
30 mm long, alternate or rarely opposite, elliptic to obovate, apex mucronate, shortly
petiolate; inflorescence, an axillary raceme, up to 15 cm long, the peduncle 1 to 4 cm
long, the pedicels, about 0.5 mm long; calyx, 2 to 3 mm long, divided nearly to the
base; corolla, 4 to 5 mm long, salmon pink; pods, 11 to 25 mm long and about 2 mm in
diameter, straight, cylindrical, appressed-strigulose or pubescent; seeds, 5 to 8. Roadside
weed on coastal strip and plateau. 4(123N), 6, 7.

Leucaena leucocephala (Lam.) de Wit "leucaena", "koa haole"(Hawaii), "lead tree",
"wild tamarind" bin ("bean")(N); te kai tetua (K)

syns. L. glauca (L. ex Willd.) Benth.; Mimosa leucocephala Lam.; Mimosa
glauca sensu L.; Acacia leucocephala (Lam.) Link

Pre-World War II introduction. Trop. America. Common. Erect slender shrub or
small tree, 1 to 5 m high, young growth puberulent; leaves, alternate, bipinnate, to 25
cm long, with 3 to 10 pairs of opposite pinnae, 5 to 10 cm long, each with 7 to 20 pairs

150

of leaflets; petioles, up to 10 cm or longer; leaflets, 6 to 19 mm long and 1.5 to 5 mm
wide, opposite, lanceolate to oblong-lanceolate, acute, asymmetrical, inequilateral at
base, sessile, somewhat dull grayish-green, glabrous; flowers, 1.5 to 3 cm in diameter, in
dense, axillary globose, head-like, pedunculate clusters; peduncles, up to 4 to 6 cm long;
calyx, 2.5 mm long, tubular, short-dentate with 5 small lobes at top; petals 5, 4 to 5 mm
long, linear, free, pale green or white; stamens, 10, nearly 1 cm long, long-exserted,
filaments white, anthers hairy, pale yellow; pods, 8 to 20 cm long and 1 to 2.2 cm wide,
clustered, 2 -valved, strap-shaped, linear, flat, beaked at apex, not septate, dark brown,
more or less dehiscent; seeds, 15 to 25, about 6 mm long, transverse, oval-oblong, flat,
glossy brown. Spontaneous in disturbed habitats on escarpment slopes below plateau and
in isolated stands on coastal strip. Used as firewood. 3(58638), 4(161N), 5(9), 6, 7,
8(9564).

Mimosa pudica L. var. tetrandra (HBK ex Willd.) DC. "sensitive plant"
syn. M. tetrandra HBK ex Willd.

Recent introduction. Trop. America. Rare. Decumbent or prostrate subwoody,
loosely branching perennial creeper, up to 50 cm high, with reddish-brown stems bearing
scattered curved prickles; leaves, alternate, bipinnate, with 1 to 3 pairs of pinnae,
palmately arranged at the end of the rachis, each with 12 to 25 pairs of leaflets; leaflets,
6 to 15 mm long and 1.2 to 3 mm wide, linear or oblong, acute, asymmetrical, sessile,
bristly, sensitive with leaves immediately folding at pulvinate joints if touched or jarred;
flowers, 1 to 2 cm in diameter, in small globose, head-like, axillary pedunculate clusters;
peduncles, up to 2.5 cm long, hairy; calyx, absent or very small; corolla, 4-lobed,
sympetalous, very small; stamens 4, 4 to 6 mm long, showy, rose-pink to purple; pods,
0.8 to 2 cm long and 2 to 4 mm wide, clustered, thin, flat, prickly-bristly, indented
between the seeds, splitting at maturity from the undivided margins into 2 to 5 one-
seeded segments; seeds, 2 to 5, about 2 mm across, rounded, brown. Roadside weed near
Buada Lagoon. 4(156N), 6(190).

Peltophorum pterocarpum (DC.) Backer ex Heyne "yellow poinciana", "copperpod",
“golden flamboyant", "yellow flame tree"
syns. P. inerme (Roxb.) Naves; P. ferrugineum (Dcne.) Benth.; Inga

pterocarpa DC.; Caesalpinia inermis Roxb.; C. ferruginea Dene.

Recent introduction. Malaysia to N. Australia. Rare. Medium-sized, heavily-
foliaged, broad-crowned tree, 8 to 15 m high; leaves, bipinnate, 30 to 50 cm long,
rachises brown puberulent, with 4 to 15 pairs of pinnae, each with 8 to 20 pairs of
leaflets; stipules small, caducous; leaflets, about 0.8 to 3 cm long and 3.5 to 10 mm
wide, opposite, oblong, rounded or emarginate, base oblique, dark green; flowers, each
about 2.5 cm across, fragrant, borne in large terminal racemes aggregated into panicles;
bracts, lanceolate, bracteoles none; bracts, buds and axes brownish-reddish pubescent;
calyx tube short, 5-lobed, lobes imbricate; petals, 1 to 2 cm long, orbicular to ovate,

151

slightly unequal, imbricate with frilly margins, subequal, bright yellow; stamens 10, free,
the filaments curved; pods, 5 to 11.5 cm long and 2 to 2.7 cm wide, 2-valved, oblong-
lanceolate, flattened, straight or curved, slightly winged along both margins, copper-
colored, red-brown or brown when ripe, blackening eventually but persistent on the tree,
veiny on the sides, indehiscent; seeds, 1 to 4, transverse, oblong, flattish, pale brown.
Planted ornamental tree on Military Ridge. 5, 6.

Samanea saman (Jacq.) Merr. "rain tree", " monkeypod tree"
syns. Albizia saman (Jacq.) F. v. Muell.; Mimosa saman Jacq.; Enterolobium
saman (Jacq.) Prain ex King; Pithecellobium saman (Jacq.) Benth.; Inga

saman (Jacq.) Willd.

Recent introduction. Trop. America. Rare. Large to massive tree, 7 to 25 m high,
the trunk up to 1 m in diameter, the crown rounded, usually broader than tall; leaves,
bipinnate, with 2 to 9 pairs of pinnae, each with 2 to 10 pairs of leaflets; leaflets, 2 to 8
cm long and 1 to 4.5 cm wide, the distal ones larger than the proximal ones, ovate to
obovate or oblong-elliptic, asymmetric, subsessile, shining above, downy beneath, folding
(closing) in late afternoon for the night and in cloudy or rainy weather; flowers, in
axillary long-pedunculate heads, the central flower larger than the others; calyx, 6 to 7.5
mm long, short lobed; corolla, up to 13 mm long, petals connate up to the middle, pink
with greenish or yellowish lobes; stamens many, 2 to 4 cm long, basally connate,
exserted, white proximally, shading to pink or crimson distally; pods, 9 to 24 cm long,
1.3 to 2.2 em wide and 1.5 to 2.5 mm thick, straight or slightly curved, thick but
compressed, semi-succulent, internally septate, with thickened sutures, brown turning
black when mature, indehiscent, containing sweet, sticky brown pulp within; seeds 15 to
20, exarillate, brown. Ornamental tree in home gardens on Meneng Terrace and at
Buada. 5, 6, 7.

Vigna marina (Burm.) Merr. "beach pea"
erekogo (N); te kitoko (K); saketa (T)
syns. V. lutea (Sw.) A. Gray; Phaseolus marinus Burm.; Dolichos luteus Sw.

Indigenous. Pantropical. Common. Prostrate, creeping, sometimes climbing,
subglabrous, weak-stemmed, long-trailing herbaceous perennial vine; leaves, alternate,
trifoliate; stipules, 2 to 3 mm long, ovate to lanceolate, inconspicuously bilobate at base,
early caducous; petioles, 5 to 10 cm long; leaflets, 4 to 10 cm long and 3 to 8 cm wide,
broadly ovate or suborbicular to obovate, apex rounded to emarginate, base obtuse, 3-
nerved from base, midrib branched upward, lateral leaflets somewhat oblique, in-
conspicuously appressed-pilose on both surfaces but soon glabrate, somewhat fleshy;
flowers, 1.5 to 2 cm long, short-pedicellate, in few-flowered racemose clusters crowded
distally on axillary peduncles, up to 15 cm or longer; petals and stamens, pale to bright
yellow, the standard usually 12 to 14 mm in diameter, obovate, the keel not much longer
than other petals, incurved for about half a complete turn or less; pods, 4 to 8 cm long

152

and 5 to 7 mm wide, linear-oblong or subcylindrical, slightly curved, inflated, slightly
contracted between the seeds, splitting open along the two sides when mature, green
turning brown; seeds, 2 to 10, up to 7 by 6 by 5 mm, ellipsoidal, brown, with an oblong
hilum and undeveloped rim-aril. Found on beaches and in open sites and waste places
behind beaches; reported by Burges (1935) to be "growing plentifully wherever soil is
fairly heavy and moist, e.g., as at Buada. The natives have always considered that other
plants grew better if near Vigna . . ..". Plant used medicinally; leaves crushed to bathe
young girls’ hair and to make adult hair grow long and black; leaves used to cover
earthen oven. 1(30.R), 2, 3(58610), 4(120N), 5(33), 6, 7, 8.

Vigna unguiculata (L.) Walp. ssp. sesquipedalis (L.) Verdc. "long bean", "yard-long
bean", "snake bean", asparagus bean", "asparagus pea"
bin ("bean")(N); te bin ("bean")(K); tau kok (C)
syns. V. sesquipedalis (L.) Fruw.; Dolichos sesquipedalis L.; Vigna sinensis (L.)
Endl. ex Hassk. var. sesquipedalis (L.) Koern.

Pre-World War II introduction. Trop. Africa. Twining, climbing annual herb with
glabrous stems, up to 2 to 4 m high; leaves, pinnately trifoliate; stipules, large, produced
below the base attachment, more or less persistent; leaflets, 5 to 16 cm long and 4 to 11
cm wide, the terminal leaflet with a long petiolule, ovate to rhombic-ovate, acute, entire
or sometimes inconspicuously lobed, the veins sometimes purplish; inflorescence, an
axillary long-pedunculate few-flowered raceme, with flowers in clusters of 3 to 6; the
bracts and bracteoles small, caducous; pedicels, short, shorter or about as long as the
calyx; calyx, campanulate, bilabiate, the two upper lobes completely or partly united, the
lower lip 3-lobed; corolla, much exserted, petals, white or greenish, tinged with yellow,
blue or purple, the standard orbicular with inflexed auricles, the wings slightly shorter
than the standard, the keel petals about as long as the wings or longer, obtuse or beaked,
sometimes incurved, truncate; stamens 10, alternately slightly longer and shorter, the
filaments of 9 connate into a sheath, the vexillary filament free, the anthers uniform;
pods, variable depending on the cultivar, 20 to 100 cm long and 0.3 to 1.1 cm broad,
pendent, linear, subterete, straight or somewhat inturned or twisted, sometimes rather
flaccid, not septate, more or less inflated and sometimes flabby when young, dehiscent,
the style dehiscent; seeds, many, usually 8 to 12 mm long, elongate reniform, variable in
color. Common. Food plant in Chinese gardens at Location and Topside workshops. 5,
6.

153
GENTIANACEAE (Gentian Family)

Fagraea berteroana A. Gray ex Benth. pua (Polynesia); "pua kenikeni" (Hawaii)
eljinut? (B)(N)
syns. Carissa grandis Bertero ex. Guill.; Fagraea berteriana A. Gray ex Benth.;
F. berteriana Benth. ex Seem.; F. grandis Pancher & Sebert

Indigenous? Pacific Islands, from New Caledonia to as far east as the Marquesas
and Hawaii, although possibly an aboriginal introduction in these areas. Extinct; reported
by Hambruch in 1910, but not seen in 1933 or thereafter. Medium to large glabrous,
often-branching tree, 1.5 to 20 m high, branches with conspicuous leaf scars; sometimes
a scrambling, climbing or epiphytic shrub; leaves, up to 15 cm long and 8 cm wide,
opposite, oval-ovate or oblong, rounded, obtuse or sub-acute-acuminate blunt, base
cuneate or decurrent, entire, rather thick, pinnately nerved, midrib stout, lateral veins
slender and obscure; petiole, up to 2.5 cm long; flowers, nearly 5 cm wide, very
fragrant, fleshy, in axillary cymose clusters; calyx, 12 to 15 mm long, rather deeply-
lobed; corolla, 5-parted, tubular, the tube 3 to 5 cm long, the lobes imbricate spreading,
overlapping to the right, creamy-white turning yellow; style greenish; fruit, 3 to 5 cm
long and 2 to 4 cm wide, ellipsoidal, obtuse to acute at apex, rather succulent, yellow
turning orange to bright red when ripe; seeds, very numerous, embedded in pulp.
Fragrant flowers used in garlands and to scent coconut oil in Polynesia and Melanesia. 1.

GERANIACEAE (Geranium Family)
Pelargonium X hortorum "geranium", "fish geranium"
syn. P. hortorum Bailey
Recent introduction. So. Africa. Rare. Tender perennial subshrub, up to 50 cm

high; leaves, 7 to 13 cm in diameter, rounded, scalloped, downy; flowers, not seen. Pot
plant. 6.

GESNERIACEAE (Gloxinia Family)

Columnea gloriosa Sprague "showy column flower", "columnea"

Recent introduction. C. America. Rare. Trailing or pendent perennial herb, stems
up to over | m long; leaves, about 2.5 cm long, pale green, pubescent; flowers, 3.5 to
7.5 cm long, borne on weeping stems, solitary; calyx, about 1.5 cm long, deeply 5-

154

parted; corolla, showy, bright scarlet with a yellow throat and underside of tube. Pot
plant. 6.

Columnea sp. "columnea"

Recent introduction. Trop. America. Rare. Pot plant. 6.

Episcia cupreata (Hook.) Hanst. "episcia", "peacock plant"
syn. Achimenes cupreata Hook.

Recent introduction. Colombia and Venezuela. Occasional. Spreading or creeping
pilose perennial herb with long stolons and short, stout stems, rooting at the joints, up to
30 cm or higher; leaves, 10 to 13 cm long to 6 to 8 cm wide, including the petiole, blade
about 7.5 cm long, opposite, broadly elliptic to ovate, edges scalloped, pinnately-nerved,
succulent, hairy, variable in color from coppery to reddish green or clear green, often
variegated; flowers, about 2.5 cm long, axillary, solitary, borne on long pubescent
peduncles; calyx, deeply 5-lobed; corolla, much larger than calyx, tubular, with 5
subequal, rounded lobes spreading to about 2 cm in diameter, tube red on upper surface,
yellow with red spots on lower surface and within, the limb orange-red, the 3 lower
lobes the longest; fruit, a 2-valved capsule; seeds, ellipsoid, smooth, brown. Pot plant.
5,0:

Saintpaulia ionantha Wendl. "African violet", "Saintpaulia"

Recent introduction. Trop. E. Africa. Rare. Stemless perennial pubescent herb;
leaves, 3.5 to 8 cm long and wide, long-petiolate, alternate to subopposite, suborbicular
to oblong-ovate, succulent, velvety, edges scalloped, often purplish beneath, forming a
rosette or broad tuft of leaves at the base; petioles, pilose; inflorescences, axillary, 2- to
10-flowered cymose peduncles, rising above the leaves; calyx, deeply 5-lobed, the lobes
linear or lanceolate, erect; corolla, 2 to 3.5 cm across, rotate to broadly campanulate,
much longer than the calyx, the tube short, the limb 5-lobed, 2-lipped, the upper lip 2-
lobed and shorter, color variable depending on the cultivar, white or pink to violet, dark
purple or blue; fruit, about 7 mm long, longer than the calyx, a narrow-oblong to
subglobose capsule, dehiscing by decay of the pericarp; seeds, ellipsoid. Pot plant. 5,6.

155
GOODENIACEAE (Naupaka Family)

Scaevola taccada (Gaertn.) Roxb. "scaevola", "saltbush",
"half-flower"; "beach naupaka" (Hawaii)
emet, emed, emit (B)(N); te mao (K); gasu, gahu (T)
syns. SS. sericea var. taccada Makino; S. frutescens sensu auct. non (Mill.)
Krause; S. frutescens var. sericea (Forst.f.) Merr. (nom. nud.); S. koenigii
Vahl; S. lobelia Murr.; Lobelia taccada Gaertn.; L. koenigii (Vahl) Wight

Indigenous. Trop. Asia to Hawaii. Very abundant. Erect freely branching,
spreading, somewhat succulent, soft-wooded, pithy-stemmed glabrous to pubescent shrub,
up to about 2 m high, with leaves spiralled or crowded near the ends of the branches;
leaves, 8 to 26 cm long and 3 to 12 cm wide, alternate, obovate to oblong-spathulate,
apex rounded (obtuse) or emarginate, base cuneate-decurrent, margins somewhat wavy or
obscurely toothed or crenate, glabrous or somewhat puberulent, slightly fleshy, light
bright green, the midrib sometimes faintly purplish, venation obscure; petiole, short, up
to 15 mm long, broad, winged by decurrent blade margins with a basal tuft of sulky
white hairs in the leaf axil; inflorescence, a fragrant axillary forking 3- to 9-flowered
cymose cluster, 2 to 5 cm long, the peduncles 0.5 to 2 cm long; calyx, 5 to 12 mm long
including acute lobes, 5 to 10 mm long; corolla, 12 to 22 mm long, the tube, 10 to 15
mm long, white to greenish or purplish, the limb, 5 to 10 mm long, spreading, white or
pale green without, and purple-veined or brown-bordered, the lobes with membrana-
ceous, sometimes fimbriate or erose margins, asymmetrical, appearing to be split in two
with only half the petals remaining; fruit, 10 to 18 mm in diameter when fresh, drying to
7 to 13 mm, 2-celled, subglobose, bluntly costate, fleshy, white; seeds, 1 or 2. Abundant
in strand vegetation; dominant species and one of first colonizers on strip-mined areas.
Wood considered good for smoking (cooking) fish and the black noddy bird (an important
delicacy at feasts); hollow sticks used as "guns" to shoot gum balls (egato) and small
balls carved from pandanus; inner bark used in the past to make headbands which
resembled noddy-bird feathers and which were worn for traditional dances; leaves used to
wrap food in and to cover the earth oven (com, eyom); Scaevola and Guettarda speciosa
(iut) flowers the first flowers smelled by returning sailors; flowers used in garlands and
either added directly, or boiled with coconut oil to scent it; leaves crushed to yield a
juice to retards loss of hair and cure rashes; inner bark scraped to yield medicine for
abscesses or boils, and white ripened fruit squeezed into eyes as a "pre-eye-drops" cure
for conjunctivitis. 2, 3(58622, 58761), 5(30), 6, 7(27801).

156
HERNANDIACEAE (Hernandia Family)

Hernandia nymphaeifolia (Presl.) Kubitzki "lantern tree"
etiu, yetiu, etsiw (N); te nimareburebu, te bingibing (K); puka, puka vaka (T)
syns. H. sonora L.; H. peltata Meissn.; H. ovigera senus auct. non L.; Biasolet-

tia nymphaeifolia Pres}

Indigenous. Trop. Asia to Pacific Is. Rare. Medium to large tree, up to 20 m
high, with a shortly buttressed trunk and smooth grayish, slightly fissured bark; leaves,
12 to 40 cm long and 10 to 30 cm wide, alternate, rounded-ovate, acute-acuminate,
subpeltate to peltate near base, glabrous, softly leathery, medium green (the ventral
insertion of the petiole usually red), palmately 5- to 9-nerved from petiole; petiole, 5 to
17 cm long; inflorescence, a densely-flowered axillary and terminal long-stalked,
tomentose cymose panicle of numerous, unisexual but monoecious, white to yellowish-
white flowers, 12 to 30 cm long; involucre, green to whitish or pinkish, slightly fleshy,
subtended by 4 bracteoles; each cyme composed of 3 pedicellate white to pale green
flowers, the 2 laterals males, the center one female; male flowers, about 7 mm long, 3-
merous; female flowers, 4-merous; style up to 5 mm long; fruit, about 2.5 cm in
diameter, broadly obovoid-subcompressed, nut-like, brown, nearly smooth, somewhat
ribbed, enclosed in an enlarged, fleshy, white or grayish to reddish succulent balloon-like
vesicle with a circumscissile opening; seed 1. Tree on or near base of escarpment. Very
light wood, which is sometimes found in the form of driftwood, used for canoe outrig-
gers, pull-floats for fishermen to tie fish to, and corks for bottles; fruit rubbed against
rock to burn each other in traditional games. 5(10), 6, 7(27819).

LAMIACEAE OR LABIATAE (Mint Family)

Coleus amboinicus Lour. "Indian borage"
syns. Plectranthus amboinicus (Lour.) Spreng.; Coleus aromaticus Benth.

Recent introduction. Africa and India to Indonesia. Rare. Very aromatic, densely
pubescent, somewhat succulent, low sprawling or ascending perennial herb, up to 1 m
high, with sub-quadrangular to subterete stems; leaves 2 to 11 cm long and 1.5 to 9 cm
wide, opposite, broadly ovate to suborbicular, acute to broadly rounded (obtusish), base
cuneate or rounded to truncate, coarsely crenate to dentate (toothed), succulent, finely
pubescent on both sides, fragile; petioles 1 to 5 cm long, densely pubescent; inflores-
cence, a long simple (unbranched) terminal spicate panicle up to 5O cm or longer bearing
numerous ( up to 30 or more) whorls of flowers on short pedicels at intervals of about 1
to 2 cm; bracts, 3 to 4 mm long; pedicels slender, hirsute, up to 5 mm long; calyx 1.5 to
4 mm long, campanulate, hirsute and glandular, the upper lip erect, broadly ovate-
oblong, the other teeth narrow, very acute; corolla, 7 to 12 mm long, pale blue or violet
to pink, the tube 3 to 4 mm long, declinate, expanding distally, pubescent without, the

157

upper lip to 4.5 by 3 mm, erect, puberulent, the lower lip to 5 to 6 by 4 mm, concave;
fruit, about 0.5 by 0.7 mm, a nutlet, smooth, pale brown. Pot plant at Location and at
Indian residence at Meneng Terrace. Leaves used as a spice in curries and medicinally.
6(211), 7.

Coleus pumilus Blanco "creeping coleus"
syn. C. repens Gurke

Recent introduction. W. Africa. Occasional. Pot plant. 2(59673, 58781), 5, 6.

Ww "

Mentha piperita L. "mint", "peppermint"
syn. M. X piperita L. (M. aquatica L. X M. spicata L.)

Recent introduction. Europe. Rare. Strongly-scented erect to sprawling rhizoma-
tous perennial herb, the stems 4-angled, glabrous to sparsely pubescent, often purplish;
leaves, 1 to 6 cm long and 1 to 3.5 cm wide, opposite, lanceolate-elliptic to oblong-
ovate, ovate or suborbicular, somewhat rugose, sharply acute to apiculate, cuneate to
rounded or cordate at base, serrate at margin, pubescent to glabrous; petioles, 3 to 6 mm
long; inflorescence, a terminal spikelet at stem tips with flowers arranged in whorls
(verticils) which are congested terminally and more distantly-spaced basally, mostly
glabrous except on calyces; calyx, 3 to 4 mm long, tubular, purplish and dotted with oil
glands, the teeth subequal, ciliate, narrowly acuminate, much shorter than calyx tube at
anthesis; corolla, 4 to 6 mm long, pale violet, glabrous or puberulent, the tube exserted;
stamens 4, small, included; fruit, about 0.75 mm long, a brown nutlet. Pot herb planted
in old oil drum on Command Ridge and at Indian residence at Meneng Terrace. 6, 7.

Ocimum basilicum L. "basil", "sweet basil"
dementsi (N); te marou (K); mili (T)

Pre-World War II introduction. Africa to Pacific Is. (Paleotropics). Occasional.
Freely-branching aromatic perennial herb or subshrub, often cultivated as an annual, up
80 cm or higher, the stems 4-angled, the branchlets glabrous to hispid; leaves, 2 to 8 cm
long and 1 to 3 cm wide, opposite, elliptic-lanceolate to ovate and oblong, acute, base
cuneate to attenuate, entire or obscurely dentate, glabrous to scattered-pubescent;
petioles, up to 2 cm long, slender; inflorescences, terminal racemes, 10 to 15 cm or
longer, at stem tips bearing many short-pedicellate, 6-flowered whorls (verticils) at
intervals; bracts, petiolate, green to rich purple, the bracteoles, 2 to 6 mm long; calyx, 2
to 3 mm long at anthesis, enlarging to 5 to 9 mm long in fruit, bilabiate, 10-nerved, with
long hairs inside, greenish, purplish-tinged; corolla, 7 to 9 mm long, tube 3.5 mm long,
upper lip 4-toothed, lower lip entire, white, glabrous to hispid; fruit, of 4 nutlets, 1.5 to
2.5 mm long, obovoid or ellipsoid, brown or blackish, pitted, swelling in water and
becoming slimy. Planted in home gardens; common in I Kiribati and Tuvaluan gardens at

158

Location. Fragrant flowers and leaves used in garlands and for scenting coconut oil. 2, 3,
5(141), 6, 7(22305).

Ocimum sanctum L. "sacred basil", tulsi (Hindi)
demere (N)?; te marou (K)
syn. O. tenuiflorum L.

Pre-World War II introduction. Trop. Asia, now Pantropical. Rare. Erect,
aromatic, much-branched herb or subshrub up to | m high, the branchlets soft-pubescent;
leaves, 3 to 7.5 cm long and 1 to 2.8 cm wide, elliptic to elliptic-oblong or rhomboidal,
obtuse to acute, base cuneate to attenuate, margin entire or remotely serrate, upper bract-
like leaves ovate to cordate, green to grayish-purple, pubescent on both surfaces but
especially on nerves beneath; petioles, up to 2.5 cm long; inflorescences, slender racemes
at stem tips, 8 to 10 cm long, bearing whorls (verticils) of flowers at intervals; brac-
teoles, 2 to 3 mm long, ovate, acuminate, ciliate; pedicels, up to 4.5 mm long; calyx,
2.5 to 3.5 mm long at the anthesis, enlarging to 5 mm in fruit, often purplish, glabrous
within except occasionally puberulent near the base of upper lobe, the upper lip subor-
bicular, reflexed, short-apiculate, the lower lip longer than the upper lip, the teeth 4,
lanceolate; corolla, 3 to 5 mm long, the tube 1.5 to 2 mm long, upper lip with rounded
lobes, lower lip entire, purplish to pale pink or white; stamens 4; fruit, 0.8 to 1.5 mm
long, nutlets, subglobose-compressed or broadly ellipsoid, purple-green to brown, smooth
to minutely pitted. Planted ornamental. Fragrant flowers and leaves used in garlands and
headbands and for scenting coconut oil. 5, 6, 7.

Plectranthus oertendahlii Fries "Swedish ivy", "prostrate coleus"

Recent introduction. S. Africa. Rare. Prostrate or trailing, tender or somewhat
succulent, perennial herb, up to 50 cm high, with creeping reddish stems; leaves,
suborbicular, velvety, bronze-green with silver along the veins and purplish beneath;
inflorescences, erect or ascending racemes or panicles, 10 to 15 cm long, bearing whorls
(verticils) of flowers at intervals; corolla, 2-lipped, pink. Pot plant. 3(58673, 58781), 5,
6.

Plectranthus scutellarioides (L.) R. Br. "painted nettle", "coleus", "painted-
leaf plant"
syns. Coleus scutellarioides (L.) Benth. and C. blumei Benth.; Ocimum scutel-
larioides L.; Solenostemon scutellarioides (L.) Codd

Recent introduction. Malaysia. Occasional. Erect or ascending, tender or
somewhat succulent, aromatic brown-pubescent perennial herb up to 1.5 m high, the
stems 4-angled and branches glabrous to pubescent; leaves, opposite, 4 to 17 cm long and
3 to 10 cm wide, broadly ovate to ovate-cordate, acute to acuminate, base truncate to

159

rounded or cuneate and attenuate, crenate, doubly crenate to laciniate, scabrid to
subglabrous, variegated, highly ornamental, variously colored, green, red, purple, yellow
or white; petioles, 1 to 8 cm long, slender; inflorescences, erect terminal racemes or
panicles, 5 to 40 cm long, bearing whorls (verticils) at intervals; bracts, 4 by 5 mm,
ovate, long-acuminate, deciduous; pedicels, 3 to 4 mm long; calyx, 2 to 4 mm long and
2 to 3 mm wide, enlarging to 7 mm long in fruit, bilabiate, obliquely campanulate,
pubescent and gland-dotted, 10-nerved; corolla, 8 to 18 mm, long, infundibular, blue to
purple or mauve, the tube, about 5 mm long, paler, puberulent, abruptly decurved, the
upper lip about 1.5 mm long, erect, the lower lip deeply concave, up to 6 mm long;
stamens 4; fruit, nutlets, 0.75 to 1.2 mm long, lenticular to broadly ovoid or subglobose,
brown, smooth, glossy. Planted ornamental and pot plant. 5, 6, 7.

LAURACEAE (Laurel Family)

Cassytha filiformis L. "beach dodder", "giant dodder","devil’s twine"
denuwanini, denuwenini, eduwinini (B)(N); te ntanini (K); fetai (T)

Indigenous. Pantropical. Abundant. Slender, filiform, branching, twining or
climbing light green to yellowish-green or yellowish-orange, glabrous parasitic herb, up
to 3 to 8 m long, attaching to host plants by means of sucker-like haustoria, often
forming sense, tangled mats; leaves, alternate, spirally arranged, reduced to minute
scales; inflorescences, erect solitary pedunculate spike-like clusters, 1.5 to 5 cm long,
borne in the axils of minute bracts; rachis, rather thick, glabrous, finely brown-hairy;
bracts and bracteoles ovate-circular; flowers, bisexual, the perianth greenish-white to
yellowish-white, somewhat fleshy, tepals 6, white, the 3 outer tepals, connate, broadly
ovate-circular, about 0.75 mm across, the 3 inner ones broadly ovate, glabrous, 2 to 2.5
mm long; stamens 9; staminodes, few, yellow; fruit, 4 to 7.5 mm in diameter, subglob-
ose, white when mature, enclosed in a fleshy perianth tube, 1-seeded; seed with a
membranaceous or coriaceous testa. Parasite on other plants, found generally on natural
vegetation at all elevations. Entire plant used as garlands and headbands; plant used for
"black magic" by I Kiribati and other islanders, a practice occasionally copied by
Nauruans; tender tips used at times in the past for scenting coconut oil; fruit eaten by
children in the past. 2(23.5), 3(58590), 4(163N), 5(6), 6, 7, 8(9566).

Persea americana Mill. "avocado", "avocado pear", "alligator pear"
syns. Laurus persea L.; Persea gratissima Gaertn.f.

Recent introduction. Mexico. Rare. Medium to large evergreen tree, up to 12 m
or taller; leaves, 7 to 30 cm long and 2.5 to 20 cm wide, alternate and spirally arranged,
elliptic, ovate-oblong or obovate-oblong, acute-acuminate, chartaceous to somewhat
coriaceous, glaucous beneath, downy when young, nerves alternate, pinnate, prominent;
petioles, 1.5 to 5 cm long; inflorescences, axillary many-flowered downy panicles

160

crowded near the ends of branches; peduncles and pedicels, yellowish-green, pubescent;
flowers, bisexual, somewhat fragrant, subtended by lanceolate deciduous hairy brown
bracts, 4 mm long; the perianth of 6 tepals, 10 to 15 mm in diameter, greenish to
yellowish white, short-pedicellate; stamens 9, in 3 whorls, the staminodes, conspicuous,
orange to brown; pedicels cylindric or enlarged and fleshy when in fruit; fruit, 7 to 25
cm long and 5 to 12 cm in diameter, subglobose to pear-shaped (pyriform) fleshy, 1-
seeded drupe, skin light-green to purplish, flesh light-green to yellow-green, of butter-
like consistency, edible; seed, single, subglobose, up to 5 cm in diameter, with 2 brown
seed coats and 2 fleshy, whitish or pink cotyledons. Seedling planted in garden at
Meneng Terrace. 5.

LECYTHIDACEAE (Brazilnut Family)

Barringtonia asiatica (L.) Kurz "fish-poison tree", "barringtonia"
kwenbabai, kwenababai, eijinut (B)(N); te baireati (K); futu (T)
syns. Mammea asiatica L.; Barringtonia speciosa Forst.; B. butonica Forst.

Indigenous. Indo-Pacific. Occasional. Large, spreading, round-crowned tree, 10 to
20 m high, with gray bark and rather stout branches; leaves, 15 to 50 cm long and 8 to
24 cm wide, alternate, clustered toward the ends of branches, obovate or oblong-obovate,
obtuse or slightly emarginate, base cuneate, subcoriaceous, glabrous, glossy-green,
subsessile; petioles, 1 to 5 cm long; inflorescences, terminal more or less erect racemes,
2 to 15 cm long and 15 cm broad; pedicels, 2 to 9 cm long; calyx, 2-lobed, sepals, 2 to
4 by 2 to 3 cm, concave, obtuse, persistent on fruit; corolla, 4-parted, sweetly fragrant,
petals, about 4 to 8 cm long and 2.5 to 4.5 cm wide at anthesis, elliptic to ovate, white,
early deciduous; stamens numerous, 4 to 12 cm long, white at base, pink- or red-tinged
distally, anthers yellow; style, 6 to 15 cm long, white at base, pink- or red-tinged
distally; fruit, 8 to 12 cm long and broad, 4-angled, 1-seeded, pyramidal or subturbinate,
exterior fibrous, crowned by calyx (2 large persistent sepals), pendent and heavy when
ripe, buoyant; seed, 4 to 5 cm long and 2.5 to 4 cm wide, ovoid or ellipsoid. Spon-
taneous or planted on coastal strip, often in home gardens; larger concentrations on
escarpment leading to plateau above Anibare Bay. Wood a favoured timber and fuelwood
for cooking toddy syrup (kamwaerara); fruit possibly used to poison fish in the past.
Fruit commonly used as a fish poison or stupefacient elsewhere in the Pacific. 1(48.R),
2, 3(58665), 5(36), 6, 7.

161
LYTHRACEAE (Lythrum Family)

Lagerstroemia indica L. "crape myrtle"

Recent introduction. S. China. Rare. Glabrous, deciduous shrub or small tree, 2
to 5 m high, with 4-angled branchlets; leaves, 3 to 10 cm long and 2 to 5 cm wide,
alternate, elliptic to oblong-subobovate, obtuse to acute, subsessile; inflorescences,
terminal or axillary panicles, 5 to 20 cm long, bearing clusters of flowers, each about 2.5
to 3 cm across; calyx, 6-lobed, campanulate, glabrous; petals 6, 12 to 15 mm long,
ovate-suborbicular, wrinkled, clawed, scentless, white, pink or lavender to rich purple or
blue; stamens about 15 to 40; fruit, about 0.5 to 1 cm long, ovoid, capsular; seeds,
winged. Planted ornamental on Command Ridge. 3(58712), 5, 6.

MALPIGHIACEAE (Malpighia Family)

Stigmaphyllon ciliatum (Lam.) Juss'Brazilian golden vine", "golden cup", "orchid vine"

Recent introduction. W. Indies to Brazil. Rare. Slender, woody vine; leaves, 2.5
to 8 cm long and nearly as wide, cordate, glabrous with pubescent margins; inflorescen-
ces, axillary racemose clusters of 3 to 7 flowers, each about 3.7 cm in diameter; sepals
5; petals 5, bright yellow, free, stalked at base, unequal, wavy, orchid-like; stamens 10,
unequal, only 6 bearing anthers; styles 3; fruit, 3-parted, each part with a broad wing
about 2.5 cm long. Planted ornamental climber. 3(58725), 6.

Tristellateia australasiae Rich. "bagnit"
syn. Tristellateia australis Rich.

Recent introduction. Malaysia and Australia. Occasional. Glabrous, long woody
climber with stems dotted with raised lenticels; leaves, 2 to 12 cm long, opposite or
verticillate, ovate, acute or obtuse, base rounded or subcordate with 2 glands, glabrous,
rather pale green; petiole, 5 to 20 mm long; inflorescences, terminal racemes at branch
tips bearing 12 or more flowers, each nearly 2.5 cm in diameter; sepals 5; petals 5,
bright yellow, 4 of them slightly falcate, equal, each about 1 cm long excluding the 2
mm long pinkish-orange claw; stamens, usually 10 of unequal length, yellow turning red;
style 1, curved, undivided; fruit, samaroid, with a lateral wing with 5 to 8 slender
flattened lobes each up to about 1 cm long. Planted ornamental climber. 5, 6(179, 205),
1s

162
MALVACEAE (Mallow Family)

Abutilon asiaticum (L.) Sweet var. supraviride Fosb.
ekaura, inen ekaura (N); te kaura ni Banaba (K)

Indigenous. S. E. Asia to the Pacific Is. Occasional. Erect velvety-pubescent
subshrub, 0.5 to 2 m high; leaves, 5 to 12 cm long, alternate, suborbicular-ovate cordate
at base, coarsely crenate-serrate or lobed, palmately-nerved, downy gray-green, long-
petiolate; flowers, 1.5 to 2.5 cm across, axillary, solitary, on long pedicels, 4 to 7 cm
long; calyx 5-lobed; corolla, 1.5 to 2.5 cm across, petals 5, about 1 cm long or longer,
imbricate, deltoid-obovate, orange-yellow; fruit, circular, a capsular schizocarp of 11 to
20 radiating carpels, hirsute, brown when dry, each carpel flattened, somewhat boat-
shaped, apiculate by the short persistent style-remnant, about 8 mm _ long; seeds,
reniform, stellate-pubescent. Found in waste places and ruderal habitats along coastal
strip, especially near the Ijuw-Anibare boundary, and among pioneering weeds in topsoil
in areas recently cleared for phosphate mining. Tender meristem used to scent coconut
oil; flowers used in garlands and headbands. 2, 3(58807, 58805), 4(107N), 5(107), 6,
7(22306).

" "

Gossypium barbadense L. "sea-island cotton", "cotton"
duwoduwo (N); te baubau (K)
syns. G. brasiliense Macf.; G. peruvianum Cav.

Pre-World War I introduction. Trop. America. Extinct? Erect branching shrub,
up to 3 m or higher; leaves, 7 to 13 cm long, alternate, rotund to ovate, cordate, deeply
palmately 3 to 5 (rarely 7)-lobed or laciniate, glabrous with short, fugacious hairs, long-
petiolate; stipules foliaceous, lanceolate or ovate, auriculate; inflorescences, terminal or
on short axillary branches, solitary or 2 to 4-flowered racemes; bracteoles of epicalyx 3,
much longer than the calyx, with cordate base and a deeply incised or laciniate apical
lobe; top of pedicel with or without glands below the bracteoles; corolla, 5 to 8 cm
across, obovately 5-petaled, petals, about 5 to 8 cm long, yellow, becoming streaked with
red or purple on fading, usually with a basal reddish or purplish spot; fruit, a 3-valved
ovoid, leathery, pitted capsule dotted with black oil glands; seeds, numerous, ovoid,
black, covered with fine, long, pure white easily-removed woolly fibers or hairs (cotton).
Collected by Burges in 1935, but not seen by subsequent collectors. Grown by Nauruans
in the past for the cotton which was used to stuff mattresses and pillows. The Nauruan
name for cotton is the same as for kapok. 2.

163

Hibiscus esculentus L. "okra", "gumbo", lady’s finger"; bindi (Fiji Hindi)
syn. Abelmoschus esculentus (L.) Moench.

Recent introduction. Asia. Coarse, erect suffruticose, annual herb, 1 to 3 m high,
with green or red-tinged stems; leaves, 10 to 25 cm long and 10 to 35 cm wide,
alternate, broadly cordate, palmately 3- to 7 lobed, serrate, hirsute, pale green beneath;
petioles, 15 to 35 cm long, hispid, often red-tinged; flowers, solitary, axillary; peduncle
about 2 cm long; epicalyx of up to 10 narrow bracteoles, up to 1.5 cm long, usually
falling before fruit reaches maturity; calyx completely fused as flower develops, splitting
longitudinally as flower opens, falling with the corolla after anthesis, 2 to 3 cm long;
petals 5, yellow with crimson spot on claw, obovate, 5 to 7 cm long; staminal column
united to base of petals, 2 to 3 cm long, with numerous stamens; stigmas small, 5 to 9,
deep red; fruit, 10 to 30 cm long and 2 to 3. cm across, a pyramidal-oblong, beaked
capsule, longitudinally furrowed, hirsute or glabrous, dehiscing longitudinally when ripe;
seeds, 5 mm in diameter, dark green to brown, rounded, tuberculate.

Hibiscus manihot L. "push spinach", "edible hibiscus"; "bush hibiscus spinach";
"pele" (Polynesia); mbele (bele in Fiji); nambere (K); pele (T); kabis, slippery
kabis (SI)

syn. Abelmoschus manihot (L.) Medik

Recent introduction. S. E. Asia. Occasional. Erect, perennial, often suffruticose
woody herb or subshrub, 1.5 to 3 m or higher, with young stems, petioles and pedicels
glabrous or short-hairy, often accompanied by longer hairs; leaves, 10 to 60 cm by 5 to
60 cm, alternate, extremely variable in shape, orbicular to ovate-oblong, base cordate,
either contracted into an elongated-triangular acumen from a broad base or palmately 3 to
7-lobed (or 3 to 7-fid) to subentire, lobes or segments triangular-ovate, oblong-lanceolate,
obovate-spathulate or linear, entire, dentate or serrate, pellucid-dotted, more or less
crenate, glabrous or pubescent; petiole, 2 to 50 cm long; stipules, 2 to 8 cm long,
filiform, caducous, flanked by paired, variable lobed or entire leaflets, 2 to 6 cm long;
inflorescences, axillary, solitary, on pedicels, 1 to 5 cm long; bracteoles of epicalyx, 4 to
6 (-8), 1 to 3 cm long and 0.5 to | cm broad, ovate to oblong, not rarely coherent in
pairs, pubescent on both sides; petals 3, 5 to 8 cm by 3 to 6 cm, yellow with purple
centers; fruit, 3.5 to 6 cm long and 2 to 2.5 cm broad, capsular, pentagonous, with 5
prominent costas, concave between the costas, loculicidally dehiscent; seeds, many, about
3 to 4 mm long, subglobose, asymmetrical, dark brown, pubescent. Food plant in
gardens at Location and Topside Workshop and occasionally in other home gardens.
Nutritious slippery green leaves cooked as a green vegetable by Solomon Islanders,
Tuvaluans, I-Kiribati and Fijians. Along with taro leaves, one of the two most important
leafy green vegetables in the Pacific Is. 5, 6(105), 7.

164

Hibiscus mutabilis L. "changeable rose mallow", “changeable rose", "variable rose"
dorot ("the rose")(N)

Recent introduction. S. China. Rare. Large, downy pubescent shrub, up to 5 m
tall, with both simple and stellate hairs; leaves, 10 to 25 cm long and broad, alternate,
orbicular, cordate, 3 to 7-lobed, long-acuminate, 5 to 11-nerved, dark green above and
light green and tomentose beneath; petioles, 2 to 8 cm long; inflorescences, axillary
(occasionally terminal), usually single but sometimes double; pedicels, 7 to 10 cm long,
accrescent, articulate; epicalyx, 6-segmented, well-developed, persistent, segments about
2 cm long, linear-lanceolate; calyx, 5 to 7 cm across, deeply divided, 5-lobed, lobes 2 to
3.5 cm long, abruptly widened above the base; corolla, 8 to 12 cm across, opening white
changing to pink, many-petaled, petals 4 to 5 cm long; fruit, about 2.5 cm wide,
subglobose, capsular, 5-locular, setose-lanate; seeds, reniform, tomentose, the hairs 2 to
4 mm long, spreading to subspreading. Planted ornamental along road to Meneng
merrace. S(15i7);, 7-

Hibiscus rosa-sinensis L. "hibiscus", "red hibiscus"
dorot ("the rose")(N); te roti ("the rose")(K); aute (T)

Pre-World War II introduction. Trop. Asia. Common. Many branching, glabrous
shrub, up to | to 4 m tall, with erect to drooping branches; leaves, about 6 to 15 cm long
and up to 12 cm wide, alternate, variable, ovate-elliptical, acute or acuminate, crenate-
serrate, subcordate, obtuse or subacute, green on both sides, palmately 5-nerved;
petioles, up to 5 cm long; inflorescences, axillary, near the ends of branches on elongate
pedicels, 1.5 to 7.5 cm long; epicalyx of 5 or more ovate- to linear-lanceolate bracts 5 to
18 mm long; calyx, green, S-lobed for about half its length, persistent; corolla, mostly 8
to 12 cm across, single or in some forms double, campanulate or broadly funnelform,
petals entire, showy, usually bright red, but also pink, yellow, light-orange or white;
staminal tube slightly longer than the corolla, often nodding or pendulous, crimson; fruit,
capsular, ovoid-rounded, 5-valved, 15-seeded (or less); seeds, subglobose, rarely seen.
Planted ornamental and hedge plant; flowers used in garlands and for decoration. One of
the commonest and most widespread of all ornamental plants in tropical regions.
3(58791), 5(126), 6, 7.

Hibiscus schizopetalus (Mast.) Hook.f. "coral hibiscus", "dragon flower"
dorot (N)

Recent introduction. E. Africa. Rare. Glabrous shrub, 2 to 4 m high, very much
like H. rosa-sinensis; leaves, alternate, ovate-elliptical, acute, dentate; inflorescences,
solitary, pendulous on long pedicels at the ends of slender, drooping branches; pedicels,
8 to 10 cm long; segments of epicalyx 1 to 2 mm long; calyx irregularly 2- to 4-lobed,
sometimes spathaceous; corolla, deeply dissected (laciniate), reflexed, petals pink-and-
white to coral-red, sometimes with white or yellow margins; staminal tube about twice as

165

long as petals, slender, flaccid, red, curved at the tip. Planted ornamental. 3(58770),
5(125), 6.

Hibiscus tiliaceus L. "beach hibiscus", hibiscus tree", "hau" (Hawaii)
ekwane (N); te kiaiai, te rao (K); fou, fau (T)
syns. Pariti tiliaceus (L.) A. St. Hil.; P. tiliaceum Britt.; Paritium tiliaceum (L.)
A. St. Hil.

Indigenous. Pantropical. Very abundant. A small, dense, broad-crowned, often
scrambling tree, 3 to 10 (rarely 15 to 18) m high, intricately branched when mature,
with many spreading low branches from a short trunk; youngest branches gray-pubescent,
older ones glabrate with grayish rather smooth very fibrous and mucilaginous bark and
bast fiber; leaves, up to 15 or 20 cm long and nearly as wide, round-ovate, abruptly
acuminate, base deeply cordate, velvety pubescent when young, glabrate and dark green
above, grayish-white stellate-tomentose beneath and slightly glaucous, palmately 7- to 9-
nerved with linear basal glands dorsally; petioles, up to 12 or 15 cm long; stipules, 2 to
4 cm long, broadly attached, ultimately leaving annular scars; inflorescences, terminal or
axillary, solitary or in few-flowered open cymes or panicles; epicalyx conspicuous,
shorter than calyx, with 8 to 12 deltoid, acute segments (bracts); pedicels 1 to 3 cm long;
calyx 5-lobed; corolla, about 10 cm across, cupular-campanulate, yellow with a maroon-
purple eye, aging (after falling) to salmon red; staminal tube shorter than corolla, yellow;
style rich purple distally; fruit, 1 to 2.5 cm long and 2 cm wide, ovoid globose, capsular,
5-celled, pubescent, grayish-brown, splitting at top when mature; seeds, generally 15,
glabrous, brownish black. In thickets and forest on escarpment surrounding plateau and
on inner and outer edges of coastal strip. Timber used for house construction and
considered good for canoe outriggers, poles for noddy bird nets, and the best wood for
the construction of frigate bird nesting platforms (efeo, eftea); very soft pieces of wood
rubbed together in the past to make fire by friction in the procedure known as ikumo;
inner bark (bast fiber) used to make white fiber, which after stripping off outer bark and
soaking in mud and sea water, is used to make skirts (ridi), special hula skirts (ingung),
and baskets (ebwer, eber); fibre used for straining coconut cream and for lashing house
rafters; leaves used for parceling pig and other foods for cooking in the earth oven
(eyom, eom); leaves cooked with water as a cure for diarrhea. 1(22.R), 2, 3(58747),
4(169N), 5(28), 6, 7.

Hibiscus ornamental hybrids "hybrid hibiscus"

Recent introduction. Rare. Origin? Variable shrubs, up to 2 m or higher; leaves
smooth to rough; flowers, single or double; bracts, few to several; calyx of single forms
5-lobed, showy, more or less campanulate; staminal column more or less covered with
stamens surrounding all but the tip of a longer S-lobed style; fruit a 5-valved capsule
containing 15 or more seeds. Planted ornamentals. 3, 5, 6, 7.

166

Malvastrum coromandelianum (L.) Garcke "false mallow"
syns. M. tricuspidatum A. Gray; Malva coromandeliana L; Malva tricuspidata
Re Br

Recent introduction. C. America to S. United States. Common. Tough-stemmed,
woody-rooted, widely-branching annual herb or subshrub up to 1 m high (usually less);
stems, pubescent with 4-rayed hairs; leaves, 2 to 6 cm long and 1 to 2.5 cm wide,
alternate, ovate to ovate-elliptic or ovate-lanceolate, acute or obtuse, serrate, appressed-
pubescent, 3-nerved from the base, midrib branched upward; petioles, 1.5 to 4 cm long,
slender; stipules, 0.5 to 1 cm or more long, lanceolate to very narrow; inflorescences,
axillary, solitary or finally appearing in short axillary and terminal clusters; peduncles,
about 5 mm or longer; epicalyx of 3 free narrow subulate bracts; calyx, deeply 5-lobed,
the sepals ovate, acute, pubescent; corolla, about 1.2 to 2 cm across, broadly cam-
panulate, 5-petaled, opening after mid-day, finally falling with the staminal tube; petals
slightly unequally bilobed at apex, yellow to salmon-orange; staminal column shorter than
the corolla, divided at apex into numerous filaments; fruit, about 6 mm across, a discoid
(wheelshaped) schizocarp breaking up at maturity into 10 one-seeded segments (meri-
carps), each about 3 mm long, with a few stiff hairs and 2 stubby points on the convex
side; seeds, 1.5 mm long, obliquely reniform, black with a dull surface. Weed in settled
areas, roadsides and ruderal sites. 3(58694, 58728, 58766), 5, 6(207), 7, 8.

Malvaviscus arboreus Cav. var. penduliflorus (Moc. & Sesse ex DC.) Schery
"sleeping hibiscus", "Turk’s cap"
syns. M. penduliflorus Moc. & Sesse ex DC.; M. conzatti Greenm.; M. gran-
diflorus Hort. non H.B.K.

Recent introduction. Trop. America. Occasional. A shrub, 1 to 3 m high, with
fibrous bark; leaves, 7 cm to 20 cm long and up to 10 cm wide, alternate, narrowly
ovate to ovate-oblong or ovate-lanceolate, acute to obtusely acuminate, base broad
cuneate to rounded, serrate; petioles, usually 5 to 12 cm long, puberulent; stipules, about
7 mm long, linear; flowers, solitary, axillary, pendulous, hibiscus-like; pedicels, 4 to 6
cm long or longer, slender, pubescent; epicalyx of usually 6 to 7 linear oblong bracts,
about 1 to 1.5 cm long, apex obtuse, margins ciliate; calyx, 5-lobed, exceeding bracts of
epicalyx by up to | cm; corolla, about 4 to 7 cm long, S-petaled, bright red (occasionally
pink), petals, unequally obovate, slightly keeled, flattened at apex, base suricular,
remaining rolled, overlapping and closed; staminal tube slightly exserted, slender, reddish
or pale pink with dull purple anthers; fruit, a 5-seeded schizocarp, globose, fleshy, red.
Planted ornamental. 5(124), 6.

167

Sida acuta Burm.f. "spiny-headed sida", "broom weed"
"coffee bush" (N)
syns. S. carpinifolia L.f.; S. glomerata Cav.

Recent introduction. Pantropical Common. Low, freely-branching perennial
subshrub, up to | m tall, with slender stems and a strong taproot; leaves, 1.5 to 7.5 cm
long and 0.5 to 2 cm wide, alternate, lanceolate or linear-lanceolate, acute, obtuse,
cordulate or rounded at base, sharply serrate, yellowish-green, more or less stellate-
pubescent or glabrous beneath, palmately 3-nerved from base, pinnately nerved above the
base, midrib branched upward; petioles, 3 to 5 cm long; stipules linear-acute or linear-
subulate, up to 12 mm long; inflorescences, solitary or paired, axillary or sometimes
terminal; pedicels, 2 to 10 cm long; calyx, about 5 mm across, 5-lobed, lobes pointed;
corolla, about 4 to 15 mm across, 5-petaled, petals united at base, obovate, rotate,
yellow; stamens numerous, fused into a column (monadelphous); fruit, 4 to 5 mm broad,
a 6 to 10-carpellate schizocarp, carpels 2-awned, grooved dorsally; seeds, about 1.5 to 2
mm long, rounded-wedge-shaped, dark reddish-brown. Weed on coastal strip, along
roadside, and in and near thickets. Two forms, a narrow ovate-lanceolate-leaved form
(22303) and a broader-leaved form (96) exist. 3(58615, 58649, 58806), 5(23, 96), 6,
7(22303), 8(9558, 9582).

Sida fallax Walp. "lima" (Hawaii)
ekaura, idibin ekaura (N); te kaura (K)

Indigenous. Indo-Pacific. Rare. Small downy, often almost prostrate, shrub, up to
50 cm high; leaves, about 2 to 4 cm long and | to 3 cm wide, alternate, oblong or
cordate, acute or obtuse, serrate or scalloped-edged, glaucous, downy; petioles, about 1
to 2 cm long ??; flowers, axillary or terminal, solitary or 2.or 3 near ends of branch tips;
pedicels, 2 to 5 cm long, slender ??; calyx, about half as large as corolla, 10-ribbed,
downy; corolla, about 2 to 2.5 cm across, 5-petaled, yellow to rich orange, reddish near
the center; stamens numerous, fused into a column (monadelphous); fruit, a wheel-like,
7- to 12-carpellate schizocarp with 1-seeded, short-awned seed cases or carpels; seeds ??.
Found in ruderal habitats on coastal strip and in areas cleared recently for phosphate
mining. Unopened flower buds used, after soaking in coconut oil to retard their opening
and make them last, to make headbands and necklaces worn by dancers and sportsmen
during special occasions; dried and treated leaves used by I Kiribati, in Kiribati, as a
very strong fertilizer and mulch in ceremonial giant swamp taro (Cyrtosperma chamis-
sonis) gardens. 1(5.R), 2(7.5), 5(108), 6(160, 162).

Sida rhombifolia L. "broomweed", "broom plant", "Cuba jute", "Paddy’s lucerne"
"coffee bush", itsi (tea) (N)

Recent introduction. Pantropical. Common. Tough-stemmed, much-branching,
erect, stellate-pubescent shrub, 30 cm to 1.5 m high; leaves, 1.5 to 8 cm long and 0.7 to

168

2.5 cm wide, alternate, elliptic or rhombic-ovate to subspatulate or lanceolate, acute to
obtusish, base cuneate to very narrowly obtuse at petiole, serrate near apex, somewhat
glaucous-green above, pale beneath, glabrous above, densely stellate-pubescent beneath;
petioles, 2 to 6 mm long (rarely longer); stipules 3 to 10 mm long, narrow; flowers,
solitary or occasionally paired in axils; pedicels 1 to 5 cm long, slender; calyx, about 15
mm wide, 5-lobed, lobes deltoid-apiculate; corolla, about 6 to 18 mm wide, 5-petaled,
petals free, obovate, unequally bilobed, rotate, yellow to pale orange; stamens numerous,
fused into a column (monadelphous); fruit, about 8 mm in diameter, an 8- to 10 (rarely
12)-carpellate schizocarp, carpels about 3 to 4.5 mm long, deltoid, apiculate, stellate
pubescent dorsally, with 2 sharp awns, about 1 mm long at apex; seeds, 1 per mericarp,
about 2 mm long, rounded wedge-shaped or almost pyramidal-triangular, dark brown or
black. Weed of roadsides, waste places and semi-shaded areas. Tea made from leaves
during World War II; leaves boiled in water used to treat blisters. 3(58621), 4(138N), 5,
GiiieeS(O575):

Sida spinosa L. var. angustifolia (Lam.). Griseb. "prickly sida"

Recent introduction. Pantropical. Occasional. Soft pubescent herb, up to 1 m
high, the young stems covered with minute soft hairs; leaves, 2.5 to 5 cm long and 0.4 to
2 cm wide, alternate, ovate-lanceolate, or oblong, acute, base rounded, obtuse or
subcordulate, serrate; petiole, up to 2 cm or longer, slender; stipules, small, pointed, one
at base and 2 lateral, usually curved downward; flowers solitary or in axillary or terminal
paniculate clusters of 2 or 3; corolla, about 1 cm across, 5-petaled, light yellow; fruit, an
ovoid 5-carpellate schizocarp, carpels 2-beaked; seeds, triangular, smooth, dark brown.
Weed in waste places. 4, 6.

Thespesia populnea (L.) Sol. ex Correa "milo" (Hawaii, Polynesia)
itira, itirya (N); te bingibing (?)(K); milo (T)
syns. Hibiscus populneus L. (at least in part); H. bacciferus Forst.f.; Malvavis-
cus populneus (L.) Gaertn.

Indigenous. Paleotropics. Occasional. Medium tree, 3 to 15 m (rarely 20 m)high,
with a fairly stout trunk, rough corrugated bark, dense round crown, and glabrous
branches; branch tips silvery brown lepidote or scurfy, glabrescent; leaves, 5 to 15 cm
long and almost as broad, alternate, rather crowded, ovate, base cordate with an open
sinus, acuminate, entire or rarely somewhat lobed, green above, only slightly paler
beneath, glabrous, slightly fleshy-coriaceous, glossy, usually at right angles to petiole,
the tip pointing down, midrib yellowish, palmately 7- to 7-nerved; petioles, 3 to 12 cm
long; inflorescences, solitary, axillary; pedicels, about half as long as the petioles, stout;
epicalyx, 3 to 5-bracteate, bracts, 4 to 17 mm long, oblong to lanceolate, caducous;
calyx, about 18 mm in diameter, rim- or disc-like, unlobed or scarcely toothed, persis-
tent; corolla, 8 to 10 cm across, 5-petaled, campanulate, rather persistent, pale yellow
with a maroon or reddish center of "eye", fading to purple or pinkish-purple; staminal

169

column fused (monadelphous), shorter than corolla, cylindric, toothed at apex, pale
yellow; stigmas yellow; fruit, about 2 to 4.5 cm across, depressed-pentagonal-globose,
usually 4- to S-celled, coriaceous-woody, glabrous, green becoming purplish or brown at
maturity, with a yellowish gum, indehiscent, irregularly crumbling upon aging, calyx
persistent at base; seeds, 4 per cell, usually 7 to 8 mm long, ovoid to obovoid, glabrous
or silky pubescent (villous). Growing along edges of mangroves in Anetan; planted on
golf course. Specimens (73.R) have leaves scarcely cordate, almost subtruncate at base,
as in Thespesia populneodes, but have seeds villous on angles, as in 7. populnea.
Considered the best wood for house construction, woodcarving, furniture and canoe
outriggers; wood also used in traditional stick games. 1(73.R), 2, 3(58745), 4(165N),
5(57), 6, 7(27821).

MELIACEAE (Mahogany Family)

Melia azedarach L. "Indian lilac", "China berry", "Persian lilac", "pride of India"
gadong, gadung (N)

Pre-World War I introduction. Trop. Asia. Common. Small to medium tree, 3 to
12 m (rarely 15 m) high, with wide-spreading branches and coarse weak wood; leaves,
up to 50 cm or longer, alternate, bipinnate, with 3 to 7 pairs of leaflets, the lowest
pinnae sometimes once more divided (thus partly tripinnate); leaflets, 3 to 9 cm long and
1 to 3 cm wide, opposite, oblong-ovate or lanceolate, acute to acuminate, base somewhat
inequilaterally acute, serrate, glabrous; petiolules, up to 8 mm long; inflorescences,
axillary, open, many-flowered panicles, 10 to 30 cm long; calyx, 5- to 6-lobed, puberu-
lent; corolla, nearly 2 cm across, 5 (or 6)-petaled, the petals about 8 mm long, oblan-
ceolate, purple to lavender or nearly white, fragrant; staminal tube cylindric, as long as
petals, toothed, lilac, anthers 10; fruit, about 13 to 20 mm long, 3- to 8-locular, a
subglobose to ellipsoid, fleshy, yellow drupe, the endocarp thick, bony; seeds, 1 (or 2)
per locule, laterally compressed, the testa crustaceous. Planted ornamental and spon-
taneous on coastal strip, in strip-mined area on plateau, near the Topside Oval, and on
escarpment slopes near phosphate processing plant. 2, 3(58732), 4(122N), 5(74), 6, 7.

Sandoricum koetjape (Burm.f.) Merr. "santol" (Philippines)
syns. S. indicum Cav.; Melia koetjape (Burm.f.) Merr.

Recent introduction. Malesia. Extinct? Medium to rather bulky, somewhat
deciduous tree, up to 12 m or higher, with velvety-pubescent young branchlets and brown
to gray smooth or slightly flaky bark; leaves, spiralled, trifoliate; leaflets, 10 to 20 cm
long, oblong-ovate, acute, rounded at base, slightly asymmetric or not, velvety-pubescent
or leathery, withering yellow or red; inflorescences in many-flowered axillary panicles,
10 to 20 cm long; calyx, 5-lobed, pubescent; corolla, 5-petaled, the petals about 1 cm

170

long, linear, yellowish, fragrant; stamen tube elongate, anthers 10, stigma 5-lobed; fruit,
8 to 10 cm across, a globose, fleshy drupe with velvety-pubescent -skin, usually 5-celled,
1 seed to each cell; pulp, brownish-white, acid, edible; seeds, large, inedible. Small
recently planted seedling growing in Filipino home garden at Location in 1980. No
longer present in 1981. 5.

MORACEAE (Mulberry Family)

Artocarpus altilis (Park.) Fosb. "breadfruit"
deme (N); te mai (K); mei (T)
syns. A. incisus (Thunb.) L.f.; A. communis Forst.

Aboriginal introduction. Malayo-Pacific. Common. Medium to large, round-
topped tree, 10 to 20 m or higher, with thick milky sap; leaves, 30 to 60 cm (rarely up
to 1 m) long and 20 to 40 (rarely up to 65 cm) cm wide, alternate, deeply pinnately S- to
7-lobed or -incised, lobes long-acuminate, oblong, acute, or in some varieties entire or
shallowly divided with 2 or 4 short triangular lobes, base broadly cuneate or obtuse, dark
green above, paler beneath, thick-coriaceous, glabrous above, sometimes hairy on the
nerves beneath or on both surfaces; midrib raised strongly beneath; petioles, 3 to 5 cm
long, stout; stipules, 10 to 25 cm long, dorsally pilose; male inflorescences, 7 to 30 cm
long and 1.5 to 4 cm wide, solitary, axillary, dense, yellow, somewhat spongy, cylindri-
clavate (somewhat club-shaped), drooping spikelike clusters on stout peduncles 3 to 8 cm
long; female inflorescences, solitary, axillary, globose to rounded-oblong or ellipsoid
headlike clusters, 8 to 10 cm long and 5 to 7 cm wide, which develop into fruits, stiffly
upright on stout peduncles 4 to 8 cm long; fruit, a syncarp, 10 to 30 cm in diameter,
globose or rounded-oblong, weighing up to 6 kg, studded with slightly conic or pyrami-
dal, hexagonal carpel apices, yellowish-green to brownish; inner flesh yellowish-white to
creamy, usually seedless, but sometimes with seeds, about 2.5 cm long. Planted staple
tree crop on coastal strip. Fruit cooked as a staple food; sap (denda) used as an adhesive
for caulking canoes and a chewing gum; leaves used for wrapping food for cooking, for
parcelization of fresh food, and as plates; medicine for curing ear aches made by
crushing juice out of tender meristems. Two main cultivars of A. altilis are deme and
modenewe or modenawe ("modern way"). Important staple food in Kiribati, Tuvalu. and
other atoll countries, and an important supplementary staple in other areas of the Pacific.
Ji, | Shea) y Son Te

171

Artocarpus heterophyllus Lam. "jakfruit", "jackfruit"
te mai rekereke (K)
syns. A. integrifolia sensu L.f.; A. integer sensu (Thunb.) Merr.

Pre-World War II introduction. Indomalaysia. Rare. Medium tree, 10 to 20 m
high, with thick milky sap and young twigs with many long, rigid hairs which are often
hooked at the apex; leaves, 5 to 25 cm long and 3 to 12 cm wide, alternate, oval or
oblong, entire (sometimes 3-lobed on young trees), acuminate, acute or obtuse, base
cuneate or obtuse, non-decurrent, dark-green, coriaceous, with stiff hairs; petioles, about
3.5 cm long; stipules, 1.5 to 8.5 cm long; male inflorescences, axillary or terminal
spikelike clusters, 2.5 to 10 cm long and 1 to 3 cm in diameter, ellipsoid-clavate, terete,
yellow or greenish yellow; female inflorescences, in globose or oblong heads which turn
into fruit and which are borne on the trunk or older branches (cauliflorous or ramiflor-
ous); fruit stalk 7 to 12 cm long; fruit, a syncarp, usually 30 to 40 cm long but oc-
casionally up to 60 to 90 cm long and about half as wide, oblong cylindric, golden
yellow to yellow-green, weighing up to 20 kg, rind studded with hexagonal bluntly conic
carpel apices; inner flesh pulpy, waxy, whitish-yellow, acid or sweetish or fetid when
over-ripe; seeds, 2 to 3 cm long, rounded, brown, enclosed in a slippery pulpy jacket.
Planted fruit tree reported present by Burges in 1933; young tree seen in Topside
Workshop food gardens in 1987. Ripe fruit eaten; immature fruit cooked as a supplemen-
tary staple in curries by Indians. A. integrifolia and A. integer are incorrect botanical
names for the jakfruit and according to Corner (in Gard. Bull. Straits Settlem. 10:56-81,
1939), A. integer refers to a separate species, the champedak which is not present in
Nauru, and A. integrifolia is an illegitimate name (Smith, 1981). 2,6.

Artocarpus mariannensis Trec. "Marianas breadfruit"
damenkamor (N); te mai kora (K); matua mei (T)

Aboriginal introduction? Micronesia. Occasional. Tree similar to A. altilis; leaves
smaller, 10 to 30 cm long, half as wide, broadly obovate to broadly elliptic, acute to
acuminate, entire to variously lobed in the upper part, but cutting usually not more than
halfway to the midrib, usually less, with some leaves often quite entire, base cuneate,
glabrous and somewhat shiny above, brown hairs on the veins and midrib beneath;
petiole, up to 4.5 cm long, usually half this length; male spike, up to about 8 to 10 cm
long; fruit, rather small, shortly cylindric, usually with several large seeds and somewhat
scanty pulp. Planted staple fruit tree. Same uses as for A. altilis, but fruit of A. marian-
nensis eaten raw and cooked. 3(58755), 5, 6, 7.

Ficus benghalensis L. "banyan", "Indian banyan", "east Indian fig", "Vada tree"
Recent introduction. India. Occasional. Large spreading tree, up to 20 m or

higher and often several times as wide, with milky latex and massive pillar roots,
descending from branches, which extend the tree laterally, sometimes almost indefinitely;

172

leaves, 10 to 30 cm long and 5 to 20 cm wide, elliptic to ovate, obtuse, base usually
cordate, glabrous (downy when young), leathery, with intercostal venation, the lateral
nerves in 5 to 7 distinctly spaced pairs, the basal nerves elongate in 2 to 4 pairs; petioles,
1.5 to 7 cm long, not articulate to lamina; male inflorescences disperse; female inflores-
cences similar to fig (gall); fruit, a fig about 14 to 25 mm in diameter, axillary, paired,
sessile or the body pedicellate, depressed-globose, orange-red, edible, the basal bracts
well-developed, 10 to 14 by 3 to 7 mm; seeds, small, smooth. Planted ornamental tree on
coastal strip; common on golf course. 3(58749), 5(53), 6, 7.

Ficus elastica Roxb. "Indian rubber tree", " rubber plant", "Indian rubber fig"

Recent introduction. India to Nepal and Malaya, perhaps southward to Java.
Occasional. A large spreading, fast-growing banyan tree, up to 35 m high (usually much
smaller as an ornamental), with smooth gray bark, copious aerial roots and milky latex;
leaves, 10 to 40 cm long and 4 to 22 cm wide, elliptic to subobovate, short-acuminate,
thick, stiff, leathery, glabrous, dark-green and shiny above, paler beneath, primary lateral
nerves usually in 15 to 22 pairs, the secondary lateral nerves almost as prominent as the
primary, leaves in bud enclosed in pointed, rolled, rosy, caducous sheaths; petioles, 4 to
12 cm long, not articulate to lamina; stipules large and conspicuous, pink to red; male
flowers, disperse; figs, up to 12 by 9 mm, in pairs, short-ellipsoid or oblong, greenish-
yellow, with a short thick peduncle usually 3 to 5 mm long and 4 to 6 mm thick, the
basal bracts 3, early caducous. Planted ornamental tree or house plant in its juvenile
fOGM 5.6, 7

Ficus prolixa Forst.f. var. carolinensis (Warb.) Fosb. "native banyan"
eaeo, eyayo, yayo (N)

Indigenous. Micronesia. Very abundant. Medium to large tree, 2 to 20 m_ high,
with many aerial roots descending from branches; leaves, 8 to 16 cm long and 6 to 8 cm
wide, alternate elliptic-oblong or oblanceolate, short acuminate, base cordate or subcor-
date, glabrous, primary nerves in 5 to 10 pairs; petioles, 0.8 to 3 cm long; male
inflorescences ostiolar and disperse; figs, 5 to 10 mm in diameter, axillary, mostly
solitary, subglobose, ripening white to pink and purple-black, with 3 slightly coherent
rounded basal bracts, sessile or borne on peduncles up to 3 mm long. Common on
plateau in areas of unmined forest, in older strip-mined areas, and on the escarpment and
coastal strip, primarily on coral-limestone pinnacles and outcrops. F. prolixa seems to be
one of the only species capable of long-term colonization of residual pinnacles in strip-
mined areas and could become dominant in the disclimax vegetation. Berry-like fruit
(moduru) eaten cooked and mixed with boiled sap (toddy or karawai) from the coconut
flower spathe (kamerara) to make a dish known as dedangan or dedengan which can
keep for two to four weeks, and, if cooked and dried in the sun, will keep for years if
stored in a dry place; sap used as chewing gum (ikumi, kumi). 1, 3(58663), 5(19), 6, 7.

173

w "

Ficus tinctoria Forst.f. var. neo-ebudarum (Summerh.) Fosb. "Dyer’s fig", "native fig"
debero (N); te bero (K); felo (T)

Recent introduction. S. E. Asia to Polynesia and Micronesia. Rare. Small
dioecious tree, up to 8 m high, commonly with long aerial roots or prop-roots; leaves,
about 8 to 15 cm long and 5 to 8 cm wide, ovate, with a slightly asymmetric base, dark
green withering to a bright yellow with purplish veins, lateral nerves in 3 to 9 pairs;
tepals, white; figs, about 12 mm in diameter, globose, yellowish turning dull reddish,
borne on peduncles about 10 mm long. Planted food tree in Rev. J. Aigimea’s garden.
Fruit eaten cooked; fruit also cooked and mashed and mixed with boiled coconut syrup
(kamirara, kamerara) to make a pudding (dedengan). Plant introduced from Kiribati,
where it is a supplementary staple in many areas and a major staple in the drier islands of
southern Kiribati. Use learned by some Nauruans from I-Kiribati, but recipe essentially
the same as used by Nauruans for F. prolixa. 5(104), 6, 7(27808).

Ficus sp.

Recent introduction. Planted ornamental. Rare. 5 (197).

MORINGACEAE (Moringa Family)

Moringa oleifera Lam. "horseradish tree", “drumstick tree", "saijan", "seijan" (Hindi),
malunggay (P)

syns. Guilandina moringa L.; M. moringa (L.) Millsp.; M. pterygosperma
Gaertn.

Recent introduction. India. Occasional to common. Small soft-wooded tree up to
10 m high, with thick stems, corky, gummy bark and pungent roots; leaves, 20 to 60 cm
long, alternate, bi-tripinnate, the pinnae and leaflets opposite; leaflets, 1 to 3 cm long and
5 to 18 mm wide, oval to obovate, sometimes faintly notches, somewhat grayish-green;
inflorescences, shorter than the leaves, axillary, many-flowered, paniculate; calyx, 5-
lobed, lobes imbricate, subequal, spreading or reflexed, white; corolla, about 1.5 to 2.5
cm across, 5-petaled, petals imbricate, shortly connate at base, unequal, the lowermost
the largest, erect, the other reflexed, white, greenish proximally, fragrant; stamens 10, 5
with yellow anthers, perfect, epipetalous, the filaments free; fruit, 15 to 4 cm long, a 3-
valved, pendent, podlike capsule, elongated, beaked, subtorulose, the valves thick,
spongy with 9 blunt ribs; seeds, many, about 10 mm in diameter, 3-angled, winged at
angles. Food plant in Indian home gardens on Meneng Terrace and in Filipino Gardens at
Location. Nutritious leaves, fruit, and flowers cooked as vegetables by resident Filipino
and Indian families. Very common food tree planted by Indians in Fiji. 5, 6, 7.

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MYRTACEAE (Myrtle Family)

Eucalyptus sp. "eucalyptus", "gum tree"

Recent introduction. Australia. Rare. Tree with peeling bark; leaves, aromatic;
flowers, petals absent, with numerous showy stamens; fruit, a woody capsule, opening by
slits; seeds, small and numerous. Planted ornamental tree in Nauruan houseyard garden.
6.

Pimenta dioica (L.) Merr. "allspice"
syn. P. officinalis Lindl.; Myrtus pimenta L.; M. dioica L.

Recent introduction. C. America and W. Indies. Rare. Small to medium tree, 6 to
12 m high, with nearly smooth silvery-brown bark and brittle wood; leaves, 5 to 20 cm
long and 2 to 8 cm wide, opposite, narrowly elliptic to elliptic-oblong, bluntly acute,
obtuse to rounded at base, thick, coriaceous, glabrous, dark green above and paler and
gland-dotted beneath, pinnately veined, mid-rib impressed above and prominent below,
highly aromatic; petioles, usually 1 to 1.5 cm long; inflorescences, many-flowered,
subterminal axillary (in axils of the upper leaves) cymose panicles, 5 to 15 cm long;
pedicels, about 1 cm long, pubescent, with small brownish bracteoles; calyx, 4-sepaled
campanulate with spreading persistent lobes about 2 mm long; corolla, about 8 to 10 mm
in diameter, 4-petaled, petals, about 4 mm long, rounded, reflexed, white, spreading,
caducous; stamens numerous, free; fruit, 5 to 7 mm in diameter, a small subglobose
berry, dark purple when mature, with sweet pulpy mesocarp; seeds, usually 2, sub-
globose with a spiral embryo. Planted ornamental seedling in home garden. 6.

Psidium guajava L. "guava"
kuwawa (N); tekuwawa (K); kuava (T)
syn. P. pomiferum L.

Recent introduction? Trop. America. Common. Shrub or small, shallow-rooted
tree, 2 to 10 m high, with smooth green, light reddish-brown or copper-colored bark,
wide-spreading branches, and pubescent 4-angled or -winged young branches, often
producing suckers from roots near base of trunk; leaves 5 to 15 cm long and 3 to 7 cm
wide, opposite, ovate-elliptic or oblong-elliptic, acute-acuminate, base obtuse to rounded,
dull green, pubescent beneath, often rather brittle, lateral veins 7 to 20 per side, slightly
sunken above, prominent below; petioles, 2 to 10 mm long; inflorescences, axillary,
solitary or in 2- to 3-flowered cymes; peduncle, about | to 2 cm long, pubescent; calyx,
4- to 6-lobed, campanulate, splitting irregularly, lobes 1 to 1.5 cm long, reflexed,
pubescent, persistent; corolla, about 2.5 to 3 cm across, petals 4 or 5, elliptic to obovate,
slightly concave, reflexed, white, slightly fragrant, fugaceous; stamens, numerous (about

175

200 to 250), white, about as long as petals, with yellow anthers; style, 1.5 to 2 cm long,
filiform, greenish-yellow, exserted above stamens; fruit, 3 to 12 cm long, globose, ovoid
or pyriform, surmounted with remnants of calyx lobes, shining pale green, whitish
yellow or faintly pink when ripe, pulpy; pulp, granular-juicy, sweet-sour, light pink to
reddish-pink or cream-colored, aromatic; seeds, many, 3 to 5 mm long, bony, reniform,
yellowish to light brown, embedded in pulp. Occasionally planted or protected in home
gardens; spontaneous on coastal strip and locally abundant in unmined forest on plateau
and in mined areas. Wood an excellent firewood and makes good fishing poles; leaves
used to treat diarrhea; ripe fruit eaten and made into jams. 3(58650, 58767), 4(105N),
5(94), 6, 7, 8.

Syzygium malaccense (L.) Merr. & Perry "Malay apple", "mountain apple"
apolo Solomona ("Solomon Is. apple")(T)
syns. Eugenia malaccensis L.; Carophyllus malaccensis (L.) Stokes; Jambosa
malaccensis (L.) DC.

Recent introduction. S. E. Asia. Rare. Medium tree, 6 to 15 m tall; leaves, 10 to
38 cm long and 5 to 22 cm wide; opposite, elliptic to oblong-obovate, bluntly acuminate,
base obtuse to acute, then abruptly decurrent on petiole, glossy, glabrous, paler beneath,
obscurely punctate, thick-coriaceous, pinnately nerved, lateral nerves submarginally loop-
connected 4 to 7 mm from margin; petiole, 8 to 15 mm long, stout, in young leaves
reddish; inflorescences, dense few-flowered cymes, about 5 cm long, on defoliate older
branches and on trunk; pedicels, 1 cm long or less; calyx tube, 1.5 to 2 cm long,
turbinate, 4-lobed, lobes rounded, wider than long; corolla, 4 to 7 cm in diameter across
expanded stamens, petals 4, 8 to 18 mm long, obovate, concave, red, fugaceous;
stamens, many (about 125), 1 to 3.5 cm long, erect, red to vivid crimson-pink; anthers,
small yellowish; style, nearly 2 cm long, dark red; fruit, about 6 to 8 cm long, obovoid
or pyriform, crowned by inflexed calyx-lobes, glossy and slightly waxy in appearance,
crisp-succulent, light green maturing to red or greenish-white or striped or mottled,
usually 1-seeded; flesh, white, watery; seeds, about 2 cm across. Planted fruit tree
seedling in Tuvaluan garden at Location. Common aboriginal introduction throughout
most of high-island Melanesia and Polynesia, where the fruit is eaten. 5, 6.

NYMPHAEACEAE (Waterlily Family)
Nymphaea sp. "water lily"

Recent introduction. E. and S. Africa and Madagascar. Rare. Aquatic perennial
herb with submerged rhizomes; leaves, peltate, floating, long-petiolate, arising from the
rhizome; flowers, cup-like, solitary, regular, bisexual; calyx, 4-sepaled, the sepals,
nearly free, intergrading into the petals; corolla, many-petaled, petals, overlapping,
showy, variously colored from rich blue, pink, yellow to creamy-white; stamens,

176

numerous, anthers introrse, opening by slits; carpels united into a multilocular ovary
bearing a flat radiate stigma; seeds, hard, operculate, arillate. Planted ornamental in
water tanks and in small household fish ponds. 3(58684), 4, 5(251).

NYCTAGINACEAE (Four-o’clock Family)

Bougainvillea glabra Choisy "bougainvillea", "red bougainvillea"
tsita, tsitta (N); te akanta (K); akanta (T)

Recent introduction. Brazil. Stout, woody scrambling or climbing vinelike shrub,
with spiny, but less thorny than B. spectabilis; leaves, alternate, elliptic, acute at both
ends, sparsely puberulent on both surfaces; inflorescences, axillary, nearly 2.5 cm long,
in threes within 3 involucral bracts; bracts, ovate, commonly red, but variable in color,
persistent; perianth S-lobed, swollen, 5-angled below constriction, yellow to yellowish-
white; fruit, 7 to 13 mm long, a glabrous anthocarp. Occasional. Planted ornamental.
Flowers used by I-Kiribati in garlands. 3(58726), 5, 6, 7.

Bougainvillea Spectabilis Willd. "bougainvillea", "purple bougainvillea"
tsita, tsitta (N); te akanta (K)

Recent introduction. Brazil or Peru. Occasional. Stout, woody scrambling or
climbing vinelike shrub with stout axillary spines and the young growth finely pubescent;
leaves, alternate, ovate, acute at both ends, tomentose beneath and often above; inflores-
cences, axillary, nearly 2.5 cm long, in threes within 3 involucral bracts; bracts, ovate,
rich purplish-magenta to rosy pink, less commonly orange, pale yellow or white, veiny,
persistent; perianth S-lobed, tubular, indistinctly angled, yellow to yellowish white;
stamens usually 8, unequal; fruit, 11 to 14 mm long, a 5-angled anthocarp, densely
pilose, not sticky. Planted ornamental. Flowers used by I Kiribati in garlands. 5, 6, 7.

Mirabilis jalapa L. "four-o’clock", "marvel of Peru", "false jalap"
teoua, teowa (N); te aoaaua, te awaava (K)

Recent introduction? Mexico. Occasional. Erect, glabrous, somewhat succulent
annual herb, up to 1 m tall, with tuberous roots; leaves, 5 to 10 cm long, opposite,
ovate-lanceolate, apex long attenuate-acuminate, base broadly cordate; petioles up to 2.5
cm long; inflorescences, bisexual, in terminal clusters; perianth, 2.5 to 5 cm long, 5-
parted, funnel- or trumpet-shaped, red, purple, yellow or various, fragrant, opening late
in the afternoon; stamens, 5; fruit, 8 mm long, an anthocarp, hard, ribbed, black, not
prickly or sticky. Planted ornamental; spontaneous in some places on coastal strip.
Flowers used in garlands. 3(58784), 5, 6, 7.

WIT

Pisonia grandis R. Br. "pisonia"
yangis, yangys, yangits (N); te buka (K); puka, puka vai (T)
syns. P. alba Span.

Indigenous. Indopacific. Uncommon. Medium to large, soft-wooded tree with a
stocky trunk and brittle, often wind-broken, branches; leaves, up to 20 cm or longer and
10 cm wide, opposite, elliptic or ovate, obtuse to acutish, base acute, thin, pale green,
glabrous, pubescent along the midrib beneath, with numerous lateral veins; petiole, up to
5 cm or longer; inflorescences, small, in terminal cymose clusters; perianth 4- to 6-
lobed, fragrant; stamens 5 to 13, unequal; style present; fruit, about 1 cm long, an
anthocarp, fusiform or short-cylindric, with rows of short spines, glandular, very sticky.
Tree in unmined plateau forest bordering escarpment above Anibare Bay and on unmined
residual rocky limestone outcrops. Very brittle wood used in past as an inferior fuel;
most important roosting habitat for noddy birds which are an important ceremonial food.
GN).

OLEACEAE (Olive Family)

Jasminum multiflorum (Burm.f.) Andr. "jasmine", "star jasmine"
rimone (N)
syns. Nyctanthes multiflora Burm.f.; J. pubescens Willd.

Recent introduction. India. Rare. Pubescent spreading, sometimes climbing shrub,
1 to 2 m high, with copiously pilose or tomentose branchlets; leaves, 2 to 6 cm long and
1.5 to 4 cm wide, opposite, ovate, acute, rounded or truncate to subcordate or cordate at
base, pubescent on both surfaces or sometimes glabrate above; petiole, short; inflorescen-
ces, short, compact terminal clusters; calyx, 4- to 8-lobed, lobes, 8 to 18 mm long,
subulate, densely pubescent with spreading yellowish hairs 8 to 18 mm long; corolla,
about 2.5 cm across, tube 19 to 28 mm long, 4- to 9- lobed, lobes, 12 to 20 mm long,
acute, white, tube greenish, fragrant to odorless; fruit, a berry, 2-lobed, 2-locular and 2-
seeded. Planted ornamental. 5, 6.

Jasminum sambac (L.) Ait. "jasmine", "Arabian jasmine", "pikake" (Hawaii)
rimone (N); te bitati (K); pitasi (T); sampagita (P)
syn. Nyctanthes sambac L.

Pre-World War II introduction. India. Common. Pubescent, or somewhat downy,
scrambling or sometimes climbing (scandent) shrub, up to 1 m or higher, with angular
branchlets; leaves, 2 to 10 cm long and 1.5 to 6 cm wide, opposite, elliptic to ovate,
acute or obtuse, base rounded or cuneate, nearly glabrous, with evident pinnate nerves,

178

blades with axillary hair tufts beneath; petiole, short, arched, pubescent; inflorescences in
few-flowered axillary clusters; calyx, about 2.5 cm across, 4- to_10-lobed, lobes about 6
to 7 mm long, linear, ciliate or glabrous; corolla, tubular, often double, tube about 15
mm long, the lobes, oblong to nearly orbicular, obtuse, as long as tube, white, fragrant;
fruit a berry. Planted ornamental, especially in I-Kiribati and Tuvaluan gardens at
Location; also in Nauruan gardens. Flowers used in garlands and for scenting oil.
3(58719), 5(18), 6, 7.

ONAGRACEAE (Evening Primrose Family)

Ludwigia octovalvis (Jacq.) Raven "swamp primrose", "willow primrose"
te mam (K); titania? (T)
syns. Jussiaea suffruticosa L.; Oenothera octovalvis Jacq.

Recent introduction. Pantropical. Rare. Erect, coarse, slender, branching
perennial herb, 50 cm to 2 m high, with sparingly to densely pubescent and longitudinal-
ly-grooved stems which are somewhat woody at the base; leaves, 3 to 15 cm long and
0.5 to 2.8 cm wide, alternate, linear-lanceolate to ovate, acuminate or acute, base cuneate
to acute, with 11 to 20 pairs of lateral nerves; petioles, up to 1 cm long; inflorescences,
solitary, axillary, 2.5 to 3 cm across, borne on short pedicels, about 1.2 cm long with 2
small bracts near top; calyx, 4-lobed, lobes, 6 to 15 mm long and 1 to 7.5 mm wide,
ovate-lanceolate to linear, persistent, sometimes red-tinged; corolla, 4-petaled, tubular,
tube linear, about 2 cm long, petals, 5 to 17 mm long and 4 to 17 mm wide, broadly
obovate or cuneate, emarginate (notched), yellow, caducous; stamens 8, 1 to 4 mm long
(excluding anthers), alternately unequal; fruit, 1.7 to 6.5 cm long and up to 5 to 7 mm
wide, a longitudinally 8- to 10-ribbed cylindrical 4-celled capsule with sepals persistent at
apex, late dehiscent along the sides; seeds, 0.6 to 0.75 mm long, several-rowed in each
cell, free, not embedded in endocarp. Weed in swampy area bordering mangroves in
Meneng. 4 (150).

OXALIDACEAE (Wood Sorrel Family)

Averrhoa bilimbi L. "belimbi"

Recent introduction. Malaya to India. Rare. Small tree, 3 to 10 m high; leaves, up
to 60 cm long, odd-pinnate with 15 to 41 pairs of leaflets; leaflets, 3 to 11 cm long and
1.5 to 3 cm wide, oblanceolate, acute-acuminate, a few proximal ones smaller than the
distal leaflets; inflorescences, cauliflorous or ramiflorous, borne in fascicles on branches
or trunk; calyx, 5-sepaled, sepals imbricate, yellowish-red to purple; corolla, 5-petaled,

179

petals, 1 to 2 cm long, dark red, glabrous within, not cohesive; stamens 10, all with
anthers; fruit, 5 to 10 cm long and 2 to 5 cm across, oblong-cylindric (cucumber-
shaped), smooth, subterete, faintly 5-angled, yellowish-green, acid, crisp; seeds, 6 to 8
mm long and 4 to 6 mm across, embedded in pulp, exarillate. Three large trees planted
near Indian homes on Meneng Terrace. Fruit eaten ripe; green and ripe fruit made into
pickles by Indian families. 7.

Oxalis corniculata L. "yellow wood-sorrel", "creeping wood-sorrel"
syn. O. repens Thunb.

Pre-World War I introduction; reported by Schumann & Lauterbach (1901) as
collected by Finsch. Paleotropical and paleosubtropical; now cosmopolitan. Rare.
Procumbent, creeping, freely-branching, pubescent perennial herb, ascending to 10 to 20
em high or decumbent, with several stems radiating from the main root, the individual
stems rooting at the nodes and rarely exceeding 50 cm in length; leaves, usually less than
2.5 cm across, alternate, palmately trifoliate, clover-like, the terminal leaflet much larger
than the lateral leaflets; leaflets, 4 to 20 mm long and wide, obocordate, deeply notched
at apex, base obtuse, nearly pubescent; stipules, up to 3 mm long, united to base at
petiole, inconspicuous; petioles, 1 to 8 cm long; inflorescences, 1- to 6-flowered umbels
on axillary peduncles, 2 to 7 cm long; pedicels, 4 to 15 mm long, slender; bracts and
bracteoles, 0.5 to 3 mm long, deltoid-linear; calyx, 5-sepaled, sepals, 2.5 to 6 mm long,
lanceolate to narrowly ovate; corolla, 5-petaled, petals, 4 to 8 mm long, oblanceolate,
yellow; stamens 10, in two series; fruit, 5 to 20 mm long and 2 to 4 mm across, an
oblong, 5-locular, 5-angled, acute-tipped, pubescent capsule, opening by longitudinal
valves; seeds, 5 to 10 per locule, each about 1.5 mm long, compressed, cross-ridged or
wrinkled, brown. Weed at MQ 40 Command Ridge. 1, 6(163).

PASSIFLORACEAE (Passion Flower Family)

Passiflora coccinea Aubl. "scarlet passion flower"

Recent introduction. Trop. America. Rare. Climbing, tendril-bearing vine; leaves,
up to 15 cm or longer, simple, ovate, toothed, woolly beneath; petioles, purplish, with
no glands or 2 at base; inflorescences with striking scarlet petals, yeilowish on reverse,
with filaments white at proximally, shading through pink to purple distally; fruit, about 5
cm in diameter, orange or yellow with green lines and spots, edible. Planted ornamental.
6.

180

Passiflora edulis Sims "passionfruit"

Recent introduction. Trop. America. Rare. Vigorous woody perennial climbing
vine, up to 15 m long, with glabrous, grooved stems and axillary, spirally coiled tendrils;
leaves, 10 to 15 cm by 12 to 25 cm, alternate, broadly ovate in outline, deeply palmately
3-lobed, lobes ovate-oblong, sharply acute to acuminate, sinus rounded, base cordate,
serrate; petioles, 2 to 5 cm long, usually glabrous, grooved on upper surface, with 2
conspicuous glands near base of blade; stipules, about 1 cm long, lanceolate; inflorescen-
ces, axillary, 7 to 10 cm in diameter; peduncle, 2 to 5 cm long, triangular; bracts 3, 1 to
3 long and 1 to 2 cm wide, near apex of peduncle, leafy, ovate to lanceolate, serrate-
glandular; calyx, 5-lobed, tubular at base, sepals, 2 to 3 cm long and 1 to 2 cm wide,
ovate-oblong, spreading, reflexed, white above, yellowish-green below, spongy, fleshy,
with thorn-like appendage near tip and 0 to 4 glands on margins; corolla, 5-petaled,
petals, 2.5 to 3 cm long and 0.5 to 1 cm wide, elliptic, free, white, alternating with
calyx, inserted on throat of calyx; corona of 2 outer rows of wavy, threadlike, radiating
filaments, 2 to 3 cm long, white distally and purple or pink proximally, with several
rows of short, purple-tipped papillae; stamens 5, filaments united in a tube around
gynophore for about 1 cm then widely parted for 1 cm; anthers, 1 to 1.5 cm long,
versatile, transverse, 2-celled, pale yellow, hanging downwards below level of ovary;
fruit, 4 to 6 cm long, subglobose or ellipsoid, yellow or purple depending on the variety
or form, rind hard, endocarp white; seeds, many, about 5 by 3 mm, compressed, testa
blackish, 3-toothed at base, attached to peg-like funiculi on the ovary wall and sur-
rounded by yellowish-orange, aromatic pulpy and juicy aril with a tart but pleasing
flavor; seeds, pulpy aril and juice edible. Two small seedlings in home garden. Sup-
plementary food plant in many parts of the Pacific and currently or formally an important
cash crop in Niue, Fiji, W. Samoa, Hawaii and Papua New Guinea. 6.

Passiflora foetida L. var. hispida (DC.) Killip "stinking passion flower",
"love-in-a-mist" oatamo, watamo (N); te biku (K)
syn. P. hispida DC. ex Triana & Planch.

Pre-World War II introduction. Trop. America. Occasional. Creeping or climbing
perennial herbaceous vine, 1.5 to 5 m long, with weak, densely long-hispid, longitudinal-
ly-grooved stems, axillary coiled tendrils and fetid (bad smelling) foliage; leaves, 4 to 12
cm long and almost equally wide, simple, ovate or ovate-cordate in outline; palmately 3-
lobed, the lobing shallow to halfway to the mid-vein, lobe apices acute, base subcordate,
margins unevenly dentate or entire, both surfaces hispid-hirsute; petioles, 2 to 6 cm long,
pubescent; stipules, laciniate, the divisions filiform; inflorescences, 2.5 to 5 cm across,
solitary, axillary; pedicels 2 to 7 cm long; involucral bracts prominently 2- or 3-bipin-
natifid; calyx, S-lobed, tubular, sepals oblong, mucronate, inside white; corolla, 5-
petaled, petals, 1.5 to 2.5 cm long, oblong, white; stamens 5, pale green, surrounded by
a ring-like corona of filaments, 9 to 15 mm long, white distally and purple to rich-blue
proximally; fruit, 1.5 to 2.5 cm in diameter, subglobose to ovoid, yellowish-orange to
red-orange, often with faint greenish vertical lines, leathery, thin-walled, surrounded by

181

pinnatifid involucral bracts, tardily dehiscent; seeds, many, surrounded by scanty, slimy,
sweetly-tart yellowish-orange pulpy aril, edible. Weed, generally on plateau, but also on
escarpment and coastal strip, along roadsides and in ruderal habitats. Scanty pulpy aril
and seeds eaten by children. 2, 3(58592), 4(117N), 5(54), 6, 7, 8(9581).

PIPERACEAE (Pepper Family)

Peperomia obtusifolia (L.) A. Dietr. "jade plant", "baby rubber plant"
syn. Piper obtusifolia L.

Recent introduction. W. Indies and Florida. Rare. Erect or decumbent, spreading,
branching, succulent perennial herb, up to 20 cm high, with reddish stems; leaves, 3 to
10 cm long and 2 to 6 cm wide, alternate, oval to inverted-ovate, obtuse, apex sometimes
notched, base decurrent to cuneate, glabrous, shiny, dark green above pale below,
succulent; petioles, 1 to 3 cm long, reddish; inflorescences, erect spikes, 5 to 15 cm or
longer, green, bearing minute sessile flowers; stamens 2. Pot plant. 3(58691), 6, 7.

Peperomia pellucida (L.) HBK. "peperomia"
syns. P. pellucidum L.; P. lineata Miq. ex Yuncker

Recent introduction. Trop. America. Rare. Pale, erect or reclining, fleshy,
glabrous, weak-stemmed perennial herb, 15 to 30 cm high, with branched stems; leaves,
1 to 3.5 long, alternate, ovate-cordate, acute to acuminate, base rounded to cordate or
cordate-truncate, glabrous, shiny above dull beneath; petioles, 3 to 12 mm _ long;
inflorescences, slender leaf-opposed or terminal spikes, 2 to 5 cm long, bearing minute,
well-separated green sessile flowers; calyx and corolla absent; stamens 2; ovary sub-
tended by a minute bract; fruit, less than 1 mm in diameter, a green globose, rostellate
(beaked) drupe with longitudinal ribs, containing a single warty seed. Weed of pot plants
in homes. 5(7), 6, 8.

POLYGALACEAE (Polygala Family)
Polygala paniculata L.

Recent introduction. Trop. America. Rare. Slender, erect, profusely-branching
annual herb, 10 to 60 cm tall, with glandular-pubescent stems and a tuberous, yellow,
strongly aromatic tap-root with the scent of peppermint or wintergreen; leaves, 1 to 2.5
cm long and 2 to 4 mm wide, linear-lanceolate, in whorls of 5 caducous leaves below
and spirally arranged above; inflorescences, slender terminal racemes, 2 to 15 cm long,

182

bearing numerous crowded small flowers, 2 to 3 mm long; calyx, 5-parted, the two inner
sepals resembling wings and much larger than the others and resembling petals; corolla,
3-petaled, the upper pair rather narrow, the lower (keel) boat-shaped; petals and inner
sepals at first white, becoming purplish or pinkish with age; stamens 8; fruit, about 2 to
3 mm long, oblong, glabrous, not winged; seeds, about 1.5 mm long, oval-oblong, black
with tiny white hairs and a 2-lobed whitish caruncle. Weed along Topside running track
and in other waste places. 5, 6(203), 7.

POLYGONACEAE (Buckwheat Family)

Antigonon leptopus Hook. & Arn. "Mexican creeper", "mountain rose",
"confederate vine", "chain of love", “love vine", "hearts on a chain", "coral vine"

Recent introduction. Mexico. Occasional. Perennial herbaceous climbing to
scrambling vine with slender glabrous stems, axillary tendrils and tuberous roots; leaves,
5 to 12 cm long, alternate, broadly ovate, cordate, apiculate to acuminate, net-veined,
wrinkled and wavy-edged; petioles, | to 5 cm long; inflorescences, long, slender, loose,
6- to 15-flowered racemes which end in branched tendrils; pedicels about 1 cm long;
flowers about 13 mm long; tepals 5, rose pink or white with a darker center, becoming
membranous, greenish, reticulate-veined; fruit, up to 1.5 cm long, a triangular, brownish
achene; seed, longitudinally grooved. Planted ornamental and spontaneous in waste places
and ruderal habitats. 3(58679), 5(59), 6, 7, 8(9562).

Coccoloba uvifera (L.) Jacq. "sea grape"
syn. Polygonum uvifera L.

Recent introduction. Trop. America. Rare. Small glabrous tree, up to 8 m or
higher, with thick branchlets and spreading branches; leaves, 10 to 20 cm across, usually
broader than long, alternate, suborbicular, cordate, firm, coriaceous, glossy, glabrous,
the midrib and lateral nerves reddish; inflorescences, erect spike-like racemes, about 15
cm long; individual flowers with 5 tepals, 8 stamens and 3 styles, greenish-yellow,
fragrant; fruit, about 1 cm across, a subglobose or pyriform, reddish berry-like drupe,
sweetish-acidic, astringent, edible. Planted ornamental near Buada Lagoon. 3(58787), 6.

183
PORTULACACEAE (Purslane Family)

Portulaca grandiflora Hook. "portulaca", "purslane", "pigface"

Recent introduction. Brazil. Rare. Decumbent, succulent, annual herb, up to 15
cm high, with tufts of white silky hairs at the nodes and just below the flowers; leaves,
12 to 35 mm long and 1 to 4 mm wide, narrowly cylindric or subterete; inflorescences,
terminal; calyx, 2-lobed, sepals, 5 to 12 mm long, connate; corolla, 2 to 3 cm across, 4-
to 6-petaled, petals, 12 to 30 mm long, pink, red, yellowish, white or striped; stamens,
many (40 to 75); fruit, about 5 mm in diameter, a membranous capsule, circumscissile
(dehiscent by a hemispherical lid). Planted ornamental. 3(58659), 5, 6, 7.

Portulaca oleracea L. “pig weed", "purslane", "wild purslane"
debois, doboiy (N); te boi (K); katuli (T)

Pre-World War II introduction. Europe. Occasional. Prostrate, spreading, fleshy,
glabrous herb, up to 30 cm high, branching at base with branches sometimes reddish;
leaves, 4 to 30 mm long and 3 to 12 mm wide, alternate or clustered at branch ends,
obovate, spathulate or oblong-cuneate, obtuse to slightly notched, base cuneate, fleshy,
flattened, dull green or reddish, subsessile; inflorescences, terminal, sessile, few-
flowered; calyx, 2-lobed, sepals connate; corolla, usually 5-petaled, petals yellow,
withering early; stamens 7 to 15; fruit, about 4 mm long, a membranous capsule,
circumscissile (dehiscent by a hemispherical lid); seeds, numerous, small, black, slightly
roughened. Weed in gardens and waste places, especially in sandy, hydromorphic soils of
the shores of Buada Lagoon. Cooked leaves and stems eaten after pounding and mixing
with coconut flower spathe syrup (kKamerara); important famine food during World War
II; plants fed to pigs; leaves and stems boiled with water being used to cure scabies.
3(58624, 58734), 5(4), 6, 7, 8.

RHAMNACEAE (Buckthorn Family)

Colubrina asiatica (L.) Brongn. "soapbush", "hoop withe"
ewongup (N)
syns. Ceanothus asiaticus L.; Ceanothus capsularis Forst.f.

Indigenous. Paleotropical. Common. Sprawling or climbing, much-branched shrub
or small tree, 1 to 5 m high; leaves, mostly 4 to 11 cm long and 1 to 6.5 cm wide,
alternate, ovate, acuminate, base rounded or subcordate, finely crenate-dentate, glossy
green, nearly glabrous, palmately 3-to S-nerved, the midrib branched upward; petioles, 1

184

to 3 cm long; inflorescences, small, axillary short-pedunculate, 3- to 7-flowered,
compact, cymose clusters; peduncles less than 3 mm long; pedicels, about 3 mm long,
slender; calyx, 5-sepaled, sepals acute, white to pale or dull yellow; corolla, 5-petaled,
petals hoodlike, green to yellowish-green or whitish; stamens 5, less than 1 mm long;
disc, broad, saucer-like, greenish-yellow to yellow-orange; fruit, 6 to 8 mm in diameter,
depressed-globose, 3-celled, green or brown; seeds 3 (1 per cell), nearly 5 mm long,
dark brown or black. Found in forests on plateau and on cliffs and slopes and at base of
escarpment. Rolled leaves used with flowers of other species in garlands. A traditional
source of soap in other Pacific islands, although not reportedly used for this purpose on
Nauru. 2, 3(58641), 4(113N), 5(69), 6, 7.

RHIZOPHORACEAE (Mangrove Family)

Bruguiera gymnorrhiza (L.) Lam. “brown mangrove"
etum, etam (N); te tongo, te tongo buangui (K)

syns. Rhizophora gymnorhiza L.; R. conjugata L.; Bruguiera gymnorhiza

Savigny; B. rheedii Bl.; B. eriopetala W. & Arn.; B. conjugata (L.) Merr.

Indigenous. Indopacific. Occasional. Medium tree, 3 to 15 m tall (in Nauru rarely
over 4 m), with an inconspicuously buttressed trunk, knee-like pneumatophores, dark
fissured trunk bark, and smooth-barked slightly nodose stems; leaves, 9 to 20 cm long
and 4 to 9 cm wide, opposite, elliptic-oblong, acute-acuminate at both ends, base
decurrent, thick-coriaceous, quite glabrous, somewhat glossy dark green; petioles, up to
4 cm or longer; stipules, to 3.5 cm long, linear-lanceolate, reddish, sheathing the new
leaves, caducous; inflorescences, solitary, axillary, somewhat nodding, up to 3 or 4.5 cm
across, on pedicels 1 cm long; calyx, 10- to 14-lobed, firm, glossy red to dull yellow or
yellow-green, lobes, about 1.5 cm long, slender-subulate or very narrowly-lanceolate, as
long as tube; corolla, 10- to 14-petaled, petals 3 to 5 cm long, narrowly oblong, notched
at apex, white fading to brown, each lobe bristly; stamens, many (or as few as 20), about
1 cm long; anthers, 4 to 5 mm long, linear; pollen dirty white; styles nearly 2 cm long,
pale green; fruit, 2 cm long and 1.5 cm thick, turbinate, crowned by calyx limb; seed
germinating on tree to form an elongated, somewhat extruded, cylindric radicle, reaching
15 to 25 cm long before falling. Localized in system of brackish lakes or lagoons near
base of escarpment in Meneng, Anabar and Anetan District (lake in Anabar known as
Araro); reportedly present in Buada Lagoon in the past. Strong wood excellent for house
construction; pre-germinated seed (fruit) eaten cooked, after scraping, drying in the sun
and then boiling; the Nauruan delicacy known as efum or etam, is prepared by mixing the
grated pre-germinated seed with coconut milk and then baking; skin of seed used to
prepare a black dye for traditional skirts (ridi). 2, 3(58746), 4(167N), 5(103), 6, 7.

185
ROSACEAE (Rose Family)

Filipendula rubra (J. Hill) B.L. Rob. "queen of the prairies"

Recent introduction. Central U.S. Hardy, clump-forming, perennial herb, up to 60
cm high; leaves, compound, up to 20 cm or longer; inflorescences, large, feather-like,
compact, many-flowered, irregular panicles, 15 to 30 cm wide, bearing many small
peach-pink flowers. Rare. Planted ornamental. 6.

Rosa damascena Mill. "damask rose"
dorot ("the rose")(N); te roti ("the rose")(K)

Recent introduction. W. Asia. Occasional. Erect, robust, aculeate shrub, up to 2
m high, with stems bearing numerous prickles or small thorns; leaves, pinnate, with 5 to
9 leaflets; leaflets, 7 by 5 cm, ovate-oblong, puberulent beneath on the midrib; inflores-
cences, clustered, 6- to 12-flowered corymbs, with bristly pedicels and receptacles;
calyx, 5-sepaled, imbricate in bud; corolla, 5-petaled (sometimes more), red or pink,
fragrant; stamens and ovaries numerous; hypanthium pilose within; fruit, a 1-seeded
achene included by the fleshy, colored fruitlike hypanthium. Planted ornamental. 5(80),
6.

RUBIACEAE (Coffee Family)

Aidia cochinensis Lour.
enga, enguh (N)
syns. Randia_ cochinchinensis (Lour.) Merr.; R. racemosa (Cav.) F.-Vill.;
Stylocoryna racemosa Cav.; Randia graeffei Reinecke; Stylocoryna den-
siflora (Wall.) Miq.; Randia densiflora (Wall.) Benth.

Indigenous. Trop. Asia to Pacific Is. Rare, possibly extinct. Small, glabrous tree
or shrub, 1 to 2 m high, with smooth stems; leaves, 9 to 18 cm long and 3 to 6 cm wide,
opposite, oblong-lanceolate to elliptic-ovate, acute or acuminate, base acute-decurrent or
attenuate, thinly coriaceous, rather glossy, pinnately nerved; petioles, 0.5 to 2 cm long,
ridged by decurrent leaf-blade margins; stipules, small, connate; inflorescences, axillary,
rather dense, many-flowered, rather richly-branched, short-pedunculate, cymose clusters,
up to 4 to 6 cm or longer, but not longer than the adjacent leaf, glabrous or minutely
puberulent; bracts shortly deltoid; calyx, short, 5 (rarely 4-)-toothed; corolla, usually 5-
lobed, tube and lobes about equally long, about 1.5 by 3 mm, creamy-white, throat
pubescent; stamens 4 to 5 , inserted on corolla tube; anthers, linear, exserted, nearly as
long as corolla lobes; style, rather thick, exserted, 7.5 mm long; fruit, 5 to 9 mm long,
globose, reddish to purplish, edible, with several to many seeds, crowned by calyx

186

remnant. Rare shrub with edible fruits in escarpment forest. Ripe fruit eaten, especially
by children. 2(K9), 6(165).

Gardenia augusta (L.) Merr. "gardenia"
syns. Varneria augusta L.; Gardenia florida L.; G. jasminoides Ellis; G.
radicans Thunb.

Recent introduction. China. Rare. Erect shrub, 1 to 2 m or higher; leaves, 4 to 12
cm long, and 1 to 4 cm wide, opposite, ovate, oblong-lanceolate or elliptic obovate,
acute or somewhat acuminate, decurrent at base on short petiole, somewhat glossy, dark
green; petioles, 1 to 8 mm long, winged by decurrent leaf-blade margins; stipules, often
split nearly to base with a single unilateral lobe, but sometimes bilobed; inflorescences,
axillary, solitary; calyx 5- to 7-winged, terminated by linear-lanceolate spurs, 15 to 30
mm long and 1.5 to 6 mm wide; corolla, 6 to 8 cm across, 6-petaled (double cultivars
with additional petals), tube 5 to 9 cm long, white aging yellow-brown, very sweetly
fragrant; stamens 5 to 11; fruit, 2 to 3 cm long, ovoid, 5- to 6- ribbed, crowned by
calyx, many-seeded. Planted ornamental. 6, 7.

Gardenia taitensis DC. "Tahitian gardenia", tiare Tahiti (Tahiti)
te tiare (K); tiale, siale (T)

Recent introduction. Pacific Is. Rare. Large shrub to small tree, up to 5 m tall;
leaves, 6 to 18 cm long and 3 to 10 cm wide, opposite, elliptic-obovate, obtusely short-
pointed, base cuneate, somewhat glossy or shiny, bright green, prominently pinnately
nerved, midrib very prominent; petiole, winged by decurrent leaf-blade margins, the
petiole bases of leaf-pairs joined together by stipules; petioles, usually less than 5 mm
long; stipules, connate, bilobed; inflorescences, solitary, in upper leaf axils; calyx,
prominently 4-lobed, lobes 11 to 30 cm long and 2 to 8 mm wide, lanceolate; corolla, 4
to 8 cm across, 4- to 8-petaled, petals, up to about 3 or 4 cm long, spreading from a
tubular base, about equalling tube, bright white, very fragrant; fruit, about 2 to 2.5 cm in
diameter, globose, longitudinally ridged. Planted ornamental in Tuvaluan garden at
Location. Fragrant flowers used in garlands and to scent coconut oil. 6, 7.

Gardenia sp. "Professor Pucci gardenia"

Recent introduction. Origin. Rare. Planted ornamental. 6.

187

Guettarda speciosa L. "guettarda"
iut, yut (N); te uri (K); pua, puapua, uli (T)

Indigenous. Trop. Asia to the Pacific Is. Small spreading tree, 4 to 15 m_ high,
with thick branchlets; leaves, 10 to 25 cm long and 6 to 18 cm wide, opposite, broad-
obovate, acute-subacuminate to obtuse, rounded or emarginate, base obtuse to subcordate,
rather dark green above, paler and usually puberulent beneath, with 7 to 10 pairs of
prominent lateral nerves; petioles, 2.5 to 4 cm long, pubescent, leaving large scars on the
twigs when they fall; stipules, small, ovate, leaf-like, at nodes between leaf-pair petiole
bases; inflorescences, compact, forked, many-flowered cymose clusters on axillary
peduncles, 5 to 10 cm long, usually forming after leaf-fall; calyx, about 1 cm long,
cupulate, truncate, pubescent; corolla, tubular, 2.5 to 5 cm long and 2 to 3 cm across the
limb, 5- to 6-petaled, dull or yellowish white, fragrant, villous within; flowers dimor-
phic, some with short styles not reaching the anthers, others with exserted styles; fruit, 2
to 3 cm in diameter, a woody, depressed-globose, 4- to 9-chambered drupe with a
circular rim at apex, whitish, pinkish or green when ripe, faintly ribbed, usually 4- to 6-
seeded. Occasional in woods and thickets on coastal strip, but less common in forest on
plateau near escarpment; common in regrowth in older strip-mined areas; planted in
home gardens. Straight pieces of timber make excellent house rafters and used for canoe
parts and handicrafts; bark scraped and mixed with other plant extracts, eg., Scaevola
taccada (emet) and Cordia subcordata (eongo) to produce a medicine for beriberi; leaves
used to parcel food and as plates; flowers used in garlands and decorations, often after
soaking in coconut oil; flowers considered the best for scenting coconut oil; juice from
flowers used as a deodorant; flowers and young leaves soaked in water to make a love
potion, which is drunk by women and which when sweating, makes men go crazy
(reportedly learned from I-Kiribati); juice of flowers mixed with coconut juice used to
keep hair healthy. 2, 3(58759), 5(70), 6, 7(27806).

Hedyotis corymbosa (L.) Lam.
syn. Oldenlandia corymbosa L.

Recent introduction. Pantropical. Rare. Much-branched, annual herb, up to 20 cm
high, with slender ascending or erect, bluntly 4-angled stems; leaves, 1 to 4 cm long and
1.5 to 8 mm wide, opposite, linear-oblong or narrowly elliptic, acute at both ends, blade
pale beneath, midrib prominent, chartaceous, subsessile or petiole very short; stipules
fused to petioles; inflorescences, 2- to 8-flowered axillary cymes which are shorter than
the leaves; pedicels, 4 to 8 mm long, slender; calyx, about 2 mm long, 4-lobed, not
exceeding ovary; corolla, about 2 mm long, white or faintly pinkish-purplish; stamens
inserted just above the base of the tube; fruit, about 2 mm by 2 mm, a 2-celled capsule,
exalate, flattened at apex, slightly laterally compressed, with several seeds per cell,
dehiscent by valves at apex. Weed in waste places and ruderal habitats. 4(140N), 5(143),
6, 8(9560).

188

Ixora casei Hance "ixora"
te katuru, te katiru (K); suni (T)
syns. J. duffii Baine; J. duffii T. Moore; I. carolinensis Hosok.; I. confertiflora
Val. non Merr.; /. volkensii Hosok.; I. pulcherrima Volk.

Recent introduction. Caroline Is. (Southeast Asia?). Occasional. Large shrub, up
to 3 m or higher; leaves, up to 30 cm long, opposite, narrow-oblong, acute, dark-green
glossy, glabrous, short-petiolate; stipules, interpetiolar, entire, acuminate; inflorescences,
many-flowered, terminal cymose clusters, about 20 cm in diameter, not at all fragrant;
calyx, tiny, 4-toothed; corolla, about 5 cm long, tubular, 4-lobed, lobes, ovate, more or
less acute, spreading, twisted spirally in bud, deep vermilion-red to scarlet; stamens 4;
fruit (rare), a 2-seeded, berry-like drupe. Planted ornamental in Nauru, but reportedly
wild in Palau, Chuuk (Truk) and Pohnpei (Ponape) and Kosrae (Kusaie). 3(58707), 5, 6,
te

Ixora coccinea L. "ixora", "flame of the woods"
syn. I. fraseri Hort. ex Grant

Recent introduction. Southeast Asia. Occasional. Small shrub, up to 2.5 m high;
leaves, 3 to 10 cm long and 2 to 5 cm wide, opposite, oblong-obovate or subcordate and
slightly amplexicaul, obtuse, coriaceous, sessile; inflorescences, dense terminal corym-
bose clusters; calyx, 4-toothed, puberulent; corolla, up to 5 cm long, tubular, 4-lobed,
lobes 10 to 15 mm long, ovate-lanceolate, acute, red, orange-pink or yellow; stigmas, 3
to 4 mm long, red; fruit, a 2-seeded berry-like drupe. Planted ornamental. 5, 6(202).

Ixora sp. "ixora"

Recent introduction. Rare. Planted ornamental. 5, 6(196).

Morinda citrifolia L. var. citrifolia "beach mulberry", "Indian mulberry"
deneno (N); te non (K); nonu (T)
syn. M. indica L.

Indigenous. Trop. Asia and Australia to S.E. Polynesia. Common. Shrub or small
tree, up to 6 m or higher, with quadrangular branchlets; leaves, 15 to 30 cm long and 6
to 18 cm wide, opposite, broadly elliptic or slightly obovate, acute or obtuse, dark green,
glossy, pubescent when young; petioles, 1 to 2 cm long, narrowly ridged by decurrent
leaf-blade margins; stipules, interpetiolar, sheathing, large, connate, deltoid, sometimes
bifid; inflorescences, axillary, solitary, fleshy, headlike clusters, up to 2.5 cm long,
irregularly subglobose on peduncles, 2 to 3 cm long; calyx, truncate; corolla, about 1 cm
long, pilose in the throat, usually with 5 acute lobes about 5 mm long, pure white;
anthers included or slightly exserted; style about 1.5 cm long; fruit, 4 to 12 cm long, a

189

globose-ovoid, fleshy, compound drupe (syncarp) containing 1-seeded pyrenes, somewhat
waxy, shiny, creamy to yellowish-white; flesh, strongly fetid and gelatinous, but edible
when ripe; seed, bony. Common to abundant on coastal strip, often protected or planted
around homes and in waste areas; occasional in plateau forest and in older strip-mined
areas. Plants kept around homes to ward off evil spirits; small pieces cut up and rubbed
on hands, face, etc., to ward off evil spirits; roots ground to provide a yellow dye; ripe
fruit eaten raw, but far more common eaten in the past; fruit cooked and mixed with
coconut syrup to make pudding (dedangan); fruit and branches cooked to eliminate evil
smells, especially after war; roots and branches crushed and squeezed to cure rashes,
fruit and roots ground and cooked to cure headaches, tender leaves heated up with
coconut oil and used as a poultice to suck puss out of boils (ibir, ibur), raw fruit ground
and drank as a cure for diabetes, and fruit cooked and used as a cure for dysentery. Plant
also a very important medicinal and multi-use plant in Kiribati and Tuvalu. In Kiribati,
where there have been recent outbreaks of vitamin A-deficiency-induced nightblindness
among young children, programs encouraging mothers to feed children the cooked leaves
of vitamin A-rich M. citrifolia have reportedly improved the situation. 2, 3(58754),
5(98), 6, 7(27807).

Mussaenda erythrophylla Schum. & Thonn. "red mussaenda"

Recent introduction. Trop. Africa. Rare. Shrub or liana, 2 to 4 m high, often with
drooping or climbing branches and copiously pilose branchlets; leaves, 5 to 12 cm long
and 3 to 8 cm wide, oval-ovate to almost cordate (when young), acuminate, base obtuse
to slightly decurrent, copiously pilose, prominently veined; petioles, 1 to 6 cm long;
stipules, interpetiolar; inflorescences, terminal, several-flowered, branched, corymbose
cymes; calyx, 5-lobed, one of the lobes (rarely 2) enlarged, petal-like, up to 9 cm long
and 6.5 cm wide, ovate-lanceolate, lax-pendant, bright red above and somewhat paler
beneath, palmately veined, pubescent, the other lobes, up to about 1 cm long and 2 mm
wide, narrowly linear-lanceolate, caducous, bright red, calyx tube reddish-pubescent;
corolla, about 2.5 cm long, tubular, 5-lobed, red-pubescent outside with tube white or
pale-yellow within distally; stamens 5, included; fruit, a berry with numerous minute
seeds; Planted ornamental. 6(195).

Mussaenda frondosa L. "mussaenda"
syn. M. sericea Bl.

Recent introduction. Trop. Africa and Madagascar to S. Asia. Rare. Erect shrub,
up to 4 m or higher, with drooping or rambling branches; leaves, 5 to 18 cm long and 3
to 7 cm wide, opposite, ovate to lanceolate, acuminate, base obtuse, rounded or decur-
rent, puberulent; petioles, 1 to 3 cm long; stipules, concealing a band of hairs, caducous;
inflorescences, terminal corymbose cymes; bracts and bracteoles caducous; calyx 5-lobed,
one of the lobes (rarely more) enlarged, petal-like, foliaceous, colored, white or pink;
corolla, mostly 2 to 3 cm long, tubular, 5-lobed, dark-yellow to orange; stamens 5,

190

included, anthers normal but always devoid of pollen in "female", long-styled flowers;
fruit, 1 to 1.5 cm long, a berry, ellipsoid, truncate at base, black when ripe, seeds,
numerous, reticulate. Planted ornamental. 6(248).

Pentas bussei K. Kr. "red pentas", "red star cluster"

Recent introduction. Trop. Africa? Rare. Planted ornamental. 3(58675), 6.

Pentas lanceolata (Forssk.) K. Schum. "pentas", "Lady Fletcher", "Egyptian star cluster"
syns. Ophiorrhiza lanceolata Forsk.; Pentas carnea Benth.

Recent introduction. Trop. Africa, Madagascar, Comorro Islands and Arabian
Peninsula. Occasional. Erect, pubescent, sometimes straggling, perennial herb or sub-
shrub, up to 80 cm high; leaves, 2.5 to 15 cm long, opposite, ovate, acute to acuminate,
pubescent; inflorescences, up to 7.5 cm across, terminal, open, headlike, corymbose
clusters; corolla, long-tubular, commonly 5-lobed, lobes, acute, mauve or rich purple to
pink and white, with a white-bearded throat; fruit, about 4 mm long, obovoid, capsular
with minute seeds. Planted ornamental. 3(58689), 5, 6(233,166), 7.

Spermacoce assurgens R. & P. "buttonweed"
syns. S. suffrutescens Jacq.; Borreria laevis sensu auct. plur. non (Lam.) Griseb.

Recent introduction. S. Asia. Uncommon. Decumbent or ascending, perennial
herb, up to 40 cm high, with slender, wiry, somewhat angled stems and slightly woody
and branching near the base; leaves, 2 to 6 cm long and about 0.5 to 2 cm wide, elliptic-
lanceolate, sharply acuminate, base cuneate, pinnately nerved, nerves prominent, scabrid
near margin but otherwise glabrous, subsessile; stipules, connate, with filamentous
processes, interpetiolar, fused to the leaf-stalks; inflorescences, in axillary verticillate
headlike clusters; calyx, small, 4-lobed, lobes ovate; corolla, 1.5 to 3.5 mm _ long,
infundibular, 4-lobed, lobes, spreading, white or pink-tinged; stamens 4, exserted from
the corolla, with white filaments and pale blue or bluish-tinged anthers; style, pink and
white-tinged; fruit, 1 to 4 mm long, 2-seeded, enclosed in calyx, finally splitting; seeds,
1.5 to 2 mm long, oblong, brown, transversely ridged, still clasped by the half-calyx
with part of the septum. Weed in waste places and houseyard gardens. 5(2), 6.

Tarenna sambucina (Forst.f.) Dur. ex Drake
syns. Coffea sambucina Forst.f.; Stylocoryna sambucina (Forst.f.) A. Gray;
Tarenna glabra Merr.

Indigenous? New Caledonia to S.E. Polynesia and Micronesia. Extinct? Small
glabrous or minutely puberulent shrub or small tree, 1.5 to 12 m high, with 4-angled

191

young branches; leaves, 10 to 20 cm long and 5 to 9 cm wide, opposite, narrowly-elliptic
or ovate-elliptic, acute at both ends, attenuate to petiole, membranaceous or thin-
coriaceous, lateral nerves usually 7 or 8 pairs; petioles, 1 to 4 cm long; stipules,
interpetiolar, deltoid, entire, persistent; inflorescences, terminal, many-flowered corym-
bose cymes; calyx, about 2 cm long, 5-lobed; corolla, 6 mm long, funnelform, creamy-
white turning yellowish, villous within, fragrant; stamens 5; style exserted 3.5 mm long,
anthers white, the style greenish to white; fruit, 5 to 6 mm in diameter, a globose berry,
green turning purplish-black; seeds, about 8, each 2 mm long, 4-sided. Reported present
by Burges in 1935, but not collected since. 2.

RUTACEAE (Rue Family)

Citrus aurantifolia (Christm.) Swingle "lime"
derem, deraim (N); te raim (K); laim, tipolo (T)
syn. Limonia aurantifolia Christm.

Pre-World War II introduction. Malesia. Occasional. Small, much-branched tree,
2 to 6 m high, with copious short, stiff, sharp spines; leaves, 4 to 10 cm long and 4 to 6
cm wide, alternate, usually ovate-elliptic but variable in different cultivars, margins
somewhat crenulate; petioles, 5 to 18 mm long and 1.5 to 6 mm wide, spathulate,
narrowly-winged, leaflike, articulate with leaf blades; inflorescences, axillary, 1- to 7-
flowered, produced over an extended period, 2 to 2.5 cm in diameter; corolla, small, 4-
to 5-parted, 8 to 12 mm long and 2.5 to 4 mm wide, white to pale pink; stamens 20 to
25; ovary 9- to 12-locular; fruit, 2.5 to 6 cm in diameter, ovoid to subglobose, green to
yellowish when ripe depending on the cultivar; peel, thin, 1 to 3 mm thick, glandular,
adherent; pulp greenish, juicy, very acid but pleasantly fragrant in some cultivars; seeds,
small, oval, flattened angular, pale yellow or whitish. Planted fruit tree. Juice of fruit
used to marinate raw fish and to make drinks. 3(58695), 5, 6(221), 7.

Citrus limon (L.) Burm.f. "lemon"
te remen, te remon (K); moli, laim (T
syns. C. medica var. limon L.; C. limonum Risso; C. limonia Osbeck

Pre-World War II introduction. E. Asia. Rare. Small tree or shrub, 2 to 9 m high,
with stout, stiff spines; leaves, 5 to 10 cm long and 3 to 6 cm wide, alternate, long-
ovate, acute, serrate; petioles, 0.5 to 1 cm long, very narrowly winged or margined,
cordate, clearly articulate with leaf blades; inflorescences, axillary, solitary or few, 3.8
to 5 cm in diameter; corolla, usually 5-parted, petals white within but pinkish or
purplish-tinged outside and when in bud; stamens 20 to 40; ovary 8- to 10-locular; fruits,
5 to 12 cm long, ovoid or ellipsoid, 8- to 10-locular, with a broad apical papilla or
nipple; peel, about 5 mm thick, adherent, prominently gland-dotted, slightly rough, green

192

to bright yellow when ripe depending on the cultivar; pulp, pale yellow, juicy, sour;
seeds, ovoid, white. Planted fruit tree. Reportedly more abundant in the past. Juice used
to marinate raw fish and to make drinks. 5, 6, 7.

Citrus reticulata Blanco "tangerine", "mandarin orange"
syns. C. deliciosa Ten.; C. nobilis Lour. var.

Recent introduction. Rare. E. Asia. Small, usually spiny tree, 3 to 8 m high;
leaves, 4 to 8 cm long and 1.5 to 4.5 cm long, alternate, ovate or elliptic to lanceolate,
usually crenate, dark shining green above, yellowish-green beneath; petioles, 5 to 15 mm
long and 1.5 to 3 mm wide, alternate, narrowly winged or margined, 1.5 to 3 mm wide
across the wings, clearly articulate with leaf blades; inflorescences, axillary, small 1.5 to
2.5 cm in diameter; corolla, usually 5-parted, petals, white; stamens about 20; ovary 10-
to 15-locular; fruit, 5 to 8 cm in diameter, depressed-globose to subglobose, greenish
with yellowish patches to bright orange; peel, thin, loose, separating easily from the
segments; pulp, orange, sweet, juicy; seeds, small. Immature planted fruit tree. 5, 6.

Citrus sinensis (L.) Osbeck “orange”, "sweet orange"
te aoranti (K)
syn. C. aurantium var. sinensis L.

Recent introduction. S. Asia. Rare. Small to medium tree, 4 to 12 m high, often
with stout spines; leaves, 5 to 15 cm long and 2 to 8 cm wide, alternate, ovate to oval-
elliptic, glabrous, dark green above, sometimes slightly serrate; petioles, 1 to 2.5 cm
long and 3 to 5 mm wide, winged, clearly articulate with leaf blades; inflorescences,
axillary, solitary or in small racemes, 2 to 3 cm in diameter; corolla, usually 5-parted,
petals white, fragrant; stamens 20 to 25, united into groups; ovary 10-to 14- locular;
style slender with globose stigma, soon deciduous; fruit, 4 to 12 cm in diameter,
subglobose, greenish-yellow or orange, juicy, edible; peel up to 5 mm thick, tightly
adherent to segments; pulp, yellow-orange to dark orange, juicy, slightly acidic, sweet;
seeds, none to many, obovoid, white. Immature planted fruit tree; reportedly more
common in past and planted by the Japanese during World War II. 6, 7.

Murraya paniculata (L.) Jack "mock orange"," orange jessamine", "orange jasmine"
syns. Chalcas paniculata L.; Murraya exotica L.

Recent introduction. Trop. Asia, Malesia and Australia. Rare. A shrub to small
tree, 1 to 8 m high, but usually kept under 3 m by pruning; leaves, alternate, pinnate;
leaflets, 3 to 7, usually 5, 3 to 10 cm long and 2 to 4 cm wide, ovate to subobovate,
glossy, thin-coriaceous, nitid; petiolules, very short; inflorescences, 1- to 8-flowered,
compact paniculate clusters; calyx, S-parted, deeply-lobed, sepals basally somewhat
united; corolla, 5-parted, petals, usually 10 to 15 mm long and 3 to 6 mm wide,

193

imbricate, pure white, fragrant; stamens 10, the filaments 6 to 12 mm long, the style 5 to
9 mm long; ovary 2-locular, styles with capitate stigma; fruit, 1 to 1.3 cm long, an ovoid
to ellipsoid reddish-orange berry; seeds, 1 to 2, villous. Planted ornamental and pot plant
at Location. 6(184), 7.

SAPINDACEAE (Soapberry Family)

Dodonaea viscosa (L.) Jacq. “native hop bush"
eteweo, eteweau (N)
syns. Ptelea viscosa L.

Indigenous. Pantropical. Common. Shrub or small tree, 0.5 to 6 m high, with
angular branchlets and slightly sticky resinous exudate; leaves, 5 to 12 cm long and 1 to
3 cm wide, alternate, narrowly lanceolate or oblanceolate, acute at both ends, long-
attenuate at base, margins somewhat wavy, firm to somewhat coriaceous, olive green
above, paler beneath, somewhat viscid appearing varnished, subsessile; petioles, 1 to 2
mm long, winged; inflorescences, in short, lax terminal or axillary panicles about half as
long as the leaves, individual flowers about 4 mm across, greenish-yellow; calyx, 5-
parted; petals none; disk, small or obsolete; stamens, usually 8, distinct; fruit, 15 to 20
mm in diameter, inflated, papery, reddish or orange to brown, 2- to 4-winged (lobed)
capsule, the wings membranous, veiny, glabrous; seeds, 1 or 2 per lobe, black, with a
spiral embryo. Common in scrub and occasional in plateau forest; occasional in revege-
tated older mined areas. Strong but flexible trunks and branches used for fishing rods and
for frames for flying-fish and noddy nets; leaves occasionally used to scent coconut oil.
23(08598,'98637);, (73), 63:7; 80578).

Nephelium litchi Camb. "lychee", "lychee nut"
syn. Litchi chinensis Sonn.

Recent introduction. S. China. Rare. Small to medium tree, up to 10 m high, with
a heavy trunk, spreading crown and thick foliage; leaves, paripinnate with 2 to 4 pairs of
leaflets; leaflets, 8 to 15 cm long and 3 to 4 cm wide, elliptic-oblong, coriaceous, glossy;
inflorescences, large terminal panicles; individual flowers, small, pale greenish-yellow;
calyx, 5-parted; petals none; disk, annular; stamens 6 to 10; fruit, 3 to 3.5 cm in
diameter, pendent in loose clusters, covered with small pyramidal tubercles, usually
bright red to purplish when ripe; pulp-like aril, white, translucent, fleshy, somewhat
gelatinous, juicy, sweet, fragrant, edible, with a grapelike consistency; seed, dark brown,
covered by aril. Planted fruit tree seedling. 6.

194
SAPOTACEAE (Sapodilla Family)

Chrysophyllum cainito L. "star apple", "cainito"

Recent introduction. W. Indies. Rare. Medium tree, 5 to 12 m high, with
puberulent, golden-brown young branchlets and white latex; leaves, 8 to 20 cm long and
3 to 9 cm wide, alternate, 2-ranked, elliptic to elliptic-oblong, sharply acute or obtuse,
base acute, coriaceous, pinnately nerved, lateral veins very slender, shiny, dark-green
above, densely silky-pubescent and golden-brown beneath; petiole, 1 to 2 cm long;
inflorescences, axillary, often many-flowered, pedicellate to sessile clusters; calyx, 5 (4
to 6)-parted, spirally arranged; corolla, usually exserted from calyx, 3 to 5 mm long,
tubular, 5 (4 to 11)-lobed, lobes equal to or longer than tube, purplish-white; stamens, as
many as corolla lobes, inserted; fruit, 5 to 12 cm in diameter, subglobose to ovoid,
smooth, with light green to purple skin; pulp, white, sweet, often juicy, edible; seeds, 4
to 8, hard, obliquely obovoid, compressed, with a broad lateral scar, glossy, brown,
embedded in a star-like arrangement in the pulp. Planted fruit tree seedling. 6.

SAXIFRAGACEAE (Saxifrage Family)
Saxifraga sarmentosum L.f mother of millions", "mother of thousands", "roving sailor",
"strawberry geranium"
syn. S. stolonifera ?? authority

Recent introduction. E. Asia. Rare. Tufted perennial herb, up to 30 cm high, with
many slender, strawberry-like, branching red runners upon which numerous new plants
are formed; leaves, 2.5 to 10 cm in diameter, rounded, cordate, scalloped or dentate,
softly-pubescent, green and white- or silvery-veined or variegated above and pale green,
suffused with red or pink beneath; inflorescences, erect, loose, long-stalked, spikelike
clusters, up to 15 to 50 cm high; corolla, 5-petaled, star-like, 2 petals about 12 mm long
which far exceed the other 3, irregularly-shaped, white to flesh-pink; stamens 10. Pot
plant. 4 (144).

SCROPHULARIACEAE (Snapdragon Family)
Angelonia angustifolia Benth. “angelonia", "monkey face"
syns. A. gardneri auct. non Hook.; A. salicarifolia auct. non H. & B.

Recent introduction. Trop. America. Occasional. Erect perennial herb, up to about
60 cm high; leaves, up to 7.5 or 9 cm long and 1 cm wide, opposite or verticillate (or

195

distal one alternate), narrow-oblong or lanceolate, minutely glandular-serrulate to
subentire, sessile or subsessile; inflorescences, axillary, solitary or paired, slender-
pedicellate; calyx, deeply 5-lobed; corolla, 1.5 to 2 cm across, short-tubular, 2-lipped,
the limb widely cupuliform, about 2 cm across, short-spurred, 5-lobed, 4 broad and flat,
the lower one cupuliform in the lower half, the throat broadened into a sac-like cup, blue
or purplish; stamens 4, borne on the dorsal side of the cupuliform part of the corolla, the
filaments short, thick; fruit, a globose or ellipsoid, 2-valved capsule, rarely indehiscent.
Planted ornamental. 3(58723), 5, 7.

Angelonia biflora Benth. “angelonia"

Recent introduction. Trop. America. Rare. Erect perennial herb, up to 1 m high;
very similar to A. angustifolia, except that it is slightly taller and has a slightly larger and
paler, often white, corolla. According to Smith (1991), it is questionable whether these
two taxa constitute separate species. 6(256). Planted ornamental

Bacopa procumbens (Mill.) Greenm.
syns. Erinus procumbens Mill.); Herpestris chamaedryoides HBK.; Bacopa
chamaedryoides (HBK.) Wettst.

Recent introduction. Trop. America. Suberect, perennial herb, with quadrangular
glabrous stems, rooting at the nodes; leaves, | to 2.5 cm long and 6 to 12 mm wide,
opposite, elliptic or shortly ovate-elliptic, crenulate-serrulate; inflorescences, axillary,
usually solitary; pedicels, 4 to 6 mm long, extending in fruit to 18 mm; calyx 5-parted,
upper segment broadest; corolla, slightly longer than the calyx, subregular, cylindric-
tubular, 5-lobed, yellow; stamens, 5; fruit, a 2-valved capsule, enclosed by the calyx.
Weed in lawns and disturbed places. 2 (58813).

Russelia equisetiformis Schlecht. & Cham. "firecracker flower", "coral plant"
dokaibangi, dugaibangi, dogaibwangi (N); te kaibaun (K)
syn. R. juncea Zucc.

Recent introduction. Mexico. Occasional. Shrub or subshrub, up to 1.5 m high,
with long, straggling, longitudinally-ridged, whorled or drooping green branches or
stems; leaves, up to 2 cm long and 1.5 cm wide, ovate, dentate, whorled in groups of 5
to 8 on main stem, but more often, reduced to minute, opposite, linear, awl-shaped scales
on branches; inflorescences, narrow, terminal, loose, drooping paniculate cymes; pedicels
about 1 cm long; calyx 5-parted; corolla, 1.5 to 2.5 cm long, long-tubular, with a small
5-lobed limb, lobes about 3 mm long, bright red, nodding; stamens 4, inserted; fruit,
about 5 mm long, globose or broadly ellipsoid, 4-valved, rostrate; seeds, many, oblong.
Planted ornamental. 3(58678), 4(157N), 5, 6(213), 7.

196

Russelia sarmentosa Jacq.

Recent introduction. Mexico. Rare. Shrub or subshrub, up to 1.5 m high; leaves,
ovate to triangular, in whorls; inflorescences, in clusters of 30 to 40 individual flowers;
calyx S-parted nearly to base; corolla, tubular, 5-lobed, scarlet; stamens 4, inserted;
fruit, globose or broadly ellipsoid, 4-valved, rostrate; seeds, many, oblong. Planted
ornamental. 3 (58795).

Scoparia dulcis L.

Recent introduction. Trop. America. Rare. Branched, subligneous, perennial herb,
20 to 80 cm high, with wiry, angular-ribbed stems; leaves, 3 to 4 cm long and 1 to 1.5
cm wide, opposite or whorled in threes, narrowly elliptic, oblong-obovate or oblan-
ceolate, subsessile, distally serrulate or serrate, gland-dotted beneath; inflorescences,
axillary, solitary, pedicellate; calyx 4-lobed nearly to base, lobes, imbricate; corolla, 5 to
8 mm across, 4-lobed nearly to base, the lobes spreading, subequal, obtuse, white to pale
blue or pale purple with a darker center, pubescent within; stamens 4, subequal,
greenish; stigma, slender; ovary green; fruit, an ovoid or subglobose, 2-valved capsule;
seeds, many, tuberculate or scrobiculate. Weed of waste places. 6(159), 8(9557).

SOLANACEAE (Nightshade Family)

Capsicum annuum L. var. annuum "chilli pepper", "red pepper"
epeba (N); te beneka (K); tili, polo feuu (T)

Recent introduction. Trop. America. Erect, glabrous, annual or short-lived, much-
branched perennial herb or subshrub, up about 1 m high; leaves, 1.5 to 12 cm long and
0.5 to 7.5 cm wide, alternate, broadly lanceolate to ovate-elliptic or acute-acuminate,
base cuneate or acute, thin, subglabrous; petioles, 0.5 to 2.5 cm long; inflorescences,
usually solitary or paired, terminal, pendulous, but because of the form of branching
appear somewhat axillary; pedicels, up to 1 or 1.5 cm long, usually 1 per node after first
flowering; calyx, about 2 mm long, campanulate, shortly 5-dentate, 10-ribbed, persistent,
usually enlarging to enclose the base of the fruit; corolla, 0.8 to 2 cm in diameter, rotate-
campanulate with a very short tube, deeply S- to 6-parted, white, sometimes with a
greenish or bluish tinge; stamens, 5 to 6, inserted near base of corolla, anthers bluish or
purplish drying green, dehiscing longitudinally, style simple, white or purple; fruit, 3 to
6 cm long and 2 to 3 cm wide, oblong to subglobose, somewhat irregularly shaped,
wrinkled or twisted, a many-seeded pendent berry, with a leathery skin, green turning
bright red when ripe, indehiscent, hot-spicy, pungent; seeds, 2 to 4 mm long, subor-
bicular, compressed, pale yellow. Planted spice plant in expatriate garden at Meneng
Terrace. Fruit used to spice food. 6, 7.

197

Capsicum annuum L. var. grossum (L.) Sendtn. "bell pepper", "sweet pepper",
"sweet capsicum", "paprika", "pimento" pepa (T)
syn. C. grossum L.; C. dulce Hort. ex Dun.

Recent introduction. Trop. America. Rare. Similar to C. annuum L. var. annuum,
but bearing a larger, oblong to linear, almost quadrangular, or sometimes broadly
ellipsoid-acuminate fruit, 6 to 14 cm long and 5 to 10 cm across, green to bright red or
occasionally yellow, partially hollow, usually sweet and non-pungent. Food plant in
Topside workshop gardens. Fruit eaten raw in salads and cooked as a green vegetable. 5,
6.

Capsicum frutescens L. "tabasco", "bird chilli", "perennial chilli"
epeba (N); te beneka (K); tili, polo feuu (T)
C. minimum Roxb.

Pre-World War II introduction. Trop. America. Rare. Coarse, perennial,
suffrutescent, much-branched subshrub, 0.6 to 2 m high, leaves, 2 to 9 cm long and 1 to
4 cm wide, alternate, ovate-elliptical to ovate-lanceolate, acute-acuminate, base acute or
obtuse, entire or slightly undulate, glabrous or slightly pubescent; petioles 1 to 3 cm
long; inflorescences, terminal and axillary, commonly paired, sometimes more, long-
pedicellate; pedicels, 1 to 2.5 cm long, 2 (rarely 3) per node after first flowering; calyx,
2.5 to 3 mm long, tubular, shortly 5-dentate, persistent, usually enlarging to enclose the
base of the fruit; corolla, 1 to 1.5 cm in diameter, deeply 5-lobed, white to greenish-
white or purplish-white; stamens 5, anthers purple; fruit, 1 to 1.5 cm long and about 5
mm across, ellipsoid-lanceolate, ellipsoid-ovoid or subconic, an erect, many-seeded
berry, with a leathery skin, green or greenish-yellow turning bright red when ripe,
indehiscent, hot-spicy, pungent; seeds, 2 to 3 mm long, suborbicular, compressed, pale
yellow. Planted or protected in Tuvaluan garden at Location and in Topside Workshop
gardens. Reportedly much more common in the past. Fruit used to spice foods. 6.

Cestrum nocturnum L. "night cestrum", "night flowering cestrum", "night-flowering
or night-blooming jasmine", "lady of the night" fafine o te po (T)

Recent introduction. W. Indies. Rare. Erect, glabrous shrub, 2 to 5 m high;
leaves, 7 to 20 cm long, spiralled, ovate-oblong to lanceolate-elliptic, petiolate; inflores-
cences, cymose racemes longer than petioles; calyx, about one-third as long as the
corolla, 5-toothed, green, persistent; corolla, 1.5 to 2.5 cm long, tubular, 5-lobed,
narrowed at throat, widened at apex, with a small spreading limb, lobes, 5 to 6 mm long,
obtuse to subacute, erect or spreading, greenish-yellow, pale-yellow or cream-white,
extremely fragrant, mostly at night; stamens 5, included, puberulent at base; fruit, 6 to
10 mm long, a subglobose berry, white, 1- to 3-seeded. Planted ornamental. 5(15), 6.

198

Datura metel L. "datura", "cornucopia", "jimson weed"
syn. D. fastuosa L.

Recent introduction. S.E. Asia. Rare. Erect herb or subshrub, | to 2 m high;
leaves, 4 to 25 cm long and 2 to 20 cm wide, narrowly ovate to angular-ovate, acute,
base oblique, sinuate-repand to sinuately-lobed, major lobes 2 or 3 per side, glabrous to
slightly puberulent; petiole, 1 to 16 cm long; inflorescences, solitary, terminal, single or
double (or triple); pedicels, 8 to 15 mm long, erect or nodding; calyx, 3 to 9 cm long,
tubular, with 5 (-9) conspicuous longitudinal veins, usually 5-lobed, lobes 8 to 25 mm
long, circumscissile after the anthesis; corolla, commonly 12 to 18 cm long, often double
or triple, tubular, trumpet-like, 5-lobed, the lobes with acumens 1 to 2.5 cm long, purple
without, pale lavender to white within; stamens 5, at apex of tube, anthers 1 to 2 cm
long; style, 10 to 14 mm long; fruit, 3 to 4 cm in diameter, a capsule, deflexed,
subglobose, prickly or spiny with 100 to 200 conical tubercules, each 2 to 5 mm long,
irregularly dehiscent; seeds, 4 to 5 mm long, many, compressed, yellow-brown. Planted
ornamental. 6.

Nicotiana tabacum L. "tobacco"
te kaibake (K)

Pre-World War I introduction. Trop. America. Rare. Coarse, erect herb up to 2
m or higher; leaves, 30 to 60 cm long, spiralled, the lowest ones forming a rosette,
oblong-ovate to lanceolate, acute, sometimes shortly-decurrent on stem, surfaces
pubescent, subsessile; inflorescences in terminal racemose panicles, pubescent; calyx, 1
to 2 cm long, tubular-campanulate, S-lobed, accrescent, puberulent; corolla, 3 to 5 cm
long, tubular, 5-lobed, lobes, ovate, puberulent, pink to red; stamens 5, included or
slightly exserted, filaments puberulent at base; ovary glabrous; fruit, 15 to 17 mm long, a
brown capsule with persistent calyx at base; seeds, numerous, minute, subreniform,
tuberculate. Planted in home gardens; reportedly more common in the past. Leaves dried
and cured for smoking. 3, 5, 6.

Physalis angulata L. “cape gooseberry", "bladderberry", "ground cherry"
watamo, oatamo (N); te baraki (K); pini (T)

Pre-World War II introduction. Trop. America. Common. Erect, nearly glabrous,
branching annual herb, up to 80 cm or higher, with hollow angular-ribbed stems; leaves,
3 to 14 cm long and 2.5 to 9 cm wide, alternate, ovate to oblong-lanceolate, acute-
acuminate to obtuse, base obtuse or acute (somewhat asymmetrical), irregularly toothed
or sinuate-entire, very short-puberulent on both sides; petioles, 2 to 11 cm long;
inflorescences, axillary, pedicellate; pedicels, 0.7 to 1.5 cm long, slender, enlarging to
2.5 cm in fruit; calyx, 3 to 6 mm long, enlarging to 2 to 4 cm long in fruit, short-
puberulent; corolla, 0.6 to 1 cm long, pale-yellow, yellowish-green near base, partly
puberulent externally; stamens 5, 2.5 to 4 mm long, capitate; anthers, 1 to 2.5 mm long,
blue; style, 4 to 5 mm long; fruit, 1 to 1.6 cm in diameter, subglobose or ellipsoid,

199

fleshy, enclosed in an inflated, balloon-like, ovoid calyx, up to about 3 cm long, with an
apical opening; seeds, about 1.7 mm long, flat, reniform, yellowish. Weed in low ground
near Buada Lagoon, on road fill in currently mined areas, and in disturbed soil and
dumps on coastal strip. Ripe fruit eaten by children. 2, 3(58645, 58768), 4(130N), 5(14),
6, 7(22321), 8(9541).

Physalis lagascae R. & S. "ground cherry", “wild cape gooseberry", "bubble fruit"
watamo, oatamo (N); te baraki(K); pini (T)
syn. P. minima L.

Recent introduction. Trop. America. Rare. Low, much-branched, somewhat
sprawling annual herb; leaves, 1.5 to 6 cm long and | to 3 cm wide, mostly alternate,
oval-ovate, acute at both ends, margin entire or somewhat wavy, spreading pilose on both
surfaces; petioles, 0.5 to 3 cm long, slender; inflorescences, solitary, axillary; pedicels, 2
to 5 mm long, enlarging to 9 mm in fruit; calyx, short-spreading-pilose without,
glabrescent, green, inflated, enclosing but free from fruit; corolla, 4 to 6 mm across,
yellow, with a darker center with 5 very distinct brownish blotches; fruits, 5 to 7 mm in
diameter, berrylike, sessile, enclosed in a papery, inflated, bag-like calyx, 12 to 25 mm
long, ovoid acuminate, prominently 5-angled. Weed in low ground near Buada Lagoon.
3(58646).

Solanum lycopersicum L. var. lycopersicum "tomato"
te tomato (K); tomato (T)
syns. Lycopersicon esculentum Mill.; L. lycopersicum (L.) Karst.

Pre-World War II introduction. Trop. America. Rare. Erect, pilose, glandular
herb, 0.5 to 2 m high, with capitate hairs on the younger parts, slightly succulent, solid
green stems, which sometimes becoming prostrate and spreading, a strong tap-root, and a
distinctive strong odor; leaves, 10 to 30 cm long and 6 to 20 cm wide, spirally arranged,
pinnate or bipinnate, with a terminal leaflet and equal or unequal smaller lateral leaflets
alternating with larger leaflets; petiole, 3 to 6 cm long; larger leaflets (major pinnae) 7 to
9, 5 to 10 cm long, opposite and/or alternate, usually irregularly toothed or incised,
sometimes pinnatifid, puberulent; inflorescences, extra axillary (opposite or between the
leaf axils), 4- to 12-flowered racemes, the individual flowers, about 2 cm in diameter,
pendent; pedicels, 1 to 2 cm long; calyx, 5- to 8-lobed, sepals, about 1 cm long, slender,
persistent, enlarging in fruit; corolla, about 1 cm long, rotate, 5- to 8-lobed, lobes,
lanceolate, stellate, later reflexed, yellow; stamens, 5 to 8 or 10; anthers, 5 mm long,
apical appendage 2 mm; ovary glabrous; fruit, 2 to 12 cm in diameter, subglobose,
usually depressed at both ends, a 3 to 12-celled berry, bright red or yellowish when ripe,
juicy, edible; seeds, many, 3 to 5 mm long and 2 to 4 mm wide, reniform, compressed,
silvery-pubescent, light brown or yellowish. Food plant at Location and in expatriate
home gardens; spontaneous in waste heaps. 5, .

200

Solanum melongena L. “egg plant", "aubergine", "brinjal"

Recent introduction. S. Asia. Rare. Erect, stellate-tomentose, weakly-perennial,
branching herb or subshrub, 0.5 to 1.5 m high, the stems sometimes prickly and all parts
covered with a gray tomentum; leaves, 6 to 25 cm long and 4 to 15 cm wide, alternate,
ovate to elliptic oblong, obtuse to acute, base obtuse to subcordate, usually shallowly
sinuately 2- to 4-lobed, both surfaces densely or sparingly stellate-tomentose; petioles, 1
to 10 cm long; inflorescences, solitary or 2- to 5-flowered cymes, opposite to subopposite
the leaf axils, the lower flowers bisexual the upper ones male; pedicels, 1 to 3 cm long,
elongating and enlarging in fruit; calyx, about 2 cm long, narrowly 5- to 7-lobed, lobes,
1 to 1.5 cm long, acuminate, enlarging and sometimes splitting in fruit, stellate-hairy,
persistent; corolla, 3 to 5 cm in diameter, 5- to 6-lobed, campanulate, lobes about the
length of the tube, broadly-triangular, incurved, hairy beneath, glabrous within, pale
violet to purplish-violet; stamens, 5 to 6, about 1 cm long, free erect, yellow, with short
filaments; anthers, 5 to 8 mm long, yellow; fruit, 5 to 30 cm long and 3 to 8 cm across,
pendent, highly variable in shape depending on the cultivar, subglobose, oblong,
cylindric or obovoid or pyriform, a fleshy berry with leathery skin, smooth, shiny,
purple-black to purple, white, yellowish or striped; seeds, many, compressed, pale
yellow or light brown, Food plant in home gardens. 5, 6.

Solanum tuberosum L. "potato", "Irish potato"

Recent introduction. S$. America-Andes Mts. Rare. Branched, sprawling annual
herb, up to | m high, with a mass of fine, fibrous, adventitious roots and swollen stem
tubers borne on underground stolons; leaves, 10 to 25 cm long, pinnate, with small
interjected leaflets between larger leaflets (main pinnae); larger leaflets, 5 to 9, ovate;
inflorescences, leaf-opposed, few-flowered cymose panicles; peduncles, 5 to 10 cm long;
calyx lobes, 5 to 8 mm long, lanceolate; corolla, about 2 cm in diameter, subrotate,
white or pale violet to purple or bluish; fruit, 1.5 to 2 cm in diameter, subglobose, a
small inedible berry; seeds, suborbicular to subreniform, green or yellowish. Immature
food plant at Location. 5, 6, 7.

STERCULIACEAE (Cocoa Family)

Waltheria indica L. "waltheria"
syns. W. americana L.; W. elliptica Cav.

Recent introduction. Pantropical. Occasional. Downy perennial subshrub, 0.3 to 1
m high, with erect, branched tomentose stems and woody near base; leaves, 1 to 6 cm
long and 0.5 to 3 cm wide, alternate, ovate-oblong, crenate or serrate, velvety-pubescent
with mixed stellate and simple hairs, prominently veined; petioles, about 1 to 1.5 cm
long; stipules, narrow, caducous; inflorescences, axillary, dense cymose clusters;

201

peduncles, up to 4 cm long; bracts 3, linear, caducous; calyx, about 6 mm long, 5-lobed;
corolla, slightly longer than calyx, 5-lobed, clawed, yellowish-orange to yellow,
persistent; stamens 5, connate at base; ovary, l-celled, sessile; fruit, a 2-valved capsule;
seeds 1, rarely 2, exalate. Weed in waste places, vacant lots and ruderal habitats on
coastal strip. 3(58668), 4(112N), 5(5, 109a), 6, 7(22301), 8(9599).

SURIANACEAE

Suriana maritima L.
gie", gie cool? (T)

Indigenous. Pantropical. Rare. Shrub or small tree with pubescent young growth,
1 to 4 m high, often dwarfed and wind-trimmed when exposed to salt spray; leaves, 1.5
to 3 cm long and 2 to 6 mm wide, alternate, crowded, narrowly obovate-oblong or
linear-spathulate, acute, base decurrent, very short-petiolate; inflorescences, axillary
racemose clusters; bracts, green, persistent; calyx, 5-lobed, persistent; corolla, 5-lobed,
petals, about 8 mm long, imbricate, yellow; stamens 10; fruit, a 5-parted, 5-angled
capsule, pubescent, brownish, enclosed in persistent calyx, Rare coastal plant. 7(27820)

TILIACEAE (Linden Family)

Muntingia calabura L. "Panama cherry", "Panama berry"
bin (bean)(N)

Recent introduction. Trop. America. Occasional. Small pubescent tree, 3 to 12 m
high, with tiered, slightly drooping branches; leaves, 5 to 10 cm long and 2 to 3.5 cm
wide, alternate, crowded, oblong-acuminate, obliquely subcordate, strongly asymmetrical,
thin, serrulate, sticky-pubescent, soon wilting; stipules, about 5 mm long, linear, paired,
unequal, caducous; inflorescences, supra-axillary, 1 or few-flowered clusters; pedicels,
about 2 to 3 cm long; calyx, 5 (rarely 6 or 7)-parted, sepals, about 1 cm long, lan-
ceolate-caudate, tomentose-hirsute; corolla, 5-parted, petals about 12 to 13 mm long,
broadly spathulate-deltoid, white or pink-tinged; stamens about 75, yellow; filaments,
about 6 mm long, slender, distinct, white; anthers, small, rounded, yellow; fruit, 1 to 1.5
cm in diameter, subglobose, berry-like (baccate), S-celled, light-red to yellowish, sweet,
juicy, edible; seeds, many, about 0.5 mm long, elliptic, grayish-yellow, embedded in
pulp. Planted ornamental fruit tree; naturalised and spreading in strip-mined areas in
interior Baiti District. Ripe fruit eaten as a snack food, primarily by children. 3(58752),
4(139N), 5(91), 6, 7.

202

Triumfetta procumbens Forst.f. "beach burr"
ikiau, ikiow, igiau, giau (B)(N); te kiaou (K); tolotolo, kiaou (7)
syn. T. fabreana Gaud.

Indigenous. Paleotropics. Rare. Pubescent prostrate subshrub with long-trailing,
densely-stellate pubescent branches up to 3 m long; leaves, 1.5 to 6 cm long and nearly
as wide, alternate, broadly ovate to suborbicular or shallowly 3-lobed, rounded to obtuse,
base rounded to subcordate, crenate-serrate, densely stellate-woolly beneath, palmately 5-
to 7-nerved; petioles, up to 6 cm long; inflorescences, axillary, pedunculate, subumbel-
late clusters; calyx, 5-lobed, with a cornate protuberance at tip; corolla, 10 to 12 mm
across, 5-parted, petals bright yellow; stamens, numerous and distinct; fruit, 6 to 12 mm
in diameter, subglobose, spiny, burr-like, stellate-pubescent. Growing along coastal strip
and in open areas in coastal thickets. Probably incorrectly reported as 7. semitriloba
Jacq. by Burgess in 1935. Juice from crushed leaves used medicinally to cure filariasis
and fever, to retard hair from falling out, and for a gelatinous post-natal medicine which
is drunk by mothers to help rid them of afterbirth; crushed leaves and stems also mixed
with toddy (kerawai) and used as poultices on boils. 1, 2, 4(111N), 5(109), 6, 7(27803).

URTICACEAE (Nettle Family)

Laportea ruderalis (Forst.f.) Chew
luna, aluna, pakisikisi (T)
syns. Urtica ruderalis Forst.f.; Fleurya ruderalis (Forst.f.) Gaud. ex Wedd.;
Schykowskya_ ruderalis (Forst.f.) Endl.; Boehmeria paniculata Gaud.;
Fleurya paniculata Gaud.

Indigenous. Malayo-Pacific. Rare. Erect, un- or few-branched, slightly fleshy,
glabrous herb, up to 40 cm high; leaves, 1.5 to 8 cm long and 1 to 4 cm long, alternate,
obtuse, obtuse or acute, base truncate or subcordate, coarsely crenate; petiole, as long as
blade; inflorescences, axillary, pedicellate, in open paniculate clusters, shorter than
petiole; male flowers, 4-rarely, 3- or 5-merous; female flowers, 4-merous, perianth with
unequal lobes; fruit, obliquely ovate, an achene partly enclosed in the persistent perianth.
Growing in shady areas near caves and in moist habitats at the base of the limestone
escarpment. 5(99), 6, 7(27809).

Pellionia sp.

Recent introduction. Trop. Asia. Small, creeping, succulent, perennial herb;
leaves, 2.5 to 7 cm long, oval, attractive, variegated with combinations of purple, light
green, bronzy-green or gray depending on the species and cultivar; inflorescences,
inconspicuous, greenish. Pot plant. 6.

203

Pilea cardieri Gagn. & Guill. “aluminum plant"

Recent introduction. Vietnam. Occasional. Erect, branching, somewhat succulent
when young, perennial herb, up to 40 cm high, with somewhat 4-angled stems and
swollen internodes; leaves, up to 9 cm long and 5 cm wide, ovate, paired, dentate near
apex, dark-green with silvery blotches above, light green beneath; petioles, 2 to 3 cm
long; inflorescences, axillary, long-pedicellate, globose clusters; perianth segments, dull
white or slightly pink-tinged. Pot plant. 5, 6.

Pilea microphylla (L.) Liebm. "artillery plant"
syns. Parietaria microphylla L.; Pilea muscosa Lindl.

Recent introduction. Trop. America. Occasional. Low, delicate, much-branched,
succulent, glabrous, perennial herb, up to 30 cm high, with tender greenish or purplish
stems; leaves, 3 to 6 mm long and 1 to 2 mm wide, opposite, 2-ranked, crowded, elliptic
to spathulate or obovate; inflorescences, axillary, cymose clusters; individual flowers,
minute, unisexual, minute, greenish, sometimes red-tinged; female flowers, 3-tepaled,
tepals, unequal; stamens, opening explosively. Weed in pot plants and under planted
ornamentals. 3(58713), 5(8), 6, 7.

Pilea nummularifolia (Sw.) Wedd. "creeping Charlie"

Recent introduction. Tropical America. Occasional. Creeping perennial herb, up
to 20 cm high, rooting at the nodes; leaves, up to 2.5 cm in diameter, oval, crenate,
succulent, pubescent, 3-veined from base; inflorescences, rose-red. Pot plant; ground
cover in shaded areas. 5, 6.

VERBENACEAE (Verbena Family)

Clerodendrum inerme L. var. oceanicum A. Gray "beach privet"
eamwiye, eamwije, eyamwiye, eyamwije (N); te inato (K); inato (T)
syns. Volkameria inermis L.; Clerodendrum nereifolium Wal.; C. commersonii
(Poir.) Spreng.

Indigenous. Indomalaysia, Australia and the Pacific Is. Common. Erect, trailing
or scrambling, sometimes shrubby, glabrous perennial, up to 5 m high; leaves, 3 to 13
cm long and | to 7 cm wide, opposite, or sometimes in whorls of 3, elliptic to obovate
or narrowly lanceolate, short-acuminate or cuspidate to somewhat blunt, base acute,
glabrate, thinly fleshy, pinnately nerved, dotted beneath with a few sunken glands

204

basally; petiole, 0.4 to 2 cm long, slender, leaf scars prominently somewhat raised;
inflorescences, axillary, supra-axillary or pseudoterminal, 3- to 12-flowered cymose
clusters; peduncle, 1 to 4 cm long, slender, subglabrate; pedicels, 3 to 6 mm long;
bracteoles, small, few; calyx, 4 to 6 mm long, campanulate, subtruncate and minutely 5-
denticulate, enlarging somewhat as fruit matures; corolla, up to 3.5 cm long, hypocrateri-
form or slender-tubular, 5-lobed, lobes, 5 to 8 mm long, ovate, subequal, somewhat
expanded distally, bluntly acute at apex, glabrous white or pinkish along tube, fragrant;
stamens 4, | cm long, exserted, inrolled in bud, filaments red, purple or pinkish;
anthers, yellow; style, exserted to 20 mm, purple to rich pink distally, paler proximally;
fruit, about 1 to 1.5 cm in diameter, obovoid, drupaceous, separating into four 1-seeded
nutlets, yellow-green to dark-brown, turning black at maturity, subtended by a indurate,
striate-venose calyx; seeds, oblong. Abundant on limestone cliffs and pinnacles, forming
luxuriant curtains near edge of escarpment and on parts of coastal strip in Anetan
District; occasionally a planted ornamental. Fragrant flowers used in garlands; leaves
reportedly pounded and used as a cure for leprosy in the past (cure reportedly received in
a dream). 2, 3(58664), 4(103N, 166N), 5, 6, 7(27817).

Clerodendrum paniculatum L. "pagoda flower"

Recent introduction. E. Trop. Asia. Rare. Perennial shrub, up to 1 to 2 m high,
with medullose to hollow, bluntly quadrangular branchlets, the nodes with a broad band
of hairs; leaves, 4 to 15 (-40) by 7 to 20 (-38) cm, opposite, broadly ovate, prominently
3 to-7-lobed or the uppermost ones entire, base cordate, the lobes acute to short-acu-
minate and remotely dentate to crenate or entire, thin chartaceous to membranaceous,
minutely strigillose to glabrous above, squamose with peltate scales beneath; inflorescen-
ces, axillary or terminal, many-flowered panicles, up to 45 cm long and broad; pedicels,
4 to 15 mm long, filiform, reddish; calyx, 3 to 5 mm long, campanulate, deeply-divided,
red to orange-red; corolla, hypocrateriform or narrow-tubular, the tube, up to 2 cm long,
slender, the limb spreading to 1 cm across, orange-red to vivid scarlet (rarely white),
puberulent; stamens and style exserted 2.5 to 3.5 cm beyond corolla throat; fruit,
drupaceous, greenish-blue to black at maturity. Planted ornamental. 5, 6(235), 7.

Clerodendrum thomsoniae Balf.f. "bleeding heart"

Recent introduction. W. Africa. Rare. Perennial shrub or climbing vine, up to 3
m or higher, with slender, distally 4-angled and terete proximal branchlets and twining
older branches; leaves, 5 to 15 cm long and 2.5 to 7 cm wide, elliptic to elliptic-ovate,
acute to short-acuminate, base subcordate to subacute or rounded, glabrate on both
surfaces or puberulent on venation; petioles, 8 to 30 mm long; inflorescences, terminal or
distally axillary, cymose clusters, to 12 by 15 cm; peduncles, 2.5 to 6.5 cm long,
slender; pedicels, 7 to 16 mm long, slender; calyx, 2 to 2.7 cm long, deeply 4- to 5-
lobed, lobes sharply acute, pale yellow-green, becoming white or pinkish in flower, red-
violet in fruit, prominently veined; corolla, 2.2 cm long, hypocrateriform or tubular, the

205

tube, 1.5 to 2.5 cm long, slender, greenish-red, 5-lobed, lobes up to 1 cm long, deep red
to scarlet or crimson distally, the limb spreading or reflexed, to 2 cm broad; stamens and
style, long-exserted, pale green; fruit, a drupe, glossy, black, with a brilliant red aril
uniting the 4 pyrenes; seeds, 4, nutlets. Planted ornamental. 3(58794), 5, 6, 7(27817).

Duranta repens L. "golden dewdrops", "golden eardrops"
D. erecta L.; D. plumieri Jacq.

Recent introduction. Trop. America. Rare. Erect perennial shrub or small tree, up
to 4 m or higher, with slender, arching, drooping or trailing branchlets and sometimes
bearing axillary or suppressed spines; leaves, 1.5 to 7 cm long and 0.5 to 4 cm wide,
opposite or in whorls of 3, elliptic or ovate, rarely elliptic-obovate, obtuse to acute to
acuminate or apiculate, base cuneate, entire or distally serrate, usually thin-textured,
glabrate on both surfaces; petioles, 1 to 8 mm long, slender; inflorescences, axillary or
terminal, erect or recurved, loose, many-flowered paniculate clusters or racemes, 5 to 30
cm long; pedicels, 1 to 5 mm long; bracteoles, minute or occasionally subfoliaceous;
calyx, 3 to 4.5 mm long at anthesis, tubular, angular, minutely toothed or subulate;
corolla, about 8 mm long, surpassing calyx by about 2 to 5 mm, tubular, 5-lobed, 7 to
14 mm across the limb, pale purple to bluish-purple, 2 slightly smaller lobes with a
purple stripe and 3 slightly larger plain lobes, fragrant, puberulent on both surfaces;
stamens 4; fruit, 6 to 12 mm in diameter, a globose, 8-seeded drupe, juicy, yellow to
orange, enclosed by the enlarged, curved-beaked, yellowish calyx. Planted ornamental. 5,
Oni

Lantana camara L. var. aculeata (L.) Mold. "lantana"
magiroa (N); te kai buaka (K); kaipuaka (T)

Pre-World War II introduction. Trop. America. Occasional. Erect or rambling,
more or less pungent-odorous, perennial shrub, | to 5 m high, the stems and branches
usually armed with short, recurved prickles; leaves, 2 to 12 cm long and 2 to 6 cm wide,
opposite, decussate or whorled, ovate to oblong-ovate, acute to short-acuminate, base
abruptly rounded and cuneate, crenate-serrate, stiff chartaceous, reticulate-rugose and
scabrous above, downy or hispid,below, pinnately nerved, spicy pungent if rubbed;
petioles 0.7 to 2 cm long; inflorescences, axillary, crowded, hemispherical, many-
flowered, corymbose heads, 2 to 4 cm across; peduncle, 2 to 9 cm long, slender;
bracteoles, 4 to 9 mm long and | to 1.5 cm wide, oblong to lanceolate, acute to subulate
at apex; calyx, about 3 mm long, 2-lobed, inconspicuous; corolla, 10 to 12 mm long,
tubular or hypocrateriform, 4 to 5-lobed, 5 to 8 mm across the limb, pubescent internal-
ly, white, pink, orange and red with a darker eye, the youngest flowers central, pale, the
older flowers orange, the oldest ones red (or white); stamens 4; fruit, 3 to 4 mm in
diameter, a fleshy, slightly juicy drupe, purple-black; seeds 1, a 2-celled pyrene with an
air cavity between the cells. Planted ornamental; naturalized in ruderal sites and in
unmined open plateau forest; occasional in older revegetated strip-mined sites. Flowers

206

used in garlands. The first Nauruan to plant lantana was reportedly a woman by the name
of Magiroa who stole it from a garden of an expatriate British Phosphate Company
employee, hence the Nauruan name. 2, 3(58599, 58798), 4(101N), 5(42), 6, 7, 8(9563).

Lantana camara L. var. "drap d’or" “cloth of gold lantana"

Recent introduction. Trop. America. Rare. Cultivar with single-colored yellow
flowers. Planted ornamental. 3, 6.

Premna serratifolia L. "prermna"
idibiner, idibinerr (N); te ango (K); aloalo, valovalo, te ango (T)
syns. P. obtusifolia R. Br.; P. gaudichaudii Schauer; P. mariannarum Gaud. ex
Schauer; P. integrifolia L.; P. taitensis Schauer; P. corymbosa (Burm.f.)
Rottl. & Willd.; P. alba Lam.; P. angustifolia Lam.; P. paulobarbata
Lam.

Indigenous. Indopacific. Common. Small, nearly glabrous shrub or small tree, up
to 5 m or higher, with minutely puberulent youngest branches; leaves, 2 to 15 cm long
and | to 9 cm wide, opposite, elliptic to oblong-ovate or suborbicular, subacuminate to
blunt, sometimes short-mucronate, base rounded or obtuse to subcordate, chartaceous,
shiny, usually ventrally concave, glabrate or minutely puberulent dorsally on nerves, 4 to
7 pairs of lateral nerves; petioles 0.5 to 6 cm long; inflorescences, terminal, sometimes
also axillary, richly-branched, many-flowered, corymbose clusters, 3 to 10 cm or more
across; peduncles, up to 2 cm long; pedicels, up to 1 mm long; bracts, to 5 mm long,
paired at each node, subulate to narrowly lanceolate, persistent; calyx, 1.5 to 2 mm long
at anthesis, cupular, bilabiate, minutely 4- to 5-toothed or subtruncate, minutely puberu-
lent on both surfaces; corolla, 2.5 to 4 mm long, tubular or short-hypocrateriform to
subrotate, tube up to 2.5 cm long, 4- to 5-lobed, somewhat bilabiate, the longest lobe to
1.5 cm long, greenish; stamens 4 or rarely 5, 1.2 to 2.4 mm long, 2 longer, 2 shorter,
very slightly exserted or included; fruit, 2 to 4 mm across, subglobose, glabrous, green
turning purple-black at maturity. Common tree on coastal strip and in escarpment forest;
common in home gardens. Timber used for house rafters in the past; wood considered to
be among the best firewood for cooking pandanus; leaves boiled with coconut oil to scent
it; flowers used in garlands; young leaves used as poultices to help wounds heal. 1(49.R),
2(8:5)93685975,58633)6"D, 6577810);

Stachytarpheta jamaicensis (L.) Vahl "Jamaica vervain", "blue rat’s tail"
edidubai, edidubaiy (N); te uti ("lice")(K)
syns. Verbena jamaicensis L.; Stachytarpheta indica (L.) Vahl

Pre-World War II introduction. Trop. America. Extinct? Erect, sparsely-
branching perennial subshrub, up to about | m high; leaves, 4 to 11 cm long and 2.5 to 5

207

cm wide, opposite, elliptic-oblong, acute or subobtuse, distally crenate-serrate, decurrent
on the petiole, flat, pale, somewhat glossy, silvery- or irridescent-green, lateral nerves
not prominent dorsally; inflorescences, erect, elongated purplish spikes, 15 to 40 cm
long; individual flowers, axillary to a bract, a few opening at a time, evanescent and
soon caducous, sessile; bracts, 4 to 5 mm long, with a scarious margin; calyx, dorsally
compressed, 4- to 5-toothed; corolla, 5 to 10 mm long and 4 to 12 mm wide tubular, 5-
lobed, pubescent internally, pale violet to purple or whitish; fertile stamens 2, included;
fruit, 4 to 5 mm long, 2-segmented, enclosed in the calyx. Weed reported present by
Burges in 1935, but not reported since. 2.

Stachytarpheta urticifolia Sims "blue rat tail", "false verbena"
edidubai, edidubaiy (N); te uti ("lice")(K)
syn. Cymburus urticifolius Salisb.

Pre World War II introduction. Trop. America. Common. Erect, sparsely-
branching perennial subshrub, 0.5 to 1.5 m high; leaves, 4 to 12 cm long and 2 to 7.5
cm wide, opposite, decussate, ovate-elliptic or ovate, acute or subobtuse, base cuneate-
decurrent, distally crenate-serrate, leaf teeth ciliate, rich green, subglossy, pinnately
nerved, lateral nerves prominent dorsally, bullate (convex between the veins); inflores-
cences, erect, elongated, greenish spikes, 14 to 50 cm long, bearing numerous flowers in
groove-like depressions on the rachis; peduncle up to 3 cm long; individual flowers,
axillary to a bract, a few opening at a time, evanescent and soon caducous, sessile;
bracts, 4 to 7 mm long, erect, lanceolate, aristate at apex; calyx, 5 to 7 mm long,
dorsally compressed, 4- to 5-toothed; corolla, up to 8 mm long, tubular, 5-lobed,
pubescent internally, medium violet to dark blue-violet; fertile stamens 2, included,
white; style, white; fruit, 2-segmented pyrene, enclosed in a persistent calyx. Common
weed in waste places and ruderal habitats, especially along roadsides. Mature dried black
fruits reportedly eaten by some children. 3(58632), 4(109N), 5(12), 6, 7, 8(9543).

Vitex negundo L. var. bicolor (Willd.) H.J. Lam "blue vitex"
dagaidu, dogaidu (N); te kaitu? (K)
syn. V. trifolia L. var. bicolor (Willd.) Mold.

Indigenous. E. Africa to the Pacific Is. Occasional. Erect, aromatic, freely-
branching shrub or small tree, 1.5 to 8 m high, with 4-angled puberulent branches;
leaves, opposite, 3- to 5- (rarely 7-) foliate; leaflets, oblong-elliptic to ovate-lanceolate,
acute to subacuminate, the central one longest, up to 11 cm and 3 cm wide, with
petiolules up to 1 to 2 cm long, the other leaflets smaller on shorter petiolules, pinnately
nerved, dark above, densely white-tomentose beneath, aromatic; petioles, 2.5 to 6 cm
long; inflorescences, terminal or axillary, narrow, branching, puberulent, paniculate
clusters; calyx, about 3 mm long, 5-toothed, gray-puberulent; corolla, about 4 mm long,
bilabiate, 5-lobed, blue-violet, puberulent; filaments and style blue to purple; fruit, 5 to 6
mm long, subglobose drupe, 1-seeded, black. Occasional in some forest stands on coastal

208

strip in low-lying areas near base of escarpment and in some home gardens. Branches
used for fishing rods for small fish; flowers and leaves used in garlands and other body
ornamentation; seeds used to make garlands; young leaves and meristem crushed with
coconut oil as a cure for fever blisters; juice of fruit drunk as a cure for fits and
convulsion. 2, 3(58635), 4(119N), 5(19, 56), 6, 7, 8(9584).

VITACEAE (Grape Family)

Cissus sp. "cissus"

Recent introduction. Trop. Asia? Rare. Thick-stemmed vine; leaves, cordate;
flowers not seen. Planted ornamental. 6.

Vitis vinifera L. "grape"

Recent introduction. S.E. Europe to India. Rare. Tendril-bearing, woody vine;
leaves, 5 to 10 cm across, subcordate in outline, usually palmately lobed, usually serrate,
more or less glabrate above, white- or reddish-tomentose beneath; inflorescences, leaf-
opposed panicles; flowers bisexual and male, 5-merous; calyx, subtruncate; corolla, 5-
parted, petals coherent distally, the calyptra caducous before full anthesis; fruit, 1 to 2
cm in diameter, and ovoid to suglobose, 2- to 4-seeded berry, juicy, edible; seeds,
rostrate at base, light brown. Single immature plant at Location. 5, 6.

209

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216

Appendix I. Class, subclass, family and antiquity status of the vascular flora of Nauru.

Class/Subclass Indi- Abori- Recent Total
Family genous ginal

PTERIDOPHYTA (Ferns and Fern Allies)

Aspleniaceae
Davalliaceae
Ophioglossaceae
Polypodiaceae
Psilotaceae
Pteridaceae

=e NOR NOR
' 1
' '
Were NRK We

Subtotal 8 - 3 11

GYMNOSPERMAE

Araucariaceae - - 1 1
Cycadaceae - - 1 1

Subtotal - - 2 D,

ANGIOSPERMAE (Flowering Plants)
MONOCOTYLEDONS

Araceae - -
Arecaceae - 1
Bromeliaceae = =
Cannaceae - -
Commelinaceae - -
Cyperaceae 2 -
Dioscoreaceae = =
Iridaceae - =
Liliaceae - <
Marantaceae - -
Musaceae - -
Orchidaceae -
Pandanaceae 1

Poaceae 3 -
Pontederiaceae -

eBerawvaPunauHanad
SS we ~
MmMANNAWARWNWMNYE HIG

Strelitziaceae
Taccaceae
Zingiberaceae

Subtotal

DICOTYLEDONS

Acanthaceae
Amaranthaceae
Anacardiaceae
Annonaceae
Apiaceae
Apocynaceae
Aquifoliaceae
Araliaceae
Aclepiadaceae
Asteraceae
Balsaminaceae
Basellaceae
Begoniaceae
Bignonicaceae
Bombacaceae
Boraginaceae
Brassicaceae
Cacataceae
Capparidaceae
Caricaceae
Casuarinaceae
Chenopodiaceae
Clusiaceae
Combretaceae
Convolvulaceae
Crassulaceae
Cucurbitaceae
Ericaceae
Euphorbiaceae
Fabaceae
Gentianaceae
Geraniaceae
Gesneriaceae
Goodeniaceae
Hernandiaceae
Lamiaceae

137,

Ww
—

ON NK IF WORK NNOD ! RBPWWENFEWIREFPONANGS

145

—
—

QS Se ee OM comm OS est ome wm Oem pO a

27,

218

Lauraceae
Lecythidaceae
Lythraceae
Malpighiaceae
Malvaceae
Meliaceae
Moraceae
Moringaceae
Myrtaceae
Nymphaeaceae
Nyctaginaceae
Oleaceae
Onagraceae
Oxalidaceae
Passifloraceae
Piperaceae
Polygalaceae
Polygonaceae
Portulacaceae
Rhamnaceae
Rhizophoraceae
Rosaceae
Rubiaceae
Rutaceae
Sapindaceae
Sapotaceae
Saxifragaceae
Scrophulariaceae
Solanaceae
Sterculiaceae
Surianaceae
Tiliaceae
Urticaceae
Verbenaceae
Vitaceae

Subtotal

GRAND TOTAL

1 Zoe — — 1!

45

59

1 2
- 1
ll 1
2 2
12 16
2 2
5 8
1 1
+ 4
1 1
3 4
2 2
1 1
1 1
3 3
2 2
1 1
2 2
2 2
- 1
- 1
2 2
12 16
5 5
1 Z
1
1
1
10 10
1
- 1
1 2
4 2,
6 2
1
287 335
429 493

219

Appendix II. Nature and ecological and cultural (ethnobotanical) importance of coastal
plant species of the tropical Pacific Ocean (Notes: 1) Under "Habitat", O = outpost
strand zone, I = inner littoral zone, M = mangrove habitats, W = coastal wetland or
marshes; N = also found naturalized or wild in non-coastal habitats, and C = cultivated
or planted; 2) Under "Origin", I = indigenous, A = aboriginal introduction, R = recent
post-European contact introduction, and ? = status unsure; 3) Under "Importance",
"Eco" = ecological importance in coastal plant communities and "Cult" = cultural
importance in terms of a species’ range throughout the Pacific islands or its overwhel-
ming importance in some localities, with +++ = very important in most island groups,
with multiple usage in terms of cultural importance, ++ = of considerable importance
in some island groups or some important uses locally, + = present in some island
groups or of some use in restricted localities, and - = of minor ecological importance or
no cultural uses reported from Melanesia, Polynesia, or Micronesia).

Latin Name Habitat Origin Importance
Eco Cult

FERNS
Acrostichum aureum I,W,M I ++ +
Asplenium nidus I,N I apap SP op
Davallia solida I,M,N,C I + +
Nephrolepis spp. I,M,N,C I span) SP
Polypodium scolopendria I,N I +E tt
Pteris spp. I,N I + ap
Pyrrosia adnascens I,N,M I + +
Stenochlaena palustris I,N,M,W I ap +
Tectaria spp. I,N I =P ala
Thelypteris spp. W,I,N I jaf emaets

HERBS
Achyranthes spp. I A? oe iieataiats
Boerhavia spp. O,I I ap Spar
Crinum asiaticum OFC I?,A?,R aF ++
Dendrobium spp. I,M,N,C I 3 +
Hedyotis spp. O,I I aR -
Heliotropium spp. O,I I ap ar
Hymenocallis littoralis OC R + +
Laportea spp. I,N I te ots

220

Lepidium bidentatum
Peperomia spp.
Portulaca australis
Portulaca pilosa
Procris pedunculata

Sesuvium portalucastrum

Tacca leontopetaloides
Taeniophyllum fasicola
Triumfetta procumbens

GRASSES AND SEDGES

Cyperus javanicus
Cyperus laevigatus
Cyperus polystachyos
Digitaria setigera
Eleocharis spp.
Fimbristylis cymosa
Ischaemum spp.
Lepturus repens
Paspalum spp.
Sporobolus spp.
Stenotaphrum spp.
Thuarea involuta

VINES AND LIANAS

Abrus precatorius
Canavalia cathartica
Canavalia rosea
Canavalia sericea
Cassytha filiformis
Derris trifoliata
Entada phaseoloides
Epipremnum pinnatum
Hoya australis
Ipomoea littoralis
Ipomoea macrantha
Ipomoea pes-caprae
Mucuna gigantea
Vigna marina

I,M
I,N
O,I,N,C
O
O,I,N
OFM
I,N
I,N
I,N
O,I,N
O,I,N
O,W
I,N
OC

Llosa lelalali tet) L
~~

De A ee A eee A cree BO cee ce BO cee cee ee ce
-

ate
= hot eee

++eeeettt
+

+ +
+

+ +a+4+ +
sb ee ot acta ae
a

+ +
+ +
++ ++
+ +
++ ++
++ ++
++ +++
+ ++
+ +
++ +
++ ++
+++ ++
+ +
+++ ++

SHRUBS

Abutilon spp.
Acanthus ebracteatus
Allophylus timoriensis
Bikkia tetrandra
Caesalpinia bonduc
Canthium spp.
Capparis spp.
Clerodendrum inerme
Colubrina asiatica
Desmodium umbellatum
Dalbergia candenatensis
Dodonaea viscosa
Euphorbia chamissonis
Gardenia taitensis
Geniostoma spp.
Jossinia reinwardtiana
Nypa fruticans
Pemphis acidula
Scaevola taccada

Sida fallax

Sophora tomentosa
Suriana maritima
Timonius spp.
Tephrosia purpurea
Wollastonia biflora
Wikstroemia spp.
Ximenia americana

TREES

Acacia simplex

Aidia cochinchinensis
Avicennia maritima
Barringtonia asiatica
Barringtonia racemosa
Bruguiera gymnorhiza
Calophyllum inophyllum
Casuarina equisetifolia
Cerbera manghas
Ceriops tagal

Cocos nucifera

>

co re ce ee re BO cree BO ce ee ee en ee cee ee ce BO ee ee ce en cen ee ee Bl ee ee le |

| ee eee cee cee ee |

.

w

.

nw

De es
ds

—
~~

221

+ +
Aiea et abo

those ab ohh
+
+
+

+
pais se ae ee
+
+t +4

eae ee hort
1a

++ 44+
++++4++
+++4+4+4+
+ ++

++ ++
+ +
++ ++
+++ ++
+ +
+++++4+
++++++
++++++
+ ++
+ +
++++++

222

Cordia subcordata
Cycas circinalis
Cynometra spp.
Diospyros spp.
Dolichandrone spathacea
Erythrina fusca
Erythrina variegata
Excoecaria agallocha
Ficus obliqua

Ficus prolixa

Ficus scabra

Ficus storkii

Ficus tinctoria
Glochidion spp.
Grewia crenata
Guettarda speciosa
Gyrocarpus americanus
Heritiera littoralis
Hernandia sonora
Hibiscus tiliaceus
Inocarpus fagifer
Intsia bijuga
Leucaena insularum
Lumnitzera littorea
Mammea odorata
Manilkara spp.
Metroxylon spp.
Morinda citrifolia
Neisosperma oppositifolia
Pandanus tectorius
Phaleria disperma
Pipturus argenteus
Pisonia grandis
Pittosporum arborescens
Planchonella costata
Polyscias spp.
Pongamia pinnata
Premna serratifolia
Rhizophora spp.
Santalum spp.
Serianthes spp.
Sonneratia alba
Soulamea amara

Syzygium spp.

O,1,C
I,N,C
M,C
I,N

M,I
M,W,I
O,I,N,C
M,O,W

O;1,M, NéP
M,W,N,C
I,N,M

O

M,lI

I,M

I,N

WN
I,N,P

Ol
O,I,N,W,P
I,N,C

I,N

OC

I,N

I,N

I,N

Le A cere ee cee BO eee cee ce BN ee BE ee ee oe ee

++++4+
+. ++
+ +
+ +
+ +
+ +
+++4++
++ +
++ ++
++ +++
+ +
+ +
++ +++
+ ++
+ ++
+++++
+ ++
+ +
++ ++
+++4+++4+
+++++
+ ++
+ +
++ ++
+ +
+ +
++ ++
+++4++
++ +4
++++4++
+ ++
++ ++
++ ++
+ +
+ +
+ +
+ +
+++4++
++++4++
+ ++
+ +
+ +
+ +
+ +

223

Terminalia catappa O,I,N,C I,A,R fee ee
Terminalia littoralis O,I,C I + +
Thespesia populnea O,I I arr) SRSA
Tournefortia argentea O,I,C I TP oP SP APSE aR
Vavaea amicorum I,N I + Te
Vitex spp. O,I,N,P I ae ae) ear
Xylocarpus spp. M,I I + ++

Sources: An extensive review of the available literature and personal records and
observations by the author; for full listing of sources, see Thaman, 1992.

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ATOLL RESEARCH BULLETIN

NO. 393

FLORA OF THE PHOENIX ISLANDS, CENTRAL PACIFIC

BY

F. RAYMOND FOSBERG AND DAVID R. STODDART

ISSUED BY
NATIONAL MUSEUM OF NATURAL HISTORY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C., U.S.A.
FEBRUARY 1994

AM 2) MURS LAAOITAr
rrr7yen“i AMMA Tee
uM YOroriteaw
or YEAURERS

FLORA OF THE PHOENIX ISLANDS, CENTRAL PACIFIC

BY F. RAYMOND FOSBERG! AND DAVID R. STODDART2

ABSTRACT

This paper lists the vascular flora of the three atolls (Canton, Gardner, Hull) and
five small islands (Birnie, Enderbury, McKean, Phoenix, Sydney) of the Phoenix Group,
located in the arid equatorial belt of the Central Pacific, based on extensive collections
made in 1973 and 1975 and on previous records and collections. The flora includes 87
species in 36 families. Only 28 of the species (32%) are considered native. A further 60
species have been recorded in the literature, many of them deliberate introductions which
have not persisted. These are mentioned but not described.

INTRODUCTION

The Phoenix Islands lie in the geographic center of the Pacific Ocean, at
approximately 170°W longitude and 5°S latitude (Figure 1). They comprise eight islands
and atolls (Figure 2). Canton, Gardner (Nikumaroro) and Hull (Orona) are atolls;
McKean, Birnie, Sydney, Phoenix and Enderbury are smaller islands with residual
enclosed lagoons. Locations and areas are as follows:

Longitude Latitude Land area

Atolls:

Canton 171°41'W 2°49'S 10.9 sq km

Gardner 174°31'30"W 4°40'30"S 6.7 sq km

Hull 172°11'W 4°31'S 4.3 sq km
Islands:

Birnie 171°31'W 3°35'S 62 ha

Enderbury GAUSS IIS WAS) 3°77 45S 745 ha

McKean 174°7'30"S 3°35'45"S 91 ha

Phoenix 170°42'45"W 3°43'15"S 94 ha

Sydney 171°14'45"W Ailes 1067 ha

| Department of Botany, National Museum of Natural History, Smithsonian Institution,
Washington, D.C. 20560, U.S.A.

2 Department of Geography, University of California at Berkeley, Berkeley, California
94720, U.S.A.

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PHOENIX ILS canton
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Figure 1. Location of the Phoenix Islands in the Central Pacific.

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4

Land areas are wholly formed of reef derived materials and mostly rise only a few meters
above sea level. The highest measured elevations are 7.0 m on Enderbury, 6.3 m on
Sydney, 6.0 on Birnie, 5.5 m on Gardner, 5.4 m on Canton, McKean and Phoenix, and
4.0m on Hull. The group is located east of Kiribati (of which it forms a part) and
Tuvalu, north of the Tokelau Islands, and south of the equatorial islands of Howland and
Baker. The nearest high islands are those of Samoa and Fiji to the south and southwest.

All the Phoenix Islands are dry, with great fluctuations in annual rainfall and
periods of severe drought. Canton in the north has a mean annual rainfall of 710 mm
(1942-67, 1972-75), and the central tier of small islands (McKean, Birnie, Phoenix,
Enderbury), though lacking instrumental records, are probably equally dry. The southern
group of Hull, Sydney and Gardner are rather wetter and support scrub forest and coconut
groves. Mean annual rainfall at Hull (1952-63) is 1171 mm, with a highest recorded
annual total of 2599 mm and a lowest of 565 mm; Sydney has a mean (1948-49, 1952-61)
of 985 mm (Highest 1846 mm, lowest 463 mm); and Gardner a mean (1951-61) of 1319
mm (highest 2722 mm, lowest 275 mm).

From May 29 to June 12, 1973 a party from the Smithsonian and the Royal
Society was able through the courtesy of the U. S. Air Force SAMTEC Project, to visit
and study the flora and vegetation of six of the eight islands in the group, and to make
collections needed to document the vascular floras of these islands. The party was
accommodated on Canton, and Birnie, Enderbury, Phoenix, Hull and Sydney were visited
for periods of one to several days. The visit occurred toward the end of a major El Nifio
event: the total rainfall at Canton over the nine-month period July 1972 to March 1973
was 2629 mm. The vegetation was perhaps at its best during this visit and excellent
collections were made, including a number of species not previously recorded from
certain of the islands. The best sets of the specimens collected have been deposited in the
U. S. National Herbarium and in the herbaria of the B. P. Bishop Museum, Honolulu, and
the Royal Botanic Gardens, Kew, England. Others have been sent to several other
institutions with Pacific interests.

In 1975, from March 19 to April 2, the same party revisited the Phoenix Islands,
again under USAF/SAMTEC auspices, and was able to fill gaps in the 1973 collections,
and to observe changes in vegetation and in the abundance of some species. This was a
normal arid year in the Phoenix Islands. Canton total rainfall for the nine months July
1974 to March 1975 was 255 mm (for the same period in 1973-74 it had been only 155
mm). The effects of dry conditions were studied, as well as the responses of some of the
species to extreme drought. All eight of the islands in the archipelago were visited,
including especially Gardner and McKean, which could not be reached in 1973.
Excellent collections were gathered on these two islands.

The climatic conditions in the Phoenix Islands are marginal for human settlement,
and the group is at present uninhabited. There is archaeological evidence of Polynesian
settlement, but it was probably sporadic. The islands were visited by whalers and guano
diggers during the nineteenth century, when small settlements were established on the
wetter southern islands and efforts made to establish coconut plantations. Hull, Sydney
and Gardner were the site of a Gilbertese colonization project beginning in 1938-40.
Climatic conditions proved marginal for cultivation, however, and following a severe

5

drought in 1960 the scheme was abandoned in 1962 and the colonists resettled in the
Solomon Islands. Canton was occupied by British and American personnel in the late
1930s, pursuing the territorial claims made to the group by both countries, and when
Canton began to be used by trans-Pacific Pan-American flights. In World War II it
became a major air force base, and the U.S. Air Force was still in occupation of Canton
(with small facilities on Enderbury, Hull and Sydney) at the time of our visits. Military
use ended shortly afterwards.

The attempts at colonization and the recent occupation by military and aviation
personnel are responsible for the introduction of a large number of the plant species
recorded below. Experience has shown, however, that the majority of these exotic
species will not persist long without the help of man. In 1950-51 Otto Degener, at the
request of the U. S. Civil Aeronautics Administration, then in charge of Canton Island,
introduced 129 species of plants to the atoll (Degener and Gillaspy 1955, pp. 35-39). Of
these only 14 were observed to be persisting in 1973, and only two, Coccoloba uvifera
and Conocarpus erecta, showed any tendency to spread out of the continually disturbed
areas of present human occupation. Certain other exotic species persist at the sites of
abandoned Gilbertese settlements, and a very few around old guano operations. Some
weeds have been brought in accidentally and have persisted, including Cenchrus
echinatus, Eleusine indica, Portulaca oleracea, Euphorbia hirta, E. hypericifolia,
Pluchea carolinensis, P. indica, Tridax procumbens, and Vernonia cinerea.

Botanical investigations in and collections from the Phoenix Group have been
few. The Wilkes Expedition visited Enderbury in 1840 and 1841 and possibly also
Gardner. John Arundel sent a few specimens to Kew Herbarium in 1882. E. H. Bryan
visited the group in 1924 and again in 1938. His collections are in the B. P. Bishop
Museum, Honolulu. F. R. Fosberg and E. H. Walker collected a few plants by moonlight
during a one hour refueling stop enroute to New Zealand in early February 1949, and F.
R. Fosberg got another small collection during a slightly longer stop on his return trip in
March of the same year. These collections are deposited in the U. S. National Herbarium,
as are a few collected by Dr. Leonard Schultz in 1939. Otto Degener and W. H.
Hatheway collected on Canton in 1950 and 1951, and Dr. and Mrs. Degener in 1958. S.
H. Lamb made a few collections in 1938 which are in the Bishop Museum. Under the
auspices of the Pacific Biological Survey Program of the Smithsonian Institution, in
1964, C. R. Long made extensive collections and observations on the vegetation on all
the Phoenix Islands, and P. Marshall and P. W. Woodward a few more. These are
divided between the U. S. National Herbarium, B. P. Bishop Museum and the University
of Hawaii. A small collection was made by Dr. Katherine Luomala in 1949-1950, now
housed in the Bishop Museum. Roger Clapp brought a small collection from Canton and
Hull Atolls in 1971. On our 1973 expedition 196 numbers were gathered, mostly by
Fosberg and Stoddart, a few by Fosberg and Clapp, and in 1975, 97 numbers were
collected by Fosberg, with at least five sets of duplicates of most of them.

All specimens that we have seen, and those listed by Degener and in manuscripts
by C. R. Long are cited below. If we have actually examined and verified them the
herbaria where they are deposited are cited by Index Herbariorum abbreviations.
Unfortunately, time has not been available to examine many of the rich collections in the
Bishop Museum and the University of Hawaii.

Species considered exotic are indicated by an asterisk *.

A few doubtful species are noted in the discussion. Species mentioned as
growing on Canton by Degener, but not found on this survey are mostly mentioned but
not described. Of great utility during the present investigation were manuscript lists of
species and notes on vegetation made by C. R. Long during the Pacific Biological Survey
and furnished to us by Mr. Roger Clapp, who was also a most agreeable companion and
an encyclopaedic source of information on the Phoenix group during the expeditions.

The late Dr. Marie-Héléne Sachet furnished many records from herbaria and
literature that she had examined and contributed greatly to the organization of the
information included in the paper.

A full account of the Phoenix Islands is near completion and includes a
description of the vegetation of each island. This flora though published separately forms
an integral part of that larger account. The flora is organized alphabetically by family,
and the species named are indexed at the end of the paper.

ACANTHACEAE

*Pseuderanthemum carruthersii var. atropurpureum (Bull) Fosb.

Upright, branched purplish or purplish-green shrub, stems white speckled, leaves
elliptic to elliptic oblong; flowers borne in terminal spikes, corollas salverform, slightly
irregular, limb thickly speckled with dark magenta-crimson.

A single sterile shrub that is probably this plant persists in the old Gilbertese
village on Hull. In sterile condition this is very hard to distinguish with certainty from
Graptophyllum pictum. Planted, also, on Canton in U.S.A.F. area, 1975.

CANTON: Eosberg 55715 (US, BISH). HULL: Fosberg & Stoddart 54795 (US,
HAW, K); Clapp P-71-42 (US).

AIZOACEAE

Sesuvium portulacastrum L.

Fleshy, prostrate, mat-forming herb, rooting at nodes; leaves opposite, very
fleshy but somewhat flattened, linear-oblong to obovate, blunt, sessile or narrowed to a
very short thick petiole; flowers axillary on short pedicels, 4-parted, with one whorl of
perianth parts that are expanded and petaloid, white or colored within, somewhat
appendiculate and slightly cucullate at apices; capsule closely invested by sepals; seeds
compressed, orbicular, black.

Two Pacific varieties are easily distinguished; both have been reported from the
Phoenix Islands.

*Sesuvium portulacastrum L. var. portulacastrum

This was reported from Canton near the dock by Degener in 1959 but was not
seen in 1973 or 1975. It was also collected by Long on McKean. It is red-stemmed, with
shiny green leaves and purplish flowers.

CANTON: Degener & Degemer 24651. McKEAN: Long 2035 (US), 2430
(US).

Sesuvium portulacastrum var. griseum Degener & Fosberg

Stems pale green, not at all reddish; leaves thick, dull grayish green, surface
under a lens, when fresh, covered with distended turgid epidermal cells; flowers white to
pale pink.

This variety is known only from the Central Pacific atolls, including the Phoenix
group, while the other is pan-tropical. This plant forms dense mats along the shores of
lagoons and other low areas where salinity is high and drainage poor.

CANTON: Fosberg & Walker 30207 (US, BISH), Fosberg 30203 (BISH), 30204
(BISH), 30205 (BISH); Degener & Hatheway 21305, 21313, Degener 21451 (BISH,
type); Bryan 24 (BISH). BIRNIE: Fosberg & Stoddart 54725 (US, HAW, K); Long
2635 (US). ENDERBURY: Fosberg & Stoddart 4756 (US, HAW, K); Lamb in 1938
(BISH); Marshall 2 ; Long 2097 (US), 2112, 2653 (US). PHOENIX: Eosberg &
Stoddart 54763 (US, HAW, K); Bryan 19 (BISH); Long 2083 (US), 2088 (US), 2625
(US). HULL: Fosberg & Stoddart 54822 (US, HAW, K). SYDNEY: Stoddart 54865
(US, HAW, K); 55713 (US, BISH,K). GARDNER: Long 2452 (US); Eosberg 55767
(US). McKEAN: Fosberg 55792 (US, BISH, K).

Refs.: Degener and Fosberg (1952), pp. 45-47; Degener and Gillaspy (1955), p.
21; Degener & Degener (1959), p. 9; Maude (1952), p. 70; Pickering (1876), pp. 240,
241, 243.

*Tetragonia tetragonioides (Pallas) O. Ktze. New Zealand Spinach. Mentioned
by Hatheway (1955, p. 5) as planted on Canton in 1951 but not seen in 1973 or 1975.

AMARANTHACEAE

*Alternanthera bettzickiana (Regel) Nicholson

Low, bushy intricately branched herb with reddish stems and reddish or
variegated small ovate to elliptic leaves and small tight axillary clusters of white chaffy
flowers.

Planted sparingly on Canton among buildings as an ornamental foliage border
plant. A recent import.

CANTON: Fosberg 55722 (US, BISH, K).

*Amaranthus dubius Mart. ex Thell. Pigweed
Reported in 1941 by Van Zwaluwenburg as introduced to Canton from Oahu, and

by Degener as growing sparingly in 1951, but as not seen in 1958. We did not find it in
1973 or 1975, but it may well persist around dwellings and installations.

CANTON: Degener & Hatheway 21295.
Refs.: Van Zwaluwenburg (1941, 1943); Degener & Gillaspy (1955), p. 21;

Degener & Degener (1959), p. 9.
*Amaranthus viridis L. Pigweed; Chinese spinach

Small herb with alternate ovate-triangular leaves, greenish spikes of small
flowers, with tiny green deeply rugose fruits, shiny black seeds.

Very rare, established in terminal area on Canton Island.

CANTON: Eosberg 55726 (US).

ANACARDIACEAE

*Schinus terebinthifolius Raddi, reported by Degener as present in 1958 on
Canton, was not seen in 1973 or 1975.

APOCYNACEAE

*Catharanthus roseus (L.) G. Don Madagascar periwinkle

Erect herb, sparingly branched below; leaves opposite, oblong obtuse, sessile;
flowers on very short pedicels or in very condensed cymes in upper axils; sepals 5, small;
corolla salverform, tube somewhat swollen below summit, limb very patent, white to
deep rose or white with a red eye, anthers sessile near top of tube surrounding stigma
which terminates a filiform style; fruits paired linear terete follicles.

A native of Madagascar now at home in all tropical countries, planted but easily
naturalizing itself. Occasional in residence area of U.S.A.F. establishment on Canton.

CANTON: Fosberg & Stoddart 54903 (US).

*Nerium oleander L. was mentioned but noi collected by Luomala, 1951, pp. 166,
173:

*Ochrosia sp. [probably O. elliptica Labill.] was seen cultivated on Canton in
1950, but has apparently not been recorded since, nor was it seen in 1973 or 1975 by us.

*Plumeria rubra L., the Frangipani or Plumeria, grown everywhere in the tropics,
has been tried a number of times in Canton without much success (Luomala 21 (BISH),
Degener 21298 (BISH)). One small, poor- looking plant (Fosberg & Stoddart 54910
(US)) was seen in 1973 and one or two more in 1975 growing in the U.S.A.F. residence
area. When well-grown the species is a small tree wlth very thick branchlets, abundant
latex, and spirally arranged elliptic acuminate leaves, white and veiny beneath; very
fragrant tubular funnelform white, yellow, pink or red flowers with recurved obovate
obtuse coralla lobes.

Refs.: Degener & Gillaspy (1955), p. 27; Degener & Degener (1959), p. 12;
Luomala (1951), pp. 167, 173.

A small * Plumeria obtusa L. was also seen in 1975.

ARACEAE

*Anthurium and *Philodendron mentioned as house-plants in Canton by Degener in
1955, not seen in 1973.

*Cyrtosperma chamissonis (Schott) Merr., the “Babai’” or Giant Taro as reported (as
Alocasia indica p. 156 and C. chamissonis p. 153) by Laxton (1951), pp. 153 and 156,
from Gardner. It has in all likelihood not persisted after the Gilbertese left the atoll.

10

Alocasia macrorrhiza (L.) Schott was seen in a pot at the U.S.A.F. installation on Canton,
1975.

*Dieffenbachia seguine (Jacq.) Schott (Dieffenbachia picta Schott) Dumb Cane

This was seen in 1975 around a house in the U.S.A.F. residence area on Canton.
It will probably not persist here without shelter and irrigation.

*Syngonium angustatum Schott cv. “Albolineatum,” planted close to buildings in U.S. Air
Force settlement on Canton in 1973, one sterile individual climbing on a residence

(Fosberg & Stoddart 54909). This may survive as long as it is watered regularly.

ARALIACEAE

*Polyscias guilfoylei (Bull) Bailey, the Hedge Panax of Hawaii and other tropical
countries was found in Canton in 1953 by Katherine Luomala (7). It was still growing in
1958 according to Degener, collected by Clapp (P-71-15 (US)), and was seen but not
collected by Fosberg and Stoddart in 1973 beside a building in the old British Settlement.
It is a tall bush with a strong odor when broken, alternate pinnately compound leaves,
with few large elliptic leaflets usually irregularly white-margined, the margins toothed. It
seldom flowers and is always propagated by cuttings. A sterile bush of the similar *P.
fruticosa (L.) Harms was found beside a building in the U.S. Air force residence area,

Fosberg & Stoddart 54908 (US); Clapp P-71-16 (US). These are not regarded as part of

the established flora.

Refs.: Degener & Gillaspy (1955), pp. 26-27; Luomala (1951), pp. 168, 173.

ARECACEAE (PALMAE)

*Cocos nucifera L. Coconut

Tree, often very tall, with a columnar trunk ringed by leaf scars, surmounted by a
rosette of gigantic pinnately compound leaves; inflorescences axillary, paniculate, at first
enclosed in a woody sword-shaped spathe or surrounding bract which is caducous as the
panicle expands; staminate flowers distal on panicle branches, pistillate toward bases,
perianth of ivory-colored scale-like segments; fruit a large oval 3-sided fibrous drupe, one
of 3 locules developed and with a single gigantic seed, this with a thin brown testa, a
layer of white endosperm, jelly-like when young, oily when mature, surrounding a large
cavity, when young filled with a very palatable water which later largely disappears.

A most useful tree, planted on all the islands except Birnie and Phoenix (and
McKean?). Those earlier reported on Enderbury are now gone, but seedlings were
planted around U.S.A.F. installation and one naturally started from a drift nut on the

11

beach. Very abundant on Hull and Sydney and Gardner, forming planted forests,
especially around abandoned Gilbertese village sites. Able to tolerate considerable
salinity and some drought, but disappearing on small islands and islets during extreme
dry periods. 600 were said to have been planted on Birnie by Maude in 1952 but have
vanished if indeed they ever grew.

SYDNEY: Bryan 42.

Refs.: Degener & Gillaspy (1955), p. 10; Degener & Degener (1959), p. 7;
Laxton (1951), pp. 138 et seq.; Groves (1951), p. 5.

* Phoenix dactylifera L., the Date Palm, was planted on Canton by Degener in
1950; two small palms survived in 1958 (Degener & Degener (1959), p. 8); none were
seen in 1973 or 1975.

ASCLEPIADACEAE

*Calotropis gigantea (L.) Ait. Crown Flower

Strong erect shrub with abundant milky sap; leaves orbicular to broadly oblong,
opposite, cordate at base, round at apex, arachnoid tomentose beneath; flowers in
pedunculate umbels in upper axils, light dull purple or (in forma wilderi Deg.) white,
calyx 5-parted, corona a handsomely scultured structure surrounding the anther tube
which closely surrounds the large fleshy stigma; fruit a large follicle, borne in pairs.

Introduced on Canton in 1940 or earlier (Van Zwaluwenburg 1942) and by
Degener in 1950-51. Both white and purple color forms were growing on Canton in
1958, according to Degener, and at least one large bush of forma wilderi was seen in the
U.S.A.F. residence area in 1973 and again in 1975. The purple-flowered form had
apparently not survived.

CANTON: Degener & Hatheway 21294 (BISH); Clegern 4 (BISH).

Refs.: Wan Zwaluwenburg (1942), p. 50; Degener & Gillaspy (1955), p. 27;
Degener & Degener (1959), p. 112; Luomala (1951), pp. 166, 173.

ASTERACEAE (COMPOSITAE)

*Conyza canadensis (L.) Cronqg. (Degener & Hatheway 24961), * Gaillardia picta
Sweet, and *Tagetes sp. were reported on Canton by Degener as present in 1958, and
*Emilia sonchifolia (L.) DC., by Van Zwaluwenburg in 1941 but none of them were seen
in 1973. *Borrichia arborescens (L.) DC. had persisted from 1951 to 1958 (Degener &
Degener 24615 (POM)), but was not seen in 1973 or 1975.

12

*Pluchea carolinensis (Jacq.) G. Don (P. odorata (L. ) Cass.)

Resinous aromatic shrub to 2 m tall, with grayish oblong-lanceolate leaves,
corymbiform flat-topped terminal clusters of dull purplish heads, fruit with a crown of
capillary pappus enabling it to be carried readily by wind.

Locally abundant on Canton near the docks and air strip on very-much-disturbed
ground. Considered by Degener to have been of recent introduction in 1951; found by
Fosberg & Walker in 1949.

CANTON: MacDaniels in 1950 (BISH); Fosberg & Stoddart 54888 (US, HAW,

K); Fosberg & Walker 30210 (US); Degener & Hatheway 21295 (BISH, US, NY, A);
Luomala 27 (BISH).

Refs.: Degener & Gillaspy (1955), p. 31; Degener & Hatheway (1952), p. 34;
Degener & Degener (1959), p. 17; Luomala (1951), p. 174.

*Pluchea indica (L.) Less.

Erect shrub to | m tall; leaves green, somewhat dentate, erect, with a resinous
odor, terminal rounded corymbose clusters of purplish pink narrow heads; fruits with a
crown of short pappus bristles.

Very local on Canton, on very disturbed ground near the docks, in 1950-51
introduced from Hawaii, where it is abundant.

CANTON: Fosberg & Stoddart 24889 (US, HAW, K).

Ref.: Degener & Degener (1959), p. 17.

Pluchea x fosbergii Coop. & Gal.

Rather depressed, much branched grayish shrub; leaves oblong, distally toothed;
heads dusty dull purplish in rather large, open, flattopped clusters; fruits lacking fertile
seeds.

A spontaneous, sterile hybrid between the two preceding species, that seems to be
so successful here that locally it is more abundant than either of its parents. It is
interesting in that its rather depressed habit and lanky branching are characters not found
in either parent.

CANTON: Fosberg & Stoddart 54890 (US, HAW, K); Fosberg 55719 (US,
BISH, K); Degener 24637 (BISH).

13
*Tridax procumbens L.

Decumbent leafy herb with erect flowering scapes; leaves angular or somewhat
lobed, dull dark green; heads with whitish rays, yellowish disk; receptacle with prominent
chaffy bracts; fruits with scale-like pappus.

Apparently not present in 1958 but present in 1971 according to photographs by
Roger Clapp, now very abundant in disturbed areas around the U.S.A.F. establishment on
Canton. Found in 1975 on Hull at abandoned U.S.A.F. site. All plants seen were
removed.

CANTON: Fosberg & Stoddart 54887 (US, HAW, K). HULL: Fosberg 55742
(US, BISH, K).

*Vernonia cinerea (L.) Less. Little Iron-weed
Erect, simple to sparsely branched herb with a basal rosette of obovate leaves and
leaves on stem decreasing in size upward, with terminal open clusters of small narrow

purple heads with white bristly pappus.

Abundant in weedy disturbed areas on Hull and Sydney around old Gilbertese
village sites, not known elsewhere in the group.

HULL: Fosberg & Stoddart 54798 (US, HAW, K); Long 2006 (US), 2012 (US),
P-71-43 (US). SYDNEY: Fosberg & Stoddart 54851 (US, HAW, K); Long
2546 (US), 2585, 2562 (US), 2563 (US); Bryan 41 (BISH).

*Zinnia elegans Jacq. Zinnia

An erect herb with opposite leaves and conspicuous heads with bright colored
spreading spatulate ligules.

This was, in 1975, seen growing in a sheltered garden at the U.S.A.F. base on
Canton.

BORAGINACEAE

*Cordia sebestena L. Geiger Tree

Small tree; leaves alternate, ovate, subcordate to cordate at base, obtuse at apex,
very rough above, somewhat stiff, petiolate; flowers in terminal rather compact cymes,
calyx cylindric, striate, toothed, corolla funnelform-salverform, brilliant scarlet, about 2
cm across; stamens included, style twice branched; fruit a large soft fleshy white drupe.

14

Originally from sea-beaches in the Caribbean, this beautiful tree is widely planted
in the tropics. Introduced on Canton before 1949, it persists in the old British Settlement
site and here and there in the U.S.A.F. establishment.

CANTON: Fosberg 30874 (US); Degener 21374; Luomala 16 ; Fosberg 55721
(US, BISH, K); Clapp P-71-24 (US), P-71-20 (US).

Refs.: Degener & Gillaspy (1955), p. 28; Degener & Degener (1959), p. 13;
Luomala (1951), pp. 167, 174.

Cordia subcordata Lam.

Small tree with low sweeping branches; leaves alternate, broadly ovate to broadly
elliptic, tending to be acuminate, base obtuse, upper surface of blade slightly scabrous to
almost smooth, petiolate, petioles often yellow; flowers in loose few-flowered cymes,
calyx cylindric, not ribbed, 4-5 toothed at apex, teeth hirsute within; corolla funnelform,
limb flaring, intense reddish orange, somewhat crispate, stamens in throat, anthers
oblong; style bifid, the branches again bifid just below stigmas; fruit a green drupe with
thin flesh and a narrow neck, formed by the calyx remaining around the corky rugose
stone.

A tree native to most tropical Pacific islands, especially coral islands, found on all
the islands studied except Birnie and Phoenix; rare on Gardner; forming a small forest on
Sydney's lagoon shore.

CANTON: Bryan 21 (GH), 1332 (A); Degener 21374; Luomala 1, 11; Degener
& Hatheway 21287 (A); Fosberg 55724 (US); Clapp P-71-9 (US); Browne in 1939
(BISH, A). ENDERBURY: Fosberg & Stoddart 54751 (US, HAW, K); Bryan 1336;
Lamb s. n.; Marshall 10; Long 2091 (US), 2093 (US), 2100, 2662 (US). HULL: Fosberg
& Stoddart 54812 (US, HAW, K). SYDNEY: Fosberg & Stoddart 54867 (US, HAW,
K); Long 2577 (US), 2590 (US). GARDNER: Fosberg 55765 (US, BISH, K), 55784
(US, BISH, K). "Gardner's and Birney's Islands", U.S. Expl. Exped. (US).

Refs.: Bryan (1942), pp. 50, 60, 71; Degener & Gillaspy (1955), pp. 28-29;
Hatheway (1955), pp. 3-9; Luomala (1951), pp. 167, 174; Laxton (1951), p. 143; Maude
(1952), p. 70; Pickering (1876), pp. 240, 242, 243; Degener & Degener (1959), pp. 13-14.

*Heliotropium procumbens Mill. (H. ovalifolium var. depressum (Cham.) Mert.)

Low herb, stems branching, prostrate to ascending; leaves narrowly oblong or
spatulate to obovate, appressed hairy; flowers in terminal scorpioid cymes, tiny, white,
fruit depressed globose, somewhat 4-lobed, eventually falling into 4 parts.

Recently introduced on Canton probably from Guam or Wake, found only in area
of U.S.A.F. establishment, at least as early as 1971 from a photo by Roger Clapp, but
now common there; introduced from there to the enclosure around the U.S.A.F.
installation on Hull, still present in 1975.

15

CANTON: Eosberg & Stoddart 54897 (US, HAW, K); Fosberg & Clapp = 898
(US, HAW). HULL: Fosberg & Stoddart 54842 (US, HAW); Fosberg 55741 (US,
BISH); Clegern 119 (BISH) .

Tournefortia argentea L. f. (Messerschmidia argentea (L. f.) Johnst.) | Tree Heliotrope

Rounded shrub or small tree, young growth and leaves sericeous-strigose, giving
leaves a grayish frosty green appearance; leaves alternate, elliptic to obovate, obtuse,
thinly fleshy, narrowed to short, thick petioles; flowers in dense, 1-2 times branched
scorpioid cymes, these terminal but very soon becoming axillary by growth of branches
at base of peduncle; flowers white, very fragrant, corolla 5-lobed; fruiting clusters
pendent, fruit a drupe with 4 stones, the flesh of which dries to a firm highly aerogenous
pithy material causing the pea-size fruits to float.

A widespread Indo-Pacific strand plant, one of the first colonists of sand and
gravel bars, not reproducing in shade, but abundant in beach ridge scrub and in other
peripheral vegetation. Known from Canton, Enderbury, Hull, Sydney, also on Gardner.

CANTON: Eosberg 30884 (US); Luomala 13 (BISH); Bryan 22 (BISH); Murphy
in 1949 (BISH). ENDERBURY: Fosberg & Stoddart 54752 (US, HAW, K); Lamb in
1938 (BISH); Marshall 1; Long 2095 (US), 2681. HULL: Fosberg & Stoddart 54816
(US, HAW, K); Schultz in 1939 (US); Clegern 94 (BISH). SYDNEY: Fosberg &
Stoddart 54843 (US, HAW, K). GARDNER: reported by Laxton; Fosberg 55744 (US,
BISH, K).

Refs.: Bryan (1942), pp. 46, 50, 60; Degener & Gillaspy (1955), p. 29; Hatheway

(1955), pp. 4-5, 7-9; Luomala (1951), pp. 164, 165, 174; Degener & Hatheway (1952), p.
34; Laxton (1951), p. 143; Maude (1952), p. 70; Pickering (1876), pp. 240, 242, 243.

CANNACEAE

A hybrid *Canna was growing in a sheltered spot in the U.S.A.F. installation in
1975:

CARICACEAE

*Carica papaya L. Papaya or Pawpaw

A gigantic tree-like herb, with a stem 8-20 cm thick, a crown of great palmately
incised orbicular leaves on long petioles, milky sap; large cream-white pistillate flowers
sessile on the stem, smaller staminate and hermaphrodite flowers in open elongate

16
panicles; large melon-like orange edible fruit with a large central cavity containing many
globose rough black seeds, each in a juicy globose envelope.

The papaya was planted and growing at various times on Canton (Luomala 15),
but did not prosper (Degener & Gillaspy (1955), p. 26; Degener & Degener (1959), p. 12;
Luomala (1951), p. 173). A single plant was seen in the U.S.A.F. establishment in 1973.
Long collected it on Hull in 1964, 2025 (US) and also on Gardner 2474 (US). Clapp also
collected it on Hull, P-71-41 (US).

CARYOPHYLLACEAE

*Spergularia marina (L.) Griseb., introduced on Canton by Degener in 1958,
probably never became established. It was not seen in 1973 or 1975.

CASUARINACEAE

*Casuarina equisetifolia L. Ironwood

Erect tree with very heavy hard wood, dark green, jointed, cylindric
photosynthetic branchlets or “needles, each segment bearing a whorl of tiny scale-like or
tooth-like, vestigial leaves; male and female flowers borne in separate catkins, reduced to
stamens and pistils enclosed in small bracts, the pistils rose-purple; the pistillate
inflorescence accrescent, the bracts enlarging, becoming woody, the whole inflorescence
superficially resembling a diminutive pine cone, the actual fruit small, winged.

Growing sparingly around the U.S.A.F. residence area and at the abandoned
British settlement.

CANTON: Degener & Hatheway 21303.
Refs.: Degener & Gillaspy (1955), p. 20; Degener & Degener (1959), p. 8.

*Casuarina glauca Sieb.

Similar to the above, but more a large shrub than a tree, producing root sprouts,
the needles coarse, stiffer, glaucous green, less prominently striate.

Persisting in a healthy condition around the old British Settlement on Canton.

17

CANTON: Degener 21372; Fosberg 30876 (US); Clapp P-71-11 (US); Fosberg
& Stoddart 54777 (US, HAW, K).

Refs.: Degener & Gillaspy (1955), p. 20; Degener & Degener (1959), p. 8.

COMBRETACEAE

*Conocarpus erectus L. Button-wood or Button Mangrove

Small tree or bushy shrub, leaves alternate, rather small, elliptic, either bright
green or silvery-sericeous; flowers small, with inferior ovaries; crowded in catkin-like
oval heads which are borne in terminal panicles; fruits flat and somewhat winged, borne
packed together in ellipsoidal or oval heads.

Both the green and silvery forms of this species are well-established on Canton,
resulting from Degener's introductions in 1950-51. This, and Coccoloba uvifera, are the
only ones of Degener's introductions that have tended to spread at all beyond the areas of
intensive human activity, past or present. They are both very sparingly established part
way east on both north and south sides of the atoll.

CANTON: Fosberg & Stoddart 54885 (US, HAW, K) (green form), 54886 (US,
HAW, K) (silvery form); Clapp P-71-35 (US) (silvery form); P-71-34 (US) (green form),
P-71-33 (US) (silvery form), P-71-29 (US) (green form).

Refs.: Degener & Gillaspy (1955), p. 26; Degener & Degener (1959), p. 12.

*Terminalia catappa L. Tropical almond
Pp

Tree with conspicuously horizontal whorls of branches, dark brown velvety
pubescence on young tips, and a characteristic style of branching with a branchlet from
below the terminal growing point of a horizontal branch assuming dominance, dipping
downward and arching up level with the turned up apical rosette of the original branch,
then the process repeating itself; leaves spirally crowded into rosettes at the branchlet-
tips, large, obovate, stiff, glossy, turning bright red before falling off; flowers in axillary
slender spikes, cream-white, ovary inferior, perianth campanulate, of one series, 5-lobed,
caducous from the top of the ovary, stamens 10, with subulate filaments, conspicuously
exserted, pistil 1, conspicuously exserted, hooked at summit, soon caducous; fruit about 4
cm long, 2.5 cm wide, a somewhat compressed ovoid drupe with two keels, the flesh
drying rather corky, the seed like an almond, only one or two fruits usually maturing on a
spike.

This widespread tropical cultivated and spontaneous strand and lowland tree was
growing about the old Pan American Hotel in 1949, but was not seen in 1973, or 1975,
although the following species was flourishing.

18

CANTON: Eosberg 30875 (US); Luomala 2; Clapp P-71-30 (US).

Refs.: Degener & Gillaspy (1955), p. 26; Degener & Degener (1959), p. 12;
Luomala (1951), pp. 167, 173.

*Terminalia muelleri Benth. (T. melanocarpa sensu Luomala and Degener & Gillaspy,
non F. v. M.).

Well-shaped tree with a spreading crown, leaves somewhat similar, darker green
than in T. catappa, glaucous beneath, similar branching, similar inflorescence and
flowers; fruiting spikes maturing a number of fruits, these ovoid, somewhat compressed,
about 2 cm long, with dark blue epidermis and dark red juicy flesh, rather sour and
astringent.

Several trees are in very healthy condition, bearing flowers and fruits in the old
British Settlement.

CANTON: Eosberg & Stoddart 54783 (US, HAW, K), 54784 (US, HAW, K);
Luomala 22; Fosberg 30879 (US); Clapp P-71-22 (US), P-71-13 (US).

Ref.: Degener & Gillaspy (1955), p. 26; Luomala (1951), pp. 167, 173.

Terminalia samoensis Rech.

Shrub or small tree, young growth with yellowish close pubescence; leaves
alternate, obovate, short-petioled, thinly coriaceous; flowers on axillary spikes, white,
small, fruits fleshy, dark red when ripe, ovoid, somewhat compressed, 1-1.5 cm long, 1
cm wide.

Known in the Phoenix Islands only from Gardner. The record from Canton
(Luomala (1951), p. 173), based on Fosberg 30879, was a misidentification of T.
muelleri.

GARDNER: Long & Woodward 2475 (US); Long 2494 (US); Fosberg 55762
(US, BISH, K), 55768 (US, BISH, K).

COMMELINACEAE

*Rhoeo spathacea (Sw.) Stearn and *Setcreasia purpurea Boom, mentioned by Degener
& Degener (1959, p. 8) as doing well in gardens on Canton in 1958, were not seen in
1973; however, Rhoeo was still present in 1971, as it was photographed then by Roger
Clapp.

19

CONVOLVULACEAE

*Ipomoea batatas (L.) Poir., Sweet Potato, was observed by Degener in 1958 as
planted and growing, but chlorotic, "in the housing area” (Degener and Degener (1959),
p. 13), but had disappeared by 1973.

Ipomoea macrantha R. & S. (I. tuba (Schlecht.) Don; Calonyction sp.; Ipomoea
turpethum sensu Pickering, non Operculina turpethum (L.) Manso; Ipomoea grandiflora
Lamarck). Moon flower

An extensive and climbing twiner; leaves alternate, large, orbicular cordate,
shortly pointed, petiolate, flowers in few-flowered axillary cymes, with 5 overlapping
orbicular concave sepals; coralloa long tubular-salviform, to 10-12 cm long, limb
spreading, delicate, white, contorted in bud, opening in the evening, collapsing the
following day when sun gets hot; stamens 5, included; style filiform, stigma capitate,
included; calyx strongly accrescent, fleshy, closely investing the capsule, when mature
becoming reflexed, the capsule thin, dehiscent, with four black hairy large seeds with one
angle.

Indigenous on most coral atolls, pan-tropical, elsewhere climbing trees, in the
Phoenix Islands as often spreading over large patches of herbaceous and shrubby
vegetation or on bare coral rubble, dying back to a thick root-crown in very dry periods,
sending out new creeping, twining stems when water is available. The thickened fruiting
calyces are known elsewhere to serve as a source of water for rats, and may well do the
same in the Phoenix Islands. Found on Canton, Enderbury, Hull, Sydney and Gardner.
Mentioned from these islands as J. grandiflora Lam. by Hemsley and in notes with a
specimen, Arundel 7 (K), collected October 1882.

CANTON: Degener & Hatheway 21309, 21310; Degener 24359 (BISH), 24360
(BISH); Luomala 42 (BISH), 43 (BISH); Bryan 25 (BISH); Clegern 41 (BISH); Eosberg
55725 (US, BISH, K). ENDERBURY: Fosberg & Stoddart 54735 (US, HAW, K);
Bryan 13, 31 (BISH), 1333 (BISH); Lamb in 1938 (BISH); Marshall 11; Long 2099
(BISH, US), 2655 (US), 2678 (BISH, US); Clegern 138 (BISH). HULL: Fosberg &
Stoddart 24793 (US, HAW, K); Long 2071 (BISH, US); Clegern 85 (BISH). SYDNEY:

Fosberg & Stoddart 54858 (US, HAW, K); pone 2538 (BISH, US), 2584 (BISH, US).

GARDNER: Woodward & Long 2523 (US); Long 2503 (BISH, US); Fosberg 55774
(US, BISH, K).

Refs.: Degener & Gillaspy (1955), p. 27; Degener & Degener (1959), p. 13
Luomala (1951), pp. 164, 171, 173; Degener & Hatheway (1952), p. 34; Pickering
(1876), pp. 240, 242, 243; Hemsley (1885), p. 116.

Ipomoea pes-caprae ssp. brasiliensis (L.) van Ooststr. (Convolvulus maritimus sensu
Pickering non Gouan, nec. Lam., nec Pall., prob. Desr.). Beach Morning-glory

20

Extensive creeper, rooting at nodes; leaves alternate, ovate-cordate, deeply
bilobed at apex, petiolate, subcoriaceous; flowers in few-flowered ascending axillary
cymes; sepals 5, orbicular, closely imbricate and concave; corolla deeply campanulate,
with short tube and long throat, limb flaring, rich rose purple, darker in center, open at
night, collapsing about midday; stamens 5, included; style 1, filiform, included, stigma
capitate, white; fruit a hard dehiscent capsule; seeds 4, dark brown, hairy.

A pan-tropical beach species, occasionally inland, found in the past very sparingly
on Canton by Hatheway in 1951; a single plant was seen by us at an old fortified spot on
a high beach ridge on the north side and some in front of the U.S.A.F. residence area on
the lagoon beach. On Hull it was found on an open sand flat back of the beach and in an
opening in the coconut plantation in the old Gilbertese village, and seedlings on a dry
channel beach with many other drift seeds, on the north side.

CANTON: Eosberg & Stoddart 54880 (US, HAW, K); Degener in 1951; Bryan
1326 (BISH), 27 (BISH). HULL: Fosberg & Stoddart 54800 (US, HAW, K), 54806

(US); Clegern 67 (BISH).

Refs.: Pickering (1876), p. 243; Hatheway (1955), p. 7; Degener & Gillaspy
(1955), pp. 27-28; Luomala (1951), pp. 166, 173.

*Ipomoea quamoclit L.

Slender twiner with pinnately dissected leaves, and small bright red trumpet-
shaped flowers with strongly exserted stamens and pistil.

Planted in shelter of buildings and seeding itself in the immediate vicinity, on
Canton in the U.S.A.F. residential area. Not likely to persist.

CANTON: Eosberg & Stoddart 55802 (US).

*Merremia tuberosa (L.) Rendle. A vigorous extensive twining vine with
palmately divided leaves, clusters of bright yellow morning-glory-like flowers; the
fruiting calyxes become enlarged and hardened surrounding the globose capsule, forming
a "wood rose."

Seen planted on Canton, climbing on a trellis, but not yet flowering in March
LOTS:

21

CRASSULACEAE

*Kalanchoe pinnata (Lam.) Pers. (Bryophyllum pinnatum (Lam.) Kurz), was said to be
present on Canton in 1958 (Degener & Degener (1959), p. 10) but was not seen in 1973
or 1975.

CRUCIFERAE

*Lepidium bidentatum var. o-waihiense (C. & S.) Fosb. (L. o-waihiense C. & S.) was
introduced to Canton by Degener in 1950-51. It was said to be abundant in 1958
(Degener & Degener 24636), but was not seen at all in 1973 or 1975.

CUCURBITACEAE

*Cucurbita pepo L., the pumpkin, was collected by Bryan on Sydney in 1938
(Bryan 44 (BISH)). It was not seen there in 1973, nor in 1975.

Three other cucurbits were reported on Canton by Degener (Degener & Gillaspy
(1955), p. 30; Degener & Degener (1959), p. 16), but do not seem to have persisted. They
were *Citrullus lanatus var. caffer (Schrad.) Mansf. (C. vulgaris Schrad.), the
watermelon, collected by Luomala (20) (seen again by Fosberg in the U.S.A.F. residential
area in 1975); *Cucumis dipsaceus Ehrenb., the Teasle-gourd, introduced by Degener in
1951 and abundantly naturalized in 1958; and *Cucumis melo L., the Muskmelon,
growing spontaneously in 1950, in a garden in 1958. None of them were seen in 1973.
Cucumis melo was collected on Hull by Long (2003 (US)) in 1964, but we did not see it
in 1973. *Cucumis sativa L., the cucumber, was collected on Canton by Luomala 28
(BISH) in 1950, but was not seen by us in 1973 nor in 1975.

CYPERACEAE

Cyperus javanicus Houtt.

Loosely tufted rough-leafed sedge with grayish-green color, ascending to erect
stout peduncles, umbelloid inflorescences with very long bracts, spikelets falling intact.

Introduced on Canton by Degener in 1950-51, persisting in 1958, but not seen in
1973 or 1975. Common on Sydney, in coconut plantation, not on wet ground.

22

CANTON: Degener & Degener 24650. SYDNEY: Fosberg & Clapp 54850
(US, HAW, K); Long 2557 (US).

Refs.: Degener & Gillaspy (1955), p. 34; Degener & Degener (1959), p. 7.

*Cyperus polystachyos Rottb. was reported from Canton (Degener & Degener
24655) by Degener in 1959, but not seen in 1973 or 1975.

*Cyperus rotundus L. Nut-grass

Plant spreading by underground rhizomes and persisting by small nut-like tubers,
narrow bright green leaves arching close to ground, peduncle erect with clusters of linear
brown spikelets.

Introduced on Canton at least as early as 1941, seen in 1951 and 1959 by Degener
and colleagues, and again by us near the terminal in 1973 and in 1975. It had increased
its area considerably between 1973 and 1975. Seen only sterile.

CANTON: Degener 21413 (US); Fosberg 55717 (US); Clapp P-71-32 (US).

Refs.: Van Zwaluwenburg (1941), p. 19; Degener & Gillaspy (1955), p. 19;
Degener & Degener (1959), p. 7.

Fimbristylis cymosa R. Br. (F. diphylla sensu Degener 1955 and F. dichotoma sensu
Degener 1959, non (L.) Vahl; F. pycnocephala Hbd.)

Dense clumps of short branches and stiff narrowly linear blunt leaves, fibrous
roots with a pleasantly peppery odor, erect peduncles with dense to open clusters of ovoid
to fusiform spikelets, dark brown or black achenes.

Common on Canton, perhaps introduced, very abundant on Sydney, local near
desiccating pond on Hull, very recently introduced in U.S.A.F. Station enclosure on
Enderbury and about the U.S.A.F. station on Hull. Varying enormously in stature, also
varying in inflorescence from dense and button-like (var. pycnocephala (Hbd.) Kiik. ex F.
Br.) to open and spreading, spikelets ovoid to lanceolate.

CANTON: Degener & Hatheway 21290, 21288, 21289 (UC, NSW); Fosberg &
Stoddart 54882 (US, HAW, K), 54901 (US, HAW, K), 54902 (US, HAW, K); Clapp P-

71-28 (US), P-71-3 (US). ENDERBURY: Eosberg & Stoddart 54743 (US, HAW),
54744 (US, HAW). HULL: Fosberg & Stoddart 54826 (US, HAW, K), 54837 (US,
HAW, K); Long 2048, 2059 (US), 2021 (US). GARDNER: Long 2459 (US); Fosberg
55746 (US, BISH, K). SYDNEY: Fosberg & Stoddart 54848 (US, HAW, K), 54849
(US, HAW, K); Bryan s.n.; Long 2543 (US), 2548 (US), Woodward & Long 2575 (US).

Refs.: Degener & Gillaspy(1953), p. 19; Degener & Degener (1959), p. 7.

23

Scirpus subulatus Vahl. Bulrush

This was not found in 1973 or 1975, and is not clear where a habitat for such a
marsh plant could exist on Canton. However, a specimen was supposedly collected in
1964.

CANTON: Holway in 1964 (BISH)

EUPHORBIACEAE

*Acalypha wilkesiana M.-A.; a colored-leafed ornamental, was seen cultivated on Canton
in 1950-51 by Degener, but had disappeared by 1958 and was not seen by us in 1973, or
L975:

Euphorbia L.

A protean genus of a great many species, found almost everywhere except in the
Polar and Sub-Polar regions, differing widely in habit and other vegetative characters, but
united in the reduction of the inflorescences to cup-like fused involucres or cyathia,
which may resemble flowers because of petaloid appendages on glands on their rims.
This genus is divided into several or many by some authors, and there is something to be
said for this course. However, there seems no logical place to stop dividing the aggregate
and, on the whole, it seems more convenient to maintain the genus in the broad Linnaean
sense.

*Euphorbia cyathophora Murr. (Poinsettia cyathophora (Murr.) Kl. & Garcke; E.
heterophylla var. cyathophora (Murr.) Griseb.). Wild or Fiddle-leafed Poinsettia

Erect branched herb with abundant milky sap; alternate leaves, oblong in general
outline but deeply several-lobed on the sides; clusters of cyathia terminal, surrounded by
a whorl of leaves similar to those below but usually with a red blotch on the upper surface
at the base. Fruit a 3-celled capsule; seeds rather cubical, blackish, very rough.

Introduced on Canton, Hull, Gardner, and Sydney, found in weedy places, noted
on Canton by Degener as in the British Settlement, but on this survey seen only around
the buildings of the U.S.A.F. establishment, more abundant in 1975. One plant, only,
seen on Hull in 1973, locally abundant in edges of coconut plantation on Sydney and
around the native village site on Gardner.

CANTON: Eosberg 30885 (US); Degener & Hatheway in 1951; Foale 5 (K);
Clegern 46 (BISH); Luomala 24 (BISH). HULL: Fosberg & Stoddart 54797 (US, HAW,
K); Bryan 49 (BISH); Long 2018 (BISH, US). SYDNEY: Fosberg & Stoddart 54873

24

(US, HAW, K); Long 2536 (BISH, US). GARDNER: Long & Woodward 2466 (US);
Fosberg 55753 (US, BISH, K).

Refs.: Degener & Gillaspy (1955), p. 24; Degener & Degener (1959), p. 10;
Luomala (1951), pp. 168, 169, 173; Degener & Hatheway (1952), p. 34.

*Euphorbia hypericifolia L. (Chamaesyce hypericifolia sensu Degener, Euphorbia
glomerifera (Millsp.) Wheeler

Slender erect or ascending herb, arching at tips, rarely reaching 0.5-1 m tall and
then rather woody at base or a slender shrub, sap milky; leaves oblong, opposite,
minutely toothed; cyathia in small clusters in upper leaf axils, gland-appendages white;
seeds gray.

A widespread tropical weed, in the Phoenix group known only from Canton,
where it is common around U.S.A.F. installations and from Enderbury. We have no
information as to exactly where on Enderbury the Clegern specimen was found in 1973.

CANTON: EFosberg & Walker 30216 (US); Degener & Hatheway 21300 (BISH);
Fosberg & Stoddart 54780 (US); Fosberg 55718 (US, BISH, K): Clapp P-71-23 (US);
Clegern 55 (BISH). ENDERBURY: Clegern 173 (BISH).

Refs.: Degener & Gillaspy (1955), p. 23; Degener & Hatheway (1952), p. 34;
Degener & Degener (1959), p. 10; Luomala (19 51), p. 166.

*Euphorbia hirta L. (Chamaesyce hirta (L.) Millsp.) Hairy Spurge

Small generally pubescent herb, erect to ascending, arching at tips; leaves
opposite, pointed, ovate, serrate, up to 2 cm long, reddish or brownish-green; cyathia in
dense axillary clusters, gland appendages whitish.

One of the most common tropical weeds, having followed man almost
everywhere in the tropics and subtropics except where it is extremely dry. It is
facultatively annual or perennial. In the Phoenix group known from Canton, Enderbury,
Hull and Sydney. On Enderbury it has been established locally for a long time and was
probably introduced by the guano-diggers. One tiny plant of it was found by us in the
enclosure of the U.S.A.F. station, far from any other places where it is found. This
probably represents a second accidental introduction from Canton. On Hull it was found
around both the U.S.A.F. station, and the abandoned Gilbertese village, on Sydney
around the abandoned Gilbertese village, more abundant in 1975 than in 1973 and had
become extremely abundant there and throughout the surrounding coconut plantation.

CANTON: Eosberg 30873; Degener & Hatheway 21298; Fosberg & Stoddart
54786 (US, HAW, K); Luomala 17; Clapp P-7-1-4 (US), P-71-6 (US), P-71-21 (US).
ENDERBURY: Bryan 30 (P); Fosberg & Stoddart 54745 (US), 54757 (US, HAW, K).
HULL: Fosberg & Stoddart 54786 (US, HAW, K); Long 2028 (US). SYDNEY:
Fosberg & Stoddart 54852 (US, HAW, K); Long 2545 (US), 2571.

25

Refs.: Degener & Gillaspy (1955), p. 23; Degener & Degener (1959), p. 10;
Degener & Hatheway (1952), p. 34.

*Euphorbia prostrata Ait. (Chamaesyce prostrata (Ait.) Small). Prostrate Spurge

Very prostrate very slender plant forming tiny purplish green mats, sap milky;
leaves orbicular or nearly so, very small, opposite; cyathia on axillary pedicels or in
several-flowered clusters; capsules 3-angled, the angles hairy.

Common on bare ground around U.S.A.F. installations and old British settlements
on Canton and Hull, and in abandoned Gilbertese villages on Hull and Sydney, not seen
on Sydney in 1975. Luomala’s (1951, p. 169) description of a change in habit from
prostrate to erect suggests that either the plants were infected by a rust, causing the erect
habit, or that the erect plants were really E. hypericifolia misidentified as E. prostrata.

CANTON: Degener & Hatheway 21299 (BISH); Fosberg & Stoddart 54779 (US,
HAW, K); Luomala 29 (BISH); Clapp P-71-5 (US); Clegern 43 (BISH); Fosberg 30880
(US); Fosberg 55715 (US, BISH). HULL: Clegern 130 (BISH); Jenkin in 1865 (K);
Fosberg & Stoddart 54788
(US, HAW, K); Bryan 48 (BISH, K). SYDNEY: Fosberg & Stoddart 54860 (US, HAW,
K).

Refs.: Degener & Gillaspy (1955), p. 23; Degener & Degener (1959), p. 10;
Luomala (1951), pp. 169, 173.
*Euphorbia pulcherrima Willd. (Poinsettia pulcherrima (Willd.) R. Grah.) Poinsettia
This common ornamental was reported by Degener & Hatheway (1952), p. 35, as
present on Canton in 1951, but was not seen in 1973.
*Pedilanthus tithymaloides (L.) Poit. Shoe-flower

Fleshy erect shrub or stiff herb with zig-zag branches; leaves alternate, ovate,
acute, distichous; cyathia red, slipper-shaped, pointed.

Widely planted tropical ornamental, one bush seen in U.S.A.F. establishment.

CANTON: Fosberg & Stoddart 54912 (US, HAW, K). GARDNER: Long 2506
(US).
*Phyllanthus amarus Schum. & Thonn. (P. niruri sensu Degener, non L.).

Erect pale green herb with horizontal branches with tiny pinnately arranged leaves
which hang together and "go to sleep" at night; flowers tiny, greenish, in axils of leaves,

26

male and female in same axil; capsules small, depressed globose; seeds are like sections
of a depressed sphere, strongly striate.

A common pantropic weed, said by Degener to have been introduced to Canton
probably in soil from Fiji, also abundant around the abandoned Gilbertese village on
Hull.

CANTON: Eosberg & Stoddart 54715 (US); Clapp P-71-25 (US); Degener &
Degener 24646 (BISH). HULL: Fosberg & Stoddart 54803 (US, HAW, K); Long 2005
(US); Clegern 79 (BISH). GARDNER: Woodward & Long 2471a (US).

Refs.: Degener & Gillaspy (1955), p. 23; Degener & Degener (1959), p. 10.

GOODENIACEAE

Scaevola taccada (Gaertn.) Roxb. (S. frutescens sensu Degener, non (Mill.) Krause; S.
sericea Vahl)

Tall shrub, rounded and mound-like when growing in open stands or isolated, to
3-4 m tall; leaves obovate, entire, narrowed to a sessile base, rounded at apex, bright
green, exstipulate, with few to abundant white axillary hairs; cymes axillary,
dichotomous, flowers white, gamopetalous, tube split to base dorsally, lobes 5, patent,
arranged in a semicircle like a small fan, margins very thin; fruit a drupe with white flesh,
a ribbed corky stone that floats very well.

Common generally on Canton, Hull and Sydney; one individual bush on
Enderbury at the top of the east beach; occurs as almost impenetrable thickets on Sydney.
Forms with glabrous and pubescent leaves were seen in mixed populations, in different
proportions. Var. fauriei Deg. & Deg. was introduced from Oahu by Degener in 1950-
51, but was not seen in 1973 or 1975.

CANTON: Fosberg 30872 (US); Bryan 1339 (BISH, NY); Degener 24644 (NY);
Degener & Hatheway 21301 (G, BISH), 21302 (G, BISH, NY, A); Fosberg & Stoddart
54877 (US, HAW, K); Luomala 12 (BISH); Long 2409 (US). ENDERBURY: Clegern
134 (BISH); Fosberg & Stoddart 54737 (US, HAW, K). HULL: Fosberg & Stoddart
54810 (US, HAW, K), 54841 (US, HAW, K), 54824 (US, HAW, K); Bryan in 1924
(BISH); Long 2002; Clegern 92 (BISH). SYDNEY: Fosberg & Stoddart 54845 (US,
HAW, K), 54846 (US, HAW, K); Bryan 43 (BISH, A); Long 2586 (US), 2559 (US),
2560 (US). GARDNER: Long 2453 (US, BISH), 2488 (US), 2495 (US); Fosberg 55745
(US, BISH, K), 55778 (US, BISH, K).

Refs.: Bryan (1942), pp. 46, 60; Degener & Gillaspy (1955), pp. 30-31;
Hatheway (1955), pp. 2-9; Luomala (1951), pp. 164, 165, 174; Degener & Hatheway
(1952), p. 34; Maude (1952), p. 70; Laxton (1951), p. 143; Pickering (1876), pp. 240,
241; Degener & Degener (1959), pp. 16-17.

Pa)

GUTTIFERAE (CLUSIACEAE)

*Calophyllum inophyllum L. Portia Tree or Kamani

Large tree with milky sap; leaves opposite, leathery, large, oblong to elliptic,
rounded at ends, petiolate; flowers in axillary panicles, white to slightly pinkish, sepals
small, petals 4, orbicular, stamens many, pistil 1; fruit a pendent globose drupe 2.5-3 cm
in diameter, corky within, seed much smaller.

Introduced on Canton before 1950, and still growing well in 1958 (Degener &
Degener (1959), p. 11), but not located in 1973 nor 1975. Several fairly large trees on
Hull in the old Gilbertese village site; probably planted here, although presumably native
or at least of spontaneous introduction in some other islands of the South Pacific.

CANTON: Luomala 18; Clapp P-71-36 (US). HULL: Fosberg & Stoddart
54794 (US, HAW, K); Long 2033 (US).

Refs.: Degener & Gillaspy (1955), p. 25; Degener & Degener (1959), p. 11;
Luomala (1951), pp. 168, 173.

LAURACEAE

Cassytha filiformis L. Love-vine

Leafless green to orange tangled string-like twining stems, sending haustoria or
penetrating organs into the tissues of other plants (or itself) when it touches them; flowers
very small, white, in short spikes; fruits globose drupes, greenish white turning purple
when very ripe, with a hard globose stone.

Native on most tropical beaches and on most Pacific Islands including two of the
Phoenix group, Canton and Enderbury; parasitic on many plant species including, in the
Phoenix group, at least Scaevola, Portulaca, Boerhavia, Suriana, Sida, and Triumfetta.
On Enderbury droppings, presumably of curlews, were found to be packed with Cassytha
stones, suggesting one of the means by which the plant may be dispersed from island to
island.

CANTON: Bryan 26 (BISH), 1338 (BISH) in 1938; Degener & Hatheway
21282; Luomala 31 (BISH). ENDERBURY: Marshall 4; Long 2106 (US), 2682 (US ).

Refs: Degener & Gillaspy (1955), p. 22; Degener & Degener (1939), p. 9.

28
LECYTHIDACEAE

Barringtonia asiatica (L.) Kurz

Tree reaching a large size, with large alternate obovate leathery entire leaves,
large flowers with many long showy pink and white stamens, the filaments coherent at
base, and pendent large square fruits each containing a single large poisonous seed.

This species has not been known from the Phoenix Islands, but in 1975 a 1 m
seedling was seen in the shelter of a building in the U.S.A.F. area on Canton Island. In
all probability a drifted fruit was picked up on the beach and planted.

LEGUMINOSAE (MIMOSACEAE)

*Desmanthus virgatus (L.) Willd. was introduced to Canton in 1950-51 by Degener. A
few plants survived in 1958 (Degener & Degener 1959, p. 10) but were not seen in 1973
or 1975.

*Intsia bijuga (Colch.) O. Ktze Degener & Degener 24687 (US). Drift seed only.

*Leucaena leucocephala (L.) deWit (L. glauca sensu auct. plur., non (L.) Benth.)

Shrub with alternate twice pinnately compound leaves with many small oblong-
lanceolate leaflets; flowers in globose heads on axillary peduncles, white, with many
stamens; fruit a thin flat, linear-oblong dehiscent pod with transversely arranged flattened
dark brown hard seeds.

This was introduced to Canton with soil from Oahu before 1941 and still persisted
around the old British settlement in 1975.

CANTON: EFosberg 30881 (US); Degener & Hatheway 21296; Luomala 3; Clapp
P-71-14 (US).

Refs.: Wan Zwaluwenburg (1941), p. 19; Degener & Gillaspy (1955), p. 22;
Degener & Degener (1959), p. 10; Degener & Hatheway (1952), p. 35; Luomala (1951),
pp. 169, 172.

*Prosopis pallida (H. H. K.) Willd. (P. chilensis sensu Degener, non (Mol.)
Stuntz) was seen by Degener as a pot plant in 1958 but evidently has not persisted.
Seedlings of *Delonix regia (Boj.) Raf. were seen in a bath house on Canton in 1975.

29
LILIACEAE (sensu latissimo, including AGA VACEAE and AMARYLLIDACEAE)

A sterile Aloe, possibly *A. barbadensis Mill. and a sterile *Yucca appear in
photos of Canton made by Roger Clapp in 1971 and were seen by us, around dwellings in
the U.S.A.F. establishment there in 1973 and in 1975. They were not collected or
identified.

*Cordyline fruticosa (L.) Chev. (C. terminalis (L.) Kunth) Ti

Erect stick-like woody stem with a feather-duster-like rosette of large elliptic or
oblong leaves. Various red-variegated horticultural forms as well as the normal green-
leafed one are commonly planted as ornamentals. Several color forms seen planted about
USAF buildings on Canton in 1975. A small plant of the green form of *Cordyline
fruticosa (L.) Chev. was seen in a bath-house on Canton in 1975.

*Crinum asiaticum L. Crinum lily

Large rosettes of soft lanceolate leaves, often caespitose and reproducing by
offsets, with axillary scapes overtopping leaves with subumbellate clusters of large white
flowers, perianth tubular with linear-oblong recurved lobes, 6 long-exserted stamens with
linear anthers, a single long pistil; fruit fleshy, with very large fleshy seeds.

A common ornamental, persisting around abandoned village sites on Hull and
Sydney atolls; reported from Canton in 1955 and 1959 but not seen there in 1971; unless
Clapp P-71-17 (US) is this species; persisting in the old village site on Gardner in 1975.

HULL: Long 1998. SYDNEY: Bryan 45; Fosberg & Stoddart 54874 (US).
GARDNER: Fosberg 55769 (US, BISH).

LYTHRACEAE

Pemphis acidula Forst.

Erect stiff shrub with many intricate slender branches and a very hard wood;
leaves opposite, small, 1-2 cm long, elliptic, thick, astringent to taste; flowers axillary,
calyx united, campanulate, striate, with 12 teeth, petals 6, white, clawed, stamens 12, in 2
series, styles of different lengths on different plants; fruit a dark reddish brown capsule,
closely invested by calyx to give the appearance of an inferior ovary; seeds many, small.

One of the most characteristic coral island plants, usually growing on coral rock,
rarely, as on Canton, on loose coral sediments; apparently recorded earlier from Canton
(Degener and Gillaspy (1955), p. 23) but the records discounted by Degener as
misidentifications of Suriana, which sometimes looks similar but has yellow flowers.

30

The present Canton plant was a single bush growing along the road on the lagoon beach
and is positively Pemphis. It is abundant on Hull Atoll. Not recorded previously nor
found on this survey on Birnie, Enderbury, Phoenix or Sydney. Reported by Hemsley as
collected on Hull by J.T. Arundel.

CANTON: Eosberg & Stoddart 54717 (US, HAW, K). HULL: Fosberg &
Stoddart 54805 (US, HAW, K); Fosberg 55731 (US, BISH, K), 55738 (US, BISH, K);

Bryan 54; Schultz 43; Long 2054, 2014 (US).
Ref.: Hemsley (1885), p. 116; Pickering (1876), p. 241.

MALVACEAE

*Gossypium brasiliense Macf. and *G. tomentosum Nutt. were introduced on
Canton by Degener in 1950-51 (Degener & Degener (1959), p. 11) but have apparently
not persisted, as they were not seen in 1973. *Hibiscus rosa-sinensis L., reported as
present on Canton in 1951 and 1958 by Degener, likewise was not found in 1973. A
single plant of a pink garden hybrid Hibiscus (Fosberg & Stoddart 54802 (US, HAW, K);
Clapp _P-71-39 (US)) was found flowering abundantly but not fruiting in the old
Gilbertese village on Hull. It is not likely to spread. Another hybrid, red flowered
(Fosberg & Stoddart 54904 (US)), grows near a house in the U.S.A.F. area, Canton.

*Hibiscus tiliaceus L. (Pariti tiliaceum (L.) Britt.) Hau (Hawaii)

A fair-sized tree, often low and with weak, tangled branches, and tough inner
bark; leaves orbicular, cordate, slightly pointed, to 10-15 cm across, dark green above,
white beneath, long petiolate, with large oblong stipules that enclose the terminal buds;
flowers in few-flowered open
terminal racemes, base of each flower subtended by a whorl of linear involucral bracts;
calyx short, toothed, corolla large, showy, campanulate, petals imbricate rolled, united at
base, yellow with maroon base, turning reddish and falling toward evening; stamens
many, filaments united into a fleshy column, their tips, with the anthers, free; ovary
densely hairy, style filiform, 5-branched, stigmas hairy, capitate; fruit a 5-valved
loculicidal hairy capsule; seeds dark brown, small.

A few trees in the U.S.A.F. establishment on Canton, but not seen in the old
British settlement where Degener reported it. It was "recently introduced" according to
Van Zwaluwenburg in 1943. Several large, but slightly chlorotic plants grow in the old
Gilbertese village on Hull.

CANTON: Fosberg & Stoddart 54900 (US, HAW, K); Fosberg 30888 (US);
Degener & Hatheway 21284; Luomala 45; Clapp P-71-37 (US) (leaves green beneath,
possibly var. sterilis F. Br.), P-71-27 (US). HULL: Fosberg & Stoddart 54799 (US).

31

Refs.: Van Zwaluwenburg (1943); Degener & Gillaspy (1955), p. 24; Degener &
Degener (1959), p. 11; Luomala (1951), pp. 167, 173.

*Sida acuta Burm f. (Sida carpinifolia L. f.) introduced on Canton prior to 1951
in soil imported from Fiji, but not reported since 1951, and not seen in 1973 or 1975.

Ref.: Degener & Gillaspy (1955), p. 25.

Sida fallax Walp. Tlima (Hawaii)

Shrub varying in habit from prostrate and mat-forming to erect and 2.5 m tall;
leaves alternate, dull green to gray-green, tomentulose, orbicular, cordate, margin
crenulate, petioles slender; flowers on axillary pedicels, sepals united, with 5 acute lobes,
corolla orange, rarely (in the Phoenix Is.) reddish in center, 1.5-2 cm across, petals united
at base, closing in the evening and falling, stamens united into a column, style with 5
branches; fruit splitting into 5 segments or carpids, these shortbeaked, the beak splitting
into a pore for the ultimate escape of the seed, sides of carpids reticulate rugose. Some
specimens (Long 2670, 2540) have very small leaves.

Generally distributed, one of the most abundant plants on all the Phoenix Islands,
where it dominates the vegetation in many areas, except on Birnie where there was in
1973 only a very small colony of less than a dozen plants. This had apparently
disappeared by 1975. It was mentioned from Canton, Enderbury, Hull and Gardner in
notes with a specimen, Arundel 11 (K) collected Oct. 1882.

CANTON: FEosberg & Walker 30201 (US); Lister in 1889 (K); Degener &
Hatheway 21329, 21330, 21331, 21332 (NSW); Degener & Degener 24661 (NY);
Fosberg & Stoddart 54761 (US, HAW, K), 54914 (US, HAW, K), 54915 (US, HAW, K);
Long 2408 (US); Schultz 22 (US); Luomala 34. BIRNIE: Fosberg & Clapp 54720 (US,
HAW, K), (not found in 1975). ENDERBURY: Fosberg & Stoddart 54733 (US, HAW,
K), 54736 (US, HAW, K); Marshall 9; Long 2098 (US), 2650 (US), 2670 (US); Lamb in
1938 (K). PHOENIX: Fosberg & Stoddart
54761 (US, HAW, K); Fosberg 55801 (US; BISH); Long 2077 (US), 2079, 2080, 2089
(US), 2614 (US), 2634 (US). HULL: Fosberg & Stoddart 54815 (US, HAW, K), 54835
(US, HAW, K); Long 2007, 2067 (US); Schultz 43 (US). SYDNEY: Eosberg &
Stoddart 54847 (US, HAW, K); Long 2540 (US), 2589 (US). GARDNER: Fosberg
55750 (US, BISH, K); U.S. Expl Exped. (NY). McKEAN: Recorded in Long's notes;

Fosberg 55794 (US, BISH, K).

Refs.: Bryan (1942), pp. 50, 53, 54; Laxton (1951), p. 147; Degener & Hatheway
(1952), p. 34; Maude (1952), p. 70; Degener & Gillaspy (1955), p. 25; Degener &
Degener (1959), p. 11; Hatheway (1955), pp. 5-7; Luomala (1951), pp. 162, 163, 171,
173; Pickering (1876), pp. 240, 241, 243; Van Zwaluwenburg (1942), p. 49.

*Thespesia populnea Sol. ex Correa, a pantropical strand tree but not native in the
Phoenix Islands, was introduced to Canton before 1942 (Van Zwaluwenburg (1943), p. 3;
Degener & Gillaspy (1955), p. 25) and was growing around the wharf and the Pan
American Hotel in 1951 (Degener & Hatheway 21308 (NY) and still in 1958 (Degener &

32

Degener (1959), p. 11), also shown on a 1971 photograph by Roger Clapp, but was not
seen in 1973 or 1975. Hemsley reported it as having been collected by J.T. Arundel on
Enderbury before 1884. :

Ref.: Hemsley (1885), p. 116; Luomala (1951), pp. 166, 170, 173; Degener &
Degener (1959), p. 11.

MORACEAE

*Artocarpus altilis (Park.) Fosb. (or A. altilis X mariannensis). The breadfruit was seen
and collected in the Gilbertese village on Hull, presumably planted, by Long [1999 (US)],
but was not seen in 1973. The species was seen as a pot plant on Canton in 1975.

*Ficus microcarpa L.

The Chinese banyan usually starts as an epiphyte, becoming a strangler,
surrounding the trunk of the host tree with roots, these anastomosing to form a very
irregular trunk with the remains of the host tree within; leaves alternate, blade obovate to
broadly elliptic, obtuse to acuminate, thick, veins rather obscure except basal pair;
stipules acuminate; figs small, axillary, globose.

A plant, probably this, seen in photograph taken on Canton in 1971, not seen in
1973 or 1975. Clegern 170 (BISH), labeled as from Enderbury, could scarcely have
come from there, and was probably from a pot plant, or from around buildings on Canton.

MUSACEAE

*Musa nana and, presumably, *Musa sapientum were reported by Degener & Gillaspy
(1955), p. 20 and Degener & Degener (1959), p.8, as introduced and and cultivated on
Canton, but in 1973 and 1975 they were not seen. A species of *Musa was identified on
a photograph of Canton taken by Roger Clapp in 1971 and Clapp P-71-38 (US), leaf only,
is presumably the same; likewise a young plant on a photo taken on Canton by Robert
Clegern in September 1973. In 1975 a small but healthy banana was seen in a sheltered
spot around a house in the USAF residence area on Canton. Such plants will doubtless
grow as long as they are sheltered from salt spray and kept well watered.

33

NYCTAGINACEAE

Boerhavia albiflora Fosb.

Similar to B. tetrandra below but stems green, leaves irregularly oval or ovate,
peduncles from beside the leaf axils rather than in them, bearing umbellate branches, the
flower clusters glomerate, perianth white, campanulate, 5-lobed, lobes emarginate, not
recurved, stamens 1-3, mostly 2, style curved.

A widespread species found on all eight islands studied on this survey, growing
abundantly in different habitats, in places in pure stands, forming large mats or a
continuous ground-cover. The thickened tap roots reach 8 cm or more in diameter and
produce long horizontal branch roots, especially in sandy places. Plants similar to this
occur from Wake and the Northern Marshalls to the Phoenix Islands, with a variety on the
Great Barrier Reef of Australia, where it was recently found on Heron Island. It has
generally been referred to B. diffusa or B. repens, both of which seem to be different.

CANTON: Fosberg & Walker 30207; Degener & Hatheway 21305, (NY);
Fosberg & Stoddart54892 (US, HAW, K); Clapp P-71-10 (US), P-71-2 (US); Browne in
1939 (BISH); Murphy in 1949 (BISH); Lamb in 1938 (BISH); Long 2404 (BISH);
Holway in 1964 (BISH); MacDaniels in 1950 (BISH); Luomala 35 (BISH), 1OA (BISH);
Clegern 30 (BISH); Bryan 1327 (BISH). BIRNIE: Fosberg & Stoddart 54724 (US,
HAW, K); Long 2636. ENDERBURY: Fosberg & Stoddart 54747 (US, HAW, K),
54748 (US, HAW, K), 54759 (US); Bryan 32 (BISH); Lamb in 1938; (BISH); Marshall
13; Long 2094 (BISH), 2096 (BISH), 2102 (BISH), 2119, 2121 (BISH), 2651 (BISH),
2666 (BISH); Clegern_150 (BISH). PHOENIX: Fosberg & Stoddart 54762 (US, HAW,
K), 54767 (US, HAW, K), 54768 (US, HAW), 54770 (US); Bryan 17 (BISH); Long 2077
(HAW) (slightly infected with Albugo, habit condensed, leaves small), 2081 (US, BISH),
2083 (US, BISH), 2087 (US, BISH), 2622 (US). HULL: Fosberg & Stoddart 54823
(US, HAW), 54832 (US, HAW, K); Bryan_52 (BISH, P); Long, 2010, 2061, 2062, 2064,
2065, 2066, 2068, 2069; Clegern 102 (BISH). SYDNEY: Fosberg & Stoddart 54854
(US), 54859 (US, HAW, K), 54864 (US, HAW, K); Long 2541 (US, BISH), 2580 (US),
2588 (US, BISH). GARDNER: Fosberg 55761 (US, BISH, K), 55772 (US, BISH, K).
McKEAN: Long 2030 (US) (very large leafed form), 2046 (US), 2028 (BISH), 2041
(US, BISH), 2034 (US), 2037 (US, BISH), 2428 (US, BISH) (peduncles and their
branches very heavy), 2439 (US, BISH), 2443 (US, BISH); Fosberg 55788 (US, BISH,
K), 55795 (US; BISH, K).

Two collections, Long 2612 (US, BISH) and 2622 (BISH) from Phoenix, and two
from Hull, Long 2068, 2010, have the appearance of this species, but have pink flowers
according to the labels. They do not appear to be B. tetrandra, as neither the leaves nor
the inflorescence resemble that species.

Griffe (1864, p, 207), mentions a plant with thick heart shaped leaves, small
white flowers and thick insipidly sweet root which is cooked and eaten by colonists.
Plant described as probably cruciferous. McKEAN. This may be Boerhavis albiflora.

34

Ref.: Degener & Gillaspy (1955), p. 21; Hatheway (1955), pp. 4-6;
Pickering(1876), pp. 240, 242, 243; Fosberg (1978), pp. 11-12.

Boerhavia tetrandra Forst. f.

Prostrate reddish stems radiating from a thickened vertical tap-root, elongate,
leaves oval or broadly oblong, rounded at both ends, white beneath; inflorescences
axillary, umbelloid, at least twice branched, flowers in open clusters, pink, perianth
scarcely lobed, its margin tending to be turned back, stamens 3-4, mostly 4, pistil 1; fruits
5-ribbed, club-shaped, the ribs glandular viscid.

One patch, only, found on a bare cobbly flat on Hull; reported by Degener from
Canton, but since he specifically says the flowers are white, he probably collected B.
albiflora. Long, (m.s.), reports both pink and white flowered plants from Phoenix, but
his pink-flowered specimens (2612, 2622) are not B. tetrandra. His 2120 (BISH), from
Enderbury may be B. tetrandra but has acute leaves which are not right. It is not
disposed of here. None but white-flowered plants were seen by us on any but Hull and
Gardner Islands. A single small pink flowered plant was found in the village site on
Gardner. Bryan (1942, pp. 54, 57, 238) indirectly reports B. tetrandra from Phoenix and
Birnie, but says nothing that would permit one to think he distinguished 2 species.
Hemsley reported this species as collected on Canton, Hull, and Enderbury by J. T.
Arundel, but we have not seen the specimens.

HULL: Fosberg & Stoddart 54820 (US, HAW, K); Bryan 53 (BISH); Long 2065
(BISH). GARDNER: Eosberg ,55777 (US).

Ref.: Pickering (1876), p. 242? Hemsley (1885), p. 116.

*Bougainvillea glabra Choisy Bougainvillea

A white form of this thorny vine was growing by one of the buildings in the
U.S.A.F. establishment in 1973 (Eosberg & Stoddart 54911 (US, HAW, K)) but will not
likely persist unless it is kept watered during dry periods. The same form was seen in
another sheltered spot in 1975.

*Mirabilis jalapa L., the garden four-o'clock, was planted in the Gilbertese village
on Hull (Long 2032 (US), 2039 (US)), but was not found there in 1973 or 1975.

Pisonia grandis R. Br. (Calpidia ovatifolia sensu Pickering probably an error for
Calpidia ovalifolia Bojer which is totally different.) Buka

Large tree with pale smooth often enormous trunk, soft brittle pulpy wood;
opposite large ovate to oblong, obtuse to somewhat acuminate light green leaves,
deciduous during drought periods, dioecious umbelloid cymes of small greenish flowers,
open panicles of clavate, 5 ribbed, spinose very glutinous fruits.

35

Common on Hull, collected once on Sydney by Bryan and by Long, but not seen
by us in 1973 or 1975; locally abundant on Gardner (1975). This is one of the
characteristic atoll trees, found from eastern Polynesia to the western Indian Ocean, its
fruits admirably adapted to being carried around by sea birds, which use the tree as a
nesting site. A specimen collected by the U.S. Expl. Exped. s.n, (US) has a label that
seems to read "Enderbury's Island." This species has not been seen or collected on
Enderbury in recent years. The Expedition called there in 1840 and 1841.

Canton: Degener & Hatheway 21285 (BRI), 21283 (BISH); Luomala 30 (BISH);
Cranwell Smith in 1956 (BISH). HULL: Fosberg & Stoddart K); Clegern 99 (BISH).
SYDNEY: Bryan 38 (BISH). Long 2015 (US). GARDNER: Long 2507 (US); Fosberg
55764 (US, BISH, K), 55779 (US, BISH, K), 55783 (US, BISH, K). ENDERBURY: [?]
Clegern 154 (BISH), 153 (BISH).

Refs.: Bryan (1942), pp. 60, 64, 71; Laxton (1951), p. 136; Maude (1952), p.
70; Pickering (1876), pp. 240, 242.

PANDANACEAE

*Pandanus tectorius Park. (sensu lato) Screw Pine

Small tree with long linear prickly-margined leaves and subglobose to crylindric-
oval heads of fruits.

In all likelihood not native to the Phoenix Group. Introduced from Hawaii to
Canton in 1950 and said by Degener to have been well-established in gardens in 1958.
Several healthy trees were seen at the abandoned settlement, on the lagoon shore in 1973.
This is a small-fruited form. In the coconut plantations and around the abandoned village
sites on Hull (planted according to Long), Sydney, and Gardner several of the large-
fruited Micronesian forms (P. pulposus Mart.) are common. These are edible forms
doubtless introduced and propagated vegetatively by the Gilbertese settlers. Quite a few
seedlings of them were seen, but none large enough to give any indication of the types of
fruits they would bear. Information from Marshallese informants suggests that seedlings
of the large-fruited forms usually bear small inedible fruits, at least in the Marshall
Islands, where the edible varieties are propagated vegetatively. On Sydney Pandanus
trees were seen sparingly in the old village site, much more abundantly toward the lagoon
from the village.

CANTON: Eosberg & Stoddart 54878 (US, HAW, BISH); Luomala 25. HULL:
Long 2004 (US), 2041; Fosberg & Stoddart 54827 (US, BISH). SYDNEY: Long 2535

(US); Fosberg & Stoddart 54844 (US, HAW, BISH, K). GARDNER: reported by
Pickering and Laxton; Fosberg 55771 (US, BISH, K).

Refs.: Laxton (1951), p. 144; Pickering (1876), p. 240; Degener & Gillaspy
(1955), pp. 16-17; Degener & Degener (1959), p. 4; Degener & Hatheway (1952) p. 35;
Luomala (1951), p. 171.

36

PASSIFLORACEAE

*Passiflora foetida L. was observed growing in cultivation on Canton in 1950 and
more seeds were sown (Degener & Gillaspy 1955, p. 26), establishing the plant locally,
as observed by Degener (1959) in 1958. The species was not seen in 1973 nor in 1975.

POACEAE (GRAMINEAE)

*Cenchrus echinatus L. Sand-burr

An annual or perennial grass with racemes of prickly bur-like fruits about the size
of a small pea.

Abundant on Canton around present and former installations, introduced at least
prior to 1949. Common on Sydney in the plantation, very local on Gardner, rare in Hull,
seen by us there only at the U.S.A.F. installation, but found earlier by Long. Not known
previously from Enderbury where in 1973 it was well established around the U.S.A.F.
installation, doubtless brought accidentally from Canton. We recommended in 1973 to
the Air Force authorities that it be eliminated from the stations on Hull and Enderbury. In
1975 a few plants persisted around the abandoned sites. All seen were pulled up.

CANTON: Fosberg & Walker 30202 (US, BISH), 30217 (US, BISH, K);
Degener & Hatheway 21252; Luomala 9, 38 ENDERBURY: FEosberg & Stoddart
54732 (US, HAW, K). HULL: Long 2005 (US), 2052 (US), 2034 (US); Fosberg &
Stoddart 54840 (US, HAW). SYDNEY: Long 2583 (US); Fosberg & Stoddart 54885
(US, HAW); Fosberg 55730 (US, BISH, K). GARDNER: Eosberg 55752 (US, BISH,
K).

Refs.: Degener & Hatheway (1952), p. 34; Degener & Gillaspy (1955), p. 17;
Laxton (1951), p. 143; Hatheway (1955), p. 1; Degener & Degener (1959), p. 4; Luomala
(1951), pp. 169, 173.

*Chloris inflata Link Finger Grass
Loosely tufted grass with purplish digitate inflorescences, these rather hairy.
Introduced to Canton in 1950, now abundantly naturalized around the U.S.A.F.

installations and housing. Recently established and spreading about the U.S.A.F.
enclosure on Hull.

37
CANTON: Degener & Hatheway 21251; Luomala 40. HULL: Fosberg &
Stoddart 54787 (US, HAW, K).

Refs.: Degener & Gillaspy (1955), p. 17; Degener & Degener (1959), p. 5;
Degener & Hatheway (1952), p. 34; Luomala (1951), p. 172.

*Cynodon dactylon (L.) Pers. Bermuda Grass

Prostrate, tightly rooted, mat-forming fine grass with erect slender peduncles and
digitate inflorescences.

Naturalized at least before 1941, on Canton; still persisting here and there about
U.S.A.F. installations but only on Canton through our visit in 1973, but what was
probably this was seen sterile at the abandoned U.S.A.F. site on Hull in 1975.

CANTON: Degener & Hatheway 21286.

Refs.: Van Zwaluwenburg (1941), p. 19; Degener & Hatheway (1952), p. 34;
Degener & Gillaspy (1955), p. 17; Degener & Degener (1959), p. 5; Luomala (1951), p.
PX

*Digitaria ciliaris (Retz.) Koel. (D. sanguinalis sensu Degener, non (L.) Scop., D.
timorense sensu Degener, non (Kunth) Bal.) Crab Grass

A slender depressed grass with digitate inflorescences. Found on Canton by
Degener, not seen there in 1973 or 1975, but found by us on Hull in the old village site.

CANTON: Hatheway 518 (US); Degener & Hatheway 21315 (US); Degener & Degener
24645 (US), 24648 (US); Clegern 42 (BISH). HULL: Fosberg & Stoddart 54813 (US,
HAW); Clegern 113 (BISH).

Refs.: Degener & Hatheway (1952), p. 34; Degener & Degener (1959), p. 5.

*Digitaria ciliaris ssp. chrysoblephara (Fig. & De Not.) S.T. Blake

HULL: S.W. part of I. 16/8/65, Jenkin 4 (K) det. Veldkamp 1970.

*Digitaria henryi Rendle Henry’s Crab-grass

A prostrate slender slightly glaucous grass with digitate inflorescences on short
pedicels, the racemes not spreading much.

Introduced on Canton by Degener in 1950, noted by him as established in 1958
(Degener & Degener 24647 (US)), not seen on Canton in 1973 or 1975. Possibly seen
near old village in 1975.

38
Refs.: Degener & Gillaspy (1955), p. 34; Degener & Degener (1959), p. 5.

Digitaria pacifica Stapf (D. stenotaphrodes sensu central Pacific authors, non (Nees)
Stapf)

A robust, broad-leafed usually tufted grass, sometimes forming spreading masses;
inflorescences digitate, racemes very stiffly erect. A very distinctive grass, endemic to
the Central Pacific atolls, with its closest relative in the Tuamotu and Society atolls of
southeastern Polynesia. It is to be expected on all of the Phoenix group, but has not been
found on Birnie, Phoenix, or Sydney. Like many Digitaria species, it behaves in a weedy
fashion when opportunity affords, and the continued disturbance on Canton has enabled it
to become much more abundant than it probably was before.

Some specimens of D. pacifica approach D. stenotaphrodes of Southeast
Polynesia in habit and number of racemes, e. g. Bryan 46 (US). However the elliptic
spikelets of this sheet suggest that it is merely a depauperate specimen of D. pacifica. It
is on this type of material that recent records of D. stenotaphrodes from the Central
Pacific are based.

CANTON: Eosberg 30886 (US, BISH); Degener & Hatheway 21316 (US),
21318 (US); Luomala 5, 39, 41; Degener & Degener 24589 (US), 24638; Fosberg &
Stoddart 54899 (US, HAW, K); Clegern 49 (BISH); bet. wharf & airfield, M.A, Foale 3
(K). ENDERBURY: Eosberg & Stoddart 54729 (US, HAW); Long 2107 (US), 2680
(US, K); Marshall 6; Clegern 160 (BISH); A.C. Brown, s n. (US). HULL: Long 2063;

Fosberg & Stoddart 54829 (US, HAW, K); W. Side, Bryan 46 (US, K). GARDNER:
Long 2476 (US), 2292 (UA); Fosberg 55775 (US, BISH, K). McKEAN: Long 2045
(US), 2426 (US), 2031 (US); Fosberg 55990 (US, BISH, K).

Refs.: Degener & Gillaspy (1955), p. 17; Degener & Degener (1959), p. 5;
Luomala (1951), pp. 163, 172; Degener & Hatheway (1952), p. 33.

*Digitaria setigera Roth ex R. & S. (D. pruriens Fisch. ex Trin., D. microbachne sensu
auct. non Presl). Crab Grass

Slender weak-stemmed diffusely spreading grass, flowering and fruiting culms
ascending, racemes digitate or almost so, 4-7, appressed- erect in fruit, second glume
usually less than half the length of spikelet.

A widespread Indo-Pacific species, here recorded for the first time from the

Phoenix Islands, only known from Gardner, where it may possibly have been introduced
by the Gilbertese. It is, however, a native plant in Polynesia.

GARDNER: Fosberg 55782 (US, HAW, K); Long 2504 (US).

39
*FEleusine indica (L.) Gaertn. Goose-grass

A tough wiry grass with ascending stems and 2-4 digitately arranged spike-like
flowering panicles and tiny rough grains.

Occasional in paths, roadsides, and around buildings on Canton and in old village
site on Hull, not seen on Gardner in 1975.

CANTON: Degener & Hatheway 21254; Fosberg & Walker 30211 (US, BISH,
K). HULL: Eosberg & Stoddart 54814 (US, HAW, K); Long 2003 (US, K).
GARDNER: Long & Woodward 2468 (US).

Refs.: Luomala (1951), p. 172; Degener & Hatheway (1952), p. 34; Degener &
Gillaspy (1955), p. 18.

*Eragrostis amabilis (L.) W. & A. (E. tenella (L.) Beauv.) Love-grass

A fine, spreading soft grass, annual or in wet years possibly perennial, stems
tufted, decumbent; inflorescences very finely much-branched, spikelets very small,
compressed.

A pantropic weed of pioneer situations and waste ground, abundant on Canton,
well established on Hull, especially abundant near the U.S.A.F. station; one tuft only seen
on Sydney in 1973 in the coconut plantation. This seems to be the first record from
Sydney; rare on Gardner in 1975 in coconut-Morinda forest, also a first record there.

CANTON: Degener & Hatheway 21297 (US); Clegern 62 (BISH). HULL:
Bryan 47 (US); Long 2024 (US), Fosberg & Stoddart 54836 (US, HAW, K). SYDNEY:
Fosber toddart 54856 (US, HAW, K). GARDNER: Fosberg 55751 (US, BISH, K).

Refs.: Degener & Hatheway (1952), p. 34; Degener & Gillaspy (1955), p. 18;
Degener & Degener (1959), p. 5.

*Eragrostis pectinacea (Michx.) Nees

This was collected near the wharf on Canton in 1951 (Degener & Hatheway
21312 (US, BISH)) and reported by Degener and Hatheway (1952), p. 34 and Degener &
Gillaspy (1955), p. 18, but was not relocated in 1973 or 1975. The specimen in US has
been examined by Leroy Harvey in 1980 and confirmed as E. pectinacea, rather than E.
pilosa (L.) Beauv. which is very similar but with narrower spikelets. There is some
question as to whether these are actually distinct species, but for our purposes we will
accept Harvey’s determination.

40
Eragrostis whitneyi Fosb.

Tiny tufted, many-stemmed annual grass, rarely producing bulbils in the
inflorescence, which is of few very elongate usually curved spikelets. Another of the few
coral atoll endemics, confined to the dry Central Pacific Atolls, except for a variety in the
Leeward Hawaiian Atolls. Grows on sand, especially on low-lying fine compact sand,
with Sesuvium, for example near the Enderbury Lagoon and in drying lagoonlets on the
southwest corner of Hull. Related to the Hawaiian E. paupera and to the Australian E.
dielsii and its allies. Rare on Sydney, as it was not found on either of our visits, 1973 and
1975. A habitat for it exists on Gardner, to the east of the passage into the lagoon, not
visited and seen only from the air on leaving.

CANTON: Bryan 1343 (BISH); Degener & Degener 24614, 24586 (US, BISH);
Degener & Hatheway 21319 (US, BISH); Fosberg & Walker 30206 (US); Hatheway 517
(US); Van Zwaluwenburg in 1940 (BISH). ENDERBURY: Fosberg & Stoddart 54734
(US, HAW, K); Long 2090, 2108 (US, HAW, BISH), 2111 (US, HAW, BISH), 2656,
2661 (US, BISH), 2668 (US, BISH), 2677 (US, BISH), 2683 (US, BISH); Woodward 1,
2, 3, 4 (US), 7; Fosberg 55710 (US, BISH, K); Browne in 1939 (BISH); Clegern 140
(BISH). HULL: Fosberg 55732 (US, BISH). SYDNEY: Long 2591 (US, HAW,
BISH).

Refs.: Pickering (1876), pp. 240, 242?; Fosberg (1939), pp. 39-41; Degener &
Hatheway (1952), pp. 33-34; Degener & Gillaspy (1955), p. 18; Degener & Degener
(1959), pp. 5, 6; Luomala (1951), pp. 171-172.

Lepturus R. Br.

The populations of this genus in the Phoenix Islands are most complex. It is one
of the dominant herbs in many open areas and is common in woods and groves that are
not too dense. In addition to the rather distinctive L. pilgerianus, forms that fit four of the
described varieties of L. repens (Fosberg, 1955) have been found in the Phoenix Group.
This may very well reflect the central Pacific location of the group as well as the fact that
the islands harbor such concentrations of sea-birds. Seeds of this genus are admirably
adapted both for floating and for “hitch-hiking” attached to bird feathers. The main areas
of distribution of the four varieties concerned are arranged radially in several directions
from the Phoenix islands. Such variability is a perfect example of the complexity that
may be found in common strand species.

Lepturus pilgerianus Hansen & Potztal (Lepturus repens var. palmyrensis F. Br.)

A coarse erect bunch-grass, rather stiff, basal branches, if produced, ascending to
erect, not prostrate and rooting at nodes, inflorescence a terete jointed spike with the
flowers and fruits borne in lateral depressions, covered by elliptic acute glumes; rachis
disarticulating at maturity.

After seeing this grass in the field, we now agree that it is separable from L.
repens, though scarcely on the basis of its being annual, as claimed by the authors of the

41

species. All Lepturus species behave as annuals in extremely dry conditions, but can
grow and produce new shoots indefinitely if the moisture supply holds out. L.
pilgerianus is closer to L. gasparricensis, of Wake and Pokak atolls, than to L. repens,
from which latter it differs in its erect or ascending basal branches and broader, elliptic
glumes with scabrous margins at tips, usually not exceeding the joints. It is, actually,
somewhat intermediate between the two, as occasionally an adventitious root or two will
appear on the lowest nodes of the branches. Possibly these and the occasional prostrate
branches of L. pilgerianus may result from hybridity with L. repens, with which it is
occasionally found. Fosberg 55793, from McKean, is somewhat loose and depressed and
has glumes very slightly acuminate. Rare cases of such hybridity are known between L.
repens and L. gasparricensis. There is some question whether or not L. gasparricensis
might better be treated as a variety of L. pilgerianus but further study may settle this.

Possibly L. repens var. maldenensis F. Br. from Malden, belongs here, but the
spikes are more slender and the glumes only about half as long. Some of the Phoenix Is.
specimens approach this but we see no break in the series. L. pilgerianus is not endemic
to Canton, but is found also on Phoenix, Hull, Sydney. On Phoenix it forms extensive
dense stands. It is common in Enderbury and parts of Hull, but rare on Sydney. It is by
no means rare on Canton as stated by Degener & Gillaspy, but abundant in places. It has
not been found on Gardner.

In 1975, after a severe drought, the tall dense stands seen on Phoenix Islands in
1973 were lying flat, dry and appearing dead except that under the influence of some
recent moisture a few green sprouts and branches were appearing from some clumps. In
a slightly lower spot these were more numerous. These may be taken as an indication
that the species is not an obligate annual.

CANTON: Degener & Hatheway 21291 (MO, US, isotype); Fosberg & Stoddart
54718 (US, HAW, K), 54719 (US, HAW), 54881 (US, HAW, K). ENDERBURY:

Fosberg & Stoddart 54740 (US, HAW, K), 54731 (US), 54741 (US, HAW, K); Long
2104 (US), 2109 (US), 2115 (US), 2118 (US), 2654 (US). PHOENIX: Fosberg &
Stoddart 54765 (US, HAW, K), 55800 (US, BISH, K), 54766 (US, HAW, K); Long 2078
(US), 2079 (US), 2079b (US), 2086 (US), 2099 (US), 2615 (US), 2623 (US). HULL:
Fosberg & Stoddart 54830 (US, HAW, K); Long 2017 (US), 2050 (US) (spikes very
slender), 2057 (US). SYDNEY: Fosberg & Stoddart54861 (US, HAW, K); Long 2544
(US). McKEAN: Long 2027 (US), 2031 (US), 2032 (US), 2042 (US), 2047 (US), 2432
(US); Fosberg 55791 (US, BISH, K), 55796 (US, BISH, K), 55793 (US, BISH).

Refs.: Hansen & Potztal (1954), p. 268; Fosberg (1955), pp. 292-293; Degener &
Gillaspy (1955), p. 18; Degener & Degener (1959), p. 6; Fosberg (1968), pp. 496-498;
Pickering (1876), p. 243.

Lepturus repens var. cinereus (Burcham) Fosb. (Lepturus cinereus Burcham).

Lax, trailing slender plant, spikes 1 mm or less thick, glumes about 4 mm long,
acute.

42

Uncommon but rather widely distributed in the western Pacific, found on Sydney
and Gardner, not previously known from the Phoenix Islands.

SYDNEY: Fosberg 55714 (US, BISH, K). GARDNER: Fosberg 55785 (US, K).

Lepturus repens (Forst. f.) R. Br. var. repens

Slender tufts, frequently bearing slender prostrate stolons rooting at nodes, slender
jointed spikes, the glumes lanceolate, acute, not subulate or awned.

This form of the species, that originally described by Forster, is the common form
in Southeastern Polynesia and only occurs sparingly in other parts of the Pacific. It was
only found in 1973 near the old guano pit in the north third of Enderbury Island, also on
Gardner in 1975.

ENDERBURY: Fosberg & Stoddart 54755 (US, HAW, K); Browne in 1939
(BISH). GARDNER: Fosberg 55748 (US, BISH, K), 55776 (US, BISH, K).
Lepturus repens var. septentrionalis Fosb.

More slender than var. swbulatus, forming small hemispherical tufts, leaves a few
cm long, spikes very slender, distinctly less than 1 mm thick.

This form is most abundant in the Northern Marshalls and Wake Island, but
reaches Gardner Island. Common very locally on coral gravel near the west end and on
the north side of Canton Island and on Gardner.

CANTON: Eosberg 55723 (US, BISH, K), 55736 (US, K). GARDNER: Long
2508 (BISH).

Lepturus repens var. repens appr. var. subulatus

Canton: Lister in 1891.

Lepturus repens var. subulatus Fosb.

Slender tufted grass usually producing runners or stolons from the bases of the
tufts, which root and give rise to secondary tufts at the nodes; inflorescence a slender teret
jointed spike with the flowers and fruits sunken in lateral depressions in the rachis,
glumes strongly subulate awned, rachis disarticulating into cylindrical floating sections at
maturity.

Less common than L. pilgerianus except on Sydney, not found on Birnie or
Phoenix. In the Phoenix islands usually forming large soft bunches, sometimes lacking
stolons but easily distinguished from L. pilgerianus by the subulate glumes.

43

ah

CANTON: FEosberg & Walker 30212 (US, BISH, K); Degener & Hatheway
21311 (US); Schultz 23 (US) (sterile); Fosberg 55706 (US, BISH, K). ENDERBURY:
Fosberg & Stoddart 54730 (US, HAW, K), 54738 (US, HAW, K), 54755 (US, HAW, K);
Fosberg 55708 (US), 55707 (US); Long 2092 (US), 2649 (US). HULL: Fosberg &
Stoddart 54808 (US, HAW, K); 54821 (US, HAW, K), 54834 (US, HAW, K); Long 2013
(US), 2056 (US). SYDNEY: Fosberg & Stoddart 54869 (US, HAW, K). GARDNER:
Long 2472 (US), 2489 (US); Fosberg 55749 (US, BISH, K), 55780 (US).

Refs.: Pickering (1876), p. 240?; Fosberg (1955), p. 290; Degener & Gillaspy
(1955), p. 18; Hatheway (1955), pp. 4-6; Degener & Degener (1959), p. 6.

*Panicum distachyon L. (Brachiaria distachya (L.) Stapf)

Prostrate mat-like grass, rooting at nodes, inflorescence a branched raceme of
clavate spiklets.

Established on Canton around the Air terminal and here and there in the U.S.A.F.
housing area. Collected and reported by Degener in 1958.

CANTON: Fosberg & Stoddart 54884 (US, HAW, K); Degener & Degener
24649 (US, BISH), 24656 (US, BISH).

Ref.: Degener & Degener, (1959), p. 6.

*Panicum miliaceum L.

This was not seen in 1973, though reported by Degener & Gillaspy (1955, p. 18)
as locally present on Canton in 1950 (Degener & Hatheway 21314 (US).
*Pennisetum ciliare (L.) Link

Rather low, much-branched annual (?), with long spike-like, often purplish
panicles, the spikelet, surrounded by an involucre of bristles, giving the panicle a foxtail-
like appearance.

Introduced on Canton by Degener in 1950-51 (as P. setosum L. Rich.), this grass
had become locally abundant in 1958. It still persisted as occasional tufts in 1973 and

IOS

CANTON: Fosberg & Stoddart 54771 (US, HAW, K), 54905 (US, HAW, K).

Ref.: Degener and Degener (1959), pp. 6-7.

44

*Setaria verticillata (L.) Beauv., reported from Canton in 1955 (Degener_ &
Hatheway 21253) and 1959, was not seen in 1973. Degener & Gillaspy (1955), pp.18-
19; Hatheway (1955), p. 4; Degener & Degener (1959), p. 7. i

* Tricholaena rosea Nees, introduced on Canton by Degener in 1950-51 and
reported by him in 1959 as widely naturalized in 1958, was not seen in 1973 or 1975.

POLYGONACEAE

*Coccoloba uvifera L. (Coccolobis uvifera L.) Sea-grape

Coarse shrub or small tree with orbicular entire very leathery leaves on extremely
short petioles, with short, sheathing stipules; terminal racemes of small white flowers
with 6 similar perianth segments, 6 stamens and a single pistil; fruiting perianth much
enlarged, becoming fleshy, closely investing the nut-like fruit so as to form an orbicular
dark reddish black drupe, these borne in pendent spike-like racemes.

Common around U.S.A.F. installations on Canton, a few isolated plants near road
as much as 7-8 miles froin station: not seen by us on Hull in 1973 or 1975, one tiny plant
in abandoned Gilbertese village on Gardner.

CANTON: FEosberg 30878 (US). HULL: Long 1998 (US). GARDNER:
Fosberg 55770 (US).

Refs.: Van Zwaluwenburg (1943), p. 3; Degener & Gillaspy (1955), pp. 20-21;
Degener & Degener (1959), pp. 8-9.

PORTULACACEAE

*Portulaca cyanosperma Egler

Prostrate, fleshy; leaves alternate, lanceolate or lance-linear, thickened, with
abundant stipular hair; flowers in few-flowered terminal heads subtended by involucral
leaves; sepals 2, petals obovate, purple; seeds iridescent, bluish.

Introduced on Canton by Degener in 1950-51, reported by him as established in
1958, still persisting sparingly in 1973. Native in the Hawaiian Islands. A specimen said
to be from Enderbury, collected by Clegern in 1973, originally determined as P.
samoensis Vv. Poelln., seems rather to be P. cyanosperma. There may be a confusion of
labels, as there is no other record of its being introduced on Enderbury. On
reexamination, the specimen seems unusually hairy, and may be Portulaca australis (P.
samoensis), which could be native on Enderbury.

45

CANTON: Fosberg & Stoddart 54913 (US, HAW, K). ENDERBURY: 1973,
Clegern 166 (BISH).

Ref: Degener & Degener (1959), p. 9.

The similar but much larger-flowered * Portulaca grandiflora Hook. was seen in a
sheltered garden on Canton in 1975.

Portulaca lutea Sol. ex Forst. f. Giant Purslane

(Here taken in the broadest sense, including large-flowered, thickstemmed mostly
upright plants, probably comprising more than one species).

Tap root rather thick, stems several to many from base, thick, green or often
grayish with a corky epidermis, branched, usually ascending; leaves opposite, usually
appearing to be more or less distichous, obovate, rounded at apex, cuneate at base, fleshy,
with a few stipular hairs; flowers in few-flowered terminal heads subtended by several
involucral leaves; sepals 2, coherent, the resulting calyptra compressed and sharply
keeled, caducous; petals 4-7, obovate 7-15 mm long, emarginate, yellow; stamens many,
15-45, style 1, its stigmatic branches 3-6, spreading, curved, hairy, ovary semi-inferior,
one-celled, with many ovules on basal placentae; fruit a pyxis or circumscissile capsule,
bearing many seeds on a cluster of stalks from the base of the capsule; seeds glossy,
black.

Probably the most abundant plant on all the 8 islands studied, varying in flower
size and other characters from island to island; flowers opening about 10 a.m., closing
about 5 p.m. or later, the petals and stamens turning jelly-like as they shrivel. Degener
has pointed out (1955) that on Canton the stamens and ovaries are eaten by hermit crabs.
We found it impossible on Birnie and Sydney to find any fruits and seeds, though large
numbers of plants were examined.

CANTON: Fosberg & Walker 30208; Degener & Hatheway 21285 (BM, BRI);
Fosberg & Stoddart 54775 (US, HAW, K), 54893 (US, HAW, K); Fosberg 55735 (US,
BISH, K); Clapp P-71-7 (US); Lamb in 1938 (BISH). BIRNIE: Fosberg & Stoddart
54721 (US, HAW, K); Long 2647 (US). ENDERBURY: Fosberg & Stoddart 4726 (US,
HAW, K), 54727 (US, HAW, K), 54728 (US, HAW, K); Lamb in 1938 (BISH); Marshall
3, 8 ; Long 2613, 2667 (US), 2686 (US). PHOENIX: Fosberg & Stoddart 54764 (US,
HAW, K), 54767 (US, HAW, K), 55798 (US, BISH, K), 55799 (US, BISH, K); Bryan
18; Long 2624 (US), 2082 (US). HULL: Fosberg & Stoddart 54828 (US, HAW), 54833
(US, HAW, K); Long 2046, 2043 (US) (very slender). SYDNEY: Fosberg & Stoddart
54870 (US, HAW, K), 54866 (US, HAW, K), 54875 (US, HAW); Long 2547 (US);
Fosberg 55712 (US, BISH, K). GARDNER: reported by Catala (1952, p. 159) as a
staple in the diet of the Gilbertese settlers (as P. oleracea). Long 2455 (US), 2499 (US);
Fosberg 55759 (US), 55757 (US, BISH, K). McKEAN: Long 2038 (US), 2029 (US),
2026 (US); Fosberg 55787 (US, BISH, K), 55797 (US, BISH, K).

46
* Portulaca oleracea L. Common Purslane

Fleshy plant with slender tap root and radiating prostrate branched red stems;
leaves small, opposite, obovate, apex rounded to subtruncate, with few stipular hairs;
flowers in several-flowered terminal heads subtended by several involucral leaves, sepals
2, petals 3-5 mm long, emarginate, yellow, stamens 10-12, style with 3-4 branches,
capsule about 3 mm across, circumscissile about at middle, seeds black, stellulate-
tuberculate. Flowers open about 10 a.m. and close about noon.

Naturalized on Canton and presumably from there introduced around the U.S.A.F.
stations on Enderbury and Hull. Probably introduced by the Gilbertese on Sydney, Hull
and Gardner. It flourishes on bare coral sand and gravel and forms hybrid swarms with P.
lutea when they grow together as on Canton (Fosberg & Stoddart 54772, 54773, 54774,
54906 (all US, HAW, K), Fosberg 55738, 55739 (both at US, BISH, K)), possibly also on
Hull (Long 2043) and Sydney, and almost certainly on Gardner (Fosberg 55758 (US,
BISH, K), 55760 (US)). The hybrids vary in color of stems, stature and flower size. The
swarms have not been analyzed in detail, hence remain in rather doubtful status.

The Canton Island plants, at least, seem to correspond fairly well to var.
granulato-stellulata v. Poelln., with tuberculate seeds, but as yet we have rather vague
ideas as to the definition and limits of the varieties within P. oleracea L. Adequate study
would necessitate several days observation on the ground, noting flower opening and
closing times, number of stamens, stature, stem color, stem thickness, and seed
characters.

CANTON: Eosberg 30881 (US), 30209 (US); Degener & Hatheway 21283;
Fosberg & Stoddart 54817 (US, HAW, K), 54883 (US, HAW, K), 54907 (US, HAW, K);
Fosberg 55737 (US, BISH, K). ENDERBURY: Fosberg & Stoddart 54746 (US, HAW,
K). HULL: Fosberg & Stoddart 54817 (US, HAW, K). SYDNEY: Fosberg & Stoddart
54871 (US, HAW, K), 55711 (US, BISH, K). GARDNER: Long 2501 (US); Fosberg
55754 (US, BISH, K), 55760 (US).

Ref: Degener & Gillaspy (1955), p. 22.

POTAMOGETONACEAE

Ruppia maritima var. pacifica St. John & Fosb.

A slender, submerged, rooted-aquatic with filiform leaves, slender stems and
umbelloid clusters of asymmetric small fruits on long stipes. Found in Enderbury lagoon,
not previously known from the Phoenix group. Tolerates brackish to highly saline water.
Not found though looked for, in 1975, when the lagoon water was so low that most of the
lagoon bottom was dry.

ENDERBURY: in lagoon, Fosberg & Stoddart 54749 (US, K, HAW).

47

RUBIACEAE

*Casasia clusiifolia (Jacq.) Urb., an attractive shrub introduced from the Bahamas
by Degener in 1950-51 (Degener & Degener 1959, p. 15), and apparently doing well in
1958, was not seen in 1973 or 1975.

Guettarda speciosa L.

A handsome tree with dark bark and a dense crown, branchlets rather thick; leaves
opposite, 10-15 or more cm long, broadly oval to broadly oblong, obtuse at apex, cordate
at base, on short petioles, with large flat obovate caducous stipules; flowers very fragrant
at night, in axillary secund symmetrical cymes, calyx small, cup-shaped, corolla large,
white, salverform, lobes 5-8, with membranous crispate margins, these infolded in bud,
opening in evening, falling the next day when exposed to hot sun or, on overcast days,
after noon, anthers attached below sinuses, included; style filiform, of two types, either
subequal with corolla tube and with a broadly cylindric stigma with a drop of very sticky
material on top, or 1/2-2/3 as long as corolla tube and with a more narrowly cylindric
stigma; fruit a depressed-globose white drupe with coarse fibers in the flesh and a large
corky stone with a number of cells each with a single seed.

Common in forests on Hull, Gardner and Sydney, appearing more or less
chlorotic, yellowish green, on the latter island. The stones float very readily and it is
likely that the absence of the species from an island means that it cannot survive there, at
least for a long time. Found once, by Luomala, on Canton. Reported by Hemsley as
collected on Hull by J.T. Arundel.

CANTON: Luomala 19; Clapp P-71-31 (US). HULL: Fosberg & Stoddart
54807 (US, HAW, K), 54809 (US, HAW, K); Fosberg 5733 (US, BISH, K), 55729 (US,
BISH, K); Long 2000 (US), 2001 (US), 2070 (US), 2023 (US). SYDNEY: Fosberg &
Stoddart 54868 (US, HAW, K); Long 2587 (US, BISH); Bryan 35 (BISH). GARDNER:
Fosberg 55743 (US, BISH, K), 55754 (US, BISH, K), 55766 (US, BISH).

Ref.: Bryan (1942), p. 60; Luomala (1951), pp. 167, 174; Pickering (1876), pp.
240, 241; Fosberg (1937), p. 262; Hemsley (1885), p. 116.

Morinda citrifolia L. Noni (Hawaii)

Shrub or small tree, smooth, branchlets rather thick; leaves large, glossy, broadly
elliptic to ovate, petiolate, base obtuse, apex acutish, veins with pits or domatia in axils
beneath; stipules fairly large, caducous; flowers in axillary pedunculate heads, only 1-3 in
bloom at a time, as more flowers open the head tends to elongate, ovaries fused into a
syncarp, calyx lobes very small, corollas salverform, 4-5 lobed, lobes linear-oblong;
syncarp enlarging with maturity into a rather irregular large whitish potato-like
compound drupe which develops a very disagreeable rancid odor when past maturity.

48

Native in the Pacific, but how far east is not known with any certainty. It was in
all likelihood spread far beyond its original range by the aboriginal peoples, who had
many uses for it. Locally abundant in coconut groves on Hull, one of the commonest
trees or shrubs on Sydney and Gardner, forming the dominant understory in some of the
coconut plantations. Found also on Canton.

CANTON: Degener 21412; Bryan 23 (BISH), 1341 (BISH), 1342 (BISH); Clapp
P-71-1 (US). HULL: Eosberg & Stoddart 54838 (US, HAW, K); Long 2055 (US).
SYDNEY: Fosberg & Stoddart 54853 (US, HAW, K); Long 2539 (US), 2570; Bryan 36
(BISH). GARDNER: Long 2458 (US); Fosberg 55747 (US, BISH, K), 55786 (US,
BISH, K).

Refs.: Bryan (1942), p. 60; Degener & Gillaspy (1955), p. 30; Degener &
Degener (1959), p. 15; Luomala (1951), pp. 166, 170 , 174; Laxton (1951), p. 143;
Maude (1952), p. 70; Groves (1951), p. 19.

SOLANACEAE

*Capsicum frutescens L., reported by Degener (1959, p. 14) as growing in a
garden on Canton in 1958, was not seen in 1973. However, by 1975 several varieties
were again in cultivation in very sheltered gardens. Several varieties of *Solanum
lycopersicum L. (Lycopersicum esculentum Mill.), also reported by Degener (Degener &
Hatheway 21307), and not found in 1973 were seen on our visit in 1975, doubtless
recently planted. These were variety commune Bailey, the large-fruited common garden
tomato, and var. galeni (Mill.) Duckwill, the small-fruited cherry tomato, introduced on
Canton in 1950-51, by Degener. *Solanum melongena var. esculenta Nees was reported
by Degener & Degener 1959, p. 15, as growing in a garden on Canton in 1958, not seen
by us in 1973, but was flourishing around a house in the U.S.A.F. residence area in 1975.
*Petunia hybrida Nilm, reported by Degener, had also disappeared, as might have been
anticipated, but was again seen in a pot in 1975. *Nicotiana glauca R. Grah. might more
likely have survived, but was not seen anywhere. It was collected in 1951 (Degener &

Hatheway 21305).

*Physalis angulata L. Ground Cherry or Husk Tomato

A spreading, much-branched herb with thin ovate, usually more or less toothed or
scalloped petiolate leaves, small solitary axillary rotate whitish or pale cream-yellow
flowers with brownish or greenish centers, and a small fleshy tomato-like berry
enveloped in a much enlarged papery loose fruiting calyx.

Very rare in weedy disturbed places on Canton and Hull, on the latter in the
abandoned Gilbertese village. The record from Enderbury may have been a temporarily
established weed, as it was not seen by us in 1973 or 1975. It was found in September
1973 by Clegern.

49

CANTON: Degener & Hatheway 21411 (BISH); Fosberg & Stoddart 54894 (US,
HAW, K); Fosberg 55803 (US, BISH); Degener & Degener 24654 (BISH).
ENDERBURY: Clegern 168 (BISH). HULL: Fosberg & Stoddart 54701 (US, HAW,
K) (this collection possibly P. lanceifolia Nees).

Refs.: Degener & Gillaspy (1955), p. 30; Degener & Degener (1959), p. 15;
Degener & Hatheway (1952), p. 34.

SURIANACEAE

Suriana maritima L.

Densely leafy green shrub up to 3 m tall; leaves alternate, crowded, linear-oblong,
bright green, in loose rosettes on small branchlets, spreading in sunlight, closing,
appressed to each other in rainy weather and at night; flowers axillary, 5-parted, petals
yellow; stamens 5; pistils 5, fruits 5, arranged in a circle, each subglobose, about 1.5 mm
across.

Common, especially near and on lagoon shores but also inland and on seaward
beach-crests, present on Canton and Hull, only, among the islands studied.

CANTON: Degener & Hatheway 21305; Bryan 1328 (BISH), 1340 (BISH);
Luomala 14 (BISH). HULL: EFosberg & Stoddart 54818 (US, HAW, K).

Refs.: Pickering (1876), p. 241; Bryan (1942), p. 46; Degener & Gillaspy (1955),
p. 23; Hatheway(1955), pp. 6, 9; Luomala (1951), pp. 165, 173; Degener & Hatheway
(1952), p. 34.

TAMARICACEAE

*Tamarix aphylla Karst. Tamarisk, Athel

Much-branched tree of a pale grayish green color; leafless, the fine branchlets
terete, fleshy and containing the photosynthetic tissue; flowers in complex terminal
panicles, small, pink, with oblong petals; fruit a small capsule.

An Old-World tree of Mediterranean and other dry or semi-dry regions,
introduced to Canton prior to 1949, it is still growing well about the site of the old British
Settlement. It does not seem to be expanding its range any, in spite of its wind-blown
seeds.

CANTON: Eosberg 39877 (US); Degener & Hatheway 21306; Fosberg &
Stoddart 54781 (US, HAW, K); Luomala 23 (BISH); Clapp P-71-12 (US).

50

Refs.: Degener & Gillaspy (1955), p. 26; Degener & Degener (1959), p. 11;
Luomala (1951), pp. 168, 173.

TILIACEAE

Triumfetta procumbens Forst. f.

Prostrate trailing elongate branched herb, stems and leaves pubescent; leaves
alternate, thick, varying from ovate or orbicular to deeply 3-5 lobed, lobes rounded;
flowers axillary, with 5 sepals, 5 thin yellow oblong petals, many stamens with capillary
separate filaments, and a single pistil; fruit a large dry bur with weak, non-pungent
spinous processes.

A native plant on most Pacific coral atolls, present on beaches and in the interior
of all the Phoenix Islands studied except Birnie. Not found in recent years on Phoenix.
Mentioned from Canton and Enderbury in notes with a specimen, Arundel 6 (K) collected
October 1882.

CANTON: Lister s.n, (CGE); Schultz 19 (US); Fosberg & Walker 30215 (US);
Degener & Hatheway 21281 (US, G), 21282 (G); Luomala 32 (US). ENDERBURY:
Fosberg & Stoddart 54742 (US, HAW); Bryan 1334; Lamb s.n.; Marshall 8; Long 2103
(US), 2105 (US), 2113 (US), 2116 (US), 2652 (US), 2684, 2685. PHOENIX: Bryan in
1924? HULL: Fosberg & Stoddart 54791 (US, HAW, K); Long 2011, 2047 (US), 2051
(US), 2060, 2022 (US). SYDNEY: Eosberg & Stoddart 54872 (US, HAW, K); Long

2569. GARDNER: Long,2487 (US); U.S. Expl. Exp. s. n. (P); Fosberg 55773 (US,
BISH, K).

Refs.: Degener & Gillaspy (1955), p. 24; Hatheway (1955), pp. 1, 7; Luomala
(1951), pp. 171, 173; Pickering (1876), pp. 240, 241, 243; Van Zwaluwenburg (1942), p.
49.

URTICACEAE

Laportea ruderalis (Forst. f.) Chew (Fleurya ruderalis (Forst. f.) Gaud. ex Wedd.)

Succulent stemmed erect branched herb; leaves alternate, petiolate, ovate, strongly
dentate, acuminate, with three strong nerves; flowers usually monoecious, borne in
axillary cymes, individual flowers very small and inconspicuous; fruit a minute flattened
pointed achene, with thickened margin, whitish.

Si

Widespread on coral islands throughout the tropical Pacific, especially on coral
gravel and sand. It is only found on three of the Phoenix Group, Enderbury, Gardner, and
Hull. On the latter it is very abundant. It was mentioned as from these three islands in
notes on a specimen, Arundel 8 (K), collected Oct. 1882.

ENDERBURY: Eosberg & Stoddart 54758 (US, HAW, K); Bryan 29 (BISH);
Long 2117 (US). HULL: Fosberg & Stoddart 54789 (US, HAW, K). GARDNER:
Long 2502 (US); Fosberg 55756 (US, BISH, K). “Fanning: Suwarrow: Enderbury:
Hull: Gardner Islds.”, Arundel s.n. (GH).

Ref.: Pickering (1876), pp. 240, 242.

*Pilea microphylla L. Artillery-plant

This was seen planted in a sheltered spot on Canton in 1975.

VERBENACEAE

*Clerodendrum inerme (L.) Gaertn.

Arching to sprawling glabrous shrub; leaves opposite, elliptic, blunt to acute;
flowers in few-flowered axillary cymes; calyx gamosepalous, corolla white, salverform,
somewhat zygomorphic, stamens and style long-exserted, maroon; fruit an obovoid to
sub-globose drupe with thin black flesh, deeply 4-grooved and eventually drying and
separating into 4 woody nutlets, about 1-1.5 cm high, 1 cm wide.

Probably not native in the Central Pacific east of the Marshall and Gilbert islands,
extending west to the western Indian Ocean, growing as an ornamental in Canton around
old British Settlement and planted in U.S.A.F. area, also in abandoned Gilbertese villages
on Hull and Sydney. The Clegern specimen determined as var. oceanicum Gray.

CANTON: FEosberg & Stoddart 54776 (US, HAW, K); Fosberg 55720 (US,
BISH, K); Clapp P-71-9 (US). HULL: Long 2008 (US); Clegern 111 (BISH).
SYDNEY: Long 2592 (US, BISH).

*Lantana camara var. aculeata (L.) Mold.

Prickly, branched, aromatic shrub; leaves ovate, acute, crenate, bullate-rugose,
petiolate; flowers in axillary, pedunculate, corymbose heads, corollas orange turning red
toward outside of head, bilabiate, lower lip large on peripheral flowers; fruit a small
globose lead-colored drupe, borne in heads.

Introduced as an ornamental, the Lantana has persisted in surprisingly healthy
condition but has not spread as it does in wetter, less saline climates. It was found on

52

Canton and Hull, on the former in the old British Settlement and in the U.S.A.F.
residential area, on Hull in the abandoned Gilbertese settlement.

CANTON: Eosberg & Stoddart 54777 (US, HAW, K); Clapp P-71-8 (US).
HULL: Fosberg & Stoddart 54804 (US).

*Premna serratifolia L.

Aromatic shrub with ovate or oblong leaves and flat-topped paniculate cymes of
small greenish flowers.

Seen only on Canton, planted around the dispensary. Although it is a common
widespread atoll plant, this seems to be the only record of it from the Phoenix Islands. It
was obviously brought in, but is not a plant often seen in cultivation.

A tree which may be Premna serratifolia L.appears on photos taken on Canton in
1971 by Roger Clapp, but the photos are not good enough for certain identification.

CANTON: Fosberg 55737 (US, BISH).

*Stachytarpheta indica (L.) Vahl

Spreading to ascending herb, stems appearing to branch dichotomously with a
spike of flowers or fruits in each forking, leaves opposite, ovate, with obtuse apex,
margins with obtusish teeth; flowers in slender spikes, sunken into grooves in the slender
rachises, corolla bluish, zygomorphic, fruits lying in grooves in rachises; resembling but
more slender than the following species.

Seen only in the dock area, in 1973, not noted previously.

CANTON: Eosberg & Stoddart 54896 (US, type of Stachytarpheta jamaicensis
var. parviflora Mold.; HAW, K).

This collection has been made the basis of Stachytarpheta jamaicensis f.
parviflora Moldenke. It is obviously closely related to that species but requires further
study before it can be finally disposed of. Meanwhile we will leave it under the name by
which we know it as a widespread weed. It has much more slender spikes and smaller
flowers than S. jamaicensis.

*Stachytarpheta jamaicensis (L.) Vahl

Spreading slightly woody herb, the stems appearing repeatedly dichotomous with
a spike of fruits or flowers in each forking; leaves opposite, ovate, with base decurrent
into a petiole, apex obtuse, margin serrate with low obtuse teeth; flowers in terminal
spikes with the calyx and ovary sunken in a groove in the thick rachis, the corolla light

53

purplish blue, curving out from the rachis, strongly zygomorphic and 2-lipped with lower
lip longer; fruit lance-ellipsoid, sunken in the rachis.

Introduced on Canton by Degener in 1951, locally common in weedy places
around the U.S.A.F. installation in 1973.

CANTON: Fosberg & Stoddart 54895 (US, HAW, K).

Ref.: Degener & Degener (1959), p. 14.

*Vitex trifolia L.

A shrub with leaves of 3 leaflets, white beneath, clusters of small bluish-purple 2-
lipped flowers, and pea-like brownish green fruits. A small bush of this was seen in
1975, planted by a house in the USAF residence area on Canton. It is native in the
Western Pacific, but probably will not survive without protection during dry periods on
Canton.

ZYGOPHYLLACEAE

Tribulus cistoides L.

Stems herbaceous, hairy, prostrate to ascending, radiating from a root crown,
sparsely or not branched; leaves opposite, pinnately compound, leaflets small, 1 cm long
or less; flowers on axillary pedicels, 5-parted, petals yellow, 1-2 cm long, obovate,
stamens 5, pistil one, ovary stiffly hairy, style short, stigma slightly lobed; fruit of 5 parts,
united into a button-like thick disk with 10 spines, separating at maturity into 5 segments,
each with 2 strong divergent spines.

Growing in open areas on Canton, Enderbury, Hull, Sydney, and McKean, often
found where seabirds nest. It is very common on Canton, despite the fact that it was not
seen by Degener and Gillaspy (1955, p. 23). It densely covers conspicuous areas on the
lagoon coast of Sydney, and several patches including a large strip from beach top to
lagoon on McKean.

CANTON: Eosberg & Stoddart 54716 (US, HAW, K). ENDERBURY: Eosberg
& Stoddart 54750 (US, HAW, K); Fosberg 55709 (US, BISH, K). HULL: Fosberg &
Stoddart 54790 (US, HAW, K). SYDNEY: Bryans. n., 37; Woodward 2564 (US); Long
2537 (US); Long & Woodward 2574 (US); Fosberg & Stoddart 54857 (US, HAW, K);
Arundel 52 (K). GARDNER: Long 2496 (US); Fosberg 55781 (US, BISH, K).
McKEAN: Long 2044 (US), 2033 (US), 2441 (US); Fosberg 55789 (US, BISH, K).

Refs.: Van Zwaluwenburg (1941), p. 17; Degener & Gillaspy (1955), pp. 22-23;
Luomala (1951), pp. 166, 173.

54

REFERENCES CITED
Bryan, E. H. 1942. American Polynesia and the Hawaiian Chain. Honolulu. 1-253.
Catala, R.L.A. 1952. Rapport sur les iles Gilbert. Noumea. 277 pp.

Degener, O. & Degener, I. 1959. Canton Island, South Pacific (Resurvey of 1958). Atoll
Res. Bull. 64: 1-24.

Degener, O. & Gillaspy, E. 1955. Canton Island, South Pacific. Atoll Res. Bull. 41: 1-
=)

Degener, O. & Hatheway, W. 1952a. Die Flora des Canton Atolls in stillen Ozean.
Revista Sudam. Bot. 10(2): 33-37.

Fosberg, F. R. 1937. Some Rubiaceae of southeastern Polynesia. Occ. Pap. Bishop
Mus. 13: 245-293.

Fosberg, F. R. 1939. Notes on Polynesian grasses. Occ. Pap. Bishop Mus. 15: 37-48.

Fosberg, F. R. 1955. Pacific forms of Lepturus R. Br. (Gramineae). Occ. Pap. Bish.
Mus. 21: 285-294.

Fosberg, F. R. 1968. Systematic notes on Micronesian plants. 3. Phytologia 5: 469-
502.

Fosberg, F. R. 1978. Studies in the genus Boerhavia L. (Nyctaginaceae), 1-5.
Smithsonian Contr. Bot. 39: 1-20.

Graffe, E. 1864. Eine Reise nach der Mac-Keans-Insel. Vierteljahrsschrift der
Naturforschenden Gesellschaft in Zurich, 9: 205-217.

Groves, K. E. [1951]. Report of an expedition to Polynesia...{Loma Linda, Calif.], 24
Pp-

Hansen, I. & Potztal, E. 1954. Beitrage zur Anatomie und Systematik der Leptureae.
Bot. Jahrb. 76: 250-270.

Hatheway, W. H. 1955. The natural vegetation of Canton Island, an equatorial Pacific
atoll. Atoll Res. Bull. 43: 1-9.

Hemsley, W.B. 1885. List of plants from various Pacific Islands, in Challenger Repts.,
Bot. \(IV): 116.

Laxton, P. B. 1951 [1952]. Nikumaroro. Jour. Polyn. Soc. 60: 134-160.

55

Luomala, K. 1951. Plants of Canton Island, Phoenix Islands. Occ. Pap. Bishop Mus.
20: 157-174.

Maude, H. 1952. The colonization of the Phoenix islands. Jour. Polyn. Soc. 61: 62-89.

Pickering, C. 1876. Geographical distribution of animals and plants. Philadelphia. 524
pp. (see pp. 221-248).

Van Zwaluwenburg, R. H. 1941. Canton Island. Hawaiian Pl. Rec. 45: 15-24.

Van Zwaluwenburg, R. H. 1942. Notes on the temporary establishment of insect and
plant species on Canton Island. Hawaiian Pl. Rec. 46: 49-52.

56
Index of Specific Names

Acalypha wilkesiana.:..2s2 beste aE EE, Accent ec oe ee 23
PLO COSI INGICG «x2 csusicn2snetcintesaedsssnad sncetedececbestingedeuesee sees edt eezockans aed ee 9
AlOCASIG NACTOTT NIZA 2.5. ROG ei a 10
Alternanithera Dettzickiana ........:...scesosecesscsvecessssceecredansssoesestl eee ene ee 8
Amaranthus dubius . 2. ccsxsecicdts nk AE awe 8
Amaranthits. Viridis> ..2.5.2...5.5. SRR IR eee Na a eset ccc eee 8
AlratHienriitehit Shs Lesaes22i 2020s deeecds cc bvoe cosa. chsblacdzad cn sdttbuasobsansitetuassecsseses<tceeteS eee es aeeeeeee ae 5
Artocar pus Gliilissn 20) 28 KOR hk. oh 32
BQrringtOnia GSIAtCOL ER... eRe soot se Detect eee nce teat een aed ante LRT eRe 28
Boerhavid alDiflor aces. viascelsceseseoseancssicicctecteendutccennctesinbeeots Gerke reco eeee eee eee 33
BOCFNGVIG TEPENS: < vccasscscsscsccscanscaesvcecteenocedeseseesee ccs toestece sets sce seccunctsed deter eee eee 33
IBOCTRGVIG LELTOMAIG. coisccccocesoccoosecrszoccooseuanesoedesnes epdetenatcgecevenves couest cece eset eee eee 34
Borrichia, ArbOreSCens:; siee.iiscsok lcccsdesss ote hee. ee ee 11
Bougainvillea glabra 2ssiac2 8 ss. Bove ocvcna een cctacees toasts cautsni ne eeaaesersi eee 34
IBY ACHIGTIA GISTACHYG. ..dasisacs seine cesaseds cond anteecesep tense ctensvavaccsh au ete teaten To nee ee 43
Bryophyllum pinnadtum 3.0.22. 2204 26th. oe ee ee eee 21
COLON YGHOM GUD GAS oI cee scsssens Sere se stk eee ee ee 19
Calophyllim inophyllam ccc ca.sscoucieescseclevcecces (oecteee cdot seasons env we 27
Calotropis Biganted .$2.:.20.is.. Ae Re eek sen 11
Calpidia OVALIPONA: ..:5.:..ccccnseseecsessscovsenesceenesstacsetstesvesetssevenesesstevis scvedesessssteteststee ee eam 34
CaM pid O VAUIfOUI Gx. Seah dasasis Tipit aaa ket saa cteees aes ease nes oak cocoa aco 34
COMMAS: ci hoa ccc ass ceside cadena deadad csccsesbucctagenssacssoteessoseseassstseeeueseeasasetaen eae eee aaa 15
COPSiCUIN JTULCSCONS sco ccbccadecescdcccesesecoesccancsesdcacezct casesdse sotaeuevecdi a daneeeect- as 48
GQHIGA PAPGYG iii. cae Oe a ee ee 15
COSGSIG CLUSUPOVG cssiacssskcccexesccteees tess Backes nse eas Ree asec tee ee ee 47
Cassythi filiformis ccsccenctesssesisettioiaasin tetesaig Scene ee PA
Casuarina equisetifolia ......2cds cnn ti. BE ee ee 16
Casuarina plana... esc Betis Rass tacc tek sasndecustacessetue cos seuccossssseses ates vee 16
COINGFARINUS TOSCUS: 5... ssscocvssenesscecsecxsccnse=ceseocseces ttcceoascecto sueceeko cates Sates tee eeeee 9
Genchrus echinatus Seis. Rasheed. oon. Sa ae eee 36
CHAMGESY COMING: bis ek SRR i eee 24
CHAMGCSYCE NYPCTICUOlIG. cscs cvcniscascndcstecesestcescendd ca sdatusoesseseet ceantaotedeeis=sq ce ee 24
GhamaéSyCe PrOStrata «2. c0c2002.68.taceecn pests ie ea ee 25
CRIOr tS inflata. cccss ss cacésndestioesssusisssenussesenauddesgeceoadssoaeesssdescassvscateendecdeceleee eat eee ee 36
Gitrallis LARGIUS: «. c.scscicecedeccccese decd scetasdaskcossataceieucetece testi waseduacneee eee eee 21
Gitrllus vil Saris: xi, .5:..012 kee. kei Reese Ee ee ee PA |
Glerodendrnmithertie, 25.5. 22 .ii secs ssscccnscsscstecsaccesssgedeneseseseUiseoks ate ee 51
COCCOLODG UVIFCTE 5c35-.205:0sosisececssadccnccesecsnsgsecstuecstsrestctestieess Beate ee 44
Coccolobisiuvifera 2:5... 2he.ckieal ace ee ee ee eee 44
Cocosmiuciferdal). Mn. Binds hte belies vcses chest ices oe 10
GCONOCOTPUS CLOCIUS oo sus cszaceséan ce sucaovcnseshentted Geass eocacetee ly
GConyza' canadensis, 's...8.:5:.0 SARs. Se ee ee eee 11
COnGIGIS CDOSTET IG LEE 8. a. cocie sooo ote Sic castes eats scen donseeccenancacstighe tee ee 13,14
GOTAYMINE FFUTICOSA sic. cen2.s0issncssensesnsedsndadnsannncecdesscedcocteseess0scsvcesoee eae ee 29
Gordyline terminalis v2... Stan dR RE eee 29

GU UUTT GSUQUIGUIN oic5 oisnccdecicccccsecseadeoacescecedsesssenseessccksde Sevessee ee 29

OU GUIS ITIDSACCUS Nee teniels Moteier role a ani ea cuue cece tuetee tu aeeta tech do) fe Hou llasal meena apis 21
GUGUINIS HELO Ne TE resins Notas st els takcewsba shte ansausdecsusceuaensbeass coueasdeedies couse: cea uaeLn ret 21
(GT CUTTUS) SQUIV Qt ota sean ete ce sense sic Ve susessst cesses vesunssceteasesausecusbeisasist MUNA Dae 21
GU CUT DILGID OD Osteria crass Pega ties ciatec cca saceasktataate cdl aabaes save toncedecevescasbasaded srveest MONUSLG BE TN 21
Gyn Od OMMGACKY OM perce resets as 28S a oss sovastesus tessa veduests ot vacttvaess sich taseievieceevisetestez REMI 37
Gy PeruUSNGVANICUS HP eer We tee toate cat sac vik ss wee luesesuuts stsssteloct Usdbby ccasdevecaseds 2h OMIDODUL SMMC: 21
Oy Pen POI STACIVOS Pris aie leao:tese,stasecnciucesea seus seu ee Suacsstseeted de, adel seus eM MONS ENG Jue teks 22
Gy POTS HT OTUTIT US NUN, BL is eso o asus xn cave salle sad goah sant aden'zcvacsecas duae sh TeSahSedRte AC oes MeeaSe ee 22
GY TOS ETINGICH GIS ONS fu 20 ss ce: 2ce.vse sschcces <cuese2csessceszesesstass4idetexen dussscneM tenes teseoel DREN RE 9
ID CV OVEN AT CUTIE cc nen eee ecee tate cc resco a tees ncn su ada dy dade eu sasaeudl vac cuessaescxeo et Nees BNA 28
DCS IN AIUAUSIV IG GATES Ime ac tose ts tck bes osc ee i iscis.escervsttxesnssscusuctedtaseceeueteseasocectestxee SE RASS 28
Die fEMVACHIG)PiCla MM mer. c esse ttehc sesccatgustvai seas cukd gah steveseswexesdocescsdesatee heh eases Se sgands 10
Ve ffEMDACHIG SEQUIM Chee ee ver dete es fatto cccaccadensiet di tuasaecnsdiceucsseeqasdtcedaiscsases cand Tet iaesee eh Reo Ate 10
DUS ULAG UG) CULL S pies errene sy SN leezes Let aa esse sscle) cca seeuuh (ost sacs vassaa,cedeaetaeeesvasseaucacacs suet oe Uee one ee i)
DUC UEATUCHIVCTTIaY Utena ss he cc cette save sated tess ice xcectctVestcraacthsass sah shsasssepsteeess SANRIO, Vase es Si
DIG IAIN CKODACICM cane ecssccccrcascxxesssstertsccnsassceckeveceneeascdpocsiaacstteas re MIAO EE 38
DIG TLGIMGIPACIUCG Natasa ares sosecssnstes tus ccaraaeasiaasScrasssnsadessanesandedsavvecteasntaesslt SU OM Doe eas EMO 38
DU GULAT UND IUATLCTIS ates tee asec cee cae eh hcg ae Casco e Sada Saas acdsee candi Casnaessuencs FEN WO 38
DD UCTECTUGIS ATI QULINGLIS pers te coss oral ty he oN Leta csn tl dets Cacte rate san eds la ven sa ueme a ra NAREUED San Suara E 32
DUC TEATAOGISCLEG CT eee eee cde eae vadocleecaeede cuentas iN oon ells kl a AL AA 38
MDI GULCTIORSECNOLAPUTOACS ee sete. Ne ae sce cce nets, Van cecead cosh cn sca tarsus idienascsde ee eeNnte Raweeete 38
DU QUEGT UA CTUINOT CUS Ciena eae a tn eas ose le ste Sn sn ene 4 ARSE YOR EO 37
HEV CUISETICRULCNC Cpe eee ene Nod ose cae Seana clans Seuss Gu sane nN stacate dunes saessta als eet seuns NRRL gun ate 39
ETUDES OFICHILTO LUG ane ee aaa ova na sen ened ea snasccenn cde cauensisae sve sed dgaS dna ua Seat he SASS EDR EREND 11
PT LOT OSTUSHAINGDULIS ects chee one abe de Pesca nna eee ca) a gde ae eiak cceneeueetancnasnaes scccuss eats so AOO NRO: She)
TERRES TORS ERI OLA As ces CER Ree EER Oe EP RE OEE Eat reset et Tay Ee 40
EG AQTOSIUSI DE CHMAGCEE at iNssech rose csnss uedaden-asadnadacedaca ted toashentertsnrasnet esl casace soe Sel ULL Node 39
FEA QTOSTUS DQUD CTA eran aha outsells lode oscalscee sc unewiceseehencs duce venenceleapniitesaciesu cu Saree uta 40
EEO OSTUSIDILOS Ctr eters estan Ne PRN iia a NK ate See tacan soa batanceiooseaRNsa See case eseaes AMEN NSD 39
ET ORTOSUWSIFCTCL UC puters sons se osu chde saws asaciasee silasenedcesesontavont sted cons sunctdeteveccres eee MOS. 39
ET UOTOSUSIWRUEEY lite eccn tee cOhrnasi ee seusti nice tonke sae iacesiaessapeipndsskcasdenevacentdeeres Meee RA On UA 40
TE UDHOWDVGIGY GEO DION ONE ion cosh caentl sc ncucce acest siee eect. -ctdcecceranss once ate dc swore 23
EFUDNOV DIG SIOMCT UCT ise iaaceg ie dete too le rcnesteteecnyhceedorede des uccuoncossceeme tat cSueee NG gee 24
UD ONDIG:MELCHOPIIY Ly oo ean e esse oe cece as etc recn eve tdave? Vic actcccesee sn cuasaseds SSSA au ORES SI ARES 23
EU OT DUG FUTGE CEs soes ese cx face Se ohsh es Sa Soe eRe R sce casd cs rsieoatuncecdeuvonectcpesvo7 RROD SUN ORN 24
EU DHOUDUGUIY CHUGH OL nce cece ces cre cet necsccuke eeve arene csob se ectlonsesseesoercoresi See NO EO 24
FETS OROUAYDT OSU reco cee ce sadn cts 0ou ston atelieds dt ecs con casdoscs incase sv ssyss. SUSAN ate ANN 25
EUPOTDIGyDUl CNCTILING eee. iae ccc fore me veccstecc vate cover vosviadncscusseder drietsescascen OSM ee EN 25
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ATOLL RESEARCH BULLETIN

NO. 394

THE HOA OF HULL ATOLL AND THE PROBLEM OF HOA

BY

DAVID R. STODDART AND F. RAYMOND FOSBERG

ISSUED BY
NATIONAL MUSEUM OF NATURAL HISTORY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C., U.S.A.
FEBRUARY 1994

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YOCYTRIL MARU A AO SSA

THE HOA OF HULL ATOLL AND THE PROBLEM OF HOA

BY

DAVID R. STODDART! AND F. RAYMOND FOSBERG?

ABSTRACT

This paper describes the channels known as hoa which are characteristic of atoll
land rims and of some islands on barrier reefs, where they often dissect previously more
continuous reef-top sediment accumulations and conglomerate platforms. They are
especially common in the central Pacific, and are particularly well developed in some atolls
of the Tuamotu Archipelago and on some of the Society Islands. Published hypotheses
accounting for the origin and development of hoa of different kinds are outlined.
Contemporary hoa on the north side of Hull Atoll in the Phoenix Islands are described and
surveyed, as are older, probably Holocene hoa, here termed ‘paleohoa’, on the same atoll.
Regional and local distributions of hoa and paleohoa and differences in hoa morphology
are used to assess theories of hoa formation. Both hoa and paleohoa are attributed to
hurricane activity, and the presence of paleohoa may thus give an indication of the
distribution of hurricanes in the Holocene.

Department of Geography, University of California at Berkeley, Berkeley, California
94720, U.S.A.

2Department of Botany, National Museum of Natural History, Smithsonian Institution,
Washington, D.C. 20560, U.S.A.

INTRODUCTION

The Polynesian term hoa was first introduced into the reef literature by Danielsson
(1954) and later incorporated into a comprehensive listing of Polynesian topographic terms
by Vallaux (1955, 1991). Danielsson (1954, p. 94) defined hoa as a ‘shallow channel
beginning on the lagoon side [of an atoll rim], separating completely or partially two motu
[reef islands].’ He added the terms tairua for a ‘closed hoa’ and poehoga for the ‘inner part
of a closed hoa.’ As defined by Danielsson hoa are one of the most striking and
conspicuous geomorphic features of most central Pacific atolls, and as such had been
mentioned by many previous investigators. Thus Agassiz (1903, p. 41-43) described and
figured many ‘cuts’, as he termed them, on the north side of Rangiroa Atoll, Tuamotu
Archipelago, as well as on other atolls in that group. But it was Newell’s (1956) account
of Raroia Atoll, also in the Tuamotu, and especially his spectacular aerial photographs,
which brought the phenomenon to general attention. Though in recent years hoa have
attracted increasing interest, especially in French Polynesia, they remain enigmatic features.
It is the purpose of this paper to draw attention to problems of existing interpretations by
reference to the hoa of Hull Atoll, Phoenix Islands, in eastern Kiribati, central Pacific.

CHARACTERISTICS OF HOA

The first detailed descriptions of hoa were made by Agassiz during the Albatross
expedition through the Tuamotu Archipelago in 1899. In a preliminary paper he describes
‘the narrow cuts which divide this part [of the northern rim of Rangiroa, between Tiputa
and Avarua] into a number of smaller islands. These secondary passes leave exposed the
underlying ledge, full of fossil corals. In some cases there is left a clear channel extending
across from the lagoon to the northern [seaward] side through which water flows at high or
half tide. In other cases the cuts are silted up with coral sand blown in from the lagoon
side. In others the cut is shut off by a high sand-bank, or a bank composed of broken
fragments of corals, leaving access to the water from the northern shore only; and finally
the cuts are also shut off on the northern side by sand and broken coral banks, the
extension of the north-shore beach leaving a depression which at first is filled with salt-
water and gradually silted up both from the lagoon side and the sea side, and forms the
typical north-shore land of the lagoon’ (Agassiz 1900, p. 38). More detailed accounts of
hoa on Rangiroa and many other atolls are given in his main report (Agassiz 1903).

At Raroia Newell (1956, p. 330) described ‘approximately 260 shallow channels
across the [atoll] rim which are drained or are awash during low water at the seaward ends
and are filled by 2 to 2.5 meters of water near the lagoon ends’; he did not however use the
term hoa for these channels. ‘In addition to the shallow channels or spillways between
islets, there are some 160 deep, angular clefts or notches (incomplete channels) in the
lagoon shore similar to the channels except that they do not extend across the island to the
seaward side’; these are the tairua of Danielsson, though this term, unlike hoa, has not
entered the reef literature.

Newell saw the incomplete channels as a stage in the formation of hoa: they
‘clearly are being lengthened headward as storm waters cross the islets towards the lagoon,
and they represent various steps in the formation of shallow channels. It is concluded that

the shallow channels are all of very recent origin and were formed chiefly by mechanical
(hydraulic) erosion of the uplifted rim’ (Newell 1956, p. 330).

Stoddart (1969, p. 8) described very similar features at Rangiroa Atoll, also in the
Tuamotus: ‘The islands are separated by shallow, narrow channels known as hoa. These
are Clearly erosional, and represent breaks in formerly more extensive islands. In the walls
of islands transected by hoa, conglomerate rock is exposed underlying the island clastics
and rising above the level of the reef flat. This conglomerate forms a ledge in the walls,
and also floors the hoa itself. Channels cut in the conglomerate by water passing from sea
to lagoon are extending seaward by headward erosion, often terminating in small waterfalls
with plunge pools. ... The lagoonward mouths of hoa are often almost closed by spits
and bars of fresh small coral shingle. On Mahereretiatae a relict hoa was seen in the
process of being recolonised by vegetation, after being sealed at both ends by beach
ridges.’ Stoddart gave schematic diagrams of the features he described.

Likewise at Borabora, Society Islands, Guilcher et al. (1969, p. 8) found examples
of different kinds of shallow passages or hoa on the eastern barrier reef. Some of them are
permanently open; others are closed during periods of moderate surf by spits built by
lagoon waves at their inner ends and become tairua; still others are active only during
exceptional storms, hurricanes or tsunamis. Aprons of sand are prograding into Borabora
lagoon through the hoa and are beginning to fill it. Hoa, tairua and prograding sand aprons
in lagoons are common elsewhere in the Society Islands and in other atolls of the
Tuamotus. Further details are given by Guilcher et al. (1969, p. 28, 46-47), and
comparisons made between hoa and tairua on Borabora and on Tahaa, Maupiti and
Rangiroa.

Newell, Stoddart and Guilcher all viewed hoa as relatively unproblematic features
(see also Guilcher, 1988, p. 148-152); none however surveyed them instrumentally.
CHEVALIER’S TYPOLOGY OF HOA
Chevalier et al (1968, p. 48-49) described hoa in some detail at Mururoa Atoll,
Tuamotus, in similar terms. Chevalier in this paper, in his general survey of French
Polynesian reefs (Chevalier, 1973), and especially in a paper devoted solely to hoa

(Chevalier, 1972), proposed a general classification of hoa:

Type I: open on the lagoon side, closed on the ocean side, permitting throughput of
water only during storms. This he found to be the most common type.

Type II: ‘Functional hoa’, comprising
(a) those open to the sea at high tide;
(b) those open to the sea at low tide during strong wave action;

(c) those fully open to the sea, notably where the algal ridge is unusually
low; he quoted examples from Pukarua and Marutea Sud, Tuamotus.

Type III: hoa where the lagoon end is blocked by sediment deposits and the hoa is
open only at the seaward end. This form is common on atolls with high sedimentation
rates such as Reao. On Mururoa some hoa become enclosed pools open only during
storms.

Type IV: hoa of Type I but completely closed by lagoon sediments. Examples are
given from Marutea Sud and Fangataufa where as a result the channel becomes
hypersaline.

Type V: dry hoa, comprising

(a) hoa of Type IV but completely blocked by sediment, dried out and
vegetated. Chevalier (1972, p. 481) described this type as ‘hoa colmaté’, figuring an
example from Reao.

(b) hoa emersed by a fall in sea-level leaving the floor at 20-40 cm above
lagoon high water, examples being cited from Mururoa, Tureia and Maturei-Vavao.

Type VI: hoa open only to the sea, with the lagoon exit closed by conglomerate, as
at Reao and Pukapuka (Tuamotus). These were interpreted as similar to Type III hoa but
with the lagoon barrier lithifed.

Types II-VI Chevalier grouped as ‘non-functional hoa’, the others as ‘functional’.
At Mururoa he described 79 out of a total of 288 hoa, or 27 per cent, as functional
(Chevalier et al. 1968, p. 51).

Chevalier’s typology systematised and extended previous observations on hoa
morphology. But he also drew attention to problems in their distribution, both between
and within atolls. Thus he noted the paucity of functional hoa on Reao and Pukapuka and
their frequency on Marutea. On Mururoa he found functional hoa concentrated on the
southwest to southeast sectors of the atoll rim, with 34 per cent in the southwest compared
with 2.4 per cent in the east. Chevalier constructed a rose diagram of the frequency
distribution of hoa on the rim in comparison with wind direction and velocity (Chevalier et
al. 1968, p. 12, 16): the graphs showed an inverse correlation between wind strength and
hoa distribution. It was noted, however, that whereas the dominant winds are easterly the
strongest swells are from the south and southwest.

Chevalier noted the probability of a Holocene fall in sea-level of ca. 1.5 m over the
last 3000-4000 years. He also proposed that hoa had their origin in transverse fissures on
the reef rim formed before the fall in sea-level. Finally, he suggested that lithification
processes in reef-top sediments were more effective on the seaward side, and that this was
the reason that hoa began eroding in the less consolidated sediments of the lagoon side,
cutting back along the lines of fissures. He did not discuss the spacing or apparent
clustering of such fissures, nor did he mention the rdle of catastrophic events such as
hurricanes or tsunamis in hoa genesis.

Chevalier concluded his analysis by suggesting an evolutionary sequence in hoa
development, given in outline in Chevalier et al. (1968, p. 53) and in more complex form
in Chevalier (1972, p. 487) (Figure 1). Chevalier’s work focussed attention on the

IV Va

EVSssunres,.2 22 SSSA a rag BES IIa,b ES IIc

Sedimentation

[femrnetes SS Emersion

Figure 1. Evolutionary sequence in the development of hoa proposed by Chevalier
(1972, p. 487).

6

diversity of hoa, but it also made clear that many problems eae in explaining their
morphology, local and regional distribution, and history.

LATER WORK IN FRENCH POLYNESIA

Subsequent work in French Polynesia has provided additional information on hoa,
especially in the context of Holocene sea-level history.

The 150 hoa of Rangiroa Atoll are classified by Ricard et al. (1985) into
permanently functioning hoa; partly obstructed hoa, including those ‘partly isolated from
the ocean by coral boulders but open to lagoon influences’ and those ‘largely open to
oceanic influences, but more or less isolated from the lagoon by a sandy strip’; and entirely
obstructed hoa. At Takapoto Atoll Salvat and Ricard (1985) use the terms ‘opened hoa’
and ‘unworking hoa’. At Borabora (Pirazzoli et al. 1985) there are both functional and
non-functional hoa. The latter are subdivided into filled-in hoa, invaded by storm debris;
emerged hoa, resulting from relative sea-level change; and ‘obturated hoa’, closed on the
seaward side by coral conglomerate and sometimes on the lagoon side by spits of
sediment.

In spite of terminological differences these classifications can all be clearly related
to Chevalier’s typology of hoa.

The Tuamotu atolls show great variability in frequency of hoa. Taiaro and Anaa
have rather few (Salvat et al. 1977, Pirazzoli et al. 1988), and with one exception those at
Taiaro are non-functional. Conversely hoa are frequent at Temoe and Reao (Pirazzoli
1987, Pirazzoli et al. 1987). In both cases they are highly concentrated on the west and
southwest sides of the atolls, with few or none on the north and east sides. On the other
hand at Tikehau the hoa are open in the east but closed in the northwest (Harmelin-Vivien
et al. 1985).

Various factors have been suggested to account for the features and origins of the
Tuamotu and Society Islands hoa. In particular Pirazzoli and Montaggioni (1988) have
summarised a great deal of evidence from corals, conglomerate platforms, beachrock and
intertidal notches that mean sea-level stood 0.8-1.0 m above its present level in the period
5000-1250 B.P., and that it did not fall below 0.7 m above present between 4500 and 1250
B.P. Emerged reef flat material in the floors of hoa has been described from Taiaro (+0.6
m, with dates of 1140480 and 1110+80: Salvat et al. 1977) and Reao (+0.5 m, 4250+100:
Pirazzoli et al. 1987). Pirazzoli (1987) has also described ‘obturated hoa’ blocked on the
seaward side by continuous conglomerate platform with a date of 3170+60 B.P., attributed
to a Holocene high sea-level stand, at Temoe. Other relevant dates are quoted by Pirazzoli
and Montaggioni (1988) for Rangiroa, Takapoto, Pukarua, Mururoa, Borabora, Raiatea-
Tahaa, and other localities with hoa.

Although Chevalier did not consider high-magnitude events such as hurricanes as
CauSative agents in the formation of hoa, they have been specifically considered (together
with other high-energy events such as tsunamis) by Bourrouilh-Le Jan and Talandier
(1985) in the explanation of megablocks on the reef flat and the alternation of motu and hoa
at Rangiroa. The megablocks they consider to be fragments of reef flat previously

7

delineated by fractures and dislodged and transported during such events. Hoa they
suggest are activated in sectors of reef rim not subject to previous fracturing. They discuss
particularly the major hurricanes of January 1903, March 1905 and February 1906, and the
six El Nifio-related storms between December 1982 and April 1983. Both sets of storms
affected Rangiroa and the northwestern Tuamotu atolls (Tikehau, Mataiva). The 1983
storms were particularly badly felt at Mataiva, where some defunct hoa were reactivated.
These they call storm hoa (‘hoa d’ouragan’) (Bourrouilh-Le Jan and Talandier 1985, p.
316). While they give no detailed analysis of the hoa on any particular atoll, they suggest
that their distribution and frequency results from the interaction of hurricane waves, their
magnitude and angle of incidence, and reef topography and orientation.

Other workers have suggested that sediment transportation during hurricanes has
been responsible for the transformation of functional into non-functional hoa. Thus at
Taiaro, where the hoa are non-functional, Salvat et al. (1977) draw attention to the
formation of storm embankments more than 2 meters high between 1878 and 1906, and
Pirazzoli et al. (1987) suggest that the non-functional hoa on the northern rim of Reao were
closed by the hurricane of 1903.

Chevalier’s suggestion about the importance of diagenesis of reef-top sediments in
controlling motu accretion and hoa erosion has been discussed in general terms by
Bourrouilh-Le Jan et al. (1985), with specific reference to Rangiroa, Tikehau and Mataiva
Atolls.

Lenhardt (1991) has studied the hydrodynamics of hoa at Tikehau, but in the
context of their function as normal flow conduits rather than in terms of their formation
during extreme events.

QUESTIONS ABOUT HOA

This review readily generates a number of questions about hoa, some of which are
not often addressed in the literature. These include the following:

° why are hoa apparently regionally concentrated, especially in the central Pacific?
[this is part of the larger question: why do some atolls have a considerable part of their reef
rim occupied by land and others do not?].

° why are hoa locally concentrated in particular parts of atolls?

° why do the reef sectors on which hoa are most numerous vary between atolls?

¢ why do some atolls (for example Diego Garcia, Canton, Gardner) apparently
have no hoa?

¢ why do some atolls (such as Raroia, Temoe, Reao, Rangiroa) have abundant
hoa?

° why do some atolls have areas with abundant hoa and others apparently devoid
of hoa?

¢ why on the same atoll can one find both active hoa (both functional and non-
functional) and fossil hoa (paleohoa), often in close juxtaposition?

Answers to these and similar questions must include consideration of:

* erosive processes resulting from wave activity generated by trade winds, swell of
more distant origins, and episodic storms.

° current sedimentary processes on both seaward and lagoon shores of motu.
y/

* possible fall in sea-level from a Holocene high stand over the past few thousand
years (a regional effect).

* possible tectonic deformation consequent on lithospheric flexure.
* the restricted spatial extent of even the most extreme catastrophic storm events.

These issues form the background to our consideration of the hoa of Hull (Orona) Atoll,
Phoenix Islands, Central Pacific.

HULL HOA

Hull (or Orona) Atoll, in 172°11'W, 4°31'S, is the central atoll of the southern tier
of the Phoenix Islands in the Central Equatorial Pacific. It was discovered by the U.S.
Exploring Expedition in August 1840 (Wilkes, 1845, III, p. 369-370). Between 1938 and
1963 it was occupied by settlers from the Gilbert Islands, but has since been uninhabited
except for a small U.S. military presence in the early 1970s. The atoll is located 185 km
south of Canton Atoll and 98 km due west of Sydney Island; Gardner Atoll lies 245 km
further west (Figure 2). It is 10.8 km long, 4.2-5 km wide, and has a total area of 40.5 sq
km (Figure 3); of this islands occupy 6.7 sq km, peripheral reef and channels 7.8 sq km,
and lagoon 26.0 sq km. There is no pass into the lagoon, and no bathymetric survey has
been made of it; it is said to have depths of 15-18 m (Bryan, 1942, p. 63). The seaward
reef flats are 80-200 m wide, being narrowest on the east and west sides and widest on the
north and south.

The atoll has a mean annual rainfall (1952-1963) of 1171 mm (maximum 2599.2
mm, minimum 245.6 mm). Spring tidal range is approximately 80 cm.

The land rim varies in width from less than 50 m at the eastern point, where it
consists of gravel ridges, to a maximum of 650 m on the west side; the average is 250-300
m. The total shoreline periphery on the seaward side, including hoa, is approximately 26.5
km. Of this 5 km (19 per cent ) on the northwest side comprises islands with hoa. The
definition of the hoa is somewhat problematic but there are some 20 intersecting the
conglomerate platform as well as lesser channels between islands on the platform. In
addition there are four hoa in the otherwise continuous southern rim, and none on the
northeast and southwest sides. There is, however, a sector of ca 1500 m near the south
point of former hoa (paleohoa), to be described.

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Profile 4 (Figure 4) was surveyed at the easternmost point of the atoll, near the
former settlement, using a Kern automatic level. The land rm at this point is ca 600 m
wide, and the profile extends inland for 280 m from the seaward shore. Extensive
cemented conglomerate forms steep bastions rising to 3.4 m at the seaward beach, which is
backed by a broad sand and rubble seaward ridge rising to over 3.0 m.

Aerial observation shows that much of the continuous land rim consists of parallel,
presumably storm-deposited, shingle and rubble ridges (Figure 6). Lagoon beaches are
low and sandy.

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The hoa on the north side of Hull are channels intersecting the broad conglomerate
platform underlying the reef-rim motu. They are normally widest toward the lagoon end
and narrow and sometimes bifurcate seawards (Figures 7, 10, 13 and 14). They rarely
penetrate through the seaward reef rim, but terminate in a pronounced cliff of conglomerate
over which water pours at high tide and during storms. The conglomerate platform usually
outcrops along both sides of each hoa (Figures 11 and 12). The lagoon mouths are often
partially closed by sandspits (Figures 7 and 8), and the lagoon ends of larger hoa are
marked by lobate deltas and aprons of sand transported through the hoa and building out
into the lagoon (Figures 9 and 10). The sectors of conglomerate platform defined by the
hoa carry individual vegetated islands or motu; frequently there is more than one such
vegetated island to a sector of conglomerate platform (Figures 7, 8, 9 and 10). The
channels between the vegetated islands may be carpeted with sediment and thinly
vegetated, and are less frequently occupied by water than the hoa.

The hoa and motu vary considerably in size. Most of the hoa are not more than 50-
100 meters in maximum width, and most of the motu vary from less than 100 up to about
600 meters in maximum east-west dimension. Profile | in Figure 5 gives a section through
one of the northern hoa. The seaward reef-flat surface rises to 1.1 m, with ponded living
microatolls on the outer reef flat at 0.15 m. Sand dunes on the seaward side of the motu
reach over 3 m. The conglomerate surface on which the motu stands declines from 1.1 to
0.4 m in a distance of 370 m from the seaward edge of the conglomerate to the lagoon.

Paleohoa

The term ‘paleohoa’ is here used to describe features now no longer active but which
clearly originated in the same way as modern hoa in the southwest sector of the atoll rim
(Figure 15). Profile 2 (Figure 5) crosses the island rim near the southwest point, where the
seaward perched beach ridge reaches a height of 4.0 m. The floor of the paleohoa, which is
completely dry, stands at approximately 1.5 m for its entire extent; it is scattered with
undercut storm boulders, especially on its seaward side, and these must date from the time
when the channels were active (Figures 17 and 18). The seaward end of the former

12

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Figure 9. Larger active hoa with through passage to the seaward side and well-developed
sediment lobe on the lagoon side, northern rim of Hull.

16

Figure 10. Motu with adjacent functional and non-functional hoa, northern rim at Hull.
This sector of conglomerate platform appears on hydrographic charts as a single island.

Figure 11. Conglomerate platform on the sides of a hoa, looking toward the lagoon,
northern rim of Hull.

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Figure 12. Conglomerate platform on the seaward side of a motu, northern rim of Hull.

2

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Figure 13. Incompletely eroded (non-functional) hoa, blocked by the conglomerate
platform.

18

Figure 14. Non-functional hoa which fails to transect the seaward conglomerate
platform, northern rim of Hull.

channel is now closed by a steep high ridge of storm rubble (Figures 18 and 19). The top
of the conglomerate platform along the channel walls stands at 1.8 to 1.94 m, and the level
of the island surface above the platform at 2.65 m. These elevations may be compared
with those of the active hoa in Profile 1, and with the crest of the algal ridge at 0.67 m,
which may be taken to approximate a datum in the range of Mean Low Water/Mean Sea
Level (i.e. a range of perhaps 30 cm).

DISCUSSION

We now consider the bearing of the hoa of Hull Atoll on the general questions of
hoa previously discussed.

First: the distribution of the hoa. Active hoa of the kind found on Hull are not
found elsewhere in the Phoenix Islands. They are notably absent from the other two
‘typical’ atolls of Canton and Gardner. Second, on Hull, the hoa are confined to a sector of
the northern rim.

i)

Figure 15. Sequence of paleohoa near the southwest point of Hull Atoll. The darker
areas are vegetated former motu.

Figure 16. Detail of motu and paleohoa shown in the foreground on Figure 15.

Figure 17. Basally-eroded storm blocks on the dry floor of paleohoa at Hull.

21

Figure 18. The dry floor of a paleohoa blocked in the background by a vegetated seaward
storm ridge, southwest rim of Hull.

Figure 19. Coarse coral rubble forming the storm ridge invading the seward ends of a
paleohoa at Hull.

22

This indicates to us that their primary cause must be local rather than regional. For
this reason we do not believe that a Holocene fall in sea-level, whether or not linked to
diagenetic changes in reef-top sediments as proposed by Chevalier and Bourrouilh-Le Jan,
is either a necessary or a sufficient condition for hoa formation. The most probable agency
for eroding the hoa is overtopping storm water associated with hurricanes, the action of
which has often been described. A higher Holocene sea-level may have been instrumental
in making possible the stranding of extensive spreads of sediment on reef tops as sea-level
fell, and which then became available for dissection into smaller motu by the cutting of hoa
during overtopping storms. Bourrouilh-Le Jan and Talandier’s (1985) suggestion of
erosion by tsunami-generated waves seems less likely, both because these are more
infrequent than hurricanes and also usually generate waves of negligible magnitude on
open-ocean atolls (see Stoddart and Walsh, 1992, p. 13).

The close spatial association of hoa on the north side and paleohoa on the west side
of Hull is also significant. The floors of the paleohoa and the upper surfaces of the
conglomerate platforms which margin them and which underlie the motu are substantially
higher (by the order of a meter) in the case of the paleohoa than in the case of the hoa.
Comparable paleohoa have been identified at Canton Atoll (Guinther, 1978) and at
Enderbury in the Phoenix Islands, and they thus appear to be a more widespread
phenomenon than currently active hoa. While paleohoa could be broadly categorized as
non-functional hoa in Chevalier’s terms, and while at Hull they are conspicuously blocked
on their seaward sides by a contemporary beach ridge of coarse storm sediments, their
non-functioning nature stems from their elevation rather than from sediment-blocking:
indeed the lodging of the beach ridge on the edge of the conglomerate platform was
doubtless aided by its elevation. At Enderbury the paleohoa are associated with lagoonal
emergent reefs of Holocene age (Tracey, 1972, 1980), and their floors stand at
approximately the same level as those at Hull (Stoddart and Fosberg, unpublished
surveys).

We suggest that these paleohoa were formed by the same kind of storm action that
is responsible for the formation of contemporary hoa, but at a period when sea-level stood
higher in the Holocene. They were subsequently abandoned when sea-level fell to its
present level and storm beaches accumulated along the seaward edges of emergent
conglomerate platforms and on emergent hoa floors.

We see no evidence that hoa are initiated by reef-flat fracturing, though doubtless
their development may be guided by joints. If fissuring or fracturing is involved, the
regional and local distribution of hoa raises considerable problems about surficial reef
structure.

Hoa are thus characteristic of reef rims with high proportions of land exposed by
the Holocene fall in sea-level and affected subsequently by major hurricanes. They are not
characteristic of reefs outside the hurricane seas, e.g. Diego Garcia Atoll in the Chagos
Archipelago, and indeed also Gardner and Canton Atolls in the Phoenix Islands. Historic
hurricanes are in fact rare in the Phoenix group, and the storm which formed the hoa on the
north side of Hull must have been an unusual event. Conversely both hurricanes and hoa
are common in the Tuamotu.

23

The presence of paleohoa at Enderbury and Canton as well as Hull suggests that
hurricanes may have been more frequent in this sector of the central Pacific in mid-
Holocene times. It would be of interest to search for paleohoa on the margins of the
modern hurricane belts to see if these features could serve as a general index of Holocene
hurricane extension.

ACKNOWLEDGEMENTS

Observations on Hull Atoll were made possible at the invitation of the U.S. Air
Force SAMTEC Project based on Canton Atoll. This provided transportation between and
logistic support on each island in the group. Fosberg’s participation was made possible by
the Smithsonian Institution, Washington, D.C., and Stoddart’s by the Royal Society of
London. We were accompanied in the field for much of the time by Mr Roger Clapp, then
of the U.S. Fish and Wildlife Service.

24
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Bourrouilh-Le Jan, F. G., Talandier, J. and Salvat, B. 1985. Early diagenesis from 6,000
years ago and the geomorphology of atoll rims in the Tuamotu. Proceedings of the
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Bryan, E.H., Jr. 1942. American Polynesia and the Hawaiian chain. Honolulu: Tongg
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Chevalier, J.-P. 1972. Observations sur les chenaux incomplets appelés hoa dans les atolls
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Chevalier, J.-P. 1973. Geomorphology and geology of coral reefs in French Polynesia. O.
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Chevalier, J.-P. and Salvat, B. 1976. Etude géomorphologique de |’ atoll fermé de Taiaro.
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Guilcher, A. 1988. Coral reef geomorphology. Chichester: John Wiley. 228 pp.

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Guinther, E. B. 1978. Observations on terrestrial surfaces and subsurface water as related
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25

Harmelin-Vivien, M., Charpy, L. and Morize, E. 1985. Atoll de Tikehau, Archipel des
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26

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ATOLL RESEARCH BULLETIN

NO. 395

STORY OF AN OCEANIC ARCHIPELAGO

BY

F. RAYMOND FOSBERG

ISSUED BY
NATIONAL MUSEUM OF NATURAL HISTORY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C., U.S.A.
FEBRUARY 1994

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STORY OF AN OCEANIC ARCHIPELAGO

BY;

F. RAYMOND FOSBERG

[Editor’s note: This was given as the Third Carl O. Sauer Memorial Lecture by Dr.
Fosberg on Thursday 23 October 1980 in the Alumni House Lounge of the University of
California at Berkeley. Carl Sauer (1890-1975) was Professor of Geography from 1923-
1957 and Chairman of the Department for 31 years. A group of colleagues, students and
friends established the fund to support the annual lectures.

The text was never published and is included here to show Dr. Fosberg’s ideas about
the past, present and future of oceanic islands. The text is from a typed copy corrected
in Dr. Fosberg’s hand. Among the corrections was a change of the announced title, "The
story of an oceanic mountain range". ]

From the time, at least several billions of years ago, when the outer layers of a molten,
or at least somewhat fluid, earth became solid enough to be called a crust, the interior,
or mantle on which this crust rested or floated, has apparently been in a state of
extremely slow but constant convective motion. If the crust ever formed a continuous
solid layer, it could not have persisted long with this slow-motion turbulence going on
in the mantle beneath. Inevitably fragmentation took place. Segments of different sizes,
but very large, formed, some with layers of lighter rock forming their upper surfaces. to
be called continents, others without. Convection in the mantle very slowly moved these
segments, or plates, of more rigid rock this way and that, jostling and grinding their
edges together. Their margins slid over and under each other, causing local melting,
volcanic and tectonic activity. Of the early eons of this stage in the planet’s history we
can know little or nothing, though recently a brash and scarcely convincing, but possibly
not too far off, attempt has been made to reconstruct the pre-Mesozoic movements of the
lighter or continental layers of those plates known to have such layers.

There seems to be some agreement that during the Permian period (280-225 million
years ago) these continental plates were united into a single principal mass, now called
Pangaea, leaving the remaining area of heavier, submerged crust as Panthalassa, the
World-Ocean or Pre-Pacific Ocean. Continued motion in the mantle pulled Pangaea apart
into major fragments corresponding in some slight
measure to the present continents or their precursors. Enclosed by them was a

Department of Botany, National Museum of National History, Smithsonian
Institution, Washington, D.C., U.S.A.

2

major portion of the World Ocean which became our vast Pacific.

Bathymetric maps and diagrams of the sea-bottom suggest that this great body of
water anything but deserved the name given it by Magellan. Great rifts, ridges, and
fracture zones, evidence of movements of plates continuing even to the present time, can
be seen. Some of these patterns on the sea-bottom show as ridges and chains of undersea
volcanoes. Whether these mountain ranges represent cracks in the sea floor, through
which vast amounts of molten rock have been extruded, or whether, as seems to be the
latest idea, permanent stationary weak points or *’hot spots" in the mantle cause local
melting of the plates as they slowly pass over them, resulting in successions of volcanoes,
oriented in the direction of movement of the plates, is not for the non-geophysicist to say.
Suffice it to know that outpourings of basaltic lava have slowly built enormous mountains
on the sea-floor. Some of these became high enough to extend for thousands of meters
above sea-level as, at first, bare, black, slaggy oceanic islands.

The present distribution of islands in the Pacific gives some idea of the widespread
occurrence of volcanic activity on the sea-bottom through the later geologic ages.
However, detailed study of the sea floor by means of echo-sounding apparatus reveals
hundreds, if not thousands of other submarine mountains that do not, at present, show on
the surface, even as reefs or banks. These are called sea-mounts. Many of them, given the
special name of guyots, have curiously flat tops, off of some of which have been dredged
the skeletons of shallow-water corals. These guyots are interpreted as volcanic islands,
which have once extended above the sea surface, that have been eroded down and have
gradually subsided, allowing coral reef platforms to grow on their tops. Continued
subsidence has in some cases been faster than reef-growth and now some have their flat
tops at 1000 to 2000 meters below present sea level, others still reach the surface as coral
atolls. Few people are aware of how many sea-mounts and guyots are known, as the
various navies of the world regard this as highly important strategic information and give
it a high security classification.

It is essential to the further development of our story to understand that these
enormous masses of volcanic material extruded on the sea-floor seem mostly to show a
tendency toward very slow subsidence. While there is general agreement on the fact of
this subsidence, its mechanism has been the subject of much controversy. Some
authorities favor the idea, first proposed by Charles Darwin, in support of his theory of
the formation of coral atolls, of widespread subsidence of large parts of the earth’s crust.
Others, such as Molengraaf and Davis, with perhaps more convincing arguments, favor
the idea of local isostatic adjustment, to compensate for the great weight of volcanic
material pressing on the immediate portion of the slightly fluid earth’s mantle beneath a
volcano. The result is very slow subsidence of each individual mountain or mountain
range until an isostatic equilibrium is reached, as with an iceberg floating in the sea.
Tectonic movement may, of course, elevate individual islands or portions of the crust so
that occasional islands or areas of sea bottom show features indicating elevation rather
than subsidence. Such exceptions to a general pattern do not necessarily invalidate the
pattern. Most islands and oceanic mountain ranges, as well as continental mountains,
show evidence of subsidence after early orogenic phases are over or slowed down.

For our story we could choose one of several existing Pacific island groups or
mountain chains - the Hawaiian, Samoan, Society, Austral, Tuamotu, or Marianas, all of

3

which show some of the features to be discussed. However, to make our story more
complete maybe it is better to create an imaginary mountain range, extending northwest
to southeast, somewhere on the Pacific sea-floor, that will exhibit all of the features and
kinds of islands that we want to discuss. Suffice it to be reasonably sure that all of the
phenomena to be described actually have existed or taken place somewhere. The thing to
remember is that the high peaks of our mountain range tower above the sea-floor as the
high Himalayas do over the Gangetic Plain, and that they emerge above the sea-surface
as islands, some of them for thousands of meters. The geomorphological, biological, and
cultural history of these islands is the subject of our lecture this evening.

We know all too little of the actual birth of submarine volcanoes, of the first extrusion
of molten basalt from vents in the sea floor, to form "pillow" lavas, which pile up into
masses of enormous volume and grow into eminences that herald their approaches to the
surface by "sulfur boils" discolored steaming patches on the sea surface. Such occurrences
have been witnessed a considerable number of times and even photographed. Breaking
of the surface by lava, itself, has less often been seen, especially by geologists and
volcanologists. The cream of Japanese volcanologists sailed forth to study one such
occurrence, Myojin Volcano, south of Japan. They apparently got too close, as they did
not return.

Myojin, and also Falcon island in the Tonga Group, have appeared above the surface
and then disappeared several times. The sudden cooling of the molten rock on contact
with sea-water causes a shattering of the glassy masses of material and the infant islands
are loose, unstable piles of cinders, clinkers, and pumice, which are soon reduced to sea-
level or below by wave-action, only to be pushed up again by more magma rising from
below. Finally there may be a big enough mass of this material to resist the attacks of the
waves long enough to build a permanently emergent subaerial volcano.

These basaltic oceanic volcanoes are usually of a gentler character than the explosive
Vesuvius, Lamington, and St. Helens type, found on continental masses and along plate
boundaries. Once a sufficiently broad base is built up a bit above sea-level, basalt flows
can be poured out on its surface, weighing it down and compacting it. Flow after flow
alternating with bed after bed of ash and cinders, added to the platform, gradually form
a wide dome-shaped slaggy black surface.

We have no way of knowing what the oceanic climatic patterns were many millions
of years ago, but given the vast expanse of the Pacific, then as now, uninterrupted by
continents or very large islands, there seems no reason to think that the climate then was
very different from now, except perhaps warmer during the Miocene. In all probability
there were prevailing northeast and southeast trade-winds, dry doldrums and _ horse-
latitudes, violent hurricanes, and powerful high elevation antitrades, and "jet-stream"
winds blowing eastward. We can observe the effects of these now on islands and infer
that they were similar a long time back. Trade-wind rains, latitudinal rainfall gradients,
orographic rainfall, rain-shadows, convection rains, widespread "kona" rainfall, heavy rain
that often accompanies hurricanes, occasional droughts, all doubtless occurred then as
now, making the climatic patterns probably similar to those we can now observe.

It can be safely assumed, I think, that from the very beginning, some of the same
dispersal agents as are active now brought occasional potential plant and animal colonists
to our young volcanic islands. Strong winds would have carried small seeds with

4

adaptations for wind dispersal, small flying insects, drifting spiders in their webs, from
other islands and continents. Typhoons, hurricanes, and their included tornadoes would
have picked up and carried heavier seeds and other propagules up to where the jet-streams
could have scattered them to great distances.

Wide ranging sea-birds would have visited our islands from the very first, depositing
guano and improving the habitats for plants, perhaps also bringing an occasional seed.
Storm-carried or off-course migrating land-birds might be expected occasionally, judging
by modern observations. They also may have occasionally brought seeds, either carried
internally or clinging to feet or feathers. If conditions were right the birds may even have
succeeded in establishing themselves, if they arrived in pairs or flocks. These events, of
course, happened extremely infrequently. Certain lowland or strand species, especially
plants, salt-tolerant and adapted to floating arrived, probably much more often carried by
currents. Marine organisms with free-floating or free-swimming stages doubtless
colonized as soon as suitable marine habitats were available, very shortly after stable
consolidated lava shores had formed.

Since the complete history of an oceanic island can never be observed, we are forced
to reconstruct its probable course from observation of many existing islands in various
stages, a method that provides ample opportunity for error. However, extensive
observation and experience, along with an element of judgment and estimation of
probabilities, may yield a fairly plausible scenario.

The ages of existing islands are matters of great uncertainty. Many years ago, when
I was concerned with the origin of the Hawaiian flora, I asked a Pacific geologist about
the probable age of the Hawaiian Islands. He said the geologists had no reliable means
of estimating this, except that, hopefully, the biologists might give some clues. He
suggested a tentative 10,000,000 years, taking into account the probable age of the
ancient, subsided northwest chain. For my own purposes, I "conservatively" adopted a
figure of 5,000,000 years. Since then drill cores have given a suggested late Cretaceous
age for the lowest levels of sediments in the Marshall Islands and at least earliest Miocene
for Midway, in the Hawaiian Leeward Group.

In considering the time that has been available for the development of the biota of a
closely associated group of islands it is essential to take into account the earliest land
surfaces, as it was most likely there that the first successful colonizations took place and
that the biota had its beginnings. Even islands now disappeared or reduced to sea-level
coral cays or atolls may well have been, in their earlier histories, the sites of significant
evolutionary events, of the origins of important lines of insular plants and animals. A
pertinent observation is that many complexes of plants and animals in particular
archipelagoes have related species on different islands in the groups that they inhabit. In
other words, inter-island colonization has been a rather more frequent occurrence. The
history of an island group must be looked at as an intricately connected and related series
of events. Many things were happening simultaneously.

As mentioned above, as soon as a stable cool surface and shore line were available,
colonization could begin. Planktonic larvae of benthic marine organisms could settle and
develop on the hardened rock of the shore-line. Birds could use the islands as resting and
nesting places, and initiate soil development with the nitrogen, calcium and phosphorus
compounds in their guano. Weathering of the ash and rock surfaces would begin at once.

5

Very soon (in a geologic sense) water-borne strand-plants, bird- and wind-carried seeds
and spores of pioneer rock-inhabiting plants could find a habitat. Ordinary rain showers
would occur, and on the windward slopes, orographic rainfall would be more abundant
the higher the island became. Plant-cover or vegetation would begin to appear. As soon
as this took place it was possible for small-animal colonization to start. In addition to the
sea-birds and their parasites, spiders and small insects probably came first, borne by the
wind. Spiders could arrive but not survive until sufficient populations of insects had built
up to provide them with food.

A feature of almost any early colonization would have been the extremely
impoverished genetic stocks represented by the single or very few individual colonists.
One might think that further evolution, requiring genetic variability as a base, would be
most unlikely. Yet much evolution did take place. The key to this paradox is that these
early habitats were completely open. In most or all species of organisms over repeated
generations, mutations of various sorts occur. Because so few of the total offspring
normally produced by an organism survive, the chances of even a beneficial mutation
surviving are almost infinitesimal under ordinary conditions. In an open habitat, however,
where almost any seed that falls in a favorable place is likely to produce a mature plant,
non-lethal mutations can rapidly accumulate in a population, providing the variability
requisite for evolution. The rigid control by competition that normally maintains the
identity of species is temporarily relaxed until the population numbers reach the saturation
point in a particular habitat and relative stability is achieved. By this time a normally
variable or heterozygous (potentially genetically diverse) condition may have come about.

In the earliest stages of Pacific basaltic islands there was, judging by modern examples
such as Mauna Loa, relatively little habitat diversity. Windward sides were wet, leeward
sides dry. Temperatures were lower at higher elevations. There were rough and smooth
lava surfaces and areas of ash and pumice. Strong winds occurred on high summits.
Isolating factors within an island were presumably almost entirely ecological. Populations
of organisms were free to spread over as much of the island surface as their ecological
amplitude permitted, and large, relatively uniform, though often genetically heterozygous,
populations may have been almost the rule.

However, the processes of weathering and soil-formation undoubtedly commenced as
soon as the volcanic material cooled and was exposed to sun, air, water, and, at high
elevations, frost. Loose material, whether ash and cinders or weathering products,
immediately started to be moved downward by water and wind, and to serve as abrasive
agents. This abrasion started to carve the volcanic land-forms and to produce erosion
features.

Those in the audience who have seen the windward coasts of the Hawaiian Islands
know that the heads and sides of valleys and gulches eroded in layered basalt are vertical
cliffs. Ridges, peaks, plateaus, and valleys bounded by such escarpments serve as very
effective isolation for most populations of plants and animals. Isolation of populations is,
as is well-known, the first step toward, and one of the essential conditions for, speciation.

By this erosion the widely distributed populations that formed on the gentle slopes of
the young volcanic domes were segregated into many isolated subpopulations. Inevitably
differentiation took place. Since this isolation was geographic, and effective, the resulting
varieties and species need not have developed genetic isolation or sterility barriers.

As this diversity of taxa developed, habitat diversification also proceeded. Where, in
a continental situation, with its wide spectrum of families, genera, and species, pre-
adapted forms would have been available for almost any habitat that developed, this was
not true on an oceanic island. Many taxa were developing but as variations in relatively
few families and genera. Dispersal was occurring and a new habitat sooner or later was
colonized, not necessarily by a form especially well suited to it, but by one able to
survive. Natural selection, even in a genetically relatively impoverished stock, would
gradually result in better adaptation to the habitat, even eventually leading to the often
strikingly well-adapted forms described in discussions of "adaptive radiation," a term
applied to the evolution of differing populations to occupy different available habitats.

As the normal geomorphic cycle proceeded, the sharp topography that maintained the
isolation of the small, mainly still interfertile, populations or taxa mentioned above,
became gradually worn down. Slopes became gentler, isolation became less effective, and
ranges of related taxa began to come together. Some of the previously discrete taxa
hybridized in these zones of contact. Their distinctions began to become fuzzy. A new
type of evolutionary process became prevalent. Occupation of existing habitats and newly
formed ones became more complete, and the patterns of adaptive radiation became more
intricate. By now the vegetation, except on new lava flows and landslide scars, had
become closed forest except at very high elevations and on exposed crests.

The birth, growth, erosion, and gradual subsidence of an island, its slow colonization
by organisms and the evolution and eventual decline of its biota are continuous, if
incredibly slow, processes, not really episodic as my description may have suggested. We
must remember that during the enormous span of time during which the first islands to
appear in our mountain range reached the maximum development of their biotas, one
after another, a whole series of younger islands emerged from the sea. Thus, at any one
time all or most of the principal stages or conditions described earlier were in existence
simultaneously. In addition to occasional successful new colonizations from outside the
young archipelago, more frequently plants and animals from islands within the group
succeeded in crossing the smaller water barriers between the islands. Thus the distinctive
biota of the archipelago was evolved and maintained.

As the millions of years passed, new volcanoes arose, extending the mountain range
to the southeast, and enlarging the archipelago as the Pacific plate slowly moved
northwestward.

The oldest islands on the northwest end of the chain gradually eroded away and
subsided, giving an opportunity for the myriads of calcium-carbonate-secreting, reef-
building organisms to grow and build the wondrous structures and communities of
organisms that we call coral-reefs. These reefs, at least ones of any large extent, are
formed on slowly subsiding coast-lines. Darwin, in his well-known theory of atoll
formation, showed the gradual change from fringing reefs, closely lining the island shores
to more distant barrier reefs ringing the partly subsided, eroded remnants of volcanoes
with coral just breaking the surface and carrying sand-cays and islets of coral sand and
debris. Eventually, when the central volcanic peak or peaks had subsided out of sight, a
ring of coral reef and coral islets, called an atoll, remained. The term atoll is modified
from the language of the Maldive Islands, a notable archipelago of these coral rings or
"necklaces" in the central Indian Ocean.

7

As this geological process took place, the relatively rich biota of the high island
became attenuated and impoverished in species as the terrain became more flattened and
simplified. The loss of diversity of habitats, as well as the increasing prevalence of salt-
spray, resulted in a drastic reduction of the biota. This was more conspicuous on dry than
on wet atolls. For example, the two driest of the Marshall Islands have only nine
flowering plant species each, while the wet southern atolls of the same group have up to
five to eight times as many.

Occasional tectonic activity results in elevation of islands already ringed with reefs,
and even of atolls. On elevated volcanoes fossil reefs may be seen as limestone terraces
or benches on the slopes, or as dissected walls of rough coral limestone at the bases of
the slopes. A few examples exist of elevated atolls and of extensive raised reef-tracts. The
elevated atolls appear as flat- or concave-topped limestone plateaus, and more extensive
tracts became eroded into weird karsts, often protruding ruggedly above the sea surface.
Such may now be seen in the southern islands of Palau, Micronesia, and in the Lau
Islands of Fiji. Elevated atolls may have vast deposits of calcium phosphate in their
summit depressions, as seen on Makatea and Nauru, presumably derived from the guano
of innumerable prehistoric seabirds.

In their pristine condition all of the variations of tropical oceanic islands described
above, except a few of the very driest, were clothed with forests, largely made up of
endemic species of trees and their accompanying endemic smaller plants, insects, and
other invertebrate animals, as well as birds, bats, and a few reptiles. Many of these
organisms had evolved weird and remarkable forms comparable to the tree lobelias and
the silver-sword of Hawaii and the dodo of Mauritius.

Of particular importance from our viewpoint, looking back from the future, were
several related features of the biota of these, and other oceanic islands. The plants and
animals had mostly evolved in the absence of large herbivores. Except for features which
the original colonists may have brought with them, the island plants had evolved no
means to protect themselves from the normal continental hazards of grazing, browsing,
or trampling by large animals. Even the thorns, spines, stinging hairs, bitter or poisonous
substances that may have characterized the species which originally colonized the islands
were mostly lost as the new biota evolved. Only some of the most recent arrivals were
still prickly, acrid, or otherwise disagreeable.

Root systems were frequently very shallow and had no adaptations to deal with
trampling, as there were no large animals to damage them by trampling.

The entire biota had evolved to take full advantage of a comparatively benign
environment, harsh in some respects, but without many of the hazards and kinds of
competition normal in continental situations. In many ways the equilibria achieved on the
islands were very delicately adjusted to make maximum use of existing resources by the
available range of organisms. Through adaptive radiation the limited original biota had
diversified to fill most ecological niches. All of this took many millions of years and was
prior to the arrival of man.

Humans clearly first came to oceanic islands in small parties, in sailing canoes,
possibly blown far out to sea by storms, or merely wandering, to see what was there. The
first-comers undoubtedly found very poor picking, except in the matter of sea-food and
birds and their eggs. The vegetation of oceanic islands contains few edible plants, indeed.

8

The popular idea of south-sea islands fringed with coconut palms is, as to pre-human
days, certainly a myth. Possibly the coconut is indigenous in the Seychelles, in the Indian
Ocean, as Jonathan Sauer has suggested, but it certainly came with man to the oceanic
Pacific islands.

Ancient Pacific peoples, probably even on casual canoe voyages, carried stocks of a
few edible plants, at least as provisions. Any serious colonizing party undoubtedly
brought an assortment of useful plants, and the knowledge of their culture. At least pigs,
dogs, and chickens came, as domestic animals, not necessarily at the same time. Thus the
early arrivals in our island group came well prepared to initiate the series of profound
changes in the island geography and biology that followed their advent.

Dependent, at first, on the resources of reef and ocean, the early settlers established
themselves near the sea, especially around the mouths of large valleys on the high islands,
and probably along lagoon margins on barrier islets and on atolls. Coconuts, bananas,
taro, breadfruit, and possibly sweet potatoes, were doubtless planted at once, as well as
some other plants. This involved a certain amount of clearing of native vegetation, but
for many years the effects were relatively minor. There were doubtless slight disturbances
in the reef fauna. Small flat areas back of the beaches and in valley bottoms were cleared
for villages, coconut and breadfruit groves, taro pits and other forms of garden culture.
A few cultivated plants and some "camp-followers" or weeds were added to the floras of
the islands. A few insects and other invertebrates (parasites on man and his domestic
animals and plants), some commensals and incidental accompanying animals (possibly
including rats) established themselves and become naturalized and part of the fauna.

While the human population remained small, changes in the island ecosystems were
slow and the results mostly not catastrophic. Most plants and animals that came with the
human colonists did not succeed in invading closed vegetation except where it was
opened up by man. An exception, perhaps, was the candlenut or kukui tree, probably
brought by man, which on some high, deeply dissected islands, occupied the 60-90 inch
rainfall belt on steep slopes almost to the exclusion of other plants.

Characteristic of oceanic islands is that their resources useful to man are limited both
in quantity and in kinds. Furthermore, islands of different geographical nature offer
different assortments and proportions of resources. Volcanic islands with considerable ash
in their composition weather more rapidly and yield richer soils than do those of hard
flow-basalt. On dry islands water may be limiting. On very wet ones severe leaching and
erosion may deplete soils more rapidly than soil-building processes develop them. Coral
atolls, especially drier ones, present a poor terrestrial environment, but are likely to be
rich in marine resources, especially if there is a good lagoon with active water circulation.
Elevated atolls are so well drained that water may be scarce, although the phosphatic soils
may be rich.

Cultural development and population growth depend both on the culture the colonists
brought with them and on the nature and abundance of resources occurring on the island.

In all likelihood, limited resources if not too extreme, served as a stimulus to cultural
development. To flourish it was necessary to learn to utilize fully and in diverse ways,
the range of resources provided by the island. Many island cultures show remarkable
ingenuity in their adaptations to their environments and utilization of everything available.
The diversity shown by the cultures found on our chain of islands in varying stages of

9

island "life histories" reflected the diversity of the islands. Settled by people of similar
Polynesian stock as diverse, of course, as the islands from which they came, the nature
of the island imposed each its own stamp on the further cultural evolution of its people.
Peoples on fertile, moderately wet islands tended to emphasize agriculture, or, rather,
horticulture, making a comfortable living. Populations grew rapidly. Atoll peoples
developed great skill and ingenuity in catching and using reef animals and often pelagic
fish. They also soon discovered the possibilities of pits excavated to the fresh-water lens
lying a few feet below the island surface. Culture of the several taros, and even of
bananas could be carried on in such pits, protected from salt-spray by windbreaks of
natural vegetation left when clearing for coconut and breadfruit groves, and as gardens
were established. The diversity of uses for coconuts and Pandanus, learned long before
in their ancestral homes, was increased in various directions.

A phenomenon surprising to us, now, was the population size achieved on the more
favorable islands. Since the colonists undoubtedly were a healthy lot, to be able to endure
the vicissitudes of long voyages and the hardships of settling new islands, and since their
numbers were small, the number of diseases they brought with them was small. The
inherent biotic potential of man being high, dense populations were reached in relatively
short periods. Undoubtedly the limits of carrying capacity of the different islands were
soon reached. Then the obvious controlling factor in many islands was warfare. Some
cultures, while able to defend themselves, found other means of population control.
Contraception was apparently fairly widely known. Infanticide was certainly not
unknown. Emigration and search for new island homes was a means of relieving
population pressure and a factor in the almost complete settlement of the habitable islands
in pre-European time. There were a few uninhabited Pacific islands at the time of arrival
of Europeans, but in almost every such case, the island was for one reason or another
unsuitable for habitation by Polynesians and Micronesians. Most of them showed signs
of having been visited and temporarily occupied. Lack of dependable water-supply was
the most frequent reason for lack of human habitation. Most populations on inhabited
islands, by the time Europeans arrived, seem to have achieved, but by one means or
another exceeded. reasonably sustainable levels. The few records made by the early
Europeans visitors show little or no evidence of over-population, depletion of resources,
or environmental degradation. For a few island cultures, such as that of the Marshall
Islands, we have information on actual conservation practices, usually woven into the
religious fabrics and apparently quite effective. What seems to have been a satisfactory,
functioning blend of material and social culture had usually been achieved. The occasional
very war-like societies may have been exceptions to this generality. Or, perhaps these
warring tendencies may have been merely the means evolved for population control. We
really know rather little of this period in island history.

One condition, very generally prevailing, was a lack of immunity to many of the
important human diseases. Apparently the early colonists in the Pacific came from peoples
who had enjoyed long isolation from the dense centers of human population where the
common diseases developed. Lack of exposure to these maladies resulted in lack of
selection pressure to evolve resistance to them. In a way this was comparable to the lack
of defenses, in the indigenous island vegetation, against the effects of large grazing and
trampling animals.

10

The centuries and millenia of presumably orderly and non-catastrophic evolution of
island cultures and their adjustment to their insular environments came to an abrupt end
in the second half of the Eighteenth Century. Up to that time, though European ships had
crossed the Pacific in several directions, the only actual beach-head of European culture
was the savagely destructive Spanish settlement in the Marianas Islands. The rest of
Micronesia and all of Polynesia were almost completely uninfluenced. Iron was known
to many island peoples but only as tiny bits presumably gleaned from rare floating ship-
timbers and other wreckage. A few other suggested traces of continental civilization are
known, such as the possible presence of pineapples, American in origin, and the existence
of these is still very conjectural and controversial. The presence of the American sweet
potato in Polynesia was certainly pre-European.

This slow evolution of highly developed cultures ended abruptly with the penetration
by English, French, and other explorers, especially Wallis, Cook, Bougainville, and
Vancouver, followed by traders, whalers, missionaries, and settlers. These brought
domestic herbivorous animals, steel tools, guns, alcohol and diseases. The results were
catastrophic.

A steady flow of exotic plants, both cultivated and weeds, followed. The domestic
herbivores escaped and became feral on most islands, opening up, eating and degrading
the highly vulnerable vegetation, providing new habitats for the exotic species. The steel
tools and the trading rewards for provisions for ships encouraged the clearing of more
land than had been needed to support the native populations; great changes in the
landscape became apparent. Tuberculosis, measles, colds, influenza, as well as over-
indulgence in alcohol decimated many populations. Mosquitoes, fleas, and lice came in,
and the diseases they spread became established.

Missionary activity destroyed the religious fabric that supported the indigenous
cultures. Fanatical new beliefs were introduced that altered vital features of these cultures.

Sailors and other visitors mixed with the natives and greatly altered the genetic
composition of the island populations. At the same time they brought venereal diseases
to further contribute to the population decline that was an outstanding feature of this
period. Some islands were virtually depopulated.

Meanwhile, land, the basis of the island civilizations, came more and more into the
hands of settlers from outside. Plantation agriculture, especially coconuts and sugar cane,
became widespread, usually on the best of the land. The children of the missionaries and
other settlers saw opportunities for Western-style enterprise and eagerly grasped them.
The second Missionary generation became sugar planters.

These events and activities had profound effects on the landscape, the biota, and the
social structure of the human populations of our islands. The low-island landscape, even
in its undisturbed state not very diverse or varied, changed from an appearance of
pleasant informality, dominated by groves of coconut palms and stately breadfruit trees,
with thatched huts, to a monotony of coconut plantations. The trees were planted in
strictly orderly rows, the houses had sheet-iron roofs, the ground kept free of
undergrowth, with all trash and humus neatly burned. Soil fertility tended to decline, as
the nuts were turned to copra and exported, taking nutrients along. Little attention was
given to replacing them. The humus content of the already poor soils became lower, both
from natural causes and from burning.

11

The level lowlands and gentle lower slopes of the high islands were likewise covered
by plantations, both sugar cane and coconut, later also pineapple. Towns and, later, cities
appeared, attracting the natives away from the villages, to populate the poor parts of the
cities, while the well-to-do outsiders built exclusive, beautiful homes in the better parts,
away from the crowded and ugly down-town sections.

The mountainous or hilly areas were devastated by the charcoal burners, feral goats,
sheep, cattle, horses and pigs. The forests were cut and degraded to poor scrub
increasingly dominated by lantana, guava, kiawe (Prosopis), and other weedy shrubs and
small trees. Fires became frequent in the drier leeward areas. Erosion was accelerated,
exposing bedrock on steeper slopes, and depositing fine red silt in the shallow water,
smothering the corals and degrading the reefs. Soil covering, with its important soil-biota
and accumulated nutrients that took millenia to develop, were lost in a few decades. The
capacity to support, not just humans, but populations of many kinds of organisms was
seriously reduced.

On each high island, a wondrous biota, largely composed of unique species, even
genera endemic to one or a few islands, suffered depletion and impoverishment. Many
of these species disappeared before they were known to science, or even known to man.
Such a loss is both an ecological tragedy of reduced diversity, and an impoverishment of
the interest and the aesthetic quality of the habitat of man. Some might say that if man
did not know what he lost it made no difference to him. I do not subscribe to this notion.

Loss of native species was not the only change in the biota brought about by the
stepped-up human activity and by the new human immigrants. People have a tendency
to introduce plants wherever they go. Economic or possible economic plants, food, forage,
fiber, timber plants, ornamentals, and "camp-followers" or weeds, came in, in great
numbers. Most were planted or established in fields, gardens, orchards, and around
dwellings. However, many plants have effective means of dispersal, mechanisms to take
advantage of wind, birds, people, and feral animals. The native vegetation tends to resist
invasion by exotic plants while in good condition. The degradation and opening up of this
closed forest vegetation by feral herbivores, fire, clearing, road-building and other kinds
of disturbance so effectively provided opportunities and habitats for exotics that, before
many years, the vegetation at low and moderate elevations on many of our islands was
dominated by these immigrants. Scrubby forests of kiawe (Prosopis), opiuma
(Pithecellobium), guava (Psidium), ironwood (Casuarina), mango (Mangifera), and
Eucalyptus, scrub of Lantana, Acacia, Melastoma, Leucaena, and Schinus, and rank
grasses (Pennisetum, Panicum, Andropogon, Saccharum, Melinis, and Sorghum) covered
vast areas in the lowlands and lower slopes, wherever the land was left uncultivated or
fallow. It soon became a fact that a visitor could travel in the islands for months without
seeing a native plant. The habitats of native species of both plants and animals were
usurped by these aliens, and the native biota, just as the native peoples, declined rapidly.
Hundreds of native plants, as well as snails, insects, and other invertebrates, along with
numbers of bird species, disappeared, mostly never to be seen again. Gentler upper slopes
were converted to pasture, making large livestock ranches. Even on steeper slopes, native
forests were invaded by exotic vines (Passiflora, Paederia) and aggressive shrubs
(Eupatorium, Lantana, Clidemia), or logged off and replaced by Eucalyptus or Pinus. The
landscape was altered beyond recognition and the natural diversity seriously reduced.

12

Due to erosion, water became scarcer. Irrigation of plantations became more and more
expensive. During dry cycles and on some of the drier islands, droughts occurred. Even
domestic water consumption at times had to be regulated. As the native vegetation, a
superb mechanism for absorbing and holding rain-water, became reduced, the water
supplies for the constantly augmented populations became more uncertain and precarious.

What of the native peoples? With introduced public-health measures, sanitation and
available medical services, the population decline was, in most islands, arrested, and
native peoples, mostly now of mixed blood, began to increase. However, because of their
period of decline, and often because of their disinterest in working for the plantation
owner, workers were scarce and the planters had to look elsewhere for their field and
factory labor. Large numbers of foreign workers were brought in. They stayed and
multiplied, in time becoming majorities of island human populations. They rapidly
adopted the dominant western cultures; the native peoples tended to do likewise. Thus the
remarkable local cultures that had evolved in the islands, disappeared. The peoples tended
to be more or less effectively amalgamated, but much of what had made them unique was
lost. A monotonous tropical version of western "civilization," with a few artificial,
imitated "native" forms of entertainment were what remained of the fabulous "South Seas"
for the tourists to flock to see.

We will pass over the period of war, with its profound and traumatic effects and
accelerated change.

Much of the scenery remains beautiful. Much of the planted exotic vegetation is
showy. The swimming is good. Away from the cities the pace and the atmosphere are
relaxing. The climate is pleasant. Visitors who come tend to want to stay.

In the cities the pace is frantic. Traffic is worse than in many continental cities. The
people are money-mad. Taxes are high. Everything is expensive. There seems to be no
limit to the engineering projects and construction of all sorts. Sky-scrapers sprout up
everywhere. Nothing is too beautiful to be sacrificed if it happens to be in the way of
"progress." Entertainment and comfort are provided for the tourists and retirees. If they
stopped coming, the metabolism of this frenetic society, fueled by tourist money, would
fall to a subsistence level that would barely be sustained by such plantation agriculture
as survived the immigration and tourist floods. So far, this has not happened, and fuel
energy has not yet become so expensive as to bring the mechanized civilization to a stop.

The South Sea idyll, which attracts the tourists and retired people, is now an artificial
and false counterfeit, but still convincing. Hordes of visitors come, many stay, and the
population grows. There are not small islands enough to supply the demands of the
wealthy who want private islands of their own. Solitude is becoming a memory only, but
it is not what the crowds want, though it is what the advertising promises. Air-travel
makes practically every island accessible.

What of the future? If energy supplies remain adequate, even though expensive, I can
see no future except that the islands will become more and more uniform, a tropical
version of standard western civilization. The diversity that still makes them fascinating
will more and more rapidly become lost. People will become even more ubiquitous than
now, as they will everywhere else.

If energy supplies fail, as they very well may, travel and communication will become
difficult and isolation will result. The amenities that have become necessities will become

13

more and more scarce, and more and more concentrated in the hands of fewer and fewer
people. The social structure will become more and more unstable. Local strong-men will
probably emerge. Violence and repression will prevail. Island versions of Ibanez’ Four
Horsemen may appear. Sooner or later the isolated, now impoverished and degraded
island ecosystems will have their human, animal, and plant populations reduced to the
levels of the real carrying capacities of their habitats. Perhaps limited but stable
subsistence societies may again become the rule. Or perhaps man may eliminate himself
and nature may start a new experiment.

PERTINENT LITERATURE

Carlquist, S. 1965. Island Life. Amer. Mus. Nat. Hist., Garden City, N.Y. 451 pp.

Carlquist, S. 1974. Island Biology. Columbia Univ. Pr., N.Y., London. 660 pp.

Dalrymple, G. B., E. A. Silver & E. D. Jackson. 1973. Origin of the Hawaiian

Islands. Amer. Scientist 61: 294-308.

Darwin, C. 1896. The Structure and Distribution of Coral Reefs. Appleton, N. Y. 344
Pp.

Davis, W. M. 1928. The Coral Reef Problem. Amer. Geogr. Soc. Sp. Pub. 9, 344 pp.

Fosberg, F. R., ed. 1963. Man’s Place in the Island Ecosystem. Bishop Mus. Press,
Honolulu, 264 pp.

Gressitt, J. L., ed. 1963. Pacific Basin Biogeography. Bishop Mus. Press, Honolulu,
563 pp.

MacDonald, G. A. & W. Kyselkon. 1967. Anatomy of an Island. Bishop Mus. Press,
Honolulu, 36 pp.

MacDonald, G. A. & A. T. Abbott. 1970. Volcanoes in the Sea... Univ. Hawaii Pr.,
Honolulu, 441 pp.

McKenzie, D. P. 1972. Plate Tectonics and Sea Floor Spreading. Amer. Scientist 60:
425-435.

Moore, J. G. 1975. Mechanism of Formation of Pillow Lava. Amer. Scientist 63: 269-
PLT,

Sauer, J. 1967. Plants and Man on the Seychelles Coast. Univ. Wisconsin Press,
Madison, 132 pp.

Zimmerman, E. C. 1948. Insects of Hawaii, vol. 1: Introduction. Univ. Hawaii Press,
Honolulu, 206 pp.

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ATOLL RESEARCH BULLETIN

NO. 396

COMMENTS ON ATOLL PHOSPHATE ROCK

BY

F. RAYMOND FOSBERG

ISSUED BY
NATIONAL MUSEUM OF NATURAL HISTORY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C., U.S.A.
FEBRUARY 1994

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COMMENTS ON ATOLL PHOSPHATE ROCK

BY

F. RAYMOND FOSBERG

Dr. K. A. Rodgers, in a series of previous articles (1987, 1989a, 1992), has gathered
together what is known, published or previously unpublished, on the occurrence of
terrestrial phosphate rock and phosphatic soil on the Tuvalu (Ellice) Archipelago of coral
atolls in the central Pacific, (the southern extension of the Marshall-Gilbert Chain). These
are typical low oceanic atolls, with no indication of any former connection with
continental land or large islands. From his text and maps of the individual atolls it is
evident that substantial areas of the "coral" sands and gravels that cover the surfaces of
these atolls have been phosphatized by whatever process or processes. This is the most
thorough treatment of phosphate occurrence on any non-elevated coral island group
anywhere. It is of great value in demonstrating the existence and importance of what is
a widespread geological and pedological phenomenon throughout the Indo-Pacific coral
island region.

Rodgers’ descriptions of the phosphatic material, though not always expressed in terms
completely familiar to me, indicate that all or most of the phosphate in Tuvalu
corresponds to what I have earlier (Fosberg, 1954, 1957; Fosberg and Carroll, 1965)
termed atoll phosphate rock, phosphatic hard-pan, or the Jemo Soil Series. In the above
cited papers, I presented a description of the bedded phosphatic rock or hard-pan found
commonly on all but the driest low coral islands in the Indo-Pacific. To account for the
origin and existence of this rock, I described a most interesting situation involving one
of the (formerly) most common forest types on these islands, a practically pure stand
forest of the large tree, Pisonia grandis R. Br. Under this forest, where its canopy is
layer or horizon of "raw humus" similar to the "mor" of northern conifer forests, with an
acidity of pH 6 to pH 4.5, or even lower. This occurrence of this type of "mor" is
otherwise very uncommon at low elevations in the tropics. Pure stands of any tree
species in the lowland tropics are also uncommon.

Sea-birds of several sorts tend to nest or roost gregariously in this forest, and where
this happens, the surface of the humus layer is stained white by the bird droppings, or
"guano". This is composed principally of finely comminuted fish-bones, largely a calcium
phosphate mixture. Beneath such white patches is commonly found an uppermost layer
of the coral-sand substrate cemented together, to varying degrees, by a bright brown
phosphatic cement, giving a very characteristic brown surface abundantly speckled with
white, the coarse sand or fine gravel carbonate particles. This hard-pan, when broken up,
can be detected by being notably lighter in weight than similar fragments of lithified coral

Department of Botany, National Museum of Natural History, Smithsonian Institution,
Washington, D.C. 20560, U.S.A.

2

sand or gravel. These observations suggested a process where the fine phosphatic powder
in the bird excrement was washed down into the porous acid raw-humus, acidified and
dissolved, the solution percolating down until it reached the alkaline calcareous sand
underlying the humus layer. It then, neutralized, precipitated out, forming a hardening
deposit, holding the sand-particles together. All degrees of this lithification were observed,
even involving small pebbles of carbonate, which tended to become friable under the
influence of the acid percolate, where part of the carbonate may have been replaced by
phosphate, rather than being washed off by the rain-water where no acid humus was
present.

Since this phenomenon was first observed by me on Jemo Island, northern Marshalls,
in 1951, I have, by watching for it in likely situations, seen it many times, in localities
from Polynesia to the Seychelle Islands, and occasionally in places where no Pisonia
remained. In such places, questioning the local people, I sometimes found that there had
formerly been Pisonia. In one or two instances, questions as to where there had formerly
been Pisonia, revealed unsuspected deposits of the speckled phosphatic rock, the surfaces
weathered to a dull gray color.

The earliest case of the association of such phosphate, mis-called "guano", to come to
my attention, was in a paper by Lipman and Taylor (1924), on Rose Atoll, where the
material was collected by A.G. Mayor (Setchell, 1924) and studied by C.B. Lipman and
J.K. Taylor. Lipman misinterpreted the nature of the phosphatic material, but there is
little doubt that it belonged to what I have called the Jemo Soil Series.

In 1970 I spent a few days on Cousin Island, in the Seychelles, Western Indian Ocean
(Fosberg, 1983). Over half of the 68 acre surface is a flat area, just about 2 m above sea-
level, now planted with coconuts growing in holes excavated to 1 m depth. Examination
of many of these holes and much of the flat surface showed this entire flat portion to be
a continuous layer of brown, white speckled, rather soft rock, exactly like the atoll
phosphate rock described from the Marshall Islands. Abundant young Pisonia grandis,
sprouting after clearing of what must have been an imposing forest, suggests that this
entire flat is a "beheaded" Jemo soil, the humus layer decomposed away after clearing.
The only other occurrence of atoll phosphate comparable in thickness to this to come to
my attention is a 5-foot-deep layer on Gaferut Island, Western Carolines, mentioned by
Niering (1961).

Sea-birds have been observed nesting in large numbers in forests of other tree species
than Pisonia, in bushes, and on the ground, on coral sand substrates. In no such situation
has there been found a raw-humus layer, or any brown, white speckled rock. One
exception to this was a description by Catala (1957) of a similar situation in the Gilbert
Islands, where the trees were cited as Guettarda speciosa forests, but none with a raw-
humus soil horizon. I can only assume that Catala, who is no botanist, mistakenly
identified Pisonia as Guettarda.

Rodgers (1992), mentions Pisonia trees wherever they occur in connection with the
phosphate deposits he describes. However, in only one place does he mention Pisonia as
dominant in the plant cover, on Nui Atoll, quoting Woodroffe (1985). He discusses the
problem in his section on geobotany, admitting that there may have been a Pisonia-

3

phosphate association in Tuvalu in the past, "man-induced changes make it impossible to
recognize such a relationship today throughout most of the archipelago, or indeed to
ascertain whether it was present in the virgin environment." He makes a considerable
point of the nonoccurrence of Pisonia on five islands today, also of the lack of bird
colonies associated with any known phosphate deposit in Tuvalu. One might expect 2000
years of human occupation to have obscured most evidences of natural conditions. He
concludes in this section, as well as in his abstract, that "the presence or absence of
Pisonia in the present day should not be taken as a geobotanical indicator." One can only
agree with this, without argument.

However, he says nothing about whether the presence of atoll phosphate rock can be
taken as an indicator of former Pisonia forests with colonies of roosting or nesting sea-
birds. I have in a number of instances assumed that the presence of such rock, as for
example, on Canton Island, Phoenix Group, a bed of weathered atoll phosphate rock, as
a very good indicator of former Pisona forests, where none occur today. This I maintain
even to the extent that a single well rounded, but unmistakable atoll phosphate pebble,
collected by the late Wayne Gagné on Laysan Island, indicates to me than a Pisonia forest
formerly existed on Laysan. Until recently, Pisonia grandis was not even considered to
be a member of the Hawaiian flora, but recent collections from Pearl and Hermes Atoll,
as well as two very old specimens in the Kew herbarium show that the species does occur
in the Hawaiian group.

One further observation, just to round out this discussion, and to indicate the need for
caution about sweeping generalizations, follows. That I can not insist, categorically that
Pisonia grandis is essential for the formation of atoll phosphate rock is shown by the
following. Samples collected by David Stoddart in 1961, on Lighthouse Reef, an atoll off
Belize, where Pisonia grandis is not even known in the same hemisphere, do not seem
to differ substantially from the atoll phosphates in the Pacific. And, in 1971, on Glover’s
Reef, another atoll off Belize, in the interior of Long Cay, in an area long ago cleared
and planted to coconuts, I located an area of many square meters of a weathered, but
when broken, bright orange brown white-speckled rock. The bed was not thick, but was
firm (Stoddart, Fosberg, and Sachet, 1982). Later, on Northeast Cay, I found small
boulders of similar rock in a pile of boulders, but none in place.

This Belize phosphate merits further study, but I have no doubt as to its nature. My
only suggestion as to its origin is that Neea choriophylla Standl., a small tree related to
the genus Pisonia, is fairly common to locally abundant on these Belize sand cays. I did
not see it forming pure stands, but before the cays were cleared and changed to coconut
plantations, such stands may have existed, and conceivably may have formed raw-humus
horizons on the soil.

Finally, to give some idea of how much change has taken place in recent years on
atolls throughout Polynesia, observations recorded in the voluminous reports of the U.S.
Fisheries Commission Albatross cruises written by Alexander Agassiz (1903), show that
around the beginning of the twentieth century Pisonia grandis forests were one of the
commonest and most conspicuous forest types on coral islands throughout the areas of
ocean visited by these expeditions. It would be surprising , indeed, if there were not
"fossil" occurrences of atoll phosphate rock on almost any of the not-too-dry Pacific
atolls.

Mention should be made of the very comprehensive review of Phosphate Rock on
Coral Reef Islands, by Stoddart and Scoffin (1983). This surveys the nature, occurrence
and petrology of phosphate rocks of a number of kinds, and undoubtedly of various
origins, on coral islands both high and low. These authors discuss what I have called
"atoll phosphate rock" along with several other forms of lithified phosphate under their
heading "recent phosphate rock". They discuss these deposits at considerable length, but,
in my opinion, do not clearly indicate whether rocks of one, several, or even many
origins are discussed in their two sections headed "Recent phosphate rock" (pp. 376-378,
and 380-382). Nowhere do they indicate that acidity is an essential factor in the process
of formation, nor do they emphasize the bright brown color of the matrix or cement in
which the white limestone grains are embedded. To me, the presence of acidity is
essential for the solution of the comminuted fish-bones of the guano and its downward
percolation through the soil. There also seems to be no other obvious origin for the bright
orange-brown color of the cement than the brown raw-humus produced under the Pisonia
forest. I would restrict the definition of atoll phosphate rock to occurrences where the.
matrix is brown and inclusions are white or pale calcareous particles, or their
pseudomorphs in very altered examples.

With this restricted definition, I strongly maintain that such rock is an indicator of a
practically pure stand of Pisonia grandis (or equivalent such as Neca) with nesting or
roosting gregarious sea-birds, on "coral" sand or gravel. None of the other suggested
processes of origin than that described above seems adequate, or necessary, to explain the
origin and occurrence of the rock (or soil) under discussion.

Also, I would scarcely expect to find, on atolls as long and densely populated by
humans as Tuvalu, more than casual persisting remnants of Pisonia vegetation. I have
noted a strong inverse correlation of present-day seabird rookeries with human
populations. Finally I have seen few, if any, uninhabited coral islands that do not have
large or enormous colonies of fish-eating birds. Lack of birds, and only a few remnant
occurrences of Pisonia grandis, are exactly what Rodgers describes for Tuvalu, where
persisting beds of what seems to be typical atoll phosphate rock seem to be common. To
me, this suggests that dense stands of Pisonia and vast numbers of seabirds were
characteristic of the Tuvalu atolls in pre-human times.

References

Agassiz, A. 1903. The coral reefs of the Tropical Pacific. Mem. Mus. Comp. Zool.
Harvard. 28: 1-410, (3 vol. plates)

Catala, R.L.A. 1957. Report on the Gilbert Islands: Some aspects of human ecology. Atoll
Res. Bull 5: 1-187.

Fosberg, F.R. 1954. Soils of the Northern Marshall Atolls, with special reference to the
Jemo Series. Soil Sci. 78: 99-107.

Fosberg, F.R. 1957. Description and occurrence of atoll phosphate rock in Micronesia.
Amer. J. Sci. 255: 584-592.

Fosberg, F.R. 1983, Natural History of Cousin Island. In Sachet, Stoddart, and Fosberg,
eds., Floristics and ecology of Western Indian Ocean islands. Atoll Res. Bull. 273:7-38.

Fosberg, F.R. and D. Carroll. 1965. Terrestrial sediments and soils of the northern
Marshall Islands. Atoll Res. Bull. 113: 1-156.

Lipman, C.B. and J.K. Taylor. 1924. Bacteriological studies on Rose Islet soils. Carnegie
Inst. Marine Biol. Pap. 19: 201-208.

Niering, W.A. 1961. Observations on Puluwat and Gaferut, Caroline Islands, with
historical and climatic information of Gaferut Island by M.H. Sachet. Atoll Res. Bull.
OR, NS

Rodgers, K.A. 1987. The mineralogy of a phosphatic horizon, Amatuku islet, Funafuti
atoll, Tuvalu. S. Pac. J. Nat. Sci. 9: 49-56.

Rodgers, K.A. 1989a. Phosphatic limestones from Tuvalu (Ellice islands). Econ. Geol.
84: 2252-2226 [sic].

Rodgers, K.A. 1989b. Dahlite and Whitlockite from Amatuku, Tuvalu. Mineral Mag. 53:
123-125.

Rodgers, K.A. 1992. Occurrences of phosphate rock and associated soil in Tuvalu, Central
Pacific. Atoll Res. Bull. 360:1-31.

Setchell, W.A. 1924. American Samoa. Part III. Vegetation of Rose Atoll. Carnegie Inst.
Pub. 341: 225-261.

Stoddart, D.R., F.R. Fosberg, and M.H. Sachet. 1982. Ten years of change on the
Glover’s Reef cays. Atoll Res. Bull. 257: I-II, 1-17, f. 1-15, pl. 1-18.

Stoddart, D.R. and T.P. Scoffin 1983. Phosphate rock on coral reef islands in Goudie,

A.G. and K. Pye eds. Chemical sediments and geomorphology. Academic Press, London
pp. 369-400.

Woodroffe, C.D. 1985. Vegetation and flora of Nui Atoll, Tuvalu. Atoll Res. Bull. 283:
1-18.

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ATOLL RESEARCH BULLETIN
NOS. 390-396

NO. 390.
NO. 391.
NO. 392.
NO. 393.
NO. 394.
NO. 395.

NO. 396.

LISTS OF TAXA NAMED FOR F. RAYMOND FOSBERG
BY DAN H. NICOLSON

ANNOTATED LIST OF THE KNOWN PUBLICATIONS OF F. RAYMONI
BY ANNA L. WEITZMAN

THE FLORA OF NAURU
BY R.R. THAMAN, F.R. FOSBERG, H.I. MANNER, AND D.C. HASSALL

FLORA OF THE PHOENIX ISLANDS, CENTRAL PACIFIC
BY F. RAYMOND FOSBERG AND DAVID R. STODDART

THE HOA OF HULL ATOLL AND THE PROBLEM OF HOA
BY DAVID R. STODDART AND F. RAYMOND FOSBERG

STORY OF AN OCEANIC ARCHIPELAGO
BY F. RAYMOND FOSBERG

COMMENTS ON ATOLL PHOSPHATE ROCK
BY F. RAYMOND FOSBERG

We

% 7 7 As.

THE NATURAL HISTORY OF CAROLINE ATOLL,
SOUTHERN LINE ISLANDS

Issued by

NATIONAL MUSEUM OF NATURAL HISTORY

SMITHSONIAN INSTITUTION
WASHINGTON, D.C. U.S.A.
FEBRUARY 1994

ATOLL RESEARCH BULLETIN NOS. 397-398

RESEARCH
BULLETIN

ATOLL RESEARCH BULLETIN

NOS. 397-398

THE NATURAL HISTORY OF CAROLINE ATOLL,
SOUTHERN LINE ISLANDS

EDITED BY

ANGELA K. KEPLER AND CAMERON B. KEPLER

ISSUED BY
NATIONAL MUSEUM OF NATURAL HISTORY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C., U.S.A.
FEBRUARY 1994

VITRLIUG &

QO TSI LAMUTAM 9O MuaetiM JAROITAM
POPTUTTTeh! WAINORITTIME
CEU 2 SRTOVERIAW P
poet VRAIS

ATOLL RESEARCH BULLETIN

NOS. 397-398

NO. 397. PART I. HISTORY, PHYSIOGRAPHY, BOTANY, AND ISLET

DESCRIPTIONS
BY ANGELA K. KEPLER AND CAMERON B. KEPLER

NO. 398. PART If. SEABIRDS, OTHER TERRESTRIAL ANIMALS, AND

CONSERVATION
BY CAMERON B. KEPLER, ANGELA K. KEPLER, AND

DAVID H. ELLIS

ISSUED BY
NATIONAL MUSEUM OF NATURAL HISTORY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C., U.S.A.
FEBRUARY 1994

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ACKNOWLEDGMENT

The Atoll Research Bulletin is issued by the Smithsonian Institution to
provide an outlet for information on the biota of tropical islands and reefs
and on the environment that supports the biota. The Bulletin is supported by
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The Bulletin was founded in 1951 and the first 117 numbers were issued by
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reports resulting from the Pacific Science Board’s Coral Atoll Program.

All statements made in papers published in the Atoll Research Bulletin are
the sole responsibility of the authors and do not necessarily represent the
views of the Smithsonian nor of the editors of the Bulletin.

Articles submitted for publication in the Atoll Research Bulletin should
be original papers in a format similar to that found in recent issues of the
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accepted, the author will be provided with a page format with which to prepare
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COORDINATING EDITOR

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Smithsonian Institution
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Wayne N. Mathis (MRC-169) Washington, D.C. 20560

Victor G. Springer (MRC-159)

Joshua I. Tracey, Jr. (MRC-137)

Warren L. Wagner (MRC-166)

Roger B. Clapp (MRC-111) National Museum of Natural History

National Biological Survey
Smithsonian Institution
Washington, D.C. 20560

David R. Stoddart Department of Geography
501 Earth Sciences Building
University of California
Berkeley, CA 94720

Bernard M. Salvat Laboratoire de Biologie & Ecologie
Tropicalé et Méditerranéenne
Ecole Pratique des Hautes Etudes
Labo. Biologie Marine et Malacologie
Université de Perpignan
66025 Perpignan Cedex, France

CAROLINE ATOLL

0) 1000 2000
Cp st
METERS

Frontispiece. Airphoto mosaic of
Caroline Atoll, RNZAF 6569.
Reproduced by permission of the
Lands and Survey Department,

New Zealand.

Preface

This study is primarily a result of the research efforts of the
First Joint US-USSR Central Pacific Expedition (Line and Phoenix Groups,
Gilbert Islands, Micronesia, inland Philippine seas, and South China
Sea). In turn, it was part of the second phase of a longer expedition
from Vladivostok, USSR, via Dutch Harbor, Unalaska, Aleutian Islands to
Singapore (26 July to 31 October 1988), which included oceanographic and
seabird investigations in arctic, temperate and tropical waters.

The authors boarded the Soviet Research Vessel (R/V) Akademik
Korolev (7,000 tons, 124 m in length) in Hilo, Hawaii, bound for
Christmas Island (02°N, 157°W), where we picked up Katino Teeb’aki, a
conservation officer for the Republic of Kiribati, who represented his
government and helped our land-based research efforts. The next stop
was at little-known Caroline Atoll (10°S, 150°W), on the southeastern
edge of the Line Group. After landing on Caroline on 22 September, we
camped in 2 locations for 7 nights, surveying the terrestrial plants and
animals on all 39 islets.

A small amount of data was also added from 2 visits of the ICBP
(International Council for Bird Preservation) 1990 Line and Phoenix
Islands Expedition (2 March to 31 May 1990) aboard the New Zealand
cutter, R/V Te Manu (7.5 tons, 10.5 m).

Caroline is a relatively untouched atoll with its native plant
communities nearly intact on all but 3 islets, and teeming seabird
communities that, collectively, are second in the Line Group only to
Christmas Island (Kiribati) in diversity. For several historical
reasons, the natural values of this spectacular blend of marine and
terrestrial resources have been overlooked.

These papers attempt to synthesize what is known about the atoll.
We have analyzed our own data in the light of several important
historical accounts from the "gray literature," including records from
early navigators, guano and copra enterprises, incidental travellers,
and a solar eclipse expedition, some of which were critical to
understanding the present situation. We include a new map of the atoll
(the first for over 100 years, with additional islets and shoals), the
first descriptions of physiography, plant communities and floristics,
and individual islets, the first published climatic records and the
first photograph and measurements of its intact, ancient Tuamotuan marae
(religious site). We also report 14 new species records (5 plants, 2
seabirds, 1 shorebird, 1 land bird, 3 lizards, 1 possible turtle, 1
marine mammal).

In Part I we have attempted to reconstruct and describe Caroline’s
botanical history, including ecological succession, species-area
relationships, and the remarkable recovery of its indigenous forests
(Pisonia grandis, Cordia subcordata, Tournefortia argentea). In
Part II, discussions of seabirds and coconut crabs are rounded off by
bringing to light current threats to the atoll’s well-being, and ongoing
conservation efforts.

An expedition of this magnitude entailed the help of many people,
and it gives us great pleasure to thank them. We are indebted to Hal
O’Connor and Randy Perry, Patuxent Wildlife Research Center, for making
possible our participation. Steve Kohl, FWS Office of International
Affairs, and Terry Whitledge aided immensely by handling innumerable
details with their Soviet colleagues. Members of the Fish and Wildlife
Service Mauna Loa Field Station, especially Jim Jacobi, Julie Williams,
Jack Jeffrey, and Martha Moore, provided welcomed logistical support in
Hilo, and Paul Sykes willingly shouldered additional responsibility that
freed CBK to join the expedition.

On the Soviet side, we thank Professor Alla V. Tsyban, Chief
Scientist of the expedition, for extensive help and friendship during
the voyage, Capt. Oleg Rostovtsev, Yevgeniy Nelepov, Yuri Volodkovich,
and the crew for ship-to-shore transport, and Svetlana Petrovskaya and
Valeriya Vronskaya for translation. We thank Greg Smith and Chuck
Stafford for Zodiac transport at Caroline.

Katino Teeb’aki shared the transect work; his skills at obtaining
and opening coconuts often energized us. Abureti Takaio, former
Minister for the Line and Phoenix Groups, permitted us to work on
Caroline and, with the residents of Christmas Island, arranged a
memorable evening of dancing and food, despite the fact that their last
supply ship was 10 months previous.

Financial assistance for the 1988 expedition and for writing the
manuscript was provided by the U.S. Fish and Wildlife Service, Patuxent
Wildlife Research Center, and the Natural Environment and Climate
Monitoring Laboratory, Goskomgidromet USSR.

Grateful thanks are extended to Derral Herbst for identifying and
preparing plant specimens (deposited in the B. P. Bishop Museum,
Honolulu, Hawaii) and George Zug for identifying lizards (deposited in
the U.S. National Museum). The Royal New Zealand Air Force supplied the
aerial photos. Roger Clapp, Ray Fosberg, Gene Helfman, Ernst Reese, and
David Stoddart shared unpublished manuscripts and other information. We
are grateful to Lynda Garrett and Wanda Manning (Patuxent Wildlife
Research Center Library, Laurel, Maryland) for digging out obscure
historical references. Harry Maude of the Australian National
University supplied plantation records and other literature
indispensable in understanding Caroline’s past and present ecology. The
libraries and herbarium at the University of Georgia were also helpful.
Thanks to Dorothy Schaumberg, Mary Lea Shane Archives of the Lick
Observatory, University of California, Santa Cruz, for the letter from
W. W. Campbell (1908) and extracts from E. B. Campbell’s unpublished
diary (1908). We especially thank Bonnie Fancher for her efficiency,
enthusiasm, and hard work, often late at night and on weekends, on the
computer and in other clerical matters in preparing the monograph. The
manuscript has benefitted from reviews by Roger Clapp, Ron and Anne
Falconer, Ray Fosberg, Ian Macintyre, Mark Merlin, Pat Roscigno, Betty |
Ann Schreiber, Fred Sibley, Thomas Spencer, Terry Whitledge, and Stephen
Zeeman. |

AKK, as co-leader of the ICBP 1990 expedition, expresses much
gratitude to Christoff Imboden, International Council for Bird
Preservation, Cambridge, U.K., and co-leader/expedition initiator Martin
Garnett, for sharing finances. Thanks also to Annie Garnett, John
Phillips, and Mark Linsley for help with field work, and to Alve
Henricson for his sailing skills. The expedition would not have been
successful without the dedication of Capt. Graham Wragg, skipper/owner
of Te Manu, who transported us 7,585 km in the central Pacific
(including 2 visits to Caroline), helped with field work, shared
finances, and whose competence and consideration in many areas eased the
varied hardships associated with 3 months at sea in a 10.5-m ketch.
Thanks also to Sandy Bartle, National Museum, Wellington, New Zealand,
for support, and to Scott Miller for providing insect vials, and David
Preston for preparing and depositing insect specimens in the Bishop
Museum, Hawaii.

On remote Caroline, the Falconers were exceptionally hospitable
hosts, helping us with field work during, and after, the expedition.
Special thanks go to 7-year old Alexandre, who discovered the first
Blue-gray Noddy nest for the island and 3 new plant records. French
Polynesian residents who assisted in various ways include Jacques
Florence, Les and Gloria Whiteley, Rick Steger, Michael Poole, Jean
Roudeix, and friends who supplied us with fresh provisions.

We particularly thank those who have aided us in follow-up
conservation efforts: Kelvin Taketa, Jim Maragos and staff of The
Nature Conservancy-Hawaii, Christoff Imboden and staff at ICBP, Alex
du Prel, Jean-Michael Chazine, Philippe Siu, George Monet, Graham Wragg,
Papeete Customs, George Ariyoshi, Kaiarake Taburuea of the Ministry of
the Line and Phoenix Islands, John Claasen, New Zealand High Commission,
Tarawa, and Peter Timeon, Ministry of Foreign Affairs, Tarawa.

AKK, CBK, DHE

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ATOLL RESEARCH BULLETIN

NO. 397

PART I. HISTORY, PHYSIOGRAPHY, BOTANY, AND
ISLE DESCRIPTIONS

BY

ANGELA K. KEPLER AND CAMERON B. KEPLER

ISSUED BY
NATIONAL MUSEUM OF NATURAL HISTORY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C., U.S.A.
FEBRUARY 1994

ACTA LIUe

‘ Pane |
(POTS GAMUTAM TO MORAN ARORA
VATU TTA MADOOCEEIAA

2.0 2 OTORTHAAW
mel YSAUREAN

Table of Contents

[BeSiE ORM RI GUGES aes come mreets see tes mieten wets Come me lare Qreerene tree i | MAING Soa UE
INSEE Of MPa eS ee eewremenne es, miscais Wane che meted Aterere Seked = SEIN pent man ar eG
Sia OnlabileStvramevemeers eects ee eee eee eM ry See ER ibs Bay
[NOS UEIRENG Baikes creates oltatiatd Gini ctanicie herent paren Aakash Rom rh air A de ee
emu ROGUGTMON rm crrretin tates sees hs ot Soult a Sra ecas tia Gere tines bb cece ere ts
Behistorny-OF “Caroline wALOl Maras tdcnt estes esss coe ces cree sales oc
Pre-European History: Tuamotuan Period..............ceceeccees
Post-European History: 17th to 19th Centuries.................
MNESGUANOMEV Aree tke cette ck he Sooke Seca tienen ects eee ean diese bo
SollapEGhinsemexpeditions. tk vse come eese to eee sete see os
likemLatewlOth_and=z0th Centuries ss. ores st ee sess Vee ee et ce ea ee

CRNCENOUS eit ce on eee ek Ne roc y se wee Tete yee eS ft ED EE NS
ICTIGEC CH MGUCSmrm erm hres Siete Ce tenes ce en es SMe Ve OL MN oss
Naming*GaroltinesseMotuse cist. Vest sta st ot Sh Sees es eee Seca ses

Deereiaminary Structurevand TOpogrvaphyss2 sc. cscs eter esse ets ces oes
Backgrounds Geollogical#Settingi.. 6. sts seats suns tee es Sete es fas

Sear LevedaChangess eat, sen oes ees eee ek peek oe pe eae eee.
General ACCOUNT eae Fe ee ee eee eee eG ee ee gos 53
Reetasl auSeymunntrien cet he tic oe weet ete eens Lee nated lao e g
SSIS) NS ar arene eth nga Re ae ee i ed Sec Lb ee
AG OOM see eerie e Sa eet ee ria oe eee ee eas Seen eM ARES Sik S

agoons HyYdBOlOGycemccr nie ee cee Chee Grete ce iecea ars

Pacha Reenisrmmaeac aa racrs Nile tte create ee ee ee ene eo ee aisle

inidacna-AcroporaeReenss: ssc ans ttc tee ot ete te eee see t es
aGOOnMREeielatSee mercer cc eects cc ret est ote ee cecete aueueng

Pagoon=Reets Patina "A Brier “SUMMANSY vce cietrs + t.cte's oe eee te sc.ee s+
SLOSS CH Ge bess reese tate ten COR Br mokciir AC RRC ICL Pr eRe RCIC RA RAGA ERC aac Mk eral are
HY CRONOGY Eee nen emtr tea che eee ye RIS CM abe Ree CME 8 eee etre
Cara ORE arts sina 9 aes sek tatoos, emer cond aires,» Men AM Sarno eR TEN oc

Windtand=Ratintallilrsses tn sites ee) RU Je OREN. sce sae

CY CHONMERS EONS Peeters ter teeter fe tare ee tote Tote tere cote otete tet ialicvettnns

Seas COnadiGonser so cee create et etcetera ee ta wma Rn

EmlOnds= sVaSciillanerlantseand PlOniSticSs ss. c: cece ce steclae ts sore et cs
Botanical e HiStowyere-tcst ise sorte cite etetere ere eels wiee ome eae rene eels chine cers
Vascular Plants ot CarolimevAtolliins.) ss ccc ste Metis cre tie olers ott one's

PilanteCoimMectionsee see este oe os ete ee fore cle wale w ela wlceeletes
Species Lists, Annotated Checklist and Maps of
MEBKESCRI al sVaASCUlAaneP tantSi ces os coe Seidole Sere be sls crspoleeels
Flomisvics and -Ecology#of thesMotuse.....-+-scs5. cos sede. cece.
SIZE" Of SENOHPI OFA Siero k nc et cd ads caedtacsoese as sede es oa
Numbenssofelindigenous@Plants).... icc. cmon o -clecis cc in el
Compos itiOnvor “CHEMPlLONA sts ..« caci< o cice eo cle ee ie ors cco seus wie ove

Fee, ECOMOGIICASUCCESSITIONe « cise sie ccie cos doe tee code eesa ceed sde che es
BaSiiGwSeiralleeSiva gO See sevarrcke orcpet cc forcvere rel she sicieisieichere viele te viele Gol ajalsreie ieee
Ecological Succession on Motus of Different Size Classes.......
SpeenescArca tREVatiOnsnipSadces cea. e ches dace dsccceaeheccss eee d

Comparisons of Species-Area Relationships with Other
OMNES fererere seve yone creas siierv otcuctcleucas tar Gre shane: erelieievel eo etesteieeshel crarsieue'e: ss shaile

il

The Question of Fresh Water... uts7.-0..+osee ae oe eee 87
Motu Size in Relation to the Distribution-of Trees,

Shrubs ‘and Herbs. .....ch2< cc c.ce nec se be cme One 89
Gee Plant Communit less sic... <c c:c.c:0 s.0,010 ceases of oe Oeiehe eee eee 90
General ACCOUNT «:..¢6 6 s-scs « o:s.6.0' ois 6% 0) 5-0 6 ove one ons oes een enn a 90
Natural Herb (Mat... 03.000 sc ss Gere soles os sistolosleleioe Cre eee 90
Seabird USO. ss... dsc. soe ws erseisiclos sicrerciols See er eae 93
Beach Scrub: with .Surtianal.:...ca.06ees ces soil cen eee eee 93
Pandanus “FOVE@SU s.s.5.c 0:6 0.4:s 0 0:5 016 dsle'd oo ous oe RINSE Sik eo ee 93
jiournefortia Scrub and) Forest. .c..sseme ower bi eee eee Ceo 94
General Distribution. ~cee ..¢.eems oocun eee eee ene 94
Species Diversity in Tournefortia Woodlands............... 96
Stature and Area Coverage. . ....<5. sjc,0% +,0.0)5 © Bats ons .o Sint olen 97
ECOTOGY .. 0: os <:s0.0 0: 0:5 075 0: 0:0 fogs neye Guae?sle AATEC shales oie ee Ten ee 98
Seabird US@:........004 6 sagstawet c0e% & Dew biG oom ROR Eee 99
Corda. FOvests..2.6 6.00 ness oo eiowlers oe 06ers Sle Ae Oeee ee 99
General DistributiOn).<...%.00..000 «<0 oom ieee Ree 99
HUSTON Y ss s5.0 0050.8 sae 6 4 o-susie orale oreie a eye sie eIG Ie b RRC ene 100
Abundance and Distributilonien.ces shee ene 100
Seabird Use. occc. ssc ccus cs coteb tees hes bem tee Cen 100
Pisontia FOVOS bs. oc.66.00.600.5.6.0 08 46 bcssie's 3.05 SRE ROR ERED. 101
General Distribution... sos. .s% 0% «os 0 o> sceysnseern Oe een 101
An Historical Perspective... ......s000s00-s0+s 0p eh eeen 101
Annual Growth Rates. <.occ d.cie.e«: si0.0.0a00 ae 0-6 o1s 3 o16s Or eee 108
Species Diversity in Pisonia Forests........... >.< 109
ECO) OGY «0:0: 0 <0 0.0 cisia'se wie ic bers © 0 ae) «ks peys Byanpiuchs “Ree 110

Relationships Between Pisonia Forest Height and Motu
DiAMENS TONS .:0. 00 < 0:0-%.550, 0 15.0, soso eke opens pepoxcnouste Gkenor RRO Ieee 110
Pisonta-Seabird Relationships. « «+ os sic Gte otorsse(s epoveyeregels helene liz
Remnant, Pisonia Forestsp in, the PaciifiiCen et -rcvercnse einen 113
Coconut. Wood Tangs «.<...5 3 :c.0.0:0 ssccccece hen ose'o 61505 Gisls «See lS Ouse OR ene 13}
General’ Distribution... << 6 c.cco0 + 0000s ecleore see eee 113
HAS COPY evs cos sons chars So arens. s Sjosie eels wis Sue sie wis el Shekels See ae C eae 115
Distribution and Abundance. ........t6.7cKem er fob 1 a7/
S@abiird WSE.. oo. 6.63 osc sie ave, 6 01018 6 ere 6:08 wuomonopottey omeRskeye emere ier cnenenene 118
Absent Plant Communities... <6.) <0/s <.c::0-crele.& ssopore te toe Poppet meee rene ener 118
H.: Description and Ecology of the Motus........... +.» sspolsih ose 119
Tee CONCTUSTON: <<... 00's 00 viscetere ce openele beeline BS depens ome pie Ree nee IWy/
dee leiterature: Cited: osc sccscaoe sos so eecerere 61a aco e oie 0s) Spenepetewe eGo eae een 172
Appendix I: Reef Information for Navigators... s 2%... his «mise 183

Appendix II: Weather Data, 1989-1990. 2. ... . . <ccsretevorcuste lope’ = lepeucleteieleneienene 184

List of Figures

Frontispiece Air-mosaic of Caroline Atoll, RNZAF 6569.
Reproduced by permission of the Department of
Lands and Survey Information, Wellington,
New Zealand.

1 Line Islands: geographic location in the Pacific Ocean.....

2 Caroline Atoll, Republic of Kiribati, with newly-named

3. Main marae on Nake Island, Caroline Atoll, based ona
plan published by the Solar Eclipse Expedition (Holden &

QuaTiEROUGI SSD ie wie ack parce st svsreteresc) Weber se awers-cnevclerer stor siatanalawawonavevanonadats

4 Caroline Atoll, as surveyed by John Arundel, 1883, and

still used as the standard hydrological chart...............

5 Caroline Atoll, as charted by the Solar Eclipse Party,

alisomini 1883 “(Holden & -Qual:trough 1884) 2228s. Sec Meo ees:

6 Caroline Atoll, by Bryan (1942), a modified version of

EheeSollapseelipsesPartyasemap ClSS3 er cc o.c 2 oe miei sce els cree

7 A map by Clapp & Sibley (1971la), based on Bryan’s 1942

Map (Rig).4 M6!) SSA AA RAS ARMAS NSS USNR MONG SI HALA ST Oe wT

8 Caroline Atoll: survey transects. The distance covered

Walsegl' 3.23: Skinla Maes SOS SSA INOS MOS Ltr G Ah a Pee SNE ob

9 Caroline Atoll: perimeter surveys. The distance covered

Weal SHeliQi G3 TKI coc Ratan amici seek ID GF 2 NE EEO SEL RGR RPE EY

10 Relationship between mean annual rainfall and
distribution of phosphate islands in the tropical Pacific

(reproduced *fromestoddart:s Scoftin 983): ene. S22 1a eee, 1s

11 Entire distribution map of rare and/or localized plants
on Caroline Atoll: Hibiscus tiliaceus, Lepidium
bidentatum, Pandanus tectorius, Phyllanthus amarus,
Psilotum nudum, Scaevola taccada, Sida fallax, Tacca
leontopetaloides, Thespesia populnea, Tribulus cistoides,

Ximeniiavamerticanals vandSpecie@swAss Bois. Me NS oo. OO:

12 + Transect distribution map of the fern Phymatosorus

SCOOPENGriaronaGaroline #ACOll ltete dea clnsrdlsttcrdds < telebel. seas cisions

13. ~Transect distribution map of the grass Lepturus repens on

Carolin eA COMM ty wek-ece tka yest ceA hadnt ck haere Aotchdcr chit Har ahobal hota oleae ces

ili

iv

14

13)

16
17

18

19

20

21

22
23
24
25

26

Ze

30

31

32

Entire distribution map of the coconut Cocos nucifera on
Caroline AtoV:. 0c. Fa letsas. Late LPS. SAAR ese ae ee eee

Transect distribution map of Laportea ruderalis on
Caroline ‘Atos. \s:.) 1 fob, ., aed Taare hI ale-: Sate area dover er eae

Transect distribution map of Achyranthes canescens.............

Transect distribution map of Boerhavia repens, Caroline
Ato reed src. iete: Siasetarhs, ticle GERAD Goisisssusisjoithe eiceee tel eer RR eee

Nee) enh es Coes HOO me Oo 6 oa G coo boon OD oS oocKc

AGO i, Sac Pidbssd,sysyscdeepocsesvoysusyevexS serous oxs.sy ayes ae bree ne

Transect and perimeter survey distribution map of Suriana
MALT EDMA S565, .2s, he IS SHAR oad. Ha AAs HAR, Bee

Transect distribution map of Ipomoea macrantha. Entire
distribution is shown for South, Island)... .).. 04.051). Shas ..ctnonnete

Entire distribution map of Cordia subcordata. .). ... 63.) .ests dquacke
Entire distribution map of Heliotropium anomalum...............
Entire distribution map of Tournefortia argentea...............

Transect distribution map of Morinda citrifolia. The
outlined area on Tridacna Island (northeast of South
Island) encloses Tournefortia-Morinda forest..............se0--

Evidence for the indigenous status of Morinda citrifolia
on Caroline Atoll: percentage cover on transects within
natural and anthropogenic «forestsis:.:<.. acsrsr. <teicsopelenel-. ced da el eee

Plant communities and amount of indigenous vegetation on
the different size classes of motus, as illustrated by
Vpie") di hagwaMms ds). ens daycsilverenavah gee. Qetstepeions ct kesh eet ek eae

Total numbers of plant subcommunities (upper graph) and

species (lower graph) in relation to motu area,

demonstrating plant succession on the different sized

motus encircling Caroline’s lagoon. Fine dots indicate

UPON VANES... «cscs 6 iee.sreiene wisi le eitelelet ole loheseuetel titlrettte telnet eee ana

Schematic profile through Arundel Islet, recovering from
disturbance over 60 years ago, showing natural herb mats,
Tournefortia scrub and forest, and 5 species of breeding
SC aD UGS i, 0.5, «<5: 54 8 5 8i5)8)e:5)'e.0y olen, 5) meilek sey ol area eee

i i i -

33

34

35

36

37

38
39

40

41

42

43

44

45

Maximum heights of Pisonia forests in relation to width
Os EHEAMORUSHE: “PERS henna hee hon Rhainhe Se Re EN oder bis Goble 3 102

Maximum canopy heights of Pisonia forests in relation to
MOE UAV Cae cae ess elle sv snsen at ala\-coelesle set's) "cite cilecasore.'s egies slice cose wieiel eve eis igie dis 8 siese 102

Schematic profile through Long Island, Transect 0.

Although Long Island has been formed in the recent past

by a merger of 5 smaller islets, this section of the

islet is very mature, containing natural herb mats,

Tournefortia scrub and forest, and tall Pisonia forest......... 103

Schematic profile through South Island, where >90% of the
land surface is covered with Cocos forests, primarily in

aedyAng: sitahete nese Be hoses SiG bisa hOB AOOENG sek kalars « 114
Nake Island: “vegetation and physiography..:...-.s.ee0.s%.sc000 121
Long Island: vegetation and physiography............cceceecees 124

Long Island: north-south transect showing division into

former islets, floristic composition, relative abundance

of plant species, degree of species overlap, and canopy

henGihitismaeet Aveo beeen ategee aerate emttnedts shake ds Sa eeniaees. 125

Long Island: east-west cross-section through Transect C,

a former interislet channel, showing floristic

composition, relative abundance of plant species, degree
Ofespeciestoverlape andsicanopy: NEIGMUS. ic J.teicte wiser 01. ois lorels cheiefate « 126

Long Island: east-west cross-section through Transect 8,

which passes through mature interior Pisonia forest of

largest of Long’s coalesced motus. Data include floristic
composition, relative abundance of plant species, degree

of Species: overlap, fandicanopy! NetghtSisc).3.... 65. ere einer 126

Vegetation and physiography of Windward Islet no. 1:
BOMSUNPBieGerliSiteitperscuvevettets cis s.cncts crerstctsiciane: certo aiaile Glvteyarecsiagneueieveneiers 131

Vegetation and physiography of Windward Islets nos. 2, 3
and 4: Windward and Crescent Islets, and Motu Atibu
GiConalp Rubbilesisiletsy).£ eek nvecd pete. Behe el. tehie. te tects: bchah:. 133

Vegetation and physiography of Windward Islets nos. 5
through 9: North Pig, Pig, Skull, North Brothers, and
BROENERSHISIGtSE Beppe tpn! hy, ltt ccdeteratitetiste fails eo! altekorte alelotstel steers. slhtieietins « 135

Pig Islet: east-west cross-section through center of

islet. Data includes floristic composition, relative

abundance of plant species, degree of species overlap and

GANOP YMG Strsres is, cyetercs cya eciciisracatrenssovels loverencce ci citslevelie iste s:roleue) Gel svelare 137

vi

46

47

48

49

50
51

52

53

54

55

56

Brothers Islet: east-west cross-section through center

of islet. Data includes floristic composition, relative
abundance of plant species, degree of species overlap and
canopy” heiqhts:.i5% 13.. ke VE... Ce Oe ARES Hee eee

Vegetation and physiography of Windward Islets nos. 10
through 12: Noddy Rock, North Arundel, and Arundel
Islets A889 288281 284. RIE Ree RRR REE EE ease

Vegetation and physiography of Windward Islet no. 13:
Tridacna’ Tslet 228842. ..084 BRE. Rk RtR% AR REA Se ee ee

Tridacna Islet: east-west cross-section through lower

center of motu. Data includes floristic composition,

relative abundance of plant species, degree of species

OVE ap and CAaNOPy ME TGHES 4.6.50. << ccvercienexonensyevcueresore onereverer One pohmeneaeaaan

South Island: ‘vegetation and physiography sc. :)1. </)s\ctelels olcldenne

South Island: distribution and abundance of plant

species along Transect 2, which runs at an angle of 60°

from the lagoon to the south shore through the western

center of the islet. Data includes floristic

composition, relative abundance of plant species, degree

of species overlap and canopy heights. .«......«.<. «+. ss osc

Vegetation and physiography of the 7 South Nake Islets:
Pandanus, Danger, Booby, Coral, and Lone Palm Islets,

Motu Kota ("Red-footed Booby Islet"), and Motu Mouakena
("Masked ‘Booby islet ras. af. See telks Stelel al shtel Mateo catatonia

Vegetation and physiography of Central Leeward Islet
no. 13) Motu! Mannikiba("Seabirdvislet™!.cn8 2. 24 ie eee

Vegetation and physiography of the Central Leeward Islets
nos. 2 through 4: Blackfin Islet, Motu Matawa ("Fairy
Tern Islet™),. ‘and ‘Emerald stie. ..0. VOR Ne, SO

Vegetation and physiography of the Central Leeward Islets

nos. 5 through 11: Shark and Scarlet Crab Islets, Motu
Nautonga ("Sea Cucumber Islet"), Azure Isle, Reef-flat,

Bird andi \Fishbal.| -Isiletisi. isc sic ite crchetere fetishes Pole ach taeda t-te

Fishball Islet (no. 11, Central Leewards): east-west
cross-section through the center of this young motu,

which exhibits early stages of geological and biological
evolution. Data includes floristic composition, relative
abundance of plant species, degree of species overlap and
Canopy: heights 2... SSS RD. MANA SDS Ae

57

Vil

Vegetation and physiography of the 5 Southern Leeward

Islets: Motus Raurau ("Blue-gray Noddy Islet"), Eitei
("Frigatebird Islet"), Pisonia Islet, Kimoa ("Rat

rset) andsAnasAnaw(tAnnerswisilety ees 222 Se beekes cose luk.. 168

List of Plates

Page
A-dawnawiewrote Carol ineswlOSSin ce sete ee Beads Skee lek neue 186
lihewmaineeleawinguon:Southwisiand,) 1883.25. tsk eee ccs eee os 186
One of 3 European-style houses built on Caroline, 1883......... 187
Anivanvistssynender ingioheCaroline, 1883s ce ce eesek. oe. eds. 187
Mapnofethesesettiement. South sland, 1883. 2242. fel. ste eee 188
WO MAG OOM MVAEWSE ABCOMTUNY) 1AGO/. ie, o's :aila crepes o's sie ereleteliotes's) suererecele’siaye ahs 188
Motu Ana-Ana, in 1883 essentially identical to today........... 189
A Tournefortia tree along South Island’s lagoon edge........... 189
The junior author in Ipomoea macrantha thicket, South
Milan det sh... SH Sno ale chee che SE MRA RRR ell. Snel BAER 6 189
Rat, Citycebasescamp,:, Longilsiand: teen ee: een meee). caret. 190
Bliack=tippedtineefi shank, b Carol inelis® lagoon: )...ii3n naee ose «. 190
Laterally continuous upraised older reefs and wide reef
filiats:. Southwest, NakenMsil andes) cca? Wee Uses pbc ne streamer 191
Floating a small boat across the reef flats from
MENL GAN CEun CO NANGIINGsar. crstsctte ccltiee celawie ee OeR OR AR e Lue.’ 191
Beach crest, rubble, seaward "lagoon" and reef flats off
SOUCNe AS tH INAkey Tisai thas wictetevcie tate kere tcuels! akousiore Wtele\giee li oot ape lots oe 192
Extensive lagoon reef flats south of Arundel Islet............. 192
An incipient motu south of Motu Mouakena..................0000- 193
Ridgeshofconalinubbille:,. Nakel sis lama: sytem: ve stcre <letote ete fr clolsvedet® «:« 193
Channel between the 2 northern islets, Long and Nake........... 194
Noddy tRocksuwindwardi vee fi fil aitsvoys. S05. cE. Eo Se Blows ee 194

Windward beach with flotsam and jetsam, Long Island............ 195

Vill

20

21

22

23

24

25

26

27

28

29
30
31
32

33

34

35
36
37
38

39
40

Conglomerate platform, Suriana and Russian vessel
Akademik Korolev, northwest point, South Island................

A large coconut crab shelters in a subterranean cavity in
TNE FOO... 5:5.0eec0s so ie ie oe wale sels e iellele Srotloiei's foie leteltele acai cite lteiateenaaeaneanan

Sandy Inlet, with foraging Bristle-thighed Curlews,
extends its mudflat. into. Nakes..........2s00.005 snes cee ee eee

South Island’s Cocos plantation, looking west along the
VaQOON. 2.02. obese sss ose eee csee eee sob bese ee whe ee SRR RC Renn

Submerged reefs adjacent to: Emerald Isile..2.228 (Re... ree
A highly productive, cross-lagoon reef of coral and clam
shells (Acropora, Tridacna) joins Tridacna Islet with

Motu. KiMOa oi)... 6: oie 26% sie e's Wiese winisiie sect w © oe nie (0 sera ener els telnet ante
Kou (Cordia subcordata) forest, Pig Islet........5.0 00. sccneneee

Sand, silt, rubble and hardpan mingle on Long Island’s
Tagoon’ edgeis 0/52). 50% 5 SRS ARN 2 8 CSR ORE RRR eee

Caroline’s sandiest beach lines the lagoon shore of Shark

1) Se a nes ree ee EC So SS oS:
Sheltered. bay,. Brothers: Tsllet... oc... che 085 ow oe ele ene ehetee een
Narrow lagoon: beach,’ BlackfiniIsletecs. . (cok 22S" REk eee eee
Recent sand additions to South Island’s northeast point........

An old interislet channel, in 1988 filling in with herbs,
Tournefortia. scrub. and Cocos. (Long Island): ..2ew0ic2 . 20m eee

The same area as Pl. 32 in March 1990, 2 weeks after
CYCTONTC: weathers .6..5% 00.05 cle wiawew aie ele pes otek hee aan

A large clearing within dying Cocos-Ipomoea forest, South
TST ANG ki wee oe ook eee Ws whe lee ee leitele Nie olelale lee ole eae nana

Pandanus. forest, south’ Nake.4. ...6.... 2 eee ReRE ee eee
The north end wall of an ancient Tuamotuan marae, Nake.........
Mixed forest wiith Cocos, Nake Tsilandt). 22. sec eee iceeeee

Orange, scarlet and green phalanges of Pandanus, foraged
by Coenobita, rest on a clump of Portulaca: .1.\ <1 ='rtteieiee

Inner-edge of lagoon, South Island; 1988: ......2 20.02). see
Inner edge of lagoon, South Island, 1965. .....\.. 2.3250. meee

4]

42

43
44
45
46

47

48

49
50

51

52

53
54
55

56

57

58
59
60

61

Achyranthes canescens in a clearing adjacent to Pisonia
ORCS wa pal Gp SUC URP Ele tevsyecie1syercezeueeieLeusieislegsitgscess ceaclie sols. eneile e..eve, oie iesG 10.4

Boerhavia fruits on feathers and bill of a Great
APC IENBSY 8) TT et ko ch A Gol cea eed NO el RO eRe Coit ARP INL Me aE a a

Inside a mature Pisonia grandis forest, Nake Island............
Fringe of Suriana, northeast point, South Island...............
Well developedenatunalniherb) Mati ns cisetsc tee et eiciciels ccs ceed e lee eicte ons

Heliotropium anomalum with tropicbird skull (first
evidenceon this Species) haSkuilil, eset: ii5.5 fertile wots loveless ue © eysce

Tournefortia scrub, fringed by a natural herb mat, and
OCCUD Eds DymagGollOny Ore SOOtYPTOrnSe ioe sce. ries 2 oebohavs oisiaiete sos auele

Secondary Tournefortia-Morinda forest with nesting Brown
Noddilesho midaenasrsieterccscr ste mero ce ee aie etlccctee eet meee g

Skul] Islet looking east to the windward reef...............06-

Mature Pisonia grandis canopy with incubating Black
Noddiiestandaar White m@mennanP iG TSl@tescsc.cs6< coc cuetsies cies ceguere ss

Azure Isle, an example of a motu containing a single
AUS OND Ma ENCE Cra cyeta cele recuse ecira eine calicieraeveievetcreieneite overs) suese alniels silo: dua eee gat

Caroline’s single clearing with Tahitian-style huts, Motu

NIDENSI ALINE esate Patani Beas ell ull Bd hea Pei ol Ut al Sabre alm De Mi der one Ra pt
PHlesmonhemibLOUS.shdaVingS--COCONUL ChaD SIGN:c-s cscs ote. +.
Beachnock s4OnG sisilandaeeas ka et ee eek, Mtl tee trace ae emai:

Windward Islets nos. 5-9: North Pig, Pig, Skull, North
Brothensmand=Brothensprcces seri crea ene ae a ean

Arundel Islet (foreground), looking south-southwest
EICTOSS WRICGECHE ISTE OcSOU JESTIETGlasiGcaitcio's Gewmrcitel ore God c

Pandan usmlsiliety (COntOrs) ici avaroteapotesaie: -xeleytearorecedepes i evsueieuone snsieysiciasoly eyes
DanGeremeltsinGicepacters cesta atcle a caere ore ccater reenter eee ee MS TORS ates

South Nake Islets nos. 3-6: Booby, Coral, Lone Palm, and

Most i MOU AK eM alate. terspe here sarcjrenone texenohonehencfoke exe kake d= sh-Nak nek pee
North vend, Motu ManniiKibals:(. rr. -peretcleiars ieeterate Ot eee
Motu Mannikiba, looking east along Transect 1...............
Motu Mannikiba, looking east along Transect 2...............
Mixed Pandanus-Tournefortia forest, Emerald Isle............
Emerald: TSV@.. <0is.c: 60:00. siee.06 0 5 0) dhguete votekenle sfeloqaietal uence at eee
Shark. USM SU wpotsgepcrspsvokoresajons despaicdoaskensnbods (otehehsne ogc exskegsaioceke tel aeons

Southern Leeward Islets nos. 1-5: Raurau, Eitei,

PiSONT as sKAMOa ANG HAN ASAN aha seus ous ayetorevoxsienchoychor<toiekoychcuolot oleae oNeneneneae

View of Motu Kimoa from) Pisoniiia, Isilet. «02-0. e cm ere

Motu Ana-Ana in 1988. Compare with Plate 7, taken in

The "blind passage," looking west from its inner end to

ENE, VAQOON «101.5: sf2%ersnexchstenogs joxenehis iootshauerepensnehexoss:opehSdsyshogsuecen shaaysReleRek ieee

List of Tables

Plants reported from Caroline Atoll but considered to be

transient or extinct, members) of: the Fllorvalc<cc.-.-1te cree acters

Vascular flora of Caroline Atoll: relative abundance of
each species within the major ecosystems, with data on

Seabird: Ubi ZaE VOM. c crsysssyeroncreveic ors a oro ce Saenoncten cine

PCW sa tacassoaiee veresh oud feseerrm sr ac ceva.ce, ou oure ho Haereu ete waSes Cite ORS oR Onna

Sizes of Pacific atoll floras, with emphasis on the

percentages of Indigenous: Plant Sepsis. «ce obanchapchsrevageucleuspeu-tsl eae

The distribution of plant communities, together with the
numbers of plant species on the motus of Caroline Atoll,

illustrating seral stages of plant succession. ..... 2...

Distribution and abundance of plant species in relation

to islet area and the primary mode of seed dispersal........

Species-area relationships of 6 Pacific islands with

entirely, Andi GenOus: FOV A;.. ofacetassps,s: 0 sotspet elaiain seacel-\e letter

Area of plant communities on the islets of Caroline Atol]

39

66

70

71

88
91

10

il

12

13

14

Widths of pioneer herb mats on seaward- and lagoon-facing

SORES sae Garonne vA isles rise, ceissenate lero pe isas evoeseetetst tare ors; Shaciicve acess

Stature and extent of Tournefortia in the major habitats

Ofee Garcolleine@m Aoi heieeetraiar, enone metre mn i uch eee uy somtte chac Us A

Distribution of well-developed (210 m height) Pisonia

forests on the motus of Caroline Atoll......................

Area and dimensions of Pisonia grandis on Vostok, Flint,

andinS sisletsnoh Garonne SALONA. Hac scale ie Sichsle ote Sense hlovaie sens

Number of trees and areas planted in Cocos on Caroline’s
islets during the major planting era (1916-1920), also

ShowingeremnantsGocos datadfon l988h. 5. cs e.c Nasiees. eos

Species diversity in Pisonia forests of decreasing

MacUIaity. <cagOlniMes Atoll te cis sie. Sites, Sosy. cpeeleac ees olersaiers oceverete « o%

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PARTI. HISTORY, PHYSIOGRAPHY, BOTANY, AND
ISLET DESCRIPTIONS

BY
ANGELA K. KEPLER' AND CAMERON B. KEPLER’

ABSTRACT

Caroline Atoll (Frontispiece) is situated at 10°00’S latitude and
150°13’W longitude in the south-central Pacific Ocean. Caroline is the
southeasternmost of the Southern Line Islands, a group of 3 islands
which also includes Vostok and Flint, lying 230 km to its west and
southwest, respectively. Although archaeologically and geographically
within Polynesia, Caroline is owned by the Republic of Kiribati
(formerly Gilbert Islands).

Caroline, 9.7 km long, 2.3 km wide at its widest point, and
26.9 km in circumference, is a crescentic coral ring with 39 islets
(motus) centered on a continuous reef enclosing a relatively shallow
lagoon. Its total land area above high water is 399 ha., with motus
ranging in size from 0.02 to 107.5 ha. Motus extend along 55% of the
reef perimeter. The closed lagoon, rich in marine life, contains a maze
of patch reefs and impeccably clear water.

The atoll, uninhabited, was "discovered" by de Quiros in 1606.
Although traces of an ancient Tuamotuan culture still exist, the atoll
apparently never supported a long-term permanent population and has been
less affected by man than most Pacific islands. Its European history
includes guano export, a multinational expedition to observe a solar
eclipse, and copra production. It has been uninhabited since the early
1930s (a factor contributing to its relatively undisturbed ecology),
except for the presence of one family from 1987 to 199i. The primary
factors responsible for its lack of permanent settlement are remoteness,
apparent absence of usable ground water, repeated failure of its coconut
plantations (diseases, destruction by coconut crabs, rats and seabirds,
smothering by vines), absence of a passage into the lagoon, and a
paucity of safe boat anchorages.

Until the 1988 USSR/USA expedition, only an 1883 chart was
available, which named 7 islets. We drafted an accurate map based on
field work and recent aerial photographs, naming 32 previously unnamed
islets, 4 islet groups and an inlet. During 8 days’ intensive field
work, we surveyed 38 islets, walking 33 km in systematic cross-islet
transects and around islet perimeters. This paper presents much new
data on Caroline from 3 visits in 1988 and 1990, and attempts to
summarize, expand and synthesize previous information in the light of
new findings.

1 400 Snapfinger Drive, Athens, Georgia, 30605, USA
2 National Biological Survey, School of Forest Resources
The University of Georgia, Athens, Georgia, 30602, USA

Manuscript received 8 April 1991; revised 26 October 1993

Soils, principally of coral, mollusc, and algal origin, are
categorized into 5 types, from barren coral rubble to rubble mixed with
humus and guano. Caroline provides an excellent example of soil
development through different age and size classes of motus.

Preliminary descriptions of the atoll’s reef, motu and lagoon
morphology are given, including the background geological setting of the
Line Islands. Physiographic features include inland upraised reefs
(feo) and deep sand deposits, coalesced islets, exposed older reefs,
lithified beachrock, a conglomerate platform, a "perched lagoon," a
nonfunctional hoa, and changes in motu size and shape during the past
century.

Caroline’s lush vegetation supports 26 species of plants organized
into 7 plant communities, 6 natural and one anthropogenic. The atoll-
wide distribution of each plant species is mapped. Plant species
(including 5 new island records) and communities are detailed,
emphasizing the atoll’s past history. The atoll’s insular flora,
although impoverished due to its geographical location, is 89%
indigenous (possibly 92%), an extremely high figure for anywhere in the
world. Although Caroline’s motus are covered with extensive tracts of
indigenous plant communities, the Pisonia grandis forests, up to 21 m
high and covering 22% of the woodlands, are particularly notable as they
constitute some of the best groves left in the Pacific. Pisoniza is
treated in detail, including data on rapid recovery and growth rates
during the past 70-odd years. Tournefortia argentea (43% of the
woodlands) is abundant, and Cordia subcordata, becoming quite rare
elsewhere, occurs on 21 motus (54%). Cocos is present, but only
dominates one islet; 22 islets harbor wholly indigenous vegetation.

Motus of varied age and size classes provide excellent examples of
substrate and vegetation development, accompanied by an increasing
diversity of bird life. On account of its relatively low human
disturbance and rapid forest recovery to a more natural state,
especially since 1920, Caroline is one of the few Pacific islands that
is truly an "outdoor ecological laboratory": many motus have recovered
so remarkably they are almost indistinguishable from those that have
remained pristine, while others are in different stages of recovery
resulting from varied management (or non-management) practices. Exotic
plant species are very few: ancient Polynesian-introduced (Cocos,
possibly Pandanus), recent Polynesian-introduced (Hibiscus tiliaceus,
Thespesia populnea, Tacca leontopetaloides, Ximenia americana), 20th
century exotics (Phyllanthus amarus, in one small area). Some garden
species, cultivated from 1987-1990 have an uncertain future and are not
treated as part of the atoll’s viable flora.

An analysis of ecological succession on motus of increasing size
reveals that by the time a motu reaches 0.8 ha in size, all the natural
plant communities, most plant species, and most species of seabirds are
present. This is in striking contrast to species-area relationships on
inhabited atolls with more introduced plant species, for example
Kapingamarangi.

Each motu is individually mapped and the main physiographic
features, known history, vegetation patterns (including species-area
relationships), seabirds, and miscellaneous biota (coconut crabs, rats,
lizards) are detailed. Appendices provide weather data and describe the
practicalities of anchoring boats, landing, and moving small boats
around the reefs and lagoon.

Permanent protection of Caroline is currently underway as The
Nature Conservancy of Hawaii negotiates with the government of Kiribati
for a Southern Line Islands Wildlife Preserve, which includes Caroline,
Vostok and Flint.

A. INTRODUCTION

Caroline Atoll! (Frontispiece; Figs. 1, 2) is a small, low coral
island situated at 10°00’S latitude, 150°13’W longitude in the south-
central Pacific Ocean. Its maximum height is less than 3 m above MSL.
It lies 2,800 km south of Hawaii, 830 km north of Tahiti, and 1,030 km
west of the Marquesas Islands. Its nearest neighbors are Vostok and
Flint, 230 km to the west and southwest, respectively.

Recent measurements by the ICBP 1990 Line and Phoenix Islands
Expedition, using a compact satellite navigation computer "Magellan" NAV
1000, indicate that the atoll lies one nautical mile east of its
previously charted longitude position, 150°14’W. Its range of
coordinates are: 09°54’ to 10°01’S latitude, 150°12’ to 150°14’ W
longitude. The given coordinates, 10°00’S and 150°13’W, intersect in
the lower lagoon just west of the "blind passage."

Although archaeologically and geographically within Polynesia, the
Line Group was uninhabited when discovered by Europeans; its islands
were variously claimed by the United States and England. With the
exception of U.S.-owned Jarvis, Palmyra, Johnston and Kingman Reef, all
are now governed by the Republic of Kiribati (formerly Gilbert Islands).

Caroline, 9.7 km long by 2.3 km wide at its widest point, is a
crescentic coral ring 26.9 km in circumference. Composed of 39 islets
(Fig. 2, Table 9) and three incipient islets, it is centered on a
continuous reef flat, submerged at high tide, that encloses a relatively
shallow lagoon. Most are well-wooded, but 4 tiny ones, less than 0.1 ha
in size, are scarcely more than coral rubble piled on the reef,
supporting sparse patches of Journefortia and Heliotropium. One islet,
Noddy Rock (Pl. 18), is a vestige of a former reef segment. The total
land area above high water is 399 ha. One of the oceanic islands
contributing to Darwin’s theory of atoll formation (Darwin 1842), its

Icaroline Atoll is neither physically, geographically, nor politically
associated with the Caroline Islands, now part of the Federated States of
Micronesia, more than 6,000 km to the northwest. Because of this confusion,
we use the name "Caroline Atoll" instead of "Caroline Island."

tropic of Cancer

! b |

170° x ) 150 140 W
a

® 20°

200 400 4600 S!-MI.

ae HAWAII

(Oe 2) eee
(0) 320 640 960 KM.

SCALE AT EQUATOR

1o.—
_.KINGMAN REEF
. PALMYRA ATOLL
< . WASHINGTON IS
Z| * FANNING ISS\
i4 Bk
| a KIRITIMATI >
! < |ICHRISTMAS !S.]\_
| ne
equator a a a i °°
“JARVIS IS Sq
2 eee ee ee ee se ee aa cs
a
Dae
PHOEN! x o
ta KIRIBATI
MA hie |
for LDEN.
GROUP = O . FILIPPO REEF
STARBUCKIS. . £xUI- x === >= Soo cee
= Se Se SS ne aS ce os le
TOKELAU | Se A j v :
‘a y een “aa | e-
i i VOSTOK IS. ; tvs 161
= Vs CAROLINE u al
3 Es AT
! | FLINT IS ATOWL FRENCH
AMERICAN! WL === = -------
SAMOA | 7 A POLYNESIA
td |
mere i oe Ho, :
WESTERN ==! COOK ISLANDS} reais | eae
samoa | ©O0 Fai ape tah “elena” ty
| “0, , eg TAMITI iat bes 4 % :
! i) me ‘a
NIUE ; ! /@ ce) > =
“a 8 | oLenBaraisteny “a 2 ee
= \ | @20_
! | °
' RAROQTONGA
' Be
_ tropic of Capricorn — |

t !

Figure 1. Line Islands: geographic location in the Pacific Ocean.
After map by the Hawaii Geographic Society (1981).

NAKE ISLAND
PANDANUS ISLET
DANGER ISLET—___
SOUTH NAKE BOOBY ISLET
CORAL ISLET
ISLETS LONE PALM ISLET -
MOTU KOTA
MOTU MOUAKENA
LONG ISLAND
lagoon
MOTU MANNIKIBA
t BLACKFIN ISLET BO'SUN BIRD ISLET
CEN RAL MOTU MATAWA
LEEWARD EMERALD ISLE WINDWARD ISLET
ISLETS SHARK ISLET
SCARLET CRAB ISLET CRESCENT ISLET
MOTU NAUTONGA MOTU ATIBU
AZURE ISLE WINDWARD
REEF-FLAT ISLET NORTH PIG ISLET
BIRD ISLET ISLETS
FISHBALL ISLET PIG ISLET
SKULL ISLET
NORTH BROTHERS ISLET
BROTHERS ISLET
SOUTHERN

MOTU RAURAU

LEEWARD MOTU EITE!

PISONIA ISLET
ISLETS MOTU KIMOA
MOTU ANA-ANA

NODDY ROCK
NORTH ARUNDEL ISLET
ARUNDEL ISLET

TRIDACNA ISLET

blind channel

SOUTH ISLAND

CAROLINE ATOLL

SCALE 1:34,000

300 ° soo 1000 2000
meters

Figure 2.
islets.
1986).

Caroline Atoll, Republic of Kiribati, with newly-named
Based on photos by the Royal New Zealand Air Force (RNZAF

6

geology, soils, climate, and vegetation are typical of low latitude
atolls and are relatively unmodified by man. :

This paper presents much new data on Caroline from 3 visits in
1988 (September) and 1990 (March, May) and attempts to summarize, expand
and synthesize widely scattered information in the light of new
findings. The only significant previous biological information on
Caroline was from the Smithsonian’s Pacific Island Biological Survey
Program’s 2-day visit in 1965 (Clapp & Sibley 1971).

B. HISTORY OF CAROLINE ATOLL

Pre-European History: Tuamotuan Period

Centuries before Europeans encountered Caroline, the atoll was
inhabited by Polynesians. No oral traditions of this occupation are
known, but evidence of former habitation was evident when de Quiros
found the atoll in 1606. He noted "an old canoe, lying on her side,"
and a small grove of coconuts planted on South Island (Bennett 1840,
Markham 1904).

No further clues were unearthed until Messrs. Brown, Brothers, and
Arundel exposed about 50 ancient Polynesian sites in the 1870s while
digging for guano (Holden 1884, Arundel 1890). Polynesian inhabitants
at that time called them "marai" (marae). Arundel photographed and drew
plans of them (Fig. 3): depicted are a platform of coral and
conglomerate rock, surrounded by 10 smaller slabs resembling
gravestones, all arranged in a rectangular plan. Although the largest 2
were marked as "graves" on Arundel’s 1883 map (Fig. 4), no bones, ashes
or human remains were found. Their findings were later identified as
Tuamotuan marae (Emory 1947). Marae, according to ancient belief,
"bound the ancestral spirits and gods of the kindred to the land,
putting it under their eternal guardianship" (Emory 1947). The largest
marae was on northwest Nake Island, and a smaller one was found near the
southern tip of Long Island. Both locations conform to such
prerequisites for building marae as nearby shorelines and birds (see
Sect. H.1), which Tuamotuans believed housed divine spirits (Emory 1947,
p. 123). Although AKK, G. Wragg and R. Falconer located, photographed
(Pl. 36), and measured these in 1990 (A. Kepler 1990d), no field work by
archaeologists has been conducted. Our data and numerous photographs
have been sent to Dr. Jeff Irwin, Auckland University, New Zealand.

Post-European History: 17th to 19th Centuries

On 21 February 1606, the Portuguese explorer Pedro Fernandez
de Quiros, employed by Spain, "discovered" Caroline Atoll (Markham 1904,
Stevens & Barwick 1930), naming it San Bernardo. Despite its
remoteness, Caroline was encountered early in Pacific history, long
before Tahiti, Rarotonga, and Hawaii. This is possibly due to its
location, for early navigators tended to sail due west from South

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8

America along lines of latitude, and 10°S was an obvious choice.

De Quiros, the last adventurer in the Spanish age of discovery, was
leading his second major trans-Pacific expedition with 3 ships and 150
men obsessed with finding the fabled "Terra Australis Incognito." The
descriptions of Caroline by his crew, although at variance with one
another, still apply today (Pl. 1). Their first at-sea impression was
that it was "divided into four or five hummocks, and all the rest
submerged. Its circumference appeared to be ten leagues" (Markham
1904). After landing, they found that

"There was a great number of fish inshore, and, owing to the
water being very shallow, they were killed with swords and
poles. There were great numbers of lobster and crawfish, and
other kinds of marine animals. They found a great quantity of
cocoa-nuts in a heap at the foot of the palm trees, many
large, and of different sizes. There were a great quantity of
sea birds of several kinds, and so importunate that they
seemed to want to attack the men. We took plenty of all these
things...It seemed to the Captain that on an island where
there are so many trees there could not fail to be water"
(Markham 1904).

Fresh water was crucial to de Quiros and his crew, who were
suffering from lack of food and water. Despite their efforts, however,
they failed to obtain water. Disappointed and lacking energy, they
continued their voyage the following morning. Their demoralized state
may explain one statement that Caroline "consisted of twenty-two islets,
uninhabited and without water, trees or scrub for wood."

In 1795, Capt. W. R. Broughton, on the British sloop Providence,
rediscovered and named the atoll while voyaging from Tahiti to Hawaii
(Broughton 1804):

"The southern extremity was the highest part, covered with
trees, most probably cocoa-nut from their appearance, as they
stood in detached clumps along the shore. The island...
appeared to be low, and covered with trees, and if I am right
in its estimated distance, its length will be about five miles
in a north and south direction. I named it Carolina Island in
compliment to the daughter of Sir P. Stephens of the
Admiralty."

Because early navigation techniques and communication were far
less sophisticated than today, especially with regards to longitude,
Caroline was sighted or "discovered" by several more explorers who were
unsure of its identity. By 1821 the atoll had amassed a collection of
coordinates and names: San Bernardo, Island of Fish, Thornton, Hurst’s,
Clark’s, Independence, and Carolina (which later became Caroline). Some
navigators equated Caroline with an island named "San Bernardo" by the
Spanish explorer Mendana in 1595. "San Bernardo" has recently been
verified as Puka-Puka, northern Cook Islands (Maude 1968).

The best early description of the atoll comes from an 1835 visit
by F. D. Bennett, who was reasonably well versed in natural history
(Bennett 1840). He noted that the islets then, as now, were "covered
with verdure...surprisingly luxuriant, when compared to the arid soil it
covers." Although Bennett had visited many atolls, he was particularly
impressed with the quality of Caroline’s coral reefs. His party
observed "rats of a red-brown color" and various birds but no reptiles
(Pt. II). Although he discusses "land lobsters (Coenobita species)," no
mention is made of coconut crabs (Birgus Jatro): perhaps the latter
were lumped with the former.

First Occupation: The existence of 2 small coconut groves on Caroline
prompted 2 British entrepreneurs, representing the Tahitian firm Collie
and Lucett, to establish a stock raising venture there in 1846. This
first known settlement was located adjacent to the main coconut grove on
the northwest peninsula of South Island; a smaller grove evidently
existed "on the south-south-west side" of the same island (Lucett 1851).
Tahitian laborers tended pigs, hens, turkeys, and grew many food plants,
including pumpkins and melons. They dried and salted fish, planted
coconuts, and extracted coconut oil (Maude 1942, Garnett 1983), and were
evidently still there in May 1852 (Ellsworth 1990).

Political Annexation: Though inhabited in prehistory by Tuamotuans,
officially "discovered" by the Spanish, and visited by British, French,
and American ships, it took centuries for Caroline to acquire a
political identity. It was formally annexed to Britain by Captain
Nares, R. N., who arrived in the H.M.S. Reindeer in 1868, finding 27
residents.

Caroline was under the control of various merchants in the late
19th century: Lionel Brown, Captain Brothers, and later John Arundel, a
well-known businessman, trader, and guano merchant in the Pacific.
Arundel’s 1883 map (Fig. 4) of Caroline is the only reasonably correct
chart published prior to this paper.

The Guano Era

Though bonded under the American Guano Act in 1860, no phosphate
was dug on Caroline until Arundel was granted a 7-year license in 1874.
A few months earlier, a set of moorings were laid off the lee side of
South Island, allowing ships of up to 1,000 tons to lie safely during
trade wind weather. Guano was the only successful business venture at
Caroline: approximately 10,000 tons were shipped to California and
Australia between 1873 and 1895, when supplies became exhausted (Young
ca. 1922). We have no direct information on which islands were mined
except South and Nake. We strongly suspect that Tridacna, Arundel,
Mannikiba, and perhaps others also yielded guano. The tonnage extracted
from Caroline was small compared to that from dry, barren, more
northerly islands with little vegetation, where populations of Sooty
Terns, an important guano species (Hague 1862), number in the millions.

10
Solar Eclipse Expedition

In 1883 Caroline received international publicity when astronomers
calculated that it lay directly under the path of a pending solar
eclipse. Three parties of astronomers (American, British, French)
arrived in the U.S.S. Hartford and set up camp on South Island, making
detailed observations of this celestial event (Pl. 2). At that time
Caroline was more famous, and housed more people, than before or since:
7 "natives," scientists and crewmen totalled 51 occupants. Legacies
from former inhabitants included 3 houses (Pl. 3), 2 sheds, 3 huts on
smaller motus, nautical flotsam and jetsam, and 2 shallow wells. To
this they added tents, observatory frames, a marble slab, flagpole, and
brick "piers" for their telescopes, most of which remained temporarily
as technological litter.

This expedition (Dixon 1884, Holden 1884, Holden & Qualtrough
1884, Trelease 1884, Young 1884) also marked the first attempt to
describe the topography, climate, flora, and fauna of the atoll.
Drawings included an artist’s rendering of Caroline, "settlement" map
(P1. 5) and views along South Island’s lagoon shore (Pls. 6, 7). An
atoll map was drafted but is highly inaccurate. Their biological
observations were sketchy (Dixon 1884, Butler & Strecker 1884). For
example, Dixon, the zoologist, listed such organisms as "shrimp,"
"hermit crabs," "gnat." As with Bennett, there was no mention of
coconut crabs, even though they were evidently abundant on South Island
in 1910 (Young ca. 1922).

The Late 19th and 20th Centuries

In 1875, C. D. Voy, a naturalist from California, visited
Caroline, collecting molluscs (Pilsbry & Vanatta 1905a, b) and fish
(Fowler 1901).

In 1885, Arundel began to clear land and plant coconuts, but his
planned copra industry was unsuccessful. In 1897 he sold his business
to the Pacific Islands Company, Ltd., which also failed. The
plantations suffered from disease and poor vitality, coconut crabs
chewed on seedlings, seabirds destroyed the developing nuts, Ipomoea
vines strangled young trees, and populations of Polynesian rats
apparently exploded, causing further damage to both intact nuts and
drying copra (Young ca. 1922, Maude ca. 1938, 1942). By 1904, when the
H.M.S. Icarus visited Caroline, only 6 Polynesians lived there. A few
months later they were repatriated to Niue, and Caroline remained
uninhabited until 1916, when a new effort was made to develop the
coconut plantation by Messrs. S. R. Maxwell and Co., Ltd.

During the uninhabited years, South Island’s vegetation and
wildlife began to recover from the earlier forest felling (Pls. 2-6).
When Mr. J. L. Young, then managing director for S. R. Maxwell and Co.,
Ltd. (Young ca. 1922), visited the atoll in July 1910, he described it
as a wilderness, teeming with Sooty Terns, fish and coconut crabs:

ll

"The ground was covered with nests of seabirds which
latter rose like a cloud when disturbed: the noise of their
shrieking was so great that one had to shout to enable oneself
to be heard by his companions. Hundreds of great Coconut
Crabs were seen: 40 large ones were caught by the crew of the
schooner in an hour. The reef and the lagoon swarmed with
fish and small sharks."

From 1916 to 1929, Caroline was altered more than before or since.
All the available land on South was deforested to make room for
thousands of palms, and laborers demolished huge numbers of coconut
crabs and seabirds (Young ca. 1922). In addition, coconuts were planted
on all of the main windward islets, southern Nake, and on Mannikiba.
(The windward islets recovered their forests remarkably quickly, see
Sect. G). Plantation workers in great part lived off the land, feasting
on fresh fish, seabirds, seabird eggs, turtles, and coconut crabs. Most
of the leeward islets escaped alteration for plantations.

Copra exports averaged around 14 tons per year from 1929 to 1934,
after which the company ran into debt. Concurrently, the French
government forbade further recruitment from Tahiti; by 1936 only a few
families were left (N.1.D. 1943). In 1941 the atoll carried a price tag
of 600 English pounds (Maude, pers. comm.), but was never purchased.

Occupation leases for Caroline were cancelled in 1943, after which
the British Western Pacific High Commission repossessed it (Maude 1953).
However, new "queen’s leases" were granted to M. P. A. Bainbridge of
Papeete, Tahiti, 1951-1964 (Nicholson & Douglas 1969), then to
Capt. Omer Darr of Moorea, French Polynesia, from 1964 to 1989. When
the British granted independence to the Gilbert and Ellice Islands in
1979, a new country, the Republic of Kiribati, assumed ownership of
Caroline, along with most of the Line and Phoenix Islands.

Apart from occasional parties of Tahitians cutting copra and a
shipwrecked sailor in the early 1980s, the atoll remained uninhabited
for over 50 years. During this time Caroline’s vegetation and wildlife
recovered to such an extent that, were it not for unpublished
manuscripts (Maude ca. 1938, ca. 1942, and no date, Young ca. 1922) and
comparisons with Flint and Vostok (St. John & Fosberg 1937, A. Kepler
1990b-d, and Kepler, in prep.), we would have been unaware of the extent
of previous human interference or of the rapidity of forest recovery
(the fact that 60% of Caroline’s motus harbored wholly indigenous
vegetation, and the presence of one small patch of one exotic plant
aside from a few standard Polynesian-introduced plants, seemed to
indicate a relatively pristine atoll).

In 1987, the Office de la Recherche Scientifique et Technique
Outre-Mer (ORSTOM, a French scientific research agency) was requested by
the Kiribati government to conduct a short study at Caroline on the
feasibility of pearl-shell culture (G. Monet, pers. comm.). Their
results concluded that the atoll would be inappropriate for this type of
development.

12

Also from 1987 to 1991, a Scotsman, Ron Falconer, his French wife
Anne, and 2 small children settled on Caroline. From October 1989 to
November 1990 a new lease was under negotiation by Felix Urima, a French
businessman, who planned to blast a channel through the reef, construct
an airstrip, build a small hotel, cut timber, and engage in various
commercial ventures including fishing, a turtle farm and pearl-shell
culture. In April 1990, Urima’s workers began commercial fishing,
killing turtles and coconut crabs, and clearing land (A. Kepler 1990a).
This was a major new insult to the atoll which, in spite of its long
history of intermittent human occupation, remains to this day "possibly
one of the least spoiled of true atolls in the Pacific" (Stoddart 1976).
Reports from our expeditions to Caroline (Kepler & Kepler 1989,

A. Kepler 1990a and d) resulted in the short-lived cancellation of
Urima’s tentative lease in November 1990 by the government of Kiribati.
As of this writing, Urima has returned to unlimited fishing of
Caroline’s reefs. Since 1988, conservation efforts have been underway
for Caroline to become part of a triple-island wildlife preserve with
Vostok and Flint (Pt. II, Sect. G).

20th Century Scientific Studies

In June 1965, a field party from the Smithsonian Institution’s
Pacific Ocean Biological Survey Program (POBSP) visited Caroline for 2
days (Clapp & Sibley 197la). Their survey and specimens added much to
the previous botanical and ornithological knowledge of the island.

Other quick visits were made by Gilbert and Ellice Island officials and

Drs. H. & H. Grossman, ornithologists from Germany, and Mr. W. Cooke, a

graduate student in corals from the University of Hawaii, comprising the
Line Islands Expedition on 9-10 September 1974, and by Roger Perry, then
Wildlife Warden of the Line and Phoenix Islands, on 12-13 November 1977,
from which no reports can be found other than a short popular account of
the Southern Line Islands (Perry 1978).

In 1990, AKK also visited Caroline twice with the ICBP 1990 Line
and Phoenix Islands Expedition (A. Kepler 1990d). These visits were
primarily to discuss conservation matters with the Falconers; introduce
Caroline to Dr. and Mrs. M. Garnett, representatives from ICBP; confirm
the illegal taking of fish, turtles, and coconut crabs; collect
invertebrates; and fill in gaps from the 1988 expedition.

C. METHODS

Field Techniques

From 22-29 September 1988, Drs. A. K. Kepler, C. B. Kepler, D. H.
Ellis (U.S.A.) and Mr. Katino Teeb’aki (Republic of Kiribati) surveyed
all 39 motus at Caroline Atoll (Fig. 2), gathering detailed information
on plants, seabirds, land birds, mammals, reptiles, coconut crabs, and
human disturbance. Some incidental data have been added from the 2
visits in 1990 (10-13 March, 18-28 May) by Dr. A. K. Kepler,

13

Capt. G. Wragg, A. Garnett, M. Linsley, J. Phillips (March), and
Dr. M. Garnett (May).

Prior to the first expedition, a series of transects and known
botanical information were mapped to ensure that 5% of each motu was
sampled, and to maximize our chances of encountering all known plant
species. Transects on the 3 larger motus were spaced approximately
400 m apart and, with one exception, were perpendicular to the long axes
of each islet (Tr. 3 on Nake extended first from east to west, then ran
south parallel to the west coast). On motus longer than 400 m, we used
2 transects. Transects on the smaller motus passed through their widest
points. Their lengths ranged from 77 m (Azure) to 2,000 m (Tr. 3,
Nake).

Considerable modifications were required when we realized that all
previous maps (Figs. 4-7) were incorrect. We redrew the transects on
Arundel’s 1883 map (Figs. 4, 8), secured just prior to the expedition.
On South Island, due to impenetrable draperies of Ipomoea vines,
Transect 3 was omitted, Transect 5 ran only from the lagoon south to the
Ipomoea curtain (75 m), and Transects 4 and 6 ran north and south until
we reached an impasse (Pl. 8).

Compass headings were determined by the configuration of each
island. Beginning at high water mark, all distances (islet dimensions,
widths of reef flats and substrates, and plant communities) were
measured using hip chains with biodegradable cotton thread. These
parameters were later checked against aerial photographs in stereoscopic
pairs (RNZAF 1986), which provided 3-dimensional overviews of most
islets. Vegetation maps, reef and islet areas, and areas of plant
communities were derived by outlining on graph paper, enlarging, then
counting dots.

Data were collected in a 30 m swath along each transect (15 m to
each side) and recorded on field forms. Within each plant community we
took photographs, assessed the relative abundance of each plant species
(see Sect. E), measured notable trees, and recorded plant community
width, plants collected, and substrate type. We also estimated the
maximum height of the dominant vegetation and percentage of ground area
covered by each species. Data on seabirds, land birds, reptiles,
mammals and coconut crabs is reported in Part II.

In addition to the linear transects, an additional 19,300 m of
perimeter surveys were conducted on 21 islets (Fig. 9). The combined
distance for linear and perimeter transects was 32.6 km. Seven tiny
islets (Noddy Rock, Skull, Atibu, Bo’sun Bird, Coral, Reef-flat,
Fishball) were surveyed completely.

In 1988 we camped on the atoll for 7 nights, establishing base
camps (Fig. 8; Pl. 9) on the northwest point of South (22-24 September)
and southwest Long (25-28 September). We relocated camps using a Zodiac
with outboard motor, and an inflatable Sevylor canoe. All transects
were surveyed during daylight hours, beginning at dawn. Walking the
interislet channels was relatively easy at low tide, but became

14

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16

PERIMETER
SURVEYS

CAROLINE ATOLL

SCALE 1:34,000
500 ° S00 1000 2000

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meters

Figure 9. Caroline Atoll: perimeter surveys. The distance covered
was 19.3 km.

17

hazardous at incoming or high tide on account of numerous aggressive
black-tipped reef sharks, Carcharhinus melanopterus (Pl. 10).

During the 1990 visits, we stayed on Motu Ana-Ana with the
Falconers. Work was not intensive, as in 1988. We walked or motored an
inflatable Lancer, visiting 20 motus gathering incidental data, locating
the marae, etc.: Ana-Ana, Kimoa, Pisonia, Eitei, South, North Arundel,
Noddy Rock, Brothers, North Brothers, Skull, Pig, North Pig, Bo’sun
Bird, Long, Nake, Mouakena, Shark, Scarlet Crab, Bird, and Fishball.
Insects were preserved in ethyl alcohol. We used a "Magellan" NAV 1000
to obtain accurate geographical coordinates of Caroline.

Naming Caroline’s Motus

Previous literature has provided vague or incomplete data on
Caroline’s constituent motus (Bennett 1840, Markham 1904, Stevens &
Barwick 1930, Holden & Qualtrough 1884, Bryan 1942, Clapp & Sibley
197la, Garnett 1983). This confusion resulted because most previous
visits had been brief. The only charts available were a quite accurate
survey by Arundel, a guano merchant who mapped the atoll in 1883
(Fig. 4, Admiralty Chart, No. 979, 1965), and a map, greatly in error,
drafted by an international Solar Eclipse Party, also in 1883 (Fig. 5).
There are no hydrological navigation charts. Unfortunately, the
astronomers’ map has been used in all subsequent scientific, historical,
military and sociological publications [N.I.D. 1941, Bryan 1942
(Fig. 6), Maude 1968, Clapp & Sibley 197la (Fig. 7), Garnett 1983]. It
shows only 25 of the 39 motus and many shapes are distorted. The 38
motus on Arundel’s map are similar to those in the RNZAF (1986) aerial
photographs. Only a few appear to have changed in minor ways since
1883: major discrepancies in Arundel’s map, we believe, are due to
difficulties involved in the accurate rendition of small land areas
(i.e. the South Nake Islets). Maude (ca. 1938) counted 36 islets, but
never published his information.

To aid our survey we named 32 islets and 4 islet groups (Fig. 2).
Our names reflect appropriate aspects of islet biology. Etymology is
provided in Section H. Gilbertese names are prefixed with motu (see
next section). Any name not appearing on Arundel’s map (Fig. 4) was
given to the islets by us. They have been sent to the British Admiralty
and U.S. Hydrographic Office (along with corrections to the Pacific
Pilot) for official recognition.

D. PRELIMINARY STRUCTURE AND TOPOGRAPHY

Our geological terminology is based on Tracey et al. (1955) as
cited by Wiens (1962), to which we add motu (Polynesian for "islet"),
now a technical term for detrital reef islands (Danielsson 1954,
Stoddart & Steers 1977). In this paper the terms motu and islet are
used interchangeably.

18

Background Geological Setting

Caroline is the southeasternmost of the Line Group (Fig. 1), a
major volcanic lineament in the Pacific, comparable in size to the
Hawaiian-Emperor chain to the north and the Marshall-Gilbert-Ellice
chain to the west. This quasi-linear chain, 4200 km long, is composed
of dozens of simple and complex seamounts and linear ridges, 6 atolls, 5
islands, and 2 submerged reefs. The Line Islands are now considered to
include Johnston Atoll (Duncan 1983, Schlanger et al. 1984), and thus
lie between 17°N and 12°S latitude, and 169° and 150°W longitude. The
name, Line Group, reflects its equator-straddling location.

The geological complexities of the East Pacific in general and
Line Islands in particular were poorly understood until a few years ago.
However, recent remote-sensing technologies, deep-drilling techniques,
improved sea-floor mapping and multidisciplinary studies of sea-level
changes have clarified much previous speculation (Jarrard & Clague 1977,
Orwig & Kroenke 1980, Montaggioni & Pirazzoli 1984, Schlanger et al.
1984, Pirazzoli & Montaggioni 1988, Spencer 1989).

The Line Islands are now known to exhibit changing geomorphology
from north to south: elongated submarine ridges and coalesced seamounts
merge into a broad central high plateau, then progress to scattered
isolated seamounts in the south (Duncan 1983, Schlanger et al. 1984).

This long chain of geologically related and unrelated islands
exhibits a complex history of volcanism involving activity from multiple
"hotspots" and overprinting events dating as far back as 93 million
years (Jarrard & Clague 1977, Orwig & Kroenke 1980, Crough & Jarrard
1981, Haggerty 1982, Duncan 1983, Schlanger et al. 1984). Furthermore,
although numerous studies, using high technology, have been conducted
from oceanographic vessels, very little data has been gathered from the
islands themselves (M. 0. Garcia & J. A. Haggerty, pers. comm.), and a
clear understanding of the myriad interacting processes which formed the
Line Islands has not yet emerged. This is particularly true of the
central and southern Line Islands.

However, recent palaeoecological research in the Tuamotu
Archipelago includes generalities which are applicable to the Line
Islands (Montaggioni & Pirazzoli 1984, Pirazzoli & Montaggioni 1988).

The known geological history of the Line Islands can be summarized
as follows:

1) During the Cretaceous period (140-65 m.y. B.P.), ridge-building
volcanic events occurred, giving rise to scattered volcanoes, older
in the north. This period of mountain-building was a worldwide
phenomenon, in the eastern Pacific supported by hotspot activity in
the vicinity of Easter Island.

2) A second eruptive phase during the Palaeocene-Eocene period (65-38
m.y. B.P.), and age-progressive from north to south, overprinted

19

earlier volcanoes. This was either a result of "hotspot" activity
along the Line-Marquesan Swell or part of Pacific-wide volcanism.

3) The history of reef growth and subsidence is complex and has not been
studied in detail. The Line chain was in latitudes amenable to reef
growth throughout its history from the Late Cretaceous (100 m.y.
B.P.) to the present (Schlanger et al. 1984). A general pattern for
the northwest Tuamotus, close to the southern Line Islands, shows
primarily Holocene reefs 6000-3000 years old (Pirazzoli & Montaggioni
1988). However, a few older reefs do exist, and dredge hauls near
Caroline (Schlanger et al. 1984) recovered reef limestones of Eocene
through Pleistocene age (54 m.y. to 10,000 years B.P.).

We found no visible fragments of Caroline’s volcanic heritage,
summarized above.

Sea Level Changes

Data on the history of sea levels for the Southern Line Islands is
lacking. However, studies in French Polynesia (Pirazzoli & Montaggioni
1988), and which appear to have been a general phenomenon in the South
Pacific, indicate that:

1) A stable sea level occurred, slightly less than 1 m above its present
level, from 5000-1500 B.P. This peaked at approximately +1.0 m
between 2000 and 1500 years ago.

2) Since then the level has been dropping gradually to its present
position, reached only recently (Pirazzoli & Montaggioni 1988).

General Account

No geomorphological or geophysical studies have been carried out
at Caroline. However, 2 deep undersea dredge hauls near the atoll
uncovered reef limestones dated at Eocene through Plio-Pleistocene
(Schlanger et al. 1984), and recent studies in the northwestern Tuamotus
date the exposed coral reefs in the Holocene around 6000 to 3000 years
B.P. (Pirazzoli & Montaggioni 1988). There is much scope for research
within Caroline’s reef matrix, varied shorelines and upraised reefs.

Caroline’s overall shape resembles a flattened crescent, 9.7 km
long on its north-south axis (Fig. 2), with outer perimeter 26.9 km and
greatest breadth 2.3 km. The longest islet, Long, extends 4.23 km
north-south, while South Island, extending 1.2 km east-west, claims the
widest stretch of land.

The motus, lying upon a wide, continuous reef flat which encloses
an elongated, relatively shallow lagoon, fall naturally into groupings
of 3 large islands (South, Nake, Long) and 4 groups of smaller islets
(13 Windwards, 5 Southern Leewards, 11 Central Leewards, 7 South Nakes).

20

There are 4 basic motu shapes, molded by the prevailing easterly winds
and currents, periodic storms, overall atoll shape, etc:

}) long, linear, and parallel to the reef axis, e.g. Long Island.

B) small, linear or oval, and perpendicular to the reef axis, e.g.
Southern Leeward Islets.

3) triangular or crescentic, with the apex facing the seaward reef, e.g.
most of the Windward Islets.

4) large and quadrangular, occupying the ends of the atoll, e.g. South,
Nake.

Caroline’s motus have similar length-width ratios as those
elsewhere (Stoddart & Steers 1977) and are similarly situated on the
inner half of the reef flat, having their lagoon beaches close to the
lagoon reef slope. Individual motus are discussed in detail in
Section H.

It is hoped that the following preliminary observations of
Caroline Atoll will inspire further research. As well as exhibiting
features similar to many atolls, its 39 islets also present individual
details that pose interesting questions which may help in deciphering
sea-level changes in the Eastern Pacific and in unraveling the somewhat
speculative geological history of the Line Islands (Jarrard & Clague
1977, Schlanger et al. 1984). Examples include the presence of an
inland, vegetated reef substrate on Long; deep inland sand on Nake;
conglomerate rock on South; hardpan on Mannikiba, Nake and Long; an
emergent reef platform (Noddy Rock); and exposed older reefs of uniform
height (Nake). Aerial photographs have indicated that Long Islet has
been formed most recently from the coalescence of 5 former islets, which
show even older subdivisions. Nake, previously at least 2 islets, has
changed shape on both its north and south ends by the addition of gravel
ridges and silt, respectively, in the past 100 years. Brothers Islet
has incorporated a small motu within its confines since 1883; several
other motus have also added bars and spits, and the lagoon has filled in
further during that short time period. The questions of phosphatic
hardpan beneath Pisonia forest and the extent of ground water lenses
need attention.

Reef Flats

Caroline’s peripheral reefs, which completely surround the lagoon,
and upon which the motus rest, are consistently wide (average 562 m,
range 396-759 m, N = 100). The windward and leeward reefs differ in
structure and dimensions. Neither are entirely dry, even at the lowest
tides. They consist primarily of barren calcareous rock, frequently
smooth, which on other atolls generally represents the erosional surface
of an older reef. Jagged "mushrooms" of exposed newer (but dead) reef
framework dot the leeward reefs, forming an open platform off
southwestern Nake (Pl. 11). Their structure and uniform height (-0.3 m)

21

are similar to those on Hikueru Atoll, Tuamotus, which have been dated
as 2565 + 55 years B.P. (Pirazzoli & Montaggioni 1988). There are no
passes from ocean to lagoon, a typical feature of central Pacific atolls
(Wiens 1962). In the Southern Hemisphere, reef flats tend to be widest
in the southwest sector and narrowest in the northeast (Wiens 1962), a
generality which Caroline fits (Pls. 12, 13).

The reef rim, irregularly dentate and 26.9 km in circumference, is
surmounted by motus for 55% of its length. On 72% of all Pacific
atolls, less than half the reef circumference is occupied by land (Wiens
1962); Caroline lies within a 28% minority in which one-half to two-
thirds of the reef rim contains land. Corresponding values for 2
Tuamotuan atolls, Rangiroa and Raroia, are 33% and 35% (Stoddart &
Sachet 1969). Where motus exist, the reef flat is divided into the
seaward reef flat (Pl. 12), motu, and lagoon reef flat (Pl. 14).

At low tide all reef flats can be waded. Black-tipped reef sharks
were highly aggressive in 1988 but by 1990 dozens had been killed. The
South Nake and Central Leeward channels were particularly hazardous,
reflected in motu names such as Blackfin, Shark, and Danger.

Windward Reef Flats: Constantly pounded by surf, the windward reefs are
typically narrower than those to leeward, averaging 519 m (range 396-
759 m), though this is less evident from a map than in the field.

The windward reef is 13.5 km long, surmounted by 16 motus that
total 63% of its length. This fits a recurrent pattern on central
Pacific and Tuamotuan atolls where motus are more frequent along
windward reef rims (Thomas 1961, Wiens 1962), due to active movement of
debris associated with the prevailing easterly winds, waves and storms.
The longest islets are Nake (1,980 m) and Long (4,226 m), both formed
from the coalescence of 2 or more smaller islets. The rest vary from
18 m to 858 m in length.

The character of these reef flats differs, depending on the
presence or absence of land, interisland distances, lagoon depth, and
recent weather conditions. In February 1990, severe cyclonic weather
rearranged tons of sediments, especially to windward, uprooting
Tournefortia scrub, obliterating extensive sections of herb mats,
exposing beachrock, depositing storm blocks, and altering the shape and
slope of the beach crests. Since Caroline has been essentially
uninhabited for 60 years, no data exists on the frequency of such
storms, although it is well-known that windward beaches worldwide are
undergoing erosion and retreat, and are thus characterized by beachrock
outcrops and other lithified sediments (Stoddart & Steers 1977).

Reef Rim with Motu: The width of the seaward flats is quite uniform,
averaging 307 m (range 193-396 m), occupying 57% of the rim width. It
consists of a slightly raised algal ridge bearing the brunt of incessant
wave action, and a rubbly reef flat, partly drying at low water, which
sweeps up to the motu’s beach (Pl. 13).

22

The motus differ considerably in width, ranging from the narrow
tip of Long, merely 30 m wide, to Windward, 290 m wide. Nake and South
Islands, forming "caps" to the atoll at its upper and lower ends,
respectively, exhibit characteristics more typical of windward than
leeward motus. Whenever atoll reefs turn sharply, debris-loaded waves
become deflected around the points, thus depositing more gravel than on
a straight shoreline. Hence, these 2 motus are the widest on the atoll
(Pl. 16). A comparison of maps a century apart (Figs. 2, 4) indicates
that several layered ridges of coral debris have accumulated on northern
Nake since 1883.

Reef Rim without Motu: Zonation within the reef flat is less marked
where there is no land. Within these interislet reef flats, however,
areas of high water transport have carved surge channels and erosional
grooves, and tidal fans extend into the lagoon, especially at its
northern end where sedimentation is most active. Caroline has no deep
pass or navigable channels into the lagoon, nor a ship anchorage beyond
the reef, though small boats may anchor within the close lee of South
Island during normal trade winds and low seas. Landing in an inflatable
is best made across the reef slightly north of the "boat entrance,"
marked by an upright anchor (App. I).

The reef flat between Tridacna and South Island, serrated with 6
erosional grooves, one labelled "blind passage," is of particular
importance to navigators (App. I). The blind passage, the most
southerly channel, is a narrow diverticulum 380 m long within a reef
430 m wide. On all previously published maps this passage is drawn as
though it completely connects ocean and lagoon (Figs. 4-7). However, it
is a nonfunctional hoa or tairua, an erosional channel cut only partly
across a continuous atoll rim. Its lagoon end (Fig. 50, Pl. 72) serves
as a sheltered anchorage for motored yachts, but it can only be entered
or exited during high winds or moderate-to-high surf. Chevalier (in
Stoddart & Steers 1977, p. 77) has suggested that hoa features are
partly a result of sea-level changes altering the balance of
sedimentation and erosion.

Leeward Reef Flats: These are wider, flatter, gentler, more
consolidated, and less filled with rubble than the windward reefs

(Pl. 11). An orange, semi-transparent alga, blanketing the coralheads,
chunks of upraised coral (Pl. 11), carbonate rock, and giant clams, is
abundant. This alga is found on many atolls, for example Enewetak and
Rangiroa (U.S. Department of Energy 1987, Stoddart & Sachet 1969).
Living coral is sparse.

Surge Channels: These occur in a variety of shapes and sizes, depending
on the distances between motus, the extent and buildup of reef flats
adjacent to land, and lagoon depth. Surge channels and reef grooves are
deeper on the windward side. The vigorous currents washing daily into
Caroline’s lagoon have created larger debris fans between windward motus
than between those to leeward (Frontispiece).

23
Beaches

Caroline’s beaches--the zones lying between low water mark and the
inland limit of wave-deposited debris--are entirely of reef origin.
There is, however, considerable variation in their composition. The
windward beaches and surge channels, in a constant state of erosion or
deposition, support the greatest variety of sediments: well sorted
sands (indicated by grain-size distribution); gravels of coralline,
algal (including Halimeda) and molluscan origin; and a wide assortment
of coral fragments. The atoll’s prime stretch of sand (Shark Islet) is
thickly overlain with pink granules, possibly Foraminifera tests, which
are abundant in the Tuamotus (Stoddart & Steers 1977).

Almost all exposed rubble on Caroline is colored gray, a
consequence of penetration by cyanobacteria. Typically the oldest
rubbles, highest up the beach and extending into the interior, are
darkest. A marked beach crest rises, gently or abruptly, from the
windward beaches, at the crest of which is deposited an assortment of
flotsam and jetsam: bottles, plastic, wood, coconuts, etc. (Pl. 19).
No storm blocks were found in 1988, but in 1990 many littered the
windward reefs and shores, the result of recent cyclonic weather.
Similarly, in 1990, thick deposits of coarse sand had overlain the
rubbly windward beaches and interislet channels of 1988 (Pl. 33).

Alterations to Caroline’s beaches provide the major changes in
motu shape. Aggradation occurs principally on the lagoonward points of
the larger windward islets, for example, Brothers (which is now joined
with a separate islet mapped by Arundel), and Windward and Tridacna
(which have added more sediments to their southwest points during the
last century).

Beachrock: These elongated strata of eroded reef, brown consolidated
sands and reef detritus, from one to a few meters wide, are not abundant
on Caroline. Occurring as seaward dipping strips at the low water mark,
they flank the windward beaches of Nake, Long and South (Figs. 37, 38,
50; Pl. 54) and a few of the leeward islets. Beachrock results from
lithification of tropical intertidal sediments by calcium carbonates, in
part due to seawater evaporation during low tide (Scoffin & Stoddart
1983). Beachrock outcrops become more exposed after storms, indicating
that some cementation may occur beneath a shallow sediment cover. A
coarser conglomerate platform (Pl. 20), possibly a relic of a former,
high sea level, occurs on the northwest point of South Island, creating
the "landing" (Pl. 20). This appears to be of similar age to the
remnant reefs of southwest Nake, which are less consolidated (Pl. 11).
Although the platform’s upper surface rests slightly above high water
level, we do not know if it represents a former reef exposed by a recent
fall in sea level. The origins of such platforms are controversial
(Stoddart & Steers 1977).

Upraised Reef: In a few areas, jagged, eroded upraised reef (feo)
comprises some of the islet’s interior--for example, the lower quarter
of Long. A thin soil cover supports a forest of lower stature than
would otherwise be expected. The rocky substrate is pitted with

24

cavities and undermined with subterranean tunnels in which at least 2
species of land crabs (Birgus latro, Cardisoma spp.) shelter (Pl. 21).
Noddy Rock (Pl. 18), the smallest motu (0.02 ha), and many jagged
coralline "mushrooms" found on the reef flats (Pl. 11) are probably
remnants of former reef flats formed when sea levels were several feet
higher than present. It is hoped that further investigation will
determine whether these older limestones are from the Holocene or
Pleistocene.

Lagoon

Caroline’s lagoon, 8.9 km long, is closed; its total area is less
than that of the combined reef flats. The lagoon is relatively shallow,
tapering in shape and depth at each end, containing both reticulate and
patch reefs of living coral. Its bathymetry is unknown.

In the north the lagoon is more sheltered, as the presence of
continuous vegetated land buffers the easterly trades, and silty
sediments increase. At its northern extremity, merging reef flats
squeeze the lagoon until it disappears east of Pandanus Islet. A
filled-in portion of the former lagoon penetrates Nake for 300 m as a
fishhook-shaped mudflat, Sandy Inlet (Fig. 37, Pl. 22), before
succumbing to encircling vegetation. At the lagoon’s southern end,
where winds whip through the "blind channel," it is choppy, having more
sediment and slightly less visibility. However, within the lee of South
Island’s north-central curve the lagoon is frequently quiet and
reflective (Pl. 23).

Lagoon Hydrology

Although Caroline’s hydrology has not been studied, the south end
of the lagoon and "blind passage" (Fig. 50, Pl. 72) were closely
observed for 2 years by Ron Falconer. He noticed that the lagoon is
typically "perched" at a level above that of all but the daily high
tides. High tide water flows rapidly over the reef flats into the
lagoon, but is held back by the reefs as the tide lowers. Lagoon water
at low tide is about 0.3 m higher than water in the "blind passage."
Water moves out of the lagoon through a few channels that, although deep
in places, form broad, shallow troughs over the reef flats. A major
channel with a current flowing west at several knots passes along the
northwest point of South Island, although water passage is impeded by
the reef flats west of South Island. If a channel were to be blasted
through the reef flats, as has been proposed, this delicate hydrology
would be disrupted. For example, the high tide water is never more than
20 cm above the coral heads and reefs in the lagoon. A man-made reef
channel for vessels could lower water levels 30-40 cm, thereby exposing
and killing the extensive Acropora-Tridacna reefs within the lagoon.

The "blind passage" is sustained by a powerful northward flow of
water along the east coast of South Island and a strong southward flow
of water along the seaward reefs of Tridacna Islet. The South Island

25

flow is augmented by water draining from a large shallow basin on its
windward reef flats. Water spills into the "blind passage" and drains
east at about 4 knots against the prevailing trade winds and surf.
There is minimal current at the west (inner) end of the passage, where
less water is collected, and throughout the passage at low tide when
there is essentially no water flow out of the lagoon.

Patch Reefs

Darwin (1842) recognized 3 basic reef types (fringing and barrier
reefs, atolls). A fourth type, patch reef, is now widely accepted.
Patch reefs are smaller than the other 3 reef types, lack a lagoon but
are located within lagoons, are submerged up to low tide level, and,
unlike other oceanic reefs, have foundations of sediments or sedimentary
rocks. They are subcircular to irregular in shape when viewed from
above, and in their smaller sizes merge with coral knolls, coral
thickets, or coral heads (Ladd 1977, Fagerstrom 1987).

A complex series of patch reefs and coral knolls (primarily
Acropora spp.), circular and elongated, flank the smaller motus and
crisscross much of Caroline’s lagoon (see Frontispiece). They are
particularly evident in the southern two-thirds of the lagoon. Coral
limestone bedrock, surmounted by abundant living coral, molluscs, and
other invertebrates, provides their basic structure (Sirenko & Koltun,
in press). The atoll’s perimeter reefs provide shelter from storms,
surf, and excessive erosion. In shallow areas they tend to be
curvilinear (Central Leeward Islets), while in deeper water, coral
knolls and pinnacles are more characteristic (Pl. 63).

Caroline’s lagoon is gradually filling in with ever-expanding
patch reefs and debris washed in from the fringing reefs. Since
Arundel’s time, the effects of detrital deposition can be discerned as
changes in the shapes of islets such as Nake, Danger, and Arundel
(compare Figs. 2 and 4).

Such change is typical of atoll evolution. Geologically, Caroline
is a few steps behind one of its "neighbor" Line Islands, Christmas,
where sediments and coral growth have converted the original lagoon into
a maze of supersaline, mini-lagoons and tiny islets, mostly cut off from
the sea. Further steps in evolution are exemplified by completely
filled-in atolls such as Jarvis and Vostok, where not even salty pools
remain.

Tridacna-Acropora Reefs

Though the giant clam (Tridacna maxima) is an abundant component
of Caroline’s lagoonside reefs, exceptional aggregations flank the most
southerly windward motus (Brothers through Tridacna, Figs. 44 to 48).
Two especially outstanding reefs extend across the lagoon from Tridacna
to Ana-Ana (Fig. 10, Pl. 25) and Tridacna to Kimoa (Fig. 48), where
Tridacna attach to Acropora spp. corals, a favored substrate (Braley

26

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27

1987). Abundant inshore Tridacna on all these islets suggest that their
density is similar to phat on the main reef: up to 20/.25 m° (i.e
80/m"), averaging 35/m” for the entire area surveyed (Sirenko & Koltun,
in press). This density exceeds the highest known aggregations of
Tridacna: up to 60/m° at Reao Atoll, Tuamotu Archipelago (Richard
1985). Densities of 6-20 clams per square meter, as found at Takapoto
Atoll (Tuamotus), are considered high. Throughout Caroline, the clams
averaged 18 x 10 cm in size. Several species of Indo-Pacific Tridacna
have suffered greatly from poaching and overharvesting, leaving few
undisturbed populations (Braley 1987). Caroline is thus a special
refuge for 7. maxima.

Lagoon Reef Flats

These vary considerably but are narrower and more gently sloping
than the seaward reef flats. They are typically covered with fine coral
gravel and coarse sand. In sheltered areas (lower Long, Windward,
Crescent, South, upper end of lagoon) lush shrubbery--Cordia,
Tournefortia, Pisonia, Cocos--overhangs the lagoon. Here fine silt,
sand and/or an algal slime are common (Pl. 27). In 1988, narrow, sandy
beaches were limited to northern South (Pl. 23) and eastern Shark
(P1. 28), but in 1990, sand occurred throughout Caroline.

Where the lagoon shorelines are less sheltered and vegetation does
not overhang the lagoon, unvegetated rubble and sparse herb mats are
typical. Here, lagoonside rubble averages less than 2 m wide (Pl. 30).
This contrasts with their seaward reef flats, which average 21 m wide
(Plea)

Lagoon Reef Fauna: A Brief Summary

Caroline’s marine environment is rich yet essentially
undocumented: knowledge is limited to preliminary lists of fish,
invertebrates and lagoon plankton (Dixon 1884; Fowler 1901; Pilsbry &
Vanatta 1905a, b; Tsyban & Smith 1988, Sirenko & Koltun, in press). All
early travelers remarked on the beauty, abundance and variety of
Caroline’s reefs (Markham 1904, Bennett 1840); today they are still
relatively untouched.

The usual assemblage of reef invertebrates--echinoderms, molluscs
(Turbo, Nerita, Cypraea), crustaceans, porifera, tunicates, etc.--are
present. Corals include several important Holocene reef-constructors:
Acropora, Pocillopora, Porites, and Montipora. Calcareous algae include
Halimeda, Porolithon and Lithothamnion (Sirenko & Koltun, in press).
Large numbers of black sea cucumbers (Ludwigothuria sp.), about 20 cm
long, are particularly abundant lagoonward of the southern windward
islands (Pl. 10). Conspicuous fish families include parrot fish
(Scaridae), butterfly fish (Chaetodontidae), surgeonfish (Acanthuridae) ,
damselfish (Pomacentridae), pufferfish (Tetraodontidae), and wrasses
(Labridae).

28
Substrata

Throughout the atoll, substrates reflect a reef origin. There is
little "soil" in the accepted sense. Various grades of jagged, eroded
coral and molluscan rubble (from fist-sized to tiny pebbles), together
with sand, coralline algae, and relatively small proportions of organic
litter, humus and guano, are present. Such accumulations of reef and
terrestrial debris are similar to those of other low, coral atolls
(Fosberg 1953, Stone 1953, Wiens 1962, Niering 1963, Stoddart & Sachet
1969, Garnett 1983, Reese 1987, Gessel & Walker 1992).

Generally speaking, atoll soils are calcareous and extremely
immature, a consequence of their limited age and frequent disturbance by
storms. Barely modified beyond the reef that spawned their presence,
they are rich in calcium and magnesium carbonates. Water retention, if
any, is due to accumulated organic matter and its associated chemical
changes. Accumulations of guano react with the calcium carbonate of
reef sands and elevated limestones to form nitrogen-rich "soils" and
phosphatic hardpan (Hutchinson 1950, Fosberg 1953, Stoddart & Scoffin
1983).

Reese (1987) categorizes atoll "soils" into 5 types, all of which
occur, in different proportions, at Caroline. The degree of organic
matter, decomposition, amount of humus, and the depth of the "soil"
strata are directly correlated with age and size of Caroline’s motus.

1) Accumulations of coral rubble, mainly of stone size. These youngest
of "soils" are most evident around the edges of the motus, acting as
a substrate for natural herb mats. Often extending well inland, they
can support surprisingly lush Journefortia scrub.

2) Unaltered coral sand and gravel. Although exposed sand was uncommon
at Caroline in 1988, this substrate occurred intertidally where the
lagoon was filling in and on actively growing sandbars, primarily in
the upper lagoon (Pls. 22, 27), northeast and northwest South
(Pls. 23, 31), and the lagoonward edge of Shark (Pl. 28). In 1990, a
single storm deposited tons of sand on Caroline.

3) Soils with a weakly developed A-horizon, with color only slightly
darker than the unaltered sand below, but with no evidence of
structural development. Especially evident in 1988 within the
ancient interislet channels that compose Long Island (Pl. 32), much
of this substrate is now storm-eroded and overlain by fresh sand
CPIR9133))R

4) Soils with a more developed A-horizon, deeper and darker than above,
with some structural development. This stage defines areas where the
rubbly/sandy substrate approaches a true, but poor, "soil." As such,
it represents older, more stable parts of each island. It is common
within the islet interiors where Pisonia is (or was) present. Its
composition may be likened to a coarse mixture of gravel, sand,
bones, humus, eroded coral and shells, all mixed with sparse amounts

29

of partly-decomposed litter. Land crabs are particularly numerous,
further breaking down organic matter into finer particles.

5) Soils with an accumulation of raw humus on the surface and with a
relatively deep A-horizon. During this stage phosphatic hardpan may
develop. These true soils, though somewhat depleted by guano
diggers, cover significant areas on South and Nake. Cocos and/or
Pisonia debris heightens their dark coloration and moisture content.
This earthy substrate is composed primarily of rotting Cocos fronds
and nut fibers shredded by coconut crabs. Patches of blackish muck
on South Island support local patches of Tacca Jeontopetaloides.

On Caroline, we noticed a type of hardpan (Pl. 67) in several
areas (primarily South, Nake, Long, Emerald, and Mannikiba). In each
case it resembled a flat sheet of old asphalt road, present within, or
adjacent to, herb mats and peripheral Tournefortia scrub. Since hardpan
forms when phosphatic derivatives from guano interact with permeable,
reef-derived carbonate sediments (Stoddart & Scoffin 1983), it is
possible that these barren clearings represent areas of high guano
concentration. Worldwide, the major guano-producing seabirds are
boobies, terns and frigatebirds (Hutchinson 1950). Adjacent to, or
nesting on, these areas of hardpan which we observed were Red-footed
Boobies (all motus), Great Frigatebirds (all except South), Lesser
Frigatebirds (Nake), Masked Boobies (Long), and Sooty Terns (only 1990
on Long, but recorded from open areas of Emerald and Mannikiba, June
1965 and July 1990, see Pt. II, Fig. 11).

We did not take soil profiles within Pisonia forests to ascertain
the presence of subsurface phosphatic hardpan. However, AKK and John
Phillips found both surface and subsurface phosphatic hardpan on nearby
Vostok, having the typical "pepper-and-salt," coloration and crumbly
texture characterized by the Jemo soils found in the Marshall Islands
(Fosberg, 1954 and pers. comm.).

Caroline provides an excellent example of the progression of soil
development through islets of different age and size classes (see Sect.
F). From a wave-washed mound of coral rubble, barely above sea level
(Fig. 5), the substrate gradually improves in texture and fertility as
the emerging islet ages and organic matter accumulates. Pioneer plants
are hardy, salt-tolerant, low-lying mats consisting primarily of
Heliotropium, and later, Tournefortia. Increasing numbers of shrubs
provide shade and branches for nesting seabirds. Larger trees (Pisoni7a,
Cordia, Morinda) add more shade and thereby increase humidity, as well
as provide opportunities for additional organic "fallout": leaves and
bird remains (nests, eggs, chicks, droppings, regurgitated food, dead
adults).

Each stage of substrate development accelerates the accumulation
of organic material and helps to define an emerging, deeper A-horizon.
Soil maturity is indicated by more organic matter, improved soil
texture, and a lowered volume of coralline and molluscan debris.
Caroline’s soils barely exceed several centimeters in depth and are

30

always intermingled with coral fragments. As a result, they are
unsuitable for burrowing seabirds such as petrels and shearwaters.

Hydrology

Hydrological information is essentially lacking. No standing
fresh water exists. The quality, extent, and salinity of the freshwater
lenses, as well as their variability according to tide, season, and
rainfall, are unknown. At the time of Caroline’s "discovery" (1606),
de Quiros and his party were desperate for fresh water. After noting
how lush and green Caroline was, they expected to find good water
supplies, but there was "nothing but salt water in the holes they dug"
(Markham 1904). Maude (1968) suggested, in hindsight, that had they
waited longer the salt water in their shallow wells might have run
fresh, as has been his experience on some other atolls. During the 19th
century, 3 wells were used--one on Nake and 2 on South (Holden &
Qualtrough 1884). One South Island well contained fresh water at 1.5 m
depth in 1974 (Garnett 1983). We saw no wells, but located 2 concrete
cisterns, one built in 1937 near the "landing" and rebuilt by the
Falconers in 1989, and another uncovered one (dating from 1938) within a
Cocos-Pisonia grove along Transect 2, about 200 m east of the southwest
corner of Nake.

Caroline’s paucity of fresh water may be partly responsible for
the lack of a permanent population. The annual rainfall in 1989
(App. II) was 1,242 mm (48.9"). However, like the similarly lush
Nikumaroro and Orona (Phoenix Islands), Caroline’s rainfall may vary
greatly from year to year, resulting in undependable water supplies.
Residents always relied on rainfall catchment for fresh water (Maude ca.
1938; R. Falconer, pers. comm.).

The relationships between fresh water and Pisonia forests are
uncertain (see Wiens 1962 and Sect. F).

Climate

Meteorological records for Caroline were sparse until 1989, when
Ron Falconer began daily records of rainfall and wind direction
(App. II). Some data is available from the plantation years 1916-1920
(Young ca. 1922) and during the 1883 Solar Eclipse Expedition
(20 April - 8 May) (Upton 1884), when 203.2 mm (8") fell.
Generalizations on regional weather conditions are found in N.I.D.
(1943), Seelye (1950), Fosberg (1956), Newell (1956), Wiens (1962),
Taylor (1973), Stoddart & Scoffin (1983), and various papers on the
Tuamotus (Stoddart & Sachet 1969, Sachet 1983). Islands in the Line
Group experience a wide range of climates. In general, those near the
equator are dry, with rainfall increasing with increasing latitude north
or south (Fig. 10).

Caroline experiences a tropical oceanic climate with little annual
variation. Temperatures are uniformly warm to hot, normally tempered by

3h

trade winds from the southeast to northeast. Falconer (pers. comm.) did
not record daily temperatures but estimated an annual average of 29° C
(range 26°-31° C). Mean annual temperatures for the Central Equatorial
Islands lie between 24° and 29° C. Surface temperatures increase
rapidly in early morning and remain hot throughout the day. Forest
interiors are humid. The daily range of temperatures exceeds the annual
fluctuation in the daily mean.

Atmospheric pressure, sunshine, and cloud cover are probably
similar to the northern Tuamotus, uniform except during storms.

Wind and Rainfall

Caroline is dominated by trade winds. As on all low atolls, land
topography has very little appreciable effect on weather. Although it
lies within an area primarily influenced by southeast trades, there is a
small annual oscillatory movement northward and southward. Appendix II
indicates that, at least for 1989 and 1990, winds blow primarily from
the north and northeast, and rarely from the southeast (April - August).

The atoll lies within a belt of variable rainfall, along with
Vostok, Flint, and the Northern Tuamotus. Young (ca. 1922) measured
rainfall for 1919 (2,172 mm) and 1920 (1,854 mm), noting that Caroline’s
rainfall is "certainly less than that of Flint." He estimated "probably
not more than 50" (1270 mm)" during 1916, 1917 and 1918, figures
extrapolated from Flint (1,600, 1,346, 1,295 mm, respectively).

Falconer measured 1,242.1 mm (48.9") in 1989 and 2,209.8 mm (87") in
1990. An unusually wet February in 1990 brought 640 mm (25.2") of rain,
due to cyclones "Peni" (centered near Vostok) and "Ofu" (centered
further west). Rainfall distribution isohyets (Taylor 1973, Stoddart &
Scoffin 1983) assign Caroline an approximate annual precipitation of
1,500 mm (60"), a perfect average of the above 6 years (x = 1,513 mm).
In general, "winter" (May - October) corresponds roughly to a dry
season, and "summer" (November - April) to a wet season.

Cyclonic Storms

Atoll motus are active structures, undergoing repeated death and
rebirth. Violent storms contribute to ongoing erosional and rebuilding
processes. Storms deposit debris along windward shores (Pls. 17, 19),
into the lagoon and often far inland.

Although the south-central Pacific is relatively free of cyclonic
storms (cyclones, typhoons, hurricanes), they occur with enough
frequency and devastating force that any discussion on climate should
include them. Although detailed records of hurricanes and tropical
storms exist for the inhabited Tuamotus since European discovery, we
know little of those that have affected Caroline. The following
evidence suggests that Caroline experienced 3 major hurricanes since the
1820s and that periodic violent storms modify the atoll substantially:

32

1) Between 1822 and 1825. When de Quiros visited Caroline in 1606, the
northwesterly Cocos plantation on South Island was healthy. When
Bennett arrived in 1834, he noted that all the palms were "of dwarf
stature," and that "amidst the original groves, the number of vigorous
seedlings fully confirmed Captain Stavers’ statement [who had visited
the atoll in 1828] that these palms had increased greatly since his last
visit to the spot" (Bennett 1840).

A few years before 1828, therefore, something had affected the
palms. By 1834 they were all of an even height and quite short, yet
bore nuts. French records indicate that 2 devastating storms whipped
through the Tuamotus during this time--in 1822 and 1825 (Sachet 1983).
At least one of these could have affected Caroline.

2) The 1878 cyclone. The first unambiguous record of major devastation
at Caroline comes from the letter of a certain J. M. Salmon, dated 1883
and reproduced in Holden (1884). Speaking of the time when

Messrs. Brown and Brothers took possession of Caroline (somewhere
between 1865 and 1872), he stated that "it seemed as if there had been a
storm or hurricane at some short period previous, which had desolated
the place." Arundel (1890) attributed this to a tidal wave that swept
across the Pacific from South America to New Zealand and Australia in
1868 (Arundel 1890), but atolls do not generally suffer greatly from
tsunamis because they lack focusing relief. The Hydrographer of the
Navy (1931, Vol. III, p. 154), however, referring to Caroline, clearly
states that in "1878 a cyclone passed over the islands, destroying most
of the coconut trees."

The Great Britain Naval Intelligence Division (N.I.D. 1943,
p. 490), in reference to Caroline, also states that "in 1878 a hurricane
wrought great destruction." This was possibly the violent storm of 6-
7 February 1878, which killed 117 persons on Kaukura Atoll, 750 km
southeast of Caroline in the Tuamotus (Sachet 1983).

3) The 1906 hurricane. Serious storms occurred in the Tuamotus during
1903 and 1906 (T. Spencer, pers. comm.). There is no record of the
effects of the latter storm on Caroline. However, it affected Flint
Island (Campbell 1908, p. 2), hence must have passed over Caroline as it
passed westward from the Tuamotus. Campbell, leader of the Solar
Eclipse Expedition to Flint (1907-1908) states that "the great hurricane
of 1906 February or March drove water to (the manager’s residence,
native huts and copra warehouses, all located 22 feet ASL and slightly
inland) and the water threw the warehouses off their post foundations."

Ten or more coconut palms on Flint were struck by lightning in
March of 1917 and 1918 (Young ca. 1922).

4) The 1990 storms. Our second visit to Caroline was 2 weeks after
cyclone "Peni," centered near Vostok (February 1990), affected the
atoll. Violent winds, torrential rain, and high seas had defoliated and
uprooted vegetation in some windward areas (P1. 33) and greatly altered
Caroline’s shorelines, interislet channels, tidal fans, and incipient
islets from our 1988 visit. Tons of sand and rubble were rearranged on

33

both windward and leeward islets, Motu Atibu virtually disappeared, and
the main interislet channel that divides Long Island lost its herb mats
and many Tournefortia shrubs, becoming smothered with fresh sand.

We note here that nothing is known of the effects of the 1982-83
El Nino Southern Oscillation at Caroline. This phenomenon is
characterized by the appearance and persistence, for 6 to 18 months, of
anomalously warm water in the equatorial waters of Peru and Ecuador.
Its biological consequences are dramatic and large-scale, extending far
into the central Pacific Ocean: diminished plankton production, reduced
fish stocks, starvation and mass breeding failure of seabirds, heavy
rainfall, growth of vegetation and disappearance of nest sites (Barber &
Chavez 1983, Cane 1983, Stoddart & Scoffin 1983, Schreiber & Schreiber
1984). Our only comments are that our seabird population figures were
either similar to or greater than those in 1965 (Pt. II) and that the
only dead birds we saw were in March 1990, victims of the cyclonic
weather described above.

Because islets on coral atolls rarely exceed 5 m in elevation, the
tidal surges associated with Class IV or Class V hurricanes, often
exceeding 5 m in depth, can overwhelm them, not only altering or
destroying the vegetation, but in extreme cases completely removing them
from the coral rim (Frisbie 1944). It is essential to consider the
ephemeral nature of Caroline’s motus in the discussions that follow.

Sea Conditions

Because the most extensive coral rubble deposits occur around
northern Nake and southern South Island, and because the Cocos
plantation of northwest South was so badly hit by storms last century,
the following Tuamotuan generalities (Newell 1956) probably also apply
to Caroline:

1) Prevailing trade winds from the east give heavy seas on the northeast
or windward side;

2) Southern ocean swells generated in the sub-Antarctic break heavily on
the south or seaward side; and

3) Occasional hurricanes or tropical storms strike in the northwest or
stormward quarter.

Tides

Only scanty data are available. In May 1883 (Holden & Qualtrough
1884), the greatest daily variation ranged from 475 mm (1’7") to 125 mm
(5"), i.e. 350 mm (1’2"). The standard hydrological chart (Fig. 4)
states 1.5 feet (0.5 m), which we use in the schematic profiles
(Figs. 32, 35, 36) as the difference between low and high spring tidal
levels. For Flint, W. Campbell (1908) guesses "about two feet" (0.6 m),
while Ward (1974) estimates 1.5 feet (0.5 m). Tidal fluctuations are

34

similar to those occurring in the Tuamotus, around 0.7 m (2’) (Newell
1956, Stoddart & Sachet 1969).

E. FLORA: VASCULAR PLANTS AND FLORISTICS

Botanical History

All early visitors to Caroline described a well-wooded atoll with
numerous islets whose vegetation extended to the shoreline. It has
changed little in the 384 years since its Western discovery. The first
botanical collection and notes were those of Bennett in 1835 (Bennett
1840), who recorded 10 flowering plants and a fern, and planted Tahitian
chestnut, sweet potato, and Polynesian arrowroot. He noted that "some
of the loftier trees" on South Island and the Southern Leewards, were 20
feet high, perhaps a consequence of cyclonic weather in the 1820s. The
location of his plant collection, if it still exists, is unknown (Clapp
& Sibley 197la).

The only indication of tall, native forests is given by Arundel
(who mined guano and later cleared land for coconuts) in an unpublished
manuscript to shipowners, where he states that "the trees on the extreme
northern and southern isiets (i.e. Nake, South) are about 80 to 100 feet
high" (Arundel 1875). Beginning in 1885, coconuts were planted on about
half of the motus, but the copra industry failed twice, and from 1929 to
1987 the atoll was essentially uninhabited.

Dixon made the first true botanical collection in 1883 during the
Solar Eclipse Expedition (in Trelease 1884). All specimens were from
South Island except Laportea ruderalis. His collection included several
Ornamentals and vegetables grown by early settlers (Lucett 1851) but not
reported since, an important point as these temporary introductions have
since been cited in the literature as part of Caroline’s 35 plant
species. Many were not found by the POBSP party, yet because no
scientific investigations had been conducted for 80 years, they were
counted as part of the atoll flora (Clapp & Sibley 197la). Two more
visits to Caroline, plus periodic searching by the Falconers, have also
failed to uncover most of these ornamentals.

Vascular Plants of Caroline Atoll

Plant Collections

To avoid duplicating Long’s plant collection (Clapp & Sibley
1971a), we collected only 5 specimens in 1988 and 33 in 1990.
Dr. D. Herbst assisted with identification, prepared and deposited the
specimens with Long’s in the Bernice P. Bishop Museum, Honolulu, Hawaii.
Collection numbers preceded by ‘K’ were collected by A. Kay Kepler;
those preceded by ’L’ are those of the late C. R. Long. Earlier

35

collections of Bennett in 1835 (Bennett 1840) and Dixon in 1883 (in
Trelease 1884) are noted by date only.

Working with Long’s location records for some species has proven
difficult. He was working with an incorrect map (Fig. 7), which showed
only 25 islets instead of thirty-nine. Much of his work was done at
night, which in some places would have made it hard for him to determine
his exact location. His references to South, Long, and Nake are
undoubtedly correct, and presumably the following: "second islet south
of Long" = Crescent; "islet northeast of South Island" = Tridacna; and
"fourth islet north of Bird Islet" = Emerald. Long records Pandanus on
the "second islet south of Nake Island," which lacked Pandanus when we
surveyed the island. Moreover, the first islet south of Nake supports
an extensive grove of large Pandanus trees on its eastern (lagoon)
shore, and we feel confident in ascribing Long’s specimen to this
island, which we had named "Pandanus" because of this grove. To be
consistent, we have ascribed all his other "second islet" specimens to
Pandanus Islet as well, and assume he made no collections on the actual
second islet (Danger). However, Sibley (pers. comm.), the
ornithologist, told us that the party visited every islet.

Species Lists, Annotated Checklist and Maps of Terrestrial Vascular
Plants

Following recent authors (Lamberson 1987, Sachet & Fosberg 1983),
we do not consider Caroline’s transient or extinct vascular species
(Table 1) or the vegetables and ornamentals in the Falconer’s garden as
part of Caroline’s viable flora. Table 2 summarizes the current flora,
detailing the relative abundance of each species within each plant
community. These tables are based on sight records supplemented by all
collections, past and present. No beach drift seeds are known from
Caroline apart from those species already represented. English and
Gilbertese names in Tables 1 and 2 are from Thaman (1987), St. John
(1973), and Perry & Garnett (n.d.). If no common name is available, the
Hawaiian name, familiar to many students of Pacific botany, is used.

Table 3 lists the distribution and abundance of plant species
(with subdivisions into tree, shrub and herb components) on all motus.
Figs. 11-25 map the entire atoll distribution of each species according
to data from transects and aerial maps.

Families are arranged phylogenetically, according to Fosberg &
Sachet (1987), with species arranged alphabetically within each family.
The taxonomy of vascular plants follows W. Wagner et al. (1990), and
ferns H. Wagner (pers. comm.). "% cover" means the percentage of the
ground area covered by a particular plant species. In all text and
tables the following symbols apply:

°

New record for Caroline

td Indigenous--plants native but also occurring elsewhere (1)

36

Table 1. Plants,reported from Caroline Atoll,

flora.

but considered to be transient or extinct members of the

Comments

One clump by cistern, South
Is., on recently disturbed

Introduced for cultivated fruit

Introduced ornamental. One
small specimen found on South

English and Date Last
Scientific Name Gilbertese Names Reported
CLASS ANGIOSPERMAE
Family Graminae
Eleusine indica (L.) Gaertn. goosegrass, te uteute 1884 Introduced weed
Eragrostis amabilis jovegrass, te uteute 1884 Introduced weed
(L.) H. & A
Family Cyperaceae
*Kyllinga brevifolia Rottb. kyllinga
ground
Family Bromeliaceae
Ananas comosa L. pineapple, te bainaboro 1884
Family Liliaceae
Crinum sp. lily, te kiebu 1884
Presently

Family Moraceae

*Artocarpus altilis
(Park.) Fosb.
Ficus carica lL.

Family Basellaceae
Boussingaultia gracilis Miers

Family Leguminosae

Inocarpus fagifer
(Parkins. ex Z) Fosb.

= I. fagiferus)
Vigna marina (?)

Family Euphorbiaceae
Euphorbia hirta
(= E. pilulifera)

Family Guttiferae
Calophyllum inophyllum L.

Family Caricaceae
Carica papaya

Family Cucurbitaceae
Cucurbita pepo L.

Family Convolvulaceae
Ipomoea batatas L.

1. pes-caprae brasiliensis
(L.) v. Ooststr.

Family Scrophulariaceae

Russelia equisetiformis
Schlecht.

cultivated

breadfruit, te mai Presently
cultivated

common fig, te biku 1884

Madeira vine 1884

Tahitian or Pacific 1840

chestnut, mape (Tahiti),

te ibi

beach pea

garden spurge, sleeping 1884

plant, te kaimatu

Alexandrian laurel, 1884

tamanu (Tahiti), te itai

Papaya, pawpaw, Presently

te babaia, te mwemwera cultivated

Pumpkin, te baukin, Presently

te bamakin cultivated

Sweet potato, te kumara Presently

cultivated

beach morning glory,
pohuehue (Hawaii),
te ruku

Coral plant, te kaibaun 1884

("golden plant")

Icince 1988, the Falconers have added more vege
beans, lemon grass, peppermint, okra, banana,

etc.

*Not previously reported from Caroline Atoll.

tables and ornamentals to t
Tahitian gardenia (tiare),

Is. by Anne Falconer, 1990.
Collection no. K-90-14

Not yet established, 2 trees on
South and Ana-Ana
Introduced for cultivated fruit

Introduced “vine climbing over
portico" (Trelease 1884)

Unsuccessful introduction in
1834. Food plant

"Vigna luteola in clearings on
South Island" (Line Island
Expedition Report 1974). Not
noted or collected otherwise.
Perhaps a misidentification of
Ipomoea macrantha?

Introduced weed, unsuccessful

In the 1940s, a "few taller
Calophyllum and Pisonia”
(N.1.D. 1943). No other
reference; did observer confuse
Calophyllum with Cordia?

Cultivated for fruit in 1884,
not seen in 1965. In garden on
Ana-Ana, one on South Is. by
cistern

Cultivated in 1884, not found
in 1965

Introduced in 1840, not
reported again until this
expedition (tubers brought in
1988). Collection nos. K-159,

-160

Found in 1965 by copra shed;
extensive searching on 3
expeditions in 1988 and 1990
failed to find it

Unsuccessful introduction in
19th century

heir ever-expanding garden: green
tomato, breadfruit, red hibiscus,

Table 2.

Vascular flora of Caroline Atoll:

relative abundance of each species
within the major ecosystems, with
data on seabird utilization.

Scientific Name

TREES
Pisonia grandis

Morinda citrifolia

Cocos nucifera
Cordia subcordata

Pandanus tectorius

Hibiscus tiliaceus

Thespesia populnea

SHRUBS

Tournefortia argent.
Suriana maritima

Ximenia americana

*Scaevola taccada
HERBS

Heliotropium anomalum
Boerhavia repens
Portulaca lutea
Laportea ruderalis

Achyranthes canescens

Phymatosorus scolopendria

Lepturus repens
Ipomoea macrantha

Tacca leontopetaloides

Lepidium bidentatum
Psilotom nudum

Phyllanthus amarus
Sida fallax

+Digitaria sp.
Iribulus cistoides

lexcludes transient and extinct species (Table 1).
*New records for Caroline.
*Not seen on this expedition,

Common & Gilbertese
Names

pisonia, puka tree,

te buka

Indian mulberry,
te non

coconut, te ni

sea trumpet, kou
(Hawaii), te kanawa

Pandanus, screwpine,
te aroka, te kaina
beach hibiscus, hau
(Hawaii) te rau

milo (Hawaii),
te bingibing

tree heliotrope,
te ren

bay cedar, te aroa,
te marou

monkeyplum.

scaevola, saltbush,
half-flower, te’ mao

“sand rose,” hinahina
(Hawaii)

pigvine, te wao

yellow portulaca,
seaside purslane, te
bointari, te boi

“nettle,” te ukeuke,
te nekeneke

maile-scented fern,
laua’e or Jawai fern,
te raukota, te keang

bunchgrass, te uteute

morning glory, wild
moon-flower, te ruk

Polynesian or island
arrowroot, pia (Hawaii

& Tahiti), te makemake
peppergrass
upright psilotum,

“reed fern,” te
kimarawa

‘ilima (Hawaii), te
kaura

crabgrass, te yteute

puncture vine, te

Kiebuy

Seabird
Utilization
B R
r °
e t)
e s
d t
i i
n fn
9 g
x x
x x
x x
x x
x x
Xx x

Sar ceas
ee

coz

0-UC

R-VC

R-UC

LR

Natural Ecosystems

Coastal
Baws] ate S
Geb) Gye
apt Aur ir
Cathy eeu)
h nb
S |e
SG
Cer wo
(ere) Us
uajt
bon] i
ala
R uc
c R-C
R-0
uc
0 (
vc A
VC-A 0
§
uc vc
R-C R-A
R-UC | UC-C
uc C-A
R-C
R tc
UC-A | L,R-A
R
LR

last seen 1965 (Clapp and Sibley 197la).

Inland
TFIP F
(0); 40) |) Aten)
Ue esa on:
rejoe
nos]nis
en etulinieit
f a
0
r
t
1
a

c A
R-VC R-A
uC 0
uc-C | R-VC
Cc 0
A UC-A
R-A R-A
0 L,0
Cc uc
c c

uc C-A,L
R R-UC

Anthropogenic
Ecosystem:
Coconut Woodlands
CeePhil ROL SCE PA MW
OoU se oo] ‘tient
(PAR We 2eW ake oi
on}|nonjeh

Saath ([eQhess otallud
a HEN Cc
t Tea ES<o
i pi Oh
0) o o}|ro
n monje s
t) s
e t
a
L,uCc L,uc R-C
R-C c 0
A A C-A
L,uc R
0 A
LR
L,R
uc Cc
LA
UC-A uc C-A
R-C R
Lc R-UC UC-C
0 A
R-A R-UC C-A
R
uC A
LA
LR
LR
LR
(S)

Species arranged according to their overall abundance on the atoll.

37

38

=x Aboriginal introduction--useful plants brought by Polynesians in
pre-historical times (AI)

# Recent _introduction--plants of accidental or deliberate
introduction after Western discovery of the atoll (RI)

A Abundant--the major or dominant species in a given area

VC Very common--often seen but not quite as abundantly as above

C Common--generally distributed in large numbers
UC Uncommon--observed uncommonly but >10 times in a given area
0 Occasional--here and there, often widely scattered but not
forming a major component of the vegetation
R Rare--observed 2-10 times in a given area
S Single--only one specimen observed
L Local--found only or principally in one or more restricted areas
D Drift seedling--plant derived from a water-borne seed
+ Not seen 1988-1990 but probably still present
PSILOTACEAE
* Psilotum nudum (L.) Beauv Fig.

Formerly Known Distribution: L-3233 from Nake.

Present Distribution: Cosmopolitan, common on remote islands, rare on
Caroline. K-90-15 from South. In 1965 common on wet base of Cocos only
on Nake. In 1988 and 1990 a few clumps found similarly on South in
shady, damp locations, close to lagoon, under 18 m canopy, northwest
sector.

POLYPODIACEAE

* Phymatosorus scolopendria (Burm. f) Pichi-Sermol1i Fig... 12;5Pleeoe
Phymatodes scolopendria (Burm. f.) Ching
Polypodium phymatodes L.
Polypodium scolopendria Burm. f.
Microsorium scolopendria (Burm.) Pichi-Sermolli

Formerly Known Distribution: Recorded 1840, collected 1884; L-3244, L-
3250, L-3287 from Nake, Long and South Islands.

39

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42

Present Distribution: Range extension from 3 to 1] motus. Rarely a
continuous ground cover, locally rare to abundant. Commonest on Nake,
10-80% cover. Well represented on South, especially in open areas of
the interior, where soils are moister. On other motus local
distribution varied from less than 1 to 80%; accurate mapping is
difficult. Absent from motus less than 0.6 ha in size, where habitats
cannot provide appropriate cover, moisture, and substrate.

Ecology: Hardy. Leaves burn in sun but can withstand very dry
conditions. Primarily in Tournefortia scrub, mixed forests with Pisonia
and Pandanus, or Cocos plantations. Associated with Cordia, Morinda,
Suriana. In open clearings within dying Cocos forests occurs in dense
mats intermingled with Boerhavia, Ipomoea, and Portulaca. Sometimes
gathers in thick bands at the interface of Tournefortia and Pisonia
forests. Prefers shelter from winds, high humidity, "soil," and
relative lack of wind, but absent from deeply shaded forests. Rhizomes
never exposed on ground surface or epiphytic on trunks, as in wetter
islands such as Hawaii or Samoa (pers. obs.) or in the moister Line
Islands (Wester 1985), indicating that Caroline’s habitats are
suboptimal. Although most ferns are not halophytic, this species grew
(rather stunted) in 1988 amongst sparse herb mats (1% cover) on older
beach sands of an ancient reef channel on Long Island (Tr. C), where
rainfall provides the sole fresh water, but was (temporarily ?)
obliterated in February 1990. Rare to uncommon in outer beach strand,
and beach scrub with Suriana (South, Arundel, Shark).

Substrata: Dry coral rubble, sand and gravel, rubble with sparse humus,
lagoon mud, relatively fertile humus, older beach sands.

PANDANACEAE
* **?2 Pandanus tectorius Park. Fig. 11; Pls. Ses

Formerly Known Distribution: Recorded 1840, unidentified Pandanus;
L-3227, Pandanus Islet, seen on Nake by Long.

Present Distribution: A minor plant community (Sect. G), Pandanus is
primarily associated with Tournefortia or Pisonia on the leeward motus.
Most common on Nake. Range extension from 2 to 7 motus.

Phenology: Flowers and fruit in October, March, and May.

Substrata: Variable. Prefers lagoon mud, pure sand and rubble-humus,
but survives in almost pure rubble.

43
GRAMINAE
* +? Digitaria species

Collected 1883 and recorded as ?Panicum (Digitaria) marginata. Examined
by Long, who believes it a Digitaria identical to his L-3235. Not found
by the authors.

* Lepturus repens (Forst. f.) R. Br. Figs 135. Pl. <2

Formerly Known Distribution: Collected 1883; L-3211, 3221, 3236, 3238,
3247, 3259, 3286 from Windward, Tridacna, Nake, Long, Emerald, Crab and
South Island, respectively.

Present Distribution: Many Pacific atolls. K-88-4, 5; 90-1, 2, 19 to
21, 25 from South, Tridacna, and Ana-Ana. On Caroline, range extension
from 6 to 26 motus.

Ecology: Patchy, rare to locally common. Usually in exposed herb mats
with Heliotropium, Laportea, Portulaca, and low Journefortia scrub.
Abundance 1-5% cover where not in thick patches. Occasionally inland
under Tournefortia, Cordia or Cocos, persisting as forest undergrowth.
Tufts tiny (few centimeters), dry and scrappy in exposed areas, but to
3 dm where shaded. Never in tall, upright clumps, as turf, or in the
same abundance as on the drier, filled-in equatorial atolls or islands
with sandier habitats (Christopherson 1927, Fosberg 1953, pers. obs.).

Substrata: Able to survive in coral rubble of varying coarseness, down
to high water mark, but preferred habitat is part sand. L-3286 was from
"numerous clumps under Suriana scrub on South Island," perhaps the low,
sandy portion of the northwest point (Pl. 45), our best Lepturus site.
Comparison of Arundel’s chart (1883), recent aerial surveys, and earlier
photographs indicate that several motus have altered shape since 1883.
The amount of open area on South Island has also decreased markedly
since 1883. The distribution of Lepturus parallels these changes; there
is clearly much less on South Island, and more in newly-created islet
fringes.

Since 1965 the lagoon shore of South Island has become overgrown
by Cocos, so much that both Surana and Lepturus are much less common
than previously (Pls. 39, 40). However, sand and debris will always be
shifting, so that Lepturus will move from place to place, establishing
wherever conditions permit. In the second situation, a comparison of
Plates 2 and 23 from 1883 and 1988, respectively, shows that a century
ago the lagoon-facing shores of South Island were far more open than the
dense Cocos plantations of today. The clumped grass in the foreground
of Plate 2 is undoubtedly Lepturus, probably mixed with introduced
grasses not seen since that time (Eleusine indica, Eragrostis plumosa),
and the dubious Digitaria sp.

44
PALMAE

** # Cocos nucifera L. Figsc= 145936, SilsnPilis ee,
3 wlhie 234 Gn oi ns

Formerly Known Distribution: Recorded 1840, 1884; L-3285 from South
Island, extensive groves on South and Nake, scattered on north portion
of Long.

Present Distribution: Range extension from 3 to 15 motus. Planted
groves on South, Nake, and Long; the rest derived from drift.

Phenology: Flowers and fruit year-round.

Ecology: Forms a major vegetation type (Sect. F). Primarily South and
Nake, where closed canopy forests average 21 m high.

TACCACEAE

# **2 Tacca leontopetaloides (L.) 0. Kuntze Fiiginy Dall
Tacca pinnatifida Forster

Formerly Known Distribution: Normally an aboriginal introduction on
Pacific islands, but on Caroline is first mentioned as planted in 1834
(Bennett 1840); L-3213, 3219, and K-90-7, 16 from moist muck, South
Island. L-3234, common under Cocos and numerous patches found in muck,
south end, Nake.

Present Distribution: Common in northwest South. None in flower; each
plant had 2-3 leaves, possibly dying back. None found on Nake, despite
searching the south end. Has large underground tubers, dies back, and
though cultivated, still occurs spontaneously in Cocos groves on many
atolls. Harvested by the Falconers.

Ecology: Needs fine, moist soil and shade. Though its seeds float for
months (Guppy 1906), it has not established itself on other motus in 150
years, probably due to the prevalence of rubbly substrates.

Phenology: Flowers and fruit in March and May, dies back in October.

URTICACEAE

* Laportea ruderalis (Forst. f.) Chew Fig. lS
Fleurya ruderalis (Forst. f.) Gaud. ex Wedd

Formerly Known Distribution: Reported 1840, collected 1884. L-3215
common in shady areas South Island; L-3229 scattered on exposed coral
and sand, west side Crescent Islet. L-3253 under shade of Cocos and
Pisonia on north side of Long Island.

45

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46

Distribution and Abundance: K-88-3 South Island, Transect 1, elevation
0.3 m, under old Cocos plantation, in humus and rubble. Range extension
from 3 to 32 islets (Table 3). Commonest and most widespread ground
cover, patchily distributed. Rare to locally abundant, percentage cover
from less than 1% in herb mats of tiny motus to 60% in tall Pisonia
forest. Best represented on Nake, Long, Brothers, South, Pisonia, Eitei
and Mannikiba, where coverage exceeded 50% in appropriate habitats. To
1.1 m tall on Kimoa.

Ecology: Largest (to 0.5 m) specimens found under Journefortia,
Pisonia, Cocos, or Pandanus. Tiny (1-2 cm) and tougher in sunny,
exposed sites. Halophytic, pioneering in herb mats on islets less
than 0.75 ha in size (e.g. Fishball). Optimum habitat is Tournefortia
scrub, in sunny clearings or belts behind beach scrub. Uncommon in
Pisonia forest. Occurs in both windward and leeward sites, but in
greater density leeward. Will persist through several stages of plant
succession if given adequate shade.

Phenology: Flowers and fruit in October, March, and May.

Substrata: Primarily beach gravel or coarse rubble. Also rubble-sand
mixtures; not lagoon silt.

OLACACEAE

* (#2) Ximenia americana L. Figzee

Never previously collected. K-90-170 South Island, 50-100 m north of
cistern, elevation 0.3 m, 10-20 m from coastal Journefortia fringe,
within Cocos plantation. Collected by crew of the yacht Amanita and
posted by Anne Falconer to AKK. K-90-23 and 24, single sterile shrub
2.5 m tall, southwest Motu Mannikiba, leathery leaves, arching stems 4-
5 m long. Found by John Phillips, collected by JP and AKK, identified
by D. Herbst and R. Fosberg.

Distribution and Abundance: Locally abundant in one location, about 50
bushes (3-4 m high, 2-3 m wide) spread over about 100 m. Adjacent to
indigenous scrub, on edge of Cocos plantation near old settlement.
Phenology: Flowering in July 1990.

Substrata: Moist soil (South), coarse strand rubble (Mannikiba).

AMARANTHACEAE
° * Achyranthes canescens R. Br. Fig. 16,9Plteae

Never previously collected. K-88-1 South Island, Transect 5, to 0.7 m,
elevation 0.3 m, in Journefortia fringe, coral rubble.

47

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48

Distribution and Abundance: Quite widespread, primarily in interior
scrub and forest of 19 motus (Table 3), from tiny, barely vegetated
Fishball (0.73 ha) to the largest, South (106 ha). Density variable:
from less than 1% in Tournefortia scrub to 50% local ground cover in
mixed Pandanus forest. Primarily associated with Tournefortia. May be
locally abundant in clearings in Pisonia forests, pure or mixed. Often
in a zone dividing Tournefortia and Pisonia trees, especially on Pig,
Brothers, and Nake.

Ecology: Never in natural herb mats. Needs shade but requires some
direct sun; rare in pure stands of Cocos and Pisonia. Prefers small,
sunny openings in forest or scrub. Drought-resistent and probably
partly halophytic. Dies back annually in the dry season and reappears
with winter rains (Anne Falconer, pers. comm.). To 1.5 mtall. Little
or no capacity for dispersal by sea. On other islands, seeds carried by
birds, especially fruit pigeons (Guppy 1906), but pigeons are absent
from the Line Islands. Perhaps dispersed by the Long-tailed Cuckoo
(Ellis et al. 1990).

Phenology: Flowers and fruit present in October, March, and May.

Substrata: Lushest growth in humus soils of forest interiors. Often
grows in pure rubble.

NYCTAGINACEAE

* Boerhavia repens L. Fig. 17, Pileee4
Boerhavia diffusa L.
Boerhavia hirsuta: sensu Bennett 1840
Boerhaavia species: Dixon 1884

Formerly Known Distribution: Reported 1840, collected 1884; L-3210,
3324, 3239, 3225, 3252, 3262, 3289, 3291 from Windward, Tridacna, Nake,
Long, Emerald, and South, respectively.

Present Distribution: Cosmopolitan, widespread in the Pacific. K-90-
164 and 165 from Ana-Ana. Range extension on Caroline from 6 to 33
motus (Table 3).

Abundance: Present in every habitat, leeward and windward, ranging from
less than 1 to 80% cover. Often in large patches. Best locations (>50%
cover) on Nake, Long, Windward, Pig, Brothers, Arundel, Tridacna, South,
Ana-Ana, Pisonia, and Pandanus Islets.

Ecology: Mostly found beneath Tournefortia, either in pure scrub or
mixed with Pisonia, Cordia, Morinda, Suriana, or Cocos. Not in deep
Pisonia shade; rarely in herb mats. Thick ground cover in indigenous
scrub (Shark) or within clearings in old Cocos-Ipomoea forest (South),
where it mingles with Phymatosorus, reaching a high density (Pl. 34) and
large size (rooting at nodes, vines exceeded 1 m long).

BOERHAVIA REPENS

CAROLINE ATOLL

SCALE 1:34,000
300 0 S00 1000 2000

meters

Figure 17. Transect distribution map of Boerhavia repens, Caroline
Atoll. Arrows indicate areas of highest density.

49

50

GMB Pisonia
PISONIA-CORDIA
[_] PISONIA-TOURNEFORTIA
Be] MIXED FOREST

PISONIA GRANDIS

CAROLINE ATOLL

SCALE 1:34,000
soo ° $00 1000 2000

inn! SS eS

moters

Figure 18. Entire distribution map of Pisonta grandis, Caroline
Atoll. Arrows indicate forests from 10 to 21 m tall.

51

BIRDS: Bristle-thighed Curlews fed within the Boerhavia mat in old
Cocos forests, South. Sticky fruits (3 mm long) found entangled in
preened down and adhering to contour feathers of a juvenile Great
Frigatebird (Pl. 42). Species is customarily dispersed around large
oceanic areas and within atolls by tree-nesting seabirds (Guppy 1906,
Ridley 1930).

Phenology: Small mauve flowers and seeds present in October, March, and
May.

Substrata: Coral rubble with sand or humus, rarely pure beach rubble.
Lushest growth in humus-and-guano-laden rubble clearings where Pisonia
forest once grew.

* Pisonia grandis R. Br. Fig. 18; Pls 43

Formerly Known Distribution: Collected 1884; L-3280 4 m tree, north
shore, South. Small grove, north end, Long.

Present Distribution: Indo-Pacific. Caroline range extension from 2 to
29 motus (Table 3).

Abundance: A major plant community (Sect. G), of special conservation
value.

Substrata: Occupies, and contributes to, best soils on atoll: mixture
of rubble, humus, and guano.

PORTULACACEAE
* Portulaca Jutea Solander ex Forster F. Fig. 19smPis. 54,38

Formerly Known Distribution: Reported 1840 and 1884; L-3233 and 3292,
3231, 3237, 3255, 3257, from South, Pandanus, Nake, Long, and Emerald,
respectively, in open coral, rubble, gravel and exposed areas, to 1.5 dm

high.
Present Distribution: Range extension from 5 to 33 islets (Table 3).

Abundance: Along with Heliotropium anomalum is a component of the plant
community, Natural Herb Mat (Sect. G). Widespread, predictable on coast
and former reef channels but local inland. Covered from one to 60% of
land area on almost every transect, windward and leeward, especially
facing lagoon. Best areas are Long, Transect 4 (36 m wide meadow) ;
South, north end of Transect 6 (50 m wide); Brothers, lee, almost pure
mat covering 20% ground (6 m wide); Kimoa, north side (8 m wide), 10 cm
high; Eitei, north side, 5 cm high.

Ecology: Primarily occurs along edges of motus in rubble mat and open
Tournefortia scrub, averages 12 cm high. Prominent in sparsely
vegetated areas, extending seaward to high tide level. Halophytic;
highly tolerant of sun. A flat mat in exposed areas but lusher inland,

52
rising to 2 dm tall. Generally found with Heliotropium, Lepturus,
Boerhavia, or Laportea, but may form pure mats. Uncommon in
Tournefortia scrub, patchy in clearings within Pisonia forests up to
13 m high. Exceptionally common in old Cocos groves with Boerhavia,
etc. (Pl. 34); otherwise rare or absent from closed canopy Cocos
plantations. Pinker stems found in sunny sites. BIRDS: Provides
nesting cushion for Masked Booby, Sooty Tern, Brown Noddy. On Noddy
Rock, Brown Noddies nest on a thick mat of pure Portulaca. Feeding
location for shorebirds.
Phenology: Flowers and fruit October, March, and May.
Substrata: Coral rubble and gravel, fine to very coarse. Healthier on
older sands and coral-humus.

ZYGOPHYLLACEAE
* Tribulus cistoides L. Fig. ll

Formerly Known Distribution: Collected 1884. L-3245 in open sandy area
among Journefortia shrubs, Long Island. Not seen elsewhere on atoll.

Present Distribution: Not seen on our surveys, but present in 2 sites

on west-central Long Island. K-90-161 (collected by Anne Falconer),

probably from one of same sites as 1965 collection. Flowers in March.
SURIANACEAE

* Suriana maritima L. Fig. 20; Pls. 20, 39, 40, 44

Formerly Known Distribution: Collected 1884. L-3220, shrub to 1.8 m,
east edge of Tridacna Islet.

Present Distribution: K-90-5, 6 from South Island. Range expansion
from one to 9 motus (Table 3).

Abundance: Occasional. Forms a vegetation unit, Beach Scrub with
Suriana (Sect. G).

Phenology: Flowers in March and May.

Substrata: Best sites in sand but also on coral rubble.

EUPHORBIACEAE
# Phyllanthus amarus Schum. and Thonn. F igi!

Formerly Known Distribution: Collected 1884. L-3283, herb Phyllanthus
niruri L. (Trelease 1884) to 4 dm, common on north side of South Island.

53

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“mung TTD DuDums 4O dew uoLzNGL4szSip AaAuns sazawisad pue yoasues, “O02 aunbs4

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*paqn] DoD]NZdoOd 40 dew uOLyNgisazSip ZI9sue4s| “61 sunbi 4

V3iN1 wOvVINivoOd

54

Present Distribution: K-90-10-13, herb, 2 small patches, South Island.
Limited to a few square meters in the atoll’s only weedy area, less than
10 m“ in 2 small clearings by the recently-renovated cistern, South. A
fairly common weed in the Society and Tuamotu Islands, probably arriving
with 20th century copra-cutters and perhaps again within the last 2
years. Caroline’s only established "weed" (excluding Polynesian
introductions such as Cocos).

MALVACEAE
o* (**?) (#2) Hibiscus cailiaceus iE. Fig. an

Never previously collected. K-90-8, 9 from South Island, northwest
peninsula, in Cocos plantation near old settlement and "landing," in
coral rubble and humus, 0.6 m in elevation.

Present Distribution: Two or 3 large spreading trees in heavy Cocos
shade, 10 m tall, with recumbent branches forming an impenetrable
thicket, similar in size and form to specimens in Flint’s settlement.
Since Flint was evidently first settled in 1872 (i.e. no aboriginal
population was present, see Kepler, in prep.), Caroline’s trees,
restricted to the old settlement area, are most likely recent Polynesian
introductions.

° * (**?) (#2) Thespesia populnea (L.) Soland. ex Correa Figiaaal

Never previously collected. K-90-22, 154, 155 from South Island, in
Cocos plantation and in lagoon strand, northwest peninsula, near
"landing."

Present Distribution: Two trees (10 m tall), one near the cistern, the
other in a fringe of native vegetation bordering the lagoon. The
history of this species is probably the same as Hibiscus tiliaceus.

* Sida fallax Walp. Figi: Ey

Formerly Known Distribution: Collected by Dixon, 1884, who found one
specimen.

Present Distribution: Not seen for 106 years. K-90-156, 157, 158 from
South Island, at edge of cistern, north side. One clump located in a
sunny clearing, recently enlarged by the Falconers.

CONVOLVULACEAE

* Ipomoea macrantha R & S Fig. 21; PIS. 34agmen
Ipomoea tuba (Schlecht.) G. Don

Formerly Known Distribution: L-3228 and 3293, 3242, 3251 on South,
Nake, and Long, respectively. Trailing vines, white flowers, stems to
25 m long climbing over Tournefortia, Morinda, and Cocos.

55

IPOMOEA MACRANTHA

CAROLINE ATOLL

SCALE 1:34,000
800 0 300 1000 2000

Pir ered CRE (a) Le eel (ee et

meters

Figure 21. Transect distribution map of Ipomoea macrantha. Entire
distribution is shown for South Island. Arrows indicate areas having
significant amounts of this vine.

56

History: Not collected last century, though plantation records indicate
that it was a major reason for the abandonments of the coconut
plantations: "The Pohue Vine“, which is the worst pest on the island,
was reported in 1921 to be under control" (Young ca. 1922). Today it
strangles about 54 ha, two-thirds of South Island’s plantation.

Present Distribution: Range extension from 3 to 7 motus, 5 Windward and
2 Southern Leeward Islets (Table 3).

Abundance: Forms part of a vegetation subunit, Dying Cocos-Ipomoea
Forest (Pl. 34, Sect. F). An indigenous, nonparasitic vine, abundant in
disturbed areas. Rampant growth over most of the interior of South
Island, where it forms dense tangles up to 25 m high. Less dense
thickets on southern Nake drape Pandanus, Tournefortia, Morinda, and
Cocos to 10 m. Coverage 2-5% elsewhere, except in 2 Pisonia sites,
where its coverage was 20% (Long Island, Tr. B; Windward Islet, Tr. 1).

Ecology: Lush in dying Cocos forests and mixed forest with Pandanus,
aided by relatively fertile soils, moisture, humidity, and partly sunny
clearings. Strangles all but the tallest Pisonia and Cordia. Typically
sea-dispersed to atolls (seeds germinate after floating up to 1 year in
seawater), crawls inland, progressively dropping seeds, to attain full
size in interior forests (Guppy 1906, Ridley 1930). Seeds of J. pes-
Caprae are known to be ingested by White Terns in the Marshall Islands,
perhaps as gizzard stones (Fosberg 1953). Possibly these same terns,
abundant at Caroline, once aided the seed dispersal of I. macrantha.
Also characteristic of Cocos plantations elsewhere in the Pacific
(Lamberson 1987, Stoddart & Sachet 1969, Fosberg 1965).

Substrata: Prefers humus-laden rubble, but can grow in coarse rubble
and sand, especially in leeward areas.

BORAGINACEAE
* Cordia subcordata Lam. Fig. 22, Pipes
Formerly Known Distribution: Collected in 1884. L-3213 and 326la,
3228, 3246, and 3261b on South, Pandanus, Long, and Emerald,
respectively; flowering trees to 4.5 m high in leeward coral rubble or

along lagoon.

Present Distribution: Africa to Polynesia. K-90-3 from South Island,
lagoon edge. Range expansion on Caroline from 5 to 23 motus (Table 3).

Phenology: Peak flowering November through April, fruits collected in
March and May.

emisidentified as Tuumfetta [= Triumfetta] procumbens.

CORDIA SUBCORDATA

with Tournefortia or
Pisonia

[= 4 Pisonia
ES with cocos

CAROLINE ATOLL

SCALE 1:34,000
500 t) $00 1000 2000

i aS a ey

meters

Figure 22. Entire distribution map of Cordia subcordata. Arrows
indicate small, but monotypic, stands.

57

58

Abundance: A separate, though minor, plant community (Sect. F).
Although groves are small and mixed with other emergents, individual
trees attain 13 m tall. Cordia has special conservation value.

* Heliotropium anomalum H. & A. Fig. 233 Pls. 16, 32; 45-37

Formerly Known Distribution: Recorded (mistakenly as H. curassavicum)
in 1840, collected in 1884. L-3222 and 3288, 3240, 3248, 3256, 3288 on
South, Danger, Long, and Emerald, respectively.

Present Distribution: Central and Eastern Pacific. K-90-17 from Ana-
Ana. In coral gravel, leeward and windward shores. Range extension on
Caroline from 4 to 34 motus (Table 3).

Abundance: Forms part of a major vegetation unit, Natural Herb Mat
(Sect. G), often associated with Laportea, Lepturus, or Boerhavia. Area
coverage ranges from less than 1% to 50%. Widespread, predictable on
wind- and salt-blown, low flats where vegetation does not overhang edge
of motu. Also in ancient reef channels and newly evolving land
connecting islets. Covers major areas of islets, that is those less
than 1.0 ha (e.g. Fishball, Skull, and Bo’sun Bird). Best developed on
Skull, Tridacna, South, Emerald, and Mannikiba (50% coverage, western
seaward rim).

Ecology: Halophytic pioneer. Heights to 22 cm, averaging 7 cm.
Thrives in heat and exposure.

Phenology: Flowers and fruits year-round.

Substrata: Primarily coral rubble and rubbly sand. Marginal habitats
extend down to high tide line in areas of coarse coral chunks, where it
is tiny and leathery.

* Tournefortia argentea L. Fig. 24; Pls. 9, S7peyeuae
Messerschmidia argentea (L.f.) Johnston

Formerly Known Distribution: Collected 1884. L-3216, 3226, 3241, 3249,
3258 from South, Tridacna, Nake, Long, and Emerald Isle; shrub to 3 m
high, edge of lagoon and above high tide, with white flowers.

Present Distribution: Range extension from 5 to 38 motus (Table 3).
Widespread in the Pacific, especially on small islets, but rarely
inland. Reaches Ducie Atoll, the most southeasterly island in
Polynesia. Caroline’s large tracts are excellent examples of relatively
undisturbed, pure Tournefortia scrub and forest.

Abundance: Dominates the atoll woodlands, forming the major vegetation
type (Sect. G). On almost every motu ranging from a spattering of
exposed shrubs within herb mats, through scrublands and taller forests
to 14 m high.

59

HELIOTROPIUM
ANOMALUM

CAROLINE ATOLL

SCALE 1:34,000
soo 0 $00 1000 2000

Eee

meters

Figure 23. Entire distribution map of Heltotropium anomalum. Arrows
indicate areas of highest density.

60

WJ TOURNEFORTIA
[=] “SAVANNA”-HERB MAT
fie) MIXED FOREST
TOURNEFORTIA-MORINDA

TOURNEFORTIA ARGENTEA

CAROLINE ATOLL

SCALE 1:34,000
soo ° 500 1000 2000

Ltrs ee ee

meters

Figure 24. Entire distribution map of Tournefortia argentea. Because
this shrub dominates Caroline's woodlands, there are no individual
arrows to indicate areas of high density.

61

Ecology: Supports 7 species of breeding seabirds; provides feeding
habitats for Reef Herons (Egretta sacra), shorebirds, land crabs, and
rats.

Phenology: Flowers and fruits year-round.

Substrata: Pure coral clinker; mixtures of rubble, gravel, sand, and
humus.

BRASSICACEAE
* Lepidium bidentatum Montin Fig. ll

Formerly Known Distribution: Reported in 1825: "a boat load of
pepper-grass and pursley" (Paulding 1831) and in 1835, "a Lepidium of
luxuriant growth" (Bennett 1840). Collected by Dixon as L. piscidium
Forst in 1883.

Present Distribution: Widely distributed throughout the North and South
Pacific. K-90-169 and 171 (collected by Alexandre Falconer), on
Tridacna and Pisonia, most probably in coastal Tournefortia scrub.
Entire and serrated leaf forms present, which have also been collected
on Flint (St. John & Fosberg 1937).

RUBIACEAE
* Morinda citrifolia L. Fig.5'25,. Pl. 48

Formerly Known Distribution: Reported 1840, collected 1884. L-3214,
3217 and 3282; 3232; 3254 on South, Nake, and Long, respectively.

Present Distribution: K-90-4, 18 from South’s lagoon edge and Ana-Ana,
respectively. Range extension on Caroline from 3 to 30 motus (Table 3).

Abundance: Coverage 2% to 50%. Basically an inland species, widespread
and predictable in scrub and forest understory. Rarely a canopy
component, except on Raurau, where 12 m tall in a 13 m Pisonia forest.
Associated with established Tournefortia woodlands on motus greater than
one hectare in size. Quite common on South despite major disturbance,
occurring within beach strand, Cocos, and dying Cocos-Ipomoea interior.
Best locations (40-60% coverage): Nake, Transect 3; Tridacna, both
transects; Long, Transect 8; Raurau and Ana-Ana.

Ecology: Appears early in plant succession: in Tournefortia scrub as
an early pioneer (Stage I), then from Stages II to IV, progressively
becoming more common and robust. Not in pure Pisonia forest (Stage V).
Much less common in Pandanus stands. Thrives in light to heavy shade,
preferably growing in moist substrata.

Biogeographical Note: Morinda is generally considered a naturalized
aboriginal introduction in the Pacific. Although possibly introduced to

62

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63

Caroline by early Tuamotuan settlers, its present distribution strongly
suggests that it is indigenous, as theorized for the northern Line
Islands (Wester 1985). Throughout the atoll] Morinda occurs in the
greatest densities on motus with no anthropogenic forests or in areas
distant from historical settlements (Fig. 26). Furthermore, it is
present on (30) 77% of the motus, virtually all those larger than 0.4 ha
in size, and many of which are presumed virgin. On Nake, Morinda occurs
frequently--in places abundantly--within the interior indigenous
forests, yet its coverage is only 5-10% in the mixed Cocos forests of
the southern sector. It also appears to be part of natural biological
succession (Table 6). However, on nearby Flint, which was probably
never settled in prehistoric times and where our 1990 surveys found
Morinda in all habitats (mixed woodland, native coastal scrub, Cocos
plantations, and abandoned settlement), the largest tree and highest
abundance was close to the old settlement (Kepler, in prep.).

Originating in southeast Asia, Morinda has been widely dispersed
by man, but has spread, unaided by man "widely by sea in the Malayan and
Polynesian Islands" (Ridley 1930). Fosberg (1974) notes that it is
always difficult to determine its true status. Its air-filled, buoyant
pyrenes can float for at least 53 days and "its seeds are almost
certainly disseminated by birds and bats" (Guppy 1906). It could also
be disseminated by Coenobita crabs and rats within and between motus, as
has been found elsewhere by Ridley.

Phenology: Flowers and fruits year-round.

Substrata: Coral rubble, gravel, sand, and humus. Rarely found in
coarse clinker. On larger motus, prefers moist soils under tall
forests.

GOODENIACEAE

° * Scaevola taccada var. sericea (Vahl) St. John Fig
Scaevola sericea var. sericea Vahl

Never previously collected. K-88-2, Windward Islet, central-windward
side, elevation 0.3 m.

Distribution and Abundance: One wind- and salt-sheared "hedge," found
by K. Teeb’aki on Windward Islet, was growing on a coarse rubble beach.
"The saltbush..., being recorded for the first time too from the
island...covered approximately 3% of the islet’s land area,~ occupying
the mid-windward side. The patch grew very low--only up to 2’ high with
its foliage forming an extended raised mat canopy all along the area it
occupied" (Teeb’aki 1988). We have been unable to return to this spot
to observe and photograph it directly.

3This probably translates as "3% of the area covered at that location on
the transect," as we understood from Teeb’aki’s description that it was small.

64

Because Scaevola is hardy, halophytic, and widespread in the
Pacific, it is surprising that it is so rare on Caroline. However, none
occur on Vostok, and only one clump is known from Flint (St. John &
Fosberg 1937). Fosberg (1953) noted that Scaevola seeds are transported
by Bristle-thighed Curlews (Numenius taitensis) in the Marshall Islands:
curlews are common on Caroline (Pt. II) and could have brought seeds
from elsewhere.

Substrata: Coarse rubble, windward beach.

° * Scaevola taccada tuamotensis St. John Figs
Scaevola sericea var. tuamotensis (St. John) Fosb.

Never previously collected. K-90-168 (collected by Alexandre Falconer),
northeast peninsula, South Island, in coral rubble.

Present Distribution: One individual, of unknown size, with Suriana and
Heliotropium, northeast peninsula, South Island, facing the inner side
of the "blind passage."

Floristics and Ecology of the Motus

Size of the Flora

Atoll floras characteristically lack diversity. Numbers of
species range from 1 (Ducie Atoll) to around 150 in the Pacific and 284
in the Indian Ocean. Tiny gravel banks such as Kingman Reef (Line
Islands) and Motu-One (Marquesas) have no vascular flora at all (Fosberg
1974). The flora of the Southern Line Islands is particularly
impoverished because of 1) their easterly location (far from the major
source areas of Australasia), 2) low profiles (most only rise a few
meters above sea level), 3) lack of topographic diversity (most have a
very limited range of habitats), 4) low to medium rainfall
(approximately 1,500 mm p.a.), and 5) edaphic factors such as salinity,
highly calcareous soils, etc. Long-distance dispersal and hardiness are
important factors in establishing a flora, especially since the closest
high island, Tahiti, is 830 km away, and the ultimate source of its
flora, the Malayan-Melanesian region, is over 8,000 km away. South
America, the closest continent, is approximately 9,000 km distant. The
motus of Aitutaki, for example, at a similar latitude but further west
and wetter, are considered depleted with 45 species. Fanning, at a
similar longitude but wetter, has 123 species. Tarawa, 3,900 km to the
northwest, receives a similar rainfall but supports 109 species.

Where an atoll’s potential flora is larger, the increased shade
and greater protection from wind, salt spray, and storms result in a
greater number of natural plant species on its larger motus. However,
such atolls are generally inhabited and alterations by both aboriginal
and modern man have modified their original flora. Caroline’s
isolation, variety of motu areas, and minimal human disturbance all
contribute to its excellence for the study of atoll evolution.

65

The number of species presently established on Caroline’s 39 motus
is 26 (Tbs. 2, 3). The previous expedition in 1965 (Clapp & Sibley
1971a) collected 20 species, of which 4 were new to the atoll. Their
total of 35 species, however, incorporating reports and collections from
the 1800s, is misleading. Our total, 5 of which were new records, would
have brought the atoll total to 43 (plus about 15 more unestablished,
mostly garden, plants). However, following recent custom (see Sect. E),
we have listed transient or extinct members of the flora separately
(Table 1).

The 1883 drawings of the South Island settlement, inhabited when
most of Caroline’s species were catalogued, shows that the island was
vastly different then (compare Pls. 2, 3 and 23). A century ago homes
were set amidst large grassy clearings; now the site is completely
obliterated beneath shady 21-m-tall coconut palms. Nine exotic species
have not been seen for over a century (Table 1). Evidently most
ornamentals and domestic vegetables perished during uninhabited periods.
The Falconers struggled to keep garden plants alive because of poor
soils, irregular rainfall, and foraging land crabs. This situation has
been noted for other atolls (Fosberg 1949). A few native species might
also have been eliminated during the guano and copra-harvesting years.

Numbers of Indigenous Plants

A comparison of the percentage of indigenous species between
ea island groups (Table 4) shows that Caroline, with 85%

23)“ indigenous, is unusually high. Only 12 of 45 Pacific islands
Sead have more than 75% of their species indigenous. Of these, 9
(including Caroline) are remote and lack permanent human occupation.
Vostok, Caroline’s nearest neighbor (243 km west), is one of only 2
islands in the world which have less than 4 species (Fosberg, pers.
comm.) .

The Tuamotu Islands (149° to 134°W) lie east and south of
Caroline, yet they harbor considerably larger floras. Rainfall is
similar. Three of them average 121 species (Table 4), averaging 42
indigenous species. When the variables rainfall and distance from a
colonization source to the west are considered, the proximity of the
Tuamotus to the diverse high islands of the Societies seems to play a
major part in determining their indigenous flora. A similar situation
exists in the southern Cook Islands. Caroline and other remote Line and
Phoenix Islands are sufficiently isolated from high volcanic and raised
reef (makatea) islands that they exhibit a much simpler flora. Tahiti,
the closest high island (830 km south), is in the wrong direction for
prevailing currents, winds, or vagrant birds to bring seeds to Caroline.

4 Perhaps as high as 92%; the Digitaria sp., if still extant, is of
unknown identity and origin.

66

Table 4.

Sizes of Pacific a

indigenous plants.

Island Group

Caroline Is.
(Fed.
States of
Micronesia)

Cook Is.
(New
Zealand)

Gilbert Is.
(Rep. of
Kiribati)

Northwest
Hawaiian Is.
(UESSA™))

Line Is.
(Kiribati)

Marshall Is.
(Fed.
States of
Micronesia)

Atoll

Kapingamarangi

Aitutaki
(motus)
Rarotonga
(motus)

Onotoa
Tarawa

Kure

Laysan

Caroline

Christmas
(Kiritimati)

Fanning

Flint

Malden
Palmyra
Starbuck

Vostok
Washington

Ailuk
Arno
Enewetak
Jaluit

Jemo
Kwajalein

Lae
Likiep

fol

species indigenous.

Total@
No.
Species

98

109

No.
Species
Indigenous

38

50
28
23
27
23(247)
19
23
18
9

21
4

%

Indigenous

39

Vl

1 floras, with emphasis on the percentages of
Islands in bold have more than 75% of plant

Source

Niering 1962

Stoddart &
Gibbs 1975

Stoddart &
Fosberg 1972

Moul 1957
Catala 1957

Lamoureux 1961,
Clay 1961

Ely & Clapp
1973

This paper
Garnett 1983

Wester 1985
Kepler

(in prep.)
Garnett 1983
Wester 1985
Garnett 1983

(incomplete,

little known)
Kepler 1990c
Wester 1985

Fosberg 1955
Hatheway 1953
Lamberson 1987
Fosberg &
Sachet n.d.
Fosberg 1955
Fosberg 1955,
1959

Table 4. Continued.

Island Group Atoll
Marshall Is. Taka
(cont. ) Ujae
Ujelang
Utirik
Wotho
Phoenix Is. Kanton
(Kiribati) (Abariringa)
Birnie
Enderbury
Nikumarero
Orona
McKean
Phoenix
Manra
Society Is. Tetiaroa
Solomon Is. Ontong Java
Tokelau Is. Nukunono
(NG 745)
Tuamotu Is. Rangiroa
(France)
Raroia
Takapoto
Outlyers Clipperton
(U.K.)
Ducie
Oeno
Wake
>An updated version of Table ll, p.

usually included, but certain ornamentals may not be.

Species
23
61
50
55
40

164

17
94

No.
Species
Indigenous

18
32

29

26
28

20

%

Indigenous

100

67

Source

Fosberg 1955
Fosberg 1955,
1959

Degener &
Gillaspy 1955,
Garnett 1983
Fosberg &
Sachet (n.d.)

Sachet &
Fosberg 1983

Bayliss-Smith
1973
Parham 1971

Stoddart &
Sachet 1969

Doty 1954
Sachet 1983

Sachet 1962

Rehder &
Randal] 1975
St. John &
Philipson 1960
Fosberg &
Sachet 1969

105, Stoddart and Gibbs (1975).
Number of species of those indigenous are not always comparable. Ferns are

Artocarpus, Morinda,
and Pandanus may be indigenous, aboriginal introductions, or both. Without
its full scientific name, a species has an unknown biogeographical status.

68
Composition of the Flora (Tbs. 2, 3)

Caroline’s botanical affinities lie with other Southern Line
Islands and the Tuamotus. Although the strand and inland floras consist
of pan-Pacific or pan-tropical species, there are several widespread
species and communities that are notably absent (see below). Those that
survive have withstood the atoll tests of time--poor soils, scarcity of
fresh water, periodic inundation by salt water, intermittent cyclonic
storms and hurricanes, harsh climate, high seedling mortality, and human
impacts. Caroline provides an excellent ecological laboratory in which
floristic correlations with variations in habitat, motu size,
vegetational zonation, and leeward/windward aspect may be studied.
Fosberg (1985) and Sachet (1967) have noted the importance of such
details in understanding the biogeography and taxonomy of Pacific
plants.

Caroline’s present established flora includes very few introduced
species: ancient Polynesian introductions (Cocos, possibly Pandanus and
Morinda), recent Polynesian introductions (Hibiscus tiliaceus, Thespesia
populnea, Tacca leontopetaloides, Ximenia americana), and 20th century
exotics (Phyllanthus amarus). This latter is restricted to one tiny
patch <2 sqm in area.

The number of indigenous plants is complicated by the fact that 3
species most likely introduced by recent Polynesians (Hibiscus,
Thespesia, Ximenia) could also be indigenous, as is the case with
Pandanus and Morinda. In Table 4 we have counted these 5 as indigenous
until later research proves otherwise. The unknown Digitaria sp.
accounts for the query in Table 4.

Trees: Seven species present. Only 3--Pacific-wide natives--are
widespread: Pisonia grandis, Morinda citrifolia, and Cordia subcordata.
Cocos nucifera and Pandanus tectorius are locally abundant, while
Thespesia populnea and Hibiscus tiliaceus are rare and limited to the
old settlement site. The absence of typical Pacific species such as
Calophyllum inophyllum and Guettarda speciosa is notable, as they are
both present on nearby Flint (with a similar plantation history to
Caroline) and occur naturally on more easterly atolls such as Rangiroa
(Stoddart & Sachet 1969).

Shrubs: Four species present, at least 4 indigenous. Only Journefortia
argentea is abundant; its most abundant size class is under 4 m.
Scaevola and Suriana, tough and widespread elsewhere, are poorly
represented on Caroline. It is noteworthy that 2 varieties of Scaevola
taccada are present. Ximenia americana is represented by a single,
large patch and one individual on South and Mannikiba, respectively.
Pemphis acidula, though common on atolls of similar latitude and
climate, is typically absent from most of the Line and Phoenix Groups
(Stoddart & Gibbs 1975, Fosberg & Sachet n.d.). This may be due to the
paucity of its preferred habitats: low rocky substrates (reef or
conglomerate rock) and sand-gravel ridges.

69

Herbs: Fifteen species present, at least 12 indigenous. Of these only
7 are common: Heliotropium anomalum, Boerhavia repens, Portulaca Jutea,
Laportea ruderalis, Achyranthes canescens, Lepturus repens, and
Phymatosorus scolopendria. Ipomoea macrantha and Tacca leontopetaloides
are locally abundant, while Phyllanthus amarus, Tribulus cistoides,
Lepidium bidentatum, and Psilotum nudum are rare and localized.
Digitaria sp. may be extinct. The fact that Sida fallax has only been
recorded twice in 106 years is curious.

F. ECOLOGICAL SUCCESSION

We have attempted to trace the development of Caroline’s flora
from the smallest to largest motus, using field data and aerial photos,
which reveal past geological processes of unknown dates or duration.
The general processes involved in motu formation are treated in Stoddart
& Steers (1977, p. 95).

Three tables provide our analysis of ecological succession:
Table 5 presents Caroline’s motus in order of ascending size, together
with the numbers of plant species and major plant communities. Since
the atoll’s total land area is small, our data provides relatively
complete floristic lists for each islet and detailed maps of their plant
communities (Figs. 37-57). The number of species varied from 3 growing
on 4 tiny islets (0.02 ha each) to 23 on South (104.41 ha). Because the
total number of species for the entire atoll (27) is also small, the
addition of one or 2 rare species contributes significantly to the total
flora. Such additions must be kept in perspective when evaluating plant
succession.

Table 3 provides a summary of plant species distribution by islet,
and Table 6 is a summary of plant species distribution and relative
abundance with respect to islet area and the primary mode of seed
dispersal.

Basic Seral Stages

Islets appear, grow, mature ecologically or vanish in violent
storms. Many interacting factors, including geographical (islet area,
atoll shape, distance from high islands and continents), geological
(changes in sea level, reef growth and destruction), chemical (nitrates
from bird droppings, leaf fall, etc.), climatological (wind, droughts,
storms, microclimates), and biological (seabirds, rats, land crabs, reef
bioerosion, and man, both aboriginal and modern) constantly interact to
change conditions. The relative influence of some of these factors is
evident when comparing the floras on motus of different sizes.

Seed-dispersal mechanisms (Table 6) and the presence of
underground fresh water are also vital. Unfortunately, the
relationships between groundwater salinity, species distribution, and
vegetation patterns on atolls are poorly understood (Fosberg 1985). The
presence and relative salinity of permanent water depends on

70

Table 5. The distribution of plant subcommunities, together with the numbers of plant species on the motus
of Caroline Atoll. Motus are arranged according to increasing area, illustrating seral stages in
plant succession.

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Skull Islet 0.02
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Reef-flat Islet 0.09

Azure Isle
Motu Nautonga
Scarlet Crab
Fishball Islet
Motu Kota

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Booby Islet
Bo’sun Bird Islet
North Arundel Islet
Motu Mouakena
Motu Eitei

Coral Islet

Motu Matawa
North Brothers Islet
Motu Kimoa

Lone Palm Islet
Motu Ana-Ana
Pisonia Islet
Blackfin Islet
Danger Islet
Crescent Islet
Motu Raurau

Bird Islet
Brothers Islet
North Pig Islet
Pandanus Islet
Pig Islet

Arundel Islet
Shark Islet
Emerald Isle
Tridacna Islet
Windward Islet
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72

Ghyben-Herzberg lenses of varying thickness on different islets, and
this in turn depends upon island dimensions (especially width), soil
porosity, rainfall, tidal fluctuation, and other hydrological factors.
Though groundwater supplies have been studied on many atolls (Wiens
1962, Maude 1953), each island group is so unique that it is unwise to
extrapolate information from one to the other.

Caroline’s 39 motus fall naturally into 4 size classes: motus
with areas of: a) <0.2 ha, b) 0.2 to 0.7 ha, c) 0.8 to 25.0 ha, and
d) >25.0 ha. These size groupings harbor all 5 of the seral stages
identified on Enewetak Atoll (Lamberson 1987), tailored to reflect
Caroline’s particular geography, geology, and impoverished flora. Each
stage may be the sole example of ecological succession on an islet or
may occur as one of several stages. Typically the early stages cover
the peripheral rubble and scrubby outer zones, while the later ones
appear as a series of roughly concentric bands progressing inland.

Stage I Early pioneers on sandbars, spits or small rubbly islets
subject to storm damage and washover. Harsh conditions,
intense sun, drying winds, salt spray. High salt
concentration in the substrate. Lack of fresh water and
nutrients. Plant genera present include Heliotropium,
Portulaca, Lepturus, Boerhavia, and seedling or open
Tournefortia scrub. No Cocos. This stage covers many smal]
motus (Noddy Rock, Fishball) and former interislet channels
(e.g. Long Island) or occurs peripherally on larger motus.

Stage II Thick scrub of mixed genera, often impenetrable. Its
protective barrier allows for the development of vegetation
on the larger islets. Seabirds begin to contribute to the
soil (guano, eggs, regurgitated fish, decaying nesting
material). Plant genera include Journefortia, Suriana,
Cordia, and Laportea. If Cocos present, accompanied by
coconut crabs. Very common around the periphery of most
motus just inland of the natural herb mats or flanking
sheltered shores adjacent to the lagoon (South, Kota).

Stage III Trees larger, seabirds add further to soil fertility. Open
grassland may develop in sunny clearings (Tridacna Islet).
Added plant communities are Cordia-Tournefortia,
Tournefortia-Morinda, and Pisonia-Tournefortia forests.
Occurs in the next inner concentric zone of vegetation to
Stage II on larger motus (Nake, Long) or, more commonly, the
entire interior of smaller ones (Pandanus, Southern Leeward
Islets).

Stage IV Pisonia dominates the older mixed forest. Morinda and
Tournefortia mature. Forests are more open. Undergrowth
mostly a ground cover of Laportea, Boerhavia, Lepturus and
Portulaca. Covers the main portion of larger islets. If
Cocos and Pandanus present, forms a mixed forest with vines
(southern Nake, Shark). Coconut crabs common. A widespread

73

stage in the center of most motus (Central Leeward, Windward
Islets).

Stage V Pisonia takes over. Other trees are confined to the forest
edges. Always in the deep interior of the larger islets.
Little or no ground cover. Abundant nesting Black Noddies
(Anous minutus). A more restricted stage (Brothers, Raurau,
central Nake, Pig).

Ecological Succession on Motus of Different Size Classes

To assist discussions of succession on Caroline’s motus, see
individual vegetation maps and graphs (Figs. 27-57) and photographs
(Pls. 13-70). Figures 27-30 summarize the amounts of each islet’s
surface covered by each major plant community and provide the numbers
and percentages of indigenous species for each islet.

1) Motus with Areas <0.2 ha Figseaevnesilisy Pll: 495 Tbs: 45, 6

Caroline has 4 motus in this category, 3 windward and one leeward,
whose combined area totals 0.15 ha. There are also 3 incipient islets
which, because of their temporary character, have not been counted in
Caroline’s overall total (Fig. 2, Pl. 15). With the exception of Noddy
Rock, a jagged, upraised limestone plateau, all consist predominantly of
coarse coral rubble (75-98% coverage). These tiny motus are the
simplest ecosystems on the atoll, representing early Stage I in plant
succession. The number of plant species per motu averages 3, all hardy,
sea-dispersed and salt-tolerant pioneers (Heliotropium, Portulaca,
Lepturus, Tournefortia). The sole plant community is a natural herb mat
of varying thickness and extent. Tournefortia, though stunted and
scattered, is not sufficiently common to form a separate scrub habitat.
Indigenous vegetation covers 2 to 25% of the islet areas. Seabirds,
especially Brown Noddies and Red-tailed Tropicbirds, may attempt to
nest.

2) Motus with Areas 0.2 to 0.7 ha BIGSe02o.ol3. 0DS. 5, 6

There are 5 leeward motus in this category, whose combined areas
total 2.21 ha. Their vegetative cover is more extensive and diverse
than in size class a, with herb mats and Journefortia scrub and forest,
but open rubble is still abundant (30-55% cover). Plant succession
corresponds to late Stage I and Stage II. The average number of species
is 8.2 (range 6-11), one-third of Caroline’s total. All vegetation is
indigenous except for a few Cocos palms. Seeds are dispersed by sea,
wind and birds.

With the appearance of shrubs, the number of species increases
markedly, and woodlands, primarily of Tournefortia, form and expand to
create dense thickets averaging 5 m tall and covering 25% of the land
area. Canopies of 10 m occur on Motus Nautonga and Kota. Seabird
colonies of up to 6 species (Brown and Red-footed Boobies, Great
Frigatebirds, Black and Brown Noddies, White Terns) are present.

74

LEFT CIRCLE RIGHT CIRCLE

MAJOR PLANT COMMUNITIES PERCENT INDIGENOUS
UNVEGETATED INDIGENOUS

BM Hers mat [__] ABORIGINAL INTRODUCTION
(__] TourNerorTa ===] RECENT INTRODUCTION
PISONIA

3 cocos

SURIANA

FEB: PANDANUS

HOUSE SITE

MMM sxe FOREST WITH COCOS

3
(100%)

3
(100%)

3
(100%)
|
3
(100%)

Fig. 27. Plant communities and indi j
sat Sees sere Soak ay fe vegetation on motus less than 0.1 ha. Caroline Atoll. The left ~pie™ depicts the relative amount of amotu’s
Siecacin gan ave a Hs 3 plant community. numbers indicate actual arca in hectares. The right “pie™ depicts the numbers and percentages of
species per motu. Data is based on the vegetation maps for each motu (Figs. 37-57) and Tables 2 and 9. ‘

0.02 ha

0.09 ha

1.01 (5%

AZURE ISLE
0.20 ha
0.11 0.04
(55%) (20%)

MOTU NAUTONGA

0.34 ha cau

0.13 (38%)
0.02 (6%) 70.08 (24%)

SCARLET CRAB ISLET
0.46 ha

0.11 (24%

0.18 (39%)

FISHBALL ISLET
0.57 ha

0.16 (28% 0.24

(42%)

MOTU KOTA
0.64 ha
0.31
(49%)

6
(100%)

ii
(100%)

(89%)

8
(100%)

WS

Fig. 28. Plant communities and amount of indigenous vegetation on motus 0.2 to 0.7 ha, Caroline Atoll. See Fig. 27 for explanation of the figure.

76

BOOBY ISLET
0.84ha 0.12
(14%)

0.41
(49%)
0.27
(32%)

0.04
(5%)

BO'SUN BIRD ISLET
0.86 ha

0.41
(47%)

NORTH ARUNDEL ISLET

0.91 ha
0.33 0.27
(36%) (30%)

Fig. 29a. Plant communities and amount of indigenous vegetation on motus 0.8 to 25.0 ha, Caroline Atoll. See Fig. 27 for explanation of the figure.

9
© 00%)

4
«> 00%)
10
E> j

Uh

MOTU MOUAKENA
1.00 ha 8

0.36 (100%)
(36%)
MOTU EITEI
1.41 ha a p
hier (100%)
(27%)
Nase :
1.07 ha Ss 9
5 (100%)
0.72 &
(42%) Ee Be

0.13 (8%) 0.07

(4%)
MOTU MATAWA
1.71 ha 9
0.52 2
0.68 (30%) oe
(40%)
Fs Sees
Rana p 0.44
0.07 (4%) (26%)

Fig. 29b. Plant communities and amount of indigenous vegetation on motus 0.8 to 25.0 ha, Caroline Atoll. See Fig. 27 for explanation of the figure.

78

NORTH BROTHERS ISLET
1.71 ha

0.68
(40%) 9

12
(100%)

LONE PALM ISLET 0.39 (9%)
1.99 ha

0.43
(25%)

(287 ica)

MOTU KIMOA
1.80 ha

0.50
(28%)

0.95
(48%)

MOTU ANA-ANA (7%)

2.16 ha
A | 44
| ] (93%)

Fig. 29c. Plant communities and amount of indigenous vegetation on motus 0.8 to 25.0 ha, Caroline Atoll. See Fig. 27 for explanation of the figure.

0.93
(43%)

0.21 (10%)

79

PISONIA ISLET 1 (7%)
2.45 ha 0.75
(31%)
13
0.15 (93%)
(6%)
BLACKFIN ISLET 0.63 1 (11%)

2.62 ha

8
(89%)

DANGER ISLET
2.71 ha
1.25 iG
(47%)
(100%)
CRESCENT ISLET 0.61
3.10 ha
1.56 10
(50%) (100%)

Fig. 29d. Plant communities and amount of indigenous vegetation on motus 0.8 to 25.0 ha, Caroline Atoll. Sce Fig. 27 for explanation of the figure.

80

MOTU RAURAU 1.71
3.48 ha

1.07
(31%)

0.04 «0.66

(1%) (28%)
BIRD ISLET 1.71
4.05 ha (42%)

0.22 (6%)
be >”'0.41 (10%)
(42%) 0. ot (0.03%)

BROTHERS ISLET

4.31 ha Ose
2.0
(47%)
0:37 a
(8%) (9.2%)
NORTH PIG ISLET 0.97
5.44 ha (18%)
1.31
(24%)

(24%)

Fig. 29e. Plant communities and amount of indigenous vegetation on motus 0.8 to 25.0 ha, Caroline Atoll. See Fig. 27 for explanation of the figure.

(10%)

9
(90%)

1
(8%)

be
7

(1 ous

PANDANUS ISLET Bs
7.20 ha ¥:
0.53
(7%)
2.09
(28%)
2.28
(32%)
1.61
PIG ISLET (23%)
7.25 ha

3.36 %
(46%)

0.03 (0.4%)

ARUNDEL ISLET
7.34 ha

4.36
(59%)

SHARK ISLET :
7.98 ha (ag)
2.92
(37%)

0.12 (1%)
2.60 (33%)

81

10

(100%)

©

(9%)
10
(91%)
11
(100%)
;
(8%)
11
(92%)

=

Fig. 29f. Plant communities and amount of indigenous vegetation on motus 0.8 to 25.0 ha, Caroline Atoll. See Fig. 27 for'explanation of the figure.

82

EMERALD ISLE 1.5 (18%)
8.34 ha
3.20
(38%) 1.82
(22%)

0.69 1.13
(8%) (14%)

TRIDACNA ISLET

9.08 ha
0.18 (2%)

7.97 0.21 (2%)
(88%) 0.72 (8%)

WINDWARD ISLET
11.42 ha
5.7,
(50%)

MOTU MANNIKIBA

21.49 ha
2.60
15.69 M27)
(73%)

= 2.04 (10%)
90.03 (0.1%)

1 (8%)
1
(92%)
11
(100%)
11
(100%)
1 (8%)
12
(92%)

S

Fig. 29g. Plant communities and amount of indigenous vegetation on motus 0.8 to 25.0 ha, Caroline Atoll. See Fig. 27 for explanation of the figure.

83

14
LONG ISLAND aa
75.90 ha 18.6 (25%) (93%)
32.2
(42%) ,
=) 7.7 (10%) (7%)
2.4 (3%)
15.0
(20%)
SOUTH ISLAND 4.2 (4%)
104.60 ha SSS 41 (1%)
13.6 (13%)
4.9 (5%)
80.8
(77%)
2
NAKE ISLAND (13%)
91.71 ha 18.3
30.65 (20%)
(33%) on
87%
6.8 (7%) (87%)

20.8
(23%) 5.75

Fig. 30. Plant communities and amount of indigenous vegetation on motus over 25.0 ha. See Fig. 27 for explanation of the figure.

84

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A low herb mat, dominated by Heliotropium, Portulaca, Boerhavia,
and, more rarely, Lepturus, develops first, after which Tournefortia
quickly becomes established. Shade, producing locally humid conditions,
and better "soils" derived from guano, decomposing leaves, and the
activities of land crabs and rats, provides appropriate habitat for
Laportea and occasional Phymatosorus and Achyranthes. The major tree
species--Pisonia, Morinda, Cordia, and Cocos--subsequently appear, but
are relatively rare. Pisonia, typically an inland species assumed to
need underground water (Spicer & Newbery 1979, Wiens 1962), could well
be salt tolerant as it occurs on motus as small as 0.2 ha (Tbs. 5, 6).
In this size class Pisonia occupies only 2-6% of the total islet areas.

3) Motus with Areas from 0.8 to 25.0 ha Figs.\292 13137 1bs.15;°6

All 27 motus in this category share a similar complement of
species and plant communities (Tbs. 5, 6). Their combined area totals
124.35 ha. They are well-wooded (Fig. 29), although the leeward motus
have a higher proportion of rubble and herb mats, and forests are higher
to windward. Unvegetated rubble covers less land area (21%) than in
size classes a and b (87% and 39%, respectively). Within the woodlands
of these motus, substrates mature from basic rubble to primitive "soils"
with small, but significant, structural development. Their flora shows
increasing diversity with size, and almost the full complement of
seabirds may nest.

All natural ecosystems are firmly established; canopy heights
range from 4 to 21 m. On small Booby Islet (0.84 ha), Pisonia is very
common, and the Pisonia forests on North Brothers (1.71 ha) and Pig
(7.25 ha), at 21 m, are the tallest on Caroline. As rich guano and dead
foliage accumulate, a layer of phosphate-rich humus enables those
species already present but poorly represented on the small motus
(Pisonia, Morinda, Boerhavia, Laportea, Achyranthes) to increase in
abundance and stature (Table 6). Additional species are Suriana,
Pandanus, Ipomoea, Lepidium, and Ximenia.

Plant succession, ranging from Stage III to Stage V in the
interior, primarily involves forest maturity rather than the addition of
large numbers of species. On the larger islets, the number of plant
Species increases by relatively small increments, filling out the sub-
canopy layers and, in the cases of Cordia and Pandanus, adding variety
to the canopy.

The average number of plant species is 11.0, ranging from 4 to 15.
If we divide the motus into smaller size classes, we find that their
species numbers show a slight overall increase with increasing size:
8.0 species for areas 0.8-1.0 ha, 9.8 species for areas 1.1-2.0 ha, 11.5
species for areas 2.1-4.0 ha, 11.3 species for areas 4.1-10.0 ha, and
12.0 for areas 10.1-22 ha. An increase in herbs (range 3-9) is
primarily responsible for these higher averages (Table 5).

Despite the large range of motu sizes in this category, plant
communities are essentially natural (Table 5). Their overall species
composition is 96% indigenous. Seventeen of the motus lack Cocos, the

86

only introduced species in this area category, which is represented by
small, isolated clumps or individual palms.

On the larger motus, and within the taller forests, more species
of birds, especially Red-footed Boobies, Great Frigatebirds, White
Terns, and Black Noddies, nest in increasingly large colonies,
furnishing more minerals to the developing soils, especially where
Pisonia covers large areas.

In summary, by the time a motu on Caroline has reached 0.8 ha in
size, al] the natural plant communities, most species of trees, shrubs
and herbs, and most species of seabirds are present, assuming no major
intervening disturbances (for example, hurricanes) have occurred. In
Caroline’s depauperate flora there are few species left to increase
floral diversity on the larger islets, regardless of their size. This
is very different from the inhabited atolls such as Kapingamarangi (see
next section).

Although we do not know when true freshwater lenses develop, they
may occur in motus of this size class. If we assume that Pisonia is not
specially salt-tolerant, limited fresh water must be available on motus
as small as 0.2 ha, and actual freshwater lenses may begin forming at
ca. 0.7 ha, as indicated by the sudden proliferation of Pisonia forest
(Tbs. 5, 6). However, the Falconers were unsuccessful in locating
underground fresh water on Motu Ana-Ana (2.16 ha), which suggests that
Pisonia may be somewhat salt-tolerant.

4) Motus with Areas >25.0 ha Figs. *30;,3ls0ibseaoge6

On Caroline no motus fall between 22 and 75 ha in size. Thus the
3 motus in this category (Nake, South, Long) cover a limited range:
75.98 to 107.50 ha. They average 18.0 plant species. The floral
components and forest heights of these larger motus (Figs. 33, 34;
Table 5) are essentially the same as for class c. There are no
additional natural ecosystems (mangrove swamps, salt flats, grasslands,
etc.) or understory layers. One additional plant community (3
subcommunities) exists, dominated by the introduced coconut palm. Ten
species, all rare or uncommon, are present only on the larger motus
(Table 3): Scaevola, Tribulus, Hibiscus, Thespesia, Ximenia, Psilotum,
Tacca (introduced in 1834), Phyllanthus, Sida, and the dubious
Digitaria. Four, possibly as many as 8, are indigenous. In 1965, one
vine of the indigenous Ipomoea pes-caprae was also found, but 3
subsequent surveys failed to locate it.

Species-Area Relationships

The relationship between the numbers of plant species and island
size has long attracted interest (Fosberg 1949, Wiens 1962, MacArthur &
Wilson 1967, Whitehead & Jones 1969), yet data from uninhabited islands
is scant. The studies from Kapingamarangi (Niering 1956, Wiens 1956)
and Aitutaki (Stoddart & Gibbs 1975) treat atolls with long histories of
human occupancy. Some of the villages on Kapingamarangi’s 23 motus date

87

to 1200 A.D. Aitutaki’s 16 uninhabited motus lie adjacent to a
westernized volcanic island in an "almost-atol]." People on both these
atolls have profoundly influenced their flora.

Caroline provides an opportunity to compare the numbers of species
on motus of different sizes in an uninhabited atoll, then to compare the
results with Kapingamarangi, Aitutaki, and uninhabited islands in the
Line and Phoenix Groups that have no introduced species and have
experienced minimal human contact (Table 8). Comparison of Caroline
with remote, uninhabited Oeno and Ducie Islands, also having entirely
indigenous flora (Fosberg et al. 1989) would also be beneficial
(T. Spencer, pers. comm.).

Comparisons of Species-Area Relationships with Other Atolls

Studies of Kapingamarangi (Niering 1956) contributed greatly to
theories of island biogeography (MacArthur & Wilson 1967). Because its
motus cover the same range of sizes as Caroline, the 2 atolls might be
expected to exhibit similar patterns. However, their species-area
relationships are completely different. On Kapingamarangi, islets less
than 1.4 ha showed a constant, small number of species, after which
islets up to 100 ha showed a direct correlation of area with numbers of
species. On Caroline, a motu of 1.4 ha supports almost two-thirds of
the total number of species, and plant diversity on islets up to 107 ha
shows only a slight, but not necessarily steady, increase (Table 5).

Species-area relationships on the motus of Aitutaki (Stoddart &
Gibbs 1975, Figs. 33 and 34 of that paper) conformed to the Caroline
model: the number of species increased only slightly on motus from 4 to
71 ha. Unfortunately, Aitutaki had only one motu less than 1.4 ha, so
comparisons for smaller islets cannot be made. The floras of all 3
atolls have been impacted by man, but Caroline far less so than the
others. Much of the floral diversity on larger islets at Kapingamarangi
is derived from plants introduced by man and cannot be considered
natural.

Six islands in the Line and Phoenix Groups (Malden, Starbuck,
McKean, Phoenix, Vostok, Birnie) are uninhabited, with entirely native
flora. All are Caroline’s "neighbors" in an oceanic sense, and all
except Vostok are dry, receiving about 750 mm (30") of rain p.a. They
are old, essentially filled-in atolls, containing hypersaline central
lagoons or no lagoon at all. Although the largest island (Malden) has
the greatest diversity, there is only a very small linear increase in
plant species with increasing area (Table 7). Plant diversity is more a
function of climate (hot and dry) and distance from source areas, than
size, similar to the situation on Caroline.

The Question of Fresh Water

The Kapingamarangi data were analyzed with availability of fresh
water in mind (Wiens 1962, Whitehead & Jones 1969). These authors

88

Table 7. Species-area relationships of six Pacific islands with
entirely indigenous flora.

Island Area No. Species
Malden 39.3 sq km 9
Starbuck 16.2 sq km 6
McKean 57 ha 7
Phoenix 49 ha 6
Vostok 24 ha 3
Birnie 20 ha 4

Islands are arranged according to decreasing area. Data is from Garnett
(1983), Fosberg and Sachet (n.d.), Clapp and Sibley (1971b), and pers.
obs.

suggested that 1.4 ha is the threshhold at which a freshwater lens can
develop. Below this size only halophytes can survive. They argue that
as there are only a limited number of salt-tolerant species, the floral
composition on islets below 1.4 ha is relatively constant. On larger
islets, species numbers increase in direct proportion to land area,
because permanent groundwater promotes the survival of an increasing
variety of nonhalophytic plants.

The groundwater vs. plant model does not apply to depauperate
Caroline for a number of reasons: first, the number of plant species is
not constant on islets below 1.4 ha: in fact, species are added faster
on motus from 0.02 to 1.4 ha than between any other size range.

Second, on Kapingamarangi, the number of species increased in
direct relation to islet size from 1.4 ha to 100 ha. On Caroline,
species numbers increased only slightly from 1.4 to 22 ha and exhibited
another minor increase from 70 to 108 ha (see Fig. 31; Tbs. 5, 6; and
Sect. F, Ecological Succession). Thus, Caroline’s data do not support
the area-diversity theory.

Third, Whitehead and Jones (1969) argue that the flora on "small]"
motus lacking a freshwater lens (i.e. <1.4 ha) consists only of salt-
tolerant strand species. This is not true on Caroline (Table 6). In
addition to harboring the usual strand species (Tournefortia, Portulaca,
Laportea, Heliotropium, Boerhavia, Lepturus), Caroline’s "small" motus
also support inland species that are generally considered non-halophytic
(Pisonia, Morinda, Achyranthes, Cordia, Phymatosorus). Either these
latter 5 species are moderately salt tolerant, or on Caroline the
neon islet size with a freshwater lens is much less than 1.4 ha, or

oth.

Fourth, Whitehead and Jones (1969) postulate that the non-
halophytic species are those that control overall species-area

89

associations. This may be a good generalization for less remote
islands, but does not hold up for atolls with depauperate floras

(Table 6). For example, on Caroline the halophytic Ipomoea macrantha,
I. pes-caprae, Scaevola taccada, Sida fallax, Lepidium bidentatum,
Hibiscus tiliaceus, Thespesia populnea, and Tribulus cistoides, which
theoretically should only occur as strand species on the smaller islets,
occur only on larger islets.

Fifth, the authors do not mention bird-dispersal of seeds, which
is probably a factor that needs to be taken into account on remote
islands: at Caroline, Pisonia and Boerhavia occur on islets from 0.2 ha
to 108 ha. In addition, the numbers and diversity of seabirds and
migrant shorebirds are much greater on unoccupied islets/atolls than on
inhabited ones, strongly affecting the distribution of certain plant
species.

Sixth, Caroline does not have an assemblage of nonstrand plants
that only occur on larger motus; the only naturally occurring, nonstrand
plant is Psilotum.

Seventh, the greatest factor complicating our understanding of
Kapingamarangi’s natural evolutionary processes is the presence of
numerous exotics: of its 98 vascular plants, only 38 (39%) are
indigenous. Its exotics include numerous weedy herbs and food plants
which occupy gardens, abandoned house sites, taro patches, and
plantations (Cocos, Pandanus, Artocarpus). These man-made habitats are
particularly prevalent on larger islands. Such an abundance of exotics,
both in species and area covered, renders a discussion of natural
processes on Kapingamarangi almost impossible. Relatively undisturbed
habitats such as those on most of Caroline’s motus, and on other
uninhabited Pacific islands such as Ducie and Oeno, whose quota of
indigenous plants exceeds 75%, provide far better data on species-area
relationships.

Motu Size in Relation to the Distribution of Trees, Shrubs and Herbs

As one progresses from small to large islets (Table 5), the number
of tree species rises from zero to 7, the number of shrubs from one to
4, and the number of herbs from 2 to 12. Caroline’s trends are similar
to those at Aitutaki (Stoddart & Gibbs 1975), where the numbers of trees
and shrubs are relatively constant over a wide range of motu sizes (3.8-
71 ha), while the number of herbs shows a slight increase. There are
too many recent exotics on Kapingamarangi for comparisons to be valid.
We believe that if Niering’s data were reanalyzed, using only indigenous
species, similar generalizations would be found, viz: most species on
atolls establish rapidly on small motus, after which a few additions
occur on motus of increasing size until the maximum number of
potentially available species is reached. Cursory examination of
Niering’s Figure 31, detailing the breakdown of total species numbers
into indigenous and non-indigenous components, bears out this
hypothesis.

90
G. PLANT COMMUNITIES

General Account

The total area covered by vegetation on Caroline is 357.55 ha, 90%
of the combined areas of all the motus. Of this, two-thirds (289.82 ha)
is woodland. Substantial areas of Caroline’s native woodlands and herb
mats are relatively pristine, and 89% (possibly 92%) of its plant
species are indigenous. Twenty-three (60%) of its 39 motus harbor
wholly indigenous vegetation (Figs. 27-30). Atolls supporting
substantial areas of native forest are typically remote and uninhabited.
Where people are present, native vegetation is usually confined to the
smallest motus or the extremities of larger ones--areas with marginal
human usefulness.

Overall vegetation patterns strongly support the theory that the
original vegetation of many atolls is arranged in concentric or parallel
belts according to salinity and ground water gradients, drainage, and
exposure to salt spray (Fosberg 1976). Because of its impoverished
flora, Caroline has no mixed broadleaf forest per se but is rich in pure
stands or simple combinations of 2 or 3 species (Figs. 17, 21, 23, 26-
29). Monotypic stands of shrubs and trees are common on atolls, but
unusual for the continental tropics, where species diversity is
considerably larger.

The present vascular flora of Caroline, 26 species, is organized
into 7 plant communities (11 subcommunities) defined principally by
dominant species (Fosberg 1953, 1976), whose areas are given in Table 8.
Eight subcommunities are natural, 3 are anthropogenic (Table 5). The
subcommunities include a mix of dominant species, which are discussed in
the major community sections below.

NATURAL COMMUNITIES:

Natural Herb Mat

Beach Scrub with Suriana
Pandanus Forest

Tournefortia Scrub and Forest
Cordia Forest

Pisonia Forest

ANTHROPOGENIC COMMUNITY:
Coconut Woodlands

Natural Herb Mat (67.73 ha) Figs. 19;0235vPiseelee
32, 34, 45, 46, 47

Widespread and predictable on wind- and salt-blown coastal coral
rubble and incipient motus, these mats are composed primarily of
Heliotropium and Portulaca. They are pioneers on newly emergent motus,
cover most of the ground area of small motus, extend inland along
ancient reef channels, and typify newly evolving land which connects or
augments established islets. Natural herb mats may persist through al]

91

Table 8. Areas of plant communities on the islets of Caroline Atoll.

% Of
Total Total
Area Land Area
(ha) Area (ha) %
Unvegetated Habitats 41.39 10.37
Coral Rubble and Sand 41.39 10.37
Natural Plant Communities 261.41 65.53
Natural Herb Mats 67.73 16.98
Beach Scrub with Suriana 1.49 0.37
Pandanus Forest 3.38 0.85
Lournefortia scrub and Forest 25.25 31.40
Cordia Forest 1.39 0.35
Pisonia Forest 62.17 15.58
Anthropogenic Community 96.14 24.10
Coconut Woodlands 96.14 24.10
Total Area Above High Water 398.94

Pure Pandanus only. Also mixed with Pisonia, Tournefortia, and Cocos.
Cordia, where mixed with Pisonia and Tournefortia, is included in
totals for those forest communities.

5 stages of plant succession as long as sunny openings occur.

Caroline’s motus illustrate 2 general principles: 1) the smaller the
area the more extreme is the strand character of its vegetation, and its
corollary, 2) as areas enlarge, strand flora becomes less important
(Fosberg 1949).

The following species are present (see Table 2 for abundance
indices):

Trees: Morinda citrifolia (1 drift seedling on 1 motu);

Shrubs: Tournefortia argentea, Suriana maritima, Scaevola taccada;
and

Herbs: Heliotropium anomalum, Portulaca Jutea, Boerhavia repens,

Lepturus repens, Laportea ruderalis, Lepidium bidentatum,
Ipomoea macrantha.

Near the high water mark, the herbs are recumbent, leathery, and
somewhat desiccated. As environmental conditions improve further
inland, they spread more laterally and average up to 7 cm tall. Their
rubbly habitat, often sprinkled with Tournefortia, resembles a low

92

savannah. Although these prostrate herbs can tolerate intense sunlight,
they grow optimally in slight shade, sandy soils, and higher relative
humidity, when they may reach 22 cm tall, forming a fairly thick mat
(P1. 45). With too much shade the mats disappear or their species
proportions and abundance changes according to the presence or absence
of sunny clearings. Thus, natural herb mats may be found in patchy
clearings within forests up to 13 m tall. They are common in the
abandoned Cocos plantations of South Island, where Boerhavia
proliferates into thick mats which completely cover the substrate, vying
with Phymatosorus and Ipomoea for "lebensraum" (P1. 34). A thick,
exposed mat of succulent herbs, primarily Portulaca, is found on Noddy
Rock.

Herb mats occurred on almost every transect, windward and leeward,
ranging from 1% to 60% coverage (Figs. 19, 23), predominating in
sparsely vegetated areas. The most extensive areas (coverage 35-50%)
were on Skull, Tridacna, interior South, Emerald, and Mannikiba. Their
widths varied according to the age, shape, exposure, and geographic
position of the motu but were widest on seaward-facing shores. Wide
bands of herb mats may encircle an entire motu; to windward they average
36 m (Table 9), while, bordering the relatively placid and
intermittently shaded lagoon, they shrink to a mere 0.9 m. On leeward
motus, the corresponding figures are 18.5 m and 4.2 m.

Table 9. Widths of natural herb mats on seaward- and lagoon-facing
shores, Caroline Atoll.

Average Width of Herb Mat (m)

Seaward Lagoonward
Leeward Motus 18.5 4.2

(3-81) (0-28)
Windward Motus 36.0 0.9

(24-69) (0-3)

Although reef flats widen where islets turn sharply, it is not
unexpected that these perimeter bands are the most extensive on the
extremely exposed shores of northern Nake (Pl. 16) and southern South
Island. On the latter, they are up to 59 m wide. Similarly, on small
exposed motus (e.g. Skull, Noddy Rock), they carpet most of the area
(Fig. 27). Under such conditions, Portulaca and Boerhavia develop much
redder stems, possibly due to the presence of a chemical "sunscreen" or
introgression with P. oleracea (Fosberg, pers. comm.).

93
Seabird Use

Whether bordering the edges of established islands or composing
the entire ground cover of tiny motus and ancient reef channels, herb
mats are nesting sites for Red-tailed Tropicbirds, Masked and Brown
Boobies, Sooty Terns, and Brown Noddies. They provide sites for the
development of phosphatic hardpan (Sect. D, Substrata). Herb mats also
provide foraging grounds for shorebirds.

Beach Scrub with Suriana (1.49 ha) Fig. 20; Pls. 23, 39, 44

Uncommon on Caroline, Beach Scrub with Suriana is typically found
on sand or sandy rubble bordering Tournefortia or Cocos. On Caroline,
it is evidently limited by the paucity of low-lying sand and gravel
sheets, with which it is normally associated elsewhere (Fosberg 1953,
Wiens 1962, Stoddart & Gibbs 1975).

The following species are present (see Table 2 for abundance
indices):

Shrubs: Tournefortia argentea, Suriana maritima; and

Herbs: Heliotropium anomalum, Boerhavia repens, Portulaca lutea,
Laportea ruderalis, Phymatosorus scolopendria, Lepturus
repens.

This plant community was found on 9 motus (Fig. 20), either in
thick bands or as scattered shrubs. Suriana is most robust on sandy
substrates, especially fringing the lower lagoon on South Island
(Fig. 36; Pls. 23, 39) and on windward Tridacna. The fringe, repeatedly
interrupted by other species, grows to 12 m wide and 1.8 m high. Here
the shrubs are closely appressed and slightly entangled, forming dense
shade which supports a sparse understory. On South, where its roots are
submerged at high tide, it is being shaded out by overhanging Cocos
(compare Pls. 39 and 40), having retreated since 1965. Suriana also
occurs as scattered individuals or in open bands in coarse rubble.

Beach strand up to 60 m wide, containing herb mats, Tournefortia, and
scattered Suriana, were found on South (Tr. 1, Pl. 20), Long (Tr. C),
Brothers, Matawa, Long, and the Southern Leeward Islets.

Pandanus Forest (3.38 ha?) Fig. 11; Pls. 17, 35-38

Although several species of Pandanus are native to the Line
Islands, and their seeds are common components of Pacific sea-drift
(Ridley 1930, Stone 1968), it is possible that the groves of
P. tectorius on Caroline represent both natural forests and cultivars
transported by early Polynesians. However, we have no data to verify

“This figure is pure Pandanus forest. Mixed forests containing Pandanus
account for a further 14.96 ha.

94

this, except that its present and past distribution on Flint Island
suggests that it is indigenous there (Kepler, in prep.). Its largest
acreages are on 2 islands that contained historical settlements (Nake,
South). However, its presence within the interior forests of a few
motus lead us to conclude that it may have experienced a dual
introduction. On Emerald, 3.20 ha (38% of the islet) supports a mixed
forest of Tournefortia, Pisonia, and Pandanus. Similarly, Shark’s
interior woodlands of Journefortia, Pisonia, and Cordia (5.52 ha, 70% of
the islet’s area) also contain a substantial amount of Pandanus, but
there was possibly a hut on Shark last century. The occurrence of
Pandanus groves or lone trees on other islets (Fig. 11) is easily
attributable to drift seedlings. Dried Pandanus phalanges are the most
conspicuous litter along Caroline’s lagoon beaches (Pl. 38); its seeds
last for months in seawater (Guppy 1906) and are probably distributed
locally by rats and land crabs, as noted elsewhere (Ridley 1930).
Phalanges from Nake’s southern mixed woodlands undoubtedly established
the grove on Pandanus Islet.

The mixed forest with Pandanus on south Nake (with Cocos, Cordia,
Pisonia, and Tournefortia) contains up to 50% Pandanus attaining heights
Of 27m (High) 37k

Many Pandanus trees were felled on South Island during the coconut
planting era (ca. 1873-1925), as we know that they were "somewhat
numerous" in 1834 (Bennett 1840), but only "one or more of the screw
pines were found growing in various parts of the island" in 1883
(Trelease 1884). A drawing in this latter paper depicts a grove from
South, where today Pandanus is uncommon in the beach scrub bordering the
Cocos plantation.

Trees were fruiting abundantly in September 1988. The green
phalanges, 17.5-20 cm in diameter, ripen to yellow and orange when they
fall, to be eaten by hermit crabs, Coenobita perlatus (Pl. 38).

Tournefortia Scrub and Forest (125.25 ha) Figs. 24,232;9Piseade
9, 51957295 a47

General Distribution

Characteristic of many Pacific islands, Journefortia, a
broadleafed evergreen, dominates the wooded motus of Caroline, forming
40% of its vegetative cover and 31% of the total land area (Fig. 24,
Table 8). Its pale foliage and hemispherical canopies (to 14 m tall)
typically surround the taller, darker canopies of Pisonia and Cordia.

A hardy halophyte, Tournefortia occurs on every motu and in every
habitat except pure Pisonia forest. It is tallest, widest and lushest
on the windward motus, particularly on those where Pisonia is also best
developed. Without direct sun though, as under dense Pisonia or Cocos,
it withers (Pl. 23).

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On other atolls Journefortia forms a narrow or interrupted belt
inland of the beach, or is a component of mixed scrub (Fosberg 1953).
However, given the floristic poverty on Caroline, especially of shrubs
and trees, Journefortia not only has expanded into niches which might
elsewhere be occupied by combinations of Scaevola, Pemphis, Suriana,
Terminalia, Hernandia, Thespesia, Hibiscus, etc., but frequently occurs
in pure stands (113.03 ha) that extend well inland. It thus occupies a
much higher percentage of the islet areas on Caroline than on atolls
with greater biodiversity. For example, Nake, the largest islet, has
the greatest amount of Journefortia (79.68 ha): 28.9 ha of pure scrub
and forest, 18.28 ha of "savannah," 17.48 ha with Cordia, 8.99 ha with
Pisonia, and 6.03 ha mixed with Cocos, Pandanus, and Pisonia.

Overall, we classify Tournefortia as a shrub (Stoddart & Gibbs
1975). However, following Mueller-Dombois et al. (1981, p. 58), we also
distinguish between its shrub (scrub) and tree communities. Because
they intergrade, we may lump them together (vegetation maps and
schematic profiles of the motus), or treat them separately (Tbs. 2, 5,
10 and ecological discussions):

1) Tournefortia Scrub: <5 m high (x = 2 m), <60% canopy coverage
(Pls. 19, 29, 32, 47). This open scrub growth is typically confined
to islet perimeters or emergent reef channels and covers much of the
vegetated rubble on smaller islets. Its species composition is
similar to that of the taller forest, except that herbs are more
prominent.

2) Tournefortia Forest: >5 m high (x = 8 m), >60% canopy coverage
(Pl. 48). This taller, closed forest, with maximum height 15 m,
develops as a second belt of woody vegetation approaching the
interior of the larger islets. Figure 32 depicts a schematic profile
through pure Journefortia scrub and forest, while Figure 35 diagrams
a profile of a larger islet where Journefortia is represented only on
its periphery.

Species Diversity in Tournefortia Woodlands

The following species occur in both scrub and forest. Those
marked "*" occur primarily in the scrubland (Table 2).

Trees: Pisonia grandis, Morinda citrifolia, Pandanus tectorius,
Cocos nucifera, Cordia subcordata;

Shrubs: Suriana maritima, Tournefortia argentea, Scaevola taccada,
Ximenia americana; and

Herbs: *Heliotropium anomalum, *Boerhavia repens, *Portulaca lutea,
*lepturus repens, *Laportea ruderalis, *Achyranthes
canescens, Phymatosorus scolopendria, Ipomoea macrantha.

Caroline’s tallest Tournefortia stands (12-15 m) occur on Nake.
On all other windward motus, the Tournefortia canopies vary between 6

97

and 9 m tall, shorter than expected if their forests were virgin. This
has historical significance: we do not know the extent of forest
felling (if any) on the Windward Islets (Crescent through Tridacna)
during the guano era, but we do know that 4,587 coconut palms were
planted during 1919-20, and that "misses" (dead seedlings) were
fastidiously replaced over the following 2 years (Young ca. 1922).
Thus, their forests, though weed-free today, comprise secondary growth
around 60 years old. It is not surprising that Achyranthes canescens
and Lepturus repens, both weedy (though indigenous), are particularly
common inland on some windward motus (Figs. 13, 16). Tournefortia’s
rapid recovery illustrates that ecosystems in the pioneer stage
generally recover their original condition rapidly when left alone
(Fosberg 1983).

Stature and Area Coverage

Forming an umbrella-like canopy, a typical Journefortia forest is
very simple. Its twisted branches and gnarled trunks stretch untidily
over an open understory. The lower branches die off as the trees
increase in stature. Sometimes a scant herbaceous cover develops in
localized pockets of better soil, such as a clearing where a dead tree
fell or a semishaded spot beneath a colony of seabirds.

Tournefortia is abundant throughout the atoll. Areas with 90%-
100% canopy cover were found on Nake (Tr. 4), Long (Trs. B, C, 4, 6, 10,
12), North Pig, Pig, North Brothers, Brothers, Crescent, Arundel
(Fig. 32), Tridacna (Trs. 1, 2), South (Trs. 1, 4), all 5 Southern
Leeward Islets, all Central Leewards over 0.5 ha, and Pandanus Islet.
Tournefortia is present across the entire width of some small motus,
e.g. Fishball (144 m wide). Even on larger motus such as Mannikiba
(280 m wide), Tournefortia blankets nearly all the land (Pl. 63). Long
(75.98 ha) is a composite motu: long, narrow, and derived from the
coalescence of at least 5 former islets. Because Tournefortia encircled
the perimeters of these ancient islets, it is now present in 5 sets of
concentric circles, connected by herb mats, down the length of the
island (Fig. 39).

In the herb mats, Tournefortia is small (x = 1.4 m) and widely
scattered (Table 10). It may be of typical hemispherical shape or
irregularly windshorn (Pls. 13, 45). On windward coasts they typically
form a tight wind barrier, one or 2 trees thick. Moving inshore from
the seaward fringe, the trees become progressively taller (x = 6 m) with
a more open understory. Cordia often mixes with Tournefortia, either as
scattered individuals in the understory or canopy, or as small groves.
On the Southern Leeward Islets such belts border the seaward scrublands.

Although still widespread in the Pacific, Tournefortia is far less
abundant than formerly. On inhabited islands it exists primarily in
relict patches or as edging around anthropogenic forests. It rarely
covers most of the land area of islets; exceptions are Taongi and Bikini
(Marshall Islands), Gaferut (Caroline Islands), and Ducie Atol]
(Pitcairn Islands) (Fosberg 1956, Wiens 1962). The finest quality

98

Table 10. Stature and percentage cover of Journefortia in the major
habitats of Caroline Atoll.

%
Av. Hgt. Av. Width Tournefortia No. No.
(m) (m) Cover Motus Transects

Natural 1.4 49 25 14 20
Herb Mat (0.3 - 1.8) (3 - 198) (5 - 95)
Tournefortia 6 55 81 38 71
Scrub & (053°—" 15) (2>=" 287) (5 - 100)
Forest
Tournefortia- 9.5 98 47 18 27
Pisonia (5 - 15) (8 - 284) (5 - 90)
Forest

Tournefortia on Caroline (15 m tall, 80% cover) occupy central and
northern Nake (Fig. 37), but even this islet was completely felled for
Cocos (Table 13). These 15-m Journefortia compare favorably with 18 m
specimens found at Jemo Island by Fosberg (1956). Perhaps Jemo’s trees
are at the upper size limit for the species, as Tournefortia is
generally recorded as 3 to 6 m tall (Wiens 1962).

Ecology

Tournefortia is an integral part of the atoll’s evolution and
ecology. Bearing seeds capable of floating for at least 4 months in the
sea (Guppy 1906), it is the first woody plant to establish on Caroline’s
tiny motus (<0.1 ha), appearing immediately after the native herbs have
begun to germinate in the coarse coral rubble. It is the only plant
species on tiny Ducie Atoll (Fosberg et al., 1989). Requiring little or
no soil and adequate rainfall, it can grow up to 2 ma year (Fosberg
1959). Tournefortia’s leaves contribute to soil development, paving the
way for plant succession from Stages I through IV, for it only persists
in soils that are conducive to the growth of its mesophytic competitors
(Fosberg 1953). The most mature trees (x = 9.5 m) occur at the
Tournefortia-Pisonia interface, but die off as Pisonia expands. When
Tournefortia has reached its maximum height, most of its lower branches
have fallen, leafage is reduced, and flowers and fruits are few.
Tournefortia usually drops out after one generation. Seedlings are
rarely seen in heavy shade, and fallen trees are fairly common on the
edge of the interior forests where Pisonia replaces it.

An example of complete replacement of Tournefortia by Pisonia is
illustrated by nearby Vostok. It has heretofore been assumed that
Vostok’s sole tree species was Pisonia grandis (Fosberg 1936, Bryan

99

1942, Clapp & Sibley 1971b, Garnett 1983). However, Young (ca. 1922)
stated that when Capt. J. Larsen, of the schooner Papeete, planted 100
coconuts there on 31 May 1922, he found "Pukatea and Tauhinu trees, etc.
60 to 80 feet high," that is, Pisonia grandis and Tournefortia argentea,
but no "Tou" trees (Cordia subcordata). By 1935 only Pisonia remained
(Fosberg 1936); hence, the last Tournefortia must have been eliminated
naturally by Pisonia.

Along some coasts (Long, Nake, South), Tournefortia overhangs the
water, its roots immersed at high tide. We found floated debris up to
20 m inland within dense Journefortia forest, indicating that this hardy
shrub can withstand periodic storms and high tides. If a rosette of
Tournefortia leaves is placed in fresh water, it droops within an hour,
indicating that its tissues require a high salt concentration in order
to maintain turgidity (pers. obs.). Perhaps decreased salinity in the
ground water, coupled with reduced light intensity in advanced seral
stages, contribute to the eventual disappearance of Journefortia in the
center of coral islands.

Seabird Use

Tournefortia is a favored roosting and breeding site for most of
Caroline’s seabirds. The taller the trees, the greater the bird
diversity they harbor: scrub contained 4 species (36%) and forest, 9
(82%). Sooty Terns nest in tight colonies in its shade, its canopies
support large populations of Red-footed Boobies and Great Frigatebirds
(Pt. II), and its branches are favored by White Terns (Figs. 32, 35,
36). Tournefortia leaves provide nesting material for noddies.

Cordia Forest (1.39 ha) Fags 225 RiseZ26

General Distribution

Cordia does not form "the main native woodland" on Caroline Atoll,
as implied by Clapp & Sibley (197la) and stated by Stoddart & Gibbs
(1975, p. 104). It occupies far less area than Journefortia or Pisonia
(Table 9). Cordia is generally mixed with other emergents: monotypic
Cordia forest covers only 1.39 ha, while Tournefortia or Pisonia
containing substantial amounts of Cordia total 25.89 ha. En toto, this
is less than 10% of Caroline’s woodlands, and Cordia is usually
subdominant. These "mixed" forests of Cordia mixed with Pisonia or
Tournefortia (rarely all 3), occasionally with a Morinda understory, are
the closest equivalent to Mixed Broadleaf Forests of other coral
islands. This widespread Pacific plant community (Fosberg 1953, 1976;
Wiens 1962) is conspicuous by its absence on Caroline. We treat Cordia
forest as a separate plant community because of its increasing rarity on
Pacific atolls, which makes Caroline’s groves an increasingly important
resource in need of conservation. Cordia forest occurs primarily on
Nake, Windward, Crescent, North Pig, Pig, Danger, Shark, and the
Southern Leeward Islets.

100
History

Bennett (1840) recorded "two species of Tournefortia" on Caroline,
possibly referring to Tournefortia and Cordia. There are no other 19th-
century records. From Cordia’s present distribution we can infer that
it was formerly more extensive on South and Nake. Scattered trees
within and bordering the Cocos plantations suggest that its history is
similar to the species on Flint, which was "in 1872...covered with a
forest of ‘Tou’ trees Cordia subcordata)" (Maude ca. 1942). Both Flint
and Caroline were worked simultaneously by the same companies for guano
(1872-1890) and copra (into the 1930s). Pisonia and Cordia forests were
felled to make room for coconuts. From Flint, several hundred Cordia
logs were exported to San Francisco for furniture and panelling. The
last logs were exported in 1896, 6 years after the guano supplies were
depleted, but coconuts were still being planted (Young ca. 1922), and
some large Cordia trees were still present in the southern 20% of the
island still covered with virgin forest (E. Campbell 1908, Kepler, in
prep.). Today, Flint’s recovering forests contain much Cordia
(A. Kepler 1990b), unlike Caroline, where Cordia is rare in similar
habitats. Some of Flint’s present windward Cordia trees may be those
"few tiny, struggling...trees...recently planted" (St. John & Fosberg
19OSZ)

Abundance and Distribution

Cordia seeds are dispersed by ocean currents and can germinate
after 40 days in sea water (Guppy 1906). On Caroline this species
develops both as an understory shrub and forest emergent (to 15 m high).
It typically occupies the woodland periphery, occurring in small
circular or linear groves, or mixing with Journefortia and/or Pisonia
(Table 5). Cordia may form tall, straight-trunked trees (Pl. 26) or
sprawl like Hibiscus tiliaceus. In dry rubble sites it may become
chlorotic or semideciduous. The tallest groves are on Pig (Pl. 26),
where 6 trees averaged 12.6 m tall, 116 cm circumference at 1.5 m (cbh),
and 99.8 cm base circumference. Lush Cordia groves occur in sheltered
parts of the upper lagoon on Long Island (Tr. 10).

Flowering times are unpredictable. Two flowers were seen in
September 1988. In November 1989, flowers were abundant, extending
through March, yet in November 1990 not one flower was observed (pers.
obs., Anne Falconer and AKK).

Seabird Use

Black and Brown Noddies, frigatebirds, and White Terns nest in
Cordia wherever it is a forest component. Great Frigatebirds and Red-
footed Boobies favor roosting in the lush, lagoonside forest of Cordia
and Pisonia near the south end of Long Island.

101

Pisonia Forest (62.17 ha) FIGS 1855.55), 345 305 39, 415
Pls 430750). 513) Tbs. l-14

General Distribution

Although Pisonia grandis was previously recorded as "present"
(Trelease 1884, Clapp & Sibley 1971la), the quality and extent of its
forests has not been recognized. Some stands are prime representatives
of a major ecosystem that was formerly far more widespread in the
Pacific.

Common throughout the atoll, Pisonia occurs on 29 motus, covering
22% of the woodlands. Well developed groves, 10-21 m tall and up to
359 cm circumference at 1.5 m, are present on 23 of these (Table 11).
Although present on motus less than one hectare in size (Table 5), it
typically occupies interior forests (schematic profile, Fig. 35), with
individual trees or groves contributing from 5% to 100% of the canopy.
In general, Caroline’s windward motus support the lushest forests: the
maximum height of windward Pisonia forests is 21 m, of leeward forests,
15 m.

Mature Pisonia forests are monocultures of grandeur. The trees
bear one to several stout boles of irregular shape, whose rotting
cavities often harbor large coconut crabs or miniponds alive with
mosquito larvae. Their scraggly branches occasionally bend over and
reroot. It is dark and humid, but open except for exposed roots and
scattered broken branches. Few seedlings occur. Polynesian rats scurry
underfoot.

In September 1988 we saw no flowers or fruit. Anne Falconer
reported flowers on Motu Ana-Ana in August 1990. Pisonia was beginning
to bloom on Vostok in March 1990 (A. Kepler, in prep.).

An Historical Perspective

Some of Caroline’s most mature Pisonia groves (to 21 m tall,
660 cm circumference at 1.5 m, multiple trunks) appear to be virgin,
(Pl. 43) but are most likely only 60-70 years old. Overall dimensions
of the trees, the low species diversity, and general character of the
plant community compare favorably to virgin groves on Vostok (Table 12;
A. Kepler 1990c, d).

Despite the maturity of many groves, especially those to windward,
planting records from 1916-22 indicate that Cocos was planted throughout
not only South, but also on Nake, Long, and all the major Windward
Islefs (Young ca. 1922). Given the standard planting density of
73 m°/palm (Young ca. 1922), we calculated the approximate area on each
islet given over to Cocos plantations, based on the number of coconuts
planted times the area required for each tree. We then compared this to
the usable areas based on today’s forest cover (Table 13). On the 9
Windward Islets, collectively, 36.36 ha were planted in Cocos, fully 92%

102

(m)

HEIGHT

CANOPY

50 100 200 300 400 500 600 700 800

WIDTH OF mOTU (m)

Figure 33. Maximum heights of Pisonia forests in relation to width of the motus. Stars represent forests with
90-100% canopy cover; dots represent forest or scrub with less than 90% cover.

(ma)
=

CAMOPY HEIGHT

0.01 0.1 1 10 108
AREA OF MOTU (HA)

Figure 34. Maximum canopy heights of Pisonia forests in relation to motu area. Stars represent forests with 90-100% canopy cover; dots
represent forest or scrub with less than 90% cover.

103

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104

Table 11. Distribution of well-developed (210 m height) Pisonia forests on
the motus of Caroline Atoll. Motus and transects are arranged
according to the decreasing height of their Pisonia groves.
Capitals indicate those motus whose forests were felled for Cocos
plantations from 1916-20.

Area of
Pisonia Pisonia Motu Area

Motu & Transect Height (m) (ha) (ha)

PIG Ole 330 Tse
NAKE, Transect 4 20* 20.79 107.46
Booby 20* 0.12 0.84
NORTH PIG 20* 1.83 5.44
NORTH BROTHERS 18* 0.43 7A
NAKE, Transect 3 (central) 1\5* 20.79 107.46
LONG, Transect 0 15% 15.00 75.98
BROTHERS 15* 0.37 4.31
Ana-Ana nS 0.93 2.16
Danger 5* 0.39 2.71
NAKE, Transect 2 14* 20.79 107.46
Bird 14* 1.70 4.05
WINDWARD, Transect 2 14° 2.97 11.42
Raurau 14* 1.07 3.48
CRESCENT 13% 0.51 3.10
Mannikiba, Transect 1 1225 I sals} 21.49
Shark 128 2.60 7.98
NAKE, Transect 3 (west) 124 20.79 107.46
LONG, Transect 12 2s 15.00 75.98
Pisonia Niles 0.86 2.45
Matawa 1] 0.07 Wel
Nautonga We 0.02 0.34
NAKE, Transect 3 (southwest) 11° 20.79 107.46
Kimoa Lis 0.59 1.80
Emerald iil 3.20 8.34
Eitei tS 0.38 1.42
LONG, Transect B 10° 15.00 75.98
LONG, Transect 8 10 15.00 75.98
NAKE, Transect 1 10° 20.79 107.46
WINDWARD, Transect 1 10 2.97 11.42
Blackfin 10° 0.41 2.62
NORTH ARUNDEL 10° 0.18 0.91

nnn

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107

of all usable ground; in several cases the amount calculated for Cocos
by Young exceeded our estimates of potentially usable ground. Thus,
Cocos was planted so intensively on the Windward Islets that virtually
all Pisonia and most Tournefortia woodlands were felled.

Two remarkable points emerge from Table 13: 1) scarcely any Cocos
remains today on the 9 Windward Islets; not one palm exists on 7 motus
(Figs. 43, 44, 47, 48), and 2) the recovery of indigenous plant
communities, Stages I through V (Sect. F) on the windward side has been
rapid and, at least on Brothers Islet (Fig. 46), reasonably complete
with regard to ecological succession and species diversity of plants and
seabirds. Today the Windward Islets have the lushest and tallest plant
communities, with a higher species diversity than the leeward islets
(Table 3), which have evidently experienced far less human disturbance.

This differential disturbance on the windward and leeward sides of
the atoll explains enigmas such as 20 m tall Pisonia forest on the
leeward Booby Islet (0.84 ha), taller than most of the windward forests;
the absence of Pisonia on windward Tridacna Islet (9.08 ha), which,
being close to South Island, probably supported Cocos which was managed
longer than the more distant windward islets; and the patchy
distribution of Pisonia in the interior of several islets (e.g.
Windward, Arundel). This last point also applies to Mannikiba
(21.49 ha), the largest leeward islet. According to Young (ca. 1922),
6,000 seed sets were brought from Flint to Caroline in 1920 and kept on
Mannikiba. This "nursery stock" was used to replant "misses" on other
islets, due mostly to destruction by coconut crabs and poor planting.
Today, Mannikiba’s total acreage of Pisonia (Fig. 53) is very small and
fragmented relative to the islet’s size: 1.13 ha, 5% of the total land
area. Compare this with Bird Islet (Fig. 55) which, as far as we know,
has never been disturbed: 1.70 ha Pisonia, 42% of the islet’s land
area.

On both Caroline and Flint there is much variation in the quality
of the regenerated Pisonia forests (Table 12). Some trees bear
enormous, partly rotting boles, black algae smothering the bark,
multiple trunks, and few or no understory herbs. Other trees are tall,
straight-trunked, with characteristic whitish bark, and bear no rotting
holes in their bases. These observations suggest that when their
indigenous forests were felled, only minimal cutting was done, and many
Pisonias were able to regenerate quickly by sprouting from rooted stumps
and fallen branches. This speculation is supported by the fact that
some of Vostok’s Pisonia trees regenerated similarly. Maude (1953,

p. 96) states that "there is room for 8,000 palms on Vostok, but only
100 have been planted and most of these have been choked in the
luxuriant ’buka’ (Pisonia grandis) forest: no attempt having been made
to exploit the island since the initial planting."

Pisonia, a soft, pulpy wood, has a well-known ability to sprout or
send up suckers from dismembered branches or fallen trunks (Fosberg
1953), and it has been noted that older trees are virtually
indestructible, fire being the only effective means of clearing forests

108

(Wiens 1962, p. 397). The senior author has photographed leaf sprouts
from partly burned twigs as small as 1 m long and 5 to 6 cm in diameter.

Since the existing Cocos plantations on South and southwest Nake
contain few Pisonias, it seems that forest clearing was more thorough on
the atoll’s larger islets than on the smaller ones, which today manifest
scant traces of their former history. Fortunately for Caroline, its
coconut plantations were plagued by a number of problems, which resulted
in their double abandonment: coconut crabs, seabirds, rats, Ipomoea
vines, and an unknown disease (see under Coconut Woodlands, this
section).

A footnote in Young (ca. 1922, p. 15) states that "the larger
portion of the 30,000 trees planted were either badly planted or smitten
with some disease as in 1927 it was reported by Mr. Bunckley that most
of them had perished." In 1929 only 13,215 trees were left, and more
were being planted. Considering the distribution of both palms and
natural forests today, it appears that plantations continued on South
and Nake and were abandoned on the smaller islets, allowing for a better
recovery than might be expected had the Cocos grown to maturity.
Tridacna (close to South) and Mannikiba (a nursery) were likely the most
intensely managed of the smaller islets, as their Pisonia today is
meagre compared to their overall areas.

Once a Cocos plantation has been well established and subsequently
abandoned, Pisonia regrowth is more difficult. This is characteristic
of many tropical islands. For example, on Cousin Island (Seychelles
Islands, Indian Ocean), an ICBP wildlife perserve since 1968, Pisonia is
currently reestablishing within a deteriorating Cocos plantation.
Phillips & Phillips (1990, p. 37) envisioned "centuries rather than
decades before something like a natural ecosystem develops." We predict
a similar time frame for Caroline’s South Island, sooner for Flint.
Forest recovery on islands elsewhere has evidently not been studied in
detail (Fosberg, pers. comm.).

Annual Growth Rates

Data on Pisonia grandis growth rates is very limited. A 7-year
study on Kabelle Island, Rongelap Atoll, Marshall Islands, disclosed
mean diameter growth rates of 1.32 and 0.39 cm/yr at 2 sites (Gessel &
Walker 1992). On Cousin Island, vegetation changes, including Pisonia
and Cocos, have been monitored since 1974, but no growth rates are yet
available (Phillips 1984, Phillips & Phillips 1990).

Because of this paucity of data on Pisonia, and because its
forests have diminished significantly this century, we present the
following data in the hopes that it might inspire more research.

One point is clear: on all 3 of the Southern Line Islands Pisonia
grandis has recovered fast from disturbance (except for total forest
elimination), reaching close to its maximum height and ecological
maturity in 70 years or less. Mature Pisonia, under optimal conditions

109

of soil, temperature and rainfall, may attain 35 m, as on Fanning and
Washington (Garnett 1983 and pers. comm.). However, in the Southern
Line Islands, canopies of similarly virgin Pisonia on Vostok rarely
exceed 25-30 m tall (A. Kepler 1990c), and only 16-18 m on the Great
Barrier Reef (Walker 1991).

Caroline’s prime groves, 21 m tall, with circumferences to 660 cm,
and bearing multiple trunks and root suckers, we now know date back only
to the 1920s. The largest trees, 21 m high and 660 cm cbh, appear to
have averaged annual growth rates of 0.32 m in height and 3.4 cm in
diameter.

Further evidence of fast growth rates is provided from Flint. In
1934 only one small Pisonia and a few tiny, struggling Cordia, recently
planted, were recorded (St. John & Fosberg 1937). Virtually the entire
island (324 ha) was a Cocos plantation. In 1990 Pisonia, quite common
on the windward side, attained maximum heights of 30 m, with base
circumferences and at 1.5 m (cbh) of 1,000 cm and 200 cm, respectively
(Table 12). These compare favorably with 2 trees of similar heights and
cbh to 510 cm on Vostok and a large Pisonia, presumably virgin, measured
on Atafu Island (Tokelaus) by the U.S. Exploring Expedition in 1840,
which was more than 600 cm in base circumference and about 12 m tall
(Wilkes 1845, Vol. V, p. 9). Furthermore, pure indigenous mixed
broadleaf forests (Pisonia, Cordia, Guettarda), to 25 m tall, covered
57 ha, 23% of Flint’s vegetated area (Kepler, in prep.), with a further
65 ha (26% of the woodlands) in mixed forests containing less than 50%
Cocos. Thus, Pisonias have established themselves since the plantation
was abandoned in the late 1930s. The largest trees, 30 m high and
200 cm cbh, now indicate a mean annual increment of 0.6 m height and
1.32 cm diameter over the past 50 years. A faster growth in height than
on Caroline is likely due to Flint’s higher rainfall and greater
relative humidity due to the presence of a more successful coconut
plantation inland: Caroline’s annual output of copra was 15 tons,
compared to 230 tons for Flint (Young ca. 1922, Maude 1953).

Species Diversity in Pisonia Forests

Caroline’s motus harbor every stage in the development of a
Pisonia forest, from stately monotypic groves to a single tree. The
plant communities between these extremes harbor the greatest species
diversity and most luxuriant growth on the atoll. The following species
are present (Table 2):

Trees: Morinda citrifolia, Cordia subcordata, Cocos nucifera,
Pandanus tectorius, Pisonia grandis;

Shrubs: Tournefortia argentea; and
Herbs: Boerhavia repens, Portulaca lutea, Laportea ruderalis,

Lepturus repens, Achyranthes canescens, Phymatosorus
scolopendria, Ipomoea macrantha.

110

The number of species within Pisonia forests ranges from one to 14
(Table 14). As Pisonia becomes more dominant, their trees are taller
(21 m), and species diversity is less (Table 14). Here the average
number of species is 3.4. Species diversity is also very low at the
other extreme of Pisonia development: in one young motu (Azure), only a
single 6-m-tall Pisonia tree is present (x = 4.0 species). The smallest
islet on which we found Pisonia, Azure is only 0.20 ha in area and 77 m
wide (Fig. 55, Pl. 51); more than half of it is rubble. The width of
its scrub is only 38 meters. Along a transect within the majestic
Pisonia grove (100% canopy cover) on Brothers (Fig. 46), we found no
other plant species, an extreme case of the barrenness of Pisonia
understory. This grove, 13 m tall and extending 42 m from east to west,
was sharply delineated from the 6-m-high Tournefortia forests on both
sides and provides a striking example of complete ecological succession
since its Cocos plantation days of the 1920s.

The highest species diversity occurred with mixed co-dominants
(Tournefortia, Cordia) and Pisonia coverage 25-50% (Fig. 34, Table 14).
Here, the average number of species was 6.2 (range 3-10). Regardless of
the area or width of the motu on which they occurred, these mixed stands
(x = 7 m tall) were always shorter than pure Pisonia forest.

Ecology

On Caroline, most plant species establish early in the evolution
of individual motus, increasing in abundance and stature while the land
area is quite small. Pisonia typifies this pattern: single trees occur
on 2 motus whose areas are only 0.2 ha (Table 6), suggesting that
Pisonia is partly salt-tolerant, at least in its early growth stages.
In general, however, motus less than 0.7 ha on Caroline have little
Pisonia (Table 6). It is difficult to imagine a freshwater lens on Motu
Nautonga (1 ha), where an 1l-m-tall Pisonia forest is found (Table 11).
Further evidence for the salt-tolerant nature of Pisonia comes from
Vostok, where a Pisonia forest, the sole woodland, extends to the edge
of the shoreline rubble and herb mat. The trees, tightly pruned by wind
and salt, have no buffer of coastal scrub. During storms, seawater
reaches Vostok’s interior forest, yet this 24-ha island supports one of
the largest and tallest (25-m-high) groves in the Pacific (Clapp &
Sibley 1971b, Fosberg 1977 and pers. obs.).

Many Pisonia trees were heavily infested with scale insects
(Coccidae) and Neuropteran larvae (Chrysopa sp.), identified by
Dr. Scott Miller (Bishop Museum, Honolulu, Hawaii). This appears to be
a natural phenomenon, as they were also abundant on the virgin Pisonia
forests on Vostok and also on secondary Pisonias at Flint.

Relationships Between Pisonia Forest Height and Motu Dimensions
Contrary to expectations, the tallest, most mature forests did not

all occur on the largest motus (Table 11). The 3 prime forests (90-100%
Canopy cover) were on Nake (total land area 107.46 ha), Pig (7.21 ha),

111

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112

and Booby (0.84 ha). Trees on Booby measure less in girth than those on
Nake and Pig, but their height (20 m) is impressive. Evidently, Booby
was never exploited for guano or planted in Cocos. Fine forests occur
on other small, undisturbed motus; for example, Pisonia grew to 14 m on
Raurau (3.48 ha) and to 11 m on Kimoa (1.80 ha).

A positive correlation exists between Pisonia height and island
width (Fig. 33). Motus appear to reach a minimum width of 90 m before
closed canopies of 13 m develop, and canopy height increases to 21 m as
motu width enlarges to 200 m (Pig, topmost star in Fig. 33). Further
increases in motu width did not result in taller trees. However, even
on motus with sufficient width, Pisonia did not develop unless other
environmental conditions were suitable. For example, on Long, Pisonia
only occurred in the centers of its former islets, not in the scrubby
areas where coalescence is more recent. Tridacna and Mannikiba, both
ideal for Pisonia, have not yet recovered fully from their Cocos
plantations.

Pisonia-Seabird Relationships

Seabirds are an integral part of Pisonia ecology. Its sticky
fruits adhere to the feathers of, and are thus dispersed by, seabirds
such as terns, boobies, and frigatebirds; thus, its early appearance on
small motus is not surprising.

On Caroline, 6 species of seabirds nest in its branches, dropping
considerable amounts of guano to the ground below. Black Noddies,
amassing in dense colonies, nest almost exclusively in Pisonia, along
with Brown Noddies, White Terns, Great and Lesser Frigatebirds, and Red-
footed Boobies. Pig Islet, with 7.25 ha of excellent Pisonia forest,
supported a dense colony of nearly 2,000 pairs of Black Noddies
(Pt. II). Bristle-thighed Curlews feed on the ground beneath its open
understory, and the Long-tailed Cuckoo forages within its canopy.

Seabirds may be so much a part of Pisonia ecology that a debate
exists as to whether Pisonia actually requires guano for successful
germination and establishment of seedlings (Shaw 1952, Fosberg 1953,
Wiens 1962). Very high phosphate and nitrogen levels are associated
with mature Pisonia, and concurrently the development of Pisonia forest
results in greatly modified soils that perpetuate its existence (Wiens
1962, Spicer & Newbery 1979). The formation of a highly acid raw humus
on the surface of the ground, sometimes in association with phosphatic
hardpan, has also been documented on several atolls by Fosberg (1953,
1956, no date; Stoddart & Scoffin 1983), including Vostok (AKK and John
Phillips, pers. obs.). We have no information on phosphatic hardpan in
Caroline’s Pisonia forests. For further discussion, see Section D,
Substrata.

113
Remnant Pisonia Forests in the Pacific

Though naturally and widely distributed throughout Indo-Pacific
islands (excluding Hawaii), Pisonia grandis forests have been subjected
to great destruction and are now rare. Pure Pisonia forest was formerly
the most widespread indigenous forest on Pacific atolls, and may have
formerly covered the greatest area of any tree species in the Pacific
(Wiens 1962, Fosberg 1976). Shaw (1952), summarizing its distribution,
stated that it only occurs on remote, generally uninhabited islands
ranging from the western Indian Ocean to the eastern Pacific, including
Malaysia. However, more recent studies, particularly by Fosberg,
indicate that because its habitat occupies, and is in part responsible
for, the most fertile areas of inhabited islands, its formerly extensive
forests have been largely replaced by coconuts. Though Pisonia’s soft
wood is of little use to either atoll inhabitants or to the timber
industry, its soils were rich sources of phosphate fertilizer and were
thus greatly disturbed during the guano mining era. Pisonia is highly
adapted for growth on coralline substrates, also having developed unique
morphological and physiological characteristics associated with seabird
colonies and mycorrhizal fungi (Walker 1991). Caroline, with 62.73 ha
in Pisonia forest (36.94 ha in monotypic groves) holds some of the
finest representatives of this ecosystem in the Pacific, even though
much of it is not virgin.

One of the prime Pacific Pisonia stands (13.5 ha on Vostok) was
partly burned in 1977 (Fosberg 1977). The Royal New Zealand Air Force
found it smoldering 3 months later (Fosberg 1977, pers. comm.). Ina
March 1990 visit to Vostok, we found that approximately 1.5 ha were
completely cleared (A. Kepler 1990c) and a further unknown amount of
land was affected. Cays of the southern Great Barrier Reef have
recently been found to harbor ca. 160 ha of uncut Pisonia forest. The
largest stand (94 ha) is on Northwest Island (Walker 1991). Other fine
groves exist on Palmyra and Washington (Northern Line Group), Rose Atol]
(American Samoa), Bikar, Taongi, and Jabwelo (Marshall Islands), and
Fanna (Southwest Palau Islands). Flint (Southern Line Group); Christmas
(Northern Line Group); Nikumaroro (Phoenix Group); Jemo and Ujae
(Marshall Islands); and Aitutaki, Penrhyn, Suwarrow, and Manihiki (Cook
Islands) have relatively small stands. Not all are healthy. For
example, groves on Bikar, Jabwelo, and Palmyra were recently devastated
by typhoons (Flint et al. 1992, IUCN 1992) and that on Taongi is
unhealthy (Thomas et al. 1989).

Coconut Woodlands (96.14 ha) Pies AM OR A Salis 425 s7e8h5
21-29, 325234, Sin 39, 40), 44

General Distribution
Cocos, although present on 15 motus and known to be planted

intensively on at least 13, covers significant areas only on the 2
largest islets, South and Nake (Table 13). Individual trees and smal]

114

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115

groves elsewhere are drift-derived or remnants of plantings made from
1916 to 1920.

The following species occur in habitats containing Cocos
(Table 2):

Trees: Pisonia grandis, Morinda citrifolia, Pandanus tectorius,
Cordia subcordata, Cocos nucifera, Thespesia populnea,
Hibiscus tiliaceus;

Shrubs: Tournefortia argentea, Ximenia americana; and

Herbs: Boerhavia repens, Portulaca Jutea, Laportea ruderalis,
Achyranthes canescens, Phymatosorus scolopendria, Ipomoea
macrantha, Lepturus repens, Tacca leontopetaloides, Psilotum
nudum, Phyllanthus amarus, Sida fallax.

The distribution of Cocos (Fig. 14) in order of decreasing
abundance is as follows: SOUTH: Forests old and neglected. Living
palms line the lagoon, currently shading out strip of native scrub.
NAKE: Southern forests (50-80% Cocos) healthier, younger, with more
native trees and Pandanus than on South; grove of about 50 palms on
northeast. LONG: Range from <1% cover (Tr. C) to dense fringe adjacent
to lagoon. EMERALD: Northeast and west-central patches. MANNIKIBA:
Main grove, northeast: 40 palms, 20 m high, another patch in south
center. ANA-ANA: House site, northeast point. BIRD, BLACKFIN,
BROTHERS, NAUTONGA, NORTH BROTHERS, PIG, PISONIA, RAURAU, SHARK: Few
trees each, primarily in Tournefortia. LONE PALM: One tree, central
forests.

History

A relatively small coconut grove was planted on South Island prior
to the 16th century by Tuamotuan settlers (Emory 1947, Maude 1968). In
1606 de Quiros noted "plenty of palms" and "many cocoa-nuts" (Markham
1904). Since then, every visitor has recorded them since they grew, and
still grow, adjacent to the boat "landing." A smaller grove evidently
also existed in the south-southwest portion of South Island (Lucett
1851). Palms were also periodically planted--and destroyed--"by whalers
and other chance visitors to the island" (Maude ca. 1938).

Until Arundel’s arrival in 1885, Cocos was basically confined to
this single grove in the northwest sector of South (Maude ca. 1942). In
1885, land clearing began, and from then till 1929 nearly 38,000 palms
were planted, 29,480 between 1916 and 1920 and another 7,000 young trees
after 1927 to replace thousands that had perished (Young ca. 1922).
Arundel’s initial license gave him the exclusive rights to occupy
Caroline and Flint, planting coconuts and other trees for 21 years, in
return for an annual rental of 50 pounds sterling (Maude ca. 1942). In
1929 13,215 trees remained, after which no one has counted them. Our
field work and scrutiny of aerial photographs indicate that far fewer
exist today.

116

Caroline’s plantations produced copra periodically from 1873 to
1934, but never profitably. They suffered greatly from the atoll’s
abandonment from 1901-1916. Dying and poorly planted palms presented
continual setbacks (Young ca. 1922); in 1878 a hurricane wrought great
destruction (N.I.D. 1943), and although no record exists of the effects
of the 1906 hurricane on Caroline, waters reaching well inland on Flint
threw warehouses off their foundations and flooded all buildings within
the settlement (Campbell 1908). In addition, plantation managers
lamented their poor productivity due to choking "by undergrowth and
Pohue Vine,~ destruction of inflorescences by great numbers of seabirds
which roosted in the tops and broke off the flowers as they appeared,"
disease, and ruination of nuts by Polynesian rats and coconut crabs. As
a result of this, the laborers slaughtered many crabs, and "greatly
reduced the numbers of sea birds, who migrated to unoccupied islets."
The rat problem was never resolved and appears to be the major reason
for the eventual abandonment of plantations on both Caroline and Flint.
Their enormous numbers and voracious eating habits greatly reduced both
the crops of potentially healthy nuts as well as the volume of dried
copra. In 1920, 4,600 were trapped on South Island, and hundreds more
were killed by small terriers introduced specifically to control them
(Young ca. 1922). Maude (pers. comm.) recalls that one terrier still
survived in the 1940s. Rats still abound, especially within coconut
groves and Pisonia forests. Another serious problem was due to coconut
crabs digging up recently planted nuts and pinching off young shoots.
After the palms had attained one year’s growth this was no longer a
problem (Young ca. 1922).

Before abandonment (1902 to 1916, and after 1934), Caroline’s
plantations were owned by several companies whose average annual copra
output was approximately 14 tons. From 1934 to the 70s copra was
harvested sporadically by small parties from Tahiti (Garnett 1983), but
within the last 2 decades it stopped altogether.

Despite the relatively fertile soils of South Island, the problems
in the plantations hampered the establishment of permanent settlements
on Caroline. In the 1930s Maude estimated that the atoll could support
400 Gilbertese, increasing to over 1000 "when the island has been fully
planted" (Maude ca. 1938). Colonists were never established though,
since the failure of the plantations was never cured (Maude 1953),
leaving Caroline "one of the least spoiled islands in the Pacific"
(Stoddart 1976). However, as plantation information in Young’s (ca.
1922) unpublished "Memoranda" indicates, Caroline is not as pristine as
it appears, but the rapid recovery of most of its windward forests is
remarkable (see Pisonia Forests, this section).

Ssaid to be Tuumfetta (= Triumfetta) procumbens, most likely a
misidentification of Ipomoea macrantha.

117
Distribution and Abundance

We recognize 4 subdivisions of the coconut woodlands: Cocos
Plantations, Dying Cocos-Ipomoea Plantation, Scattered Groves on Small
Motus, and Mixed Forest with Cocos.

1) Cocos Plantations (34.07 ha)

Superannuated palm forests dominate South Island and southwestern
Nake. Although the planting of Cocos on South eliminated most of its
original habitats, Nake escaped with less damage: Cocos covers 77% of
the area on South but only 6% of Nake (11% including mixed forests).

The 60 to 100-year-old trees form tall, closed canopy woodlands (Pl. 23)
21-25 m high, the customary maximum height recorded for old plantations
(Fosberg 1953). Figure 51 shows the distribution and abundance of plant
species along a transect running centrally through the island, while
Figure 36 depicts a schematic profile of the same swath.

Pure coconut plantations harbor relatively few species: up to 7
trees, zero to 2 shrubs, and 5-11 herbs. The understory layers are
almost exclusively indigenous, an unusual feature. However, skirting
the edge of the lagoon, tall palms overhang the water, crowding native
plants; Suriana and Tournefortia were less abundant in 1988 (Pl. 28)
than in 1965 (Pl. 40).

2) Dying Cocos-Ipomoea Plantation (53.92 ha)

Mature plantations characteristically become overgrown with shrubs
and vines (Fosberg 1953, 1956). Ipomoea macrantha, the sole vine on
Caroline, forms tangled, impenetrable thickets. Indigenous,
nonparasitic, and widely dispersed by ocean currents, it forms a very
minor component of Caroline’s natural habitats, but grows rampantly in
disturbed areas. Vine-covered coconut woodlands cover two-thirds of
South Island’s interior (Fig. 50). This moribund forest is bordered by
a belt of living palms, which in turn are sheltered by a narrow rim of
indigenous vegetation (Figs. 36, 51).

While traversing the South Island transects, the authors stomped
over intertwining thickets up to 3 m high (Pl. 8) and crawled through
tightly-knit masses of vines descending from the crowns of old palms,
Pisonia, and Morinda bushes, until this too, proved impenetrable. In
sunny clearings dotted with dead or disintegrating palms, Ipomoea,
Boerhavia, and Phymatosorus proliferated luxuriantly. Choking of the
palms by Ipomoea, one of the prime reasons for the twice-abandonment of
the copra enterprises, continues to destroy the coconuts, encouraging
natural ecological succession to begin anew.

3) Scattered Groves on Small Motus (0.82 ha)

Drift-derived palms were observed as long ago as 1834 (Bennett
1840). In 1916, when planting operations were commenced after a break
of 14 years, about 40 trees grew beyond the plantations (Maude ca.
1942). Today, small Cocos groves, up to 50 palms, drift-derived and

118

plantation remnants, generally close to the shoreline (Pls. 28, 29),
occur on 11 motus.

4) Mixed Forest with Cocos (6.24 ha)

This forest type is a simplified version of more complex and
varied mixed forests that occur on most inhabited atolls. Composed of
anthropogenic and indigenous elements, it contains a high proportion of
Cocos (50-80%) mingled with variable proportions of Tournefortia,
Pisonia and Pandanus. This forest type occurs primarily in southern
Nake (Fig. 14), but also on Emerald, Shark, and southwest Long, where it
mixes with Cordia and Tournefortia.

House Site: A clearing on Motu Ana-Ana, approximately 40 m x 70 m,
contains a few Cocos, a vegetable garden and thatched former living
quarters (Pl. 51).

Seabird Use

Cocos-dominated habitats were ornithologically the most
depauperate on Caroline: only Brown Noddies and White Terns breed. The
noddies nested high within the frond and inflorescence bases, whereas
the White Terns preferred lower sites, occasionally atop an arching
frond. The absence of other species suggests that anthropogenic Cocos
forests seriously inhibit seabird use and may continue to do so for
decades until they are replaced by native vegetation.

Absent Plant Communities

Caroline’s impoverished flora and relatively simple physiography
and geology has resulted in a limited variety of ecosystems. The atoll
is thus notable not only for its Pisonia forests, extensive monotypic
stands of Tournefortia, and Cordia groves, but also for the absence of
several ecosystems that are generally considered typical of Pacific
atolls:

1) Sesuvium Flats;

2) Pemphis, Scaevola, and Sida Scrub (2 Scaevola plants are present, and
the only 2 Sida records are from 1884 and 1990); and

3) Mixed broadleaf forests including Barringtonia, Calophyllum,
Guettarda, Hernandia, and Neisosperma.

4) Plant associations (except Cocos) typical of native cultures on
atolls: breadfruit groves (Artocarpus altilis), taro pits
(Cyrtosperma chamissonis, Colocasia esculenta, Xanthosoma
sagittifolia), cultivated ornamentals (Hibiscus rosa-sinensis,
Plumeria spp., etc.), or weedy grasslands/wastelands (Paspalum,
Sporobolus, Wedelia, Vigna, etc.). Even widespread introduced strand

119

species such as Jerminalia catappa and Casuarina equisetifolia are
absent.

In addition, there are no mangroves, peat bogs, marshes, ponds, salt
flats, or other habitats associated with fresh or brackish water.
Poorly represented are:

1) Lepturus Grassland. Although Lepturus is present in coastal herb
mats, and occasionally in patches within the forest understory, it
does not form a separate plant community. However, it may once have
covered the extensive clearings on South Island (Pl. 2, 3).

2) Mixed Forest. Though 6.24 ha of Mixed Forest (with Cocos) occurs
(primarily on Nake), it is of such minor importance to Caroline’s
overall vegetation that it is treated as a subsection of Coconut
Woodlands.

H. DESCRIPTION AND ECOLOGY OF THE MOTUS

These islet accounts synthesize the history, physiography,
vegetation patterns, ecology, seabird colonies, miscellaneous biota, and
the effects of human activity (if any) on Caroline’s 39 motus (Fig. 2).
Mapping is based on the coast-to-coast transects, perimeter surveys,
complete surveys (smaller motus), color transparencies, and aerial
photographs.

All motus are detrital reef islets representing many evolutionary
stages from barely emerged coral rubble to large islets with relatively
fertile "soils" supporting lush vegetation. There is one tiny old reef
platform in its final stages of erosion.

We discuss and map them in geographic order beginning in the north
with Nake and progressing down the windward reef through Long and the 13
Windward Islets to South Island. Beginning anew in the north, we move
south through 7 South Nake Islets, 11 Central Leeward Islets, and
finally the 5 Southern Leeward Islets.

Because of the variety of islet shapes, "long" or "length" refers
to the longest dimension lying parallel to the outer reef edge (normally
north-south) and "wide" or "width" to the longest dimension
perpendicular to the outer reef edge (normally east-west). South
Island, the only exception, is considered to lie adjacent to the
southern reef edge, so its "length" is measured east-west. Seabird
numbers are from Part II, Table 1. For convenience in locating
particular islets, the order is as follows:

120
1) NAKE (Fig. 37)

WINDWARD ISLETS
3) Bo’sun Bird (Fig. 42)
5) Crescent (Fig.43)
7) North Pig (Fig. 44)
9) Skull (Fig. 44)
11) Brothers (Fig. 46)
13) North Arundel (Fig. 47)

2) LONG (Fig. 38)

4) Windward (Fig. 43)

6) Atibu (Fig. 43)

8) Pig (Fig. 44)

10) North Brothers (Fig. 46)
12) Noddy Rock (Fig. 47)

14) Arundel (Fig. 47)

15) Tridacna (Fig. 48) 16) SOUTH (Fig. 50)
SOUTH NAKE ISLETS

17) Pandanus (Fig. 52) 18) Danger (Fig. 52)
19) Booby (Fig. 52) 20) Coral (Fig. 52)
21) Lone Palm (Fig. 52) 22) Kota (Fig. 52)

23) Mouakena (Fig. 52)

CENTRAL LEEWARD ISLETS

24) Mannikiba (Fig. 53) 25) Blackfin (Fig. 54)
26) Matawa (Fig. 54) 27) Emerald (Fig. 54)
28) Shark (Fig. 55) 29) Scarlet Crab (Fig. 55)
30) Nautonga (Fig. 55) 31) Azure (Fig. 55)
32) Reef-flat (Fig. 55) 33) Bird: (Fige.55)

34) Fishball (Fig. 55)

SOUTHERN LEEWARD ISLETS

35) Raurau (Fig. 57) 36) Eitei (Figs 57)
37) Pisonia (Fig. 57) 38) Kimoa (Fig. 57)
39) Ana-Ana (Fig. 57)

1) NAKE ISLAND (91.72 ha) Figs..30, 37; Pls..17,. 22, 35=37—mens
History: Nake’s large size and underground water lens, coupled with
topography and soils more varied than elsewhere on Caroline, attracted
Polynesian settlers. Because early European visitors stayed primarily
on South Island, there is only a single reference to Cocos prior to the
late 19th century (1 tree seen in 1825 by Paulding 1831).

The far northwest of Nake (also North Island in Young ca. 1922)
houses the only true archaeological site on Caroline--a large marae
(Figs. 3, 37; Pl. 36). Discovered during the guano era, the site is
marked as "graves" on Arundel’s map. Arundel, who was living on the
atoll when the marae was discovered, describes it thus: "On the north-
west end of Caroline are some curious old native remains, whether places
of burial or of sacrifice I cannot determine. I opened one of these,
but could find no indication whatever to guide me in a decision"
(Arundel 1890). AKK, R. Falconer, and G. Wragg located, measured, and
photographed this marae in 1990. The entire courtyard was approximately
18 m long by 14 m wide. All 10 peripheral stones and the central one
were easily identifiable from the 1883 plan (Fig. 3), although a few had
fallen over or broken due to encroaching vegetation. The lower wall,
partly destroyed by Arundel, had not been reconstructed. It is probable

2

marae

ocean

ANCIENT
CHANNEL

~
J

Figure 37. Nake Island:

121

RUBBLE
WB HERB Mar
CORDIA

[__] TOURNEFORTIA
PISONIA

[==] Cocos
PANDANUS
MIXED

BEACHROCK

ocean

vegetation and physiography.

122

that this marae had not been seen since the 1880s; though discussed by
Emory (1947), he never visited Caroline personally.

Northwest Nake is particularly suitable for a place of worship and
sacrifice: it fits most of the environmental criteria indispensable to
ancient Tuamotuan religious ritual (Emory 1947). First, flat ground was
necessary, preferably lying at right angles to, or parallel to, the
lagoon. Second, it was important to have the wind blowing across the
marae to waft away the smells of sacrificed animals. Third, ceremonial
items included branches of the Pisonia tree, leaves of Cocos (for leaf
charms/"rosaries"), and the aerial roots of Pandanus. Fourth, feathers
from "black terns" (Black Noddy), frigatebirds, and Red-tailed
Tropicbirds were also necessary for rituals. Rather than a smooth
substrate, the early Polynesians would have had to be content with
leveled coral rubble and distance from the lagoon. The only organism
not living near the marae today is the tropicbird; however, their
elongated tail feathers could have been plucked from adults nesting on
nearby motus.

Since marae are sacred places, there is possibly a significance to
the location of the main "courtyard" close to the atoll’s northern tip.
It is well known that the northern extremities of islands were
auspicious places for all Polynesians; in such places, they believed,
disembodied spirits were whisked to the netherworld.

In 1938 a cistern was built in southwest Nake, which is still
visible (see Sect. D, Hydrology). We failed to find evidence of
occupation, but in September 1974 there was "a small, barely-furnished
thatched hut," a stack of approximately 3 tons of copra on a raised
platform, and a "three fathom canoe of Polynesian construction" (Ward
1974).

Physiography: Largest in area, Nake is the northernmost motu, separated
from Long by a 40-m channel (Pl. 17). With maximum dimensions 2,000 m
long and 685 m wide, it is basically rectangular with rounded corners
and a peninsula-like extension in the southeast.

Nake lies north of the lagoon, having a southern, dry "bay" (Sandy
Inlet), in which silt, sand, and fine coral debris are being actively
deposited (Pl. 22). This hard, flat expanse of silty and sandy
sediments is 145 m wide at its mouth, extending 200 m north into the
main islet. Its 3.50 ha provide a favorite feeding location for
shorebirds, especially Bristle-thighed Curlews. If Arundel’s chart
(Fig. 4) is correct, Sandy Inlet has increased its land area during the
last century.

On the reef flats off the west side are extensive remnants of
jagged upraised reef of unknown age (Pl. 11) and occasional beachrock.
The exposed beaches and reef flats at Nake’s north point are especially
broad, characteristic of reef flats at the exposed corners of islands.
Comparisons of the northern sweep of rubble on recent aerial photos with
Arundel’s map indicate that much coral debris, curved shingle ridges
(Pl. 16), has been added since 1883. This area, the northernmost tip of

123

the atoll, is subjected to heavy wind, wave and swell action. It is
possible that particular ridges can be attributed to individual storms
as has been documented for some other atolls (Stoddart & Steers 1977).
In 1990, the deep, fine coral rubble mixed with sand east of the marae
yielded 3 old turtle nests. Overall, only 6% of the land area was
unvegetated. However, sparsely vegetated expanses of hardpan occupied
the south-central sector (immediately inland of the coast within a belt
of Tournefortia forest), and pure sand at least to 0.5 m deep bordered
Sandy Inlet.

Nake’s windward coast, complete with a peaked beach crest and
discontinuous beachrock, is 30 m wide in the north, narrowing to 3 m in
the south. Offshore, submerged reef flats form a sandy moat.

In the distant past, Nake consisted of 2 separate motus: aerial
photos (Frontispiece) reveal an oblique, ancient channel about two-
thirds of the way down the isletwhich is now well vegetated centrally
but scrubby peripherally.

Vegetation: Before the major clearing for coconut plantations, Nake’s
native forests were "80 to 100 feet high" (Arundel 1875). Today there
are 16 plant species (5 trees, 1 shrub, 10 herbs), 62% of Caroline’s
flora. It is the lushest motu, with woodlands (82.39 ha) about 80%
native and 20% Cocos (Pl. 37). Although in 1916 there were about 260
palms, and the entire island was evidently planted with 10,544 palms in
1918-1919 (Young ca. 1922, Table 13), substantial tracts of each major
vegetation type occur today. Its interior is rich in Pisonia, with the
largest acreage (20.79 ha) and some of the tallest trees (20 m high) and
largest trunks on the atoll (Pl. 43, Table 11). In addition, Cordia is
well-represented: 2 major groves of Cordia-Tournefortia forest occupy
11.8 ha, 2% of Nake’s area. Extensive pioneer herb mats, flanked on
their inner sides by Tournefortia scrub, occur in the north and east.
The remaining Cocos, essentially in the southern quarter, comprise
Caroline’s second largest coconut grove.

Birds: Nake, with 80% of Caroline’s breeding seabird species, shows a
direct correlation between islet size and bird species diversity. Nine
species of seabirds breed, all with larger populations (pairs) than
previously reported (Clapp & Sibley 1971la): Masked Booby (105), Brown
Booby (1), Red-footed Booby (496), Great Frigatebird (522), Lesser
Frigatebird (56), Brown Noddy (390), Black Noddy (814), Sooty Tern
(nesting in 1989; Anne Falconer, pers. comm.), and White Tern (1,094).

2) LONG ISLAND (75.98 ha) Rigs 30; 35) esS8—4il's)/Piis7 29%
WW MNES alls Irae Ts ty

Third largest in area, this longest of motus covers nearly one-
third of the atoll’s windward side. In the north it is separated from
Nake by a narrow channel; from its southern tip a chain of smaller motus
extends south along the windward reef.

124

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Cocos nucifera
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floristic composition, relative abundance of plant species, degree of species overlap, and canopy heights.
Vertical height is exaggerated.

Tournefortia- Pisoria Tournetortia rubdie

Boerhavis repens
Cocos nucifera

Cordia subcordate
Heliotropium anomalum
Lepturus repens
Morinda citrifolie

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Pisonia grandis
Portulece tutes

Tgurnetortia argentea

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Figure 41. Long Island: east-west cross-section through Transect 8, which passes through
mature interior Pisonia forest of largest of Long's coalesced motus. Data includes floristic
composition, relative abundance of plant species, degree of species overlap, and canopy
heights. Vertical height is exaggerated. Note the absence of low veyetation on the leeward
shore.

127

Physiography and History: Long, 4,226 m long and 330 m wide, is
somewhat snake-shaped, with an enlarged northern "head" and attenuated
"tail." From a distance its vegetation appears as a series of humps.
Long has experienced a fairly complex geological history, noted by the
Solar Eclipse Party: "On some of the islands there are spaces void of
vegetation, extending from lagoon to beach, which indicate the existence
at a former time of a water separation" (Holden & Qualtrough 1884).

At present, Long is composed of 5 distinct former motus separated
by sparsely vegetated channels of coarse sand and coral gravel. Aerial
photographs also reveal further, older subdivisions (see below).
Coalescence and fracturing of the original motus probably occurred
repeatedly. Since erosion proceeds faster on an atoll’s windward reefs,
providing coral fragments, coralline algae, and pulverized molluscs, it
is no surprise that the first series of Caroline’s motus to fuse were
those facing this rich source of parent material.

Long’s coarse rubble beaches (Pl. 19) are a mirror image of those
on Nake: they widen progressively southward. The swath of unvegetated
rubble above high tide line in the upper two-thirds of Long averages 8 m
wide, while in the lower third it is 40 m wide. Unvegetated coral
debris accounts for 10% of the island’s area (Fig. 30). Beachrock,
flanking the windward shoreline for most of its length, is more abundant
than elsewhere (Pl. 54).

Long’s lagoon flank is edged with submerged sand and silt, and is
one of the most sheltered parts of Caroline. Sand and rubble deposition
off the south point has formed a lagoon islet (Bo’sun Bird), which
could, in the future, coalesce with Long’s south point to form a hook.

An uncommon substrate on Caroline, upraised reef forms a low
rampart (generally <1 m high) paralleling the ridge crest inside the
vegetation for much of the lower quarter of Long. Although we camped in
this area and conducted 4 transects through this upraised reef, we found
no plant species that indicated the presence of feo, such as can be
found in the Tuamotus (Fosberg, pers. comm.).

In 1990 G. Wragg found some scattered large stones, similar to
those of the Nake marae, located centrally 100 m north of the southern
tip of Long, confirming the report of the remains of a smaller marae on
Long Island (Holden & Qualtrough 1884). Wragg noted that the marae was
small, measuring approximately 3 m wide by 8-10 m long. Its orientation
appeared to be northeast-southwest. The wall on one end was evidently
smashed by storm waves. Only 2 of the peripheral upright stones were
standing, of similar size to those on Nake. The platform was in
reasonable condition, with a huge Pisonia tree growing through it. Some
rock slabs were large (2 x 2m). The entire marae was situated within a
Pisonia grove, with nearby Cocos. We do not know if this coconut grove
(1.6 ha) was present before 1,343 palms (20% of the islet’s area) were
planted in 1918-19 (Young ca. 1922). The sheltered location and a
Pisonia-Cocos forest, which suggests an old clearing, further indicate
prior occupation.

128

In 1990, Wragg also uncovered an RNZAF survey marker just inland
of Long’s southernmost tip.

Vegetation: There are 15 plant species (4 trees, 2 shrubs, 9 herbs), on
Long, 58% of the total flora. Long’s variety of habitats, vegetation
heights, substrata, and birds make it the most diverse islet on
Caroline. Only 3% of its area remains in Cocos. All the atoll’s
seabirds have bred here. Its ecology is best understood with reference
to Figures 35 and 39-41.

Within the basic pattern of 5 coalesced motus, it may be seen
that:

1) From north to south (measured from the midpoint of each former
channel) the motus, of divergent size and shape, are approximately
320, 620, 700, 1,840, and 100 m long.

2) Each former motu, crowned by Pisonia forest, contains concentric
rings of decreasing fertility around its core and is morphologically
similar to motus surrounded by water, except that the coarse coral
gravel along the former perimeter is less marked. More specifically,
beach sands and gravel extend for 200-300 m north and south of the
old channels, after which they increasingly accumulate coral rubble,
humus, and guano.

3) Tournefortia dominates, interspersed with 4 patches of taller Pisonia
forest and scattered clumps of Cocos and Cordia. Interrupted herb
mats parallel the windward coast and often extend across the island
along former interislet channels (Pl. 32). Vegetation height varies
from 2 cm to 15 m.

4) Plant species diversity is highest in Tournefortia-Pisonia and lowest
in Pisonia forests.

5) Long’s tallest, most mature Pisonia groves (up to 100% Pisonia) occur
on the largest of the former islets. The Pisonia forest near the
south end (Tr. 10), although healthy, is only 12 m tall. This may be
due to its impoverished upraised pitted reef substrate barely covered
with "soil." Since it lies adjacent to Long’s most luxuriant Cocos
grove, its land could well have been cleared in 1918-19, with the
Pisonia forest taking longer than elsewhere to recuperate. Because
tern guano increases soil fertility, contributing to Pisonia growth,
it is of interest that neither Black nor Brown Noddies nested here.

6) Deep dips in Figure 39 (lower graph) correspond to east-west
corridors formed from old channels. Vegetation in these relatively
infertile, sandy flats is low, similar to that on small developing
motus (i.e. native herbs with scattered Tournefortia <2 m high). One
sandy channel (Tr. C, Pl. 32) supported sparse Suriana. During the
February 1990 cyclone, all vegetation was either uprooted, washed
away, or smothered with fresh sand and coral gravel along Transects A
and C (Pl. 33). Storm erosion was particularly marked within the
channel that almost bisects the island (Tr. A).

129

7) Secondary dips mark even older interislet channels ("ancient
channels"), visible on aerial photographs (Frontispiece) but barely
recognizable in the field. They are overgrown with Journefortia
and/or Pisonia.

8) Sharp dips within established forests or herb mats denote relatively
recent channels gouged out by storms ("recent storm cuts"). These
were also altered during the winter 1990 storm.

Figures 40 and 41 illustrate some differences between the windward
and leeward coasts. Transect C (Fig. 40) crosses the north end of Long
through an old interislet channel now filled with sand and rubble. Its
low profile reflects the simple habitat harboring halophytic herbs and
Tournefortia shrubs less than 2 m high. Although the shrubs are
scattered, the lagoon half of the transect passes through slightly
higher ground, which encourages denser Journefortia. This transverse
section is similar to that of a formative motu such as Fishball
(Fig. 56). This exposed, scrubby swath, 300 m wide, harbors Red-footed
Boobies, Great Frigatebirds, and a discrete population of Masked
Boobies. Approximately 127,000 pairs of Sooty Terns nested in a similar
sandy channel 740 m to the south (Tr. A) in 1988.

Transect 8 (Fig. 41) crossed the islet nearer the southern tip
(Fig. 8). This profile departs significantly from the usual parabolic
cross-section seen on most of the small motus and which exists further
north on Long Island. From east (windward) to west, there is first a
wide expanse of coarse, unvegetated rubble, followed by rubble dotted
with herbs, then Tournefortia scrub increasing to 9 m high. Further
inland, a forest of 10-m-high Tournefortia, Pisonia and Cordia continues
westward to the lagoon. This leeward margin of Long, extending
southward nearly to its tip, is the only location on Caroline where
tall, indigenous vegetation overhangs and shelters the lagoon. No herb
mat is present.

In summary, Long contains examples of all major plant communities,
as well as 2 minor ecosystems, Pisonia-Cordia (3.2 ha) and Cocos-Cordia
(0.82 ha). Its woodlands total 49.60 ha. Coconut crabs inhabit al]
areas containing Cocos and Pisonia; our rough population estimate is
200.

Birds: In 1988 Long supported 9 (10 in 1965) species of breeding
seabirds, as follows (pairs): Red-tailed Tropicbird (5), Masked Booby
(69), Brown Booby (12), Red-footed Booby (659), Great Frigatebird (808),
Sooty Tern (179,800), Brown Noddy (207), Black Noddy (986), and White
Tern (751). From 1988 through 1990, Sooty Terns occupied 19 large
colonyysites (hig.. lls. Pt. Mil).

Comments: Polynesian rats were abundant, especially in Cocos and
Pisonia habitats. It was often possible to see 3 or 4 simultaneously
while conducting daily surveys, and 20 or more around camp. At night,
their numbers increased substantially. Azure-tailed skinks (Emoja
cyanura) were noted.

130
WINDWARD ISLETS

This chain of 13 islets occupies the southern half of Caroline’s
east coast. All rest on the same reef flat, separated by channels
varying in width and depth. They can be waded with care at low tides,
but most harbor black-tipped reef sharks: up to 4 were visible in the
shallows within 50 m of an observer. Several motus have altered shape
since 1883, including Brothers, which has incorporated a small cay into
its present confines.

The motus range in size from Noddy Rock (0.02 ha) to Windward
(11.42 ha). They support every major vegetation type from simple herb
mats to Pisonia forests, 21 m tall. Because of their constant exposure
to trade winds, the seaward vegetation is wind- and salt-shorn. Though
appearing primarily untouched, all of the Windward Islets were planted
with Cocos (Table 13) from 1916-20 (Young ca. 1922). However, these
incipient plantations experienced difficulty and appear to have been
abandoned within a few years, and their vegetation recovered remarkably
(see Sect. G).

Flanking the lagoon of the southern motus (Brothers through
Tridacna), and extending westward, are reefs densely studded with
Tridacna clams, which add to Caroline’s outstanding natural assets
(PT. 253 Pt. 11; 4Sect. a) -

3) BO’SUN BIRD ISLET (0.86 ha) Figs. 29, 42

We named this motu for its Red-tailed Tropicbirds, commonly called
Bo’sun birds. The sizeable population is the largest on Caroline. In
addition, our 1988 records constituted the first known breeding of this
species on the atoll.

Physiography: Bo’sun Bird Islet, 165 m west of Long’s southern tip, is
the only motu lying within Caroline’s lagoon. It shares the same reef
as Long, however, and is not a true "lagoon motu."

Amoeboid in shape, Bo’sun Bird is greatly affected by the tidal
waters that spread across the shallow reef flats and gush through the
channels separating Long and Windward. Because it sits near the inner
edge of a wide windward reef flat, the layering of sediments around it
is complex and transitory; our observations indicate that more rubble
was deposited on the islet’s western edge since the aerial photos were
taken in 1985. Its western shoreline rises gradually to a high water
mark, and slight changes in water level greatly change its overall size
and shape. At high tide its perimeter is ovoid with a long westerly
extension. The "head" is approximately 70 m wide and 115 m long, while
the "nose" is 45 m long and 15 m wide.

Vegetation and Birds: Bo’sun Bird Islet, composed of coral rubble and
sand, supports only natural herb mats (Heliotropium, Portulaca,
Lepturus) and Tournefortia scrub (to 4m tall). These 2 simple plant
communities cover 35% and 55% of the land area, respectively. For its

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132

size, the motu is sparsely vegetated, with only 4 plant species (1
shrub, 3 herbs), 15% of Caroline’s total flora. There are no
introductions. 7

Bo’sun Bird’s most notable attributes are its 4 species of
breeding seabirds: Red-tailed Tropicbird (47 pairs in 1988, 130 pairs
in 1990), Sooty Tern (8,400 pairs), Brown Noddy (10 pairs), and White
Tern (6 pairs).

4) WINDWARD ISLET (11.42 ha) Figs. 29, 43

We named this "Windward" because it is the first major, and
largest, Windward Islet.

Physiography: Broadly crescentic in shape, 508 m long by 287 m wide, it
parallels the reef’s longitudinal axis and is set close to the lagoon.
Its seaward beach is quite narrow (3 m wide); there is no lagoon beach.

Vegetation: Windward has 11 species of plants (3 trees, 1 shrub, 7
herbs), 42% of the total flora. A windward crescent of halophytic herbs
borders a zone of Journefortia scrub, which mixes quite densely with
Pisonia and Cordia over most of the interior in a bilobed pattern.
These latter forests, reaching 14 m high in the south and 9 m in the
north, total 8.67 ha. This unusual distribution of central forests
undoubtedly reflects Pisonia’s recovery from 100% land clearing for
Cocos--1,299 palms--in 1920 (Young ca. 1922, Table 13). It is
remarkable that not one Cocos remains as a legacy of this disturbance.
The east-west profile of Windward, similar to that of Transect 8,
Long Island (Fig. 41), is typical of most motus, except that lagoon-
facing herb flats are almost nonexistent. Scaevola taccada var.
taccada, a new plant record for the atoll, was only found on this motu,
although S. t. var. tuamotensis was found on South Island in 1990.

Birds: Five species of breeding birds were present, all in appreciable
numbers (pairs): Red-footed Booby (163), Great Frigatebird (207), Brown
Noddy (20), Black Noddy (28), and White Tern (134).

Comments: In May 1990, AKK noted a possible motu midway between
Windward and Crescent Islets during midtide. It appeared an upraised
reef platform like Noddy Rock, but because of extensive shallow
reticulate reefs in this area, its presence at high tide has not yet
been confirmed.

5) CRESCENT ISLET (3.10 ha) Figs. 29, 43
We named this islet for its cupped shape.

Physiography: Crescent Islet is 190 m long by 225 m wide. It is almost

entirely composed of coral rubble, with a little humus in the interior.

The seaward beach is variable (up to 50 m wide), the lagoon beach,
insignificant.

133

Seq RUBBLE Ps
ee HERB MAT \
CORDIA in
[__] ToOuRNEFORTIA ie

f a Fy

PISONIA nex
TOURNEFORTIA-CORDIA ~

WINDWARD ISLET

ocean

lagoon

CRESCENT ISLET

MOTU ATIBU

Figure 43. Vegetation and physiography of Windward Islets
nos. 2, 3 and 4: Windward and Crescent Islets, and Motu
Atibu ("Coral Rubble Islet"). Atibu appears to have been
severely damaged during the February 1990 storm.

134

Vegetation: There are 10 species (3 trees, 1 shrub, 6 herbs), 39% of
Caroline’s flora. No introduced plants occur. Plant diversity is
poorer than on Windward, a reflection of small size, poor soils, and
scant herb mats. However, woodlands cover two-thirds of its area, and
the central stand of Pisonia and Cordia is 87 m wide and up to 13 m
high. Crescent was heavily planted (80% of total area, 228 palms) in
Cocos in 1920 (Table 13), but today none remain.

Birds: Crescent Islet was used by the following numbers of breeding
pairs: Red-footed Booby (28), Great Frigatebird (5), Brown Noddy (36),
Black Noddy (60), and White Tern (8).

6) MOTU ATIBU ("Coral Rubble Islet") (0.02 ha) Figs. 275.42

Motu Atibu was Caroline’s smallest and least vegetated islet.
Third in the windward chain, it measured 13 x 18 m. We named it for its
basic rubble character. Vegetation covered only 2% of the land surface
and consisted of a few Journefortia shrubs (<1 m high) encircled by
narrow swaths of low herbs and rubble. Its 3 plant species (1 shrub, 2
herbs)--12% of Caroline’s flora--were among the most meager on the
atoll. Atibu’s profile was similar to that of Fishball (Fig. 56).
There were no breeding birds.

Comments: Since a February 1990 storm, Atibu has apparently
disappeared, having been reduced to a thin strip of coral gravel below
high tide level.

7) NORTH PIG ISLET (5.44 ha) Figs. 29, 445) \Plisias

We named the fourth windward islet "North Pig" for its location
immediately north of Pig Islet.

Physiography: Classically crescentic, North Pig is 350 m long and 230 m
wide. Though approximately half of Pig’s area and less wooded overall,
North Pig has a similar distribution of sediments (including sand on the
lee side), vegetation, and breeding birds. Profiles of the 2 motus are
nearly identical (Fig. 45).

Vegetation: There are 11 plant species (3 trees, 1 shrub, 7 herbs), 42%
of Caroline’s flora. No introduced plants are present. Proceeding
south along the windward islets, lagoon-side herb mats develop and islet
cross-sections assume a more perfect symmetry--low at the edges and
forming a hump in the middle.

North Pig’s 3 vegetation zones are predictably symmetrical: a
peripheral band of herbs (more extensive on the "horns"), curved belts
of Tournefortia, and a spacious central forest of mixed Pisonia, Cordia,
and Tournefortia. The latter (to 20 m tall) covers more than one-half
the islet’s width and one-third its area, and includes fine Cordia
groves (Fig. 44). This excellent forest is surprising because 402 Cocos
palms were planted on 93% of North Pig’s usable land in 1920 (Young ca.

RUBBLE

HERB MAT
CORDIA
TOURNEFORTIA
PISONIA

Cocos
SURIANA

ae
HOBOS

NORTH PIG ISLET

PIG ISLET

lagoon

SKULL ISLET

NORTH BROTHERS
ISLET

BROTHERS ISLET

meters

135

ocean

Figure 44. Vegetation and physiography of Windward Islets
nos. 5 through 9: North Pig, Pig, Skull, North Brothers,
and Brothers Islets. Note the reefs extending westward

into the lagoon.

136

1922, Table 13). Measurements from 25 Pisonia trees (main trunks)
averaged 19 m in height, 221 cm cbh and 261 cm in base circumference
(Table 12).

Birds: Five species of seabirds bred: Red-footed Booby (31 pairs),
Great Frigatebird (17 pairs), Brown Noddy (76 pairs), Black Noddy (3,199
pairs), and White Tern (110 pairs). The largest colony of Black Noddies
on Caroline nested in the tall Pisonias.

Comments: Rats and coconut crabs were common.

8) PIG ISLET (7.21 ha) Figs. 295) 44; Pls» 26, 41, 50ju5a

Number 5 down the chain, Pig was named prior to 1883. Domestic
pigs were introduced to Caroline in 1828 by Captain Stavers but
evidently died out before 1834. Reintroduced in 1848 with the first
recorded settlers, it is not known how long they lasted. One would
expect that they were only on South Island, but the statement that
"about one-third the distance up the lagoon a canvas hut exists on one
of the smaller islets on the eastern side of the lagoon" (Holden &
Qualtrough 1884) suggests that perhaps domestic animals also inhabited
Pig. Though this is weak evidence, there must have been some reason for
this curious name. Today no pig devastation is evident anywhere on the
atoll.

Physiography: Bean-shaped, Pig is 330 m long and 255 m across. It is
separated from North Pig by a channel 60 m wide.

Vegetation: The islet has 11 plant species (4 trees, 1 shrub, 6 herbs),
42% of Caroline’s flora. Cocos, the only introduction, is rare

(0.03 ha). In 1920, 538 palms were planted (Young ca. 1922), which
covered approximately 79% of Pig’s usable area (Table 13).

Pig’s vegetation profile (Fig. 45) is classic: a wide, windward
herb mat, bordered by Tournefortia and Cordia, which, in turn, grades
rapidly into an outstanding Pisonia forest (to 21 m tall, 3.36 ha), one
of Caroline’s best groves. Measurements from 5 trees, mostly multiple-
trunked, averaged 16 m in height, 338 cm in circumference (at 1.5 m),
and 282 cm in base circumference (Table 12). This Pisonia also occupies
the largest proportion (46%) of any islet area. It is striking that
such quality forests could regenerate in about 65 years (see Sect. G).
In the Cordia forest (Pl. 26), also the finest on Caroline, 6 trees
averaged 12.6 m in height, 116 cm in circumference (at 1.5 m), and
99.8 cm base circumference. On the lee side of Pig, JTournefortia
extends directly to the lagoon shore.

Birds: Five species of seabirds bred: Red-footed Booby (14 pairs),
Great Frigatebird (118 pairs), Brown Noddy (82 pairs), Black Noddy
(1,928 pairs), and White Tern (164 pairs).

137

Tournefortia Pisonia Tournefortia Herb Mat

Achyranthes canescens
Boerhavia repens

Heliotropium anomalum

Ipomoea macrantha

Morinda citrifolia
1 !
Pisonia grandis

|
|

|

Phymatosorus scolopendria

Sam

|
{ 1
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Tournefortia argentea

€

—

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S

WW

x 20

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5 10

Zz

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(0) 100 200

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Figure 45. Pig Islet: east-west cross-section through center of
islet. Data includes floristic composition, relative abundance of
plant species, degree of species overlap and canopy heights.
Vertical height is exaggerated. Pig's profile is especially
symmetrical. It is remarkable that this islet was totally felled
for coconuts in 1920.

138

Comments: Rats and coconut crabs were common. In 1990 a grayish gecko
(possibly mourning gecko, Lepidodactylus lugubris) was seen by
A. Garnett. ;

9) SKULL ISLET (0.02 ha) Figs. 27, 44; Pls. 46, 49

Sixth in the windward chain, we named Skull] Islet after finding
the skull, tail feather, and eggshell of a Red-tailed Tropicbird, the
first evidence that this species bred on the atoll. A low shelf of
coral rubble and sand, barely above high tide mark, this motu is barren
except for a small herb mat under 5 Journefortia bushes (1 m high) on
the lagoon side. Only 2% of the surface area is vegetated. There are 3
plant species (1 shrub, 2 herbs), 11% of the atoll’s flora. Although
appearing young, the motu is marked on Arundel’s chart (Fig. 4). After
February 1990, several large storm blocks rested in the channel close to
Skull Islet.

In 1988 there were no birds. However, in March 1990, a colony of
150 Brown Noddies was in a prelaying phase, accompanied by 6 Sooty
Terns, a Brown Booby and a Wandering Tattler.

10) NORTH BROTHERS ISLET (1.71 ha) Figs. 29,°°44) "Fi asa

The seventh windward motu, we named this islet North Brothers
because of its location directly north of the named motu, Brothers.

Physiography: North Brothers is shaped like an oval that curves
lagoonward toward Brothers, 40 m away. The concave shorelines and lack
of herb mats on the opposite shorelines of these 2 islets suggest that
they might have been formerly connected. Composed primarily of rubbly
substrates, with slightly better soils centrally, it is 95 m long and
250 m wide.

Vegetation: Plant species number 10 (3 trees, 1 shrub, 6 herbs), 39% of
Caroline’s flora. A few Cocos trees are present, remains of the 180
planted in 1920 (Young ca. 1922), which covered 100% of all available
land on the islet (Table 13). Plant communities on North Brothers are
simple: Tournefortia (more open in the west) rises to an excellent
Pisonia forest, 80 m wide and 18 m tall, on the east end. Average
measurements from 3 Pisonia trees were: height 18 m, base circumference
314 cm, and number of trunks, 2.3 (Table 12).

Birds: Five species of seabirds bred on the islet in 1988 (pairs):
Red-footed Booby (25), Great Frigatebird (9), Brown Noddy (23), Black
Noddy (40, plus hundreds of old nests), and White Tern (69). In
September 1989, Sooty Terns nested on the windward beach (Anne Falconer,
pers. comm.), and in May 1990, a prebreeding swirl of thousands of Sooty
Terns swarmed above Brothers and North Brothers.

Comments: Gecko eggs were seen on Pisonia trunks in 1990.

139
11) BROTHERS ISLET (4.31 ha) Figs. 29,°44, 46; Pls. 29, ‘55

The eighth windward motu, Brothers Islet was named last century
after Captain Brothers, who managed a stock-raising venture on Caroline.
In 1873 his rights to the atoll passed into the hands of John Arundel.

Physiography: Crescentic in shape, with longish horns extending toward
the lagoon, Brothers Islet lies about two-thirds of the way down
Caroline’s windward reef. It is 198 m long x 178 m wide through the
center. A Tridacna reef extends westward almost completely across the
lagoon.

An interesting aspect of Brothers’ structure is that Arundel’s
chart (Fig. 4) indicates a tiny, separate motu off the southwest point.
Our survey and the 1986 aerial photos show that this motu is now joined
to Brothers Islet. Its former identity is marked by a small patch of
Tournefortia, around which the recently deposited sand and rubble is
sparsely dotted with native herbs.

Vegetation: There are 11 plant species (4 trees, 2 shrubs, 5 herbs),
42% of Caroline’s flora. Cocos, along the leeward shore, is the only
introduced plant. Three distinct plant communities are present:
peripheral herb mats (including leeward Portulaca with Suriana),
Tournefortia scrub and forest (to 6 m high) bordered with Cordia, and a
central Pisonia forest. Larger trees had up to 15 trunks and multiple
root suckers. Measurements of 10 trees (main trunks) averaged 15 m in
height, 140 cm in circumference (at 1.5 m), and 243 cm base
circumference. Distances to nearest neighbor for 10 trees averaged 4.2
meters. As on its neighbor islets, the Pisonia forest on Brothers is
striking, especially since it has matured to a closed-canopy monotypic
stand devoid of any subcanopy species (Fig. 46), evidently in about 65
years. In 1920, Brothers Islet was planted with 315 Cocos palms, which
covered approximately 97% of the usable land area (Table 13).

Birds: Four species of seabirds bred: Red-footed Booby (25 pairs),
Brown Noddy (8 pairs), Black Noddy (15 pairs), and White Tern (50
pairs). In May 1990, large numbers of Sooty Terns swirled over Brothers
and North Brothers.

Comments: Many mature Pisoni7a trees contained capacious cavities in
their boles that housed large coconut crabs. In March 1990, several of
these holes had feathered skeleta of Sooty Terns (and possibly also
Brown Noddies) outside their entrances, along with freshly-snipped
Pisonia branches (see Pt. II, Sect. F).

12) NODDY ROCK (0.02 ha) Figs. 27, 47; Pl. 18

We named this ninth motu in the Windward Islets for its only known
breeding seabird, the Brown Noddy. In September 1988, at least 80 pairs
were incubating their eggs on the Portulaca mat that covers its central
lee section.

140

Portulaca mat

: Tournefortia

Tournefortia

Pisonia
Herb mat

Boerhavia repens
Cocos nucifera
Heliotropium anomalum
Laportea ruderalis
Lepturus repens
Morinda citrifolia
Pisonia grandis

Portulaca lutea

Suriana maritima

Tournefortia argentea

ey 1
=
o
= 20
a
O 10.1
z
<_<
cS)
oO 100 200
pecan TRANSECT WIDTH (m| OCEAN
(lee| (windward)

Figure 46. Brothers Islet: east-west cross-section through
center of islet. Data includes floristic composition, relative
abundance of plant species, degree of species overlap and canopy
heights. Vertical height is exaggerated. Note the central
monotypic stand of Pisonia forest. This islet's forests were
totally felled in 1920.

141

RUBBLE

GB here mat N
(eens) TOURNEFORTIA
=] PISONIA
L-] cocos
lagoon Bw NODDY ROCK

rm

eo NORTH ARUNDEL ISLET 7

Acropora-Tridacna
Reet Di

. SS
Sa mS See

MP ~

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TS
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jz 0 500

meters

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£
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Figure 47. Vegetation and physiography of Windward
Islets nos. 10 through 12: Noddy Rock, North Arundel,
and Arundel. See text for explanation of the relatively
small amount of Pzsonia cover (Sect. H).

142

Noddy Rock, an eroded limestone plateau of unknown age, is 26 m
wide by 9 m long and 0.5 m above high water. It is windswept and salty,
with waves generally splashing over its eastern edge. During storms it
is completely awash (Anne Falconer, pers. comm.). Only 3 species of
plants (11% of Caroline’s flora) grow here, thinly covering the western
(leeward) third of the island in the following proportion: 75%
Portulaca, 20% Lepturus, and 5% Tournefortia.

13) NORTH ARUNDEL (0.91 ha) Discussed below
14) ARUNDEL (7.34 ha) Figs. 29325°473—Pils 14a

Arundel Islet was named last century in honor of John T. Arundel.
A British trader and guano digger, Arundel was one of the leading
figures in the Pacific phosphate industry, directing guano and coconut
planting operations on Caroline and other islands from 1873 to 1897.
His most valuable contributions, however, were his excellent surveys and
maps of several central Pacific islands, including Caroline (Fig. 4).
The islet immediately to its north, Arundel’s "cap," we named North
Arundel.

Physiography: Arundel is crescentic, with wedge-shaped North Arundel
lying across a short channel immediately to its north. North Arundel is
80 m long x 130 m wide, while Arundel is 375 m long x 210 m wide. They
are composed almost exclusively of coarse coral rubble, flanked on their
inner edges by Acropora-Tridacna reefs. Arundel’s lagoonward "horns"
have evidently added more sand and rubble since 1883 (Figs. 4, 47).

Vegetation: There are 11 plant species (3 trees, 1 shrub, 7 herbs) on
Arundel, 42% of Caroline’s total. There are no introductions. North
Arundel also has 11 (4 trees, 1 shrub, 6 herbs), 42% of Caroline’s
flora, including one introduction, Cocos.

The vegetation on the motus, along with Tridacna to the south,
consists of extensive herb mats, low scrub and small interior forests
(Fig. 32), slightly less lush than the more northerly windward motus.
Their woodlands are primarily Tournefortia, with thin belts of Cordia
and central Pisonia groves (a bilobed pattern on Arundel). Morinda is
unusually common on Arundel, and Achyranthes abundant on North Arundel.
Pisonia occupies only 13% of the land area on Arundel, compared to 46%
on Pig. Their poor soils are a possible legacy of the guano era.

Both motus were heavily planted with Cocos in 1919-20 (69 and 646
palms, respectively). All usable land was cleared (Table 13). Despite
today’s paucity of Cocos, the relatively scant Pisonia, compared to
motus further north, suggests that their plantations were more
successful and maintained more frequently.

Birds: Five species of seabirds bred on Arundel: Red-footed Booby (37
pairs), Great Frigatebird (on territory, September 1988; breeding
confirmed, early 1989 by Anne Falconer), Brown Noddy (11 pairs), Black
Noddy (249 pairs), and White Tern (227 pairs). In May 1990, thousands
of Sooty Terns swarmed both motus.

143

15) TRIDACNA ISLET (9.08 ha) Figs. 29, 48, 49; Pls. 1,
25, 48, 56, 57

The 13th and southernmost motu in the windward chain was named by
the present authors and Boris Sirenko for its outstanding coral reef
densely studded with giant clams (Tridacna maxima).

Physiography: Somewhat crescentic, measuring 446 m long and 250 m wide,
Tridacna is one of the largest motus on Caroline. Its terrain is
heavily littered with coral rubble, having a sandy strip above the beach
crest on the windward edge.

Vegetation: There are 13 plant species (2 trees, 2 shrubs, 9 herbs),
50% of the atoll’s flora. For its size, Tridacna’s vegetation is
surprisingly lacking in tall forests, a legacy of the 910 Cocos palms
planted on 82% of its available land area (Table 13). Vegetation
patterns follow the usual concentric zonation: peripheral herb mats
border a discontinuous belt of Suriana (windward side), while the large
central mass is dominated by scrubby JTournefortia-Morinda woodlands,
which cover 88% of the islet’s area, yet only attain 7 m in height. In
cross-section (Fig. 49), the short woodlands are particularly
noticeable. Compare the present lack of Cordia, paucity of Pisonia, and
richness of herbs, both in species numbers and abundance, with Pig
(Fig. 45) and Brothers (Fig. 46). Although there are no introduced
plants, thick patches of Lepturus also reflect past forest clearing.

Birds: Four species of seabirds were nesting in 1988: Red-footed Booby
(111 pairs), Brown Noddy (11 pairs), Black Noddy (249 pairs), and White
Tern (227 pairs). Tridacna is periodically a major breeding area for
Sooty Terns. Clapp & Sibley (1971la) found 4 main colonies totalling
250,000 birds, and large numbers nested along the windward beach in
August 1989 (Anne Falconer, pers. comm.). Nests were located under
shrubs, or in open areas bordering them, and were evidently preyed upon
by coconut crabs.

16) SOUTH ISLAND (104.41 ha) Figs. 305036, SO;Plisepl-8, 12;
23, 34, 39, 40, 44, 45, 56

History: The history of South Island (called Rimapoto in Young ca.
1922) is essentially the history of Caroline, for most information about
the atoll prior to 1965 is from here. It is the second largest islet,
and the staging area for trips up-lagoon as it lies adjacent to both the
"boat landing" and "blind passage."

South Island was inhabited in prehistory by Tuamotuans, who
planted the first small coconut grove on its northwest point. The first
Europeans to land, in 1606, found coconuts, fish, lobsters, and seabirds
in abundance. They dug for fresh water in vain. Two hundred years
later, in the decade after a cyclone in 1825, pigs, sweet potatoes,
arrowroot, and South Sea chestnut were introduced. However, "the
unfriendly character of the soil, and the number of land crabs that
infest it, gave us but little hope of the experiment succeeding"

144

N RUBBLE

| MB ers mat
[__] TOURNEFORTIA/MORINDA
[- *] SURIANA

lagoon

Acropora -Tridacna

Reefs

ocean

ay
i TRIDACNA ISLET
> Y
SS a ea en ee
| W meters 500
rey
a

Figure 48. Vegetation and physiography of Windward Islet no.

13: Tridacna Islet. The best quality Acropora-Tridaena reefs
extend clear across the lagoon from this motu. See Section H

for explanation of unusual forest cover.

145

Tourneforstia

Tournefortia Morinda Tournefortia Herb mat

Boerhavia repens
Heliotropium anomalum
Laportea ruderalis
Lepturus repens

Morinda Citrifolia
Phymatosorus scolopendria
Pisonia grandis
Portulaca lutea

Tournefortia argentea

CANOPY HGT im!

ppc TRANSECT WIDTH Im! OCEAN

lee} (windward |

Figure 49. Tridacna Islet: east-west cross-section through
lower center of motu. Data includes floristic composition,
relative abundance of plant species, degree of species overlap
and canopy heights. Vertical height is exaggerated. Note the
absence of well-developed interior forests, unusual for a

motu of this size (see Sect. H).

146

RUBBLE
N GHB HERB Mat

([__] TOURNEFORTIA

f= cocos

[_+] DYING COCOS-IPOMOEA
[= *] SuRIANA

{wag PANDANUS

Fa BEACHROCK

r

reef “entrance” —>

“landing”

old settlement yo lagoon

submerged
sand

ocean

blind
passage

:

1s

ig SOUTH ISLAND

£2

ap | tele en i J

a ry Soe 500

Figure 50. South Island: vegetation and physiography.
Note the accepted landing route across its leeward reef
flats.

147

(Bennett 1840). The pigs soon expired. The arrowroot, tenacious and
adapted to island environments, still exists today (unless later
immigrants brought it). Of the others, plus many other later food
plants and ornamentals, no traces exist (Table 1). Tropical heat,
droughts, storms, excessive shade from Cocos, poor germination, poor
soils, terrestrial crabs, and lack of care all undoubtedly contributed
to their demise.

The first recorded settlement on Caroline, and first for the Line
Islands was in 1846, on the northwest point. These settlers, as well as
subsequent ones, eked out a spartan living by raising stock, drying fish
and copra, and digging for guano. Their managers built "proper"
dwellings, so when U.S., British, and French astronomers arrived to
observe the solar eclipse in May 1883, South Island was quite
"civilized," far more than it is today. Three houses and 2 sheds "were
in good repair," and a variety of "anchors, chains, spars, and pieces of
the woodwork of vessels" littered its reefs (Holden & Qualtrough 1884).
Large grassy clearings adjacent to the lagoon accommodated several
European-style houses (Pls. 2-6). The astronomers’ account of South,
illustrated with pen-and-ink drawings (Pls. 2-7), is the only record of
buildings on Caroline, apart from mention of the manager’s house,
reported in 1936 by the "H.M.S." Wellington to be "in excellent
condition and spotlessly clean" (Maude ca. 1938), and a copra shed
(Clapp & Sibley 197la). Arundel took photographs, including the marae
on Nake, which we have not examined.

Today, the houses, sheds, brick piers (constructed in 1883 for
telescopes and observatory frames), signboard, flagpole, marble slab
with inscription "U.S. Eclipse Party, 1883, May 6," and all but one of
the introduced plants have disappeared. In 3 trips we found no traces
of the copra shed, nor have the Falconers, after repeated visits over 2
years. All that remains of the formerly extensive open areas are 2
small palm-shaded clearings, in 1988 used by the U.S. and Soviet
scientists for a base camp and work area. In 1987, the Falconers lived
in one clearing, and in 1990, fishermen expanded the other by burning an
area 35 x 22 m, then erecting a tin shack, cookhouse, and fishtrap, al]
destroyed in a summer 1990 storm.

Our "civilization list" probably covered all that could be seen on
South Island without digging: a 26-foot wrecked sloop (AK 6691 J.),
complete with trail to a "Robinson Crusoe-type" campsite strewn with
remnants of radio and navigational equipment, sail, cans, clothing, etc.
(southeast coast); assorted flotsam and jetsam (whisky bottles, Japanese
fishballs, plastic debris, etc.); a large rubber ship fender; a bench
mark from the 1985 RNZAF survey team; a recently renovated concrete
cistern (by the landing); and an old wooden canoe lying on its side just
like de Quiros found in 1606.

7xrundel’s memorabilia (photos, letters, diaries, a microfilm, etc.) are
in the Rare Book Collection, National Library and Pacific Manuscripts Bureau,
Research School of Pacific Studies, Australian National University, Canberra,
A.C.T., Australia.

148

We assume that all the Polynesians, ancient and recent
(Tuamotuans, Tahitians, Niueans as far as is known), lived in native
thatched huts similar to the ones on Ana-Ana today.- Fashioned from
coconut palms and pandanus trees, they disappear quickly when abandoned.
The largest number of inhabitants recorded for Caroline (probably all on
South Island) was "two managers and 52 laborers" in 1873 (The Commercial
Advertiser 1873).

The history of South Island’s coconut plantations from 1885-1901,
and from 1916-1929, is discussed under Coconut Woodlands (Sect. G).

Physiography: South Island forms the base of the thinly crescentic
isosceles triangle whose limits define the atoll. Its own shape is that
of an irregular parallelogram 858 m wide x 1,254 m long at its longest
points (Fig. 50). The north coast, a curved bay, forms the lower
boundary of the lagoon. This palm-lined shore, along with the adjacent
northwest peninsula, has been the most trodden by man, one of the most
picturesque spots on the atoll (Pls. 23, 39).

The reef flats surrounding the outer 3 sides of South are the
widest on the atoll, averaging 231 m, 578 m, and 363 m on the east,
south and west, respectively. The windward and leeward reefs
immediately to its north are 530 m wide. To leeward is the small boat
"landing," on a conglomerate platform (Pl. 20), and to windward, the
"blind passage," a non-functional hoa (Pl. 72).

Vegetation: There are 23 plant species (7 trees, 3 shrubs, 13 herbs),
89% of the atoll’s flora. Cocos dominates South Island, occupying 77%
of its area. The superannuated closed-canopy plantations (21 m tall)
that border its coastlines give no indication of the vast overgrown,
dying groves that occupy 80 ha, two-thirds of its interior (Fig. 50;
Pls. 8, 34). Here, 3 species of herbs (Boerhavia repens, Portulaca
lutea, Phymatosorus scolopendria) have proliferated unnaturally to form
dense carpets, and the vine (Ipomoea macrantha) climbs in tangled,
strangling masses to the tops of the highest palms.

The natural communities that prevail on other motus are only minor
components on South (Fig. 50): herb mats (13% of the total area),
Coastal Scrub with Suriana (1%), and Tournefortia Scrub (4%).
Conspicuously absent are prime scrublands and forests of Tournefortia,
Pisonia, and Cordia, which undoubtedly once swept in a lush expanse from
shore to shore, stratified and zoned as on other motus, and which were
"80 to 100 feet high" before extensive cleaning began (Arundel 1875).
Canopy heights of the plantations are uniform (21 m), and the outer
fringe of indigenous scrub (Tournefortia, Cordia, Suriana) and herbs
(Heliotropium, Portulaca) occupy a small proportion of the island’s
width (Figs. 36, 51). Note the abrupt transition of canopy heights as
they drop to the level of coastal scrub on both sides of the plantation.
Pandanus, too, is less extensive than formerly: Bennett (1840) called
it "somewhat numerous." During our survey, we observed only one small
Pandanus grove and a few scattered trees. Bennett also noted that the
island was "covered with verdure," with "trees attaining the height of
twenty feet." However, 9 or 10 years previous to Bennett’s visit a

149

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violent storm had whipped over the atoll. By the early 1870s, trees
were approximately 30 m high again (Arundel 1875). Remnant Journefortia
and Pisonia are illustrated in drawings of South Island’s lagoon shore
in- 1883, (Pls. 45.05).

Apart from the coastal buffer zone, little native forest remains.
Other sizable trees (Pisonia, Cordia), up to 17 m tall, are rare, but
Morinda, tolerant to both sun and shade, is still quite common. Though
we have not been able to trace any records to Caroline, it is possible
that shiploads of Cordia logs were exported to San Francisco on guano
ships, as was the case on Flint, worked simultaneously by Arundel’s
company (Young ca. 1922).

A final noteworthy aspect of South Island is that, despite its
history of sporadic occupation and extensive forest felling for coconut
plantations, only one exotic (a tiny patch of Phyllanthus amarus) and no
vegetable or garden ornamentals (excluding Polynesian introductions)
have survived.~ The 19th century gardens, once drenched in sunshine,
have long been buried beneath the deep shade of palm groves (compare
Pls. 2, 3 and 23). In addition, rare hurricanes, periodic storms,
droughts, irregular rainfall, nutrient-poor soils, rats, land crabs, and
the harsh salty environment must have contributed to the eradication of
alien species except traditional native food and medicinal plants, which
are specifically adapted for atoll environments. Studies on other
atolls, even those near high islands (Stoddart & Gibbs 1975, Stoddart &
Fosberg 1972), have demonstrated also that many exotics do not survive,
despite the proximity to source areas containing garden ornamentals and
weed plants. On our last 2 visits (March, May 1990), however, we
discovered a weed not previously reported (Kyllinga brevifolia) near the
Phyllanthus, a consequence of recent clearing around the cistern. This
area was an extension of the Falconers’ vegetable garden on Motu Ana-
Ana. Kyllinga is listed as a temporary species (Table 1).

Birds: Only 2 species of birds bred on South in September 1988, a
reflection of its paucity of natural habitats: Brown Noddy (163 pairs)
and White Tern (381 pairs). Bristle-thighed Curlews are very common,
gathering in small flocks on the rubbly shores (Pt. II). They also
forage in the open Ipomoea-Cocos forest, perching on dead coconut stumps
6-10 m high, then flying down to feed in the thick herb mats.

Terrestrial Crabs: Caroline’s highest population of coconut crabs,
having many huge individuals decades old, occupies the open Cocos
forests (Pls. 21, 59) - A crude minimum estimate for South Island is 500
mature individuals.~ We also found a fist-sized blue hermit crab within

BWe are unsure of the status of Hibiscus tiliaceus, Thespesia populnea or
Ximenia americana. Although indigenous to the area, a recent analysis of the
vegetation of Flint Island (Kepler, in prep.) indicates that these species
were most likely introduced to both islands in the late 19th century.

since March 1990, these have become much reduced due to killing and
preserving in formalin for curios.

151

a Turbo argyrostomus shell, possibly Coenobita brevimanus (Yaldwyn &
Wodzicki 1979; E. Reese, pers. comm.). As elsewhere on the atoll, land
crabs such as the reddish-purple Cardisoma sp. and scarlet hermit crabs,
Coenobita perlatus (in T. argyrostomus shells) were abundant (Pi38)2
Geograpsus sp., closer to the shore, was less common.

Rats: Polynesian rats were abundant, constantly afoot in broad

poupiaht, and at night flashlight beams often revealed a half dozen at a
ime.

SOUTH NAKE ISLETS Rid... 52

This chain of 7 islets extends 1,500 m south from Nake on the west
side. They range in size from 0.64 ha (Kota) to 7.36 ha (Pandanus).
All are well-wooded and support every natural plant community.
Proceeding south, the overall plant cover thins somewhat, but not to the
dryness and openness of the Central Leeward Islets. The herb mats are
more extensive than on the windward islets, especially to seaward.
Aboriginal introductions (Cocos, Pandanus) are sparse. We have found no
historical records indicating human disturbance to these islets, thus
their vegetation, with the possible exception of Pandanus Islet, is
evidently natural. The 2 scrawny Cocos are probably drift-derived.

On the Solar Eclipse Party’s map of Caroline (Fig. 5), only the
top 2 islets of this group are drawn. The South Nake Islets constitute
the only cluster of motus that show appreciable differences between
Arundel’s chart (Fig. 4) and the 1985 aerial photos: most were shown as
smaller, and with slightly different shapes, by Arundel. The interior
vegetation on these motus includes mature forests of Journefortia,
Pisonia, and Pandanus, so it is unlikely that these differences reflect
changes to the center of the motus. However, since the islets now
appear larger, accretions of coral rubble and sand that may have
occurred in the past 105 years, and are now barren or covered only with
herb mats, could account for most of the differences (see Coral Islet
discussion).

Although we have no actual records of Sooty Tern colonies on this
chain of islets, in May 1990 AKK observed pre-breeding swirls over Lone
Palm, Kota, and Mouakena (Pt. II, Fig. 11).

17) PANDANUS ISLET (7.36 ha) Figs. 29, 52; Pl. 58

This motu was named by the present authors for its coastal
Pandanus grove, probably a drift-derived offshoot from a parent colony
on Nake.

Physiography: Pandanus Islet, first in the chain, is irregularly oval,
400 m long and 258 m across. It is nearly twice the size shown on
Arundel’s map (ca. 3.4 ha). It occupies a sheltered spot at the apex of
the lagoon. Sand, actively filling in the adjacent lagoon, is an
important substrate component on Pandanus, extending one-third of the

152

RUBBLE : i SS
HERB MAT) 9 ;
TOURNEFORTIA
CORDIA \
PISONIA
PANDANUS \

cocos )

Re
=

PANDANUS ISLET

DANGER ISLET

BOOBY ISLET

CORAL ISLET

ocean

LONE PALM ISLET ©

© MOTU KOTA
we
Ne}

MOTU MOUAKENA

s SOUTH NAKE ISLETS
{>
a N\ a ee Se a
= 0 meters 500

Figure 52. Vegetation and physiography of the 7 South
Nake Islets: Pandanus, Danger, Booby, Coral, and Lone
Palm Islets, Motu Kota ("Red-footed Booby Islet"), and
Motu Mouakena ("Masked Booby Islet").

153

way across the islet. Although tidal reef flats are absent on the
lagoon edge, they average 75 m wide on the seaward side, producing a
fairly high proportion of rubble compared to the total land surface
(32%).

Vegetation: Plant species total 10 (3 trees, 1 shrub, 6 herbs), 39% of
Caroline’s flora. Cocos is absent, despite the motu’s close proximity
to Nake. Pandanus Islet has 4 basic vegetation zones: natural herb
mats, Tournefortia Scrub (with Pandanus), Tournefortia-Pisonia Forest,
and pure Pisonia. Woodlands cover 62% of its area. The widest pioneer
mats (13 m) of any leeward motu occupy its east edge and though sparsely
vegetated (20% Heliotropium, 5% Lepturus, 5% Portulaca) reflect active
growth toward the lagoon. Proceeding west across the island,
Tournefortia scrub (2 m high), with pockets of pure Pandanus (10 m
high), merges into Tournefortia-Pisonia Forest (to 14 m high), whose
bimodal distribution suggests that the islet was once divided. The
seaward coast supports open Journefortia (5 m high), beneath which herbs
eventually thin out onto the extensive reef flats.

Birds: Five species of seabirds breed: Masked Booby (2 pairs), Red-
footed Booby (32 pairs), Great Frigatebird (26 pairs), Brown Noddy (26
pairs), and White Tern (52 pairs).

Comments: Skinks and rats were observed, along with the ubiquitous
Coenobita and Cardisoma land crabs.

18) DANGER ISLET (2.71 ha) Pigs G5 be HS. Be)

We named Danger Islet to commemorate the deep, shark-infested
channel to its north, a barrier that aborted our first attempt to survey
the South Nake Islets.

Physiography: Danger, of rounded shape, is approximately 150 m long and
215 m wide. It is composed almost entirely of coral rubble; interior
humus is scant. Its reef-channel flats are 21 m (north) and 14 m
(south) wide. The east and west beaches, narrow and wide respectively,
are typical of all the leeward motus.

Vegetation: Danger has 10 plant species (3 trees, 1 shrub, 6 herbs),
39% of the total flora. There are no introductions. The vegetation is
concentrically zoned: herb mats, Tournefortia scrub and forest, central
Pisonia, and Cordia in the southwest. The herb mats are wide, extending
22 m and 15 m on the north and south shores, respectively.

Birds: Four species of nesting seabirds were present in 1988: Red-
footed Booby (139 pairs), Great Frigatebird (26 pairs), Brown Noddy (26
pairs), and White Tern (52 pairs).

154
19) BOOBY ISLET (0.84 ha) Figs. 295 5257Pi-voe

We named this motu, third in the chain, for its 2 species of
boobies, the common Red-footed and rarer Masked Booby.

Physiography: Booby, shaped like a teardrop, is 70 m long and 125 m
wide. Its coral rubble flats extend 10 m and 30 m on the north and
south sides, respectively.

Vegetation: Despite its small size, the most notable feature of Booby
is its Pisonia forest, 20 m tall and undoubtedly virgin. It occupies
the exact center of the islet in a circle about 40 m in diameter.
Surrounding this is Tournefortia scrub (to 8 m tall), thinning out to
peripheral bands of coral rubble. Although less than one hectare in
size, Booby Islet’s woodlands occupy two-thirds of this area. Booby
Islet has 9 species of plants (2 trees, 1 shrub, 6 herbs), 35% of
Caroline’s flora, and no introductions.

Birds: Five species of seabirds breed: Masked Booby (7 pairs), Red-
footed Booby (52 pairs), Brown Noddy (2 pairs), Black Noddy (1 pair),
and White Tern (6 pairs).

20) CORAL ISLET (1.70 ha) Figs. 29, 52; Pl. 60

Fourth from the north, Coral Islet was named for its reef-derived
coralline substrate.

Physiography: Arrowhead-shaped, Coral is approximately 130 m long by
200 m wide, more than 3 times the size mapped by Arundel (Fig. 4). Most
of its area is barely higher than the surrounding inter-islet channels.
The shallow reef flats between Coral and its 2 southern motus are only
several centimeters deep at low tide; all 3 may be destined to unite.
Unless closely inspected, they appear to have already merged, a fact
which, together with Bryan’s incorrect map (Fig. 6), helps account for
the widely differing number of motus attributed to Caroline.

Vegetation: There are 9 species of plants (2 trees, 1 shrub, 6 herbs),
35% of Caroline’s flora, and no introductions are present. Plant
communities comprise a small Pisonia forest (0.13 ha), which is
surrounded by the predominant Tournefortia, in turn fringed with a
narrow band of native herbs. "Soils" are extremely coarse.

Birds: Five species of seabirds bred in 1988: Masked Booby (1 pair),
Red-footed Booby (28 pairs), Great Frigatebird (2 pairs), Brown Noddy (6
pairs), and White Tern (15 pairs).

21) LONE PALM ISLET (1.99 ha) Figs. 29, 52; Pls. 60=e0

We named Lone Palm, fifth in the chain, for its single coconut
palm which towers, flag-like, above a dense mound of Journefortia.

155

Physiography: Similar to Kota (to its south), Lone Palm is sausage-
shaped, 97 m long and 240 m wide, and 4 times the size mapped by
Arundel. Although composed almost entirely of coral rubble, some sand
borders the lagoon. Following a pattern prevalent on all leeward motus,
its lagoon beach is 2 m wide, while the seaward beach is 17 meters.

Vegetation: Eleven species of plants are present (3 trees, 1 shrub, 7
herbs), 42% of Caroline’s flora. Plant communities are simple: a wide
band of herb mats and open Journefortia flanks an oval of Tournefortia
forest (to 10 m tall). A line of Pisoni7a trees, with a lone Cocos
surmounting the scrub, identifies this islet from lagoon or ocean.

Birds: Three species of seabirds bred in 1988: Masked Booby (2 pairs),
Red-footed Booby (48 pairs), and White Tern (9 pairs). In May 1990, we
saw a large pre-breeding swarm of Sooty Terns.

22) MOTU KOTA "Red-footed Booby Islet" (0.64 ha) FAQS 428 0152s
Pls. 60

We named this motu for its high density of Red-footed Boobies
(kota in Gilbertese).

Physiography: Sixth in line south of Nake, sausage-shaped Motu Kota is
50 m long and 175 m wide. At low tide it is almost connected to Motu
Mouakena. Both surveys indicate that coral rubble, the islet’s
predominant substrate, had further accumulated on its south side since
the 1985 aerial photos, and also since 1988.

Vegetation: Though barely wooded, Kota has 11 species of plants (3
trees, 1 shrub, 7 herbs), 42% of Caroline’s flora. One introduced
species is present, one tattered Cocos. Plant communities include:
peripheral herb mats and a central Tournefortia scrub (to 10 m tall),
with a few Pisonias.

Birds: Three species of seabirds bred in 1988: Brown Booby (1 pair),
Red-footed Booby (12 pairs), and White Tern (3 pairs). In May 1990, a
single Masked Booby was on territory, and Sooty Terns swirled overhead.

23) MOTU MOUAKENA "Masked Booby Islet" (1.00 ha) Figs. ¢29bn 525
Pilisisy 15,9162

This islet was named for its nesting Masked Boobies, a relatively
uncommon seabird on Caroline.

Physiography: Somewhat U-shaped, Motu Mouakena is seventh, and
southernmost, in the South Nake chain of islets. Both sides of the "U"
were, in the recent past, separate islets. By joining on the west, a
narrow, V-shaped inlet was created on the lagoon side. Motu Mouakena,
100 m long and 160 m wide, is extremely rubbly and infertile; much
rubble was reorganized during the February 1990 storm. Seventeen meters
to its south lies a newly emerging shoal of sand and gravel (Pl. 15),

156

perhaps destined to be Caroline’s fortieth motu. Since the above storm,
rubble has further accumulated on this shoal, its adjacent reef flats,
and in the channel separating it from Mouakena. It already supports one
Tournefortia shrub, 2 dozen Heliotropium plants, and scant Lepturus and
Portulaca.

Vegetation: Mouakena has 8 species (1 tree, 1 shrub, 6 herbs), 31% of
Caroline’s flora, with no introductions. It is thinly vegetated with

open Journefortia scrub (to 9 m tall, 26% cover), a few small Pisonia,
and sparse herb mats.

Birds: This motu supports less vegetation and fewer birds than its
overall area implies, since 38% of the land consists of unshaded, coarse
coral rubble (Fig. 29b). Although unproductive botanically, this
provides ideal nesting grounds for Masked Boobies, one of its 2 species
of 1988 breeding seabirds: Masked Booby (3 pairs) and Red-footed Booby
(8 pairs). In May 1990, we also saw one Great Frigatebird nest with
eggs and a swirl of Sooty Terns.

CENTRAL LEEWARD ISLETS

This chain of 11 motus occupies the central west side of Caroline.
All are separated by channels, wadable only at low tide but prowled by
belligerent sharks. Approximately 1,600 m south of Motu Mouakena lies a
sandy shoal (0.5 m high, 7 m wide, 4 m long), close to the lagoon edge
of the reef flats and connected only by a thin thread of rubble to Motu
Mannikiba to its south.

The islets range in size from Mannikiba (28.50 ha), the most
northerly, to Fishball (0.46 ha), the most southerly. All support good
seabird populations. Although most are well-wooded, they are
nonetheless the least lush motus on Caroline. Historical records are
meagre: much of Mannikiba’s forest was felled to make room for a Cocos
seedling "nursery" (Young ca. 1922). The bulk of "40 trees on other
islets," in Young’s plantation totals, were most likely from Shark and
Emerald. The rest of this group is evidently pristine; Bird Islet is
particularly notable.

In common with all western motus on Caroline, the lagoonside
beaches are narrow and leeward reef flats wide. The latter exhibit a
greater variety of substrata than the former, including older raised
reefs and beachrock. Periodically, thousands of nesting Sooty Terns
occupy their open spaces (Clapp & Sibley 1971a; AKK, pers. obs.;

Anne Falconer, pers. comm.).

24) MOTU MANNIKIBA "Seabird Islet" (21.49 ha) Figs.) 2957 156;
Pls. 63-65

We named this motu for its teeming seabirds, mannikiba in
Gilbertese.

RUBBLE

HERB MAT
TOURNEFORTIA
PISONIA
COCOS
BEACHROCK

lagoon

MOTU MANNIKIBA

[oxe)
0 meters 2

Figure 53. Vegetation and physiography of
Central Leeward Islet no. 1: Motu Mannikiba
("Seabird Islet").

157

158

Physiography: Largest and most northerly of the Central Leeward Islets,
Mannikiba is somewhat rectangular with rounded corners. Its reef flats,
supporting an incipient islet, stretch 2.0 km north to the South Nake
Islets.

Mannikiba’s maximum dimensions are 700 m long and 375 m wide. On
the lagoon side, the scrub skirts high water, but when the tide drops, a
strip of blinding white sandy coral lines the lagoon. To seaward,
upraised reef, beachrock, and successive layers of rubble stretch in a
wide swath (40 m) toward the outer reef, 130 m distant. The primary
inland substratum is coral rubble with hardpan in the northeast.

Vegetation: Mannikiba, the fourth largest motu, harbors 13 plant
species: (4 trees, 2 shrubs, 7 herbs), 50% of Caroline’s flora. The
only introduction is Cocos, occupying 0.1% of the land area. The
single, sterile Ximenia americana is indigenous, but may have grown from
a seed derived from the other patch of this species on South.

Mannikiba’s vegetation, denser toward the north end, is clearly
zoned: herb mats, Tournefortia scrub and forest, and scattered Pisonia
groves. The few clumps of peripheral Cocos are probably not drift-
derived, but the remnants of 6,000 "seed sets" brought from Flint Island
in June 1920. These were stored on Mannikiba and "used to replant
misses on other islets" (Young ca. 1922).

Pisonia, though present, occupies only 5% of the land area, a
small percentage for such a large islet. This suggests that large areas
of the interior forests were felled to accommodate the coconut "sets."
This is also confirmed by the presence of several old cut stumps
(Cordia?) in the interior, undoubtedly a legacy of S. R. Maxwell and
Co., Ltd., who erected huts around 1920 (Young ca. 1922). Although
nothing more is known of Mannikiba’s history, collection of guano from
its numerous seabirds may account for further past disturbance.

Transect 1 (north-central sector, Pl. 64) passed through the heart
of a fine interior forest, while Transect 2 (south-central sector)
passed through scrub and herb mats which may represent part of the
former Cocos "nursery." Profiles through these 2 cross-island transects
resemble those from Brothers (Fig. 46) and an old interisland channel on
Long (Fig. 40), respectively.

The low, peripheral herb mats (absent from the lagoon side) are
composed of 30% Heliotropium, 20% Boerhavia, 15% Tournefortia, and less
than 1% of Portulaca and Laportea. They are best represented in the
southern sector. The Journefortia forest, 6 m high on both sides, is
thick, having 95% canopy coverage. The Pisonia forests, though
fragmented (12 m high, 100% canopy cover), contain Morinda, Boerhavia,
Achyranthes, Laportea, and Phymatosorus, but none cover more than 10% of
the ground area.

Seabirds: Six species are known to breed: Red-footed Booby (184
pairs), Great Frigatebird (287 pairs), Brown Noddy (161 pairs), Black
Noddy (176 pairs), and White Tern (195 pairs). No Sooty Terns nested on

159

this islet in 1988, but Clapp & Sibley (197la) estimated 2,500 pairs in
1965, and the Falconers reported large colonies on Mannikiba, Blackfin
and Matawa in July - August 1990.

Comments: Coconut crabs live in the Cocos grove. Azure-tailed and
snake-eyed skinks (Cryptoblepharus poecilopleurus), as well as a gecko,
were noted in 1990 (DHE, G. Wragg, pers. obs.).

25) BLACKFIN ISLET (2.62 ha) Figs.29, 54; Pls. 29

We named this motu, second in the Central Leeward chain, for 2
shark attacks (near misses) within its northern channel.

Physiography: Blackfin, shaped like conjoined ovals, is 140 m long and
190 m across. Coral rubble covers 30% of its surface; all beaches and
upper reef flats are of variable widths, due in part to the fact that it
has recently incorporated a smaller, circular motu into its northern
confines.

Vegetation: Blackfin Islet has 9 species of plants (3 trees, 1 shrub, 5
herbs), 35% of Caroline’s flora. The only introduction, Cocos, is rare.
Four plant communities were present. Herb mats are well represented,
especially around the newly incorporated islet. The Tournefortia scrub,
21 m wide in the east, is short (to 2 m), but approaches the stature of
a forest (to 6 m) in the west. The central forests of Cordia and
Pisonia (0.41 ha) are 9 m high.

Birds: Three species of seabirds bred in 1988: Great Frigatebird (4
pairs), Brown Noddy (37 pairs), and White Tern (11 pairs). In May 1990,
one Red-footed Booby sat tight on a nest, while 2 months later large
numbers of Sooty Terns began laying.

26) MOTU MATAWA "White Tern Islet" (1.71 ha) Figs. 29, 54;
Pies) (Pt 11)

On arriving at this motu, the authors were greeted by 15 White
Terns, matawa in Gilbertese.

Physiography: Of oval shape, Motu Matawa is third from the north in the
Central Leeward chain. It is 105 m long and 190 m wide. The entire
motu is composed of coral rubble of varying grades, whose unvegetated
portion comprises one-fourth or more of the land area. Its lagoon beach
is 2.5 m wide, while the seaward beach (sparsely vegetated) is 6 m wide.

Vegetation: Matawa has 10 species of plants (4 trees, 2 shrubs, 4
herbs), 39% of Caroline’s flora. There are no introductions.
Vegetation is less lush and more open as one progresses south on the
leeward side. Journefortia (to 7 m) covers half the islet, surrounding
an east-central Pisonia-Cordia forest (to 8 m).

160

N
RUBBLE
GMB vers mat
(__] TourNEFORTIA
PISONIA
fees PANDANUS-TOURNEFORTIA
ESBS | CORDIA
[===] cocos
BLACKFIN >
ISLET 7
MOTU
MATAWA
ocean }
lagoon

EMERALD ISLE

U g ° meters 200
e
+
i?

Figure 54. Vegetation and physiography of the
Central Leeward Islets nos. 2 through 4: Blackfin
Islet, Motu Matawa ("Fairy Tern Islet"), and Emerald

Isle.

161

Birds: In 1988 four species of seabirds bred: Red-footed Booby (5
pairs), Great Frigatebird (1 pair), Brown Noddy (3 pairs), and White
Tern (13 pairs). Most conspicuous were White Terns, with 9 pairs
breeding on the 30-m-wide transect swath. One dark morph Reef Heron
fished in the shallows. In summer 1990, Sooty Terns also bred.

27) EMERALD ISLE (8.34 ha) Figs. 295054; Plissiv24, 66, 67

Fifth down the chain, we named Emerald for the richly colored,
translucent lagoon waters that fringe its shorelines.

Physiography: Crescentic Emerald, 330 m long and 240 m wide has
lagoonside reefs, patch reefs, and coral knolls irregularly patterned
with sandy channels.

Vegetation: Emerald Isle has 12 species of plants (5 trees, 1 shrub, 6
herbs), 46% of Caroline’s flora. The only introduction is Cocos. Four
plant communities, with a fairly high species diversity, are present:
the herb mats, covering one-fourth of its land area, are composed almost
exclusively of Heliotropium (35% cover) with scattered low Journefortia
(30% cover). The Tournefortia attains a maximum height of 8 m and, for
variety, is mixed about equally with Pandanus over most of its seaward
width (144 m).

The interior forest (to 11 m tall) is also mixed, with Pandanus,
Tournefortia, Pisonia, and a little Cordia (Pl. 66). This 3.20 ha mixed
forest, as on Shark, suggests that Pandanus may be both native and
Polynesian-introduced, although we do not have specimens to verify this.
Cocos is present as 2 small groves, complete with coconut crab sign
(mounds of shredded fibers, Pl. 53), beside the east and midwest shores.
We have been unable to trace the history of Emerald’s forests; the Cocos
and fragmented Pisonia suggest past disturbance.

Birds: Six species of breeding seabirds were present: Red-tailed
Tropicbird (1 pair), Red-footed Booby (3 pairs), Great Frigatebird (230
pairs), Brown Noddy (7 pairs), Black Noddy (150 pairs), and White Tern
(83 pairs).

Although we did not locate any Red-tailed Tropicbird nests, 2
adults circled steadily overhead. Two Reef Herons (1 dark morph, 1
light) foraged in the inshore reef shallows.

28) SHARK ISLET (7.98 ha) FigQS-129,.95s3m PIS.) 26, 68

We named this islet to commemorate a particularly pugnacious shark
charged shoreward and leaped to the beach toward our feet.

Physiography: Stoutly crescentic, Shark Islet is 280 m long and 310 m
wide in the center. The sandy lagoon beach and rubbly seaward beach are
each 3 m wide. The former is Caroline’s prime stretch of sand, overlain
by numerous pink granules, possibly due to Foraminifera tests, as in

162

fCOS} Russle
eal HERB MAT
N [__] TOURNEFORTIA
CORDIA
f=] cocos
(sas] PANDANUS

PISONIA

blew 4
a o.
<i

B7 ix
Cc
~

SHARK ISLET

iP
SCARLET CRAB _- —
ISLET

RG

~

L

lagoon

% MOTU NAUTONGA

AZURE ISLE

ocean

REEF-FLAT ISLET ye

ay

i

BIRD ISLET

= Sa

} FISHBALL ISLET ~ eS
Ld? A
{fe ;
vs CENTRAL LEEWARD ISLETS 5-11
M (ee a es eee
S 0 meters 400

Figure 55. Vegetation and physiography of the
Central Leeward Islets nos. 5 through 11: Shark
and Scarlet Crab Islets, Motu Nautonga ("Sea
Cucumber Islet"), Azure Isle, Reef-flat, Bird and
Fishball Islets.

163

common in the Tuamotus (Stoddart & Steers 1977). Beyond high water the
seaward reef flats extend for 280 m.

Vegetation: There are 12 species of plants (5 trees, 1 shrub, 6 herbs),
46% of the atoll’s flora. One introduction, Cocos, forms 3 clumps along
the lagoon beach (1% of the islet’s area). Shark’s rings of vegetation
approximate the islet’s outline. Herb mats dot the fine sand
lagoonward, while to seaward they grow in coarse rubble. The
Tournefortia (to 7 m tall) eventually gives way to a 12-m-high Pisonia
forest studded with Cordia and Pandanus. Centrally this mixed forest is
unnaturally open, suggesting past disturbance.

Birds: Four species of seabirds bred in 1988: Great Frigatebird (118
pairs), Brown Noddy (37 pairs), Black Noddy (125 pairs), and White Tern
(44 pairs). Red-footed Boobies were nesting in 1990. The notable
colonies of Great Frigatebirds and Black Noddies are due in part to the
extensive Pisonia forest, covering one-half of the islet.

29) SCARLET CRAB ISLET (0.46 ha) FAS. 12858 05

This motu was named by the authors in honor of Coenobita perlatus,
the scarlet, fist-sized hermit crab that is abundant both here and on
the entire atoll.

Physiography: Scarlet Crab, sixth in the chain and only 40 m long by
125 m wide, is a young oval motu. It skirts the southern shore of
Shark, from which it is separated by a channel 16 m wide. Because its
eastern end points into the lagoon, there is no true lagoon beach.
Together with the next 3 islets, Scarlet Crab’s seaward reef flats
(480 m) are the most extensive on Caroline’s lee side.

Vegetation: Vegetative cover is slight: less than 1% area coverage of
Heliotropium and Laportea, interspersed with 10 small Journefortia (to
1.5 m). Its species count is 6 (1 shrub, 5 herbs), 23% of Caroline’s
flora. There are no introductions.

Birds: Although during storms this motu is undoubtedly awash, 2 species
of seabirds were breeding in 1988: Brown Noddy (1 pair, on ground) and
White Tern (2 pairs, in low scrub).

30) MOTU NAUTONGA "Sea Cucumber Islet" (0.34 ha) Fatgis' 3 *28)2 255

We named this motu for the Gilbertese word for the black sea
cucumbers or "beche-de-mer" (Ludwigothuria sp.) that are strewn
ubiquitously within the lagoon shallows (Pl. 10).

Physiography: Semicircular in shape, Nautonga is seventh in the Central
Leeward chain, measuring 70 m long and 80 m wide. Situated close to the
lagoon, it is one of 3 small motus that barely protrude above the reef
flats. Nautonga’s perimeter beaches are all narrow (2 m), while its
seaward reef flats are wide (495 m).

164

Vegetation: There are 9 indigenous species (3 trees, 1 shrub, 5 herbs),
35% of the atoll’s flora. Though small, Nautonga’s vegetation is
concentrically zoned, comprising herb mats (10-14 m wide) and a central
forest of Tournefortia and Pisonia (84 m wide), to 10 m tall.

Birds: Five species of seabirds bred in 1988: Red-footed Booby (11
pairs), Great Frigatebird (2 pairs), Brown Noddy (7 pairs), Black Noddy
(32 pairs), and White Tern (10 pairs). Lesser Frigatebirds appeared to
be preparing to nest in May 1990. One pair of Blue-gray Noddies, flying
south, was seen in May 1990.

31) AZURE ISLE (0.20 ha) Figs.::28,1553 "Paden

We named this small, wedge-shaped motu for the striking colors of
its nearby lagoon.

Physiography: Eighth from the north, this small, elongated triangle of
land is 30 m long and 66 m wide. Its seaward reef flats are wide
(512 m), while the adjacent channels are narrow and shallow.

Vegetation: Azure has only 7 species (1 tree, 1 shrub, 5 herbs), 27% of
Caroline’s flora. A young motu, Azure is a superb example of an early
stage of biological succession. Its plant cover consists of a single
mound of Tournefortia scrub crowned by a single Pisonia tree (6 m tall),
growing only one meter above sea level. Only 45% of the motu is
vegetated; the rest, primarily to seaward, is coarse rubble. Azure Isle
illustrates the minimum width of vegetation (38 m) in which Pisonia
develops on Caroline.

Birds: This motu illustrates the speed at which seabirds will utilize
newly available habitats. Within its dozen or so Tournefortia shrubs
(to 4 m tall), 3 species of seabirds nest: Red-footed Booby (7 pairs),
Great Frigatebird (2 pairs), and White Tern (2 pairs). A pair of Blue-
gray Noddies were seen in May 1990.

32) REEF-FLAT ISLET (0.09 ha) Figs. 27/2as

We named this young motu for its primary characteristic: reef
flats. Ninth in the Central Leeward chain, this curved strip of coarse
rubble lies parallel to the channels that surround and spawned it. It
measures about 20 m long and 60 m wide. Three species of plants (1
shrub, 2 herbs), 12% of Caroline’s flora, cover less than one-fourth of
its area and are distributed so sparsely that not one bird was present.

33) BIRD ISLET (4.05 ha) Figs. 29, 55

This is one of the motus named on Arundel’s chart (Fig. 4),
probably because of numerous Black Noddies and/or Sooty Terns.

165

Physiography: Bird is ovoid, measuring 230 m long by 200 m wide. It
sits close to the inner edge of the lagoon reef, whereas 400 m of
seaward reef flats stretch westward.

Vegetation: There are 12 species of plants (4 trees, 2 shrubs, 6
herbs), 46% of Caroline’s flora. A small Cocos grove is the only
introduction. It is well-wooded, with very narrow herb mats (6% of
total area). Journefortia (to 8 m) and Pisonia (to 14 m) each cover 42%
of its surface; the rest is rubble. The Pisonia forest is of good
quality (90-95% canopy cover), having scattered Morinda, Boerhavia, and
Achyranthes as an understory. One large clump of Suriana (14 x 14 m,
2.5 m high) grows centrally (A. Garnett, pers. comm.). Bird Islet shows
very few signs of past disturbance, having prime plant communities, rich
in breeding seabirds.

Birds: Five species of seabirds nested in 1988: Red-footed Booby (29
pairs), Great Frigatebird (6 pairs), Brown Noddy (42 pairs), Black Noddy
(329 pairs), and White Tern (48 pairs). In June 1990, many thousands of
Sooty Terns laid on Bird and adjacent Fishball.

34) FISHBALL ISLET (0.57 ha) Figs. 28, 55, 56

Eleventh and southernmost in the Central Leeward chain, we named
Fishball after finding a large glass fishing float with a broken bottom,
decorously placed in the islet’s center amidst coral slabs.

Physiography: Paramecium-shaped, Fishball lies close to the lagoon and
is separated from Bird by a shallow, rubble strewn channel 100 m wide.
The motu is 45 m long by 144 m wide, with seaward reef flats 595 m in
extent. South of the islet, the reef flats--wadable at very low tide--
stretch 1.4 km to the Southern Leeward Islets.

Vegetation: The number of plant species is 8 (1 seedling "tree," 1
shrub, 6 herbs), 31% of Caroline’s flora. Figure 56 depicts an east-
west cross-section of Fishball, showing a vertical profile and the
relative abundance and distribution of each species. Fishbal]
exemplifies a young motu. All plants are low and halophytic; most are
herbs. The motu is half-covered with a sparse herb mat of Heliotropium
(10% cover), with scattered Laportea, Lepturus, and Portulaca (each <1%
cover). Small Tournefortia shrubs (to 2 m tall) are scattered in the
central sector, while a tiny drift seedling of Morinda, 7 cm high,
struggled to gain a foothold in the exposed, salty rubble.

This motu is a fine example of the initial stages of islet
formation and colonization, demonstrating that sea-dispersed, halophytic
herbs first appear later becoming shaded out by Journefortia, enabling a
greater plant species diversity to establish. It is very unlikely that
a ground water lens is present.

Birds: Two species of seabirds bred in 1988: Red-tailed Tropicbird (3
pairs) and Brown Noddy (5 pairs). In May 1990, many thousands of Sooty
Terns covered the ground and swirled in the air, day and night. We

166

Rubble

Herb

mat
Tournefortia
Herb

mat

rubble

Achyranthes canescens
Boerhavia repens
Heliotropium anomalum
Laportea ruderalis
Lepturus repens
Morinda citrifolia
Portulaca lutea

Tournefortia argentea

0 50 100

TRANSECT WIDTHi(m),, ooo,
lleel [windward]

Figure 56. Fishball Islet (no. 11, Central Leewards):
east-west cross-section through the center of this
young motu, which exhibits early stages of geological
and biological evolution. Data includes floristic
composition, relative abundance of plant species,
degree of species overlap and canopy heights. Vertical
height is exaggerated.

167

found no eggs, but laying occurred in June here and on adjacent Bird
Islet (Anne Falconer, pers. comm.).

SOUTHERN LEEWARD ISLETS Pils: 69

This chain of 5 small motus lies along the southwestern edge of
the lagoon. All are built upon piles of rubble about 3 m high, oriented
east-west, and separated by shallow, narrow channels. From 1.51 to
3.67 ha in size, their topography, vegetation, and breeding seabirds are
similar. Although situated on the leeward side of the atoll, the
Southern Leeward Islets exhibit some windward characteristics; they lie
opposite and slightly north of a wide break in the windward reef which
allows trade winds to sweep, uninterrupted, across the lagoon. This
promotes their 60-80% cover of scrub or forest. Ana-Ana, the
southernmost, was periodically occupied from 1987-1991 by the Falconer
family.

Of particular botanical interest are the interior forests,
composed of Pisonia mixed with more Cordia than elsewhere on the atoll.
Pure Cordia groves (mostly too small to map accurately) typically occupy
the forest peripheries.

Their history (previous to 1987) is unknown; all appear to harbor
virgin plant communities mingled with occasional drift-derived Cocos or
Pandanus.

35) MOTU RAURAU "Blue-Gray Noddy Islet" (3.48 ha) Figs.) 29. 57;
Pls. 14, 69

Northernmost of the Southern Leeward Islets, we named this motu
for the Blue-gray Noddies (raurau in Gilbertese) observed there.

Physiography: Raurau is ovoid, with a small lagoonside bay, and maximum
dimensions of 180 m long and 231 m wide. It has the most expansive
rubble of all the Southern Leeward Islets. This coarse coral clinker
extends, apron-like, around the islet, widest (40 m) closest to the
lagoon, and narrower (10 m) to seaward. The seaward reef flats extend
446 m to the ocean.

Vegetation: The number of plant species is 10 (5 trees, 1 shrub, 4
herbs), 39% of the atoll’s flora. Raurau’s 2 plant communities are
simple: a very scant herb mat is sprinkled with Tournefortia, which
rises to 6-m-high scrub all around the islet. Laportea forms a narrow
band at the interface between coral rubble and scrub. Centrally a
Pisonia forest (to 13 m), dotted with Cordia on the periphery, harbors
much Morinda in the understory, including the tallest Morinda (13 m)
seen on the atoll. A handful of drift-derived Cocos and Pandanus, the
only introductions, dot the scrub.

168

RUBBLE

WM ters mat
fae] TOURNEFORTIA
CORDIA

"J PISONIA
(=) cocos

[a] HOUSE SITE
[=] PaNnpAaNnus
oe)

SURIANA

>_>:

MOTU RAURAU

lagoon

MOTU EITE}

PISONIA ISLET Acropora -Tridacna Reefs

MOTU KIMOA
ocean

<<“
af

MOTU ANA-ANA

nr

\ SOUTHERN LEEWARD ISLETS
ji
Ms Da ee ep ee
2 o meters 400
+
{ie

aoe

Figure 57. Vegetation and physiography of the 5 Southern
Leeward Islets: Motus Raurau ("Blue-gray Noddy Islet"),
Eitei ("Frigatebird Islet"), Pisonia Islet, Kimoa ("Rat
Islet"), and Ana-Ana ("Anne's Islet").

169

Birds: No seabirds were found on transect, but a perimeter walk in 1988
revealed 4 species breeding in the leeward Journefortia: Red-footed
Booby (10 pairs), Great Frigatebird (31 pairs), Brown Noddy (1 pair),
and White Tern (2 pairs). This islet, for its size, is particularly
rich in frigatebirds.

Comments: Polynesian rats are present.

36) MOTU EITEI "Frigatebird Islet" (1.41 ha) RigSen29), 57;
Pls. 14, 69

Second in line from the north, we named this motu for its nesting
Great Frigatebirds, e7te7 in Gilbertese.

Physiography: Motu Eitei is rounded, 105 m long and 280 m wide. Lying
perpendicular to the reef axis, it touches the lagoon edge on its inner
side. To seaward, the reef flats are 644 m wide.

Vegetation: There are 8 species of plants (3 trees, 1 shrub, 4 herbs),
31% of the atoll’s flora, with no introductions. Eitei is carpeted with
3 plant communities in the usual concentric arrangement. However, there
is a slight difference in the species composition of the herb mats: on
transect, the southern mat (2 m wide) consisted solely of Portulaca,
while the north side contained a 3-m swath of Heliotropium, Laportea,
and scattered Suriana. Inside the mat is a ring of Tournefortia scrub
(to 5 m) and a central Pisonia-Cordia forest (to 11 m). Laportea is
particularly abundant, while Portulaca, normally confined to the edges,
abounds in small openings within the interior woodlands.

Birds: Four species of seabirds bred on Motu Eitei in 1988: Red-footed
Booby (17 pairs), Great Frigatebird (14 pairs), Brown Noddy (6 pairs),
and White Tern (18 pairs). The atoll’s first Blue-gray Noddy nest was
found in 1990 (Pt. II).

37) PISONIA ISLET (2.45 ha) EigSia 29), 7 Se Pilisce ain 69
We named this motu for its fine Pisonia forest.

Physiography: Pisonia, third in the chain from the north, is almost
circular, and lies closely appressed to its neighbor islets. Its
maximum dimensions are 140 m long and 220 m wide. Like Raurau, it
possesses a wide "apron" of coral rubble and sparse herbs on the lagoon
side. Its seaward reef flats are 300 m wide.

Vegetation: The number of plant species is 15 (5 trees, 2 shrubs, 8
herbs), 58% of the atoll’s flora. The only introduction is Cocos (few,
scattered, north and south shores). Well-wooded, Pisonia harbors the
customary 3 plant communities: the herb mat is almost pure
Heliotropium, dotted with Suriana. One specimen of Lepidium bidentatum
was found in 1990. The Tournefortia scrub and forest, covering half of

170

the motu’s length and width, grows to 9 m, while the Pisonia-Cordia
forest, covering 0.86 ha (35% of the islet’s area), reached 10 meters.

Birds: Despite the fine Pisonia forest, no Black or Brown Noddies
nested. Only 3 species of seabirds bred in 1988: Red-footed Booby (26
pairs), Great Frigatebird (14 pairs), and White Tern (10 pairs). Best
represented were Red-footed Boobies; a perimeter count yielded 18 tended
nests, all in Journefortia scrub. A Long-tailed Cuckoo was heard in the
interior.

Comments: Rats were common: 6 were noted on a mid-morning transect
survey.

38) MOTU KIMOA "Rat Islet" (1.80 ha) Figs. 29, 05m
Pls. 14, 69,470

Fourth from the north, we named this motu for Caroline’s single
mammalian inhabitant, the Polynesian rat, kimoa in Gilbertese.

Physiography: Kimoa, smallest of the Southern Leeward Islets and shaped
like a flared teardrop, is squeezed between its neighbor motus. Its
maximum dimensions are 92 m long and 218 m wide, almost 4 times the size
mapped by Arundel (Fig. 4). The southeast rubble and herb mats are
wide. The distance to the outer reef edge is 307 meters. Of special
note is the emergent Tridacna-Acropora reef which stretches completely
across the lagoon to Tridacna Islet. This reef is 15-20 m wide

(Fig. 48; Pls. 25, 57) and 1,023 m long, which, together with an equal
length in blind diverticulae, totals over 2 kilometers. The Tridacna
clams aggregate in densities up to 80/sq m (Sirenko & Koltun, in press).

Vegetation: Kimoa has 11 species of plants (3 trees, 2 shrubs, 6
herbs), 42% of Caroline’s flora. There are no introductions. Though
small and narrow, Kimoa is well-vegetated. Its herb mats are composed
of Heliotropium on the south side and Portulaca (plus Suriana) on the
north. The interior Tournefortia-Pisonia-Cordia forests (to 11 m) cover
nearly half the islet’s area.

Birds: Four species of seabirds bred in 1988: Red-footed Booby (21
pairs), Great Frigatebird (3 pairs), Black Noddy (2 pairs), and White
Tern (7 pairs). Red-footed Booby nests occupied perimeter sites.

39) MOTU ANA-ANA “Anne’s Islet" (2.16 ha) Figs. 29. 575: Plsiawiar
14; 525w69,0 08

This motu includes a small settlement with 3 thatched huts
(cooking, eating, sleeping), a water tank, chicken coop, and garden. It
was occupied from 1987-1991 by Anne and Ron Falconer, 2 small children,
chickens, Muscovy ducks, and a dog. A wooden sign marked "Ana-Ana"
indicated that the islet had been named.

171

It is interesting to compare Plates 7a and 71, identical profiles
of Ana-Ana 105 years apart.

Physiography: Ana-Ana is the southernmost motu in the Southern Leeward
Islets, 120 m long by 222 m wide at its widest point. Approximately 3 m
high, it is roughly oval, with a hooked point and curved bay facing the
lagoon. This point is actively growing as more and more rubble is
deposited by the large flow of water passing through the channel (430 m
wide) that separates Ana-Ana and South Island. This channel contains
abundant clams that amass into an extensive Acropora-Tridacna reef
stretching approximately 900 m across the lagoon to Tridacna Islet. The
outer reef flats measured 281 meters.

Vegetation: Ana-Ana has 15 species of plants (5 trees, 2 shrubs, 8
herbs), 58% of Caroline’s flora. Introductions include Cocos,
vegetables, a few ornamentals and, as yet, no weedy exotics. Ana-Ana’s
vegetation is typical of the other Southern Leeward Islets except for
the settlement. Narrow trails from the southern channel lead to a neat
clearing, approximately 40 m x 70 m, the only inhabited portion of the
atoll. We advised the Falconers against introducing alien plants with
spreading seeds and requested them to destroy all introductions when
vacating the island permanently.

Ana-Ana has sparse herbaceous mats: Suriana, Heliotropium,
Portulaca, Laportea, and Lepturus. The Tournefortia strand includes
Cocos, Cordia, and Pandanus. Quality Pisonia forest, 15 m high, covers
43% of the islet’s area.

Birds: No breeding seabirds were found on any of the 3 visits to
Caroline. However, the Falconers found a few White Terns and one Great
Frigatebird nesting in the perimeter scrub, as well as groups of Brown
Noddies sitting on the beach. Long-tailed Cuckoos were seen around the
huts in March, April, and May 1990. It is to be hoped that seabirds
return now that the motu is again uninhabited.

Comments: Rats and large cockroaches are abundant. Despite the tidy
site, 12 rats occupied a pile of coconut debris, while others scurried
amongst the forest litter. The Falconers trapped over 1,300 rats one 2-
year period. Several pale geckos with a few spots and largish heads
were seen in and around the huts (probably mourning geckos).

I. CONCLUSION

Lushly wooded Caroline Atoll, with the majority of its 39 islets
(399 ha of land) either in near-pristine condition or having recovered
remarkably from past disturbance, is one of the least spoiled atolls in
the Pacific. Uninhabited, it harbors plant ecosystems and breeding
seabirds (Pt. II) of national and international importance. Its marine
and terrestrial ecosystems are prime outdoor ecological laboratories for
research on geological processes including ground water, sea level
changes, the dynamics of motu formation, fish poisoning, and numerous
facets of ecology including plant succession and Pisonia growth rates.

172

Caroline boasts prime coral reefs thickly studded with Tridacna clams,
substantial numbers of coconut crabs, breeding sites for green turtles,
wintering grounds for shorebirds including the rare Bristle-thighed
Curlew, ancient Tuamotuan marae, and a crystalline, unpolluted lagoon.
The variety, abundance, and quality of its flora and fauna qualify it
for status as an officially recognized international preserve (Pt. II,
Sect. G). Efforts toward its conservation have thus far been
unsuccessful: in 1992 it was leased to a private French businessman who
is currently fishing the reefs for commercial profit, as well as
disturbing seabird, turtle and coconut crab populations.

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Regional Environment Programme, Noumea, New Caledonia, and East-
West Center, Honolulu, Hawaii. 133 pp.

Thomas, W. L. 1961. The variety of physical environments among Pacific
islands, 7n Man’s place in the island ecosystem (F. R. Fosberg,
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iimaceyand. ols Jr., RP. Er Cloud) & Ki (0. Emery.) 19552. Conspicuous
features of organic reefs. Atoll Res. Bull. 46. Wash. DC:
National Research Council.

Trelease, W. 1884. Botany of Caroline Island, in Report of the eclipse
expedition to Caroline Island May 1883. Nat. Acad. Sci. Mem.
2:87-90.

Tsyban, A. V. & G. J. Smith. 1988. Protocol of the Third
Soviet-American Joint Expedition. Part II, The Central Pacific
Ocean and South China Sea, aboard the R/V Akademik Korolev,

9 September - 31 October, 1988. Unpublished Report. 6 pp.

Upton, W. 1884. Meteorology of Caroline Island, jn Report of the
eclipse expedition to Caroline Island May 1883, pp. 41-86.

U.S. Dept. of Energy. 1987. The natural history of Enewetak Atoll,
Vols. I and II, 228 and 348 pp., respectively. Office of
Scientific and Technical Information, U.S. Dept. of Energy.

Wagner, W. L., D. R. Herbst & S. H. Sohmer. 1990. Manual of the
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Walker, T. A. 1991. Pisonia Islands of the Great Barrier Reef, Pt. I.
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182

Wester, L. 1985. Checklist of the vascular plants of the Northern Line
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Yaldwyn, J. C. & K. Wodzicki. 1979. Systematics and ecology of the
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235:1-53.

Young, C. A. 1884. Introduction, in Report of the eclipse expedition
to Caroline Island May 1883. Nat. Acad. Sci. Mem. 2:11-20.

Young, J. L. Ca. 1922. Notes on Flint, Caroline and Vostock Islands,
7n Memoranda re Tahitian business. S. R. Maxwell & Co., Ltd.,
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183
APPENDIX I

Reef Information for Navigators

We include this section because no accurate hydrological chart
exists, and the Pacific Islands Pilot (Hydrographer of the Navy 1982)
section for Caroline is incomplete. Arundel’s 1883 map (Admiralty Chart
No. 979, Fig. 4) is still used today.

Caroline is visible "from aloft" at about 23 km (Ward 1974). It
has neither a deep pass, nor navigable channels into the lagoon, nor a
ship anchorage beyond the reef. In 1873 a set of moorings was placed
off the west coast of South Island for the convenience of guano ships,
approximately "a mile north of the south-west point, in about 60 fathoms
of water and some distance from the shore" (Arundel 1875). These are
long gone, although small boats can still anchor within the close lee of
South Island during normal trade winds. Today’s ships, however, must
drift well offshore after approaching the atoll from the west (Pl. 12).

Of special note is a possible extension of the perimeter reef
south and southwest of Caroline. Arundel’s map notes: "Reef reported
to extend four cables from southeast point." This information probably
originated in Findlay’s South Pacific Directory, quoted by Holden
(1884). Evidently the windward reef of South Island extends
approximately 1.7 km from its southeast point. From here "this reef
sends out two branches to a distance of 2.5 km, one toward the
southeast, the other toward the southwest and is consequently dangerous
to approach at night." Arundel’s map does not include this bifurcation
which, according to Findlay (1884), extends at least across the width of
South Island. He also states that "a landing (not always safe) may be
effected on the north side of the southwest bifurcation, described
above." No trace of these submerged reefs is evident on the RNZAF
aerial photos.

The "boat entrance" (Figs. 4, 50), a narrow nick in the outer
leeward reef, marked by the stock and ring of an anchor and immediately
to the west of South Island’s northwest point, is not necessarily the
easiest route to the lagoon. Landing is possible across the steep-to
reef at many locations along the leeward reef; opposite the southern end
of Ana-Ana is good.

Landing adjacent to the anchor is fairly straightforward in calm
seas, especially when one becomes familiar with the crooked notch which
narrowly pierces the outer reef. After negotiating a powerful backwash,
one’s boat is swept onto the shallow reef flats--liberally laced with
chunks of jagged reef--which is exposed at low tide and barely covered
at high tide. A swift current passes west out of the lagoon between
South Island and Ana-Ana, sweeping over the reef at the notch. Only
small craft with virtually no draft can effect the 500-m journey to
South Island. Because the shallows are unchanneled and not navigable
even at high tide, skiffs must be carefully hauled through the water to
a sheltered landing spot adjacent to South’s northwest point (Pl. 12).

184

An alternative landing method used by yachts in calm weather is
via the "blind passage" (Sect. D, Fig. 50), between northeast South and
Tridacna Islet. Despite the fact that the inner one-third of this
narrow diverticulum is calm, the outer two-thirds are rough most of the
time. Its channel leading to and from the open sea is particularly
turbulent and should not be attempted without land-based assistance, and
only at first light.

Landing is also possible across the reef flats off leeward Nake,
but there is no boat passage into the lagoon. Ward (1974) states that
at high water, light draught boats can land over the reef opposite the
middle of the western side of South Island. This would be the only
cross-reef landing which does not involve walking a boat across the
uneven reef flats.

APPENDIX IT

Weather Data, Caroline Atoll, 1989-1990
A. Wind Direction and Speed (mph), 1989

Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec
ee a EEE eee
1 NNE 15) NW 18 E 25 E 12 E 20 E15 E 25 E 20 - NE 10 =—N 20 NE 8
2 NE 10 ws E15 E 16 E 20 NE 18 SE 20 E 12 - WE 120) NW O15-17 NE 10

Variable
3) METI NE 20 E 12 SE 14 S$ 15-20 E 18 E 20 SE 25 - NE 15 WE 15 tw 20
4 NE 15 NE 15 Enl2 SE 15 E15 E15 E 20 SE 20 - WE 15 NE 12 Wd 20
5 NE 20 NE 15 NE 15 E12 NE 8 E15 E 18 E 18 - NE 12 E12 big Nw 18
swells
6 NE 20 NE 20 SE 10 NE 8 E15 E 10 E18 - NE 12 E 10 W 10
7 NE 12 SE 18 E12 E14 NE 10 NE 25 - E 12 - NE 15 WE 12 Nw 18
E 10
8 NE 12 Eas calm SE 10 E 10 NE 20 SE 12 NE 18 - NE 15 cale Ww 18
squalls
9 NE 12 E18 SE 10 E 20 Ears NE 18 SE 20 E 12 - NE 12 N10 NE 10
10 NE 15 E 16 NW 10 E 20 Eas ESS SE 16 E 12 - NE 10 NE 12 NE 10
11 NE 10 NE 16 calm SE 25-E 10 E15 NE 15 E15. calm = NE 10 E 18 NE 5
squalls E15
12 NE 20 SE 14 E 10 Wo14 Eis En15 Enl2 E 12 - NE 16 E17 NE 12
13. NE 15 eul2 E 12 E 16 NE 16 NE 16 E 12 - - NE 11 — 14 NE 12
14 NE 15 E 10 NE 10-18 €E 20 NE 16 NE 8 E 12 calm - ENE 10 E10 NE 14
15 NE 15 NE 8 E8 E 18 E15 E10 E 10 - - NE 10 calm NE 14
16 NE 15 SE 8 E12 E18 E 22 E 10 NE 10 - - NE 12 cale WE 15
squalls
17 ‘NE 18 E 20 NE 15 E 18 E 25 E15 E 10 - - NE 14 cale ME 12
18 NE 12 E 12 E8 E18 NE 18 E15 big E15 - - NE 15 N NE 14
swells
19 NE 12 E 12 E10 E18 NE 14 E 10 E 16 - - NE 13 Nw 10 E12
20 «NE 18 E 12 E10 E18 NE 14 E15 E18 - - E15 Ww 15 E10
21 =~NE 15 NE 10 SE 8 E14 E 12 NE 20 E 20 - - E 16 N 16 E 10 big
squalls swells
22 ~=-NE 20 E15 calm E14 E 12 E 10 E15 - - NE 14 WE 14 E 12-NE 10
23. «NE 18 thunder N 18 SE 14 SE 10 E 10 —15 - NE 20-30 E 20 WE 16 cale
squalls thunder
24 «=NE 15 E 25 N12 SE 14 SE 10-25 € 10 E 12 - E 20-30 NE 15 NE 12 WE 15
25 (NE 12 E15 N12 E 12 E 18 SE 17 E 10 = E 20 NE 15 NE 12 NE 16
a ee ee a ee ee eee
26 «NE 12 E15 N 10 E 10 NE 14 E10 calm = E 20 NE 15 N10 WE 15
27. —=NE 12 E 25 SE 10 NE 10 E18 E10 cala NE 17 E9 —9 WE 15
28 «NE 12 E14 E 12 E 13 E 18 E10 E 15-35 E15 NE 10 E8 WE 15
29 (NE 15 E14 E 18 big E15 E 18 15 N10 NE 8 EG, ME 12
swells

30 =NE 12 E 12 E 13 E 18 £15 calm cals - E 10

B. Rainfall, 1989-1990

Month/Year

Jan

Feb

Mar

Apr

May

Jun

Jul

Aug

Sep

Oct

Nov

Dec

Annual

Source:
aBasedon 16 days’ data.
Based on 9 days’ data.

b

"89
‘90

89
"90

89
‘90

89
‘90

89
‘90

‘89
‘90

89
590

"89
"90

"89
90

"89
‘90

89
‘90

"89
‘90

89
‘90

Mean Monthly Rainfall (mm)
Te
177.

160.
640.

259.
215.

190.
48.

66.
325%

48.
78.

45.
68.

35).
109.

50.
81.

13%
Wie

78.
134.

162.
154.

1,242.
2209!

Ron Falconer, Caroline

“Based on 18 days’ data.

HO Cor

Ore

NW Wo

ine op)

Atoll (pers. comm.).

185

Mean Number of Rain Days

186

ore ; ~
et 2

Plate 1. A 1988 dawn view of Caroline as seen from its leeward seas,
very similar to that seen by the atoll’s Western "discoverer,"

de Quiros, in 1606. Left to right: Motu Ana-Ana, Tridacna Islet,
South Island. Open reef area in the foreground is the 1883 chart’s
"boat entrance."

7m _ 2 Ny Bees ie v2 ;
Plate 2. A clearing on South Island from which the Solar Eclipse

Party made their observations in 1883. Today the area is covered with
dense Cocos forest (from Holden & Qualtrough 1884).

187

Plate 3. One of the 3 European-style houses that have ever been built
on Caroline, drawn in 1883 (ibid.).

Bee

UR ee can a
RPE ots ="

Plate 4. An artist’s very free rendering of Caroline in 1883 (from
Holden & Qualtrough 1884).

188

Pade rae : (3
Plate 5. Map of the "settlement" on South Island, as drawn by the
Solar Eclipse Party (ibid.).

Pl. 6a
Plate 6. Two
lagoon views
a century ago
along the
north coast
of South
Island (from
Holden &
Qualtrough
1884).
Compare these
drawings with
Plate 23.
Pl. 6b

189

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age

Plate 10. Black-tipped reef shark (Carcharhinus melanopterus),
numerous and aggressive in Caroline’s lagoon. Note the abundant sea
cucumbers (Ludwigothuria sp.).

19]

Plate 1l.

Laterally continuous, upraised older reefs of unknown age,
southwest Nake. Note the wide reef flats.

9 Sa ny:
se pita

Plate 12. Floating a small boat across the southwest reef flats in
calm weather from the "boat entrance" to the "landing" on South
Island.

Note the wide reef flats. The Akademik Korolev drifts
offshore.

192

Plate 13. Beach crest, sandy rubble, seaward moat and narrow reef
flats off southeast Nake Island.

Plate 14. Extensive lagoon reef flats south of Arundel Islet on the
windward side. Note the 5 Southern Leeward Islets in the distance.

193

Plate 15. An incipient motu, barely connected to Motu Mouakena’s
southern shore. See also Plate 62.

Plate 16. Successive ridges of coral rubble deposited by past storms,
northeast Nake Island.

Plate 17. Channel between the 2 northern islets, Long and Nake. Note
the mixed forest with Cocos and Pandanus, and filled-in upper lagoon.

Plate 18. Noddy Rock (0.02 ha), an emergent reef platform of unknown
age along the windward reef flats. A northward view. Islet is
completely awash during storms.

Plate 19. Windward beach, Long Island, showing wide rubble flats
inland of the beach crest, rimmed by oceanic flotsam and jetsam.

a et
Plate 20. Conglomerate platform, northwest point, South Island.

Russian vessel Akademik Korolev drifts offshore and a lone Suriana
maritima dots the blinding coral beach.

196

a

Plate 21. A large coconut crab (Birgus latro) shelters ina
subterranean cavity in the feo. Some males were among the largest

ever measured.

Plate 22. Sandy Inlet, a filled-in portion of the lagoon, extends its
fishhook-shaped mudflat 300 m northward into Nake’s landmass. Here
grow the healthiest and most productive Cocos on Caroline. Note the
Bristle-thighed Curlews in the foreground.

197

Plate 23. South Island’s pure Cocos plantation, looking west along
the lagoon. This extensive grove has now obliterated all traces of
the former "settlement" (Pls. 2-6). Note the dead Journefortia bush,
killed by excessive shade.

Plate 24. Crystalline lagoon waters adjacent to Emerald Isle (Central
Leeward Islets) are studded with submerged reefs and sandy channels.

198

Plate 25. A highly productive cross-lagoon reef of Acropora spp.
corals and Tridacna maxima clam shells joins Tridacna Islet with Motu
Kimoa. Sirenko & Koltun (in press) estimate 300,000 living
Tridacna/km.

Plate 26.

Cordia Forest (to 12.6 m tall), Pig Islet.

199

200

Pes

Plate 27. Sand, silt, rubble and hardpan mingle on the upper reaches
of Long Island adjacent to the lagoon.

Plate 28. Caroline’s sandiest beach flanks the lagoon shore of Shark
Islet.

201

Plate 29. Sheltered bay, Brothers Islet. Raurau Islet lies across
the lagoon. Note the sparse herb mat and silty shallow waters.

Plate 30. Narrow lagoon beach lined with Tournefortia scrub, Blackfin
Islet (Central Leewards).

202

Plate 31. Recent sand additions to South Island’s northeast point,
partly covered with excellent natural herb mats and healthy Sur7ana
scrub (right).

Plate 32. An old inter-islet channel (Tr. C, Long Island) filling in
with herbs, Tournefortia scrub and Cocos in 1988, but smothered with
fresh sand during the February storm of 1990 (see Pl. 33). Note the
nesting Masked Boobies in middle right. A westerly view toward the
lagoon.

Plate 33. The same area as Pl. 32, March 1990, 2 weeks after the
severe cyclonic weather. The herb mats had been smothered with sand,
and a large percentage of Journefortias were partly uprooted and
defoliated. An easterly view, toward the windward ocean.

i FA

ig

&

Plate 34. A clearing within the dying Cocos-Ipomoea forest, interior
South Island. Note the prolific mats of Boerhavia, Phymatosorus and
Ipomoea.

204

Plate 35. Pandanus forest, south Nake.

Plate 36. The north end wall of the ancient Tuamotuan marae,
northwest Nake.

ee 7 0

Plate 37. Mixed forest with Cocos, southwest Nake Island.

Plate 38. Orange, scarlet and green phalanges of Pandanus rest on a
clump of Portulaca. The ubiquitous Coenobita perlatus forage on their
stringy flesh.

Plate 39. Inner edge of lagoon, South Island, 1988. Cocos is
progressively shading out the beach scrub with Suriana maritima.

Plate 40. Inner edge of lagoon, South Island, 1965, taken from
approximately the same location as Plate 39. Note the greater extent
of sand and Suriana coverage above high water than today, due to less
encroachment and shading by the palms.

Plate 41. Heavy understory of Achyranthes canescens, Boerhavia repens
and Phymatosorus scolopendria in a clearing adjacent to Pisonia
forest, Pig Islet.

208

Plate 42. Boerhavia fruits on feathers and bill of a Great
Frigatebird.

Plate 43. Inside a mature Pisonia grandis forest, interior Nake
Island. Note the barren, dark aspect, virtually devoid of undergrowth
except root suckers. Appearing virgin, this quality stand is possibly
only 60-70 years old.

209

Plate 44. Fringe of Suriana, northeast point, South Island.

Plate 45. Well-developed natural herb mat, primarily Heliotropium
anomalum and Lepturus repens. Scattered Tournefortia forms a
"savannah." Here sandy soils support a lush Heliotropium cover,
northeast point, South Island.

Plate 46. Detail of Heliotropium anomalum, Skul] Islet, with remains
of the first evidence of tropicbirds on Caroline.

Plate 47. Tournefortia scrub, fringed by a natural herb mat, and
occupied by a colony of Sooty Terns. An old inter-islet channel,
northern Long Island. Note the nesting Red-footed Boobies.

2M

2t2

Plate 48. Tournefortia-Morinda forest, with nesting Brown Noddies,
interior Tridacna Islet. This is secondary growth, as this motu was
heavily planted with coconuts in the 1920s.

213

Plate 49. Skull Islet (0.02 ha), with Brown Noddy terns, looking east
to the windward reef.

Plate 50. Mature Pisonia grandis canopy with incubating Black Noddies
and a White Tern, Pig Islet. With a canopy height of 21 m, this was
one of the most majestic interior forests on the atoll, although it is
only approximately 65 years old.

214

Plate 51. Azure Isle (Central Leewards)--an example of a motu
containing a single Pisonia tree. Note the narrow, but still shark-
patrolled, inter-islet channel. View east from Motu Nautonga, with
Brothers Islet in the distance.

Plate 52. Caroline’s sole clearing, with now-abandoned Tahitian-style
huts, Motu Ana-Ana.

215

Plate 53. Piles of fibrous shavings--coconut crab sign.

Plate 54. Beachrock at the lower, windward tip of Long Island,
typically found at low water.

216

Plate 55. Windward Islets nos. 5-9 (left to right): North Pig, Pig,
Skul] (not visible), North Brothers and Brothers.

Plate 56. Arundel Islet (foreground), looking south-southwest across
Tridacna Islet to South Island. Distant Motu Ana-Ana lies on the
right.

Plate 57. Detail, Tridacna maxima reefs, lagoonside of Tridacna
Islet. This dense aggregation of giant clams amassed up to 80 per
square meter.

Plate 58. View of Pandanus Islet (center) west down the channel
separating Nake (right) and Long (left) Islands.

218

Plate 59. Danger Islet (South Nake no. 2), looking due west across
the shallow upper lagoon from Long Island.

Plate 60. South Nake Islets nos. 3-6 (right to left): Booby, Coral,
Lone Palm, and Kota. Westerly view across the shallow upper lagoon
from Long Island.

ZS

Plate 61. Lone Palm Islet (South Nake no. 5): a southerly view from
the shallow tidal flats of Coral Islet.

Plate 62. Motu Mouakena (South Nake no. 7), with connected cay. A
westerly view from the upper lagoon. Compare with Plate 15.

220

Plate 63. North end, Motu Mannikiba "Seabird Islet" (Central Leewards
no. 1) showing mounds of Pisonia and a closer Cocos grove. Note the
circular patch reef at right.

Plate 64. Motu Mannikiba, looking east along Transect 1. Natural
herb mats on coral rubble give way to Journefortia, then a distant
patch of Pisonia. The extensive interior forests of this motu were
felled 70 years ago to support a well-maintained coconut nursery.
Forest recovery has been much slower than on islets where the Cocos
was not managed as intensively.

221

Plate 65. Motu Mannikiba, looking east along Transect 2. Low
Tournefortia scrub covers a coarse rubbly substrate, probably a former
inter-islet channel.

222

Plate 66. Mixed Pandanus-Tournefortia forest, interior Emerald Isle.

223

Plate 67. Emerald Isle, looking west across the open lagoonside scrub
and hardpan to a densely vegetated interior.

es :

ig itt
ceca

—

3

. gs — = ~~

ee :

aa
_ .

“3
aie -

Plate 68. Shark Islet (Central Leewards no. 5): view across patch
reefs to a sandy beach.

224

Plate 69. Southern Leeward Islets nos. 1-5 (right to left): Motus
Raurau and Eitei, Pisonia Islet, Motus Kimoa and Ana-Ana. View
northwest from Tridacna Islet.

Plate 70. View of Motu Kimoa ("Rat Islet") from Pisonia Islet. Its
central forest, typical of the Southern Leewards, is a mixture of
Pisonia and Cordia.

Plate 71. Motu Ana-Ana ("Anne’s Islet"): a view with giant ray, from
the shallows of the lagoon’s southern end adjacent to South Island.
Note the similarity to Plate 7, dating from 1883.

Plate 72. The "blind passage" (non-functional hoa), looking west from
its inner end across the shallow reef flats to the lower lagoon.

Ba, " coe ye fs i “re

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ATOLL RESEARCH BULLETIN

NO. 398

PART I. SEABIRDS, OTHER TERRESTRIAL ANIMALS, AND
CONSERVATION

BY

CAMERON B. KEPLER, ANGELA K. KEPLER, AND
DAVID H. ELLIS

ISSUED BY
NATIONAL MUSEUM OF NATURAL HISTORY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C., U.S.A.
FEBRUARY 1994

: en Gp reai
S (oTeni JecatTAR 7 MONE
moet TEAL :
ARS Se WO
_ sate ema 1

Table of Contents

ISIS GOf- PIUGURES IRM er Mates bevel cn fonrceh. cy Ai oie MEM eeter cwtohtnels Aa,
LSE OR TPE N CLs be Rat NCU Bi 8 lee ie AE EY Bap BAD pe 2 Te Aa a OM et
TUES HES O ful ta LS seet a ree dev car cercecons iy AN caret cea A anarchy Orgs Neaueatie ales teyaledvau sam ray ol yletcey al laVvastehati

RE HiSORYMOtmONME NOMOGHGANESCUGTES cdots ttat.1erer/eveyercreueneweloncealtes: oaPare ake: «
MEO diSiecrstesestetercteaseneie reece stans fee stan eiseirscy ws cetang arose nw lonehay eve nie ete enaitce wre shes Ges
Seabind: Species ACCOUNTS. SF. WT LER es RLS RAL
Other BiindssonaGarolhiner Atovlilic Sasirarrecstsietes ccteeleitor leis. «c telteleie.
OURS REVEGLED RACES retary crete cle, rau ciells) careers a Sicce\enele feel aloes aue(eiare| ol ater sbeuae les

LEWZIAVG'Stastste Vole posseits coede ratchet eataneohectins Rete cits Geegiece Cs 5 Ck. KURI CNR US ENES UN LA EEE.

moONw

Qn

. Conservation: Attributes of International Significance..........
EackotiaM al oma DmSiGURDAMGES rari ese vocsvsccr coker cner-bonete Sia en cele eliston ot dante oreaes
MERGES EVAIANMECOSYSILEMSRiees LRG et he SE ee LOA
EMYSMO GMA Mivsasrsus terse te ersgoweUstcmetese tae cchis Wota rain Gals cl atenntantehaahalametenehe 'ovetatel tel ers
Fala raped teste ne eae a lea aunt ley cy sl ice awiatrboat cist eclgh sud st bates stated shel statial on atlahal al Slate

COCOMUIM Crial SERS rercrororee eas reek tote ost eta Deiat ey ai AUR oelletnnalelsraiaceness
MU vail @'Svcnc sedans voncncncterts te rciccne eReader eee oa la eos eae MTC hw uot Cae teats REDE Fatah BE
AVCHACON OGY INS aectehotie Sikorski aioli cre . (alster ane a Sista Sa Gd we cietendire
Curent ConsenviatttiloneSibaituisiyy te tec i ee TR UO eed
Heme iO Ra UICeL Cie G Mei tewencu vteter ecesarevcrseeteneden a eet etemacetine MORN CANCER a ECS Pe.
Appendix: aPreliminayy aise woh Arthropods... 1... sa. oe cass ee se ces 6

i

List of Figures

Caroline Atoll, Republic of Kiribati, with newly-named
TSILCES 6 oscse 5 0.00 Wa pu Wd Oem wewerdle Seeeewbiewten SCE eee

Distribution map of breeding Red-tailed Tropicbirds and
BY OWN, BOODICS.. x... 00.30.00 wos oe wtebeleiele wie oleleus Oke tated Lfe eee eee

Breeding phenology of the Red-tailed Tropicbird on Caroline
Atoll,,. May-September, 1988............3\......CG0% .GRELR&ho fie pier eee

Distribution map of breeding Masked Boobies................008-

Breeding phenology of the Masked Booby on Caroline Atoll,
April-September 1988. . «.. oo siweiceree sie ors 00 oie ee ease sone ehetele ihe eee tena

Distribution map of breeding Red-footed Boobies................

Breeding phenology of the Red-footed Booby on Caroline
Atoll,. June-September 1988). ............. 0:01.00 .8 ue .ose s¥ehelelete et eee

Distribution map of breeding Great and Lesser Frigatebirds.....

Approximate breeding phenology of the Great Frigatebird on
Caroline Atoll, February-September 1988... 2. ..< 5 aj0u sessed

Breeding phenology of the Lesser Frigatebird on Caroline
Atol1,. February-September 1988.........0.... 0.0 0.0.0.» obese ies oe

Distribution map of breeding Sooty Terns..............0-s sere

Breeding phenology of the Sooty Tern on Caroline Atoll,
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Distribution map of breeding Brown Noddies...............-eee0e-
Distribution map of breeding Black Noddies...............sssenee
Distribution map of breeding White Terns.......... <2... eee
Distribution map of the Long-tailied Cuckoo...:...... 0.2
Distribution map of coconut crabs. .. 5...» sims. 0 «cle or olelsietetnineaene

Map showing seabird breeding species diversity by islet,
Caroline Atol Vis. .0.0.6:s06%5 see 00.0.0 ea 6 oie 6 0 ore aisle ee See

Page

List of Plates

Incubating Red-tailed Tropicbird, Bo’sun Bird Islet..........

Adult Masked Booby with egg, coral rubble with Portulaca

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List of Tables

Estimated number of breeding seabird pairs on Caroline

AGO September WlOSB. ce crate: sraic-oreie on cisiee oo oie)o eles dlsloleie else. 0 8 os

Stages in the breeding cycle of the Red-tailed Tropicbird,
Caroline Atoll, 27-29 September 1988 (ages after

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Stages in the breeding cycle of the boobies of Caroline

Atollie 2129) September tiS88eec oo bs dsecess sc csle eg cles beseees eos

Density of Red-footed Booby nests in occupied Tournefortia

habitats on islet groups, Caroline Atoll, September 1988.....

Stages in the breeding cycle of frigatebirds on Caroline

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Sooty Tern colonies on Caroline Atoll, 27-28 September

1988 eos eee eee eo ese eee eee eee eee eee ee eee e eee eo eee eee eoe eee e ee eee @

Density of White Terns on occupied islets by islet group,

Caroline AtonleeSeptembernklose. sync fa. eo cto. Sete eee ce
Lizards collected on Caroline Atoll, 1965-1988...............

Comparative abundance of Caroline’s breeding seabirds in

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PART Il. SEABIRDS, OTHER TERRESTRIAL ANIMALS, AND
CONSERVATION

BY
CAMERON B. KEPLER', ANGELA K. KEPLER’, AND
DAVID H. ELLIS?

ABSTRACT

Approximately 1,000,000 seabirds of 11 species bred on Caroline
Atoll between September 1988 and June 1990. The most abundant species,
with over 900,000 birds, was the Sooty Tern. Two species (Red-tailed
Tropicbird, Blue-gray Noddy) are reported breeding for the first time.
The known seabird fauna now includes one tropicbird, 3 boobies, 2
frigatebirds, and 5 terns.

Seabird distribution on Caroline is determined by the distribution
of plant communities and nonvegetated substrates, the prevailing trade
winds, and to a lesser extent, rats and coconut crabs. Red-tailed
Tropicbirds and ground-nesting Brown Noddies nested on small islets
relatively free of rats and coconut crabs. Masked and Brown Boobies
preferred exposed windward beaches, primarily on Long and Nake. The
tree-nesting Red-footed Booby and frigatebirds attained their highest
nest densities in areas with reduced wind speed. Black Noddies were
found in dense colonies, generally high in pisonia trees, while the
uncommon Blue-gray Noddies nested solitarily on open coral rubble.
Sooty Terns nested in large colonies, generally near or under relatively
open Journefortia scrub, but also in open areas under Journefortia and
closed-canopy Pisonia forests. Not all sites were utilized annually.
Tree-nesting Brown Noddies and White Terns, found throughout the native
forests, were the only species utilizing anthropogenic forests. The
lowest seabird population densities were found in the disturbed forests
on South and southwest Nake, and no seabirds nested on inhabited Motu
Ana-Ana.

About 300 Bristle-thighed Curlews overwinter on Caroline, foraging
in all terrestrial habitats, including Pisonia and disintegrating Cocos-
Ipomoea forests. We extended the known winter range of the Long-tailed
Cuckoo by discovering a small population on the atoll, the first record
for the Southern Line Islands.

The known lizard fauna was increased from 3 to 6 species.
Approximately 2,200 coconut crabs inhabited 12 islets on Caroline.
Although primarily associated with Cocos, we also found them in Pisonia
and Tournefortia. Pacific green turtles breed in small numbers; we
found the first known nests (old) in 1990. Polynesian rats are
abundant, and Pacific bottlenose dolphins were seen close to the
windward side in 1990.

National Biological Survey, School of Forest Resources,

The University of Georgia, Athens, Georgia, 30602, USA

400 Snapfinger Drive, Athens, Georgia, 30605, USA

National Biological Survey, Patuxent Wildlife Research Center,
Laurel, Maryland, 20708, USA

Manuscript received 8 April 1991; revised 26 October 1993

The populations of seabirds and coconut crabs on Caroline Atoll
are of national and international importance. The Black Noddy (17,000
birds) and White Tern (8,000 birds) populations are the largest in the
Republic of Kiribati, while the Red-footed Booby population (7,000
birds) is one of the largest in the world.

Caroline’s conservation attributes are numerous: large, varied
seabird populations, wintering grounds for several species of migratory
shorebirds and the Long-tailed Cuckoo, plant ecosystems of national and
international importance (lushly wooded motus with significant groves of
Pisonia grandis, Cordia subcordata, Tournefortia argentea), virtually
pristine coral reef ecosystems exceedingly rich in giant clams (7Tridacna
maxima), a coconut crab population of Pacific-wide importance, ancient
Tuamotuan marae (religious sites), and a breeding site for green
turtles. The marine ecosystems, terrestrial biota, and geology of its
39 motus of varied size, 22 of which are pristine or near-pristine,
provide prime outdoor ecological laboratories for numerous areas of
research. These qualities, plus its exceptionally clear, essentially
unpolluted lagoon, and its relative lack of disturbance, render it
worthy of permanent protection.

A. HISTORY OF ORNITHOLOGICAL STUDIES

"There were a great quantity of sea birds of several kinds, and so
importunate that they seemed to want to attack the men" (Markham 1904).
So wrote the Portuguese explorer de Quiros on 21 February 1601, the
first European to see Caroline Atoll.

Precisely which seabirds were present remained a mystery until the
island was surveyed 364 years later by the Pacific Ocean Biological
Survey Program (POBSP) (Clapp & Sibley 197la). Prior to that visit,
accounts of the avifauna had been incomplete. Bennett (1840) described
Red-footed Boobies, a frigatebird (species ?), White Terns, Bristle-
thighed Curlews, tattlers, and "a great number of small pigeons" with
white heads (certainly noddy terns, perhaps both A. minutus and
A. stolidus). The "shoal birds" that greeted him were probably Sooty
Terns. His most unusual contribution was mention of a possible
flightless rail: "The other birds of the coast were a kind resembling a
GOOGer ca: (Dee ST2)\.

The 1883 Solar Eclipse Party (Pt. I, Sect. B) published a few
sketchy notes, adding Lesser Golden-Plover, Reef Heron, and Masked Booby
("gannet") to the bird list. Of dubious identity were 2 species of
"seagull" and a "snipe" (Dixon 1884). Holden, one of the astronomers,
heard "the notes of a singing bird," which prompted us to add mist nets
to our equipment in the hopes of capturing an Acrocephalus warbler.

This resulted in our discovery of the Long-tailed Cuckoo (Ellis et al.
1990) and piqued our curiosity about what Holden might really have
heard.

The POBSP expedition spent 3 days on Caroline in June 1965. They
found 10 species of seabirds (9 breeders), 4 migrant shorebirds, and a
Reef Heron (Clapp & Sibley 1971a), providing rough population estimates
for each species. This work laid the foundation for later expeditions.
Brief visits to Caroline by the Kiribati government in 1974 (Gilbert &
Ellice Islands Government 1974, Vickers 1974) and Roger Perry in 1977
(Perry 1974, Garnett 1983) added no further information.

The 1988 expedition to Caroline was longer and more extensive than
all former visits. We found 3 new island records: a breeding seabird
(Red-tailed Tropicbird), a shorebird (Sanderling), and a migratory land
bird (Long-tailed Cuckoo), and mapped islet-by-islet distributions for
each species. Our population estimates, calculated from numerous
transect surveys, aerial photographs, and detailed vegetation analysis,
indicate that Caroline’s avifauna is far more important than had
previously been suspected (King 1973, Garnett 1983). In March and May
1990, the ICBP 1990 Line and Phoenix Islands Expedition (Pt. I,

Sect. C), of which AKK was co-leader, filled in minor gaps in our
knowledge. Seven-year-old Alexandre Falconer, then living on Caroline,
added another breeding seabird, the Blue-gray Noddy, in summer 1990.

B. METHODS

From 22-29 September 1988, C. B. Kepler, A. K. Kepler, D. H.
Ellis, and K. Teeb’aki surveyed all of Caroline’s 39 islets except North
Arundel Islet, naming most of them (Fig. 1; see also Pt. I, Sect. C).
We established 50 linear transects, extending 13,300 m x 30 m, laid out
to ensure that at least 5% of each islet was sampled for birds and
plants (see Pt. I, Sect. C and Fig. 9). Sampling was increased with
19,300 m of perimeter surveys along the windward and leeward coasts of
dilmaisiets (Pts 1, %Figse9) a" OnaNoddy Rock; Skull; Atibu, ‘Bo’sun Bird);
Coral, Reef-flat, and Fishball (Fig. 1) we made total counts of the
breeding seabirds. All surveys were conducted during daylight hours.
Some incidental data have been added from the 1990 ICBP expedition.

Distribution and habitat preference: We described 7 major plant
communities on Caroline Atoll (Pt. I). With the use of aerial photos

and the transect data, we mapped the communities found on each islet.
Bird distribution was determined and plotted using these islet
vegetation maps. If a species nested within a particular plant
community, it was plotted on the distribution maps as occurring
throughout that community unless determined otherwise.

Population sizes and breeding phenology: We measured transect distances
for each islet using a hip-chain and biodegradable cotton thread. We

recorded all birds seen within the 30-m-wide strips; transect width was
estimated visually. We assigned birds to one of several mutually
exclusive categories: adults present, adults on territory, adults on
nests (contents unknown), eggs, naked chicks, downy chicks, chicks with
remiges erupting, chicks with scapular feathers, or chicks in juvenile
plumage. We created a range of possible laying dates for each egg and

PANDANUS iSLET

DANGER iS Cera tS | \
SOUTH NAKE soosy ister ——__ \
Ch (i) ——— Ss \
ISLETS | LONE PALM ISLET i \
| MOTO. en)

mote MOUAKENA Tees iF. @|

—— LONG !ISLAND

— lagoon
MOTU MANNIK!BA ———————
} amt
BLACKFIN ISLET BO SUN BIRD ISLET
CENTRAL | MOTU MATAWA aR —
LEEWARD
EMERALD ISLE ————_____/ WINDWARD ISLET
ISLETS SHARK {‘SLET ————________/ SB
SCARLET CRAB ISLET —- CRESCENT ISLET
i i.
Ragin ! ESS MORPAANGS WINDWARD
REEF -FLAT ISLET i= ©@ »— NORTH PIG ISLET
BIRD ISLET ISLETS
FISHBALL 1SLEy ————~~—— e aw ISLET
/ = ——— SKULL ISLET
Zz & —t— NORTH BROTHERS ISLET
f —+— BROTHERS ISLET
SOUTHERN a7 ~— 7 noopy rocK

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ISLETS MOLU KINOA

MOTU ANA-ANA ———_]__

NORTH ARUNDEL ISLET
—~— arunvet ISLET

a TRIDACNA ISLET

f blind channel
}

ja ee ISLAND CAROLINE ATOLL

SCALE 1:34,000
800 ° soo 1000 2000

WE —E———EeEE ee

meters

Ng

Figure 1. Caroline Atoll, Republic of Kiribati, with newly-named islets.

chick using known growth parameters for each species (C. Kepler 1978,
Kepler & Kepler 1978). This enabled us not only to estimate seabird
populations, but also to determine and plot a rough breeding phenology
for each species (Figs. 3, 5, 7, 9, 10, 12). In these figures, the
height of the bar for each category ("downy," "scapulars," etc.)
represents the number of nests found or estimated with that development
stage in September 1988. The bar width represents the approximate time
span over which eggs could have been laid to produce that stage, while
the "no. days" is a count back from the survey dates to accommodate
growth and development that had occurred. Thus, while each figure shows
what breeding stages we found, we extend those nests back in time to
show roughly when they would have begun. The number of clutches begun
per day is determined by dividing the number of nests per stage by the
time span in days over which those eggs were laid.

Sooty Terns pested in dense colonies. Each colony was mapped, and
its total size (m~) was calculated. A minimum of 10 plots (3 m x 3 m or
3 m x 6 m), within which all eggs and chicks were counted, were randomly
located along a compass line in each colony. The population size of
each colony was estimated from these plot densities.

Mist nets: We operated 4 ATX 4-shelf 36 mm mesh mist nets (2.6 x 12 m)
for 43.5 net hours, according to the following schedule: 14.5 net hours
(daylight) beneath a 10-15 m Cocos canopy on South, 27.5 net hours (day
and night) in Pisonia-Cocos interface (12 m tall) near Transect 10 on
Long, and 1.5 net hours in Pisonia-Tournefortia within a 4-6 m canopy on
Transect 4, Long. One cuckoo was collected (USNM 607191).

Collecting other vertebrates: Lizards that were active and conspicuous
were collected at base camps on South and Long, either by hand or with a
blowgun firing steel darts. No attempt was made to search for reptiles
under coral, litter, or in other concealed locations. Rats were
collected with a blowgun or snap traps baited with coconut, the former
proving far more effective because most traps were sprung by hermit
crabs. We preserved all specimens in formalin and sent them to the U.S.
National Museum.

C. SEABIRD SPECIES ACCOUNTS

Eleven species of seabirds occur at Caroline, most of which breed
in large numbers. They include one tropicbird, 3 boobies, 2
frigatebirds, and 5 terns.

RED-TAILED TROPICBIRD (Phaethon rubricauda) FICS 45 Svs

Red-tailed Tropicbirds breed at widely scattered locations
throughout the tropical Pacific and Indian Oceans. In the Line Group,
they nest from Palmyra south to Starbuck (Perry 1980), with a large
population (8,500 birds) on Christmas Island (Clapp 1967). Prior to our
expedition it was unrecorded from Caroline, Vostok, or Flint.

* BROWN BOOBY
© RED- TAILED TROPICBIRD

CAROLINE ATOLL

SCALE 1:34,000

500 ° S00 1000 2000
meters

Figure 2. Distribution map of breeding Red-tailed Tropicbirds and
Brown Boobies on Caroline Atoll, September 1988. In this and the
following distribution maps, arrows indicate concentrations of
breeding birds.

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Distribution and habitat preference: Our first indication that Red-
tailed Tropicbirds nested on Caroline was the discovery of a skull, tail
feather, and broken egg (Pt. I, Pl. 46) under a small Journefortia bush
on a previously unnamed islet between Pig and North Brothers Islets. We
named this sparsely vegetated collection of rubble "Skull Islet" (Pt. I,
Pls. 46, 49). We later found 47 nests on another islet, naming it
Bo’sun Bird (Fig. 1) after the species’ common name.

All nests were jocated under relatively open Journefortia scrub
less than 3 m tall in open, windy locations, with the majority (91%) on
small islets (Q.24-0.86 ha). All nests were under shrubs with few stems
within a 0.5 m“ nest space, and most had peripheral cover on the sides
of the shrubs, both important factors in nest-site selection (Clark,
Ricklefs, & Schreiber 1983). All nests were in areas relatively free of
Polynesian rats (Rattus exulans) and coconut crabs (Birgus Jatro): five
nests on Long were within 50 m of the island’s south point.

There are large populations of Polynesian rats and coconut crabs
on Caroline’s bigger, more wooded islets. This rat, though basically
vegetarian, is an effective seabird predator (C. Kepler 1967, Norman
1975) that in some years has taken 65% of the Red-tailed Tropicbird eggs
and 100% of the chicks on Kure Atoll (Fleet 1972). Coconut crabs are
also known bird predators (Helfman 1979, Reese 1987): on Caroline in
1965 they preyed upon Sooty Terns (Clapp & Sibley 197la), and in 1990
AKK photographed the aftermath of predation or scavenging on at least
one species of tern on Brothers Islet. It may be no accident that
tropicbirds on Caroline occur only on small, relatively open islets that
harbor few, if any, rats and crabs, and the southern tip of Long Island,
where predator densities are low. We saw no rats on Bo’sun Bird Islet.
Although rats could swim the 165 m to the islet, the nearly continuous
presence of black-tipped reef sharks (Pt. I, Pl. 10) in the channels
surrounding the islet provides protection to its nesting tropicbirds.

Numbers: In September 1988, we found 56 active nests on 5 islets

(Fig. 2, Table 1) and estimated a minimum population of 60 pairs. The
May 1990 expedition found 130 nests on Bo’sun Bird; our revised estimate
for Caroline is approximately 300 birds. Bo’sun Bird Islet was surveyed
by POBSP in June 1965, and no tropicbirds were located on the ground or
in the air (F. Sibley, pers. comm.). It is unlikely that Red-tailed
Tropicbirds were present but overlooked at that time, suggesting that
they have colonized the atoll only recently. The Caroline population is
now the second largest colony known from the Line Group, and Caroline is
only one of 5 islands in the archipelago where Red-tailed Tropicbirds
are known to breed.

Phenology: Of the 56 nests found in 1988, 54 contained eggs or chicks
(Table 2). The 33 chicks were divided into 4 age classes (Fleet 1974,
Diamond 1975a) which, together with the 21 eggs, provided an indication
of laying phenology for 140 days prior to our arrival (Fig. 3). Eggs in
surviving nests had been laid at a fairly even rate from early May
(possibly starting earlier) through September. The finding of only 2
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Table 2. Stages in the breeding cycle of the Red-tailed Tropicbird,

Caroline Atoll, 27-29 September 1988 (ages after Stonehouse
1962). -

Nest Stage: Juv. Remiges Scapulars Downy Egg Pairs on Territory

Approximate

Age in Days

From

Laying: 90-133 69-89 58-68 44-57 =0-43 -
No. Nests: 18 4 5 6 21 2

that laying was ending. On 24 May 1990, many nests contained eggs and
downy chicks (75% nests with chicks) and pairs were still courting.

On Christmas Island, peak laying generally occurs from June to
October (Schreiber & Ashmole 1970), later than those parts of the 1988
and 1990 breeding seasons we observed on Caroline.

MASKED BOOBY (Sula dactylatra) Figs. 4, 53) Pitez

The Masked Booby is widely distributed in the Atlantic, Indian,
and Pacific Oceans. Clapp (1967) estimated that 19,100 Masked Boobies
bred in the Line and Phoenix Islands, with about 13,000 of them in the
Line Islands, mostly (ca. 9,000) on Jarvis.

Distribution and habitat preference: Eighty-four percent of Masked
Booby nests (159) were on the windward, rubbly shores of Long and Nake
Islands, extending to the north end of the atoll. Fifteen additional
nests were scattered along the lagoon edges of 5 South Nake Islets
(Table 1). Nests consisted of bare scrapes with exposed sand, usually
within a sparse ground cover of Portulaca and Heliotropium (Pl. 2).

Over half the nests were amassed in one open colony on Nake that
extended nearly 1,000 m, beginning approximately 150 m south of

Transect 2 and extending about 50 m north of Transect 4 (Pt. I, Fig. 8).
Here a nearly unbroken Heliotropium mat 30-80 m wide, with patches of
Tournefortia, occupied the area between the leading edge of the
Tournefortia scrub and the beach crest. Nests were 20-30 m apart in the
densest section (near Tr. 3). All nests were exposed to the sun, unlike
those of the Brown Boobies. Some adults and juveniles roosted under the
scrub; guano deposits indicated regular occupancy.

A loose group of 7 breeding pairs was scattered on a broad plain
of low herbs along a partially filled old interislet channel 370 m south
of the north end of Long Island (Tr. C, Pt. I, Figs. 8, 40). Four more
pairs nested in coral rubble along the channel separating Nake and Long,

11

50% of population

MASKED BOOBY

CAROLINE ATOLL

SCALE 1:34,000
500 ° 500 1000 2060

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Figure 4. Distribution map of breeding Masked Boobies on Caroline Atoll,
September 1988.

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one pair with a downy young only 2-3 cm above high-tide flow on an
"islet" between fingers of the channel, a precarious location where
nesting surely must fail in stormy periods. No birds were seen there in
March and May 1990, following a severe storm in February 1990. Four
pairs nested singly along a leeward 1,000 m stretch of lagoon shore on
the northern end of Long (Fig. 4); hardpan was the primary substrate.

Numbers: In September 1988 we found 189 Masked Booby pairs (Table 3),
including those on territory (with or without nest scrapes) and
juveniles (with or without attending adults). We found no "clubs" of
nonbreeding birds. We covered most of the habitat favored by this
species except the northern 300 m of Nake Island; in 1990 a few
scattered pairs nested there. Our population estimate, including pairs
we might have missed, was approximately 200 breeding pairs. Other
population estimates were "ca. 10" birds (Clapp & Sibley 197la) and 50 +
15% (Grossman & Grossman 1974). In 1965 POBSP biologists (F. Sibley,
pers. comm.) surveyed all locations where we found breeding pairs.
Thus, 200 pairs represents a major increase in the population on
Caroline Atoll.

Phenology: In June 1965 only 4 Masked Booby nests containing eggs were
found (Clapp & Sibley 1971a), indicating that nesting began in May or
June. On 9-10 September 1974, Grossman & Grossman counted 23 nests
containing "eggs and nestlings" on windward Long and part of Nake. We
found nests in all stages in September 1988 (Table 3, Fig. 5). The
large age class in April may include some juveniles that could fly (i.e.
were older than 180 days). We may have undercounted naked chicks, not
wishing to expose them to the sun by frightening the brooding adult.
Laying began in April or earlier, peaked in June and July (Fig. 5), and
continued until our survey in late September. The 34 pairs on
territory, many with nest scrapes (Table 3), indicated that laying was
still in progress and would continue into October.

Table 3. Stages in the breeding cycle of the boobies of Caroline Atoll,
21-29 September 1988.

Nest stage /Approximate age in days from laying
Flying Pairs on

Species Juv. Juv. Scapulars Remiges Downy Naked Eggs Territory

Masked >164 145-164 115-144 89-114 55-88 45-54 0-44
Brown >164 144-164 114-144 88-114 54-88 44-54 0-44
Red- footed = >150 T11-150_ 975-110 =54-74 45-53 0-45

Masked - 40 12 22 38 4 33 34
Brown 1 - - - - - 3 8
Red-footed many - - - 29 - 919 1,270

l For descriptions of nest stages see C. B. Kepler (1978).

14

In March 1990, 31 pairs were on territory or were attending nests,
eggs, or older chicks, indicating that a new breeding season was
underway as the previous season was ending. By May 1990 there were 63
nests, mostly with eggs, and there were no older chicks. Thus, the 1990
season augments the 1988 data and suggests an annual cycle with egg
laying beginning slowly in February and March, peaking in June and July,
and declining to a low ebb from December to February.

The large number of fledged juveniles and nests with older chicks,
in both September 1988 and in March 1990, indicated that the 1988 and
1989 breeding seasons were very successful. It also suggested that
potential predators (rats and coconut crabs) posed little hazard to this
hardy species.

BROWN BOOBY (Sula Jeucogaster) FiGuet

This widely distributed pantropical species has an estimated
population in the Line and Phoenix Islands of about 3,200 (Clapp 1967,
Perry 1980), with over half of them (2,000) recently found on Malden
Island, in the Southern Line Group. However, all other estimates of
this species on Malden from 1964 to 1980 (the 16 years after pigs were
eliminated) are below two hundred.

Distribution and habitat preference: Breeding Brown Boobies on Caroline
were restricted to the windward edges of Journefortia scrub and forest,
generally within 15-20 m of high water. In 1988 we found nests on 4
islets (Fig. 2, Table 1). Long, with 12 pairs, was the only islet
supporting more than a single pair. They were located on the northern
two-thirds of the island: 4 pairs formed a loose colony near the head
of Transect A (Pt. I, Fig. 8). All nests were under Journefortia bushes
approximately 3 m tall. In March 1990, we found 20 pairs of Brown
Boobies, all on windward Nake as far as the islet’s northern extremity.
There was no evidence of nesting on Long Island. On May 22, 1990, only
3 nests, all with eggs, were found on Nake.

On 22 September 1988, we saw 2 birds plunge-diving with Masked and
Red-footed Boobies approximately 500 m west of South Island. On the
atoll, flying Brown Boobies were observed soaring only along the
windward beaches. Two birds roosted on the south-central beach of
South, and another was found roosting on Kota.

Numbers: We counted 15 pairs during perimeter surveys in 1988, yet
found none on the transects. Since we covered virtually all the
windward beaches (Pt. I, Fig. 9), we are confident that fewer than 20
pairs nested on the atoll. Our population estimate for 1990 was 25
pairs.

The POBSP (Clapp & Sibley 197la) found 3 nests on Nake in June
1965, estimating a population of 15 birds, while the Grossmans (1974)
found 8 nests on Long Island, estimating a similar population. Even
though our surveys triple the known population, the Brown Booby remains
a rare seabird on Caroline.

15

Phenology: With the exception of one recently fledged juvenile, all
nests contained eggs in September 1988 (Table 3). Clapp & Sibley
(1971a) found eggs in June; the Grossmans found eggs in September. In
March 1990, the 20 pairs were all on nests whose contents ranged from
eggs to an older juvenile. However, 2 months later, only 3 nests,
containing eggs, could be found. These data from 4 years suggest that
the species may have trouble rearing young. More juveniles should have
been encountered, especially in May 1990. However, cyclonic weather in
February 1990 brought torrential rains and severe winds (Falconer, pers.
comm.) which defoliated and uprooted the strand vegetation of Long and
Nake (Pt. I, Pl. 33), deposited storm blocks on the windward reef flats
and tons of sand over the existing beaches and old interislet channels
of Long (AKK, pers. obs.). Brown Booby eggs and chicks would have
experienced great difficulty at this time, as the region hardest hit was
their sole nesting area. During February 1990, 640 mm (25.2") of rain
fell in 10 days (Pt. I, App. II). Predation by Polynesian rats or
coconut crabs could also limit reproduction on the atoll.

RED-FOOTED BOOBY (Su7a su7a) Figs. 6, 7

This pantropical booby numbers over 55,000 individuals in the Line
Group (Clapp 1967, Perry 1980), making it one of the most important
regions in the world for this species. Caroline holds the fifth
largest known Red-footed Booby colony (see Nelson 1978). The largest
known colony (140,000 pairs) is found on Tower Island (Galapagos):
three of the 5 biggest colonies occur in the Line Group.

Distribution and habitat preference: In 1988, the Red-footed Booby bred
on 28 islets, ranging in size from Nautonga (0.34 ha) to Nake

(107.46 ha) (Fig. 6). On the Windward Islands, Red-foots occurred from
Nake to Tridacna, absent only from the smallest islets (Noddy Rock,
Skul] Islet, Motu Atibu). The species was also widespread on the
leeward islets, extending from Pandanus to Eitei. The tiny islets
(Fishball, Azure, Reef-flat) were not occupied.

Red-foots are tree nesters whose distribution on Caroline closely
matched that of Tournefortia scrub and forest. They sometimes utilized
smaller Pisonia or Cordia trees where they intermingled with
Tournefortia, and occasionally built nests in the tallest (>15 m)
Pisonia. They nested in smaller Tournefortia patches within the
peripheral scrublands, especially those not directly exposed to the
trade winds. They clearly avoided smaller islets because of the lack of
suitable Tournefortia in which to breed. They nested inward from the
vegetated edges of the islets, generally at 3-6 m in height, and were
distributed in broken rings around the smaller motus in areas of
moderate winds. A higher percentage of the population occurred on
perimeter surveys than on cross-island transects.

Red-foots were absent from South Island, which was primarily
covered with Cocos (Pt. I, Figs. 50, 51). Even though Journefortia
occurred on all its coastlines, no boobies nested in them. Ana-Ana was
also unoccupied: the presence of a family of 4 people, a cat, and a dog
(all removed in 1991) undoubtedly discouraged nesting attempts. Red

16

RED- FOOTED BOOBY

CAROLINE ATOLL

SCALE 1:34,000
soo ° 500 1000 2000

Le A AY Ga a

meters

Figure 6. Distribution map of breeding Red-footed Boobies on Caroline
Atoll, September 1988.

17

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18

foots also avoided the mixed forests of south Nake, which contained much
Cocos and Pandanus (Pt. I, Fig. 37). Red-footed Boobies were thus found
only in Caroline’s indigenous woodlands, primarily -in Tournefortia >2 m
tall; they avoided anthropogenic plant communities and man.

Red-foots used a wider range of habitats for roosting.
Nonbreeding birds were found throughout the taller indigenous trees,
even in leeward situations where Pisonia and Cordia overhung the lagoon
(as on Long Island).

Numbers: The POBSP (Clapp & Sibley 197la) estimated 5,000 + 25% Red-
foots on Caroline in June 1965, with about 2,000 + 25% nesting pairs.
The Grossmans estimate was 3,000 + 25% and 2,500 pairs in September
1974. In 1988 we sampled systematically more than 7% of the available
habitat on all motus except Crescent (4.6% sampled) and North Arundel,
and estimated that 2,221 pairs of Red-footed Boobies nested on 27 of
Caroline’s islets (Table 1). We found an additional 1,234 roosting,
nonbreeding birds. We know (C. Kepler 1969, Nelson 1978) that fewer
boobies remain in their colonies during the day than at night. Thus an
unknown fraction of the population was at sea when we conducted our
counts. Impressive flights of Red-footed Boobies returned each evening:
3-4 birds arrived for each one that had remained behind, many
undoubtedly mates of incubating birds. To approximate the number of
returning nonbreeding birds, we doubled the number of roosting adults to
allow for an additional 1,234 adults and juveniles. Thus, our
conservative estimate was at least 7,000 individuals.

Because Red-footed Boobies were so dependent upon Journefortia, we
determined the nesting population on each islet by multiplying the
number of nests found on transects by the ratio of sampled to total
Tournefortia area. Perimeter counts (Pt. I, Fig. 9) were used if the
number of Red-foots observed exceeded the number calculated from the
cross-island transects.

Long Island held the greatest number of nests (659), mostly in the
leeward Journefortia and Tournefortia-Pisonia edge. Bird densities were
typically highest on the largest islets: Windward and Tridacna, the
largest Windward Islets, held 163 and 111 nests, respectively; and
Mannikiba, the biggest leeward islet, harbored the largest population
(184) of the entire leeward side. There were exceptions, however:
Pandanus, with 4 times the area of Journefortia of any of the South Nake
Islets, held fewer birds than 3 much smaller islets (Table 1).

Tournefortia scrub and forest covered approximately 125.25 ha
(Pie all, Table,9) . Overall, there were 1.75 Red-footed Booby
nests/1,000 m° of Tournefortia forest. Nest densities for occupied
islets by island groups (Table 4) showed that Red-foots favored areas
less exposed to the trade winds: most nests on the windward motus were
protected by well-developed Pisonia forests. The exposed Centra
Leeward Islets held the lowest nest densities (1.2 nests/1,000 m*), far
less than on the South Nake Islets (5.3/1,000 m“), which are protected
by the northern edge of Long. The greatest densities (7.8
nests/1,000 m°) occurred on the South Nake Islets south of Pandanus.

19

Table 4. Density of Red-footed Booby nests in occupied Journefortia
habitats on islet groups, Caroline Atoll, September 1988.

Number Estimated Area of Nests/1,000 me
Occupied Number foucpeponet® of Available
Islet Grou Islets Nests m Habitat
Nake 1 496 300,650 16
Long i 659 322,000 2.0
Windward Islets 8 434 251,900 ea
South Nake Islets if 319 59,800 BS
Central Leewards 6 239 197,500 eZ
Southern Leewards 4 74 39,600 19
Total Gi ence 1% 17051550 1.9

Broadly speaking, Red-foots brged in well-dispersed colonies. A
record density of 600 nests/1,000 m° on Trome] in Island (Indian Ocean)
is exceptional. Elsewhgre, 53 pairs/1,000 m° on Tower Island 9
(Galapagos), 40/1,000 m“ on Moku Manu (Oahu, Hawaii), and 27/1,000 m* on
Half Moon Cay (Honduras) are more consistent high-density colonies
(Nelson 1978) . Only on tiny Motu Kota (Pts eekig. 52)5 swith. !2onests
in 303 m° of Tournefortia (40/1,000 m“), did we find such density, and
for this reason we named the islet "Kota" (Gilbertese for Red-footed
Booby).

Phenology: In September 1988, we located 339 nests. Of the 152 whose
contents could be determined, 87 were empty, 63 contained eggs, and 2
held downy chicks. We saw dozens of flying juveniles along the windward
coasts. Most pairs were building or guarding their nests during a
prelaying stage that lasts from 11-35 days (Nelson 1969). Of the pairs
with nests, 57.2% had yet to lay and 41.4% had laid their eggs between
mid-August and late September (Fig. 7). Applied to the total breeding
population, approximately 1,270 nests were in the prelaying stage and
would be expected to produce eggs throughout October. An additional 919
nests had a mean laying date in early September (Fig. 7). Red-footed
Boobies were synchronous with Brown Boobies but delayed relative to
Masked Boobies.

In June 1965, nests containing prelaying adults, eggs, and young
in all stages indicated that the birds were in the midst of a protracted
breeding season extending from January to June. In September 1974, eggs
and "young at nearly all stages" were present. Our data reveal that no
successful nesting occurred in May-June 1988. Data from March and May
1990 indicate that nest-building began in January (or earlier), with
eggs laid from January to May. However, perimeter counts and cross-
island transects on 11 windward motus found virtually all Red-footed
Boobies either nest-building or sitting on eggs. Chicks were found only
on the leeward islands. This asynchronous breeding suggests that

20

breeding activities were curtailed by the cyclonic weather two weeks
earlier, which was particularly violent on the windward beaches
harboring Brown Booby nests. Red-footed Boobies in other tropical
locations have variable, opportunistic breeding seasons that depend upon
food availability (Nelson 1978; F. Sibley, pers. comm.); our data
suggests that similar pressures could be operating at Caroline.

Color morphs: Red-footed Boobies are polymorphic (Nelson 1978). The
basic plumages are brown or white, with brown morphs having many
combinations of tail, back, scapular, foot, and bill colors. A variety
of brown forms and white forms occurred on Caroline, with a ratio of 9:1
(337 brown to 35 white), which contrasts sharply with Nelson’s (1978)
statement that "in the Line and Phoenix Islands all birds are white
morphs." Most of the dark morphs were the "white-tailed" form (see
Nelson 1978, pp. 660-661). The variations and proportions of plumage
types show clinal change in the Line and Phoenix Islands (F. Sibley,
pers. comm.), thus the question of plumage morphology needs much more
study in the Central Pacific.

GREAT FRIGATEBIRD (Fregata minor) Figs: 8,9 anduPtoel maples

The Great Frigatebird breeds at widely scattered locations
throughout tropical waters in the Atlantic, Pacific, and Indian Oceans.
It is known to breed on all of the Line Islands except Starbuck (Perry
1980).

Distribution and habitat preference: Great Frigatebirds nested on 25
islets, including Nake, Long, and most of the larger islets (Fig. 8,

Table 1), ranging in size from Azure (0.20 ha) to Nake (107.46 ha).
Every occupied islet had some Pisonia forest, even if only a single tree
(Azure). The larger islets lacking Pisonia forest (Arundel, 7.34 ha,
Tridacna, 9.08 ha) lacked frigatebirds in 1988, although frigatebird
chicks were present on Arundel in early 1989 (Anne Falconer, pers.
comm) .

Although Great Frigatebirds were similar in nest requirements to
Red-footed Boobies, there were significant differences: the frigates
tended to nest higher in, and closer to the outer edge, of the canopy
(although nests were found as low as 1.3 m). Nest sites were more
sheltered from the wind than those of Red-foots, and in locations where
the birds could take flight easily. Such site preferences may explain
the association with Pisonia. Pisonia reaches 21 m on Caroline, taller
than other tree species, providing a windbreak on most islets. The
largest colonies (Nake, Long, Pig, Mannikiba) were found leeward of
these stands. We found nests in Tournefortia, Pisonia, and Cordia.
They were often in the Tournefortia-Pisonia interface, generally in the
taller Tournefortia. One colony on south Long overhung the lagoon in a
dense Pisonia stand. Frigates were not found in any anthropogenic
forests and were absent from then-inhabited Ana-Ana.

Numbers: The previous population estimate for Great Frigatebirds on
Caroline was 10,000 birds (Clapp & Sibley 197la, Perry 1980). The
Grossmans (1974) estimated 5,000-8,000 birds, with 4,000-6,000 nests.

B® GREAT FRIGATEBIRD “)

> LESSER FRIGATEBIRD
1988, 90

CAROLINE ATOLL

SCALE 1:34,000
seo ° 800 1000 2000

meters

Figure 8. Distribution map of breeding Great and Lesser Frigatebirds
on Caroline Atoll, September 1988.

21

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23

We calculated that 2,427 pairs bred or attended territories. An
additional 617 birds roosted, thus the entire population was
approximately 5,471 individuals. A large but undetermined number of
birds soared over the atoll throughout the day, and an uncountable
number of birds, including fledged juveniles that would ultimately
return to the island to nest (Diamond 1971), were undoubtedly at sea.
Because this species is difficult to count accurately, it is unclear if
the population has changed since 1965.

Phenology: In frigatebirds, the scapulars, which first appear at 81]
days in Fregata magnificens (Diamond 1973), erupt before the primaries.
Because we lack chick stage data for F. minor and F. ariel, we have
modified ages from Diamond (1973) for magnificens, using the hatching
times for F. arvel and F. minor from Nelson (1976), and fledging ages
from Diamond (1975b), to construct very approximate development stages
for the species on Caroline. Since they fledge at an earlier age than
F. magnificens, we have reduced the ages for chicks with erupting
primaries for F. ariel] and F. minor, kept the duration of the earlier
stages approximately the same, and reduced the period in juvenile
plumage.

We found 214 nests in 1988. Of the 144 in which we determined
contents, 49 contained eggs or young chicks, 27 held chicks with
developing scapular feathers, and 68 contained older chicks (Table 5).
The additional 70 adults occupied nests of unknown contents. We saw
fewer than 10 displaying males and a high proportion (87%) of nests with
chicks, many of them old, indicating that the breeding season was nearly
over. A major laying effort had begun in March-April (Fig. 9) and
continued into September. In March 1990, an abundance of flying
juveniles and occasional larger chicks down to the downy stage indicated
that the previous year’s breeding season was ending. Of all the seabird
species affected by the February 1990 storm, Great Frigatebirds suffered

Table 5. Stages in the breeding cycle of frigatebirds on Caroline
Atoll, 21-29 September 1988.

Nest stage/Approximate age in days from laying

Species Juv. Primaries Scaps. Downy Naked Eqgs
Great 191-220 101-190 81-100 56-80! 0-55
Lesser 181-210 91-180 71-90 56-70 46-55 0-45

No. nests in each stage

Great 22 46 27 30! 19
Lesser 4 13 4 5 0 0

1 duration of naked and downy chick stages are lumped because it was
often impossible to see into canopy nests.

24

the most obvious mortality. We found at least 10 adults and flying
immatures recently dead, either draped in partly defoliated Journefortia
shrubs or lying on the ground. A small number of males were beginning
another courtship cycle. By May 1990, courtship and egg-laying were
still underway, and nests contained eggs or small chicks up to the
"remiges" stage. Peak laying on Christmas Island (Pacific Ocean) occurs
from March-May (Schreiber & Ashmole 1970), the same laying cycle
observed on Caroline in 1988 and 1990.

LESSER FRIGATEBIRD (Fregata ar7e/) Figs. 8, 10

The Lesser Frigatebird is a pantropical species. It breeds and
disperses widely within the tropical Pacific (Sibley & Clapp 1967). One
of the largest populations in the world (30,000-85,000) breeds on McKean
Island, in the Phoenix Group (Garnett 1983). Lesser Frigatebirds breed
on 4 of the Line Islands, with the population on Malden (7,000) the
largest in the archipelago (Perry 1980).

Distribution and habitat preference: In June 1964, Lesser Frigatebirds
were found nesting in one compact colony on the leeward north end of
Long (Clapp & Sibley 197la). In September 1974, adults and flying
immatures were observed flying and resting (Grossman & Grossman 1974),
but no nests were found, most likely because the western side of the
atoll was not surveyed. A population of 200 was estimated. We found a
single colony in leeward Pisonia forest on western Nake (Fig. 8), both
in September 1988 and May 1990. The birds nested high (to 18 m) in the
Pisonia and Pisonia-Cordia edge facing an open Journefortia savannah.
Although primarily composed of F. arje], a few F. minor were scattered
along all but the eastern edge of the colony. West of the birds, across
the open forest, F. minor and Sula sula nested in a mixed colony in a
denser stand of Tournefortia. Birds were seen soaring over Nake, Long,
and the leeward islets but were not found roosting or nesting away from
the colony on Nake. However, in March 1990, approximately 650 Lesser
Frigatebirds were swarming above and roosting on Motu Nautonga in a
tight cluster, possibly preparing for nesting.

Numbers and phenology: POBSP biologists estimated a population of 1,000
Lesser Frigatebirds on Caroline in June 1964, with 400 + 10% breeding:
only eggs were found (Clapp & Sibley 197la). On Christmas Island,

F. ariel laid in May and June in 1959, 1963, 1964, and 1967 (Schreiber &
Ashmole 1970). Of 46 nests found in 1988, we could inspect the contents
of only 26: all contained feathered chicks (Table 5). Laying dates
ranged from March through July (Fig. 10), with a peak from April to
June. Our limited data on Caroline’s Lesser Frigates indicates,
therefore, that they may be synchronous with those on Christmas. In
larger Central Pacific colonies, large subpopulations of this species
have differing breeding regimes (F. Sibley, pers. comm.).

Because we did not determine the colony limits, we cannot provide
a population estimate. There was a minimum of 200 birds in 1988 (46
nests, plus roosting and flying individuals) and ca. 1,000 in 1990.

25

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26
SOOTY TERN (Sterna fuscata) Figs ell; 12

This tern is the most widespread and abundant tropical seabird in
the world. Under favorable conditions it forms immense colonies
numbering into the millions. It is known to breed on 7 of the Line
Islands: the largest population in the Pacific is found on Christmas
Island (15,000,000 at highest count), and 3,000,000 have been recorded
on Starbuck (Perry 1980).

Distribution and habitat preference: To date, 21 colonies from 10
islets are known for the years 1965, 1974, 1988, 1989, and 1990

(Fig. 11). In September 1988, we found 3 colonies, 2 on the northern
half of Long and one on Bo’sun Bird Islet; all fit the general habitat
description in Clapp & Sibley (197la). Colony A, nearly square, was

210 m on a side. Eggs were placed under a savannah-type Tournefortia
scrub, from 1-4 m tall with approximately 60% canopy cover. The
substrate was coral rubble mixed with sand, covered by Heliotropium
(5%), Portulaca (1%), Laportea (<1%), and Lepturus (<1%), typical of old
interisland channels. Colony 1 was located in a broad sandy corridor
with 2 large "groves" of Tournefortia. The northern subpopulation
extended 116 m along the windward beach, but 248 m along the lagoon.

The southern subpopulation began 28 m further south along the beach,
fronted the seaward reef for 86 m, and was shaped like a blunt triangle,
its apex pointing toward the lagoon. Most chicks were under
Tournefortia, which consisted of shrubs 2-4 m high with 80% canopy
cover. The substrate was also older beach sands mixed with coral
rubble, and covered with Portulaca (40% cover), Lepturus (<5%), and
Heliotropium (<5%). The Bo’sun Bird colony, a rough oval approximately
55 m wide by 70 m long, was under 2-3 m high Tournefortia with 75%
cover, on coral rubble/sand sparsely carpeted with Portulaca and
Heliotropium.

Numbers: Populations were determined by measuring colony dimensions,
then counting eggs and/or chicks in 9 m° sample plots located at random
points along a compass line. Because juveniles moved as we approached,
they were counted 6 m ahead of us in estimated 3 m x 6m plots. The
Colony 1 subcolonies (North, South) were treated separately.

Colony size (rounded) in 1988 ranged from 127,000 + 30,000 "nests"
(Colony A) to 1,500 + 750 new eggs on Bo’sun Bird Islet (Table 6).
There were an additional 6,900 + 1,600 nearly-fledged chicks in the
Bo’sun Bird colony, resulting from eggs laid 3 months earlier.

The total number of eggs and chicks was 188,000 + 40,000. Actual
numbers of adults are difficult to estimate, but in other studies have
exceeded the number of eggs and young by factors of more than 2 because
innumerable eggs and chicks were lost, colonies often overlapped, and
many nonbreeding adults joined the prebreeding swarms or associated with
breeding birds. Schreiber & Ashmole (1970), relying on POBSP data from
Johnston Atoll (North-central Pacific), estimated that 4 adults were
present for each egg laid. POBSP data from Johnston (Amerson & Shelton
1976) indicated that about 600,000 adults were present in a colony with
105,000 eggs, or approximately 5.7 adults/egg. If we assume that real

rail

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CAROLINE ATOLL

SCALE 1:34,000

aN juveniles

N number of colonies

meters

Figure 11. Distribution map of breeding Sooty Terns on Caroline Atoll,
September 1988 to July 1990.

28

>
SOOTY TERN 1988 (=
SS

180

LONG IS.
“a”

125

100

75

LONG IS.
div

NO. EGGS LAID |x1000)

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JULY AUG. SEPT.

Figure 12. Approximate laying dates for Sooty Tern young found on
Caroline Atoll in September 1988. See Fig. 3 for explanation.

29

Table 6. Sooty Tern colonies on Caroline Atoll, 27-28 September 1988.

Calculated
Population Nest Approx. Weeks
Colony Location Area Mean Pairs + SE Stage From Layin
Long Island, A 44,100 me 127,449 + 30,429 hatching eggs, 4-5
downy chicks
Long Island, 1 N 24,200 me 41,382 + 5,808 chicks with 7-10
short tails,
juv. plumage
Long Island, 1 S 6,400 m2 10,944 + 1,536 ‘ 7-10
Bo’sun Bird Islet, 9
old REY A) m5 6,883 + 1,575 fledglings 11-12
new 3,375 m 1 538ut) 758 new eggs 1-2
Total 75,075 m2 188,196 + 40,106

numbers of terns in our colonies lay midway between 4:and 5.7 times the
number of eggs and chicks, then the number of Sooty Terns using Caroline
Atoll would have ranged between 720,000 and 1,100,000 birds (911,800 +
21%). This is twice the estimate provided by Clapp & Sibley (1971a),
even though we found fewer colonies. However, if the POBSP had used the
criterion of 4 adults present for each egg laid, their total population
figures would have exceeded ours, hence the 2 estimates are not strictly
comparable (F. Sibley, pers. comm.).

In March 1990, laying was just beginning in 2 colonies on Long
Island, (625 m x 150 - 315 m wide and 180 m long x 160 m wide).
Enormous numbers of birds, both on the ground in densities up to 9 or 10
pairs/m” and in the air, made it impossible to calculate a reasonable
population figure. According to Anne Falconer, these 2 colonies were
very successful. Similarly, counting was difficult in May 1990 when 6
large prebreeding swirls hovered like huge clouds of gnats over discrete
islets and islet groups (Fig. 11). Our 1988 estimate of approximately
one million birds is probably a conservative count for the atoll as a
whole on an annual basis.

Phenology: The incubation period in Sooty Terns is about 4 weeks
(Dinsmore 1972). Young fledge 7-8 weeks after hatching, although
fledging ages, dependent upon food supply (Schreiber & Ashmole 1970),
are highly variable.

Four separate Sooty Tern colonies had been started over the 12-
week period prior to our study in 1988 (Table 6). On Bo’sun Bird Islet
a new wave of laying was just beginning in an open area immediately
southwest of most of the colony, while nearly fledged chicks scurried
about beneath the Tournefortia. Undoubtedly many young had already
fledged, so many more eggs would have been laid in early July by this
colony than indicated (Fig. 12). The 2 colonies on Long were
established at different times: the short-tailed juveniles in Colony 1
preceded the large number of eggs, hatching eggs, and downy chicks of
Colony A by 3-4 weeks.

30

The July-September laying period on Caroline in 1988 is very
different from the bimodal breeding (May-June, December-January)
reported from Christmas Island, Pacific Ocean (Schreiber & Ashmole
1970), and the May laying dates noted for Caroline by the POBSP in 1965.
In September 1974, "innumerable large unfledged chicks and juveniles"
were present in colonies at the tip of Long Island and on Tridacna Islet
(Grossman & Grossman 1974). Additional data (Anne Falconer, pers.
comm.) indicate that Sooty Terns may lay any time (Fig. 11), certainly
January through September (1988 to 1990). Severe storms, which
destroyed large Long Island colonies in February 1990, were likely
responsible for reinitiating breeding activities on the leeward side of
the atoll within the next few months. A great deal more research will
be needed on Caroline before the breeding seasons for this species are
fully understood. The phenomenon of several colonies on each island
breeding at different times is considered the norm for the Line and
Phoenix Groups (F. Sibley, pers. comm.), so the situation on Caroline is
not unusual.

BROWN NODDY (Anous stolidus) Fig@, 13

This tern, primarily a tree nester, is widely distributed
throughout the warm oceans of the world. It is abundant in the Line and
Phoenix Groups, with an estimated total population exceeding 40,000
birds. Brown Noddies are most abundant on Palmyra Island (10,000
birds).

Distribution and habitat preference: The Brown Noddy is second only to
the White Tern in the number of motus (28) upon which it is known to

breed (Fig. 13). It utilized the smallest (Noddy Rock, 0.02 ha) and
largest (South, 104.41 ha) motus, nesting upon coral rubble and in plant
communities ranging from the simplest herb mats to Journefortia,
Pisonia, Cordia, Cocos, and the mixed anthropogenic forests of South and
Nake. Most pairs were well dispersed, nesting from the outer edges of
Tournefortia to the central, inner branches of Pisonia, and from the
ground to the crowns of 25-m Cocos. When nesting sympatrically with
Black Noddies in Pisonia, the Brown Noddies typically occupied portions
of branches closest to the trunk. Brown Noddies nested almost
solitarily in the Cocos canopy on South, were found within dense
colonies of Black Noddies and White Terns in tall Pisonia forests, with
Red-footed Boobies and Great Frigatebirds in Tournefortia, and amidst
Sooty Terns and Red-tailed Tropicbirds (Bo’sun Bird Islet). Apart from
a few ground nesters on Raurau and Fishball, the only ground-nesting
colony (80 nests) was located on a Portulaca mat on Noddy Rock--a site
free of predators, although flooded during storms.

Brown Noddies often formed loose roosting "clubs" on the atoll’s
beaches. Aggregations of 15-20 birds were found on the west coast of
South and on Sandy Inlet, south-central Nake.

Numbers: Clapp & Sibley (197la) estimated a population of 1,000 birds
in June 1965, with about 800 birds breeding (with eggs and young). The
Grossmans estimated 400 + 15% in September 1974. We estimated a total
population of 1,491 breeding pairs (Table 1). Because nests high in

—_~
=a) \

BROWN NODDY ee

CAROLINE ATOLL

SCALE 1:34,000
S00 ty) S00 1000 2000

Ce SS SS eee

meters

Figure 13. Distribution map of breeding Brown Noddies on Caroline
Atoll, September 1988.

31

32

Cocos palms were difficult to detect, we undoubtedly overlooked many,
thus our estimate of approximately 3,000 birds is conservative.

Although larger than the population estimated by POBSP (Clapp & Sibley
1971a), uncertainties about the 1965 survey coverage (F. Sibley, pers.
comm.) prevent us from knowing if Caroline’s population has changed over
the past 25 years.

Phenology: This is another species whose breeding cycle is irregular in
the Central Pacific: during the 1963-1965 POBSP surveys, egg-laying was
found in every month of the year somewhere in the Line and Phoenix
Groups (F. Sibley, pers. comm.). On Christmas Island, which has
received more attention than any other Central Pacific island, the
timing of egg laying varies between colonies. In general, peak laying
occurs from March to May, and from November to December. On Caroline,
mating and nest-building were found in March 1990, but by May only a few
eggs had been laid. Eggs and young were found in June 1965 (Clapp &
Sibley 197la) and in September 1988 (present study). Only eggs were
present in September 1974. We found 246 nests in September 1988 and
determined the contents of 106: 103 held eggs, 3 held downy chicks.

The incubation period is 35-37 days (Dorward & Ashmole 1963), so all
viable eggs had been laid within the previous 40 days--mid-August to
late September. Because many nests were being built, we feel confident
that laying continued into October. Clearly more research is needed to
determine whether laying occurs in regular cycles.

BLACK NODDY (Anous minutus) Fig. 24

The Black Noddy is widely distributed in the tropical Atlantic and
Pacific. It is abundant in the Line and Phoenix Groups, with
populations of 16,000 estimated in the Phoenix Islands (Clapp 1967) and
over 46,000 in the Line Group. Centers of abundance are Palmyra
(20,000) and Christmas (14,500) (Perry 1980).

Distribution and habitat preference: The Black Noddy is a tree-nesting
species that on Caroline prefers tall stands of Pisonia. The largest

colonies (61% of the population) were found in the grand Pisonia forests
(to 25 m) on Pig and North Pig. We found breeding birds on 18 motus,
with populations exceeding 800 pairs in the Pisonia on Nake, Long, North
Pig, and Pig (Fig. 14). The only significant colony not primarily
associated with Pisonia was found on Tridacna, where approximately 230
pairs nested in the tallest (ca. 8 m), most central Tournefortia-Morinda
forest. Black Noddies always nested in dense colonies near islet
centers and were integral components of these plant communities: their
droppings, coating the ground with a film of guano, constantly enriched
the islet’s meager soils.

Numbers: Clapp & Sibley (197la) estimated that 7,000 + 25% birds were
on Caroline. The Grossmans (1974) estimated 500 + 20%. During our
visit the population was much larger: 5,122 pairs were estimated for
Pig and North Pig alone (Table 1). Basing our numbers primarily on the
densities of sampled colonies in Pisonia, we estimated that nearly 8,400
pairs were nesting during our 1988 visit. Our population estimate
approached 17,000 birds, to which an unknown number of nonbreeding birds

Figure 14.

—_
=~

BLACK NODDY ]

CAROLINE ATOLL

SCALE 1:34,000

soo ° 300 1000 2000

iin Ge Le Ey

meters

Distribution map of breeding Black Noddies on Caroline
Atoll, September 1988.

33

34

could be added. These values place the Caroline population far above
that for Christmas, making it the largest known population in Kiribati.

Phenology: Black Noddies were just beginning a new breeding season, as
was also found in September 1974. On 27 September we observed hundreds
of birds gathering Tournefortia leaves floating along the windward shore
(Long) or flying with fresh leaves to their nests (Pig, North Pig). Of
the 1,085 pairs counted on transect, 536 (49%) perched as pairs, were
defending nest sites, or were building nests. An additional 273 pairs
were attending nearly-completed nests but were not incubating. The
remaining 276 pairs were incubating, thus 75% of the pairs had not laid
eggs. The contents of 230 nests were unknown, although we assumed they
contained eggs because of the incubating positions of the adults. Of 46
nests whose contents were visible, 45 held a single egg, and one
contained a downy chick less than 5 days old.

The breeding seasons for Black Noddies on Christmas Island and
Johnston Atoll peak in April and May (Schreiber & Ashmole 1970, Amerson
& Shelton 1976), where pairs are highly synchronous, laying most of
their eggs within a 2-3 month period. The Caroline colony, also
synchronous, but beginning egg-production in September, would be
expected to peak in October/November, 6 months out of phase with the
colonies further north. In 1990, however, Black Noddies were just
beginning to mate and nest in March, and by May some were still sitting
tightly on nests, while others had chicks in all stages. This was most
likely the result of the stormy weather which affected all seabirds.

BLUE-GRAY NODDY (Procelsterna cerulea)

Blue-gray Noddies nest widely across the Pacific from the Kermadec
Islands to Hawaii. They are scattered throughout the Line and Phoenix
Groups. In the Line Islands, they were formerly known to breed only on
Christmas and Malden (Perry 1980). Eggs are placed in nests minimally
provided with twigs and may be on coral rubble, sheltered under
vegetation, or under coral slabs to depths of one meter (Rauzon et al.
1984).

The Blue-gray Noddy was recorded as "present" on Caroline by Perry
(1980). Clapp & Sibley (197la) noted birds over the lagoon but saw none
on land. When we approached Caroline, we saw 2 from the ship and later
observed 3 flying across the lagoon. We also saw 3 birds perched on
reef flats of the leeward motus Nautonga and Eitei. A third bird
flushed repeatedly from a small clearing around a pile of bottles on
Raurau, but we failed to find a nest. In March and May 1990, we
observed Blue-gray Noddies on all of the Southern Leewards, plus Azure
and Nautonga in the Central Leewards.

In summer 1990, Alexandre Falconer, resident on Caroline at that
time, found one small chick, attended by its parents, on an open expanse
of coral rubble on Motu Eitei, the first breeding record for Caroline.
Eitei is adjacent to Raurau, which we predicted was the most likely
breeding location for this species.

35

Blue-gray Noddies must breed in very small numbers on Caroline.
Nests are hard to find, given their cryptic placement, the small number
of birds present, and the extent of open habitat (67.7 ha of herb mats
and 41.4 ha of consolidated coral rubble). F. Sibley (pers. comm.)
suggests that Blue-gray Noddies are extremely vulnerable to predators,
since they were observed on 12 of 20 islands visited in the Line and
Phoenix Groups, but nested on only three. Where they did nest, Blue-
gray Noddy colonies contained hundreds or thousands of birds.

WHITE TERN (Gygis alba) Figie 1S yaP le. 3

The White Tern is a widely-distributed pantropical species
occurring in moderate numbers throughout the Line and Phoenix Groups.
Perry (1980) estimated 17,050 birds for the Line Islands, and Clapp
(1967) estimated 10,000 birds in the Phoenix Group.

Distribution and habitat preference: White Terns, the most widely
distributed breeding bird on Caroline, nested on 32 of the 39 motus

(Fig. 15), avoiding only those which were tiny and sparsely vegetated.

White Terns nested from one to 15 m above ground, wherever a
branch or frond provided a relatively stable platform in Journefortia
(Pl. 3), Pisonia, Cordia, Pandanus (Pt. I, Pl. 35), or Cocos. They did
not form dense colonies but were scattered throughout each motu,
normally selecting sites sheltered from the prevailing trade winds.
They utilized isolated trees, scrub, or forest. An unusual departure
from the White Tern’s usual mode of "nesting" was an egg laid in an old
Black Noddy nest, 6 m up in an 8-m-tall Journefortia on Tridacna Islet.

White Tern densities varied from islet to islet Stable 7). At one
extreme we found only 2 nests on Raurau (0.07/1,000 m°~). Densities on

Table 7. Density of White Terns on occupied islets by islet group,
Caroline Atoll, September 1988.

# White Density

# Vegetated Tern (pairs
Islet Grou Islets Area (ha Pairs 1,000 m
Nake 1 66.63 1,094 1.64
Long ] 49.60 751 ae il
South 1 86.10 381 0.43
Windward Islets 9 36.09 1,164 BR28
South Nake Islets 6 8.50 122 1.44
Central Leeward Islets 9 33.56 408 122
Southern Leeward Islets 4 6.47 37 O}.5i7

All Occupied Islets 31 286.88 395i) 1338

36

WHITE TERN (—. )

CAROLINE ATOLL

SCALE 1:34,000
soo ° $00 1000 2000

meters

Figure 15. Distribution map of breeding White Terns on Caroline Atoll,
September 1988.

37

other islets ranged from 0.75/1,000 me (Shark) ton 6. 7/1, 000 mé

(Nautonga) with a mean density of 1.38 pairs/1,000 m° of woodland.
Overall, the lushly vegetated Windward Islets supported the highest
densities. Although White Terns also nested in anthropogenic forests,
their densities were low: we believe that their low densities on South
Island and the Southern Leewards (Table 7) are attributable to man. Of
South’s 104.47 ha of vegetated land, only 4.2 ha (4.4%) was native
woodland (Pt. I, Fig. 50); fully 84% was either Cocos (18.3 ha) or dying
Cocos-Ipomoea (62.5 ha) forest. Although most of the Southern Leewards
are covered in virgin forests, central Ana-Ana has been partly cleared
(0.21 ha) to accommodate thatched huts and a garden. The activities of
a family of 4, with a dog and cat (until October 1990), apparently
depressed the White Tern population on Ana-Ana and, perhaps, even on
nearby islets. We found no White Terns on Ana-Ana during our visit,
although the Falconers, who vacated the atoll in summer 1991, assured us
that they occasionally nested.

Numbers: We used the total woodland area of each islet, coupled with
transect data, to calculate bird populations (Table 1). More birds were
found on the largest islets except South Island (see above). We
estimated 1,094 pairs for Nake, 751 pairs for Long, and nearly 400 pairs
for Tridacna; these 3 islets accounted for over half the population (and
over half the indigenous woodlands). We estimated that 3,957 pairs bred
on Caroline, which doubles the numbers of Clapp & Sibley (197la) and
Perry (1980) and exceeds by 3,000 the largest population formerly known
for the Line Islands.

Phenology: Of 569 pairs of White Terns recorded on transect, 437 were
roosting without obvious signs of eggs or chicks, 107 were incubating,
and 25 had chicks (often adults were not present). Of the 25 chicks
recorded, 17 were downy, 7 retained extensive traces of down with
remiges, and one was almost ready to fly. Incubation takes about 36
days (Ashmole 1963); young may require from 40-96 days to fledge
(Gibson-Hill 1950, Ashmole 1968). Nearly all chicks were less than 4
weeks old.

On Christmas Island, Schreiber & Ashmole (1970) found that peak
laying occurred in April-August each year, with some laying in each
month. On Caroline, Clapp & Sibley (1971a) noted that about half of the
birds had eggs, half had young in June 1965. In September 1974, White
Terns had eggs only (Grossman & Grossman 1974). In March 1990, we found
very few eggs and downy chicks, but in May a larger number of pairs were
breeding, with eggs and chicks in all stages. Again, the February storm
had most likely interrupted breeding activities, as only 2 juveniles
were found on the windward side. These were in the interior Pisonia
forests of Brothers and North Brothers Islets, which suffered less
damage than motus further north. Therefore, although White Terns on
Caroline do lay during the peak period on Christmas Island, laying
appears to be heaviest after mid-August.

38
D. OTHER BIRDS ON CAROLINE ATOLL

Seven species other than seabirds have now been recorded on
Caroline. Six of them are migrants (5 shorebirds and Long-tailed
Cuckoo). The few shorebirds encountered (except for Bristle-thighed
Curlews) and their lack of increased numbers in the fall suggest that
there is only a small migration to Caroline. The Reef Heron is
apparently resident, although no nest has been found.

REEF HERON (Egretta sacra)

We found 15 Reef Herons scattered on 8 islands: Nake (1), Long
(2), Pig (1), Brothers (3), South (2), Mannikiba (2), Matawa (1), and
Emerald (2), as well as on the open reef flats (1). Although birds were
found on both the seaward and lagoonward sides of the islets, most were
along the lagoon edge, as also found by POBSP in 1965 (Clapp & Sibley
197la) and the Grossmans (1974). We estimated that approximately 30
birds were using the atoll. We found no signs of breeding. Of the 15
individuals we observed, 5 were dark, 8 were white, and 2 were of the
pied morph.

LESSER GOLDEN-PLOVER (Pluvialis dominica)

This plover used the beaches and herb mats, generally to seaward.
In September 1988, we found them on Nake (1), Long (4), Tridacna (4),
and Mannikiba (1), estimating a total population of 20-30 birds, the
same number found by POBSP (Clapp & Sibley 197la). In March 1990, we
observed 8, and in May, 3, all in winter plumage. In September 1974, 3
were seen on the windward coast (Grossman & Grossman 1974).

WANDERING/GRAY-TAILED TATTLER (Heteroscelus incanum or H. brevipes)

In September 1988, we located 18 tattlers on 6 different islets:
Nake (3), Long (3), Crescent (1), Arundel (2), South (7), and Emerald
(2). All birds were either alone or in pairs and generally remained in
the intertidal zone, although they often foraged on herb mats close to
the beach scrub. The total population was approximately 40 birds.
Those few birds heard were all H. incanum. We saw 6 tattlers in March
1990 and several in May of the same year. The Grossmans (September
1974) observed 12 on the windward coast and around the lagoonward shore
of the windward islets and South.

RUDDY TURNSTONE (Arenaria interpres)

One turnstone was found on the windward beach of Motu Mannikiba in
September 1988, and 5 on atoll beaches in March 1990. The Caroline
population probably does not exceed 15 birds.

BRISTLE-THIGHED CURLEW (Numenius tahitiensis) Pt. I, Pilesge

The Bristle-thighed Curlew, common in the Line and Phoenix Groups,
is a widespread migrant to atolls of the Central and South Pacific

39

during the boreal winter (Pratt et al. 1987). The species is considered
rare (Johnsgard 1981, Marks et al. 1990) and is a candidate for the U.S.
Fish and Wildlife Service Endangered Species List (Gill 1990). Clapp &
Sibley (197la) estimated 20 birds for Caroline Atoll in June 1965; the
Grossmans (1974) saw 4 on the windward coasts.

We counted 83 birds on 12 of Caroline’s islets in 1988, including
the 3 large islands (Nake, Long, South) and motus in the Windwards,
Central Leewards, and Southern Leewards. In
March 1990, we saw 20 curlews on 10 islets during incidental
observations throughout the atoll, bringing the total number of islets
on which they have been recorded to sixteen (41%). On our return trip
(May 1990) we only saw 3 curlews (8 motus visited). Undoubtedly,
curlews occur throughout, utilizing essentially all plant communities
(for details, see Pt. I, Sect. E). Although they are most conspicuous
on the beaches and reef flats, higher numbers may actually forage inland
during the day. Small numbers of curlews remain all year, being least
common from April to August and most abundant after September/October
(R. and Anne Falconer, pers. comm.). This correlates with preliminary
information from Rangiroa Atoll, Tuamotu Archipelago (Gill 1990; Gill &
Redmond, in prep.).

Unvegetated perimeter habitats: On a complete perimeter count of South
Island in 1988, we found 29 curlews. Twenty-one were foraging and
loitering on the windward east coast, principally above the beach crest
on coral rubble interspersed with herb mat. Similarly, 14 of 20 curlews
found on Long and the Windward Islets foraged along the windward beach
crest, with only 6 birds found on the lagoonward shores. Curlews were
equally common on windward and leeward shores in the leeward islets,
occupying habitats composed of coral rubble and sand. While the numbers
indicate that curlews preferred windward shores, they may be biased
because most birds were seen there in the late afternoon (13, 14, 19).
Perhaps they use the relatively open areas for roosting and foraging at
dusk. Our largest flock (14, Sandy Inlet, Nake) was found at 16:00,
foraging on compacted, silty sand at the lagoonward end of the inlet,
while single curlews dotted the interislet channels and shallow tidal
reef, fats \(Pts DP lin22):.

Vegetated habitats: We found Bristle-thighed Curlews on natural herb
mats, in Tournefortia scrub, Pisonia forest, and in Cocos habitats, both
in the healthy peripheral plantations and within the dying Cocos-Ipomoea
woodlands (Pt. I, Fig. 36, Pl. 34). One was captured in a mist net
under a dense Cocos canopy. Disintegrating plantations in the center of
South (54 ha) held a large population: calculated numbers produced an
estimate of 154 curlews. They foraged over the Ipomoea-strewn ground
(interspersed with Boerhavia and Phymatosorus), frequently using broken-
topped coconut trunks as lookouts. We also found 5 curlews on transects
in Pisonia forests up to 20 m tall on Nake (calculated population, 41).
They were foraging on the relatively open, although dimly lit, forest
FloomaCR mle PAli4i2))e:

Numbers: From the 1988 data we estimated a population of 300+ curlews:
41 birds in Pisonia, 154 in Cocos-Ipomoea, 43 on the beaches of South,

40

14 at Sandy Inlet, Nake, and 62 scattered over the remainding motus.
Because 154 were calculated from a single flock of 7 curlews on one
transect on South, there may be a bias in our population estimate.
However, this habitat covers 80 ha, 77% of South’s total area, and
incidental observations made off-transect indicated that curlews
commonly foraged in Cocos-Ipomoea woodlands. We believe that our
estimated density, about 1.5 birds/ha, is reasonably correct.

Bill length: Bristle-thighed Curlews show great variation in bill
length immediately after the breeding season. Because birds of the year
migrate south before their bills reach adult length (R. Gill, pers.
comm.), the ratio of "long" to "short" bills provides a rough estimate
of juvenile survival. Of
31 curlews seen in September, 20 were clearly adult length, 7 were
conspicuously shorter, and 4 were "intermediate" (probably young birds).
All March and May birds had long, adult-sized bills.

Some subadults remain on their Pacific wintering grounds for up to
3 years, during which time they pass through a flightless phase (Gill
1990, Marks et al. 1990). No flightless birds were seen.

Foraging: We saw one curlew chase and capture a small Polynesian rat at
dusk on the south shore of South Island. The bird bashed the rat on the
coral rubble, then ran rapidly about with the rat dangling from its
bill. After about 5 minutes, the bird swallowed the rat with vigorous
gulps.

Polynesian rats, abundant on Caroline (especially in Pisonia and
Cocos-dominated habitats), remain within the forest during the day, but
many move to the beach crest and tide line at dusk. They provide
abundant potential prey for curlews, which can easily capture them on
the open rubble. The synchronous appearance of rats and curlews at the
beach-woodland interface at dusk may be part of the foraging strategy of
this large shorebird. The presence of curlews beneath the forest canopy
may also be partly associated with this source of food.

SANDERLING (Crocethia alba)

One Sanderling in winter plumage was seen at water’s edge on the
windward beach of Long Island on 27 September 1988. Although
Sanderlings are well-known fall migrants in the Line and Phoenix Islands
(Clapp & Sibley 1967, 1968), this is the first record for Caroline
Atoll.

LONG-TAILED CUCKOO (Eudynamis taitensis) Fig. 16

The Long-tailed Cuckoo breeds in New Zealand and winters in the
southwest Pacific. The center of its winter range lies in central
Polynesia, but birds have been recorded as far as Palau in the northwest
and Pitcairn Island in the southeast. Although occurring throughout
French Polynesia and the Cook Islands, it had not been recorded from the
Line Islands prior to our expedition (Bogert 1937; Clapp & Sibley 197la,
1971b; Pratt et al. 1987; Ellis et al. 1990).

4l

LONG- TAILED CUCKOO

CAROLINE ATOLL

SCALE 1:34,000
500 ) $00 1000 2000

bh a Ee

meters

Figure 16. Preliminary distribution map of the Long-tailed Cuckoo on
Caroline Atoll. The species most likely utilizes all well-wooded motus.

42

We found Long-tailed Cuckoos on 4 of Caroline’s 39 motus
(Fig. 16). We heard its distinctive mono- and disyllabic call notes on
South, Long, and Pisonia, identified one on Nake, and on 28 September
collected a male in a mist net on Transect 4, Long Island (USNM 607191).
Soon after our return home we sent a description and photograph of this
species to the Falconers: they, and AKK, have since seen them several
times on Motu Ana-Ana in March, April, and May 1989-90.

All the cuckoo sightings were at canopy or subcanopy level, and 3
of the 4 1988 birds were in Pisonia forests. The individual on South
foraged in a Cocos canopy 21 m tall. The netted male flitted
secretively within an undisturbed, tangled low-canopy (4-6 m) Pisonia-
Tournefortia interface. We suspect that this elusive migrant occurs
throughout the mid-to-upper levels of Caroline’s forest canopy.

These records establish the Long-tailed Cuckoo as a winter visitor
to Caroline Atoll. Our observations on 4 motus in 8 days, including the
southernmost, northernmost, windwards, and leewards, indicate that many
individuals were present. A March 1990 first sighting on Vostok
(J. Phillips, pers. comm.) further suggests that the species disperses
regularly to the Southern Line Group.

E. OTHER VERTEBRATES

Lizards

Although "small lizards" were observed on Caroline in 1825
(Paulding 1931), it wasn’t until 1965 that the first collections were
made (Clapp & Sibley 197la). We collected 4 lizard species, increasing
the known terrestrial herpetofauna from 3 to 6 (Table 8). Although all
are indigenous, the azure-tailed skink (Emoia cyanura) is particularly
widespread in Oceania, and is suspected of being partly dispersed by man
(Brown 1956, Crombie & Steadman 1986). Recent data from Flint suggest
that E. cyanura may have been introduced by Tahitian copra laborers this
century (Kepler, in prep.). The same situation may be true on Caroline,
especially since this easily-indentifiable species was not seen or
collected by POBSP personnel in 1965 (Table 8). All but 2 of the lizard
species known from the Line Islands (Crombie 1990) have now been found
on Caroline. It is perhaps noteworthy that we added 3 species to the
atoll list but failed to find 2 species collected in 1965. Since both
groups collected lizards opportunistically, rather than systematically,
there is evidently much need for further study in this area.

Turtles

We found 3 Pacific green sea turtles (Chelonia mydas), a
threatened species (McKeown 1978), at Caroline in 1988. Two were
swimming over the lagoon reef flats, one west of Arundel, the second
east of Ana-Ana. The third was in the open sea about 100 m west of
South Island near the "boat entrance." Ron Falconer has seen up to 7

43

Table 8. Lizards collected on Caroline Atoll, 1965-1988.

Specimens:
Species Clapp & Sibley 1971a Present Study
Mourning gecko USNM 158355-57 USNM 299773
Lepidodactylus luqubris
Polynesian gecko USNM 158353-54 --
Gehyra oceanica
Snake-eyed skink -- USNM 299772
Cryptoblepharus poecilopleurus
Moth skink USNM 158358

Lipinia noctua!

-- USNM 299768-70
Emoia impar

Azure-tailed skink -- USNM 299771
Emoia cyanura

1 USNM 158358 has recently been reidentified by R. I. Crombie as Lipinia
noctua, not Emoia nigra, as reported in Clapp & Sibley (197Ia).

turtles in the lagoon in a single day. In April and May 1990, AKK saw
workers from Tahiti capture and kill a minimum of 4 green turtles in the
lagoon, and 2 more were in fishtraps on our departure. Two others
entered the lagoon during the following 4 months (R. Falconer, pers.
comm.).

In March 1990, AKK and G. Wragg found 3 old nests, presumably of
this species, on the northwest coast of Nake within 100 m of the
northern tip of the islet. These are the first known turtle breeding
records for the atoll. Young (ca. 1922) notes that the copra plantation
laborers ate green turtles from September to December each year, and
members of the Line Islands Expedition (Grossman & Grossman 1974,
Vickers 1974) saw fresh turtle tracks either on South or Nake. The
February 1990 storm added -large amounts of sand to Caroline’s
shorelines, providing potential new habitat for turtle nesting.

Terrestrial Mammals

None of the terriers (see Pt. I) that were introduced to control
rats on South Island early this century (Young ca. 1922) have survived
(F. Sibley, pers. comm.; R. Falconer, pers. comm; pers. obs.). In May
1990 the Falconers kept a dog and cat on Motu Ana-Ana. The dog
regularly visited all the Southern Leewards and accompanied the family
in their sailing canoe throughout the atoll. As a result of our

44

recommendations, the cat was removed from Caroline in October 1990. The
dog, with the family, vacated Caroline in summer 1991.

Bennett (1840) noted "rats of a red-brown color," the first
reference to rodents on Caroline. Dixon (1884) found that rats were
"not numerous" and that they nested "just at the base of the fronds" of
the coconuts. Two specimens collected by the POBSP proved to be Rattus
exulans (Clapp & Sibley 197la), an uncommon mammal restricted to South
Island.

The 19th and 20th century settlers found rats (presumably
R. exulans) to be extremely abundant and very destructive to the coconut
plantations; they contributing greatly to the twice-abandonment of copra
enterprises on Caroline (Young ca. 1922, Maude ca. 1938). They
voraciously devoured both growing and fallen nuts, as well as dried
copra. Being arboreal, they also lapped the juices of the flower
stalks, preventing nut development. In 1920 alone, over 4,600 were
trapped on South (Maude ca. 1938). Thousands more were killed by
terriers introduced in a vain attempt to control them.

We found rats on almost every islet, especially in or near coconut
palms and Pisonia trees. We recorded them during daylight hours on most
transects. Each night at our campsites on Long and South we noticed
groups of 10-20, so tame as to approach within one meter while we were
eating. Rats evidently undergo wide population fluctuations, as they
were less abundant in March and May 1990 than in September 1988.

We suspect that rats periodically reach most motus, and that those
apparently lacking rats (such as Noddy Rock) are too small and/or
depauperate to support a resident population. Because R. exulans is a
known seabird predator (C. Kepler 1967, Fleet 1972, Norman 1975), the
restriction of some species (i.e. Red-tailed Tropicbird) to small islets
may be due to rat populations on larger islets.

Rats were an abundant nuisance on Ana-Ana; the Falconers trapped
over 1,300 animals in 2 years and, like the pioneers before them, relied
upon a dog to help keep them at bay.

Marine Mammals

On March 14, 1990, members of the Line and Phoenix Islands
Expedition observed a minimum of 10 Pacific bottlenose dolphins
(Tursiops gilli) in the open sea about 500 m off the southeast corner of
South Island.

F. COCONUT CRABS

The coconut crab (Birgus Jatro, Coenobitidae), the largest
terrestrial invertebrate on earth, ranges throughout the tropical Indo-
Pacific (Pt. I, Pl. 21). It is highly esteemed as a source of food

45

throughout its range, and for this reason is rare or absent on or near
most inhabited islands. Because it is heavily exploited by man, it is
under consideration for endangered species status (E. Reese, pers.
comm.).

Since March 1990, dozens of Caroline’s coconut crabs have been
killed for food and for preservation in formalin as curios for the
Tahiti tourist market. Because of the increasing numbers of visitors to
Caroline, it is important that Caroline’s coconut crabs receive
protection.

History: A Californian malacologist, C. D. Voy, was the first to
collect coconut crabs (Birgus latro) on Caroline in 1875 (Pilsbry &
Vanatta 1905). They are not mentioned again until 1910, when Young (ca.
1922) wrote that "hundreds of great Coconut Crabs were seen: 40 large
ones were caught by the crew of the schooner in an hour" on South
Island. Young also noted that coconut crabs were considered a great
nuisance by plantation laborers, who killed them mercilessly. Evidently
they dug up newly planted nuts and snipped off emerging shoots. On
smaller motus, visited less frequently than South, Nake, and Long, these
depredations were difficult to control. Thus the small motu plantations
were abandoned soon after initial planting, resulting in a remarkably
rapid recovery of the original vegetation (see Pt. I, Sect. G). Today
large crabs once again burrow beneath the boles of well-maturing Pisonia
forests which their ancestors helped recreate less than 70 years ago.

It is hardly credible that these enormous crabs, the dominant
terrestrial animal of the atoll environment, could have been overlooked
by almost all visitors prior to the 20th century. The only plausible
explanation is that coconut crab populations were reduced drastically
each time Caroline was inhabited: 1846 to at least 1852, 1885 to 1901,
and 1916 to 1929. Voy collected them in 1875, 10 years before initial
land clearing began; Young noted them in a 1910 visit 9 years after the
first abandonment of the plantations, and again from 1916 on when copra
enterprises were begun anew. From 1916 to 1920 land clearing was far
more extensive, involving most of the area of the Windward Islets and
Nake (Pt. I, Table 13), and thus a very large number of homeless coconut
crabs would have been evident at that time. Again, mass slaughter
reduced their numbers until the main group of copra-cutters left in
1929. Since then, occasional Polynesian and other visitors have taken
crabs, but since the island was basically uninhabited for 60 years,
their numbers have recovered substantially. "Great numbers" were seen
in 1974 (Gilbert and Elllice Is. Govt. 1974). However, activities since
1990 do not bode well for the species (see Sect. G). It is of interest
in this regard that members of the 1934 Mangarevan Expedition saw no
coconut crabs on nearby Flint Island (R. Fosberg, pers. comm.), nor were
they mentioned in an historical summary paper on Flint by Maude (ca.
1942), but some were found at the southern tip of the island in 1906,
which at that time was covered in virgin mixed broadleaf forest,
including large Pisonia grandis trees, within whose boles the crabs
burrowed (E. Campbell 1908; A. Kepler, in prep.). Today Flint has
perhaps the greatest density of coconut crabs in the world (A. Kepler

46

1990b). Human pressures have likely operated on Flint as on Caroline
(A. Kepler, in prep.).

Distribution and habitat preference: In 1988 and 1990, coconut crabs
were abundant in the Cocos plantations of South and Nake, and present,
in varying densities, on 12 other motus (Fig. 17). Although generally
associated with Cocos, we found them in woodlands of Pisonia, Cordia,
and Tournefortia, as well as on rubble beaches (especially after dusk).
Although capable of surviving without coconut palms, these crabs appear
to seek them out. In the open understory of the tall plantations, or in
groves of only one or 2 palms, telltale piles of shredded coconut husk
fibers (Pt. I, Pl. 53) disclosed the crab’s presence.

Because the prevalent coarse rubble substrates on Caroline are
hard to burrow into, coconut crabs occupied a variety of shelters:
mounds of fallen coconuts and rotting palm fronds (to 1.5 m high), piles
of rubble pushed against tree roots, sand burrows, tunnels within the
feo (Pt. I, Pl. 21), or large cavities in the boles of mature Pisonia
trees. Coconut crabs also use a variety of shelters on the Tokelau
Islands (Yaldwyn & Wodzicki 1979), Flint, and Palau (AKK, pers. obs.).

Numbers: Though conspicuous and slow-moving, coconut crabs are very
difficult to count. Environmental variables such as rainfall, tide,
lunar cycle, and population size and age classes all affect their
activity (Reese 1965; Helfman 1977a, b). Although unable to conduct
mark-recapture studies, we did make incidental observations on the
numbers of individuals seen during transect and perimeter surveys.
Coconut crabs are generally nocturnal, but we often found them during
daylight, at times exposed on coral gravel beaches close to the
waterline. E. Reese (pers. comm.) suggests that the abundance of rats
occupying the same habitat may "force" the crabs to be more diurnal, as
has been reported from the Indian Ocean. Our estimate of the population
on Caroline is approximately 2,200 individuals, based on the number of
daytime observations, the area covered, and the fact that only one out
of every 3 or 4 individuals may be present on any given night (Helfman
1977b, Reese 1987).

Foraging: Since the first detailed description of coconut crabs in
1705, their shy, curious habits have been the subject of folklore,
speculation, and misinformation (see Reyne 1939). No scientist has yet
published a documented account of a coconut crab actually opening a
coconut (Helfman 1979), which is widely held to be their consummate
foraging behavior. Helfman is convinced that they do so, as he has
found piles of coconut fiber and observed crabs walking with husked,
opened nuts in places where he was the only other possible coconut
husker. We repeat Helfman’s (1979) assertion that coconut crabs do husk
fallen coconuts. The piles of finely separated fibers (Pt. I, Pl. 53)
we encountered are totally different from those produced by stick or
machete husking, the 2 methods commonly employed by Pacific peoples.

The crab tears virtually every fiber off individually, a process so
painstakingly slow it probably takes days. We did not observe this on
Caroline, but in March 1990 AKK, on uninhabited Flint Island, observed a
large male coconut crab that had just husked a coconut and was enlarging

COCONUT CRAB Gr

Figure 17.

a

Distribution map of coconut

CAROLINE ATOLL

SCALE 1:34,000
soo ° $00 1000 2000

——— Ee

meters

crabs on Caroline Atoll.

47

48

a small crack in the center of the smooth nut in a manner similar to
that described by Gardiner (1907) in Reyne (1939, p. 297). In June
1992, AKK also encountered large males eating coconut meat and husking
fibers in the Southwest Palau Islands, Micronesia.

On Caroline we observed the aftermath of coconut crab-Sooty Tern
predation or scavenging. On Brothers Islet, several entrances and
pathways leading to coconut crab holes were strewn with the feathered
skeleta of adult Sooty Terns (and possibly Brown Noddies), along with
numerous, freshly snipped branches of Pisonia up to 0.7 m long (Pt. I,
Sect. H). This was also recorded on Tridacna Islet by Clapp & Sibley
(1971a) for Sooty Tern eggs and chicks, and by Helfman (1979) and Reese
(1987) on Enewetak, Micronesia.

Sizes: Living in a rich environment free of predators, coconut crabs
attain huge sizes on Caroline. The bodies of the largest males were as
wide as a full-sized, unhusked coconut, giving them weights of at least
4 kg (G. Helfman, pers. comm.). Thorax widths for 10 crabs (2 females
with eggs, 8 males) averaged 129 mm. The thorax of the largest male
measured 200 mm across, making it, along with many measured on Flint in
1990 (A. Kepler 1990b), one of the largest recorded coconut crabs in the
world (the previous record was 178 mm in Helfman 1977a), with an age
estimated to exceed 40 years (E. Reese, pers. comm.).

G. CONSERVATION: ATTRIBUTES OF INTERNATIONAL SIGNIFICANCE

Caroline’s exceptional attributes need to be elucidated, for the
atoll has remained essentially unknown, even to some who have evaluated
its worth (King 1973; Stoddart 1976; Garnett 1983, 1984). There are
few, if any, islands remaining in the Pacific that can claim the
impressive array of natural features exhibited by Caroline (Nicholson &
Douglas 1969). We believe that it is imperative that this atoll, which
has managed to escape large-scale, permanent human disturbance, should
remain undeveloped.

Currently Caroline is uninhabited and has been since ca. 1930
except for a single family from 1987-1991. There are no roads,
vehicles, stores, jetties, or services (water, sewage, or food), and no
communication. There is no passage into the lagoon or safe sea
anchorage.

Lack of Major Disturbances

Man’s presence anywhere--especially on pristine or near-pristine
islands--generally brings rapid, often irreversible, changes. One of
the most important of Caroline’s attributes is its relative lack of
disturbance and very few exotic plant species. Aside from obvious human
impacts on South, Nake, and Ana-Ana, the majority of its motus are
dominated by indigenous vegetation and its reefs are basically pristine.
There is no obvious pollution to alter the chemistry of the lagoon,

49

beyond the flotsam and jetsam that litter the beaches. It is thus an
exceptionally clear and clean ecological laboratory that presents a
lagoon ecosystem before extensive disturbance by man, providing marine
biologists with opportunities to study undisturbed natural communities;
for example, the maze of patch reefs in the lower half of,the lagoon has
the highest recorded density of living Tridacna (20/.25 m™) ever
recorded (Sirenko & Koltun 1992). This is one of the few undisturbed
world populations of this species (Pt. I, Pl. 25). In addition to
conventional ecological studies, biomedical research could investigate
the causes and treatment of ciguatoxicity of fishes and crabs. Such
topics are increasingly important as more islands are subjected to
disturbance and pollution. For example, the abundant red snapper
(Lutjanus vaigiensis) and red spotted crab (Carpilius maculatus), both
notoriously poisonous, are safe to eat on Caroline. Caroline, lacking
the problems and pollution that beset many other Pacific islands, could
serve as a "control island."

Terrestrial Ecosystems

Caroline’s motus of varied age and size classes provide excellent
examples of substrate and vegetation development, accompanied by an
increasing diversity of bird life. On account of its relatively low
human disturbance and rapid forest recovery, especially since 1920,
Caroline is thus also an outdoor laboratory for terrestrial ecosystems:
many motus have recovered so remarkably they are almost
indistinguishable from those which have remained pristine, while others
are in different stages of recovery resulting from varied management (or
non-management) practices.

Caroline’s concentric pattern of plant community development and
the relationships of these communities to motu size, shape, and location
on the atoll rim could continue to provide insight into evolutionary
processes on atolls that are left undisturbed.

Physiography

Caroline offers many opportunities for geological research, under
reasonably unmodified conditions. Valuable clues as to the nature of
underground water supplies may lead to a better understanding of the
regulation of water supplies on inhabited islands. Notable
physiographic features include inland upraised reefs (feo) and deep sand
deposits, coalesced islets, exposed older reefs, lithified beachrock, a
conglomerate platform, a "perched lagoon," patch reefs, a nonfunctional
hoa, and changes in motu size and shape during the past century.

Flora

Caroline’s insular flora, typical of central equatorial islands in
their natural state and covering 70% of the atoll’s land area, is of
both national and international importance. The 26 extant plant species

50

are 89% indigenous (possibly 92%), an extremely high figure for anywhere
in the world. Six of the 7 plant communities are natural. Lushly
wooded, Caroline possesses tall Pisonia grandis) forests (Pt. I,

Pl. 43), reaching 21 m and occurring on 29 islets. Although less
majestjc than the prime forests of Washington and Fanning (Northern Line
Group), which enjoy a heavier rainfall, those on Caroline are notable:
Caroline’s 62 ha of Pisonia forest may well cover a larger area than on
any other Pacific atoll.

Caroline also possesses significant stands of the hardwood kou
(Cordia subcordata), which ranges from Africa to Polynesia, but is now
rare in the Pacific. Caroline’s groves (Pt. I, Pl. 26), although smal]
and often mixed with other indigenous trees, total 26 ha, possibly the
greatest area on any Pacific atoll. The extensive presence of tree
heliotrope (Tournefortia argentea) is also notable: scrub and forests
of this species form 40% of the atoll woodlands (Pt. I, Pl. 47).
Caroline’s large groves are some of the most unmodified in the Pacific,
as elsewhere Journefortia is typically restricted to coastal fringes
surrounding anthropogenic plantations (personal observations; Wiens
1962; R. Fosberg, pers. comm.).

Caroline, and its neighbor Flint, are also ideal islands in which
to monitor the transition from Cocos plantations to a more natural
state: rainfall regimes, intensity of management, underground water
reserves, slightly differing plant communities, and a wide range of
island/islet sizes have resulted in marked differences in revegetation
pattern only 60 years after the atoll was abandoned.

Seabirds

Associated with Caroline’s plant communities are 11 species of
breeding seabirds numbering well in excess of 1,000,000 individuals.
Almost every islet harbors nesting populations of up to 9 species
(Fig. 18). The populations of most of these species are of national
importance (Table 9). For example, Caroline has the fifth largest Red-
footed Booby colony in the world. Its Black Noddy and White Tern
(Pl. 3) populations are the largest in Kiribati. Under the 1975
Republic of Kiribati Wildlife Conservation Ordinance (amended in 1979),
all known seabirds, migrant shorebirds, and endemic land birds are
"fully protected throughout the Gilbert Islands" (Garnett 1983, p. 128).
However, their protected status is in doubt on Caroline, due to attempts
to commercially develop the island. Caroline deserves protection
similar to the 5 closed areas on Christmas Island and the 7 island
sanctuaries in the Line and Phoenix Groups (Garnett 1983), preferably
accompanied by on-site enforcement.

lin 1992, these forests were evidently altered considerably by immigrant
Gilbertese and cyclonic weather.

51

SEABIRD BREEDING

SPECIES DIVERSITY

CAROLINE ATOLL

SCALE 1:34,000
500 0 $00 1000 2000

meters

Figure 18. Seabird breeding species diversity by islet, Caroline Atoll.

52

Table 9. Comparative abundance of Caroline’s breeding seabirds in the Line Group.

Species Estimated Population Comparative Abundance in the Line Group
Red-tailed Tropicbird 3002 Second largest population
Masked Booby 400 Fourth largest population
Brown Booby 40 Third largest population
Red-footed Booby 7,000 Third largest population
Great Frigatebird 6,100 Third largest population
Lesser Frigatebird 200+ --
Sooty Tern 912,000 Third largest population
Brown Noddy 3,000 Third largest population
Black Noddy 17,000 Largest population (largest in Kiribati)
Blue-gray Noddy <10 ==) 4°
White Tern 8,000 Largest population (largest in Kiribati)

4 Based upon nest count in 1990.

Shorebirds

Caroline is an important wintering ground for the Bristle-thighed
Curlew, a rare shorebird and candidate for the U.S. Fish and Wildlife
Service Endangered Species list. Some subadults remain all year on the
atoll. As adult curlews pass through a flightless phase on their
Pacific wintering grounds, islands such as Caroline provide a predator-
free environment for this vulnerable phase of their life history.

Coconut Crabs

Caroline is exceptional in harboring a robust population of
coconut crabs (Fig. 17; Pt. I, Pl. 21). These large invertebrates are
abundant in the Cocos plantations of South and Nake and are found in
good numbers in the indigenous Pisonia forests on most larger motus.

Turtles

Although green turtles are not abundant on the atoll, worldwide
populations of these marine reptiles have suffered so greatly from
overexploitation that remote, predator-free islands such as Caroline
provide important, though small, sanctuaries. Since 1978 the Pacific
green sea turtle has been reclassified by the United States Department
of the Interior as threatened and the Pacific hawksbill sea turtle as
endangered.

Archaeology

From an archaeological point of view, Caroline houses one intact
Tuamotuan marae (ancient religious site) and another smaller site,
partly destroyed by storms. The main site (Pt. I, Fig. 3, Pl. 36),
basically undisturbed since the 1870s, is a relic of prehistoric
occupation worthy of protection and study, being the only one of its
kind in the Line and Phoenix Islands.

53
Current Conservation Status

Caroline Atoll is owned by the government of the Republic of
Kiribati and does not enjoy any legal protection (Garnett 1983, Ministry
of the Line and Phoenix Islands, pers. comm.). Over the last 50 years
it has been leased to private individuals who have scarcely altered the
atoll. The benign management of the past is no guarantee for the
future, and from October 1989 to the present, pressures to develop the
atoll have mounted rapidly. Proposed schemes included an airstrip, a
blasted channel through the reef, a hotel, a casino, logging, and
commercial harvest of fish and lobsters. In March 1990, commercial
harvesting of fish, the taking of coconut crabs, and illegal killing of
green turtles began, emphasizing that no island, however remote, is
guaranteed protection through isolation.

In addition, during the past 3 years Caroline has become more
visited than ever before, mostly without the knowledge or consent of the
Kiribati government. During 1990 yachts were present almost the entire
year, and in 1989 a cruise ship landed tourists who visited several
seabird colonies during midday heat. At this time Polynesian crewmen
presented some passengers with fresh tail feathers from Red-tailed
Tropicbirds ({B. Danielsson, pers. comm.). There are many reasons why
Caroline is inappropriate for resident tourists or development
(remoteness, distance from medical aid, no regular water supply, no
passage into the lagoon, mosquitos, etc.; see A. Kepler 1990a).
Caroline could support a limited number of ship-based ecotourists each
year, but indiscriminate visitation by yachts or other craft, such as
developed into an increasing problem at Suvarov, in the Cook Islands
(G. MacCormack, pers. comm.), should be discouraged.

Recommendations for an international preserve began in January
1989. During the 1990 ICBP expedition to the Line Islands, team leaders
discussed conservation matters with Kiribati government officials and
key scientists in French Polynesia. We obtained photodocumentation of
illegal land clearing and wildlife disturbance during the last 2 visits
to Caroline (A. Kepler 1990a, b, c). As a result, the Kiribati
government considered altering their plans for the development of
Caroline in favor of wildlife preservation.

In 1990 and 1991, The Nature Conservancy of Hawaii attempted to
establish a triple-island preserve on Caroline, Vostok, and Flint
through negotiations with Kiribati officials on Tarawa. These
negotiations failed, and the Kiribati government leased Caroline (and
Flint) for 55 years to the person responsible for the schemes and
illegal activities noted above.

54
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56

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57
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235: 12534

59
APPENDIX

Preliminary List of Arthropods

The knowledge of Arthropods on Caroline is limited to 15
Lepidopterans collected by Dr. Palisa during the Solar Eclipse
Expedition (Dixon 1884), a scant list of common insects (beetle, gnat,
etc.) in the same paper, and a preliminary collection of 82 specimens
made by Graham Wragg and AKK in 1990. The latter, identified by David
Preston and Scott Miller (B. P. Bishop Museum, Honolulu, Hawaii),
contained no endemics, consisting primarily of widespread Pacific
species and immatures identifiable only to family level.

Although our collection is also scant, it is the second collection
of Arthropods from Caroline and the only one containing species other
than Lepidoptera: JIsometrus maculatus (scorpion), Scolopendra
subspinipes (centipede), Anoplolepis longipes (long-legged ant),
Isopoda, lepidopterans (larvae and pupa), dermestid beetles (larvae and
adults), cockroaches (Dictyoptera), Hibboboscidae adult (Diptera),
Isopoda, Nitidulidae (immature and adults, Coleoptera), Curculionidae
(Coleoptera), immature Hemiptera (Lygaeidae), Scolopendriidae immature,
spiders (Arachnida).

of he FOB = a

Plate 1. Incubating Red-tailed Tropicbird, Bo’sun Bird Islet,
Caroline Atoll, 25 September 1988. The nest scrape is in fine coral
rubble under a Journefortia shrub.

ern

*. sila) Sta
> > me »
a a. = eS
aad > SB as oa hid a" .

Plate 2. Masked Booby adult with egg on coarse coral rubble substrate
with Portulaca mat, Nake Island, Caroline Atoll, 26 September 1988.

61

Plate 3. White Tern adult with egg in typical nest site, a dead
Tournefortia branch, South Island, Caroline Atoll, 23 September 1988.

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